ABSTRACT

We present here the Synopsis of South American Angiosperms (SSAA), a broad text that intends to summarize in a succinct way all the native diversity of angiosperms on South America, including Malvinas as part of Argentina, and excluding southern Panamá. This work is structured by order, families, subfamilies/tribes/subtribes (if present), genera, subgenera/clades/sections/subsections/series (if present), with a small entry for each.

NUMBERS

In these numbers, we identified in November 29, 2023, exacts 4,151 native genera of angiosperms in South America (4,149 formally described and one, in Violaceae, counted but not described), 2,767 in Brazil (358 endemics) and 1,384 absent (see TABLES below).

REFERENCES

To prepare this synopsis, the author used the following general references:

VPA: VASCULAR PLANTS OF AMERICA as of April 3, 2023 (86,072 spp. of vascular plants in South America, including lycophytes, monilophytes, gymnosperms and angiosperms, DOWNLOAD DATA), for general primary information on all genera mentioned here, the vast majority of which follows the information collected on the aforementioned date.
POWO: PLANTS OF THE WORLD ONLINE, counterpoint to VPA and heavily used in the circumscription of many of the genera listed here.
ANGIOS BERGIANSKA, for definition of subfamilies and tribes and many families, and the distribution of outsiders (any taxa absent in South America).
APG: ANGIOSPERMS PHYLOGENY GROUP WEBSITE, for the exact and precise definition of families and orders used throughout this text.
THE FAMILIES AND GENERA OF VASCULAR PLANTS, which provided inspiration for the existence of this project and much of the information of genera and families present throughout the text.
an incalculable quantity with precision, but ranging between 5,000 to 7,000, of articles that were read and considered in the creation of this text.

ACESSORIES

VPA LIST RECONSIDERATIONS FOR SSAA LISTING, which brings c. 746 observations used here to the 1st version of this page.

VPA LIST RECONSIDERATIONS FOR SSAA LISTING FOR FIRST VERSION

NEWS AND UPDATES, a list of new species (1,077 since April, 2023) and taxonomic changes that update and modify the VPA primary list, organized and expanded with each new update of this work.
an incalculable quantity with precision, but ranging between 5,000 to 7,000, of articles that were read and considered in the creation of this text.

OTHER WORKS

SIDERASIS ANGIOSPERMS, which, in conjunction with this work, provides a large amount of information, with emphasis on infrafamilial and infregeneric notes, in addition to the excellent explanation of morphological and ecological notes.

COLORS

BLACK genera refer to Brazilian non endemic genera; BROWN genera refer to Brazilian absent genera; and BLUISH refer to unoficially genera (only one in this list, in Violaceae).

ANGIOSPERM PHYLOGENY

ANGIOSPERM PHYLOGENY TREE, BI LI ET AL (BMC BIOLOGY, 2021)

1. AMBORELLALES

AMBORELLALES DOES NOT OCCUR IN SOUTH AMERICA, AND IS COMPOSED OF A SINGLE FAMILY, AMBORELLACEAE (1/1).

2. NYMPHAEALES

FAMILIES ABSENTS IN SOUTH AMERICA: HYDATELLACEAE (1/13).

NYMPHAEACEAE

§ FULLY CHLOROTROPHIC

Genera/species 5/70–90 Distribution Nuphar: North America, Europe, E Asia; Barclaya: SE Asia, Malesia; Nymphaea almost cosmopolitan; Ondinea: northern W Australia; Victoria: northern and central South America; Euryale: northern India, China, Japan. Habit bisexual, usually perennial (rarely annual) herbs. Aquatic. Usually with rhizome or tuberous stem (some species of Nymphaea stoloniferous).

Nympheaceae is a predominantly temperate family, although V. amazonica Sowerby and V. cruziana Orbign. are found in the Amazon river. Many species are used in horticulture. Nuphar is introduced (cultivated). Nymphaeaceae are often pollinated by beetles, and sometimes their flowers open at night. Protogyny has been studied in Victoria amazonica. The wide distribution of species in the family encloses tropical and temperate regions, occurring in rivers, ponds, lakes, and other freshwater wetlands. Nymphaeaceae are aquatic plants fixed and rooted on the substrate with floating leaves, sometimes submerged, long petiolated.

Nymphaeaceae are closely related to the Cabombaceae, sharing with them: the rhizomatous stem; involute, peltate leaves with palmate secondary veins that are actinodromous, festoon brochidodromous, margin toothed or entire; flowers are single along the stem. However, Nymphaeaceae has relatively large, showy flowers with 4-6(-12) sepals and 6(-70) petals, while Cabombaceae have considerably fewer perianth parts and the flowers are less conspicuous.

Hydatellaceae aside, for a morphological phylogeny of [Cabombaceae + Nymphaeaceae], see D. W. Taylor (2008). Y.-L. Liu et al. (2005) provide an ITS phylogeny focussing on Nymphaeaceae, but with some at first sight rather surprising relationships - [Nuphar [Cabomba + Brasenia] [Nymphaea [Euryale + Victoria]]]. Nelumbo, which was included in the analysis, did at least stay outside this clade. However, this position of Nuphar is recovered in other analyses, too (e.g. Borsch et al. 2008; Löne et al. 2007; D. W. Taylor & Gee 2014: some analyses; Gruenstaeudl et al. 2017) and so its position below as sister to Nymphaea, etc., may be provisional. He et al. (2018: chloroplast data) found that Nuphar was sister to Nymphaea and its associates, and with quite strong support; interestingly, a UPGMA dendrogram of RNA editing sites linked Nuphar with Cabombaceae. Moreover, in a renalaysis of the data in He et al. (2018), Gruenstaeudl (2019) found that how the data were partitioned affected the results, and individual gene trees also gave different results, thus almost as many supported a relationship of Cabombaceae with Nuphar as supported an association of Nuphar with Nymphaea and friends. H.-T. Li et al. (2019) found a [Nuphar + Nymphaea] association.

SYSTEMATICS two tribes, Nuphareae (1/20, North America, Europe, E Asia) does not occur in South America; among Nymphaeea, with Nymphaea expanded, Barclaya (4; SE Asia, Malesia) is a outsider.

1. Nymphaea L. Herbaceous, rhizomatous perennials; leaves mostly floating. 67 spp., cosmopolitan, 30 in New World; six subgenera, Anecpjya (9, Australia), Confluentes (7, tropical areas of Australasia) and Lotus (4, Africa, Philippines and in a hot springs in Romania and Hungary) absent in New World, and three in this Hemisphere:

§ subg. Brachyceras ‣ 20 spp., inc. N. ampla (Salisb.) DC., N. elegans Hook. and N. pulchella DC. in South America, only the third in Brazil.

§ subg. Hydrocallis Planch. ‣ a neotropical clade with nocturnal flowers, stamens inserted in series with the petals, entire leaves or dentate-sinuate with obtuse teeth, includes 21 spp., all in South America, 19 in Brazil, six endemics.

§ subg. Nymphaea ‣ 6 spp: N. candida C.Presl (Europe to Siberia and W Himalaya), N. alba L. (Europe to W. Himalaya, NW Africa), N. leibergii (Morong) Rydb. (Canada to N U.S.A.), N. tetragona Georgi (N Europe to Korea and Himalaya, Subarctic America to NW U.S.A.), N. odorata Aiton (Central & E Canada to Nicaragua, Bahamas, Cuba), N. mexicana Zucc. (Oklahoma to SE U.S.A. and Mexico).

2. Victoria Lindl. Annual or short lived perennial aquatic herbs; mature blade floating; flowers nocturnal, white, charging to pinkish red; very numerous petals, thorns or spines in the outside side of its leaves, petioles, peduncles and outer sepals, the leaf margin is distinctly folded upwards. Three spp., V. amazonica (Poepp.) Klotzsch from W Peru to N Bolivia and low Amazon river in northern Brazil with isolated records in Roraima state in Brazil, also Guyana; absent in Ecuador and Venezuela, V. cruziana Orb. in low Paraguay river and subsequent Paraná river, in Argentina and Paraguay, being absent in Brazil; and V. boliviana Magdalena & L.T.Sm, endemic to flood plains of the Llanos de Moxos, Mamoré watershed, E Bolivia. Two morphotype species occur in Brazil, both up to Bolivia, one also in Paraguay.

V. boliviana has the third species-by-species width largest flowers in New World (after two Pachira species), up to 50 cm diameter, and their circular floating leaves have a documented diameter of up to 2.4 meters (8 feet), would have an area of 50 square feet.

CABOMBACEAE

§ FULLY CHLOROTROPHIC

Genera/species 2/6 Distribution tropical, subtropical and temperate parts of North, Central America and South America, Caribbean, Africa, China (inc. Taiwan), Korean Peninsula, Japan, Manchuria, northern India, and NE and SE Australia Habit bisexual, perennial herbs. Aquatic, with rhizome and usually floating stem; submersed organs of Brasenia covered by a gelatinous layer (absent in Cabomba); leaves of Brasenia isomorphous (monomorphous), in Cabomba anisomorphous (dimorphous).

SYSTEMATICS two genera, both in South America.

1. Brasenia Schrb. Herbs, rhizomes horizontal, producing upright shoots with alternate, oval, peltate leaves. Only one sp., B. schreberi J.F.Gmel., a wind pollinated species with reduced flowers, distributed in Africa, temp. & subtrop. Asia, E Australia, Canada to Costa Rica, Caribbean, Colombia, Guyana, Venezuela (Merida, Tachira, Trujillo, Zulia) and French Guiana (very closed to Brazilian borders), in ponds, lakes, and slow-flowing rivers and streams, and cultivated for its edible shoots.

2. Cabomba Aubl. Herbaceous aquatic with slender branching stems; submersed leaves opposite and whorled palmately dissected into linear segments; flowers axillary, white to yellow or purple. 6 spp., widely distributed from the U.S.A. to Argentina, slightly centered in South America, 5 in South America, all occur in Brazil, with C. schwartzii Rataj. endemic to Brazilian Amazon (Amazonas and Roraima states).

3. AUSTROBAILEYALES

AUSTROBAILEYALES DOES NOT OCCUR IN SOUTH AMERICA , AND IS COMPOSED OF THREE FAMILIES, AUSTROBAILEYACEAE (1/1), SCHISANDRACEAE (3/80), TRIMENIACEAE (1/8).

4. CANELLALES

TWO FAMILIES, BOTH IN SOUTH AMERICA.

CANELLACEAE

§ FULLY CHLOROTROPHIC

Genera/species 5/13–15 Distribution 5 genera: two in Africa (Cinnamosma: Madagascar; Warburgia: tropical E Africa and northern Transvaal) and three in New World. Habit aromatic trees (or rarely shrubs); leaves simple, alternate, leathery; stipules absent; sepals 3; petals fleshy, imbricate (in 1-4 whorls); ovary superior, syncarpous, 1-locular; placentation parietal; style 1, short; stamens connate in a tube; fruits berries. The Greater Antilles is the center of diversity and endemicity for the family, with three genera (Canella, Cinnamodendron and Pleodendron). In the Neotropics, most species have local distribution and are very rare, except for Canella winterana (L.) Gaertn.

The Cinnamodendron species from the Greater Antilles (5) and from South America (6) do not form a monophyletic taxon and each group is characterized by the number of floral parts (petals, stamens, carpels, ovules) and also by differences in the seeds. Pleodendron has two spp. of Caribbean (one in Haiti and Puerto Rico) and one in Costa Rica; the widely distributed Canella winterana is traditionally used for medicinal, fishing, aphrodisiac, ritualistic, and aromatic purposes.

In Winteraceae, the wood lacks vessels, the leaves are usually glaucous abaxially, the stamens are free, the gynoecium is apocarpous (carpels 1-24) and the placentation is marginal. On the other hand, in Canellaceae, the wood has vessels, the leaves are usually glossy, the stamens are connate in a tube, the gynoecium is syncarpous (carpels 2-6) and the placentation is parietal.

SYSTEMATIC outsiders Cinnamosma (3; Madagascar), Pleodendron (3; Costa Rica; Hispaniola; Puerto Rico) and Warburgia (4; tropical E Africa, NE South Africa).

1. Canella P. Brown. Trees, leaves punctate; inflorescence terminal broad panicles, petals 5 basally connate, stamens 10, style 2-3-lobed. Only one sp., C. winterana (L.) Gaertn. in SE U.S.A. (southern part of Florida Keys, Cape Sable), Caribbean and N Venezuela, Mexico and Central America.

2. Cinnamodendron Endl. Trees. Inflorescence axillary racemes, or solitary flowers axillary; petals 6-10 (in 2 whorls), stamens 8-10, style indistinctly lobed. 14 spp., 4 in Caribbean (Haiti, Cuba, Jamaica and Dominican Republic, one each) and 8 in tropical South America, all national endemics, in Venezuela (1), Suriname (1) and Brazil (6), all of these endemic to SE region, two of them, from São Paulo and Rio de Janeiro states, are rare plants in Brazil, by Plantas Raras do Brasil’s book.

WINTERACEAE

§ FULLY CHLOROTROPHIC

Genera/species 5/c. 80–110 Distribution northern Madagascar, Philippines, Malaysia, Indonesia, New Guinea, Moluccas to Solomon Islands, eastern Australia, Tasmania, Lord Howe, New Caledonia, New Zealand, Mexico to Magellan’s Strait, Juan Fernández Islands, with their highest diversity in Queensland and New Caledonia, mainly in subtropical montane forests. Habit usually bisexual (sometimes monoecious or polygamomonoecious; Tasmannia usually dioecious), evergreen trees or shrubs (rarely epiphytic; Tasmannia piperita (Hook.f.) Miers with lignotuber). Wood aromatic; alternate, simple leaves arranged spirally and somewhat irregularly clustered; leaf blade with secondary veins that do not extend to the leaf margin but link up with the neighbouring one; underside of leaf waxy and seemingly white/blue in colour; during or after flowering the twig elongates monopodially from the apical bud of the inflorescence; flowers bisexual with the sepals forming a cap-like structure which covers the stamens and carpels in bud; stamens spirally arranged; superior ovary; fruits indehiscent.

Due to the basal position of Winteraceae in the angiosperm phylogenetic tree the flower morphology is variable and not as fixed as more recently evolved families; there is considerable polymorphism and variation in both foliage and floral characters. This instability can lead to difficulty in determining the species of the specimen.

SYSTEMATIC subfamily Takhtajanioidea (1/1, Madagascar) does not occur in South America; among the other subfamily, Wintereoideae, outsiders are Tasmannia (c 40; Philippines, Borneo, Sulawesi, New Guinea, E Australia), Pseudowintera (3; New Zealand, Stewart Island) and Zygogynum (c 60; New Guinea to New Caledonia and Queensland).

1. Drimys J. R. 7 G. Forster. Evergreen shrublets to tall trees; twig elongating monopodially after flowering. 7 spp. generally grow in moist montane tropical environments, from Veracruz in E & C Mexico to Peru and Venezuela, E Brazil, and southern South America: D. roraimensis (A.C. Sm.) Ehrend. & Gottsb. in Venezuelan montane and cloud forests, dwarf forests, slopes of tepuis, along streams and in swamps, at 1800-2500 m alt. and adjacent areas in Roraima state in N Brazil; D. granandensis L. f. from Mexico through to Peru and Venezuela; D. winteri in Argentina, Chile and Bolivia; D. angustifolia Miers in nebular mountains forests in south Brazil; D. brasiliensis Miers endemic to Brazil; D. confertifolia Phil. only in endemic to the Juan Fernandez Islands; and D. andina (Reiche) R.A.Rodr. & Quez., a shrub restricted to higher parts of the Coastal and Andes mountains in Chile.

5. PIPERALES

FAMILIES ABSENTS IN SOUTH AMERICA: SAURURACEAE (4/6).

ARISTOLOCHIACEAE

§ PARASITIC (Prosopanche – Cassytha - ... – OROBANCHACEAE - Cuscuta)

Genera/species 8/511 Distribution tropical, subtropical and temperate Eurasia, Africa, America, and Australasia (southwards to northern Australia), dry regions of Africa, Réunion, Madagascar, and the southwestern Arabic Peninsula (Hydnora), and Costa Rica, southern South America and NE Brazil (Prosopanche). Habit usually bisexual (rarely unisexual), usually perennial herbs with monopodial growth (usually scrambling or climbing; sometimes suffrutices, shrubs or lianas, e.g. Aristolochia arborea); branches often with somewhat swollen nodes. Hypocotyledonary tuber often present. Achlorophylous herbs in two genera.

Plants in the family Aristolociaceae contain aristolochic acids. These have been used in traditional Chinese medicines e.g. Isotrema manshuriense (Kom.) H. Huber or in Chinese Mu Tong or Guan Mu Tong for the treatment of mouth ulcers, oedema, urinary problems and arthritis; the use of this species has been banned in Europe since 2001 and in China since 2003 due to the aristolochic acids being found to be carcinogenic, nephrotoxic and mutagenic.

In European herbal medicine Aristolochia clematitis L. was traditionally used to aid child birth. In America species of Aristolochiaceae have been used to treat snake bites which lead to one of its common names of Snake Root. Aristolochiaceae are fly pollinated which probably led to the evolution of their floral shape.

The structures of their flowers are similar to the flytrap mechanisms found in the carnivorous genus Nepenthes L. However, whereas Nepenthes digests the insects as a means of adapting to environments with poor soil, Aristolochia and Isotrema simply make use of them for pollination. The flies are attracted by the 'alluring', putrid scent of the flowers and pass through the smooth, waxy 'slide zone' on the limb of the flower. At the base of this zone they become trapped in the utricle, an inflated part of the perianth, by the retrorse hairs at its entrance. The insects pollinate the stigmas as they enter the utricle, remaining there until the stamens have ripened and deposited pollen on them as the flower withers. Meanwhile the flies are kept alive with supplies of nectar from trichome nectaries that are usually present in the utricle. Most Aristolochia and Isotrema flowers wither after about 24 hours, and as they do so the retrorse hairs that trapped the insects lose their turgidity, releasing them to pollinate more flowers (Petch, 1924).

Caterpillars of some swallowtail butterfly species and several other species of butterflies feed on the leaves, shoots and flowers of Aristolochia and Isotrema. The caterpillars can tolerate and sequester the aristolochic acids which are believed to render them poisonous to predators.

SYSTEMATIC lineage Asaroideae (2/120, Northern temperate regions, with their largest diversity in E Asia) does not occur in South America.

1. HYDNOROIDEAE CLADE (2/10) ‣ outsider Hydnora (6; arid and semiarid regions in tropical and southern Africa, Madagascar, Réunion, SW Arabian Peninsula).

1. Prosopanche de Bary. Herbaceous achlorophyllous endophytic root holoparasites mainly on Fabaceae (in, e.g., Acacia and Prosopis) and Euphorbiaceae, leafless, roots rhizome –like; flowers from rooting body (frequently semi-subterranean), large, solitary, bisexual, regular, thick, fleshy, brownish, rugoes on outer surface, coloured on innder surface; fruit sweet-smelling, coriaceous berry, woody pericarp splitting irregularly; seeds small and numerous (500 - 90,000). 7 spp., P. americana (R. Br.) Kuntze. in dry areas of Bolivia to Argentina; P. bonacinae Speg. in dry areas of Argentina, Uruguay and, until now, only one specimen of this species was recorded from Brazil, collected by Jürgens in Rio Grande do Sul state in a tobacco plantation; P. caatingicola R. F. Machado & L.P.Queiroz on mimosoid Fabaceae, known only from a small area in the municipality São Gabriel, Bahia, in a small area of ca. 200 m²; P cocuccii Tav. de Carvalho, Záchia & Mariath endemic to center Rio Grande do Sul state, Brazil; P. costaricensis L. D. Gómez of forests of Costa Rica; P. demogorgoni Funez, known only from the type locality in Urubici municipality, Santa Catarina state, S Brazil; and P. panguanensis C.Martel & Rob.Fernandez in Amazon rainforest of Peru.

Lytton J. Musselman defined as the strangest plant in the world; it had been described as a group of fungi (‘Hydnorinae’), even though the author recognised the floral nature of its reproductive structures in 1821.

2. LACTORIS CLADE (1/1) ‣ a single genus.

2. Lactoris Phil. Small herbaceous shrub. Only one sp., L. fernandeziana Phil., mostly restricted to windy fogand rain-swept mountain forests on steep slopes, where it occurs as an inconspicuous member of the understory (but the canopy over this species is often only at 2–4 m), although it is occasionally found in full sunlight on near vertical exposed cliff faces, in the island of Masatierra (Robinson Crosue I.) in the Juan Fernández archipelago.

3. SUBFAMILY ARISTOLOCHIODEAE (3/480) ‣ outsiders are Isotrema (98, 81 in Asia and 17 from North to Central America), Thottea (35, India to SE Asia) do not occur in South America.

3. Aristolochia L. (inc. Euglypha) Lianas or woody vines, rarely herbs or shrubs, sometimes with xylopodium, odorous; none petals in neotropical members, and with aristolochic acids. 540 spp., 277 in New World, 168 in South America, 98 in Brazil, 51 endemics; in South America all species of Aristolochia belong to Aristolochia subg. Aristolochia sect. Gymnolobus subsect. Hexandrae; this subsection is currently divided into two series:

§ serie Hexandrae ‣ inflorescences in racemes, often reduced to a single flower, petiole and floral peduncle with no basal zone of abscission, fruit with entire septa, seeds without wings or 1-winged; two subseries:

§ subser. Hexandrae ‣ bears axillary, solitary and ebracteate flowers.

§ subser Anthocaulicae ‣ ramiflorous to cauliflorous racemes with extremely short internodes and flowers subtended by reduced bracts; 22 spp. mainly from pluvial lowland forests from Belize to Bolivia, only 5 spp. in Brazil, mainly Amazonic: A. bahiensis F. González, A. cauliflora Ule, A. klugii O.C. Schmidt, A. lanceolatolorata S. Moore and A. ruiziana (Klotzsch) Duch.

§ serie Thyrsicae ‣ thyrsoid inflorescences, petioles and peduncle with basal zone of abscission, fruit with lattice-like septa, 2-winged seeds; only 5 spp. in Brazil, A. acutifolia Duch., A. dalyi F. González, A. paramarimbensis Duch., A. silvatica Barb. Rodr. and A. sprucei Mast.

A. rojasiana (Chodat & Hassl.) Hosseus (syn. Euglypha rojasiana Chad. & Hassl.), is a subherbaceous or slightly woody twiner without pseudostipules; grows on SE Bolivia, Paraguay, N Argentina and Mato Grosso do Sul state, Brazil; A. holostylis F. González (syn. Holostylis reniformes Duch.), is a coarsely herbaceous, procubent or ascending, without pseudostipules; grows on SE Bolivia, Paraguay, and WC Brazil to Maranhão state in northeastern; the largest flower in this family possibly belongs the over Brazilian endemic A. gigantea Mart. & Zucc., reaching up to 36 cm diameter, with natural populations only Bahia and Minas Gerais mountains.

PIPERACEAE

§ FULLY CHLOROTROPHIC

Genera/species 5/3,825 Distribution pantropical: Central America and the Caribbean to Argentina, Central Africa to Madagascar, southern India and Sri Lanka, southern China to E Australia, Tasmania and New Zealand, islands in the Pacific. Habit bisexual, monoecious or dioecious, evergreen trees, shrubs or lianas, erect or climbing, usually perennial (in Peperomia sometimes annual) herbs (Peperomia often epiphytes), often succulent, often with articulated stems and swollen nodes. Aromatic and with sharp (peppery) taste. CAM physiology present in many species of Peperomia.

Piperaceae is pantropical, with major concentrations and centres of diversity in Central and South America and in Malaysia. The members of the family most frequently inhabit rain forests. Many species of Piperaceae are rich in secondary metabolites, including alkaloids, amides, flavonoids and terpenes, having economic and medicinal importance: Piper nigrum L. (black pepper), Piper umbellatum L. (pariparoba) and Piper methysticum L. (kava-kava); species of Peperomia are commonly used as ornamental house plants. Zippelia is a monotypic genus of Asian distribution, with floral morphology considered intermediate between Piperaceae and Saururaceae. The megagametophyte is eight-celled. Investigated species of Piper contain a huge amount of alkaloids, amides, lignans, neolignans and terpenes. Piper occur in various habitats but are best represented in lowland tropical rainforests, frequently in formations below 1,500 m elevation and also commonly found in disturbed areas. Some spp. are widely distributed but several are restricted to specific centres of diversity and each country has a large number of endemics.

Saururaceae differs from Piperaceae by having (2-)3-4 distinct or fused carpels (vs. 1), if distinct then each carpel with 2-4 ovules, if fused then ovary 1-locular and each carpel with 6-13 ovules, placentation parietal (vs. basal), and fruit dry, schizocarp or apically dehiscent capsule (vs. berry or drupe).

Key to genera of Neotropical Piperaceae

1. Herbs with alternate, opposite or whorled leaves; stamens 2; stigma 1 ------------ Peperomia

1. Shrubs or small trees, sometimes lianas or climbing shrubs with alternate leaves; stamens 2-6; stigmas 3-5 - 2.

2. Shrubs or small trees, glabrous or indumented; inflorescence a solitary spike or raceme, leaf-opposed or terminal ------------ Piper

2. Lianas, glabrous; inflorescence a solitary or paired spike, axillary or occasionally terminal ------------ Manekia

SYSTEMATIC two tribes, both in South America.

1. TRIBE ZIPELIEAE (2/4) ‣ outsider Zippelia (1; SE Asia to Java).

1. Manekia Trelease. Vines, trailing along forests floor or climbing via rooting at noted of monopodial axes; leaves simple; inflorescences spicate, on sympodial axes only, solitary or 2 together, appearing axillary or terminal, erect at anthesis but usually pendulous in fruit. Three spp., M. urbanii Trel. endemic to SW Haiti, M. obtusa (Miq.) Arias endemic to rain forests of SE Brazil (50-1,500 m), and M. incurva (Sieber ex Schult.) T. Arias, Callejas & Bornst. from Costa Rica to Peru, N Venezuela and Lesser Antilles.

2. TRIBE PIPEREAE (2/c. 3,500) ‣ both genera in South America.

2. Peperomia L. Herbs erect, prostate, pendent or assurgent, usually succulent (mainly in Andes), terrestrial, rock-living or epiphytic; leaves alternate, opposite or whorled; venation palmate, pinnate or plinerved; inflorescence axillary, terminal or leaf-opposed, in spikes or racemes, solitary or paniculate, sometimes on junction of the petiole and the leaf blade (true epiphylly); fruit in depression of the rachis or stipitate, commonly verrucose and viscid. c. 1,300 spp. worldwide, 1,200 in New World, 957 in South America, 406 in Peru, 267 in Colombia; about 17/20 of Peperomia species are from Neotropical to Argentina regions, followed by SE Asia, Africa, Australia, and New Zealand; Brazil harbors 172 species, of which 112 are endemics; the species are mostly found in humid forested regions and few in dry area; 23 Peperomia species from Brazil are considered rare, by Plantas Raras do Brasil’s book.

Modern analyzes indicate 8 distinct lineages in this genus; Brazilian members are confirmed for all clades except Clade A. Peperomia subg. Tildenia is one of the most easily recognizable of these groups, with 30 geophytic species with two easily observable synapomorphies: an entirely or partially underground tuber, and with petioles and peduncles originating directly from the tuber; all these species occur in seasonal habitats of Neotropical mountain areas from Mexico to Argentina, with an almost equally high diversity in Mexico and Guatemala in the northern hemisphere and in Peru and Bolivia in the southern hemisphere. In between these two hotspots, there are only a few species known from a limited number of localities.

3. Piper L. [7th BR, inc. Ottonia] Shrubs, subshrubs, lianas, occasionally subherbaceous or small trees with nodose stems; leaves alternate, gland-dotted, petiole usually present; venation palmate or pinnate; inflorescence a solitary spike or raceme (subg. Ottonia) leaf-opposed or terminal, erect, pendent or recurved, fruit in depression of the rachis or stipitate sometimes laterally compressed, glabrous or with trichomes. c. 2,100 spp., pantropical, mainly in the New World (1,922), followed by SE Asia (300), 15 in Africa, where the economically important species P. nigrum L. (black pepper) and P. betle L. (betel leaf) originated. 1,218 spp. in South America, 450 in Peru, 425 in Colombia, and the seventh largest genus in Brazil, with 301 spp., 203 endemics; 40 spp. of Piper are considered rare in Brazil, by Plantas Raras do Brasil’s book. 8 subgenera of Piper are recognised in the Neotropics:

§ subg. Enckea ‣ c. 120 spp., from the humid forests of the Amazon, to the much drier forest west of the Sierra Madre Occidental in México, and the calcareous outcrops of the Greater Antilles; species in this clades are shrubs or small trees with very characteristic palmately nerved leaves.

§ subg. Macrostachys ‣ 200–250 spp., distributed throughout the Neotropics, but most species occur in Central America and the Northern Andes, a few species in the Amazon and a handful in the Atlantic Forest of Brazil. Species of this clade are medium-sized shrubs to small trees; leaves are pinnately nerved and basally lobulate.

§ subg. Ottonia ‣ c. 50 species and is distributed primarily in the Atlantic Forest of Brazil and the Amazon, but has one species, P. darienense C. DC., that occurs west of the Andes; species of Ottonia are small shrubs that thrive in the understory of lowland tropical forests; leaves are pinnately nerved, ovate or lanceolate in shape, and with two basal callosities.

§ subg. Pothomorphe ‣ c. 10 spp., and is distributed throughout the Neotropics; species in this clade are herbs or shrubs, and the latter can be either self-supporting or climbers.

§ subg. Peltobryon ‣ c. 80–100 spp., widespread in South America; most species are restricted to the western Amazon and the eastern Andean slopes in Bolivia, Ecuador, and Perú; a significant portion of this clade occurs in the lowland forests of Costa Rica and Panamá; few species are known north of Guatemala and apparently none is known from the Caribbean Islands; species of this clade are shrubs or treelets that thrive in the forest understory; they occur from sea level to 2000 m in elevation.

§ clade Piper cinereum | P. auritum ‣ two spp., from Mexico to Ecuador, Venezuela, Suriname and French Guiana; absent in Brazil.

§ subg. Radula ‣ c. 450 spp., is broadly distributed in the Neotropics.

§ subg. Schilleria ‣ c. 200 species found in Central America and in the Atlantic Forest of Brazil, where they are more diverse.

Patterns of distribution of Piper species vary from being locally endemic to widely distributed. There are several species restricted to a specific center of diversity (e.g., Andes, Central America) and others occur throughout the Neotropics or the Paleotropics. Piper is often a dominant element in the understory of tropical forests and found to be one of the five most speciose genera in select Neotropical forests; not surprisingly, Piper species are of great ecological importance and have been considered ‘key’ species on the basis of their association with frugivorous bats. In New World 18 spp. of Piper are mymecophyes, two in Brazil.

6. MAGNOLIALES

FAMILIES ABSENTS IN SOUTH AMERICA: DEGENERIACEAE (1/2), EUPOMATIACEAE (1/3), HIMANTANDRACEAE (1/1).

MYRISTICACEAE

§ FULLY CHLOROTROPHIC

Genera/species 21/ c. 640 Distribution pantropical, with the largest diversity in tropical Asia. Habit usually dioecious (rarely monoecious), usually evergreen (rarely deciduous) trees (rarely shrubs or lianas). Often aromatic. All Myristicaceae are woody, medium sized trees, seldom shrubs in South America Virola and lianas in a Old World genera. None genus occur in more of one continent: five genera in Asia, five genera in America and eight in Africa and Madagascar.

The flowers are unisexual. Parenchyma, wood rays and phloem contain canals with red, orange or yellow resins. The nucleus of the immature sieve element has one or rarely two protein crystals which are released inte the cytoplasm as the nucleus degenerates and persist within the mature nucleus-free sieve element. The androecium is very variable, although usually the stamens are borne in a single whorl and connate into a synandrium surrounding a central sterile column. Mauloutchia has free stamens, in some species spirally arranged and numerous, a small aril and distinctly granular exine.

In the Neotropics, the Myristicaceae is found mostly in lowland in the Amazon rainforest; all species of the family were placed in the single genus Myristica for a long time; later, the family was divided in many genera, based mainly in the variations of the androecium, and also in inflorescence, aril and pollen; some authors consider the perianth parts as sepals (petals absent). The exudate of several species of Virola is hallucinogenic.

KEY DIFFERENCES FROM SIMILAR FAMILIES

ü Myristicaceae differs from Magnoliaceae mainly by: the presence of exudate, the absence of stipules, the inconspicuous flowers with few perianth parts and the flowers aggregated in inflorescences. In Magnoliaceae, the exudate is absent, the stipules are present, the flowers are large, solitary and with numerous perianth parts.

ü Myristicaceae can be mistaken for Annonaceae due to the myristicaceous branching and the alternate, distichous and exstipulate leaves; however the latter differs in lacking the red exudate (with rare exceptions), in having fibrous bark, numerous stamens and an apocarpous gynoecium.

ü Myristicaceae differs from Canellaceae mostly by: the presence of exudate, the unisexual flowers with few perianth parts, basal placentation and the capsular fruits.

SYSTEMATIC three well defined clades, Knema clade (5/c. 325, tropical Asia and Africa to tropical Australia) absent in South America.

1. CLADE OTOBA (8/40) ‣ outsiders Coelocaryon (4; tropical Africa), Pycnanthus (4, tropical Africa), Staudtia (2, tropical W Africa to W Uganda), Cephalosphaera (1; mountains in tropical E Africa), Doyleanthus (1; Madagascar), Brochoneura (5, E Madagascar), Mauloutchia (10; E Madagascar).

1. Otoba (A. DC.) Karsten. Trees dioecious; flowers ebracteolate; usually has glaucous leaves on the lower surface; stamens are few (3(-4)), the anthers free of the column (or basally adnate); fruits are globose to ellipsoid; aril is white and laciniate. 11 spp., all in northern South America, two up to Central America, two in Brazil, none endemics.

2. CLADE VIROLA (6/c.120) ‣ outsiders Endocomia (4, India, Sri Lanka, SE Asia, Malesia), Haematodendron (1; Madagascar).

2. Compsoneura (A. DC.) Warb. Trees dioecious; flowers ebracteolate; inflorescences 1-2 in leaf axills; reddish latex; differs from all the other Neotropical genera by having the tertiary venation close-parallel and perpendicular to the midrib; it is also unique in having its seed-coat black or purple patterned; stamens are few to many (4-10), the anthers adnate to column or free, fruits are ellipsoid, and the aril is red or white and entire (rarely laciniate). 18 spp. from Mexico and Central America to northern South America, 5 in Brazil, C. racemosa Ducke endemic. A unnamed species from W Colombia has the largest fruit in ths genus.

3. Iryanthera Warb. Dioecious, rarely monoecious; inflorescence facicle-racemose, in laef axils or in leafless shoots; 2-branched hairs, although the leaves appear to be glabrous; stamens are few (3-4(-6)), the anthers adnate to the column or free; fruits are broader than wide (transversally ellipsoid); aril is red and entire (or apically laciniate). 24 spp. from N South America (from Guianas to Peru and Bolivia), 21 in Brazil, three endemics, inc. I. campinae W.A. Rodrigues from Roraima state, a rare species in Brazil, by Plantas Raras do Brasil’s book. I. megistophylla A.C. Sm., from Choco region of W Colombia, has the largest leaves of Myristacaceae. A undescribed species also from W Colombia has the largest fruit in ths genus.

4. Osteophloeum Warb. Trees dioecious, up to 40 m tall; flowers bracteolate; fruit ellipsoid, straw-colored exudate, leaf apex distinctively round and petioles somewhat long; stellate indumentum is present; stamens are numerous (12(-20)), the anthers adnate to the column; fruits are broader than wide (transversally ellipsoid) having the aril entire. Two spp., O. platyspermum (A. DC.) Warb,. from over N South America (except Bolivia) and Panamá, and O. sulcatum Little from Ecuador.

5. Virola Aubl. Trees dioecious up to 40 m tall, inflorescences never ramiflorous or cauliflorous; stellate indumentum, also present in the Osteophloeum; leaves are usually oblong; the stamens are few (3(-4-6)), the anthers adnate to the column or apically free; fruits are globose to ellipsoid; aril is laciniate (as in Otoba) and red. 59 spp., Mexico, Central and South America, centerd in Amazon rainforest; 37 spp. in Brazil, 12 endemics, including V. bicuhyba (Schott ex Spreng.) Warb. from E country, considered some persons as endemic monotypic genus, Bicuhyba de Wilde, because of their inflorescence structure; two spp. from Amazonas state are rare plants in Brazil, by Plantas Raras do Brasil’s book.

MAGNOLIACEAE

§ FULLY CHLOROTROPHIC

Genera/species 2/360 Distribution southern India, Sri Lanka, E Himalaya, Assam, E Asia to Korean Peninsula and Japan, SE Asia, Malesia, New Guinea, New Britain, SE North America, Mexico, Central America, Caribbean, scattered areas in northern and central South America from Colombia to S Brazil. Habit usually bisexual (in Magnolia sect. Kmeria monoecious), evergreen or deciduous trees or shrubs. Only one genus in South America.

Key differences from similar families- the families below differ from Magnoliaceae in the following characters:

ü Myristicaceae: connate filaments, seeds arillate with reddish exudate in trunk, unisexual flowers

^üAnnonaceae: exstipulate, perianth of sepals and petals.

SYSTEMATIC outsider Liriodendron (2; E North America; China).

1. Magnolia L. Evergreen trees (up to 30 m tall in Brazilian Magnolia ovata) or shrubs, occasionally with indumentum of simple hairs; leaves alternate, spirally arranged, simple, petiolate, entire; flowers terminal, solitary, large deciduous ensheathing bracts leaving scars, actinomorphic, hermaphrodite, very rarely unisexual, showy, pedicellate; tepals numerous, petaloid, free, primarily spirally arranged in groups of three, sometimes outer whorl sepaloid; fruit follicles, woody, dehiscent; seed one or two, red fleshy exotesta, funicle long. 140 spp. in New World, 71 in South America, in Mexican (25), Brazil (4), Caribbean, Bolivia, Colombia (41), Ecuador (21), Guianas, Peru and Venezuela; three sections in New World:

§ sect. Talauma ‣ 93 spp. in four subsections (including all 68 South America species):

§ subsect. Talauma ‣ 59 spp. from 20° N in W and E Mexico and the Caribbean to 24º S beyond the Tropic of Capricorn in the Atlantic Forest of Brazil, in tropical rainforests from near sea level to circa 2,800 m, including 45 spp. in South America. M. ovata (A. St.-Hil.) Spreng., a Brazilian endemism, known as baguaçu, is the southermost distribution in this family, so far 33º S in Rio Grande do Sul state, common in Brazil Central Highlands; another species in Brazil are M. amazonica (Ducke) Govaerts, only in Amazon rainforest (also in Ecuador), M. brasiliensis C.O.Azevedo, A.F.P. Machado & J.A. Vázquez from S Bahia and N Minas Gerais states, and the extremally rare M. paranaensis J.A. Vázquez endemic to Paraná state.

§ subsect. Cubenses ‣ 10 spp. in the Antillean mountains.

§ subsect. Chocotalauma ‣ 3 spp. in Ecuador and 3 in Colombia.

§ subsect. Dugandiodendron ‣ with 22 spp., Andes from Colombia to Peru, and the Guiana Shield in Venezuela.

§ sect. Macrophylla ‣ six species, three in the tropics of Mexico and three in temperate areas of the SE U.S.A. and NE Mexico.

§ sect. Magnolia ‣ 27 spp., usually at middle elevations in the mountains of Mexico and Central America and only two species of the SE U.S.A., one of them also occurring in Cuba.

ANNONACEAE

§ FULLY CHLOROTROPHIC

Genera/species 108/c. 2,200–2,400 Habit usually bisexual (sometimes dioecious, rarely monoecious), evergreen or deciduous trees, shrubs or lianas. Often aromatic. None sp. is a truly herbaceous or annual.

A pantropical family, with two genera in temperate North America (Asimina, Deeringothamnus near Great Lakes, closed related sometimes uniteed only in first). Only three genera in New World occur outside: Xylopia from Asia up to Fiji, Anaxagorea occur in Asia and three spp. in Annona occur in Africa. Only one neotropical sp. (Annona glabra L.) reaches up Africa. In South America has also xylopodial members in this family (Duguetia). Lianas occur almost exclusively in Asia and Africa, and tall trees occur in over tropics, of which in Guatheria and Xylopia in South America.

All genera are native in the Neotropics. Several species are cultivated all over the Neotropics because of their edible fruits: some species of Annona, ex. A. muricata L., A. cherimola Miller, A. mucosa Jacq., and A. reticulata L. Young stems of Annonaceae are often used as fishing rods.

826 spp. in New World, 669 in South America, 375 in Brazil.

SYSTEMATIC 5 subfamilies, all in South America except Meiocarpidioideae (1; tropical W Africa).

KEY TO GENERA OF NEOTROPICAL ANNONACEAE

1. Leaves spirally arranged; flowers 4(-6)-merous ------------ Tetrameranthus

1. Leaves arranged in two rows along the stem (distichous); flowers 3-merous (very rarely 2-merous) - 2.

2. Indumentum composed of stellate or scale-like hairs; fruit pseudo-syncarpous and carpels strongly coherent, but only partly connate ------------ Duguetia

2. Indumentum composed of simple hairs or lacking (hairs sometimes stellate in Annona and Anaxagorea); fruit either apocarpous or syncarpous - 3.

3. Flowers and fruit axillary (sometimes leaf-opposed in Anaxagorea brevipes) or sometimes arising from the trunk (cauliflorous); fruit apocarpous - 4.

4. Upper leaf surface with impressed midrib - 5.

5. Petals often thick and fleshy (2-10 mm thick) and covered with microscopic, brownish hairs ------------ Anaxagorea

5. Petals generally much thinner and covered with much larger hairs or glabrous - 6.

6. Pedicels with a supra-basal articulation ------------ Guatteria

6. Pedicels with a basal articulation - 7.

7. Sepals 2; petals 4; carpels 1(-2) ------------ Tridimeris

7. Sepals 3; petals 6; carpels generally numerous - 8.

8. Monocarps dehiscent, seeds distinctly arillate ------------ Xylopia

8. Monocarps indehiscent, aril absent or indistinct - 9.

9. Monocarps several-seeded - 10.

10. Petals often maroon, distinctly veined ------------ Asimina

10. Petals mostly white, not veined ------------ Diclinanona

9. Monocarps 1-seeded - 11.

11. Upper leaf surface with distinctly impressed venation ------------ Ephedranthus

11. Upper leaf surface with flat, not or slightly impressed venation - 12.

12. Pedicels with 3-6 bracts ------------ Oxandra

12. Pedicels with 1-2 bracts - 13.

13. Pedicels with 1 bract above the articulation ------------ Pseudomalmea

13. Pedicels without a bract above the articulation ------------ Klarobelia

4. Upper leaf surface with raised midrib - 14.

14. Petals often fleshy (2-10 mm thick) and covered with microscopic, brownish hairs ------------ Anaxagorea

14. Petals generally much thinner and covered with much larger hairs or glabrous - 15.

15. Leaves with distinct marginal vein, almost touching the margin (except in P. espirito-santensis) ------------ Pseudoxandra

15. Leaves without a marginal vein (except in Oxandra p.p., but then much further removed from the margin) - 16.

16. Lower leaf surface glaucous; petals 25-35 mm long. western part of South America ------------ Ruizodendron

16. Lower leaf surface not glaucous; petals 4-20 mm long - 17.

17. Petals 4-8.5 mm long, with a small, incurved, tail-like, apical appendage ------------ Onychopetalum

17. Petals 2.5-20 mm long, without an apical appendage - 18.

18. Leaves asymmetrical; petals 2.5-4.5 mm long ------------ Bocageopsis

18. Leaves symmetrical; petals mostly more than 5 mm long - 19.

19. Upper leaf surface with a distinctly grooved midrib ------------ Cremastosperma

19. Upper leaf surface with a non-grooved midrib - 20.

20. Petals 10-15 mm long; seeds 25-30 mm long ------------ Pseudephedranthus

20. Petals 4-10 mm long; seeds 8-20 mm long - 21.

21. Inflorescences often 1-flowered ------------ Oxandra

21. Inflorescences often branched ------------ Unonopsis

3. Flowers and fruit terminal, leaf-opposed, or supra-axillary, rarely arising from the trunk; fruit apocarpous or syncarpous - 22.

22. Bracts absent - 23.

23. Flowers pendent on long (40-60 mm) pedicels ------------ Cymbopetalum

23. Flowers non-pendent and pedicels much shorter - 24.

24. Leaves asymmetrical; monocarps 20-90 x 30-40 mm, wall 2.5-4 mm thick ------------ Porcelia

24. Leaves symmetrical; monocarps smaller, wall thinner - 25.

25. Petals basally connate - 26.

26. Petals basally connate into a 2-4 mm long tube; monocarps dehiscent, falciform ------------ Cardiopetalum

26. Petals free; monocarps indehiscent, fusiform to linear, ------------ Froesiodendron

25. Petals free - 27.

27. Leaves tri-plinerved at the base; monocarps 3-9; stamens c. 40 ------------ Trigynaea

27. Leaves not tri-plinerved at the base; monocarps 1-3; stamens fewer 20 - 28.

28. Flower buds conical; petals linear, recurved to patent; calyx without distinct lobes ------------ Hornschuchia

28. Flower buds globose; petals ovate to elliptic, erect; calyx distinctly 3-lobed ------------ Bocagea

22. Bracts present - 29.

29. Upper leaf surface with raised midrib - 30.

30. Fruit apocarpous, the monocarps distinctly stipitate; seeds pitted ------------ Mosannona

30. Fruit pseudo-syncarpous, the carpels strongly coherent; seeds smooth ------------ Duckeanthus

29. Upper leaf surface with impressed midrib - 31.

31. Flowers winged; fruit syncarpous (very rarely apocarpous) ------------ Annona p.p.

31. Flowers not winged; fruit apocarpous or syncarpous - 32.

32. Outermost stamens staminodial; fruit provided with an almost woody basal collar ------------ Fusaea

32. All stamens fertile; fruit without a basal collar - 33.

33. Fruit syncarpous, each carpel 1-seeded; petals often thick and fleshy ------------ Annona

33. Fruit apocarpous, each monocarp 1-several-seeded; petals thin - 34.

34. Pedicels often provided with 1 leafy bract ------------ Desmopsis

34. Pedicels without leafy bracts - 35.

35. Monocarps 1-seeded; petals yellow to cream, margins ciliate ------------ Malmea

35. Monocarps 1-several-seeded; petals red, purple, brown, or rarely cream, not ciliate - 36.

36. Petals distinctly veined, seeds with lamellate rumination ------------ Sapranthus

36. Petals not distinctly veined, seeds with spiniform rumination ------------ Stenanona

1. SUBFAMILY ANAXAGOREOIDEAE (1/c 30) ‣ a single genus.

1. Anaxagorea A. St-Hill. Trees or shrubs with simple hairs or stellate hairs in floral structures. 22 spp. in tropical America, except Caribbean; 20 in South America, 14 in Brazil, 12 only Amazon rainforest (none endemics), A. silvatica R.E. Fr. endemic to Atlantic Forest, and A. dolichocarpa Sprague & Sandwith widely distributed, and 4 in Sri Lanka to W Malesia and Moluccas, unique by Asia/America range.

2. SUBFAMILY AMBAVIOIDEAE (9/52) ‣ two tribes, Canangeae (4/36, tropical E Africa, tropical Asia to tropical Australia and Fiji) do not occur in South America; among Tetramerantheae, outsiders are Mezzettia (3; the Malay Peninsula to Moluccas), Ambavia (2; Madagascar), Cleistopholis (4; tropical Africa).

2. Tetrameranthus R.E. Fries. Small trees, leaves spirally arranged (unique among Annonaceae), often concentrated towards the end of the branches, primary vein impressed to slightly raised on the upper side; inflorescences axillary, 1-flowered to several-flowered and umbel-like, bracts up to 4 below the articulation; flowers bisexual, 4-merous or less often 5–6-merous, white to yellow or cream. 7 spp., Amazon rainforest of Venezuela, Colombia, Brazil (4, none endemic), Peru, and Ecuador, but also in the Colombian state of Choco and in French Guiana and neighboring Amapá, Brazil, and Amazon rainforest and French Guiana in the north and the Pacific coast of Colombia in the west.

3. SUBFAMILY ANNONOIDEAE (48-49/c. 1,400) ‣ the tribes Monodoreae (11/c 90, tropical and S Africa, Madagascar) and Uvarieae (14/465–470, tropical regions in the Old World) do not occur in South America.

3.1 ANNONOIDEAE ▸ TRIBE BOCAGEEAE (8/65) ‣ outsider Mkilua (1; Kenya, Tanzania).

3. Bocagea A.St.-Hil. Small trees or shrubs with simple hairs. 4 spp., both very narrow endemics, very rare, in fewer seven collections, possibly extincts, from Bahia, Espírito Santo, São Paulo and Rio de Janeiro states in SE in Brazil.

4. Cardiopetalum Schltdl. Small trees with simple hairs. Three spp., all Brazil (one endemic), from southwards Amazon rainforest and eastwards Madeira river to Minas Gerais, Tocantins, and SE Piauí, one up to Suriname and French Guiana, another up to Bolivia.

5. Cymbopetalum Benth. Small trees or shrubs with simple hairs. 27 spp., all over the Neotropics (15 in South America), S Mexico to Peru and Brazil (only 5 spp., 3 endemics).

6. Froesiodendron R.E. Fr. Trees or shrubs with simple hairs; flowers axillary or supraxillary. Three spp. from Colombia, Peru and Acre state in Brazil (only one, no endemic).

7. Hornschuchia Nees. Trees or shrubs with simple hairs (0.5 to 8m tall); morphologically variable genus, often cauliflorous; flower buds conical; petals linear, recurved to patent; calyx cup-shaped, without distinct lobes; flowers solitary or in many-flowered inflorescences; inflorescences often flagellate and produced from the main stem, axillary or terminal. 14 spp. endemics to Atlantic Forest and adjacent areas from Pernambuco to Rio de Janeiro state, centered in S Bahia.

8. Porcelia Ruiz & Pavon. Trees with simple hairs; flowers bisexual; very large, many-seeded monocarps, reaching a length of up to 13 cm. 7 spp., 6 in tropical South America and one in Costa Rica in the north; only two in Brazil, one endemic.

9. Trigynaea Schltdl. Trees or shrubs with simple hairs. 9 spp., over tropical South America; 4 in Brazil, T. duckei (R.E. Fr.) R.E. Fr. from Ecuador to Bolivia and Rondonia, N Colombia and Venezuela; Guyana to Piauí, and small area in Minas to São Paulo, in SE Brazil, and remaining three endemics.

3.2 ANNONOIDEAE ▸ TRIBE XYLOPIEAE (2/260) ‣ outsider Artabotrys (102; tropical regions in the Old World to tropical Australia).

10. Xylopia L. Trees or shrubs with simple hairs; flowers bisexual, solitary or in fascicle. 160 spp., the only genus in the family occurring in Africa (70), Asia and Oceania (40) and America (63 spp., 53 in South America, 36 in Brazil, 16 endemics, nine in Atlantic Forest); X. atlantica Mello-Silva & J.C.Lopes, endemic to Bahia, has leaves up to 41 cm long, larger than those of any other Brazilian Xylopia.

3.3 ANNONOIDEAE ▸ TRIBE GUATTERIEAE (1/185) ‣ a single genus.

11. Guatteria Ruiz & Pav. Trees or shrubs, only two remarkable lianas; axillary flowers with a pedicel which has a suprabasal articulation; inflorescences often cauliflorous. 183 spp., all over the Neotropics, the largest genus in this area, 159 in South America, some in shrubby areas in dry regions in Brazilian savannas (possibly only three spp. from Minas Gerais and Goias states in Brazil); 73 spp. in Brazil, 33 endemics; 4 spp. of Guatteria, three in Rio de Janeiro state and one in Bahia state (all not recognized by VPA) are rare species in Brazil, by Plantas Raras do Brasil’s book.

3.4 ANNONOIDEAE ▸ TRIBE ANNONEAE (7–8/c 325) ‣ outsiders Anonidium (5; tropical Africa), Neostenanthera (4; tropical Africa, tropical South America), Goniothalamus (c 135; India, Sri Lanka, SE Asia, W Malesia, New Caledonia), Disepalum (9; SE Asia, W Malesia), Asimina (17; U.S.A.), Deeringothamnus (2; Florida).

12. Annona L. Trees or shrubs, sometimes up to 40 m tall, sometimes with woody rhizomes or xylopodium; flowers often cauliflorous 166 spp., all over the Neotropics, three also occurring in Africa, A. glabra L. also in North America; 125 spp. in South America, 80 in Brazil, 25 endemics; it occur all over the Neotropics, also in savannas of C Brazil (cerrado), where most other genera of Annonaceae are lacking; A. saffordiana R.E. Fr. from Minas Gerais state is a rare species in Brazil, by Plantas Raras do Brasil’s book. A.warmingiana Mello-Silva & Pirani has an underground stem and is a smallest of all Magnoliales; this species occur in dry savannas of C Brazil.

13. Diclinanona Diels. Small tres with simple hairs, lowers polygamous. Three spp., restricted of Amazon rainforests of Guianas, E Peru, SE Colombia, S Venezuela and adjacent areas in W Brazil (all species, one endemic).

3.5 ANNONOIDEAE ▸ TRIBE DUGUETIEAE (5/100) ‣ outsiders Pseudartabotrys (1; tropical W Africa), Letestudoxa (3; tropical W and C Africa).

14. Duckeanthus R.E.Fr. Trees with simple hairs; flowers bisexual. Only one sp., D. grandiflorus R. E. Fries, endemic to Guiana Shield from Amazon rainforest of Brazil (Upper Rio Negro), in elevation ranges 100 m.

15. Duguetia A. St. Hill. Trees or shrubs with stellate or lepidote indumenta, sometimes with xylopodium; inflorescences often cauliflorous. 91 spp. from Costa Rica in the North to Bolivia and Paraguay in the South (89 in South America, 67 in Brazil, 31 endemics), and 4 spp. of this genus which were formerly placed in Pachypodanthium Engl. & Diels, which occur in Africa; two spp. from Tocantins and Bahia states are considered rare species in Brazil, by Plantas Raras do Brasil’s book.

16. Fusaea Saff. Small tres or shrubs with simple hairs. Three spp., widely distributed in Amazon rainforest of Guyana to Peru and N Brazil (two spp., none endemics).

4. SUBFAMILY MALMEOIDEAE (47/c. 730) ‣ tribes Piptostigmateae (6/35, tropical Africa), Maasieae (1/6, SE Asia, Malesia), Monocarpieae (1/6, SE Asia, W Malesia), Fenerivieae (1/9–10, Madagascar), Phoenicantheae (1/2, Sri Lanka) and Dendrokingstonieae (1/3, Thailand, the Malay Peninsula, Sumatra) do not occur in South America.

4.1 MALMEOIDEAE ▸ TRIBE MILIUSEAE (23/490–495) - outsiders Mitrephora (47; SE Asia, W Malesia), Platymitra (2; W Malesia), Orophea (c 50; S India, Hainan, SE Asia, Malesia to Moluccas), Trivalvaria (6; Assam to W Malesia), Pseuduvaria (57; India, Sri Lanka, SE Asia, Malesia to New Guinea and tropical Australia), Popovia (26; tropical Asia to tropical Australia), Huberanthus (27; tropical E Africa, Madagascar, the Comoros, S India, Sri Lanka, SE Asia to New Guinea, Queensland, New Caledonia, Fiji), Alphonsea (25; tropical Asia to S China and Malesia), Miliusa (c 50; Hainan and Malesia to Queensland), Wangia (1; Yunnan), Phaeanthus (9; India, Sri Lanka, SE Asia, Malesia), Sageraea (9; India, Sri Lanka, SE Asia to Philippines), Stelechocarpus (1; Java), Winitia (2; Vietnam, Thailand, the Malay Peninsula, Borneo), Polyalthia (45–50; tropical regions in the Old World to Queensland), Marsypopetalum (6; Hainan, SE Asia, W Malesia), Neouvaria (7; W Malesia), Monoon (56; India, Sri Lanka, Ryukyu Islands, Hainan, SE Asia, W Malesia, New Guinea, N Australia), Meiogyne (24; SW India, Hainan, SE Asia, Malesia to New Guinea, Queensland, New Caledonia, Micronesia, Polynesia), Wuodendron (1; Assam and Yunnan to Vietnam and Cambodia), Tridimeris (2; Mexico).

New World members composes Sapranthinae subtribe.

17. Desmopsis Saff. Trees or shrubs with simple hairs; flowers bisexual, few flowered. 15 spp., 13 from Mexico to Panamá, two of then up to Colombia, one endemic to Colombia, and one endemic to Cuba.

18. Sapranthus Seem. 6 spp., 5 from Mexico to Central America, and S. isae J.G. Vélez & Cogollo endemic to Caribbean coast of Colombia.

19. Stenanona Standl. 11 spp., 10 from Mexico to Central America, and S. narinensis G.E. Schatz & Maas endemic to Colombia.

4.2 MALMEOIDEAE ▸ TRIBE MALMEEAE (13/178) ‣ all genera occur in South America.

20. Bocageopsis R.E. Fr. Trees with simple hairs. 4 spp. in Mato Grosso and Pará states in N Brazil, and Guyana, in tropical South America, none is a national endemic.

21. Cremastosperma R.E. Fr. Trees with simple hairs; flowers bisexual. 34 spp., three in Costa Rica and Panamá, and remaining 31 in South America, to Bolivia in the south, eastwards up to the Guianas, and S of the Amazon River; only 4 in Brazil, none endemics.

22. Ephedranthus S. Moore. Trees or shrubs with simple hairs. 7 spp., most species are restricted to rainforest habitats in Amazon rainforests; 5 spp. in Brazil (3 endemics); two occur in riparian forest in savannas of C Brazil (cerrado) and the transition zone between savannas and Amazonian rainforest in NE Brazil; E. dimerus J. C. Lopes, Chatrou & Mello-Silva, is the only species of the genus from the Atlantic Forest in Bahia, Minas Gerais and Espirito Santo states, and the only species of the genus with dimerous flowers.

23. Klarobelia Chatrou. 12 spp. from tropical South America (11) to Costa Rica in the north; 4 spp. in Brazil, one endemic, the remaining in adjacent Colombia and Bolivia.

24. Malmea R.E.Fr. Trees or shrubs with simple hairs. 6 spp., 5 in Amazon rainforests in over N South America, M. manausensis Maas & Miralha of then in Brazil, and the sixth M. obovata R.E. Fr., very narrow endemic and a rare plant in Brazil, by Plantas Raras do Brasil’s book, in Atlantic Forest of Bahia state.

25. Mosannona Chatrou. 14 spp. from Mexico to Peru and Brazil, 10 spp. in South America, also Guianas; in Brazil only M. raimondii (Diels) Chatrou, shared with Peru.

26. Onychopetalum R.E. Fr. Trees with simple hairs. Two spp. from Brazil, both reaching into Venezuela, Bolivia and Peru.

27. Oxandra A.Rich. Trees or shrubs with simple hairs; flowers bisexual, in solitary or in few flowered fascicles, axillary. 27 spp., in over the Neotropics, 23 in South America, mainly Amazon rainforest, 14 in Brazil, 5 endemics, including the only 4 in Atlantic Forest.

Oxandra is not monophyletic; all sampled species of this genus belong to the tribe Malmeeae, subfamily Malmeoideae, but they emerge in two different clades. These clades seem to be related to Ephedranthus S.Moore, Ruizodendron ovale, Pseudomalmea Chatrou, Klarobelia Chatrou and, Pseudephedranthus Aristeguieta, although there is no strong support for the phylogenetic relationships; moreover, the exact placement of O. venezuelana R.E.Fries within Malmeeae is not known.

28. Pseudephedranthus Aristeg. Trees; young twigs glabrous; inflorescences axillary, 1–4-flowered with 2nd order flowers originating from axils of lower bracts (or possibly also through accessory buds); flowers actinomorphic, bisexual or staminate (androdioecious), 3-merous, perianth consisting of one whorl of sepals and two whorls of petals; fruit apocarpous, composed of few, indehiscent monocarps; seed 1, pale brown, rumination lamellate in 2–4 parts, raphe a distinct groove. Two spp., endemic to Guiana Shield in Brazil (both, none endemics), one up to Guyana and S Suriname, another up to Venezuela, in elevation ranges 100 – 200 m.

29. Pseudomalmea Chatrou. Three spp. from forests of Venezuela, Colombia, Ecuador, Peru, Bolivia and N Brazil (only the widely P. diclina (R.E. Fr.) Chatrou).

30. Pseudoxandra R.E.Fr. Trees or shrubs with simple hairs; flowers bisexual, in solitary or in few flowered fascicles, axillary. 24 spp., all confined in tropical South America, 14 in Brazil, 9 endemics.

31. Ruizodendron R.E. Fr. Trees, often with buttresses; leaves distichous, simple, entire, shortly petiolate; lamina medium-sized, mostly elliptic; flowers actinomorphic, bisexual, perianth consisting of one whorl of sepals and two whorls of petals; fruit apocarpous, composed of few indehiscent monocarps, these transversely ellipsoid, the stipe laterally attached; seed 1, filling the whole monocarp, orange-brown, rumination lamellate in 2 parts, raphe a fine, shallow groove. Only one sp., R. ovale (Ruiz & Pav.) R.E. Fr., from Colombia, Ecuador, E Peru, Bolivia and Acre state in N Brazil.

32. Unonopsis R.E. Fr. Trees with simple hairs; sepals 3, valvate. 52 spp., from Oaxaca to S Brazil and Caribbean; 41 spp. in South America, only 16 in Brazil (8 endemics).

7. LAURALES

FAMILIES ABSENTS IN SOUTH AMERICA: CALYCANTHACEAE (3/11).

LINEAGE 1 of 2: SIPARUNOIDS

SIPARUNACEAE

§ FULLY CHLOROTROPHIC

Genera /species 2/54 Distribution Siparuna: southern Mexico, Central America, tropical South America, Caribbean; Glossocalyx (G. longicuspis): W Cameroon and Gabon. Habit monoecious or (secondarily) dioecious, evergreen trees or shrubs (sometimes lianas). Often aromatic. 20 spp. in Brazil. In lowland and montane forests to 3,800 m elev. This is a Neotropical family with one other genus (and species) species in Africa (Glossocalyx longicuspis Benth.).

Entire plant has a lemon odour, especially when crushed (but not in herbarium material); rarely confused except Siparuna decipiens DC., which is often mistaken for Pera benensis Rusby (Euphorbiaceae).

SYSTEMATIC outsider Glossocalyx (1; tropical W and Central Africa).

1. Siparuna Aubl. Dioecious or monoecious shrubs, treelets, or trees (to 40 m tall and to 120 cm in dbh); leaves opposite, decussate or in whorls of 3, 4, or 6, petiolate, sparsely or densely pubescent; inflorescences mono- or dichasial cymes, sometimes bifid or fasciculate, axillary and/or on leafless nodes; flowers unisexual, pedicellate; mature fruiting receptacles fleshy and 1.5-4 cm in diam., globose, smooth, spiny, or with tubercles, when fresh and mature reddish or yellow. 80 spp. from tropical Mexico south through Central America, the Lesser Antilles, Trinidad, and northern South America to Bolivia and Paraguay; 20 in Brazil, 4 endemics; only S. glabrescens (C. Presl) A. DC. from Caribbean does not occur in South America. Some species in South America (inc. Brazil) are myrmecophytes.

GOMORTEGACEAE

§ FULLY CHLOROTROPHIC

Genera/species 1/1 Distribution central Chile. Habit bisexual, evergreen tree. Aromatic. Young branches quadrangular in transverse section. Use fruits, timber.

SYSTEMATIC a single genus

1. Gomortega Ruiz & Pav. Trees, bisexual, evergreen; aromatic; young branches quadrangular in transverse section. Only one sp., G. keuke (Molina) Baill., endemic to central Chile.

ATHEROSPERMATACEAE

§ FULLY CHLOROTROPHIC

Genera /species 6/16 Distribution New Guinea and Queensland, New Caledonia, E and SE Australia, Tasmania, New Zealand, central Chile and W Patagonia. Habit evergreen trees (rarely shrubs). Aromatic. Use tea (Atherosperma moschatus), fruits (‘Laurelia’), timber, carpentry.

SYSTEMATIC Daphnandra (6; Queensland, New South Wales, Tasmania), Doryphora (4; Queensland, New South Wales, New Caledonia, Tasmania), Atherosperma (1; New South Wales, Victoria, Tasmania), Nemuaron (1; New Caledonia), Dryadodaphne (3; New Guinea, Queensland).

1. Laurelia Juss. Two spp., L. sempervirens (Ruiz & Pav.) Tul. in Chile and L. novae-zelandiae A. Cunn. in New Zealand.

2. Laureliopsis Schodde. Only one sp., L. philippiana (Looser) Schodde, restricted to Chile and patagonian Argentina.

LINEAGE 2 of 2: LAURIDS

HERNANDIACEAE

§ FULLY CHLOROTROPHIC

Genera/species 4/70 Distribution Hernandioideae: pantropical, with the largest diversity in Madagascar and in SE Asia and Malesia; Gyrocarpoideae: tropical and subtropical regions, particularly in coastal areas and also on oceanic islands: E and S India, Sri Lanka, Indochina, parts of Malesia, northern Australia, with the highest diversity in Central America and South America. Habit bisexual, monoecious, andromonoecious, polygamomonoecious, or dioecious, evergreen trees, shrubs or lianas. All aromatic.

Key differences from similar families

ü In Lauraceae, the anthers dehisce via hinged flaps and the fruits are usually subtended by a cupule (thickened receptacle).

ü In Monimiaceae, the leaves are opposite (or rarely whorled), the ovary is apocarpous and superior.

SYSTEMATIC Illigera has 18 spp. in tropical regions in the Old World.

1. SUBFAMILY GYROCARPOIDEAE (2/14) ‣ both genera in South America.

1. Gyrocarpus Jacq. Dioecious trees; leaves entire or 3-5 lobed; inflorescence axillary, dichasial, ebracteate; flowers polygamous, with 4-8 tepals, stamens 4-5 with dorsal appendages, fruits with two large wings (samaroid). 4-5 spp. worldwide, three in Mexico and Central America with G. americanus Jacq. reaching into Colombia and Venezuela.

2. Sparattanthelium Mart. Shrubs or lianas scadent with recurving hooks; leaves simple, mostly triplinerved; inflorescence dichasial, ebracteate, flowers perfect, 4-5 (-8) –merous, bisexual, tepals 4-5(-8), stamens 4-5(-7) without appendages, fruits without wings and not subtended by bracteoles (nucular). 12 spp. of tropical South America, most of which rare, 10 occur in Brazil (exceptions are two spp. restricted from Guyana and Venezuela), S. borororum Mart. endemic.

2. SUBFAMILY HERNANDIOIDEAE (2/44) ‣ outsider Illigera (18; tropical regions in the Old World).

3. Hernandia L. Evergreen monoecious trees with simple leaves; thyrses with long penduncles and 3-flowered cymes with 2 male and 1 female flower, male flowers with 3-5 stamens with basal appendages, fruits subtended by bracteoles or a cupule formed by them (nucular). 25 spp. of mainly evergreen, forest trees; 10 are neotropical, and half of these are restricted in Caribbean; 4 reaches in South America: H. guianensis in Venezuela, Brazil (Pará and Amazonas) and Guyana, two from Mexico to Ecuador, and one endemic to Ecuador.

MONIMIACEAE

§ FULLY CHLOROTROPHIC

Genera /species 27/c. 280 Distribution tropical and subtropical regions in Central and South America, southern South America (Patagonia), Central and E Africa, Sri Lanka, Malesia, E Australia, New Caledonia and other islands in Melanesia, New Zealand, with the largest diversity in Malesia, on islands in the SW Indian Ocean, in Madagascar, and in South America. Habit evergreen scambling shrubs, treelets or trees, with spherical oil cells in all parts of the plants.

Currently 28 genera and 195-200 species, mostly in the Malagasy floristic region, New Guinea (this has 75 species), and the Neotropics (six genera and 28 spp., four of then genera are endemic to Brazil and adjacent Argentina; one is widely distributed, and another occurs disjunct in central Chile), with a few species in the rest of Malesia, E Australia, and New Zealand. A single species, Xymalos monospora Baill., occurs from South Africa to Kenya, Uganda, and Cameroon. 5 genera and about 40 spp. in Brazil; All the neotropical genera, except Peumus, and more than half the known species (many of which are endemic to only small areas) are found in the Atlantic forest biome.

SYSTEMATIC three clades, Hortonioidea (1/3, Sri Lanka) does not occur in South America.

1. SUBFAMILY MONIMIOIDEAE (3/19) ‣ outsiders Monimia (3; Mauritius, Réunion), Palmeria (15–20; New Guinea, E Queensland, New South Wales, Victoria).

1. Peumus Molina. Small trees, dioecious, sometimes with lignotuber. Only one sp., P. boldus Molina, in sclerophyllous forests of C & S Chile.

2. SUBFAMILY MOLLINEDIOIDEAE (18/180) ‣ outsiders Lauterbachia (1; New Guinea), Austromatthaea (1; Queensland), Decarydendron (4; Madagascar), Ephippiandra (7; Madagascar), Faika (1; New Guinea), Hedycarya (c 30; Sulawesi to New Guinea, Queensland, New South Wales, Victoria, King Island, New Caledonia, Fiji, New Zealand), Kibaropsis (1; New Caledonia), Levieria (7; Sulawesi to Queensland), Kairoa (2; Papua New Guinea), Kibara (43; Nicobar Islands, Malesia to tropical Australia), Matthaea (6; Malesia), Parakibara (1; Moluccas), Steganthera (c 30; E Malesia to New Guinea and Solomon Islands, Queensland), Tambourissa (18; Madagascar, Mascarene Islands), Tetrasynandra (3; Queensland), Wilkiea (8; Papua New Guinea, Queensland, New South Wales), Xymalos (1; E and S Africa).

2. Grazielanthus Peixoto & Per-Moura. Shrub semi-decumbent, 2 – 4 m tall, dioecious, male inflorescence in axillary tri-floral dichasium. Only one sp., G. arkeocarpus Peixoto & Pereira- Moura, very narrow endemic - a rare plants in Brazil, by Plantas Raras do Brasil’s book - known only from the central region of the coastal plain in the State of Rio de Janeiro, Brazil, within the sub-canopy of lowland forests, on alluvial soils that are humid or flooded during the rainy season; only 32 adult individuals at their deiscovery.

3. Hennecartia Poisson. Small tree, leaves often verticillate. Only one sp., H. omphalandra J. Poiss., gallery forests of NE Argentina and S Brazil, also in Uruguay

4. Macropeplus Perkins. Dioecious shrubs or treelets (1.5 - 15m tall), stems cylindrical, much branched; leaves opposite, simple, slightly variable in same branche; inflorescence male as trichasia, female as monochasia, mainly axillar, flowers actinomorphic, 4-merous. 4 spp., three restricted to mountain ranges of Atlantic Forest of SE region of Brazil (two very narrow endemics, both rare plants in Brazil, by Plantas Raras do Brasil’s book), and M. ligustrinus (Tul.) Perkins ranging in savannas of C Brazil (cerrado) and Atlantic Forests up to mountains of C Bahia, Goias and Distrito Federal.

5. Macrotorus Perkins. A dioecious shrub; female flowers not known. Two species, restricted to lowland, submontane and montane forests of the Brazilian Atlantic coast in the states of Bahia, Espírito Santo, Rio de Janeiro and São Paulo.

6. Mollinedia Ruiz & Pav. Staminate flower with nearly rounded buds, perianth lobes with ratio about 1:1 in relation to the length of the receptacle, stamens ovate or rounded, locules either with two longitudinal openings and an extended connective or confluent at the apex, making the anther horseshoe-shaped. 69 spp., in understorey lowland moist forest, from Mexico and Central America to South America (60), to the Amazon rainforest and in gallery forests in Atlantic Forests of Brazil (42, 39 endemics) and Paraguay (absent in Argentina); 5 spp. (4 in Rio de Janeiro, and one in Santa Catarina state) are rare plants in Brazil, by Plantas Raras do Brasil’s book. Two sections:

§ sect. Appendiculatae ‣ c. 38 spp., ove rrange of genus.

§ sect. Exappendiculatae ‣ c. 25 spp., Colombia, Peru and SE Brazil.

LAURACEAE

§ PARASITIC (Prosopanche – Cassytha – Bdalophytum – Krameria - ... – Lennoa – OROBANCHACEAE - Cuscuta)

Genera/species 56/2,500-3,500. Distribution tropical and subtropical regions, with the largest diversity in SE Asia and tropical America, and some genera in warm-temperate areas. Habit bisexual, monoecious, polygamomonoecious or dioecious, usually evergreen (rarely deciduous) trees or shrubs, all woody (except Cassytha). Aromatic. Cassytha consists of parasitic, climbing herbs with more or less photosynthesizing stems. A few spp. are subscandent.

Seven genera in South America also occur in Old World: Caryodaphnopsis, Cinnamomum, Cassytha, Beilschmeidia, Lindera, Ocotea and Cryptocarya. The avocado, Persea americana Mill., is widely cultivated, and this fruit in many popular in World; its native Mesoamerican region in Mexico to El Salvador; several Lauraceae are important sources of timber, e.g. Chlorocardium (Greenheart). One species, Dicypellium caryophyllaceum (Mart.) Nees, has been heavily exploited because of its aromatic bark, known as pau-cravo; many with pleasantly aromatic yellow wood (pau-rosa oil is distilled from Aniba rosaeodora Ducke).

Key differences from similar families

ü the hemiparasitic Cassytha is often confused from Cuscuta L. (Convolulaceae) and flowers must be consulted to distinguish the two; the Lauraceae are otherwise not easily confused with other families.

ü valvate anthers occur also in Berberidaceae, which are small spiny shrubs of high altitudes in the Neotropics.

SYSTEMATIC six tribes; Hypodaphnideae (1/1, W Africa, H. zenkeri (Engl.) Stapf, only member of Lauraceae with an inferior ovary) and Neocinammomeae (1/7, tropical Asia) do not occur in South America.

1.1 TRIBE CRYPTOCARYEAE (13/c. 770) - outsiders Aspidostemon (ca. 30 spp. in Madagascar), Dahlgrenodendron (1; South Africa), Eusideroxylon (1; Greater Sunda Islands), Potoxylon (1;Greater Sunda Islands), Ravensara (10-20 spp. in Madagascar and Comoro Islands), Endiandra (ca. 100 spp.; SE Asia, E Australia, and the W Pacific Islands), Triadodaphne (1; Papua New Guinea), Potameia (3-10 spp. in Madagascar), Sinopora (1; China), and Syndiclis (10-20 spp. in India and SE Asia).

1. Beilschmiedia Nees. Leaves alternate to opposite; the fruit lacks a cupule. 250 spp., 31 from Mexico to southern Brazil and in the Caribbean; five groups in Neotropics, the Curviramea group consists of nine species, B. curviramea (Meisn.) Kosterm., from Guyana and E Venezuela, and the eight Brazilian species, all endemics, one of then, from Rio de Janeiro state, is a rare plants in Brazil, by Plantas Raras do Brasil’s book; 19 spp. in South America.

2. Cryptocarya R.Br. Trees, sometimes with lignotuber; fruit almost completely enclosed in the cupule, which has only a small, apical pore. 350 spp., pantropical, 17 spp. in New World, mostly in S Brazil (13, 11 endemics) to Chile, but also known from French Guyana and adjacent Brazil, Andean Venezuela, Colombia, Ecuador, and Peru, only one not occur in South America; two spp. from Minas Gerais and Espírito Santo states are rare plants in Brazil, by Plantas Raras do Brasil’s book.

3. Yasunia van der Werff. Trees, leaves opposite, subopposite, or alternate; inflorescences in axils of leaves, paniculate; flowers subsessile, clustered, when 3 flowers terminate an inflorescence branchlet, the 2 lateral flowers not strictly opposite; the only neotropical spp. of Lauraceae with dimerous flowers. Two spp., Y. quadrata van der Werff known only from the type collection made in E Peru near frontier with Brazil and Y. sessiliflora van der Werff from Ecuador.

1.2 TRIBE CASSYTHEAE (1/20) ‣ a single genus.

4. Cassytha Osb. ex L. Vines with scale leaves, parasitic. 17 spp., pantropical, more species occur in Australia and South Africa; with C. filiformis L. widely distributed, from Mexico to Brazil, apparently absent in Central America and Peru southwards in Andes, often found near the coast, but also locally common in Mauritia swamps.

Diference between Cuscuta (Convolvulaceae) and Cassytha: Cassytha has stems with fine longitudinal rugae or ridges, sometimes with trichomes; Cuscuta has smooth stems without trichomes; among fflowers, Cassytha has 3-merous with perianth elements free; Cuscuta has 4-5-merous (rarely 3-merous), with perianth elements fused.

1.3 TRIBE CARYODAPHNOPSIDEAE (1/8)

5. Caryodaphnopsis Airy Shaw. Strongly unequal tepals and opposite leaves; fruits in neotropical species are round or pearshaped, and seated on a naked pedicel. 8 spp., one only in Central America, 7 remainings from forests of Colombia to Bolivia, C. inaequalis (A.C. Sm.) van der Werff & H.G. Richt. up to N Brazil, none endemic.

1.4 TRIBE CINNAMOMEAE (40-45/c. 2,000) - a bulk of Lauraceae family, 8 linegaes, Laurus Clade (10/c. 500, Europe, tropical and subtropical areas of both hemispheres), Cinnamonum s.s. + Kuloa (2/?; Old World), Cinnamonum sect. Camphora + Sassafras (2/?; Old World, North America) does not occur in South America. Unpalced genera are Cinnadenia (2; SE Asia), Parasassafras (1; Asia), Mocinnodaphne (1; North America, Central America), Povedadaphne (1; Costa Rica), Iteadaphne (20-50; Asia) .

∎ MEZILAURUS CLADE ‣ all genera in South America.

6. Anaueria Kosterm. The seeds are said to be edible after roasting; a cupule is lacking in fruit; flowers have green tepals and dark red stamens, very unusual in Lauraceae. Only one sp., A. brasiliensis Kosterm., known from Amazonian rainforest near the tripoint of Brazil (Amazonas state), Colombia and Peru.

7. Chlorocardium Rohwer, H.G. Richter & van der Werff. Leaves opposite. Three spp., C. rodiei (R.H. Schomb.) Rohwer, H.G. Richt. & van der Werff from Guyana and Suriname, and two from Amazonian rainforest in Ecuador (one endemic) and adjacent Colombia.

8. Clinostemon (A. Samp.) Kuhlm. & A. Samp. Mezilaurus-like, except for stamens with downward recurved anthers and one pair of glands at the base of the filaments, while in Mezilaurus the anthers are erect and glands are absent. Only one sp., C. mahuba (A. Samp.) Kuhlm. & A. Samp., disjunct from Pará and S Amapá state, and NE Peru and W Amazonas state, in flooded and ‘terra-firme’ regions.

9. Mezilaurus Kuntze ex Taubert. Trees up to 40 m tall; leaves alternate, clustered at tips of branches, flowers trimerous; the fruit is seated on a small, platelike cupule; the flowers are very similar to those of Licaria, but the two genera differ in cupule shape, leaf position, inflorescence type and wood anatomy. 24 spp. occurring in the Amazon rainforest including N Bolivia, Peru, SE Colombia, Venezuela, and the three Guianas, slightly disjucnt in SE Brazil (18 in over country, 14 endemics); six Brazilian species (5 in Amazonas and 1 in Rio de Janeiro state) are rare plants in Brazil, by Plantas Raras do Brasil’s book.

10. Sextonia van der Werff. Tres with leaves congested in axis of the branches. Two spp., S. rubra (Mez) van der Werff in Guianas, Brazil, Venezuela and Peru, and S. pubescens van der Werff endemic to Peru.

11. Williamodendron Kubitzki & H.G. Richter. The cupule is small and platelike, resembling that of Mezilaurus; leaves are clustered at the tips of branches. 5 spp., infrequently collected, one known from Costa Rica, another from N Colombia, and three in forested areas in southern Brazil, one of then from Espírito Santo state is a rare plants in Brazil, by Plantas Raras do Brasil’s book.

∎ PERSEA CLADE ‣ outsiders Alseodaphnopsis (1; China), Alseodaphne (40; Asia), Apollonias (1; Azores, Canary Islands, and Madeira), Dehaasia (50; Asia and islands of Borneo, New Guinea), Nothaphoebe (40; Asia), Phoebe (100; Asia), Machilus (100; Asia)

12. Persea Mill. Shrubs or small trees. 107 spp. in Neotropics, 73 in South America but better represented in Asia and with one species in Canary Islands; two subgenera:

§ subg. Eriodaphne ‣ remaining of genus, includes all native species known to occur in Brazil (23, 19 endemics, 3 of then, in Amazonas, Minas Gerais and São Paulo states are rare plants in Brazil, by Plantas Raras do Brasil’s book).

§ subg. Persea ‣ two spp. from Mexico to Panamá, P. americana L. and P. albida Kosterm.

∎ LINEAGE AIOUEA ‣ a single genus.

13. Aiouea Aubl. (exc. Damburneya p.p., Cinnamomum) Leaves alternate, penninerved, rarely triplinerved. 78 spp., ranging from S Mexico to S Brazil and Paraguay, 44 in South America; most species have the fruit seated on a shallow cupule with a thickened pedicel; 31 in Brazil, 20 endemics; A. macedoana Vattimo-Gil from Tocantins state is a rare plants in Brazil, by Plantas Raras do Brasil’s book.

∎ LINEAGE OCOTEA S.S. ‣ included Endlicheria, Pleurothyrium, Rhodostemonodaphne, Nectandra, Damburneya, and most of the species currently included in Ocotea., inc. the type species of this genus; al genus occur in South America.

14. Andea van der Werff. (off Ocotea) Ocotea-like shrubs and trees; 25 spp., Colombia (14, 7 endemics), Ecuador (9, 3 endemics), Peru (6, 3 endemics), Venezuela (3, two endemics) and Costa Rica (1, shared with Colombia).

15. Damburneya Raf. (off Nectandra, inc. Aiouea p.p.) Trees or shrubs; leaves evergreen, alternate, penninerved, with ± evenly reticulate to mixed (not scalariform) tertiary venation; flowers trimerous, bisexual. 24 spp., mainly Central America, with a few species reaching North America (Sinaloa and Tamaulipas; Florida), the Antilles and Bahamas, as well as northern and NW South America, and D. purpurea (Ruiz & Pav.) Trofimov reaching Bolivia and EspÍrito Santo in Brazil; including recently four species previously reported in Aiouea.

16. Endlicheria Nees. 61 spp. reported from Costa Rica, the Lesser Antilles south to Paraguay, and SE Brazil (42, 7 endemics), only one does not occur in South America; E. coriacea Chanderb. from Manaus region is a rare plants in Brazil, by Plantas Raras do Brasil’s book.

17. Nectandra Rolander ex Rottb. (exc. Damburneya p.p.) Trees up to 45 m tall; fruits are seated in a cupshaped (sometimes small) cupule; tepals are spreading at anthesis. 103 spp., from from southern Florida and Mexico to Argentina, c. 90 spp. in South America; 44 spp. in Brazil, 16 endemics, 3 of then in Pará, Minas Gerais and Rio de Janeiro states are rare plants in Brazil, by Plantas Raras do Brasil’s book.

18. Ocotea Aubl. (exc. Mespilodaphne, Andea) Trees up to 45 m tall. 472 spp. in the Neotropics from Mexico to Argentina, plus ca. 50 in Madagascar, 7 in mainland Africa and one in Canary Is., mainy in montane forests, but also on dry environments and Atlantic sandy coastal shrublands (restingas); 357 spp. occur in South America, 174 spp. occurs in Brazil (109 endemics); 13 spp. of this genus in several Brazilian states are rare plants in Brazil, by Plantas Raras do Brasil’s book. Some species in South America (inc. Brazil) are myrmecophytes.

19. Pleurothyrium Nees. Leaves alternate, inflorescences thyrso-paniculate; the fruit is seated in a rather deep, cupshaped cupule, which is often warty or lenticellate; several species have a prononced marginal vein, an unusual feature in Lauraceae, or clustered leaves. 48 spp., known from Guatemala south to Peru, Bolivia, and Brazil (37 in South America, 13 in Brazil, 4 endemics); P. undulatum (Meisn.) Rohwer from Amazonas state is a rare plants in Brazil, by Plantas Raras do Brasil’s book. Some species in South America (inc. Brazil) are myrmecophytes.

20. Rhodostemonodaphne Rohwer & Kubitzki. Inflorescense thyrso-paniculate; the cupule is deeply cupshaped and rather large. 40 spp., two of which are rather common, while several species are in need of description, all in South America, one reaching into Costa Rica; 17 spp. in Brazil, 8 endemics, three of then all from Amazonas state, are rare plants in Brazil, by Plantas Raras do Brasil’s book.

∎ ANIBA S.S. CLADE ‣ included Umbellularia, Licaria, Phyllostemonodaphne, Dicypellium, Urbanodendron, Aniba, Mespilodaphne, and some species of Ocotea. All of the species of the Ocotea complex with a distribution outside of the Neotropics were included in this lineage. Species of Paraia, Kubitzkia, and Gamanthera, when sampled, are also expected to belong to this clade; outsiders Gamanthera (1; Costa Rica) and Umbellularia (1; North America).

21. Aniba Aubl. Shrubs to trees, fruit seated in a rather deep cupule, which is often lenticellate or warty; several species have clustered leaves and pleasantly aromatic yellow wood (rosewood oil is distilled from A. rosaeodora Ducke from Brazil and Colombia). 53 spp. from Costa Rica to S Brazil (30, 9 endemics) and Bolivia; most species occur in the lowlands, but it has also been collected in the Andes up to 1,800 m; only Caribbean A. ramageana Mez. does not occur in South America; two spp. from Amazonas and one of Rio de Janeiro state are rare plants in Brazil, by Plantas Raras do Brasil’s book.

22. Dicypellium Nees & Mart. Leaves alternate, inflorescences bothroyd, few flowered. Two spp., rarely collected species, D. manausense W.A. Rodrigues restricted to Amazonas state in N Brazil (a rare plants in Brazil, by Plantas Raras do Brasil’s book) and D. caryophyllaceum (Mart.) Nees also in Guiana region, the latter heavily exploited because of its aromatic bark.

23. Kubitzkia van der Werff. (inc. Systemonodaphne) Trees; the cupule is cupshaped, with a double margin and persistent, reflexed tepals. Two spp., K. mezii (Kosterm.) van der Werff known from French Guiana, Guyana, Suriname and Amazon rainforest in Brazil, and K. macrantha (Kosterm.) van der Werff from Venezuela.

24. Licaria Aubl. The cupule is cupshaped and has a double margin; however, in some species the double margin is scarcely noticeable; few species have opposite leaves. 65 spp. from S Florida and Mexico to S Brazil and Bolivia, 43 in South America, 21 in Brazil, 3 endemics.

25. Mespilodaphne Nees & Mart. ex Nees. (off Ocotea) Trees os shrubs; leaves evergreen, alternate, lanceolate to elliptic, glabrous or very sparsely pubescent; inflorescences axillar, few flowered; tepals six. 9 spp., three restricted from Mexico to Central America, one only in Jamaica, remaining five in northern South America up to Brazil (3, one endemic), inc. M. veaguensis (Meisn.) Rohwer up to Mexico.

26. Paraia Rohwer, H.G. Richter & van der Werff. Leaves alternate, inflorescenses apparently dibothroyd. Only one sp., P. bracteata Rower, Richter and van der Werff, known from Amazonian rainforest of Brazil in Pará, Amapá and Amazonas states.

27. Phyllostemonodaphne Kosterm. Leaves alternate; the fruit is seated in a double-rimmed cupule. Only one sp., P. geminifolia (Mez) Kosterm, from the Atlantic rainforests in southern Brazil.

28. Urbanodendron Mez. Leaves alternate; fruit with a bowl shaped, double-rimmed capsule; the cupules are known in two species: double-rimmed, with more or less persistente tepals. Three spp. restricted to S Brazil, with U. macrophyllum Rohwer from Rio de Janeiro state is a rare plants in Brazil, by Plantas Raras do Brasil’s book.

8. CHLORANTHALES

A SINGLE FAMILY, PRESENT IN SOUTH AMERICA.

CHLORANTHACEAE

Genera/species 4/75 Distribution Madagascar (Ascarina), tropical Himalaya, South, E and SE Asia, Borneo, New Guinea, Melanesia to Marquesas Islands, Vanuatu, Fiji, New Caledonia, North Island of New Zealand, tropical America. Habit bisexual (Chloranthus, Sarcandra), monoecious or dioecious (Ascarina, Hedyosmum), evergreen trees or shrubs, or perennial herbs (sometimes woody at base, rarely annual). Aromatic.

An interesting family from an evolutionary point of view, lending its name to the Chloranthoid Hypothesis, which suggests that the first flowering plants had the similar small, basic, unisexual, wind-pollinated flowers.

SYSTEMATICS outsiders Ascarina (12; Madagascar, Borneo and E Malesia to Melanesia, North Island in New Zealand, Marquesas and other SW Pacific islands), Chloranthus (18; S and E Asia), Sarcandra (2; India, Sri Lanka, China, Japan, Indochina, Malesia).

1. Hedyosmum Sw. Evergreen trees (2-30 m tall) and scandent shrubs, all parts strongly aromatic; leaves simple; staminate inflorescence in dioecious species axillary or terminal, 1 to several spikes, axis racemose or paniculate; in monoecious species solitary spikes on pistillate inflorescence axis or within a cymules (along with pistillate flowers); pistillate inflorescence axillary or terminal, cymules, spikes, racemes or panicles; bracts present; staminate flowers basic, 60-300 flowers per spike, without perianth; pistillate flowers simple, perianth lobes 0.5-3mm, free at the base or partially to completely united forming a small tube, adnate to the ovary; fruit a drupe, fleshy, aromatic and brightly coloured; seed 1 per fruit, small, brown or black smooth or minutely papillate, endosperm well developed, oily. 43 spp., Mexico to Cono Sur and West Indies, with H. orientales Merrill & Chun in China and SE Asia; growing throughout the Neotropics, predominantly in wet, cool, montane forest at altitudes anywhere between 500 m and 3,000 m; 33 spp. in South America, 9/10 of the species are found in Andean South America from Venezuela to Bolivia, primarily in disturbed sites; Brazil has three spp., two restricteds of mountains in north: H. neblinae Todzia from Brazil and Venezuela; H. racemosum (Ruiz & Pav.) G. Don in northern Amazon rainforest; and H. brasiliense Miq., widespread in over country, also in Venezuela and Argentina, which has medicinal property; H. mexicanum C. Cordem. provides edible, sweet tasting fruit.

9. ALISMATALES

FAMILIES ABSENTS IN SOUTH AMERICA: APONOGETONACEAE (1/60), BUTOMACEAE (1/2), MAUNDIACEAE (1/1), POSIDONIACEAE (1/9) AND SCHEUCHZERIACEAE (1/1).

LINEAGE 1 of 3: ARACEAE

ARACEAE

§ FULLY CHLOROTROPHIC

Genera/spp. 142/4,000 Distribution mainly tropical but also subtropical regions, and approx. ten genera in temperate regions in the Northern Hemisphere. Habit bisexual, monoecious (often with female flowers in lower part of inflorescence and male flowers in middle and upper parts), andromonoecious, gynomonoecious, polygamomonoecious, dioecious, or gynodioecious, usually perennial (rarely annual) herbs (some are giant herbs, whereas others are extremely small [e.g. Wolffia]), often epiphytic or climbing with stem or roots (rarely suffrutices). In the aquatic Lemnoideae, leaves are absent and the stem is reduced to a usually flat thalloid floating structure with or without roots on the lower side. Some genera have a starchy stem tuber (e.g. in Lasioideae and Aroideae). Many representatives are aquatic (e.g. Pistia) or hygrophilous. Turions are formed in, e.g., Spirodela, Wolffia and some spp. of Lemna.

Gigantic to minute herbs, terrestrial, epiphytic, hemiepiphytic or epilithic, rarely aquatic (free-floating or rooted). Roots often aerial, in the Lemnoideae hair-like or absent. Stem aerial and erect (e.g. Dieffenbachia Schott and Rhodospatha Poepp.), creeping, subterranean and rhizomatous or tuberous (e.g. Caladium Vent. and Homalomena Schott), or appressed-climbing, frequently scandent, rarely erect, hardened, and armed (Montrichardia), or not differentiated into stem or leaf (Lemnoideae), in this case the plant is reduced to a minute, fleshy or flattened plant body bearing hair-like roots on the undersurface or roots absent. Cataphylls often variously ribbed and persistent, may remain intact or weather into fibres. Leaves alternate, sometimes distichous, simple, basal or cauline, rarely solitary (e.g. Dracontium L.); petioles often elongate, sheathed at least basally, the apex often geniculate, blades often variegated, often oblong, cordate, sagittate to hastate, sometimes perforated, the margins entire, or pinnately to palmately lobed, main veins often radiate from petiole attachment, rarely parallel, the primary lateral veins (in some families referred to as secondary veins) pinnate, sometimes joining into collective veins along margins, the lesser veins parallel or reticulate. Inflorescences terminal or axillary, solitary or clustered in axils, an unbranched spadix (spike) subtended by a single spathe (bract), in Lemnoideae the inflorescence is within a minute dorsal cavity of the plant body (Wollfia Horkel ex Schleid., Wolffiella Hegelm.), or in paired lateral budding pouches (Spirodela Schleid., Landoltia Les & D.J. Crawford, Lemna L.); spathe herbaceous, free or adnate to spadix, spreading, reflexed or convolute, sometimes constricted below middle and differentiated into tube below and blade above, often persistent, sometimes (especially when convolute) reclosing over spadix after anthesis, the blade sometimes deciduous, green, or blade and tube of spathe differently coloured; spadix often cylindric, flowers dense, the lower part often pistillate, this part often protected by accrescent spathe tube until maturity of fruits, some parts with sterile flowers or without flowers, the upper part often staminate, this part (e.g. Syngonium Schott and Xanthosoma Schott) often caducous, the apex frequently tapered, less often clavate or ellipsoidal. Flowers sessile, small, bisexual or unisexual (the plants usually monoecious), 2-3-merous, protogynous, lacking floral bracts; perianth (=perigone) lacking in unisexual flowers; tepals 4-6, usually free, rarely united; androecium of (1-)3-6(-9) stamens, the stamens free or united into synandria, the thecae terminal or lateral, extrorse, dehiscent by longitudinal slits or apical pores; staminodes sometimes present, free or united into synandria; gynoecium syncarpous, the ovary usually superior, sessile or immersed in the spadix, often entire, rarely lobed, usually 1-3 locular, rarely more, the style usually inconspicuous, the stigma wet at anthesis, sometimes distinctly lobed; placentation axile, parietal or basal, the ovules 1-many per locule, often with funicle, less often sessile. Fruits berry-like, sometimes juicy, free or rarely fused into syncarps (Syngonium), often colourful, rarely utricles (Lemnoideae); seeds 1-many; endosperm copious to absent.

Amorphophalus titanus (Becc.) Becc. ex Arcang. has one the largest inflorescences of Earth; Wollfia, Spirodela, Lemna and Pistia is pantropical; three are Holarctic (Arisaema, Calla and Lysichiton). Wollfiella are New World and Africa. Some genera have not leaf vernation more typical of monocotyledons. Araceae contains crystals of calcium oxalate, a irritant compound. About 35 genera and 450 spp. in Brazil. More then 4/5 are Anthurium or Philodendron; true epiphytes in Neotropics occur only in Anthurium, Philodendron and Stenospermation.

Use Ornamental plants, aquarium plants (Pistia etc.), starch sources and vegetables (Colocasia esculenta (L.) Schott, Xanthosoma sagittifolium (L.) Schott, Amorphophallus paeoniifolius), fruits (Monstera deliciosa Liebm.), gels in food-stuffs (A. konjac), medicinal plants, arrow poisons, fibres for basketry, animal forage (Lemnoideae). Several Aroids are cultivated for their edible tubers and leaves; some species are also used in folk medicines and as a source of fibre; aroids are widely used as ornamental plants across the world, both as house plants ands garden plants especially in the tropics.

SYSTEMATIC Gymnostachydoideae (1/1, E Queensland, E New South Wales), Orontioideae (3/8, temperate E Asia, W and E North America), and Stylochaeton clade (Stylochaeton + Zamioculcadoideae, 3/26, tropical Africa) do not occur in South America.

1. SUBFAMILY LEMNOIDEAE (5/38) ‣ all genera occur in South America.

Root hairs absent; Herbs very reduced a leaflike frond. Lemnoideae are the smallest and most reducing flowering plants; the smallness of the plants requires good binoculars magnification and some technical preparations for identification. Lemnoideae are easily distributed by birds over shorts distances. In many places, they live only so long as conditions are favorable; afterwards they disappear.

A Key To The Genera Of Lemnoideae

1. Plant body with 1 - several roots.

2. Root one ------------ Lemna
2. Roots 2–12 - 3.

3. Roots 7 - 12 (or more); plant 10 mm long ------------ Spirodela
3. Roots 2 - 3 (up to 5); plant 3 - 6 mm long ------------ Landoltia

1. Plant body without roots - 4.

4. Plant body flattened; 3 - 10 mm long ------------ Wolffiella
4. Plant body globose-ovoid; 0.6 - 1.2 mm long ------------ Wolffia

Lemnaceae are the smallest and most reduced flowering plants. Therefore, closely related species have but few distinguishing characteristics that are easy to recognize. In addition, most of these features are strongly modificable and overlap considerably. Occasionally Lemnaceae have small, 1--5-celled papillae on the upper surface of the frond. Some species have turions (compact fronds reduced in size and structure, filled with starch grains, forming under unfavorable conditions). Easily recognized by their darker color, tTurions remain for several days with the mother frond before they sink to the bottom.

1. Landoltia Les & D.J.Crawford. Only one sp., L. punctata (G.Mey.) Les & D.J.Crawford, cosmopolitan (most probably having originated in SE Asia and Australia), occurring in regions with mild winters and not very hot summers, from sea level to 2,400 m; it occurs in bodies of fresh or low salinity water, protected from strong winds and currents, having a preference for waters rich in nutrients, especially nitrogen and phosphorus; in New World grows only in Caribbean, Guianas, Colombia, Brazil and Ecuador.

2. Lemna L. Fronds veined; one root per frond. 13 spp. worldwide; cosmopolitan, 7 spp. in New World, 5 in South America, four very widely (all in Brazil) and one only in North America to Mexico and Ecuador.

3. Spirodela Schleid. Frods veined, 2-21 roots per frond. Two spp., both over widely distributed, but the former restricted of tropical America.

4. Wolffia Horkel ex Schleid. Roots absent; fronds thick, ovoid, globose; air spaces lacking; flowers inside one cavity near median line of upper frond; the smallest angiosperm worldwide. 11 spp., cosmopolitan, six restricted for larges landmass: Indian region (2, India, Pakistan, Sri Lanka, Bangladesh), Australia and New Zealand (1), Africa (1), W. elongata Landolt in NW South America, Cuba and Curaçao, W. borealis (Engelmann) Landolt only in North America, W. brasiliensis Wedd. and W. columbiana H. Karst. Widely distributed in New World, and two widely in Old World.

The smallest vascular plant species is W. angusta Landolt, native to the Malay Peninsula, New Guinea and Australia; the seven smallest species are all native to the Old World, and the remaining 4 are all New World, and the smallest New World species is W. borealis (Englem.) Landolt., followed by W. brasiliensis Wedd. - threse the smallest angiosperm species in South America and Brazil.

5. Wolffiella Helgelm. Roots absent; fronds thin, apalainated; orbicular, oblong to linear and falcate, with one layer of air space; only one cavity (two in W. welwitschii). 10 spp., 4 only in Africa (some with restricted ranges), one from Africa disjunct in Central America, W. caudata Landolt endemic to Bolivia, one only in North America, W. neotropica Landolt in Brazil and Suriname - a very rare species, known only c. 10 collections, mainly on northern Brazil, however some records in Suriname and Rio de Janeiro state; and two are very widely in tropical and temperate ranges in New World.

2. SUBFAMILY POTHOIDEAE (4/1,100) ‣ two tribes, Potheae (4/60; Madagascar, tropical Asia, NE Australia, Melanesia) does not occur in South America; Anthurium clade is monotypic.

6. Anthurium Schott. Stem creeping to erect, sometimes root climber; blade entire linear to cordate, and any others formats; usually true epiphyte or climbing hemiepiphyte, less often lithophyte (many E Brazil species grws in exposed rockies) or terrestrial; stem aerial, not tuberous or rhizomatous. 1,267 spp., Mexico to Argentina, 959 in South America, 153 in Brazil (121 endemics, 17 in several states are rare plants in Brazil, by Plantas Raras do Brasil’s book), 411 in Colombia, from humid to dry tropical lowland to lower montane forests, normaly epiphytic or climbing hemi-epiphytic, also terrestrial, lithophytic or rarely rheophytic. 19 sections.

CLADE 1 - POLYPHYLLIUM

§ Polyphyllium - two spp., A. flexile Schott and and A. clidemioides Standley, Mexico to Colombia; most basal member of genus.

CLADE 2 - U2D CLADE

§ Urospadix (inc. Chamaerepium) - 60 spp., endemic to E Brazil; within this clade, A. lucidum Kunth, endemic to E Brazil, with a chromosome number of ca. 124 is the highest known chromosome number in Anthurium.

§ subclade 2 - Pachyneurium p.p., possibly absent in Brazil.

§ Dactylophyllium - 25 spp., tropical America, 10 in Colombia, 7 in Brazil.

CLADE 3 - REMAINING TAXA

§ Andiphilum - 25 spp., mainly Central America.

§ Calomystrium - 184 spp.

§ Leptanthurium - a single sp., A. gracile (Rudge) Schott, over tropical America.

§ Tetraspermuim - 35 spp., scandent hersb, over tropical America.

§ Belolonchium - 220 spp.

§ Cardiolonchium - 175 spp.

§ Decurrentia - 45 spp., from Central America to N Brazil.

§ Digitinervium - 41 spp., Costa Rica to tropical Andes.

§ Gymnopodium - only one sp., A. gymnopus Griseb. from Cuba.

§ Multinervia - 16 spp.

§ Pachyneurium - 156 spp. (inc. several species formerly placed in Urospadix section), birds’s nestings, over tropical America. A. bovinii Camelo & Nadruz, belonging this section, from SW Amazonas state in northern Brazil, has the smallest spadix in genus, with only 0.5cm length.

§ Polyneurium - tropical America.

§ Porphyrochitonium - 375 spp., a large group from Costa Rica to Ecuador, mainly in Colombia; only one sp. in Brazil, A. bakeri Hook.f. (SEE).

§ Semaeophyllium - 23 spp., Nicaragua to Peru.

§ Xialophyllium - 108 spp., tropical America.

3. SUBFAMILY MONSTEROIDEAE (12/365–370) ‣ three tribes, all in South America.

3.1 MONSTEROIDEAE ▸ TRIBE SPATHIPHYLLEAE (2/41) - outsider Holochlamys (1; New Guinea, New Britain).

7. Spathiphyllum Schott. Herbs, penducle subequal to or longer than leaf; inflorescences adnate in leaf blade (true epiphylly). 49 spp., western Pacific, Malesia. 46 spp. in the Neotropics, few known, mostly in forests; 36 in South America, 10 in Brazil, two endemics.

3.2 MONSTEROIDEAE ▸ TRIBE HETEROPSIDEAE (4/100) - all genera in South America.

8. Alloschemone Schott. Leaves distant, climbers or hemiepiphytes in rainforests; leaf blade pinnately lobed, subcordate, lobes acute, falcate. Two spp. from N & C Brazil, one of then up to NE Bolivia.

9. Heteropsis Kunth. Bushy root climbers or hemiepiphyts; main shoot monopodial. 19 spp., Central and South America (all species) disjunct between the Amazon rainforest and the Brazilian Atlantic Forest; 15 spp. in Brazil, 5 endemics.

10. Rhodospatha Poepp. Climbers or hemiepiphytes in rainforests; leaf blade entire, venation parallel-pinnate, spadix mainly pinkish to white. 30 spp., Mexico and Central America to Bolivia and E Brazil (5), 28 in South America, highly disjunct in Amazon and Atlantic Forest, absent in central dry savanas. R. arborescens Temponi & Croat, unique endemic to Brazil and endemic of Atlantic Forest, is the unique arborescent species in its genus; the arborescent habit is scarce, occurring only in Montrichardia and Philodendron.

11. Stenospermation Schott. Herbs sometimes true epiphytic, sometimes very large tank-like; shots usually shorts; leaf blade entire, epiphytic or on the ground. 50 spp. from Guatemala to SE Brazil, 48 in South America, centered to Amazon rainforest, only 4 in Brazil, none endemics.

3.3 MONSTEROIDEAE ▸ TRIBE RHAPHIDOPHOREAE (6/220) - outsiders Rhaphidophora (c. 100; tropical Africa, Madagascar, tropical Asia to islands in the Pacific), Anadendrum (12; S China, SE Asia, Malesia to Sulawesi and Philippines), Scindapsus (c 35; tropical Asia from India to Hainan and Solomon Islands), Epipremnum (15–17; Himalayas, S China, SE Asia, Malesia to tropical Australia and islands in W Pacific), Amydrium (5; S China, SE Asia, Malesia to New Guinea).

12. Monstera Adans. Climber hemiepiphytes or scadent, main spp. with perforations in leaves. 47 spp. from S Mexico to S Brazil, mainly in Central America, but 25 in South America. Brazil has 8 spp., most of them occurring only in Amazon rainforest. M. praetermissa E. G. Gonc. & Temponi is the only endemic to Brazil, in vast area from N to S country from Ceará up to S Brazil.

4. SUBFAMILY LASIOIDEAE (10/c 55) ‣ outsiders Pycnospatha (2; SE Asia), Anaphyllum (2; Laccadive Islands), Cyrtosperma (12; New Guinea to islands in the Pacific), Lasiomorpha (1; tropical W and C Africa), Podolasia (1; W Malesia), Lasia (2; India, Tibet, China (inc. Taiwan), SE Asia, Malesia to New Guinea).

13. Anaphyllopsis A.Hay. Terrestrial or rooted aquatic herbs. Three spp., A. americana (Engl.) A. Hay in tropical South America, mainly N Amazonas river in Pará state and S Suriname, A. pinnata A. Hay only in Venezuela (unique truly pinnate compound leaf in New World in Araceae), and A. cururuana A. Hay endemic to Brazil.

14. Dracontioides Engl. Terrestrial or rooted aquatic herbs; rhizome hypogeal, erect, leaf blade strongly sagittate, black spadix. Two spp., endemics to E Brazil, from Pernambuco to Rio de Janeiro state, common in humid forests or in shaded places along of Atlantic Forest.

15. Dracontium L. Sometime gigantic, tuber depressed-subglobose; leaf usually 1, blade dracontioid; inflorescence 1; tallest peciole of leaves. 26 spp. from Mexico to Paraguay and Bolivia, in forest and savanna, ground-dwelling, seasonally dormant or evergreen; 24 in South America, 13 in Brazil, 7 endemics, only D. nivosum (Lem.) G.H. Zhu in dry seasonal scrubland of NE Brazil (caatinga), at wet shady mountains areas of Ceará state, also in Amazon rainforest in Maranhão e Pará states.

16. Urospatha Schoot. Rhizome hypogeal, horizontal to vertical, leaves several. 10 spp., Guatemala to SE Brazil (7, two endemics), 9 in tropical South America, open aquatic habitat, swamps, along rivers, in brackish waters.

5. SUBFAMILY AROIDEAE (79/1,900–1,935) ‣ 12 small lineages absent in South America: Callopsideae (1/1, Tanzania), Anubiadeae (1/8, tropical W and C Africa), Aglaonemateae (2/22, tropical Asia), Nephthytideae (3/14, tropical Africa, tropical Asia), Culcasieae (2/37, tropical W and C Africa), Zantedeschia clade sensu stricto (1/8, E tropical and S Africa), Cryptocoryneae (2/65, India and Sri Lanka to S China and SE Asia, Malesia to New Guinea), Schismatoglottideae (12/C. 180, tropical Asia, with their highest diversity on Borneo), Thomsonieae (1/c 200, tropical regions in the Old World), Arisareae (2/4, Macaronesia except Cape Verde Islands, Mediterranean to the Caucasus), Typhonodoreae (2/3, SE Canada, E and SE U.S.A., Cuba, coastal regions in E Africa, Madagascar, Mascarene Islands) and Arophyteae (3/11, Madagascar).

5.1 AROIDEAE ▸ TRIBE MONTRICHARDIEAE (1/2) - a single genus.

17. Montrichardia L. Helophytic herbs, arborescent, up to 6 m high. Two spp. from Central and over tropical South America and Caribbean; both species can reach up to 6 meters in height and which form dense populations in the beds of water courses.

5.2 AROIDEAE ▸ TRIBE PHILODENDREAE (4/c. 620) - outsider Homalomena (100–105; tropical Asia).

18. Adelonema Schott. Rarely robust; inflorescences 1-6; leaf blade lanceolate to sagittate, rarely peltate, some maculate. 16 spp. from Costa Rica to Bolivia and Brazil (4, none endemics) eastwards to French Guiana; 12 in South America; two sections, both in Brazil:

§ sect. Curmeria ‣ 6 spp., three in Brazil.

§ sect. Adelonema ‣ 10 spp., only one in Brazil.

19. Philodendron Schott. (exc. Thaumatophyllum) Climbing hemiepiphytes, true epiphytes or terrestrial herbs (some arborescent) with petiolar sheath much reduced; if petiolar sheath well-developed then plants climbing; leaf blades highly variable – ranging from linear-lanceolate to complexly bipinnatifid. 518 spp., throughout Neotropics, 457 in South America, 154 in Brazil, 74 endemics (10 of then rare spp. in several Brazilian states, by Plantas Raras do Brasil’s book), with numerous very narrow endemisms from E Brazil, grouped into three subgenera according to its floral and vegetative morphology and anatomy, namely:

§ subg. Philodendron ‣ 435 spp., the bulk of genus. P. myrmecophilum Engl. endemic to W Brazil is the only myrmecophte Araceae worldwide.

§ subg. Pteromischum ‣ 75 spp., two centers of diversity, one in Central America and another in West Amazonia, but is poorly represented in extra-Amazonian South America; only 8 spp. occur in Atlantic Forest in Brazil.

20. Thaumatophyllum Schott. (off Philodendron) Arborescent habit, very much thickened spathe, well developed sterile intermediate zone in the inflorescence equal or longer than the staminate zone, the gynoecium always having stylar lobes and an axial vascular system independent of the funicle supply. 21 spp., 19 in Brazil (13 endemics), several up to Cono Sur or Amazon rainforest in Colombia and Guianas; Paraguay and Venezuela one endemic each; terrestrial to hemi-epiphytic, but can be rupicolous, aquatic or subaquatic in freshwater swamps at lowland sites; all the extant species have a notable preference for open habitats and the ability to tolerate a certain degree of drought.

5.3 AROIDEAE ▸ TRIBE SPATHICARPEAE (13/80) - all genera in South America.

21. Asterostigma Fisch. & C.A.Mey. (exc. Incarum) Large terrestrial herbs, seasonally dormant, leaf usually 1; leaf blade pinnatisect or rarely entire; inflorescence 1-3, appearing before or with leaves. 11 spp. from lowland to mountains areas in S & SE Brazil in Atlantic Forest, one reaching to northeast coast and another into central dry savannas.

22. Bognera Mayo & Nicolson. Stem creeping, epigeal, rhizomatous; foliage leaf 1 per article, erect. Only one sp., B. recondita (Madison) Mayo & Nicolson known only small area in E banks of Rio Javari in forests floor, W Amazonas state, Brazil.

23. Croatiella Schott. Geophytic herb; stem hypogeous, tuberous, with 3-6 connected tubers forming an elongated structure; leaf solitary, erect; petiole with a long sheath, blade ovate-cordate; inflorescence solitary(?); spathe erect, not at all constricted at middle; spadix adnate to the spathe at the basal portion of the female zone. Only one sp., C. integrifolia (Madison) E.G. Gonç., only known from the Zamora River valley, E Ecuador, on the eastern slope of the Andes, between 2,800-3,150 m in cloud forest and seems to be common locally along the edges of roads.

24. Dieffenbachia Schott. Sometimes robust; stems erect or decumbent; leaves several per article. 51 spp., Mexico to W and C South America, Caribbean, mainly in Central America rainforests, evergreen, terrestrial or helophytic, in humid forets or damp sites; 35 spp. in South America, 12 in Brazil, two endemics.

25. Gearum N.E.Br. Herbs, seasonal, hibernat in dry season, flowering with fall rain; rhizome vertical, leaves 1-2, appearing soon after inflorescence, pedates, polyploid. Only one sp., G. brasiliense N.E.Br., restricted to savannas of C Brazil (cerrado) in S Tocantins and N Goiás state in center Brazil.

26. Gorgonidium Schott. One leaf, blade pinnatifid, pinnatisect or bipinnatifid; inflorescence 1-2, appearing before leaf. 8 spp., Peru to N Argentina.

27. Incarum E. G. Gonc. (off Asterostigma) Seasonally dormant or evergreen; leaf 1, erect; inflorescence 1-2, appearing soon after leaves. Only one sp., I. pavonii (Schott) E. G. Gonç., from Andes from Ecuador to Bolivia, at 1,200-3,300 m on eastern slopes or in inter Andean valleys.

28. Lorenzia E. G. Gonc. Chamerophytic herbs; stem rhizomatous, usually hypogaeous; foliage leaves sparsely scattered along the stem, petiole erect, pilose; blade deflexed, sagittate; inflorescence 1–2, erect at anthesis; spathe erect, not at all constricted; flowers unisexual, naked, all pilose; infructescence unknown. Only one sp., L. umbrosa E. G. Gonç., known only from a single remote locality in Serra do Navio, Amapá state in N Brazil; although it is possible that it also occurs in adjacent French Guiana; it is moderately common locally, always occurring in deep shade under large trees.

29. Mangonia Schott. Small, leaves several. Inflorescence 1, appearing before leaves; spathe tube to subcilyndric. Two spp., both in Rio grande do Sul state in S Brazil to Uruguay, in seasonally wet places.

30. Spathantheum Schott. Leaves 1; petiole sheath very short; inflorescence 1-2, inflorescences adnate in leaf blade (true epiphylly). Only one sp., S. orbignyanum Schott, from Peru to N Argentina.

31. Spathicarpa Hook. Small, tuber horizontal; leaves several; Inflorescence 1, adnate in leaf blade (true epiphylly). 4 spp., two endemics to E & NE Brazil; and two remaining reaching to Paraguay and Argentina.

32. Synandrospadix Engl. Leaves several; petiole sheath long; inflorescence 1-2, appearing before or with leaves. Only one sp., S. vermitoxicus, Peru to NW Argentina and Paraguay, in dry thorn forests.

33. Taccarum Brongn. Often robust; leaf 1; inflorescence 1-2, leaves many formats, some dracontioid. 6 spp., Peru to N Argentina and C & S Brazil (all species, 3 endemics), but absent in mainly Amazon rainforest.

5.4 AROIDEAE ▸ PHILONOTION CLADE (1/3) - a single genus.

34. Philonotion Schott. Stems rhizomatous or epigeal, shortly erect; inflorescence 1-3. Three spp. from Colombia to French Guiana and Peru, and N Brazil (2, none endemics), in forest floor, terrestrial or rheophytic.

5.5 AROIDEAE ▸ CALADIEAE (11/c. 210) - outsider Hapaline (8; Burma to Borneo).

35. Caladium Vent. Geophytes, seasonally dormant or evergreen, stem a subglobose tuber; leaf blade usually peltate, cordate, trisect ot sagitate. 18 spp. from Costa Rica to N Argentina, mainly in forests of N South America; 9 spp. in Brazil, two endemics; C. humboldtii (Raf.) Schott, nearly white leaves, mainly in between Madeira and Tapajós Rivers.

36. Chlorospatha Engl. Stem caulescent, decumbent to erect; leaves 1-few; inflorescence 3-18, spethe tube-narrow. 68 spp., one in Costa Rica and Panamá, another in Panamá to N Colombia, 45 in Colombia (43 endemics) and 25 in Ecuador (22 endemics), with greatest in the Andes of frontier of Colombia and Ecuador; three shared species between Ecuador and Colombia.

37. Filarum Nicolson. Tuberous, geophytes, leaves few; inflorescences 1-3, appearing with leaves. Only one sp., F. manserichense Nicholson, Amazonian rainforest of Peru.

38. Idimanthus E.G.Gonç. Geophytic herb, glabrous; stem tuberous, then producing a foliage leaf or an inflorescences; foliage leaves one or three per plant, petiole spreading; leaf blade sagittate-cordate, venation reticulate, commonly silvery variegate; inflorescence solitary, erect at anthesis, peduncle usually shorter than the petiole. Only one sp., I. amorphophalloides E.G. Gonç., so far known only from the forests fragments in a single marble outcrop in the Italva municipality, Rio de Janeiro states, Brazil, very close to the Cardoso Moreira municipality.

39. Jasarum G.S.Bunting. Aquatic herbs, completely submerged (only the inflorescence appears over the water surface) in blackwater streams, stem hypogeal, rhizomatous, erect; leaves several. Only one sp., J. steyermarkii G.S.Bunting, in acidic blackwater from Bolivar state in Veneuzela and Mazaruni River Basin in Guyana, the unique genus of Araceae endemic to Guiana Shield and only submerged aquatic species in non-Lemnoid Araceae in New World (unlike tropical Asia, very rich in this habit for Araceae), ranging from 500-1,200 m high; the aquatic nature of this aroid is very remarkable because no true aquatic were previously known for the subfamily Colocasioideae.

40. Scaphispatha Brongn. ex Schott. geophyte, seasonally dormant, stems a subglobose tuber; leaves 1-2. Inflorescence 1, appearing before leaves. Two spp. from Brazil, one of then up to E Bolivia.

41. Syngonium Schott. Climbers or hemiepyphites with elongate intenodes. Leaves blade mainly sagitate. Inflorescence several, pendent in fruit. 43 spp. from Mexico to tropical America, 19 in South America, 5 in Brazil, none endemics.

42. Xanthosoma Schott. Sometimes gigantic to arborescent; tuberous; leaves-blade in many formats. 197 spp., Mexico and Cuba to S Brazil and N Argentina, on forest floor, or seasonally flooded sites; 180 spp. in South America, 26 in Brazil, 15 endemics.

The presence of both monads and tetrads in the same genus is very rare but not unique in the flowering plants (monads in three spp. from Venezuela to Peru, and tetrads elsewere genus); for instance, in the monocots Typha minima Funck ex Hoppe (Typhaceae) has the pollen in monads while all other spp. have it in tetrads, in the dicots Epilobium L. (Onagraceae) has both types, and also Podophyllum spp. such as P. hexandrum Royle (syn. P. emodi Hook. f. & Thompson) release the pollen in tetrads, whereas P. peltatum L. and P. pleianthum Hance have their pollen in monads.

43. Ulearum Engl. Rhizomatous, leaves few, leaf blade usually variegated; inflorescence 1. Only one sp., U. sagittatum Engler, W Acre state of Amazonian rainforest of Brazil and E Peru, on forest floor.

44. Zomicarpella N.E.Br. Geophyte, seasonally dormant; stems rhizomatous, leaves 1-few. Leaves blades variegated; inflorescence with leaf. Two spp., Z. amazonica Bogner in Amazon rainforest of N Brazil and Z. maculata N.E.Br. in Colombia.

45. Zomicarpa Schott. Geophytes, evergreen to seasonally dormant, stem cormous, usually cylindrical, contractile roots present; leaves 2 – 3 per plant; petioles sheathed up to the middle; leaf blade sagittate-cordate to pedatissect, higher order venation reticulate, often variegated in variable patterns; inflorescence 2 – 5 per article, appearing together with the leaves, rarely before. Two spp. from NE Brazil, in Atlantic sandy coastal shrublands (restingas) and to dry seasonal scrubland of NE Brazil (caatinga), also occurring within savannas of C Brazil (cerrado), sometimes as a deep shade understory herb in well-drained places.

5.6 AROIDEAE ▸ TRIBE PISTIEAE (17/435–440) ‣ outsiders are all tropics and subtropics of Old World to W Pacific except Arisaema (180–185; E Africa, S Arabian Peninsula, India, Himalayas, Tibet, E and tropical Asia to New Guinea, SE Canada, E and SE U.S.A., Mexico).

46. Pistia L. Small, free-floating, densely pilose aquatic plant, becoming fixed in shallow water, 7.9-13.5 cm tall, main stem very reduced, producing slender stolons; roots forming a pendent, feathery mass from the plant base with abundant secondary branching, white and brown; stolons 2.6-10.1 cm long, pilose, leafless, light green. Only one sp., P. stratioides L., floating spp. pantropical and subtropical, common in Brazil, known in over tropical and subtropical New World.

LINEAGE 2 of 3: TOFIELDIACEAE

TOFIELDIACEAE

§ FULLY CHLOROTROPHIC

Genera/spp. 4/26–28 Distribution temperate Eurasia, with the largest diversity in E Asia, NW South America (Venezuela, Brazil, Guyana, Colombia, Bolivia, Ecuador, and Peru), SE U.S.A. (Florida). Habit bisexual, perennial rhizomatous herbs.

SYSTEMATIC outsiders Pleea (1; Alabama, Florida, North and South Carolina), Tofieldia (7–10; temperate regions on the Northern Hemisphere), Triantha (4; Japan; North America).

1. Harperocallis. McDaniel. (inc. Isidrogalvia) Rhizomatous; foliage mostly basal; flowers solytary or in a compound inflorescence. 11 spp., one from Apalachicola River lowlands on the Coastal Plain of the Florida panhandle, four in Andes from Venezuela to Bolivia, and six in Guiana Shield, three known only from Mount Neblina, all in Venezuelan or Brazilian sides of masssif, with H. paniculata (L.M. Campb.) L.M. Campb. & Dorr, known only from Brazilian side, at 2,600–2,700 m, in Cauabari River margin; this spp. may occur also in the Venezuelan part of the massif; H. schomburgkiana (Oliv.) L.M.Campb. & Dorr also collected in Mount Roraima.

H. paniculata differs from others of genus in the basal portion of inflorescence included in the leaves, and by its largers inflorescence bracts; inflorescence compound, the branches subtended by chlorophyllous cataphylls; its unique in Tofieldiaceae in having a compound inflorescence.

LINEAGE 3 of 3: PALUDICOLA/THALASSIIDS

HYDROCHARITACEAE

§ SEA GRASSES (Heterozostera – Ruppia – Halodule - Syringodium – Halophila – Thalassia)

Genera/spp. 14/110–115 Distribution mainly in tropical and subtropical but also temperate regions: North America, Eurasia, Australia, coastal marine areas in the Caribbean, and the Indian Ocean to the Pacific. Habit usually monoecious, polygamomonoecious, or dioecious (sometimes bisexual), usually perennial (sometimes annual) herbs. Aquatic, often submersed. Some genera are growing in salt or brackish water. Roots usually unbranched (in Najas fibrous). Stems monomorphic, dimorphic or polymorphic. 14 genera, three marine. All submersed herbs in many forms of growth. Centres of diversity is Asia (for species) and Africa (for genera).

Aquatic herbs, sometimes emergent. A family of 18 gg. and c.115 spp. 10 genera comprising 21 spp. occur in the Neotropics. Najas is the largest genus in the Neotropics with 7 spp., sometimes armed with prickles on internodes, leaf margins usually serrate to minutely serrulate. Six genera and 15 spp. in Brazil. Two spp. Hydrilla verticilata (L.f.) Royle and Najas graminea Delile are naturalised. Hydrocharitaceae occupy streams, rivers, lakes, bays and oceans, and are of limited economic use, except as aquarium plants; two Neotropical genera are strictly marine (Halophila and Thalassia), the rest occupy freshwater habitats. Their invasive nature, however, is more important.

SYSTEMATIC 4 subfamilies, three in South America, Stratiotoideae (1/1, Europe, temperate Asia) absent.

1. SUBFAMILY HYDROCHARITOIDEAE (1/5) - a single genus.

1. Hydrocharis L. (inc. Limnobium) Freshwater; stems contracted, roots branched; leaves uniforme. 5 spp., three in Old World, and two in American fresh waters, H. laevigata (Humb. & Bonpl. ex Willd.) Byng & Christenh in the Neotropics, from Mexico to tropical South America, other from SE U.S.A.

2. SUBFAMILY ANACHARIDOIDEAE (7/51-52) ‣ outsiders Appertiella (1; Madagascar), Lagarosiphon (9; tropical and S Africa, Madagascar), Blyxa (12; tropical Africa, Madagascar, tropical and subtropical Asia to tropical Australia and islands in W Pacific).

2. Elodea Michx. (inc. Apalanthe, Egeria) Freshwater, roots unbranched; stems horizontal or stoloniferous; leaves uniforme, leaves sometimes submerse. 10 spp., three of which are widely distributed in the fresh waters of the Andes and temperate South America south of Ecuador, three from North America, two in subtropical and temperate fresh waters in S & SE Brazil (Minas Gerais and Espírito Santo states) to E Argentina, Uruguay and Paraguay, E. heterostemon (S.Koehler & C.P.Bove) Byng & Christenh. occurs only in Araguaia Basin center Brazil in Mato Grosso and Goiás states, and E. granatensis Humb. & Bonpl. from Ecuador to French Guiana, Bolivia and Brazil up to Rio de Janeiro and São Paulo states.

E. densa (Planch.) Casp. is almost cosmopolitan due to its invasive nature.

3. Ottelia Persson. Freshwater; roots unbranched, stems contracted; leaves differentiable in petiole and blade, linear at juvenile; stipules absent. 21 spp., 20 in Old World and one in New World, O. brasiliensis (Planch.) Walp., native to the fresh waters of the S South America, in over Brazil, Paraguay and NE Argentina.

3. SUBFAMILY HYDRILLOIDEAE (8/60–65) ‣ three tribes, Vallisnerieae (3/16, tropical and subtropical regions on both hemispheres, with their largest diversity in the Old World) does not occur in South America.

3.1 HYDRILLOIDEAE ▸ TRIBE NAJADEAE (2/c. 40) - outsider Hydrilla (1; temperate regions in Europe, North and E Africa, Asia and N and E Australia).

4. Najas L. Monoecious or rarely dioecious, perennial or rarely annual herbs erect, caulescent, glabrous, entirely submersed; leaves uniforme. 37 spp. in fresh waters worldwide, 10 spp. are native to the New World: three Central and South America; one from Caribbean and Brazil; two over Canada to Argentina; one in W South America; two Holarctic; one endemic to SE U.S.A.; 5 in Brazil, none endemics.

3.2 HYDRILLOIDEAE ▸TRIBE HALOPHILEAE (3/22) ‣ outsider Enhalus (1; tropical Asia to W Pacific).

5. Halophila Thouars. Marine perennials; stems creeping or stoloniferous; leaves differentiable in petiole and blade; stipules present. 19 spp., 4 spp. in Neotropics, mainly Caribbean, two in South America: H. baillonis Asch. ex Dickie, from Costa Rica to N. Venezuela, Caribbean, NE Brazil (Paraíba, Pernambuco and Piauí states), and H. decipiens Ostenf., the only seagrass that can be considered to be pantropic.

Both species occur in normal seawater (3,5%), and are restricted to warmer waters and were not found further south than Rio de Janeiro State, being especially common on the NE coast at depths down to 62 m; Den Hartog (1970a) refers to H. decipiens as a sciaphilous spp., occurring down to a depth of 85 m. We have examined material collected at depths ranging from 0 to 62 m below LWST level; when found among the extensive populations of Halodule wrightii, the spp. forms very dense, but small, colonies.

6. Thalassia Banks ex König. Marine herbs, leaves uniforme, submerged, long but strait; marine perennials dioecious. Two spp., T. hemprichii (Solms) Aschers. in Indo-Pacific Ocean and T. testudinum Banks & Sol. ex K.D.Koenig in to the salt waters of the Caribbean from Florida to Caribbean, Colombia and Venezuela.

ALISMATACEAE

§ FULLY CHLOROTROPHIC

Genera/spp. 18/c. 90 Distribution almost cosmopolitan, with the highest diversity in North and South America. Habit usually bisexual (sometimes unisexual: in Sagittaria monoecious or polygamomonoecious, in Limnophyton polygamomonoecious, in Burnatia dioecious), usually perennial (rarely annual) herbs. Aquatic or helophytic. Tuberous stem, corm or stolons sometimes present.

4 genera and 37 spp. in Brazil. All marsh or aquatic plants, leaves erect or floating. 8 genera in New World. The center of diversity of Echinodorus is the Neotropical region. All spp. of this genus occur in the Neotropics. Helianthium is a genus endemic to the Neotropics, occurring mainly in South America; Sagittaria is a genus distributed predominantly in the western hemisphere, with about 12 species occurring in the Neotropics. Some North American species are cultivated as pond ornamental plants.

Use Ornamental plants, aquarium plants, vegetables (corms and roots of Sagittaria, leaves of Limnocharis), forage plants. Some native spp. includes aquarium and pond ornamental plants. Some spp. of Sagittaria have an edible rhizomatous stem. Leaves of Echinodorus grandiflorus (Cham & Schltdl) Micheli are used in popular medicine. Pollen collecting bees pollinate many spp. of Echinodorus. Sagittaria and some spp. of Echinodorus present floral nectaries.

Key to genera of Neotropical Alismataceae

1. Yellow or yellowish flowers - 3

3. Leaves glaucous; carpels numerous, semicircular; styles absent; seeds with transversal ribs. ------------ Limnocharis

3. Leaves light green; carpels 3-8, linear-lanceolate; styles conspicuous; seeds glandular-pubescent ------------ Hydrocleys

1. White or more rarely pink flowers - 4

4. Pseudostoloniferous herbs. Inflorescence umbelliform with up to three whorls. ------------ Helanthium

4. Rhizomatous herbs. Inflorescence a raceme-like or panicle-like cyme or a panicle-like cyme with several whorls - 5

5. Flowers hermaphrodite or pistillate (gynodioecious species) ------------ Echinodorus

5. Flowers unisexual (monoecious plants), sepals without longitudinal ribs ------------ Sagittaria

SYSTEMATIC three clades, clade Alismateae (4/19, temperate regions on the Northern Hemisphere) does not occur in South America.

A. LIMNOCHARIS CLADE ‣ outsiders are Burnatia (1; tropical to S Africa), Butomopsis (1; tropical Africa, South Asia to N Australia), Albidella (1; tropical Africa, coastal areas along the Indian Ocean to tropical Australia, Yucatán and Cuba), Ranalisma (2; tropical Africa; India, SE Asia, S China), Caldesia (3; C, S and E Europe, Africa, Asia to tropical Australia), Limnophyton (7; tropical Africa, Madagascar, S and SE Asia, Malesia, Australia).

1. Hydrocleys Rich. Plants submersed; stems short, stolons; leaves basal; inflorescence few to many on long septate scape. 5 spp. from Mexico to South America; three of then are restricted to Brazil (4, none endemics) and adjacent Argentina, Paraguay, Uruguay and Bolivia.

2. Limnocharis Humb. & Bonpl. Emersed herbs; stems short, rhizomatous, often stoloniferous; inflorescence up to 10, on aseptate scapes, occasionally proliferating. Two spp., both all widely distributed from U.S.A. to S South America, and collected in Brazil.

B. TROPICAL CLADE ‣ all genera in South America.

3. Echinodorus Rich. Monoclinous emersed herbs; leaves erect. 38 spp. from New World., 32 occur in South America, 28 in Brazil, 8 endemics, some amphi-tropical; three spp. are rare plants in Brazil, by Plantas Raras do Brasil’s book.

4. Helianthium Benth & Hook. f. Monoclinous emersed herbs; leaves erect. Two spp., one restricted to Jamaica and Guadaloupe and H. bolivianum (Rusby) Lehtonen & Myllys from Mexico or U.S.A. to South America, up Brazil.

5. Sagittaria Rupp. ex L. Herbs perennial or rarely annual, submersed, emerse or floating leaves; inflorescence erect, floating or submersed, flowers imperfect. 29 spp. occurring in the New World.; 10 in South America, 6 in Brazil, none endemics; some North American spp. are cultivated as pond ornamental plants.

JUNCAGINACEAE

§ FULLY CHLOROTROPHIC

Genera/spp. 3/35 Distribution cold-temperate regions in the Northern and Southern Hemispheres. Habit bisexual, monoecious or polygamomonoecious (in Tetroncium dioecious), usually perennial (rarely annual) herbs. Aquatic or helophytic, in fresh or brackish water.

The Juncaginaceae occupy a variety of habitats. Triglochin usually occurs in marshes that are brackish or have high concentrations of marl, and in high mountain meadows of the Andes. A subcosmopolitan family of 5 genera and c. 13 spp.

SYSTEMATIC outsiders Cycnogeton (9; W Australia, Victoria), Maundia (1; Queensland, New South Wales).

1. Tetroncium Willd. Emeresed perennial with delicate, fibrous roots. Only one sp., T. magellanicum Willd., in Tierra del Fuego, Patagonia, and the Falkland Islands.

2. Triglochin L. (inc. Lillaea) Emersed perennial, annual or perennial ‘‘grass-like’’ herbs, occasionally tuberous; stems short, leaves few, characterised by sheathing leaves with ligules or auricles and spike-like inflorescences. c. 24 spp., cosmopolitan, five spp. native to the coastal marshes of the New World, one in Bolivia and Cono Sur; two scattered of Northern Hemisphere and W South America, and remaining two spp. in Brazil, both widely distributed in New World: T. scilloides (Poir.) Mering & Kadereit, native to temperate and tropical montane marshes, from Canada to Chile, Argentina and S Rio Grande do Sul state in S Brazil, mainly in high mountain meadows of the Andes, and T. striata Ruiz. & Pavon from S Africa, Oceania, North America, Caribbean, Peru to Uruguay and S Brazil.

Triglochin is probably best known for its coastal salt marsh species found in most temperate regions of the world; however, species of Triglochin occur in a wide variety of wet to dry habitats such as estuaries, seasonal wetlands (vernal pools, rock pools), and semi-desert inland habitats from sea level to up to ca. 5,000 m altitude, extremely abundant in soils so heavily laden with marl or salts that the soil surface is white, especially on highs.

RUPPIACEAE

§ SEA GRASSES (Heterozostera – Ruppia – Halodule - Syringodium – Halophila – Thalassia)

Genera/spp. 1/11 Distribution cosmopolitan except polar areas. Mainly in coastal marine environments. Habit bisexual, usually annual (rarely perennial) herbs; Aquatic in salt or brackish (rarely fresh) water. Submersed. Shoot apices usually not transformed into turions (present in Ruppia tuberosa J.S.Davis & P.B.Tomlinson).

A subcosmopolitan family of 1 genus (Ruppia L.) and 11 species, with about 1-7 species native to fresh and brackish waters of coastal as well as inland Neotropics. Ruppiaceae are useful in stabilizing substrates and removing suspended particles from the water column.

SYSTEMATIC a single genus.

1. Ruppia L. Annual or rarely perennial submersed glabrous herbs growing in fresh and brackish waters; stems slender, often dimorphic. 11 spp., 4 in New World, two in South America: R. filifolia (Phil.) Skottsb. from Ecuador to Cono Sur and R. maritima L., distibutelly over Brazilian coast, Piauí to Ceará, Pernambuco to Alagoas; Rio de Janeiro to Rio Grande do Sul, in brackish water ponds and lakes, with salinities varying from 0.3 to 28 ppm, and a widely distributed species worldwide.

CYMODOCEACEAE

§ SEA GRASSES (Heterozostera – Ruppia – Halodule - Syringodium – Halophila – Thalassia)

Genera/spp. 6/16 Distribution tropical and subtropical coastal marine areas; some spp. in warm-temperate seas (around Australia, in Mediterranean). Habit monoecious or dioecious, perennial herbs. Marine. Submersed. Rhizome in Cymodocea, Halodule and Syringodium herbaceous, monopodially branched, in Amphibolis and Thalassodendron lignified, sympodially branched.

A sub cosmopolitan family of 5 genera and c.16 spp. Two genera comprising 5 spp. occur in the Neotropics. Submerged marine aquatics growing in shallow waters. Leaves alternate (distichous), simple. Commonly known as sea-grasses or manatee grasses. Cymodoceaceae occur in shallow, coastal, tropical or subtropical waters. They can form extensive submarine meadows in shallow, clear waters with minimal wave action. Some are important in the stabilization of shallow marine sediments, nutrient recycling, and as food sources for grazing marine animals. Monoecious or dioecious, perennial herbs. Marine. Submersed. Rhizome in Cymodocea, Halodule and Syringodium herbaceous, monopodially branched, in Amphibolis and Thalassodendron lignified, sympodially branched.

Commonly known as sea-grasses or manatee grasses. Cymodoceaceae occur in shallow, coastal, tropical or subtropical waters. They can form extensive submarine meadows in shallow, clear waters with minimal wave action; some are important in the stabilization of shallow marine sediments, nutrient recycling, and as food sources for grazing marine animals.

Key to genera of Neotropical Cymodoceaceae

1. Leaf blade distinctly terete; many flowers in conspicuous cymose inflorescences ------------ Syringodium

1. Leaf blade linear, flat; inflorescence 1-flowered or 2 flowers resembling a single flower ------------ Halodule

SYSTEMATIC outsiders Cymodocea (4; coastal marine waters along Canary Islands, Mediterranean, W Africa, tropical Asia to the Pacific); Amphibolis (2; coastal marine waters along W and S Australia), Thalassodendron (3; coastal marine waters along E Africa including the Red Sea, W Indian Ocean, East Malesia and W and NE Australia).

1. Halodule Endl. Rhizome monopodially branched, herbaceous seagrasses; leaf blade narrow, linear with dentate apex. 6 spp., two in Old World, and 4 in New World, one endemic to Panamá; H. beaudettei (Hartog) Hartog, in the Madagascar, Gulf of Mexico, the Caribbean, and along the Pacific coast from Panamá to Mexico, and 5,500 km distant population on NE Brazilian coast; H. emarginata Hartog, endemic to SE Brazil (unique sea-grass endemic to Brazil), from central Piauí to the northern part of São Paulo states, on sheltered shores, on fine sediment substrate, in salinities around 35 ppml; and H. wrightii Asch., from W. trop. Africa, SE. U.S.A., Mexico to Venezuela and Colombia, Caribbean, E Brazil, and is the only sea grass found in all continents except Antartida, occurring around Atlantic Ocean, and only sea grass that occur in E, W Africa and Fernando de Noronha Is., Brazil (discovery in 2020).

On the northeast coast of Brazil, H. wrightii forms extensive beds on large areas behind the reef edges, being associated with several spp. of algae, particularly Siphonales, Dictyotales and Ceramiales. In some places, such as Ilha de Itamaraca, drifted leaves of this species pile on the beaches in large quantities.

2. Syringodium Kutz. Rhizome monopodially branched, herbaceous seagrasses; leaf blade terete. Two spp., S. isoetifolium (Asch.) Dandy in the Indo-West Pacific, and S. filiforme Kütz. from U.S.A. to Venezuela, Caribbean, common in Colombia coasts forming vast meadows.

ZOSTERACEAE

§ SEA GRASSES (Heterozostera – Ruppia – Halodule - Syringodium – Halophila – Thalassia)

Genera/spp. 3/21 Distribution temperate coastal marine areas in the Northern and Southern Hemispheres; some spp. in subtropical and tropical seas; almost absent around South America. Habit monoecious (Zostera) or dioecious (Phyllospadix), usually perennial (rarely annual) herbs. Marine (sometimes in brackish water). Submersed. Main stem with monopodial growth. Branches leaf-opposite.

The family generally occur in shallow coastal waters of temperate to subtropical regions, all three genera just reaching into the Neotropics; perennial plants can form extensive stands (often called submarine meadows) in shallow brackish and saltwater areas with clear water and minimal wave action. Zosteraceae play an important role in stabilization of shallow marine sediments.

SYSTEMATIC outsiders Phyllospadix (6; marine coastal waters from China, Korean Peninsula and Japan to W North America and NW Mexico); Nanozostera (7; marine coastal waters on both hemispheres).

1. Heterozostera L. Monoecious perennials with monopodially branched herbaceous rhizomes; two unbranched roots and a vegetative shoot with two to five leaves at each node; leaves alternate, distichous; male flower with two free bilocular anthers, female flower with ovary containing one ovule; fruit, an achene, ovoid to ellipsoid with a scarious pericarp; seed ovoid to ellipsoid. 4 spp., three in Oceania and New Zealand and H. chilensis J. Kuo in Chile, and is now found only in three bays: Herradura Bay, Salado Bay and Tongoy Bay.

POTAMOGETONACEAE

§ FULLY CHLOROTROPHIC

Genera/spp. 5/90–95 Distribution cosmopolitan except polar areas, western Europe, northern Africa, SW Asia; southern Australia, Tasmania, New Zealand; SW South Africa. Habit aquatic freshwater or brackish herbs; A family of 4-7 genera depending on generic and familiar circumscriptions adopted, and c. 90-95 spp. (an additional 40-50 putative hybrids of many Potamogeton have been described), occurring in fresh and brackish waters worldwide.

Fruits provide the most important features in separating species of this family; consequently one should always collect fruiting specimens; some species are important foods for waterfowl and are also grown in aquaria; environmentally some species are very important in substrate stabilization and in removing particles from the water column.

SYSTEMATIC outsiders Groenlandia (1; W Europe, N Africa, SW Asia), Lepilaena (4; S Australia, Tasmania, New Zealand), Althenia (2; Atlantic coasts of Morocco and SW Europe northwards to France, Mediterranean coasts, Türkiye, C Siberia, coastal waters in Namibia and SW South Africa to E Cape), Pseudalthenia (1; coastal areas in W Cape).

1. Potamogeton L. Submersed, emersed, or floating herbs, some in semiaquatic habits. c. 80 spp. worldwide, 41 in New World, up to Argentina; 12 in South America, 10 in Brazil, two endemics, some in high mountains of central to Brazil, e.g. P. scerocarpus K. Schum from high mountains of E Brazil, known only four collection by SW Bahia, NE Goiás and n Minas Gerais, and the only sp. of this family in Serra do Cipó.

2. Stuckenia Borner. Scale-like leaves in long sheats. 6 spp., subcosmopolitan, 5 occur in the Neotropics up to Argentina; two spp. occur in Brazil, each composedly of a unique recent record: S. filiformis (Persoon) Borner, in Junco Lake, Bahia state, and another of S. pectinata (L.) Börner, collected in the state of Rio Grande do Sul, both in 2011.

3. Zannichellia L. Submersed helophytes herbs in freshwaters and brackish pounds; leaves scale-like. 5 spp. near cosmopolitan, with two spp. in the Neotropics up to Argentina: Z. andina Holm-Niels. & R.R. Haynes from Ecuador and Bolivia, and Z. palustris L. from Flora N America, Mexico, Peru, Bolivia, Cono Sur and Brazil (S region, Paraná and Rio Grande do Sul states).

10. ACORALES

ACORALES DOES NOT OCCUR IN SOUTH AMERICA, AND IS COMPOSED OF A SINGLE FAMILY, ACORACEAE (1/2).

11. PETROSAVIALES

PETROSAVIALES DOES NOT OCCUR IN SOUTH AMERICA, AND IS COMPOSED OF A SINGLE FAMILY, PETROSAVIACEAE (2/3).

12. DIOSCOREALES

FAMILIES ABSENTS IN SOUTH AMERICA: AFROTHISMIACEAE (1/16).

Merckx and Smets (2014) found that Afrothismia was sister to Tacca plus other Thismiaceae, based on ML and Bayesian analyses of sequences of nrDNA 18S and mitochondrial atp1. Thus, three families (Taccaceae, Thismiaceae, and an undescribed one including Afrothismia) may ultimately need to be segregated from Dioscoreaceae s.l., which would swell the number of monocot families to 80.

LINEAGE 1 of 2: NARTHERCIACEAE

NARTHECIACEAE

§ FULLY CHLOROTROPHIC

Genera/spp. 5/c. 45 Distribution W and E U.S.A., W Europe, E Himalaya, E Asia, parts of W Malesia; Nietneria: Venezuela, Brazil and Guyana. Habit usually bisexual (rarely polygamous), perennial herbs. In one sp. of Aletris corm; in Lophiola stolons.

SYSTEMATIC outsiders Aletris (20–25; E Himalayas, E Asia, W Malesia, Canada, U.S.A.), Metanarthecium (1; Japan), Lophiola (1; SE U.S.A.), Narthecium (7; U.S.A., Japan, Türkiye to the Caucasus, W and N Europe, Corsica, Balkan Peninsula)

1. Nietneria Klotzsch ex Benth. Herbs perennial and rhizomes short; leaves distichous and unifacial, linear; inflorescence a loose terminal corymbe, six petaloid tapals; fruit a loculicidal capsule. Two spp., N. corymbosa Klotzsch & M.R. Schomb. ex B.D. Jacks and N. paniculata Steyerm., both widely distributed in the Guiana Shield in Venezuela, Guyana, and Mount Aracá in Amazonas state, Brazil, in savannas and elfin forests on top of table mountains, at elevations of 1,000-2,800 m.

LINEAGE 2 of 2: BURMANNIA + MYCOHETEROTROPHS + TACCA + DIOSCOREA

DIOSCOREACEAE

§ FULLY CHLOROTROPHIC

Genera/spp. 3(own data, excludes Tacca)/637 Distribution pantropical; some spp. in subtropical and warm-temperate regions (Stenomeris in W Malesia). Habit bisexual (the ‘Stenomeris clade’), or monoecious or dioecious, usually perennial herbs (often twining, creeping or climbing). Rhizome usually tuberous, starchy. Roots sometimes transformed into spines.

Key differences from similar families

ü from Smilacaceae:

No tendrils (tendrils from petiole in Smilacaceae).

Basal and apical pulvini (none in Smilicaceae).

Inflorescence a spike/raceme or derived form (not an umbel).

Dioecious

Ovary inferior (not superior).

Fruit a capsule or leathery/indehiscent (not a berry).

ü from Menispermaceae:

Campylodromous (not pinnate) venation.

Parts in 3s/6s, (Menispermaceae not so).

Closest relative Burmanniaceae is morphologically very reduced and usually heteromycotrophic.

SYSTEMATIC outsiders Stenomeris (2, Malesia, Indonesia, Philippines), Trichopus (2; Madagascar, S and C India, Sri Lanka, peninsular Thailand, the Malay Peninsula).

1. Dioscorea L. All geophytes with aereal short-lived branches, stem climbing, diecious. 633 spp., native throughout the tropical and warm temperate regions of the world, 421 in New World, 312 spp. in South America, with Brazil has the highest diversity (138, 93 endemics), follws by Mexico (75), China (48), Chile (40) and Madagascar (37); species of open areas occurs in southern Brazil and adjacent Argentina; D. dodecaneura Vell. has edible tubercules, feature rare in Neotropics. By recent phylogenetic works, two subgenera and 10 major clades.

§ subg. Dioscorea ‣ exclusive from Old World.

§ subg. Helmia ‣ 10 major clades, with New World members falling in four of them:

§ New World clade I ‣ former D. subg. Dioscorea, restricted to the Neotropics, with three small clades: Epipetrum, Microdioscorea and Nanarepenta. Whithin the species of NWI clade is possible to highlight D. perdicum Taub., one of the dwarf species endemic from Brazil that does not fit any specific infrageneric classification devotaded to this issue.

§ New World clade II ‣ corresponding to D. subg. Helmia restricted to the Neotropics; withn this group, sect. Dematostemon is one of the biggest sections of the Neotropical region, comprising 45 taxons of great morphological diversity and high distribution range, going from dwarf species (eg.: D. anomala (Kunth) Griseb. and D. maianthemoides Uline ex R. Knuth) endemic to the savannas of C Brazil (cerrado) to typical Atlantic Forest species (D. campanulata Uline ex R. Knuth and D. cinnamomifolia Hook.). D. basiclavicaulis Rizzini & Mattos-Filho, within sect. Chondrocarpa for this clade, is endemic to NE Brazil, and is the unique in this genus for its thickened succulent perennial stems.

§ New World clade III ‣ includes D. dodecaneura Vell., D. stegelmanniana R.Knuth, possibly D. trifida L.f., and species representatives of D. section Rajania form Caribbean.

§ sect. Opsophyton ‣ group exclusive from Old World except by D. mollis Kunth, endemic to forests of SE & S Brazil.

A multi-lineage group of smaller erect species, dwarf, usually less than 50cm, entirely or occasionally missing a climbing stem, appearing as self-supported or prostate herbs, are rare, with some spp. Mexico (2), Chile and mainly Africa savannah, with no parallel evolution in Asia; in Brazil most of them occur in high altitude open vegetation from central-northern to SE savannas of C Brazil (cerrado); D. anomala Griseb., D. maianthemoides Uline ex. R. Knuth and D. stenophylla Uline, in Goiás and Espinhaço Range of Minas Gerais state; D. perdicum Taub. in the Serra do Mar highlands of Minas Gerais and Rio de; D. sphaeroidea R. Couto & J. M. A. Braga, from high altitude grasslands in Rio de Janeiro; and D. compacta D. Araujo from Jalapao region, Tocantins.

TACCACEAE

§ FULLY CHLOROTROPHIC

Genera/spp. 1(absent in POWO)/19 Distribution tropical regions in the Old World east to Southwest Pacific islands, N South America. Habit bisexual, perennial herbs. Rhizome usually tuberous, starchy. Leaves basal.

SYSTEMATIC a single genus.

1. Tacca Seem. Herbs, perennials, from a solid, stemless, starchy, tuberous, globose, or elongate, vertical or horizontal rhizome, inflorescence a cymose pseodo-umbel, large foliaceous and pendent filiform bracts, flowers perfect and ovary 1-locular. 12 spp. intropical zone, only one, T. parkeri Spruce in New World, highly variable in blade of leaf, widely distributed in the Guiana Shield region and the Amazonian lowlands in Colombia, Venezuela, Guyana, Peru and shady places of northern Brazil, reaching into central savannas, in sand soils from lowlands; the anther wall development in Tacca, according to the ‘Dicotyledon type’ is otherwise known only in Acorus among the monocotyledons.

BURMANNIACEAE

§ MYCOHETEROTROPHICS (Arachnitis – TRIURIDACEAE – BURMANNIACEAE – THISMIACEAE – ORCHIDACEAE – Voyria - Voyriella - Monotropa)

Genera/spp. 8(excludes Thismiaceae and Afrothismiaceae)/106 Distribution tropical and subtropical regions, with the largest diversity in South America: SE North America, Caribbean and Central America southwards to Peru and Uruguay, central and SE Africa, Madagascar, E and SE Asia, Malesia to SE Australia. Habit bisexual, perennial or annual herbs. Green autotrophs or hemiparasites (Burmannia) or achlorophyllous holoparasites (mycoheterotrophic) on fungi. Roots often tuberous, sometimes coral-like.

Small herbs, mostly mycotrophics centered in N South America. From the southern U.S.A. and Mexico in the North to S Brazil and N Paraguay in the South, also in Caribbean; almost all genera are American. Could be confused with Triuridaceae, both sharing alternate, scale-like leaves and being almost exclusively myco-heterotrophic, but Triuridaceae have flowers with many, free carpels (apocarpous) and flowers are often unisexual. All genera, except for most species of Burmannia, are myco-heterotrophic herbs without chlorophyll; reduced, scale-like leaves.

SYSTEMATIC outsiders is Marthella (1; Mount Tucuche on Trinidad).

1. Apteria Nutt. Small herb, leaves scale-like; cincinnus 1-5 flowered, flowers purple, funnel-shaped to campanulate. Only one sp., A. aphylla (Nutt.) Barnhart ex Small., S U.S.A. to Bolivia, Argentina, Paraguay and SE Brazil, also in Caribbean, mainly in wet forests, highly variable, particularly in its flower shape and size.

2. Burmannia L. Herb with ciccinus bifurcate, 1-27 flowered, flowers bicolor, yellow, white or blue, tubular to salver-shaped. c. 63 spp., Asia (30, 19 holomycots), Africa (11, 3 holomycots), America (20, 1 holomycots) and Australia (two autotrophic); all spp. are terrestrial, except for B. kalbreyeri Oliv. from Costa Rica to S Ecuador and Venezuela, growing epiphytically on various trees; 20 neotropical species, 19 in South America, 12 in Brazil, 4 endemics; B. tenella Benth. from N South America up to Brazil and Bolivia is the only entirely achlorophyllous member of its genus in the New World; however, closely related species also show remarkable reductions according to number and surface area of leaves, representing a conclusive model of how evolution may have taken place; it is very likely that many members of the New World species depend more on their mycotrophy than on own photosynthetical activity.

3. Campylosiphon Benth. Small, pale blue to white herbs; ciccinnus bifurcate, 1-14 flowered, salver-shaped, pale blue to white. Two spp., C. purpurascens Benth. in Venezuela, Colombia, Guianas, Brazil, Peru, and C. congestus (C.H.Wright) Maas Guinea to RD Congo; in rain forests, often on marshy spots.

4. Dictyostega Miers. Small whitish herb with bifurcate flowers; ciccinnus bifurcate, 6-63 flowered, flowers tubular, white, nodding. Only one sp., D. orobanchoides (Hook.) Miers, from Mexico to Bolivia and SE Brazil, absent in Caribbean.

5. Gymnosiphon Blume. Small whitish herbs up to 30 cm, flowers salver-shaped, white to yellowish. 16 spp. in Neotropics (12 in South America, 7 in Brazil, none endemics), 8 in Africa (inc. Comores and Madagascar) and 8 in Asia; in forests, often between the roots of large trees and close to streams; G. capitatus (Benth.) Urb. from Uaupes riveside in Amazonas state, is a rare species in Brazil, by Plantas Raras do Brasil’s book; however, VPA assignated this species also in Guianas and Colombia.

6. Hexapterella Urb. Small herb with purplish stems; ciccinnus bifurcate, 1-8 flowered, salver-shaped to tubular, white to purple. Two spp., H. gentianoides Urb. from Trinidad, the Guianas, Colombia, Brazil, Venezuela, and H. steyermarkii Maas & H. Maas endemic to Venezuela, in rain forests, rarely in swampy savannas.

7. Miersiella Urb. Small herb up to 20 cm, leaves scale-like, almost peltate; inflorescence umbel-like, 4-22, flowers tubular, white to purple. Only one sp., Miersiella umbellata (Miers) Urb., growing in dense rainforest in SE Brazil, and Amazon Guyana to Peru.

THISMIACEAE

§ MYCOHETEROTROPHICS (Arachnitis – TRIURIDACEAE – BURMANNIACEAE – THISMIACEAE – ORCHIDACEAE – Voyria - Voyriella - Monotropa)

Genera/spp. 3(absent in POWO)/60 Distribution tropical West Africa, Sri Lanka, Southeast Asia, Malesia, southeastern Australia, New Zealand, locally in eastern North America, tropical Central and South America to Bolivia, with their largest diversity in Brazil. Habit bisexual, perennial or annual herbs. Achlorophyllous mycoheterotrophic holoparasites.

SYSTEMATIC outsiders are Oxygyne (4; Japan; Cameroon), Haplothismia (1; W Ghats in S India).

1. Thismia Griff. Myco-heterotrophic small herbs, typically growing among leaf litter on the shady wet forest floor; root system tuberous or rarely cylindrical, roots filiform; stems unbranched; leaves (2-)4, alternate, scale-like, sometimes forming a rosette just below the flower; flowers terminal, solitary, 5-20 mm long, bisexual, actinomorphic to sometimes zygomorphic, fruit cup-shaped, fleshy; seeds many, small and ‘dustlike’, (narrowly) ellipsoid to ovoid. 59 spp., 37 in tropical and temperate Asia, 24 in the New World, 23 in South America, 20 in Brazil, 13 endemics; two subgenera.

§ subg. Ophiomeris ▸ Neotropical Thismia; 21 spp. in three sections; all species in Brazil except T. saulensis H.Maas & Maas and T. andicola Aguilar-Cano, S. Guzman-Guzman & Lopera-Toro.

§ sect. Myostoma ▸ a single species, T. hyalina (Miers) Benth. & Hook.f. ex F.Muell. in Brazil (Acre, Amazonas, Mato Grosso, Pará, Rio de Janeiro and São Paulo states) and Peru.

§ sect. Ophiomeris ▸ 16 spp., 13 in Brazil, 8 endemics: T. espiritosantensis Brade, T. mantiqueirensis Engels & E.C.Smidt., T. iguassuensis (Miers) Warm., T. janeirensis Warm., and T. macahensis (Miers) F.Muell. are restricteds to São Paulo, Rio de Janeiro and Espírito Santo states in SE Brazil, T. luetzelburgii Goebel & Suess. occur in SE Brazil disjunct in Panamá, T. glaziovii Poulsen disjunct in SE Brazil and SE Colombia, T. prataensis Mancinelli, C.T.Blum & E.C.Smidt and T. cordata D.F.Silva & J.M.A.Braga endemic to Paraná state, SE Brazil; T. variabilis D.F.Silva, Honório & J.M.A.Braga are endemic to Acre state, N Brazil; T. calcarata D. F. Silva, Honorio & J. M. A. Braga in Acre state in Brazil up to Peru; T. panamensis (Standl.) Jonker known in Brazil in Paraná state, also in Bolivia, Ecuador, Colombia, Costa Rica and Panamá; T. paradisiaca Guzman & Toro is known only from Pacific slopes in Colombia; T. violacea D.F.Silva & J.M.A.Braga endemic to Goiás state (Brazil); T. andicola Aguilar-Cano, S. Guzman-Guzman & Lopera-Toro endemic to Andean in N Colombia, and T. saulensis H.Maas & Maas endemic to center French Guiana.

§ sect. Pyramidalis ▸ 7 spp., all but one endemics to Brazil: T. caudata Maas & H.Maas and T. fungiformis (Taub. ex Warm.) Maas & H.Maas are rare species in SE Brazil; T. melanomitra Maas & H.Maas known from the Ecuador, Mato Grosso and Pará states in Brazil; T. pseudomelanomitra D.F.Silva & J.M.A.Braga endemic to Mato Grosso state; T. ribeiroi Engels, D. Ferreira-da-Silva & Soares-Lopes and T. petasiformes D.F.Silva & J.M.A.Braga are endemics to N Mato Grosso state in Brazil, in Amazon forest; and T. singeri (de la Sota) Maas & H.Maas is known from Pará and N Mato Grosso in Brazil, Mamore Basin in Bolivia and Napo region of Ecuador.

§ subg. Thismia ▸ 37 spp., one endemic to U.S.A., T. americana N.Pfeiff. collected few times near Chicago, U.S.A. (latter in 1917), remaining in tropical Asia to Australia and Tasmania.

2. Tiputinia P.E. Berry & C. Woodw. Herb with 2 cm tall, olive yellow; produces a foul, rotten fish-like odor. Only one sp., T. foetida P.E. Berry & C. Woodw., known from Amazonian forest of Ecuador and also disjunct in S Peru (collected in 2020).

13. PANDANALES

FAMILIES ABSENTS IN SOUTH AMERICA: PANDANACEAE (5/947), STEMONACEAE (4/39).

VELLOZIACEAE

§ FULLY CHLOROTROPHIC

Genera/spp. 5(own data, excludes Nanuza)/c. 280 Distribution Panamá and NW South America, Peru to Argentina, Madagascar, tropical and southern Africa, the SW Arabic Peninsula (one spp. from Yemen and Saudi Arabia), with the largest diversity in SE Brazil; Acanthochlamys: SE Tibet, W Sichuan (China). Habit usually bisexual (in Barbaceniopsis unisexual), usually perennial herbs (sometimes shrubby or almost tree-like). Xeromorphic. Stem more or less woody, covered with persistent leaves and leaf sheaths usually intermingled with extensive adventitious roots.

This family has six genera and 224 spp. in South America and Central America and only 29 in Africa; more than 70% of the spp. (173) are concentrated in Minas Gerais, essentially on the Diamantina Plateau and Serra do Cipó. The data presented by A. Giulietti et al. (1987) show that among the 59 spp. of Velloziaceae found on the Serra do Cipó, 46.5% are restricted to that range, and 91.5% are endemic to the Espinhaço Range; 27 have been described in the last 23 years. In the Grão-Mogol area, 19 spp. of Velloziaceae include nine endemics and three new taxa (Mello-Silva 1989).

The Velloziaceae form a phytogeographical link between E African and Malagasy inselbergs and South American ones. Certain spp. show a broad range in elevation. In Brazil, Vellozia plicata Mart. colonizes rocky substrates from the sea-border up to more than 1,500 m. Altogether between 250 - 300 spp. of desiccation-tolerant vascular plants out of 13 families (4 monocots) are found on inselbergs; Velloziaceae contains more desiccation-tolerant spp. than any other family, with over 200 spp. These spp. occur mainly on rock outcrops, and almost all of them seem to be desiccation-tolerant. The resinous stems of the larger spp. of Vellozia (‘canela-de-ema’) were used as fuel for locomotives on the Curvelo-Diamantina railway and these days are stacked to sell as an effective fire lighter for wood stoves and to make torches.

The Velloziaceae are quite distinct from every other family. The stem could resemble that of some Agavaceae, Dracaenaceae or Pandanaceae, but in those families it is not covered by adventitious roots along with either persistent leaf sheaths or entire marcescent leaves. Some Bromeliaceae and few Cyperaceae could present a stem like that of the Velloziaceae, but those families, as well as the Agavaceae, Dracaenaceae and Pandanaceae, have conspicuous inflorescences.

SYSTEMATIC outsiders Xerophyta (c 30; tropical Africa, Madagascar, SW Arabian Peninsula), Acanthochlamys (1; SE Tibet, W Sichuan).

Key to genera of neotropical Velloziaceae

1. Flowers diclinous, style shorter than stigmas ------------ Barbaceniopsis

1. Flowers monoclinous, style longer than stigmas - 2

2. Corona present; stamens 6; stigmas vertical, fused at apex or free, lateral ------------ Barbacenia

2. Corona absent; stamens 6-76; stigmas horizontal, fused at center ------------ Vellozia

1. Barbacenia Vand. Plants of small or medium stature; caudex usually simple or few-branched; leaf-blades longpersistent; flowers perfect; perianth forming more or less of a tube above the ovary; coronoid appendages free from the tepals but usually adnate to the filaments and making them appear broad and flat, usually bilobed. 103 spp., 102 endemics to Brazilian Plateau, highly centered in Espinhaço Range of Minas Gerais state (74 spp. in this state, also in mountains and coast of Espírito Santo), only 6 reaching to NE region (only in Bahia state), 10-12 into WC region, and one into S region (Paraná state, endemic), three only in Tocantins state, and one only single spp., B. celiae Maguire, restricted of south Venezuela (Cerro Yavi) and W Roraima state, common on conglomerate outcrop, northwest escarpment of Serra Tepequem, Roraima, Brazil, 860-960 m alt; all tested species in dry-resistence studies are desiccation tolerants; 26 spp. are considered rare in Brazil, by Plantas Raras do Brasil’s book.

2. Barbaceniopsis L.B.Smith. Xerophytic shrubs; stems simple or few branched; flowers solitary a long pedicels. 4 spp., Bolivia, Argentina (Jujuy) and Peru (Arequipa and Pasco) one endemic each, and B. boliviensis (Baker) L.B. Sm. from Bolivia/Argentina; this species is the highest known site in the family, in the Andes of Cochabamba, Bolivia, at 2,900 m; all tested species in dry-resistence studies are desiccation tolerants.

3. Vellozia Vand. Stemless herbs from 6 cm to tree-like up 6 m tall, mostly shrubs, some Agave-like and Dracaena-like (all tested species in dry-resistence studies are desiccation tolerants), a half from this ones from vascular diversity worldwide; leaves sometimes persistent after dry; latex inflammable in some spp. 1118 spp., 113 endemics to Brazil (22 are considered rare in Brazil, by Plantas Raras do Brasil’s book), V. andina Ibisch, Vasquez & Nowicki endemic to Bolivia, V. caruncularis Mart ex. Seub., V. sellowii Seub and V. variabilis Mart., known in Brazil and Bolivia, mainly Noel Kempff National Park and vicinities; and V. tubiflora Mart. ex Schult. & Schult.f., widely distributed from Panama to SE Brazil.

V. tubiflora occur in two disjunct areas: N Brazil (Mount Neblina and Mount Aracá), Colombia (only in Içana Valley), Guyana (westernmost point), Venezuela (Bolivar and Amazonas states) and N Panamá (eastern flanks of the cerro Chiriqui, where the vegetation is similar to that seen in the high mountains in Colombia), near closed Costa Rica; Brazilian Shield in mountains in Goiás and in the Distrito Federal, mountains of W Minas Gerais, reaching the region around Serrana and Cajuru, in north-central São Paulo State, in the mountains which separate the states of Ceará and Piauí, in center Diamantina Range, Bahia, in the Serra do Estrondo, in Tocantins, in the Serra do Cachimbo and the waterfalls of the Juruena and Cururu rivers, both along the border between the states of Mato Grosso and Pará, as well as in Bolivia (Chiquitos sandstone mountain chains in the eastern lowlands), in the lower elevations in the Paraguá River Basin).

V. auriculata Mello Silva & N.L.Menezes rather weak stems tend to collapse and, where there is ground contact, take root and form a new tree - a rather unusual form of vegetative propagation for the genus.

V. pyrantha A.A.Conc is a dominant species in extensive areas of rocky grasslands (campos rupestres); this dominance is so great that the name of one of the mountains where it occurs is called Serra do Candombá; these shrublands are fire-prone, and the species is resistant to fire, resprouting after only a few days, exhibiting mass flowering 30–50 days after burning; the species has been collected with flowers only after fires, which is a clear evidence of fire-dependent flowering; the rapid and abundant flowering after fire has also been observed in other species of Vellozia, such as V. peripherica Mello-Silva and V. gigantea N.L. Menezes & Mello-Silva, but only in V. pyrantha the flowering seems to be fire-dependent, as in V. alata L.B. Sm. and V. piresiana L.B. Smith.

By extrapolating year growth rates and measuring the height of individuals in the field, Alves (1994) calculated an age of 551 years for a V. kolbekii Alves specimen 3 m tall.

TRIURIDACEAE

§ MYCOHETEROTROPHICS (Arachnitis – TRIURIDACEAE – BURMANNIACEAE – THISMIACEAE – ORCHIDACEAE – Voyria - Voyriella - Monotropa)

Genera/spp. 8/c. 50 Ditribution Central America, northern and central South America, tropical W Africa, N Madagascar, E and SE Asia, Malesia, NE Australia, islands in the SW Pacific. Habit usually monoecious, polygamomonoecious or dioecious (in spp. of Sciaphila bisexual), usually perennial (rarely annual) herbs. Achlorophyllous mycoheterotrophic holoparasites.

Peltophyllum caudatum (Poulsen) R.Schmid & M.D.Turner and Triuridopsis peruviana H.Maas & Maas are both only known from a single collection each!.

SYSTEMATIC three tribes and thrre unplaced genera; tribe Kupeeae (2/2; Bakossi Mountains in W Cameroon, Udzungwa Mountains in Tanzania) does not occur in South America.

Key to genera of neotropical Triuridaceae

1. Flowers bisexual; stamens placed in the center of the flower within the carpels ------------ Lacandonia

1. Flowers unisexual or rarely bisexual, but than the stamens always placed outside and around the carpels - 2

2. Plants monoecious; flowers unisexual (2 Sciaphila bisexual); tepals bearded; fruit dehiscent or indehiscent - 3

3. Staminate flowers with 2 stamens and 4 tepals; fruit an indehiscent achene ------------ Soridium

3. (Staminate) flowers with 3 or 6 stamens, and 4 or 6 tepals; fruit a dehiscent follicle ------------ Sciaphila

2. Plants dioecious; flowers unisexual; tepals long-tailed; style apical; fruit indehiscent - 4

4. Staminate flowers with anthers sessile on a conical androphore ------------ Triuris

4. Staminate flowers without conical androphore - 5

5. Staminate flowers with 3 tepals, 6 filamented, monothecal anthers and a sterile projection in the centre of the flower ------------ Triuridopsis

5. Staminate flowers with 6 tepals, 3 sessile, dithecal anthers and no sterile projection in the centre of the flower ------------ Peltophyllum

UNPLACED GENERA

1. Peltophyllum Gardn. Unbranched herbs with filiforme rhizomes, hyaline, flattened stems and scale-like leaves, dioecious, inflorescence a simple raceme. Two spp., P. luteum Gardner in SE Brazil and adjacent Argentina and Paraguay (possibly in Guyana), and P. caudatum (Poulsen) R. Schmid & M.D. Turner very narrow endemic, possibly extinct, in Alto do Macaeh, Rio de Janeiro, as a rare plant in Brazil by Plantas Raras do Brasil’s book, known only the type collection made by Glaziou.

2. Soridium Miers. Unbranched or basally branched herbs with rhizomes and scale-like leaves, monoecious; inflorescence a simple raceme. Only one sp., S. spruceanum Miers, from Guatemala in the North to Brazil (Mato Grosso) in the South.

3. Triuridopsis H. Maas & Maas. Unbranched dioecious herbs with white flowers in a raceme; scalilike leaves. Two spp., Peru and Bolivia one endemic each.

1.1 TRIBE TRIURIDEAE (1-2/3-6) ‣ all genera in South America.

4. Lacandonia E.Martínez & Ramos. Hyaline herbs with rhizomes, simple stems, and scale-like leaves, monoecious, flowers in a simple raceme, made unique in the flowering plants by its stamens which are placed in the centre of the flower, surrounded by the apocarpous carpels. Two spp., L. schismatica E. Martínez & Ramos in lowland rainforsts in Chiapas state in Mexico, and L. brasiliana A. Melo & M. Alves in lowlands in Atlantic Forest in Paraíba and W Ceará state in NE Brazil, with relatively few morphological differences separate both spp.

Angios Bergianska: ‘Lacandonia, with two species described, represents a mutation in Triuris causing shift of organ positions (heterotopy). The androecium is central and the gynoecium peripheral, i.e. the carpels are extrastaminal. The carpellary and staminal precursors in Lacandonia develop from a common primordium. In Triuris, the precursors are formed from compound primordia.’

5. Triuris L. Unbranched herbs with short rhizomes, hyaline, flattened stems and 0-2 scale-like leaves, dioecious, inflorescence a raceme; mostly single (occasionally 2-3) mushroom-like, whitish flowers at the end of a leafless hyaline stem; the most distinctive feature of the flowers is the three long tails that hang down and spread out from the ‘cap’. 4 spp., T. hyalina Miers is disjunct in Mexico, Amazon rainforest from Colombia and Peru to C Brazil in Amazonas, Pará and Mato Grosso states, and from Bahia to Paraná states; T. hexophthalma Maas and T. alata Brade are only known from the type locality in the Parakaima mountains in Guyana (possibly populations in Peru) and Itatiaia Massif in SE Brazil (is a rare plant in Brazil, by Plantas Raras do Brasil’s book), respectively; and T. brevistylis Donn. Sm., restricted for Central America.

1.2 TRIBE SCIAPHILEAE (1-2/3-6) ‣ a single genus.

6. Sciaphila Blume. Simple or rarely branched erect herbs with elongated and often reddish stem, rhizome and scale-like leaves, up to 1.5 m, monoecious; bearing a spicate or narrow racemose inflorescence of small multi-pistillate flowers, the fruits are densely clustered minute, round, reddish follicles. 36 spp., 8 in New World, all in South America, six in Brazil, with recent and abundant records of N Mato Grosso state, Brazil; two spp. are known in W Africa, and 19 occurs in Asia (inc. Fiji and New Caledonia); S. purpurea Benth. (Peru to French Guiana and Brazil) can reach a height of almost 1.5 m, the tallest of all holomycotrophs; it is often found growing on termites nests; the two endemic Brazilian species are S. oligantha Maas e S. rubra Maas, both rare plants in Brazil, both highly restricted of Reserva Ducke, Manaus, Amazonas state, Brazil, by Plantas Raras do Brasil’s book.

CYCLANTHACEAE

§ FULLY CHLOROTROPHIC

Genera/spp. 12/228. Distribution tropical Central and South America, Caribbean. Habit monoecious, perennial herbs, suffrutices or lianas. Some spp. are epiphytes, leaves bifid. Use ornamental plants, textile plants, carpets, thatching, basketry, handicraft, medicinal plants (Asplundia). The famous Panamá hat is made of leaf fibres of Carludovica palmata Ruiz & Pav. leaves.

Cyclanthaceae is certainly monophyletic, despite considerable morphological and anatomical differences between the subfamilies Cyclanthoideae (monotypic) and Carludovicoideae, and well delimited from the closely related families in Pandanales. Similarities with the unrelated Arecaceae and Araceae are only superficial.

SYSTEMATIC two subfamilies, both in South America.

Key to genera of neotropical Cyclanthaceae

1. Leaf blades not plicate; staminate and pistillate flowers in alternate cycles (subfam. Cyclanthoideae) ------------ Cyclanthus

1. Leaf blades plicate; staminate and pistillate flowers in spirally arranged groups (subfam. Carludovicoideae) - 2

2. Leaves arranged in 2 rows (distichous); petiole ± elliptic in c.s.; placentation subapical or apical - 3

3. Leaf blades entire ------------ Ludovia

3. Leaf blades bifid - 4

4. Pistillate flowers and fruits free; placentation subapical ------------ Chorigyne

4. Pistillate flowers and fruits connate; placentation apical - 5

5. Placenta one; seeds broadly ellipsoid to narrowly oblong ------------ Sphaeradenia

5. Placentas four; seeds fusiform ------------ Stelestylis

2. Leaves spirally arranged; petiole adaxially flattened; placentation parietal - 6

6. Pistillate flowers and fruits free - 7

7. Leaf blades tricostate, not scabrous; stamens with basal bulb ------------ Schultesiophytum

7. Leaf blades unicostate or subtricostate, scabrous; stamens ± lacking basal bulb - 8

8. Staminate flowers with perianth lobes in two rows; tepals acute to shortly acuminate; seeds strongly flattened ------------ Evodianthus

8. Staminate flowers with perianth lobes in one row; tepals-acuminate; seeds somewhat flattened ------------ Dianthoveus

6. Pistillate flowers and fruits connate - 9

9. Adult leaves with four segments; fruits in a layer irregularly splitting from rachis ------------ Carludovica

9. Adult leaves with two segments, each segment sometimes secondarily split; fruits not in an irregularly splitting layer - 10

10. Spathes clustered; seeds terete ------------ Dicranopygium

10. Spathes dispersed; seeds strongly flattened - 11

11. Spathes diminishing in size upwards; seed coat smooth ------------ Asplundia

11. Spathes diminishing in size downwards; seed coat striated ------------ Thoracocarpus

1. SUBFAMILY CARLUDOVICOIDEAE (11/230–235) ‣ outsider Chorigyne (7, Costa Rica and Panamá).

1. Asplundia Harling. Roots climbing lianas or perennial herbs, with short to long stems. 100 spp., S Mexico to Bolivia and SE Brazil, and the Lesser Antilles; 90 in South America, 26 in Brazil, 13 endemics.

2. Carludovica Ruiz & Pav. Tall, terrestrial, and short stemmed plants; leaves spiral; leaf blade flabelliform-parted, trichostate. 4 spp., S Mexico to center Bolivia and Venezuela, also in Guianas, centered in Central America; C. drudei Mast. and C. palmata Ruiz & Pav. in South America, both widely distributed.

3. Dianthoveus Hammel & Wilder. Terrestrial, tall herbs with rather short stem; leaf blade bifid, unicostate or indistinctly subtricostate. Only one sp., D. cremnophilus Hammel & G.J. Wilder, SW Colombia to N Ecuador.

4. Dicranopygium Harling. Usually small to medium sized terrestrial herbs, with more or less short, fleshy stem. 53 spp. of S Mexico to S Peru and French Guiana, and the Lesser Antilles; 49 spp. in South America, two in Brazil (Roraima, Amazonas and Acre), one endemic.

5. Evodianthus Oerst. Roots climbing lianas with slender, branched stem, or terrestrial plants, mostly unbranched stem. Only one sp., E. funifer (Poit.) Lindm., from S Nicaragua to S Peru and E Brazil, and the Lesser Antilles; disjunct populations occurs in S Bahia state, Brazil.

6. Ludovia Brongn. Root climbing vines, epiphytes, or terrestrial herbs. Three spp., S Nicaragua to C Peru and French Guiana, two only from Central America to Ecuador; L. lancifolia Brongn. is only a truly epiphytes in family, and occur in all genus range.

7. Schultesiophytum Harling. Terrestrial plant, stem rather short; branching moopodial; leaf bifid, tricostate. Only one sp., S. chorianthum Harling., S Colombia and Ecuador to C Peru.

8. Sphaeradenia Harling. Medium-sized to tall terrestrial or epiphytic herbs, rarely vines. 51 spp., S Nicaragua to W Bolivia and Venezuela, and adjacent Brazil in Amazonas state (2, none endemics), 40 spp. in South America.

9. Stelestylis Drude. Short stemmed terrestrial, or epiphytic herbs; branching monopodial; leaf blade bifid, monocostate. 4 spp., S. anomala Harling endemic to Venezuela, S. coriacea Drude endemic to Roraima state in N Brazil, S. surinamensis Harling restricted for Guianas, and S. stylaris (Gleason) Harling in all three regions (in Brazil also only in Roraima state).

10. Thoracocarpus Harling. Root climbing lianas, sometimes hemiepiphytes. Only one sp., T. bissectus (Vell.) Harling, from Costa Rica to Bolivia and SE Brazil (so far São Paulo state), and the Lesser Antilles; only true liana in family.

2. SUBFAMILY CYCLANTHOIDEAE (1/1) ‣ a single genus and species.

11. Cyclanthus Pit. Ex A. Rich. Tall, usually terrestrial herb with short stem, terrestrial; leaf deeply bipartite. Only one sp., C. bipartitus Poit., S Mexico to NW Bolivia and French Guiana, and the Lesser Antilles; in Brazil occurs mainly in wet soils.

14. LILIALES

FAMILIES ABSENTS IN SOUTH AMERICA: CAMPYNEMATACEAE (2/4), COLCHICACEAE (15/281), LILIACEAE (15/860), PETERMANNIACEAE (1/1), RIPOGONACEAE (1/6).

LINEAGE 1 of 3: CORSIIDS

CORSIACEAE

§ MYCOHETEROTROPHICS (Arachnitis – TRIURIDACEAE – BURMANNIACEAE – THISMIACEAE – ORCHIDACEAE – Voyria - Voyriella - Monotropa)

Genera/spp. 2/26 Distribution S China (extinct), New Guinea, Solomon Islands, northern Queensland (Australia), southern South America. Habit bisexual (extremely protandrous), perennial herbs; achlorophyllous mycoheterotrophic holoparasites.

SYSTEMATIC outsider Corsia (c. 24, S China, New Guinea, Solomon Islands, N Queensland).

1. Arachnitis Phil. Herbs, mycoheterotrophic with short, tuberous roots; leaves alternate, 4-6, reduced to closed sheaths; flowers bisexual, zygomorphic, terminal, solitary; tepals 6, petaloid; fruits reflexed capsules, opening with 1 terminal triangular aperture; seeds many, dust-like. Only one sp., A. uniflora Phil., temperate forests of Patagonia in Andes and Tierra del Fuego, only mycoheterotrophic in Chile, sandy areas of the Falkland Is., and semi-humid and cloud forests of the Bolivian Andes, and in Peru; in this country, it is only known from Machu Picchu, where two populations are known, the presently recorded and another on the other margin of the Vilcanota river close to the Wiñay Wayna locality.

LINEAGE 2 of 3: LILIIDS

SMILACACEAE

§ FULLY CHLOROTROPHIC

Genera/spp.: 1/261 Distribution tropical, subtropical and warm-temperate regions in the Northern and Southern Hemispheres. Habit usually dioecious, twining evergreen shrubs, suffrutices or perennial herbs. Tuberous stem present in some spp. Lateral ligules and/or spines or pairwise petiolar tendrils present. Aerial roots absent. Rhizome woody.

Smilacaceae is similar to Dioscoreaceae in comprising vines with net-veined leaves; however, Smilacaceae has:

ü Leaves with petiolar sheaths terminating in a pair of tendrils.

ü Inflorescences an umbel or panicle of umbels.

ü Fruit a berry with 1-3 seeds.

Whereas family Dioscoreaceae has:

ü Leaves without petiolar sheaths nor tendrils.

ü Inflorescences mostly spicate, racemose, or paniculate.

ü Fruit a 3-winged loculicidal capsule.

SYSTEMATIC a single genus worldwide.

1. Smilax L. Vines, usually climbing by paired tendrils, or rarely erect herbs, often with rhizophores; stems rounded or sometimes quadrangular, armed with spines or unarmed, the surface smooth, scabrous, villose or setose; leaves simple, alternate, margin entir; inflorescences pedunculate, axillary, an umbela or panicle of umbels; flowers unisexual (the plants dioecious); fruit a fleshy and globose berry, indehiscent, smooth, orange, yellow, or black when mature; seeds 1-3. 261 spp., tropical, subtropical, and temperate areas of both hemispheres, with most spp. concentrated in the New World - 124 spp. - and tropical Asia, throughout the Neotropics, up to Argentina, occupying several habitats at elevations from 0 to 3,200 meters; 73 spp. in South America, 32 in Brazil, 15 endemics; in the Neotropics the rhizomes of most spp. of Smilax are used medicinally, but especially S. officinalis Kunth, S. longifolia Rich., S. siphilitica Humb. & Bonpl. ex Willd., S. aristolochiifolia Mill., S. moranensis M.Martens & Galeotti and S. domingensis Spreng. ex A. DC; the stems of some spp. such as S. tomentosa Kunth are used in basket-making.

MELANTHIACEAE

§ FULLY CHLOROTROPHIC

Genera/spp. 14/c. 170 Distribution temperate Eurasia, Mediterranean, Himalaya, NE India, Burma, E Asia, Indochina, northern Andes. Habit bisexual, andromonoecious, polygamous, dioecious, androdioecious, or gynodioecious, evergreen or deciduous perennial herbs (rarely somewhat lignified at base). Sometimes with a bulb without nutrient-storing scales (in some spp. of Schoenocaulon a corm). Schoenocaulon (c 25) Northerly temperate regions southwards to the northern Andes (one sp. in South America, disjunct from N Venezuela).

The seeds of Schoenocaulon contain alkaloids which are used medicinally; they are dissolved in acetic acid and frequently used as insecticides.

SYSTEMATIC groups Helonias Clade (3/13, E U.S.A., Himalayas, E Asia), Chionographideae (2/7, E U.S.A., E Asia), Xerophylleae (1/2, North America) and Parideae (3/c 73, temperate and mountainous regions on the Northern Hemisphere) do not occur in South America; among Melanthieae, outsiders are Zigadenus (1; SE U.S.A.), Toxicoscordion (9; S Canada, W and C U.S.A.), Anticlea (11; Mongolia, China, Korean Peninsula, Japan, Russian Far East, Canada, U.S.A., Mexico, Central America to Guatemala), Stenanthium (5; SE and E U.S.A.), Melanthium (4; C and E U.S.A.), Veratrum (25–30; temperate regions on the Northern Hemisphere), Amianthium (1; E U.S.A.).

1. Schoenocaulon A. Gray. Herbs, perennial, scapose, from tunicate bulb-rhizomes; leaves narrow, grasslike; inflorescences racemose, simple; racemes spikelike; flowers with tepals withering-persistent, 6, distinct, equal, leathery to slightly petaloid, yellowish green, green, or greenish purple; fruits capsular, 1–3-locular; seeds 1–8 per locule. 25 spp., one in South America, S. officinale (Schltdl. & Cham.) A. Gray ex Benth. from Mexico and Central America with disjunct populations in Venezuela (Aragua, Distrito Capital, Miranda, and Trujillo states); this species seems to originate from Mexico and was spread southward by native American Indians; however, little is known about the geographical distribution of the plant which is a prerequisite to undertake wild seed colection.

PHILESIACEAE

§ FULLY CHLOROTROPHIC

Genera/spp. 2/2 Distribution southern Chile and adjacent Argentina. Habit bisexual, suffrutex (Philesia) or liana (Lapageria); aerial roots absents; rhizome, when present, short, woody, sometimes branched. Use ornamental plants.

SYSTEMATIC both genera in South America.

1. Philesia Comm. ex. Juss. Sufflutex. Only one sp., P. magellanica J.F. Gmel, known only in Chile and Argentina.

2. Lapageria Ruiz & Pav. Fragile lianas, tubular flowers (white to red), succulent fruits. Only one sp., L. rosea Ruiz & Pav., endemic to Chile.

LINEAGE 3 of 3: ALSTOEMERIIDS

ALSTROEMERIACEAE

§ FULLY CHLOROTROPHIC

Genera/spp. 4/156 Distribution Neotropics to Tierra del Fuego, Australia, Tasmania, New Zealand. Habit bisexual, perennial (in Alstroemeria rarely annual) herbs, often with more or less lignified stem. Twining, climbing or epiphytic.

The family is distributed in New World from Central Mexico and Antilles to Patagonia, living in a wide range of habitats from cloud forests to swamps and desert areas, and from high Andes to marsh lands. Some species of Bomarea and several Alstroemeria species and hybrids have gained world-wide importance as cut flower crops due to their attractive flowers and the excellent keeping quality of them. Nevertheless, they are also satisfactory for growing in pots and for garden and landscape uses. The storage roots of some species of Bomarea and Alstroemeria are edible. The recognized taxa have been morphologically studied in detail and data focused on floral characters, the presence or absence of resupination in the leaf, the plant height and the characteristics of the inflorescence. Most of the floral characters have taxonomic importance. Several of them are also of ornamental weight such as the basic colour (which varies from yellow, orange through red-purple and violet to pink and white) the disposition, the form of the tepals and the presence or absence of spots or streaks on them.

SYSTEMATIC two tribes, both in South America.

1. TRIBE ALSTROEMERIEAE (2–3/210–240) ‣ both genera in South America.

1. Alstroemeria L. Herbs, mostly glabrous perennials with erect, foliate, sterile and fertile shoots; flowers showy in lax umbel-like, usually zygomorphic with patterned tepals; fleshy root tubers. 87 spp. from South America, from Peru (6) to Chile (35), Brazil (44, 39 endemics) up Argentina (10) and Bolivia, with two in Guiana Shield in Venezuela, both also Brazil.

In Brazil common in high mountains, sometimes abundant, near weeds, some also very narrow endemic, mainly in Minas Gerais state, at different habitats: forest, savanna, high fields, marsh, rocky grasslands (campos rupestres) and dry seasonal scrubland of NE Brazil (caatinga), from a height of 300 m in the Amazon rainforest up to 2,300 m at Serra do Itatiaia; 5 spp. from Pará, Mato Grosso, Espirito Santo and Minas Gerais are rare in Brazil, by Plantas Raras do Brasil’s book.

2. Bomarea Mirb. Stems twining, erect or procubent, leafy. Inflorescences pendulous or nodose. 114 spp., 10 exclusively from Mesoamerica (Mexico only 2), three from Central America to Ecuador, the last also in Caribbean, B. edulis (Tussac) Herb. in over range of genus, and unique species in Brazil, and 100 only in South America; four subgenera:

§ subg. Baccata ‣ 3 spp., B. diffracta Baker (Colombia), B. bracteolata Gereau (Panama) and B. carderi Mast. (Panama to Ecuador; large twining lianas, up to 10 m long, growing in lowland and montane forests.

§ subg. Bomarea ‣ 79 spp.; four sections are recognized within this subgenus: Multiflorae, Edules, Goniocaulon, and Pardinae; 16 spp. do not fit into any of these sections; all range of genus.

§ subg. Wichuraea ‣ 18 spp., distributed in the high Andes ranging between 2,500 m to 5,000 m; in the north of Argentina one species descends to 1,800 m; the most northern spe-cies is found in Ecuador, in the south the subgenus reaches the north of Argentina and Chile.

§ subg. Sphaerine ‣ erect or pendant, up to 1 m long, growing in cloud forests and páramo; 12 spp., Venezuela to Bolivia, high diversity in Peru (9).

2. TRIBE LUZURIAGEAE (2/5) ‣ outsider Drymophila (2; E & SE Australia, Tasmania).

3. Luzuriaga Ruiz & Pavon. Shrubs with a short rhizome, hemiepiphytic, scandent or pendant; leaves oblong-ovate; inflorescence axillary cincinnus. 4 spp., one in New Zealand and three in Atacama to Tierra del Fuego and Falklands, the latter also in Peru, in moss covered thunks and rockes, or in Empetrum heaths.

15. ASPARAGALES

FAMILIES ABSENTS IN SOUTH AMERICA: BLANDFORDIACEAE (1/4), BORYACEAE (2/13), DORYANTHACEAE (1/2), IXIOLIRIACEAE (1/4), LANARIACEAE (1/1), XERONEMATACEAE (1/2).

LINEAGE 1: ORCHIDACEAE

ORCHIDACEAE

§ MYCOHETEROTROPHICS (Arachnitis – TRIURIDACEAE – BURMANNIACEAE – THISMIACEAE – ORCHIDACEAE – Voyria - Voyriella - Monotropa)

Genera/spp. 704/22,000–23,000 Distribution cosmopolitan except polar regions, with the highest diversity in tropical and subtropical Asia and tropical Central and South America. Habit herbs or rarely vines, perennial, rarely annual, some fully strongly mycotrophic, epiphytic, terrestrial, lithophytic, or rarely aquatic or subterranean, usually green and photosynthetic, some without chlorophyll and holomycotrophs. Many spp. have rhizome, root tubers or tuberous stem. Numerous spp. (c. 70%) are epiphytes, often with internodes modified into swollen water-storing pseudobulbs, often with contractile aerial roots covered with a thick layer of dead water-absorbing tissue, velamen, formed from epidermis; other spp. are climbing. Some genera consist of achlorophyllous holoendoparasites on fungi (intracellular modified ectomycorrhiza with mostly basidiomycetes).

Several genera (especially within Vandeae, e.g. Campylocentrum) lack photosynthesizing leaves, which are replaced by photosynthesizing roots. The leaves of some epiphytic Epidendroideae-Vandeae may be very small and not photosynthetic and/or soon deciduous. The vegetative plant then consists largely of photosynthetic roots. These roots may be stout (ca 5 mm across) and terete, as in Dendrophylax, while the roots of the aptly named Taeniophyllum are distinctively flattened. There are over 200 species of leafless Epidendroideae, all epiphytes, with an estimated 20 or more independent losses of leaves. How carbon dioxide and water flux are controlled in leafless epiphytes is unclear, especially because there are no stomata in the roots, although the aeration units may be stomata analogues; roots of leafless orchids like Campylocentrum tyrridion Garay & Dunst., which lack stomata, also carry out CAM. Photosynthesis in orchid roots is poorly understood.

Orchids range vegetatively from Lilliputian plants a few millimeters long (Bulbophyllum and Platystele) to gigantic clusters weighing several hundred kilograms (Grammatophyllum) to some as much as 13.4 m in height (Sobralia altissima D. E. Bennett & Christenson from Peru); the world’s smallest orchid is a Campylocentrum insulare C. E. Siquiera & E.M. Pessoa, a rare aphyllous species restricted of a small area in S Brazil, possibly extinct in wild; plants of species reaches to 10 mm in maximous size. Nearly all temperate orchids are terrestrial. Likewise, flowers vary in size from less than 1 mm and barely visible to the naked eye (Platystele, Campylocentrum), to 15–20 cm diameter (some Paphiopedilum, Phragmipedium, and Cattleya), and ultimately to 76 cm (Phragmipedium caudatum (Lindley) Rolfe). Weight can vary from a fraction of a gram (many Pleurothallis) to nearly 100 grams (Coryanthes).

North American/Central America genera 60 genera in New World outside South America; united, these genera has 319 spp. in New World.

SYSTEMATICS five subfamilies, Apostasioideae (2/16, Sri Lanka, NE India, Japan, SE Asia, Malesia, NE Queensland, Melanesia.) does not occurs in South America.

1. SUBFAMILY VANILOIDEAE (c 15/180–190) - two tribes, both in South America.

1.1 VANILLOIDEAE ▸TRIBE POGONIEAE (5/75) - outsiders Cleistesiopsis (3; E and SE U.S.A.), Isotria (2; E and C U.S.A.), Pogonia (5; China (inc. Taiwan), Korean Peninsula, Japan; Moluccas Islands; E North America).

1. Cleistes Rich. ex Lindl. (exc. Cleistopsis) Terrestrial, upright, perennial herbs; stem upright, glabrous, terete, and hollow; leaves solitary and inserted halfway up the stem or several; inflorescence a single, terminal flower or a terminal raceme of 2-6 flowers in some tropical spp.; flowers showy, resupinate, and subtended by a large, leafy bract. 48 spp., all in South America, 4 up to Central America or Caribbean, 28 in Brazil, 20 endemics, growing in moist, welldrained, sandy, acidic soils; the spp. are also most frequently found in open savannas or at the edges of savannas and forests, collected at low elevations, but a few spp. are native to the high-elevation tepuis of E Venezuelan.

2. Duckeella Porto & Brade. Solid stems with basally linear leaves, lateral, ramified and multi-flowered inflorescences, yellow flowers, and a lip poorly differentiated from the petals. 4 spp., terrestrial from dense Amazon rain forest of N Brazil (3, none endemics), Colombia, Venezuela and possibly Guyana, mainly in open areas, on sandy soils of Amazonian white sand vegetation.

1.2 VANILLOIDEAE ▸TRIBE VANILLEAE (9/c. 180) - outsiders Lecanorchis (c 20; tropical Asia, Japan), Clematepistephium (1; New Caledonia), Eriaxis (1; New Caledonia); Erythrorchis (2; SE Asia, Malesia, Taiwan (China), the Ryukyu Islands; Queensland, New South Wales), Pseudovanilla (8; Malesia to tropical Australia and islands in the Pacific), Cyrtosia (5; India, S China, SE Asia, Japan, Ryukyu Islands), Galeola (6; India and Himalayas to China (inc. Taiwan) and SE Asia, Malesia to New Guinea, Madagascar and the Comoros).

3. Epistephium Kunth. Terrestrial, sympodial, erect herbs, stem elongate, robust, simple or branched; inflorescence a terminal raceme, rarely axillary; flowers showy, resupinate, typically light pink to dark purple in colour, short-lived. 29 spp. distributed throughout much of N South America and Bolivia, Argentina and Paraguay, two up to Caribbean or Central America; 100 – 1,200m. 13 spp. in Brazil, 6 endemics.

NE Brazil is a centre of diversity for the genus as a whole, but a number of spp. are also found in Colombia, Venezuela, the Guianas, and Paraguay. E. amabile Schltr. extends the range of the genus westward to Peru; the spp. are typically found growing in open, grassy areas, savannas or thickets. In some cases they are found in the unique white sand ecosystems of the Guiana Shield; some spp. (e.g. E. hernandii Garay) can grow to heights of 5 m or more and scramble through neighbouring vegetation. In contrast, E. ellipticum R.O. Williams & Summerh. is a diminutive spp. only 20 cm tall.

4. Vanilla Plum ex. Mill Epiphytic, hemiepiphytic or terrestrial, herbaceous vines; stems scandent, sparsely or profusely branched, terete or quadrangular, sometimes sulcate; inflorescence axillary, racemose to paniculate, rarely a complex cyme; flowers usually showy, short-lived, sometimes strongly fragrant; pseudobulb absent. 111 spp. pantropical in over tropical New World (90 rom Florida to Argentina and Caribbean), SE Asia-New Guinea (28) as in Africa-Madagascar (24). 76 spp. in South America, 40 in Brazil, 21 endemics, mainly in northern Amazon and Atlantic Forest; one sp. in Brazil occurs only in small marsh in SE Brazil, and V. palmarum (Salzm. ex Lindl.) Lindl. is found exclusively up in palms.

There are no Vanilla spp. in montane habitats, and those that have been collected at higher elevations usually have been found in savanna-like vegetation, dry forest on intermountain valleys, and occasionally in lower mountain rain forest; leafless spp. may be found in xeric formations, and at least Mascarene and Caribbean groups are found in tropical deciduous forest, thorn scrub, and coral buff vegetation; leafy spp. are usually found in more mesic environments, although some of them may reach tropical deciduous forest and coastal dune vegetation. spp. with membranaceous leaves are found in moister habitats. Therefore, aromatic fruits are known or expected from 35 spp., some of them cultivated locally or gathered from the wild, but they are not of economic importance. In Mexico, Indian women hold the fruits of V. pompona Schiede in great esteem and prepare them with oil before applying to the hair. The fruits of widely distributed V. odorata C. Presl are also collected in many American countries as a V. planifolia Andrews substitute and give fragrance to rums.

2. SUBFAMILY CYPRIPEDIOIDEAE (5/160–165) ▸ outsiders Cypripedium (51; temperate regions on the Northern Hemisphere), Paphiopedilum (80–85; India to S China and SE Asia, Malesia to New Guinea, Bismarck Archipelago and Solomon Islands), Mexipedium (1; Mexico).

5. Phragmipedium Rolfe. 32 spp. (and three hybrids) from SW Mexico to Bolivia and Brazil, eastwards French Guiana, absent in Caribbean; 29 spp. in South America, 7 in Brazil, two endemics.

6. Selenipedium Rchb.f. Herbs reaching some meters in height resembling in habit grasses or bamboos, rather than orchids; slipper orchids, with a sac-like labellum, unattractive and small flowers, huge size of the plants, and difficulty in their cultivation; they are often overlooked in the field while not in flower, as plants. 11 spp., disjunct: 4 spp. from Panamá to N Peru, four in Trinidad & Tobago to along E Venezuela and French Guiana up to Brazil (three of them, none endemics), and two endemics to Brazil, one in Amapá and N Pará state.

S. chironianum Sambin. & Braem., endemic to French Guiana, reported by Guy R. Chiron, Aurelien Sambin and Guido J. Braem. as Apedium Chiron, Sambin & Braem, new genus, with Apedium chironianum (Sambin & Braem) Chiron, Sambin & Braem their single species, not is recognized within.

3. SUBFAMILY ORCHIDIOIDEAE (194/4,925–5,000) - 4 tribes, only Diurideae (35/c. 1,000, Himalayas, China, Japan, tropical Asia, New Guinea, Australia, Tasmania, New Caledonia, New Zealand, Macquarie Island, Polynesia) absent in South America.

3.1 ORCHIDIOIDEAE ▸ TRIBE CODONORCHIDEAE (1/2) - a single genus.

7. Codonorchis Lindl. Perennial, terrestrial, sympodial herbs, reproducing from seed and also form local colonies by the production of daughter tubers on the end of stolons; the plants have distinct growth and dormancy phases, surviving extremes of cold as fleshy tubers; leaves 2-4, small; inflorescence erect, terminal, one-flowered; flower resupinate, petals free, as long or shorter than the dorsal sepal, erect. Two spp., C. canisioi Mansf. restricted S in Brazil, low mountains near the Atlantic coast, around São Leopoldo municipality in Rio Grande do Sul state (known only type collection), and C. lessonii (Brongn.) Lindl. in Argentina, Falkland Islands (Islas Malvinas), and Chile, occupying temperate to cold regions, along the Andes to Tierra del Fuego and Malvinas (Falkland) Islands; in this latter area, the habitats include the more humid areas in the sub-Antarctic rain forest.

3.2 ORCHIDIOIDEAE ▸ TRIBE ORCHIDEAE (61/c. 2,150) - 5 small lineages, Brownleeinae (2/86, Africa, Madagascar, Mascarene Islands, India, Sri Lanka, Thailand, China (inc. Taiwan), Japan, the Ryukyu Islands, Philippines, Java, the Lesser Sunda Islands, New Guinea, the Caroline Islands), Coryciinae (3/40, S Africa to Malawi and southern Tanzania, with their highest diversity in W Cape and the Drakensberg in KwaZulu-Natal), Pachites clade (1/2, W Cape) and Disinae (2/175, Africa, Madagascar, Mascarene Islands, Arabian Peninsula) do not occur in South America. Among Orchidinae (53/c. 1850), outsiders are all in Old World to Pacific (high diversity in South Africa, Madagascar and China) except Gymnadenia (23; Europe, temperate Asia, E North America), Coeloglossum (1; temperate regions on the Northern Hemisphere), Dactylorhiza (c 40; Europe, Macaronesia, Mediterranean, temperate Asia, Alaska), Pseudorchis (1–2; Europe, N Asia to Kamchatka, E North America, Greenland), Galearis (10; Himalayas, Tibet, Burma, China, Korean Peninsula, Japan, Russian Far East, Kamtchatka, Canada, U.S.A., W Greenland), Platanthera (c 135; temperate to tropical regions on the Northern Hemisphere south to mountains in Malesia).

8. Habenaria Willd. (inc. Habenella, Rhinorchis) Herbs perennial, terrestrial or semiaquatic, often rather succulent; leaves basal, abruptly reduced to bracts or scattered, gradually reduced toward inflorescence; inflorescences terminal; flowers few to many, resupinate, sometimes showy and nectar spur; petals 2-lobed, lateral lobe on abaxial margin; lip 3-lobed, spurred at base; fruits capsules, ellipsoid. 894 spp., pantropical. temperate and pantropical distribution (Canary Is., Africa, Madagascar, Middle East to Japan and Pacific, U.S.A. to over coutries South America, Caribbean), and its main centers of diversity are in Brazil, southern and central Africa, and E Asia. 325 spp. occurs in New World, 236 in South America, 170 in Brazil, 107 endemics, centered in savannas of C Brazil (cerrado); Minas Gerais, with about 81 species, has been the state with the highest number of recorded species.

3.3 ORCHIDIOIDEAE ▸ TRIBE CRANICHIDEAE (99/c. 1850) - eight small lineages, three absent in South America: Pterostylidinae (2/210, E Malesia, Queensland to South Australia, Tasmania, New Caledonia, New Zealand), Galeottiellinae (1/2, Mexico, Guatemala) and Manniellinae (1/2, tropical W Africa).

∎ SUBTRIBE CHLORAEINAE ‣ all genera in South America.

9. Bipinnula Comm. ex Juss. Terrestrial herbs; leaves 2–6, green with light-green veins, erect or prostrate; inflorescence terminal, 1-flowered or exceptionally 2-flowered, predominantly greenish, with reticulate green veining and labellum differently coloured with respect to other perianth segments; labellum clawed or sessile, dark-coloured, articulated basally, entire to trilobed, ecalcarate; capsule erect, ellipsoid, dehiscent via two dorsal sutures. 10 spp., Argentina, Chile, Uruguay and S Brazil (4, none endemics).

10. Chloraea Lindl. Roots thick, cylindrical, with evident absorptive hairs; leaves several in a basal rosette; inflorescence few-to many-flowered, slender to robust, acute, reticulate, sterile bracts, similar to the leaves; floral bracts elliptic-lanceolate, acute, much longer than the flower at the base of the inflorescence, then as long as the ovary; flowers white, greenish, yellow; orange, or reddish, sessile. 53 spp., 51 in Andes from and S Argentina (inc. Tierra del Fuego), Peru, Bolivia, Chile, and the Falkland Islands, except by and C. bella Hauman endemic to Entre Rios, Argentina and neighbouring areas of Uruguay, and C. membranacea Lindl. in SE South America up Paraná state in S Brazil.

11. Gavilea Poepp. 17 spp., Juan Fernandes, S Chile to Falkland Is in Argentina from Missiones to Tierra del Fuego.

∎ SUBTRIBE GOODYERINAE ‣ outsiders Aenhenrya (1; S India), Anoectochilus (43; tropical Asia to Hawaii), Chamaegastrodia (3; Himalayas, E Asia), Cheirostylis (53; tropical and subtropical regions of the Old World), Cystorchis (c 20; Thailand and eastwards to Micronesia), Danhatchia (1; New Zealand), Dossinia (1; Borneo), Erythrodes (26; tropical Asia, New Caledonia to Samoa and Tonga), Eurycentrum (7; New Guinea, Solomon Islands, Vanuatu), Gonatostylis (2; New Caledonia), Goodyera (100–105; temperate regions on the N Hemisphere, Mozambique, Madagascar, tropical Asia), Halleorchis (1; Cameroon, Gabon), Herpysma (1; S and SE Asia to Sumatra), Hetaeria (c 30; Sri Lanka, SE Asia, Malesia to Tahiti), Hylophila (7; Thailand to Solomon Islands), Kuhlhasseltia (10; Korean Peninsula, Japan to New Guinea), Lepidogyne (1; Malesia), Ludisia (1; SE Asia, W Malesia), Macodes (11; Japan, Vietnam, Malesia to Vanuatu), Myrmechis (17; Himalayas, Tibet, SE Asia, Malesia to New Guinea, Taiwan (China), Korean Islands, Japan, the Kuril Islands), Odontochilus (c 25; Himalayas, tropical Asia, Hawaii), Orchipedum (3; SE Asia, Java, Philippines), Pachyplectron (2; New Caledonia), Papuaea (1; New Guinea), Platylepis (17; tropical and S Africa, Madagascar, Mascarene Islands, Seychelles, Moluccas, Tahiti), Rhomboda (22; Himalayas, S China (inc. Taiwan), SE Asia, Malesia to New Guinea, Bismarck Archipelago, Solomon Islands, New Caledonia, Queensland, Japan, the Ryukyu Islands), Schuitemania (1; Philippines), Vrydagzynea (43; SE China (inc. Taiwan), SE Asia, Nicobar Islands, Malesia to New Guinea, Solomon Islands, Vanuatu, Queensland, Fiji, Samoa, Tonga), Zeuxine (c 75; tropical and subtropical regions in the Old World), Zeuxinella (1; Vietnam).

12. Microchilus C. Presl. (inc. Aspidogyne, Kreodanthus, Ligeophila, Platythelys, Stephanothelys) Terrestrial or occasionally lithophytic, rarely epiphytic herbs; stem with few to many, scattered to subrosulate leaves; leaves coloured green to brownish, sometimes with silver to greenish maculation; inflorescence pubescent; flowers resupinate, the outer surface of sepals glabrous to pubescent. 246 spp. from Mexico to Bolivia, eastwards French Guiana, Argentina, Paraguay, Brazil (40, 18 endemics), the Caribbean, and Galapagos Islands, in lowland to mid-elevation evergreen forest from sea level to 2,000m; 160-224 spp. in South America. One species, M. pedicellatus (Cogn.) E.C.Smidt & M.W.Chase from Brazil and Paraguay, has leaves reduced to bracts and may be mycoheterotrophic.

∎ SUBTRIBE SPIRANTHINAE ‣ outsiders Funkiella (27; Texas, Mexico, Guatemala, Costa Rica), Sotoa (1; W Texas, N and C Mexico), Svenkoeltzia (4; Mexico), Aulosepalum (8; Mexico to Costa Rica), Physogyne (3; Mexico), Pseudogoodyera (1; Central America, Cuba), Greenwoodiella (3; Texas, Mexico, Central America, Caribbean), Kionophyton (4; Mexico, Guatemala), Schiedeella (22; SW U.S.A., Mexico, Central America, Caribbean), Dichromanthus (4; Texas, Arizona, Mexico, Central America), Deiregyne (18; Mexico, Guatemala).

13. Aracamunia Carnevali & I.Ramirez. Terrestrial herbs; rhizome creeping, slender; roots fleshy, cylindrical, pubescent; stems short but conspicuous, to 15 mm long; leaves 7-11, forming a loose rosette; inflorescence a loose raceme with 3-6 flowers; flowers white, ascending, non-resupinate, narrowly tubular. Only one sp., A. liesneri Carnevali & I. Ramírez, endemic to Pantepui Life Zone, known so far only from the summit of the Cerro Aracamuni in Venezuela, growing on mosses of stream banks in closed forest at 1,550m.

14. Beloglottis Schltr. 7 spp., Florida Mexico, Belize to Panamá, Lesser Antilles, Suriname, Venezuela, Colombia to Bolivia, C Brazil (only the widely B. costaricensis (Rchb. f.) Schltr., in Amazonas state) and W Argentina; 3 spp. in South America.

15. Brachystele Schltr. Terrestrial, acaulescent herbs; leaves one or several arranged into a loose rosette; sometimes some 'cauline' leaves, or foliaceous bracts, are present on the lower part of the inflorescence; inflorescence inconspicuously many-flowered, all-sided to more or less onesided raceme; flowers held horizontally, more or less tubular, fragrant in some spp., white, yellowish or greenish, sometimes sepals and petals green and labellum white to orange-yellow. 19 spp. distributed in Mexico, Nicaragua, Costa Rica, Colombia, Venezuela, Suriname, Guyana, French Guiana, Peru, Chile, Paraguay, Uruguay, Brazil (10, 3 endemics), and Argentina, absent in Ecuador; terrestrial, thriving in open habitats including dry to marshy grasslands, savanna, rocky fields, xerophilous scrub, tropical deciduous forest, dry pinelands, and grassy openings in pine-oak forest, usually in clay, sandy or rocky ground and sometimes in areas subject to periodic fires, from sea level to 2,300m. 17 spp. in South America.

16. Buchtienia Schltr. 4 spp., one sp. in Peru to Bolivia, N to C Brazil and Paraguay, and Brazil, Peru and Ecuador one endemic each.

17. Coccineorchis Schltr. 8 spp., Mexico to Colombia, one up to Bolivia; 5 spp. in South America.

18. Cotylolabium Garay. Terrestrial, acaulescent herbs; roots fleshy, flexuose; leaves few (3), cauline, distantly spaced; inflorescence almost completely covered by the leaf sheaths, above with tubular, acute, spaced bracts, apex condensed, one-sided; flowers held horizontally, yellow. Only one sp., C. lutzii Garay, known only from Serra do Caparaó, from 2,600 to 2,800m above sea level, in shallow soil among high altitud grasslands (campos de altitude), occupying an area less than 200 m² and consisting of few tens of individuals, on the bordering area between the states of Minas Gerais and Espírito Santo, in SE Brazil.

19. Cybebus Garay. Only one sp., C. grandis Garay, Colombia and Ecuador.

20. Cyclopogon C Presl. (exc. Quechua, inc. Stigmatosema) Terrestrial, lithophytic or epiphytic, acaulescent herbs; leaves one or several forming a basal rosette, present or less commonly absent at flowering time; Inflorescence with flowers on all-sides to more or less onesided, few to many; flowers slightly descending or nodding, less commonly held horizontally, fleshy, fragrant; sepals and petals green, olive green, greyish green, bronzy green, greenish white, or creamy, sometimes suffused with red, brown or purple, labellum white, often with green, brown or reddish marks. 91 spp. SE U.S.A. (Florida), Mexico, Central America, the Caribbean, and South America (79, excluding Chile); predominantly terrestrial or lithophytic, but some spp., such as C. prasophyllus (Rchb.f) Schltr. and C. epiphyticus (Dodson) Dodson, are usually, perhaps exclusively, epiphytic.

Cyclopogon spp. thrive in varied environmental conditions, including moist to wet tropical and subtropical, cloud, and pine-oak forests, ravines, and riparian vegetation in tropical deciduous and semi-deciduous forests, woody and shrub by savannas, grasslands, rocky fields, marshy areas, and bogs from sea level to about 3,000m. Brazil has 38 spp., 23 endemics.

21. Degranvillea Determann. Terrestrial, rhizomatous, leafless herbs; roots apparendy absent; inflorescence white, pubescente; flowers opening simultaneously, held horizontally, more or less tubular, white. Only one sp., D. dermaptera Determann, seems to be an obligate mycoparasite; it has been found growing in humus soil in seasonally dry tropical forest between 400 and 650m, in French Guiana and Suriname.

22. Eltroplectris Raf. 14 spp. from South America (Suriname, Venezuela, Colombia to Bolivia, over Brazil (9, 6 endemics), Argentina and Paraguay), one up to North America.

23. Espinhassoa Salazar & J.A.N. Bat. Rosette of several iridescent, elliptic, shortly petiolate leaves, flowers with a narrowly tubular base, flared perianth segment apices, comparatively simple labellum, and short pollinarium with the viscidium located at the center of its ventral surface. Two spp., both endemics to mountains of Minas Gerais state, SE Brazil.

24. Eurystyles Wawra. Obligate epiphytic, acaulescent herbs, growing on bare or moss-covered branches, tree trunks, and vines up to a few metres above the ground; leaves several, forming minute, often grouped rosettes, do not wither at the end of the growing season; blades green, often greyish, glossy; inflorescence erect to pendulous; capitate with numerous minute flowers; floral bracts larger than the flowers, imbricating, ovate; flowers tubular, non-resupinate. 23 spp. distributed from SE Mexico throughout Central America, the Greater Antilles, and South America (17, except in Chile and Uruguay and Guianas). 11 spp. in Brazil, 8 endemics.

25. Hapalorchis Schltr. Terrestrial or epiphytic, rhizomatous herbs; leaves one or (more commonly) several, forming a loose rosette; inflorescence several-flowered, one-sided; flowers held horizontally, loosely tubular; coloration described as white or yellowish green, often with sepals pale green sometimes suffused with pink, petals white with a green or brownish green vein near the apex, and labellum white with green veins. 13 spp. from South America, two up to Mexico and Central America, also Caribbean; 6 spp. in Brazil, 5 endemics; delicate terrestrials or humus epiphytes in shady, wet locations in tropical and sub-tropical evergreen, cloud, and riparian forests and marshy areas, from near sea level to about 2,350m; flowering occurs throughout the year.

26. Helonoma Garay. 4 spp., one in Colombia to Peru, three from Venezuela, Guyana and Amazonas state in N Brazil (1, none endemic).

27. Lankesterella Ames. Epiphytic, acaulescent herbs, epiphytes growing with mosses and lichens on trunks and branches of small trees and shrubs; leaves several, forrning minute, often grouped rosettes; inflorescence ebracteate, densely pilose (the hairs translucent); flowers held horizontally to descending, tubular, large in proportion to the size of the plant; petals white, translucent with darker veins, labellum white with green veins on the interior. 11 spp. distributed in Cuba, Hispaniola, Costa Rica, Panamá, Colombia, Venezuela, Ecuador, Peru, Bolivia, Brazil (7, 6 endemics), Paraguay, and Argentina – 8 in South America, from near sea level to 2,700m.

28. Lyroglossa Schltr. Two spp. in Mexico and Central America, one up to to Trinidad, Brazil, French Guiana to Venezuela, Colombia.

29. Mesadenella Pabst. & Garay. Terrestrial or lithophytic, usually in leaf mould or humus, acaulescent herbs; leaves several, forming a basal rosette; Inflorescence glabrous below, sparsely pubescent above, flowers on all sides; Flowers ascending, scentless, white or greenish white, often with brownish suffusion near the base of sepals and a yellow area on labellum. 10 spp., distributed from Mexico through Central America to Colombia, Venezuela, Suriname, French Guiana, Guyana, Peru, Ecuador, Bolivia, Brazil (4, two endemics), Paraguay, and Argentina; 9 in South America, in the shade in tropical and subtropical, evergreen or semi-deciduous forests, which are wet to seasonally dry, and often in seasonally inundated areas or riparian vegetation, from near the sea level to about 800m.

30. Microthelys Garay. 5 spp., 4 from New Mexico to C. America, and M. intagana (Dodson & Dressler) Szlach. endemic to Ecuador.

31. Nothostele Garay. Terrestrial, caespitose herbs, acaulescent, roots tuberous; leaves absent at anthesis; inflorescence terminal, elongate, secund, several-flowered. Two spp., both found on calcareous banks in a canyon at 1,400m, from savannah in center Brazil, one in Minas Gerais, another in Distrito Federal and Goiás state.

32. Odontorrhynchus M.N.Correa. 8 spp., one in Peru and 7 from Bolivia to Chile and Argentina.

33. Pelexia Poit. Ex. Rich. (inc. Pachygenium) Terrestrial, lithophytic or rarely epiphytic, sometimes humus epiphytes, acaulescent herbs; leaves several, usually forming a basal rosette, rarely cauline; inflorescence glabrous below; flowers ascending to held nearly horizontally, sepals and petals green, yellow or greenish white, rarely reddish brown. 91 spp., Mexico, Central America, the Antilles, and over South America (80, except in Chile); they live in a variety of habitats, including tropical and subtropical evergreen and semi-deciduous, cloud, oak, and tropical deciduous forests, wet to seasonally dry grassland and scrub, marshy areas, and rocky fields from sea level to about 3,100m. Brazil has 39 spp., 29 endemics.

34. Pteroglossa Schltr. (inc. Ochyrella, Callistanthos) 22 spp. from South America (Venezuela, Colombia, Ecuador, Peru, Brazil, Argentina and Paraguay), one up to Mexico; 11 spp. in Brazil, 7 endemics.

35. Quechua Salazar & L.Jost. (off Cyclopogon) Only one sp., Q. glabrescens (T.Hashim.) Salazar & L.Jost, Andean Ecuador and Peru, growing on steep rocky slopes and canyons of limestone in foothill and mountain rain forest, from 1,100–1,700m elevation.

36. Sacoila Raf. Terrestrial, acaulescent herbs; leaves several, forming a basal rosette; Inflorescence glabrous below, pubescent above, many-flowered, all-sided; Flowers relatively showy, fleshy, odourless, pale green, yellowish tan, pink, brick red or coral red. 8 spp. from South America, two up to SE U.S.A. (Florida) through Mexico, Central America and the Antilles; 5 spp. in Brazil, two endemics; in general, Sacoila inhabits areas of grassland, dry scrub, rocky fields, savannas, and oak, pine-oak, and tropical deciduous to semi-evergreen forests from sea level to 2,200m. In tropical Mexico and Central America, S. lanceolata is common in abandoned fields, cattle pastures, roadsides, and gardens, and they have been observed even in the middle of paths and dirt roads.

37. Sarcoglottis C. Presl. Terrestrial, acaulescent herbs; leaves several, forming a basal rosette; inflorescence many-flowered, rarely only 1-2 flowers or the flowers densely arranged in a subcorymbose short raceme; flowers held horizontally or slightly nodding to strongly ascending, fleshy, green, yellow, white, rosy, bronze, brownish or often a combination of these colours, commonly with dark green veins in labellum and sometimes petals. 49 spp. found from Mexico throughout all of Central America, Trinidad and the Lesser Antilles to South America (35, except in Chile); exclusively terrestrial, found in a variety of habitats that include tropical and sub-tropical evergreen and semi-deciduous, tropical deciduous, cloud, oak and oak-coniferous forests, dry or wet, marshy grasslands, savannas, xerophilous scrub, and rocky fields from sea level to about 2,700m; 24 spp. in Brazil, 15 endemics.

38. Sauroglossum Lindl. Terrestrial, acaulescent herbs; leaves several, forming a basal rosette; inflorescence glabrous below, glandular-pubescent above, many-flowered, either all-sided or sub-secund; flowers erect to held nearly horizontally, yellow, orange, brick red, or the sepals and petals green, yellowish green or whitish, sometimes with a reddish tinge, and the labellum white or yellow. 12 spp. restricted to South America, in two major areas: one with 10 spp. includes the Andes of Colombia, Ecuador, Peru, Bolivia, Chile, Argentina and W Brazil (two in this portion of country), and the other with two species the region of the Serra do Mar in SE Brazil; this plants grow in leaf litter in dense to open, moist to relatively dry and riparian forests, grassland, sclerophyllous scrub, and open grassy, rocky or gravelly areas with shallow soil from near sea level to about 3,700m.

39. Skeptrostachys Garay. Terrestrial herbs; leaves several, forming a basal rosette or, more commonly, distributed along the inflorescence; inflorescence many-flowered, all-sided; floral bracts green, red or brownish; flowers showily coloured, fleshy, slightly ascending, white, greenish yellow, pale yellow, fleshcoloured, pale brown-orange, brick red or bright red. 14 spp. distributed in the Brazilian Shield (12, 5 endemics) and adjacent areas of Paraguay, Uruguay, and Argentina, with one record in Suriname; all spp. grow in open areas, including moist or wet grasslands, marshy places, rocky grasslands (campos rupestres), and savannas of C Brazil (cerrado), often in sandy soil and areas subject to periodic fires from sea level to about 1,600m.

40. Spiranthes Rich. 35 spp. from temperate regions on the Northern Hemisphere, Malesia, Australia, Pacific islands, tropical America, 26 spp. in New World, with S. torta (Thunb.) Garay & H.R. Sweet widely distributed in North America, Central America and Caribbean, with records in French Guiana.

41. Stalkya Garay. Only one sp., S. muscicola (Garay & Dunst.) Garay, endemic to Andes of W Venezuela.

42. Stenorrhynchos Rich. ex Spreng. 8 spp. Mexico to South America (6, Venezuela to Bolivia).

43. Thelyschista Garay. Terrestrial, acaulescent herbs; roots numerous, fleshy, cylindrical, pilose; leaves 4-7, forming a basal rosette; inflorescence pubescent, many-flowered on all sides; flowers tubular, sepals pale green, petals and labellum white. Only one sp., T. ghillanyi (Pabst) Garay, in rocky fields, growing in sandy places on conglomerate and sandstone substrate, at 700 to 850m, in Diamantina Range, center Bahia state, Brazil.

44. Veyretia Szlach. 11 spp. Trinidad, French Guiana to Colombia, over Brazil (9, 5 endemics), Argentina and Paraguay.

∎ SUBTRIBE DISCYPHINAE ‣ a single genus.

45. Discyphus Schltr. Acaulescent, deciduous geophytes with fasciculate roots; a single orbicular, cordate leaf lying on the substrate and clasping the base of the scape; inflorescence densely glandular-pubescent; flowers campanulate, resupinate, petals free from the dorsal sepal but adnate to the proximal half of the column, labellum free. Only one sp., D. scopulariae (Rchb.f.) Schltr., highly disjunct, in Panamá, Trinidad, Venezuela and NE Brazil.

∎ SUBTRIBE CRANICHIDINAE ‣ outsiders Fuertesiella (1; Cuba, Hispaniola), Galeoglossum (3; Mexico).

46. Aa Rchb. f. Terrestrial, caespitose herbs; leaves basal, rosulate; inflorescence lateral, erect, elongate, many-flowered, narrowly racemose; flowers small; petals narrower than sepals, glabrous, free. 26 spp. from Andes, from Venezuela to Argentina, one sp. up to Chile, also Costa Rica, open scrub and woodland, damp mountain meadows, paramo, among scrub and grass between sandstone or limestone rocks, peaty areas between rocks, peat swamps, and river gravel; 3,100-4,400m.

47. Altensteinia Kunth. 8 spp., Venezuela to Bolivia.

48. Baskervilla Lindl. Terrestrial herbs with a short ascending rhizome and fleshy horizontal hairy roots; leaves ovate to lanceolate, petiolate; inflorescence elongate, many-flowered; peduncle remotely sheathed; bracts glabrous; sepals free, spreading; petals with a claw adnate to the column and a spreading or reflexed free blade. 10 spp., 9 native to Nicaragua, Costa Rica, Panamá, and the Andes (Guyana to Bolivia), with one sp. in S & SE Brazil and Bolivia (B. paranaensis (Kraenzl.) Schltr.); terrestrial in cloud and elfin forest, less commonly in secondary forest; 1,000-3,300m.

49. Cranichis Sw. Terrestrial, caespitose herbs; leaves one-several, usually basal in a rosette, rarely cauline, fleshy, ovate to lanceolate, often longly petiolate, present at anthesis; inflorescence terminal, elongate; peduncle and rachis puberulent; bracts leafy, shorter than the flowers; sepals free; petals free, much narrower than the sepals, often ciliolate on the margins, sub-spreading. 73 spp. native from Florida, Mexico and Caribbean south to Bolivia and Argentina, the greatest diversity in the Andes from Venezuela to Peru, but only 5 in Brazil (3 endemics); 55 in South America. Terrestrial or lithophytic in primary and secondary lowland and montane woodland and forests, in humus and on sphagnum tussocks, rarely in meadows; 360-3,000m.

50. Gomphichis Lindl. 36 spp. from Venezuela and Guyana to Bolivia, two up to Central America.

51. Myrosmodes Rchb.f. Small terrestrial herbs with fleshy, fusiform, often pubescent, fasciculate roots; leaves fasciculate, small, narrowly elliptic to ovate-elliptic, shortly petiolate, often fleshy; inflorescence lateral, erect, many-flowered, racemose, anthesis often before leaves emerge; peduncle completely enclosed by infundibuliform, scarious, often ustulate, at first imbricating then spreading sheaths; bracts scarious, often ustulate, shorter than flowers; flowers non-resupinate, small, glabrous. 16 spp. native to Andean South America from Venezuela to Bolivia and Argentina; plants are commonly seen among tussocks of Azorella Lam., swampy ground near lakes and streams, in stream beds, and rocky areas; 3,800-4,600m, the highest habitat known to support orchids in Earth (4,900m in Andes)

52. Ponthieva R. Br. (inc Exalaria) Terrestrial, caespitose herbs; leaves several, ovate to lanceolate, petiolate; inflorescence terminal, elongate; flowers delicate in texture; sepals thin-textured; petals adnate in basal part to sides of column, asymmetrical. Labellum fleshy, small, concave, entire to three-Iobed, adnate in basal part to the column. 74 spp. widely distributed in the tropical and subtropical Americas from Florida and Texas to northern Argentina, east up to French Guiana, Caribbean; 58 in South America; plants grow on mossy trunks and on the forest floor in cloud forest, along streams in dry forest, and moist banks in oak and oak/ pine woodland from sea level to 3,000m; only three spp. in Brazil, one endemic.

53. Porphyrostachys Rchb.f. Two spp., Ecuador to Peru.

54. Prescottia Lindl. Erect, terrestrial herbs. Rhizome absent. Stems simple, erect, pubescent distally; leaves non-articulate, basal, petioles sheathing at base, lamina elliptic to ovate; inflorescences terminal, racemes many-flowered; flowers non-resupinate; sepals thin, basally connate forming a short cup, rarely free, erect; petals thin, narrow, adnate to the sepaline cup. 23 spp. of Florida, the Caribbean, Mexico to Argentina, Uruguay, 22 in South America, mainly in Brazil (13, 9 endemics); found from the subtropical regions. Prescottia is more widely distributed and may be found from sea level to 3,000m; most plants occur in the shade of shrubs, broadleaf forests or pine woodlands in soil or leaf-litter.

55. Pseudocentrum Lindl. 11 spp., 11 from Colombia to Bolivia, two in Caribbean and one in Central America.

56. Pterichis Lindl. 37 spp., one in Jamaica and 36 in South America (Venezuela to Cono Sur), three up to Central America.

57. Solenocentrum Schltr. 4 spp., two in Central America, one in Colombia and Ecuador, and one in Bolivia.

58. Stenoptera C. Presl. Terrestrial herbs with fleshy, fibrous, fasciculate roots; leaves basal or cauline; inflorescence terminal, erect, many-flowered; flowers non-resupinate, small; sepals subequal, basally connate into a rather slender tube, wide opening, apices spreading; petals free, not connivent with dorsal sepal, often linear, spreading. 7 spp. native to Colombia to Peru, one sp. in N Brazil; found on open rocky slopes and open woodland in paramo and cloud forests; 800-3,600m.

4. SUBFAMILY EPIDENDROIDEAE (590–600/c 18,000) ▸ 16 tribes; Thaieae (1; Thailand), Arethuseae (23/c 760; S Canada, E U.S.A., Caribbean, India, S China, SE Asia, Malesia to New Guinea, Australia and islands in W Pacific, Japan), Podochileae (27/c. 1,300; tropical Africa, India, Himalayas, S China, SE Asia, Malesia to New Guinea, Australia, Melanesia and islands in W Pacific) and Nervilieae (3/84, Europe, temperate Asia to Himalayas and Kamchatka, tropical and S Africa, Madagascar, Arabian Peninsula, tropical Asia to New Guinea and N Australia, islands in the Indian and Pacific oceans) do not occur in South America.

EPIDENDOID LINEAGE 1 of 4: NEOTTTIEAE

4.1 EPIDENDROIDEAE ▸ TRIBE NEOTTIEAE (6/178) - outsiders Cephalanthera (19; temperate regions on the Northern Hemisphere), Neottia (c 65; Europe, temperate Asia, North America), Aphyllorchis (22; Sri Lanka to Japan and E Queensland), Limodorum (3; Mediterranean to Iran), Epipactis (70–75; Europe, tropical Africa, temperate Asia, Himalayas, SE Asia).

59. Palmorchis Barb. Rodr. Terrestrial herbs; roots fasciculate, slender. Stem erect, reed-like, occasionally shrubby, leafy above; leaves plicate, elliptic to ovate-elliptic, acute to acuminate, petiolate, spreading, usually few in number; inflorescence axillary or terminal, racemose or paniculate, several-flowered; floral bracts conduplicate; flowers small for the size of the plant, resupinate; sepals and petals subsimilar, free, connivent to spreading; petals shorter and narrower than sepals. 39 spp. tropical regions of Central and South America (French Guiana to Peru, N Brazil), including Trinidad, with the centre of diversity in northern South America (35, 7 in Brazil, three endemics), in shade in mixed lowland forests on lateritic- and ironstone-derivative soils as well as in riverine, cloud, and ‘crabwood’ forests, from sea level to about 1,000m.

EPIDENDOID LINEAGE 2 of 4: SOBRALIEAE

4.2 EPIDENDROIDEAE ▸ TRIBE SOBRALIEAE (4/269) - all genera in South America.

60. Brasolia (Rchb.f.) Baranow, Dudek & Szlach. (off Sobralia) Tall erect plants with reed-like, unbranched stems; leaves narrowly lanceolate to ovate, distichous, often somewhat plicate; nflorescence axillary from the base of one of uppermost leaves or terminal, few- to many-flowered, often branched, with distinct internodes of rachis and its branches. 23 spp., Guianas to Bolivia and N Brazil (two species, no endemics).

61. Elleanthus C. Presl. Epiphytic or terrestrial, caespitose herbs; roots fleshy. Stems simple or branched; leaves distichous, plicate or occasionally conduplicate; inflorescence terminal, racemose or subcapitate, spiral or distichous, the floral bracts often conspicuous in size or colouring; flowers concave, usually brightly coloured; sepals and petals free, similar. 130 spp., collectively range from Mexico and Caribbean to Bolivia and S Brazil (14, none endemics), eastwards to French Guiana, especially diverse in the Andes (108 in South America). Elleanthus are epiphytes or lithophytes in wet or cloud forests from close to sea level up to about 3,000m.

62. Sertifera Lindl. ex Rchb.f. 11 spp., NW Venezuela to Peru.

63. Sobralia Ruiz. & Pavon. (exc. Brasolia) Terrestrial, lithophytic, or epiphytic herbs; roots fleshy; stem unbranched or rarely branched, from a few centimetres to 14m; leaves cauline with basal sheaths, plicate; inflorescence terminal or axillary, racemose, often condensed; flowers resupinate; sepals and petals free, similar. 128 spp. in mainland tropical America (81 in South America) from Mexico to Peru, Bolivia, and S Brazil (21, 8 endemics), up to French Guiana. S. altissima D.E. Benn. & Christenson, endemic to Amazonian Peru, is tallest of all orchids, reaching up to 13.4m tall.

EPIDENDOID LINEAGE 3 of 4: HOLOMYCOTROPHICS VICINITY

4.3 EPIDENDROIDEAE ▸ TRIBE GASTRODIEAE (6/91) - outsiders Auxopus (4; tropical Africa, Madagascar), Didymoplexiella (8; SE Asia, W Malesia to Japan and the Ryukyu Islands), Didymoplexiopsis (1; Thailand, Laos, Vietnam, Hainan), Didymoplexis (17; tropical and SE Africa, Madagascar, Himalayas, SE Asia, China (inc. Taiwan), Malesia to New Guinea, northern Australia, Melanesia, and islands in western Pacific), Gastrodia (c 65; tropical Africa, E Asia to Japan, the Ryukyu Islands and Taiwan (China), Himalayas and SE Asia to New Guinea, Melanesia, Australia, Tasmania and New Zealand).

64. Uleiorchis Hoehne. Terrestrial, holomycotrophics, achlorophyllous herbs. Stem erect, glabrous; leaves scale-like, membranous, sessile; inflorescence erect, 1–6-flowered, glabrous; Flowers resupinate, patent, glabrous, sepals and petals pale brownish white, sometimes also faintly tinged with pale violet, labellum dull yellow with brown or sepia markings near base. 4 spp. known from Central America and tropical South America. U. ulei (Cogn.) Handro, the more widely distributed, has been collected in Honduras, Panamá, Colombia, Venezuela, Peru, the Guianas, and SE Brazil (absent in Ecuador); U. liesneri Carnevali & I.Ramírez is only known from a single collection from Amazonian Venezuela; U. pratënsis M.E.Engels & E.C.Smidt is known from few collections in the Atlantic Rain Forest of Serra da Prata, S Brazil, growing from 30 to 400m; and U. longipedicellata A.Cardoso & Ilk.-Borg is only known from tree sites in the Carajás Range, Pará state.

Plants occur in sheltered locations in humus-rich substrates, such as leaf mould or forest hummocks; less commonly, they are found growing on moss-covered stumps or decaying logs; plants are usually solitary or sometimes occur in small groups; according to a note by Ule attached to one of his herbarium collections at Brussels, some populations may persist at the same location for many years.

4.4 EPIDENDROIDEAE ▸ TRIBE TRIPHOREAE (4/30) - all genera in South America.

65. Monophyllorchis Schltr. Single-leaf herbs. 5 spp. from Nicaragua to Ecuador, two of then endemic to Colombia and one endemic to Ecuador.

66. Pogoniopsis Rchb.f. Terrestrial, achlorophyllous, mycoheterotrophic herbs; stem thick, fleshy, pale yellow in colour, erect; leaves reduced to scale-like bracts, brown; inflorescence a congested raceme of a few flowers; flowers subtended by large, yellow bracts; sepals yellow, oblong-lanceolate, 11-12 mm long; petals white, similar to the sepals in shape and size; labellum lanceolate, equal in size to the sepals, white with orange striations; it´s only genus of mycoheterotrophic endemic to Brazil. Two spp., P. nidus-avis Rchb.f. from Minas Gerais and the area around Rio de Janeiro, and P. schenckii Cogn. from Pernambuco de Santa Catarina states; collections have been made at elevations around 1,100m.

67. Psilochilus Barb. Rodr. Terrestrial herb with a creeping rhizome; stem erect, purple; leaves sheathing stem, petiolate, upper surface light or dark green sometimes with light green or white markings, lower surface purple or purplegreen; inflorescence terminal, racemose or paniculate; flowers erect; sepals free, keeled; petals free, not keeled. 15 spp. found throughout the Greater and Lesser Antilles, Central America, and tropical South America (11) in Guyana to Peru and Brazil (two spp., none endemics). All spp. are terrestrial, herbaceous plants of wet montane forests up to 1,600m and rare or uncommon throughout their range.

68. Triphora Nutt. Terrestrial herbs; leaves greenish purple, alternate, often reduced and bract-like, ovate to cordate, convolute or plicate, the margins entire or slightly serrate. Stems one or two, relatively thick, succulent; inflorescences racemose to corymbose, lateral or terminal; flower resupinate or non-resupinate. 21 spp. distributed in the eastern U.S.A. and southern Ontario (Canada), Caribbean, and through Mesoamerica to Ecuador, east to French Guiana, Brazil (8, 6 endemics); 21 in South America (absent in Peru). All members of the genus are terrestrial, understorey herbaceous plants of mesic or montane forests, found at elevations ranging from sea level to 3,000m.

They grow under generally low-light conditions in moist, well-drained, rich soils among leaf litter. Because of their small size and inconspicuous habit, field botanists often overlook them. As a consequence of periodic dormancy with fluctuations in the number of individuals observed from season to season, Triphora populations leave an impression of being elusive or having an annual life cycle; the factors that cause Triphora to enter and emerge from dormancy are not known, but at least in the temperate regions of its distribution such factors may be related to the amount of rainfall; flowering generally occurs in late summer, which may (at least in the temperate regions) help to increase the chances of outcrossing because few other flowering plants are blooming at that time, and therefore competition is relatively low for pollinators.

4.5 EPIDENDROIDEAE ▸ TRIBE XERORCHIDEAE (1/2) - a single genus.

69. Xerorchis Schltr. Terrestrial herbs; roots fine, wiry; leaves distichous, delicate and thin-textured, gradually becoming smaller toward the apex of the stem; inflorescence terminal; flowers not opening widely, white to greenish yellow. Two spp., Colombia, Brazil, the Guyanas, Venezuela, Ecuador, and Peru, in forest-floor terrestrials, growing in humus or on fallen trees in deep shade in dense lowland and lower montane forests, from sea level to 700m.

4.6 EPIDENDROIDEAE ▸ TRIBE TROPIDIEAE (3/39) - outsider Kalimantanorchis (1; W Borneo)

70. Corymborkis Thouars. Terrestrial, rhizomatous herbs; stem reedlike, usually unbranched, to several metres tall; leaves several; inflorescence an axillary raceme or panicle, much shorter than the leaves, fewto many-flowered; flowers sometimes showy and fragrant; sepals and petals approximately subequal, connivent or connate at base. 6-7 spp., pantropical, 2 spp. occur from Guinea to South Africa, the Comoro Islands, Madagascar, and Mascarenes; 1-2 spp. range from tropical Asia to NE Australia and the South Pacific islands (to Samoa); 5 spp., all in South America, are distributed in the Mexico, Caribbean, Central America and northern South America, with one sp. also in Bolivia, Paraguay, Brazil (C. flava (SW.) Kuntze, unique in Brazil, no endemic), and Argentina; all spp. are grow in shaded localities in evergreen tropical forests.

71. Tropidia Lindl. Terricolous herbs, green or holomycotrophic; with erect stem, often branched; two or many plicate leaves; inflorescences axillary or terminal, few- to manyflowered; flowers resupinate or non-resupinate, sometimes with a spur; and column with prominent rostellum, and a dorsal and erect anther with two elongated sectile pollinia. 20 spp., tropical & subtropical Asia to SW Pacific, and T. polystachia (Swartz) Ames in Florida to tropical America, from Mexico to Ecuador, Venezuela, Caribbean, and a single and recent contacted population in W Mato Grosso state, in center Brazil.

4.7 EPIDENDROIDEAE ▸ TRIBE WULLSCHLAEGELIEAEE (1/2) - a single genus.

72. Wullschlaegelia Rchb. f. Leafless, terrestrial herbs; roots fusiform (arising from the stem base) or filiform; stem ascending; inflorescence elongate, subtended by scale-like bracts, pubescent; flowers white, pubescent, resupinate or not, subtended by bracts; dorsal sepal and petals similar in shape, elliptic to ovate to broadly lanceolate, acute; petals falcate-basally expanded, rounded. Two spp. from Caribbean and Mexico/Central America through tropical South America (except Bolivia and Chile), both in Brazil. It is uncertain which fungal groups are associated with Wullschlaegelia.

EPIDENDROID LINEAGE 4 of 4: CORE EPIDENDROID

4.8 EPIDENDROIDEAE ▸ TRIBE MALAXIDEAE (17/c. 4,800) - three lineages, all in South America.

∎ SUBTRIBE VARGASIELLINAE (1/4) ▸ a single genus in South America.

73. Vargasiella C.Schweinf. 4 spp., S. Venezuela, Colombia, Peru and Bolivia, absent in Ecuador.

∎ SUBTRIBE MALAXIINAE (14/c. 1,400) ▸ outsiders Alatiliparis (5; Sumatra, Java), Crepidium (c 260; islands in the Indian Ocean, India, China, SE Asia, Malesia to New Guinea, northern Australia, islands in the Pacific), Crossoliparis (1; Central America), Dienia (2; India, S China, SE Asia, Malesia to northern Australia and Melanesia, Japan), Hammarbya (1; temperate and subarctic regions on the Northern Hemisphere), Hippeophyllum (c 10; Malesia), Oberonia (c 325; tropical and S Africa, Madagascar, Mascarene Islands, India, SE Asia, Malesia to New Guinea and northern Australia), Oberonioides (2; China, Thailand), Orestias (4; tropical Africa), Stichorkis (8; islands in the Indian Ocean, India, SE Asia, Malesia to New Guinea, islands in the Pacific), Tamayorkis (4; Mexico, Central America).

74. Crossoglossa Dressler & Dodson. 43 spp., from Nicaragua southward into Colombia and Bolivia along the Andes, 33 in South America, in extremely humid forests, at elevations of (500-)1,500-2,500(-2,850) m where they can be epiphytic or terrestrial; in South America, the majority of the species occurs in Colombia and Ecuador, fewer species occur in Peru and Bolivia, and one is known from Venezuela.

75. Liparis Rich. Terrestrial in fens, marshes, wet grassland, and among rocks, usually in light to deep shade, or epiphytic on tree trunks and branches in shade in lowland, hill, lower, and upper montane foress, occasionally on rocks, from sea level to 3,600m; rhizomatous herbs, rarely mycotrophic and leaves reduced to scales; stem pseudobulbous, clustered or not, covered when young by sterile bracts; leaves one to several; inflorescence erect, racemose; flower yellow, green, orange, or purple, often translucent, usually resupinate. 424 spp., mainly in Asia to SW Pacific islands, subtropical and tropical New World (45, 32 in South America, only three in Brazil, one endemic), except Chile and Uruguay, with a single spp. in Europe and two in North America. L. alpina P.Royen was recorded on wet, moss-covered slopes in alpine grasslands between 3,350 and 3,600m.

76. Malaxis Sol. ex. Sw. Terrestrial or rarely epiphytic, occasionally holomycotrophic herbs; roots hairy. Stem cylindrical to pseudobulbous, fleshy, often creeping and rooting in basal part; leaves, if present, thin-textured to fleshy, usually pleated, petiole sheathing at base; inflorescence erect, racemose, unbranched; Flowers green, brown, yellow, pink, or purple. 164 spp., found throughout the tropics and subtropics of the Old and New World - 144, Mexico to Argentina, Paraguay, Brazil (12, 8 endemics), eastwards French Guiana, Caribbean, with a few in temperate regions of Europe, Asia; 55 spp. in South America; species are terrestrial in marshes, grassland, woods, and forests, usually in shade, or wet paludiculous areas (e.g. M. cipoensis fron C Brazil) from sea level to 3,000m.

∎ SUBTRIBE DENDROBIINAE (3/3376) ▸ outsider Genyorchis (10; tropical W and C Africa); Dendrobium Sw is the largest flowering genus absent in New World, 1586 spp. ranges from Sri Lanka throughout tropical Asia and the Pacific region, north to Japan, east to Tahiti, and south to New Zealand.

77. Bulbophyllum Thouars. Epiphytic, occasionally lithophytic, or rarely terrestrial herbs; leaves usually persistent, sometimes deciduous, duplicate, inarticulate, usually petiolate, thinly herbaceous to coriaceous, usually glabrous; inflorescences one- to many-flowered, apical or along distal part of rhizome. 2,168 spp. (5^th largest worldwide) and is widely distributed from continental tropical Africa, the Comoros, Madagascar, Seychelles, Reunion and Mauritius (c. 200), India to New Zealand and the tropical Pacific islands as far east as Tahiti (c. 1,600) to the Neotropics (81, 77 in South America); the main centres of diversity are Madagascar (200) and New Guinea (600); 55 spp. in Brazil, 41 endemics. Six sections among Neotropical species based on floral characters of species:

§ sect. Bulbophyllaria ▸ three spp., B. aristatum (Rchb.f.) Hemsl., the largest range for any spp. of Bulbophylum in New World, from Cuba, Costa Rica, Dominican Republic, El Salvador, Guatemala, Honduras, Mexico (also with 1 endemic), Nicaragua, Panamá, and Venezuela; a third from over northern South America.

§ sect. Didactyle ▸ 7 spp., occurring in Paraguay (only one sp., also Brazil), Brazil (3 endemic), Colombia (1 endemic), one in Brazil reaching into Guyana and Venezuela, another to Venezuela and Peru across Colombia and Ecuador.

§ sect. Furvescens ▸ 5 spp., Mexico to N Bolivia, east to French Guiana, 2 over northern South America, ramaining in Mexico (1 endemic), Venezuela (1 endemic) and Suriname and French Guiana (1 endemic).

§ sect. Micranthae ▸ 12 spp., 11 occurring in C & SE Brazil (8 endemic), Bolivia (3, 1 endemic) and Paraguay (1, also in Brazil).

§ sect. Napelli ▸ 12 spp., one endemic Venezuela, remaining in Brazil, one of then reaching into Argentina.

§ sect. Xiphizusa ▸ 25 spp., Mexico to Venezuela and Ecuador, Jamaica, and S Peru to Bahia, southwards to NE Argentina; 15 spp. endemics to Brazil.

4.9 EPIDENDROIDEAE ▸ TRIBE CYMBIDIEAE (154/c. 3,800) ▸ 10 subtribes, nine in South America (three small, three medium, three larges), Cymbidiinae (11/c. 130, tropical and S Africa, Madagascar, SE Asia to Solomon Islands, Japan and Australia) absent.

∎ SUBTRIBE CYRTOPODIINAE (1/47) ‣ a single genus.

78. Cyrtopodium R. Br. Mostly terrestrial or lithophytic, rarely epiphytic, caespitose herbs; pseudobulb when young entirely concealed by distichous, scarious sheaths, eventually naked; leaves several, coriaceous; inflorescence lateral, arising from near base of pseudobulb, erect, racemose to paniculate, many-flowered; flowers resupinate, often showy; sepals and petals membranous, spreading, often conspicuously undulate. 54 spp. is distributed from Florida (North America), Caribbean and Mexico through all of Central America to S Brazil (39, 23 endemics) and Argentina but is well represented in central Brazil; 53 spp. in South America. Most spp. are terrestrial or lithophytic, and these are rarely found established on trees; relatively few spp. are epiphytic; the terrestrial spp. occupy a broad range of habitats, including open grasslands, savannas of C Brazil (cerrado), dry rocky slopes, and seasonally-to-permanently wet areas; plants in the genus have a marked seasonal growth; pseudobulbs of terrestrial spp. can be entirely above ground, but those of many others are partially or completely underground.

∎ SUBTRIBE EULOPHIINAE (3/288) ‣ outsiders Grammangis (2; Madagascar), Orthochilus (c 35; tropical Africa, Madagascar, tropical and subtropical America).

79. Eulophia R.Br. 245 spp. Expanded genera, mainly Old World, only three spp. in New World, from U.S.A. to Panamá and Caribbean, French Guiana to Venezuela, Colombia to Bolivia, over Brazil (only the wider E. alta (L.) Fawc. & Rendle), Argentina, Uruguay and Paraguay.

∎ SUBTRIBE CATASETINAE (8/c. 354) ‣ all genera in South America.

80. Catasetum Rich. ex Kunth. Epiphytic (often on rotten tree trunks), lithophytic, or sometimes terrestrial herbs. Pseudobulb of several internodes, slenderovoid or fusiform, when young entirely concealed by distichous, scarious sheaths, eventually naked; leaves several, distichous, convolute, plicate, articulate, deciduous at end of growing season; inflorescences one to several, lateral, arising from base of pseudobulb, erect, arching or pendent, racemose; flowers unisexual, often sexually dimorphic, rarely also intermediate; floral bracts lanceolate, funnel-shaped.

192 spp. and several documented natural hybrids is distributed from Mexico to S Brazil and Argentina, Paraguay, up to French Guiana, and in Caribbean (Cuba and Trinidad) but is best represented in Brazil (123, 93 endemics); 190 in South America, 9 in Caribbean; most spp. of Catasetum are found on trees or palms, but some grow on sand (e.g. C. discolor) or on thin organic soils on granite (e.g. C. bergoldianum Foldats) or sandstone outcrops (e.g. C. discolor). Their range extends from sea level (e.g. C. macrocarpum Rich. ex Kunth along the coast of Brazil, the Guianas, and Venezuela) up to 2,200m (e.g. C. discolor, growing on tepui summits in the Guiana Shield). Natural hybridization is relatively common in the genus. Epiphytic spp. are often found in rotten wood, and it is apparent that some plants first grow mycotrophically for a period of time, developing an extensive root system, and then produce pseudobulbs that can increase in size rapidly within a few years. Plants can bear either male or female flowers, usually in separate inflorescences. Male (staminate) flowers tend to be colourful, resupinate or not, with a saccate or open labellum, and they forcibly attach a relatively large pollinarium onto their pollinators. Female (pistillate) flowers, on the other hand, are always green and non-resupinate (Romero 1990); inflorescences with both male and female flowers are rarely found in nature; it is not known why they are encountered more frequently in cultivated plants. Intermediate flowers, ranging variously from the typical morphology of male and female flowers, are rarer in nature, and even bilaterally half male-half female flowers have been documented; these intermediate flowers are, however, reproductively non-functional.

81. Clowesia Lindl. Epiphytic herbs on trees or palms in tropical rain forests at 60–1,700m. Pseudobulb of several internodes; ovoid to slender-ovoid, when young entirely concealed by distichous, scarious sheaths, eventually naked; leaves several, distichous, convolute, plicate, articulate; deciduous at end of growing season; inflorescence one to several, lateral, arising from base of pseudobulb, arching or pendent, racemose; flowers resupinate, bisexual, protandrous, fragrant, often showy; bracts lanceolate, funnel-shaped; sepals and petals membranous. 7 spp., six distributed from Mexico to Panamá, two up to South America, C. warczewitzii (Lindl. & Paxton) Dodson up to Brazil, and one endemic to Ecuador, but it is particularly well represented in Mexico, where at least five spp. have been reported (4 endemics).

82. Cyanaeorchis Barb.Rodr. Three spp. from S Brazil, one up to NE. Argentina.

83. Cycnoches Lindl. Epiphytic or rarely terrestrial herbs; leaves several, distichous, convolute, plicate, articulate, deciduous at end of growing season; inflorescences one to several, lateral, arising from base of apical internodes, arching or pendent; flowers non-resupinate, unisexual, staminate and pistillate ones often sexually dimorphic, rarely also with polymorphic intermediate ones; bracts lanceolate, funnel-shaped. Staminate flowers whitish green or various colours, spotted or not; sepals and petals membranous, lateral sepals strongly refl exed or not. 26 spp. is distributed from Mexico to S Brazil (4, one endemic) and northern Argentina, east up to French Guiana, but it is well represented in Mesoamerica and the northern Andes; 22 spp. in South America.

84. Dressleria Dodson. 12 spp., Nicaragua to Peru, absent in Venezuela, 8 in South America.

85. Galeandra Lindl. Caespitose epiphytic or terrestrial herbs; leaves several, distichous, coriaceous; inflorescence terminal, erect or arching to pendent, racemose or paniculate, with 1–20 flowers; flowers resupinate, often showy; bracts concave to cymbiform, linear to narrowly ovate, acuminate, not as long as ovary; sepals and petals membranous, erect to spreading; sepals narrowly elliptic to narrowly obovate, lateral sepals sometimes oblique; petals similar to sepals, oblique. 31 spp. from Florida, Caribbean and Mexico to S Brazil and Argentina; 26 spp. in South America; Brazil is the country with the largest number of spp., 17 spp. in two main areas, but only four endemics: equatorial Amazon region (mainly epiphytic, mostly on palms (Arecaceae) in gallery forests and permanently flooded places) and the South American savannas (mainly terrestrial, at umid grasslands and sandy areas). G. montana Barb.Rodr. occurs in sandy soils in montane habitats on the Brazilian Shield and in sand dunes near the sea in Bahia and Espírito Santo. Among terrestrials, G. beyrichii Rchb. f. grows in thick rain forests and seasonal forests; many populations of this spp. appear to be holomycotrophics and flower over many years without producing leaves.

86. Grobya Lindl. Epiphytic, sympodial herbs; leaves 3–7, linear, with 1–3 prominent veins on abaxial surface, base sheathing; inflorescence lateral, raceme, few to many-flowered; peduncle with numerous bracts, basal ones tubular, apical ones laminar; flowers resupinate, generally yellow to pale yellow with brownish dots; sepals and petals membranaceous. 5 spp., epiphytes in wet forests of E Brazil, mainly in the Brazilian Atlantic Forest; G. cipoensis F. Barros & Lourenco is an epiphyte exclusively on Vellozia gigantea, which grows on rocky grasslands (campos rupestres).

87. Mormodes Lindl. Epiphytic, rarely terrestrial, saprolignophilous herbs; leaves several; inflorescences 1–several, lateral, erect, Flowers showy, fragrant, perfect but always dichogamous, resupinate or non-resupinate. 73 spp. that are widely distributed in tropical America, from Mexico to Bolivia and Brazil (27, 20 endemics) east to French Guiana, but excluding the Caribbean; 75 spp. in South America. All spp. of Mormodes, as for most other members of subtribe Catasetinae, are saprolignophilous, that is, they live on the rotting wood of stumps and fallen trees or branches, as well as other substrates that provide them with a continuous supply of nutrients and moisture, as in the humus trap formed by the leaf bases of some palms; these plants are mycotrophic, and seedlings are always found associated with fungal hyphae that spread on and into the rotting wood. The mycoheterotrophic phase of at least some of the spp. seems to be relatively long; large masses of achlorophyllous rhizomes and roots can apparently live for years embedded in the wood before an aerial shoot is visible, which then usually grows rapidly and is able to produce flowers and seeds within only a couple of years; Mormodes occur in evergreen and semi-deciduous tropical forests and cloud forests from sea level to about 2,200m elevation.

∎ SUBTRIBE ONCIDIINAE (64/c. 1,610) ‣ outsiders Amparoa (1; S Mexico, Central America), Cuitlauzina (7; Mexico, Central America), Hintonella (1; Mexico).

88. Aspasia Lindl. Up to 30 cm tall, epiphytic, caespitose, sympodial herbs; leaves bifacial, conduplicate, articulate, eventually deciduous, epetiolate; inflorescences one or two, lateral, glabrous, never branched, shorter than leaves, 1–6-flowered, bracts shorter than pedicel; flowers resupinate, spurless, cream to pinkish or brownish with darker spots; sepals and petals free, similar in size and shape, but sepals immaculate and petals usually with spots or bars of dark brown. 7 spp. known from Guatemala and Belize south to Panamá in Central America and from Venezuela and Colombia south to Brazil (4, one endemic) and Paraguay, growing epiphytically at lower elevations (800m or less), often in seasonally dry forests.

89. Brassia R. Br. (inc. Mesospinidium) Moderate to large (15–70cm tall), epiphytic, caespitose, sympodial herbs; leaves bifacial, conduplicate, articulate, eventually deciduous, epetiolate; inflorescences one or two, lateral, glabrous, rarely branched, often longer than leaves, 5–30-flowered, bracts usually shorter than pedicels, rarely covering part of flower bases; flowers resupinate, spurless, cream, yellow, orange-yellow or greenish yellow; sepals and petals free, similar in shape but petals often shorter than sepals, often both ‘spidery’ and elongate-curving, usually with dark brown to reddish brown spots concentrated on their basal halves. 67 spp. from Florida (North America) through the Caribbean, Mexico, and all of Central America through South America to Brazil (14, 4 endemics) and Bolivia, east up to French Guiana, growing epiphytically at a wide range of elevations, from sea level in seasonally dry forests up to 2,500m or more in cloud forests. 60 spp. in South America.

90. Caluera Dodson & Determann. 4 spp., scattered in Venezuela, Ecuador, French Guiana, N Brazil (two spp., both in Pará state, one endemic) and Suriname.

91. Capanemia Barb.Rodr. Small (less than 8 cm tall), epiphytic always on small branches, caespitose, sympodial herbs. Pseudobulb, when present, abbreviated, with a single terminal leaf and 1–3 leaf-bearing bracts. Leaf conduplicate or terete, articulate, eventually deciduous, epetiolate; inflorescences one or two, lateral, glabrous, rarely branched, shorter than leaves, 1–2-flowered, bracts shorter than pedicels; flowers resupinate, spurless, white to cream with a bright yellow spot in the centre of the labellum; sepals and petals free or lateral sepals fused up to 3/4 their length, similar in size, shape, and colour. 6 spp. in Bolivia, Argentina, Paraguay, Brazil (5, 4 endemics), and Uruguay but centred in SE Brazil, growing epiphytically at low elevations (ca. 300m) in seasonally dry forests; they occur on introduced spp. of genera such as Citrus L. (Rutaceae) and Hibiscus L. (Malvaceae), as well as in primary forests.

92. Caucaea Schltr. 20 spp., Venezuela to Peru.

93. Chytroglossa Rchb.f. Small (up to 3 cm), epiphytic, caespitose, sympodial herbs; leaves duplicate, bifacial, articulate, eventually deciduous, epetiolate; inflorescences 1–3, lateral, glabrous, unbranched, usually longer than leaves, 1–8-flowered, bracts nearly as long as pedicels; flowers usually resupinate, spurless, yellow-green to yellow, often with reddish purple spots on labellum, lateral lobes white or yellow with reddish purple spots. Three spp., restricted to the Atlantic Forest of E Brazil in the states of Paraná, São Paulo, Rio de Janeiro, Espírito Santo, and Minas Gerais, (ca. 600–1,800m) in wet forests.

94. Cischweinfia Dressler & N.H.Williams. 12 spp., Costa Rica to Bolivia, absent in Venezuela; 10 spp. in South America.

95. Comparettia Poepp. & Endl. (inc. Neokoehleria, Scelochilus) Small to moderate (up to 20 cm), epiphytic, caespitose, sometimes partially psygmoid, sympodial herbs. Pseudobulb elongate to orbicular; leaves 1–4, mostly bifacial, conduplicate, but some unifacial, articulated, sometimes darkly (red) pigmented; inflorescences one or two, lateral, glabrous, mostly unbranched, usually exceed- ing length of leaves, 5–25-flowered, bracts shorter than pedicels; flowers resupinate, red, orange, yellow, pink, carmine, greenish yellow, often with red and white markings on labellum; sepals and petals free, except for lateral sepals that are at least basally fused to form a nectar spur, the others similar in size, shape, and colour, lanceolate. 84 spp. collectively occur from Caribbean through Mexico (only two) south through Central America to Brazil (5, none endemics), Bolivia, and Peru; 83 spp. in South America. Only one species, C. falcata, is widely distributed. Most species grow on the smallest branches of their host trees in wet montane forests up to 2,700m, but C. falcata and others can grow as low as 100m. In certain areas, they move onto cultivated plants, and most occur in sites in which there are often mosses and lichens.

96. Cuitlauzina Lex. 8 spp. Mexico to Colombia, two spp. in South America.

97. Cypholoron Dodson & Dressler. Two spp., Venezuela and Ecuador one endemic each.

98. Cyrtochiloides N.H.Williams & M.W.Chase. Three spp., SE. Mexico to Peru, one endemic to Central America.

99. Cyrtochilum Kunth. (inc. Trigonochilum) Large (up to 2 m), epiphytic to terrestrial, caespitose to longscandent, sympodial herbs; pseudobulb ovoid, usually round in cross-section, with 2–4 terminal leaves and 2–6 leaf-bearing bracts; leaves conduplicate, bifacial, articulate, eventually deciduous, epetiolate; inflorescences one or two, lateral, glabrous, usually branched and often thigmatropic (actively vining) or lax, longer than leaves, 10–60-flowered, bracts shorter than pedicels; flowers resupinate, spurless, white, yellow, pink, brown or purple, often in unusual mixtures of colours with brown or pinkish spots or bars; sepals and petals free, similar in size and shape or petals much wider, reflexed in some spp. 187 spp. ranging in the high Andes of Venezuela, Colombia, Ecuador, Bolivia, and Peru, with one sp. also occurring outside of this area on Guadeloupe, Hispaniola, and Puerto Rico (the Caribbean); these spp. grow in wet forests throughout the Andes at middle to high elevations (1,200m and up); many of them grow on or near the ground, and their long, vining inflorescences climb up through the vegetation to reach brighter areas. Some of the smaller spp. grow on small branches or twigs (e.g. C. meirax (Rchb.f.) Dalström) and approach being twig epiphytes; there are a number of spp. that grow in high-elevation grasslands or páramos.

100. Eloyella P.Ortiz. (inc. Dunstevillea) 10 spp., from Colombia to Peru, one of them up to Panamá, Venezuela and Guyana, French Guiana, one in Roraima and Pará states in N Brazil.

101. Erycina Lindl. Small (2–10 cm), epiphytic, caespitose, psygmoid monopodial or sympodial herbs; leaves 2–9; inflorescences 1–5, lateral, glabrous, unbranched or branched, usually longer than leaves, sometimes successively flowering, 1–25-flowered, bracts shorter than pedicels; flower orientation irregular but mostly resupinate, unspurred, uniformly bright yellow; sepals and petals free or lateral sepals fused basally, otherwise similar in size and shape or petals much wider. 7 spp. occurs widely throughout the American tropics, up to Brazil (3, all wider), absent only from the Antilles and Florida, although it does occur on Trinidad; six spp. in South America in wet, lower elevation forests from 200–600m on smaller axes of their host trees, except for E. hyalinobulbon, these spp. grow in. E. hyalinobulbon (La Llave & Lex. ) N.H.Williams & M.W.Chase, which grows in oak-pine forests at 1,200–2,000m on the Pacific slopes of Mexico; it is usually found on trunks and other larger axes.

102. Fernandezia Ruiz & Pav. 94 spp., 89 only South America (Venezuela to Bolivia), two of this continent to Central America, and three only Central America and Mexico.

103. Gomesa R.Br. (inc. Alatiglossum, Baptistonia, Coppensia) Small to large (up to 70 cm tall), epiphytic or terrestrial, sympodial herbs, usually caespitose but in some cases with an elongated rhizome or erect stem; leaves three or four; inflorescences one or two, lateral, glabrous, unbranched to branched, usually exceeding length of leaves, 3–60-flowered; flowers usually resupinate, spurless, yellow, green or white, often with brown markings. 124 spp. with a centre of distribution in Brazil (120, 106 endemics), particularly in the Atlantic Forest, but extending to Argentina, Bolivia, Paraguay and Uruguay, with two outliers in Colombia and Venezuela and one in both areas; most are epiphytes, but several grow terrestrially in swampy areas in full sun (e. g. G. montana Barb.Rodr.). Most grow at moderate elevations, rarely at sea level, and some come from cloud forest conditions.

104. Grandiphyllum Docha Neto. Small to large (up to 50 cm tall), epiphytic (rarely lithophytic), sympodial, herbs; leaves one per sympodium, often greygreen, coriaceous, dorsiventrally flattened, bifacial, articulate, epetiolate; inflorescence lateral, usually branched, usually longer than leaves, 4–80-flowered, bracts shorter than pedicels; flowers resupinate, spurless, pale yellow, white, brown, pale cream, often with brown to reddish brown spots; sepals and petals free, oblanceolate, similar in size and shape or petals in some spp. larger and broader. 9 spp. endemics to SE & S Brazil except three found into N Argentina (Corrientes and Misiones Provinces) and Paraguay; in spite of their coriaceous leaves, Grandiphyllum grows mostly in relatively shady sites from sea level to 1,000m, also on trunks and other larger axes.

105. Hofmeisterella Rchb.f. Two spp., Venezuela to Bolivia.

106. Ionopsis Kunth. 5 spp., two very widely in tropical America (both in Brazil), plus Colombia, Brazil and Ecuador one endemic each.

107. Leochilus Knowles & Westc. 11 spp., S Florida, Mexico, Belize to Panamá, Caribbean, Venezuela to Ecuador and Brazil (1, wider); only three in South America.

108. Lockhartia Hook. Small to moderate (10–35 cm), epiphytic, caespitose, sympodial (appearing at times monopodial) herbs. Stem erect to pendulous, covered in short leaves (pseudobulbs present only in L. genegeorgei D.E.Benn. & Christenson, elongate with conduplicate, articulate leaves); leaves 5–50, unifacial, non-articulate (except in L. genegeorgei), overlapping, imbricate; inflorescences 1–25, lateral or terminal, glabrous, sometimes branched, usually not exceeding leaves, 1–4-flowered, bracts shorter than pedicels, open, orbicular; flowers greenish yellow, yellow or white with darker red or red-brown markings; sepals and petals free, similar in size and shape, obovate. 32 spp., which occur throughout much of the Neotropics, from Mexico through Central America to Panamá and from Venezuela and French Guiana to Bolivia, Brazil (8, 3 endemics), Peru and Trinidad; 28 spp. in South America, found over a wide range of elevations from sea level to 2,000m in relatively shaded sites with high humidity. They flower repeatedly from the same sympodia over a period of 2–3 years.

109. Macradenia R.Br. 13 spp. from South America (French Guiana to Venezuela, Colombia to Peru, over Brazil, Paraguay), two up to Florida and Mexico; 11 spp. in Brazil, 6 endemics.

110. Macroclinium Barb.Rodr. 41 spp., Mexico, Belize to Panamá, French Guiana to Venezuela, Ecuador to Bolivia, N to SE Brazil (8, 6 endemics); 25 in South America; absent in Colombia.

111. Miltonia Lindl. (inc. Phymatochilum) Moderate to large (25–85 cm tall without the infl orescence), epiphytic, mostly caespitose but sometimes long-rhizomatous (e.g. M. spectabilis), sympodial herbs. Pseudobulb laterally compressed, ancipitous; leaves 2–6, bifacial, conduplicate, articulate; inflorescences 1–3, lateral, glabrous, unbranched to branched, arching-erect, usually longer than leaves, 1–150-flowered, bracts shorter than pedicels; flowers resupinate, cream, yellow, orange, purple, pink or brown with darker pink to reddish brown or white markings; sepals and petals free, similar in size and shape or petals broader, oblanceolate to lanceolate and spidery. 12 spp. endemics to Brazil except one up to Argentina and Paraguay, at lower elevations (200–1,000m) in relatively shaded sites with high humidity; with this circumscription (excluding Miltoniopsis), Miltonia does not occur in Central America, Bolivia, Colombia, Ecuador or Peru.

112. Miltoniopsis God.-Leb. 6 spp. from Venezuela to Bolivia, two up to Central America.

113. Nohawilliamsia M.W.Chase & Whitten. Small to medium-sized (up to 25 cm tall) epiphytic, sympodial herbs with obovoid, longitudinally furrowed pseudobulbs subtended by 1–3 foliaceous bracts; leaves one or two apically plus 1–3 basally per sympodium; inflorescence one or two, lateral, unbranched to sparsely branched, erect, much longer than leaves, 5–35- flowered; flowers resupinate, spurless, sepals and petals reddish purple with a yellow margin, labellum bright yellow with a few reddish purple spots on the callus; sepals and petals free, oblanceolate, similar in size and shape. Only one sp., N. pirarensis (Rchb. f.) M.W. Chase & Whitten, known only from southern Venezuela (Amazonas and Bolivar), W Guyana and NW Brazil in Roraima state, an unusual distribution for a member of Oncidiinae; occurs more or less terrestrially or lithophytically in the mountains (tepuis) of the Guiana Shield and the Amazon basin at about 300–1,600m.

114. Notylia Lindl. Small (5–20 cm tall), epiphytic, caespitose, sympodial herbs; leaves 2–8; inflorescences 1–3, lateral, glabrous, unbranched, pendent, usually longer than leaves, 10–70-flowered, bracts shorter than pedicels; flowers resupinate, cream, white, yellow or green with rose and yellow markings in some spp.; labellum often white; sepals and petals lanceolate, free except lateral sepals fused in many spp., otherwise similar in size. 50 spp. found from Mexico south through Central America to Panamá, and from Colombia and Venezuela to Brazil (25, 10 endemics), Bolivia, Paraguay, and Peru; 44 spp. in South America, from nearly sea level to about 900m in relatively shaded sites with high humidity; they are twig epiphytes and occur on the smallest axes of their hosts.

115. Notyliopsis P.Ortiz. Only one sp., N. beatricis P. Ortiz, endemic to Colombia.

116. Oliveriana Rchb.f. 14 spp., 13 from Guyana to Peru, and one endemic to Bolivia.

117. Oncidium Sw. (inc. Heteranthocidium, Odontoglossum) Small to large (5–120 cm) epiphytic, mostly caespitose, sympodial herbs; leaves 2–4, bifacial, articulate; inflorescences 1–3, lateral, glabrous, branched or unbranched, erect to pendent, usually longer than leaves, 2–120-flowered, bracts shorter than pedicels, some spp; flowering repeatedly from the same inflorescence (e. g. some spp. of the former genus Sigmatostalix); flowers resupinate, cream, white, yellow or pink with reddish brown, rose and yellow markings in some spp. 435 spp. found from Mexico and Florida (North America) through Central America and the Caribbean to Bolivia, and Peru; 372 spp. in South America, 14 in Brazil (only one endemic); there are two centres of diversity, Mexico and the Andes, from sea level to 2,500; some occur in exposed sites, whereas others grow in highly shaded places; nearly all are epiphytic, but some spp. grows in soil (e.g. O. graminifolium Lindl. in heavy clay soils in Mexico).

118. Ornithocephalus Hook. Small (up to 10 cm) epiphytic, monopodial, sometimes elongate, psygmoid herbs without pseudobulbs; leaves 4–30, unifacial, laterally flattened, usually articulate, eventually deciduous, epetiolate; inflorescences 1–5, lateral, sometimes hairy, unbranched, usually shorter than leaves, 8–50-flowered, bracts shorter than or same length as pedicels; flower orientation variable, usually resupinate, spurless, white, yellow-green to yellow, often with orange-brown spot(s) on labellum; sepals and petals free, petals usually wider and falcate with a minutely erose or papillate margin, sometimes with a papillate reverse. 56 spp., collectively ranging from Mexico through Central America and the Caribbean to Brazil (7, two endemics), Bolivia, and Peru, east up to Caribbean; 40 spp. in South America; some spp., such as O. gladiatus, occur over most of this distribution; Ornithocephalus grow epiphytically at low to middle elevations, ca. sea level to 2,000m in wet forests; flowering has been reported throughout the year.

119. Otoglossum (Schltr.) Garay & Dunst. (inc. Brevolongium, Ecuadorella) 29 spp., Costa Rica, Panamá, Venezuela, Colombia to Bolivia, Amazonas state in N Brazil (only one species); 28 spp. in South America, centered in Ecuador.

120. Phymatidium Lindl. Small (up to 10 cm tall), epiphytic, caespitose, monopodial herbs without pseudobulbs; leaves 5–30, spirally arranged, bilaterally flattened, unifacial, terete to triquetrous, unarticulate, epetiolate, except in P. falcifolium Lindl. with bifacial leaves; inflorescences 1–5, lateral, unbranched, longer than leaves, 1–10-flowered, bracts shorter than pedicels; flowers mostly resupinate but variable, spurless, green, greenish yellow or white; sepals and petals free, obovate, similar in size and shape. 10 spp. found in SE Brazil; one sp. edges into Uruguay and another into Argentina and Paraguay, growing epiphytically at 150–1,850m in humid forests.

121. Platyrhiza Barb.Rodr. Small (up to 8 cm tall), epiphytic, caespitose, sympodial herbs; leaves 3–5, distichous, conduplicate, bifacial, articulate, epetiolate; inflorescences 1–3, lateral, unbranched, longer than leaves, 1–10-flowered, bracts shorter than pedicels; flowers mostly resupinate but variable, spurless, greenish yellow to greenish white with a yellow spot in middle of the labellum callus; sepals and petals free, oblanceolate, similar in size and shape. Only one spp., P. quadricolor Barb. Rodr., found only in SE Brazil in the wet Atlantic Forest, growing epiphytically in shady sites at 150–1,000m in humid forests just a few metres off the ground.

122. Plectrophora H.Focke. 8 spp., from French Guiana to Venezuela, Colombia to Bolivia, N to C Brazil (5, two endemics), one up to Guatemala.

123. Polyotidium Garay. Small (up to 18 cm tall), epiphytic, caespitose, sympodial herbs. Pseudobulb unifoliate, cylindrical. Leaf bifacial but nearly terete, articulate, eventually deciduous; inflorescence one (rarely two), lateral, unbranched to weakly branched, longer than leaves, 5–25-flowered, bracts shorter than pedicels; flowers resupinate, spurless, red to red-orange; sepals and petals free, but lateral sepals fused most of their length, obovate. Only one sp., P. huebneri (Mansf.) Garay, reported from the Guiana Shield from Amazonian and Orinocan regions of Colombia, Venezuela, and Amazonas state in Brazil, growing epiphytically in shady sites at 100–200m in wet forests.

124. Psychopsis Raf. 5 spp., Costa Rica to French Guiana and Trinidad, Colombia to Peru, Roraima state in N Brazil (3, one endemic).

125. Pterostemma Kraenzl. (inc. Hirtzia) 5 spp., Colombia to Ecuador.

126. Quekettia Lindl. 7 spp., northern South America up to Peru and Brazil (3, none endemics).

127. Rauhiella Pabst & Braga. Small (up to 3 cm tall) epiphytic, caespitose, sympodial herbs; leaves duplicate, bifacial; inflorescences 1–3, lateral, glabrous, unbranched, pendent, usually longer than leaves, 1–10-flowered; bracts nearly as long as pedicels; flowers usually resupinate or variously arranged, asymmetrical, spurless, yellow-green to yellow, often with reddish purple spots on labellum; sepals and petals free, lanceolate, similar in size and shape except for longer lateral sepals, which are also falcate. Three spp. restricted to the Atlantic Forest rain forest of E Brazil in the States of Bahia and Rio de Janeiro, growing epiphytically at low to middle elevations, ca. sea level to 1,300metres, in wet forests. Where they grow in the littoral zone, conditions are much drier.

128. Rhynchostele Rchb.f. 17 spp., Mexico to Panamá, two disjuncts in NW Venezuela.

129. Rodriguezia Ruiz & Pav. Moderate (up to 25 cm high), epiphytic, caespitose to scandent, sympodial herbs, sometimes partially psygmoid; leaves 1–4, mostly bifacial; inflorescences 1–4, lateral, 5–30- flowered, bracts shorter than pedicels; flowers resupinate or of irrgular symmetry, yellow, pink, carmine, greenish yellow or white, often with red and yellow markings on all parts, particularly yellow on labellum callus. 52 spp. from Mexico south through Central America and from Colombia and Venezuela south to Argentina, Brazil (24, 14 endemics), Bolivia, and Peru, east up to French Guiana. R. lanceolata Ruiz & Pav. is widely distributed throughout this range and also found in many parts of the Caribbean; 50 spp. in South America; most spp. grow on smaller branches of their host trees, in wet forests, from sea level up to 1,500m.

130. Rossioglossum (Schltr.) Garay & G.C.Kenn. 11 spp., Mexico to Peru and Venezuela, two in South America.

131. Sanderella Kuntze. Small (5–8 cm tall), epiphytic, caespitose, sympodial herbs. Leaf bifacial, articulate, and eventually deciduous; inflorescences one or two, lateral, glabrous, unbranched, roughly length of leaves, 5–10- flowered, bracts shorter than pedicels; flowers mostly resupinate (although orientation often irregular), yellow-green with a few reddish brown markings on labellum; dorsal sepal and petals free, often similar in size and shape, lateral sepals fused to one-half their length, lanceolate. Two spp., in a restricted area of S Brazil (both, none endemic) and Misiones Province in Argentina up to Minas Gerais and Bolivia, growing at low to middle elevations (200–900m), generally on small branches near water.

132. Santanderella P.Ortiz. Only one sp., S. amado-rinconiana P. Ortiz, endemic to Colombia.

133. Saundersia Rchb.f. Small (5–8 cm tall), epiphytic, caespitose, sympodial herbs. Pseudobulb unifoliate, ellipsoidal, often covered by bracts. Leaf conduplicate, bifacial, articulate, coriaceous, and eventually deciduous; inflorescences one or two, lateral, glabrous, unbranched or branched, roughly the length of leaves or less, 5–40-flowered, bracts as long or longer than pedicels; flowers mostly resupinate although orientation often irregular, yellow-green, reddish to white, often with a few reddish brown markings on sepals and petals. Sepal and petals free, similar in size and shape, lanceolate. Two spp., from SE Brazil. These two spp. grow epiphytically at low to middle elevations (200–900m).

134. Schunkea Senghas. Small (4–8 cm tall), epiphytic, caespitose, sympodial herbs; pseudobulb nearly terete, unifoliate; leaf conduplicate, bifacial, coriaceous, articulate, and eventually deciduous; inflorescences one or two, lateral, glabrous, unbranched, shorter than leaves, 2–8-flowered, bracts shorter than pedicels; flowers resupinate, white to cream with red-pink central stripes on perianth; sepals and petals free, similar in size and shape, oblanceolate. Only one sp., S. vierlingii Senghas has only been collected once, in Espírito Santo State in SE Brazil, around 600m in open, wet forests.

135. Seegeriella Senghas. Two spp., Ecuador to Bolivia.

136. Solenidium Lindl. Small (8–15 cm tall), epiphytic, caespitose, sympodial herbs; leaves one or two, bifacial, articulate and eventually deciduous; inflorescences one or two, lateral, glabrous, unbranched, longer than leaves, 5–30-flowered, bracts shorter than pedicels; flowers resupinate, yellow-green, yellow to white, with reddish brown markings on sepals and petals; sepals and petals free, similar in size and shape, oblanceolate. Two spp. occurring in Amazonian/Orinocan South America, from Colombia to Peru and Bolivia and Venezuela and Guyana to Brazil (both spp., none endemics), in wet forests at elevations of 500– 2,000m.

137. Suarezia Dodson. Only one sp., S. ecuadorana Dodson, Ecuador.

138. Sutrina Lindl. Two spp., Peru and Bolivia one endemic each.

139. Systeloglossum Schltr. 5 spp., three in Costa Rica to Panamá, and two in Colombia, Ecuador to Peru.

140. Telipogon Kunth. (inc. Stellilabium) Small to medium (4–25 cm tall), epiphytic, rarely terrestrial, caespitose, sympodial herbs without pseudobulbs; leaves 2–15; inflorescences one or two, lateral (sometimes appearing terminal), unbranched, arching to erect, longer than leaves, 1–10-flowered, but often with single flowers opening successively; flowers resupinate, yellow-green to yellow, often suffused with red-purple and with dark veins, nearly always with a series of spots, hairs, and other markings particularly on labellum; sepal and petals similar in size and shape or petals larger, oblanceolate to nearly triangular. 245 spp. from Costa Rica and Panamá in Central America to Colombia, Venezuela, Ecuador, Bolivia, and Peru in South America (188); the greatest species diversity is found in the Andes of Colombia and Ecuador, with fewer than 4 species are widely distributed in distribution, e.g., Venezuela to Peru; only one Central American species occur in South America; these species occur on all axes of trees and shrubs and occasionally in nearly terrestrial conditions in mossy areas at 1,400–3,000m in perpetually wet forests, often in near-páramo conditions; flowers having the appearance of an insect in the center of the usually roundish flower; many species are brightly colored with yellow, brown and red spots, blotches, vein-lining and reticulations.

141. Thysanoglossa Porto & Brade. Epiphytic, sympodial herbs; pseudobulb petiole-like, unifoliate. Leaf dorsiventrally flattened, bifacial, V-shaped or semi-terete, articulate, epetiolate; inflorescences one or two, lateral, unbranched, longer than leaves, 1–6-flowered, bracts shorter than pedicels; flowers mostly resupinate but variable, spurless, greenish yellow to bright yellow; sepals and petals free, oblanceolate, similar in size and shape. Three spp. found only in SE Brazil in Atlantic Forest, growing epiphytically in shady sites at 150–1,500m in humid forests.

142. Tolumnia Raf. 29 spp. from Caribbean, one up to Florida, another up to Colombia, Venezuela, Guyana and disjunct in French Guiana.

143. Trichocentrum Poepp. & Endl. (inc. Cohniella) Small to large (up to 70 cm tall), epiphytic (rarely lithophytic), sympodial herbs; leaves one or rarely two per sympodium, coriaceous, dorsiventrally flattened or terete, articulate, epetiolate; inflorescences one or two, lateral, 1–80-flowered, sometimes weakly branched; bracts shorter than pedicels; flowers resupinate, spurless to long-spurred, pink, purple, yellow, white, dark brown, reddish brown, pale cream with brown to reddish brown spots; sepals and petals free, oblanceolate, similar in size and shape or petals in some spp. much larger and broader. 85 spp. found throughout the Neotropics, from Florida and Mexico through Central America, the Caribbean to northern Argentina, Uruguay, S Brazil (14, 4 endemics), Paraguay, and Peru, 45 in South America. In spite of their seemingly xerophytic habits, the 65 spp. of Trichocentrum s.l. mostly grow in relatively shady and humid sites from sea level to 2,800m, often growing on trunks and other larger axes, in many cases found in deep shade and nearly completely buried in moss.

144. Trichoceros Kunth. 7 spp., Colombia to Bolivia

145. Trichopilia Lindl. 39 spp., Mexico, Belize to Panamá, Caribbean, Venezuela to Suriname, Colombia to Bolivia, N to SE Brazil (6, two endemics); 28 spp. in South America.

146. Trizeuxis Lindl. Only one sp., T. falcata Lindl., Trinidad, Costa Rica to Guyana, Colombia to Bolivia, Brazil.

147. Vitekorchis Romowicz & Szlach. Two spp., Venezuela to Peru.

148. Warmingia Rchb.f. 5 spp. very disjunct, one in Central America, one in Colombia and Ecuador, another in Peru and two in Brazil, one of then up to Argentina.

149. Zelenkoa M.W.Chase & N.H.Williams. Only one sp., Z. onusta (Lindl.) M.W. Chase & N.H. Williams, Colombia to Peru.

150. Zygostates Lindl. (inc. Centroglossa) Small (up to 10 cm tall), epiphytic, sympodial herbs. Pseudobulb globose, subtended by 1–15 foliaceous bracts on a leafy stem, sometimes branched; roots glabrous or pilose; leaves one or two per sympodium, coriaceous, bifacial, articulate, usually petiolate; inflorescences 1–5, lateral, generally unbranched, equal to or longer than leaves, 1–15-flowered; bracts shorter than pedicels; flowers resupinate or not, sometimes asymmetrical, spurless, some parts green, others white to cream, orange or yellowish green with green to orange spots on the labellum callus; sepals and petals free, similar in size and shape or petals broader. 27 spp. found from Colombia, Ecuador, Venezuela, and Guyana to S Brazil (20, 18 endemics), Argentina (Misiones), and Paraguay, especially rich in spp. in SE Brazil, occasionally seen in some specialist collections, grown for their floriferousness (in spite of their small size).

∎ SUBTRIBE ZYGOPETALINAE (35/c. 430) ‣ outsider Stenotyla (9; S Mexico, Central America).

151. Aetheorhyncha Dressler.Only one sp., A. andreettae (Jenny) Dressler, endemic to Ecuador.

152. Aganisia Lindl. Epiphytic or terrestrial; leaves plicate, weakly coriaceous, narrowed at base into a conduplicate, channeled petiole, clear to dark green, shiny on upper surface; inflorescence lateral, a few(2)- to many(10)-flowered raceme; Flowers resupinate, sepals white to pale bluelavender or blue-mauve with adaxial surfaces flushed pink, petals white to pale blue-lavender or bluish mauve, pale pink-flushed white within; labellum white with yellow centre or blue-lavender with dark purple-lavender margins or gold-bronze with a mauve red or bluish purple centre. 3 spp. from Trinidad and tropical South America, where it has been recorded from Venezuela to central Brazil in Mato Grosso state (all spp., one endemic), and from the Amazonian regions of Frech Guiana to Colombia, Ecuador, Peru. Species of the genus occur as epiphytes in hot, tropical to premontane rain forests, usually at elevations of 100–500m, rarely up to 1,000m.

153. Batemannia Lindl. Epiphytic, caespitose herbs; leaves membranaceous to subcoriaceous; inflorescence lateral, a 1–7-flowered raceme; flowers resupinate, sepals and petals greenish yellow or olive green, often suffused with brown, or with purple-brown blotches, labellum white or pale yellow, often finely spotted with purple, column white, spotted purple underneath. 5 spp. from tropical South America, ranging from Trinidad to northern Brazil (2, none endemics) and from Venezuela to the Amazonian regions of Colombia, Ecuador, Peru, and Bolivia; epiphytes in hot, tropical wet forests at elevations of 100–600m.

154. Benzingia Dodson. (inc. Chondrorrhyncha p.p.) 11 spp. from Colombia to Peru, one up to Costa Rica.

155. Chaubardia Rchb.f. Three spp., French Guiana to Venezuela, Colombia to Bolivia, N to SE Brazil (2, none endemics).

156. Chaubardiella Garay. 8 spp. from Colombia to Peru, two up to Central America, C. tigrina (Garay & Dunst.) Garay up to French Guiana.

157. Cheiradenia Lindl. Only one sp., C. cuspidata Lindl., endemic to Guiana Shield, in elevation ranges 100 – 900m, French Guiana to Venezuela, and Amapá state in N Brazil (only one record, never found in live form).

158. Chondrorhyncha Lindl. (off Benzingia p.p., Ixyophora p.p.) 7 spp., Venezuela to Ecuador.

159. Chondroscaphe (Dressler) Senghas & G.Gerlach. 12 spp., Costa Rica to Peru and Venezuela, 12 in South America.

160. Cochleanthes Raf. 3 spp., one wider in Mexico to Panamá, Caribbean, Venezuela to Ecuador and Brazil, and two restricted in Central America and Trinidad & Tobago.

161. Cryptarrhena R.Br. Three spp., all wider, Mexico to Panamá, Jamaica, Trinidad, French Guiana to Venezuela, Colombia to Bolivia, Brazil (3, none endemic).

162. Daiotyla Dressler. 4 spp., two in Costa Rica and Panamá and two endemics to Colombia.

163. Dichaea Lindl. Epiphytic, caespitose or scandent herbs without pseudobulbs; inflorescences lateral, single-flowered, supraaxillary, produced sequentially or rarely simultaneously; flowers resupinate, ringent to spreading, often scented, sepals and petals ivory white to greenish white or orange-grey, sepals mostly flecked with purple-violet toward base, petals commonly heavily spotted or blotched with purpleviolet. 130 spp., distributed from Mexico through Central America and Caribbean to Bolivia and Brazil (31, 14 endemics), with greatest diversity in Andean South America (100); epiphytes (rarely terrestrials), mostly restricted to shady places in the understorey of wet forests from 0–2,500m. With a few exceptions, spp. of Dichaea are shade-loving plants, invariably growing in subdued light on the main trunk (often near the soil) and large, shaded branches. Most spp. are not exposed to direct sunlight, and plants usually establish themselves among thick layers of mosses, which ultimately cover large portions of stems and leaves.

164. Echinorhyncha Dressler. 5 spp., Colombia to Ecuador.

165. Euryblema Dressler. Two spp., Panamá and Colombia one endemic each.

166. Galeottia Ness. Epiphytic, pseudobulbous, caespitose herbs; leaves subplicate-venose, membranaceous to subcoriaceous, lanceolate-elliptic to oblanceolate, acute, abruptly acuminate, abaxially obscurely carinate along nerves, narrowed at base into a distinct, conduplicate petiole, clear to dark green, shiny on upper surface; inflorescence lateral, one or two per shoot, a 2–8- flowered raceme; flowers showy, resupinate, sepals and petals yellowish green to cream, mostly longitudinally striped with reddish brown or with purple-brown stripes and blotches, rarely suffused with chestnut brown, labellum usually creamy white, rarely almost solid red or with yellow lateral lobes, midlobe often striped with purple, callus red or variously striped with purple, column white, striped purple on the underside. 12 spp. from South America, one ranging up to S Mexico through Central America, with highest diversity on the western slopes of the Peruvian and Colombian Andes also in Guianas; the spp. of the genus occur as epiphytes or terrestrials in wet forests at elevations of 150–2,500m in shady to partially exposed spots; 5 spp. in Brazil, none endemics.

167. Hoehneella Ruschi. Epiphytic, caespitose, pseudobulbous herbs; leaves conduplicate, articulate, membranaceous, dark velvety green, paler abaxially; inflorescence lateral, single-flowered, produced from axils of lower sheaths; flowers resupinate, parts not completely spreading, sepals and petals greenish yellow, labellum white with a violet callus. Two spp., one in São Paulo, another in Espírito Santo state, as shade-epiphytes on mossy branches in evergreen, wet forests at 600–1,100m; flowering has been recorded in January, but may occur at other times.

168. Huntleya Bateman ex. Lindl. Epiphytic, often large caespitose herbs without pseudobulbs; roots terete, produced from rhizome; stem enclosed by 7–16 imbricating sheaths, upper ones foliaceous; leaves conduplicate, articulate, membranaceous to coriaceous, oblanceolate to narrowly obovate, acute to acuminate, abaxially carinate, narrowed at base into an indistinct, conduplicate petiole, grass green to dark green; inflorescence one or more per shoot, lateral, single-flowered, produced from axils of lower sheaths, peduncle terete, erect, provided with 1–3 conduplicate, clasping bracts; floral bract double, conduplicate, membranaceous, shorter than ovary, the external one loose, widely ovate, acute, the subopposite internal bractlet narrowly lanceolate to ligulate; flowers resupinate, usually scented, sepals and petals glossy cream to yellow, frequently white in basal third. 15 spp. from South America (centre of diversity in the northern Andean regions of Colombia and Ecuador), three ranging from Nicaragua; mostly restricted to extremely wet forests, usually at medium and low elevations (from sea level to 800–1,000m); Huntleya mostly grow as large epiphytes on trunks and large branches, where the thick rhizome often assumes a creeping habit; two spp. in Brazil, none endemics.

169. Ixyophora Dressler. (inc. Chondrorrhyncha p.p.) 7 spp., 5 from Colombia to Peru and two endemics to Bolivia.

170. Kefersteinia Rchb. f. Epiphytic, caespitose herbs without pseudobulbs. Stem abbreviated, Leaves membranaceous; inflorescence lateral, 1–20 per shoot, single-flowered, produced from axils of lower sheaths, peduncle terete, arching or pendent, provided with one or two basal, triangular, membranaceous to scarious bracts; Flowers resupinate, spreading or campanulate, sepals and petals pure white or yellow to cream white or pale green, rarely pale pink to pale purple, often spotted with purple to dark red-brown or tan- brown, heavily marked with orange-red or finely spotted purple toward base, rarely covered with large purple- brown blotches, labellum white or pinkish white to cream or yellow, spotted and flecked with purple to almost solid purple, rarely with a large, central, purple blotch, callus white to yellow, mostly finely spotted with purple, rarely solid purple or pink.

64 spp. collectively ranging from S Mexico through Panamá and Venezuela and Colombia to French Guiana and Bolivia in South America (50), K. mystacina Rchb. f. up to Brazil (by a single non academic collection, with a unknown range), most diverse along the Andes of Ecuador and Peru (with 21 and 17 spp. recorded, respectively), and it is particularly well represented in the mountainous areas of Central America up to Costa Rica, where 11 spp. have been recorded so far; diversity rapidly diminishes toward the north, with a single spp. (K. tinschertiana Pupulin) known from Guatemala and S Mexico; epiphytes in shady habitats, often growing on the trunks and mossy oldest branches of trees alongside streams; the habitat of Kefersteinia ranges from tropical warm to evergreen and wet forests at elevations of 100–2,500m, although most spp. are found in premontane forests at mid-elevations (900–1,500m). Some spp. (namely K. bengasahra D.E.Benn. & Christenson, K. expansa Rchb.f., K. retanae G.Gerlach, K. stevensonii Dressler) are restricted to warm, tropical forests under 500m, whereas K. guacamayoana Dodson & Hirtz, K. aurorae D.E.Benn. & Christenson, K. pusilla (C.Schweinf.) C.Schweinf., K. pellita, and K. tolimensis Schltr. have been exclusively recorded from forests at elevations of 1,700–2,500m; flowering has been recorded throughout the year, although some spp. seem to flower consistently during the dry season and others during the rainy months.

171. Koellensteinia Rchb. f. Terrestrial, rarely epiphytic, caespitose, mostly pseudobulbous herbs; leaves membranaceous to subcoriaceous; inflorescence a lateral, few-flowered; flowers resupinate, sepals pale cream to pale green or clear light yellow, sometimes flushed with pink or transversely marked with rose, violet or red-brown toward base, petals pale cream to pale green or clear light yellow, sometimes sparsely dotted or transversely striped with purple, labellum white, mostly with transverse, purple stripes or with purple spots toward base, often flushed with yellow around callus. 17 spp. distributed in Puerto Rico and Trinidad, Central America (Belize and Panamá), and tropical South America (all species) from Venezuela to Brazil (10, 6 endemics) and from Colombia to Peru and Bolivia along the Amazonian watershed of the Andes; the centre of diversity is NE Brazil; terrestrials in humid soils and among mossy rocks on the floor of bright, open forests and grasslands and less frequently epiphytes on shaded tree trunks in evergreen forests at 100–2,300m.

172. Neogardneria Schltr. ex Garay. Epiphytic, caespitose, pseudobulbous herbs; leaves medium to dark green, shiny on upper surface; inflorescence lateral, one or two per shoot, a 2–6-flowered raceme produced from base of developing pseudobulb and emerging from axils of lower sheaths, peduncle terete, suberect to erect, rachis fractiflex; floral bracts lanceolate, conduplicate-cymbiform, loose, shorter than ovary; flowers resupinate, spreading, greenish to light yellow, labellum white to greenish white, marked with red-purple freckles, callus and column yellow. Only one sp., N. murrayana (Gardner ex Hook.) Schltr. ex Garay, from SE Brazil, recorded from Espírito Santo and Río de Janeiro states. Epiphytes in the shade on mossy old tree trunks in evergreen cloud forests at 1200–1,600m, mostly along welldrained ridges of the Atlantic chains of central and S Brazil; flowering has been recorded from December through March. (FP).

173. Otostylis Schltr. Terrestrial, caespitose, pseudobulbous herbs; leaves membranaceous; inflorescence lateral, emerging from axils of lower sheaths, a successively flowered raceme, rarely branching at base, with 5–20 flowers open simultaneously; flowers resupinate, with sepals and petals white to pale cream, sometimes flushed with pale pink toward the apex, labellum white to pale cream, disc pale yellow to yellow, sometimes spotted purple toward base, column white, variously marked with purple-violet at base. Two spp. ranging from Trinidad to Venezuela and Brazil (both, none endemics) and from Colombia to Peru, with most taxa recorded from the Guiana Shield; terrestrial in moist and swampy peaty soils in open areas of evergreen, damp rain forests and semi-dwarf forests at 100–1,700m; flowering has been recorded at least from April to June.

174. Pabstia Garay. Epiphytic, lithophytic (occasionaly terrestrial), caespitose, pseudobulbous herbs; leaves subcoriaceous; inflorescence lateral, 1–4 per shoot, one few – flowered; flowers scented, fleshy, sepals creamy white to green, rarely striped with red-brown, lateral sepals sometimes sparsely spotted with red-purple, petals creamy white to green, heavily spotted/blotched or striped with purple red, labellum white, blotched/striped with purple to solid violet apically, sometimes flushed with green toward apical margins, disc and callus greenish white to pale violet, column white flecked with purple to pale violet. 5 spp. endemics to the coastal regions of SE Brazil (Espírito Santo, Río de Janeiro, Santa Catarina) in South America; epiphytes and lithophytes in shady and humid locations in forests along the coastal mountains of SE Brazil at 200–1,500m; flowering occurs from July to December.

175. Paradisanthus Rchb.f. Terrestrial, caespitose, pseudobulbous herbs; leaves plicate, submembranaceous; inflorescence lateral, a successively few-flowered, rarely branching raceme; flowers resupinate, with creamy white to pale green sepals and petals, mostly transversely striped with brownish red toward base, labellum white, sometimes spotted violet along margins of lateral lobes, column white, with ventral surface blotched red to dark purple-brown. 4 spp., native to the coastal regions of central and S Brazil, where they have been recorded from the states of Bahia to Rio Grande do Sul; terrestrial plants in the open shade of tropical and subtropical, low-elevation rain forests, where they are usually found in dry soils with the roots growing among surface detritus.

176. Pescatoria Rchb.f. Epiphytic, caespitose herbs without pseudobulbs; leaves membranaceous to coriaceous, erect to erect-spreading; inflorescence one or more per shoot, lateral, single-flowered; flowers resupinate, fleshy, sepals and petals white to creamy yellow, greenish cream or pink to solid purpleviolet, mostly flushed with lavender toward apex, rarely striped with lavender or purple in basal half, labellum white to pink or yellow, midlobe often flushed lavender or purple apically, rarely solid violet, callus yellow (rarely white) to dark purple, ribs of callus usually purple. 22 spp. and two named natural hybrids, ranging from Costa Rica to Brazil and Peru, up French Guiana, centred in NW South America, mainly diverse along the Andes of Colombia and Ecuador, with 16 and 9 spp., respectively. spp. diversity rapidly diminishes toward the north (a single spp. recorded for Costa Rica) as well as toward the southern end of the Andes (only one sp. in Peru) and in E South America (2, wider); two spp. are recorded from Venezuela and the Guyanas to N Brazil. Most spp. of Pescatoria are native to wet forests at medium elevations (around 800–1,500m), but P. hemixantha (Rchb.f.) Dressler is found in the warmer forests of the tropical belt at 200–500m; Pescatoria are large epiphytes, usually restricted to trunks and large branches of their hosts.

177. Pridgeonia Pupulin. Epiphytic, caespitose herbs without pseudobulbs; roots terete, thick, 2.5–3.0 mm in diam., produced from the short rhizome. Only one sp., P. insignis Pupulin, endemic to SE Ecuador, near Peruvian border.

178. Promenaea Lindl. Epiphytic, caespitose, pseudobulbous herbs; leaves subplicate-venose, grey-green to medium green, paler abaxially; inflorescence lateral, 1–4 per shoot, 1-(rarely 2-)flowered, flowers resupinate, large for plant, spreading, sepals and petals creamy white to pale greenish yellow to bright yellow, immaculate or variously spotted or transversely striped with red-purple to violet-brown, labellum white to cream with yellow apex, greenish yellow to bright yellow, or solid black-purple, often heavily spotted with red purple, mostly in basal portion. 15 spp., all endemics to E Brazil, Bahia to Rio Grande do Sul; epiphytes in shady and humid forests, mostly along the Atlantic slopes of Serra do Mar in SE Brazil, at 1,000–2,000m.

179. Stenia Lindl. 22 spp., Trinidad, Guyana, Venezuela, Colombia to Bolivia and Brazil (1, wider).

180. Warczewiczella Rchb.f. 11 spp., from French Guiana to Venezuela, Colombia to Bolivia, N to SE Brazil, two up to Honduras and Panamá; 3 spp. in Brazil, two endemics.

181. Warrea Lindl. Terrestrial, rarely epiphytic, caespitose, pseudobulbous herbs; leaves plicate, membranaceous; inflorescence lateral, a 4–14-flowered raceme; flowers resupinate, usually nodding, subglobose, sepals white to pale orange-yellow on inner surface, sometimes blotched with rose toward base, abaxially cream to orange, flushed purple, petals white to yellowish cream, often flecked with rose or pale purple, labellum white to yellow, with an orange to purple basal blotch or blotch restricted to disc, or flushed rose-purple and striped with purple at apex, the callus white to yellow, apically purple or solid purple. 3 spp., one wider in tropical South America, one only in Mexico and Central America and one endemic to Peru; terrestrials (rarely epiphytes) in decaying leaf matter, mostly in the shaded understorey of dense, wet forests and occasionally on drier slopes and in more exposed conditions. Records of Warrea distribution indicate that plants are mostly restricted to premontane forests, occasionally ranging to the tropical belt of moist to wet forests at 600–1,500m.

182. Warreella Schltr. Two spp., Colombia and Venezuela one endemic each.

183. Warreopsis Garay. 4 spp., one in Panamá and three from Venezuela to Ecuador.

184. Zygopetalum Hook. Terrestrial, epiphytic or lithophytic, pseudobulbous (rarely without, in which case with a prostrate, indeterminate rhizome becoming erect apically), caespitose to scrambling herbs; leaves plicate-venose, membranaceous to subcoriaceous; inflorescence lateral, one or two per shoot, a 3–10- flowered racem; flowers resupinate, showy, fragrant, sepals and petals yellowish green to green, heavily blotched or spotted with crimson or chestnut brown, rarely solid reddish brown, labellum white to purplish violet, heavily striped along veins with purple, violet or red, callus white to purple-violet. 14 spp. from Brazil, two also ranging to Paraguay, northern Argentina, Bolivia, and Peru; terrestrial or epiphytic (facultatively terrestrial and lithophytic) in wet forests, mostly at elevations of 1,000–1,800m in shady to partially exposed situations. Z. maxillare Lodd. is almost exclusively restricted to tree-ferns in wet and shady conditions.

185. Zygosepalum Rchb. f. Terrestrial or epiphytic, pseudobulbous, rhizomatous-creeping or caespitose herbs; leaves membranaceous to subcoriaceous; inflorescence lateral, a 1–7-flowered raceme produced from base of immature or mature pseudobulb; flowers resupinate, showy, sepals and petals pink or pale reddish brown to deep purple with cream margins, or yellowish green to green blotched with red-brown, labellum white to cream with a few purple lines at base to striped with purple-violet, or blotched purple at base, callus white striped purple to solid purple-violet, column white or yellowish to pale green, variously flecked with purple on ventral surface. 7 spp., mostly native to the Amazon and Orinoco basins and the Guiana Shield, which range from Venezuela to northern Brazil (4, none endemics) and from Colombia to Ecuador and Peru, in the Amazonian watershed of the Andes; Zygosepalum is centred in Venezuela, with five recorded spp.; epiphytic or terrestrial (facultatively epiphytic) plants in tropical warm to montane wet forests at elevations of 200–2,500m in shady to partially exposed situations. Z. labiosum (Rich.) C. Schweinf. and closely related spp. are almost exclusively restricted to lower portions of trees, close to the water along streams and rivers in shady conditions. spp. of tepuis, such as Z. tatei and Z. angustilabium, grow as terrestrials or epiphytes in dense, tangled, dwarf, wet forests, often in open conditions.

∎ SUBTRIBE ERIOPSIDINAE (1/6) ‣ a single genus.

186. Eriopsis Lindl. Epiphytic, lithophytic or terrestrial, erect herbs; leaves to 1 m long, usually with several light green, conspicuous veins on the lower surface; inflorescence lateral, erect or arching, racemose, multiflorous (sometimes more than 50 flowers); flowers resupinate, 3–4 cm in diameter; sepals and petals free, similar, spreading, dull yellow to orange with brownish flush and maroon margins, elliptic–ovate. 6 spp. distributed from Guatemala and Belize in Central America south to the W Andes of Peru and also in the lowlands of Amazonia in Venezuela, Colombia, Ecuador, Peru, Bolivia, and Brazil (2, none endemics), between sea level and 2,400m.

E. biloba Lindl. and E. rutidobulbon Hook. mostly grow as lithophytes or terrestrials in savannas, preferring open places in direct sun; in Ecuador, they are often terrestrial on steep roadbanks. Plants of E. biloba are adapted to seasonally burned habitats in the Gran Sabana in southern Venezuela; here they survive fi res with their pseudobulbs partly buried in the sand. E. biloba is a dominant orchid on the summit of Auyan-tepui in southern Venezuela. E. rutidobulbon occurs in W Colombia and Ecuador at elevations from 1200–1,700m. E. sceptrum Rchb. f. & Warsz. occurs in lowland Amazonia of Venezuela, Colombia, Ecuador, and Peru; it often forms large specimens close to the water and is also found in inundated forests just above the water-line.

∎ SUBTRIBE MAXILLARIINAE (12/c. 820) ‣ all genera in South America.

187. Anguloa Ruiz & Pav. 13 spp., Venezuela to Bolivia.

188. Bifrenaria Lindl. Epiphytic, lithophytic or rarely terrestrial, sympodial herbs to 60 cm tall, caespitose to long-rhizomatous; leaves membranaceous to coriaceous; inflorescence racemose, lateral, erect or pendent, 1–10-flowered; flowers resupinate, zygomorphic, fragrant or not, of various colours, often long-lived; sepals concave, ovate, rarely lanceolate or oblong, rarely attenuate or mucronate, lateral sepals parallel to column or divergent, sometimes basally prolonged into a spur that is fused to the column foot; petals obovate, rarely rhombic or lanceolate, asymmetric and always smaller than sepals. 22 spp. occuring in Brazil (20, 15 endemics), Colombia, Ecuador, Guyana, Peru, and Venezuela as well as the islands of Trinidad and Tobago. The centre of diversity is the coastal Atlantic rain forest in SE Brazil, typically epiphytes from 50–1,500m, but some spp. are facultative terrestrials (B. atropurpurea Lindl. and B. longicornis Lindl.) or lithophytes (B. harrisoniae (Hook.) Rchb.f., B. inodora Lindl.). B. tyrianthina (Loudon) Rchb. f. is the only spp. that is exclusively lithophytic; most spp. are epiphytes in the coastal Atlantic forest of Brazil or less frequently as lithophytes in rocky grasslands (campos rupestres). Two spp. occur only in Amazonia.

189. Guanchezia G.A.Romero & G.Carnevali. Caespitose, terrestrial or rarely lithophytic or epiphytic, sympodial herbs. Leaf articulate, long-petiolate, up to 60 cm long; inflorescence arising from base of the most recent pseudobulb, erect, racemose, 1–3 flowered; flowers fleshy, resupinate, nutant, showy; floral bracts concave, narrowly elliptic, acute, shorter than the ovary, sepals and petals yellowish, labellum white, heavily veined with dark purple; sepals oblong-lanceolate, lateral sepals oblique and adnate to the column foot; petals oblong-lanceolate, smaller than sepals. Only one sp., G. maguirei (C. Schweinf.) G.A. Romero & Carnevali, endemic to the Guiana Shield Venezuela, Colombia and N Amazonas states in Brazil, in open tepui associations, often in bogs, at 1,200–1,600m; also reported growing on Cerro Avispa and Cerro Autana, Venezuela, in poorly drained, boggy, acid, sandy, or peaty soil at 1,500m with a minimum night temperature of 14–17° C. The pseudobulbs are frequently buried to their apices in the boggy soil. Smaller plants were rarely found growing epiphytically on trees in ravines.

190. Horvatia Garay. Only one sp., H. andicola Garay, Ecuador.

191. Lycaste Lindl. Epiphytic, lithophytic or terrestrial, caespitose, erect, sympodial herbs; leaves distichous, plicate, elliptic, with 3–5 pairs of prominent veins plus midvein, basally attenuate forming a variously developed pseudopetiole, in some spp. deciduous after maturation of the pseudobulb and vascular bundles of abscission layer forming sharp spines on the apex of pseudobulb; inflorescences lateral, 1-(rarely, 2-) flowered; flowers showy, resupinate, fleshy, spreading, usually long-lived; bracts hooded, partially or completely concealing the terete ovary. 43 spp. from Mexico to South America (26, only two in Brazil, none endemics); epiphytic plants of Lycaste occur in light woodland, usually on trees with rough bark where the roots can run in the moist crevices occupied by coexisting mosses. Terrestrial or lithophytic plants are found in leaf litter and shade on the forest floor, often associated with limestone boulders or cliffs. They flower better at woodland margins, light woodland or in coppiced areas.

192. Maxillaria Ruiz. & Pav. (inc. Brasiliorchis, Christensonella, Cryptocentrum, Mormolyca, Heterotaxis, Trigonidium, Xanthoxeranthella, Pseudocymbidium) Epiphytic, lithophytic, or terrestrial, sympodial herbs, caespitose or rhizomatous, erect to (rarely) pendent; inflorescence single-flowered; flowers inconspicuous to showy, campanulate to spreading, resupinate or non-resupinate, usually long-lived, occasionally fragrant; bracts shorter or longer than the pedicel plus ovary; sepals and petals free, submembranous to fleshy, fibres evident when torn; possibly nectar spur species occur South America.

688 spp., ranging from northern Mexico, Central America, and the Lesser Antilles (Guadeloupe, Dominica) south to S Brazil (106, 39 endemics), Bolivia, and northern Argentina; 586 spp. in South America, mainly Colombia and Ecuador. Most Maxillaria spp. are epiphytes in montane forests with highest diversity in the Andes of Colombia, Ecuador, and Peru. In these habitats, the distinction between epiphytes and terrestrials is often vague because the same spp. may occur in tree branches as well as on steep slopes or roadbanks. Some spp. inhabit dry tropical forests or forests with a strong dry season, but we are unaware of any deciduous spp. spp. of Maxillaria can be found from near sea level up to 3,500m in subpáramo vegetation in the Andes (e.g., M. rotundilabia C.Schweinf.).

193. Neomoorea Rolfe. Only one sp., N. wallisii (Rchb. f.) Schltr., Panamá to Colombia.

194. Rudolfiella Hoehne. 7 spp. from French Guiana to Venezuela, Colombia to Peru, N Brazil, two up to Panamá and Trinidad; 3 spp. in Brazil, one endemic.

195. Scuticaria Lindl. 12 spp., French Guiana to Venezuela, Colombia to Peru, Brazil (9, 7 endemics).

196. Sudamerlycaste Archila. 33 spp., Caribbean, Venezuela to Peru, one, S. rossyi (Hoehne) Archila, disjunct spp. from SE Brazil; 31 in South America.

197. Teuscheria Garay. 7 spp., SE. Mexico to Ecuador and Venezuela, possibly Suriname; 7 spp. in South America.

198. Xylobium Lindl. 18 spp., Mexico, Belize to Panamá, Caribbean French Guiana to Venezuela, Colombia to Bolivia, over Brazil (4, none endemics); 17 spp. in South America.

∎ SUBTRIBE COELIOPSIDINAE (3/19) ‣ three genera, all in South America.

199. Coeliopsis Rchb.f. Only one sp., C. hyacinthosma Rchb. f., Costa Rica to Colombia and Ecuador.

200. Lycomormium Rchb.f. 5 spp., Colombia, Ecuador and Peru.

201. Peristeria Hook. Terrestrial or epiphytic herbs; roots tomentose, lanate; pseudobulbs clustered, 2–5 leaved; leaves elliptic-lanceolate, plicate, often deciduous; inflorescence basal, erect or pendent racemes; flowers fleshy, subglobose, fragrant, pure white to yellowish green. 13 spp. from northern South America to the Amazonian lowlands of Peru, Bolivia and Brazil (5, 3 endemics) up to Trinidad and Tobago. P. elata Hook. is terrestrial with large, round pseudobulbs up to 17 cm in diameter, growing mostly in full sunlight; other spp. are mainly epiphytic in the shady, humid conditions of the lower canopy but are also found in loose humus on the ground.

∎ SUBTRIBE STANHOPEINAE (20/c. 300) ‣ outsiders Horichia (1; H. dressleri; Panamá), Lacaena (2; L. bicolor, L. spectabilis; Central America).

202. Acineta Lindl. 10 spp., Mexico to Suriname, Colombia to Peru, N Brazil (1, no endemic); 7 in South America.

203. Archivea Christenson & Jenny. Only one sp., A. kewensis Christenson & Jenny, cited by author as a single species from Brazil, but without indication of locality.

Possibly placed hare in Stanhopienae, Archivea was proposed by Christenson & Jenny in Orchids 65 : 497 in 1996, named in reference to the fact this has been described only based on a watercolor found in the archives at Kew; it is doubtful genus until now because they never matched the plant illustrated here in his description have been collected. The species of this genus are supposed epiphytes; the authors; we observed that this species may not be Brazilian or even that it was a mistake for illustration.

Vegetatively closely resembles some species of Coryanthes and Cirrhaea, but its flowers are presented in upright basal inflorescences; we note that the upright position of the inflorescence in the illustration looks more like a freedom that the illustrator took portraying the plant and it is possible that this is pending; the inflorescence holds about ten pale yellow flowers with sepals and petals narrow and acuminate; the flowers resemble those of Horichia, which differ primarily by the lip with two lateral claws, whose lateral lobes fuse together forming a keel in the center of the lip, but much shorter and curved spine.

204. Braemia Jenny. Only one sp., B. vittata (Lindl.) Jenny, French Guiana to Venezuela, Colombia to Peru, N Brazil.

205. Cirrhaea Lindl. Lithophytic or epiphytic herbs; leaf plicate, elliptic-lanceolate, acuminate, petiole canaliculated; inflorescence wiry, glabrous, angular with some triangular, amplexicaul bracts; flowers variously coloured (even within a species), pedicel bent 90° so that flowers face away from inflorescence axis; sepals free, ovate-lanceolate, much longer than broad, concave, reflexed, lateral sepals sigmoid, asymmetric; petals smaller than sepals, narrowly spathulate, sigmoid with their broadest part above the middle. 7 spp., distributed in SE Brazil (Atlantic Forest) in the states Rio Grande do Sul, Santa Catarina, Paraná, São Paulo, Espírito Santo, and southern Bahia, as lithophytes or less frequently epiphytes on bases of tree trunks in humid evergreen or semi-deciduous forests of Atlantic Forest; they prefer sites that are semishaded with high humidity.

206. Coryanthes Hook. Epiphytic herbs, often in ant nests; leaves lanceolate or elliptic-lanceolate, plicate, conspicuously veined, subcoriaceous, acute or acuminate; inflorescence 1–10- flowered, mostly pendent but in some spp. spreading or erect (often depending on number of flowers), with some triangular, clasping bracts bearing reddish brown-bordered extrafl oral nectaries along their midvein; flowers fleshy, heavy (the heaviest orchid flower is Coryanthes bruchmuelleri Rchb.f. at more than 100 g), fragrant, often brightly coloured, with a complex morphology; flower: Sepals membranaceous, free, spreading then reflexed and inrolled distally, dorsal sepal broadly triangular, lateral sepals two or three times larger than dorsal, obliquely subfalcate; petals membranaceous, smaller than sepals, linear-lanceolate to ligulate, margins undulate. Labellum fleshy with a terete or compressed basal claw (except for Coryanthes macrocorys Rolfe), tripartite: hypochile helmet-like, entire-flat or bilobed-flat, often with hairy patches, these sometimes annulate or banded, with or without one or more horn-shaped osmophores below; mesochile canaliculate or terete, glabrous in C. sect. Coryanthes, warty or with transversely orientated lamellae in C. sect. Lamellunguis; epichile bucket-shaped, oval in longitudinal section, high at base tapering abruptly lower to tip, filled with liquid covering the callosity, apex tridentate with a larger, broader, obtuse central tooth and falcate, incurved lateral teeth, each of these with fleshy tooth-like projection at the base, inner surface slippery because of imbricate cell layers. Column ventrally flattened, without a foot, basally with two prominent, subquadrate to long-falcate, liquid-secreting glands (pleuridia), apex reflexed-clavate, with wings closely nestled next to vertical margins of epichile, clinandrium also bearing two lateral, curved, long-triangular wings; anther terminal, pollinarium consisting of two yellow, flat, incompletely fused, ovoid pollinia, a ligulate, sigmoid stipe, and a laterally tailed viscidium; stigma transverse, slit-like; rostellum ligulate, stiff.

48 spp. collectively ranging from Mexico (Oaxaca and Veracruz) through Central America southward to Bolivia (Cochabamba) and Brazil (23, mainly Amazon Basin, 12 endemics) in South America (44). Coryanthes plants grow as obligate epiphytes in humid evergreen forests from sea level up to 1,200m; in lowland forests they often grow on branches hanging over water or close to rivers, but this could be an observation artefact because they are easier to see there; near the seashore in Colombia they are frequently found in mangrove swamps. Coryanthes plants are dependent on ants of several genera (e.g. Azteca, Campanotus, Crematogaster) and form a mutualistic relationship in so-called ant gardens with other plants (myrmecophites), such as Codonanthe (Gesneriaceae), Epidendrum (Orchidaceae), Anthurium (Araceae), and some Cactaceae and Piperaceae. Ants construct a constantly growing nest, which can easily reach the size of 1 m in diameter, with a special substrate containing humus, ground, and chewed plant material. They can favour plants they want in their ‘garden’ by pruning unwanted plants with their mandibles and feeding desired plants with vertebrate feces collected on the ground. Abundant provision of nutrients allows plants to grow rapidly. In return, ants benefit from sugar solutions offered in the extra-floral nectaries of Coryanthes plants and fiercely defend them from herbivores. One negative effect on the growth of Coryanthes plants is caused by scale insects and aphids, which are milked for nutrients by ants. The damage is caused not so much by sucking actions of insects but by associated transmission of bacteria (Erwinia) in the process. Also, in their natural habitat Coryanthes plants often attract weevils (Curculionidae). Larvae of these insects live inside pseudobulbs, well protected against the omnipresent ants. Here, too, secondary infection by bacteria is more dangerous for the plant. Seed maturation in Coryanthes plants is rapid. All spp. investigated needed 60 to 70 days to produce about 600,000 seeds per capsule. Seldom are seedlings or young Coryanthes plants found without ants, indicating that normally Coryanthes seeds reach an ant nest and then begin to germinate. On the other hand, ant nests without Coryanthes are common. Quick seed-ripening, together with high growth rates and frequent observation of dying plants in abandoned ant nests, indicate a short life cycle in Coryanthes. In cultivation, within two years one can obtain plants of flowering size, and in nature similar conditions hold because of maintenance by ants. Brief life cycles in orchids are otherwise known only in twig epiphytes, but they produce less biomass and smaller flowers. (GG).

207. Embreea Dodson. Only one sp., E. rodigasiana (Claess. ex Cogn.) Dodson, Colombia to Ecuador.

208. Gongora Ruiz & Pav. Epiphytic herbs; leaves plicate, elliptic-lanceolate, shortly petiolate; inflorescence lateral, pendent (except G. erecta Whitten & D.E.Benn.); sepals free; petals partially adnate to column. Labellum fleshy and above column, bi- or tripartite. 57 spp., ranges from Mexico throughout Central America to Bolivia and SE Brazil (11, 5 endemics) and also in Trinidad and Tobago; 51 spp. in South America; Gongora spp. are epiphytes in wet and hot evergreen forests; the plants mostly occupy lower regions of their host trees, growing in deep shade, but they may also be found more exposed to direct sunlight. Frequently they grow in ant gardens, but this is not obligate as in Coryanthes spp. Although they often develop an ample root system nurtured by ants, they do not offer extra-floral nectar.

209. Houlletia Brongn. 9 spp. from over tropical South America; two species in Brazil, one endemic.

210. Lacaena Lindl. Only one sp., L. bicolor Lindl., Mexico to Colombia.

211. Lueckelia Jenny. One sp., L. breviloba (Summerh.) Jenny, Bolivia, N Brazil and Peru.

212. Lueddemannia Linden & Rchb.f. Two spp., NW Venezuela to Peru.

213. Kegeliella Mansf. 4 spp., one in Mexico and three in N South America, one in Brazil, none endemic.

214. Paphinia Lindl. 14 spp. from French Guiana to Venezuela, Colombia to Peru, N Brazil, one up to Trinidad; three species in Brazil, one endemic.

215. Polycycnis Rchb.f. 13 spp., Venezuela to Suriname, Colombia to Bolivia, Brazil (3, none endemics).

216. Schlimia Planch. & Linden. 7 spp., Venezuela and Bolivia.

217. Sievekingia Rchb.f. 11 spp., 9 in Colombia, Ecuador and Peru, one in Bolivia and one in Venezuela to French Guiana.

218. Soterosanthus F.Lehm. ex Jenny. Only one sp., S. shepheardii (Rolfe) Jenny, Colombia to Ecuador.

219. Stanhopea Fros ex Hook. Epiphytic or, less frequently, terrestrial herbs; leaves broadly elliptical with prominent veins, plicate, distinctly petiolate; inflorescence pendent, with some papery, clinging sheaths, thick and fleshy, 2–15-flowered; flowers facing downward, heavily perfumed; sepals and petals patent or reflexed (petals close to labellum only in S. tricornis Lindl.), dorsal sepal free, lateral sepals united at their base, dorsal sepal and petals smaller than lateral sepals. 71 spp. is similar to that of the subtribe, from Mexico through Central America to northern Argentina and S Brazil (7, 4 endemics), 43 in South America; subg. Stanhopeastrum range from the W Andes of Ecuador and Colombia to Guatemala; subg. Candida occur only in the Amazon basin; subg. Stanhopea extends over the entire range of the genus. There are two ‘hot spots’ of spp. richness, one in Mexico and another in Colombia. Large epiphytes in lower parts of the canopy from sea level up to 700m for members of S. subgen. Stanhopeastrum and S. subgen. Candida. spp. of S. subgen. Stanhopea prefer higher elevations, up to 1,800m. Habitats are usually wet, but in some cases there is a pronounced dry period, as in the pine and oak forests of Mexico. If there is enough light, they can also occur on moss-covered rocks or loose humus layers; infl orescences then occur on the substrate. It is questionable if those plants can reproduce because pollinators then cannot fall through the exit formed by lip and column.

220. Trevoria F.Lehm. 6 spp. from Colombia and Ecuador, one also in Bolivia.

221. Vasqueziella Dodson. Only one sp., V. boliviana Dodson, Peru to Bolivia.

10. EPIDENDROIDEAE ▸ TRIBE EPIDENDREAE (97/c. 7590) - six subtribes, only Agrostophyllinae (2/107; Seychelles to islands in W Pacific) absent in South America.

∎ SUBTRIBE CALYPSOINAE (13/78) ▸ outsiders Coelia (5; S Mexico, Central America, Caribbean), Dactylostalix (1; Japan), Ephippianthus (2; Korean Peninsula, Japan, Sakhalin), Changnienia (1; C and E China), Tipularia (7; Assam, Himalayas, Burma, Tibet, China (inc. Taiwan), Korean Peninsula, Japan, E U.S.A.), Calypso (1; temperate regions on the Northern Hemisphere), Yoania (4; NE India, Himalayas, China (inc. Taiwan), Vietnam, Japan), Aplectrum (1; Japan, North America), Cremastra (5; Himalayas, Tibet, China (inc. Taiwan), Korean Peninsula, Japan, Russian Far East), Danxiaorchis (1; Danxiashan in Guangdong), Oreorchis (16; W Himalayas to Japan and Taiwan (China)), Corallorhiza (11; temperate regions on the Northern Hemisphere, with their highest diversity in North America).

222. Govenia Lindl. Terrestrial herbs; leaves 1–3, oblanceolate to broadly oblanceolate, acute, plicate, arising from the base of the stem or corm; inflorescence lateral, subcapitate to elongate, manyflowered; flowers variously coloured; dorsal sepal concave, hooded over column, broadly lanceolate. Lateral sepals falcate, down-curved, acute; petals falcate, arching over column. 32 spp., ranging from Florida, Mexico, and Caribbean south to over South America (11, except Chile). spp. occur in montane, pine-oak, tropical, and second-growth forests and jungles. Dressler (1965) reported a plant with retained coralloid mycorhizome, as is seen in many groups in this tribe; only one sp. in Brazil, wider.

∎ SUBTRIBE BLETIINAE (4/60) ▸ outsiders Basiphyllaea (7; Florida, Caribbean), Hexalectris (10; southern U.S.A., Mexico).

223. Bletia Ruiz & Pav. Terrestrial herbs; roots few, fleshy; leaves plicate, basal, 1–4 (6) present or absent during flowering, the base tapering into the petiole; inflorescence lateral on the corm, one or rarely two, 2–50 flowers per inflorescence; flowers open or tubular, variously coloured, orange, yellow, greenish, magenta, lilac, or pale-pink to whitish. 42 spp., from northern Mexico, Florida, and Caribbean to NW Argentina, French Guiana to Venezuela; 2 spp. in South America, both wider; commonly grow among grasses on steep hillsides at low to middle elevations, pinelands, pine-oak forests or even in cloud forests, from sea level to 2,700m. A few spp. occur in tropical forests or xerophytic shrubby vegetation.

224. Chysis Lindl. 10 spp., 7 only from Mexico and Central America, one from Mexico to Bolivia, one only in Colombia and Venezuela, and C. guimaraensis Benelli & E. Pessoa endemic to Mato Grosso state, WC Brazil.

∎ SUBTRIBE PONERINAE (4/23) ▸ outsiders Helleriella (2; Central America), Ponera (2; Mexico, Central America).

225. Isochilus R. Br. Epiphytic or lithophytic, caespitose or creeping sympodial herbs; leaves numerous; inflorescence apical, sessile, raceme usually several-flowered; flowers tubular - campanulate, odourless; sepals and petals concolorous in tones of white, pink, lilac, magenta, or orange, labellum white below middle and coloured as other parts above. 11 spp. found in Mexico (center of diversity), Central America and the Antilles, with only the very wider I. linearis (Jacq.) R. Br. up to South America until SE Brazil and Argentina; evergreen and semi-deciduous tropical forests, mangrove swamp, pine-oak and various types of cloud forest from sea level to 2,900m elevation. Only one sp. in Brazil.

226. Nemaconia Knowles & Westc. 7 spp., Mexico to Panamá, and two up to tropical South America until Brazil (1, non endemic).

∎ SUBTRIBE LAELIINAE (38/c. 2340) ▸ outsiders Acrorchis (1; Central America), Alamania (1; Mexico), Amoana (2; Mexico), Artorima (1; Mexico), Barkeria (17; Mexico, Central America), Broughtonia (6; Greater Antilles, the Bahamas), Dinema (1; Mexico, Central America, Cuba, Jamaica), Domingoa (4; Central America, Cuba, Hispaniola), Hagsatera (2; Mexico), Meiracyllium (2; Mexico, Central America), Microepidendrum (1; Mexico), Psychilis (14; Caribbean), Quisqueya (4; Caribbean), Rhyncholaelia (2; Mexico, Central America), Tetramicra (13; Caribbean).

227. Adamantinia Van den Berg & C.N.Gonç. Epiphytic herbs, pseudobulb fusiform to ellipsoid, erect, green suffused of brownish purple; leaves one or seldom two, fleshy; inflorescence a raceme with up to eight sequential flowers, spathe lacking; flowers resupinate, 6–7 cm in diameter; sepals dark pink, membranaceous, lanceolate; petals the same colour and texture as the sepals, with the median portion larger than sepals, elliptical-ovate; labellum also dark pink. Only one sp., A. miltonioides van den Berg & C.N.Gonç., recorded only twice, only from a restricted mountain range in the Diamantina Range, in the central portion of Bahia State, NE Brazil; the first was a photographic record of it growing in the sun on a small tree of a rock cliff at about 1,400m; the second record was from where the plant was described, growing on an isolated tree in full sun at about 950m in a mountain pass; both locations were windy and wet montane habitats with a distinct dry season in the winter.

228. Arpophyllum Lex. 5 spp. from Mexico and Central America, one up to Venezuela, Colombia and Jamaica.

229. Brassavola R. Br. Epiphytic or rupicolous herbs, commonly erect, more rarely pendulous, caespitose or shortly creeping; leaves fleshy; inflorescence terminal, racemose with 2–7 flowers; flowers generally showy, the sepals and petals white or creamy white to pale green or yellow, with strong nocturnal fragrance. 20 spp. from Caribbean and Mexico into South America (18) as far south as northern Argentina and S Brazil. Many spp. occur in Brazil (10, 8 endemics) with fewer in Central America and the Andean countries.

The distribution of Brassavola appears to have two centres of diversity, with little sympatry, and only a few spp. have broad ranges. A group of spp. including B. cucullata (L.) R. Br. and all spp. of the Cuneilabia complex occur in a wide arc around the Caribbean and on the Pacific coast of Central America but are virtually absent from Caribbean, with the exception of the Jamaican endemic, B. subulifolia Lindl. and the widely distributed B. cucullata. A second centre occurs in South America, where at least 10 of the spp. grow in and around the Amazon Basin; Brazilian spp. tend to be Amazonian in distribution (B. angustata Lindl., B. martiana Lindl., and B. gardneri Cogn.) or occur in SE Brazil or the combined Paraguay–Bolivia–northern Argentina–S Brazil area (where up to seven spp. occur).

230. Cattleya Lindl. (inc. Brasilaelia, Cattleyella, Hoffmannseggella) Epiphytic herbs, more rarely rupicolous or terrestrial; leaf one, two, or rarely three, green, sometimes suffused with dark pink, terminal on pseudobulb; inflorescence generally a few-flowered raceme, more rarely multiflorous (up to 25 flowers); flowers resupinate, fragrant; sepals pink, yellow to golden, brown to dark red or green, sometimes spotted; petals of the same colour, texture, and shape as the sepals. 137 spp., 135 in South America, two only Central America; 119 spp. in Brazil, 111 endemics. 4 subgenera:

§ subg. Cattleya ▸ 91 spp. in three sections:

§ sect. Cattleya ▸ 17 spp., 7 in Colombia (5 endemic), one reaching to Peru, one to Panamá, another to Venezuela; one endemic to Peru and Venezuela each; two endemic to Brazil; 1 from Ecuador and Peru, 1 from Venezuela to Guyana; 3 more widely distributed.

§ sect. Crispae ▸ 72 spp., includes Microlaelia, Hoffmansegella, Sophronitis.

§ sect. Lawrenceanae ▸ 3 spp., plants from low elevation in the Amazon of northern Brazil and Venezuela, or from lowland, dry habitats in Venezuela.

§ subg. Cattleyella ▸ only one sp., C. araguaiensis Pabst, from S Pará to Tocantins state, in center Brazil.

§ subg. Intermediae ▸ 21 spp. all endemic to Brazil except by C. intermedia Graham ex Lindl. (Brazil to Uruguay and Paraguay), C. loddigesii Lindl. (Brazil to Argentina), C. nobilior Rchb.f. (Brazil to Bolivia) and C. violacea (Kunth) Rolfe (over South America).

§ subg. Maximae ▸ only one sp., C. maxima Lindl., from Venezuela to Peru.

231. Caularthron Raf. Epiphytic herbs of wet and seasonally dry forests, more rarely rupicolous; leaves at the upper nodes of the pseudobulb, generally 2–5, conduplicate; inflorescence terminal on pseudobulbs, a many-flowered raceme of up to 15 flowers, the peduncle covered with bracts, flowers opening successively; flowers up to 5 cm diameter; sepals and petals similar, elliptic to obovate, white or light pinkish. Two spp. occurring mainly in the Amazon and northern South America to coastal Ecuador and the Caribbean, both in Brazil. C. bilamellatum (Rchb.f.) R. E. Schultes also extends northward throughout Central America to the Mexican state of Chiapas, and provides one of the classical examples of myrmecophily in orchids, along with spp. of Myrmecophila.

232. Constantia Barb. Rodr. Epiphytic or rupicolous herbs; leaves two, green with white ridges, fleshy; flowers resupinate, up to 3 cm in diameter; sepals and petals white, pinkish or brownish red, fleshy. 6 spp., SE & S Brazil, all restrcited of only one state: 4 in Minas Gerais, in Rio de Janeiro and Santa Catarina states one each. C. cipoensis Porto & Brade occurs in Serra do Cipó, Minas Gerais, but as an specialized epiphyte on V. piresiana L. B. Smith and V. compacta Mart. ex. Schult. f.

233. Dimerandra Schltr. 7 spp. from South America, in French Guiana to Venezuela, Colombia to Peru, Brazil, two up to Mexico, Central America and Jamaica; two species in Brazil, none endemics.

234. Encyclia Hook.Epiphytic or rupicolous, rarely subterrestrial herbs; leaves generally two, more rarely one or three or four, green to purple, terminal on pseudobulb; inflorescence most commonly a panicle but sometimes a raceme, 1 to more than 50 flowers, spathe lacking, scape variable but most often erect or suberect, rarely subpendent, smooth or verruculose to warty, scape bracts generally present, commonly much shorter than the internodes; floral bracts usually inconspicuous; flowers resupinate, fragrant, inconspicuous to showy. 198 spp. occurring throughout the Neotropics, from Mexico through Central America, Caribbean, and the Andean range to S Brazil (61, 45 endemics), Paraguay, and northern Argentina; 107 spp. in South America; the southern limit of occurrence of Encyclia is around 29º S in Rio Grande do Sul State, Brazil, and the northern limit around 29º N in Sonora State, Mexico.

235. Epidendrum L. (inc. Takulumena) Epiphytic, lithophytic, or rarely terrestrial herbs, caespitose, sympodial or rarely monopodial; stem usually cane-like, simple or branching, rarely pseudobulbous; leaves one to numerous per stem, green to variously blotched, lined or tinged with purple; inflorescence apical, lateral, or rarely basal, single-flowered, racemose to paniculate; flowers mostly greenish yellow to white, sometimes brightly coloured yellow to red or pink to purple or black, often fragrant during a few hours of the day or night. 1,764 spp., the largest genera endemic to New World in Angiosperms; distributed from SE U.S.A. (North Carolina) to northern Argentina and Caribbean, Paraguay and Galapagos; 1,395 spp. in South America, 164 in Brazil, 90 endemics; highly diverse in Colombia (546) and Ecuador (577). Ubiquitous in the forests of tropical America, some even found in treeless areas such as sand dunes, scrub, and páramos. E. magnoliae Muhlenberg is the only epiphytic orchid that occurs in the warm temperate, deciduous forests of SE North America; many spp. are entirely terrestrials; plants of E. radicans Lindl. and E. secundum Jacq. are known to produce flowers when they are only one year old, and their seeds are among the largest in the orchid family.

236. Guarianthe Dressler & W.E.Higgins. 4 spp. from Mexico and Central America, one up to Venezuela, Colombia and Trinidad.

237. Homalopetalum Rolfe. 9 spp., mainly Mexico to Panamá; 4 spp. in South America, in Ecuador (1), Brazil (2, both endemics, from Rio de Janeiro to Santa Catarina states) and Venezuela (1).

238. Isabelia Barb. Rdr. Epiphytic, more rarely rupicolous herbs; leaf one, acicular or linear; inflorescence 1–2-flowered; sepals white, pink, or dark magenta, widely elliptical to ovate; petals the same colour as the sepals, narrower and oblong, or broader and elliptic. 3 spp., broadly distributed in E Brazil (Bahia from Rio Grande do Sul states), with I. virginalis also in Paraguay and northern Argentina. One natural hybrid, I. x pabstii, is known from Paraná State, Brazil, where both parents (I. pulchella (Kränzl.) van den Berg & M.W. Chase and I. violacea (Lindl.) van den Berg and M.W. Chase) occur. Grows on Podocarpus lambertii (Podocarpaceae), as an epiphyte in the Atlantic forest at sea level, drier forests in the Brazilian Shield, as epiphyte in small trees along streams in rocky grasslands (campos rupestres) and in higher montane areas, where it grows directly on rocks.

239. Jacquiniella Schltr. 12 spp., Mexico, Belize to Panamá, Caribbean, French Guiana to Venezuela, Colombia to Peru, N to SE Brazil (3, none endemics); 7 spp. in South America.

240. Laelia Lindl. (inc. Schomburgkia) Epiphytic or rupicolous, caespitose to scandent herbs; leaves 1–3, coriaceous to chartaceous, frequently fleshy and stiff, green and tinged with purple; inflorescence apical, racemose, the flowers arranged in a helical; flowers showy, resupinate or not, white, yellow, lilac-magenta, bronzy brown to chocolate-purple, sometimes with a varnished appearance, whitish mottled with purplish brown in L. lyonsii (Lindl.) L.O. Williams; a capsule of L. speciosa has 250,000–1,000,000 seeds; the circumscription of Laelia used here is different from that of other authors. It excludes the Brazilian Laelia spp. that have been transferred to Sophronitis. On the other hand, it includes those spp. previously included in Schomburgkia except for section Chauno-schomburgkia Schltr., now treated as the genus Myrmecophila. These changes make Laelia a genus of 32 spp. distributed more or less continuously from northern Mexico to Brazil (3, one endemic) and Bolivia, and the Caribbean (Cuba and Jamaica); 12 spp. in Mexico, 11 endemics; 14 spp. in South America.

241. Leptotes Lindl. Epiphytic herbs; stem abbreviated, cylindrical; leaf one, rarely two, terete, channeled; inflorescence a raceme up to 7 flowers, but generally 2–3, spathe lacking; sepals white to pinkish, oblong; petals the same colour as sepals and narrower. 8 spp., all from Brazil, although L. bicolor Lindl. extends to Paraguay and L. unicolor Barb.Rodr. to Argentina and Paraguay, mainly in the Atlantic forest of SE Brazil, from Bahia to Rio Grande do Sul State.

242. Loefgrenianthus Hoehne. Epiphytic herbs; rhizome pendent; stem abbreviated; leaf one, fleshy and flat; inflorescence uniflorous; flowers with sepals and petals white and yellowish lip. Only one sp., rare L. blanche-amesiae (Loefgr.) Hoehne, SE Brazil, from Rio de Janeiro to Paraná, in Podocarpus and Araucaria forests.

243. Myrmecophila Rolfe. 10 spp., 8 from Mexico to Panamá, one of then up to Venezuela and Colombia, another only in Venezuela and Colombia, and one restricted for W & S Caribbean; all spp. are myrmecophilous.

244. Nidema Britton & Millsp. Two spp., Mexico to Panamá, Caribbean, Guyana, Venezuela, Colombia to Bolivia and N Brazil (only one sp., N. ottonis (Reichb.) Britton & Millsp, in W Acre, and in Manaus region, Amazonas state).

245. Oestlundia W.E.Higgins. 4 spp., Mexico to Costa Rica, one up to Venezuela and Peru in South America, absent in Ecuador.

246. Orleanesia Barb. Rodr. Epiphytes or lithophytic herbs, several times on granite outcrops; leaves conduplicate; inflorescence erect or rarely arching, panicles or racemes; flowers usually green or yellowish green, variously tinged or spotted with purple, brown, maroon, or orange, with spreading perianth or the petals subparallel to the column. 9 spp. distributed in Colombia, Venezuela, the Guianas, Ecuador, Peru, Bolivia, and Brazil (6, 3 endemics).

247. Prosthechea Knowles & Westc. Epiphytic or lithophytic herbs. Pseudobulbs fusiform, often flattened; leaves 1–5; inflorescence racemose, often with a prominent spathe; flowers usually non-resupinate. 119 spp. from Florida (U.S.A.) to Caribbean, and Mexico southward through tropical South America to French Guiana and Paraguay, Brazil (35, 26 endemics). Members of this genus are epiphytic or lithophytic and prefer a moist habitat in woodlands including swamps and forests from sea level to 2,600m; 68 spp. in South America.

248. Pseudolaelia Porto & Brade. Epiphytic herbs, some specialized epiphytes on Velloziaceae, or terrestrial, mainly in rocky grasslands and also in large rocky outcrops; pseudobulb erect; leaves two or more; inflorescence a raceme of up to 15 flowers with sequential flowering, spathe absent; flowers resupinate, 2–5 cm in diameter; sepals and petals pink or yellowish. 17 spp. in E Brazil, with endemic spp. in Minas Gerais (5), Espírito Santo (3), Bahia (1), Rio de Janeiro (1), and remaining in Espírito Santo and adjacent Bahia and Minas Gerais states.

249. Pygmaeorchis Brade. Epiphytic herbs. Rhizome inconspicuous; leaves two, terminal on pseudobulbs, conduplicate, linear, oblong; inflorescence one-flowered, from half to the same length of the leaves, spathe absent; flowers resupinate, purple or greenish. Two spp., found at about 1,500m elevation in Serra dos Órgãos and Serra do Mar or as an epiphyte on Velloziaceae among rocky grasslands (campos rupestres) of Minas Gerais (as for spp. of Constantia and Pseudolaelia).

250. Scaphyglottis Piftzer. Epiphytic, caespitose herbs; leaves 1–3, apical on stems, papery, chartaceous to coriaceous or fleshy; inflorescence fasciculate or racemose, often successive; sepals similar, free, porrect or spreading; petals usually wider than sepals; labellum articulate to column foot or rarely firmly united, simple or trilobed, with or without calli. 86 spp., ranging from tropical Mexico and Caribbean to Peru, east up to French Guiana, Bolivia, and S Brazil (15, two endemics), 56 in South America; the greatest diversity is in Costa Rica and Panamá, where about c. 60 spp. occur.

∎ SUBTRIBE PLEUROTHALLIDINAE (44/c. 5,100) ▸ nine lineages, Dilolimis group (3/8, Cuba, Hispaniola, Jamaica, Puerto Rico) not occur in South America.

OCTOMERIA GROUP ▸ outsiders Atopoglossum (8, Cuba).

251. Brachionidium Lindl. Epiphytic, lithophytic, or terrestrial, erect or repent herbs. Rhizome sometimes branched, entirely covered by glabrous to scurfy sheaths. Stem abbreviated, erect, sheathed at base, lacking an annulus. Leaf coriaceous, elliptical, acute to obtuse, petiolate; inflorescence solitary-flowered; floral bract infundibular, acuminate; flower usually non-resupinate; sepals membranous. 82 spp. distributed throughout the Antilles (Cuba, Jamaica, Hispaniola, Puerto Rico, Martinique, Grenada, Guadeloupe, Dominica, St Kitts, Montserrat, St Lucia, and St Vincent) as well as in Guatemala and Costa Rica south to Venezuela and Guyana, S Brazil (6, one endemic), and Bolivia, Epiphytic, lithophytic, or terrestrial in humid, cloud, or elfin forests up to an astonishing 3,900m, often growing among bryophytes and leaf litter. Many spp. are common on embankments of road cuts; 64 spp. in South America.

252. Madisonia Lindl. (inc. Sansonia, Anathalis p.p., Specklinia p.p., Pabstiella p.p., Pleurothallis p.p.) Small, epiphytic or rupicolous, long-repent; rhizome slender, segmented between the ramicauls, nodes with ribbed, slightly ciliate sheaths. 10 spp., 7 endemics to E Brazil, and three collected in N Brazil, Bolivia, Peru, Colombia, Ecuador, French Guiana, Guyana, Jamaica, Suriname, Trinidad and Tobago and Venezuela.

253. Octomeria R. Br. Epiphytic or lithophytic, caespitose, scandent or repent herbs. Stem enclosed by one or more tubular, imbricating, glabrous sheaths, without an annulus. Leaf coriaceous; inflorescence fasciculate; floral bracts infundibular; flower resupinate. 158 spp., mainly epiphyte, lithophytes, or terrestrials from lowland scrub to cloud forests and elfin forests, up to 3,000m in elevation from Cuba and the Lesser Antilles and from Belize and Nicaragua south to Argentina and French Guiana; of the Brazilian spp. (97, 73 endemics), most occur in the states of Santa Catarina, Paraná, São Paulo, Rio de Janeiro, Minas Gerais, and Espírito Santo. The centers of diversity of the genus are in the Guianas, the Amazon basin and in particular, SE and S Brazil. Central American, Caribbean, Andean and the South American species which grow north and west of Amazonian Brazil are treated as a northern group, comprising c. 60 species; the remaining group of southern species occur in Brazil and neighboring countries to the south, with ca. 100 species.

RESTREPIA GROUP ▸ all genera in South America.

254. Barbosella Schltr. Epiphytic, lithophytic, or terrestrial, caespitose to long-repent herbs, sometimes forming dense, cushion-like colonies. Stem erect, enclosed by one or more sheaths, lacking an annulus. Leaf coriaceous, elliptical to suborbicular or semiterete to terete, acute to obtuse, sessile or petiolate; inflorescence solitary-flowered on an erect peduncle; floral bract oblique, acute, enclosing pedicel; flower resupinate; sepals membranous. Dorsal sepal linear to triangular, acute or caudate, free. Lateral sepals ovate to triangular, acute to obtuse, connate at base into a synsepal; petals elliptical to linear-ovate, acute. 19 spp., occurs in Caribbean (Hispaniola, Guadeloupe, and Martinique) and also from Guatemala to S Brazil (10, 8 endemics) and northern Argentina, east up to French Guiana. Epiphytic, lithophytic, or terrestrial from lowland forests (down to 100m in Paraná, Brazil) to wet elfin forests (up to 3,500m elevation in the Colombian Andes). A few spp. such as B. miersii form dense, moss-like colonies (Miller et al. 1994); 17 spp. in South America.

255. Chamelophyton Garay. Only one sp., C. kegelii (Rchb.f.) Garay, disjunct in Venezuela, Suriname and French Guiana.

256. Dresslerella Luer. 12 spp., Guatemala to Peru, 7 in South America, absent in Venezuela.

257. Echinosepala Pridgeon & M.W.Chase. 11 spp., only three in South America, Belize to Suriname, Jamaica, Colombia to Bolivia, N Brazil (2, both wider).

258. Myoxanthus Poepp. & Endl. (exc. Restrepiella p.p.) Epiphytic, lithophytic, or terrestrial, caespitose, repent or scandent herbs. Rhizome usually enclosed by scurfy sheaths. Stem erect, enclosed by several tubular, usually scurfy to pubescent sheaths, lacking an annulus. Leaf coriaceous, linear to elliptical, acute to obtuse, sessile or petiolate; inflorescence fasciculate, flowers produced successively or simultaneously; floral bracts tubular; flower resupinate. 50 spp. comprise this genus, which ranges from S Mexico (Chiapas) and Belize to Bolivia, Venezuela through Suriname, and Brazil (8, 5 endemics); some Myoxanthus are epiphytes, lithophytes, or terrestrials from lowland scrub to cloud forests and from 3 m (Brazil) to 3,200m (Colombia), one of the largest elevational ranges of any genus in Pleurothallidinae; 49 spp. in South America.

259. Pleurothallopsis Porto & Brade. 19 spp., 18 from Costa Rica to Venezuela, Colombia to Bolivia, and one, P. nemorosa (Barb. Rodr.) Porto & Brade, endemic from SE Brazil.

260. Restrepia Kunth. Epiphytic, caespitose herbs. Stem erect, enclosed by infundibular, imbricating, distichous, often speckled sheaths, without an annulus. Leaf coriaceous, elliptical to ovate, acute to obtuse, petiolate; inflorescence a fascicle of successive, solitary flowers appearing along the lower (abaxial) leaf surface; floral bracts tubular; flower resupinate. 58 spp. ranging from S Mexico (Chiapas) to Bolivia and Venezuela, 54 in South America; epiphytes from rain forests at 350m through cloud forest to paramo at 3,500m in elevation; highest diversity in the Andes of Colombia and Ecuador.

261. Restrepiella Garay & Dunst. (inc. Myoxanthus p.p.) 5 spp., one wider from S Florida, Mexico to Colombia, and Guatemala, Colombia, Brazil and Costa Rica one endemic each; Brazilian species R. ovatipetala (Chiron & Xim.Bols.) Rojas-Alv. & Karremans is endemic to Espírito Santo state.

ACIANTHERA GROUP ▸ two genera, Dondonia (1, Hispaniola) is a outsider.

262. Acianthera Scheidw. (inc. Kraenzlinella) Epiphytic or lithophytic (rarely terrestrial), caespitose or repent, rarely pendent herbs. Stem terete or laterally compressed with two or three edges, lacking an annulus, enclosed by one or more glabrous or pubescent sheaths. Leaf coriaceous, round to elliptical and obovate, acute to obtuse, sessile or petiolate, decurrent on stem in some spp; inflorescence racemose or single-flowered, from apex of stem or (rarely) directly from rhizome, emerging from a spathe in many spp., peduncles sometimes triquetrous; sepals generally fleshy and pubescent on outer surfaces. 225 spp. from the Antilles and Mexico to Argentina and Uruguay but is particularly well represented in Brazil (129, 104 endemics), 195 in South America. A. unguicallosa, endemic to Socorro Island in the Revillagigedo Archipelago, is probably the westernmost spp. of Pleurothallidinae. Generally humid-forest or cloud-forest epiphytes from 250 to 2,600m, frequently along the banks of rivers or streams. Some spp. occur in evergreen or dry forests on plains as in Bolivia. Others are common on trees in plantations of cacao, citrus, etc., and also on quartzite rocks in rocky grasslands (campos rupestres) of Brazil.

LEPANTHES GROUP

263. Anathallis Barb.Rodr. (exc. Lankesteriana p.p., Madisonia p.p.) Epiphytic, lithophytic, or terrestrial, caespitose or repent herbs. Leaf coriaceous, elliptical to obovate or oblanceolate, acute to obtuse, petiolate; inflorescence racemose (rarely solitary flowered), often arising from within a small spathe; floral bracts tubular or infundibular and oblique, acute to apiculate; sepals membranous or fleshy, often pubescent or papillose on adaxial (inner) surfaces. 133 spp. (118 in South America) from the Greater Antilles (Cuba, Jamaica, Hispaniola, Puerto Rico) and S Mexico to Brazil (98, 81 endemics), Bolivia, and Argentina, up eastwards to French Guiana, generally humid-forest or cloud-forest epiphytes from 200 to 2,700m (rarely to 3,000m). Some spp. such as A. rubens are terrestrial in leaf litter.

264. Draconanthes (Luer) Luer. Three spp., NW Venezuela to Bolivia.

265. Frondaria Luer. Only one sp., F. caulescens (Lindl.) Luer, Colombia to Bolivia.

266. Gravendeelia Bogarín & Karremans. (off Trinchosalpinx) Long-prolific, pendent habit, the few-flowered inflorescence, the cupped flower with extremely long sepals, the elongate lip with two central keels, the elongate column with a distinct foot, the incumbent anther and ventral, entire stigma. Only one spp., G. chamaelepanthes (Rchb.f.) Bogarín & Karremans, from Colombia to Bolivia.

267. Lankesteriana Karremans. (inc. Anathallis p.p.) 19 spp., S Mexico, through Central America, the Andes, and all the way down to Bolivia and Brazil; Costa Rica, Ecuador and Colombia contain the largest number of species.

268. Lepanthes Sw. (exc. Andinia p.p.) Epiphytic, lithophytic, or terrestrial, caespitose, repent or scandent herbs; stem erect or pendent, sometimes proliferating, leaf coriaceous; inflorescence racemose, fasciculate or solitary-flowered; flower resupinate or nonresupinate; sepals membranous; petals membranous or fleshy, triangular to transversely ovate to elongate or bilobed. 1,094 spp. ranging from Caribbean (Cuba, Jamaica, Puerto Rico, Hispaniola, Dominica, Guadeloupe, St Lucia, St Vincent) and S Mexico to Bolivia and N Brazil (6, Pará, Amazonas, Maranhão, Amapá, two endemics), 762 in South America, highly centered in Colombia (328) and Ecuador (367); mainly epiphytes, lithophytes, or terrestrials in moss and leaf-mould, principally in middle to high elevations, from wet to cloud and elfin forests with a high degree of endemism; many are twig epiphytes; vertical distribution ranges from 100 to over 3,300m.

269. Lepanthopsis (Cogn.) Ames. Epiphytic or lithophytic, caespitose or scandent herbs. Leaf coriaceous, suborbicular to elliptical, acute to obtuse, petiolate; inflorescence racemose, occasionally solitary-flowered; Flower resupinate or nonresupinate; sepals membranous, ovate, obtuse to long-acuminate, dorsal sepal free or only basally connate to lateral sepals, lateral sepals variously connate; petals membranous, suborbicular to elliptical, obtuse to acuminate. 48 spp., ranges from SW Florida, throughout the Greater Antilles, and S Mexico to Venezuela, Bolivia, and S Brazil (5, 3 endemics), a exact half in South America; many spp. are endemic to Hispaniola. Epiphytes, sometimes twig epiphytes, in moist lowland to cloud forests (occasionally semi-dry, scrub forests), generally between 1,000 and 2,000m elevation. However, L. melanantha grows at only 3 m in the Fahkahatchee Swamp in Florida, whereas L. peniculus (Schltr.) Garay and L. farrago (Luer & Hirtz) Luer are reported from up to 3,200m in Colombia and Ecuador, respectively.

270. Opilionanthe Karremans & Bogarín. (off Trichosalpinx). Two spp., endemics to Peru.

271. Pseudolepanthes (Luer). Archila. (off Trichosalpinx). 11 spp. from Colombia, 1-2 up to NW Ecuador.

272. Pendusalpinx Karremans & Mel.Fernández. (off Trichosalpinx). 6 spp. Colombia and Venezuela to Bolivia and Peru, also in French Guiana; they are not present in Central America, the Antilles and Brazil.

273. Stellamaris Mel.Fernández & Bogarín. (off Trichosalpinx) Only one sp., S. pergrata (Ames) Mel.Fernández & Bogarín, from Costa Rica to Colombia.

274. Trichosalpinx Luer. (exc. Gravendeelia, Opilionanthe, Pendusalpinx, Stellamaris, inc. Karma) Epiphytic, lithophytic, or terrestrial, caespitose, repent or pendent herbs. Stem erect or descending, often proliferating, enclosed by several sclerotic, tubular or infundibular (‘lepanthiform’) sheaths, ridges and margins pubescent, with an annulus. Leaf coriaceous, elliptical, oblanceolate to obovate, acute to obtuse, petiolate; inflorescence racemose or rarely solitary-flowered; floral bracts tubular or infundibular, often spiculate. 103 spp., Caribbean (Cuba, Jamaica, Hispaniola, Puerto Rico, Grenada, St Kitts, Martinique, Guadeloupe) and from Mexico to Bolivia, Venezuela and Trinidad and Tobago to French Guiana, and Brazil (12, 3 endemics); 80 spp. in South America. Epiphytes, lithophytes, or terrestrials in wet or cloud forests to elfin forests and subpáramo, from near sea level to 4,000m in elevation.

275. Zootrophion Luer. 27 spp., Nicaragua to Panamá, Caribbean, Colombia to Bolivia, Brazil (1, non endemic); 25 spp. in South America.

MASDEVALLIA GROUP ▸ 5 genera, all in South America.

276. Diodonopsis Pridgeon & M.W.Chase. 6 spp. from Colombia to Bolivia, two up to Costa Rica.

277. Dracula Luer. Epiphytic or terrestrial, caespitose to repent or ascending herbs. Stem enclosed by tubular sheaths, with an annulus. Leaf thincoriaceous, carinate or subplicate, linear to elliptical, acute or subacute, petiolate; inflorescence racemose, usually successive, or rarely solitary-flowered; floral bracts tubular; flower resupinate; sepals ovate, acuminate to obtuse, caudate, variously connate; petals cartilaginous, oblong, apex bivalvate, verrucose between valves. 133 spp. from Central America to Ecuador, one up to Peru, another up to state of Chiapas in S Mexico (1); most occur in the Andes of Colombia and Ecuador; as yet, no spp. of Dracula have been found in Bolivia, Brazil, or Venezuela; generally epiphytic, occasionally terrestrial, in mossy humus of humid, wet or cloud forests at elevations of 300–2,800m but mostly between 1,500 and 2,500m.

278. Masdevallia Ruiz & Pav. (inc. Spilotantha) Epiphytic, lithophytic, or terrestrial, caespitose to repent herbs. Stem erect or rarely descending, enclosed at base by imbricating sheaths, with an annulus. Leaf coriaceous, elliptical to obovate, acute to rounded, petiolate; inflorescence solitary-flowered or racemose, the peduncle terete or triquetrous; floral bracts tubular to cucullate; flower resupinate; sepals membranous or fleshy, showy, triangular to obovate, acuminate to obtuse, often caudate, free or variously connate; petals reduced, elliptical to oblong, acute to rounded or dentate, usually with a longitudinal callus and a rounded projection (‘tooth’) at the base.

616 spp. in New World, 583 in South America primarily Andean spp., ranging from S Mexico and Belize to Bolivia, Venezuela to French Guiana and Brazil (17, 6 endemics), highly centered in Colombia (155) and Ecuador (162). Masdevallia are epiphytes, lithophytes, or terrestrials, principally in cloud forests and subparamo up to 4,000m, but a few occur in warm humid forests as low as 10 m in elevation (e.g. M. lata Rchb.f.).

279. Porroglossum Schltr. Epiphytic or terrestrial, caespitose or repent herbs; stem erect, enclosed at base by imbricating, tubular sheaths, with an annulus; leaf coriaceous, sometimes rugose, elliptical to obovate, acute to obtuse, petiolate; inflorescence racemose, peduncle sometimes hirsute; floral bracts tubular; flower resupinate or non-resupinate; sepals connate up to middle, forming a cup, the apices usually contracted into caudae; dorsal sepal elliptical to obovate, rounded to acuminate; lateral sepals ovate, oblique, acute, often forming a mentum below column foot; petals ovate to oblong, obtuse to rounded. 48 spp., ranging from Colombia to Venezuela and Bolivia, mainly epiphytes or terrestrials in cloud forests at elevations of 1,000–3,200m.

280. Trisetella Luer. 26 spp., Costa Rica to French Guiana, Colombia to Bolivia, N Brazil (1, non endemic); 24 spp. in South America.

PHLOEOPHILA GROUP - a single genus.

281. Ophidion Luer. 16 spp., Panamá, Venezuela to Ecuador, disjunct in Bolivia.

282. Phloeophila Hoehne & Schltr. 6 spp., Belize to Panamá, Ecuador, Cuba, Brazil (2, none endemics), and Bolivia; 4 spp. in South America.

SPECKLINIA GROUP

283. Andinia (Luer) Luer. (inc. Pleurothallis p.p., Lepanthes p.p.) 77 spp., Venezuela (1, no endemic), Ecuador (29, 19 endemics), Colombia (26, 16 endemics), Peru (8, 7 endemics) and Bolivia (3, 1 endemic), where they are found in very humid forests, at elevations from 1,200 to 3,825 m, growing mostly under shady conditions.

284. Dryadella Luer. Epiphytic, caespitose herbs. Leaf coriaceous, linear-oblong or obovate, acute to obtuse, petiolate; inflorescence racemose or solitaryflowered; floral bracts imbricating; flower resupinate; sepals fleshy; species of this genus occur as epiphytes, sometimes in large populations, in cloud forests up to 2,600m but often at much lower elevations in humid forests. 57 spp. ranging from S Mexico, Belize, and Guatemala to Venezuela, Peru, and S Brazil (23, 20 endemics); 51 spp. in South America.

285. Muscarella Luer. (inc. Specklinia p.p.) 48 spp., tropical America, 29-48 in South America, only two in Brazil, endemism?

286. Platystele Schltr. Epiphytic, lithophytic, or terrestrial in wet, cloud, or subpáramo forests, some possibly two species, from Ecuador, among smallest orchidaceae worldwide, caespitose to repent herbs, from almost sea level in Belize (P. stenostachya (Rchb.f.) Garay) up to 3,200m in Ecuador (P. altarica Luer); stems erect, enclosed by imbricating, tubular to infundibular (but not sclerotic) sheaths, with an annulus; inflorescence racemose, sometimes prominently fractiflex; flower resupinate; sepals membranous, elliptic to ovate, rounded to long-acuminate, sometimes caudate, one-veined (or none), dorsal sepal free, lateral sepals free or basally connate only; petals membranous, filiform to ovate, rounded to acuminate. 117 spp. from Cuba and S Mexico (Veracruz, Oaxaca, Chiapas) to French Guiana and Trinidad and Tobago, S Brazil (6, two endemics), and Bolivia; 94 in South America.

287. Scaphosepalum Pfitzer. Epiphytic, lithophytic, or terrestrial, caespitose or repent herbs; stem erect, enclosed by two or three imbricating sheaths, with an annulus; leaf coriaceous; inflorescence racemose, secund or distichous, often flexuous; flower non-resupinate; sepals fleshy, glabrous to pubescent; petals ovate or ventricose, acute to obtuse; labellum oblong to pandurate; column semiterete, winged, hooded, with a foot; anther apical, incumbent; pollinia two; stigma entire. 50 spp., ranging in 4 well defined areas: Mexico to Panamá, NW Venezuela to S Ecuador, N Venezuela to N Guiana, and S Peru to C Bolivia, as epiphytic, lithophytic, or terrestrial in wet or cloud forests at elevations of 500–3,200m; only one species cited by Brazil, S. breve (Rchb. f.) Rolfe, described for Brazil only in 2008, only in small summit of Monte Roraima, northern Roraima state; 46 spp. in South America.

288. Specklinia Lindl. (exc. Muscarella p.p., Madisonia p.p.) Epiphytic, lithophytic, or terrestrial, caespitose or repent herbs in humid, wet, cloud, or elfin forests and from sea level to 3,000m in elevation; stem erect, enclosed by one or more tubular, imbricating sheaths, with an annulus. Leaf coriaceous, orbicular to obovate, acute to obtuse, usually petiolate; inflorescence racemose (rarely solitaryflowered); flower resupinate (rarely non-resupinate). 123 spp. ranging from Caribbean (both Greater and Lesser Antilles) and Mexico to Bolivia, Venezuela to French Guiana, and Brazil (23, 10 endemics); 87 spp. in South America.

289. Teagueia (Luer) Luer. 18 spp., 14 in Ecuador, 3 in Colombia and one in Peru.

PLEUROTHALLIS GROUP

290. Pabstiella Brieger & Senghas. (exc. Madisonia p.p.) Epiphytic, caespitose or creeping herbs. Stem erect, with 1–3 tubular or infundibular sheaths and an annulus. Leaf coriaceous, elliptical, acute to obtuse; inflorescence solitary-flowered or racemose; floral bracts infundibular, acute to acuminate; flower non-resupinate or resupinate. Dorsal sepal ovate, rounded to acuminate, shortly connate to lateral sepals. Lateral sepals connate (rarely free), ovate to tubular, obtuse to acuminate, often forming a conspicuous mentum with column foot; petals oblong to obovate, obtuse to acuminate. 124 spp., ranging from southern Colombia to Peru, Venezuela, Suriname, Brazil (120, 111 endemics), and Argentina (Misiones), two of then up to Central America. Species of Pabstiella are cloud-forest epiphytes from 1,000–2,500m, although P. yauaperyensis and P. determannii occur at lower elevations on trees in scrub or grasslands. 10 sections, mainly endemics to E Brazil forests.

291. Pleurothallis R. Br. (exc. Andinia p.p., Madisonia p.p.) Epiphytic, lithophytic, or terrestrial, caespitose to repent (rarely scandent) herbs. Stem erect (rarely pendent), sometimes laterally or apically compressed, enclosed by tubular sheaths, with or without a conspicuous annulus. Leaf coriaceous, linear to ovate (rarely semi-terete), acute to acuminate, base often cordate or rounded, sessile or petiolate, petiole sometimes twisted, sometimes decurrent on stem; inflorescence racemose, fasciculate or solitaryflowered, usually arising within a foliaceous spathe; floral bracts tubular or infundibular; flower resupinate or non-resupinate.

Pleurothallis today composes 716 spp., 605 in South America, even after more than 300 transfers to Acianthera, Anathallis, Andinia, Kraenzlinella, Pabstiella, Phloeophila, Specklinia, and Stelis; this vast genus ranges throughout the Neotropics, from the whole of Caribbean and Mexico to Bolivia, Venezuela, Colombia (204), Ecuador (354), Guyana, Suriname, and French Guiana and also to Brazil (17, 7 endemics), Paraguay and Argentina. P. ruscifolia (the type) is probably the most widely distributed spp. in the subtribe, extending virtually over the entire range of the genus; epiphytes, lithophytes, or terrestrials in a wide variety of habitats, from scrub near sea level to cloud forests and subpáramo at 3,000m and above.

292. Stelis Sw. Epiphytic, lithophytic, or terrestrial, caespitose to repent herbs; stem erect, enclosed by tubular or infundibular sheaths, with an annulus; leaf coriaceous; inflorescence racemose or (rarely) solitary-flowered, sometimes arising from a conspicuous spathe; flower usually resupinate. 1,210 spp. distributed from SW Florida (Collier County), both the Greater and Lesser Antilles, and S Mexico to Bolivia, Venezuela to French Guiana, and to S Brazil (51, 31 endemics); 1,072 in South America, 523 in Ecuador and 434 in Colombia; plants from swamps and warm, lowland scrub or broad-leaved forests close to sea level up to cloud forests and páramo (over 4,000m).

§ subg. Stelis ▸ 1,030 spp., over range of genus

§ subg. Crocodeilanthe ▸ 84 spp. in two groups:

o sect. Crocodeilanthe ▸ of 84 spp. of which 90% are found at high elevations in the Andes of Bolivia, Colombia, Ecuador, Peru and Venezuela, many local endemics. A few species are known from Costa Rica and Panamá, and a single species is reported from the Greater Antilles, another from the Lesser Antilles and yet another from Brazil.

o sect. Pseudostelis ▸ only one species, the common and widespread S. deregularis Barb. Rodr. which is found at mid elevations from Mexico to Brazil, through Central America.

§ subg. Physothallis ▸ 31 spp., mostly found at high elevations in the Andes of Bolivia, Colombia, Ecuador, Peru and Venezuela; three sections: sect. Acuminatae (27, that generally have rather narrow distributions in the Andean countries, especially Bolivia and Peru, a single species from Central America, and a couple are reported from Mexico, Guyana and Brazil; sect. Physothallis (3, endemics to Ecuador); sect. Rubens (a sole member, widely distributed from Colombia to Bolivia and Brazil).

§ subg. Niphantha ▸ includes two species of whitish, hirsute flowers. S. gelida (Lindl.) Pridgeon & M.W.Chase is a common species with the widest distribution in the genus, it is found from Florida and Mexico, through Central America and the Antilles, down to Peru, Bolivia and Brazil. S. pidax (Luer) Karremans is only known from Ecuador.

§ subg. Physosiphon ▸ six spp., distributed from Mexico and Guatemala, where the highest diversity is found, through Central America, and down to Bolivia; absent in Brazil.

§ subg. Dracontia ▸ 40 spp., c. 30 are endemic to Costa Rica and Panamá; a few species extend northwards into Mexico and Guatemala, a couple are known from the Antilles, and three make it downwards into the Andes; absent in Brazil.

§ subg. Uncifera ▸ 42 spp., from Mexico to Bolivia and Peru, they are especially diverse in Middle America and no records exist for the Antilles or Brazil.

§ subg. Condylago ▸ two spp., Panamá and Colombia.

§ subg. Umbralia ▸ 5 spp., mostly found from Costa Rica to Ecuador, with the highest diversity in Colombia; this subgenus includes the very variable S. imraei (Lindl.) Pridgeon & M.W.Chase, which is most like a species complex distributed from Costa Rica to Peru and Bolivia, the Guyanas, Brazil and the Lesser Antilles.

4.11 EPIDENDROIDEAE ▸ TRIBE VANDEAE (137/c. 2,300) – four subtribes, Adrorhizinae (3/33, Sri Lanka and Burma to E Queensland) and Aeridinae (83/c. 1325, tropical Africa to Asia and Pacific) do not occur in South America.

∎ SUBTRIBE POLYSTACHIINAE (3/c. 240) ▸ outsiders Hederorkis (2; Mascarene Islands, Seychelles), Imerinaea (1; Madagascar).

293. Polystachya Hook. Epiphytic, lithophytic or occasionally terrestrial herbs, absent from drier regions, particularly deserts and semi-deserts; leaves coriaceous to thin-textured; Inflorescence terminal, racemose, simple or branching, one- to many-flowered; peduncle glabrous or pubescent, often bearing one or more sterile bracts; flowers usually non-resupinate, pubescent or glabrous on outer surface. 230 spp., pantropical, centered in Africa and Madagascar, some in Asia; few spp. are found in the tropical Americas - 21 spp. Florida, Mexico, Belize to Panamá, Caribbean, Lesser Antilles, French Guiana to Venezuela, Colombia to Bolivia, over Brazil (12, 9 endemics), Argentina and Paraguay. 21 spp. in South America.

∎ SUBTRIBE ANGRAECINAE (47/c. 744) ▸ outsiders mainly in Africa and Madagascar up to Mascarenes Is. except by two up to Sri Lanka and Dendrophylax (14; Florida, Caribbean).

294. Campylocentrum Benth. Epiphytic herbs; roots photosynthetic in all spp., forming the main plant body in most leafless spp.; stem erect to pendent, sprawling in leafy spp., erect and less than 1 cm in most leafless spp; leaves articulate, reduced to nonphotosynthetic scales in most leafless spp.; inflorescence axillary, unbranched, usually congested with many open flowers; flowers white, occasionally tinged with orange; sepals and petals connivent, free, lanceolate, equal in size. 75 spp., from Florida and the Caribbean throughout most of Central and South America (66) as far south as Argentina. More than half of the spp. diversity is in Brazil (39, 13 endemics), from sea level to about 2,940m. Many are found in shaded sites with high humidity but may also be found in xeric habitats on twigs of shrubs or small trees or on limestone rocks. 5 sections.

§ sect. Campylocentrum ▸ 38 spp., over range of genus.

§ sect. Dendrophylopsis ▸ 13 spp., over range of genus; in this section, C. insulare C. E. Siquiera & E.M. Pessoa from Santa Catarina, S Brazil, is the world smallest species of orchids in flower size. This species was found growing as an epiphyte in remnants of the Atlantic Forest; it probably occurs in adjacent areas but due to the extremely reduced size of the plant, it is difficult to find.

§ sect. Laevigatum ▸ 15 spp., almost restricted to eastern South America, except for C. jamaicense from the Antilles.

§ sect. Pseudocampylocentrum ▸ only one sp., C. poeppigii (Rchb.f.) Rolfe, from Antilles, Central America and northern South America up to C Brazil.

§ sect. Teretifolium ▸ six spp., endemic to E Brazil.

4.12 EPIDENDROIDEAE ▸ TRIBE COLLABIEAE (20/443) ▸ outsiders Acanthephippium (13; India and SE Asia to China (inc. Taiwan), Japan, Malesia to New Guinea and islands in SW Pacific to Tonga), Ancistrochilus (2; tropical Africa), Ania (11; India, S China, SE Asia, Malesia to New Guinea, Solomon Islands, Queensland), Chrysoglossum (4; Himalayas, Tibet, Taiwan (China), SE Asia, Malesia to New Guinea and east to Fiji and Samoa), Collabium (14; Himalayas, S China, SE Asia, Malesia to New Guinea and Melanesia), Devogelia (1; Moluccas, New Guinea), Diglyphosa (3; Himalayas, SE Asia, Malesia to New Guinea), Eriodes (1; India to Vietnam), Hancockia (1; Yunnan, Vietnam, Japan, the Ryukyu Islands, Taiwan (China)), Ipsea (3; India, Sri Lanka, Thailand), Nephelaphyllum (11; Himalayas, S China, SE Asia, W Malesia to Borneo and Philippines), Pachystoma (2; Himalayas, S China (inc. Taiwan), SE Asia, Andaman and Nicobar Islands, and Malesia to New Guinea, New Caledonia and tropical Australia), Paraphaius (3; subtropical regions in E and SE Asia), Pilophyllum (1; Peninsular Thailand, Malesia to New Guinea and Solomon Islands), Plocoglottis (41; SE Asia, Malesia to New Guinea, Bismarck Archipelago and Solomon Islands), Risleya (1; Himalayas, W China), Spathoglottis (c 50; India, SE Asia, Malesia to New Guinea, Solomon Islands, NE Queensland, New Caledonia, Samoa, Niue), Tainia (23; India and Sri Lanka to China and Japan, SE Asia, Malesia to New Guinea and NE Queensland).

295. Calanthe R.Br. c. 270 spp., Tropical & Subtropical Old World to Pacific, only one in New World, C. calanthoides (A. Rich. & Galeotti) Hamer & Garay, from Mexico to Colombia, Caribbean.

LINEAGE 2: HYPOXIDIOIDS

ASTELIACEAE

§ FULLY CHLOROTROPHIC

Genera/spp. 3/37 Distribution Mascarene Islands, New Guinea, New Caledonia, Fiji, Samoa, the Society Islands, the Marquesas Islands, the Hawaiian Islands, temperate regions in southeastern Australia, Tasmania, New Zealand, Auckland, Campbell and Chatham Islands, southern Chile and Argentina, the Falkland Islands. Habit bisexual (Milligania), dioecious or gynodioecious, perennial herbs, rarely epiphytic.

SYSTEMATIC outsiders Neoastelia (1; NE New South Wales) and Milligania (5; Tasmania).

1. Astelia Banks & Sol. ex R.Br. Herbs, sometimes epiphytic and cushions. 26 spp., 25 in Mascarene Islands, New Guinea, New Caledonia, Fiji, Samoa, Society Islands, Marquesas Islands, Hawaii, temperate parts of SE Australia, Tasmania, New Zealand, Auckland, Campbell and Chatham Islands, and A. pumila (G. Forst.) R. Br. in S Chile, Argentina and the Falkland Islands.

HYPOXIDACEAE

§ FULLY CHLOROTROPHIC

Genera/spp. 5/161 Distribution Africa, Mascarene Islands, Seychelles, tropical and subtropical regions in Asia northwards to E Himalaya, SW China and Japan, New Guinea, Australia, Tasmania, New Zealand, E U.S.A. to Caribbean and South America southwards to Uruguay, with the largest diversity in South Africa. Habit usually bisexual (in Curculigo rarely unisexual), perennial herbs. Tuberous rhizome or corm.

SYSTEMATIC outsiders Empodium (9–10; South Africa, Swaziland, Lesotho), Pauridia (c 55; Namibia, South Africa, Lesotho, Australia, Tasmania, New Zealand), Molineria (7; tropical Asia), Hypoxidia (2; Seychelles); the largest genus of the family is Hypoxis L., but this has many taxonomic problems with delimitation of its spp., as they have a high degree of apomixy. It is found in tropical and subtropical areas of all continents; the family is found mainly in Africa, but Hypoxis and Curculigo have a few native spp. in the Neotropics.

1. Curculigo Gaertn. Medium-sized robust plants, which has a flower that appears at ground level and subterranean fruits, characterized by the umbel with a sessile or almost sessile flower. 20 spp., subcosmopitan, only one in New World, C. scorzonerifolia (Lam.) Baker, from Mexico, Central America, Caribbean, Trinidad & Tobago, Venezuela, Guianas, Colombia, Bolivia, Brazil and Cono Sur.

Ravenna created genus Heliacme Ravenna (based on C. scorzonerifolia Lam.) for the Neotropics, on the basis of a unilocular ovary and dry fruit (as compared with the rest of Curculigo Gaertn. which is otherwise African).

2. Hypoxis L. Small to medium-sized robust plants with aerial flower scape and dry fruit. 90 spp., pantropical, 18 in New World, 4 spp. in South America: H. decubens L. widely distributed, H. humilis Kunth known only rocky outcrops and grasslands in high mountains in Andes from Venezuela to Argentina, H. catamarcensis Brackett endemic to Argentina, and H. atlantica Funez, Hassemer & J.P.R. Ferreira, known only two sites (one in Bahia state, another in Santa Catarina), only in sea level sand soils near coast, in humid depressions.

LINEAGE 3: TECOPHILIDS

TECOPHILEACEAE

§ FULLY CHLOROTROPHIC.

Genera/spp. 9/23 Distribution tropical and S Africa, Madagascar, Chile, California. Habit bisexual, perennial herbs. Rhizome developed into a tuber or a corm (with basal innovation).

SYSTEMATIC outsiders Cyanastrum (3; Central and tropical E Africa), Kabuyea (1; Kenya, Tanzania); Odontostomum (1; California), Cyanella (7–8; Namibia, South Africa, with their highest diversity in W Cape) and Walleria (3; tropical and S Africa, Madagascar).

1. Conanthera Ruiz & Pav. 5 spp., all endemics to N Chile.

2. Tecophilaea Bertero & Colla. (inc. Distrepta). Two spp., T. cyanocrocus Leyb. endemic to Chile, and T. violiflora Bertero ex Colla from N Chile and S Peru.

3. Zephyra D. Don. Two spp., endemics to Chile.

LINEAGE 4: HIGH TEPALOIDES

IRIDACEAE

§ FULLY CHLOROTROPHIC

Genera/spp. 69/c. 1,750 Distribution tropical, subtropical and temperate regions in the Southern and Northern Hemispheres, with the largest diversity in S Africa, the eastern Mediterranean area, SW and E Asia, and parts of Central and South America. Habit bisexual, usually perennial (some spp. of Sisyrinchium annual) herbs (Klattia, Nivenia and Witsenia are suffruticose with a more or less lignified stem base). Usually with a rhizome (often tuberous) or a tunicated corm (with basal innovation; rarely bulb). Geosiris aphylla Baill. in Madagascar is an achlorophyllous mycotrophic holoparasite with scale-like membranous leaves. Several genera in South America.

The Iridaceae are a prominent family, forming characteristic components of several ecoregions such as fynbos. Genera such as Crocus and Iris are significant components of the floras of parts of Eurasia, and Iris also is well-represented in North America. Gladiolus and Moraea are large genera and major constituents of the flora of sub-Saharan and S Africa. Crocosmia crocosmiiflora (Lemoine ex Morren) N.E.Br. native in South Africa and naturalized in Brazil. Sisyrinchium, with more than 140 spp., is the most diversified Iridaceae genus in the Americas; the family occur in a variety of habitats from savannas, seasonally dry forests, to high altitude paramo and rocky grasslands (campos rupestres); many neotropical Iridaceae genera are adapted for seasonally dry habitats and some prefer truly xeric environments in which their underground systems may remain dormant for a long period; the lack of flowering material in herbaria (flowers often being destroyed in the process of herborization), creates a problem for floral studies as often only materials with fruits are available in herbaria; many species of Iridaceae are endemics with small populations and/or few occurences, which make them very vulnerable to extinction.

SYSTEMATIC subfamilies Isophysidoideae (1/1, Tasmania), Patersonioideae (1/25, Sumatra, Philippines, northern Borneo, New Guinea, Australia, Tasmania, New Caledonia), Geosiridoideae (1/3, Madagascar, Mayotte in the Comoros, Queensland), Aristeoideae (1/c 55, South Africa to Senegal and Ethiopia, Madagascar, with their highest diversity in the Cape Provinces), Nivenioideae (3/15, W Cape) and Crocoideae (31/1.260–1.305, Europe, Mediterranean, Africa, Madagascar, Socotra, SW Asia to W China) do not occur in South America.

Among the only South America subfamily, Iridoideae, tribes Diplarrheneae (1/2, New South Wales, Victoria, Tasmania) and Irideae (4/c. 550, temperate regions on the Northern Hemisphere, Mediterranean, southern tropical Africa to South Africa) do not occur in South America.

Key to genera of Neotropical Iridaceae

1. Plicate leaves, underground systems covered by membranaceous cataphylls - 2

2. Subequal tepals - 3

3.Stamens opposite to the styles ------------ Itysa

3. Stamens alternate to the styles - 4

4. Styles branches deeply divided into 2 or 3 arms ------------ Nemastylis

4. Styles undivided - 5

5.Stems terminated by a large foliaceous cauline bract subtending the inflorescence ------------ Eleutherine

5. Stems terminated by a linear cauline bract subtending the inflorescence ------------ Calydorea

2. Unequal tepals - 6

6. Panduriform anthers with broad connective ------------ Alophia

6. Linear anthers with slender connective - 7

7. Large foliaceous cauline bract subtending the inflorescence ------------ Cipura

7. Terminal inflorescence, not as described above - 8

8. Stamens and styles being held erect in a long tube at the center of the flower ------------ Tigridia

8. Styles deeply divided not forming a tube with the stamens at the center of the flower - 9

9. Inner tepals shorter than the outer but never less than half the size of them ------------ Cypella

9. Inner tepals smaller than half the size of the outer, to minute in size ------------ Mastigostyla

1. Linear or cylinndrical leaves, underground systems covered by fibrous cataphylls or absent, in this case in form of a rhizome - 10

10. Linear leaves, stamens alternating with the styles - 11

11. Tepals united at least basally and sometimes for a considerable distance, secretion of sugar nectar from the filaments ------------ Olsynium

11. Tepals free, secretion of oils from the filaments or rarely sugar nectar - 12

12. Ovary and capsules borne on slender pedicels, well exserted from the subtending bracts, tepals blue, lilac, white or yellow ------------ Sisyrinchium

12. Sessile or subsessile ovary and capsule not exserted from the subtending bracts - 13

13. Ovary hairy, tepals subequal, always blue ------------ Orthrosanthus

13. Ovary never hairy, tepals desequal, outer tepals smaller than the inner, flowers white, only one species with blue flowers ------------ Libertia

10. Linear or cylindrical leaves, stamens opposite the styles - 14

14; flowering stems winged, rarely inconspicuously so, leaf-like; often growing in forested habitats, never amongst rocks ------------ Neomarica

14; flowering stems wingless, rarely inconspicuously winged [only in Deluciris rupestris], never leaflike; often growing in open habitats, sometimes amongst rocks - 15

15; leaves cylindrical or plicate, sometimes absent during anthesis ------------ Pseudotrimezia

15; leaves dorsiventrally flattened, often present during anthesis - 16

16. First proximal node on the flowering stem ebracteate or with two bracts similar in length or one at most twice as long as the other - 17

17; flowers yellow to orange ------------ Pseudotrimezia

17; flowers blue to purple ------------ Deluciris

16. First proximal node on the flowering stem with one bract or rarely with two, one at least three times longer than the other - 18

18. Pseudo-midrib present, flowers yellow, style apex entire or with non-petaloid crests ------------ Trimezia

18. Pseudo-midrib absent, flowers blue to purple, style apex with petaloid crests ------------ Pseudiris

1.1 IRIDOIDEAE ▸ TRIBE SISYRINCHIEAE (6/238) - all genera occur in South America.

1. Libertia Sprengel. Small to medium evergreens rhizomatous perennials. 12 spp., one from Colombia to Bolivia, one in Araucania to Falklands also Juan Fernandez, 5 Chilean endemics, and 9 in New Guinea, Australia (one endemic), and New Zealand (7 endemics).

2. Olsynium Rafinesque. Leaves linear to terete; flowering stems aerial or subterranean, flowers pink, orange, white, yellow or red; seasonal or evergreen herbs. 14 spp., 13 in South America, from Venezuela to Falkland Islands (9 confined in Cono Sur), only O. acaule (Klatt) Goldblatt at Peru northwards, and O. douglasii (A. Dietr.) E.P. Bicknell in W North America.

3. Orthrosanthus Sweet. Perennials herbs. 10 spp., 5 in Australia, and 5 in over region from Central America and Mexico to Argentina and Bolivia, also Venezuela, none a national endemics, mainly mainly at high elevations.

4. Sisyrinchium L. Small to medium herbs; flowers usually yellow or blue to purple with yellow centre. 204 spp., S. acre H. Mann. in Hawaii, remaining 203 in South, Central and North America, with one sp. in Greenland, highly centered in S Brazil, Chile and Argentina; 136 in South America, 71 spp. in Brazil, 34 endemics. Nine sections:

§ sect. Cephalanthum ▸ c. 20 spp., Brazil, Bolivia and Cono Sur.

§ sect. Echthronema ▸ 7 spp., W North to Colombia, Peru and Bolivia, absent in Brazil.

§ sect. Hydastylus ▸ 6 spp., North America, Central America, western South America and Hawaii, absent in Brazil.

§ sect. Morphanthus ▸ two spp., North America to Argentina inc. Brazil.

§ sect. Rhizilineum ▸ 3 spp., widely in New World, inc. Brazil.

§ sect. Segetia ▸ 6 spp., Central America to Cono Sur, absent in Brazil.

§ sect. Sisyrinchium ▸ c. 60 spp., North and S South America, inc. Brazil.

§ sect. Spathirhachis ▸ 9 spp., Peru, Chile and Argentina.

§ sect. Trichoparcus ▸ 6 spp., Brazil, Bolivia and Cono Sur.

§ sect. Viperella ▸ c. 27 spp., Brazil, Bolivia and Cono Sur.

Sisyrinchium plants are usually found in open areas, grasslands, rocky formations, or wet environments and are sometimes found in ruderal habitats; some spp., such as S. micranthum Cav. are widely distributed in S Brazil, whereas others, such as S. rambonis R.C. Foster, occur only in environments with specific ranges of humidity and altitude.

5. Solenomelus Miers. Seasonal perennials with short rhizomes; leaves lanceolate to linear; flowers second, yellow or blue. Two spp. from Chile and Argentina.

6. Tapeinia Comm. ex. Juss. Small, evergreen cushions rhizomatous perennial; flowering stems axillary, with a single terminal flower, pale-pink, greenish externally. Only one sp., T. pumila (G. Forst.) Baill., from S Chile and Argentina.

1.2 IRIDOIDEAE ▸ TRIBE TRIMEZIEAE (5/79) - all genera occur in South America; high diverse in Brazil (69 spp., 67 endemics).

7. Deluciris A.Gil & Lovo. Underground stem vertical, corm-like with short internodes wrapped in a fibrous tunic; leaves, 1 to multiple, always present (non-hysteranthous), dorsiventrally flattened, flowers predominantly bluish to violet, always with various spots and transverse stripes; style apex with non-petaloid crests, stigma transverse. Two spp., along the Espinhaço Range in the Brazilian states of Bahia and Minas Gerais, mostly associated with rocky grasslands (campos rupestres), in open, dry meadowsor marshy environments.

8. Neomarica Sprague. Medium to large, occ. small, evergreen perennial with a creeping rhizome; stems erect to inclined; flowers crem, yellow to blue; compressed (leaf-like) flowering stem (peduncle + first bract) and a creeping or erect rhizome not covered by persistent fibrous tunic-like leaf bases. 31 spp., 29 endemics to forested coast of Brazil, N. candida (Hassl.) Sprague reaching into Paraguay, Argentina and Uruguay, and N. variegata (M.Martens & Galeotti) Henrich & Goldblatt, restricted to Mexico and Central America.

9. Pseudiris Chukr and A. Gil. Perennial herbs, rhizome corm-like, compact, erect, globose to cylindrical; leaves plane, linear-ensiforme to ensiforme, light-green; flowering stems simple or 2-3 branched, erect; flowers showy, lilac to blue, fugacious; capsules green, ovoid to oblong. Only one sp., P. speciosa Chukr & Gil, known only from the Diamantina Range on rocky grasslands (campos rupestres) in the municipalities of Lençois, Mucugê and road from Palmeiras to Capão, center Bahia state, Brazil.

10. Pseudotrimezia R. Foster. Small seasonal perennials herbs, with a bulb or bulb-like rootstock comprising a thick erect rhizome surrounded by leaf basis; leaves (one to many) cylindrical (shaped in a single), erect or strongly recurved; flowers yellow, very gold in one sp. 26 spp. mainly to the Espinhaço Range in Minas Gerais State, Brazil, from Serra de Grão-Mogol, in the north, to Serra do Cipó, in the south, mainly in the savannas of C Brazil (cerrado), the Espinhaço Range is covered by typical xeromophorphic vegetation, growing on rocky outcrops and shallow white sands, in altitudes above 900; many spp. restricted to single locations with particular environmental conditions, mostly in the Diamantina highs (central region of Espinhaço Range in Minas Gerais); a few species are more broadly distributed (e.g., P. cathartica (Klatt) Ravenna, P. juncifolia (Klatt) Lovo & A.Gil).

11. Trimezia Salisb. Ex Herb. Medium to large herbs. 19 spp., two only Mexico/Central America, 16 only South America (12 endemics to Brazil, and 4 restricted from E Venezuela to Guianas), and the widely distributed T. martinicensis (Jacq.) Herb. in Mexico, Central America, Caribbean, Trinidad & Tobago, Venezuela, Guianas, Colombia, Bolivia and Brazil.

1.3 IRIDOIDEAE ▸ TRIBE TIGRIDIEAE (20/191) - outsiders Nemastylis (6; SE U.S.A., Mexico, N Central America) and Cobana (1; Guatemala, Honduras).

12. Alophia Herb. Small to medium seasonal perennial with bulbs with dark brown papyraceous tunics. 6 spp., two endemics to Mexico, A. silvestris (Loes.) Goldblatt from Mexico to Nicaragua, A. drummondii (Graham) R.C. Foster from North America to Central America and scarce records in Brazil, Bolivia and Guianas, and two remaining endemic to C Brazil, in savannas and outcrops.

13. Calydorea Herb. Small seasonals perennials with bulbs with dark brown, papyraceous tunics; flowers pale to dark blue to purple with yellow marks. 19 spp., C. venezolensis (Ravenna) Goldblatt & Henrich in Venezuela; remaining 18 from Bolivia (3) to Chile and Uruguay, more than half from Brazil (11, 5 endemics) and adjacent Platine Basin; in S Brazil, the genus presents an interesting distribution, with typical spp. belonging to different biomes: C. basaltica Ravenna, C. crocoides Ravenna, and C. longipes Ravenna occur mainly in the Atlantic Forest biome, whereas C. alba Roitman & A. Castillo and its allies are endemic of the Pampa biome.

Goldblatt and Manning (2008) proposed that Calydorea includes provisionally Catila, Itysa, Lethia Ravenna, and Tamia Ravenna; the most recent study of Iridaceae phylogeny included only two Calydorea species, which did not group together in the phylogenetic trees (Goldblatt et al., 2008); considering the number of species involved in the analysis, phylogenetic relationships still need to be clarified.

14. Catila Ravenna. Only one sp., C. amabilis Ravenna, from C Rio Grande do Sul state, Brazil, up to Buenos Aires province in Argetina, and Uruguay.

15. Cipura Aubl. Cypella-like; flowers yellow to blue, white or red. 8 spp., two only in Colombia and Venezuela, one in Cuba, 4 in Brazil (two endemics), and C. campanulata Ravenna from Mexico to Venezuela.

16. Cypella Herbert. (exc. Larentia, inc. Phalocallis p.p.) Small or medium-sized plants perennials with 1–2 flowered spathes, often yellow or blue to purple and orange flowers with broadly clawed tepals. 39 spp., endemics in Mexico (2), Peru (1), Bolivia (2), C. linearis (Kunth) Baker scattered in over South America, and remaining 33 restricted from S Brazil (23, 8 endemics) to Argentina, Paraguay, Uruguay and Chile.

In the most recent and most complete phylogenetic overview of Iridaceae, Goldblatt et al. (2008) analyzed three species of Cypella and the genus appeared as polyphyletic, including Cipura Aubl., Nemastylis Nutt., Herbertia, Onira, and Calydorea; the relationships among Cypella, Phalocallis, and the monotypic genera Kelissa and Onira remain to be determined using molecular data (Goldblatt and Manning, 2008).

17. Eleutherine Herb. Small perennial seasonal; leaves few, plicate; flowering stem branches; flowers white. Two spp., E. bulbosa (Mill.) Urb. in Mexico, Caribbean, Guianas, Colombia, Ecuador, Peru, Bolivia, Brazil and Cono Sur, and E. latifolia (Standl. & L.O. Williams) Ravenna from Mexico, Central America, Bolivia and Cono Sur.

18. Ennealophus N.E.Br. Cypella-like, but flowers only shadeds of blue to purple with contrasting markings. 5 spp., 4 only in Andean region of Bolivia and Argentina, and E. foliosus (Kunth) Ravenna up to Amazonian rainforest of Brazil and Ecuador.

19. Gelasine Herbert (inc. Sphenostigma). Cypella-like but flowers blue to purple, often with white or darker margins. 8 spp., one endemic to Bolivia, remaining 7 from C & S Brazil (4 endemics), three of then also into NE Argentina, Uruguay.

20. Herbertia Sweet. Small seasonal perennial perennials with bulbs with dark brown papyraceous tunics; leaves few, ensiform to linear, flowers blue to purple with dark or white markings. 9 spp., H. tigridioides (Hicken) Goldblatt in Bolivia and Argentina and remaining 8 in Brazil, four up to adjacent Cono Sur (Argentina, Uruguay and Paraguay, one also in Texas, Louisiana) and four endemics.

21. Hesperoxiphion Sweet. Cypella-like. 5 spp., H. huilense Ravenna endemic to Colombia, and all remaining restricted of Peru except H. peruvianum (Baker) Baker up to Bolivia.

22. Kelissa Ravenna. Only one sp., K. brasiliensis (Baker) Ravenna, endemic to Rio Grande do Sul state, Brazil.

23. Larentia Klatt. (off Cypella). Three spp., two endemics to Mexico, and L. linearis (Kunth) Klatt) from Venezuela, Colombia, Bolivia and Paraguay to southern Brazil.

24. Lethia Ravenna. Only one sp., L. umbellata (Klatt) Ravenna, disjunct in Bolivia and E Brazil, in Minas Gerais, Bahia and Paraíba states.

25. Mastigostyla I.M. Johnston (inc. Cardenanthus). Bulbous caulescent or sub-acaulescent herbs, producing flowers with a short basal perianth tube tepals different in size and shape. 28 spp., M. cardenasii R.C. Foster in Peru and Bolivia, M. cyrtophylla I.M. Johnst. in Peru and Argentina, remaining endemic from Argentina (8), Bolivia (9) or Peru (9).

Based on molecular evidence, Goldblatt and Manning (2008) included Cardenanthus in Mastigostyla, a synonymy further supported by later molecular phylogenies (Chauveau et al. 2012), which also suggested that T. philippiana I. M. Johnston should be included in Mastigostyla.

26. Onira Ravenna. Only one sp., O. unguiculata (Baker) Ravenna, Rio Grande do Sul state in Brazil and NE Uruguay.

27. Phalocallis Herb. (exc. Cypella p.p.) Only one sp., P. coelestis (Lehm.) Ravenna, native from S Brazil to NE Argentina.

28. Tigridia Juss. Small to medium seasonal perennial with bulbs with darkery papery tunics. 50 spp., 39 from Mexico to El Salvador, two from Mexico scattered up Bolivia, 8 endemics to Peru and T. philippiana I.M. Johnst. restricted of N Chile.

ASPHODELACEAE

§ FULLY CHLOROTROPHIC

Genera/spp. 41/c. 895-910 Distribution Central, southern and E Europe, Africa, Madagascar, Mascarene Islands, W and C Asia eastwards to Japan and China, Australia, Tasmania, New Zealand, Polynesia (including Hawaii), Fiji, New Caledonia, New Guinea, Norfolk Island, northern Andes and the Guiana Shield. with the largest diversity in South Africa. Habit bisexual, usually perennial herbs (some spp. of Aloe and Kniphofia are pachycaul shrubs or trees), often evergreen, arborescent with aerial stem, or fruticose with subterranean stem. Stem thick, often tuberous, with leaves concentrated at apex. Rarely with bulb. Many representatives are leaf succulents; roots often somewhat succulent, sometimes contractile, sometimes swollen.

SYSTEMATIC subfamilies Xanthorroeoideae (1/30, Australia, Tasmania) and Asphodeloideae (18–19/790–830, S and E Europe, Mediterranean, Africa, Madagascar, Socotra, Arabian Peninsula, Mascarene Islands, W and C Asia, Australia, New Zealand, with their largest diversity in South Africa) do not occur in South America; among Hemerocalloideae, the only South American subfamily, tribes Hemerocallidoideae (2/20, Eurasia) and Johnsonioideae (7/c 40, South Africa, Madagascar, New Guinea, Australia, with their highest diversity in SW W Australia) do not occur in South America; among the unique South American tribe, Phormioideae, outsiders are Phormium (2; New Zealand incl. Stewart Island, Norfolk Island, Chatham Islands), Agrostocrinum (1; W Australia), Geitonoplesium (1; Philippines, Central and E Malesia to New Guinea, Queensland, New South Wales, Victoria, Norfolk Island, Lord Howe, New Caledonia, Fiji), Thelionema (3; Queensland, New South Wales, Victoria, Tasmania), Rhuacophila (1; Malesia, New Caledonia, Fiji), Herpolirion (1; New South Wales, Victoria, Tasmania, New Zealand), Stypandra (1; W Australia, South Australia, Queensland to Victoria, New Caledonia), Dianella (c 40; E Africa, Madagascar, E Asia to Japan, tropical Asia from India to Malesia, Australia, New Zealand, islands in SW Pacific, Hawaii, with their highest diversity in Australia).

1. Ecrremis Willd. Ex Bak. Rhizomatous or short-stemmed perennials to 1 m tall; leaves basal; leaves bases strongly keeled; inflorescence a loose corymbose panicle. Only one sp., E. coarctata (Ruiz & Pav) Baker, across highr elevations of South America including the middle and northern Andes, the Cordillera de la costa (Venezuela), and the summits of several tepuis in the Guiana Shield of Venezuela (in Amazonas and Bolivar states) and adjacent Brazil (only in Mount Neblina, at 2,500m, in a small field in Bacia do Gelo trail, Amazonas state)

2. Pasithea D. Don. Shortly rhizomatous perennial herbs to 15 cm tall; stem unbranched; leaves narrow-linear, mostly basal, bases sheating; inflorescence a racemose panicle terminating a leafless scape, bract short. Only one sp., P. caerulea (Ruiz & Pav) D. Don, from lower elevations and drier habitatas in N Chile and S Peru.

AMARYLLIDACEAE

§ FULLY CHLOROTROPHIC

Genera/species 71/700–820 Distribution tropical and subtropical regions in the N and S Hemispheres northwards to W Europe and eastwards to E Asia, with the largest diversity in South America, Mediterranean area and S Africa. Habit bisexual, perennial or biennial herbs, usually with a bulb surrounded by membranous scales (rarely a bulb-like corm; in Tulbaghia a tuberous rhizome) rich in polysaccharides (in Clivia, Cryptostephanus and Scadoxus corm or rhizome). Rarely epiphytic or aquatic. Often with a strong characteristic odour (‘onion smell’). New bulb developing in axile of uppermost leaf.

SYSTEMATIC three subfamilies, Agapanthoideae (1/7, S and SE South Africa from W Cape northwards to just south of Limpopo River, S Mozambique) not occur in South America.

1. SUBFAMILY ALLIOIDEAE (12/800) - tribes Allieae (1/750–780, temperate regions on the Northern Hemisphere, N, NE and S Africa, S Asia southwards to Sri Lanka, Mexico, with their highest diversity in Mediterranean area, Central Asia and SW North America) and Tulbaghieae (1/20, tropical and S Africa) do not occur in South America; the two others tribes are highly centered in South America.

1.1 ALLIOIDEAE ▸ TRIBE LEUCOCORYNEAE (7/152) - all genera occur in South America.

1. Atacamalium R. Pinto & Nic. García. Bulbous hebrs, flowers minute, greenishs. Only one sp., A. minutiflorum R. Pinto & Nic. García, coastal Atacama desert of N Chile.

2. Beauverdia Herter. Perennial herbs, bulb indeterminate; simple or prolific; leaves few; inflorescences 1-flowered, rarely bi-flowered; flowers bisexual, hypogynus, trimerous or tetramerous, actinomorphic, yellow or white. 4 spp., Argentina, Brazil (3, none endemics), and Uruguay, where they occur in open areas, usually in fields modified by livestock or human uses, in lowland or in gentle slopes with rocky and/or exposed soil.

B. hirtella (Kunth) Herter comprises two subspecies: subsp. hirtella and subsp. lorentzii, the former occurs in Uruguay and southern Rio Grande do Sul state, Brazil, while the second subspecies spreads in southeastern Corrientes and eastern Entre Ríos provinces, Argentina. Based on field surveys, study of nomenclatural types and other specimens, and review of literature, the new subspecies B. hirtella subsp. glabrata is here proposed, being endemic to rocky grasslands in south-central Rio Grande do Sul state, Brazil.

3. Ipheion Raf. Alliaceous odour absent or present. 5 spp. from Cono Sur (Argentina, Chile, Uruguay, Paraguay), I. recurvifolium (C.H.Wright) Traub, I. sessile (Phil.) Traub and I. uniflorum (Lindl.) Raf. up to S Brazil.

It is probable that I. setaceum (Baker) Traub (Argentina endemics) would be a previous name for Beauverdia vittata, or nevertheless, the fact that the two plants of the holotype are glabrous (Beauverdia vittata is generally papillose) and the locality of the type collection is probably wrong, Ipheion setaceum is here considered a doubtful species.

4. Latace Philippi. Two spp. from C & S Argentina and coast of Chile.

5. Leucocoryne Lindl. Herbs; leaves linear, scape slender; absent of alliaceous odor; inflorescence 1-12 flowers, mainly blusih. 46 spp. endemics to Chile.

6. Nothoscordum Kunth. Geophytes; alliaceous odour absent or present. 80 spp., Argentina, Uruguay, Paraguay, Chile, Bolivia, Peru, and Brazil (32, 26 endemics), only N. bivalve (L.) Britton, N. gracile (Aiton) Stearn and N. gramineum (Sims) P. Beauv. Peru northwards; N. bahiense Ravenna from Bahia state is a is a rare plant in Brazil, by Plantas Raras do Brasil’s book.

7. Tristagma Poeppig. Geophytes; inflorescence 1-8 flowers, alliaceous odour absent. 14 spp., Argentina and Chile.

1.2 ALLIOIDEAE ▸ TRIBE GILLESIEAE (4/23) - all genera occur in South America; the latest revision of Gilliesiinae (Escobar, 2012) accepted the genera Ancrumia Harv. ex Baker, Gethyum Phil., Gilliesia Lindl., Miersia Lindl., Schickendantziella Speg., Solaria Phil., Speea Loes. and Trichlora Baker, but POWO from Kew not recognizes Ancrumia, Gethyum, Solaria and Speea.

8. Gilliesia Lindl. (inc. Gethyum, Ancrumia, Solaria) Leaves usually 2, slender towards at base; scape slender, longer than the leaves; flowers very strongly zygomorphic; alliaceous odor recorded. 13 spp. from Chile, two up to and Argentina.

9. Miersia Lindl. (inc. Speea) Leaves 3-5 linear; inflorescence 3 – 7 flowers, zygomorphic; tepals 6, free, acuminate; corona of 6 narrow scapes. 10 spp., 9 endemics to from Chile and one endemic to Bolivia.

10. Schickendantziella Speg. Only one sp., S. trichosepala (Speg.) Speg., from S Bolivia to NW Argentina.

11. Trichlora Baker. Bulb narrow, tunicated; leaves 1-3, linear; scape slender; spathe bract 2, free from each other; inflorescence with 5 flowers. Two spp., endemics to Peru.

2. SUBFAMILY AMARYLLOIDEAE (c 80/870–900) - 12 clades, of them Cyrtantheae (1/c 50, tropical and S Africa, especially South Africa), Calostemmateae (2/7, Malesia, tropical and E Australia), Gethyllideae (6/c 84, tropical and S Africa (with their largest diversity in South Africa)), Lycorideae (2/25–32, temperate to subtropical E Asia to Iran, Central Asia and Burma), Pancratieae (2/28–29, SE Europe, Canary Islands, Mediterranean, W Africa, Namibia, SW and S Asia) and Narcisseae (7/c 72, Europe, Mediterranean, NW Africa, Crimea, the Caucasus, W Asia to Iran) do not occur in South America.

In the Neotropics, the family occurs from Canada through Central America and Caribbean to Chile and Argentina in South America; notable areas of diversity throughout this range include E Brazil, N & C Chile, and the central Andes of Ecuador and Peru; Hippeastrum Herb. is primarily found in the Andes and E Brazil, Hymenocallis occurs mostly in Mesoamerica, Clinanthus Herb. is largely endemic to Peru, and Zephyranthes Herb. is broadly distributed.

The greatest generic diversity is found in Peru. The Neotropical genera of Amaryllidaceae are chiefly adapted for seasonally dry habitats and some prefer truly xeric environments in which their bulbs may remain dormant for a period longer than they are in active growth (e.g., Leptochiton, Paramongaia, some Eucrosia); at the other extreme, species have colonized the understory of rain forests (Urceolina, Griffinia) and aquatic habitats (a number of Hymenocallis, Hippeastrum angustifolium Phil., Crinum). The family has also adapted to the high montane tropical climates of the Andes, with certain genera are primarily found at elevations in excess of 2,000m; Clinanthus humilis (Herb.) Meerow is found above 4,000m. This species has adapted to high elevations by retaining the scape (and developing fruit) inside the bulb until the seeds are ripe; only 11 spp. of Amaryllidaceae are epiphytic, three in southern Africa, three of Pamianthe in Ecuador, Peru and Bolivia one each, and 5 spp. of Hippeastrum endemic to S Brazil.

2.1 AMARYLLOIDEAE ▸ TRIBE AMARYLLIDEAE (11/c. 160) - outsiders Amaryllis (2; N and W Cape), Nerine (c 23; S Africa), Brunsvigia (c 20; S Africa), Crossyne (2; N and W Cape), Hessea (14; Namibia, N and W Cape), Namaquanula (2; Namibia, N Cape), Strumaria (28; S Africa, especially Namibia and SW South Africa), Boophone (2; E to S Africa), Ammocharis (7; tropical and S Africa).

12. Crinum L. Leaves often perennial, sometimes forming a pseudostem; flowers zygomorph to actinomorph; perigone tube long, cylindrical. 65 (-112) spp., 8 in New World, in terrestrial or shady places: one in Central America, two only in Caribbean, two Peruvian endemics, C. kunthianum M. Roem. from Colombia and Ecuador, and C. americanum L. and Cr. erubescens Aiton widely distributed, the former in Brazil.

2.2 AMARYLLOIDEAE ▸ TRIBE GRIFFINEAE (3/23) - tribe endemic to Brazil; this species, along with Lycoris Herb. (E & SE Asia, tribe Lycorideae), the only members of Amarylloideae to develop blue flowers.

13. Cearanthes Ravenna. Small herb, rounded dark-green leaves, blue flowers. Only one sp., C. fuscoviolacea Ravenna, known only four populations in forested environments of Ibiapaba Massif, NW Ceará and NE Piauí state, Brazil.

14. Griffinia Ker Gawl. Herbs, bulbous; inflorescence is 2–20-flowered, flowers with a true hypanthium in some species, feature unique among New World Amarylloideae, formed by the continuation of the perigonal tube over the ovary in some of the species, and white to bluish. 23 spp., two subgenera:

§ subg. Griffinia ▸ inflorescence is 4–20-flowered; flowers diurnal, unscented, lilac and/or white (only in G. alba K.D. Preuss & Meerow). 21 spp. from SE and NE Brazil (Bahia to Rio de Janeiro states, two of them, in Bahia and Pernambuco states, are rare plantd in Brazil, by Plantas Raras do Brasil’s book), mainly in forests of Atlantic Forest and its inland extensions.

§ subg. Hyline ▸ inflorescence is 2–3-flowered, flowers large, nocturnal, scented, and white. Two spp., G. nocturna Ravenna and G. gardneriana (Herb.) Ravenna, in seasonally dry regions, from NE Brazil, the former reaching into central savannas in W.C. region up to N Mato Grosso state.

15. Worsleya Traub. Large herbs with falcate leaves; roots thick, with velamen; bulb mostly exposed. Only one sp., W. rainieri (Hook.) Traub & Moldenke, very narrow endemic to 12 inselbergs of Petropolis and region, Rio de Janeiro state, Brazil, above 1,200m in high humidity forests.

2.3 AMARYLLOIDEAE ▸ TRIBE HIPPEASTREAE (13/185-200) - two subtribes, all genera of both occur in South America.

∎ SUBTRIBE TRAUBIINAE ▸ all genera occur in South America.

16. Paposoa Nic.García. (off Rhodophiala) Herbs usually over 10 cm tal; bulb ovoid; leaves annual, present while blooming, lorate, 30–60 cm long, 5–9 mm wide, flat and slightly fistulous; inflorescence pseudo-umbellate, 2–5-flowered, each flower subtended by a lanceolate bracteole; flowers slightly zygomorphic, nodding. Only one sp., P. laeta (Phil.) Nic.García, the single species within this new genus, inhabits fog oases or loma vegetation in the coastal desert of N Chile, between the Antofagasta and Atacama Regions.

17. Phycella Lindley. (inc. Placea, Famatina p.p.) Plants 20–100 cm tal; bulb ovoid; leaves annual, parallel venation, 30–60 cm long; inflorescence usually pseudo-umbellate, 1–9-flowered, each flower subtended by a linear-lanceolate bracteole; flowers slightly to strongly zygomorphic, various tones of red, dark pink, white, cream, or rarely yellow, either with red to magenta longitudinal stripes or the basal half green yellowish. ca. 13 spp. and mostly restricted to central Chile, in various habitat types such as desert scrub, sclerophyllous scrub and forest, high-Andean vegetation, including bogs and creeks, with a single population known from Neuquén, Argentina.

18. Rhodolirium Phil. (off Rhodophiala) Herbs usually 15 to 25 cm tal; bulb ovoid; leaves annual, absent or emerging while flowering, linear, parallel venation, 15–30 cm long, 2–5 mm wide, flat, canaliculate, sometimes pruinose, apex obtuse. Two spp., inhabits high-Andean to Patagonian steppe habitats from C & S Chile and adjacent Argentina.

19. Traubia Moldenke. Leaves linear, hysteranthous; cape hollow, spathe bracts two; flowers zygomorphic, funnelform, tube very short. Only one sp., T. modesta (Phil.) Ravenna, inhabits hilly areas close to the coast of central Chile, between the localities of Huentelauquén and Salamanca (Coquimbo Region), and Rapel (Metropolitan Region).

∎ SUBTRIBE HIPPEASTRINAE ▸ both genera occur in South America.

20. Hippeastrum Herb. (inc. Tocantinia) Terrestrial or epiphityc bulbous herbs, roots often with velamen; leaves usually anual; flowers 2-23, large, zygomorphic, funnelform. 89 spp. from South America, all countries of South America except Chile, centered in E Brazil (43, 34 endemics) and Andean of Peru to Argentina; two subgenera:

§ subg. Hippeastrum ▸ scape multi-flowered; flowers pedicellate; perigone variously colored, mostly without fragrance, in many vegetation types and habitats, mostly from Argentina to Colombia, very diverse in E Brazil in Atlantic Forest and savannas of C Brazil (cerrado) and central Andes of Peru and Bolivia; includes 5 epiphytes, inc. H. aulicum Herb.; H. calyptratum (Ker Gawl.) Herb., pollinated by bats, which grows as epiphytes attached to mossy trees, or grow as lithophytes on rock surfaces in bright light at around 1,200m elevation in Organ Mountains within the highly endangered eco-region of the Atlantic Forest of Brazil; ocasionally they are also found growing in shade; H. papilio (Ravenna) Van Scheepen, native at Santa Catarina state, S Brazil, and was considered extinct in its natural habitat until the 1990s, when Fred Meyer, observed it growing in tall trees in Rio Grande do Sul state; this species survives now as a population of approximately 50 plants within a 4-square-mile patch of Atlantic Forest habitat, fragmented by roads and drains.

§ subg. Tocantinia ▸ scape single-flowered; flower sessile; perigone white, sweet-scented nocturnally; three spp., plants in savannas of C Brazil (cerrado) in Bahia, Minas Gerais and Tocantins states, on sandy soils, within semideciduous dry forests.

21. Zephyranthes Herb. (inc. Famatina p.p., Rhodophiala p.p., Eithea, Sprekelia, Habranthus) Bulb generally small, single flowered, flower suberect, actinomorphic. 162 spp. of tropical and subtropical America, from U.S.A. to Argentina and Caribbean, 103 in South America; in Brazil (39, 25 endemics, one of them, from Bahia state, is a rare plant in Brazil, by Plantas Raras do Brasil’s book) occurs mainly in sourhern region; some species known as rain lilies because the flowers appear in through fall after the first rain; five subgenera:

§ subg. Habranthus ▸ three spp. found in northern Argentina, Uruguay, and S Brazil; in Argentina, Z. jamesonii (Baker) Nic.García & S.C.Arroyo inhabits sand dunes at sea level to high-Andean scrubs at 3,000m altitude range.

§ subg. Eithea ▸ two spp., Z. blumenavia (K.Koch & C.D.Bouché ex Carrière) Nic.García & Dutilh. inhabits the Brazilian Atlantic forest in the states of São Paulo, Paraná, and Santa Catarina, in very humid conditions within the forest, such as along rocky banks of small rivers or among large plants, while Z. lagopaivae (Campos-Rocha & Dutilh) Nic. García & Dutilh is known from only two semideciduous forest fragments within areas with well-defined seasonality in the state of São Paulo.

§ subg. Zephyranthes ▸ 139 spp., distributed widely in the Neotropics, but also occurs in subtropical to temperate regions, from central Argentina to the S U.S.A., also in the Caribbean islands; 80 in South America, 34 in Brazil, 23 endemics; Z. americana (Hoffmanns.) Herter, from NE Argentina, Uruguay and S Brazil, is a geophyte geocarpic herbs, unique among this quality in South American Amaryllidaceae.

§ subg. Neorhodophiala ▸ only the polymorphic Z. bifida (Herb.) Nic. García & Meerow from Uruguay, NE Argentina, S Brazil, and Paraguay (Itapúa); it is a component of the Pampas ecoregion, inhabiting open environments such as grasslands, savannahs, and between rocks in low hilly areas.

§ subg. Myostemma ▸ ca. 17 spp. from Chile and Argentina between, inhabiting a range of habitats including desert, sclerophyllous mediterranean-type scrub, deciduous nothofagid forests, and high-Andean scrubs, rarely in Patagonian steppe.

2.4 AMARYLLOIDEAE ▸ TRIBE EUSTEPHIEAE (3/20) - all genera occur in South America.

22. Chlidanthus Herb. Herbs; leaves with scabrous margins; flowers suberect to decliante; funnelform or funnelform to tubular; actinomorphic or slightly zygomorphic. 7 spp., Bolivia (4 endemics), Argentina (2 endemics) and C. fragrans Herb. from Peru and Bolivia.

23. Eustephia Cav. Herbs tuberous, with lowers declinate to pendent; perigone tubular, variously colored, but usually tipped green; tube short. 6 spp., 5 endemics to Peru and one to Bolivia.

24. Hieronymiella Pax. Herbs; flowers suberect to declinate; perigone funnelforme or tubular, white, yellow, rose or purple, often fragrant. 7spp., from Argentina (6, 5 endemics), H. marginata (Pax) Hunz. reaching to Bolivia, and H. bedelarii R. Lara & Huaylla endemic to Bolivia, chefly to high elevation.

2.5 AMARYLLOIDEAE ▸ TRIBE CLINANTHEAE (3/15) - all genera occur in South America.

25. Clinanthus Herb. (inc. Crocopsis) Stenomesson-like. 18 spp., mainly Peru (17, 15 endemics), one up to Bolivia, Argentina and Chile, C. incarnatus (Kunth) Meerow also in Ecuador and Bolivia, and one Bolivian endemic.

26. Pamianthe Stapf. Epiphytic; roots with velamen, leaves persistent; flower large and shortly pedicellate, fragrant; bulbs mostly aerial, prolonged above to a long neck. Three spp., one Peru and Bolivia, and Colombia and Ecuador one endemic each.

27. Paramongaia Velarde. Herbs; leaves anual, narrowly lorate, keeled, deeply glaucous, somewhat succulent. Two spp. from W Andes from Peru (both) and Bolivia; this plant has very short season of growth.

2.6 AMARYLLOIDEAE ▸ TRIBE HYMENOCALLIDEAE (11/159) - all genera occur in South America.

28. Eucrosia Ker Gaul. Herbs; leaves hysteranthous; flowers without fragrance, petiolate; perigone showy, zygomorphic. 7 spp., W Ecuador (4 endemics) and NW Peru (two endemics), E. bicolor Ker Gawl. in both countries.

29. Hymenocallis Salisb. Herbs, leaves annual or persistent, white fragrant flowers, crateriform, actinomorphic, fragrant, white. 61 spp., mainly center Mexico (33, 31 endemics), from Caribbean to Peru and Amazon rainforest of Brazil, also in coastal mangroves; only 10 up to South America, national endemics in Venezuela (3), Peru (2) and Brazil (1, H. schizostephana Worsley), one Venezuela to Caribbean, H. pedalis Herb. in northern Andes, H. littoralis (Jacq.) Salisb. and H.tubiflora Salisb. in coasts from Mexico to Peru and N Brazil, the first common in mangroves, the two lasts occurs in Amazonian rainforest.

30. Ismene Salisb. Herbs; scape ancipitous; leaves annual, forming a long pseudostem, appearing before the flowers, sometimes succulent. 12 spp., all from Peru, one up to Ecuador.

31. Leptochiton Sealy. Herbs, ephemeral geophytes; leaves anual, linear, lax, fragile, keeled, appearing before the flowers; flowers large, fragrant, sessily, yellow or white. Two spp. inhabiting seasonally very dry lowlands dioecious forests from SW Ecuador and NW Peru (both).

32. Mathieua Klotzch. Herbs; leaves long petiolate, lamina ovate. Only one sp., M. galanthoides Klotzsch, from a fragmantary type, the remains of a collection from dry forests of Piura, Peru, presumed extinct.

33. Phaedranassa Herb. Herbs, leaves mostly hysteranthous; flowers mostly tubular. 10 spp., Central America to Peru, mainly in Ecuador (8, 6 endemics); large but local populations, along roadsides.

34. Plagiolirion Baker. Herbs; leaves persistente; flowers white, non fragrant, 10-40, perigone zygomorphic, small. Only one sp., P. horsmannii Baker, endemic to the slopes of the Cauca Valley in Andean Colombia.

35. Pyrolirion Herb. (inc. Leucothauma) Herbs, leaves annual, linear or linear to lanceolate; scape hollow, single flowered, sometimes retained inside in the bulb. 8 spp. in Peru (6 endemics) and Bolivia (two endemics).

36. Rauhia Traub. Herbis with bulbs large, leaves annual, shortly or widely petiolate, succulent, unique among this family in New World. 5 spp. from seasonaly dry, open woods or rock slopes of the middle Maranon and Utcumbamba Valleys in N Peru.

37. Stenomesson Herb. Herbs, leaves annual; flowers actinomorphic, funnelform-tubular, tubular, campanulate or ventricose. 28 spp., almost all in Peru (23 endemics), three up to Bolivia, one up to Colombia, and one a Ecuador endemic.

38. Urceolina Reichb. (inc. Eucharis, Caliphruria) Herbs with leaves hysteranthous; flowers yellow to orange tipped green, not fragrant; perigone actinomorphic, urceolate. 28 spp., U. bouchei (Woodson & P.Allen) Traub. in Guatemala, Panamá and Costa Rica, and remaining 27 from Colombia to Bolivia and N Brazil (3, U. ulei (Kraenzl.) Traub., U. cyaneosperma (Meerow) Christenh. & Byng. and U. castelnaeana (Baill.) Christenh. & Byng., none endemics), absent in Venezuela, mainly are endemic to western Amazon rainforest and adjacent lower slopes of E Andes, mainly in Peru (10 endemics) and Colombia (7 endemics).

ASPARAGACEAE

§ FULLY CHLOROTROPHIC

Genera/spp. 121/2,500 Distribution worldwide except Arctic Habit bisexual, perennial herbs. Rhizome developed into a tuber or a corm (with basal innovation).

SYSTEMATIC seven subfamilies, 3 in South America and 4 outsiders: Aphyllanthoideae (= Aphyllanthoideae, 1/1, Mediterranean Basin), Asparagoideae (= Asparagaceae s.s., 2/165–195(–300?), Eurasia, Africa, Australia, Mexico), Brodiaeoideae (= Themidaceae, 12/62, from Canada to Guatemala) and Nolinioideae (= Ruscaceae, 24/740–765, temperate to tropical regions on the Northern Hemisphere south to W Malesia and Central America, with their largest diversity in E and SE Asia, North America, Mexico and South Africa). 668 spp. in New World.

1. SUBFAMILY LOMANDROIDEAE (14/175–180) ▸ three tribes, Lomandreae (5/65–70, New Guinea, Australia, New Caledonia) does not occur in South America.

1.1 LOMANDROIDEAE ▸TRIBE LAXMANNIOIDEAE (8/c 95) - outsiders Arthropodium (14; Madagascar, Australia, New Caledonia, New Zealand), Murchisonia (2; Australia), Thysanotus (50; China to New Guinea, Australia), Eustrephus (1; New Guinea, Queensland, New South Wales, Victoria, New Caledonia), Laxmannia (14; Australia), Sowerbaea (5; Australia), Chamaescilla (4; Australia, Victoria, Tasmania).

1. Trichopetalum Lindl. Herbs with erect rhizomes and somewhat fleshy roots; flowers in loose racemes panicles. Two spp. from SW Argentina (in Neuquen) and center Chile, one endemic each.

1.2 LOMANDROIDEAE ▸TRIBE CORDYLINEAE (1/17) - a single genus.

2. Cordyline Comm. Ex R. Br. Woody shrubs or trees to 10 m with thick rhizomes thick, somewhat fleshy roots. 24 spp., Mascarenes, to Pacífico, South America, and C. sellowiana Kunth from Bolivia, S & SE Brazil, Paraguay, N Argentina and Uruguay.

2. SUBFAMILY SCILLOIDEAE (38/805–845) ▸ tribes Urgineoideae (2/c 60, Mediterranean, Africa, Madagascar, Arabian Peninsula to Sri Lanka and Burma), Ornithogaloideae (1/270–300, Europe, Mediterranean to the Caucasus, subtropical and S Africa, Madagascar, S Arabian Peninsula, Socotra, SW Asia to India and Sri Lanka, with their largest diversity in South Africa) and Hyacinthoideae (34/470–480, Europe to SW Asia, North Africa, tropical and S Africa, Madagascar, Arabian Peninsula, S India, Sri Lanka, E Asia) do not occur in South America; the single tribe in continent, Oziroideae, is monotypic and endemic.

3. Oziroe Raf. Herbs, bulbous, bulb leaves imbricate, cauline leaves few, thick and gloved, synanthous, few white flowers. 5 spp. in W South America, Peru to Paraguay and Chile, Bolivia, Argentina, with O. argentinense (Lillo & Hauman) Speta collected recently in S Mato Grosso do Sul state, Brazil.

3. SUBFAMILY AGAVOIDEAE (23/637) ▸ three tribes, one in South America; the remaining two occur only in China, Korea (1/1, Anemanrrhena) and S Africa (1/1, Behnia).

3.1 AGAVOIDEAE ▸TRIBE AGAVEAE (10–13/350–360) - outsiders Hesperocallis (1; California, Arizona); Hosta (23; China, Korean Peninsula, Japan, Russian Far East); Hesperoyucca (2; Arizona; California, Baja California), Schoenolirion (3; Texas, SE U.S.A.), Hesperaloe (8; SW U.S.A., NW Mexico); Hastingsia (4; California and SW Oregon), Camassia (6; Canada, U.S.A.), Chlorogalum (5; California, Baja California); Yucca (c 50; C and W U.S.A., California, Mexico, Central America, Caribbean), Beschorneria (8; C and S Mexico, N Central America).

4. Agave L. Small to gigant; acaulescent or with short trunk, perennial, poly-or monocarpic. 270 spp. from Caribbean, SW U.S.A. to N Venezuela and Ecuador, mainly in montane areas; 9 spp. in South America, mainly in N Andes

5. Furcraea Vent. Rosettes, massive, acaulescent or with trunk to 6 m tall; leaves stiff or flexible. 27 spp. from C Mexico to Bolivia and French Guiana (9 in South America, some endemic to Andean Colombia), with F. foetida (L.) Haw. up to N Amapá state, Brazil.

3.2 AGAVOIDEAE ▸TRIBE ANTHERICEAE (8/280–290) - outsiders Anthericum s.s. (c 65; Europe, North and E Africa to Tanzania, SW Asia), Paradisea (2; mountains in southern Europe), Chlorophytum (190–200; Africa, Madagascar, India to northern and E Australia), Leucocrinum (1; SW U.S.A.) and Eremocrinum (1; Utah, N Arizona).

6. Anthericum L. Herbs. 65 spp., predominantly in Mediterranean and southern Central Europe, Middel East, N and E African south to Tanzania; possibly in South America. 9 spp. in South America, all national endemics in Peru (5), Bolivia (2) and Argentina (2).

Generic limits of Anthericum relative to the South American species are still incertain and the subject of ongoing research; in particular, the generic delimitation of the Old World Anthericum species in Kativu and Nordal (1993) and Nordal and Thulin (1993) excludes several of the New World species as defined by Ravenna (1987, 1988); at least one (and possibly up to 15) of these S American species represents an undescribed genus with afffinities to Echeandia (R.W. Cruden and P. Ravenna, pers. comm.).

7. Diamena Ravenna. Herbs with a short vertical rhizome and tuberous roots. Only one sp., D. stenantha (Ravenna) Ravenna, endemic to upper Cerro de las Cabras near Trujillo, Peru, and may be extinct in wild.

8. Diora Ravenna. Herbs with a short vertical rhizome and tuberous roots. Only one sp., D. cajamarcaensis (Poelln.) Ravenna, Cajamarca and Junin regions in Peru.

9. Echeandia Ortega. Herbs with short rhizomes and tuberous roots; flowers in racemes or thyrses. 84 spp. from Texas to Argentina; 8 spp. in South America, 5 national endemics in Colombia (1), Venezuela (1), Peru (2) and Ecuador (1), E. macrophylla Rose ex Weath. and E. pittieri Cruden scattered from México to Colombia and Venezuela, and E. ciliata (Kunth) Cruden from Mexico to Venezuela and Cono Sur.

10. Hagenbachia Ness & Mart. Herbs with a horizontal rhizome and thick, fleshy but non tuberous roots; flowers on loose racemes and panicles. 6 spp., three from Costa Rica to Ecuador, H. matogrossensis (Poelln.) Ravenna from Brazil, Paraguay and Bolivia, H. hassleriana (Baker) Cruden from Bolivia and Paraguay, and H. brasiliensis Nees & Mart. from Bolivia and E Brazil, in below 400m in dry seasonal scrubland of NE Brazil (caatinga), evergreen forest, and disturbed sites.

11. Trihesperus Herb. Two disjunct spp., T. latifolius (Kunth) Herb. from Colombia and Ecuador, T. glaucus (Ruiz & Pav.) Herb. from Peru to Bolivia, Argentina.

3.3 AGAVOIDEAE ▸TRIBE HERRERIEAE (2/9) - outsider Herreriopsis (1; Madagascar).

12. Clara Kunth. Stemless rosette herbs (non-scadent); leaves filiforme to linear; inflorescences in raceme simple or double composed. Three spp. of grasslands, all in Brazil, with C. gracilis R. C. Lopes & Andreata a narrow endemic, recorded in the grasslands of the ‘campanha gaucha’, in Rio Grande do Sul, Uruguay, Paraguay and NE Argentina (only in border with Paraguay).

13. Herreria Ruiz and Pavon. Climber herbs; leaves clustered in short lateral shots, ovate-lanceolate to linear-lanceolate; inflorescence raceme or paniculate. 9 spp., 8 in Brazil (4 endemics), two up to Bolivia, one up to Bolivia, Argentina, Paraguay and Uruguay, and another up Argentina, Paraguay and Uruguay; H. stellata Ruiz & Pav. is a Chilean endemic; Atlantic sandy coastal shrublands (restingas), dry xeric seasonal of NE Brazil (caatinga), and forest formations.

16. ARECALES

FAMILIES ABSENTS IN SOUTH AMERICA: DASYPOGONACEAE (4/20).

ARECACEAE

§ FULLY CHLOROTROPHIC

Genera/species 182/c. 2,440 Distribution chiefly pantropical, some species in subtropical regions and a few species in warm-temperate areas. Habit usually monoecious, polygamomonoecious or dioecious; male and female flowers isomorphic or heteromorphic (sometimes bisexual); evergreen, woody (trees, shrubs or lianas), usually with monopodial growth (rarely branched). Roots or leaflets sometimes modified into spines (spine roots). Aerial roots (stilt roots, prop roots or pneumatophores) present in many species. Nypa consists of mangrove plants. Hyphaene has dichotomous branching.

Palms are among the best known and most extensively cultivated plant families. They have been important to humans throughout much of history. Many common products and foods are derived from palms, and palms are also widely used in landscaping for their exotic appearance, making them one of the most economically important plants. In many historical cultures, palms were symbols for such ideas as victory, peace, and fertility. Today, palms remain a popular symbol for the tropics and vacations.

MORPHOLOGY Whether as shrubs, trees, or vines, palms have two methods of growth: solitary or clustered. Palms have large, evergreen leaves that are either palmately ('fan-leaved') or pinnately ('feather-leaved') compound and spirally arranged at the top of the stem. The inflorescence is a panicle or spike surrounded by one or more bracts or spathes that become woody at maturity. Most palms are distinguished by their large, compound, evergreen leaves arranged at the top of an unbranched stem. However, many palms are exceptions, and in fact exhibit an enormous diversity in physical characteristics. As well as being morphologically diverse, palms also inhabit nearly every type of habitat within their range, from rainforests to deserts. The morphological diversity of palms is greater than that of any other monocotyledoneous family; indeed, the may represent one the most diverse of seed plants as a whole.

The raffia palm (Raphia regalis Becc.) of tropical Africa has huge pinnate leaves up to 80 feet (24 m) long. The leaves of the Amazonian palm (Manicaria saccifera Burret.) are nearly 30 feet (8 m) long, and have been listed by some authors as the longest undivided leaf of any plant. However, according to Chuck Hubbuch of Fairchild Tropical Garden, Coral Gables, Florida, the leaf is typically divided shallowly at the tip and is not truly entire. The golf ball-sized fruits of Manicaria palms, called ‘sea coconuts’, commonly wash ashore on beaches throughout the Caribbean and southern Florida. There are two additional palm candidates for the record of longest undivided leaf: Marojejya darianii J.Dransf. & N.W.Uhl, a palm native to Madagascar with a leaf up to 5 meters in length that is divided only once at the tip; and joey palm (Johannesteijsmannia altifrons (Rchb.f. & Zoll.) H.E.Moore), a palm native to Thailand with a leaf up to 4 meters long that is completely undivided.

Arecaceae are notable among monocots for their height and for the size of their seeds, leaves, and inflorescences; The coco de mer (Lodoicea maldivica (J.F.Gmel.) Pers.) has the largest seed of any plant, 40–50 cm in diameter and weighing 15–30 kilograms each. Endemic to Seychelles; The Corypha species have the largest inflorescence of any plant, up to 7.5 meters tall and containing millions of small flowers.

Key differences from similar families

Rarely confused with other families except Cyclanthaceae.

ü palms are never root climbers (some Cyclanthaceae are root climbers).

ü palms do not have perianth parts in fours (Cyclanthaceae perianth parts, where present, in fours).

ü Cyclanthaceae leaves do not develop like palm leaves.

ü Cyclanthaceae have distinctive inflorescence structure/organization (spicate, alternating male and female flowers) not found in palms.

RANGE AND HABITAT They are abundant throughout the tropics, and thrive in almost every habitat therein. Their diversity is highest in wet, lowland tropical forests, especially in ecological ‘hotspots’ such as Madagascar, which has more endemic palms than all of Africa. Only an estimated 130 palm species grow naturally beyond the tropics, mostly in the subtropics. The northernmost native palm is Chamaerops humilis L., which reaches 44°N latitude in southern France. The southernmost palm is the Rhopalostylis sapida H.Wendl. & Drude, which reaches 44°S on the Chatham Islands where an oceanic climate prevails. More than two-thirds of palm species live in tropical forests, where some species grow tall enough to form part of the canopy and shorter ones form part of the understory; the rarest palm known is the Hyophorbe amaricaulis Mart. The only living individual that remains is at the Botanic Gardens of Curepipe in Mauritius.

MAINLY USES in economic importance palms are second only the grassesand possibly legumes. Species in the five genera Acrocomia, Astrocaryum, Attalea (including Maximiliana and Orbignya, mainly A. speciosa Mart. ex Spreng., known as babaçu), Elaeis and Oenocarpus comprise the most important oil-bearing palms of the region. Indigenous peoples depended upon these palms as a source of vegetable oil and subsistence utilization continues to this day. These palms produce high quality oil; Oenocarpus oil has been compared to olive oil. But the quantity of oil-bearing fruit in these wild palms is low. In Brazil, palm leaf base fibers are collected from Attalea funifera Mart. (Bahia piassava) and Leopoldinia piassaba Mart. (Pará) and primarily used to manufacture brushes and brooms. Palm wood from the genera Bactris, Iriartea, Socratea and Wettinia is reported to be the highest quality. There are many abundant palms species in these four genera which could be exploited for specialized wood products.

USEFUL TIPS FOR GENERIC IDENTIFICATION

Spiny/armed palms:

§ Acoelorraphe (petiole only)

§ Acrocomia

§ Aiphanes

§ Astrocaryum

§ Bactris

§ Brahea (teeth on petiole)

§ Copernicia (petiole)

§ Cryosophila (root spines on stem)

§ Desmoncus

§ Elaeis (petiole)

§ Mauritiella (root spines on stem)

§ Rhapidophyllum (leaf sheath spines)

§ Serenoa (teeth on petiole)

§ Trithrinax (leaflet tips, sheath fibre spines)

§ Washingtonia (teeth on petiole)

§ Zombia (sheath fibre spines).

Fan-leaved palms:

§ Acoelorraphe

§ Brahea

§ Chelyocarpus

§ Coccothrinax

§ Colpothrinax

§ Copernicia

§ Cryosophila

§ Hemithrinax

§ Itaya

§ Lepidocaryum

§ Leucothrinax

§ Mauritia

§ Mauritiella

§ Rhapidophyllum

§ Sabal

§ Schippia

§ Serenoa

§ Thrinax

§ Trithrinax

§ Washingtonia

§ Zombia

Climbing palms:

§ Desmoncus

§ Chamaedorea elatior

Stilt rooted palms:

§ Iriartella

§ Dictyocaryum

§ Iriartea

§ Socratea

§ Wettinia

Vegetable ivory palms:

§ Ammandra

§ Aphandra

§ Phytelephas

Flowers emerging from pits:

§ Welfia

§ Pholidostachys

§ Calyptrogyne

§ Calyptronoma

§ Asterogyne

§ Geonoma

Fruit scaly (like reptile scales):

§ Mauritia

§ Mauritiella

§ Lepidocaryum

§ Raphia

SYSTEMATIC 831 spp. in New World. 532 spp. in South America in 55 genera, 16 of then absent in Brazil, all except two in Colombia. 282 spp. in Brazil and 257 in Colombia; only 102 in Mexico; subfamily Nypoideae (1/1, Sri Lanka, Bengal, SE Asia, Malesia, Melanesia to New Guinea, N Queensland and Melanesia) does not occur in South America.

1. SUBFAMILY CALAMOIDEAE (17/540) ‣ three tribes, Eugeissoneae (1/6, Thailand, the Malay Peninsula, Borneo) and Calameae (9/485, tropical Africa, tropical Asia to Samoa, with their largest diversity in W Malesia) do not occur in South America; among South American Lepidocaryeae, subtribe Ancistrophyllinae (3/22, tropical W and C Africa) not occur in South America.

∎ SUBTRIBE RAPHIINAE (1/20) - a single genus.

1. Raphia P.Beauv. Solitary or caespitose. 28 spp. of Africa, one in Madagascar, and one in New World: R. taedigera (Mart.) Mart., why exibes bizarre distribution: Nigeria to Cameroon, inundated riverine habitats of Central America (Nicaragua, Costa Rica, Panamá) as well as in the estuaries of the Atrato and Amazon rivers of Colombia and Brazil.

Raphia spp. have the largest leaves of any plant, up to 25 meters long and 3 meters wide.

∎ SUBTRIBE MAURITIINAE ‣ all genera in South America.

2. Lepidocaryum Mart. Slender, ceaspitose, unarmed, spreading slender rhizomes. Only one sp., Lepidocaryum tenue Mart., of Guyana to Peru and N Brazil.

3. Mauritia L.f. Massive, unarmed, solitary fan-leaved tree palm, often forming dominant stands in swampy areas. Moriche, buriti palms. Two spp. from tropical South America, M. carana Wallace restricted in Amazon region, and M. flexuosa L. f. in over continent.

Mauritia flexuosa L. is the Neotropical most abundant palm, forming population with million on individues in center Brazil, mainly in wet places, seasonal soils and riversides; known as buriti occurring as dense stands in permanently swampy areas, particularly in the Amazon rainforest; management of natural stands could enhance fruit and leaf production to provide food items and fiber.

4. Mauritiella Burret. Moderate, ceaspitose; stems bearing spinelike adventitious roots. 5 spp. in northern South America, 4 in Colombia, two endemics; three spp. in Brazil, one endemic.

2. SUBFAMILY CORYPHOIDEAE (47/480–490) ‣ tribes Chuniophoeniceae (4/6, Arabian Peninsula to Vietnam), Corypheae (1/5, India, Sri Lanka, Burma, Andaman Islands, SE Asia, Malesia to New Guinea and tropical Australia), Caryoteae (2/c 40, tropical Asia, tropical Australia, Melanesia), Borasseae (8/22, tropical regions in the Old World) and Phoeniceae (1/14, Canary Islands, Mediterranean, tropical and subtropical regions in northern and central Africa and Asia east to S China, the Malay Peninsula and Sumatra) do not occur in South America.

2.1 CORYPHOIDEAE ▸ TRIBE SABALEAE (1/14) - a single genus.

5. Sabal Adans. 15 spp. in C U.S.A. to Panamá, Caribbean, with S. mauritiiformis (H. Karst.) Griseb. & H. Wendl. also in northern coasts of Colombia and Venezuela.

2.2 CORYPHOIDEAE ▸ TRIBE CRYOSOPHILEAE (11/80) - outsiders are Thrinax (4; Cuba; Jamaica; Central America, Caribbean), Schippia (1; Belize), Zombia (1; Hispaniola), Hemithrinax (3; Cuba), Leucothrinax (1; Florida Keys, Caribbean).

6. Chelyocarpus Dammer. Solitary or clustering, unarmed; leaves palmate, or costapalmate; flowers solitary, petals like sepals. 4 spp. from over Amazon rainforest up to Central America, two in Brazil, none endemic.

7. Coccothrinax Sarg. Slender to moderate palms with leaves unarmed and petiole base not cleft. 52 spp. in Caribbean Basin, incl. Florida, Mexico and Central America, two up to South America in Colombia and Venezuela; most species rich fan-leaved genus in Neotropics, largely restricted to Caribbean islands and Central America.

8. Cryosophila Blume. Solitary palms, hermaphroditic; stem bearing root spine; leaves palmate, sheaths splitting basally; flowers solitary. 8 spp., six from Mexico to Panamá, one endemics to Colombia, and one in Colombia and Panamá.

9. Itaya H.E.Moore. Moderate, solitary, unarmed, leaves palmate, flowers solitary. Only one rare sp., I. amicorum Moore, found only in Amazon valleys in Colombia, Amazonas state in N Brazil, and Peru.

10. Sabinaria Galeano & R. Bernal. Leaf blades with a single deep, medial, abaxial split, and short abaxial splits in each segment, mostly unisexual flowers with biseriate perianth, calyx connate with the corolla at a single place on its margin, large, tightly appressed, persistent rachis bracts that hide the pistillate flowers, and fruits tightly packed and hidden among leaf bases, often covered by litter. Only one sp., S. magnifica Galeano & R. Bernal., restricted to rainforests near Colombia and Panamá frontier, in wet low elevations.

11. Trithrinax Mart. Solitary or clustering, hermaphroditic; leaves palmate or costapalmate; flowers solitary. 5 spp. in Bolivia to N. Argentina and Brazil (4, three endemics); T. campestris (Burmeist.) Drude & Griseb. is only Cono Sur. T. schizophylla Drude occur in Porto Murtinho municipality, SW Mato Grosso do Sul state, Brazil.

2.3 CORYPHOIDEAE ▸ TRIBE TRACHYCARPEAE (19/300–310) - outsiders are Colpothrinax (3; Central America; W Cuba, Isla de la Juventud), Washingtonia (2; SW U.S.A., NW Mexico), Pritchardia (25–35; Solomon Islands, Fiji, Samoa, Tonga, the Cook Islands, the Tuamotu Islands, Hawaii), Brahea (11; Mexico, Central America), Rhapidophyllum (1; SE U.S.A.), Maxburretia (3; peninsular Thailand, the Malay Peninsula), Chamaerops (1; W Mediterranean), Trachycarpus (11; Himalayas, China, northern Thailand), Guihaia (3; southern China, northern Vietnam), Rhapis (10; S China, SE Asia), Serenoa (1; SE U.S.A.), Acoelorraphe (1; Central America), Livistona (c 30; NE Africa, Arabian Peninsula, Ryukyu Islands, tropical and E Asia to tropical Australia), Johannesteijsmannia (4; Hainan, Vietnam, Thailand, the Malay Peninsula, northern Sumatra, Borneo), Lanonia (8; Indochina), Pholidocarpus (6; peninsular Thailand, the Malay Peninsula, Sumatra, Borneo, Moluccas), Saribus (9; Philippines, Raja Ampat Islands, Banggi Island, Sulawesi, Moluccas, New Guinea, Nggela in Solomon Islands, southernmost New Caledonia), Licuala (c 150; Himalayas, southern China, SE Asia, Malesia to New Guinea and tropical Australia, Vanuatu).

12. Acoelorraphe H.L.Wendl. Moderate, clustered, leaves briefly costapalmate. Only one sp., A. wrightii (Griseb. & H. Wendl.) H. Wendl. ex Becc., in S Florida, Caribbean, Mexico to Colombia.

13. Copernicia Mart. Solitary trees; leaves palmate to shortly costapalmate. 28 spp., Hispaniola (2), Cuba (24), C. prunifera (Mill.) H.E. Moore in NE Brazil, C. alba Morong in Chaco region, C. tectorum (Kunth) Mart. in Venezuela and Colombia.

The carnaúba palm C. prunifera represents the regions chief commercial source of hard vegetable wax, constitute almost pure stands in seasonally- flooded river valleys in NE Brazil. Leaves of this fan palm have a coating of hard wax which is obtained by cutting and drying the leaves and then mechanically chopping them into small pieces to dislodge the wax particles; C. alba in Paraguay and adjoining parts of Brazil, Argentina and Bolivia forms huge stands on areas subject to periodic flooding and periodic drought; the largest populations have been estimated to contain half a billion individuals.

3. SUBFAMILY CEROXYLOIDEAE (8/46) ‣ three tribes, Cyclospatheae (1/4, SE Florida, E Mexico, Caribbean) does not occur in South America.

3.1 CEROXYLOIDEAE ▸ TRIBE CEROXYLAE (4/34) - outsiders are Oraniopsis (1; NE Queensland), Ravenea (20; Madagascar, the Comoros).

14. Ceroxylon Humb. & Bonpl. Tall; inflorescences solitary; the Andean wax palms - conspicuous in Andean forests, pinnate -leaved, often very tall, stems waxy. 13 andean spp. from coastal Venezuela to Bolivia, centered in Colombia (7), Ecuador (6) and Peru (7) - conspicuous in Andean forests, often very tall, stems waxy.

C. quindiuense (H. Karst.) H. Wendl., Colombia's national tree, is the tallest monocot of the world, reaching up to 60 meters tall; for palm trees, C. parvifrons (Engel) H. Wendl., grows up to 3,500 m (altitude in Ecuador) from Venezuela to Bolivia, the highest place for palms worldwide.

15. Juania Drude. Leaflets discolorous; inflorescence solitary. Only one sp., J. australis (Mart.) Drude ex Hook. f., endemic to Juan Fernandez Islands, in Pacific Chile.

3.2 CEROXYLOIDEAE ▸ TRIBE PHYTELEPHATHEAE (3/8) - all genera in South America; the vegetable ivory palms, tagua. Dioecious with short-lived inflorescences, the males elongate and comprising numerous reduced flowers with up to 1,000+ stamens, the females congested and consisting of the largest of all palm flowers (up to c. 20 cm long); fruits warty containing numerous seeds with hard ivory-like endosperm.

16. Ammandra O.F.Cook. Solitary or caespitose; stem prostrate or decumbent. Only one sp., A. decasperma O.F. Cook, in Colombia and Ecuador.

17. Aphandra Barfod. Stem erect; petiole less than 2 m; leaflets regularly arranged. Only one sp., A. natalia (Balslev & A.J. Hend.) Barfod, rare disjunct in forests, in Ecuador and Acre state in W Brazil.

18. Phytelephas Ruiz & Pav. Stem erect or procubent; petiole less than 2m. 6 spp. Colombia to Bolivia and N Brazil (only P. macrocarpa Ruiz & Pav., non endemic); vegetable ivory is the hardened endosperm of palms in this genus.

4. SUBFAMILY ARECOIDEAE (111/1.260–1.310) ‣ largest subfamily, 14 tribes and more 100 genera; all tribes have pinnate or bipinnate leaves and flowers arranged in groups of three, with a central pistillate and two staminate flowers; tribes Podococceae (1/1, Central Africa), Oranieae (1/28, Madagascar, southern Thailand to New Guinea), Sclerospermateae (1/3, tropical W Africa), Pelagodoxeae (2/2, New Guinea, Melanesia, Marquesas Islands) and Areceae (61/595–600) do not occur in South America.

4.1 ARECOIDEAE ▸ TRIBE IRIARTEEAE (5/31–35) - all genera occur in South America.

19. Dictyocaryum H.Wendl. Solitary, rarely clustered; leaves plumose. Three spp., Colombia to Ecuador, Peru, Bolivia, Guyana, Venezuela and Acre state in N Brazil (only the Amazon widely distributed D. ptarianum (Steyerm.) H.E. Moore & Steyerm.), from low elevations up to 1,800 m, mainly in montane forests.

20. Iriartea Ruiz & Pav. Solitary, robust. Only one sp., I. deltoidea Ruiz & Pav., which is found from Nicaragua south into Bolivia and a great portion of western Amazon rainforest.

21. Iriartella H.Wendl. Caespitose, slender; inflorescence solitary, bissexual, branching to 1 order. Two spp., from Amazon rainforests of Guyana to Colombia, Bolivia, Brazil (both spp., none endemics).

22. Socratea Karst. Solitary, inflorescence solitary, branching 1 order. 5 spp. from Nicaragua to South America, four in smaller ranges, two in Brazil, none endemics.

23. Wettinia Poepp. ex Endl. Solitary or clustered, moderate to robust. 21 spp., one in Panamá, an the 20 remaining in Amazon rainforests of South America, only three in Brazil, none endemic.

4.2 ARECOIDEAE ▸ TRIBE CHAMAEDOREEAE (5/110–120) - outsiders Hyophorbe (5; Réunion and Rodrigues), Gaussia (5, S Mexico, Central America, the Greater Antilles).

24. Chamaedorea Willd. Small, erect, procubent, climbing, or acaulescent, solitary, or clustered; most species rich palm genus in the Neotropics, most diverse in Central America, mostly slender understorey dioecious palms, pinnate- or entire-leaved, with flowers solitary (not in clusters) on the inflorescence branches. 104 spp., most species rich palm genus in the Neotropics, most diverse in Central America and Mexico (51, 20 endemics); only 16 species in South America, three in Brazil, none endemics, absent in Guianas and Caribbean; the smallest leaves of palms are the C.tenella W. Endl. and C. tuerckheimii (Dammer) Burret, both Mexico (Veracruz, Oaxaca) to Honduras, wiht blades sometimes less 15 cm long.

25. Synechanthus H.Wendl. Slender solitary or clustered, monoecious. Three spp., two in from Mexico and Central America, and S. warscewiczianus H. Wendl. from Mexico to Colombia and Ecuador.

26. Wendlandiella Dammer. Dwarf, slender clustering, dioecious. Only one sp., W. gracilis Dammer, with three varieties, in Peru to Bolivia and Acre state in N Brazil.

4.3 ARECOIDEAE ▸ TRIBE ROYSTONEEAE (1/11) - a single genus.

27. Roystonea O.F.Cook. 10 spp., 7 endemic to Caribbean, one only in Mesoamerica, one in Florida to Central America and Caribbean, and R. oleracea (Jacq.) O.F. Cook, from Lesser Antilles to NE Colombia and Venezuela.

4.4 ARECOIDEAE ▸ TRIBE REINHARDTIEAE (1/6) - a single genus.

28. Reinhardtia Liebm. 6 spp., Mexico, Central America and Caribbean, three of then from Mexico or Central America to NW Colombia.

4.5 ARECOIDEAE ▸ TRIBE EUTERPEAE (5/31) - outsider Neonicholsonia (1; N. watsonii; Central America).

29. Euterpe Mart. Tall, moderate to robust, solitary or clustered pinnate-leaved tree palms, assai palms; leaf sheaths tubular and forming a cylinder (crownshaft); inflorescence resembles a horse's tail. 8 spp., Central and South America, from Belize southward to Brazil (5, none endemic), Peru and Argentina; these palms grow mainly in swamps and floodplains.

E. edulis Mart., known as palmito juçara is a single-stemmed palm restricted to the Atlantic Forest in South America, with high exploitation for commercial palm heart production in Brazil, Argentina and Paraguay.

30. Hyospathe Mart. Small, rarely moderate, solitary or clustered. 7 spp. from Costa Rica to Brazil in Amazon region in Brazil (only the widely distributed H. elegans Mart.), highly centered in Colombia to Peru.

31. Oenocarpus Mart. Solitary or clustered, moderate to very large pinnate -leaved palms with inflorescences similar to a horse's tail. 8 spp. all in South America, also in Trinidad, one up to southern Central America; 5 in Brazil, none endemic.

32. Prestoea Hook.f. Small to moderate, clustered. 9 spp. from Costa Rica to Bolivia and Brazil, also Caribbean; all in South America, only two in Brazil, none endemics.

4.6 ARECOIDEAE ▸ TRIBE LEOPOLDINIEAE (1/3) - a single genus.

33. Leopoldinia Mart. Solitary or clustered; inflorescence interfoliar, solitary. Three spp., endemics to the Guiana Shield from NW Brazil, SE Colombia and S Venezuela, one extends to C Brazil (all spp., none endemic).

4.7 ARECOIDEAE ▸ TRIBE MANICARIEAE (1/1) - a single genus.

34. Manicaria Gaertn. Solitary or caespitose, leaves undivided or variously divided. Two spp., M. saccifera Gaertn. found in Trinidad, Central and South America, and M. martiana Burret only from Brazil and Colombia; it has one of the largest known leaves in the plant kingdom (up to 8 meters in length).

4.8 ARECOIDEAE ▸ TRIBE GEONOMATEAE (6/c 100) - outsider Calyptronoma (3; Jamaica; Cuba, Hispaniola; Puerto Rico).

35. Asterogyne H.Wendl. ex Benth. & Hook.f. Small to medium-sized, solitary, unarmed, pleonanthic, monoecious palm; stem smooth, brown to brownish cream, erect, sometimes basally decumbent, occasionally with basal and/or lateral vegetative branches, unarmed; inflorescence interfoliar, solitary, branched to one order, rarely spicate or forked, erect at anthesis, becoming pendente in fruit, protandrous. 5 spp., one from Belize to Ecuador, three endemics to Venezuela, and A. guianensis Granv. & A.J. Hend. endemic to a single locality in the E French Guiana, close to Brazilian frontier; Henderson and Balick (1987) suggested that this species, at that time not yet described, may occur also in N Brazil.

36. Calyptrogyne H.Wendl. 9 spp., 7 from Mexico to Panamá, one from Central America and Colombia, another endemic to Colombia.

37. Geonoma Willd. Small, solitary or clustered; inflorescence spicate or branched with flowers emerging from pits., most frequently encountered understorey palm genus in South America, mostly rather small, leaves pinnate or often entire; easily confused with related genera (e.g. Calyptrogyne, Calyptronoma, Asterogyne). 78 spp., distributed from Mexico to C Bolivia and NE Paraguay and Haiti, 70 in South America; the country with the highest diversity are Colombia (40), Ecuador (29) and Brazil (23, only 4 endemics).

The areas with the highest number of species are situated along the mountain regions and adjacent areas of C and W South America, in Costa Rica (Alajuela), Panamá (Chiriquí, Coclé), Colombia (Antioquia, Chocó, Valle, Caquetá, Putumayo), Ecuador (Sucumbios), and Peru (Amazonas, Pasco); no species of Geonoma occurs in areas with less 1,000 mm annual precipitation; barely in savannas of C Brazil (cerrado) and dry seasonal scrubland of NE Brazil (caatinga), and in Llanos region.

38. Pholidostachys H.Wendl. ex Benth. & Hook.f. Solitary, inflorescences spicate or branched to 1(-3) orders. 7 spp., P. pulchra H. Wendl. ex Burret from Costa Rica to Colombia, remaining all in northern South America, including the only Brazilian species, the over widely distributed P. synanthera (Mart.) H.E. Moore.

39. Welfia H.Wendl. Solitary tree; inflorescences branched to 1(-2) orders. Two spp., from Costa Rica to Peru (one endemic).

4.9 ARECOIDEAE ▸ TRIBE COCOSEAE (20/340–350) - three subtribes, all in South America.

∎ SUBTRIBE ELAEIDINAE (2/3) ‣ both genera in South America.

40. Barcella Drude. Solitary, acaulescente, unarmed; petiles margun shaped. Only one sp., B. odora (Trail) Drude, confined a small area of the banks of the Rio Negro and tributaries, Amazonas state in north Brazil, recently collected in SW Roraima state, also in Brazil.

41. Elaeis Jacq. Solitary, erect; petioles armed with fiber spines and spinelike eroded leaflet basis. Two spp., E. guineensis Jacq. in Africa and E. oleifera (Kunth) Cortes in Neotropics, from Costa Rica to N Brazil up Ecuador, Peru and Guianas, in rainforests.

Being one of only two genera which occurs in both hemispheres; internationally, the most significant contribution of the American oil palms thus far concerns E. oleifera, which is being used as a source of germplasm for a breeding program to improve disease resistance in E. guineensis.

∎ SUBTRIBE BACTRIDINAE ‣ outsider Hexopetion (2; Mexico to Panamá).

42. Acrocomia Mart. Acaulescent ot erect, solitary; spines present, reaching up to 20 cm. 7 spp., two from Brazil, Bolivia and adjacent Cono Sur, A. aculeata (Jacq.) Lodd. ex Mart. widely distributed, and four endemics to Brazil.

43. Aiphanes Willd. Solitary or caespitose, acaulescent to erect. 36 spp. from Costa Rica to Amazon rainforest of Bolivia and W Brazil (only Acre state, 3 spp., none endemics) and Caribbean, 35 in South America (one restricted for Caribbean), mainly Colombia (29, 18 endemics).

44. Astrocaryum G.Mey. Acaulescent to erect, solitary or clustered; inflorescence branching to 1 order; leaflets single-fold; viciously spiny stemless to canopy pinnate-leaved tree palms, leaf undersurface white. 38 spp. over neotropics, 35 in South America; some spp., are many thorned. 22 spp. in Brazil, eight endemics.

45. Bactris Jacq. ex Scop. Solitary or caestipose, acaulescent or erect; most species rich of the spiny genera, undergrowth to canopy pinnate-leaved palms (sometimes entire-leaved), includes the peach palm B. gasipaes Kunth (pejibaye, chonta, pupunha). 85 spp., from Mexico and Caribbean south to Paraguay, with the greatest diversity in Brazil (45, 13 endemics); 72 in South America; two species from coast Bahia and Amazonas state are rare plants in Brazil, by Plantas Raras do Brasil’s book.

46. Desmoncus Mart. Slender, caespitose, erect or climbers, New World counterpart to the true rattans of the Old World; spiny climbing palm with pinnate leaves, rattan-like. 22 spp. distributed from Mexico to Paraguay and Brazil, 18 in South America, highest diversity in Colombia, with 11 species, followed by Brazil with 7 (one endemic).

Not all Desmoncus species are climbers, and not all Neotropical climbers are Desmoncus. One other genus, Chamaedorea Willdenow contains a single species with climbing stems, C. elatior Mart. Stems are mostly clustered. Only one species, D. giganteus A.J. Hend. has solitary stems, although this is recorded from only two specimens; stems in all but one species, D. stans Grayum & de Nevers, are not free-standing, and climb by means of acanthophylls. However, D. mitis Mart. subsp. ecirratus is recorded on several specimen labels as being non-climbing; despite the climbing habit of most species, stems of Desmoncus seldom reach great heights; the mean plant height of all climbers is only 6.2 m.

∎ SUBTRIBE ATTALEINAE ‣ outsider Beccariophoenix (1; E Madagascar), Jubaeopsis (1; Pondoland in E Cape), Voanioala (1; Madagascar), Paschalococos (1; Easter Island, extinct), Cocos (1; tropical and subtropical coastal areas).

47. Allagoptera Nees. Ceaspitose, acaulescent or very short stems, stems also branching aerially by forking. 6 spp. in Brazil (all species, 4 endemics), Bolivia, Paraguay and Argentina (Missiones), in sandy and rocke well-drained soils.

48. Attalea Kunth. Small to massive, solitary, acaulescent or erect. American oil palms, often massive tree palms with huge pinnate leaves, visible in agricultural lanscapes. 35 spp., from Mexico (2), Central America (4) to South America (34) up Bolivia, Paraguay, and southern Brazil (23, 8 endemics, one a rare palm, by Plantas Raras do Brasil’s book, endemic to northern Diamantina Range, Bahia state), also Caribbean (3, two in Trinidad and Tobago, along the southern edge of the region, and one in Haiti).

A variety of species in Attalea remain important sources of edible oil, thatch, edible seeds and fibre; the leaves of A. butyracea (Mutis ex L. f.) Wess. Boer and A. maripa (Aubl.) Mart.are used extensively for thatching; several species are oil palms, with A. speciosa Mart. ex Spreng. among the most important economically. Piassava fibres, extracted from the leaf bases of A. funifera Mart., are commercially important. A. cuatrecasasiana (Dugand) A.J. Hend., Galeano & R. Bernal, endemic to Bajo Calima region in Choco from W Colombia, is third largest fruit among all palms, exceeded in fruit size among palms only by coconut (Cocos) and double coconut (Lodoicea); the fruits are up to 14cm ✕ 10 cm in diameter, besides some sources take Brazilian A. speciosa Mart. as the largest fruit in this genus (12cm ✕ 10cm).

49. Butia Becc. Solitary, acaulescent or erect, petiole usually armed; inflorescences branched 1 order. 21 spp., cooler, dry areas of southern Brazil (19 in Brazil, 11 endemics), Paraguay, Uruguay and Argentina.

B. lallemantii Deble & Marchiori, also known as ‘butiá-anão’ due to its reduced height, is among the species that stand out in sandy patches; the species occur only in the sandy grasslands of SW Rio Grande do Sul, where it usually constitutes populations with large number of individuals, standing out in the landscape.

50. Jubaea Kunth. Solitary, massive tree; petiole unarmed. Only one sp., J. chilensis (Molina) Baill., central Chile; a rare non tropical genus of Arecaceae.

51. Parajubaea Burret. Solitary trees; petiole unarmed; inflorescence branched to 1(-2) orders. Three spp., one in Ecuador and two in Bolivia.

52. Syagrus Mart. (inc. Lytocarium) Solitary or clustered, acaulescent to trees; petiole rarely armed; inflorescence branched to 1 order, rarely spicate. 75 spp., 74 in South American (the exception is one endemic to Caribbean; only this and S. romanzoffiana (Cham.) Glassman make outside continent - both in Caribbean), mainly in Brazil (68, 59 endemics). S. wedermanii Burret from Diamantina Range in Bahia state, and S. leptospatha (non recognized by VPA) from center Mato Grosso do Sul state are rare plants in Brazil, by Plantas Raras do Brasil’s book.

17. COMMELINALES

FAMILIES ABSENTS IN SOUTH AMERICA: HANGUANACEAE (1/22), PHYLIDRACEAE (3/6).

PONTEDERIACEAE

§ FULLY CHLOROTROPHIC

Genera/spp. 2/40 Distribution pantropical, with the largest diversity in tropical America, some spp. in subtropical and warm-temperate regions in both Hemispheres. Habit bisexual, usually perennial (sometimes annual) herbs. Aquatic (sometimes freely floating) or helophytes. Vegetative stems with indeterminate growth. It is a small family of heterostylous aquatic plants, occurring in tropical and subtropical waters.

Common in over Brazil, notably in Pantanal, Amazon rainforest, and lagoons in center Brazil; some wartergarden ornamentals, including Pontederia (water hyacinth) which is a devastating weed of warm waterways. Charles Darwin was interested in the specialized form of heterostyly found in the family, known as tristyly. It is best known for the water hyacinth (Pontederia crassipes Mart.), which is an invasive spp. in many waterways.

SYSTEMATIC both genera in South America.

1. Heteranthera Ruiz & Pav. (inc. Hydrothrix) Submerged, floating or emergente spp., annual or perennial; inflorescence a spike or paired, often solitary. 18 spp., 16 from North America up to Argentina (13 in Brazil, 4 endemics) and two in Africa and Madagascar; the bizarre H. gardneri (Hook.f.) M. Pell. is endemic to water bodies in dry NE Brazil.

2. Pontederia L. (inc. Eicchornia) Floating, submersed or rooted, creeping or ascending perennial with long rhizomes or stolons; stems trailing to erect, delicate to spongy, branching at the base, rarely branching at the upper half, rooting at the basal nodes or along the whole stem; internodes reduced to elongate, producing stolons or not; some species commonly found as free -floating in watercourses throughout the topics, has leaves with inflated petioles. 26 spp., over Neotropics, 15 in New World, all in Brazil, two endemics; almost all Paleotropical species belong to Pontederia subg. Monochoria except for Pontederia natans P.Beauv., which is restricted to Africa and is a member of Pontederia subg. Eichhornia (Kunth) M.Pell. & C.N.Horn; species in Pontederia can range from paludal to free-floating plants, thus occurring in a wide range of water bodies, from perennial to temporary, but most commonly in slow or stagnated water.

§ subg. Oshunae ‣ herbs erect emergent or free-floating, stems inconspicuous; sessile leaves early deciduous, petioles generally inflated; only P. crassipes Mart., over South America; this species is probably the most aggressive aquatic weed ever known in the tropics.

§ subg. Monochoria ‣ flowers pedicellate, enantiostylous, perianth only basally connate, campanulate; 10 spp., Old World.

§ subg. Eichhornia ‣ herbs procumbent-emergent, stems elongate; sessile leaves late deciduous; 4 spp., 3 Neotropical and P. natans P.Beauv. restricted to continental Africa and Madagascar.

§ subg. Pontederia ‣ flowers sessile, non-enantiostylous; ovary 1-locular by abortion, fertile locule 1-ovulate, placentation pendulous; fruit an achene; 8 spp., all neotropical.

§ subg. Cabanisia ‣ herbs erect emergent or free-floating, stems inconspicuous; petioles never inflated; 3 spp., Brazil to Argentina, scatered from NW South America, Central America and Mexico.

COMMELINACEAE

§ FULLY CHLOROTROPHIC

Genera/spp. 36/650-660; Distribution tropical and subtropical regions, with the largest diversity in Africa, South Asia, Mexico and northern Central America; some spp. in temperate E Asia, E and S North America and Australia. Habit usually bisexual (often monoecious or andromonoecious, rarely polygamomonoecious), usually perennial (rarely annual) herbs. Usually somewhat succulent, sometimes twining, very rarely epiphytic. Bulb rarely present. Nodes swollen.

Most genera cited here are native to the Neotropics. Only Murdannia has spp. introduced from the Old World. Spp. of Dichorisandra, Tradescantia and Gibasis geniculata (Jacq.) Rohweder are used as ornamentals. Several spp. of Commelina are weedy; leaves of Siderasis fuscata are a rich green with a lighter green mid-rib vein. Upon close inspection, you will notice the leaves are covered with tiny brownish hairs.

This family has two subfamilies: one occurs from in Zimbabue and Australia; another is cosmopolitan. Faden and Hunt (1991) accepted 2 subfamilies: Cartonematoideae (raphide canals absent or next to the leaf veins), which was divided into 2 tribes: Cartonemateae and Tricerateleeae; and Commelinoideae (raphide canals present and mostly between leaf veins, etc), also comprising 2 tribes: Tradescantieae and Commelineae; to identify the genera it is important to observe flower characters in the field, e.g.the colour and shape of the petals, and the number of stamens, their disposition and the dehiscence of the anthers; the plant habit is also important and may vary between rosulate, rhizomatous or stoloniferous, trailing plants, and erect or semi-scandent herbs to 4 m tall.

Commelinaceae comprises 42 genera and ca. 720 species; diversity centers for the family can be recognized in Brazil with 46 of 50 species of Dichorisandra Mikan, in India with 27 of 54 species of Murdannia Royle, in Africa with 57 of 62 species of Aneilema R. Brown and Asia with 170 species of Commelina; 277 spp. in New World, 146 in South America.

Key differences from similar families - differs from:

ü Pontederiaceae, which are aquatic and floating or immersed and have inflated petioles.

ü Haemodoraceae, where perianth whorls are fused basally and septal nectaries are present.

SYSTEMATIC three subfamilies, one in South America and two outside: Cartonematoideae (1/1, Zimbabue and Mozambique, known only two collections) and Tricellateroideae (2/12, Australia); among Commelinoideae, three tribes, two in South America, Palisoteae (1/30, tropical Africa) absent, and some genera unplaced within.

1.1 COMMELINOIDEAE ▸ TRIBE COMMELINEAE (11/c. 340) ‣ outsiders Anthericopsis (1; tropical E Africa), Dictyospermum (5; India, Sri Lanka to New Guinea), Pollia (17; tropical regions in the Old World, tropical Australia; one species in Panamá), Polyspatha (3; tropical W Africa), Pseudoparis (3; Madagascar), Tricarpelema (8; tropical Asia, Cameroon, Gabon and Equatorial Guinea).

1. Aneilema R.Br. Perennial or annual herbs; inflorescences terminal and axillary (rarely all axillary) thyrses (sometimes reduced to a single cincinnus); flowers strongly zygomorphic. 64 spp., pantropical, but mainly in Africa; widely distributed in tropics, only two in New World, A. brasiliense C.B. Clarke from Brazil and Venezuela, and A. umbrosum (Vahl) Kunth widely distributed in Central and South America.

2. Buforrestia C.B.Clarke. perennial herbs; roots fibrous; leaves spirally arranged; inflorescence axillary, perforating the sheaths, contracted or elongate, composed of 1-5, sessile contracted cincinni; floers zygomorphic. Three spp., in forest understory, two in W Africa and B. candolleana C.B. Clarke in Amazon rainforest in Guyana, French Guiana, Brazil and Suriname.

3. Commelina L. Perennial or annual herbs; roots usually fibrous; inflorescence terminal or leaf-opposed, composed of 1-2 cincinni enclosed spathe; flowers strongly zygomorphic. 215 spp., over 45 spp. in tropical America, cosmopolitan, some a weeds, others very narrow endemic; 37 in New World, 19 in South America, 9 in Brazil (4 widely distributed, two more restricted, and 3 endemics).

The cream-yellowish petals of C. catharinensis Hassemer, J.P.R. Ferreira, Funez & J.D. Medeiros is by far the most striking and unique feature of this species, as no other species in South America present this character; this spp. is somewhat similar to C. rufipes Seub., which nevertheless has white flowers (instead of cream-yellowish) and whitish-silvery indehiscent fruits (instead of light brown dehiscent fruits); all other species of Commelina in South America have fully white, blue, or purple petals.

4. Floscopa Lour. Perennial or annual herbs; leaves spirally arranged; inflorescence terminal and axillary thyrses, commonly forming compound inflorescences; flowers zygomorphic. 20 spp., pantropical, mainly in aquatic situations, either in forests or open habitat; 5 spp. in New World, all widely distributed in tropical America (and in Brazil) except by F. perforans Rusby endemic to Bolivia.

5. Murdannia Royle. Herbs, perennial or annual, rhizomatous or not, with a definite or indefinite base, terrestrial to paludal to rooted emergent aquatics. Roots thin and fibrous or tuberous and fusiform. Rhizomes short to elongate. Stems trailing and ascending at the apex or erect, unbranched to densely branched, rooting in the rhizome and at the basal nodes, rarely at the distal ones when they touch the substrate. 60 spp., pantropical, 6 native in New World, all in Brazil (two endemics) and adjacent Bolivia, Venezuela, Paraguay and Guyana except a unnamed spp. known only one collection from Venezuela; although few in number, the Neotropical species of Murdannia exhibit all the extremes in inflorescence morphology found in Murdannia as a whole.

1.2 COMMELINOIDEAE ▸ TRIBE TRADESCANTINAE (15/C. 280) - outsiders tribes Cyanotinae (1/ 50; tropical and subtropical regions of the Old World, from Africa to N Australia), Coleotrypinae (3/27, tropical regions in the Old World, Madagascar), unplaced tribe Spatholirion (6; S China, SE Asia), Belosynapsis (6; Madagascar, tropical Asia from India to New Guinea), Aetheolirion (1; Thailand), Sauvallea (1; Cuba), Streptolirion (1; E Himalayas to SE Asia and Korean Peninsula), Stanfieldiella (4; tropical Africa).

∎ UNPLACED GENUS

6. Tripogandra Raf. Annual or perennial herbs, often succulents; roots fibrous; inflorescences composed of pairs of sessile cincinni subtended by small bracts; flowers moderately zygomorphic. 22 spp., Mexico to Uruguay and Caribbean, a exact half in South America, 7 in Brazil, 4 endemics; T. elata D.R. Hunt endemic to Distrito Federal and Goiás, and T. warmingiana (Seub.) Handlos endemic to Minas Gerais state are rare species in Brazil by Plantas Raras do Brasil’s book; however, the latter was collected also in Rio de Janeiro and Bahia recently.

∎ SUBTRIBE DICHORISANDRINAE ‣ all genera in South America.

7. Cochliostema Lem. Tank or creeping epiphytes, unique among New World Commelinaceae, rarely terrestrial; inflorescence axillary thyrses with cincinni subtended by large, colored bracts; flowers zygomorphic, fragrant. Two spp. from forests of Costa Rica to Ecuador and Colombia (one endemic).

8. Dichorisandra J.C.Mikan. Perennial herbsup to 1.5 m tall, with diverse habits; inflorescence terminal, axillary or basal, thyrsiform or reduced; flowers slightly zygomorphic, showy, mainly blue or purple. 53 spp., highly centered in Brazil: one only in Central America and Mexico, six from Guianas up Bolivia outside Brazil, 4 in Brazil and other countries (including the widely distributed D. hexandra (Aubl.) C.B. Clarke), and remaining 42 endemics to Brazil; four species, one in each SE Brazilian state, are rare species in Brazil by Plantas Raras do Brasil’s book.

9. Geogenanthus Ule. Perennial herbs, dracaenoid; roots with distal tubers; shoots produced by subterranean rhizomes; inflorescence basal, perforating bladeless sheats, consisting of 1-2(-4) cincinni; flowers slightly zygomorphic. 3 spp., forests understory in Acre state in Amazon rainforest of Brazil (only G. poeppigii (Miq.) Faden, in Acre and Amazonas states), Colombia, Bolivia, Peru and Ecuador.

10. Plowmanianthus Faden & C.R. Hardy. Decumbent, perennial ornamental rosette herbs to 40 cm tall; roots thin, fibrous, lacking tubers, the younger with indument of root hairs initially white or yellow but soon turning orange-brown; inflorescences axillary, borne among the lower (older) leaves, consisting of 1 (-3) 1- many flowered cincinnus subtended by a small lanceolate bract; flowers bisexual, cleistogamous or chasmogamous, monosymmetric. 5 spp., two endemic to Panamá and 3 restricted to portions of Amazonian rainforest in Colombia, Ecuador, Peru, and W Brazil (only one).

P. grandifolius Faden & C. R. Hardy is known from the Peruvian departments of Amazonas and Loreto, and the Brazilian states of Amazonas and Acre, in spatially diffuse, poorly defined populations, shallowly rooted in humus-rich and leaf-litter layers in well-shaded understories and streamsides of primary rainforest between 100–300 m elevation.

11. Siderasis Raf. Herbs or vines, perennial, with a definite base, terrestrial or rupicolous; roots thin, fibrous, sometimes forming terminal, small, fusiform to oblongoid tubers; rhizomes present or not; subterraneous stems present or not, when present buried deep in the soil, unbranched, produced directly from the short rhizome; internodes moderately elongate to elongate; inflorescence terminal, consisting of a simple cincinnus or thyrsiform. 6 spp., S. almeidae M. Pell. & Faden in southern Bahia state, 3 restricted of Espírito Santo state, and two endemics to Rio de Janeiro state, inc. S. fuscata (Lodd.) H.E. Moore, a rare species in Brazil by Plantas Raras do Brasil’s book, known only Rio de Janeiro and Niteroi cities.

∎ SUBTRIBE THYSANTHEMINAE ‣ outsiders Gibasoides (1; Mexico), Matudanthus (1; Mexico), Thyrsanthemum (3; Mexico), Weldenia (1; Mexico, Guatemala).

12. Elasis D.R.Hunt. Procumbent or scadent perennial herbs rooting at the nodes; inflorescence consisting of up to 5 simple cincinni, terminal, and solitary from the upper axils, forming a loose cluster; flowers actinomorphic. Two spp., E. hirsuta (Kunth) D.R. Hunt, endemic to high mountains in Ecuador, and E. guatemalensis (C.B.Clarke ex Donn.Sm.) M.Pell. from Mexico and Guatemala.

13. Tinantia Scheidw. Annual herbs; inflorescence paniculiforme or umbeliforme thyrses or composed of 1-2 cincinni, elongate; flowers zygomorphic. 13 spp., U.S.A. to Argentina, Caribbean, 6 in South America, two in Brazil, none endemics.

∎ SUBTRIBE TRADESCANTIINAE ‣ all genera in South America.

14. Callisia Loefl. Perennial or annual herbs, often succulent; roots thin, rarely tuberous; inflorescence terminal and/or axillary, composed of pairs of sessile sinsinni subtended by small bracts usually less than 1 cm long, often aggregated into compound inflorescences; flowers actinomorphic. 20 spp., Mexico to Paraguay and Caribbean; 6 in South America, 4 in Brazil, none endemics.

15. Gibasis Raf. Annual or perennial herbs; roots fibrous or tuberous; inflorescences terminal, composed of pairs or pseudo-umbels of stipitate cincinni; flowers zygomorphic. 14 spp. from Mexico (high diversity) to Paraguay and Caribbean, only the widely distributed G. geniculata (Jacq.) Rohweder in South America.

16. Tradescantia L. Herbs chamaephytes or geophytes, base definite or indefinite, perennial, frequently succulent, terrestrial, rupicolous or epiphytes; rhizomes absent; stems prostrate with ascending apex or erect, herbaceous to succulent, rarely fibrous, unbranched to branched only at base or little to densely branched, rooting at the basal nodes or at the distal ones when they touch the substrate. 86 spp., all American; the genus is currently circumscribed into 12 taxonomic sections and four series, and native to the Neotropics, but with a center of diversity in Mexico and Southern U.S.A. 5 subgenera in this genera:

§ subg. Austrotradescantia ‣ 16 spp., Bolivia, Brazil (15, 11 endemics), Paraguay, Uruguay and Argentina.

§ subg. Campelia ‣ c. 15 spp., Mexico to N South America; only one sp. in Brazil, T. zanonia (L.) Sw., geographically disjunct between the Amazon rainforest and the Atlantic Forest domains. T. zebrina Heynh. ex Bosse, an aggressive invasive species widely distributed in Brazil, belongs this subgenus.

§ subg. Mandonia ‣ c. 20 spp., disjunctively distributed across the American continent, with species occurring in North America, Central America, and South America; only one sp. in Brazil, T. ambigua Mart. ex Schult. & Schult.f., which is restricted to dry seasonal scrubland of NE Brazil (caatinga) and savannas of C Brazil (cerrado) domains.

§ subg. Setcreasea ‣ c. 10 spp., U.S.A. and Mexico.

§ subg. Tradescantia ‣ c. 30 spp., Canada to Mexico.

HAEMODORACEAE

§ FULLY CHLOROTROPHIC

Genera/spp. 15/c. 100 Distribution South Africa, New Guinea, Australia, Tasmania, E and SE North America, Central America, and northern South America, with the largest diversity in Australia. Habit bisexual, perennial herbs. Tuberous stem (Pyrrhorhiza, Tribonanthes) or bulb (Haemodorum) and often stolons. Roots and subterranean stems often intensely red to reddish-brown. All over the Neotropics, but also in the U.S.A. and Canada (Lachnanthes). The red-coloured pigment often present in the underground parts of 10 spp. in 8 genera (inc. Xiphidium caeruleum in New World) - arylphenalenone haemocorin - is unique in the flowering plants.

SYSTEMATIC two subfamilies, Conostylidoideae (6/62–70) only Australia, mainly southwestern; among Haemodoroideae, outsiders are Dilatris (4; Western Cape), Haemodorum (20; SW, N and E Australia, Tasmania), Cubanicola (1; Cuba), Lachnanthes (1; Nova Scotia and Massachusetts to Louisiana and Florida, Cuba), Barberetta (1; E Cape, KwaZulu-Natal), Wachendorfia (4; Western and E Cape).

1. Pyrrorhiza Maguire & Wurdack. Herbs to 1,5m tall with orange red corms aggregated in clusters. Only one sp., P. neblinae Maguire & Wurdack, endemic to Pantepui Life Zone, in venezuelan siade of Mount Neblina and adjacent areas in extreme south Venezuela; cloud forests, at elevations of 1,800 – 2,100 m.

2. Schiekia Meisn. Rhizomatous herbs to ca. 1 m tall; flowers cream white. Three spp. from Colombia to French Guiana, N Brazil (all of them, none endemics) and Bolivia; mountain savannas and woodlands.

3. Xiphidium Aubl. Rhizomatous or stoloniferous herbs to 0.5 m tall; white tepals. Two spp., X. caeruleum Aubl. in Mexico to Brazil, Bolivia up to French Guiana, and X. pontederiiflorum M. Pell., Hopper & Rhian J. Sm. from Panamá to Ecuador, in moist soils, often along watercourses.

18. POALES

FAMILIES ABSENTS IN SOUTH AMERICA: ECDEIOCOLEACEAE (2/3), FLAGELLARIACEAE (1/5), JOINVILLEACEAE (1/4).

LINEAGE 1 of 6: BROMELIACEAE

BROMELIACEAE

§ CARNIVOROUS (Brocchnia - Catopsis - Paepalanthus - Drosera - Heliamphora - Philcoxia - Genlisea - Utricularia - Pinguincula)

Genera/spp. 76/3,492 Distribution tropical and subtropical regions of America from Virginia in the U.S.A. to Patagonia in South America; one sp. of Pitcairnia in tropical West Africa. A few spp. are found on islands in the Pacific (Racinaea insularis (Mez) M.A.Spencer & L.B.Sm. of the Galápagos Islands, and Greigia berteroi Skottsb. and Ochagavia elegans Phil. of the Juan Fernandez Islands, in Chile).

All of the large genera have broad Neotropical distributions with the exception of Puya, which is largely restricted to the Andes, and Dyckia, which is mostly found in southern South America (mainly Brazil). Atlantic Forest is a well-known center of diversity for the family and holds ca. 900 spp., mainly Vriesea (166), Aechmea (136) and Neoregelia (96)), followed by high-altitude grasslands (campos de altitude) vegetation distributed along the Espinhaço Range in Minas Gerais and Bahia, sandy coastal plains, grasslands on rocky soils, dry seasonal scrubland of NE Brazil (caatinga) and certain parts of the Amazonian forest; other ecosystems such as the mangrove swamps and the Pantanal wetlands have a more modest presence of bromeliads. Bromeliaceae occupy a wide range of habitats and are one of the most characteristic elements of Neotropical forests. They occur from arid to very wet regions, from sea level to above 4,000 m in the Andes, and from rich volcanic soils to nutrient-poor white-sand savannas. 1,387 species in Brazil, of which 1,218 are endemics.

Herbaceous and evergreen, or rarely shrubby perennials (some Dyckia); terrestrial, saxicolous, or epiphytic; three patterns of growth, sympodial branching with determinant ramets, monocarpy, or monopodial with axillary flowering; stem usually very short and hidden by sheaths, sometimes long (many Tillandsia species) or rarely columnar, gigantic; there are species that live in isolation, or form extensive colonies, dense or sparse clumps, most grow towards the light, but others turn towards the ground (Tillandsia reclinata E.Pereira & Martinelli); roots functional but serving merely as holdfasts in many epiphytic species, reduced both structurally and functionally, or completely lacking; leaves spirally arranged, usually rosulate or distributed along a stem, distichous in few species, in a wide variety of colours (maroon, through shades of green, to gold; varieties may have leaves with red, yellow, white, and cream variations; others may be spotted with purple, red, or cream, while others have different colors on the tops and bottoms of the leaves), shapes, sizes, and textures, simple, with margins entire or spinose-serrate or serrulate, rarely deciduous and heterophyllous (some Pitcairnia species), generally leaf sheath wider than the blade; almost always bearing minute multicellular structures consisting of a stalk and a shield called peltate scales or trichomes, serving to absorb moisture and nutrients, sometimes hairy, or appear to be covered with an epicuticular powder wax; inflorescence terminal, rarely lateral or pseudo-lateral, scapose or sessile, simple or compound, panicles, racemes, spikes, head-like, or solitary pseudo-lateral flowers, usually bearing brightly coloured conspicuous bracts, coloration of inflorescence may fade in a few days, or last for months; flowers diurnal or nocturnal, sessile, or pedicellate, perfect or sometimes functionally unisexual, regular or nearly so, normally trimerous; perianth heterochlamydeous, 3 sepals are twisted left (sinistrorse) and 3 petals twisted right (dextrorse), free or connate forming a basal tube of varying length; sometimes inner surface has double longitudinal ridges, or double strap-shaped appendages (ligules), located at the base of petals, generally membranaceous, or fleshy, and takes a number of shapes; androecium 6 in 2 series, inner facing petals and outer facing sepals; filaments rounded, flattened or thin, free, or joined to petals or to each other; anthers dorsifixed or basifixed, with 2 locules and 4 pollen sacs that open through a longitudinal slit to release pollen; gynoecium a single pistil of a trilocular ovary, trifurcate stigma usually spiral-conduplicate; ovary superior to inferior, 3-carpellate and 3-locular; ovules obtuse to long caudate; placentation axile, extending length of cell or variously reduced. Fruit capsular, septicidal, or indehiscent but then hard and never pulpy or baccate, berries flaccid and often sweet. Seeds winged with entire appendage, plumose with an appendage at base or apex (or both) or naked; embryo small, situated at base of copious mealy endosperm; seeds generally abundant, as well as offshoots (underground rhizomes, aerial stolons); some bromeliads are faintly scented, while others are heavily perfumed.

PSEUDOVIVIPARY is documented for some species of Orthophytum subg. Orthophytum and rarely for Rokautskyia, being always related to taxa with well developed peduncles; this type of propagation is comparatively rare and described for approximately 50 species of aquatic or terrestrial angiosperms, covering families like Agavaceae, Alliaceae, Bromeliaceae, Eriocaulaceae, Poaceae, Polygonaceae, Posidoniaceae, and Saxifragaceae, just to name few; in Bromeliaceae, pseudovivipary is reported for genera like Alcantarea, Ananas, Orthophytum, Tillandsia and Vriesea.

CARNIVORY three spp. of bromeliads belonging to two genera are currently seen to be carnivorous; Brocchinia hechtioides Mez, B. reducta Baker and Catopsis berteroniana (Schult. & Schult. f.) Mez); in both cases, each genus consists of around 20 spp., the overwhelming majority of which are non-carnivorous regular tank bromeliads.

This in itself is unusual since all other genera (except possibly Philcoxia) of carnivorous plants consist exclusively of carnivorous spp. - perhaps this is an indication of a the recent evolution of carnivory among bromeliads; perhaps mistaking the foliage for flowers, visiting insects explore the interior of the plants' rosetes perhaps in search of nectarines. The surface of the leaves of B. hechtioides, B. reducta and C. berteroniana is extremely waxy and very slippery. The UV-reflective white powder that coats the leaves is crumbly and loose and greatly hinders the ability of insects in securing a firm footing; the slightest movement of the plants in the wind or any falter on the part of the insect, causes it to flip and fall into the water filled leaf axils of the bromeliad; trapped by the surface tension of the liquid contained within, the trapped prey is unable to climb up the slippery leaf exists, and eventually drowns. It is not clear the degree to which enzymes are secreted by B. hechtioides, B. reducta and C. berteroniana however at least simple enzymes such as phosphatases are produced directly in the case of B. reducta. Bacteria and various microorganisms assist the digestion process and break down the soft remains of trapped prey releasing nutrients into the liquid contained within the bromeliads' reservoirs; the resultant nutrient soup is absorbed directly by the bromeliads leaves.

USE several bromeliads, particularly spp. of Aechmea, Bromelia, and Greigia, have edible fruits but are consumed only locally. Bromelia spp. are known macambira in Brazil. Aechmea magdalenae (André) André ex Baker, Ananas lucidus Mill., Neoglaziovia variegata (caroá) Mez, and several Bromelia spp., are, or have been, cultivated for the long, strong fibers in their leaves that are used to make hammocks, fishing nets, and twine.

SYSTEMATIC eight subfamilies, Hechtioideae (1/65–70; Texas, Mexico, Central America, with Mesoamerantha and Bakerantha inside the broadest Hechtia) does not occur in South America.

1. SUBFAMILY BROCCHINIOIDEAE (1/17) ‣ a single genus.

1. Brocchinia Schult. & Schult.f. (inc. Ayensua). Rosulate herbs in many adaptations traits, including carnivorous species (B. reducta Baker and B. hechtioides Mez), ant-fed myrmecophytes (B. acuminata L.B. Sm.), tank epiphytes, impounding treelets (B. micrantha (Baker) Mez), N₂-fixers (heterocystous cyanobacteria in its tanks of one terrestrial population of B. tatei L.B.Sm.), and terrestrial forms that absorb nutrients primarily through their roots (mainly). 17 spp., endemic to Guiana Shield, at elevations of 100-2,800 m, from Colombia to Guyana and N Brazil (5, one endemic), almost all species occurring on sand or sandstone on the tepuis or sand plains at low elevations; a few species also occur on granite outcrops at the edge of the Shield. B. tatei L.B. Sm., which can reach enormous dimensions (about 1.5 m in diameter) may be carnivorous; inflorescences of B. micrantha (Baker) Mez (Venezuela and Guyana) is the largest in Guiana Shield, up to 10m tall.

2. SUBAFAMILY LINDMANIOIDEAE (2/41) ‣ both genera endemic to South America.

2. Connellia N.E.Br. Stemless or short-caulescent; leaves rosulate, coriaceous; blades spine-serrate to entire. 6 spp., with larger and more brightly colored petals restricted to Guiana Shield from the tepuis of SE Venezuela and adjacent Guyana, at elevations of 1,300 – 2,800 m, four collected in Roraima state, Brazil, one endemic.

3. Lindmania Mez. Small to large heliophytic herbs; leaves rosulate; flowers white to greenish. 35 spp., endemic to Guiana Shield from Venezuela (35, 28 endemics), Amazonas state in N Brazil (5, none endemics, mainly in Mount Neblina) and Guyana (5, none endemics), mainly in open areas, at elevations of 1,000-2,800 m.

3. SUBAFAMILY TILLANDSIOIDEAE (21/1362) ‣ four tribes, all in South America.

3.1 TILLANDSIOIDEAE ▸ TRIBE CATOPSIDINAE (1/20) - a single genus.

4. Catopsis Griseb. Xeric canopy epiphitics; leaves densely utriculose-rosulate, often cretaceous-coated; scape compiscuous; populations in some taxa are diecious. 20 spp., Florida, Mexico, the Caribbean, Belize, Guatemala and south into Peru, Guyana, Venezuela, and Brazil (only two, very widely, in Bahia, Paraná, Santa Catarina states); only 5 in South America.

3.2 TILLANDSIOIDEAE ▸ TRIBE GLOMEROPITCAIRNIEAE (1/2) – a single genus.

5. Glomeropitcairnia Mez. Stemless, leaves rosulate; scape erect; petals free. Two spp., heliophytic habitats, Lesser Antilles, Trinidad, one in adjacent Venezuela.

3.3 TILLANDSIOIDEAE ▸ TRIBE TILLANDSIEAE (8/942) - outsider Pseudalcantarea (3; Mexico to Nicaragua).

6. Barfussia Manzan. (off Tillandsia) & W. Till. Usually epiphytic, rarely terrestrial herbs, acaulescent, forming impounding rosettes; leaves mesomorphic; leaf sheaths distinct; leaf blades lingulate, acute; inflorescence pink, compound, once or twice branched, rhachis usually alate; floral bracts glabrous; flowers distichously arranged. Three spp., Andes from Colombia to Bolivia.

7. Gregbrownia W. Till & Barfuss. (off. Mezobromelia) Epiphytic or terrestrial herbs, large sized, forming impounding rosettes; leaves mesomorphic; leaf blades lingulate; inflorescence compound, twice or rarely triple branched, a laxly flowered panicle; flowers distichously, rarely spirally arranged. 4 spp., Andes of Peru and Ecuador.

8. Guzmania Ruiz & Pav. Stemless or rarely caulescent; leaves densely polystichous. 205 spp., 184 in South America, mainly mesophytic habits, from Florida to Bolivia, Brazil to Mexico, few spp. in E South America; 9 spp. in Brazil, almost all only in northern Amazon rainforest, G. lingulata (L.) Mez widely distributed in northern country, none endemics.

9. Lemeltonia Barfuss & W. Till. (off Tillandsia) Plants epiphytic herbs, acaulescent or rarely caulescent, forming non-impounding rosettes; leaves usually semi-xeromorphic, rarely mesomorphic or xeromorphic and densely lepidote; leaf blades very narrowly triangular. 7 spp. from Colombia to Ecuador, one up to Peru and L. monadelpha (E. Morren) Barfuss W. & Till in Central America, Trinidad & Tobago, Venezuela, Guianas, Colombia, Ecuador and N Brazil

10. Racinaea M.A. Spencer & L.B. Sm. (inc. Tillandsia p.p.) Plants usually epiphytic or rarely terrestrial herbs, acaulescent or rarely caulescent, forming impounding, or non-impounding, often pseudobulbous rosettes; inflorescence usually compound, once or twice branched, occasionally triple branched, or simple. 80 spp., Andean, mainly Ecuador, extending to Central America, Bolivia, the Greater Antilles and the Guianas, and SE Brazil (3, two endemics); 77 spp. in South America.

11. Tillandsia L. (exc. Barfussia, Cipuropsis, Lemeltonia, Racinaea p.p., Wallisia, inc. Vriesea p.p.) Plants epiphytic or epilithic, rarely terrestrial herbs, much variable in size and habit, caulescent or acaulescent, forming mostly non-impounding to occasionally impounding, sometimes pseudobulbous rosettes; some species in E Brazil grows towards the ground. 673 spp., from Virginia in U.S.A. to Uruguay and Caribbean, highly centered from NW South America and Mexico (254); some spp. has long stems, anothers is rootless, mainly epiphytic, some terrestrial. 417 spp. in South America, 99 in Brazil, 61 endemics, 7 of them, from several states, are rare plants in Brazil, by Plantas Raras do Brasil’s book. Some species in South America are myrmecophytes, unique among this family in continent.

12. Wallisia (Regel) E. Morren. Epiphytic or terrestrial herbs, usually acaulescent, rarely caulescent, forming impounding or rarely non-impounding rosettes; leaf sheaths often silvery grey when dry; leaf blades ligulate to linear-triangular; inflorescence simple or compound, once or rarely twice branched, usually with reduced spikes. 90 spp., Central America and Antilles extending to Ecuador and Bolivia, 26 in South America, very rarely in Brazil (3, one endemic).

3.4 TILLANDSIOIDEAE ▸ TRIBE VRIESEAE (11/398) - two subtribes, both in South America.

■ SUBTRIBE CIPUROPSIDINAE (7/115) ‣ outsider Zizkaea (1-5; Greater Antilles).

13. Cipuropsis Ule. (inc. Tillandsia p.p., Mezobromelia p.p., Vriesea p.p.) Plants epiphytic or terrestrial herbs, acaulescent, forming impounding rosettes; leaves mesomorphic; leaf blades lingulate; inflorescence usually compound, once or twice branched, rarely simple; floral bracts carinate, 1.5–2.5(–3) cm long. 16 spp., northern Andes from Venezuela to Peru, three up Trinidad, Central America and Caribbean, C. rubra (Ruiz & Pav.) Kessous & A.F.Costa up to N Brazil..

14. Goudaea W. Till & Barfuss. (off Vriesea) Epiphytic or terrestrial herbs, usually medium sized, forming impounding rosettes; leaves mesomorphic; leaf blades lingulate, appearing glabrous; inflorescence simple or compound, once branched; floral bracts imbricate, ecarinate, concealing the sepals completely. Two spp., Trinidad & Tobago, Colombia, Ecuador, Peru and Bolivia.

15. Jagrantia Barfuss & W. Till. Epiphytic herbs, medium sized, forming impounding rosettes; leaves mesomorphic; leaf blades lingulate; inflorescence simple; floral bracts imbricate, 3 times the length of the sepals, laterally strongly compressed and sharply carinate, deciduous along a basal transversal line after anthesis when dry. Only one sp., J. monstrum (Mez) Barfuss & W. Till, from Costa Rica to Ecuador.

16. Josemania W. Till & Barfuss. Epiphytic or rarely terrestrial herbs, small to medium sized, forming impounding rosettes; leaves mesomorphic; leaf sheaths very dark, usually becoming silvery-grey when dry, rarely castaneous; leaf blades lingulate. 5 spp., Costa Rica, Panamá, Colombia, Ecuador and Peru; 4 in South America.

17. Lutheria Barfuss & W. Till. Epiphytic or terrestrial herbs, medium to large sized, forming impounding rosettes; leaf blades lingulate, cross-banded or uniformly colored, appearing glabrous; inflorescence simple or compound, once branched; flowers distichously arranged, often second, appearing at one side of the inflorescence. 4 spp., Colombia to French Guiana, Trinidad & Tobago, and N Brazil in Roraima and Pará states.

18. Werauhia J.R. Grant. (inc. Vriesea p.p.) Epiphytic or terrestrial herbs, usually acaulescent, rarely caulescent, forming impounding or rarely non-impounding rosettes; leaf sheaths often silvery grey when dry; leaf blades ligulate to linear-triangular; inflorescence simple or compound, once or rarely twice branched, usually with reduced spikes. 92 spp., Central America and Antilles extending to Ecuador and Bolivia, 27 in South America, very rarely in N Brazil (two spp., bot very widely).

■ SUBTRIBE VRIESEINAE (4/283) ‣ all genera in South America.

19. Alcantarea (E. Morren) Harms. Very large saxicolous rosettes (often reddish leaves), usually reaching 2-5 m high while flowering, mainly in open places and mountains areas. 42 spp. near sea level up to 1,900 m, mainly on gneiss-granitic inselbergs inside the Atlantic rainforest habitats of E Brazil, from Bahia to S. Paulo states, and in on quartzite outcrops in the Espinhaço Range, in the states of Bahia and Minas Gerais; inflorescences of A. imperialis (Carrière) Harms. is the largest in Brazil, up to 5 m tall; 4 spp., in Minas Gerais and Rio de Janeiro states, are rare plants in Brazil, by Plantas Raras do Brasil’s book.

20. Stigmatodon Leme, G.K. Br. & Barfuss. (inc. Vriesea p.p.) Epilithic herbs in vertical, bare granitic surfaces, usually forming inconspicuously impounding or rarely non-impounding rosettes, vegetatively propagating by short basal axillary shoots, or sometimes regular-leaved adventitious offsets produced at the old portion of the stem not covered by leaves and monocarpic, occasionally never forming any shoots or offsets. 33 spp., endemic to SE Brazil, mainly in rock outcrops.

21. Waltillia Leme, Barfuss & Halbritter. Plants terrestrial (including saxicolous) growing along the margins of running streams or in periodically soaked terrains; leaves coriaceous, forming non-impounding rosetes. Two spp., both endemics to Minas Gerais state, with few subpopulations, in an ecologically similar micro-habitat.

22. Vriesea Lindl. (exc. Goudaea, Cipuropsis p.p., Stigmatodon p.p., Werauhia p.p., Tillandsia p.p., inc. Mezobromelia p.p.) Acaulescent, usually epiphytic, leaves rosulate, usually reddish. 233 spp., 227 in South America, E Brazil (196, 184 endemics) extending to NW Argentina, SE Bolivia, Peru (V. oxapampae Rauh, V. ochracea Rauh & E. Gros), Venezuela and the Greater Antilles, some in Central America; 5 spp., from several states, are rare plants in Brazil, by Plantas Raras do Brasil’s book.

4. SUBAFAMILY PITCAIRNIOIDEAE (3/850–855) ‣ all genera occur in South America.

23. Dyckia Schult. f. (inc. Encholirium, Deuterocohnia) Rosette terrestrial spp., leaves spinose serrate, rigid, sometimes ring formations or forming cushions or pollsters; leaves densely rosulate, spinose-serrate, sometimes silvery, highly xeropmorphic, Hechtia-like or Agave-like; some species dioecious. 219 spp., C, E & S Brazil (197; 185 endemics, highly endemics in southern Brazil and Espinhaço Range), N Peru, northern Chile, and center Bolivia to center Argentina, SE Bolivia, over Paraguay and Uruguay, and vast area in N Argentina, mainly xeric in dry areas, some found only in wet areas, in inundated riverisdes and lagoons. 21 spp., from several states, are rare plants in Brazil, by Plantas Raras do Brasil’s book.

24. Fosterella L.B.Sm. Mesophytic terrestrial herbs, stemless or near so; leaves rosulate; blade entire or weakly serrulate. 35 spp., one Mexico to El Salvador, and 34 from center Peru to N Argentina and Paraguay (the bulk of genus endemic to Bolivia) and Brazil (8, 5 endemics); Brazilian species are mostly from Mato Grosso and Mato Grosso do Sul states, but F. batistana Ibisch, Leme & J. Peters occurs in high dense forest in Amazonas and Pará states and F. atlantica Leme & Forzza is endemic to Minas Gerais state.

25. Pitcairnia L'Hér. Mostly terrestrial herbs but sometimes saxicolous or epiphytic; mostly stemless to long-caulescent; leaves usually strongly dimorphic, sometimes deciduous and petiolate (unique among some Bromelia, Disteganthus and Cryptanthus in family); inflorescences raceme to panicle. 409 spp., high diverse habitats, from Mexico and Caribbean to Brazil (54, 45 endemics) and Africa (1), 331 in South America; one sp., from Minas Gerais state, is a rare plant in Brazil, by Plantas Raras do Brasil’s book.

P. feliciana (A. Chev.) Harms & Mildbr in center Guiné, Africa, one the most remarkable disjuncions in PLANT KINGDOM, found in some 100 km from the coast, at the base of Mt. Gangan near Kindia, in a few square km in size, near the side of the primary slope, grouped in two subpopulation, one a few km from Kindia, on the edge of the road to Telimele, the other in the gorges of the Samou River.

P. azouryi Martinelli & Forzza can be differentiated from all other Brazilian species of the genus by possessing large flowers (10-18 cm long), pale green, thus cryptic, however. P. encholirioides L.B. Sm. of SE Brazil has the unique bulb (succulent leaves under dry leaves in basal part of plant) of family.

5. SUBAFAMILY NAVIOIDEAE (5/c 105) ‣ capsular fruits; seeds winged to naked; petals minute; sepals cochlear, with the two adaxial overlapping the abaxial; stellate chlorenchyma absent; leaves entire, serrulate, or spinose, but not succulent; inflorescence paniculate to capitate. Included genera: Brewcaria, Cottendorfia, Navia, Sequencia, Steyerbromelia

26. Brewcaria L.B.Sm., Steyerm. & H.Rob. Large heliophyitc herbs; leaves rosulate, blades narrowly triangular, spinose serrate. 6 spp. from Venezuela, one up to Colombia.

27. Cottendorfia Schult.f. Heliophytic herbs; leaves rosulate; blades entire, pungent; inflorescence tripinatelly paniculate, lax; dioecious. Only one sp., C. florida Schult. f., restricted of xeric habitats in Bahia, Piauí and Paraíba states, Brazil.

28. Navia Schult. & Schult.f. 97 spp., the largest genus endemic to Guiana Shield, ranging from Colombia to Suriname and northern Brazil (7, 4 endemics, three of then only in summit of Mount Aracá), at elevations of 100-2,500 m, mostly endemics to a single summit; 68 endemics of Venezuela.

29. Sequencia Givnish. Herbs; leaves linear, to roughly 1.8 m long, with dark curved spines near the base and serrulate toward the tip, and arranged in a rosette about a twisted, prostrate stem; inflorescence amply paniculate; flowers perfect. Only one sp., S. serrata (L.B. Sm.) Givnish, known only from low sandstone mesetas (Cerro de Circasia, Cerro Yapoboda, Cerro de Canenda) in Vaupes region in SE Colombia.

30. Steyerbromelia L.B.Sm. Large heliophytic herbs, leaves rosulate. 9 spp., six in mountains areas of 600-2,700m in s Venezuela and N Brazil (only S. plowmanii (L. B. Sm., Steyerm. & H. Rob.) B. Holst, in Mount Neblina), in Guiana Shield, in open summits, and three restricted sandstone mountains in E Colombia.

6. SUBFAMILY PUYOIDEAE (1/228) ‣ a single genus.

31. Puya Molina. Stemless or more generally long-caulescent, up to 7m tall, all terrestrial; leaves coriaceous; blades never constricted at base, coarsely spinose-serrate. 230 spp., one only in Costa Rica and Panamá, 229 in South America (one also in Central America), 227 in open mountains areas in Andes from NW Venezuela to NW Argentina, two in Guiana Shield, with P. floccosa (Linden) E. Morren ex Mez, found in Tepequen Range in N Roraima state, Brazil. P. raimondii Harms (Bromeliaceae) from mountains of Peru and Bolivia is the most massive inflorescence of the Earth, with 8-12m tall, and have a diameter of up to 2.4m; extra-large specimens can grow as tall as 15m; the inflorescence can bear approximately 8,000 small white flowers. P. alpestris (Poepp.) Gay endemic to Chile and Schwartzia brasiliensis (Choisy) Bedell ex Gir.-Cañas. (Marcgraviaceae) endemic to E Brazil are the unique blue-nectar plants known.

7. SUBAFAMILY BROMELIOIDEAE (33/765–770) ‣ fruits indehiscent, baccate; genera endemic to Brazil ranges from dry-scrub vegetation (Neoglaziova) up to nebulous high mountains (Fernseea); seven informal but monophyletic lineages: (Bromelia + (Greigia Clade + (Fernseea + (Acanthostachys + (Ananas Clade + (Neoglaziova + Core Bromelioid)))))). Aechmea marie-reginae H. Wendl. and the two Androlepis, all from Mesoamerica, have unisexual flowers and are the unique Bromelioideae with this characteristic. Outsiders Androlepis (2; southern Mexico to Nicaragua) and Hohenbergiopsis (1; Mexico to Central America).

■ BROMELIA CLADE (1/69) ‣ a single genus.

32. Bromelia L. Coarse terrestrial or rarely epiphytics, spreading by rhizomes; leaves coarsely spinose along the margins, sometimes petiolate (unique among some Disteganthus, Cryptanthus and Pitcairnia in family); sepals with soft usually broad apices; inflorescences compound; sepals mostly free or nearly so. 69 spp., mostly in dry open ground, but sometimes submesophytic, Central and South America (65), mainly in Brazil (52, 41 endemics), in dry savanas has the highest diversity with about 30 species, in contrast to the general rule for the subfamily, which reaches its highest diversity in the Atlantic Forest; one sp. from Bahia state is a rare plant in Brazil, by Plantas Raras do Brasil’s book.

■ GREIGIA CLADE (4/41) ‣ all genera in South America.

33. Deinacanthon Mez. Only one sp., D. urbaniana (Mez) Mez., from Paraguay to N Argentina and Bolivia.

34. Fascicularia Mez. Terrestrial or rarely tree thunks at tall trees, stemless; leaves very densely rosulate, linear, some reddish, petals white. Only one sp., F. bicolor (Ruiz & Pav.) Mez, endemic to Chile.

35. Greigia Regel. More or less caulescent; leaves imbricate along the stem, partially or completely serrate. 35 spp., saxicolous in open ground or terrestrial in forest paramo, Mexico to Panamá, Venezuela to Bolivia, Chile and Juan Fernandes Is; 30 spp. in South America.

36. Ochagavia Phill. Suffruticose, amply proliferating; leaves thunk-forming. 4 spp., Juan Fernandez and Chile.

■ FERNSEEA CLADE (1/2) ‣ a single genus.

37. Fernseea Baker. Stemless; leaves forming a dense pseudobulb, leaves apparently rigid; inflorescence panicles; sepals narrowly elliptic, 7mm long; flowers subsessile or pedicellate. Two spp. restricted of two small high nebulous peaks to coast areas in E Brazil, SE Minas Gerais, NE Sao Paulo and SW Rio de Janeiro, one of them, from São Paulo state, is a rare plant in Brazil, by Plantas Raras do Brasil’s book.

■ ACANTHOSTACHYS CLADE (1/2) ‣ a single genus.

38. Acanthostachys Klotzch. Rhizomatose, epiphytic or saxicolous; leaves long and narrow, pendulous; scape well developed or lacking. Three spp., E Brazil (all species, two endemics), Paraguay, NE Argentina.

■ ANANAS CLADE (2/6 + 2; Aechmea fernandae (E.Morren) Baker from Peru and Brazil, and and Aechmea tayoensis Gilmartin from Ecuador belongs this clade)

39. Ananas L. (inc. Pseudananas) Rosettes, not producing stoons, or producing; leaves rosulate; inflorescense densely strobiliform, usually crowned with a tuft of sterile foliaceous bracts, often producing slips at the base. Two spp., A. macrodontes Morren from Bolivia, Paraguay, Argetina and S Brazil, and A. comosus (L.) Merrill, a seedless cultigen (a plant that has been altered by humans through a process of selective breeding); because it has been in cultivation for thousands of years, its exact origins are unknown.

The pineapple (A. comosus (L.) Merr.) is the third most important tropical fruit in world production after banana and citrus; cultivated pantropically since the 1500s for its edible fruits, is by far the most commercially important spp. of Bromeliaceae; the protein-digesting enzyme, bromelain, is extracted from pineapple fruits for use in meat tenderizers and as an anti-inflammatory.

Within A. comosus, five botanical varieties are recognized, two wild [A. comosus var. microstachys (formerly A. comosus var. ananassoides) and A. comosus var. parguazensis] and three domesticates [A. comosus var. comosus (the edible pineapple), A. comosus var. erectifolius (the curagua, a fiber crop) and A. comosus var. bracteatus, now an ornamental pineapple].

40. Disteganthus Lem. Herbs, rosettes, caulescent or acaulescent, propagating by stolons or rhizomes; leaves sometimes petiolate (unique among some Bromelia, Cryptanthus and Pitcairnia in family), armed. 4 spp., swampy forests, Suriname, French Guiana, and adjacent Amapá state in N Brazil (only D. basilateralis Lem., no endemic).

■ CRYPTANTHOID COMPLEX (9/166) ‣ a fully Brazilian group.

41. Cryptanthus Otto & A. Dietr. Terrestrial or saxicolous, andromonoecious, usually stemless or sometimes caulescent, propagating by short axillary shoots or long and slender stolons or rhizomes, leaves sometimes petiolate (unique among some Bromelia, Disteganthus and Pitcairnia in family). 65 spp., Rio de Janeiro, Minas Gerais and Espírito Santo to the NE states of Bahia, Sergipe, Alagoas, Pernambuco, Paraíba and Rio Grande do Norte; 4 spp. are rare plants in Brazil, by Plantas Raras do Brasil’s book.

42. Forzzaea Leme, S. Heller & Zizka. Saxicolous, homogamous, stemless, propagating by short axillary shoots or rarely stoloniferous, then bearing thick stolons, clustering or solitary; leaves succulent, thick coriaceous, forming non impounding rosetes. 8 spp., endemic to the W-NE part of the meridional highs of the Espinhaço Range in Minas Gerais state; one sp. is a rare plant in Brazil, by Plantas Raras do Brasil’s book.

43. Hoplocryptanthus Leme, S. Heller & Zizka. Saxicolous or epilithic, homogamous, stemless or long caulescent, propagating by short axillary shoots; leaves coriaceous, forming non-impounding rosetes. 9 spp., endemic to the Minas Gerais state, growing mostly in the Iron Quadrangle (Quadrilátero Ferrífero) region, which is a well known Precambrian terrain with approximately 7,000 km² and significant mineral resources, particularly iron and gold.

44. Krenakanthus (Leme, S. Heller & Zizka) Leme, Zizka & Paule (off Orthophytum). Plants terrestrial, monoecious, long caulescent, flowering 30–60 cm tall, propagating by elongated shoots developing near the base of the inflorescence. Two spp. known from E Minas Gerais state; plants terrestrial, forming large and dense groups in organic-rich, shallow soils accumulated on rocky surfaces in shady spots inside fragments of humid Atlantic Forest.

45. Lapanthus Louzada & Versieux. Herbs rupicolous, stoloniferous, stem an inconspicuous rhizome, covered by the leaf sheaths; leaves rosulate, arcuate; inflorescence sessile, compound with two flowers per branch or pseudosimple; flowers with a tubular corolla; sepals white-hyaline, elliptic to triangular, inconspicuous, reaching ca. 1/3 of the flower length; petals free, white-hyaline or orange-yellow. Three spp. endemic to Minas Gerais state, in transitional vegetation between semideciduous seasonal forest and campo rupestre in the Serra do Cipó area or in humid quartizitic rock outcrops near to waterfalls and gallery forest, between 700–900 m, two of them are rare plants in Brazil, by Plantas Raras do Brasil’s book.

46. Orthocryptanthus (Leme, S. Heller & Zizka) Leme, Zizka & Paule (off Orthophytum). Plants saxicolous, monoecious, usually long caulescent or sometimes stemless, propagating by short axillary shoots or stolons. Two spp. from Minas Gerais state, in the Atlantic Forest domain; saxicolous or epilithic, living on rocky outcrops in open to partially shaded sites, 1,000 to 1,440 m elevation.

47. Orthophytum Beer. (exc., Krenakanthus, Orthocryptanthus, Sincoraea, Siqueiranthus) Rosette or prostrate (sometimes erect) rupicolous, saxicolous or sometimes terrestrial herbs with grater patterns of leaves colour; floral bracts leaf-like, petals with reflexed lobes. 64 spp. endemic to NE and SE Brazil, in areas of soil on top of granitic-gneiss inselbergs (Atlantic Rainforest and dry seasonal scrubland of NE Brazil (caatinga), from Paraíba to Espírito Santo states) or quartzite-sandstone in rocky grasslands (campos rupestres, Espinhaço RANGE, in Bahia and Minas Gerais) forming mats or sometimes as isolated individuals.

§ subg. Capixabanthus ‣ 8 spp., Espírito Santo state and in neighboring areas in Minas Gerais (Nanuque) and Bahia (Lagedão), in the Atlantic forest domain, on granitic inselbergs in low elevated flatlands, or on shallow soils in shaded sites of mountainous areas to 600 m elevation; one sp. is a rare plant in Brazil, by Plantas Raras do Brasil’s book.

§ subg. Clavanthus ‣ 9 spp. endemic to the Espinhaço Range in Minas Gerais state, growing in the southern part of the range and extending through the septentrional highs near the border with Bahia state, at altitudes of 650 to 1,300 m; two spp. are rare plants in Brazil, by Plantas Raras do Brasil’s book.

§ subg. Orthophytum ‣ 48 spp., Ceará and Rio Grande do Norte to Espírito Santo and Minas Gerais states; Atlantic Forest, rocky grasslands (campos rupestres) and dry seasonal scrubland of NE Brazil (caatinga); low to médium altitude (to 500 m; few spp. higher than 600-1,200 m); axillary shoots, stolons/rhizomes & pseudovivipary.

48. Rokautskyia Leme, S. Heller & Zizka. Plants terrestrial or saxicolous, homogamous, stemless to long caulescent, propagating by short axillary shoots, long and slender stolons or rarely pseudoviviparous with shoots developed on the inflorescence. 14 spp., an exclusive dweller of the hygrophilous Atlantic Forest of the montainous central region of Espírito Santo state.

49. Sincoraea Ule. (off Orthophytum) Plants stemless; primary bracts and the basal portion of the inner leaves turning white, yellow or red, forming a colorful ring around the inflorescence in contrast with the color of the distal portion of leaves. 11 spp., endemic to the rocky grasslands (campos rupestres) of the septentrional highs of the Espinhaço Range, with most species found in the state of Bahia and a single species in the north of Minas Gerais state; three Minas Gerais’s species and six from Bahia are rare plants in Brazil, by Plantas Raras do Brasil’s book.

50. Siqueiranthus Leme, Zizka, E.H. Souza & Paule. (off Orthophytum) Andromonoecious, saxicolous, long caulescent usually by the successive growth of a single apical shoot, stems 18–160 cm long, 1.7–2.8 cm in diameter, rigid, erect to prostrate, propagation by a single shoot at the base of the inflorescence and by few shoots along the stem, rhizomes not detected. Only one sp., S. cinereus (D.M.C. Ferreira & Louzada) Leme, Zizka, E.H. Souza & Paule, known from the type population, as a saxicole, growing at the top of a granitic outcrop in the Atlantic Forest domain in the municipality of Ibateguara, Alagoas state, Brazil.

■ NEOGRAZIOVA CLADE (1/3) ‣ a single genus.

51. Neoglaziovia Mez. Terrestrial and saxicolous herbs; stemless; leaves few; inflorescense simple, racemose, flowers pedicellate. Three spp. endemic to dry areas in NE Brazil, used for their fibrers.

■ HOHENBERGIA CLADE (2/57) ‣ both genera in South America.

52. Hohenbergia Schults & Schults f. (exc. Wittmackia p.p.) Stemless, leaves rosulate; blades spinose-serrate; scape well developed; inflorescenses of strobilaceous spikes. 52 spp., 50 endemics to E Brazil (5 of them, all in Bahia state, are rare plants in Brazil, by Plantas Raras do Brasil’s book), H. stellata Schult. & Schult.f. in Trinidad-Tobago to Venezuela, Curaçao, Martinique and NE Brazil, and H. andina Betancur endemic to Colombia.

53. Karawata Schults & Schults f. Epiphytic, terrestrial or saxicolous, shortly rhizomatous; leaves spiraled, erect; leaf-sheaths elliptical to triangular, castaneous, contrasting in colour to the leaf-blades, margins entire, imbricate, tank-forming; leaf-blades green, both sides with argenteous indument, apex pungent; inflorescence terminal, polytelic, simple strobilioid, ovate or capituliform. 7 spp. endemic to the Brazilian Atlantic Forest, from Pernambuco to Rio de Janeiro states.

■ CORE BROMELIOIDEAE ‣ Aechmea rubiginosa Mez (Venezuela, Colombia, Ecuador, Brazil) is sister of all remaining sepcies in this group. Androlepis Alliance is composed of an assemblage of morphologically dissimilar species, which have been classified in previous treatments within six genera or subgenera: Aechmea subg. Podaechmea Mez, A. subg. Pothuava (Baker) Baker, Androlepis Brongn. ex Houllet, Billbergia Thunb., Hohenbergiopsis L.B. Sm. & Read and Ursulaea Read & Baensch.

54. Aechmea Ruiz & Pav. (inc. Ursulaea) Terrestrial or epiphytic spp., acaulescent, frequently propagating by rhizomes; leaves usually forming a tank, usually spinose-serrate. 277 spp., from Mexico to South America (260), highly diverse in Brazil, with 181 spp., 153 endemics, 10 of them, in several states, are rare plants in Brazil, by Plantas Raras do Brasil’s book. This genus has sixth largest diversity of variegated leaves worlwide (36).

55. Araeococcus Brongn. (exc. Pseudaraeococcus) Stemless; inflorescence laxlay paniculate, mesophytic epyphites; flowers perfect, petals naked. 6 spp., 5 in Brazil (none endemics) up to Venezuela, Guianas, Colombia and Bolivia, and one from Costa Rica to N Colombia.

56. Billbergia Thunb. Terrestrial, saxicolous or epiphytic herbs; stemless; leaves forming broadly funnelform to long-tubular rosettes; scapebracts large, mostly red; petals zygomorphic or tightly recoiled and flowers sessile, epigynous tube usually well developed. 66 spp., Mexico, Central America, along the Atlantic coast, through Uruguay, and as far south as Argentina, mainly in Brazil, and along the Pacific coast through Ecuador and Peru; 64 spp. in South America, 49 in Brazil, 37 endemics, one of them, from Bahia state, is a rare plant in Brazil, by Plantas Raras do Brasil’s book.

57. Canistrum E. Morren. (inc. Wittrockia p.p.) Mostly epiphytic; leaves rosulate, armed; scape evidente; inflorescence compound, densely corymbose, involucrate with the bracts covering all but the apices the flowers. 20 spp. endemics to Atlantic Forest in E Brazil, 6 of them, from several states, are rare plants in Brazil, by Plantas Raras do Brasil’s book.

58. Edmundoa Leme. 3 spp., endemic to E Brazil.

59. Eduandrea (Mez) Leme, W. Till, G.K. Br., J.R. Grant & Govaerts. Hygrophilous perennial herbs, small-sized, terrestrial, propagating by long and stout, underground rhizomes; leaves forming a very narrow funnelform rosette with limited water holding capacity; inflorescence tripinnate, densely corymbose, distinctly elevated above the leaf-rosette, inconspicuously involucrate; flowers perfect, sessile or subsessile, odorless. One rare sp., E. selloana (Baker) (Mez) Leme, W. Till, G.K. Br., J.R. Grant & Govaerts, endemic to the gallery forests (white quartz sandy soils, often between rocks) that transverse the rocky grasslands (campos rupestres) in the regions of Ouro Preto and adjacences of Minas Gerais state.

60. Hylaeicum (Ule ex Mez) Leme, Forzza, Zizka & Aguirre-Santoro. (off Neoregelia) Plants epiphytic, homogamous, stemless, but offset distinctly stoloniferous; stolons slender to stout, 10–40 × 0.3–2 cm, covered by distinct cataphylls; leaves coriaceous or thick-coriaceous, forming water impounding rosettes. 12 spp., Venezuela to Peru and NW Brazil (5, 2 endemics).

61. Lymania R.W. Read. Slenderly stoloniferous or stout-rhizomatous; leaves few, froming a crateriform or utriculate rosete, minutely serrate; inflorescense variable. 10 spp. from E Brazil.

62. Neoregelia L. B. Sm. Terrestrial, saxiculous or epiphytic rosette herbs, usually spinose serrate; this genus has the third world diversity of variegated leaves (62); inflorescence sunk in the center of the rosete, very densely captiform. 114 spp. endemics to Brazil, most of them grow in sandy soils, 10 of them, in several states, are rare plants in Brazil, by Plantas Raras do Brasil’s book; the largest national endemic genera in new World.

63. Nidularium Lem. (inc. Canistropsis) Epiphytc or terrestrial herbs, stemless; leaves forming funnelform tanks, serrate or serrulate, petals erect and apex distinctly obtuse cucullate, connate or agglutinated in a tube the height of the sepals. 56 spp. endemic to Atlantic Forest from Bahia to Rio Grande do Sul states in E Brazil; the largest genus endemic to Atlantic Forest; 16 spp., in several states, are a rare plant in Brazil, by Plantas Raras do Brasil’s book.

64. Portea Brogn. ex. K. Koch. Terrestrial, saxicolous or epiphytic; leaves rosulete, serrate; scape bracts bright colored; inflorescence compound, sepals more or less connate, long-mucronate; petals appendaged. 8 spp. from forets of Atlantic Forest, E Brazil.

65. Pseudaraeococcus (Mez) R.A.Pontes & Versieux. (off Araeococcus) Epiphytic, eventually terrestrial or saxicolous, propagating by thin stolons, usually long, and involving the phorophytes; rosette lageniform or infundibuliform, tank conspicuous; leaves of 3 to 10, concolor or discolor, chartaceous. 6 spp. endemics to Atlantic Forest of NE Brazil, almost all restricted of Bahia state except some records in Pernambuco and Alagoas states.

66. Quesnelia Gaudich. Terrestrial, saxicolous or epiphytic spp., stemless or rarely long-stemmed; leaves serrulate; scape developed; inflorescence usually simple, flowers sessile. 24 spp. from Santa Catarina to Bahia states, with diversity centers in the Rio de Janeiro coastal region and the rainforest of S Bahia; one sp., from São Paulo state, is a rare plant in Brazil, by Plantas Raras do Brasil’s book.

67. Ronnbergia E. Morren & Andre. (exc. Wittmackia p.p.) Epiphytic spp.; leaves fasciculate or rosulate, entire or serrulate. 23 spp. from NW & W South America, some up to Costa Rica

68. Wittmackia Mez. (inc. Ronnbergia p.p., Hohembergia p.p.) Terrestrial, rupicolous or epiphytic, cespitose or solitary, stoloniferous or acaulescent; rosette broad, forming phytotelmata. 46 spp., 28 endemics to E Brazil, 16 only in Caribbean (11 endemics to Jamaica), one in Mexico and W. lingulata (L.) Mez widely from Costa Rica to Brazil, Caribbean.

LINEAGE 2 of 6: TYPHACEAE

TYPHACEAE

§ FULLY CHLOROTROPHIC

Genera/spp. 2/62 Distribution Typha (40) is cosmopolitan except polar areas, although most frequent in the Northern Hemisphere; Sparganium, 15 spp., is distributed in extratropical parts of the Northern Hemisphere, 1-2 spp. occurring in SE Asia, New Guinea, SE Australia, and New Zealand. Habit Monoecious, perennial herbs. Aquatic or helophytic. Rhizome rich in starch. Two genera in Neotropics.

SYSTEMATIC both genera in South America.

1. Sparganium L. 20 spp., temperate and arctic regions, one or two species in SE Asia, Malesia, New Guinea, Australia, the S Hemisphere south to New Zealand; 9 spp. in New World, one in South America, S. americanum Nutt., C & E Canada to C & E U.S.A., Mexico (Sinaloa, Durango), disjunct in Colombia.

2. Typha L. Perennial tall plants with starchy rhizome, emergent from shallow water or growing in wet soil, sometimes completely submersed and floating, with typical cylindrical-brownish inflorescence at the end of a long and unbranched axis or scape; sometimes vast marsh perennial with sheating leaves. 42 spp., 4 in New World, widely distributed in lagoons, rivers and wet areas, submerged; cccurs in almost all considerable watercourses and lakes of neotropics, in few members to gigantic-vast-dominat populations. often forms extensive monospecific stands; like the other species, it is a rapid colonizer of disturbed, polluted or newly formed wetlands; 3 spp. occur in Brazil, all widely distributed; T. subulata Crespo & R.L. Pérez-Mor. is restricted from Argentina to Uruguay.

LINEAGE 3 of 6: RAPATEACEAE

RAPATEACEAE

§ FULLY CHLOROTROPHIC

Genera/spp. 17/95 Distribution tropical South America, with their highest diversity in the Guiana Shield; Epidryos distributed northwards to Panamá; Maschalocephalus in tropical W Africa (Sierra Leone and Liberia). Habit bisexual, perennial herbs. Usually hygrophytic (a few spp. are epiphytic). Rapateaceae is restricted to the Neotropics with the exception of Maschalocephalus dinklagei Gilg & Schumann that occurs in W Africa. The center of spp. diversity of Rapateaceae are the lowlands Guiana Shield and the Amazon rainforest.

SYSTEMATIC 4 clades, all in South America.

1. SPATHANTHUS CLADE (1/2) ‣ a single genus.

1. Spathanthus Desv. Herbs with leaves distichous; inflorescence 1-several, solitary in leaf axills, spicate. Two spp., Guyana, Venezuela, N Brazil (both Brazil, none endemics), Colombia and French Guiana; in stream-and riverbanks, open inundated forests, and in scrub, north-amazonic white-sandy savannas (campinaranas).

2. SUBFAMILY RAPATEOIDEAE (2/31) ‣ all genera occur in South America.

2. Cephalostemon Schomb. Herbs, leaves gramineous; inflorescence subglobose to cilyndrical, appendaged seeds. 5 spp., in Amazonian savannas, north-amazonic white-sandy savannas (campinaranas), similar habitats in Brazilian Shield of Brazil (all species), Colombia, Guyana, Suriname and Venezuela, mainly in lowland and upland sites, mainly on open, herbcovered sites on wet, extremely infertile sands; C. riedelianus Korn only occurs in the campo rupestre of Minas Gerais.

3. Duckea Maguire. Herbs, leaves gramineous; inflorescence subglobose to cilyndrical, nonappendaged seeds. 4 spp. in Venezuela, Colombia and Brazil (3, none endemics).

4. Rapatea Aubl. herbs, rhizomes short; leaves dorsiventrla to ensiforme; inflorescence many each 2 involucral bracts. 22 spp., mucky, inundated forests of northern South America, into the Amazon rainforest, coastal Brazil (Bahia), and Pacific Colombia, and Bolivia, Ecuador, Guyana, French Guiana, Peru, Suriname and Venezuela. 13 spp. in Brazil, two endemics.

3. SUBFAMILY MONOTREMATOIDEAE (4/7) ‣ outsider is Maschalocephalus (1; Guinea, Sierra Leone, Liberia, Ivory Coast; most probably due to late long distance dispersal).

5. Monotrema Koern. Herbs gramineous, leaves linear; inflorescence axillary, 1-2 per leaf axill, monocephalous, small, subglobose, 1-2 bracteate. 4 spp., 3 endemic to the Guiana Shield from Amazonian savannas, open inundated forests and scrub of Brazil, Colombia and Venezuela, and M. aemulans Körn. extending up to southern Amazon rainforest of Brazil (two spp., none endemics).

6. Potarophytum Sandw. Herbs, rhizomes inbranched, inflorescence 1-2 axillary, shorter than the leaves, with prophylloid bracts. Only one sp., P. riparium Sandwith, endemic to the Guiana Shield of streamside forests on white quartz gravel and sand of Kaieteur Mountain Range in Guyana, at 100-400 m elevation range.

7. Windsorina Gleason. herbs, penduncles 3-9 per leaf axills, each penduncle subentended by 1-2 prophyllous bracts; inflorescence cymose; petals free. Only one sp., W. guianensis Gleason, endemic to the Guiana Shield of in inundated forests of Potaro river in Guyana, at 50-300 m elevation range.

4. SUBFAMILY SAXOFRIDERICIOIDEAE (9/54) ‣ all genera occur in South America.

8. Amphiphyllum Gleason. Herbs, leaves equitant; thyrse bractes several, the outer 2 rigid, more or less valvate and connate. Only one sp., A. rigidum Gleason, endemic to the Guiana Shield of in tepui bogs and meadows of Cerro Duida and Cerro Marahuaca in Venezuela, 1,200-2,000 m elevation range.

9. Epidryos Maguire. Herbs, epiphityc or occasionally terrestrial herbs; axillary penduncles numerous. 4 spp., E. guayanensis Maguire and E. matheusii Sch. Rodr. ex A.S. Flores from Venezuela, Brazil (only the former, at known only by a single collection in the summit of Mount Caburaí) and Guyana; E. allenii (Steyerm.) Maguire endemic to Panamá; and E. micrantherus (Maguire) Maguire in Colombia and Ecuador.

10. Guacamaya Maguire. Herbs, leaves glacous; penduncles axillary; inflorescence hemisphaerical, solitary. Only one sp., G. suberpa Maguire, endemic to the Guiana Shield, known from a small area of Amazonian savannas along the Guainía and Atabapo in SE Colombia and SW Venezuela, 100-200 m elevation range.

11. Kunhardtia Maguire. Herbs, penduncles solitary; inflorescence spheroid or hemisphaerical. Two spp., endemics to the Guiana Shield of tepui bogs and meadows in Sierra Maigualida, Sipapo, Autana, Cuao, Guanay, Yutaje, Coro-Coro and nearby lowlands in northern venezuelan Amazonas, 100-2,100 m elevation range.

12. Marahuacaea Maguire. Herbs, leaves auriculate; inflorescence many, axillary. Only one sp., M. schomburgkii (Maguire) Maguire, endemic to Pantepui Life Zone, in tepui bogs and meadows of Cerro Marahuaca in Gran Sabana, Venezuela, at 2,400-2,800 m elevation range.

13. Phelpsiella Maguire. Small herbs; leaves linear, penduncles subterminal, solitary; inflorescence small, compressed. Only one sp., P. pterocaulis Maguire, endemic to the Guiana Shield of from tepuis bogs and meadows of Cerro Paru in Venezuela, at 1,300-2,000 m elevation range.

14. Saxo-fridericia Schomb. Herbs, usually robust; inflorescence globbose or hemisphaerical, spikelets numerous. 10 spp., almost endemic to the Guiana Shield (Colombia, French Guiana, Guyana, Suriname, and Venezuela), except one which occurs in C Brazil (6, S. brasiliensis P.E. Berry & Krahl endemic, known only from white-sand campina vegetation north of Manaus along the Caracaraí highway in Amazonas State), in ecotones between tepui bogs/meadows and surrounding scrub, and between riverbanks and lowland inundated forests; peat pockets over granitic outcrops of Brazil.

15. Schoenocephalium Seub. Herbs, sheats partially open; spikelets radiate. 4 spp., Amazonian savannas on white sand in nearby portions of W Amazonas of Venezuela, and adjacent Colombia (all four species) and Brazil (two spp., none endemics), in the drainages of the Orinoco, Guainia, Atabapo, Ventuari, Inírida, Vaupés, Caquetá, and Rio Negro, 100 – 800 m elevation range.

16. Stegolepis Klotzsch. Herbs, caudex unbranched; leaves gramineous or ensiform; penduncles several, axillary, 1-3 m long; heads 1-many spikelets. 33 spp. endemic to the Guiana Shield of tepuis and meadows of Brazil (5, one endemic), Guyana (6, one endemic) and Venezuela (31, 25 endemics), 500-2,800 m elevation range.

LINEAGE 4 of 6: JUNCACEAE/CYPERACEAE/THURNIACEAE

THURNIACEAE

§ FULLY CHLOROTROPHIC

Genera/spp. 2/4 Distribution NE South America (mainly the Guiana Shield, N Brazil), S South Africa (W Cape province, Pondoland and southern Kwazulu-Natal). Habit bisexual, perennial herbs. Graminids. Often helophytic or aquatic. Culm angular in cross-section, erect.

Key differences from similar families

1. Perianth present; fruit a capsule - 2

1. Perianth mostly lacking or if present consisting of hairs, scales or bristles; fruit a nutlet ------------ Cyperaceae

2. Inflorescence a dense and large globose head on a long peduncle; leaf margins serrate ------------ Thurniaceae

2. Inflorescence differently shaped; leaf margins never serrate ------------ Juncaceae

SYSTEMATIC outsider Prionium (1; W and E Cape, S Kwazulu-Natal).

1. Thurnia Hook. f. Large perennial rhizomatous aquatic herbs, resembling some Cyperaceae; leaves all basal, V – or M – shaped in cross section; inflorescence globose, subtented by leafy bracts. Three spp., T. jenmanii Hook. f. is a narrow endemic species found only in Suriname, T. polycephala Schnee occurs in Venezuela and Guyana while T. sphaerocephala (Rudge) Hook. f., occur from Colombia to French Guiana, and Amazon rainforest in Brazil, also outside Guiana Shield, reaching in some places in C Brazil, in or along creeks, on white sand, in dense canopy forests, in habitats flooded by acidic (black) waters such as streams and lakes in the Amazon rainforest; seeds are dispersed by fishes.

JUNCACEAE

§ FULLY CHLOROTROPHIC

Genera/spp. 8/440-450 Distribution cosmopolitan, with their highest diversity in cold and temperate regions in the Northern and Southern Hemispheres. Habit usually bisexual (rarely monoecious, dioecious or gynodioecious), usually perennial (sometimes annual) herbs. Graminids. Many spp. are aquatic or hygrophytic. Culm terete or flattened in cross-section, smooth or with longitudinal ridges and furrows.

Juncaceae are native in alpine meadows and in grasslands, especially in the highlands, where they may be weedy. Juncus is most common in south Brazil, some endemic spp., the remaind are widely distributed. Some are endemic to the Andes (all Distichia, Oxychloe, Patosia and spp. of Juncus and Luzula), whereas other are widely distributed, often temperate spp. in the tropics members of the family are often found above (800-)-1,500 m above msl.

SYSTEMATIC all genera occur in South America.

1. Distichia Nees & Meyen. Cushion-forming pickly herbs to several meters in diam.; dioecious, flowers lateral, in axills of foliage leaves. Three spp. from high Andes from Colombia to N Argentina, one endemic to Ecuador.

2. Juncus L. Herbs, perennials and rhizomatous or occasionally annual, mainly halophytes; leaves spiral, fully glabrous; inflorescence few-to-many flowered, separate or congested in headlike clusters. c. 250-275 (up to 315 in some autors) spp., cosmopolitan; 131 spp. in New World, 47 in South America, 22 in Brazil, mainly southern region with some of them occurring at high altitude areas such as the Mantiqueira Range, which is located along the border of the states of São Paulo, Rio de Janeiro and Minas Gerais, 6 endemics; 10 sections worldwide, 7 of then in Brazil; outsiders Brazil are a North American and Asian, one only in semi-polar areas in high altitudes in N & S hemisphere, and another in Mediterranean Basin.

Juncus in the current circumscription is heavily non-monophyletic, with the two species forming a sister-group to the remaining Juncaceae. Luzula was sister to a main clade comprising Juncus pro parte, Distichia, Marsippospermum, Oxychloe, Patosia, and Rostkovia. Furthermore, J. capensis and J. lomatophyllus formed a sister-group to Oxychloe, Distichia and Patosia, where Distichia was paraphyletic relative to Patosia. Additional molecular analyses, including even more taxa of Juncus, are desirable.

3. Luzula L. Herbs, perennials, rarely annual rhizomatous; leaves spiral, margin ciliate; inflorescence many flowered, paniculate, spicate or anthelate. 123 spp., cosmopolitan, 43 in New World, 19 in South America, only one in Brazil, L. ulei Buchenau, endemic to highlands of SE Brazil in Minas Gerais, Santa Catarina and Rio Grande do Sul states.

4. Marsippospermum Desv. Herbs to 60 cm tall; rhizome creeping, scale bearing, sympodial; flowerinf stems erect, flowers terminal, solitary. 4 spp., one in New Zealand, and remainig three in Patagonian of Chile and Argentina, also Falklands Is.

5. Oxychloe Phil. Cushion-forming pickly herbs. to several meters in diam, less dense than Patosia or Distichia; usually dioecious or rarely hermaphrodite flowers, this lateral in axills of foliage leaves. 5 spp., in high Andes from Argentina and Chile up to Patagonia, O. andina Phil. reaches in northern up to Peru and Bolivia.

6. Patosia Buchenau. Cushion-forming herbs. to several meters in diam.; dioecious or rarely hermaphrodite flowers, this lateral in axills of foliage leaves. Only one sp., P. clandestina (Phil.) Buchenau, from the Andes from Bolivia to northern Chile and Argentina.

7. Rostkovia Desv. Herbs to 30 cm tall; flowers stems erect; flowers solitary, terminal. Two spp., R. tristanensis Christoph. endemic to Tristan da Cunha and R. magellanica (Lam.) Hook. f. from subantartic islands from New Zealand to South Georgia, and South America mainland Patagonian, disjunct in Ecuador.

CYPERACEAE

§ FULLY CHLOROTROPHIC

Genera/species 94/5,300–5,500 Distribution cosmopolitan, with their largest diversity in cold and temperate regions. Habit usually bisexual or monoecious (rarely andromonoecious, gynomonoecious, dioecious, androdioecious or gynodioecious; unisexual flowers sometimes in bisexual pseudanthia), usually perennial (sometimes annual) herbs (rarely shrubs, lianas or epiphytes; Microdracoides consists of small trees). Graminids. Some species have bulb-like or tuberous swollen internodes or stem bases. A few species possess stilt roots. Fimbristylis fusca xeromorphic, with adventitious roots (with well developed velamen) running down along envelope formed by persistent leaf bases.

3^rd largest family in the Monocots and 7th among the angiosperms; of the 5,400 species described in 106 genera. Carex and Cyperus are the most diverse genera. Carex comprises ca. 2,000 species, mostly from subtropical and temperate zones in the Northern Hemisphere and less well represented in the Neotropics while Cyperus s.l. has ca. 1,000 species and is a cosmopolitan genus with numerous species associated with flooded areas and many others considered weeds. The Neotropical region supports ca. 1,500-2,000 species based on several non-integrated checklists, and Brazil accounts for almost 700 of these.

SYSTEMATIC two subfamilies, both in South America.

1. SUBFAMILY MAPANIOIDEAE (11–12/c 170) ‣ outsiders Capitularina (1; New Guinea), Exocarya (1; Papua New Guinea, Queensland), Lepironia (1; Madagascar and eastwards to Polynesia), Chorizandra (5; Australia, Tasmania, New Caledonia), Chrysitrix (4; W Cape; W Australia), Paramapania (7; Malesia), Scirpodendron (2; India and Sri Lanka to tropical Australia and Polynesia), Principina (1; Principé in tropical W Africa).

1. Diplasia Rich. Tall perennial with creeping rhizomes. Only one sp., D. karatifolia Rich., from Costa Rica to N Peru and Bolivia, Brazil (Amazon rainforest in W Maranhão state), in rain forests.

2. Hypolytrum Rich. Perennials, often on a thick woody rhizome. 58 spp., pantropical, mainly equatorial, in forests; 28 in New World, all also in Brazil except for H. pallidiceps S.S. Hooper & T. Koyama, which is known from the Guyana side of the Roraima Mountains on the border between Brazil and Venezuela; 14 endemics, 7 of them, in several states, are rare plants in Brazil, by Plantas Raras do Brasil’s book.

3. Mapania Aubl. Herbs, rhizomatous or stoloniferous perennials, sometimes medium-sized to tall (tufted or rhizomatous) perennials with culms few-many-noded. 70 spp., pantropical, mainly equatorial, in tropical rainforests and along open swamps; 20 spp. in New World, all of then in South America, three up to Central America, 7 in Brazil, none endemics.

2. SUBFAMILY CYPEROIDEAE (c 98/c 4,300) ‣ 15 tribes, all in South America except Dulichieae (2/5, North America, Europe, temperate Asia) and Sumatroscirpeae (1/4, N Burma, S China, N Vietnam, N Sumatra).

CYPEROIDEAE ▸ UNPLACED CYPEROIDEAE - outsiders Afrotrilepis (2; tropical W and C Africa), Nelmesia (1; N Congo), Neoscirpus (1; Korean Peninsula), Reedia (1; W Australia), Trichoschoenus (1; Madagascar).

4. Rhynchocladium T. Koyama. Only one sp., R. steyermarkii, endemic to Guiana Shield in S Venezuela and Guyana, and in Mount Neblina (absent in Brazilian side), in woodlands and savannas, at elevations of 500-2,400 m.

2.1 CYPEROIDEAE ▸ TRIBE TRILEPIDEAE (3/14) - outsiders Coleochloa (8; tropical and S Africa, Madagascar), Microdracoides (1; tropical W and C Africa).

5. Trilepis Nees. Tufted perennials or dwarf shrubs with a caudex. 5 spp., T. kanukuensis Gilly from the Guiana Shield (Venezuela and Guyana), three endemics to SE Brazil (all are rare plants in Brazil, by Plantas Raras do Brasil’s book, in Rio de Janeiro and Minas Gerais states), and T. lhotzkiana Nees ex Arnott is the most widely distributed species, occurring in Venezuela and the Brazilian states of São Paulo, Rio de Janeiro, Espírito Santo, Minas Gerais, and Bahia.

Among Neotropical Cyperaceae, desiccation tolerance is confined to three Trilepis from SE Brazil.

2.2 CYPEROIDEAE ▸ TRIBE CRYPTANGIEAE (5/43) - all genera occur in South America.

6. Cephalocarpus Nees. Tufted or isolated caudex forming dwarf shrubs (sometimes epiphytic). 21 spp., from Guiana Shield and surrounding areas in northern South America, in Venezuela, Guyana, Colombia, Brazil and Suriname; additionally some species occurs in mountains adjacent to the Andes, in Peru and Ecuador, with similar edaphoclimatic conditions (Cordillera del Condor); its species are more frequent and diverse at higher elevations on nutrient poor sandy or rocky soils.

7. Cryptangium Schrad. ex Nees. (off Lagenocarpus) Monoecious herbs, with short and/or creeping rhizome; caudex absent; inflorescence terminal, solitary, panicle-like. Only one sp., C. verticillatum (Sprengel) Vitta, in non-Andean South America regions, from of Guyana and Brazilian Shields southwards to southern areas of Brazil (probably São Paulo state), from western Brazilian coast to eastern Colombian Vaupés, on sandy soils.

8. Didymiandrum Gilly. (off Lagenocarpus) Dioecious herbs, with short creeping rhizome; caudex absent; inflorescence terminal, solitary, panicle-lik; involucral bracts leaf-like, with elliptic to oblong-elliptic blades. Only one sp., D. stellatum (Boeckeler) Gilly, from Guyana Shield, in Colombia, Venezuela and Guiana and Brazil in borders of rain forest areas.

9. Everardia Ridl. ex Thurn. Dwarf shrubs with woody caudex. 13 spp., almost endemic to Guiana Shield, N South America, terrestrial or epphytic, on bare sandstones rocks, escarpments, in crevices; three spp. in Brazil, all in frontier of Venezuela and Brazil and widely in northern South America to Peru and Ecuador, in Roraima and Amazonas states.

10. Exochogyne C.B.Clarke. (off Lagenocarpus) Monoecious herbs, with short creeping rhizome; caudex absent; inflorescence terminal, solitary, congested or laxly spike-like. Involucral bracts leaf-like, with linear-lanceolate to lanceolate blades, expanded at base and enclosing the paracladia. Two spp., northern South America, on the eastern Guyana Shield and in the Amazon basin, mainly in open vegetation, but is also found in forested white-sand savannas, on nutrient poor sandy soils and among rocks.

11. Krenakia S.M.Costa. (off Lagenocarpus) Monoecious herbs, with short creeping rhizome; caudex absent; inflorescence terminal, panicle-like. Involucral bracts leaf-like, with linear- lanceolate to lanceolate blades. 10 spp. from Brazil and is especially species rich in the campos rupestres (SE Brazil), with two cases of disjunction, one in the Venezuelan highlands and one in the Caribbean (Cuba), mainly in mountain top habitats, mainly in open or riparian vegetation on nutrient poor soils.

12. Lagenocarpus Nees. (exc. Cryptangium, Exochogyne, Didymiandrum, Krenakia) Herbs perennials, rhizomatous or stoloniferous. 15 spp., Caribbean to southern South America (Rio Grande do Sul state, Brazil), in different non-Andean biomes but mainly in soils with low nutrient availability, like sandy, rocky or waterlogged soils. 3 species, in Minas Gerais and Espírito Santo states, are rare plants in Brazil, by Plantas Raras do Brasil’s book.

2.3 CYPEROIDEAE ▸ TRIBE SCLERIEAE (1/250–260) - a single genus.

13. Scleria P. Bergius. Small annuals or medium sized to tal tufted, rhizomatous, or even scrabing perennials. c. 250 spp., cosmopolitan, 121 in New World, 101 in South America, 76 in Brazil, 23 endemics, one of them, from Mato Grosso state, is a rare plant in Brazil, by Plantas Raras do Brasil’s book.

2.4 CYPEROIDEAE ▸ TRIBE BISBOECKELEREAE (4/24) - all genera in South America.

14. Becquerelia Brongn. Thufted or shotly rhizomatous, rarely stoloniferous or perennials. 5 spp., two over N South America (one of then in Brazil) and three Brazilian endemics.

15. Bisboeckelera Kuntze. Tufted, rarely shortly rhizomatous perennial. 4 spp. from N South America (one highly dubious record of Cono Sur), forests; in Brazil two confirmed spp., both in Amazon River northwards, none endemics.

16. Calyptrocarya Nees. Herbs, rhizomatous or stoloniferous perennial. 8 spp. in Central and South America, forests; in Brazil only in Amazon rainforest, up to Maranhão state; six spp., only one endemic.

17. Diplacrum R. Br. Annuals or tufted, rarely stoloniferous herbs. 7 spp., pantropical, in forests, oftem in savanas, with white sands; 5 spp. in New World, 4 in South America, two in Brazil, none endemics.

2.5 CYPEROIDEAE ▸ TRIBE KOYAMAEEAE (1/1) - a single genus

18. Koyamaea W. Thomas & Davidse. Loosely tufted to shortly rhizomatous perennial. Only one sp., K. neblinensis, endemic to Mount Neblina in Guiana Shield (Amazonas state in Brazil and Venezuela), at elevations of 500-2,000 m, in small field place.

2.6 CYPEROIDEAE ▸ TRIBE SCHOENEAE (24/c. 390) - outsiders Trianoptiles (3; N, W and E Cape); Gymnoschoenus (2; S Australia, Tasmania), Tricostularia (5; Australia, one species also in Malesia, S Asia and New Caledonia), Xyroschoenus (1; Seychelles), Morelotia (2; Hawai), Tetrariopsis (1; W Australia), Tetraria (c. 50; Africa, Australia, New Zealand), Epischoenus (8; W and E Cape, KwaZulu-Natal), Lepidosperma (c 65; Malesia to Australia, New Caledonia, New Zealand), Neesenbeckia (1; W Cape), Cyathochaeta (5; W Australia, SE New South Wales), Gahnia (c 40; E Asia, Malesia to Australia, New Caledonia and islands in the Pacific incl. Hawaii), Mesomelaena (5; W Australia), Ptilothrix (1; Queensland, New South Wales), Evandra (2; W Australia), Caustis (5; Australia, Tasmania), Cyathocoma (3; W Cape to KwaZulu-Natal, Mozambique), Capeobolus (1; W and E Cape), Chamaedendron (5; New Caledonia), Costularia (20; SE Africa, Madagascar, Mascarene Islands, Seychelles).

19. Carpha Banks & Sol. ex R. Br. Tufted, mat-forming (rarely stoloniferous) perennials, sometimes cushions. 15 spp., 14 in South Africa, Africa mountains, Madagascar, Mascarenes, New Guinea, Japan, Australia, New Zealand, and C. schoenoides Banks & Sol. ex Hook. f. in Chile and Argentina.

20. Machaerina Vahl. 50 spp., Madagascar and Mascarenes to SE Asia, Malesia, SE Australia, New Zealand, New Caledonia, Caribbean, Pacific islands to North America and South America; in this continent are 5 spp., Juan Fernandes in Pacific Chile and Guyana one endemic each, and three endemics to S & SE Brazil, in swamps and open fields from mountains in Minas Gerais and Rio de Janeiro state, in wetlands, sometimes as floating mats, or in woodlands, often at high altitudes.

21. Oreobolus R. Br. Low cushions or tufted plants. 15 spp., Malesia to SE Australia, Tasmania, New Zealand, Tahiti, Hawaii, Juan Fernandes, Falklands, 5 in W South America and Central, in wet alpine and subantartic vegetation; O. goeppingeri Suess. occurs in Brazil in Mount Neblina and sand dunes in Bahia coast.

22. Schoenus L. Herbs, perennial, cespitose, rhizomatous. c. 100 spp., centered in Australia, S. nigricans L. subcosmopolitan, inc. North America to Mexico and Caribbean, often in humid grasslands or woodlands; a few species occurs in Africa, Eurasia and South America (4): three restricted of Cono Sur, and S. lymansmithii M.T. Strong, very narrow endemic in Morro da Igreja in Santa Catarina state in S Brazil.

2.7 CYPEROIDEAE ▸ TRIBE RHYNCHOSPOREAE (1/379) - a single genus.

23. Rhynchospora Vahl. (inc. Pleurostachys) Herbs, annual or perennial, cespitose or not, often scaly-rhizomatous. c. 379 spp., third largest genera in family, 360 spp. in New World, 246 in South America, 174 in Brazil, 76 endemics; with a marked concentration in (sub)tropical America, most species preferring humid to wet habitats in savannah vegetation, slightly centered in savannas of C Brazil (cerrado). Only seven spp. of flowering plants has the low chromosome number 2n = 4, of which only R. tenuis Willd. ex Link occurs in South America (also in Caribbean); four spp., in four states, are rare plants in Brazil, by Plantas Raras do Brasil’s book.

2.8 CYPEROIDEAE ▸ TRIBE CYPEREAE (14/c. 1,200) - outsiders Scirpoides (4; Mediterranean, S Africa, tropical and subtropical regions in Asia), Hellmuthia (1; S coast of W Cape), Ficinia (c 75; tropical and S Africa, Madagascar, New Zealand, with their highest diversity in the Cape Provinces), Erioscirpus (2; N India, Himalayas, N Burma), Dracoscirpoides (2; South Africa), Afroscirpoides (1; South Africa), Actinoscirpus (1; tropical and subtropical Asia to northern Australia and islands in the Pacific), Pseudoschoenus (1; N, W and E Cape, Free State, Lesotho).

24. Bolboschoenus (Asch.) Palla. Rhizomatous perennials, rhizome often forming hardovoid tubers. 15 spp., over subcosmopolitan, 5 in New World, B. robustus (Pursh) Soják widely distributed, two only in North America, one from North America to Colombia, and B. fernaldii (E.P. Bicknell) Soó ex Govaerts endemic to Brazil.

25. Cyperus L. (inc. Androtrichium, Kyllinga) Annuals, or tufted, rhizomatous, stoloniferous, or bulbiferous perennials. 956 spp., well represented in the tropics, extending towards the temperating regions, ranging from very dry to permanently submerged, from open sunny places to dark forest, from acid bogs to brackish marshes, several species on disturbed ground and becoming weed; in the New World (277), areas of high diversity and endemism includes Mexico, the Greater Antilles, and NE Brazil; 194 spp. in South America, 129 in Brazil, 36 endemics, two of them, from Bahia state, are rare plants in Brazil, by Plantas Raras do Brasil’s book.

The spikelet morphology of C. prophyllatus A.R.O.Ribeiro, Pereira-Silva & M.Alves (endemic to Espirito Santo state, Brazil) is unique among the genus in having both a conspicuous spikelet prophyll and a corky rachilla articulation, which remain persistent at the base of the spikelet after disarticulation.

26. Fuirena Rottb. Small to medium sized annual or rhizomatous perennials. 30 spp., perennials, concentrated in America (a half in South America) and Africa, mainly in open places; 6 spp. in Brazil, two endemics, one of them, from Pernambuco state, is a rare plant in Brazil, by Plantas Raras do Brasil’s book.

27. Isolepis R. Br. Annuals, or mat-forming perennials. 60 spp., subcosmopolitan, with concentration in S Africa and Australia, 10 spp. in New World, mostly in wetlands, or subaquatic, 8 in South America, two in Brazil, I. fluitans (L.) R. Br. endemic.

28. Schoenoplectiella Lye. 51 spp., cosmopolitan; 6 in New World, only S. supina (L.) Lye in South America (Santa Catarina state in S Brazil and NE Argentina), which also occurs in Old World.

29. Schoenoplectus (Reichb.) Palla. Herbs usually perennial, sometimes annual, cespitose or not, rhizomatous or not. 11 spp., cosmopolitan; 4 spp. in South America, scattered from North America to Cono Sur; two in Brazil, both very widely.

2.9 CYPEROIDEAE ▸ TRIBE ABILDGAARDIEAE (7/c. 520) - outsiders Nemum (8; tropical Africa), Trachystylis (1; coastal areas in E Queensland and NE New South Wales), Arthrostylis (1; tropical Australia), Actinoschoenus (3; Madagascar, Sri Lanka, China, N and NW Australia), Crosslandia (1; N Australia).

30. Bulbostylis Kunth. Annual or tufted perennials, rarely with a elongate rhizome. c. 100 spp., tropical to warm temperate, 91 in New World, centered in Africa and South America (78); 64 spp. in Brazil, 28 endemics, six of them, in several states, are rare plants in Brazil, by Plantas Raras do Brasil’s book.

31. Fimbristylis Vahl. (inc. Abildgaardia) Annual or tufted, more rarely creeping. c. 300 spp., pantropical to warm-temperate, heavy concentration in Australasia; mostly in sunny, moist to wet places; 41 spp. in New World, 31 in South America, 24 in Brazil, 11 endemics, two of them, both from Bahia state, are rare plants in Brazil, by Plantas Raras do Brasil’s book.

F. fusca (Nees) Benth. ex C. B. Clarke (Asia to Oceania) is xeromorphic, with adventitious roots (with well developed velamen) running down along envelope formed by persistent leaf bases; among Neotropical Cyperaceae, desiccation tolerance is confined to one Trilepis and F. dichotoma (L.) Vahl., pantropical.

2.10 CYPEROIDEAE ▸ TRIBE ELEOCHARIDEAE (1/280–290) - a single genus.

32. Eleocharis R. Br. Small to medium sized tufted annuals or rhizomatous, stoloniferous, rarely bulbiferous, perennials herbs, sometimes submerged stoloniferous perennials, with floating to partly emerged culms. c. 300 spp., subcosmopolitan with a marked concentration in New World (196), 128 in South America, 83 in Brazil, 29 endemics, 4 of them, all from Bahia state, are rare plants in Brazil, by Plantas Raras do Brasil’s book; wet lands, often emergent, and sometimes submerged aquatic.

2.11 CYPEROIDEAE ▸ TRIBE CLADIEAE (1/4) - a single genus.

33. Cladium P. Browne. Rhizomatous, or stoloneferous perennials. 5 spp., two in North America, C. costatum Steyerm. in wet areas in the extreme N Brazil along the border with Venezuela; C. mariscus (L.) Pohl subcosmopolitan, in New World only in some places in South America inc. Brazil, and C. jamaicense Crantz occurring along the coast in marshes forming dense population, widely in New World.

2.12 CYPEROIDEAE ▸ TRIBE SCIRPEAE (9/74) - outsiders Khaosokia (1; peninsular Thailand); Calliscirpus (2; S Oregon, California, NW Mexico); Eriophorum (20; temperate and arctic regions on the Northern Hemisphere, South Africa).

34. Amphiscirpus Oteng-Yeb. Small perennial from a deep rhizome. Only one sp., A. nevadensis (S. Watson) Oteng-Yeb., W Canada, U.S.A, Bolivia, Chile and Argentina, in sandy, semisaline soil, often in a dried-up lake beds.

35. Oreobolopsis T. Koyama & Guaglian. Small, tufted perennial; culms subscapose. Two spp. from Ecuador to Bolivia, in high mountains.

36. Phylloscirpus C. B. Clark. Small perennial on slender horizontal or vertical rhizome, sometimes cushions. Three spp. in Andean S America, from Colombia to Argentina and Chile, in marsh pina grasslands, and bogs at high altitudes.

37. Scirpus L. Tufted or rhizomatous perennials. 35 spp. from New World (30), Mexico, Eurasia, Australia, Pacific Islands; 11 in South America, 7 in Chile and Argentina, two endemics to Peru, one Peru to Chile and Bolivia, and one endemic to Amazonas state in Venezuela.

38. Trichophorum Pers. Tufted or a creeping rhizomatous perennials, culms scapose. 7 spp., six to Holartic region and tropical mountains in Asia and T. rigidum (Boeckeler) Goetgh., Muasya & D.A. Simpson, from Ecuador to Bolivia and Argentina, in tundra zone or acid bogs, rarely on woodylands.

39. Zameioscirpus Dhooge & Goetgh. Cushions. Three spp. from Peru and Bolivia to NW Argentina, Chile.

2.13 CYPEROIDEAE ▸ TRIBE CARICEAE (1/1,800) - a single genus.

40. Carex L. Herbs, perennial, cespitose or not, rhizomatous, rarely stoloniferous; culms usually trigonous, sometimes round; leaves basal and cauline, sometimes all basal; inflorescences terminal, consisting of spikelets borne in spikes arranged in spikes, racemes, or panicles; bracts subtending spikes leaflike or scalelike; flowers unisexual; achenes biconvex, plano-convex, or trigonous, rarely 4-angled. 2,003 spp. (6^th largest worldwide), cosmopolitan, centered in some cold holartic áreas; it´s absent in tropical lowlands except for a few species in SE Asia; 723 spp. in New World, 193 in South America, 32 in Brazil, 8 endemics; Carex displays the most dynamic chromosome evolution of all flowering plants (n = 6 to n = 66); seven subgenera:

§ subg. Soderostrica ‣ 30 spp., endemic to E Asia, with a centre of diversity in SE Asia.

§ subg. Carex ‣ 1374 spp., cosmopolitan, only absent from Antarctica.

§ subg. Euthyceras ‣ 124 spp., widely distributed in the Northern Hemisphere, with disjunct centres of diversity in North America and Himalayas, with C. arctogea Harry Sm. disjunct in Argentina an Chile, some specie also in New Zealand.

§ subg. Psyllophorae ‣ 53 spp., Western Palearctic, South America, sub-Saharan Africa, and New Zealand, reaching Arabian Peninsula, with centres of diversity in Patagonia and Cape Region.

§ subg. Uncinia ‣ 99 spp., primarily the Neotropics, Australia and New Zealand, but also lineages endemic to North America, the former genus Uncinia also present in Malesia and Pacific and sub-Antarctic archipelagos; it is the only group of Carex present in a true Antarctic region [C. meridensis (Steyerm.) J.R.Starr, South Georgia archipelago]; disjunct centres of diversity in South America and New Zealand.

§ subg. Vignea ‣ c. 330 spp., cosmopolitan, with a centre of diversity in North America, some in S Brazil.

LINEAGE 5 of 6: MAYACACEAE/XYRIDACEAE/ERIOCAULACEAE

MAYACACEAE

§ FULLY CHLOROTROPHIC

Genera/spp. 1/5 Distribution SE U.S.A., the Caribbean, Central and South America; one sp. (Mayaca baumii Gürke) in SW Africa (Angola and possibly Congo and Zambia; due to recent distance dispersal). Distribution bisexual, perennial rooted herbs. herbs. Aquatic or helophytic. Amphibious herbs with ericoid leaves.

Mayacaceae belongs within the Order Poales, but differ vegetatively from most of the other members of this order. Anthers in some spp. are monothecal, the nucellar epidermis is basally thickened and the outer layer of the endosperm has proteins.

Key differences from similar families Differs from other aquatic plants such as:

ü Pontederiaceae, which have inflated petioles and flowers arranged in dense inflorescences.

ü Haemodoraceae, where the calyx and corolla are fused and septal nectaries present.

SYSTEMATIC A single genus.

1. Mayaca Aubl. Aquatic to telmatic perennial, rooted herbs, leaves in umbel or simples, spirally arranged along this stems; flowers solitary, axillary, emergent, actinomorphic, perfect, pink or white; aquatic submerged or amphibious, inner or along watercourses, low deep. 5 spp., M. baumii Gürke from SW Africa in Congo, S. DR Congo to Angola, and possibly Zambia, due to recent distance dispersal, and 4 native to tropical and warm-temperate America, M. fluviatilis Aubl. SE U.S.A., the Caribbean, Central and South America up to Argentina, M. longipes Mart. ex Seub. and M. sellowiana Kunth over South America, the latter also in Costa Rica, and M. kunthii Seub. more restricted, from Brazil, Venezuela and Uruguay, found nearly throughout the country, growing in rivers and perennial or temporary lagoons; species differentiation is quite a hard task, relying on anatomical features of the androecium, presence or absence of inflorescence and corolla color.

ERIOCAULACEAE

§ (Brocchnia - Catopsis - Paepalanthus - Drosera - Heliamphora - Philcoxia - Genlisea - Utricularia - Pinguincula)

Genera/spp. 16(own data)/c. 1,270 Distribution tropical and subtropical regions on the Southern and Northern Hemispheres, with their largest diversity in the Guiana Shield and SE Brazil; a few spp. of Eriocaulon in temperate parts of Europe, E Asia and North America; Mesanthemum: tropical Africa, Madagascar. Habit usually monoecious (rarely dioecious or bisexual), perennial or annual herbs. Many spp. are helophytes and some are aquatic.

1,200 species with ca. 850 species and nine genera concentrated in South America, especially in Brazil; the main centers of taxonomic and morphological diversity are the Espinhaço Range and the Guiana Shield, most of them are distributed in the Brazilian campo rupestre vegetation, comprising more than 80% of the species; secondary centers of diversity are located in the central savannas, especially in Goiás and Mato Grosso, and in the Amazon rainforest domain; in these two areas diversity is greatly underestimated, as they have been poorly collected and barely studied.

A fairly recent development has been expansion of the trade in everlasting plants (‘sempre-vivas’) - collection and trade in dried inflorescences of some 40 wild spp., several of them rare endemics, that are in demand particularly for export to U.S.A., Japan and Europe. About 300 metric tonnes dry weight are being obtained yearly. The spp. most prized for their beauty are the Eriocaulaceae Comanthera elegans (Bong.) L.R. Parra & Giul., C. brasiliana (Giul.) L.R. Parra & Giul., C. suberosa (Giul.) L.R. Parra & Giul., C. magnifica (Giul.) L.R. Parra & Giul., Syngonanthus xeranthemoides (Bong.) Ruhland, Actinocephalus macrocephalus (Bong.) F.N.Costa & Sano and Leiothrix flavescens (Bong.) Ruhland; the scapes and inflorescences of these spp. are collected and dried in the sun, to be sold as ornamental objects and often exported from Brazil to different countries; some spp. in the states of Bahia, Minas Gerais, and Tocantins are critically endangered due to over-exploitation.

SYSTEMATIC two subfamilies, both in South America.

1. SUBFAMILY ERIOCAULOIDEAE (2/c. 500) - outsider is Mesanthemum (16; tropical and subtropical Africa, Madagascar).

1. Eriocaulon L. Annual or perennial herbs, mostly rosulate, highly variable in size, submerged or floating spp. 480 spp. worldwide (truly pantropical, with at least one species reaching temperate areas in Europe, also in Japan and Australia), 113 in New World, 78 in South America, 63 in Brazil, 47 endemics (9 are rare plants in Brazil, by Plantas Raras do Brasil’s book, in several states, mainly in Minas Gerais).

2. SUBFAMILY PAEPALANTHOIDEAE (17/c 770) ‣ four clades, all in South America.

2.1 PAEPALANTHOIDEAE ▸ CLADE RONDONANTHUS (1/5) - a single genus.

2. Rondonanthus Herzog. Herbs, perennial, rosulate; flowers trimerous, unisexual, hermaphrodite or male. 5 spp., endemic to Guiana Shield, at elevations of 100-2,800 m, two restricted of Venezuelan tepuis, remaining three up to Brazil (none endemics) and Guyana, with R. capillaceus (Klotzsch ex Koern.) Hensold & Giuliett, which occurs also in Mount Aracá in Amazonas and Oiapoque River in Amapá state.

2.2 PAEPALANTHOIDEAE ▸ LEIOTHRIX CLADE (1/50) - a single genus.

3. Leiothrix Ruhland. Eriocaulon-like, sometimes cushion-like. 50 spp., L. flavescens (Bong.) Ruhland occurs in Brazil, Bolivia, Venezuela, Guyana, Peru and Colombia, L. celiae Moldenke is exclusively to Venezuela, L. arechavaletae Ruhland is restricted for Cono Sur, and the remaining 47 taxa are endemic to small mountainous areas in the Brazilian states of Minas Gerais and Bahia (13, one in Bahia and 12 remaining in Minas Gerais state, are rare plants in Brazil, by Plantas Raras do Brasil’s book). L. fluitans (Mart.) Ruhland is the only aquatic taxon in the genus.

2.3 PAEPALANTHOIDEAE ▸ CLADE SYNGONANTHUS (2/161) - both genus in South America.

4. Comanthera L. B. Smith. Herbs, rosulate or caulescent spp. 34 spp., all in Brazil, only C. kegeliana (Körn.) Moldenke outside, up to Venezuela and Guyana, in two subgenera, subg. Comanthera (c. 20), and subg. Thysanocephalus (c. 15), with its diversity concentrated in the Espinhaço Range in SE Brazil and in the Guiana Shield; most spp. are restricted to rocky grasslands (campos rupestres) of quartzitic origin, an ecosystem composed of rocky outcrops and white-sand savannas mixed with gallery forest and influenced by the surrounding biomes, notably the Atlantic Forest and the savannas of C Brazil (cerrado).

5. Syngonanthus Ruhland. Herbs, rosulate or caulescent; inflorescence in the center of rosette, short to long. 128 spp., two exclusively Caribbean, S. flavidulus (Michx.) Ruhland in SE U.S.A., 6 exclusively from Mexico and Central America, and 119 in South America, 116 exclusive, few spp. in Africa one; the 102 Brazilian species (68 endemics, 20 in several states are rare plants in Brazil, by Plantas Raras do Brasil’s book) are distributed all over the country, but concentrated in the savannas of C Brazil (cerrado); it is the most diverse genus of Eriocaulaceae in the Amazon rainforest. Some populations of S. umbellatus (Lam.) Ruhland in northern Pará state, Brazil, have strog remarkable blue anthers, a feature unique in this family.

2.4 PAEPALANTHOIDEAE ▸ PAEPALANTHUS COMPLEX (12/474) - outsiders Nisius (7, Cuba) and Lachnocaulon (6; SE North America to Cuba).

Classfication below excludes 13 spp., Paepalanthus balansae Ruhland from Brazil to Paraguay, possibly a new genus, P. bosseri (Morat) Stützel and P. itremensis (Morat) Stütze from Madagascar, both possibly new genera each; P. extremensis Silveir from C Brazil, possibly a Floralia species; P. guaraiensis Moldenke, P. scholiophyllus Ruhland and P. lundii Körn. from C & SE Brazil, these poor related and possibly a new genus; P. magistrae Sano from Piauí state, possibly a new genus; P. mellii Moldenke from Mexico and P. pauper Moldenke from Guyana, these placed in Syngonanthus; P. lamarckii Kunth from Gambia to Zambia and Tanzania, Madagascar, Belize and Caribbean to Bolivia and Brazil, P. obtusifolius (Steud.) Körn. from Brazil [BA], and P. tortilis (Bong.) Körn. from over northern South America, these very strong related and possibly united in a new genus.

6. Actinocephalus (Körn.) Sano. Perennial or monocarpic perennial herbs.Stems short or elongated, with rosulate or spiral leaves. 54 spp., endemic to Brazil, from Alagoas to Rio Grande do Sul and Tocantins states.

7. Cora Andrino & Sano. Perennial herbs; stems short with rosulateleaves; third-order inflorescence consisting of scapes with capitula emerging from the apex of the fertile main axis; flowers dimerous, pistillate flowers with free petals and free stigmatic branches. 22 spp., all endemics to Brazil except C. chiquitensis (Herzog) Andrino also in Bolivia, Colombia and Venezuela.

8. Cryptanthella (Suess.) Andrino. Cushion plants orplants with flat mats of rosettes, outer capitulum whorl withtrimerous pistillate flowers, staminate flowers central. 14 spp., 8 from Venezuela (4), Colombia (6), Ecuador (3), Peru (4), Bolivia (2), and six endemics to Brazil.

9. Floralia Andrino & F.N.Costa. Perennial herbs; stems short with rosulateleaves or stems elongated, branched, erect and flexible, withspiral leaves along the stem. 15 spp., endemic to E Brazil, in states of Minas Gerais, Bahia, Goiás and São Paulo, with only one species known from Rio Grande do Sul, only from the type collection.

10. Giuliettia Andrino & Sano. Annual herbs; stems elongated, un-branched, rarely branched. Second-order inflorescences con-sisting of scapes and capitula emerging from the same pointat the stem apex. 30 spp., 25 widely distributed in tropical South America, almost in over Brazil (19, 9 endemics), reaching the Guayana Shield in Venezuela, Colombia and Bolivia, and two in Belize and three in W Cuba.

11. Gnomus Andrino & Sano. Herbs, annual or perennials, which never exceed 20 cm in height, with some speciesbeing among the smallest in Eriocaulaceae, with only 1 cm. 6 spp., endemics to highlands of central Brazil (Veadeiros Chain), Espinhaço Range and Atlantic sandy coastal shrublands (restingas), in Goiás, Minas Gerais and Bahia states, Brazil.

12. Hydriade Andrino. Herbs, perennials, distinguished from all othergenera of Eriocaulaceae by the elongate, erect to ascendingstem, covered by persistent leaves, and by terminal inflorescences in which one or two lateral ramifications develop afterthe emergence of the scapes with capitula, with continuous growth of the vegetative parts. 7 spp., 5 endemics to Brazil and two in Ecuador and Peru.

13. Monosperma (Hensold) Andrino. Herbs, perennials, single-seeded indehiscent fruits, in contrast to the capsular fruits produced by most Eriocaulaceae. 24 spp., all endemics to the Pantepui, in the Guiana Shield of Venezuela (22, 15 endemics), Brazil (3, M. septentrionalis (Trovó) Andrino endemic), and Guyana (6, one endemic), with M. gleasonii (Moldenke) Andrino in all three countries; most species are narrow endemics to the summits of one or few tepui formations.

14. Paepalanthus Mart. Perennial or annual herbs; stems short orelongated, branched or unbranched; first-order inflores-cences, rarely second order; trimerous flowers, pistillate flowers with free petals and free stigmatic branches. 257 spp., all endemics to Brazil - largely restricted to the rocky grasslands (campos rupestres) of the Espinhaço Range and adjacent mountains, with few species in the Serra da Mantiqueira complex, and in Atlantic sandy coastal shrublands (restingas) - except by 15 spp., all in subg. Platycaulon, one of them in Brazil and Peru (P. planifolius (Bong.) Körn.), and 14 fully absents, from Costa Rica to Colombia (all) and Venezuela.

15. Tonina Aubl. Annual herb. Stems elongated, branched,erect and flexible, with spiral leaves along the stem. Only one sp., T. fluviatilis Aubl. widely distributed in the Neotropics, occurring in coastal flooded areas of Central and South America.

XIRYDACEAE

§ FULLY CHLOROTROPHIC

Genera/spp. 5/c. 380 Distribution tropical and subtropical regions on the Southern and Northern Hemispheres, with the largest diversity in northern South America; few spp. in temperate areas. Habit bisexual, usually perennial (sometimes annual) herbs. Often helophytic (rarely aquatic); some spp. are xerophytic.

Plants of seasonally or permanently wet habitats, with the major generic diversity occurs in Guiana Shield with three small endemic genera, although the number of spp. in the largest and most widely distributed genus Xyris is fewer there than in the rocky grasslands (campos rupestres) of Brazil.

SYSTEMATIC two subfamilies, both (and their all genera) in South America.

1. SUBFAMILY XYRIDEAE (1/250–260) ‣ a single genus.

1. Xyris L. Usually rosulate (less often caulescent), solitary or caespitose, sometimes rhizomatous or bulbous herbs, also cushions; few are true aquatic; yellow epedicellate flowers in spikes, subssessile, solitary in bracts axil. ca. 400 spp., pantropical, reaching to Canada Argentina and China, 264 in New World, 235 in South America. 179 found in Brazil (132 endemics, 31 of then are rare plants in Brazil, by Plantas Raras do Brasil’s book, in Amazonas, Goiás, São Paulo, Bahia and Minas Gerais (mainly) states), 46 are endemic to the Espinhaço Range and 14 exclusive to the Serra do Cipó, often associated with wetlands and habitats affected by periodic fires, such as bogs and savannas and also humid rock outcrops at high elevations; three sects:

§ sect. Nematopus ‣ the most diverse with ca. 250 species mainly in South America and basal or central placentation.

§ sect. Pomatoxyris ‣ the smallest section with 22 species restricted to Australia and has axillary placentation.

§ sect. Xyris ‣ pantropical and has ca. 140 species with parietal placentation.

2. SUBFAMILY ABOLBODOIDEAE (4/c 25) ‣ all genera in South America (with their largest diversity in the Guiana Shield).

2. Achlyphila Maguire & Wurdack. Perennial from slender, branched rhizomes; inflorescence usually with 2 approximate, distichously arranged leaflike spathes subtending 1-3 flattened-pedicelled flowers, sepals 3, lanciform, chaffy. Only one sp., endemic to Pantepui Life Zone, A. disticha Maguire & Wurdack, known only from summit elevations of Mount Neblina on the Venezuelan side of Venezuela/Brazilian frontier, at elevations of 1,700-2,300 m.

3. Abolboda Humb. & Bonpl. Cespitose or solitary, often cushion-like, mostly glabrous perennials, rarely annuals; inflorescence subsessile to strongly scapose; leaves mostly basal. 24 spp., highly centered in Guiana Shield (15 restricted of this area) in N South America, few reaching to S & SE Brazil (10, two endemics) and Bolivia (three spp., one endemic), boggy, cold to warm habitats, mainly in open places.

4. Aratitiyopea Steyerm. & P. E. Berry. Robust, Navia-like, perennials from a thick rhizome; dowstem and upperstem leaves sub-dimorphic, up a ‘leaf-cephalia’ subtending flowers (this largest, with colorful bracts), this actinomorphic, 7-10 cm, white to pale purple. Only one sp., A. lopezii (L. B. Smith) Steyem. & Berry, highly ornamental, from NW Brazil in Amazonas state, S Venezuela and SE Colombia, and one (possibly the same species) disjunct record of Cordillera del Condor, northern Peru.

5. Orectanthe Maguire. Glabrous perennias; stems very short (basal rosetes) or erect short; scapes 1-several, naked, variously elongate; inflorescence a large, terminal burrlike head; flowers pale yellow, rarely purple. Two spp. endemics to Guiana Shield in Venezuela and Guyana, O. sceptrum (Oliv.) Maguire reaching in N Brazil (Roraima and Amazonas states), in sandstones of region, in medium to high elevation, at elevations of 500-2,800 m.

LINEAGE 6 of 6: POACEAE and RELATED GRAMINIDS

RESTIONACEAE

§ FULLY CHLOROTROPHIC

Genera/spp. 48/500-505 Distribution southern Africa, Madagascar, Indochina, Malesia, New Guinea, Australia, Tasmania, New Zealand, New Guinea and SE Asia northwards to Hainan and Indochina (Chordifex), with the largest diversity in SW Australia and the W Cape Province in South Africa; Apodasmia: Chile; southernmost South America (one sp. of Gaimardia in Tierra del Fuego and the Falkland Islands). Habit Usually dioecious (in Coleocarya, Centrolepis, Gaimardia, Aphelia and Lepyrodia monoecious, andromonoecious or polygamomonoecious; in, e.g., some spp. of Lepyrodia sometimes bisexual), perennial (sometimes annual, usually annual in Centrolepis, Gaimardia and Aphelia) herbs. Graminids. Often xeromorphic. Culm photosynthesizing, simple or branched (branches sometimes verticillate), usually terete (rarely quadrangular or flattened in cross-section), smooth, verrucose, striate, furrowed or pitted, medullated or with a hollow centre, often with compact swollen nodes.

Use Ornamental plants (Elegia sp. etc.), thatching (Thamnochortus insignis Mast., etc.), forage plants.

SYSTEMATIC 4 subfamilies, Restionoideae (c 17/350–355, Africa south of Sahara, Madagascar, Australia) and Sporadanthoideae (3/22, Australia, Tasmania, New Zealand, Chatham Islands) do not occur in South America.

1. SUBFAMILY LEPTOCARPOIDEAE (23/c 90) ‣ outsiders are 22 genera, 20 restricted of Australia, Empodisma (2; Australia, Tasmania, New Zealand) and Dapsilanthus (4; Hainan, Thailand, Indochina, the Malay Peninsula, Aru Islands, New Guinea, Northern Territory, N Queensland).

1. Apodasmia B.G.Briggs & L.A.S.Johnson. Three spp., one in Australia, one in New Zealand, and A. chilensis (Gay) B.G.Briggs & L.A.S.Johnson in Chile.

2. SUBFAMILY CENTROLEPIDOIDEAE (3/36) ‣ outsiders are Aphelia (6; Australia) and Centrolepis (26; Hainan, Indochina, Malesia to Australia).

2. Gaimardia Gaudich. Herbs, sometimes cushions. 4 spp., two in Tasmania, one in New Guinea, Tasmania, New Zealand South Island, and G. australis Gaudich. in Falkland Islands, Magellan Strait region in Chile and Argentina.

POACEAE

§ FULLY CHLOROTROPHIC

Genera/spp. 793/c. 12,074; Distribution cosmopolitan including polar areas; Habit usually bisexual (sometimes monoecious, andromonoecious, gynomonocious, polygamomonoecious, dioecious, androdioecious, or gynodioecious), usually perennial, biennial or annual herbs (sometimes woody, up to c. 40 m tall). Graminids. Sometimes helophytes, rarely aquatic. Numerous spp. are xerophytes. Culm terete to elliptic in cross-section, usually with hollow (fistulose; sometimes medullated solid) internodes and solid swollen nodes.

The Poaceae has at least 11,000 species recorded in ca. 700 genera which makes it the fifth largest plant family and the second among the Monocots (a global checklist is available and continuously updated by Soreng et al. 2014). Grasses are distributed throughout all continents and are found in almost all continental environments of the planet. Festuca, Poa, Stipa, Eragrostis and Paspalum are the most diverse genera.

In the Neotropics, more than 4,000 species of grasses in almost 300 genera are known, with Paspalum and Eragrostis being especially diverse. Poaceae is a sister group of a clade composed of two small families, Ecdeiocoleaceae (1/3, Australia) and Joinvilleaceae (1/4, Pacific zone), and comprises the graminid clade with Flagellariaceae (1/4, Paleotropics).

DIVERSITY some regions in the Neotropics must be highlighted for the diversity and taxonomic importance they represent for Poaceae. The Central Brazilian savannas holds an impressive diversity, especially of C4 species of Panicoideae and Chloridoideae; the Atlantic Forest, Guiana Shield and Andes are home to several endemic species; the Amazon Forest, although having not been satisfactorily inventoried, is also important for its diversity of habitats and species; and finally the Pampas is also highly diverse. In the savannas of C Brazil (cerrado) ca. 700 species in 126 genera occur, which represents almost 50% of the total number cited to the country.

Savannas of C Brazil (CERRADO) - this amazing grass diversity can be associated with the variety of habitats which can be found in the savannas of C Brazil (cerrado), such as riparian forests, grasslands, rocky outcrops, wetlands, among others, some of which are subjected to fire; the three richest genera belong to Panicoideae: Paspalum (153), Axonopus (41) and Panicum (34), followed by Eragrostis (33), Digitaria (27) and Aristida (26); the rocky grasslands (campos rupestres) of the Espinhaço Range supports at least 36 endemic species besides peculiar genera such as Apochloa, Dichanthelium, Renvoizea and the monotypic Plagiantha tenella Revoinze; the quartzitic mountains of Veadeiros National Park and the serpentine soils of Niquelândia region (state of Goiás) are also remarkable for their high levels of endemism. Filgeuirasia species, Actinonocladum verticillatum (Nees) McClure ex Soderstr. and Aulonemia effusa (Hack.) McClure are a few bamboos adapted to fire.

ATLANTIC FOREST - Atlantic Forest is one of the centers of diversity of bamboos in the world, with more than 160 native species, 135 of them endemic; it also houses a high diversity of Panicoideae (C3 taxa) adapted to shady habitats, such as Ichnanthus, Parodiophyllochloa, and Ocelochloa. In the high-altitude grasslands (campos de altitude) some odd plants such as Glaziophyton mirabile Franch.; Cambajuva ulei (Hack.) P. L. Viana, L. G. Clark & Filg., Cortaderia modesta (Döll) Hack., Apochloa lutzii (Swallen) Zuloaga & Morrone, as well as several species of Chusquea sect. Swallenochloa and Dicahnthelium are found; the high-altitude forests associated with these mountains are also rich in endemic species of woody bamboo, mainly from Merostachys, Chusquea and Aulonemia, and some Panicoideae, such as Paradiophyllochloa penicillata (Nees ex Trin.) Zuloaga & Morrone, Ocelochloa latissima (J.C.Mikan ex Trin.) Zuloaga & Morrone, and Hymenachne condensata (Bertol.) Chase. Colanthelia and Apoclada, endemic genera of bamboo, are also found in high-altitude forests; the central part of the Atlantic Forest (southern Bahia and Espírito Santo) is relevant to the diversity of grasses including the narrow endemic and threatened basal grass Anomochloa marantoidea Brongn.; herbaceous bamboo, such as Raddia, Piresia, Diandrolyra, Olyra, Parianella, and Eremitis, and woody ones in Alvimia, Atractantha, and Eremocaulon.

GUIANA SHIELD - Davidse et al. (2007) cited 526 species to the Guiana Shield and most of the endemic ones are related to the Tepuis, such as Cortaderia roraimensis (N.E.Br.) Pilg. and other species in Arthrostylidium, Aulonemia, Dichanthelium, Chusquea, Myriocladus and Trichanthecium.

AMAZON RAINFOREST - The Amazon Forest has not been satisfactorily inventoried but some typical components of the agrostological flora can be highlighted. The lowland forests are the main center of endemism of herbaceous bamboos in Pariana (ca. 30), besides Guadua (at least 10), Olyra (14), Raddiella (5) and the poorly known genera Agnesia and Froesiochloa. Other species are typical from open, white-sand savanna vegetation, especially of the Neotropical genera Axonopus and Paspalum and the monotypic Arundoclaytonia dissimilis Davidse & R.P. Ellis and Steyermakochloa angustifolia (Spreng.) Judz, which seem to be endemic to this habitat; the scarce mountains in the region are also diverse and include several species endemic to the iron rocky-ferruginous grasslands (canga) in the Serra dos Carajás, the sandstone formation of Serra do Cachimbo and the granitic rocky outcrops of Serra do Tumucumaque in Brazil, French Guiana, Guyana and Suriname.

ANDES - The Andes are highly diverse in woody bamboo genera including the endemic genera Aulonemia, Arthrostylidium, Chusquea, Elytrostachys and Rhipidocladum, which are often associated with high-altitude forests and the Páramos (especially Chusquea and Aulonemia). According to these authors, 90% of the bamboo species in the Andes are endemic. Some groups of grasses which are especially rich in temperate regions and belong mainly to Poideae are also well represented in the region, such as Poa, Festuca, Piptochaetium, and Cortaderia.

South America conspectus with genera:

Other aspects of the eco-physiology of Poaceae to be taken into account:

Cold Tolerance. The ecological success of Poaceae is not just because some adopted C4 photosynthesis; cooler temperate grasslands in the northern hemisphere are dominated by Poöideae, all of which are C3 grasses; thus although about 16% of all species growing in Quebec and Labrador north of 54º N are Cyperaceae, it is Poaceae that are next at 11%, and all are members of Poöideae. Poöideae have a complex relationship between freezing tolerance, day length, vernalization, and flowering. Core Poöideae evolution may be linked with the cooling at the beginning of the Oligocene ca 33-27 m.y.a., gene families implicated in low temperature-induced stress response expanding prior to Poöideae diversification; the genes seem to have been under positive selection; proteins that inhibit ice recrystallization are known from the group; fructans are probably also involved in cold tolerance. Low levels of fructan - specifically levans - accumulation have been noted in many Poaceae, but notably high levels are found only in Poöideae, although not in taxa of the ‘basal’ pectinations like Nardus, Stipa and Phalaridinae; fructans may enable Poöideae that accumulate them to survive drought or frost better, and they have been implicated in stabilizing cell membranes at low temperatures.

As mentioned, the evolution of Poöideae may initially be linked with cooling at the onset of the Oligocene, but much subsequent diversification is later and associated with Pleistocene cooling. Thus the Poa alliance, whose 775 species are about 1/5th of all Poöideae, may have begun diversifying in the Miocene ca 15 m.y.a., but most is Palaeocene and younger, occurring within the last 4 m.y. Furthermore, the diversification rates are very high, up to 3.93 species/million year, although this depends on dating, and much is not connected with obvious island-like areas.

Other grasses also tolerate cooler condition, including the more northerly temperate bamboos (Bambusoideae: Arundinarieae) and the austral Danthonioideae. In the latter, evolution of cold tolerance is estimated to have begun ca 25 m.y.a. during the late Oligocene in Africa. The two species of Danthonioideae studied (Chionochloa) seemed to tolerate cold conditions by controlling ice nucleation.

Some Poaceae show allelopathic reactions with other plants, Sorghum roots producing an allelopathic quinone (an oxygen-substituted aromatic compound) and Festuca roots meta-tyrosine, a non-protein amino acid. Benzoxazinoids like DIMBOA (2,4-dihydroxy-7-methoxy-1,4-benzoxazin-3-one), cyclic hydroxamic acids, are largely restricted to Poaceae, and are found in both Panicoideae and Poöideae. They confer resistance to fungi, insects (volatilized, they attract wasps that parasitize the herbivorous insects earting them), and even herbicides, and are also allelopathic, but less so to other grasses than other plants.

Poaceae such as Spartina (= Sporobolus sect. Spartina) and Puccinellia are major elements of salt marshes; the C4 Sporobolus sect. Spartina (Chloridoideae) is a particularly prominent component of temperate salt marshes where it dominates large areas; there has been past hybridisation in the genus, and hybridization also occurs between introduced and native species, some of the products being very invasive. Salt tolerance in grasses is quite widely distributed, and two thirds of the species are also C4 plants. Some 200+ species are involved, and weak salt tolerance - tolerance of salinity up to ca 80mM NaCl - has evolved some 76 times, possibly being preceed by the acquisiition of C4 photosynthesis. Euhalophytes, tolerating at least 200mM NaCl, about half the salinity of sea water, have evolved some 43 times, and in both cases the clades involved are young and small. Functioning salt glands are known only from Chloridoideae. A number of grasses in different subfamilies accumulate glycine betaines and other compounds commonly associated with allowing plants to grow in saline conditions. Bambusoideae and Danthonioideae are notable for lacking even weak halophytes.

Woody bamboos tend to colonize forest gaps and edges and can dominate in the canopy and understory of both temperate and tropical forests, particularly in mountainous regions. In western Amazonia around 160,000 km² of forests is dominated by two species of Guadua, possibly because of the activities of the geoglyph builders who became active around 4,000 years ago. Even herbaceous bamboos (Olyreae) may dominate understory vegetation. However, although bamboos are the second most important woody monocot clade (after palms), they do not appear in the very top ranks of any of the important ecological traits studied.

At least some species of Micraira (Micrairoideae) are resurrection plants (Sanchez-Ken et al. 2007).

SYSTEMATIC the most updated infrafamilial classification for the Poaceae proposed by the Grass Phylogeny Working Group accepts 12 subfamilies and a long list of s and subs. Anomochloideae, Pharoideae, and Puelioideae comprise the basal grass clade; Panicoideae, Aristioideae, Chloridoideae, Micrairoideae, Arundinoideae, and Danthonioideae make up the PAC-MAD clade; and Pooideae, Errhartroideae and Bambusoideae are the subfamilies of the BEP clade.

All five occur in the Neotropics, except Puelioideae (endemic to Africa). The first four subfamilies occur mainly in forests, frequently with pseudopetiolate leaves, and their photosynthetic pathway is C3 (Bambusoideae) or presumed C3. The representatives of the other subfamilies occur predominantly in open savannas and grasslands and present generally linear and not pseudopetiolate leaves, with C3 or C4 photosynthesis, depending on the subfamily.

1. SUBFAMILY ANOMOCHLOOIDEAE (2/4) ‣ the most basal family; two monotypic tribes, both in South America.

1.1. ANOMOCHLOIDEAE ▸ TRIBE ANOMOCHLOEAE (1/1) - a single genus.

1. Anomochloa Brongn. Rhizomatous perennial, herbaceous; looking more like a member of zingiberales than a grass. Only one sp., A. marantoidea Brongn., a rare herbaceous grass (by Plantas Raras do Brasil’s book) that grows exclusively in shaded, tropical forests of Una municipality, in Bahia state, NE Brazil.

1.2 ANOMOCHLOIDEAE ▸ TRIBE STREPTOCHAETEAE (1/3)

2. Streptochaeta Schrad. ex Nees. Herbs perennial, rhizomatous (the internodes crowded), with inflorescence spike-like. Three spp., from Mexico to Argentina, S. angustifolia Soderstr. endemic to Espírito Santo state, Brazil, S. sodiroana Hack. from Mexico to NW South America, and S. spicata Schrad. ex Nees widely distributed.

2. SUBFAMILY PHAROIDEAE (3/12) ‣ outsiders Leptaspis (3; tropical Africa, Madagascar, the Comoros, tropical Asia from India and Sri Lanka to Queensland and Melanesia), Scrotochloa (2; India, Sri Lanka, Burma, SE Asia, Malesia to New Guinea, Bismarck Archipelago, Solomon Islands, Queensland).

3. Pharus P. Browne. 8 spp., Mexico to Argentina, Florida and Caribbean, 7 in South America, 4 in Brazil (all very distributed); most spp. are widely distributed, although there are two, P. ecuadoricus Judz. in Ecuador and P. vittatus Lem. in Mesoamerica and Colombia, that are more geographically restricted.

3. SUBFAMILY ORYZOIDEAE (19/115) ‣ three tribes, Ehrharteae (1/c 35, South Africa to Ethiopia, Mascarenes, Malesia, Philippines, Java to Australia and islands in the Pacific, mountains in New Zealand, W and E Cape) absent in South America.

3.1 ORYZOIDEAE ▸ TRIBE STREPTOGYNEAE (1/2) - a single genus.

4. Streptogyna P. Beauv. Two spp., one sp. with epizoochorous spikelets, S. americana C. E. Hubb., is widely distributed in the American tropics, from U.S.A. to Bolivia and Brazil, and a second spp. occurs in the tropics of Africa and Sri Lanka.

3.2 ORYZOIDEAE ▸ TRIBE ORYZEAE (13–14/72–73) - three subtribes, Phyllorachideae (3/5, tropical E Africa, Madagascar) does not occur in South America.

■ SUBTRIBE ORYZINAE ‣ outsiders Maltebrunia (4; tropical to S Africa, Madagascar), Prosphytochloa (1; E Cape, KwaZulu-Natal, Mpumalanga, N Province).

5. Leersia Soland. Annual or perennial, rhizomatous, stoloniferous or ceaspitose herbs; hermaphroditic spikelets. 20 spp., 6 in the New World, widely distributed; three in South America, all of then in Brazil, none endemics.

6. Oryza L. 25 spp., two cultivated (one of Asian origin, another in Africa) and 21 wild, in Asia (10), Africa (5), Australia (2) and in New World (4), all of then occurs in wet areas in N & W Brazil, with O. glumipatula Steud. endemic; hermaphroditic spikelets.

■ SUBTRIBE ZIZANIINAE ‣ outsiders Chikusichloa (3; China, Japan, the Ryukyu Islands, Sumatra), Hygroryza (1; S and SE Asia to China), Potamophila (1; N New South Wales), Zizania (4; E India to E Asia, North America).

7. Luziola A. L. Juss. Annual, or perennial; stolonferous or decubent; herbaceous, branched above. 10 spp. from U.S.A. to Argentina and Caribbean; 9 spp. in South America, 7 in Brazil, two endemics.

8. Rhynchoryza Baillon. Perennial, culms herbaceous, helophytic and glycophytic. Only one sp., R. subulata (Nees) Baill, found in swamps in austral South America, S Brazil, Argentina and Paraguay.

9. Zizaniopsis Doell & Aschers. 6 spp., one in North America and Mexico, one in Colombia, one only in Argentina, Z. bonariensis (Bal. & Poitr.) Speg., from Argentina, Uruguay, and S Brazil, and two endemics to Brazil.

4. SUBFAMILY BAMBUSOIDEAE (118/1.555–1.580) ‣ three tribes, Arundinarieae (30/c 550, Madagascar, Himalayas to E Asia, North America, with their largest diversity in China) does not occurs in New World. Bambusoideae representatives occur predominantly in forests, although some genera present species growing in open grasslands especially in high altitudes, e.g. the genus Chusquea Kunth, and are C3, as the three subfamilies mentioned above. Bambusoideae presents a very high richness, with ca. 110-115 genera and 1,400 spp. and includes three traditional groups, in the lignified bamboos (Bambuseae and Arundinarieae), very diverse in the Neotropics, and the herbaceous bamboos (Olyreae), almost restricted to this region, except by Olyra latifolia L., also occurring in Africa (native or introduced?) and Buergersiochloa, monospecific endemic from N. Guinea. Representatives occur predominantly in forests, although some genera present spp. growing in open grasslands especially in high altitudes, e.g. the genus Chusquea Kunth. In New World, only five genera outside South America, three herbaceous from Cuba, one from Mexico to Honduras, and one from SE U.S.A.; Bamboos are highly diverse in the Atlantic Rain Forest, Brazil, including four genera endemic from Olyreae and only one from Bambuseae. It is well represented in the Amazonian Forest as well, but more collections are needed in this area.

4.1 BAMBUSOIDEAE ▸ TRIBE BAMBUSEAE (67/890–915) - 11 tribes, genera about equally distributed between the Old and New World, with somewhat greater spp. diversity in the Old World; four subtribes are native to the New World; Arundinariinae are the only subtribe native to both hemispheres. 155 spp. in Brazil, 129 endemic.

■ SUBTRIBE CHUSQUEINAE ‣ a single genus.

10. Chusquea Kunth. 193 spp., from Mexico to South America (153), making the largest genus of bamboos, centered in central and N Andes from Colombia to Peru (c. 87), C and SE Brazil (53, 46 endemics, 5 of them from Minas Gerais to Santa Catarina state, are rare plants in Brazil, by Plantas Raras do Brasil’s book), and Mexico, and Central America (35); the genus has 15 spp. in southern South America, and single disjunct spp. each in the Caribbean (C. abietifolia Griseb.), the Juan Fernandez Islands in the S Pacific Ocean (C. fernandeziana Phil.), and southern Venezuela and the Guiana Shield (C. linearis N.E.BR.). 5 subgenera.

Chusquea has the widest altitudinal (sea level to over 4,000 m) and latitudinal range of any bamboo genus; species are characteristic of montane forests, including cloud and elfin forests, but a number also grow in humid subpáramos, páramos, and grasslands; a few species (e.g., C. simpliciflora Munro) are found in lowland tropical forests, while other species at higher latitudes (both north and south) grow in more temperate forests, such as the S beech forests of Chile and the pine-oak forests of Mexico.

§ subg. Chusquea ‣ c. 90 spp., widely range of genus.

§ subg. Magnifoliae ‣ 10 spp., Central America, northern and central Andes, Trinidad, Guyana, and N Brazil.

§ subg. Platonia ‣ 11 spp., upper montane forest and páramos of Ecuador and Colombia, with C. fimbriligulata extends into N Peru; some Chusquea from this subgenus are most remarkable for the incredible leaf sizes attained in some spp.; in C. spectabilis L.G.Clark., from Venezuela to Ecuador, leaf blades can reach 3–4 m in length, the largest leaves known in the grass family; C. aristata Munro from Colombia to Peru has the highest known elevation for any bamboo, growing at altitudes up to 4,300 m in Ecuador.

§ subg. Rettbergia ► 11 spp., Brazilian montane and Atlantic forests, except C. arachniformis L.G.Clark & Londoño, which is endemic to montane forests of NW Colombia.

§ subg. Swallenochloa ‣ 52 spp., usually open habitats, often at high elevation; widespread, but most diverse in subpáramos and páramos of the northern Andes, Central America, and campos de altitude of E Brazil.

■ SUBTRIBE ARTHROSTYLIDIINAE - outsider Tibisia (3; Caribbean).

11. Actinocladum McClure ex Soderstrom. Fire-resistant bamboo. Only one sp., A. verticillatum (Nees) McClure ex Soderstr., from the savannas of C Brazil and Bolivia (cerrado) inhabiting open grasslands; it is adapted to survive a prolonged dry season and fire due to its thick-walled, pith-filled culms, thick coriaceous leaves, and indurate scales which protect the specialized buds.

12. Alvimia C.E.Calderón ex Soderstr. & Londoño. Slender and vining bamboo, with fleshy fruits (unique with some Guadua and Olmeca among bamboos in New World) and pseudospikelets. Three spp., which inhabits the atlantic sandy coastal shrublands (restingas) or white sand coastal forest in the state of Bahia in Brazil between 40-100 m elevation, two of them are rare plants in Brazil, by Plantas Raras do Brasil’s book.

13. Athroostachys Benth. (inc. Atractantha p.p.) Rhizomes pachymorph, short-necked; culms initially erect, becoming scandent to clambering on the neighboring vegetation. Two spp., endemics to E Brazil in Bahia, Minas Gerais, Rio de Janeiro and Espírito Santo states.

14. Atractantha McClure. (exc. Athroostachys p.p.) Vining and scandent culms, rhizomes pachymorph and short-necked; culms slender and hollow to solid and with peripheral air canals in some species. 5 spp., four inhabit the atlantic sandy coastal shrublands (restingas) and Atlantic forest from Bahia to Espírito Santo, at elevations from 0 to 650 m, and A. amazonica Judz. & L.G. Clark occurs in the Amazon region of Brazil, Colombia and Venezuela, at elevations from 80-100 m in wet, lowland, seasonally flooded forest (igapo).

15. Arthrostylidium Rupr. Scandent, slender culms and erect culm leaf blades. 30 spp., nine endemics to Caribbean, three only in Central America and Mexico, and 18 remaining in South America, from Venezuela to Bolivia and Brazil (5, three endemics, one of them, from Pará state, is a rare plant in Brazil, by Plantas Raras do Brasil’s book).

16. Aulonemia Goudot. (exc. Aulonemiella p.p.) Reflexed culm leaf blades, well-developed fimbriae on the culm lead margins and foliage leaf sheaths, paniculiform inflorescences, and multi-flowered spikelets. 49 spp., one in Mexico, one in Central America, and 47 in South America (one of then also in Central America), in wet, usually montane forests and paramos from S Mexico through Central and N South America to Bolivia, disjunct in SE Brazil; 16 spp. in Brazil, A. deflexa (N.E.Br.) McClure in forests of N Brazil, Venezuela and Guyana, and 15 endemics to SE Brazil (one of them, from Minas Gerais state, is a rare plant in Brazil, by Plantas Raras do Brasil’s book), mainly restrictes of Atlantic Forest.

All the 11 spp. from SE Brazil are endemic, and other occurs in northern region, shared with Guyana; among them, only A. xerophylla P.L. Viana & Filg. from Distrito Federal and Goias, A. aristulata (Döll) McClure from Bahia to Santa Catarina state, and the unusual A. effusa (Hack.) McClure from Bahia and Minas Gerais, are not considered endemic to the Atlantic forest biome.

17. Aulonemiella L.G. Clark, Londoño, C.D. Tyrrell & Judz. (inc. Aulonemia p.p.) Delicate, scandent to scrambling wood bamboos; rhizomes not seen. Two spp. from high mountains of Colombia and Ecuador.

18. Cambajuva P.L. Viana, Filg. & L.G. Tall bamboos, rhizomes pachymorph, short necked; culms woody, erect; culm leaves clearly differentiated from the foliage leaves; synflorescence paniculate, spiciform; spikelets consisting of two glumes. One woody bamboo, C. ulei (Hack.) P. L. Viana, L. G. Clark & Filg., endemic to coastal highlands (often with snow) along the Serra Geral of Santa Catarina and Rio Grande do Sul states in S Brazil, in peaty bogs or rarely in riparian vegetation.

19. Colanthelia McClure & L.B.Sm. Delicate, climbing and hanging bamboos. 10 spp., endemic to Atlantic Forest (sea level to taller mountains) in SE & S Brazil, one of them, from Minas Gerais state, is a rare plant in Brazil, by Plantas Raras do Brasil’s book, with C. rhizantha (Hack.) McClure in Missiones Province in adjacent NE Argentina.

20. Didymogonyx (L.G. Clark & Londoño) C.D. Tyrrell, L.G. Clark & Londoño. (off Rhipidocladum). Two spp. from mountains of Colombia and Venezuela.

21. Elytrostachys McClure. Culms thin-walled, erect below and clambering and vining above. Two spp., distributed from Honduras to Colombia and Venezuela.

22. Filgueirasia Guala. Strongly caespitose bamboos, vegetative clumps 0.30-3.21 m in diameter; rhizomes compact; culms erect, green or glaucous; inflorescence branches secund, short spikelets, the florets 3-15 per spikelet. Two spp., in open areas in semi-arid savannas from Bahia to Mato Grosso do Sul states, inc. Minas Gerais.

23. Glaziophyton Franchet. Herbs up to 2 m tall, culms typically leafless, almost herbaceous, and aggregated in dense clumps, Juncus-like. Only one sp., G. mirabile Franch., highly very narrow endemic and a rare plants in Brazil, by Plantas Raras do Brasil’s book, known only from seven small populations in the misty summits of mountains in the Serra dos Orgãos, at grasslands fields, in Rio de Janeiro state in SE Brazil, in Petroplis and Nova Iguazu municipalities.

The peculiar juncoid nature of the culm internodes septate lumina in Glaziophyton apparently has not been described for any other known bamboo species.

24. Merostachys Spreng. Erect, arching apically, reflexed culm leaf blades, fan-shaped branch complements, and racemiform inflorescences. 59 spp., one only Mexico, another only Central America, one from Brazil and Central America, 58 only South America, 53 in Brazil, 50 endemics (two shared Argentina and Paraguay and 11 as rare plants in Brazil, by Plantas Raras do Brasil’s book, in several states from Rondonia to Rio Grande do Sul); others species occur also in Guyana, Venezuela, Peru and Bolivia.

M. tatianae Santos-Gonçalves, Carvalho-Okano & Filgueiras (from Rio Doce State Park, Minas Gerais) has the longest internodes so far recorded for the genus (37–108 cm long).

25. Myriocladus Swallen. Erect habit and thick leaves. 12 spp., 11 in Venezuela: two up to Mount Neblina in Brazil, two up Mount Araca also in Brazil, one up to Guyana, and M. caburaiensis E. Afonso & P.L. Viana endemic to Mount Roraima in Roraima state in N Brazil, near Guyana border.

26. Rhipidocladum McClure. (exc. Didymogonyx) Erect, arching apically or clambering and hanging, and are characterized by fan-shaped branch complements, racemiform inflorescences and thin-walled culms. 21 spp. from NE Mexico and Trinidad to NW Argentina and central Brazil, 14 in South America, two in Brazil, both widely distributed, none endemics.

■ SUBTRIBE GUADUINAE ‣ outsiders Tibisia (3; Caribbean), Olmeca (5; Mexico).

27. Apoclada McClure. Only one sp., A. simplex McClure & L.B. Sm., a beautiful tall and luxuriant bamboo of the mesic forests of SE Brazil (Sao Paulo state and one enclave of Araucaria forest in Santa Catarina state).

28. Eremocaulon Soderstr. & Londoño. 5 spp. from Bahia to Sao Paulo states, Mato Grosso, and Amazon rainforest, all endemics to Brazil, two of them, one in Bahia and another in São Paulo state, are rare plants in Brazil, by Plantas Raras do Brasil’s book.

29. Guadua Kunth. Wood bamboos, sometimes with fleshy fruits (unique with some Alvimia and Olmeca among bamboos in New World). 34 spp., the largest in Neotropcs; Mexico to Argentina, 30 in South America, 20 spp. in Brazil, 5 endemics, two of them, from Goiás and Minas Gerais states, are rare plants in Brazil, by Plantas Raras do Brasil’s book.

Guadua boasts the tallest of the native New World bamboos (and Poales) in G. chacoensis (Rojas) Londono & P. M. Peterson (Paraná Basin) and G. angustifolia Kunth. from Mexico to Peru, and the most economically important bamboo in the Western Hemisphere. G. sarcocarpa Lodoño & Peterson is a tropical rainforet bamboo from Peru, highly expected occur in Brazil (Acre state), unknown for indigenous peoples in Amazon, and it’s the first report a one edible bamboo in New World; however, a variety of G. weberbaueri Pilger in same area also has fleshy fruits.

Guadua constitutes the most extensive bamboo forest in New World, covering more than 180,000 km² (approximately sized of UK) in the western Amazon rainforest; the largest area of bamboo in Brazil is in Acre state (8 millions hectares), with G. weberbaueri Pilger is the dominant species.

30. Otatea (McClure & E.W.Sm.) C.E.Calderón ex Soderstr. 13 spp., all endemic to Mexico and Central America region except by O. colombiana Ruiz-Sanchez & Londoño endemic Norte Santander in Colombia, and O. fimbriata Soderstr., in seasonally dry, tropical deciduous, subdeciduous, or oak forests, often on limestone-derived soils, from Mexico to Costa Rica, with populations also in N Colombia.

4.2 BAMBUSOIDEAE ▸ TRIBE OLYREAE (21/c 115) - three tribes, Buergersiochloinae (4/5; one genus in New Guinea and three endemics to Cuba) not occur in South America; members of Olyreae are small- to medium-sized, non-lignified, clumpforming, stoloniferous, or occasionally scandent plants (Olyra latifolia L.), with restricted vegetative branching and unisexual spikelets, endemic to the New World with two exceptions: Buergersiochloa is a rare monotypic endemic of New Guinea, while O. latifolia is a widely distributed American spp. that has presumably been introduced into Africa, where it is also widely distributed, and Sri Lanka.

■ SUBTRIBE PARIANINAE ‣ all genera in South America.

31. Eremitis Döll. Herbaceous bamboos, producing aerial flowers and scaly, cleistogamous subterranean synflorescences from the tips of whiplike ‘stolons’ that bury themselves in the leaf litter or soil. 16 spp. from Bahia, Espírito Santo, Minas Gerais and Rio de Janeiro states, in dense ombrophilous and semi-deciduous forests along the Atlantic forest; one of them, from Espírito Santo state, is a rare plants in Brazil, by Plantas Raras do Brasil’s book.

Only two genera of Poaceae - Amphicarpum Kunth fro E U.S.A. and Eremitis Döll - produce spikelets underground from positively geotropic shots, as well as above ground, from open panicles; these genura composes the only fully amphicarpic genus endemic to a country in New World. E. afimbriata F.M. Ferreira & R.P. Oliveira (S Espírito Santo state) and E. magnifica F.M. Ferreira & R.P. Oliveira (only E Minas Gerais state), species which display blue iridescence in their leaf blades, unique among bamboos worldwide.

32. Pariana Aubl. Cylindrical synflorescences with numerous (up to 36 or more) showy yellow or white stamens per male spikelet; it is generally agreed that these facilitate insect pollination by phorid flies and gall midges. 36 spp. from Nicaragua to Bolivia and Amazon Brazil (absent in tepuis, however); 33 in South America, 29 in Brazil, 13 endemics, two of them, from Pará e Maranhão states, are rare plants in Brazil, by Plantas Raras do Brasil’s book.

33. Parianella Hollowell, F.M. Ferreira & R.P. Oliveira. Monomorphic stems, glabrous to pilose; leaf blades 3–8 per culm, green on both blade surfaces; inflorescence one per culm, terminal, monomorphic, solitary and spiciform; male spikelets 2.0–3.2 mm long, glumes oblong-triangular, 0.9–2.0 mm wide. Three spp., endemics to Atlantic Rain Forest of S Bahia and C Espírito Santo state, Brazil, one of them a are rare plants in Brazil, by Plantas Raras do Brasil’s book.

■ SUBTRIBE OLYRININAE ‣ all genera in South America.

Although predominantly anemochoric, Poaceae have evolved other ways to disperse their diaspores, such as zoochory and also ballistochory (forcible dispersal), which is considered rare within this family, only known for the sister genera Raddia Bertol. (8, Brazil, one up to Venezuela), Sucrea Soderstr. (2, E Brazil) and Piresia (Reitzia) Swalllen. (9, northern South America), all South American berbaceous bamboos.

LINEAGE 1

34. Diandrolyra Stapf. Herbs; flowering culms typically bearing only one fully developed and specialized leaf blade, which is appressed to and overtops the single contracted inflorescence; this inflorescence is inconspicuous and completely hidden under the abaxial surface of the leaf blade when the plant is viewed from above, which makes Diandrolyra species appear to always be sterile. Three spp. endemic to the Atlantic Forest of Brazil in Bahia, Espírito Santo, Rio de Janeiro, Sao Paulo and Minas Gerais states.

35. Parodiolyra Soderstr. & Zuloaga. (exc. Taquara). 4 spp. from Costa Rica to Bolivia and Bahia, one widely distributed, three restricted (Venezuela, Colombia, Brazil); mesophytic; shade spp. and spp. of open habitats; in low and mid-elevation forests and savannas; 3 spp. in Brazil, one endemic.

36. Raddiella Swallen. 8 spp. from Caribbean and tropical America; three only in Brazil (one of them, from Rondônia state, is a rare plant in Brazil, by Plantas Raras do Brasil’s book); Colombia and French Guiana one endemic each, one from Panamá to SE Brazil and Bolivia, and two from Guiana Shield (one of then in Brazil). R. vanessieae Judz., endemic to French Guiana, is smallest known bamboo, flowering at only 2 cm high; the only known annual bamboo is found in this genus is R. minima Judz. & Zuloaga, from the state of Mato Grosso state in C Brazil.

37. Taquara I.L.C.Oliveira & R.C.Oliveira. (off Parodiolyra) Perennials, culms erect, sometimes learning on sorrounding vegetations. Two spp. widely distributed in South America, from E Colombia and Venezuela to the Atlantic coast of Brazil (only T. micrantha (Kunth) I.L.C.Oliveira & R.C.Oliveira) and the E Andes along forest edges and interior of forests.

LINEAGE 2

38. Brasilochloa R.P.Oliveira & L.G.Clark. (off Sucrea) Herbs with tuberous roots and synflorescences congested to spiciform, perennial, monoecious, caespitose, erect; leaves distributed along the culms. Only one sp., B. sampaiana (Hitchc) R.P.Oliveira & L.G.Clark, known only from Atlantic Forest of Rio de Janeiro and Espírito Santo states in E Brazil, and a rare plants in Brazil, by Plantas Raras do Brasil’s book.

39. Raddia Bertol. Herbs; after flowering, the drying margins and bases of the female glumes contract and twist, ejecting the mature floret (with enclosed fruit) up to a meter from the parent plant in a dispersal mode known as ballistochory. 9 spp., 8 endemics to Brazil (in forest understories at elevations of 0-500 (-1000) m, one of them, from Espírito Santo state, is a rare plants in Brazil, by Plantas Raras do Brasil’s book), and R. guianensis (Brongn.) Hitchc. within and reaching from French Guiana, Trinidad and Venezuela, and also NE Brazil.

40. Sucrea Soderstr. (exc. Brasilochloa) Rhizomatous grass (appearing cespitose) of shaded forests; culms herbaceous, erect, not branching, with broad persistent papyraceous sheaths and thick prop roots. Two spp., endemic to Atlantic Forest in E Brazil.

LINEAGES 3 + LINEAGE 4

41. Agnesia Zuloaga & Judz. Two spp., A. lancifolia (Mez) Zuloaga & Judz, from Colombia and N Brazil, and A. loretensis (Mez) J. R. Grande from Brazil, Peru, Colombia and Bolivia.

42. Arberella Soderstr. & C. E. Calderon. 8 spp. from Costa Rica to N South America (4), two in Brazil, none endemics – however, one is assignated as rare plant in Brazil, by Plantas Raras do Brasil’s book, known only in Bahia state.

43. Cryptochloa Swallen. 9 spp., 7 from Mexico to Ecuador, one from over N South America, and one from Brazil and adjacent French Guiana; 5 spp. in South America.

44. Froesiochloa G.A. Black. Herbs, perennials; the flowering culms leafy, culms herbaceous. Only one sp., F. boutelouoides G.A.Black, endemic to the Guiana Shield, in Pará state in N Brazil and Guyana.

45. Lithachne P. Beauv. Small herbs, with tooth-shaped female florets that are unique in the grass family. 4 spp., L. pauciflora (Sw) P. Bauv. widely distributed (Mexico to Paraguay and Caribbean), and Cuba, Brazil and Honduras one endemic each.

46. Maclurolyra C.E.Calderón ex Soderstr. Only one sp., M. tecta C.E.Calderón ex Soderstr, from Panamá to N Colombia.

47. Olyra L. 23 spp., Mexico to Argentina, and Africa (a single disjunct, Benin to Zimbabea, Comoros, Madagascar); 22 spp. in South America, 19 in Brazil, 8 endemics, six of them, from several states, are rare plants in Brazil, by Plantas Raras do Brasil’s book.

48. Piresia Swallen. (inc. Reitzia) Herbs; dimorphic culms; aerial ones bear broad and flat leaf blades at the top and rarely develop an inflorescence, and decumbent ones usually bear reduced leaves or bladeless leaf sheaths, with inflorescences that are raceme-like, few-flowered, and often hidden under the litter. 7 spp., French Guiana to Peru and Brazil up to east coast; all species in Brazil, two endemics.

49. Rehia Fitjen. Only one sp., R. nervata Fijten, from Guyana to French Guiana and N Brazil.

5. SUBFAMILY POOIDEAE (186–188/3.630–3.680) ▸ 12 lineages, Brachyelytreae (1/3; Japan, SE China, E U.S.A.), Nardeae (1/1; Europe except Mediterranean, W Asia), Lygeeae (1/1; Mediterranean, North Africa), Duthieeae (7/11; temperate and subtropical regions in the Old World, Mexico), Phaenospermateae (1/1; Assam, SE China (inc. Taiwan), Korean Peninsula, Japan), Ampelodesmeae (1/1; Mediterranean), Diarrheneae (1/5; E Asia, North America) absent in South America.

The species are known as the ‘cool season’ or ‘pooid’ grasses and all are C3 and distributed in temperate climates; 121 genera in the New World, 82 native and 39 cultivated. A small area of 490 km² of Sao Joaquin national Park in S Santa Catarina state in S Brazil has 55 spp. of this subfamily; Filgueiras et al. (2012) reported 135 spp. for Brazil, 25 of which are endemic (in genera Melica, Piptochaetium, Nassella, Agrostis, Calamagrostis, Poa, and several Calothecinae; the highest spp. richness occurs in the natural grasslands of Rio Grande do Sul state, a few spp. extending to the high-altitude grasslands (campos de altitude) or, in smaller numbers, to the rocky grasslands (campos rupestres) of SE Brazil; the main winter crops of southern Brazil belong to this subfamily, e.g. wheat, barley, rye and oat.

5.1 POOIDEAE ▸ TRIBE MELICEAE (8/133) - two subtribes; Brylkiniinae (E Russia, China, Japan) does not occur in South America; among Melicinae, outsiders are Lycochloa (1; Lebanon), Schizachne (1; arctic and temperate Russia, Siberia, Central Asia, Mongolia, China, Korean Peninsula, temperate and arctic North America, mountains in SW U.S.A.), Pleuropogon (6; W U.S.A., one species, arctic circumpolar), Koordersiochloa (2; tropical and S Africa, Réunion, S India, mountains in Indonesia and Philippines).

50. Glyceria R.Br. 40 spp., temperate regions on the Northern Hemisphere, with their largest diversity in North America; 18 spp. in New World, three from Argentina southwards to S Brazil (G. multiflora Steud., none endemic), Colombia one endemic.

51. Melica L. Perennial rhizomatous, herbaceous, scadent or not, no tuberous. 80 spp. of temp. Northern Hemisphere to Mexico, W & S South America to SE & S Brazil; 48 spp. in New World, 31 spp. in South America (12 in Brazil, three endemics, one of them, from Rio Grande do Sul state, is a rare plant in Brazil, by Plantas Raras do Brasil’s book), all but one (this up to Colombia) in Bolivia southwards; mesophytic to xerophytic. Absent in Venezuela.

52. Triniochloa Hitchc. 6 spp. from tropical America, two in South America, T. stipoides (Kunth) Hitchc from Mexico to Bolivia and Venezuela, and T. andina Luces from Colombia and Venezuela.

5.2 POOIDEAE ▸ TRIBE STIPEAE (23–25/545–550) - outsiders Macrochloa (1; Mediterranean), Stipa (c 150; Europe, North Africa, temperate and drier subtropical regions in Asia, with their highest diversity in SW and S Asia), Psammochloa (1; the Gobi Desert), Trikeraia (4; Pakistan to China), Ptilagrostis (13; Russia to China), Piptatheropsis (5; Canada, U.S.A.), Hesperostipa (5; Canada, U.S.A., northern Mexico), Patis (3; temperate regions on the Northern Hemisphere), Achnatherum (c 40; Europe, North Africa, temperate regions in Asia east to Japan), Eriocoma (3; North America), Celtica (1; W Mediterranean), Anemanthele (1; New Zealand), Austrostipa (c 60; Australia, one species also in New Zealand), Oloptum (1; Mediterranean, SW Asia), Orthoraphium (1; Himalayas), Stipellula (5; warm-temperate and subtropical regions in the Old World), Timouria (4; Pakistan and Central Asia to Mongolia and China).

Among the genera considered, only Piptochaetium, Austrostipa, and Hesperostipa were resolved as monophyletic, while Achnatherum, Amelichloa s.l., Anatherostipa, Jarava, and Nassella were polyphyletic, and Aciachne was polyphyletic or paraphyletic. As a result, Amelichloa can be restricted to a monophyletic group if including A. brachychaeta, A. ambigua, A. clandestina, and A. caudata, or it should be considered within Nassella. The phylogenetic position of species of Aciachne suggests inbreeding and outbreeding events with species of Anatherostipa, Ortachne, and Hesperostipa.

53. Aciachne Benth. Low cushions. Three spp., high mountains from Costa Rica to NW Argentina and Venezuela.

54. Amelichloa Arriaga & Barkworth. 5 spp., 4 spp. found Chile, Argentina and Uruguay, and one endemic to N Mexico.

55. Anatherostipa (Hack. ex Kuntze) Peñailillo. Cespitose plants; prophyllum 2–5 cm, 2-awned; fundamental cells of lemma epidermis with thick, sinuous walls; paleas subeqalto lemmas, veined, often with hairs. 10 spp., nine from Peru to Chile and Argentina, and A. hans-meyeri (Pilg.) Peñail. up to Central America.

56. Jarava Ruiz & Pav. (inc. Stipa p.p.). 29 spp. from W & S South America, one up to Mexico; only J. plumosa (Spreng.) S.W.L. Jacobs & J. Everett in Brazil (only in Santa Catarina and Rio Grande do Sul states), no endemic.

57. Nassella (Trin.) É.Desv. 117 spp., 112 in South America, 21 in Brazil (5 endemics), centered in Peru southwards, some up to Venezuela and Costa Rica, Central and North America; it is well represented in two South American regions: the central Andean region (Peru, Bolivia, N and C Chile, and NW Argentina), and from N Patagonia, the Pampas, central and NE Argentina to Uruguay, and S Brazil; Argentina includes 70 spp., and more than half of them are concentrated in the NW region of the country (Jujuy, Salta, Tucuman, Catamarca, and La Rioja).

58. Ortachne Nees ex Steud. (inc. Lorenzochloa) Glumes shorter than floret; awn poorly demarcated; leaves filiform, not stiff; paleas subequal to lemmas, veined, sometimes with hairs. Three, one from Venezuela to Bolivia, and two in from Cono Sur.

59. Pappostipa (Speg.) Romasch., P.M.Peterson & Soreng. 30 spp., 28 spp. from Chile and Argentina, three of then up to Bolivia and Peru, and two in NW Mexico and SW U.S.A.

60. Piptochaetium J.Presl. Mesophytic to xerophytic. 36 spp. from semi-arid grasslands of E. Canada to Guatemala, 29 spp. in Venezuela to W. South America, SE & S Brazil to S. South America; 10 spp. in Brazil, two endemics, one of them, from Santa Catarina state, is a rare plants in Brazil, by Plantas Raras do Brasil’s book.

5.3 POOIDEAE ▸ TRIBE BRACHYPODIEAE (1/18) - a single genus.

61. Brachypodium P.Beauv. 17 spp., 15 in Old World, two in New World, B. mexicanum (Roem. & Schult.) Link from Mexico to Bolivia, and one endemic to N Mexico.

5.4 POOIDEAE ▸ TRIBE TRITICEAE (20/510–520) - 4 subtribes, Littledaleinae (1/4; Central Asia and Tibet to W China) and Triticinae (4/c. 55; Canary Islands, Mediterranean and SW Asia to China) do not occur in South America.

■ SUBTRIBE BROMINAE ‣ a single genus.

62. Bromus L. Mesophytic, or xerophytic; shade spp. and spp. of open habitats. 150 spp., temperate regions on the Northern Hemisphere, Mediterranean, southern Africa, tropical Mountains in South America; commonly adventive; 58 spp. in New World, 25 in South America (17 from Peru to Argentina and Chile, two of then reaches Juan Fernandes; five from tropical Andes to southern South America, one scarsely disjunct), only 3 native to Brazil (two platine, one over neotropics), none endemic.

■ SUBTRIBE HORDEINAE ‣ outsiders Agropyron (13; temperate regions in the Old World), Australopyrum (5; New Guinea, E Australia, Tasmania, New Zealand), Crithopsis (1; Crete and Libya to Iran), Eremopyrum (4; S Europe, Mediterranean, Morocco to W China), Henrardia (2; Türkiye and Iran to Central Asia), Heteranthelium (1; Türkiye to Pakistan), Hordelymus (1; Europe and North Africa to the Caucasus), Peridictyon (1; the Balkan Peninsula), Psathyrostachys (10; E Mediterranean to Central Asia), Secale (9; E Europe and eastwards to Central Asia, Mediterranean, the Middle East, Roggeveld in W Cape), Taeniatherum (1; the Iberian Peninsula, Mediterranean to Pakistan and C Asia).

63. Elymus L. c 155 spp., temperate regions on the Northern Hemisphere to C & S America; 49 spp. in New World, 8 in South America, 6 in Argentina, 3 up to Chile; one from Costa Rica to Peru, other only Peru, a third from Peru to S Argentina.

64. Hordeum L. Annual or perennial, ceaspitose herbaceous, unbranched above. 40 spp. of temp. Eurasia, Macaronesia, N & S Africa, N. America to Guatemala, Bermuda, Peru to S. South America, commonly adventive; mostly in dry soils. 24 spp. in New World, 18 from South America, 12 only in Argentina and Chile, sometimes reaching Juan Fernandes Is, two Peru southwards, H. euclaston Steud., from S Brazil to Argentina; H. flexuosum Nees ex Steud., from Uruguay to Argentina; H. stenostachys Godr. from S Brazil to Argentina, S. Africa, and another disjunct northern hemisphere and Argentina.

65. Leymus Hochst. (inc. Eremium). 55 spp., subarctic & temp. Northern Hemisphere to N. Mexico, 13 in New World, with L. erianthus (Phil.) Dubc. in Chile and Argentina.

5.5 POOIDEAE ▸ TRIBE POEAE (121/2,395–2,430) - 7 unplaced genera (only one in New World) and 27 subtribes; of them, Brizinae (2/5; Europe, NW Africa, temperate Asia, Mediterranean), Scolochloinae (2/2; temperate regions on the Northern Hemisphere, SE New South Wales, Victoria, Tasmania), Sesleriinae (6/40; Europe, NW Africa, Libya, W Asia to the Caucasus and Iran), Dactylidinae (2/3, Mediterranean, the Middle East to Pakistan, temperate Asia, Macaronesia), Cynosurinae (1/9, Europe, Mediterranean, SW Asia), Ammochloinae (1/3, Mediterranean, SW Asia), Parapholiinae (8/22, Mediterranean, Macaronesia, SW Asia to Iran, southern Asia, temperate Eurasia), Miliinae (1/6, Europe, temperate Asia, E North America), Beckmanniinae (4/6, temperate regions on the Northern Hemisphere) and Ventenatinae (5/20, southern Europe, Mediterranean to the Caspian Sea, SW Asia to Afghanistan) not occur in South America.

■ UNPLACED

66. Nicoraepoa Soreng & L.J.Gillespie. 7 spp., restricted in wetlands of Chile and Argentina.

■ SUBTRIBE TORREYOCHLOINAE ‣ outsider Torreyochloa (5; NE Asia, Canada, U.S.A.).

67. Amphibromus Nees. (inc. Helictotrichon) 12 spp., mainly Australia and New Zealand, with A. quadridentulus (Döll) Swallen and A. scabrivalvis (Trin.) Swallen in S Brazil to NE. Argentina, Peru, Chile and Bolivia.

■ SUBTRIBE PHALARIDINAE ‣ a single genus.

68. Phalaris L. Annual or perennials herbs, tuberous or not. 17 spp., temperate regions on the Northern Hemisphere, Mediterranean, Andean and S South America; 7 spp. in South America, two in S Brazil, none endemics; absent in Venezuela.

■ SUBTRIBE AVENINAE ‣ outsiders Arrhenatherum (7; Europe, Mediterranean, N and W Asia), Avena (22; Europe, Mediterranean, North Africa to Ethiopia, SW Asia), Sphenopholis (7; Canada, U.S.A., Mexico, Hispaniola, Hawaii), Trisetopsis (20–25; tropical to S Africa, Madagascar, Yemen, Himalayas, C China, southern India and Sri Lanka, Sumatra, Java), Trisetaria (16; Mediterranean to W Himalayas), Gaudinia (4; Azores, Mediterranean), Graphephorum (2; North America, Mexico, Central America), Tzveleviochloa (3; Assam, N Burma, C China), Lagurus (1; Mediterranean), Tricholemma (2; Morocco, Algeria).

69. Koeleria Pers. 47 spp., temp. Eurasia, NW Africa, Cameroon, Ethiopia to S. Africa, New Zealand, N. America and S. South America, Falkland Is. 10 spp. in South America, from Colombia to Argentina and Uruguay, Chile and Bolivia, mainly Argentina.

70. Peyritschia E.Fourn. (inc. Calamagrostis p.p., Deyeuxia p.p.). 31 spp. throughout Mexico and Central America with 8 extending into South America as far south as Bolivia.

71. Rostraria Trin. 12 spp., 11 in Macaronesia, Medit. to Sahara and India, Caucasus to C. Asia and W Himalaya, and R. trachyantha (Phil.) Soreng from Peru to N & C Chile.

72. Trisetum Pers. Perennials, caespitose, sometimes shortly rhizomatous; culms 5-300 cm tall, erect to geniculate at base, glabrous or pubescent. 68 spp. in both hemispheres, 33 in New World, 15 in South America from Peru to Argentina.

■ SUBTRIBE ANTHOXANTHINAE ‣ a single genus.

73. Anthoxanthum L. (inc. Hierochloe). 47 spp., temperate and alpine regions on the Northern Hemisphere, tropical mountains in Africa and Asia, 11 in New World, 7 from North America to Colombia and Venezuela (A. davidsei (R.W. Pohl) Veldkamp and A. mexicanum (Rupr. ex E. Fourn.) Mez up to South America) and four from Ecuador to Tierra del Fuego, mainly in Argentina.

■ SUBTRIBE ECHINOPOGONINAE ‣ outsiders Ancistragrostis (1; New Guinea, tropical Australia), Dichelachne (9; E Malesia to New Guinea and Australia, New Zealand), Echinopogon (7; New Guinea, Australia, New Zealand), Pentapogon (1; SE South Australia, SE New South Wales, Victoria, Tasmania).

74. Relchela Steud. Only one sp., R. panicoides Steud, from C & S Chile to S. Argentina.

■ SUBTRIBE CALOTHECINAE ‣ ten genera, all in Brazil.

75. Boldrinia L.N.Silva. (off Chascolytrum) Only one sp., B. parodiana (Roseng., B.R.Arrill. & Izag.) L.N.Silva, from S Brazil to Uruguay.

76. Chascolytrum Desv. (exc Boldrinia, Calotheca, Chascolytrum, Erianthecium, Lombardochloa, Microbriza, Poidium, Rhombolytrum, Rosengurttia) 10 spp., Bolivia, Chile and Argentina one endemic each, 4 endemics to Brazil, two from S Brazil to NE Argentina, Paraguay and Uruguay, and C. subaristatum (Lam.) Desv. from Mexico to Guatemala, Colombia to Chile and Argentina, Paraguay, Uruguay and S Brazil.

77. Condilorachia P.M.Peterson, Romasch. & Soreng. Three spp., C. brasiliensis (Louis-Marie) P.M.Peterson, Romasch. & Soreng, and C. juergensii (Hack.) P.M.Peterson endemics to S Brazil, and C. bulbosa (Hitchc.) P.M. Peterson, Romasch. & Soreng from Argentina and Chile.

78. Erianthecium Parodi. (off Chascolytrum) Only one sp., E. bulbosum Parodi, from S Brazil to Uruguay.

79. Calotheca Desv. (off Chascolytrum) Only one sp., C. brizoides (Lam.) P.Beauv., S Brazil to Argentina, Chile and Urugauy

80. Laegaardia P.M. Peterson, Soreng, Romasch. & Barberá (off Calamagrostis). Only one sp., L. ecuadoriense (Lægaard) P.M. Peterson, Soreng, Romasch. & Barberá, endemic to Ecuador where it has been found in the Provinces of Azuay, Chimborazo, Loja Napo, and Pichincha.

81. Lombardochloa Roseng. & B.R.Arrill. (off Chascolytrum) Only one sp., L. rufa (J.Presl) Roseng. & B.R.Arrill., from Peru, S Brazil to NE Argentina.

82. Microbriza Parodi ex Nicora & Rúgolo. (off Chascolytrum) Two spp., M. brachychaete (Ekman) Parodi ex Nicora & Rúgolo and M. poimorpha (J.Presl) Parodi ex Nicora & Rúgolo, S Brazil to Argentina, Paraguay adn Uruguay.

83. Paramochloa P.M. Peterson, Soreng, Romasch. & Barberá (off Calamagrostis). Two spp. from paramos of Colombia, Ecuador and Venezuela.

84. Poidium Nees. (off Chascolytrum) Six spp., three endemics to SE & S Brazil, P. calotheca (Trin.) Matthei in Colombia, SE & S Brazil to NE Argentina, Paraguay and Uruguay, P. juergensii (Hack.) Matthei disjunct in Colombia and SE & S Brazil, and P. uniolae (Nees) Matthei from Bolivia to S Brazil and Argentina

85. Rhombolytrum Link. (off Chascolytrum) Two spp., R. monandrum (Hack.) Nicora & Rúgolo from S Brazil to Uruguay, and R. rhomboideum Link endemic to Chile.

86. Rosengurttia L.N.Silva. (off Chascolytrum) Only one sp., R. monandra (Hack.) L.N.Silva, Colombia to NW Argentina, S Brazil.

■ SUBTRIBE AGROSTIDINAE ‣ outsiders Gastridium (2; W Europe, Macaronesia, Mediterranean to the Caucasus and Iran), Hypseochloa (2; Mount Cameroon, mountains in Tanzania), Limnodea (1; S U.S.A.), Triplachne (1; Mediterranean). In a molecular phylogeny based on four gene regions (ITS, rpl32-trnL spacer, rps16-trnK spacer, and rps16 intron), the South American species of Calamagrostis and Deyeuxia form a clade sister to the Mexican and Central American species of Trisetum, Calamagrostis, and Peyritschia (most transferred below to Peyritschia), within the Koeleriinae clade B of the Aveninae. Within the Cinnagrostis clade are two subclades, one containing species formerly placed in Calamagrositis sect. Chamaecalamus Pilg. [more recently placed in Deyeuxia sect. Chamaecalamus], which is sister to the remaining species in the genus.

87. Agrostis L. (exc. Podagrostis p.p., inc. Bromidium) Herbaceous, annual or perennial, stoloniferous, ceaspitose or decumbent. 180 spp., temperate regions on both hemispheres, tropical mountains, Macquarie Island, subantarctic South America, Falkland Islands, Crozet Islands, Kerguelen, Prince Edward Islands; 75 spp. in New World, 48 in South America, widely distributed; 6 spp. in Brazil, two endemics.

88. Calamagrostis Adans. (exc. Peyritschia p.p., Laegaardia, Paramochloa, Cinnagrostis). 252 spp. in temp. & subtrop. to trop. mts. worldwide; several species in New World, six in Latin America, two in South America, from Venezuela to Ecuador.

89. Cinnagrostis Griseb. (inc. Calamagrostis p.p., Leptophyllochloa, Deyeuxia p.p.). 77 spp. from South and Central America (only one up to S Mexico), 61 in South America, 4 up to S Brazil, one endemic, known only from Santa Catarina state, also as a rare plant in Brazil, by Plantas Raras do Brasil’s book.

90. Lachnagrostis Trin. 32 spp., 31 in Eritrea to S Africa, SW Arabian Pen., Lesser Sunda Is. to Australasia, Easter I., and L. sodiroana (Hack.) Rúgolo & A.M. Molina restricted to C. Andes in Ecuador and Peru.

91. Polypogon Desf. c 18 spp., Old World, Australia, U.S.A. to C. America, Venezuela to W South America, SE & S Brazil to S. South america, Tristan da Cunha. 11 spp. in New World, all South America (three up to Central and North America), mostly spp. widely distributed, 3 in southern South America, and 4 in Brazil, none endemics.

92. Podagrostis (Griseb.) Scribn. & Merr. (inc. Agrostis p.p.). 11 spp., four from North America, three from Mexico and Central America, and four in South America: three in tropical Andes from Venezuela to Peru, another in Cono Sur; absent in Bolivia.

■ SUBTRIBE AIRINAE ‣ outsiders Aira (8; Europe, Mediterranean to Iran), Antinoria (2; Mediterranean), Corynephorus (5; Europe, Mediterranean to Iran), Helictochloa (22; C and S Europe, Mediterranean, Canary Islands, North Africa, the Balkan Peninsula, E Europe to Crimea and the Caucasus, the Middle East, one sp. in Russia, Ukraine, Central Asia, Mongolia, China and North America), Periballia (3; Mediterranean).

93. Avenella Bluff ex Drejer. Only one sp. in New World, A. flexuosa (L.) Drejer, from Europe to Japan, tropical Africa, Greenland to C & E U.S.A., S. South America to Falkland Is.

■ SUBTRIBE HOLCINAE ‣ outsider Holcus (10; Europe, Mediterranean, North and S Africa, SW Asia).

94. Vahlodea Fr. Only one spp., V. atropurpurea (Wahlenb.) Fr., disjunct from Greenland to U.S.A., and Argentina and Chile.

■ SUBTRIBE ARISTAVENINAE ‣ a single genus.

95. Deschampsia (L.) P. Beauv. (inc. Deyeuxia p.p., Calamagrostis p.p.) Plants perennial or annual, often forming tussocks, sometimes cushions; culms erect, usually, 150 cm tall, sometimes slightly bent at the base, slender to stout. 47 spp., temperate and polar regions on both hemispheres, in Andes to Tierra del Fuego, Falkland Islands, Antarctic Peninsula and adjacent islands, high altitude tropics or restricted range endemics: New Zealand (4), Tristan da Cunha Islands (4), Madeira Islands (2), Pamir Mountains (2), mountains of Indonesia and Papua New Guinea (1), Cordillera Central of Hispaniola (1), Azores Islands (1), mountains of Central Mexico (1), Hawaii (1) and Kilimanjaro mountains in tropical Africa (1).

22 spp. in South America, mainly Cono Sur, Peru and Ecuador one endemic each, D. eminens (J. Presl) Saarela up to Colombia, D. caespitosa (L.) P. Beauv. in wet meadows, bogs, and along streams and creeks in the Andes from Bolivia (?) to Tierra del Fuego; isolated populations of this species can also be found in high places of SE Brazil, at about 1,000 m elevation.

D. antarctica E. Desv. is one of only two vascular plant species known to be native to Antarctica, thus the southernmost monocot of the World, and several adjacent southern islands and in the southern part of the American continent.

■ SUBTRIBE LOLIINAE ‣ outsiders Castellia (1; Macaronesia, Mediterranean, North Africa to Pakistan), Leucopoa (10; temperate Asia, Canada, U.S.A.), Drymochloa (5; Europe, W Mediterranean, North Africa), Lolium (12; temperate regions on the Northern Hemisphere, Canary Islands, Mediterranean), Patzkea (1–5; Europe, Mediterranean), Pseudobromus (6; tropical and subtropical Africa).

96. Festuca L. (inc. Dielsiochloa, Megalahcne, Podophorus) Graminoids, commonly adventive; helophytic (rarely), mesophytic (mostly) or xerophytic (rarely); halophytic, or glycophytic. 629 spp., temperate, polar and alpine regions on both hemispheres, Tierra del Fuego, Falkland Islands, South Georgia Is., Kerguelen Is., Juan Fernández Is, Macquarie Island, tropical mountains; 221 spp. in New World, 156 in South America, three in Brazil, none endemics; hillsides, mountains, plains, meadows.

■ SUBTRIBE COLEANTHINAE ‣ outsiders Coleanthus (1; temperate Northern Hemisphere), Sclerochloa (2; Mediterranean, North Africa and the Middle E to C Asia and China).

97. Catabrosa P.Beauv. 7 spp., temp. Northern Hemisphere, one in Lesotho and KwaZulu-Natal, and two spp. in New World, only C. werdermannii (Pilg.) Nicora & Rúgolo in South America, from Bolivia to N & C Chile and Argentina.

98. Phippsia (Trin.) R. Br. Three spp., one in Subarctic to WC U.S.A., one in India, and P. wilczekii Hack. endemic to Mendoza in NW Argentina.

99. Puccinellia Parl. 120 spp., temperate and Arctic regions on the Northern Hemisphere, southern Africa, one in Australia, 11 in New World, all from Peru to Uruguay and Tierra del Fuego except one endemic to Ecuador.

■ SUBTRIBE POINAE ‣ a single genus.

100. Poa L. Annual or perennial; rhizomatous, stoloniferous, ceaspitose or decumbent herbs unbranched above. c. 500 spp., reich in weeds, high pastures, from coast up tallest mountains of world, few in sand places; 206 spp. in New World, 127 in South America (high diverse in western continent, eg. 89 in equatorial and tropical Andes, and 62 in Cono Sur); six spp. in Brazil, two endemics. Recent molecular phylogenetic studies have substantially influenced the classification of Poa, currently divided into five subgenera:

§ subg. Ochlopoa ‣ 13 spp., worldwide, mostly in N Africa, C and SW Asia, and Europe; two sections in New World: Alpinae and Parodiochloa.

§ subg. Pseudopoa ‣ 5 spp., NE Africa, C and SW Asia, Europe.

§ subg. Poa ‣ c. 400 spp., over range of genus; two clades corresponding to supersections Poa and Homalopoa. Poa supersect. Homalopoa is a large and diversified clade in terms of both species and sections, and includes about half of the species in Poa; it is currently divided into ten sections worldwide along with the Punapoa informal assemblage. The New World represents a major centre of diversity with 144 endemic Homalopoa species.

§ subg. Stenopoa ‣ c. 40 spp., Asia, Europe, North America, a few species in New World, within sect. Pandemos, Secundae and Stenopoa.

§ subg. Sylvestres ‣ endemic to North America.

■ SUBTRIBE PHLEINAE ‣ a single genus.

101. Phleum L. Annual or perennials, rhizomatous, ceaspitose, stoloniferous herbs. 15 spp. of temperate Eurasia, America, temperate regions on the Northern Hemisphere, temperate South America; open habitats, in weedy places, damp soils ans swamps; only one sp. in South America, P. alpinum L., subarctic & temp. northern Hemisphere to Guatemala, Argentina and Chile, South Georgia.

■ SUBTRIBE CINNINAE ‣ outsiders Aniselytron (2; N India to Japan), Cyathopus (1; Himalayas), Simplicia (2; New Zealand).

102. Agrostopoa Davidse. 3 spp., endemic to Colombia.

103. Cinnastrum E.Fourn. Only one sp., C. poiformis (Kunth) E.Fourn, from Mexico up to W. Bolivia and Venezuela.

■ SUBTRIBE ALOPECURINAE ‣ outsiders Cornucopiae (2; E Mediterranean to Iraq), Limnas (3; Central Asia to NE Siberia).

104. Alopecurus L. c 35 spp., temperate regions on the Northern Hemisphere, temperate South America; 9 spp. in New World, six in South America, scattered from Colombia to Argentina and Uruguay, some amphitropical.

6. SUBFAMILY ARISTIDOIDEAE (3/350–360) ‣ outsiders Sartidia (6; C to S Africa, Madagascar), Stipagrostis (c 55; arid and semi-arid subtropical regions in Mediterranean, Africa and Asia to China).

105. Aristida L. ca. 250 spp. widely distributed in the tropics of the Old World and in the Neotropics; 140 in over New World, 81 in South America, 38 in Brazil, 10 endemics, one of them, from Amazonas states, is a rare plant in Brazil, by Plantas Raras do Brasil’s book.

7. SUBFAMILY PANICOIDEAE (217/3.075–3.105) - four main lineages, with 13 tribes, eight in South America, and Thysanolaeneae (1/1, S and SE Asia, S China), Centotheceae (2/4, tropical regions in the Old World), Cyperochloeae (2/2, W Australia), Chasmanthieae (1/6, S Canada, E U.S.A., N Mexico, Tanzania to Angola) and Lecomtelleae (1/1, Andringitra mountain range in S Madagascar) absents. Panicoideae presents the highest richness of species within Poaceae, comprising ca. 3,270 species widely distributed in the tropics and subtropics, many C4, with three biochemical subtypes, according to Brown and Hattersley, but also including several C3 species. These species belong especially to the tribe Paniceae, with most members in open areas and some groups from forests, and the tribe Andropogoneae, mainly from open areas. Paniceae from forests are often C3, but some species from open and arid areas are also C3 in this group, as in the rocky grasslands (campos rupestres) of SE Brazil, e.g. in Apochloa Zuloaga & Morrone and Renvoizea Zuloaga & Morrone. Likewise, Echinolaena inflexa (Poir.) Chase, a species typical of the savannas of C Brazil (cerrado), is also C3. In the Neotropics there are ca. 120 genera.

In Panicoideae, Parodiophyllochloa, Cyphonanthus, Apochloa, Renvoizea, Stephostachys, Morronea, Ocellochloa, Osvaldoa, Trichanthecium, Rugoloa and Louisiella have been segregated from the polyphyletic Panicum and described as new genera; Thrasya was merged within Paspalum and Ophiochloa was synonymized into Axonopus. Pennisetum was transferred to Cenchrus, and Rupichloa was recircumscribed to accommodate two species previously classified in Urochloa.

The bulk of this subfamily is Paspaleae and Paniceae.

7.1 PANICOIDEAE ▸ TRIBE GYNERIEAE (1/1) - a single species.

106. Gynerium Willd. ex P. Beauv. Tall grasses, distichous leaves, leathery blades with wide midribs, dioecy, and plumose pistillate inflorescences. Only one sp., G. sagittatum (Aubl.) P. Beauv., from Mexico and the Caribbean to NW Argentina, and it is found along river margins or in swampy places.

7.2 PANICOIDEAE ▸ TRIBE ZEUGITEAE (4/18) - outisders Chevalierella (1; Congo), Lophatherum (2; India, Sri Lanka, Himalayas, China (inc. Taiwan), Korean Peninsula, Japan, SE Asia, Malesia to New Guinea and Queensland).

107. Orthoclada P. Beauv. Two spp., O. africana C.E.Hubb. from Tanzania to Zambia, and O. laxa (Rich.) P.Beauv., from S Mexico to tropical South America, Bolivia to French Guiana.

108. Zeugites P. Brown. 12 spp., mainly in Mexico and Central America, three in South America, from Venezuela to Bolivia, one endemic to Ecuador.

7.3 PANICOIDEAE ‣ TRIBE STEYERMARKOCHLOEAE (2/2) - a tribe endemic to South America.

109. Arundoclaytonia Davidse & R.P. Ellis. Perennial 2-3 m tall, erect; vegetative culms usually densely covered for 2-70 cm to a thick- ness of 1. 5-6 cm by aerial roots tightly appressed to the culm and by remnants of leaf sheath bases; internodes numerous, 2-15 mm long, 1-1.5 cm diam., solid, lignified; flowering culms to 1 cm diam., consisting of many, often elongated internodes. Only one sp., A. dissimilis Davidse & R.P. Ellis, restricted from Amazonian campinas in Amazonas and Pará states, south central Amazonian Brazil, in white sand savannas.

110. Steyermarkochloa Davidse & R. P. Ellis. Dimorphic culms and leaves, with solid sheaths and cylindrical and flattened blades, a unique foliar morphology within grasses. Only one sp., S. angustifolia (Spreng.) Judz, grows in the Amazonian llanos of Colombia, Venezuela, and adjacent Amazonas state in N Brazil, in seasonally flooded savanas of Guiana Shield, in elevation ranges 100-200m.

7.4 PANICOIDEAE ▸ TRIBE TRISTACHYIEAE (6/c 70) - outsiders Danthoniopsis (16; Africa, Arabian Peninsula to Pakistan), Gilgiochloa (1; tropical Africa), Trichopteryx (5; tropical and southern Africa, Madagascar).

111. Loudetia Hochst. ex Steud. 25 spp., trop. & S. Africa, Madagascar, S. Arabian Pen., and one sp., L. flammida (Trin.) C.E.Hubb, disjunct from Bolivia to C. Brazil and Paraguay, Guinea and Congo.

112. Loudetiopsis Conert. 11 spp., mainly W. trop. Africa to Chad, with L. chrysothrix (Nees) Conert disjunct Bolivia to Brazil and Paraguay, W. trop. Africa to Chad.

113. Tristachya Nees. 22 spp., tropical and southern Africa, Madagascar, and six spp. in New World, 5 from Mexico to Central America, and T. leiostachya Nees from Brazil and adjacent Paraguay.

7.5 PANICOIDEAE ▸ TRIBE PASPALEAE (38/595–600) - one unplaced genera and three subtribes, all in South America.

■ SUBTRIBE PASPALINAE ‣ outsiders Aakia (1; S Mexico, Central America), Hopia (1; SW U.S.A., NW Mexico), Lecomtella (1; Madagascar).

114. Acostia W.D. Clayton, M. Vorontsova, K.T. Harman & H. Williamson. Perennial, culms 25-40 cm long, ligule a ciliolate membrane. Only one species, A. gracilis Swallen, endemic to Ecuador.

115. Anthaenantiopsis Mez ex Pilg. 4 spp. in Brazil (3, none endemics), Bolivia, Argentina, Paraguay and Uruguay.

116. Axonopus P. Beauv. (inc. Ophiochloa) 101 spp., 99 in tropical and subtropical regions in America (over countries except Canada and Chile), one species in Africa; 90 spp. in South America, 61 in Brazil, 27 endemics, five of them, two from Bahia, two in Goiás, and one from Pará state, are rare plants in Brazil, by Plantas Raras do Brasil’s book.

117. Echinolaena Desv. (off Ichnanthus p.p., Oedochloa p.p.) Annual or perennial; rhizomes present; culms erect, decumbent or scandent. Two widely spp. from Belize, Bolivia, Brazil (both, none endemics), Colombia, Costa Rica, French Guiana, Guatemala, Guyana, Honduras, Mexico, Nicaragua, Suriname, and Venezuela, inhabit open field areas.

118. Gerritea Zuloaga, Morrone & Killeen. Only one sp., G. pseudopetiolata Zuloaga, Morrone & T. Killeen, endemic to Bolivia.

119. Hildaea C. Silva & R.P. Oliveira. Annual or perennial; rhizomes absent; culms stoloniferous, decumbent or scandent, sometimes erect via stout and rigid adventitious roots which emerge from the lower nodes. 6 spp. from tropical America (all in South America) except H. pallens (Sw.) C. Silva & R.P. Oliveira, also in Africa, Asia to NE. Australia; 5 spp. in Brazil, H. parvispiculata C. Silva & R.P. Oliveira endemic to northern and has the smallest spikelets of genus.

120. Ichnanthus P. Beauv. (inc. Echinolaena p.p.) Annual or perennial; rhizomes present or absent; culms erect, decumbent or scandent. 24 spp., distributed in the tropics of the New World where they usually inhabit forests, or less commonly open field areas, all in South America and Brazil, 17 endemics.

121. Ocellochloa Zuloaga & Morrone. Perennial, delicate to robust, stoloniferous or decumbent, rooting and branching at the lower nodes to erect, leaning or not on vegetation; culms hollow, rarely solid, simple or branching. 12 spp. from Mexico to Argentina, 10 in South America, 8 in Brazil, 4 endemics.

122. Oedochloa C. Silva & R.P. Oliveira. (inc. Echinolaena p.p., Ichnanthus p.p.) Annual or perennial; rhizomes present or absent; culms erect, decumbent or scandent. 9 spp., 6 in South America (all in Brazil except one endemic to Ecuador, three endemics), one in Caribbean and two in Mexico and Central America.

123. Osvaldoa J. R. Grande. Caespitose, shortly rhizomatous perennials; culms erect, many-noded; blades linear-lanceolate, shortly pilose toward the base on adaxial surface; inflorescence oblong and open panicle. Only one sp., O. valida (Mez) J. R. Grande, S Brazil, NE Argentina and NW Uruguay; border of rivers and streams.

124. Paspalum L. c 400 spp., tropical and subtropical regions on both hemispheres, with their largest diversity in tropical America; 373 spp. in New World, 298 in South America, 217 in Brazil, 80 endemics, nine of them, all from central savannic states, are rare plants in Brazil, by Plantas Raras do Brasil’s book.

125. Renvoizea Zuloaga & Morrone. Caespitose perennials, rhizomatous, forming robust tussocks; basal leaf-sheaths tough and persistent; ligule membranous-ciliate, ciliate or absent; spike dense, narrowly oblong; spikelets elipsoid, biconvex, glabrous or pilose. 10 spp., endemic to Brazil, growing in savannas of C Brazil (cerrado), atlantic sandy coastal shrublands (restingas), and high-altitude grasslands (campos de altitude) in the states of Bahia (6 endemic species in this state), Minas Gerais, Espírito Santo and Rio de Janeiro (two endemic species).

126. Spheneria Kuhlm. Only one sp., S. kegelii (C.Muell.) Pilg., restricted from Guyana, Suriname and N Brazil.

127. Streptostachys Desv. Two spp., one from N Brazil to Bolivia, Venezuela, Guianas and Trinidade & Tobago, the latter endemic to Bahia state, where it grows in sand dunes of the northern Espinhaço Range.

■ SUBTRIBE OTACHYRIINAE ‣ all genera in South America.

128. Anthenantia P. Beauv. 4 spp. in New World, only one in South America, A. lanata (Kunth) Benth, Mexico to Uruguay, east to French Guiana, Caribbean.

129. Hymenachne P. Beauv. 10 spp., C. trop. Africa, Assam to S. China and Indo-China, trop. & Subtrop. America; 5 spp. in New World, 4 in South America, two over widely distributed in Neotropics, two only Brazil and adjacent Argentina, Paraguay and Uruguay.

130. Otachyrium Nees. 8 spp. over South America except Chile (one up to Trinidad & Tobago), 7 in Brazil, 4 endemics - two of them, in Bahia and Goiás states, are rare plants in Brazil, by Plantas Raras do Brasil’s book.

131. Plagiantha Renvoize. Inflorescence lax, without spikelets unilaterally arranged, glabrous, the lower palea expanded at maturity with two prominent wings, lower lemma 2-4-nerved, and upper anthecium cartilaginous, covered with compound papillae all over its surface. Only one sp., P. tenella Renvoize, in damp ground in shady places in woodland or savannas of C Brazil (cerrado); 640-1,000 m, known only in Serra do Curral Feio, Bahia state in Brazil.

132. Rugoloa Zuloaga. 5 spp., two in Asia to Australia and 3 all over widely distributed in New World, all in Brazil.

133. Steinchisma Raf. 8 spp., from U.S.A. to Ecuador, Brazil, Guianas, Caribbean; 6 in South America, 5 in Brazil, none endemics.

■ SUBTRIBE ARTHROPOGONINAE ‣ outsiders Phanopyrum (1; SE U.S.A.), Triscenia (1; Cuba).

134. Altoparadisium Filg., Davidse, Zuloaga & Morrone. Caespitose perennials; culms erect; internodes hollow; leaves primarily cauline; sheaths round on the back, the margins free; auricles absent; ligule a minute fringe of hairs; collar undifferentiated; blades linear-lanceolate; inflorescence a contracted, terminal panicle. Two spp., A. scabrum (Pilg. & Kuhlm.) Filg., Davidse, Zuloaga & Morrone from C Brazil and Bolivia, and A. chapadense Filg., Davidse, Zuloaga & Morrone endemic to Brazil (Goias state), a rare plant in Brazil, by Plantas Raras do Brasil’s book.

135. Apochloa Zuloaga & Morrone. Caespitose perennials, forming robust tussocks, rarely decumbent and rooting at the lower nodes, culms simple, blades basal; blades lanceolate to linear-lanceolate, frequently inrolled, pungent, densely pilose to glabrous; inflorescence an open, lax, and terminal panicle; spikelets ovoid to ellipsoid, glabrous, occasionally pilose. 15 spp., A. chnoodes (Trin.) Zuloaga & Morrone in N Brazil, Venezuela and Guyana, 4 only in Venezuela and 10 endemics to Brazil, growing in open, rocky soils of the Guiana Shield and in savannas of C Brazil (cerrado) and rocky grasslands (campos rupestres), between 1,000-2,500 m elevation.

136. Arthropogon Nees. 4 spp., two endemics to SE Brazil, another high disjunct in Mexico, Brazil and Bolivia, and one endemic to Colombia.

137. Canastra Morrone et al. Plants perennial, caespitose; culms 50-60 cm tall, simple or branched at the basal nodes, erect; internodesc ompresseds, triate, glabrousn; odes glabrous. Two spp. endemic to Brazil, one of them known only from Minas Gerais state and a rare plants in Brazil, by Plantas Raras do Brasil’s book, typically among rocks, or very close to the margins of puddles, streams, or small rivers; they are found in shallow and extremelly sandy soils; plants robust, tussock-like, with compiscuous upright purples inflorescence.

138. Coleataenia Griseb. Plants perennial, cespitose, short to long rhizomatous, the rhizomes short or long; culms simple or branching at the middle and upper nodes, internodes terete or compressed; inflorescence exerted, panicles lax to contracted, few or multiflowered. 7 spp., U.S.A. to Brazil and Argentina, 4 in South America, all of them in Brazil, none endemics.

139. Cyphonanthus Zuloaga & Morrone. Short-rhizomatous perennial with simple culms decumbent, rooting at the lower nodes; ligules ciliate; blades lanceolate; inflorescences lax, spikelets ovoid, gibbous, glabrous. Only one sp., C. discrepans (Döll) Zuloaga & Morrone, Cuba and Belize to C Brazil, from sea level to 1,200 m, absent from Ecuador to Argentina, and rarely in Costa Rica and Colombia.

140. Homolepis Chase. 5 spp., Mexico to Uruguay, French Guiana and Caribbean, all in Brazil, H. longispicula (Döll) Chase endemic to dry areas in sandy soils in Brazil.

141. Keratochlaena Morrone & Zuloaga. Caespitose, shortly rhizomatous perennials; blades lanceolate, rigid, sparsely pubescent to glabrous, pungent; inflorescences racemose to subpaniculate; spikelets narrowly ellipsoid, solitary. Only one sp., K. rigidifolia (Filg., Morrone & Zuloaga) Morrone & Zuloaga, only known from the type locality (a rare plants in Brazil, by Plantas Raras do Brasil’s book), in Loreto municipality, Maranhão state, Brazil, in open savannas.

142. Mesosetum Steud. 27 spp., tropical America, 23 in South America, with their highest diversity in Brazil, with 22, 13 endemics, two of them, from Mato Grosso and Goiás states, are rare plants in Brazil, by Plantas Raras do Brasil’s book.

143. Oncorachis Morrone e Zuloaga. Plants perennial, shortly rhizomatous; inflorescences terminal, lax; spikelets ellipsoid, greenish and tinged with purple, pilose or glabrous, the rachilla manifest and thickened between the lower glume and the upper anthecium. Two spp. from savannas of C Brazil (cerrado) at Bahia to Minas Gerais and Mato Grosso states, and Sierra de Amambay in Paraguay, reaching to 1,200 m in elevation.

144. Oplismenopsis Parodi. Only one sp., O. najada (Hack. & Arechav.) Parodi, in Rio Grande do Sul state in S Brazil to NE. Argentina, and Uruguay.

145. Stephostachys Zuloaga & Morrone. Plants perennial, rhizomatous, rhizomes short; culms erect, 1–3 m tall, some culms decumbent and rooting at the lower nodes; internodes 7–20 cm long, terete, solid toward the basal portion, glabrous, pale. Only one sp., S. mertensii (Roth) Zuloaga & Morrone, widely distributed from Mexico and Cuba to Paraguay, Bolivia, and northern Argentina; it grows in or on the edges of forest, usually in inundated areas.

146. Tatianyx Zuloaga & Soderstr. Only one sp., T. anthracites (Trin.) Zuloaga & Soders, from Bahia and Minas Gerais states.

7.6 PANICOIDEAE ▸ TRIBE ARUNDINELEAE (4/80) - outsiders Garnotia (c 30; S and E Asia to islands in the Pacific), Chandrasekharania (1; Kerala in India), Jansenella (1; India, Sri Lanka); the positions of Chandrasekharania and Jansenella are uncertain.

147. Arundinella Raddi. c 60 spp., tropical and subtropical regions on both hemispheres; three spp. in New World, all widely distributed and in Brazil.

7.7. PANICOIDEAE ▸ TRIBE ANDROPOGONEAE (82/1.110–1.130) - nine subtribes and six unplaced genera (all Old World); subtribes Arthraxoninae (1/27; tropical and subtropical Africa, Madagascar, islands in the Indan Ocean, India and Himalayas to S China and SE Asia, Malesia to New Guinea and N Australia), Chionachninae (1/12; India to Burma, S China and SE Asia, Malesia to New Guinea, Solomon Islands and N Australia; C. cookei (Trilobachne cookei): the Caribbean), Coicinae (1/4; India to S China and SE Asia, N Queensland) and Germainiinae (4/31; Madagascar, Arabian Peninsula, India to China and Japan, SE Asia, Malesia to New Guinea and Queensland) do not occur in South Amarica

■ SUBTRIBE TRIPSACINAE ‣ outsiders Oxyrhachis (1; tropical E and S Africa, Madagascar), Urelytrum (7; tropical and S Africa, Madagascar), Vossia (1; tropical Africa to Namibia and N Botswana, E India to Burma), Zea (6; Mexico, Central America).

148. Elionurus Humb. & Bonpl. ex Willd. c 15 spp., tropical and subtropical Africa and eastwards to Sind in NW India, Australia, tropical and subtropical America; 7 spp. in New World, 6 in South America, three in Brazil, none endemics.

149. Rhytachne Desv. ex Ham. 12 spp., tropical and S Africa, Madagascar, tropical South America, three in New World, R. gonzalezii Davidse, (N. South America to N Brazil), R. guianensis (Hitchc.) Clayton, (S. Mexico, Colombia to Suriname and N Brazil) and R. subgibbosa (Winkler ex Hack.) Clayton, (SE. Mexico, S Brazil to NE. Argentina, N Zambia).

150. Tripsacum L. 16 spp., S U.S.A., Mexico, Central America, South America to Paraguay, 5 in South America, two in Brazil, none endemics.

■ SUBTRIBE ROTTBOELLIINAE ‣ outsiders Chasmopodium (3; tropical W and C Africa), Eremochloa (14; India, Sri Lanka, SE Asia, Malesia to New Guinea and tropical Australia), Hemarthria (12; tropical and subtropical regions in the Old World), Ophiuros (4; the Horn of Africa, India and E Himalayas to Burma, S China and SE Asia, the Ryukyu Islands, Malesia to New Guinea and Australia), Hackelochloa (2; sub-Saharan Africa, Arabian Peninsula, India, E Himalayas to Japan and SE Asia, Malesia to New Guinea, Micronesia), Phacelurus (10; tropical and subtropical Africa, SW and E Asia, tropical Asia to Indochina and Japan, one species, P. digitatus, on the Balkan Peninsula to Syria), Rottboellia (6; tropical Africa, Madagascar, tropical and subtropical Asia and Australia, Melanesia), Glyphochloa (12; central and SW India), Heteropholis (3; Central and tropical E Africa; Madagascar; Sri Lanka), Lasiurus (1; Morocco and Mali to the Middle East and NE India), Loxodera (5; tropical Africa), Thaumastochloa (9; New Guinea, Australia).

151. Mnesithea Kunth. 33 spp., in trop. & subtrop. to C & E U.S.A., 12 in New World, six in South America, 4 in Brazil, none endemic; all South American species are widely.

■ SUBTRIBE ISCHAEMINAE ‣ outsiders Andropterum (1; tropical Africa), Dimeria (c 60; Madagascar, India and Sri Lanka to China, Korean Peninsula, Japan and SE Asia, Malesia to New Guinea and tropical Australia, Micronesia), Kerriochloa (1; SE Asia), Pogonachne (1; Mumbay area in India), Triplopogon (1; W India).

152. Ischaemum L. c 70 spp., tropical and subtropical regions on both hemispheres, especially Asia, 4 spp. in New World, all in South America: one fully over Neotropics, one from N South America, another endemic to S Venezuela, and one disjunct in Ogasawara-shoto to Philippines, SE & S Brazil to NE. Argentina.

■ SUBTRIBE SACCHARINAE ‣ outsiders Asthenochloa (1; C Malesia), Euclasta (2; tropical Africa, Madagascar, the Comoros, Oman India, Burma), Pseudodichanthium (1; W India), Cleistachne (1; tropical E and SE Africa, Oman, India), Sorghum (28; Old World, India, Sri Lanka, Burma, SE Asia Australia, one species, S. trichocladum, in Mexico and Central America), Eulalia (34; Africa, Madagascar, Asia and Australia), Homozeugos (6; tropical Africa), Miscanthus (16; tropical and subtropical regions in the Old World, S Africa, E Asia), Polytrias (1; SE Asia), Pseudosorghum (2; Himalayas, Burma, Yunnan, SE Asia, Java, Philippines), Tripidium (3; SE Europe, Mediterranean, SW and S Asia), Veldkampia (1; SE Asia).

153. Agenium Nees. 4 spp., all widely from Bolivia to Brazil (all spp., none endemics) and NE. Argentina, Paraguay and Uruguay.

154. Eriochrysis P. Beauv. Grass with inflorescences with golden-brown to light-brown trichomes, and heterogamous spikelets: sessile spikelets with a bisexual flower and pedicelled spikelets with a pistillate flower. 7 spp., tropical and southern Africa, India, tropical America, six in South America, all in Brazil (one endemic), mainly C & S South America.

155. Imperata Cirillo. 11 spp., tropical and subtropical regions on both hemispheres, 6 spp. in South America: two widely in Neotropics, two from Ecuador to SE Brazil and Argentina, two only Chile and Argentina; none species is endemic to Brazil.

156. Saccharum L. (inc. Erianthus). 35–40 spp., in tropical and subtropical regions on both hemispheres. 12 spp. in New World, 7 in South America, 5 in Brazil, none endemic.

157. Sorghastrum Nash. 18 spp., tropical and subtropical regions in America, 13 spp. in South America, 11 in Brazil, 4 endemics; one spp. is disjunct tropical America and Africa.

158. Trachypogon Nees. 4 spp., one tropical and southern Africa, Madagascar and three in tropical America, all in Brazil (none endemics), one of then in both Africa and New World.

■ SUBTRIBE ANDROPOGONINAE ‣ outsiders Bhidea (3; India), Diheteropogon (4; tropical and S Africa), Anadelphia (21; W to SE tropical Africa), Capillipedium (18; E Africa, tropical Asia to Australia and New Caledonia), Clausospicula (1; Northern Territory), Cymbopogon (55–60; tropical and subtropical regions in Africa, Madagascar, Asia and Australia), Dichantium (19–23; tropical and subtropical regions in the Old World), Eremopogon (1; tropical Africa, the Middle East to India and Sri Lanka), Exotheca (1; tropical E Africa, Vietnam), Hyperthelia (7; tropical and S Africa), Iseilema (c 30; India, Sri Lanka, SE Asia, Malesia to New Guinea and tropical Australia), Monocymbium (3; tropical and S Africa), Parahyparrhenia (6; tropical W and C Africa, India, Thailand), Pseudanthistiria (4; India, Thailand, southern China), Spathia (1; tropical Australia), Spodiopogon (16; Türkiye, SE, Central, E and tropical Asia to Russian Far East, Japan, Taiwan (China) and the Malay Peninsula), Themeda (26; tropical and subtropical regions in Africa, Asia and Australia); Lakshmia (1; Sri Lanka, W Ghats).

159. Andropogon L. 120 spp., over cosmoopolitan, all countries of New World except Chile; 66 spp. in New World, 49 in South America, 29 in Brazil, 11 endemics, one of them, from Minas Gerais state, is a are rare plant in Brazil, by Plantas Raras do Brasil’s book.

160. Bothriochloa Kuntze. 21 spp. in tropical and subtropical regions in America, 18 in South America, mainly amphitropical; 12 in Brazil, two endemics, one of them, from Mato Groso do Sul state, is a rare plants in Brazil, by Plantas Raras do Brasil’s book.

161. Elymandra Stapf. 6 spp. restricted from tropical and S Africa except E. lithophila (Trin.) Clayton, from SE Brazil, RD Congo and Zimbabwe, probably due to long distance dispersal.

162. Heteropogon Pers. 5 spp., three only Old World (mainly S Asia), and H. contortus (L.) P.Beauv. ex Roem. & Schult. and H. melanocarpus (Elliott) Benth., both pantropical.

163. Hyparrhenia Andersson ex E. Fourn. c 65 spp., Macaronesia, Mediterranean, Africa, Madagascar, SW and S Asia eastwards to Pakistan and India, tropical America, with their highest diversity in Africa, only one sp. in New World, H. bracteata (Humb. & Bonpl. ex Willd.) Stapf., occuring in Old World and from Mexico to Bolivia and Brazil.

164. Schizachyrium Nees. 60 spp., trop. & S. Africa, W. Indian Ocean, trop. & subtrop. Asia to W. Pacific, America, 36 spp. in New World (all countries), 22 in South America, 17 in Brazil, only two endemics.

7.8 PANICOIDEAE ▸ TRIBE PANICEAE (74/1.190–1.210) - Paniceae from forests are often C3, but some species from open and arid areas are also C3 in this group, as in the rocky grasslands (campos rupestres) of SE Brazil, e.g. in Apochloa and Renvoizea. Likewise, Echinolaena inflexa (Poir.) Chase, a species typical of the savannas of C Brazil (cerrado), is also C3. Six unplaced genera (five absent in South America) and eight subtribes, only Neurachninae (4/14, arid regions in Australia) absent.

■ SUBTRIBE UNPLACED TAXA

165. Cnidochloa Zuloaga. (off Panicum) Short-rhizomatous perennial with culms decumbent, rooting at lower nodes; inflorescence a lax and diffuse panicle, spikelets long, ellipsoid, chasmogamous. Only one sp., C. longipedicellata (Swallen) Zuloaga, restricted to Brazil, where it grows along margins and in clearings of forests in the states of Rio de Janeiro, Paraná, and Santa Catarina, between 700 and 1,400 m elevation.

■ SUBTRIBE ANTHEPHORINAE ‣ outsiders Chlorocalymma (1; Tanzania), Chaetopoa (2; Tanzania), Taeniorhachis (1; Somalia), Tarigidia (1; Namibia, NW, Free State), Thyridachne (1; tropical Africa), Trachys (1; coastal areas in S India, Sri Lanka, Bangladesh and Burma).

166. Anthephora Schreb. 11 spp., 10 in tropical and southern Africa, Arabian Peninsula, and one species, A. hermaphrodita (L.) Kuntze, also in tropical America, widely distributed.

167. Digitaria Haller. Spikelets with a reduced lower glume, upper anthecium cartilaginous, and upper lemma margin thin and not inrolled. 250 spp. worldwide, 84 spp. in New World, 54 in South America, 26 in Brazil, 12 endemics, three of them, from Minas Gerais, Mato Grosso do Sul and Santa Catarina states, are rare plants in Brazil, by Plantas Raras do Brasil’s book.

■ SUBTRIBE DICHANTHELIINAE ‣ outsider Adenochloa (14; tropical and subtropical Africa, Madagascar).

168. Dichanthelium (Hitchc. & Chase) Gould. 68 spp. from Canada and the U.S.A. to center of Argentina and Chile, from sea level to approximately 3,000 m, frequent in forest edges, in wet habitats, some species in open places, on moist or dry sandy soils; 33 spp. in South America, 26 in Brazil, 15 endemics, three of them, from Bahia and Minas Gerais states, are rare plants in Brazil, by Plantas Raras do Brasil’s book.

■ SUBTRIBE BOIVINELLINAE ‣ outsiders Setiacis (1; Hainan), Alloteropsis (5; tropical and subtropical regions in the Old World), Amphicarpum (2; SE U.S.A.), Cyphochlaena (2; Madagascar, the Comoros), Cyrtococcum (14; Africa, Asia and Australia), Entolasia (6; Africa, tropical Asia to E Australia, New Caledonia), Mayariochloa (1; Cuba), Microcalamus (1; W Africa), Ottochloa (3; India, S China, SE Asia, Malesia to N Australia), Poecilostachys (c 20; tropical Africa, Madagascar).

169. Acroceras Stapf. 24 spp., 4 in New World, one widely in Neotropics and thee only in South America, all four in Brazil, one endemic.

170. Echinochloa P. Beauv. 45 spp., warm-temperate to tropical regions on both hemispheres, 11 spp. in New World, 7 in South America, 4 in Brazil, one endemic.

171. Lasiacis (Griseb.) Hitchc. Scandent habit. 16 spp., tropical and subtropical regions in America, 13 in South America, all widely distributed, 5 in Brazil, none endemics.

172. Morronea Zuloaga and Scataglini. Perennials with decumbent culms; ligules membranousciliate; inflorescence a lax and spreading panicle spikelets ellipsoid, glabrous; caryopsis with hilum punctiform. 6 spp. inhabiting humid and shaded places from Mexico to Argentina, 4 only in Mexico and Central America and two widely in tropical America, only one in Brazil.

173. Oplismenus P. Beauv. 7 spp., tropical and subtropical regions on both hemispheres, 4 in New World, three in South America and two in Brazil, all very highly widely distributed.

174. Parodiophyllochloa Zuloaga & Morrone. Perennials with decumbent culms; ligules membranous; panicles lax and spreading, generally with chasmogamus or cleistogamus spikelets, the chasmogamus inflorescences usually terminal on the culm. 6 spp., Mexico to Uruguay (all in South America and in Brazil), east up to French Guiana, four widely distributed, one in S Brazil and Argentina, and one endemic to Brazil, in dense colonies at forest edges.

175. Pseudechinolaena Stapf. 6 spp., mainly Madagascar, with P. polystachya (Kunth) Stapf. pantropical, ver highly widely distributed.

■ SACCIOLEPIS CLADE ‣ both genera in South America.

176. Sacciolepis Nash. c 30 spp., tropical and subtropical regions on both hemispheres, with their largest diversity in tropical Africa; 5 spp. in New World, all widely distributed and in Brazil.

177. Trichanthecium Zuloaga, Morrone & Scataglini. 38 spp., Africa and New World, 25 in New World, all in South America, 18 in Brazil, six endemics.

■ SUBTRIBE MELINIDINAE ‣ outsiders Thuarea (2; Madagascar, tropical and E Asia to tropical Australia, Melanesia, Micronesia and Hawai), Moorochloa (3; tropical and subtropical Old World), Tricholaena (4; Canary Islands, Mediterranean, Africa, Madagascar), Leucophrys (1; S Namibia, N Cape), Melinis (22; Africa, Madagascar), Eccoptocarpha (1; S tropical Africa), Yvesia (1; Madagascar).

178. Chaetium Nees. Three spp., one in Mexico to Central America, one in Cuba, and C. festucoides Nees, from Colombia to Venezuela and NE Brazil.

179. Eriochloa Kunth. 33 spp., tropical and subtropical regions on both hemispheres, 20 spp. in New World, 14 in South America, six in Brazil, none endemics.

180. Rupichloa Salariato & Morrone. (inc. Urochloa p.p. in some sources) Rhizomatous or caespitose perenials, rooting at the lower nodes; blades lanceolate to linear lanceolate. Two spp., endemics to Bahia and Minas Gerais state in SE Brazil, both rares by Plantas Raras do Brasil’s book, up to limestone soils 1,000-1,500 altitudinal range.

181. Urochloa P. Beauv. 86 spp., tropical and subtropical regions on both hemispheres; 19 spp. in New World, 7 only from North America to Mexico, 5 from Colombia and Ecuador to NW Argentina, six are scattered from North America or Caribbean up to Brazil, sometimes Argentina: U. adspersa (Trin.) R.D.Webster, U. fusca (Sw.) B.F.Hansen & Wunderlin, U. mollis (Sw.) Morrone & Zuloaga, U. plantaginea (Link) R.D.Webster, U. platyphylla (Munro ex C.Wright) R.D.Webster and U. polystachya (Kunth) Mabb.; and one is endemic to Brazil, U. megastachya (Nees ex Trin.) Morrone & Zuloaga known only in Serra do Cipó in Minas Gerais state.

■ SUBTRIBE PANICINAE ‣ both genera in South America.

182. Louisiella C. E. Hubb. & J. Leonard. Aquatic perennials; culms prostrate, decumbent and rooting at the lower nodes, then ascendent, succulent; internodes spongy, hollow, greenish to pale, glabrous. Three spp., L. elephantipes (Nees ex Trin.) Zuloaga from Mexico to Argentina, a common aquatic grass growing in ponds and shallow water; L. fluitans C.E.Hubb. & J.Léonard from Cameroon, Central African Republic, Southern Sudan, and the Democratic Republic of Congo; and L. paludosa (Roxb.) S. N. Landge, from Iran, Pakistan, India, Tropical Asia, Indo-China, North and East Australia.

183. Panicum L. (exc. Cnidochloa) Caespitose, annual or perennial plants; ligules membranous-ciliate or ciliate; blades lanceolate to linear lanceolate; inflorescences lax and open, with spikelets arranged in long pedicels; spikelets with lower glume present, 3–9- nerved, upper glume and lower lemma subequal, 7–13-nerved, lower palea and lower flower present or absent. 450 spp. worldwide, 78 in New World, 53 in South America, 41 in Brazil, 11 endemics, 8 of them, from several states, are rare plants in Brazil, by Plantas Raras do Brasil’s book.

■ SUBTRIBE CENCHRINAE ‣ outsiders Acritochaete (1; tropical Africa), Alexfloydia (1; New South Wales), Chamaeraphis (1; N Australia), Dissochondrus (1; Hawai), Holcolemma (3; tropical E Africa, India, Australia), Hygrochloa (2; N Australia), Paractaenum (2; Australia), Pseudochaetochloa (1; N Australia), Pseudoraphis (7; India to China, Japan and Australia), Spinifex (4; India to E Asia and islands in the Pacific), Stereochlaena (4; tropical E to South Africa), Streptolophus (1; Angola), Uranthoecium (1; tropical Australia), Whiteochloa (6; Aru Islands, tropical Australia), Xerochloa (3; Australia), Zygochloa (1; arid regions of central Australia).

184. Cenchrus L. 108 spp., SE. Europe, Africa to W. Indo-China, N & E Australia to Pacific, and New World (36); 25 in South America, 8 in Brazil, none endemics.

185. Ixophorus Schltdl. Only one sp., I. unisetus (J. Presl) Schltdl., Mexico to Colombia, disjunct in Caribbean.

186. Paratheria Griseb. Two spp., tropical W and C Africa, Madagascar, P. prostrata Griseb. from W. trop. Africa to Ethiopia and Namibia, Madagascar, disjunct in Costa Rica, Caribbean, N South America, Bolivia and Brazil.

187. Setaria P. Beauv. 180 spp. worldwide, mostly in the tropics and subtropics of Africa, Asia, and the New World (60), 43 spp. in South America, 24 in Brazil (only a single endemic), mainly of spp. restricted of a only to few countries.

188. Setariopsis Scribn. Two spp., one and endemic to Mexico, and S. auriculata (E.Fourn.) Scribn from Arizona to Nicaragua, disjunct in Colombia and Venezuela.

189. Stenotaphrum Trin. 7 spp., tropical and subtropical regions on both hemispheres, only one in New World, S. secundatum (Walter) Kuntz, from SE U.S.A. to E & S South America, tropical Africa to Chad.

190. Zuloagaea Bess. Plants perennial, caespitose or single-stalked, erect; culms 20-200 cm tall, 1–8 mm thick, glabrous, lowest internode often thickened into a hard, corm-like base; blades linear. Only one sp., Z. bulbosa (Kunth) Bess, from Arizona and New Mexico to N South America (between Colombia and Ecuador); it occurs at elevations from 900-3,000 meters, primarily on rocky or sandy soil in drainage areas or in deep leaf mulch in forests of oak and/or pine, under dry to very moist conditions.

8. SUBFAMILY ARUNDINOIDEAE (11-13/35-37) ‣ three tribes, Arundineae (4/18, Tropical and subtropical regions of the Old World) and Crinipedeae (4/8, tropical and subtropical regions of the Old World, with their largest diversity in S and E Africa) do not occur in South America; among Molinieae, outsiders are Hakonechloa (1; Japan), Leptagrostis (1; Ethiopia), Molinia (2; Europe, temperate Asia), Piptophyllum (1; Angola).

191. Phragmites Adans. 4 spp. cosmopolitan, one in neotropics; the only one representative in South America, P. australis (Nees) Döll, considered cosmopolitan, of swamps and riverbanks, occur in all coutries of New World except Brazil, Paraguay and some Caribbean Is.

9. SUBFAMILY MICRAIROIDEAE (9/c 185) ‣ three tribes, Micraireae (1/16; tropical Australia) and Eriachneae (1/c 50; Sri Lanka and southern China to southern Malesia and Australia) do not occur in South America; among Isachneae, outsiders are Coelachne (11; tropical Africa, Madagascar, tropical Asia from India to New Guinea), Heteranthoecia (1; tropical Africa), Sphaerocaryum (1; India to China (inc. Taiwan), SE Asia and W Malesia), Limnopoa (1; S India), Hubbardia (2; W India), Zenkeria (5; India, Sri Lanka).

192. Isachne R. Br. Plants annual or perennial, trailing culms which can reach a length of several meters; ligule setose; inflorescence a panicle, contracted to lax; spikelets with two florets, disarticulating above the glumes and deciduous together, with the glumes later also deciduous. 95 spp., mainly in tropical Asia; 14 spp. in New World, 7 in South America, 4 in Brazil, three endemics, one of them, from Bahia state, is a rare plant in Brazil, by Plantas Raras do Brasil’s book; mainly in marshy, swampy and wet places, along rivers, or associated with gallery forests.

10. SUBFAMILY DANTHONIOIDEAE (19/c 285) ‣ outsiders Merxmuellera (7; tropical and S Africa, Madagascar), Capeochloa (3; S Africa), Geochloa (3; tropical and S Africa), Pentameris (c 80; Africa, Madagascar, S and E Asia), Chionochloa (26; New Zealand, Auckland and Campbell Islands, Lord Howe, New Guinea), Chaetobromus (1; S Namibia, Northern and W Cape), Pseudopentameris (3; mountains in W Cape), Austroderia (5; New Zealand, Chatham Islands), Notochloe (1; New South Wales), Plinthanthesis (3; SE Australia), Chimaerochloa (1; tropical Asia to New Guinea), Tenaxia (8; Africa, temperate to tropical Asia), Schismus (5; Mediterranean, Africa, SW Asia to NW India and China, especially S Africa), Tribolium (14; S Namibia, N, W and E Cape), Phaenanthoecium (1; mountains in NE tropical Africa), Danthonidium (1; India).

193. Cortaderia Stapf. Gynodioecious, dioecious, hermaphrodite or apomictic perennials, ranging from rounded vegetable hedgehogs less than 0.5 m tall to erect 4 m tall tussocks; innovations intravaginal; spreading stolons rare. 21 spp. from Tierra del Fuego in the south to Venezuela in the north (two up to Central America), and from the Atlantic coastal mountains near Rio de Janeiro to the Ecuadorian Andes (4 in Brazil, two endemics, one of them, from Santa Catarina state, is a rare plants in Brazil, by Plantas Raras do Brasil’s book), and from sea level at the southern extreme to over 4,500 m at the equator. C. selloana (Schult. & Schult. f.) Asch. & Graebn., which is globally cultivated as a garden ornamental, but which is also an aggressive invader in many warm-temperate regions.

194. Danthonia DC. Perennial ceaspitose herb up to 1m tall, unbranched above. 27 spp. worldwide, Macaronesia, Europe to Caucasus, N. Africa, Russian Far East, temp. & subtrop. America; 21 spp. in New World, 12 in South America, 4 in Brazil, none endemic, shared with Argentina, Uruguay and Bolivia, two also endemic to Caribbean.

D. secundiflora J. Presl is found all the way to the high altitude grasslands (campos de altitude) of Bahia (Morro do Chapéu), disjunct in southern country, Uruguay, and Colombia; this is the northernmost limit known for microthermic grasses in Brazil.

195. Rytidosperma Steud. Perennial ceaspitose herb up to 1m tall, unbranched above. 68 spp. from SE Asia along mainly Australasia, New Zealand, Hawaii, and perhaps also South America, with six spp. from Chile and Argentina.

11. SUBFAMILY CHLORIDOIDEAE (125–126/1.405–1.420) - five tribes, only Centropodieae (2/6, S Africa and one species in North Africa eastwards to India) absent in South America.

11.1 CHLORIDOIDEAE ▸ TRIBE TRIRAPHIDAE (4/14) - outsiders Habrochloa (1; Central Africa), Neyraudia (4; tropical and subtropical regions in Africa and Asia), Nematopoa (1; Zambia, Zimbabwe).

196. Triraphis R. Br. 8 spp., six in Africa and Arabian peninsula one in Australia, T. devia Filg. & Zuloaga in Goias state in center Brazil - with only three endangered populations in mountains Veadeiros National Park, and a rare plant in Brazil, by Plantas Raras do Brasil’s book, always in savanna habitats, often in sandy or stony soils; probably only Poaceae genus with their distribution disjunct type.

11.2 CHLORIDOIDEAE ▸ TRIBE ERAGROSTINAE (13/410–415) - three subtribes, all in South America.

■ SUBTRIBE COTTEINAE ‣ outsiders Kaokochloa (1; NW Namibia), Schmidtia (2; Africa, Cape Verde Islands, Pakistan)

197. Cottea Kunth. Only one sp., C. pappophorides Kunth; southern U.S.A. to central Mexico and from Ecuador to Bolivia and NW Argentina.

198. Enneapogon Desv. ex P. Beauv. 26 spp., dry warmer regions on both hemispheres; one sp., E. desvauxii P.Beauv., amphitropical disjunct of WC U.S.A., Peru to N Argentina and Chile.

■ SUBTRIBE ERAGROSTIDINAE ‣ outsiders Cladoraphis (2; Namibia, N and W Cape), Richardsiella (1; Zambia).

199. Eragrostis Wolf. c 350; tropical to warm-temperate regions on both hemispheres; non-monophyletic. 95 spp. in New World, 67 in South America, over widely distributed, 36 in Brazil, 6 endemics.

200. Steirachne Ekman. Two spp., S. barbata (Trin.) Renvoize and S. diandra Ekman, from Venezuela, Guyana and N Brazil (both spp., none endemics).

■ SUBTRIBE UNIOLINAE ‣ outsiders Entoplocamia (1; Namibia), Fingerhuthia (2; S tropical and S Africa, Arabian Peninsula, Afghanistan to India), Tetrachaete (1; Ethiopia, Tanzania, Arabian Peninsula), Tetrachne (1; E Cape, Free State, Lesotho, Pakistan).

201. Uniola L. Strongly laterally compressed spikelets with 6–20 florets and coriaceous, sharply keeled lemmas. 5 spp., two in Caribbean and Mexico to North America, Peru and Ecuador one endemic each, and U. pittieri Hack. from Mexico to Peru and Venezuela.

11.3 CHLORIDOIDEAE ▸ TRIBE ZOYSIEAE (4–5/185–190) - two subtribes, Zoysiinae (2/10; Sudan and Somalia to Sind, Mauritius to Polynesia) absent in South America; among South American Sporobolinae, outsiders is Psilolemma (1; E Africa).

202. Sporobolus R. Br. c 160 spp.; warm-temperate to tropical regions on both hemispheres; 80 spp. in New World, 40 in South America, 25 in Brazil, 8 endemics - three of them, from Paraná, Goiás and Minas Gerais states, are rare plants in Brazil, by Plantas Raras do Brasil’s book.; S. atrovirens (Kunth) Kunth, endemic to Mexico, is one of three desiccation tolerant Poaceae in New World.

11.4 CHLORIDOIDEAE ▸ TRIBE CYNODONTHEAE (95/785–800) - 25 subtribes, 14 in South America, Triodiinae (1/c 70; Australia), Allolepiinae (1/1; S U.S.A., Mexico), Hilariinae (1/10; S U.S.A., Mexico, Guatemala), Kaliniinae (1/1; SW U.S.A. to central Mexico), Orcuttiinae (2/6; California), Zaqiqahinae (1/1; Ethiopia, Somalia, Socotra, SW Arabian Peninsula), Aeluropodinae (2/8; Mediterranean, SW Asia to India and N China, coasts of the Red Sea, tropical and southern Africa), Orininae (2/19; S Europe, Türkiye to temperate E Asia), Perotidinae (3/18; tropical and subtropical Africa, S Madagascar, temperate to tropical Asia east to China, New Guinea and N Australia), Farragininae (2/10; S tropical and S Africa), Sohnsiinae (1/1; Mexico) do not occur in South America.

■ CYNODONTEAE INCERTAE SEDIS: Kampochloa (1; Zambia, Angola; possibly close to Ctenium?), Lepturidium (1; Cuba), Vietnamochloa (1; Vietnam).

■ SUBTRIBE JOUVEINAE ‣ all genera in South America.

203. Jouvea E. Fourn. Two spp. from Mexico to Colombia and Ecuador in South America (1).

■ SUBTRIBE DACTYLOCTENIINAE ‣ outsiders Acrachne (3; tropical and S Africa, Madagascar, SE Asia, Australia), Dactyloctenium (13; tropical and subtropical regions on both hemispheres), Brachychloa (2; KwaZulu-Natal, Mozambique).

204. Neobouteloua Gould. Two spp. endemics to N Argentina.

■ SUBTRIBE ELEUSENINAE ‣ outsiders Dinebra (6; tropical E and NE Africa, Madagascar, SW Asia to India, S U.S.A.), Astrebla (4; Australia), Tetrapogon (11; Canary Islands, Africa, Arabian Peninsula, the Middle East to India and Central Asia), Lepturus (c 15; coasts in E Africa, South Africa, Sri Lanka, Australia to Polynesia), Micrachne (5; Central and tropical E Africa), Chrysochloa (4; tropical Africa), Stapfochloa (6; tropical and subtropical Africa), Cynodon (10; tropical and subtropical regions of Old Hemispheres), Afrotrichloris (2; Somalia), Schoenefeldia (2; tropical and S Africa, Madagascar, India), Apochiton (1; Tanzania), Austrochloris (1; Queensland), Coelachyrum (1–8; tropical and southern Africa, Arabian Peninsula, Pakistan), Daknopholis (1; E Africa, Madagascar, Aldabra), Oxychloris (1; Australia), Harpochloa (1; S tropical and S Africa), Neostapfiella (3; Madagascar), Pommereulla (1; S India, Sri Lanka), Sclerodactylon (1; coast of E Africa, islands in the Indian Ocean).

205. Chloris Sw. c 55 spp., warm-temperate to tropical regions on both hemispheres; 32 spp. in New World, 20 in South America, 9 in Brazil, C. exilis Renvoize endemic.

206. Diplachne P.Beauv. (off Leptochloa). Two spp., one over widely distributed, D. fusca (L.) P.Beauv. ex Roem. & Schult., trop. & subtrop. to N. America, inc. almost all countries of New World.

207. Disakisperma Steud. 4 spp., three in Ethiopia to S. Africa, Arabian Pen., and D. dubium (Kunth) P.M.Peterson & N.Snow, WC & S U.S.A. to Honduras, Colombia to Bolivia and N Argentina.

208. Eleusine Gaertn. 9 spp., trop. & subtrop. Old World, Brazil to S. South America, only one sp. in South America, E. tristachya (Lam.) Lam., from Brazil, Chile, Argentina, Paraguay and Uruguay.

209. Eustachys Desv. 15 spp., U.S.A., Mexico, Central America, the Caribbean, South America (12), one sp. in E and S Africa, Oman and Yemen, and one in SE Asia to Taiwan (China) and New Guinea; 11 spp. in Brazil, one endemic.

210. Leptochloa P.Beauv. (inc. Trichloris, Enteropogon, exc. Diplachne). 29 spp., tropical and S Africa, Australia, Marquesas Islands, North and South America; 19 spp. in New World, 13 in South America, 5 in Brazil, none endemics.

211. Microchloa R.Br. 6 spp., Africa, one sp. in tropical and subtropical regions on both hemispheres; two spp. in New World, over region from North America to southern South America, one widely in Brazil.

M. indica Desv. is one of three desiccation tolerant Poaceae in New World, with dry specimens collected in NE Brazil and in Argentina.

212. Rheochloa Filg. et al. Perenials, culm 60-75 cm, sometimes flexuous to decumbent, rroting at the lower nodes, unbranched at vegetative portions. Only one sp., R. scabriflora Filg., P.M.Peterson & Y.Herrera, central Brazil, endemic to SE Goias state, mesophytic (in humid grassland), a rare plant in Brazil, by Plantas Raras do Brasil’s book, possibly extinct.

213. Stapfochloa H.Scholz. 8 spp., one in Africa; 3 from Brazil, Argentina,Paraguay and Uruguay; 2 from Argentina, Bolivia and Paraguay absents in Brazil; and 2 widely distributed in New World.

■ SUBTRIBE PAPPOPHORINAE ‣ all genera occur in South America.

214. Neesiochloa Pilg. Small herbs up to 26cm high. Only one sp., N. barbata (Nees) Pilg., endemic dry seasonal scrubland of NE Brazil (caatinga), in Bahia, Ceará and Piauí states.

215. Pappophorum Schreb. 9 spp., S U.S.A. to Argentina, 8 in South America, 5 in Brazil, one endemic.

216. Tridens Roem. & Schult. 14 spp., Angola, E U.S.A. and southwards to Argentina; 5 spp. in South America, four restricted in Brazil (with one endemic) and adjacent Argentina, Uruguay and Paraguay, and T. flaccidus (Döll) Parodi, from Colombia to Guyana and Brazil; T. eragrostoides (Vasey & Scribn.) Nash in J.K.Small, occur in S. U.S.A. to Mexico, Cuba and Venezuela; absent in Ecuador, Peru, Bolivia and Chile.

■ SUBTRIBE BOUTELOUINAE ‣ a single genus.

217. Bouteloua Lag. 57 spp., Canada, U.S.A. and Mexico, Central America, the Caribbean, South America to Argentina, centered in the SW U.S.A and Mexico; 13 spp. in South America, mainly widely distributed, four of then reaches from North and Central American up to Caribbean of Venezuela and Colombia, 4 in Brazil, one endemic.

■ SUBTRIBE MONANTHOCHLOINAE ‣ a single genus.

218. Distichlis Raf. 11 spp., 1 sp. in Australia, remaining in U.S.A., Mexico, the Caribbean, South America; 6 spp. from Venezuela to Uruguay, also in Paraguay, and one from North America to W South America, absent in Paraguay and Brazil.

■ SUBTRIBE MUHLENBERGIINAE ‣ a single genus.

219. Muhlenbergia Schreb. Herbs, sometimes scandent. 170 spp. in Canada, U.S.A., Mexico, Central America, tropical and subtropical South America, few species in Central to NE Asia, Himalayas and SE Asia to New Guinea, centered in N Mexico and the SW U.S.A. 36 spp. in South America, 7 in Brazil, none endemics.

■ SUBTRIBE SCLEROPOGONINAE ‣ outsiders Swallenia (1; Eureka Valley Sand Dunes in Inyo County in California).

220. Blepharidachne Hack. 4 spp., two only W U.S.A. and N Mexico, and two only N Argentina.

221. Erioneuron Nash. Three spp. in S U.S.A. and Mexico, two also in Bolivia and Argentina.

222. Munroa Torr. 5 spp., one in W Canada to W & C U.S.A., NE Mexico and four in Peru to N. Argentina and Chile.

223. Scleropogon Phil. Only one sp., S. brevifolius Phil., SW & WC U.S.A. to Mexico, disjunct in NW Argentina.

■ SUBTRIBE TRAGINAE ‣ outsiders Polevansia (1; Cape, Lesotho), Pogononeura (1; tropical Africa), Monelytrum (1; S Angola, Namibia), Orthacanthus (1; S Africa).

224. Tragus Haller. 8 spp., 7 in Old World and a single native species, T. andicola M.A.Zapater & Sulekic, endemic to NW Argentina.

225. Willkommia Hack. 4 spp., three in Africa and W. texana Hitchc. from North America (Texas, Ohlahoma) and coastal Argentina and Uruguay.

■ SUBTRIBE CTENIINAE ‣ a single genus.

226. Ctenium Panz. 20 spp., tropical and subtropical regions of Africa and North and South America, America, Madagascar. 11 spp. in New World, 7 in South America, all in Brazil, two endemics, also in Colombia, Paraguay and Suriname.

■ SUBTRIBE GOUININAE ‣ outsiders Triplasis (2; C and E U.S.A. to Costa Rica), Tridentopsis (2; S U.S.A., Mexico), Vaseyochloa (1; Texas).

227. Gouinia E. Fourn. ex Benth. & Hook. f. (exc. Schenckochloa). 10 spp. from Mexico to Brazil, Bolivia and Argentina; 5 in South America, all in Brazil, one endemic.

228. Schenckochloa J.J.Ortíz. (off Gouinia). Only one sp., S. barbata (Hack.) J.J.Ortíz, endemic to NE Brazil in Rio Grande do Norte, Paraíba and Pernambuco states.

229. Triplasiella P.M.Peterson & Romasch. (off Tridentopsis) Plants caespitose perennials with knotty, short rhizomatous bases; culms 20-100 cm tall, nodes often bearded, the hairs up to 2 mm long, soft. Only one sp., T. eragrostoides (Vasey & Scribn.) P.M.Peterson & Romasch., in S U.S.A., E and S Mexico, the Caribbean, and Venezuela, in shrubby grasslands, desert scrub, open ground, cliffs, and rocky sites; 0-2,300 m.

■ SUBTRIBE HUBBARDOCHLOINAE ‣ outsiders Dignathia (5; tropical E Africa, W India), Leptocarydion (1; N Namibia, KwaZulu-Natal, Mpumalanga, N Province, Botswana, E Africa), Bewsia (1; N Namibia, N South Africa, Swaziland), Lophacme (2; S tropical and S Africa), Hubbardochloa (1; mountains in Central Africa), Decaryella (1; Madagascar).

230. Gymnopogon P. Beauv. 14 spp., tropical and subtropical regions in over America, one also from India to Thailand; 10 spp. from South America, 7 in Brazil, one endemic.

231. Leptothrium Kunth. Three spp., two from Cape Verde to Egypt and Tanzania, Arabian Pen. to Pakistan, and L. rigidum Kunth from Caribbean, Colombia and Venezuela.

■ SUBTRIBE TRICHONEURINAE ‣ a single genus.

232. Trichoneura Andersson. 7 spp., 4 in tropical Africa and Arabian Peninsula, one in Texas, one from Galapagos and one from Peru and Chile.

■ SUBTRIBE TRIPOGONINAE ‣ outsiders Desmostachya (1; North Africa, SW Asia to India and SE Asia), Melanocenchris (3; Chad and tropical NE Africa to India and Sri Lanka), Halopyrum (1; coasts along the Indian Ocean), Eragrostiella (6; E Africa, Sri Lanka to N Australia), Oropetium (6; arid and semiarid regions in tropicl W Africa to India and SE Asia).

233. Tripogon Roem. & Schult. (exc. Tripogonella). 52 spp., 50 in Old World and two from Bolivia, Chile, Argentina, Paraguay and Uruguay.

234. Tripogonella P.M.Peterson & Romasch. (off Tripogon). Three spp., one in Australia and New Guinea, another in tropical and southern Africa, and T. spicata (Nees) P.M.Peterson & Romasch. from Texas to C Argentina, Venezuela, S, C & SE Brazil and Caribbean; this species is one of three desiccation tolerant Poaceae in New World, with dry specimens collected in Brazil and Argentina.

19. ZINGIBERALES

FAMILIES ABSENTS IN SOUTH AMERICA: LOWIACEAE (1/32) AND MUSACEAE (3/91).

The basal branching in Zingiberales is still unclear, although Musaceae are sometimes identified (with low support) as sister to the rest. Heliconia (Heliconiaceae) is part of the basal polytomy. Strelitziaceae and Lowiaceae appear to be sister-groups. According to Stevens (2001 onwards) these share the following potential synapomorphies: pseudopetiole with adaxial and abaxial series of air canals; perianth whorls coloured, distinct; adaxial stamens of inner whorl sterile; floral column (sterile apex of ovary) present; stigma trilobate; aril hairy; and exostomal aril lobed or fimbriate. Three distinct lineages in New World.

LINEAGE 1 of 3: HELICONIACEAE

HELICONIACEAE

§ FULLY CHLOROTROPHIC

Genera/spp. 1/198 Distribution tropical and subtropical regions of Mexico, Central and South America, some spp. in E Malesia (New Guinea and eastwards), Melanesia and eastwards to Samoa. Habit bisexual, perennial (often giant) herbs. Rhizome starchy, pseudostems present, buds axillarys.

Key differences from similar families:

ü One ovule per locule (vs. many in Musaceae and Strelitziaceae).

ü Bisexual flowers (vs. unisexual flowers in Musaceae).

ü 5 fertile stamens (vs. 1 fertile stamen in Marantaceae, Costaceae, Cannaceae and Zingiberaceae).

SYSTEMATIC a single genus.

1. Heliconia L. Medium to very large sized rhizomatous herbs (up to 10 m tall), habit musoid, cannoid or zingiberoid; inflorescences are almost always terminal, but in some spp. may emerge directly from the rhizome; inflorescence erect or pendulous, arrangement of bracts along the rachis is plane or spirally; fruit berries, green or yellow when immature and turning deep blue or purple when mature. 198 spp., throughout the Neotropics, 6 from Molucas to Vanuatu in Pacific Ocean and 183 in New World, 138 in South America, highly centered in Colombia (101), in montane rainforests, in shaded moist forests or open disturbed habitats, up to 2000 m, in extensive, conspicuous stands or as individuals; often pioneer spp.; hummingbird pollinated; 29 in Brazil (only 5 endemics), only in Amazon rainforest and Atlantic Forest, barely or absent in others areas; 5 subgenera:

§ subg. Heliconia ‣ erect inflorescence, flowers non-resupinate; musoid; Neotropics.

§ subg. Heliconiopsis ‣ erect or pendant inflorescence, flowers non-resupinate; musoid; Sulawesi to Fiji and Samoa.

§ subg. Stenochlamys ‣ erect inflorescence, flowers resupinate; musoid, canoid, zingibeoid; Neotropics.

§ subg. Taeniostrobus ‣ erect inflorescence, flowers resupinate or non-resupinate; musoid; Neotropics.

§ subg. Griggsia ‣ pendant inflorescence, flowers non-resupinate; musoid; Neotropics.

LINEAGE 2 of 3: STRELITZIACEAE

STRELITZIACEAE

§ FULLY CHLOROTROPHIC

Genera/spp. 3/12 Distribution Phenakospermum: tropical South America; Strelitzia: temperate and subtropical regions in South Africa; Ravenala: Madagascar. Habit bisexual, perennial (often giant) herbs (aerial stem in Phenakospermum, Ravenala and Strelitzia nicolai lignified). Rhizome rich in starch, with short and thick branches. Dichotomous branching present in Strelitzia. Strictly, one Neotropical genus Phenakospermum with, but treated here to include both the bird of paradise plant Strelitzia reginae Aiton and the travellers palm Ravenala madagascariensis Sonn., both of which are introduced into the Neotropics. Strelitzia reginae Banks ex Aiton cultivated and naturalized in the Neotropics; Ravenala madagascariensis (travelers palm), introduced from Madagascar.

Key differences from similar families

ü Pseudostem absent, staminodes lacking and 4 to many ovules per locule vs. pseudostem present, 1 petaloid staminode and 1 ovule per locule in Heliconiaceae.

ü Leaves distichous and flowers bisexual vs. leaves spiral and flowers unisexual in Musaceae.

ü 5 fertile stamens vs. 1 fertile stamen in Marantaceae, Costaceae, Cannaceae and Zingiberaceae.

ü Zingiberaceae differ by being aromatic vs. Strelitziaceae is non-aromatic.

Useful tips for generic identification:

ü Phenakospermum: terminal inflorescence; 5 stamens.

ü Ravenala: inflorescence lateral; 6 stamens.

SYSTEMATIC outsiders Ravenala (1; Madagascar) and Strelitzia (4–5; temperate and subtropical regions in southern Africa).

1. Phenakospermum Endl. Arborescent, suckering plants; leaves in fanlike clusters on the top of the trunk; inflorescence terminal. Only one sp., P. guyannense (Rich.) Endl. ex Miq., found in the Neotropics east of the Andes from Guianas and Maranhão state to Ecuador in western and Bolivia and south; particularly abundant in forests bordering seasonally flooded savannas, along river margins, in swamp forests, forest edges, and disturbed areas, 50-700 m; know as ‘sororoca’ in Brazil; pollinated by bats; flowers of this spp. stay receptive for only one night, and seeds are cooked and eaten by some indigenous people.

LINEAGE 3 of 3: CORE ZINGIBEROID

MARANTACEAE

§ FULLY CHLOROTROPHIC

Genera/spp. 29/520–530 Distribution pantropical except Australia, with their largest diversity in tropical America. Habit bisexual, perennial (sometimes giant) herbs (rarely climbing; sometimes with woody rhizome). Rhizome rich in starch. Marantaceae leaves are held vertically during the night!

Phylogenetic studies at the family level have shown Neotropical diversity to primarily involve two large groups: the Maranta clade with nine genera and ca. 70 spp. is concentrated in SE Brazil; and the Calathea clade with five genera and ca. 370 spp. is distributed throughout the Neotropics but with highest diversity in northwestern South America. Another character exclusive to this family in the order is the presence of paired flowers, in which th two appear as mirror images of each other. Only sp. in the genus Monotagma have single flowers. These plants are easy to grow in tropical and subtropical climates where they are cultivated as landscape plants and as potted plants in temperate climates.

There is a growing demand for species of Calathea, Ctenanthe, Maranta, and Stromanthe as ornamentals. These plants are easy to grow in tropical and subtropical climates where they are cultivated as landscape plants and as potted plants in temperate climates. This family includes a number of edible spp. such as Calathea barbata Petersen and Calathea allouia Lindl., from which the tubercules are cooked and eaten by indigenous people in central Brazil and northern South America, respectively. However, only a single spp., Maranta arundinacea L. (arrowroot) is economically important. Fibre plants are abundant in the family; the inflorescence peduncles of species of Ischnosiphon are used for making baskets, mats, strings for musical instruments and ornaments in the Amazonian region and the Guianas; species with larger leaf blades are used for wrapping foods, to cover cargo and as bottle stoppers.

SYSTEMATIC six lineages, the three most basal not occur in South America: Sarcophrynium Clade (1/6; tropical Africa), Donax Clade (4/9; tropical W and C Africa, SE Asia, Malesia to Vanuatu), Hypselodelphys Clade (3/14; tropical W and C Africa; New Guinea; Japan); Monophyllanthe unplaced, but accepted here near Maranta.

UNPLACED

1. Monophylanthe K. Schum. Aerial shoots with a single basal leaf each, small to medium sized plants (up to 1,5 m tall), flowers are small, white, with or without the external staminodes, disposed in both sides of the inflorescence, with very small or absent bracteoles. Two spp. from N Amazon rainforests of Brazil (both species, none endemics), Colombia and Guianas.

THALIA CLADE (1/6) ‣ a single genus.

2. Thalia L. Rosulate or caulescent marsh plants; inflorescence a very richly branched synflorescence; spathes caducous; sepals very short; floral tube indistinct; outer staminode solitary; cucullate staminode with two appendages; style with a very long ventral projection of the stigmatic orifice. 6 spp., T. geniculata L. widely distributed, weed confined of open marshes, reported also in Africa, three almost confined to Brazil (two endemics and one up to Cono Sur), Ecuador and U.S.A. one endemic each.

CALATHIDS (6/c. 380) ‣ outsider Phrynium (35–40; India to southern China and SE Asia, New Guinea, Vanuatu).

3. Calathea G. Mey. Herbs, usually with several basal and one or more cauline leaves; leaves uniformly green or paler below; inflorescence terminal on a leafy shoot, compound, usually composed of 2-several similar, partial inflorescences, rarely simple; flowers open; corolla tube elongate, lobes reflexed or recurved; fruit usually 3-seeded, dehiscent. 58 spp. from Mexico and the Caribbean islands to Brazil and Bolivia, 38 in South America, 4 in Brazil, one endemic.

4. Goeppertia Nees. Herbs with basal and/or cauline leaves; leaves uniformly green or with various types of patterning (seventh largest diversity of variegated leaves worlwide, 28), this sometimes present only in juvenile plants; inflorescence terminal on a leafy shoot or more rarely borne on a separate leafless shoot, simple, or rarely compound and composed of up to 9 similar, partial inflorescences; flower groups brachyblastic; flowers open or remaining closed, rarely self-fertile; corolla tube elongate; fruit 3-seeded. 201 spp. from Mexico and the Carribean islands to Argentina and Paraguay, 168 in South America, 93 in Brazil, 47 endemics.

5. Ischnosiphon Korn. Rosulate or caulescent, often very tall herbs (up to 10 m height), with aerial stems and resembling shrubs or lianas; inflorescence simple or distinct synflorescence; sclerotic bracts; floral tube very long and narrow; outer staminode solitar. 36 spp., distributed over the Neotropics, 35 in South America, 20 in Brazil, 4 endemics.

6. Monotagma K. Schum. Leaf blades firm and chartaceous when dry; inflorescence a richly-branched synflorescence, often much congested; bracts tough and coriaceous, often woody in texture when dry, conduplicate; cymules one-flowered; flower tube very long and narrow; outer staminode solitary. 39 spp., 38 in South America (only three up to Central America and Trinidad e Tobago), mainly restricted to the Amazon rainforest; 22 spp. in Brazil, 10 endemics; the Manaus region is recognized as one of the centers of diversity for this genus.

7. Pleiostachya K. Schum. Laterally flattened inflorescences with conduplicate and much overlapping bracts; bracts markedly fibrous and persistent; floral tube very long and narrow; outer staminode solitary; callose staminode entirely firm and fleshy. Three spp. from Pacific lowlands of Mexico, Central America (one restricted), Colombia (one endemic) and Ecuador.

MARANTIDS (11/c. 110) ‣ outsiders Halopegia (3; Congo; Indochina, Java; Madagascar), Marantochloa (18; tropical Africa, Madagascar, the Comoros), Stachyphrynium (8; India and Sri Lanka to Tibet, Yunnan, SE China, Andaman Islands, SE Asia, W Malesia to Sulawesi), Afrocalathea (1; Central Africa).

8. Ctenanthe Eichler. Rosulate or caulescent plants with antitropic leaves; cauline leaves mostly clustered; inflorescence a sparsely to moderately branched; corolla tube very short and wide; bracts long-persistent; two outer staminodes slightly unequal; callose staminode distally petaloid and showy. 16 spp. from South America, only two up to southern Central America and Trinidad e Tobago, with the highest diversity in the Brazilian Atlantic forest; 15 spp. in Brazil, 11 endemics.

9. Maranta L. (inc. Hylaeanthe, Koernickanthe, Myrosma) Rhizomes usually with some form of specialization, root tubers usually present; inflorescence usually lax, simple or more complex; floral tube moderately long and narrow (rarely short); two outer staminodes subequal to unequal; sepals persistent on the fruit. 54 spp., all in South America (5 up to Mexico and Trinidad & Tobago), 43 in Brazil, 26 endemics; M. gigantea N. Luna & E. M. Pessoa and M. zingiberina L. Andersson, from Bahia, Paraíba, Pernambuco and Alagos states in NE Brazil are the tallest species in this genus, up to 1,8 m tall.

10. Saranthe (Regel & Körn.) Eichler. Hygrophytic habit; bracts deciduous (rarely long-persistent), herbaceous to membranaceous, thin-papiraceous when dry; floral tube short and wide; two outer staminodes, equal; fertile stamen with a petaloid, narrowly oblong appendage equalling or somewhat exceeding the anther. 9 spp., 8 from Atlantic Forest of SE Brazil, with S. eichleri Petersen up to adjacent Argentina, and S. urceolata Petersen endemic to Peru.

11. Stromanthe Sond. Usually caulescent herbs, usually clustered leaves; inflorescence a richly branched, but often diffuse, synflorescence; leaves antitropic; flower tube very short and wide to inconspicuous; sepals very large and fibrous; outer staminodes completely absent or two rudimentary to petaloid and showy. 20 spp. from Mexico, Caribbean, Central and tropical South America (15), centered in Central America and SE Brazil (9, 7 endemics).

CANNACEAE

§ FULLY CHLOROTROPHIC

Genera/spp. 1/12 Distribution over tropical New World; Habit herbs. An originally Neotropical family with the single genus Canna L., wdely cultivated all over the tropics and tubtropics, and often escaped and naturalized. Canna are devoid of hairs, but often the leaves, sheaths, inflorescence, bracts, and sepals are described as glaucous or woolly, scurfy, lanuginose, or waxy. This aspect is caused by an epicuticular wax layer.

SYSTEMATIC a single genus.

1. Canna L. Perennial, often large-sized, glabrous herbs, with rhizomes; stems unbranched; leaves distichously to spirally arranged; lamina with closely set, pinnately arranged, parallel secondary veins; inflorescence terminal, bracteate, a thyrse, with 2- or 1-flowered cincinni; flowers asymmetric; sepals 3, free; corolla composed of 3 unequal petals; fruit a capsule; seeds numerous, without an aril, with a so-called imbibition lid. 10 spp., 3 up to almost 3,000 m in Andes of Peru to Bolivia, C. flaccida Salisb. endemic to U.S.A., C. pedunculata Sims restricted of SE Brazil in Rio Grande do Sul, Rio de Janeiro, and Santa Catarina, in swampy areas and forest margins; remaining five widely distributed (3 of then in Brazil), some in low, open and wet vegetation, others prefer forested areas; several spp. thrive well in secondary vegetation, along roads, in coffee plantations, and as weed in cultivated areas.

ZINGIBERACEAE

§ FULLY CHLOROTROPHIC

Genera/spp. 57/1,440-1,460 Distribution tropical and southern Africa, Madagascar, Mascarene Islands, Seychelles, S, E and SE Asia, Malesia, New Guinea, northern and E Australia (especially Queensland), Pacific islands; some spp. of Renealmia in tropical America (Central America, Caribbean, northern and central South America). Habit bisexual, perennial (sometimes giant) herbs, often with a pseudostem consisting of superimposed leaf sheaths. Sometimes with stilt roots. Aromatic. Rhizome and roots usually rich in starch (root cells sometimes with starch nodules).

Neotropical Zingiberaceae can be distinguished from all other families in this order by being aromatic and in having an indument often composed of stellate hairs. Aframomum melegueta K.Schum. is cultivated in the Guianas for its spicy fruit. Several spp. of Curcuma and Zingiber are cultivated for their spicy rhizomes. Hedychium coronarium J.Koenig is naturalized all over the Neotropics, and in the Andes it is even a pest, replacing the original vegetation along roads. The other genera are cultivated as ornamentals, important spp. being Alpinia purpurata K.Schum. and Etlingera elatior (Jack) R.M. Sm.

SYSTEMATIC four clades, Siphonochiloideae (1/11, tropical and southern Africa, Madagascar), Tamijioideae (1/1, NW Borneo) and Zingiberoideae (33/610–630, Madagascar, Himalayas, E and tropical Asia, tropical Australia) absent in South America; among the South American group, Alpinioideae, 26 genera occur only in tropical and southern Africa, Madagascar, Mascarene Islands, Seychelles, tropical Asia, tropical Australia and islands in the Pacific.

1. Renealmia L. f. Herbs with well-developed rhizomes and pseudostems; inflorescence usually lax, sometimes branched, terminal on a separate leafless shoots, erect or prostrates, or rarely terminal in a leafy shoot; 1-several-flowered cincinni; flowers white to yellow, tubular to tunicate. 84 spp., 20 in S Africa and 64 spp. in New World, from Mexico to Bolivia and S Brazil, also present in Caribbean, mainly forests; 56 spp. in South America, 21 in Brazil (8 endemics); some spp. in Brazil are very narrow endemic, centered in Amazon rainforests; R. aceana Maas from Acre state, R. chrysotricha Petersen from Rio de Janeiro municipality, and R. matogrossensis Maas from Mato Grosso state are rare species in Brazil by Plantas Raras do Brasil’s book, being the unique rare Zingberales in Brazil.

COSTACEAE

§ FULLY CHLOROTROPHIC

Genera/spp. 8/c. 155–160 Distribution pantropical, with their largest diversity in Central and South America, New Guinea and northern Australia. Habit bisexual, perennial (sometimes giant) herbs, often with aerial stem. Rhizome rich in starch. Root nodules rarely present. Rarely epiphytic. Rarely with axillary bulbils. Stem usually simple (in Tapeinochilos branched, with lateral branches penetrating leaf sheaths). Non-aromatic herbal plants.

Costaceae have various flower characters in common with Zingiberaceae, but they differ by having non-aromatic leaves and closed leaf sheaths.

Key to genera of the Neotropical Costaceae:

1. Flowers solitary in the axils of the upper leaves ------------ Monocostus

1. Flowers in a terminal spike - 2

2. Ovary 2-locular; bracteole tubular ------------ Dimerocostus

2. Ovary 3-locular; bracteole folded or tubular - 3

3. Bracteole tubular, bicarinate at the abaxial side; stigma cup-shaped ------------ Chamaecostus

3. Bracteole folded; stigma 2-lamellate, always provided with a dorsal 2-lobed appendage ------------ Costus

SYSTEMATIC two well defined clades.

BASAL LINEAGE (4/11) ‣ all genera in South America.

1. Chamaecostus C.D. Specht & D.W.Stev. Low or very small, occasionally acaulescent plants, never exceeding 1 m in height; bracts herbaceous to chartaceous, green or yellow; bracteole tubular, bicarinate at the abaxial side; stigma cup-shaped. 8 spp. from French Guiana to Colombia, Brazil (7, three endemics), Bolivia and Peru.

2. Dimerocostus Kuntze. Very tall herbs. Leaves large, usually long-acuminate. Three spp., D. strobilaceus Kuntze from Honduras to Peru, Bolivia and Acre state, Brazil; two remining in Colombia to Bolívia.

3. Monocostus K. Schum. Low herbs; is unique in the family in having solitary, axillary flowers. One sp., M. uniflorus (Poepp. ex. Petersen) Maas, restricted to low forests near the town of Tarapoto in the San Martin districts in E Peru.

COSTUS COMPLEX (4/120-124) ‣ outsiders Paracostus (2; tropical W and C Africa; Borneo), Hellenia (2; SE Asia, W Malesia; India, Andaman and Nicobar Islands, S China, SE Asia, Malesia to New Guinea, Bismarck Archipelago and Queensland), Tapeinochilos (16–20; Moluccas, New Guinea to tropical Australia).

4. Costus L. Low to tall terrestrial herbs, usually over 1 m tall; sometimes branched; the leaves are arranged in a spiral along the pseudostem or the pseudostem itself may be spiraled; in most spp. the inflorescence is at the end of a leafy stem, but sometimes it may emerge directly from the rhizome. 117 spp., pantropical; 74 in New World, 56 in South America, 20 in Brazil, mostly in Amazon rainforest, 4 endemics, with C. atlanticus E. Pessoa & M. Alves endemic to Pernambuco state, highly diverse in Colombia (33).

20. CERATOPHYLLALES

A SINGLE FAMILY, PRESENT IN SOUTH AMERICA

CERATOPHYLLALES

§ FULLY CHLOROTROPHIC

Genera/species 1/4–6 Distribution cosmopolitan except polar areas. Habit monoecious, perennial herbs. Aquatic, with submersed stems and leaves, sometimes anchored in the sediment through rhizoid branches. Branches extra-axillary, alternating with leaves. Roots absent. Submersed aquatic, rootless, glabrous herbs. Throughout the Neotropics in non-marine aquatic habitats. Use Aquarium plants, medicinal plants.

Distinguished from Myriophyllum L. (Haloragaceae) by the branched leaves and achene fruit.

SYSTEMATIC a single genus, Ceratophyllum (6, cosmopolitan).

1. Ceratophyllum L. Submersed aquatic, rootless, glabrous herbs; leaves whorled, simple, finely divided, lobes often toothed; inflorescences extra-axillary and alternating with leaves, spikes or racemes; flowers solitary, unisexual, monoecious, actinomorphic, pedicels short or flowers sessile, bracts foliaceous; calyx of (7)9-12(15) toothed sepals; corolla lacking; stamens 3-many, spirally arranged, free of perianth, filaments short, anthers adnately fixed, dehiscing via full-length slits; fruits achenes, papillose, spiny or smooth; seeds 1, tiny. 6 spp., three only Old World, C. echinatum A. Gray in North America, C. demersum L. very widespread (almost cosmopolitan) and C. australe Griseb. in New World northern of Ecuador Line, the two last in Brazil.

21. RANUNCULALES

FAMILIES ABSENTS IN SOUTH AMERICA: EUPTELEACEAE (1/2), CIRCAEASTERACEAE (2/2).

LINEAGE 1 of 3: LARDIZABALIDS

LARDIZABALACEAE

§ FULLY CHLOROTROPHIC

Genera/species 7/29–34 Distribution Lardizabalaceae except Lardizabala and Boquila: E Himalaya, northern Indochina, China (inc. Taiwan), Hainan, Korean Peninsula, Japan; Lardizabala and Boquila: Chile between the Andes and the Pacific. Habit usually monoecious or dioecious (in Decaisnea polygamomonoecious), usually climbing or scrambling evergreen or deciduous shrubs or lianas (Decaisnea an upright shrub).

SYSTEMATIC subfamily Sargentodoxoideae (1/1, China, Laos, Vietnam) does not occur in South America; among Lardizabaloideae, outsiders are Decaisnea (1; Himalayas to C China), Sinofranchetia (1; W and C China), Akebia (4; China (inc. Taiwan), Korean Peninsula, Japan), Stauntonia (20–25; NE India, Himalayas, China (inc. Taiwan), Korean Peninsula, Japan).

1. Boquila Decne. Lianas, trifoliate leaves, flowers white, fruit one seeded. Only one sp., B. trifoliata (DC.) Decne, Chile and Argentina.

2. Lardizabala Ruiz & Pav. Wood vines, leaves trifoliate, flowers black-purple, male flowers in cymes, female flowers solitary, fruit many seeded. Only one sp., L. biternata Ruiz & Pav., endemic to Chile.

LINEAGE 2 of 3: PAPAVERIDS

PAPAVERACEAE

§ FULLY CHLOROTROPHIC

Genera/species 46/c. 755 Distribution temperate and subtropical parts if the Northern Hemisphere, Central America, northern South America, E African mountains, southern Africa, Macaronesia, with their largest diversity in Mediterranean, W, C and E Asia, and the SW U.S.A. Habit usually perennial, biennial or annual herbs (sometimes climbing; some species of Argemone, Bocconia, Dendromecon, Hunnemannia, and Romneya evergreen shrubs or small trees). Some species have tuberous rhizome or tuberous roots. Stem sometimes pachycaul. Use ornamental plants, baking (seeds from Papaver somniferum), seed oils for soap (Argemone, Glaucium, Papaver), medicinal plants, narcotics (opium from Papaver somniferum and P. bracteatum). Only subfamily Papaveroideae (23/210–240) in South America.

SYSTEMATIC subfamilies Pteridophylloideae (1/1, Japan) and Fumarioideae (20/c 520, mainly temperate regions in the Northern Hemisphere, southern Africa) do not occur in South America; among Papaveroideae, tribe Eschscholzieae (3/13–15, W North America, northern and E Mexico) does not occur in South America.

1.1 PAPAVEROIDEAE ▸ TRIBE PAPAVEREAE (8/c. 260) - outsiders are Arctomecon (3; Mojave Desert in the U.S.A.), Papaver (223; Europe, the Cape Verde Islands, Mediterranean, S Africa, temperate and subtropical Asia, W North America), Romneya (1–2; S California, N Baja California), Canbya (2; W U.S.A.; W Mojave Desert), Hesperomecon (1; S Oregon, California), Meconella (3; W U.S.A.), Platystemon (1; W U.S.A., Baja California).

1. Argemone L. Annual or biennial herbs, spine, with yellowish latex; flowers solitary, terminal, 4-6 white petals. 28 spp., about 25 in North America from California to Florida and N Mexico; one in Hawaii; and three in South America, all currently native from to Chile : A. crassifolia Ownbey, known only of Quebrada de Cañas, Atacama; A. rosea Hook., known only regions III and IV, in dry rivers, rock slopes, 0-900 m; and A. subfusiformis G.B. Ownbey, reaching from Peru and W Argentina, Ecuador, S Paraguay and S Uruguay.

1.2 PAPAVEROIDEAE ▸ TRIBE CHELIDONIEAE (9/45–50) - outsiders are Sanguinaria (1; S Canada, E U.S.A.), Eomecon (1; E China), Dicranostigma (8; Himalayas, W China), Eomecon (1; E China), Glaucium (c 25; W Europe, Mediterranean, SW to Central Asia), Chelidonium (1; Europe, temperate and subarctic Asia), Hylomecon (2; NE China, Korean Peninsula, Japan), Macleaya (2; temperate China (inc. Taiwan), Japan), Stylophorum (3; temperate China, E U.S.A.).

2. Bocconia L. Perennial herbs to small trees, pachycaulous 2-6 m, purplish to greysh lates; inflorescences lateral or terminal paniculates; flowers perfect. 8 spp., Mexico (7, 4 endemics) to Bolivia and Venezuela; two spp. in South America, B. frutescens L. from Mexico to Colombia and Caribbean, and B. integrifolia Bonpl. restricted at Andes from Venezuela to Bolivia.

LINEAGE 3 of 3: RANUNCULIDS

BERBERIDACEAE

§ FULLY CHLOROTROPHIC

Genera/species 13/650-700 Distribution Eurasia from W Europe eastwards to Malesia, North Africa (the Atlas Mountains), E African mountains, North and Central America, mountain areas in South America. Habit bisexual, evergreen or deciduous shrubs or biennial to perennial rhizomatous or tuberous herbs (rarely trees). Roots and rhizome often bright yellow inside due to presence of berberine. All herbs or woody shrubs; Northern hemisphere and South America. One genus in Neotropics and South America.

Berberis is easy to recognize at generic level, however species are sometimes difficult to delimit especially due to the great variation in leaf size and shape and the great number of species described. Berberin (an alkaloid) is used as a dye (yellow). Fruits are eaten locally in jams.

Use Ornamental plants, fruits (Berberis), medicinal plants, dyeing (yellow) substances.

SYSTEMATIC subfamilies Podophylloideae (6–8/c 90, temperate regions on the Northern Hemisphere) and Nandinoideae (4/15, E Europe to E Asia) do not occur in South America; among Berberidoideae, Ranzania (1; Japan) is a outsider.

1. Berberis L. Shrubs or rarely small trees, often spiny, up to 10 m; spines as reduced leaves of long shoots, palmate to leafy, simple or 3-parted; leaves alternate, along the stem or often in rosettes, simple (always in South America) or 1-odd-pinnately compound; inflorescences terminal, usually racemes, rarely umbels or flowers solitary; flowers 3-merous, yellow to orange, sometimes tinged with red; fruit a berry, less than 1 cm, reddish-brown turning dark purple to black, usually juicy; seeds 1 to few. 621 spp., diverse worldwide in the northern Hemisphere, 179 in New World it reaches South America along the Andes (with 133 spp. in continent, common in paramos and punas), extending to Tierra del Fuego and east to S & SE Brazil (4): B. laurina Thunb. up to Cono Sur, and three endemics, inc. B. kleinii Mattos, endemic to high mountains of Santa Catarina state, and as a rare plant in Brazil, by Plantas Raras do Brasil’s book; all species in South America belong to simple-leaved Australes group., with 16 sections:

§ sect. Actinacanthae ‣ 20 spp., Argentina, Chile.

§ sect. Agapatenses ‣ 2 spp., Peru, Bolivia.

§ sect. Buxifoliae ‣ 13 spp., Argentina, Chile, Uruguay, Brazil.

§ sect. Confertae ‣ 12 spp., Peru, Bolivia, Ecuador, Colombia.

§ sect. Corymbosae ‣ 4 spp., Chile, Juan Fernández Islands.

§ sect. Goudotiae ‣ 7 spp., Venezuela, Colombia, Ecuador, Peru.

§ sect. Illicifoliae ‣ 15 spp., Argentina, Chile, Uruguay, Brazil.

§ sect. Latifoliae ‣ 2 spp., Peru, Costa Rica.

§ sect. Laurinae ‣ 21 spp., Argentina, Chile, Bolivia, Peru, Brazil, Uruguay, Ecuador, inc. B. laurina Thunb. from Brazil.

§ sect. Microphyllae ‣ 2 spp., Argentina, Chile, Falkland Islands.

§ sect. Montanae ‣ 5 spp., Argentina, Chile.

§ sect. Paniculatae ‣ 22 spp., Colombia, Ecuador, Peru, Bolivia, Venezuela.

§ sect. Quindiuenses ‣ 5 spp., Bolivia, Chile, Colombia, Ecuador.

§ sect. Trigonae ‣ 2 spp., Argentina, Chile.

§ sect. Truxillenses ‣ 25 spp., Peru, Colombia, Ecuador, Venezuela, Bolivia.

§ sect. Virgatae ‣ 12 spp., Peru, Bolivia, Ecuador.

MENISPERMACEAE

§ FULLY CHLOROTROPHIC

Genera/species 78/c 430 Distribution tropical and subtropical regions in the Northern and Southern Hemispheres, and a few species in temperate E North America and temperate E Asia. Habit dioecious, usually evergreen lianas, or scrambling and climbing perennial herbs (rarely shrubs or trees (Burasaia, Penianthus, Sphenocentrum); one species of Stephania an erect herb). Widely distributed throughout the Neotropics, particularly in the humid lowlands, although some genera (e.g. Cissampelos) also occur in arid areas. Usually vines or lianas with a few exceptions: Abuta grandifolia (Mart.) Sandwith is a tree, and a few species such as Cissampelos ovalifolia DC. are herbaceous. Seventeen genera recorded in the Neotropics. Epiphytes, herbs or trees are barely absent in New World. In North America occurs only Cocculus, Menispermum and Calycocarpum. Use ornamental plants, medicinal plants (Anamirta cocculus (L.) Wight & Arn., Jateorhiza palmata (Lam.) Miers etc), fish- and arrow poisons (curare from Chondrodendron, Curarea, Sciadotenia etc., coccel kernels and picrotoxine from Anamirta), timber. Most species of this family contain powerful alkaloids with biochemical activity. Several are also used in the preparation of arrow and dart poisons, including Curarea, Chondrodendron, Abuta, Telitoxicum. Chondrodendron tomentosum Ruiz & Pav. is the original natural source of the alkaloid tubocurarine, used as a muscle relaxant in surgical procedures. Some sp. (e.g. Abuta grandifolia (Mart.) Sandwith) have edible fruits.

In a sterile condition certain Menispermaceae may be confused with Cucurbitaceae (from which they differ in the lack of tendrils) or Dioscoreaceae (which never have apically flexed pulvinae and whose branchlets often have swollen nodes). They may also be confused with Aristolochiaceae (which also lack the apically flexed pulvinae and sometimes have leafy stipules), and possibly Sparattanthelium (Hernandiaceae), which differs in its ranalean odour and simple stem structure. The leaves of some lianas (e.g. Abuta) may perhaps be confused with those of Loganiaceae (Strychnos), from which they differ in their alternate (rather than opposite) arrangement.

SYSTEMATIC Two subfamilies, both in South America.

1. SUBFAMILY CHASMANTHEROIDEAE (29/c. 160) - two tribes, Coscinieae (3/6, tropical Asia) absent in South America; among Burasaieae, outsiders are Calycocarpum (1; E North America); Parabaena (6; SE Asia, Malesia), Aspidocarya (1; NE India to SW China), Tinomiscium (1; SE Asia, Malesia), Fibraurea (3; India; S China, Indochina; SE Asia, Philippines, Borneo, Sulawesi), Paratinospora (2; China inc. Taiwan), Penianthus (4; tropical W and Central Africa), Sphenocentrum (1; tropical W Africa), Burasaia (4; Madagascar), Orthogynium (1; Madagascar), Dioscoreophyllum (3; tropical Africa), Jateorhiza (2; tropical Africa), Tinospora (36; tropical Asia, tropical Australia), Kolobopetalum (4; tropical Africa), Rhigiocarya (2; tropical W and Central Africa), Hyalosepalum (10; tropical Africa), Sarcolophium (1; tropical Africa), Chasmanthera (2; tropical Africa; Congo), Leptoterantha (1; tropical Africa), Syntriandrium (1; tropical W and Central Africa), Dialytheca (1; Angola), Chlaenandra (1; New Guinea), Platytinospora (1; tropical W and C Africa).

1. Borismene Barneby. Climbers with simple leaves; flowers in pseudopanicles or pseudoracemes. Only one sp., B. japurensis (Martius) Barneby in Bolivia, Brazil, Peru, Colombia, Ecuador and Venezuela.

2. Disciphania Eichler. Climbers with entire or lobed leaves, sometimes leaf-palmately (D. cubijensis (R. Knuth) Sandwith from Brazil, Peru and Bolivia, only at family in South America with no-simple leaves); flowers in pseudospikes or pseudoracemes. 26 spp., C Mexico to NE Argentina, high centered in Amazon rainforest, 20 spp. in South America, 10 in Brazil, 5 endemics.

3. Odontocarya Miers. Climbers with simple leaves; flowers in pseudopanicle. 37 spp. in New World, 35 in South America, four extending to Central America and Lesser Antilles; 21 in Brazil, 5 endemics.

2. SUBFAMILY MENISPERMOIDEAE (45/380–385) - tribes Menispermeae (2/3, E Asia, E North America), Limacieae (1/3, Burma to W Malesia) and Spirospermeae (4/10, tropical Africa (one species), Madagascar) do not occur in South America.

2.1 MENISPERMOIDEAE ▸ TRIBE AMONOSPERMEAE (13/c 80) - outsiders are Diploclisia (2; China; India, Sri Lanka, Burma, S China, SE Asia, Malesia to New Guinea), Sarcopetalum (1; S New Guinea, E Queensland, E New South Wales, E Victoria), Legnephora (5; New Guinea, E Queensland), Parapachygone (1; NE Queensland), Hypserpa (10; SE Asia, Malesia to Polynesia), Pericampylus (3; China (inc. Taiwan), tropical Asia from India to Malesia), Echinostephia (1; SE Queensland, NE New South Wales).

4. Abuta Aubl. Climbers or rarely erect trees with simple leaves. 35 spp. in tropical America (31 in South America); A. grandifolia (Mart.) Sandw. is the unique tree and one of few edible fruit in this family; 18 spp. in Brazil, 5 endemics.

5. Anomospermum Miers. (exc. Elissarrhena, Rupertiella, inc. Orthomene) Climbers with simple leaves; flowers in cymes and pseudoracemes. 9 spp., northen South America, three up to Panamá; 8 in Brazil, A. matogrossense Krukoff & Barneby endemic.

6. Caryomene Barneby & Krukoff. Climbers with simple leaves; flowers in few flowered cymes. 5 spp. in Colombia, Venezuela, Guyana and northern Brazil (all species, none endemics) to Bolivia.

7. Elephantomene Barneby & Krukoff. Climbers with simple leaves; flowers in few flowered cymes. Only one sp., E. eburnea Barneby & Krukoff, rare, N Brazil, Ecuador and Guianas.

8. Elissarrhena Miers. (off Anomospermum). Two spp. in Central and South America, both in Brazil, none endemics.

9. Rupertiella Wei Wang & R. Ortiz. (off Anomospermum) Lianas, dioecious, leaves spiral; blade elliptic to broadly elliptic or obovate, usually plinerved 3–5; finer veins prominent on both surfaces; flowers: sepals 6, fleshy, elliptic or oblong- elliptic; petals 6, submembranous, obovate or flabelliform, margins inflexed, enveloping the individual stamens. Only one sp., R. boliviana (Krukoff & Moldenke) Wei Wang & R. Ortiz, known in Ecuador, Peru, Bolivia and Brazil.

10. Telitoxicum Mold. Climbers with simple leaves; flowers in racemes or panicles. 8 spp. in Brazil (7, 3 endemics), Venezuela, Ecuador, Peru, Colombia and Guyana.

2.2 MENISPERMOIDEAE ▸ TRIBE CISSAMPELIDAE (5/c 130) - outsiders are Stephania (c 70; tropical regions in the Old World), Perichasma (2; Central Africa, Angola), Antizoma (3; arid regions in S Africa), Cyclea (32; China to Philippines).

11. Cissampelos L. Climbers or rarely erect shrubs or perennial herbs (rare in Menispermaceae) with near peltate leaves, sometimes with xylopodium. c. 20 spp., 12 in Central and South America (11), Africa and Asia; 9 in Brazil, C. sympodialis Eichler endemic.

2.3 MENISPERMOIDEAE ▸ TRIBE PACHYGONEAE (4/45) - outsiders are Haematocarpus (2; Himalayas, SE Asia to Sulawesi), Cocculus (9; tropical and S Africa, Madagascar, Socotra, tropical and subtropical Asia to N Australia, subtropical to temperate North America), Pachygone (12; S China, SE Asia, Malesia to islands in W Pacific).

12. Hyperbaena Miers ex. Benth. Climbers or erect shrubs or trees with simple, entire or serrate leaves. 22 spp. in Central and South America (only 4); 3 spp. in Brazil, none endemics.

2.4 MENISPERMOIDEAE ▸ TRIBE TILIACOREAE (16/111) - outsiders are Syrrheonema (3; tropical W and C Africa), Carronia (4; New Guinea, E Queensland, E New South Wales), Pycnarrhena (9; SE Asia, Malesia to tropical Australia), Beirnaertia (1; tropical Africa), Triclisia (15; tropical Africa, Madagascar), Albertisia (19; tropical and subtropical Africa), Anisocycla (5; tropical Africa, Madagascar), Tiliacora (22; tropical regions in the Old World), Eleutharrhena (1; Assam, Yunnan), Macrococculus (1; New Guinea), Pleogyne (1; E Queensland), Synclisia (1; C Africa).

13. Chondrodendron Ruíz & Pavón. Climbers with often slighly peltate leaves; flowers in cauliflorous fascicles. Three spp., C. tomentosum Ruiz & Pav. from Panamá to Ecuador, Bolivia and Brazil, and two endemics to Brazil.

14. Curarea Barneby & Krukoff. Climbers with simple leaves; flowers in panicles to pseudoracemes. 9 spp., Central America to Bolivia (all in South America), Brazil (5, C. crassa Barneby, a large liana, rare, endemic to coastal Bahia) and Guianas.

15. Sciadotaenia Miers. Climbers with simple leaves; flowers in solitary or in simple flowered cymes. 20 spp. in South America, only S. nitida (L. Riley) Krukoff & Barneby into Central America; 15 spp. in Brazil, 7 endemics.

16. Ungulipetalum Mold. Herbaceous climbers with simple leaves; flowers in cymes. Only one sp., U. filipendulum (Mart) Mold., in SE Brazil, in Rio de Janeiro and São Paulo states.

RANUNCULACEAE

§ FULLY CHLOROTROPHIC

Genera/species 51/2,000-2,200 Distribution cosmopolitan except Antarctica, with their largest diversity in temperate and polar regions in the Northern and Southern Hemispheres. Habit usually bisexual (rarely dioecious), usually perennial, biennial or annual herbs (in Xanthorrhiza suffrutices; in Clematis lianas); main root often early withering and replaced by adventitious roots; some species with tuberous roots; roots and rhizomes in Xanthorhiza and Coptis and bark in Xanthorhiza intensively yellow-coloured due to presence of berberine (a major alkaloid).

Although Ranunculaceae species are distributed worldwide. its members are most common in the temperate and cold areas of the northern hemisphere. The diversity is reduced in the tropics: of nearly 62 genera in total, only 11-13 are native to the Neotropics, with 90-100 species in total. 11-13 native genera. 101 spp. in South America.

SYSTEMATIC five subrfamilies, Glaucidioideae (1/1, Japan), Hydrastioideae (1/1, Central and E North America) and Coptoideae (2/10–15, temperate regions on the N Hemisphere) do not occur in South America.

1. SUBFAMILY THALICTROIDEAE (7–8/280–340) ‣ outsiders are Leptopyrum (1; W Siberia to E Asia), Paraquilegia (4; W Iran to Himalayas and W China), Urophysa (2; China), Aquilegia (100–110; temperate regions on the Northern Hemisphere), Dichocarpum (18; Himalayas, E Asia), Enemion (6–7; NE Asia, SW Canada, W U.S.A.), Isopyrum (3; Central Europe, temperate Asia).

1. Thalictrum All. Perennials, usually with sympodial rhizome; inflorescence variously format; flowers bisexual or unisexual. 10 spp., Mexico through Central America and the Andes to southern Argentina (7 in South America, from Venezuela to Cono Sur, slightly diverse in Peru, absent in Chile and Brazil), also in Africa, S Europe and W China.

2. SUBFAMILY RANUNCULOIDEAE (c 35/1.440–1.540) ‣ two tribes, both in South America.

2.1 RANUNCULOIDEAE ▸ TRIBE ANEMONEAE (3/460-520) - all genera in South America.

2. Anemonastrum Holub. (off Anemone) Small herbs. 14 spp., mainly northern hemisphere, but one in New Zealand and A. antucense (Poepp.) Mosyakin & de Lange from N Chile and Argentina.

3. Anemone L. (inc. Berneoudia, Oreithales, Knowltonia, exc. Anemonastrum) Rosettes of basal leaves with a variety of perennating structures, inflorescences with involucral leaves on the peduncle, a perianth composed of petaloid sepals of variable number, and achenes. 114 spp., almost cosmopolitan; with 15 spp. in South America, mainly in temperate areas and zones of high elevation in Ecuador (1), Peru (3), Bolivia (1), Chile (9), Argentina (5), Brazil (3, two endemics), Paraguay and Uruguay; Brazilian A. sellowii Pritz. and A. assisbrasilica Kuhlm. & Porto composed a endemic subsection Sellowi of subgenus Anemone, in the coastal mountains of southeast country.

4. Clematis L. Shrubs, half-shrubs, sometimes perennial with woody base; stems scadent, sometimes erect; inflorescence terminal or axillary, trichotomously compound principally, sometimes single flowered. 295 spp., c. 26 in New World, 15 spp. in South America mainly in Colombia (2), Ecuador (4), Peru (8), Bolivia (7), Argentina (5) and Brazil (5, C. ulbrichiana Pilg. endemic), mostly temperate and subtropical.

2.2 RANUNCULOIDEAE ▸ TRIBE RANUNCULEAE (c. 32/c. 1,000) - outsiders are Calathodes (4; Himalayas, China inc. Taiwan), Adonis (c 30; Europe, temperate Asia), Trollius (c 30; temperate regions on the Northern Hemisphere), Callianthemum (14–15; Central Europe to Central Asia), Helleborus (c 20; W, C and S Europe, Mediterranean to the Caucasus and northern Syria, Tibet and W China); Delphinium (300–320; temperate regions on the Northern Hemisphere), Staphisagria (3; Mediterranean), Aconitum (250; temperate regions on the Northern Hemisphere), Gymnaconitum (1; Tibet, W China), Nigella (18; Europe, Mediterranean, temperate Asia); Actaea (29; temperate regions on the Northern Hemisphere), Anemonopsis (1; Honshu in Japan), Beesia (2; W and SW China, N Burma), Asteropyrum (2; China), Eranthis (8–9; Europe, temperate Asia), Kumlienia (1; California), Peltocalathos (1; S Africa), Beckwithia (3; temperate and polar regions on the Northern Hemisphere), Oxygraphis (4–5; temperate Asia), Arcteranthis (1; Alaska, NW Canada), Trautvetteria (1; Japan, Sakhalin, SW Canada, W and E U.S.A.), Coptidium (2; cold-temperate and arctic regions on the Northern Hemisphere), Ficaria (4; Europe, Mediterranean to Central Asia), Ceratocephala (5; nearly cosmopolitan), Paroxygraphis (1; E Himalayas).

5. Callianthemoides Tamura. Perenials, stems subscapose; leaves semibasal, 3-7 times pinnately ternate, flowers 1-2, terminal; petals 10-20, white. Only one sp., C. semiverticillata (Phil.) Tamura, Chile and Argentina.

6. Caltha L. Perennial herbs; stems simple or sparsely branched in the upper part; basal and cauline leaves; flowers in corymbiform cyme. 12 spp., temperate regions on both hemispheres; three spp. of South America, in the Andes to Ecuador from Tierra del Fuego and Falkland Islands, also in SE Australia, Tasmania, New Zealand.

7. Halerpestes Greene. (off Ranunculus) Small herbs. 9 spp., 6 from SE. European Russia to temp. Asia, H. cymbalaria (Pursh) Greene from Alaska to Tierra del Fuego and Uruguay (absent in Brazil), and two restricteds for Peru to Cono Sur.

8. Hamadryas Comm. ex Juss. Dioecious perennial with thick rhizomes; stems scapose, or sparsely branched with bracts; flowers bisexual, in loose cymes. 5 spp., only in Chile and Argentina.

9. Laccopetalum Ulbr. Perennials with short, thick rhizome; stem simple; flowers terminal, bisexual, subglobose. Only one sp., L. giganteum (Wedd.) Ulbr., in alpine zone of Andean Peru.

10. Myosurus L. Small, scapose annual with a persistent, elongate hypocotyl; flowers terminal, bisexual. 15 spp. in Laurasia, Mexico, two in Peru to Argentina and Chile, one of then also in North America.

11. Ranunculus L. (exc. Halerpestes) Perennial or annual; stem branched or simple, sometimes cushions. 1,690 spp., almost cosmopolitan; in New World 125 spp. in areas with high humidity, 53 in South America, mainly in Colombia (12), Ecuador (12), Peru (33), Bolivia (18), Argentina (26), Chile (19); 3 spp. in Brazil, R. apiifolius Pers., R. bonariensis Poir. and R. flagelliformis Sm., none endemics.

22. PROTEALES

FAMILIES ABSENTS IN SOUTH AMERICA: PLATANACEAE (1/10).

SABIACEAE

§ FULLY CHLOROTROPHIC

Genera/species 3/162 Distribution Himalaya, southern, E and SE Asia, Malesia, New Guinea, New Britain, Solomon Islands, tropical America northwards to central Mexico (Ophiocaryon in tropical South America). Habit bisexual or polygamodioecious, usually evergreen (two species of Meliosma deciduous) trees, shrubs or lianas (in Sabia japonica with short spines).

Meliosma can be confused with Sapotaceae, but it lacks milky latex; in the Neotropics only Meliosma alba (Schltdl.) Walp. has compound leaves; Ophiocaryon could be confused with Sapindaceae but differs in having a well developed terminal leaflet, in contrast with the rudimentary terminal leaflet in Sapindaceae.

Key to the genera of Neotropical Sabiaceae

1. Pinnately compound leaves; two stigmas ------------ Ophiocaryon

1. Simple, very rarely pinnately compound leaves; one stigma ------------ Meliosma

SYSTEMATIC outsider Sabia (c 30; SE Asia, Malesia).

1. Meliosma Bl. Trees, evergreens or dioecious. 126 spp., disjunct between SE Asia (40) and tropical America (85), mostly south of Mexico and Central America (including Atlantic coast) and tropical Andes; 51 spp. in South America, 4 spp. in Brazil, three confined to Atlantic Forest of E Brazil (M. itatiaiae Urban a rare plant in Brazil, by Plantas Raras do Brasil’s book, known only in Mantiqueira Range, on the border between Rio de Janeiro, Minas Gerais, and São Paulo states) and M. herbertii Rolfe in Brazilian Amazon rainforest and over northern South America; all Neotropical species are included in section Lorenzanea, except M. alba (Schltdl.) Walp., disjunct in Mexico and Asia.

2. Ophiocaryon R. H. Schomb. ex. Endl. Evergreen small trees or shrubs. 10 spp. restricted to the rainforests of northern South America, and one sp. in Guiana Shield; mainly along rivers; six spp. in Brazil, none endemics.

NELUMBONACEAE

§ FULLY CHLOROTROPHIC

Genera/species 1/2 Distribution Nelumbo nucifera: E, S and SE Asia, Malesia southeastwards to northern Australia; Nelumbo lutea: E North America southwards to the Great Antilles and Colombia. Habit bisexual, perennial herbs. Aquatic. Rhizome rich in starch. Internodes near growth points forming fleshy banana-shaped nutrient-storing tubers.

Nelumbonaceae used to be associated with Nymphaeaceae, the two having superficially similar flowers and vegetative parts; the differ in habit, as Nelumbo has leaves and flowers elevated above the water surface by stalks, while Nymphaeaceae leaves and flowers float on the surface.

SYSTEMATIC a single genus.

1. Nelumbo Adans. Herbaceous aquatic perennials producing latex with adventitious roots; leaves in groups of three along stem, petiole to 2m in length; leaf emergent or floating, simple, large, 10-100 cm across, bluish green adaxially and extremely water repellent; flowers solitary, actinomorphic, showy (10 or more cm diam.), pink to white or yellowish, elevated above water on terete peduncles up to 2 m in length; petals ca. 20-30; fruits globose or elongate ovoid; seeds exalbuminose. Two spp. worldwide, N. nucifera Gaert. native to S Russia, Asia, India and Australia, and N. lutea Willd. from Canada to Honduras, disjunct Colombia, Jamaica and Cuba.

PROTEACEAE

§ FULLY CHLOROTROPHIC

Genera/species 76/c. 1,700 Distribution South and Central America, Africa south of Sahara, Madagascar, southern India and Sri Lanka, and eastwards to E China (inc. Taiwan), southern Japan, Indochina, Malesia, islands in the SW Pacific, Australia and Tasmania; with their largest diversity in Mediterranean climates of Australia and South Africa. Habit usually bisexual (sometimes monoecious, andromonoecious or dioecious), evergreen trees or usually shrubs (rarely perennial rhizomatous herbs, some species with lignotuber). Most species are xerophytic.

Eight genera and 98 species occur in the Americas distributed from Mexico to Chile and Argentina of which five occur in the tropical region, with the greatest diversity is in the Andes and E Brazil, and belongn Grevilleoideae subfamily (45/855. Australia, Tasmania, SE Pacific to SE Asia, India and Sri Lanka, South America, South Africa, and Madagascar).

KEY TO GENERA OF NEOTROPICAL PROTEACEAE

1. Adult leaves pinnate - 2

2. Fruit a follicle, seeds winged; style erect ------------ Roupala

2. Fruit indehiscent, seeds not winged; style curved ------------ Euplassa

1. Adult leaves entire, simple or pinnatifid, never pinnate - 3

3. Fruit indehiscent with thick hard or fleshy pericarp, seeds not winged ------------ Panopsis

3. Fruit a dehiscent follicle, the pericarp thin not fleshy, seeds winged - 4

4. Hypogynous glands 3 lobed, broad, truncate; ovules many; flowers zygomorphic ------------ Oreocallis

4. Hypogynous glands 4; ovules 2; flowers actinomorphic or only weakly diagnonally zygomorphic - 5

5. Ovules ascending; young inflorescences conical due to overlapping bracts subtending flower pairs; seed lateral to wing ------------ Orites

5. Ovules pendulous; young inflorescences not conical; bracts subtending flower-pairs small; seed central to wing ------------ Roupala

SYSTEMATIC five lineages, Bellendenoideae (1/1, Tasmania), Persoonioideae (2/110, Australia, New Caledonia, New Zealand), Symphionematoideae (2/3, SE Australia, Tasmania) and Proteoideae (25/640, Africa south of Sahara (with their highest diversity in the Cape region), Madagascar, Australia) lineages do not occur in South America; subfamily Grevilleoideae (45/855), the only in South America, has four lineages, tribe Banksieae (3/175, N, SW and E Australia, Tasmania) does not occur in South America

1.1 PROTEOIDEAE ▸ TRIBE MACADAMIEAE (15/92) ‣ outsiders Macadamia (9; Madagascar, Sulawesi, Queensland, New South Wales, New Caledonia), Brabeium (1; W Cape), Malagasia (1; Madagascar), Catalepidia (1; Queensland), Virotia (6; New Caledonia), Athertonia (1; Queensland), Heliciopsis (14; China, Burma, Thailand, Indochina, Malesia to Philippines), Cardwellia (1; Queensland), Sleumerodendron (1; New Caledonia), Bleasdalea (5; New Guinea, Queensland), Hicksbeachia (2; Queensland, New South Wales), Kermadecia (4; New Caledonia).

1. Euplassa Salisb. Trees, less frequently shrubs; leaves spirally arranged, paripinnate; inflorescence generally unbranched, pseudo-racemose, axillary or rarely terminal, solitary, occasionally with 2 inflorescences per leaf axil; flowers in pairs, sessile or pedicellate, weakly zygomorphic, each pair subtended by a small common bract; tepals 4; fruit a nut, rarely a drupe, 1-2-seeded. 21 spp. confined to South America, 14 in Brazil (11 endemics), 6 spp. endemic to Venezuela to Peru, with E. inaequalis (Pohl) Engl. widely distributed in South America.

2. Panopsis Salis. Shrubs or trees; conflorescence terminal or lateral, a raceme of flower pairs, or a panicle of such racemes. 27 spp., widely distributed in Central and South America (24), sparsely in Brazil (5, two endemics).

3. Gevuina Molina. Trees; adult leaves imparipinnate to tripinnate, often with roots crown; conflorescence lateral, a raceme of flower pairs. Only one sp., G. avellana Molina, rainforests and disturbed sites, Chile and Argentina.

1.2 PROTEOIDEAE ▸ TRIBE ROUPALEAE (13/177) ‣ outsiders Megahertzia (1; Queensland), Knightia (1; New Zealand), Eucarpha (2; New Caledonia), Triunia (4; Queensland, New South Wales), Neorites (1; Queensland, New South Wales), Lambertia (10; W Australia, New South Wales), Xylomelum (7; Australia), Helicia (c 100; South, E and SE Asia, Malesia to New Guinea, Australia), Hollandaea (4; Queensland), Darlingia (2; Queensland), Floydia (1; Queensland, New South Wales).

4. Orites R.Br. Shrubs or trees; adult leaves simple; conflorescence terminal or lateral, a raceme of flower pairs, or a panicle of such racemes; flower pair subtended by an ovate to linear, caduceus bract. 9 spp., 7 endemics to Australia and Tasmania and two in South America, Chile and Bolivia one endemic each.

5. Roupala Aubl. Shrubs or trees up to 30 m tall, often with roots crown; conflorescence lateral; flower pair subtended by a scale-like, sometimes caducous bract; flowers pedicellate or sessile. 42 spp., Mexico to Argentina, slyghtly centered in northern Andes, only 15 in Brazil (11 endemics, R. sculpta Sleumer is a rare plant, known only in São Paulo municipality, by Plantas Raras do Brasil’s book); the widely distributed R. montana Aubl. is a excelent woody, and known as ‘Brazilian oak’, and occur in over Neotropics, in Brazil except coast of NE region.

1.3 PROTEOIDEAE ▸ TRIBE EMBOTHRIEAE (12/560) ‣ outsiders Alloxylon (4; New Guinea, Aru Islands, Queensland, New South Wales), Telopea (5; New South Wales, Victoria, Tasmania), Stenocarpus (21; New Guinea, Aru, Australia, New Caledonia), Strangea (3; W Australia, Queensland, New South Wales), Opisthiolepis (1; Queensland), Buckinghamia (2; Queensland), Grevillea (c 360; Malesia to New Guinea, Australia, New Caledonia), Hakea (c 150; Australia), Finschia (3; New Guinea, Bismarck Archipelago, Solomon Islands, Aru, Palau, Vanuatu).

6. Embothryum J. R. Forst & G. Forst. Shrubs or trees; leaves simple, entire. Only one sp., E. coccineum R. Forst. & G. Forst., alpine shrublands to rainforests of Argentina and Chile.

7. Oreocallis R.Br. Shrubs or trees; leaves simple, entire; conflorescence terminal or lateral, not subtended by an involucre of bracts. Two spp., in Andes of Ecuador and Peru, in montane shrublands to forests.

8. Lomatia R.Br. Shrubs or trees; leaves in many formats. 12 spp., nine in shrublands to rainforests, E Australia including Tasmania, and 3 spp. in Chile, Argentina, one reaching into Peru and Ecuador, in montane shrublands to forests.

23. TROCHODENDRALES

TROCHODENDRALES DOES NOT OCCUR IN SOUTH AMERICA, AND IS COMPOSED OF A SINGLE FAMILLY, TROCHODENDRACEAE (2/2).

24. BUXALES

A SINGLE FAMILY, PRESENT IN SOUTH AMERICA.

BUXACEAE

Genera/species 6/122 Distribution tropical and S Africa, Madagascar, Socotra, NE Africa, Macaronesia, W and C Mediterranean, E Türkiye, Caucasus and N Iran, Afghanistan, Himalaya, S India, Sri Lanka, E Asia to Korean Peninsula and Japan, SE Asia, W and C Malesia, southern North America, Central America, NW South America (Andes). Habit usually monoecious (rarely dioecious, polygamodioecious or bisexual), evergreen trees or shrubs; Pachysandra consists of perennial herbs with a rhizome. Use Ornamental plants (Buxus sempervirens, Pachysandra terminalis etc.), timber, carpentry, engravings (Buxus).

Hexaporotricolpites pollen, from the late Albian of Gabon and Brazil, is similar to recent buxaceous pollen and is even more similar to pollen of the sister family Didymelaceae, which is endemic to Madagascar. Buxus is the only genus of the family that has an almost worldwide distribution. It occurs in Africa (including Madagascar) and Central America and in the Caribbean region, Mediterranean region, and Asia. The other smaller genera are geographically restricted, Sarcococca to E Asia, Pachysandra to E Asia and E North America, and Styloceras to South America. Sarcococca is a mainly E Asian genus with 1 species in Guatemala and Mexico, still of doubtful generic affiliation.

SYSTEMATIC two subfamilies, both in South America.

Key to genera of Neotropical Buxaceae

1. Tepals absent, stamens numerous ------------ Styloceras

1. Tepals present, stamens usually four - 2

2. Leaves decussate, fruit a 3-valved capsule ------------ Buxus

2. Leaves alternate, fruit subdrupaceous ------------ Sarcococca

1. SUBFAMILY BUXEAE (1/101) ‣ a single genus.

1. Buxus L. Shrubs or small trees, tetragonal branchlets, leaves decussate; flowers in lax to glomerate racemes with a terminal female flower; staminate 4-merous tepals and stamen; tepals decussate; stamens opposite tepals, inserted around a pistillode; fruit a 3-horned capsule; dehiscing loculicidally into 3 spreading 2-horned valves. 101 spp., widely distributed on all continents except Australia; 51 spp. in New World, with more than 30 spp., Cuba is main centre of diversity; following by Jamaica with only 4 -5 spp., Porto Rico with two, and Martinique and the Bahamas, each with one sp.; in Central America and Mexico are 4 spp.; and in S. America only B. citrifolia (Willd.) Spreng., with the few reports of the species in Panama indicate that it is uncommon and restricted to a narrow range, occurring in the Canal area and in the provinces of Panamá and Colón; in Colombia it is known from Cartagena and Sucre, where it appears to be rare; one larger Venezuelan population is recorded; records also in Suriname.

2. SUBFAMILY STYLOCERATEAE (4/21) ‣ outsiders Haptanthus (1; Matarras in Honduras); Sarcococca (c 15; tropical and subtropical Asia from Afghanistan and the Himalayas to China, SE Asia and Malesia including Philippines, Mexico and Guatemala), Pachysandra (5; India, Nepal, Burma, China (inc. Taiwan), Japan, SE U.S.A.).

2. Styloceras Kunth. ex. Juss. Trees or shrubs, dioecious, rarely monoecious; staminate flowers in short pendent spikes; tepals absent; several stamens inserted on an angular-ovate bract; pistillode absent; fruit globose, yellow, dupraceous, slightly fleshy, indehiscent or tardily dehiscent. 6 spp., only South America. S. penninervium A. Gentry & G. Aymard known only from the eastern slope of the C Peruvian Andes 1,800 – 1,850 m; S. brokawii A. Gentry & R. Foster. in lowland N Bolivia, S Peru and Acre state in western Brazil; the first non Andean spp. of this genus discovery; S. kunthianum Jussieu endemic to Ecuador, in Upper Pastaza valley; S. columnare Miiller Arg. in Sorato area in Bolivia and Peru; S. connatum Torrez & P. Jørg., known only in the cloud forests of Madidi National Park and Pilón Lajas Biosphere Reserve and Communal Lands in the Bolivian Andes; and S. laurifolium (Willdenow) HBK in Venezuela to S Peru, above 2,200 m.

25. GUNNERALES

FAMILIES ABSENTS IN SOUTH AMERICA: MYROTHAMNACEAE (1/2).

GUNNERACEAE

§ FULLY CHLOROTROPHIC

Gener/species 1/63 Distribution E and South Africa, Madagascar, Malesia, New Guinea, Solomon Islands, Tasmania, New Zealand, Hawaii, Central America, South America (above all in the Andes), Juan Fernandez, Falkland Islands. Habit perennial herbs, either with ascending or creeping pachycaulous stems, covered with large leaf scars, apically with large to gigantic, long-petioled leaves reaching up to c. 5m in height (G. magnifica H. St. John), and between these often covered with conspicuous bracts protecting the inflorescence and vegetative buds, or stoloniferous and mat-forming, with short, upright stem portions bearing leaf-rosettes, reaching from 4 cm to about 1 m in height, or in one case (G. herteri Osten), diminutive annuals.

The massive inflorescence of small, reddish flowers is up to 2.3 m (7 ft 6 inches) long and weighs about 13 kg. Several small species are found in New Zealand, notably G. albocarpa (Kirk) Cockayne, with leaves only 1–2 cm long, and also in South America, with G. magellanica Lam. having leaves 5–9 cm wide on stalks 8–15 cm long.; some species commonly known as ‘giant rhubarb’; these organs are covered by leaf-like scales. Among tracheophytes, the eudicot Gunnera L. is the only known angiosperm that harbors a cyanobacterial symbiont namely Nostoc.

SYSTEMATIC a single genus

1. Gunnera L. Caracters of family. 56-57 spp., subtropical or montane genus, of which most occur in South (all countries except Guianas and Paraguay) and Central America; 5-6 spp. inhabit New Zealand, while SE Asia, Tasmania and Africa have a single endemic species each; two species grow on Hawai; 4 occur in Mexico and Central America, and 42 in South America; six sections: Gunnera (1, Africa and Madagascar), Pseudogunnera (1, SE Asia to New Guinea), Milligania (6, one in Tasmania and remaining in New Zealand) and the three of New World:

§ sect. Ostenigunnera ‣ only G. herteri Osten, from S Brazil and E Uruguay, growing near coastline, in sand soils; is only glabrous species, unique annual specis in family; this species seems to be sister to the remaining species of genus.

§ sect. Panke ‣ 43 spp. neotropical (mainly in Andes, e.g. 25 in Colombia) and two spp. in Hawai, only G. manicata Linden ex André in Brazil, native to the Serra do Mar mountains of SE & S Brazil, is perhaps the largest species, with leaves typically 1.5-2 m (5-6 ft) wide, but exceptionally long, up to 3.4 m (11 ft), borne on thick, succulent leaf stalks (petioles) up to 2.5 m (8 ft) long.; all species of this section are similar in morphology, being gigantic herbs with stout and erect rhizomes surrounded by palmately lobed leaves that sometimes reach up to several meters in diameter; the inflorescences of Panke are compound spikes with many small, dimerous and wind pollinated flowers and the fruits are drupes, which are fleshy and brightly colored in some species while greenish and dry in others.

§ sect. Misandra ‣ two spp., G. magellanica Lam. and G. lobata Hooker f., from Colombia to Tierra de Fuego.

26. DILLENIALES

A SINGLE FAMILY, PRESENT IN SOUTH AMERICA.

DILLENIACEAE

§ FULLY CHLOROTROPHIC

Genera/species 11/c 430 Distribution almost pantropical, with the largest diversity in tropical Asia and Australia; Hibbertia also in warm-temperate parts of Australia including Tasmania. In South America the high diversity occurs in Amazon rainforest. Habit usually bisexual (rarely dioecious), usually evergreen (rarely deciduous) trees or shrubs (rarely suffrutices; Tetracera consists of lianas; Acrotrema costatum Jack (possibly nested in Dillenia) is a perennial herb with woody rhizome). Bark often intensely brown. Lianas, shrubs, small trees with tortuous branches (Curatella), or evergreen trees up to 30 m (Dillenia); lianas generally with stems >5 cm diameter, the vascular bundles disposed in bands or concentric rings separated by abundant parenchyma (papyraceous bark, often asperous: Doliocarpus, Neodillenia, Pinzona, several spp. of Davilla).

Key to genera of Neotropical Dilleniaceae

1. Erect shrubs to trees up to 15 m tall ------------ Curatella

1. Scandent to creeping climbers or lianas, or else small erect shrubby, generally < 3 m tall - 3

3. Carpel 1 per flower - 4

4. Sepals unequal in size, the 2 inner ones larger, covering the fruit entirely ------------ Davilla

4. Sepals ± equal in size, never covering the fruit - 5

5. Stamens free, not forming a ring around the carpel, aril white ------------ Doliocarpus

5. Stamens connate at the base, forming a ring around the carpel, aril red ------------ Neodillenia

3. Carpels 2-5 per flower - 6

6. Carpels 3-5 per flower; fruits follicles or berries - 7

8. Carpels free; aril white ------------ Davilla

8. Carpels connate ventrally from the base to the apex of the ovary; aril orange ------------ Pinzona

6. Carpels 2 per flower; fruits capsules - 8

7. Inflorescence paniculate; fruits follicles, the aril fimbriate or deeply laciniate ------------ Tetracera

7. Inflorescence racemose or flowers solitary; fruits berries, the aril entire ------------ Neodillenia

SYSTEMATICS two subfamilies, Hibbertieae (1/170–180, Madagascar, Malesia to New Guinea, Australia, New Caledonia, Fiji, with their highest diversity in Australia) and Dillenieae DC. (4/c 80, Tropical Asia to tropical Australia) are only Old World.

1. SUBFAMILY DELIMEAE (1/c 50) ‣ a single genus.

1. Tetracera L. Lianas or scandent shrubs, rarely lignotuberous subshrubs, inflorescences paniculate. 47 spp., pantropical, 21 in New World, 15 in Brazil (7 endemics); unique species of this family in Argentina.

2. SUBFAMILY DOLIOCARPEAE (5/c 75) ‣ all genera occur in South America.

2. Curatella Loefl. A small to medium-sized tree. Only one sp., C. americana L., from Mexico to Brazil; common in savannas of C Brazil (cerrado), known as ‘lixeira’.

3. Davilla Vand. Scandent shrubs or lianas; leaves sometimes with amplexicaul petiolar wings. 29 spp. of lianas and erect shrubs, from E Mexico to Paraguay and Brazil, with center of diversity in NE Brazil; 28 spp. in Brazil, 22 endemics; D. bilobata (not recognized in VPA) from Bahia is the unique species in this genus with lobulate petals; D. glaziovii Eichler from São Paulo, D. sellowiana from Rio de Janeiro (not recognized in VPA) and D. sessiliflora Fraga from Bahia state are rare plants in Brazil, by Plantas Raras do Brasil’s book.

4. Doliocarpus Rolander. Shrubs, mostly scandent, or lianas; unique in family with ramiflorous, fasciculate or glomerate inflorescences, an unicarpellate, one-celled ovary, a berry as a fruit, sometimes opening irregularly, and seeds completely covered by a white aril. 59 spp., from Mexico to Paraguay, 57 in South America, 37 spp. in Brazil, 18 endemics; D. prancei Kubitzki, from near Manaus in Amazonas state, is a rare plant in Brazil, by Plantas Raras do Brasil’s book.

5. Neodillenia Aymard. Lianas with successive cambia; trichomes simple; inflorescences axillary (and sometimes also ramiflorous). Three spp., one endemic to Venezuela, N. coussapoana Aymard and N. peruviana Aymard from Colombia, the latter also in Ecuador, and the former also in Peru and Amazonas state in Brazil.

6. Pinzona Mart. & Zucc. A high-climbing liana, largely glabrous at maturity except for inflorescence axes. Only one sp., P. coriacea Mart. & Zucc., throughout Central and South America from Belize to NE Brazil and Antilles.

27. SAXIFRAGALES

FAMILIES ABSENTS IN SOUTH AMERICA: ALTINGIACEAE (1/15), APHANOPETALACEAE (1/2), CERCIDIPHYLLACEAE (1/2), CYNOMORIACEAE (1/1), DAPHNIPHYLLACEAE (1/30), ITEACEAE (2/24), PAEONIACEAE (1/36), PENTHORACEAE (1/2), TETRACARPAEACEAE (1/1).

LINEAGE 1 of 4: PERIDISCACEAE

PERIDISCACEAE

§ FULLY CHLOROTROPHIC

Genera/species 4/11 Distribution tropical South America, tropical W Africa. Habit. bisexual, evergreen trees or shrubs.

Trees, deciduous; leaves alternate, simple, lamina leathery, margins entire, tri-nerved, large pit on the underside in each axil of basal lateral veins; pulvinate at both ends of petiole; stipules present, intrapetiolar, caducous (leaving prominent scars); inflorescence axillary racemes (Peridiscus) or fascicles (Whittonia), bracteoles large and persistent; flowers regularly symmetrical, bisexual; sepals 4-5(-6) (Peridiscus) or 7 (Whittonia), imbricate, reflexed when open; apetalous; stamens numerous; ovary superior, syncarpous, half immersed in disc and glabrous (Peridiscus) or woolly and on top of disc (Whittonia); carpels 3-4; styles 3(-4), free, ovules 6-8, pendulous; fruit drupaceous (unknown in Whittonia). Seed 1 per fruit.

SYSTEMATIC outsider Soyauxia (7–9; tropical W and C Africa).

1. Peridiscus Benth. Trees, dioecious, glabrous except for the stipules and inflorescences (axillare racemes); flowers pale green to yellow or yellowish; apetalous. Only one sp., P. lucidus Benth, from south Venezuela, W Guyana, Amazonas and Amapá states in N Brazil; and in Guainia region in E Colombia; in Brazil this species occurs mainly in Upper Rio Negro to near Manaus.

2. Whittonia Sandwith. Small dioecious trees with long, golden brownish hairs on toung shoots; fascicles; apetalous. Only one sp., W. guianensis Sandwith, endemic to the Guiana Shield of Guyana, is possibly extinct, being known from only one specimen collected below Kaieteur Falls in Guyana, at 50 – 100 m elevation range; an attempt to rediscover it in 2006 was not successful.

LINEAGE 2 of 4: HAMAMELIDS

HAMAMELIDACEAE

§ FULLY CHLOROTROPHIC

Genera/species 26/95–115 Distribution E and SE Africa, Madagascar, southern Türkiye, SE Trans-Caucasus, N Iran, W and E Himalaya, Assam, Manipure, E and SE Asia to Korean Peninsula and Japan, Malesia, New Guinea, NE Australia (Queensland), E North America, Central America, NW South America. Habit usually bisexual (sometimes monoecious, andromonoecious or polygamomonoecious), evergreen or deciduous trees or shrubs. Use Ornamental plants, medicinal plants, timber.

SYSTEMATIC tribes Exbucklandioideae (3/10, Assam to SE Asia and S China, W Malesia) and Disanthoideae (1/1; Japan) do not occur in South America; among Hamamelidoideae, outsiders are Chunia (1; Hainan), Corylopsis (c 25; E Himalayas, China, Korean Peninsula, Japan), Dicoryphe (13; Madagascar, the Comoros), Distyliopsis (6; Burma, Thailand, Indochina, Malesia, Taiwan (China)), Distylium (c 15; SE Asia, Malesia), Embolanthera (2; SE Asia, Philippines), Eustigma (3; S China, SE Asia), Fortunearia (1; China), Fothergilla (2–3; SE U.S.A.), Hamamelis (6; SE Canada, E U.S.A., Mexico; China; Japan), Loropetalum (3; N and NE India to China, Japan), Maingaya (1; Malay Peninsula), Molinadendron (3; Mexico, Central America), Neostrearia (1; Queensland), Noahdendron (1; NE Queensland), Ostrearia (1; Queensland), Parrotia (2; SW Caspian area; E China), Parrotiopsis (2; Himalayas), Sinowilsonia (1; W and C China), Sycopsis (2; Assam, China inc. Taiwan), Trichocladus (6; tropical and S Africa).

1. Matudaea Lund. Evergreen trees, multicellular glandular hairs in the blade margin of primary leaves, stipules and bracts; branches with 2 prophylls; leaves triplinerved; inflorescences condensed panicles or botryoids (each axis terminated by a flower); flowers bisexual; sepals absent; petals absent; 2 fused bracts subtend the individual flower; stamens 12-24, polyandrous; anthers opening by 2 valves; ovary superior; carpels with 1 ovule; large, decurrent stigma. Two spp., M. trinerva Lundell from Mexico to Central America, and M. colombiana G. Lozano-C endemic to Andean Colombia.

LINEAGE 3 of 4: CRASSULIDS

HALORAGACEAE

§ FULLY CHLOROTROPHIC

Genera/species 8/c. 145 Distribution cosmopolitan, largest diversity in the Southern Hemisphere, particularly in Australia. Habit usually monoecious or polygamomonoecious (rarely bisexual or dioecious), perennial or annual herbs, suffrutices or evergreen small shrubs (some Glischrocaryon consists of shrubs and small trees). Numerous species are aquatic, other representative are amphibious, hygrophytes or terrestrial mesophytes. Myriophyllum in the Northern Hemisphere produces turions (condensed reproductive and hibernating shoots). The main root in aquatic and amphibious species is replaced by adventitious roots (without root hairs), anchoring the plant to the substrate (adventitious roots in Haloragis inserted between the leaves).

Centre of diversity in Australia with c. 60% of species; the four South American genera grow in slow moving water or in marshy, littoral habitats; some species are found at greater than 3,000 m; only two fully extra-australian genera, both native to neotropics; 5 spp. in Brazil.

Key differences from similar families - the features listed are not present in the Haloragaceae:

ü Gunneraceae: petiolate (often long), zygomorphic flowers, fleshy drupaceous fruits or leathery dehiscent fruits.

ü Penthoraceae: superior ovary, many seeds.

ü Tertracarpaeaceae: superior ovary, free carpels, many seeds, endemic to Tasmania.

SYSTEMATIC outsiders Glischrocaryon (5; S W Australia, SW South Australia, New South Wales, Victoria), Haloragodendron (5–6; Western Australia, SE New South Wales, E Victoria, Tasmania); Meionectes (2; Western Australia, South Australia, S Victoria, Tasmania), Trihaloragis (1; Western Australia), Gonocarpus (c 40; SE Asia and Malesia to Japan, Australia and New Zealand).

1. Haloragis Forst. & G. Forst. Annual or perennial herbs or subshrubs from taproots or stolons, glabrous, scabrous or with simple hairs; stems ascending or creeping, some growing in water. 28 spp. confined almost entirely to Australia and New Zealand, with two spp. on S Pacific Islands eastward to Juan Fernandez Islands: H. massatierrana Skottsb. and H. massafuerana Skottsb., both absent in continental Chile, in a wide variety of terrestrial habitats.

2. Laurembergia Bergius. Perennial herbs with woody rhizomes, some helophytic; leaves subwhorled, opposite or alternate, not heterophyllous; flowers unisexual in dichasia, the distal positions occupied by male or hermaphroditic flowers which stand out on a long pedicel from the almost sessile female flowers; 8 stamens in 2 whorls with the outer fertile whorl alternating with the sepals; the ovary initially 4-loculate, becoming 1-4-loculate through the collapse of the columella; one-seeded fruit with variable structure. 4 spp., 3 exclusively paleotropical (excluding Australia) and the polymorphic L. tetrandra (Schott) Kanitz, amphi-atlantic (i.e., also Africa), in New World from Atlantic of Argentina to Colombia, mainly in Brazilian southern coast and absent in Guianas.

3. Myriophyllum L. Perennial, rarely annual, aquatic or littoral herbs, free floating or rhizomathous; whorled heterophyllous leaves; flowers frequently unisexual; sepals less than half the length of the petals (frequently absent); fruit schizocarpic, ornamented and splitting at maturity into 2-4 mericarps. 61 spp., almost cosmopolitan, with centres in Australia (36, 31 endemic), North America (13, 4 endemics), and India/Indo-China (10, 7 endemic), although absent from most of Africa, the Middle East and much of S Asia and NE South America; 4 spp. in South America, all widely distributeds os scattered, three in Brazil, none endemics.

4. Proserpinaca L. Submerged, emergent of seasonally terrestrial rhizomatous herbs; alternate heterophyllous leaves; inflorescence solitary or in dichasia of up to 3 flowers per axil; flowers hermaphroditic; fruit 3-seeded nutlet. Two spp., P. palustris L. in North America, Mexico, Central America, Caribbean, Colombia and Brazil, and P. pectinata Lam. from North America, Mexico, Caribbean and Central America.

CRASSULACEAE

§ FULLY CHLOROTROPHIC

Genera/species 19(own data)/1,400–1,410 Distribution mainly in dry and warm areas in the Southern and Northern Hemispheres, with their largest diversity in Mexico (325) and southern Africa (250). Habit usually bisexual (rarely unisexual), usually perennial (rarely annual or biennial) herbs, suffrutices or evergreen shrubs (rarely trees or epiphytes). Leaf succulents. Usually xerophytes. Rarely aquatic (i.a. Crassula aquatica (L.) Schönland). Many species produce adventitious roots from broken leaves; some species form bulbils along leaf margins or in inflorescences.

Crassulaceae is the 3ª largest group of succulent species worldwide, c. 1,400 succulents. Mostly plants of dry, rocky habitats, usually terrestrial but rarely epiphytic and very rarely aquatic. Although the distribution of Crassulaceae is nearly worldwide, most species are found in five centers of diversity: Mexico (ca. 330), Mediterranean basin (ca. 100), Macaronesia (ca. 63), S Africa (ca. 250) and E Asia (ca. 300). Species usually grow in arid to semi-arid rocky and mountainous environments. There are several species of Kalanchoe that are cultivated, some naturalized; they are best recognized by flowers with (4) fused petals forming a tube, and many species have lobed, crenate or serrate margined leaves; a number of species of Kalanchoe, formally in the genus Bryophyllum, have plantlets on the leaf margins.

SYSTEMATIC three subfamilies, Kalanchoideae (4/185–190, Africa, Madagascar, Arabian Peninsula) does not occur in South America.

1. SUBFAMILY CRASSULOIDEAE (1/211) ‣ Subcosmopolitan, mostly restricted to southern Africa except for a small group of aquatic Crassula species that are distributed worldwide.

1. Crassula L. Herbs or shrubs, annual or perennial, aquatic or terrestrial, not viviparous, 0.1-5 dm, glabrous or pubescent, typically with isostemonous flowers. 211 spp. worldwide, 13 spp. in New World, 12 of then in South America, 7 restricted of this continent, and three only Cono Sur/North America range; C. penduncularis (Sm) Meigen, unique Brazilian nayive species in this family, occurs only in Santa Catarina and Rio Grande do Sul states, Peru, Bolivia and Cono Sur.

Some highly reduced annual Crassula are morphologically aberrant: C. aphylla Schönland & Baker f. from South Africa forms leafless, globular shoots reaching maturity at about 3mm; it may represent the smallest succulent plant.

2. SUBFAMILY SEMPERVIVOIDEAE (18/970–1,000) ‣ 4 clades, Telephium Clade (9/58, temperate regions on the Northern Hemisphere, one genus and species in Madagascar), Aeonium clade (5/100, alpine and mountain areas on the Northern Hemisphere, with their highest diversity in Himalayas, Tibet and W China) and Semperviveae (3/35–40, Europe, Mediterranean, Morocco, W Asia) do not occur in South AmericaSedeae, the fouth group, has only Sedum L.; the highest base chromosome number known for any flowering plant (n = 270) belongs to Sedum suaveolens Kimnach endemic to Mexico.

2. Sedum L. (inc. Villadia, Echeveria) Annual to perennial, monocarpic to polycarpic herbs orsubshrubs, rarely monocarpic rosette plants; roots usuallyfibrous, less commonly tuberous or as taproots; stems usuallywith branched non-flowering shoots or sessile rosettes, rarelyrhizomatous. c. 755 spp., mostly in temperate and subtropical regions of North America, Europe, north Africa, Near Eastand Asia, a few species in Central and South America and Central to E Africa, 484 spp. in New World, mainly in Mexico. Two groups:

§ Leucosedum clade ‣ 200 spp., Sedeae p.p., inc. Pistorinia, Rosularia, Prometheum, Afrovivella, Sedella, Dudleya, Sedum p.p. (ca. 120).

§ Acre clade ‣ c. 550 spp., Sedeae p.p., Cremnophila, Echeveria, Graptopetalum, Lenophyllum, Pachyphytum, Thompsonella, Villadia, Sedum p.p. (ca. 345), includes all 45 spp. of in South America, from Venezuela to N Argentina, mainly in Peru (34, 31 endemics).

LINEAGE 4 of 4: SAXIFRAGIDS

GROSSULARIACEAE

§ FULLY CHLOROTROPHIC

Genera/species 1/193 Distribution temperate parts of Eurasia, Mediterranean, NE Africa, North and Central America, the Pacific coast of South America, the Andes southwards to Tierra del Fuego. Habit usually bisexual (sometimes dioecious), usually deciduous (rarely evergreen) shrubs, upright, creeping (often with subterranean stems) or occasionally somewhat climbing; branches differentiated into long shoots and short shoots. Leaves sometimes modified into spines.

SYSTEMATIC a single genus.

1. Ribes L. Shrubs, sometimes lianescent (only in South American species) and very rarely cushion-forming (only Peru), bark often exfoliating; plants usually glandular and/or pubescent; often 3-lobed, usually pubescent and/or glandular leaves, sometimes aromatic; inflorescences terminal, racemose, sometimes reduced to 1-2 flowers; fruit a berry crowned with persistent flower remnants; seeds with gelatinous exotesta and dense, brown/black endotesta. 193 spp. in eight subgenera, occurring mainly in the north-temperate zone, 115 spp. in New World, 51 in South America (all confined to continent), all in the dioecious subgenus Parilla, section which into southern Central America with a single species, in two, apparently unrelated, clades:

§ sect. Parilla ‣ ca. 10 spp. in temperate South America.

§ sect. Andina ‣ c. 42 spp., two in S Central America, 13 are restricted of N Andes in Venezuela, Colombia and Ecuador, and c. 27 Peru southwards up to Terra del Fuego; Peru has 24 spp. (20 endemics), more than twice as many species of this genus overall as any other single South American country, sometimes found growing in vertical rock faces; R. frankei Weigend & Breitkopf. is possibly the most aberrant species of Ribes - it is a small, cushion-forming dwarf-shrub to 30 cm tall, with erect inflorescences that are virtually immersed in the leafy cushions; it is the smallest species of Ribes known so far and has been collected on vertical rock faces in the high Andes of Pasco in central Peru.

Ribes is important for browsing by Andean animals; birds avidly devour its fruit and, due to its dense branching, it provides excellent nesting sites for birds; the fruits of South American species are apparently all edible, but usually insipid. R. cucullatum Hook. & Arn. reaches elevations of 4,700 m or more in the Cordillera Blanca and is one of the highest-growing woody plants in this area; many species are very narrowly endemic, some only known from the type collection, some on the brink of extinction, e.g., R. lehmannii Jancz. in Ecuador, R. contumazensis Weigend and R. ovalifolium Jancz. in Peru.

All species are native, and many species are very narrowly endemic; R. rubrum L. (red currant), R. nigrum L. (black currant) and various cultivars of subg. Grossularia (gooseberries) are occasionally cultivated in temperate South America and may be present as species.

SAXIFRAGACEAE

§ FULLY CHLOROTROPHIC

Genera/species 40/c. 670 Distribution mainly in northern temperate and polar areas; a few species in southern temperate regions, and on tropical mountains (in the Andes). Habit usually bisexual (rarely androdioecious), usually perennial herbs (sometimes annual or biennial, rarely somewhat lignified). Some species are succulent and/or xerophytic. Use ornamental plants, medicinal plants.

Vegetatively resembles the Gentianaceae, Apiaceae, Violaceae and other alpine plants found in the Andes, being distinguished from them by its corolla with five distinct petals, stamens 1 or 2 x the number of petals; it can be confused with the Crassulaceae, but these are succulent and have carpels totally distinct and generally present in larger number (4 or +).

SYSTEMATIC nine tribes, Micratheae (1/80, fully circumboreal, in North America extending south extensively in montane regions to northern Mexico and the Appalachians; in Eurasia extending south sporadically into southern Europe, Mongolia, Korea, and Japan), Dermereae (6/19; China to Korea, W U.S.A., Himalayas), Leptarrheneae (2/2; China, Japan, Alaska to U.S.A.), Heuchereae (15/83, Asia, North America), Astilbeae (2/10; temperate Asia to New Guinea, North America) and Saniculiphylleae (1/1; China) does not occur in South America.

TRIBE CASCADIEAE (2/2) ‣ outsider Cascadia (1; W U.S.A.).

1. Saxifragodes DM. More. Stems slender, branched at base. One poorly understood species, S. albowiana (Kurtz ex Albov) D.M. Moore, from center Chile to Tierra del Fuego, and Santa Cruz in Patagonia.

TRIBE CHRYSOSPLENIEAE (2/57) ‣ outsider Peltoboykinia (2; Japan).

2. Chrysosplenium L. Plants stoloniferous; leaves glabrous to pilose, small, opposite or alternate, petiolate, crenate, estipulate. 55 spp., in moist areas of temperate to arctic North America and Eurasia, mainly in Asia, two in South America, C. macranthum Hook. and C. valdivicum Hook., both in southern Chile and Argentina.

TRIBE BOYKINIEAE (8/19) ‣ outsiders Telesonix (2; Canada to New Mexico), Jepsonia (3; California to Baja California), Sullivantia (3; Rocky Mountains to Virginia), Boykinia (7; North America and Japan), Suksdorfia (1; W North America), Bolandra (2; W North America), Hemieva (1; W North America).

3. Hieronymusia Engler. Plants from a slender rhizome; leaves round cordate, shallowly lobed and serrate. One poorly known species, H. alchemilloides (Griseb.) Engl., Sierra de Tucuman in Argentina and in Bolivia on damp, shady humus and damp rock cliffs at 3,000–4,000 m.

TRIBE SAXIFRAGEAE (1-2/c. 370) ‣ both genera in South America.

4. Saxifragella Engl. Cushions. Only one sp., S. bicuspidata (Hook f.) Engl., Chile and Argentina; this species should be included within the large genus Saxifraga, which is consistent with the results phylogenetic works.

5. Saxifraga L. Perennials or more rarely delicate annuals or biennials, often cushions; 470 spp., widely distributed but primarily of temperate or Arctic regions of the N Hemisphere; many of the species are circumboreal; two spp. in South America, S. magellanica Poir. from southern South America reaching up to Peru and Ecuador, and S. boussingaultii Brongn. in Bolivia (possibly a mistake) and Cotopaxi region in Ecuador.

28. VITALES

A SINGLE FAMILY, PRESENT IN SOUTH AMERICA.

VITACEAE

§ FULLY CHLOROTROPHIC

Genera/species 18/920-960 Distribution mainly in tropical and subtropical regions, some species in warm-temperate areas; Leea: SE Asia, Malesia, New Guinea and Australia, two species in Africa and Madagascar. Habit bisexual, monoecious, polygamomonoecious, dioecious or polygamodioecious, usually evergreen (sometimes deciduous) lianas (sometimes climbing, perennial herbs) with leaf-opposed, simple or branched branch-tendrils with small adhesive cushions attaching plant to trees or cliffs etc. (rarely small succulent trees with swollen stem; Leea consists of evergreen trees, shrubs or perennial herbs without tendrils). Lenticels often abundant and significant. Some native species have economic potential; fruits of Mesoamerican Ampelocissus are used by local people for vinegar preparation and as table fruit.

Key differences from similar families leaf-opposite tendrils and/or inflorescences (axillary in Cucurbitaceae and Sapindaceae).

SYSTEMATIC two clades; Leeoideae (1/34-70, SE Asia, Malesia to New Guinea, tropical Australia, one species in Africa and Madagascar) does not occur in South America; all New World Vitaceae are Vitoideae; among this group, are five tribes; outsiders tribes Cayratieae (5/300-320; tropical and subtropical regions on both hemispheres, tropical regions in the Old World east to E Queensland) and Pathernocisseae (2/10-18; temperate Asia, North America, Mexico).

1. VITOIDEAE ▸ TRIBE AMPELOPSIDEAE (3/31) ‣ outsiders are Ampelopsis (c 16; temperate and subtropical regions of Asia and America), Nekemias (9; India, E and SE Asia, Java, Sulawesi, E North America).

1. Clematicissus Planchon. (inc. Cissus p.p.) Lianas, often with xylopodium; tendrils leaf-opposed, mostly dichotomously branched, lacking adhesive discs; leaves alternate, palmate, petiolate, stipulate, stipule adnate to petiole base; inflorescences compound cymes, leaf-opposite; flowers pedicellate, bisexual, 4- (South American species) or 5-merous (Australian species); fruit berry, globose or spheroidal, purple, purple-black or green, 1–4 seeded; seeds obovoid, chalaza marked on dorsal surface, two furrows on ventral surface. 6 spp., two from Australia, 4 from South America, centered in Bolivia, one up to Peru, Cl. pruinata (Weinm.) C. A. Zanotti & A. M. Panizza and Cl. striata (Ruiz & Pav.) Lombardi up to Brazil and Cl. tweedieana (Baker) Lombardi up to Cono Sur in Chile.

2. VITOIDEAE ▸ TRIBE CISSEAE (1/c. 350) ‣ a single genus.

2. Cissus L. (exc. Clematicissus p.p.) Woody or herbaceous climbing or scrambling lianas, or sometimes erect shrubs, hermaphroditic to polygamo-monoecious, sometimes with xylopodium; stems terete, winged or striated, often succulent; sometimes tuberous roots present. 350 spp., widely distributed in all tropical regions, a few extending into the temperate zone; 135 species in Africa, 85 species in Asia, 10 species in Australia, and 80 species in the Neotropics, mainly South America (60), 46 in Brazil (17 endemics); two spp. are rare in Brazil, one from Amapá state, another in Sergipe state, by Plantas Raras do Brasil’s book.

The berries of some South American Cissus e.g. C. stipulata Vell. and C. trigona Willd. ex Schult. & Schult. f. (both Brazilian, with the latter also in Bolivia), which have the biggest fruits of all Neotropical species of genus, are reported as sweet in Herbarium labels, whereas the small-fruited species usually have unpleasant fruits, filled with stinging raphides. C. verticillata (L.) Nicolson & C.E.Jarvis subsp. verticillata has the widest geographic and altitudinal distribution of all the Neotropical species; it occurs in almost all American countries, except Canada and Chile, at altitudes ranging from sea level to 2,500 m. In Brazil it is the only species under intense pharmacological study because of its reported medicinal properties. It is also cultivated around the world as an ornamental, although it is a potential weed, as in the Florida orange groves.

3. VITOIDEAE ▸ TRIBE VITEAE (5/180-200) ‣ outsiders are Acareosperma (1; Laos), Pterocissus (1; Hispaniola), Parthenocissus (10–18;), Ampelocissus (95–120; tropical regions on both hemispheres).

3. Vitis L. Vine-like shrubs or high climbing polygamo-dioecious vines, often with large woody stems. 78 spp., mostly temperate regions of the northern hemisphere, 29 in New World, mainly in North America, with V. tiliifolia Humb. & Bonpl. ex Roem. & Schult. extending into Venezuela, Colombia and Ecuador (with a doubtful record in French Guiana), and V. novogranadensis Moldenke endemic to Colombia.

29. ZYGOPHYLLALES

TWO FAMILIES, BOTH IN SOUTH AMERICA.

KRAMERIACEAE

§ PARASITIC (Prosopanche – ... – Krameria - Mitrastemon – APODANTHACEAE – SANTALALES - Lennoa – OROBANCHACEAE - Cuscuta)

Genera/species 1/18 Distribution New World Habit bisexual, usually shrubs or suffrutices (Krameria ixine Loefl.is a perennial herb, usually with woody rhizome); root hemiparasites (no specific host); the oils secreted by the 2 modified petals are collected by visiting female bees of the genus Centris, and mixed with pollen (and nectar from other taxa as Krameria does not produce nectar) into a paste, and fed to their developing larvae; Krameria species are dependent on the bees for pollination; this two modified petals are unique within the angiosperms.

SYSTEMATICS Only one genus.

1. Krameria L. ex. Loefl. Characters of family. 16 spp., 8 in South America: K. argentea Mart. ex Spreng., endemic to the Brazilian Shield in the states of Distrito Federal, Goias and SW Bahia; K. bahiana B.B. Simpson., endemic disjunct spp. from Bahia, in to coastal regions and in rock fields in Diamantina Range; K. cistoidea Hokk. & Arn. only in Chile in Antofagasta region (one of the driest places on Earth) at 100 to 1550 m; K. grandiflora A. St. Hill., in Brazil and adjacent Paraguay and Uruguay; K. ixine Loefl. from Sinaloa, Mexico across Venezuela, Colombia, Guianas and Caribbean; K. lappacea (Dombey) Burdet & B.B. Simpson can be found at altitudes up to 3,600 m from Ecuador to Jujuy in Andean Argentina; K. spartioides Klotzsch ex O. Berg grows in Colombia, Venezuela, N Brazil and Suriname; and K. tomentosa A. St. Hill., from Guyana to Bolivia and across Brazil in dry areas.

ZYGOPHYLLACEAE

§ FULLY CHLOROTROPHIC

Genera/species 22/c. 155 Distribution mainly tropical and subtropical regions in the Northern and Southern Hemispheres; some species in warm-temperate regions, with their largest diversity in arid and subarid areas. Habit usually bisexual (rarely dioecious), evergreen shrubs, suffrutices, or perennial or annual herbs (rarely trees), sometimes with axillary, simple or branched, spines. Nodes often swollen and/or articulated. Bark often bitter. Some species are succulent. Many species are xerophytes or halophytes. C₄ photosynthesis present (almost all species in Kallstroemia have C₄ photosynthesis; nearly all in Zygophyllum are C₃ photosynthetic).

Use Ornamental plants, timber and carpentries (extremely hard wood, Lignum vitae, from Guaiacum), fruits (Balanites aegyptiaca Del.), medicinal plants, edible flower buds. Some of the South American species have been used for their timber, notably Guaiacum which has extremely strong, hard, resinous wood; G. sanctum L. is known as lignum vitae.

Certainly, one of the world's heaviest and hardest ironwoods is the Caribbean tree called lignum vitae (G. officinale), with a specific gravity of 1.37. The name lignum vitae means ‘wood of life’, owing to the medicinal properties of the sweet-smelling resin; the density and high resin content of the wood make it extremely resistant to friction and abrasion and account for its remarkable self-lubrication properties. Under certain conditions it actually wears better than iron. In fact, the highly-prized wood was used for end grain thrust blocks which lined the propeller shafts of steamships.

SYSTEMATICS five subfamilies, Morkillioideae (3/4, all only in Mexico) and Seetzenioideae (1/1, North Africa to Afghanistan and India, N and E Cape) do not occur in South America.

1. SUBFAMILY BALANITOIDEAE (6/80-90) ‣ outsiders Tribulus (40–45; tropical and subtropical regions on both hemispheres), Kelleronia (3; NE Africa, S Arabian Peninsula), Sisyndite (1; S Namibia, N and W Cape), Neoluederitzia (1; S Namibia), Balanites (9; tropical Africa, Arabian Peninsula to India and Burma).

1. Kallstroemia Scop. Prostrate to decumbent herbs, mainly in dry areas; leaves opposite, parapinnate, entire folicules; inflorescences as isolated flowers, axillary or supraxillary, pendunculate; petals white to orangish. 21 spp. of New World, centered in U.S.A. of up to south Mexico (9), 8 in South America: K.adscendens (Anderss.) Robins., endemic to the Galápagos Islands, Ecuador; K. boliviana Standi., disturbed areas in the semiarid valleys on the eastern face of the Cordillera Oriental of Bolivia, and known from a single similar locality in Peru; K. maxima (L.) Hook. & Arn., from U.S.A. to Colombia east to Guianas and Caribbean; K. parviflora Norton, U.S.A. to Peru; K. pennellii D. M. Porter., know only from the type locality in Cajamarca in Peru; K. pubescens G. Don. occur in the Lesser Antilles, Puerto Rico, and Jamaica; across N South America and north through Central America to Yucatán on the east and Sinaloa, Mexico; south through Colombia and Ecuador to N Peru; K. tucumanensis Desc, O’Don & Lourt., in semiarid S Bolivia and NW Argentina; and K. tribuloides (Mart.) Steud., semiarid NE Brazil, S Bolivia, and NW Argentina; apparently native to Argentina and Bolivia and introduced into Brazil; occur in open sandy places, riverbanks, railroad embankments, and roadsides from 300 to 1,800 m; sympatric over much of its range with K. tucumanensis; Brazilian collections were mainly from sandy places along the Rio São Francisco, the largest river in the area; this species is found in similar situations in Argentina; flowers from November to May.

2. SUBFAMILY LARREOIDEAE (9/32–36) ‣ all genera occur in South America.

2. Bulnesia Gay. Shrubs to medium sized trees. 4 spp. from Peru, Bolivia and Cono Sur.

3. Gonopterodendron (Griseb.) A.C.Godoy-Bürki. Trees. 5 spp., two in Colombia and Venezuela, remaining three in Bolivia, Brazil (only G. sarmientoi (Lorentz ex Griseb.) Godoy-Bürki., reported in small areas at the state of Mato Grosso do Sul, in wetland riverbanks) and Cono Sur.

4. Guaiacum L. 5 spp., mainly in Mexico (two endemics) Atlantic coast of Central America, G. officinale L. also N South America, in coasts of Colombia, Venezuela and Guianas, and Caribbean, although present in Trinidad and Guyana, it may not be native there; because of development, harvest, and fires, G. officinale is extinct or near extinction on several Caribbean islands.

5. Izozogia G. Navarro. Only one sp., I. nellii G. Navarro, endemic to Bolivia.

6. Larrea Cav. Herbs, allelopatics. 5 spp., one in North America and remaining four in South America (Bolivia, Argentina, Chile and Peru).

7. Metharme Phil. ex Eng. One little-known species, M. lanata Phil., endemic to Tarapacá in N Chile.

8. Pintoa Gay. Only one sp., P. chilensis Gay, endemic to the Atacama province of Chile.

9. Plectrocarpa Gillies ex Hook. & Arn. Two spp., P. rougesii Descole, O'Donell & Lourteig and P. tetracantha Gillies ex Hook. & Arn., endemics to NW Argentina.

10. Porlieria Ruiz & Pav. 4 spp. Argentina, Uruguay, Chile (one endemic), Bolivia (one endemic) and Peru.

3. SUBFAMILY ZYGOPHYLLOIDEAE (6/183) ‣ most recent phlogeny indicate the following classification: Roepera A. Juss. with c. 60 spp. in Australia and S Africa, Zygophyllum L. with c. 50 species in Asia, Tetraena with c. 40 spp. mainly in Africa and Asia, Augea with a single species in southern Africa, Melocarpum (Engl.) Beier & Thulin with two spp. in the Horn of Africa region, and Fagonia L. with c. 30 species in both the Old and the New Worlds.

11. Fagonia L. c. 30 spp., Old World, in arid regions bordering Mediterranean in Europe, Asia and Africa (Yemen to India, south into Kenya, two spp. disjuncts in Namibia, Botsuana and South Africa), nine in deserts about the Gulf of California in North America (6 endemic to Mexico), and one sp., F. chilensis Hook. & Arn., in western coast of South America in N Chile and S Peru.

30. FABALES

ALL FAMILIES IN SOUTH AMERICA.

The sister-group relationships within Fabales are not unambiguously resolved and the bootstrap or bayesian support for any branch is more or less weak. Hence, Fabaceae, Polygalaceae, Quillajaceae or Surianaceae may be identified as sister to the remainder, depending on the characters and analysis methods used.

Bello & al. (2007, 2009) found the following topologies based on matK, rbcL or combined matK and rbcL, respectively: [Polygalaceae + [Fabaceae + [Quillajaceae + Surianaceae]]], [Fabaceae + [Quillajaceae + [Polygalaceae + Surianaceae]]] and [Quillajaceae + [Surianaceae + [Fabaceae + Polygalaceae]]]. Persson (2001) received the topology [Polygalaceae + [Surianaceae + [Fabaceae + Quillajaceae]]]. Qiu & al. (2010) found the following topology: [Quillajaceae + [Fabaceae + [Polygalaceae + Surianaceae]]]. Soltis & al. (2011) received [[Quillajaceae + Polygalaceae] + [Fabaceae + Surianaceae]]. Finally, Moore & al. (2011) found [Surianaceae + [Quillajaceae + [Fabaceae + Polygalaceae]]].

SURIANACEAE

§ FULLY CHLOROTROPHIC

Genera/species 5/9 Distribution Tropical beaches (Suriana maritima L., pantropical along sea coasts, except in West Africa, but truly native of New World), subtropical Australia (Cadellia), NW (Stylobasium), NE and E (Guilfoylia) C Australia, Mexico (Recchia, 4 spp. endemic). Habit Usually bisexual (in Stylobasium polygamomonoecious), evergreen small trees or shrubs. Some species are xerophytes.

SYSTEMATICS outsiders Recchia (3; Mexico), Cadellia (1; SE Queensland, NE New South Wales), Guilfoylia (1; E Queensland, NE New South Wales), Stylobasium (2; mainly W and C Australia)

1. Suriana L. Evergreen small trees or shrubs, usually reaching a hight of 1-2m. Only one sp., S. maritima L., in coastal regions and limestone coastal outcrops; in New World occur Atlantic and Caribbean Coast from Florida to E Brazil; in Brazil, this species is found in few and rare places in coast, very narrow restricted of beaches in Piauí Alagoas and Bahia states, in psamophylous habitats.

POLYGALACEAE

§ FULLY CHLOROTROPHIC

Genera/species 29/1,050–1,250 Distribution cosmopolitan except polar regions, Polynesia and New Zealand. Habit usually bisexual (in Balgoya functionally unisexual), perennial or annual herbs, evergreen or deciduous? shrubs, trees (rarely lianas; Salomonia comprises achlorophyllous root holoparasites and Epirixanthes holomycotrophs). Some species are xerophytes. Often with paired crateriform glands (extrafloral nectaries) or spines at nodes (sometimes elsewhere).

628 spp. in New World, 460 in South America.

Key differences from similar families

Species with papilionaceous flowers differ from Fabaceae subfam. Papilionoideae in having:

Simple leaves (vs. usually compound).

A 3-merous corolla with a standard of two petaloid sepals (vs. usually a 5-merous corolla including a standard of one petal).

One ovule per locule in 2-8 carpellate ovaries (vs. usually several ovules per locule and 1-carpellate ovaries).

SYSTEMATIC monogeneric Xanthophyllum Clade (1/110, S India, Sri Lanka, SE Asia, Malesia, New Guinea, Solomon Islands, Queensland) does not occur in South America; all New World species are Polygaloideae subfamily, with three tribes, Carpolobieae (2/7, tropical Africa, Madagascar) does not occur in South America.

1. POLYGALOIDEAE ▸ TRIBE MOUTABEEAE (c 5/15) - outsiders are Balgoya (1; New Caledonia) and Eriandra (1; New Guinea).

1. Barnhartia Gleason. Only one sp., B. floribunda Gleason, restricted to the Guianas, S Venezuela and Amazonas state in N Brazil.

2. Diclidanthera Mart. 5 spp. from Amazon rainforest, from Colombia, Venezuela, Guyana, Brazil (4, D. laurifolia Mart. endemic), Peru and Bolivia.

3. Moutabea Aubl. 11 spp., from Costa Rica (2) to Bolivia, in Amazon rainforest, few outside this area; all occur in Brazil, 4 endemics.

2. POLYGALOIDEAE ▸ TRIBE POLYGALEAE (c 13/950–1.150) - outsiders Comesperma (34, Australia), Epirixanthes (5, tropical Asia), Badiera (7, Caribbean), Muraltia (121, Africa), Rhinotropis (17, SW U.S.A., Mexico), Phlebotaenia (2, W Indies), Heterosamara (17, tropical and S Africa, tropical and subtropical Asia), Polygaloides (6, Europe, Mediterranean, northern Africa, W Asia, E North America), Pteromonnina (c 30), Salomonia (5, tropical Asia, tropical Australia to E Queensland). Epirixanthes consists of holomycoheterotrophs, whereas Salomonia comprises root holoparasites.

Key to genera of neotropical Polygaleae

1. Fruit 1-, 2-, or 4-winged, New World distribution (except for a few Securidaca) - 2

2. Fruit a symmetric 2-winged samara ------------ Monnina (in part)

2. Fruit a strongly asymmetric 1-, 2-, or 4-winged samara or samaroid capsule- 3

3. Tree, 4-winged (2 large, 2 small) samaroid capsules ------------ Phlebotaenia

3. Subshrub or woody liana, rarely shrub or treelet, 1–2- winged samara - 4

4. Subshrub (Polygala-like), keel with 2 reflexed orbicular appendages ------------ Monrosia

4. Woody liana, rarely shrub or treelet, keel without appendages ------------ Securidaca

1. Fruit a 2-locular capsule or a drupe, worldwide distribution - 5

5. Seeds comose (hairs often longer than seed) - 6

6. Inflorescences panicles ------------ Bredemeyera

6. Inflorescences racemes, sometimes with short internodes and resembling an axillary fascicle, or solitary axillary flowers - 7

7. Plants unarmed, Australia ------------ Comesperma

7. Plants armed with thorns - 8

8. Lower external sepals free, capsules glabrous ------------ Hualania

8. Lower external sepals connate, capsules tomentose ------------ Rhamphopetalum

5. Seeds lacking a coma, usually with short trichomes (almost always shorter than seed) or glabrous - 9

9. Calyx deciduous in fruit - 10

9. Calyx persistent in fruit - 15

10. Fruit a drupe ------------ Monnina (in part)

10. Fruit a 2-locular capsule - 11

11. Keel ecristate, New World - 12

12. Shrubs 2 m or more tall (rarely flowering when smaller) ------------ Badiera

12. Herbs (but often suffrutescent) usually less than one m tall ------------ Hebecarpa

11. Keel cristate, Old World (except one sp. of Polygaloides in North America) - 13

13. Keel with a 2-lobate crest; pollen heteropolar ------------ Heterosamara

13. Keel with a plurilobate crest; pollen isopolar - 14

14. Petioles 1–1.5 cm long ------------ Senega subg. Chodatia

14. Petioles nearly lacking to 4 mm long------------ Polygaloides

15. Achlorophyllous annual herbs ------------ Epirixanthes

15. Chlorophyllous herbs (annual to perennial) and shrubs - 16

16. Fruit a drupe or horned capsule ------------ Muraltia

16. Fruit a capsule lacking horns (margin sometimes fringed) - 17

17. Capsule margins fringed with sharp teeth, Australasia ------------ Salomonia

17. Capsule margins almost always entire (rarely slightly crenate or shallowly undulate, but not fringed with sharp teeth) - 18

18. Flowers with keel crested ------------ Senega

18. Flowers with keel ecristate (but often 3-lobate) - 19

19. Lower external sepals connate (the resulting structure often 2-lobate) ------------ Asemeia

19. Sepals all free, none 2-lobate - 20

20. Stipular nectaries on stem at base of petiole ------------ Caamembeca

20. Nectaries lacking on stem (and in inflorescence) ------------ Gymnospora

4. Acanthocladus Klotzsch ex Hassk. Shrub to tall tree; branches usually ending in sharp thorns; leaves subopposite or alternate; inflorescence usually an axillary, congested brachyblast, sometimes appearing fasciculate or rarely a raceme with a conspicuous rachis; flowers papilionaceous, pedicellate; petals usually whitish purple. 8 spp. in seasonally dry to moist forests in South America in Amazon rainforest (1), Paraná Basin in Brazil (1), Atlantic Forest (2, Bahia to Rio de Janeiro), Ecuador (1), Colombia (1) and one widely distributed in center South America (up Bolivia and Argentina); A. dukei (Barringer) J.F.B. Pastore & D. Cardoso occur in Darien region in Panamá.

5. Asemeia Raf. Erect herbs or shrubs, roots fleshy to lignous, stems subaphyllous to frondose, glabrous to pubescent; racemes terminal, axillary or extra-axillary; keel petal pale lilac, apex yellow; capsules herbaceous, oblong elliptic to elliptic, glabrous or cilate at margins. 32 spp. (and one variety) in New World, occurring in seasonally dry, open savannas (or comparable vegetation), or less frequently associated with forest vegetation; two subgenera:

§ subg. Asemeia ▸ 28 spp., South and Central America, except for A. grandiflora (Walter) Small from North and Central America and the Caribbean; 21 spp. in Brazil, 19 endemics.

§ subg. Apopetala ▸ 7 spp., endemic to Central America and Mexico with one species, A. echinosperma (Görts) J. F. B. Pastore & J. R.Abbott, endemic to Suriname.

6. Bredemeyera Willd. (exc. Hualania, Rhamphopetalum). 12 spp., B. divaricata (DC.) J.F.B. Pastore distributed from Mexico to temperate parts of Argentina, and all remaining 11 restricted of South America, 9 in Brazil, 7 endemics.

7. Caamembeca J.F.B.Pastore. Corolla papilonaceous, trimerous, sometimes with two rudimental petals; fruit a samara, berry or capsule, fruit dry, samara or capsule, calyx persistent in fruit, keel non-crested, sometimes emarginated, external sepals free, ovary glabrous, glands at petiole base, capsules elliptic. 13 spp., three occur only in Peru, Bolivia and Paraguay, one in Amazon rainforest, and all remaining species are endemic to E Brazil; C. martinelli (Marques & E.F. Guim.) J.F.B. Pastore from coast Bahia state is a rare plant in Brazil, by Plantas Raras do Brasil’s book.

8. Gymnospora (Chodat) J.F.B.Pastore. Pubescent pedicels, pubescent and free external sepals, calyx persistent in frutification, noncristate carina, puberulent and chartaceous capsules, and minute seed appendages or caruncles. Two spp., endemic to C, S & E Brazil, up to Bahia and Mato Grosso do Sul, apparently restricted to forest margins and savannas; G. blanchetii (Chodat) J. F. B. Pastore, has been reported principally from SE Bahia near Ilheus, although the taxon was collected twice from other Brazilian states (Minas Gerais and Rio de Janeiro).

9. Hebecarpa (Chodat) J. R. Abbott. Perennial herbs or shrubs, usually multi-stemmed, stems lack thorns; racemes terminal; flowers with the entire periant deciduous in fruit. 19 spp., SW U.S.A., Mexico (high diversity) to Andean regions in South America, and Caribbaen region; three in South America, from Venezuela to Bolívia.

10. Hualania Phil. (off Bredemeyera) Shrubs, 2–3 m tall, densely branched, cylindrical, stems terminating in spines, young branches flexible; leaves 1–3 × one mm, sessile, caducous, triangular, apex acuminate, margin entire, both surfaces glabrous. Only one sp., H. colletioides Phil., remarkably adapted to dry areas being restricted to the Del Monte Province, in Catamarca, La Rioja, Mendoza, and San Juan at 1,400–1,575 m altitude range, usually growing along seasonal rivers, and producing odoriferous flowers and fruits during the dry season

11. Monnina Ruiz & Pav. Herbs to shrubs. papilionaceous flowers and drupes, sometimes with xylopodium. 178 spp. distributed from Mexico to Bolivia and Argentina, 166 in South America, Monnina s.s. is highly centered in Andes of Peru (46), Ecuador (39) and Colombia (32); 14 spp. in Brazil, 6 endemics, some very narrow; nine in southern region.

12. Monrosia Grondona. (off Polygala/Senega) Prostrate subshrubs, to 10 cm, with several branches, the base woody and glabrous, distally herbaceous and tomentose, stems not terminating in spines, young stems green. Only one sp., M. pterolopha (Chodat) Grondona, restricted to the Argentinian provinces of Catamarca, La Rioja, and San Juan, at 2,000 to 3,000 m altitude range.

13. Rhamphopetalum J.F.B.Pastore & M.Mota. (off Bredemeyera) Shrubs, 15–50 cm, branches intensely green to greyish, tomentose, stems terminating in spines, young stems green; leaves all alternate, sessile, coriaceous, lamina 4–8×1–2 mm, elliptic to obovate, base acute, margin entire, apex attenuate to rounded, both surfaces tomentose. Only one sp., R. microphyllum (Griseb.) J.F.B.Pastore & M.Mota, in the provinces of Catamarca, Chubut, La Pampa, La Rioja, Mendoza, Neuquén, and San Juan, with the distribution ranging from 27°S in Catamarca to 45°S in Chubut.

14. Securidaca L. Papilionaceous flowers and unilaterally winged samaras. 55 spp. in the Caribbean and from Mexico to Paraguay, 49 in South America, 26 in Brazil, 7 endemics; S. acuminata A. St.-Hil. from Minas Gerais state is a rare plant in Brazil, by Plantas Raras do Brasil’s book.

15. Senega Spach. (off Polygala; exc. Monrosia) Usually herbaceous and with papilionaceous flowers and capsule. 587 spp., 278 spp. widely distributed in the New World, 174 in South America, 110 in Brazil, c. 62 endemics; 20 Senega species from several states are rare plants in Brazil, by Plantas Raras do Brasil’s book. Three subgenera.

§ subg. Clinclinia ‣ 26 spp. endemic to Argentina and/or Chile, but with two species, S.[P.] aspalatha L. and S.[P.] cyparissias A.St.-Hil. & Moq., also occurring in Brazil.

§ subg. Monninopsis ‣ nine species occurring mainly in Mexico and the adjacent U.S.A., S.[P.] cisandina Chodat (Bolivia) and S.[P.] darwiniana A.W.Benn. (Cono Sur).

§ subg. Senega ‣ c. 175 spp., with most of them in the Americas, but with 9 species native to Africa, and with S.[P.] paniculata L. naturalized around the Paleotropics.

QUILLAJACEAE

§ FULLY CHLOROTROPHIC

Genera/species 1/2 Distribution southern South America Habit evergreen glabrous tree with saponaceus bark; nodes unilacunar. In young axes, cork in iniated subeiderm, and t rhytidome is scaly. An endodermis is lacking, and the secondary phloem dilates diffusely.

SYSTEMATIC Only one genus, endemic to South America.

1. Quillaja Molina. Caracters of family, sometimes with lignotuber. Two spp., Q. saponaria Molina grows in the winter-rain region from 31 to 38ºS in center Chile, from sea levelup to 2,000 m altitude, and in the valleys once formed dense forests which have largely been replaced by pastures; and Q. brasiliensis (A.St.-Hil. & Tul.) Mart. is a tree of mixed mesophytic forests and extends from São Paulo, Brazil, to Uruguay and Misiones, Argentina; it is also reported from Peru (Depto. Cuzco) but it is questionable whether it is indigenous there.

FABACEAE

§ FULLY CHLOROTROPHIC

Genera/species 793/18,300-18,400; Distribution cosmopolitan except Antarctica. Habit usually bisexual (rarely monoecious, andromonoecious or polygamomonoecious), evergreen or deciduous trees, shrubs, lianas or suffrutices, perennial, biennial or annual herbs. Numerous species are xerophytes, whereas some are aquatic. Petiole or branch sometimes modified into photosynthesizing phyllodia or phyllocladia, respectively; the tallest tropical tree, Dinizia excelsa Ducke, is in this family.

Legumes have been gathered, cultivated, eaten and used in a multitude of other ways by humans for millennia and are arguably as important as grasses in global terms. Certainly the range of uses of legumes is broader than that of the grass family. Legume products contribute enormously to the world`s economy through food (for animals and humans) and drink, pharmaceuticals and medicine, bio-fuels, biotechnology (as industrial enzymes), building and construction, textiles, furniture and crafts, paper and pulp, mining, manufacturing processes, chemicals and fertilisers, waste recycling, horticulture, pest control, and ecotourism.

DRINKS. across the globe legumes have been used as substitutes for coffee, tea, tobacco, betel-nut, hops, garlic, ebony wood, cocaine, soap and liquorice (true liquorice, Glycyrrhiza glabra L. is also a legume).

FERTILE SOILS ancient cultures were aware of the ability of many legumes to improve the soil, even if they did not then appreciate that this results from symbiotic nitrogen fixation. Some 40 to 60 metric tons of nitrogen are fixed annually by agriculturally important legumes; many species are used as soil improvers and stabilisers and in reforestation programmes.

POISONOUS natural accumulation of nitrogen has also resulted in predation of legumes by a wide range of animals and insects. To combat this, the family has evolved a wide repertoire of chemical defences based on secondary compounds, especially alkaloids; humans have exploited the chemistry of legumes by utilising many species as medicines, insecticides, molluscicides, abortifacients, purgatives, fish, arrow and ordeal poisons, anti-fungal agents, aphrodisiacs and hallucinogens. Some legumes have been used as antidotes to poisons, as anti-inflammatories and antiseptics; the senna pod is a well known laxative.

MATERIALS several legumes are rich in gums used as glues and food thickeners (e.g. Acacia Mill. s.l., Astragalus L.), resins used in paints, polishes and varnishes (e.g. Hymenaea, Copaifera, Prioria) and oils used in lubricants and cosmetics; important dyes, such as brasil, indigo and dyer`s greenweed all come from legumes, and several species are used as inks, and for tanning leather.

CROPS grain and forage legumes are grown on approximately 180 million hectares (12 to 15 percent) of the Earth`s arable surface and account for 27 percent of the world`s primary crop production with grain legumes alone contributing 33 percent of the dietary protein nitrogen needs of humans. The main dietary legumes (the pulses) include several species of bean (Phaseolus), the pea (Pisum sativum L.), chickpea (Cicer arietinum L.), broad bean (Vicia faba L.), pigeon pea (Cajanus cajan (L.) Huth), cowpea (Vigna unguiculata (L.) Walp.) and lens (Lens culinaris Medik.). Legumes (mainly soybean, Glycine max (L.) Merr., and peanut, Arachis hypogaea L.) also contribute more than 35 percent of the world’s processed vegetable oil.

FORAGE legumes provide the protein, fibre and energy that have underpinned dairy and meat production for centuries. In temperate regions, alfalfa (Medicago sativa L.) is an important forage for cattle. Other important temperate pastures used for forage, hay, silage and green manure, include clovers (Trifolium), trefoil (Lotus corniculatus L.) sweetclovers (Melilotus) and vetches (Vicia). In the tropics, species of Aeschynomene, Arachis, Centrosema, Desmodium, Macroptilium and Stylosanthes are all being used to improve tropical pasture systems, with Stylosanthes the most geographically widely distributed.

ORNAMENTAL in the tropics, avenues and parks are invariably adorned with a representation of Albizia, Cassia, Delonix, Senna and Tipuana; spectacular, showy-flowered species of Caesalpinia, Calliandra, Mucuna and Strongylodon have also become popular in tropical gardens. Multipurpose trees and shrubs have long been selected and refined by local communities for shade, ornament, forage, fodder, fuelwood, bee forage for honey production, and soil enrichment. Regional favourites include Calliandra, Gliricidia, Inga and Leucaena in Central America; some are grown as impenetrable spiny hedges, living fence-lines and windbreaks.

TIMBER species of Acacia, Anadenanthera, Dalbergia, Erythrina, Prosopis and Pterocapus are all important woody tree legumes in forestry; legume timber and wood from many species has long been put to a multitude of uses, ranging from heavy construction (house and boat building, railway sleepers and cart wheels), to paper and plywood manufacture, and fine furniture production, carpentry, marquetry and veneer work. Woods high in silica (e.g. Dicorynia guianensis Amshoff) are of particular value in marine construction.

EXTINCTION some species (e.g. Dalbergia nigra (Vell.) Allem. ex Benth.) are now considered rare and endangered because of over-exploitation due to their commercially valuable timbers. Paubrasilia echinata (Lam.) E. Gagnon, H. C. Lima & G. P. Lewis (Brazil Wood or pau-brasil, the tree from which the country Brazil took its name), once a source of an important red dye and still the preferred wood for violin bows, has been reduced, by major habitat destruction, to a few populations along the Atlantic coast of Brazil.

SYSTEMATIC six subfamilies, only Duperquetoideae (1/1, tropical West and Central Africa) absent in South America; 4859 spp. in South America.

1. SUBFAMILY CERCIDIOIDEAE (14/300–460) - outsiders Cercis (10; Mediterranean, E Asia, North America to NE Mexico); Adenolobus (2; S Angola, Namibia, N Botswana, N Cape); Brenierea (1; Madagascar); Griffonia (4; tropical West Africa); Gigasiphon (4–5; tropical Africa), Tylosema (4; Africa southwards to South Africa), Barklya (1; NE Queensland), Lysiphyllum (8; Malesia and eastwards to tropical Australia), Lasiobema (15–20; E Asia from E Himalaya to Japan, Indochina, Java).

1. Bauhinia L. Trees and shrubs (sometimes semi-scandent), sometimes with xylopodium. 150-300 spp., 138 spp. in New World, 109 in South America, 68 in Brazil, 39 endemics, three of them are rare plants in Brazil, by Plantas Raras do Brasil’s book; seasonally dry tropical bushland, woodland, wooded grassland (savannas of C Brazil, known as cerrado), dry seasonal scrubland of NE Brazil (caatinga) and coastal forest; several on sand or limestone; nine sections at Bauhinia s.s., Bauhinia, Pauletia, Amaria, Alvesia, Micralvesia, Telestria, Pseudophanera, Afrobauhinia and Gigasiphon, only three firsts in New World. Some South American species possibly have nectar-spurs.

§ sect. Pauletia ▸ c. 70 spp.; tropical America and two in Southern Asia, South China, and Malesian Area.

§ sect. Bauhinia ▸ 17 spp., Mexico, adjacent U.S.A. (Texas), northern Central America, Greater Antilles, and NE Brazil.

§ sect. Amaria ▸ c. 15 spp. from N South America to Mexico.

2. Phanera Lour. 60-65 spp., 18 in New World, all in Brazil (12 endemics), six of them up to Peru and from Venezuela to French Guiana.

3. Piliostigma Hochst. Shrubs to trees. Six spp. in two sections:

§ sect Piliostigma ▸ 4 spp., Africa, Southern Asia, and Australia.

§ sect. Benthamia ▸ two spp., B. (Piliostigma) glaziovii Taub. and B. (Piliostigma) uruaguayensis Benth., from Brazil (former endemic) to Argentina and Uruguay.

4. Schnella Raddi. Liana or scandent shrub with tendrils. 16 spp., 15 in South America (a single only in Central America), two of then up to Mexico, Central America and Caribbean, S. microstachya Raddi is the only species that exists as far south as Paraguay and N Argentina; 9 spp. in Brazil, 8 endemics.

Wet tropical rainforest, including vàrzea and terra firme forest, although some are also found in seasonally dry forest such as dry seasonal scrubland of NE Brazil (caatinga), and savannas of C Brazil (cerrado); the species occupy a range of elevations and have been found at 1000 m above sea level.

2. SUBFAMILY DETARIOIDEAE (84/c. 760) ▸ six lineages, tribe Schotieae (1/5, southern Africa) and Danieleae (2/6, Africa and Madagascar) not occur in South America.

2.1 DETARIOIDEAE ▸ TRIBE BARNEBYDENDREAE (2/2) - two genera, both in South America.

5. Barnebydendron J. H. Kirkbr. Deciduous tree 5 - 40 m; single or multiple-stemmed; leaves once pinnate, 6 – 23 cm long; inflorescence racemose, 3 – 17.5 cm long, cauliflorous, flowers bright scarlet; flowers 2.3 – 3.4 cm long with exserted stamens and style; not scented; petals 3 (occasionally with 2 vestigial petals), fleshy, obovate, scarlet, pink-red, pink or pale pink to almost white, fruit a flattened, pendent, samaroid pod. Only one sp., B. riedelii (Tul.) J.H. Kirkbr., disjunc from Guatemala, Costa Rica, and Panamá, small populations in S Venezuela, Peru (in floodplains and ravines from 350 – 850 m) and adjacent areas in Acre state in Brazil, over Purus River Valley, and E Brazil from southern Bahia up to Rio de Janeiro state (on dry, rocky hillsides, from 50 – 800 m).

6. Goniorrachis Taub. Evergreen tree (5– 15(–30)-m high), flowers characterized by their long hypanthium with an adnate gynoecium on the rim, and by the radially symmetric corolla and diplostemonous androecium superficially resembling Rosaceae flowers. Only one sp., G. marginata Taub., restricted of dry seasonal scrubland of NE Brazil (caatinga) and dry forests, especially on richer soils and along temporary rivers, from Sergipe and Bahia to Minas Gerais, Espírito Santo and northern Rio de Janeiro. Used for timber (but known to cause allergic responses).

2.2 DETARIOIDEAE ▸ PRIORIA CLADE (3/16) - outsiders Colophospermum (1; tropical southern Africa), Hardwickia (1; drier regions of India).

7. Prioria Griseb. 16 spp., tropical Africa, tropical Asia to islands in the Pacific, one endemic to Costa Rica, P. copaifera Griseb. from Nicaragua to NW Colombia and Jamaica.

2.3 DETARIOIDEAE ▸ TRIBE DETARIEAE (14/163–170) - outsiders Stemonocoleus (1; C Africa), Augouardia (1; Gabon), Neoapaloxylon (3; Madagascar), Baikiaea (7; tropical to S Africa in Namibia and Botswana), Detarium (3; tropical and subtropical West Africa to Sudan), Gilletiodendron (5; tropical Africa), Hylodendron (1; coasts of the Gulf of Guinea), Sindora (18–20; SE Asia, Malesia), Sindoropsis (1; Gabon), Tessmannia (12; tropical Africa), Eurypetalum (3; Cameroon to Gabon).

8. Copaifera L. Trees or shrubs up to 40 m tall in several species, containing resin; leaves imparipinnate; leaflets 10–20, alternate to subopposite, elliptic to obovate-oblong, with translucent dots; inflorescences borne in axils of terminal leaves; flowers bisexual, distichous; pod suborbicular, flattened; seed solitary. 42 spp., 34 in New World, most diverse in S America (all species, centred in Brazil (25, 18 endemics); c. 10 spp. in Amazonian Guianas, Brazil, Venezuela, Colombia and Bolivia; c. 15 spp. in drier habitats in Brazil to Bolivia and Paraguay; three spp. in the Caribbean (one disjunct to E Brazil, one extending from Panamá to Venezuela); two spp. in C America (Panamá and Costa Rica). 4 spp. in Africa (3 in W and WC regions and one sp. in Zambezian south tropical Africa (Angola, Zambia and Congo (Kinshasa)); a single species occurs in Malesia (Borneo); tropical lowland rain forest (sometimes inundated), seasonally dry forest, thorn forest, woodland and shrubland.

Commercially produced high quality resins (copal, copaiba, copaiva, Jesuit’s balsam) are used for medicines (scented gums are also used as unguents), varnishes, lacquers, paints and fuel (the oleo-resins are said to be a substitute for diesel); other uses are timber, e.g., for construction, bridges, shipbuilding, furniture, joinery, panelling, turnery and blockboard.

9. Eperua Aubl. Trees, often in monodominant stands. 17 spp., NE Amazonian S America in Brazil up to Cerrado (14, 6 endemics), Guianas, E Venezuela and NE Colombia, in tropical lowland rain forest, often along rivers and in inundated areas, and in seasonally dry forest, woodland scrub and wooded grassland; planted as ornamentals; several species (wallaba, apa, wamara) are used for timber in heavy construction, roofs (shingles), carpentry, joinery, furniture and flooring; for firewood and charcoal, and for resin or wallaba oil (e.g., E. oleifera Ducke).

10. Guibourtia Benn. Trees or shrubs. 14-15 spp., Africa (Guinea-Congolean WC Africa (7); dryland S tropical Zambezian (c. 3) and N tropical Sudanian (1) regions, and c. 2 spp. in Zanzibar-Inhambane E Africa), with a single neotropical species, G. hymenaeifolia (Moric.) J. Léonard, disjunct between Cuba and seasonally dry E Brazil, Paraguay and Bolivia, in tropical lowland (sometimes swampy or seasonally inundated) rain forest, seasonally dry forest, woodland, bushland and thicket, often along rivers and on sandy soils.

Various species used for timber (bubinga, akume, ovangkol, hyedua), e.g., in high quality furniture, cabinet work, joinery, panelling, veneers, heavy carpentry, implement handles, boat masts, for firewood and charcoal; gum-copal (for incense, coating pills, varnishes, illuminants and mosquito repellents), medicine, fish poisons and (in at least one species) edible seeds.

11. Hymenaea L. Unarmed evergreen trees up to 40 m tall; bark grey to greyish-white, smooth or rough; young branchlets puberulous to glabrous; leaves with a single pair of leaflets with many pellucid gland dots; inflorescence paniculate; flowers white; petals 5; pod indehiscent, thick, woody, ± resinous-warty with a pithy endocarp, 1–3-seeded; seeds ± ellipsoid, hard. 21 spp. from New World, 18 in tropical S America with centres in Amazonia and coastal Atlantic forest (c. 6) and dryland NE to SE Brazil (5 in this region; one of which extends to Paraguay and Argentina); one sp. widely distributed in the Neotropics (extending to C America, the Caribbean and Mexico); one sp. endemic to Cuba, allied to a single species in coastal E Africa, Madagascar and Mascarene Islands. 18 spp. in Brazil, 11 endemics, one of them, from Maranhão state, is a rare plant in Brazil, by Plantas Raras do Brasil’s book.

Tropical riverine and inundated forest to seasonally dry forest, woodland, thorn forest, bushland and thicket, often on slopes; resin (referred to as S American or Zanzibar copal, depending on the species) is used for incense, glue, varnish, shellac and as traditional medicines; other uses include edible fruits, timber (algarrobo, jatoba, courbaril, guapinol) for construction, high quality furniture, cabinetry, veneers, joinery, panelling, turnery, musical instruments and boat building, the bark is used for canoes, and some species are cultivated as ornamentals.

12. Peltogyne Vogel. Trees often in monodominant stands up to 40 m tall. 27 spp., all in South America, centred in Amazonian (Brazil (also Atlantic forest and c. 3 spp. to the drier NE), Guianas, Venezuela, Colombia and Bolivia), 3 spp. in C America, one extending to Mexico; one S American species to Trinidad. 25 in Brazil, 16 endemics, five of them (4 in Amazonas and one in Rio de Janeiro state) are rare plants in Brazil, by Plantas Raras do Brasil’s book.

Typically in tropical lowland rain forest, often along rivers; less commonly in seasonally dry forest, woodland or thorn scrub. Several species are utilised for their high quality timber (amaranth, amarante, purpleheart, violetwood, pau roxo), e.g., in furniture, cabinet work, flooring, construction, turnery, decorative veneers, marquetry, musical instruments and carving.

2.4 DETARIOIDEAE ▸ TRIBE AMHERSTIEAE (c 56/c. 600) - three subtribes, Saraceae (4/16, India, Sri Lanka, southern China, SE Asia, Malesia to Sulawesi) absent in South America.

∎ SUBTRIBE AFZELIEAE (3/18) ▸ outsiders Afzelia (13; tropical Africa, tropical Asia), Intsia (4; SE Asia to Pacific coasts).

13. Brodriguesia R. S. Cowan. Small tree, yellow flowers. Only one sp., B. santosii R.S. Cowan, endemic to Atlantic Forest of Sergipe and Bahia state in NE Brazil, and a rare plant in Brazil, by Plantas Raras do Brasil’s book.

∎ SUBTRIBE AMHESTIEAE ▸ outsiders Lebruniodendron (1; coasts of the Gulf of Guinea), Normandiodendron (2; Central Africa), Plagiosiphon (5; Central Africa), Zenkerella (6; coast of Guinea, tropical E Africa), Neochevalierodendron (1; Gabon), Gabonius (1; Gabon), Micklethwaitia (1; Mozambique), Annea (2; tropical West and Central Africa), Scorodophloeus (2; coasts of Guinea and E Africa), Hymenostegia (7–13; along the Gulf of Guinea); Librevillea (1; Gabon), Gilbertiodendron (c 30; tropical and subtropical Africa), Didelotia (10–11; Central Africa), Anthonotha (c 30; tropical Africa), Oddoniodendron (6; along the Gulf of Guinea), Englerodendron (1; Usambara Mountains in Tanzania), Isoberlinia (5; tropical Africa; polyphyletic), Berlinia (17; tropical Africa), Microberlinia (2; coasts of the Gulf of Guinea), Brachystegia (25–30; C tropical Africa to Botswana), Icuria (1; C Mozambique), Aphanocalyx (14; tropical West and Central Africa), Julbernardia (11; tropical to S Africa), Bikinia (c 10; W Central Africa), Tetraberlinia (7; Central Africa); Cryptosepalum (12; tropical Africa), Humboldtia (7; southern India, Sri Lanka), Paramacrolobium (1; tropical Africa), Polystemonanthus (1; tropical West Africa); Tamarindus (1; tropical Africa), Loesenera (2; tropical West Africa), Talbotiella (5–8; tropical West Africa), Leonardoxa (1; Cameroon), Hymenostegia (3; coast of the Gulf of Guinea); Amherstia (1; Burma), Michelsonia (1; E Congo), Pseudomacrolobium (1; Congo).

14. Brachycylix (Harns) R. S. Cowan. Trees. Only one sp., B. vageleri (Harms) R.S. Cowan, N & C Colombia, in tropical lowland rain forest, often riverine; used for ornamentals and timber.

15. Brownea Jacq. Trees or shrubs. 23 spp., W South America in Venezuela, Colombia, Peru and Ecuador with two spp. more widely distributed to Brazil (none endemics), the Guianas and two extending to C America (Panamá and Costa Rica) and the Caribbean, in understorey in tropical lowland rain forest. Several species (palo de cruz, bois rose, palo rosa, monta del rosa) are used for timber, medicine, handicrafts (from dried pods and seeds) and cultivated as ornamentals.

16. Browneopsis Huber. Trees. 7 spp., W S America in Peru, Ecuador, Colombia and Acre state in N Brazil (only B. peruviana (J.F. Macbr.) Klitg., not endemic), principally tropical lowland rain forest (terra firme and inundated), but also in sub-montane forest on the foothills of the Andes; used for timber.

17. Crudia Schreb. Trees; leaves imparipinnate; leaflets ± 7–10, alternate, entire; petiolules twisted; inflorescences terminal or lateral, lax but many-flowered racemes; flowers bisexual, small, arranged in 2 rows on long, slender pedicels; petals absent; seeds usually one or 2, large, orbicular or kidney-shaped. c. 50 spp., tropical lowland rain and swamp forest, thicket and scrub forest, often along rivers, most diverse in Asia, particularly Malesia (c. 30 spp., extending to Papuasia) and c. 3 spp. endemic to Indo-China, Andaman-Nicobar Islands and Sri Lanka; 10 spp. in S America (c. 9 spp. in Amazonia and one sp. each in the Caribbean and C America, 6 in Brazil, none endemics); c. 7–9 spp. in W and WC Africa (Senegal, Guinea, Sierra Leone, Liberia, Ivory Coast, Ghana, Nigeria, Cameroon, Equatorial Guinea, Gabon, Congo (Kinshasa) and Angola). The timber is extremely hard and heavy, with limited known uses.

18. Cynometra L. Trees, rarely shrubs; leaves paripinnate; leaflets 2–30, opposite; inflorescences many-flowered racemes or panicles, axillary or sometimes terminal, flowers small, arranged in more than 2 rows, sometimes articulated at apex of pedicel; bracteoles 2, caducous; pod indehiscent or dehiscent, sometimes swollen; seeds one or 2 (–4). c. 26–28 spp. in Africa (Guineo-Congolian W and WC regions (c. 16), Zanzibar-Inhambane E Africa (c. 9) and one sp. each in the Sudanian and Lake Victoria regions); 10 spp. endemic to Madagascar and 2 spp. to the Comoros; c. 26 spp. in SE Asia (c. 22 spp. centred in Malesia and Papuasia to Indo-China and Australia and the Pacific, c. 4 spp. endemic to SW and E India); 24 spp. in the Neotropics (c. 7 spp. in Mexico, Caribbean and C America; c. 17 spp. in S America centred in Amazonia, 12 in Brazil, 4 endemics); tropical lowland rain and swamp forest, often along rivers and sublittoral; seasonally dry forest, woodland, bushland or thicket, often on white sands; some species grow gregariously forming dominant stands.

Various species used for timber (kekatong, guapinol negro, nganga, baraka, muhimbe) in construction, ships keels, railway sleepers, flooring and carpentry; also used for firewood, as shade trees and bee forage.

19. Dicymbe Spruce ex. Benth. Trees, making ECM symbioses with fungi. 20 spp. from Guiana Shield, one up to Colombia, and one from this area to middle Amazon Brazil (9 in Brazil, 6 endemics), often monodominant in tropical lowland rain forest, gallery forest, in valleys and (rarely) on rocky slopes; used for timber.

20. Ecuadendron D. A. Neill. Trees. Eperua-like. Only one sp., E. acosta-solisianum D.A. Neill, W Ecuador, lower Andean foothills, trropical lowland wet to moist forest (with considerable fog-associated precipitation).

21. Elizabetha R.H.Schomb. ex Benth. Trees. 11 spp., SE Colombia, SW Venezuela, Amazonian Brazil (9, one endemic) to the interior of Guyana and Suriname, tropical lowland rain forest (terra firme and inundated) often along rivers.

The bark of E. princeps Schomb. ex Benth. is burned to produce an ash mixed with snuffs or hallucinogens (known as ebena), prepared from the resin of the genus Virola Aubl. (Myristicaceae).

22. Heterostemon Aubl. Small trees or shrubs. 8 spp., endemic to the Guiana Shield of Colombia to French Guiana, N Brazil (4, none endemics), 100 – 300 m. elevation range, in tropical lowland (sometimes inundated) rain forest, understorey and forest margins, often riverine and on sandy soils; used as ornamentals and for timber.

23. Macrolobium Desf. Trees or shrubs. 76 spp., centred in S America (73 spp., most diversity in Amazonia with two spp. to the Atlantic coast in SE Brazil, also west from Venezuela, Columbia, Peru to Ecuador in foothills on either side of the Andes); 5 up -or in - to in C America and Caribbean (Trinidad & Tobago, Costa Rica, Honduras and Panamá); tropical lowland rain forest, often along rivers and in seasonally inundated places, or in seasonally dry woodland and wooded grassland; used for timber, fish poisons and as ornamentals; 37 spp. in Brazil, 15 endemics. M. archeri Cowan from W Colombia and Ecuador possibly has the largest leaves among Fabaceae. M. acaciifolium (Benth.) Benth. from northern South America is myrmecohyte.

24. Paloue Aubl. Trees and shrubs. 5 spp., Amazonian Brazil (4, none endemic) Guyana, Suriname, French Guiana, in tropical lowland rain forest (terra firme and inundated), often along rivers.

25. Paloveopsis R. S. Cowan. Trees. Only one sp., P. emarginata R.S. Cowan, Guyana and Amazonas state in Brazil; tropical lowland rain forest (terra firme), often along rivers.

3. SUBFAMILY DIALIOIDEAE (16/c 83) ▸ oustiders Petalostylis (2; drier regions in Australia), Labichea (14; Australia except central parts), Baudouinia (6; Madagascar), Distemonanthus (1; tropical West Africa), Storckiella (4; NE Queensland; New Caledonia; Fiji), Zenia (1; S China, Thailand, Vietnam), Kalappia (1; Malili in Sulawesi), Eligmocarpus (1; SE Madagascar), Koompassia (3; Malesia), Mendoravia (1; SE Madagascar).

26. Androcalymma Dwyer. Tree up to 30 m tal, yellow flowers. Only one sp., A. glabrifolium Dwyer, fro upper Amazon Basin in Brazil, a rare plant in Brazil, by Plantas Raras do Brasil’s book, known only in São Paulo da Olivença municipality.

27. Apuleia Mart. Trees up to 50 m tall. Only one sp., A. leiocarpa (Vogel) J.F. Macbr., widely distributed across S America in Amazonian and extra-Amazonian Brazil, Venezuela, Peru, Paraguay, Bolivia and Argentina, in a wide range of tropical and subtropical habitats, from rain forest, riverine and gallery forest to dry seasonal scrubland of NE Brazil (caatinga). The timber is used commercially for, e.g., high quality furniture, heavy construction and flooring.

28. Dialium L. Trees up to 30 m tall. 40–70 spp., pantropical (from 19°N in S Mexico and Belize to 23°S in Madagascar) with one sp. extra-tropical in Mozambique and S Africa; not extending E of Wallace’s Line (totally absent from Australia and New Guinea); main centres of diversity in WC Africa (14; 3 also in E coastal Africa, 3 endemic to Madagascar) and W Malesia (4 in S Thailand, Laos, Cambodia and S Vietnam, 2 in Borneo, Sumatra and Malay Peninsula, 1(–2) spp. in S India and Sri Lanka), and 4 sp. widely distributed in the Neotropics (all in South America) from S Mexico through C America to the Amazon and Atlantic forest in Brazil (3, one endemic), extending into Colombia, Peru, Ecuador and Bolivia, in tropical rain forest (mainly lowland evergreen), occasionally in peat, swamp or heath forest, or coastal; a few species in monsoon forest and wooded grassland (savanna), but in drier vegetation types the species frequently occur on river margins or deeps.

The edible fruits are used mainly in chutneys, the timber of various species (keranji, jutahy) is used for boat building, house construction, flooring, tools, firewood and charcoal (some species have a high silica content and resist borer attack). The bark and leaves are widely used in folk medicine in W Africa (e.g., for fevers and toothache); the bark of D. cochinchinense Pierre is a substitute for betel nut in Indochina; the bark of D. pachyphyllum Harms has a poisonous red gum-resin used for arrow-poison in Congo (Kinshasa).

29. Dicorynia Benth. Trees. Two spp., D. guianensis Amshoff exclusive from Guianas and D. paraensis Benth. in Amazonian Brazil (especially common along the Rio Negro) and just extending to the Colombian-Venezuelan border, essentially a riparian genus, typically of inundated rain forest or swamp forest (várzea), but also on terra firme and in ‘evergreen seasonal forest’ on well-drained sites; below 600 m.

The excellent, durable timber of D. guianensis (angelique, basralocus) has been widely used for furniture and cabinet making, and in heavy construction, general carpentry, railroad ties and cooperage, and especially in marine construction due to its high silica content which results in a high natural resistance to wood borers.

30. Martiodendron Gleason. Trees. 5 spp., S America in the Guianas, Amazonian Brazil (Roraima, Para, Amazonas, Acre states), S Venezuela, and NE Brazil (Bahia, Piauí, Maranhão), in tropical rain forest (often periodically inundated) to seasonally dry forest and wooded grassland (savanna), below 600 m.; all species occur in Brazil, one endemic.

31. Poeppigia C. Presll. Trees. Two spp., P. procera C. Presl, Mexico, Honduras, Guatemala, El Salvador, Cuba, Venezuela, Colombia, Peru, Bolivia, circum-Amazonian Brazil (inc. Acre, Rondônia, Mato Grosso, Minas Gerais, Bahia, Pernambuco and Piauí), in dry seasonal scrubland of NE Brazil (caatinga); locally used for construction timber, firewood and medicine; and P. densiflora Tul., endemic to dry seasonal scrubland of NE Brazil (caatinga).

4. SUBFAMILY FABOIDEAE (460/14,315–14,700) ▸ 25 lineages, 18 in South America; Dermatophyllum clade (1/6, U.S.A., Mexico), Baphieae (5/58–63, tropical Africa to E Asia), Hypocalypteae (1/3, Western and Eastern Cape), Mirbelieae (32/715 - 745, Australia, with their highest diversity in southwestern and southern parts), Disynstemon clade (1, southwestern Madagascar), Wisterieae (3/14, Eastern North America, East, South and SE Asia, New Guinea, Australia, New Caledonia), Hedysareae (14/475–520, Temperate regions in Eurasia, northern Africa to Ethiopia) are lineages (all in tribal level) are the eight absents in South America.

4.1 FABOIDEAE ▸ CLADRASTIDOIDS CLADE (3/18) – outsider Cladrastis (9; E Asia, SE U.S.A.), Pickeringia (1; California)

32. Styphnolobium Schott. 8 spp., one from E Asia, and 7 from Mexico to Costa Rica, with S. sporadicum M. Sousa & Rudd up to NW Colombia.

4.2 FABOIDEAE ▸ TRIBE SWARTZIEAE (8/c 215) - outsider Bobgunnia (2; tropical and southern Africa).

33. Ateleia (DC)Benth. Trees and shrubs. c. 24 spp., Mexico, C America & Caribbean (c. 17), 4 in S America (Venezuela to N Argentina; 3 spp. in Brazil, none endemics); one sp. widely distributed, in tropical to subtropical seasonally dry forest, woodland, bushland and xerophytic shrubland, often in rocky areas or riverine; one sp. of high humid forest. Used for timber, insecticides and wildlife-food.

34. Bocoa Aubl. Trees. 4 spp., S America (one sp. from French Guiana and Surinam, 3 in Brazil, one in over Amazon rainforest, one from Pará and Amazonas extending to French Guiana and Surinam, the other recorded only from Maranhão - this as a rare plant in Brazil, by Plantas Raras do Brasil’s book, in tropical Amazonian (non-inundated) rain forest (2), savannas of C Brazil (cerrado) and bushland (1). Used for timber (furniture, musical instruments), medicine, cosmetics and considered to have magical powers

35. Candolleodendron R. S. Cowan. Trees. Only one sp., C. brachystachyum (DC.) R.S.Cowan (saboneteira) from S America (French Guiana, N and NE Brazil), in tropical Amazonian (non-inundated) rain forest; now considered to be closest to Bocoa and Swartzia.

36. Cyathostegia (Benth.) Schery. Trees and shrubs. Two spp., inter-Andean valleys of S Ecuador and N Peru, in seasonally dry tropical bushland and shrubland, often on rocky hillsides.

37. Swartzia Schreb. Shrubs, trees and lianas, up to 30 m tall. 195 spp., Mexico, C America, Caribbean (c. 10); S America (182, to SE Brazil), in tropical rain forest and seasonally dry forest, wooded grassland, often riverine, to high mountain slopes, dry sandy plains and rocky beaches. from tropical rain forest and seasonally dry forest, wooded grassland, often riverine, to high mountain slopes, dry sandy plains and rocky beaches. 112 spp. in Brazil, 59 endemics - 12 of them, from several states, are rare plants in Brazil, by Plantas Raras do Brasil’s book.

Used for timber (Swartzia species variously known as bastard rosewood, wamara, bannia, coração de negro, are used for inlay, parquet flooring, turnery, furniture, cabinetwork, violin bows, specialty items; suggested as a substitute for ebony); also used in medicine. 111 spp. in Brazil, 62 endemic.

38. Trischidium Tul. Trees. 5 spp., all spp. in Brazil (3 endemics), one sp. extending to Paraguay. one sp. to Guyana, Bolivia and Peru, in tropical Amazonian (non-inundated) rain forest (2), coastal forest (1) and seasonally dry woodland and bushland (dry seasonal scrubland of NE Brazil (caatinga) and savannas of C Brazil (cerrado)).

4.3 FABOIDEAE ▸ TRIBE DIPTERYGEAE (17/c 80) - three lineages, all in South America.

∎ ANGYROCALYCIEAE CLADE (4/11) ▸ outsiders Angylocalyx (6; tropical Africa), Xanthocercis (3; Gabon and Zambia to S Africa, Madagascar), Castanospermum (1; W New Britain, E Queensland, E New South Wales, New Caledonia, Vanuatu).

39. Uleanthus Harms. Small trees up to 15 tall. Only one sp., U. erythrinoides Harms, from small area in Amazonas state near Manaus up to center Pará state, in tropical lowland forest, along margins of rivers and waterfalls. This genus has never been included in any comprehensive molecular or morphological analyses; it has some inflorescence and floral features in common with Angylocalyx, but synapomorphies to assert relationships with any other genera are as yet lacking

∎ DYPTERIGIEAE CLADE (4/19) ▸ all genera in South America.

40. Dipteryx Schreb. Trees up to 40 m tall. 15 spp., mostly Amazonian (4 in Venezuela); 2 spp. up to C America (Panamá to Honduras); one sp. to drier areas of C Brazil (13, 4 endemics), E Bolivia and Paraguay, in tropical rain forest (riverine and usually non-inundated) to seasonally dry forest (some on white sand) and savannas of C Brazil (cerrado).

A tiroup of c. 3 species produce fragrant coumarin-yielding seeds, e.g., D. odorata (Aubl.) Willd. (cumaru, tonka bean, sarrapia, almendro) is a major source of coumarin whose vanilla-like fragrance is used for scenting tobacco, snuff and confectionery, and is an ingredient of perfumes and cosmetics; the beans also yield a high percentage of solid fat or tonka butter which is used to flavour food; plants produce balsam resins (oleoresin) and red gums from the leaves, stems and bark; one species bears pods with a sweet, edible pericarp; the hard wood is generally used for heavy construction, flooring, shafts, bearings, veneers and in craft objects; also used for charcoal and medicine

41. Monopteryx Spruce ex. Benth. Trees. 3 spp., Colombia, Venezuela, French Guiana and NW Amazonian Brazil (all species, none endemics), in non-inundated tropical lowland rain forest, on sandy soil; potentially useful wood for furniture and construction; the wood is resinous and has a balsamic smell; seeds edible when roasted

42. Pterodon Vogel. Trees. 4 spp. from EC states of Brazil, one up to and E Bolivia, in seasonally dry tropical forest, savannas of C Brazil (cerrado) and dry seasonal scrubland of NE Brazil (caatinga), often associated with rocky outcrops. The very hard wood is suitable for items requiring durability; also used for medicine, firewood, fodder, as ornamentals and for reforestation

43. Taralea Aubl. Trees and shrubs. 4 spp., Amazonian S America in Venezuela (4), the Guianas, Amazonian Brazil (3, none endemics), Columbia and Peru, in tropical lowland rain forest (riverine and non-inundated or inundated plains) to seasonally dry forest (sometimes on white sand), woodland and marshes; also 1–2 spp. in montane forest and open moist woodland to scrub on sandstone.

The very hard wood is suitable for construction and firewood; T. oppositifolia Aubl. (cumaru-da-praia) contains an unscented oil extracted for industrial use.

∎ AMBURANEAE CLADE (8/29) ▸ outsdiers Dupuya (2; Madagascar), Cordyla (6; tropical Africa to South Africa, Madagascar).

44. Alexa Moq. Trees up to 40 m high. 10 spp., Venezuela, Guyana, Surinam, French Guiana, Amazonian Brazil (5, one endemic), in tropical lowland (occasionally upland) rain forest. A. imperatricis (R.H.Schomb.) Baill. (haiari, haiariballi) is used as a commercial timber (construction, crating and plywood); species’ also used for fish poisons and medicine. A. cowani Yakovlev, from Venezuela and Guyana, is a myrmecophyte.

45. Amburana Schwacke & Taub. Trees up to 40 m tall. Three spp., non-Amazonian Brazil (all species, one endemic), Peru, Bolivia, Paraguay, N Argentina, in seasonally dry tropical to subtropical forest (often on rocky hillsides or in ravines), bushland and dry seasonal scrubland of NE Brazil (caatinga). Used for timber, e.g., A. cearensis (Allemão) A.C.Sm. (cherry wood, amburana, cerejeira, ishpingo), for furniture, construction, frames, cabinetry, veneers and carpentry; also used as medicine

46. Dussia Krug & Urb. ex Taub. Trees. 11 spp., c. 4 in S Mexico, C America and Caribbean and 7 in S America in Venezuela, Colombia and the Guianas to C Peru and Amazonian Brazil (2, none endemics), in tropical lowland and upland rain forest. Used for timber (local construction); the blood-red sap is sometimes used as medicine

47. Myrocarpus Allemao. Trees up to 35 m tall. 5 spp., Venezuela, S and E Brazil (4, one endemic and rare in Brazil, by Plantas Raras do Brasil’s book, restricted from São Paulo state), Bolivia, Paraguay and N Argentina, in wet to seasonally dry tropical lowland forest and woodland. Used for timber (durable, fragrant and of good quality for construction, furniture and wood carving); essential oils (balsam) from the bark (e.g., of M. frondosus Allemão (cabreuva)) are used in aromatherapy, also as medicine, perfumes and incense.

48. Myrospermum Jacq. Trees and shrubs. Two spp., N & C Mexico to Colombia and Venezuela, in seasonally dry tropical to subtropical woodland and bushland, on stony hillsides and along rivers. Used (sparingly) for timber

49. Myroxylon L. f. Trees up to 30 m tall. Two spp., both widerspread, from Mexico to western and northern S America to N Argentina, both in Brazil, in tropical lowland rain forest to seasonally dry forest and woodland. Used for timber (furniture, turnery, interior trim and construction); balsam extraction for medicine, perfumery, thickening agents, etc. (cultivated in e.g., Sri Lanka and W Africa)

50. Petaladenium Ducke. Tree 8–30 m tall; bark grey brown to pale brown, smooth or flaking slightly, deeply vertically fissured and often with lenticels; slash pale redbrown, producing small quantities of thick, red exudate; twigs pubescent, lenticellate; stipules not seen; leaves 5–9(–11)foliolate, imparipinnate; inflorescenc 9–22 cm long, ramiflorous or axillary in simple lateral 1–4branched racemes or panicles. Only one sp., P. urceoliferum Ducke, endemic to the Guiana Shield of Rio Negro in N Brazil (in a very small dense forest in NW Amazonas state in Brazil, at 100 m elevation range; it is unique among the whole legume family (except Duparquetioideae) in having fimbriate-urceolate-glandular wing petals.

4.4 FABOIDEAE ▸ TRIBE EXOSTYLOIDEAE (6/21) - all genera in South America.

51. Exostyles Schott. Trees or shrubs. 4 spp., one sp. EC Brazil, two spp. SE Brazil, both from Atlantic Forest of Brazil (non-inundated Atlantic coastal forest and Atlantic sandy coastal shrublands (restingas), and seasonally dry tropical submontane forest), and one in Amazonian Brazil and adjacent Suriname. Used for timber

52. Harleyodendron R. S. Cowan. Shrubs or often small tress 5-10 m tall, the thunks slender, the bark pale gray. Only one sp., H. unifoliolatum R. S. Cowan, endemic to tropical coastal forest on steep rocky slopes of Atlantic Forest of Bahia state in NE Brazil.

53. Holocalyx Michelli. Trees. Only one sp., H. balansae Micheli, from Paraguay, S Brazil, Bolivia and Argentina and inhabits most forests or gallery forets in savannas of C Brazil (cerrado). Used for its resistant and durable timber (for construction, beams, furniture and carpentry), as ornamentals and shade trees, but the leaves are toxic (containing cyanogenic glycosides).

54. Lecointea Ducke. Trees. 5 spp. in Colombia, Venezuela, N Brazil (3, one endemic), Ecuador and Peru, one up to Mexico, in tropical Amazonian inundated forest to seasonally dry, riverine and hillside forest.

55. Uribea Dugand & Romero. Trees. Only one sp., U. tamarindoides Dugand & Romero, from Costa Rica, Panamá and Colombia, in tropical lowland rain forest.

56. Zollernia Wied-Neuw. & Nees. Trees and shrubs. 10 spp. from S America, one sp in Brazil, Venezuela and Guianas; 8 endemics to NE to SE Brazil from Amazon basin south to Santa Catarina State (two of them, from Rio de Janeiro and Minas Gerais states, are rare plants in Brazil, by Plantas Raras do Brasil’s book), and one in Cono Sur, in tropical Amazonian rain forest and seasonally dry forest, savannas of C Brazil (cerrado) and dry seasonal scrubland of NE Brazil (caatinga) on Atlantic sandy coastal shrublands (restingas). Used as timber; potential as ornamentals

4.5 FABOIDEAE ▸ TRIBE ORMOSIOIDEAE (4/c 140) - all genera in South America.

57. Clathrotropis (Benth.) Harms. Trees. 6 spp., Caribbean, W and N S America in Peru, Colombia, Venezuela, Surinam, Guyana, French Guiana and Brazil (2, no endemics), in tropical lowland (often riverine and seasonally inundated) rain forest, occasionally upland or montane forest; also seasonally dry woodland and shrubland.

Used as timber, e.g., C. brachypetala (Tul.) Kleinhoonte (aromata; alma negra) for construction, boat building, furniture, flooring and railway sleepers); also used for medicine and a mammal poison

58. Ormosia Jacq. Trees and shrubs up to 40 m tall. 132 spp., 65 in Mexico, C America, Caribbean, northern and western S America (55) to Brazil (35, 16 endemics) and Bolivia; 50 spp. in SE Asia (India, Indo-China, China, Malesia, Micronesia, Solomon Is.) and Australia; tropical rain forest (sometimes riverine) or seasonally dry forest, bushland, shrubland and xerophytic woodland, on sand over limestone.

The seeds are used as beads A number of species (variously known as tento, kokriki, sirari and huayruru) are used for their durable timber (construction, furniture, carpentry and veneer); also used for medicine and ornamentation (bright red seeds used as necklaces and charms; species variously known as lady bug tree, jumby bead or necklace tree) . O. macrophylla Benth. from northern South America is a myrmecophyte.

59. Panurea Spruce ex. Benth. Trees. Two spp., endemic to the Guiana Shield of Colombia and Amazonas state in N Brazil, at 100 m elevation range, in tropical lowland forest in north-amazonic white-sand savannas (campinaranas).

60. Spirotropis Tul. Trees and lianas. Only one sp., S. longifolia (DC.) Baill., endemic to the Guiana Shield of Venezuela to French Guiana, 400 - 800 m elevation range, in tropical lowland to montane, riverine forest

4.6 FABOIDEAE ▸ TRIBE GENISTOIDEAE (62–70/2.265–2.280) - 10 lineages, Camoensieae (1/2, coasts of the Gulf of Guinea and south to Angola), Thermopsideae (5/61–62, Macaronesia and Mediterranean to Central and E Asia, North America), Euchresteae (1/4, India, the Himalayas, China (inc. Taiwan), Korean Peninsula, Japan, SE Asia, Java), Podalyrieae (9/138–152, South Africa, few species to NE Africa and Arabian Peninsula) and Oberholzeria (1/1, Namibia) absents in South America.

∎ SUBTRIBE BRONGNIARTIEAE (14/c 160) ▸ outsiders Behaimia (1; Cuba), Hovea (c 40; Australia), Templetonia (11; Australia), Plagiocarpus (1; N Australia), Cristonia (1; W Australia), Lamprolobium (2; NE Queensland), Thinicola (1; Western Australia).

61. Amphiodon Huber. Trees up to 20 m tall; leaves alternate, pulvinate; stipules minute, caducous; corolla papilionaceous, petals 5, glabrous, clawed; the wing petals slightly longer than the keel, their upper base auriculate; keel petals weakly adherent along part of their lower margins. Only one sp., A. effusus Huber, Amazon Brazil, N Bolivia, E Peru, Suriname and French Guiana, widely distributed in Amazonian non-flooded forests (terra-firme), especially in secondary forests and clearings.

62. Brongniartia Kunth. Shrubs os small trees. 52 spp., tropical and subtropical America, amphitropical and somewhat disjunct: SC U.S.A. (only one, B. minutiflora S. Watson, in Texas), N and C Mexico (50 endemics, almost restricted from Pacific Coast up to center country), with two in W Andean of Bolivia (one endemic) and Chile (disjunct with Mexico), in tropical to warm-temperate (often montane) forest, woodland, bushland, thicket, shrubland and grassland, mostly in dry areas. Used as ornamentals.

63. Cyclolobium Benth. Small trees. Only one sp., C. brasiliense Benth. in Brazil (from Rondônia and Goiás east to Bahia and south to São Paulo, Mato Grosso do Sul and Paraná), Paraguay (Cordillera de Altos and Ybycuí National Park), Bolivia (Santa Cruz)), in seasonally dry tropical forest, often along riverbanks. Used as ornamentals.

64. Harpalyce Sessé & Moc. ex DC. Shrubs or small trees, sometimes with xylopodium; large peltate glands on many parts of the plant, largely resupinate flowers, strongly bilabiate calyx and helically contorted keel (secondarily straightened in 3 to 4 species), orange to red petals, the keel helically contorted through 180º, dimorphic anthers, and compressed, elastically dehiscent pods with seed chambers delimited by transverse septa. 38 spp. in seasonally dry tropical to warm-temperate humid forest, woodland, bushland and thicket, shrubland and grassland; most species are evergreen and flower during the dry season; the wood is used for small turnery item. Three disjunct secions:

§ sect. Brasilianae ▸ 12 spp. from C & C Brazil, one up to Bolivia, two of them, from Minas Gerais and Bahia state, are rare plants in Brazil, by Plantas Raras do Brasil’s book.

§ sect. Harpalyce ▸ 9 spp. in Mexico and Mesoamerica.

§ sect. Cubensis ▸ 17 spp. endemics to Cuba.

65. Limadendron Meireles & A. M. G. Azevedo. (off Poecilanthe) Trees; leaves alternate, pulvinate, unifoliolate or very rarely mu1tifoliolate with 3–7 clearly opposite leaflets; inflorescences solitary racemes, cauliflorous or axillary, pendant; flowers bilaterally symmetrical, pedicellate; corolla papilionaceous, petals 5, glabrous, unguiculate; legume dehiscent with woody valves. Two spp., L. amazonicum (Ducke) Meireles & A. M. G. Azevedo (Brazilian and Venezuelan Amazonian forests, especially in sandy soil along black water, seasonally flooded forest (igapó) and in shrubby riparian vegetation), and L. hostmannii (Benth.) Meireles & A. M. G. Azevedo (Amazonian forest in Brazil (Amapá and Amazonas), Colombia, Guyana, French Guiana, and Surinam, especially along small rivers, ‘igarapes’, (but in areas that are not susceptible to seasonal flooding) and in ‘‘terra firme’’ forests (also non-flooded).)

66. Poecilanthe Benth. (exc. Limadendron) Trees and shrubs. 10 spp., S America, mostly Brazil (8, 5 endemics), Surinam, Bolivia, in tropical lowland rain forest (Amazonian basin species group); seasonally dry tropical forest to woodland and thicket, or riverine forest (E Brazil to Uruguay and Argentina species group). Used as timber, medicine, anti-fungal agents and ornamentals.

67. Tabaroa L. P. Queiroz, G. P. Lewis & M. F. Wojc. Tree 5-8 m, canopy widely elliptic c. 3 – 5 m diam., foliage deciduous in dry season; Inflorescence a terminal, almost pendent, pyramidal panicle; comprising 10 – 16 racemose inflorescence branches; f lowers 4.5 – 5 mm long, lemon-scented, zygomorphic, papilionoid, hypanthium short, campanulate; fruit indehiscent. Only one sp., T. caatingicola L. P. Queiroz, G. P. Lewis & M. F. Wojc., known only from a very narrowly restricted area in Southwestern Bahia (at c. 12 km²), Brazil, on the lower slopes of the Rio de Contas mountain range.

∎ SUBTRIBE LEPTOLOBIEAE (6/27–32) ▸ all genera in South America.

68. Bowdichia Kunth. Trees and shrubs. Two spp., Colombia, Venezuela, Guyana, Surinam to C Brazil (both spp., none endemics) in tropical, seasonally dry lowland wooded grassland, grassland and bushland, occasionally lowland forest. B. nitida Benth. (sucupira) is an important timber used in construction (flooring, beams and frames), also used for charcoal and as fish poisons

69. Diplotropis Benth. Trees and shrubs up to 30 m tall. 12 spp., S America, mostly Amazonian basin in Colombia, Venezuela, Surinam, Guyana, French Guiana, Brazil (8, 5 endemics - one of them, from Amazonas states, is a rare plant in Brazil, by Plantas Raras do Brasil’s book), Ecuador, Peru and Bolivia, in tropical rain forest (riverine, inundated and non-inundated) to seasonally dry forest, wooded grassland and bushland. Used for timber, e.g., D. purpurea (Rich.) Amshoff (sucupira; tatabu; botonallare) in heavy construction, shipbuilding, flooring, turnery and manufacture of furniture, tool handles and railway sleepers); also used as a mammal poison.

70. Guianodendron Schutz Rodrigues & A. M. G. Azevedo. Trees to 35 m, with obtriangular crown, base buttressed, trunk to ca. Leaves alternate, imparipinnate, 5- to 9- foliolate; inflorescence paniculate, axillary or terminal, usually 2 or 3 panicles forming an axillary fascicle. Flowers 5–6 mm long; hypanthium straight, 1–1.3 mm long; corolla actinomorphic, petals 5, white, shortclawed. Only one sp., G. praeclarum (Sandwith) Sch. Rodr. & A.M.G. Azevedo, occuring in Guyana and N Brazil in different forest types (wallaba forest, Mora forest, mixed lowland forests, and marsh forest), which range from high-canopied forests (canopy at 30–45 m) to lower-canopied forests (canopy at 10–17 m), usually on sandy, alluvial, loamy, and periodically inundated or well-drained soils; also north-amazonic white-sand savannas (campinaranas) from upper Rio Negro, on sandy and wet soils, in the base of the Guiana Shield. The wood is used for house posts and furniture

71. Leptolobium Vogel. Trees and shrubs. 11 spp., L. panamense (Benth.) Sch. Rodr. & A.M.G.Azevedo from S Mexico to Venezuela, and 10 in Brazil (8 endemics and two up to adjacent Bolivia and Venezuela); L. panamense (Billy Web, carboncillo) is an important hardwood noted for its strength and durability; the inner bark (cascara amarga) contains an alkaloid used for medicine and as na anti-malarial; it is also the main ingredient of ‘Sweet Blood’, a commercial remedy sold for diabetes.

72. Orphanodendron Barneby & J.W.Grimes. Trees. Two spp., both endemic to Andean Colombia, possibly placed within Papilionoideae.

73. Staminodianthus D.B.O.S. Cardoso, H.C. Lima, L.P. Quieroz. Small trees to 6 m tall, to large trees to 40 m; leaves imparipinnately compound; inflorescence an axillary raceme or a terminal, congested corymbose panicle; flowers bilaterally symmetrical; petals 5, entirely pinkish, glabrous, crimped or nearly so. Three spp. from the Amazon basin of northern Brazil (all spp., one endemic) and adjacent areas in SE Colombia, S Guyana, NE Peru, and S Venezuela, in gallery forests, highland savannas, non-flooded terra-firme forests on sandy or sand-loam soils

∎ SUBTRIBE SOPHOREE (6–11/64–76) ▸ outsiders Ammodendron (4–5; W and C Asia to NW China), Ammothamnus (2; C Asia), Maackia (8; E Asia), Salweenia (1; SE Tibet), Amphimas (3–4; W C Africa).

74. Sophora L. Unarmed herbs, shrubs or small trees; leaves once odd-pinnately compound; stipules very small, falling early. 45 spp., SE Europe to W, C & E Asia and south through tropical regions to Australasia and the Pacific; 15 spp. in New World, 9 in South America, c. 3–4 spp. in sect. Edwardsia (Chile, Argentina and Juan Fernandez Is.), only the wider S. tomentosa L. in coastal E Brazil; largely introduced in Africa; one sp. endemic from coastal Kenya south to S Africa and Madagascar; in seasonally dry tropical to warm temperate lowland and upland forest or dry vegetation types and sand dunes.

Cultivated as ornamentals (in sect. Edwardsia, with yellow flowers, e.g., S. tetraptera J.F.Mill. (kowhai) and S. microphylla Aiton (small-leaved kowhai); and in sect. Pseudosophora, with blue to white flowers, e.g., S. davidii(Franch.) Skeels); also used for its durable timber (for bearings, turnery and cabinet work), as medicine (e.g., S. flavescens Aiton (ku shen), S. tonkinensisGagnep. (shan dou gen) and S. microphylla); some species are toxic and used as insecticides. number are cultivated as ornamentals including the Chinese and Korean S. japonica L.

∎ SUBTRIBE CROTALARIEE (19/1.320–1.325) ▸ hardly South Africa, few species in E and NE Africa and SW Asia, only Crotalaria in New World.

75. Crotalaria L. Annual or perennial herbs with stems sometimes winged above, sometimes also with xylopodium; leaves sessile or subsessile, apparently simple in our species; flowers usually in axillary or terminal, 2 – many flowered racemes, long-pedicelled. 600 spp., most spp. in Africa and Madagascar (c. 510, c. 34 of which endemic to Madagascar and 5 widely distributed in Old World); c. 18 widely distributed in tropical Asia; c. 60 spp. endemic to the Indian subcontinent; c. 5 spp. to W Asia, c. 15 spp. to Indo-China, c. 12 spp. to China, c. 6 spp. to Malesia and 9 spp. to Australia; 60 spp. endemic (and a further 15 introduced) in the New World, 37 in S America (mainly Brazil with 31, 16 endemics - 4 of them, from Bahia, Goiás and Minas Gerais states, are rare plants in Brazil, by Plantas Raras do Brasil’s book), 20 spp. in N and C America (mainly Mexico) and c. 5 spp. widely distributed in New World. Some are used as sources of fodder or fiber.

∎ SUBTRIBE GENISTEAE (c 6/c 500) ▸ outsiders Cytisus (c 140; Europe, Africa, Asia), Genista (c 130; Europe, Macaronesia, Mediterranean, Asia), Adenocarpus (c 15; Canary Islands, Mediterranean, tropical African mountains).

76. Anarthrophyllum Benth. Shrubs, sometimes cushions. 15 spp. from Andes in Argentina and Chile, in dry shrubland, bushland and grassland in cold parts of the Andes, on sandy soils and in rocky valleys. Used for forage, windbreaks and firewood

77. Lupinus L. Annual and perennial herbaceous species, as well as a few soft-woody shrubs and small trees. c. 110 spp. Mediterranean basin in Europe to Türkiye, Middle East and N Africa (c. 13), plus montane E tropical Africa (2); 484 spp. in New World, 331 in South America, mainly Colombia (86), Peru (171) and Bolivia (50), only 31 in Brazil, 21 endemics; some spp. are widely introduced elsewhere. Mostly open habitats, in disturbed places, in poor (often acid) soils.

Important as a domesticated human food crop; L. albus L. (white lupine, lupini bean) and L. luteus L. (yellow lupine) from Mediterranean and L. mutabilis Sweet (pearl lupine) from the New World are grown for edible, high protein seeds (also for flour); several species widely grown for livestock fodder, green manure, cover crops, fish poisons, medicine, oils and as attractive ornamentals (e.g., Russell hybrids of L. polyphyllus); a number of species are variably toxic and lupinosis causes death in animals, due largely to ingestion of foliage containing quinolizidine alkaloids

78. Sellocharis Taub. Shrub or herb; 5–7 leaflet-like structures in a whorl at the nodes, feature unique among Fabaceae. Only one sp., S. paradoxa Taub., collected at grasslands of Rio Grande do Sul state, S Brazil, known only from the type collection until re-collected recently in the Rio Grande do Sul region.

4.7 FABOIDEAE ▸ VATAIREOIDEAE CLADE (4/21) - all genera in South America.

79. Luetzelburgia Harms. Trees and shrubs, leaves imparipinnately compound and without stipels, flowers papilionoid with five petals sericeous outside. 14 spp., all national endemics in Brazil (11, two of them, from Bahia state, are rare plants in Brazil, by Plantas Raras do Brasil’s book), Bolivia (2) and Colombia (1); in seasonally dry tropical, lowland woodland and wooded grassland, occasionally lowland rain forest. Used for timber, fuelwood; roots milled for flour as a famine food.

80. Sweetia Spreng. Trees. Two spp., Bolivia, Brazil (only one, no endemic) and Paraguay, in tropical rain forest (sometimes riverine) and seasonally dry forest, bushland and thicket to subtropical forest; S. fruticosa Spreng. (mani) is considered a future crop tree for logging in Bolivia

81. Vataireopsis Ducke. Small to emergent trees up to 30 m tall. 4 spp., Amazonian S America from Ecuador to French Guiana to Bolivia and N Brazil (3, one endemic), in tropical lowland Amazonian and coastal Atlantic rain forest. Used medicinally in psoriasis treatment; in 1876 chrysarobine was extracted from V. araroba (Aguiar) Ducke and in 1916 a derivative, dithranol, was first synthesised; other species also used for treatment of leishmaniasis and dermatitis

82. Vatairea Aubl. Tall emergente trees up to 40 m tall. 8 spp., Neotropics, centre of diversity in Amazonia; 7 spp. confined to northern S America, one in C America, from S Mexico to Panamá; the majority of species inhabit tropical lowland rain forest, in seasonally flooded forest (igapó), V. macrocarpa Ducke occurs in seasonally dry forest, savannas of C Brazil (cerrado) and dry seasonal scrubland of NE Brazil (caatinga); 7 spp. in Brazil, two endemics. The majority of the species (known variously as faveira amargosa, angelim amargosa, amargo, bitter angelim or danto) produce good quality wood used for construction, furniture, cabinet work, flooring, joinery and veneers; also used for medicine

4.8 FABOIDEAE ▸ TRIBE ANDIREAE (3/c 47) - all genera in South America.

83. Aldina Endl. Trees, making ECM symbioses with fungi. 17 spp., endemic to the Guiana Shield of Colombia (1) to Guyana and N Brazil (8, 3 endemics), with 9 endemics to Venezuela, 100 – 1300 m elevation range, in tropical montane and lowland Amazonian (either inundated or riverine) forest, and seasonally in north-amazonic white-sand savannas (campinaranas).

Used for timber, e.g., A. heterophylla Spruce ex Benth. (macucu de paca, angelim da Campina) for heavy construction, sleepers, flooring, exterior panelling, joinery and fencing.

Although Aldina is a small genus, some species such as A. heterophylla Spruce ex Benth. and A. latifolia Spruce ex Benth. are among the most widely distributed angiosperms in Amazonian campina (sandy) and seasonally flooded forest (igapó), respectively. A recent ecological estimate listed A. heterophylla among the dominant 227 species from an assemblage of ca. 16,000 woody species that make up the Amazon forest, whereas ca. 11,000 species are rare or only narrowly distributed.

84. Andira Juss. Trees and shrubs, sometimes with xylopodium. 30 spp., Neotropics, most diverse in wet forest Amazonia (c. 12) and in the Brazilian Atlantic forest (8); c. 5 spp. in seasonally dry S America; 3 spp. in C America, Caribbean and Mexico; A. inermis (W.Wright) DC. widely distributed in the Neotropics and in WC Africa, in tropical rain forest (most species), seasonally dry forest, savannas of C Brazil (cerrado), Atlantic sandy coastal shrublands (restingas); often on stream-banks and in swampy areas. 17 spp. in South America, 20 in Brazil, 11 endemics, two of them, from Bahia state, are rare plants in Brazil, by Plantas Raras do Brasil’s book.

Used for timber (e.g., A. inermis subsp. inermis (cabbage tree, partridge wood or black plum), for construction, furniture, cabinet work, railway sleepers and fence posts), fish poisons, ornamentals, shade plants for coffee plantations and medicine (anti-helminthic drugs are obtained from the bark (known as worm bark) and seeds, but these are poisonous in high doses); also used as fungicides (Goa or Bahia powder). A. inermis (W.Wright) Kunth ex DC. and A. vermifuga Benth. from tropical New World, both in Brazil, are myrmecophytes.

85. Hymenolobium Benth. Trees, very tall up to 50 m tall, usually emergent in tropical humid lowland rain forest. 16 spp., concentrated in Brazil (15, 7 endemic), the Guianas, and Venezuela; but one extends into Peru, one into Ecuador, and one occurs only in C America (Panamá, Costa Rica, and Nicaragua); all species produce hardwoods (e.g., H. excelsum Ducke (para-angelim, angelim amarelo, angelim rosa or angelim da mata), is used for heavy and light construction, furniture, flooring, panelling, joinery and veneers)

4.9 FABOIDEAE ▸ TRIBE DALBERGIOIDEAE (53-54/1.520-1.525) - two subtribes, both in South America.

∎ SUBTRIBE AMORPHINAE (c 8/240–245) ▸ outsiders Amorpha (16; North America, with their highest diversity in SE U.S.A.), Parryella (1; Mexico), Eysenhardtia (12–15; Central America), Psorothamnus (9; arid regions in SW North America), Marina (c 40; SW U.S.A., Mexico, Central America).

86. Apoplanesia C. Presl. Small trees. Two app. one from S Mexico to Nicaragua and another endemic to Venezuela, seasonally dry tropical forest and woodland, hillsides, river valleys and roadsides; the wood of A. paniculata C.Presl (chulul) is very hard and used for cabinetry, crafts and bows; the bark yields a yellow dye; also used as ornamentals and in agroforestry.

87. Dalea Mill. (inc. Parosela) Unarmed herbaceous perennials or low shrubs; leaves once odd-pinnately compound; leaflets few to many, resin-dotted; stipules slender, falling early; flowers in terminal spikes, these often dense and cone-like; bracts often resin-dotted; Xerophytic herbs or shrubs. 191 spp., SW Canada, U.S.A., C America, Caribbean and south to Argentina, mostly Mexico (142, 102 endemics) and U.S.A. to Canada (c. 23 exclusives), and a centre of radiation with 31 spp. in the Andean region from Colombia, Peru, Ecuador (and Galapagos Is.), Bolivia, to NW Argentina and coastal N Chile; one sp. naturalised in Philippines. Temperate, continental temperate, mediterranean, subtropical and montane tropical forest, woodland, thorn-thicket, grassland, shrubland and desert, often in gullies and on rocky hillsides.

Used as soil stabilisers, green manure, stock forage, medicine, ornamentals, brooms and in making baskets; D. purpurea Vent. (purple prairie clover) roots are chewed for their pleasant taste and the dried leaves are a tea substitute

88. Errazurizia Phil. Xerophytic shrubs. 4 spp., two in SW U.S.A., one in NW Mexico and E. multifoliolata (Clos) I.M. Johnst. in coastal Chile; subtropical and mediterranean fog-deserts (Baja California, Sonoran and Atacama deserts), on rock or sand.

∎ SUBTRIBE DALBERGIINAE (45–46/1.280) ▸ outsiders unvailable.

89. Acosmium Schott. Trees and shrubs. 5 spp., Brazil (4, two endemics) to Paraguay and N Argentina, also in, in seasonally dry tropical woodland and wooded grassland, occasionally lowland forest and dunelands.

90. Adesmia DC. Annual and perennial herbs and shrubs, sometimes cushions. 204 spp., from Andes of Peru to Tierra del Fuego, S Brazil (19, 13 endemic) and Uruguay, centred in Andean Chile and Argentina (171), in subtropical to temperate semiarid and arid montane grassland and shrubland (chaparral); second largest genus endemic to South America after Dyckia (Bromeliaceae); the dead, spiny stems of some species are piled and stacked to make impenetrable fences; used as ground cover, erosion control, human food (e.g., roots of A. lotoides Hook.f.), ornamentals and medicine.

91. Aeschynomene L. Shrubs or herbs, in very few cases climbers, sometimes with xylopodium; seasonally dry tropical woodland, wooded grassland, bushland and grassland (sometimes montane), often in rocky or sandy areas; many species are hydrophytes occurring in marshes, at the edges of water holes, in swampy areas and flood plains. 100-120 spp., c. 34 occur in the Neotropics and subtropics, centred in Mexico to C America; in the Old World the distribution is principally African-Madagascan (c. 90–95), with one sp., A. aspera L., endemic to Asia and Australia; c. 3–4 spp. are widely introduced in the Palaeotropics from the New World; 29 spp. in South America, 23 in Brazil, 9 endemics. Used ecologically in management of inundated areas, as ornamentals, for fodder and green manure; A. aspera L. (sola, shola, ambatch wood, joint vetch) and A. elaphroxylon (Guill. & Perr.) Taub. are major sources of pith, a white spongy wood used for paper, fibre, helmets (solar topi), art work, handicrafts and artificial flowers (e.g., sola rosario flowers); the wood is also used for floats, rafts and canoes

92. Amicia Kunth. Perennial herbs or spindly to multiple-stemmed shrubs. 7 spp., one in Mexico and Central America, remaining in Andes S America from Ecuador, Peru, Bolivia, and Argentina, weedy along roads, and in seasonally dry tropical montane grassland to 3,000 m; occasionally used as cattle fodder in Argentina; A. zygomeris DC. is an attractive ornamental.

93. Arachis L. Annual, perennial or biennial, erect, decumbent or procumbent, sometimes rhizomatous or stoloniferous; tetrafoliolate leaves, sometimes trifoliolate; pauciflorous axillary spikes, arranged along the length of the branches or grouped around the collar of the plant; flowers sessile, hypanthium well developed; corolla orange or yellow; subterranean fruit (all species, the largest geocarpic genera off all flowering plants). 80 spp., C & S South America, centered in Brazil (63, 44 endemics) and N Argentina, Bolivia (second center of diversity), Uruguay and Paraguay; some spp. are narrow endemic to dry places in NE Brazil, in seasonally dry tropical to subtropical, dry and well-drained wooded grassland and grassland.

A. hypogaea L., the peanut or groundnut, is a major human food and source of vegetable oil (second only to soybean in importance among legumes), and animal fodder (mainly leaves); also used as a soil fertiliser and ground cover; it´s ancient natively endemic to Bolvia; other species are used similarly, but on a more local or regional basis (e.g., A. pintoi Krapov. & W.C.Greg., native from dry areas in E Brazil).

94. Cascaronia Griseb. Spindly shrubs or multiple-stemmed trees. Only one sp., C. astragalina Griseb., endemic to S Bolivia, NW Argentina and Paraguay, in seasonally dry subtropical forest, forest margins and along rivers; the wood is used for fuel and turnery

95. Centrolobium Mart. ex Benth. Trees up to 35 m tall. 7 spp., 3 in Ecuador, Colombia, Venezuela, Guyana and N Brazil, extending into Panamá; 4 in SE Brazil and E Bolivia, in tropical rain forest and seasonally dry forest at low elevations, dry seasonal scrubland of NE Brazil (caatinga), Atlantic sandy coastal shrublands (restingas). Various species (porcupine wood, canary wood, zebra wood, amarillo guayaquil, arariba) used for timber in house construction, flooring, ship components, decorative veneers, fine furniture and cabinet work; also planted as hedgerows, as ornamentals and shade trees in plantations; known as porcupine trees in some areas because of their large samaroid fruits with prickly seed chambers; 5 spp. in Brazil, 3 endemics.

96. Chapmannia Toor. & A. Gray. (inc. Pachecoa) Perennial herbs, small shrubs or trees. 7 spp., two in New World (C. prismatica (Sessé & Moc.) Thulin in Mexico, Guatemala, Colombia and Venezuela and one sp. in Florida), one sp. in Somalia and 4 spp. in Socotra; seasonally dry tropical to subtropical woodland, bushland, grassland and open scrubland, often in rocky or sandy areas and on roadsides. Used as an ornamental and for fodder in Somalia

97. Cranocarpus Bentham. Slender erect shrubs. 3 spp., endemic to E Brazil, in seasonally dry tropical humid forest (2); the other species in Atlantic sandy coastal shrublands (restingas).

98. Ctenodon Baill. Herbs, pyrophytic or fire-sensitive subshrubs, shrubs, treelets or a small tree reaching up to 8 m tall; erect or prostrate; the stems covered by glandular hairs or simple pubescence, or the plants sometimes glabrous; 66 spp., over tropical New World, 45 in South America, 36 in Brazil, 24 endemics, six of them, from Bahia and Goiás states, are rare plants in Brazil, by Plantas Raras do Brasil’s book.

99. Dalbergia L.f. Shrubs, trees up to 40 m tall, and climbing lianas, some species vary in habit from scandent shrubs in dry habitats to robust lianas in humid areas. Over 100 species distributed pantropically, but with centers of diversity in Amazonia and Indo-Asia. pantropical, centred in the Old World with 60–70 species in Africa (one sp., D. ecastaphyllum (L.) Taub., reaches India), 43 spp. in Madagascar, of which 42 are endemic; about 80 species in Asia with 33 species in India (19 endemic), 44 in Indo-China, and eight in New Guinea. Two of the Asian species (D. candenatensis (Dennst.) Prain and D. densa Benth.) reach Australia; 61 spp. occur in the neotropics with 44 spp. in S America (centred in Amazonia, 39 in Brazil, 21 endemics) and c. 15–20 spp. in tropical Mexico to C America and the Caribbean. Tropical rain forest to seasonally dry tropical to subtropical humid and dry forest, woodland, bushland, thicket and wooded grassland.

Many Indian, African, and Brazilian species of Dalbergia, including D. nigra(Vell.) Allemão ex Benth., D. sissoo Roxb., D. latifolia Roxb. and D. melanoxylon Guill. & Perr. are known to produce high quality timber that is used for construction, fine furniture, cabinet work, marquetry and inlay, pianos and other musical instruments, tool and cutlery handles, turnery, carving and various specialty items. Common names include rosewood, blackwood, tulipwood, kingwood, ebonywood (not true ebony), cocobolo, nambar, palisandro and sisam. The term rosewood most probably refers to the pleasant smell of the wood. Since the wood burns well, it is also often the preferred wood for cooking and charcoal-making (e.g., D. sissoo in Asia). Several species are cultivated in the Old World tropics as multipurpose trees for timber, fibre, fodder, fuelwood and medicine, or as ornamentals; contact dermatitis is reported from the wood.

100. Diphysa Jacq. Shrubs and trees. 20 spp., Mexico and C America, with two sp. extending into S America (Venezuela and Colombia), and another into S U.S.A. (Arizona), in seasonal tropical dry to humid forest and thorn scrub forest, often along rivers. Used as living fences and occasionally as ornamentals in Mexico; the wood is strong and has been used as clubs in battle

101. Discolobium Benth. Spindly shrubs or perennial herbs. 7 spp., Bolivia, NE Argentina, Brazil (6, Mato Grosso to Bahia, 3 endemics) and Paraguay, in seasonally dry tropical to subtropical wooded grassland or grassland, often hydrophytes in permanently inundated and seasonally flooded areas, in swamps and along rivers. Used as forage and medicine

102. Etaballia Benth. Trees. Only one sp., E. dubia (Kunth) Rudd, neotropical, occurring in Guyana, Venezuela, and N Brazil, in tropical lowland rain forest, often along riverbanks.

103. Fissicalyx Benth. Trees. Only one sp., F. fendleri Benth., Venezuela, Guyana, Brazil (only in Roraima state) and Panamá, in lowland rain forest (on terra firme) to lower-level montane forest (evergreen to semi-deciduous), forest margins and wooded grassland.

104. Fiebrigiella Harms. Perennial herbs. Only one sp., F. gracilis Harms, Andean S America from S Ecuador, Peru and Bolivia, in tropical montane grassland, shrubland and thicket to 3,500 m, often in rocky areas.

105. Geoffroea Jacq. Trees or shrubs. 3 spp., two from N Chile, Argentina (one endemic), Bolivia, Paraguay and Uruguay, and G. spinosa Jacq. from Argentina and Paraguay to Bolivia, Peru, Ecuador, N Colombia, Venezuela, NE Brazil and the Antilles, characteristic component of dry open chaco vegetation in southern S America, and in disjunct areas of tropical seasonally dry forest in northern S America. Used for human food (the edible fruits are made into jam or used to flavour wine); the timber is used for carpentry and furniture making; the bark and leaves have medicinal properties

106. Grazielodendron H.C.Lima. Only one sp., G. riodocensis H.C.Lima, restricted of tabuleiros region in S Bahia and N Espírito Santo states in Atlantic Forest of SE Brazil.

107. Machaerium Pers. Shrubs or small trees with firm glabrous branches with pairs of sharp, recurved stipular spines; leaves imparipinnate, 5– 9-foliolate; leaflets opposite or subopposite; inflorescence axillary and terminal, sparsely branched panicles; pod strongly falcate to curved so that point touches base, indehiscent, flattened, coriaceous, 1-seeded; upper suture prominent, lower one much arched; seeds reniform, flattened. 130 spp., Neotropics, centred in Amazonian S America (116, 74 in Brazil, 36 endemics), with c. 15 spp. from Mexico (2 endemics) and C America (2); M. lunatum is also amphi-atlantic, extending to the west coast of Africa.

Tropical inundated and non-inundated rain forest, seasonally dry forest, dune forest, low woodland, thicket, thorn scrub, shrubland and remnant shade trees in cocoa plantations; Machaerium species are widely used in forest management as shade plants and for the recovery of degraded areas; the wood of many species (caviuna, pau ferro, santos rosewood, morado), is hard and used for fine furniture, cabinetry, decorative veneers, panelling, pianos, flutes, axe handles and fence posts (the wood is resistant to decay); the reddish sap of some species has been used by native tribes to treat snakebite and the leaves of other species are a cocaine substitute; contact dermatitis is reported from the wood.

108. Maraniona C.E.Hughes, G.P.Lewis, Daza & Reynel. Small trees. Only one sp., M. lavinii C.E. Hughes, G.P. Lewis, Daza & Reynel in northern Peru (Cajamarca, Amazonas), in tropical seasonally dry forest and thorn scrub on steep rocky slopes, in a narrow elevational zone between 1,400 and 1,600 m.

109. Nissolia Jacq. (inc. Chaetocalyx) Perennial twining herbs and shrubs. 32 spp., centered in Mexico, 19 in South America, 13 in Brazil, 7 endemics, in seasonally dry tropical to warm temperate forest, shrubland and grassland, often in mesic sites along forest margins, on stream banks, hillsides and in ravines. Used as a fish poison and as an antidote to snakebite.

110. Paramachaerium Ducke. Trees. 5 spp., 4 South American in Peru, N Brazil (3, none endemics) and Guianas, one endemic to Panamá, tropical periodically inundated or non-inundated rain forest and seasonally dry woodland, often along rivers.

111. Platymiscium Vogel. Tall trees. 20 spp. from Mexico and C America to tropical S America; 11 spp. in South America, 7 in Brazil, two endemics. Used as ornamentals; most species (variously called trebol, macawood, macacauba, granadillo, coyote, tarara colorado) are highly valued regionally for their hardwood timber, used in house construction and for fine furniture, cabinet making, decorative veneers, joinery and musical instruments; a half of this genus are myrmecophites.

112. Platypodium Vogel. Trees 20 - 30 m. Only one sp., P. elegans Vogel, Panamá, Colombia, Venezuela, Brazil, Bolivia and Paraguay, in seasonally dry tropical or humid gallery or riverine forest, thicket and woodland; Used for timber, in house construction, furniture, tools and for decoration.

113. Poiretia Vent. Twining, scandent or erect, perennial herbs or shrubs. 13 spp. confined to the Neotropics, one endemic to Venezuela, two in Brazil and neighborig countries, 9 endemics to Brazil (three of them, from Minas Gerais and Bahia states, are rare plant in Brazil, by Plantas Raras do Brasil’s book), and P. punctata Desv., extending into northern S America, C America, Caribbean and Mexico, in seasonally dry tropical to subtropical riverine forest, savannas of C Brazil (cerrado), grassland and shrubland. One species is said to be toxic to cattle and may be a source of rotenones.

114. Pterocarpus Jacq. Tall trees, some species are buttressed rain forest emergentes. 35-40 spp., distributed pantropically, with the greatest diversity in Africa; c. 20 spp. endemic to Africa, one sp. (P. santalinoides L’Her. ex DC.) also native to the Neotropics, and one sp. (P. indicus Willd.) native to Asia; 5 spp. restricted to S Asia, Indo-China and Madagascar; c. 13 spp. endemic to the Neotropics (10 in South America, 7 in Brazil, 3 endemics - one of them, from Bahia state, is a rare plant in Brazil, by Plantas Raras do Brasil’s book), in tropical lowland evergreen rain forest (less than 10) to seasonally dry forest, woodland, thicket and wooded grassland.

P. indicus (narra, Solomon’s padauk, Papua New Guinea rosewood, amboyna) may be the most economically important legume timber species. The wood of several species including P. angolensis DC. (African teak, bloodwood, kiaat, muninga) and P. soyauxii Taub. (padauk) is highly valued and used for fine furniture, cabinet making, panelling, joinery, musical instruments, implements and curios. The resin supplies dyes (e.g., P. tinctorius Welw.), and in the Amazon is known as ‘sangre de drago’ (= dragon ’ s blood); the resin is also a widely used folk medicine (the source of Kino gum); other uses are in revegetation and soil improvement, as shade trees, ornamentals, cosmetics and baskets (from the inner bark). P. amazonum (Benth.) Amshoff from northern South America is a myrmecophyte.

115. Ramorinoa Speg. Xerophytic shrub or tree, aphyllous thorn. Only one sp., R. girolae Speg., endemic to W Argentina (La Rioja, San Juan, and San Luis provinces), in subtropical lowland or lower montane dry forest or shrubland in rocky or sandy areas (stands of this spp. are called ‘Chicales’). The beautiful and durable wood (chica) is used for musical instruments and furniture; seeds are eaten and also ground up and used as a coffee substitute.

116. Riedeliella Harms. Lanky, scandent shrubs. 3 spp. endemic to SE Brazil except one up to Paraguay, in seasonally dry tropical forest, savannas of C Brazil (cerrado) and dry seasonal scrubland of NE Brazil (caatinga) - one of them, from Minas Gerais state, is a rare plant in Brazil, by Plantas Raras do Brasil’s book. Used locally as firewood and as ornamentals.

117. Soemmeringia Mart. Prostrate shrub or perennial herbs. Only one sp., S. semperflorens Mart., from Brazil, Bolivia, Colombia, Guianas and Venezuela, in seasonally dry tropical forest, savannas of C Brazil (cerrado), wooded grassland and scrub, often along rivers, in floodplains and disturbed areas.

118. Steinbachiella Harms. Only one sp., S. leptoclada Harms, endemic to E Bolivia, closed Brazilian border (Mato Grosso state).

119. Stylosanthes Sw. Subshrubs or perennial herbs, sometimes with xylopodium, seasonally dry tropical to warm temperate woodland, wooded grassland, thicket, shrubland and grassland, on sandy or rocky soils, along streams and sometimes weedy in old cultivated lands and on roadsides. 46 spp., native to New and Old World, mainly neotropical, centred in S America (41, 32 in Brazil, 15 endemics) with 4 spp. in N & C America and 6 spp. widely distributed in the Neotropics, and 2 spp. widely distributed in Africa, Madagascar, India and Sri Lanka (S. fruticosa (Retz.) Alston and S. erecta P.Beauv). Several New World spp. have been cultivated and are naturalised in Africa, Asia and Australia (e.g., S. guianensis (Aubl.) Sw., S. humilis Kunth and S. viscosa Sw.). Major livestock fodder plants in warm temperate and tropical areas of the world (e.g., S. guianensis or Brazilian lucerne); several species are planted as soil stabilisers and improvers, and as ground cover, in e.g., coffee plantations; also used for medicine.

120. Tipuana (Benth.) Benth. Erect or spreading trees. Only one sp., T. tipu (Benth.) Kuntze, of subtropical forests in Bolivia and NW Argentina and Brazil, characteristic component of seasonally dry subtropical (gallery to montane) forest and open shrubland. T. tipu (tipu, rosewood, racehorse tree, pride of Bolivia, palo mortero) is widely used as an ornamental and street tree in subtropical to warm temperate areas; also in reforestation, as windbreaks and for erosion control; for fodder, timber (furniture and cabinet making), firewood and charcoal

121. Weberbauerella Ulbr. Perennial herbs or shrubs. Three spp., two coast of Peru and one in N Chile, in seasonally dry tropical coastal forest, in vegetation on sands and sandy hills (lomas).

122. Zornia J.F.Gmel. Usually erect or prostrate herbs, sometimes shrubs, in spiciform inflorescences. 75 spp., pantropical; 55 spp. in New World, centred in S America with 47 spp. (37 in Brazil, 19 endemics - three of them, from Goiás, Minas Gerais and Bahia, are rare plants in Brazil, by Plantas Raras do Brasil’s book), but also c. 14 spp. in U.S.A., Mexico and C America; c. 30 spp. in the Old World (c. 13 in Africa, one sp. endemic in Madagascar, c. 8 spp. in Asia and c. 8 spp. endemic in Australia); dry tropical to warm temperate woodland, bushland, wooded grassland, grassland and shrubland, often in rocky areas and weedy in cultivated lands; Used as forage, a cover crop (green manure), for medicine and soap; some species may be toxic

4.10 FABOIDEAE ▸ TRIBE INDIGOLFEROIDEAE (6/ c.800) - outsiders Cyamopsis (4; drier regions in Africa southwards to South Africa and eastwards to India), Indigastrum (9; tropical and S Africa, one pantropical species), Phylloxylon (7; Madagascar), Microcharis (c 35; Africa southwards to South Africa, Madagascar, Arabian Peninsula), Rhynchotropis (2; S Central Africa).

123. Indigofera L. c 730 spp., Africa–Madagascar (c. 490); Asia to Pacific (c. 115); Australasia (c. 30–40); c. 13 spp. widely distributed in the Palaeotropics; c. 6 spp. pantropical; New World has 63 spp. U.S.A. to Argentina (c. 30 in N & C America, c. 20 spp. in S America, 12 in Brazil, three endemics); seasonally dry tropical to warm temperate forest, woodland, wooded grassland and grassland, sclerophyllous shrubland, forest margins and disturbed areas. c. 25 sections in Africa-Madagascar where Indigofera is most diverse, increasing to c. 30 sections worldwide; all species belong to one of four well-supported and biogeographically distinctive clades in the analyses of Schrire et al. (2003).

Used as dyes (important species are I. arrecta Hochst. ex A. Rich., I. articulata Gouan, I. suffruticosa Mill. and I. tinctoria L.), medicine, fodder, cover crops, green manure, human food, erosion control and ornamentals; some species are toxic to livestock, others have insecticidal qualities.

4.11 FABOIDEAE ▸ TRIBE PHASEOLOIDEAE (164/3,210–3,300) – nine lineages, Kennediinae (4/21-23, tropical Asia, New Guinea, Australia) and Abrinae (1/13–18; tropical and southern Africa, Madagascar, Arabian Peninsula, tropical Asia to China and tropical Australia) not occur in South America.

∎ SUBTRIBE CLITORIINAE ▸ outsiders Clitoriopsis (1; Congo, Sudan), Austrosteenisia (4; New Guinea, N Australia).

124. Centrosema (DC.) Benth. Climbing herbs or subshrubs, sometimes with xylopodium; leaves pinnately 3-foliolate; stipules small, striate; stipels present; inflorescences (1) few– many-flowered, falsely racemose; bracts paired, striate; bracteoles appressed to calyx, striate; corolla showy. 42 spp., S America (38, 32 in Brazil, 11 endemics), C America, Caribbean, Mexico and SE U.S.A.; 3 spp. widely introduced in the Old World; tropical and subtropical bushland formations derived from seasonal forest, woodland, wooded grassland and shrubland, also forest margins.

Important forage and pasture crops with a broad range of tolerance of extreme conditions; also used as cover crops and green manure; C. rotundifolium Mart. ex Benth., from Brazil and Bolivia, is a amphicarpic species, unique in this genus.

125. Clitoria L. Shrubs, lianas, herbs and trees, someties with woody rhzomes and xylopodium; seasonally dry to wet tropical lowland and montane forest, woodland or scrubland and wooded grassland. 58 spp., 44 in S America, C America, Caribbean and Mexico (27 in Brazil, 3 endemics), Africa (4), Madagascar (2), Indian subcontinent (1), Indo-China, China, Malesia (6), Australia (1). Used as ornamentals, forage, green manure, cover crops and medicine.

126. Periandra Mart. ex. Benth. Shrubs, subshrubs or twining herbs, sometimes with roots crown, seasonally dry and wet tropical riverine forest, woodland or wooded to shrubby grassland, often in rocky areas. 7 spp., one in Caribbean and six in Brazil, three of then up to Bolivia. The sweet-tasting root is used as a substitute for liquorice.

∎ SUBTRIBE MILLETTIEAE ▸ Dalbergiella (2; tropical Africa), Millettia (c 150; tropical and subtropical regions in the Old World, especially Africa and Madagascar), Leptoderris (c 40; tropical Africa), Philenoptera (13; tropical Africa, Madagascar), Ophrestia (15–20; Africa, SE Asia), Mundulea (13–14; S Africa, Madagascar), Ptycholobium (3; drier regions in NE and S Africa, Arabian Peninsula), Pongamiopsis (3; Madagascar), Fordia (18; SE Asia, West Malesia), Burkilliodendron (1; the Malay Peninsula), Solori (9; SE Asia), Derris (c 65; tropical regions of the Old World to New Guinea); Kunstleria (11; Kerala in SW India, West and Central Malesia, tropical Australia); Apios (8–10; E Asia, S Canada, U.S.A.); Craibia (2; tropical to S Africa), Ostryocarpus (2; tropical W and C Africa), Xeroderris (1; tropical to S Africa), Platysepalum (7–8; tropical Africa), Schefflerodendron (3–4; tropical Africa), Sylvichadsia (4; Madagascar), Hesperothamnus (5; Mexico), Aganope (3–6; tropical Africa, SE Asia), Pyranthus (6; Madagascar), Chadsia (c 17; Madagascar), Requienia (3; W to NE Africa, S Africa), Dewevrea (1; tropical W Africa).

127. Apurimacia Harms. Shrubs. Three spp., Peru and Bolivia, and Cordoba Hills, Argentina, seasonally dry tropical montane shrubland on rocky slopes; used as fish poisons and insecticides.

128. Dahlstedtia Malme. Small trees or (sometimes scandent) shrubs. 13 spp., Bolivia, S Brazil (12, 9 endemic - 4 of them, from Bahia, Minas Gerais and Rio de Janeiro states, are rare plants in Brazil, by Plantas Raras do Brasil’s book), N Argentina, in tropical (mostly Atlantic) rain forest, mainly riverine, to seasonally dry forest on rocky slopes. Used as a fish poison (roots) and for insecticides.

129. Deguelia Aubl. (inc. Milletia) Lianas. 22 spp., Panamá to Amazonia Amazonian S America (21), mostly Brazil (17, 9 endemics, one of them, from Amazonas state, is a rare plant in Brazil, by Plantas Raras do Brasil’s book), in tropical Amazonian and Atlantic rain forest, often seasonally flooded or riverine; used as insecticides (rotenones).

130. Lonchocarpus Kunth. Mostly trees, less often shrubs. c 164 spp., S America (40, c. 22 Amazonian and c. 20 non-Amazonian, 11 in Brazil, two endemics - one of them, from Minas Gerais state, is a rare plant in Brazil, by Plantas Raras do Brasil’s book); C America, Caribbean and Mexico (c. 120) with one amphi-Atlantic sp. (L. sericeus) widely distributed also in Africa, in tropical rain forest and seasonally dry forest, woodland or rocky shrubland; various species used as timber (e.g., black cabbage bark, machiche, sindjaple), for furniture, construction and flooring; also used as insecticides (rotenones), fish poisons, dyes, ornamentals, fodder, fibre and medicine.

131. Muellera L. f. (inc. Margaritolobium, Coronila) Trees or (sometimes) shrubs. 27 spp. in tropical South America (26) to Mexico (one endemic); northern species in tropical swamp or riverine forest and mangrove; southern species in seasonally dry tropical shrubland along rivers and streams. Used as fish poisons; 19 spp. in Brazil, 11 endemics - one of them, from Mato Grosso do Sul state, is a rare plant in Brazil, by Plantas Raras do Brasil’s book.

132. Tephrosia Pers. Clump-forming perennials with tough roots; stems densely short-pilose, sometimes with xylopodium; leaves once odd-pinnately compound. 400 spp. mainly in seasonal areas of the tropics, especially Africa, concentrated in C and tropical America (74), Africa-Madagascar (c. 170), Asia (c. 40) and Australia (c. 90); seasonally dry tropical woodland, bushland, thicket and grassland, often in open and disturbed sandy or rocky areas. 18 spp. in South America, 10 in Brazil, 3 endemics; used as fish poisons, cover crops, livestock fodder, insecticides, ornamentals and for medicine, e.g. T. virginiana (L.) Pers. (goats rue, devil’s shoestring, hoary pea), T. vogelii Hook.f. (fish-poison pea), T. purpurea (L.) Pers. and T. candida (Roxb.) DC. (white tephrosia).

133. Piscidia L. Trees, rarely shrubs. 7 spp., SE U.S.A. (Florida), Mexico, C America, Caribbean, with P. carthagenensis Jacq. to Ecuador, Peru, Colombia, Venezuela, seasonally dry tropical forest, woodland, or bushland, often on rocky hills (some restricted to limestone). Used as fish poisons, fodder, medicine (the bark and essential oil of P. erythrina L., the Jamaica dogwood, are used in commercial herbal remedies) and timber (construction)

134. Platycyamus Benth. Trees. Two spp., P. regnellii Benth. endemic to Brazil, and P. ulei Harms from small area in S Acre state in Brazil, Peru (near easternmost point) and Bolivia (northermost point), in tropical rain forest (Amazonian and E coastal Brazil). Used as ornamentals.

∎ SUBTRIBE DESMODIINAE ▸ outsiders Pseudarthria (6; S Africa, Madagascar, Mauritius, Réunion, tropical Asia), Lespedeza (c 40; tropical and E Asia, Australia, temperate North), Grona (41; tropical and subtropical regions), Alysicarpus (30–35; tropical and subtropical Africa, S Asia), Melliniella (1; Africa), Leptodesmia (3; Madagascar, India), Codariocalyx (2; India, Sri Lanka, China (inc. Taiwan), Indochina, Malesia to tropical Australia), Uraria (c 20; tropical and subtropical regions in the Old World), Mecopus (1; India, Hainan, Indochina, Java), Christia (2; India to China and Indochina, Malesia to N Australia), Eleiotis (2; India and Sri Lanka to Burma), Ototropis (13; tropical Asia, S China inc. Taiwan), Pycnospora (1; tropical E Africa to Somalia, India, E and SE Asia to N Australia), Trifidacanthus (1; Hainan, S Vietnam, Philippines, Lombok, Flores); Ougeinia (1; India, Nepal), Aphyllodium (4; India, Sri Lanka, Hainan, Indochina, Malesia to New Guinea, N Australia), Tadehagi (4; tropical Asia), Akschindlium (1; SE Asia), Droogmansia (6; tropical Africa), Dendrolobium (c 20; tropical Asia, Indian Ocean islands, Australia), Phyllodium (8; tropical and subtropical Asia to N Australia), Ohwia (1; India, China (inc. Taiwan), Japan, Indochina, Malesia), Hanslia (2; Malesia to New Guinea, Queensland, Vanuatu), Nephrodesmus (5–7; New Caledonia), Arthroclianthus (c 20; New Caledonia, one species also in Vanuatu), Verdesmum (1; Yunnan).

135. Desmodium Desv. (exc. Grona) Perennial herbs or shrubs, rarely small trees; leaves pinnately compound; leaflets mainly 3; stipules broad or narrow, persistent or falling early; flowers small, usually many, in erect, narrow racemes or panicles; petals pink, lavender, red-purple, purple, or whitish, often drying bluish or a striking blue-green. 231 spp. of warm areas of the world (163 in New World), especially E Asia, Brazil, and Mexico, most diverse in SE Asia (at infrageneric level) and Mexico to S America (at specific level). In New World occur in warm temperate N America (c. 33), Mexico (c. 80, c. 50 endemic), C America, Caribbean and tropical to subtropical S America (60 in continent, 26 in Brazil, 3 endemics, one of them, from Mato Grosso state, are rare plants in Brazil, by Plantas Raras do Brasil’s book) and Australia (c. 14). A number of species are widely distributed in the Old and New Worlds and are often cultivated. Seasonally dry to wet tropical, warm temperate and temperate forest, woodland, thicket, wooded grassland, bushland and grassland, usually common in open or seasonally wet and riverine areas, also in disturbed and ruderal vegetation; important livestock forage and cover crops (e.g., the tick clovers D. intortum (Mill.) Urb. and D. uncinatum (Jacq.) DC.), also grown as intercropping insect repellents; widely used for medicine; cultivated as ornamentals (e.g., D. elegans DC.), and used for fibre

136. Grona Lour. (off Desmodium) Desmodium-like. 44 spp., mainly from tropical Africa to N Australia and China; 5 spp. in New World, G. barbata (L.) H.Ohashi & K.Ohashi, G. triflora (L.) H.Ohashi & K.Ohashi and G. adscendens (Sw.) H.Ohashi & K.Ohashi pantropical, G. orinocensis (DC.) H.Ohashi & K.Ohashi from Colombia to Venezuela, and G. juruenensis (Hoehne) H.Ohashi & K.Ohashi, a rare plants in Brazil, by Plantas Raras do Brasil’s book, endemic to Mato Grosso state in C Brazil.

∎ SUBTRIBE CAJANINAE ▸ outsiders Butea (2; tropical Asia), Cajanus (37; tropical regions in the Old World to Australia), Adenodolichos (22; tropical Africa), Paracalyx (6; Ethiopia, Somalia, Socotra, India, Indochina), Bolusafra (1; W Cape), Carrissoa (1; Angola), Dunbaria (20–25; India, SE Asia to China, Malesia to New Guinea, N Australia), Flemingia (c 30; tropical regions in the Old World).

137. Eriosema (DC.) Rchb. Herbs or subshrubs, with xylopodial or taproot tubers. 151 spp., Africa and Madagascar (c. 100–110), N and S America (47, Mexico to N Argentina, 40 in South America, 36 in Brazil (highly centered in central savannas), 23 endemics) and SE Asia to Australia (2); seasonally dry tropical to subtropical forest margins, woodland, thicket and wooded grassland, or open grassland in rocky or swampy areas, or in old cultivations and waste ground. Used as human famine food, medicine and fish poison.

138. Rhynchosia Lour. Herbs, vines or subshrubs, 230 spp., pantropical, c. 140 spp. in Africa-Madagascar; 63 spp. in tropical and subtropical America (c. 28 in N and C America, 32 in S America, 22 in Brazil, 4 endemics) and c. 30–35 spp. in warm temperate to tropical Asia to N Australia (2 endemic); seasonally dry forest, forest margins, woodland, thicket, wooded grassland, shrubland and grassland, often in open rocky areas or along streams and in disturbed areas; many species are pyrophytes. Used as pasture plants and several species, commonly called rosary bean, have attractive red, blue, black, mottled or bicoloured seeds used for necklaces etc.; seeds also used as weights or as narcotics; plants also famine foods.

∎ SUBTRIBE ERYTHRININAE ▸ outsiders Otoptera (2; Africa, Madagascar), Psophocarpus (7; tropical and subtropical regions in the Old World), Spatholobus (c 30; SE Asia to Central Malesia), Meizotropis (2; India, the Himalayas, W Indochina), Cochlianthus (2; Nepal, W China).

139. Erythrina L. Trees or shrubs up to 40 m tall, with red to orange flowers and are apparently all bird-pollinated. 112 spp. of warm areas of the world, 64 in the Neotropics (c. 50 spp. in Mexico, C America and Caribbean; c. 24 in S America (11 in Brazil, one endemic), a number of which may comprise the basally branching elements of the genus); 38 in Africa and Madagascar; c. 12 in Asia to Australia), variously used as shade trees, ornamentals with showy flowers, for their colorful seeds used as beads, or to shade coffee on plantations. Seasonally dry tropical and subtropical lowland to upland forest (sometimes coastal, in inundated areas or riverine), woodland, wooded grassland, bushland, thicket and grassland.

Used as ornamentals (coral or lucky bean trees), shade trees, timber (construction, implements), living fences and enclosures, green manure, livestock fodder, medicine and seeds are used for necklaces. Trunks, stems, and even leaves are characteristically armed with spines. Some species are also soft wooded or have tuberous roots and can be grown as container plants or trained as bonsai. Two species, E. mulungu Mart. ex Benth. and E. velutina Willd., are common in dry seasonal scrubland of NE Brazil (caatinga), and while erythrinas were encountered at Porto Novo, Itinga, and Milagres, the plants were invariably leafless, flowerless, and in some cases without seeds, so species identification was difficult.

∎ SUBTRIBE PSORALEEAE ▸ outsiders Pseudovigna (1; tropical Africa), Neorautanenia (3; S tropical Africa), Neonotonia (2; tropical and S Africa, tropical Asia from S Arabian Peninsula to Malesia); Pueraria (17–20; S and E Asia); Amphicarpaea (5; Africa, E Asia, North America), Glycine (c 25; Africa, S Asia to Australia), Sinodolichos (2; Burma, S China), Bituminaria (4; Mediterranean to the Caucasus and the Middle East), Orbexilum (11; the U.S.A., Mexico), Psoralea (30–35; S Africa, with their largest diversity in Western Cape), Cullen (c 35; Mediterranean, Africa, S Asia to New Guinea and Australia), Pediomelum (20–25; North America), Ladeania (2; W U.S.A.), Rupertia (3; SW Canada, W U.S.A.), Herpyza (1; W Cuba), Teyleria (3; SE Asia, West Malesia), Dumasia (c 10; Africa to S Asia), Nogra (4; India to S China and Thailand), Eminia (4; Zambesian region in Africa), Pseudeminia (4; tropical Africa), Phylacium (2; SE Asia to Queensland), Neocollettia (1; Burma, Java), Mastersia (1; Assam, Central Malesia), Diphyllarium (1; Indochina).

140. Calopogonium Desv. Herbs or scandent shrubs. 9 spp., Mexico, C America and Caribbean to Paraguay and Argentina; 6 spp. in South America, 4 in Brazil, none endemics; seasonally dry tropical and subtropical forest, woodland or thicket, often on margins, usually near rivers or swamps, or in bushland, shrubland and disturbed grassy areas. Used as fodder and cover crops; C. mucunoides Desv. is widely used as a green manure; also a weedy escape in the Palaeotropics.

141. Otholobium C. H. Stirt. Shrubs, trees or herbs. 61 spp., 53 in S and E of southern Africa (Cape and Afromontane regions, especially common in mediterranean S and SW Cape, with outliers to Angola and E mountain areas of Kenya); possibly including 8 spp. from S America, in the middle to high altitude Andes, from Colombia and Venezuela southwards to Chile and Argentina; seasonally dry tropical and mediterranean lowland to montane shrubland, forest margins, grassland and seepage areas (Andean species in montane forest, woodland, shrubland and xeric grassland); used as medicine and for flavouring food by smoking

142. Pachyrhizus DC. Herbs or shrubs. 6 spp., Mexico, C America, Colombia, Venezuela, Ecuador, Peru, Bolivia, inc. 3 species widely cultivated in tropical and subtropical America, one also extensively in the Palaeotropics, seasonally dry tropical forest and thicket, often on margins, and in scrub vegetation and open grassy areas. Used for human food, e.g., P. erosus (L.) Urb. (yam bean, jicama), pods also edible; other parts used for fibre and medicine.

143. Teramnus P. Browne. Climbing herbs or shrubs. 9 spp., principally Old World tropics (Africa, SE Asia (Indian subcontinent, S China, Indo-China, to Malesia)); one sp. neotropical and 2 pantropical, the three in South America, all widely distributed, two of then in Brazil; mainly seasonally dry tropical bushland and thicket, grassland, wooded grassland and forest clearings, often in open and rocky dry areas. Used for ground cover, has potential as pasture (forage) plants

∎ SUBTRIBE PHASEOLINAE ▸ outsiders Wajira (5; Africa, SW Arabian Peninsula, India, Sri Lanka); Sphenostylis (5; Africa, India), Decorsea (6; tropical to S Africa, Madagascar), Macrotyloma (c 25; tropical and subtropical Africa and Asia), Alistilus (3; southern tropical Africa, Madagascar), Dolichos (c 60; tropical and subtropical Africa to South Africa to India and E Asia), Nesphostylis (4; tropical Africa, tropical Asia); Dipogon (1; W and E Cape), Lablab (1; tropical and subtropical Africa), Spathionema (1; tropical Africa), Vatovaea (1; tropical E Africa to Oman), Physostigma (2–4; tropical Africa), Ramirezella (7; Mexico, Central America), Strophostyles (3; S Canada, U.S.A.), Pseudoeriosema (5; tropical Africa), Austrodolichos (1; N Australia), Dysolobium (6; E India, SW China, Indochina, Malesia), Strongylodon (14; Madagascar, tropical Asia to Polynesia).

144. Ancistrotropis A. Delgado. Small to medium size flowers (less than 2 cm) with a pleated calyx tube, a standard petal forming a hood, wing petals much longer than keel, and a hooked keel beak similar to that of Sigmoidotropis but with the distalmost portion splayed open. 7 spp., from South America, one up to North America; 6 in Brazil, 4 endemics.

145. Barbieria DC. Erect or scandent shrubs. Only one sp., B. pinnata (Pers.) Baill., S Mexico, C America, Caribbean and W South America to E Brazil, in tropical riverine forest, thicket and forest margins or in areas of secondary growth and open areas. Used as ornamentals and ground cover crops.

146. Cochliasanthus Benth. (off Vigna) Climbing herbs; flowers are characteristically large (e.g., the total calyx length measures ca. one cm on average) and very showy in that the standard and wing petals are twisted (spiraled) in a right-handed fashion, and the keel is distally twisted with up to fi ve loose right-handed coils. Only one sp., C. caracalla (L.) Trew., secondary and primary wet forests, mostly without a dry season, from southern Mexico to N Argentina and Uruguay.

147. Cologania Kunth. Perennial climbing herbs. 12 spp., tropical America, centred in Mexico (11, 6 endemics) to SW U.S.A., C America, South America (3) in Colombia, Venezuela, Ecuador, Peru, Bolivia and Argentina, largely confined to montane habitats, in tropical to warm temperate montane forest, woodland and thicket to wooded grassland and open bushland on mountain slopes.

148. Condylostylis Piper. (inc. Vigna p.p.) 4 spp., scattered, only one the widely distributed in Brazil, secondary and primary wet to semideciduous forests from southern Mexico south to Argentina and Uruguay (0 – 1,500 m altitude range); absent from the Caribbean.

149. Delgadoa F.S. Santos, C. Snak & L.P. Queiroz. Woody lianes, flowers white. Only one sp., D. bambuicola F.S. Santos, C. Snak & L.P. Queiroz, only two localities in SE Bahia state, Brazil.

150. Dolichopsis Hassl. Herbs. Two spp. known from the Chaco vegetation of Paraguay and adjacente Argentina and Brazil (only the latter, ca. 300 m altitude range, non endemic).

151. Helicotropis. A. Delgado. Perennial twining vines, from a thick taproot up to 30 cm deep; stems hollow, striate, with adventitious roots at nodes; flowers violet-purple, sometimes lavender-blue; calyx campanulate, upper lip broad, emarginated. 3 spp. from Mexico to Bolivia and S Brazil, two in South America, both in Brazil, none endemics.

152. Leptospron (Benth) A. Delgado. Two spp., one endemic to Mexico and L. adenanthum (G. Mey.) A. Delgado distributed throughout Mexico, Central America, the Caribbean, and in South America south to Argentina and Uruguay (0-2,000 m altitude range), also naturalized in limited areas of the paleotropics including Africa and Asia.

153. Macroptilium (Benth.) Urb. Herbs, prostrate or twining, sometimes with xylopodium; leaves pinnately 3-foliolate; inflorescences axillary, falsely racemose, long-pedunculate; flowers usually fairly small, white or red to blackish purple. 20 spp., N to S America (17, 11 in Brazil, one endemic) from S U.S.A. to Argentina, mainly concentrated in tropical S America, growing seasonally dry tropical and subtropical disturbed forest, woodland or thicket, scrubland and grassland, often weedy; major pasture and fodder legumes, e.g., M. atropurpureum (DC.) Urb. (siratro) and M. lathyroides (L.) Urb. (phasy bean); two spp. widely naturalised in E and S tropical Africa.

154. Mucuna Adans. Mostly lianas, sometimes erect shrubs, flowers arranged in pseudopanicles, pseudoracemes, or umbel-like pseudoracemes, and the inflorescence peduncle ranges from 2 cm to more than 2 m long; 114 spp., pantropical, c. 77 spp. in Asia and China, 1–2 spp. in Australia; c. 19 spp. in Africa, Madagascar and Mascarenes; 26 spp. in the Neotropics, centred in C America & the Caribbean (20 in South America, 8 spp. in Brazil, two endemics); tropical wet lowland to montane, often coastal rain forest, to seasonally dry forest, woodland and thicket; M. pruriens (L.) DC. (velvet, Bengal or Mauritius bean), is widely grown for forage and as a cover crop and green manure; the hairs on many species (often known as buffalo bean) are an intense irritant; some species are ornamentals.

155. Mysanthus G.P.Lewis & A.Delgado-Salinas. Herbaceous twining climber with terete pubescent stems, often climbing to 3 m; leaves pinnately trifoliolate; inflorescence an axillary, erect, nodose pseudoraceme up to 90 cm long; corolla glabrous, all petals fading to a yellowish colour; standard greenish cream near base, inner face of blade very pale pink or lilac, outer face dark pink or greyish purple, sometimes speckled or streaked dark purple. Only one sp., M. uleanus A.Delgado & G.P.Lewis, disjunct distribution: dry seasonal scrubland of NE Brazil (caatinga) at Bahia state (M. uleanus var. uleanus) and forests of E São Paulo state (M. uleanus var. dolicopsoides); possibly one sp. from savannas of Bolivia (cerrado Chiquitano).

156. Oryxis A. Delgado & G. P. Lewis. Robust twiner or semi-erect subshrub to c. 3 m; stems terete, becoming woody, up to 5 mm in diameter, minutely white or fulvous strigillose or tomentose, later glabrescent; leaves pinnately trifoliolate; inflorescence an erect, axillary, nodose pseudoraceme up to 45 cm long; corolla glabrous; standard outer face creamish with wine-red slightly thickened or inrolled margin, inner face lilac with a central yellow blotch bordered wine-red and with lilac coloured insect guides. Only one sp., O. monticola (Mart. ex Benth.) A.Delgado & G.P.Lewis, of the rocky grasslands (campos rupestres) and savannas of C Brazil (cerrado) in east central Brazil (Minas Gerais).

157. Oxyrhynchus Brandegee. Climbing herbs to shrubs. 4 spp., N and C America (3 from S U.S.A., Mexico, C America, Caribbean to NW Colombia) and SE Asia (one sp. from Papuasia and E Malesia), seasonally dry to wet tropical and subtropical, coastal and montane forest. Used for human food and forage.

158. Phaseolus L. Climbing herbs. 75 spp., concentrated in Mexico with a secondary diversification in the central Andes, most commonly inhabits upper elevation pine or oak forests but with some notable exceptions, such as P. viridis Piper from tropical wet forests and P. microcarpus Mart. from seasonally dry tropical forests; c. 3 spp. in S America endemic to the N and C Andes and Galapagos. Used for human food as major pulse and vegetable crops; also used for fodder, green manure and as ornamentals; two clades:

§ clade A ‣ 33 spp., 5 unplaced and remaining in three lineages: Pauciflorus (10, undisturbed pine-oak forests of Mexico, and barely enters Guatemala and SW U.S.A.), Pedicellatus (9, mainly in central and N Mexico and adjacent Texas, and S New Mexico and Arizona), and Tuerckheimii (9, throughout Mexico and Central America except Belize) groups, and the four weakly resolved species, and not occur in South America.

§ clade B ‣ comprises the Filiformis (3, Baja California to Coahuila, and adjacent S California), Vulgaris (9, throughout Mexico, Central America, and Andean South America, including four of the five cultivated species), Lunatus (7, Mexico, Bermudas, Peru and Galapagos one endemic each, P. augusti Harms from Ecuador to Argentina, and the cultivated P. lunatus L.), Leptostachyus (3, Mexico, Guatemala, El Salvador, Honduras, Nicaragua, and NW Costa Rica), and Polystachios (17, U.S.A. to E Texas, and throughout Mexico south to Oaxaca) groups.

Despites the five cultivated species (P. acutifolius A.Gray (tepary bean, U.S.A. and Mexico), P. coccineus L. (scarlet runner bean), P. lunatus L. (lima bean), P. vulgaris L. (common or kidney bean) and P. dumosus Macfad.), South America has only four wild species, one in Galapagos and three from Ecuador to NW Argentina.

159. Sigmoidotropis (Piper) A. Delgado. (inc. Vigna p.p., Macroptilium p.p.) 7 spp., S. speciosa (Kunth) A.Delgado widely in tropical America, two endemics to Peru, and remaining in Mexico, Central America and Caribbean, inhabiting secondary and primary forests, with or without a dry season, as well as coastal thickets and riparian forests throughout much of the neotropics (0 – 2250 m altitude range).

160. Vigna Savi. Herbs; Seasonally dry tropical woodland, wooded grassland and grassland, often in well drained sites with low fertility. 105 spp., palaeotropics and subtropics, c. 80–85 spp., mostly in Africa (c. 55–60 spp.; 4 spp. endemic to Madagascar) and SE Asia (c. 21), 11 in New World, inc. three pantropical (V. luteola (Jacq.) Benth., V. umbellata (Thunb.) Ohwi & H.Ohashi and V. vexillata (L.) A.Rich., two of then in Brazil), 4 in over tropical America (V. juruana (Harms) Verdc., V. lasiocarpa (Mart. ex Benth.) Verdc., V. longifolia (Benth.) Verdc., V. trichocarpa (C.Wright) A.Delgado, all in Brazil and in subg. Lasiospron), V. halophila (Piper) Maréchal, Mascherpa & Stainier endemic to Brazil, V. lonchophylla Piper endemic to Mexico, V. myrtifolia Piper endemic to Bolivia, and V. truxillensis (Kunth) N.Zamora disjuntc in Mexico, Bolivia and Central America; five subgenera: Ceratotropis, Haydonia, Lasiospron (6), Plectrotropis, and Vigna.

Many species are major pulse, vegetable, fodder and green manure crops, e.g., V. angularis (Willd.) Ohwi & H.Ohashi (azuki or adzuki bean); V. mungo (L.) Hepper (urd bean, black gram); V. radiata (L.) R.Wilczek (mung bean, green gram); V. umbellata (Thunb.) Owhi & H.Ohashi (rice bean); V. aconitifolia (Jacq.) Maréchal (moth bean); V. unguiculata (L.) Walp. (cowpea, yard long bean) and V. subterranea (L.) Verdc. (bambara groundnut, bambara bean).

4.12 FABOIDEAE ▸ TRIBE DIOCLEAE (13/200) - three lineages, all in South America:

∎ CANAVALIA CLADE ▸ a single genus.

161. Canavalia DC. Lianas to slender vines, in tropical flooded or swamp forest, river banks and forest margins to seasonally dry coastal vegetation, thicket, open woodland, wooded grassland and rocky hillsides, often climbing over or trailing through other vegetation. 60 spp., 38 in Neotropics (29 in S America, 16 in Brazil, 7 endemics); c. 10 spp. in Pacific Is.; 8 spp. in Asia and one sp. in Madagascar; c. 5 spp. widely distributed in Palaeotropics or pantropical; Used for human food, cover crops and green manures, e.g., C. gladiata (Jacq.) DC. (sword bean) and C. ensiformis (L.) DC. (jack bean); also as ornamentals, medicine, human food (vegetable) and forage; common drift seeds; one sp., fro Bahia state, is a rare plant in Brazil, by Plantas Raras do Brasil’s book.

∎ GALACTIA CLADE ▸ outsiders Lackeya (2; SE U.S.A., Mexico), Rhodopis (4; Hispaniola, Cuba, Porto Rico, Jamaica).

162. Betencourtia A. St.-Hil. (inc. Galactia p.p., Collaea p.p.) Herbs or subshrubs, either prostate or twining vines, sometimes with xylopodium; inflorescence an axillary and umbelliform pseudoraceme, pedunculate. 8 spp., 7 in Brazil (two endemics, four only from Bolivia and Cono Sur, and one scattered from South America up to Venezuela) and one only in Cono Sur, mostly from tropical (savannas of C Brazil (cerrado) and rocky grasslands (campos rupestres)) and subtropical (Pampas) open fields, with one species (B. scarlatina) from montane and gallery forests.

163. Bionia Mart. ex Benth. (exc. Mantiqueira) Low shrubs with virgate, little ramified branches, with or without underground woody rhizomes, less frequently twinning vines with branches woody or slightly woody; inflorescence a nodose pseudoraceme; flowers 2.5-4.0cm long, showy; petals dark red to orange-red, glabrous. 4 spp. widely distributed in mountains of E Brazil (Ceará, Pernambuco, Bahia, Minas Gerais and Espírito Santo) to C Brazilian Shield (states of Goiás, Tocantins, Maranhão and Distrito Federal); 4 undescribed species.

164. Caetangil L.P. Queiroz. (off Camptosema) Prostrate or twinning subshrubs; inflorescence an axillary elongate pseudoraceme, with a long and slender peduncle, and remote 2–3-flowered nodes, flowers shortly pedicellate and provided with a pair of bracteoles just below the calyx. 4 spp., two accepted and two undescribed, from seasonally dry vegetation in Chaco (Bolivia, Paraguay WC Brazil), dry seasonal scrubland of NE Brazil (caatinga), and open fields in S Guyana and N Brazil.

165. Camptosema Hook. & Arn. (exc. Caetangil) Lianas, vines or shrubs, tropical seasonally dry to wet forest or upland woodland. Only one sp., C. rubicundum Hook. & Arn., in Uruguay, N Argentina and the Brazilian state of Rio Grande do Sul, in thickets and gallery forests along the streams.

166. Cerradicola L.P. Queiroz. (off Galactia) Mostly perennial erect subshrubs, sometimes prostate but not twinning, frequently with hard underground rhizomes or xylopodia; inflorescence an axillary elongate pseudoraceme, flowers shortly pedicellate and provided with a pair of bracteoles just below the calyx. 17 spp., all in central Brazilian Shield (except one from Bolivia and NW Argentina), three extending to the E Andean slopes of E Bolivia and three up to NW Argentina. Some species also occur in similar environments in the rocky grasslands (campos rupestres) of the Espinhaço Range (in the Brazilian states of Bahia and Minas Gerais).

167. Collaea DC. Subshrubs, sometimes with xylopodium. 8 spp., S America from Brazil (6, 4 endemics, northward to the Brazilian states of Minas Gerais and Espírito Santo), Bolivia, Paraguay to N and C Argentina, also in Peru, in seasonally dry tropical and subtropical lowland to montane shrubland and grassland; potential as ornamentals.

168. Cratylia Mart. ex. Benth. (inc. Camptosema p.p.) Lianas or erect to scandent shrubs. 7 spp., all in Brazil (3 endemics), southward from the Amazon basin, to Paraguay, E Bolivia and Peru, in seasonally dry forests and woodlands, rain forests and savanna vegetation.

169. Galactia P. Browne. (exc. Cerradicola, Betencourtia p.p., Nanogalactia) Prostrate perennials herbs or shrubs; with trailing or twining stems, pubescent to nearly glabrous; leaves pinnately compound. 84 spp., distributed in North America (31), the Caribbean (45), Central America (9), South America (16-19, 5 in Brazil, fewer than 3 endemics (possibly only one)), Australia (3), Asia (2), and Africa (1).

170. Mantiqueira L.P. Queiroz (off Bionia). Vines, flowers with a tubular calyx, red petals and the standard petal spreading and longitudinally folded, delicate inflorescence with linear 2(–3)-flowered nodes. Only one sp., M. bella (Mart. ex Benth.) L.P. Queiroz, restricted to the Mantiqueira mountain range of E Brazil.

171. Nanogalactia L.P.Queiroz. (off Galactia) Erect or twinning herb, usually under 40 cm tall, the stems usually under one mm diam., often with xylopodium; inflorescence mostly an axillary, few-flowered axillary fascicle, sometimes a slender pseudoraceme with 1–2 nodes, each node vestigial, 1–3-flowered, flowers pedicellate and provided with a pair of bracteoles just below the calyx. Three spp., one in Mexico, another in North America, and N. pretiosa (Burkart) L.P. Queiroz in S Brazil and Cono Sur.

∎ DIOCLEA GROUP ▸ all genera in South America.

172. Dioclea Kunth. Climbing shrubs; leaves pinnately 3-foliolate; leaflets ovate-elliptic, rounded at base; inflorescences stiff, woody; corolla: vexillum orbicular, reflexed, auriculate at base; wings obovate or oblong, free; carina incurved, subrostrate; pod linear-oblong to ovate-oblong; 13 spp., all in South America up to N Argentina and Paraguay, slightly centered from Venezuela to Ecuador (5), three up to Central America or Mexico, in tropical lowland rain forest, riverine or swamp forest and thicket; seasonally dry tropical to subtropical woodland, wooded grassland and scrub.; 7 spp. in Brazil, two endemics.

173. Cymbosema Benth. Vines or lianas. Only one sp., C. roseum Benth., Amazonian Brazil, Colombia, Peru, Venezuela; C America; Mexico, in tropical lowland rain forest and riverine forest.

174. Cleobulia Mart. ex. Benth. Vines or lianas, or shrubs. 4 spp., one endemic to Mexico and three endemics to Brazil, in tropical Amazonian rain forest, seasonally dry forest, oak and pine forest.

175. Macropsychanthus Harms. Stout, high-climbing lianas with twining stems, less frequently shrubs or woody vines in open habitats.; leaves pinnately trifoliolate, stipellate or estipellate; inflorescence a stout, woody, erect pseudoraceme, nodes multiflorous, woody, stalked and secundiflorous; bracteoles fleshy. 48 spp., mainly in the New World (38-42, 23-27 in Brazil, 6-10 endemics - three of them, from Pará and Rio de Janeiro states, are rare plants in Brazil, by Plantas Raras do Brasil’s book), 11 spp. from Philippines and Indonesia to New Guinea and two Pantropical sea-drifted species extending to continental Africa and Madagascar.

4.13 FABOIDEAE ▸ TRIBE ROBINIEAE (11/75–80) - outsiders Robinia (4–5; North America), Genistidium (1; Texas, Mexico), Peteria (4; SW North America), Olneya (1; SW North America), Sphinctospermum (1; SW North America), Lennea (5; Central America), Hebestigma (1; Cuba), Poitea (12; the Caribbean).

176. Coursetia DC. (inc. Cracca) Trees, shrubs and herbs. 37 spp., SW U.S.A., Mexico, C America, Caribbean and S America (16), in Guyana, Venezuela, adjacent Brazil and the Andean cordillera to N Argentina; also with extensions to Paraguay and EC Brazil (5, 3 endemics - one of them, from Bahia state, is a rare plant in Brazil, by Plantas Raras do Brasil’s book); seasonally dry tropical and subtropical forest and montane woodland.

177. Gliricidia Kunth. Trees and shrubs. 5 spp., Mexico and C America, with G. brenningii (Harms) Lavin in Ecuador to Peru; G. sepium (Jacq.) Steud. is widely introduced in tropical areas throughout the world; seasonally dry tropical, often lowland forest, thicket and thorn scrub, on hillsides and in exposed or disturbed areas; one sp. in montane woodland.

The economically important G. sepium (madre de cacao, madricacao) is widely used for living fence systems, shade trees for crops, weed and erosion control, livestock fodder, medicine, insecticides, fuelwood, poles, green manure, ornamentals and bee forage

178. Poissonia Baill. Trees, shrubs and herbs. 5 spp., three in Peru and two in Argentina and Bolivia, in tropical and subtropical seasonally dry forest and shrubland, along river and stream banks, also in open arid vegetation; used for edible roots and as a soil stabiliser

4.14 FABOIDEAE ▸ TRIBE SESBENIEAE (1/c 60) - a single genus.

179. Sesbania Scop. Annual or perennial herbs or subshrubs, unarmed; stems long, green, glabrous, unbranched below; leaves once even-pinnately compound, up to 30 cm long. 60 spp., Africa-Madagascar (c. 30); Asia to Pacific (c. 9); Australia (c. 7); 2 spp. widely distributed in Palaeotropics; 12 spp. in New World (8 in South America, 5 in Brazil, one endemic); c. 3 spp. pantropical; seasonally dry tropical, subtropical and warm temperate areas, in seasonally wet, flooded or swampy habitats; riverine forest, woodland, wooded grassland and grassland, on lake margins, river banks and in coastal areas.

Four sections are recognised and only in warm temperate to tropical N America are all four represented; used for forage, fibre (from bark), wood, pulp & paper, dye, gum, cover crops, green manure, human food (e.g., flowers of S. grandiflora (L.) Pers.), medicine, ornamentals and fish poisons (useful species include S. grandiflora (agati or corkwood tree); S. bispinosa (Jacq.) W.Wight (dhaincha) and S. exaltata (Raf.) Cory (Colorado River hemp)); some species are invasive weeds and are toxic to livestock (e.g., S. punicea Benth.).

4.15 FABOIDEAE ▸ TRIBE LOTEAE (19/c 295) - outsiders unvailable, mainly Africa to Pakistan and North America.

180. Acmispon Raf. Herbs. 29 spp., 28 in S Canada to Mexico and A. subpinnatus (Lag.) D.D.Sokoloff, endemic to in Chile; warm temperate and mediterranean desert, grassland, shrubland, woodland and ruderal habitats.

181. Ornithopus L. Herbs. 5 spp., mainly W Europe (north to S Scotland) and Mediterranean region (incl. N Africa) to Macaronesia, east to Caucasus and Iran, O. micranthus (Benth.) Arechav. from N Argentina, Uruguay and Rio Grande do Sul state in S Brazil; mediterranean to temperate or subtropical grassland and shrubland, often on sandy soils.

Loteae with lomentaceous fruits are absent from the New World, so Ornithopus is unusual in this respect; this is the only genus of Loteae that includes both Old and New World species; O. sativus Brot. (serradella; common birdsfoot) and other species are cultivated for forage, soil cover, green manure and for hay and silage

4.16 FABOIDEAE ▸ TRIBE GLYCYRRHIZEAE (1/18) - a single genus.

182. Glycyrrhiza L. Herbs. 20 spp., mainly Eurasia; Europe (and Mediterranean including N Africa) to W, C and E Asia, also N America (1), G. astragalina Gillies ex Hook. & Arn. in Argentina and Chile and Australia (1); mainly mediterranean, warm temperate and continental temperate grassland, shrubland, bushland and woodland, often in sandy, marshy and disturbed areas.

G. glabra L. (liquorice, licorice) roots and its extracts are used in medicine, flavouring of confectionery and tobacco, human food (roots) and drink (tea), soil binders and as foaming agents (in industrial products)

4.17 FABOIDEAE ▸ TRIBE HEDYSAREAE (25–27/c. 4,300) - outsiders mainly in Northern Hemisphre (Oxytropis also in North America), with radiations in s Africa (eg. Lessertia, 55 spp.) and Australasia (eg. Swainsona, 85 spp., Australia).

183. Astragalus L. Herbs, vary from short living annual herbs (ca. 80) to perennial rhizomatous or hemicryptophytic herbs (ca. 2,500) and to cushion forming spiny shrubs (ca. 300) in habit; leaves once odd-pinnately compound, stipulate, estipellate; flowers in peduncled, terminal or axillary heads or head-like or spike-like racemes; petals various shades of pink, lavender, or purple to white or yellowish herbs or shrubs. c. 3,063 spp. (second argest worldwide) c. 2,400 in S and C Europe (chiefly Mediterranean region, including N Africa, A. atropilosulus (Hochst.) Bunge subsp. abyssinicus (Hochst.) Gillett up to S Africa, Middle East, SW Asia and Sino-Himalayan region to W China (c. 1,200, most diverse in Türkiye, Iran and Afghanistan); also E Europe to C Asia, Mongolia, Siberia, NE China and Japan (c. 620, mostly in former USSR) and 526 spp. in New World, in W N America (349 in U.S.A., 94 in Mexico, two in Guatemala) and S America, with 117 spp., mainly in Argentina (70, from NW to Patagonian coast), Peru (25), Chile and Bolivia (23), A. distinens Mackloskie reaching to W Uruguay, and 4 up to Ecuador.

4.18 FABOIDEAE ▸ TRIBE FABEAE (10/c 870) ▸ three subtribes, Parochetinae (1/1, Asia to Africa) is absent in New World.

∎ SUBTRIBE TRIFOLIINAE – outsiders Ononis (c 30; Europe, Canary Islands, Mediterranean, Ethiopia, Iran), Medicago (85–90; Europe, Mediterranean, Ethiopia, S Africa, Asia), Trigonella (35–40; Macaronesia, Mediterranean, S Africa, Australia), Melilotus (19; Europe, Mediterranean, northern Africa, Ethiopia, temperate and subtropical Asia).

184. Trifolium L. Annual or perennial herbs; leaves alternate, palmately or pinnately compound, 4th largest diversity of variegated leaves worlwide. 291 spp. of bee-pollinated herbs, principally N temperate Eurasian (c. 150, with c. 130 in Mediterranean region including Türkiye) and New World (87, mostly temperate N America); also tropical and subtropical montane areas: c. 36 in Africa, 18 in S America, mainly Andes, 4 up to S Brazil, three restricted of Cono Sur and S Brazil and one widely distributed; mainly mediterranean, temperate and tropical montane grassland; introduced worldwide for fodder, soil and pasture improvement, as honey plants, hay and silage, and in horticulture.

T. repens L. (white clover) is probably the most widely grown species with the greatest impact on agriculture of any cultivated forage plant; T. pratense L. (red clover) is also used for medicine; some species are eaten as human food; trefoil dermatitis associated with photosensitisation is a problem caused by animal ingestion of some Trifolium species. T. polymorphum Poir and T. argentinense Speg. are amphicarpic species, unique among this genus in New World.

∎ SUBTRIBE VICIINAE ▸ outsider Cicer (c 45; Greece, Canary Islands, Morocco, Ethiopia, W and C Asia).

185. Lathyrus L. Annual or perennial, trailing, sprawling, or ascending to climbing herbs with winged stems (except in L. venosus); leaves pinnately compound, the rachis terminating in a tendril that is often branched. 160 spp., mostly N temperate regions: Europe and Asia (c. 100 spp., principally Mediterranean and Irano-Turanian, some spp. to China, Korea and Japan) and N to E Africa (c. 5), with additional centres in N America (c. 30) and temperate S America (31, 14 in Brazil, two endemics); temperate, mediterranean and tropical montane grassland, shrubland and woodland.

Many species are widely introduced and naturalised: used extensively as cover crops, for fodder; as ornamentals (e.g., L. odoratus L. (sweet pea), L. latifolius L. (everlasting pea) and L. sylvestrisL.) and as human food (e.g., L. sativus (grass pea, Indian pea, chickling vetch), L. ochrus (L.) DC. and L. montanus Bernh. (with edible root tubers) and also for erosion control, as green manure and for medicine; toxins are present in some species, causing lathyrism

186. Vicia L. Annual, decumbent, trailing to climbing herbs; leaves once pinnately compound, the rachis tip usually terminating in a simple or branched tendril, stipulate. 140 spp., Europe and Asia (c. 110, principally from Mediterranean and Irano-Turanian regions, some spp. to China, Korea and Japan); N to E Africa (c. 15); additional centres in N America (c. 17 spp., including one in Hawaii) and temperate S America (30, 10 in Brazil, one endemic). Many species are widely introduced and naturalised (some are weeds); vetches are used extensively as cover crops, for forage, hay, silage, erosion control and green manure; V. faba L. (broad bean, fava bean) is a major pulse crop, also eaten green, with many cultivars in the trade; V. narbonensis L. is a minor pulse crop; several other species (e.g., V. sativa L., V. ervilia (L.) Willd. and V. villosa Roth) are cultivated as fodder; V. sativa (common vetch) is also used for medicine; in various species occasional toxicity (causing favism) is found from amino glucosides in seeds

5. SUBFAMILY CAESALPINIOIDEAE (148/4,100–4,200) ▸ three major clades, all in South America, with 9 smaller clades.

CLADE CAESALPINIOIDEAE one of 3: CERATONIEAE

5.1 CAESALPINIOIDEAE ▸ TRIBE CERATONIEAE (7/27) - outsiders Ceratonia (2; SE Mediterranean to Somalia and Arabian Peninsula), Acrocarpus (1; India, Sri Lanka, SE Asia, Malesia), Tetrapterocarpon (2; Madagascar); Arcoa (1; Hispaniola); Gymnocladus (6; E and SE Asia, E North America), Umtiza (1; E Cape).

187. Gleditsia L. Polygamous or dioecious trees or shrubs, usually armed with straight or branched thorns; leaves alternate, deciduous, once or twice pinnately compound, sometimes a leaf partly once pinnate, partly two times pinnate. 14 spp., with 2-3 in E North America, G. amorphoides (Griseb.) Taub. restricted from S Brazil, Bolivia and Cono Sur, one Caspian area, and the rest India and Japan to New Guinea.

CLADE CAESALPINIOIDEAE 2 of 3: PTEROGYNE/CASSIA-CAESALPINIA

5.2 CAESALPINIOIDEAE ▸ TRIBE PTEROGYNEAE (1/1) - a single genus.

188. Pterogyne Tul. Trees up to 30 m tall. Only one sp., P. nitens Tul. tropical South America. E Brazil, Paraguay, Bolivia, N Argentina, seasonally dry tropical to subtropical woodland and thorn scrub (including dry seasonal scrubland of NE Brazil (caatinga)), semi-deciduous subtropical forest and liana forest. P. nitens (viraro, amendoim, tipa or tipa colorado) yields an excellent timber for cabinetwork, furniture, interior finishing, turnery, flooring, railway crossties and cooperage.

5.3 CAESALPINIOIDEAE ▸ TRIBE CASSIEAE (c 19/c 700) - outsiders Latrobea (6; SW W Australia), Distemonanthus (1; tropical West Africa), Eligmocarpus (1; SE Madagascar), Koompassia (3; Malesia), Mendoravia (1; Madagascar), Storckiella (4; NE Queensland, New Caledonia, Fiji), Zenia (1; S China, Thailand, Vietnam), Kalappia (1; Sulawesi).

189. Batesia Spruce ex. Benth. Only one sp., B. floribunda Spruce ex. Benth., from Amazonia of N Brazil, Colombia, Brazil and French Guiana.

190. Cassia L. Trees up to 30 m tall. 12–13 spp. native in the Americas, mostly of the Amazon Basin but one sp. endemic to Mexico (remaining all in South America, all of then in Brazil, 4 endemics), two in SE Brazil, one in NE Brazil and two widely distributed on either side of the Panamá isthmus, including in the Antilles; 10 spp. in Africa S of the Sahara, one extending into Madagascar; one sp. endemic in Madagascar; one sp. endemic to Myanmar and Thailand; 3 spp. native to India, Sri Lanka and SE Asia are widely cultivated throughout the tropics and subtropics; 2 spp. endemic in Australia.

191. Chamaecrista Moench. Erect annuals, sometimes with xylopodium; leaves 2-ranked, once even-pinnately compound; petioles with a sessile to stalked, ± disc-shaped gland; petals yellow, some of which sometimes with a reddish spot at base. c. 350 spp., Africa, E Asia, temperate and tropical regions in New World (295, 278 in South America), highly centered in dry areas in Brazil (271, 232 endemics); 52 spp., in several states, are rare plant in Brazil, by Plantas Raras do Brasil’s book.

Chamaecrista is the largest genus of the Leguminosae subfamily Caesalpinioideae found in the savannas of C Brazil (cerrado) biome; it encompasses 117 spp., of which 43 are endemic; in this biome, species of this genus usually grow in flat areas, slopes, or hilltops, and also in rocky fields, where they stand out due to their asymmetrical flowers with lively yellow petals, viscous inflorescence or not, and elastically dehiscent fruits. Four sections in this genus.

§ sect. Absus ▸ three subsections.

§ subsect. Absus ▸ 20 spp., 17 endemics to Brazil.

§ subsect. Viscosa ▸ 169 spp., c. 160 in Brazil.

§ subsect. Zygophyllum ▸ 8 spp., Mexico to South America, 5 endemics to Brazil.

§ sect. Apoucouita ▸ 22 spp., Atlantic Forest of NE and SE Brazil and in Amazonia region of Brazil and surrounding countries.

§ sect. Baseophyllum ▸ 10 spp. 9 in rocky fields (campo rupestre) vegetation in NE and SE Brazil, with the exception of the peculiar, little-known and infrequently collected C. bucherae (Moldenke) H.S.Irwin & Barneby which is endemic to Cuba.

§ sect. Chamaecrista ▸ 134 spp., 70 in New World and 64 extra-New World, in Africa with 36 spp., 3–4 extending to Madagascar; 6 spp. endemic in Madagascar and one in Aldabra; 12 native in Australia, 5 spp. endemic in India, 3(–4) spp. in continental SE Asia, Java and New Guinea, one sp. endemic in Philippines, the genus extending into Korea and Japan.

192. Melanoxylum Schott. Small tree 5-12 m, canopy, with showy yellow flowers; inflorescence a terminal panicle; leaves imparipinnate, fruits, oblong, curved. Only one sp., M. brauna Schott, restricted from Bahia to Rio de Janeiro states, reaching in dry forests of Minas Gerais.

193. Recordoxylon Ducke. Trees. 3 spp., Amazonian Brazil (2, one endemic), Guyana, French Guiana and Venezuela, in non-flooded rain forest on terra firme, montane forest, seasonally flooded riverine forest (várzea).

194. Senna Mill. Annual or perennial, unarmed herbs or shrubs; leaves once even-pinnately compound; petiolar glands present or absent; corollas yellow or orange; 300 spp. from warm-temperate to tropical regions, especially America. 215 spp. native to the New World, c. 45 restricted to C and N America, 10 in the Greater Antilles, c. 30 restricted to NW South America (Venezuela, Colombia, Ecuador, Peru), 3 restricted to the Guianas and Venezuela, c. 30 in southern S America (Argentina, Uruguay, Paraguay, including 8 endemics in Chile), 5 endemics in Bolivia, c. 20 restricted to SE Brazil and 16 to NE Brazil, c. 25 spp. widely distributed across S America, and c. 25–30 on either side of the Panamá isthmus (either disjunctly or widely distributed); c. 18–20 spp. native to Africa (many in Somalia-Masai E and NE Africa (including 2 extending to Socotra; one sp. endemic on Socotra and 3 extending to the Middle East and Asia), others more widely distributed in dryland phytochoria), 9 spp. endemic in Madagascar, 3 spp. indigenous in Malesia, 3 spp. restricted to India, one sp. in Myanmar (Burma) and Thailand (and possibly also native in Cambodia, Laos and Vietnam); 33 spp. endemic in Australia plus 16 endemic so-called ‘form taxa’; c. 3–5 spp. widely cultivated and of unknown certain origin; Arid scrubland, rocky hillsides and deep desert sands, deciduous woodland, thorn woodland (including dry seasonal scrubland of NE Brazil (caatinga)), savannas of C Brazil (cerrado), few in Amazonian forest; some endemics on limestone (especially in Madagascar), coastal tropical.

163 spp. in South America, 82 in Brazil, 38 endemics, one of them, from Bahia state, is a rare plant in Brazil, by Plantas Raras do Brasil’s book.

S. alexandrina Mill., endemic to Brazil, or true senna is well-known as a laxative; many species are multi-purpose shrubs with numerous medicinal uses, especially as purgatives; S. alata (L.) Roxb. is universally used against parasitic skin diseases; timber of some species is used in construction and for charcoal; occasionally seeds are roasted and ground as a coffee substitute, less often used as fish poisons and for tanning leather; many are ornamentals and shade trees widely introduced throughout the tropics.

195. Vouacapoua Aubl. Trees up to 40 m tall. 3 spp., Guianas and Amazonian Brazil (2, one endemic); tropical America; tropical terra firme rain forest. V. americana Aubl., partridgewood in the N American timber trade, has rot-resistent wood, is used for house beams and is a valued commercial wood in Pará, Brazil; the wood of all species is used for cabinetwork, parquet flooring and furniture.

5.4 CAESALPINIOIDEAE ▸ TRIBE CAESALPINIEAE (27/224) - outsiders Hererolandia (1, Namibia), Gelrebia (8, C & S Africa), Hultholia (1, China to S Asia), Moullava (4, Africa, SAsia), Biancaea (6, India to Japan and Malaysia), Pterolobium (10, Africa, Arabia, SE Asia), Mezoneuron (24, Asia, Africa, Madagascar, Australia, Pacific Islands, Hawaii), Cordeauxia (1, Somalia, Ethiopia), Stuhlmannia (1, Kenia, Tanzania, Madagascar), Ticanto (15, Asia).

196. Arquita E. Gagnon, G. P. Lewis & C. E. Hughes. Small to medium-sized, often decumbent shrubs, 0.3–2.5 m in height, slender in stature, usually with glandular trichomes on various parts of the plant; young stems and inflorescence rachises red-orange to maroon. 5 spp., Ecuador to Argentina, in high Andes.

197. Balsamocarpon Clos. Shrub 1–2 m tall, with long terete branches with thin, straight, 3–5 mm long, often caducous spines. Only one sp., B. brevifolium Clos., from desert scrub, rocky hillsides in Chile; the fruit resin is used from tanning.

198. Caesalpinia L. (exc. Erythrostemon p.p., Cenostigma p.p., Denisophytum, Hoffmannseggia p.p., Libidibia p.p., Paubrasilia, Tara p.p.) Shrubs or small trees, usually 1–6 m tall, armed with curved deflexed prickles, these either in pairs at the base of leaves, or scattered along the shoots (or both), or sometimes on woody protuberances at the base of trunks and stems; young shoots terete, glabrous and eglandular. 9 spp., C. cassioides Willd. from dry valleys of Colombia, Ecuador and Peru, C. pulcherrima (L.) Sw. from S Guatemala and Mexico, and remaining 7 endemics to Caribbean.

199. Cenostigma Tul. (inc. Poincianella p.p., Caesalpinia p.p.) Unarmed multi-stemmed shrubs, small compact trees, (0.3–) 0.5–6 m, or large trees to 35 m tall, the larger trees with fluted trunks at maturity; bark smooth, or occasionally rough and flaking, brown, grey, or mottled silver or grey; young shoots terete, glabrous to pubescent, glandular to eglandular. 15 spp., 9 in South America, all of them in Brazil, 5 endemics, the majority of species are found in central and NE Brazil, including parts of the Amazon; 2 species extend around the circum-Amazonian arc of dry forests and adjacent savannas of C Brazil (cerrado), including in Paraguay, Argentina and Bolivia, and one taxon is also found in the seasonally dry inter-Andean valleys of Peru; species are also found throughout Central America, from Panamá northwards and in Mexico, extending to the Caribbean, with endemics in Cuba and Hispaniola.

200. Coulteria Kunth. Trees or shrubs, 3–20 m tall, unarmed; young twigs with a dense velvety-bronze pubescence, glabrescent; stipules not seen. 7 spp., Mexico and C America, one extending to Cuba, Jamaica and Curaçao, C. mollis Kunth to Venezuela (including Isla Margarita) and Colombia; seasonally dry tropical forest, deciduous woodland and dry thorn scrub, some species on limestone.

201. Denisophytum R. Vig. (off Caesalpinia) Shrubs to small trees, 0.5–2 (–5) m tall, armed with straight or curved, deflexed prickles, scattered along shoots and also in pairs at the petiole base; young twigs glabrous to pubescent, eglandular. 8 spp., found across North America, South America and Africa, including Madagascar, a classical highly disjunct trans-continental distribution typical of lineages occupying the succulent biome; three species are distributed in Mexico, Florida, and the Caribbean, D. stuckertii (Hassl.) E. Gagnon & G. P. Lewis is endemic to Paraguay and N Argentina, one is endemic to northern Madagascar, and the other three occur in northern Kenya, Somalia and Arabia. An evaluation of species limits is needed in this group.

202. Erythrostemon Klotzsch. (inc. Caesalpinia p.p.) Shrubs or small to medium-sized trees varying from (0.5–) 1–12 (– 20) meters tall, occasionally suffrutices, rarely armed; bark variable, smooth or rough, sometimes exfoliating, grey, greyish white, pale brown or reddish brown, often with white or black pustular lenticels. 31 spp., 22 in S U.S.A. and Mexico, one in Caribbean, eight occur in South America, with E. calycina (Benth.) L.P.Queiroz endemic in dry seasonal scrubland of NE Brazil (caatinga) in Bahia and Pernambuco states, and the other seven in Argentina, Bolivia, Chile, and Paraguay, in seasonally dry tropical and subtropical semi-arid thorn scrub (including caatinga), spiny cactus scrub, woodland and grassland. E. gilliesii (Hook.) Klotzsch (yellow, or desert bird of paradise) is widely cultivated as a garden ornamental and is used in revegetation.

203. Guilandina L. Lianas and scandent shrubs characterised by unisexual flowers (morphologically the flowers of at least some species appear to be hermaphrodite, but lack pollen in the anthers and are thus cryptically pistillate. 7-18 spp., pantropical, from as far N as Japan south to S Africa, one in China (inc. Taiwan), India, Myanmar, Indochina, Hong Kong, one endemic to Madagascar, one in Australia, and 13 spp. in New Word, 9 in the Caribbean, two in South America (both in Brazil); coastal sands and thicket, secondary forest, lowland rain forest, some on limestone, 0 – 1770 m.

204. Haematoxylum L. Multi-stemmed shrubs to 3 m, to medium-sized trees, 3–15 m in height, armed with scattered straight conical spines, 0.5–1.5 cm long on shoots, and the short, lateral shoots spinescent; mature trees with conspicuously fluted trunks, shrubs often with ribbed branches; young stems reddish brown to grey, glabrous to pubescent, eglandular (some stalked glands). 5 spp., two in Central America (El Salvador to Costa Rica), Mexico, Colombia and Venezuela (only H. brasiletto H. Karst.), and the Caribbean (perhaps introduced), two endemic to Mexico and one disjunct in Namibia.

The heartwood of H. campechianum L. (campeche, logwood), is the source of a colourless chemical haematoxylin which on oxidation turns to haematein, a commercial dark violet dye used for wool, silk, cotton, fur, leather, bone and synthetic fibre dying, and with iron chromium mordants to obtain red and black; also used as a stain in microscopical preparations (particularly to show up cell nuclei), an ink for writing and painting and the rich red colour has been used to adulterate wine. The species has minor medicinal uses; H. brasiletto H.Karst. (brazilette), produces brasil in used as a bright red dye; both New World species used as ornamentals and living hedges (H. campechianum cultivated in Africa and Madagascar); minor uses for furniture and carpentry; bee flowers yield a high quality honey.

205. Hoffmannseggia Cav. (inc. Caesalpinia p.p.) Perennial woody herbs, most species forming a basal rosette, or subshrubs to 3 m, unarmed, often arising from bud-bearing and tuberous roots, shoots pubescent and with gland-tipped trichomes. 26 spp., disjunct: H. burchellii (DC.) Oliv. in S Africa, 11 in SW U.S./N Mexico, 13 within Peru to Chile and Argentina, and one in both areas; some SW U.S. species have tubers that are edible when roasted.

206. Libidibia (DC.) Schltdl. (inc. Caesalpinia p.p.) Small to medium-sized or large unarmed trees, 6–20+ meters in height; bark hard, smooth, with a patchwork of shades of grey, white and pale green, often referred to as snake skin bark. 7 spp., one species in Mexico, one widely distributed in Brazil, one in Colombia, Venezuela and the Antilles, one in Colombia, Ecuador and Peru, one in Paraguay, Bolivia, Argentina and SW Brazil, one (L. monosperma E.Gagnon & G.P.Lewis, previously in the monospecific genus Stahlia) endemic to Puerto Rico and the Dominican Republic, and L. coriaria (Jacq.) Schltdl. widely distributed throughout Mexico, Central America, the Caribbean and NW South America.

Used as ornamental park and street trees; pods rich in tannins and used commercially in the tanning industry, pods of some species also used for animal fodder, ink and local medicines (due to astringent properties of the tannins); wood prized in turnery, and for parts of guitars and violins, that of Caesalpinia glabrata Kunth and C. paraguariensis (D.Parodi) Burkart (both species of Libidibia (partridge wood) but without combinations in the genus) used in decorative inlay and cabinet work; some species used in heavy construction (railway sleepers, beams, bridge supports), for tool handles and as firewood; L. coriaria differs in having white (not yellow) flowers; exserted stamens with red anthers; rough, fissured bark which flakes away in vertical strips (not smooth, patchwork bark exfoliating in woody plates), and flowers lacking the tentacle-like papillae of the other species; all Libidibia species have indehiscent woody, black or dark brown pods, but those of L. coriaria are distinct in being curled and twisted.

207. Lophocarpinia Burkart. Shrub. Shrub 0.5 (– 3) m tall, armed with scattered straight, conical, 2–5 mm long spines on shoots; leaves and inflorescences crowded on brachyblasts; shoots glabrous, reddish, the lateral ones sometimes, spinescent. Only one sp., L. aculeatifolia (Burkart) Burkart; Paraguay, N Argentina, possibly also occurring in Mato Grosso do Sul, Brazil, in seasonally dry tropical to subtropical woodland (chaco).

208. Paubrasilia E. Gagnon, H. C. Lima & G. P. Lewis. (off Caesalpinia) Medium sized to large trees, 5 – 35 m tall, armed with small to large upturned prickles, these usually arising from woody protuberances, one – 20 mm long (the prickles often sparse or lacking on more mature specimens and larger, older branches); bark chestnut brown to almost black with greyish pustular lenticels, flaking in large woody plates; heartwood red, with the trunk exuding a red sap when injured; stipules lanceloate, acute to acuminate, caducous. Only one sp., P. echinata (Lam.) E. Gagnon, H. C. Lima & G. P. Lewis, endemic to E Brazil, from Rio Grande do Norte to Rio de Janeiro in coast, also in Minas Gerais. Widely cultivated in Brazil as an ornamental street or park tree, and sometimes in plantations; occur in dry coastal cactus scrub often on rocky outcrops, inland in Atlantic Forest, and in tall Atlantic sandy coastal shrublands (restingas) on well-drained sandy soil.

209. Pomaria Cav. Small shrubs, subshrubs or perennial herbs, with a moderate to dense indumentum of simple curled hairs, sometimes also scattered plumose trichomes, intermixed with sessile, oblate glands (drying black) on stems. 16 spp., 9 in SE U.S.A., C and N Mexico, 4 in S America (dry open areas, SE Brazil (3, one endemic), Uruguay, Paraguay and Argentina), 3 in S Africa (Namibia, Botswana and S Africa), mainly in subtropical dry areas of grassland and wooded grassland and in degraded sites, many on limestone.

210. Stenodrepanum Harms. Suffrutescent shrub, (10–) 20–40 cm tall, with bud-bearing and occasionally tuber-forming roots; glabrous, with globose sessile glands scattered along the branches. Only one sp., S. bergii Harms, endemic to C and W Argentina, in subtropical wooded grassland and scrub, especially on salt pans.

211. Tara Molina. (inc. Caesalpinia p.p.) Shrubs or trees, 3–5 (– 8) m tall, armed with deflexed prickles on the shoots; twigs glabrous to puberulent. 3 spp., one restricted to Mexico, one in Mexico (Yucatan), Nicaragua, Cuba, Haiti, Jamaica and the Bahamas, T. spinosa (Molina) Britton & Rose in S America (Ecuador and Peru) and cultivated both within and outside (including Malta, Canary Islands, India, E and NE Africa) its native range; seasonally dry tropical forest to semi-arid thorn scrub.

Planted as living fencelines, the bark of two species with medicinal properties, that of T. spinosa (Molina) Britton & Rose (tara, spiny holdback) rich in tannins and used in Peru for tanning leather, dyeing and ink production (various commercial products sold), also for firewood and gums.

212. Zuccagnia Cav. Shrubs, 1–5 m. Stipules caducous; leaves alternate, pinnate, (2–) 3–5 (– 6) cm long. Only one sp., Z. punctata Cav., in the Andes in Chile and Argentina, in dry temperate upland and montane brushland, thicket and sandy plains. Minor local medicinal uses; the leaves yield a yellow dye.

CLADE CAESALPINIOIDEAE 3 of 3: MIMOSA/DIMORPHANDRA ALLIANCE

5.5 CAESALPINIOIDEAE ▸ DIMORPHANDRA CLADE A (6/43–63) - outsiders unvailable.

213. Campsiandra Benth. Trees up to 40 m tall. 20 spp., S America, mainly in the Amazon and Orinoco basins (13 endemics to Venezuela, 5 in Venezuela and Brazil (of which 2 extend to Colombia, one to Bolivia and one to Colombia, Bolivia and Peru), one endemic to Peru, one restricted to the Guianas); tropical riverine and swamp forest (both black and white water rivers) on alluvial plains, white sand beaches and embankments; 7 spp. in Brazil, none endemics.

214. Dimorphandra Schott. Trees up to 50 m tall. 26 spp. from S America (14 in Amazonian Brazil (6 of which extend to C Brazil, the Guianas, Venezuela, Colombia or Peru), one endemic to Peru, one to Venezuela, 5 centred in the Guianas (extending into N Brazil, Venezuela and Colombia), one in E and NE Brazil, 2 in SE Brazil, and 2 widely distributed throughout Brazil and extending into Bolivia and Paraguay); 22 spp. in Brazil, 11 endemics.

Tropical Amazonian rain forest on terra firme, periodically inundated forest, permanently humid forest, savannas of C Brazil (cerrado), dry seasonal scrubland of NE Brazil (caatinga) and open sandy sites in forest. The wood of some species (e.g., D. exaltata Schott) used in cabinetwork and for tool handles; the fruits of some species highly toxic to cattle (e.g., D. mollis Benth. and D. gardneriana Tul.); the bark of D. mollis used in tanning; several species used as medicine (e.g., for bioflavonoids) and ornamentals.

215. Dinizia Ducke. Trees up to 88m, several specimens up to 88.5m tall. Two spp., D. excelsa Ducke from Amazonian S America (Brazil, Guyana), and D. jueirana-facao G.P. Lewis & G.S. Siqueira endemic to small area in Atlantic Forest of Espírito Santo state, Brazil; non-inundated (often riverine) rain forest, known only by 24 individuals.

D. excelsa is known as angelim vermelho, red angelim, angelim pedra, parakwa, and is used for timber (heavy construction, flooring, stairs, ship building, posts and railway sleepers) and in reforestation; this species reaches up to 88.5m tall, making the tallest vascular plant in South America, second largest plant in Neotropics, the tallest angiosperm of New World and the tallest non-Malvid angiosperm worldwide.

216. Mora R.H.Schomb. ex Benth. Trees up to 35 m tall. 6 spp., C America, N South America and the Greater Antilles: two from W Surinam through Guyana to the Orinoco Delta in Venezuela (one extending to Trinidad), M. oleifera (Triana ex Hemsl.) Ducke from Nicaragua to Ecuador, M. paraensis (Ducke) Ducke from the delta of the Amazon in Brazil, and two endemic to the Hispaniola, in tropical riverine forest, periodically inundated or not, or swampy areas, broad-leaved humid forest, one sp. just behind the mangrove zone, hill slopes.

M. excelsa Benth. and M. gonggrijpii (Kleinhoonte) Sandwith (mora, morabukea, nato, pracuuba) are major timber trees in the Guianas, especially used in heavy construction, industrial flooring and for charcoal; the seeds of M. oleifera are thought to be the largest (18 x 15 x 8 cm) dicotyledonous seeds known, and third in over flowering plants (after Lodoiscea maldivica and Cocus nucifera L., both Arecaceae) they are a local source of a red dye; the seeds of M. excelsa are also large (up to 12 ✕ 7 cm), being the eighth of all plants; none of this spp. occur in Brazil, however the Brazilian endemic M. paraensis (Ducke) Ducke is the largst of all seeds in country, with 8.3 cm long ✕ 4.5 cm wide ✕ 3.8 cm thick.

5.6 CAESALPINIOIDEAE ▸ SCHIZOLOBIEAE (Delonix Clade, 8/45–49) - outsiders unvailable.

217. Parkinsonia L. (inc. Cercidium) Shrubs and trees. 11-12 spp., 3 in S U.S.A. and Mexico (two extending to Baja California and one widely cultivated throughout the tropics), two from Peru, Bolivia and Argentina, three widely distributed from Mexico through Venezuela, Ecuador and Peru to N Argentina, two of them in Brazil, none endemics; 4 in Africa (one endemic to Kenya, one to Somalia, one more widely distributed in the E and NE and one in S and SW Africa); seasonally dry tropical bushland, semidesert scrub, coastal dunes and flood plains.

P. aculeata L. (Jerusalem thorn, paloverde) is widely cultivated throughout the tropics as an ornamental and thorn hedge but can escape to become a weed; also used for medicine and human food (flowers eaten in Senegal, the seeds of P. africana Sond. used as a coffee substitute).

218. Peltophorum (Vog.) Benth. Trees up to 40 m tall. c 5 spp., tropical and subtropical regions pantropical (into the subtropics) with three native to the Neotropics (one in E Brazil, N Uruguay, NE Argentina, Paraguay, Bolivia and the Caribbean, the two restricted to Venezuela, one of them although this probably conspecific with the widely distributed), one native to S Africa, one in Malesia (Sabah, Sarawak, Kalimantan), and 2 widely distributed in Asia (Thailand, Cambodia, Laos, Vietnam, and Peninsula Malaysia, one extending to Sumatra and Java, the other to Sri Lanka, Philippines, and N Australia); many species widely cultivated, seasonally dry tropical and evergreen (mainly lowland) forest, beaches and mangrove forest, coastal monsoon vine thicket, flood plains and tidal flats, bushveld and woodland.

Widely cultivated for ornament and shade; some species used for carpentry, planking, furniture and fuelwood; leaves used for livestock fodder; some medicinal uses and dyes.

219. Schizolobium Vogel. Trees up to 30 m tall. Only one sp., S. parahyba (Vell.) Blake, from tropical America. Atlantic SE Brazil (one variety), Amazonian Brazil, Bolivia, Paraguay, Peru, Ecuador, Colombia and Venezuela, through C America to SE Mexico (the other, more widely distributed variety); tropical rain forest, mixed and secondary forest.

S. parahyba (Vell.) S.F.Blake var. amazonicum (Ducke) Barneby (quamwood, guapuruvu, reach-for-the-sky, feather-duster tree) is widely planted as an ornamental (including outside the New World) and as shade trees (e.g., for coffee).

5.7 CAESALPINIOIDEAE ▸ TRIBE TACHIGALEAE (3/c 70) - all genera in South America.

220. Arapatiella Rizzini & A. Matos. Tree. Only one sp., A. psilophylla (Harms) R.S.Cowan, endemic to Atlantic Forest of Bahia state in NE Brazil, one of them is a rare plant in Brazil, by Plantas Raras do Brasil’s book.

221. Jacqueshuberia Ducke. Trees. 7 spp., two in Amazonian Brazil (3 in Brazil, two endemics), one in Guyana, 2 in E Venezuelan, one in Colombia and one in Peru, in tropical seasonally inundated riverine forest on white sand, montane forest on sandstone, savanna, open sandy areas by black-water streams of the Amazon (north-amazonic white-sand savannas known as campinaranas). A wide range of flower colour, including red, purple and yellow exists in this small genus.

222. Tachigali Aubl. Trees, sometimes monocarpic, up to 40 m tall. 84 spp. of Neotropics, 83 in S America with main centre of diversity in the Amazon Basin, but extending S to Paraguay and Argentina and N into C America (only two, one endemic in Costa Rica, one more widely distributed); 15 in E Venezuelan, of which 3 endemic, 14 in Peru, of which 5 endemic, several spp. widely distributed in S America), mainly of tropical rain forest, lowland montane forest, seasonally flooded and non-flooded evergreen lowland forest and woodland, gallery and riparian forest, sometimes on white sands, also in savannas of C Brazil (cerrado) and rocky grasslands (campos rupestres). 60 spp. in Brazil, a exact half endemics - 5 of them, in Amazonas and Rio de Janeiro states, are rare plants in Brazil, by Plantas Raras do Brasil’s book. 25 spp. of this genus, some from South America, are myrmecophytes.

Various species used for timber (djedoe, yawaredan, suicide tree (because some species monocarpic)) for construction, canoes and charcoal; the bark of T. tinctoria (Benth.) Zarucchi & Herend. used in tanning and as a dye.

5.8 CAESALPINIOIDEAE ▸ TRIBE DIPTYCHANDROIDEAE (2/10) - both genera in South America.

223. Diptychandra Tul. Two spp., D. aurantiaca Tul.C, E and NE Brazil, Paraguay and Bolivia, and D. granadillo C. Romero & Arbeláez from mountains of Colombia; seasonally dry tropical forest, gallery forest, savannas of C Brazil (cerrado), dry seasonal scrubland of NE Brazil (caatinga), rocky grassland and chaco.

224. Moldenhawera Schrad. Trees or shrubs with an indumentum of principally biramous hairs; stipules compound or rarely setaceous; leaves compound, pinnate, bipinnate or partially bipinnate; inflorescences subcorymbose racemes, generally aggregated into pseudopanicles; second order bracts caducous; flowers tetramerous or pentamerous, with fleshy reflexed sepals; petals yellow or rarely pink, clawed, the limb with the margin crenate or lobed, usually fringed. 12 spp., scattered in several states in E Brazil, centerd in Bahia state, 5 of them, in Bahia, Maranhão e Espírito Santo states, are rare plants in Brazil, by Plantas Raras do Brasil’s book.

5.9 CAESALPINIOIDEAE ▸ TRIBE MIMOSEAE (82/3.035–3.150) - 78/3.270–3.320. Tropical, subtropical and warm-temperate. In August 2011 the long discussion on the split of the genus Acacia into five new genera has been ended; Acacia s.s. has c. 1045 spp. in Australia (of which 941 endemic and 7 extending to Asia, and c. 100 spp. new and yet to be described fide Maslin et al., 2003), 7 in the Pacific, 3 confined to Asia, 2 in Madagascar and Mascarenes. Archidendron Clade (8/1,000-1,100, Madagascar, tropical Asia, Australia, islands in the Pacific) do not occur in South America.

∎ IN PARAPHYLETIC GRADE OR ISOLATED PHYLOGENETIC POSITION - outsiders Cylicodiscus (1, Central Africa), Fillaeopsis (1, W and SW Africa), Mariosousa (13, SW U.S.A. to Central America), Newtonia (12-13, tropical Africa and America), Xerocladia (1, Namibia, Northern Cape).

225. Lachesiodendron P.G.Ribeiro, L.P.Queiroz & Luckow. Tree (2)3–20 m tall, usually multi-stemmed from the base, rarely flowering as shrubs ca. 1.5 m tall. Only one sp., L. viridiflorum (Kunth) P.G.Ribeiro, L.P. Queiroz & Luckow, NE Brazil, N Paraguay and the adjacent Brazilian Mato Grosso do Sul State, Bolivia, NW Argentina, NW Peru, W Ecuador, the Caribbean coast of Colombia and Venezuela, S and W Mexico and N Guatemala.

226. Neltuma L. (inc. Prosopis p.p.) Spiny, erect to prostrate subshrubs, shrubs and small trees, (0.1–) 4–10 (–20) m high, usually with a short trunk to 40–60 (-100) cm diameter. 34 spp. in New World, 30 spp. in South America, centred in Argentina, Chile, Paraguay to E Bolivia, Uruguay and S Brazil (4, none endemics) with a minor centre in the tropical Andean region from Peru, Ecuador and Colombia; 2 spp. widely distributed in Neotropics; 4 spp. in Mexico–SW U.S.A.); cultivated and naturalised worldwide; tropical to warm temperate seasonally dry forest, woodland, wooded grassland, semi-xerophytic woodland and shrubland, thorn scrub and desert, on sandy plains or hills, ravines and along dry stream beds.

Used since ancient times as human food and drink (from the sweet fleshy pods), timber (e.g., for construction, fence posts and furniture), livestock fodder, charcoal, firewood, shade trees, ornamentals, gums, dyes for tanning, medicine, coffee substitute, bee plants and desert reforestation; major species are N. juliflora (Sw.) Raf. and N. pallida (Humb. & Bonpl. ex Willd.) C.E. Hughes & G.P. Lewis, known as mesquite, algarroba and screw bean; some species are invasive and pernicious weeds; one sp., N. rusciflora (Griseb.) C.E. Hughes & G.P. Lewis, occurs in Chaco of Argentina and Bolivia, with one population with two trees in W Pernambuco state (uncommon distribution).

227. Parasenegalia Seigler & Ebinger. (inc. Lysiloma) Trees, shrubs, or lianas, some to 25 m; prickles and stipular spines absent; leaves alternate, bipinnate; inflorescence a globose head or cylindrical spike, solitary to small clusters in leaf axis or in large pseudoracemes or pseudopanicles of globose heads; flowers bracteate, actinomorphic. 7 spp., tropical areas of the New World, three species restricted to the Caribbean, one in Belize and Guatemala, one in Guianas, two in western South America, and P. santosii (G.P. Lewis) Seigler & Ebinger in savannas from near sea level to 400 m in Bahia and Minas Gerais, Brazil.

228. Plathymenia Benth. Trees up to 30 m tall. Only one sp., P. reticulata Benth., from Brazil, Paraguay, E Bolivia, and Argentina, seasonally dry tropical forest (and secondary vegetation derived from moist forest; savannas of C Brazil (cerrado), rocky grassland (campo rupestre), bushland and scrub).

Used commercially for timber (vinhatico) in flooring, construction, posts and furniture; also for reforestation, shade trees, firewood and medicine.

229. Pseudosenegalia Seigler & Ebinger. Shrubs or small trees, some to 12 m; prickles and stipular spines absent; leaves alternate or clustered on short shoots, bipinnate; inflorescences a cylindrical spike, one to 2 clustered in leaf axils; flowers bracteate, sessile, actinomorphic, sympetalous, 5-merous. Two spp., known in dry scrub and thorn-scrub vegetation 1,300 – 3,300 m altitude range in Bolivia.

230. Senegalia Raf. (inc. Acacia) Shrubs, trees, or lianas, unarmed or armed with prickles, but without stipular spines. 219 spp., pantropical, 69 in Africa and Madagascar, of which 7 extend to Asia, 36 restricted to Asia, 2 in northern Australia and the Pacific (one extending to Asia); 99 in New World of which 35 in N and C America, 72 in S. America (18-19 spp. in Peru and Colombia each), 69 in Brazil, 40 endemics – 3 of them, all in Bahia state, are rare plants in Brazil, by Plantas Raras do Brasil’s book.

231. Strombocarpa (Benth.) Engelm. & A. Gray (inc. Prosopis p.p.). Low spiny, sometimes creeping, shrubs or small trees, 0.15–3 (–18) m high, multi-stemmed from the base or sometimes with a short trunk to 10–30 (–45) cm diameter. 10 spp., markedly bicentric amphitropical distribution in arid and semi-arid regions of southern U.S.A. and northern Mexico (3), and in south-central Peru to Argentina and Chile (7).

∎ PHYLOGENETIC POSITION UNCERTAIN - outsiders Chidlowia (1, W tropical Africa), Indopiptadenia (1, India, Nepal).

232. Cedrelinga Ducke. Tree with a open, rounded crown up to 50 m tall. Only one sp., C. cateniformis Ducke, from French Guiana to Bolivia, N Brazil, mainly Amazonian lowland tropical rain forest to seasonally dry forest, especially along streams, 50–250 (–800) m; mature trees can attain 60 (–66)m.

This species (cedrorana, tornillo, huayracaspi) is used in agroforestry for nitrogen fixing, fuelwood, medicine and timber (for construction, boat building, frames and furniture; a substitute for mahogany).

233. Pseudosamanea Harms. (inc. Albizzia p.p.) Trees. Three spp., one endemic to Cuba, and two other widely distributed from SW Mexico to Peru and Venezuela, seasonally dry tropics. P. guachapele (Kunth) Harms occurs in drought-deciduous woodland to 1,000 m, P. cubana (Britton & Rose) Barneby & J.W.Grimes in wooded grassland (palm-savanna) and along water courses below 50 m. P. guachapele (guachapele, frijolillo) is used for timber in shipbuilding (planking, ribs, decking), railway sleepers, general construction, flooring, decorative veneers and furniture, and as livestock fodder.

∎ XYLIA CLADE (6/37-39) ▸ outsiders Adenanthera (13; tropical Asia and Australia, islands in the Pacific), Amblygonocarpus (1, tropical Africa to Namibia and Botswana), Calpocalyx (11, tropical W Africa), Pseudoprosopis (7, tropical Africa), Tetrapleura (1-2, tropical Africa), Xylia (1-2, tropical to S Africa, tropical Asia).

234. Pentaclethra Benth. Trees. 3 spp., two in WC Africa and P. macroloba (Willd.) Kuntze in S and C America (Amazonas north to C America and Caribbean), tropical lowland rain forest (often riparian) and secondary vegetation types; seasonally dry woodland and wooded grassland. Considered a multi-purpose tree suitable for agroforestry; P. macrophylla (oil bean tree, ugba, owala) is used for timber (construction), firewood, charcoal, medicine, fibre, dyes, oils, as soap substitutes, fish poisons, human food (cooked seeds) and ornamentals; the strongly elastic pods are used as soles for footwear; timber of P. macroloba (Willd.) Kuntze (gavilán, paroa-caxi) is used as a substitute for mahogany.

∎ ENTADA CLADE (3/44) ▸ outsiders Aubrevillea (2, tropical W and C Africa), Piptadeniastrum (1, C Africa).

235. Entada Adans. Trees, shrubs and lianas. 41 spp., 28 only in Africa and Madagascar, 8 endemic to Indo-China and Malesia and Australia, two widely distributed and dispersed by ocean currents (E. rheedii Spreng. throughout the Palaeotropics and E. gigas (L.) Fawcett & Rendle in Africa to Neotropics), E. polystachya (L.) DC. and E. polyphylla Benth.from Mexico to C Brazil and Caribbean, and E. simplicata (Barneby) Sch. Rodr. & A. S. Flores, endemic to Roraima state in N Brazil; tropical lowland and riverine rain forest, seasonally dry forest, woodland and wooded grassland, bushland, thicket and dry scrub.

Various species used as livestock fodder, ground cover, green manure, fibre (in rope and storage bins), medicine, fish poisons, soap substitutes, firewood, charcoal and in traditional ceremonies.

The large round drift seeds of the species listed above emerge from some of the biggest pods in the Leguminosae (over one metre in length) and are often used to make jewellery (‘ sea hearts ’); the world's largest legume fruits (bean pods) are produced by this genus; the longest pods of the Central American E. gigas may be up to 5 feet long (1.5 m); this gigantic woody vine is truly like Jack's fabulous bean stalk. In Costa Rica it is called ‘monkey ladder’ or ‘escalera de mono’; the woody seeds of E. gigas are called ‘sea hearts’ and are often washed down streams to the sea where they drift across the ocean to distant continents.

∎ DICHROSTACHYS CLADE (13/c. c. 105) ▸ outsiders Alantsilodendron (9), Calliandropsis (1, Mexico), Dichrostachys (25, tropical Africa, Madagascar, S and SW Asia to Australia), Gagnebina (5, Madagascar, the Comoros, Mascarene Islands), Kanaloa (1, Hawaii), Lemurodendron (1, Madagascar), Schleinitzia (2, Malesia, Vanuatu to Tahiti, Guam).

236. Desmanthus Willd. Unarmed herbaceous or semi-shrubby perennials, sometimes with xylopodium; leaves twice even-pinnately compound, with a petiolar gland between the lowest air of pinnae (gland sometimes minute). 26 spp. of warm areas of the Americas, U.S.A. to Mexico and C America (14 in Mexico (7 endemic); 3 endemic to U.S.A.); S America (8, inc. 2 from SE Brazil, Paraguay, N Argentina; 2 disjunct between SE U.S.A.–E Mexico and the SE Brazil, Paraguay, N Argentina region); 3 more widely distributed in the New World including the Caribbean, but absent from the Amazon Basin. 5 spp. in Brazil, none endemics.

D. pernambucanus (L.) Thell. (long misinterpreted as D. virgatus (L.) Willd. (which is nevertheless still a good species)) is a widely distributed pantropical weed; genus also used as ornamentals, cattle fodder and in erosion control; D. illinoensis (Michx.) MacMill. (Illinois or prairie bundleflower, prairie mimosa, spider bean) is used for human food (leaves, cooked seeds), medicine and is a potential pulse crop.

237. Leucaena Benth. Trees or shrubs. 16 spp., Mexico (10 endemics with 2 extending to S U.S.A. (Texas and New Mexico) and 4 to C America); 4 endemic spp. in C America; two sp. in S America, from N and W coastal regions S to Peru; one sp. pantropical; tropical and subtropical seasonally dry forest, semi-arid thorn scrub forest, to warm temperate open habitats.

L. leucocephala (Lam.) de Wit is cultivated pantropically and has become naturalised and weedy in many areas; this and other species are used for livestock feed, green manure, timber (for construction, firewood and charcoal), small wood products, soil conservation (ground cover and reforestation) and human food (unripe pods and seeds).

238. Mimozyganthus Burkart. Shrubs or small trees. Only one sp., M. carinatus (Griseb.) Burkart, SE Bolivia, SW Paraguay, Argentina, in tropical and subtropical arid and semi-arid scrub bushland (chaco vegetation), often associated with cacti.

239. Neptunia Lour. Herbs. Some have sensitive leaves (leaflets folding together on being touched) as in Mimosa. 12 spp., Australia (4 endemic), Papuasia through Malesia to Indo-China (2); India and Sri Lanka (1), one sp. pantropical; New World has 5 spp. from S U.S.A. (1), Mexico, C America, Caribbean south to Paraguay and N Argentina, tropical to warm temperate open woodland, wooded grassland and grassland, floodplains, swamps and other wet areas. 5 spp. in South America, two in Brazil, both widely distributed.

N. oleracea Lour. is often a troublesome weed (with sensitive leaves); used for medicine, human food (young stems, leaves and pods are eaten) and as soil binders; produced as a cash crop in SE Asia. Neptunia is morphologically unlike any other mimosoid because of its (semi-)aquatic lifestyle.

240. Piptadeniopsis Burkart. Shrubs or small trees. Only one sp., P. lomentifera Burkart, endemic to Paraguay, subtropical seasonally dry thorn forest, thicket and scrub (chaco).

241. Prosopidastrum Burkart. Shrubs. 7 spp., disjunct between Baja California in Mexico (1) and Argentina up to Bolivia (6), subtropical xerophytic bushland, thicket, grassland and semi-desert.

∎ PARKIA CLADE (3/197) ▸ all genera in South America.

242. Anadenanthera Speg. Trees. Two spp., through N S America to Peru, Bolivia, Argentina and Paraguay, both Brazilian; probably introduced in the W Indies.

A. peregrina (L.) Speg. (cohoba, niopo or yopo) is one of the classic hallucinogens of the Americas, taken as snuff made from the crushed seeds; A. colubrina (Vell.) Brenan (angico preto, cebil, curupay) is used for timber (heavy construction, flooring, railway sleepers, turnery) and leather tanning; seasonally dry tropical to subtropical riverine forest and forest margins to savannas of C Brazil (cerrado) and wooded grassland (savanna), from a wide range of habitats; often planted near villages. A. peregrina (L.) Speg. (cohoba, niopo or yopo) is one of the classic hallucinogens of the Americas, taken as snuff made from the crushed seeds; A. colubrina (Vell.) Brenan (angico preto, cebil, curupay) is used for timber (heavy construction, flooring, railway sleepers, turnery) and leather tanning.

243. Parkia R. Br. Trees without spines or prickles, up to 50 m tall; leaves 2-pinnate; pinnae (3)4–11(–14); leaflets in 10–18(–28) pairs; inflorescence capitate, shortly claviform, solitary or paniculate; flowers in upper part of heads bisexual, in lower part male or neuter. 34 spp., pantropical, but with three disjunct centres of diversity in S America (20, centred in Amazonia (but extending from Honduras in the north to SE coastal Brazil in the south), Africa-Madagascar and the Indopacific region; 3 spp. in WC to SE Africa and one sp. in Madagascar; c. 12 spp. in Asia from NE India (1), Indo-China and China (1), Malesia (c. 5, some extending to Indo-China), Papuasia (1), Micronesia (2) and Fiji (1); one sp. widely distributed in SE Asia. Tropical lowland rain forest (riparian, swamp and non-inundated), hill forest, seasonally dry forest, savannas of C Brazil (cerrado), wooded grassland and Atlantic sandy coastal shrublands (restingas). Of the three sections in Parkia only sect. Parkia is pantropical, the other two being restricted to the Neotropics. Brazil has 18 spp., 5 endemics.

P. speciosa Hassk. is used as human food in SE Asia; mature and slightly immature green seeds are eaten as a vegetable (petai, sataw bean, chou dou), sold fresh as bunches of strap-shaped pods in markets and as pods or loose seeds, eaten fresh or tinned, from supermarkets; seeds are also fermented and the pulpy endocarp makes a refreshing drink; other species are used extensively as food in W Africa (e.g., P. biglobosa (Jacq.) G.Don, or néré, African locust bean); also used as cattle fodder, cordage, shade trees, medicine and the timber in plywood manufacture, construction, utensils and as firewood.

244. Vachellia Wight. & Arn. Shrubs to trees, sometimes with roots crown; c. 161 spp., pantropical, 73 in Africa and Madagascar of which c. 15 extend to Asia, 21 restricted to Asia, 7 in Australia and the Pacific; 61 spp. in the New World, of which c. 35 in N and C America and c. 14 in South America, 3 in Brazil, one endemic: V. ibirocayensis (Marchiori) Deble & Marchiori; this species is a shrub 1-2 m high, bark brown, grows on stony grasslands and ravines near to Ibirocai and Ibirocai-mirim rivers in Rio Grande do Sul state, being probably endemic in this region; the populations are fragmented and with few individuals.

∎ STRYPHNODENDRON CLADE (7/45) ▸ all genera in South America.

245. Gwilymia A.G. Lima, Paula-Souza & Scalon (off Stryphnodendron). Trees 2.5–40 m tall; branches unarmed, not odoriferous, smooth, usually lenticellate, young shoots and leaves glabrescent, pubescent, or tomentose and covered with reddish granular trichomes. 7 spp., Amazon rainforest, seasonal forests and savannas of Bolivia, Brazil, French Guiana, Guyana and Suriname.

246. Marlimorimia L.P. Queiroz, L.M. Borges, Marc.F. Simon & P.G. Ribeiro (inc. Pseudopiptadenia p.p.). Unarmed trees; Leaves bipinnate; petiole with an extrafloral nectary well below the first pair of pinnae, close to the pulvinus, always below mid-petiole; pinnae 5–10 to many pairs per leaf. 6 spp., three endemic to coastal forests of SE Brazil, and remaining three from N Brazil to Costa Rica.

M. pittieri (Harms) L.P. Queiroz & L.M. Borges is used for timber (carbonero) in heavy construction, furniture, flooring, posts, turnery and railway sleepers; tannins are extracted from the bark.

247. Microlobius C.Presl. Shrubs or small trees. Only one sp., M. foetidus (Jacq.) M. Sousa & G. Andrade, disjunct in W and S Mexico to Honduras, and in S Brazil (only Mato Grosso do Sul state), Bolivia, Venezuela, Paraguay and Argentina, tropical to subtropical seasonally dry forest, thorn scrub and seasonally wet wooded grassland. Used for traditional medicine; the plant has a strong garlic or onion-like smell.

248. Naiadendron A.G. Lima, Paula-Souza & Scalon (off Stryphnodendron). Trees 8–30 m tall; branches unarmed, strongly striate, castaneous, apex yellow-tomentose and covered with ferruginous granular trichomes, not odoriferous. Only one sp., N. duckeanum (Occhioni) A.G. Lima, Paula-Souza & Scalon, endemic to the Amazon rainforest, being recorded from the Brazilian states of Acre, Amazonas and Rondônia, on clay or sandy soil in ombrophilous and terra firme forests.

249. Parapiptadenia Brenan. Trees and shrubs up to 35 m tall. 6 spp., SE & E Brazil (all spp., three endemics), Bolivia, Paraguay, Uruguay, and Argentina, tropical coastal and Atlantic sandy coastal shrublands (restingas), secondary growth forest, woodland and scrub. All species are used as commercial timber (chari, angico, anchico colorado) in high quality furniture and construction (e.g., P. rigida (Benth.) Brenan); also used for gums, a mucilage constituent of medicines and in agroforestry.

250. Pityrocarpa Britton & Rose. (inc. Pseudopiptadenia p.p.) Small trees or shrubs. 7 spp. from Mexico to Brazil (all spp., 3 endemics) and Paraguay, mainly in Brazilian Atlantic rainforests (3), in the northern Amazonian rainforests (1), in seasonally dry tropical forests and woodlands in dry seasonal scrubland of NE Brazil (caatinga, 3), western Mexico (1) or in Venezuelan savannas and Paraguayan Chaco (1), in tropical coastal rain forest, gallery forest, secondary forest, woodland, savannas of C Brazil (cerrado) and dry seasonal scrubland of NE Brazil (caatinga). 9 spp. in Brazil, 6 endemics.

251. Stryphnodendron Mart. (exc. Gwilymea, Naiadendorn) Trees and suffrutices, sometimes basal burls. 28 spp. of S America (14 in Amazonia, with one sp. to Nicaragua; 14 spp. in C & SE Brazil, one sp. to Bolivia and Paraguay; 3 in NE Brazil); tropical rain forest and riparian forest, seasonally dry forest, savannas of C Brazil (cerrado), wooded grassland and dry seasonal scrubland of NE Brazil (caatinga). 22 spp. in Brazil, 15 endemics.

The bark of S. adstringens (Mart.) Coville (barbatimão) is used for medicine (possibly effective against leishmaniasis); other uses in agroforestry are for timber (construction), nitrogen fixing and reforestation (e.g., S. microstachyum Poepp. & Endl., or vanillio); some species are toxic to livestock.

∎ MIMOSA CLADE (3/645) ▸ outsider Adenopodia (7, tropical Africa, Mexico, Central America).

252. Mimosa L. [4th BR] Prostrate, low spreading, or trailing, herbaceous perennials to 4 m long or erect shrubs, usually armed, sometimes with xylopodium, basal burls, roots crown, or woody rhizomes; leaves twice even-pinnately compound, the ultimate leaflets folding together at night, in rain, or on being touched, sensitive to touch or not so; flowers sweet-scented, in our species in pink to purple (rarely white), globose heads. Used as ornamentals, living fences, soil binders, fodder, green manure, shade plants, fuelwood and medicine (e.g., M. pudica L., the sensitive plant, sleeping grass, humble plant); many species are weedy, causing problems in agricultural land.

600 spp., Mexico (72 endemics, 15 extending to U.S.A. and 7 to C America (3 and 2 endemic to each region respectively); Caribbean (8 endemic); 7 spp. disjunct between Mexico–C America–Caribbean and S America; 20 spp. widely distributed in New World, 3 of which are pantropical weeds; 481 spp. in S America (inc. the widely distributeds) in Brazil (379, 286 endemics) and adjacent Paraguay, Argentina and Uruguay; Ecuador, Peru and Bolivia; Orinoco basin; 35 spp. endemic in the Palaeotropics (33 in Madagascar (31 endemic); 2 in SE tropical Africa and 3 spp. endemic to the Indian subcontinent). Seasonally dry tropical and subtropical forest, woodland, wooded grassland, thorn forest, tropical montane woodland, temperate grassland and desert. 26 spp., from several states, are rare plants in Brazil, by Plantas Raras do Brasil’s book. 5 sections and 41 series.

§ sect. Batocaulon ▸ 174 spp., tropical America.

§ sect. Calothamnos ▸ 26 spp., S & SE Brazil, Uruguay, NE Argentina and Paraguay, one extending into Bolivia.

§ sect. Habbasia ▸ 78, all (but two) native in and all (but nine) endemics to South America, ranging from warm temperate Argentina to the Gulf States of U.S.A. and the Caribbean; one (M. pellita Humb. & Bonpl. ex Willd.) circumtropical.

§ sect. Mimadenia ▸ 15 spp. from S. America, the two extending only feebly into Central America, and one endemic to W Central America to S Mexico.

§ sect. Mimosa ▸ c. 310 spp., over range of genus.

253. Piptadenia Benth. (inc. Adenopodia) Trees up to 30 m tall, shrubs (erect and scandent) and lianas. 33 spp., S America with centres in Amazonia linking to the Atlantic forests of Brazil (c. 10), NE & E Brazil drylands with extension to Venezuela (c. 4, one extending to C America and Mexico); sub-Andean drylands in Peru, Ecuador and Bolivia (c. 2); SE Brazil (c. 4); 2 spp. widely distributed in drylands from Mexico and C America to Argentina, in tropical lowland rain forest, coastal and riparian forest, tropical to subtropical seasonally dry forest, secondary forest, woodland, dry seasonal scrubland of NE Brazil (caatinga) and rocky wooded grassland. 20 spp. in Brazil, 14 endemics.

Various species used for timber (surucuçu, icarapé, soroca, jurema preta, pau jacaré, angico de bezerro) for construction, posts, tool handles, carving and firewood; tannins are extracted from the bark.

∎ CALLIANDRA CLADE (3/164) ▸ all genera in South America.

254. Acaciella Britton & Rose. Mostly slender shrubs or small trees and less frequently perennial herbs, unarmed, glabrous or pubescent; stipules fugacious or persistent, glabrous or ciliate. 15 spp., U.S.A. to Argentina, east up to W Venezuela, highly centered in Pacific Mexico, but 3 occur in South America: A. angustissima (Mill.) Britton & Rose, from U.S.A. to Bolivia and Argentina; A. glauca (L.) L. Rico, from Caribbean and N Venezuela; and A. villosa (Sw.) Britton & Rose, from Mexico to Ecuador, absent in Venezuela.

255. Calliandra Benth. Trees and shrubs, sometimes with xylopodium. 153 spp., one in Somalia and Kenya, another in South Africa, and remaining endemics to the Americas, ranging from SW U.S.A. to Uruguay, warm temperate Argentina and N Chile; 30 spp. restricted to N America, 4 extending from N America into northern S America, 6 endemic to the Caribbean, a concentration of taxa (66) from NE to SE Brazil (with 31 endemic to Bahia), 26 spp. restricted to N, NW and W S America (from the Guianas to S Peru and W Bolivia), mostly seasonally dry tropical forest (many riparian), woodland, dry seasonal scrubland of NE Brazil (caatinga); savannas of C Brazil (cerrado) and rocky grasslands (campos rupestres), some tropical submontane, several adapted to desert environments. 112 spp. in South America, 77 in Brazil, 62 endemics – 28 of them, in several states, are rare plants in Brazil, by Plantas Raras do Brasil’s book. Two subgenera, Afrocalliandra for African members (2), and subg. Calliandra in New World, with six sections:

§ sect. Androcallis ▸ 76 spp., U.S.A. to Uruguay.

§ sect. Calliandra ▸ 10 spp., Caribbean, Mexico, Guatemala.

§ sect. Microcallis ▸ 6 spp., 3 in NE Brazil, one in Chile, one U.S.A./Mexico, and one in C & S South America.

§ sect. Monticola ▸ 37 spp., all in Espinhaço Range of E Brazil

§ sect. Septentrionalis ▸ 6 spp., U.S.A. to Mexico.

§ sect. Tsugoides ▸ 4 spp., Guiana Shield and sand savannas in the Guianas, Venezuela, the Colombian Amazon and the northern state of Amazonas, Brazil.

Widely cultivated as ornamentals (calliandra, powder puff); used in agroforestry (e.g., C. houstoniana (Mill.) Standl. var. calothyrsus (Meisn.) Barneby, as livestock fodder, fuelwood, green manure, pulp for paper production, erosion control, firebreaks, reforestation, bee forage, leaf meal (protein source), medicine and as shade trees.

∎ ZAPOTECA CLADE (5/46) ▸ outsiders Faidherbia (1, E Mediterranean, tropical to southern Africa), Sanjappa (1), Thailentadopsis (3, Sri Lanka, Thailand, S Vietnam), Viguieranthus (18, Madagascar, tropical Asia).

256. Zapoteca H. M. Hern. Shrubs. 23 spp., Neotropics, 11 from SW U.S.A. and N Mexico through C America and the Caribbean (6 endemics to Mexico with a concentration in Oaxaca, 3 endemic in C America and one in Hispaniola); 10 in South America, 6 restricted to S America and largely circum-Amazonian (4 endemics in Colombia, Ecuador, Peru and Brazil one each), 3 widely distributed through N and S America (one extending to Paraguay and N Argentina); common in open sites derived from seasonally dry tropical forest, and in arid and semiarid scrubby vegetation and wet evergreen forest; Brazil has 5 spp.

∎ COJOBA CLADE (3/34) ▸ outsiders Hesperalbizia (1, Mexico), Lysiloma (11, Florida, Mexico, the Caribbean).

257. Cojoba Britton & Rose. Trees and shrubs. 22 spp., Greater Antilles, NW Andean and trans-Andean S America, SE Mexico and C America, 6 in South America; C. arborea (L.) Britton & Rose nearly coextensive with the genus, tropical, mostly wet lowland or submontane forest; three Caribbean species in seasonally dry scrub forest or in sclerophyllous shrubland (chaparral). Cojoba arborea (ardilla, jolote beard) is used as timber, ornamentals and shade trees; in Brazil occur only C. chazutensis (Standl.) L. Rico, in Acre state.

∎ PITHECELLOBIUM CLADE (5/c. 27) ▸ outsiders Ebenopsis (2, Texas, Mexico), Havardia (6, Mexico to Central America), Painteria (?).

258. Pithecellobium Mart. Trees and shrubs up to 40 m tall. 16 spp. restricted to Mexico, C America and the Greater Antilles (3 extending N into subtropical Mexico, the Bahamas and Florida, one circum-Caribbean), 10 in South America, two in Brazil, one endemic; in lowland seasonally dry tropical woodland and thorn scrub below 500 m, some to 1550 m, a few on coastal dunes or in riparian woodland.

P. dulce (Roxb.) Benth. (Madras thorn, Manila tamarind, guayamochil), is a common ornamental, hedging and shade tree, where it is also grown for the edible pulp in the seed aril (also made into beverages); other uses include timber (construction and posts), livestock fodder, firewood, bee forage (honey), seed oils (soap) and bark for tanning; plants can be tenacious weeds.

259. Sphinga Barneby & J.W.Grimes. Trees and shrubs. Three spp., Cuba (1), Mexico (2, one endemic), Guatemala, and NW S America (only S. platyloba (Bertero ex DC.) Barneby & J.W. Grimes Colombia, Venezuela and extending into the Caribbean island of Aruba), seasonally dry tropical to arid lowland and submontane forest, thicket, wooded grassland and thorn scrub.

∎ SAMANEA CLADE (2/14) ▸ both genera in South America.

260. Chloroleucon (Benth.) Britton & Rose. Trees and shrubs, characterised by axillary spines (modified sterile peduncles), striate resting buds and flowering preceding leafing. 11 spp., America, from NW Mexico (1) to the Antilles (2), remaining species in S America, Venezuela and French Guiana to S to Argentina and SE Brazil (7, 3 endemics, one of tem, from Bahia state, is a rare plant in Brazil, by Plantas Raras do Brasil’s book), in warm temperate and tropical lowland and less often submontane seasonally dry forest, xeromorphic brush-woodland, coastal thicket, wooded grassland, shrubland and desert.

261. Samanea (Benth.) Merr. Trees up to 35 m tall. Three spp., mostly circum-Amazonian S America (Venezuela, Colombia, Peru, NE Bolivia, Paraguay, S, E and N Brazil) to C America (El Salvador); seasonally dry tropical deciduous to moist evergreen forest, woodland and wooded grassland; two spp. in Brazil, none endemics. S. saman (Jacq.) Merr., from Belize to Venezuela and Ecuador, is a myrmecophyte.

S. saman (Jacq.) Merr. (rain tree, saman, monkeypod, coco or cow tamarind) is widely planted in the Old and New World tropics for shade, as an ornamental and for its nutritious pods (for livestock fodder and the sweet pulp is a human food, also made into a beverage); also used for medicine, timber (furniture, general construction, interior trim, boxes and crates, panelling, plywood and veneer), hats (made from wood shavings) and bee forage (honey).

∎ JUPUNBA CLADE (6-7/88) ▸ outsiders Albizia obliquifoliata + Albizia rhombifolia clade (Africa, posibly in Hydrochorea).

262. Pseudalbizzia Britton & Rose. Trees, shrubs, and lianas. 17 spp. (and 5 varieties) ranging in distribution from northwestern Mexico to northern Argentina and including the Caribbean. Tropical, mostly lowland (sometimes inundated) or low-montane seasonally dry riparian forest, woodland, wooded grassland, bushland and thicket; some restricted to rain forest; others extratropical, rarely in desert foothills, often in secondary vegetation. Tropical, mostly lowland (sometimes inundated) or low-montane seasonally dry riparian forest, woodland, wooded grassland, bushland and thicket; some restricted to rain forest; others extratropical, rarely in desert foothills, often in secondary vegetation.

Used for timber (construction, furniture, cabinet work, veneers, general carpentry), livestock fodder, human food (fruit pulp and seeds), bark (fibre and pulp for paper), medicine, firewood, gums, tannins, dyes, ink, soaps, fish poisons, ornamentals, street and plantation shade trees and for reforestation.

263. Punjuba Britton & Rose. (off Abarema) Unarmed trees; stipules usually deciduous; branches with indumentum present or glabrous; leaves bipinnate, alternate, one or two (three) pairs of pinnae; nectaries sessile between pairs of pinnae and leaflets. 6 spp., 5 from Panamá to Ecuador, and one endemic to Bolivia.

264. Hydrochorea Barneby & J. W. Grimes (inc. Balizia). Shrubs and trees, unarmed; branches grey to brown pilosulous to glabrescent, cylindrical; stipules persistent or caducous. 10 spp., 7 from S America in the Orinoco and Amazon basins, the Guianas and in Brazil (all of them, one endemcis) extending into Maranhão and Mato Grosso do Sul, one sp. from Mexico to Honduras, and two in W Africa from Seneal to DR Congo, in tropical, often inundated, riparian forest and woodland.

265. Jupunba Britton & Rose. (off Abarema) Unarmed trees; stipules usually deciduous; branches with indumentum or glabrous; leaves alternate, bipinnate, with one to 12 (–16) pairs of pinnae; nectaries sessile or elevated, between the pairs of pinnae and leaflets; leaflets sessile or pedicellate, symmetric or asymmetric, obovate, oval, rhombic, oblong or linear, chartaceous, coriaceous or membranaceous. 37 spp., tropical America, 29 in South America, 21 in Brazil, 10 endemics.

∎ INGA CLADE (6-9/c. 370) ▸ outsiders Albizia dinklagei + Albizia altissima (Africa).

266. Abarema Pittier. Trees. (exc. Punjuba, Jupunba) Two spp., A. cochliacarpos (Gomes) Barneby & J.W. Grimes and A. diamantina E. Guerra, Iganci & M.P. Morim, endemics to E Brazil.

267. Blanchetiodendron Barneby & J.W.Grimes. Small tree with whitish flowers. Only one sp., B. blanchetii (Benth.) Barneby & J.W.Grimes from NE Minas Gerais and SE Bahia states, in seasonally dry tropical forest, savannas of C Brazil (cerrado), liana forest and wooded grassland, 400 – 1,000 m.

268. Inga Mill. (inc. Zygia p.p.) Trees. 291 spp., Neotropics from in Mexico to Uruguay, including two spp. restricted to the Caribbean, highest diversity in Peru, Ecuador, Colombia and C America. 40 spp. confined to Mexico and southern C America, c. 80 spp. essentially Amazonian, 39 spp. restricted to coastal Brazil, c. 60 spp. extra-Amazonian within an arc from the Guianas through Colombia to Peru and Bolivia, the remaining spp. (c. 40–45) extending across the Panamá isthmus (several of these widely distributed especially in S America), mostly of lowland and montane rain forest throughout the humid tropics, 0-3,000 m, half in rain forest on non-flooded land, half in riparian habitats on periodically flooded land, rarely in seasonally dry areas. 138 spp. in Brazil, 47 endemics, one of them, from Bahia state, is a rare plant in Brazil, by Plantas Raras do Brasil’s book.

Used as multi-purpose soil restoration and agroforestry trees (e.g., I. edulis Mart., or ice cream bean, guava machete, ingá cipó), for edible fruits (the white pulp is eaten or used in flavouring desserts), shade for crops, leaf mulch, nitrogen fixing properties, timber, fuelwood and medicine.

269. Leucochloron Barneby & J.W.Grimes (exc. Boliviadendron). Trees, occasionally shrubby. 4 of E Brazil, from Bahia to Paraná states), in seasonally dry tropical, mainly wooded grassland, woodland and dry seasonal scrubland of NE Brazil (caatinga).

270. Macrosamanea Britton & Rose. Trees and shrubs. 11 spp., S America, most diverse and numerous in the Amazon basin extending N into the Orinoco valley and the Guianas, in tropical, mostly riparian and seasonally flooded forest, two spp. of seasonally wet sandy wooded grassland (savanna); 9 spp. in Brazil, 3 endemics.

271. Zygia P. Browne. (exc. Inga p.p.) Trees and shrubs, often cauliflorous. 67 spp., Neotropics, from Mexico and the Greater Antilles to S America (55), most diverse in C America and S Mexico (c. 10), Amazonia (c. 10), N and NW S America (c. 20 spp., most diverse in Colombia and the Guianas), in t ropical, mostly riparian forest and coastal habitats, generally below 900 m, but a few species reach 2,800 m. Used for soil stabilisation and erosion control (e.g., Z. longifolia (Willd.) Britton & Rose, or sotocaballo); 19 spp. in Brazil, 3 endemics.

∎ ALBIZIA CLADE (3/+12) ▸ outsiders Albizia s.s. (Asian, African, Madagascan and Pacific species; incl. Cathormion).

272. Boliviadendron E.R. Souza & C.E. Hughes. (off Leucochloron). Trees. Only one sp., B. bolivianum (C.E. Hughes & Atahuachi) E.R. Souza & C.E. Hughes, endemic to Bolivia recorded from just a small number of localities on the eastern flanks of the Andes at mid-elevations in interior Andean valleys in the Departments of La Paz, Cochabamba and Santa Cruz.

273. Enterolobium Mart. Trees up to 50 m tall. 11 spp., 4 spp. Amazonian Brazil to neighbouring countries, one of which more widely distributed to C America and Mexico; 5 spp. in Atlantic forest to seasonally dry vegetation in E Brazil, Paraguay, N Argentina, Uruguay and Bolivia; one sp. in dryland Venezuela to Colombia and one sp. widely distributed in C America, Caribbean and Mexico extending to Venezuela and Colombia, in tropical lowland and submontane rain forest (terra firme), often along rivers, and seasonally dry low forest, savannas of C Brazil (cerrado), wooded grassland, dry seasonal scrubland of NE Brazil (caatinga), Atlantic sandy coastal shrublands (restingas) and inundated grassland (pantanal). 9 spp. in Brazil, two endemics.

The timber of several species (guanacaste, batibatra, tamboril, corotu, ear pod tree) is valued for high quality furniture, cabinet work, joinery, panelling, veneers and water-resistant construction (canal sides and troughs for example); also used as shade trees, ornamentals and for pasture improvement, livestock fodder (although the fruits of some species are toxic to cattle), fibre (for paper), cork, soap substitutes, gums and medicine.

31. ROSALES

FAMILIES ABSENTS IN SOUTH AMERICA: BARBEYACEAE (1/1), DIRACHMACEAE (1/2) AND ELAEAGNACEAE (3/102).

LINEAGE 1 of 3: ROSACEAE

ROSACEAE

§ FULLY CHLOROTROPHIC

Genera/species 108/3,400-3,500 (more or less amphimictic species) Distribution Cosmopolitan except Antarctica, with their largest diversity in subtropical and temperate regions in the Northern Hemisphere. Habit usually bisexual (sometimes monoecious, dioecious or polygamous), evergreen or deciduous trees, shrubs or suffrutices, or usually perennial (rarely annual) herbs. Some species are xerophytes.

SYSTEMATIC three higher lineages with 14 smaller lineages, subfamily Dryadoideae (4/21) does not occur in South America. 278 spp. in South America.

1. SUBFAMILY ROSOIDEAE (28–31/2.080 - 2.840) - two clades, Filipendula clade (1/12; Europe, temperate Asia, NE North America) does not occur in South America; among supertribe Rosodae, with five tribes, only tribe Roseae (1/100–150, temperate regions on the Northern Hemisphere, tropical mountains in Ethiopia, Philippines and Mexico) does not occur in South America.

1.1 ROSOIDAE ▸ TRIBE RUBEAE (1/250(–700)) ‣ a single genus.

1. Rubus L. 230 species worldwide, 133 in New World, 55 in South America. 21 spp. and Colombia, mainly 2,000 – 3,000 m high in tropical Andes; 7 spp. in Brazil, none endemics.

Rubus has been divided into 12 subgenera of which only few species have been domesticated; the subg. Orobatus is exclusive to South America; representatives of the subgenera Rubus and Idaeobatus also are found in the South America; R. rosifolius Sm. as weed in world. R. glaucus Benth. fruits are produced from Mexico to Ecuador; they are consumed fresh and processed for products such as jellies and beverages.

1.2 ROSOIDAE ▸ TRIBE SANGUISORBEAE (12/300–310) - two subtribes, both in South America.

► SUBTRIBE AGROMONIINAE ‣ outsdier Leucosidea (1; South Africa, Lesotho, Zimbabwe), Aremonia (1; S and SE Europe), Spenceria (1; W China), Hagenia (1; Central African mountains, Sudan, Ethiopia to Zimbabwe)

2. Agrimonia L. Perennial rhizomatous herbs; leaves interruptedly imapripinnate. 18 spp., widely distributed in Europe and Asia southwards to Sri Lanka and Java, 8 spp. in North America and Mexico, A. parviflora Aiton in Haiti, Brazil and North America, A. villosa Cham. & Schltdl in Brazil, grows in 900-1,100 m in mountains forests of São Paulo to Rio Grande do Sul states, also in NE Argentina, and A. hirsuta Bong. ex C.A. Mey. endemic to S Brazil.

► SUBTRIBE SANGUISORBIINAE ‣ outsiders Cliffortia (c 120; S Africa to Angola and Kenya, with their highest diversity in the Cape Provinces), Poterium (13; Europe, Madeira, Canary Islands, Mediterranean, North Africa, temperate Asia), Poteridium (1; W U.S.A., NW Mexico), Sanguisorba (c 15; temperate regions on the Northern Hemisphere).

3. Acaena Mutis. (inc. Margyricarpus, Polylepis, Tetraglochin) Perennial herbs either tufted or stoloniferous, sometimes cushions, slightly woody at the base, shrubs, erect or creeping, sometimes spiny, possibly evergreen, or trees up to 25 m tall; leaves are imparipinnate, the leaflets mostly lobed or toothed, the flowers are small, more or less sessile, clustered in heads or interrupted spikes, petals are absent and the solitary carpels produce an achene enclosed in a spiny hypanthium. 108 spp., A. pinnatifida Ruiz & Pav. in California and Cono Sur, South Africa and Hawaii one endemic each, 18 in Australia and N. Zealand, and other 87 confined to South America and adjacent Antarctic Islands, 5 of then in Brazil, three endemics (two confiend to Santa Catarina state), Acaena [M]. pinnatus (Lam.) Kuntze, widely distributed in Colombia to Argentina, S Brazil, central Chile, and SE Uruguay, and A. eupatoria Cham. & Schltdl. from S Brazil, Argentina and Uruguay.

Although some members of the genus are components of the upper montane forest, others occur in woodlands at elevations as high as 5,200 m, completely surrounded by puna vegetation and well isolated from any other type of forest; populations of Acaena [Polylepis] tarapacana Phil. (Bolivia and Cono Sur) near Nevado Sajama, Bolivia, grow at about 5,600 m in altitude, making the highest record of trees in the world, displacing records of Abies squamata Masters (Pinaceae) in SW China.

1.3 ROSOIDAE ▸ TRIBE POTENTILLEAE (11–13/c. 1.360) ‣ two subtribes, both in South America.

► SUBTRIBE POTENTILIINAE (1/493) ‣ a single genus.

4. Potentilla L. 493 spp., 5 in South America, Argentina, Peru and Bolivia one endemic each, P. heterosepala Fritsch occur from Mexico to Colombia and Venezuela, and P. dombeyi Nestl. in Ecuador and Peru.

► SUBTRIBE FRAGARIINAE ‣ outsiders Argentina (c 65; Eurasia to New Guinea), Comarum (1; temperate regions on the Northern Hemisphere), Farinopsis (1; Himalayas, Central Asia, Siberia, Mongolia, China), Sibbaldia (2–4; arctic and alpine regions on the Northern Hemisphere), Sibbaldianthe (2; Europe to Siberia and W China), Sibbaldiopsis (1; Pakistan, Kashmir, India, Nepal, Bhutan, Sikkim, China); Dasiphora (3; temperate regions on the Northern Hemisphere), Chamaerhodos (7; Central and E Asia, W North America), Drymocallis (3; Europe, Mediterranean, temperate Asia), Chamaecallis (1; Afghanistan, Himalayas, SW China, Burma)

5. Alchemilla L. (inc. Aphanes, Lachemilla). Perennial herbs and shrubs. 250 (- 1,000) spp., mainly Holarctic distribution with a centre of species richness in western Eurasia but occurs also in S India, Sri Lanka, Java, China and Japan and on the mountains of Africa and Madagascar, and c. 85 in Neotropics, distributed in South and Central America from Mexico and the Greater Antilles (Hispaniola) to the Andes of N Chile and Argentina. 68 spp. occur in South America, highly centered in northern Andes, one of the most important and most species rich groups of plants in the andean páramos, between 2,200 and 5,000 m in elevation, where they can form dense stands; only one sp. occur in E South America, A. parodii (I.M.Johnst.) Rothm., from S Brazil to S Argentina.

6. Fragaria L. Perennial rhizomatous herbs, often stoloniferous. 22 spp., almost all of Eurasia, highly centered in E Asia (including narrow endemic spp. in India and Hymalaias, China, Japan), two restricteds for North America, F. vesca L. widely in North America, and F. chiloense (L.) Mill., in W North America, Bolivia, S Chile, SW Argentina and Hawaii.

1.4 ROSOIDAE ▸ TRIBE COLURIEAE (2–3/c 60) ‣ outsider Fallugia (1; SW U.S.A., NW Mexico).

7. Geum L. Perennial small rosettes, leaves imparipinnate. 30-50 spp., widely distributed in Europe, Asia, and North and South America: 9 spp., 6 only in Cono Sur, G. boliviense Focke and G. involucrata Juss. ex Pers. up to Bolivia and Brazil, and G. peruvianum Focke from Colombia to Peru; one sp. in South Africa, Tasmania, New Zealand and adjacent islands in subantartic region; in open vegetation and forests, some weedy.

2. SUBFAMILY SPIRAEOIDEAE (45/1.040 - 1.330) ‣ 7 tribes, Lyonothamneae (1/1, California), Neillieae (2/25-32, Asia and North America), Sorbarieae (4/13, Asia, U.S.A., Mexico) and Kerriodae (7/10, Asia, U.S.A., Mexico) do not occur in South America.

2.1 SPIRAEOIDEAE ▸ TRIBE AMYGDALEAE (1/230 - 490) - a single genus.

8. Prunus L. Trees or shrubs, rarely with thorns, evergreen or leaf-shedding. c. 200 spp., cosmopolitan genus, relatively few species in Neotropics and tropical Africa; 99 in New World, 48 in South America, 7 in Brazil, 5 of then from Brazil and Cono Sur, P. myrtifolia (L.) Urb. widely distributed in New World, and P. ulei Koehne endemic to Brazil.

2.2 SPIRAEOIDEAE ▸ TRIBE SPIRAEEAE (8/90-110) ‣ outsiders Aruncus (1; temperate regions on the Northern Hemisphere), Kelseya (1; W U.S.A.), Eriogynia (1; NW North America), Petrophytum (3; W North America), Sibiraea (4; the Balkan Peninsula, Siberia, Central and E Asia), Spiraea (80–100; temperate regions on the Northern Hemisphere south to Himalayas and Mexico), Xerospiraea (1; Mexico).

9. Holodiscus (K. Koch) Maxin. Shrubs, unarmed, leaf sheding, up to 20 m tall. Only one sp., H. argenteus (L.f.) Maxim., from NW Mexico to Colombia; montane habitats, in forests and rocks.

2.3 SPIRAEOIDEAE ▸ TRIBE PYRODAE (27/700-730) - two subtribes, Gillenieae (1/2, U.S.A.) does not occur in South America; among the unique lineage in South American, Pyreae, are three lineages, two in South America, and Vauquelinia Clade (1/2, Mexico/North American).

► KEGENECKIA LINEAGE (2/4) ‣ outsider Lindleya (1; Mexico)

10. Kageneckia Ruiz & Pavon. Shrubs or small tres; leaves serrate. Three spp. from Peru, Bolivia, Chile and Argentina.

► PYRINAE (18/690 - 720) ‣ outsiders are Dichotomanthes (1; SW China), Eriobotrya (11–15; Himalayas, E Asia, mountains in W Malesia), Rhaphiolepis (5; E and SE Asia), Sorbus (130–140; temperate regions on the Northern Hemisphere), Amelanchier (c 20; temperate regions on the Northern Hemisphere south to Guatemala), Pyrus (15–20; Europe, Mediterranean, temperate Asia), Aronia (c 40; Himalayas to Japan and Sumatra, E North America, Central America), Malus (c 40; temperate regions on the Northern Hemisphere), Cydonia (1; the Caucasus, Kurdistan), Docynia (1; E Himalayas, S China, N Thailand), Pseudocydonia (1; China), Chaenomeles (3; China, Korean Peninsula, Japan), Cotoneaster (260; temperate regions in the Old World), Pyracantha (7; NE Spain to northern Iran; Himalayas; SW to central China inc. Taiwan), Crataegus (140–150; temperate regions on the Northern Hemisphere), Osteomeles (1–3; China (inc. Taiwan), the Ryukyu islands, the Cook Islands, Tonga, Pitcairn Island, Rapa Iti, Hawai), Chamaemeles (1; Madeira).

11. Hesperomeles Lindl. 15 spp. all restricted from Colombia to Peru and Bolivia, except one which ranges up to Central Amerioca

LINEAGE 2 of 3: RHAMNOIDS

RHAMNACEAE

§ FULLY CHLOROTROPHIC

Genera/species 63/940–990 Distribution cosmopolitan except polar areas, with their largest diversity in tropical and subtropical regions. Habit usually bisexual (rarely monoecious, dioecious or androdioecious), evergreen or deciduous trees, shrubs or lianas (sometimes with tendrils or hooks; Crumenaria decumbens is a perennial herb), often with spines, often xeromorphic. Some species have phyllocladia and reduced leaves.

They are characterized by flowers with petal-opposed stamens (obhaplostemony) and a tendency towards xeromorphism. Obhaplostemony is a relatively rare feature in angiosperms, and this has resulted in Rhamnaceae being associated with other families such as Vitaceae and Cornaceae exhibiting this arrangement. The xeromorphic adaptations exhibited by some members of the family include reduced or absent leaves, crowding of leaves, shortening of branch axes, presence of thorns or spines, and a low, shrubby habit. There are few plants of economic value in Rhamnaceae, the most notable being the jujube (Ziziphus jujuba Mill.), a fruit tree, and the ornamental shrubs Ceanothus and Colletia.The xeromorphic adaptations exhibited by some members of the family include reduced or absent leaves, crowding of leaves, shortening of branch axes, presence of thorns or spines, and a low, shrubby habit. Colletia paradoxa (Spreng) Escal from Brazil to Argentina and Chile, Uruguay and Paraguay exibs curious thorns as leaves, and know as Crucifixion thorn or Barbd Wire Bush (for obvious reason). Z. joazeiro is known as joazeiro in NE Brazil.

SYSTEMATIC ten small lineages do not occur in South America: Ventilagineae (2/20–45, tropical regions in the Old World), Maesopsideae (1/1, tropical Africa), Bathiorhamneae (1/7, Madagascar), Doerpfeldieae (1/1, Cuba), Lasiodiscus clade (1/12, tropical Africa, Madagascar), Emmenosperma clade (1/5, New Guinea, N and E Australia, New Caledonia, Fiji), Schistocarpaea clade (1/1, NE Queensland), Phyliceae (3–4/190–195, S Africa, Madagascar, Mascarene Islands, Amsterdam Island, St. Helena, Tristan da Cunha, Gouth Island), Ceanothus clade (1/c 55, North America, Mexico, with their highest diversity in California and NW Mexico) and Pomaderreae (10/210–220, Australia, Tasmania, New Zealand, with their highest diversity in Western Australia). 345 spp. in New World, 112 in South America.

UNPLACED GENERA

1. Araracuara Fern. Alonso. Shrubs or small trees, few ramificated; leaves clustered, up to terminal branches; inflorescences thyrso-paniculates. Only one spp., A. vetusta Fern. Alonso, known only rocky fields from Caqueta region, SE Colombia.

1. TRIBE RHAMNEAE (14/265–270) ‣ outsiders Rhamnus (c 70; temperate to tropical regions on the Old World), Berchemiella (2; Japan; Hubei in China), Ventia (6; North and Central America, Mexico), Rhamnella (c 10; N Pakistan to China, Korean Peninsula and Japan, New Guinea, E Queensland, Fiji and Tonga), Dallachya (1; New Guinea, islands in W Pacific), Berchemia (c 20; E Africa to E Asia, New Caledonia, W North America), Auerodendron (10; the W Indies), Reynosia (c 15; Florida, Central America, Caribbean).

2. Condalia Cav. 21 spp., mainly U.S.A. to Guatemala, six in South America, one endemic to Colombia, another in Peru and Ecuador, remaining four in C Bolivia to C Argentina and SE Brazil (only C. buxifolia Reissek, non endemic).

3. Frangula Mill. (off Rhamnus) Herbs to shrubs or small trees, 5-merous flowers. 40 spp., widely distributed, Eurasia and Magreb; 20 in South to North America, 12 in South America, mainly in Colombia and Venezuela, some up to Cono Sur; two in Brazil, no endemics.

4. Karwinskia Zucc. 15 spp., from Texas to Panamá and Caribbean, with K. colombiana Dugand & M.C. Johnst. is endemic to Colombia and K. humboldtiana (Schult.) Zucc. Ranges from U.S.A. to Panamá and Venezuela.

5. Krugiodendron Urb. Evergreen shrubs or small trees; leaves opposite, flowers in axillary, sessile or shortly pendunculate. Only one spp., K. ferreum (Vall.) Urb., Caribbean, S Florida, Mexico to Colombia.

6. Rhamnidium Reissek. 11 spp., highly disjunct, 6 only in Caribbean, four in Cono Sur, R. elaeocarpum Reissek up to Ecuador and R. glabrum Reissek up to Brazil, and two endemics to Brazil.

7. Sageretia Broingn. 35 spp., mostly Asia (China), three in North America, S. elegans (Kunth) Brongn. in Mexico to Bolivia, and S. lehmannii (Hieron.) Radlk. confined to Cono Sur inc. Paraguay.

8. Scutia Comm. ex Brongn. 5 spp., one in tropics Old World, and 4 in tropical South America, two from Colombia to Ecuador, S. arenicola (Casar.) Reissek endemic to Brazil, and S. buxifolia Reissek from Brazil, Bolivia and Cono Sur.

2. TRIBE AMPELOZIZIPHEAE (1/1) ‣ a single genus.

9. Ampelozizyphus Ducke. Vining shrubs or high-climbing lianas. Three spp., A. amazonicus Ducke, from Amazon rainforest of Brazil, Colombia (also in Chocó), Ecuador, Perú, the Orinoco river basin in Venezuela (Bolívar, and southern Apure states), and Guianas; A. guaquirensis Meier & P. E. Berry, a tree from Venezuela’s Coastal Cordillera; and A. kuripacorum Aymard & Castro-Lima, known from Guianía department, Colombia.

A. amazonicus is well known as an Amazonian medicinal plant used to treat infections, diseases (i.e., malaria, stomach pain, liver disorders, gastritis, inflammation of the prostate, and rheumatism), snake bites, and also as a fortifying tonic and even as an aphrodisiac.

3. TRIBE GOUANIEAE (6/c 63) ‣ outsider Helinus (5; tropical and S Africa, Madagascar, NW India).

10. Alvimiantha Gray-Wilson. Evergreen climbing shrubs; leaves triplinerved. Only one species, A. tricamerata Gray-Wilson, endemic to dry regions of NE Brazil.

11. Crumenaria Mart. Annual or perennial herbs, leafy or almost leafless, sometimes with xylopodium; leaves alternate, with stipules; inflorescence a 1-14 flowered umbel, terminal or axillar; flowers pedicellated, with campanulate or turbinate-campanulate floral tube. 4 spp. from Brazil, one endemic, the remaining up to Paraguay to Argentina, C. decumbens Mart. also in Honduras and Guatemala.

12. Gouania Jack. Climbing shrubs or lianes; branches provided with coiled tendrils, pubescent, glabrescent later; leaves alternate, petiolate; inflorescence a paniculate thyrse; flowers usually bisexual; sepals 5; petals 5, cucullate; inserted below margin of disc; fruit a schizocarp, longitudinally 3-winged (or 3-angled), septicidal, separating into 3 woody or coriaceous, indehiscent cocci; seeds compressed or planoconvex. 50 spp., pantropical, and need of revision; 39 spp. in New World, 28 in South America, 15 in Brazil, 5 endemics.

13. Jonstonalia Tortosa. Climbers with tendrils in the flowering branches; leaves sparse, elliptic, entire, 3-nerved, petiolate; flowers in axillary, sessile, 2-flowered inflorescences; fruit schizocarp, giving rise to three. Only one sp., J. axilliflora (M.C. Johnst.) Tortosa, endemic to Cajamarca region in Peru.

14. Reissekia Endl. Evergreen shrubs, flowers in terminal or in axillary cymes; hypanthium shallow. Only one sp., R. smilacina (Smith) Endl., dry areas in NE Brazil.

4. TRIBE PALIUREAE (5/110–160) ‣ outsiders Hovenia (3; Himalayas, N Burma, China, Korean Peninsula, Japan), Paliurus (5; Mediterranean to Japan), Ziziphus (70-120; tropical and subtropical regions on Old World), Pseidoziziphus (2; North America and Mexico).

15. Sarcomphalus L. (off Zizyphus) Shrubs or small to medium-sized trees, deciduous, 1–10(–15) m tall, spinose; stems and primary branches with brownish or greyish bark, branchlets glabrous to pubescent, often with short, leaf-baring shoots; plants armed by usually paired, non-leafy spines. 37 spp., California to N South America, E Brazil (8, 4 endemics), and adjacent Andes in Bolivia and Argentina; 14 in South America.

5. COLUBRINA CLADE (4/c 52) ‣ outsiders Jaffrea (2; New Caledonia), Alphitonia (c 15; Malesia to New Guinea, N and E Australia, islands in W Pacific, Hawaiian), Granitites (1; SW Australia)

16. Colubrina Rich ex Brongn. 31 spp., Asia and Africa, U.S.A. to Argentina, Caribbean; 27 in New World, 8 in South America, three in Brazil, one endemic.

6. TRIBE COLLETIEAE (6/29) ‣ outsider Adolphia (2; SW Unites States, NW Mexico)

17. Colletia Comm. ex Juss. 5 spp., Ecuador, Peru, Bolivia, Chile (also Juan Fernandes), Argentina, Uruguay and Brazil (2, one endemic).

18. Discaria Hook. 5 spp., Australia and New Zealand (2), three in Cono Sur, with D. americana Gillies & Hook. up to S Brazil.

19. Kentrothamnus Suess. & Overkott. Shrubs 1-4 m. tan; leaves decussate (sometimes slightly offset, not perfectly opposite); flowers solitary or in few-flowered fascicles, bisexual, 5-merous, fruit a 3-parted schizocarpous pedestalled capsule ca. 5 mm. long, the 3 parts at maturity separating from each other and from the pedestal and each splitting in a ventral-apical midline to release the seed. Only one sp., K. weddellianus (Miers) M. C. Johnst), growing in very arid scrub on rocky mountain-slopes, and that it grows taller near stream banks than in the more xeric places (1-2 m.), in S Bolivia and NW Argentina.

20. Ochetophila Poepp. ex Endl. Prostrate to erect, leafy, often spiny shrubs or trees up to 8 m tall, with actinorhizal root nodules. Two spp., from Argentina and Chile.

21. Retanilla (DC.) Brongn. 5 spp., 4 in Chile and Argentina, one also in Peru, and one endemic to Peru.

22. Trevoa Miers ex. Hook. Only one sp., T. quinquenervia Gillies & Hook., endemic to Chile.

LINEAGE 3 of 3: CORE MORA/URTICALEAN

ULMACEAE

§ FULLY CHLOROTROPHIC

Genera/species 7/43–50 Distribution subtropical and temperate regions in North America and Eurasia, southern Central America, parts of Malesia, with their highest diversity in temperate regions, also in tropical Africa. Habit bisexual, monoecious, andromonoecious or polygamomonoecious, evergreen or deciduous trees or shrubs. Prophylls usually basal. Use ornamental plants, timber, carpentries.

Family of about 7 genera and 40 species, distributed in temperate and tropical regions; in the South America there are two genera and about 12 species. Ulmus mexicana (Liebm.) Planch., from Mexico to Panamá, is the largest tree in Mexico and Central America, reaching into 87 m tall.

Key to genera of the South American Ulmaceae

1. Fruit a samara with 2 unequal, falcate wings ------------ Phyllostylon

1. Fruit drupaceous ------------ Ampelocera

1. Samara 1-winged ------------ Ulmus

1. Samara with 2 unequal and falcate wings ------------ Phyllostylon

Key differences from similar families

ü Plants without latex, unarmed (sometimes armed in Cannabaceae, usually with latex in Moraceae).

ü Leaves simple, alternate, distichous, sometimes with cystoliths, stipules lateral (leaves simple or compound, alternate or opposite, without cystoliths, stipules lateral in Cannabaceae; Moraceae generally with simple leaf, alternate,

ü rarely opposite, without cystoliths, usually terminal stipule).

ü Perianth imbricate at pre-anthesis (imbricate in Cannabaceae, imbricate or valvular in Moraceae).

ü Fruit samara or drupaceous (drupe or achene in Cannabaceae and Moraceae, sometimes forming compound fruit (syconium) in the latter).

SYSTEMATIC outsiders Hemiptelea (1; N China, Korean Peninsula), Holoptelea (2; W, C and SW tropical Africa; India), Planera (1; SE U.S.A.), Ulmus (25–30; temperate regions on the Northern Hemisphere and southwards to N Mexico), Zelkova (6; Crete; E Türkiye, the Caucasus, N Iran; China (inc. Taiwan), Korean Peninsula, Japan, Kuril Islands; Sicily; China).

1. Ampelocera Klotzsch. Small to large, monoecious trees; branches without spines; inflorescences axillary. 10 spp. from tropical America, 8 in South America, 4 in Brazil, A. glabra Kuhlm. endemic; bark Ampelocera are hardwood and are used in building homes, furniture or dormant; the bark of specific A. edentula Kuhlm. is astringent, toxic and ulcerative, and it is used by the population of the Pichis valley, Peru, for tattoos.

2. Phyllostylon Capanema ex Benth. Andromonoecious trees or shrubs, with sriff irregular branches without spines. Two spp., P. brasiliense Capan. ex Benth. & Hook. f. endemic to E coast of Brazil, and P. rhamnoides (J. Poiss.) Taub. Widely distributed from Central America and Caribbean to Cono Sur.

CANNABACEAE

§ FULLY CHLOROTROPHIC

Genera/species 10/130–180. Distribution tropical and subtropical regions, sometimes temperate parts of the Northern Hemisphere. Habit usually monoecious, polygamomonoecious or dioecious (rarely bisexual), usually evergreen (sometimes deciduous) trees or shrubs (some species of Celtis are lianas), perennial or annual herbs (Humulus is twining and climbing by means of specialized hairs). Use ornamental plants, fibre plants (ropes and paper from phloem of male plants of Cannabis sativa L.), beer spices (Humulus lupulus L.), seed oils, medicinal plants and narcotics (Cannabis sativa), timber, carpentries.

The Celtis species are hard to identify because their flowers are very similar and the morphological characters usually cited by different authors have great variation. The taxonomy of the genus requires extensive revision, because the available studies do not adequately account for the status of the many published names. Trema micrantha (L.) Blume is a species of wide geographical distribution and great morphological variability. Studies on its biology in two natural populations revealed that several plants do change their sex during the breeding season; these studies revealed also the occurrence of rare bisexual flowers. We accept two species for the Neotropical region but the genus, like Celtis, also needs extensive revision. Some species of Celtis provide hardwood, some are ornamental or have edible fruits. The fruits of Aphananthe monoica are also edible. Trema micrantha is an important species in the regeneration of deforested areas; its fruits are enjoyed by birds; the bark can be used to make handmade paper and its timber for manufacturing industrial paper.

Key to genera of South American Cannabaceae

1. Leaves opposite; stipules united, leaving a conspicuous interpetiolar scar ------------ Lozanella

1. Leaves alternate; stipules free or united only at their base -2

2. Trees, unarmed; tepals induplicate-valvate in bud, stigmas 2, simple ------------ Trema

2. Trees or shrubs, sometimes scandent, sometimes armed; tepals imbricate in bud, stigmas 2, simple or bifurcate ------------ Celtis

Key differences from similar families

ü Plants without latex, sometimes armed, never urticating (latex in all parenchymatic tissues in Moraceae or restricted to bark in Urticaceae; plants unarmed in Ulmaceae, Moraceae (except Maclura) and Urticaceae; sometimes urticating in Urticaceae).

ü Leaves simple, alternate, distichous, rarely opposite (leaves simple or lobed, alternate, distichous, spiral or opposite in Moraceae and Urticaceae).

ü Leaves usually 3-nervate from base (penninerved with secondary venation ending in the teeth of the margin in Ulmaceae; penninerved or palmate in Moraceae and Urticaceae).

ü Flower unisexual with apical placentation (flower unisexual or bisexual in Ulmaceae; basal placentation in Urticaceae).

ü Fruit drupaceous (samara or nut in Ulmaceae; drupe or achene in Moraceae and Urticaceae, sometimes forming a compound fruit).

SYSTEMATIC outsiders Gironniera (6; Sri Lanka, SE Asia, S China to islands in the Pacific), Cannabis (1; C Asia), Humulus (2–3; temperate regions on the C Hemisphere), Pteroceltis (1; N and C China), Chaetachme (2; tropical and S Africa, Madagascar).

1. Celtis L. Monoecious or rarely polygamoecious tres, shrubs or rarely lianas, scandent shrubs with spines. 70 spp., worldwide, 29 from Mexico and Caribbean islands to Argentina, 19 in South America, 11 in Brazil, 7 endemics.

2. Lozanella Greenm. Two spp., L. enantiophylla (Donn. Sm.) Killip & C.V. Morton from Mexico to Bolivia and L. permollis Killip & C.V. Morton from tropical Andes.

3. Trema Lour. Monoecious, dioecious, or polygamous tres or shrubs; branches without spines. 4 spp., all in New World, two widely distributed in South America, from U.S.A., Mexico, and Caribbean islands to N Argentina, both in Brazil.

MORACEAE

§ FULLY CHLOROTROPHIC

Genera/species 48/1,100-1,150 Distribution tropical and subtropical regions; some species in temperate regions. Habit monoecious or dioecious (rarely gynodioecious), usually evergreen (sometimes deciduous) trees, shrubs, lianas, suffrutices or perennial herbs (Dorstenia and other tuberous geophytes or succulents; in Fatoua annual herbs). Often with large plank buttresses. Certain species of Ficus are epiphytes, some of which, the ’stranglers’, anchor in the soil by means of adventitious roots and then ‘strangle’ the host tree by the growth of these roots causing the death of the host.

The family is distributed throughout the Neotropics, with a major center of diversity in northern South America, specifically the Amazon rainforests, where all Neotropical genera are represented. Dorstenia has a concentration of species mainly in northern Central America and the Greater Antilles and in the Atlantic forests of SE Brazil. All genera are endemic and restricted to the neotropics with the exception of Morus, Trophis, Dorstenia and Ficus which have a pantropical distribution pattern.

Use Ornamental plants, fruits (Ficus carica L., Artocarpus altilis (Parkinson) Fosberg, Artocarpus heterophyllus Lam., Morus nigra L.), textile (tapa) and paper (Antiaris, Broussonetia papyrifera (L.) L'Hér. ex Vent.), nutrient substrate of silk moth larvae (Bombyx mori on Morus alba L.), narcotics (Brosimum acutifolium Huber), medicinal plants, arrow poison, rubber (Castilla elastica Sessé, Ficus elastica Roxb. ex Hornem.), timber. Castilla elastica is known as caucho in South America, have util latex. The genus Artocarpus J.R. & G. Foster and Morus alba L. are naturalized in the Neotropics, grown for their edible fruits.

The jackfruit (Artocarpus heterophyllus) produces massive seed-bearing fruits on its branches. Native to the Indo-Malaysian region, this tree is grown throughout the tropics for its pulpy, edible fruit. It belongs to the same genus as the famous breadfruit (A. altilis). According to Charles Heiser (Seed to Civilization, 1973), jackfruits may reach nearly three feet (0.9 m) and weigh up to 75 pounds (34 kg), thus making them the largest tree-bearing fruits on earth.

The inflorescences of the family are among the most complex in Angiosperms; these are often associated with pollination systems in which plants provide breeding sites as a reward for the pollinator. For the genus Ficus entomophily is well documented where the pollination is carried out by wasps of the family Agaonidae (Chalcidoidae, Hymenoptera). The two neotropical sections Americana and Pharmacosycea are pollinated by the wasp genera Pegoscapus and Tetrapus respectively. The Neotropical Castilla elastica Sessé, the New Guinea Antiaropsis decipiens K. Shum. and probably all species of tribe Castilleae are pollinated by thrips (Thysanoptera). Although the family always have been cited, even in the recent scientific literature, as an example of a wind-pollinated group, probably only 6% of all species of the family are really anemophilous.

SYSTEMATIC only one lineage absent in South America: Parartocarpus + Hullettia (2/6; S Burma, Peninsular Thailand, the Malay Peninsula, Sumatra). 371 spp. in New World, 324 in South America, a half in Ficus.

1. TRIBE ARTOCARPEAE (4/c 65) ‣ outsider Artocarpus (c 60; tropical Asia east to islands in W Pacific).

1. Acanthinophyllum (off Clarisia) Allemão. Trees. Only one sp., A. ilicifolium (Spreng.) W.C.Burger, widely in northern South America inc. Brazil.

2. Batocarpus H.Karst. Canopy tree, stems and the shallow roots covered with elevated red lenticels; inflorescences axillary; pistillate inflorescences multiflorous and globose-capitate; fruiting perianth green. Three spp. from Costa Rica to Amazonian rainforest of Bolivia, all widely distributed, all in Brazil.

3. Clarisia Ruiz & Pav. Ruiz & Pav. (exc. Acanthinophyllum) Trees or shrubs up to 40 m tall, dioecious; fruit adnate to is fleshy perianth; inflorescences often cauliflorous, or if axillary then the bark at the base of the trunk reddish; pistillate inflorescences uniflorous or multiflorous and discoid-capitate; fruiting perianth red, orange, pale yellow, or greenish yellow). Two spp. from S Mexico through Central America into Venezuela, Colombia, Ecuador, Peru, Bolivia, Guianas, the Brazilian Amazon rainforest, and E Brazil; all widely distributed, all in Brazil.

2. TRIBE MOREAE (c 6/65–70) ‣ outsiders Milicia (2; tropical Africa), Streblus (14; tropical Africa, Madagascar, SE Asia, E Queensland, E New South Wales, Solomon Islands, Norfolk Island, New Zealand, Micronesia, Polynesia).

4. Bagassa Aubl. Trees, opposite leaves (unique at Moraceae). Only one sp., B. guianensis Aubl., knows as tatajuba, in Guianas and N Brazil, in dense canopy rainforest.

5. Morus L. Trees, leaves alternate distichous, the margin crenate to serrate. 13 spp., 8 in Asia, M. mesozygia Stapf. in S and C Africa, two only North America, M. celtidifolia Kunth. from Mexico to Central America, with dubious records in Peru, and M. insignis Bureau. in Central America to Andean South America up to Argentina, from Argentina to south Mexico.

Phylogenies based on separate data sets were not statistically incongruent, and the combined tree reveals that Morus, as currently circumscribed, is non-monophyletic; subg. Morus is resolved as a clade and consists of two well supported clades: one of Asian taxa and one of North American taxa. Sampled members of the genus Trophis (two, including the type) form a clade sister to subg. Morus. M. mesozygia from Africa, of subg. Afromorus, and M. insignis from Neotropics, subg. Gomphomorus, which have not been included to date in other phylogenetic studies of the family, are placed outside the Trophis + subg. Morus clade.

6. Sorocea A. St-Hil. Lamina is slightly to pronouncedly inequilateral; leaf margin mostly dentate to denticulate, sometimes the teeth and acumen spinulose. 23 spp., all in South America, 4 up to Mexico or central America; slightly centered in Atlantic Forest of Brazil; 16 spp. in Brazil, a half endemics.

7. Trophis P.Browne. 11-12 spp., 4-5 in Asia, 7 in tropical America, three in South America, two in Brazil, both in Acre state; all spp., except one, has perianth fruits enclosing as indeiscent fruits.

3. TRIBE MACLUREAE (1/11) ‣ a single genus.

8. Maclura Nutt. Climbers, trees or shrubs up to 37 m tall, dioecious, straight to curve spines, axillary thorns; inflorescence often with glands with yellow dye; bracts of male flowers not peltate; bracts of female flowers sometimes peltate; stigmas usually two, often unequal in length. 11 spp., mainly tropical, three in New World, M. pomifera (Raf.) C.K. Schneid. in North America; and M. brasiliensis (Mart.) Endl. and M. tinctoria (L.) D. Don ex Steud. in Neotropics, both in Brazil, no endemics.

4. TRIBE DORSTENIEAE (12/140–145) ‣ outsiders Fatoua (2; Madagascar, E and SE Asia, N and E Australia, New Caledonia), Broussonetia (8; Madagascar, tropical and subtropical Asia), Malaisia (1; Malesia), Bleekrodea (4; Madagascar, the Malay Peninsula, Borneo), Sloetia (1; W Malesia, Philippines, Sulawesi), Utsetela (2; Gabon, Congo), Trilepisium (1; tropical and S Africa, Madagascar, Mascarene Islands), Treculia (3; tropical W and C Africa, Madagascar), Bosqueiopsis (1; tropical Africa), Scyphosyce (2; tropical W and C Africa).

9. Brosimum Sw. (inc. Helianthostylis, Trymatococcus) Trees, monoecious and dioecious up to 40 m tall, inflorescences usually bisexual and globose, covered with peltate bracts. 19 spp., in Central and South America (18); 17 spp. occur in Brazil, two endemics.

10. Dorstenia L. Herbs, subshrubs or shrubs, often succulent, oftem with taproots tubers, monoecious; tepals absent in the pistillate flower, interfloral bracts simple, expanded receptacle with several minute flowers crowded and fused with each other, the peripheral ones fused with the margin of the receptacle; herbaceous habit; inflorescence a receptacle with variable shape - orbicular, elliptic, obovate, angulate, stellate or furcate. c. 105 spp., 64 in New World, 44 in South America (39 in Brazil, 36 endemics, 12 spp. of this genus in Bahia, Espirito Santo, Rio de Janeiro and Minas Gerais states are rare plants in Brazil, by Plantas Raras do Brasil’s book), more 60 spp. in Africa and only one sp. in Asia; the neotropical species falls all in the exclusivey New World section Dorstenia:

5. TRIBE FICEAE (1/750–800) ‣ a single genus.

11. Ficus L. Trees, subshrubs, climbers, stranglers, or woody epiphytes, monoecious or dioecious, one of the most diverse genera with regard to habit and life form, with both deciduous and evergreen free standing trees (up to 40 tall in Brazil), small shrubs, stranglers, root climbers and creepers, with inflorescence in urceolate receptacles, often cauliflorous, the syconium enclosing the flowers, with a narrow opening called an ostiole allowing the pollination by wasps (Agaonidae, Chalcidoidea, Hymenoptera). 750 spp., 500 in SE Asia/Australasia, 186 spp. in New World, 170 in South America (81 in Brazil, 28 endemics, 5 of then in Acre, Roraima, Mato Grosso and Pernambuco (Fernando de Noronha) states are rare plants in Brazil, by Plantas Raras do Brasil’s book) including three unvailabke in distribution (F. llanensis Dugand., F. llewelynii standl., F. mitrophora Warb.), and 100 in Africa. Six subgenera:

§ subg. Ficus - Malesian region and mainland Asia.

§ subg. Pharmacosycea - New World, Pacific to West Africa; two or more sections.

§ sect. Oreosycea - 55 spp., centered in Oceania.

§ sect. Pharmacosycea - free standing terrestrial trees except subsect. Carautae; ca. 35 spp., 6 in Atlantic Forest; chloroplast genome data confidently resolved relationships among major groups of figs and largely support current understanding based on nuclear sequence data including passively pollinated Neotropical section Pharmacosycea as sister lineage to all other Ficus; three subsections.

o subsect. Carautae - only one sp., F. crassivenosa W.C. Burger, from Costa Rica to N Brazil, common in lowland region, plants initially hemi-epiphytic with adventitious roots unique among neotropical subg. Pharmacosycea.

o subsect. Bergianae.

o subsect. Petenensis.

§ subg. Urostigma - New World, Pacific to West Africa.

§ sect. Americana – tree or shrubs, often hemi-epiphyte; inflorescence a syconium; with a waxy glandular spot at the base of the midrib of the lower surface of the leaf; ca. 130 spp.

§ subg. Sycidium - West Africa to Pacific region.

§ subg. Synoecia - New World, Pacific to West Africa.

§ subg. Sycomorus - West Africa to Pacific region.

6. TRIBE CASTILLEAE (11/60–65) ‣ outsiders Sparattosyce (1; New Caledonia), Antiaropsis (2; New Guinea), Antiaris (1; tropical regions in the Old World), Mesogyne (1; tropical Africa).

12. Castilla Cerv. Trees, monoecious and dioecious. Three spp. from Mexico to Bolivia and Brazil; two of them have been economically important as a source of rubber; only C. ulei Warb. in Brazil, a over Amazon rainforest tree.

13. Helicostylis Trécul. 8 spp., Central and South America up to Bolivia and Brazil (6, no endemics).

14. Maquira Aubl. Trees, mostly dioecious; fruit adnate to is fleshy perianth; stipules free, not fully amplexicaulis. 5 spp. from N South America, one up to Central America; 4 occur in Brazil (one endemic to Peru), no endemics.

15. Naucleopsis Miq. 26 spp., all in South America, 4 up to Mesoamerica, 17 in Brazil, 5 endemics.

16. Perebea Aubl. Tress or shrubs, monoecious and dioecious; stipules free, amplexicaulis. 10 spp., all in South America, 4 up to Mesoamerica, 8 in Brazil, no endemics.

17. Poulsenia Eggers. Trees, monoeciuous, aculeate. Only one sp., P. armata (Miq.) Standl., from Mexico to Bolivia up Venezuela, and Acre state in N Brazil.

18. Pseudolmedia Trécul. 12 spp., tropical America, 9 in South America, 5 in Brazil, one endemic.

URTICACEAE

§ FULLY CHLOROTROPHIC

Genera/species 59/c. 1,300 Distribution cosmopolitan except polar areas, with their largest diversity in tropical Asia. Habit usually monoecious, polygamomonoecious or dioecious (rarely bisexual), usually evergreen (sometimes deciduous) trees, shrubs, lianas or suffrutices, perennial or annual herbs (in Coussapoa and Poikilospermum usually epiphytes). Young stems and branches often quadrangular in cross-section. Some arborescent species have buttresses. Cecropia often live in symbiosis with ants.

The family is cosmopolitan with c. 49 genera, 2,000 spp. total; and 16 genera, c. 450 spp. in the Neotropics. Leaves with cystoliths, stipules, inflorescences bracteate, perianth reduced to a single whorl of tepals. 684 spp. in New World, 377 in South America.

A poorly studied family where little is known about generic relationships and where the monophyly of most genera remains to be tested. Understanding the floral anatomy, especially with regards to the hypanthium, also needs much work. In addition, Andean Pilea is in great need of revision where a large number of new species (c. 50-100) await description. Where Poikilospermum Zipp. ex Miq. is described to the Cecropiaceae there is strong evidence for the inclusion of Cecropiaceae within a monophyletic Urticaceae. Where Poikilospermum is excluded and ascribed to the Urticaceae, however, the relationship between the two groups is ambiguous.

Key differences from similar families:

Ulmaceae: gynoecium 2-3 carpelled in flower (vs. 1), ovule apical (vs. basal); anthers not dehiscing explosively (vs. dehiscing explosively).

Moraceae: gynoecium 2-3 carpelled in flower (rarely fruit) (vs. 1), ovule apical (vs. basal); anthers not dehiscing explosively (vs. dehiscing explosively).

Key to genera of Neotropical Urticaceae

1. Leaves opposite, occasionally strongly unequal at each node and appearing alternate - 2

2. Stems and leaves with stinging hairs; stipules forked; stigma capitate ------------ Urtica

2. Stems and leaves without stinging hairs; stipules not forked; stigma capitate or threadlike - 3

3. Herb, shrub, small tree or vine; leaves never succulent, with punctiform cystoliths; stipules narrowly ovate with a length to width ratio > 1:4; stipules free, lateral; stigma thread-like ------------ Boehmeria

3. Herbs, epiphytes or shrublets; leaves always succulent, with fusiform, 'V' or 'X' shaped cystoliths; stipules fused, intrapetiolar, deltate, cordiform, auriculate, ovate, obovate or oblong with a length to width ratio < 1:4; stigma capitate ------------ Pilea

1. Leaves alternate - 4

4. Stipule scars prominent and completely encircling the stem; trees or shrubs, rarely a vine; leaves > 50 mm, entire, 3-lobed, palmate or compound pinnatifid, or palmatifid; secondary and third order venation consistently parallel, tertiary veins spaced < 1 mm apart; where leaves weakly incised, a tree - 5

5. Leaves palmately lobed or compound pinnatifid, or palmatifid - 6

6. Flowers borne in clusters on a cymose inflorescence; seeds > 5 mm diameter ------------ Pourouma

6. Flowers borne in compact racemes borne on a cymose inflorescence; seeds < 2 mm diameter ------------ Cecropia

5. Leaves entire or lobed, where lobed, entire leaves also frequently found on the same branch - 7

7. Leaves lobed, although entire leaves also found on the same plant; flowers borne in clusters on a cymose inflorescence ------------ Pourouma

7. Leaves entire, never lobed; flowers borne in capitate globose heads borne on a cymose inflorescence ------------ Coussapoa

4. Stipule scars not encircling the stem, prominent or not; herbs, and or epiphytes, shrubs and or vines or trees; leaves 2-400 mm, entire, rarely lobed, never palmate, compound pinnatifid or palmatifid; the secondary and third order venation parallel or not, where parallel not consistently so, where leaves > 50 mm, tertiary veins spaced > 1 mm apart - 8

8. Inflorescences strongly congested; flowers obscured by numerous or prominent bracteoles - 9

9. Pistillate inflorescences or portions of inflorescence with 2-4 prominent green bracts and bracteoles -10

10. Shrubs; leaf margins dentate or serrate ------------ Hemistylus

10. Herbs; leaf margins entire - 11

11. Inflorescences enveloped by two prominent cordiform bracts, which completely obscure the flowers ------------ Rousselia

11. Inflorescences subtended by 3-4 linear bracts, which do not completely obscure the flowers ------------ Parietaria

9. Pistillate inflorescences with >10 brown, grey-brown, brown or green-brown bracts and bracteoles - 12

12. Leaf margins entire ------------ Pouzolzia

12. Leaf margins dentate, serrate or crenate for some part of its length - 13

13. Leaves not strongly unequal-sized at consecutive nodes; pistillate perianth tube absent, bracteoles conspicuous, glossy, much longer than the pistillate flowers ------------ Phenax

13. Leaves of unequal sized at consecutive nodes or not; pistillate perianth tube present, bracteoles inconspicuous, papery less than half length of pistillate flowers - 14

14. Leaves of almost equal size at consecutive nodes or, where of unequal size, not consistently so; achene easily released from the perianth in fruit, shiny ------------ Pouzolzia parasitica

14. Leaves of strongly unequal size at consecutive nodes; achene not easily released from the perianth in fruit, not shiny - 15

8. Inflorescences not strongly congested, although flowers can be borne in compact glomerules or on a few-branched spike-like panicle; flowers not obscured by prominent bracteoles even though frequently very small - 15

15. Upper leaf surface with punctiform cystoliths - 16

16. Tree > 3 m; young stems and leaves with stinging hairs ------------ Discocnide

16. Shrub < 3 m; young stems and leaves without stinging hairs ------------ Hemistylus

15. Upper leaf surface with fusiform cystoliths - 17

17. Young stems and leaves with stinging hairs; stipules forked for 1/2 or more of length ------------ Laportea

17. Young stems and leaves with or without stinging hairs; stipules forked or not forked, where forked, divided for 1/3 or less of length - 18

18. Stipules forked ------------ Urera

18. Stipules not forked - 19

19. Pistillate inflorescences pendulous, thread-like; stigma foot-shaped; achene in fruit dry, not surrounded by a fleshy tepal or panicle branch; pistillate inflorescences with flowers borne singly along peduncle; plants without stinging hairs; stems without spines ------------ Myriocarpa.

19. Pistillate inflorescences erect, not thread-like; stigma penicillate; staminate inflorescence with flowers borne in clusters of 3-10 - 20

20. Plants without stinging hairs; pistillate infructescence branches becoming fleshy in fruit giving the appearance of a spadix ------------ Gyrotaenia

20. Plants with or without stinging hairs; pistillate infructescence not becoming fleshy, tepals becoming fleshy ------------ Urera

SYSTEMATIC 3 subfamilies, all in South America.

1. SUBFAMILY CECROPIEAE (c 32/c 500) ‣ outsiders Leucosyke (c 50; Malesia to Polynesia), Musanga (2; tropical Africa), Myrianthus (4–7; tropical Africa), Gesnouinia (1; Canary Islands), Soleirolia (1; Italy, islands in W Mediterranean); Forsskaolea (6; SE Spain, Canary Islands, the Cape Verde Islands, Africa, Arabian Peninsula, India), Didymodoxa (3; S and E tropical Africa to N Ethiopia), Droguetia (7; tropical to subtropical Africa, NE India to Yunnan and Taiwan in China, and Java), Australina (2; Ethiopia, Kenya, SE Australia, New Zealand); Oreocnide (c 18; China, Japan, tropical Asia to New Guinea); Chamabainia (1; tropical Asia, Taiwan in China), Gonostegia (5; S China, SE Asia to N Australia), Neodistemon (1; tropical Asia), Pipturus (35–40; Mascarene Islands, tropical Asia to New Guinea, N Territory and Queensland, Melanesia, Polynesia incl. Hawai), Neraudia (5; Hawai); Debregeasia (3; Ethiopia, tropical and subtropical Asia), Astrothalamus (1; W and C Malesia), Archiboehmeria (1; SE Asia, S China), Sarcochlamys (1; tropical Asia), Cypholophus (15–30; Malesia to New Guinea and islands in W Pacific, Taiwan in China).

1. Boehmeria Jacq. Shrubs or small trees, sometimes woody-based herbaceous perennials, monoecious or dioecious; leaves opposite, lamina triplinerved; inflorescences axillary, usually spike-like, with flowers gathered in unisexual glomerules along inflorescence axis (rarely sessile and globular). 50 spp., pantropical, 19 in New World, 16 in South America, 5 in Brazil, no endemics.

2. Cecropia Loefl. Trees, pioneer species characteristic of disturbed forest, trunks frequently with stilt or buttress roots, hosting extensive ant colonies and bearing nodes encircled by prominent stipule scars, leaves palmately lobed or compound. 71 spp., 69 neotropical, 67 in South America, highly centered and Andean forests (37 in Colombia), 2 in Africa; 20 spp. in Brazil, 4 endemics. 47 spp. of this genus, many in South America and Brazil, are myrmecophytes.

3. Coussapoa Aubl. Trees or shrubs, either hemiepiphitic, with aerial roots, or terrestrial, with stilt-roots, leaves entire; mainly in lowland forests. 48 spp., from Central and South America (41 here), 22 spp. in Brazil, 7 endemics, C. arachnoidea Akkermans & C.C.Berg from Amapá state and C. floccosa Akkermans & C.C.Berg from Minas Gerais state are rare plants in Brazil, by Plantas Raras do Brasil’s book. C. asperifolia Trécul from norther South America is a myrmecophyte.

4. Hemistylus Benth. Small trees or shrubs, leaves alternate, long-petiolate, entire or dentate. 5 spp., 4 from S to Venezuela to Ecuador, with H. odontophylla Wedd. up to Mexico, and the fifth, H. brasiliensis Wedd. ex Warm., endemic to Brazil.

5. Parietaria L. Herbs, annual or perennial, sparsely to densely pubescent with hooked and straight, nonstinging hairs on all parts of plant, stinging hairs absent; Stems often branched from base, erect, ascending, or decumbent; Leaves alternate; stipules absent; Inflorescences axillary; flowers bisexual, staminate, or pistillate, proximal flowers usually bisexual and staminate, distal flowers pistillate. 24 spp., subcosmopolitan distribution, 7 spp. in New World, all North and Central America except P. floridana Nutt., native from U.S.A. to Argentina, S Brazil, and Cuba.

6. Phenax Wedd. Shrubs or subshrubs, not urticants. 26 spp., neotropical, 15 in South America, only two spp. in Brazil, no endemics.

7. Pouzolzia Gaudich. 70 spp., pantropical; 13 spp. in New World, 11 in South America, only three in Brazil, P. saxophila Friis, Wilmot-Dear & A.K. Monro, from southern Bahia state, endemic.

8. Pourouma Aubl. Tress, often with stilt-roots, often aromatic; leaves entire or palmate. 32 spp., neotropical, 31 in South America (except a single species endemic to Mexico), 19 of them occurs in Brazil, three endemics. P. formicarum Ducke and P. myrmecophila Ducke from northern South America are myrmecophytes.

9. Rousselia Gaudich. 4 spp., Mexico, Central America, Caribbean, R. erratica Standl. & Steyerm. up to N Colombia.

2. SUBFAMILY URTICEAE (c 11/c 210) ‣ outsiders Nanocnide (2; China (inc. Taiwan), N Vietnam, Korean Peninsula, Japan), Zhengyia (1; Hubei province in C China), Dendrocnide (37; tropical Asia to New Guinea, E Queensland, E New South Wales and islands in the Pacific), Discocnide (1; Mexico, Guatemala), Girardinia (3; E and NE Africa, tropical and subtropical regions in Africa and Asia); Obetia (19; tropical and S Africa, Madagascar, Mascarene Islands), Poikilospermum (c 20; E Himalayas, SE Asia, Malesia).

10. Laportea Gaudich. Herbs or shrubs, urticant. 23 spp., pantropical, 12 only in Africa and Madagascar, three in New World, only one in South America, the pantropical aggressive weed L. aestuans (L.) Chew.

11. Urera Gauduch. Shrubs, small trees and lianas of riparian and disturbed vegetation, occasionally with stinging hairs on the leaves and inflorescence, fruit fleshy and brightly coloured or white, urticant. 24 spp. New World, 13 in South America, 6 in Brazil, one endemic.

12. Urtica L. Herbs to shrubs. 80 spp., subcosmopolitan, centered in Northern Hemisphere, 29 in New World, 20 spp. in South America, mainly in mountains of Andes, highly centered in Peru and Chile, all in W South America except U. circularis (Hicken) Soraru from Bolivia, Argentina to S Brazil, and U. spathulata Sm., from Argentina, S Brazil, Paraguay and Uruguay.

3. SUBFAMILY ELATOSTEMATEAE (7–8/950-1,000) ‣ outsiders Elatostema (c 300; tropical Old World, 1 in New Zealand), Procris (c 15; tropical regions in the Old World); Gyrotaenia (5; Caribbean), Meniscogyne (1; Laos, Vietnam), Lecanthus (3; tropical regions in the Old World), Petelotiella (1; Indochina).

13. Myriocarpa Benth. Shrubs or trees, small trees of riparian and disturbed vegetation, inflorescences few-branched spikes, often long and pendent. 15 spp., Neotropical, centered in Mexico and Central America, 9 in South America, only two in Brazil, both widely distributed.

14. Pilea Lindl. Understory herbs, epiphytes or shrublets, opposite succulent leaves with intrapetiolar stipules (the later rarely much reduced or even absent, in some spp. very aberrant) in each axil. 716 spp., 379 in New World, 142 in South America, largest centered in Andes (eg. 67 in Colombia); only 14 in Brazil, 8 endemics; some spp. are very narrow endemic, e.g. the fully glabrous P. carautae M. D. M. Vianna & R. J. V. Alves, known only from the Cabo Frio municipality, Rio de Janeiro, SE Brazil.

P. brasiliensis Gaglioti, Romaniuc & A.K. Monro, endemic to savannah of C Brazil, belongs to the Dentatae-group of Weddell (1869) and the Fallaces group of Killip (1936); it represents the first record of the Fallaces group in Brazil; it is characterised by being glabrous, with toothed pinnately veined leaves, petioles of up to 3 cm long and paniculate inflorescences.

P. cavernicola A.K. Monro, C.J. Chen & Y.G. Wei from NW Guangxi Province, ca 500–1,000 m, caves in limestone karst, growing at any point from the back to the entrance of the cave, PAR 0.02-1.39 mmol/m2/sec (ca 0.04-2.78 % full daylight), possibly the shadiest plant worldwide.

32. FAGALES

FAMILIES ABSENTS IN SOUTH AMERICA: CASUARINACEAE (4/91) AND TICODENDRACEAE (1/1).

LINEAGE 1 of 4: NOTHOFAGACEAE

NOTHOFAGACEAE

§ FULLY CHLOROTROPHIC

Genera/species 4(own data)/36 Distribution southern South America (including Staten Island) south of 33ºS, New Zealand, Tasmania, E and SE Australia, New Caledonia, New Guinea, the D’Entrecasteaux Islands (Goodenough, Normandie), New Britain. Habit monoecious, evergreen or deciduous trees or shrubs. Horizontal lenticels often abundant. Bud scales decussate.

SYSTEMATIC outsider Trisyngyne (25; New Guinea, New Caledonia).

1. Fuscospora (R.S.Hill & J.Read) Heenan & Smissen. (off Nothofagus) Trees up to 30 m high; dichasia with 1 central dimerous flower and 2 lateral trimerous flowers, or 1 or 0 dimerous flower and 1 trimerous flower; fruits 0–1 dimerous, 1–2 trimerous, or 4–7. 6 spp., in New Zealand (4 endemics), F. alessandri (Espinosa) Heenan & Smissen endemic to center Chile, and one in Tasmania.

2. Lophozonia Turczaninow. (off Nothofagus) Trees up to 40 m high; dichasia with 1 central dimerous flower and 2 lateral trimerous flowers; fruits 1 dimerous or 2 trimerous. 7 spp., New Zealand (2 endemics), Australia (1 endemic), and 4 in Chile, two up to Argentina.

3. Nothofagus Blume. (exc. Lophozonia, Fuscospora) Trees up to 45 m high; dichasia with 1 central dimerous flower and 2 lateral trimerous flowers, or 1 trimerous flower; fruits 1 dimerous or 1–2 trimerous. 5 spp. in southern South America, all in both Argentina and Chile.

LINEAGE 2 of 4: FAGACEAE

FAGACEAE

§ FULLY CHLOROTROPHIC

Genera/species 8/983 Distribution southern Canada southwards to NW South America and Cuba, temperate parts of Europe and SW Asia, Mediterranean, Himalaya, E Asia to Russian Far East and Japan, SE Asia, Malesia, New Guinea. Habit usually monoecious (rarely dioecious), evergreen or deciduous trees or shrubs. Use ornamental plants, fruits (Castanea sativa Mill.), timber, carpentries, barrels, cork (Quercus suber L.), tanning (tannin from oak galls).

Fagaceae could be confused with some other members of the Fagales, but differ from them in possessing the folllowing characters:

Myricaceae: inferior ovary, absence of pellucid punctuations and peltate scales, nutlike fruit.

Betulaceae: fruit subtended by cupule.

Juglandaceae: simple leaves.

Ticodendraceae: nutlike fruit, absence of circular stipule scar.

Lauraceae: Quercus fruits could perhaps be confused with those of the Lauraceae, but there are many significant differences from that family (lack of typical Lauraceous odour, generally serrate leaf margins, stipules, stellate hairs etc.).

SYSTEMATIC two subfamilies, Fagoideae (1/10) absent in South America; among Quercoideae, outsiders are Castanea (8; temperate regions on the Northern Hemisphere, E Mediterranean to N Iran), Castanopsis (143; tropical and subtropical regions in Asia, with their highest diversity on Borneo), Lithocarpus (343; India, Sri Lanka, SE Asia, Malesia), Chrysolepis (2; W North America), Notholithocarpus (1; SW Oregon, California).

1. Quercus L. Trees or shrubs with alternate (often closely bunched), some exceptionally large, dominant overstory trees, perhaps an almost equal number of species are shrubs or small trees, making ECM symbioses with fungi, simple, deciduous or evergreen leaves. 461 spp. in northern temperate zone to south up in Colombia and Malaysia, particularly in drier habitats such as chaparral, in edaphically challenging environments, and in some higher elevation forests; 202 species in New World, one notes a gradual reduction of oak species diversity from Mexico into Central America: 4 in Canada, 91 in the U.S.A., one in Cuba, 137 in Mexico (nearly absent in Yucatan and coastal Pacific and Atlantic, 83 endemics), 13 in Central America, and the single Q. humboldtii Bonpl., subdivided into 2–3 species by some authors, in Colombia, 1,400 m to 3,300 m altitudinal range. Two subgenera:

§ subg. Quercus ‣ 5 sections.

§ sect. Protobalanus ‣ 5 spp. from U.S.A. to NW Mexico.

§ sect. Ponticae ‣ two spp. in mountainous areas of NETürkiye and W Georgia (Transcaucasia) and in W North America (N California, S Oregon)

§ sect. Virentes ‣ 7 spp. in SE North America, Mexico, the West Indies (Cuba), and Central America.

§ sect. Quercus ‣ c. 150 spp. in North America, Mexico, Central America, western Eurasia, East Asia, and North Africa.

§ sect. Lobatae ‣ 120 spp. in North America, Mexico, Central America, and Colombia in South America.

§ subg. Cerris ‣ three sections.

§ sect. Cyclobalanopsis ‣ ca. 90 spp. in tropical and subtropical Asia including the southern Himalayas.

§ sect. Ilex ‣ c. 35 spp. in Eurasia and North Africa.

§ sect Cerris ‣ c. 10-12 spp. in Eurasia and North Africa.

2. Trigonobalanus Forman. Three spp., T. doichangensis (A.Camus) Forman from China (SW & S Yunnan) to N Thailand, T. verticillata Forman from Hainan, Peninsula Malaysia, Sumatera, N. Borneo, Central Sulawesi, and T. excelsa Lozano, Hern. Cam. & Henao, from central Colombia; this species is a small tree known only three sites at 1650 to 2,100 m altitudinal range.

LINEAGE 3 of 4: JUGLANDIDS

JUGLANDACEAE

§ FULLY CHLOROTROPHIC

Genera/species 9/70 Distribution North America, Caribbean, Central America, Andes, SE Europe, northern Türkiye and eastwards to Himalaya, Assam, E Asia southwards to northern Vietnam and northwards to Russian Far East; Taiwan (China), SE Asia, Malesia, New Guinea. Habit usually (sometimes dioecious; in Platycarya occasionally bisexual), usually evergreen or deciduous trees (rarely shrubs). Sometimes aromatic. Buds covered by brown hairs, often scaly. Use ornamental plants, seeds and seed oils (Juglans, Carya), medicinal plants, timber, carpentries.

Although closely related to other members of the Fagales, this family is readily distinguished from all of them by its compound leaves.

Key to genera of South American Juglandaceae

1. Leaves alternate ------------ Juglans

1. Leaves opposite ‣ 2

2. Fruits with samaroid wings ------------ Oreomunnea

2. Fruits without samaroid wings ------------ Alfaroa

SYSTEMATIC two subfamilies, both in South America.

1. SUBFAMILY JUGLANDOIDEAE (5/47) ‣ outsiders Cyclocarya (1; E China), Pterocarya (6; Caucasus; E and SE Asia), Carya (18; E Asia south to N Vietnam, E North America to Central America), Platycarya (1; C and E China, Korean Peninsula, Japan, N Vietnam).

1. Juglans L. 22 spp. in 4 sections, sect. Trachycaryon monotypic endemic to E North America, sect. Juglans with J. regia L. from Europe to China and the Himalayas and J. sigillata Dode endemic to China, sect. Cardiocaryon with three spp. from China, Japan and Korea; and sect. Rhysocaryon, the black walnuts, 16 spp., endemic to the New World and includes nine North American, three Central American and four South American taxa: J. australis Griesb. (S Bolivia and NW Argentina), J. boliviana (C.DC.) Dode. (Bolivia), J. venezuelensis Mann (Venezuela) and J. neotropica Diels. (W Venezuela to Peru, in altitudes of 1,600 to 3,100 m).

2. SUBFAMILY ENGELHARDIOIDEAE (3/19) ‣ outsider Engelhardia (8; the Himalayas to China (inc. Taiwan), SE Asia to New Guinea).

2. Alfaroa Standl. 7 spp., six from Mexico to Panamá and A. williamsii D. E. Stone., from Nicaragua to N Colombia, 1,680 m to 2,300 m altitudinal range, Antioquia, Huila and Santander.

3. Oreomunnea Oerst. Three spp., two in Central America and O. munchiquensis Lozano & F.González endemic to Andes from 1,500 m to 2,500 m from Cauca Cordillera Occidental, Colombia.

MYRICACEAE

§ FULLY CHLOROTROPHIC

Genera/species 4(includes Morella)/51 Distribution.tropical and temperates regions of the world, absent in E South America. Habit monoecious, andromonoecious or dioecious, evergreen or deciduous trees or shrubs. Usually aromatic. Use spices (Myrica gale), aromatic waxes, tanninic acid, medicinal plants.

SYSTEMATIC outsiders Myrica (2, one circumboreal, another endemic to California), Comptonia (1, E North America); Canacomyrica (1, New Caledonia)

1. Morella L. (inc. Myrica p.p.) Trees or shrubs, frequently aromatic, often with woody rhizomes; flowers usually dioecious in bracteates spikes; male spikes axillary, solitary, usually densely flowered; female spikes axillary, longer or shorter than the male spikes; perianth 0; drupe small, globose or ovoid, usually warted and covered with white wax; endocarp hard. Seed erect; testa membranous; albumen 0; embryo straight with planoconvex fleshy cotyledons and a short radicle. 56 spp., distributed throughout the Northern Hemisphere, 6 in Mascarenes, 11 in tropical Africa and 9 in South Africa; 8 in South America: M. cerifera (L.) Small (Alaska to Panamá, Caribbean, and San Andreas Caribbean Is. off Colombia), M. chavalieri C. Parra-O. (puna montane vegetation in Argentina and Bolivia), M. funckii (A. Chev.) C. Parra-O. (tropical Andes), M. parvifolia (Benth.) C. Parra-O. (tropical Andes), M. pavonis (C. DC.) Parra-O. (dry coastal areas in N Chile and montanes of S Peru), M. pubescens (Humb. & Bonpl. ex Willd.) Wilbur. (tropical Andes), M. rotundata (Steyerm. & Maguire) Parra-Os. (Chimantá Massif of Guiana Shield) and M. singularis (C. Parra-O.) C. Parra-O. (tropical Andes).

LINEAGE 4 of 4: BETULIDS

BETULACEAE

§ FULLY CHLOROTROPHIC

Genera/species 6/170 Distribution temperate and polar regions in the Northern Hemisphere and southwards to northern Argentina, North Africa, Himalaya, Indochina and Sumatra. Habit monoecious, usually deciduous (rarely evergreen) trees or shrubs. Horizontal lenticels often abundant. Use ornamental plants, fruits (nuts from Corylus), timber, carpentry, charcoal, brushes and besoms, birch bark (Betula).

All members of this family have associations with nitrogen-fixing bacteria in their roots (e.g. Frankia species); the wood of Alnus acuminata Kunth is very hard and is used for many purposes including construction, furniture, matches, musical instruments, and tool handles. This species is planted as a timber tree and has also been used for agroforestry in Costa Rica (as a shade tree for coffee crops).

SYSTEMATIC subfamily Coryloideae (4/70) does not occur in South America; among Betuloideae, Betula (c 35; temperate, boreal and polar regions on the Northern Hemisphere south to SE Asia) is a outsider.

1. Alnus Mill. Trees, leaves simple, alternate (sometimes almost distichous), with pinnate venation and serrate or dentate margins, sometimes with indumentum of simple hairs. Intra-petiolar stipules present; flowers unisexual (plants monoecious), borne on catkins, subtended by bracts; petals absent; fruits are one-seeded nuts: flattened, winged and borne in cone-like structure. c. 25 spp. from temperate Northern Hemisphere to Himalaya and Andes; only one sp. in South America, A. acuminata Kunth, from Mexico to N. Argentina, with three subspecies disjunct:

§ A. acuminata subsp. acuminata - NW Venezuela to N Argentina, widely distributed through South and Central America between 1,200 and 3,200 m altitude, generally associated with damp ground and occurs along the banks of rivers and marshes.

§ A. acuminata subsp. arguta - NW Mexico to SW Panamá.

§ A. acuminata subsp. glabrata - endemic to Mexico.

33. CUCURBITALES

FAMILIES ABSENTS IN SOUTH AMERICA: CORYNOCARPACEAE (1/5), DATISCACEAE (1/2) AND TETRAMELACEAE (2/2).

LINEAGE 1 of 3: ANISOPHYLIDS

APODANTHACEAE

§ PARASITIC (Prosopanche – ... - Mitrastemon – APODANTHACEAE – SANTALALES – Lennoa – OROBANCHACEAE - Cuscuta)

Genera/species 2/12 Distribution California, Florida, Caribbean, Mexico, Central America, South America southwards to central Argentina, E Mediterranean, from SE Türkiye to northern Iran, tropical E Africa, SW Australia. Habit usually monoecious or dioecious (rarely bisexual), achlorophyllous herbaceous endophytes without rhizome or normal roots. Root or stem holoendoparasites (Apodanthes on Salicaceae [e.g. Casearia and Xylosma], Burseraceae and Meliaceae; Pilostyles on Fabaceae).

Vegetative body resembling fungal mycelium (endophytic tissues in host plant). Stem and leaves absent; flowers are the only visible part outside the host (solitary, in groups or rows); flowers unisexual and very small, a few mm in diametre, subtended by scalelike bracts; the structures around the flowers are referred to here as bracts, but they have also been called leaves (scale-like), scales or sepals (the inner whorl) by different authors; isophasic parasitism.

SYSTEMATIC both genera in South America.

1. Apodanthes Poit. Isophasic trunk holoparasitic, flowers less than 10mm; the bracts are whitish and yellow to orange, brown or red, free in the inner and outer whorls and connate in the middle ones. Only one sp., A. caseariae Poit., from Guatemala, Honduras, Costa-Rica, Panama, Colombia, Venezuela, Suriname, French Guiana, N & E Brazil, Peru and Bolivia, parasiting only plants of the families Salicaceae (Casearia and Xylosma), some Burseraceae and Meliaceae; A. minarum of Minas Gerais state is a rare species in Brazil by Plantas Raras do Brasil’s book, but it’s non-valid species.

2. Pilostyles Guill. Isophasic trunk holoparasitic, flowers less than 10mm, the smallest of all parasitic plants; dioecious (rarely monoecious), bracts are red to brown and free, cleistogamous in mexican endemics. 12 spp., 5 in Old World (Iran and Syria, Zimbabue, Zambia, Tanzania, Angola, Malawi and SW Australia) and 7 in New World, four only from U.S.A. to Mexico, one reaching into Honduras, and three in South America: P. berteroi Guillemin in Adesmia of Chile, Argentina, Peru, and Bolivia; P. blanchetii (Gardner) R.Br. in Mimosa, Bauhinia, Cassia, Dioclea, Galactia and Schnella at Jamaica, Cayman Is., Costa Rica to Venezuela, Guyana, Brazil, Argentina and Uruguay; and P. boyacensis González F, Pabón-Mora in dry interandean valleys of the Colombian E Cordillera, 2,000-2,415 m elevation range, parasiting Dalea cuatrecasasii Killip ex Barneby (Fabaceae).

ANISOPHYLLEACEAE

§ FULLY CHLOROTROPHIC

Genera/species 4/c. 35 Distribution northern South America (Amazonas), tropical Africa, southern India, Sri Lanka, W Malesia. Habit usually monoecious (sometimes polygamomonoecious or bisexual; in Combretocarpus dioecious), evergreen trees or shrubs. Most species are anisophyllous (Combretocarpus is isophyllous).

Often formerly placed in Rhizophoraceae but differs in the spirally arranged exstipulate leaves and distinct styles and in the petals not aristate.

Key to genera of Neotropical Anisophyllaceae

1. Petals laciniate or clawed; fruit a capsule ------------ Anisophyllea

1. Petals simple not clawed; fruit a samara ------------ Polygonanthus

SYSTEMATIC outsiders Poga (1; Guinea to Congo), Combretocarpus (1; W Malesia).

1. Anisophyllea R. Br & Sabine. Trees or shrubs to 38 m, monoecious; inflorescence an axillary panicle; flowers hermaphrodite or unisexual; petals 3-laciniate; fruit a berry or drupe. 67 spp., of which 30 are in Malesia, 26 in mainland Africa, 5 in Madagascar, and two in South America, A. manausense Pires & W.A.Rodrigues, known only in Central Amazon rainforest around Manaus and west to the Rio Demeni, between Manaus and Itacoatiara (Amazonas state) and Loreto region in Peru; and A. guianensis Sandw., from Guyana and Amazonas state, Brazil.

2. Polygonanthus Ducke. Small trees, leaves distichous; inflorescence racemose but appearing spicate, winged samaras. Two spp., extreme rares tress in the Brazilian Amazon rainforest, in valleys of rivers: P. amazonicus Kuhlm. occurs in municipality of Maues, and reports in Acre and Pará states, in with river’s beach; P. punctulatus Ducke is from Rio Negro, collected only one time.

LINEAGE 2 of 3: CORIARIIDS

CORIARIACEAE

§ FULLY CHLOROTROPHIC

Genera/species 1/15 Distribution Mexico, Central America, W South America (the Andes from Colombia southwards to central Chile), W Mediterranean, Greece, temperate and subtropical Himalaya, Japan, Taiwan in China, Philippines, New Guinea, New Zealand, islands in the SW Pacific (Solomon Islands, Vanuatu, Fiji, Samoa Islands, Society Islands, etc.). Habit usually dioecious (sometimes monoecious, andromonoecious, gynomonoecious or polygamomonoecious, rarely bisexual), evergreen or deciduous shrubs (sometimes suffrutices or small trees). Some branches with limitied growth similar to pinnate leaves. Buds usually perulate.

SYSTEMATIC a single genus.

1. Coriaria L. Suffruticose erect or scandent, evergreen shrubs; branching (resembling compound leaves), leaves opposite, simple, coriaceous, abaxial surface puberulent, margins entire, primary venation palmate; inflorescences terminal or from previous season's growth, racemose; flowers many, small, bracteate, bisexual, actinomorphic, pentamerous; fruit a pseudo-drupe or achene, small ca. 2-3mm long; seed 1-10, compressed. 13 spp., 12 in W Mediterranean, Himalayas to Japan, Taiwan in China, Philippines and New Guinea, Solomon Islands, Vanuatu, Fiji, New Zealand (7 endemics), Samoa, Society Islands and other islands in the South Pacific, and C. ruscifolia L., in two taxa, subsp. microphylla (Poir.) J.E.Skog, New Guinea to New Zealand, Mexico to Peru, Venezuela; subsp. ruscifolia, New Zealand to S. Pacific, C & S Chile to SW Argentina; recorded growing as part of the pioneer community following volcanic activity; nitrogen-fixing roots with characteristic nodules; check under leaves for the pendent racemose inflorescence; fruits enclosed by fleshy corolla, are reported to contain poisonous compounds; flowers are wind pollinated.

LINEAGE 3 of 3: CUCURBITIDS

BEGONIACEAE

§ FULLY CHLOROTROPHIC

Genera/species 2/2,027 Distribution tropical and subtropical regions in the Southern and Northern Hemispheres, especially northern South America and tropical Asia. Habit usually monoecious (male flowers are produced ab initio, and subsequently female flowers; rarely dioecious), usually more or less succulent perennial herbs with tuber (in Hillebrandia round) or rhizome (in Begonia rhizome, rarely tuber), sometimes climbing, sometimes somewhat woody below and frutescent. Nodes swollen.

Begoniaceae includes the monotypic genus Hillebrandia, distinguished by a semi-inferior ovary, incompletely closed ovary, fruits that dehisce between the styles, and more numerous, more highly differentiated sepals and petals. It is the only member of the family native to Hawaii. Begonia is extensively widely distributed.

Key differences from similar families

ü in Datiscaceae fruits dehisce between the styles, pistillate flower tepals are absent in Datiscaceae.

ü in Begoniaceae fruits dehisce between the wings, pistillate flower tepals (2-)3-5(6-8), never absent.

SYSTEMATIC outsider Hillebrandia (1; Hawaii).

1. Begonia L. Annual or perennial herbs, shrubs or subshrubs, frequently succulent, erect or climbing, rarely epiphytic; stems erect or creeping, with rhizomes or tubers, generally fleshy; leaves alternate, simple, entire or lobed, margins serrate, asymmetric, venation palmate or pinnate, the genus with largest diversity of variegated leaves worldwide (221); petiolate; inflorescences cymous or thyrsoid; flowers unisexual (plants monoecious or dioecious), white, pink or reddish; fruits usually loculicidal capsules, frequently chartaceous, seeds numerous, small (235-)300-600(-1450). c. 1,500 spp., largely distributed in the tropical and subtropical areas of the world, and within this area is absent only from Australia and Polynesia; 661 spp. in New World, 477 in South America, the eleventh largest genus in Brazil, with 242 spp. (221 endemics) in almost all ecosystems, except in the mangroves, with approximately 180 species in the Atlantic Forest, and about 177 restricted to this domain; 27 spp., all in SE region and some in Santa Catarina state, are rare species in Brazil by Plantas Raras do Brasil’s book. 33 sections in New World:

Due to its many forms and ranges of habits, and variation of leaves and flowers, Begonia is one of the most diverse genus worldwide.

§ Parietoplacentaria (3, Mexico to Panama), Urniformia (1, Guatemala to Panama) and Quadriperigonia (20, Mexico to Honduras) has 24 spp. do not occur in South America.

§ Barya (2, Ecuador and Peru), Australes (20, Argentina, Bolivia, Ecuador and Peru), Gobenia (16 Colombia to Peru), Hidristyles (11, Colombia, Argentina, Peru and Bolivia), Eupetalum (16, Venezuela to Argentina), Microtuberosa (1, Peru), Pilderia (6, Guyana to Peru and Trinidad e Tobago), Semibegoniella (15, Colombia and Ecuador), Warburginna (1, Bolivia) have together 88 spp., are restricted to South America but do not occur in Brazil. B. elachista Moonlight & Tebbitt from E Peru, unique member of sect. Microtuberosa, is the world smallest species of genus, that reaches maturity at fewer than 5 cm in height.

§ Astrothrix (5, Espirito Santo and Rio de Janeiro states), Gaerdtia (8, Espírito Santo, Minas Gerais, Pará, Rio de Janeiro, São Paulo), Kollmannia (2, Espirito Santo state), Latistigma (5, Espírito Santo, Minas Gerais, Rio de Janeiro, São Paulo), Pereirae (5, Minas Gerais, Rio de Janeiro, São Paulo), Solananthera (3, E Brazil), Stellandrae (1, São Paulo), Tetrachia (16, E Brazil) and Trachelocarpus (5, Paraná, Rio de Janeiro, Santa Catarina, São Paulo) have 50 spp together. and they are all endemic to Brazil - with emphasis on largest Tetrachia.

§ Lepsia (8, Panamá and tropical South America), Pritzelia (148, tropical South America to Costa Rica), Rossmannia (1, Brazil, Colombia, Ecuador and Peru) and Donaldia (3, over tropical South America) together have 170 spp. and occur exclusively in South America (all including Brazil) except for a few that extend as far as Costa Rica.

§ Ephemera (14, over tropical South America), Knesbeckia (52, over tropical New World), Wageneria (7, over tropical New World) occur from Mesoamerica to Brazil and together contain 73 spp.

§ Gireoudia (111, Mexico to Ecuador and Venezuela), Casparia (33, Costa Rica to Peru) and Ruizopavonia (32, Mexico to Bolivia) occur from Mexico (or Costa Rica) to the Central Andes and Venezuela (but not in Brazil) and together have 171 spp.

§ Cyathocnemis has 18 spp. from Colombia to Bolivia, but 4 spp. Brazil and Venezuela may belong to this group.

§ 41 spp are allocated in the section Begonia endemic to the Caribbean and 5 spp. quite isolated from E Brazil and Paraguay.

§ finally, the Doratometra section has 8 spp. spread across tropical America, inc. B. wallichiana Lehm. also collected from Vietnan (!), possibly due to human dispersion.

CUCURBITACEAE

§ FULLY CHLOROTROPHIC

Genera/species 101/945–975 Distribution tropical and subtropical regions, especially rainforests in South America and drier parts of Africa, relatively few species in Australasia and in temperate regions. Habit monoecious, andromonoecious, gynomonoecious, polygamomonoecious, dioecious, androdioecious, and gynodioecious (in Actinostemma and Schizopepon sometimes bisexual), usually perennial herbs, almost never self-supporting, mostly climbing or winding (rarely lianas, shrubs or tree, secondarily woody, or annual herbs; Dendrosicyos extremely pachycaul and secondarily arborescent with soft juicy stem). Many species are xerophytes.

Approximately 120 genera and 760 spp. with mostly tropical distribution, as they do not tolerate sub-zero temperatures; mainly herbaceous or vines lianas, few trees or shrubs. Due to its economic importance, the genus Cucurbita has been studied in detail from the point of view of its domestication. Easy to recognize at family level, the difficulties of determining Cucurbitaceae to genus and species are mainly caused by their unisexual flowers borne in different types of inflorescences (female flowers tend to be large, solitary and axillary, while the male inflorescences are racemose or paniculate and the flowers are smaller and early deciduous). Fruit normally a hard-skinned berry known as a pepo, green, white, yellow, orange or tinged with red, often spotted or striped, sometimes soft-walled, small berries (Melothria, some Cayaponia), sometimes fleshy capsules with irregular, explosive dehiscence (Cyclanthera), rarely capsular or samaroid, 1-many seeded.

Key differences from similar families differs from Begoniaceae in climbing habit, presence of tendrils and bilaterally symmetric leaves; from other climbers and vines such as Vitaceae, Sapindaceae and Passifloraceae in unisexual flowers and inferior ovary.

Momordica charantia L invasive. Cultivated genera of Neotropical origin (Cucurbita) and introduced ones (Citrullus lanatus (Thunb.) – watermelon; Cucumis L.) are of major importance as food and forage crops. South America has about 360 species of Cucurbitaceae that descend from just a few transoceanic dispersal events, mostly from Africa to South America. Of course, the undisputed record for the world's largest fruit is a 1337 pound pumpkin, a member of the gourd family (Cucurbitaceae).

SYSTEMATIC ten small lineages are absent in South America: Gomphogyneae (5/56, E and tropical Asia, E Queensland, Fiji, with their largest diversity in SE Asia), Alsomitra (1/1, SE Asia, Malesia to New Guinea), Actinostemmateae (1/4, India, China (inc. Taiwan) and E Siberia to Korean Peninsula and Japan,, Laos, Vietnam), Indofevilleeae (1/1, Assam, Bhutan, Tibet), Thladiantheae (2/30–35, temperate Asia, E and tropical Asia to Taiwan in China and Malesia), Siraitieae (1/5, E Himalayas, S China, Thailand, Vietnam, Malesia, tropical Africa), Momordiceae (1/35–40, tropical and subtropical Africa, Arabian Peninsula, tropical Asia to E Queensland), Joliffieae (3/9, tropical Africa, Madagascar), Bryonieae (3/18, Europe, Mediterranean, North Africa, Canary Islands, SW and Central Asia, NW and C Australia) and Schizopeponeae (2/9, Himalayas, Tibet, China, Japan). 435 spp. in New World, 268 in South America.

1. TRIBE TRICERATIEAE (5/23) ‣ outsider Cyclantheropsis (3; tropical Africa, Madagascar).

1. Anisosperma Silva Manso. (off Fevillea) Dioecious, perennials, woody climber, to several meters long. Only one sp., A. passiflora (Vell.) S. Manso, from gallery forest in SE Brazil from Bahia to Santa Catarina states.

2. Fevillea L. (exc. Anisosperma). 7 spp., F. cordifolia L. wider from Central America to South America, F. trilobata L. in E Brazil, Guianas and adjacent NE Argentina, F. bahiense G. Rob. & Wunderlin in Bahia state in Brazil (a rare plants in Brazil by Plantas Raras do Brasil’s book), F. anomalosperma M. Nee in NE Bolivia, and remaining three in larger areas in South America.

3. Pteropepon (Cogn.) Cogn. 6 spp., 5 from Central America to Argentina, and P. deltoideus Cogn. in Brazil, Peru, Ecuador and Colombia; in humid ravines close to rivers, primary rainforest, and secondary scrub.

4. Sicydium Schltdl. 11 spp. in Central to tropical South America (7, 3 in Brazil, none endemics), and the Caribbean and Mexico; in disturbed tropical and deciduous forest and along rivers, in dry forest and among shrubs of coastal lowlands; flowering and fruiting all year.

2. TRIBE ZANONIEAE (4/11–13) ‣ outsiders Gerrardanthus (5; tropical and southern Africa), Zanonia (1; tropical Asia), Xerosicyos (3; Madagascar).

5. Siolmatra Baill. Two spp., S. brasiliensis (Cogn.) Baill. and S pentaphylla Harms (both in Brazil, none endemics), over Amazon rainforest up to Argentina.

3. TRIBE SICYOEAE (12/c 265) ‣ outsiders Nothoalsomitra (1; Queensland); Trichosanthes (83; tropical Asia to Queensland and in the Pacific), Hodgsonia (2; Assam, Bhutan, China, Burma, Thailand, Indochina, Malesia, tropical Asia), Linnaeosicyos (1; Hispaniola), Echinocystis (1; North America), Marah (8; U.S.A., Mexico), Frantzia (6; Central America).

6. Cyclanthera Schrad. Delicate climber, flowers small and white or cream, fruit clavate and explosive, spiny, seeds ruminated at the edges. 48 species in S U.S.A., Mexico, Central and South America (24), one species extending into the Galapagos archipelago; roadsides, forest clearings, on riverbanks and cultivated ground, hedges, tropical deciduous forest, in humid lowland forest, dry xeric forest, and montane cloud forest; C. pedata (L.) Schrad. is cultivated in Asia; 9 in Brazil, C. eichleri Cogn. and C. tenuifolia Cogn. endemics.

7. Echinopepon Naudin. 18 spp., S U.S.A. States to Northern Argentina, in forest clearings, semi-deserts and ravines, on hillsides, roadsides, sand dunes and seaside gravel shores, some are weeds of cultivated ground; two spp. in South America, E. disjunctus Pozner from Cono Sur, and E. racemosus (Steud.) C. Jeffrey widely distributed in tropical New World.

8. Hanburia Seem. (inc. Elateriopsis). 8 spp. from Mexico to Peru and Venezuela; 6 in South America; in primary and disturbed rainforest, deciduous forest, and cloud forest.

9. Luffa Mill. Monoecious, annual, prostrate or scandent herbs; leaves simple, palmately 5–7-lobed, petiolate, asperulose or scabrid, petiolate. fruit globose to cylindric, rostrate, smooth, ribbed or spiny, dry, brownish, fibrous, dehiscent by an apical operculum. 8 spp., three in the New World, L. quinquefida (Hook. & Arn.) Seem. occurring from Gulf of California to Nicaragua, L. operculata (L.) Cogn. from Panamá to S Brazil, absent in Bolivia and barely in Amazon rainforest; and L. astorii Svenson in coastal Venezuela, Ecuador, and Peru; 4 in Asia, some shared with North Africa), and L. saccata F.Muell. ex I.Telford in N Australia.

10. Sicyos L. (inc. Sechium, Sicyocaulis) Monoecious, annual, climbing or trailing herbs; stems usually ± hairy; tendrils 3–5-fid; leaves simple, palmately 5- lobed, broadly ovate- or reniform-cordate in outline. 79 spp., 58 mostly from Mexico to Argentina (29 in South America), 14 in Hawaii, two in North America, 2 in Australia, one in New Zealand, Norfolk and Lord Howe, two in Galapagos (known only from the type collection and apparently extinct), S. polyacanthus Cogn. introduced in Africa; forest margins, hillsides, clearings, roadsides, pastures, seabird colonies; three spp. in Brazil, S. martii Cogn. endemic.

4. TRIBE CONIANDREAE (19/147–156) ‣ outsiders Bambekea (1; tropical Africa), Eureiandra (8; tropical and subtropical Africa, Madagascar, Socotra), Dendrosicyos (1; Yemen, Socotra), Seyrigia (5; Madagascar), Trochomeriopsis (1; Madagascar), Corallocarpus (13–16; tropical Africa, Madagascar, Arabian Peninsula, Pakistan, India), Kedrostis (c 20; Africa, Madagascar, Arabian Peninsula, India, Sri Lanka, W Malesia), Tumamoca (2; Arizona, Mexico), Ibervillea (10–11; Texas to Guatemala).

11. Apodanthera Arn. Monoecious or dioecious herbaceous climbers or trailers up 5m long. 32 spp., from Texas to Mexico and disjunct in South America; 11 in Brazil, 9 endemics; A. sagittifolia (Griseb.) Mart. Crov. from S Brazil, Argentina and Uruguay is the southernmost record of Cucurbitaceae in New World, and maybe extinct in Brazil; three spp. from Bahia state are rare plants in Brazil by Plantas Raras do Brasil’s book; three sections:

§ sect. Apodanthera ‣ 20 spp., Ecuador to Cono Sur and S Brazil (only 5 spp., 2 endemics).

§ sect. Pseudoapodanthera ‣ six species endemic to NE Brazil.

§ sect. Cucurbitopsis ‣ 6 spp., U.S.A. to Mexico.

12. Ceratosanthes Adans. 5 spp., all in Brazil, two up to adjacent Cono Sur, C. palmata (L.) Urb. up to northern South America, C. tomentosa Cogn. endemic to mainland Brazil, and C. rupicola Ridl. endemic to Fernando de Noronha island; semi-arid plains and mountain slopes, roadsides, cultivated ground. Three spp. of this genus from Fernando de Noronha Is. are rare plants in Brazil by Plantas Raras do Brasil’s book, but C. angustiloba Ridl. and C. cuneata Ridl. are recently invalid, remaining only C. rupicola Ridl.

13. Cucurbitella Walp. Only one variable sp., C. asperata (Gillies ex Hook. & Arn.) Walp., in Argentina, Brazil, Uruguay, Paraguay; in dry bushland, along roadsides, and on disturbed ground.

14. Doyerea Grosourdi. Only one sp., D. emetocathartica Grosourdy, Caribbean, Central America, Venezuela and Colombia.

15. Gurania Forest liana with showy orange/yellow flowers arranged in pseudo-umbel; some species are many ornamental. 50 spp., in Central to South America (47), in tropical forests; 29 spp. in Brazil, 18 endemics; 5 spp., one lack origin (G. velutina Cogn.), others in Pará, Bahia and Rio de Janeiro states, are rare plants in Brazil by Plantas Raras do Brasil’s book.

16. Halosicyos Mart. Crov. Dioecious perennial, herbaceous climber or trailer with large woodt rootstock. Only one sp., H. ragonesei Mart. Crov., endemic to C Argentina; on sand soils and in halophilous bushlands on the border of salinas.

17. Helmontia Cogn. Appearing dioecious, but almost certainly monoecious, with plants first male, then family, perennial, herbaceous, or woody climbers, to several meters long. Three spp., in rainforest of Guianas, Venezuela amd Brazil (two spp., none endemics).

18. Melothrianthus Mart. Crov. Dioecious, herbaceous climbres or trailers, foetid. Only one sp., M. smilacifolius (Cogn.) Mart.Crov, endemic to E Brazil, maily Espirito Santo to Rio Grande do Sul, also Bahia, Goiás and Distrito Federal; humid places; possibly 2–3 additional as yet undescribed species.

19. Psiguria Neck. ex Arn. Monoecious, perennial, vines with simple tendrils, herbaceous but some becoming woody with greater stem size and age; leaves petiolate; inflorescences pedunculate racemes or corymbs; staminate flowers axillary, corolla rotate, orange, pink, or red. Pistillate flowers axillary or terminal; corolla rotate, orange, pink, or red; fruit a pepo, oblong, indehiscent, solid green or with lighter green or white stripes, 18–80 mm long, 10–24 mm wide, wall 0.7–3.0 mm thick. 6 spp. in Central to South America, in tropical forests, 5 in South America, three in Brazil: P. ternata (M. Roem.) C. Jeffrey also in Peru, Bolivia and Paraguay; P. triphylla (Miq.) C. Jeffrey in over Neotropics; and P. umbrosa (Kunth) C. Jeffrey in Caribbean Venezuela, French Guiana and some populations in E Brazil.

20. Wilbrandia Silva Manso. Monoecious or dioecious, perennial climber or trailer with wood rootstock. 5 spp., from Argentina, Brazil (all species, two endemics, with W. glaziovii Cogn. from Rio de Janeiro state a rare plant in Brazil by Plantas Raras do Brasil’s book) and Paraguay, and rainforest and secondary scrub; W. hibiscoides Silva Manso also in Guianas (probably a mistake).

5. TRIBE BENINCASEAE (24/204–214) ‣ outsiders Citrullus (4; Mediterranean, North Africa, tropical and S Africa, W Asia), Peponium (c 20; tropical and subtropical Africa, Madagascar, Seychelles), Lagenaria (6; tropical Africa, Madagascar, one species, L. siceraria, pantropical), Acanthosicyos (1; Angola, Namibia, Botswana, South Africa), Raphidiocystis (4; tropical Africa, Madagascar), Cephalopentandra (1; NE tropical Africa), Lemurosicyos (1; Madagascar), Solena (3; tropical Asia), Borneosicyos (1; Sarawak, Sabah), Benincasa (2; New Caledonia, New Ireland, New Guinea, Queensland; known only from cultivation), Ctenolepis (1; Madagascar), Dactyliandra (2; the Namib desert in Angola and Namibia, the Thar desert in Pakistan and India; Kenya), Khmeriosicyos (1; Cambodia; probably extinct), Papuasicyos (c 8; New Guinea), Scopellaria (2; Yunnan, SE Asia, W Malesia to Philippines), Trochomeria (8; subtropical and tropical Africa), Indomelothria (2; SE Asia, W Malesia), Ruthalicia (2; tropical W Africa), Muellerargia (2; Madagascar; the Lesser Sunda Islands, Timor, tropical N Australia), Cucumis (c 55; subtropical and tropical regions in the Old World), Zehneria (c 60; tropical regions in the Old World), Diplocyclos (4–5; subtropical and tropical Africa, tropical Asia, tropical Australia), Coccinia (c 25; tropical and southern Africa, one species, C. cordifolia, also in tropical Asia).

21. Melothria L. Annual or perennial herbs, scadent or prostate, fragile. 14 spp., Central and South America; 12 spp., 8 in Brazil, M. hirsuta Cogn. endemic; M. sphaerocarpa (Cogn.) H. Schaef. & S.S. Renner disjunct of Central and South America and W tropical Africa; roadsides, and cultivated ground, arid plains, clearing, forets margins, grass or woodlands.

6. TRIBE CUCURBITEAE (11/100–110) ‣ outsiders Polyclathra (1; Mexico, Central America), Peponopsis (1; Mexico), Sicana (3; Central America, Caribbean), Penelopeia (2; Hispaniola), Schizocarpum (11; Mexico, Guatemala), Cionosicyos (4; Central America, Cuba, Jamaica).

22. Abobra Naudin. Dioecious, herbaceous, perennial trailer, to 7m long, with feshy rootstock. Only one sp., A. tenuifolia (G. in Hook) Cogn. in Argentina and Uruguay; in xeric bushlands and dry soils.

23. Calycophysum Trianna. Monoecious, perennial or woody climbers, root non-tuberous, to 10 m long. 4 spp. from Venezuela to Bolivia, in Andean cloud forests and lowland rainforests.

24. Cayaponia Silva Manso. Monoecious or rarely dioecious, herbaceous or suffrutescent, often much-branched climbers or creepers, to 20 m long, with perennial roots; leaves triangular-ovate, shallowly 3-palmatilobed, 3-nerved; fruit dry. 71 spp., mainly in South America (66), few in Central America, Mexico, U.S.A.; one sp. endemic to Fernando de Noronha Island; 1-2 spp. from W Africa and Madagascar; 48 spp. in Brazil, 18 endemics, 4 of them from Minas Gerais, Bahia, Goias and C. noronhae C. Jeffrey, endemic to Fernando de Noronha islands in Atlantic Ocean, are rare plants in Brazil by Plantas Raras do Brasil’s book.

25. Cucurbita L. Monoecious, perennial or annual, herbaceous climbers or trailers to 6m long. 14 spp., 9 wild spp. and 5 domesticated (C. argyrosperma Huber, C. ficifolia Bouche, C. maxima Duchesne, C. moschata Duchesne, and C. pepo L.), in tropical and subtropical America; disturbed places, humid ravines, floodplants, tropical deciduous forests, grasslands, deserts, rocky hillsides and oak-pine forests; five of the wild Cucurbita species are xerophytic perennials that occur in Mexico and the SW U.S.A.; seven wild species are mesophytic annuals that occur in Mexico, the SE and SC U.S.A., and Central and South America.

5 clades occur within genus, only one with species in South America, Group Maxima, with C. ecuadoriensis H.C. Cutler & Whitaker endemic to W Ecuador and C. maxima Duchesne, with their wild form C. maxima subsp. andreana endemic to center Bolivia; subfossil records of C. pepo L. and C. moschata Duchesne ex Poir. from Central America and the northern Andes indicate that squashes are among the oldest neotropical domesticated plants.

26. Tecunumania Standl. & Steyerm. Two spp., one from Mexico to Costa Rica, and T. stothertiae Cornejo & Schaefer. endemic to Ecuador.

34. CELASTRALES

TWO FAMILIES, BOTH IN SOUTH AMERICA.

LEPIDOBOTRIACEAE

§ FULLY CHLOROTROPHIC

Genera/species 2/2 Distribution tropical W Africa, Central America and northern South America to Peru. Habit dioecious, evergreen trees (or shrubs?).

SYSTEMATIC outsider Lepidobotrys (1; tropical W Africa).

1. Ruptiliocarpon Hammel & N.Zamora. High tree, in dense rainforests. Only one sp., R. caracolito Hammel & N.Zamora from C Suriname, W Colombia (50 m to 800 m, in Antioquia, Valle), NE Peru (center Loreto), Costa Rica and Nicaragua, and recently discovery in Acre and Rondona states in NW Brazil.

CELASTRACEAE

§ FULLY CHLOROTROPHIC

Genera/species 98/1.280–1.290 Distribution mainly tropical and subtropical regions in the Northern and Southern Hemispheres; some species in temperate areas; Pottingeria acuminata: Assam (Naga Hills), upper Burma, NW Thailand. Habit bisexual, monoecious, andromonoecious, polygamomonoecious?, dioecious, gynodioecious, or polygamodioecious?, usually evergreen or deciduous trees, shrubs or lianas (rarely herbs, suffrutices, or ericoid or epiphytic shrubs; Stackhousia usually annual or perennial herbs, sometimes succulent). With or without spines (Acanthothamnus and Canotia with glandular stems). Branches rarely photosynthesizing phyllocladia. Bark often yellow (due to triterpenic compounds).

Several Neotropical species (Anthodon decussatum Ruiz & Pav., Cuervea crenulata Mennega, Elachyptera festiva (Miers) A.C. Sm., E. floribunda (Benth.) A.C. Sm., E. micrantha (Cambess.) A.C. Sm., Hippocratea volubilis L., Prionostemma aspera (Lam.) Miers, Pristimera celastroides (Kunth) A.C. Sm., P. nervosa (Miers) A.C. Sm., P. verrucosa (Kunth) Miers., Semialarium mexicanum (Miers) Mennega, S. paniculatum (Mart.) N. Hallé) are large lianas that grow both along borders and inside forests where wind may or may not help with dispersal; other Neotropical species (Cuervea kappleriana (Miq.) A.C. Sm., Hylenaea comosa (Sw.) Miers, H. praecelsa (Miers) A.C. Sm., and Pristimera tenuiflora (Mart. ex Peyr.) A.C. Sm.), are water-dispersed and have been reported from riparian and floodplain forests. 19 genera and about 130 spp. in Brazil.

Nicobariodendron has dioecy, distichous leaves, no stipules, mucilage cells, racemose inflorescence, intrastaminal disc, two stamens, basal placentation and drupe. It is known only from the type material from Nicobar Islands. 439 spp. in New World.

SYSTEMATIC two high clades, both in South America.

1. CLADE PARNASSIOIDEAE (2/70) - outsider Parnassia (c 70; cold-temperate regions on the Northern Hemisphere south to Morocco, Sumatra and Mexico, with their largest diversity in Himalayas, W China and NW North America).

1. Lepuropetalon Elliott. Many small annual succulent herbs ca. 2 cm tall, rarely larger, the smallest terrestrial angiosperm from several countries as Brazil and Chile, winter growing, leaves spathulated, flowers incompiscuous, corolla absent, usually solitary, if not then paired; tannin sacs visible as red lines or dots on the leaves and calyx, especially when dry. Only one sp., L. spathulatum (Muehl.) Eliot, found by looking very carefully in disturbed areas, growing amidst grasses and other small annual herbs, mosses and liverworts, some times in damp and often sandy soil and outcrops., in disjunct areas of SW U.S.A., Mexico, chefly along the Andes of Ecuador, central Chile, S Brazil (recorded of Porto Alegre municipality [Morro São Pedro region], in Rio Grande do Sul state), NE Argentina, Paraguay and Uruguay.

2. SUBFAMILY CELASTROIDEAE (102/1,180 – 1,210) - 98 genera within 14 lineages (four genera unplaced); six outside South America: Pottingerioideae (2/5, SE Asia, SW U.S.A., northern Mexico), Monimopetalum clade (1/1, China), Gymnosporia Clade (8/113, Tropical and subtropical regions in the Old World, tropical Asia to tropical Australia and New Caledonia, Caribbean), Old World Maytenus Clade (6-7/8-?, Old World), Salaciopsis Clade (1/6, New Caledonia) and Sarawakodendroideae (1/1, Borneo).

CELASTROIDEAE ▸ UNPLACED GENERA

2. Goniodiscus Kuhlm. Trees, glabrous, tree, fruit a large drupe, seed exalbuminous, oily. Only one sp., G. elaeospermus Kuhlm., endemic to Amazon rainforest (only Amazonas state) in Brazil; the oil extracted from its seeds supported a small business at the type location.

3. Prionostemma Miers. Tall, glabrous scandent shrubs; without latex; leaves opposite, entire, rarely dentate; inflorescences axillary, rarely terminal, thyrsoid; flowers bisexual, 5-merous, disk fleshy, annular-pulvinate, entire, extrastaminal. 4 spp., P. asperum (Lam.) Miers from Mexico, Central America, Trinidad & Tobago, Venezuela, Colombia, Bolivia, Brazil, and three in Africa and India, savannas and forests.

2.1 CELASTROIDEAE ▸ MICROTROPIS CLADE (3/94) - outsiders Microtropis (66; tropical Asia), Quetzalia (11; S Mexico, Central America).

4. Zinowiewia Turcz. Trees or shrubs, leaves opposite; fruit like Plenckia, a samara with a lateral-apical wing, oblanceolate or obovate, curved like a sable. 13 spp., Mexico to Peru, 6 in South America, Venezuela (one endemic), Colombia (2 endemics), Ecuador (one endemic), Peru and Bolivia, with Z. australis Lundell also collected in Roraima state in N Brazil.

2.2 CELASTROIDEAE ▸ STACKHOUSIOIDEAE (18/89) - outsiders are Wilczekra (1; Central Africa); Hexaspora (1; NE Queensland); Menepetalum (4; New Caledonia), Dinghoua (1; NE Queensland), Apatophyllum (5; Queensland, New South Wales), Psammomoya (4; W Australia); Tripterococcus (4; W Australia), Macgregoria (1; C Australia), Stackhousia (16; Malesia to New Guinea, Australia, Tasmania, New Zealand, Micronesia); Denhamia (10; N and E Australia), Brassiantha (1; New Guinea), Hedraianthera (1; Queensland, New South Wales), Dicarpellum (4; New Caledonia), Hypsophila (3; NE Queensland), Siphonodon (6; tropical Asia to New Guinea, E Queensland, NE New South Wales), Peripterygia (1; New Caledonia), Xenodrys (1; Madagascar).

5. Crossopetalum P. Browne. Small trees or shrubs, inflorescence axillary, cymose, ovary (2-) 4-locular with ovules erect, small drupes. 3 in Old World, inc. C. serrulatum (Loes.) I.Darbysh., the most geographically widely distributed species in the entire Celastraceae family, and 20 spp. in Florida, Caribbean to Venezuela, two in South America, C. rhacoma Crantz, Central America to Caribbean Is. of Colombia and Venezuela, Antilhas, and C. eucymosum (Loes. & Pittier) Lundell endemic to Peru.

2.3 CELASTROIDEAE ▸ BREXIOIDEAE (15/108–123) - outsiders are all in Madagascar except Brexia (1 or 12; coastal regions in tropical E Africa, Madagascar, Seychelles), Kokoona (10; Sri Lanka to Malesia), Lophopetalum (18–20; tropical Asia to tropical Australia); Pleurostylia (5; tropical and S Africa, Madagascar, Mascarene Islands, tropical Asia to New Guinea, N Australia, New Caledonia).

6. Elaeodendron Jacq. Dioecious evergreen trees, pistillate flowers with petaloid staminodes (Neotropics), fruit a large indehiscent drupe with a hard stone enclosing seeds. 6 spp. in New World, two only in Mexico, three only in Caribbean, and E. xylocarpum (Vent.) DC. from Mexico, Central America, Caribbean and Venezuela.

2.4 CELASTROIDEAE ▸ SALACIODEAE (5/270) - outsiders are Salacighia (2; tropical W and C Africa to Angola), Thyrsosalacia (2; Central Africa).

7. Cheiloclinium Miers. Lianas os scandent trees or shrubs, glabrous or glabrescent; leaves opposite, entire, crenate, or serrate; inflorescence axillary, thyrsoid or cymose; flowers bisexual, 5-merous. 16 spp., all in South America, 4 up to Mesoamerica (only one in Mexico) to SE Brazil (11 in Brazil, 4 endemics), humid forests.

8. Peritassa Miers. Lianas os scedent trees or shrubs, often with xylopodium; glabrous (some inflorescence pilose); leaves opposite, subopposite, or alternate, entire, crenate, or serrate; inflorescences axillary, thyrsoid or cymose; flowers bisexual, 5-merous. 20 spp., humid forests, of Costa Rica to Paraguay, all in South America, 16 in Brazil, 9 endemics; two spp., both collected in Espírito Santo state, are rare species in Brazil by Plantas Raras do Brasil’s book.

9. Salacia L. Tall unarmed lianas or scadent shrubs, rarely completely erect trees or shrubs, often with xylopodium, glabrous or glabrescent; leaves opposite or suboppposite, rarely alternate, entire, crenate, or serrate; inflorescence axillary or caulifllrous, thyrsoid, cymose or fasciculate. 200 spp., Old and New World Tropics, Australia, humid forests to dry scrub and grasslands, 40 spp. in New World, 35 in South America, 27 in Brazil, 8 endemics; S. nemorosa Lombardi is a rare species in Brazil, collected in Espírito Santo state, by Plantas Raras do Brasil’s book.

10. Tontelea Miers. Lianas os scedent trees or shrubs, glabrous or puberulent, often with xylopodium; leaves opposite to subopposite, entire, crenate or serrate; inflorescence axillary, thyrsoid, rarely cymose; flowers bisexual, 5-merous. 24 spp., all in South America, two up to Mexico and Central America, from lowands to 1,800 m alt; 14 spp. in Brazil, 5 endemics.

2.5 CELASTROIDEAE ▸ HIPPOCRATEOIDEAE (18/106) - outsiders are Plagiopteron (1; S China, S Burma, Thailand); Helictonema (1; tropical Africa); Arnicratea (3; India to SE Asia), Bequaertia (1; tropical Africa), Apodostigma (1; tropical Africa, Madagascar), Reissantia (7; tropical regions in the Old World), Loeseneriella (16; tropical regions in the Old World), Campylostemon (10; tropical Africa), Tristemonanthus (2; tropical W and C Africa), Simicratea (1; Angola), Trochantha (2; tropical Africa).

11. Anthodon Ruiz & Pavon. Lianas, glabrous; leaves opppsite or subopposite, crenulate or serrulate; inflorescences axillary, cymose; flowers bisexual, 5-merous; petals regularly serrate; disk fleshy, copular, entire, extrastaminal. Two spp., A. decussatus Ruiz & Pav. from Central America, Venezuela, Guianas, Colombia, Ecuador, Peru, Bolivia, Brazil, Cono Sur, and A. panamensis A.C. Sm. from Colombia and Ecuador.

12. Cuervea Triana ex Miers. Tall-growing, unarmed lianes; latex present or absent; leaves opposite or subopposite, entire, crenulate, or denticulate; inflorescences axillary, rarely terminal, cymose or thyrsoid; flowers bisexual, large, 5-merous; disk membranous, copular, entire or lobed. 7 spp., two in Africa, Caribbean islands (three endemics, St Vincent, Cuba, and Jamaica), C. kappleriana (Miq.) A.C. Sm. from Mexico to Bolivia, Guianas and Caribbean, and C. crenulata Mennega endemic to Brazil, and at humid to gallery forests.

13. Elachyptera A.C.Sm. Glabrous lianes or scrambling shrubs; latex absent, glabrous (inflorescences sometimes puburulent); leaves opposite or subopposite, entire, crenulate, or serrate; inflorecences axillary or terminal, cymose or thyrsoid; flowers bisexual, 5-merous, disk almost fleshy. 7 spp., 3 in Africa and one in Madagascar, 4 of Mexico to SE Brazil (all 4, slightly centered within up to Bolivia and Argentina, E. coriacea Lombardi from Bahia statet a rare species in Brazil, by Plantas Raras do Brasil’s book), in humid forests to dry thickets, mangroves swamps.

14. Hippocratea L. Lianas, puberulent or glabrescent; leaves opposite, entire, crenate, or serrulate; inflorescences axillary, cymose or thyrsoid; flowers bisexual, 5-merous; disk fleshy, pulvinate or cupular, entire, extrastaminal. 3 spp., two in tropical Africa, and H. volubilis L. from Central and South America, scatered in almost all countries, at humid to gallery forests, 0-1,800 m.

15. Hylenaea Miers Trees or lianas, glabrous; leaves oppsoite or subopposite, entire; inflorescences axillary or terminal, thyrsoid; flowers bisexual, 5-meroys; disk membranous or more or less fleshy, cupular, entire. Three spp., Central and South America, forests, all well distributed (all in Brazil) but H. unguiculata Mennega more restricted, from Brazil and Guianas.

16. Pristimera Miers. Lianas or scandent shrubs, glabrous or puberulent; older stems normally terete, with pale yellow wood; latex usually absent; leaves opposite or subopposite, entire or serrulate; inflorescences axillary, rarely terminal, cymose, rarely thyrsoid; flowers bisexual, small, 5-merous, fleshy. 21 spp., Indonesia, Madagascar, Africa (12), 11 in New World tropics from Mexico to Cono Sur and Caribbean, humid forests to dry tickets; 9 spp. in South America, 4 spp. in Brazil, P.a sclerophylla Lombardi endemic.

17. Semialarium N.Hallé. Trees or lianas, puberulent or glabescent; leaves opposite, rarely subopposite, crenulate or serrulate; inflorescences axillary, rarely terminal, thyrsoid; flowers bisexual, 5-merous, fleshy, entire. Two spp., S. mexicanum (Miers) Mennega from Mexico, Venezuela and Central America, and S. paniculatum (Mart.) N. Hallé from Brazil, Colombia and Cono Sur, forests, to 1,300 m.

2.6 CELASTROIDEAE ▸ AMERICAN MAYTENUS CLADE (5/145-160) - all genera in South America.

18. Fraunhofera Mart. Trees or shrubs, pubescents, sessile and pubescent leaves, axillary and terminal racemose inflorescences with strigose peduncle, fruit possibly a drupe. Only one sp., F. multiflora Mart., endemic to NE Brazil, in deciduous forets.

19. Maytenus L. (exc. Monteverdia p.p.). Shrubs to trees, often with woody rhizomes; 36 spp., Florida and Texas to subantarctic regions in Tierra del Fuego, Argentina, with a large altitudinal variation from sea level to ca. 3,900 m elevation in the Andean mountains. 28 spp. in South America, only the southern South American M. boaria Molina occur in Brazil, in high elevations in Minas Gerais to Santa Catarina states.

20. Monteverdia A. Rich. (inc. Maytenus p.p.) Membranaceous, chartaceous or coriaceous leaves that are spirally or distichously arranged; axillary fasciculate or cymose inflorescences that are sometimes reduced to an individual flower; flowers with two carpels, two ovules per carpel; capsular fruits with coriaceous pericarp that open by two reflexing or (rarely) upright valves; white arils that completely cover the seeds. 123 spp., over New World, 81 in South America, 51 in Brazil, 34 endemics; 7 spp. of this genus are rare in Brazil, all treelets in Bahia, Minas Gerais and Rio de Janeiro states, by Plantas Raras do Brasil’s book. M. multicostata Cornejo & Biral (Ecuador) has the largest fruits in the genus (to 3.5 cm long).

21. Plenckia Reissek. (inc. Viposia) Trees or shrubs, glabrous, often dioecious, leaves alternate, fruit a samara with an apical wing, surrounding seed, elongate or oblong, symmetrical, and eventually septicidally dehiscent capsule with age. 5 spp. from Bolivia, Brazil (3, P. bahiensis Loes. endemic) and Cono Sur.

22. Tricerma Liebm. Thick and fleshy leaves that are spirally arranged; axillary fasciculate inflorescences; flowers with three carpels, one ovule per carpel; capsular fruits with coriaceous pericarp that open by three reflexing valves; red arils that completely cover the seeds. 5 spp., two from North America to Mexico and Caribbean, one from Ecuador and Peru, and two from Bolivia, Paraguay and Argentina.

2.7 CELASTROIDEAE ▸ CELASTROIDEAE (10/200) - outsiders are Tripterygium (1; China inc. Taiwan); Paxistima (2; North America), Wimmeria (12; Central America); Acanthothamnus (1; Mexico), Canotia (2; SW U.S.A); Euonymus (c 130; temperate regions on the Northern Hemisphere, E Australia, Tasmania), Glyptopetalum (c 20; tropical Asia), Torralbasia (1; Caribbean), Xylonymus (1; W New Guinea).

23. Celastrus L. Scadent shrubs, glabrous, rarely pubescents. 41 spp., 13 in New World, C. scandens L. in North America, and remaining from Mexico to South America (8), also Madagascar and SE Asia to Australia, only the widely distributed C. liebmannii Standl. in Brazil.

2.8 CELASTROIDEAE ▸ SCHAEFFERIA CLADE (5/32) - outsiders are Orthosphenia (1; Mexico), Rzedowskia (1; Mexico).

24. Haydenoxylon M. P. Simmons. Dioecious trees, inflorescence large, cymose with 4-merous flowers, fruit a loculicidal capsule, 2-4 lobed. 4 spp., one in Colombia to Peru; two in Mexico and Central America (inc. H. calzadae (Lundell) Biral off VPA); and H. urbanianum (Loes.) M. P. Simmons from Colombia to Bolivia and Acre state in Brazil.

Gymnosporia magnofolia (Loes.) Lundell from Peru is a dubious taxa, possibly in Haydenoxylon and rejected here.

25. Myginda Jacq. Unisexual shrubs, flowers white, 2-locular, ovules pendulous, fruit 2-locular. Two spp., one endemic to Mexico, and M. uragoga Jacq. from Mexico, Central America, Caribbean and Colombia.

26. Schaefferia Jacq. Trees or shrubs, glabrous or rarely puberulent. 15 spp. from Mexico to Argentina and Uruguay, 4 spp. in South America, only the platine species S. argentinensis Speg. in Brazil, dry, rochy woodlands, hummocks, thickets.

35. OXALIDALES

FAMILIES ABSENTS IN SOUTH AMERICA: CEPHALOTACEAE (1/1) AND HUACEAE (2/4).

LINEAGE 1 of 2: OXALIDS

CONNARACEAE

§ FULLY CHLOROTROPHIC

Genera/species 13(own data, with Berberdinia)/160-190 Distribution pantropical, southwards to Uruguay, South Africa and Queensland. Habit usually bisexual (sometimes monoecious, rarely dioecious), usually evergreen (sometimes deciduous) shrubs or lianas (sometimes trees). Many species are poisonous. All five genera are native. Three out of the 5 native genera are endemic to Neotropics. Their geographic distribution within the region is given above under "Distribution in the Neotropics". About 70 spp. in Brazil.

The family is pantropical and includes 16 genera and about 300-350 species. Its center of distribution is W Africa. In the Neotropics there are 5 genera and 111 species. While most species are found mostly within the tropics of Cancer and Capricorn, two taxa extend past 27° S in the state of Santa Catarina, Brazil: Connarus rostratus (Vell.) L.B.Sm. and Rourea gracilis G.Schellenb. Both Rourea (Rourea induta Planch.) and Connarus (Connarus suberosus Planch.) are important components of savannas of C Brazil (cerrado).

Key differences from similar families - the lack of stipules is a key difference from the Leguminosae in sterile herbarium material.

Use Timber, medicinal plants, seed oil (Connarus).

SYSTEMATIC outsiders are Ellipanthus (6; coastal regions in tropical E Africa, Madagascar, tropical Asia), Hemandradenia (2; tropical W and C Africa), Burttia (1; C Tanzania), Vismianthus (2; Mlinguru in SE Tanzania; SW Burma); Jollydora (3; E Nigeria to Angola); Manotes (4–5; tropical Africa); Cnestis (13; tropical Africa, Madagascar, one species in tropical Asia), Agelaea (8; tropical Africa, Madagascar, tropical Asia)

Keys to genera of Neotropical Connaraceae

1. Carpel solitary at flowering stage; petals with glandular punctuations ------------ Connarus

1. Carpels 5 at flowering stage; petals without glandular punctuations - 2

2. Sepals valvate or only narrowly imbricate; inflorescence (and fruit) densely pilose ------------ Cnestidium

2. Sepals imbricate; inflorescence (and fruit) usually glabrous or villous but not densely pilose - 3

3. Ovary slightly stipitate; leaves consistently trifoliolate; leaflets papillose below ------------ Pseudoconnarus

3. Ovary sessile; leaves uni-, tri- or multifoliolate; leaflets usually without - 4

4. Flowers produced before or approximately at the same time as the leaves ------------ Bernardinia

4. Flowers produced after the leaves ------------ Rourea

1. Bernardinia Planch. Only one sp., B. fluminensis (Gardner) Planch., endemic to E part of Brazil, in states of Bahia, Minas Gerais, Rio de Janeiro, extending north to Pará and south to São Paulo.

2. Cnestidium Planch. Lianas with imparipinnate leaves, inflorescence a panicle. Three spp., C. guianensis (G. Schellenb.) G. Schellenb. in the Guianas and E Venezuela, C. rufescens Planch. is known from Mexico to Colombia, W Venezuela, Ecuador and Cuba, and C. froesii Pires in known only N Brazil (only type collection, Amazonas state) and Venezuela.

3. Connarus L. Lianas or lianescent shrubs with cylindric branches; leaves trifoliolate or imparipinnate; leaflets opposite or not, often acuminate, often glandular-punctate; inflorescence an axillary panicle, often appearing terminal and more compound due to reduced leaf development; flowers bisexual, heterostylous; pedicels with a distinct joint; sepals 5; petals 5, longer than sepals, free or coherent, usually white; follicle red, opening lengthwise, usually along ventral suture; seed solitary, attached to ventral side of follicle. 75-80 spp., pantropical, 63 in Neotropics, 54 in South America, 38 in Brazil, 23 endemics, 6 of then from Pará, Rio de Janeiro, Bahia and Minas Gerais states are rare plants in Brazil, by Plantas Raras do Brasil’s book.

4. Pseudoconnarus Radlk. Trifoliolate leaves, always with papillae on the lower surface of the leaflets. 5 spp., Colombia, Venezuela, Guyana, Suriname, Peru, and the states of Acre, Amazonas and Pará in Brazil (3, none endemics).

5. Rourea Aubl. Shrubs to trees, often with roots crown; 55-70 spp., pantropical, 51 in Neotropics, 47 in South America, 40 in Brazil, 24 endemics, 7 of them from Bahia, São Paulo, Distrito Federal, Mato Grosso and Maranhão states are rare plants in Brazil, by Plantas Raras do Brasil’s book.

OXALIDACEAE

§ FULLY CHLOROTROPHIC

Genera/species 5/667 Distribution Tropical and subtropical regions, few species of Oxalis in temperate areas (some species of Oxalis are widely distributed weeds). Habit Usually bisexual (in Dapania androdioecious), evergreen shrubs or trees (rarely lianas) or perennial (rarely annual) herbs, often with root tubers, sometimes succulent. Some species are xerophytes or helophytes. Juice often bitter. Some species of Oxalis have CAM physiology. Two neotropical genera: Oxalis and Biophytum.

359 spp. in New World. Oxalis corniculata L. and O. pes-caprae L. are naturalized, and O. tetraphylla (L.) DC. is cultivated. O. tuberosa Molina, known as maca, is an herbaceous perennial plant that overwinters as underground stem tubers. The plant was brought into cultivation in the central and southern Andes for its tubers, which are used as a root vegetable. The plant is not known in the wild, but populations of wild Oxalis species that bear smaller tubers are known from four areas of the central Andean region.

Distinguishing characters (always present)

ü compound (or unifoliolate) leaves.

ü determinate inflorescences.

ü perfect, radial, hypogenous flowers.

ü ten stamens with two different lengths.

ü fruits loculicidal capsules.

SYSTEMATIC outsiders Averrhoa (5; Malesia), Dapania (3; Madagascar, Malesia), Sarcotheca (12; W Malesia)

1. Biophytum DC. Herbs up to 1m, stems sometimes woody; imparipinnate leaves clustered at the branch apices, and the terminal leaflet is reduced to a bristle-like mucro. 81 spp., pantropical, 37 neotropical, 31 in South America, in rain forests or disturbed areas from sea level to about 2,000 m alt., slightly centered in Andes and Guiana Shield; 3 spp. in Brazil, all from mountains of northern Amazon rainforest, none endemics.

2. Oxalis L. Annual or perennial herbs, bulb-like tubers, bulbs or fleshy rhizomes, sometimes succulent, with shrubs/vines in South America, one aquatic in South Africa, often cushions in Andes; trifoliolate leaves, either pinnate or palmately arranged. 566 spp., widely distributed, 325 in New World, from which 276 species are found throughout South America, found in both temperate and tropical habitats, growing from sea level to an altitude of more than 3,700 m - with the largest diversity in terms of Oxalis growth forms, including shrubs, herbs, annuals, vines and geophytes; c. 210 species are endemic to South Africa, all bulbous perennials with above-ground plant parts borne on seasonal rhizomes emergent during the rainy season; 97 spp. in Brazil, 57 endemics; 10 species of this genus in several Brazilian states rare plants in Brazil, by Plantas Raras do Brasil’s book. 4 subgenera and 28 sections, based mainly on characters of the leaf:

§ subg. Oxalis ‣ 413 spp., worldwide, highly centered in the Andean mountains from Colombia to Bolivia, also very diverse in South Africa; 19 sections. 42 spp. in Brazil.

§ subg. Monoxalis ‣ two spp. herbaceous, O. dichondrifolia A. Gray and O. robusta Kunth, from the SW U.S.A. and Mexico.

§ subg. Thamnoxys ‣ 72 spp., mainly in Brazilian Shield in central and E Brazil (54); nine sections have been recognized:

§ sect. Foliosae ‣ 5 spp.

§ sect. Hedysaroideae ‣ 6 spp.

§ sect. Holophyllum ‣ 8 spp.

§ sect. Phyllodoxys ‣ 2 spp.

§ sect. Pleiocarpa ‣ only one sp., O. glaucescens Norlind, endemic to Brazil.

§ sect. Polymorphae ‣ 14 spp., from Atlantic Forest except O. alstonii Lourteig and O. roselata A. St.-Hil. up to dry seasonal scrubland of NE Brazil (caatinga) and savannas of C Brazil (cerrado).

§ sect. Psoraleoideae ‣ 5 spp.

§ sect. Robustae ‣ 9 spp.

§ sect. Thamnoxys ‣ 27 spp., two in Cuba, 6 in Andes, and 19 in Brazil.

§ subg. Trifidus ‣ two spp., O. sleumeri Lourteig and O. tacorensis Burtt, from NE Chile, NW Argentina and SW Bolivia.

Oxalis is the only eudicot genus to evolve geographically, morphologically, and taxonomically diverse clades of bulb-bearing species, and two major radiations of bulb-bearing species exist within the genus. More than 200 species of caulescent, tunicate-bulbed (onion-like) Oxalis are centered on the Cape Floristic Region in southern Africa. In addition, around 60 species of acaulescent, imbricate-bulbed (lily-like) Oxalis occupy the Americas with a broad distribution in mountainous areas from Patagonia to the NE U.S.A. O. gigantea Barnéoud, endemic to Chile, is largest species of genus.

LINEAGE 2 of 2: CUNONIIDS

BRUNELLIACEAE

§ FULLY CHLOROTROPHIC

Genera/species 1/60 Distribution Mexico to Bolivia and Caribbean. Habit evergreen trees, unarmed.

The presence of interpetiolar stipules and stipels on the rachis of the compound leaves or on the petiole of the unifoliolate leaves are diagnostic and are particularly useful to distinguish them from high altitude species of Turpinia (Staphylaceae). The latter present the same variation of the leaves but without stipules and stipels.

SYSTEMATIC a single genus.

1. Brunellia Ruiz & Pavon. Tree or treelets; plants dioecious or gynodioecious; stem and branches terete, unbranched in lower part; branch nodes swollen; leaves petiolate, stipulate, unifoliolate or multifoliolate with 3-26(30) leaflets, opposite or whorled; inflorescences proliferating monotelic thyrso-paniculate, generally pedunculate; flowers apetalous, 4-16 mm in diameter, usually largest in unifoliolate species; fruit of various follicles; seeds red, 2 to 3 per follicle. 62 spp., one only to Mexico/Central America, 6 shared from Central and South America (B. comocladifolia Bonpl. also in Antilles), 54 from Venezuela to Bolivia (montane forests, either at lower elevations such as from 600 to 2,000 m, or much higher, usually between 2,800 and 3,800 m, a single spp., B. hygrothermica Cuatrec., grows in the superhumid sea-level region, high diversity in Colombia, 36), and B. neblinensis Steyerm. & Cualtrec, in high shrub land in Mount Neblina, Venezuela, possiby in Brazil.

CUNONIACEAE

§ FULLY CHLOROTROPHIC

Genera/species 27/315–325 Distribution mainly the Southern Hemisphere between 13ºS and 35ºS, with their largest diversity in Australia, New Caledonia and New Guinea, few species northwards to Philippines, Mexico and Caribbean. Habit usually bisexual (rarely andromonoecious, polygamomonoecious, dioecious, androdioecious, gynodioecious, or polygamodioecious), usually evergreen (rarely deciduous) trees or shrubs (some species of Weinmannia are lianas or hemi-epiphytes). Some species are xerophytes. Bark provided with numerous lignified cells and usually with lenticels. Species in several genera, including Weinmannia and Eucryphia, are sources of nectar for honey.

Key differences from similar families - Weinmannia resembles Brunellia and Staphylea in having opposite, usually toothed leaves (simple or pinnate in Brunellia, imparipinnate in Staphylea) and stipules (interpetiolar in Brunellia, free-lateral in Staphylea), and all three genera occur in montane neotropical forest. Weinmannia can be distinguished as follows:

ü it has a syncarpous ovary (2 carpels) with free stylodia (vs carpels free, 4-6 [-8] in Brunellia; carpels 2-3, partially free or united at level of ovary, stylodia fused distally in Staphylea);

ü axillary racemes (vs axillary panicles/thyrses in Brunellia; axillary or terminal panicles in Staphylea);

ü stipels absent at base of leaflets in compound leaves (vs stipels present in Brunellia; small glands present in Staphylea);

ü petals caducous, often falling like a cap (vs absent in Brunellia; present and not caducous in Staphylea);

ü seeds hairy (vs arillate in Brunellia; without aril or hairs in Staphylea).

Key to genera of Neotropical Cunoniaceae:

1. Flowers large (3-5 cm diameter) with showy petals, solitary ------------ Eucryphia

1. Flowers small (< 1 cm diameter) with or without petals, several to many per inflorescence - 2

2. Inflorescence paniculate/thyrsoid; stipules free-lateral (4 per node); venation craspedodromous ------------ Caldcluvia

2. Inflorescence of axillary racemes (panicle in one species of Weinmannia from Colombia); stipules various; venation semi-craspedodromous - 3

3. Leaves palmately compound; stipules free-lateral (4 per node); corolla absent ------------ Lamanonia

3. Leaves simple or imparipinnately compound; stipules interpetiolar (2 per node); corolla present but caducous ------------ Weinmannia

SYSTEMATIC tribes Spiraeanthemeae (2/20, Moluccas, New Guinea, NE Queensland, New Caledonia, Fiji, Samoa, Solomon Islands, Vanuatu), Hooglandiaclade (1/1, New Caledonia), Bauereae (1/4, SE Australia, Tasmania), Davidsonia clade (1/3, E Queensland, New South Wales), Aistopetalumclade (1/2, New Guinea), Schizomerieae (4/14, Cape, Moluccas, New Guinea, E Queensland, NE New South Wales, Solomon Islands, New Zealand, W Tasmania), Acrophyllum clade (1/1, New South Wales), Gillbeeaclade (1/3, New Guinea, Queensland) and Codieae (3/16, Moluccas, Queensland, New South Wales, New Caledonia) do not occur in South America.

1.1 TRIBE EUCRYPHIEAE (1/7) ‣ a single genus.

1. Eucryphia Cav. 7 spp., 5 in Australia and Tasmania and two in Chile and Argentina.

1.2 TRIBE GEISSOIEAE (4/25) ‣ outsiders Pseudoweinmannia (2; Queensland, New South Wales), Karrabina (2; Queensland, New South Wales), Geissois (16; New Caledonia, Vanuatu, Fiji, Santa Cruz Islands).

2. Lamanonia Vell. Small to medium-sized trees or occasionally shrubs characterized by opposite and decussate, palmately compound leaves, each usually composed of three or five leaflets that always have toothed margins. 5 spp., almost all restricted of forests in Central, E and S Brazil, except two reaching to Argentina and Paraguay; L. brasiliensis Zickel & Leitão from Distrito Federal and L. chabertii (Pamp.) L.B. Sm. from São Paulo state are rare plants in Brazil, by Plantas Raras do Brasil’s book.

1.3 TRIBE CALDCLUVIEAE (4/10) ‣ outsiders Opocunonia (1; E Malesia to New Guinea, Solomon Islands), Ackama (10; Sulawesi, Philippines, Moluccas, New Guinea, Solomon Islands, Queensland, New South Wales, North Island of New Zealand).

3. Caldcluvia D. Don. 11 spp., 10 in Australasia and C. paniculata (Cav.) D. Don in central Chile and Argentina; forest; remaining in Australasia.

1.4 TRIBE CUNONIEAE (4/210-220) ‣ outsiders Vesselowskya (2; Queensland, New South Wales), Pterophylla (68, Madagascar, Comoros, Malesia, Pacific Islands), Pancheria (c 30; New Caledonia), Cunonia (c 25; New Caledonia, one species in W and E Cape and KwaZulu-Natal).

4. Weinmannia L. 91 spp., two in Mascarenes Islands and 89 in New World, from Mexico to Cono Sur, Caribbean, South America (83), with the greatest concentration of species in Colombia (35), Ecuador (30) and Peru (43), mostly in forest, especially montane tropical forest and subtropical forest, extending into temperate forest in the south, often a significant component of cloud forests in the Andes where several species can co-occur at one locality or in a single altitudinal zone; absent from the lowlands of the Amazon rainforest; 5 spp. in Brazil, three endemics, W. pinnata L., a very variable and widely distributed tree, ranging from S Mexico to Brazil, and W. organensis Gardner also in Peru.

ELAEOCARPACEAE

§ FULLY CHLOROTROPHIC

Genera/species 12/c. 550 Distribution Mexico, Central America, Caribbean, South America southwards to southern Chile, Madagascar, Mauritius, Socotra, E Himalaya, E and S India, Sri Lanka, E Asia to Japan, SE Asia, Malesia, New Guinea, Melanesia, Samoa, Tonga, and other Pacific islands, Australia, Tasmania, New Zealand. Habit usually bisexual (rarely monoecious or dioecious), evergreen trees, shrubs or suffrutices; a large number of species are xerophytes.

Trees, shrubs or herbs. Trunks sometimes with buttresses. Sloanea is only genus without petals. All taxa listed are native. The genus Elaeocarpus L. is cultivated in Brazil. Two genera and 41 spp. in Brazil. The fruits of Sloanea are armed or smooth woody capsules; in Crinodendron the fruits are dehiscent and winged, and in Vallea they are fleshy and berry-like.

Crayn et al. (2006) demosntrated that Sloanea is a monophyletic sister-group of the clade formed by Vallea and Aristotelia. In the same study, Crinodendron was also considered as monophyletic, and forming a sister-group with Peripentadenia.

SYSTEMATIC two tribes, both in South America.

1.1 TRIBE ELAEOCARPEAE (9/c. 435) ‣ outsiders Peripentadenia (2; Queensland), Dubouzetia (11; Moluccas, New Guinea, Northern Territory, Queensland, New Caledonia), Tremandra (2; W Australia), Platytheca (2; W Australia), Tetratheca (c 40; W Australia, S Australia to Queensland, Tasmania), Elaeocarpus (c 350; Madagascar, tropical and subtropical Asia and eastwards to New Caledonia, New Zealand and Hawaii), Sericolea (16; E Malesia), Aceratium (c 20; E Malesia and eastwards to Queensland, Solomon Islands and Vanuatu).

1. Crinodendron Molina. 4 spp., two endemics to in Chile, C. tucumanum Lillo in Bolivia and Argentina, and C. brasiliense Reitz & L.B. Sm. in south Brazil, a shrub with white flowers, very narrow endemic of Bom Retiro municipality, Santa Catarina state, in nebular forests.

1.2 TRIBE SLOANEEAE (3/165) ‣ all genera occur in South America.

2. Aristoelia L’ Her. 5 spp., A. chilensis (Molina) Stuntz from Chile, remaining 4 in New Zealand, Tasmania and New South Wales.

3. Sloanea L. Small to large trees up to 40 m tall, often butteressed; occasionally shrubby; leaves sometimes subopposute; inflorescence cymose, thyrsoid, a fascicle or raceme, or flowers solitary; fruit woody. 150 spp. in Central and South America (123), 70 in Old World, Madagascar (3), NE India & Nepal (4), Burma (7), China (12-15), Vietnam (6), Borneo (2), New Guinea (18), New Caledonia (9) and Australia (4); 48 spp. in Brazil (16 endemics), widely distributed in country. One undeterminated sp. fom tropical America is myrmecophyte.

4. Vallea Mutis ex L.f. Shrubs or trees, pink flowered. Two spp. in Venezuela, Colombia, Ecuador, Peru, Bolivia and Argentina, in the lower slopes of Andes.

36. MALPIGHIALES

FAMILIES ABSENTS IN SOUTH AMERICA: BALANOPACEAE (1/9), CENTROPLACACEAE (2/9), CTENOLOPHONACEAE (1/2), IRVINGIACEAE (4/12), LOPHOPYXIDACEAE (1/1), PANDACEAE (3/17) AND RAFFLESIACEAE (3/49).

LINEAGE 1 de 6: RHIZOPHORIDS

RHIZOPHORACEAE

§ FULLY CHLOROTROPHIC

Genera/species 15/150–160 Distribution Pantropical, Pacific and Atlantic coasts of Central and South America, the Caribbean, Atlantic and eastern coasts of Africa, Pacific and Indian Ocean islands, with their highest diversity in Madagascar and tropical Asia. Habits Usually bisexual (rarely polygamomonoecious), evergreen trees or shrubs. Many species are mangrove trees. Pneumatophores (in Gynotrocheae and Rhizophoreae) and stilt roots often present. The family consists of 14 genera and about one hundred species distributed around the tropics. Three genera occur in Amazon rainforest, Rhizophora that is confined to coastal mangrove forest.

SYSTEMATIC three tribes; Gynotrocheae (4/30–32, Madagascar, tropical Asia to tropical Australia, islands in W Pacific) does not occur in New World.

Key to genera of Neotropical Rhizophoraceae

1. Ovary inferior; fruit a viviparous drupe; plants with stilt roots ------------ Rhizophora

1. Ovary superior; fruit a capsule, plants without stilt roots - 2

2. Flowers hermaphrodite; leaves opposite; petals fimbriate or laciniate; inflorescence of solitary flowers or sessile clusters ------------ Cassipourea
2. Flowers dioecious; leaves verticellate; petals laciniate; inflorescence a corymobose panicle with long peduncle ------------ Sterigmapetalum

1. TRIBE MACARISIEAE (8/105–115) ‣ outsiders Macarisia (8; Madagascar), Anopyxis (3; tropical Africa), Blepharistemma (1; SW India), Comiphyton (1; Gabon to E Congo).

1. Paradrypetes Kuhlm. Dioecious trees; leaves opposite, entire or more often serrate with few to many spinose teeth, without imbedded glands; inflorescences epipetiolar (the leaf blade sometimes aborting and leaving only a scar), ebracteate, cymose (dichasial); staminate flowers numerous; fruit a glabrous drupe. Two spp., P. ilicifolia Kuhlman, along rivers in the Atlantic rain forest of Brazil, in the states of Espírito Santo and Minas Gerais, and P. subintegrifolia G.A. Levin, from Iaco and Tarauaca drainages of SW Amazonas and Acre, Brazil, and Ecuador (Napo region), in varzea forest.

2. Cassipourea Aubl. Trees or shrubs, leaves opposite and decussate; inflorescences solitary flowers or fasciculate groups, axillary, sessile or pedicellate; flowers hermaphrodite; disk fleshy or membraneous, dentate; calyx tube campanulate, 4-5-lobed, the lobes erect, valvate; petals 4-5, unguiculate, fimbriate, folded in bud, white; fruit an ovoid capsule. 70 spp., pantropical, mainly African, 16 from Belize to Bolivia, 10 in South America, three of which occur in Amazon rainforest and two in central Amazon rainforest; 3 spp. in Brazil, one endemic.

3. Sterigmapetalum Kuhlm. Small to large trees; leaves 2-5 verticellate; stipules interpetiolar or axillary, coriaceous; inflorescence of pedunculate, corymbose, dichotomous panicles, with bracts and bracteoles; flowers dioecious; calyx 6-7 lobed in female flowers, 5-6 lobed in male flowers; fruit an obovate-oblong capsule, 5-6 locular with 2 seeds in each locules, septicidal. 8 spp., in Colombia, Guianas and Venezuela (mainly species), and central Amazon rainforest west to Iquitos, Peru; only two in Brazil, none endemics.

2. TRIBE RHIZOPHOREAE (4/18–19) ‣ outsiders 4/19. Bruguiera (6; coasts of tropical East Africa and east to Samoa), Kandelia (2; coasts of India and Bangladesh to Borneo, China and Kyushu in S Japan), Ceriops (5; coasts of East Africa and India to Queensland, Melanesia, Micronesia and N to SE China).

4. Rhizophora L. Mangroves, with rhizophores. 8-9 spp. and 4 hybrids, two species and one hybrid in New World: R. mangle L. from Florida to southern Brazil, C Mexico to northern Peru, Fiji, New Caledonia, Tonga, Samoa, and American Samoa - Bermuda is the most northerly extent of its range; presence on the mid-Atlantic islands (St. Helena and Ascension) has been reported for this species, but this is not confirme; R. racemosa G.Mey occur from Costa Rica to Maranhão state in northern Brazil in Atlantic coast, and Costa Rica to Pacific Coast of Colombia, also W Africa Guinea Gulf, from S Mauritania to Angola; and the hybrid Rhizophora × harrisonii Leechm. from Mexico, Central America, Trinidad & Tobago, Venezuela, Guianas, Colombia, Ecuador, and Brazil (only Pará and Maranhão states).

ERYTHROXYLACEAE

§ FULLY CHLOROTROPHIC

Genera/species 4/c. 260 Distribution pantropical (especially the Andes, Amazonas and Madagascar), southwards to C Chile and Argentina, South Africa and eastern India. Habit usually bisexual (rarely dioecious), evergreen small trees or shrubs. Branches often covered with distichous scale-like rudimentary leaves. Buds perulate.

Recent molecular studies, supported by various morphological and anatomic characters, have shown affinity between Erythroxylaceae and Rhizophoraceae, and suggest that they belong to the order Malpighiales, sensu APG I (1998); in the classification proposed by the APG II (2003), the large affinity and the set of characters shared by both families led Erythroxylaceae and Rhizophoraceae to be considered, optionally, as a single Family; the main characters shared by the two families are: alkaloids from the tropane and pyrrolidine groups, the presence of colleters, terminal buds protected by stipules and green embryos.

Use Medicinal plants, cocaine (Erythroxylon coca Lam., E. novogranatense (D. Morris) Hieron.), timber, tar, dyeing substances.

SYSTEMATIC outsiders all in Africa: Aneulophus (2; tropical W and C Africa), Nectaropetalum (5; tropical and southern Africa, Madagascar), Pinacopodium (2; tropical Africa).

1. Erythroxylum P.Browne. Glabrous shrubs or small trees, sometimes with woody rhizomes or xylopodium; branchlets erect to spreading; leaves simple, entire, alternate, pinnately veined; inflorescences fasciculate at nodes, sometimes short-pedunculate, with 1-many flowers; small, actinomorphic, bisexual or unisexual, calyx lobes 5, persistent, valvate, sepals united below; petals 5, free, alternate with sepals, imbricate in bud; fruits small, drupaceous and one-seeded. Seeds with straight embryo, with or without endosperm. c. 230 spp., 212 spp. native in New World, from Mexico, Mesoamerica, Caribbean, South America (182); 136 spp. in Brazil, 92 endemics, highly centered in Bahia state with 25 spp. 7 spp. in seven states are rare plants in Brazil, by Plantas Raras do Brasil’s book.

Some native species of Erythroxylum, referred to in the literature as having pharmacological potential because they contain alkaloids, flavonoids and terpenoids, are widely used in medicine; among these is E. coca Lam., from which the cocaine alkaloid is extracted. It has been used since 1879 as a local anaesthetic and for terminally ill patients (Brompton cocktail), but mainly illegally as a recreational drug sold in major urban centers to a considerable addict population; this species has long been used as a ritualistic psychoactive medicinal plant by a host of South American tribes. In Brazil, E. vacciniifolium Mart., popularly known as catuaba, is used to stimulate the central nervous system in addition to exhibiting aphrodisiac properties; E. pelleterianum A. St.-Hil. is used to treat stomach pains; E. myrsinites Mart. and E. suberosum A. St.-Hil. are used in the tanning industry.

LINEAGE 2 de 6: OCHNOIDS

OCHNACEAE

§ FULLY CHLOROTROPHIC

Genera/species 36/565–575 Distribution pantropical (few subtropical species), mainly in Central America, the Caribbean and tropical South America, with their largest diversity in Amazon rainforest. Habit usually bisexual (sometimes polygamomonoecious, dioecious, androdioecious, or polygamodioecious), evergreen trees, shrubs or lianas, rarely perennial herbs. Almost all Ochnaceae are evergreen shrubs or small trees; none genera in New World are tall trees, and herbs occur only in Sauvagesia.

SYSTEMATIC three subfamilies, Medusagynoideae (1/1, Mahé in Seychelles) does not occur in South America.

1. SUBFAMILY QUIINOIDEAE (4/50–55) ‣ all genera occur in South America.

1. Froesia Pires. Shrubs or trees, unbranched stems bears a rosette of huge leaves and is terminated by inflorescences. 5 spp. in northern South America up to Peru, 3 in Brazil, F. crassiflora Pires & Fróes endemic, a rare plant in Brazil, by Plantas Raras do Brasil’s book.

2. Lacunaria Ducke. Shrubs or medium-sized trees with a more or less regular branching pattern; leaves of adult plants simple; leaves are simple and disposed in whorls of (3–)4(–6); inflorescences are bracteate and mostly axillary, sometimes terminal; flowers actinomorphic, small, and subglobose in bud; petal color white or yellow; fruits are berry-like and possess a fibrous pericarp with longitudinal large ‘resiniferous’ lacunae. 7 spp. from northern South America, L. crenata (Tul.) A.C. Sm. widely distributed up to Nicaragua and Atlantic Forest of southern Brazil (where all species occur, two endemics, both rare plants in Brazil, by Plantas Raras do Brasil’s book) and Bolivia, mostly in Amazon rainforests.

3. Touroulia Aubl. Trees; leaves of adult plants compound. Two spp. restricted to northern South America (except Ecuador and Peru), mostly in the Amazon forests, both in Brazil, none endemics.

4. Quiina Aubl. Trees; leaves of adult plants simple. 32 spp., one in Jamaica, and 31 in mainland (all in South America), from Belize and Jamaica to Bolivia and southern Brazil (22, 8 endemics), moist in the Amazon rainforest.

2. SUBFAMILY OCHNOIDEAE (24/515–520) - four lineages; Testulea clade (1/1, Gabon) does not occur in South America.

2.1 OCHNOIDEAE ▸ TRIBE LUXEMBURGIOIDEAE (2/23) - both genera ocur in South America.

5. Luxemburgia A. St.-Hil. 20 spp., mainly in Espinhaço Range in Minas Gerais state; a few spp. in Goiás, Bahia, and in granitic outcrops in Espírito Santo and Rio de Janeiro states, all above 1,000 m elevation range; eight species (seven in Minas Gerais and one in Bahia state) are rare plants in Brazil, by Plantas Raras do Brasil’s book.

6. Philacra Dwyer. 4 spp., all in the Guiana Shield of Venezuela, two up to Amazonas state in northern Brazil, 800-2,000 m elevation range.

2.2. OCHNOIDEAE ▸ TRIBE SAUVAGESIOIDEAE (16/c 85) - outsiders Neckia (1; W Malesia, Philippines), Schuurmansia (18; C Malesia to New Guinea), Schuurmansiella (1; NW Borneo), Euthemis (2; SE Asia to Borneo), Indosinia (1; southern Vietnam).

7. Adenarake Maguire & Wurdack. Two spp., endemic to Pantepui Life Zone, Mount Neblina in Venenzuela and Brazil (only A. muriculata Maguire & Wurdack, no endemic) up to Guyana, 1,500-2,800 m elevation range.

8. Blastemanthus Planch. Two spp., endemic to the Guiana Shield of Colombia to Guyana and northern Brazil (all species, two endemics), 100-200 m elevation range.

9. Cespedesia Goudot.Only one sp., C. spathulata (Ruiz. & Pav.) Planch., Central America, Venezuela, Colombia, Ecuador, Peru, Bolivia and Brazil.

10. Godoya Ruiz & Pav.Two spp., G. antioquiensis Planch. and G. obovata Ruiz & Pav., Colombia to Bolivia.

11. Krukoviella A.C.Sm.Only one sp., K. disticha (Tieghem) Dwyer, western Amazon rainforest, in Brazil (Amazonas state), Peru and Ecuador (Zamora-Chinchipe).

12. Poecilandra Tul.Two spp., restricteds for to the Guiana Shield of Colombia to Guyana and Brazil (only P. retusa Tul., non endemic), 500-2,800 m elevation range.

13. Rhytidanthera (Planch.) Tieghem. Only genus of Ochnaceae with compound leaves. Only one sp., R. splendida (Planch.) Tiegh., in small populations restricted to the primary forest of the Colombian and Venezuelan Andes and the sandstone hills of La Macarena and Chiribiquete.

14. Sauvagesia L. Herbs, perennial or annual, completely glabrous, with erect shoots and frequently elongate-ascending branches from base; leaves blade scale,acicular, bristle or latifolius, in some species in end of the branches; with actinomorphic flowers with pink or white petals; inflorescence paniculate to racemose and terminal or reduced to solitary or rarely paired flowers in axils of foliage leaves. 47 spp., all in South America (37 in Brazil, 20 endemics; 4 spp., 3 in Bahia state and one in Goiás state are rare plants in Brazil, by Plantas Raras do Brasil’s book), 5 up to Mexico, Central America and Caribbean, inc. S. erecta L., with a worldwide distribution; most species either endemic to white-sand forests and remnant areas of the Guiana Shield in Venezuela, Colombia, Brazil and Guyana, or to campos along the Espinhaço Range in Minas Gerais and Bahia, Brazil. Two sections:

§ sect. Sauvagesia ‣ two subsections:

§ subsect. Sauvagesia ‣ comprises the type, S. erecta L., and allies, including representatives of the former separate genera Pentaspatella, Roraimanthus and Leitgebia; this clade is sister to Adenarake.

§ subsect. Vellozianae ‣ some species of the formerly independent Lavradia (S. capillaris (A.St.-Hil.) Sastre, S. glandulosa (A.St.-Hil.) Sastre), one of many spectacular radiations in the rocky grasslands (campos rupestres) of the Espinhaço Range, including the monocot genera Vellozia (Velloziaceae) and Leiothrix (Eriocaulaceae), the Asclepiadoideae genera Hemipogon and Minaria (Apocynaceae), Merianthera (Melastomataceae), Oocephalus (Lamiaceae), Richterago (Asteraceae), and the legumes Calliandra sect. Monticola and Baseophyllum clade of Chamaecrista.

§ sect. Imthurnianae ‣ 3 spp., Colombia to Guinanas and N Brazil (only one).

Sastre's (1978, 1981) infrageneric classification of Sauvagesia might provide a solution to the issue of the two clades with a nested Adenarake. Thus, Adenarake might be kept as a distinct genus if clade A is established as a different genus (e.g., as Lavradia). However, without a comprehensive taxonomic treatment of the genus at hand, it is beyond the scope of the present study to judge if this is a sound solution to the problem or whether a broad concept is to be preferred, including both clades plus Adenarake in Sauvagesia.

15. Tyleria Gleason, Steyerm. & Wurdack (inc. Adenanthe). 13 spp., all endemic to the Guiana Shield of Venezuela but one up to Guyana and two up to northern Brazil (only Amazonas state), 1,200-2,500 m elevation range.

16. Wallacea Spruce ex Benth. & Hook.f. Two spp. endemic to the Guiana Shield of white-sand forests in SE Colombia, southern Venezuela and northern Brazil (both species in all three countries), 100-200 m elevation range.

2.3 OCHNOIDEAE ▸ TRIBE OCHNOIDEAE (9/c 385) - outsiders Lophira (2; tropical W and C Africa), Campylospermum (c 65; tropical C and E Africa, Madagascar, tropical Asia), Idertia (2; W and C tropical Africa, São Tomé), Brackenridgea (10; tropical regions in the Old World), Rhabdophyllum (4; W and C tropical Africa), Ochna (c 85; tropical regions in the Old World).

17. Elvasia DC. 14 spp., 10 in Brazil, 4 endemics (three of then, all in Espírito Santo state, are rare plants in Brazil, by Plantas Raras do Brasil’s book); two subgenera:

§ subg. Elvasia ‣ 4-5 spp., star-shaped fruits and a distribution concentrated in the Amazon rainforest.

§ subg. Hostmannia ‣ 9-10 spp., globular fruits and a circum-Amazonian distribution, one up to Central America; E. kollmannii Fraga & M.M. Saavedra has elliptic-coriaceous leaves, with acute to mucronate apex, and flowers with white petals, a unique character in Elvasia; E. gigantifolia Fraga & M.M. Saavedra has the largest leaves in Elvasia, up 35 cm long.

18. Ouratea Aubl. Shrubs or small trees, remarkable leaves with the secondary veins strongly curving upwards near the usually serrate margin and continuing almost as submarginal veins, with pseudoapocarpic gynoecium with a gynobasic style and a gynophore that develops, in the ripe fruit, into a reddish carpophore that displays black mericarps derived from fertilized 1-ovulate units, and mainly yellow flowers. 273 spp. in Central and South America (238); 127 spp. in Brazil, 88 endemic (4, in Tocantins, Bahia and Minas Gerais states, are rare plants in Brazil, by Plantas Raras do Brasil’s book).

O. cauliflora Fraga & Saavedra, narrow endemic to Espírito Santo state is the only cauliflorous species in this genus, and one by two of white flowers in this genus together O. cassinefolia (DC.) Engl., endemic to Pará and Maranhão states.

19. Perissocarpa Steyerm. & Maguire. Three spp., one endemic to at high-elevation areas in the Andes in Peru, and two remaining from Coastal Cordillera of Venezuela, and the western portion of the Guiana Shield in Amazonas state of northern Brazil (both species, none endemics), Guyana and Venezuela.

BONNETIACEAE

§ FULLY CHLOROTROPHIC

Genera/species 3/c. 35 Distribution Archytaea and Bonnetia: northern South America, the Caribbean; Ploiarium: Cambodia, W Malesia, Moluccas, New Guinea. Habit bisexual, evergreen subpachycaul trees or shrubs. Use ornamental plants. Three genera and about 40 spp., northern South America, Caribbean, SE Asia, W Malesia, Moluccas and New Guinea. Two genera in New World, all endemic.

SYSTEMATIC outsider Ploiarium (5; Cambodia, W Malesia, Halmahera, New Guinea).

1. Archytaea Mart. Two spp., endemic to the Guiana Shield region and adjacent lowlands in Colombia, Venezuela, Brazil (only A. triflora Mart.) and Guyana, ranging from 50-2,000 m elevation range.

2. Bonnetia Mart. Trees or shrubs. 30 spp., 28 in Guiana Shield (Colombia, Brazil (5), Venezuela (19 endemics), one up to along the Andes to Peru, B. stricta (Nees) Nees & Mart. from rocky grasslands (campos rupestres) of Bahia to Atlantic sandy coastal shrublands (restingas) of Rio de Janeiro state, and B. cubensis (Britton) Howard in Cuba and Porto Rico.

CLUSIACEAE

§ FULLY CHLOROTROPHIC

Genera/species 18/860–870 Distribution pantropical. Habit usually bisexual (rarely polygamomonoecious, in e.g. Clusia and Garcinia also dioecious), evergreen trees or shrubs (in Clusia sometimes lianas or epiphytes, also with CAM physiology).

The family occurs in most Neotropical habitats, from sea level to 3,500 m altitude (spp. of Clusia). Diversity of genera is highest on the Guiana Shield and surrounding areas whereas species diversity is highest in and near the northern Andes. The Caribbean are poor in species. Most species are found in rainforests but species of Clusia and Garcinia also occur in dry, open habitats. Pollination mechanisms are highly diverse and pollinators include bees, wasps, perching birds, bats, flies, beetles, moths, cockroaches and perhaps small arboreal mammals. Several species are cultivated as fruit trees, e.g. Platonia insignis Mart. (known as bacuri) and Garcinia spp. The latex has several traditional technical and medicinal uses, and contains compounds with antibiotic and antitumor activity.

SYSTEMATIC three tribes, all in South America.

1. TRIBE CLUSIEAE (6/c. 430) ‣ all genera occur in South America.

1. Arawakia L. Marinho. (off Tovomita) Trees or shrubs; exudate white; leaves clustered at the apex of the branches; leaf blades fleshy; inflorescence terminal; flowers with buds enclosed by the outer sepals; petals white to yellowish; fruit a capsules purplish-red when mature; petals and staminodes caducous, sepals persistent and adpressed to the fruit; seeds one per locule, arillate, the aril orange. 18 spp., Nicaragua to Bolivia, usually in highlands in the Andes and the Guiana Shield (mainly elevations from 100-1,700 m), although some species reach lowlands in Central America and Colombia, only the widely distributed A. weddelliana (Planch. & Triana) L. Marinho in Brazil, known only in Amazonas state.

With at least 4 species restricted of Central America, South America has a maximum of 14 spp.

2. Chrysochlamys Poepp. & Endl. Shrubs or trees; latex o ten milky white or yellow. 38 spp. of Mexico, Central America (9 restricted, one shared with South America), Lesser Antilles, northern South America (26); only 5 in Brazil, none endemics.

3. Clusia L. (inc. Havetiopsis, Oedematopus, Quapoya) Hemiepiphytes, lianas, small trees and shrubs with fleshy capsules and seed less than 6 mm long with an orange aril; flowers often with resin; their leaves are reference because their curious venation. 296 spp., all Neotropical, 257 in South America, 74 in Brazil, 28 endemics.

Clusia L. (Clusiaceae), Clusiella Planch & Triana (Calophyllaceae) and Dalechampia Plum. ex L. (Euphorbiaceae) are the only known genera to offer resin as a reward for some groups of bees that use it in nest construction.

4. Dystovomita (Engl.)D’Arcy. (exc. Tovomita p.p.). Two spp. from Nicaragua to Colombia, Ecuador, Guyana, Peru, and Venezuela, both in South America.

5. Tovomita Aubl. (inc. Tovomitidium, Dystovomita p.p., exc. Arawakia) Trees or shrubs, sometimes hemiepiphytics, glabrous the latex often white or yellowish. 76 spp., of South and Central America, 75 in South America (one restricted for Caribbean), 43 in Brazil, 18 endemics, typically found in lowland rainforests and lower montane forests, very rarely in in other environments, as white sand forests of Peru and Colombia.

6. Tovomitopsis Planch. & Triana. Two spp., restricted of SE Brazil, from Minas Gerais to Paraná state.

2. TRIBE GARCINIEAE (1/100-300) - only one genus.

7. Garcinia L. (inc. Rheedia) Shrubs or trees, generally glabrous. 405 spp., pantropical, many species has sweet edible fruits; 14 sections, only one occur in New World, sect. Rheedia, with c.40 spp. from New World and Madagascar; 37 spp. in New World, 18 in South America, 7 in Brazil, one endemic.

3. TRIBE SYMPHONIEAE (7/50-52) ‣ outsiders Pentadesma (c 15; tropical Africa), Montrouziera (5; New Caledonia).

8. Lorostemon Sw. Largest trees, bark with latex white to yellow. 4 spp. from Guyana to Colombia and northern Brazil (4 (one endemic), in Amazonas and Pará states).

9. Moronobea Aubl. Medium to large trees, glabrous; bark with latex yellow; edible fruits. 7 spp., endemic to the Guiana Shield of Veneuela, Guianas, Colombia and northern Brazil (5, two endemics, M. pulchra Ducke, from Amazonas state, is a rare plant in Brazil, by Plantas Raras do Brasil’s book), except by M. coccinea Aubl., which occurs in central Amazon rainforest of Brazil up to Bolivia.

10. Platonia Raf. Large tree to 40 m tall, fruit edible with a tart but sweet. Only one sp., P. insignis Mart., endemic to the Guiana Shield in Colombia, Venezuela, Guyana, Suriname, French Guiana and northern Brazil, from Roraima and Amazonas to N Piaui state.

11. Symphonia L. Medium or large trees, with a straight bole and a crown of short horizontal branches; leaves opposite, petiolate, entire, ± coriaceous; inflorescence a terminal, one to many-flowered, corymbose or umbellate cyme; flowers bisexual, pedicellate; fruit a 1–3-seeded berry with tough epidermis; seeds large, with a fibrous aril. c. 16 spp., all confined to Madagascar except S. globulifera L.f., found in Africa and from Mexico to Bolivia, Brazil and Caribbean; this species yields a resin of economic importance.

12. Thysanostemon Maguire. Two spp., endemics to the Guiana Shield in Guyana, 900-1,000 m elevation range

CALOPHYLLACEAE

§ FULLY CHLOROTROPHIC

Genera/species 13/c. 480 Distribution pantropical. Habit usually bisexual (occasionally cryptic-dioecious, rarely andromonoecious), usually evergreen trees (sometimes shrubs or epiphytes). The family comprises some important timber trees, particularly Calophyllum spp. Several species are cultivated as fruit trees, e.g. Mammea americana L. Wood of Calophyllum brasiliense Cambess., known as guanandi in Brazil, that is very used in Brazil by the traditional communities. The wood is ideal for the production of canoes, masts for ships, beams, for civil construction, internal works, floors, woodwork and carpentry.

Use Ornamental plants, fruits (Mammea americana L. etc), perfumes (Mammea siamensis T.Anders.), medicinal plants, cosmetics, dyeing substances, seed oils, timber.

SYSTEMATIC: two clades; Endodesmia Clade (2/2, W Africa) does not occur in South America; among Calophylleae, three tribes, all in South America.

1. CALOPHYLLUM GROUP (6/290) ‣ outsiders are Mesua (c 40; tropical Asia), Mammea (c 50; tropical regions on both hemispheres), Kayea (7; SE Asia, Malesia to New Guinea), Agasthiyamalaia (1; W Ghats), Poeciloneuron (2; S India).

1. Calophyllum L. Trees up to 40 m tall, rarely shrubs, glabrescent, buds finely tomentose; latex sulfur yellow with a fain-greenish tint; flowers bisexual, terminal or axillary, in few- to many-flowered racemes or paniculate cymes or rarely reduced to 1–3. ca. 180 spp., most abundant in tropical Asia and Australasia, but also occurring in Madagascar, Mascarene Islands, East Africa, 10 in New World, a half in Caribbean, another half from Mexico to Bolivia, southern Brazil, and Guianas, all in Brazil, C. angulare A.C. Sm. endemic.

In 1810, the imperial government reserved to the state the monopoly of exploration of C. brasiliense Cambess wood for use exclusively in the production of masts and crossjacks for ships, and, therefore, it is the country’s first hardwood (law of January 7th, 1835).

2. KIELMEYERA GROUP (3/70) ‣ all genera occur in South America.

2. Caraipa Aubl. Shrubs or trees with sparse latex, alternate leaves, or shrubs in tepuis. 43 spp., Colombia to French Guiana, Peru, and northern and NE Brazil (26, 12 endemics), from terra-firme forest, flooded forest, seasonally wet forest, and white-sand vegetation to the top of mountains and tepuis, rare in Andes; two spp. from Mount Aracá are rare plants in Brazil, by Plantas Raras do Brasil’s book.

3. Haploclathra Benth. 4 spp. from Amazon rainforest of Brazilian (3, one endemic), Guianas and Peru (2).

4. Kielmeyera Mart. & Zucc. Trees, shrubs or subshrubs, sometimes with xylopodium; glabrous or with simple, bifid or dendritic trichomes and with spiraled alternate leaves; inflorescences are terminal, cymose, with monoclinous or staminate flowers, a pentamerous perianth, white or pinkish corolla, and usually asymmetric petals; fruits are septicidal woody capsules with numerous winged seeds. 56 spp., five in Brazil up to Bolivia, K. coriacea Mart. & Zucc. in Brazil, Bolívia and Paraguay, Peru and Bolivia one endemic each, and remaining 48 endemics to Brazil (11, in several states, are rare plants in Brazil, by Plantas Raras do Brasil’s book).

3. MAHUREA GROUP (4/26) ‣ all genera occur in South America.

5. Clusiella Planch. & Triana. Slender epiphytic lianas or shrubs. 8 spp., 7 in northern South America (one only Central America), mainly in Colombia and Ecuador; from Costa Rica to Peru, Venezuela, Guyana and northern Brazil (two spp., Amazonas, Acre and Pará, none endemics).

6. Mahurea Aubl. Shrubs to medium sized trees; wood reddish. Two spp., M. exstipulata Benth. of Guyana to Colombia, northern Brazil and Peru, M. palustris Aubl. from coastal French Guiana and adjacent Brazil.

7. Marila Sw. Shrubs or trees; leaves opposite; latex white, brown, yellow, or clear. 19 spp. from Mexico to Bolivia, 15 in South America, only two spp. in Brazil, none endemics.

8. Neotatea Maguire. Shrubs or small trees with few, thick branches. Three spp. endemics to the Guiana Shield in Colombia and Venezuela, and possibly in Brazil (Mount Neblina), 200-2,000 m elevation range.

HYPERICACEAE

§ FULLY CHLOROTROPHIC

Genera/species 6/c. 600 Distribution cosmopolitan except polar areas. Habit bisexual, evergreen or deciduous trees, shrubs or herbs. Species in dry areas sometimes with lignotuber. Cosmopolitan family with four neotropical genera, two restricted from montane of south Mexico and Guatemala; Hypericum is most non-lowland habitats, Vismia prefer this regions.

SYSTEMATIC three tribes, Cratoxyleae (2/7, Madagascar, Burma, S China, SE Asia, Malesia to the Lesser Sunda Islands) does not in South America.

1. TRIBE HYPERICEAE (1/370) ‣ only one genus.

1. Hypericum L. Shrubs or perennial (some annual) herbs, plants glabrous, with pellucid plands, sometimes with roots crown; mainly yellow flowers. 430 spp., nearly cosmopolitan, 193 in New World, with main centers of diversity in Eurasia (more than 230) and South America (107, c. 70 in paramos in Andes), and smaller ones in North America (c. 40), SE Asia (c. 47), and Africa (c. 30), mostly temperate areas, as Argentina; 24 in Brazil, nine endemics, H. pedersenii N.Robson, from Rio Grande do Sul state, is a rare plant in Brazil, by Plantas Raras do Brasil’s book; Hypericum has two subgenera:

§ subg. Brathys - three sections:

§ sect. Brathys ‣ 88 spp., Andes S to Peru & Bolivia, C & E North America.

§ sect. Myriandra ‣ 30 spp., E North America, Mexico, Greater Antilles, Bahamas, Bermuda.

§ sect. Trigynobrathys ‣ 59 spp., New World, E & S Asia, Australia, New Zealand, Hawaii, tropical & S Africa.

§ subg. Hypericum ‣ 27 sections, Old World and six from North America up to Central America, one endemic to Mexico.

Although predominately a high-elevation group in the Neotropics, a small number of herbaceous Hypericum species has adapted to lower elevations, distributed below 3,000 to less than 1,000 m in lowland regions of temperate South America.

2. TRIBE VISMIEAE (3/110) ‣ outsiders Harungana (1; tropical Africa, Madagascar, Mauritius), Psorospermum (40–45; tropical Africa, Madagascar).

2. Vismia Vand. Small trees, shrubs or lianas; leaves opposite, branches near spreading, with simple trichomes, ferruginous; yellow to orande latex, darkening to reddish on exposure; inflorescences terminal or axillary cymes, cymose panicles or glomerules; flowers bisexual, regular. 61 spp. in two subgenera:

§ subg. Vismia ‣ 54 spp., Mexico to Brazil, 53 in South America, 30 in Brazil, 9 endemics.

§ subg. Afrovismia ‣ 7 spp. in Africa.

PODOSTEMACEAE

§ FULLY CHLOROTROPHIC

Genera/species 52/295–300 Distribution tropical, subtropical and warm-temperate regions in the Southern and Northern Hemispheres, with their largest diversity in Central America and tropical South America. Habit annual or perennial, aquatic herbs, often bizarre in form, sometimes resembling lichens, bryophytes, seaweeds, or unlike any other plants; Leaves with variable size and shape, sessile or petiolate, generally with sheath, linear, lanceolate, palmate, reniform, cordate, obovate, reduced to capillary filaments, or thalloid, margin entire or dichotomically subdivided; uninerved, penninerved ou palminerved, surface smooth, papilose or spinulose, phyllotaxy opposite, spiraled, distichous, tristichous, rosulate or fasciculate. haptophytes, attached by adhesive hairs to rock or other hard objects in flowing freshwater, mostly in rapids and waterfalls; roots usually photosynthetic, creeping or partly floating, thread-like, ribbon-shaped, crustose (foliose), sometimes short-lived or absent. Shoots nearly always arising as endogenous buds from roots; stems reduced or elongate, simple or branched, sometimes dimorphic, occasionally only present when flowering.

The interpretation of the vegetative body is controversial; many Podostemaceae have a flattened photosynthetic body which adheres to a hard substrate; it has been called a ‘thallus’ because the conventional demarcation into stem, leaf and root is usually not obvious, and various botanists have denied or doubted the homology of this vegetative body with stems (caulomes), leaves (phyllomes) and roots of other angiosperms. In the Neotropics (20 genera and 150 spp.) it ranges from C Mexico to northern Paraguay and Argentina .

SYSTEMATIC three subfamilies, all in South America, one restricted.

1. SUBFAMILY TRISTICHOIDEAE (6/15–16) ‣ outsiders Indotristicha (3; India), Dalzellia (6; Sri Lanka to S China and SE Asia), Indodalzellia (1; S India), Cussetia (2; Thailand, Cambodia, Laos; possibly extinct); Terniopsis (1–5; S China, Thailand, the Malay Peninsula).

1. Tristicha Touars. (inc. Heterotristicha) Herbs with branching roots, adhered to the substrate; branches prostrate, attached or floating, 0.2–15 cm long; leaves with phyllotaxy tristichous, laxly or densely arranged along the stem, membranous, sessile, entire, rarely lobed; flowers axillary or terminal, solitary or not, surrounded by membranous leaves. Only one polymorphic sp., T. trifaria (Bory ex Willd.) Spreng., widely distributed in tropical and subtropical America (Nicaragua to Colombia, Brazil, Guianas, Argentina and Paraguay), Africa and Asia; possibly two different species.

2. SUBFAMILY WEDDELLINOIDEAE (1/1) ‣ only one genus.

2. Weddellina (Warm.) Engl. Plant scaly; flowers terminal, solitary. Only one sp., W. squamulosa Tul., northern to C South America in Colombia to Suriname and northern Brazil, 100-400 m elevation range.

3. SUBFAMILY PODOSTEMOIDEAE (50/278) ‣ two subgroups, one monotypic, another the core.

3.1 PODOSTEMOIDEAE ▸ DIAMANTINA CLADE (1/1) - a single genus.

3. Diamantina Novelo, C.T. Philbrick & Irgang. Roots prostrate, elongate, dorsiventrally flattened, sometimes intertwined, green and photosynthetic, branched, with asymmetric root cap, attached to solid substrata via holdfasts (haptera) and adhesive hairs; unique Podostemaceae in New World with digitate leaves. Only one sp., D. lombardii Novelo, C.T. Philbrick & Irgang, endemic to the state of Minas Gerais, in the basin of the Rio Preto, and is known from only two localities: Rio Preto (Parque Estadual do Rio Preto) and Rio do Peixe.

3.2 PODOSTEMOIDEAE ▸ CORE PODOSTEMOIDEAE (49/287) - outisiders Noveloa (2; Mexico, Central America), Inversodicraea (4; tropical Africa), Monandriella (1; Cameroon), Saxicolella (1; tropical W and C Africa), Ledermanniella (45–50; tropical and S Africa), Letestuella (1; Kunene River in Namibia), Stonesia (2; tropical W and C Africa), Macropodiella (4–6; tropical W and C Africa), Leiothylax (2; tropical Africa), Winklerella (1; tropical W Africa), Dicraeanthus (4; tropical W and C Africa), Djinga (1; Cameroon), Endocaulos (1; Madagascar), Thelethylax (2; Madagascar), Angolaea (1; Angola), Paleodicraeia (1; Madagascar), Sphaerothylax (2; tropical and S Africa, Madagascar), Maferria (1; SW India), Zehnderia (1; Cameroon); Cladopus (10; S China, S Japan, SE Asia, Malesia to New Guinea and NE Queensland), Paracladopus (2; Thailand); Hydrodiscus (1; Laos), Hydrobryum (5–10; S India, E Nepal, Assam, China, S Japan), Hanseniella (2; Thailand), Thawatchaia (1; Thailand); Willisia (1–2; Kerala in S India), Zeylanidium (5; India, Sri Lanka), Griffithella (1; W Ghats in India), Farmeria (1; Sri Lanka), Polypleurum (4; India, Sri Lanka, NE Thailand), Diplobryum (4; Laos, S Vietnam).

4. Apinagia Tul. 54 spp., a third of New World Podostemaceae, Colombia to and Uruguay, mainly in Guiana and Brazil Shields (32 in Brazil, 19 endemics).

5. Autana C. T. Philbrick. Aquatic herbs, presumed perennial, attached to rocks in river rapids and waterfalls; roots not seen; stems prostrate or upright, the flattened stems repeatedly dichotomously or subdichotomously branched. Only one sp., A. andersonii C. T. Philbrick, known from four rivers that drain into the Orinoco River, Venezuela; the species is common where it occurs.

6. Castelnavia Tul. & Wedd. Herbs, often with dense, moss-like growth or forming dense carpet like growt; stems (shoot system) prostrate, tightly attached to substratum throughout, cylindrical to flattened, repeatedly dichotomously, subdichotomously, or irregularly branched. 6 spp., from Tocantins River complex (Para, Tocantins, Goiás and Mato Grosso), two reaching into Minas Gerais and C. pendulosa (C.T. Philbrick & C.P. Bove) C.T. Philbrick & C.P. Bove up to E Bolivia.

7. Ceratolacis (Tul.) Wedd. Two spp., endemics to Brazil, scattered in Tocantins, Minas Gerais and Pernambuco states.

8. Cipoia C.T.Philbrick, Novelo & Irgang. Roots prostrate, elongate, dorsiventrally flattened to elliptical in cross-section, sometimes intertwined, green and photosynthetic branched, with asymmetric root cap, attached to rocks via holdfasts (haptera) and adhesive hairs. Two spp., C & SE region of the state of Minas Gerais.

9. Lophogyne Tul. (inc. Apinagia p.p., Jenmaniella, Maranthrum p.p., Monostyles) 13 spp., 11 in Brazil (9 endemics, two up to Guianas and Venezuela), one exclusively to Guianas, another in Venezuela, Suriname and disjunct in N Ecuador.

10. Marathrum Humb. & Bonpl. 15 spp., 7 in Mexico and Central America, two of then span this region into NW Venezuela, N Colombia and Brazil, national endemics in Venezuela (1), Cuba (1), Guyana (1), Peru (1), Colombia (2), and remaining three widely distributed in Brazil (none endemics), Argentina, Colombia, Venezuela and Guianas.

11. Mourera Aubl. (inc. Tulasneantha). 8 spp. from Brazil (5 endemics), three of them up to Colombia through Guiana Shield and Bolivia, Paraguay and northern Argentina.

12. Oserya Tul. & Wedd. 5 spp. from northern South America in Guyana, Suriname, Venezuela to center Brazil (4, one endemic).

13. Podostemum Michx. (inc. Crenias, Devillea). 12 spp., P. rutifolium Warm. widely distributed from Mexico to Argentina, one from North America to Caribbean and Central America, one only in Guianas, and remaining nine in S & SE Brazil (6 endemics), four of then up to Argentina, Paraguay and Uruguay.

14. Rhyncholacis Tul. (inc. Macarenia). 26 spp., endemic to the Guiana Shield of northern Brazil (12, 6 endemics), Colombia, Guyana, Suriname, and Venezuela, 100-1,200 m elevation range.

15. Wettsteiniola Suess. Thalloid irregular, peltate, 5–10 mm diam.; leaves bipinnate, 8–10 cm long, petiole 4–5 cm long, 3–5mm diam., rachis flattened; first division of the sheet up to 1.5 cm long, secondary divisions repeatedly furcated, last divisions numerous, filiform, 2–4 cm long, stipules up to 3 mm wide; flowers in fascicules, arising from the thalloid base. Three spp., endemics of Paraná River Basin, one very rare from Argentina, W. pinnata Suess. narrow endemic to São Paulo and Paraná states, and W. accorsii (Toledo) P. Royen widely distributed from Paraná state in Brazil to northern Argentina and Paraguay.

LINEAGE 3 de 6: CLADE MALPIGHIIDS

PUTRANJIVACEAE

§ FULLY CHLOROTROPHIC

Genera/species 2/225 Distribution pantropical, subtropical East Asia, SE Africa, Madagascar, New Caledonia and eastern Australia. Habit Usually dioecious (some species of Drypetes monoecious or polygamodioecious), evergreen trees or shrubs. Trees or rarely shrubs, lacking latex and extrafloral nectaries. The Putranjivaceae are represented by one genus in the Neotropics, occurring in several vegetation types but usually in tropical moist forests: In the New World the majority of species are Caribbean.

SYSTEMATIC outsider Putranjiva (4; Pakistan, India, Sri Lanka, Himalayas, S China (inc. Taiwan), SE Asia, Malesia to New Guinea, Japan, Ryukyu Islands).

1. Drypetes Banks ex Lam. Trees or treelets, dioecious, leaves with entire blades, inflorescences axillary or cauliflorous, fruits drupaceous. 215 spp., 120 in Asia, 75 in Africa, and 17 in New World, 8 in South America, three spp. in Brazil, two only in Amazon rainforest, both no endemics, and D. sessiliflora Allemão, endemic to Atlantic Forest of E Brazil, from Bahia to Rio de Janeiro states.

CARYOCARACEAE

§ FULLY CHLOROTROPHIC

Genera/species 2/c. 25 Distribution Central America from Costa Rica to Paraguay, tropical South America, with their largest diversity in Amazon rainforest. Habit bisexual, evergreen trees or, sometimes, shrubs or suffrutices. Confined to the Neotropics, from Costa Rica to South America.

SYSTEMATIC both genera occur in South America.

1. Anthodiscus G.F.W.Meyer. Trees or shrubs; leaves alternate, trifoliate; stipels and stipules 0; flowers medium sized. 10 spp., in Guyana, Venezuela, Colombia, Peru and W Amazon rainforest of Brazil (4, none endemics).

2. Caryocar L.Trees, sometimes large up to 50 m tall, one of them the tallest Malphighiales from Brazil, or rarely shrubs or sufrutices, sometimes with xylopodium; leaves opposite, also trifoliate; flowers large. 16 spp., from Costa Rica (up 3 in Central America), Colombia, Peru, Ecuador, Venezuela, the Guianas to central Atlantic coastal Brazil (12, two endemics); C. brasiliense Cambess. is common throughout the savannas of C Brazil (cerrado); C. gracile Wittm. is endemic to white sand areas of NW Amazon rainforest; C. cuneatum Wittm. in the dry seasonal scrubland of NE Brazil (caatinga); C. montanum Prance occurs at about 1,000 m in the Guiana Shield.

ELATYNACEAE

§ FULLY CHLOROTROPHIC

Genera/species 2/c. 50 Distribution cosmopolitan except polar areas, with their largest diversity in tropical regions. Habit bisexual, usually perennial or annual herbs (Bergia suffruticosa is suffrutescent). A large number of species are aquatic, whereas others are amphibious helophytes.

The family has historically been considered closely related to the Clusiaceae. However, recent molecular phylogenetic studies suggest that it is sister to Malpighiaceae, in the order Malpighiales. Some members of Elatine are cultivated in aquaria, and a few species of Bergia employed in traditional medicine. Otherwise the family is of little economic importance.

SYSTEMATIC both genera occur in South America.

1. Bergia L. Annual [or perennial], herbaceous [or suffrutescent], simple to much-branched procumbent to ascendent plants of moist, disturbed soils: often occurring on sand bars along rivers, but never truly aquatic. 25 spp., mostly Old World, centered in S Africa, also in SE Asia (5) and Malesia and Australia (2); two spp. in the New World, B. texana Seub. ex Walp. from northern Mexico, and California to Washington in U.S.A., and the little-known B. arenaroides Fenzl is endemic to Brazil, known only two sets in northern Minas Gerais state, collecteds in 1816-1821 and the last in 1914, possibly extinct.

2. Elatine L. Small, aquatic or emergent, opportunistic annual plants (except E. alsinastrum L.), rarely suffrutescent, herbaceous annuals or short-lived perennials of marshes, streambanks, shores of lakes and ponds, mud flats, pools, ditches, and rice fields, with life cycle either while completely submersed under water; flowers solitary in the axils of the upper leaves, [2 or] 3 [or 4]-merous. 31 spp., cosmopolitan, North America and Mexico (7), Eurasia and North Africa (12), India/Malesia (2), S Africa (2, Zimbabwe and Namibia), Australia, New Zealand, and Fiji (1), 7 spp. from N & C Andes, from Venezuela to Argentina and Chile (one also in North America), and E. lindbergii Rohrb. endemic to SE and S Brazil, from Minas Gerais to Paraná states.

MALPIGHIACEAE

§ FULLY CHLOROTROPHIC

Genera/species 77/1,035-1,040 Distribution tropical and subtropical regions, with their largest diversity in tropical South America. Habit usually bisexual (rarely polygamomonoecious), evergreen or deciduous trees, shrubs, suffrutices or lianas. Malpighiaceae are an angiosperm family of trees, shrubs, and vines in the tropical and subtropical forests and savannas of the New and Old Worlds. They comprise 1,300 spp. in 77 genera, with 150 species belonging to 17 exclusively Old World genera. The majority of the genera and species are found in the New World, and the only members of the family found in both hemispheres are two species of New World genera that also occur in coastal W Africa.

About 45 genera and 530 spp. in Brazil, 90 Byrsonima and 95 Heteropterys. Malpighia emarginata D.C.is a tropical fruit-bearing shrub or small tree in this family. Common names include acerola, Barbados cherry, Caribbean cherry and wild crepemyrtle. Acerola is native to Yucatan, and found in South America, southern Mexico and Central America, but is now also being grown as far north as Texas and in subtropical areas of Asia, like India. Byrsonima species is popularly known as ‘murici’. There are several properties attributed to the leaves of Byrsonima species including febrifuge, to treat gastrointestinal dysfunctions and skin diseases. Ayahuasca is any of various psychoactive infusions or decoctions prepared from the Banisteriopsis spp. is vine, usually mixed with the leaves of dimethyltryptamine (DMT)-containing species of shrubs from the genus Psychotria.

Only two shared Old/New World: Heteropterys and Stigmaphyllon. Bunchosia, Malpighia and Byrsonima ate the three arborescent genera of Malpighiaceae with fleshy, bird-dispersed fruits.

SYSTEMATIC two subfamilies, both in South America.

1. SUBFAMILY BYRSONIMOIDEAE (10/215–240) ‣ three tribes, all in South America.

1.1 BYRSONIMIOIDEAE ▸ TRIBE BYRSONIMEAE (3/c 180) - all genera occur in South America.

1. Blepharandra Griseb. Trees or shrubs; leaves eglandular; inflorescence terminal, unbranched or basally ternate, each branch a pseudoraceme composed of 1–several-flowered cincinni; corolla bilaterally symmetrical; petals white, pink, or red, the posterior petal pale yellow in two species; fruit dry, indehiscent. 6 spp., endemic to the Guiana Shield of Colombia, Guyana, southern Venezuela and Amazon rainforest of northern Brazil (5, two endemics), at 100-2200 m elevation range.

2. Byrsonima H. B. K. Trees, shrubs, or subshrubs, sometimes with woody rhizomes or xylopodium, never vines; inflorescence terminal, mostly a pseudoraceme of 1-flowered non-decussate cincinni but in some species a raceme of few-flowered cincinni; corolla bilaterally symmetrical; petals light or medium yellow, white, pink, or red, glabrous in most species; fruit a drupe, the thin flesh green turning yellow, orange, red, purple, blue, or blue-black at maturity. 150 spp., from southern Mexico, SE Florida, and the Caribbean to SE Brazil, absent in Argentina, centered in South America (131), in wet forests, but the genus is most diverse in savannas and other relatively open types of vegetation. 100 spp. in Brazil, 54 endemics, 9 of then from several states are rare plants in Brazil, by Plantas Raras do Brasil’s book.

Fruits of B. crassifolia (L.) H. B. K. sens. lat., which are yellow and about the size of a cherry, are gathered from wild and cultivated trees from Mexico to C South America and sold in markets under the names chaparro, chaparro manteco, muricí, murucí, nance, nanche, and nancite; the fruits are eaten as is or used to make ice cream (Belém) or wine (Costa Rica), and in the Yucatan Peninsula of Mexico they are sold preserved in a sweet syrup; the astringent qualities of the bark, for which the genus was named, are no longer important in tanning leather, but one does occasionally see a comment on a plant label that the bark is a febrifuge.

3. Diacidia Griseb. Trees, shrubs, or subshrubs; vegetative hairs mostly basifixed or sub-basifixed; leaves eglandular; inflorescence terminal, a simple or compound thyrse composed of several-flowered non-decussate cincinni or a pseudoraceme of 1-flowered cincinni; corolla bilaterally symmetrical; petals light yellow, often with red claws. 11 spp., collected among granitic outcrops and in meadows, savannas, scrub, and rocky tepui slope forests, at 100-2,300 m elevation range, in two subgenera:

§ subg. Sipapoa ‣ 10 spp., on mountains of southern Venezuela (7 endemics) and adjacent Brazil (3, only Amazonas state, D. aracaensis W.R. Anderson endemic).

§ subg. Diacidia ‣ D. galphimioides Griseb., widely distributed in the drainages of Río Negro and Río Vaupés in Venezuela, Colombia, and Amazonas state in northern Brazil.

1.2 BYRSONIOIDEAE ▸ TRIBE ACMANTHEREAE (3/14) - all genera occur in South America.

4. Acmanthera (Adr. Juss.) Griseb. Trees, shrubs, or subshrubs; inflorescence an axillary shoot with 1 internode and 1 node below a pseudoraceme of single or clustered non-decussate flowers; corolla bilaterally symmetrical; petals mostly white, occasionally described as pinkish or pale yellow, glabrous or abaxially hairy; lateral petals entire, erose, or denticulate; fruit dry. 7 spp., all Brazilian endemics, most from near rivers in forests that are periodically flooded, in the Amazon rainforest; one species is known from north-amazonic white-sand savannas (campinaranas) in Amazonas state, another from savannas (cerrado) of Piauí state; this species, with another three in Amazonas state are rare plants in Brazil, by Plantas Raras do Brasil’s book.

5. Coleostachys Adr. Juss. Shrub or treelet 1.5–4 m tall, monopodial; leaves eglandular; inflorescence an elongated axillary spike (unique spikes among Malpighiaceae); petals white, abaxially sericeous in center, the posterior also sparsely sericeous adaxially on claw and base of limb; lateral petals entire; fruit dry. Only one sp., Coleostachys genipifolia Adr. Juss., imperfectly known from few collections in wet forests of the Amazon rainforest of French Guiana and adjacent Brazil (Amapá, Pará and Maranhão states).

6. Pterandra Adr. Juss. Trees, shrubs, or subshrubs; vegetative hairs medifixed in most species, basifixed in one; inflorescences lateral, a series of 2–6-flowered sessile or subsessile fascicles; corolla almost radially symmetrical; petals white or greenish yellow or green, becoming yellowish in age, or pink, abaxially sparsely to densely hairy; fruit dry. 15 spp., two in Panamá and remaninig in South America, mainly Colombia, Venezuela, and Brazil (8, 7 endemics, three of then are rare plants in Brazil, by Plantas Raras do Brasil’s book), mainly in wet or mesic forests, but a few (including the only species that is widely distributed and well collected, P. pyroidea A.Juss.) inhabit campos and savannas of C Brazil (cerrado).

1.3 BYRSONIMIOIDEAE ▸ TRIBE GALPHIMIEAE (5/41) - all genera occur in South America.

7. Andersoniella C. Davis & Amorim. (off Lophanthera). Small- to medium-sized shrubs or trees, stems densely or loosely sericeous to glabrate, hairs copper-colored to reddish when present, lenticels present or not; inflorescence terminal, rarely axillary, pendent, rarely erect, with a pair of sterile and much-reduced leaves at base or not, thyrse composed of cincinni of 1–3(–4–5) bisexual flowers; petals generally pink or white to pink. Three spp., in Central America (Nicaragua and Costa Rica) and southward to NW South America in the Amazon region in Colombia and the Brazilian state of Amazonas (only A. spruceana (Nied.) C. Davis & Amorim, non endemic).

8. Galphimia Cav. Shrubs or subshrubs, occasionally small trees; inflorescence terminal, erect, a pseudoraceme of single flowers; corolla varying from moderately bilaterally symmetrical to nearly radial; petals bright yellow, often suffused with red or turning red in age, glabrous or sparsely hairy on abaxial midrib, minutely denticulate or slightly erose. 26 spp., 22 in North American, mostly Mexican but extending as far north as Texas and as far south as Nicaragua, and 4 in South America, in seasonally dry, open habitats of in Paraguay (one endemic), Argentina, Uruguay, Bolivia and over Brazil (4, G. brasiliensis (L.) A. Juss. endemic, in Paraíba, Pernambuco, Piauí, and Bahia).

9. Lophanthera Adr. Juss. (exc. Andersoniella) Shrubs or trees; leaves bearing glands on petiole or immersed in abaxial surface of lamina or both; inflorescence terminal, often pendent, an unbranched raceme of few-flowered cincinni or dichasia or a pseudoraceme of single flowers [1-flowered cincinni]; corolla bilaterally symmetrical; petals bright yellow, entire or minutely denticulate, glabrous or minutely pilose on margin. Three spp., northern Brazil, two extending into the Alto Orinoco drainage in Venezuela, Colombia and Peru, along rivers in wet forests or in sandy open places near rivers, and the cocci are very probably dispersed by water. L. lactescens Ducke, a only Brazilian endemic, is an ornamental street tree, which is interesting given the fact that the species is rare in nature, known from only a few collections in Xingu Valley in Pará state, northern Brazil.

10. Spachea A.Juss. Shrubs or trees; leaves usually bearing glands immersed in both surfaces of lamina; inflorescence terminal or lateral, sometimes pendent, an unbranched raceme of few-flowered cincinni or a pseudoraceme of single flowers; corolla bilaterally symmetrical; petals pink or white, minutely denticulate or fimbriate, glabrous or with a few hairs at base of claw. 6 spp., one in Cuba, two only Colombia, another only in southern Central America, and two in northern South America (including Trinidad), both in Brazil, none endemics; wet forests, mostly at low elevations.

11. Verrucularina A. Juss. Shrubs; leaves eglandular; inflorescence terminal, erect, an unbranched, often corymbose raceme of few-flowered cincinni or single flowers; corolla bilaterally symmetrical, the posterior petal somewhat larger than the lateral petals; petals bright yellow, glabrous or sparsely red-tomentose abaxially on claw or narrow crest of limb, denticulate to subentire. Two spp. in Brazil, but very high disjuncts: V. piresii (W.R. Anderson) Rauschert from sandy savannas on Mount Aracá in Amazonas state, near Venezuela, and V. glaucophylla (A. Juss.) Rauschert from shrubby vegetation on sandstone hills in central Bahia.

2. SUBFAMILY MALPIGHIOIDEAE (c 63/1.170-1.205) ‣ 11 lineages, Acridocarpus clade (2/c 33, tropical regions in the Old World) is sister to the remaining Malpighioideae e only lineage among this subfamily absent in South America.

2.1 MALPIGHIOIDEAE ▸ TRIBE MCVAUGHIOIDEAE (3/7) - all genera in South America.

12. Burdachia Mart. ex Endl. Trees or shrubs; lamina bearing several–many glands in abaxial surface; inflorescence terminal, single or 2 or 3 together, each usually divided near base into 3 (–5) axes, each axis a raceme of short 1–6-flowered cincinni; corolla bilaterally symmetrical; petals pink or white, glabrous; limb of outermost petal concave; lateral petals with the margin entire; fruit dry, indehiscent. Three spp. in Guyana and Amazon rainforest of Venezuela, Colombia, Peru and Brazil (all, one endemic), and in lowland forests near rivers or in low, periodically flooded places; the dry indehiscent fruits are well adapted for dispersal by water.

13. Glandonia Griseb. Trees or shrubs; inflorescence terminal, mostly unbranched, a raceme of short 2–5-flowered cincinni; corolla without a plane of symmetry; petals white or lateral petals white and posterior petal yellow, glabrous or bearing a tuft of hairs abaxially at base of claw; fruit dry, indehiscent. Three spp., of Amazonas state in northern Brazil (two endemics), with G. williamsii Steyerm. up to Colombia, Venezuela, in lowland forests along rivers or in areas periodically flooded.

14. Mcvaughia W. R. Anderson. Shrubs; inflorescence terminal, unbranched but sometimes ternate, each axis a raceme of short 2–7-flowered cincinni; bracts and bracteoles persistent; lowest bracteole and alternate subsequent bracteoles bearing 1 large eccentric abaxial gland; pedicel straight in bud. corolla bilaterally symmetrical; petals bright yellow, glabrous; fruit a dry, indehiscent nut. Three spp. in dry seasonal scrubland of NE Brazil (caatinga) on sandy soils of lowland NE Bahia and SE Piauí states, and sandy coastal plains in Sergipe state, all in NE Brazil.

2.2. MALPIGHIOIDEAE ▸ BARNEBYA CLADE (1/2) - a single genus.

15. Barnebya W. R. Anderson & B. Gates. Trees or woody vines; leaves alternate or appearing opposite when leaves are crowded; inflorescence terminal, unbranched or with a dominant axis and many weaker axes from near base; flowers with the calyx, corolla, and androecium perigynous; corolla bilaterally symmetrical; petals yellow, glabrous or thinly sericeous abaxially on claw; fruit dry. Two spp. of lowland wet and dry forests and dry seasonal scrubland of NE Brazil (caatinga), from Piauí and Pernambuco to São Paulo.

2.3 MALPIGHIOIDEAE ▸ TRIBE PTILOCHAETOIDEAE (4/5) - outsider Lasiocarpus (5; Mexico)

16. Dinemandra A. Juss. Ericoid shrubs. Only one sp., D. ericoides A. Juss., endemic to N Chile.

17. Dinemagonum A. Juss. Ericoid shrubs. Only one sp., D. gayanum A. Juss., endemic to N Chile.

18. Ptilochaeta Turcz. 4 spp. from Brazil (P. glabra Nied. endemic), three also in northern Argentina, Bolivia and Paraguay.

2.4 MALPIGHIOIDEAE ▸ TRIBE TRISTELLATOIDEAE (6/105–110) - outsiders Echinopterys (2; Mexico), Tristellateia (c 20; Madagascar, one in tropical Africa, another in tropical Asia and eastwards to tropical Australia and New Caledonia), Henleophytum (1; Cuba).

19. Bunchosia H. B. K. Trees or shrubs; inflorescence mostly unbranched but ternate in a few species, axillary; corolla bilaterally symmetrical; petals lemon-yellow or occasionally whitish, usually glabrous; fruit indehiscent, a berry of 2 or 3 1-seeded pyrenes in a common fleshy exocarp, yellow, orange, or red at maturity, the pyrenes free from each other at maturity, with a smooth, brittle, cartilaginous wall. 84 spp., from Mexico and the Caribbean to SE Brazil (11, 5 endemics, one of then from Bahia state is a rare plant in Brazil, by Plantas Raras do Brasil’s book) and adjacent Argentina, in habitats including relatively dry woodlands, savannas, and wet forests, mainly in the periphery of the Amazon rainforest and the Brazilian Shield, also in wet forests of eastern Ecuador and Peru; 48 spp. in South America; the flesh of the fruits is edible, with the fruits of some species reaching the size of small plums.

20. Heladena Adr. Juss. Woody vine, occasionally described as a shrub or small tree; vegetative hairs medifixed; leaves opposite; inflorescence unbranched, terminating a leafy branch with full-sized leaves or lateral, axillary to a full-sized leaf and then usually bearing 1 pair of much-reduced leaves, these often deciduous; pedicels pedunculate; corolla bilaterally symmetrical; fruit dry. Only one sp., H. multiflora (Hook. & Arn.) Nied., southern Brazil, Paraguay, and NE Argentina, in gallery forest or woodland, often near rivers.

21. Thryallis Mart. Scandent shrubs and woody vines; vegetative hairs stellate; leaves bearing 1 or 2 pairs of glands at apex of petiole and/or on base of lamina; inflorescences terminal and lateral, variously grouped; corolla bilaterally symmetrical; petals glabrous, the limb oblate (wider than long); lateral petals with the margin erose or irregularly dentate to nearly lacerate; fruit dry. 5 spp. of Brazil (three endemics, T. parviflora C.E.Anderson from Distrito Federal is a rare plant in Brazil, by Plantas Raras do Brasil’s book), two of then up to adjacent Paraguay and Bolivia, mainly of open habitats, thickets, dry seasonal scrubland of NE Brazil (caatinga), secondary forest, roadsides, but they have occasionally been reported from gallery forest.

2.5 MALPIGHIOIDEAE ▸ TRIBE HIRAEOIDEAE (5/50) - outsider Psychopterys (8; southern Mexico, Belize, Guatemala).

22. Adelphia W. R. Anderson. Woody vines; leaves opposite; inflorescence an open, elongated, axillary or terminal panicle or pseudoraceme with the flowers borne singly or (most often) in 2s, 3s, or 4s; floriferous peduncles well developed; samaras separating from a short pyramidal torus. 4 spp., Colombia and Peru one endemic each, one from in wet evergreen or semideciduous forests of Jamaica, SE Mexico to Venezuela and Ecuador, and A. macrophylla (Rusby) W.R. Anderson in Peru, Bolivia and Acre state in Amazon rainforest of Brazil.

23. Excentradenia W.R.Anderson. Woody vines; leaves opposite, subopposite, or alternate; Inflorescence a single short axillary raceme of 3–7 (–9) 4-flowered umbels, with 1 umbel terminal and the other 1–4 pairs axillary to bracts bearing stipules and often petiole glands; corolla bilaterally symmetrical, petals bright yellow, glabrous; samaras with a large, membranous. 4 spp. from northern South America, two only in Venezuela and adjacent Guianas, another only in Bolivia, and E. primaeva (W.R. Anderson) W.R. Anderson, a rare plant in Brazil, by Plantas Raras do Brasil’s book, only in Amazonas state in northern Brazil.

24. Hiraea Jacq. Woody vines, sometimes shrubby; leaves mostly opposite, sometimes ternate; inflorescences axillary, usually 1–several umbels of 4–many flowers, the umbels when 4-flowered often borne in a cyme; corolla bilaterally symmetrical; petals mostly yellow or yellow turning orange or red, glabrous; samaras separating from a very low pyramidal torus. 75 spp., from W Mexico to Paraguay and adjacent Argentina and SE Brazil (21, 13 endemics), also in Lesser Antilles (Grenada, St. Lucia), in diverse habitats but avoiding very dry vegetation types; 68 spp. in South America.

25. Lophopterys Adr. Juss. Woody vines or shrubs (small trees); inflorescence paniculate, rarely simple, the flowers ultimately borne in pseudoracemes; corolla bilaterally symmetrical; petals bright yellow, glabrous or only very sparsely sericeous abaxially; samaras separating from a short pyramidal torus. 7 spp., from Colombia to SE Brazil, Bolivia to French Guiana, wider in tropical continent; three in Brazil, L. floribunda W.R. Anderson & C. Davis endemic.

2.6 MALPIGHIOIDEAE ▸ TRIBE TETRAPTERYDIDEAE (c 15/c 330) - outsiders Hiptage (30–35; Mauritius, Sri Lanka, northern Pakistan, southern Himalayas to S China inc. Taiwan, SE Asia, Malesia to Fiji), Flabellariopsis (1; tropical Africa), Flabellaria (1; tropical Africa).

26. Alicia W.R.Anderson. Vines, white, pink, or lilac corolla. Two spp., both over South America and in Brazil, none endemics.

27. Callaeum Small. Woody vines, or shrubs with scadent or trailing branches. 11 spp., 8 from Texas to Honduras, one in Andes to Ecuador to Peru, C. antifebrile (Griseb.) D.M. Johnson in northern South America up to Brazil, and C. psilophyllum (A. Juss.) D.M. Johnson in Brazil, Bolivia and Paraguay.

28. Carolus W.R.Anderson. Woody vines. 6 spp., one in Central America, another in Ecuador, two endemics to Brazil, C. chlorocarpus (A. Juss.) W.R. Anderson in Brazil and adjacent Brazil and Cono Sur, and C. sinemariensis (Aubl.) W.R. Anderson in over tropical neotropics.

29. Christianella W.R.Anderson. Woody vines, or shrubby in open habitats. 5 spp., scattered in Mexico to Guianas and Cono Sur, in forests, roadsides thickets, and shrubby savannas; 4 spp. in Brazil, C. paludicola (W.R. Anderson) W.R. Anderson endemic.

30. Dicella Griseb. Woody vines; inflorescences a decompound panicle the flowers borne ultimately in short pseudoracemes with decussate bracts; petals yellow, abaxially densely sericeous; fruit composed of a dry, hard, indehicent, nut-like structure. 7 spp., one in Costa Rica, two endemics to Brazil, three from Brazil to Bolivia and Paraguay, and one from Colombia and Peru to center Brazil.

31. Glicophyllum R.F.Almeida. (exc. Tetrapterys) Woody vines, scandent shrubs or subshrubs. 27 spp., in evergreen, seasonally dry forests and savannas from Mexico, Central and South America.

32. Heteropterys Kunth. Woody vines, shrubs or small trees; leaves usually bearing glands; flowers borne in umbels, corymbs, or pseudoracemes; these single or grouped in racemes or panicles, axillary or terminal; petals mostly yellow or pink. 155 spp., from Mexico to Argentina (except Chile), Caribbean and Africa, with one widely distributed, mostly Caribbean species, H. leona (Cav.) Exell, is also found in low wet places along the coast of W Africa from Senegal to Angola; 144 spp. in South America, 101 spp. in Brazil, 58 endemics, 11 of then, from several states, are rare plants in Brazil, by Plantas Raras do Brasil’s book; two subgenus, four sections and six subsections (Aptychia, Metallophyllis, Parabanisteria, Rhodopetalis, Stenophyllarion and Xanthopetalis), all in Brazil, belongs this genus.

H. aphrodisiaca O. Mach., endemic to the Brazilian scrubland regions, is traditionally used in folk medicine as an aphrodisiac, a stimulant and in the treatment of nervous weakness; the herb has been found to reduce the toxic effects of cyclosporin A on rat testis, increase tendon strength with endurance training in rats, and improve memory in aged rats. Additionally, an aliphatic nitro compound extracted from the root was found to have antimicrobial and antiviral effects in vitro.

33. Jubelina A.Juss. Woody vines; leaves with the petiole eglandular; inflorescence axillary and terminal, decompound, thyrsiform, containing much-reduced bract-like leaves below the floriferous bracts; petals yellow or pink or pink and white; fruit breaking apart into 3 samaras. 6 spp. from northern South America in French Guiana to Peru, northern Brazil (2, none endemics), one up to Central America.

34. Malpighiodes Nied. 4 spp. in northern South America from Colombia to French Guiana and Brazil (two, Amazonas and Pará states, M. bracteosa (Griseb.) W.R. Anderson endemic).

35. Mezia Schwacke ex Nied. Woody vines, shrubs, or small trees; leaves with the petiole eglandular; inflorescences tighly reddish- or brown-sericeous throughrout, axillary and terminal, often decompound; petals yellow; fruit breaking apart into 3 samaras. 15 spp. of South America to Bolivia and French Guiana, 7 in Brazil, three endemics, mainly forests.

36. Niedenzuella W.R.Anderson. (inc. Aenigmatanthera). 18 spp. from South America, one up to Central America, 15 in Brazil, 6 endemics.

37. Tetrapterys Cav. (exc. Glicophyllum) Woody vines or shrubs, occasionally described as small trees, oftem with xylopodium; leaves usually bearing glands; flowers borne in umbels, corymbs, or pseudoracemes, these often grouped in panicles; petals yellow or pink; fruits breaking apart into 3 samaras. 53 spp., Mexico to South America (except Chile and Uruguay); 34-62 in South America, 6-33 in Brazil, 1-16 endemics; T. cordifolia W.R.Anderson from Mount Aracá in Amazonas state is a rare plant in Brazil, by Plantas Raras do Brasil’s book.

38. Tricomaria Hook. & Arn. Multi-branched shrub. Only one sp., T. usillo Hook. & Arn., endemic to dry vegetation in W Argentina.

2.7 MALPIGHIOIDEAE ▸ TRIBE STIGMAPHYLLOIDEAE (15/445–450) - outsiders Sphedamnocarpus (18; tropical and southern Africa, Madagascar), Philgamia (4; Madagascar), Cottsia (3; southern U.S.A., northern Mexico).

39. Aspicarpa Rich. Perennial herbs (rarely in this family). 11 spp., U.S.A. up to Paraguay, 7 in South America, 4 in Brazil, A. harleyi W.R. Anderson endemic.

40. Banisteriopsis C.B.Rob. ex Small. Vines, shrubs, or rarely small trees; leaves bearing glands on petiole or abaxial surface of blade or both; petals yellow, pink, or white, usually lateral 4 spreading or reflexed and the posterior erect; fruit breaking apart into 3 samaras. 60 spp. from South America except one from Caribbean, six up to Mexico and Central America; 46 in Brazil, 30 endemics, 10 of then, from Goiás, Minas Gerais and Rio de Janeiro states, are rare plants in Brazil, by Plantas Raras do Brasil’s book.

41. Bronwenia W.R.Anderson & C.C.Davis. Woody vines or shrubs, when shrubby the branches often twining; leaves decussate; inflorescence axillary or terminal, paniculate or dichasial; corolla bilaterally symmetrical; petals yellow, glabrous, the posterior petal different in size, shape, and stance from the lateral 4; fruit dry, breaking apart at maturity into 3 samaras. 10 spp. of South America, three up to Mexico and Central America, mostly in wet forests and savannas in South America, often in drier vegetations in Central America and Mexico; Brazil has 8 spp., five endemics, two of then, from Amazonas and Mato Grosso do Sul states, are rare plants in Brazil, by Plantas Raras do Brasil’s book; moist forests, seasonally dry forests, tropical deciduous forests, dry thickets.

42. Camarea A.St.-Hil. Erect or climbing shrubs, sometimes with ericoid leaves and xylopodium. 6 spp. from Brazil (4 endemics), two also outside, C. affinis A. St.-Hil. highly disjunct in southern Guyana, and savannas of Brazil and Paraguay, and C. axillaris A. St.-Hil. also in Cono Sur.

43. Diplopterys A.Juss. Woody vines; inflorescences axillary, shorter then the subtending leaves, of 1-several simple 4-flowered umbels or 1-several racemes of up to 7 4-flowered umbels; petals yellow, long-fimbriate. 31 spp., one endemic to Mexico, three from Central and South America, remaining 27 only South America, over tropical regions of continent, 21 in Brazil, 11 endemics, two of then (both in Bahia state) are rare plants in Brazil, by Plantas Raras do Brasil’s book.

44. Gaudichaudia Hicken. Lianas and subshrubs. 19 spp., 18 spp. Mexico (16 endemics) and Central America, one extends into NW South America in Venezuela and Colombia, and one endemic to Caribbean.

45. Janusia A.Juss. (exc. Peregrina) 15 spp. of Brazil (10 endemics, one of then from Bahia state is a rare plant in Brazil, by Plantas Raras do Brasil’s book) from southern Amazonas to coast of NE region and Rio Grande do Sul, five of then up to Argentina, Bolivia, Paraguay and Uruguay and Venezuela.

46. Mionandra Griseb. (inc. Cordobia, Gallardoa) 4 spp. from N Argentina, Bolivia and Paraguay (one endemic).

47. Peixotoa A.Juss. Shrubs, subshrubs, or vines; vegetative hairs medifixed and mostly stalked, the stalk nearly or quite absent in a few species; inflorescences terminal and/or axillary, with flowers borne in umbels of 4 - 8, the umbels variously grouped in compound inflorescences; corolla bilaterally symmetrical; petals yellow, glabrous, fimbriate or denticulate on the marginar; fruit dry. 30 spp., 29 spp. of Brazil, only three extending of adjacent Bolivia and Paraguay and one endemic to Paraguay; the shrubby spp., occur mostly in savannas of C Brazil (cerrado), while the vines are in woodylands, gallery forests margins; 10 spp. in several center and southern states, are rare plants in Brazil, by Plantas Raras do Brasil’s book.

48. Peregrina W. R. Anderson. (off. Janusia) Perennial with many stems from woody rootstock; stems slender, erect, non-twining, 15- 45 cm tall, sericeous, eventually glabrescent; leaves opposite or subopposite; inflorescence a terminal umbel of (2-)4-6(-12) flowers, the flowers all chasmogamous; petals 5, orange-yellow, glabrous, flat, short-fimbriate; fruit breaking apart into 3 samaras borne on a short pyramidal torus. Only one sp., P. linearifolia (St. Hill) W. Anderson., grassy sandy fields of Paraná state in S Brazil, Paraguay and NE Argentina, at 800-900 m.

49. Stigmaphyllon A.Juss. Woody or herbaceous vines, a few species shrubby; leaves with the blade entire or lobed; inflorescence unbranched or more commonly a dichasium (or occasionally a small thyrse) of congested pseudoracemes, these corymbose or umbellate; petals yellow or yellow and red; fruit dry, breaking apart into 3 samaras. 115 ssp. in two subgenera:

§ subg. Stigmaphyllon ‣ 94 spp., mostly twining woody vines or rarely shrubs native to the Neotropics from southern Mexico to northern Argentina, except Chile and the high Andes, 13 in Caribbean. S. bannisterioides (L.) C.E.Anderson. is also found in seashore vegetation along the Atlantic Coast from southern Mexico to northern Brazil, in the Caribbean, and along the coast of W Africa (Guinea Bissau, Guinea, Sierra Leone); 81 spp. in South America, 50 in Brazil, 35 endemics, 11 of then, in several states, are rare plants in Brazil, by Plantas Raras do Brasil’s book.

§ subg. Ryssopterys ‣ 21 species of woody vines of the Taiwan in China, Ryukyu Islands, Sunda Islands (except Borneo and Sumatra), New Guinea, Queensland (Australia), New Caledonia, Vanuatu, Solomon Islands, Micronesia, Palau, and Philippines.

2.8 MALPIGHIOIDEAE ▸ ECTOPOPTERYS CLADE (1/1) - a single genus.

50. Ectopopterys W.R.Anderson. Woody vine; inflorescence a terminal and lateral panicle, the flowers borne in decussate 4-flowered umbels, 6-flowered corymbs, or pseudoracemes; floriferous peduncle well developed; bracts and bracteoles eglandular; corolla bilaterally symmetrical, the posterior petal very different from the lateral 4; petals light yellow, glabrous; lateral petals with the margin dentate; fruit dry. One woody vine, E. soejartoi W.R. Anderson, in lowlands wet forests of Colombia, Ecuador, and Peru.

2.9 MALPIGHIOIDEAE ▸ AMORIMIA CLADE (1/10) - a single genus.

51. Amorimia W.R.Anderson. Woody vines, sometimes described as shrubs; leaves usually opposite, sometimes subopposite or alternate; inflorescence with the flowers borne in elongated terminal or axillary pseudoracemes or panicles; corolla bilaterally symmetrical; petals yellow or yellow turning red-orange in age; fruit dry, breaking apart into samaras separating from a short or moderately high pyramidal torus. 15 spp. from South America, found in diverse habitats from Argentina and northern Colombia to Rio Grande do Sul state in Brazil (12, 8 endemics, one of then from Minas Gerais state a rare plant in Brazil, by Plantas Raras do Brasil’s book). Two subgenera:

§ subg. Amorimia ‣ 8 spp., all in Brazil and endemics except by A. exotropica (Griseb.) W.R.Anderson up to E Paraguay.

§ subg. Uncinae ‣ 7 spp., Colombia, Ecuador and Peru one endemic each, one in northern Brazil, Peru and Bolivia, and remaining three endemics to Brazil.

2.10 MALPIGHIOIDEAE ▸ TRIBE MALPIGHIEAE (c 10/120–125) - outsiders Calcicola (2; Mexico), Aspidopterys (c 25; tropical Asia), Caucanthus (3; E and NE Africa, Arabian Peninsula), Triaspis (c 15; tropical and southern Africa), Digoniopterys (1; Madagascar), Rhynchophora (2; Madagascar), Madagasikaria (1; Madagascar), Microsteira (c 25; Madagascar).

52. Malpighia L. Shrubs or trees up to 10 (–24) m tall; inflorescence axillary, an unbranched pseudoraceme congested in most species into a dense corymb or umbel, in some species with only 1-2 flowers; bracts and bracteoles eglandular; pedicels pedunculate; flowers bilaterally symmetrical in calyx, corolla, and androecium; petals pink, lavender, or white, glabrous or nearly so; fruit fleshy, mostly indehiscent, drupes or berries, usually red at maturity. 108 spp., 23 only in Mexico (21 endemics) and Central America, 59 exclusively Caribbean, three in Caribbean and mainland Mesoamerica, and 4 up to South America, two only up to northern Colombia, and two others up to Colombia, Ecuador and Venezuela.

M. glabra L., in relatively dry deciduous or semi-deciduous forests, occasionally in the understory of lowland evergreen forests, 200-300 m, from southern Texas to Ecuador, Greater Antilles, and Venezuela, and M. emarginata DC., which occus in South America in dry areas with xerophytic vegetation, near sea level to 200 m, common in the coastal lowlands of northern Venezuela, where perhaps native, and elsewhere cultivated; apparently native from Mexico to Honduras, elsewhere in Central America and the Caribbean probably escaped from cultivation, apparently native in NW Colombia.

53. Mascagnia (Bertero ex DC.) Colla. Twining vines with woody stems varying from slender to stout, occasionally described as shrubby; leaves decussate; inflorescences elongated or occasionally congested pseudoracemes, single or grouped in panicles; flowers bilaterally symmetrical in all whorls, bisexual; petals yellow, pink, white, or various shades of lilac, blue, or purple, glabrous, entire, erose, or dentate; fruit dry. 42 spp. from northern Mexico to northern Argentina and SE Brazil (19, 5 endemics, two of then, in Mato Grosso and Espírito Santo states, are rare plants in Brazil, by Plantas Raras do Brasil’s book) and in the Caribbean, growing in diverse habitats; 32 spp. in South America.

LINEAGE 4 de 6: CLADE CHRISOBALANIDS

TRIGONIACEAE

§ FULLY CHLOROTROPHIC

Genera/species 5/c. 30 Distribution Madagascar, W Malesia, Central America, tropical South America. Habit bisexual, evergreen trees, shrubs or lianas. Trigonia nivea Cambess. and Trigonia villosa Aubl. var. villosa are used as ornamentals. As medicine has T. nivea var. candida used in the cure of renal diseases in Paraty municipality, Rio de Janeiro state, and Trigonia microcarpa Sagot ex Warm. is used by the indians in the Amazon rainforest to increase uterine contractions in childbirth.

SYSTEMATIC outsiders Humbertiodendron (1; E Madagascar) and Trigoniastrum (1; W Malesia).

1. Isidodendron Fernández-Alonso Pérez & Idarraga. Tall trees; papilionaceous flowers. Only one sp., I. tripterocarpum Fern. Alonso, Pérez Zab. & Idárraga, in tropical moist forests, primary or secondary and altered zones of Colombia.

2. Trigonia Aubl. Treelets, shrubs, scandent shrubs or lianas; papilionaceous flowers. 29 spp., all in South America, two up to Central America and Mexico, inhabiting primary forests, others are colonial, pioneer being among the contingent that arises in degraded areas in the three Americas, up to Argentina; 19 in Brazil, 7 endemics.

3. Trigoniodendron E.F. Guim. & Miguel. High trees; papilionaceous flowers. Only one sp., T. spiritusanctense E.F. Guim. & Miguel., in the Atlantic Forest of Espírito Santo state in SE Brazil.

DICHAPETALACEAE

§ FULLY CHLOROTROPHIC

Genera/species 3/160-165 Distribution pantropical, southwards to SE and southern Africa. Habit usually at least morphologically bisexual (at least sometimes functionally monoecious or dioecious), evergreen trees, shrubs or lianas. Sometimes xerophytic. Lenticels often numerous. Trees, shrubs, lianas, or suffruticose subshrubs. A tropical family of about 240 species in three genera, distributed throughout the lowland tropical regions of both hemispheres (but absent from Polynesia and Micronesia), extending into the subtropics in Africa and India.

Use Medicinal plants, arrow poisons, mammal pesticides. Some species are poisonous, especially to cattle; many members of this family has true epiphylly, with inflorescences in many positions at leaves.

SYSTEMATIC a phylogeny is carried out by a research group in Wageningen. Tapura is nested in Dichapetalum, according to Yakandawala & al. (2010); all genera occur in South America.

1. Dichapetalum Thouars. Mainly lianas, a few small trees or shrubs; some spp. may be either shrubs or lianas or even small trees. 142 spp., 90 in Africa, 7 in Madagascar, 16 in SE Asia to Pacifico, 29 in New World, 19 in South America, 8 in Brazil, two endemics.

2. Stephanopodium Poepp. Small to medium-sized trees, small incompiscuous inflorescences. 15 spp., 14 in South America, 7 from Venezuela to Peru, and 7 endemics to Atlantic Forest of SE Brazil from Bahia to Rio de Janeiro and Minas Gerais states.

3. Tapura Aubl. 34 spp., 8 in tropical Africa and 26 in tropical America, 21 in South America, 12 in Brazil, 5 endemics, including 4 in Atlantic Forest, all endemic within.

EUPHRONIACEAE

§ FULLY CHLOROTROPHIC

Genera/species 1/3 Distribution northern tropical South America (Guiana Shield). Habit bisexual, evergreen tree or shrub. Confined to Amazon rainforest of Brazil, Colombia and Venezuela; a single genus; the unique family endemic to the Guiana Shield.

The family Euphroniaceae is placed in the order Malpighiales, closest to Chrysobalanaceae in the APG III classification (Stevens, 2008; APG III, 2009). Takhtajan (1997) placed the family in the Chrysobalanales, together with Trigoniaceae, Dichapetalaceae and Chrysobalanaceae. It was considered by Cronquist (1982) as part of the Vochysiaceae, while Marcano-Berti (1989) segregated the genus Euphronia and gave it family status.

Key differences from similar families

Superficially resembling Vochysiaceae and Trigoniaceae, but keyed out as follows:

1. Leaves alternate (but see also Trigoniodendron) ------------ Euphroniaceae

1. Leaves opposite or in whorls - 2

2. Calyx spurred; petals 0-5, without a spur ------------ Vochysiaceae

2. Calyx without a spur; petals 5, one of which is spurred ------------ Trigoniaceae.

From Chrysobalanaceae it can be keyed out as follows:

1. Petals 3; staminodes present; ovary 3-locular; fruit a 1-seeded capsule ------------ Euphroniaceae

1. Petals 5; staminodes absent; ovary 1-2-locular; fruit a drupe ------------ Chrysobalanaceae

SYSTEMATIC only one genus from South America.

1. Euphronia Mart. & Zucc. Shrubs or small trees; leaves alternate, simple, lower surface covered with greyish white hairs; inflorescences terminal or axillary racemes; flowers bisexual, zygomorphic; sepals 5, imbricate, connate at the base, unequal; petals 3, purplish, contorted, free, clawed; fruits septicidal capsules, with persistent sepals and androecium; seeds 1 per locule, slightly winged at the base. Three spp., two from Colombia to Guyana and E. hirtelloides Mart. in Venezuela, N Brazil, Colombia, in savannas on white sand or rocky areas, or on exposed sandstone outcrops, or in riparian forests; at elevations of 100-1,400(-2,000) m, abundant in north-amazonic white-sand savannas (campinaranas) in Rio Anauá and Upper Rio Negro in Jau national Park in Amazonas state in northern Brazil.

CHRYSOBALANACEAE

§ FULLY CHLOROTROPHIC

Genera/species 27/525–530 Distribution tropical and subtropical regions in the Northern and Southern Hemispheres, including SE U.S.A., SE Africa, with their largest diversity in Central and South America and the Caribbean. Habit usually bisexual (rarely andromonoecious or gynomonoecious), evergreen trees or shrubs. Lenticels abundant. Tress or shrubs; from SE U.S.A., the Caribbean and Mexico to southern Brazil and Paraguay. Originally classified subfamily of the Rosaceae. Almost all taxa are native and endemic except of two taxa which occur outside the Neotropics: Chrysobalanus icaco L.and Parinari excelsa Sabine. Cultivated taxa include Neocarya macrophylla (Sabine) Prance ex F. White and Maranthes corymbosa Blume.

SYSTEMATIC outsiders Kostermanthus (3; W Malesia, Sulawesi), Neocarya (1; tropical W Africa), Bafodeya (1; tropical W Africa), Geobalanus (2; SE U.S.A., Mexico, Central America), Magnistipula (12; tropical Africa, Madagascar), Parastemon (3; Nicobar Islands, Malesia to New Guinea), Grangeria (2; Madagascar, Mauritius, Réunion), Dactyladenia (c 30; tropical Africa), Atuna (8; tropical Asia to Samoa), Maranthes (12; ten species in tropical Africa, one species, M. corymbosa, in tropical Asia and east to islands in the Pacific, one species, M. panamensis, in Nicaragua, Costa Rica and Panamá), Angelesia (3; SE Asia, Malesia), Hunga (11; New Guinea, New Caledonia, the Loyalty Islands), Afrolicania (1; tropical W and C Africa).

1. Acioa Aubl. 6 spp. from Colombia, Peru, Brazil (all species, two endemics) and French Guiana.

2. Chrysobalanus L. Shrubs or trees; leaves suborbicular to lanceolate-elliptic, petiolate, lower leaf surface glabrous or with a few stiff appressed hairs; inflorescence a short, few-flowered raceme of cymules or cymose throughout, or a false raceme or a subsessile fascicle; flowers regular; petals 5, longer than sepals; fruit a small, glabrous drupe. Three spp., C. prancei I.M. Turner from Brazil and Venezuela, C. icaco L. from Florida and Mexico to Caribbean and South America, also in Africa, and C. cuspidatus Griseb. ex Duss in Lesser Antilles.

3. Cordillera Sothers & Prance. Trees; leaves with caducous stipules; inflorescences terminal little-branched panicles; rachis and branches with a few minute appressed hairs. Only one sp., C. platycalyx (Cuatrec.) Sothers & Prance, in higher altitude Andean and Central American forests, from Costa Rica and Panamá to Colombia, Ecuador and Venezuela, from 1,000-2,700 m, and has been recorded as low as 250 m.

4. Couepia Aubl. Trees or shrubs. 62 spp., widely distributed throughout continental Neotropics, 56 in South America, 51 in Brazil, 30 endemics, 14 in several states are rare plants in Brazil, by Plantas Raras do Brasil’s book.

5. Exellodendron Prance. 5 spp., Colombia, Venezuela, Guianas and Brazil (all species, three endemics, E. gracile (Kuhlm.) Prance from Espírito Santo state is a rare plant in Brazil, by Plantas Raras do Brasil’s book).

6. Gaulettia Sothers & Prance. Trees or shrubs; leaf venation reticulate on the abaxial surface and with hair-filled stomatal cavities, leaves often with a whitish bloom on abaxial surface; inflorescence and axis densely ferrugineous (except for G. elata); inflorescence panicles or racemes; calyx lobes acuminate or rounded; petals 5, white. 9 spp., throughout the Amazon rainforest of Peru, Ecuador, Colombia, Venezuela, Brazil (7, Acre, Amazonas, Para, Amapá and Rondonia, one endemic) and the Guianas.

7. Hirtella L. Trees, sometimes flowering as shrubs; leaves lanceolate or lanceolate-elliptic, petiolate, lower surface glabrous or with a few strigose or strigulose hairs; stipules subulate, persistent; inflorescence many-flowered, usually a lax raceme or an elongate thyrse; flowers slightly irregular; petals 5, shorter than sepals; fruit a drupe with scanty mesocarp and smooth. 108 spp. widely distributed throughout Neotropics, including Caribbean (101 in South America), Madagascar and mainland Africa one exclusive each; 68 spp. in Brazil, 26 endemics, 9 in several states are rare plants in Brazil, by Plantas Raras do Brasil’s book. 7 spp. from tropical America (inc. Brazil) are myrmecophites. H. magnifolia Prance (N Brazil, Peru and Colombia) has the largest leaves of any species in this genus, up to 40 cm in fertiles branches.

8. Hymenopus (Benth.) Sothers & Prance. Trees; leaf lower surface glabrous or hirsute, never tomentose, pulverulent, farinaceous nor with stomatal cavities; inflorescences terminal and subterminal panicles or racemose panicles. 28 spp. from Costa Rica, Panamá, Trinidad and Tobago, to northern South America (25) in Colombia, Peru, Ecuador, Bolivia, Venezuela, the Guianas, and in Brazil (19, 4 endemics), mainly in the Amazon rainforest.

9. Leptobalanus (Benth.) Sothers & Prance. Trees; leaf lower surface glabrous, tomentose or lanate, stomatal cavities present or absent. Petiole with or without glands. Inflorescence panicle, racemose panicle or panicle of cymes. 31 spp., from Mexico, Central America and the Caribbean through to northern South America (26), the Guianas, Peru, Ecuador, Bolivia and SE Brazil (15, 3 endemics); L. maguirei (Prance) Sothers & Prance from Mato Grosso and Mato Grosso do Sul states is a rare plant in Brazil, by Plantas Raras do Brasil’s book.

10. Licania Aubl. Trees, shrubs, sometimes geoxylic; leaf lanate, pulverulent-furfuraceous, pulverulent farinaceous, or glabrous, with or without stomatal cavities; petiole with a pair of glands present or not; inflorescences racemose panicles, panicles or spikes; petals absent. 105 spp., two of which imperfectly known, distributed throughout Central and South America (103) and in the Leeward Islands, from Mexico and Costa Rica to SE Brazil (73, 26 endemics); 11 spp. from Amapá, Bahia, São Paulo, Rio de Janeiro, Espírito Santo and Amazonas states are rare plants in Brazil, by Plantas Raras do Brasil’s book

11. Microdesmia (Benth.) Sothers & Prance. Trees; leaf lower surface lanate and deeply reticulate; stomatal cavities present; petals 5. Two spp., M. arborea (Seem.) Sothers & Prance widely distributed from Mexico and Central America to NW South America (Venezuela, Colombia, Bolivia, Peru, Ecuador) and in Brazil in the state of Acre, and M. rigida (Benth.) Sothers & Prance restricted to NE Brazil and Venezuela.

12. Moquilea Aubl. Trees up to 40 m tall; leaf lower surface glabrous or lanate, never with stomatal cavities. Petiole with or without glands; inflorescence panicles, racemes or racemose panicles. 51 spp. distributed from Mexico, Central America, the Leeward and Windward Islands, and throughout South America (45, Guianas, Venezuela, Colombia, Ecuador, Peru, Bolivia, Brazil); the largest leaves of Chrysobalanaceae occurs in M. gentry (Prance) Sothers & Prance of Bajo Calima region in Colombian Chocó. 15 spp. in Brazil, 7 endemics, two of then in Pará and Maranhão states are rare plants in Brazil, by Plantas Raras do Brasil’s book.

13. Parinari Aubl. Shrubs or trees, sometimes with woody rhizomes. 33 spp., 14 in Africa and SE Asia and Australasia, 19 from Central America and throughout South America (all species), and one, P. excelsa Sabine, also in Africa; 16 spp. in Brazil, 5 endemics; two spp., in Bahia state, are rare plants in Brazil, by Plantas Raras do Brasil’s book.

14. Parinariopsis (Huber) Sothers & Prance. Trees; leaf lower surface deeply reticulate and lanate; inflorescence panicles. Only one sp., P. licaniiflora (Sagot) Sothers & Prance, from Guianas, Venezuela, Colombia, Peru and Amazon rainforest of Brazil in Acre, Amazonas and Pará states.

LINEAGE 5 de 6: CLADE SALIC/VIOLOIDS

HUMIRIACEAE

§ FULLY CHLOROTROPHIC

Genera/species 8/51 Distribution southern Mexico, Central America, tropical South America; one species of Sacoglottis in tropical W Africa. Habit bisexual, evergreen trees or shrubs. Young branches angular in cross-section. Often with aromatic juice; wood hard, aromatic, often with balsamic sap, heartwood reddish, alburnum yellow or yellowish.

SYSTEMATIC all genera occur in South America.

1. Duckesia Cuatrec. (inc. Humiriastrum p.p.). Two spp., D. verrucosa (Ducke) Cuatrec. from to northern Brazil and E Peru and D. liesneri K.Wurdack & C.E.Zartman endemic to Venezuela.

2. Endopleura Cuatrec. Tree (up 30 m tall) with a red trunk with wood very hard. Only one sp., E. uchi (Huber) Cuatrec., Bolivia (Pando), Brazil (Amapá, Amazonas, Pará, Rondonia), Guyana, Suriname, Venezuela; 50–620 m elevation, terra firme; their edible and medicinal fruits are known as uxi.

3. Humiria Aubl. 4 spp., Colombia, Venezuela (all species, one endemic), Guyana, Surinam, French Guiana, Peru and Brazil (three spp., none endemics).

4. Humiriastrum Cuatrec. (exc. Duckesia p.p., Vantanea p.p.). 15 spp., all in South America, one up to Costa Rica, reaching French Guiana and Brazil (7, 3 endemics).

5. Hylocarpa Cuatrec. Only one spp., H. heterocarpa (Ducke) Cuatrec, largest tree endemic to the Guiana Shield of Upper Rio Negro, in a very small dense forest in NW Amazonas state in Brazil.

6. Sacoglottis Mart. Trees up to 40 m tall. 11 spp., S. gabonensis (Baill.) Urb. in W Africa (Liberia, Côte d’Ivoire, Nigeria, Cameroun, Gabon and the DRC) and remaining 10 from Costa Rica, Trinidad, St. Vincent, Panamá, Colombia, Venezuela, Guyana, Surinam, French Guiana and Brazil (5, none endemics); S. ovicarpa of Bajo Calima region in Colombian Choco is possibly the largest-fruited Humiriaceae. 8 spp. in South America.

7. Schistostemon Cuatrec. 9 spp., Venezuela, Guyana, Surinam, French Guiana, Peru, northern & C Brazil (4, none endemics), in Amazon rainforest.

8. Vantanea Aubl. (inc. Humiriastrum p.p.) 22 spp., Costa Rica to Bolivia, reaching French Guiana and southern Brazil (13, up to Santa Catarina state, 9 endemics); 20 in South America.

ACHARIACEAE

§ FULLY CHLOROTROPHIC

Genera/species 31/c. 145 Distribution pantropical, with few species in southern Africa. Habit usually dioecious (rarely monoecious), evergreen trees or shrubs (Acharieae consist of climbing herbs); fruits often highly ornamented (tubercles, warts, or thick spines in Lindackeria; thin bristles in Mayna; wings or lacerate wings in Carpotroche).

Achariaceae were traditionally a small South African family of flowering plants. The family is not very diverse in the Neotropics; more genera and spp. occur in tropical Africa and Asia. Four genera and 17 spp. in Brazil. Recent phylogenetic analyses using DNA sequence data affirm that Achariaceae s.s. are nested within the cyanogenic taxa of the former Flacourtiaceae (tribes Pangieae, Lindackerieae [Oncobeae, in part], and Erythrospermeae). Occasionally, the taxa of the former Flacourtiaceae have been segregated from Achariaceae into the family Kiggelariaceae, but more recent studies do not support this exclusion.

Key differences from similar families Achariaceae are a heterogeneous spices, but spices the Neotropical taxa (except for Chiangiodendron) belong to tribe Lindackerieae, which are usually distinctive in having longer and more numerous petals than sepals, linear -elongate anthers, and ornamented fruits; most Achariaceae, including Chiangiodendron, also have flexed petioles with thickened bases and spices.

SYSTEMATIC 4 subtribes, Pangieae (10/29, 7 monotypic genera, Old World, Mexico, Central America), Acharieae (3/3, S Africa) and Erythrospermeae (7/32, Old World) do not occur in South America; outsiders in Lindackerieae, the sole tribe in South America, are Buchnerodendron (2; C and tropical East Africa), Caloncoba (c 10; tropical Africa), Camptostylus (2; tropical W and C Africa), Grandidiera (1; tropical E Africa), Peterodendron (1; tropical E Africa), Poggea (4–6; tropical W and C Africa), Prockiopsis (3; Madagascar), Xylotheca (10–13; E and S Africa).

Key to genera of South American Achariaceae

1. Style 1; fruit surface with warts or coarse, fibrous spines ------------ Lindackeria

1. Styles (2-)3-8(-10); fruit surface smooth, with vertical ridges or wings, or covered with slender bristles - 2

2. Leaf venation Clusia-like ------------ Kuhlmanniodendron

2. Leaf venation not Clusia-like - 3

3. Fruit berry-like, with a thin fruit wall, covered with slender bristles; styles (2-)3-4(-5) ------------ Mayna

3. Fruit capsular, with a thick fibrous fruit wall, with vertical ridges or wings; styles (4-)6-7(-8 ------------ Carpotroche

1. Carpotroche Endl. Trees, fruits winged, sometimes wings lacerate, some species cauliflorous. 12 spp. from South America, one up to Guatemala; 8 in Brazil, two endemics, only C. brasiliensis (Raddi) Endl. in Atlantic coast of Brazil; C. froesiana Sleumer, from Amazonas state, is a rare plant in Brazil, by Plantas Raras do Brasil’s book

2. Kuhlmanniodendron Fiaschi & Groppo. Treelets to trees 7-14 m tall, 2-8 until first ramification; inflorescence botryoid, axillary, usually restricted to the terminal portion of the branches, but sometimes cauliflorous below leaves; flowers unisexual, 1.0–1.5 cm in diameter, axillary to supra-axillary to the bracts; fruits baccate, dry, indehiscent. Two spp. from rainforests of Espírito Santo and cocoa region in Bahia states in Atlantic Forest of E Brazil, found at northern sea-level forests as well as in the more interior montane and submontane forests.

3. Lindackeria C.Presl. Unarmed shrubs or trees; leaves simple; margins usually toothed; petioles sometimes elongate; stipules present; flowers bisexual or male by abortion, in racemes or solitary in axils; sepals 3, imbricate, concave; petals 6–12, imbricate, not much longer than sepals; fruit a globose, woody; seeds 1–3, with cordate cotyledons. 13 spp., 6 in South America, one up to Mexico, 7 remaining in Africa; 5 spp. in Brazil, three endemics.

4. Mayna Aubl. Trees or shrubs, fruits with slender bristles. 7 spp. from South America, two up to Mexico and Central America; three in Brazil, none endemics.

GOUPIACEAE

§ FULLY CHLOROTROPHIC

Genera/species 1/1 Distribution northern tropical South America (Guyana, Suriname, northern Brazil), Central America. Habit bisexual, evergreen trees or shrubs. Goupiacea is a Neotropical family with sole one genus.

SYSTEMATIC a single species in this family.

1. Goupia Aubl. Tree, large, fast-growing tree growing up to 40 m tall with a trunk up to 1.32 m diameter, often buttressed at the base up to 2 m diameter, with rough, silvery grey to reddish-grey bark; aluminium-accumulating, and disagreeable smell. Only one sp., G. glabra Aubl., from Guatemala (one collection in forested area), northern Brazil (very common in northern Mato Grosso and Pará states), Colombia, French Guiana, Guyana, Surinam, Bolivia, Peru, and Venezuela; it is a other names include Cupiuba (Brazil), Saino (Colombia) and Kopi (Suriname); has been used in carpentary and for railway sleepers.

VIOLACEAE

§ FULLY CHLOROTROPHIC

Genera/species 24/c. 750 Distribution cosmopolitan except polar areas. Habit usually bisexual (in Melicytus dioecious), perennial or annual herbs, evergreen or deciduous shrubs or trees (rarely lianas). Herbs, subshrubs, shrubs, treelets, trees (Rinoreocarpos), less frequently lianas (Hybanthopsis). Violaceae a widely distributed family, comprising 23 genera and approximately 900 spp. Roughly half the species of the species belongs to the predominantely herbaceous genus Viola, which are concentrated the northern Hemisphere and mountains areas oat the tropical regions.

SYSTEMATIC two subfamilies, both in South America.

1. SUBFAMILY FUSISPERMOIDEAE (1/3) ‣ a single genus.

1. Fusispermum Hekking. Trees, leaves oblanceolate; flowers in axillary paniculate cymes. Three spp., one in Panamá, and two in Colombia, Ecuador and Peru.

2. SUBFAMILY VIOLOIDEAE (31/c. 750) ‣ 31 monophyletic lineages, 28 named genera and 3 not yet; of 28 named genera, 8 do not occur in New World (Agathea, Melicytus, Isodendrion, Afrohybanthus, Pigea, Decorsella, Allexis and Scyphellandra, with 66 species in total); 5 occur only in North/Central America (Cubelium, Hybanthus, Mayanaea, Mexion and Ixchelia, with 9 spp. in total); remaining 15 are South American.

2. Amphirrhox Spreng. Two spp., A. longifolia (A. St.-Hil.) Spreng., from Central America to C & E razil, and A. latifolia Mart. & Eichler endemic to Guianas.

3. Anchietea A.St.-Hil. Liana or reclining shrubs; fruits as membranaceous, inflated capsules, usually prematurely exposing the strongly flattened seeds to maturation; flowers with nectar spurs. 8 spp., widely distributed in South America, two endemics to Peru, one in the Andes from Colombia to Bolivia, five in extra-Amazonia South America - all of then in Brazil, three endemics, inc. Bahia and Espírito Santo states narrow endemics.

4. Bribria Wahlert & H. E. Ballard. (off Rinorea) Trees to 30 m tall; leaves alternately arranged, inflorescences axillary, lateral to terminal, 1–5-fascicled, thyrsoid, lateral cymules 1–9-flowered, sometimes accompanied by aborted flower buds; pedicels to 7.5 mm long; flowers weakly zygomorphic; buds orbicular, ellipsoid, or ovoid. Three spp., Central America and Venezuela one endemic each, and B. apiculata (Hekking) Wahlert & H. E. Ballard from Central America, Ecuador, Peru, Bolivia and northern Brazil.

5. Calyptrion Gring (inc. Corynostylis). Vines, large, with zygomorphic leaves, flowers with nectar suprs. 4 spp. native to Amazon rainforests, one extending north up to Mexico, mainly grow along seasonally flooded forest (igapó); three spp. in Brazil, none endemics.

6. Gloeospermum Triana & Planch. Trees por treelets; corolla white. 16 spp., from Central America and northern South America (10), only the widely distributed G. sphaerocarpum Triana & Planch. in Brazil.

7. Hekkingia H.E.Ballard & Munzinger. Small tree, to 7 m tall; bark grey, warty; leaves simple; inflorescences long-pedicellate racemes, cauline, produced only at the base of the trunk, with 30 or more flowers; flowers bisexual, yellow, ca. 3 cm long and 2 cm in diameter; fruit an capsule, 3–3.5 cm long and 2–2.5 cm wide, pink or red. Only one sp., H. bordenavei Ballard & Muzinger, from NE French Guiana (near Cayenne).

8. Hybanthopsis Paula-Souza. Twining herbs; leaves serrate, the teeth glandular, stipules 2; flowers solitary, axillary, sepals 5, free, lilac or purplish, sometimes with darker linear markings, solitary, axillary; corolla strongly zygomorphic, petals 5, free, the anterior petal clawed, concave or saccate at base; capsule thinwalled, chartaceous, opening by a single longitudinal slit. Only one sp., H. bahiensis Paula-Souza, endemic to dry seasonal scrubland of NE Brazil (caatinga) of southern and SE Bahia state, E Brazil, near Itaberaba municipality, and is commonly found in disturbed areas such as forest borders and roadsides.

9. Hybanthus thiemei group Herbs with axillary solitary flowers on very long filiform pedicels, strongly zygomorphic corollas with a strongly clawed bottom petal, medially positioned globose staminal glands, and pale yellow minutely alveolate seeds. 5 spp., two in Hispaniola, two in Central America, and H. nanus (A. St.-Hil.) Paula-Souza from Brazil and adjacent Argentina and Uruguay.

10. Leonia Ruiz & Pav. Trees, leaves oblanceolate. 6 spp. from forests of northern South America, only L. glycycarpa Ruiz & Pav. up to Central America; 4 spp. in Brazil, none endemics.

11. Noisettia Kunth. Subwoody, strongly zygomorphic and conspicuously spurred to long-spurred flowers. Only one sp., N. orchidiflora (Rudge) Gingins, Peru to E coast of South America in Brazil and French Guiana.

12. Orthion Standl. & Steyerm. Trees to shrubs, leaves lanceolate or oblonceolate. 6 spp. from Mexico and Central America, O. guatemalense Lundell up to northern Colombia.

13. Paypayrola Aubl. Trees or shrubs; leaves lanceolate or oblonceolate, sometimes cauliflorous. 8 spp., southern Central America, northern and E South America (7); 5 in Brazil, one endemic.

14. Pombalia Vand. Annual or perennial herbs, subshrubs, shrubs or small trees, sometimes with a woody and gemmiferous well-developed root system; plants glabrous or variously hairy; leaves either all alternate, or all opposite, or the basal ones opposite turning alternate towards the apex of the branches, sometimes with very reduced internodes and clustered in short shoots; flowers strongly zygomorphic due a very differentiated anterior petal, bisexual. 43 spp., over New World, only one sp. worldwide; 34 in South America, 23 in Brazil, 9 endemics.

15. Rinoreocarpus Ducke. Small tree with 1-3 fascicled cymose inflorescences in the axils of the leaves, weakly zygomorphic corolla, red-orange petals, placentas with 6–10 ovules, and a coriaceous capsule containing 6-12 seeds. Only one sp., R. ulei (Melch) Ducke, restricted to humid forests of Amazon rainforests of northern South America.

16. Rinorea Aubl. (inc. Phyllanoa, exc. Bribria). 225-275 spp., pantropical, including 46 spp. in Mexico, Central America, northern South America (40) and E coast of South America; 25 spp. in Brazil, 6 endemics (3, from Maranhão, Amazonas and Espírito Santo states are rare plants in Brazil, by Plantas Raras do Brasil’s book); two sects. in New World:

§ sect. Pubiflora ‣ 33 spp., distribution similar to that of sect. Rinorea (see below), but species of this also occur in Mexico, Mesoamerica, Bolivia, Guyana, Surinam, and Trinidad.

§ sect. Rinorea ‣ 13 spp., from Panamá to the greater Amazon rainforest of South America and the Coastal Cordillera in Venezuela, and disjunctly so in the Atlantic Coastal Forests in Espírito Santo and Bahia States of Brazil, where they commonly grow in primary or disturbed rainforest on terra firma or in periodically inundated areas, or in drier savanna, gallery, and seasonally dry forests.

17. Schweiggeria Spreng. Shrub; strongly zygomorphic and conspicuously spurred to long-spurred flowers. Only one sp., S. fruticosa Spreng., endemic to E Brazil.

18. Viola L. Annual or perennial acaulescent or caulescent herbs, often cushions (unique among Malpighiales), shrubs or very rarely treelets; flowers axillary and solitary, rarely in cymes; corolla white to yellow, orange or blue or multicolored with or without yellow throat, strongly zygomorphic, bottom petal slightly to much shorter than others and weakly differentiated, spur scarcely exserted to very long; nectar spurs in some species, inc. Brazilian ones. 664 spp., widely distributed in temperate regions and montane areas in tropics worldwide, 230 in New World, mainly Central America and W South America (153, 110 in Cono Sur); extant chromosome numbers range from diploid 2n = 4 in V. modesta Fenzl. from C Asia, among lowest number known in angiosperms and also found in five other genera unrelated to Viola and Violaceae, to at least 20-ploid 2n = ca. 160 in V. arborescens L. Two subgenera, 31 sections, and 20 subsections.

§ subg. Neoandinium ‣ 139 spp. in 11 section, 7 only from Argentina and Chile, sect. Inconspicuiflos (8) is endemic to Peru, and three remainig widely in W South America.

§ subg. Viola ‣ 20 sections, of which 7 occur in South America: sect. Chilenium (2, Colombia to Patagonia), sect. Leptidium (18, Mexico to Bolivia, Lesser Antilles, SE Brazil), sect. Nosphinium subsect. Mexicanae (10, Mexico to Ecuador), sect. Rubellium (3, endemic to C Chile), sect. Tridens (1, Chile and Argentina), sect. Viola subsect. Rostratae (51, north-temperate, except for V. huidobrii Gay in Argentina and Chile, and one in New Guinea), sect. Xanthidium (2, Peru to N Argentina).

Sect. Leptidium includes the three Brazilian species, all endemic, V. cerasifolia A.St.-Hil., V. gracillima A.St.-Hil. and V. subdimidiata A.St.-Hil., very genetically related and small perennial herbs, endemic of country, mainly in montane forests and grasslands in highlands.

In broader comparisons with the world’s known violet flora the V. lilliputana H. H. Iltis & H. E. Ballard (endemic to C Peru) also appears to be one of the world’s smallest violets (if not the smallest) and surely stands as one of the smallest terrestrial dicots, with the entire aboveground plant body scarcely topping 1.1 cm in height - rivaled in stature only by sporadic and particularly diminutive individuals of certain species in the annual V. parvula Tineo complex (wild pansies of sect. Melanium W. Becker) of SE Europe and adjacent W Asia.

PASSIFLORACEAE

§ FULLY CHLOROTROPHIC

Genera/species 30/c. 924 Distribution tropical, subtropical and warm-temperate regions in the Northern and Southern Hemispheres (Malesherbioideae only W South America), with their highest diversity in tropical America (nearly all Turneroideae) and Africa. Habit usually bisexual (in, e.g., Adenia dioecious), usually perennial or annual herbs, climbing and twining with tendrils, or evergreen shrubs or lianas (rarely small trees). Tendrils simple or branched, axillary, usually consisting of modified pedicels or inflorescences, often oblique relative to the branch and evil-smelling. Some species are xerophytes.

SYSTEMATIC four subfamilies, all in South America, two restricteds.

1. SUBFAMILY MALESHERBIOIDEAE (1/24) ‣ a single genus.

1. Malesherbia Ruiz & Pav. Shrubs, subshrubs, or annual or perennial xerophytic herbs, often cyanogenic, erect or procumbent, from 15 cm to 2 m tall; inflorescences axillary or terminal, usually racemes or leafy panicles, rarely fasciculate or flowers solitary; flowers campanulate or funnel-shaped; fruits stipitate capsules, enclosed by the persistent floral tube; seeds 1 to many, ovoid, exarillate, reticulately ridged, pitted, with an oily endosperm. 27 spp., 11 endemic to Peru in Andean valleys east of Lima and to the C Peruvian Andes, and in the coastal fog zones and lomas vegetation; and 16 in Argentina and Chile, in mediterranean, desert and dry montane habitats of varying aridity; most species prefer rocky, sunny areas inhabited by few other plants.

2. SUBFAMILY TURNEROIDEAE (13/227) ‣ outsiders Erblichia (1; Central America), Mathurina (1; Rodriguez), Arboa (2; Madagascar), Stapfiella (4; tropical Africa), Hyalocalyx (1; SE tropical Africa, Madagascar), Tricliceras (11; tropical and subtropical Africa), Loewia (1; NE tropical Africa), Afroqueta (1; South Africa, Swaziland), Streptopetalum (3–6; tropical and S Africa), Adenoa (1; E Cuba).

Brazil holds the top number of species in Turneroideae (157, 82% of the American species are native, and 73% of them are endemic to this country), the highest number of endemics in being found in the mountains of the states Bahia, Goiás and Minas Gerais; all genera in New World are native, some species are cultivated, as T. subulata Sm. (buttercup, chanana, guarujá); sometimes weedy.

Key to South America genera of Turneroideae

1. Indument with porrect-stellate tector trichomes; corona on petals and sepals ------------ Piriqueta

1. Indument never por-rect-stellate tector trichomes; corona absent, rarely a membranous ligule, but then only on petals - 2

2 Sepals free from each other or almost so ------------ Pibiria

2. Sepals fused to each other, usually by 1/3 of their length - 3

3. petals white with yellow base or, when yellow, pink, or red, the stamens are pilose and only basi-dorsi-ventrally adnate to the floral tube ------------ Oxossia

3. petals usually yellow, when white, the stamens are glabrous and/or with margins adnate to the petal claws, forming nectariferous pockets ------------ Turnera

2. Oxossia L. Rocha. (off Turnera) Subshrubs to shrubs, rarely trees,up to 3m high; leaves medium to large (2–21.5cm long); extrafloral nectaries rarely absent; inflorescences elongated to abbreviated racemes, head-like, spike-like, terminal or lateral, rarely solitary flower; flowers heterostylous,usually small; fruit globose to ovoid; seeds obovoid to pyriform. 17 spp. of Brazil, two up to Venezuela and Guyana, seven of then are rare plants in Brazil, by Plantas Raras do Brasil’s book, mainly in Bahia state; most in rainforests in the Atlantic forest and Amazon rainforest, with some species occurring in dry seasonal scrubland of NE Brazil (caatinga), rocky grasslands (campos rupestres) and the savannas of C Brazil (cerrado).

3. Piriqueta Aubl. Herbs, subshrubs or shrubs, sometimes with xylopodium; leaves rarely with nectaries; flowers solitary, sometimes gathered in cymose inflorescences, hardly ever in an apical raceme; peduncle and pedicel developed, bracteoles generally wanting; capsule granulose, in a few species smooth; seeds obovoid, reticulate. 45 spp. from southern U.S.A to northern Argentina and Uruguay, 43 in South America, 39 in Brazil (27 endemics, 15 of then are rare plants in Brazil, by Plantas Raras do Brasil’s book, 10 only in Bahia), centered in Diamantina Range in center Bahia state.

4. Turnera L. (exc. Oxossia) Herbs, subshrubs, shrubs or treelets with simple hairs; leaves often with nectaries; flowers solitary or gathered in cymose or racemose inflorescences; peduncle absent or developed, often adnate to the petiole (true epiphylly); petals ligulate in a few species; capsule granulose or smooth; seeds obovoid. 138 spp., two in Africa (T. thomasii (Urb.) Story in Kenya and T. oculata Story in Angola and Namibia), 137 in New World, from southern U.S.A. to Argentina, 131 in South America, and centered in Brazil (112, 81 endemics, 26 are rare plants in Brazil, by Plantas Raras do Brasil’s book), extreme highly centered in Minas Gerais, Bahia and Goiás states. 11 series:

§ ser Annularis ‣ 3 spp., Mexico and northern Central America (e.g., T. odorata Richard), but is also found from Venezuela to Brazil in Amazonian savanna and rainforest.

§ ser. Anomalae ‣ 14 spp., South America, mainly in Brazil, SE Bolivia and NW Paraguay. It is mostly found in dry seasonal scrubland of NE Brazil (caatinga) and in Chaco, and wet Amazonian tropical forests, as well as occasionally in the savannas of C Brazil (cerrado) and Atlantic Forest.

§ ser. Cristatae ‣ a single species, T. sidoides L., from montane areas in Bolivia to sea level in Brazil, Uruguay and Argentina.

§ ser. Leiocarpeae ‣ 56 spp., Mexico to Argentina and mostly comprises narrowly distributed species, including many that are known only from the type specimen; most species occur in Brazil (50), mainly in mountain ranges in the states of Minas Gerais and Bahia and in C Brazil in Goiás State, in savannas of C Brazil (cerrado), rocky grasslands (campos rupestres) and in dry seasonal scrubland of NE Brazil (caatinga).

§ ser. Microphyllae ‣ 3 spp., NE region of Brazil and Minas Gerais, mainly in dry seasonal scrubland of NE Brazil (caatinga) and occasionally in rocky grasslands (campos rupestres); however, T. diffusa Willd. ex Schult is widely distributed in dry forests throughout Mesoamerica, the Caribbean Islands, Mexico, and the southern U.S.A. (Texas).

§ ser Papilliferae ‣ two spp., NE Brazil, in dry seasonal scrubland of NE Brazil (caatinga), but T. chamaedrifolia Cambess. is widely distributed and also occurs in areas of the savannas of C Brazil (cerrado), Atlantic Forest, and disturbed vegetation.

§ ser. Rinoreifoliae ‣ 6 spp., from Panamá to Argentina, in seasonally inundated forests, gallery forests dry seasonal scrubland of NE Brazil (caatinga), Atlantic Forest in the South and SE regions of Brazil, and eastern portion of the Andes from southern Ecuador to NE Argentina, as well as the western portion of the savannas of C Brazil (cerrado).

§ ser. Salicifoliae ‣ 6 spp., South America, mainly in wet Amazonian tropical forests, occasionally in the Amazonian savanna, in the Guianas, Suriname, Venezuela, and Brazil.

§ ser. Sessilifoliae ‣ two spp., restricted to Brazil in rocky grasslands (campos rupestres) and savannas of the Espinhaço Range, in Minas Gerais.

§ ser. Stenodyctieae ‣ 10 spp., South America, in wet forests, floodplains and sometimes savannas (a few species), in northern Amazonia, in Colombia, Venezuela, Guiana, Suriname, NW Peru, northern Bolivia, and northern Brazil.

§ ser Turnera ‣ 28 spp., in the New World and Africa (2), is the most geographically widely distributed subseries, and occurs in all tropical biomes including very dry and disturbed habitats.

3. SUBFAMILY PIBIRIOIDEAE (1/1) ‣ a single genus from Guyana. In 1993, a shrub Pibiria flava Maas with yellow flowers reminiscent of but obviously distinct from Turnera was found near Mabura Hill in C Guyana, which was recollected in 2000. The mysterious species belongs to Passifloraceae, in which it is well supported as sister to Turneroideae, and because it lacks several floral characters typically (but not universally) found in this subfamily (heterostylous, fused sepals/petals, adnation of the stamens to the calyx, presence of a corolla and clawed petals), proposed a new subfamily, Pibirioideae, to accommodate it.

5. Pibiria Harms. Shrub with yellow flower. Only one sp., P. flava Maas, endemic to Mabura Hill in C Guyana.

4. SUBFAMILY PASSIFLOROIDEAE (c 16/c 620) ‣ two clades with a status of subfamilies if Passforaceae s.s. is desmembered of Turneraceae and Maleseherbiaceae; Paropsioideae (6/20–27, Tropical Africa, Madagascar, East Malesia) absent. Among Passifloroideae, outsiders are Adenia (90–95; tropical regions in the Old World, with their highest diversity in tropical Africa); Basananthe (37; tropical and S Africa), Deidamia (5; Madagascar), Efulensia (2; tropical Africa), Schlechterina (1; tropical East Africa), Crossostemma (1; tropical Africa).

Distribution Tropical, subtropical and warm-temperate regions in the Northern and Southern Hemispheres, with their highest diversity in tropical America and Africa. Habit Usually bisexual (in, e.g., Adenia dioecious), usually perennial or annual herbs, usually climbing and twining with tendrils, or evergreen shrubs or lianas (rarely small trees). Tendrils simple or branched, axillary, usually consisting of modified pedicels or inflorescences, often oblique relative to the branch. Some species are xerophytes. Only Passiflora in Argentina.

6. Ancistrotryrsus Harms. Woody vines with hooks; flowers in cymose inflorescences. Three spp., from tropical Amazon rainforest up Guiana Shield, all in Brazil, no endemics.

7. Dilkea Mast. Woody vines or small trees, without tendrils, or rarely with a few poorly developed ones; flowers in axillary or terminal glomerules or short-spicate, rarely solitary, hermaphrodite, red or white; sepals 4-5; petals 4-5; fruit globose or ovoid. 12 spp., Panamá to Tropical South America in Amazon rainforest, 4 spp., in Brazil, two endemics; 2 subgenera:

§ subg. Epkia ‣ 6 shrubs or small trees with a strongly rhythmic growth and usually without tendrils, centered in Guiana Shield.

§ subg. Dilkea ‣ 6 spp. of lianas or climbing shrubs, or small shrubs in the case of D. margaritae Cervi, with a continuous growth and thick tendrils trifid at the apex, from Panamá to Amapá (Brazil) and from Amazon rainforest of Bolivia to the Guianas, centered in Ecuador and Peru.

8. Mitostemma Mast. Scandents shrubs; flowers usually in short, terminal or axillary racemes; calyx much reduced; sepals 4; petals 4; corona in 3 series of filaments; stamens 8 or 10, inserted on the floor of the hypanthium near the base of the ovary, free, or united close to the base; gynophore erect; ovary 1-celled, with 4 parietal placentae; styles 4, distinct to the base; stigmas reniform-capitate; fruit ovoid. Three spp., one only in Guyana, M. glaziovii Mast. in Brazil and Guianas, and M. brevifilis Gontsch. endemic to vast area in C & E Brazil.

9. Passiflora L. Mostly perennial climbers, a few small trees up to 20m tall, sometimes with xylopodium, shrubs, herbaceous vines and even annuals; Passiflora possibly contains the greatest variation in leaf blade of any plant genus. 579 spp., 24 from India to New Zealand and 562 in New World, 444 in South America, and 159 in Brazil, 87 endemics; 12 spp. of Passiflora are rare plants in Brazil, by Plantas Raras do Brasil’s book, in over country; 5 subgenera:

§ subg. Astrophea ‣ 62 spp. in five sections, with their center of diversity being in lowland areas of northern South America. 31 spp. in Brazil in section Capreolata (4), Leptopoda (2), Pseudoastrophea (17) and Botryastrophea (8, in series Botryastrophea and Carneae); section Astrophea, which includes all tree members of this genus (9 spp., all but one from Colombia, 4 exclusives), occur only from Costa Rica to Venezuela and Peru.

§ subg. Decaloba ‣ 230 spp. in 8 subsections, Auriculata, Bryonioides, Cieca, Decaloba and Multiflora occur in Brazil, remaining absents: Disemma (all 21 Old World members in subgenus), Hahniopathanthus (5, Mexico to Colombia and Venezuela) and Pterosperma (4, Mexico to Panamá); only 19 spp. in Brazil.

§ subg. Deidamioides ‣ 13 spp. in five sections: Polyanthea (1, Venezuela, Guianas, N Brazil), Mayapathanthus (2, Mexico to Costa Rica), Tetrastylis (4, endemic to Atlantic Forest of Brazil), Tryphostemmatoides (4, Nicaragua to Ecuador) and Deidamioides (1, endemic to Brazil).

§ subg. Passiflora ‣ endemic to New World, 255 spp. in six supersections, Passiflora, Stipulata, Laurifolia, Coccinea and Distephana occur in Brazil, Tacsonia (62-64, all restricted to the high Andes at 1,700 to approximately 4,000 m), the trumpett Passiflora, Venezuela to Bolivia, and comprises mainly species with showy flowers, is absent; within this group; P. antioquiensis H.Karst, endemic to Colombia, is possibly the largest flower of genus.

Passiflora supersect. Tacsonia is characterized by several morphological traits, suggesting that the group might be monophyletic, although this has not really been tested. The best-sampled phylogeny so far included only seven Tacsonia species, which formed a clade. While most species of the supersection have hummingbird-adapted flowers, the longest tubed-flowers are restricted to 37 species pollinated by the sword-billed hummingbird, Ensifera ensifera, whereas the 19 species with shorter tubed red flowers (hypanthium 1–3 cm long) are pollinated by shorter billed hummingbirds. Bats pollinate another seven species that have greenish or white flowers. Like most Passiflora, Tacsonia species are self-incompatible and depend on cross-pollination to set seed.

§ subg. Tetrapathea ‣ 3 spp., Australasia to New Zealand.

LACISTEMACEAE

§ FULLY CHLOROTROPHIC

Genera/species 2/15–25 Distribution Caribbean (Jamaica), southern Mexico southwards to SE Brazil, Paraguay, Uruguay and northern Argentina. Habit bisexual, evergreen trees or shrubs. Use Timber.

SYSTEMATIC both genera occur in South America.

1. Lacistema Sw. Small trees. 12 spp. from forests of South America (mainly Amazon rainforest), L. aggregatum Rusby is a fairly common and widely distributed species in the Neotropics, from Mexico to SE Brazil, Uruguay, and NE Argentina; 11 in Brazil (exception is one from Ecuador to Bolivia), 5 endemics.

2. Lozania Mutis ex Caldas. 5 spp., Costa Rica to Peru and Amazonas state in northern Brazil (only L. klugii (Mansf.) Mansf., no endemic).

SALICACEAE

§ FULLY CHLOROTROPHIC

Genera/species 52/1,080-1,090 Distribution mainly tropical and subtropical areas, some genera in temperate regions, few in Australia, absent from New Zealand; Salix and Populus mainly in temperate regions and some species of Salix only in arctic areas in the Northern Hemisphere. Habit usually bisexual (sometimes dioecious, e.g., Salix, Populus, Chosenia, and Scyphostegia), evergreen or deciduous trees or shrubs (some genera climbing; few species of Salix dwarf shrubs); Casearia sometimes with phyllanthoid branching: orthotropic branches with reduced spiral leaves, and plagiotropic branches sylleptic with normal leaves distichous).Salicaceae were traditionally a mostly temperate family consisting of 2 genera, Populus (poplars, cottonwoods) and Salix (willows, vime in Portuguese). In general, tropical members of the family Salicaceae have few economic uses. The mostly temperate genus Salix is one of the early sources of aspirin precursors.

Use Ornamental plants, matches (Populus), baskets (withies from Salix), bioenergy (Salix), timber, protective plantations (against wind and erosion), fruits (Dovyalis), medicinal plants.Salix species (willows) are very important in management of ecosystems such as prevention of erosion, bioremediation of soil and they provide a specialised habitat for other organisms. Both Salix and Populus are potential biofuel sources. In the Neotropics, genera like Casearia and Macrohasseltia are occasionally used for wood, and the genus Ryania has toxic compounds used in poisons and insecticides. Despite their lack of use in the Neotropics, Salicaceae are common elements of tropical forests.

18 genera and about 100 spp. in Brazil.

SYSTEMATIC clades Scyphostegieae (2/2, N and C Borneo, Yunnan) and Scolopieae (5/43, tropical regions in the Old World) do not occur in South America.

1. SUBFAMILY SAMYDEAE (14/c 230) ‣ outsiders Ophiobotrys (1; tropical W and C Africa), Osmelia (4; Sri Lanka, Malesia), Pseudosmelia (1; Moluccas), Trichostephanus (2; tropical W and C Africa).

1. Casearia Jacq. (inc. Laetia p.p., Hecatostemon and Zuelania, exc. Piparea, Irenodendron) Shrubs to trees, sometimes basal burls. 239 spp. worldwide, 97 in New World, in Antilles, Mexico, Central and South America (75); 52 spp. in Brazil, 19 endemics, C. neblinae Sleumer is a rare plant in Brazil, by Plantas Raras do Brasil’s book, endemic to region of Mount Neblina.

2. Euceraea Mart. Three spp., endemic to the Guiana Shield, E. nitida Mart. in Colombia, Venezuela, Guyana, Suriname, Brazil, at 100-2,100 m elevation range; E. rheophytica P.E. Berry & M.E. Olson occur only in deep canyon in Venezuelan side of Mount Neblina, and E. sleumeriana Steyerm. & Maguire also only in Venezuela.

3. Irenodendron M.H. Alford & Dement. (off Caesarea) Unarmed shrubs or trees, to 20 m tall and 30 cm dbh; leaves simple, alternate, pinnately veined; inﬂorescences axillary, rarely supra-axillary, fascicles of 3–10 ﬂowers; flowers bisexual, sepals (4–)5(–6), imbricate, pale green to white to pink, petals absent; fruits capsules; seeds 1–few, arillate. Three spp., Amazon rainforest of Brazil (all spp., none endemics), Colombia, Guyana, Peru, and Venezuela; mixed evergreen forest, terra ﬁrme, often along streams, north-amazonic white-sand savannas (campinaranas), and Paciﬁc of Colombia, collection from Colombia possibly a new species.

4. Lunania Hook. 14 spp., southern Mexico, Central America, Cuba, Jamaica, only L. parviflora Spruce ex Benth. in South America, in over Amazon rainforest.

5. Neoptychocarpus Buchheim. Three spp., Colombia to Guianas and northern Brazil (two, none endemics).

6. Piparea Aubl. (off Caesarea). Trees. Three spp. from Mexico to Bolivia and northern Brazil (all, none endemics).

7. Ryania Vahl. 8 spp. from South America, one up to Central America; 7 spp. in Brazil (the exception in one species from Colombia and Venezuela), one endemic.

8. Tetrathylacium Poepp. 6 spp., endemics in Ecuador (3) and Peru (1), T. johansenii Standl. from Central America to Colombia, and T. macrophyllum Poepp. from Central America to Bolivia and Brazil. T. macrophyllum Poepp. from Costa Rica to Brazil is a myrmecophite.

2. SUBFAMILY PROCKIEAE (9/67) ‣ outsider Hasseltiopsis (1; Central America).

9. Abatia Ruiz & Pav. Small to médium-sized shrubs, many stamens. 10 spp., mountain regions in tropical America, nine in South America; 4 spp. in Brazil, three endemics (all are rare plants in Brazil, by Plantas Raras do Brasil’s book, in Minas Gerais and Rio de Janeiro states), A. angeliana M.H. Alford up to Cono Sur.

10. Azara Ruiz & Pav. Shrubs to small trees, sometimes with lignotubers. 10 spp., all confined to Chile, Argentina and Uruguay but A. salicifolia Griseb. up to Bolivia and A. uruguayensis (Speg.) Sleumer up to southern Brazil.

11. Banara Aubl. 33 spp., tropical America, 19 in South America, 10 spp. in Brazil, only B. trinitatis Sleumer endemic, restricted of Trindad Is. in Atlantic Ocean, a rare plant in Brazil, by Plantas Raras do Brasil’s book.

12. Hasseltia Kunth. 6 spp. from southern Mexico to Brazil and Bolivia, 5 in South America, Venezuela, Colombia and Peru one endemic each, only the widely distributed H. floribunda Kunth in Brazil.

13. Neosprucea Sleumer. 10 spp., one in center Guyana, and remaining nine from Colombia to Peru.

14. Macrothumia M.H.Alford. Trees with 30 m tall; leaves simple, chartaceous, alternate; inflorescence a terminal, congested, fascicle or umbel-like raceme; flowers creamy-yellow, 1 cm diameter; fruits capsular, globose. Only one sp., M. kuhlmannii (Sleumer) M.H.Alford, from Atlantic forest of Alagoas, Bahia, Minas Gerais and Espírito Santo states in E Brazil.

15. Pineda Ruiz & Pav. Two spp. from Andes of Ecuador, Peru and Bolivia.

16. Prockia L. 7 spp., southern Mexico to northern Argentina, three in South America, Venezuela and Peru one endemic each, and the widely distributed P. crucis P. Browne ex L., unique in Brazil.

3. SUBFAMILY SALICEAE (12/545–550) ‣ oustiders Populus (35–40; Northern Hemisphere, P. ilicifolia, in East Africa), Bennettiodendron (7; S China, tropical Asia), Olmediella (1; C America), Idesia (1; China, Japan), Carrierea (4; S and SW China, SE Asia), Itoa (1; S China, tropical Asia), Lasiochlamys (11; New Caledonia), Ludia (27; tropical E Africa, Madagascar, Mascarene slands), Poliothyrsis (1; China), Tisonia (14; Madagascar).

17. Macrohasseltia L.O. Williams. Only one sp., M. macroterantha (Standl. & L.O. Williams) L.O. Williams, from southern Belize to NW Colombia.

18. Salix L. 450 spp., distributed across the temperate to arctic regions of the Northern Hemisphere, entering tropical regions along montane ranges; 275 are found in China, 107 in the former Soviet Union, 65 in Europe, and 113 in North America north of Mexico. The only species native to South America is S. humboldtiana Willd., a small to medium-sized tree, which grows wild along watercourses in from Mexico to Argentina and southern Brazil (in several many ecossystems), Chile, Uruguay and Caribbean.

4. SUBFAMILY HOMALIEAE (8/c 200) ‣ outsiders Bartholomaea (3; Central America), Bivinia (1; tropical East Africa, Madagascar), Byrsanthus (1; tropical W Africa), Calantica (10; Madagascar), Dissomeria (2; tropical Africa), Neopringlea (3; southern Mexico, Guatemala), Bembicia (1; Madagascar).

19. Homalium Jacq. c 180 spp., tropical and subtropical regions on both hemispheres, three in the New World, one endemic to Mexico and two remaining widely distributed in Neotropics.

5. SUBFAMILY ONCOBEAE (6/120–140) ‣ outsiders Oncoba (c 30; tropical and S Africa, Arabian Peninsula), Flacourtia (22; tropical and S Africa, Madagascar, SE Asia, Malesia to Fiji), Dovyalis (17; tropical and S Africa, Madagascar, tropical Asia from Sri Lanka to New Guinea), Trimeria (2; tropical East Africa to S Africa).

20. Pleuranthodendron L. O. Williams. Only one sp., P. lindenii (Turcz.) Sleumer, from Mexico, Central America, Colombia, Ecuador, Peru and northern Brazil.

21. Xylosma G. Forst. 85-100 spp., E and SE Asia, Malesia to New Guinea, eastern Queensland, New Caledonia, Vanuatu, Polynesia, Guam, and 58 spp. in New World, Central America, the Caribbean including Hispaniola, northern South America (30); 10 spp. in Brazil, one endemic.

LINEAGE 6 de 6: EUPHORBIIDS REALM

PHYLLANTHACEAE

§ FULLY CHLOROTROPHIC

Genera/species 60/1,950-2,000 Distribution mainly in tropical regions, with their largest diversity in Malesia. Habit usually monoecious or dioecious (rarely bisexual), usually evergreen (sometimes deciduous) trees or shrubs (sometimes climbing), perennial or annual herbs. Some species are xeromorphic shrubs or herbs with ericoid leaves (some species of Phyllanthus have phyllocladia, others are limnic and floating); branches often dimorphic.

Phyllanthaceae are represented by 19 genera in South America, and occur in several vegetation types, especially in rainforest, savanna and associated ecosystems. Only Margaritaria and Phyllanthus in Argentina. A number of taxa are regionally cultivated for their fleshy edible fruits [e.g., P. acidus (L.) Skeels, P. emblica L., Baccaurea spp., Antidesma spp.].

SYSTEMATIC two subfamilies, both in South America.

UNPLACED PHYLLANTHACEAE ‣ Ashtonia (2), Chonocentrum (1).

1. Chonocentrum Pierre ex Pax & Hoffm. Only one sp., C. cyathophorum (Müll. Arg.) Pax & Hoffm., endemic to the Guiana Shield of Upper Rio Negro in northern Amazonas state in Brazil, in areas with 100 m elevation range.

1. SUBFAMILY PHYLLANTHOIDEAE (30/1,500-1,550) ‣ outsiders Notoleptopus (1; Malesia to New Guinea and N Australia), Poranthera (14; Australia, Tasmania, New Zealand), Pseudophyllanthus (1; S Africa), Phyllanthopsis (2; Texas, Mexico), Actephila (c 35; S China, tropical Asia to Queensland, New South Wales and islands in the Pacific), Leptopus (10; the Caucasus, Iran and Himalayas to China and SE Asia), Heywoodia (1; tropical Africa, KwaZulu-Natal, E Cape), Chascotheca (1; Cuba, Hispaniola), Dicoelia (1; W Malesia), Wielandia (13; Kenya, Madagascar, the Comoros, Seychelles), Lingelsheimia (6; C Africa to Tanzania, Madagascar), Plagiocladus (1; C Africa), Richeriella (1; India, Hainan, Thailand, W and C Malesia), Heterosavia (4; Florida Keys, C America, Cuba, Bahamas, Grand Cayman), Securinega (5; Madagascar, Mascarene Islands), Lachnostylis (3; Cape), Keayodendron (1; Ivory Coast to Cameroon), Cleistanthus (c 140; tropical regions in the Old World), Pseudolachnostylis (1; tropical and S Africa), Pentabrachion (1; Cameroon, Gabon), Bridelia (c 50; tropical Africa and Madagascar to N Australia and islands in the Pacific).

2. Amanoa Aublet. Trees, glabrous in all parts; leaves alternate, petiolate; flowers in solitary, subglobose, sessile, terminal and axillary heads surrounded by coriaceous bracts, capsule separating into 2-valved cocci; seeds solitary by abortion, shining, testa crustaceous; endosperm thin or absent. 17 spp., through the tropics of Africa (2) and the Americas (15), mainly in the northern portion of South America (Brazil, Colombia, Ecuador, Venezuela and French Guiana) or along the rainforests, montane and gallery forests and savanna, one up to Belize and one restricted of Lesser Antilles. Brazil has 11 spp., two endemics.

3. Andrachne L. 22 spp., with a primarily Tethyan distribution from Persia through Mediterranean to the Caribbean, two in New World, A. brittonii Urb. in Caribbean, and A. microphylla (Lam.) Baill. disjunct from Baja California in Mexico and pacific coast of Peru.

4. Astrocasia B.L.Rob & Greenm. 5 spp., three endemics to Mexico, A. jacobinensis (Mull. Arg.) G. L. Webster from E Brazil disjunct in Bolivia (its transfer from Phyllanthus is supported by genetic data) and A. tremula (Griseb.) G.L. Webster, distributed from Mexico and Cuba to Bolivia and E Brazil, absent in Guianas; its species grow mainly in savanna vegetation in open areas.

5. Cicca L. (off Phyllanthus) Herbs, shrubs or trees, monoecious or dioecious, branching (non-)phyllanthoid, branchlets (bi)pinnatiform (sometimes further ramified), rarely opposite (subg. Menarda (Comm. ex A.Juss.) R.W.Bouman), sometimes specialized in vegetative and floriferous branchlets. 42 spp. in 4 subgenera, all from Old World except Cicca subg. Cicca sect. Cicca, with 3 spp., C. acida (L.) Merrill of a unknown origin, C. chacoensis (Morong) R.W.Bouman in Bolivia, Brazil and Southern Cone, and C. elsiae (Urban) R.W.Bouman from Mexico, Central America, Trinidad & Tobago, Venezuela, Guianas, Colombia, Brazil.

6. Croizatia Steyerm. Shrubs. 5 spp. from Colombia, Venezuela and Ecuador, one up to Panamá.

7. Discocarpus Klotzsch. 5 spp. from Brazil (all species, one endemic), French Guiana, Guyana, Suriname, Colombia, Peru, and Venezuela, distributed through the rainforest and swamp forest of Amazon rainforest up to semideciduous forests of eastern (Bahia, Maranhão) and central Brazil (Goiás).

8. Flueggea Willd. 16 spp., widely distributed in tropical to warm temperate regions, only three in New World: F. acidoton Griseb. (Cuba), F. elliptica (Spreng.) Baill. (Ecuador), and F. schuechiana (Müll.Arg.) G.L.Webster in Bahia and Pernambuco states in NE Brazil, where they grow in the edges of secondary forest.

9. Gonatogyne Klotzsch ex Müll. Arg. Only one sp., G. brasiliensis (Baill.) Müll. Arg., of SE Brazil.

10. Margaritaria L. f. 13 spp., distributed thorough tropical America, Africa, Asia, and Australia; 4 spp. in New World, three restricted of Caribbean and M. nobilis L.f., the most widely distributed species of genus in the Neotropics in almost all countries, growing in seasonal, rain and gallery forest.

11. Meineckia Baill. Monoecious or dioecious, glabrous shrubs or subherbaceous undershrubs; leaves alternate; flowers axillary, in unisexual or bisexual fascicles; male flowers on long, very slender pedicels, sepals 5, imbricate, petals 0; female flowers on very long, slender pedicels distinctly articulate near base; sepals and disc as in male flowers; fruits 3-lobed; seeds often 1 per locule. 30 spp., almost all in Old World except three in New World, two from Mexico to Honduras and M. neogranatensis (Müll. Arg.) G.L. Webster, highly disjunct in Nicaragua, Colombia and E Brazil.

12. Moeroris Raf. (off Phyllanthus) Herbs, (sub)shrubs or small trees, monoecious or dioecious, branching (sub-)phyllanthoid (nonphyllanthoid in M. arenaria (A.Gray) R.W.Bouman), branchlets pinnatiform (rarely bi-pinnatiform in subgenus Tenellanthus. 197 spp. in three subgenera.

§ sub. Moeroris ‣ 179 spp., 154 in Old World, 15 endemics to Caribbean, M. stipulata Raf. and M. caribaea (Urban) R.W.Bouman widely in tropical New World, two endemics to Mexico; M. minutula R.W.Bouman, M. microphylla (Kunth) R.W.Bouman, M. lindbergii (Müller) R.W.Bouman wider in South America, one in West Indies up to North America; M. vichadensis (Croizat) R.W.Bouman endemic to Colombia; and M. leptophylla (Müller) R.W.Bouman endemic to Brazil.

§ subg. Swartziani ‣ 4 spp., M. abnormis (Baillon) R.W.Bouman endemic to North America, and 3 in Africa.

§ subg. Tenellanthus ‣ 16 spp., Old World.

13. Phyllanthus L. (exc. Cicca, Moeroris) Small herbs to treelets, filocladoid in some spp., also aquatic, microphyllous, palm-like, with phyllanthoid branching (often absent), staminate flowers with a generally segmented glandular disk, pistillate flowers with an entire glandular disk, and diversely sculptured seed coats; it has a diversity of growth forms (annual, arborescent, aquatic, pachycaulous, and phyllocladous), chromosome numbers, and pollen types rivalling that of any genus of flowering plants. 208 spp., New World with some weeds in Old World, from the SE of the U.S.A. to Argentina, including the Caribbean and Chile, c. 146 in South America, 93 in Brazil, 68 endemics, in several ecosystems, being more frequent in open vegetation, e.g. dry savannas, dry seasonal scrubland of NE Brazil (caatinga), and rocky grasslands, or in disturbed areas, some widely distributed over Neotropics, but majority restricted. 5 subgenera, listed below:

§ sect. Ciccopsis (unplaced subgenus) ‣ one sp., endemic to Cuba.

§ sect. Omphacodes (unplaced subgenus) ‣ one sp., endemic to Caribbean.

§ subg. Ciccastrum ‣ two spp., over tropical New World.

§ subg. Conami ‣ 12 spp., endemic to tropical America, with three sections: Apolepsis (1, tropical America), Calodyction (1, Mexico and Guatemala) and Conami (10, tropical America).

§ subg. Microglochidium ‣ 23 spp., tropical America.

§ subg. Phyllanthus ‣ 5 sections, 72 spp.

§ sect. Antipodanthus ‣ 6 spp. from Brazil, one up to Bolivia and Paraguay.

§ sect. Choretropsis ‣ two subsections, 13 spp., all endemics to Brazil; the absence of leaves has led to the formation of palisade-like tissue in the entire stem and branchlets, and sometimes to the enlargement of these organs to supply the photosynthetic requirements. Such modified branchlets are the main distinctive characteristics of the species in sections Xylophylla (L.) Baill. (Caribbean) and Choretropsis Müell. (Brazil). The diversity of branchlet shapes in the Brazilian species occur at two subsections in section Choretropsis: subsect. Choretropsis (P. goianensis L.J.M. Santiago, P. choretroides Müll. Arg., P. spartioides Pax & K. Hoffm.) characterized by cylindrical branchlets, and subsect. Applanata (P. angustissimus Müll. Arg., P. klotzschianus Müll. Arg., P. edmundoi L.J.M. Santiago, P. flagelliformis Müll. Arg., and P. gladiatus Müll. Arg.) with develops flattened branchlets.

o subsections Applanata ‣ 8 spp., all endemics to Brazil.

o subsect. Choretropsis ‣ 5 spp., all endemics to Brazil.

§ sect Loxopodium ‣ 8 spp., tropical America. P. fluitans Benth. ex Mull.Arg., from this section, it’s the unique aquatic plant of Euphorbiaceae and related families known, growing in rivers and lagoons in tropical areas from Mexico to Brazil; it is a free-floating aquatic perennial that grows to 2” tall and spreads to 5” or more, sometimes a aggressive weed in temperate regions, such as Florida peninsula.

§ sect. Phyllanthus ‣ three subsections, 44 spp., 3 of them, all endemics to Brazil, unplaced at subsection level.

o subsect. Almadensis ‣ only one sp., P. almadensis Muller, endemic to Brazil.

o subsect. Clausseniani ‣ 31 spp., tropical South America.

o subsect. Phyllanthus ‣ 9 spp., P. niruri L. wider in tropical America, remaining endemics in Brazil, Bolivia and Trinidad & Tobago.

§ sect. Pityrocladus ‣ 6 spp., South and Central America.

§ subg. Xylophylla ‣ 14 sections, 92 spp., and 4 unplaced at section level, inc. P. bahianus Mull. Arg.

§ sect. Adianthoides ‣ 7 spp., tropical America.

§ sect. Asterandra ‣ two spp., wider in tropcal America.

§ sect. Callitrichoides ‣ one sp., endemic to Cuba.

§ sect. Cyclathera ‣ 4 spp., Caribbean.

§ sect. Diplocicca ‣ only one sp., P. octomerus Mull. Arg., endemic to Brazil.

§ sect. Elutanthos ‣ 24 spp., widely distributed in tropical America.

§ sect. Epistylium ‣ 3 spp., Caribbean.

§ sect. Glyptothamnus ‣ one sp., Cuba.

§ sect. Hylaeanthus ‣ 8 spp., tropical America.

§ sect. Orbicularia ‣ 13 spp., Caribbean.

§ sect. Oxalistylis ‣ only one sp., P. salviifolius Kunth, from Costa Rica to Peru and Venezuela.

§ sect. Thamnocharis ‣ 3 spp., endemics to Cuba.

§ sect. Williamiandra ‣ 4 spp., Caribbean.

§ sect. Xylophylla ‣ 16 spp., Caribbean.

14. Savia Willd. Two spp., S. sessiliflora (Sw.) Willd. occur is highly disjunct in Mexico, northern Venezuela, Caribbean and Brazil and S. dictyocarpa Müll. Arg. occur in SE Brazil and adjacent Bolivia, frequently in the Atlantic Forest.

15. Tacarcuna Huft. Trees; the previously unrecorded high stamen number in T. amanoifolia Huft. (14–19, among the highest in Phyllanthaceae) suggests this taxon is derived. Three spp. from Venezuela, Colombia and Peru, one reaching to Panamá.

2. SUBFAMILY ANTIDESMATOIDEAE (20/435-440) ‣ outsiders Bischofia (2; China, India to E Asia, Melanesia and Polynesia to Samoa and Niue), Spondianthus (1; tropical E and C Africa), Uapaca (c 50; tropical Africa, Madagascar), Protomegabaria (3; tropical W and C Africa), Aporosa (75–80; tropical Asia to Solomon Islands), Maesobotrya (c 20; tropical Africa), Baccaurea (c 45; tropical Asia, islands in W Pacific), Distichirhops (3; Borneo, New Guinea), Nothobaccaurea (2; Solomon Islands, Fiji), Leptonema (2; Madagascar), Apodiscus (1; tropical W Africa), Martretia (1; tropical W and C Africa), Antidesma (c 170; tropical and subtropical regions in the Old World), Thecacoris (c 25; tropical Africa, Madagascar), Hymenocardia (6; tropical and S Africa, SE Asia, Sumatra).

16. Celianella Jabl. Only one sp., C. montana Jabl., endemic to the Guiana Shield of Venezuela (Amazonas, Bolivar) where it inhabits scrub-savanna on tepui summits, 1,000-1,500 m elevation range.

17. Didymocistus Kuhlmann. Only one sp., D. chrysadenius Kuhlm., of Amazonas state in northern Brazil, Colombia and Peru.

18. Hieronyma Allemão. Trees up to 30 m tall. 27 spp., New World, 17 in South America, two in Brazil, both are widely distributed in the rainforest and submontane forest of the over Neotropics.

19. Jablonskia Webster. Only one sp., J. congesta G.L. Webster, confined to Amazon rainforest of Guianas to Peru and Brazil; largely associated with watercourses or within.

20. Richeria Vahl. Shrubs or trees (2-25 m tall), dioecious; leaves simple, alternate, chartaceous or coriaceous; inflorescence in spikes or racemes, cauliflorous, sometimes axillary, solitary or adensed; fruits capsule. 4 spp., two scattered from Costa Rica to Ecuador, R. dressleri G. L. Webester in Costa Rica, Panamá, Ecuador and in a small-forested area in Ducke Reserve in Amazonas state, Brazil; and the widely distributed R. grandis Vahl; mainly in riparian forest of the over Neotropics.

PICRODENDRACEAE

§ FULLY CHLOROTROPHIC

Genera/species 25/90–94 Distribution mainly tropical regions, with their largest diversity in New Guinea, Australia, New Caledonia and Madagascar. Habit usually monoecious or dioecious (rarely bisexual), evergreen or deciduous trees or shrubs, perennial herbs; many species are stem succulents or twining.

The Picrodendraceae are represented by 5 genera in the Neotropics, all endemic with quite restricted distribution, occurring in several lowland vegetation types including humid evergreen forests, dry deciduous forests or arid scrubland. Picrodendron is endemic to Caribbean. Picrodendron (embryos with ruminate cotyledons, bitter taste of bark etc.) are very unusual by the cited characters, but seem to be part of the Picrodendraceae, supported by the characteristic pollen. Podocalyx is still insufficiently known. Therefore the exact circumscription of the family is not completely settled.

SYSTEMATIC outsiders are Tetracoccus (5; SW U.S.A., NW Mexico), Hyaenanche (1; W Cape), Austrobuxus (c 28; W Malesia, E Queensland, New Caledonia, Fiji), Dissiliaria (6; Queensland), Sankowskya (1; Queensland), Whyanbeelia (1; Queensland), Choriceras (2; New Guinea, Northern Territory, Queensland), Petalostigma (5; Papua New Guinea, Queensland, New South Wales, Northern Territory, W Australia), Kairothamnus (1; New Guinea), Scagea (2; New Caledonia), Neoroepera (2; Queensland), Micrantheum (4; South Australia to Queensland, Tasmania), Stachystemon (9; W Australia), Pseudanthus (9; Australia, Tasmania), Picrodendron (1; the Caribbean), Oldfieldia (4; tropical Africa), Aristogeitonia (7; tropical Africa, Madagascar), Mischodon (1; India, Sri Lanka), Voatamalo (2; Madagascar), Androstachys (1; tropical Africa, Madagascar).

1. Parodiodendron Hunz. Only one sp., P. marginivillosum (Speg.) Hunz., in drier forests of Bolivia and NW Argentina.

2. Piranhea Baill. 4 spp., one endemic to Mexico and three in Amazon rainforest of SE Colombia, Venezuela and northern to E Brazil (all three species, P. securinega Radcl.-Sm. & J.A. Ratter endemic).

3. Podocalyx Klotzsch. Shrubs to treelts, dioicous; leaves eliptic, membranaceous; inflorescence terminal, in peseduo-spiciforme. Only one sp., P. loranthoides Klotzsch, restricted to Guianas, Colombia, Peru, Venezuela and northern Brazil, with a notorious collection (unique extra-amazonic) of Alagoas state in E Brazil.

LINACEAE

§ FULLY CHLOROTROPHIC

Genera/species 10 (inc. Cliococca)/c. 265 Distribution Linoideae: cosmopolitan except polar areas, mostly temperate and subtropical regions; Hugonieae: pantropical. Habit bisexual, evergreen trees, shrubs, suffrutices or lianas, perennial, biennial or annual herbs. Some species are xerophytes. Hugonieae are often lianas with tendrils formed by basal branches of the inflorescences.

Use Ornamental plants (Linum, Reinwardtia), textile and paper, seed oils, timber, medicinal plants, fruits (Hugonia). Moreover, Linum flowers are more showy with convolute yellow, blue or red petals, which are larger than the sepals.

SYSTEMATIC two tribes, both in South America.

1. TRIBE HUGONIEAE (3/50-57) ‣ outsider Hugonia (c 40; Africa and Madagascar to New Caledonia and Fiji).

1. Hebepetalum Benth. Trees, shrubs or often lianas with branch tendrils. Three spp. in Amazon rainforest of South America, all in Brazil, none endemics.

2. Roucheria Planch. 8 spp. from Amazon rainforest of South America, mainly in Colombia (6), 3 in Brazil (none endemics), one up to Central America.

2. TRIBE LINEAE (4-8/c 200) ‣ outsiders of Linum s.l: Anisadenia (2–3; Himalayas to C China and N Thailand), Reinwardtia (1; N Pakistan, N India, S Himalayas, China, SE Asia), Tirpitzia (2–3; SW China, northern Thailand, Vietnam).

3. Cliococca Bab. Herbs, perennial, glabrous with extensevely branched underground rootstock; leaves alternate, linear, 1-nerved; solitary flowers terminal in branches, sepals 5, petals 5; fruit a capsule, 10 one-seeded segments. Only one spp., C. selaginoides (Lam.) Rogers & Mildner, from southern Brazil, Bolivia, Uruguay, Argentina and Chile, in grasslands.

4. Linum L. Annual or perennial herbs; leaves alternate, opposite or whorled, sessile, simple, entire; stipular glands present or not so; flowers in terminal cymes or sometimes in racemose or paniculate inflorescences. ca. 200 spp., subcosmopolitan, 65 in the New World tropics and subtropics; 18 spp. in South America, all from Ecuador to southern Brazil (5, one endemic) and Chile.

IXONANTHACEAE

§ FULLY CHLOROTROPHIC

Genera/species 4/c. 22 Distribution Himalaya, NE India, S China, SE Asia, Malesia, New Guinea, NE South America, tropical Africa. Habit usually bisexual (rarely unisexual), evergreen trees or shrubs. Trees or shrubs; family with five genera and about 30 spp.; Ixonanthes (3) occurs from China to New Guinea. Two genera in New World.

SYSTEMATIC outsider Ixonanthes (5; NE India, E Himalayas, S China, SE Asia, Malesia to New Guinea).

1. Cyrillopsis Kuhlm. Two spp. from Venezuela to northern Brazil (only C. paraensis Kuhlm.) and French Guiana, 100-1,100 m elevation range.

2. Ochthocosmus Benth. Nine spp., two in tropical Africa and 7 spp. in New World, 6 endemics to Guiana Shield in E Colombia, Guyana, French Guiana, Venezuela, Brazil (three spp., no endemics), and two extending to southern Amazon rainforest of Brazil up to Bolivia.

PERACEAE

§ FULLY CHLOROTROPHIC

Genera/species 5/123-140 Distribution pantropical. Habit usually dioecious (sometimes monoecious, especially in Pera), evergreen trees or shrubs (sometimes perennial herbs, more or less lignified at base).

SYSTEMATIC outsiders Clutia (c 75; tropical and subtropical Africa to South Africa, Arabian Peninsula), Trigonopleura (3; W and C Malesia).

1. Chaetocarpus Thwaites. Trees or shrubs, dioecious, with a hirsute and sericeous indumentum; leaves alternate, simple, entire, penninerved, eglandular; flowers in axillary fascicles, the male fascicles many-flowered, the female fascicles few-flowered; flowers small; petals absent; fruit ellipsoid-subglobose, dehiscing into bivalve cocci; seeds compressed-ovoid. 12-15 spp., pantropical (1 African), 9 of New World, in the Antilles and from Venezuela to Bolivia, 4 in northern South America, two in Brazil, none endemics.

2. Pera Mutis. Trees or shrubs, cymose inflorescence, which can be either unisexual or bissexual with flowers minute enveloped by a fragrant, somewhat globose, showy (yellow, cream, white, and sometimes red) involucral bract. 33 spp., from southern Mexico to the Antilles and Bolivia, mostly Cuba, 22 in South America; 15 in Brazil, 8 endemics.

Two spp. are known to have ethnobotanical uses: P. glabrata (Schott) Baill., which is used to make wooden clogs in Brazil, and P. benensis Rusby, used by the Chimane Indians from Bolivia to treat cutaneous leishmaniosis.

3. Pogonophora Miers ex Benth. Trees or shrubs, 1.5 – 25 m tall; leaves simples, alternate; inflorescences in panicles, the female more long than male; fruits capsules. Two spp., one in W Africa (Gabon) and P. schomburgkiana Miers ex Benth. in the rainforests of Colombia to the Guianas, disjunct reaching in SE Brazil.

EUPHORBIACEAE

§ FULLY CHLOROTROPHIC

Genera/species 228/c. 6,745 Distribution cosmopolitan except polar areas, but mainly tropical, with their largest species diversity (Euphorbia) in S Africa, Mediterranean and the irano-turanian regions, and southern North America. Habit monoecious or dioecious, evergreen or deciduous trees or shrubs, perennial or annual herbs (rarely lianas; many species are stem succulents and xerophytes). CAM and C₄ physiologies present in many species (i.a. Chamaesyce subclade of Euphorbia).

Four, originally paleotropical, genera are cultivated with a single species each: Aleurites (candle nut), Codiaeum (garden croton), Ricinus (castor oil), Vernicia (tung oil). Hevea brasiliensis L., the Pará rubber tree, often simply called rubber tree (knows as seringueira in Brazil), is the most economically important member of the genus Hevea. It is of major economic importance because its sap-like extract (known as latex) is the primary source of natural rubber.

Euphorbiaceae are pantropical and comprise c. 230 genera and c. 5,700 spp.; in the Neotropics they are represented by 85 genera and c. 2,600 spp.; 79 of the 82 genera are native; 61 of these are endemic. The flowers are very small, and in many cases, it is not necessary to study their characters for generic identification; the family is very rich in extrafloral nectaries (on floral bracts, stipules and leaves), and shows a large variation in the type of hairs.

SYSTEMATIC six lineages; basal clades Cheilosoideae (2/7, S Burma, Nicobar Islands, Malesia to New Guinea, Solomon Islands) and Suregada clade (1–2/c 30, tropical regions in the Old World) do not occur in New World.

1. SUBFAMILY ADENOCLINOIDEAE (6/67) - outsiders Endospermum (11; SE Asia, Malesia to Fiji); Klaineanthus (1; Nigeria to Gabon), Adenocline (8; S Africa north to Malawi), Ditta (2; Cuba, Hispaniola, Puerto Rico).

1. Omphalea L. 20–25 spp., tropical regions on both hemispheres, 8 in New World, two in South America, O. diandra L., widely distributed in Neotropics, and O. brasiliensis Mull. Arg., endemic to Bahia state, Brazil.

2. Tetrorchidium Poepp. & Endl. Shrubs, lianes or trees, exuding a milky sap; leaves alternate or sometimes opposite on main axes, simple; inflorescences leaf-opposed; male flowers in simple, dense spikes; female flowers in 3–5-flowered umbels; fruit (2)3- locular and -lobed, seeds compressed-ellipsoid, sarcotesta orangered, endotesta foveolate, black. 23 spp., 5 in tropical center Africa, 18 in tropical America up to Argentina, 12 in South America, 5 in Brazil, two endemics.

2. SUBFAMILY ACALYPHOIDEAE (c. 98/1.940–1.975) - 12 tribes; tribes Sphyranthereae (1/2, Andaman Islands, Nicobar Islands), Pycnocomeae (6/51, Africa and Madagascar), Erismantheae (3/5, SE Asia, Hainan, W Malesia, Sumatra, Borneo, N Papua New Guinea), Epiprineae (8/33, tropical Africa, Madagascar, Socotra, Sri Lanka, S India to Hainan and New Guinea, Queensland), Ampereae (2/19, Australia) and Agrostistachydeae (4/11, tropical Africa, S India, Sri Lanka, Thailand, Malesia to New Guinea) do not occur in South America.

2.1 ACALYPHOIDEAE ▸ TRIBE ACALYPHEAE (c. 53/1,250-1,300) - outsiders Claoxylon (c 115; Madagascar, tropical Asia, N and E Australia, Melanesia, Hawaii), Crotonogynopsis (2; tropical Africa), Discoclaoxylon (4; tropical Africa, São Tomé); Erythrococca (c 40; tropical and S Africa, S Arabian Peninsula), Micrococca (12; tropical Africa, Madagascar, Arabian Peninsula, tropical Asia to the Malay Peninsula); Sampantaea (1; Thailand, Cambodia), Wetria (2; Burma, Thailand, W Malesia, Philippines, Papua New Guinea, Queensland); Lasiococca (5; Himalayas, Hainan, Vietnam, the Malay Peninsula), Spathiostemon (2; Peninsular Thailand, Malesia to New Guinea), Homonoia (3; Malesia New Guinea); Lobanilia (8; Madagascar); Macaranga (305–310; tropical regions in the Old World); Mareya (4; tropical W and C Africa); Mercurialis (12; Europe, Mediterranean, temperate Asia to N Thailand), Seidelia (2; N and W Cape, Free State), Leidesia (1; S Africa); Ricinus (1; E and NE Africa, Arabian Peninsula, SW Asia), Adriana (2; Australia); Mallotus (120–125; tropical Asia to tropical Australia and islands in the Pacific, two species in tropical Africa and Madagascar); Blumeodendron (5; SE Asia, Andaman Islands, Malesia to New Guinea), Podadenia (1; Sri Lanka), Ptychopyxis (13; Peninsular Thailand, Malesia to New Guinea), Botryophora (1; SE Asia, W Malesia); Afrotrewia (1; Cameroon, Gabon).

3. Acalypha L. Herbs to shrubs with variable hairs but not urticating, sometimes with xylopodium, leaves simple and serrate-crenate, inflorescences simple, bracts of pistillate flowers accrescent, conspicuous and often serrate to lobed, stigmas laciniate -multifid. 442 spp., pantropical, of which 221 are from New World, 83 in South America, 40 in Brazil, 17 endemics.

4. Avellanita Phil. Only one sp. A. bustillosii Phil., restricted of Chile.

5. Cleidion Blume. c. 25 spp., tropical regions on both hemispheres, 4 spp. in New World, from Mexico to Bolivia and Brazil (2, none endemics).

6. Dysopsis Baill. Small herb with suborbicular crenate leaves. Three spp., one fom Costa Rica to Colombia and two remaining from Ecuador to Chile, Argentina and Juan Fernandez, being a in montane habitats.

2.2 ACALYPHOIDEAE ▸ TRIBE ADELIEAE (4/c. 70) - outsiders Enriquebeltrania (2; Mexico), Garciadelia (4; Hispaniola), Lasiocroton (c 25; Cuba), Leucocroton (26–27; Cuba); 50 only in Cuba

7. Adelia L. 10 spp. from southern Texas to Argentina, Caribbean, Brazil (two, none endemics); 4 in South America, scattered.

2.3 ACALYPHOIDEAE ▸ TRIBE ALCHORNEEAE (7/c. 90) - outsiders Bocquillonia (14; New Caledonia), Orfilea (4; Madagascar, Mauritius); Aubletiana (2; Cameroon, Gabon), Mareyopsis (2; C Africa).

8. Alchornea Sw. Shrubs to trees up to 30 m high, rarely lianas, mainly dioecious; leaves are often coriaceous and distinctly triplinerved and/or distinctly dentate, the infloresces usually compound. 50–60 spp., tropical regions on both hemispheres, with 32 in New World, 30 in South America, 8 in Brazil, none endemics.

9. Aparisthmium Endl. Dioecious trees (2-25 m tall), rarely monoecious; leaves peninerved; inflorescences male as spikes or receme spiciform; inflorescences female in racemes terminal or axillary; fuits capsules, seeds 3. Only one sp., A. cordatum (A. Juss.) Baill., over tropical South America.

10. Conceveiba Aubl. Shrubs to trees, leaves simple, alternate; inflorescence panicle or corymbiform, flowers subsessile, sessile or pedicelate; calyx gamosepalous. 18 spp., two in tropical W Africa, 16 in South America, three up/or to Central America, 8 in Brazil, two endemics. C. martiana Baill. from northern South America is a myrmecophyte.

2.4 ACALYPHOIDEAE ▸ TRIBE BERNARDIEAE (3/80) - outsider Discocleidion (1; C China, the Ryukyu Islands).

11. Adenophaedra (Müll.Arg.) Müll.Arg. Dieocious shrubs or trees; leaves alternate, penninerved; staminate inflorescences in spiciform racemes or panicles, pistillate in panicles, often racemes; petals absent; fruit a capsule. Three spp. from Central America to Guianas, Ecuador and Brazil (all spp., with A. cearensis Secco endemic and rare plant in Brazil, by Plantas Raras do Brasil’s book, known only from Ceará state).

12. Bernardia Houst. ex Mill. 77 spp., California to Uruguay, Caribbean, 42 in South America, with their largest diversity in Brazil (29, 22 endemics).

2.5 ACALYPHOIDEAE ▸ TRIBE CARYODENDREAE (4/8) - outsider Discoglypremna (1; tropical Africa).

13. Alchorneopsis Müll.Arg. Dioecious trees (6-28 m tall), leaves alternate, trinerved; all inflorescences in spiciform racemes; fruit a capsule. Only one sp., A. floribunda (Benth.) Müll. Arg., Central America, Caribbean, Trinidad & Tobago, Venezuela, Guianas, Colombia, Ecuador, Peru, Bolivia, Brazil.

14. Bahiana J.F.Carrión. Shrub to trees, flowers diminute, trilocular fruits. Two spp., B. pyriformis J.F.Carrión, known only from dry mountainous region in center Bahia state, NE Brazil, and B. occidentalis K. Wurdack, endemic to Peru.

15. Caryodendron H.Karst. 5 spp. from northern South America, one up to Central America; three in Brazil, one endemic.

2.6 ACALYPHOIDEAE ▸ TRIBE CHROZOPHOREAE (11/c. 165) - outsiders Chrozophora (11; Mediterranean, tropical East Africa, SW and S Asia to Thailand); Doryxylon (1; Luzon, the Lesser Sunda Islands), Melanolepis (2; SE Asia to islands in the Pacific, Taiwan in China), Sumbaviopsis (1; Assam, SE Asia, W Malesia), Thyrsanthera (1; SE Asia); Speranskia (3; S China, N Burma)

16. Argythamnia P.Browne. 32 spp., 31 from Caribbean, Texas to Panamá, two of then up to Venezuela and Colombia, and one endemic to Bolivia.

17. Caperonia A.St.-Hil. Herbs to small shrubs, characterized by leaves with a typical venation, the straight side veins running directly into the serrate margin of the blade (craspedodromous). 42 spp., tropical Africa, Madagascar, 29 in tropical America, 27 in South America, 18 in Brazil, 11 endemics.

18. Chiropetalum A.Juss. 22 spp., two in Mexico and Texas, remaining 20 in Peru to Chile, up to Paraguay and southern Brazil (9, 6 endemics).

19. Ditaxis Vahl ex A.Juss. 33 spp., tropical and subtropical America, 25 in South America, 13 in Brazil, 10 endemics.

20. Philyra Klotzsch. Shrubs, dioiceous, stems spinescent, alternate leaves, chartaceous; inflorescence axillary, racemose. Only one sp., P. brasiliensis Klotzsch, from Paraguay, NE Argentina, southern Brazil, and areas in Bahia and Alagoas states.

2.7 ACALYPHOIDEAE ▸ TRIBE PLUKENETIEAE (17/c. 340) - outsiders Ctenomeria (2; South Africa), Megistostigma (5; Yunnan, SE Asia, W Malesia), Cnesmone (11; Assam, SE Asia, W Malesia), Platygyna (7; Cuba), Acidoton (6; Hispaniola and Jamaica), Zuckertia (2; Mexico, Central America), Pachystylidium (1; India, SE Asia to C Malesia), Sphaerostylis (2; Madagascar).

21. Angostylis Benth. Two spp., A. longifolia Benth. from Brazil in Amazonas, in areas of 100 m elevation range, and A. tabulamontana Croizat from Suriname.

22. Astrococcus Benth. Only one sp., A. cornutus Benth., endemic to the Guiana Shield of Colombia, Venezuela and Amazonas state in Brazil, 100-200 m elevation range.

23. Bia Klotzchia. Monoecious herbs, twining vines or suberect perennial shrubs, and subshrubs; leaves alternate, simple, membranaceous, petiolate; Inflorescences oppositifolious, terminal, a bifurcated raceme with one branch staminate and the other pistillate; flowers pedicellate; staminate flowers in threes or solitary; pistillate flowers always solitary. 5 spp. from South America, all in Brazil, B. capivarensis D.Medeiros, L.Senna & R.J.V. Alves endemic.

24. Chicomendes W.Cordeiro & M.F.Sales (off Tragia). Herbs with lanceolate leaves. Only one sp., C. rubiginosus (Huft) W.Cordeiro, Athiê-Souza & A.L.Melo, known only two locations, one in Loreto in N Peru, another from W Amazonas state, N Brazil.

25. Dalechampia Plum. ex L. Twining vines, erect or decumbent herbs, and rarely subshrubs, with stinging hairs (trichomes) and pseudanthal inflorescences composed of two involucral bracts, a staminate pleiochasium with 7–15 staminate flowers and three pistillate flowers, often clusters of resiniferous glands. 99 spp. from New World, mainly in South America (89), 74 in Brazil, 39 endemics, one of them, from Amazonas state, is a rare plant in Brazil, by Plantas Raras do Brasil’s book.

26. Gitara Pax & K. Hoffm. Only one sp., G. nicaraguensis (Hemsl.) Card.-McTeag. & L.J. Gillespie, in Central America, Venezuela, Colombia, Ecuador, Peru, Bolivia and N Brazil.

27. Haematostemon (Müll.Arg.) Pax & K.Hoffm. Two spp., endemic to the Guiana Shield, H. coriaceus (Baill.) Pax & K. Hoffm. in Venezuela, Brazil (only Amazonas state) and Colombia, and H. guianensis Sandwith only Guyana, 100-200 m elevation range.

28. Monadelpha L.J.Gillespie & Card.-McTeag. (off Tragia) Climbing vines, apparently monoecious; latex absent; stems twining; stems, leaves and inflorescences with stinging and simple hairs. Only one sp., M. guayanensis (L.J.Gillespie) L.J.Gillespie & Card.-McTeag., known only three collectons in southern Venezuela (two) and one a single imprecise in Amazonas state, northern Brazil, along Jurua Valley.

29. Plukenetia L. Climbing vines to lianas, hairs simple, leaves simple, conspicuously glandular at base, inflorescences leaf-opposed, racemose-paniculate, flowers and fruits 4-merous (the 4-merous flowers distinguish it from several similar genera). c. 20 spp., tropical Africa, Madagascar, P. corniculata Sm. in tropical Asia, 17 in tropical America, 15 in South America, 9 in Brazil, two endemics.

30. Romanoa Trevis. Vines. Only one sp., R. tamnoides (A. Juss) Radcl.-Sm; E & southern Brazil and adjacent areas of Paraguay and Bolivia.

31. Tragia L. (exc. Monadelpha, Chicomendes) Herbs to subshrubs, often twining, sometimes with xylopodium; indumentum of simple and urticating hairs, leaves simple and often serrate, inflorescences often leaf-opposed, racemose, staminate flowers with 2-50 stamens; some smaller similar genera differ in minor floral characters, and the generic limits are not completely solved yet. 152 spp., tropical and subtropical regions on both hemispheres, with 49 in New World, 24 in South America, 14 in Brazil, 4 endemics.

3. SUBFAMILY CROTONOIDEAE (60/2,010-2,020) ‣ 10 subtribes, Ricinocarpeae (8/79, Australia, Borneo, New Caledonia) and Ricinodendreae (5/11, tropical East Africa, Madagascar, India, Sri Lanka, New Guinea) do not occur in South America.

3.1 CROTONOIDEAE ▸ TRIBE ALEURITIDEAE (15/60) - outsiders Aleurites (2; tropical Asia to islands in W Pacific), Vernicia (3; Burma, SE Asia, Malesia and S China to Japan); Benoistia (3; Madagascar); Cyrtogonone (1; tropical W Africa), Crotonogyne (16; tropical Africa), Manniophyton (1; W and C tropical Africa to Angola); Grossera (8; tropical Africa, Madagascar), Cavacoa (3; tropical Africa), Tannodia (9; tropical Africa, Madagascar), Tapoides (1; Borneo); Neoboutonia (3; tropical Africa); Deutzianthus (2; northern Vietnam; Sumatra), Oligoceras (1; Vietnam), Paracroton (4; S India, Sri Lanka, Malesia to New Guinea); Mildbraedia (5; W and C tropical Africa to Mozambique).

32. Garcia Vahl ex Rohr. 6 spp., 4 only in Caribbean and two from Mexico, G. nutans Vahl ex Rohr up to Colombia.

3.2 CROTONOIDEAE ▸ TRIBE CODIAEEAE (15/c. 180) - outsiders Alphandia (3; New Guinea, New Caledonia, Vanuatu), Baliospermum (5; Himalayas, Tibet, Yunnan, Indochina, the Malay Peninsula, Sumatra, Java, Sumbawa), Baloghia (15; Queensland, New South Wales, Norfolk Island, New Caledonia), Blachia (11; India, Andaman Islands, S China, SE Asia, Philippines), Codiaeum (16–17; Malesia to New Guinea, tropical Australia and New Caledonia), Dimorphocalyx (17; India, Sri Lanka to Philippines, New Guinea and tropical Australia), Fontainea (9; New Guinea, Queensland, New Caledonia, Vanuatu), Hylandia (1; Queensland), Ostodes (2; Assam, Himalayas, SE Asia to Java and Borneo), Pantadenia (3; Madagascar; Thailand, Indochina), Strophioblachia (1; Yunnan, Hainan, Thailand, Cambodia, Vietnam, Philippines, Sulawesi); Trigonostemon (c 85; India and China to Philippines, New Guinea, Queensland and Fiji).

33. Dodecastigma Ducke (inc. Anomalocalyx). Three spp. of South America, from French Guiana to northern Brazil (2, none endemics), Peru and Bolivia

34. Pausandra Radlk. Shrubs to trees, leaves alternate, penninerved; inflorescences axillary or terminal, in spikes with solitary flowers to 3-chasia; staminate flowers incompiscuous, pistillate flowers medium-sized; fruit a capsule. 8 spp. from South America (all in Brazil, two endemics), only one up Honduras.

3.3 CROTONOIDEAE ▸ TRIBE CROTONEAE (6/c. 1250) - all genera in South America.

35. Astraea Klotzsch. Similar to Croton but usually with lobed leaves and with staminate flowers with a glabrous receptacle. 12 spp., A. lobata (L.) Klotzsch widely distributed from Florida, Mexico to trop. America, nine endemics to Brazil and two restricted in Bolivia and Paraguay.

36. Acidocroton Griseb. 14 spp. distributed in Caribbean (10), Mexico (3) and N Colombia (A. gentryi Fern. Alonso & R. Jaram.).

37. Brasiliocroton P.E.Berry & Cordeiro. Monoecious tree 3–15 m tall, diameter of main trunk to 30 cm; trunk sulcate with smooth bark; inflorescence a terminal bisexual panicle, with a main terminal rachis and several subterminal axillary branches; flowers cream to white, greenish in bud, mostly staminate. Two spp. from nonflooded, primary or secondary forests and in low, at the edge of rain forests, seasonally dry forests, and secondary forests, coastal forests at elevations of 50–800 m, in remnants of the Atlantic Forest in southern Bahia, Espírito Santo, and E Minas Gerais states, and disjunct in northern Maranhão state.

38. Croton L. [6th BR] Herbs to trees or lianas, sometimes with xylopodium; branches sometimes resinous, leaves vriated; hairs dendritic or lepidote leaves usually simple with a pair of basal glands, inflorescences mostly simple, staminate flowers with a pubescent receptacle and with numerous stamens inclinate in bud; fruit a capsule. 1,141 spp., cosmopolitan, pantropical, 748 species in New World, 540 in South America, 334 in Brazil, 248 endemics, 5 of them are rare plants in Brazil, by Plantas Raras do Brasil’s book, in Minas Gerais, Amazonas and Pará states; 31 sections in New World, Croton subg. Croton is exclusively from Old World.

§ subg. Quadrilobi (Müll. Arg.) Pax in Engl. & Prantl

§ sect. Olivacei ‣ a single sp., forests in Ecuador and Peru.

§ sect. Sampatik ‣ 4 spp., widely distributed in South America in the Amazon-Orinoco rainforests to the foothills of the Andes and in the Atlantic Forests of SE Brazil, from 100-900 m.

§ sect. Quadrilobi ‣ a single sp., only one sp. endemic to low-elevation moist forests in southern Bahia, Brazil.

§ sect. Pachypodi ‣ 5 spp., a mainly South American group that reaches its northern limit in Costa Rica; it is found in humid montane and lower montane forests in Costa Rica, Panamá, Colombia, Venezuela, Ecuador, Peru, Brazil, and Bolivia, from 50-1,200 m.

§ sect. Moacroton ‣ 8 spp., endemic to Caribbean.

§ sect. Nubigeni ‣ a single sp., endemic to Honduras and Nicaragua.

§ sect. Alabamenses ‣ a single sp., endemic to U.S.A.

§ sect. Corinthii ‣ a single sp., endemic to Costa Rica.

§ sect. Corylocroton ‣ 11 spp., deciduous forests of the Antilles, southern Mexico, and South America, and evergreen cloud forests of Central America, from sea level to 1,600 m.

§ subg. Adenophylli

§ sect. Cyclostigma ‣ 41 spp.; secondary vegetation, roadsides, river banks, and landslide areas of dry to wet forests from central Mexico and Central America to tropical and subtropical South America, from sea level to 3,000 m. C. amenthiformis Riina from Ecuador and N Peru is unique within Croton because of its peculiar pendant inflorescence with congested flowers and dimorphic bracts.

§ sect. Adenophylli ‣ 223 spp., in a wide variety of habitats, many of them semiarid and often in secondary vegetation, in the southern U.S.A., Mexico, Central America, the Caribbean, and all countries of South America, from sea level to 3,000 m.

§ subg. Geiseleria

§ sect. Cupreati ‣ a single sp., Andean cloud forests of northern Ecuador and SE Colombia.

§ sect. Eremocarpus ‣ a single sp., Mexico and U.S.A.

§ sect. Cuneati ‣ 11 spp., entirely South American section, including the adjacent Caribbean island of Trinidad, consists of rainforest trees found from seasonally flooded riparian Amazon rainforest lowland forests on the slopes of tepuis in the Guiana Shield and the Venezuelan Coastal Range, from 100 - 2500 m.

§ sect. Eluteria ‣ 22 spp.

o subsect. Eluteria ‣ 15 spp., widely distributed in the Caribbean, and from Mexico to tropical South America; dry, open vegetation to rainforests, from 0- 1,700 m.

o subsect. Cubenses ‣ 4 spp., endemic to Cuba; dry, scrubby vegetation from sea level to 500 m.

o subsect. Jamaicenses ‣ 3 spp., endemic to Jamaica; dry, scrubby vegetation on limestone and moister upland areas, from sea level to 700 m.

§ sect. Crotonopsis ‣ a single sp., endemic to U.S.A.

§ sect. Argyranthemi ‣ 2 spp., Mexico and U.S.A.

§ sect. Drepadenium ‣ 6 spp., a primarily North American group which would be endemic to Mexico and the U.S.A. except for the widely distributed, sand dune species C. punctatus Jacq., which extends along the Atlantic coast to northern South America (absent in Brazil).

§ sect. Prisci ‣ 3 spp., occurring in moist forests of the Brazilian states of São Paulo, Rio de Janeiro, and Bahia.

§ sect. Pedicellati ‣ 20 spp., distributed disjunctly between Mexico and South America, with its greatest diversity in eastern Brazil; it is found in scrubby tropical deciduous forests in Mexico and Brazil, as well as in dry inter-Andean valleys in Colombia, Peru, and Bolivia, from 200-2,500 m.

§ sect. Lamprocroton ‣ 37 spp.

o subsect. Lamprocroton ‣ 17 spp., a South American group is most diverse in Brazil, but extends into adjacent Paraguay, Argentina, Uruguay, and Bolivia; it is found most commonly in open vegetation on sandy or rocky ground, including rock outcrops, from 100-2,800 m, but some members are also known from riparian and swampy habitats

o subsect. Argentini ‣ 20 spp., entirely South American, but its center of diversity appears to be south of Brazil in Uruguay and Argentina.

§ sect. Luetzelburgiorum ‣ a single sp., C. luetzelburgii Pax & K. Hoffm., restricted to the Diamantina Range of center Bahia.

§ sect. Cleodora ‣ 18 spp., inhabit moist or seasonally dry forests in tropical South America, Central America, and into central Mexico.

§ sect. Cordiifolii ‣ a single sp., dry seasonal scrubland of NE Brazil (caatinga), in Bahia.

§ sect. Eutropia ‣ a single sp., endemic to Atlantic sandy coastal shrublands (restingas) of E Brazil from sea level to 150 m.

§ sect. Geiseleria ‣ 82 spp., fields, roadsides, waste places, open and mostly dry vegetation, deciduous to occasionally moist forests, widely distributed in the New World from the U.S.A. to Argentina, from sea level to 1,800 m. C. glandulosus L. and C. hirtus L’Hér have been introduced as weeds in Africa and Australia.

§ sect. Barhamia ‣ 84 spp.

o subsect. Barhamia ‣ 19 spp., widely distributed in the Caribbean and Central and South America, with its greatest diversity in Brazil. It is found in mostly open, dry vegetation from 100-1,200 m.

o subsect. Astraeopsis ‣ 4 spp., Caribbean and the Yucatan peninsula of Mexico and adjacent parts of Belize. Open, scrubby vegetation, usually on limestone, from sea level to 300 m.

o subsect. Medea ‣ 45 spp., Brazil, Paraguay, Uruguay, and Argentina; temperate and subtropical, generally open vegetation, from sea level to 1,000 m.

o subsect. Micranthi ‣ 13 spp., Mexico, Caribbean, and Central and South America; sand barrens, coastal regions, waste places, and other open vegetation, from sea level to 1,500 m.

o subsect. Sellowiorum ‣ 3 spp., sandy soils in open vegetation in eastern Brazil. C. sellowii Baill. occurrs at lower elevations in Atlantic sandy coastal shrublands (restingas) from sea level to 100 m. C. myrsinites Baill. and C. schultesii Müll. Arg. occupy rocky grasslands (campos rupestres) habitats from 800 to 1,500 m.

§ sect. Luntia ‣ 19 spp., widely distributed across tropical South America and extends into Central America in Panamá, Costa Rica, and Nicaragua. Its members are found in primary to secondary moist forests and disturbed sites, often near streams, from sea level to 3000 m.

§ sect. Julocroton ‣ 41 spp., primarily South American group with its greatest diversity in Brazil, but a few species extend into the rest of tropical South America, and one species, C. argenteus L., extends throughout Central America, the Caribbean, and up to the southern U.S.A.

§ sect. Lasiogyne ‣ 43 spp., widely distributed in the New World tropics and subtropics, occurring mostly in dry forest and scrub vegetation, from sea level to 1,000 m.; 11 spp. in Brazil, 6 endemics.

§ sect. Heptallon ‣ 9 spp., Mexico and U.S.A.

39. Sagotia Baill. Two spp., from Central America and northern Colombia to Mato Grosso (and Maranhão state) and Bolivia – both in Brazil.

40. Sandwithia Lanj. Trees; leaves simple; inflorescence in panicle or fascicle; all flowers pedicelate, leaves present. Two spp. from Colombia to French Guiana to northern Brazil in Amazonas, Roraima and Amapá states.

3.4 CROTONOIDEAE ▸ TRIBE ELATERIOSPERMEAE (2/3) - outsider Elateriospermum (1; Peninsular Thailand, W Malesia).

41. Glycydendron Ducke. Dioecious trees, 15-30 m tall; white latex; leaves palmatinerved; inflorescences stamineta in panicles, inflorescences pistilate in racemes; fruits a drupe. Two spp., G. amazonicum Ducke, in over Amazon rainforests, and G. espiritosantense Kuhlm. endemic to Espírito Santo state in E Brazil.

3.5 CROTONOIDEAE ▸ TRIBE HEVEEAE (1/10) - a single genus.

42. Hevea Aubl. Trees, sometimes emergent canopy up to 50 m tall, one of them the tallest Malpighiales from Brazil; hairs simple, leaves 3-foliolate, inflorescences pseudopaniculate, flowers without petals. 11 spp. from the Amazon rainforest, all in Brazil (three endemics, one of them, from Pará state, is a rare plant in Brazil, by Plantas Raras do Brasil’s book).

3.6 CROTONOIDEAE ▸ TRIBE JATROPHEAE (3/190) - all genera occur in South America.

43. Jatropha L. Herbs to trees, sometimes succulent, often with a thick rootstock, hairs simple, leaves variously shaped but often palmately lobed, stipules often conspicuous and multifid, inflorescences dichotomous, corymbiform, flowers with distinct sepalas and petals. 155 spp., tropical and subtropical regions on both hemispheres, North America, with c. 102 in New World, 46 in South America, 14 in Brazil, 9 endemics.

44. Joannesia Vell. Trees up to 30 m tall. Two spp. J. heveoides Ducke from Amazonas rainforest region and J. princeps Vell. in E Brazil, fro Maranhão to Paraná state.

45. Vaupesia R.E.Schult. Only one sp., V. cataractarum R.E. Schult., Colombia, Amazonas and Roraima states in NW Brazil.

3.7 CROTONOIDEAE ▸ TRIBE MANIHOTEAE (2/c. 200) - both genera occur in South America.

46. Cnidoscolus Pohl. Herbs to trees, similar to Jatropha, but plants with urticating hairs or bristles, leaves biglandular at base, flowers with a petaloid perianth but without separate petals. 99 from U.S.A. to Argentina and Caribbean, mainly Mexico (27, 21 endemics) and Brazil (44, 40 endemics); 68 in South America; C. vitifolius (Mill.) Pohl from Brazil can have a distinct, though short, swollen trunk and relatively few basal branches, but it branches copiously thereafter, leaving the general impression of a shrub rather than a tree. Eight sections:

§ sect. Acrandrae ‣ two spp., Cuba and Rep. Dominicana.

§ sect. Calyptrosolen ‣ 20 spp., mainly in Mexico and Central America but with a few species in Colombia, Ecuador and Venezuela.

§ sect. Cnidoscolus ‣ 10 spp., 8 endemics to Brazil and two endemics to Peru.

§ sect. Graminifolius ‣ only one sp., C. graminifolius Fern. Casas, known only from the state of Tocantins, Brazil.

§ sect. Jussieuia ‣ 25 spp., vast majority endemic or microendemic to Brazil, but some are restricted to the Northern Hemisphere (C. texanus (Müll. Arg.) Small, C. stimulosus (Michx.) Engelm. & A. Gray); this section also includes the most widespread species in the genus, C. urens (L.) Arthur, reported from Argentina, Bolivia, Brazil, Colombia, Costa Rica, Ecuador, French Guiana, Guatemala, Guyana, Mexico, Nicaragua, Panama, Paraguay, Peru, Suriname, Venezuela and Caribbean.

§ sect. Vitifoliae ‣ 25 spp., mainly in Brazil and Paraguay but with a few species reaching Argentina, Bolivia, and Peru.

§ sect. Oligandrae ‣ 3 spp., all endemics to Brazil.

§ sect. Platyandrae ‣ two spp., endemics to Cuba.

47. Manihot Mill. Acaulescent shrubs (in center Brazil) to medium sized trees, sometimes vine-like (7, mainly in Brazil), including perennial subshrubs with large taproot tubers or xylopodium; and whose stems frequently die back to the root crown in response to dry periods or fires; somes species lactescents and rubber-producing; not urticating, leaves simple (all in South America) to palmately lobed, often pruinose-glaucescent, eglandular but stipellate at base, inflorescences racemose-paniculate, flowers without petals; lactiferous spp., mainly in tree-habit types. 151 spp., tropical and subtropical America, centered in Goiás/Minas Gerais in C Brazil (38-40), Mexico (21, 14 endemics), NE Brazil (16), Mato Grosso and adjacent Bolivia (6), mainly dry places, barely in forets, often dominat; 131 spp. in South America, 115 in Brazil, 100 are endemics and 40 have been reported from the state of Goiás.

§ sect. Anisophyllae ‣ two spp., Bolivia, Argentina and Paraguay.

§ sect. Brevipetiolatae ‣ 6 spp., C Brazil.

§ sect. Caerulescentes ‣ two spp., Brazil to Paraguay.

§ sect. Carthaginensis ‣ two spp., N Venezuela to N Colombia.

§ sect. Crotalaria ‣ 4 spp., Paraguay to Brazil.

§ sect. Foetidae ‣ 6 spp., endemic to Mexico.

§ sect. Glazioviinae ‣ 7 spp., NE Brazil.

§ sect. Graciles ‣ 10 spp., C Brazil to Paraguay and NE Argentina.

§ sect. Grandibracteatae ‣ only one sp., M. tomentosa Pohl., endemic to C Brazil.

§ sect. Heterophyllae ‣ 15 spp., Colombia to N Pará state, Andes from Peru an Bolivia, and Bahia to N Argentina.

§ sect. Manihot ‣ only one sp., M. esculenta Cranz., a cultigen possibly native from Brazil. M. esculenta Crantz, the cassava or mandioca, sometimes escribed as the ‘bread of the tropics’, is a woody shrub of the spurge family native to Brazil and Paraguay, one of the most drought-tolerant crops, extensively cultivated as an annual crop in tropical and subtropical regions for its edible starchy, tuberous root, a major source of carbohydrates, providing a basic diet for around 500 million people; just is a 15 crop plants provide 90 % of the world’s food energy intake, and cassava is the fifth most produced food in world, with 233 milion tons in 2008, after rice, maize, wheat and potato, and given the highest yield of carbohydrates per cultivated area among crop plants, except for sugar cane and sygar beets; wild populations of Manihot esculenta subspecies flabellifolia, shown to be the progenitor of domesticated cassava, are centered in WC Brazil.

§ sect. Parvibracteatae ‣ 11 spp., Arizona in U.S.A. to Panamá.

§ sect. Peruviana ‣ 3 spp., Ecuador and Peru to NE Brazil.

§ sect. Peltata ‣ 3 spp., Brazil to Paraguay.

§ sect. Quinquelobae ‣ 14 spp., endemic to C Brazil.

§ sect. Sinuatae ‣ two spp., Peru to N Argentina and C Brazil.

§ sect. Stipularis ‣ 5 spp., Goiás and Minas Gerais states in C Brazil.

§ sect. Tripartitae ‣ only one sp., M. tripartita (Spreng.) Müll.Arg., from Brazil, Paraguay and Bolivia.

§ sect. Variifoliae ‣ two spp., Paraguay and Mato Grosso do Sul state in Brazil.

3.8 CROTONOIDEAE ▸ TRIBE MICRANDREAE (2/12) - both genera occur in South America.

48. Micrandra Benth. Monoecious trees, leaves alternate, penninerved, stipulate; inflorescences bissexual, in panicles simple or composed, axillary or terminal, flowers apetalous; fruit a capsule. 12 spp., Amazon rainforest of tropical South America, 8 in Brazil, none endemics.

49. Micrandropsis W.A.Rodrigues. Only one sp., M. scleroxylon (W.A.Rodrigues) W.A.Rodrigues, from Brazil (Amazonas) and SE Colombia.

4. SUBFAMILY EUPHORBIOIDEAE (38/2.410–2.420) ‣ 3 tribes, all in South America.

4.1 EUPHORBIOIDEAE ▸ TRIBE STOMATOCALYCEAE (4/9) - outsiders Plagiostyles (2; Nigeria, Gabon, Congo), Pimelodendron (4; Malesia to New Guinea, Queensland); Hamilcoa (1; Nigeria, Cameroon).

50. Nealchornea Huber. Dioecious trees, icthiotoxic; latex whitish or yellowish, sweet; leaves alternate; Inflorescense male in panicles, with flowers apetalous; female inflorescences in racemes, flowers also apetalous; fruits indehicent. Two spp. from E. Colombia to Peru and Amazonas state in northern Brazil (both, one endemic)

4.2 EUPHORBIOIDEAE ▸ TRIBE EUPHORBIEAE (5/c. 2060) - outsiders Anthostema (3; tropical W Africa, Madagascar), Dichostemma (2; C Africa; Cameroon); Neoguillauminia (1; New Caledonia), Calycopeplus (5; northern Australia, W Australia).

51. Euphorbia L. Herbs to shrubs or succulents, small trees, prostrate or decumbent, all produce a mostly white latex which they exude when cut, and this sap is often toxic, characterized by the unique pseudanthium of much-reduced flowers (cyathium); recently all cyathia-bearing species were united into one single genus; thus, this feature is present in every species of the genus but nowhere else in the plant kingdom. 2,046 spp. (5^th largest worldwide), fully cosmopolitan; succulent spurges are most diverse in southern and E Africa and Madagascar, but they also occur in tropical Asia and the Americas; 496 in New World, 246 spp. in Mexico (129 endemic), 182 in South America, 69 in Brazil (41 endemics). 4 subgenera, three in New World (59 section joined) and South America (12 sections within):

The most morphological diverse genus of angiosperms in Brazil, and the 6ª largest group of succulent species worldwide, with c. 650 spp. E. laevigata Lam. (endemic to Brazil) unplaced.

§ subg. Athymalus ‣ 7 sections, only Old World.

§ sect. Antso ‣ only one sp., E. antso Denis., from W and SW Madagascar (Toliara Region), in semiarid, deciduous forests and scrub, often on soils derived from limestone, from sea level to 500 m.

§ sect. Pseudacalypha ‣ 11 spp., Ethiopia, Somalia, Sudan and the southern Arabian Peninsula (Oman, Yemen), extending to Angola, Egypt, Kenya and Tanzania; the non-succulent species occur in open, disturbed habitats, rocky terrain, sheltered among stones or under small shrubs; from sea level to 1,800 m.

§ sect. Lyciopsis ‣ 19 spp., WC to E and NE Africa and the Arabian Peninsula (Oman, Saudi Arabia, Yemen, Socotra), extending into Angola, Botswana and Namibia; dry stony areas among deciduous shrubs and trees, gypsum, limestones, granites, or sand dunes, from sea level to 2,100 m.

§ sect. Crotonoides ‣ 8 spp., S to E Africa, in Angola, Botswana, Namibia, South Africa, Zimbabwe, Malawi, Zambia, Tanzania, Kenya, Sudan and Ethiopia; on grassy hillsides, in leaf-litter under trees in dry open woodlands, on sand or loam, sometimes in disturbed patches, 200–3,000 m.

§ sect. Somalica ‣ 13 spp., Tanzania, Kenya, Ethiopia, Somalia and Yemen (Socotra); dry, largely deciduous bushland in open flats, on slopes of rocky hills and escarpments, from near sea level to 1,000 m.

§ sect. Balsamis ‣ 9 spp., S Arabian Peninsula up into Iran, also to Sudan (Red Sea Hills), Somalia, Socotran Archipelago (Abd al Kuri only), as well as in West Africa and the Canary Islands, on dry stony slopes or stony flat plains and they sometimes form the dominant component of the otherwise very scanty vegetation.

§ sect. Anthacanthae ‣ 87 spp. found in southern Africa to very sparsely distributed over the rest of sub-Saharan Africa; bare, dry, stony plains with little other vegetation to deep Kalahari sand among scattered clumps of grass and trees to among fynbos vegetation, from sea level to 2,700 m.

§ subg. Esula ‣ 457 spp., worldwide, 21 sections, only sect. Helisocopia in South America.

§ sect. Aphyllis ‣ 23 spp., Macaronesian islands (except for the Azores), southern coast of Portugal, and the Atlantic coast of Morocco and Western Sahara, Africa, S Arabian Peninsula, Socotra, and Madagascar.

§ sect. Arvales ‣ 7 spp., Mediterranean region and neighboring areas, Canary Islands, Iranian highlands, central Asia to N Afghanistan, W China, and India; on open stony or gravelly slopes, in steppes and deserts, in fields and by roadsides.

§ sect. Biumbellatae ‣ 3 spp., endemic to the W Mediterranean region (Spain, France, Italy, Algeria, Morocco, Tunisia).

§ sect. Calyptratae ‣ two spp., deserts and steppes of N Africa (E. calyptrata Coss. & Durieu) and Iran (E. connata Boiss.).

§ sect. Chylogala ‣ 4 spp., Mediterranean region, Arabian Peninsula, SW and C Asia (to N PamirAlay and W Tien-shan); in deserts and steppes, on stony mountain slopes and by dry streams, or in fallow fields.

§ sect. Esula ‣ 96 spp., widespread in Europe and Asia, also in Africa, Madagascar, Réunion, Indonesia (Java), New Zealand, and Samoa, introduced in the New World and elsewhere; in meadows (lowland to montane), steppes, semideserts, riparian and littoral vegetation, sometimes in forests, weedy in arable lands, along roads and other disturbed areas.

§ sect. Exiguae ‣ 5 spp., S Europe, northern Africa, Asia, Macaronesia, Mauritius, the Comoros, and Madagascar; in grasslands, rocky or sandy subdeserts, rock fissures in mountains, weedy areas, abandoned cultivated areas, from sea level to 2,200 m.

§ sect. Guyonianae ‣ only one sp., E. guyoniana Boiss. & Reut., from NW Africa (Morocco to Libya); dunes and sandy areas on the northern edge of the Saharan Desert.

§ sect. Helioscopia ‣ 136 spp., most diverse across Eurasia and North Africa, but a few species also present in eastern Africa, Macaronesia, and the New World (26); the range of habitats is very diverse, from montane forests to steppes, rocky outcrops and wetlands, with a few weedy species; most species are associated with mountains and are fairly mesic, although some are widely distributed in lowland areas. 2 spp. in South America, E. philippiana (Klotzsch & Garcke) Boiss., widely disjunct to central Chile and is the only species in subg. Esula endemic to South America, and one in Brazil, E. spathulata Lam. disjunct from Canada to Mexico, and southern Brazil, NE Argentina and Uruguay.

§ sect. Herpetorrhizae ‣ 12 spp., Asia, from Türkiye and Caucasus to Kuwait, Pakistan, and Afghanistan, and into W China; on sand, clay and saline soils in deserts and steppes, and on stony slopes of mountains.

§ sect. Holophyllum ‣ 27 spp., mountainous areas of E and C Asia, the Himalayas, S Siberia, NE China, Korea, and Japan, with one species disjunct in the Iberian Peninsula; in forests, steppes, on rock outcrops, and in alpine meadows.

§ sect. Lagascae ‣ 3 spp., Canary Islands, Mediterranean region, and western part of Iranian highlands, in grasslands and ruderal habitats.

§ sect. Lathyris ‣ only one sp., E. lathyris L., found mostly in cultivation or near human settlements, in two main areas: eastern Asia and the Mediterranean, but also widely distributed by humans and found in temperate or subtropical regions worldwide.

§ sect. Myrsiniteae ‣ 14 spp., Mediterranean region, Caucasus to the Iranian highlands, usually growing in dry, exposed, rocky habitats.

§ sect. Pachycladae ‣ two spp., Circum-Mediterranean region; both species are mainly littoral, but can penetrate up to 100 km inland, in summer-arid areas, on coastal sands in littoral scrub, on edges of roads and trails, on substrates of siliceous, schistose, gypseous, volcanic, or calcareous origin, from sea level to 800 m

§ sect. Paralias ‣ 12 spp., Mediterranean region, Balkans, Caucasus, Macaronesia (including Azores), with E. tricotoma Kutnh in S Florida, SE Mexico to Belize, Bahamas, Cuba, Cayman Islands; often in coastal sandy habitats but some species farther inland and upland.

§ sect. Patellares ‣ 14 spp., Mediterranean region and mountains of central Europe, the Balkan Peninsula, Caucasus, and Iranian highlands; in forests and montane meadows or rocky habitats.

§ sect. Pithyusa ‣ 50 spp., mainly Mediterranean and neighboring areas, Iranian highlands and montane regions of central Asia, with a few species in the steppe zone of Eurasia and in the mountains of eastern Asia (at least as far east as Thailand); on rocky (calcareous) substrates, in steppes and steppe-like grasslands, deserts, dry scrub, and gypsum hills, sometimes weedy.

§ sect. Sclerocyathium ‣ 9 spp., SW and central Asia, including the Himalayas; on dry stony mountain slopes, sometimes in steppes.

§ sect. Szovitsiae ‣ only one sp., E. szovitsii Fisch. & C.A. Mey., from Caucasus, Türkiye, Iranian highlands, southern part of central Asia; on gravel slopes, screes, stream beds, or in steppes.

§ sect. Tithymalus ‣ 35 spp., 7 native to and restricted to the Old World from the eastern Mediterranean region to Iran and the Arabian Peninsula; E. peplus L. is also presumably native to the Mediterranean region but is now widespread worldwide; the remaining species are native to the New World from U.S.A., Canada, Mexico, Central America, Hispaniola. Many of them are restricted to relatively high montane habitats, but others occur at lower elevations and are either widespread or local.

§ subg. Chamaescyce - 566-574 spp. worldwide, 15 sects., 4 in South America; subg. Chamaescyce is the only taxonomic group at the lower level than the genus with photosynthesis C3, C2, C4 and CAM, with C2 in this group occurring only in E. acuta Engelm. (U.S.A.) and E. johnstoni Mayfield (Mexico).

§ sect. Alectoroctonum ‣ 115 spp., widespread in the New World from Canada to Argentina, with a center of diversity in Mexico and Central America; tropical and subtropical forests, desert scrub, and disturbed areas, sea level to 3,000 m; three species in Brazil, E. estevesii N. F. A. Zimmermann & P. J. Braun and E. sarcoceras O.L.M. Silva & Cordeiro endemics, and E. insulana Vell. up to over South America.

§ sect. Anisophyllum ‣ two subsections.

o subsect. Acutae ‣ two spp., W Texas and Mexico (Coahuila, Durango, Tamaulipas); grassland to desert scrub in and around the Chihuahuan Desert on sandy or gravelly limestone substrates, 200–1,500 m.

o subsect. Hypericifoliae ‣ 365 spp., warm, arid and semi-arid vegetation or disturbed habitats, and summer annuals in temperate areas; worldwide, from sea level to 4,000 m; 30 spp. in Brazil, 11 endemics.

§ sect. Articulofruticosae ‣ 18-26 spp., most diverse in the arid winterrainfall region of W South Africa and S Namibia, extending into southern Angola and Botswana and east to KwaZulu-Natal, South Africa; growing in sandy soils or on rock outcrops, in low shrublands to deserts and consolidated dunes, from sea level to ca. 2,000 m

§ sect. Bosseriae ‣ 3 spp., SW and S Madagascar, in xerophytic vegetation, ca. 50–500 m.

§ sect. Cheirolepidium ‣ two spp., from N Africa through Central Asia; fallow fields and dry, open habitats, 500–1,500 m.

§ sect. Crossadenia ‣ two subsections.

o subsect. Apparicianae ‣ 4 spp., E. flaviana Carn.-Torres & Cordeiro; E. sobolifera O.L.M. Silva & P.J.Braun; E. appariciana Rizz. and E. teres M. Machado & Hofacker, all endemics to Bahia state, Brazil, growing on granitic domes (inselbergs) or sandstone outcrops, 250–1,200 m.

o subsect. Ephedropepplus ‣ 8 spp., 7 endemics to NE Brazil in Tocantins, Bahia, Goiás, Minas Gerais, Distrito Federal, Pernambuco and Piauí states, in rocky grasslands vegetation on sandy substrates and Atlantic sandy coastal shrublands (restingas), sea level to 1,400 m, and E. riinae V.W.Steinm. from Bolivia and Mato Grosso state in Brazil; E. gymnoclada Boiss. is the only aphyllous in this group.

§ sect. Denisiae ‣ two spp., southern Madagascar, in xerophytic vegetation, sea level to 200 m.

§ sect. Eremophyton ‣ 3 spp., Australia, New Caledonia, Vanuatu; coastal sands to inland desert and scrub, sea level to 600 m.

§ sect. Espinosae ‣ two spp., S and E Africa (Angola, Botswana, Namibia, N South Africa, Kenya, Malawi, Tanzania, Zambia, Zimbabwe); hilly, deciduous woodlands, 300–1,400 m.

§ sect. Frondosae ‣ 7 spp., E to southern Africa (Angola, Botswana, Ethiopia, Kenya, Malawi, Mozambique, Namibia, South Africa, Tanzania, Uganda, Zambia, Zimbabwe) and Arabian Peninsula (Oman, Saudi Arabia, Yemen); open to dense bushland, forest, 400–2,700 m.

§ sect. Gueinziae ‣ only E. gueinzii Boiss., South Africa (Mpumalanga, Free State, KwaZulu-Natal, E Cape), Lesotho, and Swaziland; grasslands on rocky slopes and above sandstone cliffs, 200-2,000 m.

§ sect. Plagianthae ‣ two spp., southern Madagascar, in xerophytic and semi-xerophytic vegetation; sea level to 1,000 m.

§ sect. Poinsettia ‣ 4 subsections.

o subsect. Lacerae ‣ two spp., W & C Mexico; xerophytic scrub, wooded ravines, 900–2,500 m.

o subsect. Erianthae ‣ only E. eriantha Benth., SE U.S.A. (Arizona, California, New Mexico, Texas) and northern Mexico (Baja California, Baja California Sur, Chihuahua, Coahuila, Durango, Sonora); desert scrub and thorn scrub on rocky slopes and along washes, sea level to 900 m.

o subsect. Extipulatae ‣ two spp., SW U.S.A. to S Mexico; desert scrub, grasslands, oak forest, riparian areas.

o subsect. Stormieae ‣ 21 spp., widespread in the New World, from Canada to Argentina, but with a center of distribution in Mexico; in a wide variety of habitats from desert scrub to moist montane forests, sea level to 2,700 m; two spp. in Brazil, E. zonosperma Müll. Arg endemic and the widely E. heterophylla L.

§ sect. Scatorhizae ‣ 7 spp., Africa (Angola, Ethiopia, Kenya, Somalia, Tanzania), Arabia (Saudi Arabia, Yemen), India; rocky or sandy scrub or deserts, 200–1,850 m.

§ sect. Tenellae ‣ 4 spp., Southern Africa (Angola, Botswana, Namibia, South Africa, Zimbabwe); in open desert areas, exposed gravelly or sandy soils and rocky slopes, ca. 100-1,100 m.

§ subg. Euphorbia - 661 spp. worldwide, 21 sects., eight occur in South America.

§ sect. Brasilienses ‣ 5 spp. from Espírito Santo, Bahia, Minas Gerais, Paraiba and Pernambuco, in rocky uplands (campos rupestres), grasslands, thorn scrub, and tropical deciduous forest; E. phosphorea Mart. is the largest growing succulent Brazilian spurges.

§ sect. Calyculatae ‣ 2 spp., Mexico (Jalisco, Michoacán, Guanajuato, Guerrero, Mexico, Oaxaca, Puebla); deciduous montane woodlands, oak and pine oak forest, sometimes on lava flows, 1,600–2,300 m.

§ sect. Crepidaria ‣ 15 spp., Mexico (highest diversity), Central America, Caribbean, northern South America; in South America only one, E. tithymaloides L., in over Caribbean Basin up to Mexico, Florida, Colombia, Venezuela and Guyana; desert scrub, as well as dry and wet tropical forests, sea level to ca. 800 m; unique members of genus with nectar spurs.

§ sect. Cubanthus ‣ two subsections.

o subsect. Cubanthus ‣ 6 spp., Caribbean (Bahamas, Cuba, Jamaica); scrub forest mostly on limestone, sea level to 500 m.

o subsect. Moa ‣ 3 spp., E Cuba; scrub and forests on serpentine soils, sea level to 800 m.

§ sect. Denisophorbia ‣ 13 spp., Madagascar, Mayotte and Comoro Islands, Seychelles, Mauritius.

§ sect. Deuterocalli ‣ 3 spp., widespread in Madagascar, mostly in dry scrub or among rocks in moister habitats, sea level to ca. 500 m.

§ sect. Euphorbia ‣ 244 spp., widespread across most of Africa, Arabian Peninsula, and in southern Asia from Pakistan to Malaysia, Indonesia and Papua New Guinea; in a wide variety of habitats, but especially in arid landscapes, open areas, dry forests, scrub, rock outcrops, with some species occurring in moist forests.

§ sect. Euphorbiastrum ‣ 6 spp., Venezuela, Colombia, Ecuador, Peru, to southern Mexico and Caribbean (Windward Islands); montane scrub and forest edges, lowland moist forests.

§ sect. Goniostema ‣ 76 spp., widespread across the island of Madagascar; in a wide variety of habitats from sea level to upper mountains.

§ sect. Lactifluae ‣ only E. lactiflua Phil., Northern Chile, rocky areas of Atacama Desert from the region of Antofagasta south to the vicinity of Copiapó; nearly sea level to 700 m.

§ sect. Mesophyllae ‣ only E. sinclairiana Benth., from Mexico (Chiapas), to Colombia, Peru and Brazil (Acre); understory of wet, lowland forests but also extending to higher elevations in cloud forest, from near sea level to ca. 800 m

§ sect. Monadenium ‣ 90 spp., E, C, and SE tropical Africa; in a variety of mostly arid habitats: thickets, rock outcrops, and other open areas.

§ sect. Nummulariopsis ‣ 37 spp., SE U.S.A., southern South America (Argentina, Bolivia, Brazil, Chile, Paraguay, Peru, Uruguay); high montane habitats, grasslands, and sand dunes (South America), pine woodlands and sandy areas (U.S.A.), sea level to ca. 3,500 m; 10 spp. in Brazil, 7 endemics.

§ sect. Pachysanthae ‣ 4 spp., Madagascar; in a variety of habitats, such as thickets in xeric bush of the southeast, tsingy (karst) formations in the north, remnant forests in the high plateaus in the northwest, and rainforest relicts of the central east.

§ sect. Pervilleanae ‣ 7 spp., widespread across Madagascar.

§ sect. Pacificae ‣ 11 spp., Australia, Indonesia, New Guinea, Philippines, and Pacific Islands (Norfolk Island, Fiji, Solomon Islands, Hawaii); the Australian herbaceous or pencil-stemmed species occur in dry interior or coastal habitats, whereas the remaining species occur in tropical or subtropical moist forests or scrub.

§ sect. Portulacastrum ‣ 2 spp., Chile and Bolivia, montane habitats.

§ sect. Rubellae ‣ 3 spp., NE Africa (Sudan, Ethiopia, Kenya, Uganda); open sites in limestone crevices on well-drained soil, or under bushes in evergreen or deciduous bushland.

§ sect. Stachydium ‣ 6 spp., northern South America to E Brazil, absent in Mexico; low deciduous forest and thorn scrub on sandy or loose stony soils or rocky outcrops; 200-1,200 m; two spp. in Brazil, E. comosa Vell. in Brazil, Colombia and Venezuela and E. heterodoxa Mull. Arg. is endemic.

§ sect. Tanquahuete ‣ two spp., W & C to S Mexico; subtropical deciduous forests in montane canyons and lava flows; near sea level to 2100 m, species that are by far the tallest members of Euphorbia in the New World, and among the largest in the genus.

§ sect. Tirucalli ‣ 25 spp., Widespread and most diverse in Madagascar, also native in Arabian Peninsula (Oman and Yemen, including Socotra) and Africa (Angola, Namibia, Somalia, and South Africa). E. tirucalli L. is widespread across Africa, but it is not clear where the species is native (probably Madagascar and southern Africa). It is widely cultivated in India and in other tropical countries.

4.3 EUPHORBIOIDEAE ▸ TRIBE HIPPOMANEAE (29/340) - outsiders Homalanthus (23; tropical Asia to New Guinea, N and E Australia, New Caledonia, New Zealand, Polynesia); Grimmeodendron (2; Caribbean), Bonania (7; Caribbean), Balakata (2; S China, tropical Asia, New Guinea, tropical Australia), Falconeria (1; Himalayas, SE Asia, W Malesia), Sclerocroton (6; tropical Africa, Madagascar), Triadica (3; East and tropical Asia), Dalembertia (4; Mexico, Guatemala), Excoecaria (c 40; Africa to Australia and Melanesia).

52. Actinostemon Mart. ex Klotzsch. Monoecious trees or shrubs; latex scanty; inflorescence terminal and sometimes axillary. 19 spp., one only in Caribbean, remaining 18 in South America, two up to Central America and Caribbean, 17 in Brazil, 11 endemics.

53. Adenopeltis Bertero ex A.Juss. Monoecious shrubs, latex non recorded. Only one sp., A. serrata (W.T. Aiton) I.M. Johnst., endemic to C Chile.

54. Algernonia Baill. Monoecious trees or shrubs, stems and branches unarmed; latex white; inflorescense terminal. 12 spp., 11 restricted of dense forests of SE Brazil, centered in Rio de Janeiro state, and A. amazonica (Emmerich) G.L. Webster restricted of Amazon rainforest from Brazil and adjacent Peru in Huanuco departament.

55. Colliguaja Molina. 5 spp., C. brasiliensis Klotzsch ex Baill. from southern Brazil, Paraguay and Uruguay, and four only in Chile and adjacent Argentina.

56. Dendrothrix Esser. 4 spp., two only from Venezuela to Peru, D. yutajensis (Jabl.) Esser from Venezuela to Ecuador and Brazil and D. wurdackii Esser known only in Brazil, from Amazonas state, and is a rare plant in Brazil, by Plantas Raras do Brasil’s book.

57. Gradyana S. M. Athiê-Souza, A. L. Melo & M. F. Sales. Strictly axillary sinuous thyrsoid staminate inflorescences, solitary pistillate flowers, apical glands in the bracts, long-stipitate glands between pistillate sepals, and staminate flowers with three stamens and three sepals. Only one spp., G. franciscana S. M. Athiê-Souza, A. L. Melo & M. F. Sales, found in the São Francisco river valley in Alagoas and Sergipe states, in the semiarid region of NE Brazil.

58. Gymnanthes Sw. c 25 spp., tropical Africa, Malesia, 21 in tropical America, 11 in South America, 8 in Brazil, 4 endemics.

59. Hippomane L. High poisonous trees. Three spp., two restricted from Hispaniola and H. mancinella L. from Florida, the Caribbean to Venezuela, Colombia, Galápagos Islands.

60. Hura L. Monoecious trees up to 40 m tall, thunk spiny; latex white, common in lowland dioecious forests. Two spp., H. polyandra Baill. from Mexico to Nicaragua, with a disjuntc record in Ecuador, and H. crepitans L. from Nicaragua to Bolivia, Brazil and Caribbean.

61. Incadendron K. J. Wurdack & Farfán. Only one sp., I. esseri K. J. Wurdack & Farfán, from southern Ecuador and northern Peru.

62. Mabea Aubl. Monoecious trees and shrubs; latex whitish; hairs dendritic and brownish to reddish, staminate flowers pedicellate with numerous (up to 90) stamens each, ovaries and fruits tomentose, pistillate flowers with long styles. 40 spp., restricted from lowland in Mexico to SE, Brazil (27, 4 endemics), 37 in South America.

63. Maprounea Aubl. Monoecious (dioecious) trees or shrubs; latex whitish; inflorescence terminal. 5 spp., two in Africa and three in over tropical South America, all in Brazil, with M. brasiliensis A. St.-Hil. endemic.

64. Microstachys A.Juss. Monoecious perennial herbs with several short erect stems arising from a thick, much-branched xylopodium; latex white; inflorescences terminal on main axes; fruit 3-lobed, septicidally dehiscent into 3 bivalved cocci, each valve with a small appendage; seeds subcylindric, smooth. 22 spp., 5 in Africa, one in SE Asia and Australia, and 16 in South America, 15 in Brazil (one endemic to Paraguay), with nine endemics, one up to also Mexico and Caribbean.

65. Ophthalmoblapton Allemão. Monoecious trees, stems and branches unarmed; latex white or yellow, highly toxic; indumentum absent; inflorescence axillary and spiciform. 4 spp. from E Brazil, only from Bahia to Rio de Janeiro, in Atlantic Forest, one of them, from Bahia state, is a rare plant in Brazil, by Plantas Raras do Brasil’s book.

66. Pachystroma Müll.Arg. Monoecious trees and shrubs; latex white. Only one sp., P. longifolium (Nees) I.M. Johnst., from southern Brazil, Bolivia and southern Peru.

67. Pleradenophora Esser. 6 spp., five in Mexico, one of then up to southern U.S.A., and P. membranifolia (Müll. Arg.) Esser & A.L. Melo, in Amazon rainforest lowlands and mesophilous forests of Peru and Bolivia, extending into Brazil in Goiás, Mato Grosso, Mato Grosso do Sul, Minas Gerais and São Paulo states.

68. Pseudosenefeldera Esser. Only one sp., P. inclinata (Müll. Arg.) Esser, in Amazon rainforest, from Venezuela to Peru and northern Brazil.

69. Rhodothyrsus Esser. Two spp. in northern South America, from Suriname to Peru and northern Brazil (only R. macrophyllus (Ducke) Esser, non endemic).

70. Sapium Jacq. Woody species, sometimes with xylopodium; leaves simple with petiolar glands, staminate flowers subsessile with 2 stamens each, seeds with a red aril. 25 spp. from tropical America, 13 in South America, 11 in Brazil, only two endemics. S. laurifolium (A.Rich.) Griseb. from S Mexico to Brazil is a myrmecophite.

71. Sebastiania Spreng. Woody plants with simple elongate inflorescences, the staminate flowers subsessile and with 3 stamens and 3 free sepals, the fruits regular with a notably thin wall. 32 spp., tropical America, 22 in South America, 11 in Brazil, 6 endemics.

72. Senefeldera Mart. Monoecious trees; latex white; inflorescence axillaty. 4 spp., two from Venezuela to Peru, S. triandra Pax & K. Hoffm. in Amazon rainforest of Brazil and Peru, and S. verticillata (Vell.) Croizat in Atlantic Forest of E Brazil.

73. Senefelderopsis Steyerm. Monoecious trees and shrubs; latex white. Two spp. restricted to the Guiana Shield, from Colombia to Guyana.

74. Spegazziniophytum Esser. Monoecious, succulent shrubs, branches transformed into thorns. Only one sp., S. patagonicum (Speg.) Esser, endemic to dry areas of Argentina.

75. Stillingia Garden ex L. Sapium-like but with more variation in habit (subshrubs to trees or succulents), the petioles eglandular, the fruits with a characteristic woody persistent cornute base, seeds dry without an aril. 30 spp., Madagascar, Mascarene Islands, East Malesia, Fiji, 25 in tropical and subtropical America incl. southern U.S.A.; 12 in South America, 10 in Brazil, 8 endemics.

37. GERANIALES

TWO FAMILIES, BOTH IN SOUTH AMERICA.

GERANIACEAE

§ FULLY CHLOROTROPHIC

Genera/species 7(excludes Rhynchotheca)/c. 835 Distribution cosmopolitan except polar areas, especially in temperate and subtropical regions, with their largest diversity in South Africa; few species in tropical regions. Habit bisexual, usually perennial or annual herbs (sometimes suffrutices or shrubs; in Geranium section Neurophyllodes trees), some species are xerophytic stem succulents and a large number are geophytes, often with root or stem tubers. Family compound only herbs. No have simple leaves. Flowers of five petals and five sepals.

In Geranium the inflorescence is a cyme sometimes reduced to 1-flowered cymules born on a reduced stem, or more commonly composed of 2-flowered cymules alongated stem; in some cases these cymules are grouped in pseudoumbel aggregates towards the end of the branches; in Erodium inflorescences are composed of pseudoumbels at the end of each branch; all Neotropical Geranium species have 10 fertile stamens; Erodium has 5 fertile stamens (alternate with the petals) and 5 staminodes (opposite the petals).

SYSTEMATIC two clades, both in South America, one exclusive.

1. HYPSEOCHARIS CLADE (1/7) ‣ a single genus.

1. Hypseocharis Remy. Perennial acaulescent herbs with thick taproots or tubers. 7 spp., from the Andes, from S Peru to Argentina.

2. TRIBE GERANIEAE (5/c 830) ‣ outsiders 5/c 830. Pelargonium (c 250; Africa, Madagascar, Asia, St. Helena, Tristan da Cunha, Australia, New Zealand, with their highest diversity in South Africa), Monsonia (c 40; Africa, Madagascar, SW Asia, with their highest diversity in southern Africa), California (1; S Oregon, California, Baja California in NW Mexico).

2. Erodium LHér. Annual or perennial herbs, ocasioanlly shrubs. 74 spp. and is distributed on all continents, excluding Antarctica; a major centre of diversity is observed in Mediterranean region region (62), whereas, the other continents harbour only a few native species: one in North America; 5 in Australia; 4 in Asia; and E. geoides A.St.-Hil. in South America, disturbed sites on sandy soils, 0–1,100 m, Peru to Rio Grande do Sul state in S Brazil and Argentina.

3. Geranium L. Annual or perennial herbs, mostly herbaceous perennials with aerial stems, ocasioanlly shrubs, some annuals, and some form small cushions or dwarf shrubs; leaves usually palmatifid or palmatisect, although a few species have digitate, tridentate, or even entire leaves; all species have actinomorphic, pentamerous flowers with ten stamens and the distinctive beaked fruit typical for the family. c. 350 spp., distributed in almost every continent and ecosystem, being absent only in the poles, arid deserts, and low-altitude tropical areas; it is present in most important mountains of tropical America, and some dry areas near the sea, although it is absent in lowland tropical forest; two species of subg. Geranium are native in the New World as well as elsewhere: G. erianthum DC. occurs also in E Asia, and G. sylvaticum L. is widely distributed in Europe and W Asia. 126 spp. in New World, 84 spp. in South America, only 7 in Brazil; the greatest diversity is found in Mexico (36) and Peru (34). The Andes in Colombia and Ecuador also harbor many endemic species, but fewer species are found in other parts of South America as well as in Central America and North America north of Mexico; in New World occur three subgenera:

§ subg. Geranium ‣ c. 150 spp. in South America, only three in Brazil, G. core-core Steud., scattered in South America, G. albicans A. St.-Hil. from S Brazil and Uruguay, and the endemic G. thunbergii Siebold ex Lindl. & Paxton; G. dissectum L. is a weed distributed worldwide.

§ subg. Robertium ‣ none species in South America; 8 sections.

§ subg. Erodioidea ‣ 22 spp. in with four section: one in E Africa, two from Europe and adjacent Asia, and Brasiliensia, with 3 spp., Brazil and adjacent Uruguay: G. arachnoideum A. St-Hill is widely distributed in grasslands and dry places of three states of S Brazil; G. glanduligerum R. Knuth occur disjunct in marshy places of Rio Grande do Sul/Santa Catarina and São Paulo states, also in Uruguay; G. brasiliense Progel is endemic of damp places and forest margins in nebulous highlands of Rio de Janero state; all three spp. are perennial herbs with erect habit, inflorescences a monochasial cymes, purplish corollas, and grows in open places.

FRANCOACEAE

§ FULLY CHLOROTROPHIC

Genera/species 7(includes Rhynchotheca)/40 Distribution tropical and southern Africa. Greyia: South Africa, with their largest diversity in eastern Transvaal and Drakensberg; Francoa and Tetilla: mountains in central Chile; three remaining genera in South America. Habit usually bisexual (in Bersama functionally polygamodioecious), evergreen or deciduous small trees or shrubs (Bersama) or suffrutices (Melianthus). Evergreen or deciduous trees or shrubs (Greyia) or rhizomatous perennial herbs (Francoa, Tetilla), sometimes perennial or annual herbs. In Rhynchotheca and some species of Viviania many short shoots are modified into spines. Use Ornamental plants, medicinal plants, timber, carpentries, aromatic substances (incense wood).

SYSTEMATICS tribe Bersameae (2/8, Africa) and Greyeae (1/3, South Africa and Eswatini) are South Africans; Vivianeae and Francoeae are restricted of South America.

1.1 TRIBE FRANCOEAE (2/2) ‣ both genera in South America.

1. Francoa Cav. Only one sp., F. appendiculata Cav., endemic to center-south Chile.

2. Tetilla DC. Only one sp., T. hydrocotylifolia DC., endemic to center Chile.

1.2 TRIBE VIVIANIEAE (4/18) ‣ all genera in South America.

3. Balbisia Cav. Shrubs up to 2m tall; inflorescence thyrsoid; leaves opposite, entire, flowers bowl-shaped, with yellow, sometimes yellowish-green or reddish-yellow corolla, fruit capsular, subtended by pair of deeply divided bracts. 11 spp., Chile, Peru, Bolivia, Argentina.

4. Rhynchotheca Ruiz & Pavon. Shrubs up to 2m tall; leaves opposite, not-entire, apetalous flowers and may be unique anemophyllous spp. in Geraniales. Only one sp., R. spinosa Ruiz & Pav., endemic to interandean valleys of Peru and Ecuador.

5. Viviania Cav. Shrubs up to 4 m tall, or perennial or rarely annual herbs; inflorescence thyrsoid or cymose; campanulate, white to pinks flowers, pedicellate without bracts directly at base, fruit capsular. 6 spp., 4 endemics to Chile, V. albiflora (Cambess) Reiche, in Argentina, Uruguay and S Brazil, and V. linostigma R. Kunth, from SE Santa Catarina to extreme NE Rio Grande do Sul state, endemic to Araucaria Forest formations in the ‘Aparados da Serra Geral’; the two extra-Andean species the two species could be reliably differentiated by a trait such as tetramerous flowers for V. linostigma versus pentamerous flowers for V. albiflora.

38. MYRTALES

FAMILIES ABSENTS IN SOUTH AMERICA: CRYPTERONIACEAE (3/12) AND PENAEACEAE (3/29).

LINEAGE 1 of 5: COMBRETACEAE

COMBRETACEAE

§ FULLY CHLOROTROPHIC

Genera/species 10/510–520 Distribution mainly tropical regions of Africa (with their highest diversity), subtropical southern Africa, Madagascar, Seychelles, tropical South and Southeast Asia, southeastern China, Malesia, New Guinea, Melanesia, western Pacific islands, northern and eastern Australia. Habit usually bisexual (rarely andromonoecious or dioecious), evergreen or deciduous trees, shrubs, suffrutices or lianas (Lumnitzera and Laguncularia are mangrove trees with pneumatophores). Bark often exfoliating. Sometimes xerophytic.

The family stands out economically for its ornamental value, with some species commercialized by florists worldwide. Others are cited in the literature as having pharmacological potential and being widely used as popular diuretics or antipyretics. A number of species exhibit antimicrobial, antihemorrhagic and antiulcer activities. Conocarpus erectus L. and Laguncularia racemosa (L.) Gaertn. are typical of mangrove areas and are ecologically essential to the dynamics of this ecosystem and to the survival of its organisms. Laguncularia racemosa shows several adaptations to daily flooding by salt water: the leaves contain salt excretory glands, the seeds germinate while still attached to the tree (vivipary), and the roots include pneumatophores.

SYSTEMATIC two subfamilies, Strephonematoideae (1/3, tropical West and Central Africa) absent in South America; two tribes within Combretoideae, both in South America.

Key to genera of South American Combretaceae

1. Inflorescences globular clusters; fruit aggregated ------------ Conocarpus
1. Inflorescence spicate, racemose or paniculate; fruit not aggregated - 2

2. Lowers receptacle with 2 adnate bracteoles ------------ Laguncularia
2. Lowers receptacle without adnate bracteoles - 3

3. Leaves opposite; anthers versatile; trees or shrubs often scandent ------------ Combretum
3. Leaves alternate or verticillate; anthers versatile or adnate to the filaments; trees ------------ Terminalia

1. COMBRETOIDEAE ▸ TRIBE COMBRETOIDEAE (5/475–510) - outsiders Getonia (1; tropical Asia), Guiera (1; N tropical Africa).

1. Combretum Loefl. Herbs, shrubs, lianas and trees, up 25 in C. leprosum Mart. and C. glaucocarpum Mart. from Brazil. 255 spp., pantropical, in New World from Mexico, Mesoamerica, Caribbean, South America; one sp. no have petals; Africa has 163 spp., New World only 33 (28 in South America, 21 in Brazil, only two endemics, inc. C. rupicola Ridl. in rare in Brazil by Plantas Raras do Brasil’s book, known only in Fernando de Noronha island), Asia has 27 and 1 in Australia.

2. Conocarpus L. Shrubs or trees (mangroves); bark fissured; branches subangular to narrowly winged; leaves alternate, petiolate, somewhat fleshy; flowers bisexual, small, crowded in dense, cone-shaped heads borne in racemes; petals absent; stamens (5)7(10), in 2 whorls, exserted; filaments filiform, glabrous; disc long-pubescent; ovary inferior, 1-locular; ovules 2; fruit a trapezoidal, samara-like, glabrous, brownish maroon achene with a corky pericarp; seeds cylindric, curved, whitish. Two spp., C. erectus L., considered a mangrove, from North America to Peru, Brazil and Caribbean, but its lack of vivpary or pneumatophores suggests that it is best considered a ‘mangrove associate’, and C. lancifolia Engl. a tree in sandy soils of NE Africa and Arabia.

3. Terminalia L. Trees up to 45 m tall; leaves spiral, often with pocket-shaped or bowl-shaped domatia. c. 190 spp., cosmopolitan genus of tres up to 50 m tall, centered of SE Asia, in New Word from Mexico, Mesoamerica, South America; 62 spp. in New World, 47 in South America, 36 spp. in Brazil, 9 endemics.

2. COMBRETOIDEAE ▸ TRIBE LAGUNCULARIEAE (4/10) - outsiders Lumnitzera (2; coasts in tropical East Africa, Madagascar and Seychelles to islands in the Pacific), Macropteranthes (5; Northern Territory, Queensland), Dansiea (2; E Queensland).

4. Laguncularia Gaertn. Polygamous shrubs or small trees (mangroves); branchlets glabrous; leaves opposite, petiolate, oriaceous to fleshy, obovate or elliptic, glabrous; flowers bisexual or unisexual, sessile, subpaniculate; brpetals absent, or minute and caducous; stamens 10, in 2 rows, included; ovary inferior, 1-locular, crowned by an epigynous disc; ovules 2; fruit coriaceous, crowned by calyx limb, shortly club-shaped, subtrigonous, ribbed, shortly pubescent; seed oblong-ovoid. Only one sp., L. racemosa (L.) C.F. Gaertn., E and W tropical America (Florida and Mexico up to Ecuador and Brazil), W tropical Africa.

LINEAGE 2 of 5: MYRTIIDS

LYTRACEAE

§ FULLY CHLOROTROPHIC

Genera/species 28/575–585 Distribution cosmopolitan except polar areas, especially tropical and subtropical regions in the Northern and Southern Hemispheres, with their largest diversity in the Americas. Habit usually bisexual (in Capuronia dioecious; in Sonneratia and Duabanga sometimes monoecious?), evergreen trees or shrubs, perennial or annual herbs (rarely lianas). Many species are hygrophytes, some representatives are aquatic (Sonneratia comprises mangrove trees with pneumatophores, producing vertical anchor roots, and horizontal nutrition roots; Trapa is aquatic). Bark often exfoliating. Young stems and branches usually quadrangular in cross-section. Use ornamental plants, fruits (Punica granatum L., Trapa), vegetables (Trapa), dyeing substances (Lawsonia inermis L.), medicinal plants, timber.

Key differences from similar families Onagraceae differ from Lythraceae by their viscid pollen threads and, excluding Ludwigia, also by epigynous flowers with inferior ovaries.

SYSTEMATIC 3 subfamilies, all in South America.

Key to genera of Neotropical Lythraceae

1. Leaves glandular-punctate, the punctae non-secretory and orange-filled, turning black or secretory and translucent - 2

2. Leaves translucent-punctate, secretory, blades varnished by resin; inflorescences 3-flowered axillary cymes ------------ Lourtella

2. Leaves orange to black-punctate, non-secretory; inflorescences multi-flowered clusters - 3

3. Flowers in compact umbelliform clusters; floral tube campanulate, greenish; capsule indehiscent ------------ Adenaria

3. Flowers in loose cymose clusters; floral tube cyathiform, deep red; capsule loculicidally dehiscent ------------ Pehria

1. Leaves non-punctate, glabrous or variously indumented - 4

4. Floral tubes campanulate to globose or cyathiform, about as long as to slightly longer than wide, actinomorphic - 5

5. Seeds encircled by a broad, thin wing - 6

6. Floral tubes caducous, distally pleated, (8-)10-12(-16)-merous; leaves with subapical abaxial porate chamber ------------ Lafoensia

6. Floral tubes persistent, distally smooth, 6- or 8-merous; leaves normally developed at the apex, lacking a porate chamber - 7

7. Trees with strongly divaricate branching; flowers 8-merous; placenta appearing free -central ------------ Physocalymma

7. Shrubs and subshrubs with opposite branching; flowers 6-merous; placenta bipartite, septa lunate ------------ Diplusodon

5. Seeds not winged - 8

8. Trees, shrubs, or subshrubs, 1-40 m - 9

9. Stamens basifixed; petals cream-coloured; capsules indehiscent ------------ Crenea p.p.

9. Stamens dorsifixed; petals bright yellow, rose, purple, or white - 10

10. Petals bright yellow; flowers solitary; pedicels 0-4 mm ------------ Heimia

10. Petals rose, purple, or white; flowers solitary and/or in axillary clusters; pedicels 10-40 mm ------------ Ginoria

8. Annual or perennial herbs, 2 cm - 2m, commonly less than 50 cm - 11

11. Capsules septicidally dehiscent, wall microscopically (10x) transversely striated; leaves decussate or whorled; ------------ Rotala

11. Capsules splitting irregularly or initially circumscissile, then irregular; wall microscopically smooth, without striations - 12

12. Capsules splitting irregularly; leaf base auriculate ------------ Ammannia p.p.

12. Capsules initially circumscissile, then splitting irregularly; leaf base cordate ------------ Nesaea

4. Floral tubes cylindrical, at least twice as long as wide, actinomorphic or zygomorphic - 13

13. Floral tubes actinomorphic; capsules dehiscing from the apex ------------ Lythrum

13. Floral tubes slightly to strongly zygomorphic, bilateral; capsules indehiscent or dehiscent dorsally by a longitudinal slit together with floral tube - 14

14. Stamens attached near base of floral tube, anthers basifixed; capsules indehiscent, placenta and seeds retained within ------------ Pleurophora

14. Stamens attached at midlevel or higher in floral tube, anthers dorsifixed; capsules dehiscent, placenta and seeds exserted ------------ Cuphea

1. SUBFAMILY LYTHROIDEAE (5/87) ▸ outsider Decodon (1; E U.S.A.).

1. Heimia Link. Slender-stemmed, glabrous shrubs with lanceolate-linear leaves. Three spp., H. salicifolia Link, from Texas to Argentina and Caribbean (hallucenogenic, anti-inflammatory), H. montana (Griseb.) Lillo in Bolivia and Argentina, and H. apetala (Spreng.) S.A. Graham & Gandhi in Brazil adjacent Cono Sur.

2. Lythrum L. Herbs or small shrubs with opposite, alternate or verticillate leaves. 35 spp., wet places, nearly worldwide distribution especially in Europe, 11 spp. in New World, only L. maritimum Kunth in South America, along coasts from Colombia to S Brazil, Venezuela to Chile, Argentina, Asia, and North America.

3. Rotala L. Glabrous, aquatic, amphibious, or terrestrial herbs with decussate or verticillate leaves. 40 spp. worldwide, primarily African and Asian, two also in North and Central America to Argentina and Brazil, both widely distributed in almost all countries.

2. SUBFAMILY PUNICOIDEAE (14/355–360) ▸ outsiders Woodfordia (2; NE Africa, S Arabian Peninsula; Madagascar, tropical Asia to S China and Timor), Koehneria (1; S Madagascar), Pemphis (1; tropical regions in the Old World), Punica (2; Socotra; SW Asia), Capuronia (1; Madagascar), Galpinia (1; S Africa north to Zimbabwe and Mozambique).

4. Adenaria Kunth. Srubs or small trees. Only one species, A. floribunda Kunth, from Mexico to Brazil, Argentina, widely distributed, in evergreen forests.

5. Cuphea P. Browne. Herbs or subshrubs, some microphyllous and ericoid, with diverse indumentum, mainly tubular flowers, sometimes with xylopodium. 256 spp., restricted in New World, South America (162) up to Argentina, Central America, Mexico, Caribbean, and U.S.A., centered in Brazil (109 spp., 71 endemics, mainly in center dry savanas) and Mexico, mainly in open, mesophytic habitats, uplands; within Brazil, the states of Minas Gerais, Goiás and Bahia contain the largest number of species, many of which are narrowly restricted endemics (26 spp. are rare plants in Brazil, by Plantas Raras do Brasil’s book, in all regions of country).

Graham & Kleiman (1987) believe that the primary centre of speciation for Cuphea is found in the upland regions of eastern Brazil, with a major secondary centre in Mexico. Recent research developments in Cuphea shows that these plants are sources of medium-chain unsaturated fatty acids such as caprylic acid, capric acid, lauric acid and myristic acid; these substances are important new sources of raw materials for the chemical industry in the manufacture of detergents, lubricants and other products, since, at present, these substances are extracted almost exclusively from the oil of Cocos nucifera L. and other members of the Palmae.

6. Diplusodon Pohl. Shrubs or subshrubs with showy, sometimes with xylopodium, 6-merous, actinomorphic flowers and floral tubes on which the sepals are alternate with conspicuous epicalyx segments; their capsular fruits have a bipartite placenta with lunate septa, unique in the family, and the seeds are winged. 93 spp. from Brazilian savannas, almost all endemic to center Brazil (all states with cerrado vegetation except Paraná and Pará, the largest number as micro-endemics of a single mountain, but centered in dry areas and savannas in Espinhaço Range), except by two, both also reported for Brazil, in adjacent E Bolivia, D. virgatus Pohl, unique white flowered spp. of this genus, and D. bolivianus T.B. Cavalc. & S.A. Graham, first record of this genus outside Brazil, described in 1996.

43 spp. are rare in Brazil are rare plants in Brazil, by Plantas Raras do Brasil’s book, one the largest number of a single genus, in Mato Grosso, Goiás, Bahia, Tocantins and Minas Gerais, and also in Distrito Fedral.

In Diplusodon there is a high degree of narrow endemism among the species restricted to specialized microhabitats, in mountains at 1,000–1,600 m elev. with rough topography, where they grow on sandy and rocky soils among rocky outcrops and slopes; the species are isolated by geographical barriers and climate in rocky grasslands (campos rupestres), the mountains of the east and midwest of the country. Species isolation has been a significant factor in production of morphological novelties within the group.

Phylogenetic relationships inferred from nuclear ITS and ETS sequences strongly support the monophyly of Diplusodon, with four major clades, three of which are strongly correlated with endemism and are centered in the Espinhaço Range, Goiás highs, and the Diamantina highs in Bahia state; the fourth clade was designated the Cerrado clade (C), containing species from all three centers of diversity as well as widely distributed species and several species of lower-elevation cerrado.

7. Gyrosphragma T.B.Cavalc. & M.G.Facco. Srubs or small trees, spurless, weakly actinomorphic floral tubes, two deep red petals, a stipitate ovary and irregularly circumscissile dehiscence of a thickened capsule. Two spp. in rocky outcrops and grasslands, in Minas Gerais, Espírito Santo and Minas Gerais states in SE Brazil.

8. Lafoensia Vand. Glabrous shrubs or trees, the leaves with subapical, abaxial porate chamber, flowers 16-merous. 6 spp. scattered in South America, one up to Mexico, some in Andes; mainly montane forests and savannas of C Brazil (cerrado); 4 spp. in Brazil, two endemics.

9. Lourtella Graham, Baas, & Tobe. Shrubs to 4m, pith enclosing a large central secretory duct. Only one species, L. resinosa S.A. Graham, P.Baas & H. Tobe, N Peru and disjunctly in S Bolivia, in dry open deciduous woods with cacti.

10. Pehria Sprague. Shrubs or small trees, stems often flushed wine-red. Only one species, P. compacta (Rusby) Sprague, Honduras, Nicaragua, Venezuela, Colombia, in secondary vegetation, pastures, roadsides, margins of forests at low to mid-elevations.

11. Physocalymma Pohl. Trees, with strongly divaricate, subopposite, 4-angled branches, showy pink flowered. Only one sp., P. scaberrimum Pohl, restricted of central South America in Ecuador, Brazil, Peru and Bolivia, in dry forests and savannah.

12. Pleurophora D. Don. Herbs to 1m, indumentum of unicellular eglandular and multicellular glandular hairs. 8 spp. in two subgenera:

§ subg. Anisotes ▸ three spp., P. saccocarpa Koehne in Mato Grosso do Sul, Argentina, Bolivia and Paraguay, and two endemics to dry dry, rocky habitats and fields and in the seasonally dry forests of semiarid regions in NE Brazil.

§ subg. Pleurophora ▸ 5 spp. with reduced spiny parts in arid regions of Bolivia, Chile and Argentina.

3. SUBFAMILY LAGERSTROEMIOIDEAE (8/115–125) ▸ outsiders. Lagerstroemia (50–55; tropical Asia, N and E Australia), Duabanga (3; tropical Asia), Sonneratia (c 20; coasts of the Indian and Pacific Oceans), Trapa (1; warm-temperate regions in Europe, Africa and Asia); Ginoria (14; Mexico, Caribbean), Tetrataxis (1; Mauritius), Lawsonia (1; North Africa, tropical regions in the Old World).

13. Ammannia L. (inc. Crenea) Glabrous ubshrubs to small trees, annual herbs, aquatic or marsh-inhabit. c. 27 spp., worldwide tropical to temperate distribution, especially Africa to SE Asia, 9 spp. in New World, 6 in South America, all widely distributed, 4 in Brazil, with A. maritima (Aubl.) S. A. Graham, P. W. Inglis, & T. B. Cavalc. of Caribbean, Trinidad & Tobago, and northern South American coasts from Venezuela continuously into N Piauí state, and disjunct in Bahia and São Paulos states.

ONAGRACEAE

§ FULLY CHLOROTROPHIC

Genera/species 22/625–635 Distribution: cosmopolitan except Antarctica, with their largest diversity in SW North America. Habit usually bisexual (rarely unisexual), usually perennial, biennial or annual herbs (rarely trees or shrubs, e.g. in Fuchsia). Some species are aquatic. The family is cosmopolitan. Most genera are restricted to western North America or have their major distribution there. North America genus occurs mainly in Baja California.

Although these genera may occur in regions with climate similar to tropical Mexico, their geographic distribution is above the Tropic of Cancer and they were not considered here as Neotropical. The phylogenetic relationships and evolution of Onagraceae have been intensively studied and the family is considerably well-known in many aspects like: cytogenetics, palynology, physiology, ecology, anatomy and morphology. However, there is a lack of recent taxonomic treatments for many groups.

SYSTEMATIC two subfamilies, both occur in South America.

1. SUBFAMILY JUSSIAEOIDEAE (1/c 85) ▸ only one genus in this subfamily.

1. Ludwigia L. A very morphological diverse genus, usually perennial herbs of damp ground or shallow water; stems creeping or floating to erect, up to treelets; leaves entire or minutely toothed; flowers 4- or 5-merous, axillary; stipules often prominent; hypanthium absent; flowers tetramerous or pentamerous. c. 86 spp., cosmopolitan, with their largest diversity in North and South America (74 in New World), from Mexico to South America (50, mainly in wet areas); 48 in Brazil, 10 endemics; among the 22 sections of Ludwigia, 13 are monotypic, a indication of the sharp distinctions among the species; the remaining nine sections comprise more than one species:

§ nine sections, with joined 11 spp., are only Old World, mainly in Africa.

§ sections Oligospermum, Macrocarpon and Seminuda (joined 18 spp.) are mixed Old and New World, all of these from Brazil.

§ 8 sections exclusively from New World occur in South America, all in Brazil, with three very restricteds: sect. Cinerascentes (1, Suriname and N Brazil), sect. Amazonia (1, northern South America) and Anastomosans (1, endemic to Brazil, composed only by L. anastomosans (DC.) H. Hara, the largest species in genus - a treelet up 10 m tall, very rare species known from few scattered individuals restricted to black water streams of the ancient Brazilian Shield).

§ sects. Ludwigia, Isnardia and Microcarpium (joined 23 spp.) from North America to Caribbean, mainly in E U.S.A.

2. SUBFAMILY ONAGROIDEAE (21/540–550) ▸ outsiders Hauya (2; Central America); Circaea (7; temperate regions on the Northern Hemisphere), Lopezia (22; Mexico, Central America), Megacorax (1; Mexico); Gongylocarpus (2; Mexico, Central America); Chamaenerion (8; temperate regions on the Northern Hemisphere); Xylonagra (1; Baja California); Taraxia (c 10; North America, Mexico), Chylismiella (1; U.S.A.); Eulobus (4; California, Arizona, Baja California), Eremothera (7; W North America, western Mexico), Neoholmgrenia (2; W North America), Camissoniopsis (14; SW U.S.A., NW Mexico), Tetrapteron (2; W U.S.A., NW Mexico).

2. Camissonia Link. 12 spp., 11 in U.S.A. (one up to Mexico), and C. dentata (Cav.) Reiche from Peru to Chile and Argentina.

3. Clarkia Pusch. 42 spp., 41 North America south to Mexico, esp. California, and C. tenella (Cav.) H.F. Lewis & M.R. Lewis from Peru, Chile and Argentina.

4. Fuchsia L. Mostly mesic shrubs, erect, scandent or climbing. 105 spp., confined to cool, moist habitats, with nearly 3/4 concentrated in the tropical Andes, in 10 sections grouped in five near disjunct ranges.

§ sect. Eclliandra ▸ 6 spp., Mexico to Central America.

§ sect. Ellobium ▸ 3 spp., Mexico to Central America.

§ sect. Fuchsia ▸ 61 spp., tropical Andes, Hispaniola, only F. boliviana Carrière up to Argentina.

§ sect. Hemsleyella ▸ 14 spp., tropical Andes.

§ sect. Jimenezia ▸ only one sp., Panamá and Costa Rica.

§ sect. Kierschlegelia ▸ only one sp.,endemic to Chile.

§ sect. Quelusia ▸ 9 spp., 8 from highlands of SE Brazil, and a single F. magellanica Lamarck, from S Chile and Argentina; F. regia (Vell.) Munz, is widely distributed and covers the entire range of the section in Brazil; the remaining species have much more restricted distributions, several occurring on a single mountain range.

§ sect. Schufia▸two spp., Mexico to Central America.

§ sect. Skinnera ▸ 4 spp., New Zealand and Tahiti.

5. Gayophytum A.Juss. 9 spp., 7 in North America, one in both North America and Cono Sur, and one restricted for temperate South America.

6. Epilobium L. Herbaceous plants, either perennial or annual; leaves mainly opposite. c. 160 spp., temperate region, in tropics restricted of montate environments, including Neotropics up to Argentina; 50 spp. in New World, 14 spp. in South America, 11 from NW Venezuela to S Argentina and Chile, and E. hirtigerum A. Cunn., restricted from S Brazil, NE Argentina and Uruguay.

7. Oenothera L. Annual or perennial herbs; leaves forming a basal rosette; flowers in leaf axils or in ± distinct inflorescences, radially symmetrical, 4-merous, opening near sunset or near sunrise. 149 spp., from Mexico to Argentina (a few in Central America), including S Brazil (8, one endemic); 46 in South America; the smallest species is O. acaulis Cav. from Chile (10 cm), and the largest is O. stubbei W. Dietr., P.H. Raven & W.L. Wagner from Mexico (3m).

LINEAGE 3 of 5: VOCHYSIACEAE/MYRTACEAE

VOCHYSIACEAE

§ FULLY CHLOROTROPHIC

Genera/species 8(own data, excludes Ruitzerana)/c. 220 Distribution mainly tropical South and Central America; Erismadelphus and Korupodendron: tropical West Africa. Habit bisexual, usually evergreen trees or shrubs (sometimes lianas, rarely herbs). Vochysiaceae is very common in the Brazilian Shield. Some species are tolerated in urban arborization, but rarely cultivated. Aluminum accumulators.

SYSTEMATIC four tribes, all in South America.

1. TRIBE SALVERTIEAE (1/1) ▸ a single genus.

1. Salvertia A.St.-Hil. Tree or shrub, leaves verticillate, with white flowers with nectar spurs. Only one sp., S. convallariodora A. Saint Hillary, endemic to of savannah in Brazil, E Bolivia, with small populations also in Suriname.

2. TRIBE VOCHYSIEAE (1/142) ▸ a single genus.

2. Vochysia Aubl. Large trees or shrubs (sometimes few 0.3 m, with leaves at top of branches), few to many branched, sometimes xylopodium, or great trees with very beautiful yellow flowers with nectar spurs, up to 60 m tall, the tallest of all Myrtales in Brazil. 147 spp. from Mexico (2) to Brazil and Bolivia, 144 in South America, 89 in Brazil, 51 endemics; 11 spp., mainly in Amazonas and Minas Gerais states, are rare plants in Brazil, by Plantas Raras do Brasil’s book. V. vismiifolia Spruce ex Warm. (northern South America, inc. Brazil) is the unique to myrmecophyte in family.

§ sect. Vochysiella

§ subsect. Decorticantes ▸ a very well-known taxon characterized by an ovary with a well-developed indumentum, flower buds larger than 1.3 cm, and glabrous, pilose, or glaucous leaves with irregular venation; the taxon includes ten species distributed mainly in savannas of C Brazil (cerrado).

§ subsect. Calophylloideae

§ sect. Ciliantha

§ subsect. Micranthae

§ subsect. Lutescentes

§ subsect. Ferrugineae

§ subsect. Discolores

§ subsect. Chrysophyllae

§ subsect. Megalanthae

§ sect. Pachyantha ▸ nine spp.

3. TRIBE ERISMEAE (3/18) ▸ outsiders Erismadelphus (1; tropical West and Central Africa), Korupodendron (1; Central Africa).

3. Erisma Rudge. Emergent or canopy trees up to 40 m tall; hairs stellate; leaves opposite, flowers with nectar spurs. 16 spp., in Central and South America (15, one is endemic to Panamá), mostly in Brazilian Amazon rainforest, also in Venezuela to Peru, and Guianas; 14 spp. in Brazil, 3 endemics, two in Amazonas and one in Espirito Santo state, all rare plants in Brazil, by Plantas Raras do Brasil’s book. Two sections.

§ sect. Erisma ▸ near a half of species of genus.

§ sect. Rixa ▸ near a half of species of genus.

4. TRIBE QUALEEAE (3/79) ▸ three genera, all in South America.

4. Callisthene Mart. Trees or shrubs; cataphylls often present at the base of branchlets and inflorescences; flowers with, flowers with nectar spurs. 11 spp. in Brazil (all species, 7 endemics, two, in Rio de Janeiro and Minas Gerais states, are rare plants in Brazil, by Plantas Raras do Brasil’s book), remainig 4 up to Bolivia, C. hassleri Briq. also in Paraguay. Two sections.

§ sect. Callisthene ▸ all genus except C. fasciculata Mart.

§ sect. Cataphyllantha ▸ only one sp., C. fasciculata Mart., from Brazil and Bolivia.

5. Mahechadendron Cortés, Cortés & Alonso. Trees, with highly durable wood; inflorescences in compound dichasia, compact, only developed laterally; flowers tiny, the tiniest flowers known in the Vochysiaceae; the flowers also have the single petal non-clawed; fruit capsular. Only one sp., M. puntecascarillo Cortés, Cortés & Alonso, endemic to moist forests of the Middle Magdalena River in Colombia.

6. Qualea Aubl. (inc. Ruizterania). Trees to shrubs, up to 30 m tall, with only have one petal but it is very pretty, yellow, white or blue, with nectar spurs. 65 spp., all in over tropical South America, 53 spp. in Brazil, 18 endemics; 6 spp. are rare plants in Brazil, by Plantas Raras do Brasil’s book, 3 in Amazonas, 2 in Pará and one in Minas Gerais state. Two subgenera.

§ subg. Qualea

§ sect. Qualea

§ sect. Trichanthera

§ sect. Polytrias

§ sect. Costatifolium

§ subg. Amphilochia

MYRTACEAE

§ FULLY CHLOROTROPHIC

Genera/species 127/5,560–5,600 Distribution tropical, subtropical and warm-temperate regions especially in the Southern Hemisphere, with their largest diversity in Australia, rare in African tropics; only Myrtus in Europe. Habit usually bisexual (rarely andromonoecious, polygamomonoecious, dioecious, or androdioecious), usually evergreen (rarely deciduous) trees or shrubs, all species are woody, with essential oils, and flower parts in multiples of four or five. Often aromatic. Bark often exfoliating. Numerous species are xerophytes. Osbornia consists of mangrove trees.

Genera with capsular fruits are absent from the Americas, apart from the Metrosideros species in Chile and Argentina. Eucalyptus is a dominant, nearly ubiquitous genus in the more mesic parts of Australia and extends north sporadically to Philippines. Eucalyptus regnans F. Muell. is the tallest flowering plant in the world. The genera Heteropyxis and Psiloxylon have been separated as separate families by many authors in the past as Heteropyxidaceae and Psiloxylaceae. The Myrtaceae represent one of the most striking ecological and taxic radiations of the Gondwanan flora.

Mexico has 11 genera and 108 spp., only the Central America Chamguava not in Brazil; Myrcia, Myrcianthes, Psidium, Pimenta and Ugni are monotypic in this country; Chamguava, Mosiera and Myrciaria are diotypic; Calyptranthes has 19 spp., and Eugenia, 78; only 131 spp. in Colombia. All genera listed are native, and all but Eugenia are endemic to the Neotropics.

Ornamental plants, fruits (Psidium, Feijoa, Campomanesia, Eugenia, etc.), spices (Eucalyptus, Pimenta, Syzygium), gums, essential oils for perfumes, medicinal plants (Eucalyptus), timber, paper pulp (Eucalyptus), drainage of swamps (Eucalyptus, Metrosideros). Feijoa sellowiana (O. Berg) O. Berg, the Pineapple Guava or Feijoa, cultivated for its edible fruits. Psidium guajava L. (guava), Eugenia uniflora L. (pitanga, Suriname and Brazil) and Myrciaria cauliflora (Mart.) O. Berg (jaboticaba, Brazil, Argentina and Paraguay) are widely cultivated for fruit, jams and juices. Myrciaria dubia (Kunth) McVaugh (camu-camu) is a common edible fruit in Amazon rainforest region. Syzygium jambos (L.) Alston and various Eucalyptus species are introduced, cultivated (for fruit and timber/shade respectively) and are often naturalized.

Of particular note are the tall, shrubby eucalypt species known as ‘mallees’ that have multiple stems arising from an underground stem sometimes called a ‘lignotuber’, the two southern African species of Eugenia and a number of Brazilian (some also Bolivian) species from various genera found in savannas of C Brazil (cerrado, as Psidium, Campomanesia and Eugenia) that are rhizomatous subshrubs; these all appear to be adaptations to fire-prone habitats.

Metrosideros commonly has tree species that produce large numbers of adventitious roots, and certain of these have been recorded as beginning their lives as strangling epiphytes; also, a number of species of Metrosideros sens. lat. from New Zealand and Papua New Guinea are root-climbing lianes; there is only a single mangrove in the family, the monotypic Osbornia, a shrub or small tree that lacks pneumatophores.

SYSTEMATIC two subfamilies, Psiloxyloideae (2/4, Central and SE Africa, Mascarene Islands) absent in South America; among Myrtoideae, 13 small lineages do not occur in South America, all of them in SE Asia, S China, Malesia to New Guinea, Australia (high centered), Tasmania, New Caledonia (joined 79 genera and c. 2,400 spp.) except Syzygieae (2/500), which reaches to Africa and Hawaii; two tribes in New World.

1. MYRTOIDEAE ▸ TRIBE METROSIDEREAE (1/55) - a single genus.

1. Metrosideros Banks et. Gaertn. (inc. Tepualia) Shrubs or small trees, capsular fruits (unique among all New World Myrtaceae). 56 spp., mainly Pacific islands, including Philippines and New Guinea, with an outlying species in South Africa and one, M. stipularis (Hook. & Arn.) Hook. f., in southern Chile and Argentina.

2. MYRTOIDEAE ▸ TRIBE MYRTEAE (49/2.630–2.640) - all lineages in South America except Decasperminae (12/193, Austro-Pacific and Southeast Asia).

∎ GENERA INCERTAE SEDIS

2. Amomyrtella Kausel. Tree; hairs simple; inflorescences monads. Two spp. endemics to Ecuador.

3. Amomyrtus (Burret) Legrand & Kausel. Shrubs or trees; hairs simple. Two spp. from Chile, just into W Argentina.

∎ SUBTRIBE BLEPHAROCALYCINAE ▸ a single genus

4. Blepharocalyx O. Berg. (exc. Temu) Trees or shrubs; hairs simple; inflorescence uniflorous or with 3 to ca. 35 flowers in a dichasium or a panicle with dichasial subunits; flowers 4-merous; fruit crowned by a square scar; seeds 1 to ca. 11, 4-6 mm long. 4 spp. from Caribbean, Brazil (all species, two endemics), Venezuela, Bolivia and Paraguay, Chile and Argentina; in Brazil B. eggersii (Kiaersk.) Landrum occur only in Amazonas state, and B. salicifolius (Kunth) O. Berg, is a extremely variable species of SE and S Brazil up to Ecuador.

∎ SUBTRIBE EUGENIINAE ▸ outsiders Calyptrogenia Burret (6; Caribbean) and Hottea (c 9; Cuba, Hispaniola).

5. Eugenia L. (inc. Plinia p.p.) [2th BR] Trees or shrubs, sometimes with woody rhizomes; hairs simple or dibrachiate; inflorescence traitorous, a bracteate shoot, or rarely a dichasium, the bracteate shoots sometimes with a long or abbreviated axis and from 2 to many flowers; flowers 4(-5)-merous; calyx open or closed and tearing regularly or irregularly; fruit crowned by the calyx-lobes, remnants of the calyx, or by a circular scar. 1,192 spp., SE Asia and the Pacific (c. 14 and 35, respectively), Africa and neighboring spp. (c. 120), 973 in New World, only 4 in North America, mainly in South America with 592 species, and its highest diversity is found in Brazil where 471 spp. can be found, of which 378 are endemics, only 4 spp. from several states are rare plants in Brazil, by Plantas Raras do Brasil’s book.

E. reperta Sobral & Mazine (Minas Gerais state, Brazil) has the largest fruit of this genus, up to 10 cm long

§ sect. Calycorectes ▸ c. 28 spp., Mexico to South America; it is more diverse in the Amazon, Southern Brazil, Paraguay and Argentina and rare in Uruguay.

§ sect. Eugenia ▸ c. 30 spp., most of them restricted to Brazil, distributed throughout the country, mainly concentrated in the Northeast, Midwest, Southeast and South. E. ligustrina (Sw.) Willd. occurs from Central America to Southern Brazil.

§ sect. Hexachlamys ▸ c. 10 spp., S and SE Brazil, Bolivia, Paraguay, Argentina and Uruguay.

§ sect. Phyllocalyx ▸ c. 16 spp., NE Brazil to Argentina, Bolivia and Paraguay, occurring frequently in Atlantic Forest and sparsely in Cerrado biome.

§ sect. Pilothecium ▸ c. 21 spp., Guianas, Colombia and Ecuador to Argentina.

§ sect. Pseudeugenia ▸ c. 22 spp. from Guianas and Suriname to Bolivia, Paraguay and Southern Brazil in Santa Catarina state.

§ sect. Racemosae ▸ c. 59 spp., Mexico and the Antilles throughout South America to Uruguay and northern Argentina.

§ sect. Speciosae ▸ c. 15 spp., widely distributed in the Atlantic Forest, from eastern Brazil to Paraguay and N South America.

§ sect. Umbellatae ▸ c. 700 sp. from S Mexico, Cuba and the Antilles to Uruguay and Argentina.

6. Myrcianthes O. Berg. Trees or shrubs; hairs simple; inflorescence uniflorous or more often a dichasium of 3-7 flowers; flowers 4-5-merous; calyx open; ovary usually 2-1ocular; ovules a few to numerous per locule; fruit crowned by the calyxlobes; seeds 1-2. 40 spp., 38 in South America, mainly Andean, extending from Mexico to Chile, 8 reaches into Brazil, 3 endemics.

7. Pseudanamomis Kausel. Trees; branches glabrous. Inflorescence almost umbel-like, with 3 or more flowers. One widely distributed species, P. umbellulifera (Kunth) Kausel, Trinidad and Tobago, Guyana, Venezuela and Colombia.

∎ SUBTRIBE LUMINAE ▸ all genera in South America.

8. Luma A. Gray. Shrubs or small trees; hairs simple; inflorescence of monads, triads or dichasia. Two spp., from Chile and Argentina.

9. Myrceugenia O. Berg. (exc. Nothomyrcia) Trees or shrubs; inflorescence usually uniflorous, less often a dichasium or a bracteate shoot; flowers 4-merous; calyx open or, in 2 spp., closed and falling as a calyptra; fruit crowned by the calyx-lobes or by a circular scar. 48 spp. growing from temperate and subtropical Brazil and Cono Sur to S Chile; 34 spp. in Brazil (30 endemics, 4 of then, in SE & S Brazil, are rare plants in Brazil, by Plantas Raras do Brasil’s book).

10. Nothomyrcia Kausel. (off Myrceugenia) Trees. Only one sp., N. fernandeziana (Hook. & Arn.) Kausel, endemic to endemic to Massatierra, Juan Fernandez Islands, Pacific Chile.

11. Temu O. Berg. (off Blepharocalyx). Only one sp., T. cruckshankii (Hook. & Arn.) O. Berg, from Cono Sur.

∎ SUBTRIBE MYRCIINAE ▸ a single genus.

12. Myrcia DC. ex Guillemin. [3th BR] (inc. Calyptranthes, Marlierea) Trees or shrubs; hairs simple or dibrachiate; inflorescence a panicle; flowers 5-merous (rarely 4-merous in Amazonas state); calyx open, or closed in the Marlierea complex; bracteoles deciduous; ovary 2-3(-4)-locular; ovules 2 per locule; fruit crowned by the calyx-lobes or remnants of the calyx; seeds 1-2; seed coat membranous to crustaceous. 798 spp., Mexico and Caribbean south to Argentina, 600 in South America, the third largest genus in Brazil, with 455 spp., 364 endemics, only 7 spp., in several states, are rare plants in Brazil, by Plantas Raras do Brasil’s book. M. madida McVaugh, endemic to Peru, is a myrmecophyte.

§ sect. Aguava ▸ 32 spp., all tropical biomes including very wet and dry habitats.

§ sect. Aulomyrcia ▸ 140 spp., Amazon forest, the Guiana Shield, Caribbean and the Atlantic coastal forests (particularly Bahia and Espírito Santo), extending to associated drier habitats

§ sect. Calyptranthes ▸ 277 spp., moist forests (Amazon and Atlantic) and cerrado (including gallery forest) of Central and South America and throughout the Caribbean; relatively few species extending to associated drier habitats.

§ sect. Eugeniopsis ▸ 22 spp., Atlantic Forest with occasional occurrences in campos rupestres.

§ sect. Gomidesia ▸ 57 spp., Atlantic coastal and associated lowland, montane and gallery forests and cerrado, extending to the Amazon and Caribbean.

§ sect. Myrcia ▸ 120 spp., over range of genus.

§ sect. Reticulosae ▸ 23 spp, Atlantic coastal forests, cerrado and campos rupestres.

§ sect. Sympodiomyrcia ▸26 spp., Atlantic Forest, cerrado (campo rupestre and riparian forest, not savanna), and a disjunct distribution in the Guiana Shield.

§ sect. Tomentosae ▸ 9 spp., over range of genus.

∎ SUBTRIBE MYRTINAE ▸ outsiders Chamguava (3; S Mexico, Central America), Myrtus (2; M. communis: the European Mediterranean; M. nivellei: North Africa);

13. Accara Landrum. Shrubs; essentially glabrous; hairs when present simple; inflorescence uniflorous; flowers 4-merous; fruit crowned by the calyx-lobes; seeds ca. 40, ca. 3 mm long; seed coat hard, shiny; embryo C-shaped, the hypocotyl and cotyledons about equal in length. Only one sp., A. elegans (DC) Landrum, a shrub very narrow endemic to dry areas in Caraça Mountains, center Minas Gerais state, in rocky grasslands (campos rupestres), a rare plant in Brazil, by Plantas Raras do Brasil’s book.

14. Calycolpus O. Berg. Trees and shrubs; hairs simple or two-armed. 16 spp., Panamá to Peru and Trinidad, Guyana and N Brazil up to Minas Gerais state, 15 in South America, 14 restricteds; 9 spp. grows in Brazil, 4 endemics; C. australis Landrum from Minas Gerais state is a rare plant in Brazil, by Plantas Raras do Brasil’s book.

∎ SUBTRIBE PLINIINAE ▸ all genera occur in South America.

15. Algrizea Proença & NicLaugh. Shrubs; leaves opposite; inflorescences 3–flowered dichasia, long–pedunculate, solitary or in synflorescences, at upper nodes; flowers white, pentamerous, of small size for the family, with bracts and bracteoles persistent in the open flowers, sometimes persistent in young fruit. Two spp., both endemic to highland areas of the Diamantina Range, Bahia state, Brazil, sometimes amongst rocks or near streams, on sandy, clay, and laterite soil., in sandy soils of rocky grasslands (campos rupestres).

16. Myrciaria O. Berg. rees or shrubs; hairs simple; bark reddish, deciduous; inflorescence a bracteate shoot often reduced to a glomernle, often cauliflorous; flowers 4-merous; calyx open; fruit crowned by a circular scar; seeds 1-2; seed coat membranous; embryo a solid mass or with 2 separate, piano-convex cotyledons. 33 spp., Guatemala and Belize south to Paraguay and N Argentina, 31 in South America, mainly in Brazil (26, 17 endemics).

17. Neomitranthes Legrand. Trees or shrubs; hairs simple; inflorescence a bracteate shoot often reduced to a glomerule; calyx closed, falling as a calyptra, the staminal ring splitting at anthesis; petals 0- 4; ovary usually 2-1ocular; ovules 2-6 per locule; fruit crowned by a circular scar. 15 spp., Bahia to Rio Grande do Sul states of Brazil, in Atlantic forests, from Atlantic sandy coastal shrublands (restingas) to mountais.

18. Plinia L. (exc. Eugenia p.p.) Trees or shrubs, sometimes with woody rhizomes; hairs simple; inflorescence a bracteate shoot often reduced to a glomerule, the peduncles often subtended by conspicuous persistente bracts, often cauliflorous; flowers 4-merous, usually silky-pubescent; calyx closed or nearly so, tearing irregularly; fruit crowned by remnants of the calyx. 75 spp., Caribbean, Costa Rica south through tropical South America to Argentina; 43 spp. in South America, 37 in Brazil, 32 endemics; two spp. from E Brazil are rare plants in Brazil, by Plantas Raras do Brasil’s book.

19. Siphoneugena O. Berg. Trees or shrubs; hairs simple; inflorescence usually a bracteate shoot; flowers 4-merous; calyx open or closed, tearing irregularly or falling as a calyptra; fruit crowned by a circular scar; seeds 1-2. 11 spp., in Atlantic sandy coastal shrublands (restingas) from Brazil (9, 6 endemics) north to Venezuela, Guianas, two up to Caribbean to Puerto Rico and Central America, south into Argentina; S. guilfoyleiana Proença is known until now only from the Atlantic sandy coastal shrublands (restingas) in the state of São Paulo; S. delicata Sobral & Proença from Espirito Santo state is a rare plant in Brazil, by Plantas Raras do Brasil’s book.

∎ SUBTRIBE PIMENTIINAE ▸outsider Mosiera Small (34; Caribbean, Mexico, Central America and North America).

20. Acca O. Berg. (exc. Feijoa) Trees or shrubs; hairs simple; inflorescence uniflorous; flowers 4- merous; calyx open; petals reddish to pink, fleshy when mature; stamens ca. 60-90, stiff, red, more or less straight, erect in bud; fruit crowned by the calyx- lobes. Two spp. in the Andes of Peru, the latter to Bolivia.

21. Campomanesia Ruíz & Pavón. Trees or shrubs, sometimes with woody rhizomes; hairs simple; inflorescence uniflorous, a dichasium of 3(-15) flowers, or a bracteate shoot; flowers normally 5-merous; stigma usually capitate; fruit crowned by the calyx-lobes, remnants of the calyx, or by a circular scar. 45 spp., tropical and subtropical South America in Brazil (44, 33 endemics, C. paranensis D. Legrand only in Cono Sur) north to Venezuela and Trinidad, west to Colombia, Peru, and south to N Argentina. 5 spp. from E Brazil are rare plants in Brazil, by Plantas Raras do Brasil’s book.

22. Curitiba Salywon & Landrm. Trees or shrubs; hairs simple; inflorescence uniflorous or a bracteate shoot of usually no more than 2 flowers; flowers 4-merous; calyx-open; ovary 2-1ocular; ovules ca. 10 per locule; fruit elongate, angular, crowned by the calyx- lobes. Only one sp., C. prismatica (D. Legrand) Salywon & Landrum, from montane forests of Atlantic Forest of NE Santa Catarina and SE Paraná states in S Brazil (750–950 m elevation range).

23. Feijoa O. Berg. (off Acca). Only one sp., F. sellowiana (O. Berg) O. Berg, in S Brazil, Uruguay and NE Argentina.

24. Legrandia Kausel. Small tree; branchlets 4-angled; leaf lamina often with domatia below. Only one sp., L. concinna (Phil.) Kausel, endemic to Chile.

25. Myrrhinium Schott. Trees or shrubs; hairs simple; inflorescence a dichasium of 3- 7 flowers, solitary or aggregated in groups of 2 to ca. 8 on short bracteate shoots, often cauliflorous; flowers 4-merous; calyx open; petals red, pink, or purplish, fleshy when mature; fruit crowned by the calyx-lobes. Only one sp., M. atropurpureum Schott, with two varieties ranging from the Andes from Colombia to SE and S Brazil and Argentina.

26. Pimenta Lindl. Trees and shrubs; hairs simple; inflorescence a dichasium or a panicle of 3-15 flowers; flowers 4-merous; calyx open; ovary 2-1ocular; ovules 3- 6 per locule, on a subapical, stalk-like placenta; fruit crowned by the calyx-lobes. 18 spp., 17 in Caribbean from Cuba to Trinidad, Mexico (Veracruz, Oaxaca) to Panamá and Caribbean coast of Venezuela (only one), and P. pseudocaryophyllus (Gomes) Landrum disjunct in S Bolivia and C, S & SE Brazil.

27. Psidium L. Trees and shrubs; hairs simple; inflorescence uniflorous, a dichasium of usually no more than 3 flowers, or rarely a hracteate shoot; flowers normally 5-merous; fruit crowned by the calyxlobes, remnants of the calyx, or by a circular scar. 113 spp., Mexico and the Caribbean to northern Argentina, Galapagos and Revilagigedo, most of which grow in Brazil; 81 in South America, 66 in Brazil, 48 endemics; three spp. are naturalized in subtropical and tropical regions around the world. P. guajava L., ‘goiaba’, is commonly cultivated; P. macedoi Kausel from Goiás state is a rare plant in Brazil, by Plantas Raras do Brasil’s book.

∎ SUBTRIBE UGNINAE ▸ outsiders Lenwebbia (2; Queensland, E New South Wales), Lophomyrtus (2; New Zealand), Neomyrtus (1; New Zealand incl. Stewart Island)

28. Myrteola O. Berg. Subshrubs or shrubs; hairs simple. Tree spp., two from Colombia to Bolivia, and M. nummularia (Poir.) O. Berg in Colombia and Venezuela to S Chile and Argentina, as well as the Juan Fernandez and Falkland Islands, and populations in Mount Neblina in Amazonas state, Brazil.

29. Ugni Turcz. Shrubs, often densely branched, with small, coriaceous leaves; hairs simple; inflorescence uniflorous; flowers 5-merous, nodding; bracteoles persistent; calyx open; petals white, sometimes with a reddish spot in the center, together forming an ericoid, lampshade-like structure. 4 spp., U. myricoides (Kunth) O. Berg from Mexico and Guatemala to Guyana and Bolivia, growing also in high-montane habitats of Mount Neblina in Amazonas state; remaining three species in Cono Sur, Chile, Juan Fernandez Is.

LINEAGE 4 of 5: MELASTOMATACEAE

MELASTOMACEAE

§ FULLY CHLOROTROPHIC

Genera/species 167/4,955--5,035 Distribution mainly tropical and subtropical regions in the Southern and Northern Hemispheres, with their largest diversity in tropical South America. Habit usually bisexual (rarely androdioecious), evergreen trees or shrubs, perennial herbs (sometimes aquatic) or lianas. Many species are epiphytic. Young stems and branches often quadrangular in cross-section. Some microphyllous species (particularly in Microlicia) have leaves with a single mid-vein only, but can easily be assigned to this family by their distinctive stamens (poricidal anthers and prolonged, often appendaged connectives.

Nectar production is rare in Melastomataceae and most species are visited by pollen-gathering bees that use thoracic vibrations (buzz pollination) to expel the pollen through the anther pores. The characteristic anther appendages probably function as a hold for the bees legs.

Miconia calvescens DC. and Miconia crenata (Vahl) Michelang. (ex. Clidemia hirta D.Don) are aggressive weeds which have spread to the Pacific islands.

Ant domatia (myrmecophites) are known to occur in over 80 species of neotropical Melastomataceae spanning 10 different genera.

Anisophylly, or the presence of leaves of different size in each pair, in the Melastomataceae is not restricted to Miconia longidentata Michelang. & W. Meier., Miconia farinasii (Wurdack) Michelang. or Miconia flexuosa (Triana) Michelang., as it is also found in several other genera; in the Neotropics anisophylly has been reported in some species of Bertolonia, Blakea, Miconia, Macrocentrum and Triolena; the degree of anisophylly varies greatly among species, and in some cases the smaller leaves may even be caducous or apparently altogether lacking, conferring the plant an apparent alternate phyllotaxis. A recent study of anisophylly in selected species of Miconia in Eastern Brazil showed that this character varies greatly even within the same individual, and those authors stated that it may not be of great taxonomic value; however, within the species traditionally placed in Miconia at least 35 species show consistently marked anisophylly that does have taxonomic significance.

102 genera in New World; Stanmarkia and Heterocentron do not occur in South America. 3688 spp. in New World, 3011 in South America.

SYSTEMATIC three subfamilies, Kibessioideae (1/c 15, Indochina, Hainan, Malesia to N Queensland) absent in South America.

1. SUBFAMILY OLISBEOIDEAE (6/345–355) - outsiders Memecylon (105–110; tropical regions in the Old World), Warneckea (35; tropical Africa, Madagascar, Mauritius), Lijndenia (12; tropical regions in the Old World), Spathandra (1–6; tropical Africa, Madagascar).

1. Mouriri Aubl. Trees up to 30m tall, possibly the tallest of Melastomataceae in New World; peninervous leaves. 89 spp. from Mexico, C. America, Caribbean, tropical South America (74, 54 in Brazil, 24 endemics); the greatest number of species is found in the Amazon rainforest; the wood of some species is used for general construction; many species have fleshy, edible wild fruit and appear in local markets eg. M. pusa Gardner ex. Hook from Brazil.

2. Votomita Aubl. Peninervous leaves. 10 spp., 3 endemics to Brazil; Venezuela, Peru, Central America, Caribbean and Guianas one endemic each, V. orinocensis Morley in Venezuela and Colombia, and V. monadelpha (Ducke) Morley in Brazil and Guianas.

2. SUBFAMILY MELASTOMATOIDEAE (c 153/4,400–4,610) - all 18 lineages in South America except tribe Dissochateae (6/c. 190, tropical Asia); three tribal unplaced genera: Dinophora (1; tropical West and Central Africa), Stanmarkia (2; Mexico, W Guatemala) and Ochthocharis (14; tropical Africa, tropical Asia).

3. Centradeniastrum Cogn. Laxly branched shrubs or suffrutescent herbs; leaves opposite, decussate, long-petiolate, the blades of a pair slightly to markedly unequal in size, 3-7-nerved, the primary veins impressed adaxially and elevated abaxially; inflorescence a simple or compound dichasium with each flower subtended by a pair of sessile or short-petiolate bracteoles. Two spp. in Andes from Colombia to Peru.

MELASTOMATOIDEAE LINEAGE 1 of 6: CLADE ASTRONIEAE/HENRIETTEAE

2.1 MELASTOMATOIDEAE ▸ TRIBE ASTRONIEAE (4/156) – outsiders Astrocalyx (2; Philippines), Astronia (c 60; tropical Asia to islands in the Pacific), Astronidium (67; Borneo and Philippines to New Guinea, Fiji, Micronesia to Society Islands), Beccarianthus (c. 22; Borneo to New Guinea).

4. Tessmannianthus Markgr. Trees, up to 30m tall, possibly the tallest of Melastomataceae in New World. 7 spp., three of Central America, two in Colombia and two in Ecuador/Peru.

2.2 MELASTOMATOIDEAE ▸ TRIBE HENRIETTEEAE (3/77) – all genera in South America.

5. Bellucia Raf. 22 spp. from South America (only 4 reaching in Central America and Mexico), 18 in Brazil, 4 endemics; some species has wild edible fruit, and known as ‘tapirs guava’ (goiaba de anta).

6. Henriettea DC. Shrubs to trees, often cauliflorous; 67 spp. from Belize to Bolivia and Brazil, 45 in South America, 23 in Brazil, 6 endemics.

7. Kirkbridea Wurdack. Infloresences pendunculate. Two spp. endemics to Sierra Nevada de Santa Marta in Colombia, in elevation range from 1,600 m – 1,800 m, both known only by type collection.

MELASTOMATOIDEAE LINEAGE 2 of 6: LITHOBIUM

2.3 MELASTOMATOIDEAE ▸ TRIBE LITHOBIEAE (1/1) – a single genus.

8. Lithobium Bong. Herbs with rosulate leaves, rosettes less than 3cm in diameter, possibly the smallest of all Melastomataceae in New World; trimerous flowers; the most basal herb among Melastomataceae. Only one sp., L. cordatum Bong., Brazil, endemic to Minas Gerais state, in SE Brazil.

MELASTOMATOIDEAE LINEAGE 3 of 6: CLADE BLAKEEAE/MERIANIA/ERIOCNEMA/MICONIA

2.4 MELASTOMATOIDEAE ▸ TRIBE BLAKEEAE (3/c 105) – both genera in South America.

9. Blakea P.Browne. Trees, shrubs or woody vines, mainly epiphytic; anthers compressed laterally with 2 well-separated apical pores. 186 spp., tropical America, 127 in South America, highly centered in Colombia (72); only 4 spp. in Brazil (none endemics); 12 spp., from Costa Rica to Ecuador, are myrmecophites, nine in South America.

10. Chalybea Naudin. 11 spp., 7 endemics to highlands of Colombia, mainly Boyaca and Santander, and four remaining in Ecuador and Peru.

2.5 MELASTOMATOIDEAE ▸ TRIBE MERIANIEAE (c 16/230–240) - all genera in South America.

11. Adelobotrys DC. Climbings sometimes with adventicious roots, campanulate hypanthium, obtusely 5-lobed calyx, and five-locular capsule that dehisces to reveal a free central column. 33 spp., two in Central America, A. adscendens (Sw.) Triana widely distributed in tropical America, 30 only South America; 19 spp. in Brazil, 18 in Amazon rainforest (one endemic) and A. atlantica Schulman, the first and only known record of the genus in Atlantic Forest, known only city of Ilhéus collected in 1943, possibly extinct.

12. Axinaea Ruiz & Pav. Trees and shrubs. 41 spp. from Venezuela to Bolivia, one up to Costa Rica and Panamá, largely restricted to the Andes and centered in small area of N Peru and S Ecuador (25), growing mostly in very humid cloud-forest at altitudes between 1,200 and 3,800 m.

13. Centronia D.Don. 9 spp., W tropical South America up to Guiana Shield and N Brazil (3, none endemics, only in Amazonas state).

14. Graffenrieda DC. Trees, shrubs, or rarely lianas. 68 spp., only Caribbean, 61 in South America (54 restricted), slightly centered in Guiana Shield; 22 spp. in Brazil, 6 endemics.

15. Macrocentrum Hook.f. Herbs to small shrubs, sometimes droseroid, also sometimes viviparous in species from Venezuela and Guianas. 26 spp., one sp. is known only from N Venezuela, another from Andean Peru, 23 from Guiana Shield (20 endemics) from Colombia to French Guiana, three of then up to N Brazil mainly Roraima state and Ecuador, and a residual, and one endemic to Pará state, N Brazil.

16. Maguireanthus Wurdack. Only one sp., M. ayangannae Wurdack, endemic to the Guiana Shield of Guyana, at 1,000 – 1,500 elevation range.

17. Meriania Sw. Trees or shrubs; inflorescences usually terminal panicles; connective with a dorso-basal spur and sometimes with an ascending dorsal appendage also; seeds narrowly oblong-pyramidal; oblong or reniform with a foveolate testa. 131 spp. from Central America to SE Brazil, Caribbean, 121 in South America; 51 spp. in Colombia; 17 spp., in Brazil, 14 endemics (remaining three up to N South America): the centre of diversity in Brazil is mountains of Rio de Janeiro state; one spp. from Mount Aracá in northern Amazonas state, is a rare plants in Brazil, by Plantas Raras do Brasil’s book; five sections: Eumerianie, Umbellata, Pachymeriae, Davya and Adelbertia, the two lasts in Brazil:

§ sect. Adelbertia ▸ with by panicles few-flowered, calyx 5-lobed, external lobes distinct and toothed, internal lobes diminutive and appressed, and anthers with a dorsal, elongated, ascending appendage.

§ sect. Davya ▸ multifloral panicles, calyx with lobes inconspicuous and denticulate, or lobes absent, and anthers with a dorsal, elongated ascending appendage.

18. Salpinga Mart. ex DC. 11 spp. restricted in over Amazon rainforest, five in Brazil (two endemics).

2.6 MELASTOMATOIDEAE ▸ TRIBE ERIOCNEMEAE (3/7) - all genera in South America.

19. Eriocnema Naudin. Subcaulescent rhizomatous habit, rosulate herbs; capsular fruits, and anthers lacking appendages. Two spp., both rare plant in Brazil, by Plantas Raras do Brasil’s book, restricted of rocky places in Minas Gerais state.

20. Ochthephilus Wurdack. Only one sp., O. repentinus Wurdack, endemic to Pantepui Life Zone, in Mount Ayanganna; Guyana, at 1,400 – 1,600 m elevation range.

21. Physeterostemon R.Goldenb. & Amorim. 5 spp. endemics to Atlantic Forest of Bahia state in Brazil, two of them are rare plants in Brazil, by Plantas Raras do Brasil’s book; only E Brazilian genus with the combination of inferior ovaries and dry, indehiscent fruits.

2.7 MELASTOMATOIDEAE ▸ TRIBE MICONIEAE (1/1918) – a single genus.

22. Miconia Ruiz & Pav. [1th BR] Shrubs or trees, sometimes myrmercophytes, usually preferring very moist, hilly, and forested terrain, some are vines, hemiepiphytes, or epiphytic shrubs, sometimes with roots crown, and a few, such as M. poeppigii Triana (Central America to Bolivia and Brazil, possibly the tallest of Melastomataceae in New World) are large trees, up to 30 m; flowers in terminal panicles, 4-5-6, 8-merous (mainly 5-merous); inflorescense dichasia, scorpioid, glomerulate or spiciforme; sometimes with bluish fruits. 1,884 spp. (7^th largest worldwide), the largest genus in Brazil (579, 287 endemics), Colombia (541) and Venezuela (348), and the largest genus endemic to New World, and within family, ranging from N Mexico to Argentina, Caribbean and North America (only M. bicolor (Mill) Triana); 1,429 spp. in South America; eight species from SE Brazil (six only in Rio de Janeiro state, five epiphytes from ex-Pleiochiton) are rare plants in Brazil, by Plantas Raras do Brasil’s book.

Some species occur in areas with seasonal rainfall, and many grow in the perpetually wet lowland rain forests. A smaller number of species occur in deciduous forests and savannas, but Miconia is virtually absent in the dry South American Chaco and dry seasonal scrubland of NE Brazil (caatinga). Most species prefer acidic soils, but in the Antilles and northern Mesoamerica, a number of species grow on limestone or limestone-derived soils; the majority of the species are pollinated by bees that extract the pollen by vibrating the poricidal anthers from nectarless flowers, but there are records of pollination by bats, birds, flies, and wasps on nectar-producing flowers, whose anthers open by slits or large pores that do not need vibration to release the pollen. Agamospermy occur in some species and apparently is related to polyploidy and hybridization, although it is not clear how widely these mechanisms are distributed within the group.

Diocey occur in 37 spp. of Melastomataceae all in Miconia, mainly from northern Andes: 25 from Ecuador to Bolivia, 3 of them up to Colombia, 3 endemics to Venezuela, 5 from Mexico and Central America and 3 in Caribbean, one up South America, another up Central America - none in Brazil.

69 spp. of this genus are myrmecophtes, 25 in Brazil, 4 endemics.

MELASTOMATOIDEAE LINEAGE 4 of 6: CLADE BERTOLONIEAE/SONERILEAE

2.8 MELASTOMATOIDEAE ▸ TRIBE BERTOLONIEAE (1/18) – a single genus.

23. Bertolonia Raddi. Herbs, erect to prostrate, sometimes epiphytic; inflorscense scorpioid, terminal, flowers pentamerous, corolla white, sometimes pinkish or lilac in border; some species are viviparous. 34 spp., endemics to Brazil from Pernambuco, Bahia (15, center of diversity) to Santa Catarina states, mainly in shady forests.

2.9 MELASTOMATOIDEAE ▸ TRIBE SONERILEAE (38/700–790) - outsiders Amphiblemma (8–13; tropical Africa), Anerincleistus (30; India and S China to Philippines), Aschistanthera (1; Vietnam), Barthea (1; China inc. Taiwan), Blastus (9–12; Assam to West Malesia), Bredia (12–30; E and SE Asia), Calvoa (10–20; tropical Africa), Catanthera (11–17; Sumatra, Borneo, New Guinea), Cincinnobotrys (4–7; tropical Africa), Cyphotheca (1; Yunnan), Dicellandra (3; tropical Africa), Driessenia (14–18; West and Central Malesia), Fordiophyton (9–14; S China, SE Asia), Gravesia (c 110; tropical Africa, Madagascar), Kendrickia (1; S India, Sri Lanka), Kerriothyrsus (1; Laos), Medinilla (c 200; tropical Africa, Madagascar, tropical Asia to S China (inc. Taiwan) and New Guinea, NE Queensland, Fiji, Samoa), Neodriessenia (9; Borneo), Oxyspora (c 25; tropical Asia, S China), Pachycentria (10–15; Burma, Malesia), Phyllagathis (30–55; S China, SE Asia, W Malesia), Plethiandra (9; W Malesia), Poikilogyne (21; Borneo, New Guinea), Poilannammia (4; Vietnam), Preussiella (2; tropical West and Central Africa), Sarcopyramis (2; tropical Asia), Scorpiothyrsus (3–6; Hainan, SE Asia), Sonerila (c 175; tropical Asia), Sporoxeia (4–6; Burma, SE Asia), Stussenia (1; Vietnam).

24. Boyania Wurdack. Three spp., two endemic to the Guiana Shield of Guyana, 800-1,000 m elevation range, and one endemic to Colombia.

25. Neblinanthera Wurdack. Only one sp., N. cumbrensis Wurdack, endemic to the Guiana Shield of Venezuela and Amazonas state in Brazil, 1,300 – 1,800 m, of Mount Neblina.

26. Opisthocentra Hook.f. Only one sp., O. clidemioides Benth. & Hook. f. in Amazonas state in N Brazil and adjacent areas in Venezuela and Colombia.

27. Phainantha Gleason. 5 spp., tropical South America, 4 from Guiana Shield in Guyana and Venezuela, and one in S Ecuador.

28. Tateanthus Gleason. Only one sp., T. duidae Gleason, endemic to the Guiana Shield of Venezuela and Amazonas state in N Brazil, at 800 – 1,200 m elevation range.

29. Tryssophyton Wurdack. Two spp., endemics to the Guiana Shield of Guyana, at 1,100 -1,200 m, elevation range.

MELASTOMATOIDEAE LINEAGE 5 of 6: CLADE TRIOLENA/CYPHOSTILEAE/CAMBESSEDESIA

2.10 MELASTOMATOIDEAE ▸ TRIBE TRIOLENAE (2/44) - both genera in South America.

30. Monolena Triana. 16 spp., tropical America, centered in Panamá, 9 in South America, only M. primuliflora Hook. f. in Brazil, on Acre and Amazonas states.

31. Triolena Naudin. (inc. Diolena). Herbs to shrubs, sometimes viviparous in one sp. of northern South America. 28 spp., tropical America, 21 in South America, mainly Amazon rainforest; in Brazil occur only two spp., only in Acre state in Brazil territory.

2.11 MELASTOMATOIDEAE ▸ TRIBE CYPHOSTYLEAE (4/20) – all genera in South America.

32. Allomaieta Gleason. Strongly anysophyllous leaves and domatia inmmersed in the base of the leaf blade, one of Colombia is myrmecophite. 10 spp. endemics to mountains of Colombia.

33. Alloneuron Pilg. 6 spp., disjunct Colombia and Peru (3 endemics).

Within the Cyphostyleae, and among all Neotropical Melastomataceae, Alloneuron is unique in having leaves that have semicraspedodromous venation, and not the characteristic acrodromous venation found is most other species in the family. A. trinervium, is the only one in Alloneuron with three basal veins, a venation pattern that is intermediate between semi-craspedodromous (found in all other members of the genus) and acrodromous (predominant in the rest of the family).

34. Quipuanthus Michelangeli & C. Ulloa. Herbs with short stems growing on rocks, rosette-like; stems short, up to 10 cm with condensed internodes up to 5 mm, fleshy, pink inside (cut), completely concealed by a mix of densely packed coarse stramineous simple trichomes, 9–12 + ca. 0.5 mm and sparse, minute (< 0.1 mm) sessile glands; leaves with petioles 5–26 cm long, pinkish, the entire length covered by simple trichomes. Only one sp., Q. epipetricus Michelang. & C. Ulloa, known from two populations on the foothills of the E Andes mountains: one in the Cordillera de Galeras, Napo province (border with Orellana), Ecuador, and another in the Imaza district, province of Bagua, department of Amazonas, Peru.

35. Wurdastom B.Walln. Erect shrubs or trees, glabrous or with indumentum of barbellate, dendritic or amorphous trichomes; leaves op-posite, isophyllous and petiolate; venation acrodromous and regulary plinerved; inflorescences terminal or rarely axillary, thyrsoid, with helicoid branches, multiflorous. 8 spp., Colombia (5 endemics), Ecuador and Peru.

2.12 MELASTOMATOIDEAE ▸ TRIBE CAMBESSEDESIEAE (4/67) - almost endemic to Brazil.

36. Bisglaziovia Cogn. Shrubs, leaves papyraceous, flowers in a dichasia, axillary, corolla pentamerous. Only one sp., B. behurioides Cogn., very narrow endemic in two municipalities (Teresópolis and Macaeh de Cima) of center Rio de Janeiro state.

37. Cambessedesia DC. Subshrubs to shrubs, sometimes with xylopodium; inflorescence a simple dichasia, spiciform or thyrsoid, petals bicolor, red and yellow, rarely monocolor, fully red or fully yellow. 25 spp., from S Piauí to Paraná state, in E Brazil, mainly very narrow endemic to Espinhaço Range (MG), Diamantina Range (BA) and Veadeiros flatmountains (GO), mainly in rocky grasslands; only two are widely distributed, mostly endemic to savannas of C Brazil (cerrado) and especially diverse in rocky grasslands (campos rupestres) of C and NE Brazil; three occur in high altitud grasslands (campos de altitude) of the Brazilian Atlantic Forest biome; 10 are rare plants in Brazil, by Plantas Raras do Brasil’s book, all from Bahia, Minas Gerais and Goiás states.

38. Huberia DC. Shrubs to trees (rarely climbers); inflorescences terminal short panicles or depauperate cymes, or subapical solitary flowers; flowers pedicellate, tetra- to hexamerous; fruits capsular, regularly dehiscent or irregular (‘core’ Huberia), seeds pyramidate or winged and striate; H. bradeana Bochorny & R.Goldenb. is highly divergent duo be a climber with pentamerous flowers. 36 spp., 32 endemics to E Brazil, most of them are from high elevation grasslands more 1,000 m in the Atlantic Forest in E Brazil, with a few species occurring also at sea level, and 4 spp. from the Andes of Peru (1,200–3,350 m elevation, one up to Ecuador).

39. Merianthera Kuhlm. Shrubs; variable indumentum up to fully glabrous; leaves opposite, peciolate, caducous, acrodromous; flowers solitary, or in terminal panicles; 5-merous, corolla pink to purple; fruit capsular, many seeded; M. burlemarxii Wurdack is a myrmecophite. 8 spp. endemics to Brazil, largely endemic to granitic/gneissic inselbergs on rocky outcrops in E Brazil.

MELASTOMATOIDEAE LINEAGE 6 of 6: CLADE RUPSTREA/MICROLICIA/RHEXIA/MELASTOMEAE

2.13 MELASTOMATOIDEAE ▸ RUPESTREA CLADE (1/2) - A single genus; sister of Microlicieae + (Rhexieae + Melastomeae).

40. Rupestrea R.Goldenb., Almeda & Michelangeli. Shrubs, dry and indehiscent fruits that may be hydrochorous, and by the orthocampylotropous, monoembryonic seeds, each incompletely divided into two cavities by a septum. Two spp. restricted to rocky fields at the northern portion of the Diamantina Range in Bahia state, Brazil.

2.14 MELASTOMATOIDEAE ▸ TRIBE MICROLICIEAE (2/265) - both genera in Brazil, species almost exclusively found in E and C Brazil in rocky grasslands (campos rupestres); a few Rhynchanthera spp. are found farther to the north and SW and in wetter environments, and five Microlicia spp. are found in the Guiana Shield.

41. Microlicia D.Don. (inc. Chaetostoma, Lavoisiera, Stenodon, Trembleya) Shrubs or subshrubs, branched, often ericoid, erect or, more rarely, decumbent; ovary superior; fruit capsular, dehiscing longitudinally from apex to base; flowers solitary or in groups, mainly pink, often white or yellow (only in six species). 285 spp., 275 endemics to Brazil, M. benthamiana Triana ex Cogn. in Roraima state, Brazil, Venezuela and Guyana, M. insignis Cham. and M. windschii Versiane, D.Nunes & R. Romero in Bolivia and Brazil, seven outside country, endemics in Bolivia (3), Peru (2), Venezuela (1) and Colombia (1).

Pterolepis haplostemona Almeda & A.B. Martins and M. macedoi L.B. Sm. & Wurdack appear to be the only known Melastomataceae that are endemic to serpentine substrates in Brazil. 75 spp. are rare plants in Brazil, by Plantas Raras do Brasil’s book, the largest number for a single genus, mainly in Bahia, Minas Gerais or Goiás state.

42. Rhynchanthera DC. Shrubs or subshrubs, stems with glandular trichomes; leaves lanceolate to linear; inflorescense terminal and thyrsoid; flowers 5-merous, mainly pink, often white. 20 spp., Mexico to Bolivia, Paraguay and E Brazil, 19 in South America, 17 restricted within, 11 in Brazil, three endemics.

2.15 MELASTOMATOIDEAE ▸ TRIBE RHEXIEAE (3/23–26) - outsider Rhexia (11–13; E North America, S and SE U.S.A., the Caribbean), the only genera of this family in New World which occur in temperate North America, with most species being distributed from Texas to Nova Scotia; one widely distributed species is also found in Cuba, Hispaniola and Puerto Rico.

43. Arthrostemma Pav. ex D.Don. 4 spp., three only in Mexico and Central America, oe of then also in Venezuela, and A. ciliatum Pav. ex D. Don in Mexico, Caribbean, Central America, Venezuela, Colombia, Ecuador, Peru, Bolivia and N Brazil (Acre and Amazonas states), with a high dubious record in Rio de Janeiro state.

44. Pachyloma DC. 4 spp., endemics to the Guiana Shield of Amazon rainforest of Colombia to Guyana and N Brazil (2, no endemics), 50 – 1,200 m elevation range.

2.16 MELASTOMATOIDEAE ▸ TRIBE MELASTOMATEAE (c 46/840–950) - outsiders Heterotis (7; tropical Africa), Guyonia (14; tropical W and C Africa), Argyrella (6; tropical West, Central and SE Africa), Melastomastrum (4; tropical Africa), Tristemma (11–16; tropical Africa, Madagascar, Mascarene Islands), Dichaetanthera (30–35; tropical Africa, Madagascar), Dissotis (c 110; tropical Africa, Madagascar; polyphyletic), Amphorocalyx (5; Madagascar), Anaheterotis (1; Guinea, Sierra Leone), Dissotidendron (11; tropical W and C Africa), Dupineta (1–5; tropical West and Central Africa), Antherotoma (4; tropical Africa, Madagascar), Pseudosbeckia (1; tropical East Africa), Dionycha (3; Madagascar), Osbeckia (c 50; SE Asia, Malesia to New Guinea, N Australia), Melastoma (20–25; SE Asia, Malesia, N and E Australia), Heterocentron (17–28; S Mexico, Central America), Nerophila (1; tropical West Africa), Cailliella (1; tropical West Africa), Dionychastrum (1; Uluguru Mountains in Tanzania).

∎ TIBOUCHINA AND ALLIES

45. Andesanthus P.J.F.Guim. & Michelang. Trees or small trees 5–20 m at maturity or shrubs 1–3 m tall, with trichomes flattened scales of varying sizes and shapes, scabrous, trichomes bulla-based or with pustulate base. 9 spp., two in mountains of S Central America in Costa Rica and Panamá and seven from W Venezuela in Mérida to S & C Peru in Pasco region, mainly in Colombia, in cool montane forests, growing along road cuts or forest margins.

46. Brachyotum Triana. Shrubs or small trees; flowers pendulous; petals free but connivent and imbricate in a campanulate tube, often dark purple; stamens isomorphic. 55 spp. at high elevations (1,500 – 4,700 m) in the Andes from Colombia to Bolivia, one up to Argentina, either in paramo or puna vegetation, or in the surrounding cloud forests; largest Neotropical genus in Melastomataceae absent in Brazil.

47. Bucquetia DC. Three spp. restricted to the N Andes, and found in areas of páramo and subpáramo of Venezuela (Táchira), N Colombia and S Ecuador at 2,300–4,300 m.

48. Centradenia G. Don. Erect shrubs, subshrubs, or suffrutescent, (0.06–)0.2–2 (–3) m tall, perennial; puberulous, hispid or strigose on stems and leaves with smooth, unbranched, multicellular trichomes; glandular trichomes with multicellular stalks terminated by a globose cell. 4 spp., from Hidalgo, Mexico, through Central America to Panamá, with C. paradoxa (Kraenzl.) Almeda known from outlying stations in the Choco lowlands on the Pacific coast of Colombia.

Their range parallels that of the monotypic genus Schwackaea, and to a lesser extent that of Heterocentron, which is centered in Mexico and northern Central America; these distribution patterns are noteworthy because almost all other neotropical genera in the family with species in Mexico and Central America have centers of diversity in South America.

49. Chaetogastra DC. Shrubs, subshrubs, herbs or treelets covered with spreadingpubescent trichomes or villose to setulose, sometimes with xylopodium. 118 spp., from Mexico, Central America and the Caribbean through the Andes of Venezuela, Colombia, Ecuador, Peru, Bolivia and Argentina to E Brazil (41, 34 endemics), mostly in cloud forests; some herbaceous species of wide distribution, such as C. gracilis (Bonpl.) DC., occupy open areas above 100 m from Colombia and Venezuela to Bolivia, N Argentina, and C Brazil.

50. Chaetolepis (DC.) Miq. Highly branched shrubs or suffrutescent herbs, with leaves strigose or hispid and rigid with an ericoid appearance, small flowers with 4 petals yellow or purple and yellow stamen connective not or slightly prolonged at the base, less then 1 mm long, without appendages. 9 spp., 8 in paramos of Colombia and Venezuela, C. anisandra Naudin with population also in the Guiana Shield, and one endemic to the highest elevations in Costa Rica.

51. Monochaetum (DC.) Naudin. Suffrutescent herbs or shrubs, 0.5–2 m tall, villous or pubescent appressed or laxly; trichomes simple, dendritic or glandular. 60 spp., 19 in Mexico from Sinaloa to Central America and 41 species distributed in South America from Andes to Peru and the Venezuelan Coastal Cordillera, and are also present in the Guiana Shield, usually at high elevations from 900 up to 3,200 m; most South American species are found in the paramos and cloud forests of Venezuela and Colombia, with a few species extending as far south as central Peru and M. bonplandii (Kunth) Naudin widely distributed species in the Guiana Shield – and the single species in Brazil, only in Mount Neblina.

They occur in cloud forests, pine or pine-oak forests, moist thickets, volcanic crater margins, pastures, disturbed sites, trail margins, shaded roadbanks, boggy areas and rocky meadows; most Central American species are found in cloud forests in Costa Rica and Panamá, and a second, smaller group in the mountains of southern Mexico.

52. Pilocosta Almeida & Whiffin. Subshrubs, erect or decumbent, 50 cm tall or prostrate matting or trailing herbs 30–50 cm long, pubescent to strigose in all structures with trichomes simple, smooth or finely roughened. 5 spp., from Costa Rica south Panamá, P. nana (Standl.) Almeda & Whiffin up to Colombia and Ecuador, along rivers or streams, and along seepage areas or well-drained, more open areas along roadsides, exposed gravelly banks, and disturbed, often eroded slopes.

53. Pleroma D. Don. Shrubs, subshrubs erect, rarely prostrate or trees covered with trichomes; leaves opposite or rarely verticillate, blades of a pair essentially equal in size, margin entire, papery to coriaceous, variously pubescent, glutinous or nearly glabrous; inflorescence terminal. 173 spp., one only in Ecuador and Peru, another endemic to Bolivia, and remaining 171 in E Brazil in Atlantic Forest and savannas of center Brazil (cerrado), and rarely in dry seasonal scrubland of NE Brazil (caatinga), five of then outside Brazil, up to Venezuela, Colombia, Peru, Guianas, Bolívia and Central America; forests and forest margins, river banks, high altitud grasslands (campos de altitude), rocky outcrops, and Atlantic sandy coastal shrublands (restingas), from sea level to 2,650 m. P. cleistoflora (Ule) P.J.F. Guim. is cleistogamous prostrate herb. 4 spp. (all ex-Svitramia) from Minas Gerais state are rare plants in Brazil, by Plantas Raras do Brasil’s book.

54. Schwackaea Cogn. Erect annual herb, 15–40(–50) cm tall; trichotomously branched, the branchlets quadrangular, setulose on the nodes and sometimes along the angles, trichomes short and simple except for reproductive structures where they are setose and in the hypanthium where they are rarely small glandular, otherwise glabrous. Only one sp., S. cupheoides (Benth.) Cogn., from Mexico throughout Central America, chiefly on the Pacific slope, to N Colombia and on Cocos Island and grows in natural or man-made savannas from sea level to 2,000 m altitude.

55. Tibouchina Aubl. Shrubs, subshrubs or treelets, strigose in all structures with trichomes that are compressed into flattened scales of varying sizes and shapes. 39 spp. from Belize, Honduras, Nicaragua, Costa Rica and Panamá to Amazon rainforest in Bolivia and Brazil, including Venezuela and all the Guianas, 37 in South America, 23 in Brazil (15 endemics), mainly in N and W regions, with a southern boundary in savannas of C Brazil (cerrado) at Minas Gerais and Mato Grosso do Sul states; it inhabits lowland savannas (up to 1,200 m), open areas, forest edges, stream margins and rocky slopes and outcrops.

∎ CORE NEW WORLD MELASTOMATOIDEAE

56. Castratella Naudin. Two spp. restricted to paramos in W Venezuela and E Colombia.

57. Desmoscelis Naudin. Two spp., one endemic to Bolivia and D. villosa (Aubl.) Naudin over Paraguay, Brazil up to N South America, Venezuela and Guianas.

58. Loricalepis Brade. Small shrubs with tetramerous flowers and white petals. Two spp., L. duckei Brade only from white sand vegetation along the upper Rio Negro basin, in the state of Amazonas, Brazil, and in the nearby Río Guainía basin, in the department of Guainía, Colombia; and another endemic to Atlantic Forest domain in Bahia state, where it is known from a single locality in the Itacaré municipality.

59. Mallophyton Wurdack. Only one sp., M. chimantense Wurdack, endemic to Pantepui Life Zone, in Chimantá-tepui Venezuela, at 2,000 – 2,500 m elevation range.

60. Poteranthera Bong. 6 spp., two only in Bolivia, three only in Brazil, and P. pusilla Bong. over northern South America from Colombia, Venezuela, Bolivia and Brazil.

61. Pterogastra Naudin. Three spp., one in Peru, P. divaricata (Bonpl.) Naudin in Colombia, Venezuela, Guyana, Ecuador, Peru and Amazonas state in N Brazil, at altitudes between sealevel and 2,600 m in natural or disturbed grasslands; and P. minor Naudin confined to natural savannas at low altitudes in Venezuela and Colombia.

62. Pterolepis (DC.) Miq. Herbs and small shrubs (sometimes with xylopodium) with pink, purple, or white (3-)4-5-merous flowers, stamens with basal-ventrally prolonged connectives, and cochleate seeds. 16 spp., one restricted of Central America and S Mexico, and 15 remaining in Brazil, 11 endemics and four remaining reaching into the over tropical South America, some up to Mexico and Caribbean; two species, from Amazonas and Bahia state, are rare plants in Brazil, by Plantas Raras do Brasil’s book.

Pterolepis haplostemona Almeda & A.B. Martins and Microlicia macedoi L.B. Sm. & Wurdack appear to be the only known Melastomataceae that are endemic to serpentine substrates in Brazil.

2.17 MELASTOMATOIDEAE ▸ TRIBE MARCETIEAE (19/139) - all genera occur in South America.

63. Acanthella Hook.f. Erect, perennial, subshrubs to treelets, 0.5–3 m tal; stems not winged; leaves shortly petiolate, congested, blade glabrous. Two spp. endemics to the Guiana Shield in Colombia, Venezuela and Amazonas state in Brazil (only A. sprucei Benth. & Hook. f.), in fissures of granitic and sandstone outcrops, from 50 to 1,900 m.

64. Aciotis D. Don. Erect or prostate, annual or perennial, mostly herbs to subshrubs, 0.2–2 m tall. 14 spp., tropical America, all in Brazil, two endemics (one, from Pará state, is a rare plant in Brazil, by Plantas Raras do Brasil’s book); commonly found in wet areas, also in secondary forest and less often in drier habitats or sandy soils, from sea level up to 1,700 m.

65. Acisanthera P. Browne. Erect, perennial, herbs to subshrubs, 0.1–0.8 m tal; stems carinate to conspicuously winged. 7 spp., tropical America from Mexico to Cono Sur and Caribbean and Brazil (5, two endemics), usually on the margins of lakes, creeks, and rivers and in flooded areas; A. quadrata Pers. is widely distributed and occur in Caribbean.

66. Appendicularia DC. Erect, annual, herbs to shrubs, 0.7–3 m tal; stems not winged. 4 spp., SE Venezuela, Guianas, and Brazil (3, none endemics), in the state of Amapá; lowland savannas and rocky, open areas, sometimes in iron-rich mountains from 50 to 800 m.

67. Brasilianthus Almeda & Michelangelli. Delicate, wiry annual herbs 4–38 cm tall, mostly sparingly and openly branched distally; leaves opposite, isomorphic in size and shape in each pair; basal leaves early deciduous and typically not present on flowering or fruiting plants; cauline leaves membranaceous to somewhat fleshy when fresh; flowers 4-merous and haplostemonous. Only one sp., B. carajaensis Almeda & Michelangelli, restricted of Carajás Range where it is restricted to rocky-ferriginous grasslands (cangas) that form island-like lenses nestled in the Amazon rainforest of SE Pará state, Brazil.

68. Comolia DC. (exc. Acisanthera p.p., Fritzschia p.p.) Shrubs to subshrubs; stems and thunk quadrangular in cross section; leaves of many formats; flowers solitary, in dichasia or in panicles, axillary or terminal; flowers tetramerous, corolla purple, pink or white. 11 spp., lowland savannas of South America, from Colombia, SE Venezuela, Trinidad, and the Guianas to N and NE Brazil (8, three endemics, one of them is a a rare plant in Brazil, by Plantas Raras do Brasil’s book), from sea level to 1,100 m.

69. Comoliopsis Wurdack. Erect or prostate, perennial, woody shrubs to subshrubs, 0.1–3 m tall; stems not winged. Three spp., restricted to the mountain complexes of the Guiana Shield in southern Venezuela, C. neblinae Wurdack up to N Brazil, including Mount Roraima (at Roraima state in Brazil), Mount Neblina (at Amazonas state in Brazil), and Duida, at 2,000–2,400 m.

70. Dicrananthera C. Presl. Erect, annual, herbs, 0.1–0.4 m tall. Stems minutely winged. Only one sp., D. hedyotidea C. Presl., native from Guyana, Venezuela to NE Brazil, in swampy savannas or forest edges at 150–700 m.

71. Ernestia DC. (exc. Appendicularia p.p., Pseudernestia) Erect, commonly perennial, shrubs to subshrubs, 0.5–2 m tal; stems not winged. 11 spp. from Venezuela, Colombia, Peru, Ecuador, and Brazil in Amazonas state (6, none endemics), in lowland to highland habitats, often on rock outcrops and disturbed areas along lowland forest edges, at an altitudinal range of 300–1,500 m.

72. Fritzschia Cham. Prostate and decumbent or erect, perennial, herbs, shrubs to subshrubs 0.2–3 m tal; stems not winged. 9 spp., restricted mostly to the Espinhaço Range of Minas Gerais state in SE Brazil, except one which extends into Goiás and Distrito Federal, in central Brazil at elevations from 700 to 1,800 m; rocky grasslands, usually associated with sandy, humid to well-drained soils or stream edges.

73. Leiostegia Benth. Erect, perennial, shrubs to subshrubs, 0.5–2 m tal; stems not winged. Only one sp., L. vernicosa Benth., SE Venezuela, Guyana, Suriname, and Brazil, in Amazonas and Pará states, in open savannas at an altitudinal range of 100–500 m.

74. Macairea DC. Shrubs to subshrubs, often trees or small herbs; leaves peciolate; inflorescence panicles, thyrsoid, terminal; bracts leafy; flowers 4-merous, corolla pink to magenta, rare white or purple. 22 spp. from tropical South America, 17 endemics to the Guiana Shield (some strictly adapted to climate and soil conditions), two from the Guiana Shield to Ecuador, Peru, M. radula (Bonpl.) DC. restricted to the Brazilian Shield up to Bolivia (being commonly found in the savannas of C Brazil (cerrado), growing in gallery forests), and two in over Amazon lowlands, inc. M. thyrsiflora DC., the most widely distributed species. 11 spp. in Brazil, only one endemic.

75. Marcetia DC. Subshrubs to shrubs, erect, caespitose or prostrate, rarely herbs; Stems quadrangular in cross section; flowers 4-merous, solytary, corolla lilac, pink, purple, red, magenta, white or yellow. 33 spp., confined to Brazil except by and M. taxifolia (A. St.-Hil.) DC., which has a wide and disjunct distribution between E Brazil and NW South America (Colombia, Venezuela and Guyana); its main centre of diversity in Bahia, where 25 of its species occur; 14 spp. are rare plants in Brazil, by Plantas Raras do Brasil’s book, all in Bahia state except one in Minas Gerais.

76. Nepsera Naudin. Erect, perennial, usually herbs or shrubs, 0.5–2 m tal; stems not winged. Only one sp., N. aquatica (Aubl.) Naudin, widely distributed in Central America, the Antilles, and South America from the coast of Ecuador and Colombia through the inter-Andean valleys east into Brazil, Venezuela, and the Guianas; wet areas, near streams or swamps in forests, from sea level to 500 m.

77. Noterophila Mart. Erect, annual, herbs or rarely subshrubs, 0.3–0.5 m tal; stems winged and basally swollen. 6 spp., from Cuba, Trinidad, Belize, Costa Rica, Honduras, Nicaragua, Panamá, Bolivia, Colombia, Guianas, Suriname, Venezuela and Brazil (all species, mainly in Roraima and Pará states, one endemic to dry savannas in center country); humid, open areas including sandy savannas, granitic outcrops, swamps, pastures, mangroves, lake margins, and Atlantic sandy coastal shrublands (restingas), and sometimes associated with pine-oak forests, at elevations from 0 to 1,300 m alt.

78. Pseudoernestia (Cogn.) Krasser. (off Ernestia) Erect, perennial, shrubs to subshrubs, 0.5–0.8 m tal; stems not winged. Two spp., one from Colombia, Peru and Venezuela and P. glandulosa (Gleason) M.J.R.Rocha & P.J.F.Guim. in Guyana, Suriname, and Brazil (Amapá state); lowlands along streams and forests, from 200 to 400 m alt.

79. Rostranthera M. J. R. Rocha & P. J. F. Guim. Erect, perennial, herbs to shrubs, 0.2–1 m tal; stems winged. Only one sp., R. tetraptera (Cogn.) M. J. R. Rocha & P. J. F. Guim., Suriname and Amazonas, Pará, and Roraima state in Brazilian Amazon, also expected for French Guiana, in lowland rainforests and savanas, sometimes on rocky sandstone.

80. Sandemania Gleason. Erect, perennial, shrubs to subshrubs, 1–2 m tal; stems not winged. Only one sp., S. hoehnei (Cogn.) Wurdack; throughout the Amazon rainforest of Venezuela, Peru, Bolivia, and Brazil, in white sand savannas.

81. Siphanthera Pohl. Herbs to subshrubs; leaves sessile or subsessile, membranaceous; flowers 4-merous, solitary or in globose inflorescence, sometimes also panicles; corolla lavender, purple or white. 15 spp. from Colombia, Guyana, Suriname, Peru, Brazil (12, 5 endemics), and Bolivia, primarily in sandy, grassy, or rocky soil, mostly in moist areas at elevations ranging from near sea level to 2,700 m.

LINEAGE 5 of 5: OLIGOMYRTALES

ALZATEACEAE

§ FULLY CHLOROTROPHIC

Genera/species 1/1 Distribution lower Andean slopes of Peru and Bolivia, damp forests of Upper Amazonas and cloud forests of Costa Rica and Panamá. Habit bisexual, evergreen trees or shrubs (sometimes semi-epiphytic). Young stems and branches at first quadrangular in cross-section, later terete.

Relationships suggested by various authors since 1794 encompass eight families in five orders. Systematic placement has included: Santalaceae, Celastraceae, Lythraceae, Crypteroniaceae, Rhamnaceae, Melastomataceae and Flacourtiaceae. Alzateaceae is most probably a member of the order Myrtales with characters such as internal phloem, vestured pits, flavonols in the leaves and ellagic acid. Major differences from all other families of Myrtales in morphology and embryology are sufficient for Alzateaceae to stand as a separate monotypic family. DNA studies recently confirm that Rhynchocalycaceae is the most closely related family and also monotypic.

Key differences from similar families the families below differ from Alzateaceae in the following characters:

Santalaceae: No ellagic acid produced or internal phloem.

Celastraceae: Sepals and petals are mostly imbricate, has axile placentation, no internal phloem with seeds which are often arillate with abundant endosperm.

Rhamnaceae: No internal phloem, ellagic acid, vestured pitting of vessel elements nor wings on seeds plus it has drupaceous fruits (not dry capsules).

Flacourtiaceae: Leaves usually alternate.

Melastomataceae: Conspicuous venation, showy flowers and apically dehiscent anthers.

Lythraceae: No thick fleshy perianth, petals present and no distinctly bisporic Allium embryo sac.

Sonneratiaceae: Has pneumatophores, showy petals and is many seeded.

Punicaceae: Showy petals and many seeded fruit.

Myrtaceae: Superior ovary.

SYSTEMATIC a single species.

1. Alzatea Ruiz & Pav. Variable habit from small trees to slender high-climbing hemi-epiphytes with a tendency to a multi-trunked growth form, stilt roots and sometimes developing a strangler habit; leaves Clusia-like, thick and coriaceous, oval with rounded apex and base with very short petiole; secondary veins immersed or slightly prominulous below. Only one sp., A. verticillata Ruiz & Pav. which occur along the lower slopes of the Andes in Peru and Bolivia, in humid forests of the Upper Amazon rainforest of Colombia and Ecuador, and also in the cloud forests at elevations from 900 to 2,200 (-3,000) meters in Costa Rica and Panamá; two subespecies, only minimally distinct and some partially intermediate representatives do occur near the border of Ecuador and Peru:

§ subsp. amplifolia S.A. Graham - larger more oval, sessile or subsessile leaves and is distributed through Costa Rica and Panamá.

§ subsp. verticillata Ruiz & Pav. - smaller leaves with petioles and is distributed through the eastern escarpment of the Andean mountains in South America.

39. CROSSOSOMATALES

FAMILIES ABSENTS IN SOUTH AMERICA: APHLOIACEAE (1/1), CROSSOSOMATACEAE (4/7), GEISSOLOMATACEAE (1/1), GUAMATELACEAE (1/1), STACHYURACEAE (1/10) AND STRASBURGERIACEAE (2/2).

STAPHYLEACEAE

§ FULLY CHLOROTROPHIC

Genera/species 3/43Distribution temperate W and E North America, S Mexico, the West Indies, NW South America southwards to Bolivia, SE Europe, Caucasus, N Türkiye, S India, Sri Lanka, Himalaya, China, Korean Peninsula, Japan, SE Asia, Malesia, New Guinea. Habit usually bisexual (sometimes monoecious, polygamomonoecious or dioecious), evergreen or deciduous trees or shrubs, sometimes stoloniferous.

SYSTEMATICS outsiders Staphylea (11; Central and SE Europe, Türkiye, Caucasus, temperate Asia to E Asia, North America) and Euscaphis (1; temperate China, Korean Peninsula, Japan, N Vietnam).

1. Turpinia L. Deciduous shrubs or small trees; leaves (3–)7– 11-foliolate, with or without stipule-like glands at the insertion of the petioles. 30-40 spp., China to Japan, S India, Sri Lanka, Himalayas, SE Asia, Malesia to New Guinea, NW South America southwards to Bolivia; 7 spp. in New World, two in South America, T. occidentalis (Sw.) G.Don, ranging from Mexico to Bolivia and Brazil (very common in Acre state, also Amazonas and Rondonia, in Amazon rainforest), where know as ‘sabugueiro’, and T. megaphylla (Tul.) G.Don., restricted of Colombia and Ecuador.

40. PICRAMNIALES

A SINGLE FAMILY, PRESENT IN SOUTH AMERICA.

PICRAMNIACEAE

§ FULLY CHLOROTROPHIC

Genera/species 4/57 Distribution Florida, Central America, Caribbean, tropical South America. Habit Dioecious, usually evergreen trees or shrubs, bark contains bitter-tasting anthraquinones.

Useful tips for generic identification: fruit a berry in Picramnia, a samara in Alvaradoa; carpels only 1 fertile in Alvaradoa, 2 - 3 in Picramnia; ovules terminal and pendulous in Picramnia, basal and erect in Alvaradoa.

SYSTEMATIC all genera in this family occur in South America.

1. Aenigmanu W. Thomas. Trees, compount leaves, reddish stellate fruits. Only one sp., A. alvareziae W. W. Thomas Picramniaceae, known from two locations in Amazon region of Madre de Diós, E Peru, and one in W Acre state in northern Brazil.

2. Alvaradoa Liebm. Medium-sized trees to shrubs; inflorescences of long, slender racemes. 5 spp., A. subovata Cronquist from Argentina to Bolivia, one from U.S.A. to Central America and Caribbean, and remainig four in Caribbean (Cuba, Hispaniola and Jamaica).

3. Nothotalisia W. Thomas. Small tree or shrub, 0.5–8 m tall, dioecious; leaves alternate, imparipinnate, estipulate, the leaflets alternate or subopposite, (1–)3–8, the venation brochidodromous; inflorescence terminal, subterminal, or axillary, a raceme of glomerules, each glomerule a congested cyme of 1–12 flowers. 3 spp., Panamá, Colombia, Peru (one endemic), Bolivia, N. peruviana (Standl.) W.W.Thomas reaching to Acre and Amazonas states in N Brazil, all in E side of Andes, absent in Venezuela and Ecuador; populations of Colombia and Panama are odd by Amazonian in flowers and fruits.

4. Picramnia Swartz. Small trees or shrubs; inflorescences thyrses or racemes; estipulate, compound leaves with alternate leaflets, dioecy, racemose or paniculate inflorescences bearing minute flowers, antepetalous stamens, and berries with persistent, sessile stigma lobes. 48 spp., Florida, Mexico, and Caribbean to S Brazil (21, 11 endemics), Argentina and Paraguay; 35 in South America; practicalIy all species are found in forests (with the exception of a couple of species found in savannas of C Brazil (cerrado) and rocky grasslands (campos rupestres) of Brazil); P. ferrea Pirani & W.W. Thomas from Carajas Range, Pará state, and P. grandifolia Engl., near Rio de Janeiro municipality, are rare plants in Brazil, by Plantas Raras do Brasil’s book.

Gumillea has been placed as synonym to Picramnia. This is a questionable decision. Gumillea auriculata Ruiz & Pav. is known only from a single specimen collected in Peru at the end of the 18^th century (information from APG II site on internet). Stem in P. magnifolia J.F. Macbr. usually hollow and inhabited by ants of prey.

41. SAPINDALES

FAMILIES ABSENTS IN SOUTH AMERICA: BIEBERSTEINIACEAE (1/4), KIRKIACEAE (1/6) AND NITRARIACEAE (4/16).

LINEAGE 1 of 3: BURSERACEAE/ANACARDIACEAE and OUTSIDERS

BURSERACEAE

§ FULLY CHLOROTROPHIC

Genera/species 18/c. 610 Distribution tropical and subtropical regions in the Northern and Southern Hemispheres northwards to California, Himalaya and E China, and southwards to Uruguay, South Africa and northern Australia. Habit usually dioecious or polygamomonoecious (sometimes bisexual), evergreen or deciduous trees or shrubs (sometimes epiphytic). Branches usually spinose. Sometimes pachycaul. Stilt roots or plank buttresses often present. Resin often fragrant (often like almond).

Bursera dominates many of Mexico's dry forests in these terms. On dehiscence of the fruit, the valves fall away to reveal a basifixed stone (usually) partially enveloped in a fleshy pulp that is red, orange, or yellow. One of its two subgenera has a characteristic peeling bark that is often red or yellow. Protium is highly important in relative density, relative diversity or both in many of northern South America's moist to wet forests, particularly in central Amazon rainforest; it usually has a relatively smooth, grayish bark, the lateral petiolules almost always have a distal pulvinulus, and on dehiscence of the fruit the valves fall away and the 1-5 stones, each enveloped in a sweet, white (rarely red) pulp, are suspended from the apex of the fruit by an inverted V-shaped structure.

Key differences from similar families Burseraceae vs. Anacardiaceae: locules with two epitropous ovules (vs. one apotropous ovule); resin not causing contact dermatitis (vs. sometimes); corolla aestivation usually induplicate-valvate (vs. imbricate or less often valvate).

SYSTEMATIC all South American Burseraceae are Burseroideae (Beiselioideae is a monotypic endemic to Mexico); among Burseroideae, Garugeae (2/24, semiarid regions in tropical Africa, Madagascar, Socotra, Himalayas, SE Asia, Malesia to Melanesia and tropical Australia) does not occur in South America.

1. BURSEROIDEAE ▸ TRIBE PROTIEAE (1/c.180) - a single pantropical genus, foresty.

1. Protium Burm.f. Small to large trees, rarely shrubs, often buttressed. 152 spp., 143 in S Mexico and the Neotropics, 126 in South America, 83 in Brazil, 16 endemics, 4 in tropical China, India, the Malesian region, and two in Madagascar and Mauritius; it’s highly important in relative density, relative diversity or both in many of northern South Americas moist to wet forests, particularly in central Amazon rainforest. P. amplum Cuatrec. from Bajo Calima region, Choco, W Colombia, has the largest leaf among Burseraceae; three spp. from Rio de Janeiro, Goiás and Paraná states are rare plants in Brazil, by Plantas Raras do Brasil’s book.

2. BURSEROIDEAE ▸ TRIBE BURSEREAE (3/302) - outsider Aucoumea (1, Central Africa).

2. Bursera Jacq. ex L. 112 spp. from SW U.S.A., Mexico (96, 83 endemics), Central America, Caribbean, and 7 in South America, in Colombia, Ecuador, N Peru, N Venezuela, two of them up to extreme N Brazil, in Roraima and Amazonas states, and B. pereira DC. Daly disjunct, endemic to open savannah in Goiás state, center Brazil, known only three populations, and southermost point of range of genus. A highly incert taxon (B. tonkinensis Engl.) occur in Vietnan.

3. Commiphora Jacq. Shrubs or trees, sometimes caudiciform or rupicolous, (polygamo-)dioecious; bark close or more often papery and peeling. c. 185 spp., Africa, Madagascar, Arabia, peninsular India, Sri Lanka, Vietnam, Pakistan, Iran, and C. leptophloeos (Mart.) J. B. Gillett from thorn-scrub of NE Brazil but also occurs in dry formations in a broad are thet includes S Pará, Mato Grosso, Minas Gerais, Goiás, the upper rio Orinoco basin in Venezuela, and the Department of Santa Cruz in Bolivia and Paraguay; this species has a noticeably thickened trunk and branches and attractively peeling bark; it is a common species that we saw at many of our stops; it grows in sandy and rocky soils as well as in sandstone and limestone outcroppings.

3. BURSEROIDEAE ▸ TRIBE CANARIEAE (7/264) - outisiders Triomma (1; Bihar in India, W Malesia), Ambilobea (1; N Madagascar), Canarium (c 120; tropical Africa, Madagascar, islands in the Indian Ocean, tropical Asia to S China and New Guinea, N and E Australia, Fiji, Micronesia, Tonga, Samoa), Santiria (22–24; W Africa, Malesia) and Rosselia (1; SE New Guinea).

4. Dacryodes Vahl. Evergreen dioecious trees; trunk shallowly fluted; crown dense, much branched; leaves imparipinnate; leaflets 5–8-jugate, entire; inflorescences of axillary or terminal, elongated panicles; flowers unisexual; calyx 3-lobed, rotate or broadly campanulate; fruit an ovoid or ellipsoid drupe; endocarp thin and cartilaginous; seed large; cotyledons very much thickened and deeply folded or conduplicate, thus appearing palmately lobed. 103 spp., 32 in Asia, 18 in Africa, and 53 in Neotropics; 49 spp. in South America, 17 in Brazil, 3 endemics; usually lowland but some montane; D. edilsonii D.C.Daly from Acre state is a rare plant in Brazil, by Plantas Raras do Brasil’s book.

5. Trattinnickia Willd. Dioecious trees up to 40 m tall, rarely shrubby. 19 spp., mainly in Amazon rainforests and Guianas, 10-13 in overall Brazil, three endemic in Atlantic Forest (Bahia, Minas Gerais and Rio de Janeiro one each), T. aspera (Standl.) Swart in the Chocó (including S Panamá), and two in the Andes; the genus occurs primarily in lowland moist to pluvial forests but ranges from near sea level to ca. 1,000 m elevation; T. ferruginea Kuhlm. from Minas Gerais state is a rare plant in Brazil, by Plantas Raras do Brasil’s book.

ANACARDIACEAE

§ FULLY CHLOROTROPHIC

Genera/species 79/730–750 Distribution mainly tropical and subtropical regions in the Northern and Southern Hemispheres; some species in warm-temperate areas northwards to southern Canada, Central and E Europe, and NE China. Habit monoecious, andromonoecious, polygamomonoecious, dioecious, or gynodioecious (sometimes bisexual), evergreen or deciduous trees or shrubs (sometimes with spines; rarely lianas or perennial herbs to suffrutices).

Anacardiaceae (the cashew or sumac family) are a family of flowering plants bearing fruits that are drupes and in some cases producing urushiol, an irritant. There are 81 (see list above) genera of Anacardiaceae and ca. 800 species worldwide, of which 31 are native to the Neotropics. Anacardium occidentale L., the cashew, is a tree of this family; originally native to NE Brazil, it is now widely grown in tropical climates for its cashew seeds and cashew apples. Toxicodendron succedaneum (L.) Kuntze has naturalized in Brazil and possibly elsewhere. Several non-native Anacardiaceae are cultivated in the Neotropics for their edible fruits: Bouea macrophylla Griff., Harpephyllum caffrum Bernh. ex Krauss, Mangifera indica L., Schinus terebinthifolia Raddi, Sclerocarya birrea Hochst. subspecies caffra (Sond.) Kokwaro, and Spondias dulcis G.Forst. Spondias tuberosa Arruda, commonly known as Umbu, Imbu, or Brazil plum, is native to NE Brazil; it bears fruit once a year and can produce up to 300 kilos of fruit in a single tree; this tree have a robust root system, and is an important resource for one of the poorest and driest regions of Brazil, where local agriculture is based on corn, beans, sheep, and goat.

A geoxylic suffrutex habit occurs in Lannea from Zambezian region and savannas of C Brazil (cerrado, Anacardium); water storage roots occurs in some spp., eg. Spondias tuberosa endemic to dry seasonal scrubland of NE Brazil (caatinga).

Key differences from similar families Anacardiaceae can be separated from Burseraceae by the latter virtually always having opposite leaflets, whereas Anacardiaceae leaflets are usually alternate or subopposite.

SYSTEMATIC two subfamilies, both in South America, and some unplaced genera at family subdivisions: Euleria (1; Cuba), Haematostaphis (1; tropical Africa to Nigeria), Holigarna (8; India, SE Asia), Koordersiodendron (1; Borneo, Philippines, Sulawesi, Maluku, New Guinea), Pseudospondias (2; tropical W and C Africa).

1. SUBFAMILY ANACARDIOIDEAE (73/810) ‣ three tribes, all in South America, and some outsider unplaced genera.

ANACARDIOIDEAE ▸ UNPLACED ANACARDIOIDEAE – outsiders Androtium (1; W Malesia), Blepharocarya (2; Arnhem Land in Northern Territory, Queensland), Campylopetalum (1; Thailand), Drimycarpus (3–4; India to Borneo), Euroschinus (9; Malesia to New Guinea, Queensland, New South Wales, New Caledonia), Melanochyla (c 30; S Thailand, Malesia), Nothopegia (10; India, Sri Lanka), Parishia (5; Burma, Thailand, W Malesia), Pseudosmodingium (5; Mexico), Rhodosphaera (1; Queensland, New South Wales), Swintonia (12; Andaman Islands, SE Asia, W and Central Malesia), Trichoscypha (32; tropical and S Africa)

1. Apterokarpos Rizzini. Shrubs or trees, dioecious. Only one sp., A. gardneri (Engl.) Rizzini, endemic to dry seasonal scrubland of NE Brazil (caatinga).

2. Cardenasiodendron F.A.Barkley. Trees dioecious, with contact dermatitiscausing exudate. Only one sp., C. brachypterum (Loes.) F.A. Barkley, endemic to Bolivia.

3. Haplorhus Engl. Trees dioecious; leaves evergreen, alternate, simple, sessile to very short petiolate, linear to lanceolate. Only one sp., H. peruviana Engl., endemic to the dry valleys of Peru to N Chile.

4. Myracrodruon Allem. Trees dioecious up to 30 m tall, with contact dermatitis-causing exudate. Two spp., in sub-Amazonian Brazil (both spp., none endemics), Bolivia, Paraguay, and northern Argentina.

5. Ochoterenaea F.A.Barkley. Trees, dioecious with milky exudate. Only one sp., O. colombiana F.A. Barkley, in Panamá, Venezuela and Colombia south to Ecuador and Bolivia.

1.1 ANACARDIOIDEAE ▸ TRIBE TAPIRIRINAE (10/85-90) - outsiders Choerospondias (1; NE India to N Thailand, Vietnam, SE China (inc. Taiwan) and Japan), Pleiogynium (2; Indochina and Malesia to Queensland and islands in the Pacific), Dracontomelon (8; SE Asia, Malesia to Fiji), Cyrtocarpa (3, Mexico), Poupartia (c 10; Madagascar, Mascarene Islands), Harpephyllum (1; S Africa), Lannea (c 40; tropical Africa, Madagascar, Socotra, tropical Asia), Operculicarya (8; Madagascar, the Comoros, Aldabra).

6. Antrocaryon Pierre. Tall, dioecious to polygamous trees; leaves alternate, imparipinnate with 5–9 pairs of entire leaflets; inflorescence lax, subterminal-axillary panicles; fruit a subglobose, somewhat horizontally flattened, weakly 5-lobed drupe with thin fleshy mesocarp and woody endocarp forming a nut with 5 elliptic; seeds 3 or 4 per nut, oblong, flattened, curved; testa thin. Three spp., two in tropical Africa and A. amazonicum (Ducke) B.L.Burtt & A.W.Hill in Amazon rainforest of NW Brazil, SE Colombia and E Peru.

7. Tapirira Aubl. Trees up to 30 m tall, polygamodioecious; leaves evergreen, alternate, imparipinnate to paripinnate, petiolate. 10 spp. from S Mexico to SE Brazil (3, none endemics), Bolivia and Paraguay; 7 in South America.

8. Tumultinervia J. D. Mitch. & Daly. Small to medium-sized polygamodioecious. Only one sp., T. caatingae (Mitchell & Daly) J. D. Mitch. & Daly in dry thorn-scrub of Bahia, Tocantins, Goiás and Minas Gerais states, Brazil.

9. Uniostium J. D. Mitch. & Daly. Small to medium-sized polygamodioecious trees. Only one sp., U. velutinifolia (Cowan) J. D. Mitch. & Daly in NE Colombia to NE Roraima state in Brazil, including Guyana and Venezuela.

1.2 ANACARDIOIDEAE ▸ TRIBE BUCHANANIEAE (3/43-50) - oustiders Buchanania (25–30; tropical Asia to islands in W Pacific) and Pentaspadon (5; SE Asia, Malesia to New Guinea, Solomon Islands).

10. Campnosperma Thwaites. Trees, polygamodioecious with Terminalia-branching, often trunk buttressed or with stilt roots, and with contact dermatitis-causing exudate. 13 spp., 11 in Madagascar, Seychelles and tropical Asia, and two in New World, C. gummiferum (Benth.) Marchand from Honduras to Ecuador, and C. panamense Standl. in Brazil, Colombia and Venezuela.

1.3 ANACARDIOIDEAE ▸ TRIBE ANACARDIEAE (39/555–585) – three clades, all in South America.

∎ CLADE 1 ‣ outsiders Faguetia (1; Madagascar), Semecarpus (70–75; tropical Asia to Australia, New Caledonia and Fiji), Fegimanra (3; tropical W and C Africa), Gluta (c 30; Madagascar, tropical Asia), Bouea (3; SE Asia, W Malesia), Mangifera (c 70; tropical Asia to Solomon Islands).

11. Anacardium L. Shrubs to tall trees up to 30 m tall, andromonoecious subshrubs, with contact dermatitis-causing exudate, sometimes with xylopodium. 12 spp., Anacardium fruticosum J.Mitch. & S.A.Mori endemic to Guyana, A. corymbosum Barb.Rodr., A. microsepalum Loes., A. nanum A.St.-Hil. endemics to Brazil; A. amapaense J.D.Mitch. in Brazil and French Guiana; A. negrense Pires & Fróes, A. parvifolium Ducke, A. giganteum Hancock ex Engl., A. occidentale L., and A. spruceanum Benth. ex Engl. in over northern South America (all in Brazil) sometimes up to C Brazil and Trinidad; A. humile A.St.-Hil. in savanas from Brazil, Bolivia and Paraguay; and A. excelsum (Bertero & Balb. ex Kunth) Skeels in Cuba, Honduras to Ecuador and Venezuela, absent in Brazil.

Anacardium humile differs from the other species of Anacardium because it presents a subshrub habit and narrower leaves.

∎ CLADE 2 ‣ outsiders Bonetiella (1; Mexico), Comocladia (16–20; Mexico, Central America, Caribbean), Cotinus (7; Mediterranean to China; NW Yunnan; SE U.S.A.), Rhus (35–40; temperate and subtropical regions on both hemispheres), Searsia (110–115; Mediterranean, tropical and subtropical Africa, Socotra, the Middle East, Arabian Peninsula, India, Himalayas, Burma, SW China), Baronia (1; Madagascar), Malosma (1; SW California, Baja California), Laurophyllus (1; Cape), Pachycormus (1; NW Mexico), Pistacia (12; Mediterranean, Asia to Malesia, S U.S.A. to Central America), Actinocheita (1; Mexico).

12. Astronium Jacq. Trees up to 40 m tall, dioecious, with clear contact dermatitis-causing exudate. 11 spp., in Mexico south to Paraguay and northern Argentina; all spp. in South America, 8 in Brazil, 4 endemics.

13. Lithrea Hook. Trees, dioecious, with clear contact dermatitis-causing exudate, sometimes with lignotuber. Three spp. in Paraguay, Argentina, Uruguay, Chile, two up to in Brazil (both, none endemics) and Bolivia.

14. Loxopterygium Hook.f. Trees, polygamodioecious with contact dermatitiscausing exudate, clear or white and turning black with exposure to air. Three spp. with disjunct distributions from Venezuela south to Argentina, absent from Amazon rainforest, L. sagotii Hook. f. in Venezuela, Guyana, Suriname and French Guiana; L. huasango Spruce ex Engl. in SW Ecuador to NW Peru; and L. grisebachii Hieron. in Bolivia south to NW Argentina.

15. Metopium P. Br. Trees or shrubs with prominent resin ducts and contact dermatitis-causing exudate turning black with exposure to air; leaves evergreen, imparipinnate, petiolate; leaflets petiolulate, entire; mature leaflets often speckled with black spots. Three spp. in Caribbean, S Florida (US), Mexico, and N Central America, with M. brownei (Jacq.) Urb. in Caribbean islands of Colombia.

16. Mosquitoxylum Krug & Urb. Trees, polygamodioecious (androdioecious); leaves evergreen (more or less), alternate, imparipinnate; leaflets opposite or subopposite, short-petiolulate, entire. Only one sp., M. jamaicense Krug & Urb., Jamaica, and S Mexico to NW Ecuador.

17. Orthopterygium Hemsl. Shrubs or trees, dioecious with milky exudate; leaves deciduous, alternate, imparipinnate, petiolate. Only one sp., O. huaucui (A. Gray) Hemsl., endemic to W Peru.

18. Schinus L. Shrubs or trees, dioecious, rarely subshrubs, rarely with thorns, and with contact dermatitis-causing exudate. 32 spp., from Ecuador south to Patagonia but excluding Amazon rainforest, 12 in Brazil, 7 of then up to Cono Sur, three endemics, and two up to Bolivia and Peru.

19. Schinopsis Engl. Trees, dioecious or monoecious, sometimes with thorns, and with contact dermatitis-causing exudate turning black with exposure to air. 7 spp., one endemic to dry forests of N Peru, remaining in Bolivia, Paraguay to N Argentina, two up to S Brazil (none endemics), often the dominant canopy tree in Chaco forests.

20. Thyrsodium Salzm. ex Benth. Trees, dioecious with milky exudate; leaves evergreen, alternate to subopposite, imparipinnate, petiolate. 6 spp., in Colombia, Peru, Bolivia, Venezuela, the Guianas, an Amazon rainforest and E Brazil (all species, none endemics), absent from the Andes.

21. Toxicodendron Mill. Shrubs, trees, or lianas, polygamodioecious with contact dermatitis-causing, white exudate turning black with exposure to air. 22 spp., India and Nepal; Bhutan and Myanmar; and temperate E Asia to New Guinea, and six in New World, all in North America and Mexico, one up to Central America and T. striatum (Ruiz & Pav.) Kuntze up to Bolivia and Venezuela.

∎ CLADE 3 ‣ outsiders Dobinea (2; E Himalayas to S China); Loxostylis (1; South Africa), Smodingium (1; South Africa, Swaziland, Lesotho), Sorindeia (9; tropical Africa, Madagascar, Mascarene Islands), Protorhus (1; Namibia, South Africa, Swaziland, Madagascar), Abrahamia (19; Madagascar), Heeria (1; W Cape), Ozoroa (c 40; Africa, Yemen), Micronychia (6; Madagascar).

22. Mauria Kunth. Shrubs or trees, hermaphrodite, sometimes cleistogamous, or less frequently polygamodioecious with contact dermatitis-causing exudate. 16 spp., all in Andean region from Venezuela to Bolivia, M. heterophylla Kunth to El Salvador and M. thaumatophylla Loes. to extreme N Argentina.

2. SUBFAMILY SPONDIOIDEAE (4/23) ‣ oustiders Haplospondias (1; Yunnan, Burma), Pegia (2; E Himalayas, E Asia, W Malesia) and Poupartiopsis (1; E Madagascar).

23. Spondias L. Strongly protandrous trees (very rarely hemi-hepiphytic) with contact dermatitis-causing exudate. 19 spp., 11 from Mexico south to SE Brazil and Bolivia; 7 in India and Sri Lanka east to tropical China and South Pacific Islands, and one highly disjunct record in Madagascar; New World species are:

§ S. testudinis J.D. Mitch. & Daly is restricted to SW Amazon rainforest, and S. globosa J.D. Mitch. & Daly is from western Amazon rainforest.

§ S. admirabilis J.D. Mitch. & Daly and S. expeditionaria J.D. Mitch. & Daly are known from very few localities in Brazil’s Atlantic Coastal Forest, while the other two Atlantic Forest species, S. macrocarpa Engl. and especially S. venulosa (Engl.) Engl., are somewhat more broadly distributed in E Brazil.

§ S. tuberosa Arruda and S. bahiensis P.Carvalho, Van den Berg & M.Machado are endemics to dry seasonal scrubland of NE Brazil (caatinga).

§ S. mombin L. is native to moist forests through much of northern South America, although it is uncertain whether the populations in Brazil’s Atlantic Coastal Forest are native.

§ S. radlkoferi Donn. Sm. ranges from Mexico to Colombia and NW Venezuela; there is an unconfirmed report from Los Ríos in W Ecuador; S. purpurea L. is native to N Mexico and Central America and may be native to SW Ecuador.

LINEAGE 2 of 3: SAPINDACEAE

SAPINDACEAE

§ FULLY CHLOROTROPHIC

Genera/species 143/1,650-1,690 Distribution tropical, subtropical and temperate regions in the Northern and Southern Hemispheres. Habit monoecious, andromonoecious, polygamomonoecious, dioecious, androdioecious, or polygamodioecious (sometimes bisexual), evergreen or deciduous trees, shrubs or lianas (Cardiospermum consists of perennial climbing herbs with inflorescence branches modified into tendrils).

From southern U.S.A. to northern Argentina and Chile, including Caribbean. Trees, shrubs, lianas or less often herbaceous vines; 40 genera and about 800 spp. in the Neotropics; Blighia sapida Kon. is an important crop in Jamaica; it is cultivated for its edible arils. Dimocarpus longan Lour., Litchi chinensis Sonn., and Nephelium lappaceum L. are cultivated for their edible fruits in small-scale operations throughout tropical America. Acer spp., Filicium Thwaites spp., Koelreuteria spp., and Harpullia arborea Radlk. are cultivated as ornamental trees throughout the Neotropics. Sapindaceae are the source of numerous products, some of which are economically important.

In the Neotropics these include edible fruits such as keneep or genip (Melicoccus bijugatus Jacq.), wild genip (M. oliviformis Kunth) and the pitomba (Talisia esculenta Radlk.). Numerous species of Paullinia have been reported to be useful in the preparation of medicines, caffeine-rich beverages, binding and weaving material, and for fish, human and arrow poisoning. The seeds of Paullinia cupana Kunth are the source of the important Brazilian crop guaraná, a source of caffeine and flavoring of soft drinks. Almost all Sapindaceae are used around the tropics for fish poisoning.

SYSTEMATIC 19 lineages in four higher clades, subfamily Xanthoceroideae (1/1, N and NE China, Korean Peninsula) does not occur in South America; all others are present.

1. SUBFAMILY HIPPOCASTANOIDEAE (5/180–185) ‣ two tribes, Acereceae (2/c.165; temperate regions on the Northern Hemisphere, tropical mountains); among Hippocasteneae, which occur in South Ameica, outsiders are Handeliodendron (1; SW China) and Aesculus (13; the Balkan Peninsula, Himalayas to Japan and Indochina, North America, NW Mexico).

1. Billia Peyr. Trees; leaves opposite, trifoliolate; margins entire. Two spp., B. hippocastanum Peyr. from Mexico to Panamá, red to pink flowers, and B. rosea (Planch. & Linden) C. Ulloa & P. Jørg., white petals, Costa Rica to the southern extent of the range in Colombia; reddish or yellowish on the bases and/or veins as well as larger, thicker, more waxy leaves than the northern species; the two species have overlapping geographic ranges in Costa Rica and northern Panamá.

2. SUBFAMILY DODONAEOIDEAE (21/150–175) ‣ two tribes, both in South America.

2.1 DODONAEOIDEAE ▸ TRIBE DODONAEEAE (c 14/115-125) - outsiders Arfeuillea (1; SE Asia), Harpullia (25–30; tropical Asia to New Guinea, Northern Territory, Queensland, New South Wales, New Caledonia, Tonga), Majidea (4–5; tropical E Africa, Madagascar), Euphorianthus (1; Philippines and Sulawesi to Vanuatu), Eurycorymbus (1; S China inc. Taiwan), Cossinia (5; Mauritius, New Caledonia), Diplopeltis (5; W Australia, Northern Territory), Hirania (1; Somalia), Loxodiscus (1; New Caledonia)

2. Averrhoidium Baill. Trees, dioecious; leaves alternate, paripinnate; leaflets serrate or entire. 4 spp., one only in Mexico, A. paraguaiense Radlk. from Bolivia, Brazil and Parguay, and remaining two endemic to Brazil.

3. Diplokeleba N.E.Br. Trees, falsely polygamous trees; leaves paripinnate; leaflets entire or undulate. Two spp., D. herzogii Radlk. endemic to Bolivia, and D. floribunda N.E.Br. in Brazil (only Mato Grosso do Sul state), Bolivia and southern South America.

4. Dodonaea Mill. Trees or shrubs, dioecious or falsely polygamous-dioecious, with viscous glandular hairs. 67 spp., 59 endemics to Australia, 8 pantropical species, three in New World, two in Caribbean, and Dodonaea viscosa Jacq. widely distributed in tropical New World.

5. Llagunoa Ruiz & Pav. Shrubs or trees, falsely polygamous shrubs or trees; leaves alternate, simple or trifoliolate; flowers solitary or in axillary cymes. Three spp., Venezuela and Chile one endemic each, and L. nitida Ruiz & Pav. from Colombia to Bolivia.

6. Magonia Vell. Trees, falsely polygamous; leaves alternate, paripinnate; distal leaflet rudimentary; inflorescences axillary or terminal thyrses. Only one sp., M. pubescens A.St.Hil., from Brazil, Bolivia and Paraguay.

2.2 DODONAEOIDEAE ▸ TRIBE DORATOXYLEAE (8/19) - oustiders Doratoxylon (5; Madagascar, the Comoros, Mauritius), Euchorium (1; W Cuba), Filicium (3; tropical E Africa, Madagascar, one species also in India and Sri Lanka), Ganophyllum (2; Central Africa, Andaman Islands, Nicobar Islands, Vietnam, the Malay Peninsula, Sumatra, Java, Philippines, New Guinea, northern Australia, Solomon Islands), Hippobromus (1; South Africa), Zanha (3; S Africa, Madagascar), Hypelate (1; Florida, Caribbean).

7. Exothea Macfad. Trees, dioecious, falsely polygamous; leaves alternate, paripinnate. Two spp., E. paniculata (Juss.) Radlk. distributed throughout Caribbean, Mexico, Guatemala, Colombia, and Ecuador, and E. copalillo (Schltdl.) Radlk. endemic to Mexico.

3. SUBFAMILY SAPINDOIDEAE (c 110/1.615–1.640) – 14 clades, Ungnadia clade (1/1, Texas, NE Mexico), Delavaya clade (1/1, China, Vietnam), Macphersonia clade (9/c 42, Africa, Madagascar, Dhofar), Tristiropsis clade (2/6, Malesia to New Guinea, Queensland, islands in W Pacific), Koelreuterieae (5/21–22, Tropical E Africa, South Africa, Madagascar, Mauritius, SW Asia, China (inc. Taiwan), Fiji) and Schleichereae (5/10, tropical Africa, Madagascar, SW China, tropical Asia) do not occur in South America.

3.1 SAPINDOIDEAE ▸ TRIBE PANCOVIEAE (c 29/235–245) - outsiders Deinbollia (c 40; tropical and subtropical Africa, Madagascar, Mascarene Islands), Atalaya (12; tropical Africa, tropical Asia to Australia), Hornea (1; Mauritius), Thouinidium (3; Mexico, Central America, Caribbean), Lepisanthes (c 25; tropical regions in the Old World to northern Australia), Pometia (2; Sri Lanka, Andaman Islands, Nicobar Islands, Indochina, Taiwan in China, Malesia, Fiji, Samoa, Tonga), Nephelium (c 16; Assam and Yunnan to Hainan and Malesia), Dimocarpus (8; India and Sri Lanka to NE Queensland), Otonephelium (1; W India), Litchi (1; S China to W Malesia and Philippines), Lecaniodiscus (2; tropical Africa), Eriocoelum (c 10; tropical Africa), Lepidopetalum (6; Andaman Islands, Nicobar Islands, Sumatra, Philippines to New Guinea, Bismarck Archipelago, Bougainville, N Queensland), Blighia (6; tropical Africa), Cubilia (1; Borneo, Philippines, Sulawesi, Moluccas), Haplocoelopsis (1; Central and tropical E Africa), Glenniea (8; tropical Africa, Madagascar, Sri Lanka, Indochina, Malesia), Laccodiscus (4; tropical W and Central Africa), Xerospermum (c 11; Bangladesh, Indochina, W Malesia), Chytranthus (25–30; tropical W and Central Africa), Namataea (1; Cameroon), Pancovia (10–12; tropical W and C Africa), Placodiscus (c 10; tropical Africa), Pseudopancovia (1; C Africa), Radlkofera (1; tropical W Africa), Zollingeria (4; SE Asia, Borneo).

8. Porocystis Radlk. Trees, falsely polygamous; leaves alternate, paripinnate or imparipinnate; leaflets entire. Three spp., P. toulicioides Radlk. in Brazil and Guianas, P. acuminata (Radlk.) Acev.-Rodr. endemic to Brazil (both only in Amazonas state), and P. demerarae Sandwith endemic to French Guiana.

9. Pseudima Radlk. Small to large trees, falsely polygamous-dioecious; leaves alternate, pinnately compound. Only one sp., P. frutescens (Aubl.) Radlk., from Central America, Guianas, Colombia, Peru, Bolivia and Brazil.

10. Sapindus L. Trees, duodichogamous; leaves alternate, paripinnate or unifoliolate; leaflets 2–8 pairs, often falcate. 11 spp., with tropical to sub-temperate distribution, mainly in Asia, only S. saponaria L. in New World, in almost all countries of hemisphere.

11. Toulicia Aubl. Unbranched small trees, falsely polygamous-dioecious; leaves imparipinnate; leaflets usually falcate, opposite or alternate. 13 spp. from the lowlands of South America, all in Brazil, 5 endemics.

3.2 SAPINDOIDEAE ▸ BLOMIA CLADE (3/10) - oustiders Blomia (1; Mexico, Belize, Guatemala), Haplocoelum (6; tropical Africa, Madagascar).

12. Guindilia Gill. Trees, falsely polygamous trees; leaves alternate, simple, opposite, entire or tridentate at apex. Three spp., from Chile and Argentina.

3.3 SAPINDOIDEAE ▸ TRIBE MELICOCCEAE (4/64) - outsiders Castanospora (1; Queensland, New South Wales), Tristira (1; Philippines, Sulawesi, Moluccas).

13. Melicoccus P.Browne. Trees, dioecious or monoecious; leaves alternate, paripinnate; leaflets 1–2 pairs. 10 spp., M. oliviformis Kunth disjunct Mexico and tropical South America, one endemic to Dominican Republic, and 8 confined to South America; 4 in Brazil, one endemic.

14. Talisia Aubl. Unbranched shrubs or small to large trees, duodichogamous; fruits indehiscent. 55 spp., only one absent in South America, most of which occur in the dense, lowlands humid forest, 35 spp. in Brazil, 8 endemics.

3.4 SAPINDOIDEAE ▸ TRIBE CUPANIEAE (38/465–470) - oustiders Diploglottis (12; New Guinea, Queensland, New South Wales), Podonephelium (4; New Caledonia), Alectryon (c 25; E Malesia to New Guinea, Australia, Solomon Islands, New Caledonia, Vanuatu, Fiji, New Zealand, Samoa, Hawai), Elattostachys (c 20; Malesia to New Guinea, Queensland, New South Wales, Solomon Islands, Vanuatu, New Caledonia, Fiji, Tonga, Niue, Samoa), Jagera (1; E Malesia to New Guinea, Queensland, New South Wales), Guioa (c 65; Thailand, Malesia to New Guinea, Queensland, New South Wales, New Caledonia, Fiji, Samoa, Tonga), Mischocarpus (c 15; SE Asia, Malesia to New Guinea, Queensland, New South Wales), Sarcopteryx (12–13; Moluccas, New Guinea, Queensland, New South Wales), Molinaea (9; Madagascar, Mascarene Islands), Tina (17; Madagascar), Mischarytera (3; New Guinea, Queensland), Gongrodiscus (3; New Caledonia), Podonephelium (9; New Caledonia), Storthocalyx (5; New Caledonia), Rhysotoechia (c 15; Borneo, Philippines, Sulawesi, Moluccas, New Guinea, Queensland, New South Wales), Lepiderema (8; New Guinea, Queensland), Sarcotoechia (10–11; Moluccas, New Guinea, Queensland), Toechima (c 8; Flores, New Guinea, Queensland, New South Wales), Arytera (28; India, SE Asia, Malesia to New Guinea, Solomon Islands, Northern Territory, Queensland and New South Wales, Fiji, Samoa, Tonga), Synima (1; tropical Africa), Blighiopsis (1; Congo), Cnesmocarpon (4; New Guinea, Queensland), Cupaniopsis (c 60; Sulawesi, New Guinea, Solomon Islands, Australia, New Caledonia, Fiji, Samoa, the Caroline Island), Gloeocarpus (1; Philippines), Gongrospermum (1; Philippines), Lychnodiscus (7; tropical Africa), Pavieasia (3; S China, Vietnam), Phyllotrichum (1; Laos), Sisyrolepis (1; Thailand, Cambodia), Trigonachras (9; Malesia).

15. Alatococcus Acev.-Rodr. Small to medium sized trees (14–19 m tall); leaves alternate, paripinnate, leaflets entire; inflorescence distal or axillary, paniculate thyrses, with flowers in lateral subcincinni; flowers functionally unisexual; calyx zygomorphic, sepals 5, free, imbricate, the outer 2 smaller; petals 5, clawed, twice as long as the sepals; fruit of a single, woody, sub-globose, winged mericarp. Only one sp., A. siqueirae Acev.-Rodr., known only gallery and tall forests of Espírito Santo state, E Brazil.

16. Cupania L. Small trees; leaves pinnately compound; leaflets mostly with serrate margins; fruits capsular; seeds arillate at the base. 52 spp., in dense, lowlands, humid Neotropical forests; 41 spp. in South America, 33 in Brazil, 21 endemics.

17. Dilodendron Radlk. Trees, falsely polygamous-dioecious; leaves alternate, bipinnate or sub-tripinnately compound. Three spp. from Central America to Bolivia and Venezuela, D. bipinnatum Radlk. up to Brazil and Paraguay.

18. Matayba Aub. Small to large trees, falsely polygamous-dioecious,; leaves alternate, paripinnate or imparipinnate. 45 spp. in dense, lowlands, humid Neotropical forests; 38 spp. in South America, 29 in Brazil, 13 endemics; Radlkofer divided Matayba into four sections, with sect. Matayba as the largest section, with 17 species restricted to South America; in Brazil, this section is predominantly extra-Amazon rainforest, except for M. guianensis Aubl., with a widely distributed distribution, and M. atropurpurea Radlk. which is found in the Brazilian and Colombian Amazon rainforest; another species, M. elaeagnoides Radlk., is found in S Brazil and adjacent Paraguay and NE Argentina.

19. Pentascyphus Radlk. Trees or shrubs, falsely polygamous; leaves alternate, paripinnate; leaflets alternate, entire. Only one sp., P. thyrsiflorus Radlk., occurring in French Guiana, Suriname and Amazonas state in N Brazil.

20. Scyphonychium Radlk. Trees, falsely polygamous; leaves alternate, paripinnate; leaflets entire; distal leaflet rudimentary; inflorescences terminal thyrsoids. Only one sp., S. multiflorum (Mart.) Radlk., native to N and E Brazil, and French Guiana.

21. Tripterodendron Radlk. Trees, falsely polygamous-dioecious trees; leaves alternate, tripinnate. Only one sp., T. filicifolium Radlk., endemic to E Brazil.

22. Vouarana Aubl. Medium-sized trees, falsely polygamous; leaves alternate, paripinnate; leaflets entire; distal leaflet rudimentary. Two spp., from Costa Rica to Ecuador, Guianas and N Brazil (both species, none endemics).

3.5 SAPINDOIDEAE ▸ TRIBE ATHYANEAE (2/3) - both genera occur in South America.

23. Athyana Radlk. Trees, monoecious leaves alternate, imparipinnate. Only one sp., A. weinmanniifolia (Griseb.) Radlk., from Peru, Bolivia, Paraguay and Argentina.

24. Diatenopteryx Radlk. Shrubs or trees, falsely polygamous; indumentum of simple hairs and scales. Two spp., D. sorbifolia Radlk. from in Brazil, Bolivia, Paraguay, and Argentina, and D. grazielae Vaz & Andreata endemic to Brazil.

3.6 SAPINDOIDEAE ▸ TRIBE BRIDGESIEAE (1/1) - a single genus.

25. Bridgesia Bertero ex Cambess. Shrubs, falsely polygamous shrubs; leaves alternate, simple, lobed or serrate. Only one sp., B. incisifolia Bertero ex Cambess., from Andean Chile.

3.7 SAPINDOIDEAE ▸ TRIBE THOUINIEAE (3/c. 250) - all genera occur in South America.

26. Allophylastrum Acev.-Rodr. Small trees or shrubs (to 7 m tall); leaves alternate, trifoliolate; leaflets serrate; flowers solitary, axillary or in short racemes, actinomorphic; petals wanting; fruit of 1–2 basally connate, indehiscent monocarps, with fleshy exocarp, and a semi-woody endocarp; seeds exarillate, with papery testa. Only one sp., A. frutescens Acev.-Rodr., from Guyana and Roraima state in N Brazil, in terra firme forests.

27. Allophylus L. Shrubs or trees, duodichogamous or dioecious, or less often scandent shrubs; ubiquitous shrubs or small trees. c. 255 spp., pantropical, 57 in humid, lowlands to mid-elevation forests in the Neotropics; 45 spp. in South America, 27 in Brazil, 9 endemics.

28. Thinouia Planch. & Triana. Lianas; secondary growth of stems with numerous cortical steles; actinomorphic leaves with umbeliform inflorescences. 12 spp., from Central and South America (11, 9 confined, but one endemic to Mexico); 9 in Brazil, 4 endemics.

3.8 SAPINDOIDEAE ▸ TRIBE PAULLINIEAE (6/475–480) – all genera in South America, and mainly neotropical with the exception of Paullinia pinnata L., Cardiospermum corindum L., and C. halicacabum L., which in addition to their Neotropical ranges, are naturally distributed in parts of the Paleotropics.

29. Cardiospermum L. (exc. Serjania p.p.) Monoecious herbaceous vines and less frequently erect shrubs or subshrubs; inflated capsules; leaves alternate, ternately compound or biternate. 9 spp., native to the Neotropics, C. halicacabum L. and C. corindum L. are now widely distributed throughout the tropics in Africa and Asia, C. grandiflorum Sw. over tropical New World, one in Mexico and S U.S.A., 4 endemics to Brazil and one endemic to Peru.

30. Lophostigma Radlk. Woody vines; cross section of stem with a single vascular cylinder. Two spp. in Bolivia, one up to Peru and Ecuador.

31. Paullinia L. Woody vines, usually producing milky sap; coriaceous to woody capsules, opening to expose arillate seeds. 192 spp. in the Neotropics, one, P. pinnata L., extending to Africa and Madagascar; most diverse in the dense lowland to mid-elevation forests; most speciose in Brazil (104, 32 endemics), Peru and Colombia, 175 in over South America.

P. unifoliolata Perdiz & Ferrucci from S Bahia is the only even unifoliate species in this genus; others species has unifoliate and compound leaves together; Paullinia display a wide diversity of fruit morphologies.

32. Serjania Mill. (inc. Cardiospermum p.p.) Vines, woody or herbaceous, sometimes with xylopodium; duodichogamous, often producing milky sap. 241 spp., abundant in open vegetation such as savannas, Atlantic sandy coastal shrublands (restingas), shrubby savannas and forest margins; most diverse in the Brazilian Shield (with around 100) and central Mexico; 168 spp. in South America, 122 in Brazil, 68 endemics.

33. Urvillea Kunth. Herbaceous to woody vines; stems terete and lenticellate, becoming trilobate at age, producing milky sap. 18 spp. of Peru to Cono Sur and Brazil (16, 10 endemics), with U. ulmacea Kunth in H. B. K. extends into the Lesser Antilles and North America and one endemic to Mexico.

LINEAGE 3 of 3: MELIACEAE/RUTACEAE/SIMAROUBACEAE

MELIACEAE

§ FULLY CHLOROTROPHIC

Genera/species 58/590-600 Distribution tropical and subtropical lowland areas, mainly southern and SE Asia; few species in warm-temperate regions; Xylocarpus consists of mangrove trees. Habit bisexual, monoecious, polygamomonoecious or dioecious, evergreen trees and shrubs (in Munronia suffrutices, in Naregamia perennial herbs). Bark often with a bitter taste, often with a strong garlic-like or sweet smell, etc.

Trees, treelets or rarely shrubs; plants monoecious, dioecious or polygamous; Cedrela odorata L. is the most commercially important and widely distributed species in their genus. Known as Spanish-cedar in English commerce, and cedro in Brazil, the aromatic wood is in high demand in the American tropics because it is naturally termite- and rot-resistant; oil of seeds from Carapa guianensis Aubl. in highly medicinal porposes.

SYSTEMATIC two main clades can be discerned, corresponding to Melioideae and Cedreloideae, both in South America.

1. SUBFAMILY MELIOIDEAE (c 35/530–540) ‣ oustiders Azadirachta (2; tropical Asia), Melia (2; southern tropical Africa, tropical Asia to New Guinea), Astrotrichilia (c 12; Madagascar); Quivisianthe (1–2; Madagascar), Walsura (16; India and Sri Lanka to New Guinea); Sandoricum (4; Malesia, Borneo); Munronia (3; subtropical China and tropical Asia to Timor), Lepidotrichilia (4; tropical E Africa, Madagascar), Vavaea (4; Sumatra and Philippines to N Australia, Fiji, the Caroline Islands and Tonga), Pseudoclausena (1; Indochina to New Guinea), Cipadessa (1; tropical and subtropical Asia from India, Sri Lanka and Nepal to S China, Indochina and Central Malesia), Ekebergia (4; tropical and S Africa), Owenia (5; Australia), Malleastrum (23; Madagascar, the Comoros, Aldabra), Pterorhachis (2; Central Africa), Nymania (1; S Namibia, N, W and E Cape), Calodecaryia (2; Madagascar), Humbertioturraea (10; Madagascar), Turraea (c 60; tropical and S Africa, Madagascar, Mascarene Islands, Socotra, tropical Asia to N and E Australia), Synoum (1; Queensland, New South Wales), Anthocarapa (1; Queensland, New South Wales, New Guinea to New Caledonia and Rotuma), Heckeldora (7; tropical W and C Africa), Turraeanthus (3; tropical Africa), Dysoxylum (c 80; India and Sri Lanka to S China, Indochina, Malesia to New Guinea, Christmas Island, islands in S and SW Pacific to N and E Australia, New Caledonia, Norfolk Island, Lord Howe, New Zealand, Tonga to Niue), Chisocheton (53–55; Assam, S China, SE Asia, Malesia to Queensland and Vanuatu), Sphaerosacme (1; Himalayas), Aglaia (c 120; tropical Asia to islands in W Pacific), Aphanamixis (3; tropical Asia to Solomon Islands), Lansium (1; Malesia), Reinwardtiodendron (7; SE Asia, W and C Malesia).

1. Cabralea A.Juss. Tree or treelet up to 30 m tall; hairs simple; leaves usually pinnate. Only one variable sp., C. canjerana (Vell.) Mart., from Costa Rica through tropical South America to N Argentina.

2. Guarea Allam. ex L. Dioecious trees or treelets; indumentum of simple hairs; leaves pinnate, with a dormant terminal bud or more rarely a terminal leaflet; leaflets 8–15, oblong-elliptic, entire; calyx irregularly lobed; petals 5, free from staminal tube; fruit a cleistocarp, indehiscent or breaking up after falling; seeds large, completely covered by a thin yellow aril. 82 spp., 77 in tropical America (51 in South America) and 5 in tropical Africa; G. cartaguenya from Bajo Calima region, Choco, W Colombia, has the largest leaf among Meliaceae; 27 spp. in Brazil, 4 endemics; G. crispa T.D.Penn. from Amazonas state is a rare plant in Brazil, by Plantas Raras do Brasil’s book.

3. Ruagea Karsten. Trees and treelets, dioecious; bud-scales rare. 15 spp. of montane rain forest and cloud forest from South America (Venezuela to Bolivia), two up to Guatemala.

4. Trichilia P.Browne. Trees and treelets, usually dioecious, rarely polygamous. 108 spp., in mainland lowland tropical America (84, Mexico to Argentina, Caribbean, 70 in South America), 18 spp. in Africa, 6 spp. in Madagascar; frequent understory trees in Amazon rainforest; 53 in Brazil, 15 endemics; T. discolor A.Juss from Pará and T. florbranca T.D.Penn. from Bahia are rare plants in Brazil, by Plantas Raras do Brasil’s book.

2. SUBFAMILY CEDRELOIDEAE (c 10/33–35) ‣ outsiders Chukrasia (1; India, Sri Lanka and S China to W Malesia), Neobeguea (3; Madagascar), Toona (5; Pakistan to S China, SE Asia to Queensland and E New South Wales), Capuronianthus (2; Madagascar), Lovoa (3; tropical Africa), Xylocarpus (3; mangroves in E Africa to Tonga), Khaya (6; tropical Africa, Madagascar), Entandrophragma (11; tropical Africa), Pseudocedrela (1; tropical Africa), Soymida (1; India, Sri Lanka).

5. Carapa Aubl. Trees of very variable habit, up to 40 m tall; leaves usually paripinnate, and with an apical gland, mostly crowded at ends of stout branchlets; flowers unisexual, 4- or 5(6)-merous, in large, much-branched panicles; fruit a large, septifragal capsule, breaking open on hitting ground; seeds 12–20, large, subangular, with a woody but buoyant outer covering. 11 spp., 10 in South America, 3 in Brazil, none endemics, inc. C. guianensis Aubl. (sensu lato) widely distributed in the neotropics.

6. Cedrela P.Browne. Trees, deciduous up to 50 m tall; leaves usually paripinnate; leaflets entire. 19 spp. in the Neotropics, 15 in South America, only three in Brazil, none endemics; one of the world’s most important timber tree genera..

7. Schmardaea H. Karst. Trees or shrubs, deciduous. Only one sp., S. microphylla (Hook.) H. Karst. ex Müll. Hal., Andes from Venezuela to Peru, up to 2,700 m in cloud forests.

8. Swietenia Jacq. Trees, deciduous, leaves usually paripinnate, up to 45 m tall; leaflets entire. Three spp., S. humilis Zucc. in Pacific coast of Central America and Mexico; S. macrophylla King, mogno wood, in Atlantic coast of Central America, South America south to Bolivia and Brazil; and S. mahagoni (L.) Jacq., West Indian Mahogany, from S Florida, Cuba, Jamaica, Hispaniola.

RUTACEAE

§ FULLY CHLOROTROPHIC

Genera/species 152/1,650-1,700 Distribution tropical, subtropical and warm-temperate regions in the Northern and Southern Hemispheres, with their largest species diversities in southern South Africa and Australia. Habit usually bisexual (rarely monoecious, andromonoecious, polygamomonoecious, or dioecious), evergreen or deciduous trees, shrubs or lianas (rarely perennial herbs). Often strongly aromatic. Many species are xerophytes.

About 160 genera and 1,900 spp. of Rutaceae were recognised, of which 48 genera and 350 spp. occur in the Neotropics (Kallunki 2004), 44-50 of these species belong to the genus Conchocarpus (Galipeinae). Rutaceae is widely distributed in subtropical and tropical or less frequently in temperate regions of the world. In the Neotropics the family is more diverse in the understorey of moist forests, especially in Brazilian Atlantic rain forests (Galipeinae), but spp. of this group can also be found in drier areas (e.g. Casimiroa, Zanthoxylum, Balfourodendron and Helietta). Cneoridium, Thamnosma, and some Zanthoxylum, Ptelea and Choisya spp. occur in desertic or temperate areas in Mexico; and the Pitavia occurs in temperate areas in Chile. Only Thamnosma (also present in African) and (sub)tropical worldwide Zanthoxylum also occurring in other continents.

Rutaceae are best known by the exotic genus Citrus, because of its commercially consumed fruits. Other groups of the same subfamily of Citrus (Aurantioideae, see discussion above) are commonly cultivated as ornamentals in America, including species of Atalantia, Clausena, Murraya, and Swinglea. Other extra-American cultivated genera are Dictamnus, Phellodendron, and Ruta. Native species are used as commercial timbers, such as Euxylophora paraensis Huber (pau-amarelo), Balfourodendron riedelianum (Engl.) Engl. (pau-marfim), both from Brazil, and Zanthoxylum flavum Vahl (West Indian silkwood); Pilocarpus (jaborandis) are sources of the alkaloid pilocarpine, used to treat glaucoma: one of the species, P. microphyllus Stapf ex Wardlew. is commercially cultivated in Brazil. The bark of some species of Angostura, Galipea and Hortia are used to treat fevers. Casimiroa edulis La Llave & Lex. (zapote blanco, white-zapote) is cultivated for its edible fruits in Central America.

SYSTEMATIC six subfamilies, Haplophyllodeae (1/66, western Mediterranean and N and NE Africa, through Arabia and central Asia to China), Auranthioideae (27/205, Africa, Asia, Australasia), and Rutoideae (5/20, temperate and tropical regions of the Northern Hemisphere as well as southern Africa) is absent in South America.

1. SUBFAMILY AMYRIDOIDEAE (3/42) ‣ outsiders Cneoridium (1, California to NW Mexico), Stauranthus (2, S. Mexico to Central America).

1. Amyris P. Browne. Glabrous shrubs or trees, sometimes armed with short, axillary spines. 47 spp., U.S.A. (Texas, Florida), Mexico, Central America (including Caribbean) to Peru disjunct in French Guiana; 13 spp. occur in South America.

2. SUBFAMILY CNEOROIDEAE (8/35) ‣ outsiders Spathelia (3; Caribbean), Harrisonia (2; tropical Africa, tropical Asia, N Australia), Cneorum (2; Canary Islands; W Mediterranean), Ptaeroxylon (1; N Tanzania to South Africa), Cedrelopsis (8; Madagascar), Bottegoa (1; S Somalia, Ethiopia, Kenya).

2. Dictyoloma A. Juss. Monoecious small trees; multicellular oil glands only on margins of the leaflets, their cell walls not resorbed. Only one sp., D. vandellianum A.Juss., from Ecuador, Peru, Bolivia and C Brazil.

3. Sohnreyia K. Krause (inc. Spathelia p.p.). Monocarpic polygamous trees with simple unbranched trunks conspicuously marked with leaf scars, pachycaulous, palm-like. 5 spp., N South America, only S. excelsa K. Krause in Brazil, Rondonia and Mato Grosso states up to Bolivia, non endemic.

3. SUBFAMILY ZANTHOXYLOIDEAE (109/c. 1,700) ‣ more than 50 outsiders.

4. Adiscanthus Ducke. Shrubs or small unbranched trees; leaves alternate, simple. Only one sp., A. fusciflorus Ducke, lowland forests of N South America in Colombia, Venezuela, Brazil and Peru, E of Tapajos River, in Brazil.

5. Andreadoxa Kallunki. Trees; leaves alternate, 1-foliolate; inflorescence an axillary thyrse, several borne near apex of branch. Only one sp., A. flava Kalunki, a rare plant in Brazil, by Plantas Raras do Brasil’s book, with yellow flowers very narrow endemic to forests of SE Bahia, only in Itabuna municipality, known only a single specimen in a human area.

6. Angostura Roem. & Schultes. Trees or shrubs; trichomes echinoid or stellate; leaves alternate, (1)3-foliolate or sometimes digitately 4–7-foliolate. 9 spp., Central America (Nicaragua) and southwards to Venezuela, S Brazil (4, three endemics, one up to Bolivia) and Bolivia; 7 in South America.

7. Apocaulon R. Cowan. Decumbent, pubescent, rhizomatous herbs; leaves alternate, congested. Only one sp., A. carnosum R.S. Cowan, endemic to the Guiana Shield in montane forests of southern Venezuela, 600-1,300 m elevation range.

8. Balfourodendron Méllo ex Oliv. Trees or treelets up to 35 m tall; leaves opposite, digitately 3-foliolate. Two spp., both in NE to SE Brazil, one up to adjacent Paraguay and Argentina, in (semi) deciduous forests.

9. Conchocarpus Mikan. (inc. Almeidea, exc. Dryades) Shrubs or small trees, often unbranched; trichomes simple. 49 spp., all in South America, 47 in Brazil (31 endemics; two species from Central America to Venezuela do not occur in Brazil) up to Central America (4) in over tropical South America, mainly in forests; c. 14 spp. in Bahia, Espírito Santo and Rio de Janeiro states are rare plants in Brazil, by Plantas Raras do Brasil’s book.

10. Decagonocarpus Engl. Shrubs or small trees; leaves opposite, simple; inflorescence a terminal, few-flowered monochasium. Two spp., endemics to Guiana Shield in S Venezuela, Amazonas state in N Brazil and E Colombia (both in all three countries), 100-1,800 m elevation range.

11. Dryades Groppo, Kallunki & Pirani. (off Conchocarpus) Shrubs or treelets up to 6 m tall, the stems usually single or with few, orthotropic, relatively thick branches and often bearing leaves congested at apex of the stem or branch or less often diffusely branched and leaved; leaves ascending or patent, alternate, 1-foliolate, flowers bisexual, 5-merous, pedicellate; corolla tubular, zygomorphic or (sub) actinomorphic; petals creamy-white, fruit of 1–5 follicular mericarps. 5 spp., endemics to the Atlantic Forest domain in E Brazil, ranging from from the states of Pernambuco and Bahia to Santa Catarina; three species inhabit the understory of moist forests, and two species inhabit the Atlantic sandy coastal shrublands (restingas).

12. Ertela Adans. Erect herbs, sometimes suffruticose, stems dichotomously or trichotomously forked above. Two spp., E. trifolia (L.) Kuntze widely distributed from SW Mexico and disjunct in N South America to Peru, Bolivia and Rio de Janeiro state; and E. bahiensis Kuntze endemic to E Brazil.

13. Erythrochiton Nees & Mart. Shrubs or small trees, often un–few-branched, sometimes flageliflorous in N Andes. 7 spp., Costa Rica across N South America and South to Bolivia and Brazil (two spp., none endemics). E. hypophyllanthus Planch., endemic to Colombia, is the one most remarkable of all Rutaceae, with epiphyllous inflorescences in lower surface of leaves.

14. Esenbeckia Kunth. Shrubs or small trees, rarely subshrubs. 36 spp., Caribbean, and from U.S.A. and Mexico to north Argentina, rarely represented in Amazon rainforest, ranging from dry woodland to moist forest, slightly centered in Mexico and Brazil (15, 5 endemics); 20 spp. in South America.

15. Euxylophora Huber. Tall tree up to 30 m tall; wood yellow; leaves alternate, simple; inflorescence a terminal, corymbiform thyrse. Only one species, E. paraensis Huber, from SE Colombia and Pará to Maranhão states in N Brazil; found in dry land forests in the Amazon rainforest.

16. Galipea Aubl. Trees or shrubs; trichomes simple; leaves alternate, (1)3-foliolate. 13 spp., one Central America, remaining in tropical South America (11 confined to continent); 9 spp. in Brazil, 5 endemics; G. carinata Pirani from Espírito Santo state is a rare plant in Brazil, by Plantas Raras do Brasil’s book.

17. Helietta Tul. Shrubs or trees; leaves (sub) opposite, digitately 3-foliolate; inflorescence a terminal, simple or compound thyrse. 9 spp., three in Mexico and U.S.A. (Texas), one endemic to Cuba; 4 in S South America in Peru, Bolivia, Paraguay, N Argentina and Brazil (3, one endemic) and one endemic to Venezuela.

18. Hortia Vand. Trees or shrubs with simple leaves crowded near the apices of the branches, showy broad corymbose terminal inflorescences, reddish to pink flowers, and baccate fruits with abundant oil glands. 10 spp., Panamá, N South America to São Paulo state in Brazil (9, 4 endemics, three of then, all from Amazonas state, are rare plant in Brazil, by Plantas Raras do Brasil’s book).

19. Leptothyrsa Hook.f. Shrubs or small trees not ramified; leaves alternate, simple, very long, crowded at tip of stems. Only one sp., L. sprucei Hook.f., lowland forests of Amazonas state in N Brazil, Colombia and Peru.

20. Lubaria Pittier. Tree; leaves simple, opposite; inflorescence a terminal, once- or twice-forked dichasium with monochasial branches. Two spp., one in Costa Rica and Venezuela, another endemic to Colombia.

21. Metrodorea St-Hill. Shrubs or trees; leaves opposite, 1–3-foliolate. 6 spp., all in Brazil and endemic within, except M. flavida K. Krause, which also occur in adjacent Bolivia, Peru and Guianas.

22. Naudinia Planch. & Linden. Shrub, leaves alternate, 1-foliolate; inflorescence an extra-axillary cyme. Only one sp., N. amabilis Planch. & Linden, Colombia.

23. Neoraputia Emmerich. Trees; trichomes simple; leaves alternate (sometimes opposite). 6 spp., 4 endemic to E Brazil, N. magnifica (Engl.) Emmerich ex Kallunki up to Peru, and N. paraensis (Ducke) Emmerich ex Kallunki in over Amazon rainforest up to Central America.

24. Peltostigma Walp. Shrubs and small trees; leaves alternate, digitately 3–5- or 1-foliolate. Two spp., Mexico, Caribbean, Guatemala to Nicaragua, P. guatemalense (Standl. & Steyerm.) Gereau up to Colombia and Ecuador.

25. Pilocarpus Vahl. Shrubs or trees; leaves alternate. 19 spp., one in Caribbean, all remaining in South America (17 confined), with one up to Mexico, centered in Brazil (15, 10 endemics); moist and dry forest and woodland; absent from the Amazon rainforest.

26. Pitavia Molina. Dioecious small trees or shrubs; trichomes simple; leaves simple, opposite, or in trimerous whorls. Only one sp., P. punctata Mol., coastal Cordillera of Chile.

27. Raputia Aubl. Shrubs or trees; leaves opposite, 1- or 3-foliolate. 15 spp., northern South America up to Brazil (8, two endemics, both rare plants in Brazil, by Plantas Raras do Brasil’s book), mostly in lowland forest, only three outside Amazon rainforests.

28. Raputiarana Emmerich. Shrub; leaves alternate, digitately 5–7-foliolate. Only one sp., R. subsigmoidea (Ducke) Emmerich, Amazon rainforest of N Brazil, Ecuador, Colombia.

29. Rauia Nees & Mart. Shrubs or small trees; leaves alternate, 1- or 3- foliolate. 5 spp., two only in Guiana Shield, two endemics to Brazil, and R. resinosa Nees & Mart. widely distributed in E tropical South America.

30. Raulinoa R. Cowan. Shrub armed species with reddish flowers. Only one sp., R. echinata R. S. Cowan, a rare plant in Brazil, by Plantas Raras do Brasil’s book, known only in rocky riversides of Itajaíaçu River in Santa Catarina state, S Brazil.

31. Ravenia Vell. Shrubs or small trees; leaves opposite or appearing alternate due to reduction of one of two opposite leaves, simple or 3-foliolate. 10 spp., 5 only in Caribbean, three endemics to Brazil (R. pseudalterna Ducke is a rare plant in Brazil, by Plantas Raras do Brasil’s book), R. rosea Standl. from Colombia to Honduras, and R. biramosa Ducke from Amazon rainforest of Peru, Colombia and N Brazil.

32. Raveniopsis Gleason. Shrubs or small trees; trichomes simple or stellate-lepidote. 20 spp., 18 spp. endemic to Guiana Shield of S Venezuela (16, 14 endemics) and N Brazil, (300–) 1,000–2,600 m above sea level; and two spp. extending to central Brazil; two species, one from Mount Aracá and one from Rondonia, are rare plants in Brazil, by Plantas Raras do Brasil’s book.

33. Rutaneblina Steyerm. & Luteyn. Low shrubs; leaves simple; inflorescence an almost sessile, condensed or corymbiform dichasium. Only one sp., R. pusilla S. & L., endemic to Pantepui Life Zone (Venezuelan side of Mount Neblina) in Guiana Shield of S Venezuela.

34. Sigmatanthus Huber ex Emmerich. Small tree or shrub; leaves alternate, trifoliolate; inflorescence terminal, raceme-like. Only one sp., S. trifoliatus Huber ex Emmerich, NE Brazil, from Pará to Rio Grande do Norte.

35. Spiranthera A. St-Hil. (inc. Nycticalanthus) Shrubs or trees; leaves alternate, 3-foliolate; inflorescence a terminal thyrse. 5 spp., S. odoratissima A. St.-Hil., widely distributed in savannas of C Brazil (cerrado) and open formations of Brazil and Bolivia; three other species are Amazonian forest trees or treelets: S. guianensis Sandwith, from Guyana, Venezuela, and NW Brazil; S. speciosa (Ducke) L.A.Brito & Pirani, very narrow endemic to Manaus vicinity in central Amazon rainforest of Brazil, a rare plant in Brazil, by Plantas Raras do Brasil’s book; S. parviflora Sandwith, from Guyana, Venezuela, Colombia, Peru, and northern Brazil; and S. atlantica Pirani represents the only species record for the genus in the Atlantic Forest of E Brazi (two collections from Espırito Santo).

36. Ticorea Aubl. Trees or shrubs; leaves alternate, 3(4)-foliolate, domatiiferous. 5 spp., lowland forests of the eastern slopes of the Andes in Ecuador, Peru and Bolivia, and in Brazilian Amazon rainforest (4, none endemics) and the Guianas.

37. Toxosiphon Baill. Shrubs or small trees, often unbranched; leaves alternate, 1- or 3-foliolate, often clustered at apex of stem. 4 spp., T. lindenii (Planch.) Baill. ranging from southern Mexico through Panamá, and three from Ecuador to northern Bolivia and adjacent Brazil (2, none endemics), in the understorey of humid tropical forests.

38. Zanthoxylum L. (inc. Fagara) Dioecious, monoecious or polygamous, deciduous or evergreen trees or rarely scandent shrubs, often aculeate or spinose, sometimes with knobs tipped with spines (knobthorns). c.180-200 spp., (sub) tropics worldwide, 134 in New World, widely distributed, 79 in South America, absent in continental Chile, but Z. mayu Bert. present in Juan Fernandes Is, in Pacific Ocean. 27 in Brazil, 11 endemics.

Z. magnifructum Reynel (Colombia) has the largest fruits of any species in the genus in the New World.

SIMAROUBACEAE

§ FULLY CHLOROTROPHIC

Genera/species 20/140–145 Distribution tropical and subtropical regions, a few species (e.g. of Ailanthus) in temperate E Asia (Korean Peninsula, Japan). Habit monoecious, polygamomonoecious, dioecious, or polygamodioecious (androdioecious) (sometimes bisexual), evergreen or deciduous trees or shrubs. Bark, wood and seeds often very bitter-tasting. Branch medulla characteristically light-coloured and well-developed.

The Simaroubaceae are a primarily tropical family of about 200 species in 20-30 genera. The primary center of diversity for the family is in tropical America with a secondary center in tropical W Africa. Most species are dioecious trees or shrubs with alternate compound leaves, five-parted flowers and drupaceous fruits.

The anemophilous Leitneria floridana Chapman is sister to the [Brucea+Soulamea] clade. Leitneria is characterized by a combination of features which is unique to Simaroubaceae, i.e., dioecy, silky young branches, bark not bitter, catkin-like cymose inflorescences consisting of numerous reduced dichasia, reduced flowers without calyx (at least in male flowers; in female flowers sepaloid bracts/bracteoles?), disc absent, four stamens with basifixed anthers, pollen grains with reticulate exine, one carpel (in reality two carpels, one of which reduced and rarely developing), unilocular (pseudomonomerous) ovary, decurrent stigma, parietal placentation, ascending ovules, bistomal micropyle, drupe, endotegmic seed-coat, mesotegmen with reticulate thickenings, starchy endosperm, and n = 16.

SYSTEMATIC the both subfamilies occur in South America.

1. SUBFAMILY SIMAROUBEAE (19/9-100) ‣ outsiders Ailanthus (6; Turkestan, India, Sri Lanka, China (inc. Taiwan), SE Asia, Malesia to New Guinea, N and E Australia); Nothospondias (1; C Africa); Leitneria (1; SE U.S.A.), Brucea (8–10; tropical regions in the Old World), Soulamea (14; Seychelles, New Caledonia, Fiji, SE Asia and Malesia to Polynesia); Samadera (5–6; Madagascar, SE Asia to Queensland), Gymnostemon (1; Côte d’Ivoire), Perriera (2; Madagascar), Hannoa (5–7; Central Africa), Odyendea (1; tropical Africa), Eurycoma (3; tropical SE Asia, W Malesia to Philippines), Iridosma (1; C Africa), Pierreodendron (2; tropical W and Central Africa to Angola).

1. Homalolepis Hook. f. Subshrubs with aerial stem absent or very short, usualy with xylopodium, shrubs usually broadly branching, treelets with a slender, unbranched stem (palmlike habit) or large trees generally with broad canopy. 28 spp., SE Brazil (25, 21 endemics) to El Salvador in Central America, mostly concentrate in extra-Amazon rainforest regions. H. suaveolens (A. St.-Hil.) Devecchi & Pirani from Minas Gerais state is a rare plant in Brazil, by Plantas Raras do Brasil’s book; two sections:

§ sect. Homalolepis ‣ 15 spp., mainly in the extra-Amazon rainforest part of South America; only two occur exclusively in Amazon rainforest, and H. guajirensis Devecchi, W.W. Thomas & Pirani is endemic to dry formations of northern Colombia and Venezuela.

§ sect. Grandiflorae ‣ 13 spp., including the one with the broadest distribution in the genus (H. cedron (Planch.) Devecchi & Pirani), mainly geophytic subshrubs with leaves crowded at ground level, or treelets with a palmlike habit.

2. Picrolemma Hook. f. Small shrubs, up to 6 m, dioecious; leaves imparipinnate. Two spp., P. huberi Ducke known from Colombia to Peru and P. sprucei Hook. f. from the Amazon rainforest.

3. Quassia L. Trees or shrubs; branchlets glabrous; leaves imparipinnate, 2–5(–8)-jugate; not crowded at tips of branches; leaflets ± opposite, usually with pitted glands in upper (and sometimes in lower) surface; petiole and rachis not winged; flowers in terminal and axillary thyrses; fruits of 1–3(4) drupaceous mericarps; seeds 1 per mericarp. Two spp., Q. amara L. in Mexico, Central America, Caribbean, Trinidad & Tobago, Venezuela, Colombia, Brazil, another in W Africa.

4. Simaba Aubl. 5 spp., overall in N South America, 4 in Brazil, none endemics, S. monophylla (Oliv.) Cronquist restricted to Venezuela and Guianas.

5. Simarouba Aubl. Shrubs and trees to 35 m, dioecious; leaves paripinnate or imparipinnate. 6 spp., three are each endemic to one of the Greater Antilles: Cuba, Hispaniola, and Puerto Rico; S. glauca DC. found in North America, Central America and Caribbean; and two in S America, the widely distributed S. amara Aubl., S. versicolor A. St.-Hil. from Brazil and adjacent Bolivia.

2. SUBFAMILY PICRASMATEAE (2/c 23) ‣ all genera occur in South America.

6. Castela Turpin. Shrubs, erect or trailing, or small trees to 5 m, dioecious, armed with (occasionally branching) thorns. 12 spp., Caribbean, Jamaica, Hispaniola, SW U.S.A. and N Mexico, 4 in South America, C. coccinea Griseb. and C. tweedii Planch. from Bolivia and Paraguay to Argentina and Brazil (only the latter), C. galapageia Hook. f. in the Galapagos Islands, and C. erecta Turpin found from Texas to South America, Caribbean and Brazil.

7. Picrasma Blume. Small trees, sometimes to 20 m, or shrubs, monoecious or dioecious. 8 spp., two in SE Asia, three only in South America (in Brazil only P. crenata Engl., up to Cono Sur; one endemic to Ecuador) and three from N South America north to Mexico and Caribbean.

42. HUERTALES

FAMILIES ABSENTS IN SOUTH AMERICA: GERRADINACEAE (1/2) AND PETENAEACEAE (1/1).

TAPISCIACEAE

§ FULLY CHLOROTROPHIC

Genera/species 2/5–6 Distribution Tapiscia sinensis: S and SE China, N Vietnam; Huertea: Caribbean, the Andes from Colombia to Peru Habit bisexual or unisexual (male; andromonoecious), evergreen (Huertea) or deciduous (Tapiscia) trees.

Differs by Staphyleaceae by alternate leaves, wood anatomy, connate sepals, and seeds with a less well developed exotegmen.

SYSTEMATIC outsider Tapiscia (1–2; S and SE China, N Vietnam).

1. Huertea Ruiz & Pav. Evergreen trees or shrubs with reddish wood; mesophytic; leaves alternate; spirally arranged; compound; imparipinnate or trifoliolate, serrate; inflorescences terminal or from leaf axils; paniculate; flowers tiny; unisexual; perianth actinomorphic; petals free; stamens five, free of perianth; fruit a drupe; surface smooth; seeds 1-2; 1-10mm. 4 spp., H. cubensis Griseb. in Mexico, Honduras, Costa Rica, Cuba, R. Dominicana; H. glandulosa Ruiz & Pavon from Costa Rica, Colombia, Venezuela, Ecuador, Peru; H. granadina Cuatrec from Colombia and Venezuela; and H. putumayensis Cuatrec endemic to Colombia.

DIPENTODONTACEAE

§ FULLY CHLOROTROPHIC

Genera/species 2/18 Distribution Dipentodon: E Himalaya, NE India, N Burma, SE Tibet, SW China, Vietnam; Perrottetia: S China (inc. Taiwan), Malesia, New Guinea, NE Queensland, Hawaii; Mexico, Central America, northern Andes southwards to Peru. Habit bisexual or dioecious, evergreen trees or shrubs.

SYSTEMATICS outsider Dipentodon (1; NE India, E Himalayas, N Burma, SE Tibet, SW China, Vietnam).

1. Perrottetia Kunth. Bisexual or dioecious, evergreen trees or shrubs. 15 spp., two in Asia, 13 in New World, 9 only South America (7 in Colombia, 5 endemics), two shared in Mexico/Central/South America (both in Colombia), and two only Mexico/Central America; P. lanceolata H. Karst. is a Venezuelan endemic.

43. MALVALES

FAMILIES ABSENTS IN SOUTH AMERICA: NEURADACEAE (3/8), SARCOLAENACEAE (10/72), SPHAEROSEPALACEAE (2/20)

LINEAGE 1: THYMELAEACEAE

THYMELAEACEAE

§ FULLY CHLOROTROPHIC

Genera/species 52/c. 800 Distribution cosmopolitan except polar areas, although mainly in tropical Africa and Australia, Mediterranean, and West, East and SE Asia. Habit usually bisexual (sometimes andromonoecious, trimonoecious, polygamomonoecious, dioecious, androdioecious, or gynodioecious), evergreen or deciduous trees, shrubs or suffrutices (rarely lianas or perennial herbs), usually poisonous. Many species are xerophytes. Bark strongly fibrous. Use ornamental, textile and paper plants (bast fibres of Edgeworthia, Daphne, Lagetta, Thymelaea, Wikstroemia), incense (Wikstroemia) and perfumes (sesquiterpene alcohols in fungal infected wood), medicinal plants, cosmetics (Aquilaria etc.), timber.

With regards to Tepuianthus Maguire & Steyermark (previously in its own monogeneric family Tepuianthaceae), recent molecular and morphological data suggest that this genus is sister to subfamily Octolepidoideae (Horn, 2004). Aside from Tepuianthus and the closely related Octolepidoideae, molecular phylogenetic research within the family has mostly focused on the monophyletic subfamily Thymelaeoideae with emphasis on South African species of several genera and to a lesser extent, the large, mostly Australian genus, Pimelea Banks ex Gaertn.

New World members of the family account for 12 native genera and about 108 species; most groups are tropical and subtropical, found in lowland to montane environments. All American genera except Tepuianthus Maguire & Steyerm. belong to the much larger Thymelaeaoideae subfamily. Centers of species diversity are in Brazil (particularly around Rio de Janeiro), NW South America (especially Ecuador, Colombia, and Venezuela), and the Greater Antilles.

Several species of Daphne, a genus native to the Old World, are cultivated in the warm temperate zones of the New World; the tough phloem fibers are often twisted into cords and ropes, and the inner bark and fibers are sometimes used to make papers and textiles; fruit, leaves, and bark of many species are used as fish poison, and rarely also as a purgative; the family is economically important for ramin and agar wood (gaharu), with many populations threatened with extinction due to unsustainable extraction; in addition, several species are important for horticulture; Lagetta lagetto (W. Wright) Nash from the Carribbean has bark made up of many layers which has been used as gauze, lace or muslin.

Key differences from similar families The following features distinguish these families from the Thymelaeaceae:

ü Chrysobalanaceae - stipules present; glands usually present on petiole or leaf base; calyx and corolla present; petals free.

ü Clusiaceae - yellow sap present; opposite leaves usually with parallel venation; flowers unisexual; calyx and corolla present.

ü Ebenaceae - leaves distichous or spirally arranged with glands on the underside; calyx and corolla present.

ü Lamiaceae - leaves decussate; flowers zygomorphic; calyx and corolla present; stamens usually 2 or 4.

ü Rubiaceae - interpetiolar stipules present; stamens equal to the number of corolla lobes; ovary inferior.

SYSTEMATIC the isolate clade Synandrodaphne (1; Central Africa) and the subfamily Octolepidoideae (8/57–62, Tropical Africa, Madagascar, Malesia to New Guinea, NE Queensland, Melanesia) do not occur in South America.

1. SUBFAMILY TEPUIANTHEAE (1/7) - a single genus.

1. Tepuianthus Maguire & Steyerm. 6 spp. endemics to the Guiana Shield in 100 – 2,200 m elevation range, of Venezuela, adjacent Colombia, and Amazonas state in Brazil (2), with the shrubblet T. aracaensis Steyerm. & Maguire from Mount Aaracá in N Amazonas state is a rare plants in Brazil, by Plantas Raras do Brasil’s book.

2. SUBFAMILY THYMELAEOIDEAE (35/785 - 795) - outsiders Aquilaria (22; NE India, SE Asia, Malesia to New Guinea), Gyrinops (9; Sri Lanka, Laos, Malesia to New Guinea); Linostoma (3; tropical Asia, Australia), Jedda (1; Queensland), Enkleia (3; Andaman Islands, SE Asia, Malesia), Dicranolepis (c 20; tropical Africa), Synaptolepis (5; tropical and S Africa, Madagascar), Craterosiphon (11; tropical Africa), Linodendron (3; Cuba), Stephanodaphne (9; Madagascar, the Comoros), Phaleria (c 25; Sri Lanka, Malesia to New Guinea, Northern Territory, Queensland, New South Wales, the Caroline Islands), Peddiea (14; tropical and SE Africa, Madagascar), Dirca (4; S Canada, U.S.A., N Mexico), Lagetta (3; Caribbean), Daphne (90–95; Europe, Mediterranean, temperate Asia), Wikstroemia (c 90; SE Asia to islands in the Pacific incl. Hawaii), Rhamnoneuron (1; SE Asia), Edgeworthia (4; China, Japan), Thymelaea (c 35; S Europe, Mediterranean, temperate Asia, Himalayas, Tibet), Diarthron (c 15; Europe, W Asia), Stellera (1; Iran, C Asia, Himalayas, Tibet, China), Dais (2; south tropical and southern Africa; Madagascar), Struthiola (35–40; tropical and southern Africa, with their highest diversity in the Cape Provinces in South Africa), Lachnaea (35–40; N, W and E Cape), Passerina (c 20; Central to southern Africa, with their highest diversity in the Cape Provinces in South Africa), Kelleria (11; Sabah, New Guinea, Victoria, Tasmania, New Zealand), Gnidia (150–155; tropical and southern Africa, Madagascar, Arabian Peninsula, India, Sri Lanka, with their largest diversity in the Cape Provinces in South Africa), Pimelea (c 130; Philippines to Australia, Tasmania, Lord Howe, New Zealand).

2. Daphnopsis Mart. 69 spp., is by far the largest genus in the Americas, ranging from central Mexico to Chile and Argentina, also throughout most of the Antilles; 32 in South America, 12 occur in Brazil, 9 endemics.

3. Drapetes Banks ex Lam. Cushions. Only a single species, D. muscosus Lam., from S Chile, Argentina, and the Falkland Islands.

4. Funifera Leandro ex C. A. Mey. 4 spp., endemics to SE Brazil.

5. Goodallia Benth. Only one sp., G. guianensis Benth., distributed in the lowlands of Guyana and Roraima state in N Brazil, and remains poorly known.

6. Lasiadenia Benth. Two spp., L. rupestris Benth. from Amazonian rainforest of Venezuela, Colombia, Guyana and Amazonas state in Brazil, and L. ottohuberi T. Plowman & Nevling endemic to Venezuela.

7. Lophostoma (Meisn.) Meisn. 4 spp., Colombia, Venezuela and Amazonian rainforest of Brazil (3, two endemics).

8. Ovidia Meisn. Two spp., O. andina Meisn. from S Chile, Argentina and O. sericea Antezana & Z.S.Rogers from Bolivia.

9. Schoenobiblus Mart. 9 spp. from lowland tropical South America in Amazon rainforests; 3 spp. in Brazil, no endemics.

LINEAGE 2: BIXACEAE

BIXACEAE

§ FULLY CHLOROTROPHIC

Genera/species 4(includes Amoreuxia)/21–22 Distribution U.S.A., Mexico, Central America, Caribbean, tropical South America, western and central tropical Africa, Madagascar, India, Sri Lanka, northern Burma, Thailand and Indochina, northern Australia, eastern New Guinea. Habit bisexual, deciduous trees or shrubs (Cochlospermum), or perennial herbs with woody subterranean stem (Amoreuxia). Leaves in Diegodendron fragrant (like camphor). Juice coloured (orange to red). Some species are xerophytes. Juice coloured (yellow), resinous.

Testa in Bixa with red sarcotesta (with bixine, a carotenoid); Bixa orellana L., yelding a reddish food colouring commercially known as colorau, or corante, is widely cultivated and often naturalized througrouth tropics.

SYSTEMATIC outsiders Diegodendron (1; Madagascar)

1. Amoreuxia Moccino & Sesse ex. DC. Perennial herbs with woody subterranean stem. 4 spp. of U.S.A. to Central America up to Colombia (only A. palmatifida DC.), A. wrightii A.Gray disjunction of Peru and Ecuador.

2. Bixa L. Evergreen shrubs or small trees with raddish or yellow sap u to 30 m; flowers in terminal thyrsoid, large and showy (white or pink). 6 spp. in two morphological groups:

§ trees, confined to dense rainforest of Brazil, Ecuador, Bolivia and Peru: B. atlantica Antar & Sano, endemic to Bahia, Minas Gerais and Espirito Santo states in Atlantic Forest of SE Brazil; B. arborea Huber, described by Huber (1910) in Bragança, Pará; B. excelsa Gleason & Krukoff, described by Gleason and Krukoff (1934) along the Envira River, a tributary of the Juruá River, SW Amazon rainforest; and B. platycarpa Ruiz & Pav. ex G. Don, described by Ruiz and Pavón (Don 1831) in Peru.

§ shrubs, B. orellana L. which was described by Linnaeus (1753, never wild collected), and B. urucurana Willd., described by Willdenow (1809) in Brazil (without an identified locality).

3. Cochlospermum et al. Bisexual, seven species are trees and five that are suffrutescent subshrubs, often with taproot tubers or xylopodium. 12 spp., two in N Australia and S New Guinea, one in Asia, five in Africa (Senegal to Uganda and Angola), and four in New World, all in South America but two reaching Central America, one up Mexico and Caribbean; three spp. in Brazil, none endemics.

LINEAGE 3: CISTO/DIPTEROCARPOIDS

DIPTEROCARPACEAE

§ FULLY CHLOROTROPHIC

Genera /species 22/c. 550 Distribution SE Colombia, tropical Africa, Madagascar, Seychelles, tropical Asia eastwards to New Guinea, with their highest diversity in the West Malesian lowland rainforests. Habit bisexual, evergreen trees or shrubs. Often with large plank buttresses. Use timber (hardwood), resins (dammar), camphor (Dryobalanops aromatica), butter fats from fruits.

SYSTEMATIC two subfamilies, Dipterocarpoideae (10–11/c 510) only in Seychelles and Asia to New Guinea; among Monotoideae, outsiders are Marquesia (3; tropical Africa) and Monotes (c 30; tropical Africa, Madagascar).

1. Pseudomonotes Lodoño, Alvarez & Morton. Main canopy unbuttressed tree 25-30 m tall, to 70-80 cm diam.; branchlets terete, glabrous; leaves 9-23 cm long, 6-16 cm wide, entire, chartaceous; inflorescence axillary, subcymose, 5-7 cm long; flowers bisexual, actinomorphic; flower bud 5-8 mm long, 2-4 mm wide (flowers only observed in bud); corolla glabrous, greenish white, petals 5, stamens numerous, fruit a dry nut, 3-4 cm long, 1.5-2 cm wide, glabrous, ovoid, pericarp woody; seed 1 per fruit. Only one sp., P. troprenbosii Lodoño, Alvarez & Morton., known only from the type locality in the vicinity of Araracuara, Department of Amazonas, Colombia, at 200-300 m elevation; no open flowers have been seen; the plants have been collected in bud in November and in fruit in April.

CISTACEAE

§ FULLY CHLOROTROPHIC

Genera /species 8/c. 170 Distribution temperate and subtropical regions in the Northern Hemisphere eastwards to Central Asia, few species in Caribbean, Guiana Shield and southern South America. Habit bisexual, evergreen shrubs or suffrutices, or annual herbs, one sp. a tree in South America. Numerous species are xerophytes. Cistaceae grow in lowlands and mountains up to middle altitudes, with a preference of open areas on sandy or chalky soils. The distribution of the family is exclusive to the Northern Hemisphere except for 4 disjunct, endemic species in warm-temperate southern and tepuis in South America. Neotropical Cistaceae are mostly heliophytic shrubs found in Mexico and Mesoamerica at high elevations in exposed areas, dry rock slopes on alkaline or sandy soils and in mixed pine and oak forests.

Key differences from similar families

ü Cochlospermaceae - palmatifid leaves; no leaf hairs; cystoliths absent; lacking spiny stipules; dissected into distinct lobes or sections; non-fleshy; not sclerified or modified; not one-veined or pinnately veined; stipules absent; exudate colour brown, orange, red or yellow; exudate darkens rapidly on exposure. Inflorescence structure racemes; anthers dehise via pores; peduncle present.

ü Rosaceae - leaves sometimes pinnately compound; margins usually serrate; paired stipules; spines may be present on midrib of leaflets and the rachis of compound leaves; flowers arranged in racemes, spikes or heads; bases of sepals, petals and stamens are fused together to form hypanthium.

ü Malvaceae: subfamily Malvoideae - cystoliths absent in leaves; leaf margins can be serrate, dentate or crenate as well as entire; no obvious exudate when cut; bracts not present (although can be misinterpreted as being present); usually 5 calyx & corolla segments (3-5 in Cistaceae); stamens adnate to perianth; ovary locules usually 3-5; usually fewer androecial members (often 10-15); fruit surface often spiny or ridged.

ü Malvaceae: subfamily Tilioideae - usually trees or shrubs; inflorescence axillary (uncommon in Cistaceae); gynoecium 2-100-carpellate; petals sometimes bifid.

ü Malvaceae: subfamily Grewioideae - often trees or shrubs; cystoliths absent in leaves; leaf margins can be serrate, dentate or crenate as well as entire; no obvious exudate when cut; ovary locules 1-10; 4-5 calyx & corolla segments (3-5 in Cistaceae); fruit surface often spiny or ridged.

ü Turneraceae- leaf margin rarely entire; cystoliths absent; 5 calyx & corolla segments (3-5 in Cistaceae); staminodes absent; androecial members 5; no obvious exudates when cut.

SYSTEMATIC two subfamilies, very divergent in habit and habitat, both in South America.

1. SUBFAMILY PAKARAIMAEOIDEAE (1/1) ‣ a single genus.

1. Pakaraimaea Maguire & Ashton. Unbuttressed coppicing smooth-barked main canopy tree or large shrub, making ECM symbioses with fungi. Only one sp., P. dipterocarpacea Maguire & Ashton, in heath forest and savannah edges, white sand soils, of W Guyana and E Venezuela; discovery only in 1977.

2. SUBFAMILY CISTOIDEAE (8/205-210) – outsiders Fumana (15–18; Europe, Mediterranean, North Africa); Lechea (17–18; North America, Mexico, Belize, Cuba); Helianthemum (c 110; Europe, Canary Islands, Mediterranean, North Africa to Central Asia), Tuberaria (c 12; West and Central Europe, Mediterranean), Halimium (12; Mediterranean, North Africa, Türkiye), Cistus (19–20; Canary Islands, Mediterranean, Türkiye, the Caucasus).

2. Crocanthemum Spach. (inc. Helianthemum p.p.) Half-shrubs or shrubs, often appearing herbaceous, aromatic; leaves stipulate, alternate; pinnately veined; flowers chasmogamous and cleistogamous. 19 spp., 18 in North America up to Costa Rica and Hispaniola, and one, C. brasiliense (Lam.) Pers., from S Brazil, Uruguay and NE Argentina.

Good delimination characteristics between Helianthemum and Crocanthemum includeing leaf arrangement, pollen type, shape of the funicle, embryo and base of the style; Helianthemum the only genus with opposite leaves and only neotropical genus which may have stipules (though Crocanthemum has axillary tufts of leaves, sometimes mistaken for stipules); Lechea petals, of which there are 3, unlike the other two neotropical genera (5), are minute, 3 stigmas; Helianthemum style narrowly tapering from apex to a filiform base. Crocanthemum style short and erect.

LINEAGE 4: SUPERMALVIDS

MUNTINGIACEAE

§ FULLY CHLOROTROPHIC

Genera /species 3/3 Distribution Florida, Mexico, Central America, Caribbean, tropical South America and southwards to Argentina. Habit usually bisexual (rarely unisexual), evergreen trees or shrubs. Use Ornamental plants, fruits, fibres, timber (Muntingia).

Key differences from similar families The Muntigiaceae could potentially be confused with members of the family Malvaceae subfamily Tilioideae, but differ from them in the following characters:

ü Absence of mucilage cavaties or canals in stem and leaf anatomy.

ü The combined occurrence of long, simple bristles, stellate hairs, and glandular trichomes with globular heads, is unknown in subfamily Tilioideae.

ü The stipule -dimorphism shown in Muntingia and Dicraspidia is unknown in subfamily Tilioideae.

ü The supra-axillary position of the inflorescences.

ü The occurrence of inferior ovary, which is superior in subfamily Tilioideae.

SYSTEMATIC all genera in South America.

1. Dicraspsidia Standl. Only one species, D. donnell-smithii Standl., found from S Central America to N Colombia.

2. Muntingia L. Tres, white flowers. Only one sp., M. calabura L., distribution extending from Mexico and Caribbean to the Brazilian Amazon rainforest (natively in Amazonas, Acre and Rondonia states) and Bolivia; now frequently cultivated in Africa and Asia as an ornamental and for its edible fruits; one of the few tree species known from both sides of the Andes. In Brazil trees are often planted along riverbanks, where the fruits falling into the water are useful fish attractants. Muntingia provides valuable fuel wood as it ignites quickly, burns with intense heat and makes little smoke. Guatemala, 'Jam tree' in Sri Lanka and 'Jamaican Cherry' amongst many other vernacular names.

3. Neotessmannia Burret. Only one species, N. uniflora Burret, a very rare tree known only from the type collection from swamp forest in one location in E Peru, where fruiting material has yet to be collected.

CYTINACEAE

§ PARASITIC (Prosopanche - Cassytha - Bdallophytum - Krameria - Mitrastemon - ... - OROBANCHACEAE - Cuscuta)

Genera/species 2/11 Distribution Canary Islands, Mediterranean, Türkiye, southern and western Caucasus, southern Africa, Madagascar, southern Mexico, Central America. Habit monoecious or dioecious, achlorophyllous perennial herbaceous endophytic root holoparasites without rhizome or normal roots. Cytinus subgenus Cytinus on species of Cistaceae; Cytinus subgenus Hypolepis on species of at least five different clades of angiosperms; Bdallophytum mostly on Bursera (Burseraceae).

SYSTEMATIC outsider Cytinus (8; subg. Cytinus in Mediterranean, Canary Islands, Türkiye, Caucasia; subg. Hypolepis in southern Africa and Madagascar).

1. Bdallophytum Eichler. Herbs, achlorophylous, holoparasitic and andromonoecious; endophyte vegetative tissue (not visible) developed in shallow roots of plants of the genus. 4 spp., 3 from Mexico, one of them up to Costa Rica, and B. caesareum (Fern.Alonso & H.Cuadros) Byng & Christenh., known only from the Valledupar region in the center of the Department of Cesar, Colombia, in the foothills of the Sierra Nevada de Santa Marta at altitudes below 500 m.

MALVACEAE

§ FULLY CHLOROTROPHIC

Genera /species 246/4,400–4,700 Distribution cosmopolitan except polar areas, with their largest diversity in tropical forests. Habit usually bisexual (rarely monoecious, polygamomonoecious or dioecious), evergreen or deciduous trees, shrubs or suffrutices (rarely lianas), perennial, biennial or annual herbs. Some species are xerophytes. Some genera with tough fibres in bark and stem.

Ornamental plants, textile plants (seed hairs from Gossypium; endocarp hairs, kapok, from Ceiba pentandra, Bombax etc.; phloem fibres from Corchorus capsularis, Abroma augusta, Hibiscus cannabinus, H. sabdariffa, etc.), fruits (aril from Durio zibethinus, etc.), vegetables (Abelmoschus esculentus (L.) Moench etc.), spices, beverages and stimulants (Theobroma cacao, T. grandiflorum, Cola spp., etc.), medicinal plants (Althaea officinalis etc.), seed oils (Gossypium), timber, carpentries (balsa from Ochroma pyramidale etc.), forage plants (Hermannia). Abelmoschus esculentus (L.) Moench (okra) is grown throughout the world as a vegetable, the immature fruits being eaten in a variety of ways. It and A. moschatus Medik. are also used as a source of stem fibers and of seed oil. In addition, the seeds are sometimes used as a coffee substitute, and masticated seeds are used as an antidote for poisonous snake bite. A red-flowered form (A. tuberculatus Pal & Singh?) was recently introduced into U.S. horticultural use as a garden ornamental. 1610 spp. in South America.

SYSTEMATIC nine subfamilies; subfamily Dombeyoideae (21/370–375, tropical regions in the Old World, St. Helena, with their largest diversity in Madagascar) does not occur in South America.

1. SUBFAMILY GREWIOIDEAE (25/680–695) ‣ two tribes, both in South America.

1.1 GREWIOIDEAE ▸ TRIBE APEIBEAE (11/c. 125) - outsiders Ancistrocarpus (3; tropical Africa), Sparrmannia (4; tropical Africa, Madagascar), Entelea (1; New Zealand, Three Kings Islands), Clappertonia (3; tropical West and Central Africa), Glyphaea (2; tropical Africa), Pseudocorchorus (5; Madagascar), Erinocarpus (1; SW India).

1. Apeiba Aubl. Trees, often vast; large leaves, sometimes bullate-reticulate, often domatiiferous; fruit is high aculeate, small yellow flowers, 4 or 5-merous. 11 spp., Mexico to South America (all species), highly centered in northern Amazon rainforest, from Venezuela to French Guiana and Brazil (7, two endemics).

2. Corchorus L. Shrubs, subshrubs, or annual herbs; leaves usually simple. 40-100 spp., pantropical, often ruderal, 18 spp. endemic to Australia, 8 in New World, all in South America, 5 in Brazil, no endemics.

3. Heliocarpus L. Small or medium-sized tree, up to 22m, and trunk up to 40 cm in diameter, with serrate leaves. Only one polymorphic species, H. americana L., Mexico to Paraguay and Caribbean.

4. Triumfetta L. Trees, shrubs or herbs, sometimes annual. c. 204 spp., 50 spp. in Africa, 40 in Asia, 61 in Australia, 54 in over tropical New World, 18 in South America, centered in Mexico (37, 33 endemics) and Brazil (10, 4 endemics, mainly in the savannas of C Brazil (cerrado) and Atlantic Forest), in savannah, woodlands, at forest margins, absent in continental Chile, but present in Isla de Pascua.

1.2 GREWIOIDEAE ▸ TRIBE GREWIEAE (14/c. 570) - outsiders Colona (35–40; China, SE Asia, Malesia), Desplatsia (5; tropical Africa), Duboscia (2; tropical Africa), Microcos (13–15; tropical Asia to Fiji), Grewia (c 320; tropical and subtropical regions of the Old World), Eleutherostylis (1; Moluccas, New Guinea), Tetralix (6; Cuba).

5. Goethalsia Pittier. Tall tree. Only one sp., G. meiantha (Domm. Sm) Burret, from Nicaragua to Boyaca, Colombia and Venezuela.

6. Hydrogaster Kuhlm. Large dioecious tree up to 30 m tall; the trunk stores eater in cavities, with can pour forth upon injury, hence the generic name. Only one sp., H. trinervis Kuhlm, endemic to Atlantic Coasts of Bahia and Espírito Santo states, in E Brazil.

7. Luehea Willd. Trees or shrubs up to 30 m tall, leaves dentate or serrate. 19 spp. from Mexico to Argentina, all in South America, often in deciduous forests or secondary vegetation; 10 spp. in Brazil, 4 endemics.

8. Lueheopsis Burret. Trees, often vast; leaves entire or crenulate. 6 spp., tropical South America, most of primary humid forest; 5 spp. in Brazil, L. burretiana Ducke endemic.

9. Mollia Mart. Trees or shrubs; indimentum of stallate hairs and peltate scales. 17 spp. of Amazon rainforests of northen South America, 14 in Brazil, 8 endemics.

10. Trichospermum Blume. Trees, sometimes vast; leaves entire to serrate or serrulate. 41 spp., 36 in Old World, and three from southern Mexico to Cuba and Peru, three of then in South America.

11. Vasivaea Baill. Small dioecious tree, shrub, or climber; inflorescence terminal. Two spp. from Venezuela, Colombia, Peru and Amazon rainforest of N Brazil (both species, no endemics).

2. SUBFAMILY BYTTNERIOIDEAE (25/700–725) ‣ four tribes, Lasiopetaleae (9/150–155, Madagascar, New Guinea, Australia, Tasmania, New Caledonia) absent in South America.

Some groups are mainly found in humid tropical forest, while others are restricted to drier habitats. Herrania lemniscata (Schomb.) R.E.Schult. the sweet-sour pulp around the seeds is edible, tasting like Theobroma pulp; recent studies of T. cacao L. genetics seem to show the plant originated in the Amazon rainforest and was distributed by humans throughout Central America and Mesoamerica. It seeds are used to make chocolate; the cupuaçu, T. grandiflorum (Willd. ex Spreng.) K. Schum., is a closely related species also grown in Brazil; like the cacao, it is also the source for a kind of chocolate known as cupulate or cupuaçu chocolate; the cupuaçu is considered of high potential by the food and cosmetics industries.

Key differences from similar groups - the families and subfamilies listed below differ from the Malvaceae subfamily Byttnerioideae as follows:

ü Brownlowioideae - sepals fused into a campanulate or urceolate tube; stamens many, free or fasciculate.

ü Bombacoideae - usually stout trees with bottle-shaped and/or trunks armed with prickles; leaves usually palmate/palmately lobed; sepals fused into a tube; petals adnate to androecium; style branched.

ü Cochlospermaceae - anthers dehisce via pores.

ü Grewioideae - petals usually yellow or white, clawed, often with hairy basal nectaries; stamens numerous, free, sometimes grouped in antesepalous fascicles.

ü Helicteroideae - calyx tubular; epicalyx always present; petals clawed; androgynophore usually present; stamens 10-30; ovaries usually apocarpous, except Ungeria and Reevesia.

ü Malvoideae - epicalyx is present in and absent in Neotropical genera; petals adnate to the androecium; stamens fused into a staminal column.

ü Sterculioideae - petals always absent; androgynophore usually present; stamen filaments free; ovaries apocarpous.

2.1 BYTTONERIOIDEAE ▸ TRIBE THEOBROMATEAE (4/43) - outsider Glossostemon (1; Arabian Peninsula, Iraq, Iran).

12. Guazuma Mill. Trees or shrubs, leaves simple, serrate. Three spp., one in Central America, G. crinita Mart. in Brazil, Peru, Bolivia and G. ulmifolia Lam. widely in tropical areas from Mexico, Central and South America.

13. Herrania Goudot. Small trees, al shoots orthotropic, leaves digitately compound, often cauliflorous. 17 spp. in tropical South America (one up to Central America), 4 in Brazil, none endemics, in two sections:

§ sect. Herrania ‣ 3 spp., that have short ligules (less 2 cm) and a patelliform calyx with sepals connate more than half their length, giving a ‘closed’ appearance to the flower; Panamá, Colombia and Venezuela.

§ sect. Subcymbicalyx ‣ long ligules (2.5-18 cm) and has sepals that are free almost to the base, giving the flower an ‘open’ appearance; over range of genus.

14. Theobroma L. Trees with larger fruits with nutlets, often cauliflorous. 20 spp., from south Mexico to Amazon rainforest, higly centered in E Peru and NW Brazil; 11 spp. in Brazil, two endemics; six sections are recognized based on tree architecture, fruit, petal, and androecial characters.

§ sect. Andropetalum ‣ T. mammosum Cuatr. & León, rare, known only Nicaragua and Costa Rica.

§ sect. Glossopetalum ‣ T. angustifolium Moçiño & Sessé, T. canumanense Pires & Fróes, T. chocoense Cuatr., T. cirmolinae Cuatr., T. flaviflorum Aguillar, T. grandiflorum (Willd. ex Spreng.) Schum., T. hylaeum Cuatr., T. nemorale Cuatr., T. obovatum Klotzsch ex Bernoulli, T. simiarum Donn. Smith., T. sinuosum Pavón ex Hubber, T. stipulatum Cuatr., T. subincanum Mart.; over distribution of genus.

§ sect. Oreanthes ‣ 5 spp., T. bernouillii Pittier, T. glaucum Karst., T. speciosum Willd., T. sylvestre Mart., T. velutinum Benoist., Panamá to SE Brazil and Guianas.

§ sect. Rhytidocarpus ‣ only one sp., T. bicolor Humb. & Bonpl., northern South Ameica in N Brazil.

§ sect. Telmatocarpus ‣ two spp., T. gileri Cuatr. and T. microcarpum Mart., from Venezuela and Colombia to C Brazil.

§ sect. Theobroma ‣ T. cacao L., Costa Rica to N Brazil; this spp., is distinctive within the genus and is placed alone in the type section; it has several unusual charactersuch as a thick, fleshy pericarp, at least partly white flowers, and whorls of lateral branches in fives (all other species in the genus have branches in fours); cacao is also unique in having high amounts of theobromine and caffeine in mature seeds but an absence of tetramethyl urate, characters that may have played a role in the species' domestication.

Representative species from all the sections can be found in Brazil, except for Andropetalum.

Cocoa has also been used for an array of medicinal purposes. Unfermented cocoa seeds and the seed coat are used to treat a variety of ailments, including diabetes, digestive and chest complaints. Cocoa powder, prepared from fermented cocoa beans, is used to prevent heart disease. Cocoa butter is taken to lower cholesterol levels, although its efficacy is unclear. It is also used widely in foods and pharmaceutical preparations, as well as being used as a rich moisturiser for the skin. The crushed shells of cocoa beans are used as an alternative to peat mulch. Mulches are layered on to the soil surface to suppress weeds, conserve moisture, improve its visual appearance and minimize erosion. Not only does this make good use of cocoa-shell, which is a by-product of the chocolate industry, but it also helps reduce the use of peat.

2.2 BYTTNERIOIDEAE ▸ TRIBE BYTTNERIEAE (8/260–265) - outsiders Scaphopetalum (12; tropical Africa), Leptonychia (c 25; tropical regions in the Old World), Abroma (1; tropical Asia, NE Queensland), Kleinhovia (1; tropical Asia, NE Queensland), Megatritheca (2; Gabon, Congo).

15. Ayenia L. Mostly less 1m tall and, in especially dry regions, may be low with procumbent or decumbent branches, typically malvaceous palmately-veined and toothed leaves (2 of small trees (4 to 8 m) with a well-defined trunk); fruits spinose. 73 spp. from S North America to Argentina, centered in Mexico to Panamá and Caribbean (38), and S Brazil (17, 10 endemics) and adjacent Bolivia and Paraguay (10), 37 in South America.

16. Byttneria Loefl. Unarmed lianas on other woody plants for support, or shrubs that may be leaning or semi-scandent at some stages of development, often with long curving branches covered in prickles or more rarely spines that are used to scramble over nearby vegetation. 130 spp., pantropical, 88 of which are New World, 82 in South America, and 27 in Madagascar; 52 spp. in Brazil, 22 endemics.

17. Rayleya Cristóbal. Shrubs, leaves simple, serrate, with nectar near base of midvein. Only one sp., R. bahiensis Cristóbal, from central Bahia state, Brazil, in dry scrublands.

2.3 BYTTNERIOIDEAE ▸ TRIBE HERMANIEAE (4/250–265) - outsiders Hermannia (c 120; tropical and subtropical regions on both hemispheres, with their largest diversity in the Cape Provinces in South Africa), Dicarpidium (1; NW Australia).

18. Melochia L. Shrubs, more rarely herbs or trees. 54 spp., mostly neotropical, a few in the Palaeotropics - some widely distributed weeds (e.g., Chocolate weed, M. corchorifolia L); 52 spp. in New World in five sections, 41 in South America, 22 in Brazil, 11 endemics:

§ sect. Melochia ▸ 11 spp., has 8 only in New World, two anfi-atlantic, and one endemic to Africa.

§ sect. Physodium ▸ 3 spp., endemic to Mexico.

§ sect. Visenia ▸ 12 spp., Asia and the eastern Pacific.

§ sect. Mougeotia ▸ 18 spp., Mexico to Uruguay and northern Argentina.

§ sect. Pyramis ▸ 14 spp., only New World except the pantropical M. pyramidata L., known also in Mascarene Islands and Malesia to northern Australia.

19. Waltheria L. Shrubs or herbs, sometimes with xylopodium, leaves simple, serrate. 64 spp., six spp. in Africa, Madagascar, Pacific Islands, Asia, and Australia, W. indica L. native to the New World, but a pantropical weed, and 57 exclusively in New World, 40-43 in South America, mostly neotropical with centres of diversity in Mexico (16, 11 endemics) and Brazil (34, 24 endemics).

3. SUBFAMILY HELICTEROIDEAE (12/135–140) ‣ two tribes; Durioneae (6/c 53, India, Sri Lanka, Burma, Malesia) does not occur in South America; among Helictereae, outsiders are Reevesia (c 15; tropical Asia, Central America), Ungeria (1; Norfolk Island), Neoregnellia (1; Cuba, Hispaniola), Mansonia (2; tropical Africa, Assam, Burma), Triplochiton (2; tropical Africa).

20. Helicteres L. Shrubs or small trees. 60 spp. found in the tropics of South America, Asia and Australia, but is absent from Africa; 38 spp. in New World, 33 in South America, 31 spp. in Brazil, 23 endemics.

4. SUBFAMILY TILIOIDEAE (3/37) ‣ outsiders are Tilia (c 45; temperate regions on the Northern Hemisphere), Craigia (2; China, N Vietnam).

21. Mortoniodendron Standl. & Steyerm. Shrubs or small to vast trees, sometimes buttressed; leaves simple, margin entire. 15 spp. from southern Mexico to Panamá, highly centered in Central America, one of then up to NW Colombia in South America.

5. SUBFAMILY BROWNLOWIOIDEAE (9/c 80) ‣ outsiders are Diplodiscus (12; Sri Lanka, West Malesia to Philippines), Indagator (1; Queensland), Brownlowia (c 30; SE Asia, Malesia to Solomon Islands), Pentace (c 25; Burma, SE Asia, West Malesia), Pityranthe (2; Sri Lanka, S China inc. Taiwan), Jarandersonia (4; Borneo), Berrya (6; India, Sri Lanka, SE Asia, Malesia), Carpodiptera (5; tropical E Africa, the Comoros, S Mexico, Caribbean, Trinidad).

22. Christiana A.DC. Trees or shrubs up to 45-50 m tall, with stellate indumentum, often dioecious by abortion; functionally dioecious; leaves alternate, large, palmately nerved from the base; stipules falling; flowers functionally unisexual in pedunculate terminal and lateral branched corymbiform cymes; calyx campanulate, 3–5-lobed; petals 5(–7). 5 spp., one in Tahiti (extinct), two from Ecuador, Peru up to Guianas (one also in Amazonas and Pará states in N Brazil), C. macrodon Toledo endemic to SE Brazil and C. africana DC., ranging from Madagascar through tropical Africa, disjunct (possibly due to human active) of Central America, Venezuela, Guianas and N Brazil.

6. SUBFAMILY STERCULIOIDEAE (13/425–575) ‣ outsiders are Brachychiton (31; New Guinea, Australia), Cola (c 120; tropical Africa), Octolobus (3; tropical Africa), Acropogon (c 25; New Caledonia), Franciscodendron (1; NE Queensland), Argyrodendron (4; Queensland, New South Wales), Firmiana (16; tropical regions in the Old World from tropical E Africa and eastwards), Hildegardia (7; tropical Africa, Madagascar, tropical Asia, Northern Territory, Cuba), Scaphium (8; tropical Asia), Pterocymbium (c 15; SE Asia, Malesia to Fiji), Heritiera (c 35; tropical Africa, tropical Asia, Queensland, New South Wales, New Caledonia).

23. Pterygota Schott. & Endl. Tall trees with straight, smooth trunk; young branches minutely stellate-puberulous; leaves entire or palmatilobed, 5–9-nerved from base, petiolate; flowers unisexual, in axillary panicles. 11 spp., Central Africa (3), SE Asia (4), and four in New World, three in two countries range (Costa Rica and Panamá, Colombia and Venezuela, Peru and Bolivia) and P. brasiliensis Allemão, native to the Atlantic coastal forest of Brazil from Pernambuco to São Paulo states.

24. Sterculia L. Trees, rarely lepidote; leaves simple to digitely compound. 300 spp., pantropical, 34 in New World, 29 in South America, 14 in Brazil, 6 endemics.

7. SUBFAMILY MALVOIDEAE (110/1.845–1.850) ‣ eight lineages and two unplaced genera in South America; three clades are ex-Bombacaceae and are also most basal in this subfamily Fremontodendreae (2/2, California to Guatemala), unplaced genera Howittia (1; SE Queensland, New South Wales, Victoria), Camptostemon (3; coasts in Central Malesia, New Guinea and N Australia) and Lagunaria (1; Queensland, Norfolk Island, Lord Howe), and Alyogyne clade (1/5, Western Australia, South Australia, southern Northern Territory) do not occur in South America.

Key differences from similar families - The families and subfamilies listed below differ from the Malvaceae subfamily Malvoideae as follows:

ü Brownlowioideae - sepals fused into a campanulate or urceolate tube; an androgynophore is always absent; stamens are many, free or fasciculate.

ü Bombacoideae - are usually stout trees with bottle-shaped and/or trunks armed with prickles.

ü Byttnerioideae - an epicalyx is always absent; the petals are free from the androecium; the stamens are free.

ü Cochlospermaceae - the anthers dehisce via pores.

ü Grewioideae - an epicalyx is always absent; the petals are usually yellow or white; the stamens are free, sometimes fasciculate.

ü Helicteroideae - an epicalyx is always present; stamens 10-30.

ü Sterculioideae - petals are always absent; an androgynophore is usually present; the stamen filaments are free; the ovaries are apocarpous.

ü Turneraceae - stipules and epicalyx are always absent; two glands are often present at the leaf base; the stamens are free.

MALVOIDEAE ▸ UNPLACED GENERA

25. Pentaplaris L. O. Wiliams & Standll. Tall, buttressed trees; leaves simple, entire. Three spp., from Costa Rica to Ecuador, disjuntc in Bolivia.

26. Uladendron Mac.-Berti. Tall tree; leaves lightly lobed, serrate. Only one sp., U. codesuri Marc-Berti, endemic to the rainforests of central Guiana Shield in Venezuela, 100-200 m elevation range.

7.1 MALVOIDEAE ▸ OCHROMEAE CLADE (2/5) - both genera in South America.

27. Ochroma Sw. Trees up to 30 m tall; leaves simple to lobed. Only one sp., O. pyramidale (Cav. ex Lam.) Urb., from tropical America, widely distributed in clearings and secondary forests; furnishing the extremely light balsa wood, one of the world's softest and lightest woods with a specific gravity of only 0.19.

28. Patinoa Cuatrec. Trees with verticillate branches; leaves simple, with fruits edible. 4 spp. from Panamá through Colombia (two endemics) to Brazil (two spp., no endemics) and Peru.

7.2 MALVOIDEAE ▸ SEPTOTHECA CLADE (1/1) - a single genus.

29. Septotheca Ulbr. Tall tree, lepidote; inflorescences are axillary, long-pedunculate and subumbellate. Only one sp., S. tessmannnii Ulbr., from Colombia, Ecuador, Peru and Amazonas state in Brazil.

7.3 MALVOIDEAE ▸ TRIBE MATISIEAE (3/104)

30. Matisia Bonpl. Trees, with 3-verticilate branches. 56 spp., in tropical South America, 4 spp. up to Central America, some with edible fruits; 7 spp. in Brazil (only Amazon rainforest), none endemic.

31. Phragmotheca Cuatrec. Tree, rarely lepidote; leaves simple, ovate to cordate. 11 spp. from Colombia (center of diversity) to Peru; some fuits are edible.

32. Quararibea Aubl. Trees up to 30 m tall, with 4-5 verticillate branches, occasionally lepidote, leaves unifoliate, flowers small (2.0–6.4 cm in length), solitary, opposite the leaves or axillary, ramiflorous, pendulous or erect, usually cauliflorous. 42 spp., Neotropical, some with edible fruit; 34 in South America, 13 in Brazil, 7 endemics.

7.4 MALVOIDEAE ▸ TRIBE MALVEAE (71/1.210–1.220) - outsiders Meximalva (2; Mexico), Neobrittonia (1; Mexico to Panamá), Robinsonella (16; Central America), Fryxellia (1; SW U.S.A., NW Mexico), Billieturnera (1; S Texas, NE Mexico), Allowissadula (10; Texas, Mexico), Horsfordia (4; SW U.S.A., NW Mexico), Periptera (5; W Mexico, Guatemala), Phymosia (8; S Mexico, Central America, Caribbean), Malacothamnus (16; California, NW Mexico), Iliamna (8; SW Canada, U.S.A.), Napaea (1; E to C U.S.A.), Eremalche (3; California, NW Mexico), Kitaibela (1; Lower Danube in W Balkan Peninsula), Malope (2; Mediterranean), Anisodontea (20; South Africa, Lesotho), Alcea (75–80; Mediterranean to C Asia), Althaea (17; Europe, Mediterranean, temperate Asia to NE Siberia), Malva (c 30; Europe, Macaronesia, Mediterranean, tropical African mountains, temperate and Central Asia, NW Himalayas, S Australia, Tasmania, America), Callirhoe (9; S Canada, U.S.A., Mexico), Sidalcea (c 30; SW Canada, W U.S.A., NW Mexico), Hoheria (6; New Zealand), Lawrencia (5; Australia, Tasmania), Plagianthus (3; New Zealand), Gynatrix (2; SE New South Wales, Victoria, Tasmania), Asterotrichion (1; Tasmania).

33. Abutilon Mill. Subshrubs or shrubs to small trees, glabrescent or pubescent, sometimes glandular-pubescent; flowers solitary in the leaf axils or aggregated into racemes or panicles, less commonly in umbels; corolla often yellow or orange, less often white, lavender, rose, red, or purplish. 160 species in all continents, mostly neotropical, 82 spp. in New World, 43 in South America, 20 in Brazil, 14 endemics.

34. Acaulimalva Krapov. Acaulescent perennial herbs from a woody caudex; leaves petiolate, the blades suborbicular or ovate, basally cuneate; flowers axillary, long- or shortpedicellate; involucel of 2 or 3 lanceolate or filiform bracts; calyx campanulate, 5-lobed; corolla usually exceeding the calyx, white, lavender, purplish (sometimes with a dark center), or sometimes yellow. 19 spp., in the high Andes (2,900–4,800 m), from Venezuela and Colombia to Bolivia and NW Argentina, highly diverse in Peru (12 endemics).

35. Akrosida Fryxell & Fuertes. Small trees or shrubs minutely stellate-pubescent; leaves discoulored; flowers in axillary fascicles, the pedicels slender, tomentulose, articulated in upper third; petals asymmetrically emarginate, white to bluish. Two spp., notably highy disjunct, one in mountains of Dominican Republic and bluish flowered A. macrophylla (Ulbr.) Fryxell & Fuerte known only from the vicinity of Nova Friburgo in NE of Rio de Janeiro state, Brazil.

36. Allobriquetia Bovini. (off Briquetia) Subshrub or shrubs, erect; branches erect, rarely terete; leaves peciolate below, and amplexicaul bellow the inflorescence; synflorescence frondose-bracteate, pyramidal, lax or spiciform raceme; flowers pedicellate; calyx campanulate, corolla yellow; ovary 5-13 celled, 2-3 ovules per cell. Three spp., two from NW Mexico, and A. spicata (Kunth) Bovini from Mexico to Argentina and Brazil.

37. Allosidastrum (Hochr.) Krapov., Fryxell & D.M. Bates ex Fryxell. Shrubs 1-5 m tall with pubescence of both stellate and simple hairs, sometimes viscid; leaves long-petiolate; flowers in terminal, more or less leafless panicles or racemes; petals white to yellow, with or without a dark basal spot. 4 spp. from Mexico and Caribbean to South America (3) as far as Bolivia, Venezuela, Brazil (Mato Grosso) and Suriname.

38. Andeimalva J.A. Tate. 5 spp., all national endemics in Chile (1), Peru (3) and Bolivia (1).

39. Anoda Cav. Annual or perennial herbs or subshrubs, erect or decumbent, hispid or stellate-pubescent or -puberulent to glabrate; leaves petiolate, the blades linear. Flowers solitary in the leaf axils or aggregated in open apical racemes or panicles; petals spreading, yellow, white, lavender, or purplish. 24 spp., mostly Mexican, a few in the U.S.A., A. cristata (L.) Schltdl from Mexico to Argentina and Caribbean, also Venezuela, and A. hastata Cav. endemic to Cono Sur.

40. Bakeridesia Hochr. Shrubs or small trees 1-8 m tall, densely stellate-pubescent, often ferruginous; flowers solitary or paired in the leaf axils or aggregated into axillary or terminal inflorescences; involucel absent; petals usually large (1.5-6 cm long) and showy, white, yellow, or yellow-orange, often with a dark reddish spot at base. 28 spp. from Mexico and N Central America, with B. integerrima (Hook.) D. M. Bates with disjunct populations in Venezuela, Colombia and Ecuador.

41. Bastardia Kunth. Herbs or shrubs 0.5-3 m tall, pubescente with stellate, simple, and often glandular hairs; flowers solitary or paired in the leaf axils, often aggregated into terminal leafy panicles; involucel absent; calyx divided almost to the base; petals relatively small, yellow. Three spp., widely distributed in the Neotropics, two in Brazil, the endemic B. elegans K. Schum. and the widely distributed B. bivalvis (Cav.) Kunth ex Griseb.

42. Bastardiopsis (K. Schum.) Hassl. Trees or large shrubs, the (young) herbage stellate-pubescent, the hairs very small with many radii; inflorescences axillary or terminal, the flowers glomerulate or paniculate, subsessile or short-pedicellate; calyx basally rounded, ca. half-divided; petals yellow. 6 spp., one in Caribbean, three spp. in Colombia, Venezuela, Peru and Ecuador, B. grewiifolia (Ulbr.) Fuertes & Fryxell endemic to Brazil, and B. densiflora (Hook. & Arn.) Hassl. in Brazil and Cono Sur.

43. Batesimalva Fryxell. Shrubs 1-2 m tall, stellate-pubescent; flowers solitary or clustered in the leaf axils; pedicels slender and elongate; involucel absent; calyx ecostulate, often brownish at least at base; petals bluish lavender, white, or orange (the flowers sometimes cleistogamous). 4 spp., three in North America and Mexico, and B. killipii Krapov. ex Fryxell in Venezuela; therefore, the generic limits of Batesimalva need to be reconsidered, and the generic placement of the venezuelan species needs to be reviewed.

44. Bordasia Krapov. Only one sp., B. bicornis Krapov., endemic to Paraguay.

45. Briquetia Hochr. (exc. Allobriquetia) Herbs or subshrubs 0.5-1.5 m tall, usually with a single erect stem, branching only in the inflorescence, more or less soft-pubescent with stellate and simple hairs. calyx 5-lobed, small; petals yellow. Only one sp., B. denudata (Nees & Mart.) Chodat & Hassl., in Argentina, Paraguay, and Brazil (Bahia, Mato Grosso do Sul, Minas Gerais and Paraná).

46. Callianthe Donnel. Shrubs or treelets 1–8 m. tall; leaves simple, to 22 cm long and 19 cm wide; flowers solitary, paired or in clusters of 3 flowers, axillary, on pedicels to 10 cm long; corolla rotate or campanulate, exceeding calyx in length, the petals obovate and often clawed at the base, 1.5– 5.5 cm long, 0.5–5 cm wide; red, white, cream, yellow, purple, lavender, pink, or burgundy, usually prominently veined, the veins sometimes of another color; fruits schizocarpic, to 5 cm in diameter. 49 spp., some from Mexico to Colombia and Venezuela (41 in South America), and from Ecuador to Bolivia (7) and S Brazil (36, 27 endemics).

47. Calyculogygas Krapov. Erect well-branched shrubs, with minute stellate pubescence; flowers solitary in the leaf axils, slender, the pedicels subequal to the subtending leaf; calyx deeply 5-parted; petals red (drying purplish). Two spp., C. uruguayensis Krapov in grasslands, shrublands and rocky formations in the Pampa biome in Brazil and Uruguay, and with two collections from the Brazilian Atlantic Rainforest biome in Rio Grande do Sul and Santa Catarina states, and C. serrana Grings., endemic to the highland slopes of ‘Serra Geral’ highs, in the Brazilian state of Rio Grande do Sul.

48. Calyptraemalva Krapov. Shrubs, the stems reddish, minutely stellate-pubescent distally, the hairs ferruginous, glabrescent; flowers solitary in the leaf axils, the pedicel to 5.5 cm long, subequal to subtending leaf, stellate-pubescent, with a small, medial (deciduous) bract; corolla roselavender, 4 cm long, externally glabrous, stellate-pubescent and hirsute on claw. Only one sp., C. catharinensis Krapov., endemic to Santa Catarina state, S Brazil.

49. Corynabutilon (K. Schum.) Kearney. Medium to large shrubs or even small trees, stellate-pubescent or sublepidote; flowers solitary (or paired) in the leaf axils or in subcorymbose or subumbeUate axillary inflorescences; involucel absent; calyx campanulate, ca. half-divided to subtruncate, the lobes entire; corolla twice the length of calyx or more, usually lavender or purplish to almost white. 7 spp. from temperate Chile and Argentina.

50. Cristaria Cav. Annual or perennial herbs, erect or procumbent, rarely subshrubs, sparsely to densely stellate-pubescent, sometimes with glandular hairs; flowers solitary (or paired) in the leaf axils or aggregated apically in racemes or panicles; involucel absent; calyx campanulate, ca. half-divided or more; corolla ca. twice the length of calyx (sometimes smaller), white, lavender, or purple, sometimes with a dark center. 19 spp., two from Peru and remaining 17 in Chile, two up to W Argentina and most confined to the Chilean Atacama desert; mostly coastal at low elevation, but some species occurring at high elevation.

51. Dendrosida Fryxell. Shrubs or small trees 3-10 m tall, stellate-pubescent or - pubernlent to glabrate; flowers solitary in the leaf axils or in fewflowered axillary umbels, sometimes apically congested; involucel absent; calyx prominently to obscurely 10-ribbed and - angled, ca. half-divided; petals yellow or red. 6 spp., four from S Mexico (Michoacan to Chiapas) and two endemics to N Colombia.

52. Dirhamphis Krapov. Subshrubs with stellate pubescence. Leaves petiolate (except sometimes the uppermost sessile), the blades ovate, cordate, crenate-dentate, acute or acuminate, markedly discolorous; stipules filiform; flowers solitary in the leaf axils; involucel absent; calyx prominently 5-lobed; petals 8-20 mm long, yellowish; androecium included, pallid or yellowish. Two spp., one from W Mexico and D. balansae Krapov. from Bolivia and Paraguay.

53. Fuertesimalva Fryxell. (inc. Urocarpidium p.p.) Annual or perennial herbs, ascending or erect, with stellate pubescence; leaves petiolate, the blades ovate or orbicular, usually palmately lobed or parted; flowers sometimes solitary, usually in axillary scorpioid cymes; involucel of 3 filiform bractlets; calyx stellate-pubescent, 5-lobed; corolla purplish (sometimes white), shorter than to slightly longer than the calyx. 14 spp. from Venezuela to Chile and Argentina two disjunct species in Mexico, mainly in montane environments in Andes.

The type species of Urocarpidium, U. albiflorum Ulbr. is more appropriately placed in Tarasa, thus reducing Urocarpidium to Tarasa; the remaining species do not belong with Tarasa; a new genus, Fuertesimalva, is erected to hold them. F. limensis (L.) Fryxell is the type species.

54. Gaya Kunth. Erect herbs or subshrubs, sparingly branched, minutely puberulent, sometimes also pilose; leaves sometimes distichous, petiolate, flowers solitary in the leaf axils, sometimes nodding; involucel absent; calyx small, basally rounded, more than half-divided, not prominently nerved; petals usually yellow. 38 spp., from Mexico, Caribbean Islands, the U.S.A., and over tropical South America (37), with 14 spp. in Brazil (8 endemics).

55. Herissantia Medik. Herbs, subshrubs, or shrubs, erect or decumbent, pubescent or hirsute, sometimes viscid; flowers often solitary in the leaf axils or borne on several-flowered peduncles; involucel absent; calyx lobes lanceolate or ovate; petals white. 5 spp., H. crispa (L.) Brizicky pantropical, one endemic to Mexico and three remaining in Argentina, Uruguay and Brazil (all, one of them endemic).

56. Hochreutinera Krapov. Erect subshrubs with scattered to dense stellate and simple pubescence; flowers solitary in the leaf axils or in terminal racemes or lax panicles; involucel absent; calyx pubescent, half-divided (or sometimes more deeply divided), the lobes more or less cordate, acuminate, more or less accrescent; petals yellow, equaling the calyx to 3 times as long as calyx. Two spp., one from Mexico and Central America, and H. hasslerana (Hochr.) Krapov. from Paraguay, Brazil and Argentina;

57. Kearnemalvastrum D.M. Bates. Erect subshrubs, stellatepubescent; flowers in congested or lax cymes, aggregated into paniculate inflorescences; involucel of 3 inconspicuous linear bractlets; calyx 5-lobed; petals white, equalling or slightly exceeding the calyx. Two spp. from Mexico to Costa Rica, one up to N Colombia.

58. Krapovickasia Fryxell. Procumbent (but not repent), decumbent, or erect perennial herbs, sometimes with xylopodium, with simple hairs and fine stellate pubescence; flowers solitary or fasciculate in the leaf axils; petals yellowish (fading rose). 4 spp., K. physaloides (C. Presl) Fryxell disjunct in southern Texas/NE Mexico and Peru, and three remaining in Brazil (all, no endemics), Paraguay, Bolivia, Argentina and Uruguay.

59. Lecanophora Speg. Perennial herbs, erect or decumbent, sometimes rosette-forming, or subshrubs to 0.5 m tall, densely stellate-pubescent to glabrate; flowers solitary in the leaf axils, long-pedicellate, sometimes aggregated into terminal racemes or panicles; involucel absent; calyx campanulate, ca. half-divided or more; corolla 2-3 times the length of calyx, lavender. 7 spp., mainly Argentina, and one also in Chile;

60. Malvastrum A. Gray. Perennial shrubs or subshrubs (sometimes annual), erect, with patent or appressed stellate pubescence; flowers solitary in the leaf axils or aggregated in apical spikes or racemes; involucel of 3 filiforrn or spatulate bractlets; calyx 5-lobed; corolla yellow or orangish, rarely with a red center. 22 spp. from North, Central and South America (14), some in Australia, and one species adventive elsewhere; 4 spp. in Brazil, three widely distributed and one endemic.

61. Malvella Jaub. & Spach. Prostrate perennial herbs with stellate and lepidote pubescence; flowers solitary in the leaf axils, long-pedicellate; involucel present or absent; calyx lobes ovate or cordate; petals pale yellow fading pale rose. 4 spp., three in western U.S.A. and Mexico, M. leprosa (Ortega) Krapov. also in Peru to Uruguay, and one disjunct in Mediterranean.

62. Modiola Moench. Trailing herbs, usually with little pubescence; flowers solitary in the leaf axils, the pedicels usually shorter than the subtending petioles, more or less pubescent; involucellar bracts 3, distinct, lanceolate; calyx 5-7 mm long, with simple hairs 1-2 mm long; corolla dark orange, often with a red center, 6-8 mm long. Only one sp., M. caroliniana (L.) G. Don, pantropical and subtropical (inc. Central, South America and Caribbean), extending into temperate climates; trailing herbs, usually with little pubescence.

63. Modiolastrum K. Schum. Perennial herbs, decumbent or suberect, the stems more or less hirsute to subglabrous; flowers solitary in the leaf axils, longpedicellate (sometimes exceeding the subtending leaf); involucel trimerous, the bracts lanceolate to ovate, subequal to calyx or somewhat shorter; calyx 5-lobed, deeply divided; corolla 1.5-2 times the length of calyx, pale yellow to rose or purple. 4 spp. in Paraguay, Bolivia, Argentina, Uruguay and Brazil (only M. malvifolium (Griseb.) K. Schum., no endemic).

64. Monteiroa Krapov. Shrubs or subshrubs, glabrous or pubescent; flowers sometimes solitary, but usually in few-flowered axillary inflorescences, these cymose or racemose, sometimes subpaniculate, or subumbellate; involucel trimerous, the bracts ovate-lanceolate; calyx campanulate, 5-lobed; petals exceeding the calyx, sometimes auriculate, apically notched. 11 spp., all restricted of S Brazil, except by M. glomerata (Hook. & Arn.) Krapov. in adjacent northern Argentina.

65. Neobaclea Hochr. Small shrubs, minutely stellate-pubescent; flowers solitary in the leaf axils, the pedicels exceeding the corresponding petiole; involucel absent; calyx 5-lobed, the lobes undulate-crenate (like the leaves) or pinnately divided; petals lavender or pink, exceeding the calyx. Only one sp., N. crispifolia (Cav.) Krapov., endemic to temperate Argentina.

66. Nototriche Turcz. Acaulescent perennial herbs (rarely annuals), usually forming a compact cushion of crowded leaves, the underground stems usually branched, usually covered with persisting leaf sheaths; flowers sometimes subsessile and crowded among the leaves, sometimes long-pedicellate and overtopping the leaves (true epiphylly). 110 spp., Andean of Ecuador, Peru (38 endemics), Bolivia, Chile and Argentina, usually at altitudes above 4,000 m.

67. Palaua Cav. Annual or perennial herbs, erect or decumbent, usually pubescent, the perennial species often forming a rosette of large leaves at ground level, from which long flowering shoots (sometimes procumbent) arise with heteromorphic, smaller leaves; flowers solitary in the leaf axils, sometimes aggregated terminally in a leafy raceme; calyx 5-lobed (except deeply 3-parted in P. trisepala Hochr.); petals subequal to calyx and inconspicuous (especially in annual species) to large and showy, white, pink, or rose-lavender, often drying purplish. 16 spp. in Pacific coast Peru and Chile; 12 endemics to Peru, two in both Peru and Chile, and two endemics to Chile.

68. Pseudabutilon R.E. Fr. Shrubs or subshrubs, the stems usually densely stellate-pubescent (sometimes glabrescent), sometimes also with long simple hairs; flowers solitary or glomerate in the axils, or borne in small axillary umbels, sometimes aggregated into terminal inflorescences; petals yellow (rarely white), 4-15 mm long. 18 spp., U.S.A. to Argentina, 14 in South America, six in Brazil, two endemics.

69. Rhynchosida Fryxell. Perennial herbs or shrubs, procumbent, ascending, or erect; flowers solitary in the leaf axils; pedicels slender; involucel absent; calyx deeply 5-lobed, accrescent and inflated in fruit, the lobes cordate-ovate, overlapping, apiculate; petals yellowish, with or without a purple spot at the base, slightly exceeding to twice as long as the calyx. Two spp., R. physocalyx (A. Gray) Fryxell in Texas, U.S.A., northern Mexico, disjunct in Bolivia and Argentina, and Rio Grande do Sul state in southern Brazil; R. kearneyi Fryxell endemic to Bolivia.

70. Sida L. Perennial herbs or subshrubs, erect or prostrate, glabrous or pubescent, sometimes viscid; leaves petiolate to subsessile, the blades ovate (sometimes lobed), elliptic, rhombic, or linear, usually dentate; flowers solitary in the leaf axils, in axillary glomerules, or in dense or open terminal inflorescences; pedicels shorter than to much longer than the calyces; involucel absent; calyx 5-lobed, often 10-ribbed at the base and plicate in bud; corolla white, yellow, orangeish, rose, or purplish, sometimes with a dark red center. 200 or more spp., pantropical, extending into temperate zones worldwide; 187 spp. in New World, 157 in South America, 101 in Brazil, 52 endemics; currently 12 sections:

§ sect. Cordifoliae ‣ 24 spp., North to South America, two in Australia and S. cordifolia L. pantropical.

§ sect. Distichifoliae ‣ 16 spp., tropics.

§ sect. Ellipticifoliae ‣ nine spp., U.S.A., Mexico and Guatemala.

§ sect. Hookerianae ‣ two spp., one in Australia another from Ethiopia to South Africa.

§ sect. Malacroideae ‣ 24 spp., North to South America, one in Old World.

§ sect. Muticae ‣ only one sp., S. aggregata C. Presl. from Mexico to northern South America and Caribbean, inc. Brazil.

§ sect. Nelavagae ‣ 30 spp., North America to South America up Brazil, and two in Asia.

§ sect. Pseudo-Napaeae ‣ a single species in North America.

§ sect. Oligandrae ‣ four spp., high elevations of Ecuador, Peru and Bolivia.

§ sect. Sidae ‣ 37 spp., over widely in New and Old World tropics.

§ sect. Spinosae ‣ 18 spp., widely in New World.

§ sect. Stenindae ‣ two spp., one endemic to Paraguay and S. linifolia Juss. ex Cav. very widely, also in Brazil.

71. Sidasodes Fryxell & Fuertes. Shrubs ca. 3 m tall, densely pubescent with yellowish stellate (often stipitate) hairs; inflorescences axillary or terminal with subcorymbose branching, the flowers subsessile and congested at the ends of the branches; involucel absent; calyx deeply divided, densely pubescent externally, glabrous internally; petals ca. 1 cm long, white or lavender, bearded on claw. Two spp. from Colombia, Ecuador, and Peru.

72. Sidastrum Baker f. Erect subshrubs 1-2 m tall, more or less stellate-pubescent; flowers solitary in the leaf axils, in short axillary racemes, or forming ample terminal panicles or racemes; pedicels long or short, sometimes capillary; involucel absent; calyx small, cupuliform, ecostate, 5-lobed; petals small, white, yellow, or orange (less commonly rose or purple). 8 spp., Mexico and Caribbean to Argentina; 4 spp. in South America, all in Brazil but widely distributeds.

73. Sphaeralcea A.St.-Hil. Herbs or shrubs, erect (or less often procumbent), usually perennial, stellate-pubescent, frequently with numerous stems; inflorescences racemose, paniculate, or thyrsoid; pedicels usually shorter than the calyces; involucel of 3 linear bracts; calyx 5-lobed, usually pubescent; corolla about twice the length of the calyx, yellow, orange, rose, or purple (rarely whitish). 53 sp., 6 in S Africa, 33 from S Canada to N Mexico, and 14 in Paraguay, Bolivia, Uruguay, Argentina, and Chile, mainly in arid habitats of temperate climates, extending to tropical zones primarily at higher elevations; in Brazil occur only S. bonariensis (Cav.) Griseb, known only in Bage municipality, Rio Grande do Sul state, in two small populations.

74. Spirabutilon Krapov. Shrubs to subshrubs, hirsute, flowers yellow-greenish, 1-2, axillary, pendulous; fruit schizocarpic. Only one sp., S. citrinum Krapov., from E Espírito Santo state, Brazil.

Spiralate anthers is very rare character in plants, known only in fully genera Centaurium J.Hill., Chironia L. (Gentianaceae), Merremia Denst. ex Endl. (Convolvulaceae, some spp. possibly in Distimake Raf.), Spiranthera A. St-Hill. (Rutaceae), Spirotheca Ulbr. and the present genera (Malvaceae), and in Attalea speciosa Mart. ex Spreng. (Arecaceae).

75. Tarasa Phil. Annual herbs or subshrubs, decumbent, ascending, or erect, stellate pubescent; leaves petiolate, the blades triangular, rhomboid, or suborbicular, simple, lobed, or parted; flowers usually in scorpioid cymes; involucel of (2)3 usually filiform bracts; calyx more or less accrescent, 5-lobed, stellate-pubescent, the hairs often stipitate (the stipe purplish); corolla purple or blue (rarely white), the petals sometimes bi-auriculate at base. 29 spp., 27 confined to Peru to Argentina and Chile, one endemic to Mexico, and one in both regions.

Tarasa humilis (Gillies ex Hook. & Arn.) Krapov., the only species in the genus with an acaulescent habit and solitary flowers, was suggested to be a link between either the perennial species of Nototriche or the genus Acaulimalva.

76. Tetrasida Ulbr. Shrubs or small trees 1-5 m tall, with minute stellate puberulence; leaves petiolate, the blades ovate to elliptic, subcordate, entire or serrulate, usually acute, stellate-puberulent above and beneath; stipules subulate, inconspicuous, caduceus; flowers in terminal or axillary panicles, the pedicels 0.5-2 cm long; involucel absent; calyx campanulate, ecostate, halfdivided, small; petals exceeding calyx, yellow. Three spp., from Bolivia and Peru.

77. Tropidococcus Krapov. Herbs 1-2 m tall, annual, very branched, pale flowers, solitary. Only one sp., T. pinnatipartitus (A.St.-Hil. & Naudin) Krapov. endemic to SE Rio Grande do Sul state, S Brazil, in low fields, wet places.

78. Urocarpidium Ulbr. (exc. Fuertesimalva p.p.). Two spp., U. albiflorum Ulbr. from xeric areas of western Peru and U. mathewsii (Turcz.) Krapov. from Cono Sur.

79. Wissadula Medik. Medik. Herbs or subshrubs, usually erect, sometimes with xylopodium, stellate-pubescent or sometimes glabrate, sometimes with long simple hairs; leaves petiolate, the blades broadly ovate to narrowly triangular, entire or crenate- dentate; flowers sometimes solitary in the leaf axils, usually in condensed or open terminal panicles; involucel absent; calyx 5-lobed, usually small; petals usually yellowish and small, sometimes white. 42 spp., 40 in New World, in Texas, U.S.A. to Argentina, 39 in South America, 1 in Africa and Asia each, in two sections:

§ sect. Wissadula ‣ 37 spp., 18 of then in Brazil, 4 endemics.

§ sect. Wissada ‣ three spp., two from N South America to Caribbean (both absent in Brazil), and W. stipulata Bovini endemic to Mato Grosso do Sul state.

80. Woodianthus Krapov. Erect shrub up to 2 m tall; stems yellowish, pinkish flowers; urticating hairs. Only one sp., W. sotoi Krapov., endemic to Santa Cruz, Bolivia.

7.5 MALVOIDEAE ▸ TRIBE GOSSYPIEAE (9/125–130) - outsiders Cephalohibiscus (1; New Guinea), Cienfuegosia (c 30; tropical Africa, tropical and subtropical America), Lebronnecia (1; Marquesas Islands), Thepparatia (1; Thailand), Gossypioides (2; tropical Africa, Madagascar), Kokia (3; Hawaii)

81. Cienfuegosia Cav. Perennial herbs or subshrubs (sometimes shrubs) with procumbent, ascending, or erect stems, glabrous or pubescent. Leaves petiolate, the blades linear, elliptic, digitately divided, or reniform, entire or serrate, pubescent to glabrous, with or (in American species) without adaxial foliar nectaries; flowers solitary or sympodial in the leaf axils; corolla yellow (sometimes nearly white), pink, or purple, with or without a dark basal spot. 25 spp., with subg. Cienfuegosia in Africa and subg. Articulata in the New World with 21 spp. from Florida and Texas to the Caribbean Islands, and south to N Argentina; 6 spp. in Brazil, only one endemic.

82. Gossypium L. Shrubs, subshrubs, or small trees. 52 species, pantropical, in 4 subgenera:

§ subg. Gossypium ‣ 16 spp., Old World, inc. two cultigens.

§ subg. Houzingenia ‣ 14 spp. from Mexico (12, 11 endemics), Arizona (1), Ecuador (Galapagos) and Peru one endemic each.

§ subg. Karpas ‣ the tetraploid species, the cultigen G. hirsutum L. and G. barbadense L., Dominica Republic, Galapagos Is. and Hawaii one endemic each, and G. mustelinum (G.Watt) Miers endemic to NE Brazil. G. mustelinum (G.Watt) Miers is known only semi-arid region of Ceará, Rio Grande do Norte, Pernambuco and Bahia states, only in seven populations, near perennial and semi-perennial sources of water, such as ponds or pools in intermittent streams, the species has never been bred or commercially exploited, and there is no report of any kind of domestic use by the people who live in localities surrounding the plant populations.

§ subg. Sturtia ‣ 17 spp., endemic to Australia.

83. Hampea Schltdl. Trees or shrubs, mostly dioecious; leaves petiolate, the blades elliptic, ovate, or weakly lobed, entire; flowers solitary in the leaf axils or in axillary fascicles; petals whitish, glanddotted, reflexed. 21 spp. from Mexico to Colombia (3, two endemics and one up to reaching into Panamá).

84. Thespesia Sol. ex Corrêa. Tall shrubs to trees, to 30 m, with a vestiture of scales or stellate hairs; flowers perfect, showy, borne singly in leaf axils, sometimes grouped terminally by reduction of internodes; corolla campanulate or sometimes the petals spreading, usually yellow, less often white or deep rose, usually with a dark red center. 14 spp., five endemic to Africa, four endemic to the island of New Guinea, three endemic to the Greater Antilles, and the remaining two widely distributed in tropical littoral habitats around the world, with T. populnea (L.) Sol. ex Corrêa in New World, from Florida and Mexico to Suriname, and some record in NE Brazil.

7.6 MALVOIDEAE ▸ TRIBE HIBISCEAE (23/325–760) - outsiders Radyera (2; Nambia and South Africa; Australia), Hibiscadelphus (6; Hawaii), Symphyochlamys (1; tropical Africa), Megistostegium (3; Madagascar), Perrierophytum (6–9; Madagascar), Humbertiella (5; Madagascar), Helicteropsis (1; Madagascar), Humbertianthus (1; Madagascar), Cenocentrum (1; China, SE Asia), Anotea (1; Mexico), Jumelleanthus (1; Madagascar), Julostylis (3; SW India, Sri Lanka), Dicellostyles (2; Sri Lanka, Himalayas, Thailand), Nayariophyton (1; Himalayas, Yunnan), Kydia (2; Himalayas, Yunnan, SE Asia).

85. Hibiscus L. Subshrubs, shrubs, or trees (rarely herbs), glabrescent, pubescent, or hispid, sometines lianescent; leaves petiolate, the blades elliptic, lanceolate, ovate, or cordate, sometimes lobed or parted, dentate or less commonly subentire; flowers usually solitary (sometimes fasciculate) in the leaf axils, sometimes aggregated apically; pedpetals yellow (sometimes with a purple spot at base), lavender, red, or other colors, sometimes large and showy. 200 spp. almost worldwide, mostly tropical and subtropical, 87 in New World; South America has 53 species (32 in Brazil, 23 endemics), some almost weedy; all South american Hibiscus with more than 1 nectary are from Minas Gerais states (six) except H. pohlii Gürke, from Distrito Federal and adjacences.

86. Kosteletzkya C. Presl. Stout erect herbs or subshrubs, often with hispid vestiture, frequently growing in moist habitats; flowers solitary in the leaf axils or aggregated into terminal panicles or racemes; corolla rotate or tubular, whitish, yellow, or pink. 17 spp. worldwide, 7 in Africa, another in Malesia, and 9 in New World, from S U.S.A. to Colombia, Venezuela, Peru and Ecuador (only K. depressa (L.) O.J. Blanch., Fryxell & D.M. Bates in South America), with one species in temperate North America.

87. Malachra L. Erect herbs or subshrubs, sometimes puberulent, more commonly hispid, often with urticating hairs; flowers in condensed, bracteate, headlike racemes, the ‘heads’ axillary or terminal, sessile or pedunculate; corolla white, yellow, or lavender, often inconspicuous. 10 spp., pantropical, but centered in N South America (9, five exclusive); 6 spp. in Brazil, two endemics.

88. Malvaviscus Fabr. Shrubs or small trees, sometimes scandent, branching pubescent or glabrous; flowers solitary in the leaf axils or sometimes in apical cymelike groups; calyx campanulate or tubular, 5-lobed; petals red (rarely white), auriculate toward base, forming a tubular corolla; fruits a fleshy schizocarp or berry, oblate, usually red (sometimes white), with 5 carpels, each 1-seeded; only fleshy-fruited genus in the subfamily except for Anotea from Mexico, and only blue-black fruits. 10 spp., from S U.S.A. to Peru and Brazil; widely naturalised worldwide; 4 in South America, only the widely distributed M. concinnus Kunth in Brazil, in Amazonas state.

M. palmatus Urb. from western Acre state is a rare plant in Brazil (unique among Malvaceae), by Plantas Raras do Brasil’s book, but there is no record of this species in Vascular Plants of America.

89. Pavonia Cav. Prostrate perennial herbs, erect subshrubs, or shrubs (rarely arborescent, often cauliflorous), sometimes viscid or glabrate; leaves petiolate, ovate, elliptic, lanceolate, oblanceolate, deltoid, sometimes lobed, asymmetrical, dentate or crenate (rarely entire); flowers solitary or paired in the leaf axils or aggregated in racemes, panicles, or heads; involucel 4-22- parted, the bractlets distinct or basally connate; calyx 5-10bed; petals white, yellow, lavender, purple, sometimes with a dark basal spot.

291 spp., 60 in Old World and 233 from S U.S.A. to Argentina and in the Caribbean Islands, absent only from Chile; 191 spp. in South America, 140 spp. in Brazil, 89 endemics, 78 present in the NE and SE regions, in shrublands, forest edges, rocky fields, and modified areas such as roadsides and abandoned croplands; few species occur in low, dry fields, shaded environments in forest interiors or in wetlands. The center of species diversity of section Lebretonia Rio Grande do Sul and Santa Catarina states of S Brazil, where the broadest range of morphological diversity occurs.

§ subg. Asterochlamys ‣ 32 spp.

§ sect. Albae ‣ 3 spp., Mexico to Colombia and Venezuela.

§ sect. Asterochlamys ‣ 29 spp., tropical America.

§ subg. Goetheoides ‣ 16 spp., all endemics to coastal forests of Brazil.

§ subg. Malache ‣ 18 spp.

§ sect. Collicolae ‣ 12 spp. in three sections (Collicolae, Rubriflorae, Conditiflorae), all endemics to Mexico.

§ sect. Laminares ‣ only one sp. endemic to Mexico.

§ sect. Malache ‣ 4 spp., coastal environments, Florida to Pará state in Brazil in Atlantic coast (only P. paludicola Nicolson in Brazil), and from Panamá to Ecuador in Pacific coast.

§ subg. Pavonia ‣ 114 spp.

§ sect. Cancellaria ‣ 4 spp., tropical America.

§ sect. Lebretonia ‣ 4 sections, 60 spp.

o subsect. Cordifoliae ‣ 7 spp., 4 endemics to Mexico and three in Chaco regions, two up to WC Brazil.

o subsect. Exsertae ‣ 14 spp., all in Caribbean except by P. lasiopetala Scheele from SW U.S.A. and NW Mexico, P. commutata Garcke and P. pabstii Krapovickas & Cristóbal in Brazil, one up to NE Argentina.

o subsect. Hastifoliae ‣ 33 spp., all from C Brazil to N Argentina, Bolivia, Paraguay and Uruguay by except P. nepetifolia (Standley) Standley endemic to Mexico.

o subsect. Lebretonia ‣ 6 spp., C Brazil to NE Argentina.

§ sect. Lopimia ‣ 23 spp., tropical America.

§ sect. Malvaviscoides ‣ 32 spp., tropical America.

§ sect. Pavonia ‣ 15 spp., tropical America.

§ subg. Typhalea ‣ 34 spp.

§ sect. Varians ‣ only one sp., P. varians Moricandi, endemic to central Brazil in the states of Bahia, Ceará, Pernambuco, and Piauí, at 500-800 m elevation.

§ sect. Carcerariae ‣ 3 spp., two endemics to Mexico and one endemic to NE Brazil.

§ sect. Diathericae ‣ only one sp., endemic to mangroves in Pacific coast of Colombia.

§ sect. Typhalea ‣ 9 spp., tropical America.

§ sect. Urenoideae ‣ 20 spp., tropical America.

90. Peltaea (C. Presl) Standl. Shrubs or subshrubs, erect or decumbent, stellate-pubescent. Leaves short-petiolate, the blades elliptic or ovate to suborbicular, crenate-serrate, acute or obtuse; flowers solitary in the leaf axils (two species) or more commonly grouped in head-like aggregations subtended by specialized floral bracts, the bracts subsessile, cordate-ovate, serrate, the ‘heads’ subsessile to long-pedunculate; pedicels usually short; involucel of 8-13 bracts, these spatulate or bifurcate; calyx 5-lobed; petals relatively showy, yellow or rose-lavender. 20 spp. from New World, from tropical Paraguay and north to Mexico, and in the Caribbean Islands; 19 spp. in South America (the exception a Caribbean endemic), 13 in Brazil, 6 endemics.

91. Phragmocarpidium Krapov. Shrubs ca. 1 m tall, with scattered coarse stellate pubescence; leaves petiolate; flowers solitary in the axils of reduced leaves (floral bracts) on long lateral flowering branches; pedicels 1 cm long or less; involucel of 8 filiform bracts, ca. half the length of the calyx; calyx 1.5 cm long, ca. half-divided; petals 2- 3 times the length of calyx, rose (or whitish) with dark red center and venation, glabrous. Only one species, P. heringeri Krapov, from central Brazil, in Goias state and Distrito Federal.

92. Rojasimalva Fryxell. Decumbent perennial herbs, sometimes repent; inflorescences terminal, a headlike group of 2-4 flowers subtended by 2-3 floral bracts, these sessile, ovate, foliaceous; pedicels 1- 4 mm long; involucel of ca. 10 linear, setose bracts, exceeding the calyx; calyx ca. half divided, the lobes 3-nerved, the margin ciliate; petals lavender, 3-4 times the length of calyx, glabrous. Only one sp., R. tetrahedralis Fryxell, endemic to coastal mountains of N Venezuela.

93. Talipariti Fryxell. Trees. 21 spp., 20 from SE Asia to Pacific islands, and only one native in New World, T. pernambucence (Arruda) Bovini from Central and South America, along tropical coasts southwards up to SE Brazil and N Peru.

94. Urena L. Shrubs 0.5-2 m tall, more or less stellate-pubescent, leaves petiolate, the blades variable, often 3-5-angled, -lobed, or -parted, less often ovate, oblong, or lanceolate, crenate or serrate, with one or more prominent foliar nectaries on the abaxial side, these often yellow-bordered; flowers solitary or glomerulate in the leaf axils or forming terminal racemes; petals rose or lavender. Three spp., pantropical, two are cosmopolitan weeds in tropical and temperate areas of the world, and a third, U. pedersenii Krapov, with lilac flowers, is a very narrow endemic to Cristalina municipality in Goias state and Distrito Federal, in grasslands of C Brazil.

95. Wercklea Pittier & Standl. Coarse herbs, shrubs or trees up to 20 m tall, stellate-pubescent or glabrate, the stems with prominent leaf and stipule scars, sometimes with ligneous prickles on stems and spinescent prickles or pungent hairs elsewhere; calyx stellate-pubescent or sometimes also prickly; corolla 6-15 cm long, campanulate or salverform. 13 spp. from Neotropics: the Caribbean Islands, Costa Rica to Colombia and Ecuador (4 in South America), with a largely circum-Caribbean distribution.

8. SUBFAMILY BOMBACOIDEAE (18/160–175) - three subtribes, all in South America.

Subfamily Bombacoideae is mostly present in the New World tropics and contains 27 genera (c. 250 species); a few of those genera (Adansonia L., Bombax L., Camptostemon Mast., and Lagunaria (DC.) Rchb.), containing c. 19 species, are restricted to the Old World tropics. In Bombacoideae clade, general notes are: Cavanillesia arborea is the largest pachycaul tree in Brazil, know as barriguda; Ceiba pentandra (L.) Gaertn. - economically important kapok producer; introduced into and cultivated in the Old World. Chiranthodendron pentadactylon (the Devil's Hand Tree) has great potential as ornamental. Ochroma pyramidale (Cav. ex Lam.)Urb. yields the extremely light balsa wood. Phragmotheca, Matisia, Quararibea - the fruits of some species are fleshy, aromatic and edible. Patinoa spp. the pulpy fruits are edible or used locally as fish poison. For instance, Adansonia L., Cavanillesia Ruiz & Pav. and Ceiba Mill. are pachycaulous with barrel-trunks for water-storage and Pseudobombax Dugand has a photosynthetic underbark. Therefore, the massive horizontal xylopodium-like organ of E. saxicola could be interpreted as an adaptation to the harsh edaphic and microclimatic conditions of inselbergs, with their high degree of solar radiation and high evaporation rates (Porembski et al. 1998) that presumably exerted selection pressure on E. saxicola to adapt to this unique environment.

In recent phylogenetic studies, the Malvoideae is resolved as sister to the Bombacoideae with some intermediate genera (e.g. Pentaplaris and Matisia).

In the Paleotropics, it is represented by fewer than 18 native species in three genera: Adansonia L. (8-9), Bombax L. (three or four species), and Rhodognaphalon (Ulbr.) Roberty (three species); whether the disjunct distribution of Ceiba pentandra (L.) Gaertn. and Pachira glabra Pasq. in the African and American continents is natural or anthropogenic has long been controversial; ignoring these two widely cultivated species, three independent migrations from the Neotropics to the Paleotropics are usually invoked to explain the worldwide distribution of Bombacoideae.

Key differences from similar groups

The families and subfamilies listed below differ from the Malvaceae subfamily Bombacoideae as follows:

ü Araliaceae - if trees, then not bottle-shaped nor with stout prickles; stipules absent; petioles often sheating the base of the leaf; flowers in umbellate units; stamens equal in number to the petals and free, anthers dithecal; fruits usually fleshy (rarely dry) drupes or berries, never with kapok.

ü Brownlowioideae - sepals fused into a campanulate or urceolate tube; an androgynophore is always absent.

ü Malvoideae - mostly shrubs and herbs without stout, bottle-shaped and/or prickled trunks.

ü Byttnerioideae - sepals free; epicalyx always absent; petals cupped or hooded with strap-like appendages, free from the androecium; style unbranched.

ü Cochlospermaceae - anthers dehiscing via pores.

ü Convolvulaceae - stipules absent; leaves pinnately veined; stamens 5.

ü Grewioideae - without stout, bottle-shaped or prickled trunk; epicalyx always absent; sepals free; petals usually yellow or white, often with hairy basal nectaries; stamens free, rarely fasciculate.

ü Helicteroideae - calyx tubular; petals free and clawed; androgynophore usually present; stamens 10-30; ovaries usually apocarpous, except Ungeria and Reevesia.

ü Sterculioideae - petals always absent; androgynophore usually present; the stamen filaments free; ovaries apocarpous.

8.1 BOMBACOIDEAE ▸ TRIBE BERNOULLIEAE (3/8) - all genera in South America.

96. Bernoullia Oliver. Trees, leaves 3-7 digitate; flowers brownish to orange. Three spp., two in Mexico and Central America, and B. uribeana Cuatrec. endemic to Colombia.

97. Gyranthera Pittier. Tall, dioecious trees up to 65 m high, leaves digitate, possibly the tallest Malvales in South America and tallest tree in Venezuela. Two spp., one from Panamá, and G. caribensis Pittier endemic to Venezuela; one undescribed spp. from Colombia and N Ecuador.

98. Huberodendron Ducke. Tall trees up to 50 m tall, one of them the tallest Malvales from Brazil, buttressed presents, leaves simple; inflorescences cincinnate (fascicles pauciflorous). Three spp. from Guianas extending south to Brazil (two spp., one endemic), Bolivia up to Ecuador and Peru.

8.2 BOMBACOIDEAE ▸ TRIBE ADANSONIEAE (5/32) - outsider Adansonia (8; tropical Africa, Madagascar, NW Australia).

99. Aguiaria Ducke. Tall trees 50 m tall with small, one of them the tallest Malvales from Brazil, up to 50 cm tall buttresses, with also lepidote indumentum; leaves simple; small, less than 4 cm long fruits, with dehiscent exocarp that splits off into five valves, but these remain attached to the indehiscent endocarp, superficially resembling the fruits in the genus Cedrela, a remarkable morphology of its anemochoric fruits that are unique among Bombacoideae and angiosperms as a whole. Only one sp., A. excelsa Ducke, largest tree endemic to the Guiana Shield of Upper Rio Negro, in a very small dense forest in NW Amazonas state in N Brazil.

100. Cavanillesia Ruiz & Pav. Deciduous trees up to 30 m tall, often tall, sometimes with swollen trunks, buttressed. 4 spp. from Panamá to Bolivia, up Venezuela, and NE Brazil; one is endemic to Colombia, another from Nicaragua to Peru, C. hylogeiton Ulbr. occur on Brazil and Bolivia, and C. umbellata Ruiz & Pav., the Brazilian baobab, occur in Brazil, Peru, Colombia and Venezuela.

C. umbellata grows approaching 30 m in height, impressive in its native habitat as some of the baobabs; its massively thickened trunk, sometimes bearing peculiar, oddly shaped tubercles, can be more than 1.5 m thick and is topped by a rather sparse canopy of limbs and foliage; its bark ranges from relatively smooth, often with low horizontal ridges, to rugose (rough).

101. Catostemma Benth. Trees, often tall, rarely shrubs, buttressed absent, leaves unifoliate, flowers in fascicles, axillary; petals white. 15 spp., from Brazil, Venezuela, Guyana and Colombia, one up to Ecuador; 7 spp. in Brazil, 5 endemics.

102. Scleronema Benth. Tall trees, cylindrical thunks, buttressed absent; leaves simple; flowers solitary or in axillary fascicles; petals unguiculate. 6 spp., from Colombia, Venezuela, Guyana and N Brazil (5, three endemics, in Amazonas and Pará states), low high (5m).

8.3 BOMBACOIDEAE ▸ TRIBE BOMBACEAE (10/120-125) - outsiders Bombax (9; tropical regions in the Old World), Rhodognaphalon (2; tropical Africa), Neobuchia (1; Haiti).

103. Ceiba Mill. Tall trees up to 50 m tall, one of them the tallest Malvales from Brazil, with horizontally branched crown and a very straight trunk; bark pale grey, smooth, usually with scattered conical spines, commonly ventricose trunk, digitately compound and pulvinate leaves clustered toward the branch tips, leaflets articulate with the petiole, calyx opening irregularly, androecium of five stamens with filaments partially or fully connate in a tube and capsules with abundant kapok. 20 spp. from tropical America, from Sonora (Mexico) to Argentina - 17 in South America, mainly in dry areas in E Brazil (11, 6 endemics, some very narrow endemic), except for C. pentandra (L.) Gaertn., the Silk Cotton or Kapok Tree, which occur both in the Neotropics and West Africa.

104. Eriotheca Schott & Endl. Mostly medium to emergent trees with unarmed trunks, sometimes with xylopodium; leaves that are palmately compound with leaflets that are articulate at the petiole apex; relatively small flowers with a persistent calyx that is accrescent in fruit; capsules with abundant brown kapok; and numerous, relatively small, striate seeds. 27 spp. restricted to South America (21 in Brazil, 13 endemics).

105. Pachira Aubl. Trees, sometimes dioecious up to 50 m tall. 58 spp., 52 neotropical and six species (subg. Rhodognaphalon) in tropical Africa; 49 spp. in South America, slightly centered in forest of NW South America, 20 in Brazil, 9 endemics, some very narrow endemics; which are mostly evergreen and found in Amazonian white-sand forests and riparian forests, but also in Atlantic Forest and arenaceous-quartzitic rock outcrops in the Espinhaço Range in the states of Bahia and Minas Gerais. P. glabra Pasq. is often found in cultivation, mostly in eastern Brazil where it is probably native, although its original distribution area has been attributed to the African continent.

P. aquatica Aubl. and P. insignis (Sw.) Sw. ex Savigny from tropical America has the largest flowers of New World plants, with width up to 66(-71) cm if held horizontally.

106. Pochota Ram. Goyena. Only one sp., P. fendleri (Seem.) W. S. Alverson & M. C. Duarte, known from Nicaragua to N South America, and Brazil (only Roraima state).

107. Pseudobombax Dugand. Deciduous trees, trunks usually unarmed; leaves that are digitately compound, pulvinate, and clustered at branch tips, with the petiole widened at apex; leaflets that are not articulate at the petiole apex; a persistent calyx that is accrescent in fruit. 25 spp. from Mexico to South America (23), 15 in Brazil (10 endemics), seeds wind dispersed; P. minimum Carv.-Sobr. & L.P. Queiroz may be promptly recognized by its extremely reduced flowers (to 6 cm long) and fruits (to 5.5 cm long), the smallest recorded for this genus.

108. Spirotheca Ulbr. Epiphytes with strangler capacity, sending down roots and sometimes forming trees of 30 m or more (depending on size of host), or free-standing shrubs or small trees; trunk, and particularly the branches, usually aculeate; leaves 3–7-foliolate; flowers axillary, usually solitary at the ends of branches; petals 5, narrowly oblong, initially whitish-greenish or pink, becoming dark reddish with age, or consistently bright red. 6 spp., 4 in moist mid- to high elevation forests from Costa Rica and Panamá through the western Andean countries to Bolivia, and two disjuncts in coastal forests of SE Brazil: S. rivieri (Decne.) Ulbr. and S. elegans Carv.-Sobr., M. Machado & L. P. Queiroz, latter restricted of Bahia state, only found to inhabit seasonally dry tropical forest vegetation.

44. BRASSICALES

FAMILIES ABSENTS IN SOUTH AMERICA: AKANIACEAE (2/2), EMBLINGIACEAE (1/1), GYROSTEMONACEAE (5/20), LIMNANTHACEAE (2/8), MORINGACEAE (1/13), PENTADIPLANDRACEAE (1/1), RESEDACEAE (11/116), SALVADORACEAE (3/10), SETCHELLANTHACEAE (1/1) AND TIGANOPHYTACEAE (1/1).

LINEAGE 1 of 4: TROPAEOLIDS

TROPAEOLACEAE

§ FULLY CHLOROTROPHIC

Genera/species 1/94 Distribution mountain areas from S Mexico to Tierra del Fuego, S Brazil to Argentina. Habit herbs. Tropaeolaceae is a small family endemic to South and Central America, more often in the highlands. Tropaeolum majus L. is edible; its fruit and flowers taste like capers and are used to garnish salads.

SYSTEMATIC a single genus.

1. Tropaeolum L. Herbaceous climbers often with tuberous roots; plants dioecious; leaves simple, alternate, estipulate, entire and often peltate or deeply lobed, to palmate; inflorescence axillary, few-flowered or flowers solitary; flowers showy, colourful, hermaphrodite, usually zygomorphic, sepals 5, united forming a nectar spur or calcar; petals 5, clawed, 3 lowermost frequently variously marked towards center of flower; fruits fleshy schizocarp, splitting into 3 one-seeded fruticules, or fruit 3-winged; seeds rounded. 94 spp., all occur in South America, distributed mostly in Andean countries and in the highlands of other regions (e.g. Colombia, with 28 spp., 65 in over northern Andes) up to E Brazil (4, two endemics) and S Venezuela, and 4 reaches into Central America and southern Mexico; two sections:

§ sect. Tropaeolum ‣ include the formerly recognised sections Bicolora, Dipetala, Mucoronata, Schizotrophaeum, Serratociliata, Tropaeolum, and Umbellata.

§ sect. Chilensia ‣ include the formerly recognised genera Magallana and Trophaeastrum, and the formerly recognised Tropaeolum sect. Chymocarpus.

LINEAGE 2 of 4: CARICIDS

CARICACEAE

§ FULLY CHLOROTROPHIC

Genera/species 6/c. 35 Distribution Mexico, Central America, Caribbean, South America to northern Chile and Argentina, with their largest diversity in Mexico; Cylicomorpha: tropical W and E Africa. Habit usually dioecious (sometimes monoecious, andromonoecious, gynomonoecious, or polygamomonoecious; rarely bisexual), evergreen trees or shrubs (rarely perennial herbs or climbing shrubs; in Jarilla and Jacaratia corumbensis with a stout tuberous stem); stem usually unbranched, often spiny. All species produce latex that can be white or light yellow.

Carica papaya L is an important food crop, especially in NE Brazil; this species has two types of plants: one with many-flowered inflorecences comprising staminate and pistillate flowers, that sometimes produce an elongated fruit, and the other with pistillate, sessili flowers, producing abundant rounded fruits, known as Papaya, Mamão (Braz.) and Fruta-Bomba (Caribbean); the wild form occur only in Mesoamerica from southern Mexico to Costa Rica.

Use Ornamental plants, fruits, papain for medical and technical purposes (softening of meat, tanning of leather).

Key differences from similar families - differs from Brassicaceae and Tropaeolaceae in woody, shrubby to tree-like habit; from Araliaceae (similar due to its palmately compound leaves) in superior ovary and pentamerous flowers (vs. Araliaceae inferior ovary, tetramerous flowers); from Bombacaceae (similar leaf shape and sometimes spiny trunk) in succulent indehiscent fruit (vs. Bombacaceae fruits which are dehiscent capsules with woolly seeds).

Key to genera of South American Caricaceae

1. Leaves entire to deeply lobed, branches smooth ------------ Vasconcellea

2. Leaves digitate to trifoliolate, branches often spiny ------------ Jacaratia

SYSTEMATIC outsiders Cylicomorpha (2; tropical Africa), Carica (1; S Mexico, Central America), Jarilla (3; Mexico, Guatemala), Horovitzia (1; Oaxaca in Mexico).

1. Jacaratia A.DC. Pachycaul trees sometimes with spiny branches, leaves digitate, sometimes with taproot tubers. 7 spp., two only in Mexico/Central America, remainig five up to South America (three restricted); among South American species, J. corumbensis Kuntze is adapted to dry areas of Bolivia, Brazil and Paraguay, remaining are tropical rain forests dwellers; 4 spp. in Brazil, J. heptaphylla (Vell.) A. DC. restricted from coastal Bahia to Mato Grosso do Sul state.

2. Vasconcellea A. St-Hill. (inc. Carica p.p.) Unbranched or little brached tree, with pithly internodes,sometimes lianoid. 26 spp., Mexico to Argentina and SE Brazil, with one sp. from coastal Chile, only two also outside in South America; northern Andes is the center of species diversity of Caricaceae (highly centerd in S Ecuador and N Peru), and two reaching into Central America; 4 spp. in Brazil, none endemics. V. horovitziana (V.M. Badillo) V.M. Badillo endemic to Ecuador is only lianoid species in Caricaceae.

LINEAGE 3 of 4: SALVADORIDS

BATACEAE

§ FULLY CHLOROTROPHIC

Genera/species 1/2 Distribution New World and Oceania Habit herbs.

SYSTEMATIC a single genus.

1. Batis P. Browne. Perennial, herbs, shrublets or shrubs, stem prostrate or ascending, leaves fleshy, opposite, decussate, simple, sessile, linear to clavate, yellow-green to green, margin entire; dioecious or monoecious; inflorescences usually axillary, densely spicate, strobilate, conical, with flowers in four ranks; flowers bracteate (bracts cochleariform and imbricate in male inflorescences, smaller and partially incorporated in the fleshy female spike), small, unisexual, actinomorphic; fruit an aggregate or a drupe with four pyrenes; seed coat membranous. Two spp., dioecious B. argillicola P. Royen., from coasts of southern New Guinea and northern Australia, and the monoecious B. maritima L. from tropical and subtropical coasts of America and the Pacific Islands, from California to Peru and the Galapagos Islands; in the Atlantic from Florida to Brazil in Pará to Rio Grande do Norte states (disjunct records also in Sergipe) and the Antilles; both species are ecologically important for regeneration of mangrove areas.

KOEBERLINIACEAE

§ FULLY CHLOROTROPHIC

Genera/species 1/2. Distribuição SW U.S.A. to Mexico, Bolivia. Habit bisexual, evergreen shrubs or small trees. Xerophytes. Branches photosynthesizing, terminating in an elongated spine. Prophylls basal on branches and spines.

SYSTEMATIC a single genus.

1. Koeberlinia Zucc. Shrubs or small trees, strongly xerophytic; stems greenish and photosynthetic, divaricately and intricately branched, spinescent at the tips; leaves scale-like; flowers in umbel-like axillary racemes, actinomorphic, hermaphroditic, sepals 4, petals 4, imbricate, free; fruit a subglobose berry; seeds 2-4, exarillate, with scanty endosperm, embryo curved. Two spp., K. spinosa Zucc. from the S U.S.A., N Mexico, and K. holacantha W.C. Holmes, K.L. Yip & Rushing in dry forests of Santa Cruz, Bolivia, 1,700-2000 m elevation range.

LINEAGE 4 of 4: BRASSICIDS

TOVARIACEAE

§ Analyzed in 08.01.2021

Genera/species 1/2. Distribuição Mexico to Venezuela and Bolivia, Jamaica. Habit Bisexual, annual or perennial herbs, or evergreen shrubs or suffrutices (sometimes climbing or arborescent). Leaves when dry with coumarin-like scent.

SYSTEMATIC a single genus.

1. Tovaria Zucc. Shrubs or herbs, with a penetrating smell; leaves alternate, trifoliolate; stipules minute; inflorescences terminal and axillary, elongate racemes; flowers bisexual, actinomorphic; sepals (6-)8(-9); petals (6-)8(-9), free, shortly clawed, green to yellow; fruits berries; seeds many, small. Two spp., T. pendula Ruiz & Pav. common in disturbed habitats such as roadsides and landslines in mountains areas to 3,000 m from Mexico, Central America, NW South America (especially the Andes) to Peru and Bolivia, and T. diffusa (Macfad.) Fawc. & Rendle endemic to Jamaica.

CAPPARACEAE

§ FULLY CHLOROTROPHIC

Genera/species 16(own data)/460-470. Distribution cosmopolitan except continental Antarctica; Habit usually bisexual (rarely monoecious, andromonoecious or dioecious), usually perennial, biennial or annual herbs (sometimes evergreen or deciduous shrubs, suffrutices, rarely lianas or small trees). Some species are succulents and some are aquatic. Many species are xerophytes.

As currently circumscribed, it contains 33 genera and about 700 species. Based on morphological grounds and supported by molecular studies the American species traditionally identified as Capparis have been transferred to resurrected old generic names and several new genera have been recently described. Species of this subfamily are often dominant evergreen elements in dry forests. The flowers are ephemeral, mostly nocturnal. Some species (e.g. Quadrella cynophallophora (L.) Hutch., Sarcotoxicum salicifolium (Griseb.) Cornejo & Iltis) are cultivated as ornamental. Both humans and monkeys eat the pulp of the fruits of several species of Capparidastrum and Neocalyptrocalyx. The pulp of the fruits of species of Calanthea and Sarcotoxicum is highly poisonous.

SYSTEMATIC outsiders Hispaniolanthus (1; Hispaniola), Caphexandra (1; Central America), Acanthocapparis (1; Mexico to Honduras), Boscia (c 30; tropical and southern Africa, Madagascar, Arabian Peninsula), Maerua (c 75; tropical regions in the Old World), Bachmannia (1; Mozambique to KwaZulu-Natal and E Cape), Ritchiea (c 30; tropical Africa), Buchholzia (2; tropical W Africa), Euadenia (3; tropical Africa), Cladostemon (1; Mozambique, Zimbabwe, Swaziland, KwaZulu-Natal), Dhofaria (1; Oman), Cadaba (c 30; S Africa, Madagascar, Indian Ocean islands, Arabian Peninsula, India, Australia), Apophyllum (1; E Australia).

1. Anisocapparis Cornejo & Iltis. Shrubs or trees, leaves simple. Only one sp., A. speciosa (Griseb.) X. Cornejo & H.H. Iltis, ranging from Bolivia into adjacent parts of Mato Grosso do Sul state in C Brazil, Paraguay and N Argentina.

2. Atamisquea Miers. Shrubs or small tres, with densely lepidote branches. Only one sp., A. emarginata Miers ex Hooker & Arnott, disjunct in SW U.S.A., Mexico and Andes of Chile, Argentina and Bolivia, in semideserts.

3. Beautempsia (Benth. & Hook.) Gaudich. Only one sp., B. avicenniifolia (Kunth) Alleiz., restricted to the very dry woodlands and thorn scrub of W Ecuador and deserts in Peru.

4. Belencita H. Karst. Evergreens shrubs or trees up to 3m tall. Only one sp., B. nemorosa, from Colombia and Venezuela, in dry forests or scrubs in coastal region.

5. Calanthea (DC.) Miers. Two spp. restricted to the xerophytic forests of E Colombia and Venezuela.

6. Capparicordis Cornejo & Iltis. Shrubs or small trees, or rarely scramblers or lianas; leaves 4–10 × 4–10 cm; flowers few to c. 20, in short, open corymbiform racemes or rarely solitary; corolla lemon-yellow or orange; fruits dehiscent, ovoid to globose, baccate. Two spp., C. crotonoides (Kunth) Iltis & Cornejo from N Chile and S Peru, and C. tweediana (Eichler) Iltis & X. Cornejo, from NW Argentina to Bolivia and Paraguay, and extending, but barely, into SW Brazil at Corumbá, in semi-deciduous dry forests of the Chaco vegetational province.

7. Capparidastrum (DC.) Hutch. (exc. Neocapparis) Shrubs to/or trees (rarely herbaceous in C. humile (Hassl.) X. Cornejo & H. H. Iltis), glabrous or covered by simple short trichomes; leaves simple, spirally arranged; inflorescences racemose, usually terminal, rarely cauline (C. frondosum (Jacq.) X. Cornejo & H. H. Iltis, in Hispaniola and Brazil); petals 4; fruits pendulous, capsular. 22 spp., ranging from S Mexico to N Argentina, and Caribbean, centered from the lowlands to the lower slopes at both sides of the Andes of Peru, Ecuador, and Colombia; 17 in South America, 6 in Brazil, none endmemics; inflorescence racemose, cauline in C. frondosum (Jacq.) Cornejo & Iltis individuous from Brazil and Hispaniola.

8. Colicodendron Mart. Evergreen shrubs or trees; leaves simple, short-petiolate to sessile, opposite to spirally or whorled; inflorescences terminal, subterminal and/or axilar, a panicle, raceme or racemose spike, rarely a solitary axillary flower (in C. yco); flowers with calyx 1-seriate, valvate; petals 4, imbricate or torsivus in bud, sessile; fruits amphisarcous or pseudoamphisarcous, woody. 5 spp., two from Venezuela, Ecuador and Peru, C. yco (Mart.) Mart. from NE Brazil and Guyana, with solitary yellow flowers, many branched; C. bahianum Cornejo & Iltis, a unbranched or sparsely branched shrub, restricted to the Atlantic coastal rainforests of the State of Bahia, E Brazil; and C. martianum Cornejo, a shrub 1-3 m tall in SE Bahia state in Brazil.

9. Crateva L. Shrubs or trees, deciduous, glabrous; bark smooth, brown; young branches with prominent yellowish lenticels; leaves 3-foliolate, petiolate; inflorescence of axillary or terminal, corymbose racemes; flowers bisexual or unisexual by abortion; sepals 4, open in aestivation, arising from a shallow receptacle; petals 4(5), rather large, clawed, white; fruit globose or ovoid with a coriaceous pericarp borne on a stout stipe. 12 spp., pantropical, 8 in Old World, 4 in neotropics, two in South America: C. tapia L. from Mexico to Argentina and the Lesser Antilles, inc. Brazil and C. yarinacochaensis Cornejo & Iltis a local endemic of the Amazonian rainforests in the Department of Ucayali, E Peru.

10. Cynophalla (DC.) J. Presl. 17 spp., all South American, 5 up to Caribbean and North America, distributed from Southern Florida and Mexico, slightly centered in NW Peru and N Ecuador; 7 spp. in Brazil, one endemic.

11. Mesocapparis (Eichl.) Cornejo & Iltis. Lianas or scandent shrubs, with slender and ± fractiflexus terminal branches; leaves simple, petiolate, alternately arranged; inflorescence a solitary, axillary and subterminal flower; floral bracts absent; flower buds subglobose; petals 4; fruit a pendulous globose pepo, containing many seeds embedded in a white pulp. Only one sp., M. lineata (Dombey ex. Person) X. Cornejo & H. H. Iltis, found in 4 disjunct areas: in Amazon rainforest in Acre/Amazonas states; in Pará and Maranhão states; in a single locality Rio Grande do Norte state; and from Bahia to Rio de Janeiro, in E Brazil; it’s a single lianescent Capparaceae of Brazil.

12. Monilicarpa Cornejo & Iltis. Subshrubs, shrubs or small trees, unbranched or multi-branched; leaves simple. Two spp., M. tenuisiliqua (Jacq.) X. Cornejo & H.H. Iltis from N Colombia, Venezuela and Trinidad & Tobago, and M. brasiliana (Banks ex DC.), X. Cornejo & H.H. Iltis disjunct to E Brazil, in Bahia to Rio de Janeiro coastal forests.

13. Morisonia L. Shrubs or small trees, up to 7m tall; leaves simple. Three spp., M. multiflora Triana & Planch. only in Colombia, M. oblongifolia Britton in W Brazil (Acre and Amazonas states), Bolivia and Peru, and M. americana L. from Mexico and Caribbean up to Ecuador, Colombia, Venezuela and Guianas.

14. Neocalyptrocalyx Hutch. Shrubs to tall trees, with ± straight terminal branches; leaves simple, petiolated, spirally arranged; inflorescence axillary and (sub)terminal, usually corymbose to raceme, rarely a solitary flower; fruit a pendulous amphisarcum, with a subwoody thick wall, 3–10 mm thick, containing many seeds embedded in a white or orange pulp. 11 spp., ranging east to the Andes, from Venezuela (1 endemic) and the Guianas to Brazil, and recently collected in E Ecuador; in Brazil occur 6 spp., in N, NE & SE regions, 4 endemics.

15. Neocapparis Cornejo. (off Capparidastrum) Many branched trees; terminal branches, buds and young petioles with very short unbranched trichomes, otherwise glabrous; leaves spirally arranged, simple, tertiary nerves densely reticulate arranged on both sides. Two spp., one from dry forests, S Mexico (Oaxaca, Guerrero) to coastal N Colombia and Venezuela, and another in coastal Ecuador and Peru.

16. Preslianthus Cornejo & Iltis. Tall trees or shrubs; leaves spirally arranged; inflorescences terminal, unbranched and few flowered to loosely 5- to 15-branched and many flowered openly corymbose racemes or panicles or long penduncles; flowers often incompiscuous, greenish or greenish-yellow; fruit a amphisarcum, yellow at maturity. Three spp., one only in Panamá, and the two remainig restricted of dense forests of Costa Rica to Bolivia and W Brazil, disjunct in Guiana Shield and E Brazil.

17. Quadrella (DC.) J. Presl. 23 spp., U.S.A. and Mexico and Caribbean, two of then up to Colombia and Venezuela to Peru.

18. Sarcotoxicum Cornejo & Iltis. Shrubs to low trees; leaves simple, petiolate laxly spiral; inflorescence a short raceme and/or solitary flowers, terminal, subterminal and/or axilar; floral bracts absent; fruit a pendulous, spherical, cucurbitoid amphisarcum. Only one sp., S. salicifolium (Griseb) X. Cornejo & H. H. Iltis, restricted to Bolivia, Paraguay and N Argentina.

The fruits are edible at maturity if properly cooked, but extremely poisonous when immature, or even dried; it has been reported that the Matako Indians of the Argentine Gran Chaco eat them to commit suicide; during the Chaco war, many soldiers died after eating them; to make the pulp edible, the whole fruit must be boiled for 1 to 4 hours (depending on its size), changing the water once; the Matako Indians boil them six or seven times in different water to eat them; the Guaraní Indians use them as a hallucinogen or, in higher amounts, as poison.

19. Steriphoma Spreng. Shrubs up to 2 m tall, with many showy inflorescences, mainly orange. 7 spp. from Mexico to Peru and Venezuela.

CLEOMACEAE

§ FULLY CHLOROTROPHIC

Genera/species 25(own data)/c. 370. Distribution cosmopolitan except continental Antarctica; Habit usually bisexual (rarely monoecious, andromonoecious or dioecious), usually perennial, biennial or annual herbs (sometimes evergreen or deciduous shrubs, suffrutices, rarely lianas or small trees). Some species are succulents and some are aquatic. Many species are xerophytes.

SYSTEMATIC outsiders Areocleome (1; N Australia), Arivela (12; Old World, mainly Australia), Cleome (27; E Mediterranean, Spain, N and E Africa, Arabian Peninsula, and E to Afghanistan, Pakistan, Turkmenistan, and Kazakhstan), Cleomella (22; C and W North America to S and C of Mexico), Coalisina (6; Egypt, Yemen, Ethiopia, Namibia, South Africa, Angola, Sudan, Mozambique, India), Corynandra (5; India to Malaysia), Dipterygium (1; Egypt to Pakistan), Gilgella (1; North and tropical Africa, Arabian Peninsula, east to Pakistan), Gynandropsis (1; Asia), Kersia (8; Namibia, Angola, South Africa), Polanisia (6; North America, Mexico), Puccionia (1; Somalia), Rorida (12; NE Africa through Arabian Peninsula to Afghanistan, Tajikistan, and Pakistan), Sieruela (36; Kenya, Madagascar, Mozambique, Tanzania, South Africa, SW Africa, Ethiopia, Angola, Somalia, Congo, Sierra Leone, India), Styidocleome (1; tropical and NE Africa, east through Arabian Peninsula, S Iran, Afghanistan, Pakistan, and NW India), Thulinella (1; NE Africa, Arabian Peninsula, and E to S Iran).

Six neotropical clades are supported: (1) Iltisella, two spp., only in Mexico and Central America; (2) the Dactylaena clade, including Dactylaena and Physostemon; (3) the Andean clade, comprising four genera; (4) the Melidiscus clade; (5) the Cleoserrata clade, and (6) the Tarenaya clade; the last three corresponding to genera with the same name.

A. DACTYLAENA CLADE (3/13) ‣ all genera in South America.

1. Dactylaena Schrad. ex Schult. F. Annual or perennial herbs or subshrubs, highly branched; leaves alternate, compound, 3-foliolate; leaflets sessile or short-petiolulate; margin entire, ciliate or serrulate-ciliate; racemes terminal, ebracteate, bearing immature or mature fruits in lower clusters and separate clusters of flowers at apex; flowers zygomorphic; sepals 4, the lower one longer than others; petals 4, dimorphic, the upper pair typically longer than the lower. 7 spp., one from N Haiti, two from Bolivia and Argentina, and 4 in Brazil, one also in Venezuela (and three endemics), in montane forests, semideciduous to deciduous forests, border of forests, riparian forests, sandy coastal areas, riverbanks, rocky outcrops, roadsides, and waste disposal areas, at elevations from near sea level to 2550 m.

2. Haptocarpum Ule. Herb with clambering stems, perennial; leaves 3-foliate; petals 2; two functional stamens; fruits at dehiscence abscising the replum, leaving a forked base. Only one sp., Haptocarpum bahiense Ule, center Bahia state, Brazil, known only from type collection, this species is only Brassicales rare in Brazil, by Plantas Raras do Brasil’s book.

3. Physostemon Mart. & Zucc. Perennials or annuals, usually upright; leaves uni-, tri-, or pentafoliolate; stamens 6–10, most species have some stamens with apophyses. 8 spp., three only in Mexico, Central America and Caribbean, two scattered from Mexico to SE South America (both in Brazil), and two endemics to Brazil.

B. ANDEAN CLADE (4/39) ‣ all genera in South America.

4. Andinocleome Iltis & Cochrane. (inc. Cleome p.p.) Small herbs to shrubs or small trees; leaves usually 5–13-foliolate; the nectiferous disc is; usually conical or obconical and prominent; seeds usually lack an aril. 9 spp., predominantly Andean South America, with two species reaching Mexico, and one of these also occur in Hispaniola and Venezuela

5. Cochranella M.McGinty & Roalson. (inc. Cleome p.p.) Small to large unarmed annuals (0.75–1 m) to occasionally longer-lived tropical herbs, with a mixture simple and glandular-pubescent foliage; leaves 5–9-foliolate, estipulate; flowers with closed corolla aestivation, bisexual; petals glabrous, narrowly clawed, variable in color, usually white to pink or purple, but sometimes with a more green/greenish-yellow hue. Only one sp., C. pilosa (Bentham) E.M.McGinty & Roalson, W to S Mexico (Sinaloa to Chiapas) to Ecuador and Venezuela, Hispaniola; in wet areas.

6. Podandrogyne Ducke. (inc. Cleome p.p.) Unisexual flowers (monoecious plants); petals often orange or orange-red; placentas contorted and seeds have a large aril. 32 spp., 30 restricted from montanes foresty habitats from Venezuela to Bolivia, three of them up to Central America, and two restricted of Central America.

7. Pterocleome M.McGinty & Roalson. (inc. Cleome p.p.) Large, erect herbs or shrubs (1–2.5 m), simple to glandular-pubescent; leaves palmately compound 5–9- foliolate; racemes open, ebracteate, pedicels (2–5 cm); flowers with closed corolla aestivation, bisexual; sepals 1/5 the size of the petals at anthesis; petals pubescent, purple to green in color, not clawed. Only one sp., P. stylosa (Eichler) Iltis ex E.M.McGinty & Roalson, N Venezuela and Colombia; tropical to montane forests.

C. MELIDISCUS CLADE (1/2) ‣ a single genus.

8. Melidiscus Raf. Typically shrubs 2–6 m tall; leaves 5–11-foliolate; petals usually green to yellow. Two spp., M. giganteus (L.) Raf. from Mexico to Colombia, disjunct in Bolivia, and M. viridiflorus (Schreb.) Raf. from over tropical South America, disjunct in Mexico.

D. CLEOSERRATA CLADE (1/5) ‣ a single genus.

9. Cleoserrata Iltis. Annual herbs, glabrous, unarmed, with stipules diminutive or absent; leaves 3‒7(‒9)-foliolate, leaflets serrate-ciliate; inflorescences bracteate or not, bracts diminutive, caducous, or well-developed; flowers with showy petals, pinkish, purplish, ivory, or white. 5 spp., Mexico to Ecuador, Guianas and Caribbean up to E Brazil (two, one endemic).

E. TARENAYA CLADE (1/37) ‣ a single genus.

10. Tarenaya Raf. (inc. Cleome p.p., Hemiscola) Herbs or shrubs, annual; stems sparsely to profusely branched; glabrous or glandular-pubescent (sometimes spiny; inflorescences terminal or axillary (from distal leaves), racemes (flat-topped or elongated); bracts present; flowers (often appearing unisexual due to incomplete development), zygomorphic; sepals persistent, distinct, equal (each often subtending a nectary); petals equal; stamens 6; fruits capsules, dehiscent, oblong; seeds 10-30+, triangular to subglobose, not arillate, (cleft fused between ends). 38 spp., T. afrospina (Iltis) Soares Neto & Roalson in Nigeria, Cameroon, Gabon and Congo Brazzaville, unique spiny cleomid in Old World, 37 in mainland from Mexico and Central to South America (33), 22 in Brazil, 13 endemics; few in temperate region or in shallow water.

BRASSICACEAE

§ FULLY CHLOROTROPHIC

Genera/species 346/c. 3,997 Distribution cosmopolitan, with their largest diversity in temperate regions in the Northern Hemisphere. Habit usually herbs (sometimes shrubs, rarely trees, very rarely lianas), taprooted or with few to many-branched and slender or woody caudex, sometimes with slender or tuberous rhizomes, rarely stoloniferous; terrestrial or rarely submerged aquatics, with a pungent watery juice, without or rarely with multicellular glandular papillae or tubercles; petals rarely lobate or fimbriate. Stamens (two, four or) six (24 in Megacarpaea), two outer often shorter than four inner (tetradynamous), approximately as long as petals; nickel or zinc accumulated in many species; selenium accumulated in, e.g., Stanleya sp.; from the sister family Cleomaceae, the Neotropical Brassicaceae are easily distinguished by having actinomorphic corolla, fruit septum, and curved embryos without a gap between the radicle and cotyledons, as well as by lacking the stipules and palmately compound leaves.

The Brassicaceae are easily distinguished by the cruciform corolla and tetradynamous stamens; the exceptions are in some species of Lepidium that have only two stamens and a rudimentary or absent corolla; the characteristic capsule (often called silique or silicle; see above) is distinctive in the family for all species with dehiscent fruits, in which the fruit valves fall off and the 'false' septum remains; the three distinctive features above collectively occur in the vast majority of species, and when one or two features do not apply then the third would help; for example, in Lepidium spp. with two stamens and apetalous flowers, the fruit type described above is distinctive.

Centered in U.S.A., China, Irano-Turanian region. Botanical discoveries of South American Brassicaceae date back to the 1753 descriptions of the Linnaean Lepidium bonariense L. and L. didymum L. that have since become cosmopolitan weeds; native species of South America gow primarily at high altitudes, some species of Rorippa and Cardamine grow in typical tropical lowlands. One Cakile spp. occur in coast of Colombia and Venezuela. Exhalimolobus, Descurainia, Physaria and Tomostima also occur in Uruguay. In Colombia occur 42 spp., all Draba (20), Descurainia sp. and Brayopsis sp. occur above 3,200 m elevation range; 11 anothers spp. in several genera occur only above 1,800 m elevation range; the remaining Pennelia sp., Cakile sp. and some Cardamine, Lepidium and Rorippa occur above 1,800 m elevation range.

The Brassicaceae include several important crop plants grown in the Neotropics as vegetables (e.g., species of Brassica); the only native Neotropical crop is Lepidium meyenii Walp. (maca) cultivated in the high Peruvian Andes and consumed locally. Vegetable oils of various Brassica species, especially Brassica napus L. (canola), probably rank first in terms of the world's tonnage production, but have not been grown in our area. Lepidium didymum L. and L. bonariense L. are among that notable native South American weeds that have become naturalized elsewhere in the world.

SYSTEMATIC two subfamilies, Aethionomoideae (1/c 55, Mediterranean, SE Europe and SW Asia to Afghanistan), absent in New World; among Brassicoideae, with five supertribes, with 57 tribes. Cochlearieae (2/c 35, Europe, Mediterranean, North Africa, NE Asia, Alaska, northern Canada) unplaced among the supertribes.

1. SUPERTIRBE ARABODEAE (45/680 - 750) ‣ 4 tribes, Alysseae (24/150–220, mainly S and SE Europe, NW Africa and SW Asia), Asperuginoideae (1/1, Türkiye and eastwards to Central Asia and Pakistan), Stevenieae (3/8, Europe, Central and E Asia) absents in South America. Among South American Arabideae, outsiders are Abdra (2; U.S.A.), Arabis (c 60; temperate regions on the Northern Hemisphere, tropical African mountains), Athysanus (1; W U.S.A.), Aubrieta (16; S Europe to Iran), Drabella (1; Europe to Türkiye and the Caucasus, Morocco), Pachyneurum (1; Central Asia), Baimashania (2; Yunnan, Qinghai), Pseudodraba (1; Afghanistan and Pakistan), Sinoarabis (1; Xizang), Arcyosperma (1; Nepal), Borodiniopsis (1; W China), Botschantzevia (1; Kazakhstan), Dendroarabis (1; Altai), Parryodes (1; Bhutan), Scapiarabis (4; N Pakistan, Central Asia, Tajikistan, Afghanistan, Xinjiang).

1. Draba L. Annual, biennial or perennial herbs (sometimes very diminutely), rarely stoloniferous or woody stems; basal leaves present, cauline leaves often absent, sometimes cushions. 423 spp., more than 110 in Himalayas (75 in restricted region) and neighbouring Central Asia, and about 90 species in Europe and the remainder of Asia (90 in Russia, 45 in Europe, 48 in China, 37 in India, 24 in Nepal, 23 in Pakistan, 10 in Afganisthan), 7 in Africa. As currently understood, Draba is represented in the New World by 212 species, of which 82 are restricted to South America (Colombia (21), Venezuela (12), Ecuador (20), Peru (25), Bolivia (10) and Cono Sur (10); and 130 are native to North America, including only five that are endemic to Mexico and Central America.

The genus shows a tremendous diversity in the growth habit, though most of the paramo taxa have woody lower stems and often form subshrubs; all range diversity of genus occur in South America; however, plants of the Venezuelan endemic D. lindenii (Hook.) Planch. are often shrubs 50–150 cm tall and have lower stems to 1 cm in diameter; by contrast, plants of the Bolivian endemic D. inquisiviana Al-Shehbaz are tiny, scapose perennials only 0.5–1.2 cm tall; all other species of Draba worldwide fall between these two extremes, and those with typically woody lower stems are restricted to South America.

2. Tomostima L. Herbs, annuals that produce early-season chasmogamous flowers with white petals and late-season cleistogamous flowers without petals. 6 spp., 4 in North America, T. araboides (Wedd.) Al-Shehbaz, M. Koch & Jordon-Thaden in Peru and T. australis (Hook. f.) Al-Shehbaz, M. Koch & Jordon-Thaden in Argentina and Uruguay.

2. MEGACORE BRASSICODAE (111/c. 1,000) ▸ 13 tribes, 10 off South America: Calepineae (3/7, Southern Europe, Mediterranean, SW and Central Asia), Conringieae (2/8, Europe to Central Asia), Eutremeae (3/34, E Europe, temperate Asia, one species also in North America), Isatideae (5/c 34, Europe, Central and SW Asia), Sisymbrieae (1/44, Europe, Mediterranean, N and S Africa, Asia, one species in North America), Thlaspideae (12/36, Europe, Mediterranean, SW and Central Asia, Himalayas), Aphragmeae (▇▇), Fourraeeae (▇▇), Plagiolobeae (▇▇), Schrenkielleae (▇▇); and three in South America.

2.1 BRASSICEAE ▸TRIBE COLUTEOCARPEAE (3/c 127) - outsiders Coluteocarpus (1; mountains in SW Asia), Pseudosempervivum (6; Türkiye, Armenia).

3. Noccaea Moend. Annual or perennial herbs. 128 spp., distributed primarily in Europe and Asia, with one species endemic each in N. Africa (Algeria), N. magelanica (Comm. ex Poir.) Holub in Patagonia from Argentina and Chile, N. mexicana (Standl.) Holub only in Mexico, and three in the U.S.A. (one of then reaches up to Mexico).

2.2 BRASSICEAE ▸ TRIBE THELYPODIEAE (28/260) - outsiders Chaunanthus (4; Mexico), Chlorocrambe (1; Oregon, Utah), Dryopetalon (8; SW North America), Hesperidanthus (5; W North America), Phravenia (1; N Mexico), Pringlea (1; Prince Edward Islands, Crozet Islands, Kerguélen Islands, Heard Island), Romanschulzia (14; Central America), Stanleya (6–7; W North America), Streptanthus (c 55; W and S North America), Thelypodiopsis (15; North America, Mexico, Guatemala), Thelypodium (19; W North America, Central America), Thysanocarpus (5; W U.S.A.), Warea (4; S U.S.A.).

4. Chilocardamum O. E. Schulz. Herbs perennial, with a well-developed woody caudex covered with leaf remains of previous years; stems erect, simple, rarely branched; basal leaves sessile, rosulate, simple; cauline leaves sessile, not auriculate at base; racemes several to many flowered, dense or lax, ebracteate, corymbose, elongated slightly in fruit; petals white. 4 spp., Patagonia, Argentina (Neuquen, Rio Negro, Chubut, Santa Cruz).

5. Dictyophragmus O. E. Schulz. Annual, taprooted, often glacous, glabrous spp. leaves all cauline. 3 spp., two in S Peru and one in N Argentina.

6. Englerocharis Musch. Perennial, scapose, cespitose herbs, often cushion forming, with thick, simple or branched caudex densely covered with persistent leaf remains of previous years (green or silvery green), mainly with 1-3 cm diameter and 2 – 10 cm high. 6 spp., one widely distributed in Peru but narrowly distributed in Bolivia, and remaining five all Peruvian endemics known thus far only from their type collections.

7. Hollermayera O. E. Schulz. Perennial, rhizomatous, glabrous herb; cauline leaves petiolate. Only one sp., H. valdiviana (Phil.) Ravenna, endemic to Chile.

8. Ivania O. E. Schulz. Perennial, taprooted, glabrous herbs. Two spp. from N Chile, from Valparaiso and Copiapo regions.

9. Mostacillastrum O. E. Schultz. Herbs perennial or rarely annual, with a woody caudex or base. Stems erect to ascending, rarely decumbent, simple or branched basally and/or apically; cauline and basal leaves present; petals white, rarely lavender or yellowish. 31 spp., 20 spp. in South America, M. pandurifolium (Kunth) Al. Shehbaz endemic to Venezuela, 19 from Peru southwards, two spp. in Haiti, and nine from U.S.A. to Mexico.

10. Neuontobotrys O. E. Schulz. Plants subshrubs, woody at base, rarely perennial herbs; stems erect to ascending, few to many, woody at base, often many branched above; racemes several to many flowered, lax, ebracteate, corymbose, elongated considerably in fruit; petals yellow drying reddish, rarely white or pink. 14 spp., Peru (6, two endemics), Bolivia to Patagonia.

11. Parodiodoxa O. E. Schulz. Perennial, cespitose, glabrous herbs. Basal leaves rosulate. Only one sp., P. chionophila (Speg.) O.E. Schulz, NW Argentina (Catamarca, Jujuy, La Rioja, Salta, and Tucuman), where it grows on open, rocky soils at elevations of 3,500–5,100 m.

12. Phlebolobium O. E. Schulz. Perennial, glacous, glabrous or sparsely pubescent herbs; basal leaves rosulate. Only one sp., P. maclovianum (d’Urv) O. E. Schulz., endemic to Falkland Is.

13. Polypsecadium O. E. Schulz. Plants annual or perennial herbs, subshrubs, or shrubs; stems erect to ascending, branched above; basal leaves absent; racemes many flowered, lax, ebracteate, corymbose, elongated considerably in fruit; petals white to lavender or purple, obovate to spatulate, apex obtuse. 15 spp., two in N Andes, 12 from Peru to S Argentina, and P. brasiliense O.E.Schulz, endemic to Santa Catarina state, Brazil.

14. Sarcodraba Gilg. & Muschl. Perennial, taprooted, cauline leaves sessile; petals white. 4 spp., Patagonian Argentina up San Juan and La Rioja provinces.

15. Sibara Greene. Annual or perennial herbs, rarely shrubs. Stems erect, simple or many branched, sometimes woody at base. Racemes many flowered, ebracteate, lax or rarely corymbose. Petals white or lavender to purple. Anagram of Arabis. 12 spp. exhibiting amphitropical disjunction: six of these grow in California and Baja California, 6 in dry areas of N Chile (4 endemic) and Argentina.

16. Weberbauera Gilg. & Muschl. Perennial, taprooted, pulvinate, pubescent herbs. Leaves entire to dentate or rarely pinnately lobed. 19 spp., Andes from Peru (15, 11 endemic) to Argentina, Chile, following the expansion of its earlier delimitation to include Catadysia O. E. Schulz.

17. Zuloagocardamum Salariato & Al-Shehbaz. Herbs having a well-developed woody caudex with reduced leafless stems, 5 cm tall, rosulate, awl-shaped or linear, sessile, parallel-veined basal leaves conspicuously ciliate with simple trichomes, racemes much shorter than the basal leaves, torulose fruits, and mucilaginous seeds; flowers white. Only one sp., Z. jujuyensis Salariato & Al-Shehbaz, endemic to Espinazo del Diablo in Jujuy Province in Argentina.

2.3 BRASSICEAE ▸ TRIBE BRASSICEAE (c 36/c 275) - all outsiders of Macaronesia and Mediterranean Basin up to N Africa and Sahara, Middle East, C Asia, Socotra and NW India except Brassica (c 165; Europe, Mediterranean, temperate Asia), Crambe (c 35; Europe, Macaronesia, Mediterranean, mountains in N tropical Africa, W Asia) and Orychophragmus (3; China). Brassica L. has 165 spp. in the present sense includes Brassica s.str. (c 40; Europe, Mediterranean, temperate Asia), Ceratocnemum (1; Morocco), Coincya (6; southern Europe, Mediterranean), Diplotaxis (c 25; Europe, Mediterranean to NW India), Eruca (1; Mediterranean, NE Africa), Erucastrum (c 25; Europe, Macaronesia, Mediterranean), Hirschfeldia (1; Mediterranean), Raphanus (4; Europe, Mediterranean to Central Asia), Rapistrum (2; Europe, Mediterranean, W Asia), Sinapidendron (4; Madeira), Sinapis (6; Europe, Mediterranean), Trachystoma (3; Morocco).

18. Cakile Mill. Annual, often glabrous; leaves fleshy. 6 spp., two in coasts in Europe, Mediterranean, Arabian Peninsula, 4 in New World, C. lanceolata (Willd.) O.E. Schulz reaching to coastal Colombia and Venezuela, two in Old World, one endemic to Middle East.

3. MEGACORE CAMELINODAE (78/c. 1,200) - 15 lineages, Alyssopsideae (4/9, E Mediterranean to China), Boechereae (8/c 118, Russian Far East, North America), Camelineae (8/36, Europe, temperate Asia, N North America), Crucihimalayeae (3/14, NE Asia, one species also in North America and SW Greenland), Erysimeae (1/c 180, Europe, Macaronesia, Mediterranean, Africa, temperate Asia, North America), Microlepidieae (16/55, Australia, New Zealand), Oreophytoneae (2/6, SW Europe, NW Africa, tropical E Africa), Smelowskieae (1/25, Pakistan to Central and NW Asia, NW North America), Turritideae (1/4, Europe, Africa, W Asia), Yinshanieae (1/7–13, China, N Vietnam), Malcolmieae (▇▇), Arabidopsideae (▇▇), Hemilophieae (▇▇) absents in South America, and 5 presents.

3.1 TRIBE CARDAMINEAE (c 12/c 335) - outsiders Aplanodes (2; KwaZulu-Natal; E Cape, Lesotho), Armoracia (3; Europe to Siberia, E North America), Barbarea (22; Europe, N Asia, North America), Iodanthus (1; U.S.A. from Connecticut to Texas), Leavenworthia (8; S and SW U.S.A.), Nasturtium (c 15; temperate regions on the Northern Hemisphere), Ornithocarpa (2; Mexico), Planodes (2; SE U.S.A., N Mexico), Selenia (5; S U.S.A., Mexico), Sisymbrella (1; W Mediterranean).

19. Cardamine L. Often compound leaves and linear fruits with flattened replum. 160-200 spp., 66 spp. in New World, 10 in Australian and New Zealand, 3 in Africa and 4 in New Guinea, 23 in South America, C. chenopodifolia Pers., C. africana L. and C. chenopodiifolia Pers. in Brazil, none endemics. in Brazil. Some spp. are invasive cosmopolitan weeds; C. chenopodifolia Pers., from Bolivia, Argentina, S Brazil, Paraguay, Uruguay and Chile, is a amphicarpic species, unique among amphigeocarpy Brassicaceae in New World.

20. Rorippa Scop. Mainly semi-aquatic small herbs. 80-90 spp., cosmopolitan, 25 spp. in Eurasia, 21 in North America, 12 in Africa, 14 in South America, and 9 (all in endemic sect. Ceriosperma) in New Guinea, Australia, New Zealand and Polynesia; lowland temperate and tropical mountains; they are distributed primarily along the Andes, where the grow in wet habitats at altitudes of up to 4,000 m. 3 spp. in Brazil, R. bonariensis (Poir.) Macloskie, R. clandestina (Spreng.) J.F. Macbr. and R. hilariana (Walp.) Cabrera, occurs in Brazil, mainly in southern places.

3.2 TRIBE LEPIDIEAE (1/c 250) – a single genus.

21. Lepidium L. (inc. Lithodraba) Annual, biennial or perennial, rarely thorn, glabrous or pubescent herbs, subshrubs, or rarely lianas (in Australia), sometimes cushions; reduction in floral organ number is common in many species, and over half of the species have only two or four stamens, rather than the usual six; petals are reportedly absent from about 1/4 of species, and they are rudimentary in many others. c. 220 spp., cosmopolitan, including Hawaii and New Zealand, in lowland temperate areas and in tropical mountains, but also tropical lowlands and dry areas (such as Namibia). 88 spp. occur in New World, 57 in South America, only three in Brazil: L. bonariense L. from S Brazil and adjacent Argentina, Paraguay and Uruguay, L. auriculatum Regel & Korn. from Peru to Brazil and Cono Sur, and L. didymum L. also from Peru to Brazil and Cono Sur.

The cultivated Lepidium of the Peruvian highlands (also in Bolivia and Cono Sur), better known as ‘maca’, have correctly been assigned to L. meyenii Walp. by various authors; one can draw so many similar parallels among the cultivated vs. wild forms of radish (Raphanus sativus L.), turnip (Brassica rapa L.), and rape (B. napus L.), all of which produce fleshy roots in cultivation but do not produce them when they become naturalized.

3.3 TRIBE DESCURAINIEAE (6/c 45) - outsiders Robeschia (1; the Middle East), Hornungia (5; Europe, Mediterranean, one species, a widely distributed weed), Ianhedgea (1; SW Asia to China).

22. Descurainia Webb. & Bertel. Annual, biennial or perennial, taprooted, pubescent herbs or shrubs. 35 spp., of which 13 are native in North America north of Mexico, 7 in Macaronesia, 4 in Central America, one in the Caucasus, D. sophia (L.) Webb ex Prantl is a cosmopolitan weed, 10 in South America, all from Peru to Patagonia, except two up to Colombia and Uruguay (D. erodiifolia (Phil.) Prantl. ex Reiche), and D. clefii Al-Shehbaz endemic to Colombia

23. Trichotolinium O. E. Schulz. Perennial, densely pubescent herbs or shrubs; leaves all in basal rosette. Only one sp., T. deserticola (Speg) O. E. Schulz, Patagonian Argentina.

24. Tropidocarpum Hook. Annual, pubescent herbs; petals yellowish or yellow. 4 spp., three in California and adjacent Mexico, and T. lanatum (Barnéoud) Al-Shehbaz & R.A. Price restricted of C Chile.

3.4 TRIBE PHYSARIEAE (7/133–138) - outsiders Dimorphocarpa (4; SW Canada, W U.S.A.), Dithyrea (2; SW U.S.A., NW Mexico), Lyrocarpa (3; California, Mexico), Nerisyrenia (9; S U.S.A., Mexico), Paysonia (8; SE U.S.A.), Physaria (105–110; SW Canada, W U.S.A., NW Mexico), Synthlipsis (2; S U.S.A., Mexico).

25. Physaria (Nutt. ex. Torr. & A. Gray) A. Gray. Perennial, taprooted, densely pubescent herbs, sometimes cushions; basal leaves rosulate, or cauline leaves. 110 spp., mainly North America, one extending in Arctic Russia, and six in South America, all in Argentina, one in adjacent Bolivia and P. mendoncina (Philippi) O’Kane & Al Shehbaz up to Uruguay, mainly pink or purple petals.

3.5 TRIBE HALIMOLOBEAE (6/39) - outsiders Halimolobos (8; SW U.S.A. to Central America), Sphaerocardamum (4; Mexico).

26. Exhalimolobos Al-Shehbaz and C.D. Bailey. Biennial herbs; stems erect, simple or branched above; basal leaves petiolate, not rosulate, coarsely dentate or sinuate; cauline leaves sessile, base auriculate to amplexicaul, dentate or sinuate-dentate; petals white, oblanceolate or spatulate, apex obtuse; fruits dehiscent, linear or rarely oblong siliques, terete or slightly angustiseptate. 9 spp., 4 endemics to Mexico, and 5 occur in South America: E. hispidulus Al-Shehbaz and C.D. Bailey occur disjunct in Mexico and from Venezuela to Bolivia; and four in Argentina and Bolivia, with E. weddellii (E. Fourn.) Al-Shehbaz & C.D. Bailey up to Uruguay.

27. Mancoa Wedd. Annual or biennial herbs; stems few to several from base, decumbent or ascending to erect, usually branched above; basal leaves petiolate, often rosulate, 1- or 2- pinnatisect, very rarely simple, dentate or ultimate lobes subentire; cauline leaves sessile or short petiolate, auriculate or not, dentate to pinnately lobed; petals white, obovate or spatulate, apex obtuse. 8 spp., 4 in Mexico, 4 in South America, widely distributed in Peru, Chile, Argentina and Bolivia.

28. Pennellia Nieuwl. Perennial herbs. Stems erect to ascending, often branched above. Basal leaves petiolate, rosulate, entire or dentate to sinuate or runcinate; cauline leaves short petiolate to sessile, not auriculate, entire or dentate to sinuate; flowers cup shaped; petals white throughout or lavender to purple at apex. 8 spp., six only in North and Central America up to Colombia, and three in Cono Sur, one also in Bolivia.

29. Petroravenia Al.Shehbaz. Perennial, rhizomatous, densely pubescent herbs; leaves all in basal rosette. Only one sp., P. eseptata Al-Shehbaz, endemic to NW Argentina (Jujuy and Salta provinces), where it grows on volcanic sandy soils of the Central Andean Puna between 4,500 and 4,600 m.

4. MEGACORE HELIOPHILOIDAE (48/c. 400) - 13 tribes, 10 absents in South America, Anastaticeae (13/96, Macaronesia and Mediterranean to Central Asia and NW India and E African mountains), Asteae (2/2, Mexico), Biscutelleae (5/61, Europe, Mediterranean, Asia), Chamireae (1/1, W Cape), Subularieae (3/4, Temperate regions on the Northern Hemisphere), Heliophileae (1/c 90, S Africa, especially Cape), Iberideae (2/29, Europe, Mediterranean, NW Africa, SW Asia), Megacarpaeeae (2/12, E Europe to Central Asia), Notothlaspideae (1/2, S Alps in New Zealand), Hillielleae (▇▇).

4.1 EUDEMEAE ▸ TRIBE EUDEMEAE (12/42-43) – all genera in South America. While the hemicryptophyte rosette is the ancestral growth form in Eudemae, the cushion habit evolved independently twice in the tribe (Dactylocardamum and Xerodraba). Delpinophytum, the remaining cushion-former originally included in Eudemeae, was placed within the lineage I, probably within tribe Lepidieae, showing that this character evolved twice also in South American members of this tribe (Lepidium and Delpinophytum). Therefore, cushion formation in South American Brassicaceae evolved independently four times. Cushion formation is rather uncommon elsewhere in the family and is known in Draba L., Baimashania, and Lepidium. Cushion formation is one of the most conspicuous growth forms in highly exposed alpine habitats and is especially abundant in temperate and subpolar regions. This life-form represents an efficient trap for heat and water, with a maximum reduction of losses due to its lowest surface-volume ratio. It is common in various parts of South America, such as Patagonia, the subantarctic mountains, and the tropical alpine environments where ‘winter’ comes nearly every night. Indeed, the revised worldwide catalogue of cushion plants, shows the Andes and Patagonia as the richest zones worldwide in the number of cushion-former genera. Therefore, convergence in cushion formation is not rare and represents an adaptation to extreme sites.

30. Alshehbazia Salariato & Zuloaga. Herbs perennial, cushions, with slender rhizomes the branches of which terminated by rosettes with petiolar remains of previous years; basal leaves rosulate, petiolate. Three spp., along the central and S Andes of Argentina, Bolivia, Chile, and Peru, in mountain wetlands, peat bogs, shaley soil by streamside, and moist seepage areas.

31. Aschersoniodoxa Gilg. & Muschl. (exc. Gongylis) Perennial, glabrous or pubescent herbs (1-2 cm diameter of rosettes); leaves basal, fleshy. Three spp., growing at high elevations (3,800 – 5,200 m) on scree in rather inaccessible areas. A. cachensis (Speg.) Al- Shehbaz is distributed in NW Argentina, W Bolivia, and C Peru, it is relatively uncommon and is known from a total of about two dozen collections; A. mandoniana has been collected about a dozen times from the high Andes of Dept. La Paz of Bolivia, where it is endemic; A. pilosa Al- Shehbaz is narrow Peruvian endemic known only from their type collection.

32. Ancashia Al-Shehbaz, Salariato, A. Cano & Zuloaga (inc. Brayopsis p.p., Dactylocardamum p.p., Weberbauera p.p.). 6 spp., endemics to Peru (4) and Bolivia (2).

33. Borealandea Al-Shehbaz, Salariato, A. Cano & Zuloaga (inc. Brayopsis p.p., Eudema p.p.). 3 spp., Colombia to Ecuador.

34. Dactylocardamum Al-Shehbaz. Perennial, pulvinate, pubescent herbs forming cushions, woody and much branched at base, 1-3 cm high, 3-4.5 mm in diameter; ultimate stems finger-like, leaves scale-like, high densely imbricate, resembling Xerodraba, flowers solitary (1 mm long), terminating branches, unique in crucifers in fruits axillary sandwiched imbricated leaves. Two spp., both restricted of Ancash department in Andean Peru.

35. Delpinophytum Speg. Perennial, glabrous subshrubs, cushions; stems finger-like; leaves all scale-like, densely imbricated; flowers solitary, terminating branches. Only one sp., D. patagonicum (Speg.) Speg., endemic to Patagonia in Argentina.

36. Eudema Humb. & Bonpl. (inc. Brayopsis p.p., exc. Borealandea p.p.) Perennial, often pulvinate, glabrous or pubescent herbs, all rhizomatous, sometimes cushions; leaves all in basal rosetes, single flowered. 9 spp. from Peru to Cono Sur.

37. Gongylis Theophr. ex Molinari & Sánchez. (off. Aschersoniodoxa) Herbaceous plants to 6 cm tall in fruit, with few branched thick caudex; basal rosettes 4 − 6 cm in diam.; rosette leaves fleshy, 1.5 − 3 cm including petiole, blade suborbicular-obovate, 0.7 − 1.3 cm wide, adaxially hirsute with trichomes 0.1 − 0.35 mm, glabrous abaxially, margin crenate-dentate; petiole flattened, 1 − 2 cm, glabrous; cauline leaves absent; flowers unknown. Only one sp., G. peruviana (Al-Shehbaz, Navarro & A. Cano) Sánchez & Molinari, endemic to Lima, Peru.

38. Onuris Phil. perennial, taprooted, glabrous or pubescent herbs, sometimes cushions; basal leaves present, also cauline leaves. 5 spp., S Argentina and S Chile.

39. Stenodraba O. E. Schulz. (exc. Stenodrabopsis) 4 spp., S Argentina and Chile.

40. Stenodrabopsis Al-Shehbaz, Salariato, A. Cano & Zuloaga (inc. Stenodraba p.p.). 3 spp., from Argentina and Chile.

41. Xerodraba Skottsb. Perennial, taprroted, pulvinate, pubescent herb to shrub (nanoplants), sometimes cushions; stems finger-like, leaves fleshy, scale-like, densely imbricate, single flowered by rosette. 5 spp., S Argentina and Chile.

4.2 EUDEMEAE ▸ TRIBE CREMOLOBEAE (4/c 32) - all genera in South America.

42. Aimara Salariato & Al-Shehbaz. Shrubs branched with well-developed cork; reduced and fleshy leaves. Only one sp., A. rollinsii (Al-Shehbaz & Martic.) Salariato & Al-Shehbaz, restricted to the Atacama Plateau in NW Chile, between 2,500 and 3,350 m with high temperature fluctuations. Typical shrubs are uncommon in Brassicaceae, though they undoubtedly evolved independently many times in the family.

43. Cremolobus DC. (exc. Yunkea) Annual or perennial herbs, shrubs or lianas, up to 5m tall. 5 spp., from center Colombia to center Bolivia, and one isolated records from northernmost point in Chile; this genus includes the unique lianas of Brassicaceae in New World.

44. Menonvillea DC. Annuals or perennial, often stoloniferous or rhizomatous, herbs or subshrubs; basal leaves rosulate or not. 24 spp. along Chile/Argentina border from Atacama to Patagnia, and vast areas in coastal Chile.

45. Yunkia Salariato & Al-Shehbaz. (off Cremolobus) Perennial herbs or scandent vines; stems to 3m tall, erect, somewhat woody basally, herbaceous elsewhere, glabrous to sparsely puberulent with minute trichomes, or densely pubescent with conspicuous ones. Two spp from Bolivia and NW Argentina, restricted of yungas vegetation.

4.3 EUDEMEAE ▸ TRIBE SCHIZOPETALEAE (4/19) - all genera in South America.

46. Atacama Hook. & Arn. Only one sp., A. nivea (Phil.) O. Toro, Mort & Al-Shehbaz, endemic to central Chile up to S Antofagasta province.

47. Machaerophorus Schlechtendal. Herbs suffrutescent perennial, or subshrubs, tomentose, becoming puberulent or glabrescent with age; stems erect to ascending, branched, flexuous, woody below, leafy, unarmed; basal leaves not developed; cauline leaves petiolate, not auriculate at base, finely pinnatisect into narrowly linear lobes, usually grooved abaxially and densely pubescent along groves, fleshy, entire. Three spp., endemics to southern Peru, in Arequipa and Puno regions.

48. Mathewsia Hook. & Arn. Perennial shrubs or small shrubs; cauline leaves present; petals yellowish. 7 spp., 5 endemic to N Chile, and two in S Peru, with species distributed from the Arequipa district of S Peru to the Valparaiso area in Chile.

49. Schizopetalon Sims. Annual pubescent herbs; cauline leaves petiolate. 10 spp., primarily in Chile, but S. rupestre (Barnéoud) Reiche also in Argentina.

5. MEGACORE HESPERODAE (46/c. 330) – six tribes, fully absent in South America: Anchonieae (9/66–68, Europe, temperate Asia), Buniadeae (1/3, S and E Europe, Mediterranean, W Asia), Chorisporeae (4/57, Europe, temperate Asia, northern North America), Dontostemoneae (2/19, SE Europe to Siberia and E Asia), Euclidieae (28/165, Europe, temperate and arctic Asia, arctic and alpine North America), Hesperideae (2/c 30, Europe, temperate Asia), Shehbazieae (▇▇).

45. SANTALALES

FAMILIES ABSENTS IN SOUTH AMERICA: MYSTROPETALACEAE (3/3) AND OCTOKNEMACEAE (1/14).

LINEAGE 1 of 8: STROMBOSIACEAE

STROMBOSIACEAE

§ FULLY CHLOROTROPHIC

Genera/species 4/c. 40 Distribution Tropical W Africa, E Himalayas to W Malesia, tropical America. Habit usually bisexual (androdioecious in Erythropalum), usually evergreen trees or shrubs (in Erythropalum liana climbing by axillary branch tendrils). At least Erythropalum and Heisteria are autotrophic (haustoria absent).

SYSTEMATIC outsiders Scorodocarpus (1; W Malesia from Peninsular Thailand to Borneo), Strombosia (9–10; tropical Africa, India, Sri Lanka, Burma, SE Asia, Malesia to Philippines an northern Moluccas), Diogoa (2; tropical Africa), Engomegoma (1; Gabon), Strombosiopsis (3; tropical Africa).

1. Tetrastylidium Engl. Non parasitic plants. Two spp. of South America, T. grandifolium (Baill.) Sleumer in E Brazil (Atlantic Forest), and T. peruvianum Sleumer in Amazon rainforest of Peru, Brazil and Colombia.

LINEAGE 2 of 8: ERYTHROPALACEAE

ERYTHROPALACEAE

§ FULLY CHLOROTROPHIC

Genera/species 4/c 20 Distribution tropical Africa, India, Sri Lanka, Burma to Central Malesia, southern Brazil. Habit usually bisexual (rarely dioecious), evergreen or deciduous trees or shrubs. At least Scorodocarpus and Strombosia autotrophic (haustoria absent).

SYSTEMATIC outsiders Erythropalum (1; Himalayas to Sulawesi and Java).

1. Brachynema Benth. Non parasitic plants. Two spp., B. axillare R. Duno & P.E. Berry from S Venezuela (the base of Mount Neblina) and Amazonian rainforest in N Brazil, and B. ramiflorum Benth. from central Amazonian rainforest of Brazil up to E Peru.

2. Heisteria Jacq. Non parasitic plants. 37 spp., from Mexico to Bolivia, 33 in South America, 22 in Brazil (8 endemics, one of then a rare species in Brazil by Plantas Raras do Brasil’s book, from N Amazonas state, uncollected until 1940), Caribbean, and three species in Africa.

3. Maburea Maas. Non parasitic plants. Only one sp., M. trinervis Maas, endemic to the Guiana Shield of Guyana, 0-100 m elevation range.

LINEAGE 3 of 8: XIMENIACEAE/OLACACEAE/OKTONEMATACEAE

XIMENIACEAE

§ PARASITIC (Prosopanche – ... – APODANTHACEAE – SANTALALES – Lennoa – OROBANCHACEAE - Cuscuta)

Genera/species 4/13 Distribution pantropical, southern Africa, southern China. Habit usually bisexual (in one species of Ximenia often functionally unisexual), evergreen trees or shrubs (species of Ximenia). At least Ximenia are root hemiparasites (probably also Curupira, Douradoa and Malania).

SYSTEMATIC outsider Malania (1; W Guangxi, E Yunnan).

1. Curupira G.A.Black. Medium sized tree (up to 12 m), possibly non parasitic, inflorescences of umbels subtented by a whorl of small, round tipped bracts. Only one sp., C. tefeensis G.A.Black, from wet soils of small area in Amazonas states of Brazil, and from Colombia (Caquetá) to Peru.

2. Douradoa Sleumer. Large trees with vanation pinnate, possibly non parasitic. Only one sp., D. consimilis Sleumer, from Brazil in Amapá and Pará states, possibly in Acre.

3. Ximenia L. Parasitic shrubs. 7 spp., two only in Mexico, two only in Caribbean, X. intermedia (Chodat & Hassl.) DeFilipps in Brazil and Cono Sur, X. coriacea Engl. endemic to Brazil, and the widely distributed X. americana L. from over tropical America, tropical and southern Africa, tropical Asia and Australia; some representatives are xerophytes; axillary shoots often modified into spines.

APTANDRACEAE

§ FULLY CHLOROTROPHIC

Genera/species 8/33 Distribution South America, Costa Rica, Honduras, Madagascar, SE Asia Habit usually bisexual (Harmandia and Hondurodendron dioecious), evergreen or deciduous trees or shrubs. At least the majority probably root hemiparasites. Some specis are xerophytes.

SYSTEMATIC two tribes, both in South America.

1.1 TRIBE APTANDREAE (5/10) ‣ outsiders Harmandia (1; Indochina, the Malay Peninsula), Hondurodendron (1; Honduras), Ongokea (1; tropical W Africa).

1. Aptandra Miers. Trees non parasitic with thin, elongate-elliptical alternate leaves with acute apices; inflorescence terminal or axillary, simple or branched panicles; flowers bisexual, with very small, 4-toothed, calyculus becoming large and funnel-shaped or urceolate around the fruit at maturity; petals 4, fleshy, linear to tongue-shaped, recurving at anthesis. 4 spp., one in W Africa and three in tropical South America (Colombia to Bolivia, N Brazil (2, no endemics), Venezuela and Guianas.

2. Chaunochiton Benth. Small to moderate trees, non parasitic; leaves alternate, pinnately veined, petiolate, glabrous; inflorescence an axillary, short-pedunculate, corymb-like panicle with few to many flowers; flowers fragrant, with small, cupulate, 5-dentate calyx much enlarged in fruit; petals 5, distinct, linear-elongate, most of the adaxial surface pilose. Three spp., from NW South America (2 in Brazil, non endemics) to Costa Rica, Guianas and Bolivia; most forests.

1.2 TRIBE ANACOLOSEAE (3/24) ‣ outsiders Anacolosa (16; tropical Africa, Madagascar, tropical Asia), Phanerodiscus (2; Madagascar).

3. Cathedra Miers. Non parasitic plants. 6 spp. from Brazil (5 endemics, one endemic to center Rio de Janeiro state is a rare plant in Brazil, by Plantas Raras do Brasil’s book), with C. acuminata (Benth.) Miers up to Colombia, Peru, Venezuela and Guianas.

LINEAGE 4 of 8: COULACEAE

COULACEAE

§ FULLY CHLOROTROPHIC

Genera/species 3/3 Distribution tropical W Africa, W Malesia, northern tropical South America. Habit bisexual, evergreen trees. Haustoria have never been observed and the three genera are probably non-parasitic.

SYSTEMATIC outsiders Coula (1; tropical W Africa), Ochanostachys (1; W Malesia).

1. Minquartia Aubl. Non parasitic trees. Only one sp., M. guianensis Aubl., from Costa Rica to Bolivia and Brazil (only in Amazon rainforest).

LINEAGE 5 of 8: OLACACEAE

OLACACEAE

§ PARASITIC (Prosopanche – Cassytha - ... – APODANTHACEAE – SANTALALES – Lennoa – OROBANCHACEAE - Cuscuta)

Genera/species 2–3/45–50 Distribution tropical and southern Africa, Madagascar, Indomalesia, Australia, New Caledonia;tropical South America. Habit usually bisexual (in Olax rarely monoecious or dioecious), evergreen trees or shrubs (rarely lianas). Olax (including ‘Dulacia’?) and Ptychopetalum are root hemiparasites. Some species are xerophytes. Branches in Olax sometimes strongly quadrangular (with wing-like edges).

SYSTEMATIC outsider Olax (40–45; tropical and S Africa, Madagascar, tropical Asia, Australia, New Caledonia; probably incl. Dulacia).

1. Dulacia Vell. Parasitic. 13 spp. in Colombia, Peru, Brazil (10, 4 endemics), Venezuela and Guianas; branches sometimes strongly quadrangular.

2. Ptychopetalum Benth. Parasitic. 4 spp., two in Africa and two from N Brazil (both, one endemic) and S Guyana.

LINEAGE 6 of 8: BALANOPHORACEAE

BALANOPHORACEAE

§ PARASITIC (Prosopanche – Cassytha - ... – APODANTHACEAE – SANTALALES – Lennoa – OROBANCHACEAE - Cuscuta)

Genera/species 13/c. 43 Distribution Caribbean, Mexico to Chile and Brazil, W Africa to South Africa, Madagascar, Comores, New Caledonia; New Zealand; Himalaya, Vietnam, W Malesia; the Malay Peninsula; tropical Asia, Malesia, islands in the Pacific, tropical NE Australia. Habit monoecious or dioecious, perennial, whitish, yellow, brown or red, achlorophyllous herbaceous root holoendoparasites with branched or simple subterranean tuber-like structures partly of root nature, partly consisting of host tissue. Succulents.

A poorly known family in terms of distribution, pollination, and number of species, mostly due to their partially or entirely subterranean habit. Some reports of plants visited by flies, bees and wasps, but no definitive studies about their pollination biology. Also, some reports about seed dispersal by ants. Their general physiology is also little-known, as is a family without stomata; very distinctive from other parasitic taxa, by its set of unique characters; the other family of subterranean holoparasitic root parasites with subterranean habit is the Hydnoraceae, which is a characterized by a large, fleshy, solitary flower, with 3-4 fleshy tepals, and a large fruit with numerous seeds.

6 genera and 13 spp. in Brazil, 19 in over New World.

Key to genera of Neotropical Balanophoraceae

1. Style 1; tubers with wax; flowers not embedded in filiform hairs, or anthers 3, merged into a synandrium ------------ Langsdorffia

1. Styles 2; tubers with starch; flowers with both sexes imbedded in filiform hairs, or anthers 2, not merged into a synandrium - 2

2. Flowers embedded in a layer of filiform hairs; anthers merged into a usually 3-merous synandrium - 3

3. Stem and young inflorescence covered by triangular scales ------------ Scybalium

3. Stem naked, or with a few inconspicuous bracts; young inflorescence covered by hexagonal, peltate bracts - 4

4. Inflorescence emerging from buds on elongated, rhizome-like structures, ------------ Helosis

4. Inflorescence emerging directly from a ‘tuber’ connected with the host root ------------ Corynaea

2. Flowers not imbedded in hairs, on conspicuous or elongated branches, these subtended by deciduous scale-like bracts; stamens with filaments - 5

5. Inflorescence branches subtended by ovate -triangular bracts; apical part of female branch not peltately enlarged ------------ Lophophytum

5. Inflorescence branches subtended by hexagonal or irregularly-shaped bracts; apical part of female branch peltately enlarged - 6

6. Male flowers solitary, each flower subtended by a peltate bract; anthers sessile ------------ Lathrophytum

6. Male flowers in many-flowered branches, these subtended by a peltate or clavate bract ------------ Ombrophytum

SYSTEMATIC Sarcophyteae (2/2, tropical and S Africa) does not occur in South America; MYSTRAPETALACEAE (3/3; SW South Africa, North Island of New Zealand, mountains on New Caledonia) emancipated as family before APG V.

1. SUBFAMILY HELOSIDEAE (6–7/17) ‣ outsiders Ditepalanthus (2; Madagascar), Exorhopala (1; NW Malay Peninsula)

1. Corynaea Hook.f. Only one sp., C. crassa Hook. f., from Panamá, Colombia, Venezuela, Ecuador, Peru, and Bolivia.

2. Helosis Rich. Three spp., one in NW portion of the Malay Peninsula, another only in Caribbean, and H. cayenensis (Sw.) Spreng. from from Mexico, Central America, Greater Antilles, Colombia, Ecuador, Peru, Bolivia, Venezuela, Guianas, throughout S Brazil.

3. Ombrophytum Poepp. ex Endl. 7 spp., two in dry areas from SW Bolivia, NW Argentina and N Chile, French Guiana and Colombia one endemic each, three from Ecuador, Peru to Amazon rainforest of Acre and Amazonas states in Brazil (all three).

4. Scybalium Schott & Endl. 4 spp., one endemic to Jamaica, another from Colombia, Venezuela and Ecuador, and two remaining endemics to forests of SE Brazil.

2. SUBFAMILY BALANOPHOREAE (5/22–24) ‣ outsiders Rhopalocnemis (1; India, the Himalayas, S China, Thailand, Vietnam, W Malesia), Balanophora (17–19; Congo, Madagascar, the Comoros, tropical Asia to S Japan, Malesia, tropical NE Australia and islands in the Pacific), Thonningia (1; tropical Africa from Senegal to SW Ethiopia and south to Zambia).

5. Langsdorffia Mart. 4 spp. highly disjuntcs: one in Madagascar, another in Papua New Guinea, L. heterotepala is L. J. T. Cardoso, R. J. V. Alves & J. M. A. Braga known only from cloud forests of mountain slopes above 1,500 m at Itatiaia Range in SE Brazil, and L. hypogaea Mart. over widely in Central and South America up to Brazil.

6. Lathrophytum Eichl. Only one sp., L. peckoltii Eichler, in SE Brazil from Goiás to Rio de Janeiro states.

7. Lophophytum Schott & Endl. 4 spp. from N Colombia, E Peru and NW Acre state in Brazil (all 4), Peru, Bolivia, Paraguay, northern Argentina and S & SE Brazil, with L. rizzoi Delprete a rare species in Brazil by Plantas Raras do Brasil’s book, collected only two times in Goiás state.

LINEAGE 7 of 8: LORANTHIDS

MISODENDRACEAE

§ PARASITIC (Prosopanche – Cassytha - ... – APODANTHACEAE – SANTALALES – Lennoa – OROBANCHACEAE - Cuscuta)

Genera/species 1/8 Distribution southern South America. Habit hemiparasitic aerial shrubs.

SYSTEMATIC a single genus.

1. Misodendrum Banks ex. DC. Evergreen shrubs hemiparasites, usually dioecious (rarely bisexual or monoecious), with distinct sympodial growth and even as young a thick stem, almost exclusively on Nothofagus. 8 spp., cold-temperate South America (S Chile and adjacent parts of Argentina) from Magellan’s Strait northwards to c 33º S.

SCHOEPFIACEAE

§ PARASITIC (Prosopanche – Cassytha - ... – APODANTHACEAE – SANTALALES – Lennoa – OROBANCHACEAE - Cuscuta)

Genera/species 3/34. Distribution southeastern China, Hainan, southern Japan, the Korean Peninsula, Taiwan, Indochina, northern Sumatra, Mexico, Central America, the West Indies, tropical and temperate South America. Habit usually bisexual (sometimes unisexual), evergreen trees or shrubs, or perennial herbs (many species of Quinchamalium). Root hemiparasites. Use Edible tubers in Arjona.

SYSTEMATIC all genera in South America.

1. Arjona Cav. Shrubs non parasitics, with branched underground rhizomes or with underground swollen knotty runners, rarely woody below subshrub; leaves alternate, linear or lanceolate, usually rigid and pointed; flowers rather numerous sitting crowded in short terminal spikes or only few axillary, white, lilac or crimson. 6 spp., 4 of tropical & temperate S. America from Bolivia to Tierra del Fuego, Uruguay; and two in Peru and Brazil, one endemic each; Brazilian species is A. megapotamica Malme, known only from high mountains of Santa Catarina and Rio Grande do Sul states; most species are found in dry open areas in rocky and sandy soils, sometimes found in humid places such as wet meadows.

2. Quinchamalium Molina. Low, half parasitic herbs or herbaceous plants, often with plumper primary roots, small stems usually branch out from the base, branches often prostrate; flowers bisexual, 5-merous, in terminal dense racemes often numerous from the receptacle over the cupular ‘calyx’ (Becherkelch) dropping or drying persistent. Only one sp., Q. chilense Molina from southern Chile to central Peru, W Bolivia, N Argentina.

3. Schoepfia Schreber. 27 spp., 4 in S Japan and Korean Peninsula, SE China (inc. Taiwan), Hainan, Indochina, N Sumatra and tropical Africa, and 23 in Mexico, Caribbean, SE U.S.A., Central and South America (8), three in Brazil, S. velutina Sandwith endemic, a rare plant in Brazil by Plantas Raras do Brasil’s book, known only type collection from 1927 in Diamantino region, Mato Grosso state.

LORANTHACEAE

§ PARASITIC (Prosopanche – Cassytha - ... – APODANTHACEAE – SANTALALES – Lennoa – OROBANCHACEAE - Cuscuta)

Genera/species 78/c. 1,035 Distribution tropical and subtropical regions, with their largest diversity in the Southern Hemisphere, and a few species in temperate parts in the Northern or Southern Hemispheres (SE Europe, Japan, New Zealand). Habit usually bisexual (rarely dioecious; in Nuytsia monoecious), usually evergreen shrubs (sometimes lianas; Atkinsonia, Gaiadendron and Nuytsia are root parasitic trees) with distinctly sympodial growth. Usually hemiparasites (rarely leafless holoparasites) and stem parasites. Some species are hyperparasites on stem hemiparasitic shrubs. Branches rarely transformed into photosynthezising phyllocladia. Shoots rarely resembling Cuscuta in appearance.

Cacti appear to be especially resistant to attack by mistletoes; they are parasitized by only two species of mistletoe, Tristerix aphyllus (Miers ex DC.) Barlow & Wiens and Ligaria cuneifolia (Ruiz & Pav.) Tiegh. (both Loranthaceae), even though cacti often occur among parasitized trees and shrubs.

SYSTEMATIC genera Nuytsia (1; W Australia) and Atkinsonia (1; E Australia) are the sussecively most basal and do not occur in South America.

1. CLADE GAIADENDREAE (1/3) ‣ a single genus.

1. Gaiadendron G. Don. Shrub or an aerial parasite, stoloniferous, often epiphytic on trees while parasitizing others epiphytes, variable plant, with showy yellow flowers, often white, inflorescences terminal or axillary; leaves paired, shiny, lanceolate to ovate. Three spp., Venezuela and Peru one endemic each, and G. punctatum (R. & P.) G. Don., from Nicaragua to Bolivia, with isolated stations in Mount Roraima in Brazil (Roraima state), Venezuela and Guyana.

2. SUBFAMILY LORANTHOIDEAE (c 71/895–915) ‣ three tribes, Elytrantheae (14/140–145, India to New Zealand) and Lorantheae (c 40/470–480, tropical regions in the Old World) do not occur in New World; among Psittacantheae, four tribes, Tupeinae is monotipic and endemic to New Zealand.

∎ TRIBE PSITTACANTHEAE/ SUBTRIBE NOTANTHERINAE ‣ both genera in South America.

2. Desmaria Tiegh. Large mistletoes with epicortical roots sprounding new roots; showy flowers yellow to red. Only one sp., D. mutabilis (Poepp. & Endl.) Tiegh. ex Jacks, endemic to S Chile, perhaps exclusively in Nothofagus.

3. Notanthera (DC) G. Don. f. Dense leafy plants, with persistent, papillate hairs on young growth; leaves shiny, dark green; inflorescences clustered near shoot ends, one per leaf axil or as terminal one; flower bud white, with bight pink petals, six. Only one sp., N. heterophylla (R. & P) G. Don, endemic to S Chile.

∎ TRIBE PSITTACANTHEAE/ SUBTRIBE LIGARINAE ‣ both genera in South America.

4. Ligaria Tiegh. Robust large plants, showy flowers, stems rigid, single haustorial connection; isolate pendunculate inflorescences, axillary, single flowered or in small clusters, pendunculate, brilliantly red flowered. 4 spp., two endemics to Peru, L. cuneifolia (Ruiz & Pav.) Tiegh. from arid and subtemperate C Peru to C Chile up to S Brazil, Uruguay, and L. teretiflora (Rizzini) Kuijt endemic from Bahia and Minas Gerais states in E Brazil.

Only L. cuneifolia populations in Peru and Tristerix aphyllum in Chile infects Cactaceae (Corryocactus and Trichocereus in Peru /Trichocereus and Eulychnia in Chile, respectively).

5. Tristerix Mart. Glabrous to nerly leafless; inflorescense terminal, racemose or spike-like; showy flowers, inderteminate inflorescence, terminal raceme, suportade by leafy bracts. 13 spp., from subparamo elevations in N Colombia to low elevation and adjacent areas of the Andes of Argentina and south-central Chile, 4 endemics to Peru, Chile and Colombia one endemic each.

The chilean endemic, T. aphyllus (Miers ex DC.) Barlow & Wiens, parasitizes two species of cactus (unique in cacti, with Ligaria cuneifolia populations in Peru, which infects Corryocactus and Trichocereus), Echinopsis chilensis and Eulychnia acida; this species has been reported to be a nonphotosynthetic holoparasite (unique among Loranthaceae), however, the presence of green tissues in seedling radicles suggest that chlorophyll is present.

∎ TRIBE PSITTACANTHEAE/ SUBTRIBE PSITTACANTHINAE ‣ outsiders Dendropemon (c 40; Caribbean, with their highest diversity on Hispaniola), Panamanthus (1; Panamá).

6. Aetanthus (Eichl.) Engl. 18 spp. from the N Colombia and adjacent Venezuela to N Peru, limited to high elevations.

7. Cladocolea Tiegh. (exc. Peristhetium p.p.) Erect or pendulous epiphytes, glabrous or lightly pilose, epicortical roots present or absent, dioecious or monoecious. 27 spp., mainly in Mexico, others few recorded for Panamá, and Andean Ecuador and adjacent Peru; 8 spp. in South America, 3 in Brazil, two endemics (narrow endemics in Amazonas and Rio de Janeiro states) and C. micrantha (Eichler) Kuijt occur in over northern Amazon rainforest.

8. Maracanthus Kuijt (inc. Oryctina p.p.). Oryctina–like, differing only by floral structures, glabrous or partly furfuraceos plants, dioecious or with bisexual flowers; leaves paired, inflorescences simples, terminal or axillary spikes; flowers hexamerous, petals dimorphic. Three spp., two from Colombia and Venezuela and one endemic to Osa peninsula, Costa Rica.

9. Oryctanthus (Griseb.) Eichler. Leafy plants, succulent spical into monodes, phyllotaxy decussate, leaves leathery; inflorescences solitary or axillary where clustered at the nodes; flowers bisexusl, usually hexamerous, yellow to dark red; fruit several coloured. 14 spp., from Mexico to South America (13 in continent), one also in Caribbean; 4 spp. in Brazil, none endemics, mainly Amazon rainforest, up Mato Grosso, Tocantins and Piauí states.

10. Oryctina Tiegh. (exc. Maracanthus p.p., Passovia p.p.) Aereal parasitic shrubs to moderate size, fragile, stems cylindrical or quadrangular; inflorescence spical, inside monades; leaves minute to large, decussate; flowers hexamerous, with two bracteoles; stamens placed in two series; high delicate. Three spp. growing at trees of open places in dry savannah of center Brazil, in NW Minas Gerais, N Goiás, S Tocantins and W Bahia; one of then is a rare plant in Brazil, by Plantas Raras do Brasil’s book, collected in N Minas Geais state, above Aspidosperma (Apocynaceae).

11. Passovia Mart. (inc. Oryctina p.p.). 25 spp. distributed from Central and South America (23), up to state of Rio de Janeiro, in SE Brazil (16 in Brazil, 6 endemics, 4 of then rare by Plantas Raras do Brasil’s book, all formerly in Phthirusa).

12. Peristethium Tiegh. (inc. Cladocolea p.p., Struthanthus p.p.) Aerial hemiparasitic shrubs with inflorescence dioecious enclosed by dioecious bracts. 17 spp., one in Costa Rica and Panamá, and 15 spp. of South America, mainly from Ecuador and Peru, with two rare, narrow endemics on Mount Roraima and the Pakaraima Mountains; three spp. in Brazil, in Amazon rainforests up to ecotone with the central Brazilian savannas, P. reticulatum (Rizzini) Caires endemic.

13. Phthirusa Mart. Epiphytes glabrous, parasites on branches of dicotyledons, at times with diminutive feruginous scales, dioecious, sessile flowers. 11 spp., two in NE Brazil (Alagoas, Bahia and Rio de Janeiro), one of then disjunct in Guyana, and remaining nine in N Andes.

14. Psittacanthus Mart. Epiphytes, glabrous or pilose, parasites on the branches of dicotyledonous trees, connected to the host only by the initial haustorium, monoecious, flowers very showy. 115 spp., distributed from Baja California to Mexico to Argentina, Jamaica and Lesser Antilles, 87 in South America, 33 in Brazil (except southern region, 8 endemics, three of then in Amazon rainforests are rare in Brazil by Plantas Raras do Brasil’s book), in most diverse habitats.

P. longiflorus Kuijt, known only from Amazonas in Peru, may be confused with Aetanthus macranthus (Hook.) Kuijt because of its unusually long flowers (to 17 cm), the longest known for this genus; P. gigas Kuijt is a species endemic to Bajo Calima region in W Colombia, which represent one the largest simple leaves of Loranthaceae, with individual masses of tissue 1 m long and over 50 cm wide; P. nudus (Ant. Molina) Kuijt & Feuer from Mexico and Central America is the only aphyllous species in this genus.

15. Pusillanthus Kuijt. Aerial hemiparasitic shrubs stems Oryctina-like, equally high delicate. Only one sp., P. pubescens (Rizzini) Caires, highly disjunct restricted from dry areas of Venezuela, Guyana and NE Brazil, in dry areas of Pernambuco, Alagoas, Paraíba and Bahia states.

16. Struthanthus Mart. (exc. Peristhetium p.p.) Scandent, leafy, glabrous or pilose epiphytes, parasites on branches and stems of gymnosperms, dicotyledons and arborescent monocotyledons, dioecious. 101 species occurring from Mexico to Argentina, 75 in South America; 52 species in Brazil (41 endemics, 8 of then from Goiás to Rio de Janeiro states are rare in Brazil by Plantas Raras do Brasil’s book), occurring in all the states and ecosystems.

17. Tripodanthus (Eichl.) Tiegh. Epiphytic shrubs parasitic on branches or lianas and/or tree-like parasites of roots, monoecious; inflorescences terminal or axillary in racemes or corymbs; flowers small, ca. of 1.5-2.0 cm in diam., fleshy tepals, reflexed at anthesis, white or greenish-white, yellowish or red. Three spp., one endemic to Antioquia, Colombia, T. acutifolius (Ruiz & Pav.) Tiegh. from Paraguay to Ecuador, Venezuela, and S Brazil, and T. flagellaris (Cham. & Schltdl.) Tiegh. in Argentina, Uruguay and S Brazil.

LINEAGE 8 of 8: SANTALIDS

OPILIACEAE

§ PARASITIC (Prosopanche – Cassytha - ... – APODANTHACEAE – SANTALALES – Lennoa – OROBANCHACEAE - Cuscuta)

Genera/species 11/33. Habit usually bisexual (in Agonandra and Gjellerupia dioecious), evergreen trees and shrubs (sometimes lianas). Root hemiparasites in some Old World genera.

SYSTEMATIC - outsiders Gjellerupia (1; New Guinea); Anthobolus (3; C and N Australia); Champereia (1; Yunnan, Guangxi and Taiwan in China, Andaman and Nicobar Islands, Burma, Thailand, Vietnam, Malesia to Philippines and New Guinea), Melientha (1; Yunnan, SE Asia to Philippines), Yunnanopilia (1; Yunnan, Guangxi, Laos, N Vietnam); Lepionurus (1; Nepal to New Guinea), Urobotrya (7; tropical Africa, SE Asia, Malesia to Flores), Cansjera (3; India, Sri Lanka, Nepal, Andaman and Nicobar Islands, S China, S and SE Asia, Malesia to New Guinea, Solomon Islands, tropical Australia), Rhopalopilia (3; Central Africa), Pentarhopalopilia (4; tropical Central and coastal E Africa, Madagascar), Opilia (5; tropical Africa, Madagascar, India, Sri Lanka, Yunnan, SE Asia, Philippines, New Guinea, tropical Australia).

1. Agonandra Miers ex Benth. & Hook.f. Trees or shrubs, parasitics; leaves alternate, simple, margin entire; without stipules; petiolate; inflorescences axillary or cauliflorous, rarely terminal; flowers small, inconspicuous, actinomorphic, mainly bisexual, sometimes unisexual (plants then dioecious), 4-5, free or connate at the base; stamens 4-5, epipetalous; ovary superior, immersed in a lobed disk or diskless, unilocular; fruit drupaceous. 10 spp., five restricted of S Mexico and Central America, and five in South America, all in Brazil, mostly in Amazon rainforest and savannas, one restricted of S Brazil, remaining also in Panamá, Colombia, Venezuela, Guyana, Ecuador, Peru, Bolivia, Paraguay and Argentina.

SANTALACEAE

§ PARASITIC (Prosopanche – Cassytha - ... – APODANTHACEAE – SANTALALES – Lennoa – OROBANCHACEAE - Cuscuta)

Genera/species 42/c. 1,020 Distribution mostly tropical and subtropical regions, temperate parts in the Northern and Southern Hemispheres, with their largest diversity in semiarid climates. Habit bisexual, monoecious or dioecious, usually evergreen trees or shrubs, or perennial herbs, sometimes with distinct sympodial growth. Root or stem hemiparasites (Phacellaria and some species of Viscum are hyperparasites, parasitizing on Loranthaceae or Amphorogynaceae). Sometimes spiny or xeromorphic. Branches sometimes photosynthesizing phyllocladia in Viscum sp. from E Cape.

SYSTEMATIC seven clades, taken here as tribe level; Comandreae (2/2, North America, the Balkan Peninsula to Romania) and Amphorogyneae (9/54, South and SE Asia, Malesia to Australia, New Caledonia, with their largest diversity in Australia) do not occur in South America.

1.1 TRIBE NANODEOIDEAE (2/3) ‣ both genera in South America.

1. Mida L. Hemiparasitic trees or shrubs; leaves usually opposite, + leathery or fleshy, pinnately-nerved; small panicles axillary or terminal; bracts deciduous, small; flowers bisexual, 4-5-merous. Two spp., M. salicifolia A. Cunn. in New Zealand and M. fernandeziana (Phil.) Sprague et Summerhayes in the Juan Fernandez Is., in former times common on both islands, is now, however, completely exterminated; Skottsberg saw the last living specimen!

2. Nanodea Banks ex Gaertner f. Small herb with creeping, branched stems; leaves alternate, narrowly linear; flowers small, to three between the highest leaves, the lateral in the axil of flabellate bracts. Only one species, N. muscosa Gaertn. f., in Andean Patagonia, Tierra del Fuego, Staten-Islands, Argentina and Falkland-Islands.

1.2 TRIBE SANTALEAE (EREMOLEPIDACEAE, 11/61) ‣ outsiders Omphacomeria (1; New South Wales, Victoria, Tasmania), Exocarpos (27; SE Asia, Malesia to New Guinea, Australia, New Caledonia, Hawaii), Santalum (12; tropical Asia, Australia to Hawaii), Osyris (3; Mediterranean, Africa, SW Asia to India), Nestronia (1; SE U.S.A.), Colpoon (6–7; Africa to India), Rhoiacarpos (1; W Cape, E Cape, KwaZulu-Natal).

3. Antidaphne Poeppig & Endlicher. (inc. Ixidium) Erect shrubs, with epicortical roots recorded as regularly emitting haustoria, monoecious or dioecious, mite flowers, the smallest in Santalaceae. 10 spp., 7 strictly South American, one endemic to the Caribbean, two up to Mexico and Central America; 4 spp. in Brazil, three endemics, two of then are rare in Brazil by Plantas Raras do Brasil’s book, A. amazonensis Rizzini collected in Byrsonima (Malpighiaceae) in Manaus region, and A. schotii (Eichler) Kuijt from Rio de Janeiro municipality, uncollected more than 135 years.

4. Eubrachion Hooker f. Erect shrubs, without epicortical roots, glabrous, monoecious; stems and branches green-yellowish, cylindrical. Two spp., E. gracile Kuijt in Venezuela and Ecuador and E.ambiguum (Hook & Arm.) Engl. in Jamaica, Dominican Republic, Puerto Rico, Venezuela, Argentina, Uruguay and Brazil.

5. Lepidoceras Hooker f. Two spp., Chile and Peru one endemic each.

6. Myoschilos Ruiz & Pav. Rather richly branched, glabrous shrub; leaves alternate, oblong oval, membranous, entire; flowers in catkin-like racemes, which stand solitary or by twos at the nodes of the previous-year’s branches. Only one sp., M. oblongus Ruiz et Pav., in C and S Chile and adjacent Argentina; the leaves used as a light purgative.

1.3 TRIBE VISCACEAE (7/c 520) - outsiders Korthalsella (11–15; NE and E Africa, Madagascar, Mascarene Islands, the Himalayas, E Asia to Japan, New Zealand, Hawaii), Arceuthobium (40; Mediterranean, NE tropical Africa, Himalayas, China, SE Asia, W Malesia, North America, Central America, Caribbean), Notothixos (8; Sri Lanka, Burma, SE Asia, Malesia to Philippines and SE Australia, the Santa Cruz Islands), Viscum (c 75; temperate to tropical regions in the Old World), Ginalloa (9; tropical Asia).

7. Dendrophthora Eichl. Shrubs glabrous or pilose, dioecious or monoecious; leaves absent to normal; flowers in axillary spikes. 129 spp. from S Mexico, including Cuba and the Caribbean, South America to Bolivia and the SE Brazil, 83 in South America, only three in Brazil, D. fendleriana (Eichler) Kuijt (Venezuela and N Brazil, in Neblina Massif), D. elliptica (Gardner) Krug & Urb. and D. warmingii (Eichler) Kuijt in Brazil, none endemics.

8. Phoradendron L. Shrubs glabrous or pilose, dioecious or monoecious; leaves absent to normal; spikes axillary; over Neotropics up to Argentina; it’s extremely similar to Dendrophthora, the only difference being the anthers. 271 spp. of New World, 186 spp. in South America, only 41 spp. in Brazil (11 endemics, 5 of then are rare in Brazil by Plantas Raras do Brasil’s book).

Parasitic plants with sprouted from a single, interconnected endophyte growing under thunks of other plants are known isophasic parasitism, known in Apodanthaceae, Mitrastemaceae and Arceuthobium (Santalaceae, North America and Asia) and P. perredactum Rzedowski Calderon (known only Oaxaca state, S Mexico, in Bursera trees); anothers aphyllous spp. are P. aphylla Steyerm. from Venezuela and P. fragile Urb. of Brazil, but they aren’t isophasics.

1.4 TRIBE THESIOIDEAE (3/c 350) ‣ outsiders Buckleya (4; China, Japan; SE U.S.A.), Osyridicarpos (1; tropical and S Africa).

9. Thesium L. Perennial herbs with + woody, branched basic axis or subshrubs to small shrubs; leaves very small scale-like or linear, very rarely broader and large. Inflorescences simple racemose or spicate or from 3- more flowered compound cyme, sometimes at the junction of the petiole and the leaf blade (true epiphylly). 326 spp. in Europe, Africa, Asia, Australia, highly centered in southern Africa, and 3 spp. in New World (belonging a own sect., Psilothesium), T. tepuiense Steyerm. in Venezuela and Guyana, T. aphyllum Mart. ex A. DC. and T. brasiliense A. DC. endemics to C & S Brazil, one of the more notably disjunctions of South America and Old World.

T. aphyllum, with strongly woody rootstock; stem to a foot high, rigid, sharp-edged, weakly branching out; leaves scattered, diminutive; flowers in spikes, with 2 prophylls, very small - related Thesium brasiliense.

1.5 TRIBE CERVANTESIOIDEAE (8/c 21) ‣ outsiders Pyrularia (2; Himalayas, China, U.S.A.), Pilgerina (1; Madagascar), Staufferia (1; Madagascar), Scleropyrum (c 6; tropical Asia), Okoubaka (2; tropical Africa).

10. Acanthosyris (Eichl.) Griseb. Trees or shrubs, frequently with leaf axillary thorns; leaves on short shoots clustered, on young long shoots slightly spiral; flowers in small 3-5-flowered cymes, bisexual, 4-5-merous. 6 spp., Central America to Argentina and Uruguay (center of diversity), 4 in Brazil, A. paulo-alvinii G. M. Barroso endemic; the wood of some species ‘quebrachillo’ or ‘sombra del toro hembra’ is mentioned by cabinet makers etc.

11. Cervantesia Ruiz & Pavon. Small trees, non parasitic; leaves entire, oval to elliptic, on the top side weakly, on the lower surface pubescent, usually closely; young branches and inflorescences also closely pubescent; flowers in small balls at short, spike-like, to 1-2 axillary or at the end of the branches paniculate united inflorescences. Three spp., mountains from Colombia to Bolivia.

12. Jodina Hook. & Arn. ex Meissner. Shrub or tree, non parasitics; leaves alternate, glabrous, rhombic, at the corners spiny on the edges; flowers pubescent, in close, short bundles in the leaf axils. Only one sp., J. rhombifolia Hook. et Arn., in S Brazil, Bolivia, Uruguay and Argentina; leaves 3-6 cm long, rhomboid.

46. BERBERIDOPSIDALES

TWO FAMILIES, BOTH IN SOUTH AMERICA.

AEXTOXICACEAE

§ FULLY CHLOROTROPHIC

Genera/species 1/1 Distribution southern and Central Chile and adjacent parts of Argentina. Habit dioecious, evergreen tree.

SYSTEMATIC a single genus.

1. Aextoxicon Ruiz & Pav. Evergreen tree. Only one dioecious, A. punctatum Ruiz & Pav., from southern and Central Chile and adjacent parts of Argentina.

BERBERIDOPSIDACEAE

§ FULLY CHLOROTROPHIC

Genera/species 1 or 2/3 Distribution C Chile, E Australia. Habit bisexual, evergreen climbing shrubs or lianas.

SYSTEMATIC outsider Streptothamnus (1, Queensland, New South Wales).

1. Berberidopsis Hook. f. Bisexual, evergreen climbing shrubs or lianas. Two spp., B. corallina Hook. f. in a small mountain area in coastal southern Chile, and B. beckleri (F.Muell.) Veldkamp in easternmost point of Australia.

47. CARYOPHYLLALES

FAMILIES ABSENTS IN SOUTH AMERICA: ANCISTROCLADACEAE (1/20), ASTEROPEIACEAE (1/8), BARBEUIACEAE (1/1), DIDIEREACEAE (6/20), DIONCOPHYLLACEAE (3/3), DROSOPHYLLACEAE (1/1), GISEKIACEAE (1/7), KEWACEAE (1/8), LIMEACEAE (1/25), LOPHIOCARPACEAE (2/7), MACARTHURIACEAE (1/9), NEPENTHACEAE (1/187), PHYSENACEAE (1/2), SARCOBATACEAE (1/2), SIMMONDSIACEAE (1/1), STEGNOSPERMATACEAE (1/4) AND TAMARICACEAE (4/12).

LINEAGE 1: POLYGONOIDES

DROSERACEAE

§ CARNIVOROUS (Brocchnia – Catopsis - Paepalanthus - Drosera - Heliamphora - Philcoxia - Genlisea - Utricularia - Pinguincula)

Genera/species 3/254 Distribution cosmopolitan except polar and arid regions, with their largest species diversity in Australia and New Zealand. Habit bisexual, perennial or annual herbs (in Drosera sometimes climbing). Corm or tuberous rhizome present in some species of Drosera. Most species are hygrophytes; Aldrovanda is aquatic with submersed leaves. Carnivorous; Aldrovanda: ‘snaptraps’ with c. 20 trigger hairs per foliar lobe; Dionaea: ‘snaptraps’ with three trigger hairs per foliar lobe; Drosera: ‘flypaper traps’.

One genus in Neotropics. Aldovandra is a Old world rootless aquatic; the whorls of leaves are lobed as in the vênus fly trap with small trigger hairs allowing the fastest known plant movement known (0.01-0.02 sec).

SYSTEMATICS outsiders Dionaea (1; North and South Carolina), Aldrovanda (1; central and E Europe, Africa, Asia to Queensland).

1. Drosera L. Bisexual, perennial or annual herbs, sometimes climbers; modified, insectivorous leaves with enzyme-secreting, tentacle-like glandular trichomes; adapted to grow in wet areas with very few available nutrients - such as bogs and swamps. c. 180 spp., cosmopolitan genus, found from sea-level to altitudes over 3,000 m throughout the Neotropics, exceptionally diverse in SW Australia, where there are about one third of the species in the whole genus; 47 spp. in New World, 31 in Brazil, 21 endemics, centered in Guiana Shield and rocky grasslands (campos rupestres) in Espinhaço Range of Brazil, few species that occur in lowlands are mostly found in sandy coastal habitats known as restingas and only a few of them occur in inland lowland habitats, including D. capillaris Poir. in N Brazil, D. cayennensis Sagot ex Diels in W and N Brazil, and D. sessilifolia A. St.-Hil. which is abundant in C, W, NW, and the N tip of Brazil; found from sea-level to altitudes over 3,000 m throughout the Neotropics. Except for D. regia Stephens and D. arcturi Hook., two large clades emerges:

§ subg. Drosera ‣ contains the generic type, D. rotundifolia L., with 8 sections, 4 centered in Australia (Arachnopus, Stelogyne, Prolifera and Psychophila), and four with/or also Neotropical and African distribution.

§ sect Thelocalyx ‣ two spp. highly disjuncts, one in Australia and D. sessiliflora A. St-Hill. in northern South America inc. Brazil.

§ sect. Brasiliae ‣ 18 spp., all endemics to Brazil except D. montana A.St.-Hil. up to Peru and Bolivia. D. magnifica Rivadavia & Gonella is a species of sundew endemic to Pico Padre Ângelo (1,500–1,530 m) in E Minas Gerais in SE Brazil, where it grows among sandstone outcrops in herbaceous and shrubby vegetation; it is one of the third largest species of Drosera, and largest of New World - the other two being D. regia from South Africa and D. gigantea Lindl. from Australia - and was discovered in 2015 through images which appeared on the social network Facebook. Drosera magnifica has the distinction of being the first plant species to be discovered through images posted in a social media group; D. graomogolensis T. Silva is a rare plant in Brazil, by Plantas Raras do Brasil’s book.

§ sect. Drosera ‣ temperate and diploid Neotropical taxa, as well as D. spatulata Labill and allied taxa; 26 spp. in New World, 12 in Brazil (three endemics), 5 in Brazil and other some places but confined to South America; 4 in Brazil up to Central and North America; 14 outside Brazil: 4 in Guiana Shield near Brazilian borders and possibly in Brazil, 5 exclusive from North America, D. uniflora Willd. in Cono Sur, three in high Andes from Venezuela, Ecuador and Peru, and D. moaensis Panfet endemic to Caribbean; belonging this clade, D. amazonica Rivadavia, A.Fleischm. & Vicent. is probably water-dispersed (dispersal mechanism would be unique among New World Drosera) either by flowing ground water of the open seepage habitats or by the heavy tropical rainfalls common to that area; the flowers of this species are sweetly perfumed, a character this species shares with at least four more New World species of Drosera, all of which are highland species from the Guiana Shield north of the Amazon rainforests.

§ sect. Pycnostigma ‣ all species belonging to the ‘African clade’, which comprises all Drosera species occurring on the African continent, with the exception of D. regia (D. subg. Regiae) and D. indica L. (D. sect. Arachnopus).

§ subg. Ergaleium ‣ ‘Australian Clade’, comprising D. sections Coelophylla, Lasiocephala, Bryastrum, Erythrorhiza, Stolonifera and Ergaleium, with a single South America species, D. meristocaulis Maguire & Wurdack from Venezuela side of Neblina Massif, near Brazilian border.

POLYGONACEAE

§ FULLY CHLOROTROPHIC

Genera/species 55/900-1,110. Distribution cosmopolitan, with most genera and species occurring in northern temperate regions. 31 genera are found in the western hemisphere. 16 of these genera are restricted to western North America, with 3 disjunct to Chile and Argentina. Habit usually bisexual (sometimes monoecious, polygamomonoecious or dioecious), usually perennial or annual herbs (often climbing or twining) or shrubs (rarely trees or lianas). Stem and branches often sulcate, geniculate, striate and/or hollow, often with swollen nodes (branches in Muehlenbeckia and Calligonum photosynthesizing phyllocladia).

While Coccoloba, Salta, Magoniella, Ruprechtia and Triplaris are endemic to the Neotropics, Rumex, Polygonum and Antigonon have weedy status, with species of knot-weed (Polygonum) being particularly invasive in Europe. Antigonon leptopus Hook. & Arn. originates from Mexico but is cultivated worldwide because of its showy flowers.

Traditionally the Polygonaceae were placed within the Caryophyllales on the basis of the unilocular ovary and single, basal ovule. However, recent studies have revealed that the family lacks the P-type sieve-tube plastids, anatropous ovules, betalain pigments and perisperm characters that characterise the order Caryophyllales. Recent studies by the APG II (2003) still include the Polygonaceae within the Caryophyllales.

SYSTEMATICS four lineages, basal clade Afrobrunnichia (2; tropical West Africa) does not occur in South America.

1. SUBFAMILY SYMMERIOIDEAE (1/1) ‣ a single species.

1. Symmeria Benth. Shrubs or small trees, dioecious; twigs solid, myrmecophyte. Only one sp., S. paniculata Benth., disjunct from northern South America over Amazon rainforest and tropical West Africa.

2. SUBFAMILY ERIOGONOIDEAE (23/600–630) ‣ 6 tribes, Brunnichieae (2/8, SE U.S.A., Mexico, Central America), Leptogoneae (1/1, Hispaniola) and Gymnopodieae (1/2, S Mexico, Guatemala) do not occur in South America.

2.1 ERIGONOIDEAE ▸ TRIBE COCCOLOBEAE (3/125) - outsiders Podopterus (3; Mexico, Guatemala), Neomillspaughia (2; Central America).

2. Coccoloba P. Browne ex. L. Shrubs, trees, making ECM symbioses with fungi, often with scrambling branches or lianas, unisexual flowers and pedunculate fleshy fruits. 176 spp., tropical and subtropical regions in southern North America to South America, 83 in South America, 46 in Brazil, 17 endemics; C. cereifera Schwacke from Minas Gerais state is a rare plant in Brazil, by Plantas Raras do Brasil’s book.

Although records of sheet sizes are common between plants of the W Colombia, the plant with the largest simple leaves worldwide is C. gigantifolia E. Melo, C.A. Cid Ferreira & R. Gribel from in the Madeira River region, north of Rondonia and S Amazonas; these leaves can reach 2.5 m length and 1.44 in width; they are, of course, larger than the Grias (Lecythidaceae) and Pentagonia (Rubiaceae). C. excelsa Benth. from Nicaragua to South America is myrmecophyte.

2.2 ERIGONOIDEAE ▸ TRIBE TRIPLAREAE (4/56-60) - all genera occur is South America.

3. Magoniella Adr. Sanchez. (inc. Ruprechtia p.p.) Strict lianaceous habit, myrmecophytes, with hollow stems and the fruits are green with red sepals. Two spp., M. laurifolia (Cham. & Schltdl) Adr. Sanchez, is confined to SE Brazil in Atlantic Forest and in Atlantic sandy coastal shrublands (restingas), between 0–500 m, and M. obidensis (Huber) Adr. Sanchez in known from Brazil, Bolivia, and Venezuela where it occurs in secondary forests and on margins of lowland rain forests, between 100–900 m.

4. Ruprechtia C. A.Meyer. Trees to shrubs, three-winged fruits, female partial inflorescences 2-3-flowered, male flowers pedicellate, twigs often solid. 34 spp., in all Latin American countries except Chile, 28 in South America, with the highest diversity in Brazil (14, 5 endemics), followed by Venezuela (11); most of the species are restricted to seasonally dry and dioecious habitat. Two species in South America (both in Brazil) are myrmecophytes.

5. Salta Adr. Sanchez. Trees or shrubs. Only one sp., S. triflora (Grisebach) Adr. Sanchez, in Brazil, Argentina, Bolivia, and Paraguay, a common plant that grows in Chaco forests and thickets and occasionally, in seasonally inundated forests, between 200–1,500 m; it flowers and fruits in the dry season (while the plant is leafless).

6. Triplaris Loef. ex. L. Shrubs to trees, myrmecophytes, dioecious bark often peeling off; inflorescence axillary or terminal pleiothyrsi, red to purple three-winged fruits, female partial inflorescences 1-flowered, male flowers sessile, often associated with ants that live in its hollow branches trees. 19 spp., mainly Amazon rainforest, with few species in Central America, the Antilles, and N South America (18); the highest diversity occurs in Peru (13), followed by Brazil (7, one endemic) and Colombia (7), absent in Argentina, considered a pioneer plant that grows in seasonally inundated forests, along rivers, or in disturbed areas; however, there are some species found in dry forests in N Colombia and in San Martin, Peru.

2.3 ERIGONOIDEAE ▸ TRIBE ERIGONEAE (12/410-420) - outsiders Harfordia (1; Baja California, Mexico), Pterostegia (1; W and SE U.S.A.), Gilmania (1; Death Valley in SE California), Dedeckera (1; California), Stenogonum (2; W U.S.A.), Sidotheca (3; California, NW Mexico), Goodmania (1; California, Nevada), Hollisteria (1; California).

7. Chorizanthe R. Br. ex. Benth. Spreading to erect annual or perennial herbs or shrubs. 60 spp., 42 annuals in North America and Mexico and 18 perennials in dry areas of Chile and Argentina.

8. Eriogonum Michx. 253 spp., all from North America and Mexico, except disjunct populations of E. divaricatum Hook. collected in three locations in southern Argentina, and it has not been collected for several decades.

9. Lastarriaea Remy. Prostrate to ascending annual herbs. Three spp. from S California to C Baja California, with L. chilensis J. Rémy disjunct in N and C Chile.

10. Oxytheca Nutt. Erect to decumbent annual herbs; leaves basal or cauline; flowering stem erect. 3 spp. from SW U.S.A., mainly Greater Basin and Mojave, with O. dendroidea Nuttall subsp. chilensis (J. Rémy) Ertter, restricted to the Andes of Chile and Argentina; it, like another annual, Chorizanthe commissuralis J. Rémy, which is closely allied to western North America C. brevicornu Torrey, is probably a recent introduction into South America.

3. SUBFAMILY POLYGONOIDEAE (17/880–900) ‣ 6 tribes, Oxygoneae (1/c 30; tropical and S Africa, Madagascar), Fagopyreae (2/16; E Africa, Asia) and Calligoneae (1/80–85; Mediterranean to India) do not occur in South America.

3.1 POLYGONOIDEAE ▸ TRIBE POLYGONEAE (7/c. 340) - outsiders Oxygonum (22; tropical and S Africa, Madagascar); Knorringia (1; Siberia, Afghanistan, Pakistan, Central Asia, Himalayas, W China); Fallopia (c 22; temperate regions on the Northern Hemisphere); Atraphaxis (40–45; SW Europe, N Africa, W Asia to Himalayas and E Siberia), Duma (3; Australia).

11. Muehlenbeckia Meisner. Twining procumbent or erect, poygamo dioecious or dioeciuos shrubs. 20–25 spp., New Guinea, Solomon Islands, Australia, Tasmania, New Zealand, the Chatham Islands, 8 from Colombia to Cono Sur, two up to Mexico and Central America and M. sagittifolia (Ortega) Meisn. up to S Brazil.

12. Polygonum L. Prostrate or rarely semi-erect, much-branched annual herbs, flowers in spikes. 75 spp., subcosmopolitan, 53 in New World, 19 in South America, 8 Brazil, two endemics.

3.2 POLYGONOIDEAE ▸ TRIBE RUMICEAE (3/c. 260) - outsiders Oxyria (2–3; arctic and alpine regions, circumboreal south to California), Rheum (55–60; Europe, temperate and subtropical regions in Asia).

13. Rumex L. Perennial or annual herns, rarely woody shrubs. c 200 spp., temperate regions on both hemispheres, especially the Northern Hemisphere, 63 in New World, 22 in South America, only two in Brazil, R. brasiliensis Link up to Cono Sur and R. sellowianus Rech. f. endemic.

3.3 POLYGONOIDEAE ▸ TRIBE PERSICARIEAE (4/185-192) - outsiders Rubrivena (1; S Himalayas, Xizang, N Burma, W China), Bistorta (40–45; temperate and arctic regions in Europe, Asia and North America).

14. Koenigia L. Plants decumbent, ascending to erect annual or perennial herbs, arising from taproots, occasionally rooting adventitiously from proximal nodes. 60 spp., mainly in meadows, along stream banks, or on talus slopes in arctic, temperate and alpine regions of the Northern Hemisphere, with K. islandica L., single species in New World, is arctic and circumpolar spp., also in S South America in Tierra del Fuego.

15. Persicaria Mill. 150 spp., almost cosmopolitan, 18 spp. in New World, 7 in South America, 5 widely distributed (4 of then in Brazil) and two endemics to Brazil, these from São Paulo to Santa Catarina states.

PLUMBAGINACEAE

§ FULLY CHLOROTROPHIC

Genera/species 21/700–800 Distribution cosmopolitan except Antarctica, with their largest diversity in arid and saline environments of Mediterranean and SW and Central Asia. Habit bisexual, usually perennial herbs or shrubs (sometimes annual herbs, rarely lianas). Many species are xerophytes or halophytes.

Herbs or shrubs, sometimes scrambing. Secretors glands oftem presents. Some spp. are endemic to temperate South America. Plumbaga auriculata Lam. (a climber) and P. indica are commonly naturalized in the Neotropics with others cultivated throughout the region. The remaining of the family is native, with some endemic species. Pollinated by bees, flies and small beetles. From the Latin for lead ‘Plumbum’, so called by Pliny who attributed the curing of lead disease to European species.

Key differences from similar families - the following families differ from Plumbaginaceae in having the following features:

Primulaceae:

Sympetalous.

Opposite leaves (can also be alternate or basal).

Axillary inflorescence.

Often solitary flowers.

Polygonaceae:

Tepals.

Normally 3-sided fruiting structures, either a nut or achene.

Conspicuous swollen nodes on the stem.

Frankeniaceae:

Revolute leaves.

Pedicels absent.

Ligule-like appendage present adaxially at petal base.

SYSTEMATICS two subfamilies, both in South America.

1. SUBFAMILY STATICOIDEAE (24/590) ‣ two subtribes, Aegialitideae (1/2, coasts along E India to Burma, Andaman Islands, N Australia and S New Guinea) does not occur in South America; among Staticeae, outsiders are Acantholimon (290–300; E Mediterranean to C Asia), Bamiania (1; Afghanistan), Bukiniczia (1; Afghanistan, Pakistan), Cephalorhizum (6; C Asia), Ceratolimon (3; Mediterranean), Chaetolimon (3; C Asia), Dictyolimon (2; Afghanistan to India), Ghaznianthus (1; Afghanistan), Gladiolimon (1; Afghanistan), Goniolimon (c 20; Russia and the Balkan Peninsula to Mongolia), Ikonnikovia (1; C Asia, China), Limoniastrum (1; Mediterranean), Limoniopsis (2; Türkiye to the Caucasus), Muellerolimon (1; Australia), Myriolimon (2; W and C Mediterranean), Neogontscharovia (2; Afghanistan, C Asia), Popoviolimon (1; Central Asia), Psylliostachys (c 7; E Mediterranean to the Caucasus and Central Asia), Saharanthus (1; Morocco, N Sahara), Vassilczenkoa (1; Afghanistan, Central Asia).

1. Armeria (A. P. Cand) Wild. Caespitose perennial herbs, with a branched woody caudex, sometimes cushions. 98 spp., temperate regions on the Northern Hemisphere, and only one sp., A. maritima (Mill.) Willd. var. goodalliana T.R., in the Chilean Andes southwards to Tierra del Fuego.

2. Bakerolimon Linez. Shrubs, leafless; branches thin, covered of small, apressed scale. Only one sp., B. plumosum (Phil.) Lincz., endemic to deserts of N Chile; records in Peru belongs areas today in Chile lands.

3. Limonium L. Perennial, rarely annual herbs or dwarf shrubs, usually in a basal rosette. 350 spp., cosmopolitan, with their highest diversity in maritime and arid habitats on the Northern Hemisphere; 5 spp. in Neotropics, found is to be found in coastal areas; two spp. in South America, L. brasiliense (Boiss.) Kuntze in S Brazil and adjacent Argentina and Uruguay, and L. guaicuru Kuntze endemic to Chile.

2. SUBFAMILY PLUMBAGINOIDEAE (4/36) ‣ outsiders Ceratostigma (8; NE tropical Africa, Tibet, China, SE Asia), Dyerophytum (3; Namibia, N and W Cape, Socotra, Arabian Peninsula to India), Plumbagella (1; Central Asia).

4. Plumbago L. Herbs, shrubs or subshrubs, leafy. 24 spp., tropical to warm-temperate regions on both hemispheres, mainly in Africa, Mediterranean, 3 in New World; can be found throughout the Neotropics in dry scrubland, lowland tropical forest and in the Andes; two spp. in South America, P. zeylanica L. dry scrubland from Mexico to Argentina (cosmopolitan, in all New World countries except Canada, Panamá, Chile and Uruguay) and Brazil; and P. caeruela Kunth. Colombia, S Ecuador to Chile and Argentina, absent in Brazil.

P. zeylanica L. and P. scandens, both Linnaean species, have heretofore been treated as distinct, the former name applied exclusively to Old World plants, the latter to New World specimens. John Edmondson (pers. comm.) indicates that he believes this ‘could be a classic case of New World and Old World taxonomists each doing their own thing.’ Plants in herbaria under these two names appear indistinguishable.

LINEAGE 2: FRANKENIIDS

FRANKENIACEAE

§ FULLY CHLOROTROPHIC

Genera/species 1/82 Distribution northernmost and southernmost Africa, Macaronesia, St. Helena, southern Europe, SW Asia and Australia, with their largest diversity in Mediterranean and SW Asia, New World. Habit usually bisexual (sometimes polygamomonoecious), evergreen shrubs, suffrutices or perennial (sometimes annual) herbs. Usually halophytic or xerophytic, sometimes gypsophilous or calciphilous.

Key differences from similar families - differs from Plumbaginaceae in having: revolute leaves; pedicels absent; ligule-like appendage present adaxially at petal base; 1-3 carpels.

SYSTEMATICS A single genus.

1. Frankenia L. Caracters of family, sometimes cushions in South America. 78 spp., 16 spp. in New World, 9 spp. in SW South America: F. chilensis C. Presl, from coastal areas of S Peru and N Chile, in saline conditions; F. triandra J. Rémy, an inland species, also found in S Peru and N Chile, NW Argentina and the Puna of Bolivia at 3,350-4,800 m; six restricteds of Cono Sur up to Uruguay, and F. salina (Molina) I.M. Johnst. disjunct North America and Cono Sur.

LINEAGE 3: RHABDODENDRACEAE

RHABDODENDRACEAE

§ FULLY CHLOROTROPHIC

Genera/species 1/3 Distribution tropical South America (mainly the Amazon rainforest). Habit usually bisexual (rarely androdioecious), evergreen trees or shrubs; the family was often placed in or near Rutaceae and was in the Chrysobalanaceae for a while; it differs in many characters from both families; molecular and wood anatomy place it near to the Phytolacaceae in the Caryophyllales.

SYSTEMATICS A single genus.

1. Rhabdodendron Gilg & Pilg. Shrubs or small trees. Leaves entire, alternate, gland-dotted, coriaceous, with small peltate hairs on the undersurface; stipules small, subulate or obscure; inflorescence of supra-axillary racemose panicles or racemes; bracts and bracteoles small and reduced to scales; flowers hermaphrodite; petals 5. Three spp., native and endemic to the Neotropics: R. gardnerianum (Benth.) Sandwith, a shrub 6 m taller, usually smaller, known from the type, collected in 1839 on the banks of the Rio Negro in Bahia state (a rare plant in Brazil, by Plantas Raras do Brasil’s book), and so may be extinct; some records possibly in Tocantins state; R. macrophyllum (Spruce ex Benth.) Huber, a multi-trunked also with pelucid dots on the leaves; this species is a common shrub of north-amazonic white-sand savannas (campinaranas) in the immediate vicinity of Manaus and east to the Trombetas River only, and one record in Moju Island; and R. amazonicum (Spruce ex Benth.) Huber, a tree to 15 m x 20 cm diameter, usually smaller; this species is common in terra firme forests from the Manaus region eastwards to the Para-Maranhão border and northwards into the three Guianas, and two highly isolated records in Colombia and upper Rio Negro of Brazil.

LINEAGE 4: MICROTEACEAE

MICROTEACEAE

§ FULLY CHLOROTROPHIC

Genera/species 1/10 Distribution Baja California to South America America and Caribbean. Habit bisexual, usually annual herbs.

Key to families in Phytolaccaceae s.l.

1. Ovary of 5-16 carpels and seeds ------------ Phytolaccaceae s.s.

1. Ovary of a single carpel and seed - 2

2. Ovary with 2-4 stigmas ------------ Microteaceae

2. Ovary with a single stigma ------------ Petiveriaceae

SYSTEMATICS A single genus.

1. Microtea Sw. Annual herbs (occasionally perennial and somewhat lignified at base), fruit a muricate to spiny achene. 10 spp., Central America to Venezuela, E Brazil, and the Antilles to Argentina; 9 spp. occurs in Brazil (the exception is M. portoricensis Urb., restricted of Cuba, Hispaniola and Porto Rico), and half of family are endemic to this; M. debilis Sw. and M. maypurensis (Kunth) G. Don are considered as aliens in the humid tropics of Africa (Cameroon) and Asia (Indonesia), respectively.

LINEAGE 5: AMARANTO/CARYOPHILIIDS

CARYOPHYLLACEAE

§ FULLY CHLOROTROPHIC

Genera/Species 101/2,625 Distribution cosmopolitan although mainly temperate regions in the Northern Hemisphere, the Arctic, temperate parts of the Southern Hemisphere (including the Antarctic continent), tropical mountains, with their largest diversity in Mediterranean and West and Central Asia. Habit usually bisexual (rarely monoecious, andromonoecious, dioecious or gynodioecious), usually perennial, biennial or annual herbs (sometimes suffrutices; rarely shrubs, small trees or lianas).

Caryophyllaceae includes 54 locally endemic genera (many of which in the eastern Mediterranean region of Europe, Asia, and Africa). Caryophyllaceae and Molluginaceae are exceptional on this order by in processing anthocyanin pigments rather than betalains. In neotropics, this family centred in Andes of Peru to Chile. Few in Caribbean and tropical Brazil; the number of genera and species in the Southern Hemisphere is rather small.

525 spp. in New World, 254 in South America. 10 genera and 25 spp. in Brazil, three Brazilian endemics.

SYSTEMATICS two main clades, both in South America.

UNPLACED

1. Reicheëila Pax. Caespitose moss-like herbs (cushions); leaves small, scale-like, with scarious margins. Only one sp., R. andicola (Phil.) Pax, Chile, in altitudes beyond 3,000 m.

1. SUBFAMILY TELEPHIOIDEAE (2/15) ‣ outsider Telephium (5; Mediterranean, Madagascar).

2. Corrigiola L. Herbs, annual or biennial (perennial); taproots slender; stems decumbent to ascending, branched, terete. 13 spp., nearly cosmopolitan, especially Europe, Africa. 4 spp. in South America, 3 restricted for Cono Sur, and C. andina Planch. & Triana disjunct in Mexico, Peru and Bolivia.

2. SUBFAMILY CARYOPHYLLOIDEAE (c. 80/c. 2,600) ‣ 11 clades, Sclerantheae (9–10/85–100, Northern Hemisphere, Ethiopia, New Guinea, Australia), Eremogoneae (1/90–100, temperate regions on the Northern Hemisphere), Rhodalsine clade (1/1, Mediterranean, Canary Islands, Morocco to N Egypt, N Somalia) and Caryophylleae (13/665–685, Eurasia, Africa, North America and Australia and New Zealand) do not occcur in South America.

2.1 CARYOPHYLLOIDEAE ▸ TRIBE PARONYCHIEAE (c 6/110-160) - outsiders Chaetonychia (1; W Mediterranean), Gymnocarpos (10; Canary Islands to E Asia, NE Africa), Cometes (2; NE Africa and Ethiopia to NW India).

3. Herniaria L. Herbs or dwaft shrubs, often mat-forming. 48 spp., Europe, Canary Is, Mediterranean, N Africa, Somalia, southern Africa, SW Asia to India, one sp., in H. austroamericana Chaudhri & Rutish in Bolivia and N Argentina.

4. Paronychia Milller. Herbs, often woody at base, or small shrubs, sometimes cushions. 110 spp., nearly cosmopolitan (except southern Africa, SE Asia), especially Mediterranean, Türkiye, SE U.S.A. and the C Andes in Peru. 53 spp. in New World, 26 in South America, 22 from Venezuela to Cono Sur, six in Brazil (all South American exclusives), two endemics, both rare plants in Brazil, by Plantas Raras do Brasil’s book, P. fasciculata Chaudhri from Minas Gerais state, and P. revoluta C.E. Carneiro & Furlan in Rio Grande do Sul state.

5. Philippiella Speg. Cushion-forming, dioecious, small shrub; leaves imbricate, slightly fleshy, stipules interpetiolarly connate. One sp., P. patagonica Speg., Patagonia of Chile and Argentina. Oxelman & al. (2002) and Greenberg & Donoghue (2011) found P. patagonica was nested within Herniaria; the genus is retained pending additional sampling in Herniaria.

2.2 CARYOPHYLLOIDEAE ▸ TRIBE POLYCARPAEAE (20/155–165) - outsiders Dicheranthus (1; Canary Islands), Pteranthus (1; North Africa and Cyprus to Iran), Illecebrum (1; Europe, Canary Islands, Mediterranean), Loeflingia (4; S Iberian Peninsula; W Mediterranean; S Portugal; U.S.A., Mexico), Polycarpaea (40–50; warmer regions of both hemispheres),Scopulophila (2; Mexico, California, Nevada), Sphaerocoma (1; NE Sudan, S Arabian Peninsula, Iran), Pollichia (1; N Arabian Peninsula, Ethiopia, tropical E to S Africa), Achyronychia (1; SW U.S.A., Mexico), Cerdia (1; Mexico), Ortegia (1; the Iberian Peninsula), Krauseola (2; N Kenya, S Ethiopia; Mozambique), Pirinia (1; SW Bulgaria), Polytepalum (1; Angola), Stipulicida (1; SE U.S.A., Cuba).

6. Augustea Iamonico. (off Polycarpon) Shrubs, chamaephytes/nanophanerophytes, up to 50 cm tall; stem woody at the base, branched; leaves often in whorls of 4 or in whorls of 4 (rarely 6) shortly petiolate or sessile; inflorescence terminal and dichotomous cymes; flowers subsessile. 4 spp., A. anomalum Hassler endemic to Paraguay, A. suffruticosum (Griseb.) Iamonico from Argentina, Bolivia, Paraguay, and two endemics to Chile.

7. Cardionema DC. Tufted perennial herbs; stems slightly woody. 6 spp., three only Bolivia and Cono Sur, C. congestum (Benth.) A. Nelson & J.F. Macbr. endemic to Ecuador, C. camphorosmoides (Cambess.) A.Nelson & J.F.Macbr. endemic to E Brazil, and C. ramosissimum (Weinm.) A. Nelson & J.F. Macbr. widely distributed from North America to Cono Sur and SE Brazil, a aggressive spine fruited common in coast of S Brazil.

8. Drymaria Willd. ex. J. A. Schults. Annual or perennial herbs. 59 spp., 58 restricted of New World from western U.S.A. to Patagonia, and D. cordata (L.) Willd. ex Roem. & Schult. in over tropcal New World (unique species in Brazil) and tropical Africa; 22 spp. in South America.

9. Microphyes Philippi. Low herbs, often strongly pubescent; leaves mainly basally. Tree spp. in Chile, one up to S Argentina, in dry sandy habitats.

10. Polycarpaea Lam. Herbs to shrubs, sometimes with xylopodium. 74 spp., only two in New World, P. hassleriana Chodat from Bolivia and Paraguay, and P. corymbosa (L.) Lam. from over South America (collected in several Brazilian states), Panamá, tropical Africa south to Saara, India and E China to Thailand.

11. Polycarpon Loef. 7 spp., 4 from Europe to N Africa and India, make up a complex of highly similar, annual or perennial taxa mainly distributed in Mediterranean region, P. depressum Nutt. in California to Mexico (Baja California), P. prostratum (Forssk.) Asch. & Schweinf. in over tropical Old World, and P. apurense Kunth in Venezuela, Colombia, Peru, Paraguay, Brazil, and Argentina.

12. Pycnophyllum Remy. Perennial caespitose or tufted herbs with creeping rhizomes (cushions), dioecious. 29 spp., Andes from 3,000-4,600 m in Chile, Bolivia, Peru (16 endemics) and Argentina.

2.3 CARYOPHYLLOIDEAE ▸ TRIBE SPERGULEAE (3/c 65) - outsider Thylacospermum (1; C Asia, Himalayas, W China)

13. Spergula L. Annual, arrely perennial herbs, often glandular viscid. 12 spp. in New World. 9 in South America, all in Cono Sur, S. grandis Pers. and S. levis (Cambess.) D.Dietr. up to S Brazil.

14. Spergularia (Pers.) J. Presl. and C. Presl. Herbs, annual or strongly perennial with branched, woody caudex; taproots filiform to stout; stems erect to sprawling, simple to freely branching distally or throughout, terete, sometimes woody. 60 spp., coastal and saline areas, 24 in New World, W North America (including Mexico), Central America, W South America (19, 10 from Colombia to Peru, and 11 in Cono Sur, S. fasciculata Phil. and S. platensis (Cambess.) Fenzl in both areas), Europe (Mediterranean region), Africa (Mediterranean region), S. ramosa Cambess. from Bolivia to Brazil (Santa Catarina) and Argentina.

S. manicata (Skottsb) Kool & Thulin, endemic to remote San Ambrosio Island, off the coast of Chile, in the only member of Caryophyllaceae that may grow to a small tree.

2.4 CARYOPHYLLOIDEAE ▸ TRIBE SAGINEAE (c 10/210–215) - outsiders Drypis (1; Mediterranean to Lebanon); Bufonia (34; Canary Islands, Mediterranean, the Middle East), Mcneillia (5; S and SE Europe, NW Anatolia in Türkiye), Minuartia (c 55; Europe, North Africa, SW Asia, the Caucasus, N India), Habrosia (1; Syria, Iraq, W Iran), Minuartiella (4; mountain regions from Anatolia in Türkiye to Nakhichivan in Azerbaijan and N Iran), Facchinia (7; alpine areas in Central Europe).

15. Colobanthus Bartl. Glabrous, often strongly caespitose perennial herbs, sometimes cushions, usually with strong taproot. 20 spp., mountains of SE Australia and Tasmania, New Zealand and adjacent islands, four in New World: C. lycopodioides Griseb. in the Falkland Islands, the Antarctic Peninsula and adjacent islands, South Georgia, New Amsterdam, Kerguélen, C. quitensis (Kunth) Bartl. in tropical South America disjunct in Mexico, up to 3,600 m altitudinal range in Andes of Colombia, Bolivia and Peru one endemic each.

C. quitensis (Kunth) Bartl.) is the world´s southernmost dicot, which is one of only two flowering plants found in Antarctica.

16. Sabulina Rchb. (exc. Arenaria p.p.) Annual or perennial herbs, rarely subshrubs, often tufted or mat-forming, glabrous or pubescent. 65 spp., (possibly 70, including some W European and W Asian taxa not yet transferred to Sabulina), all but two found in the northern hemisphere (Europe and Asia, two in North America), one in South America, S. acutiflora (Fenzl ex Endl.) Dillenb. & Kadereit in southern Chile disjunct North America.

E. Hultén (1964) confirmed the report of Sabulina groenlandica (Retz.) Small from a mountain in southern Brazil (Morro de Igreja, Santa Catarina state, as Minuartia groenlandica (Retz.) Ostenf.); this remains the only report of ex-Minuartia in South America, but not fully accepted.

17. Sagina L. Annual or perennial herbs, dwarf, often tufted; flowers solitary or in few-flowered cymes. 15–20 spp., arctic, temperate and alpine regions on the Northern Hemisphere, mountains in E Africa and New Guinea, Himalayas, 8 spp. in New World, 6 in North America to Mexico and Caribbean, and two in Cono Sur, S. chilensis Gay and S. humifusa (Cambess.) Fenzl ex Rohrbach, the latter up to Rio Grande do Sul state in S Brazil.

2.5 CARYOPHYLLOIDEAE ▸ TRIBE ARENARIEAE (4-5/c. 200) - outsider Moehringia (c 30; temperate Europe, Asia and North America).

18. Arenaria Ruppius ex. L. (inc. Alsine, Scleranthus, Sabulina p.p.) Annual or perennial, sometimes cushions (e.g. in Andes); taproots filiform to moderately thickened; rhizomes slender; stems prostrate to ascending or erect, simple or branched, terete to ellipsoid, angular or grooved; with leaves ovate to lanceolate, petals usually with margins entire, stamens 10. 160 spp., mainly in northern temperate regions, also in the Artic, some spp. on the mountains of South America and NE Africa; 61 spp. in New World, 43 in South America, all belong to former subgen. Dicranilla and subgen. Leiosperma; A. lanuginosa (Michx.) Rohrb. occur from SE. U.S.A. to NW Venezuela and N. Argentina, S & SE Brazil, Caribbean; it’s a morphologically diverse, both in our area and southward into N South America, and is in serious need of comprehensive study; other species in subg. Leiosperma (e.g., A. gypsostrata B. L. Turner) that occur in Mexico resemble A. lanuginosa; the nature of those relationships also requires study.

2.6 CARYOPHYLLOIDEAE ▸ TRIBE ALSINEAE (12/400–410) - outsiders Lepyrodiclis (3; Türkiye to Himalayas), Pseudostellaria (17; Europe, Afghanistan, Central Asia and W China to Korean Peninsula and Japan, Canada, U.S.A.), Odontostemma (c 65; China, Tibet, Sikkim), Shivparvatia (7; N India, Himalayas, SW China), Holosteum (5; temperate regions in Europe and Asia, Ethiopia), Moenchia (4; Europe, Mediterranean), Brachystemma (1; Himalayas, China, Indochina), Pseudocerastium (1; Anhui in E China), Thurya (1; SW Asia).

19. Cerastium L. Annual or perennial herbs, rarely subshrubs, usually glandular hairy; leaves sessile or subsessile. 100 spp., with almost cosmopolitan distribution; c. 66 in New World, 38 in South America, from Venezuela to Chile, Argentina and Uruguay, 8 in Brazil, all restricteds from South America, none endemics from country.

20. Pycnophyllopsis Skottsb. Moss-like herbs (cushions), perennial evergreen, glabrous. 5 spp., 4 in Andes of Bolivia and Peru, and P. muscosa Skottsb. in montane regions of Patagonia, Argentina.

21. Stellaria L. Annual or perennial herbs, often fragile. 173 spp., mainly Eurasia, with center of diversity in E central Asia, some are afromontane, others cosmopolitan. 52 spp. in New World, 24 in South America, from Venezuela to Chile and Argentina.

2.7 CARYOPHYLLOIDEAE ▸ TRIBE SILENEAE (2/882) - outsider Agrostemma (2; temperate regions of Europe and Asia, Mediterranean).

22. Silene L. Annual, biennial or perennial herbs, rarely geophytes or small shrubs, sometimes cushions. 880 spp., subcosmopolitan, 76 spp. in New World, 23 in South America, all from S Ecuador to Chile and Argentina, 17 in Cono Sur.

ACHATOCARPACEAE

§ FULLY CHLOROTROPHIC

Genera/species 2/10 Distribution Texas, Mexico, Central America, South America to Paraguay and Argentina. Habit dioecious, evergreen trees or shrubs. Apices of short shoots often modified into spines. The Achatocarpaceae has been treated as part of the Phytolaccaceae in the past, but now it stands alone as an independent family. Recent molecular studies using the markers rbcL and matK resolve Amaranthaceae-Chenopodiaceae as sister to the Achatocarpaceae and this evidence is supported by existing morphological data.

Key differences from similar families It differs from the Amaranthaceae s.l. in having fleshy fruit with translucent skin (vs. dry fruit with different projections, dispersed mostly by wind); the Phytolaccaceae have mostly bi-many-locular ovaries, while the Achatocarpaceae has a one-locular ovary with one ovule.

SYSTEMATICS outsider Phaulothamnus (1; California, Texas, N Mexico, Tres Marias Islands)

1. Achatocarpus L. Shrubs or trees up 10 m heigh, leaves often succulent, fleshy, deciduous, drying black. 9 spp., 7 in South America: 4 in extra-Brazil Chaco region (two endemics to Paraguay); two from Mexico to Venezuela and Bolivia; and A. praecox Griseb. in Peru, Bolivia, S Brazil, N Argentina and Paraguay.

AMARANTHACEAE

§ FULLY CHLOROTROPHIC

Genera/species 184/2,400–2,500 Distribution cosmopolitan except polar areas, with their highest diversity in saline, arid and semiarid areas. Habit usually bisexual (sometimes monoecious, andromonoecious, gynomonoecious, dioecious, androdioecious, rarely polygamomonoecious), perennial, biennial or annual herbs, evergreen or deciduous suffrutices or shrubs (rarely trees or lianas), sometimes with spines. Often leaf or stem succulents. C₄ plants with c. 17 different types of foliar anatomy. Many species are halophytes or xerophytes.

Currently, 19 families of higher plants are known to contain species expressing the C4 photosynthetic pathway. In each family, the C4 pathway arose independently, producing approximately 50 distinct evolutionary lineages. Sixteen of these families are eudicots; the eudicot clade with the largest number of C4 species is the Amaranthaceae, with half of the approximately 1,400 eudicot C⁴ species.

Use Ornamental plants, vegetables (Beta vulgaris L., Spinacia oleracea L., Chenopodium quinoa Willd., etc.), sugar (Beta vulgaris var. altissima), forage-plants (Beta vulgaris, Atriplex, Chenopodium), glass production (Salicornia, Salsola, etc.), timber, wood carving, medicinal plants (Dysphania etc.).

SYSTEMATICS subfamilies Salsoloideae (c 35–37/c 370, Europe, Macaronesia, Mediterranean, Africa, SW and central Asia, with their highest diversity in central and SW Asia), Betoideae (6/17, Europe, Macaronesia, Mediterranean region, North Africa, SW Asia, SW North America), Camphorosmoideae (16/170–180, temperate and subtropical Eurasia, N and S Africa, Australia, with their highest diversity in Australia, few species in South Africa or North America) and Corispermoideae (3/c 80, Eurasia) do not occur in South America.

1. SUBFAMILY POLYCNEMOIDEAE (3/16–18) ‣ outsiders Hemichroa (1; Australia, Tasmania), Surreya (2; Australia), Polycnemum (6; C, S and E Europe, Mediterranean, NW Africa, Central Asia).

1. Nitrophila S. Watson. Herbs, perennial, suffrutescent, glabrous; stems numerous from base, erect or prostrate, not armed, not fleshy; inflorescences axillary, flowers solitary or in 2-3-flowered clusters, bibracteate;fruiting structures: fruit a utricle; pericarp free from seed, membranous, indehiscent. 4 spp., two in W North America and Baja California in Mexico, N. atacamensis (Phil.) Ulbr. in Antofagasta region in Chile, and N. australis Chodat & Wilczek in N & S Central Argentina.

2. SUBFAMILY AMARANTHOIDEAE (c 76/c 330) ‣ 5 tribes, four in South America; Aerveae (3/c. 120, 100 in drier regions in Australia, Flores and Timor, tropical Africa from Senegal to Ethiopia and N Somalia, south to S Africa, Mauritius, Pakistan, India, Sri Lanka, Burma, Mauritius, tropical and subtropical regions in the Old World) does not occur in South America.

Key to genera of South American Amaranthoideae

1. Leaves alternate - 2

2. Fruits with many seeds; flowers bisexual - 3

3. Sepals ovate or suborbicular, obtuse; stigmas 3; seeds more than 10 ------------ Pleuropetalum

3. Sepals lanceolate or elliptic, obtuse or acute; stigmas 2-3; seeds less than 10 ------------ Celosia

2. Fruits with one seed; flowers bisexual or unisexual - 4

4. Erect herb; stamen filaments free at the base ------------ Amaranthus

4. Scandent herbs or lianes; stamen filaments fused into a cup-shape - 5

5. Fruits indehiscent or irregularly dehiscent; seeds non-arillate ------------ Herbstia

5. Fruits dehiscing by circumsissile lid; seeds arillate ------------ Chamissoa

1. Leaves opposite - 6

6. Flowers or flower glomerules in elongate spikes or in cymes - 7

7. Flowers in glomerule or a many-flowered cyme; flowers sterile; bractoles terminating in a hook ------------ Cyathula

7. Flowers in spikes or cymes; flowers: one bisexual and one sterile with hook-shaped projections ------------ Pseudoplantago

6. Flowers or flower glomerules in cylindrical spikes, simple heads or arranged in complex paniculate or racemose structures - 10

8. Inflorescences in glomerules or axillary spikes ------------ Guilleminea

8. Inflorescences in simple or composite panicles, racemes or spikes - 9

9. Sepals basally connate; plants with abundant grey indumentum; fruits with or without appendages - 10

10. Sepals connate into a tube; fruits winged; flowers shorter than 5 mm ------------ Froelichia

10. Sepals only connate at base; fruits not winged; flowers longer than 8 mm ------------ Frielichiella

9. Sepals free at base; plants glabrous to densely indumented, but indumentum not grey; fruits without appendages - 11

11. Stigma penicillate; staminodes ligulate or triangular, apex acute or divided, alternating with the filaments ------------ Alternanthera

11. Stigma bilabiate or bifid; staminodes present or absent, not like Alternanthera - 12

12. Stigma bilabiate or broadly emarginated - 13

13. Inflorescences of heads; trichomes straight, surrounding the flowers ------------ Pfaffia

13. Inflorescences in long racemes; trichomes S-shaped, surrounding the flowers ------------ Hebanthe

12. Stigmas bifid - 14

14. Filaments fused into a tube; inflorescences with or without foliolose bracts ------------ Gomphrena p.p.

14. Filaments only fused at base; inflorescences without foliolose bracts - 15

15. Inflorescences of racemes ------------ Iresine

15. Inflorescences in globose or cylindrical spikes or head ------------ Gomphrena p.p.

AMARANTHOIDEAE ▸ UNPLACED GENERA - 36 unplaced genera, 23 in Africa, 2 in Australia, 5 in E & SE Asia, 1 in Arabia peninsula, 2 in over tropical Old World, 1 in Indico Ocean islands, and 2 in South America.

2. Hebstia Sohmer. Herbs or shrubs with alternate leaves; flowers in one to several condensed axillary cymules. Only one sp., H. brasiliana (Moq.) Sohmer, in Brazil, Argentina and Paraguay.

3. Lecosia Pedersen. Herbs to suffrutices, leaves opposite or alternate; flowers hermaphrodite in solitary dichasia; sorne leaves tinged with red on lower surface. Two spp., both rare plants in Brazil, by Plantas Raras do Brasil’s book, known only their type collections, both Atlantic Forest in E Brazil, L. formicarum Pedersen in coast of Bahia state (collected in ant nests), L. oppositifolia Pedersen in Espírito Santo state; in aspect, the species of this genus are not unlike certain species of Celosia, for which reason its name is an anagram of that word; on the whole, the systematic position of Lecosia is uncertain, but probably it should be placed near Celosia.

2.1 AMARANTHOIDEAE ▸ TRIBE ACHYRANTHEAE (8/81) - outsiders Achyranthes (12; tropical and subtropical regions in the Old World), Nototrichium (3; Hawaii), Calicorema (2; tropical Africa and southwards to Namibia and N Cape), Pandiaka (13; tropical and S Africa), Psilotrichum (16; tropical regions in the Old World), Pupalia (4; tropical regions in the Old World), Sericostachys (1; tropical Africa).

4. Cyathula Blume. Annuals or perennial leaves opposite; inflorescence spiciforme or capitate, with a complex and singulat terminal architecture. 27 spp., 12 in mainland Africa, two up to Madagascar, 9 endemics to Madagascar, 4 in E Asia, and two cosmopolitan, C. achyranthoides (Kunth) Moq. and C. prostrata (L.) Blume, both in tropical America and Brazil.

2.2 AMARANTHOIDEAE ▸ TRIBE AMARANTHEAE (2/c. 50) - both genera occur in South America.

5. Amaranthus L. Annual or more rarely perennial herbs (only in A. peruvianus (Schauer) Standl., A. rosengurttii Hunz. and A. vulgatissimus Speg.), erect or rarely decumbent, monoecious or dioecious, green, but sometimes pupish because betalains; inflorescence terminal, spikes or paniculate. 91 spp., 61 in New World, 32 in Mexico (six endemics), 31 in South America, being 22 restriteds from Venezuela to Chile and Argentina, three from Mexico to South America ansents in Brazil (A. australis (A.Gray) J.D.Sauer, A. crassipes Schltdl., A. tortuosus Hornem.), and six in Brazil, four widey distributeds and two with restricted distributions. Four well defined clades:

§ Eurasian/South African/Australian (ESA) + South American clade ‣ contains all of the Old World species and 11 spp. from South America, 3 in Brazil, the widely distributeds A. blitum L., and A. viridis L., and A. rosengurtii Hunz., from Uruguay, Argentina and small portion of southern Rio Grande do Sul state, Brazil, truly assignated as native in Brazil; however, in this country this species is very rare, possibly extinct.

§ Hybridus Clade ‣ monoecious; 13 spp., all endemics to New World, A. spinosus L.and A. hybridus L. in Brazil, both widely distributeds; this monophyletic group here called the Hybridus Clade consists of A. hybridus L. and its domesticated and wild or weedy relatives from the Americas, as well as two lesser-known Neotropical species, A. acutilobus Uline & W.L. Bray and A. scariosus Benth. It loosely corresponds to Mosyakin and Robertson’s subgenus Amaranthus. Includes A. palmeri from North Americ and Mexico, and possiby the Brazlian endemic A. bahiensis Mart.

§ Dioecious/Pumilus clade(S) ‣ all dioecious species of Amaranthus are included in this group, except for A. palmeri S. Watson and A. watsonii Standl., both from Hybridus clade; this clade includes one monoecious species, A. pumilus Raf. from North America.

§ Galapagos Clade ‣ include species from North America, Central America, Caribbean and Galapagos (4).

6. Chamissoa Kunth. Herbs or shrubs with arching branches, or scandent, with alternate leaves; flowers in long spike-like thyrsi. Tree spp., C. altissima (Jacq.) Kunth. widely distributed from Mexico to South America and Caribbean, and two restricted of center and E South America, from Peru to Argentina and Brazil (both).

2.3 AMARANTHOIDEAE ▸ TRIBE CELOSIEAE (5/81) - outsiders Deeringia (12; tropical and subtropical regions in the Old World), Henonia (1; Madagascar), Hermbstaedtia (c 15; tropical and southern Africa except the Cape provinces).

7. Celosia L. Perennial, annual, suffrutescent or ocasionally scadent herbs; flowers in lax or compact thyrses or spikes. 45 spp., worldwide, 10 in South America, 6 mainly Mexico, one up to North America and another up Noth America and Caribbean; C. corymbifera Didr. and C. persicaria Schinz are E Brazil and Peru endemics, C. virgata Jacq. from Mexico to Peru and Venezuela, also in Caribbean, and C. grandifolia Moq. occur from Colombia to Peru, disjunct in E Brazil.

8. Pleuropetalon Hook. f. Small subshrubs, shrubs or more rarellly small trees, with alternate leaves. Three spp., P. darwinii Hook. f. from Galapagos, and remaining two widely from Mexico to Bolivia and Caribbean.

2.4 AMARANTHOIDEAE ▸ TRIBE GOMPHRENEAE (19/c. 390) - outsiders Woehleria (1; Cuba), Tidestromia (6; SW U.S.A., Mexico), Gossypianthus (2; U.S.A., Mexico, Central America).

9. Alternanthera Forssk. Annual or perennial, sometimes scrabling or floating, with opposite leaves; inflorescence axillary, sessile or pendunculate, solitary or fasciculate, of heads or shorts spikes. 102 spp. of New World, 82 in South America, 34 in Brazil, 14 endemics; two spp. from Minas Gerais state are rare plants in Brazil, by Plantas Raras do Brasil’s book.

10. Froelichia Moench. Annuals or perennials with opposite leaves, sometimes with xylopodium; flowers hermaphrodite, solitary in the bracts of branched spikes, the branches often short and dense. 15 spp., tropical and subtropical N and S America (9), only 5 in Brazil, F. sericea Moq. endemic; F. interrupta (L.) Moq. is widely in tropical America.

11. Froelichiella R.E.Fr. Erect small herb with chartaceous opposite leaves; flowers hermaphrodite, solitary in the bracts of a short, compact or interrupted spike. Only one sp., F. grisea R. E. Fr., a rare plant in Brazil, by Plantas Raras do Brasil’s book, very narrow endemic N Goiás state in center Brazil.

12. Gomphrena L. (inc. Pseudogomphrena, Blutaparon, Lithophila) Annual or perennial with opposite leaves, sometimes with xylopodium, sometimes fleshy, mainly showy globose inflorescences, capitate to spicate, axillary or terminal, sessile or pendunculate. 129 spp., tropics and subtropics of New World (92) and Australia; 87 spp. in South America, 47 in Brazil, 30 endemics; 9 spp. in several states are rare plants in Brazil, by Plantas Raras do Brasil’s book.

13. Guilleminea Kunth in H. B. K. Perennial with opposite leaves; inflorescence, axillary, in dense often fasciculate spikes. 7 spp., one in Caribbean, G. densa (Humb. & Bonpl. ex Schult.) Moq. from North America to Argentina along W South America, 4 in Bolivia and Cono Sur, and G. fragilis Pedersen from Paraguay to SW Brazil in Mato Grosso do Sul state.

14. Hebanthe Mart. Shrubs, subshrubs, scadent or semi-scadent (some fully climbing), woody underground system; leaves opposite, ovate; inflorescences spikes, joined in panicles, axillary or terminal; flowers bisexual; fruit a capsule monospermic. 7 spp., H. paniculata Mart. in Andes of Bolivia and Peru, 5 from Brazil up Bolivia and Cono Sur (3 endemics), and H. grandiflora (Hook.) Borsch & Pedersen from Mexico to Guianas, Bolivia and Brazil, barely in Caribbean, slightly centered in E South America.

H. eriantha (Poir) Pedersen is one of the important medicinal plants of Brazil due to its adaptogenic qualities that serve to normalize and enhance body systems, increase resistance to stress, and boost overall functioning.

15. Hebanthodes Pedersen. Herbs; flowers in elongate spikes grouped in a panicle with plurinodal main axis; filaments connate at base only; stigma broadly extended, emarginated. Only one species known, H. peruviana Pedersen, endemic to Pasco region, center Peru, which is the type.

16. Iresine P. Browne. Annual or perennial herbs to subshrubs or shrubs, rarely scadent or small trees; inflorescence capitate or spike-like. 38 spp., widely distributed in New Worlds, mainly in Mexico (30, 20 endemics), 7 in South America, only two in Brazil, I. diffusa Humb. & Bonpl. ex Willd. and I. herbstii Hook., both widely scattered.

17. Pedersenia Holub. 5 spp. from Honduras to Paraguay, Venezuela and N & W Brazil (2, P. argentata (Mart.) Holub, in Amazonas and Pará states, and P. macrophylla (R.E. Fr.) Holub in Mato Grosso do Sul state), none endemics.

18. Pffaffia Mart. Herbs, perennials, erect or semi-prostrate, sometimes with xylopodium; flowers hermaphrodite, perfect, solitary in the bracts of dense and spike-like to lax and paniculate inflorescence; fruit as capsule monospermic. 36 spp., P. completa (Uline & W.L. Bray) Borsch only Central America, two both Central and South America, 33 only South America, 20 in Brazil, 15 endemics, centered in mountains dry areas of center Brazil; only one Brazilian spp. not occurs in savannas of C Brazil (cerrado); two spp. from Minas Gerais state are rare plants in Brazil, by Plantas Raras do Brasil’s book.

19. Pseudoplantago Suesseng. Herbs with opposite leaves, perennials. Only one sp., P. friesii Suess, in Argentina and S Brazil.

20. Quaternella Pedersen. Three spp., Q. confusa Pedersen, Q. ephedroides Pedersen and Q. glabratoides (Suess.) Pedersen, all endemic to Brazil, in over eastern country.

21. Xerosiphon Turcz. Herbs, highly fragile, erect, subglabrous, tuberous; quadrangular stems; sometimes aphyllous; inflorescences simple or cymes of spikes. Two spp., endemic to dry areas in Brazil, one from Minas Gerais to Mato Grosso, and other in dry seasonal scrubland of NE Brazil (caatinga) at Piauí to Bahia and Paraíba, some specimens apparently leafless.

3. SUBFAMILY CHENOPODIOIDEAE (23/460–470) ‣ four tribes, two in South America, Axyrideae (3/10, Mediterranean, E Europe, SW to Central Asia, temperate Asia, Korean Peninsula, western North America) and Anserineae (2/15; Europe, N and E Asia, Australia, Canada, U.S.A.) absents.

3.1 CHENOPODIOIDEAE ▸ TRIBE ATRIPLICEAE (14/c. 400) – outsiders Lipandra (1; Europe, Mediterranean), Chenopodiastrum (5; temperate regions on the Northern Hemisphere), Microgynoecium (1; Tibet), Archiatriplex (1; Sichuan), Exomis (1; Namibia, N, W and E Cape, Free State), Extriplex (2; California), Grayia (3; W U.S.A.), Manochlamys (1; Namibia, N and W Cape), Proatriplex (1; the Navajo Basin in SW U.S.A.), Stutzia (2; W U.S.A.).

22. Atriplex L. Annual or perennial herbs, subshrubs or shrubs; flowers solitary or in clusters. 247 spp., worldwide, 110 in New World, 45 in South America in Cono Sur, Paraguay, Uruguay, Bolivia and Peru, plus A. peruviana Moq. up to Ecuador and three from coasts of Colombia and Venezuela (with A. oestophora S.F. Blake endemic); common in W South America.

23. Chenopodium L. Annual or perennial, nonaromatic (but sometimes foetid) herbs, shrubs or small trees, young stems and leaves often densely farinose, i.e. covered with vesicular globose trichomes, which later collapse forming a cup shaped structure mostly persistent; monoecious or (rarely) dioecious. c. 90 spp., cosmopolitan, nearly absent in E South America; 43 spp. in New World, 25 only in North America and Mexico, Central America and Caribbean; 19 spp. in South America, all from N Peru to Patagonia, Paraguay and Uruguay, some disjuncts in North America, except by C. petiolare Kunth up to Colombia, and C. hircinum Schrad. and C. quinoa Willd. up to Ecuador.

24. Oxybasis Kar. & Kir. (inc. Chenopodium p.p.) Non-aromatic annual herbs; their stems grow erect to ascending or prostrate and are branched with usually alternate, basally sometimes nearly opposite branches; the alternate leaves consist of a petiole and a simple blade; the leaf blade is thickish oder slightly fleshy, and may be triangular, triangular to narrowly triangular, hastate, rhombic, or lanceolate, with entire to dentate margins. 8 spp., subcosmpolitan, mainly Northern hemisphere, 4 in South America, O. macrosperma (Hook. f.) S. Fuentes, Uotila & Borsch from Peru to Argentina and Chile, Paraguay, Uruguay, and Falklands Is, O. rubra (L.) S. Fuentes-B., Uotila & Borsch disjunct in Cono Sur and North America, and two endemics to Cono Sur.

25. Holmbergia Hicken. Climbing shrub. Flowers clustered in leaf axils or arranged in terminal spikes. Only one sp., H. tweedii (Moq.) Speg., from Bolivia, Paraguay, Uruguay and Argentina.

3.2 CHENOPODIOIDEAE ▸ TRIBE DYSPHANIEAE (4/43) - outsiders Suckleya (1; Rocky Mountains), Neomonilepis (1; North America), Teloxys (1; Mongolia).

26. Dysphania R.Br. Annuals or short-lived perennial herbs,more or less covered with simple multicellular hairs andstalked glandular hairs/subsessile glands, sometimes glabres-cent, usually aromatic; stems rarely somewhat woody in lowerpart, erect, ascending, decumbent or prostrate, branched (rarely± simple), not jointed, not spiny, not fleshy. 46 species in five sections.

§ sect. Adenois ‣ 13 spp., two in North America, one in Tristan de Cunha, and remaining 10 in South America, two in S Brazil, none endemics: D. retusa (Moq.) Mosyakin & Clemants ex Brignone and the widely D. ambrosioides (L.) Mosyakin & Clemants.

§ sect. Botryoides ‣ 10 spp., mainly in Himalayas and adjacent China, Central Asia to Arabian Peninsula and Mediterranean Europe and Africa, and E Africa and the adjacent Arabian Peninsula.

§ sect. Dysphania ‣ 17 spp., Australia, one in New Zealand.

§ sect. Incisa ‣ two spp., D. graveolens (Willd.) Mosyakin & Clemants in southern North America and D. mandonii (S.Watson) Mosyakin & Clemants in Argentina, Bolivia, Chile, Peru.

§ sect. Margaritaria ‣ 4 spp., West-Australian and Africa to SW of Arabian Peninsula, with secondary distribution in Europe and West Asia.

4. SUBFAMILY SUAEDIOIDEAE (2/75-80) - outsider Bienertia (3; SE European Russia and Kazakhstan to W Central Asia).

27. Suaeda L. Glabrous herbs, fleshy; flowers perfects and perfects. 91 spp., cosmopolitan, 19 in North America, Mexico and Caribean, and 9 in South America, from Argentina and Chile, S. divaricata Moq. and S. foliosa Moq. up to Bolivia and Peru.

5. SUBFAMILY SALICORNIOIDEAE (10–11/c 90) ‣ outsdiers Halopeplis (3; Mediterranean, North Africa; Arabian Peninsula to Pakistan; the Caucasus and Iraq to Central Asia and Xinjiang), Halostachys (1; Russia to Central Asia), Halocnemum (2; central Mediterranean to Central Asia), Kalidium (6; Mediterranean to Central Asia), Microcnemum (1; Mediterranean to the Caucasus), Arthrocaulon (2; Mediterranean, Macaronesia, NW and NE Africa, SW Asia), Arthroceras (1; California, N Mexico), Tecticornia (44; Australia, one species, T. indica, along coasts of E Africa, S Asia and N Australia).

28. Allenrolfea O. Kuntze. Succulent, articulate shrubs without distinct leaves; flowers 3 to 5 in the axil of free bracts. Three spp., A. occidentalis (S. Watson) Kuntze from U.S.A. and N Mexico; A. patagonica (Moq.) Kuntze and A. vaginata (Griseb) Kuntze are restricted from dry areas in Argentina.

29. Heterostachys Ung. Sternb. Succulent shrubs with cylindrical leaves; flowers solitary, free. Two spp., H. olivascens Speg., endemic to southern coast of Argentina, and H. ritteriana (Moq.) Ung. from Argentina, Paraguay, Colombia, Venezuela, Peru, and Hispaniola (Haiti, Dominican Republic), mainly Caribbean coast.

30. Mangleticornia P. W. Ball, G. Kadereit & Cornejo. Shrubs; stems opposite-branched, jointed and fleshy when young, becoming woody and not jointed, not armed; leaves opposite, sessile, joined at base, eventually deciduous; leaf blades fleshy projections at distal end of each joint; inflorescences terminal, branched, spike-like thyrse. Only one sp., M. ecuadorensis P.W. Ball, G. Kadereit & Cornejo, in SW Ecuador, primarily in Guayas province, extending south to departamento Tumbes in N Peru.

31. Salicornia L. (inc. Sarcorcornia) Annual or perennial fleshy herbs or subshrubs, erect to prostrate, sometimes creeping and rooting at the nodes. 47 spp. in 4 subgenera:

§ subg. Amerocornia ‣ 8 spp., 3 in North and South America and 5 in South America: S. cuscoensis Gutte & G.K. Müll. ex Freitag, M.Á. Alonso & M.B. Crespo (endemic to Andean Peru), S. andina Phil. (saline soils of high plateaus on Atacama Desert in Chile and in Argentina and Bolivia), S. magellanica Phil. (coastal salt marshes on Patagonia and Tierra del Fuego in Argentina), S. neei Lag. (saline deserts in Pacific Coast of Chile and Peru, through the N half of Argentina, to the Atlantic coast of Argentina up to S Brazil) and S. pulvinata R.E. Fr. (strongly saline soils on the edges of temporary, endorheic salt pans of Altiplano Andino in Peru, Bolivia, Argentina and Chile).

§ subg. Arthrocnemoides ‣ 7 spp.; W-Europe, Mediterranean Basin and W-Asia.

§ subg. Afrocornia ‣ ca. 16 spp.; S-Africa and Australia.

§ subg. Salicornia ‣ ca. 17 annual spp.; worldwide except South America and Australia.

LINEAGE 6: AIZOA/NYCTAGINIIDS

AIZOACEAE

§ FULLY CHLOROTROPHIC

Genera/species 120/c. 1,800 Distribution arid tropical and subtropical regions including western and southern Australia, with their highest diversity in S and SW Africa. Habit predominantly succulent, annual to perennial herbs, subshrubs or shrubs.

Aizoaceae is the largest group of succulent species worldwide. Of the several ornamental genera grown, the most notable are the succulents Lithops and Conophytum, the stone plants, which consist of a pair of truncate leaves, often with translucent windows to allow sunlight in. Amongst the members of Aizoaceae with showy flowers, the anthesis is determined by the presence of sunlight, with flowers opening at noon and closing after a few hours to re-open the next day, lasting a few days. Mesembryanthemoideae and Ruschioideae dominate much of the succulent vegetation in the Karroo areas of South Africa, where they constitute more than half the number of species and more than 90% of the biomass.

Key differences from similar families Unlike Cactaceae, where succulence is found on stems and leaves (which are alternate if present) in the Aizoaceae the succulence is confined to their (generally opposite) leaves; Aizoaceae differs from Portulacaceae in opposite leaves and many strap-like perianth segments of staminodial origin; it can be separated from the Molluginaceae because of its succulence and generally colourful and showy flowers, while Molluginaceae are not succulent and have white, small flowers.

SYSTEMATICS only eight genera outside Ethiopian Region: Australian endemics Sarcozona and Gunniopsis, one also in Australia and South America (Carpobrotus), and other (Disphyma) also in Australia and New Zealand, and the widely distributeds Tetragonia, Aizoon (Africa, Mediterranean and Middle East.), Sesuvium (22) and Trianthema (28); subfamilies Acrosanthoideae (1/6, W Cape) and Mesembryanthemoideae (6/c 95, Southern Africa) do not occur in South America.

1. SUBFAMILY SESUVIOIDEAE (5/63) ‣ two tribes: Sesuvieae and Anisostigmateae (2/4, Somalia, Namibia); among Sesuvieae, outsiders Zaleya (7; NE and E Africa, Madagascar, India, Sri Lanka, the Lesser Sunda Islands, northern Australia).

1. Sesuvium L. (inc. Cypselea) Erect to procumbent herbs with opposite leaves that often bear conspicuous sheath-like lateral appendages on the petioles (pseudostipules). 15 spp., 5 endemics to Africa, 3 endemics to Caribbean up to U.S.A. and Mexico, and remaining seven in South America: S. curassavicum Sukhor. in Curaçao and Bonaire Islands, N Colombia e N Venezuela; S. humifusum (Turpin) Bohley & G. Kadereit highly disjuct in Caribbean and N Argentina, S. mezianum (K. Müll.) Bohley & G. Kadereit is endemic to Paraguay, S. edmonstonei Hook.f. is endemic to Galapagos, S. americanum (Gillies ex Arn.) A.I. Jocou & C.R. Minué (= S. verrucosum Raf.) from Mexico and U. S. A., also in Argentina and Peru; S. sessile Pers., endemic to Cono Sur; and S. portulacastrum (L.) L., distributed worldwide in tropical and subtropical zones between approximately 35°N and 42°S, inc. Brazil.

2. Trianthema L. Herbs with leaves with membranes at the base of the petiole. 28 spp., Africa, tropical and subtropical Asia, Australia (12), three spp. in New World, T. argentina Hunziker & Coccuci endemic to Argentina, T. hactandra Wingfield & M. F. Newman endemic to Venezuela, and T. portulacastrum L., pantropical, a weedy very common in NE Brazil, in cities, houses, gardens, etc.

2. SUBFAMILY AIZOOIDEAE (7/124) ‣ outsiders Aizoanthemopsis (1; Mediterranean, northern Africa, the Middle East to Iran), Gunniopsis (14; Australia), Aizoanthemum (4; southern Angola, northern Namibia), Aizoon (c 30; southern Angola to South Africa, one species, A. canariense, in Macaronesia, Mediterranean, Zimbabwe, northern Kenya and Socotra to India).

3. Tetragonia L. Fleshy to succulent, erect to weak annual or perennial herbs or low shrubs, somewhat papillose, flowers axillary, soli-tary to fascicled or cymose, subsessile to stipi-tate or pedunculate. 62 spp., Marocco (1), Australia and New Zealand (5), Africa including Socotra (35), subtropical Pacific coast and temperate coasts of W South America, 5 in Peru and Chile, 4 only in Chile, and T. pedunculata Phil. endemic to Peru.

3. SUBFAMILY RUSCHIOIDEAE (106/1,410–1,430) ‣ all genera are african except Dysphima from W Cape, Australia and New Zealand, and Carpobrotus in Africa, Australia and South America.

4. Carpobrotus N.E.Brown. Subshrubs, succulent, glabrous; stems trailing, mat-forming; rooting at nodes; inflorescence branches ascending; leaves cauline, opposite; inflorescences terminal or axillary, flowers solitary, showy, tubular, 3-10(-15) cm diam.; petals (including petaloid staminodia) to 250, distinct, magenta, pink, yellow, cream, or white. 13 spp., S Africa, Australia, and C. chilensis (Molina) N.E.Br, from Chile and adjacent Argentina.

PHYTOLACCACEAE

§ FULLY CHLOROTROPHIC

Genera/species 4/33. Distribution tropical and subtropical regions, with their largest diversity in South America, also in Chile. Habit usually bisexual (rarely dioecious), evergreen trees, shrubs or lianas (Ercilla), perennial or annual herbs. perennial or annual herbs (Anisomeria is a succulent). Anisomeria and some species of Phytolacca with napiform roots.

The family is of little economic importance. Phytolacca dioica L., ombú tree, is occasionally cultivated in warmer regions of the world, mostly as a fast-growing shade tree having a peculiar trunk and soft, spongy wood. The high saponin content of some species makes them useful for washing clothes, and in eastern Africa Phytolacca dodecandra LHér., endod or soap berry plant, is employed locally for this purpose. This species also has molluscicide properties and is used to control schistosomiasis.

Although Phytolaccaceae clearly belongs to the Caryophyllales, there is much debate about its circumscription and exact position. Some genera historically considered within the family have recently been segregated as separate families. For example, the North American genus Stegnosperma Benth. is now considered in a monogeneric family Stegnospermataceae, as too are the Old World genera Barbeuia Thouars. (Barbeuiaceae) and Gisekia L. (Gisekiaceae).

The New World genera Achatocarpus Triana. and Phaulothamnus A. Gray were recognized by some specialists as members of Phytolaccaceae, but these are now treated in Achatocarpaceae. Even with these genera removed, controversy exists, and the core of the family is sometimes further divided into two families: Phytolaccaceae (ovary of 3-16 carpels, corresponding to subfamilies Agdestioideae and Phytolaccoideae) and Petiveriaceae (ovary of one carpel, corresponding to subfamilies Rivinioideae and Microteoideae).

A definitive conclusion as to their status awaits further comprehensive studies, and the recommendations for a broad Phytolaccaceae are followed here.

Key to families in Phytolaccaceae s.l.

1. Ovary of 5-16 carpels and seeds ------------ Phytolaccaceae s.s.

1. Ovary of a single carpel and seed - 2

2. Ovary with 2-4 stigmas ------------ Microteaceae

2. Ovary with a single stigma ------------ Petiveriaceae

Key to genera of South American Phytolaccaceae

1. Climbers ------------ Ercilla

1. Herbs, shrubs or trees - 2

2. Sepals unequal and somewhat fleshy; carpels distinctly free ------------ Anisomeria

2. Sepals subequal and not fleshy; carpels (at least in the neotropical species) united ------------ Phytolacca

SYSTEMATICS outsiders Agdestis (1; S U.S.A., Mexico, Central America southwards to Nicaragua, introduced in Caribbean and Brazil), Nowickea (2; central Mexico, ‘monstruous forms’ of Phytolacca?).

1. Anisomeria D. Don. Herbs or shrubs, often succulent; flowers in spikes or racemes. Only one sp., A. littoralis (Poepp. & Endl.) Moq., endemic to Chile.

2. Ercilla Adr. Juss. Woody climbers; flowers in dense axillary spikes or spike-like racemes. Two spp., endemic to temperate rainforest in Chile.

3. Phytolacca L. (inc. Nowickea). Herbs, shrubs or trees; flowers in most terminal spikes and racemes. c 25; subcosmopolitan, 14 species in the New World, Mexico to Argentina, 12 in South America, only P. dioica L., P. rivinoides Kunth & C.D. Bouché and P. thyrsiflora Fenzl ex J.A. Schmidt in Brazil, all widely distributeds.

Nowickea J.Martínez & J.A.McDonald, reported as a genus with two Mexican endemic species from Phytolaccaeae, is now reduced as synonimous of Phytolacca icosandra L.

PETIVERIACEAE

§ FULLY CHLOROTROPHIC

Genera/species 9/13–20 Distribution Florida, Caribbean, Central and South America, tropical Africa (some species of Hilleria), Australia, Vanuatu and New Caledonia (Monococcus). Habit usually bisexual (in Ledenbergia and Monococcus dioecious), evergreen trees (in Seguieria with spines), shrubs or lianas, perennial herbs (sometimes with lignified base).

Petiveria and some species of Gallesia are used in traditional medicine.

Key to genera of families in Phytolaccaceae s.l.

1. Ovary of 5-16 carpels and seeds ------------ Phytolaccaceae s.s.

1. Ovary of a single carpel and seed - 2

2. Ovary with 2-4 stigmas ------------ Microteaceae

2. Ovary with a single stigma ------------ Petiveriaceae

Key to genera Petiveriaceae at South America

1. Flowers slightly zygomorphic, upper tepal free, the lower three tepals connate at the base into a three-lobed lip ------------ Hilleria

1. Flowers actinomorphic, all tepals free - 2

2. Ovary with 4-6 recurved, awn -like protuberances ------------ Petiveria

2. Ovary without protuberances - 3

3. Inflorescence paniculate; fruit a samara - 4

4. Plants usually spiny; tepals 5, herbaceous and reflexed in fruit ------------ Seguieria

4. Plants unarmed; tepals 4, woody and erect in fruit ------------ Gallesia

3. Inflorescence spiciform or racemose; fruit a drupe or utricle - 5

5. Fruit a drupe - 6

6. Stamens 4; stigma capitate ------------ Rivinia

6. Stamens 8-25; stigma penicillate ------------ Trichostigma

5. Fruit a utricle - 7

7. Racemes pendulous; bract placed halfway or above between the flower and the base of the pedicel ------------ Ledenbergia

7. Racemes erect; bract placed at the base of the pedicel ------------ Schindleria

SYSTEMATICS outsider Monococcus (1; SE Queensland, NE New South Wales, New Caledonia, Vanuatu).

1. Gallesia Casar. Trees, up to 30 m tall; smelling of garlic. Only one sp., G. integrifolia (Spreng.) Harms, South America in Bolivia, Brazil, Ecuador, and Peru, widely distributed, mainly most forests and gallery forests.

2. Hilleria Vell. Herbs, sometimes suffrutescent at the base; flowers in racemes. Three spp., one endemic to Peru, H. secunda (Ruiz & Pav.) H. Walter in Andes from Venezuela to Bolivia, and H. latifolia (Lam.) H. Walter in tropical South America to S Brazil and Argentina, also in tropical Africa, Madagascar and Mascarene Islands.

3. Ledenbergia Klotzsch. ex. Moq. Trees or shrubs, flowers in racemes. Three spp., L. macrantha Standl. in Mexico to Central America, L. peruviana O.C. Schmidt in Peru and Ecuador, and L. seguierioides Klotzsch ex Moq. endemic to Venezuela.

4. Petiveria L. Herbs or subshrubs, slightly woody at the base; flowers in spike to spike-like racemes. Only one sp., P. alliacea L., widely distributed in tropical and subtropical America from Florida and the Antilles to Argentina and Brazil, two subsp., one diploid and another tetraploid.

5. Rivina L. Herbs or subshrubs, woody at the base. Only one sp., R. humiliis L., widely distributed in tropical and subtropical America from the southern U.S.A. and the Antilles to Argentina and Chile, also in Brazil.

6. Schindleria H. Walter. Shrubs, rarely subshrubs, flowers in racemes, bisexual. Three spp., Peru, Bolivia, Paraguay and N Argentina.

7. Seguieria Loefl. Trees, slightly scandent shrubs, lianas, nearly always spine. 6 spp., distributed from Trinidad and Central America to Argentina and Guianas, mainly open areas; 5 spp. in Brazil (one endemic), S. brevithyrsa H. Walter endemic to Bolivia.

8. Trichostigma A. Rich. Shrubs or vines, rarely tree-like; flowers in racemes, bisexual. Three spp., two more restricted from Central America to Ecuador, and T. octandrum (L.) H. Walter distributed from Mexico and the Antilles to Argentina and Brazil.

NYCTAGINACEAE

§ FULLY CHLOROTROPHIC

Genera/species 31/300–400 Distribution tropical, subtropical and warm-temperate regions in the Northern and Southern Hemispheres, with their largest diversity in North and South America; Phaeoptilum in SW Africa. Habit Usually bisexual (rarely monoecious, andromonoecious, gynomonoecious or dioecious), evergreen or deciduous trees (Leucastereae), shrubs or lianas, perennial or annual herbs. Roots sometimes fleshy or tuberous.

The Nyctaginaceae are distributed mostly in the tropics and subtropics of the New World and comprise approximately 32 genera and 400 species. Of the 23 genera only Boerhavia, Commicarpus, Phaeoptilum, Pisonia and one species of Mirabilis also occur in the Old World. Only one genus is endemic to Old World: Phaeoptilum, monotypic in Africa. In the Neotropics there are 27 genera and approximately 200 species. Selinocarpus is highly disjunct from N Mexico and SW U.S.A. (8), and Somalia (1).

As in many caryophyllid families, e.g., Amaranthaceae and Portulacaeae, Nyctaginaceae is a propensity in many Nyctaginaceae to be tolerant of, or specialists of, gypseous soils. Outcrops of gypsum (hydrous calcium sulfate) are quite common in arid North America, especially in the Chihuahuan Desert. These areas have a flora characterized by gypsophiles, which never occur on other substrates, and gypsum-tolerant species, which are found on both gypseous and nongypseous soils. In the U.S.A. and Mexico, Nyctaginaceae are well represented in gypsum communities. At least 25 species in seven genera are known to occur on gypsum. Of these, roughly half are known gypsophiles, found only on gypsum soils.

Distinguishing characters (always present): sepals united to form a tube; inflorescences often subtended by conspicuous involucre or flowers sometimes subtended by sepal -like bracts; corolla absent; fruits often surrounded by accrescent perianth tube, with a single seed (anthocarps); inflorescences terminal or axillary, variously branched but usually ultimate branches cymose, paniculate or sometimes capitate pseudanthia; bracts and bracteoles (1-3) present, sometimes very small and early caducous, calyx-like and subtending a single corolla-like calyx (Mirabilis) or corolloid and subtending a cluster of flowers (Bougainvillea).

SYSTEMATICS all subfamilies occur in South America.

1. SUBFAMILY LEUCASTEROIDEAE (4/5) ‣ all genera are South American endemics.

1. Andradea Allemão. Tall trees; inflorescense paniculate; flowers petaloid. Only one sp., A. floribunda Fr. Allemao, endemic to SE Brazil and Bahia state.

2. Leucaster Choisy. Scandent shrubs, densely covered with stellate hairs on younger parts; flowers rotate. Only one sp., L. caniflorus Choisy, endemic to SE Brazil.

3. Ramisia Glaz. ex Baill. Tree; flowers in short axillary paniculate cymes; flowers petaloid. Only one sp., R. brasiliensis Oliv., endemic to SE Brazil.

4. Reichenbachia Spreng. Shrubs or small trees; flowers tomentose, tubular. Two spp., R. hirsuta Spreng. from Colombia to Bolivia, and R. paraguayensis (D. Parodi) Dugand & Daniel of Mato Grosso do Sul state in C Brazil, Argentina, Paraguay and Uruguay.

2. SUBFAMILY BOLDOOIDEAE (2/6) ‣ both genera in South America.

5. Salpianthus Bonpl. Herb, up to 2 m high, campanulate perianth (2.0-3.5 mm long) with glandular and uncinated hairs. 5 spp. from Mexico to Costa Rica, with S. purpurascens (Cav. ex Lag.) Hook. & Arn. up to Ecuador and Venezuela, also Caribbean.

6. Cryptocarpus H. B. K. Shrubs; leaves thicksh. Only one sp., C. pyriformis Kunth, in Galapagos, Ecuador, Peru and Bolivia, disjunct in Mexico.

3. SUBFAMILY NYCTAGINOIDEAE (25/390) ‣ 4 tribes, all in South America.

3.1 NYCTAGINOIDEAE ▸ TRIBE COLIGNONIEAE (1/6) - a single genus in this tribe.

7. Colignonia Endl. Subshrubs perennial, tuberous roots; flowers in cymose umbel-like partial inflorescense. 6 spp. from Colombia, Ecuador, Peru, Bolivia, Paraguay and Argentina.

3.2 NYCTAGINOIDEAE ▸ TRIBE BOUGAINVILLEAE (2/c 20) - outsider Phaeoptilum (1; Namibia, northern South Africa, Botswana).

8. Belemia Pires. Climbing shrubs with small potato-like tubers; flowers pink, bisexual. Two spp., B. fucsioides Pires from Espírito Santo and Minas Gerais states, and B. cordata Harley & Giul. endemic to Tocantins state, C Brazil.

9. Bougainvillea Comm. ex. Juss. Coarse climbers up to 25m long, often with superaxillary spines; inflorescence sorroundedeb by three sessile showy cordate bracts (true epiphylly). 11 spp., 9 from Ecuador to S Agentina and Paraguay (5 of them up to Brazil), and two endemics to Brazil.

Phylogenetic reconstructions based on plastid genomes showed that B. pachyphylla and B. peruviana (Peru, Ecuador) are basal taxa, while B. spinosa (Argentina, Bolivia, upper Paraguay, Peru) is sister to two distinct clades: the predominantly cultivated Bougainvillea clade (B. spectabilis, B. glabra, B. praecox, all form Brazil, the latter up to Argentina, Bolivia and Paraguay) and the clade containing wild species of Bougainvillea (B. berberidifolia, B. campanulata, B. infesta, B. modesta, B. stipitata, Argentina (3), Bolivia (5), Paraguay (3), Brazil (4)). B. spectabilis Willd. is native to eastern Brazil and widely cultivated throughout the world for the ornamental effect of its beautiful and colorful floral bracts.

3.3 NYCTAGINOIDEAE ▸ TRIBE PISONIEAE (7/c. 200) - outsdiers Ceodes (21, Africa, India to Australa), Grajalesia (1; Mexico), Neeopsis (1; Guatemala), Rockia (1, Hawaii).

10. Cephalotomandra Karst. & Triana. Shrubs or small trees; flowers yellow. Only one sp., C. fragrans H. Karst. & Triana, endemic to Colombia.

11. Guapira Aubl. (inc. Neea) Trees or shrubs, making ECM symbioses with fungi, puberulent becoming glabrous; flowers in terminal inflorescences. 157 spp., from Central to South America and Antilles, 115 in South America, 47 in Brazil, 28 endemics.

12. Pisonia L. Trees or shrubs or climbers, making ECM symbioses with fungi, with or without spines; flowers in paniculate cymes. 40 spp., pantropical, barely in Africa and Australia, 23 in New World, 6 in South America, three in Brazil, none endemics.

13. Pisoniella (Heimerl) Standl. Shrub or small tree with mostly regularly forked branches; inflorescence terminal, many flowered. Two spp., P. arborescens (Lag. & Rodr.) Standl. in Mexico and P. glabrata (Heimerl) Standl. from S Bolivia and N Argentina.

3.4 NYCTAGINOIDEAE ▸ TRIBE NYCTAGINEAE (12/240–260) - outsiders Acleisanthes (16; Sonoran and Chihuahuan deserts in SW U.S.A. and N Mexico, Somalia); Abronia (20–25; SW U.S.A., N Mexico), Tripterocalyx (4; SW Canada, W U.S.A.); Cyphomeris (2; New Mexico, Texas, N Mexico), Anulocaulis (5; SW U.S.A., N Mexico), Nyctaginia (1; New Mexico, Texas, N Mexico), Okenia (1–2; SE Florida, Mexico, Nicaragua), Cuscatlania (1; El Salvador).

14. Allionia L. Annual or perennial, prostrate herbs; flower triad forming a pseudanthium. Two spp., both highly widely distributed from North America and Mexico up to Southern Coast of western flanks of mainland South America, also in Uruguay.

15. Boerhavia L. Annual or perennial herbs, rarely wood at base. 61 spp., 30 only Old World, 21 spp. from U.S.A. to Central America, one in Hawaii, six endemics to South America, in Peru (3), Argentina (2) and Chile (1), B. coccinea Mill. and B. erecta L. native from U.S.A. to Bolivia, sometimes adventive in Brazil, and only native to Brazil, the widely distributed pantropical B. diffusa L.

Several authors (Fay 1980; Spellenberg 2001, 2003) have highlighted that at the species level this is a taxonomically difficult group due to morphological variation. Especially among annuals of the Sonoran desert and the pantropical B. diffusa Vahl and B. coccinea Mill. complex, apparently factors such as wide dispersal, hybridization, and autogamy have contributed to that variation; the genus is in need of a critical revision.

16. Commicarpus Standl. Subshrubs, leaves often fleshy; flowers in capitula, umbels or racemes. 25 spp., Africa (except extreme N and S), S Spain, Burma-China-Malaysia region, and 5 in New World, 3 restricted for Mexico, C. scandens (L.) Standl. from North America to Peru, Venezuela and Caribbean, and C. tuberosus (Lam.) Standl. in northern Andes.

17. Mirabilis L. Annual or perennial herbs. Roots sometimes tuberous; flowers in dichasias. 53 spp., North America to N South America (14, Colombia to Argentina and Chile, only three in tropical Andes), M. himalaica (Edgew.) Heimerl. in Himalaias.

LINEAGE 7: PORTULACIDS

MOLLUGINACEAE

§ FULLY CHLOROTROPHIC

Genera/species 11/c. 90 Distribution mainly tropical and subtropical regions, with their highest diversity in southern Africa; some species in warm-temperate areas. Habit usually bisexual (in Mollugo species dioecious), usually annual or perennial herbs (sometimes suffrutices). Often somewhat succulent.

All are found in dry, open Neotropical environments. Mollugo verticillata L. and Glinus radiatus (Ruiz & Pav.) Rohrb. are weedy and commonly found in disturbed habitats. Molluginaceae is more diverse in Africa, and some species of Mollugo are used as herbs in cooking.

Key differences from similar families may be confused with Galium L. and other herbaceous Rubiaceae because of its general habit, but has superior ovary and free petals (Rubiaceae has inferior ovary and gamopetalous corolla); it can be confused with Aizoaceae and Portulacaceae, but is less succulent and has smaller, less colourful flowers than those.

SYSTEMATICS outsiders Trigastrotheca (3; tropical and subtropical Asia and Australia), Paramollugo (6; one almost pantropical, one restricted to Somalia, one to Madagascar, and three to Caribbean), Hypertelis (5; SW and SE Europe, tropical and subtropical regions, South Africa), Polpoda (2; W Cape), Adenogramma (10–11; N and W Cape), Psammotropha (11; tropical and S Africa), Suessenguthiella (1; Namibia, N and W Cape), Coelanthum (3; S Namibia, N and W Cape), Pharnaceum (c 28; S Africa).

1. Glinus L. Much-branched annual herbs, often forming mats. 6 spp., three in Africa, one of then up to Arabia, another up to Australia; one endemic to Australia; one in SE Asia; and one in New World, a single weedy, widely distributed species, G. radiatus (Ruiz & Pav.) Rohrb., a prostrate semi-succulent herb with purplish leaves and stellate white flowers.

2. Mollugo L. (inc. Glischrothamnus) Annual or perennial herbs, small shrubs or subshrubs in Brazilan M. ulei (Pilg.) Thulin, rarely dioecious (also M. ulei), glabrous or with indumentum of glandular hairs; flowers in seemingly axillary, sessile or pedunculate, umbel- or raceme-like cymes, or flowers solitary; bracts small, membranous or partly herbaceous. c. 15 spp., 14 in New World: 6 restricted of Caribbean, three endemics to Brazil (M. ulei (Pilg.) Thulin is a rare plant in Brazil, by Plantas Raras do Brasil’s book), three endemics to Galapagos Is., M. snodgrassii B.L. Rob. in Galapagos and Peru; and M. verticillata L. is widely distributed.

HALOPHYTACEAE

§ FULLY CHLOROTROPHIC

Genera/species 1/1 Distribution endemic to Argentina. Habit herbs.

SYSTEMATICS a single argentine species.

1. Halophytum Speg. Annual, glabrous, leaf-succulent monoecious herbs; leaves sessile, alternate, but occasionally fascicled on short shoots; flowers unisexual, small, usually with 2 (male flowers) or 2–4 (female flowers) bracts or bracteoles, female flowers 4–5 together in the axils of upper leaves, male flowers numerous, densely aggregated in a condensed spike-like inflorescence from the axils of the upper leaves; sepaloids none; petaloids none in female flowers, 4 in male flowers, fruit a thin-walled, indehiscent, 1-seeded nutlet partly. Only one sp., H. ameghinoi Speg., from Argentina (Catamarca, Chubut, La Rioja, and Mendoza).

MONTIACEAE

§ FULLY CHLOROTROPHIC

Genera/species 16/275 Distribution cosmopolitan except polar areas (mainly temperate regions on the Northern Hemisphere), with their largest diversity in western North America, western and southern South America, southern Australia, Tasmania, New Zealand, and subantarctic islands; Habit usually bisexual (rarely unisexual), perennial or annual herbs (rarely suffrutices), often succulent. Roots sometimes tuberous. Stem frequently absent. Some species are xerophytic.

SYSTEMATIC 16 genera in four main clades, Rumicastrum Clade (1/66) is endemic to Australia, three remaining in South America, with 98 spp. in region.

1. TRIBE CISTANTHEAE (5/74) ‣ outsiders are Calyptridium (11, Canada, U.S.A. to Mexico), Thingia (1, U.S.A. to Mexico).

1. Cistanthe Spach. Herbs or shrubby succulents. 43 spp., two in North America and Mexico, 41 from Peru, Argentina and Chile, in two sections:

§ sect. Cistanthe ‣ 12 spp., all but one perennial, primarily in Chile, one extending into San Juan Province, Argentina, and two endemic to Peru.

§ sect. Rosulatae ‣ 29 annual and perennial species, mainly Chile, four extending into Argentina, one primarily in Peru but now collected in Chile, one endemic to Peru, and two in SW North America.

2. Lenzia Philippi. Small perennial acaulescent herb, chaemophyte, largely membranous awl-shaped leaves, scarcely 2-3 cm tall, solitary axillary flowers. Only one sp., L. chamaepitys Phil., Bolivia, Chile and Argentina at very high altitudes in fjaeldmark, observed to occur among the highest elevation plant species (4,200 m).

3. Montiopsis Kuntze. Perennial or annuals. 18 spp. in Chile and bordering regions of Argentina, often in high altitudes, in two subgenera:

§ subg. Montiopsis ‣ 15 spp., mainly Chile and Argentina, M. cumingii (Hook. & Arn.) D.I. Ford up to Peru, and M. polycarpoides (Phil.) Peralta up to Bolivia.

§ subg. Dianthoideae ‣ 3 spp., Argentina and Chile.

2. TRIBE MONTIOIDEAE (8/109) ‣ outsiders are Claytonia (33, temperate Asia, North America), Erocallis (1, Canada to NW U.S.A.), Hectorella (1, New Zealand), Lyallia (1, Kerguelan), Lewisia (16, Yuko to N Mexico), Lewisiopsis (1, Canada to NW U.S.A.), Rumicastrum (66, New Guinea to Australia).

4. Calandrinia H.B.K. Herbs to subshrubs, sometimes cushions. 36 spp., 3 only in North America to Mexico, two from North to South America, and 31 only in South America, 33 spp. in South America, more than 3,000 m high in tropical Andes.

5. Montia L. Annual herbs, prostrate or decumbent, rooting at the nodes, often sub-aquatic. 19 spp. from North America, mainly western, some up to E Siberia, 1 in Australia and New Zealand, and 3 in South America. Three sections:

§ sect. Australiensis ‣ 10 spp., 8 in Australia/New Zealand and two in NW North America and SW Canada.

§ sect. Montia ‣ 6 spp., evidently originally of W North America, including the widespread M. fontana L. in Europe east to Siberia, south to the Middle East and North and E Africa, also in Irian Jaya, Papua New Guinea, Australia, New Zealand, northern and western North America and western South America up to Uruguay (possibly introduced); it occurs more or less throughout Europe from the Iberian Peninsula north to Iceland, east through Scandinavia to North European Russia and south to Mediterranean, the Ukraine and the Balkans; M. biapiculata Lourteig endemic to Colombia, M. meridensis Friedrich in high mountains in Colombia and Venezuela, and the apparently adventive M. chamissoi (Ledeb. ex Spreng.) Greene and M. parviflora (Moc. ex DC.) Greene.

§ sect. Montiastrum ‣ 4 spp., NW North America and NE Asia, including the adventive M. linearis (Douglas)Greene.

3. TRIBE PHEMERANTHEAE (2/26) ‣ two genera, both in South America.

6. Phemeranthus (Raf) DC. Herbs sometimes with tuberous roots; leaves mostly basal on a branched caudex, usually linear. 25 spp., mainly restricted of NW, SW U.S.A. (10 endemics) and Mexico (5 endemics, up to Oaxaca), including also six in both countries simultaneously, and P. punae (R.E. Fr.) Eggli & Nyffeler in N Argentina and S Bolivia.

7. Schreiteria Carolin. Perennial herb with tuberous roots; flowers in cymes. Only one sp., S. macrocarpa (Speg.) Carolin, endemic to Tucuman province in N Argentina; the placement of the monotypic Schreiteria in Montiaceae is preliminary; this enigmatic genus has not been found again in the past 80 years or so, and was not available for analysis.

BASELLACEAE

§ FULLY CHLOROTROPHIC

Genera/species 4/19 Distribution tropical and subtropical regions of America (most species) and Africa, Madagascar; introduced into South Asia, New Guinea and islands of the Pacific. Habit usually bisexual (rarely monoecious; some functionally unisexual), winding or climbing perennial herbs or lianas, slightly to distinctly fleshy. Stem bases and rhizome usually swollen and tuberous; roots sometimes tuberous. Perenial, sub-fleshy to succulent, herbaceous or occasionaly suffruticose vines, or sometimes erect plants. Small flowers with 2 petaloid sepals and 5 petals, subtended by 2 opposite bracteoles.

Ullucus tuberosus Caldas is an important high Andean food crop, and Basella alba L. is cultivated for its edible leaves. A few Anredera species are grown as ornamentals, also outside their native distributions. The homologies of the (1) bracteoles, (2) sepals, and (3) petals have been much debated without being definitely settled. Three of the four genera are endemic to New world. All species of Anredera, Tournonia, and Ullucus are native, and all except a few widely distributed Anredera species are endemic; the Basella species (all 5 in Africa) found in the Neotropics is cultivated and naturalised. Most species are found in open, dry habitats.

Key to genera of Neotropical Basellaceae

1. Leaf margin dentate by glands; inflorescence a dichasium ------------ Tournonia

1. Leaf margin entire; inflorescence a raceme, spike, or panicle - 2

2. Petals caudate at apex; anthers basifixed, dehiscent by short apical slits ------------ Ullucus

2. Petals obtuse at apex; anthers dorsifixed, dehischent by longitudinal slits ------------ Anredera

SYSTEMATICS outsider Basella (5; tropical E and SE Africa, Madagascar, tropical Asia from India to New Guinea, possibly introduced in tropical America and on oceanic islands).

1. Anredera Juss. Twining or scandent succulent twining vines, producing new shoots to tickend stem base or tubers. 12 spp. in South America, 8 restricted from Colombia to Argentina, two up to Mexico and North America, A. cordifolia (Ten.) Steenis in southern and central South America up to Brazil, and A. tucumanensis (Lillo & Hauman) Sperling, known from humid forests in S Ecuador, Bolivia, N Argentina, and SE Brazil; it is expected to occur also in Peru.

2. Tournonia Moq. Scandent succulent twining vines, producing new shoots to tickend stem base. Only one sp., T. hookeriana Moq., known paramo and semi-dry scrub to more humid forest at high altitudes from W Colombia to N Ecuador.

3. Ullucus Caldas. Erect or trailing succulent herbs, producing new shoots to tubers, edible by natives. Only one sp. with two subsp., U. tuberosus subsp. aborigineus (Brücher) Sprerling from high altitudes in Peru, Bolivia and N Argentina, and U. tuberosus subsp. tuberosus (Brücher) Sprerling cultivated from Venezuela to Chile.

TALINACEAE

§ FULLY CHLOROTROPHIC

Genera/species 2/28 Distribution Africa, Madagascar, North and South America, with their largest diversity in southern North America. Habit bisexual, usually small shrubs (sometimes perennial herbs, suffrutices or small trees, sometimes lianoid) with somewhat succulent leaves. Subterraneous organs often tuberous. Aerial parts often ephemeral.

SYSTEMATICS 2 genera, both in South America.

1. Amphipetalum Bacigalupo. Xerophytic scrub with a tuberous root, up to 40 cm tall. Only one sp., A. paraguayense Bacigalupo, endemic to SE Bolivia and Paraguay.

2. Talinum Adans. Shrublets with annual branches from a perennial base, usually tuberous; leaves basally in a rosette or semirosette; inflorescence normally a compact terminal panicle composed of branched lateral cymes, less commonly of terminal branched cymes or, rarely, of solitary axillary flowers; flowers opening during the day or during the night, small or large (0.5 – 2.2 cm diam.). 17 spp., 4 restricted of S Africa, T. portulacifolium lium (Forsk.) Aschers. ex Schweinf occur in Africa to India; 12 in New World, 6 in South America: three restricted of Bolivia, Paraguay and Argentina, two widely distributed (with T. paniculatum (Jacq.) Gaertn. in Brazil), and T. triangulare (Jacq.) Willd. in Brazil, Venezuela and Peru, cultivated as a salad plant.

T. polygaloides Gillies ex Arn., from Paraguay, Argentina, Bolivia and Chile, is anomalous having woody stems c. 40 cm high with only the leaves succulent; additionally its flowers are much larger than in other species and the seeds have a different ornamentation; although it is very different from the other Bolivian species, molecular studies support its retention in Talinum.

PORTULACACEAE

§ FULLY CHLOROTROPHIC

Genera/species 1/152. Distribution cosmopolitan. Habit bisexual, perennial or annual herbs (in Portulaca suffrutescens Engelm. somewhat lignified at stem base). Roots often tuberous. Usually leaf succulents. Use Ornamental plants, vegetables, medicinal plants. Only one genus.

SYSTEMATICS a single genus.

1. Portulaca L. Usually succulent herbs mostly with tuberous roots. 116 spp., cosmopolitan, with two subgenera, one endemic to Australia, the other cosmopolitan; 75 in New World, 56 in South America, 16 in Brazil, 7 endemics. P. hirsutissima Cambess. and P. confertifolia Hauman (Argentina) has hairs in leaf limb, unique with this feature in New World Portulacaceae, like some African species; P. cryptopetala Speg. (Bolivia to Uruguay) is only species of genus with C₃ – C₄ photosyntesis. P. hatschbachii Legr from Paraná state is a rare plant in Brazil, by Plantas Raras do Brasil’s book

ANACAMPSEROTACEAE

§ FULLY CHLOROTROPHIC

Genera/species 3/63 Distribution southern and southeastern Africa, Somalia, southwestern Arabian Peninsula, southwestern United States, Mexico, Bolivia, Argentina, with their largest diversity in South Africa. Habit bisexual, small shrubs or suffrutices to thick-stemmed perennial herbs. Sometimes with a basal fleshy caudex or tuberous main-root. Often stem succulents.

SYSTEMATICS outsider Talinopsis (1; S U.S.A., Mexico).

1. Anacampseros L. 59 spp., 53 in SE and S Africa, two only in NE Africa, one in Australia, A. coahuilensis (S.Watson) Eggli & Nyffeler in N Mexico and SW U.S.A., A. vulcanensis Añon, and A. kurtzii Bucigalupo in N Argentina and S Bolivia.

2. Grahamia Hook. Only one sp., G. bracteata Hook & Arn., endemic to N and C Argentina.

CACTACEAE

§ FULLY CHLOROTROPHIC

Genera/species 150/1,862 Distribution mainly arid and semiarid regions of North and South America (British Columbia and Alberta southwards to Patagonia). Habit usually bisexual (at least in Mammillaria dioica K. Brandegee and Selenicereus innesii Kimnach functionally dioecious), usually perennial herbs (sometimes climbing or epiphytic; rarely deciduous shrubs or small trees (Pereskia)). Usually xerophytic. Cactaceae is the second largest group of succulent species worldwide (after Aizoaceae); almost all species are stem succulents with elongate and branched or unbranched, or almost globular pachycaul photosynthesizing stem; some genera with flattened almost foliaceous stem segments, phylloclades, such as the well-known Christmas cactus or Thanksgiving cactus (in the genus Schlumbergera). Stem and branch surfaces usually with areolae – modified axillary short shoots, brachyblasts – with numerous spines – modified leaves or foliar lobes. Cacti occur in a wide range of shapes and sizes. The tallest free-standing cactus is Pachycereus pringlei (S.Watson) Britton & Rose, with a maximum recorded height of 19.2 m (63 ft),(2) and the smallest is Blossfeldia liliputiana Werderm, only about 1 cm (0.4 in) in diameter at maturity.

The word ‘cactus’ derives, through Latin, from the Ancient Greek κάκτος (kaktos), a name originally used for a spiny plant whose identity is not certain. Cultivated under glass in temperate regions, naturalised outdoors in the Paleotropics (in Australia some Opuntia became weeds), especially Opuntia ficus-indica Mill. (from Mexico but widely cultivated and introduced in Mediterranean regions, nowadays part of the culture in Europe); Lophophora williamsii, commonly known as peyote, is noted for its psychotropic alkaloids.

Key differences from similar families rarely present latex (different from succulent Euphorbiaceae); spines are produced in areoles (different from other stem succulents like Asclepiadaceae, Euphorbiaceae, etc.). DNA studies show that Portulacaceae possibly cannot be separated from Cactaceae because it lacks autoapomorphies; Cactaceae however presents areoles which are not present as such in Portulacaceae.

SOME DATA One the most striking features of cacti is their high evels of endemisms at both generic and species levels. Brazil, Mexico and Pehave the highest levels of generic endemisms, at 40% and 30% respectively. Astoundingly, however, 78,9% of all cacti occurring Mexico (536), and 67,34% of Brazilian species (198) are found within these respective countries and nowhere else in the world. 182 in North America, 688 in Mexico, 279 in Peru, 254 in Brazil (in 40 genera), 233 in Bolivia. 1884 spp. in New World. 1027 cacti occur in South America.

SYSTEMATIC all lineages occur in South America.

1. SUBFAMILY PERESKIOIDEAE (1/10) ‣ a single genus.

1. Pereskia Mill. Trees, shrubs or climbers with well developed leaves, flowers without developed tube, white, orange, pink or red, fruits indehiscent with large seeds. 10 spp. from South America, P. aculeata Mill. in over tropical South America up to Mexico and Caribbean; endemics in Brazil (3), Bolivia (3), Peru (1); remaining two species in S Brazil, Argentina, Paraguay and Bolivia. P. stenantha F. Ritter has unique in the genus due to their urceolate corolla and larger nectary, indicative of hummingbird pollination; P. nemorosa Rojas Acosta of Brazil, Paraguay, NE Argentina, and Uruguay has the largest flowers in the genus.

2. SUBFAMILY LEUENBERGERIOIDEAE (1/8) ‣ a single genus.

2. Leuenbergeria Lodé. Small trees and shrubs, may be lianas, with precocious development of the cork cambium and without stomata in the stems (not xeric features in plants); yellow flowers in two spp., one in Brazil, another in Colombia/Venezuela; over all others cactus has cauline stomata. 8 spp., three in Hispaniola, one in Cuba, 1 from Pacific coastal area from S Mexico to Costa Rica in lowland dry forest from sea level to 1,000 m, and three in South America: L. aureiflora (F. Ritter) Lodé in dry seasonal scrubland of NE Brazil (caatinga), at NE Minas Gerais and S Bahia, at elevations of 300–700 m; L. bleo (Kutnh) Lodé in Panamá and Colombia along rivers and in secondary forests, from sea level to 500 m; and L. guamacho (F.A.C. Weber) Lodé in drier regions of Colombia and Venezuela, from sea level to 800 m; possibly also the Dutch Antilles.

3. SUBFAMILY MAIHUENIOIDEAE (1/2) ‣ a single genus.

3. Maihuenia (Philippi ex F. A. C. Weber) Schumann. Low, caespitose shrubs, resembling Opuntia (Maihueniopsis) spp.; stems succulent, globose or short cylindric; leaves small, terete, persistent. Two spp., Patagonia in S Chile and S Argentina.

4. SUBFAMILY OPUNTIOIDEAE (18/325) ‣ three tribes, all in South America.

4.1 OPUNTIOIDEAE ▸ TRIBE OPUNTIEAE (7/185) - outsider Consolea (9; Florida Keys, Caribbean).

4. Airampoa Fric. Clump-form shrubs (cushions) with determinate cylindrical or flattened stems. 16 spp. (2 or 5 in some sources) from S Peru to N Argentina, Chile and Bolivia.

5. Brasiliopuntia (K.Schum.) A.Berger. Tree-like; bee-pollinated, bowl-shaped flowers with yellow or greenish-yellow petals, quite unlike the hummingbird-pollinated flowers of Nopalea and Tacinga. Two spp., from Bolivia, Brazil, Paraguay, Uruguay and Argentina, growing at the periphery of dense forests.

6. Miqueliopuntia Fric. ex F.Ritter. Clump-form shrubs with determinate cylindrical stems. Only one sp., M. miquelii (Monv.) F. Ritter, endemic to N Chile.

7. Opuntia Mill. (exc. Salmiopuntia, Tacinga p.p., inc. Platyopuntia) Shrubs to tree-like plants with reduced, sometimes cushions, scale-like leaves and flattened pads with glochids and serrate spines, flower with well defined green pericarpel. 175 spp. (154 in some sources), from Canada to Argentina, 54 in South America, 5 in Brazil, none endemics; the divergent O. schickendantzii F.A.C.Weber occurs in Argentina in Catamarca, Jujuy, Salta, and Tucumán provinces, Paraguay and in Cochabamba in Bolivia; it occurs at elevations 1,000 – 2,000 m, and maybe a distinct genus.

Eight major clades have been recovered within Opuntia s. str., and the South American species are mainly nested in two of these clades: Macbridei (northern part of South America, from central Peru to central Colombia) and Elatae (southern South American lineages occupying mainly the Pampa and the Chaco regions, as well as the Galapagos Islands). Opuntia leoglossa Font & M.Köhler is known only invasive populations in Australia and Spain.

8. Salmonopuntia Fric. (off. Opuntia) Only one sp. (two in some sources), S. salmiana (J. Parm. ex Pfeiff.) Fric., from Argentina, Paraguay and Bolivia.

9. Tacinga Britton & Rose. (off. Opuntia) Prickly-pear cacti, stems alender cylindric, scrambling, cane-like; some of its species have a unique growth form among the prickly pears, that of a vine that scrambles over other plants for support. 10 spp., one endemic to NE Venezuela in Lara and Sucre states, and nine restricted for NE Brazil, one of them, from Minas Gerais state, is a rare plant in Brazil, by Plantas Raras do Brasil’s book; it is only opuntioid genus found in the semiarid regions of the E corner of Brazil.

4.2 OPUNTIOIDEAE ▸ TRIBE TEPHROCACTEAE (6/70) – all genera in South America.

10. Austrocylindropuntia Backeberg. Low plants, sometimes cushions; 7 spp. from Colombia to W Argentina, in montane dry environments

11. Cumulopuntia Rotter. Low plants, sometimes cushions; 12 spp., Andes of Peru, Bolivia and Cono Sur.

12. Maihueniopsis Spegazzini. Compact cushions, with small ovate to obconic cladodes and areoles evenly distributed upon the cladode. 20 spp. from Cono Sur, three up to S Bolivia.

13. Pterocactus Schumann. Dwarf, almost geophytic shrubs; rootstock usually large, tuberous; stems segments globose, cylindric or clavate. 10 spp., Patagonia in S and W Argentina and adjacent Chile (1).

14. Punotia R. Kiesling. Cushion. Only one sp., P. lagopus (K.Schumann) D.R.Hunt, from Peru and Bolivia.

15. Tephrocactus Lemaire. 12 spp., 11 from Argentina and one endemic to Peru.

4.3 OPUNTIOIDEAE ▸ TRIBE CYLINDROPUNTIEAE (5/70) - outsiders Micropuntia (1; Mojave Desert in SW U.S.A.), Pereskiopsis (6; Mexico, Central America), Grusonia (21; SW U.S.A., NW Mexico).

16. Cylindropuntia Rotter. 40 spp., Meixco to Central America, Caribbean, U.S.A., two up to South America, C. caribaea (Britton & Rose) F.M. Knuth also in Venezuela, and C. tunicata (Lehm.) F.M. Knuth disjunct North America and Ecuador.

17. Quiabentia Britton & Rose. Shrub or tree-like; stems often verticillate, cylindric terete; leaves flat, fleshy. Two spp., the tree Q. verticillata (Vaupel) Vaupel ex Berger in W Paraguay and nearby portions of Bolivia and Argentina, and the shrubby, Pereskiopsis-like Q. zehntneri (Britton & Rose) Britton & Rose endemic to S Bahia and N Minas Gerais, near São Francisco River, Brazil.

5. SUBFAMILY CACTOIDEAE (111/1.425–1515) – 7 lineages, all in South America.

CACTOIDEAE ▸ UNPLACED CACTOIDEAE (lack informations)

18. Cremnocereus M. Lowry, M. Winberg & J. Gutierrez. Shrub with stems branching from near the base; stems cylindrical, a few metres long with ribs bearing areoles several millimetres apart; flowers borne close to the stem apex, tubular, erect; tepals flesh coloured or white. Only one sp., C. albipilosus M. Lowry & Winberg, endemic to dry forests of Bolivia

5.1 CACTOIDEAE ▸ TRIBE BLOSSFELDIEAE (1/1) - a single genus.

19. Blossfeldia Wedermann. Tinly, button-like, only about 10-12 mm in diameter at maturity, the smallest of all cacti; stem simple or caespitose, not ribbed or tuberculate, poikilohydric; spine absent. Only one sp., B. liliputana Werderm., from the eastern Andes in W Bolivia and NW Argentina; is the one from two dissecation tolerant species dicots in New World, joined Clinopodium giliesii (Benth.) Kuntze (Lamiaceae).

5.2 CACTOIDEAE ▸ TRIBE CACTEAE (26/413) - outsiders Acharagma (2; N Mexico), Ariocarpus (7; S Texas and N Mexico), Astrophytum (6; S Texas and N Mexico); Aztekium (3; NE Mexico), Cochemiea (38, Mexico), Coryphantha (43; SW and S U.S.A., Mexico), Cumarinia (1; Mexico), Echinocactus (2; SW U.S.A., Mexico), Epithelantha (10; Texas, Arizona and N Mexico), Escobaria (17; SW Canada to Mexico, Cuba), Ferocactus (29; SW U U.S.A., Mexico), Geohintonia (1; NE Mexico), Homalocephala (3, SW U.S.A., Mexico), Kadenicarpus (2, Mexico), Kroenleinia (1, Mexico), Leuchtenbergia (1; N Mexico), Lophophora (8; S Texas, N and E Mexico), Neolloydia (5; Texas and N Mexico), Obregonia (1; NE Mexico), Ortegocactus (1; S Mexico), Pediocactus (9; W U.S.A.), Pelecyphora (2; N Mexico), Rapicactus (5; Mexico), Sclerocactus (23; SW U.S.A., N Mexico), Stenocactus (21; Mexico), Strombocactus (2; C Mexico), Thelocactus (11; Texas, Mexico), Turbinicarpus (23; NC Mexico). North America, with their highest diversity in the Chihuahuan desert in Texas and northern Mexico. 15 genera endemic from Mexico, one endemic U.S.A., ten shared with U.S.A., one reaching into Cuba (Escobaria, 23) and one reaching into South America..

20. Mammillaria Haworth. Small globose to cylindric, with tubercles and two different types of areoles, latex sometimes present, small flowers appearing in rings around the stem. 179 spp. (143 in Korotkova, 2021), SW U.S.A., Caribbean, and Mexico (172, 159 endemics), two up to N Colombia and coast of Venezuela.

5.3 CACTOIDEAE ▸ TRIBE LYMANBENSONIEAE (3/40) - all genera confined to South America.

21. Calymmanthium Ritter. Shrubby or arborescent; stems segmented, 3-4 winged, spiny, especially the old stems. Only one sp. from Cajamarca in N Peru.

22. Copiapoa Britton & Rose. Low growing or mound-forming, very small or to 1 m, sometimes cushions; rootstock fibrous or with greatly enlarged taperoot. 34 spp., coastal deserts in N Chile.

23. Lymanbensonia Kimnach. Terrestrial or epiphytic, epiphytic habit obligatory or facultative; plants usually erect at first, then spreading, pendent; flowers usually solitary, rarely 2 per areole, lateral, actinomorphic, 1.2 to 3 cm long, narrowly tubular bell-shaped, tepals not fully expanding, spreading at apices perianth intensely coloured (red, pink, orange, magenta) or white. 5 spp., from S Ecuador (Loja) to central and S Peru (Amazonas, Junín, Puno) and the eastern Andes of Bolivia (La Paz; Cochabamba, Santa Cruz).

5.4 CACTOIDEAE ▸ TRIBE PHYLLOCACTEAE (33/342) - all lineages in South America except Pachycereae (13-16/c. 170, SW U.S.A., Mexico to C Central America). Outsiders except members of Hylocereae are Bergerocactus (1; California and Baja California), Carnegiea (1; Sonoran desert in southern Arizona, S California and N Mexico), Cephalocereus (16; E and S Mexico), Isolatocereus (1; C Mexico), Echinocereus (83; South Dakota to Mexico), Escontria (1; S Mexico), Lemairecereus (2; Mexico to Honduras), Lophocereus (3; U.S.A. to Mexico), Marshallocereus (1; Central America), Morangaya (1; NW Mexico), Myrtillocactus (4; Mexico, Guatemala), Nyctocereus (1; Mexico), Pachycereus (7; SW U.S.A., Mexico, N Central America), Peniocereus (8; SW North America, Mexico, Central America), Polaskia (2; S Mexico).

∎ FRAILEA CLADE (1/18) ‣ a single genus.

24. Frailea Britton & Rose. Low growing, caespitose or unbranched; stems sometimes tuberculous (i.e., without ribs), flowers yellow, diurnal. 18 spp., in Korotkova, 2021, Colombia (one record of F. pumila (Lem.) Britton & Rose, highly dubious), Bolivia, Argentina, S Brazil (12, 4 endemics, highly centered in Rio Grande do Sul state, two of them rare plants in Brazil, by Plantas Raras do Brasil’s book), Paraguay and Uruguay.

F. castanea Backeberg, resembling a miniature Euphorbia obesa Hook.f. or Astrophytum asterias (Zucc.) Lem., with its body flat with the ground, and having many low ribs with strings of areoles bearing very short black spines; the species grows in areas of shallow reddish soil with rock outcrops of iron-rich sandstone; F. pygmaea (Spegazzini) Britton & Rose, as the name implies (pygmaeus meaning dwarf), this is perhaps the smallest of all Frailea species, some forms having heads no wider than one or two centimetres; it is also quite widely distributed, growing in S Brazil, Uruguay and Argentina.

∎ AUSTROCACTUS + EULYCHNIA CLADE ‣ both genera in South America.

25. Austrocactus Britton & Rose. 10 spp., Chile and S Argentina.

26. Eulychnia Phil. 7 spp., one in coastal deserts in Peru, and remaining six in N Chile.

∎ PFEIFFERA CLADE ‣ lack informations about outsiders.

27. Pfeiffera Salm-Dyck. Epiphytic, rarely epilithic or terrestrial; epiphytic habit mostly obligatory; flowers usually solitary, rarely 2 per areole, lateral, sometimes subterminal, actinomorphic, funnel-shaped or broad-campanulate, mostly 1 – 2 cm in diameter. 6 spp. from Bolivia (La Paz, Cochabamba, Santa Cruz, Chuquisaca and Tarija) to N Argentina (Jujuy, Salta, and Tucumán).

∎ CORRYOCACTUS CLADE ‣ lack informations about outsiders.

28. Corryocactus Britton & Rose. 13 spp., 10 from S Peru (one up to N Chile) and 3 endemics to Bolivia.

∎ ARMATOCEREUS CLADE (4/29) ‣ outsider Leptocereus (20; Cuba, Hispaniola, Puerto Rico).

29. Armatocereus Backeb. 7 spp. from Ecuador and Peru.

30. Brachycereus Britton & Rose. Shrub; lava cactus. Only one sp., B. nesioticus (K. Schum.) Backeb., endemic to Galapagos, Ecuador.

31. Jasminocereus Britton & Rose. Tree. Only one sp., J. thouarsii (F.A.C. Weber) Backeb, endemic to Galapagos, Ecuador.

∎ STROPHACACTUS + NEORAIMONDIA CLADE (2/4) ‣ both genera in South America.

32. Neoraimondia Britton & Rose. Two spp., one in Peru, another in Bolivia.

33. Strophocactus Britton & Rose. Weakly scandent, terrestrial prostrate; flowers funnelform, nocturnal, white. Three spp., S. brasiliensis (Britton & Rose) S. Arias & N. Korotkova restricted from dry areas in Bahia and Minas Gerais states in E Brazil, S. sicariguensis (Croizat & Tamayo) S. Arias & N. Korotkova from Lara, Venezuela, and Colombia, and the very odd S. wittii (K. Schum.) Britton & Rose, from Amazon rainforest in high waterline of black water rivers; this species is a water-dispersed seed, a feature unique in Cactaceae.

∎ ACANTOCEREUS CLADE ‣ lack informations about outsiders.

34. Acanthocereus (A. Berger) Britton & Rose. 15 spp., inc. Florida, S Mexico, Central America Caribbean, and A. tetragonus (L.) Hummelinck up to coastal Venezuela and Colombia.

∎ SUBTRIBE HYLOCEREINAE (8/84) ‣ outsiders Aporocactus (2; Mexico), Disocactus (16; Mexico to Central America), Deamia (3; Mexico to Nicaragua).

35. Epiphyllum Haw. Mostly epiphytic, some lithophytic, leaf-like; flowers similar to Hylocereus and Selenicereus, large white. 10 spp., mostly Central America, reaching to South America to Argentina with 5, Colombia and Peru one endemic each, and three more expanded, two of then in Brazil, E. oxypetalum (DC.) Haw, and E. phyllanthus (L.) Haw., none endemics.

36. Kimnachia S. Arias & N. Korotkova. Shrubby with pendent stems, differentiated into primary and secondary stems (dimorphic), lacking spines; flowers actinomorphic, 1–2 per areole, 7–12 mm long and for up to 15 mm in diameter, green to reddish brown, perianth whitish/yellowish; fruit globose to ovoid, 4-8 mm in diameter, whitish, pulp whitish. Only one sp., K. ramulosa (Salm-Dyck) S. Arias & N. Korotkova, Mexico to Bolivia, N Brazil and Venezuela.

37. Pseudorhipsalis Britton & Rose. Epiphytic; stems leaf-like, spineless; flowers small, diurnal, white or yellow. 5 spp., four in Central America (two up to N South America to Peru and Guianas, P. amazonica (K. Schum.) Ralf Bauer up to Amazonas, Roraima and Pará states in N Brazil), one only in Caribbean.

38. Selenicereus (A. Berger) Britton & Rose. Scandent, epilithic or epiphytic shrubs; large flowers, nocturnal; stems often 5 m or more, often producing aerial roots. 31 spp., S Mexico, Central America, N South America - 6 spp., 5 from Venezuela to Peru, and S. setaceus (Salm-Dyck) A. Berger ex Werderm. from Brazil, Bolivia and Cono Sur.

39. Weberocereus Britton & Rose. Night-blooming, epiphytic or lithophytic, climbing or sprawling shrubs. 6 spp., 5 in Central America, and W. rosei (Kimnach) Buxb. endemic to Ecuador.

∎ SUBTRIBE ECHINOCEREINAE ‣ lack informations about outsiders.

40. Stenocereus Riccobono. 24 spp., S U.S.A., Mexico, Central and N South America (2 in Colombia, Venezuela), Caribbean.

5.5. CACTOIDEAE ▸ TRIBE RHIPSALIDEAE (4/61) - all genera in South America.

41. Hatiora Britton & Rose. Epiphytic or epilitic shrubs; stems cyllindric, angled, winged or flat, segmented. Two spp., both endemics to SE Brazil.

42. Lepismium Pfeiff. Epiphytic or epilitic shrubs, creeping or pendulous; stems cylindric, ribbed, angled, winged or flat; small flowers. 4 spp., E Bolivia and NE Argentina extending to E Brazil (3, none endemics).

43. Rhipsalidopsis Britton & Rose. Two spp., both endemics to S Brazil.

44. Rhipsalis Gaertn. Epiphytic, rarely epilitic shrubs, often pendulous. Stems cylindric, ribbed, angled, winged or flat, usually segmented and spineless, with small white flowers and fleshy fruits. 45 spp. from South American, two up to Caribbean, Central America and Mexico, with R. baccifera (J.M.Muell.) Stearn in S Africa, Madagascar and Sri Lanka, 38 in Brazil (31 endemics; two of them, from Espírito Santo and Rio Grande do Sul states, are rare plants in Brazil, by Plantas Raras do Brasil’s book), with outliers endemics in Ecuador (1), Peru (2) and Bolivia (1); four extra-Brazilian many widely. Three subgenera.

§ subg. Calamorhipsalis ‣ 8 (or more) spp., all endemic to Brazil exc. R. floccosa Salm-Dyck ex Pfeiff. to Bolivia, Venezuela and Guianas.

§ subg. Erythrorhipsalis ‣ 9 (or more) spp., all endemic to Brazil exc. R. cereuscula Haw. up to Bolivia and Parguay.

§ subg. Rhipsalis ‣ 23 (or more) spp., tropical America to Africa, some absent in Brazil.

45. Schlumbergera Lem. Epiphytic or epilitic shrubs, stems segmented, in some spp. opuntioids; flowers scarcely to strongly zygomorphic, white, red, pink, or purplish; fruit baccate, globose to obconic, ribbed or terete, the perianth deciduous. 7 spp. found in the coastal mountains of SE & S Brazil, one of them, from Espírito Santo state, is a rare plants in Brazil, by Plantas Raras do Brasil’s book.

5.6. CACTOIDEAE ▸ TRIBE NOTOCACTEAE (5/124) - all genera in South America.

46. Eriosyce Philippi. (inc. Islaya) Stems usually unbranched, globose to short-cylindric, ribbed; ribs usually divided into prominent tubercles. 56 spp. from Chile and N Argentina, five up or endemic to dry coasts of SW Peru.

47. Neowerdermannia Backeberg. Low growing, with stout taproot; stem simple, globose to depressed. Two spp., Peru, S Bolivia, N Chile and N Argentina.

48. Parodia Spegazzini. Small, globose to cylindric, ribbed or tuberculate cactis, low growing, simple or clustering; flowers diurnal yellow to red or pink, short tube. 66 spp. from E South America, in S Brazil (24, 16 endemics, 6 are rare plants in Brazil, by Plantas Raras do Brasil’s book, one in Paraná and remaining in Rio Grande do Sul state), Uruguay, NE Argentina and S Paraguay, and E Andes from Bolivia and NW Argentina; one record of P. erinacea (Haw.) N.P. Taylor in Colombia, dubious.

49. Rimacactus Mottram. Only one sp., R. laui (Lüthy) Mottram, in a small barren area (less than 10 square km) in the costal desert of Tocopilla, where hardly any other plants at all are capable of surviving, in N Chile.

50. Yavia R.Kiesling & Piltz. Only one sp., Y. cryptocarpa R. Kiesling & Piltz, Bolivia and N Argentina.

5.7. CACTOIDEAE ▸ TRIBE CEREAE (39/545–575) – six lineages and a unplaced genus, all in South America.

51. Estevesia P.J.Braun. Only one sp., E. alex-bragae P.J.Braun & Esteves, endemic to Goiás state, Brazil.

∎ UEBELMANNIA CLADE (1/3) ‣ a single genus.

52. Uebelmannia Buining. Unbranched, stems usually small, globose to cylindric; flowers small, diurnal, yellow. Three spp., all rare plants in Brazil, by Plantas Raras do Brasil’s book, endemics to mountains of NE Minas Gerais state.

∎ AYLOSTERA CLADE (1/9) ‣ a single genus.

53. Aylostera Speg. Small globullar cacti, flowers showy; 9 spp., Bolivia to Venezuela.

∎ GYMNOCALYCIUM CLADE (1/61) ‣ a single genus.

54. Gymnocalycium Pfeiff. ex Mittler. Low growing, globular cacti, mostly unbranched, stems often strongly depressed, or globose to short cylindric, sometimes cushions. 61 spp., S Brazil (5, G. horstii Buining endemic), Paraguay, Bolivia (8), Uruguay and Argentina (41).

∎ SUBTRIBE REBUTIINAE (4/41) ‣ all genera in South America.

55. Browningia Cardenas. 11 spp., 9 in Peru (one up to Ecuador another up Cono Sur), one in C Andean Colombia, another in Bolivia and Paraguay.

56. Castellanosia Cárdenas. Only one sp., C. caineana Cárdenas. restricted from Bolivia and Paraguay.

57. Lasiocereus F. Ritter. Two spp., endemic to Peru.

58. Rebutia Pfeiff. ex Mittler. Low growing, stems simple or more often freely clustered, small, globose to shortly cylindric. 7 spp. from Bolivia and NW Argentina.

59. Weingartia Werderm. 20 spp., Bolivia, one up to Peru.

∎ SUBTRIBE CEREINAE (13/180 excluding Pierrebraunia) ‣ all genera in South America.

60. Arrojadoa Britton & Rose. Low shrubs, few branched; stems cylindric or slender-cylindric, sometimes segmented; one sp. with true stem-tuber. 7 spp., NE Brazil, six are rare plants in Brazil, by Plantas Raras do Brasil’s book, all in Minas Gerais or Bahia states; at maturity, stems produce terminal cephalia blanketed in white wool and brown to reddish bristles.

61. Brasilicereus Backeb. Shrubs, stems cylindric, rather woody. Three spp. from Minas Gerais and Bahia states in E Brazil, B. markgrafii Backeb. & Voll a rare plant in Brazil, by Plantas Raras do Brasil’s book, from Minas Gerais state.

62. Cereus Mill. Tree-like or shrub, usually much branched; stems erect or ascending; sweet potato cactus; naked large nocturnal flowers and fruits dehiscent by longitudinal slits. 31 spp., mostly S. American (30, only three up to Caribbean) from Colombia, Venezuela and Guianas to Argentina and E Brazil; 13 in Brazil, 10 endemics, one, from Fernando de Noronha Is., is a rare plant in Brazil, by Plantas Raras do Brasil’s book.

C. jamacaru DC. is the one of the largest tree-cacti at up to 18 m (59 ft) in height, 10 m (33 ft) crown spread and up to 102 cm (40 in) trunk thickness; It can bear spines up to 19 cm long; by reason of its succulence, these may be the most massive (heaviest) of all tree flowers.

63. Cipocereus F.Ritter. Shrubs, stems cylindric, rather woody; spines absent to numerous; fruits in some species intensely blue-waxy. 5 spp., endemic to Minas Gerais state, E Brazil, all are rare plants in Brazil, by Plantas Raras do Brasil’s book.

64. Coleocephalocereus Backeb. Branching from the base or unbranched; stems elongate-globose to cylindrical or columnar; flowers in cephalium. 6 spp., E & SE Brazil, mainly in open areas, one of them a rare plant in Brazil, by Plantas Raras do Brasil’s book, in Minas Gerais state.

65. Discocactus Pfeiff. Low growing, stem mostly simple, depressed-globose to globose; cephalium terminal, depressed; flowers small, white, and nocturnal. 12 spp., 10 in Brazil (one up to Paraguay) and two in Bolivia; two endemic species from Minas Gerais state are rare plants in Brazil, by Plantas Raras do Brasil’s book.

66. Espostoopsis Buxb. Shrubby, mainly branching near the base; stems cylindric; stems plenity cobered of white hairs. Only one sp., E. dybowskii (Rol.-Goss.) Buxb., endemic to Bahia state in E Brazil.

67. Facheiroa Britton & Rose. Shrubs or tree-like; stems cylindric; nocturnal flowers inside a lateral cephalium. Three spp. endemic to NE Brazil, Bahia, Piauí and Pernambuco states, with one is a rare plant in Brazil, by Plantas Raras do Brasil’s book, endemic to Bahia.

68. Leocereus Britton & Rose. Few-branched; rootstock woody; stems slender, erect, ribbed; flowers tubular, nocturnal. Only one variable sp., L. bahiensis Britton & Rose, endemic to E Brazil.

69. Melocactus Link & Otto. Unbranched (unless damaged); stem depressed to globosse-columnar, rarely more than 1 meter; with cephalium and small diurnal red, pink or magenta flowers, fruits white to deep red. 37 spp. (43 in Korotkova, 2021), 21 endemics to E Brazil (7 are rare plant in Brazil, by Plantas Raras do Brasil’s book, six in Bahia and one in Pernambuco state), two endemics to Roraima state in N Brazil (one of them a rare plant in Brazil, by Plantas Raras do Brasil’s book), three in Colombia to Venezuela, two in Ecuador to Peru, six in Caribbean, one up to N Colombia, two in northern South America from Colombia to Guianas and N Brazil (both), and M. curvispinus Pfeiff. from Mexico to Colombia, Venezuela and Caribbean; rocky fields, semideciduous and deciduous forest, vegetation on sandy soil and coastal dunes and vegetation on rocky outcrops.

70. Micranthocereus Backeb. Shrubby to tall columnar, unbranched or branched, from the base; stems cylindrical; flowers from the cephalium. 10 spp., endemics to Tocantins, Bahia, Minas Gerais and Goiás states in C Brazil, 5 of them (from Bahia and Minas Gerais) are rare plants in Brazil, by Plantas Raras do Brasil’s book.

71. Pilosocereus Byles & G.D.Rowley. Tree-like or shrubby, more or less branched from the base or thunk, globose to cylindric, to 10 m; fruit fig-like, flowers diurnal yellow to red or pink, short tube. 42 spp. and four heterotypic subspecies (51 in Korotkova et al., 2021), 20 endemics to E Brazil (six are rare plants in Brazil, by Plantas Raras do Brasil’s book, all in Minas Gerais or Bahia states), P. oligolepis (Vaupel) Byles & G.D. Rowley occurs in Roraima state in Brazil and Guyana, P. machrisii (E.Y. Dawson) Backeb. in Brazil and Cono Sur, two endemics to Venezuela, one in Caribbean coast of Colombia and Venezuela to Caribbean, one from Colombia to Peru, and 16 from Mexico, Central America and Caribbean.

72. Praecereus Buxb. Cereus-like. Two spp., P. euchlorus (F.A.C. Weber ex K. Schum.) N.P. Taylor from Venezuela and Colombia up to Cono Sur and Brazil, and P. saxicola (Morong) N.P. Taylor endemic to Brazil.

73. Stephanocereus A.Berger. (syn. Lagenocereus) Tree-like, sparsely branched to unbranched; stems cylindric to ovoid; flowers in ring-like cephalia. Two spp. from Bahia and Minas Gerais states in Brazil.

74. Stetsonia Britton & Rose. Tree-like, 5-10 m; stems cylindric, ribbed, spiny; flowers large, nocturnal. Only one sp., S. coryne (Salm-Dyck) Britton & Rose, from Paraguay, S Bolivia, NW Argentina, and Mato Grosso do Sul state in C Brazil.

75. Xiquexique Lavor, Calvente & Versieux. Tree-like to shrubby cacti, main stem upright, mature branches arched, running more or less parallel to the ground, apices ascending, new axes arising subapically (branching candelabriform); ribs 4-15. Three spp. from NE Brazil, along the entire NE region and in the central-north portion of Minas Gerais state.

∎ SUBTRIBE TRICHOCEREINAE ‣ a monophyletic Echinopsis would need to include Acanthocalycium Backeb., Arthrocereus A.Berger, Cephalocleistocactus F.Ritter, Cleistocactus Lem. (including Borzicactus Riccob.), Denmoza Britton & Rose, Espostoa Britton & Rose (including Vatricania Backeb.), Haageocereus Backeb., Harrisia Britton, Matucana Britton & Rose, Mila Britton & Rose, Oreocereus (A.Berger) Riccob., Oroya Britton & Rose, Pygmaeocereus H.Johnson & Backeb., Rauhocereus Backeb., Samaipaticereus Cárdenas, Weberbauerocereus Backeb., and Yungasocereus F.Ritter, all of which are part of a well-supported clade (100% bootstrap support) interspersed with species of Echinopsis. Species previously assigned to Lobivia, i.e., species with diurnal flowers, Trichocereus, i.e., columnar species, or Echinopsis s.str., i.e., globular species with funnel-shaped flowers of nocturnal anthesis, do not form clades.

Adopted here with 4 clades and a isolate genus; all genera in South America.

ISOLATE GENUS

76. Arthrocereus A.Berger. Shrubs; stems cylindric; showy nocturnal white or pinkish flowers, mainly open areas in rocky places. 5 spp., 4 from Minas Gerais and São Paulo, one from Mato Grosso states, all in Brazil, two of them are rare plants in Brazil, by Plantas Raras do Brasil’s book, both in Minas Gerais state.

CLADE I

77. Cleistocactus Lemaire. Shrubby or rarely tree like, variously habit, color flower and corolla format. 20 spp. from Colombia to N Argentina, Bolivia, Paraguay and Uruguay, only C. baumannii (Lem.) Lem. in Brazil, reported from Mato Grosso and Mato Grosso do Sul states in W Center Brazil.

78. Samaipaticereus Cárdenas. Only one sp., S. corroanus Cárdenas, from Bolivia to Peru.

79. Vatricania Backeb. Only one sp., V. guentheri (Kupper) Backeb., endemic to Bolivia.

80. Weberbauerocereus Backeb. 9 spp., 8 in Peru one endemic to Bolivia.

81. Yungasocereus F.Ritter. Only one spp., Y. inquisivensis (Cárdenas) F.Ritter ex D.R.Hunt, endemic to Bolivia.

CLADE II

82. Borzicactus Riccob. Erect, crawling or decumbent plants, with more or less sturdy, cylindrical, long stems, ribs rounded, more or less thick around the areoles, sometimes with chins; spines relatively short and often numerous, aciculate. 19 spp., Ecuador to Peru.

83. Espostoa Britton & Rose. 11 spp., 10 in Peru (8 endemics, two up to Ecuador) and one endemic to Bolivia.

84. Haageocereus Backeb. 13 spp., 12 in Peru (10 endemics and one up to N Chile) and one endemic to N Chile.

85. Matucana Britton & Rose. 17 spp., endemic to Peru, mainly along Marañon Valley

86. Mila Britton & Rose. Only one sp., central Peru.

87. Oreocereus (A.Berger) Riccob. 11 spp., Peru, Bolivia, Chile, Argentina.

88. Oroya Britton & Rose. Two spp., Peru.

89. Pygmaeocereus Britton & Rose. Two spp., Peru.

90. Rauhocereus Backeb. Only one sp., R. riosaniensis Backeb., endemic to Peru.

91. Reicheocactus Backeb. Two spp. (sometimes only R. famatinensis (Speg.) Schlumpb.), endemics to NW Argentina.

CLADE III

92. Acanthocalycium Backeb. 5 spp., four endemics to Argentina and A. rhodotrichum (K.Schum.) Schlumpb. from E. Bolivia to W. Central Brazil (Mato Grosso do Sul state) and Uruguay.

93. Denmoza Britton & Rose. Only one sp., D. rhodacantha (Salm-Dyck) Britton & Rose; W and NW Argentina.

94. Lobivia Britton & Rose. 33 spp., Peru to Argentina, Chile and Bolivia.

95. Setiechinopsis Bolivia Only one sp., S. mirabilis (Speg.) Backeb. ex de Haas, from E Argentina (Santiago del Estero, Santa Fe, San Juan, San Luis, Mendoza, La Rioja, Mendoza, Colonia Ceres), grows on the edges of salinas and in monte shrublands, often under shrubs in flat prairies up to the base of mountains.

96. Trichocereus Ricoob. 6 spp., Ecuador to Argentina, Chile and Bolivia.

CLADE IV

97. Chamaecereus Lodè & F. Carlier. 4 spp., Bolivia and Argentina.

98. Echinopsis Zucc. Tree-like to globose, large flowers white to deep red, externally with areoles and hairs, stems usually disticly ribbed, very spine to spineless. 15 spp. in South America, from Bolivia to Argentina and S & W Brazil (two, none endemics), one up to Ecuador.

The only cactus genus that is more confusing than Echinopsis is that of Opuntia.

99. Harrisia Britton. Shrubs, some tree like, to 7 m tall, or scandent; roots sometimes tuberous; flowers large, white, nocturnal. 19 spp. in two subgenera:

§ subg. Harrisia ‣ two sections.

§ sect. Adscendens ‣ only H. adscendens (Gurke) Britton & Rose, in dry seasonal scrubland of NE Brazil (caatinga).

§ sect. Harrisia ‣ 11 spp., Greater Antilles, Bahamas, and Florida, U. S. A.

§ subg. Eriocereus ‣ two sections.

§ sect. Eriocereus ‣ 6 spp.; Gran Chaco region of Argentina, Bolivia, S Brazil (2, none endemics), and Paraguay.

§ sect. Roseocereus ‣ only H. tetracantha (Labour.) D.R. Hunt in the E inter-Andean dry valleys of Bolivia at elevations of 1,200–2,600 m.

100. Leucostele Backeb. 7 spp. from Bolivia, Chile and NW Argentina.

101. Soehrensia Backeb. 21 spp., 20 from Bolivia and Argentina, one endemic to Paraguay.

48. CORNALES

FAMILIES ABSENTS IN SOUTH AMERICA: CURTISIACEAE (1/1), GRUBBIACEAE (1/3), HYDROSTACHYACEAE (1/22) AND NYSSACEAE (5/37).

CORNACEAE

§ FULLY CHLOROTROPHIC

Genera/species 2/108 Distribution temperate regions in the Northern Hemisphere, tropical Central and E Africa, Madagascar, Mascarene Islands, subtropical and tropical Asia, Malesia, New Guinea and nearby islands, E Australia, Melanesia. Habit usually bisexual (in Alangium grisolleoides Capuron and some species of Cornus dioecious), evergreen or deciduous trees or shrubs (Cornus suecica L. and C. canadensis L. are stoloniferous perennial herbs or suffrutices; some species of Alangium are lianas). Use ornamental plants, fruits (Cornus mas L. etc.), timber.

Some Viburnum species look similar to Cornaceae, but the latter often have 4-merous flowers, they lack stellate indumentum, and their lateral veins ascend towards the apex of the blade. Only one genus in South America.

SYSTEMATIC outsiders are Alangium (27; tropical Africa, NE Madagascar, China to E Queensland, NE New South Wales and New Caledonia), Nyssa (8; tropical Asia to China, E and SE U.S.A., S Mexico), Camptotheca (1; S and SE China), Davidia (1; SW China), Diplopanax (2; China, Vietnam), Mastixia (c 20; S India, Sri Lanka, E Himalayas to S China, SE Asia, Malesia to New Guinea and Solomon Islands).

1. Cornus L. Trees or shrubs; leaves opposite or alternate, simple, margins entire or denticulate, primary venation pinnate, arching; stipules very rarely present; inflorescences terminal occasionally axillary, cymose, involucres petal -like; flowers bisexual (rarely unisexual and dioecious), small, actinomorphic; sepals 4(-10); petals 4(-10), free, valvate; fruit a grooved, globose or ovoid drupe; seeds 2, flat. 60 spp., northern temperate up to Central America, only one sp. in South America, C. peruviana J.F.Macbr., found predominately at high elevations at 1,400-3,000 m above sea level, below the forest canopy in mountainous regions of Central America, Bolivia, Colombia, Ecuador, Mexico and Peru; also cultivated in Brazil as an ornamental and for its wood throughout most of the Neotropical countries.

HYDRANGEACEAE

§ FULLY CHLOROTROPHIC

Genera/spp. 9/215–225 Distribution temperate and subtropical North America, southern Mexico, Central America, W South America to central Chile (Andes), Caucasus, E Asia to Japan and Russian Far East, and a few species in SE Asia eastwards to New Guinea, with their highest diversity in China. Habit usually bisexual (in Broussaisia polygamodioecious), evergreen or deciduous trees or shrubs, often climbing and twining (rarely suffrutices or perennial herbs). Bark usually exfoliating. Use ornamental plants, medicinal plants.

The climbing habit is relatively rare in the family Saxifragaceae, the majority of the species being shrubs; all the climbing species, which are found only in Hydrangea L., utilize the same method of climbing, i.e., attachment to a supporting substrate by means of fine rootlets arising directly from the upright stems (a method of climbing similar to that of poison ivy, Rhus toxicodendron L.); one genus in Neotropics.

The opposite leaves and inferior ovary might cause confusion with families included in the Gentianales (Rubiaceae and others), Caprifoliaceae, Lamiales. The free petals and stamens twice as many as petals found in Hydrangeaceae are good characters to separate them.

SYSTEMATIC subfamily Jamesioideae (2/5, SW U.S.A., northern Mexico) and Philadelpheae (6/c 125, temperate and subtropical regions on the Northern Hemisphere, with their highest diversity in SE Asia to Philippines) do not occur in South America; in South America only the monogeneric Hydrangeoideae.

1. Hydrangea L. Evergreen root climbers growing up to 30 – 40 m high in the canopy of mostly primary forests, as tipically tropical lianas, with coriaceous leaves and hortensia-like whitish, greenish, yellowish or purplish inflorescences, in large terminal corymbs often with sterile outer flower. 208 spp., incl. Hydrangea s.str. in America and Himalayas to Japan and Philippines, Cardiandra in E Asia, Deinanthe in central China and Japan, Platycrater in Japan, Dichroa in China, SE Asia and Malesia to New Guinea, Broussaisia on Hawaii, Schizophragma in Himalayas to Korean Peninsula, Japan and Taiwan (China), Decumaria in China and U.S.A., and Pileostegia in E Asia.

Only one section in Neotropics: Hydrangea sect. Cornidia, with 26 spp., 12 only in Mexico and Central America, three spp. from Central America to Venezuela and Bolivia; three endemics to Colombia; three from Colombia and Ecuador to Peru; one endemic to Ecuador; two widely in Central Andean in Ecuador, Peru and Bolivia, H. mathewsii Briq. endemic to Peru, and H. serratifolia (Hook. & Arn.) F. Phil. from Chile and Argentina (a shrubs or robust climbers, the largest liana of southern South America); and the highly disjunct H. integrifolia Hayata from Philipinnes and Formosa. South America includes exact 14 spp.

LOASACEAE

§ FULLY CHLOROTROPHIC

Genera/species 21/c. 346 Distribution temperate, subtropical and tropical regions of North, Central and South America from SW Canada to Argentina and Chile, Caribbean, the Galápagos Islands, Marquesas Islands; one species in SW Africa; one species in Somalia, NE Ethiopia, and the SW Arabic Peninsula. Habit usually annual, biennial or perennial herbs, sometimes winding, rarely subshrubs or a succulent tree with often exfoliating bark, very rarely woody lianas or cushion-forming plants, thickend storage roots sometimes present.

Only the genera Kissenia R.Br. ex T.Anderson (one species in SW Africa; one species in Somalia, NE Ethiopia, and the SW Arabic Peninsula) and Plakothira J.Florence (Marquesas Islands) are extra-American. All species are native, and many species are very narrowly endemic. Mentzelia aspera Vell. is likely to have originated in N Central America and has probably been spread as a weed; 5 genera and 15 spp. in Brazil, 10 of then endemic; Peru is the most important centre of diversity for the family, with at least 80 spp. in 7 genera, two endemic; Mexico and Chile are secondary centres of diversity. Only Blumembachia, Caiophora, Loasa and Mentizelia in Argentina. The family is a potential source of highly unsaturated oils from the seeds. Several species are used in traditional medicine, but only one is found in international trade and only to a very limited extent (Mentzelia scabra subsp. chilensis (Gay) Weigend). Many species, especially in Aosa, Nasa, Caiophora and Loasa, have extremely painful stinging hairs. A large number of species, especially of the genus Nasa, are very narrowly endemic, some only known from the type collection, and at least two already extinct (N. hastata (Killip) Weigend from C Peru and N. humboldtiana subsp. humboldtiana (Urb. & Gilg) Weigend from S Ecuador).

Members of the Loasaceae are sometimes confused with Cucurbitaceae or Malvaceae:

ü Flowers hermaphrodite (vs. usually unisexual in Cucurbitaceae).

ü Leaves estipulate (stipulate in Malvaceae).

ü Pubescence of glochidiate/scabrid trichomes (stellate trichomes in Malvaceae)

SYSTEMATIC genera Eucnide (1/13, SW U.S.A., Mexico) and Schismocarpus (1/1, Oaxaca Province in Mexico), sucessively basal in Loasaceae, do not occur in South America.

1. SUBFAMILY MENTZELIOIDEAE (1/c 80) ‣ a single genus.

1. Mentzelia L. Annual or perennial herbs, shrubs to small trees. 101 spp. from Argentina to Canada, Caribbean, Galapagos Is., most SW U.S.A. and Mexico; 10 in South America, only the widely distributed M. aspera L. in Brazil.

2. SUBFAMILY GRONOVIOIDEAE (4/9) ‣ outsiders are Petalonyx (5; SW U.S.A., Mexico), Cevallia (1; SW U.S.A., Mexico), Fuertesia (1; Hispaniola).

2. Gronovia L. Scandent, annual herbs, stinging hairs and characteristical glochidiate hairs 0.5 mm long with two hooks at their tip always present; leaves opposite below, alternate above; flowers in terminal thyrsoids, flowers 5-merous, subsessile, calyx yellow or yellowish green; petals pale yellow; fruit an ovoidal cypsela. Two spp., G. longiflora Rose is endemic to S. Mexico and G. scandens L. is widely distributed from Mexico to Venezuela and Peru, fast-growing weeds in raingreen forests.

3. SUBFAMILY LOASOIDEAE (15–17/c 225) ‣ outsiders are Kissenia (2; Ethiopia, Somalia, SW Arabian Peninsula; Namibia, Northern Cape), Plakothira (3; Marquesas Islands).

3. Aosa Weigend. (inc. Chichicaste) Annual or perennial herbs or shrubs, often sparsely branched. 7 spp., A. plumieri (Urb.) Weigend in south Hispaniola, A. grandis (Standl.) R.H.Acuña & Weigend in Costa Rica/Choco region, and five species of Brazil, four in dry areas in NE region, and one in Atlantic Forest in Rio de Janeiro state.

A. grandis is among the largest species of Loasaceae, both in terms of absolute plant size and in the size of its organs; in the wild, plants have stems up to ca. 4 m long and 7 cm in diameter (the largest stems sometimes held almost horizontal for most of their length, with adventitious roots often developing from the nodes); the species inhabits the understory of evergreen wet forests, close to forested streams or more rarely, roadsides; the leaves are lobed as in Aosa and other Loasoideae, usually very large, reaching lengths up to ca. 50 cm, as are the bracteate inflorescences that can grow to more than 1 m long.

4. Blumenbachia Schrad. Scandent or ascending, annual or perennial herbs. 11 spp. from Brazil, Uruguay, Paraguay, Argentina, Chile; the six Brazilian species are restricted of southern region, mainly in Santa Catarina state, and B. amana T. Henning & Weigend from Minas Gerais state; sect. Gripidea has five spp., four species range from central and N Argentina to S Brazil with two wideranging species, B. insignis Schrad. and B. latifolia Cambess., and two narrow endemics, B. catarinensis Urban & Gilg in Santa Catarina, Brazil, and B. hieronymi Urban in Córdoba, Argentina.

5. Caiophora C.Presl. Rosulate, erect or scadent perennial herbs. 50 spp. from Peru to Argentina, Chile, C. arechavaletae (Urb.) Urb. & Gilg up to Uruguay and S Brazil (Rio Grande do Sul state, unique outside Andes) and C. contorta (Desr.) C. Presl up to Ecuador.

6. Grausa Weigend & R.H. Acuña. (off Loasa) Perennial, rosulate or weakly rhizomatous, or climbing herbs with diffuse root-system; leaves congregated into dense, basal leaf rosette, petiolate, sagittate, pinnatisect or palmatisect to ternately compound, very rarely pinnate, petiole distinct. 6 spp. from Argentina and Chile, from lowlands (twining species) or high Andes (acaulescent species).

7. Huidobria Gay. Erect, densely branched annual herbs or shrubs. Two spp. endemics to deserts of N Chile; diphyletic genus.

8. Klaprothia Kunth. Erect or ascendant annual or perennial herbs. Two spp., K. fasciculata (C. Presl) Poston and K. mentzelioides Bonpl. & Kunth, both widely distributed and weedy spp. from S Mexico to Brazil (both species) and Bolivia, also Caribbean, Galapagos.

9. Loasa Adans. (exc. Grausa, Pinnasa) Rosulate, ascendant, or erect annual or perennial herbs, often cushions; petals white, red or yellow. 19 spp. in Chile (mostly) and adjacent Argentina.

10. Nasa Weigend. Annual or perennial herbs, subshrubs and shrubs, 5 to 400 cm tall; stinging hairs always present, primary root short-lived, root system dominated by adventicious roots from the decumbend basal stem portion; leaves opposite or alternate, flowers in terminal thyrsoids, dichasia or monochasia; flowers pentamerous, corolla reflexed, spreading or narrowly campanulate, petals white, yellow or orange; fruits are capsules. 90 spp., N. triphylla (Juss.) Weigend from Mexico to Bolivia, two only in Central America, and remaining 87 confined to South America; 20 spp. in Colombia, 31 in Ecuador, over 50 in Peru, 2 in Bolivia, Chile, Venezuela, essentially a Andean group from the cloud forests, subparamo and the puna with only very few species at lower elevation or outside the Andes (e.g. Cordillera de Colonche, Ecuador); the vast majority of taxa is narrowly endemic to single mountains or narrow regions; morphologically the genus is enormously variable including evergreen shrubs with hummingbird-pollinated, large, orange flowers and ephemeral plants which are self-pollinated.

11. Pinnasa Weigend & R.H. Acuña. (off Loasa) Perennial, rosulate herbs with thick tap-roots, sometimes stoloniferous; leaves congregated into dense, basal leaf rosette, petiolate, deeply pinnatifid to bipinnate, petiole distinct, much shorter than lamina/rachis; inflorescences usually axillary, arising from the axils of the rosette leaves, one- to many-flowered, often with a winding axis, frondose with opposite, pinnate or pinnatifid bracts. 11 spp., Chile and Argentina (4).

12. Presliophytum (Urb. & Gilg) Weigend. Erect densely branched shrubs and tiny small herbs. 5 spp., 3 in desert of W Peru and two in N Chile, in rocky slopes.

13. Scyphanthus D.Don. Scandent annual herbs. Only one sp., S. elegans Sweet, Mediterranean scrub lands of central Chile.

14. Xylopodia Weigend. Shrub with erect branches from horizontal xylopodium. Two spp., X. klapothioides Weigend, very narrowly endemic is so far only know from in rocky slopes in Cajamarca, Peru, and X. laurensis C.M. Martín & C.A. Zanotti eastern slope of the Andes from Bolivia (Dptos. La Paz and Chuquisaca) to northern Argentina (Prov. Jujuy).

49. ERICALES

FAMILIES ABSENTS IN SOUTH AMERICA: BALSAMINACEAE (2/1,108), DIAPENSIACEAE (6/19), FOUQUIERIACEAE (1/11), RORIDULACEAE (1/2), SLADENIACEAE (2/3).

LINEAGE 1 of 6: MARCGRAVIIDS

MARCGRAVIACEAE

§ FULLY CHLOROTROPHIC

Genera/species 7/139 Distribution southern Mexico, Central America, Caribbean, northern and central South America; Habit bisexual evergreen shrubs, often epiphytic or semi-epiphytic, or lianas (rarely small trees).

Marcgraviastrum, Souroubea, Norantea and Sarcopera are missing in the Antilles (although Norantea guianensis Aubl. is rarely cultivated as an ornamental). Native and endemic to the Neotropics, Norantea guianensis sometimes cultivated in countries outside its range (e.g. Jamaica, Costa Rica, Trinidad). The elaborate inflorescences with the variously shaped bracteal ‘nectary-containers’ are frequented by a wide range of visitors (insects, lizards, birds, bats, non-flying mammals). Different pollination syndromes are exemplified in the various taxa. Whereas Ruyschia and Souroubea seem predominantly insect-pollinated, Norantea, Sarcopera and Schwartzia brasiliensis (Choisy) Bedell ex Gir.-Cañas are probably best adapted to bird-pollination, especially by perching birds. Most Schwartzia, Marcgraviastrum and Marcgravia spp. seem to be primarily bat-pollinated, but the ornithophilous syndrome seems to occur as well. Apart from occasional horticultural use in the tropics the family has no significant economic value.

Marcgravia is sister to the remaining Marcgraviaceae. The leaves in Marcgravia are distichous. The inflorescence is umbel-like, racemose, with sterile flowers. The flowers have four sepals, four petals connate into a calyptra, and nectaries adnate to the sterile aborted flowers. The remaining Marcgraviaceae have spiral leaves, inflorescence with exclusively fertile flowers, three or five sepals, five entirely or partially free petals, and free (not adnate) nectaries. Marcgraviastrum has an umbel-like racemose inflorescence. ’Sarcopera’ (non-monophyletic) has a spicate inflorescence.

Vegetatively similar to the Ternstroemiaceae (non-climbing!) this family was formerly considered to be close to Theaceae/Ternstroemiaceae. Recent molecular data associate it rather with Balsaminaceae, and the former Theaceae-satellites Pellicieraceae and Tetrameristaceae.

SYSTEMATIC all genera in South America.

1. SUBFAMILY MARCGRAVIOIDEAE (1/67) ‣ a single genus.

1. Marcgravia L. Climbing shrubs or vines with dimorphic branches: juveniles are creeping, adepressed to the substrate; fertile pendulous; leaves distichous; conspicuous heterophylly between juvenile, plagiotropous and adult orthotropous branches; perianth tetramerous, petals fully fused to form a caducous cap; only central (sterile) flowers of inflorescence fused with nectary bract, other flowers lacking nectary. 67 spp., S Mexico, Mesoamerica, South America (50), Antilles; 18 ssp. in Brazil, 8 endemics.

2. SUBFAMILY NORANTEOIDEAE (6/71) ‣ all genera occur in South America.

2. Marcgraviastrum (Wittm. ex. Szyszyl) de Roon & S. Dressler. Sprawling shrubs and lianas, often epiphytic. 15 spp., all in South America, 13 in northern South America up to S Nicaragua and up N Brazil (M. mixtum (Triana & Planch.) Bedell), plus two endemics to SE Brazil.

3. Norantea Aubl. Lianas or sprawling shrubs, often epiphytics; inflorescence long, dense racemes, red showy. Two spp., N. guianensis Aubl. in N South America to S Brazil and Bolivia, and N. goyasensis Cambess. in Brazil and Bolivia.

4. Ruyschia Jacq. Climbing shrubs or lianas; Inflorescence dense multiflorous racemes. 8 spp. confined high mounatins, 7 from Mexico up to South America (5, 3 in N Andes), and one in Lesser Antilles.

5. Sarcopera Bedell. Sprawling shrubs and lianas, occasionally small trees, often epiphytic. 8 spp. in N South America, two up to Central America (Honduras southwards), mainly in N Andes and Guiana Shield, 3 up to N Brazil (none endemics, only in Amazonas and Roraima states).

6. Schwartzia Vell. Sprawling shrubs, occasionally small trees; inflorescence short racemes. 20 spp., 16 from Costa Rica through the Andes south to Bolivia, in the Caribbean Basin, and 4 spp. in E Brazil, all endemics. S. brasiliensis (Choisy) Bedell ex Gir.-Cañas. (Marcgraviaceae) endemic to E Brazil and Puya alpestris (Poepp.) Gay (Bromeliaceae) endemic to Chile are the unique blue-nectar plants known.

7. Souroubea Aubl. Climbing shrubs or lianas, often epiphytic; inflorescence lax or dense racemes. 19 spp., Mexico to Bolivia (absent from the Antilles), 15 in South America, 5 in Brazil, one endemic.

TETRAMERISTACEAE

§ FULLY CHLOROTROPHIC

Genera/species 3/4 Distribution Neotropics and SE Asia. Two neotropical trees genera restricted Costa Rica to Brazil, and one sp. (the type) in SE Asia. Tetramerista (1-3) is endemic to SE Asia. Habit bisexual, evergreen trees or shrubs (Pelliciera consists of mangrove trees with buttresses). Aerial roots frequent along stem base in Pelliciera.

SYSTEMATIC two subtribes, both in South America.

1. TRIBE PELLICIEREAE (1/1) ‣ a single genus.

1. Pelliciera Planch. & Triana. Mangrove trees to 15m tall, 15-20cm in diameter (above buttresses); trunks with enlarged fluted boles, buttresses 1.5-2m tall (average high tide level), 1.5m in diameter, formed by the emergence of short-born roots; leaves spirally arranged, clustered at apex of branches; flowers solitary, in leaf axils or terminal, sessile; sepals 5, free; petals 5, free; stamens 5; fruit indehiscent, woody, covered with resinous pustules; seed 1. Two spp., P. benthamii (Planch. & Triana) Cornejo endemic to Panamá, and P. rhizophorae Planch. & Triana from the Gulf of Nicoya, Costa Rica to the Esmeraldas river, Ecuador and few scattered populations on the Caribbean coasts of Nicaragua, Panamá and Colombia; these species has the second largest seed in dicots (10 cm diameter) and sixth of all flowering plants.

2. TRIBE TETRAMERISTEAE (2/2) ‣ outsider Tetramerista (1; the Malay Peninsula, Sumatra, Borneo).

2. Pentamerista Maguire. Tree 3-6m tall, tortuous branches; leaves simple, axillary racemes and greenish-yellowish flowers; sepals 5, free; petals 5, free; stamens 5, free from perianth, filaments flattened, base connate, alternate with petals, anthers basifixed, dehiscing via longitudinal slits; ovary superior, carpels and locules 5, syncarpous, placentation axile, ovule 1 in each locule; style 1, undivided; stigma simple to minutely lobed; fruits berries, endocarp firm; seeds 4-5, relatively large. Only one sp., P. neotropica Maguire, in a restricted area of savannas in N South America in border of Venezuela and Colombia, and one small scarcely population in Madeira – Purus rivers interfluvial region of Amazonas state of Brazil, 1,000 km distance of both, diiscovery (in Brazil) only in 2007.

LINEAGE 2 of 6: LECYTHIDS

POLEMONIACEAE

§ FULLY CHLOROTROPHIC

Genera/species 27/c. 385 Distribution North America including polar areas, Mexico, Central America, northern and W South America southwards to southern Chile, temperate and arctic Eurasia, Himalaya, with their largest diversity in W North America. Habit bisexual, usually perennial, biennial or annual herbs (sometimes shrubs or suffrutices; rarely lianas; in Cantua trees). Often evil-smelling.

SYSTEMATIC subfamily Acanthogilioideae (1/1, Baja California in NW Mexico) do not occur in South America. South America includes 37 spp. in this families.

1. SUBFAMILY COBAEIOIDEAE (3/32) ‣ outsider Bonplandia (1; Mexico, Guatemala). 22 spp. in South America, 21 in Peru, 12 endemics.

1. Cantua Juss. ex. Lam. (inc. Huthia) Shrubs or small trees, often with dimorphic leaves. 16 spp., almost a half South American Polemoniaceae, all endemics to Peru except 5, two into Ecuador and three into Bolivia.

2. Cobaea Cavanilles. Vines with alternate, pinnatelly compound leaves, stems woody or herbaceous, to 25 m long; inflorescence cymose, 1-5 flowered; corolla campanulate; corolla green, yellow, white, or purple, sometimes greenish at the beginning of anthesis and turning pink or purple with age, often suffused with pink, purple, or red, rarely striped, actinomorphic, glabrous or variously pubescent externally, villous inter nally near stamen insertion. 18 spp., 4 sections:

§ sect. Cobaea ‣ 6 spp., 5 in Mexico and C. trianae Hemsley in Colombia, Ecuador, and Venezuela; along streams or canyon walls in Andean cloud forests.

§ sect. Pachysepalae ‣ only one sp., restricted to southern Mexico and Guatemala

§ sect. Rosenbergia ‣ 9 spp., six in South America: three in NW Peru and Ecuador; two disjuncts in Mexico to Honduras and Peru; and C. penduliflora (Karsten) Hook. f. in Ecuador, Colombia, Peru (including Loreto central region, inlowland amazonic forest, near Brazilian border) and disjunct in N Venezuela.

§ sect. Triovulatae ‣ only one sp., C Mexico, Costa Rica, and Panama.

2. SUBFAMILY POLEMONIOIDEAE (19/c 350) ‣ 4 subtribes (all in South America); 14 spp. in South America in 10 genera; two spp., Microsteris gracilis (Hook.) Greene and Polemonium micranthum Benth., are conspecific with their high disjuncts North America counterparts, while the rest are unique to South America.

2.1 POLEMONIOIDEAE ▸ TRIBE POLEMONIEAE (1/c 28) - only one genus in this tribe.

3. Polemonium L. 24 spp., P. caeruleum L. in Eurasia and 23 remaning from North America, P. micranthum Benth. disjunct also in Chile and Argentina.

2.2 POLEMONIOIDEAE ▸ TRIBE PHLOCIDEAE (3/c 125) ‣ outsiders Linanthus (c 55; W North America, Mexico), Gymnosteris (2; W U.S.A.), Phlox (c 70; NE Asia, North America).

4. Leptosiphon Benth. Annual, herbaceous, or suffrutescent perennials, or shrubs. 30 spp., California and Baja California, with L. pusillus (Benth) Greene) endemic Chile (Coquimbo, Valparaiso, O'Higgins, Reg. Metropolitana).

5. Microsteris Greene. Annual, herbaceous, or suffrutescent perennials. Only one sp., M. gracilis (Hook.) Greene, in Canada to Mexico disjunct in Argentina, Peru, Chile and Bolivia.

2.3 POLEMONIOIDEAE ▸ TRIBE GILIEAE (c 6/c 95) ‣ outsiders Saltugilia (3; California); Lathrocasis (1; W North America), Allophyllum (6; W U.S.A.).

6. Collomia Nutt. 15 spp., 14 from shrublands and woodlands of W North America, and C. biflora (Ruiz & Pav.) Brand in South America, as common annual foothills species along the Andean corridor in Argentina and Chile.

7. Gilia Ruiz & Pavon. 37 spp., W North America but four in South America: G. lomensis V. F. Grant. are endemic to coastal regions Peru, G. valdiviensis Griseb. is endemic to coastal Chile, G. laciniata Ruiz & Pav. are from S Ecuador, Peru, Bolivia, Chile, Argentina and Uruguay, near sea level to high-alpine habitats, and G. crassifolia Benth. in Cono Sur.

8. Navarretia Ruiz & Pavon. 43 spp., 42 in North America (37 endemics to U.S.A.) and Mexico, and N. involucrata Ruiz & Pav. in South America, inhabiting vernal pools and seasonally moist depressions along the Andean corridor in Chile and Argentina.

2.4 POLEMONIOIDEAE ▸ TRIBE LOESELIEAE (9/c 95) ‣ ousiders Aliciella (21; W North America), Bryantiella (2; S California), Microgilia (1; W U.S.A.), Eriastrum (14; SW U.S.A.), Langloisia (3; SW U.S.A., N Mexico).

9. Dayia J.M.Porter. (inc. Bryantiella p.p.) 5 spp., 4 in SW U.S.A. and NW Mexico, and D. glutinosa (Phil.) J.M.Porter in N Chile and S Peru.

10. Giliastrum (Brand) Rydb. Perennial (also annual?) herbs, flowering the first year, glandular puberulent, with sparse nonglandular trichomes, or glabrous; leaves alternate, entire, toothed to pinnatifid, gradual1y reduced in size in the inflorescence; flowers perfect, in dichasia. 10 spp., 8 in North America, one of them up to Caribbean, and two endemics to NW Argentina (Catamarca, Mendoza and San Juan).

11. Ipomopsis Michx. Annuals, long-lived monocarpics, or suffrutescent perennials. 30 spp., 29 in shrublands and forests of W North America, one SE North America, and I. gossypifera (Gillies ex Benth.) V. E. Grant in Andes of Argentina, Chile and Bolivia.

12. Loeselia L. Annual, suffrutescent perennials, or subshrubs. 18 spp. in North America to Central America, although a single species, L. glandulosa Cav. G. Don, extends southwards as far as Venezuela and Colombia in South America.

LECYTHIDACEAE

§ FULLY CHLOROTROPHIC

Genera/species 24/305-310. Distribution pantropical, subtropical regions of South and E Asia. Habit bisexual, usually evergreen trees (sometimes shrubs or lianas). Foetidia has evil-smelling wood. Bark often fibrous. Brazil Nut family. Many species are large trees in lowlands of South America, esp. Brazil. Bertholletia excelsa is a source of income for local populations in Amazonian rainforest in Brazil and Bolivia (Brazil-nut, castanha-do-pará).

A single 1 km² plot 90 km north of Manaus, Brazil, has 38 spp. of Lecythidaceae; continental New World has almost all Lecythidaceae except one in Jamaica (Grias cauliflora L.); only 15 spp. occur in E Brazil, nine endemics. 95 spp. in New World absent in Brazil, including 44 in Eschweilera and 30 in Gustavia; 11 spp. occur in Espírito Santo state. Actinomorphic flowers are found in all species of Old World Lecythidaceae but only in Grias and species of Allantoma and Gustavia in the New World.

SYSTEMATICS 5 subfamilies, two in New World. Napoleonaeoideae (2/c 13, tropical W and C Africa), Barringtonioideae (5/c 84, tropical E Africa, Madagascar, S and SE Asia, Malesia to New Guinea, tropical Australia, Melanesia to the Ryukyu Islands and Micronesia) and Foetidioideae (1/18, Madagascar, the Comoros, Mascarene Islands, tropical E Africa) does not occur in New World.

1. SUBFAMILY SCYTOPETALOIDEAE (6/c 20) ‣ outsiders Oubanguia (3; tropical W and C Africa), Scytopetalum (3; tropical W and C Africa), Rhaptopetalum (11–12; tropical W and C Africa), Pierrina (1; Cameroon, Gabon, Equatorial Guinea), Brazzeia (3; C Africa).

1. Asteranthos Desf. Tree or shrubs to 4-15 m tall, locally common and abundant; petals absent, but the flower have a corolla-like umbel. Only one sp., A. brasiliensis Desf., endemic to the Guiana Shield of E Colombia, SW Venezuela and NW Brazil (upper Rio Negro basin, in Amazonas state), 100 – 200 m elevation range.

2. SUBFAMILY LECYTHIDOIDEAE (10/c 200) ‣ all genera occur in South America.

2. Allantoma Miers. (inc. Cariniana p.p.) Upper canopy and emergent trees; medium to large sized leaves (6-30 cm long), glabrous, tertiary venation percurrent; inflorescence racemose or paniculate; flowers with actinomorphic androecium; 5 sepals and 5 petals. 8 spp., 4 endemic to N Brazil (3 in Amazonas and one in Rondonia, all rare plants in Brazil, by Plantas Raras do Brasil’s book), 3 remaining in Brazil up Peru, Bolivia, Ecuador, Colombia and Venezuela; from the headwaters of the Negro River to the mouth of the Amazon River in Brazil and on the eastern side of the Amazon River except A. lineata (Mart. & O. Berg) Miers; not found in the Guianas.

3. Bertholletia Bonpl. Trees to 62 m tall, exceptionally large trees may reach the age of 800-1,200 years; leaves not clustered at end of branches; trunk cylindrical with deep fissures in the bark; leaves medium to large (20-30 cm long), glabrous, obovate to oblong; flowers zygomorphic; calyx with two sepals; 6 petals. Only one species, B. excelsa Humb. & Bonpl. (Brazil Nut), Colombia, Peru, Bolivia, N Brazil to Venezuela, and Guianas.

4. Cariniana Casar. (exc. Allantoma p.p.) Small to large trees to 65 m tall, either canopy trees, the tallest of all Ericales in Brazil; flowers nearly actinomorphic to zygomorphic, the smallest at Lecythidaceae. 9 spp., 4 endemics to Brazil, 4 in Brazil up to Bolivia, Colombia, Venezuela, Peru and Paraguay, and C. pyriformis Miers from Central America, Colombia and Venezuela; two are rare plants in Brazil, by Plantas Raras do Brasil’s book.

An individual from C. legalis (Mart.) Kuntze in Santa Rita do Passo Quatro is identified by many sources as being around 3,000 years old; however, as highlighted by other sources, there is contention and controversy about this age, and cites that she may not even be 1,000 years old; SDb does not accept this supposed age of,3000 years.

5. Corythophora R. Knuth. Large trees; leaves not clustered at end of branches; flower zygomorphic. 4 spp., two in Brazil and Guianas (Guyana, Suriname and French Guiana), C. alta R. Knuth endemic to Brazil, and C. labriculata (Eyma) S.A. Mori & Prance only in Guianas.

6. Couratari Aubl. Canopy and emergent trees up to 40 m tall; leaves small to medium (3-20 cm long), glabrous or pubescent; flowers zygomorphic, with 6 sepals and 6 petals. 19 spp., distributed through Central America (Costa Rica and Panamá), the western Andes (Colombia), throughout the Amazon rainforest and in the coastal forests of the Brazilian states of Bahia, Espírito Santo, and Rio de Janeiro; 14 spp. in Brazil, 7 endemics, 4 of then are rare plants in Brazil, by Plantas Raras do Brasil’s book.

7. Couroupita Aubl. Medium to large-sized trees, cauliflorous. Three spp., C. nicaraguarensis DC. from Nicaragua to Ecuador, C. guianensis Aubl. (cannon-ball tree) over Amazon rainforest, and C. subsessilis Pilg. restricted of Brazil and E Peru; the cannon-ball tree is often used as a street-tree in Rio de Janeiro, despite the weight of its fruits that threaten parked vehicles.

8. Eschweilera Mart. ex DC. Small to large trees up to 40 m tall; trunk cylindrical, with or without large buttress roots; leaf blade glabrous with brochidodromous venation; petiole with flat-convex outline; flowers zygomorphic; calyx with 6 sepals; corolla with 6 petals. 105 spp., Trinidad-Tobago, Honduras to French Guiana, Bolivia and Brazil (51, 21 endemics, five in Amapá, Para, Bahia and Rio de Janeiro as rare plants in Brazil, by Plantas Raras do Brasil’s book); the only mexican species of this family in the endemic Eschweilera mexicana T. Wendt, S.A. Mori & Prance; 25 spp. are adapted to montane forests in Andes; 89 in South America.

9. Grias L. Trees to 30 m tall, mainly pachycaulous unbranched trees, leaves largest (possibly largest among Ericales, up to 2.2 m lenght), cauliflorous; leaves clustered at end of branches, fleshy mesocarps, edible in some species; flowers actinomorphic. 12 spp., restricted from Colombia (3 endemics), Ecuador (4 endemics) and Peru except G. multinervia Cuatrec. up to Venezuela and G. cauliflora L. also in Jamaica and Belize.

10. Gustavia L. Small to large trees, leaves often clustered at end of branches, inflorescenses axillary or cauline. 46 spp., Costa Rica to French Guiana, N Bolivia, Ecuador and Brazil (10, 3 endemics, all are rare plants in Brazil, by Plantas Raras do Brasil’s book, in Roraima and Pará states); some spp. are adapted to montane forests in Andes.

11. Lecythis S.A.Mori. Small to medium size trees up to 60 m tall; leaves not clustered at end of branches; flowers zygomorphic. 28 spp., Costa Rica to French Guiana, Peru and Brazil (22, 10 endemics, two in Amazonas state are rare plants in Brazil, by Plantas Raras do Brasil’s book), all in South America, 4 up to Central America; some spp. in savannha of Roraima and savannas of C Brazil (cerrado) are adapted to dry environments.

LINEAGE 3 of 6: PENTAPHYLLACOIDS

PENTAPHYLACCACEAE

§ FULLY CHLOROTROPHIC

Genera/species 12/c. 340 Distribution tropical and subtropical regions, Himalaya and E Asia to Korean Peninsula and Japan, SE Asia to New Guinea and Queensland, Melanesia, islands in the SW Pacific, Hawaii, Canary Islands, Mexico, Central America, Caribbean, tropical South America; Balthasaria in E Africa, two species of Ternstroemia in tropical W Africa. Habit bisexual or dioecious, evergreen trees and shrubs. Use Ornamental plants, timber.

Four Neotropical and thirteen worldwide with a total of approximately 300 species; found throughout the Neotropics in montane areas, particularly in cloud forests.

SYSTEMATIC subfamily Pentaphyllacoideae (1/1, Guangdong, Hainan, N Indochina, the Malay Peninsula, Sumatra) do not occur in South America.

1. SUBFAMILY TERNSTROEMIEAE (2/c 100) ‣ outsider Anneslea (3). Tropical, with their largest diversity in Malesia and in Central and South America.

1. Ternstroemia Mutis ex L.f. Shrubs or trees; leaf margin entire. c. 140 spp., 106 sp. in the New World (Mexico to Argentina, 65 in South America, 23 in Brazil, 12 endemics), 2 in Africa, and 40 in SE Asia. T. washikiatii Cornejo & C.Ulloa (Ecuador) the large leaves (21–33 ✕ 8.5–11 cm), and the large fruits (4.5–5.5 ✕ 6.5–7.7 cm) of this genus.

2. SUBFAMILY FREZIEREAE (9/233) ‣ outsiders Visnea (1; Madeira, Canary Islands), Adinandra (c 85; India, Sri Lanka, Burma, China, S Japan, SE Asia, Malesia to New Guinea, tropical Africa), Archboldiodendron (1; mountains on New Guinea), Cleyera (18; Mexico, Central America, Caribbean, one species, C. japonica, in Himalayas and China to Korean Peninsula and Japan); Euryodendron (1; S China), Eurya (c 70; tropical and subtropical regions in Asia, islands in the W Pacific, Hawaii), Balthasaria (1; tropical Africa).

2. Freziera Willd. (inc. Killipiodendron) Small dioecious trees or shrubs; leaf margin serrate. 64 spp., S Mexico to S Bolivia, E to Guyana, Caribbean, and E Brazil, mainly primarily montane, 56 in South America; three spp. are found in montane forests of the E Venezuela: F. calophylla Triana & Planch. off Amazonas state in N Brazil, also in W Venezuela and Ecuador; F. roraimensis Tul., endemic to the Mount Roraima off Roraima state; and F. carinata A. L. Weitzman, in Venezuela and Brazil (in N Amazonas state); the second species in Brazil is F. atlantica Zorzanelli & Amorim, discovery in 2015, known from two disjunct localities in the Atlantic Forest of Brazil, in the states of Bahia and Espírito Santo.

3. Symplococarpon Airy Shaw. Trees; leaf margin entire to serrate. Two spp., S. purpusii (Brandegee) Kobuski from Mexico to Colombia and Venezuela, and S. flavifolium Lundell endemic to Mexico.

LINEAGE 4 of 6: PRIMULOIDS

SAPOTACEAE

§ FULLY CHLOROTROPHIC

Genera/species 75/1,250-1,300 Distribution mainly pantropical; some species in subtropical regions (SE North America, central South America, SE Africa, S Asia). Habit usually bisexual (sometimes monoecious or dioecious, rarely gynomonoecious), evergreen trees or shrubs (rarely lianas). Sapotaceae is a largely tropical family of evergreen trees and shrubs. There are 53 genera and about 1,100 species in the family, but generic limits in the family are notoriously difficult and changeable. Manilkara yield edible fruit, useful wood and latex.

The best-known species are M. bidentata (A. DC.) A. Chev. (balata), M. chicle (Pittier) Gilly (Chicle) and M. zapota (L.) P. Royen (Sapoti). A South America forest family by excelence. Latex of Sapotaceae is a source of gutta-percha, balata, and chicle, either pure trans-polyisoprene polymers or a mixture of cis and trans constituents. The berries of a number of species are edible. 11 genera and 220 spp. in Brazil.

Key differences from similar families The families listed below differ from Sapotaceae as follows:

Ebenoideae: lacks white latex, corolla lobes contorted in bud, stamens usually twice as many as corolla lobes, ovary locules 2-ovulate, styles distally or completely free, seed coat without hard shiny surface.

Lissocarpoideae: lacks white latex, indumentum of simple hairs, inflorescence a cyme, corolla lobes contorted in bud, corolla throat bearing a tubular corona.

Symplocaceae: lacks white latex, indumentum of simple hairs, leaf margins serrate or dentate, fruit a drupe.

SYSTEMATICS three clades, Sarcospermatoideae (2/14, E Himalayas, S China, SE Asia, Malesia) are absent in New World.

1. SUBFAMILY CHRYSOPHYLLOIDEAE (36/660–680) ‣ outsiders outsiders: Amorphospermum (1, Australia), Aningeria (2–8, Africa), Aubregrinia (1, Africa), Breviea (1, Africa), Delpydora (2, Africa), Donella (1, Africa), Englerophytum (20, Africa), Gambeya (16, Africa), Magodendron (2, New Guinea), Malacantha (1, Africa), Neohemsleya (1, Africa), Niemeyera (1, Australia), Omphalocarpum (27, Africa), Pichonia (13, Australasia), Planchonella (110, Asia, Oceania), Pleioluma (40, Asia, Oceania), Pycnandra (59, New Caledonia), Sahulia (1, New Guinea), Sersalisia (5, Australia, New Guinea), Spiniluma (2, Africa, Arabia), Synsepalum (36, Africa), Tridesmostemon (2, Africa), Van-royena (1, Australia), Xantolis (14, Africa, Asia).

1. Achrouteria Eyma. 3 spp., from Guyana, northern Brazil (all, one endemic) and Venezuela.

2. Chloroluma Baill. Two species distributed in N Argentina, SE Brazil, Bolivia, Paraguay and Uruguay.

3. Chromolucuma Ducke. Trees; leaves subtended by large stipules, calyx of 5 imbricate sepals, corolla cup-shaped or tubular, stamens included, staminodes present, seed with dull rough testa. 9 spp., C. cespedesiiformis J.F. Morales in Costa Rica and 8 in South America, all in Brazil, 3 endemics.

4. Chrysophyllum L. Trees or shrubs, very rarely lianas; stipules absent, calyx of 5 imbricate sepals, corolla cup-shaped to tubular, stamens included, staminodes absent, seed scar lateral (adaxial). 25-30 spp. confined to the Neotropics, including Central and South America, and the Lesser and Greater Antilles.

5. Cornuella Pierre. Only one sp., C. venezuelanensis Pierre, widely distributed from southern Mexico, across Central America and the Andean countries to Bolivia. Recorded from French Guiana and Suriname, probably extending into the Brazilian Amazon.

6. Diploon Cronq. Trees with leaves spaced; stipules absent, calyx of 5 imbricate sepals, corolla rotate, stamens exserted, staminodes absent, ovary unilocular with 2 basal ovules, seed with a basal scar. Only one sp., D. cuspidatum Hoehne, Venezuela and Guianas to SE Brazil, and Ecuador to Bolivia.

7. Ecclinusa Mart. Trees (up to 30 m tall) or rarely shrubs, white latex; stipules present, flowers sessile, calyx of 5 imbricate sepals, staminodes absent, seed scar lateral (adaxial) and extending around the base of the seed. 12 spp., mainly Amazon rainforest, only one reaches to Atlantic Forest in Brazil (6 spp., two endemics).

8. Elaeoluma Baill. Trees or shrubs; stipules absent, leaves usually minutely punctate, calyx of 5 imbricate sepals, corolla cup-shaped to rotate, stamens exserted, staminodes absent, seed scar lateral (adaxial). 5 spp., 4 from Colombia, Venezuela, Guianas and Amazonian rainforest of Brazil, and E. glabrescens (Mart. & Eichler) Aubrév. in Central America, Venezuela, Guianas, Colombia, Ecuador, Peru, Bolivia, Brazil.

9. Englerella Pierre. Only one sp., E. macrocarpa Pierre, from Colombia, Guyana, French Guiana to N Brazil.

10. Gayella Pierre. Treelet 2–3(–6) m tall, with fawn indumentum on young branches, petiole, midvein, pedicels and sepals, turning greyish with age. Only one sp., G. valparadisaea (Molina) Pierre, restricted to the provinces of Choapa (Coquimbo Region) and San Antonio (Valparaíso Region) of central Chile, in rocky slopes, ravines and gullies, usually below 100 m altitude and within reach of sea mist, but with two subpopulations up to 400 m.

11. Labatia Sw. 16 spp. from Brazil (14, 8 endemics) up to over South America up to Nicaragua, one from Costa Rica to Colombia, and one restricted to Caribbean.

12. Lucuma Molina. (inc. Pouteria p.p.) 18 spp., one widely from Caribbean to Panama and Brazil, one from Mexico to Central America, one from U.S.A. to Caribbean, remaining only in South America. 15 spp. in Brazil, 5 endemics.

13. Martiusella Pierre. Trees. Leaves with spinous-serrate margin and minute translucent veins; tertiary venation oblique; flowers 5–6-merous. Only one sp., M. imperialis (Linden ex K.Koch & Fintelm.) Pierre, endemic to E Brazil.

14. Micropholis (Griseb.) Pierre. (inc. Pouteria p.p.). Trees or shrubs up to 50 m tall in rainforests, leaves spaced; stipules absent, leaves with numerous fine closely parallel secondary veins, calyx a single whorl of 5 imbricate sepals, corolla cup-shaped or tubular, stamens included or exserted, staminodes present, seed scar lateral (adaxial). 38 spp., 36 in South America, 29 in Brazil, 6 endemics (4 of then, all in Amazonas state, are rare plants in Brazil, by Plantas Raras do Brasil’s book), in two sections:

§ sect. Micropholis ‣ 29 spp. throughout tropical America from Mexico to Bolivia and Brazil, and Caribbean.

§ sect. Exsertistamen ‣ 9 spp. Guianas to Peru and Brazilian Amazon rainforest.

15. Nemaluma Baill. Two spp. from N Brazil, both up to Guianas, one up to Venezuela.

16. Peteniodendron Lundell. 4 spp., P. durlandii (Standl.) Lundell widely in Neotropics, three remaining mainly restricteds for Brazil, two endemics and one up to Guianas.

17. Pouteria Aubl. (exc. Lucuma p.p.) Trees or shrubs up to 50 m tall, rarely with root rhizomes; leaves spirally arranged, rarely opposite; stipules nearly always absent, calyx of 4 or more sepals in a single imbricate whorl, corolla cup-shaped to tubular, stamens included, staminodes present, seed scar lateral (adaxial). 6 spp., all in Brazil, one endemics and remianing widely, some up to Nicaragua.

18. Pradosia Liais. Tall canopy trees up to 40 m tall, treelets, sometimes with xylopodium, leaves opposite or verticilate, rarely less frequently spirally arranged, some buttressed. 26 spp. from South America (all), one reaching into Costa Rica and Panamá; 17 spp. in Brazil, 8 endemics.

Only P. brevipes (Pierre) T. D. Penn., from Brazil and Paraguay, has a specialized habit with most of the plant being below the ground, with only a few branches emerging above the ground. P. huberi (Ducke) Ducke has been recorded in permanent flooded-forests, or swampy land in northeast Amazon rainforest; 3 spp. are found in dry montane forest up to 1,200 m altitude in north and NW South America.

19. Prieurella Pierre. 5 spp. from Nicaragua in the north, Guianas, and the Amazon basin, 3 in Brazil, none endemics.

20. Ragala Pierre. 4 spp. from northern South America, all in Brazil, none endemics.

21. Sarcaulus Radlk. Trees; stipules absent, calyx of 5 imbricate sepals, corolla globose, carnose, staminodes present, scar lateral (adaxial). 105 spp., 5 spp. in Sarcaulus s.s., from Ecuador, Peru and E Brazil one endemic each, S. vestitus (Baehni) T.D. Penn. disjunct in Brazil and Ecuador, S. brasiliensis (A. DC.) Eyma widely distributed fom S Central America to Bolivia and Brazil; and c. 100 spp., formerly placed within Pouteria, composed the clade Q, without several data.

2. SUBFAMILY SAPOTOIDEAE (c. 23/515–520) ‣ five lineages (mainly tribe level), two in New World, Lecomtedoxa/Neolemonniera clade (2/9, Gabon, tropical W Africa), Tseboneae (3/25, Madagascar) and Isonandreae (7/c. 260, Seychelles, India, Sri Lanka, SE Asia, Malesia to New Guinea, tropical Australia) absent.

2.1 SAPOTOIDEAE ▸ TRIBE SAPOTEAE (10/c. 140) - outsiders Inhambanella (2; W and SE tropical Africa), Vitellariopsis (5; E Africa), Vitellaria (1; tropical W and C Africa), Tieghemella (2; tropical W and C Africa), Mimusops (c 40; tropical Africa, Madagascar, Mascarene Islands, Seychelles, one species to tropical Asia), Autranella (1; tropical W and Central Africa), Labramia (9; Madagascar), Labourdonnaisia (7; Madagascar, Mauritius, Réunion), Faucherea (11; Madagascar).

22. Manilkara Adans. Trees, rarely shrubs, nearly always with sympodial branching, rarely dioecious, calyx of 2 whorls of 3 sepals, outer whorl valvate, corolla lobes 6, usually divided into 3 segments, staminodes present, seed scar usually basi-ventral. 20 spp. from tropical Africa and Madagascar, 15 in Asia, Australia, Pacific, and 29 spp. from Florida, Mexico to SE Brazil (15, 11 endemics, 4 spp. in Bahia and one in Pernanbuco state are rare plants in Brazil, by Plantas Raras do Brasil’s book), Caribbean; 20 spp. in South America.

2.2 SAPOTOIDEAE ▸ TRIBE SIDEROXYLEAE (1/75–80) ‣ a single genus.

23. Sideroxylon L. Often spiny, calyx a single whorl of 5 imbricate sepals, corolla lobes divided into 3 segments or entire, staminodes present, seed scar basal or basiventral. 83 spp. worldwide, 6 in Africa, 6 in Madagascar, 8 in Mascarenes, 4 in Asia, 1 in NW Pakistan and Afganinthan, Oman, Ethiopia, Djibuti and Somalia; 55 in New World, only 4 in South America, three from Central America over Colombia/Venezuela region, and one, S. obtusifolium (Humb. ex Roem. & Schult.) T.D. Penn., widely distributed, and only spp. in Brazil.

EBENACEAE

§ FULLY CHLOROTROPHIC

Genera/species 3/802. Distribution pantropical, with their highest diversity in Malesia; some species in temperate parts of E North America, India, E China, and Japan. Habit usually dioecious (rarely monoecious, polygamomonoecious or bisexual), usually evergreen (rarely deciduous) trees or shrubs. Bark, roots and heartwood often black or, in air, blackening. Species of Diospyros are of economic importance for the wood that several produce and for their fruits. The wood, which is either uniformly dark (ebony) or variously streaked and marbled, has been much used in furniture making. The fruit (date plums, persimmons) can be very astringent if eaten before they are fully ripe.

SYSTEMATIC both genera in this family occcur in South America.

1. Diospyros L. Trees or shrubs, bark often black; inflorescence cymose in male flowers and a single flower in female flowers. 500-600 spp., pantropical, 200-300 in Asia and Pacific, 95 in Madagascar, 110 in Africa mainland, 15 in Australia and 126 in over New World (57 in Brazil, 30 endemics, 16 spp. in several Brazilian states (mainly in Amazon rainforest) are rare plants in Brazil, by Plantas Raras do Brasil’s book); Euclea (Africa) and Royena (Africa) have sometimes been identified as sister-group to Diospyros.

2. Lissocarpa Benth. Treelets, small or, less frequently, medium tall trees without latex, glabrous on all organs (except on stigmas); roots probably black; leaves alternate, simple; flowers axillary, solitary along the proximal part of long-shoots; flowers actinomorphic, 4-merous (very rarely 5-merous), unisexual (apart from many completely sterile flowers); fruit wall thin, fleshy, pink or red at maturity when alive. 9 spp. 5 from Colombia to Bolivia (one endemic to Peru); Venezuela and Guyana one endemic each; and two in Brazil: L. benthamii Gurke, in Upper Rio Negro at SW Venezuela, SE Colombia, NW Brazil (only Amazonas state, very common in some places), along or near black-water rivers (bosques bajos de rebalse en las orillas) in seasonally flooded, usually ‘low and open’ forests; and L. kating B.Walln, known from Brazil (Amazonas), Colombia (Vaupés), and Peru (Loreto), where it grows at elevations between 90 and 250 meters, in primary, non-flooded (‘terra-firme’) rain forests).

PRIMULACEAE

§ FULLY CHLOROTROPHIC

Genera/spp. 57/2040–3050 Distribution cosmopolitan except polar areas. Habit usually bisexual (rarely monoecious, polygamomonoecious or dioecious), evergreen or deciduous trees (in), shrubs, lianas or suffrutices, perennial or annual herbs (sometimes cushion-shaped and woody at base), usually with a basal leaf rosette. Some species are xerophytes. Hottonia and Samolus are aquatic. Aegiceras mangrove trees. Cyclamen has a tuber formed by the hypocotyl. Use Ornamental plants, medicinal plants.

The biggest problem for APG III was the question of how to treat Primulaceae and their immediate relatives, a closely related group that in the past has often been recognized as a separate order. Primulaceae has 58 genera and 2590 spp. in worldwide. APG III include in this family the ex-familys. Eight genus and 120 spp. in Brazil.

Locally, the wood of several species is used for rustic construction (fencing) and as fuel (wood/charcoal), and fruit with thick mesocarps are often aten. Species of Ardisia, Cyclamen, Lysimachia and Rapanea are cultivated as ornamental plants, including trees. Some species of Anagallis are introduced weeds. In other regions of world some species are used in folk medicine.

SYSTEMATIC subfamily Maesoideae (1/100-110, tropical Africa, SE Africa, Madagascar, Yemen, Sri Lanka, E Asia to Japan, SE Asia, Malesia to New Guinea, Melanesia, Queensland, islands in the Pacific) do not occur in South America.

1. SUBFAMILY PRIMULOIDEAE (6/540–550) ‣ outsiders Omphalogramma (15; Himalayas, W China), Hottonia (2; Europe, W Asia, SE Canada, E U.S.A.), Soldanella (16; mountain regions in C and S Europe), Bryocarpum (1; E Himalayas).

1. Androsace L. 100 spp., almost all restricted of Northern Hemisphere except by A. salasii Kurtz native from S. Chile to W. Central & S. Argentina.

2. Primula L. Perennial herbs, sometimes very small. 523 spp., native to the Northern Hemisphere, with the distribution extending southwards to Ethiopia, Indonesia, New Guinea, 21 spp. in New World, with two, P. magellanica Lehm. and P. comberi W.W. Sm., in the southern tip of South America, also in Falklands Is.

2. SUBFAMILY MYRSINOIDEAE (c 40/1,500–2,500) ‣ outsiders Coris (1; W and C Mediterranean, Somalia), Ardisiandra (3; mountains in tropical Africa), Cyclamen (22; S Europe, Mediterranean to the Caucasus and Iran, NE Somalia), Embelia (c 130; tropical and subtropical regions in the Old World), Grenacheria (12; Malesia), Heberdenia (1; Madeira, Canary Islands), Pleiomeris (1; Canary Is.), Solonia (1; Cuba), Emblemantha (1; Sumatra), Sadiria (4; Assam, E Himalayas), Antistrophe (6; tropical Asia from N India to SE Asia and Malesia), Aegiceras (1; coastal areas in SE Asia to islands in the Pacific), Amblyanthus (3; E Himalayas to SE Asia, Malesia to New Guinea), Amblyanthopsis (3; Himalayas to SE Asia and W Malesia), Elingamita (1; Three Kings Islands in New Zealand), Loheria (10; Philippines, New Guinea), Vegaea (1; Hispaniola), Oncostemum (c 90; Madagascar, Mascarene Islands), Badula (17; Madagascar, Mauritius, Réunion), Discocalyx (c 105; Central Malesia to tropical Australia, Melanesia and Polynesia, with their largest diversity in Philippines and on New Caledonia), Labisia (17; SE Asia, Malesia), Systellantha (3; Borneo), Monoporus (9; Madagascar), Fittingia (7; New Guinea), Conandrium (2; E Malesia to New Guinea and Bismarck Archipelago).

3. Ardisia Sw. Shrubs or treelets, often subshrubs or herbs. 450 spp., pantropical, mainly tropical America and Asia, a few in temperate Japan; is found in much of the family’s range but not in Africa; 169 spp. in New World, 56 in South America, only 5 in Brazil (all in Amazon rainforest, A. semicrenata Mart. up to Atlantic Forest), two endemics; Ardisia subg. Graphardisia is a small distinctive subgenus of 7 spp. with nine subspecies found from Mexico to Bolivia and adjacent Brazil.

4. Ctenardisia Ducke. Shrubs monocaulous or small trees; inflorescences terminal panicles. 4 spp., one endemic to Mexico, another only in Central America, C. stenobotrys (Standl.) Lundell & Pipoly in Venezuela and Amazonas state in Brazil, and C. speciosa Ducke endemic to Pará state.

5. Cybianthus Mart. (inc. Conomorpha) Shrubs or small trees, often epiphytic; vegetative and floral parts sometimes brownish lepidote. 157 spp., 152 in South America, fully neotropical, 66 in Brazil, a exact half endemics. 11 subgenera, Conomorpha, Cybianthus, Laxiflorus and Weilgetia occur in Brazil; Comomyrsine, Cybianthopsis, Grammadenia, Iteoides, Micronomorpha, Stapfia e Triadophora are absent.

6. Geissanthus Hook.f. Trees or shrubs, inflorescence in terminal panicles. 50 spp., all from Venezuela to Bolivia, two up to center and SE Brazil (São Paulo state), and one up to Mesoamerica.

7. Gentlea Lundell. Shrubs or small trees; inflorescence terminal panicles or umbel. 11 spp. from Mexico to N and NW South America, up Peru, two in South America, one endemic to Peru and G. venosissima (Ruiz & Pav.) Lundell disjunct in Mexico, Central America, Venezuela and Roraima state in N Brazil.

8. Hymenandra A. DC. ex Spacht. 16 spp., 7 spp. in Indo-Malesia region (Assan to Borneu) and 7 spp. from Nicaragua to Panamá, and two endemics to Choco region in Colombia.

9. Lysimachia L. (inc. Anagallis, Pelletiera). Annual or perennial herbs, rarely shrubs; flowers in axis or upper leaves, in panicles or racemes, sometimes cephalium. c. 210 spp. worldwide, including 4 ancient species growing in New World, with white flowers and morpholocally distinct (none in Brazil), and others moved from Anagalis and Pelletiera; 34 spp. in New World, 16 in South America, 12 in Brazil, 8 endemics (almost all in southern country).

10. Myrsine L. (incl. Rapanea, Suttonia). Shrubs or trees, sometimes with roots crown. 300 spp., pantropical; 74 spp. in New World, 61 in South America, 25 in Brazil, 15 endemics, one of them, from Minas Gerais state, is a rare plant in Brazil, by Plantas Raras do Brasil’s book.

11. Parathesis (DC.) Hook.f. Trees or shrubs, often with stellate or hairs dendroid, poliaxyal, mainly ferruginous-tomentose. 96 spp., Mesoamerica, Andes of South America and Caribbean, 20 in South America, two in Brazil, none endemics, in Amazonas, Acre and Rondonia; two subgenera:

§ subg. Parathesis ‣ three sections 54 spp., inc. both Brazilian members of genus.

§ subg. Lateralis ‣ 42 spp. within 4 sections, absent in Brazil.

12. Stylogyne AD. C. Small androdioecous, dioecious, or bissexual shrubs or trees, with small and delicate 4-merous, or more often, 5-merous campanulate flowers. 40 spp. of Neotropics, 34 in South America, 18 spp. in Brazil, 9 restricted of Amazon rainforest (Xingu eastwards, one of then endemics), and 9 restricted of Atlantic Forest (all endemics, one in Bahia, 8 from Espírito Santo to border of Santa Catarina and Rio Grande do Sul states, three of them, all from Rio de Janeiro state, are rare plants in Brazil, by Plantas Raras do Brasil’s book).

3. SUBFAMILY SAMOLOIDEAE (1/c 12) ‣ a single genus.

13. Samolus L. Herbs, short-lived perennial found in small colonies by permanently wet and often seasonally flooded springs, flushed sea-cliffs, puddles, ditches, lagoons and lake shores, reed (Phragmites) stands, wet pastures, and saline meadows, in more or less saline habitats such as small, brackish ponds near the seashore or in salt marshes. 12 spp., six in North America to Central America and Caribbean, national endemics in South Africa (1) and Chile (2); S. repens Pers occur in Australia, New Zealand, New Caledonia and Chile; S. subnudicaulis A. St.-Hil. occur in S Brazil and adjacent Cono Sur; and S. valerandi L. in southern half fo South America, Canada to Guatemala, S Africa, Australia, New Zealand, Europe to India, S China, Japan, Magred and sorrounding Sahhara desert in northern Africa.

4. SUBFAMILY THEOPHRASTOIDEAE (7–6/90–95) ‣ outsiders Theophrasta (2; Hispaniola), Neomezia (1; Cuba) and Deherainia (3; Mexico to Honduras).

14. Bonellia Colla. 20 spp., a half only in Mexico and Central America, six restricted of Caribbean, and 4 restricted of Andes from Venezuela to Peru.

15. Clavija Ruiz & Pav. Unbranched or sparsely unbranched shrubs or treelets stems spini. 56 spp., 55 in South America (5 up to Central America); and the only Antillean species, C. domingensis Urb. & Ekman, sister to all other species occurring in South and Central America; 10 spp. in Brazil, two endemics.

16. Jacquinia L. Shrubs or small trees, richly branched. 14 spp., 12 restricted of Caribbean and adjacent Mexico and Florida, J. arborea Vahl from Mexico to Panamá and Venezuela, and J. armillaris Jacq. in Caribbean, Colombia, Venezuela and N Brazil, at vast areas in Atlantic sandy coastal shrublands (restingas), inc. Fernando de Noronha Is., in sand psamophillous soils.

17. Votschia Stahl. Only one sp., V. nemophila (Pittier) B. Stahl, apparently endemic to Panamá, but collected at 3 km off Colombian border in Pacific Coast.

LINEAGE 5 of 6: THEOIDS

THEACEAE

§ FULLY CHLOROTROPHIC

Genera/species 8/250–450 Distribution SE U.S.A., Caribbean, Central America, tropical South America, E Asia to Korean Peninsula and Japan, SE Asia, Malesia, New Guinea. Habit bisexual, usually evergreen (rarely deciduous) trees and shrubs. Use Ornamental plants, stimulants, seed-oils, timber.

SYSTEMATIC three tribes, Gordonieae (3/3–60, SE Asia, W Malesia, Georgia in U.S.A.) and Stewartieae (1/9–30, C China, Korean Peninsula, Japan, SE U.S.A.) does not occur in South America; among Theeae, outsiders are Camellia (c 125; E Asia to Japan and Taiwan in China, tropical Asia), Polyspora (c 40; S China, SE Asia), Pyrenaria (42; S China, SE Asia, W Malesia), Apterosperma (1; China).

1. Laplacea Kunth. Trees, with flowers, leaves at the tip of branches, inflorescences terminal. 23 spp. from Old World, 13 endemic to Caribbean, national endemics in Brazil (2, L. acutifolia (Wawra) Kobuski and L. tomentosa (Mart.) G.Don.), Colombia (2), U.S.A. (1), Ecuador (1), L. spathulata Kobuski from Brazil to Peru, L. brenesii Standl. from Mexico to Panamá, L. barbinervis Moric. from Colombia to Ecuador, and L. fruticosa (Schrad.) Kobuski from Nicaragua to Guianas, Bolivia and Brazil.

SYMPLOCACEAE

§ FULLY CHLOROTROPHIC

Genera/species 1/406 Distribution SE U.S.A., southern Mexico, Central America, Caribbean, tropical South America, tropical and subtropical areas in South, E and SE Asia, Malesia, Melanesia, E Australia; some species in temperate regions of E Asia and E North America. Habit usually bisexual (rarely polygamomonoecious or dioecious), usually evergreen trees or shrubs (Symplocos paniculata (Thunb.) Miq. is deciduous). Use Beverages (maté), dyeing sources, carpentries, carvings.

SYSTEMATIC outsider Cordyloblaste (2, E Asia).

1. Symplocos Jacq. Trees or shrubs, evergreen or deciduous; mature current-year branchlets green or brown; leaf midvein sulcate, flat, or prominent adaxially; hermaphroditic or dioecious; inflorescences axillary or pseudoterminal, or irregularly concaulescent; bracteoles subtending flowers caducous or persistent. 346 spp., 144 in Old World, 202 in southern U.S.A., Caribbean, Central Mexico to S Brazil (45, 36 endemics, 9 of them, all in Minas Gerais or Rio de Janeiro states, are rare plants in Brazil, by Plantas Raras do Brasil’s book), Argentina and Paraguay, particularly but not exclusively in montane areas; lacking or very rare in the entral Amazon rainforest. 152 spp. in South America. Two subgenera:

§ subg. Symplocos ‣ almost over genus.

§ sect. Symplocos ‣ 178 species, tropical America.

o ser. Symplocos ‣ 171 spp., tropical America; the informal Neosymplocos group has 13 spp. from Brazil and Paraguay.

o ser. Urbaniocharis ‣ 7 spp., Caribbean.

§ sect. Barberina ‣ 25 spp., S. wikstroemiifolia Hayata in Asia and remaining in New World.

§ sect. Lohdra ‣ 142 spp., E Asia, Australasia.

§ subg. Palura ‣ only one sp., S. paniculata (Thunb.) Miq., from E Asia.

STYRACACEAE

§ FULLY CHLOROTROPHIC

Genera/species 12/c. 160 Distribution The U.S.A., Mexico, Caribbean, Central America, South America southwards to Argentina, the N and E Mediterranean, E Himalaya (Nepal to Arunachal-Pradesh), Assam, E India, E Asia to Korean Peninsula and Japan, SE Asia, Malesia, New Guinea, Solomon Islands. Habit usually bisexual (in Bruinsmia gynodioecious), evergreen or deciduous trees or shrubs.

Use Ornamental plants, medicinal plants, balsam resins (benyamin-gum, benzoin etc. from Styrax) for pharmaceutical, confectionery and perfume industries, incense.

SYSTEMATIC outsiders are Huodendron (4; S China, se Asia); Bruinsmia (2; E Himalayas, Burma, S China, Indochina, Malesia to New Guinea), Alniphyllum (3; E Himalayas, SW and C China (inc. Taiwan), SE Asia), Halesia (2; SE U.S.A.), Melliodendron (1; S China), Changiostyrax (1; S China), Perkinsiodendron (1; E China), Rehderodendron (5; S and W China, Burma, Vietnam), Pterostyrax (4; Burma; China, Japan), Sinojackia (8; S China), Parastyrax (2; Yunnan, Burma).

1. Styrax L. Trees (4-30 m), sometimes with developed xylopodia, which have a strong capacity to form new sprouts after fire. 130 spp.; widely distributed but disjunctive distribution, New World (95 spp., up to Argentina), E Asia, and Mediterranean region; 70 spp. occur in South America, two in meditarranean vegetation in California, several occurs in savannah in center Brazil and Andean paramo; some Styrax in Brazil (25, 10 endemics).

LINEAGE 6 of 6: ERICOIDES

MITRASTEMONACEAE

§ PARASITIC (Prosopanche - Cassyta - Bdalophytum - Krameria - Mitrastemon - APODANTHACEAE – ... - Cuscuta)

Genera/species 1/2 Distribution New World and Asia. Habit bisexual, achlorophyllous, herbaceous endophytes without rhizome or normal roots. Holoendoparasites on roots of Fagaceae (Castanopsis, Lithocarpus, Quercus, and Trigonobalanus); isophasic parasitism.

SYSTEMATIC two achlorophyllous spp. in single genus.

1. Mitrastemon Makino. Two spp. worldwide, M. yamamotoi Makino from NE India, Indochina, Malesia (including New Guinea), Taiwan (China), the Ryukyu Islands, Japan; and M. matudae Yamam. from S Mexico, Guatemala, NW Colombia, in fagaceous forests.

ACTINIDACEAE

§ FULLY CHLOROTROPHIC

Genera/species 3/447 Distribution S and E China, Korean Peninsula, Japan, Siberia, SE Asia, Himalayas, Malesia to New Guinea, NE Queensland, Solomon Islands, Fiji, Mexico and Central America, northern Andes to Bolivia). Habit bisexual, morphologically dioecious (Clematoclethra), functionally dioecious or monoecious, evergreen or deciduous trees, shrubs or lianas. Use Ornamental plants, fruits.

SYSTEMATICS outsiders Clematoclethra (1; W and C China), Actinidia (55; S and E China, Korean Peninsula, Japan, Siberia, SE Asia).

1. Sauraia Wild. Tall (up 30 m) to small trees or woody shrubs, usually pubescent. 391 spp., c. 300 from Himalaya to Fiji, 87 spp. from Mexico to Bolivia, and one sp. in Queensland; 60 spp. in South America (59 confined), in Venezuela (8, 1 endemic), Colombia (34, 22 endemics), Ecuador (23, 14 endemics), Peru (11, 5 endemics) and Bolivia (6, 3 endemics); only the canopy S. yasicae Loes. definitely wider range in Neotropics; 27 spp. occur only from Mexico and Central America, 22 are treated in Hunters (1966) revision. 7 sections at Soejarto DD (1980, only South Americans), with only 49 spp. at to date:

§ sect. Omichlophilae - 4 spp., all endemic to Colombia, 1,700 – 3,200 m altitudinal range.

§ sect. Gynotrichae - only one spp., S. loesneriana Busc, endemic to 2,800 – 3,000 m altitudinal range in Piura and Cajamarca in Peru.

§ sect. Laevigatae - 9 spp., 8 from Venezuela to Peru and one endemic to Bolivia.

§ sect. Parviflorae - 10 spp., Colombia (6 endemics) to Peru (1 endemic) and Bolivia (one endemic), mainly restricted.

§ sect. Pulverulentae - 9 spp., Venezuela, Colombia (5 endemic) and Ecuador (2 endemic), coastal lowland to highlands.

§ sect. Macrophyllae - 13 spp., from Venezuela (one endemic) to C Peru (three Colombian endemics), and one endemic to Bolivia.

§ sect. Lanatae - three spp., Colombia and Peru one endemic each, and S. bullosa Wawra from Colombia to Peru; at the volcano El Galeras, above Pasto, individuals of this spp. are found up to 3,600 m, probably the highest limit of tolerance of members of Saurauia.

The highest altitude which members of South American Saurauia tolerate is 3,600 m, represented by S. bullosa, and the lower limit is almost at sea level, represented by S. mexiae Killip ex Soejarto, S. parviflora Triana & Planch., S. pseudoleucocarpa Buscal. and S. yasicae Loes.; the geographical range of the South American species is limited to the Andean mountain system, from Venezuela in the north to Bolivia in the south. No species are represented in the ‘puna’ vegetation of Peru and Bolivia. A strong species concentration is located near the Colombo- Ecuadorian frontier (the Narino-Putumayo region), just before the Andes splits into the three Colombian Cordillera systems, i.e., the Occidental, Central, and Oriental.

SARRACENIACEAE

§ CARNIVOROUS (Brocchnia - Catopsis - Paepalanthus - Drosera - Heliamphora - Philcoxia - Genlisea - Utricularia - Pinguincula)

Genera/species 3/30 Distribution E U.S.A., S Canada, the Guiana Shield. Habit bisexual, perennial rhizomatous herbs. Carnivorous. Helophytes.

At Guiana Shield mountains, the uneven plateau of Cerro Duida is heavily inclined, rising north to south from around 1,300–1,400 m to a maximum of 2,358 m. But the highest point of the massif, at 2,832 m, is found on Cerro Marahuaca, this being the second-highest mountain of the entire Guiana Shield (after Mount Neblina).

SYSTEMATIC outsiders Darlingtonia (1; Oregon, N California) and Sarracenia (9–10; E U.S.A., S and SE Canada).

1. Heliamphora Benth. Pitcher herbs, with a double keel; rhizomatous, most acaulescent. H. nutans and H. minor has erect stems and can attain a dendroid habit; pitchers long with short petioles and are borne in a rosette, shaped like urns and trumpets; flowers are quite large, with free rather petal-like sepals, free petals, and usually numerous stamens. 23 spp., live in acidic, boggy habitats, in two geographical groups:

§ Western Range (6) ‣ two spp. only inland Venezuela (H. macdonaldae Gleason and H. tatei Gleason, restricted of Cerro Duida, Huachamakari and Marahuaka), and four in Mount Neblina (three only in this tepui), but only three in Brazil:

§ H. ceracea Nz., Wist., Grant., Riv., Fleisc. & McPherson - occur at 1,900 m altitudinal range, known only from several small populations on slopes of Mount Neblina in Brazil, but may have wider distribution; if not, it would be the only known species not found in Venezuela.

§ H. hispida Nerz & Wistuba - occur at 1,800–2,994 m altitudinal range, found in southern portion of Mount Neblina, on and around Mount Neblina and Pico 31 de Março (including area around the Titirico River); additional populations may grow in the largely unexplored Cañón Grande; plants near the summit of Mount Neblina (2,994 m) represent the upper altitudinal limit for the genus.

§ H. neblinae Maguire - occur at 860–2,200 m altitudinal range, known with certainty from southern portion of Mount Neblina, where it grows on and around summit up to Pirapacu Range in Brazilian side; herbarium material suggests it is also present in the northern parts, though it has not been observed in the extreme north; the range of H. neblinae extends to the two northern tepuis (Aracamuni and Avispa, the last represent the lower altitudinal limit for the genus, with populations growing at 860 m; the pitchers of H. neblinae are some of the largest in the genus, occasionally exceeding 50 cm.

§ H. parva (Maguire) S.McPherson, A.Fleischm., Wistuba & Nerz - occur at 1,750–2,200m altitudinal range, recorded from NW part of Mount Neblina at 2,000–2,200m in Venezuela side, and no other Heliamphora species is known from this area, though the largest known stands grow in the northwestern part of the massif, at lower elevations of 1,750–1,850 m; it is unknown whether the range of this species extends into the central valleys of Mount Neblina.

Mount Neblina is located in the extreme north of Amazonas and is dominated by the highly fragmented complex with many plateaus averaging 2,000–2,400 m in elevation, and is the Brazil's highest peaks (2994 m), known as Pico Phelps in Venezuela; a giant valley, Cañón Grande, runs southwest to northeast through the middle of Mount Neblina; the smaller outcrops of Cerro Aracamuni and Cerro Avispa, both reaching approximately 1,600 m, lie to the north of this complex. The name Mount Neblina not includes Cerro Aracamuni and Cerro Avispa.

Hibrids are H. ceracea × H. hispida, recorded from Brazilian border area on flanks of Mount Neblina; H. neblinae × H. parva - found in the north of Mount Neblina; complex backcrosses have been recorded; Undetermined hybrids - putative complex swarms involving H. ceracea, H. hispida, H. neblinae and H. parva are found in the southern part of Mount Neblina.

§ Eastern Range (17) ‣ 15 spp. are endemic to Venezuela (Gran Sabana, Los Testigos and Ptari Tepui two endemics each, one endemic to Auyan Massif, 5 in Middle East Tepuis), mainly encompassed by Bolívar state, Venezuela; the five remaining is from East Tepuis, and two ranges up to Brazil and Guiana:

§ H. glabra (Maguire) Nerz, Wistuba & Hoogenstrijd is found in Mount Roraima (in small, scattered populations in northern portion of summit plateau (including around tripoint). Growing at roughly 2750 m, these stunted plants represent the upper altitudinal limit of the species; lowest altitude populations (1,200 m) grow on slopes off the mountain's northern flank. Distribution may similarly extend eastwards from Mount Roraima, but this requires confirmation), in Serra do Sol (summit supports the largest known populations of this species, unusually consisting of predominantly green-pitchered plants; only known Venezuelan locality for this species is at the southern end of the plateau) and Wei Assipu Tepui (on summit. Likely to grow around base also, but this requires confirmation).

§ H. nutans Benth, the first Heliamphora to be described and is the best known species, found in Mount Roraima (mostly around edges of mountain and on upper tepui cliffs; summit plateau hosts highest-growing population of this species, at 2,700 m; originally discovered in ‘El Dorado Swamp’ off Mount Roraima's southern flank, but this population has not been relocated since its discovery and was possibly destroyed by fires); and also in Wei Assipu Tepui (on upper tepui cliffs).

§ hybrids H. nutans x H. glabra also occur in Mount Roraima (in northern part of summit plateau in apparently sterile clumps; complex backcrosses reported) and Wei Assipu Tepui (complex backcrosses reported).

East tepuis, this tepuis, known also Roraima–Ilú range, stretches in a northwesterly direction from the tripoint of Brazil, Guyana, and Venezuela, closely following the Guyana–Venezuela border, with an isolated double-peaked plateau (Serra do Sol or Uei Tepui) to the south. Moving in a northwesterly direction from Serra do Sol (2,150 m), the major summits of this chain are Mount Roraima (2,810 m), Kukenán Tepui (2,650 m), Yuruaní Tepui (2,400 m), Wadakapiapué Tepui (2,000 m), Karaurín Tepui (2500 m), Ilú Tepui (2,700 m), and Tramen Tepui; with the exception of the tiny Wadakapiapué Tepui, all of these peaks are known to support Heliamphora.

H. uncinata Nerz, Wistuba & A. Fleischm. is only known from a single collection at the type location, a narrow canyon on Amurí-tepui (the western sector of Acopán-tepui), where it grows mainly on the vertical sandstone cliff surface in shaded conditions in humus pockets and cracks at c. 1,850 m; the only other species of Heliamphora known so far that preferably grows on vertical, wet sandstone walls is H. exappendiculata (Maguire & Steyerm.) Nerz & Wistuba.

They are exquisitely constructed pitfalls that entrap insects lured to the mouth of the pitcher by nectar-secreting glands and glistening surfaces. Downward-pointing hairs in the throat of the pitcher prevent the insect’s escape, and the exhausted prey slide down the slippery throat and fall into the liquid in the pitcher, where they are either digested by enzymes secreted by glands in the pitcher or eaten by the animals living in the pitcher, their remains being excreted into the liquid.

CLETHRACEAE

§ FULLY CHLOROTROPHIC

Genera/species 2/99 Distribution E and SE U.S.A., Mexico, Caribbean, Central America, tropical South America, Madeira, E Asia to Korean Peninsula and Japan, Taiwan and Hainan in China, SE Asia, Malesia, New Guinea. Habit bisexual (rarely functionally gynodioecious), evergreen or deciduous trees or shrubs.

SYSTEMATICS both genera occur in South America.

1. Clethra L. Trees and shrubs with entire to serrate leaves. 86 spp., most diverse in the tropical humid montane habitats, reaching their upper altitudinal limit at about 3,800 m in the Andes; two sects:

§ sect. Cuellaria ‣ 57-62 spp. in two subsections:

§ subsect. Cuellaria ‣ Central and South America (22 in continent), only two spp. in Brazil, C. scabra Pers in C & S South America, and C. uleana Sleumer, endemic to mountains in S region.

§ subsect. Pseudocuellaria ‣ a single sp., C. arborea Aiton, from Madeira.

§ sect. Clethra ‣ 24-29 spp., 22 in China (inc. Taiwan), Korean Peninsula, Japan, SE Asia, Malesia to New Guinea, and two in North America.

2. Purdiaea Planch. Small trees or shrubs with sessile entire leaves. 13 spp., 11 endemic to Cuba, P. belizensis (A.C. Sm. & Standl.) J.L. Thomas in Central America, and P. nutans Planch. from Venezuela to Peru.

CYRILLACEAE

§ FULLY CHLOROTROPHIC

Genera/species 2/2 Distribution coastal plains in SE U.S.A., Caribbean, Central America, N South America. Habit bisexual, evergreen or deciduous shrubs or small trees.

SYSTEMATIC the family comprises two genera, with Cliftonia monophylla (Lam.) Britton ex Sarg. endemic to SE U.S.A. a outsider; Cyrillaceae is placed in the Ericales as sister to Ericaceae, and also closely related to Clatraceae. The Cyrillaceae have similarities with Ericaceae and Clethraceae. It differs from them by hidehiseent fruit, number of carpels and locules (3 in Clethra L.), presence of nectariferous dise and few ovules (numerous in Clethra).

1. Cyrilla L. Small shrub to canopy tree, sometimes with woody rhizomes. 10 spp., 9 restricted of Caribbean (almost all endemic to Cuba), and C. racemiflora L., from SE U.S.A., Greater Antilles, Mexico (only known in Mexico from one small isolated population in Oaxaca), Belize, Honduras, Nicaragua, Panamá, E Colombia to Suriname, and N Brazil in Amazonas and Roraima states (very common in Mount Aracá in Brazil, and along N Rio Negro, Nhamunda and Trombetas rivers); the species also grows to be a large tree in the cloud forest of Puerto Rico but it is always a much smaller plant in savanna habitats of Central and South America and the U.S.A.

J. L. Thomas (1960) acknowledges that C. racemiflora is polymorphic and should be recognized as a single variable species, on the basis of local variation in leaf and inflorescence size. Yesilyurt (2009) pointed out that during studies of the flora of the Guianas that specimens showed considerable amount of variation in size and shape and even texture in all the features of the leaves, flowers and fruits.

ERICACEAE

§ MYCOHETEROTROPHICS (Arachnitis – TRIURIDACEAE – BURMANNIACEAE – ORCHIDACEAE – Voyria - Voyriella - Monotropa)

Genera/species 121/4,100 Distribution cosmopolitan, although few species in tropical lowland regions, with their largest diversity in Himalaya to SW China, New Guinea, South Africa, Australia, and New Zealand. Habit usually bisexual (rarely monoecious or dioecious), usually evergreen (sometimes deciduous) shrubs or suffrutices (rarely trees, lianas or perennial herbs). Some genera consist of achlorophyllous and mycotrophic plants. Numerous species are xerophytic. Some species are helophytes. A starchy lignotuber is present especially in many species in mediterranean climates. They are herbs, terrestrial, epiphytic (erect or pendent), or, rarely, saxicolous shrubs and subshrubs 1-2 m tall, lianas, cushion plants (in Dracophylum, Disterigma empetrifolium and Rhododendron saxifragoides), prostrate mat-forming chamaephytes, and trees to 20 m tall. The herbs are either achlorophyllous and mycotrophic or green and autotrophic.

DIVERSITY NUMBER c. 220 spp. occur in North America in 40 genera; 44 (40 endemic) in Caribbean in 9 genera; 60 in Mexico and Central America in 20 genera; 66 (64 endemic at, Gaultheria setulosa shared with Guiana Shield and ropical Andes, another reaching into Paraguay and Uruguay) in Brazilian highlands in 3 genera; 10 (all endemic at) in temperate Andes in 2 genera; 71 in Guiana Shield in 18 genera; and 586 (519-522 endemic at) in tropical Andes in 30 genera;

46 genera and about 800 species of Ericaceae native to the Neotropics; in South America highly centered in northern Andes, with most species in Colombia and Ecuador, where approximately 50% (ca. 398 out of ca. 800) of the species occur; 32 genera and 752 spp. are Neotropical endemics.

ENVIRONMENT, HABITAT, AND HABIT Overall, Ericaceae are the sixth largest epiphytic family of vascular plants in Neotropics, following the Orchidaceae, Bromeliaceae, Araceae, Piperaceae, and Gesneriaceae. At least 30 genera and c. 340 species of neotropical Ericaceae, equaling about 65% of the genera and 45% of the sp., occur as epiphytes (and this is probably a low estimate); in the Neotropics Ericaceae grows continuously throughout the year in na seasonal habitats, producing flushes of intensely red-pigmented new leaves; this make the family is very easy to spot in the field; the great majority of the sp. have flowers with fused petals.

It should be noted that, with only two known exceptions, Gaultheria erecta Vent. (the Costa Rican populations only) and occasional individuals of Lyonia octandra (Sw.) Griseb. (a Jamaican endemic), all superior-ovaried genera of neotropical Ericaceae are terrestrial.

In the Neotropics the vertical relief often spans 0-5,000 m, giving rise to a mosaic of habitats and ecological niches. In these areas Ericaceae have evolved different life-forms and may occupy many different habitats. They are herbs, terrestrial, epiphytic (erect or pendent), or, rarely, saxicolous shrubs and subshrubs 1-2 m tall, lianas, cushion plants, prostrate mat-forming chamaephytes, and trees to 20 m tall; the herbs are either achlorophyllous and mycotrophic or green and autotrophic. They can also be quite plastic.

A few species are found as epiphytes in mangrove swamps; only two neotropical species, Cavendishia laurifolia and Comarostaphylis polifolia, have been recorded from limestone habitats; in the E Venezuela, Notopora schomburgkii Hook.f. and Vaccinium puberulum Klotzsch are typical elements in the sandy savanna shrublands; Gaultheria myrsinoides Kunth forms carpets or creeping mats over many hectares in the mountains of Mexico and Guatemala, as well as in the superparamo at Nevado del Cocuy, Colombia.

In montane regions of the Neotropics, several weedy species of Ericaceae are frequently found as pioneers following volcanic activity or recent landslides or as part of the edge community around mature forests. People probably have also had great influence on the distribution of these species by creating disturbed areas, especially along the forest edge and steep slopes of clearings after periods of logging or road building. A particularly weedy species in the high-elevation paramo, Gaultheria myrsinoides, is resistant to trampling by cattle and is a successional species in heavily grazed areas. This species invades paramo in places where the vegetation is low and open, and it persists after the vegetation recovers. Fire does not affect it directly, but actually helps spread it indirectly because it opens the vegetation (Pels & Verweij, 1992). All Ericaceae, even rhizomatous species, have a tap root initially that may develop to a depth of 2-3 m, as well as a shallow root system that spreads diffusely just below the soil surface, where there is usually abundant moisture. A few species are rhizomatous and/or mat forming. Numerous species develop bark-covered lignotubers and/ or burls that can vary in size from several centimeters to more than 1 meter in diameter.

BIOGEOGRAPHICAL Floristic assemblages at the generic and specific levels further characterize each of the five neotropical biogeographical regions. For example, the Guiana Shield, by three endemic genera Ledothamnus (7), Mycerinus (3), and Notopora (5), as well as by Vaccinium puberulum, Vaccinium euryanthum A. C. Sm., and Thibaudia nutans Klotzsch; and the SE Brazilian, by the genera Agarista (24 endemic) and Gaylussacia (36 endemic).

The five biogeographical regions, however, are structurally, vegetatively, and floristically more complex than the generalizations mentioned above imply. Each region itself may be divided into several smaller, more local floristic units of high ericad speciation, which in many cases coincide with the finer structural features as proposed by Simpson (1975) and modified by Berry (1985). Although these smaller floristic units are not as obvious as the larger biogeographical regions, they are nevertheless definable by their own clusters of genera and/or species.

The following six units are examples.

1. A southern Mesoamerican unit: Costa Rica to Panamá, consisting of low-elevation, continental divide, premontane cloud forests. It is characterized by four endemic genera, Anthopteropsis (1), Didonica (4), Lateropora (3), and Utleya (1), as well as Cavendishia series Lactiviscidae (9). Few taxa span the Middle American land bridge that arose during the Pliocene, and 49% of the species (ca. 57 of 116) are endemic.

2. A northern Andean unit: the three cordilleras of Colombia, the northwestern and eastern slopes of Ecuador, and the northeastern slopes of Peru, consisting of wet, mid- to highelevation (1,000-2,999 m) montane cloud forests. It is characterized by the genera Cavendishia, Ceratostema, Macleania, Orthaea, Psammisia, Sphyrospermum, Themistoclesia, and Thibaudia, each with 20 or more species.

3. A high Andean paramo unit: between (3,000-) 3,500-4,500 m. It is characterized by the genera Plutarchia (9 endemics to Colombia) and Gaultheria (16, essentially endemic to paramo), and Vaccinium floribundum, Gaultheria myrsinoides and Disterigma empetrifolium Kunth.

4. A south-central Ecuador-north-central Peruvian unit: coincident with the Amotape-Huancabamba Zone, consisting of a more seasonal scrub forest. It is characterized by Gaultheria reticulata Kunth, G. tomentosa Kunth, G. lanigera Hook.f. var. lanigera, and several closely related species of Oreanthes and Ceratostema.

5. A south-central Peru-northern Bolivian unit: high-elevation cloud forests. It is characterized by the following endemic genera: Demosthenesia (9), Siphonandra (2-3), Pellegrinnia (4), Rusbya (1), and Polyclita (1).

6. A ‘Choco’ unit: Pacific coastal South America from extreme SE Panamá through Colombia to north-central Ecuador, and consisting of low-elevation (mostly below 1,000 m) rain or cloud forest. It is characterized by Anthopterus wardii Ball, Macleania smithiana Luteyn, 16 species of Cavendishia, and 10 species of Psammisia. This unit was first noted by Smith (1946) and subsequently promoted as a distinct phytogeographical province by Gentry (1982a).

WIDELY DISTRIBUTED TAXA IN NEOTROPICAL VACCINIEAE Most species of neotropical Vaccinieae are narrow endemics, growing in specific habitats or altitudinal belts; the narrowly endemic species are generally distinctive, seem to be more specialized, and have narrower morphological variation than do more widely distributed species. Many are found only in a single mountain range, and a few are endemic to single peaks. But in addition to numerous narrow endemics, each large genus of Vaccinieae also has one or two widely distributed species, e.g., Cavendishia bracteata, Macleania rupestris, Psammisia guianensis, Satyria panurensis (Benth. ex Meisn.) Hook. f. ex Nied., Sphyrospermum cordifolium Benth., Thibaudia floribunda, and Vaccinium floribundum.

The following Ericaceae were also collected at Mount Aracá, all collected in Brazil in 1993: Bejaria sprucei Meissner, Cavendishia callista Donn.-Sm., Satyria panurensis (Benth.) Bentham & Hooker, Sphyrospermum cordifolium, Thibaudia formosa (Yd.) Hoerold, Thibaudia nutans Klotzsch. ex Mansfeld and Vaccinium puberulum Klotzsch. ex Meissner var. puberulum.

SYSTEMATICS eight subfamilies, Enkianthoideae (1/17, Japan, S China, N Burma, Indochina), Arbutoideae (3–5/c 85, temperate regions on the Northern Hemisphere southwards to mountains in Central America), Cassiopoideae (1/12, arctic-alpine regions on the Northern Hemisphere, Himalayas), Harrimanelloideae (1/2, arctic-alpine regions in N Europe to N Ural, Japan, the Kurile Islands, Kamchatka, Alaska, arctic Canada to Quebec and NE U.S.A.) do not occur in South America.

1. SUBFAMILY MONOTROPOIDEAE (14/55–60) ‣ tribes Pyroleae (4/c. 42, temperate regions on the Northern Hemisphere to mountains on Taiwan (China) and northern Sumatra) and Pterosporeae (2/2, S Canada, W and NE U.S.A., N Mexico) does not occur in South America; among Monotropeae, outsiders are Allotropa (1; W U.S.A.), Cheilotheca (3; Assam to W Malesia), Hemitomes (1; W U.S.A.), Monotropastrum (2; E Asia, Sumatra), Monotropsis (1; SE U.S.A.), Pityopus (1; W U.S.A.), Pleuricospora (1; SW Canada, W U.S.A.).

1. Monotropa L. Mycoheterotrophic herbs associated at Quercus sp. Three spp., M. hypopitys L. from Northern Hemisphere up to Central America, one endemic to Florida shrubby region, and M. uniflora L. from Northern Hemisphere up to NW Colombia, in native Quercus forest and introduced pine plantations in the W Cordillera.

2. SUBFAMILY ERICOIDEAE (18/c 1.770) ‣ 5 tribes, Rodoeae (1/850) and Ericeae (3/860, Europe, Africa) not in South America.

2.1 ERICOIDEAE ▸ TRIBE PHYLLODOCEAE (7/42) ‣ outsiders Elliottia (4; Alaska, Canada, U.S.A.), Kalmia (10; Canada, U.S.A., Cuba, arctic-alpine regions on the Northern Hemisphere), Epigaea (3; Türkiye, Caucasus; Japan; U.S.A.), Rhodothamnus (2; Alps, Türkiye), Kalmiopsis (1; Oregon), Phyllodoce (7; cold-temperate and arctic-alpine regions on the Northern Hemisphere).

2. Bejaria L. Shrubs to trees, corolla tubular. 15 spp., U.S.A. and Cuba one endemic each, and 13 in N South America from Colombia to Bolivia up Guyana, B. aestuans Mutis ex L. up to Mexico, also northernmost point of Brazil (4, all in Roraima state, one reaching into Amazonas state, none endemics).

2.2 ERICOIDEAE ▸ TRIBE EMPETREAE (3/6) ‣ outsiders Ceratiola (1; SE U.S.A.) and Corema (2; Portugal, W Spain, the Azores, E Canada, NE U.S.A.).

3. Empetrum L. Prostrate shrubs. 4 spp., three in northern temperate, and one, E. rubrum Vahl ex Willd., in extreme S South America, Faklands Is., Tristan de Cunha.

2.3 ERICOIDEAE ▸ TRIBE BRYANTHEAE (2/8) ‣ outsider is Bryanthus (1; Kamchatka, Japan). The discovery of the clade composed of Bryanthus and Ledothamnus also reveals an interesting biogeographic disjunction within the subfamily Ericoideae. Bryanthus is found only in Japan and Kamchatka, whereas Ledothamnus is associated with tepuis on the Guyana-Venezuala border and adjacent Brazil. To the extent that we have phylogenetic resolution in the Ericaceae, this disjunction is so far unique in the family, and especially in the subfamily Ericoideae; most intercontinental disjunctions here appear to be Asian-North American. A NE Asian- South American disjunction has been reported in the family Lardizabalaceae.

4. Ledothamnus Meisn. Terrestrial ericoid shrubs and subshrubs of small stature, of open savannas and rocky mountains tepuis in Venezuela. 7 spp., six species are each restricted to one or a few tepuis, endemics to the Guiana Shield of Venezuela, and L. guyanensis Meisn reaches up to N Brazil, with one collection in Roraima state at 1,000 – 2,800 m elevation range.

3. SUBFAMILY VACCINIOIDEAE (c 50/1,300–1,400) ‣ five tribes, Andromedeae (2/3, Northern Hemisphere) and Oxydendreae (1/1, North America) do not occurs in South America. 659 spp. in South America.

3.1 VACCINIOIDEAE ▸ TRIBE VACCINIEAE (33/1220-1250) ‣ outsiders are only eight, Didonica (4; Costa Rica, Panamá), Symphysia (15; Central America, Caribbean), Gonocalyx (9; Central America) and Anthopteropsis (1; central Panamá) in Central America and Caribbean; Agapetes (109; tropical Asia from India to New Guinea), Dimorphanthera (75–80; Malesia, with their highest diversity on New Guinea), Paphia (c 20; E New Guinea, E Queensland, New Caledonia, Fiji) in Paleotropics, and Oxycoccus (4; temperate and arctic-alpine regions on the Northern Hemisphere) in Northern Hemisphere. 34 genera, 25 in South America (only two of then also outside Neotropics, Vaccinium and Gaylussacia, the unique mainly terrestrial genera in tribe), with 480 spp. in this region. Only seven genera in Brazil, the bulk of spp. are Gaylussacia. The six remaining genera has only 16 spp. of northern Amazon rainforest, few collected.

5. Anthopterus Hook. Shrubs or small trees, rarely epiphytes. 12 spp. from Costa Rica to NE Peru, 11 in South America, mainly in Ecuador (4 endemics) and Colombia (2 endemics).

6. Cavendishia Lindley. Epiphytic (obligate or facultative, up to only a few centimeters to several meters long) or terrestrial shrubs, wiry, or scadent subshrubs, erect shrubs, and small trees to 7 meters tall. 114 spp., Mexico to Bolivia, E to the Guianas, Guiana Shield, NW Brazil (2, none endemics, from Amazonas state); 43 in Central America/Mexico, 83 in South America, 48 endemics to Colombia.

7. Ceratostema Juss. Terrestrial or epiphytic, raraly scadent shrubs. 36 spp., 35 from S Colombia to N Peru (30 endemics to E Ecuador, 3 endemics to Peru), and C. glandulifera Maguire, Steyerm. & Luteyn endemic to Guyana.

8. Demosthenesia A.C.Sm. Usually epiphytic shrubs, lignotubers frequent. 12 spp. C Peru (8 endemics) to N Bolivia (2 endemics).

9. Diogenesia Sleumer. Usually epiphityc shrubs; branches slender, often pendent, rarely climbing. 14 spp., from W Venezuela to N Bolivia.

10. Disterigma (Klotzsch) Nied. Terrestrail or epiphytic shrubs, rarely cushions. 36 spp., highly centered in Ecuador and Colombia, reaching up to Guatemala in northern, Bolivia in south, and W Guyana in east (34 in South America); D. humboldtii (Klotzsch) Nied. occur in Mount Neblina in Amazonas state, N Brazil.

D. empetrifolium (Kunth) Drude, a terrestrial, normally erect, somewhat wiry small shrub in subpáramo, may become a stoloniferous subshrub in paramo areas, a cushion plant in highly exposed paramo, or a tiny subshrub in the rocky crevices of superpairamo. This genus was officially collected in Brazil in 2015, through a project called ‘Montanhas da Amazônia’, on Mount Caburai, Mount Aracá, and Mount Neblina, in the Brazilian portion of the Guiana Shield.

11. Gaylussacia Kunth. Shrubs or subshrubs, estrictely terrestrial. 56 spp. in three section:

§ sect. Decamerium ‣ 6 spp. all in U.S.A., one reaching to Canada.

§ sect. Gaylussacia ‣ three spp. in North America and 46 in South America, G. buxifolia Kunth in Colombia and Venezuela, G. cardenasii A.C. Sm. in Bolivia and Argentina, G. loxensis Sleumer in Ecuador and Peru, G. brasiliensis (Spreng.) Meisn. in Brazil and adjacent Cono Sur, and remaining 42 endemic to Brazil, highly centered in mountains of Espinhaço Range and montane forests of Rio de Janeiro and Espírito Santo, one reported in Pará state; all of these species are important ecologically as a food source for wildlife and a componente of forests, shrub-lands, and bogs.

§ sect. Vitis-idaea ‣ only G. brachycera (Michx.) A. Gray from E U.S.A., possibly a Vaccinium species.

12. Gonocalyx Planch. & Linden. Small epiphytic, rarely terrestrial shrubs. 11 spp., 4 spp. in the Antilles, 6 in Costa Rica & Panamá, and G. pulcher Planch. & Linden endemic to N Colombia.

13. Macleania W. J. Hook. Epiphytic or terrestrial shrubs. 33 spp., from S Mexico to Peru and Bolivia (31 in South America), high diverse in W Ecuador (13 endemics).

14. Mycerinus A.C.Sm. Terrestrial or epiphytic shrubs. Three spp., endemic to the Guiana Shield of Venezuela, 1,300 – 2,700 m.

15. Notopora Hook.f. Terrestrial shrubs. 5 spp. endemic to the Guiana Shield of Venezuela, adjacent Guyana, with N. schomburgkii Hook. f. in Mount Caburai in N Roraima state, Brazil, at 400-2,500 m alt.

16. Oreanthes Benth. Terrestrial or epiphytic shrubs. 7 spp., 6 are endemic to Ecuador and there are only two collections from O. buxifolius Benth. in N Peru.

17. Orthaea Klotzsch. Epiphytic shrubs. 37 spp. in Mexico to Bolivia (34 in South America), Guyana (5) and Trinidad.

18. Pellegrinia Sleumer. Usually epiphytic. 4 spp. endemics to Peru.

19. Plutarchia A.C.Sm. Terrestrial shrubs; inflorescence a fascicle. 11 spp. from Colombia (9 endemics) and N Ecuador (one endemic), P. angulata A.C. Sm. in both countries.

20. Polyclita A.C.Sm. Terrestrial or epiphytic shrubs. Only one sp., P. turbinata (Kuntze) A.C. Sm., endemic to Bolivia.

21. Psammisia Klotzsch. Epiphytic or terrestrail shrubs, sometimes lianescent, flowers with the greatest diversity of corolla shapes and colors of any other genus of neotropical Vaccinieae (e.g., tubular, obconic, urceolate, turbinate and depressed, hemispheric; yellow, magenta, vermilion, dark wine, red, white, green; unicolor, multicolor). 67 spp. from Costa Rica to Bolivia, E to French Guiana, Trinidad (64 in South America). 4 spp. in Brazil (Roraima, Amazonas and Pará state, none endemics). P. guianensis Klotzsch has a distinctive arcuate or C-shaped distribution around the Amazon rainforest, growing in white-sand savannas in the northern parts and continue along the western part at the Andes-Amazonia rainforest ecotone. Polytomic with Macleania, Ceratostema and some Thibaudia.

22. Rusbya Britton. Epiphytic shrubs, flower solitary. Only one sp., R. taxifolia Britton, from N Bolivia.

23. Satyria Klotzsch. Epiphytic or terrestrial shrubs. 26 spp. from S Mexico to N Bolivia, Guianas, NW Brazil; 22 occurs in South America, two in Brazil: S. cerander (Dunal) A. C. Sin., the only neotropical species of Ericaceae that may be considered Amazonian; species of rare occurrence, found in the lowlands of French Guiana and adjacent Brazil in Amapá state; and S. panurensis (Benth. ex Meisn.) Benth. & Hook.f. ex Nied., with a distinctive arcuate or C-shaped distribution around the Amazon rainforest, growing in white-sand savannas in the northern parts (in Brazil in Roraima and Amazonas states) and continue along the western part at the Andes/Amazonia rainforest ecotone, also disjunct along the Caribbean lowland slopes of western Panamá to southern Mexico; it’s only epiphytic Ericaceae in Brazil.

24. Semiramisia Klotzsch. Terrestrail or epiphytic shrubs. 4 spp., Venezuela (one endemic), Colombia (two endemics), and one from Colombia to Peru.

25. Siphonandra Klotzsch. Terrestrial or epiphytic. 6 spp. Central Peru (3 endemics) to N Bolivia (2 endemics), one in both countries.

26. Sphyrospermum Poepp. & Endl. Epiphytic or terrestrial shrubs with pendent branches. 22 spp., S Mexico to N Bolivia, Guianas, Haiti and Trinidad, 21 in South America, only S. buxifolium Poepp. & Endl. in Brazil (Roraima and Amazonas state), non endemic.

27. Themistoclesia Klotzsch. Often epiphytic shrubs. 36 spp. from Costa Rica to N Bolivia and Venezuela, 33 in South America, 15 endemics to Colombia,.

28. Thibaudia Ruiz & Pav. Epiphytic or terrestrial shrubs. 72 spp., one in Costa Rica, e 71 in N South America, three up to N Brazil (Amazonas and Roraima states), none endemics.

29. Vaccinium L. Shrubs, rarely trees, lianas, or with woody rhizomes, estrictely terrestrial. c. 140 spp., worldwide, except Australia, 83 in New World, Caribbean, Mexico to Argentina, east to Guyana, Guiana Shield, NW Brazil (2, V. puberulum Klotzsch ex Meisn. in Roraima and Amazonas, also in Venezuela and Guianas, and the very narrow endemic V. pipolyi Luteyn, endemic to Mount Aracá (a rare plant in Brazil, by Plantas Raras do Brasil’s book), 26 in South America.

V. floribundum Kunth seems to be the most used native species, especially in Ecuador and Colombia, where the fruits are made into jams, drinks, and occasionally pies; it is very abundant throughout the Andes and could probably be developed by cultivation into a more valuable fruit crop; the wood of various neotropical Ericaceae is used locally for firewood or charcoal; fruits from a form of V. puberulum caused a considerable lowering of blood pressure, followed by nausea and vomiting in a number of botanists in Venezuela - symptoms similar to some of those reported for temperate-region species of Rhododendron and Kalmia and presumably caused by the toxic compound andromedotoxin. However, laboratory analyses were unable to confirm the presence of andromedotoxin in the Venezuelan collection.

3.2 VACCINIOIDEAE ▸ TRIBE LYOINEAE (4/77) ‣ outsiders Craibiodendron (5; S China, SE Asia), Lyonia (c 35; E and SE Asia, E U.S.A., Mexico, Caribbean) and Pieris (7; Himalayas to E Asia and E Siberia, SE U.S.A., Cuba).

30. Agarista D.Don. Shrubs. 32 spp., A. salicifolia G.Don in Africa, Madagascar, Raunion and Mascarenes, A. populifolia (Lam.) Judd in U.S.A. in Florida and South Carolina, three from Mexico to Honduras, and remaining 27 in South America: 5 scattered from Venezuela to Cono Sur, 19 endemics to Brazil, A. duckei (Huber) Judd in northern Amazon rainforest up to Pará state, and two from Brazil to Cono Sur.

3.3 VACCINIOIDEAE ▸ TRIBE GAULTHERIEAE (4/150) ‣ outsiders Chamaedaphne (1; temperate regions on the Northern Hemisphere), Eubotrys (2; U.S.A.), Leucothoe (5; Himalayas, Japan, North America).

31. Gaultheria Kalm ex L. (inc. Pernettya, Tepuia) Shrubs, sometimes decumbent. 292 spp., Circum-Pacific in Northern & Southern Hemispheres/Mexico-Argentina, east to Brazil; 68 in New World, 57 in South America, 13 in Brazil, 6 endemics (also a hybrid endemic); G. myrsinoides Kunth by Middleton is frequently mentioned as having intoxicating properties or to be poisonous, although this is not well documented; G. erecta Vent. occur from Mexico to Brazil, and is a large shrub or small tree.

4. SUBFAMILY STYPHELIOIDEAE (superior ovaries, c 35/510-525) ‣ 7 tribes, with outsiders only in Australia, Tasmania (24 endemics), New Caledonia and New Guinea except Leptecophylla up to the Hawaii, Leucopogon and Trochocarpa up to Malesia region.

32. Lebetanthus Endl. Scrambling shrubs, corolla suburceolate. Only one sp., L. myrsinites (Lam.) Dusén, from Patagonia and Terra do Fogo.

50. ICACINALES

A SINGLE FAMILY, PRESENT IN SOUTH AMERICA.

ICACINACEAE

§ FULLY CHLOROTROPHIC

Genera/spp. 24/160 Distribution Madagascar, W and S India, Sri Lanka, SE Asia, S China (inc. Taiwan), Japan, islands of the W Pacific, Malesia, New Guinea, Melanesia, N Queensland; a few spp. in temperate regions. Central America, Caribbean, tropical South America; Habit usually bisexual, evergreen trees, shrubs or lianas with non-axillary branch tendrils.

Use timber, medicinal plants (seed oil from Sarcostigma), starch sources (tubers of Casimirella); in America only one sp. are economically important Casimirella ampla (Miers) R.A. Howard produces a tuber weighing 5-20 kilos, rich in starch that is edible after washing out the bitter-tasting compounds.

SYSTEMATIC outsiders Cassinopsis (6; S and E Africa, Madagascar); Nothapodytes (11; E Asia, tropical Asia), Mappia (5; S Mexico, Central America, Caribbean); Alsodeiopsis (11; tropical Africa), Merrilliodendron (1; Philippines, W Pacific), Lavigeria (1; Cameroon, Gabon, Congo), Icacina (5; tropical Africa), Mappianthus (1; S China), Iodes (28; tropical regions in the Old World), Polyporandra (1; E Malesia to New Guinea, Melanesia); Desmostachys (6; tropical Africa, Madagascar), Natsiatum (1; E Himalayas to SE Asia), Hosiea (2; W and C China, Japan), Rhyticaryum (12; E Malesia to New Guinea and W Pacific islands), Sarcostigma (2; India to SE Asia and Malesia), Miquelia (7; tropical Asia), Phytocrene (11; SE Asia, Malesia), Stachyanthus (4; tropical Africa), Pyrenacantha (c 35; tropical regions in the Old World), Natsiatopsis (1; Burma, S Yunnan), Sleumeria (1; N Borneo).

1. Casimirella Hassl. 7 spp. in S America to Paraguay, with a diversity center in SW Brazil (4, 3 endemics) and Paraguay; C. ampla (Miers) R.A. Howard very widspread.

2. Leretia Vell. Vine to scandent shrub. Only one sp., L. cordata Vell., in Costa Rica to Brazil (disjunct among this country, Amazon rainforest and Atlantic Forests) and Bolivia.

3. Pleurisanthes Baill. Woody vines to climbing shrubs, stems slightly quadrangulate. 7 spp. rainforests of Venezuela (one endemic), Guyana, Suriname, French Guiana, Ecuador, Peru, and Brazil (high diversity, 6, Amazon rainforest and Atlantic Forest, with two endemics).

51. METTENIUSALES

A SINGLE FAMILY, PRESENT IN SOUTH AMERICA.

METTENIUSACEAE

§ FULLY CHLOROTROPHIC

Genera/species 10/55 Distribution pantropical Habit bisexual, evergreen trees. Most New World genera are widely distributed in the region, but Oecopetalum is confined to Southern Mexico and Central America, and Ottoschultzia to Southern Mexico, Mesoamerica and Caribbean region. Use Metteniusa edulis H. Karst. has fruits which are reportedly edible. Poraqueiba sericea Tul. has a fleshy fruit rich in oil that is eaten fresh, also seeds that can provide flour. The fruit is widely traded throughout the Amazon rainforest.

Vegetatively not easily distinguished from many tropical families with simple, alternate exstipulate leaves (e.g. some Euphorbiaceae, Flacourtiaceae, Metteniusaceae, and Olacaceae).

SYSTEMATIC 3 subfamilies, all in South America.

1. SUBFAMILY PLATEOIDEAE (2/10) ‣ outsider Platea (5, Indochina, Malesia to New Guinea).

1. Calatola Standl. 7 spp., South Mexico, Central America, and South America (4, mainly Andes), with two spp., C. microcarpa A.H. Gentry ex R. Duno & Janovec and C. costaricensis Standl., in Acre state in Brazil (the former restricted to Peru and Brazil); C. costaricensis Standl. occur from Mexico to C South America.

2. SUBFAMILY APODYTOIDEAE (3/21) ‣ outsiders Apodytes (8, tropical and subtropical Africa, Madagascar, southern India, Sri Lanka, Mauritius, Indochina, Malesia to New Guinea, Queensland, New Caledonia) and Rhaphiostylis (11, tropical W Africa).

2. Dendrobangia Rusby. Trees or shrubs; conspicuous peltate scales with fringed margins cover stems, leaves, inflorescences and external parts of the flowers; dries black. Two spp., D. boliviana Rusby from Costa Rica to Bolivia and SE Brazil (disjunct among this country, Amazon rainforest and Atlantic Forests), and D. multinervia Ducke from Ecuador, Colombia to Peru and NW Brazil.

3. SUBFAMILY METTENIUSOIDEAE (6/24) ‣ outsiders Oecopetalum (3; Mexico, Central America), Pittosporopsis (1; SE Asia) and Ottoschulzia (4, Guatemala, Caribbean).

3. Emmotum Desv. 13 spp. in Colombia, Venezuela, Guyana, Suriname, French Guiana, Peru, Brazil (9, 3 endemics), and Bolivia, mainly the Venezuelan and the Amazon rainforest.

E. nitens (Benthan) Miers grows in savannas of C Brazil (cerrado) across the Brazilian Shield, extending from Brazil into Bolivia and reaching the Amazonian rainforest in the north; E. harleyi R. Duno is only known from the Diamantina Range including the Serra do Tombador in Bahia, Brazil; it grows mainly in rocky grasslands (campos rupestres) over white crystalline sand rock, between 800 and 1200(–1450) m and, more rarely, on gallery forests at 500 m; E. affine Miers is endemic to Paraba to Bahia Atlantic Forest in Brazil coast.

4. Metteniusa Karsten. Canopy trees with alternate leaves and axillary, cymose inflorescences; the flowers are bisexual, fragrant, with five imbricate sepals; five massive, valvate petals basally fused to form a corolla tube up to 2 cm long. 7 spp., 5 endemics to Colombia, M. edulis H. Karst. disjunct in Ecuador and Sierra Nevada de Santa Marta, La Guajira, 910-2000 m elevation range, also Venezuela, and M. tessmanniana (Sleumer) Sleumer from Costa Rica to Peru.

5. Poraqueiba Aubl. Trees. Three spp., P. guianensis Aubl., P. paraensis Ducke and P. sericea Tul., Panamá to Ecuador, east up to Guyana and N Brazil (all species, none endemic).

52. ONCOTHECALES

TROCHODENDRALES DOES NOT OCCUR IN SOUTH AMERICA, AND IS COMPOSED OF A SINGLE FAMILLY, ONCOTHECACEAE (1/2).

53. GARRYALES

GARRYALES DOES NOT OCCUR IN SOUTH AMERICA, AND IS COMPOSED OF TWO FAMILIES: EUCOMMIACEAE (1/1) AND GARRYACEAE (2/27).

54. BORAGINALES

FAMILIES ABSENTS IN SOUTH AMERICA: CODONACEAE (1/2) AND WELLSTEDIACEAE (1/6).

BORAGINACEAE

Genera/spp. 81/1,545-1,665(=Boraginoideae in Angios Bergianska) Distribution temperate (especially warm-temperate) regions on the Northern Hemisphere, fewer species in subtropical areas, on tropical mountains and in the New Guinean Alps, Australia and New Zealand. Habit usually perennial or annual herbs (sometimes shrubs, rarely trees). Use ornamental plants, dyeing substances (Alkanna, Buglossoides etc.), honey flowers (Phacelia etc.), timber (especially Cordioideae), fruits, vegetables (Borago), forage plants (Symphytum), medicinal plants.

SYSTEMATIC three subfamilies, all in South America.

1. SUBFAMILY ECHIOCHILOIDEAE (3/25-30) ▸sister of all remaining Boraginoideae; outsiders Echiochilon (14; NW and E Africa to Arabian Peninsula and NE India), Ogastemma (1; Canary Islands, North Africa, Arabian Peninsula).

1. Antiphytum DC. ex. Meisn. 10 spp., 9 in Mexico, two of then up to North America, and A. cruciatum (Cham.) DC. from Uruguay and Rio Grande do Sul state in S Brazil.

2. SUBFAMILY BORAGINOIDEAE (81/1.545–1.665) ▸two tribes, both in South America.

2.1 BORAGINOIDEAE ▸ TRIBE BORAGINEAE (13/c. 140) ‣ subtribe Boragineae (11/c. 140, Europe, Mediterranean, temperate Asia, northern and southern Africa) do not occur in South America; among Moritzinea, all genera and species occur in South America.

2. Moritzia DC. ex Meisn. Three spp., one from Panamá to Venezuela, and two remaining endemics to S Brazil.

3. Thaumatocaryon Baill. Two spp., both to S Brazil, one reaching into Paraguay and Argentina.

2.2 BORAGINOIDEAE ▸ TRIBE LITHOSPERMEAE (21/c. 460) ‣ outsiders all in Europa, Cape, Africa, E Mediterranean, SW Asia, Türkiye to Himalayas and China, Thailand and Japan.

4. Lithospermum L. 79 spp., temperate regions of both hemispheres, 74 in New World (all in a restricted monophyletic clade), with their highest diversity in North America and Mexico; 11 spp. in South America, L. mediale I.M. Johnst. from Central America, Colombia and Venezuela, and remaining 10 from Ecuador, Peru and Bolivia, highly centered in Amotape-Huancabamba region.

3. SUBFAMILY CYNOGLOSSOIDEAE (81/1,545–1,665) ‣ Lasiocaryeae (3/6, northern India, Central Asia, Himalayas, W China), Asperugeae (4/c. 50, temperate regions on the Northern Hemisphere south to Afghanistan and Mexico, with their highest diversity in North America), Trichodesmeae (2/45, tropical and subtropical regions in the Old World, E Himalayas), Craniospermeae (1/5, temperate Central and E Asia) do not occur in South America.

3.1 CYNOGLOSSOIDEAE ▸ TRIBE OMPHALODEAE (4/40) - outsiders Omphalodes (20–30; Europe, temperate Asia, S Texas, Mexico), Myosotidium (1; Chatham Islands), Iberodes (5; SW Europe, NW Africa).

5. Selkirkia Hemsl. (inc. Cynoglossum p.p.) Perennial erect herbs to subshrubs, or also ascending or decumbent stoloniferous herbs or shrubs; leaves simple, alternate along stem; flowers in ebracteate (rarely few frondose bracts at base) thyrsoids, these sometimes congested and appearing paniculate. 4 spp., S. trianae (Wedd.) Holstein & Weigend from C Colombia to Ecuador; S. berteroi (Colla) Hemsl. from Robinson Crusoe Island on Juan Fernandez Islands, and two endemic to continental Chile.

3.2 CYNOGLOSSOIDEAE ▸ TRIBE ROCHELIEAE (8/c. 180) ‣ outsiders Eritrichium (c 50; temperate regions on the Northern Hemisphere), Rochelia (c 15; Europe, temperate Asia), Pseudoheterocaryum (4; the Caucasus to Iran, Afghanistan and Central Asia), Lepechiniella (4; NE Africa, SW Asia) and Rochelieae s.s. (2/2, the Caucasus, Iran, Afghanistan, Pakistan, Central Asia).

6. Hackelia Vaey ex Beal. Herbs perennial, rarely biennial, basal leaves sessile, cauline leaves peciolate; inflorescences cinccini bracteates. 38 spp., temperate regions on the Northern Hemisphere, 4 of then in mountain regions in Andes from Colombia to Chile and Bolivia.

7. Lappula Moench. 70 spp., Eurasia, Africa, North America, and Australia, 5 in New World, with L. redowskii (Hornem.) Green, of Northern Hemisphere and Chile and S & SE Argentina.

3.3 CYNOGLOSSOIDEAE ▸ TRIBE MYOSOTIDEAE (4/150-170) ‣ outsiders Trigonotis (c 60; E Europe and Central Asia to New Guinea), Brachybotrys (1; Manchuria, Siberia, Korean Peninsula), Decalepidanthus (7–9; Iran and Pakistan to Himalayas).

8. Myosotis L. 151 spp., temperate regions on both hemispheres, tropical mountains, 6 in New World: 4 in North America, and two in South America, both from Chile and Argentina.

3.4 CYNOGLOSSOIDEAE ▸ TRIBE CYNOGLOSSEAE (18/c. 480) ‣ tribes Bothriosperminae (3/9, tropical to NE Asia), Microula clade (1/30-35, Pakistan, NW India, Himalayas, Tibet, W China) and Cynoglossinae (3/c. 170, temperate (especially warm-temperate) regions on the Northern Hemisphere, fewer species in subtropical areas, on tropical mountains and in the New Guinean Alps, Australia and New Zealand) do not occur in South America; among Amsinckiinae, outsiders are Dasynotus (1; NW U.S.A.), Harpagonella (1; SW U.S.A., NW Mexico), Eremocarya (2; SW U.S.A., NW Mexico), Oreocarya (60–65; W North America), Sonnea (2; W North America).

9. Amsinckia Lehm. Herbs annual, erect, inflorescences cymose, often ebracteate, corolla tubular, cylindrical. 14 spp., mainly W U.S.A., 4 in South America, scattered from Ecuador to Argentina, A. tessellata A. Gray also in North America.

10. Cryptantha G. Don. (inc. Nesocaryum) Strigose to hispid, annual, biennial, or perennial herbs, with simple to highly branched, generally ascending to erect stems and simple, basal to cauline, generally linear, lanceolate, or oblanceolate leaves; inflorescences cinccini, single or double, lax or compact, bracts absent. Corollas are almost universally white (yellow in a few species) and are rotate to salverform, with five, often yellow fornices (invaginated, folded regions) surrounding the corolla throat. Flowers chasmogamous or cleistogamous. 107 spp., approximately 60 from Alaska to S Mexico, and from the Pacific coast and east to Texas; 47 spp. in South America: C. albida (Kunth) I.M. Johnst. disjunct North America/Argentina, and 46 reaminig endemic to continent; 39 in Chile (31 endemic), 5 up to Argentina, one of then also in Bolivia, two reaches up to Peru; 4 endemic to Argentina, one endemic to Peru; and one confined to Bolivia/Peru.

11. Greeneocharis Gürke & Harms. Two spp. in North America, G. circumscissa (Hook. & Arn.) Rydb. also in Cono Sur.

12. Johnstonella Brand. 18 spp., 14 in North America, J. albida (Kunth) M.E.Mabry & M.G.Simpson disjunct in Arizona to Texas, Mexico, NW Argentina, and three restrited from W Peru to C Argentina, absents in Bolivia.

13. Pectocarya A. DC. Small herbs, annual, thin; flowers sessile or subsessile; flowers white, short tubular. 12 spp., 8 only North America and 4 endemics to South America from Peru, Bolivia, Chile and Argentina.

14. Plagiobothrys Fisch. & C. A. Mey. Small herbs, annual or perennial; inflorescence cymose; corolla tubular or hypocrateriform, white. 59 spp. in New World, W North America, Mexico, 22 spp. from western South America in Ecuador to Chile and Argentina, 19 in Chile, one in Colombia, few spp. in E Asia and Australia.

HYDROPHYLLACEAE

Genera/spp. 12/235–240(=Hydrophylloideae in Angios Bergianska) Distribuição New World Habit usually herbs (rarely shrubs).

SYSTEMATIC all outsiders from Canada to Mexico.

1. Phacelia Juss. Erect to decumbent, herbaceous, caulescent, branching, usually pubescent and often glandular annuals or perennials from taproots or creeping rootstocks; flowers few to numerous in usually helicoid, long pedunculate to sessile; corolla blue, purple, pale lilac or white; narrowly to widely campanulate; capsule ovoid, loculicidally dehiscent. 207 perennial and annual species, North America (93 spp. in California) and 9 in temperate South America, highly centered in Chile and Argentina, P. pinnatifida Griseb. ex. Wedd. and P. nana Wedd. up to Peru, Bolivia and one up to Uruguay, and P. secunda J.F. Gmel. disjunct Mexico, Peru, Bolivia, Cono Sur.

NAMACEAE

Genera/spp. 4/c. 75(=Namoideae in Angios Bergianska) Distribuição southern U.S.A., Mexico, Central America, West Indies, South America, Hawaii. Habit usually shrubs (rarely herbs).

SYSTEMATIC outsiders Eriodictyon (11; SW U.S.A., NW Mexico), Turricula (1; SW U.S.A., NW Mexico).

1. Nama L. 47 spp. from Canada to Central America and Caribbean, one sp. in Hawaii, three in South America, all disjuct: N. dichotoma (Ruiz & Pav.) Choisy in North America, Mexico, Central America, Ecuador, Peru, Bolivia, Cono Sur; N. jamaicensis L. in North America, Mexico, Central America, Caribbean, Bolivia, Cono Sur; and N. undulata Kunth in North America, Mexico, Cono Sur.

2. Wigandia Kunth. Large, erect, suffruticose to arborescent perennials, variously pubescent and glandular throughout; flowers few to many, in terminal cymes or panicles; corolla greenish-white, yellow or lavender, open-campanulate, often partially pubescent, equal to or usually exceeding the calyx; capsule ovate-oblong, pubescent, dehiscing loculicidally or septicidally, and containing more than 200 minute, reticulate-rugose, brown seeds. 6 spp., two restricted to Caribbean, W. urens (Ruiz & Pav.) Kunth from Mexico to Venezuela, disjunct in Andean Peru; three restricteds of Andes in Ecuador and Peru.

HELIOTROPIACEAE

Genera/spp. 5/400–500(=Heliotropioideae in Angios Bergianska) Distribuição tropical, subtropical and warm-temperate regions on both hemispheres Habit trees, shrubs, lianas or herbs.

SYSTEMATIC outsider Mallotonia (1; Florida, Mexico, Caribbean).

1. Euploca Nutt. (inc. Heliotropium p.p.) Herbs or subshrubs and have usually dry fruits. 56 publicated spp., and more than 45 of Heliotropium and Schleidenia awaiting a transfer to Euploca, in North America to Argentina, Africa, Arabian Peninsula to Indochina, Australia; 42 in New World, 30 in South America, 19 in Brazil (inc. the widely distributed E. humilis (L.) Feuillet), 7 endemics.

2. Heliotropium L. (inc. Tournefortia p.p., exc. Euploca p.p.) Annual herbs (some very small, ephemeral, in dry habitats) to small shrubs, some with rhizomes and underground stolons; scorpioid inflorescence; inflorescences without bracts, exclusively scorpioideous, anthers free, two nutlets, straight embryo and kranz anatomy is absent. 350–400 spp., cosmopolitan, mainly Turanian region and South America; 177 spp. in New World, 125 in South America, 27 spp. in Brazil, 11 endemics; 10 sections in South America:

§ CLADE 1 ‣ sects Hypsogenia and Plagiomeris (both Tournefortia clade) are endemic to the Puna region and to Mediterranean Andes (high Andean zones of central Chile and Argentina) and adjacent Patagonia; all remaining sections occur in Brazil; sections Heliotrophytum and Coeloma are distributed on the eastern side of the Andes and the latter extends into Mesoamerica; sections Platygyne, Schobera and Tiaridium are widely distributed in South America and range into Mesoamerica and the Caribbean; they rarely occur in the Andes, but are present on both Andean flanks; today, they are usually found in human-disturbed areas and are sometimes considered weeds, and their present distribution patterns may thus be of limited phytogeographical relevance; Tiaridium is the only section of Heliotropium that occurs in the Amazonian rainforest, while the rest are restricted to arid, semiarid or even saline environments of the tropics and subtropics. Tournefortia s.s. is widely distributed in the Neotropics (ca. 100–120) from S U.S.A. to S Peru and N Argentina, but is also in the Indo-Pacific Region and E Africa with 12 spp.; it occurs on both sides of the Andes, including the Amazonian rainforest, as well as in the Andean region itself; it is also frequent in Mesoamerica and the Caribbean, but is absent from temperate regions and very rare in dry environments.

§ CLADE 2: sect. Heliothamnus ‣ centered of diversity in the central and northern Andes, with a single species extending into Central America

§ CLADE 3: sect. Cochranea ‣ endemic to the Peruvian and Atacama Deserts and is the only group in Heliotropium distributed only on the western flank of the Andes.

3. Ixorhea Fenzl. Shrubs 1-2.5 m tall, very branched at base; leaves alternate, flowers pedicelate, scent, in terminal panicles; corolla hypocrateriform, lilac to pinkish, inside yellow. Only one sp., I. tschudiana Fenzl, N Argentina in Tucuman and Salta provinces.

4. Myriopus Small. (inc. Tournefortia p.p., Heliotropium p.p.) Shrubs or passive climbers and have 4-lobed fleshy fruits. 24 spp., two widely distributed in Neotropics up to S U.S.A., 8 confined to Caribbean and Central America, and remaining 14 in Brazil up to Santa Catarina state (12 in country, 5 endemics), Colombia, Ecuador, Peru, Bolivia, Guianas and Venezuela.

EHRETIACEAE

§ PARASITIC (Prosopanche – ... – SANTALALES - Lennoa – OROBANCHACEAE - Cuscuta)

Genera/spp. 10/c. 150(=Ehretioideae in Angios Bergianska) Distribuição tropical and subtropical Africa, Madagascar, southern Asia to New Guinea, Australia, the Solomon Islands, southern U.S.A., Mexico, Central America, the West Indies, South America. Habit usually trees or shrubs (rarely herbs; Lennoa and Pholisma achlorophyllous root holoparasites on other Ehretiaceae).

SYSTEMATIC outsiders Halgania (18; Australia), Pholisma (3; SW U.S.A., Mexico).

1. Bourreria P. Browne. Trees or shrubs; bark smooth or slightly striate when young, light to dark gray or brown, longitudinally fissured; flowers actinomorphic, bisexual, corolla sympetalous, mostly white, predominantly membranaceous, rotate to funnelshaped, dehiscence imbricate. 61 spp., 5 in E Africa from Ethiopia to Mozambique and Madagascar, and 56 in New World, centered in the Caribbean and Central America but also occurs in NW Mexico, Florida, and N South America (6), all in Colombia and Venezuela, with B. costaricensis (Standl.) A.H. Gentry reaches to Ecuador, also in Caribbean.

2. Cortesia Cav. (inc. Ehretia p.p.) Small, xerophilic shrub, long-cuneiform leaves with 2–3 tips, small, mostly solitary flowers, and a drupe with a two-parted endocarp. Only one sp., C. cuneifolia Cav. in South America, endemic to W & C Argentina; on dry desert tracks and on saline marshy ground.

3. Ehretia P. Browne. (inc. Rotula) Shrubs, leaves alternate, sessile, cuneiform; flowers terminal, corolla white. 50 spp., pantropical, two in New World, both only in Brazil: Rotula pohlii (Kuhlm.) E.F.Guim. & Mautone endemic to Pará state, and E. lycioides (Mart.) Gottschling & Hilger from China, India, Philippines, NE Brazil (Maranhão, Piauí, Ceará, Pernambuco, Bahia), SE Brazil (Minas Gerais, Rio de Janeiro), Thailand, Vietnam, Laos, peninsular Malaysia, Sri Lanka, Borneo, Sulawesi, New Guinea, Andamans (South Andamans), Nicobars (North Nicobars), Myanmar.

4. Keraunea Cheek & Sim.-Bianch. Scandent shrub or liana, with terete, fistular axes; woody stems 4–5 mm in diameter, with bark brownish white; leaves alternate, simple, exstipulate; inflorescence terminal on short leafy spur shoots, corymbose, 7–8 mm, long, 3–5-flowered, with indumentum as petiole; corolla ca 3.6 mm long; tube ca 0.5 mm long (unusually short in the family). 5 spp. from Atlantic Forest of E Brazil, in Bahia, Minas Gerais, Espírito Santo and Rio de Janeiro states.

5. Lepidocordia Ducke. Two spp., L. williamsii (I.M. Johnst.) J.S. Mill. in of Mexico to Nicaragua, and L. punctata Ducke in Amazon rainforest of Venezuela, Guyana and Brazil in Pará and Roraima states.

6. Lennoa Lex. Annual, succulent holorhizoparasites; stems subterranean or nearly so, white, aging brown, 3- to 15- (-30) cm long, and 0.25-2 cm in diameter; leaves reduced to scales; inflorescence dense, cymose thyrsoid, variously branched, circular to elongate; flowers 8-merous; corolla 2.5- to 9.5-mm long, lavender to bluish-purple, rarely pink, usually with a yellow band in the throat. Only one sp., L. madreporoides Lex. in S Mexico, Guatemala (El Progreso), Colombia (Magdalena) and Venezuela (Falcon), from sea level to about 2,200 m elev., in a variety of habitats.

7. Rochefortia Sw. 11 spp., Mexico, Central America and Caribbean, R. spinosa (Jacq.) Urb. up to NW Colombia, Peru and Venezuela.

8. Tiquilia Pers. 28 spp., arid regions in America, 16 spp. in South America from Ecuador and Galapagos to Andean Bolivia, T. nuttallii (Hook.) A.T. Richardson up to Argentina and T. paronychioides (Phil.) A.T. Richardson up to Bolivia.

CORDIACEAE

Genera/spp. 4/365–370(=Cordioidaee in Angios Bergianska) Distribuição tropical and subtropical regions on both hemispheres, with their highest diversity in the West Indies and South America. Habit trees, shrubs or lianas (Coldenia herbaceous).

SYSTEMATIC outsiders Coldenia (1; Old World tropics), Hoplestigma (2; Cameroon, Gabon).

1. Cordia L. Usually erect shrubs, sometimes trees up o 40 m tall; bark often fibrous. 211 spp., tropical and subtropical regions on both hemispheres, 57 spp. in Old World (55 in Myxa s.s., and 2 in Sebestena s.s.), 174 in New World, mainly South America (118); 55 spp. in Brazil, 25 endemics; two spp. are rare in Brazil: C. decipiens I.M.Johnst. from Amazonas state and C. andrade-limae from Paraíba state (not recognized at VPA), by Plantas Raras do Brasil’s book. 5 spp. from tropical America (inc. Brazil) are myrmecophites.

2. Varronia P. Browne. Herbs, shrubs, subshrubs, trees, often with xylopodium. 136 spp., New World, 66 in South America, 35 spp. in Brazil, 23 endemics.

55. GENTIANALES

ALL FAMILIES IN SOUTH AMERICA.

RUBIACEAE

§ FULLY CHLOROTROPHIC

Genera/species 611/11,470–11,550 Distribution cosmopolitan except polar areas, with their highest diversity in tropical and subtropical regions. Habit usually bisexual (rarely monoecious, polygamomonoecious or dioecious), evergreen or deciduous trees, shrubs, lianas or suffrutices, or perennial, biennial or annual herbs. Hydnophytum, Myrmecodia, Myrmeconauclea, Myrmephytum, etc. are epiphytic or (Myrmeconauclea) rheophytic myrmecophytes (ant plants), in the hollow swollen stems and branches – hypocotylar bases – of which ant colonies live. Young stems and branches often quadrangular in cross-section.

HABITATS predominantly pantropical, and with a small portion of spp. of extra-tropical distribution. Almost one half of the spp. (and about one third of the genera) occur in the Neotropics. In South America, they are adapted to virtually every habitat: from páramo to arid and desertic environments. Rubiaceae are especially diverse in the Amazon Basin, Andean cloud forests, savannas of C Brazil (cerrado), dry seasonal scrubland of NE Brazil (caatinga), Atlantic sandy coastal shrublands (restingas), and the Atlantic forests of Brazil. The two main centers of endemism in the Neotropics are the Guiana Shield and the Greater Antilles.

HABIT The Rubiaceae is well represented at all layers of tropical vegetation, with all kinds of habits, as herbs, shrubs, lianas, and from small trees to tall canopy trees, and all dimensions; Rubiaceae ranges from 5 cm tall (Spermacoce) to 55 m tall (Chimarrhis); most spp. of Hillia Jacq., Cosmibuena Ruiz & Pav., and several spp. of Notopleura (Benth. & Hook. f.) Bremek. and Psychotria L. are epiphytic shrubs adapted to live in the forest canopy; Limnosipanea Hook. f. is a short-seasonal, semi-aquatic herb endemic to seasonally inundated habitats of central Brazil and the Venezuelan llanos; Many members of the Spermacoceae are herbs and subshrubs frequently found in disturbed habitats (e.g., cow pastures and forest edges); Weedy genera such as Galium L., Spermacoce G.Mey., Diodia L., Richardia L. and Mitracarpus Zucc. have distribution sometimes worldwide.

CURIOSITIES leaf blades undivided (exceptionally deeply lobed in Genipa infundibuliformis Zappi & Semir and several spp. of Pentagonia Benth.); corollas gamopetalous (except for Dialypetalanthus Kuhlm., with distinct petals); stamens commonly as many as corolla lobes (except for Dialypetalanthus, with two rows of 15–25 stamens); a few other genera of Rubiaceae present calycophylls (common in Ixoroideae, only in Kerianthera in Cinchonoideae and Oreopolus in Rubioideae); zygomorphic flowers in Coutarea, Hillia, Palicourea, Henriquezieae and Posoqueriae.

USES the most economically valuable genus is Coffea (Coffee), which is the worlds most heavily traded commodity after oil. spp., of cultivated Coffea L. commonly escape into forests near farms in Brazil. Cinchona oficinalis L. (known as quina) is cultivated for quinine have been introduced in many tropical countries; Genipa americana L., known as genipapo in Brazil, is cultivated throughout South America for edible fruits and dye; Psychotria ipecacuanha (ipecacuanha, an expectorant); Uncaria gambier (gambier, an important tannin source); Uncaria spp. (medicines); Calycophyllum spp. (lemonwood, a timber). The family also contains some of the most beautiful tropical ornamentals (Ixora, Gardenia, Mussaenda, Portlandia, Serissa). Rubiaceae also provide local indigenous populations with remedies against malaria and other diseases, body paints, edible fruits and construction wood.

Two genera are particularly odd morphologically: Dialypetalanthus: chemistry; cork cortical, phloem stratified; K free, C free, opposite K, both in two decussate pairs; A (8-)16-17(-25), not epipetalous, basally connate, anthers basifixed, dehiscing by pores, bracteoles apparently ad/abaxial. Oil glands are described as being pervasive; Theligonum: iridoids and raphide sacs +; plant monoecious, anemophilous; staminate flowers: paired, opposite, not subtended by leaves; P tubular, ridged, splitting into 2-5 segments; A 2-30, development variable; pollen 4-8-zonoporate; carpellate flowers: G 1-locular by suppression, with 1 campylotropous ovule; testa single-layered, walls thin; endosperm with starch, embryo curved. Vegetatively Theligonum is a good match with Rubioideae, where it is included by B. Bremer. The differences between it and other Rubiaceae are those that might be expected of a wind-pollinated plant, i.e., monoecy, reduced and modified perianth, many stamens, and only a single-seeded fruit.

SYSTEMATIC five lineages, Luculieae (1/5, Himalayas, Yunnan, Burma, S China, N Thailand, N Indochina) and Coptosapelteae (2/c 56, India to central China, SE Asia, West Malesia to Philippines, with their highest diversity on Borneo) do not occur in South America.

UNPLACED RUBIACEAE - outsiders Benzonia (1; Guinea), Berghesia (1; Mexico), Byrsophyllum (2; India, Sri Lanka), Clarkella (1; Himalayas, Thailand), Coptophyllum (8; India, West Malesia), Eizia (1; Mexico), Fergusonia (1; SE India, Sri Lanka), Klossia (1; Malay Peninsula), Koenania (1; China), Lecariocalyx (1; Borneo), Lorencea (1; S Mexico, Guatemala), Maschalodesme (2; New Guinea), Nodocarpaea (1; Cuba), Placocarpa (1; Mexico), Pseudodiplospora (1; Andaman Islands), Sacosperma (2; tropical Africa), Sinospiradiclis (1; China), Streblosiopsis (1; Borneo), Stylosiphonia (1; Central America), Temnocalyx (1; Tanzania), Tortuella (1; Île Tortue near Hispaniola).

1. Acrobotrys K.Schum. & K.Krause. Shrubs to small trees. Only one sp., A. discolor K. Schum. & K. Krause, endemic to Cauca region in Colombia, about 1,000 m elevation range.

2. Aphanocarpus Steyerm. Only one sp., A. steyermarkii (Standl.) Steyerm., endemic sandstone table mountains and adjacent highs of Bolivar state in Venezuela, 1,000 – 2,500 m elevation range.

3. Coryphothamnus Steyerm. Only one sp., C. auyantepuiensis (Steyerm.) Steyerm., endemic to Auyan-tepui in Venezuela, at 1,700-2,300 m elevation range.

4. Didymochlamys Hook.f. Herbs, epiphytic or terrestrial. Two spp. from Nicaragua to Guyana and Ecuador.

5. Duidania Standl. Only one sp., D. montana Standl., endemic to the sandstone mountains of Duida, Huachamacari, and Marahuaca of Guyana and Venezuela, 1,200 – 2,600 m elevation range.

6. Etericius Desv. ex Ham. Only one sp., E. parasiticus Desv., endemic to Guyana.

7. Flexanthera Rusby. Only one sp., F. subcordata Rusby, endemic to Colombia.

8. Holstianthus Steyerm. Only one sp., H. barbigularis Steyerm., endemic to the Guiana Shield of Venezuela, 1,200-1,400 m elevation range.

9. Pagameopsis Steyerm. Two spp., endemic various tepuis of the Venezuelan Guiana and adjacent northern Brazil in Amazonas state (only P. maguirei Steyerm.), 1,100 to 2,500 m elevation range.

10. Pseudohamelia Wernham. One sp., P. hirsuta Wernham, endemic to Colombia.

11. Riqueuria Ruiz & Pav. One sp., endemic to Peru.

1. SUBFAMILY CINCHONOIDEAE (c 113/1,500–1,530) ▸ 10 lineages, only Hymenodictyeae (2/25, Madagascar, tropical regions in the Old World to Sulawesi) is absent in South America.

1.1 CINCHONOIDEAE ▸ TRIBE CINCHONEAE (9/110–120) - all genera in South America.

12. Ciliosemina Antonelli. Two spp., Venezuela, Colombia (one endemic), Ecuador, Peru, Brazil (only C. pedunculata (H. Karst.) Antonelli).

13. Cinchona L. Shrubs or small trees, bark exfoliating. 24 spp., forested mountains fro Venezuela to central Bolivia, one up to Costa Rica, 8 endemics to Peru.

14. Cinchonopsis L.Andersson. Trees in forests. Only one sp., C. amazonica (Standl.) L. Andersson, restricted from Amazonia rainforest in Brazil, Venezuela, Ecuador, Colombia and Peru.

15. Joosia H.Karst. Shrubs and small trees lineolate tertiary leaf venation, its well-developed interpetiolar stipules that are quickly deciduous, its cymose inflorescences, its salverform 4- or 5-lobed corollas, fragrante, white or pink, and its slender, cylindrical, septicidal capsules containing numerous flattened seeds 18 spp. from Venezuela to northern Bolivia, one up to Panamá, centered in Ecuadorean and Peruvian Andes.

16. Ladenbergia Klotzsch. Trees medium sized to tall, bark exfoliating, Cinchona-like. 41 spp., Costa Rica to Guyana, Bolivia and N Brazil (8, 4 endemics); 37 in South America.

17. Maguireocharis Steyerm. Only one sp., M. neblinae Steyerm., endemic to the Guiana Shield of S. Venezuela (Mount Neblina) to Brazil (Serra Pirapucu, Amazonas state), 1,300 – 1,400 m elevation range.

18. Pimentelia Wedd. Only one sp. from Colombia to Peru.

19. Remijia DC. Shrubs to medium sized trees. 40 spp. from over tropical South America, 21 in Brazil, 12 endemics. R. glomerata Huber and R. physophora Benth. ex K.Schum. from N Brazil and Venezuela, are myrmecophites.

20. Stilpnophyllum Hook.f. Shrubs or small trees. 4 spp., Colombia to Peru.

1.2 CINCHONOIDEAE ▸ TRIBE ISERTIEAE (2/15) - both genera in South America.

21. Isertia Schreb. Shrubs or small trees. 15 spp. from South America, 4 up to Central America until Guatemala, Brazil, up to French Guiana, also in Cuba and Guadalupe; 10 spp. in Brazil, one endemic.

22. Kerianthera J.H.Kirkbr. Trees, leaves often 1m longer; large, terminal inflorescences with some calyx-lobes transformed into calycophylls, multi-locellate anthers and fruit with numerous winged seeds. Two spp., K. preclara J. H. Kirkb only found in a small area north of Manaus, Amazonas state, and K. longiflora Zappi & C. T. Oliveira, from the Atlantic Forest of states of Espírito Santo and Minas Gerais.

A few other genera of Rubiaceae present calycophylls, such as Pogonopus, Capirona, some species of Simira and Warszewiczia. Amongst them, due to its long corollas, Kerianthera is more likely to be confused with Pogonopus, however the multi-locellate anthers are observed only in Kerianthera and Isertia in the neotropics and in a few other African species.

1.3 CINCHONOIDEAE ▸ TRIBE RONDELETIEAE (13/c 210) - outsiders Tainus (1; Hispaniola), Acrosynanthus (7; Caribbean), Mazaea (2; Cuba), Phyllomelia (1; Cuba), Arcythophyllum (c 15; tropical America), Roigella (1; Cuba), Rovaeanthus (2; Mexico, Central America), Suberanthus (7; Cuba, Hispaniola), Acunaeanthus (1; Cuba), Blepharidium (1; Central America), Habroneuron (1; Mexico), Lindenia (2–3; New Caledonia; Fiji; Central America).

23. Rondeletia L. Shrubs or small trees. 154 spp., S. Mexico to Peru, Caribbean (136, all restricteds) to N. South America (Venezuela to Peru); 13 spp. in South America.

1.4 CINCHONOIDEAE ▸ TRIBE NAUCLEEAE (17/180–200) - outsiders Mitragyna (7; tropical regions in the Old World), Sinoadina (1; China, Japan, Burma, Thailand), Haldina (1; India, Sri Lanka, S China, SE Asia), Adina (15; China, Korean Peninsula, Japan, tropical Asia to New Guinea), Khasiaclunea (1; NE India), Neonauclea (75–80; S China, tropical Asia to islands in W Pacific), Diyaminauclea (1; Sri Lanka), Breonadia (1; tropical Africa, Madagascar), Breonia (6–20; Madagascar; paraphyletic), Gyrostipula (3; Madagascar, the Comoros), Janotia (1; Madagascar), Corynanthe (8; tropical West and Central Africa), Pseudocinchona (3; tropical Africa), Neolamarckia (2; tropical Asia to tropical Australia), Nauclea (12; tropical regions in the Old World).

24. Cephalanthus L. 6 spp., one from Tanzania to S. Africa, two from India to Taiwan in China, two from Canada to Guatemala, Cuba, and C. glabratus (Spreng.) K. Schum. in S Brazil (Mato Grosso do Sul to Ro Grande do Sul and São Paulo states), E Paraguay, Uruguay & NE Argentina.

25. Uncaria Schreb. Woody lianes or sometimes forming scrambling bushes or thickets; young stems ± quadrangular; flowering branches bearing strongly curved hooked spines; leaves paired, petiolate, elliptic, apex acuminate; inflorescences globose, solitary, axillary or terminal; flowers not fused, pedicellate; fruits fusiform, dry, dehiscent; seeds small. 34 spp., mainly Asia and Australia; two spp. in New World, from Belize to Paraguay, up to French Guiana and Caribbean, both in South America and in Brazil.

1.5 CINCHONOIDEAE ▸ COLLETERIA + CHIONE CLADE (2/3) - outsider Colleteria (2; the Caribbean).

26. Chione DC. Shrubs or medium sized trees. Only one sp., C. venosa (Sw.) Urb., Mexico to Peru, Caribbean, and in Acre state in western Amazonian Brazil.

1.6 CINCHONOIDEAE ▸ TRIBE HILLIEAE (3/29) - outsider Balmea (1; Mexico)

27. Cosmibuena Ruiz & Pav. Shrubs or small trees, epiphytic or terrestrial. 4 spp., SE. Mexico to Venezuela, Bolivia and Brazil (only the widely C. grandiflora (Ruiz & Pav.) Rusby).

28. Hillia Jacq. Scandent or shrubs hemiepiphytic or epiphytic, glabrous, fleshy. 27 spp., Mexico to Bolivia, up to French Guiana, Caribbean, 19 in South America, 4 in Brazil, one endemic.

1.7 CINCHONOIDEAE ▸ TRIBE HAMELIEAE (11/175–180) - outsiders Syringantha (1; Mexico); Pinarophyllon (2; Central America), Deppeopsis (3; S Mexico), Pseudomiltemia (2; S Mexico), Plocaniophyllon (1; Chiapas in Mexico), Omiltemia (4; Mexico), Renistipula (3; S Mexico, Central America); Cosmocalyx (1; Mexico).

29. Deppea Schltdl. & Cham. 36 spp., 35 from Mexico to Panamá, and D. blumenaviensis (K.Schum.) Lorence, restricted from S Brazil to NE. Argentina.

30. Hamelia Jacq. Shrubs or small trees. 16 spp., over Neotropics, of Florida and Mexico to Paraguay, French Guiana, Caribbean; 7 spp. in South America, two in Brazil, none endemics.

31. Hoffmannia Sw. Tiny herbs to herbaceous shrubs, Fuchsia-like; opposite or verticillate leaves, deciduous stipules that are interpetiolar or very shortly fused around the stem, axillary cymose inflorescences of varied form, 4-merous homostylous flowers, rather small corollas with the lobes imbricate in bud, generally 2-locular ovaries, and small fleshy baccate fruits containing numerous angled seeds. 121 spp., Mexico to to Argentina, up to Venezuela, Caribbean, 45 in South America, only two in Brazil, H. duckei Standl. endemic. H. vesciculifera Standl., from Panamá and Colombia, is a myrmecophite.

32. Patima Aubl. Low shrubs, mainly unbranched, both myrmecophites. Two spp., P. guianensis Aubl. endemic to W Guyana, and P. minor C.M. Taylor from French Guiana to Guyana and Pará state in N Brazil.

1.8 CINCHONOIDEAE ▸ CHIOCOCCEAE (28/190–195) - outsiders Siemensia (1; Cuba), Bikkia (c 20; East Malesia to islands in western Pacific), Badusa (3; Palawan, New Guinea, islands in western Pacific), Scolosanthus (27; the Caribbean), Schmidtottia (16; Cuba), Ceuthocarpus (1; Cuba), Eosanthe (1; Cuba), Phialanthus (22; the Caribbean), Ceratopyxis (1; Cuba), Isidorea (17; the Caribbean), Portlandia (6; Jamaica), Cubanola (2; Cuba, Hispaniola), Solenandra (22; Mexico and Central America, Cuba, Hispaniola, Jamaica to the Lesser Antilles), Cigarrilla (1; Mexico), Nernstia (1; Mexico), Osa (1; Costa Rica), Thogsennia (1; Cuba, Hispaniola), Phyllacanthus (1; Cuba), Catesbaea (17–20; Florida Keys, the Caribbean), Asemnantha (1; Mexico), Coutaportla (2; Mexico), Morierina (2; New Caledonia), Shaferocharis (3; Cuba), Ramonadoxa (1, Cuba).

33. Adolphoduckea Paudyal & Delprete. (off Exostema). Tree, 7–30 m tall, to 1 m dbh. Only one sp., A. maynense (Poepp.) Paudyal & Delprete, in Ecuador to Bolivia and Brazil, only in Acre state and possibly Amazonas.

34. Chiococca P.Browne. (exc. Salzmannia p.p.) Lianas (scandent) or shrubs, fruits often aplaned. 25 spp., North to South America (7), 3 in Brazil, one endemic.

35. Coutarea Aubl. (exc. Coutareopsis) Shrubs or small trees. Two spp., C. alba Griseb. from Venezuela to Bolivia, and C. hexandra (Jacq.) K. Schum. widely in Mexico, Central America, Trinidad & Tobago, Venezuela, Colombia, Ecuador, Peru, Bolivia, Brazil and Cono Sur.

36. Coutareopsis Paudyal & Delprete. (off Coutarea) Shrubs 1.5–5.0 m tall; branches laterally compressed or terete, puberulent to glabrous, with lateral short shoots. 3 spp., from Ecuador and Peru.

37. Erithalis P.Browne. Shrubs or small trees. 8 spp., Florida, Mexico to Colombia, Venezuela, Guyana, Caribbean, centered in Antilles, only E. fruticosa L. in South America.

38. Exostema (Pers.) Rich. ex Humb. & Bonpl. (exc. Adolphoduckea) Shrubs or small to tal trees. 32 spp., 31 in S Florida, Mexico to Central America and Caribbean, and E. corymbosum (Ruiz & Pav.) Spreng. endemic to Peru.

39. Motleyothamnus Paudyal & Delprete. Shrubs or small trees, 1–6(–10) m tall. Only one sp., M. corymbosus (Ruiz & Pav.) Paudyal & Delprete, known from open places and shrublands of the Andes in Peru, at 1000–2800 m elevation, on slopes along streams in both moist and dry areas.

40. Salzmannia DC. (inc. Chiococca p.p.) Shrubs or treelets; branches tetragonal to terete, glabrous, with copious resinous exudates. 4 spp., one in higher elevation on the costal cordillera of Venezuela, and three species growing in the coastal dunes and restinga and forested vegetation of NE Brazil.

41. Strumpfia Jacq. Only one sp., S. maritima Jacq., in Florida, Mexico, Caribbean, Central America and N Venezuela.

1.9 CINCHONOIDEAE ▸ TRIBE GUETTARDEAE (c 20/680–685) - outsiders Rogiera (c 15; S Mexico, Central America, the Caribbean), Allenanthus (2; Central America), Antirhea (c 40; Madagascar, Malesia to Samoa), Bobea (4; Hawaii), Hodgkinsonia (2; Queensland, New South Wales), Tinadendron (2; Vanuatu; New Caledonia), Timonius (c 170; Mauritius, Seychelles, Sri Lanka, Andaman Islands, Malesia to islands in the Pacific, with their largest diversity on New Guinea), Javorkaea (1; Honduras), Resinanthus (16; the Caribbean, Mexico).

42. Arachnothryx Planch. Shrubs or small trees. 100 spp., Mexico to Peru and Trinidad, Venezuela; 24 spp. in South America.

43. Chomelia Jacq. (inc. Antirhea, exc. Guettarda p.p.) Shrubs or small trees, often spiny. c. 300 spp., 78 spp. in over Neotropics, 65 in South America, 37 in Brazil, 27 endemics, one of them, from São Paulo state, is a rare plant in Brazil, by Plantas Raras do Brasil’s book.

44. Gonzalagunia Ruiz & Pav. Herbs, shrubs or small trees. 37 spp., Mexico to trop. America, 26 in South America, only three in Brazil, none endemics.

45. Guettarda L. (inc. Chomelia p.p.) Shrubs or small trees, often spiny. 156 spp., trop. & subtrop, 139 of then in tropical America, 46 in South America, 20 in Brazil, 12 endemics, two of them, both from Bahia state, are rare plants in Brazil, by Plantas Raras do Brasil’s book.

46. Machaonia. Bonpl. Shrubs or small trees. 32 spp., Mexico to South America (8), only two in Brazil, none endemics.

47. Malanea Aubl. Lianas, shrubs or trees. 40 spp. from South America (three up to Central America and Mexico) except one restricted to Caribbean, 15 in Brazil, 8 endemics.

48. Neoblakea Standl. Two spp., Ecuador and Venezuela one endemic each.

49. Pittoniotis Griseb. Medium sized to tall trees. Two spp., P. trichantha Griseb. from S. Mexico to Venezuela and Colombia, and P. rotata C.M. Taylor endemic to Ecuador.

50. Stenostomum C.F.Gaertn. Resinous young growth, leaves with the tertiary veins either not visible or arranged in generally regular rectangular areoles, axillary inflorescences, perfect flowers, 4- to 5-lobed calyx limbs, slender tubular to salverform corollas with the four to five lobes imbricate in bud. 46 spp., S. Mexico to trop. America, only two in South America, only S. acreanum (K. Krause) Achille & Delprete in Brazil (northern region, no endemic).

2. SUBFAMILY IXOROIDEAE (220–225/3,700–3,740) - [Posoquerieae + Sipaneeae] + Condamineeae + [[Mussaendeae + Sabiceeae] + [Steenisia + [Retiniphyllum + [‘Vanguerieae alliance’ + ‘Coffeeae alliance’]]]]; 12 tribes, Mussaendeae (6/225–235,tropical regions in the Old World (absent from Australia and eastwards)) and Steenisieae (1/5, Borneo, Natuna Islands) do not occur in South America.

2.1 IXOROIDEAE ▸ TRIBE POSOQUERIEAE (5/c 40) - all genera in South America.

51. Gleasonia Standl. 5 spp., all endemic to the Guiana Shield, one only in Venezuela and remaining four only in Brazil, except by one sp. in S Amazon rainforest of Brazil. G. uaupensis Ducke, endemic to Brazil, is a myrmecophite.

52. Henriquezia Spruce ex Benth. Shrubs or small trees. Three spp., endemic to the Guiana Shield, one in Guyana and two widely in Colombia, Venezuela and Amazonas state in N Brazil (both, none endemics), 50 – 300 m elevation range.

53. Molopanthera Turcz. Large tree (5 – (10) - 30). Only one sp., M. paniculata Turcz., restricted to Atlantic Forest of Bahia to Rio de Janeiro states in E Brazil.

54. Platycarpum Humb. & Bonpl. Shrubs to large trees, mainly covered by goldish hairs. 14 spp., endemic to the Guiana Shield of Colombia to Guyana and N Brazil, except by one disjunct from the Venezuelan Guiana to Amazon Peru, a single endemic to Peru, and two in W and C Amazonian Brazil; 6 spp. in Brazil, three endemics, one of them, from Acre state, is a rare plant in Brazil, by Plantas Raras do Brasil’s book.

55. Posoqueria Aubl. Shrubs or small trees. 25 spp., confined to Central and South America (19), one up to S Mexico; 10 spp. in Brazil, 6 endemics, one of them, from Bahia state, is a rare plant in Brazil, by Plantas Raras do Brasil’s book.

2.2 IXOROIDEAE ▸ TRIBE SIPANEEAE (10/c. 43) - outsider Steyermarkia (1; Chiapas in Mexico to Guatemala).

56. Chalepophyllum Hook.f. Subshrubs to shrubs; distal internodes, base of peti-oles and base of peduncles often covered by a resinous exudate produced by stipular colleters. Only one sp., C. guianense Hook. f., endemic to the Guiana Shield of SE Venezuelan and adjacent Guyana and Mount Caburai, northern Roraima state, Brazil, 300-1,430 m elevation range.

57. Dendrosipanea Ducke. Shrubs or treelets. Stipules free or basally adnate to petioles, deltoid, acuminate or bifid, with basal 2/3 cov-ered by dense colleters intermixed with hairs inside; leaves opposite, subsessile to short-petiolate. 3 spp., endemic to the Guiana Shield of Colombia (1), Venezuela (2) and Amazonas state in N Brazil (all 3), 30-110 m elevation range.

58. Limnosipanea Hook.f. Herbs, terrestrial, semi-aquatic (amphibious) or aquatic (in L. spruceana); erect, prostrate or decumbent; ephemeral to short-lived (2–4 months life span) when terrestrial or semi-aquatic, or perennial when grow-ing in permanently submersed areas. Three spp., Panamá to Venezuela and Brazil (all spp., one endemic), reported also Suriname to Bolivia.

59. Maguireothamnus Steyerm. Shrubs. 3 spp. endemic to the Guiana Shield of Guyana, Venezuela (all spp., two endemics) and Brazil (one sp., Mount Roraima and Serra do Sol in Roraima state), 1,300-2,600 m elevation range.

60. Neblinathamnus Steyerm. Shrubs. Two spp. restricteds for Neblina Massif, both in Venezuela, N. brasiliensis Steyerm. up to Brazilian side, 1,300-1,900 m elevation range.

61. Neobertiera Wernham. Perennial herbs, subshrubs or shrubs. 5 spp., all from Guiana Shield, 100-300 m elevation range, two only in Guyana, two only in French Guiana, and N. montedouradensis Delprete endemic to dense forests of Amapá and Pará states, N Brazil.

62. Pteridocalyx Wernham. Shrub 1–2 m tall, with a central, thin stem, sparsely branched towards the top; young branches reddish, older stems and branches pale gray, with calycophylls. Only one sp., P. appunii Wernham, endemic Upper Potaro River Basin in Guyana.

63. Sipanea Aubl. Herbs, erect, trailing, prostrate or decumbent, often rooting at basal nodes, sometimes mat-forming, rarely floating on water in inundated areas (S. biflora), or sub-shrubs (stems woody at base), erect or decumbent. 17 spp. from South America, two up to Nicaragua and Trinidad, mainly in Venezuela (10), Guyana (9), Suriname (9), Brazil (7, one endemic) and French Guiana (6).

64. Sipaneopsis Steyerm. Subshrubs or shrubs, rarely herbs, erect, ascending or rarely decumbent. 8 spp., endemic to the Guiana Shield of Colombia (2), Venezuela (6, 4 endemics), and . Brazil (3, two endemics), 50-800 m elevation range.

2.3 IXOROIDEAE ▸ TRIBE CONDAMINEEAE (31/355–360) - outsiders Emmenopterys (2; SW China, Burma, Thailand), Pinckneya (1; SE U.S.A.), Dolicholobium (c 30; Philippines to Fiji), Mussaendopsis (3; West Malesia), Mastixiodendron (7; East Malesia to Fiji), Picardaea (1–2; Cuba, Hispaniola), Hintonia (3; Central America).

65. Alseis Schott. Trees or shrubs with triangular to deltoid interpetiolar stipules, apical, spiciform or racemose inflorescence, protogynous flowers with a cylindricaltubular perianth, clavate, septicidal, many-seeded capsules. 19 spp., S. Mexico to Bolivia, Brazil (11, 5 endemics), east to French Guiana; 16 spp. in South America.

66. Bathysa C.Presl. Shrubs or small trees, often with calycophylls. 9 spp. from N South America up to S Brazil (7, 5 endemics, one of them, from Rio de Janeiro state, is a rare plant in Brazil, by Plantas Raras do Brasil’s book).

67. Bothriospora Hook.f. Trees. Only one sp., B. corymbosa (Benth.) Hook. f., from Guyana to Peru and N Brazil.

68. Calycophyllum DC. Medium to tall trees up to 30 m tall (tallest neotropical Rubiaceae), emergent in canopy forests, bark exfoliating, often with calycophylls. 11 spp. from South America (one up to Mexico and Caribbean), east to Guyana; 7 spp. in Brazil, two endemics.

69. Capirona Spruce. Tall trees, often emergent in canopy forest, bark exfoliating, often with calycophylls. Only one sp., C. decorticans Spruce, French Guiana to Bolivia and N Brazil.

70. Chimarrhis Jacq. Small to tall trees, with buttresses, very hard wood, often with calycophylls. 15 spp., Nicaragua to Peru, east to French Guiana, Caribbean; 10 spp. in South America, 6 in Brazil, two endemics.

71. Condaminea DC. Shrubs or small trees. 5 spp. from South America, one up to Costa Rica, only the widely C. corymbosa (Ruiz & Pav.) DC. in Brazil.

72. Dialypetalanthus Kuhlm. Trees, flowers white, with two rows of stamens. Only one sp., D. fuscescens Kuhlm., from N Bolivia, W Amazon Brazil and E Peru.

Dialypetalanthus brooken the 'normality' of flowers in this family: this genus has distinct petals (unlike all others Rubiaceae), and many time more stamens - 15 to 25 - that corolla lobes.

73. Dioicodendron Steyerm. Shrubs or small trees, many branched. Two spp., Andes from Venezuela to Bolivia.

74. Dolichodelphys K.Schum. & K.Krause, Verh. Shrubs or small trees, many branched. Only one sp., D. chlorocrater K. Schum. & Krause, Venezuela to Peru.

75. Elaeagia Wedd. Large shrubs to large trees, mainly resinous. 26 spp., Mexico to Bolivia, 21 in South America up N Brazil (only E. maguirei Standl., also in Venezuela and Guianas), east to Suriname, Caribbean.

76. Ferdinandusa Pohl. Shrubs to tall trees. 23 spp. from South America (one up to Nicaragua), 19 in Brazil, 8 endemics.

77. Hippotis Ruiz & Pav. Shrubs or small trees. 22 spp. from South America (two up to Central America), from Venezuela to Peru.

78. Lintersemina H. Mendoza-Cifuentes & A. Celis & M.A. González. Treelet, leaves opposite; inflorescences axillary, subterminal, corymbose cymes, pedunculate, many-flowered; flowers hermaphroditic, 5- or 6-merous, actinomorphic. Only one sp., L. chucuriensis H. Mendoza-Cifuentes, A. Celis & M.A. González, endemic to Magdalena Medio Region of Colombia, growing in tropical rainforest.

79. Macbrideina Standl. Small to médium sized trees. Only one sp., M. peruviana Standl., from Amazon in Colombia, Ecuador, Peru and Bolivia.

80. Macrocnemum P.Browne. Shrubs to tall trees. 8 spp., 7 from Colombia to Bolivia, M. roseum (Ruiz & Pav.) Wedd. up to Acre state in N Brazil and Costa Rica, and one endemic to Jamaica.

81. Parachimarrhis Ducke. Medium to tal trees, with buttresses, bark exfoliating; inflorescence terminal corymbose Only one sp., P. breviloba Ducke, from Amazonian Colombia, Peru and N Brazil.

82. Pentagonia Benth. Shrubs or small trees, often very tal trees. 46 spp., Guatemala to Peru, 39 in South America, Amazonian Brazil (4, none endemics), a half in Colombia. Impressive for its large leaves, Pentagonia stands out for being the genus of the only species of Rubiaceae with normal, mature leaves pinnately lobed to deeply pinnatifid; in the New World, occasional individuals of some species of Simira Aubl. may have pinnatifid leaves and a few species of Cruckshanksia Hook. & Arn. have leaves deeply and digitately 2 or 3 lobed.

83. Pogonopus Klotzsch. Shrubs or small trees, often with calycophylls. 4 spp., Mexico to Bolivia along Colombia to S Brazil (only the widely P. tubulosus (A. Rich.) K. Schum.).

84. Rustia Klotzsch. (inc. Tresanthera) Shrubs or small trees. 18 spp., 14 from Guatemala to Peru, Venezuela and N Brazil (the widely R. thibaudioides (H. Karst.) Delprete), and 4 remaining disjunct in S Brazil; 15 spp. in South America.

85. Schizocalyx Wedd. (inc. Phitopis) Trees and shrubs, often with calycophylls; flowers bisexual, homostylous, protandrous, sessile to pedicellate, fragrant; corolla campanulate to funnelform or tubular, pale green, white, pink, or white flushed with purple. 9 spp., one in Central America, seven in tropical Andes, and S. cuspidatus (A. St.-Hil.) Kainul. & B. Bremer endemic to SE Brazil, found in wet to moist forests from low to middle elevations.

86. Simira Aubl. Shrubs to large trees, often with calycophylls, easily recognized in the forest due to the characteristic red color that the wood acquires immediately after cutting. 43 spp., Mexico to Argentina, east to French Guiana, Brazil (20, 13 endemics); 39 spp. in South America; three spp., from Rio de Janeiro and Espírito Santo states, are rare plant in Brazil, by Plantas Raras do Brasil’s book.

87. Sommera Schltdl. Shrubs or small trees 11 spp., Mexico to Panamá, three up to South America in Colombia, Ecuador, Peru and N Brazil (only S. sabiceoides K. Schum.).

88. Tammsia H.Karst Only one sp., T. anomala (H. Karst.) H. Karst., from Venezuela to Peru.

89. Warszewiczia Klotzsch. Shrubs to trees, often with calycophylls. 8 spp. from South America (two up to Mexico and Caribbean), 4 in Brazil, one endemic.

90. Wittmackanthus Kuntze. Shrubs to trees up to 35 m tall, often with calycophylls. Only one sp., W. stanleyanus (Schomb.) Kuntze, Panamá to Peru, E to Guyana.

2.4 IXOROIDEAE ▸ TRIBE SABICEEAE (4/c. 140) - outsiders Tamridaea (1; Socotra), Virectaria (8; tropical Africa), Hekistocarpa (1; Nigeria, Cameroun).

91. Sabicea Aubl. Lianes, straggling shrubs or sometimes scarcely woody climbers, usually ± hairy; leaves opposite, petiolate; flowers sometimes showing limited heterostyly, small or medium-sized, paniculate or capitate axillary inflorescences, or in a few cases plant cauliflorous. 130 spp., trop. Africa, Madagascar, Sri Lanka, 49 spp. in Mexico to trop. America (47 in South America), 19 in Brazil, 6 endemics.

2.5 IXOROIDEAE ▸ TRIBE RETINIPHYLLEAE (1/23) - a single genus.

92. Retiniphyllum Bonpl. Shrubs or small trees. 23 spp., endemic to the Guiana Shield of Suriname to Colombia and Brazil, except by several species in eastern Brazil and S Amazon Brazil, and two extends to E Andes of Peru, with many spp. present in Brazil (16, 5 endemics), mostly in white-sand areas of the Amazon Basin.

THE ‘VANGUIEREAE ALLIANCE’

At Vanguiereae Alliance, Crossopterygeae (1/1, tropical and S Africa), Jackieae (1/1, Malay Peninsula, Borneo), Scyphiphoreae (1/1, Mangrove shrub, coastal areas in SE Asia, West Malesia, tropical Australia and New Caledonia) and Trailliaedoxeae (1/1, SW China) and Glionnetia (1/1, Seychelles) may be sucessive sisters of Vanguerieae (24/650-680, Tropical regions in the Old World to tropical Australia, Melanesia and Polynesia) + Greeneeae (2/10, SE Asia, West Malesia) + Aleisanthieae (3/10, West Malesia) + Ixoreae; in this group, only Ixoreae occur in New World.

2.6 IXOROIDEAE ▸ TRIBE IXOREAE (1/c. 550) - a single genus.

93. Ixora L. Shrubs or small trees; flowers bisexual, usually fragrant, 4(5)-merous, few to many; fruit a drupe, usually red, spherical or 2-lobed, slightly fleshy or coriaceous, containing 1 or 2, 1-seeded, thin-walled pyrenes; calyx limb persistent; seeds frequently undeveloped, rusty brown in colour. 350 spp., cosmopolitan, with most of them in tropical Asia and islands of the southern Pacific, ca. 37-40 spp. in continental Africa, ca. 35 spp. in Madagascar, and 61 in New World, from Belize to Argentina (a single endemic to Mexico), east to French Guiana, Caribbean, 59 in South America; 38 spp. in Brazil, 20 endemics; a total of nine species were known to occur in the Atlantic Forest, three of them, from Rio de Janeiro and Bahia states, are a rare plants in Brazil, by Plantas Raras do Brasil’s book.

THE ‘COFFEA ALLIANCE’

Topology is Airospermeae +[Augusteae +[Alberteae +[Nematostylis clade +[ [Bertiereae + Coffeeae] + [[Burchellia + Galiniera] +[Mantalania + [Didymosalpinx + Monosalpinx] + [Octotropideae + [Sherbournieae + Cordiereae + Pavetteae + Gardenieae]]]]]]]]; clades Airospermeae (2/7, Malesia to New Guinea, Fiji), Nematostylis clade (2/6, Madagascar), Coffeeae (11/305-310, Tropical Africa, Madagascar, Mascarene Islands, Andaman Islands, tropical Asia), Burchellia + Galiniera (2/3, tropical Africa, Madagascar), Mantalania clade (2/4, Madagascar), Didymosalpinx + Monosalpinx (2/5, tropical Africa), Octotropideae (27/c. 110, tropical Africa, Madagascar, Mascarene Islands, Seychelles, India, Sri Lanka, SE Asia, Malesia, with their highest diversity in tropical Africa and Madagascar), Sherbournieae (4/c. 65, tropical and subtropical Africa) and Pavetteae (c 17/c 640, tropical regions in the Old World) do not occur in South America

2.7 IXOROIDEAE ▸ TRIBE BERTIEREAE (1/c. 55) - a single genus.

94. Bertiera Aubl. Small trees or shrubs, rarely lianas; leaves shortly petiolate; petioles normally channelled above; flowers moderate in size or small, borne in terminal, spicate thyrsoid panicles, scorpioid cymes or spherical compact heads; fruit indehiscent, ovoid or globose, often coriaceous, sometimes bluish; seeds many, red-brown to black, angular, rugulose or granulose. 55 spp. mostly present in Africa, Madagascar and Mascarene Islands, and with 10 spp. in the Neotropics (9 in South America), Mexico to SE Brazil (3, none endemics).

2.8 IXOROIDEAE ▸ TRIBE AUGUSTEAE (2/c 85) - outsider Wendlandia (c 80; NE Africa, Iraq to Malesia, tropical Australia).

95. Augusta Pohl. 4 spp. in two subgenera:

§ subg. Augusta ▸ only one sp., A. longifolia (Spreng.) Rehder, endemic to Brazil, with two varieties: var. longifolia, occurring in the torrents and small rivers of the savannas of C Brazil (cerrado), and var. parviflora (Pohl) Delprete, occurring in the torrents and small rivers of the Atlantic Forest in the state of Rio de Janeiro.

§ subg. Lindenia ▸ three spp., A. rivalis (Benth.) J.H. Kirkbr. from Mexico to Colombia, and A. austrocaledonica (Brongn.) J.H. Kirkbr. from New Caledonia and A. vitiensis (Seem.) J.H. Kirkbr., endemic to Fiji.

2.9 IXOROIDEAE ▸ TRIBE GARDENIEAE (c 54/580–605) - 49 outsiders, 17 only in mainland Africa, 21 only from tropical and temperate Asia from India to Indonesia, one only in E Australia, three from tropical Asia to Oceania islands, one from Africa to Madagascar, one only in Madagascar; the others are Gardenia (140–150; tropical and subtropical regions in the Old World to islands in the Pacific), Rothmannia (40–45; tropical and S Africa to Seychelles and the Malay Peninsula), Aidia (50–55; tropical regions in the Old World), Casasia (11; Central America, the Caribbean), Catunaregam (12; tropical Africa, tropical Asia).

96. Genipa L. Medium to large tress, fruit sweet fleshy. Three spp., G. americana L., in over trop. America (edible fruits), G. infundibuliformis Zappi & Semir, SE Brazil, in three localities, one in coast São Paulo state, and two, distant, in forests Minas Gerais state, and G. spruceana Steyerm. in over N South America.

97. Randia L. Shrubs to small trees, often lianas, 1 – 3 spiny, corolla mainly white. 112 spp., Florida, Texas, Mexico to Argentina, east to French Guiana, Caribbean, in that inhabit evergreen and deciduous forest from sea level up to 3,300 m. 32 spp. in South America, 7 in Brazil, two endemics.

98. Rosenbergiodendron Fagerl. Shrubs or small trees, large with nocturnal flowers up to 20 cm long. 4 spp. in Panamá, Trinidad and French Guiana to Bolivia, Brazil (three spp., none endemics), Paraguay.

99. Sphinctanthus Benth. Shrubs or small trees; hermaphroditic flowers, cream-white, white, yellow, yellowish orange to bright orange corollas, corolla tube with a ring of hairs at the middle or near the base inside and corolla lobes contorted to the left. 9 spp., restricted of South America up to SE Brazil (7, three endemics); S. fluviidulcis Delprete & C. H. Perss is known by only two individual plants.

100. Stenosepala C.H.Perss. Shrubs or small trees, dioecious. One sp., S. hirsuta C.H. Perss., from Panamá to W Colombia.

101. Tocoyena Aubl. Shrubs or small trees. 21 spp., Honduras to Paraguay, east to French Guiana and Cuba; 20 spp. in South America, 13 in Brazil, 6 endemics.

2.10 IXOROIDEAE ▸ TRIBE CORDIEREAE (13/c 135) - outsider Glossostipula (3; S Mexico, Guatemala).

102. Agouticarpa C.H. Perss. Dioecious shrubs, treelets or trees; inflorescences terminal, heterogeneous, sessile or pedunculate, male inflorescences 3–16-flowered, thyrse-like or a 3-flowered dichasium, female inflorescences usually 1 to 3-flowered dichasium; corollas usually salverform, rarely infundibuliform; fruits globose or rarely somewhat vertically compressed, many-seeded. 7 spp. from N South America (two up to Central America), centered in Ecuador (6), only the amazon widely A. curviflora (Dwyer) C.H. Perss. in Brazil.

103. Alibertia A.Rich. ex DC. Shrubs or small trees, terminal inflorescence, corolla hypocrateriform, with or cream. 19 spp., Mexico to Paraguay and Argentina, east to French Guiana, Caribbean, 16 in South America, 10 in Brazil, one endemic.

104. Amaioua Aubl. Shrubs or small trees. 11 spp., Mexico to Bolivia, east to French Guiana, Caribbean; 10 spp. in South America, 6 in Brazil, 4 endemics.

105. Botryarrhaena Ducke. Shrubs or small tress. Two spp., Venezuela (one endemic) to Peru and N Brazil (only B. pendula Ducke), mainly centered in savanas of Colombia and Venezuela.

106. Cordiera A.Rich. ex DC. 26 spp., from South America (three up to Central America), 22 in Brazil, 11 endemics.

107. Duroia L.f. Shrubs or small trees, dioecious. 37 spp., one in Costa Rica and remaining in South America, inc. Atlantic Forest of Brazil (24, 5 endemics); three spp. from northern South America, are myrmecophites.

108. Kutchubaea Fisch. ex DC. Trees medium sized to tall, often emergent in canopy forest, resinous in young branches. 13 spp., forests of N South America; 7 spp. in Brazil, two endemics.

109. Melanopsidium Colla. Dioecious shrubs 1.5-5 m tall; bark thin, reddish brown; leaves opposite, petiolate; inflorescences terminal and solitary; both pistillate and staminate inflorescences corymbose, many-flowered; corolla short-campanulate, 5-6(-7)-lobed, 5-8 x 3-4.5 mm, cream-white; tube campanulate. Only one sp., M. nigrum Colla, of Atlantic sandy coastal shrublands (restingas) from Bahia (Ilheus) to Rio de Janeiro states.

110. Riodocea Delprete. Dioecious trees; bark exfoliating in small, papery, longitudinal strips; inflorescences terminal; corolla campanulate, 13-15-merous (highest merosity in the Rubiaceae), 12-15-lobed, greenish yellow. Only one sp., R. pulcherrima Delprete, has been found naturally occurring only in riverine forest of the southern portion of the Rio Doce Valley, in Bahia and Espírito Santo states.

111. Stachyarrhena Hook.f. Shrubs or small trees, dioecious. 9 spp., one in Panamá and 8 in South America, up to Atlantic Forest of Brazil (7 in Brazil, 4 endemics).

3. SUBFAMILY RUBIOIDEAE (193/8.270–8.460) ▸ the sucessive most basal at Rubioideae, Colletoecemateae (1/2, Central Africa) and Ophiorrhizeae (6/350–355, tropical Asia to Melanesia), not occurs in South America; [[Lasiantheae + Perameae] + [Coussareeae + [”Spermacoceae alliance” + ”Psychotrieae alliance”]]].

3.1 RUBIOIDEAE ▸ TRIBE UROPHYLLEAE (c 14?/190–250) - outsiders Temnopteryx (1; Central Africa), Urophyllum (85–150; tropical regions in the Old World and northwards to China and Japan), Pentaloncha (3; tropical West and Central Africa), Pauridiantha (c 30; tropical Africa), Praravinia (c 50; Borneo, Philippines, Sulawesi), Antherostele (4; Philippines), Crobylanthe (1; Borneo), Didymopogon (1; Sumatra), Lepidostoma (1; Sumatra), Rhaphidura (1; Borneo), Rhipidantha (1; Uluguru Mountains in Tanzania), Stichianthus (1; Borneo).

112. Amphidasya Standl. Herbs, shrurbs or shrubs, often scandent or subscandent. 13 spp., Costa Rica to Peru, Venezuela, only one in Brazil, A. neblinae Steyerm., occurring in Mount Neblina in Amazonas state; South America has 12 spp.

113. Raritebe Wernham. Shrubs or small trees. Two spp. from Costa Rica to N Peru.

3.2 RUBIOIDEAE ▸ TRIBE LASIANTHEAE (c 4/c 210) - outsiders Lasianthus (c 170; tropical Africa, tropical Asia and eastwards to tropical Australia), Trichostachys (14; tropical Africa), Saldinia (22; Madagascar).

114. Lasianthus Jack. Shrubs or rarely small trees, rarely foetid, glabrous to hairy or strigose; leaves opposite; flowers sometimes heterostylous, mostly small, in sessile axillary fascicles or glomerules or less often in pedunculate, simple or branched inflorescences; fruits succulent, very often blue but sometimes pink, purple, white or black; seeds narrowly oblong, curved, with membranous testa and fleshy albumen. 170 spp. occurring in the Old World, and three spp. in the Neotropics, two in Dominican Republic and L. panamensis (Dwyer) Robbr. in Costa Rica, Panamá and adjacent Colombia.

115. Ronabea Aubl. Three spp., R. emetica (L.f.) A. Rich. from Nicaragua to western Ecuador and N Brazil, R. isanae (J.H. Kirkbr.) C.M. Taylor endemic to Amazonian NW Brazil, and R. latifolia Aubl. from Belize to Ecuador, the Guianas to NE Brazil.

3.3 RUBIOIDEAE ▸ TRIBE PERAMEAE (1/12) - a single genus.

116. Perama Aubl. Herbs (sometimes very small, 10 cm) or erect or prostrate, small or showy, in dense populations very divergent of Rubiaceous pattern. 12 spp. tropical South America except Argentina, centered in savannas of Guyana and Brazilian Shield (11, 7 endemics; P. wurdackii Steyerm. only in Peru), P. hirsuta Aubl. up to Trinidad & Tobago.

3.4 RUBIOIDEAE ▸ TRIBE COUSSAREAE (8/410–420) - all genera in South America.

117. Bradea Standl. Herbs, shrubs or subshrubs up to 2 m tall. 6 spp., of rocky outcrops in Atlantic forest understory in Rio de Janeiro, Minas Gerais and Espírito Santo states.

118. Coccocypselum P.Browne. Prostrate, erect or scandent herbs, flowers in cephalia; many spp. have blue fruits and pinkish flowers. 24 spp. from Mexico to Argentina, Paraguay, Brazil (16, 8 endemics) and Guianas; all species occur in South America.

119. Coussarea Aubl. Shrubs or small trees. 121 spp., S. Mexico to Argentina, Brazil (57, 32 endemics), Guianas, Caribbean; 111 spp. occur in South America; 5 spp., 4 in Rio de Janeiro and one in Bahia state, are rare plants in Brazil, by Plantas Raras do Brasil’s book.

120. Cruckshanksia Hook. & Arn. Herbs often with calycophylls. 7 spp. from Chile, Argentina.

121. Declieuxia Kunth. Herbs or suffrutex plants up to 1 m tall, sometimes with xylopodium. 28 spp., all restricted of Brazil, except two up to Bolivia, one to Venezuela and Guyana, and D. fruticosa (Willd. ex Roem. & Schult.) Kuntze widely distributed neotropics.

122. Faramea Aubl. Shrubs or small trees; inflorescence terminal, rarely axillar. 185 spp., from Mexico to Uruguay, east to French Guiana, Caribbean; 165 spp. in South America, 90 in Brazil, 55 endemics.

123. Hindsia Benth. ex Lindl. 11 spp. of shrubs and subshrubs from Atlantic Forests and Atlantic sandy coastal shrublands (restingas) from states in SE Brazil, three of them, from Rio de Janeiro and Bahia states, are rare plants in Brazil, by Plantas Raras do Brasil’s book.

124. Heterophyllaea Hook.f. Two spp. from Peru to N. Argentina, absent in Chile.

125. Oreopolus Schltdl. Herbs, dense cushions. Only one sp., O. glacialis (Poepp.) Ricardi, in open mountain sites in Argentina and Chile.

126. Standleya Brade. Small herbs. 5 spp., endemic to Espírito Santo and Rio de Janeiro states in E Brazil.

THE ‘SPERMACOCEAE ALLIANCE’

[Danaideae + [[Knoxieae + Spermacoceae] + [Anthospermeae + [Argostemmateae + [Dunnieae + [Paederieae + [Putorieae + [Theligonum + Rubieae]]]]]]]]; six lineages among this clade do not occur in South America: Danaideae (3/c 60, tropical Africa and Madagascar), Knoxieae (15/c 130, W and S tropical Africa and Madagascar, Socotra, Knoxia also in tropical Asia), Argostemmateae (3/c 220, tropical regions in the Old World), Dunnieae (3/7 or 10, N India, S China, Borneo, Sulawesi), Putorieae (1/c 35, Canary Islands, Mediterranean, SW Asia to NW India) and Theligoneae (1/4, Macaronesia, Mediterranean, SW China (inc. Taiwan), Japan).

3.5 RUBIOIDEAE ▸ TRIBE SPERMACOCEAE (69/1.380–1.410) - outsiders Hedyotis (c 130; tropical and subtropical Asia to New Guinea, island in NW Pacific), Mexotis (4; Mexico), Martensianthus (6; Mexico), Terrellianthus (1; S Mexico, Central America), Agathisanthemum (4; tropical Africa, the Comoros), Kohautia (c 35; tropical and subtropical Africa, Arabian Peninsula to India, Thailand, tropical Australia), Manostachya (3; tropical Africa), Lathraeocarpa (2; Madagascar), Gomphocalyx (1; Madagascar), Phylohydrax (2; tropical East Africa, Madagascar), Amphiasma (7; tropical and SW Africa), Pentanopsis (2; NE tropical Africa), Conostomium (5; tropical East Africa), Dentella (8; tropical Asia to tropical Australia and New Caledonia), Pentodon (2; tropical and subtropical regions in Africa, Seychelles, Arabian Peninsula, tropical and subtropical America), Dibrachionostylus (1; tropical East Africa), Neanotis (c 30; tropical Asia to tropical Australia, islands in the Pacific), Involucrella (2; SE Asia), Debia (4; SE Asia, Andaman Islands, Philippine Islands), Kadua (c 30; French Polynesia, Hawai), Leptopetalum (5; tropical Asia to tropical Australia, islands in the Pacific), Scleromitrion (c 10; tropical Asia to tropical Australia, islands in the Pacific), Exallage (15; tropical Asia to tropical Australia, islands in the Pacific), Dimetia (7; tropical Asia), Cordylostigma (9; tropical and S Africa, Madagascar), Phialiphora (2; NW Madagascar), Stenaria (5; S Canada, U.S.A.), Thecorchus (1; tropical Africa), Hedythyrsus (2; tropical Africa), Pseudonesohedyotis (1; Tanzania), Mitrasacmopsis (1; tropical East Africa, Madagascar), Astiella (1; Madagascar), Amphistemon (2; Madagascar), Thamnoldenlandia (1; Madagascar), Synaptantha (2; tropical and subtropical regions in Australia), Houstonia (c 30; S Canada, U.S.A., Mexico), Hydrophylax (1; tropical and subtropical Africa, Madagascar, India, Thailand), Bouvardia (c 50; SW U.S.A., Mexico, Central America), Nesohedyotis (1; St. Helena), Pleiocraterium (4; S India, Sri Lanka, Sumatra), Carterella (1; Baja California in NW Mexico), Dolichometra (1; Tanzania), Lucya (1; Caribbean), Phyllocrater (1; Borneo), Polyura (1; Assam), Stephanococcus (1; tropical Africa), Micrasepalum (2; Cuba; Hispaniola), Spiradiclis (38; India, SW China, SE Asia, Java), Crusea (15; SW U.S.A., Mexico, Central America), Leptomischus (7; SE Asia).

127. Anthospermopsis (K.Schum.) J.H.Kirkbr. (off Mitracarpus). Small shrub. Only one spp., A. catechosperma (K.Schum.) J.H.Kirkbr., a rare plant in Brazil, by Plantas Raras do Brasil’s book, endemic to Brazil, in Atlantic sandy coastal shrublands (restingas) between Salvador and Camaçari, Bahia state.

128. Arcytophyllum Willd. (inc. Hedyotis p.p.) Prostrate or erect herbs, often shrubs, many branched. 18 spp. from SC. U.S.A. to Bolivia and Venezuela, 15 in South America.

129. Carajasia R.M. Salas, E.L. Cabral & Dessein. Herbs 2–10 cm tall, perennial, with main branches erect; stem tetragonal, glabrous, papillate on angles, wine-colored; flowering branches with two axillary flowers on each node; each flower subtended by 2 foliaceous bracts; bracteoles inconspicuous; flowers sessile, homostylous; hypanthium obovoid, pubescent; dry fruit obovoid. Only one sp., C. cangae R.M. Salas, E.L. Cabral & Dessein, known only of Canaã dos Carajás municipality, state of Pará, Brazil; they inhabit rocky-ferriginous grasslands (cangas) only on the top of the Carajás mountain range.

130. Denscantia E.L.Cabral & Bacigalupo. Scandent habit, the tubular stipule sheath, thyrsi or pleiothyrsi inflorescences, isostylous flowers, pollen grains with multiple endoaperture, and complanate seeds. 5 spp., D. calcicola R. M. Salas & E. L. Cabral in seasonally dry region inside the dry seasonal scrubland of NE Brazil (caatinga) biome of Bahia state Brazil, and 4 remaining Atlantic Forest biome of Brazil, in areas of Atlantic sandy coastal shrublands (restingas), three endemics to Bahia state and two of them rare plants in Brazil, by Plantas Raras do Brasil’s book.

131. Diadorimia J.A.M.Carmo, Florentín & R.M.Salas (off Psyllocarpus). Subshrubs, cespitose habit arising from a well-developed, woody subterranean system, triangular stipules with margins bearing four small lobes, heterostylous flowers subtended by a pair of hyaline bracteoles. Only one sp., D. densifolia (Zappi & Calió) J.A.M.Carmo, Florentín & R.M.Salas, endemic to Cipó Highs in Minas Gerais state, SE Brazil.

132. Diodia L. Annual or perennial herbs to small shrubs, erect or scandent, woody at base, may branched. 18 spp., 12 in trop. Africa and 6 in C & E U.S.A. to South America (3), two very scattreed and one endemic to Brazil, from Tocantins state, as a rare plant in Brazil, by Plantas Raras do Brasil’s book.

133. Edrastima Raf. Annual herbs. 5 spp., 4 in Africa and E. uniflora (L.) Raf., disjunct in C & E U.S.A., Caribbean, SE Brazil to NE Argentina.

134. Emmeorhiza Pohl ex Endl. Herbs to tiny subshrubs, erect or scandent. Only one sp., E. umbellata (Spreng.) K. Schuman Trinidad to Paraguay, Guianas, Brazil, disturbed areas, forests, riversides, shrublands at 0–2,500 m elevation range.

135. Ernodea Sw. 8 spp., Florida to Caribbean (highly centered in Bahamas), with E, littoralis Sw. up SE. Mexico and NW Colombia, in coastal areas in sand dunes, near mangrove communities, limestone, disturbed open areas, or pinelands.

136. Galianthe Griseb. Herbs, sometimes with xylopodium, mainly in open places. 53 spp. from Peru to S Brazil (37, 22 endemics, 7 of them, from several states, are rare plants in Brazil, by Plantas Raras do Brasil’s book), Bolivia, Paraguay and Uruguay, six up to N South America and Central America, and one endemic to Mexico.

137. Hexasepalum Bartl. ex DC. Subshrubs or shrubs, perennial, rarely annual herbs or somewhat fleshy plants; stems cylindrical or quadrangular, erect or decumbent, sometimes creeping and irregularly branched forming dense mats; leaves linear, ovate, elliptic, rarely slightly succulent. 14 spp., one in W Africa and 14 in New World, 11 in South America, 10 in Brazil, 5 endemics.

138. Leptoscela Hook.f. Herb, about 30 cm. Only one sp., L. ruellioides (Willd. ex Roem. & Schult.) J.H. Kirkbr. & Delprete, widely distributed in NE Brazil from 50 – 1,000 m of altitude range.

139. Manettia Mutis ex L. Herbs with tiny stems. 118 spp. from Mexico to Uruguay, Brazil (27, 20 endemics), Guianas, Caribbean; 107 spp. in South America; three spp., all from Rio de Janeiro state, are rare plants in Brazil, by Plantas Raras do Brasil’s book.

140. Merumea Steyerm. Two spp. endemics to Cerro Sipapo of the Venezuelan Guiana and the Merume Mountains of Guyana, 1,100 - 1,600 m elevation range.

141. Mitracarpus Zucc. (exc. Anthospermopsis) Erect or prostrate annual or perennial herbs with 4-angled stems; flowers not heterostylous or only slightly so, in dense, spherical, sessile, terminal or axillary heads; fruit a thin circumscissile capsule; seeds oblong or globose, endosperm fleshy. 69 spp. from Mexico to Uruguay, Caribbean, 37 in South America, highly centered in highlands of Brazil (29, 22 endemics, one of them, from Minas Gerais state, is a rare plant in Brazil, by Plantas Raras do Brasil’s book).

142. Oldenlandiopsis Terrell & W.H.Lewis. Only one sp., O. callitrichoides (Griseb.) Terrell & W.H.Lewis, disjunct spp., S. Mexico to Central America, Caribbean, NW Argentina.

143. Oldenlandia L. (exc. Edrastima p.p., Hedyotis p.p.) Perennial or annual herbs, erect or decumbent. Cosmopolitan, 197 spp., 18 in New World, 7 in South America, four in Brazil, one endemic.

144. Paganuccia R.M.Salas. Herbs to subshrubs, stems quadrangular, with flowers in globbose inflorescences. Only one sp., P. icatuensis R.M.Salas, known only so far from two collections from the continental dunes in the São Francisco River basin, Bahia, north-eastern Brazil.

145. Planaltina R.M.Salas & E.L.Cabral. Suffrutices with erect stems; leaves opposite, quadrangular stems; inflorescences glomerule, terminal or axillary; flowers sessile or pedicellate; corolla infundibuliforme, white; capsules indehiscent. 4 spp. in savannas of C Brazil (cerrado) at Distrito Federal, Goiás (all species) and Minas Gerais state in center Brazil (800 – 1,200 m elevation range).

146. Psyllocarpus Mart. & Zucc. (exc. Diadorimia) Herbs, sometimes ericoids. 14 spp. widely distributed in Brazil, in two disjunct areas: savannas in central highlands of Brazil (Distrito Federal, Goiás and Minas Gerais) and white‐sands vegetation in Amazon valley; four spp., in four diferent states, are rare plants in Brazil, by Plantas Raras do Brasil’s book.

147. Richardia L. Annual or more commonly perennial herbs from fibrous roots or typically from a long, sparingly branched taproot, occasionally with adventitious roots from those nodes touching the soil; stems terete, decumbent and often forming small, matted cushions, or sprawling to infrequently erect; inflorescence a pedunculate, terminal capitulum. 16 spp. in two sections:

§ sect. Richardia ▸ 4 spp., R. lomensis (Krause) Standley endemic to SW Peru, two widely distributed in South America except for the Amazon Basin, and one occurring from the S U.S.A. and Cuba to NW South America.

§ sect. Asterophyton ▸ 12 spp., 7 native to South America primarily from Peruvian Andes and central Bolivia (2 endemic) to SE Brazil (5 of then) and Uruguay, the remainder 5 endemics to North America with 1 species from Mexico to Louisiana and 3 in Cuba; R. schumannii Lewis & Oliver is endemic to the grasslands of south-eastern Brazil. R. pedicellata (K. Schumm) O. Ktze. is endemic to the shrub savannas and dry grasslands of SE Brazil and southern Paraguay at elevations up to 1,000 m.

148. Schwendenera K.Schum. Only one sp., S. tetrapyxis K. Schum., disjunct from Pará and São Paulo states in Brazil.

149. Spermacoce L. (inc. Borreria, Diacrodon). Annual or perennial herbs, up to low shrubs, erect or prostrate, stems quadrangular. c. 300 spp., 157 in New World, 128 in South America, including Chile, 92 spp. in Brazil, 49 endemics, 12 of them, in several states, are rare plants in Brazil, by Plantas Raras do Brasil’s book.

150. Staelia Cham. & Schltdl. Herbs from dense small cushion to free stemmed plants, sometimes bonsai-like. 18 spp. from Bolivia, Brazil (16, 12 endemics, one of them, from Tocantins state, is a rare plant in Brazil, by Plantas Raras do Brasil’s book) to N. Argentina, Paraguay and Uruguay.

151. Tobagoa Urb. Herbs prostrate or erects, foetid. Two spp., from Panamá, Colombia, N Venezuela and Tobago.

3.6 RUBIOIDEAE ▸ TRIBE ANTHOSPERMEAE (10/210–215) - outsiders Anthospermum (c 40; Africa, Madagascar), Nenax (9–11; S Namibia, South Africa), Galopina (4; S Africa to Malawi), Phyllis (2; Madeira, Canary Islands), Durringtonia (1; Queensland, New South Wales), Pomax (1; Australia), Carpacoce (7; W and E Cape), Opercularia (17–18; W Australia, South Australia to Queensland, Tasmania).

152. Coprosma J.R.Forst. & G.Forst. 109 spp., mainly Malesia to Pacific, and two from Juan Fernández Is., Chile.

153. Leptostigma Arn. Prostrate to scandent herbs, stoloniferous, stems tiny-fleshy. 7 spp., one in New Zealand, two in Australia, and 4 in W South America, from Colombia to Chile and Argentina, mainly in paramos.

154. Nertera Banks ex Sol. Small herbs, prostrate or decumbent, stoloniferous with tiny flexible stems. 15 spp. worldwide, only one in New World, N. granadensis (Mutis ex L.f.) Druce, which occur in Madagascar, Asia to Pacific, Caribbean, Mexico to Chile and Venezuela and Subantarctic Is.

3.7 RUBIOIDEAE ▸ TRIBE PAEDERIEAE (6/130–135) - outsiders Saprosma (45–50; tropical Asia), Spermadictyon (1; India), Leptodermis (c 50; the Himalayas to Japan), Serissa (1; S China), Pseudopyxis (3; Japan).

155. Paederia L. 30 spp., with 16 spp. in Asia, 12 spp. in Africa and Madagascar, and two spp. in tropical America, P. ciliata (Bartl. ex DC.) Standl., that occurs in SW Mexico, and P. brasiliensis (Hook. f.) Puff, found in Brazil (Mato Grosso, Tocantins and Ceará states), Peru, Bolivia, Argentina and Paraguay.

3.8 RUBIOIDEAE ▸ TRIBE RUBIEAE (4/c 700) - outsiders Kelloggia (2; S China, Bhutan; SW U.S.A., NW Mexico); Didymaea (8; Central America), Rubia (75–80; Macaronesia, Mediterranean, Africa, temperate Asia, North America).

156. Galium L. Prostrate to erect herbs, glabrous or pubescent, monoecious, dioecious, often polygamomonoecious, tiny stems. 400 spp., worldwide, 170 spp., in New World, about 50 spp. in the Neotropics, 72 in South America, 25 in Brazil, 10 endemics, centered in Santa Catarina state.

THE “PSYCHOTRIEAE ALLIANCE”

[Craterispermeae+Schradereae+[Gaertnereae+[Mitchelleae+Morindeae]]+Prismatomerideae+Psychotrieae]. Among this clade, three small lineages do not occur in South America: Craterispermeae (1/16, tropical Africa, Madagascar, Seychelles), Prismatomerideae (4/26, Sri Lanka, Assam, SE Asia, Hainan, West Malesia) and Mitchelleae (2/14, China (inc. Taiwan), S Korea, Japan; North America).

3.9 RUBIOIDEAE ▸ TRIBE SCHRADEREAE (3/c 60) - outsiders Lecananthus (3; West Malesia), Leucocodon (1; Sri Lanka).

157. Schradera Vahl. Scandent above canopy trees, rarely epiphytics, tree in a single South American species; adventive roots in juvenile individuous, stems terete in adult members. 59 spp., 16 in pen. Thailand to New Guinea, and 43 in New World, 36 in South America, 4 in Brazil, two endemics.

3.10 RUBIOIDEAE ▸ TRIBE GAERTNEREAE (2/c 85) - outsider Gaertnera (c 70; tropical regions in the Old World).

158. Pagamea Aubl. Shrubs or small trees, often ‘woody’ herbs. 26 spp. in over tropical South America, centered in Amazonia, 16 in Brazil (7 endemics), most of them occurring in white-sand areas.

3.11 RUBIOIDEAE ▸ TRIBE MORINDEAE (c 6/c 170) - outsiders Gynochthodes (c. 20; SE Asia, Malesia and eastwards to tropical Australia and Melanesia), Coelopyrena (1; E Malesia), Coelospermum (9; SE Asia, Malesia and eastwards to E Queensland and New Caledonia), Siphonandrium (1; New Guinea).

159. Appunia Hook.f. Lianas, shrubs or small trees. 13 spp., S. Mexico to Bolivia, Brazil (6, one endemic), Guianas; 12 spp. in South America.

160. Morinda L. Lianas, shrubs or small trees. 130 spp. trop. & subtrop., nine spp. in New World, Florida, Mexico to Ecuador and Venezuela; 6 spp. in South America.

3.12 RUBIOIDEAE ▸ TRIBE PSYCHOTRIEAE (26/3.480–3.580) - outsiders Schizocolea (2; West Africa); Gillespiea (1; Fiji), Hedstromia (1; Fiji), Cremocarpon (9; Madagascar, the Comoros), Pyragra (2; Madagascar), Amaracarpus (c 30; Seychelles, tropical Asia to tropical Australia and Solomon Islands), Dolianthus (13; mountains in Papua New Guinea), Myrmecodia (27; Malesia to Fiji), Hydnophytum (c 95; tropical Asia, with their highest diversity on New Guinea), Squamellaria (4; Fiji), Myrmephytum (5; Philippines, Sulawesi, western New Guinea), Anthorrhiza (9; New Guinea), Calycosia (8; Polynesia), Hymenocoleus (12; tropical Africa), Chassalia (c 110; tropical regions in the Old World, with their largest diversity on Madagascar), Puffia (1; Madagascar), Peripeplus (1; Central Africa), Streblosa (c 25; West Malesia). Anthorrhiza, Hydnophytum, Myrmecodia, Myrmephytum and Squamellaria are myrmecophilous with ant colonies living in the hollow stems.

161. Carapichea Aubl. (exc. Notopleura p.p., inc. Psychotria p.p.) 22 spp. from South America, two of them up to Nicaragua, 17 in Brazil, 5 endemics. C. ipecacuanha (Brot.) L. Andersson is the famous Brazilian ipecacuanha, a medicinal plant widely distributed in Brazil, Colombia and Central America, but increasingly rare due to excessive exploitation.

162. Coccochondra Rauschert. Only one sp., C. laevis (Steyerm.) Rauschert, from endemic to Pantepui Life Zone (Serrania Maigualida, Serrania del Peru, Cerro Yutajé), S. Venezuela, at 2,000 m elevation range.

163. Eumachia C.Wright. 27 spp., Mexico to Paraguay, Guianas, Caribbean, 16 in Brazil, only two endemics; 22 spp. occur in South America.

164. Geophila D.Don. Perennial forest floor herbs, mostly with slender creeping stems rooting at nodes; leaves opposite, with mostly long petioles; flowers bisexual, sometimes heterostylous, mostly in terminal umbels or sometimes solitary. 30 spp., trop. to S. China, 9 in New World, 7 in South America, six in Brazil, none endemics.

165. Notopleura (Hook.f.) Bremek. (inc. Carapichea p.p.) Herbs terrestrial (mainly unbranched) or epiphytic, subshrubs, shrubs or small trees, often scandent, fleshy to succulents. 102 spp., Mexico to Brazil (3, N. bahiensis C.M. Taylor endemic), 83 in South America, also in Bolivia, Guianas, Caribbean.

166. Palicourea Aubl. Shrubs or medium sized trees, sometimes basal burls, inflorescence terminal, pseudopaniculate; several species has blue fruits. 491 spp. from over Neotropics, except Chile and Uruguay, centered in Colombia; 429 spp. in South America, 91 in Brazil, 38 endemics. P. corymbifera (Müll.Arg.) Standl. from northern South America is a myrmecophite.

167. Psychotria L. (inc. Margaritopsis, exc. Capaichea p.p.) Shrubs, trees small to medium sized, often epiphytic, sometimes with bluish fruits. 1,638 spp. worldwide, c. 1,200 in Old World, 656 in New World (absent only by Chile and Canada); 471 spp. in South America, 221 in Brazil, 120 endemics.

168. Rudgea Salisb. Shrubs or small trees; inflorescence terminal, often axillar, erect or pendulous, paniculate, corymbose, capitate or cymose; corolla hypocrateriform, white. 174 spp. from Mexico to Argentina, 164 in South America, inc. Brazil (77, 48 endemics), Guianas, Caribbean.

Only R. quisquiliae Bruniera & Torres-Leite and R. axilliflora Bruniera & Torres-Leite from Espírito Santo state in SE Brazil has axillary inflorescences within this genus; besides the size of the leaf-blades, R. quisquiliae and R. macrophylla also share shortly petiolate leaves (petioles up to 10–12 mm long), obovate to elliptic blades, cordate at base, and 13–18 pairs of secondary veins, seem to be adapted to capture organic material falling from the forest canopy possibly as a nutritional resource, as seen in forest plants with similar habit, such as Agrostistachys borneensis Beccari in the Euphorbiaceae.

GENTIANACEAE

§ MYCOHETEROTROPHICS (Arachnitis – TRIURIDACEAE – BURMANNIACEAE – ORCHIDACEAE – Voyria - Voyriella - Monotropa)

Genera/spp. 103/1,650-1,700 Distribution cosmopolitan except polar and arid areas, with their largest diversity in temperate and subtropical regions and on tropical mountains; Voyria: tropical America, tropical West Africa. Habit usually bisexual (rarely polygamomonoecious), perennial, biennial or annual herbs (sometimes shrubs, rarely lianas or trees). Some genera are partially mycoheterotrophic. A few clades (Cotylanthera, Leiphaimos, Voyria, and Voyriella) consisting of achlorophyllous holoparasitic mycotrophs. Other genera are cultivated for ornamental purposes (e.g. Eustoma, Exacum). Several spp. occurring in the Neotropics are narrow endemics.

SYSTEMATIC six tribes, Exaceae (8/c 150, tropical regions in the Old World, S Australia, Tasmania, New Zealand, with their highest diversity in Madagascar) only in Old World, mainly Madagascar; 553 spp. in South America, 7/10 are only Gentianella, Halenia, Symbolanthus or Macrocarpaea.

1.1 TRIBE SACCIFOLIEAE (5/16–20) ▸ all genera occur in South America.

1. Curtia Cham. & Schldtl. Erect annual herbs, usually not very conspicuous; inflorescence a lax cyme; flowers 5-merous; corolla white, yellow, pale lilac, pink, or purplish, sometimes hairy on the inside, funnel- to salver-shaped; corolla tubes about as long as corolla lobes. 8 spp., 7 in Brazil, three endemics, only C. tenella (Mart.) Cham. outside South America (curiously disjunct Brazil, Mexico and Central America).

2. Hockinia Gardner. Herb, annual; inflorescence with solitary flowers or as cymose monochasia; flowers 5-merous; corollas pale lilac, funnel shaped; corolla tube is longer than corolla lobes. Only one sp., H. montana Gardner, in Serra dos Orgãos and adjacent mountains in Rio de Janeiro state in SE Brazil.

3. Saccifolium Maguire and Pires. Shrub up to 60 cm tall, branches with lots of corky bark; leaves tightly arranged, opposite or possibly alternate, sac-shaped, partly translucent, and with glands on the lower surface, not found in any other plant; flowers 5-merous, solitary, in the axils of leaves; corollas tubular, widened in the middle; fruits never collected as mature. Only one sp., S. bandeirae Maguire & Pires, endemic to Pantepui Life Zone, known from the single mountain of Mount Neblina in Venezuela and Amazonas state in northern Brazil, 2,700 – 2,800 m elevation range.

4. Tapeinostemon Benth. Perennial herbs, upright and sometimes slightly woody at the base of the plant; flowers 5-merous, in lax cymes or heads; corolla white, yellow, or orange, salver- to funnelshaped or tubular, with relatively short lobes. 8 spp. in N South America, 3 in Brazil, none endemics.

5. Voyriella Miq. Saprophytic herbs, 2-10 cm high, completely white; leaves 3-4 mm long; inflorescence a dense, globose, bracteate, mostly many flowered, terminal or axillary cyme; corolla white, campanulate, 5-6 mm long (hardly exceeding the calyx), lobes ovate-triangular, 1-1.5 mm long, apex acute, recurved. Only one sp., V. parviflora (Miq.) Miq., lowlands of N South America and adjacent Panamá, mainly in Guianas, over Colombia, valleys of Negro and Madeira rivers in Brazil, Guianas, Pará and Amapá coasts, border Pará and Maranhão states in Brazil, and one collection in Mato Grosso state in Brazil.

1.2 TRIBE VOYRIEAE (1/19) ▸ a single genus.

6. Voyria Aubl. Saprophytic herbs; stems mostly simple, terete; leaves small and scale-like; inflorescence a terminal few-flowered, bifurcate cyme, or the plant having a solitary flower only; flowers variously coloured, (4-)5-merous; calyx tubular to campanulate, small; corolla salverform to funnelform, far exceeding the calyx. 23 spp., one in W tropical Africa and remaining 22 in tropical and subtropical America, 18 in South America; 13 in Brazil, V. obconica Progel from Atlantic Forest (known only from three collections) are endemics; V. parasitica (Schltdl. & Cham.) Ruyters & Maas from U.S.A. to Honduras and Caribbean, and two endemics of Panamá and Costa Rica do not occur in South America; lowland rain forest, creek forest, swamp forest, montane rain forest, hammock forest and Amazonian dry forest.

They grow in decaying leaf mold, on brown to black clayey to loamy soils. A few species, like V. aphylla (Jacq.) Pers., prefer much drier vegetation types like white sand savannas and savanna forests. V. parasitica, the species with the northernmost distribution, is often reported as growing on limestone substrate; all species are terrestrial, although it sometimes happens that plants grow on dead, decaying logs. Voyriella and Voyria are confined to elevations from sea level to 1,000 m; for a few species, e.g., V. aphylla, higher elevations (up to 1,800 m), are recorded. Simultaneouly, V. spruceana Benth. and V. aphylla have been found growing as epiphytes up to 30 m high in trees in Colombia.

Voyria has two well distinct subgenera: subg. Voyria (only V. caerulea Aubl. in Brazil) and subg. Leiphaimos (all remaining species in country).

1.3 TRIBE CHIRONIEAE (27/c 165) ▸ outsiders Bisgoeppertia (3; Caribbean), Blackstonia (4; Europe, Mediterranean), Chironia (c 30; tropical and S Africa, Madagascar), Schenkia (5; Australia, Tasmania, islands in W Pacific; Hawaii; Europe, Mediterranean, North Africa), Exaculum (1; Europe), Geniostemon (5; Mexico), Ixanthus (1; Canary Islands), Orphium (1; W Cape), Gyrandra (5; Mexico, Central America), Sabatia (21; E and C North America, Mexico, Central America, the Caribbean), Canscora (9–10; tropical regions in the Old World to tropical Australia), Cracosna (3; SE Asia), Duplipetala (2; Thailand, the Malay Peninsula), Hoppea (2; India, Sri Lanka, Burma), Microrphium (1; Thailand, the Malay Peninsula, Palawan), Phyllocyclus (5; Burma, S China, Thailand), Schinziella (1; tropical Africa); Xestaea (1; Panamá).

7. Centaurium Hill. (inc. Erythraea) 20 spp., cosmopolitan, three spp. in to South America: Argentina, Chile and Peru one endemic each; Chilean species is a important medicinal herb, frequent from the Pacific litoral to the Andean pre-Cordillera and from the Atacama Chiloe.

8. Cicendia Adans. Small, annual, filiform habit; rosette leaves are dioecious and the lanceolate cauline leaves are very small; single 4-merous flowers or scarselly cymes, corolla yellow. Two spp., C. filiformis (L.) Delarbre from Mediterranean and W Europe, and C. quadrangularis (Dombey ex Lam.) Griseb. disjunct in SW & W North America and South America from Ecuador to Argentina, also in Uruguay.

9. Coutoubea Aubl. Annual, rarely short-lived perennial, sometimes suffrutescent herbs; rosette leaves are absent and the cauline leaves are lanceolate; flowers 4-merous, white, corolla salver-shaped, are born in dense spikes. 5 spp. widely distributed in N South America, one of then up to Central America and Mexico; 4 spp. in Brazil, none endemics; savannas and weedy, open places.

10. Deianira Cham. & Schltdl. 7 spp., all in Brazil, 4 endemics and three up to E Bolivia; D. damazioi E.F.Guim. from Minas Gerais is rare plant in Brazil, by Plantas Raras do Brasil’s book.

11. Eustoma Salisb. Three spp., mainly in Mexico (two endemics), and E. exaltatum (L.) Salisb. ex G. Don from North America, Mexico, Central America, Caribbean, Trinidad & Tobago and Venezuela.

12. Symphyllophyton Gilg. (inc. Schultesia p.p.) 7 spp. endemics to Maranhão, Piauí, Bahia, Tocantins and Goiás states in N & C Brazil.

13. Schultesia Mart. (exc. Symphyllophyton p.p.) 21 spp., pantropical, minus Asia; 18 spp. in New World, 17 in Brazil (S. lisianthoides (Griseb.) Benth. & Hook. f. ex Hemsl. occur from Mexico to Venezuela and Colombia), some widely distributed, 10 endemics (five from Bahia and one in Piauí state are rares by Plantas Raras do Brasil’s book), mainly in savannas.

14. Zygostigma Griseb. Perennial herbs, thiny, erect; leaves oppsoite, sessile; inflorescences cymose or solitary; corolla hipocrateriform. Only one sp., Z. australe (Cham. & Schltdl.) Griseb., S Brazil, NE Argentina and Uruguay.

15. Zeltnera Mansion. Herbs, annuals, biennials or short-lived perennial, stems single or several from the base, branched along the main stem (sparingly to profusely); flowers witish or pinkish. 25 spp., highly centered in North America and Mexico, two up to Central America, and Z. quitensis (Kunth) G. Mans. in Mexico, Central America, Caribbean, Venezuela, Colombia, Ecuador, Peru.

1.4 TRIBE POTALIEAE (13/c 155) ▸ outsiders Congolanthus (1; tropical Africa), Djaloniella (1; Guinea), Faroa (c 20; tropical Africa), Karina (1; Congo), Oreonesion (1; Gabon), Pycnosphaera (1; tropical Africa), Urogentias (1; Tanzania); Anthocleista (c 50; tropical Africa, Madagascar, Mascarene Islands), Fagraea (60–70; S India, Sri Lanka to China, SE Asia, Malesia and islands in W Pacific, with their highest diversity on Borneo).

16. Enicostema Blume. Three spp., two in Madagascar, tropical Africa and Asia, and E. verticillatum (L.) Engl. ex Gilg from Central America, Caribbean, Trinidad & Tobago, Venezuela and Guyana.

17. Lisianthius P. Browne. 31 spp., almost totally confined to Mexico, Central America and the Greater Antilles (all eight Jamaica spp. are endemics), one of then, L. seemannii (Gnseh.)0. Kuntze, ranges into NW Colombia.

18. Neurotheca Salisb. ex Benth. & Hook.f. Three spp. in tropical Africa and western Madagascar, N. loeselioides (Spruce ex Progel) Baill. also in from N South America.

19. Potalia Aubl. 9 spp., one in Costa Rica, and remaining from Colombia to Bolivia and Brazil (5, none endemics; mainly Amazon rainforests).

1.5 TRIBE HELIEAE (19–22/c 185) ▸ outsider Zonanthus (1; Cuba).

20. Adenolisianthus Gilg. Shrubs or subshrubs, leaves crowded towards branch apices; inflorescences terminal, many-flowered, with long-stalked, monochasial branches (flowers one by one on one side of the branch); flowers 5-merous, horizontal or nodding; corolla broadly funnelshaped, greenish-yellow. Only one sp., A. arboreus (Spruce ex Progel) Gilg., in a small area in the river basins of Rio Negro and Rio Vaupes of Amazonas state in NW Brazil, S Venezuela and S Colombia in South America, in lowland, white-sand savanna.

21. Aripuana Struwe, Maas & V.A. Albert. Shrubs or small tree; leaves inflorescences terminal, many-flowered, with dichasial cymes; flowers 5-merous, erect; corolla narrowly salver-shaped, up to 7 cm long, thin, white, with spreading lobes. Only one, A. cullmaniorum Struwe, Maas & V.A. Albert, rare plant in Brazil, by Plantas Raras do Brasil’s book, known only from a small area along the river Aripuanan River in in the southeastern part of Amazonas state, Brazil.

22. Calolisianthus Gilg. (exc. Chelonanthus p.p.) Unbranched, perennial herbs, sometimes woody at the base; inflorescences terminal, usually few-flowered, with short, monochasial branches, with scale-like bracts; flowers 5-merous, horizontal or nodding, slightly zygomorphic; corolla showy, large, salvershaped to funnelshaped, blue, purple, or red, with rounded or acute lobes. 4 spp. endemics to SE Brazil, from mountains of the Brazilian Highlands (Pará to Paraná), in savannas, forests, and grasslands, especially rochy grassslands.

23. Celiantha Maguire. Herbs, sometimes slightly woody; leaves lanceolate to ovate, often leathery; inflorescences terminal, few- or many-flowered, with dichasial cymes or panicles, with scale-like bracts; flowers 4 or 5-merous; corolla narrowly funnel-shaped, thin, purple, magenta, yellow, lilac, to pink. Three spp., endemic to Pantepui Life Zone in Auyan, Aracamuni, Ilu, Kukenan, Chimantá (one endemic), Roraima, Pirapucu, Mount Neblina (one endemic), all in Venezuela, Amazonas state in N Brazil (only C. bella Maguire & Steyerm.) and Guyana (only C. imthurniana (Oliv.) Maguire), 1,500 – 2,800 m elevation range, in grasslands and open, rocky areas on tepui summits.

24. Chorisepalum Gleason and Wodehouse. Shrubs or small trees; inflorescences terminal, few-flowered, cymose, with leaf-like bracts; flowers erect, actinomorphic; corolla 6-merous, salver- to funnel-shaped, thin, green, with spreading lobes. 5 spp., endemics to the Guiana Shield of Venezuela to Suriname, at 800 – 2,400 m elevation range, in grasslands and open, rocky areas on tepui summits (high-altitude).

25. Chelonanthus Gilg. (inc. Calolisianthus p.p.) Annual or perennial herbs (up to 3.5 m tall), sometimes woody at the base; inflorescences terminal, many-flowered; flowers 5-merous, often horizontal or nodding, actinomorphic or slightly zygomorphic; corolla campanulate to funnelshaped, blue, purple, green, yellowish, or white. 14 spp. from South America, one up to Mexico and Caribbean; 9 in Brazil, two endemics; forests, roadsides, savannas, and grasslands.

26. Helia Mart. Herbs, not branched; inflorescence terminal, cymose, long-stalked; flowers 5-merous, actinomorphic; corolla salvershaped, white to yellow, tube narrow, lobes rounded, persistent in fruit. Two spp., of Brazil, one up to Paraguay and Bolivia, in wet grasslands, meadows, and savanas.

27. Irlbachia Mart. Annual herbs with leaves on the stems or in a basal rosette; flowers arranged in terminal cymes with long unbranched branches; flowers 5-merous, actinomorphic with zygomorphic stamens and style; corollas salvershaped, campanulate, or narrowly funnelshaped, often with a short corolla tube, and white, pink, lavender, blue, or purple. 9 spp., I. alata (Aubl.) Maas from Mexico, Central America, Trinidad & Tobago, Venezuela, Guianas, Ecuador, Peru and Brazil, remaining only from Colombia to French Guiana and N Brazil (5, none endemics), primarily in areas north of the Amazon River in Brazil, Venezuela, Colombia, and Guyana, in savannas, grasslands, open areas, often on white-sand. Several species are endemic to the tepui mountains.

Future reclassifications should probably include I. pratensis (Kunth) L. Cobb & Maas, from Venezuela, Colombia, Brazil, inside a smaller, recircumscribed Chelonanthus.

28. Lagenanthus Gilg. Shrub, branched; flowers solitary or few, terminal, hanging; corollas long-tubular, slightly inflated, base yellow, middle with broad scarlet-red and orange bands; stamens inserted close to base of corolla tube; filaments long; fruits capsules, thick-walled. Only one sp., L. princeps (Lindl.) Gilg, restricted of high elevation areas in mountains along the Colombian-Venezuelan frontier in South America; Lindley (1849) though this was ‘one of the noblest plants in existence’.

29. Lehmanniella Gilg. Herbaceous or slightly woody plants, with scandent branches; inflorescence terminal umbels, few-flowered; flowers 5-merous; calyx campanulate; lobes elliptic, rounded at apex; corolla narrow, red, tubular, inflated; lobes short and rounded. Two spp., Peru and Colombia one endemic each, in cloud forests in high elevation areas in mountains.

30. Macrocarpaea (Griseb.) Gilg. Branched shrubs (rarely epiphytic) or small trees up to 10 m (rarely perennial herbs); inflorescences terminal, composed of few- or many-flowered dichasia or cymes; flowers 5-merous, large(2.0-7.5 cm long), slightly zygomorphic; calyces campanulate, fused at the base, thick; corollas yellow, white, to greenish, funnelshaped, thick and fleshy. 123 spp., all except M. macrophylla (Kunth) Gilg (Panamá to Colombia) exclusiverly a single bioregion: Central America (4), Caribbean (3) and South America (116); c. 100 in Andes from Venezuela to Bolivia, six in Guiana Shield, and 9 spp. in Brazil, M. piresii Maguire from Amazonas state and adjacent Venezuela, and 8 from Atlantic Forest from Bahia to Santa Catarina states, two reaching to rocky grasslands, beloging the endemic section Tabacifoliae.

31. Neblinantha Maguire. Semi-woody herbs or shrubs; flowers solitary, sometimes appearing axillary, but are terminal, 4–5-merous, actinomorphic; corolla deep pink to coral-colored, salvershaped or narrowly funnelshaped, thin; lobes circular or triangular. Two spp., both endemic to Pantepui Life Zone, in Mount Neblina, Amazonas state in N Brazil (both, none endemics) and Venezuela, at 2,200 – 2,600 m elevation range.

32. Prepusa Mart. Herbs, shrubs, or small trees; leaves basal rosette in the herbs, or crowded at the branch apices in the trees or shrubs; flower 6-merous showy with large, inflated, campanulate and thin-walled calyx, often with wings; corolla nearly hidden inside calyx, white or yellow, salver- to funnelshaped. 6 spp., all restricted to the rochy grasslands and high altitud grasslands (campos de altitude) in Bahia, Espírito Santo, Rio de Janeiro and adjacent Minas Gerais, four of then are rares by Plantas Raras do Brasil’s book.

33. Purdieanthus Gilg. Straggling vines, branches narrow branches; leaves shortly petiolated, elliptic; inflorescence terminal, cymose; calyx small with short, oblong lobes; corolla long, tubular. Only one sp., P. pulcher (Hook.) Gilg, of high elevations on the Colombian and SE Venezuelan, in cloud forests and paramos, at 2,500-4,000 m altitude.

34. Rogersonanthus Maguire & B.M.Boom. Shrubs to trees; inflorescence small, terminal cyme, 1–12-flowered; flowers 5-merous; calyx campanulate, fused at base, corollas green to yellow, thick, and leathery; stamens inserted in lower half of corolla tube; anthers lanceolate, recurved after anthesis. Two spp., in Guiana Shield in N South America, including the tepui mountains of Guyana and Venezuela, plus the island of Trinidad, in cloud forests, mountain meadows, bogs, and grasslands at high elevation.

35. Roraimaea Struwe, S.Nilsson & V.A. Albert. Suffrutescent short-lived perennial herbs up to 1 m, glabrous, woody at base; stem and branches herbaceous, with four, very narrow wings; inflorescence terminal and diffuse, cymose, 1–18-flowered; flowers 5-merous, erect to horizontal; corolla tubular or salver-shaped, orange to red, deciduous in fruit. Two spp., R. aurantica Struwe, S. Nilsson, & V.Albert in white-sand area from the S part of the state of Roraima in Brazil, and R. coccinea (Steyermark ex L. Struwe, S. Nilsson, & V. A. Albert) Struwe, S. Nilsson, & V. A. Albert, in wet savannas in Mount Neblina from Brazil and Venezuela up to 1,900 m alt.

36. Senaea Taub. Shrubs; branches erect, round or square; inflorescences with leaf-like bracts, flowres in terminal or axillary cymes; flowers 6-merous; calyx campanulate; corolla blue, funnel- to bellshaped. Two spp. in Minas Gerais and Rio de Janeiro states, one of then rare plant in Brazil, by Plantas Raras do Brasil’s book.

37. Sipapoantha Maguire & B.M. Boom. Annual herb; inflorescence terminal, cymose, 1–3 branches, 1–7-flowered; flowers 5-merous, slightly zygomorphic; corolla showy and large, funnelform, dark blue to purple, thin, deciduous in fruit, lobes elliptic to circular, obtuse, corolla bud apex rounded. Two spp., S. ostrina Maguire & B.M. Boom in Cerro Sipapo, in Amazonas State, Venezuela (1,200 – 2,200 m elevation range), and S. obtusisepala Lepis, Maas & Struwe endemic to SE Roraima state, Brazil, both in mountain summits.

38. Symbolanthus G.Donwith. Shrubs, rarely small trees or herbs; flowers over (5-)8-15 cm long; trumpet-shaped corollas with long corolla tube, white, green, yellow, pink, or red, often with white stripes in corolla mouth. 33 spp., Caribbean and Central America one restricted each, and 31 remaining in mountain areas of Colombia, Ecuador, Guyana, Peru, Venezuela, Bolivia, Brazil (two, Amazonas and Roraima states), in rainforests, cloud forests, paramo, and montane shrubby and grassy areas.

39. Tachia Aubl. Small trees or shrubs, often with hollow and yellow branches; flower in leaf axils, without bracts, solitary, slightly zygomorphic; calyx fused at base, sometimes more, yellow, tubular, sometimes keeled; corollas tubular to salvershaped, with long corolla tube, yellow, cream, orange, or greenish. 14 spp., one in a Central America, another from Costa Rica to Brazil and Bolivia; all others are exclusively from Amazon rainforest; 7 in Brazil, 3 endemics. Tachia guianensis Aubl. from Guianas is a myrmecophite.

All of then have a very peculiar feature only visible when dried: the specimens become glued to the newspapers in which they were dried, by the secretion of a sticky substance probably fromthe axils of the leaves; the way this secretion is formed and its functon is still completely unknown, and needs further investigation.

40. Tetrapollinia Maguire & B.M.Boom. Single-stemmed annual, sometimes very small and inconspicuous; inflorescence a terminal cyme with monochasial branches, with 1–20 flowers; flowers 5-merous; corolla funnelshaped, varying from white, blue, pink to purple. Only one sp., T. caerulescens (Aubl.) Maguire & B.M. Boom, from savannas of tropical South America, in Bolivia, Brazil, French Guiana, Guyana, Suriname, Venezuela.

41. Yanomamua J.R.Grant, Maas & Struwe. Herb to 1 m, glabrous throughout; stems quadrangular, 3–4 mm in diam; inflorescence composed of solitary flowers oppositely paired in the axils of the upper three leaf pairs; corolla, stamens, and pistil unknown. Only one sp., Y. araca E. Dean, Maas & Struwe, known only open summit of Mount Aracá of N Amazonas state in N Brazil; unique in the Gentianaceae as an herb with sessile subcordate pandurate (fiddle-shaped) leaves.

1.6 TRIBE GENTIANEAE (17/940–970) ▸ outsiders Kuepferia (12; N India, Sikkim, S and E Tibet, NW Yunnan, SW Sichuan, Nepal, Bhutan, N Burma), Crawfurdia (16; the Himalayas to N Burma and China), Metagentiana (14; China, N Burma, N Thailand), Sinogentiana (2; China), Tripterospermum (30–35; E Asia); Bartonia (3; E North America), Comastoma (c 25; temperate and arctic-alpine regions on the Northern Hemisphere, the Himalayas), Frasera (15; North America), Gentianopsis (c 20; temperate regions on the Northern Hemisphere), Jaeschkea (3; the Himalayas), Latouchea (1; E China), Lomatogonium (21; temperate and arctic-alpine regions in Europe and Asia), Megacodon (1; Himalayas), Obolaria (1; SE Canada, E U.S.A.), Pterygocalyx (1; E Asia), Lomatogoniopsis (3; China), Swertia (135–150; Europe, temperate Asia, Madagascar, mountain regions in Africa and Malesia), Veratrilla (2; E Himalayas, W China).

42. Gentiana L. Annual, biennial or perennial herbs. Leaves opposite, very rarely whorled; flowers 5-merous (rarely 4- to 8-merous), in simple dichasia, terminal clusters, axillary whorls, or solitary; calyx fused about halfway up, sometimes split to the base on one side, often with intracalycine membrane; corolla tubular, funnelshaped, campanulate, urnshaped, or rarely rotate. 335 spp., most of the species occur in temperate areas in Asia, Europe, North America, only two in Southern Hemisphere, both in sect. Chondrophyllae and South American: G. sedifolia Kunth, from Costa Rica, Panamá, Andes from Colombia to northern Chile, in moist alpine habitats, and G. gayi Griseb., from northern Chile and southern Bolivia to Tierra del Fuego, in moist alpine habitats and wet grasslands.

43. Gentianella Moench. Annual to perennial herbs and range from 3–100 cm in height, sometimes cushions; flowers 0.5–5 cm long, have entire petal margins, and one or two naked nectaries per petal lobe on the upper petal surface. 260 spp., in alpine or arctic habitats in Eurasia (30), NW Africa (1), North America (10) and South America (195), 3/4 confined to center Andes (50 in Bolivia (33 endemic), 28 in Ecuador (21 endemic), 104 in Peru (95 endemic)), 7 in Colombia (one endemic), 29 in Argentina (18 endemic), 32 in Australia and New Zealand.

44. Halenia Borkh. Herbs to shrubs, sometimes cushions; 62 spp., two in Asia and remaining most are high-elevation plants of Central and South America (44), one up to Argentina; unique genus within Gentianaceae with nectary spurs; recent studies indicate that this genus has only 39 spp. in Asia (2), North America (1), Central America (15) and South America (21), centered in Colombia.

GELSEMIACEAE

§ FULLY CHLOROTROPHIC

Genera/spp. 3/14 Distribution SE U.S.A., E Mexico, NE South America, tropical Africa, Madagascar, S China, SE Asia, West Malesia. Habit bisexual, evergreen tree (Pteleocarpa), shrubs or lianas (Gelsemium, Mostuea). A small family composes of trees or shrubs, many rare in Neotropics.

SYSTEMATIC outsiders Gelsemium (India, N Burma, N China (inc. Taiwan), SE Asia, West Malesia; SE U.S.A., E Mexico, Guatemala), Pteleocarpa (1; West Malesia).

1. Mostuea Didrichsen. 10 spp., eight in Africa and two in South America: M. surinamensis Benth. occurs only center Suriname and Trombetas valley in Pará state in Brazil; and M. muricata Sobral & L.C. Rossi, a undershrub to shrub 0.5 – 1.5m tall, with white flowers, collected in one municipality of NE Mato Grosso state and in four municipalities of São Paulo state in Brazil, in shady and sandy sites in the interior of mesophyllous forests, where is eventually occur in great densities in the understory.

LOGANIACEAE

§ FULLY CHLOROTROPHIC

Genera/spp. 16/395–400 Distribution tropical and subtropical regions in the Northern and Southern Hemispheres: eastern U.S.A. to Uruguay, Africa south of the Sahara, Madagascar, Mascarene Islands, from India and Sri Lanka to Korean Peninsula and Japan, SE Asia, Malesia, New Guinea, Melanesia, Micronesia, Australia, Tasmania, Lord Howe Island, New Zealand, Hawaii and other Polynesian islands. Habit usually bisexual (rarely monoecious, gynomonoecious, dioecious, or gynodioecious), evergreen or deciduous trees, shrubs or lianas (with branch tendrils), perennial or annual herbs, not very extensive in dryland vegetation.

A basal group in relation to Rubiaceae, Apocynaceae and Gentianaceae, it is not an easy family to characterize and has been divided into several different families over the years.

Key differences from similar families

Superior ovary sets it apart from Rubiaceae and Campanulaceae.

Corolla valvate to imbricate (rather than contorted) separates it from Gentianaceae.

Lack of latex separates it from Apocynaceae.

SYSTEMATIC outsiders Usteria (1; tropical W and C Africa), Gardneria (5; India to C Japan and SE Asia, Java), Neuburgia (10–12; Philippines and Sulawesi to New Guinea, tropical Australia, New Caledonia, Vanuatu and Fiji), Adelphacme (1; W Australia), Mitrasacme (c 45; tropical Asia, China, Australia, New Caledonia), Phyllangium (5; Australia, Tasmania), Schizacme (5; Australia, New Zealand), Logania (c 22; Australia, New Caledonia, New Zealand, with their highest diversity in W Australia, South Australia and New South Wales), Geniostoma (c 40; Mascarene Islands, Malesia to S Japan, and Taiwan in China, Queensland, Vanuatu, Fiji, New Zealand, Samoa, Tonga, Tahiti, Hawaii), Orianthera (13; Australia); Norrisia (2; SE Asia, West Malesia).

1. Antonia Pohl. Large shrub or small tree, with many series of bracts subtending each flower, resembling Asteraceae. Only one sp., A. ovata Pohl., from Bolivia, E Brazil to Guianas, Venezuela and Colombia.

2. Bonyunia M.R.Schomb. 7 spp. from tepuis of the Guiana region and outliers in lowland region of the Amazon River matersehed of Brazil (3, none endemics), Guyana, Venezuela and Colombia, and lowlands regions of Amazon rainforest facing Andes in Peru, throughout on white sands.

3. Liesneria M.R.Schomb. Only one sp., L. faveolata (Fernández Casas) Fernández Casas, in both sides of Ecuador/Colombia border.

4. Mitreola Boehm. Herbs, annual or perennial, erect to creeping, branched or not; branches terete to 4-angled; leaves opposite, sessile or petiolate; inflorescences terminal and/or axillary, usually long pedunculate, 2 or 3 or more branched, branches scorpioid; flowers sessile or shortly pedicellate, 5-merous; corolla urceolate to campanulate. 7 spp., three in China and adjacent regions, two in North America and adjacent Caribbean, one unknown, and M. petiolata (Walter ex J.F. Gmel.) Torr. & A. Gray from North America, Mexico, Central America, Caribbean, Trinidad & Tobago, Venezuela, Guianas, Colombia, Brazil, Ecuador, Peru and Bolivia.

5. Spigelia L. Herbs to shrubs, opposite or whorled leaves, one-sided cymose inflorescences, often brightly colored pentamerous flowers with usually funnelform or tubular corollas, articulated styles, and strongly bilobed capsules with persistent style and fruit bases. 94 spp., neotropical, from temperate South America northward into the tropics of South America, to Central America, Mexico and the Caribbean, and into the warm-temperate southern U.S.A., inhabiting mid-elevation to lowland areas, with at least 70 species found in South America; 55 spp. are distributed in Brazil (42 endemics), most species are geographically restricted, several are widely distributed from North to South America, S. anthelmia L. also naturalized in Africa and Malaysia, and cultivated as a medicinal plant.

S. genuflexa Popovkin & Struwe is a species described in 2011; it is unusual in that the stems bend down after flowering to deposit the seeds in the soil (geocarpy, the unique known case of amphigeocarpy in the Loganiaceae); it grows in residual stands of Atlantic forest in northern coast of Bahia state, at an altitude of 150 metres, flowering during the rainy season, and almost disappears during the dry season, able to self-pollinate, with anthesis lasting less than a day.

6. Strychnos L. Trees, shrubs, creeping armed or unarmed lianas, and have opposite leaves, with three to five veins starting at the same point of the leaf, inflorescences in panicles or corymbs, flowers with salverform or rotate corollas, and fruits are berries or globose capsules, with few or many seeds. 190 spp., pantropical, but very extensive in the Amazon rainforest, inc. SE Brazil and Argentina; 84 spp. in New World, 79 in South America, 69 in Brazil, 23 endemics; six Brazilian species of Strychnos from Amazonas (4), Minas Gerais (1) and Goiás (1) states are rares by Plantas Raras do Brasil’s book; three sections based mainly on differences in corolla shape and size:

§ sect. Longiflorae ▸ salverform corollas.

§ sect. Intermediae ▸ corolla tubes smaller and narrower than those of section Longiflorae.

§ sect. Breviflorae ▸ short, rotate corollas.

APOCYNACEAE

§ FULLY CHLOROTROPHIC

Genera/spp. 376/c. 4,600 Distribution cosmopolitan except polar areas and cold-temperate regions. Habit usually bisexual (rarely functionally dioecious), evergreen trees, shrubs or lianas (sometimes perennial or annual herbs). Numerous spp. are xerophytic and many are stem succulents. Some spp. (e.g. Dischidia major (Vahl) Merr. (= D. rafflesiana)) are myrmecotrophic with ant symbiosis (ant colonies present in leaves).

With about 5,000 spp., it is one of the ten largest families of angiosperms, and also one of the most popular, due to the traditional widely distributed use of some of its spp. as ornamental plants (Allamanda cathartica L., Catharanthus roseus (L.) G. Don, Nerium oleander L., Plumeria rubra L., Thevetia peruviana (Pers.) K. Schum.). Several spp. are rich in chemical compounds (Rauvolfia), others provide hardwood (e.g., Aspidosperma), and only a few are fruit spp. (Couma rigida Müll. Arg., known as mucugê, and Hancornia speciosa Gomes, known as mangaba). They show variable habit, from magna trees to vines or small herbs, including some succulent ones (Stapeliinae). Yet, they can be easily recognized by the presence of latex.

Subfamily Asclepiadoideae can be characterised by the presence of a gynostegium in which the expanded stylar head is fused to the androecium by cell fusion, and where the narrow gap between the sclerified wings of adjacent anthers form five grooves. The form of the pollinarium is also diagnostic for the subfamily, and indeed tribes within it can be determined by close observation of pollinarium structure. Pollen of one anther locule is aggregated into a pollinium encased by a waxy wall. This pollen mass is linked to another in an adjacent anther by a pair of translator arms and a single corpusculum. So two pollinia linked by translator arms to a single corpusculum form a pollinarium that can be extracted from the flower as a single unit. The corpusculum is usually black or dark brown, and sits at the top of the groove between adjacent anthers.

The family belongs to the Gentianales and can be easily recognized by the presence of latex and a style-head derived from the fusion of two carpels at the apex of the styles.

SYSTEMATIC subfamilies Periplocoideae (37/166–167, tropical, subtropical and arid temperate regions of the Old World to northern Australia) and Secamonoideae (8/200–206, tropical and subtropical regions in the Old World to northern Australia and islands in the Pacific), and tribes Alstonieae (2/47, tropical Africa, Madagascar, S China, tropical Asia, Melanesia, islands in the W Pacific, one species, A. longifolia, in Central America), Hunterieae (4/21, tropical and S Africa, Madagascar), Amsonieae (1/c 20, Greece, Türkiye, China, Korean Peninsula, Japan, the U.S.A., Mexico), Carisseae (2/15, tropical regions in the Old World to New Caledonia), Melodinae (5/29, tropical and S Africa, Madagascar, tropical Asia, islands in the Pacific), Wrightieae (2/35, tropical and subtropical regions in the Old World), Nerieae (6/74, Mediterranean, tropical and S Africa, Madagascar, Arabian Peninsula, Socotra, SW and tropical Asia), Apocyneae (24/125, Russia to East and tropical Asia, Vanuatu, North America) and Baisseeae (4/29, tropical and S Africa, Madagascar) do not occur in South America.

1. TRIBE ASPIDOSPERMATEAE (6/81) ▸ outisders Strempeliopsis (2, Jamaica, Cuba) and Haplophyton (1, Mexico, Guatemala).

1. Aspidosperma Mart. & Zucc. Shrubs and trees up to 45 m tall. 77 spp. from South America, eight up to Central America and Caribbean, 64 in Brazil, 24 endemics, two of then are rares by Plantas Raras do Brasil’s book.

2. Geissospermum Allemão. 6 spp. from N South America, from Venenzuela to French Guiana and N Brazil (all spp., one endemic), Bolivia and Peru.

3. Microplumeria Baill. Only one sp., M. anomala (Müll.Arg.) Markgr., in Amazon rainforest from N Brazil, S Venezuela and SE Colombia.

4. Vallesia Ruiz & Pav. 7 spp. from Central America, Caribbean and U.S.A., V. glabra (Cav.) Link up to N Argentina and Paraguay along western flank of South America.

2. TRIBE VINCEAE (8/c. 160) ▸ subtribes Kopsiinae (1/22, SE Asia, West Malesia, the Caroline Islands), Ochrosiinae (1/40–43, Mascarene Islands, Seychelles, SE Asia, Malesia to tropical Australia, islands in W Pacific to Hawaii), Vincinae (1/6, Europe, Mediterranean, North Africa to Central Asia) and Catharanthinae (3/17, 8; Cameroon, Madagascar, the Comoros, Sri Lanka, India) do not occur in South America.

■ SUBTRIBE TONDUZIINAE (2/4) ▸ outsider Tonduzia (2; Mexico, Central America).

5. Laxoplumeria Markgr. Trees or shrubs with white latex; leaves alternate or whorled; flowers small; calycine colleters absent; corolla salverform; fruit a pair of long, slender follicles; seeds numerous, compressed, elliptic, covered with long hairs or with clumps of shorter hairs marginally. 5 spp. from N South America, one up to Panamá, four in Brazil, two endemics.

■ SUBTRIBE RAUVOLFIINAE (1/76) ▸ a single genus.

6. Rauvolfia L. Shrubs or trees with whorled leaves and branches, hypocrateriform flowers, and generally small, drupaceous, variously syncarpous or apocarpous fruits with two pyrenes. 80 spp., tropical regions on both hemispheres, 39 spp. in New World, 30 in South America, 19 in Brazil, 9 endemics; R. blanchetii A.DC. from Bahia state is rare plant in Brazil, by Plantas Raras do Brasil’s book.

3. TRIBE WILLUGBEIEAE (18/157) ▸ subtribes Leuconotidinae (3/8, C Africa, SE Asia) and Willughbeiinae (1/17, India and Sri Lanka to Borneo and Sulawesi) do not occur in South America.

■ SUBTRIBE LACMELLEINAE ▸ all genera in South America.

7. Couma Aubl. 5 spp., slightly centered in N South America, C. macrocarpa Barb. Rodr. up to Belize to Bolivia and Brazil (all species), none is a national endemic.

8. Hancornia Gomes. Only one sp. H. speciosa Gomes, from Brazil to Paraguay, Peru and Bolivia.

9. Lacmellea H.Karst. 24 spp., highly centered in northern South America (22), three up to Central America, two only Central America; 12 spp. in Brazil, three endemics.

10. Parahancornia Ducke. 7 spp. from Amazon rainforest from Venezuela to Bolivia, all in N Brazil, two endemics.

■ SUBTRIBE LANDOLPHIINAE (10/95) ▸ outsiders Ancylobotrys (7; tropical and S Africa, Madagascar), Chamaeclitandra (1; tropical Africa), Clitandra (1; tropical Africa), Cylindropsis (1; tropical W and C Africa), Dictyophleba (6; tropical Africa), Orthopichonia (6; tropical W and C Africa), Saba (3; tropical Africa, Madagascar), Vahadenia (2; tropical W and C Africa).

11. Pacouria Aubl. Three spp. from northern South America, all in Brazil, P. paraensis (Huber) Pichon endemic.

4. TRIBE TABERNAEMONTANEAE (15/c. 154) ▸ two subtribes, both in South America.

■ SUBTRIBE AMBELANIINAE ▸ all genera in South America.

12. Ambelania Aubl. Three spp. from N. South America (Peru, Colombia, Guianas) to N & NE Brazil (all species, none endemics), mainly Amazon rainforest.

13. Macoubea Aubl. Three spp., one restricted Central America and two in N South America, both in Brazil, none endemics.

14. Molongum Pichon. Three spp. from SE Colombia, S Venezuela and in NW Amazonas state in N Brazil (all species, M. zschokkeiforme (Markgr.) Pichon endemic).

15. Mucoa Zarucchi. Two spp., M. duckei (Markgr.) Zarucchi from Peru, Colombia and in Amazonas state in N Brazil, and M. pantchenkoana (Markgr.) Zarucchi endemic to Venezuela.

16. Neocouma Pierre. Two spp. from Venezuela, Colombia and N Brazil (both species, none endemnics).

17. Rhigospira Miers. Only one sp., R. quadrangularis (Müll.Arg.) Miers from Venezuela to NE Peru and N Brazil.

18. Spongiosperma Zarucchi. 6 spp. from S Venezuela, Colombia (2) and N Brazil (4, two endemics) up to Maranhão state.

■ SUBTRIBE TABERNAEMONTANIINAE (8/c. 134) ▸ outsiders Callichilia (6; tropical Africa), Calocrater (1; tropical W and C Africa), Carvalhoa (1; E and SE Africa), Crioceras (1; Gabon to Angola), Schizozygia (1; tropical E Africa, the Comoros), Tabernanthe (2; C Africa), Voacanga (12; tropical and subtropical regions in the Old World).

19. Tabernaemontana Plum. c 110 spp., tropical regions on both hemispheres, 69 spp. in New World, 48 in South America, 30 in Brazil, 7 endemics; T. brasiliensis Leeuwenb. (syn. Stemmadenia brasiliensis Leeuwenb.) from Pará state is rare plant in Brazil, by Plantas Raras do Brasil’s book.

5. TRIBE ALYXIEAE (7/147-152) ▸ two subtribes, Alyxieae (4/113–118, Madagascar, tropical Asia to Solomon Islands, Micronesia and Polynesia) does not occur in South America; among subtribe Condylocarpinae, outsiders are Chilocarpus (14; S India and Nicobar Islands to New Guinea), Plectaneia (3; Madagascar).

20. Condylocarpon Desf. 7 spp., Nicaragua to South America (6, slightly centered in N continent), SW to Brazil (6, only one endemic, from Rio de Janeiro and Espírito Santo states, rare plant in Brazil, by Plantas Raras do Brasil’s book), and Cono Sur.

6. TRIBE PLUMERIEAE (10/80) ▸ three tribes, all in South America.

■ SUBTRIBE ALLAMANDIINAE ▸ a single genus.

21. Allamanda L. Shrubs to small trees, leaves verticillate, subverticillate, opposite or subopposite, infundibuliform corolla (often slightly zygomorphic) usually yellow (rose in A. blanchetii A. DC.), with a corona of hairs within the tube and above the anthers, dehiscent capsular fruits, winged seeds. 17 spp. from Honduras to Argentina, Brazil (14, 10 endemics, sometimes very narrow endemics to dry limestone outcrops), few in Colombia and Venezuela, some in French Guiana, Suriname and Peru, and A. cathartica L. widely distributed up to Mexico.

Many spp. ornamental worldwide, probably one of the most horticulturally popular genera in Apocynaceae.

■ SUBTRIBE PLUMERIINAE (3/39) ▸ outsider Mortoniella (1; Central America).

22. Himatanthus Willd. ex Schult. 9 spp. from South America, one up to Panamá, all in Brazil, none endemics.

23. Plumeria Tourn. ex L. 24 spp. exclusives of Caribbean except four up to Florida, Mexico, Central America and N South America (3), in Colombia, Venezuela and Guianas.

■ SUBTRIBE THEVETIINAE (6/25) ▸ outsiders Cameraria (7; Caribbean), Cerbera (6; coasts of the Indian and W Pacific Oceans), Cerberiopsis (3; New Caledonia).

24. Anechites Griseb. Two spp. from Caribbean and Central America to Venezuela (one endemic) to Peru and Ecuador.

25. Skytanthus Meyen. Three spp., S. acutus Meyen endemic of Chile from Antofagasta to Coquimbo, S. hancorniifolius (A. DC.) Miers and S. martianus (Müll. Arg.) Miers restricted of E Brazil.

26. Thevetia L. 4 spp., two from Mexico to Cono Sur and Cuba, and two remaining restricted of Brazil and adjacent Bolivia and Argentina.

7. TRIBE RHABDENIEAE (1/6) ▸ a single genus.

27. Rhabdadenia Müll. Arg. 6 spp., one in Mexico, R. biflora (Jacq.) Müll. Arg. from S Florida to trop. America, the remaining four only in South America; three in Brazil, none endemics.

8. TRIBE ODONTATENIEAE (7/41) ▸ outsiders Cycladenia (1; SW U.S.A.), Pinochia (4; Mexico, Central America, the Greater Antilles), Thyrsanthella (1; SE U.S.A.) and subtribe Pentaloninae (2/4, Florida, Central America, the Caribbean).

28. Elytropus Müll.Arg. Only one sp. E. chilensis (A.DC.) Müll.Arg., from C and S Chile and Rio Negro in Argentina.

29. Odontadenia Benth. 21 spp., from Mexico to Brazil (17, two endemics) and Caribbean, 20 in South America.

30. Secondatia A.DC. Six spp., S. densiflora A. DC. over South America and remaining more restricteds in Jamaica (1), Colombia (1), Brazil (2, one also in Colombia and S. floribunda A. DC. endemic) and mountains of Ecuador and Peru (1).

31. Stipecoma Müll.Arg. Vines, flowers pink. One sp., S. peltigera (Stadelm.) Mull.Arg., from C Brazil to E Bolivia.

9. TRIBE MALOUETINAE (9/c. 105) ▸ three subtribes, Pachypodiinae (2/32-33, Cuba, the Bahamas, S and SE Africa, Madagascar) does not occur in South America.

■ SUBTRIBE GALACTOPHORINAE ▸ a single genus.

32. Galactophora Woodson. 6 spp. endemics to the Guiana Shield from N South America, including Amazon rainforest of Brazil, except by some expecis which extends to S lowland Peru, Bolivia and Guyana; 3 spp. in Brazil, none endemics.

■ SUBTRIBE MALOUETIINAE (8/66) ▸ outsiders Allowoodsonia (1; Solomon Islands), Carruthersia (4; Philippines to Solomon Islands, Fiji to Tonga), Eucorymbia (1; Borneo), Funtumia (2; tropical Africa), Holarrhena (5; tropical Africa, tropical Asia), Kibatalia (c 15; SE Asia, Malesia to Philippines), Mascarenhasia (8; Madagascar, E and S Africa), Spirolobium (1; Thailand, Indochina, the Malay Peninsula).

33. Malouetia A.DC. Shrubs up to tall trees (25 m), with domatia in the axils where the secondary veins meet the midvein, the almost sessile, umbel-like fasciculate inflorescences, and the absence of a distinct tuft of hairs (coma) at the end of each seed. 31 spp., three in W & WC tropical Africa and 28 spp. over tropical South America, two up to Central America, 15 in Brazil (7 endemics, one of then rare plant in Brazil, by Plantas Raras do Brasil’s book, in Roraima state), mainly along the black-water rivers of northwestern Amazon rainforest, especially the tributaries of the Orinoco and Casiquiare river systems.

10. TRIBE ECHITEAE (16/c. 270) ▸ four subtribes, Parsoniinae (5/c. 140, E Asia to China (Taiwan), tropical Asia to northern Australia, New Caledonia, New Zealand, Fiji, S Mexico, Central America) do not occur in South America.

■ INCERTAE SEDIS

34. Bahiella J.F.Morales. Lianas, with old stems woody with latex; young stems terete; leaves opposite, peciolate; inflorescence cymose, axillary, paniculate, many flowered. Two spp. very narrow endemics to coastal forests of Bahia state, Brazil.

■ SUBTRIBE PELTASTINAE (4/25) ▸ a unknown genus by SDb.

35. Macropharynx Rusby. (inc. Asketantera p.p.) 16 spp., scatterd in South America (over genus) over from Costa Rica to N Brazil (4, one endemic) and Guianas.

36. Rhodocalyx Miers. Two species, both Brazil, also in several adjacent countries.

37. Temnadenia Miers. 3 spp., one in Colombia and two endemics to Brazil.

■ SUBTRIBE ECHITINAE (4/?) ▸ outsider Asketanthera (4; tropical America), Thenardia (?; ?), Thoreauea (?; ?).

38. Echites P.Browne. 10 spp., all Caribbean, Mexico, Central America and Florida, with E. umbellatus Jacq. occurring in Colombian Caribbean Is.

■ SUBTRIBE LAUBERTIINAE (2/5) ▸ both genera in South America.

39. Hylaea J.F.Morales. Two spp., H. arborescens (Monach.) J.F.Morales from S. Venezuela, and H. leptoloba (Monach.) J.F.Morales endemic to N Brazil (Amazonas state).

40. Laubertia A.DC. Three spp., L. peninsularis Woodson from Mexico to Belize and Guatemala, L. boissieri A. DC. from Venezuela to Bolivia, and L. brasiliensis J.F. Morales, endemic to Brazil to the states of Pará and Amazonas, growing in river margins and igarapó forest, from 100 to 200 m elevation range.

■ SUBTRIBE PRESTONIINAE (1/74) ▸ a single genus.

41. Prestonia R.Br. Lianas with eglandular leaves, sometimes with taproot tubers or xylopodium; axillary or terminal cymose inflorescences, sepals with a single colleter, flowers usually with an annular corona around the mouth and/or free corona lobes, follicular fruits, and truncate seeds that are comose at themicropylar end. 74 spp. from Mexico to South America and Caribbean, 58 in South America, 22 in Brazil, 6 endemics.

11. TRIBE MESECHITEAE (6/236) ▸ outisders Titinnabularia (3; Mexico to Honduras).

42. Allomarkgrafia Woodson. 10 spp., three only Costa Rica/Panamá, A. plumeriiflora Woodson from Costa Rica to Colombia, and 6 only from S Colombia, N Ecuador and Peru.

43. Forsteronia G.Mey. 44 spp., from Mexico to trop. America, 42 in South America, 28 in Brazil, 11 endemics.

44. Mandevilla Lindl. Mostly woody to fragile vines, but erect shrubs are also common, while unbranched subshrubs and epiphytes occur less frequently, sometimes with xylopodium; flower in racemose inflorescense, ranging from inconspicuous white, tubular flowers less than 1 cm long to brightly colored, showy infundibuliform flowers up to 9 cm long. 172 spp., S U.S.A, Mexico to Argentina, largest Neotropical genus in Apocynaceae, highly complex in deserts, savannas, tepuis, open grasslands, and forests, 150 in South America, 73 in Brazil, 40 endemics; three spp., all from Minas Gerais, are rares by Plantas Raras do Brasil’s book.

M. longiflora (Desf.) Pichon. from Bolivia, Argentina, S Brazil, Paraguay and Uruguay has the longest corolla tubes in Apocynaceae, reaching up to 17 cm in and are only fully open at dusk, when they produce a distinctive scent, suggesting pollination by hawkmoths; their flowers are showy and variously colored and, in most cases, have an infundibuliform corolla.

45. Mesechites Müll.Arg. 10 spp., most single endemics in Caribbean (3), Peru (3), Colombia (1), Venezuela (1), except M. trifidus (Jacq.) Müll. Arg. from Mexico to Brazil and Cono Sur and M. mansoanus (A. DC.) Woodson in Brazil and Cono Sur.

12. SUBFAMILY ASCLEPIADOIDEAE (181/2.420–2.430) - five tribes; Fockeeae (2/9, tropical and S Africa, Oman), Eustegieae (2/6, SE Africa) e Ceropegieae (11/798, 90% in Ceropegia) do not occur in South America.

12.1 ASCLEPIADOIDEAE ▸ TRIBE MARSDENIEAE (27/c 380) - outsiders are all in Africa and Madagascar (4), India, S China and Himalayas to tropical Australia and islands in W Pacific (15) except Gongronema (15; tropical regions in the Old World), Hoya (200; tropical Asia, tropical and E Australia, islands in the Pacific to Polynesia), Marsdenia (c. 100; northern to S Africa, Madagascar, tropical and subtropical Asia to Malesia, Australia, Melanesia, one species, M. erecta, in eastern Mediterranean), Telosma (10; tropical N S Africa, Madagascar, tropical Asia eastwards to the Malay Peninsula),

46. Ruehssia H.Karst. (inc. Marsdenia) Lianas, suffrutescent twiners, herbaceous twiners or shrubs, sometimes with very odd leaves, often succulents (3, Brazil, Guyana and Venezuela one endemic each); latex white, yellow, brown or colorless; corolla usually with well-developed tubular portion enclosing the gynostegium. 137 spp. in the New World, 69 in South America, 43 in Brazil (33 endemics); some species associated with inselbergs, limestone outcrops in E Brazil, Minas Gerais, Bahia and Goiás states; 11 spp., in several states from Brazil, are rares by Plantas Raras do Brasil’s book. Brazilian succulent member is R. megalantha (Goyder & Morillo) F.Esp.Santo & Rapini from NE region, unique Brazilian succulent in Apocynaceae.

12.2 ASCLEPIADOIDEAE ▸ TRIBE ASCLEPIADEAE (74/1.630–1.680) - 12 subtribes worldwide, Astephaniane (3/14, S Africa) and Tylophirinae (4/c. 120, tropical and subtropical regions of the Old World) do not occur in South America. This tribe has the 4ª largest group of succulent species worldwide, c. 1,150 spp., mainly in Old World.

■ SUBTRIBE ASCLEPIADINAE ▸ outsiders list unvailable.

47. Asclepias L. Erect herbs or shrubs, perennial or annual (rarely), to 100 cm high, sparsely branched or richly branched, latex white, sometimes with xylopodium; one succulents sp. occur in Mexico; flowers 5-merous, generally brightly coloured, red, yellow or white. c. 220 spp., 80 in Africa, mainly in South Africa, 120 in North America, 12 spp. extending into Central and South America (10); South American species are strongly supported as monophyletic and derived from North American ancestors, all restricteds of Bolivia to Argentina and S Brazil (6, A. bracteolata E. Fourn. endemic) except the widely distributed A. curassavica L. and A. woodsoniana Standl. & Steyerm. from Mexico to Colombia

■ SUBTRIBE CYNANCHINEAE ▸ outsiders list unvailable.

48. Cynanchum L. (inc. Telminostelma, exc. Petalostelma p.p., Scyphostelma p.p.) Suffrutescent twiners, erect herbs, stem succulent twiners or erect stem succulents, 30–400 cm high, unbranched or sparsely branched or richly branched; latex white, ivory or yellow; subterranean organs constituting a woody rootstock, tap roots, fibrous roots, or root tubers. 262 spp., of Africa, America, Asia, Australia, Europe; very variable, often slightly disturbed habitats; 88 spp. in New World, 76 in South America (highly centered between Venezuela and Peru), 7 in Brazil, 5 endemics.

■ SUBTRIBE PENTACYPHINAE ▸ a single genus.

49. Pentacyphus Schltr. Weakly twining subshrubs with white latex; inflorescences sciadioidal; flowers nodding; corolla campanulate; pollinia medifixed or (sub-)basally inserted on corpusculum. 5 spp. of South America, in Colombia, Ecuador, Peru, Venezuela; mountain rain forest, 1,800–4,100 m.

■ SUBTRIBE DIPLOLEPINAE ▸ a single genus.

50. Diplolepis R.Br. Suffrutescent twiners 3–4 m high or shrublets 30–100 cm high. 14 spp., Argentina and Chile.

■ SUBTRIBE ORTHOSIINAE ▸ all genera occur in South America.

51. Jobinia E. Fourn. Suffrutescent twiners; shoots glabrous; leaves petiolate. 17 spp., one from Mexico to Honduras and 16 only in South America, 9 in S & SE Brazil (6 endemics, one of then rare plant in Brazil, by Plantas Raras do Brasil’s book); almost a half of Andes in Ecuador to Bolivia, and remaining occur inlowlands.

52. Monsanima Liede-Schumann. Slender twining habit, extra-axillary inflorescences shorter than leaves, corona with fluted lobes, almost entirely connate, free only at the apex, where they arch over the gynostegium, and pollinaria with short, flattened, broader than long, hyaline caudicles. Two spp., M. tinguaensis R.Santos & Fontella is known only from the type specimen, in Serra do Mar in Rio de Janeiro state, and M. morrenioides (Goyder) Liede & Meve, which is known only from Pico das Almas in Diamantina Range, Bahia State, and rare plant in Brazil, by Plantas Raras do Brasil’s book.

53. Orthosia Decne. Suffrutescent or herbaceous twiners, 0.5–8 m high, with distinct long and short shoots, orthotropous or plagiotropous (often when young). 32 spp. of Caribbean, U.S.A. (Florida); Belize, Costa Rica, El Salvador, Guatemala, Honduras, Mexico, Nicaragua, Panamá, Venezuela, Argentina, Bolivia, Brazil (15, 9 endemics, two of them rares by Plantas Raras do Brasil’s book), Paraguay, Peru; forests, mainly in clearings, thickets, roadsides, to 2,000 m; 23 in South America.

54. Scyphostelma Baillon. 22 spp., one in Costa Rica and Panamá, and all remaining in Andes from Colombia to Bolivia.

■ SUBTRIBE METASTELMATINAE ▸ all genera in South America.

55. Barjonia Decne. Subshrubs to erect herbs, sparsely branched; leaves opposite or decussate, (sub-)sessile; inflorescences thyrsoid to pleiothyrsoid, corolla (sub-)campanulate, or rotate; lobes ovate, ovate-triangular or lanceolate, erect, sometimes curved at the apex. 7 spp. of Brazil (three endemic, mainly Goiás, Mato Grosso do Sul, Minas Gerais, Paraná, São Paulo, one is rare plant in Brazil, by Plantas Raras do Brasil’s book), four into Bolivia and Suriname, in open fields, savannas of C Brazil (cerrado), as well as in rocky grasslands (campos rupestres), mainly in central Brazil, including Serra da Canastra and the Espinhaço Range.

56. Blepharodon Decne., Herbaceous or suffrutescent twiners, rarely shrubs or herbs, to 10 m high, sometimes with xylopodium, latex white. 26 spp. of over South America, one up to Mexico, in forest margins, roadsides; dry sandy or rocky savannas, mainly Brazil (12, 5 endemics, one of them rare plant in Brazil, by Plantas Raras do Brasil’s book).

57. Ditassa R. Br. Herbaceous or suffrutescent twiners, rarely (sub-)shrubs, orthotropous, mostly with small flowers; corolla lobes mostly not obviously pubescent, corona of five 'double' lobes - each lobe has a ligule on the inner face. 110 spp. of South America in Argentina, Bolivia, Brazil (62, 52 endemics, 12 of them rares by Plantas Raras do Brasil’s book, mainly from Minas Gerais, but some also in Rio de Janeiro, Espírito Santo, Tocantins and Bahia), Colombia, Ecuador, Guyana, Paraguay, Peru, Suriname, Uruguay, Venezuela, in forest margins, thickets, some in ferriculous sites, 100–3,000 m.

58. Hemipogon Decne. Erect herbs, subshrubs, herbaceous or suffrutescent twiners, sometimes with xylopodium, 30-50 cm high, latex white, subterranean organs constituting a woody rootstock or taproots. 9 spp., one from mountains of Bolivia and Peru, remaining in Brazil (4 endemics, two rare plant in Brazil, by Plantas Raras do Brasil’s book, both in Minas Gerais state) and similar habiatats in neighbroung Peru, Bolivia, Paraguay, one also in Ecuador; H. sprucei E. Fourn. has a strikingly disjunct distribution, occurring in Brazil and also in dry Andean valleys of Peru and Bolivia

59. Hypolobus E.Fourn. Suffrutescent twiners, wiht shoots sparsely hirsute to tomentose over the whole surface; calyx longer than corolla tube, basally fused, abaxially with trichomes. Only one sp., H. infractus E. Fourn, in Brazil (Bahia), possibly extinct, rare plant in Brazil, by Plantas Raras do Brasil’s book, known only the type collection but without type-locality.

60. Metastelma R. Br. Herbaceous or suffrutescent twiners, to 4 m high, latex white to ivory, subterranean organs constituting a woody rootstock; small flowers; corolla lobes generally pubescent above, corona of 5 staminal lobes. 93 spp., North America, Caribbean, Central America, South America (21), in forest margins, disturbed areas, riverine vegetation, to 3,000 m; 5 spp. in Brazil (4 endemics, one rare plant in Brazil, by Plantas Raras do Brasil’s book, in Bahia state, exception the widely distributed M. parviflorum (Sw.) R. Br. ex Schult.).

61. Minaria T.U.P.Konno & Rapini. Shrubs or subshrubs, rarely herb-like; branches erect, rarely prostate or twining to the apex; latex white; leaves decussate, coriaceous, rarely membranaceous; inflorescences cymose, subaxillary or axillary, occasionally single-flowered; corolla white, cream or yellow, rarely pinkish, campanulate to urceolate, usually less than 5 mm long. 22 spp., one only in Bolivia, two from Brazil to Bolivia, Argentina and Paraguay, and 19 endemics to Brazil (14, one ex-Barjonia, another ex-Hemipogon, all in Minas Gerais state, are rare plant in Brazil, by Plantas Raras do Brasil’s book), most of then are confined to small areas of rocky grasslands in the Espinhaço Range, mainly in its Minas Gerais portion.

62. Morilloa Fontella, Goes & S.A.Cáceres. Hemipogon-like. 4 spp. endemics to E Brazil, occurring in rocky grasslands (campos rupestres), savannas of C Brazil (cerrado) and in forest borders in the States of Goiás, Bahia, Minas Gerais and São Paulo, two of them are rares by Plantas Raras do Brasil’s book.

63. Nautonia Decne., Prostrate herbs, many small. Only one sp., N. nummularia Decne from Argentina, S Brazil, Paraguay, in sandy areas.

64. Nephradenia Decne., Erect herbs, 30-50 cm high; shoots glabrous or almost so. 5 spp. endemic to Brazil (with one rare plant in Brazil, by Plantas Raras do Brasil’s book, in Mato Grosso state) except by N. linearis Benth. ex E. Fourn. also in Venezuela, Colombia, Bolivia and Guianas.

65. Peplonia Decne. (inc. Macroditassa) Twining vines, glabrous or almost so; leaves usually elliptic, oblong, ovate or lanceolate (narrowly ovate), often subcoriaceous and discolorous. Inflorescences axillary, usually opposite, often with a short, bifurcate peduncle. Flowers white, greenish or yellowish; corolla urceolate to rotate, abaxially glabrous, adaxially puberulent, lobes usually barbate to barbellate in the basal half. 14 spp., one from Venezuela and Guyana, P. adnata (E. Fourn.) U. C. S. Silva & Rapini from Brazil, Peru and Bolivia, one in Brazil and Guianas, and remaining 11 spp. endemics to Brazil (Bahia to Rio Grande do Sul, two of them rares by Plantas Raras do Brasil’s book); seaside scrub, Atlantic sandy coastal shrublands (restingas), roadsides, thickets, riverine vegetation.

66. Petalostelma E.Fourn. Suffrutescent twiners, to ca. 1 m high. 12 spp., Argentina, Bolivia, Brazil (9, 5 endemics, mainly in Chaco region), Paraguay, in dry vegetations.

67. Vailia Rusby. Suffrutescent twiners, to ca. 1 m high. Two spp., V. salicina (Decne.) Morillo in over N South America (inc. Brazil and Ecuador), and V. anomala (Brandeg.) W. D. Stevens from Mexico to Colombia and Venezuela.

■ SUBTRIBE TESSADIINAE ▸ a single genus.

68. Tassadia Decne. Suffrutescent twiners with distinct long and short shoots, latex white. 29 spp. from northern South America, one up to Mexico; in mountain forests, disturbed places, river shores, flood plains, to 1,900 m; Brazil has 16 spp., 3 endemics, inc. two rare plant in Brazil, by Plantas Raras do Brasil’s book, in Mato Grosso and Tocantins.

■ SUBTRIBE OXYPETALINAE ▸ all genera in South America.

69. Araujia Brot. (inc. Morrenia) Suffrutescent twiners, 5–6 m high, latex white. 12 spp. from Argentina (6), Bolivia, Brazil (7, none endemics), Paraguay, Uruguay, A. sericifera Brot. up to S Peru; mainly subtropical Chaco, dry to moist forest, often in disturbed situations.

70. Funastrum E. Fourn. Herbaceous or suffrutescent twiners, latex white, with a garlic scent, rhizomes frequently present; often with their feet in seasonally flooded pools; stems semisucculent and photosynthetic (leaves often drop off); umbels of white or cream flowers, corona mosly with a tubular outer ring, and 5 fleshy lobes on the back of the anthers. 20 spp., from S U.S.A. to Paraguay and Caribbean, 8 in South America, two in Brazil, none endemics; rheophytes, arid and semi-arid areas, plains, pampa, stony slopes, to 1,500 m.

71. Oxypetalum R. Br. (inc. Schistonema) Suffrutescent twiners or erect herbs, sometimes with xylopodium, shoots glabrous, densely hirsute, hispid, pilose, pubescent or villous; flowers moderate in size; mostly with conspicuous stylar head appendages (divided into 2 or more lobes); corona lobes arising from the corolla tube not on the back of the anthers. 136 spp. from Argentina, Bolivia, Brazil (95, 52 endemics, 13 rares by Plantas Raras do Brasil’s book), Ecuador, Paraguay, Peru, Uruguay, only two north of Ecuador, up to Mexico and Caribbean; forest, on forest and pasture margins, and in open, often slightly disturbed areas, from humid to rather dry landscapes.

72. Philibertia Kunth. Suffrutescent twiners, prostrate or erect herbs, 150-500 cm or 20-40 cm high, latex white with a garlic scent, or, rarely, colorless; corona on back of stamens not on the corolla, or absent; stylar head appendages variable or absent. 45 spp. restricted from Peru to Chile and Argentina, centered in the dry east-Andean valleys of Bolivia and northern Argentina in the southern Yungas, except for one up to Ecuador; open forests, forest margins, grasslands, rocky slopes, quebradas, 1,000–5,000 m or Paramo, 3,000–4,500 m.

73. Tweedia Hook. & Arn. Erect herbs or suffrutescent twiners, to 1 m high, orthotropous, sometimes vines; subterranean organs constituting a woody rootstock. 6 spp. from Chile and Argentina, T. brunonis Hook. & Arn. up to Bolivia and Uruguay, in open bushland with creosote bushes and cacti, in semiarid areas on both sides of the Andes, mostly in the Monte from southern Bolivia to C Argentina, in the central Andes of western Argentina, and in the Desert and Matorral of central Chile.

■ SUBTRIBE TOPEINAE ▸ a single genus.

74. Topea H. A. Keller. Twining plants with leaves oblong-lanceolate to lanceolate; basis deeply cordate to auriculate; inflorescences sciadioidal (umbelliform), pendulous, present all the year; corolla lobes with vibratile trichomes on the adaxial side. Two spp., T. aenigma (H. A. Keller) H. A. Keller endemic a small private area in Missiones, Argentina, and T. patens (H. A. Keller) H. A. Keller in Amambay Departamentn in E Paraguay, NE Argentina and Mato Grosso do Sul state, Brazil.

■ SUBTRIBE GONOLOBIINAE ▸ outsiders list unvailable.

75. Atrostemma Morillo. Twining shrubs; branches shortly pubescent; leaves 6–18 cm long, both surfaces sparsely puberulous; inflorescences 3–7-flowered; corolla rotate, yellowish green and dark-reticulate. 10 spp. from mountain forests of Colombia and Venezuela, forests and thickets; 100–1,700 m.

76. Austrochthamalia Morillo & Fontella. 5 spp., Argentina and Bolivia one each, three remaining in Brazil, one of then up to Paraguay, another up to Bolivia.

77. Brargentina Morillo. Only one sp., B. bornmuelleri (Schltr. ex Malme) Morillo & H.A. Keller, from S Brazil and N Argentina.

78. Caa H. A. Keller & Liede. Climbing plants with a thin base, not suberified; young branches with mixed pu-bescence, consisting of multiseptate acicular trichomes and capitate glandular tri-chomes, adult branches puberulous. Only one sp., C. balansae (Fontella & Morillo) H. A. Keller & Liede, with only three known occurrence locations, two in Argentina and one in Caaguazú, Paraguay.

79. Chloropetalum Morillo. (inc. Matelea p.p., Gonolobus p.p.) 4 spp., two endemics from Brazil and two over widely from Honduras to Paraguay, up French Guiana and N Brazil (both).

80. Coelostelma E. Fourn. Erect suffrutex, 30–60 cm high, glabrous all over; leaves shortly petiolate, to 9.5 cm long; inflorescences shortly pedunculate, 2- flowered; corolla rotate-campanulate, acuminate, conspicuously veined. One sp., C. refractum E. Fourn, endemic to Minas Gerais state, E Brazil, known only the type collection.

81. Cristobalia Morillo, S.A.Cáceres & H.A.Keller. Woody shrubby vines; stems 1-5 m long, prostrate or vining, flexuous, terete, not corky at the base, branches densely pubescent throughout. Two spp. from Bolivia, Argentina and Rio Grande do Sul state in S Brazil.

82. Fischeria DC. Suffrutescent twiners, 3-20 m high, latex white. 8 spp., one only in Caribbean, four from Central America to N Colombia, F. polytricha Decne. in Brazil and Bolivia, and two remaining widely distributed; forest margins, disturbed moist montane slopes, damp thickets, secondary growth, often near streams, roadsides; 0–1,000 m, sometimes up to 1,700 m.

83. Gonolobus Michx. (exc. Chloropetalum p.p.) Suffrutescent twiners or, rarely, prostrate herbs, latex white; flowers rotate with exposed discoid head to gynostegium and laterally disposed pollinia, anthers with dorsal appendages; flowers commonly green; follicles with longitudinal wings. 107 spp., widely distributed in New World, 39 in South America, 17 in Colombia, 4 spp. in Brazil, two endemics, G. dorothyanus Fontella & E.A. Schwarz endemic to Rio de Janeiro state and rare plant in Brazil, by Plantas Raras do Brasil’s book.

84. Graciemoriana Morillo. Twining vine, ca. 10 m long; leaves petiolate; blades 18–26 cm long; inflorescences pedunculate, 3–4-flowered; corolla ca. 18 mm diam., rotate-campanulate, green. One sp., G. graciae (Morillo) Morillo, French Guiana, non-flooded moist forest, 200–400 m.

85. Gyrostelma E.Fourn. Subshrubs, orthotropous or plagiotropous. Only one sp., G. oxypetaloides E. Fourn, C Brazil.

86. Ibatia Decne. Suffrutescent to herbaceous twiners or erect herbs; unflorescences extra-axillary, solitary, shorter than adjacent leaves, few- to many-flowered, simple, bostrychoid; corolla rotate to campanulate, white, yellowish-green or purple, uniformly colored or with reddish veins. 30 spp., Colombia and Venezuela, Bolivia, Brazil and Cono Sur, with Ib. maritima (Jacq.) Decne. up to Panamá; a specifc diversity by geographical area is lack duo higly discrepances between VPA and owner works at Gonolobiinae; two spp. from Bahia state are rares by Plantas Raras do Brasil’s book.

87. Lachnostoma Kunth. 9 spp., costal Venezuela to Peru; endemic in Colombia (4), Ecuador (2), Venezuela and Peru one each.

88. Lhotzkyella Rauschert. Twiner with short mixed-pubescence in 2 lines; inflorescences short-pedunculate, 4–6-flowered; corolla ca. 16 mm long, campanulate, dark purple; follicles narrowly ovoid. One sp., L. lhotzkyana (E. Fourn.) Rauschert, endemic to Mato Grosso state, Brazil.

89. Macroscepis Kunth. Suffrutescent twiners, 6-12 m high. 15 spp., Central America, Costa Rica, Guatemala, and over South America (11); 6 spp. in Brazil, 4 endemics.

90. Malinvaudia E. Fourn. Twiner, stems glabrous; leaves shortly petiolate, blades to 13 x 4.5 cm; inflorescences 6–9-flowered; corolla campanulate, throat with 5 interlobular pubescent pads; follicles unknown. Only one sp., M. capillacea E. Fourn., E Brazil and NE Argentina, below 600 m.

91. Matelea Aubl. Suffrutescent twiners (in the tropics), prostrate or erect herbs (in temperate zones) or, rarely, shrubs or caudiciforms, latex white, some succulents in Mexico and Central America. 248 spp. of North America, Central America (centered in S Mexico/Guatemala), South America (mainly); rain and cloud forest margins, thickets, disturbed habitats, to 2,500 m; a specifc diversity by geographical area is lack duo higly discrepances between VPA and owner works at Gonolobiinae; six species from Bahia, Santa Catarina nd São Paulo states are rares by Plantas Raras do Brasil’s book.

92. Orinoquia Morillo Twining subshrub; inflorescences very long-pedunculate, 3–5-flowered; corolla 50–55 mm diam., rotate. Only one sp., O. yanomamica (Morillo) Morillo, endemic to Venezuela, rain forest, headwaters of the Orinoco river, 150–400 m.

93. Peruviasclepias Morillo. Erect, prostrate or short twining herb or subshrub, 0.4–1.5 m long; leaves 2–4.5 cm long; inflorescences 6–12(–24)- flowered; corolla 6–7 mm diam., yellowish green, salverform or short-campanulate; pollinia subhorizontal to erect, oblongoid-pyriform; follicles fusiform, muricate. Only one sp., P. aliciae (Morillo) Morillo, NW Peru, deserts, 100–500 m.

94. Phaeostemma E. Fourn. Twining vines; flowers large and broadly campanulate (corolla 23–37 mm in diameter), corollas green to greenish-yellow, gynostegium stipitate, anthers subtriangular almost horizontal. 8 spp., Venezuela and Suriname one endemic each, and remaining six from N Argentina to SE Brazil (all spp., 5 endemics).

95. Pruskortizia Morillo. Woody shrubby vines, with white latex; stems thick (4-10 mm), flexuous, mixed of brown or yellowish-brown long eglandular (1–8 mm long) and short glandular trichomes (0.1–0.25 mm long); similar pubescence on leaves and inflorescence. Two spp., from W. South America to N Brazil (both species, none endemics).

96. Pseudolachnostoma Morillo. Suffrutex twinners, densely pubescent with yellow trcihomes, white latex, short reflexed calyx lobes, corolla short openly campanulate, and then reflexed, the lobes much longer than the tube, narrowly ovate to oblong. 12 spp., one in Central America, and remaining restricted of South America up to N Brazil (2, none endemics).

97. Rhytidostemma Morillo. Suffrutescent twiners, shoots slender, 2–4 mm diam., densely hispid or pubescent over the whole surface with erect or retrorse, long and flexuous trichomes, yellow. 7 spp., mainly Guiana Shield, all South America, one up to Panamá; three spp. in Brazil, one endemic (R. fontellanum Morillo in Amapá state).

98. Riparoampelos Morillo. Suffrutescent twiner; stems shortly pubescente; leaves 8–13 cm long, lanceolate to obovate-elliptic; inflorescences subsessile, 2–3-flowered; corolla 20–24 mm diam., rotate(–campanulate); lobes dark purplish, spreading, apically with long, flat and white trichomes; pollinia horizontal, asymmetrically pear-shaped; follicles narrowly ovoid or fusiform, muricate, with irregular blunt-tipped projections. Only one sp., Ri. amazonica (Morillo) Morillo, Amazon rainforest and Guianas, wet or rain forests, mainly in riparian vegetation, 100–300 m.

99. Rojasia Malme. Only one sp., R. gracilis (Morong) Malme, N Argentina, Paraguay and SE Bolivia.

100. Schubertia Mart. Suffrutescent twiners with large, urceolate white flowers. 5 spp. from Argentina, Brazil (4, 3 endemics), Paraguay and Bolivia; shady places, forest margins, along rivers.

101. Tressensia H.A. Keller. Herbaceous twiner; inflorescences long-pedunculate, 3–6-flowered; flowers long-pedicellate; corolla ca. 3 cm diam., shallowly campanulate, green-reticulate; follicles obclavate, 5-winged; seeds verrucose, with dentate wing. One sp., T. viridis H.A. Keller & S.A. Cáceres, endemic to Argentina, riverine forest fragments.

56. LAMIALES

FAMILIES ABSENTS IN SOUTH AMERICA: BYBLIDACEAE (1/8), CARLEMANNIACEAE (2/5), MAZACEAE (4/44), PAULOWNIACEAE (2/8), PEDALIACEAE (11/78), PLOCOSPERMATACEAE (1/1), STILBACEAE (12/40), THOMANDERSIACEAE (1/6), WHIGHTIACEAE (1/2).

LINEAGE 1 of 8: OLEOIDS

OLEACEAE

§ FULLY CHLOROTROPHIC

Genera/spp. 29/545–590 Distribution cosmopolitan except polar areas, with their largest diversity in SE Asia and Australia. Habit usually bisexual (rarely polygamomonoecious or dioecious), evergreen or deciduous trees, shrubs or lianas (rarely suffrutices).

The Guinness Book of World Records lists the South African black ironwood (Olea laurifolia Lam.) as the heaviest wood with a specific gravity of 1.49; this is rather doubtful since the specific gravity of pure cell wall material is 1.5 (i.e. without any cellular structure).

SYSTEMATIC 5 tribes, four absent in South America: Forsythieae (2/10, Balkan Peninsula, E Asia), Fontanesieae (1/2, Sicily, Türkiye, Syria, Lebanon; east central China), Myxopyreae (3/7, tropical Asia) and Jasmineae (1/c 200, warm-temperate to tropical regions in the Old World); among Oleeae, outsiders are Ligustrum (c 60; Europe, Mediterranean, northern Africa, E and SE Asia, Malesia to E Australia and Tasmania); Comoranthus (3; Madagascar, Mayotte Island), Fraxinus (40–45; temperate regions on the Northern Hemisphere, few species in tropical regions); Olea (32–33; warm-temperate to tropical regions in the Old World), Haenianthus (3; Greater Antilles), Notelaea (12; Australia, Tasmania), Picconia (1; Madeira, Canary Islands), Noronhia (c 45; Madagascar, the Comoros), Phillyrea (2; Madeira, Mediterranean to Syria and N Iran), Osmanthus (32; SW Asia, China to W Malesia, SE U.S.A., Mexico), Nestegis (5; New Zealand, Norfolk Island, Hawaii), Hesperelaea (1; Guadalupe Island off Baja California; probably extinct).

1. Chionanthus L. Trees or shrubs, evergreen or deciduous; leaves opposite, simple, entire or dentate; inflorescence axillary or terminal, cymose, paniculate or fasciculate; flowers regular, bisexual, rarely unisexual, usually white or yellow, rarely pink, often fragrant; fruit a drupe, 1- or 2-seeded, mesocarp fleshy, endocarp bony or crustaceous, with or without endosperm, bluish or purplish when ripe. 61 spp., C & E U.S.A. to southern America, Africa, Madagascar, Asia, Japan to Pacific; 39 spp. in New World, mainly in Caribbean and E Brazil (12, 10 endemics); 23 spp. in South America; three spp., from Minas Gerais and Rio de Janeiro states, are rare plants in Brazil, by Plantas Raras do Brasil’s book.

C. megistocarpus (Oleaceae, Colombia) has largest fruits for this genus in the Neotropics (4.5–5 cm long).

2. Forestiera Poir. 20 spp., U.S.A. to Central America (mainly U.S.A./Mexico zone), Caribbean, and F. ecuadoriensis Cornejo & Bonifaz restricted in Guayas province in Ecuador.

3. Menodora Bonpl. Herbs to shrubs, sometimes with xylopodium; 27 spp., two in Africa, 18 in North America from Colorado to Mexico, and 7 spp. restricted of Bolivia and Cono Sur except for M. integrifolia (Cham. & Schltdl.) Steud., from Bolivia to Uruguay, Rio Grande do Sul state in S Brazil, Argentina and Paraguay.

4. Priogymnanthus Green. Trees, deciduous or semi-deciduous; leaves opposite, simple, entire; inflorescence a racemoid reduced dichasium, up to 11-flowered, shortly pedicellate, flowers and subtending bracts in opposite pairs, the rachis terminated by three flowers, calyx absent, corolla 4-lobed, lobes thin, slender, very early caduceus; fruit a drupe, ellipsoid or more or less spherical. Three spp., P. apertus (B. Ståhl) P.S. Green in Ecuador, P. hasslerianus (Chodat) P. S. Green, from hillside in C Brazil (Tocantins to Mato Grosso states), Bolivia, Argentina and Paraguay, and P. saxiculous Loambardi, endemic to Minas Gerais state, Brazil.

5. Schrebera Roxb. 8 spp., highly disjunct, 7 spp. in tropical Africa, Madagascar, SE Asia, and S. americana (Zahlbr.) Gilg in Western Hemisphere, endemic to Peru.

LINEAGE 2 of 8: TETRACHONDRACEAE

TETRACHONDRACEAE

§ FULLY CHLOROTROPHIC

Genera/spp. 2/3 Distribution North America to northern South America, Patagonia and New Zealand. Habit herbs.

SYSTEMATIC all genera in South America.

1. Polypremum L. Only one sp., P. procumbens L., from SE U.S.A., Mexico, Caribbean, Central America, northern South America in Colombia and Venezuela; a single disjunct population is reported from Paraguay, but it is not known whether it is the result of human introduction.

2. Tetrachondra Petrie. Aquatic or semiaquatic herbs. Two spp., the succulent T. hamiltonii Petrie ex Oliv. from southern New Zealand and T. patagonica Skottsb. from southern Patagonia and Tierra del Fuego.

LINEAGE 3 of 8: GESNERIOIDES

CALCEOLARIACEAE

§ FULLY CHLOROTROPHIC

Genera/spp. 2/274 Distribution Calceolaria: Mexico, Central America, western South America (especially the Andes and westerly temperate regions), the Falkland Islands, one reaching into SE Brazil; Jovellana: New Zealand and Chile. Habit bisexual, perennial or annual herbs, suffrutices or shrubs.

SYSTEMATIC both genera occur in South America.

1. Calceolaria L. Annual or perennial herbs and shrubs up to 4m, usually yellow bilabiate flowers, sometimes cushions; stems escending to erect, rounded, corolla labiate with down petal sacciform, and two stamens. 270 spp. from Mexico to Terra do Fogo, the widely distributed C. tripartita Ruiz & Pavon. reaching into S Brazil, all but nine (reaching into Mexico) endemic to South America (only central-american C. irazuensis Donn. Sm. absent); occurs mainly at high altitudes along the Andes but reaches sea level on both the Atlantic and Pacific coasts of Patagonia, mainly in Peru (128, 89 endemics), Cono Sur (85, 66 restricteds) and Colombia (23). Calceolaria is among the largest oil-producing genera - nonvolatile, a very unusual floral reward that attracts particular solitary oil-collecting bees; with high diversity in Chile and Ecuador/Peru border; Calceolaria has been subdivided into three subgenera with 24 sections:

§ subg. Calceolaria ‣ 20 sections; shrubs, subshrubs, and herbs that have stamens with filaments shorter than anthers, mainly distributed in tropical regions.

§ subg. Cheiloncos ‣ two sections, Micranthera and Rugosae; also includes herbs and subshrubs, but their stamens have filaments that are several times longer than their anthers; temperate South America.

§ subg. Rosula ‣ three sections, Bellidifoliae, Corymbosae and Kremastocheilos; rosulate or subrosulate herbs that have stamen filaments about as long as the anthers; temperate South America.

2. Jovellana Ruiz & Pav. Herbs and shrubs, perennial, stems prostrate or ascending to erect, rounded. 4 spp., Chile and New Zealand two endemics each.

PELTANTHERACEAE

§ FULLY CHLOROTROPHIC

Genera/spp. 1/1 Distribution Guatemala to Bolivia.Habit bisexual evergreen small tree or shrub.

UNPLACED LAMIALES (1/1) ‣ a single genus.

1. Peltanthera Benth. Small trees, glabrous or with branched hairs, with abundant, aromatic, 5-merous, actinomorphic flowers in cymes; bracts squamiform, often obsolete; calyx equal, short, and with free sepals; corolla hypocrateriform, with valvate petals; anthers confluent, peltate; gynoecium with two locules, filiform style, and capitate stigma; leaves opposite, elliptic, wide, membranaceous, penninerved, slightly serrulate. Only one sp., P. floribunda Bent., from Costa Rica to Bolivia.

GESNERIACEAE

§ FULLY CHLOROTROPHIC

Genera/spp. 151/3,260–3,280 Distribution tropical and subtropical regions in the Northern and Southern Hemispheres; some spp. in temperate regions.Habit usually bisexual (rarely monoecious), usually perennial herbs (sometimes lianas, rarely trees, shrubs or annual herbs). Many spp. are epiphytic. Stem and leaves often more or less succulent. Root fibrous. Use some genera are cultivated as ornamentals (mostly indoor plants), and among the most popular are Achimenes, Codonanthe, Columnea, Gloxinia, Episcia, Kohleria, Nematanthus, Sinningia, Seemannia; plants are easy to propagate by cuttings or by seeds; they also hybridize easily within related groups and many cultivars are available.

About 1,200 spp., ca. 60 genera in rain forest on mountain slopes with main diversity center between Colombia and Peru (over 400) and secondary centers in Central America (ca. 250), E Brazil (over 200), Guiana shield (ca. 100) and tropical Mexico (ca. 100). Brazil has 207 spp., and 24 genera, all are Gesnerieae. Some genera are cultivated as ornamentals (mostly indoor plants), and among the most popular are Achimenes, Codonanthe, Columnea, Gloxinia, Episcia, Kohleria, Nematanthus, Sinningia, Seemannia. The flower of Nematanthus has petals fused into a pouch-like shape, with a small opening; the fancied resemblance of the flower to a goldfish gives the plant its common name, ‘goldfish plant.’ Compared to other regions that are especially rich in species of Gesneriaceae such as Colombia (400), the Guiana Shield, combined with a narrow part of northern Brazil, has a surprisingly large number of endemic genera.

All New World has inflorescence formed by an indeterminate thyrse with axillary pair-flowered cymes.

Key differences from similar families

Stipules never present, leaves never compound (vs. Bignoniaceae with compoubd leaves).

Latex never present (vs. Campanulaceae which usually has white latex).

SYSTEMATICS four subfamilies, three in South America (one endemic), Didymocarpoideae (62/1.760–1.980, southern Europe, India, Sri Lanka and southern China (inc. Taiwan) to Japan, Malesia and tropical Australia, Solomon Islands and islands in the Pacific to Hawaii, few species in tropical Africa and Madagascar) absent in New World.

1. SUBFAMILY SANANGOIDEAE (1/1) ‣ a single genus.

1. Sanango Bunting & Duke. Small tree, c. 15 m tall, with very hard wood; inflorescence a terminal, bracteose thyrse; corolla tubular, tube curved and slightly gibbous in the lower part, limb subregular, lobes rounded, with cochlear aestivation in bud; fruit a capsule, depressed at the apex, style (for a long time) persisting, dehiscence at first septicidal, then loculicidal. Only one sp., S. racemosum (Ruiz & Pav.) Barringer, in forests at 300-750 m altitude, Peru (W central Amazonas and C Junin) and Ecuador.

2. SUBFAMILY EPITHEMATEAE (7/c. 80) ‣ outsiders Whytockia (8; S China inc. Taiwan), Monophyllaea (>30; peninsular Thailand, Malesia to New Guinea), Stauranthera (3; India and southern China to SE Asia, Malesia to New Guinea), Loxonia (3; the Malay Peninsula, Sumatra, Java, Borneo), Gyrogyne (1; Guangxi in southern China), Epithema (22; NE India and Nepal to southern China (inc. Taiwan) and SE Asia, Malesia to New Guinea, one species, E. tenue, in tropical W and C Africa).

2. Rhynchoglossum Blume. Herbs, perennial or annual, monocarpic, the very strongly asymmetrical leaves; stems terete, fleshy-succulent; inflorescences terminal on main and side branches, unilateral racemes with two rows of flowers, subtending bracts small, linear; corolla strongly zygomorphic; tube cylindrical, white; fruit a globose or ovoid capsule. 3-4 spp., R. azureum (Schltdl.) B.L. Burtt from Mexico to Honduras, Costa Rica to Peru, remaining in India and southern China to New Guinea, on wet and shady (preferably limestone) rocks, in forest or open, shady places; usually in the lowlands.

3. SUBFAMILY GESNERIOIDEAE (1/23) ‣ 5 tribes, four in South America, Titanotricheae (1/1, SE China (inc. Taiwan), southern Japan, the Ryukyu Islands) does not occur in South America.

3.1 GESNERIOIDAE ▸ TRIBE NAPEANTHEAE (1/23) - a single genus.

3. Napeanthus Gardner. Terrestrial, perennial herbs with rhizomes; stem short, decumbent, rooting at the nodes, or ± absent; axillary cymes 1- to many-flowered, pedunculate, subumbellate to racemose-paniculate; flowers actinomorphic to zygomorphic; corolla white to blue, pink or lilac; fruit a capsule. 16 spp., 15 in South America (three up to Central America and Mexico), many more to be expected; low forest herb, growing in shady, damp places and on mossy rocks, throughout the neotropics; 4 spp. in Brazil, N. jelskii Fritsch and N. macrostoma Leeuwenb. up to Guianas, remaining two endemics.

3.2 GESNERIOIDAE ▸ TRIBE BESLERIEAE (9/c. 275) – two subtribes, both in South America.

■ SUBTRIBE BESLERIEAE – all genera in South America.

4. Besleria L. Perennial herbs, shrubs or small trees (to 5 m), with fibrous roots; stem terete or quadrangular; inflorescences axillary, ebracteate, fasciculate or subumbellate cymes, rarely flowers solitary; corolla yellow, orange, red, or white; tube usually cylindric, sometimes spurred or gibbous at base; limb (sub)regular or bilabiate; fruit a fleshy, globose berry; white, red or orange. 163 spp., throughout the neotropics, with centres of diversity in the Andes of Colombia (41 endemics) and Ecuador, some taxa endemic in SE Brazil (26, 13 endemics, B. meridionalis C.V.Morton from Minas Gerais state is a rare plant in Brazil, by Plantas Raras do Brasil’s book); most species are endemic with very local distribution; terrestrial plants growing in lowland and montane forests, often along streams, on river-banks, on wet rocks etc.; 139 spp. in South America; the adnate inflorescences at leaf limbe in some species from Central America is a false epiphylly. B. formicaria Nowicke, from Costa Rica to Colombia is a myrmecophite.

5. Cremosperma Benth. Small terrestrial or epiphytic perennial herbs; stems soft woody, short, erect, ascending or repent; inflorescences few, emerging from the upper leaf axils, umbellate or capitate cymes, usually longer than the leaves; ebracteate; flowers small; corolla small (2 cm or less), yellow, white or reddish, occasionally spotted; fruit a capsule, dehiscing irregularly. 24 spp., from the Andes of Venezuela to Peru, two up to Panamá, growing in wet humus or moss on slopes, rocks, or the moss cover of trees, usually at higher elevations.

6. Gasteranthus Benth. Terrestrial herbs or low subshrubs, with fibrous roots; stem succulent or woody, erect, terete; inflorescences axillary, ebracteate, one to many-flowered cymes, peduncles usually elongate; corolla ranging from funnelform with a wide limb over broad tubular to hypocyrtoid; colour whitish, pale to bright yellow, orange to red, often spotted; fruit a 2- or 4-valved capsule. 41 spp., from Guatemala and nearby Mexico (only one), Costa Rica and Panamá through western South America (37) south to Bolivia, with a centre of diversity in western Ecuador (20 endemics); terrestrial herbs of forests (including cloud forests), growing in shaded and humid places, in ravines or near waterfalls, from the lowlands to montane forests to 1,800 m.

7. Reldia Wiehler. Terrestrial herbs with creeping rhizomes; stem terete; inflorescences axillary, often several in a leaf axil, epedunculate (?), ebracteate; corolla small, funnel-shaped, usually white, often yellow in the throat; tube dorsally spurred; limb unequal; fruit a dry, bivalved capsule, surrounded by the persistent calyx. 7 spp., from Panamá (one endemic) to northern Peru, growing in montane (more rarely lowland) rainforests, in humid, shaded ravines along streams or close to waterfalls.

■ SUBTRIBE ANETHANTINAE – all genera in South America.

8. Anetanthus Hiern ex Benth. Terrestrial or saxicolous herbs; stem decumbent or pendent, slender; axillary cymes several-flowered, pedunculate; corolla white, bluish, violet or red, tube subcylindrical, limb with orbicular, slightly spreading lobes; fruit an elongate ovoid capsule, 2-valved, dehiscence septicidal (plus short loculicidal slits); seeds suborbicular, flattened, narrowly winged. Two spp. from Colombia to Bolivia, A. gracilis Hiern up to Brazil, and A. disjunctus L.E. Skog & J.L. Clark in Guyana.

9. Cremospermopsis L. E. Skog. & L. V. Kvist. Perennial herbs to subshrubs, stem usually erect; cymes axillary, pedunculate, usually congested to umbel-like aggregates, mostly with 4 lanceolate to ovate bracteoles; corolla small, white or yellow, funnelform to tubular; fruit a dry capsule. 3 spp., endemic to Colombian departments of Antioquia and adjacent Bolivar, in humid (probably low elevation cloud) forests, in shady places and often close to streams, from (0-)500-900(-1,550)m.

10. Resia H.E.Moore. Terrestrial perennial herbs or subshrubs, with erect stem (eventually becoming pendent), unbranched stem roots fibrous; cymes axillary, long-pedunculate, flowers in a dense head; corolla white, orange or yellow, infundibuliform; fruit an ovoid or subglobose, laterally compressed dry capsule. 4 spp., Colombia, Venezuela and Ecuador; the species are reported to grow on moist shaded rocks in mountains, alt. 1,000-1,350 m.

11. Shuaria D.A.Neill & J.L.Clark. Small tree, 3–5(-8) m tall, frequently with multiple trunks arising from the tree base; inflorescences axillary, emerging from the upper leaves, pair-flowered dichasial cymes, ebracteolate; corolla white, tubular, gibbous at the base and shallowly pouched on ventral side; fruit a bivalved dry capsule. Only one sp., S. ecuadorica D.A.Neill & J.L.Clark., Cordillera del Cóndor and Amazonian regions of SE Ecuador, growing in lowland rain forest and lower montane cloud forest, with an elevation range 420-1,600 m.

12. Tylopsacas Leeuwenb. Terrestrial herbs; stem ± absent; cymes unilateral (cincinni), with numerous flower pairs; corolla white, tube slightly inflated in the middle, limb slightly bilabiate, with elliptic, rounded lobes; stamens 4, didynamous, included; filaments inserted near corolla base; anthers reniform, coherent in pairs. Only one sp., T. cuneata (Gleason) Leeuwenb, endemic to the Guiana Shield, growing in montane forests (100-1,700 m elevation range) of Venezuela, Guyana, and in N Brazil.

3.3 GESNERIOIDAE ▸ TRIBE CORONANTHEREAE (9/23) - three subtribes, Coronantherinae (2/13–20; New Caledonia, Solomon Islands, eastern Queensland, North Island in New Zealand), and Negriinae (2/3, New Caledonia, NE Queensland, Lord Howe) do not occur in South America; among Mitrariinae, the solely in South America, outsider is Fieldia (1; Queensland, New South Wales, Victoria).

13. Asteranthera Hanst. Epiphytic or terrestrial, creeping-climbing shrubs; stem and branches thin, woody, rooting at the basal nodes; flowers axillary, solitary, erect; bracteoles small, lanceolate, inserted below the calyx; corolla red, with whitish lines on tube, lobes mottled, infundibuliform-galeate; fruit a fleshy berry. Only one sp., A. ovata Hanst. southern Chile and adjacent Argentina, growing as an epiphyte or terrestrially in cool rain forests.

14. Mitraria Cav. Straggling or epiphytic subshrub; stem and branches softly woody, obscurely tetragonous; flowers axillary, solitary, pendulous; bracts large, ovate, connate or one side, embracing the calyx on one side; corolla bright scarlet; tube curved; fruit a fleshy berry. Only one sp., M. coccinea Cav., Chile and adjacent Argentina, Chile and adjacent Argentina, occurring in cool temperate rain forest.

15. Sarmienta Ruiz & Pav. Suffrutescent creeper or climber, usually epiphytic; stems slender, flexuous, sparingly branched, rooting at the basal nodes; flowers axillary, solitary, pendulous; corolla scarlet; tube elongate, 2 stamens lang up outside corolla; fruit a fleshy berry. Only one sp., S. scandens (J.D. Brandis ex Molina) Pers., southern Chile and Chiloe Island, occurring in cool rain forest, growing on damp or mossy rocks and tree trunks.

3.4 GESNERIOIDAE ▸ TRIBE GESNERIEAE (53/1,500) - five subtribes, all in South America.

■ SUBTRIBE GESNERIINAE ‣ outsiders Bellonia (2; Hispaniola, Cuba), Gesneria (45–50; Caribbean), Pheidonocarpa (1; Cuba, Jamaica); Gesneria L. revised shown a only Caribbean range; their two South American species is placed now in Rhytidophyllum Mart.

16. Rhytidophyllum Mart. (off Gesneria) Shrubs to trees, usually single-stemmed; axillary cymes long-pedunculate, of several to many flowers in a double cincinnus or compound dichasium; flowers rather short-tubed, obliquely campanulate, with wide mouth, limb spreading, greenish, brownish, with darker dots; fruit a dry, bivalved capsule. 24 spp., 22 spp. restricted of Caribbean region and two in South America, R. cumanense (Hanstein) L. Skog and R. onacaense (Rusby) L. Skog., both in Colombia and Venezuela.

■ SUBTRIBE GLOXINIINAE ‣ outsiders Moussonia (11; S Mexico to Panamá), Eucodonia (2; C and S Mexico), Smithiantha (8; Mexico, Guatemala), Niphaea (2; Mexico, Guatemala), Solenophora (16; Central America).

17. Achimenes Pers. Terrestrial herbs with scaly rhizomes; stems erect or decumbent, unbranched; inflorescences axillary, of single flowers or two to three in a cyme; corolla usually showy, erect or oblique in the calyx, either tubular with a flat spreading limb, or obliquely funnelform, with a basal sac or spur on the upper side. 27 spp., usually found in wet areas in forests, on shaded banks near streams or on damp rock outcrops, from Mexico and C America, A. erecta (Lam.) H.P. Fuchs and A. longiflora DC. extending to Colombia, and A. pedunculata Benth. up to Venezuela.

18. Amalophyllon Brandegee. Terrestrial or saxicolous, subacaulescent to caulescent herbs with scaly rhizomes; stem erect or decumbent, usually unbranched; inflorescences axillary, with (1–)4 flowers congested in the axil; corolla 5-lobed, erect in the calyx, rotate to subrotate with a very short tube, white or rarely tinged pink. 6 spp., S Mexico to Panamá and South America (3, two exclusively in Colombia, A. albiflorum (Rusby) Boggan, L.E. Skog & Roalson from Central America up to Andes from the E range of Venezuela.

19. Chautemsia Chautems. Perennial herbs with scaly rhizomes; stems erect, unbranched, usually comprising only 3–4 nodes; flowers solitary in leaf axils; corolla infundibuliform-cylindrical, tube white to white-yellowish; throat not constricted, yellow; lobes white; capsule ellipsoid, apex erect, fleshy. Only one sp., Chautemsia calcicola A.O. Araujo & V.C. Souza, only found in the localities Arcos, Pains and Iguatama in the region of Formiga, in the western part of Minas Gerais, in deciduous forest remnants on limestone outcrops at 600-800 m.

20. Diastema Benth. Slender, terrestrial, perennial herbs with scaly rhizomes; stem short, unbranched, hirsute or villous; inflorescence usually a terminal, bracteose raceme; corolla white, sometimes with purple markings on the lobes, tubular or funnelform, only slightly broader toward the limb, neither spurred nor ventricose; fruit an obovoid, membraneous, 2-valved capsule, with apex convex. 17 spp., all in South America (southern into Bolivia), 4 up to Mexico and Central America, centre of diversity in Andean Colombia, Ecuador and Peru, as terrestrial herbs growing on damp rocks near streams, in wet forests at low elevations; only D. racemiferum Benth. in Brazil, in Rondonia, possibly in Acre state.

21. Gloxinella Roalson & Boggan. Terrestrial, rather weak-stemmed perennial herbs with scaly rhizomes; stem erect, indumentum villous, without uncinate hairs; flowers axillary solitary, epedicellate and ebrbracteolate, showy; corolla alvender, campanulate, limb of subequal lobes. Fruit an ovoid to elliptic fleshy capsule. Only one sp., G. lindeniana (Regel) Roalson & Boggan; for long, the species was known only from cultivated material of unknown origin, but in 1990’s it has been re-discovered in Peru (Cajamarca).

22. Gloxinia L'Her. (inc. Anodiscus) Terrestrial perennial herbs with scaly rhizomes (absent in G. xanthophylla); stem erect; leaves opposite, ± isophyllous, rarely ternate, with 5-9(-12) pairs of veins; stem and leaves almost glabrous or pilose; flowers solitary, forming together a terminal, indeterminate raceme; corolla broadly tubular to campanulate, white, pink, purple or brownish. 4 spp. G. alterniflora A.O.Araujo & Chautems endemic to Mato Grosso do Sul state, Brazil, G. xanthophylla (Poepp.) Roalson & Boggan endemic to Peru and Ecuador, and G. erinoides (DC.) Roalson & Boggan and G. perennis (L.) Fritsch widely distributed tropical America, growing in small or large colonies on rocks, on riverbanks, or in shaded damp places in forest; the plants are dormant during winter, reduced to the underground rhizomes.

23. Gloxiniopsis Roalson & Boggan. Terrestrial perennial herbs with scaly rhizomes; stem erect; flowers solitary, emerging from the axils of opposite or alternate bracts, forming together a terminal, indeterminate raceme; pedicels ebracteolate; corolla white, campanulate; fruit a subglobose fleshy capsule; seeds numerous, minute, rhombic to ellipsoid. Only one sp., G. racemosa (Benth.) Roalson & Boggan, in the Andes of Colombia.

24. Goyazia Taub. Small, perennial, saxicolous herbs with scaly rhizomes; stem thin, stiff, wiry; flowers axillary, solitary, with short pedicel; corolla with straight, subcampanulate tube, slightly widening towards the throat, limb oblique, with subequal lobes; fruit a dry, bivalved capsule. 3 spp., terrestrial herbs growing on damp and mossy rocks; dormant in winter in C Brazil from Pará to Minas Gerais and from Mato Grosso do Sul to Maranhão.

25. Heppiella Regel. Epipetric or (less commonly) epiphytic, perennial, rhizomatous, ± suffruticose herbs. Axillary cymes with or without peduncle, sometimes composing a terminal, corymboid thyrse, or reduced to single flowers; corolla red; tubular, straight, limb with small (sub)equal lobes; fruit a dry, bivalved capsule with loculicidal dehiscence. 4 spp., Peru to W Venezuela, usually growing on damp, mossy rocks, rarely epiphytic at the base of trees.

26. Kohleria Regel. (inc. Capanea) Terrestrial, perennial herbs or subshrubs with scaly rhizomes; stem usually erect, sometimes decumbent, terete; inflorescences from leaf or bract axils, in few-flowered, fasciculate to umbellate cymes, or flowers solitary; corolla somewhat zygomorphic, red or greenish; fruit a dry capsule, dehiscing into 2 valves, apex conical or rostrate. 22 spp., W Cordillera from Peru to Mexico, Venezuela, Trinidad, K. hirsuta (Kunth) Regel up to Guyana, Suriname and French Guiana; the centre of diversity in Colombia; 19 in South America; growing in sun-exposed margins or glens of rain forest, also in secondary habitats.

27. Mandirola Decne. Terrestrial perennial herbs with scaly rhizomes; stem erect; leaves opposite or ternate, petiolate, lamina with 5-6 pairs of veins; flowers in axillary, bracteolate pair-flowered cymes, often with a short peduncle, or solitary, showy; corolla pink, lavender or purple, hypocrateriform, with an oblique limb, usually distinctly toothed or fimbriate; fruit a dry rostrate capsule. 3 spp., endemic to C Brazil.

28. Monopyle Moritz ex Benth. Terrestrial, perennial herbs, with scaly rhizomes; stem erect or decumbent, terete, slender, usually branched; flowers in a terminal thyrse or raceme, emerging from small bracts; corolla somewhat zygomorphic; tube open-campanulate; limb 5-lobed, flaring, white or white with blue, sometimes yellow in the throat. 22 spp., along the W Cordillera from Bolivia to Guatemala, 16 in South America, only M. reflexa (Rusby) Roalson & Boggan in Brazil, in Acre state.

29. Nomopyle Roalson & Boggan. Terrestrial perennial herbs with scaly rhizomes; stems weak, erect or decumbent, glabrescent; flowers axillary, solitary, ebracteolate; corolla campanulate to almost rotate, white to lavender, lobes subequal, entire. Only one sp., N. dodsonii (Wiehler) Roalson & Boggan, in Ecuador and Peru, terrestrial forest plants, seasonally dormant.

30. Pearcea Regel. (inc. Parakohleria) Terrestrial, perennial herbs, usually with conspicuous indumentum, occasionally with stolons, rarely with scaly rhizomes; cymes congested, pedunculate (rarely without peduncle), emerging from the axils of frondose leaves; corolla bent downward at base, rarely bilabiate, usually villous or pilose outside; red, often basally and ventrally yellow, rarely entirely yellow. 21 spp., in E Andean slopes and adjacent lowlands from northern Colombia through Ecuador and Peru to NW Bolivia, P. sprucei (Britton ex Rusby) L.P. Kvist & L.E. Skog occurring nearly in the total range of the genus; all other spp. are much more local; centres of diversity are the northern and C Ecuadorian Amazon region and C Peru; occurring in lowland rainforest (below 700 m) to cloud forest above 2,500 m, particularly rich in species and individuals are the lower montane forests.

31. Phinaea Benth. Terrestrial, perennial herbs with scaly rhizomes; stem short, erect; axillary cymes 1- to many-flowered, congested; calyx campanulate, with spreading equal or subequal lobes; corolla subrotate or cup-shaped, tube short, limb of spreading or erect, broad, subequal lobes; fruit a somewhat fleshy or dry 2-valved capsule, opening loculicidally. 6 spp., from Mexico, Central America, Caribbean, P. albolineata (Hook.) Benth. ex Hemsl. in Colombia and Pará state in N Brazil, P. divaricata (Poepp.) Wiehler in Ecuador and Peru, and P. ecuadorana Wiehler endemic to Ecuador; terrestrial herb growing in forests, seasonally dormant.

32. Seemannia Regel. Terrestrial perennial herbs with scaly rhizomes, often produced at the tips of long stringy rhizomes; stem erect or decumbet; flowers axillary, usually solitary (except S. sylvatica), showy; corolla tubular or inflated, often constricted at the mouth, red, orange, purple, rarely yellow; with barrel-shaped multicellular trichomes at the tube entrance; fruit a dry rostrate capsule. 4 spp., from Ecuador to Cono Sur, mainly in the Andes, S. purpurascens Rusby and S. sylvatica (Kunth) Hanst. up to Brazil, the former also in Guianas; terrestrial herbs, growing on earth banks or rocks in forest; seasonally dormant.

■ SUBTRIBE COLUMNEINAE ‣ outsiders Cobananthus (1; Central America), Oerstedina (2; Central America), Rufodorsia (4; Costa Rica, Panamá); second largest group of epiphytes spp. in eudicots (first is Ericaceae); only three genera outside this tribe in Gesneriaceae has epiphytic spp. (Sinningia, and two paleotropical genera).

33. Alloplectus Mart. Epiphytic subshrubs with fibrous roots; stems terete or quadrangular, usually scandent, prolifically branched; flowers densely crowded or solitary, non-resupinate, corolla oblique to horizontal in the calyx, tubular and usually ampliate, yellow or red. 13 spp., A. calochlamys Donn. Sm. in Mexico and Central America, and 12 from Venezuela to Bolivia, obligately epiphytic, growing in moist and wet forests, usually occurring at higher elevations (500-)2,000-3,500 m.

34. Alsobia Hanst. Stoloniferous epiphytic herbs; stem creeping or pendent, with one stolon per node in the axils of alternating leaves, the subsequent stolons giving the appearance of a single pendent stem with numerous plantlets in a series; leaves with a several-layered hypodermis on the adaxial side. 4 spp. from Mexico and Central America, A. dianthiflora (H.E. Moore & R.G. Wilson) Wiehler up to Colombia.

35. Centrosolenia Benth. Herbs, terrestrial or saxicolous; stems elongate, decumbent, rooting and branching at the nodes; inflorescences axillary, 2–8-flowered; corolla oblique in calyx, narrowly tubular, gibbous basally on upper surface with slight ovate-oblong spur, white, violet, or deep red, corolla lobes orbicular. 15 spp., endemic to the Guiana Shield, 8 exclusively Venezuela, only C. hirsuta Benth. up to Brazil, most are found growing on shady banks of rivers and streams, in crevices or on wet mossy rocks, or in the understory of the rainforests; in the Guiana region, they predominantly inhabit the slopes of the high tepuis, the summit areas of low elevation tepuis, or the understory of the forested high plains at 300-1,500 m.

36. Chrysothemis Decne. Terrestrial or epiphytic perennial herbs with tubers; stems succulent, erect or rarely decumbent, subquadrangular; cymes axillary, corolla tube cylindric; fruit a fleshy, bivalved capsule, globose to ovoid, included in the persistent calyx. 9 spp., Caribbean, Mexico (Chiapas), Guatemala to Colombia, Ecuador, Venezuela, Guyana, Surinam, French Guiana and Brazil (3, only Amazonas state, C. kuhlmannii Hoehne endemic), 8 in South America; C. pulchella (Donn ex Sims) Decne. in over range of genus; growing in shaded areas along roads and streams of wet forests.

37. Christopheria J. F. Smith & J. L. Clark. (off Episcia) Terrestrial stoloniferous herb with two stolons per node; inflorescences axillary, pedunculate, flowers few to many in cymes; corolla posture oblique relative to calyx, yellow with 4–5 red spots ventrally in throat; fruit a bivalved capsule. Only one sp., C. xantha (Leeuwenb.) J. F. Smith & J. L. Clark, known from forests in French Guiana and Guyana at 50-500 m.

38. Codonanthe (Mart.) Hanst. Epiphytic shrubs, lianas or herbs, usually growing in ants' nests; stems pendent, repent, or erect, to 2 m tall, becoming woody, branches few; nodes producing sometimes adventitious roots; cymes axillary, 1- to few-flowered; corolla white, pink, lilac, yellow, or deep purplish, often with reddish lines or spots. 8 spp., in S & E Brazil all endemic of Atlantic Forest; usually growing as epiphytes in ants' nests, in lowland and montane forests; some species form a constituent element of the flora of ant's nests; the plants are associated with the ants in various ways; they often have extrafloral nectaries on the leaf undersides, between the calyx lobes, or at the nodes; the colored seeds exhibit a gelatinous surface or the funicles may play a role in ant dispersal.

39. Codonanthopsis Mansf. (inc. Codonanthe p.p.) Epiphytic lignescent herbs or subshrubs; stem terete, thickish; cymes few-flowered, shortly fasciculate; flowers small; corolla white with purple, or lilac, tube cylindrical, with a dorsal spur at the base, limb with short lobes. 13 spp., 10 in South America, along the river systems of the Amazon region from Peru to Brazil (5, none endemics), and in Central America and Caribbean, growing epiphytically in lowland and montane forests, sometimes associated with ants.

40. Columnea L. (inc. Dalbergaria, Pentadenia) Epiphytic herbs or shrubs (less commonly epipetric or terrestrial), shoots often conspicuously dorsiventral; cymes axillary, 1- to 10-flowered; corolla often red, less commonly yellow, cream or greenish, tube frequently ventricose, basally often dorsally gibbous, usually glandular hairy in throat, limb with subequal lobes to strongly bilabiate, lobes 5 or 4, usually rounded. 217 spp., from Mexico south to Ecuador and Bolivia to Brazil (6, C. ulei Mansf. endemic, from Ceará state, a rare plant in Brazil, by Plantas Raras do Brasil’s book), 141 in South America; 98 spp. in Colombia, 38 in Costa Rica.

41. Corytoplectus Oerst. Terrestrial herbs; stem suffrutescent or fleshy; leaves opposite, ± isophyllous; axillary cymes erect, several- to many-flowered, umbel-like or congested; corolla erect in the calyx, tubular with some ventral inflation, constricted in the throat; limb narrow, with short, (sub)equal lobes. Fruit a globose, shiny black or translucent berry with black seeds. 13 spp., one in Mexico, remaining 12 in South America, from Guianas to Bolivia, mainly growing in cloud forests at higher elevations of the Western Cordillera and Guiana Shield; only C. congestus (Linden ex Hanst.) Wiehler in Brazil, in Amazonas state.

42. Crantzia Scop. Epiphytic or terrestrial subshrubs or coarse herbs with fibrous roots; stems terete or quadrangular, richly branched; cymes axillary, few-flowered, bracteolate; flowers resupinate (except C. tigrina); corolla oblique to horizontal in the calyx, tubular, yellow or red, not puched, limb of usually 5 small lobes; fruit a fleshy bivalved capsule surrounded by the persistent calyx. Two spp., C. cristata (L.) Scop. in the Caribbean and C. epirotes (Leeuwenb.) J.L. Clark on E Venezuela and W Guyana, as epiphytic climbers, or terrestrials; growing in moist and wet forests.

43. Cremersia Feuillet & L. E. Skog. Terrestrial, caulescent herbs; leaves opposite; cymes axillary, pedunculate, bracteolate, flowers pedicellate; corolla salverform, bilabiate, 5-lobed; fruit a dry, bivalved capsule, dehiscing loculicidally, opening to 180°, seeds with short, thick funicles. Only one sp., C. platula C.Feuillet & L.E.Skog, known only from central French Guiana; the collections were made in a mid-elevation primary rainforest where the plants were growing on a talus of granitic boulders or a cliff.

44. Drymonia Mart. Terrestrial or epiphytic (sub)shrubs or lianas; stems quadrangular or terete, in climbers up to 5 m long, branched or not, often with spreading adventitious roots along the internodes; axillary cymes 1- to several-flowered, flowers clustered; flowers often showy and brightly coloured; corolla usually funnelform and broader toward the mouth, spurred or saccate at the base, throat broad; fruit a fleshy capsule with brightly orange- or purple-coloured reflexed valves, displaying a cone-shape mass of seeds and pulpy funicles. 82 spp., throughout the neotropics, with centre of distribution in Colombia and Ecuador, 58 in South America, terrestrial or epiphytic shrubs and subshrubs, growing in lowland and montane forests; 10 spp. in Brazil, none endemics.

45. Episcia Mart. (exc. Christopheria) Stoloniferous, terrestrial or saxicolous herbs; stem creeping, rooting at nodes, dichasially branched by the regular production of stolons; cymes of 1-6 axillary flowers, peduncles slender; corolla white, yellow, blue, purple or red, inserted horizontally in the calyx, zygomorphic salverform to campanulate, fruit an ovoid bivalved fleshy capsule. 9 spp. from tropical South America (8, 7 restricted) to Mexico and the Guianas, in tropical forests, in damp places, slopes, banks or rocks, usually at low elevations, partly forming large colonies; 3 spp. in Brazil, no endemics.

46. Glossoloma Hanst. Terrestrial (rarely epiphytic) subshrubs or coarse herbs with fibrous roots; flowers densely crowded or solitary, resupinate; corolla oblique to horizontal in the calyx, tubular and usually ampliate, yellow or red, constricted at the throat, often ventricose below the mouth, limb of usually 5 small lobes; fruit a fleshy bivalved capsule surrounded by the persistent calyx. 28 spp., from S Mexico to NW South America (24), south to Bolivia, also Venezuela, terrestrial (rarely epiphytic); growing in moist and wet forests, mostly at higher elevations.

47. Lampadaria Feuiilet & L. E. Skog. Terrestrial, caulescent herbs, with short internodes; leaves opposite; cymes axillary, long-pedunculate, bracteolate, flowers short-pedicellate; corolla campanulate, slightly bilabiate; fruit a somewhat fleshy capsule, bivalved, dehiscing loculicidally. Only one sp., L. rupestris Feuillet & L.E.Skog., known only from the north of the Potaro-Siparuni Region in Guyana. One collection was made about 10 km north of Mt. Wokomung at 650 m near a stream; the type collection about 5 km SE of Mt. Ebini, 275 - 350 m in wet rainforest where the plants were growing on rocks.

48. Lembocarpus Leeuwenb. Terrestrial, perennial herb with a small, annual, hairy subterranean tuber, from which (per season) a single leaf emerges; inflorescence separate, cymosely branched or reduced to a single flower; corolla broadly tubular to campanulate; pale blue or white with lilac or purple limb. Only one sp., L. amoenus Leeuwenb., endemic to the Guiana Shield in Surinam and French Guiana, occurring in montane rain forests (300-400 m elevation range), growing on granitic outcrops.

49. Lesia J. L. Clark & J. F. Smith. (inc. Nematanthus p.p.) Epiphytic or terrestrial subshrub; stems erect, usually branched, to 4 m tall; inflorescences axillary, flowers 1–2 in reduced cymes; corolla posture oblique relative to calyx, yellow. Two spp., L. savannarum (C. V. Morton) J. L. Clark & J. F. Smith, known from E Colombia to E Peru, Amazonas state in Brazil, Guyana, and Surinam, from 270 – 1570 m; and L. tepuiensis G.E. Ferreira & Chautems, known only Mount Aracá in Amazonas state, Brazil.

50. Nautilocalyx Linden ex Hanst. Terrestrial perennial herbs; stem erect, ascending or decumbent, ± succulent; inflorescences axillary, fascicled cymes, or flowers solitary; corolla white to yellow, usually with purple spots or lines, tube laterally broadened, with a short basal sac or spur, limb with nearly equal lobes, entire, their margins toothed or fimbriate; fruit a capsule. 41 spp., 38 in South America, throughout tropical America except SE Brazil (12, 5 endemics); terrestrial herb, growing in shady, wet places and rocks in forests; the white or yellow episcioid corolla suggests pollination by euglossine bees (gynandro-euglossophily).

51. Nematanthus Schrad. (exc. Lesia p.p.) Epiphytic or saxicolous subshrubs or herbs; stem ascendent, climbing, creeping or pendent, lignescent or woody at base, with fibrous roots from the nodes; axillary cymes 1- to 8-flowered; corolla yellow, orange, white. red, rose to dark purple, often resupinate; tube cylindrical or gradually widening, sometimes sharply bent at base, usually strongly ventricose, and making an angle of 20-90° with the pedicel, limb with short spreading or reflexed lobes, mouth constricted, orbicular or pentagonal; fruit a fleshy, white or variously coloured capsule. 32 spp., of montane forests of S & SE Brazil, mainly in Atlantic Forest, less commonly on rocks; a morphologically diverse genus that includes four types of flowers: hypocyrtoid-non-resupinate, hypocyrtoid-resupinate, pendent-resupinate; large campanulate white corollas; pollination apparently by hummingbirds or Euglossine bees; three species from São Paulo and Espírito Santo states are rare plants in Brazil, by Plantas Raras do Brasil’s book.

52. Neomortonia Wiehler. (exc. Pachycaulos) Epiphytic or saxiculous herbs; stems pendent, repent, or scrambling, to 1 m long, slender, 1-2 mm in diam.; flowers solitary in the leaf axils, showy, oblique in the calyx; corolla white, eventually reddish on upper side, tube spurred at base, funnel-shaped, limb with spreading, fringed lobes; or red, strongly pouched, mouth constricted; fruit a berry, orange. Only one sp., N. rosea Wiehler, from Central America to Ecuador, growing on damp, shady rocks or epiphytically on trees in lowland or montane forests.

53. Pachycaulos J. L. Clark & J. F. Smith. (off Neomortonia) Pendent epiphytic herb; stems creeping to decumbent, terete; leaves opposite, subequal; inflorescences axillary, of a single flower; corolla bright red with a yellow limb, fruit a berry, subglobose, laterally compressed. Only one sp., P. nummularium (Hanst.) J. L. Clark & J. F. Smith, known from Mexico to northern Peru (absent in Colombia, however), from 630-2,700 m.

54. Pagothyra (Leeuwenb.) J. F. Smith & J. L. Clark. (off Paradrymonia) Climbing vine up to 1 m, adhering to host via numerous adventitious roots along stem; stems attached to host or pendent, terete; inflorescences axillary, racemose, bracteate, of three to many flowers; corolla posture oblique relative to calyx, yellow (white), marked with bright red, brown, or purple-brown spots; fruit a capsule. Only one sp., P. maculata (Hook. f.) J. F. Smith & J. L. Clark, known from forests of French Guiana, Guyana, and Venezuela from near sea level to 500 m; it probably occurs in Surinam, but is not known from there.

55. Paradrymonia Hanst. (exc. Pagothyra) Epiphytes, facultative; stems subwoody, elongate, creeping or ascending with many adventitious roots; leaves usually clustered on a short stem; leaves of isophyllous species rather short–petiolate; inflorescences axillary, reduced pair-flowered cyme and appearing in fascicles; corolla tubular with a broad limb, base with well-developed spur, corolla lobes subequal, margins range from entire to crenate, or with fimbriations (on ventral lobe); fruit a semi-fleshy, bivalved, dehiscent capsule. 11 spp. P. ciliosa (Mart.) Wiehler from Central America to Guianas, Brazil and Bolivia, 8 only from Guianas to Ecuador, one endemic to Peru, and P. buchtienii (Mansf.) Wiehler endemic to Bolivia, distributed in the understory of rainforests, moist rocks or logs in primary and secondary rainforests.

56. Rhoogeton Leeuwenb. Terrestrial stemless plants with a small tuber, tuber with fibrous roots; cymes long pedunculate, several-flowered, condensed; corolla tubular to trumpet-shaped, oblique in the calyx, tube with a dorsal sac or blunt spur at the base, limb with spreading lobes; fruit a bivalved capsule, loculicidally dehiscent, valves opening to 180º. Two spp., endemic to the Guiana Shield of Guyana and Venezuela (1,000-1,100 m elevation range).

57. Trichodrymonia Oerst. Epiphytes, facultative; stems subwoody, usually reduced to basal rosette, with numerous adventitious roots; leaves opposite, larger leaf of anisophyllous plants semierect and extending beyond the shoot apex; inflorescences many-flowered; corolla infundibuliform, trumpet-shaped or salverform, rarely hypocyrtoid; fruit a semi-fleshy capsule, rarely a berry. 29 spp., from S Mexico to Central America, 20 in South America mainly in Colombia and Venezuela, two up to Peru and Ecuador, the Andes and the Amazon rainforest; predominantly in the understory of rainforests, on wet slopes, stream banks, or on moist rocks or logs.

■ SUBTRIBE SPHAERORRHIZINAE ‣ a single genus endemic to Brazil.

58. Sphaeorrhiza Roalson & Boggan. Terrestrial perennial rhizomatous herbs; rhizomes with tuber-like swellings, often breaking apart into propagules; stem erect or decumbent; leaves lamina with 3-7 pairs of veins; flowers axillary, solitary, showy; corolla white, lavender or purple, broadly tubular; fruit a dry rostrate capsule. 4 spp., endemic to savannas of C Brazil (cerrado) in Pará and Mato Grosso to Minas Gerais and Piauí, mainly on center states, seasonally dormant, surviving and propagating by the unique type of rhizome.

The most distinctive feature is that the plants produce ‘lumpy rhizomes’ unlike the scaly rhizomes of most members of Gloxinieae; the rhizomes have tuber-like swellings that easily break apart, with each piece capable of producing a new plant; another characteristic feature are the valvate calyx lobes sealed in bud.

■ SUBTRIBE LIGERIINAE ‣ a single genus.

59. Sinningia Nees. (inc. Paliavana, Vanhouttea) Terrestrial, often saxicolous, rarely epiphytic perennial herbs or subshrubs, suffrutesent, usually with hard-fleshy tubers (to 1m in diam.), rarely stoloniferous, often rupiculous or with xylopodium; stems erect, less frequently decumbent, mostly unbranched, arising from the tuber, rarely stemless, rosulate, and unifoliate; leaves opposite, or in whorls, or congested on top of stem; inflorescences axillary, cymes, racemes or spikes; corolla red, orange, white, greenish-yellowish, blue or purple, tube broadly tubular, cylindrical or obliquely campanulate. 93 spp., from C America to northern Argentina, but highly centered in E & S Brazil (91, 79 endemics, 6 up to Cono Sur, only S. tubiflora (Hook.) Fritsch (Argentina and Uruguay) and S. sulcata (Rusby) Wiehler (highly rare, Bolivia) are absent in Brazil); few species have a more continental distribution; S. schomburgkiana (Kunth & Bouché) Chautems in known only Amazonas, Roraima and Guyana; S. sceptrum (Mart.) Wiehler is highly disjuntc of mountains of SE Brazil and Andes of Ecuador and Peru; S. incarnata (Aubl.) D.L. Denham is the only species occurring naturally north of Panamá: its range extends into Mexico; 17 spp. in Bahia, Minas Gerais, São Paulo, Rio de Janeiro, Espírito Santo and Paraná states are rare plants in Brazil, by Plantas Raras do Brasil’s book.

S. nordestina Chautems, Baracho & J.A.Siqueira is possibly only the unique true annual Gesneriaceae of New World.

S. tuberosa (Mart.) H. E. Moore and Lembocarpus amoena (Suriname, French Guiana, N Brazil) belongs to the morphologically most remakarble spp. of Neotropical Gesneriaceae; both spp. possess a storage orgen in the form of a tuber and produce usually only one leaf per reason; S. tuberosa is endemic to Minas Gerais and Rio de Janeiro states in SE Brazil; has a perennial tuber and produces one (rarely two or more) tiny vegetative shoot(s) at its top. This shoot produces first a few pairs if tiny cataphylls and then a strongly anisophyllous leaf pair, only one of which develops into a large foliar leaf; the flowers are produced on separate short shoots.

S. minima A.O. Araujo & Chautems endemic to SE Pará state, is the smallest Gesneriaceae worldwide, growing to a maximum size of 2.5 cm across, with correspondingly small flowers.

Two epiphytes, S. cooperi (Paxton) Wiehler and S. douglasii (Lindl.) Chautems, makes Sinningia the only genus with epiphytic spp. of Gesneriaceae outside Episceae in New World.

LINEAGE 4 of 8: PLANTAGINACEAE

PLANTAGINACEAE

§ CARNIVOROUS (Brocchnia - Catopsis - Paepalanthus - Drosera - Heliamphora - Philcoxia - Genlisea - Utricularia - Pinguincula)

Genera/spp. 107/1,750–1,780 Distribution cosmopolitan except polar areas (mainly temperate regions). Habit usually bisexual (rarely monoecious, gynomonoecious, dioecious, or gynodioecious), usually perennial, biennial or annual herbs (rarely shrubs or suffrutices). Some genera are aquatic or semi-aquatic. Numerous representatives are xerophytes. At least one sp. of Philcoxia has carnivorous subterranean leaves.

In our circumscription, the family Plantaginaceae is highly heterogeneous, which is understandable considering the diverse evolutionary trends occurring in the family. It includes highly specialized aquatic plants (Hippuris, Callitriche), small anual spp., tall shrubs (Hebe, Aragoa), rainforest herbs (Tetranema), weeds (e.g., Plantago, Veronica, Cymbalaria), and alpine chasmophytic spp. (Penstemon, Veronica, Erinus, Ourisia). Plantaginaceae has a cosmopolitan distribution and include about 100 genera and 2,000 spp. Of these, 45 genera and 370-400 spp. are native in the Neotropics. Nearly half of the genera occurring in the Neotropics are endemic.

Plantaginaceae are very difficult to distinguish from Scrophulariaceae s. str., Gesneriaceae, Stilbaceae, etc., all having largish, monosymmetric flowers. However, two features common in the family, the frequent absence of regular vertical partitions in the heads of the glandular hairs and septicidal capsule dehiscence, are not that common in Lamiales. Many genera of the Plantaginaceae are used as ornamental worldwide, including coral-plant (Russelia), beardtongue (Penstemon), hebe (Hebe) and snapdragon (Antirrhinum).Some spp. of Plantaginaceae have a pantropical distribution, including some weeds, such as Scoparia dulcis L. and Bacopa monnieri (L.) Wettst. Many spp. native to the northern Hemisphere are weeds in the Neotropics. These include spp. of Linaria, Plantago and Veronica.

SYSTEMATICS tribe Chelonoideae (9/295–320, Borneo, North America, Mexico, Central America) and Globularioideae (3/52, Europe, Macaronesia, Mediterranean to Pakistan, N and NE Africa, Arabian Peninsula, Socotra) do not occur in South America.

1. SUBFAMILY GRATIOLOIDEAE (32/c 365) ‣ two tribes, both in South America.

1.1 GRATIOLOIDEA ▸ TRIBE ANGELONIEAE (6/c 75) - all genera occur in South America.

1. Angelonia Bonpl. (inc. Monopera) Herbs or subshrubs, rarely shrubs. 26 spp., from Mexico to Argentina and Caribbean, 24 in South America, mainly in NE Brazil (overall country 20 spp., 15 endemics, two of then in Minas Gerais and Goiás states are rare plants in Brazil, by Plantas Raras do Brasil’s book), with endemic also in Paraguay, Cuba, Bolivia, Argentina and Mexico

2. Basistemon Turcz. Shrubs to small trees. 7 spp., 5 from Venezuela to Argentina in western side of South America, and B. peruvianus Benth. ex B.D. Jacks. and B. silvaticus (Herzog) Baehni & J.F. Macbr. in Bolivia and Brazil both, the former also in Peru.

3. Ildefonsea Gardner. Herb with lilac flower. Only one sp., I. bibracteata Gardner, a rare plant in Brazil, by Plantas Raras do Brasil’s book, known only from remnant forests in the Rio de Janeiro municipality, SE Brazil.

4. Monttea Gay. Three spp., Chile, W Argentina.

5. Melosperma Benth. Two spp. from Chile and western Argentina.

6. Ourisia Comm. ex Juss. Herbs with zygomorphic to subrotate, pentamerous flowers with four didynamous stamens, loculicidal capsules, and numerous small, reticulate, angled seeds. 30-31 spp., 6 from Venezuela to Bolivia, two of then up to Argentina, 10 in extra-tropical Andes from Argentina and Chile, 14-15 in New Zealand and one in Tasmania.

1.2 GRATIOLOIDEA ▸ TRIBE GRATIOLEAE (27/c 315) - 9 clades.

■ CLADE 1 ‣ two genera, both in South America.

7. Darcya B.L.Turner & C.P.Cowan. Four spp. from Costa Rica and Panamá, D. mutisii (Fern. Alonso) B.L. Turner up to NW Colombia, and D. vandellioides (Benth.) Scatigna, endemic to Brazil, in in open and wet, often disturbed areas of the Atlantic Forest from Bahia to Santa Catarina states.

8. Mecardonia Ruiz & Pav. Herbs glabrous to pubescent. 11 spp., all in the Neotropics, U.S.A. to Argentina, 8 in South America; 4 in Brazil, M. pubescens Rossow endemic, a rare plant in Brazil, by Plantas Raras do Brasil’s book. known only in Vacaria municipality in Rio Grande do Sul state.

■ CLADE 2 ‣ two genera, both in South America.

9. Bacopa Aubl. (inc. Conobea) Annual or perennial herbs, stems prostrate to ascending or erect, some spp. are submerged aquatic herbs. 55 spp., pantropical, but nearly all spp. native in the Neotropics, widely distributed; 45 in South America, 31 spp. in Brazil, 8 endemics.

10. Chodaphyton Minod. (off Stemodia) Herbs. Only one sp., C. ericifolium (K.Schum.) Minod, native from C Brazil to C Argentina and Paraguay.

■ CLADE 3 ‣ three genera, endemics to Brazil.

11. Lapaea Scatigna & V.C.Souza. (off Stemodia) Subshrubs or rarely herbs, terrestrial to rupicolous, per-ennial, up to 1 m high, with variable indument on vege-tative parts, usually aromatic. 5 spp. from rocky grasslands (campos rupestres), at elevations 900-2,000 m, growing on edges of relatively humid and shaded fractures of quartzite out-crops, regionally called lapas, in Minas Gerais and Bahia states, Brazil; three spp. are rare plants in Brazil, by Plantas Raras do Brasil’s book.

12. Philcoxia P. Taylor & V.C. Souza. Annual or perennial herbs, glandular-pubescent; stems erect, rounded; subterranean stems and peltate leaves with circinate ptyxis, under or on the soil surface. 7 spp., restricted to sandy habitats of the savannas of C Brazil (cerrado) and dry seasonal scrubland of NE Brazil (caatinga) biomes, including areas of rocky grasslands (campos rupestres) within the boundaries of the Espinhaço Range and highs in Goiás, Bahia, Minas Gerais and Maranhão states; P. bahiensis V.C. Souza & Harley, P. goiasensis P. Taylor and P. minensis V.C. Souza & Giul. are rare plants in Brazil, by Plantas Raras do Brasil’s book.

The adaxial side of their leaves are provided with stalked capitate glands; protease activity and presence of phosphatases as well as digestion of nematodes and nutrient uptake have been detected; the rare case of circinate leaves in these carnivorous plants are remarkable, since the narrow leaves in both Byblis in Byblidaceae and Drosophyllum in Drosophyllaceae have circinate vernation.

Philcoxia is the most recent carnivorous plant genus to have been discovered. In basic strategy, it is a sticky-leaved plant. On the other hand, it has a little trick up its sleeve.If you were lucky enough to encounter this plant in Brazil (and very few people have!), you would no doubt only notice the short (up to about 25 cm tall), branching inflorescences. The small pink flowers are bilaterally symmetric, lack spurs, and have 5 lobes. The plant's leaves are entirely underground--just barely hidden under the sandy soil crust. The leaves emerge from the underground stem, and are placed at the end of long, filamentary petioles. Each leaf is round, to slightly elongated, and covered (on the upper surface) with short-stalked glands; in Philcoxia bahiensis not only are the glands found on parts of the lower leaf surface, but also sessile glands are seen; laboratory and greenhouse work convincingly demonstrates that these leaves attract and kill nematodes. Furthermore, the leaves digest the nematodes and absorb the nutrients. This is--so far--the only carnivorous plant that seems to specialize in nematodes--although nematodes have been observed being killed and eaten by other carnivorous plants. Also, it has been noted that this is the only sticky-leaved carnivorous plant that hunts under the sand.

13. Tetraulacium Turcz. Annual herbs, densely pilose; stems prostrate, quadrangular; bluish or purplish flowers. Only one sp., T. veronicaeforme Turcz., NE Brazil, with one collection in Rio de Janeiro another in Mato Grosso do Sul states.

■ CLADE 4 ‣ outsider Leucospora (2; E North America, Mexico).

14. Geochorda Cham. & Schltdl. (off Stemodia) Herbs. Only one sp., G. glechomoides (Spreng.) Kuntze, native from S Brazil to NE Argentina and Uruguay.

■ CLADE 5 ‣ two genera.

15. Schistophragma Benth. ex Endl. Annual herbs, glabrous. 4 spp. of Mexico, one up to North America, S. mexicanum Benth. ex D. Dietr. up to NW Colombia.

16. Scoparia L. Herbs to subshrubs, mainly glabrous. 11 spp., tropical America, 10 in South America, one sp. a pantropical weed; 7 spp. in Brazil, S. elliptica Cham. endemic, a rare plant in Brazil, by Plantas Raras do Brasil’s book, from Paraná and Santa Catarina states

■ CLADE 6 ‣ outsiders Dopatrium (14; tropical and S Africa, tropical Asia to New Guinea, tropical Australia), Hydrotriche (3; Madagascar), Limnophila (37; tropical and subtropical regions in the Old World).

17. Gratiola L. (inc. Fonkia) Herbs, with axillary flowers. 34 spp., mainly North America (13, all restricted) and Oceania, two in Eurasia, G. oresbia B.L. Rob. in Central America, G. bogotensis Cortés in Venezuela to Ecuador, G. peruviana L. in Brazil, Bolivia and Cono Sur, and G. uliginosa Phil. endemic to Argentina.

■ CLADE 7 ‣ possible outsider Adenosma (27; China, tropical Asia, N Australia).

18. Dizygostemon (Benth.) Radlk. ex Wettst. Annual herbs, densely pilose; stems erect, quadrangular; lilac flowers. Two spp., D. floribundum (Benth.) Radlk. ex Wettst. from dry areas of Piauí, Bahia and Pernambuco, NE Brazil, and D. riparius Scatigna & G.D. Colletta endemic to Maranhão state.

19. Matourea Aubl. 10 spp., one endemic to Colombia, two in N Brazil, Venezuela and Guianas (dubious records in Central America), and remaining 7 endemics in E Brazil, two of then are rare plants in Brazil, by Plantas Raras do Brasil’s book.

■ CLADE 8 ‣ only one genus.

20. Stemodia L. (exc. Lapaea, Darcya p.p.) Annual, glandular-pubescent, much-branched, often aromatic herbs, or subshrubs; leaves opposite or verticillate, occasionally alternate below, simple, subentire to variously toothed; flowers irregular, in lax terminal racemes or solitary-axillary, pedicellate; calyx 5-lobed; lobes narrow, equal or subequal; corolla tubular; tube cylindric; capsule globose, ovoid, sometimes acuminate; seeds many, small, striate. 38 spp. in New World, 25 in South America, mainly in open areas in wet places; pantropical; 14 spp. in Brazil, 5 endemics.

■ CLADE 9 ‣ only one genus.

21. Umbraria Scatigna & V.C.Souza. Herbs terrestrial to rupicolous, perennial, indument on vegetative parts composed of short, capitate trichomes, interspersed with non-glandular trichomes, sometimes glandular-punctate. Two spp. endemic to Brazil, one in rocky grasslands from Minas Gerais, another occurs in the Atlantic Forest from Rio de Janeiro to Bahia states, around 400 m a.s.l.

■ UNPLACED CLADE ‣ unplaced genera outsiders Sophronanthe (1; SE U.S.A.), Trapella (1; temperate to tropical regions in E Asia), Deinostema (2; E Asia), Cheilophyllum (8; Caribbean).

22. Anamaria V.C. Souza. Herbs, aquatic, highly dimorphic reproductive (erect) and vegetative (with floating leaves) branches. Only one sp., A. heterophylla (Giul. & V.C. Souza) V.C. Souza, endemic from the seasonally marshes and lagoons in dry seasonal scrubland of NE Brazil (caatinga).

23. Boelckea Rossow. Annual herb, pubescent; stems erect, weakly quadrangular. Only one sp., B. beckii Rossow, endemic to Bolivia.

2. UNNAMED SUBFAMILY (2/58) ‣ a single tribe.

2.1 TRIBE RUSSELIEAE (2/58) ‣ outsider Tetranema (3; Central America).

24. Russelia Jacq. 45 spp., almost entirely confined to Northem and Central America, 46 in Mexico, 36 endemics, and only R. sarmentosa Jacq. reaches South America where it is known from a single collection in Colombia and higly dubious records in French Guiana.

3. SUBFAMILY ANTIRRHINOIDEAE (26/290–300) ‣ outsiders all from SW Unites states and NW Mexico except Antirrhinum (c 20; Mediterranean, with their largest diversity in Spain), Chaenorhinum (26; Mediterranean, SW Asia), Holzneria (2; SW Asia), Albraunia (3; SW Asia), Schweinfurthia (8; arid and semiarid regions in NE Africa to India), Acanthorrhinum (1; NW Africa), Misopates (9; Europe, Macaronesia, Mediterranean to Ethiopia and NW India), Linaria (95–100; Europe, Mediterranean, N Africa, temperate and subtropical Asia), Asarina (1; southern France, NE Spain), Cymbalaria (10; C Europe, Mediterranean to Iran), Lophospermum (8; Mexico, Guatemala), Rhodochiton (3; Mexico, Central America), Kickxia (24; Europe, Macaronesia, Mediterranean, N and NE Africa, SW and S Central Asia), Nanorrhinum (29; tropical and subtropical regions in the Old World), Anarrhinum (7; Central and southern Europe, Mediterranean, Ethiopia).

25. Galvezia Dombey ex Juss. 5 spp. in Ecuador and Peru, mainly near the coast, two endemic to the Galapagos Islands.

26. Maurandya Ortega. Three spp. from Mexico, M. erubescens (D. Don) A. Gray up to Panamá, scattered in Venezuela, Ecuador and Peru, M. barclayana Lindl. disjunct in Peru, and M. scandens (Cav.) Pers. disjunct in Venezuela.

27. Nuttallanthus D.A. Sutton. Herbs, flowers with nectar spurs. 4 spp., mainly North America, N. canadensis (L.) D.A. Sutton from Canada to Mexico, Guadalupe, Dominican Republic, W & S South America and Uruguay to Venezuela, and N. subandinus (Diels) D.A. Sutton in W South America, Colombia to Ecuador.

4. SUBFAMILY DIGITALIOIDEAE (16/650–665) ‣ five tribes, three in South America, Digitalideae (3/c 28, Europe, Macaronesia, Mediterranean to Central Asia, Morocco) and Hemiphragmateae (1/1, Himalayas in northern India, Nepal, Sikkim, Assam and Bhutan to China (inc. Taiwan), Philippines, Sulawesi) absents.

4.1 DIGITALOIDEAE▸TRIBE PLANTAGINEAE (3/c 275) - all three genera in South America.

28. Aragoa Kunth. Shrubby habit, xeromorphic leaves, and actinomorphic flower with four corolla lobes and four stamens. 23 spp., Colombia (21) and Venezuela (4), endemic to the paramos.

29. Littorela P.J. Bergius. Aquatic herbs, unisexual monoecious flowers, the fruits containing a single anatropous ovule, and by the plants being capable of reproducing vegetatively by means of stolons. Three spp., L. uniflora (L.) Asch. in Europe, including Iceland and the Azores, and it grows along the margins and in shallow water (down to circa 4m below water-line) of freshwater lagoons, lakes and ponds, and also temporarily inundated depressions, L. americana Fernald in E Canada and the NE U.S.A., and grows in muddy, sandy or gravelly shorelines of lakes, ponds and slow-moving rivers; and L. australis Griseb. ex Benth. & Hook. f. distributed in southern Chile, southern Argentina and the Falkland Islands, where it grows on the margins of freshwater lagoons.

30. Plantago L. (inc. Bouguiera). Herbs or rarely subshrubs, sometimes cushions, perennial or annual, anemophilous. c. 270 spp., cosmopolitan, 84 in New World, 61 in South America, 15 in Brazil, 9 endemics, including P. trinitatis Rahn, the only shrubs spp. of this genus in Brazil, found on Trinidade island, 1,500 km east of Rio de Janeiro in the Atlantic Ocean; although some species have wide distributions, and are sometimes ruderal plants, many others have restricted distributions and occur in more specialized environments, such as those endemics to small oceanic islands. Four subgenera:

§ subg. Bouguiera ‣ only one sp., P. nubicola (Decne) Rahn. restricted from mountains from Peru to Argentina and Bolivia.

§ subg. Coronopus ‣ 11 spp., mainly in Mediterranean region, P. maritima L. also found in other parts of Europe, central Asia, North America and southern South America.

§ subg. Plantago ‣ c. 131 spp., all continents and found on many oceanic islands, often in mesic or moist habitats.

§ subg. Psyllium ‣ 67 spp., Asia, Europe, Africa, North and South America (they are absent from New Guinea, Australia and New Zealand).

4.2 DIGITALOIDEAE ▸ TRIBE VERONICEAE (7/445–455) - outsiders Kashmiria (1; Himalayas), Lagotis (c 20; E Europe, the Caucasus, northern and C Asia, Himalayas to W China), Picrorhiza (3; Himalayas), Scrofella (1; NW China), Wulfenia (4; SE Europe, Türkiye), Wulfeniopsis (1; Himalayas in E Afghanistan, Pakistan, northern India and Nepal).

31. Veronica L. (inc. Hebe). 410–420 spp., temperate and alpine regions on both hemispheres, tropical African mountains, New Zealand and surrounding islands, Tierra del Fuego, W Patagonia south latitude, the Falkland Islands; 4 spp. in South America, V. elliptica G. Forst. and V. salicifolia G. Forst. in New Zealand extending to Argentina and Chile, V. peregrina L. from Alaska and Yukon to Chile, Argentina and S Brazil, and V. serpyllifolia L. from North America to northern Andes.

4.3 DIGITALOIDEAE▸ TRIBE SIBTHORPIEAE (2/6) ‣ outsider Ellisiophyllum (1; India, SE and E Asia to Japan and E to Philippines, Taiwan (China) and New Guinea).

32. Sibthorpia L. Perennial, small, creeping herbs; stems rooting at nodes; leaves alternate, petiolate, suborbicular-reniform, crenate to incised; flowers axillary, 4- to 8-merous, solitary or fasciculate, pedicellate; pedicels ebracteate; capsule somewhatcompressed, loculicidal; seeds few, oblong-ovoid, reticulate or smooth. 5 spp., three in Old World, two in New World, S. conspicua Diels from Bolivia and Argentina and S. repens (L.) Kuntze from Mexico to Argentina.

5. SUBFAMILY CALLITRICHOIDEAE (2/20–50) ‣ both genera in South America.

33. Callitriche L. Terrestrial or aquatic, mainly rosettes; Callitriche is the only genus of angiosperms in which both aerial pollination and hypohydrophily have been documented. 60-65 spp., cosmopolitan, 34 in New World, 21 native in over tropical and temperate South America scattered across a wide range of habitats, 6 in Brazil, none endemics; 3 spp. are endemic to Colombia.

34. Hippuris L. 4 spp., three in North America up to Eurasia, and H. vulgaris L. in North America, Argentina, Chile and Australia.

LINEAGE 5 of 8: SCROPHULARIACEAE

SCROPHULARIACEAE

§ FULLY CHLOROTROPHIC

Genera/spp. 58/c 1,560 Distribution mainly subtropical and warm-temperate regions on the Northern and Southern Hemispheres, with their largest diversity in South Africa and Australia; some representatives in tropical America, tropical E Africa, Madagascar, tropical Asia, New Guinea, and tropical Australia.Habit usually bisexual (rarely functionally dioecious), perennial or annual herbs, evergreen or deciduous shrubs (sometimes suffrutices or biennial herbs, rarely trees or lianas). Many spp. are xerophytes.

SYSTEMATICS In the past it was treated as including about 275 genera and over 5,000 sp., but its circumscription has been radically altered since numerous molecular phylogenies have shown the traditional broad circumscription to be grossly polyphyletic. Many genera have recently been transferred to other families within the Lamiales, notably Plantaginaceae and Orobanchaceae but also several new families.

8 clades; the clades Colpias (1/1, Northern Cape), Teedieae (6/18–19, tropical and southern Africa, Madagascar), Camptoloma (1/3, Gran Canaria, Somalia, southern Yemen, Socotra, Kuria Muria Islands, W deserts in Angola, northern Namibia), Phygelius (1/2, southern Africa) do not occur in South America.

1. UNNAMED SUBFAMILY (5/c. 160) ‣ a single tribe.

1.1 TRIBE HEMIMERIDEAE (5/c 160) ‣ outsiders Diascia (c 70; southern Africa), Nemesia (60–65; tropical and southern Africa, especially W Cape), Hemimeris (7; N, W and E Cape), Diclis (10; tropical and southern Africa, Madagascar).

1. Alonsoa Ruiz & Pavon. Perennial herbs, glabrous except for glandular pubescent inflorescence. 15 spp., two from W and E Cape, 10 in South America from Venezuela to Bolivia (centered in Andes of Peru), one up to Cono Sur, another into Central America and Mexico.

2. SUBFAMILY MYOPOROIDEAE (10/c 300) - three tribes, Aptosimeae (3/41, Africa, Arabian Peninsula to Malesia) and Myoporeae (4/c 240, Mauritius, E and SE Asia, E Malesia to New Guinea, Australia, New Zealand to the Hawaii, Caribbean) are absent in South America; among Leucophylleae, outsiders are Eremogeton (1; S Mexico, Guatemala) and Leucophyllum (12; SW U.S.A., Mexico).

2. Capraria L. Annual or perennial herbs or shrubs; stems erect, much branched, terete. 4 spp., two from U.S.A. to Central America, C. biflora L. from SE U.S.A. (Florida), Caribbean Islands, Mexico, Central America, Galapagos Islands and South America east of the Andes and Brazil, and C. peruviana Benth., in South America on the western side of the Andes (Peru and Ecuador) and the Galapagos Islands.

3. UNNAMED SUBFAMILY (1/c. 110) ‣ a single tribe.

3.1 TRIBE BUDDLEJEAE (1/c 110) ‣ a sinlge genus, Buddleja (101).

3. Buddleja L. Shrubs or small trees; interpetiolar stipule-like lobes (rarely foliaceous) sometimes present at base; tetramerous flowers arranged in axillary cymes, with sessile and epipetalous stamens, often actinomorphic; corolla sometimes quadrilobate; stamens sometimes four; fruit sometimes a drupe. 101 spp., in seven sections, 5 only Africa and adjacent Arabia (joined 10), sect. Alternifoliae from Asia (25), and the sect. Buddleja restricted of New World, with 66 spp., 48 in South America, 15 in Brazil, 9 endemics (2 from SE Brazil states are rare plants in Brazil, by Plantas Raras do Brasil’s book).

4. SUBFAMILY SCROPHULARIOIDEAE (31/c. 1,220) ‣ a single tribe and two subtribes, Scrophularieae (6/c 570, Europe, Africa, Macaronesia, Mediterranean, temperate Asia, Himalayas and Tibet, North and tropical America) outside South America; among Limoselleae, outsiders are 22 genera confined to tropical and southern Africa, Barthlottia (1; Madagascar), and Manulea (74; southern Africa, India, with their highest diversity in W Cape).

4. Limosella L. Small, aquatic to semi-aquatic herbs. 5-16 spp., 4 spp. in the New World, all in South America, three scattered in continent with outliers in Caribbean, Mexico and North America, L. australis R.Br up to S Brazil, and L. acaulis Sessé & Moc. endemic to Venezuela.

LINEAGE 6 of 8: LINDERNIACEAE

LINDERNIACEAE

§ FULLY CHLOROTROPHIC

Genera/spp. 21/252 Distribution tropical to warm-temperate regions in the Northern and Southern Hemispheres, with their largest diversity in tropical Africa and SE Asia. Habit bisexual, annual or perennial herbs (sometimes lignified at base, rarely shrubs). Lindernia intrepida (Dinter ex Heil) Oberm. (= Chamaegigas intrepidus Dinter) from Namibia is poikilohydric aquatic resurrection plant. Cosmopolitan, with three genera in Neotropics, from N Mexico to Chile, Argentina, and Paraguay.

SYSTEMATIC outsiders Artanema (4; tropical Africa, SE Asia, Malesia), Bonnaya (12; one in Kenya, Tanzania, 11 from India to Taiwan in China, Australia and Polynesia), Chamaegigas (1; Namibia), Craterostigma (25; tropical and southern Africa, Madagascar, Yemen, Socotra, India to Vietnam), Crepidorhopalon (30; tropical and southern Africa, Madagascar), Hartliella (4; Congo, Zambia), Hemiarrhena (1; W Australia), Legazpia (1; E Asia to islands in W Pacific), Linderniella (16, tropical and southern Africa, Madagascar), Pierranthus (1; Indochina), Schizotorenia (2; SE Asia), Scolophyllum (2; SE Asia), Stemodiopsis (6; tropical Africa), Torenia (51; tropical and S Africa, Indian Ocean islands, SE Asia), Vandellia (52, Africa to Australia, one in Lesser Antilles).

1. Ameroglossum Eb. Fisch., S. Vogel & A.V. Lopes. (inc. Catimbaua, Isabelcristinia) Perennial shrub up to 2m tall or saxicolous small shrub, stems erect or pendent, occasionally prostrate, quadrangular, woody at base, densely pubescent; inflorescence thyrsic, terminal, frondose, lax, with axillary cymes, bracteoles 1.5-2 x 0.8 mm; flowers pedicellate, pedicel up to 5 mm long, bicarinate above; corolla scarlet except the yellow lower lip, cylindric, up to 41 mm long. 11 spp., all very narrow endemic to dry rockys from Rio Grande do Norte to Alagoas state, NE Brazil, in stands mixed with other arbustive species.

Ameroglossum shares with Russelia, Hemichaena and Eremogeton several characters such as the thyrsic inflorescence and the shrubby habit; it differs, however, in the long staminode, which is minute or absent in Russelia or completely absent in the other genera mentioned; a very characteristic autapomorphy of Ameroglossum is the lower lip of corolla, which looks undivided and tongue-like, but really is minutely 3-lobed at apex; in Russelia, Hemichaena and Eremogeton, the lower lip is conspicuously 3-lobed. The flowers of Russelia also differ by their much less pronounced zygomorphy, and those of Eremogeton by four stamens.

2. Cubitanthus Barringer. Perennial herb, with indumentum of long multicellular hairs; stem decumbent, 4-winged, with adventitious roots at the nodes; roots fibrous; leaves opposite; petiole slightly winged; flowers solitary in leaf axils; sepals free, lanceolate-acute; corolla cylindrical, limb bilabiate, upper lip subentire, not reflexed, lower lip 3-lobed, densely villous at base; capsule septicidal. Only one sp., C. alatus (Cham & Schltdl) Barringer, very narrow endemic, found as a weed in cacao plantations, but is known only from the municipalities of Ilheus and Itajuipe in southern Bahia state, Brazil.

3. Lindernia All. Annual or perennial herbs, shrubs of dwaft shrubs, glabrous to pubescent. 30 spp., 23 only Old World, and 7 in New World (two only Caribbean, one only SE U.S.A., one endemic to Mexico), only three in South America: L. brachyphylla Pennell ex Steyerm. in Venezuela, Guyana, Brazil, L. dubia (L.) Pennell and L. rotundifolia (L.) Alston over New World (both in Brazil).

4. Micranthemum Michx. Herbs (sometimes fewer 5 cm), glabrous; axillary flowers. 14 spp., all New World, 8 endemics to Cuba, two only Mexico, one only North America, one from Florida and Cuba, M. pilosum Ernst endemic to Venezuela and the widely distributed M. umbrosum (Walter ex J.F.Gmel.) S.F.Blake over New World, inc. Brazil, mainly in wet places.

LINEAGE 7 of 8: LAMIOIDS

LAMIACEAE

§ FULLY CHLOROTROPHIC

Genera/spp. 232/6,900-7,200 Distribution cosmopolitan except polar areas. Habit usually bisexual (sometimes gynomonoecious or gynodioecious, rarely polygamomonoecious or dioecious), usually perennial, biennial or annual herbs (sometimes evergreen trees or shrubs, rarely lianas). Many spp. are xerophytic. Young stems and branches usually quadrangular in cross-section. Usually aromatic.

The South American Labiate flora, which contains two very large genera, Salvia and Hyptis, but rather few genera in total, falls naturally into three regions: A. Andean; B. Guianan and Brazilian Shields; C Temperate South America.

ANDEAN LABIATAE are primarily derived from north temperate genera, and the various groups have many links to Central America, including Mexico. The largest group is Salvia subg. Calosphace with its Andean centre of diversity linked to a much larger one in Mexico. Scutellaria, Stachys and Clinopodium also appear to have been derived from a southern migration from Central America, but have radiated to produce endemic groups, in some cases associated with adaptations to humming-bird pollination; other Andean Nepetoid genera include the monotypic Obtegomeria from Colombia, related to Clinopodium, Minthostachys, which appears most closely related to the Macaronesian Bystropogon, and Lepechinia, which extends to Mexico with one in Hawaii; the small Neotropical genus Catoferia (Nepetoideae, Ocimeae) also links Central and southern America, but its closest relatives would seem to be eastern Asiatic members of the Old World subtribe Ociminae, perhaps the genus Orthosiphon Benth. & M.Ashby. The only other member of the subtribe native to the Americas is Ocimum, which extends from Mexico to South America. One sp. extends into lowland areas of the Andes.

GUIANAN AND BRAZILIAN SHIELDS this region, including Amazonian and eastern South America, mostly overlying much older geological formations, has a very different flora, dominated by the Nepetoid Ocimeae, subtribe Hyptidinae; this is composed of the large genus Hyptis and a number of smaller endemics, satellite genera, such as Rhaphiodon, Hypenia, Hyptidendron and Eriope. Brazil is the centre of diversity of 21 genera, several endemic and others centered in Brazil with few outliers outside Brazilian borders; the primarily Old World genus Ocimum (Nepetoideae) is represented by several spp. endemic to the area. It seems probable that Ocimum originally had an Old World tropical origin, perhaps reaching the New World via W Africa in the early Tertiary. A few spp. of Salvia (Mentheae) reach the mountains of E Brazil and extend south into temperate southern America. Other endemic genera to be found in this region include Cornutia (Viticoideae), Amasonia and Monochilus (Ajugoideae), and more widely distributed genera such as Vitex, Volkameria (Viticoideae) and Aegiphila (Ajugoideae); the latter, with over 100 spp., extends throughout tropical America.

TEMPERATE SOUTH AMERICA a number of endemic genera occur in this region, belonging to Nepetoideae, tribe Mentheae: Kurzamra, Rhabdocaulon, Glechon and Hoehnea, while Cunila and Hesperozygis are trans-equatorial, being also represented in Mexico, although this needs confirmation from molecular studies.

Key differences from similar families Separate from Verbenaceae because of cymose inflorescence, usually lipped (rather than salverform) corolla, stamens well-exserted.

Differs from Rubiaceae as fused interpetiolar stipules absent, usually zygomorphic flowers, superior (rather than inferior) ovary.

Differs from Solanaceae as usually opposite leaves, fruits with 4 nutlets or 1-5 pyrenes (not copious seed).

Separated from Apocynaceae by absence of white sap, petals not twisted in bud, no colleters on petiole base and adjoining stem, seed usually without tuft of hairs; androecium and gynoecium never fused.

Separated from Boraginaceae as flowers not regular, cymes usually not scorpioid, often not hispid.

Separate from Rutaceae (compound leaves Vitex can be confused with Rutaceae) as flowers not regular corolla and calyx lobes fused (not free), leaves usually opposite (not alternate).

Use Ornamental plants, spices, perfumes (Lavandula, Marrubium, Mentha, Pogostemon, Rosmarinus, Salvia), medicinal plants, honey (Phacelia etc.), timber (Viticoideae, Ajugoideae, Tectona grandis, etc.), seed-oils (Perilla etc.).

SYSTEMATIC Verbenaceae is now restricted to subfamily Verbenoideae of traditional classifications which has an indeterminate racemose inflorescence and a salverform corolla with stamens included; whereas in the Lamiaceae the inflorescence is cymose with determinate, usually opposite cymes and the corollas are tubular and usually bilabiate with the stamens usually exserted from the tube, but can be held within the lobes and rarely within the tube.

The cymes in Lamiaceae are arranged usually in opposite pairs along an indeterminate axis, forming a thyrse. In some Lamiaceae the cymes are reduced to single flowers though bracteoles are often present below the flower in these cases, indicating the cymose, rather than racemose, nature of the inflorescence. These gross morphological differences are supported by anatomical and pollen characters: the Verbenaceae have their ovules attached marginally on the carpel margin, and have thickened pollen exine near the apertures; the Lamiaceae have the ovules attached submarginally and have an unthickened exine.

A probable topology is (Li & al. 2016) [[Callicarpa + Prostantheroideae] + [[Symphorematoideae+Viticoideae] + Nepetoideae + [Tectona + [Premnoideae + Ajugoideae + [Peronematoideae + [Scutellarioideae + [Cymarioideae + Lamioideae]]]]]]]].

12 lineages, Prostantheroideae (14/310–320, Australia), Symphorematoideae (3/c 30, India to W Malesia), Tectona clade (1/3, tropical Asia to southern China and Philippines), Peronematoideae (4/17, southern Burma and Thailand, the Malay Peninsula, Malesia to New Guinea, Bismarck Archipelago, Solomon Islands and NE Queensland) and Cymarioideae (2/3, SE Asia) lineages do not occur in South America.

1. CALLICARPA CLADE (1/c 140) ‣ a single genus.

1. Callicarpa L. Small trees or shrubs, rarely lianas. 140 spp., temperate and tropical Asia and America (North and Central America, Venezuela to Bolivia), W. Indian Ocean to W. Pacific; 31 spp. in New World, only C. acuminata Kunth in South America, up to Venezuela and Bolivia.

2. CLADE VITICOIDEAE (4/c 275) ‣ outsiders Petitia (2; Caribbean), Pseudocarpidium (9; Caribbean, with their largest diversity on Cuba), Teijsmanniodendron (23; SE Asia, Nicobar Islands, Malesia to New Guinea, Bismarck Archipelago and Solomon Islands).

2. Vitex L. Trees or shrubs, leaves usually digitately compound with 3-8 leaflets; fruit a drupe, likened to an egg in egg-cup. 250 spp. in Old and New World (55 spp., over countries, except Canada and Chile), few in temperate regions; 41 in South America, 34 in Brazil, 19 endemics, 5 in Amazonas, Para, Maranhão and Bahia states are rare plants in Brazil, by Plantas Raras do Brasil’s book.

3. SUBFAMILY NEPETOIDEAE (105/3.540–3.840) ‣ three tribes, Elsholtzieae (6/c. 70, Caucasus, E India and Himalayas to Burma, China, Korean Peninsula Japan, the Ryukyu Islands, the Kuril Islands and Russian Far East, SE Asia, Malesia to Philippines) absent in South America.

3.1 NEPETOIDEAE ▸ TRIBE MENTHEAE (56/1.940–2.440) - outsiders all only Old World or North America to Mexico except Lycopus (19; Europe, NW Asia, temperate North America, one species, L. australis, in SE South Australia to SE Queensland, Tasmania), Mentha (18–19; nearly cosmopolitan except South America), Satureja (c 45 or 200–210; nearly cosmopolitan, with their highest diversity in temperate regions in the Old World), Piloblephis (1; Georgia, Florida, the Bahamas).

3. Clinopodium L. Herb and shrubs, perennial, rarely annual, aromatics. 100 spp., 70 in the New World, and temperate Eurasia, but a few in Africa, tropical Asia and Indomalesia; 40 spp. in South America, highly centered in Ecuador/Peru (a half of continent restricted of this zone), almost over countries, only the widely distributed C. brownei (Sw.) Kuntze reaches into S Brazil; Clinopodium giliesii (Benth.) Kuntze, endemic to Meditteranean places of Chile, is the one from two desiccation tolerant species dicots in New World, joined Blossfeldia Werdermann from Cactaceae.

4. Cumiria Colla. Small tree or shrub, aromatic. Only one sp., C. eriantha Benth, in lower montane forest, Massatierra Island, Juan Fernandes Archipelago, Chile.

5. Cunila Royen ex L. Shrubs, subshrubs, and rhizomatous perennial herbs, strongly aromatic, with simple hairs. 19 spp., 8 in C & E U.S.A. to Panamá, 11 in Brazil, 4 endemics and 7 up adjacent Argentina, Uruguay and Paraguay.

6. Eriopidion Harley Hooker. Eriope-like, diifering with its herbaceous habit, in lacking a stylopodium and a calyx with hygroscopic lobes and triquetrous nutlets. Only one sp., E. strictum (Benth.) Harley a rare plant of dry, sandy areas within the dry seasonal scrubland of NE Brazil (caatinga), reappearing in semi-arid vegetation by the Lower Orinoco River, in Venezuela.

7. Eriothymus (Benth.) Rchb., Handb. Shrubs. Only one sp., E. rubiaceus (Benth.) J.A.Schmidt, known only the type collection in a mounatins area of Minas Gerais state, Brazil, and possibly extinct.

8. Glechon Spreng. Small shrubs or subshrubs, sometimes with xylopodium. 7 spp., all from Brazil, 4 reaching to Argentina, Paraguay and Uruguay, open, damp areas.

9. Hedeoma Pers. Herbs, sometimes cushions, strongly aromatic. 43 spp., disjunct distribution: 37 in Canada and S. U.S.A. to Guatemala, and 6 spp. from Peru southwards, incl. Brazil (3, two endemics).

10. Hesperozygis Epling. Shrubs or subshrubs, with simple or rarely branched hairs. 7 spp., endemic to SE & S Brazil.

11. Hoehnea Epling. Perennial herbs, flowers deep pink to lilac or white. 4 spp. all from Brazil, only H. epilobioides (Epling) Epling reaching to Argentina and Paraguay, marshy ground.

12. Kurzamra Kuntze. Mat-forming perennial herb, dwaf, aromatic. Only one sp., K. pulchella (Clos) Kuntze, Chile (Atacama and Coquimbo) and adjacent Argentina.

13. Lepechinia Willd. (inc. Sphecele) Herbs and shrubs, perennial. 48 spp., mainly montane, Andes from Argentina and Chile northwards to Mexico and California, including Venezuela (33 in South America), with outlying spp. in Brazil (2, L. annae (Taub. ex Schwacke) Bradea from Mantiqueira Range and L. speciosa (A. St.-Hil. ex Benth.) Epling from Caparao Range, both endemic).

14. Minthostachys (Benth.) Spach. Weak-stemmed, often semi-scadent shrubs aromatic, leaves shallowly toothed to subentire. 19 spp., montane Andes from Venezuela to Argentina (absent in Chile).

15. Obtegomeria Doroszenko & P.D.Cantino. Ericoid low shrubs with simple hairs. Only one sp., O. caerulescens (Benth.) Doroszenko & P.D. Cantino, endemic to Sierra Nevada de Santa Marta, Colombia.

16. Rhabdocaulon (Benth.) Epling. Shrubs and perennial herbs, stems erects or virgate. 8 spp., primarily Brazil (6 endemics), R. stenodontum (Briq.) Epling reaching to Paraguay, R. strictum (Benth.) Epling up to Argentina, Uruguay and Paraguay; savannha and open areas.

17. Salvia L. Annual or perennial herbs or shrubs, often aromatic, recognizable by its unusual stamen structure. c. 900 spp., Old World and the Americas, with three distinct regions of diversity: 648 in New World (237 in South America); Central Asia and Mediterranean (250); Eastern Asia (90); Mexico has 333 spp. in this genus, and it’s the largest in country, and the country with the largest diversity in this genus; 62 spp. in Brazil, 47 endemics.

3.2 NEPETOIDEAE ▸ TRIBE OCIMEAE (44/1,400-1,600) - outsiders Lavandula (45–50; Macaronesia, Mediterranean, North Africa to Somalia, SW Asia to SE India); Isodon (c 105; tropical and subtropical regions in Africa and Asia), Siphocranion (2; China, Tibet, Burma, SE Asia), Hanceola (8; China), Asterohyptis (4; Mexico to Costa Rica), Syncolostemon (c 45; southern and SE Africa), Heterolamium (2; China), Fuerstia (9; tropical E and S Africa), Benguellia (1; Angola), Endostemon (20; tropical and southern Africa and Madagascar to Arabian Peninsula and India), Haumaniastrum (c 35; tropical Africa, one species, H. villosum, on Madagascar), Platostoma (c 45; tropical Africa, Madagascar, tropical Asia to New Guinea), Dauphinea (1; Madagascar), Capitanopsis (3; Madagascar), Madlabium (1; Madagascar), Plectranthus (c 350; Africa, Madagascar, tropical Asia to tropical Australia and islands in the Pacific), Thorncroftia (6; northern Province, Mpumalanga), Tetradenia (c 20; tropical and southern Africa, Madagascar), Anisochilus (17; tropical Asia from India and Sri Lanka to Indochina), Leocus (5; tropical Africa), Aeollanthus (40–45; tropical and subtropical Africa), Alvesia (3; Central Africa), Pycnostachys (c 35; tropical and southern Africa, Madagascar).

18. Cantinoa Harley & J.F.B.Pastore (inc. Oocephalus p.p.). 26 spp., two widely distributed, two from Mexico to northern Andes and 22 centered in Brazil - 4 up to neighbouring countries (Peru, Bolivia, Paraguay), and 18 endemics; two spp. have been introduced into the Palaeotropics: C. americana (syn. Hyptis spicigera Lam.) and C. mutabilis (syn. H. mutabilis (Rich.) Briq.).

19. Catoferia (Benth.) Benth. in G.Bentham & J.D.Hooker. Shrubs or herbs, woody at base or sometimes geoxylic. 4 spp., three from Mexico to Panamá, and C. spicata (Benth.) Benth. from Colombia and Peru.

20. Condea Adans. Shrub, subshrub or herbs with an often spiciform thyrse of pedunculate or sessile cymes in which the intercalary cyme-axes are contracted; the flowers are usually small and the gynoecium without a stylopodium. 27 spp., ranging from W U.S.A. and Central America to the Caribbean and South America (8, except Chile and Guianas); three spp. in Brazil and adjacent Argentina, Paraguay and Uruguay, C. thyrsiflora (Epling) Harley & J.F.B. Pastore endemic to Brazil, C. jacobi (Fern. Alonso) Harley & J.F.B. Pastore in NE Colombia, C. tafallae (Benth.) Harley & J.F.B. Pastore in Bolivia and Peru, C. verticillata (Jacq.) Harley & J.F.B. Pastore from U.S.A. to Ecuador and Venezuela.

21. Cyanocephalus (Pohl ex Benth.) Harley & J.F.B.Pastore. Hyptis-like, but with globose (even when immature), pedunculate capitula with narrowly linear involucrate bracteoles, inconspicuous at anthesis, tube of fruiting calyx usually strongly curved with oblique mouth. 27 spp., 22 mainly in savannas of C Brazil (cerrado), four extending to Paraguay and Bolivia, and C. pedalipes (Griseb.) Harley & J.F.B. Pastore endemic to Cuba. 8 spp. from several states are rare plants in Brazil, by Plantas Raras do Brasil’s book.

22. Eplingiella Harley & J.F.B.Pastore. Shrubby habit withsmall xeromorphic leaves, the flowers in 15−18-fld cymes, forming pedunculate, spherical clusters of very shortly pedicellate flowers, with narrowly linear bracteoles, not forming an involucres. Three spp., of semi-arid, sandy areas, in upland areas of NE Brazil extending at dry seasonal scrubland of NE Brazil (caatinga) and tabuleiros and in the Atlantic sandy coastal shrublands (restingas), in the states of Rio Grande do Norte, Ceará, Paraíba, Pernambuco, Sergipe and Bahia (E. cuniloides (Epling) Harley & J.F.B. Pastore endemic to Bahia is a rare plant in Brazil, by Plantas Raras do Brasil’s book).

23. Eriope Humb. & Bonpl. ex Benth. (exc. Hypenia p.p.) Trees, shrubs, subshrubs or herbs, sometimes with xylopodium. 32 spp., all in Brazil, 30 endemics, E. macrostachya Mart. ex Benth. and E. crassipes Benth. extending into Colombia, Venezuela, French Guiana, and Paraguay, mostly in savannas and mountains; 9 spp. of several states are rare plants in Brazil, by Plantas Raras do Brasil’s book.

24. Gymneia (Benth.) Harley & J.F.B.Pastore. Herbs; inflorescences forming elongate congested or interrupted spikes, composed of globose verticillasters in the axils of reduced bracts; fruiting calyx with tube strongly curved and with oblique mouth. 7 spp., all in Brazil (4 endemics), occurring in the savannas of C Brazil (cerrado), three extending to Eastern Bolivia and G. virgata (Benth.) Harley & J.F.B. Pastore also in Paraguay, with one sp. typical of waste places in dry seasonal scrubland of NE Brazil (caatinga).

25. Hypenia (Mart. ex Benth.) (inc. Eriope p.p.) Harley. Shrubs or subshrubs, often aromatic. 25 spp., H. violacea Mart. Gord. & S. Valencia is endemic to Mexico, 24 spp. in Brazil, 20 endemics (6, mainly in Goiás state, are rare plants in Brazil, by Plantas Raras do Brasil’s book); H. salzmannii (Benth.) Harley, a somewhat weedy spp. of NE Brazil in upland dry seasonal scrubland of NE Brazil (caatinga) and coastal sands, often behaves as an annual, in drier areas, extending rarely into the foothills of some of the tepuis of the Eastern Guiana Shield; H. glauca (A. St.-Hil. ex Benth.) Harley, H. macrosiphon (Briq.) Harley and H. reticulata (Mart. ex Benth.) Harley reaches from Brazil to Bolivia and Paraguay.

26. Hyptidendron Harley. Trees, shrubs or subshrubs, often aromatic. 22 sp., from E Brazil to Mato Grosso and Rondônia, three extending into E Bolivia, and H. arboreum (Benth.) Harley, of forest margins, has a disjunct distribution in the Guiana Shield, in Colombia and on the Peru/Bolivia border; three spp. from center states are rare plants in Brazil, by Plantas Raras do Brasil’s book.

27. Hyptis Jacq. (inc. Peltodon, Oocephalus p.p.). Herbs to shrubs or small trees, very rarely lianoids, some members (in center Brazil) are Cupressus-like, sometimes with xylopodium. 172 spp., almost entirely New World, from U.S.A. to Caribbean and southern to Argentina and Peru, only H. atrorubens Poit extending into Old World (Africa, from Liberia to Gabon, also in New World); 157 spp. in South America, 125 in Brazil, 87 endemics, 22 are rare plants in Brazil, by Plantas Raras do Brasil’s book.

28. Leptohyptis Harley & J.F.B.Pastore. Herbs; flowers arranged in few-flowered compact cymes in the axils of inconspicuous bracts, forming a very slender ± interrupted spike, the flowers with an actinomorphic calyx with subulate lobes, bearing a scarious, deltate flange in each sinus between them, the corolla tubular. 5 spp., restricted to the mountains of NE Brazil, in Bahia and Pernambuco; L. pinheiroi (Harley) Harley & J.F.B.Pastore from Bahia state is a rare plant in Brazil, by Plantas Raras do Brasil’s book.

29. Marsypianthes Mart. ex Benth. Shrubs or perennial herbs. 7 spp., 6 in savannas and coastal dunes, primarily in savannas of C Brazil (cerrado), 4 of them endemics; M. hassleri Briq. up to Paraguay and Argentina; the weed M. chamaedrys (Vahl) Kuntze from Mexico southwards and widely distributed in the Neotropics; and M. arenosa Brandegee endemic to Mexico.

30. Martianthus Harley & J.F.B.Pastore. Cyanocephalus-like, but with flowers in compact pedunculate, spherical capitula with an involucre of linear, membranous bracteoles, the calyx tube is often curved downwards in fruit and the gynoecium without a stylopodium. 4 spp., three restricted to dry seasonal scrubland of NE Brazil (caatinga), inc. dry montane, with an outlying sp., M. elongatus (Benth.) Harley & J.F.B.Pastore, in similar semi-arid conditions in Huarochiri, coastal Peru; M. sancti-gabrielii (Harley) Harley & J.F.B. Pastore ex Benth. from Bahia state is a rare plant in Brazil, by Plantas Raras do Brasil’s book.

31. Medusantha Harley & J.F.B.Pastore. Shrubs, sometimes with xylopodium; 8 spp., 7 endemics to E Brazil and M. eriophylla (Pohl ex Benth.) Harley & J.F.B. Pastore up to E Bolivia; M. carvalhoi (Harley) Harley & J.F.B.Pastore from Bahia state is a rare plant in Brazil, by Plantas Raras do Brasil’s book.

32. Mesosphaerum P. Browne Civ. Herbs to shrubs, inflorescence with several to many flowers in lax or more congested cincinnate cymes, with bracteoles not forming an involucre, the flowers with a calyx often with white trichomes at the throat, and the gynoecium with stylopodium absent. 23 spp., with a primarily Andean distribution (13), especially in Ecuador and Colombia, 4 only in mountain areas of Mexico and Central America, three has a wider distribution, extending also into E Brazil, and two are endemics to Brazil; M. irwinii (Harley) Harley & J.F.B. Pastore from Bahia state is a rare plant in Brazil, by Plantas Raras do Brasil’s book).

33. Ocimum L. Shrubs or herbs, sometimes with xylopodium, usually aromatic. 65 spp., 5 in New World, O. campechianum Mill. from Florida and Mexico to Uruguay and Caribbean, O. carnosum (Spreng.) Link & Otto ex Benth. in Mexico, N. Venezuela, S. Bolivia to Brazil and N. Argentina, two only in Brazil and Cono Sur, and O. transamazonicum E. Pereira endemic to Brazil.

34. Oocephalus (Benth.) Harley & J.F.B.Pastore. (exc. Hyptis p.p., Cantinoa p.p.) Herbs to shrubs, inflorescence of congested pedunculate or sessile cymes, not forming a globose or semi-globose capitulum but near ovoid in form, surrounded by an involucre of often broad, ovate or lanceolate bracteoles; corollas with an elongate tube. 22 spp., all in Brazil, typically in rocky grasslands (campos rupestres) of the Espinhaço Range of Minas Gerais and Bahia and in similar habitats in Goiás; only one also outside Brazil, O. oppositiflorus (Schrank) Harley & J.F.B. Pastore is spreading into disturbed habitats and has a wider distribution, also extending into E Bolivia; 6 spp. are rare plants in Brazil, by Plantas Raras do Brasil’s book.

35. Orthosiphon Benth. Shrubs or herbs, often geoxylic. 40 spp., highly disjunct, in Africa, Madagascar and tropical or subtropical Asia, one in New World, O. americanus Harley & A.J.Paton, endemic to Colombia.

36. Physominthe Harley & J.F.B.Pastore. Shrubs or subshrubs, often aromatic; stems virgate, erect; leaves lobed, sometimes weakly so; inflorescence thyrsoid, lax, formed of pedunculate, subumbellate cymes (with intercalary axes reduced), 3- to 6-flowered, flowers shortly pedicellate from the axils of minute, subulate bracts. Two spp., P. vitifolium (Pohl ex Benth.) Harley & J.F.B.Pastore, restricted to usually montane savannas of C Brazil (cerrado), in areas of Bahia, Minas Gerais, Goiás and São Paulo states, and P. longicaulis Harley endemic to Bahia.

37. Rhaphiodon Schauer. Herb, perennial strongly aromatic; stems prostrate from central tap-root, rooting at nodes; inflorescence a solitary spherical capitulum, many-flowered, axillary; flowers sessile; corolla strongly 2-lipped, 5-lobed (2/3), deep purple; nutlets large, usually 1-2 developing per flower, slightly flattened, glossy, mucilage absent. Only one sp., R. echinus (Nees & Mart.) Schauer, in dry sandy places subject to seasonal flooding, NE Brazil, especially along roadsides; only genus endemic to Brazil composed of only weed species.

4. SUBFAMILY PREMNOIDEAE (3/170–180) ‣ outsiders Premna (130–140; tropical Africa, Madagascar, Indian Ocean islands, tropical Asia to tropical Australia and islands in the Pacific), Gmelina (c 35; Pakistan, India, Sri Lanka, southern China, SE Asia, Malesia to New Guinea, tropical Australia, Solomon Islands, New Caledonia, Fiji).

38. Cornutia Plum. ex L. Trees or shrubs, leaves usually aromatic. 9 spp., of Mexico to Peru, east to French Guiana, Brazil and Caribbean; 5 in South America, C. pubescens Gaertn. only in French Guiana, C. australis Moldenke endemic to Minas Gerais state in SE Brazil (unique in Brazil), two only from Venezuela to Peru, and C. pyramidata L. from Mexico to Bolivia and Guianas.

5. SUBFAMILY AJUGOIDEAE (23/740–770) ‣ outsiders Karomia (9; Africa, Madagascar, Vietnam), Discretitheca (1; Nepal), Rotheca (35–60; tropical Africa, tropical Asia), Glossocarya (10; Sri Lanka, Burma, SE Asia, New Guinea, Queensland); Ajuga (35–40; regions in the Old World to Siberia, Japan, Australia and islands in Pacific), Pseudocaryopteris (3; India, Himalayas, Burma, southern China, SE Asia), Tripora (1; China, Korean Peninsula, Japan), Amethystea (1; C Asia, southern Siberia, Mongolia, China, Koreas, Japan), Trichostema (17–19; southern Canada, U.S.A., Mexico), Caryopteris (7; Central Asia, Tibet, Mongolia, China (inc. Taiwan), Korean Peninsula, Japan); Hosea (1; Borneo), Oxera (24; Borneo to New Guinea, Bismarck Archipelago, Solomon Islands, Queensland, New Caledonia, Vanuatu, Fiji, Samoa, Tonga), Ovieda (1; Hispaniola), Tetraclea (1; Arizona, Texas, N Mexico), Kalaharia (1; tropical and southern Africa), Schnabelia (5; SW and southern China), Rubiteucris (2; India, Himalayas, Tibet, Burma, China (inc. Taiwan)).

39. Aegiphila Jacq. Trees, shrubs or woody vines, frequently dioecious. 138 spp., Florida, Mexico to Argentina, east up to French Guiana, 120 spp. in South America, 33 in Brazil, 15 endemics, 4 of then in Amazonas, Pará and São Paulo states are rare plants in Brazil, by Plantas Raras do Brasil’s book.

40. Amasonia L.f. Shrubs, subshrubs and perennial herbs, sometimes with taproot tubers. 5 showy spp. from French Guiana to Peru, Brazil (all spp., two endemics), Bolivia and Paraguay.

41. Clerodendrum L. Herbs to trees. c. 150 spp., 17 in New World, 7 in South America, scattered, two in Brazil, C. ekmanii Moldenke up to Argentina, and C. rusbyi Moldenke endemic.

42. Volkameria L. Shrubs, sometimes subherbaceous, lianes, rarely small trees; branches ± tetragonal; leaves never spiny; inflorescences axillary to supra-axillary cymes; flowers usually fragrant; calyx campanulate, sometimes pink or purple, lobes unequal; fruits generally globose to obovoid. 25-30 spp., pantropical, with only 1 spp. in Asia, two in New World, V. aculeata L. from Mexico to Brazil and Caribbean, up to Guianas, and V. ligustrina Jacq. from Mexico and Central America.

43. Monochilus Fisch. & C.A.Mey. Perennial herbs or shrubs with racemose inflorescences. Two spp. endemics to Brazil, M. obovatus C. P. Cantino in savannas of Goiás state (cerrado), and M. gloxiniifolius Fischer & C. A. Mayer in Atlantic Forest of Rio de Janeiro and south Espírito Santo states.

44. Teucrium L Shrubs, subshrubs or perennial herbs (rarely annual or biennial), often aromatic; subosmopolitan but mostly extra-tropical, centred in Mediterranean. 250 spp., 11 spp. from southern U.S.A. to Chile, east to French Guiana, Brazil; 4 in South America, T. bicolor Sm. and T. nudicaule Hook. only in Cono Sur, and two widely distributed, T. cubense Jacq. and T. vesicarium Mill., both in Brazil.

6. SUBFAMILY SCUTELLARIOIDEAE (5/c 380) ‣ outsiders Wenchengia (1; Hainan); Holmskioldia (3; Himalayas), Renschia (1; northern Somalia), Tinnea (c 20; tropical and southern Africa)

45. Scutellaria L. Perennial herbs, shrubs and subshrubs, usually non-aromatic. 360 spp., cosmopolitan, poorly represented in tropical lowlands, possibly absent in southern Africa; 142 spp., in New World, over countries; 38 spp. in South America, 8 in Brazil (one endemic).

7. SUBFAMILY LAMIOIDEAE (50/1.295–1.470) ‣ only one genus occurs in Neotropics southwards Mexico; the clades Pogostemoneae (10/c. 145, southern tropical Africa, Madagascar, islands in the Indian Ocean, tropical Asia to Tibet, China and New Guinea, Korean Peninsula, Bismarck Archipelago, tropical Australia), Colquhounia clade (1/5, Himalayas, Tibet, Burma, SW China, Thailand, Indochina), Gomphostemmateae (2/57, Pakistan, NE India, Burma, southern China (inc. Taiwan), Korean Peninsula, Japan, SE Asia, Andaman Islands, the Malay Peninsula, Malesia to the Lesser Sunda Islands), Synandreae (5/19, southern and SW Canada, U.S.A., Mexico), Betonica-Galeopsis clade (2/15, Europe, Mediterranean, North Africa, W and SW Asia to the Caucasus and northern Iran), Paraphlomideae (3/32, China (inc. Taiwan), SE Asia to Central Malesia, Japan, the Ryukyu Islands), Phlomideae (2/210, Mediterranean to SE Russia and the Caucasus to W and C Asia, India, Himalayas, Mongolia, W and C China), Leonureae (6/c. 81, Europe, Russia, Türkiye and Caucasus, Iran, Afghanistan, Central Asia to NW China and Mongolia, Japan), Marrubieae (5/c. 90, Mediterranean, North Africa, Arabian Peninsula, W Asia, NW India, one species, B. africana, in southern Namibia and South Africa), Lamieae (3/50–60, Europe, Mediterranean, North Africa, temperate Asia to Kamchatka, Japan and Taiwan in China) and Leucadeae (6/c. 160, tropical and southern Africa, Ethiopia, southern Arabian Peninsula, Iran, Pakistan, southern and tropical Asia to Japan and Malesia to Queensland) do not occur in South America. Among the solely South American tribe, Stachydeae, outsiders is Melittis (1; Europe, Türkiye).

46. Stachys L. Perennial or annual herbs, or rarely small subshrubs, often stroghly aromatic. 450-600 spp., cosmopolitan, absent in Australia and New Zealand; 96 in continental New World, c. 40 in Mexico, 30 in South America, centered in NW region, 11 in southern region, 18 from Venezuela to Bolivia, only one, S. gilliesii Benth., reaches up to Brazil (also in Ecuador, Peru, Bolivia, Cono Sur).

PHRYMACEAE

§ FULLY CHLOROTROPHIC

Genera/species 15/207. Distribution Americas, Asia, Africa, and Australasia. Habit annual or perennial herbs, secondarily woody; iridoids absent, possibly sometimes present. Use ornamental plants.

SYSTEMATICS outsiders Mimulus (7, North America, Asia to Africa, and Australasia), Elacholoma (2, Australia), Glossostigma (5, mainly Australia and New Zealand, India, Indochina, and apparently southern Africa), Phryma (1, E North America and SE Asia), Hemichaena (5, Mexico and Central America), Mimetanthe (1, North America, Mexico), Diplacus (46, North America, Mexico), Peplidium (4, arid and semi-arid Australia, North Africa, India), Uvedalia (2, Australia, Papua New Guinea, Timor), Microcarpaea (2, China inc. Taiwan, India, Indonesia, Japan, Korea, Malaysia, Thailand, Vietnam, and Australia) and Thyridia (1, Australia, New Zealand).

1. Erythranthe Spach. (off Mimulus) Annual (fibrous-rooted or taprooted) or perennial (rhizomatous), terrestrial or semi-aquatic; stems herbaceous, prostrate to decumbent or erect, terete or 4-angled; flowers apparently solitary or axillary in bracteate, corymboid or racemose groupings. 111 spp., mainly in North America; 12 sections, 8 only in North America/Mexico (29), two from Asia/North America (with 18 spp. in North America and 10 in Asia), and three also in South America:

§ sect. Sinopitheca ‣ 4 spp. in Asia and E. bridgesii (Benth.) G.L. Nesom in Chile and Argentina.

§ sect. Mimulosma ‣ 18 spp., E. stolonifera (Novopokr.) G.L. Nesom from Russia, 17 spp. from U.S.A. and Mexico, E. moschata (Douglas ex Lindl.) G.L. Nesom disjunct in Chile.

§ sect. Simiola ‣ 38 spp. restricted from North America and Mexico, nine in Chile (4 of then reaching into Argentina, 1 into Bolivia) and adjacent southern coast of Peru, and one, E. glabrata (Kunth) G.L. Nesom, in U.S.A., Mexico, and disjunct in Colombia.

2. Leucocarpus D. Don. Shrubs or suffrutescent perennial herbs, erect, up to 2.5 m tal, large and thickened-succulent leaves; flowers in axillary, pedunculate cymes, on short, bracteate pedicels. Corollas yellow or white with a yellow throat, deciduous, limbs bilabiate; fruit a berry. Only one sp., L. perfoliatus (Kunth) Benth. ranging from center Mexico and Central America (Panamá, Nicaragua, Honduras, Guatemala) southward to South America (Venezuela to Bolivia), at elevations of 450-3,100 m.

OROBANCHACEAE

§ PARASITIC (Prosopanche – Cassytha - ... - Mitrastemon – APODANTHACEAE – SANTALALES - Lennoa – OROBANCHACEAE - Cuscuta)

Genera/spp. 99/2,000 Distribution cosmopolitan. Habit bisexual, usually perennial, biennial or annual herbs (sometimes climbers (Velloziella), rarely shrubs or suffrutices, e.g. Brandisia, Cyclocheilon, Asepalum). Nearly all spp. are either root hemiparasites having green assimilating leaves, or more or less succulent achlorophyllous root holoparasites with scale-like leaves. Often blackening when dry. Roots in Asepalum and Cyclocheilon at least usually red. Lindenbergia comprises autotrophic, photosynthesizing plants. This family include some damaging weeds such as the witchweeds (Striga and allies).

21 genera and 300-330 spp. are native in the Neotropics and one (Parentucellia Viv.) is naturalised; throughout the Neotropics, especially the Andes and the highlands of Brazil. Rehmania clade is divergetnt of the remaining Orobanchaceae; Two genera and nine spp.; Rehmannia: China, Korean Peninsula; Triaenophora: China. Lindebergieae subfamily absent in South America.

This family include some damaging weeds such as the witchweeds (Striga asiatica (L.) Kuntze, S. hermonthica (Delile) Benth. and allies) and they present in the agriculture in many countries, particularly in Africa and Asia, affecting very poor areas. The parasitic habit, plus the quick life cycle, and extraordinary capacity for propagation (thousands of small seeds carried by wind can be produced by a single plant), makes the species of Striga some of the worst weeds in the world.

There is no record of Striga infestation in the Neotropics, but some areas in U.S.A. have been infected for decades, with little perspective of eradication. Considering that many places where Striga species occur in Asia and Africa have a climate similar to the Neotropics and also a similarity in crops cultivated, it is realistic to be concerned about these species. Buchnera, which is morphologically similar to Striga and its sister group, has many native Neotropical species, but there is no record of Buchnera species behaving as weeds; many genera have big and attractive flowers, with ornamental potential. These include Esterhazya, Escobedia, Agalinis and Physocalyx, but cultivation are difficult since they are parasites.

Key differences from similar families

Orobanchaceae could be differentiated from Scrophulariaceae by number of locules in the ovary (1 in Orobanchaceae, 2 in Scrophulariaceae). In the current circunscription, the main difference between the two families is the parasitic habit, not always obvious, since some Orobanchaceae, such as Esterhazya, Agalinis and Physocalyx can be robust shrubs, apparently independent of other plants; from Verbenaceae, Acanthaceae and Lamiaceae, which Orobanchaceae is sometimes confused, Orobanchaceae can be differentiated by the number of ovules (many in these three families and 1 in Orobanchaceae) in addition to the parasitic habit.

SYSTEMATIC three high clades, Rehmannieae (2/7; endemic to China) and Lindenbergieae (1/12, NE Africa, tropical Asia to Philippines, with their highest diversity in India) does not occur in South America; among genera in Orobanchoideae (97/1,820–2,100) tribe Cymbarieae (6/24, E U.S.A., E Mediterranean, SW Asia, Ukraine and southern Russia to Central and E Asia) does not occur in South America.

Key to genera of South American Orobanchaceae

The morphological distintion between some genera of Euphrasinae is unclear and the key, in this sense, is provisional, waiting for a comprehensive phylogenetic and morphological work concerning to this group.

1. Plants holoparasitic, without chlorophyll; ovary unilocular ------------ Aphyllon

1. Plants hemiparasitic (mostly not evidently so), with chlorophyll; ovary bilocular - 2

2. Calyx strongly zygomorphic, sometimes spathe -like - 3

3. Climbers or scandent plants; corolla with the upper lip not domed ------------ Velloziella

3. Erect plants; corolla generally with the upper lip domed - 4

4. Leaves alternate; bracts showy (more than flowers) ------------ Castilleja

4. Leaves generally opposite or verticillate; bracts not showy - 5

5. Calyx spathe -like ------------ Nothochilus

5. Calyx campanulate, not spathe -like - 6

6. Corolla ventricous (inflated) ------------ Lamourouxia

6. Corolla not ventricous - 7

7. Corolla with upper lip laterally compressed and entire or bilobed ------------ Pedicularis

7. Corolla with upper lip not laterally compressed and always entire ------------ Neobartsia

2. Calyx actinomorphic or almost so - 8

8. Corolla salver-shaped - 9

9. Anthers 1-thecate; corolla up to 3 cm long; generally pink, purple or violet ------------ Buchnera

9. Anthers 2-thecate; corolla more than 3 cm long; white or cream ------------ Escobedia

8. Corola mostly campanulate or bilabiate, not salver-shaped - 10

10. Corolla evidently 2-lipped - 11

11. Corolla with upper lip laterally compressed ------------ Pedicularis

11. Corolla with upper lip not laterally compressed - 12

12. Corolla inflated, red or pink and over 2.5 cm long ------------ Lamourouxia

12. Corolla not inflated, yellow, white, pink, generally under 2.5 cm long - 13

13. Corolla yellow - 14

14. Calyx with multipartite lobes ------------ Agalinis

14. Calyx not with multipartite lobes ------------ Melasma

13. Corolla red, pink or orange - 15

15. Leaves with serrate margins ------------ Magdalenaea

15. Leaves with entire margins - 16

16. Stamens exsert, anthers villous ------------ Esterhazya

16. Stamens included; anthers glabrous or almost so - 17

17. Corolla rose to pink; calyx not inflated, green or purple ------------ Agalinis

17. Corolla orange; calyx inflated, also orange ------------ Physocalyx

10. Corolla tubular, campanulate, subrotaceous or subglobose, not 2-lipped - 18

18. Corolla with upper lip entire; capsule ovoid to subglobose ------------ Neobartsia

18. Corolla with upper lip generally bilobed; capsule oblong to lanceolate, flattened ------------ Euphrasia

OROBANCHOIDEAE ▸ UNPLACED GENERA - 9 outsiders from Africa, 6 from U.S.A./Mexico (some endemics to a own single country), Seymeriopsis from Cuba, 4 from Madagascar, 6 from NE, E, SE Asia, Leptorhabdos (1; Caucasus, Iran to Central Asia and Himalayas) and Micrargeria (3; tropical Africa, India) are unplaced in tribes

1. Magdalenaea Brade. Perennial shrubs, glabrous; stems erect, leaves cauline. Only one sp., M. limae Brade, a rare plant in Brazil, by Plantas Raras do Brasil’s book, endemic to the Pedra Dubois, Santa Maria Madalena in eastern Brazil; many recent expedictions were made ir order to find this spp. in field with no results; it is possible to be extincted.

2. Nothochilus Radlk. Perennial suffrutescent herbs or dwarf shrubs, pubescent; stems erect, terete, rarely with secondary branches; inflorescences raceme, with green bracts, orange flowers. Only one sp., N. coccineus Radlk., ornamental, a rare plant in Brazil, by Plantas Raras do Brasil’s book, endemic in rocky grasslands (campos rupestres) to the Caparaó Massif between Minas Gerais and Espírito Santo states in Eastern Brazil, possibly parasiting only Chusquea bamboos, in a small area of less 5 km²; highly endangered.

3. Physocalyx Pohl. Perennial suffrutescent shrubs, pubescent; stems erect, terete. Three spp., endemic to the Espinhaço Range in Bahia (only P. scaberrimus Philcox, a rare plant in Brazil, by Plantas Raras do Brasil’s book) and Minas Gerais (2), Brazil.

4. Velloziella Baill. Herbs to shrubs, mainly scadent, unique climbings among Orobanchaceae, perenial; stems erect, flexuose, terete. Three spp., two in southern & SE Brazil, and V. spathacea (Oliv.) Melch. restricted in Guiana Shield, from Roraima and adjacent areas of Venezuela and Guyana.

1.1 OROBANCHOIDEAE ▸ TRIBE OROBANCHEAE (12/c. 180) - outsiders Kopsiopsis (2; W North America), Xylanche (1; Himalayas, China inc. Taiwan), Boschniakia (1; temperate Asia to Japan, NW North America), Conopholis (3; SE U.S.A., Mexico to Panamá), Epifagus (1; North America); Orobanche (c 100; temperate and subtropical regions of both hemispheres), Boulardia (1; Spain), Diphelypaea (2–4; SE Europe, Türkiye, the Caucasus), Phelipanche (c 50; Europe, Mediterranean, Macaronesia, W and SW Asia), Mannagettaea (2; E Siberia to W China), Cistanche (16; Mediterranean, Ethiopia to W India and NW China).

5. Aphyllon Mitch. (off Orobanche) Herb, annual or rarely perennial, achlorophyllous, holoparasitic; stems fleshy; leaves reduced to scale-like bracts; inflorescences terminal racemes, spikes, corymbs, or panicles; calyx 5-toothed; corolla sympetalous, bilabiate to regular, tubular and often curved; fruit loculicidal capsules. 22 spp., 18 in North America, and 4 in South America: A. chilense (Phil.) A.C. Schneid. (Chile and Argentina, parasiting almost exclusively Grindelia chiloensis), A. tacnaense (Mattf.) A.C. Schneid. (S Peru, and probably Cuzco, parasiting mainly Franseria and Artemisia, both Asteraceae), A. tarapacanum (Phil.) A.C. Schneid. (N Chile, uncollected from 1891 to 1972) and A. weberbaueri (Mattf.) A.C. Schneid. (S Peru, and known from the type collection).

1.2 OROBANCHOIDEAE ▸ TRIBE RHINANTHEAE (15/340-510) - outsiders Pterygiella (5; southern China); Melampyrum (c 35; temperate regions on the northern Hemisphere), Rhinanthus (c 45; temperate regions on the northern Hemisphere), Lathraea (4; Europe, temperate Asia), Bartsia (1; cold-temperate and alpine regions in Europe and North America), Rhynchocorys (8; southern Balkan Peninsula, C Mediterranean to Iran), Tozzia (1; the Alps, the Carpathians), Odontitella (1; the Iberian Peninsula), Nothobartsia (2; W Mediterranean), Hedbergia (1; mountains in tropical Africa), Parentucellia (3; Mediterranean to SW Asia), Macrosyringion (2; Mediterranean), Odontites (c 30; Europe, Mediterranean to Himalayas).

6. Euphrasia L. 213 spp. northern hemisphere (16 in North America) up to New Guinea, nearly absent in Africa and U.S.A., Oceania, and South America (7, two sections, both endemic: Paradoxae (monotypic, Juan Fernandes) and Trifida (6, mainly Chile, 5 in Argentina, E. antarctica Benth. up to Bolivia, and E. pubescens Benth. endemic to Peru).

7. Neobartsia L. Annual or perennial hemiparasitic herbs or subshrubs, usually pubescent and often glandular; stems prostrate, scandent, or erect, branched mainly in the proximal parts; corolla pubescent, yellow, red, purple, or lavender with the lobes equal to unequal. 48 spp., Andean South America from Colombia to Argentina, up 2,800m elevation range.

1.3 OROBANCHOIDEAE ▸ TRIBE BUCHNEREAE (20/c. 390) - outsiders Cyclocheilon (3; Somalia, Ethiopia, Arabian Peninsula), Asepalum (1; Ethiopia, Kenya, Tanzania, Yemen), Sopubia (41; Africa, Himalayas to Indochina and Taiwan in China), Graderia (4; Africa, Socotra), Nesogenes (8; Tanzania, Madagascar, islands in the Indian and Pacific oceans), Radamaea (5; Madagascar), Bardotia (1; Madagascar), Rhamphicarpa (5; Russia, Africa, Türkiye, India, tropical Australia), Sieversandreas (1; Madagascar), Xylocalyx (5; Somalia, Socotra), Striga (40–45; tropical, subtropical and southern Africa, SE Asia, northern Australia), Cycnium (18; tropical E Africa to South Africa), Centranthera (c 13; China and S to N and NE Australia), Alectra (c 40; Africa, tropical Asia), Hyobanche (9; southern Africa), Harveya (c 30; Africa, Mascarene Islands), Aeginetia (8; India, Sri Lanka and Burma to China, Japan and SE Asia, Malesia), Christisonia (16–20; SW China, E Asia, Malesia).

8. Buchnera L. Herbs to subshrubs, sometimes with xylopodium. 140 spp., Africa and southern U.S.A. to Argentina, with 24 in New Wortld, 19 in South America, 11 in Brazil, 4 endemics.

9. Melasma P.J. Bergius (inc. Alectra). Perennial suffutescent herbs, sometimes shrubs; stems erect, terete. 40-50 spp., Africa and New World, three in New World, M. melampyroides (Rich.) Pennell and M. rhinanthoides (Cham.) Benth. in Brazil, Paraguay and Argentina, and M. physaloides (D. Don) Melch. from Central America and Mexico.

10. Escobedia Ruiz & Pav. Perennial herbs to subshrubs, glabrous or pubescent, roots yellow. 9 spp., Mexico to S Brazil and Bolivia, highly centered in Central America; only three in South America, two endemics to Colombia and the widely distributed E. grandiflora (L. f.) Kuntze in over continent, inc. Brazil.

1.4 OROBANCHOIDEAE ▸ TRIBE PEDICULARIDEAE (11/870-970) - oustiders Phtheirospermum (5; E Asia), Aureolaria (8; U.S.A., Mexico), Seymeria (c 25; U.S.A., Mexico), Cordylanthus (19; North America), Orthocarpus (8; Canada, W U.S.A., NW Mexico), Triphysaria (6; British Columbia to California).

11. Agalinis Raf. (inc. Anisantherina). Annual or perennial herbs, glabrous or scabrid; corolla mainly pink tons. 68 spp., from the coastal plain of Nova Scotia in Canada, to the puna communities above 3,000 m in the Andes of South America (28); North and Central America are represented by about 35; 13 spp. in Brazil, 10 endemics, 4 of then, all in Minas Gerais state, are rare plants in Brazil, by Plantas Raras do Brasil’s book.

Agalinis from North America are herbaceous while shrubs and mainly subshrubs are common in South America; most Brazilian species occur in the SE and S, mainly in rocky grasslands (campos rupestres) of Minas Gerais state, generally restricted to certain mountain ridges or groups of ridges; one from the species, A. hispidula (Martius) D 'Arcy, has a much broader distribution, ranging from the Antilles region to about 20° southern latitude.

12. Castilleja Mutis ex L.f. Annual or perennial herbs, glabrous or pubescent; stems erect; bracts more attractive than flowers. 208 spp., three in E North America, 5 in Eurasia, 30 in Central America and South America (22 in latter, including 7 annual species in Andes of Chile and Peru), and the rest in W North America, only the widely distributed C. arvensis Schltdl. & Cham. in Brazil.

13. Esterhazya J.C.Mikan. Perennial shrubs, stems erect; leaves cauline, opposite, flowers pink to red. 5 spp., Brazil (all species, 4 endemics, three are rare plants in Brazil, by Plantas Raras do Brasil’s book, narrow endemics in highlands of E Brazil in Minas Gerais and Rio de Janeiro states), Paraguay and Bolivia (only E. splendida J.C. Mikan).

14. Lamourouxia Kunth. Perennial suffutescent herbs. 24 spp., N Mexico to Andean Colombia, Ecuador and Peru, with L. virgata Kunth in Colombia to Peru, and L. sylvatica Kunth in southern Ecuador and Peru.

15. Pedicularis L. Annual, biennial or perennial, half rosulate or rosulate herbs, glabrous to densely villose. 677 spp., 43 in New World, 42 in North America to Panamá, and P. incurva Benth. in Andes of Colombia and Ecuador.

LINEAGE 8 of 8: BIGNO/VERBENO/ACANTHOIDS

SCHELEGELIACEAE

§ FULLY CHLOROTROPHIC

Genera/spp. 4/30–35 Distribution S Mexico, Central America, NW South America, Cuba (Synapsis). Habit bisexual, evergreen trees or shrubs (sometimes twining, often epiphytic). Bark often whitish.

SYSTEMATICS Outsider Synapsis (1; E Cuba).

1. Exarata A. H. Gentry. Large canopy trees, branchlets terete; flowers distinctly pedicellate. Only one sp., E. ilicifolia Griseb., forests of Choco, in Ecuador and Colombia.

2. Gibsoniothamnus L.O. William. Small epiphytic or rarely terrestrial shrubs; stems erect, rounded. 12 spp., restricted of Central America (six spp. endemic to Panamá), except G. alatus A.H. Gentry up to Colombia and G. cornutus (Donn. Sm.) A.H. Gentry up to Mexico.

3. Schlegelia Miq. Small epiphytic or rarely terrestrial shrubs, trees or vines; stems erect, rounded. 24 spp. from Mexico, Central America, Antilles, Guianas, Colombia, Venezuela, Ecuador, Peru and Brazil; 20 in South America, 7 in Brazil, one endemic; S. dressleri A.H. Gentry from Bajo Calima region, Choco, W Colombia, has the largest leaf among Schlegeliaceae.

MARTYNIACEAE

§ FULLY CHLOROTROPHIC

Genera/spp. 5/16 Distribution warm and arid or semiarid regions from the southern U.S.A. southwards to Argentina. Habit bisexual, usually annual (rarely perennial) herbs (rarely shrubs, in Holoregmia with fleshy young stems). Craniolaria annua L. has a large root tuber. With densely spaced glandular hairs and evil-smelling. Mycorrhiza probably absent.

Ibicella lutea (Lindl) Van Eselt,the Yellow-Unicorn Plany, is naturalized elsewhere, and possibly carnivorous Some spp. (Proboscidea, Ibicella) are possibly carnivorous; the glands are similar to those in Byblis and Lentibulariaceae, although this may be a precursor to insect-trapping. Martyniaceae are an exclusively New World family and confined to areas of Mediterranean, tropical and subtropical climate. The distribution areas ranges from the southern U.S.A. to northern Argentina. Fragnment of old Pedaliaceae. Three genera, with one sp. each in Brazil. One genera absent in South America, Martynia from Mexico and Caribbean.

SYSTEMATIC outsider is Martynia (1; Mexico, Central America, the Greater Antilles).

1. Craniolaria L. Annual herbs with xylopodium; corolla white to yellowish. Three spp., C. annua L. in Colombia to Guyana, Hispaniola, Jamaica, Porto Rico, and two in Argentina, one of them up to S Brazil and Bolivia.

2. Holoregmia Ness. Shrub, robust much-branched woody to small tree, up to 3m tall, younger stems fleshy; corolla pale ochraceous yellow; viscid leaves, large, lobed. Only one sp., H. viscida Ness., SE Bahia state in NE Brazil, with very restricted distribution in Diamantina Range, in Rio das Contas valley, sometimes in disturbed habitats; most primitive genera in this family.

3. Ibicella (Stapf) Van Eselt. Annual herbs; corolla pale to deep yellow; fruit with two long horns. Two spp., both in Argentina and Paraguay, one up to S Brazil, another up to Bolivia.

4. Proboscidea Schmidel. Annual or perennial herbs, sometimes with a thickened primarly root. 8 spp. from southern U.S.A. to Mexico, P. altheifolia (Benth.) Decne. disjunct in Peru.

BIGNONIACEAE

§ FULLY CHLOROTROPHIC

Genera/spp. 79/827 Distribution mainly tropical regions, with their largest diversity in South America; some spp. in subtropical and warm-temperate Asia. Habit bisexual, usually evergreen (rarely deciduous) trees, shrubs or lianas (Argylia, Incarvillea (inc.Niedzwedzkia) are perennial herbs, Tourrettia consists of twining perennial herbs). Often with leaf-tendrils. Lenticels often frequent on stems and branches. The family, cosmopolitan, is present in both the Old World and the New World, with Campsis and Catalpa the only genus common to both; imponent tropical family compound br usually trees and lianas. 104 genera and 800 spp., centered in Neotropics. Few herbs, such as Argylia and Tourretieae.

Some spp. of Tabebuia and Catalpa are used for timber. Jacaranda, Tabebuia and Crescentia provide ornamentals widely cultivated in subtropical and tropical regions. Several spp. have been reported to contain active compounds against major diseases such as cancer (bark of Tabebuia), malaria, hepatitis, leishmaniases, diabetes, etc; the onion and garlic-smelling spp. (Mansoa) and clove-smelling spp. (Tynanthus) are used as condiments throughouts the Neotropics. B. nocturna (Barb. Rodr.) L.G. Lohmann is used in love portion and hallucinogenous. Fridericia chica (Humb. & Bonpl.) L.G.Lohmann. produces a red dye which is used to dye basket fibres and for Body paints by native peoples in Amazon; fruits of Crescentia are used for maracas instruments.

SYSTEMATICS nine clades, eight as tribes, Catalpeae (1–2/11, E Asia, SE U.S.A., the Greater Antilles), Oroxyleae (4/6–8, tropical Asia) and Coleeae (18/140–145, tropical Africa, Madagascar, Indian Ocean islands, NW and W India, SE Asia, Malesia and eastwards to northern Queensland and SW Pacific Islands) do not occur in New World.

UNPLACED GENERA – only two, both in South America.

1. Paratecoma Kuhlm. Large trees up to 65 m tall, the tallest of all Lamiales in Brazil, stems glabrous, corolla white, salver-former tubular, inflorescence in axillary fascicles. Only one sp., P. peroba (Record) Kuhlman, in Rio Doce valley, in Minas Gerais, Espírito Santo and Rio de Janeiro states in SE Brazil.

2. Romeroa Dugand. Small to medium-sized trees, corolla yellow. Only one sp., R. verticillata Dugand, endemic to Magdalena Valley in from Colombia.

1. TRIBE JACARANDEAE (2/50) ‣ a single genus.

3. Jacaranda Juss. (inc. Digomphia) Trees or shrubs up to 40 m tall, sometimes xylopodium subshrubs (centered in savannas of center Brazil), stems glabrous pubescents, corolla blue or purple blue to magenta. 52 spp., from Guatemala to South America (46), 39 in Brazil, 24 endemics; 4 sections:

§ sect. Copaia ‣ monothecal anthers and cupular calyces; a single species, J. copaia (Aubl.) D. Don, from Central America to Bolivia, Brazil and Guianas.

§ sect. Dilobos ‣ dithecal anthers and cupular calyces, and including more than half of the species of the genus, all restricted to Brazil; 7 spp. in several states are rare plants in Brazil, by Plantas Raras do Brasil’s book

§ sect. Jacaranda ‣ monothecal anthers and campanulate calyces; 11 spp. from South America, J. caucana Pittier up to Central America, and six restricted for Caribbean; a specimen of J. decurrens Cham. (Bignoniaceae) - an endangered species from savannas of C Brazil and adjacent Paraguay (cerrado) - in Ouro Grosso range, Itutinga municipality, Minas Gerais State, Brazil, with a crown diameter of 22m; the mean age calculated for the individual was 3,801 years, making it one of the oldest known living Neotropical plants; this species is an obligate geophyte with small seasonal aerial branches growing from woody underground systems.

§ sect. Nematopogon ‣ species previously included in Digomphia and united by divided staminode apices and spathaceous calyces; 3 spp., two only in Guiana Shield of Venezuela, Guyana, Brazil, Colombia, at 100-2,500 m elevation range, another disjunct in this area and Andes (Guyana to Bolivia and Peru).

2. TRIBE TOURRETIEAE (2/4) ‣ two genera, both in South America.

4. Eccremocarpus Ruiz & Pavon. Wiry vine with multifoliate, usually tripinnatisect tendrillate leaves. 5 spp. from Andes of Colombia to Peru, E. scaber Ruiz & Pav. up to Argentina and Chile.

5. Tourretia Foug. Annual herbs (unique at family in New World plus Argylia), with vine with tendrillate leaves. Only one sp., T. lappacea (L'Hér.) Willd, Mexico to Bolivia, east up to Venezuela, and Argentina.

3. TRIBE TECOMEAE (11/72) ‣ outsiders Campsis (2; E Asia; SE U.S.A.), Incarvillea (c 16; C Asia and Himalayas to E Asia), Podranea (1; tropical and southern Africa), Deplanchea (c 8; Malesia to New Guinea, Australia and New Caledonia), Lamiodendron (1; New Guinea), Tecomanthe (5–6; Malesia to New Guinea, New Britain, Solomon Islands; New Guinea; Australia, New Zealand), Pandorea (9; Malesia to New Guinea, Australia, New Caledonia), Astianthus (1; Mexico to Nicaragua).

6. Argylia D. Don. Perennial chamaephytic subshrubs to herbs (unique at family in New World plus Tourretia), stems sometimes terminating in spine. 13 spp. from Chile and Argentina, A. radiata (L.) D. Don up to southern Peru.

7. Campsidium Seem. Liana, stem glabrous, except for a few, minute lepidote scale, composed leaves, pink flowers. Only one sp., C. valdivianum (Phil.) W. Bull, in the Valdivian rainforest of Chile and Argentina.

8. Tecoma Juss. Shrub or small trees, rarely subscandent. 8 spp., 7 from Ecuador to Chile and Argentina, and T. stans (L.) Juss. ex Kunth in all (sub-)tropical New World from Arizona and Florida in U.S.A. up to Argentina, widespread and common around the Caribbean, occurring in most of the West Indies and thence along the mountain chains from Mexico south to northern Argentina; in South America rarely far from the Andes and apparently absent as a native plant from Brazil. It is widely cultivated in Bolivia and throughout the tropics..

4. TRIBE DELOSTOMEAE (1/4) ‣ a single genus.

9. Delostoma D. Don Tree or shrubs, stems glabrous to pubescent; corolla red or magenta. 4 spp., Andes from Venezuela to Peru.

5. TRIBE BIGNONIEAE (20/415–420) ‣ all genera occur in South America.

10. Amphilophium Kunth. Lianas, stems with multiples of four phloem arms in cross section; corolla cream or purple, tubular. 46 spp., from Mexico to Caribbean to Argentina, 42 in South America, 28 in Brazil, 12 endemics; A. perbracteatum A.H.Gentry from Bahia state is a rare plant in Brazil, by Plantas Raras do Brasil’s book.

11. Adenocalymma Mart ex Meisn. Lianas, stems with four phloem arms in cross section; corolla yellow, tubular t campanulate. 78 spp. from South America (three also in Central America to Mexico), with endemics in Colombia (3), French Guiana (2) and Brazil, with 70 spp., 49 endemics, 10 of then are rare plants in Brazil, by Plantas Raras do Brasil’s book.

12. Anemopaegma Bureal ex Bail. Shrubs or lianas, stems with multiples of four phloem arms in cross section, sometimes with xylopodium; corolla bright to pale yellow, tubular to campanulate. 48 spp., all in South America, 6 up to Mexico and Central America, 36 in Brazil, 15 endemics; A. mirabile (Sandwith) A.H.Gentry from Piauí state is a rare plant in Brazil, by Plantas Raras do Brasil’s book.

In Bignoniaceae cauliflory seems to have evolved at least three times, once in the Paleotropical clade (Colea Bojer ex Meisn., Ophiocolea H. Perrier and Rhodocolea Baill.), once within the ‘Tabebuia Alliance’ clade (Amphitecna Miers, Crescentia L. and Parmentiera DC.), and once in Adenocalymma Mart. ex Meisn. in Bignonieae. A. cauliflorum L.H. Fonseca & L.G. Lohmann from the Atlantic Forest as the only fully cauliflorous of the genus, of the five species (all in Brazilian) with this characteristic.

13. Bignonia L. Lianas, stems with multiples of 4 phloem arms in cross section; corolla dull red to orange outside, yellow within. 29 spp., 28 from Mexico to Caribbean, Brazil and Paraguay, and B. capreolata L., in southern U.S.A.; 17 spp. in Brazil, one endemic; B. nocturna (Barb. Rodr.) L.G. Lohmann smelling of almond (cyanide) and containing hallucionogenic compounds; B. costata (Bureau & K.Schum.) L.G.Lohmann from Rio de Janeiro state is a rare plant in Brazil, by Plantas Raras do Brasil’s book.

14. Callichlamys Miq. Lianas, shrub when young, stems with 4 phloem arms in cross section; corolla campanulate, yellow to orange. Only one sp., C. latifolia (L. Rich) K. Schaum., from Mexico to Brazil in over tropical America.

15. Cuspidaria DC. Lianas, stems with 4 phloem arms in cross section; corolla cream to pale yellow or magenta, tubular to campanulate. 19 spp. in South America, C. inaequalis (DC. ex Splitg.) L.G. Lohmann up to Panamá, 18 in Brazil (C. weberbaueri (Sprague) A.H. Gentry endemic to Peru), 7 endemics.

16. Doliochandra Cham. Lianas, stems with many irregularly divided phloem arms in cross section; corolla red, tubular. 10 spp. from South America, 4 up to Central America and Mexico, 9 in Brazil (D. hispida (DC.) L.H. Fonseca & L.G. Lohmann only in Cono Sur), two endemics.

17. Fridericia Mart. Lianas, rarely small trees or shrubs, stems with 4 phloem arms in cross section; corolla red or pale pink to purple, campanulate. 61 spp. from Mexico to Argentina and Caribbean, 57 in South America, 53 in Brazil (19 endemics, 4 of then in Maranhão, Bahia and Rio de Janeiro states are rare plants in Brazil, by Plantas Raras do Brasil’s book).

18. Lundia DC. 14 spp., of South America, two up to Mexico and Central America, 13 in Brazil (L. cordata (Vell.) DC. endemic to Peru), 8 endemics; L. damazii DC. from Minas Gerais state is a rare plant in Brazil, by Plantas Raras do Brasil’s book.

19. Manaosella J. C. Gomes. Only one sp., M. cordifolia (DC.) A.H. Gentry, from southern Venezuela, Brazil and Bolivia.

20. Mansoa DC. 17 spp. of South America, 4 up to Mexico and Central America, 16 in Brazil (M. gentryi M. M. Silva endemic to Peru), 9 endemics; M. onohualcoides A.H.Gentry from Ceará state is a rare plant in Brazil, by Plantas Raras do Brasil’s book.

21. Martinella Baill. Lianas, stems with 4 phloem arms in cross section; corolla yellow to purplish with dark nectar guides, tubular to campanulate. 5 spp., M. obovata (Kunth) Bureau & K. Schum. widely distributed in Neotropics, all in Brazil, two endemics, one only in Brazil and Peru.

22. Pachyptera DC. ex. Meisn. Lianas, stems with 4 phloem arms in cross section. 5 spp., P. kerere (Aubl.) Sandwith from Honduras to Bolivia, Brazil and Guianas, two only in Venezuela and Colombia, P. incarnata (Aubl.) J.N.C. Franc. & L.G. Lohmann in Brazil and Guianas, and P. aromatica (Barb. Rodr.) L.G. Lohmann endemic to Brazil.

23. Perianthomega Bureal ex Bail. Lianas, stems lepidote. Only one sp., P. vellozoi Bureau, S Brazil, northern Paraguay, and Bolivia.

24. Pleonotoma Miers. Lianas with tetragonal branches, prophylls or pseudostipules of axillary bud foliaceous or stipule-like, interpetiolar glandular fields absent, ternately compound leaves, and the inflorescence a simple raceme. 16 spp. from South America, to SE Brazil and westwards from the Atlantic to Colombia, Peru and Bolivia east of the Andes, in tropical wet forests of Central and South America and also in the gallery forests and dry forests of the Brazilian and Bolivian highlands, P. variabilis (Jacq.) Miers up to Guatemala and Trinidad and Tobago, 14 in Brazil, 7 endemics; from, the centre of diversity is in Brazilian Amazonia where 8 spp.

25. Pyrostegia C. Presl. Lianas, stems with multiples of four phloem arms in cross sections, compound leaves with two leaflets and a terminal tendril; corolla orange-red, narrowly campanulate to tabulate (3 hummingbird pollinated and 1 moth pollinated). Two spp., P. venusta (Ker Gawl.) Miers widely distributed tropical South America, and P. millingtonioides Sandwith exclusive of Pará and Maranhão states in Brazil; in dry secondary growth.

26. Stizophylum Miers. Lianas, stems with four phloem arms in cross section. Three spp., all widely distributed or scattered in S Mexico to Bolivia and Brazil (all, none endemics).

27. Tanaecium Sw. Lianas; stems with multiples of four phloem arms in cross section. 22 spp., Mexico to Bolivia, Brazil and Guianas, 21 in South America, 16 in Brazil, four endemics.

28. Tynanthus Miers. Lianas, stems with four phloem arms in cross section, corolla white. 14 spp., Mexico to Bolivia and Brazil, 12 in South America, 10 in Brazil, 5 endemics.

29. Xylophragma Sprague. Lianas, stems with four phloem arms in cross section. 8 spp., all in Brazil, 5 endemics, two up to over South America, X. seemannianum (Kuntze) Sandwith up to Central America and Mexico.

6. TRIBE CRESCENTIEAE (12/c. 160) ‣ outsiders Ekmanianthe (2; Cuba, Hispaniola).

30. Amphitecna Miers. Leaves simple, often with a red pulvinus on short petiole; inflorescence terminal or cauli/ramiflorous; flowers greenish or creamy white, petal lobes united into rim; fruits oblong (if orbicular, then mangrove species). 20 spp. from Florida, Mexico and Caribbean, two up to Colombia and A. latifolia (Mill.) A.H. Gentry from Mexico to Ecuador, Venezuela, Caribbean and Florida.

31. Crescentia L. Small to medium sized tree; leaves simple or 3-foliate; fruit pepo or calabash, large, sphaerical or elliptical; inflorescences cauliflorous, flowers greenish white, sometimes with red lines. 6 spp., 4 only in Central America and Caribbean, C. cujete L. from Mexico to Colombia, Venezuela and Caribbean, and C. amazonica Ducke in northern South America, a strictly amazonic species, in Brazil, Venezuela, Colombia and Peru.

32. Cybistax Mart. ex. Meisn. Shrub or tree; stems glabrous to lepidote; corolla light green. Only one sp., C. antisyphilitica (Mart) Mart., from Ecuador to Bolivia, Argentina, Paraguay, Suriname and Brazil.

33. Godmania Hemsl. Small to medium sized trees, stems pubescent, corolla yellow or brown. Two spp., G. aesculifolia (Kunth) Standl. from Mexico to Bolivia, Brazil, up to Guyana, and G. dardanoi (J.C. Gomes) A.H. Gentry of Brazil and Venezuela.

34. Handroanthus Mattos. Trees or occasionally shrubs; wood, very dense with high specific gravity; leaves (3)5–9 foliolate (reported to be occasionally 1-foliolate in H. pumilus (A.H. Gentry) S.O. Grose and H. selachidentatus (A.H. Gentry) S.O. Grose, both from Brazil and Bolivia), inflorescence dichotomously branched; calyx coriaceous, campanulate; corolla yellow or magenta with yellow throat, tubular-infundibuliform to tubular-campanulate, fruit an elongate linear to cylindric capsule. 33 spp., all in South America, six up to Central America or Caribbean; 27 spp. in Brazil, 15 endemics, 4 of then in Bahia, Espírito Santo, Paraná and Santa Catarina states are rare plants in Brazil, by Plantas Raras do Brasil’s book.

35. Parmentiera DC. Leaves 3–5 foliolate, often with stipular spines; fruit a linear capsule, bilocular throughout, seeds small with vestigial wings, no pulp in fruit; flowers terminal and cauliflorous, greenish white; calyx membranaceous. 10 spp., 9 from from Mexico to Panamá, and P. stenocarpa Dugand & L.B. Sm. endemic to NW Colombia.

36. Roseodendron (Rose) Miranda. Trees to 35 m; leaves palmately 5–7 foliolate; inflorescence a large terminal panicle; corolla yellow, sometimes with reddish lines in throat, tubular-infundibuliform. Two spp., R. donnell-smithii (Rose) Miranda, ranging from Mexico to Ecuador and Colombia, and R. chryseum (S.F. Blake) Miranda in dry areas of Venezuela and Colombia.

37. Sparattosperma Mart. ex. Meisn. Medium sized to large trees, branchelets glandular-lepidote; corolla white or pale pink. Two spp., S. catingae A.H. Gentry endemic to Bahia state in NE Brazil (a rare plant in Brazil, by Plantas Raras do Brasil’s book), and S. leucanthum (Vell.) K. Schum. from Venezuela to Paraguay and Brazil.

38. Tabebuia Gomes ex DC. Trees or shrubs up to 35 m tall; leaves 1–7(9) foliolate; inflorescence dichotomously branching; corolla white to red, often with yellow throat, completely yellow in two species (T. nodosa (Griseb.) Griseb. and T. aurea (Silva Manso) Benth. & Hook. f. ex S. Moore) tubular-infundibuliform to tubular-campanulate; fruit na elongate linear to cylindric capsule. 75 spp., widely distributed throughout Central and South America (16) and the Antilles, only 11 in Brazil, 4 endemics; T. reticulata A.H.Gentry from Espírito Santo state is a rare plant in Brazil, by Plantas Raras do Brasil’s book.

39. Zeyheria Mart. Shrubs or subshrubs, to 3 m tall (up to 35 m tall in Z. tuberculosa (Vell.) Bureau ex Verl.); leaves 5-foliolate; corolla, yellow/brown, sometimes pinkish in throat; calyx bilabiate, split to base, densely pubescent with stellate hairs; fruit flattened orbicular, with spiny projections, also densely stellate tomentose; seeds orbicular. Two spp. in open savannhas from center Brazil (NE region to center wetlands, both species, none endemics) to E Bolivia and Parguay.

ACANTHACEAE

§ FULLY CHLOROTROPHIC

Genera/spp. 207/3,770–3,780 Distribution tropical regions, especially in S and SE Asia, Africa, Brazil and Central America, some spp. in warm-temperate regions; Avicennia and Hilairanthus: mangrove vegetation in tropical and subtropical areas in the Southern and Northern Hemispheres. Habit usually bisexual (rarely unisexual), usually perennial or annual herbs, evergreen shrubs or lianas (rarely evergreen trees; Avicennia and Hilairanthus consists of mangrove trees or mangrove shrubs with articulated branchlets). Some representatives are aquatic. Many spp. are xerophytic. Some spp. are epiphytic.

SYSTEMATIC all subfamilies occur in South America; only one gnus unplaced at subfamilies: Ritonia (4; Madagascar).

1. SUBFAMILY NELSONIOIDEAE (8/105-110) ‣ outsiders Nelsonia (1; Africa, Madagascar, tropical Asia to tropical Australia), Saintpauliopsis (1; tropical Africa), Anisosepalum (3; Central Africa).

1. Aymoreana Braz, T.F.Daniel & C.Kiel. (off Staurogyne) Herb shortly caulescent, stems up to 3 cm long, reddish, not branched, with dense multi-cellular eglandular trichomes; iInflorescence of an axillary panicle of spikes. Only one sp., A. nitida (S.Moore) Braz, T.F.Daniel & C.Kiel, restricted to the Atlantic Forest, from central Espírito Santo northward to southern Bahia states, E Brazil.

2. Elytraria Michx. 22 spp., tropical and subtropical regions on both hemispheres, 15 in New World, 8 endemics to Caribbean (inc. E. serpens Greuter & R. Rankin from Cuba, off VPA), 4 only North America/Central America, and two in South America, E. tuberosa Leonard endemic Ecuador, and E. imbricata (Vahl) Pers., widely distributed in New World.

3. Staurogyne Wall. (exc. Aymoreana) Calyx with unequal segments, with one posterior segment larger than the others, one anterior pair of segments intermediate in size, and with one lateral pair smaller than the others; distinguished by the four, usually didynamous stamens with a reduced staminode between the posterior pair and by the capsules without retinacula and usually with numerous seeds. c. 145 spp., distributed throughout tropical regions of the world, 33 spp. in New World, all in South America (two up to Caribbean and Mexico), 27 in Brazil, 21 endemics - 8 of then, all in SE region, are rare plants in Brazil, by Plantas Raras do Brasil’s book; only two in Mexico and Caribbean.

2. SUBFAMILY ACANTHOIDEAE (c 190/3.190-3,280) ‣ 7 tribes, Neuracanthus clade (1/13, tropical and subtropical E Africa, Madagascar, southern Arabian Peninsula, India to Indochina), Andrographideae (6/105–110, tropical Asia) and Whitfieldia clade (8/c 38, tropical Africa, Madagascar) do not occur in South America.

2.1 ACANTHOIDAE ▸ TRIBE ACANTHEAE (20/490–500) - outsiders Neriacanthus (1, Jamaica), Crossandra (c 55; tropical Africa, Madagascar, Arabian Peninsula, tropical Asia), Crossandrella (3; tropical Africa), Sclerochiton (19; tropical and southern Africa), Streptosiphon (1; Tanzania), Cynarospermum (1; Caribbean), Blepharis (125–130; Mediterranean, tropical regions in the Old World and southwards to South Africa), Acanthopsis (8; Namibia, South Africa), Acanthus (29; warm-temperate to tropical regions in the Old World), Salpixantha (1; Jamaica), Holographis (16; Mexico), Strobilacanthus (1; Panamá).

4. Aphanandrium Lindau. (former Neriacanthus in South America) 5 spp., four from Panamá to Peru and one endemic to Venezuela.

5. Aphelandra R.Br. (inc. Orophochilus, Rhombochlamys, Encephalosphaera) Suffrutescent herbs or shrubs, perennials, stems terete to quadrangular; leaves opposite, petiolate; inflorescence of terminal or axillary spikes, these often conspicuous with showy bracts and flowers; corolla bilabiate, pale to bright red, orange or yellow; stamens 4, usually exserted; anthers 1- thecous; capsule clavate to subglobose, 4-seeded. 206 spp., occurring from NW Mexico southeastward through regions of both wet and dry forests to SE Brazil and NE Argentina, 180 in South America, centered in the Andes of Colombia (66), Ecuador and Peru; 45 spp. in Brazil, 34 endemics, 14 of then, in several Brazilian states, are rare plants in Brazil, by Plantas Raras do Brasil’s book.

6. Cyphacanthus Leonard. Only one sp., C. atopus Leonard, endemic to Boyaca departament in Colombia.

7. Stenandrium Nees. Acaulescent or caulescent herbs lacking cystoliths, perennials; leaves opposite or quaternate; inflorescence of axillary or terminal elongated or headlike usually pedunculate dichasiate spikes, dichasia opposite or alternate, 1-flowered, sessile, subtended by bracts; flowers subtended by 2, sessile bracteoles; calyx deeply 5-lobed, lobes equal or subequal; corolla pink, purple, or white. c. 100 spp., Africa, Madagascar, tropical and subtropical New World (49), 28 in South America from Venezuela to Chile and Brazil (17, 12 endemics), also Argentina; 4 spp., from Minas Gerais and Goiás states are rare plants in Brazil, by Plantas Raras do Brasil’s book.

2.2 ACANTHOIDAE ▸ TRIBE BARLERIEAE (12–13/365–370) - outsiders Barleriola (4; Caribbean), Borneacanthus (6; Borneo), Boutonia (1; Madagascar), Chroesthes (6; southern China, SE Asia), Crabbea (14; tropical and southern Africa), Hulemacanthus (2; New Guinea), Lasiocladus (2; Madagascar), Pericalypta (1; Madagascar), Podorungia (5; Madagascar), Pseudodicliptera (2; Madagascar).

8. Barleria L. 300 spp., all in Old World except B. oenotheroides Dum. Coars., shared by Old World and New World, from Mexico to Colombia and Venezuela.

9. Lepidagathis Willd. (inc. Teliostachia, Acanthura, Lophostachys) Small decumbent or erect subshrubs, or erect to sprawling shrubs; decumbent subshrubs usually have adventitious roots at the nodes that touch the substrate; aerial branches are annual and die after flowering and fruiting; simple or compound indeterminate inflorescence; small to large corollas, showy, white, lilac or pinkish with purple, purplish-red or yellow markings. 90 spp., two thirds of which occur in Africa, India and Malesia, 26 in Central and South America, with one endemic to Mexico, L. alopecuroidea (Vahl) R. Br. ex Griseb. widely distributed, and 22 confined in South America, 16 in Brazil, 10 endemics.

2.3 ACANTHOIDAE ▸ TRIBE RUELLIEAE (47/985–990) - 8 subtribes, Erantheminae (5/41-46, Cameroon, tropical Asia, New Guinea, northern and E Australia, New Caledonia), Strobilanthinae (3/c. 190, tropical Asia), Phaulopsidinae (1/18; tropical Africa) and Mimulopsinae (4/24-29, tropical Africa, Madagascar), and several unplaced genera do not occur in South America; 4 genera incertae sedis, none in New World.

■ SUBTRIBE PETALIDIINAE ‣ outsiders Ruelliopsis (2; tropical and southern Africa), Petalidium (c 35; tropical and southern Africa, W India, W Himalayas), Duosperma (26; tropical and southern Africa), Strobilanthopsis (1; tropical Africa).

10. Dyschoriste Nees. Decumbent to erect perennial herbs with cystoliths; leaves opposite; inflorescence of dichasia in leaf axils throughout plant or restricted to axils of distal leaves or bracts and forming a spicate or capitate thyrse; dichasia alternate or opposite, 1- many flowered; corolla blue to blue-purple to white, tube gradually or abruptly expanded distally. c. 80 spp., 46 in New World, southern U.S.A., throughout the Caribbean, Mexico, Central America, and South America (21, Ecuador to Argentina and Brazil (12, 8 endemics)). Old World (OW) species are found in Africa (20), Madagascar (8), and SW to SE (SE) Asia (7-10); D. smithii Leonard from Santa Catarina state is a rare plant in Brazil, by Plantas Raras do Brasil’s book.

■ SUBTRIBE TRICHANTHERINAE ‣ outsiders Louteridium (9; S Mexico, Central America).

11. Bravaisia DC. Shrubs to tallest trees, sometimes very tall, largest corolla campanulate, fertile stamesn 4; inflorescence a thyrsi; corolla white to witish or bluish. Three spp. from Mexico and Cuba southward to Panamá, B. integerrima (Spreng.) Standl. up to coasts of Colombia and Venezuela, mainly coastal forests, inmangroves environments.

12. Sanchezia Ruiz & Pavon. Erect or climbing herbs or shrubs; stems usually glabrous; flowers solitary or more often fascicled, usually large and conspicuous, yellow, orange, red or purple, borne in heads, spikes or racemes, the flower clusters subtended by small or large, rarely partly connate bracts, these sometimes colored. 46 spp., all in South America (only 3 up to Central America), with their largest diversity in the Andes and Amazon rainforest from Peru (31 - 42 endemics in this country), Bolivia and Brazil (4, S. munita (Nees) Planch. endemic), some into southern Colombia and Venezuela.

13. Suessenguthia Merxm. Shrubs, stems erect, quadrangular, sulcate; flowers solitary or fascicled, usually large and conspicuous, borne in heads, spikes or cymes, the flower cluster subtended by small or often large and partly connate bracts, these sometimes colored; corollas lilac or red, tube cylindric, 5-lobed, the lobes equal. 9 spp., these handsome flowering shrubs or treelets are often found growing over-hanging at the edge of streams; all in E Andes in Peru and Bolivia (some endemics each), and S. trochilophila Merxm. up to lowland areas in C & W Amazon rainforests of Brazil.

14. Trichanthera Kunth. Two spp., T. gigantea (Bonpl.) Nees in Central America to Venezuela and Ecuador and N Brazil, T. corymbosa Leonard only in Venezuela and Colombia.

15. Trichosanchezia Midlbr. Herbs. Only one sp., T. chrysothrix Mildbr., endemic to Amazonas department in northern Peru.

■ SUBTRIBE HYGROPHILINAE ‣ outsider Brillantaisia (15–17; tropical Africa, Madagascar).

16. Hygrophila R. Br. c 25 spp., tropical regions on both hemispheres, 12 in New World, one only Caribbean, H. costata Nees widely distributed, 10 only South America; 8 in Brazil, 6 endemics.

■ SUBTRIBE RUELLIINAE (5/c. 320) ‣ outsiders Dischistocalyx (13; tropical Africa), Satanocrater (2; tropical Africa), Acanthopale (9; tropical regions in the Old World).

17. Ruellia L. (inc. Polylychnis, Stachyacanthus) Herbs or shrubs, perennial sometimes with xylopodium; leaves petiolate; inflorescence an axillary, occasionally long-pedicellate flowers in the upper axils combined into a terminal raceme or spike; flowers usually large and showy, regular, sometimes curved; corolla red, yellow, white, or purple (usually mauve), funnelform or salverform, contorted in bud, spreading in flower, sometimes saccate, tubular at base, widened and campanulate into throat. c 350 spp., tropical and subtropical regions on both hemispheres, 253 spp. in New World, 172 in South America, 94 in Brazil, 63 endemics; two spp. from Santa Catarina state are rare plants in Brazil, by Plantas Raras do Brasil’s book. 15 sections in New World, 11 in Brazil.

§ sect. Aphragmia ‣ 12 spp., mainly Mexico and E Brazil; less commonly, species occur in Argentina, Bolivia, and Paraguay or Ecuador and Peru.

§ sect. Blechum ‣ 12 spp., widely in tropical America, inc endemics in Brazil.

§ sect. Boreosilva ‣ 10 spp., endemic to U.S.A.

§ sect. Brasilia ‣ 6 spp., endemics to E Brazil.

§ sect. Cerradicola ‣ 10 spp., endemics to dry savanas in C Brazil (cerrado).

§ sect. Chiropterophilae ‣ 14 spp., possibly endemics to Mexico.

§ sect. Chromatoruellia ‣ 7 spp., Costa Rica to Bolivia, absent in Brazil.

§ sect. Eurychanes ‣ 4 spp., endemics to dry savanas in C Brazil (cerrado).

§ sect. Gymnacanthus ‣ 13 spp., Neotropics from Mexico to Argentina, with most species occurring in Brazil.

§ sect. Mexicanae ‣ 8 spp., mainly endemics to Mexico, R. erythropus (Nees) Lindau up to Brazil.

§ sect. Physiruellia ‣ 14 spp., widely in South America inc. Brazil.

§ sect. Ruellia ‣ 15 spp., mainly North America and northern Central America, five in South America (two in Brazil) and two presumably native in the West Indies.

§ sect. Siphonacanthus ‣ six spp., 5 in Brazil, one up to Peru, Ecuador and Bolivia, and one endemic to Costa Rica.

§ sect. Stephanophysum ‣ c. 60 spp., western South America, especially portions of Peru, Ecuador, Bolivia, as well as northern South America, primarily Venezuela and Colombia, few in Brazil.

§ sect. Strobiliformes ‣ 12 spp., northern South America, primarily Venezuela and Colombia, with a outlier in Bolivia. Absent in Brazil.

2.4 ACANTHOIDAE ▸ TRIBE JUSTICIEAE (95-100/c 1,500) - 5 subtribes, 4 in South America, and few several unplaced genera. Monothecinae (8/39-44, Madagascar, Asia, New Guinea) does not occur in South America.

■ Incertae sedis Justicieae ‣ outsiders Samuelssonia (1; Hispaniola), Ichthyostoma (1; Ethiopia, Somalia), Streblacanthus (5; Central America).

18. Tessmanniacanthus Mildbr. Only one sp., T. chlamydocardioides Mildbr., endemic to Andean region of Peru.

■ SUBTRIBE GRAPTOPHYLLINAE ‣ outsiders Afrofittonia (1; tropical W and C Africa), Ballochia (3; Socotra), Codonacanthus (3; NE India, southern China, Japan), Cosmianthemum (14; Borneo), Filetia (9; the Malay Peninsula, Sumatra), Glossochilus (2; southern Africa), Graptophyllum (c 15; tropical Old World, Australia, Melanesia), Isotheca (1; Trinidad), Linariantha (1; Borneo), Phialacanthus (5; Himalayas to the Malay Peninsula), Sapphoa (2; Cuba), Spathacanthus (3; Central America), Asystasia (c 55; Old World), Ruttya (6; tropical and southern Africa, Madagascar, Yemen), Ruspolia (5; tropical Africa, Madagascar), Wuacanthus (1; China), Chileranthemum (3; Mexico), Mackaya (2; Swaziland, South Africa, southern Asia), Thysanostigma (2; Thailand, the Malay Peninsula).

19. Chamaeranthemum Nees. Three spp., one in Central America and two endemics to SE Brazil.

20. Herpetacanthus Nees. Subshrubs to shrubs, usually anisophylous, calyx with five subequal segments, bilabiate corolla, usually the upper lip bidentate and lower lip trilobate; inflorescences of secundiflorous spikes or thyrsus with two adjacent rows of sterile bracts and two adjacent rows of fertile bracts; cleistogamy is relativaly common. 20 spp., two only in Ecuador, one only Costa Rica, H. panamensis Leonard in Honduras, Nicaragua e Panamá, H. rotundatus (Lindau) Bremek. in Guianas, Brazil, Ecuador, Peru, Bolivia, and all 15 remaining species endemic to Brazil, 14 from the Atlantic Forest in Brazil, from São Paulo State to southern Bahia State (two only São Paulo, three only Espírito Santo, two only Rio de Janeiro, three only in Bahia); one from a deciduous forest in Bahia.

21. Odontonema Ness. 30 spp., 17 in South America, 4 spp. in Brazil, all endemic, O. amplexicaule (Nees) Kuntze in Amazon rainforest and 3 remaining endemics to Atlantic Forest.

22. Oplonia Raf. 19 spp., Madagascar, 16 in tropical America, with their highest diversity in Caribbean, three in South America: two endemics to Peru, and one in Argentina and Bolivia.

23. Pranceacanthus Wassh. Wood herb, shrubs or subshrubs, 0.8 – 2.5 m tall; stems subquadrangular, ascending; leaves oblong elliptic; inflorescences axillary and terminal, pendunculate unilateral spikes, corolla bright red, 3.5 cm long, tubular. Only one sp., P. coccineus Wassh, endemic to Mato Grosso, Rondonia, Acre, Amazonas, and northern Bolivia, occasional in valleys of forests near stream and in humid, mixed forest on ‘terra firma’

24. Pseuderanthemum Radlk. Herbs to shrubs, leaves peciolate to sessile; inflorescence spikes terminal or axillary; corolla hipocrateriforme. 130 spp., tropical regions on both hemispheres, 52 spp. in New World, 41 in South America, highly centered in Andes of Colombia and Ecuador; 11 spp. in Brazil, 10 endemics, P. congestum (S. Moore) Wassh. up to Bolivia.

25. Psilanthele Lindau. Only one sp., P. eggersii Lindau, known only from three old collections, found in the provinces of Guayas, Manabi and Chimborazo, in Ecuador.

26. Pulchranthus V.M.Baum, Reveal & Nowicke. 4 spp., tropical northern South America, Venezuela, Colombia, Ecuador, Bolivia, Peru, Brazil (3, none endemics) and Guianas.

■ SUBTRIBE ISOGLOSSINAE ‣ outsiders Isoglossa (c 50; tropical Old World, Arabian Peninsula), Brachystephanus (22; tropical Africa, Madagascar), Celerina (1; Madagascar), Melittacanthus (1; Madagascar).

27. Sebastianoschaueria Nees. Herbs, inflorescence of terminal spikes; corolla violet, tube cylindric, limb deeply 5-lobed with equal lobes. Only one sp., S. oblongata Nees, endemic to Brazil, in Minas Gerais and Rio de Janeiro states.

28. Stenostephanus Nees. (inc. Kalbreyeriella, Razisea) Erect or spreading herbs or shrubs with cystoliths; leaves opposite, petiolate; inflorescence consisting of terminal and axillary dichasiate spikes, racemes, thyrses or panicles, 1-3- many flowered, flowers sessile or pedunculate; corollas 1- or 2-colored, with the colors various (usually ed, pink, yellow, cream-colored), the tube cylindric or distally gradually or abruptly expanded into a throat. 84 spp. in New World, from W Mexico (15) to Bolivia, Venezuela and Brazil (2, one endemic), 61 in South America, occurring primarily at relatively high elevations (usually between 1,700-2,450 m) with greatest diversity in Colombia (a third of genus here endemic).

■ SUBTRIBE TETRAMERIINAE ‣ outsiders Ancistranthus (1; Cuba), Angkalanthus (1; Socotra), Aphanospermum (1; NW Mexico), Chalarothyrsus (1; Mexico), Chorisochora (3; South Africa, Socotra), Chlamydocardia (3; tropical W and C Africa), Clinacanthus (2; S China, SE Asia, Malesia), Ecbolium (22; tropical regions in the Old World), Gypsacanthus (1; Mexico), Henrya (2; Central America), Hoverdenia (1; Mexico), Kudoacanthus (1; Taiwan in China), Megalochlamys (10; SW tropical Africa to Namibia and northern South Africa, Arabian Peninsula), Mirandea (6; Mexico), Populina (2; Madagascar), Trichaulax (1; Kenya, Tanzania), Yeatesia (3; SE U.S.A. to NE Mexico)

29. Carlowrightia A. Gray. (exc. Thyrsacanthus p.p.). 26 spp. from North America, Central America, and C. ecuadoriana T.F. Daniel & Wassh. endemic to Ecuador.

30. Fittonia Coem. Two spp., F. albivenis (Lindl. ex Veitch) Brummitt from Venezuela, Colombia, Ecuador, Peru, N Brazil and Bolivia, F. gigantea Linden ex André only Peru and Ecuador, but is a widely cultivated greenhouse plant in temperate countries because of its attractive foliage with brightly colored veins.

31. Pachystachys Nees. (inc. Pseuderanthemum p.p.) Herbaceous or suffrutescent plants; leaves large, petioled; spikes terminal, dense; flowers borne in terminal spike of verticillasters consisting of 3 or 4 flowers each; corolla ringent, slenderly obconic, curved, 2-lipped, lower lip 3-lobed, lobes subequal, oblong or ovate, upper lip narrow, 2- lobed apically. 18 spp., along edges of the lowland rain forest at relatively low elevations (below 1,000 m) from Caribbean (2) and northern South America southward to Peru and Bolivia and eastward to French Guiana and Amazonian Brazil (6, two endemics).

32. Schaueria Nees. (inc. Justicia p.p.) Erect perennial herbs to shrubs with cystoliths and often with conspicuously multicelullar trichomes; compound inflorescences, flowers white to yellow. 14 spp., endemic to E Brazil, Bahia to Rio Grande do Sul states, in rainforest, dense woodlands, in dense ombrophyllous forests, semideciduous forests and Atlantic sandy coastal shrublands (restingas), between 80 and 700 m, with species mostly concentrated in Rio de Janeiro and Bahia states.

33. Tetramerium Nees. 30 spp., 23 only in Mexico to Central America, T. nervosum Nees from U.S.A. to Peru, 5 endemics to Peru, and T. wasshausenii T.F. Daniel in Peru, Bolivia and Ecuador.

34. Thyrsacanthus Moric. (inc. Schaueria p.p., Carlowrightia p.p., Justicia p.p., Anisacanthus p.p.) Branched shrubs, cylindrical stems, sometimes dry xeric; inflorescences racemes, spikes or thyrses; corolla mainly red, bilabiate. 8 spp., 5 endemics to Brazil, T. boliviensis (Nees) A.L.A. Côrtes & Rapini in Brazil and adjacent Bolivia and Paraguay, T. sulcatus (Nees) C. Ezcurra & A.L.A.Côrtes from Paraguay, Argentina and Uruguay, and T. secundus (Leonard) A.L.A. Côrtes & Rapini from Venezuela to French Guiana and N Brazil.

■ SUBTRIBE JUSTICINAE ‣ outsiders Anisotes (19; tropical Africa, Madagascar), Ascotheca (1; Gabon), Dasytropis (1; Cuba), Dicladanthera (2; W Australia), Hypoestes (100-105; tropical regions in the Old World), Metarungia (3; tropical and southern Africa), Rhinacanthus (10; tropical regions in the Old World), Rungia (23; tropical regions in the Old World), Trichocalyx (2; Socotra), Vavara (1; Madagascar), Xerothamnella (2; southern Queensland, NW New South Wales, NE South Australia).

35. Cephalacanthus Lindau. Only one sp., C. maculatus Lindau, endemic to Andean region of Peru.

36. Clistax Mart. Shrubs, terrestrial, epiphytic, sometimes scandents; stems cylindrical; leaves peciolate; inflorescence a axillar thyrsi; corolla whitish to lilac, billabiate, non ressupinate. Three spp., endemic to E Brazil; C. bahiensis Profice & Leitman from Bahia state is one of three epiphytic species in this family worldwide.

37. Dicliptera Juss. Annual or perennial herbs, sometimes woody below, occasionally shrubby and scrambling, with cystoliths; stems obscurely hexagonal; inflorescence of small bracteate cymes in axils of upper leaves forming loose, terminal spikes or panicles; corolla strongly bilabiate, usually resupinate, pink, red, orange or white; stamens 2; anthers 2-thecous; capsule 4-seeded, the placenta rising from base as fruit ripens. 80-150 spp., tropical and subtropical regions of the world, 77 in New World, up to Argentina in South America (50), only six in Brazil (only one endemic); although readily recognized by its hexagonal stems and the flattened, bracted cymes, the genus presents difficulties at the specific level.

38. Harpochilus Ness. Herbs to shrubs, few branched; flowers with long emergent thyrses or panicles (each spp. with a own), whitish; large, pale lemon-green or cream-color corollas are strongly bilabiate, a shape uncommon in the chiropterophilia (bat polinizated); capsules obovate, 4 seeded. 4 spp., from rocky slopes and sandstones of Ceará, Rio Grande do Norte, Paraíba, Pernambuco, Alagoas and Bahia states in NE Brazil.

39. Justicia L. (inc. Schaueria p.p., Dianthera, exc. Thyrsacanthus p.p., Anisacanthus p.p.). Herbs or undershrubs with cystoliths, perennials; leaves sessile or petiolate, margin entire or sinuate; inflorescence of 1(-3)-flowered diachasia in leaf axils forming terminal spikes, sometimes compounded into panicles; corolla with a distinct tube, strongly 2-lipped, upper lip entire or 2-lobed, lower lip 3-lobed. c. 600 spp., worldwide; 526 spp. in New World, 350 in South America, 138 in Brazil (86 endemics); 6 spp. from Acre, Distrito Federal and Rio de Janeiro states are rare plants in Brazil, by Plantas Raras do Brasil’s book.

40. Megaskepasma Lindau. Only one sp., M. erythrochlamys Lindau, confined to Venezuela and Suriname.

41. Poikilacanthus Landau. Herbs to subshrubs, perennials, sometimes with xylopodium, inflorescemces spikes or capitula, axillary or terminal; corolla bilabiate. 11 spp., 7 only Mexico and Central America, Venezuela, Ecuador and Brazil one endemic each, and P. glandulosus (Nees) Ariza in Brazil and Cono Sur.

3. SUBFAMILY THUNBERGIOIDEAE (5/165–170) ‣ outsiders Thunbergia (c 100; tropical and subtropical regions in the Old World), Meyenia (1; India, Sri Lanka), Pseudocalyx (5; tropical Africa, Madagascar), Anomacanthus (1; Congo, Angola).

42. Mendoncia Vell. ex. Vend. Herbaceous or suffrutescent twining vines; stems articulated when young; flowers solitary or clustered in leaf axils, each pedunculate and subtended by 2 large, corolla sympetalous, hypocrateriform, not inflated above, contorted, whitish, greenish, or reddish, often with purplish markings within, tube cylindric to funnelform; fruit drupaceous, ovoid to ellipsoid, the mesocarp fleshy, the endocarp osseous; seeds 1-2. 76 from S Mexico to S Brazil and Bolivia, 72 in South America (38 in Colombia), 18 in Brazil (7 endemics), a unknown number of species also in tropical Africa and Madagascar; due to the superficial similarity of many of the species and the fact that lianas and vines are generally poorly known and collected in South America, the taxonomy of the genus is rather difficult; 4 endemics, in Acre, Amazonas and Bahia states, all rare plants in Brazil, by Plantas Raras do Brasil’s book.

4. SUBFAMILY AVICENNIOIDEAE (2/8) ‣ outsider Avicennia (5; West Indo-Pacific).

43. Hilairanthus Tiegh. (off Avicennia) Trees or shrubs, mangroves with pneumatophore roots; wood with anomalous secondary lateral growth from successive cambia. 3 species, on the eastern Paciifc coast from western Mexico to northwestern Peru (Piura) and the Galapagos Islands, and on the Atlantic coast from southern North America (southeastern U.S.A. and Mexico), Bermuda, West Indies, and Central America to South America (to S Brazil and Uruguay) - and in tropical Western Africa: H. bicolor (Standl.) Cornejo from E tropical Pacific from Mexico to Colombia (high tolerance to hypersaline conditions); H. schaueriana (Stapf & Leechm. ex Moldenke) Cornejo, a small tree, 15-20 m in sized; found from the lower Lesser Antilles and from Venezuela and Guyana south to Brazil (absent from French Guiana; Brazil contains over 90% of the estimated areal extent of this species); and H. germinans (L.) Cornejo, from tropical coasts of North and South America ranging from southern Florida and Bermuda, 32°20' to Atafona, Brazil, 21°37' and all Caribbean Islands (status on Anguilla is unknown); and from Puerto Lobos, Mexico south to Piura River, Peru including the Galapagos Islands, Cocos and Malpelo Islands; this species also noted from West Africa.

VERBENACEAE

§ FULLY CHLOROTROPHIC

Genera/spp. 31/1,200 Distribution SE North America, Mexico, Caribbean, S Brazil to Argentina, Atlantic islands, Europe, northern and NE Africa, SW and C Asia, Taiwan in China, Korean Peninsula, Japan. Habit herbs, shrubs or small trees, sometimes lianas, stems terete or quadrangular with varied indumentum, unarmed, or sometimes with prickles or spines; eglandular or with resinous glands on stems, leaves, bracts, calyx or corolla. Usually bisexual (some spp. of Citharexylum dioecious), evergreen or deciduous trees, shrubs or lianas, perennial or annual herbs (Pitraea tuberous perennial). Stem and branches often quadrangular in cross-section. Often aromatic.

Verbena, which has its greatest spp. diversity in the New World, also has spp. in Europe, Asia and North Africa. The closely-allied Glandularia has a disjunct North-South American distribution; Lantana has species in Africa, as do Lippia and Priva. The main pattern for the rest of the genera seems to be temperate and tropical America, as found in, for example, Tamonea which occurs from Mexico and the Caribbean to northern South America, Brazil and E Bolivia. Some genera are confined to the southern part of South America, i.e. Salimenaea, and Lampaya, all of which are restricted to Argentina and Chile. Apart from a few spp. of some of the largest genera (Lantana, Lippia, Priva, and Verbena) and a few small genera endemic to Africa and the Indian Ocean rim (e.g., Chascanum and Coelocarpum), Verbenaceae are New World in distribution.

Verbenaceae include forest trees, shrubs, lianas, and herbs and occur in open and forested, xeric and mesic habitats. In many arid habitats in portions of Argentina, some spp. of Junellia and Salimenaea play roles as community dominants, and in some cloud forests of Andean southern Peru, spp. of Citharexylum also may share dominance. Petrea contributes to the rich liana flora of wet Neotropical forests. Glandularia, Lippia, Lantana, and Verbena are common elements in disturbed sites in many habitats in Latin America, and a few spp., most notably Lantana camara, have become widely distributed weeds.

Key differences from similar families superficially resembling the Labiatae, the Verbenaceae can be distinguished from it by:

Racemose inflorescence.

Included stamens.

Thickened stigma lobes with conspicuous stigmatic tissue.

Colporate pollen.

SYSTEMATIC all nine lineages occur in South America.

UNPLACED GENUS

1. Dipyrena Hook. Two spp. from Argentina and Chile.

1. TRIBE PETREEAE (2/12) ‣ a single genus.

2. Petrea L. Lianas with fleshy drupaceous fruits consisting of two pyrenes derived from a unicarpellate ovary (one carpel having aborted) and large, showy calyces that exceed the corollas. 15 spp. of lianas from South America, 3 up to Mexico and Central America, 9 in Brazil, three Amazon Brazilian endemics, inc. P. brevicalyx Ducke from Amazonas state, a rare plant in Brazil, by Plantas Raras do Brasil’s book.

2. TRIBE DURANTEAE (5/185–200) ‣ outsider Chascanum (27–32; Africa, Madagascar, Arabian Peninsula to western India).

3. Bouchea Cham. 14 spp. in Texas to Argentina and Caribbean; 9 in South America, 4 in Brazil, three endemics (two only Minas Gerais, one without label place), all are rare plants in Brazil, by Plantas Raras do Brasil’s book.

4. Duranta L. Erect or subscandent shrub or small tree, often spinose, inflorescence an elongate lax spike. 31 spp. in Texas to Argentina and Caribbean, 25 in South America, two in Brazil, only one endemic.

5. Recordia Moldenke (inc. Verebenoxylum). Trees or shrubs, non-aromatic; leaves simple, sub-serrate to serrate; inflorescence terminal or axillary, spicate to spike; corolla white. Two spp., R. boliviana Moldenke, endemic to Bolivia, and R. reitzii (Moldenke) V. Thode & N. O'Leary, a small tree endemic to coastal mountains Atlantic Forest of S Brazil, in Santa Catarina and Rio Grande do Sul states.

6. Stachytarpheta Vahl. Herbs or sub-shrubs, terminal, spicate inflorescence. 125 spp., all in New World but including some weeds in tropical Africa (Stachytarpheta is probably restricted in its pre-Columbian distribution to the New World), 109 in South America, 94 in Brazil, 85 endemics, 24 of then are rare plants in Brazil, by Plantas Raras do Brasil’s book.

3. TRIBE CASSELIEAE (3/14) ‣ all genera occur in South America.

7. Casselia Nees & Mart. Perennial herbs, shrubs or low bushes up to 80 cm tall, stems erect, sometimes with xylopodium; inflorescences bracteose, arranged in an homotetic pleiobotryum; flowers 5-merous, calyx campanulated; corolla violet, blue, rose, pink, lilac, or white colored, funnelform or hypocrateriform, slightly zygomorphic. 7 spp., all in Brazil, two up to Paraguay and Bolivia, from savannas of C Brazil (cerrado) near Amazon ecotone, some also in the Atlantic Forest biogeographic province.

8. Parodianthus Tronc. Two spp., endemics to dry temperate regions C & southern of Argentina.

9. Tamonea Aubl. 6 spp., two in Caribbean, one only in Mexico, T. curassavica (L.) Pers. and T. spicata Aubl. widely distributed in tropical America, and T. juncea Schauer endemic to Brazil.

4. TRIBE CITHAREXYLEAE (2/103) ‣ outsider Rehdera (2; Mexico, Central America).

10. Citharexylum L. (inc. Baillonia) Trees and shrubs, sometimes with spines, inflorescence long-spicate, many-flowered, fruit often brightly-coloured, sitting in a cupped calyx. 56-67 spp. in over New World, including Caribbean, 28 in South America, 7 spp. in Brazil, none endemics. Three subgenera:

§ subg. Citharexylum ‣ three sections.

§ sect. Citharexylum ‣ 25-30 spp., mainly Mesoamerica, two extend into the Caribbean and South America and four are exclusive from South America, from Colombia to French Guiana; over section absent in Brazil.

§ sect. Mexicanum ‣ 4 spp., Mexico, one up to U.S.A.

§ sect. Pluriflorum ‣ 2-3 spp., Mexico to Nicaragua.

§ subg. Purpuratum ‣ a single sp., C. altamiranum Greenm, endemic to Mexico.

§ subg. Sudamericanum ‣ three sections.

§ sect. Andinium ‣ 11 spp. (plus one subspp., 12 taxa), central Andes in Peru, Bolivia, and Argentina, two up to in Colombia and Ecuador, mainly in arid to semi-arid inter-Andean valleys and high-elevation Andean grasslands, with C. montevidense reaching intolowlands of NE Argentina, S Brazil, Paraguay, and Uruguay.

§ sect. Caribe ‣ 2-7 spp., endemic to Caribbean.

§ sect. Sylvaticum ‣ 11 spp. (plus one subspp., 12 taxa, identically Andinium), throughout South America, typically in mesic forests and wet areas, from mid-elevation cloud forests in the northern Andes to moist lowlands surrounding the Amazon as well as those in southeastern Brazil (6), Paraguay, and Argentina; C. amabilis (Bocq.) Christenh. & Bing. (former Baillonia) is only species of genus to occur in the savannas of C Brazil (cerrado), also in Paraguay.

5. TRIBE PRIVEAE (1–2/c 20) ‣ both genera occur in South America.

11. Pitraea Turcz. Perennial herb, tuber-bearing (unique in Verbenaceae). Only one sp., P. cuneato-ovata (Cav.) Caro, in Peru and Bolivia to Argentina and Chile; it is frequently found in disturbed habitats and is a weed in cultivated fields in parts of South America.

12. Priva Adans. Small herb, not aromatic, inflorescence a lax spike. 23 spp., 11 in southern Africa to Myanmar and 12 in New World, 5 in South America, only three in Brazil, P. lappulacea (L.) Pers. widely distributed and two endemics.

6. TRIBE RHAPHITHAMNEAE (1/2) ‣ a single genus.

13. Rhaphithamnus Miers. Spinescent shrubs with bicarpellate ovaries that develop into a drupaceous fruit, and flowers with long straight corolla tubes, arranged into one- to five-flowered axillary racemes. Two spp., R. spinosus (Juss.) Moldenke in the Valdivian forests of Chile and Argentina and R. venustus (Phil.) B.L. Rob. on the Juan Fernandez islands.

7. TRIBE NEOSPARTONEAE (3/6) ‣ all genera occur in South America.

14. Diostea Miers. Ephedroid plants (many-branched shrubs with cylindrical, striate stems). Only one sp., D. juncea (Gillies & Hook.) Miers, from Chile and W Argentina.

15. Lampayo Philippi ex Murillo. Low spreading shrub with thick fleshy leaves, sometimes ephedroid. 3 spp. in Argentina, Bolivia and Chile.

16. Neosparton Griseb. Shrubs glabrous, non-aromatic, often ephedroid; stems cylindrical-striate. 3 spp. fom pampas of Argentina to Chile.

8. TRIBE LANTANEAE (11/c 275) ‣ outsiders Borroughsia (2; North America), Coelocarpum (7; Somalia, Madagascar, Socotra); Diphyllocalyx (7; Cuba), Isidroa (1; Hispaniola), Nashia (1; Puerto Rico, St Croix in the Virgin Islands).

17. Aloysia Palau. 36 spp., 6 spp. from N Mexico and U.S.A., disjunct 30 spp. in South America, in Venezuela to Chile, Brazil (9, 5 endemics), Argentina and Uruguay.

18. Lantana L. Aromatic shrubs, large, sometimes with xylopodium, with head-like inflorescences, usually multicoloured, fruit somewhat fleshy. 114 spp., 105 from southern U.S.A. to southern South America (68), 11 in Africa to Thailand; 34 spp. in Brazil, 11 endemics, two of then are rare plants in Brazil, by Plantas Raras do Brasil’s book, in Bahia and Rio de Janeiro states.

19. Lippia L. (exc. Salimenaea) Smaller aromatic shrubs, sometimes with xylopodium, fruit dry. 170 spp., 24 in Africa, and 146 from southern U.S.A. to southern South America (119), 85 in Brazil, 58 endemics; 18 spp. are rare plants in Brazil, by Plantas Raras do Brasil’s book, in several states, mainly in savannas.

20. Phyla Lour. Procumbent, creeping herb, indumentum of medifixed hairs. 4 spp., two restricted of North America and Mexico and two widely distributed in almost all regions of New World.

21. Salimenaea N. O’Leary & P. Moroni (off Lippia). Only one sp., S. integrifolia (Griseb.) N. O’Leary & P. Moroni, SW Bolivia, NW Argentina, NE Chile.

22. Troncosoa N. O’Leary & P. Moroni (ex Acantholippia). Only one sp., T. seriphioides (A. Gray) N. O’Leary & P. Moroni, endemic to Argentina.

9. TRIBE VERBENEAE (5/360–410) ‣ all genera occur in South America.

23. Glandularia J. F. Gmel. Herbs or suffruticose plants, erect, prostrate or decumbent, mostly with ascending floral branches; leaves with margins entire, crenate, serrate, dentate or lobed. 92 spp., 26 from U.S.A. to Honduras and 66 from Ecuador to Chile, east up to Brazil (33, highly centered in Rio Grande do Sul state, 11 endemics) and Uruguay.

24. Hierobotana Briq. Woody herb, non aromatic; procubent, leaves 3-partited. Only one sp., H. inflata (Kunth) Briq., endemic to Ecuador.

25. Junellia Modenke (inc. Urbania). Shrubs, sometimes low, cushion-forming, non aromatic; sometimes spiny; J. spathulata is a ephedroid habit. 39 spp., J. trifida (Kunth) P. Peralta & N. O'Leary endemic to Mexico, 38 remaining from Peru to Falkland Is, 27 restricted of Cono Sur, in mountain environments.

26. Mulguraea N.OLeary & P.Peralta. 11 spp. in Cono Sur, 3 up to Peru and M. asparagoides (Gillies & Hook.) N. O'Leary & P. Peralta up to Bolivia.

27. Verbena L. 57 spp., cosmopolitan, the greatest diversity in New World (54, from Canada to Tierra del Fuego – 26 in South America), and 3 in Europe, Asia and North Africa (V. supina L., V. officinalis L.), and V. dalloniana Quézel is endemic to Chad; 16 spp. in Brazil, three endemics.

LENTIBULARIACEAE

§ CARNIVOROUS (Brocchnia - Catopsis - Paepalanthus - Drosera - Heliamphora - Philcoxia - Genlisea - Utricularia - Pinguincula)

Genera/species 3/370 Distribution cosmopolitan except polar and arid regions. Habit bisexual, perennial or annual herbs. Roots present (Pinguicula) or absent (Genlisea, Utricularia). Mycorrhiza absent. Aquatic or helophytic; some species are epiphytic. Carnivorous. Stem in Genlisea and Utricularia photosynthesizing. Leaves in Utricularia with bladder-shaped suction traps (in terrestrial species with chemical attractants), in Genlisea modified into ‘eel trap’-like structures; tubers or rhizomes present in many terrestrial species of Utricularia.

Plant herbaceous annual or perennial; common in wet places of marsh or wetlands; aquatic, terrestrial or sometimes epiphytic, lacking roots, fixed in moist substrate or free-floating, vascular system often reduced, mostly carnivorous. Stem short; hairs sessile to stalked, glandular-headed, some secreting mucilage and others digestive enzymes. Leaves alternate or sometimes whorled, often in basal rosettes (Pinguicula, Genlisea), entire to much divided, heterophyllous in different periods of the development of the plant, many Pinguicula and other spp. (especially sub-tropical species) depend on seasonal climate changes for the regular development of winter rosettes, the leaves of which differ from the summer rosettes. Leaves always highly modified, flat and densely covered with sticky, mucilage-secreting and digestive hairs, and with margins rolling inward in response to contact of glandular hair with prey organism (Pinguicula), or tubular and spiralled, with downward-pointing and digestive hairs and a basal chamber (Genlisea), often dimorphic in aquatic plants (Utricularia L.), with finely divided submerged leaves and traps of complex structure; In winter, subtropical spp. of Utricularia and Pinguicula develop special organs in axes of branches (hibernacula), air leaves forming floating rosettes around floral peduncle (floats), reduced to scales, or absent in some terrestrial forms, (dimorphism is represented by rosettes of leaves and for insectivorous leaves in ascidio form (traps) (Utricularia) or not obviously foliaceous, highly dissected, bearing prey-catching bladders, each with 2 sensitive valves forming a trapdoor entrance, which opens inward in response to a stimulus conveyed by 4 sensory hairs and then immediately closes again, and lined on inside with branched digestive hairs (Utricularia), stipules lacking or tubular and inserted within the soil substrate (Genlisea), in Pinguicula leaves without special structures for the capture of prey, surface is covered with sessile or peduncular glands.

SYSTEMATIC all genera occur in South America; key to genera of Neotropical Lentibulariaceae:

1. Leaves divided into linear segments, rarely in rosette, sepals with two lobes ------------ Utricularia

1. Leaves in basal rosette; sepals with 5 lobes - 2

2 Lobes of the calyx laciniate; unequal ------------ Genlisea

2. Lobes of the calyx deeply 2-lipped ------------ Pinguicula

1. Genlisea A.St.-Hil. Annual or perennial herbs, stems short, erect, leaves dimorphic in a basal rosette, roots absent, calyx with 5 lobes, nectar-spur flowers. 31 spp., occurring in sub-Saharan Africa (10 in mainland, 1 in Madagascar), 20 in tropical and subtropical America; the Central Brazilian highlands represent the main diversity center of the genus, where both subgenera occur; in this region, most Genlisea species are endemic to the rocky grasslands (campos rupestres) vegetation type; two subgenera:

§ subg. Genlisea ‣ 21 spp., 11 spp. of Africa and Madagascar, 2 endemics to Brazil, two endemics to Venezuela, 4 in Brazil and countries of Guiana Shiled, G. repens Benj. in over tropical South America, and one widely distributed, G. filiformis A. St.-Hil. from Bolivia, Brazil, Colombia, Guianas to Mexico, also in Cuba.

§ subg. Tayloria ‣ 10 spp., endemic to center Brazil; G. lobata Fromm-Trinta is a rare plant in Brazil, by Plantas Raras do Brasil’s book.

G. tuberosa Rivadavia, Gonella & A. Fleischm. has the smallest angiosperm genome known at around 61 Mbp. G. margaretae Hutch. (Tanzania, Zambia, Madagascar) as having the second smallest known angiosperm genome size at 63.4 Mbp, just 0.2 Mbp lower than that of G. aurea A. St.-Hill, the third; threse species is also endemic to Brazil, from the states of Mato Grosso in the west to NE Bahia and down to Santa Catarina in the southeast; it typically grows on sandstone highlands at altitudes 550 m - 2550 m.

2. Pinguicula L. Annual or perennial herbs, terrestrial or rarely epiphytic; true roots present; winter and summer leaves different; insects are caught and digested by sessile and short-stalked mucilage-producing glands on the upper side of rosette leaves (‘flypaper traps’), nectar-spur flowers. 122 spp., Europe, northern Asia, Hymalaias, Japan, North America, Central America, highly centered in Mexico (47, 43 endemics), Caribbean (12 in Cuba (10 endemics) and 1 in Dominic Republic), and 12 in South America, falling in two different clades, both continental restricts:

§ subg. Isoloba ‣ 21 spp., four sections, only sect. Ampullipalatum in South America with 11 spp. all restricteds from continent, four from Venezuela, Colombia, Ecuador, Peru and Bolivia, and five simultaneously in Argentina and Chile.

§ subg. Pinguincula ‣ 25-30 spp., northern hemisphere absent in New World.

§ subg. Temnoceras ‣ 63-73 spp., four sections, only one in South America, sect. Heterophylliformis, a monotypic section, with P. elongata Benj. from Andes of Colombia and Venezuela.

3. Utricularia L. Annual or perennial herbs, terrestrial, epiphytic or aquatic; roots absent; root-like stems; vegetative parts not diferenciable; rhizomes or stolons often present; calyx with 2 lobes, nectar-spur flowers. 281 spp., the largest genus of carnivorous plants, that occur throughout the world, 5 in New World, highly centered in South America (93), mainly Brazil (78, 25 endemics), Venezuela, Guyana, W Australia (55) and India (33), with many local diversity in Brazil, e.g. Diamantina Range (20, 4 endemic) and swamps of center Roraima state; two spp., from Pará and Mato Grosso state, are rare plants in Brazil, by Plantas Raras do Brasil’s book.

57. VAHLIALES

VAHLIALES DOES NOT OCCUR IN SOUTH AMERICA, AND IS COMPOSED OF A SINGLE FAMILY, VAHLIACEAE (1/5).

58. SOLANALES

FAMILIES ABSENTS IN SOUTH AMERICA: MONTINIACEAE (3/5) AND SPHENOCLEACEAE (1/2).

HYDROLEACEAE

§ FULLY CHLOROTROPHIC

Genera/spp. 1/14 Distribution almost pantropical: the SE U.S.A., Mexico, Central America, Caribbean, tropical South America to Argentina, tropical West and Central Africa, tropical Asia, northern Australia. Habit bisexual, annual or perennial herbs or shrubs, sometimes with axillary-sublateral spines. Helophytic. The family Hydroleaceae consists of one genus, Hydrolea.

SYSTEMATIC a single genus.

1. Hydrolea L. Erect, often branching, sometimes becoming decumbent, herbaceous to partly woody, usually glandular-pubescent, sometimes spiny in leaf -axils, annuals or perennials from taproots, usually in wet soil; leaves alternate, entire, elliptical-lanceolate to obovate; flowers usually few in lateral and /or terminal cymes; corolla usually blue, sometimes white, broadly campanulate; capsule globose to nearly elliptic, dehiscing irregularly, containing numerous, minute seeds. 11 spp. of aquatic plants, subcosmopolitan; two sections:

§ sect. Hydrolea ‣ 7 spp., New World: H. spinosa L. occur over New World with three subespecies; H. elatior Schott are widely distributed in South America, with disjunct populations in Honduras and Mexico; H. nigricaulis C. Wright ex Griseb. occur only in Cuba and Jamaica; and four endemics to U.S.A.

§ sect. Attaleria ‣ 4 spp., three only in tropical Africa, and one from Africa up to tropical Asia.

CONVOLVULACEAE

§ PARASITIC (Prosopanche – Cassytha - ... – APODANTHACEAE – SANTALALES – Lennoa – OROBANCHACEAE - Cuscuta)

Genera /spp. 57/1,590-1,610 Distribution cosmopolitan except polar areas, with their largest diversity in subtropical regions in Asia and America. Habit usually bisexual (in Hildebrandtia dioecious), usually climbing and winding perennial or annual herbs (rarely evergreen trees, shrubs or lianas). Some spp. are xerophytic.

Members of the Cuscuta (also known as Dodder and Devil's Guts) are regarded as some of the worst weeds in the world. Attaching to their agricultural hosts through haustoria and twining around their victims they steal light and nutrients from valuable crops. Sweet potato (Ipomoea batatas L.) is world’s third most important root crop, and it is found only in cultivation.

Key differences from similar families - Solanaceae similar, but has:

ü Synsepalous calyx.

ü Sometimes zygomorphic corollas.

SYSTEMATIC two subfamilies, Humbertioideae (1/1; Madagascar) does not occur in South America; among Convolvuloideae, tribes Cardiochlamyeae (7/24, Madagascar, India, Himalayas to Burma, S China and SE Asia, Malesia, Mexico, Australia) and Erycibeae (1/67; tropical Asia to Japan and tropical Australia) do not occur in South America.

1.1 COLVOLVULOIDEAE ▸ TRIBE DICHONDREAE (8/58) - outsiders Falkia (3; Africa), Nephrophyllum (1; Ethiopia), Petrogenia (1; N Mexico), Metaporana (5; E Africa, Madagascar, Socotra), Dipteropeltis (3; tropical W and Central Africa), Rapona (1; Madagascar).

1. Calycobolus Willd. ex Roem. & Schult. Lianas with stems woody, glabrous or pubescent. 25 spp., two in Mexico, three spp. in South America (Colombia, Venezuela, Guyana, Peru, Bolivia, Brazil - all three, one endemic), remaining in Africa.

2. Dichondra Forst. & Forst. f. Creeping or crawling perennial herbs, often rooting at nodes. 17 spp., 14 spp. from U.S.A., Mexico, Mesoamerica, Antilles, over South America (7, three in Brazil, one endemic), and 1 in Australia and two in New Zealand.

1.2 COLVOLVULOIDEAE ▸ TRIBE CRESSEAE (11/210) - outsiders Petrogenia (1, U.S.A., Mexico), Hildebrandtia (13; Africa, Madagascar, Arabian Peninsula), Seddera (c 20; tropical and subtropical regions in Africa, Madagascar, Arabian Peninsula), Stylisma (8; U.S.A.), Wilsonia (3; Australia, Tasmania), Itzaea (1; Central America), Neuropeltis (18; tropical Africa, tropical Asia), Neuropeltopsis (1; Borneo).

3. Bonamia Thouars. Herbs, vines, erect subshrubs; style divided into two filiform arms, and the globose to reniform, relatively small stigmas; the sepals are not accrescent and the cotyledons are ovate-oblong to shallowly emarginate. 64 spp., pantropical genus; 33 spp. in New World, 26 in South America, 18 in Brazil, 11 endemics; two spp. in Mato Grosso do Sul and Mato Grosso states are rare plants in Brazil, by Plantas Raras do Brasil’s book.

4. Cressa L. Perenial shrubs and herbs. 4 spp., two in New World, C. nudicaulis Griseb. from North America and Cono Sur, and C. truxillensis Kunth from North America, Mexico, Central America, Ecuador to Chile, Paraguay, Uruguay and Rio Grande do Sul state in S Brazil; Old Worlds, mainly salt pans and coastal marshes.

5. Evolvulus L. Herbs or shrubs, annual or perennial, non-laticiferous; stems prostate, ascending or erect. 107 spp. in New World, from U.S.A. south to Brazil and Bolivia and Caribbean, two species, E. alsinoides (L.) L. and E. nummularius (L.) L., also in the Old World tropics; 92 spp. in South America, 73 in Brazil, 50 endemics, two of then, from Goiás and Bahia states, are rare plants in Brazil, by Plantas Raras do Brasil’s book.

1.3 COLVOLVULOIDEAE ▸ TRIBE MARIPEAE (3/36) - all genera in South America.

6. Dicranostyles Benth. Lianas reaching over 30 m; larger stems smooth to slightly fluted, 15 cm in diameter or larger. 16 spp., N South America, French Guiana to Peru and N Brazil (13, 5 endemics, two of then, from Amazonas state are rare plants in Brazil, by Plantas Raras do Brasil’s book), centered in Guiana Shield, D. ampla Ducke up to Central America.

7. Lysiostyles Benth. Forests liana reaching 30 m; stems up to 4 cm in diameter. Only one sp., L. scandens Benth., endemic to the Guiana Shield of French Guiana to Venezuela and Amazonas state in N Brazil, 200–1,300 m elevation range.

8. Maripa Aubl. Lianas reaching over 30 m; larger stems fluted, to 30 cm in diameter, younger stems often angled. 20 spp., 19 over N South America, French Guiana to Peru and N Brazil (12, none endemics) up to Mesoamerica, and one sp. restricted from Mexico to Nicaragua.

1.4 COLVOLVULOIDEAE ▸ TRIBE JACQUEMONTIEAE (1/120) - a single genus.

9. Jacquemontia Choisy. Perennial or annual vines, with climbing, decumbente, prostate, rarely suffrutescent stems (erect herbs), flowers never yellow. 123 spp., mostly New World (104 spp., U.S.A., over Neotropics, 84 in South America) with a few spp. occurring in tropical Africa, Asia (only one endemic) and Australia; 66 spp. in Brazil, 39 endemics, one in Minas Gerais is a rare plant in Brazil, by Plantas Raras do Brasil’s book.

1.5 COLVOLVULOIDEAE ▸ TRIBE CUSCUTEAE (1/220) - a single genus.

10. Cuscuta L. Herbs chlorophyll-lacking and parastic, has a withering terrestrial root system which is short-lived, and no internal phloem; leaves reduced to minute scales, roots absent and plants attached to the host by numerous small haustoria. 219 spp. in World; 140 spp. in New World, 65 in South America, over distribuited; 22 in Brazil, 9 endemics (C. globosa Ridl. from Fernando de Noronha Is. in Atlantic Ocean is a rare plant in Brazil, by Plantas Raras do Brasil’s book; however, none Cuscuta is endemic to islands); growing in mesophytic habitats - particularly along streams and in areas associated with anthropogenic ecosystems; occasionally found in halophytic areas.

1.6 COLVOLVULOIDEAE ▸ TRIBE ANISEAE (3/6) - all genera in South America.

11. Aniseia Choisy. (inc. Iseia) Vines; stems herbaceous, mostly trailing on ground or over other plants, becoming somewhat woody near base; leaves usually with short petioles, blades linear to ovate or elliptical; inflorescences mostly solitary, less often cymose; flowers axillary, with no detectable odor; corollas white, campanulate; fruits capsular, globose to ovoid, 2-celled, 4-val- vate, brown. Three spp., widely distributed of tropics, all in Brazil, with A. argentina (N.E.Br.) O'Donell restricted from S Brazil and Argentina.

12. Odonellia K.R.Robertson. Lianas or voluble herbs. Two spp., O. hirtiflora (M. Martens & Galeotti) K.R. Robertson from Mexico through Mesoamerica, Colombia, Venezuela, Ecuador, Peru and Brazil, another, O. eriocephala (Moric.) K.R.Robertson, restricted of Brazil.

13. Tetralocularia ODonell. Only one sp., T. pennellii O'Donell, from Colombia, N Brazil, Bolivia and French Guiana.

1.7 COLVOLVULOIDEAE ▸ TRIBE CONVOLVULEAE (1/200) - a single genus.

14. Convolvulus L. Herbs up shrubs. 200 spp., mainly temperate, centres of diversity are to de found in Mediterranean, E Africa, Central Asia and Arabia, S Africa, Australasia, 17 in New World, North America to Mexico, and Galapagos to S Brazil (15 in South America, 7 spp. in Brazil, one endemic) and Chile.

1.8 COLVOLVULOIDEAE ▸ TRIBE IPOMOEAE (c. 24/c. 870) - outsiders Polymeria (10; E Malesia to Australia and New Caledonia); Remirema (1; Thailand), Hyalocystis (2; tropical Africa), Hewittia (1; tropical Africa, Madagascar, tropical Asia to New Guinea), Xenostegia (5; tropical Africa, Madagascar, tropical Asia to tropical Australia and Solomon Islands), Decalobanthus (c 13; SE Asia, Malesia to islands in the Pacific, tropical E Africa and Madagascar), Merremia (10–20; S, E and SE Asia, Malesia to New Guinea and N Australia), Stictocardia (12; tropical regions in the Old World), Lepistemon (10; tropical regions in the Old World), Argyreia (90–95; tropical Asia to Australia), Astripomoea (12; tropical and S Africa), Blinkworthia (2; Burma, S China), Lepistemonopsis (1; tropical E Africa), Paralepistemon (1; S tropical Africa).

15. Calystegia R.Br. 25 sp., over Old World, 23 spp. in New World, from Canada and U.S.A., only three in South America, two scattered in Ecuador to Cono Sur (one also in Mexico), and C. brummittii P.P.A. Ferreira & Sim.-Bianch. endemic to Araucaria forests in Santa Catarina and Rio Grande do Sul states in S Brazil.

16. Camonea Raf. Herbaceous twiners or prostrate creepers; leaves entire or angulate-lobed; with two firm outgrowths (paired auricles) at petiole base (absent in C. vitifolia); corolla with a tuft of hairs at the apex of the mid-petaline bands, otherwise glabrous; anthers longitudinally dehiscing and curved at the apex or spirally dehiscing. 5 spp., centred in tropical Asia with C. umbellata (L.) Simões & Staples widely distributed in tropical America (almost all countries) and Africa.

17. Daustinia Buril & Simões. Vines with yellow flowers. Only one sp., D. montana (Moric.) Buril & A.R. Simões, endemic to E Brazil.

18. Distimake Raf. (inc. Merremia p.p.) Robust herbaceous climbers (rarely lianas or erect shrubs, sometimes with xylopodium); leaves usually five- to seven-palmately lobed or compound (rarely simple or reduced to scales); calyx mostly with flat sepals (not convex) appressed to the corolla tube base, accrescent in fruit; corolla often white or pale yellowish, with or without a dark red centre, entirely glabrous, drying with dark lines in mid-petaline bands. 35 spp., widely distributed in tropical America, with 34 spp. in New World, 22 in South America, 15 in Brazil, 7 endemics, D. repens (D.F. Austin & Staples) Petrongari & Sim.-Bianch. from Minas Gerais state is a rare plant in Brazil, by Plantas Raras do Brasil’s book) and tropical Africa with disjunct species in Asia and N Australia.

19. Ipomoea L. (inc. Turbina) Climbing herbs or lianas, trees, shrubs or erect herbs, perennial or less often annuals, usually with milky sap, glabrous or pubescent, sometimes with xylopodium. c. 500 spp., comopolitan; 428 spp. in New World, S U.S.A. to Chile, Uruguay, and Caribbean, 263 in South America, 177 in Brazil, 70 endemics (6 of then are rare plants in Brazil, by Plantas Raras do Brasil’s book); absent in continental Chile, but present in Isla de Pascua; 6 unplaced species and six well defined clades:

Ipomoea includes six true trees (up to 15m tall in a mexican endemic), all from Mexico, I. wolcottiana Rose, Gard. & Forest and I. pauciflora M. Martens & Galeotti up to Peru in South America.

§ CLADE A ‣ 277 spp.

§ CLADE B ‣ 105 spp.; comprises species mostly from Mexico and surrounding countries although it includes quite a few South American species.

§ CLADE C ‣ comprise a morphologically heterogeneous group of American (40) and Australasian species, which contains a number of small, well-supported and morphologically distinct clades.

§ CLADE D ‣ comprises a small clade of entirely American species (8); all species are herbaceous but show no other obvious common character.

§ CLADE E ‣ consists of 4 species, of which only one is certainly of New World origin (I. habeliana Oliv.); all other species in the clade are either African or of uncertain origin suggesting the clade is essentially African with I. habeliana having evolved from I. violacea L. in the Galapagos Islands.

§ CLADE F ‣ this is the large, essentially Old World Clade (OWC), containing 26 naturally occurring New World species as well as several Old World species which are ancient or recent introductions to the New World.

20. Operculina Silva Manso. Herbaceous or woody climbers. 18 spp., tropical worldwide, including 10 spp. in Mexico through Mesoamerica, south to Brazil (7 in South America, 5 spp. in Brazil, one endemic), and Caribbean.

SOLANACEAE

§ FULLY CHLOROTROPHIC

Genera /spp. 101/c. 2,650 Distribution tropical, subtropical and temperate regions in the Northern and Southern Hemispheres, with their highest diversity in South America. Habit usually bisexual (sometimes monoecious, andromonoecious or dioecious), evergreen trees, shrubs or lianas, perennial, biennial or annual herbs. Some spp. are xerophytes. Some spp. are succulent. Often evil-smelling (foetid).

Several genera of Solanaceae are large, epiphytic shrubs - Juanulloa has tubular usually orange flowers; Markea has flowers that are white or greenish (except Markea coccinea Rich.of the Amazon with red flowers) with large spreading lobes and berries with tiny, rectangular seeds; Trianaea, Merinthopodium and Dyssochroma are all bat-pollinated and have large, tough greenish corollas, often on long, pendant peduncles. Few spp. are cultivated in New World; these include Solanum melongena L. (eggplant or aubergine) from Asia and Solanum aethiopicum L. (scarlet eggplant) from Africa. Many Neotropical taxa are widely cultivated for food (Capsicum pepper, picant fruits, one known as pimentão), Solanum (S. tuberosum L. is a important crop cultivated in world; native ancient in Andes, mainly by Incas; S. lycopersicum L. is tomato, common in Brazil and Italia; edible fruits), medicine (Nicotiana, Brunfelsia) or as ornamentals (Petunia, Streptosolen). Solanum is one of the largest angiosperm genera, with approximately 1,500 species distributed worldwide. The genus includes important economic plants such as the tomato, potato, and eggplant, as well as a number of lesser-known cultivated species such as the pepino (S. muricatum Aiton), naranjilla (S. quitoense Lam.), cocona (S. sessiliflorum Dunal), and tree tomato (S. betaceum Cav.).

Solanaceae are little diversified in the rest of the world. For example, in Africa only a few genera are present, namely the two widely ranging genera Solanum and Lycium, three in common with Europe and Asia (Mandragora, Hyoscyamus and Withania), the extraordinary Nicotiana africana, and the endemic Discopodium. In Asia the strictly Asian Tubocapsicum, species of Solanum and Lycianthes, and species of the European genera Atropa, Mandragora, Hyoscyamus and Withania are found; finally, Nothocestrum is a geographically isolated genus on Hawaii. 2298 spp. in New World; 1767 in South America.

Except Solanoideae, diversity in Old World occcur only by Tsoala (Madagascar), Nicotianoideae (8 genera, in Namibia, Australia and New Caledonia) and Schwenckia americana D. Royen ex L. (Africa); among Solanoideae, several clades of Solanum, one clade of Lycium, Hyoscyameae, Mandragora, part of Withaninae (probably one event), Physalis alkekengi L., and part of Lycianthes (the phylogeny of Lycianthes is still poorly known, but a relatively small group of about 20 species exists in SE Asia and may represent a single lineage).

SYSTEMATIC nine high lineages, all in South America.

1. SUBFAMILY SCHIZANTHOIDEAE (1/12) ‣ a single genus.

1. Schizanthus Ruiz & Pav. Annual or biennial herbs; phellogen pericyclic; pericyclic fibres absent; flowers strongly zygomorphic, resupinate. 14 spp., Chile and Argentina.

2. SUBFAMILY DUCKEODENDROIDEAE (1/1) ‣ a single genus.

2. Duckeodendron Kuhlm. Trees ca. 30 m high, the tallest member of Solanaceae; leaves entire, coriaceous, pubescent abaxially; petiole canaliculate, with dense ochre indumentum; inflorescence terminal, cymose, ca. 15-flowered, brownish-pubescent; flowers subtended by one tiny pilose bract, pedicels articulated; calyx aestivation quincuncial, lobes broad, as long as tube; corolla greenish, tubular, slightly infundibuliform, pubescent inside, lobes rounded, aestivation quincuncial; drupe 1-seeded, seed ovoid. Only one sp., D. cestroides Kuhlm., endemic to the Central Amazon rainforest in Amazonas and Pará state in Brazil.

3. SUBFAMILY GOETZEOIDEAE (6/7) ‣ outsiders Coeloneurum (1; Hispaniola), Henoonia (1; Cuba), Espadaea (1; Cuba), Goetzea (2; Puerto Rico; Hispaniola), Tsoala (1; Madagascar, probably extinct).

3. Metternichia Mikan. Shrub to tree, many branched, simple leaves, brownish bark, with showy capanulate white flowers. Only one sp., M. princeps Miers, highly poisonous nephrotoxic for goats, endemic of coastal forests of SE Brazil and adjacent places of dry seasonal scrubland of NE Brazil (caatinga) and Atlantic sandy coastal shrublands (restingas).

4. TRIBE BENTHAMIELLEAE (3/15) – all genera in South America.

4. Benthamiella Speg. Pulvinate chamaephytes or lax cushions; stems branched, densely leafy. 12 spp., Patagonia in S Argentina and Chile.

5. Combera Sandw. Small, erect perennial herbs, often cushions; stems densely leafy, partly subterranean. Two spp. in the Andes in temperate Chile and Argentina.

6. Pantacantha Speg. Low shrubs; stems branched, densely leafy, often cushions. Only one sp., P. ameghinoi Speg., endemic to Patagonia in central Argentina.

5. SUBFAMILY CESTROIDEAE (8/c 250) - three tribes, all in South America.

5.1 CESTROIDEAE ▸ TRIBE SALPIGLOSSIDEAE (2/7) - both genera in South America.

7. Reyesia Gay. Annual or perennial herbs, dichasially branched, glutinose. 4 spp., endemic to dry areas in N Chile, one of then reaching in closed small area in adjacent in Argentina.

8. Salpiglossis Ruiz & Pav. Herbs or shrubs, densely viscid-pubescent; stems leafy or almost without leaves, sometimes xeromorphic (S. spinescens) with spine-like processes. Three spp., one in Mexico and two in Chile, one of then reaching into Argentina; S. sinuata Ruiz & Pav. is a large-flowered species from central Chile which is frequently cultivated in temperate gardens for its showy flowers.

5.2 CESTROIDEAE ▸ TRIBE BROWALLIEAE (1/17) - a single genus.

9. Browallia L. Annual herbs, inflorescence racemiform. 23 spp., B. eludens R. Van Devender & P.D. Jenkins endemic to Mexico and U.S.A., and 17 in Andes from Venezuela to Bolivia, highly centered in Peru (20, 17 endemics), two also outside this area, B. speciosa Hook. up to Central America, and B. americana L. up to Caribbean and Trinidad & Tobago.

5.3 CESTROIDEAE ▸ TRIBE CESTREAE (5/218) - all genera in South America.

10. Cestrum L. 192 spp., predominately tropical in distribution but several species range south as far as Chile and Uruguay and several range north as far as the southern U.S.A. and the Bahamas; 99 spp. in South America. Brazil (26 in Brazil, 13 endemic) is considered to have a larger species diversity, following Andean regions of Colombia to Bolivia, Peru, and the north of Argentina; two sections are recognized:

§ sect. Cestrum ‣ 181 spp., over tropical New World.

§ sect. Habrothamnus ‣ c. 8 spp. from E Mexico and ranging into Central America.

§ sect. Pseudocestrum ‣ only one sp., C. inclusum Urb., from Hispaniola.

11. Protoschwenkia Soler. Small shrubs, profusely branched. Only one sp., P. mandonii Soler., Bolivia and adjacent Mato Grosso do Sul state in C Brazil.

12. Sessea Willd. 24 spp., 21 from Andean region Venezuela to Bolivia and three in Brazil and adjacent Cono Sur, S. brasiliensis Toledo endemic.

13. Streptosolen Miers. Evergreen scabrous-pubescent shrubs, densely branched; leaves ovate, petiolate; inflorescence subcorymbose cymes; flowers pedicellate; calyx zygomorphic, tubular, 4–5-lobed; corolla infundibuliform, capsule globose to ovate; seeds 60–80, cuboidal-elongated; testa reticulate. Only one sp., S. jamesonii (Benth.) Miers, Ecuador and Peru.

14. Vestia Willd. Much branched shrubs, il-scent. Only one sp., V. foetida Hoffmanns., Chile.

6. SUBFAMILY PETUNIOIDEAE (8/c 135) - outsider Plowmania (1; Mexico, Guatemala).

15. Bouchetia Dunal. Rhizomatous or root sprouting, decumbent herbs. Basal leaves oblong-spathulate or rhombic- ovate, upper leaves narrowly elliptic; flowers solitary, axillary. Three spp., displaying a disjunct distribution, with B. anomala (Miers) Britton & Rusby in Rio Grande do Sul state in S Brazil, Paraguay, Uruguay and north to central Argentina, and two other species in North America and Mexico.

16. Brunfelsia L. Shrubs or small trees. 50 spp. in two clades., mainly in South America, especially S & C Brazil and in Antilles. Two large clades:

§ South American Clade ‣ 28 spp. from South America up to Panamá, mainly in the Amazon rainforest up Guiana Shield, E Andes and SE Brazil (20, 12 endemics), three of then, in Amazonas, Minas Gerais and Rio de Janeiro states are rare plants in Brazil, by Plantas Raras do Brasil’s book.

§ sect. Brunfelsia ‣ 23 spp. from Cuba (13, 11 endemics), Jamaica (6, all endemics), Hispaniola (4, two endemics), and Puerto Rico (5, 3 endemic), one up to Lesser Antilles.

17. Calibrachoa La Llave & Lex. Small shrubs, more rarely annual herbs, generally viscid-pubescent; stem usually with brachyblasts. 28 spp., SE and S Brazil (24, 13 endemics, 5 of then in Santa Catarina Rio Grande do Sul states are rare plants in Brazil, by Plantas Raras do Brasil’s book), Paraguay, Uruguay, and Argentina, C. parviflora (Juss.) D’Arcy disjunct between South and C Mexico.

18. Fabiana Ruiz & Pav. Chamaephytes or microphyllous shrubs, of homoblastic or heteroblastic growth, sometimes cushions; stems and leaves with dense resiniferous indumentum. 16 spp., S Peru to Argentina, especially in the Andes, reaching up to 4,900-5,000 m elevation range in F. bryoides Phil.

19. Hunzikeria D'Arcy. Small herbs up to 25 cm, usually much branched from a perennial root. 4 spp., three in SW U.S.A. and N Mexico, and H. steyermarkina D’Arcy, known only from the subdesert area of Venezuela near the Caribbean coast just west of Caracas.

20. Leptoglossis Benth. Xeromorphic herbs or shrubs, flowers are solitary or aggregated into will-defined racemes or panicles, each flower subtended by a bract. 7 spp., six endemic in coastal Peru and L. linifolia (Miers) Benth. & Hook. f. ex Griseb. in Chile and NW Argentina.

21. Nierembergia Ruiz & Pav. Small shrubs or rhizomatus stoloneferous tuber bearing prostrate from meadows and marshes. 23 spp., 20 mainly from Argentina and Chile extending into S Brazil (7, two endemics), S Bolivia, Paraguay and Uruguay, Mexico and Ecuador one endemic each, and N. repens Ruiz & Pav. from Colombia to Chile.

22. Petunia L. Annual herbs, viscid-pubescent; brachyblasts absent. 17 spp. from S and SE South America in Brazil (14, 9 endemics, 6 of then from Minas Gerais, Santa Catarina and Rio Grande do Sul states are rare plants in Brazil, by Plantas Raras do Brasil’s book), Bolivia, Uruguay, Paraguay, Argentina.

7. SUBFAMILY SCHWENCKIEAE (3/c 30) - all genera in South America.

23. Heteranthia Nees & Mart. Herb to subshrub, erect, often decumbent, 40 cm tall; stems cylindrical, uncommon small adventive roots, alterne leaves; inflorescence terminal or axillary, racemose; corolla infundibuliforme, whitish to purplish. Only one sp., H. decipiens Nees & Mart., Minas Gerais, S Bahia and adjacent Rio de Janeiro, in SE Brazil.

24. Melananthus Walp. Tiny herbs, or small shrubs. 5 spp., all in Brazil, three scattered up to Caribean, Central America and Venezuela, one up to Cono Sur, and M. fasciculatus (Benth.) Soler. endemic, as a rare plant in Brazil, by Plantas Raras do Brasil’s book, known only Espirito Santo state.

25. Schwenckia L. Annual or perennial herbs, sometimes shrubs, often with xylopodium. 22 spp. over tropical South America (21, the exception is a Caribbean endemic S. filiformis Ekman), with S. americana D. Royen ex L. also in W Africa; 16 spp. in Brazil, 6 endemics.

8. SUBFAMILY NICOTIANOIDEAE (c 8/c 105) - outsiders all in W and S Australia except Duboisia (4; E Australia, New Caledonia).

26. Nicotiana L. Herbs to small trees. 76 spp., 8 in SW North America, southern South America (43, 8 in Brazil, 4 endemics) east of the Andes, one in Namibia, 25 in Australia. 13 sections.

§ sect. Alatae ‣ 8 spp., tropical America, all in Brazil, and only section in country.

§ sect. Nicotiana ‣ only naturalized world-wide N. tabacum L., possibly native from Colombia and Peru, cultivated tobacco; a perennial herbaceous plant. It is found only in cultivation, where it is the most commonly grown of all plants in the Nicotiana genus, and its leaves are commercially grown in many countries to be processed into tobacco; it grows to heights between 1 to 2 metres.

§ sect. Noctiflorae - 6 spp., S South America.

§ sect. Paniculatae ‣ 8 spp., W South America.

§ sect. Petunioides ‣ 8 spp., U.S.A. to South America, absent in Brazil.

§ sect. Polydicliae ‣ 2 spp., U.S.A. to Mexico.

§ sect. Repandae ‣ 4 spp., U.S.A. to N Mexico.

§ sect. Rusticae ‣ only one sp., Andes.

§ sect. Suaveolentes ‣ 26 spp., Australia, New Caledonia, Namibia.

§ sect. Sylvestres ‣ only one sp., Bolivia and Argentina.

§ sect. Tomentosae ‣ 5 spp., Peru to Argentina.

§ sect. Trigonophyllae ‣ 2 spp., U.S.A. to Mexico.

§ sect. Undulatae ‣ 5 spp., Andes, Ecuador to Bolivia.

9. SUBFAMILY SOLANOIDEAE (50–55/1,700–1,800) - all clades, mainly tribes, occur in South America except Mandragoreae (1/4; Mediterranean, Central Asia, Himalayas).

9.1 SOLANOIDEAE ▸ TRIBE ATROPEAE (c. 260) - outsiders Atrichodendron (1; Vietnam); Atropa (5; N Morocco, Europe to the Caucasus, N Iran, Central Asia and Himalayas, Mongolia), Anisodus (5; Himalayas in Nepal, India and Bhutan to W China), Atropanthe (1; China), Hyoscyamus (c 17; Europe, Mediterranean, Madeira, Canary Islands, North Africa to Somalia, SW and Central Asia to China), Przewalskia (1; W China), Scopolia (2; Alps, the Carpathians, the Caucasus, Korean Peninsula, Japan), Physochlaina (6; the Caucasus to India and China).

27. Jaborosa Juss. Perennial herbs, frequently with gemmiferous roots and rhizomes. 22 spp., mainly W Andes in Peru to Chile, Bolivia to S Argentina, only two reaching SE South America, with J. integrifolia Lam. up to Rio Grande do Sul state in S Brazil and Uruguay.

28. Latua Phil. Shrubs or trees with heteroblastic growth and cauline spines, often cauliflorous. Only one sp., L. pubiflora (Griseb.) Baill., southern Chile.

29. Lycium L. (inc. Grabowskia) Woody shrubs, mainly xeromorphic, sometimes dwarf halophytic in Andes. 92 spp., warm-temperate and subtropical regions on both hemispheres, 55 in New World, South America (33, highly centered in Argentina), S Africa (ca. 20), North America (ca. 20), Eurasia (from Europe to China and Japan: ca. 10), Australia (1), and several islands in the Pacific Ocean (2), only three spp. in Brazil, none endemics.

L. boerhaviifolium L. f. has by far the widest distribution, ranging from Argentina and Chile north to Ecuador and Bolivia, S Brazil, Galapagos Islands and has been collected from several locations near Tehuacán in Puebla, Mexico; the occurrence of these isolated Mexican populations would suggest that this species has been introduced to this area.

30. Nolana L. Herbs, with blue or white flowers, sometimes succulent (unique among Solanaceae with Sclerophyllax). 94 spp., W slopes of the Andes and the deserts, from Peru (42 endemics) to Chile (42 endemics), nine in both countries, and N. galapagensis (Christoph.) Johnst. disjunc in the Galápagos Islands.

31. Sclerophylax Miers. Leaves opposite, entire, succulent (unique among Solanaceae with some Nolana), usually asymmetrical; petiole sometimes absent; stomata diacytic; flowers solitary, axillary; sepals and petals five. 12 spp. from Argentina, two up to Paraguay and Uruguay.

9.2 SOLANOIDEAE ▸ EXODECONUS CLADE (1/6) - a single genus.

32. Exodeconus Raf. Small annual viscid herbs, campanulate 5-merous flowers, yellow, white, purplish or bluish. 6 spp., the Galápagos Islands, Ecuador (2), Peru (4) to Chile, and Argentina.

9.3 SOLANOIDEAE ▸ TRIBE NICANDREAE (1/3) - a single genus.

33. Nicandra Adans. Subshrubs. Three spp., two endemics to Peru and N. physalodes (L.) Gaertn. from Peru to N Argentina; this species is an ornamental and ruderal in tropical and subtropical areas throughout the world.

9.4 SOLANOIDEAE ▸ TRIBE SOLANDREAE (10/64) - all genera occur in South America.

34. Doselia A.Orejuela & Särkinen. (off Markea) Lianas, very long branches, abundant persistent pubescence of simple eglandular hairs, membranaceous leaves, pendulous long-pedunculate inflorescences (to 50 cm long) with few (1–2(–7)) flowers, distally winged pedicels, large calyces with long-acuminate lobes, and large showy corollas 9–12 cm long. 4 spp., from Colombia to Ecuador.

35. Dyssochroma Miers. (inc. Markea p.p.) Epiphytic shrubs or small trees, occasionally ter-restrial; stems pendulous, the bark thin, flexible and exfoliating, drying a dark, reddish brown; cauliflorous; green, funnelform corollas (greenish) with long, revolute lobes, and exserted stamens; bat pollination. Three spp., endemics to E Brazil, from Ceará to Rio Grande do Sul states, in montane Atlantic Forests.

36. Hawkesiophyton Planch. & Linden. (off Markea) Small greenish or yellowish corollas. Three spp. from over tropical northern South American forests, one up to S Central America, and H. ulei (Dammer) Hunz. (a myrmecophyte species) up to Amazon rainforests of N Brazil.

37. Juanulloa Ruiz & Pav. Epiphytic shrubs or small trees, the stems often hanging and pendulous from the canopy, 1 -20 m; bark o f stems loose and exfoliating when dry, often reddish or reddish brown. 10 spp., one only in Mexico and Central America and 9 in northern South America up to Bolivia, one up to Mexico, with three reported of Amazon rainforests of N Brazil, with J. parviflora (Ducke) Cuatr. endemic to Brazil near Manaus, known only from the type.

38. Markea A.Rich. (exc. Dyssochroma p.p., Doselia, Hawkesiophyton) Hemi-epiphytic to lianas; flowers solitary or arranged in few-flowered pendant cymes, corollas tubular or campanulate; corolla bright orange or more often greenish purple or cream colored, salverform to funnelfore. 14 spp., some has association with ants; nine from Amazon rainforest, mainly in Colombia (8), some reaching into S Panamá, coastal Venezuela and Mount Roraima, M. coccinea Rich., unique in genus with red corollas and salverform flower shape, reaching into center Brazil and Bolivia, near Corumbá municipality in Mato Grosso do Sul, until now the southern limit of the generic distribution; 4 spp. in Brazil, none endemics. Two South American species (M. formicarum Dammer and M. longiflora Miers), both in Brazil, are myrmecophytes.

Two clades recovered in the phylogeny may possibly represent new genera, however their constituent species require much further taxonomic study before new genera can be described with confidence:

§ Markea antioquiensis group ‣ Colombia, included species: M. antioquiensis S. Knapp and M. pilosa S. Knapp, the two species are terrestrial shrubs of the forest understory or, if epiphytic, occur in lower parts of the host tree near the ground; both species also have very large leaves arranged in verticillate whorls, short inflorescences (< 10 cm), very large membranaceous calyces (more than half the length of the corolla), and conspicuous bracts in the inflorescence; M. spruceana Hunz. from Ecuador is morphologically similar and probably belongs to this group but was not included in the molecular analysis.

§ Markea sturmii group ‣ Colombia, included species: M. sturmii Cuatrec. and M. hunzikeri A. Orejuela & C.I. Orozco; profusely branched epiphytic shrubs with slender stems, small greenish flowers up to 3 cm long, filaments shorter than the anthers, and anther thecae confluent at the apices.

39. Merinthopodium Donn. Sm. Epiphytics hrubso r small trees, 1 -10 m, often clamberingo ver branches and hangingf romt he canopy. Three spp., one in C America, and two from Colombia and W Venezuela.

40. Poortmannia Drake. (off Trianaea). Only one spp., P. speciosa Drake, endemic to mountains of Colombia.

41. Schultesianthus Hunz. Climbing epiphytes or scandent shrubs. 8 spp., Mexico to Peru and Venezuela, 5 in South America.

42. Solandra Sw. Climbing epiphytic shrubs or lianas or small trees, glabrous or pubescent, solitary flowers, white greenish. 11 spp. from Mexico (centre of diversity) and Caribbean to Colombia to Bolivia, and S Brazil (2, none endemics); 5 spp. in South America.

43. Trianaea Planch. & Linden. (exc. Poortmannia) Shrubs or lianas, usually epiphytic, sometimes with a woody tuberosity, exceptionally small trees. 5 spp., Andes from Venezuela to Peru.

9.5 SOLANOIDEAE ▸ TRIBE DATUREAE (3/19) - outsider Datura (12; S U.S.A., Mexico).

44. Brugmansia Pers. Subshrubs or small trees; flowers pendant to inclinate, 5-merous, calyx zygomorphic. 5 spp., divided into two informal groups: B. suaveolens (Humb. & Bonpl. ex Willd.) Sweet and B. versicolor Lagerh. from the Amazon rainforest and Ecuadorian lowlands up to southern portion of Atlantic Forest of Brazil, and three remaining from high elevations in the northern and central Andes; moreover, introductions by humans have expanded the range of several species within the Americas and to other continents, and many are found commonly in disturbed areas.

According to IUCN (2019) - in a circumscription radically different from that adopted by Lockwood (1973) and Hay & al. (2012), deductions and historical point field without clear confirmation that six species are originally native to the northern Andes, with B. insignis extending to the west of Brazil; the seventh of them, B. suaveolens, is identified as endemic to the forests of the Brazilian Atlantic Forest; there are no herbarium collections of any species of this genus made from confirmed wild plants; no botanist specialising in this genus has ever reported seeing wild plants of any species; reports by non-specialist botanists of the occurrence of ‘wild’ plants are either misidentifications (usually of Datura), or misinterpretation of remnants or localised escapes from cultivation, usually along creeks and occurring by vegetative propagation from stem fragments; in all such instances investigated in Ecuador and Colombia, the plants are of the anthropogenic hybrid Brugmansia x candida; it is quite clear that such instances do not represent self-sustaining sexually reproducing populations; the complete lack of evidence of fruit dispersal or spontaneous seedlings, combined with the presence of large numbers of fruits containing viable seed, suggests their dispersers are extinct. Hence, all the species should best be regarded as extinct in the wild. They are all threatened with total extinction in their native South America because of the ongoing practice of eradicating them from gardens because of their poisonous nature, combined with the progressive loss of the traditional (indigenous) knowledge of their multiple uses (which is what appears to have been the reason for their long-term survival, perhaps over millennia).

45. Trompettia J.Dupin. Woody shrubs to 2 m tal; stems erect but arching towards apices, many of these becoming spiny, older portions glabrous, becoming pubescent towards younger portions of stem. Only one spp., T. cardenasiana (Hunz.) J. Dupin, from a small region in S Bolivia in the department of Potosí, province of Nor Chichas, north of Santiago de Cotagaita.

9.6 SOLANOIDEAE ▸ TRIBE SOLANEAE (2/c. 1,400) - both genera in South America.

46. Jaltomata Schltdl. 75 spp., from Mexico to Bolivia and Antilles, 69 in South America, highly centered in Peru (58, 56 endemics), one in Galapagos, and one a Caribbean endemic. Nine spp. and 4 morphospecies of this genus in Peru, one of them up to Bolivia, has highly remarkable red-to-orange nectars.

47. Solanum L. [8th BR] Herbs, shrubs, trees, sometimes woody lianas, often epiphytic or with wood rhizomes, usually pentamerous flowers with fused sepals and petals, stellate to pentagonal corollas, and stamens with short filaments and anthers opening by terminal pores. 1,238 spp., cosmopolitan, occurring on all temperate and tropical continents, 1,010 spp. in New World, with their largest diversity in Australia and South America (850), occuping an incredibly wide range of habitats and habits, but the highest diversity of both groups and species occurs in circum-Amazonian tropical South America; the eighth largest genus in Brazil, with 285 spp., 150 endemics (10 of then are rare plants in Brazil, by Plantas Raras do Brasil’s book) and 309 in Peru, the largest diversity; is by far the largest genus in the family, and is the one most likely to be encountered in the Neotropics. Three major clades, 11 medium-sized and 48 smaller clades within the two large major clades.

LINEAGE I

§ Thelopodium Clade ‣ three spp., two from Panamá to Ecuador and S. thelopodium Sendtn. in Colombia to Bolivia and Brazil.

LINEAGE II ‣ non-prickly herbs, shrubs or herbaceous or woody vines that lack stellate trichomes; mostly plurifoliate sympodial unit structure, many compound-leaved species; 4 major clades, and 16 end-lineages, only three in Brazil, with 17 spp. in country. All herbaceous vines and epiphytes in Solanum are found in LINEAGE II except for a single truly herbaceous vine species found in the Nemorense clade (S. hoehnei C.V.Morton).

§ VANA Clade ‣ four subclades.

§ Valdiviense Clade ‣ two woody spp. from Chile.

§ Archaesolanum Clade ‣ 8 spp., restricted to New Guinea, Australia, Tasmania, and New Zealand.

§ Normania Clade ‣ 3 spp., native to NW Africa, the adjacent Iberian Peninsula, and the Macaronesian islands.

§ African Non-Spiny Clade ‣ 14 spp.; all nine Madagascar species are endemic; S. macrothyrsum Dammer is endemic to Mayotte, 4 in continental Africa

§ Regmandra Clade ‣ herbs and shrubs, 11 spp., restricted from Peru and Chile.

§ DulMo ‣ two clades, both globally distributed.

§ Dulcamaroid Clade ‣ woody plants and vary from shrubs or lax shrubs to vines, sometimes large canopy lianas, while other vining species are woody only at the base, especially in temperate climates; 43 spp., from southern Oregon to central Argentina and Caribbean; largest diversities: Argentina (10), Peru (9), Ecuador (9) and Brazil (7).

§ Morelloid Clade ‣ non-spiny herbaceous and suffrutescent species; ca. 75 spp. in five sections (Solanum, Campanulisolanum, Parasolanum, Chamasarachidium and Episarcophyllum), with the black nightshade group (sect.Solanum) is the largest of these with ca. 52 species and is the only group to occur outside of the Americas (Africa, Australasia and Europe), including S. nigrum L., the type of the genus Solanum; more diverse in South America, mainly in Argentina (48), Bolivia (41), Peru (25), Chile (10), Ecuador (9), Paraguay (8) and Brazil (7, two endemics).

§ section Petota ‣ herbaceous perennials with dormant tubers, 107 spp., from Utah and Colorado, throughout the tropical highlands of Mexico, Central America and the Andes, to Argentina, Chile, S Brazil (2, S. commersonii Dunal and S. chacoense Bitter) and Uruguay, with the highest species richness in central Mexico and the central Andes (41 in Peru); S. morelliforme Bitter & Muench it is the only epiphytic wild potato, growing on horizontal branches of mature Arbutus L., cyprus, elm, juniper, pine, or oak trees, often in moss and organic litter central Mexico to Honduras, incredible disjunc in center Bolivia (4,000km distant), and only sp. of this section in both North and South America. Nine smaller lineages.

§ Herpystichum ‣ small herbs or herbaceous vines commonly rooting along nodes with simple to 3–5-foliate leaves, flower buds distinctly onion-shaped and fruits mostly flattened along base to tip axis Lowland rain forest and pre-montane forests from southern. 10 spp., Mexico to northern Peru.

§ Pteroidea ‣ understory herbs, herbaceous vines with adventitious roots and unbranched shrubs with unifoliate sympodial units, simple or pinnate leaves, and axillary inflorescences with small, deeply stellate corollas, fruits often pointed and warty. 10 spp., lowland rain forest and pre-montane forests from Mexico to tropical South America.

§ S. oxycoccoides ‣ somewhat woody vine with simple leaves lacking pseudostipules, and small red fruits with only a few seeds. Only one sp., Peruvian Andes.

§ Articulatum ‣ somewhat woody vines with paired pseudostipules, with large, often branched inflorescences and winged seeds, resembling Basarthrum clade but lacking bayonet hairs (2-celled hairs with a short apical cell). Two spp., montane forests in Costa Rica, Panama, and northern Colombia.

§ Basarthrum ‣ somewhat woody vines or lax shrubs, simple to pinnately compound leaves with frequent interjected leaflets, subtended by a pair of pseudostipules, all species with bayonet hairs (2-celled hairs with a shorter apical cell), with pendant fruits typically green, striped. 16 spp., dry to moist mid-elevation habitats in Central & South America.

§ Anarrhichomenum ‣ somewhat woody vines readily rooting along nodes, simple or compound leaves lacking interjected leaflets, subtended by a single (not paired) pseudostipule at most nodes, fruits orange to red and seeds winged. 12 spp., mid-elevation habitats in Central & South America.

§ Etuberosum ‣ rhizomatous herbs with pinnately compound leaves with frequent interjected leaflets subtended by paired pseudostipules, with many-branched terminal inflorescences, morphologically similar to Petota clade members that have tubers, pedicels articulated at or near the base (unlike Petota clade). 3 spp., Argentina and Chile.

§ Tomato ‣ herbs to woody vines with pinnate leaves with mostly serrate to serrulate margins, some with interjected leaflets, all subtended by paired pseudostipules, all species with strong vegetative odours (glandular trichomes) and yellow corollas, with most species with distinct apical anther modifications (appendages), fruiting pedicels articulating mostly in the distal part. 17 spp., Colombia, Ecuador, Peru and Chile.

§ Petota ‣ tuber-bearing herbs mostly with compound leaves with interjected leaflets with entire leaflet margins, subtended by paired pseudostipules, inflorescences always branched and terminal, corollas usually rotate, fruiting pedicels articulating mostly in the distal part, berries green to purple. 113 spp., North, Central & South America.

LINEAGE III ‣ 7 clades, six in Brazil. 32 end-lineages, 3 endemics to Brazil.

§ S. anomalostemon ‣ a single spp., small herb with glandular trichomes, simple to 3-foliate leaves, distinct cordate shaped anthers with apical modification (beak). Dry forests in southern Peru.

§ Brevantherum Clade ‣ non-prickly, shrubs or trees (herbs) with cylindrical anthers, some with stellate hairs; ca. 80 spp. from S U.S.A. to Argentina, with a center of diversity in the Brazilian Atlantic Forest; a few species are widely distributed and invasive in tropical regions in the Old World. 4 small lineages.

§ Trachytrichium ‣ shrubs with simple entire leaves, simple trichomes and unbranched inflorescences with deeply stellate white corollas, heteromorphic filaments in some species, fruits dull green, many-seeded. Two spp., Argentina, Brazil and Paraguay.

§ Inornatum ‣ herbs to small shrubs with simple entire leaves, simple trichomes, and unbranched inflorescences with deeply stellate white corollas, fruits translucent or dull green to red, with few large seeds. 5 spp. endemic to Atlantic Forest and savanas from C Brazil (cerrado).

§ Gonatotrichum ‣ herbs to shrubs, some species with distinctive 2-celled trichomes with a long apical cell that is bent at a 90-degree angle, inflorescences unbranched with mostly rotate white corollas, strongly heteromorphic filaments in some species, fruits translucent green, explosively dehiscent and with many seeds. 7 spp., SE U.S.A. to southern South America.

§ Brevantherum ‣ trees or shrubs, some species with distinct branching pattern where stem forks have an inflorescence or a leaf arising from it (dichasial branching) and rank fishy vegetative odour, all species with stellate trichomes (or lepidote scales), most species with highly branched inflorescences, some with upright and stout peduncle and pedicels, mostly white corollas, fruits various colours remaining green until just before maturity. 80 spp. Central and South America with many species now found globally.

§ Cyphomandra Clade ‣ non-prickly shrubs, small trees or woody vines with rank vegetative odour (burning smell), plurifoliate or trifoliate sympodial growth, tapered anthers, and with long and mostly branching inflorescences, some species with branched dendritic trichomes. 50 spp., woody shrubs or small trees found in mesic forests from Mexico to N Argentina and SE Brazil. Three small clades. S. sibundoyense (Bohs) Bohs (Colombia) produces some of the largest fruits known in Cyphomandra clade (10cm ✕ 7cm).

§ S. graveolens ‣ woody vine with pinnately compound leaves, white broadly stellate corollas, anthers with apical modification (beaks). A single spp., endemic to Brazilian Atlantic Forest.

§ Cyphomandropsis ‣ shrubs or small trees with simple entire leaves, many with dendritic trichomes, mostly white corollas, some with stone cells. 11 spp., mostly dry habitats across South America.

§ Pachyphylla ‣ shrubs (often lax and spreading) or small trees with distinct branching pattern where stem forks have an inflorescence or a leaf arising from it (dichasial branching), some species with lobed or compound leaves, corollas mostly deeply stellate, anthers with enlarged connectives, some with stone cell. 42 spp., lowland and pre-montane moist forests from Mexico to South America.

§ Geminata Clade ‣ c. 150 spp., non-prickly shrubs or trees with simple entire leaves and sometimes a rank vegetative odour (burning smell), anthers cylindrical; all but one of which occur in the New World tropics; 44 in Brazil; shrubs and small trees mostly occurring in forest understory habitats; they are often inconspicuous, rare and rarely collected, with only a few widely distributed and weedy species. Two small clades.

§ Reductum ‣ small shrubs with dendritic hairs, broadly stellate white corollas, and one species with basal anther modifications (sack). Two spp., endemic to Argentina.

§ Geminata ‣ shrubs or trees, many with geminate (paired/twinned) leaves, inflorescences mostly leaf-opposed and with deeply stellate white flowers with relatively stout, oblong anthers, and fruits that remain green at maturity. c. 150 sp., moist lowland or montane forests from Central to South America.

§ Leptostemonum Clade ‣ prickly shrubs, trees, woody vines or herbs with stellate trichomes, difoliate (geminate or non-geminate) sympodial units dominate, leaves simple or lobed, mostly andromonoecious, simple inflorescences dominate, anthers tapered. ca. 450 spp. the so-called ‘spiny solanums’; c. 250 spp. in Neotropics (111 in Brazil), 80 in Africa, c. 51 spp. in tropical Asia (excluding the island of New Guinea, and the lowlands of Nepal and Bhutan), 150 in Australia. 19 small lineages.

§ S. polygamum ‣ dioecious shrub or small tree with needle-like prickles, simple entire leaves, deeply stellate 5–6-merous white corollas, cylindrical anthers, female flowers lacking developed anthers and with large, forked stigmas, and pubescent orange fruits subtended by large leafy calyx lobes A single species, from Caribbean.

§ Lasiocarpa ‣ shrubs or small trees with needle-like prickles, with mostly repand leaves and unbranched inflorescences, mostly broadly stellate corollas and orange fruits covered in stellate usually glandular trichomes. 13 spp., 11 from Central America to W South America, S. stramonifolium Jacq. up to NE Brazil, and two in Asia and the Pacific.

§ Acanthophora ‣ shrubs and herbs with deeply stellate corollas characterised by having a mix of stellate and simple trichomes on lower leaf surfaces but only simple trichomes on stems and upper leaf surfaces. 22 spp., disturbed and open habitats from Mexico to South America (mostly E Brazil); introduced and naturalized elsewhere.

§ Gardneri ‣ slender-stemmed shrubs and herbs mostly with needle-like prickles, small simple (mostly unlobed) leaves, short and laterally directed inflorescences, deeply stellate corollas, and berries with somewhat accrescent calyces covering less than half of the fruit. 10 spp., dry habitats of eastern to central Brazil, Caribbean, Mexico and Central America, and northern Peru.

§ Thomasiifolium ‣ shrubs or woody vines mostly with broad-based recurved prickles, some species with glandular-stellate trichomes, stems often with short internodes and leaves grouped at the apex, inflorescences unbranched, corollas purple, some bilaterally symmetric, fruits either large and densely pubescent with large seeds, or small and glabrous with accrescent calyces that cover less than half of the fruit. 9 spp., E Brazil.

§ Erythrotrichum ‣ shrubs, woody vines or small trees covered in broad-based recurved prickles, with stellate glandular trichomes, trichomes dense and often reddish-brown in colour, fruits covered in stellate trichomes. 35 spp., tropical South America.

§ Sisymbriifolium ‣ shrubs or herbs with dense needle-like prickles, leaves deeply lobed (nearly pinnate), corollas broadly stellate to nearly rotate, white, fruits red with appressed spiny calyces that spread open at full maturity. Two spp., dry habitats of South America, inc. Brazil.

§ Crinitum ‣ trees, large shrubs or woody vines with scattered broad-based prickles and large flowers with bilaterally symmetric purple corollas and long, heteromorphic anthers, berries large and hardened, oxidize black when cut open, with distinctly swollen calyces. 23 spp., Mexico and South America.

§ Androceras ‣ herbs densely covered in needle-like prickles, with bilaterally symmetric corollas with heteromorphic anthers, dry fruits surrounded by accrescent spiny calyces. 16 spp., S U.S.A. and Mexico.

§ S. campechiense ‣ small shrub covered in needle-like prickles, corollas rotate, fruits with appressed calyces lacking pedicel articulation. Only one sp., dry forests from Texas to Colombia, Jamaica, Puerto Rico.

§ Carolinense ‣ rhizomatous herbs and shrubs covered in needle-like prickles, some species with tubers, corollas rotate, mostly purple, fruits green to yellow mottled, some with appressed calyces. 11 spp., four in North America and seven in South America (Argentina, Bolivia, Brazil, and Paraguay).

§ Bahamense ‣ shrubs and trees with some needle-like prickles, deeply stellate corollas, anthers with stellate trichomes on the adaxial surface, and small juicy red or black fruits on strongly recurved fruiting pedicels. 6 spp., Caribbean.

§ Micracantha ‣ scandent shrubs and woody vines that climb using broad-based recurved prickles with unbranched inflorescences, deeply stellate (mostly white) corollas, and mostly orange or red fruits. 14 spp., Florida to Bolivia and Brazil, including the Caribbean.

§ Asterophorum ‣ tuber-bearing shrubs with difoliate geminate sympodial growth, with broad-based prickles, simple leaf-opposed inflorescences with broadly stellate white corollas, and fruit with appressed calyces covering at least half of the fruit. 4 spp., Brazilian Atlantic Forest.

§ S. multispinum ‣ rhizomatous small shrub covered in needle-like prickles, with white broadly stellate corollas, and yellow berries with prickly accrescent calyces that cover less than half of the fruit. Only one sp., dry habitats of Argentina, Brazil and Paraguay.

§ Torva ‣ shrubs, small trees or woody vines with mostly branched (and stout) inflorescences, with some needle-like or occasionally broad-based prickles, usually broadly stellate corollas, and fruits on stout pedicels often held upright, most species with mucilaginous pulp. 55 spp., worldwide, mostly tropical Americas, with a few members in Asia.

§ S. euacanthum ‣ herb covered in needle-like prickles and stellate glandular trichomes, with broadly stellate white corollas, and dehiscent fruits lacking mesocarp and ‘exploding’ when ripe (probably from tension in the exocarp) with appressed calyces that cover more than half of the fruit. Only one sp., dry habitats of subtropical Argentina.

§ Elaeagnifolium ‣ rhizomatous shrubs or herbs with various degrees of both broad-based recurved and needle-like prickles, weakly bilaterally symmetric purple corollas with heteromorphic anthers and dry fruits, some with lepidote scales. 5 spp., deserts and dry habitats in North and South America (Argentina, Paraguay, Brazil, Uruguay, Chile).

§ Eastern Hemisphere Spiny (EHS) ‣ morphologically diverse group of shrubs, herbs, woody vines or small trees, mostly densely prickly (broad-based recurved and/or needle-like), some rhizomatous, many with appressed calyces that cover more than half of the fruit. 336 spp., most prickly Solanum species outside the Americas are members of this clade Africa, Madagascar, Asia, Australia, Pacific.

§ Wendlandii/Allophyllum Clade ‣ prickly to non-prickly shrubs, herbs and woody vines with tapered anthers. Two clades.

§ Allophyllum ‣ shrubs or herbs lacking prickles, with distinct branching pattern where stem forks have an inflorescence or a leaf arising from it (dichasial branching), leaves with idioblasts containing crystal sand (sand-punctate), inflorescences unbranched, few-flowered and relatively short, corollas broadly stellate or rotate, white or greenish white, some species with stone cells. 4 spp., lowland and montane moist forests of Central and South America.

§ Wendlandii ‣ woody vines with mostly broad-based recurved prickles and with large and generally many-branched inflorescences, many species with lobed or compound leaves, heteromorphic filaments, or swollen calyces in fruit. 9 spp., six in Mexico and Central America, one in S Ecuador and N Peru, and S. alternatopinnatum Steud. in SE Brazil, N Argentina and Paraguay; are all woody or semi-woody vines with small, broad-based recurved prickles.

§ Nemorense ‣ prickly vines, shrubs or herbs, andromonoecious, with tapered anthers. 4 spp., South America.

9.7 SOLANOIDEAE ▸ SALPICHROA CLADE (1/18) - a single genus.

48. Salpichroa Miers. Scandent, pendant, prostrate or straggling shrubs, with rootstocks. 18 spp., Andes from Colombia to Argentina, S. origanifolia Baill. up to S Brazil.

9.8 SOLANOIDEAE ▸ TRIBE PHYSALIDEAE (23/c. 300) - 4 small clades, all in South America.

∎ CUATRESIA CLADE ‣ a single genus.

49. Cuatresia Hunz. Shrubs or small trees. 16 spp. from tropical lowland forests in N South America, one up to Guatemala, Costa Rica and Panamá

∎ SUBTRIBE WITHANIINAE ‣ outsiders Withania (c 20; warm-temperate to tropical regions in the Old World, St. Helena), Nothocestrum (4; Hawai), Tubocapsicum (1; S and E Asia), Discopodium (1; tropical African mountains).

50. Athenaea Sendtn. (inc. Aureliana) Shrubs or small trees; flowers axillary, in fascicles. 14 spp., 13 endemics to S Brazil (3 of then from Minas Gerais and Rio de Janeiro states are rare plants in Brazil, by Plantas Raras do Brasil’s book) and A. fasciculata (Vell.) I.M.C.Rodrigues & Stehmann reaching into adjacent Argentina, disjunct N Bolivia and adjacent Acre state.

51. Deprea Raf. Erect shrubs to small trees with spreading branches, exceptionally herbs with axillary inflorescences generally with three to 15 flowers per node, calyx lobes minute or short, exceptionally long and narrowly triangular, corolla funnel-shaped to stellate, stamen petalum broadening gradually to abruptly in width basipetally, with or without auricles, anthers dorsifixed, generally exserted and mucronate, ovary glabrous, fruiting calyx always accrescent, enveloping the fleshy berry tightly or loosely. 52 spp., highly centered in montane Venezuela to Bolivia (24 endemics to Peru), with one reaching into Central America, almost all in restricted ranges.

∎ SUBTRIBE IOCHROMEAE ‣ all genera in South America.

52. Dunalia Kunth. Shrubs or small trees. 8 spp., the Andes, Colombia to Argentina and Chile.

53. Eriolarynx Hunz. Shrubs or small trees. 3 spp., two in NW Argentina and one from Bolivia and Peru.

54. Iochroma Benth. (inc. Acnistus) Unarmed shrubs or treelets, exceptionally spiny, with colorful, tubular flowers. 40 spp., higly centered in Ecuador, Peru (26, 23 endemic), Bolivia, one up to Cono Sur and three up to Colombia, and I. arborescens (L.) J.M.H.Shaw, widely distributed from Mexico to Peru and SE Brazil; mainly in montane open forested areas.

55. Saracha Ruiz & Pav. Shrubs or small trees; young stems densely hairy with deciduous brown trichomes (thickened, multiseriate, bifurcate or not, ending in a uniseriate row of cells). 5 spp., Andes from Venezuela to Bolivia, three endemics to Peru.

56. Vassobia Rusby. Shrubs or small trees, with cauline spines. Two spp., V. dichotoma (Rusby) Bitter confined in Bolivia and Peru and V. breviflora (Sendtn.) Hunz. from Bolivia, E Paraguay, S Brazil, N Argentina and NW Uruguay.

∎ PHYSALIS CLADE ‣ outsiders Cataracta (1, Mexico), Leucophysalis (5; North America, Mexico, Central America), Schraderanthus (1; Mexico, Guatemala, Venezuela), Capsicophysalis (1; S Mexico to Guatemala and Honduras), Calliphysalis (1; SE U.S.A.), Oryctes (1; SW U.S.A.), Quincula (1; SW U.S.A. to N Mexico), Chamaesaracha (13; North America).

57. Darcyanthus Hunz. Annual sticky herbs, indumentum of long glandular multicellular trichomes; inflorescences fasciculate, sessile, flowers 7–10, pedicellate, 5- merous. Only one sp., D. spruceanus (Hunz.) Hunz. ex N.A. Harriman, oriental rainforests of Peru, Bolivia and Acre state in Brazil.

58. Physalis L. Herbs, rarely shrubs, fruits with inflated calyx. 101 spp., P. alkenkege L. in Eurasia and 100 in New World, almost all in Mexico (81 spp., 48 endemics) to northern South America; 12 spp., almost all widely distributed (only P. galapagoense Waterf. more restricted) in South America (only three in Brazil, no endemics).

59. Witheringia L'Her. Herbs or shrubs. 15 spp., SE Mexico and southwards to Bolivia (7 in South America), Caribbean, Amazonas and Acre states in N Brazil (only the widely distributed W. solanacea L'Hér.).

9.9 SOLANOIDEAE ▸ TRIBE CAPSICEAE (2/172) - both genera in South America.

60. Capsicum L. Shrub or small tree habit with long branches and vigorous suckers and sprouts; flowers in most species are multiple per node, erect; the corolla is stellate, white with greenish/yellowish spots in the throat and purplish red spots in the petals lobes; fruits are small and roundish, as large as a pea, pendulous, green when immature, greenish yellow, almost translucent and deciduous when ripe, all pungent, quite hot when immature, less hot, sweet and juicy after ripening; seeds very coriaceous. 43 spp., from the SW U.S. to Argentina, slightly centered in E Brazil (16, 12 endemics) and in Bolivia. 11 lineages emerges from this genus, with only C. benoistii Hunz. ex Barboza unassignated in a clade. C. baccatum L. from Peru to S Brazil and Argentina, C. pubescens Ruiz & Pav. from Ecuador to Bolivia, and C. eximium Hunz. from Bolivia to Argentina have higlhy uncommon yellow nectars.

§ Andean ‣ 9 spp., C Mexico to N Venezuela and N Peru.

§ Annuum ‣ 4 spp., C. galapagoense Hunz. nested among the closely related C. frutescens L., C. chinense L. and C. annuum L.; whereas C. annuum var. glabriusculum is known as a weed throughout Central and northern South America up to southern North America, C. annuum var. annuum, C. frutescens and C. chinense are only known from cultivation (although it is very likely that C. chinense was domesticated in northern Brazil), and are well known for their valuable fruits, either the pungent cultivars or the sweet types used as food, condiments, and medicine.

§ Atlantic Forest ‣ 12 spp., endemic to Atlantic Forest in E Brazil. C. hunzikerianum Barboza & Bianchetti, endemic to Atlantic Forest in Brazil, has the largest flowers of genus, reache up to 1.6 cm long. Urceolate corollas among this genus occur only in two spp. from Andes and in the Brazilian endemic C. friburgense Bianchetii & Barboza.

§ Baccatum ‣ 3 spp., C. praetermissum (Heiser & P.G.Smith) Hunz. from SE Brazil, C. chacoense Hunz. from Bolivia to central Argentina and Paraguay, and C. baccatum L. from Bolivia to N Argentina and SE Brazil, reaching Colombia in the north.

§ Bolivian ‣ 5 spp., Bolivia to Peru, with C. coccineum (Rusby) Hunz. reaching in Acre state in W Brazil.

§ Caatinga ‣ two spp., from dry areas in NE Brazil, N Colombia and N Venezuela.

§ Flexuosum ‣ only one sp., C. flexuosum Sendtn, from E & S Brazil to NE. Argentina.

§ Longidentatum ‣ only one sp., C. longidentatum Agra & Barboza, endemic to NE Brazil.

§ Purple Corolla ‣ 3 spp., Bolivia to Argentina and one cultigen.

§ Pubescens ‣ only one sp., the cultivated C. pubescens Ruiz & Pav. from Ecuador to Bolivia.

§ Tovarii ‣ only one sp., C. tovarii Eshbaugh, P.G.Sm. & Nickrent, endemic to Peru.

61. Lycianthes Bitter. Shrubs, trees, vines or perennial herbs. 150-160 spp., 121 in New World, 61 in South America, 37-50 in E Asia; 13 spp. in Brazil, belonging to Lycianthes subg. Polymeris, with three subordinate sections (sects. Asaropsis, Simplicipila, and Eupolymeris), 4 endemics.

59. AQUIFOLIALES

FAMILIES ABSENTS IN SOUTH AMERICA: HELWINGIACEAE (1/4).

AQUIFOLIACEAE

§ FULLY CHLOROTROPHIC

Genera/species 1/560 Distribution eastern North America, Mexico, Caribbean, Central America, tropical South America, W, C and S Europe, Türkiye and eastwards to northern Iran, Macaronesia, tropical and subtropical parts of Africa (one species), Madagascar, India and Himalaya to Japan and Russian Far East, SE Asia, Malesia, New Guinea, Melanesia, northern Australia. Habit usually dioecious or bisexual (rarely polygamomonoecious), usually evergreen (rarely deciduous) trees or shrubs (rarely lianas). Only one genus.

Aquifoliaceae is frequently confused with Celastraceae (Monteverdia) and Symplocaceae (Symplocos Jacq.). From Monteverdia it is readily distinguished by the lack of a disc in the flowers (conspicuous in Monteverdia), and the fruit (drupe vs. capsule in Monteverdia). Leaves in species of Ilex are not pruinose (covered with a wax layer) like in many species of Monteverdia. Additionally, the top of the branches in Monteverdia is, in many species, 4-angled or striate, or even winged. In Ilex the branches are usually rounded in transverse section. Symplocos species often bear flowers with 5-100 stamens (in 1-4 whorls), with a nectariferous disc surrounding the style base (absent in Ilex). The style is clearly distinct from the ovary (short or absent in Ilex), this (sub) inferior (superior in Ilex). Leaves in Symplocos usually become light-green when dry, different from Ilex (grayish or dark-colored). Additionally, many species of Symplocos have glandular-denticulate leaves.

SYSTEMATIC a single genus worldwide.

1. Ilex L. Trees or shrubs, usually evergreen, sometimes deciduous, rarely prostrate subshrubs (only in Brazilian Ilex prostrata Groppo), often with roots crown; leaves alternate, rarely (sub)opposite, simple, usually glabrous; inflorescences axillary, derived from thyrses: cymes, thyrsoids, fascicles or apparently solitary flowers; flowers 4-6(-23)-merous, actinomorphic, petals usually white or cream, base connate to the base; fruit a drupe, with 1-6 pyrenes, with one seed each, epicarp red, purple or virtually black, mesocarp fleshy; seeds with oily, proteinaceous, abundant endosperm. 500 spp., ca. 277 in New World, 202 in South America; in the Neotropics can be found from sea level up to ca. 4,000 m (Andes); 58 spp. in Brazil, 42 endemics; Nemopanthus Raf. (one species in North America) was recently included under the synonymy of Ilex; I. auricula S. Andrews, I. blanchetii Loes. and I. mucugensis Groppo from Bahia, and I. prostrata Groppo from Minas Gerais are rare plants in Brazil, by Plantas Raras do Brasil’s book.

Several species of Ilex, e.g. I. aquifolium L., I. cornuta Lindl. & Paxton, I. crenata Thunb., and I. opaca Aiton are cultivated as ornamentals, mainly because their decorative fruits and/or foliage. However, their use in the Neotropics as ornamentals is occasional.

The species with most economic importance in the Neotropics is by far I. paraguariensis A.St.-Hil., native in Argentina, Brazil, Paraguay, and Uruguay; from the leaves of this species is made a tea, the ‘mate’, ‘erva-mate’ or ‘yerba-mate’, largely consumed in South Brazil, Argentina, Paraguay and Uruguay. Mate is a part of the popular culture in those places, being characteristically served hot in a gourd (called ‘cuia’ in Brazil) and drank with a straw (the ‘bomba’). It can be also be served as a cold beverage (‘tererê’), usually in a decorative cow horn, especially in Paraguay and Brazil.

Other species of llex, such as I. dumosa Reissek or I. theezans Mart. were sometimes used in the past as substitutes of the true mate; another species, as I. tarapotina Loes. (Colombia and Peru) and I. vomitoria Aiton (Mexico and North America) are used as teas, mainly because their emetic or stimulating effects; the wood of some species, as I. affinis Gardn., I. brevicuspis Reissek, I. cerasifolia Reissek, I. dumosa, and I. theezans is locally used in Brazil in works of carpentry, and as fuel; a unnmaed species of Bajo Calima region in Choco, W Colombia has the largest leaves of Aquifoliaceae (15-25 cm × c.10 cm).

PHYLLONOMACEAE

§ FULLY CHLOROTROPHIC

Genera/species 1/4 Distribution S Mexico (Durango) to NW Bolivia (the northern Andes); absent in Venezuela. Habit bisexual, evergreen trees or shrubs; inflorescence axillary, fasciculate or umbel-like, seemingly epiphyllous (inflorescence seemingly arising from meristem on adaxial surface of leaf) and apparently inserted adaxially on mid-vein.

SYSTEMATIC a single genus.

1. Phyllonoma Willd. ex Schultes. Shrubs or small trees, infrequently epiphytic, glabrous throughout, the young twigs variously grooved; leaves alternate, inflorescences cymose and branched or racemose and essentially unbranched above the point of insertion, arising on the midvein on the adaxial leaf surface anywhere near the middle of the blade to the base of the acumen or, less frequently, on the acumen (true epiphylly, one the most classic among New World plants); flowers 2-4 mm in diameter; sepals 5; petals 5. 4 spp., two in South America: P. ruscifolia Willdenov ex. Schultes, of inflorescence cymose (Guatemala, Costa Rica and Panamá, Colombia; N Peru to NW Bolivia), and P. weberbaueri Engler., with inflorescence racemose (in Andes of central and southern Peru and NW Bolivia, in cloud forest at altitudes over 2,000 m); their inflorescences epiphyllous, borne at various positions on the adaxial surface of the leaf blade, differentiates Phyllonomaceae from any other Neotropical Angiosperm family.

60. CARDIOPTERIDALES

TWO FAMILIES, BOTH IN SOUTH AMERICA.

CARDIOPTERIDACAE

§ FULLY CHLOROTROPHIC

Genera/species 3–5/36–41 Distribution mountain regions in Central and South America, SE Asia, Taiwan in China, Malesia, New Guinea, Melanesia, E Australia, islands in the Pacific. Habit usually bisexual (sometimes andromonoecious, gynomonoecious, polygamomonoecious, dioecious, androdioecious, or gynodioecious), evergreen trees or shrubs, or climbing and twining herbs (Cardiopteris). Trees or shrubs occasionally climbing (Citronella D. Don). Two genera in Neotropics;

New World Icacinaceae genera are found within the tropics, a few species occur in subtropical regions, and one species outside of these (Citronella mucronata at 45° S). Native with some species of Citronella cultivated.

SYSTEMATIC outsiders Cardiopteris (3; E Himalayas to SE Asia, Malesia to New Guinea, Bismarck Is. and Solomon Islands), Pseudobotrys (2; New Guinea), Gonocaryum (11; Hainan and Taiwan in China, Indochina, the Malay Peninsula, Malesia to New Guinea and Bismarck Is.), Leptaulus (6; tropical Africa, Madagascar).

1. Citronella D. Don. Trees or shrubs occasionally climbing; all parts lepidote-stellate or indumentum of simple hairs; leaves simple, alternate, distichous, margin entire, occasionally serrate with small domatia abaxially; inflorescence terminal or axillary, paniculate or cymose; flowers bisexual or occasionally polygamous, glomerulate, actinomorphic, pentamerous; fruit drupes, exocarp lepidote-stellate becoming glabrous, mesocarp fleshy, endocarp hard; seed 1, endosperm abundant. 21 spp. in Malaysia, Pacific and 10 in the New World, all in South America, from in Costa Rica to Chile and Argentina, some endemic to Chile, Colombia and Brazil (5, C. engleriana (Loes.) R.A. Howard endemic); Citronella still has many problems for recognition at specific level.

STEMONURACEAE

§ FULLY CHLOROTROPHIC

Genera/species 12/c 80 Distribution southern India (W Ghats), Sri Lanka, Assam, Burma, Indochina, Malesia, New Guinea, Melanesia, Queensland, Central and South America. Habit usually dioecious (sometimes bisexual), evergreen trees or shrubs. The family Stemonuraceae has evergreen trees or shrubs, stems with indumentum of simple hairs, and includes eleven genera from the Old World and Discophora Miers from the New World.

SYSTEMATIC outsiders Lasianthera (1; tropical W Africa); Cantleya (1; Malesia), Codiocarpus (2; Andaman and Nicobar Is., Philippines, Aru Islands), Gastrolepis (2; New Caledonia), Gomphandra (60–65; SE Asia, Malesia to New Guinea and Solomon Is.), Grisollea (2; Madagascar, Comoros, Seychelles), Hartleya (1; New Guinea), Irvingbaileya (1; NE Queensland), Medusanthera (10; Malesia to islands in W Pacific), Stemonurus (14; Sri Lanka, Andaman Islands, Vietnam, Malesia), Whitmorea (1; Solomon Islands).

1. Discophora Miers. Shrubs to trees, 3-20 m tall; bisexual; simple hairs; axillary inflorescences, supraxillary or cauliflorous, thyrsoid; flowers 5-merous, actinomorphic, corolla pale green; fruit drupaceous, white at immaturity, purple at maturity. Three spp.: D. froesii Pires is endemic to Venezuela, and has leaves densely pubescent with long and erect trichomes; blades narrowly ovate to lanceolate; secondary veins 10-13; D. guianensis Miers is a small tree from Costa Rica to almost Amazon rainforest (up to NE Pará and NE Mato Grosso states), south into NW Bolivia, disjunct in small populations in SE Bahia and N Espírito Santo states in E Brazil, near coast (possibly occurs in center Brazil); has leaves glabrous, if pubescent, trichomes shorter and more adpressed than above; blades oblong, narrowly oblong, ovate, or narrowly ovate; secondary veins 7-10; and D. montana R.A. Howard occurs in Colombia, in Antioquia, Boyaca, Cundinamarca and Meta departaments, and in Peru from Amazonas, in montane environments, and differs from D. guianensis in its smaller drupe, c. 1 cm long, and leaves with a long slender point; in lowland rainforest below 500 m alt.

61. ASTERALES

FAMILIES ABSENTS IN SOUTH AMERICA: ALSEUOSMIACEAE (4/10), ARGOPHYLLACEAE (2/24), PENTAPHRAGMATACEAE (1/31), PHELLINACEAE (1/10), ROUSSEACEAE (4/15).

LINEAGE 1 of 3: CAMPANULIDS

CAMPANULACEAE

§ FULLY CHLOROTROPHIC

Genera/species 94/c. 2,200 Distribution cosmopolitan except extreme polar regions; few representatives in Malesia, Australia and New Zealand. Habit usually bisexual (rarely dioecious or gynodioecious), usually perennial (sometimes annual or biennial) herbs (sometimes evergreen shrubs, small trees or lianas). Some species are aquatic. Others are xerophytic.

Key differences from similar families - Campanulaceae is easy to distinguish by combined presence of latex, simple leaves and inferior ovary. Material from other families with long and reddish or orange corolla -tube is often placed in Campanulaceae, mainly Acanthaceae, Lamiaceae and Rubiaceae. However, these families differing in leaves commonly opposite and free stamens and anthers. Allied families or others traditionally placed close to Campanulaceae such as Pentaphragmataceae, Stylidiaceae (including Donatiaceae), Sphenocleaceae and Goodeniaceae (including Brunoniaceae), can be distinguished from the Campanulaceae by the following features: they lack latex; Goodeniaceae have a style with apical hairy pollen -collecting indusium and stylar cup; Pentaphragmataceae and Stylidiaceae have extrorse anthers.

SYSTEMATIC five subfamilies, Nemacladoideae (2/14, SW U.S.A., Mexico) and Cyphioideae (1/c 65, Africa, the Cape Verde Islands, with their largest diversity in S Africa) do not occur in South America.

1. SUBFAMILY CAMPANULOIDEAE (44/900–950) - three tribes, Cyanantheae (10/64, Ethiopia, Canary Islands, tropical E Africa tropical and E Asia, SE Asia, Malesia) does not occur in South America.

1.1 CAMPANULOIDEAE ▸ TRIBE WAHLENBERGIEAE (16/340–350) - outsiders Siphocodon (2; W Cape), Rhigiophyllum (1; W Cape), Feeria (1; Morocco), Kericodon (1; W and E Cape), Nesocodon (1; Mauritius), Heterochaenia (3; Réunion), Berenice (1; Réunion), Microcodon (3; N and W Cape), Craterocapsa (5; South Africa, Swaziland, Lesotho, Zimbabwe), Namacodon (1; C Namibia), Gunillaea (2; tropical Africa, Madagascar), Prismatocarpus (c 30; tropical and S Africa), Roella (c 20; W Cape, one species in E Cape and KwaZulu-Natal), Merciera (6; W Cape), Treichelia (2; W Cape).

1. Wahlenbergia Schrad. ex Roth. Annual or perennial herbs, subshrubs and shrubs. 260 spp., mainly Old World to SW Pacific, 14 spp. in Colombia to S. South America; five endemics to Juan Fernandes, three endemics to Brazil, also three endemics to Peru; W. linarioides Lam. from Ecuador to SW Brazil, Argentina and S Chile; W. peruviana A. Gray from Peru to Argentina, and W. perrottetii (A. DC.) Thulin from Brazil, French Guiana and Africa.

1.2 CAMPANULOIDEAE ▸ TRIBE CAMPANULEAE (18/590 –610) - utsiders Jasione (13–16; Europe, Mediterranean, SW Asia), Musschia (3; Islas Desertas), Campanula (500 – 520; temperate regions on the Northern Hemisphere, tropical mountains), Legousia (7; Europe, Mediterranean), Petromarula (1; Crete), Phyteuma (c 40; Europe, Mediterranean, temperate Asia), Physoplexis (1; the Alps), Trachelium (2; Europe, Mediterranean), Homocodon (2; Bhutan, SW China), Favratia (1; the Alps), Zeugandra (2; Iran), Peracarpa (1; tropical Asia), Heterocodon (1; SW Canada, W U.S.A.), Githopsis (4; Vancouver Island to Baja California), Cryptocodon (1; Pamir), Cylindrocarpa (1; Kyrgyzstan), Sergia (2; Tadzhikistan; Kazakhstan).

2. Triodanis Raf. Annuals; leaves sessile or short-petiolate; flowers medium-sized, sessile, 1–3(–8) in upper axils, forming a spike-like inflorescence, lower ones smaller and cleistogamous; hypanthium elongate; calyx lobes 3 in cleistogamous flowers; corolla lavender blue, rotate. 7 spp., North America to Mexico, T. perfoliata subsp. biflora (Ruiz & Pav.) Bradley also in Ecuador to S. South America, S Brazil and Uruguay.

2. SUBFAMILY CYPHOCARPOIDEAE (1/3) ▸ a single genus.

3. Cyphocarpus Miers. Annual and perennial herbs, sometimes twining vines; leaves pinnatifid; flowers small to medium-sized. Three spp. in N & C Chile.

3. SUBFAMILY LOBELIOIDEAE (26/1,180 – 1,200) ▸ outsiders Grammatotheca (1; W and E Cape, KwaZulu-Natal), Dialypetalum (5; Madagascar), Wimmerella (10; S Africa), Dielsantha (1; C Africa), Monopsis (13; tropical and S Africa), Isotoma (12; Australia), Ruthiella (4; New Guinea), Solenopsis (7; Canary Islands, Mediterranean to Türkiye, Cyprus and Syria), Porterella (1; W N America), Howellia (1; W U.S.A.), Hippobroma (1; Caribbean), Heterotoma (1; Mexico, Central America), Sclerotheca (10; Rarotonga, Society Islands, Marquesas Islands, Rapa Island), Trematolobelia (4; Hawaii), Brighamia (2; Hawaii), Delissea (9; Hawaii), Cyanea (c 60; Hawaii), Clermontia (22; Hawaii).

The ‘CBS clade’ includes three shrubby neotropical genera, Centropogon, Burmeistera and Siphocampylus, which collectively comprise almost half of the species of Lobelioideae; Burmeistera and Centropogon have fleshy fruits and Siphocampylus is capsular type; Burmeistera is a monophyletic group primarily bat-pollinated, herbs or hemi-epiphytic herbs or subshrubs that climb nearby vegetation; tubular corollas with reproductive parts positioned above the opening; Siphocampylus is inferred to be a paraphyletic relative to fleshy -fruited Centropogon and Burmeistera, but fleshy fruits have evolved repeatedly, making Centropogon polyphyletic. Both genera are difficult to distinguish; apart from the fruit there are subtle differences regarding the lower anthers; the other representative genus in Neotropics is Lysipomia, a monophyletic group of small cushion-forming plants endemic to the high Andes; the genus is sister to the clade comprising the remaining three shrubby genera.

4. Burmeistera Karst. & Triana. Suffrutescent (sometimes hemi-epiphytic) herbs, trees or shrubs, sometimes scandent lianas, often green flowers are primarily pollinated by nectar bats, with latex. 120 spp. (42 + 78), from Honduras to Peru and Venezuela, highly centered in Colombia and Ecuador, 96 (54 + 42) in South America, 57 in Colombia, 51 in Ecuador, two up to Peru and one in Brazil, B. pallida (Drake) E.Wimm., collected in Mount Neblina.

5. Centropogon C.Presl. Suffrutescent herbs or shrubs, sometimes scandent lianas; flowers medium-sized to large, solitary, axillary, rarely forming a terminal corymb or raceme. 211 spp. over Neotropics, from S Mexico to Bolivia and Brazil, 195 in South America, two in the Lesser Antilles; only the widely distributed C. cornutus (L.) Druce occur in Brazil.

6. Diastatea Scheidw. Annuals. Flowers small, in terminal, sometimes secund 5–30-flowered racemes (rarely solitary, axillary); pedicels ebracteolate; corolla bilabiate, purplish, blue or white. 5 spp. from central Mexico to Panamá, with D. micrantha (Kunth) McVaugh extending south to Bolivia, Argentina and Venezuela.

7. Downingia Torr. Annuals; leaves much smaller than bracts, sessile; corolla bilabiate, blue (rarely pinkish or white), often marked with yellow or white on ventral lip; flowers small, sessile, in a terminal raceme. 13 spp., W Canada to NW Mexico, D. pusilla (G.Don ex A.DC.) Torr also in Chile to Argentina.

8. Legenere McVaugh. Emergent aquatic annuals; leaves sessile; flowers small, both chasmogamous and cleistogamous, pedicellate, in a terminal raceme; pedicels ebracteolate; corolla bilabiate, yellow. Only one sp., L. valdiviana (Phil.) E. Winn., highly disjunct in California, Chile to S. Argentina.

9. Lobelia L. Annual or perennial herbs, shrubs, trees or giant rosette plants (often pliestesial). 400 spp., pantropical, 160 in New World, from Mexico to Argentina, Bolivia and Brazil, Caribbean; 38 spp. from South America, 20 in Brazil, 12 endemics (one, in Distrito Federal, a rare species in Brazil by Plantas Raras do Brasil’s book).

10. Lysipomia Kunth. Dwarf perennial herbs, often forming cushions, sometimes scapose; leaves sessile or petiolate, commonly rosulate. 30 spp. in Andes of South America, from Venezuela to Bolivia; the smallest of all angiosperms is L. mitsyae Sylvester & D.Quandt, a minute glabrous short-lived monocarpic herb and also from Peru, 1.8–4(–5.5) mm tall, is smaller; this species occur only in Cuzco region.

11. Siphocampylus Pohl. Suffrutescent herbs or shrubs, sometimes scandent or twining lianas; leaves rarely opposite or whorled. 238 spp. Neotropics, from Costa Rica to Argentina and in Greater Antilles, highly centerd in Andes from Peru and Bolivia; 217 in South America, 28 in Brazil, 25 endemics (two of then, from Rio de Janeiro and São Paulo states, are rare species in Brazil by Plantas Raras do Brasil’s book).

Syphocampylus umbellatus clade has 6 spp., all restricted to the Central Andes of Peru and Bolivia, with the exception of S. umbellatus (Kunth) G.Don, whose range also extends to Brazil; this clade is composed of robust shrubs or trees that are exceptionally tall for the centropogonid clade, or rarely scandent subshrubs.

LINEAGE 2 of 3: STYLIDIIDS

STYLIDIACEAE

§ FULLY CHLOROTROPHIC

Genera/species 5/314 Distribution mainly extratropical regions of Australia, Tasmania, New Zealand, South and SE Asia, southernmost South America. Habit Usually bisexual (sometimes monoecious or polygamomonoecious), usually perennial herbs (sometimes dwarf shrubs; in Donatia and Phyllachne cushion-shaped). Some species are helophytes and numerous species are xerophytes.

SYSTEMATIC two subfamlies, both in South America.

1. SUBFAMILY DONATIOIDEAE (1/2) ▸ a single genus.

1. Donatia J. R. Forst. Much-branched short herbs, forming dense, often large, cushion-like expanses. Two spp., one in Australia and New Zealand, and D. fascicularis J.R. Forst. & G. Forst. from S Chile and Argentina.

2. SUBFAMILY STYLIDIOIDEAE (3/320) ▸ outsiders Levenhookia (10; W Australia, S Australia, Victoria, New South Wales), Stylidium (probably more than 300; Bengal, Sri Lanka, Burma, Vietnam, southern coast of China, the Malay Peninsula, Philippines, Australia, Tasmania, New Zealand incl. Stewart Island, with almost all species confined to Australia).

2. Forstera G.Forst. (inc. Phyllachne) Much-branched short herbs, forming dense, often large, cushion-like expanses. 9 spp., Tasmania, New Zealand incl. Stewart Island (3, two endemics), Auckland Islands, Campbell Island, one species, F. uliginosa Hombr. & Jacquinot ex Decne, in Chile and Argentina.

LINEAGE 3 of 3: CORE ASTERALES

MENYANTHACEAE

§ FULLY CHLOROTROPHIC

Genera/species 6/60–70 Distribution cosmopolitan except polar areas, with their largest diversity in Australia. Habit usually bisexual (rarely monoecious, dioecious or gynodioecious), usually perennial (rarely annual) herbs. Hydrophytes (aquatic) or helophytes; three genus in New World, two occurs only in North America.

The Gentianaceae differs in that their members having opposite leaves.

SYSTEMATIC outsiders Menyanthes (1; temperate regions on the Northern Hemisphere), Nephrophyllidium (1; N Japan, NW North America); Ornduffia (6; SW W Australia, S Australia, Queensland to Victoria, Tasmania), Villarsia (3; W Cape), Liparophyllum (8; SW W Australia, S Australia, Queensland to Victoria, Tasmania, New Zealand).

1. Nymphoides Séguier. Aquatic perennial herbs; leaves above the water-level, surfacing or rarely submerged, alternate, simple, entire, orbicular, occasionally slightly succulent, cordate or reniform, margins entire, dentate or crenate with hydathodal glands, venation palmate; estipulate; petiolate, petiole bases sheathing; flowers in terminal or axillary fascicles, heads, panicles or solitary, actinomorphic; sepals 5, basally connate; petals 5, basally connate, winged; fruit a capsule; seeds 1-many, dry, 0.4 - 5.2 mm, flattened or lenticular. 43 spp., 13 spp. in Africa and Madagascar, 12 in Australia, 9 in India, and one in Eurasia; 8 spp. in New World, two in E North America, one in Mexico and Guatemala, and five in South America: Colombia and Bolivia one endemic each, N. verrucosa (R.E.Fr) A. Galán & G. Navarro in Bolivia and Cono Sur, the yellow flowered N. grayana Kuntze in Cuba, Bahamas, Pantanal and Tocantins state in center Brazil, also in Africa (a remarkable disjunction), and the white flowered N. humboldtiana Kuntze in over New World.

GOODENIACEAE

§ FULLY CHLOROTROPHIC

Genera/species 7/c. 440 Distribution tropical and subtropical coastal areas along the Atlantic, the Indian and the Pacific Oceans, Australia, Tasmania, islands in the Pacific and the Indian Oceans, with their largest diversity in W Australia and Tasmania. Habit bisexual perennial or annual herbs, suffrutices or shrubs (rarely climbing or arborescent). Many representatives are xerophytic.

Members of the Neotropical Goodeniaceae are readily distinguished by an unique floral character known as the 'indusium' which is a cup-shaped specialized stylar outgrowth below the apex of the style (that attracted the interest of Charles Darwin), and which acts as a pollen presenter.

SYSTEMATIC outsiders Leschenaultia (26–27; S New Guinea, Australia, W Australia), Anthotium (4; W Australia), Dampiera (c 65; Australia, with their highest diversity in W Australia); Brunonia (1; Australia, Tasmania); Coopernookia (6; W Australia, S Australia, New South Wales).

1. Goodenia Sm. (inc. Selliera) Perennial shrubs or subshrubs, or annual or perennial herbs, sometimes stoloniferous and rooting at nodes. 235 spp., New Guinea, Australia, one species also in Malesia to S China, one on Java, and G. radicans (Cav.) Pers. in South Australia, Victoria, New South Wales, Tasmania, New Zealand, Stewart Island and continental Chile.

2. Scaevola L. Perennial herbs, shrubs, scramblers or small trees; leaves alternate or rarely opposite. 130 spp., two in New World: S. plumieri (L.) Vahl, found throughout sandy coastal plain ecosystem areas of the Neotropics, from both coasts of U.S.A. to Ecuador (absent in Colombia, however), in Pacific coast, and Brazil, Venezuela and Caribbean, in Atlantic Coast - also in Africa and India; and S. wrightii M. Gómez, endemic to Cuba.

CALYCERACEAE

§ FULLY CHLOROTROPHIC

Genera/species 8/46 Distribution South America, especially in extratropical regions (southern Peru and SE Brazil to S Chile), the Falkland Islands. Habit usually bisexual (rarely monoecious? or gynomonoecious, in Acicarpha often andromonoecious with male flowers in centre), usually perennial (rarely annual) herbs (occasionally somewhat woody at base); the greater part of the family (all genera and c. 40) are found in Argentina.

The Calyceraceae is easily separated from the Compositae by the leaf-like bracts forming the involucre rather than the involucral bracts in the Compositae that differ clearly from the cauline leaves.

SYSTEMATIC - all genera in South America.

1. Acicarpha Juss. (inc. Boopis p.p.) Annual or perennial herbs; stems more or less branched, erect or procumbent, terminating in an inflorescence overtopped by lateral branches. 7 spp., S Brazil (5, 2 endemics, one of then from Itatiaia Massif a rare plant in Brazil by Plantas Raras do Brasil’s book), S Paraguay, Uruguay, and NC Argentina, along riversides or the Atlantic seashore.

2. Anachoretes S.Denham & Pozner. (off Boopis) Hemicryptophytic, glabrous, non-glutinose herbs with decumbent lateral branches. Only one sp., A. castillonii (Hicken) S.Denham & Pozner, endemic to Sierra de Ambato, Catamarca, Argentina.

3. Asynthema S.Denham & Pozner. (off Boopis) Therophytic, small, glabrous, non-glutinose herbs with lateral, erect or decumbent, slender branches. Only one sp., A. gracile (Phil.) S.Denham & Pozner, is widely distributed in Argentina and Chile, in Monte, Patagónica (from Mendoza to Santa Cruz in Argentina) and Chilena Central biogeographic provinces, on arid soils.

4. Boopsis Juss. (exc. Acicarpha p.p., Anachoretes, Asynthema) Perennial or rarely annual herbs, glabrous; stems erect or decumbent, scapose or ramified; leaves sometimes clustered in a rosette. Only one sp., B. anthemoides Juss., Argentina and Uruguay, from sea level to 3,500m, in Pampeana, Espinal, Chaqueña, Monte, Patagónica and Prepuneña biogeographic provinces. B. anthemoides is the most widely distributed species of the family.

5. Calycera Cav. Hemicryptophytes, chamaephytes, cryptophytes or therophytes herbs, glabrous or weekly lanuginose; stems erect or decumbent; leaves alternate or in rosetes. 6 spp., from Bolivia and N Chile to W Argentina (one specimen has been collected in Peru), reaching the Atlantic seashore in Uruguay, extending northwards along coastal dunes from Buenos Aires and Uruguay.

6. Gamocarpha DC. (inc. Nastanthus, Boopsis p.p.) Habit rosulate herbs with hemicryptophytic, cushion-shaped or caespitose habit according to the growth of lateral branches; lateral branches erect or decumbent, foliose, scapiform (short and thick or long and slender), stoloniferous, glabrous, non glutinose. 13 spp., Argentina and Chile, where it is widely distributed along the Andes and Patagonia, less frequent in Monte, and in Malvinas Island.

7. Leucocera Turcz. (inc. Calycera p.p., Boopsis p.p.) Annual, chamaephytic or rosulate hemicryptophytic herbs with erect, decumbent or rhizomatous branches, glabrous or pubescent with uniseriate, simple trichomes on leaves. 7 spp., Andean regions from Chile and Argentina.

8. Moschopsis Phil. (inc. Gamocarpa p.p., Boopsis p.p.) Glabrous, usually cushion-shaped, rosulate herbs with lateral rhizomatous or stoloniform branches producing new cushion-shaped, crowded rosettes. 10 spp., Chile (Region Metropolitana) and Argentina (San Juan, Mendoza, Neuquén), to the southern portion of Patagonia and in Tierra del Fuego, from sea level to 3,000m.

ASTERACEAE

§ FULLY CHLOROTROPHIC

Genera/species 1,702/34,588 Distribution cosmopolitan except Antarctica. Habit usually bisexual (sometimes monoecious, gynomonoecious, polygamomonoecious, dioecious, androdioecious, or gynodioecious), usually perennial, biennial or annual herbs (sometimes evergreen, rarely deciduous, shrubs, trees or lianas). Some species are succulent. Numerous representatives are spiny or prickly. Many Asteraceae have stem or root nodules or lignotuber. A large number of species are xeromorphous. Some species have C₄ and/or CAM physiology. Often with a strong scent or odour.

The majority of Compositae are subshrubs, shrubs or perennial herbs, adapted to moderately xeric conditions, but most life forms are represented, from ephemeral desert annuals, pyrophytes and hemicryptophytes to trees of 30m or more, including especially in Senecioneae, leaf and stem succulents, also halophytes, marsh plants, lianas, epiphytes and aquatics, though the later two life forms are comparatively infrequent and submerged aquatics rare.

Pachycaul, megaphytic, polycarpic or sometimes monocarpic trees, shrubs or herbs occur widely on tropical mountains and oceanic islands, e.g. Centaurodendron (Cynareae, Juan Fernández Islands), Sonchus (Cichorieae, Macaronesia), Dendroseris (Cichorieae, Juan Fernández Islands), Espeletia (Millerieae/Heliantheae s.l., Andes), Argyroxiphium (Madieae/Heliantheae s.l., Hawaii), Lachanodes (Senecioneae, St. Helena) and Dendrosenecio (Senecioneae, tropical east African mountains).

Compositae are prominent among the plants utilized by peoples in native cultures in all parts of the world, especially for medicinal purposes. For example, in China 500 and in Mexico 180 species of wildgrowing Compositae are currently employed in traditional medicine. Preparations of Compositae are variously valued for their antibiotic, antifungal, antihelmintic, antiplasmodial, expectorant, sedative, diuretic, spasmolytic, haemostatic, immunostimulatory or anti-inflammatory properties. Wild Compositae species are also used as foods, fish poisons, fodder, and sources of oil and nectar. Industrial products for which Compositae (cultivated or wild) provide raw materials include insecticides, medicines, soaps, detergents, varnishes, paints, cosmetics, rubber, perfumes, food products, and flavourings and colourants for foods and drinks (Garg and Sastry 1996; Viswanathan and Singh 1996). Many Compositae are also highly valued as ornamentals in both commercial and recreational horticulture.

Among cultivated Compositae, the following are among the more important of over 260 species currently in cultivation for other than ornamental purposes: Cynara cardunculus, cardoon and globe artichoke, edible leaves and capitula; Carthamus tinctorius, safflower, edible and industrial oil; Cichorium intybus, chicory, leaf vegetable, roots provide a coffee substitute and also yield fructans; Cichorium endivia, endive, salad vegetable; Pterocypsela indica, Indian lettuce, salad vegetable; Lactuca sativa, lettuce, salad vegetable; Scorzonera hispanica, Spanish salsify, scorzonera, root vegetable; Acmella oleracea, Pará cress, culinary herb; Echinacea purpurea, purple coneflower, medicinal, immunostimulatory; Helianthus annuus, sunflower, edible and industrial oil and edible seeds, the most important crop plant of the family; Helianthus tuberosus, Jerusalem artichoke, root vegetable; Smallanthus sonchifolius, yacon, root vegetable; Artemisia dracunculus, tarragon, flavouring and oil used in perfumery; Tanacetum cinerariifolium, pyrethrum, yields the insecticidal monoterpenes called pyrethrins; Glebionis coronarium, crown daisy, green vegetable; and Petasites japonicus, butterbur, green vegetable. Among the most important of the very numerous genera cultivated as ornamentals are Gerbera (Mutisieae, Barberton daisy); Dahlia (Coreopsideae/Heliantheae s.l., dahlia); Tagetes (Tageteae/Heliantheae s.l., French and African marigolds), Xerochrysum (Gnaphalieae, everlasting), Callistephus (Astereae, China aster), Symphyotrichum (Astereae, Michaelmas daisy), Chrysanthemum (Anthemideae, florist’s chrysanthemum) and Pericallis (Senecioneae, florist’s cineraria).

A number of Compositae are of negative economic significance, inasmuch as they are more or less noxious weeds of gardens, cultivated fields, pastures and plantations. Among the more important weedy genera are Acroptilon, Carduus, Cirsium, Centaurea and Onopordum (all Cynareae), Chondrilla (Cichorieae), Parthenium, Ambrosia, Chromolaena and Mikania (all Heliantheae s.l.), Chrysanthemoides (Calendulae) and Senecio (Senecioneae). Parthenium hysterophorus causes a contact dermatitis; the pollen of Ambrosia species is a cause of hay fever, and many Senecio species are highly hepatotoxic to grazing livestock.

Aquatic plants generally are uncommon among angiosperms, yet the aquatic habit has evolved independently over 200 times (Cook 1999). Although it is difficult to apply a definition of “aquatic” universally, several genera of Asteraceae Bercht. & J. Presl tribe Eupatorieae Cass. contain species that clearly tolerate inundated soil conditions as well as periods of complete immersion, at least of their vegetative tissues. Worldwide, Cook (1999) recognized 18 Asteraceae genera (e.g. Bidens L., Cotula L., Hydropectis Rydb.), including four Eupatorieae genera (Gymnocoronis DC., Sclerolepis Cass., Shinnersia R. M. King & H. Rob., and Trichocoronis A. Gray), as including aquatic species, and he hypothesized that the latter taxa together represented a single origin of the aquatic habit in the family.

Key differences from similar families

Close examination will place all members of the Compositae within the family without any great difficulty; however, superficial examination may well place some taxa, albeit very rarely, into other families; both vegetative and fertile material of Ichthyothere is sometimes misplaced in the Commelinaceae; material from other families is often placed in the Compositae, most notably members of the Acanthaceae, Amaranthaceae, Labiatae and sometimes Rubiaceae; all of these families are instantly thrown out because they have:

Free stamens.

Capitate stigmas.

Typically distinct calyces.

No pappus -like structure.

Very distinctive fruiting structures.

SYSTEMATIC a probable topology of Asteraceae is the following: [Barnadesioideae + [Famatinanthoideae + [Mutisioideae + Stifftioideae + [Wunderlichioideae + [Gochnatioideae + [Hecastocleidodoideae + [Carduoideae + [Pertyoideae + [Gymnarrhenoideae + [Cichorioideae + [Corymbioideae + Asteroideae]]]]]]]]]]]; of these subfamilies, Hecastocleidoideae (1/1, S Nevada and adjacent California), Pertyoideae (4/95–100, Afghanistan, Himalayas to Japan and SE Asia, W Malesia to Philippines), Gymnarrhenoideae (1/1, North Africa, SW Asia) and Corymbioideae (1/9, W and E Cape) do not occur in South America.

1. SUBFAMILY BARNADESIOIDEA (10/c 92) ▸ all genera occur in South America; shrubs or trees; stems often with axillary spines; leaves alternate, apex mucronate or spiny; capitula usually homogamous and discoid, rarely heterogamous and disciform; phyllaries imbricate, multiseriate; receptacle paleaceous or epaleaceous; corollas actinomorphic or slightly zygomorphic or bilabiate, throat pubescent inside, corolla-lobes long, erect, ascending or recurved, densely long pubescent towards apices; flower colour yellow, white red or violet; anther-bases short-sagittate or conspicuously tailed; style arms short, obtuse; achenes usually brown; pappus of uniseriate plumose hairs.

1. Arnaldoa Cabrera. Shrubs, 1–4m; stems erect, with axillary spines; leaves alternate. 4 spp. endemic to northern Peru (3) and southern Ecuador (1), in xerophytic habitats.

2. Archidasyphyllum (Cabrera) Saavedra e P.L.Ferreira. Trees, gynodioecious. Two spp. endemics to Chile and adjacent areas in Argentina.

3. Barnadesia Mutis. Shrubs or trees, 0.6–20m. Stems erect, spiny. 23 spp. from the E Andes in Colombia southwards into NW Argentina and SE Brazil (only B. caryophylla (Vell.) S.F. Blake, also in Bolivia).

4. Chuquiraga Juss. Intricately branched shrubs, 0.25–2m; stems erect or compressed into cushions, often spiny. 24 spp. from the Andes of Colombia south into Argentina (15) and Chile, frequently in xeromorphic habitats.

5. Dasyphyllum Kunth. Shrubs or trees, 0.5–20m; stems erect or decumbent, with or without spines. 32 spp., distributed from Venezuela south to Argentina (5) to Brazil (16, 11 endemics, 4 of then are rare species in Brazil by Plantas Raras do Brasil’s book); absent in almost all Argentina, Amazon rainforest and Pacific Coasts.

6. Doniophyton Wedd. Herbs with some secondary growth, 2.5–8 cm. Stems ascendent or decumbent, fasciculate spines present or absent. Two spp. in N Chile and Patagonian Argentina.

7. Duseniella K. Schum. Annual herbs, unarmed, to 10 cm; leaves basally opposite, succulent. Only one sp., D. patagonica K. Schum., endemic to Patagonian Argentina.

8. Fulcaldea Poir. ex Lam. Shrubs or small trees to 8 m in height; trunks to 25 cm, usually with long paired axillary spines; inflorescences of corymbose or paniculate cymes, terminal and axillary; capitula small, sessile or subsessile, 1-flowered; florets hermaphrodite, actinomorphic, 5-merous, corolla tubular, lavender, pale pink or red. Two spp., F. laurifolia Poir. ex Lam., endemic to southern Ecuador and NW Peru, and F. stuessyi Roque & V.A.Funk, known only from the type collection, this species is restricted to Bahia occurring in seasonally deciduous forest from Diamantina highs in Bahia state, Brazil.

9. Huarpea Cabrera. Subshrubs, unarmed, perennials, c. 5 cm; leaves alternate, subrosulate, sessile. Only one sp., H. andina Cabrera, endemic to San Juan Province of Argentina.

10. Schlechtendalia Less. Perennial herbs, rhizomatous succulent, unarmed, to 1m; leaves basally rosulate, on stems opposite. Only one sp., S. luzulaefolia Less., from Rio Grande do Sul state in S Brazil, Uruguay and NE Argentina.

2. SUBFAMILY FAMATINANTHOIDEAE (1/1) ▸ a single genus and species.

11. Famatinanthus Ariza & S. E. Freire. Branched shrubs, young stems pubescent, old stems subglabrous, peridermis black with age; leaves opposite, simple, sessile; capitula solitary at tip of branches, homogamous, radiate, sessile; involucre campanulate; phyllaries 3-seriate, imbricate, gradate; receptacle naked; florets 10 or 11, dimorphic. Only one sp., F. decussatus Ariza & S. E. Freire, endemic to Sierra de Famatina, NW Argentina, 1,800 – 2,500m elevation range.

3. SUBFAMILY MUTISIOIDEAE (45/630–635) ▸ three tribes, all in South America. Herbs, shrubs or trees. Stems unarmed. Leaves alternate, usually unarmed, rarely apically spiny. Capitula radiate, subradiate or discoid, homogamous or heterogamous; phyllaries usually imbricate, multiseriate; receptacle usually naked. Corollas often bilabiate, sometimes regular or ligulate; corolla-lobes long; flower colour white, yellow; anther-bases often conspicuously tailed; style arms short obtuse, or truncate and fringed. Achenes usually brown; pappus usually of simple hairs, rarely plumose.

3.1 MUTISIOIDEAE ▸ TRIBE ONOSERINAE (7/42) - all genera in South America.

12. Aphyllocladus Wedd. Almost leafless, well-branched odoriferous shrubs; leaves alternate, simple, sessile, minute, rapidly falling, linear-spathulate, entire; capitula solitary, terminal rarely in few-headed scorpioid-like cymes, radiate or disciform; florets several to many (c. 10–40), heterogamous or homogamous; corollas lilac to purple. 4 spp., S Bolivia (1), N Chile, NW Argentina.

13. Gypothamnium Phil. Moderately branched glabrous shrubs; leaves spiralled, often ascending, simple, linear, somewhat fleshy, entire; capitula terminal, solitary, radiate, medium to large; florets heteromorphic, heterogamous, marginal florets uniseriate, female, spreading, ± reflexed; disc florets numerous, hermaphrodite; corollas glabrous, purple or pinkish-purple. Only one sp., G. pinifolium Phil., endemic to a small dry coastal region in N Chile.

14. Lycoseris Cass. Dioecious subshrubs or shrubs, usually scandent; leaves alternate, simple; Capitula solitary, terminal or few to several in corymbs or racemes, large, many-flowered, female often considerably larger than male; Florets usually numerous, heteromorphic; corollas orange to orange-red, sometimes yellow or violet; ray florets uniseriate, sterile; disc florets actinomorphic, relatively short 5-lobed. 11 spp. from Guatemala to Bolivia, Colombia, Brazil and Venezuela; in Brazil occur only L. boliviana Britton in Mato Grosso and Mato Grosso do Sul states and Bolivia; 10 spp. in South America.

15. Onoseris Willd. Annual or perennial herbs, subshrubs or suffrutices; leaves radical or alternate, capitula solitary or few to many in panicles, usually radiate, rarely discoid, erect; ray florets, when present, uniseriate, female; corollas violet or purple, glabrous; disc florets hermaphrodite, fertile, numerous; corollas usually yellow, reddish or purple, tubular. 32 spp., Mexico and Guatemala to South America (29; Colombia to Argentina, absent in Chile), one sp. in Brazil, O. brasiliensis Cabrera, endemic, colected in western outcrops in Mato Grosso state, yellow flowers.

16. Paquirea Cabrera. Branched shrubs, glabrous in old stem; leaves alternate, lamina oblanceolate, spinose-dentate with few spines, apices spiny; capitula solitary, long-pendunculate, axillary, or two capitula in divaricately branched axillary inflorescence, discoid, homogamous, slightly nodding before anthesis but erect in flower; florets actinomorphic, hermaphrodite, several; corollas 5-lobed, yellow. Only one sp., P. lanceolata (H. Beltrán & Ferreyra) Panero & S.E. Freire, endemic to Arequipa, Peru.

17. Plazia Ruiz & Pav. Shrubs, often resinous; leaves spiralled, lamina ovate or oblong, entire; capitula solitary, terminal, surrounded by leaves, radiate; involucre campanulate; florets numerous, hermaphrodite, all fertile; corollas white to pink, glabrous, marginal florets bilabiate, inner florets actinomorphic, deeply divided. 4 spp., Peru (3, two endemics), Bolivia, Argentina, Chile.

18. Urmenetea Phil. Subshrub or perennial herb; leaves alternate and loosely rosulate; capitula terminal on scape, radiate, homogamous; ray florets uniseriate, female; corollas bilabiate, white or pink, outer lip; disc florets numerous, hermaphrodite, fertile; corollas yellow, actinomorphic, short 5-lobed, lobes erect to slightly spreading; basal anther appendage caudate, short-pilose. Only one sp., U. atacamensis Phil., Chile and Argentina.

3.2 MUTISIOIDEAE ▸ TRIBE MUTISIEAE (15/c 280) - outsiders Amblysperma (2; W Australia), Gerbera (c 40; NE, tropical and S Africa, Madagascar, S Asia to China), Oreoseris (12; E Türkiye, Armenia, Central Asia, Himalayas, China, N Thailand and Vietnam), Leibnitzia (7; S and E Asia, Arizona, New Mexico, N Mexico), Perdicium (2; W Cape).

■ UNPLACED GROUP

19. Chaetanthera Ruiz & Pav. Erect or prostrate annual or perennial herbs or subshrubs, monoecious, rarely dioecious; leaves opposite and decussate with connate bases; capitula terminal, solitary, sessile, rarely 2 or 3 in a cyme, radiate or disciform, small to medium; ray florets female; corollas white, yellow, rarely orange or reddish; disc florets hermaphrodite or female; corollas yellow. 31 spp., Peruvian Altiplano (one species) along the Andean cordillera, and throughout Chile to Valdivian rainforest starts, mainly Argentina (20 of then) and Chile; only 4 outside Cono Sur.

20. Oriastrum Poepp. & Endl. 16 spp., from Peruvian Altiplano in Lima and Junín, in the Altiplano to the south of Lake Titicaca in Peru, Bolivia, Chile and Argentina (summer rains), and in the high elevation Altoandino habitats along the Chilean and Argentinean Andes, confined to high elevation habitats found above 2,000m. and most typically between 3,000 – 5,000m.

■ GROUP 1

21. Brachyclados D. Don. Dwarf caespitose or lax shrubs, sometimes cushions; leaves alternate, simple; capitula solitary, terminal, sessile or short-pedicellate, radiate, heterogamous; florets numerous, marginal female, disc hermaphrodite; corollas yellow, ofmarginal florets bilabiate, corollas of disc florets bilabiate but lacking markedly enlarged limb. Three spp., Chile and Argentina.

22. Lulia Zardini. Herbs caulescent, perennial; leaves alternate; sessile; blades linear, graminiform, long, parallel-veined, margin entire, glabrous; capitula solitary, terminal on scapes, radiate; Ray florets uniseriate, female; corollas yellow or orangish, glabrous, disc florets hermaphrodite; corollas yellow. Only one sp., L. nervosa (Less.) Zardini, endemic to Sphagnum bogs in southern Brazil (São Paulo to Santa Catarina states).

23. Trichocline Cass. Perennial scapigerous herbs or shrubs, rarely caespitose; leaves rosulate; capitula solitary on scapes, radiate, erect; ray florets female, uniseriate, yellow to orangish yellow or rarely reddish, outer lip conspicuous, spreading, apices very short 3-toothed, inner of two long linear spiralled lobes; disc florets numerous, hermaphrodite; corollas bilabiate, outer lip short 3-toothed, inner of two short linear lobes. 23 spp., 21 in Argentina, Bolivia, Brazil (9, one endemic), Chile, Paraguay, Uruguay, and Ecuador and Peru.

■ GROUP 2

24. Chaptalia Vent. (inc. Gerbera) Scapigerous perennial, rarely annual, herbs; leaves rosulate, simple, often relatively few; capitula solitary, terminal on scapes, heterogamous, nodding at first becoming erect in flower and fruit, sometimes nodding in fruit or always erect; corollas usually white or creamy white, rarely purplish, sometimes with purplish midstripe beneath limb. 65 spp., southern U.S.A., Caribbean, Central and South America, 29 in South America, 14 in Brazil, 8 endemics; includes C. hieracioides (Kunth) X.D.Xu & W.Zheng. (ex-Gerbera hieracioides (Kunth) Zardini) from Peru and Ecuador.

■ GROUP 3

25. Adenocaulon Hook. Perennial herbs; leaves alternate or forming basal rosette; capitula many in lax panicles; capitula small, few-flowered, disciform, heterogamous, hemispherical, erect; corollas actinomorphic or slightly zygomorphic (bilabiate), whitish or yellowish-white, staminodes present, style bifid; central florets few, hermaphrodite, functionally male, usually lacking achene, corollas actinomorphic, 5-lobed, style undivided. 5 spp. highly disjunct: A. chilense Less. in Argentina and Chile, one in Mexico and Guatemala, two from in Asia, and one in U.S.A. and Canada.

26. Eriachaenium Sch. Perennial rhizomatous herbs; leaves alternate, lamina obovate to elliptic, undulate, sometimes serrate; capitula solitary, axillary, small, sessile or on short-bracteolate side shoots; marginal florets female, few; corollas white (pinkish in some herbarium material), actinomorphic; central florets very few, hermaphrodite, functionally male; corollas white, actinomorphic. Only one sp., E. magellanicum Sch. Bip., Argentina and Chile (Patagonia and Tierra del Fuego).

■ GROUP 4

27. Mutisia L. f. Plants perennial subshrubs or shrubs, often climbers; leaves simple, apices with or without terminal tendril, rarely deeply pinnatisect, coarsely lobed or deeply partite, or pinnately compound with few to several pairs of leaflets and rachis alwayswith a terminal simple; capitula small to large, solitary, leaf-opposed or terminal, erect or pendulous, ‘subligulate’ and disc florets hermaphrodite and bilabiate; corollas yellow, orange, pink, purple or white, bilabiate. 65 spp., Argentina (20), Brazil (4, 2 endemics), Bolivia, Chile, Colombia, Ecuador, Paraguay, Peru, Uruguay.

28. Pachylaena D.Don ex Hook. & Arn. Prostrate, rhizomatous rosulate herbs or subshrubs; leaves simple; capitula solitary, terminal, sessile to subsessile, radiate, large; florets heteromorphic, marginal florets female, disc florets hermaphrodite; corollas of marginal florets yellowish-red to pink, bilabiate, outer lip an enlarged 3-toothed limb, corollas of disc florets yellow, bilabiate. Two spp., from Andes in Argentina and Chile.

3.3 MUTISIOIDEAE ▸ TRIBE NASSAUVINAE (24/310–315) - outsiders Berylsimpsonia (2; the Greater Antilles) and Acourtia (c 85; SW U.S.A., Mexico, Central America, the Caribbean).

29. Ameghinoa Speg. Low, densely branched shrub; leaves often in dense clusters on brachyblasts, simple, coarsely dentate; capitula fewin terminal clusters, short-pedicellate, homogamous; florets hermaphrodite, isomorphic, 20–30; corollas yellow. Only one sp., A. patagonica Speg., endemic to Argentina (Patagonia).

30. Burkartia Crisci. Dwarf caespitose shrub forming hemispherical cushions; leaves sessile, densely spiralled, acicular, margins strongly revolute; capitula solitary, terminal, usually appearing sessile although shortly pedunculate, few-flowered, radiate, homogamous; florets hermaphrodite, fertile; corollas white. Only one sp., B. lanigera (Hook. & Arn.) Crisci, Argentina (Patagonia).

31. Calopappus Meyen. Only one sp., C. acerosus Meyen, Argentina (Patagonia).

32. Cephalopappus Nees & Mart. Perennial rosulate stoloniferous herbs; leaves with ovate to narrowly obovate lamina, coarsely dentate; capitula solitary, scapose, rarely scape branched with two (or few) capitula, homogamous; florets many, hermaphrodite, all fertile; corollas white, few opening at a time and lost very rapidly, leaving green achene, bilabiate when open, lobes spreading. Only one sp., C. sonchifolius Nees & Mart., endemic to Brazil, in dense shade of the Atlantic Forest from Bahia to Rio de Janeiro, in relatively dense shade of forest floors or on mossy banks.

33. Criscia Katinas. Perennial herb or subshrub; leaves rosulate, lamina obovate to broadly obovate, entire; capitula solitary or 2–4 on scapes, large, bilabiate, homogamous; involucre hemispherical or broadly campanulate. Only one sp., C. stricta (Spreng.) Katinas, Argentina, Rio Grande do Sul state in S Brazil, and Uruguay.

34. Dolichlasium Lag. Small, moderately branched low shrub; leaves alternate, petiolate, lamina pinnatisect, pinnae opposite, ovate; capitula large, terminal, solitary, homogamous, erect; florets hermaphrodite, numerous, homogamous, fertile; corollas yellow. Only one sp., D. lagascae D. Don, Argentina (Patagonia).

35. Holocheilus Cass. Perennial rosulate herbs; leaves usually few, loosely rosulate, lamina medium and coarsely dentate, crenate or entire, or large and pinnatisect or coarsely lobed; capitula in large, few- to many-headed terminal corymbs or cymes, rarely scapose, homogamous, small to medium, erect; florets hermaphrodite; corollas white. 7 spp., Bolivia, Argentina, Brazil (6, 2 endemic), Paraguay, Uruguay. H. monocephalus Mondin from highlands of Rio Grande do Sul state is considered a rare species in Brazil by Plantas Raras do Brasil’s book.

36. Jungia L. f. Perennial herbs, rarely rosulate, subshrubs, shrubs or lianes, sometimes with xylopodium; leaves alternate, lamina cordate, usually lobed, entire, serrate, dentate or crenate; capitula in usually terminal corymbs or panicles, sometimes with dense glomerules, homogamous, appearing radiate by enlarged outer lip of marginal florets; florets few to many, hermaphrodite, all fertile; corollas white, violet pink, purple or rarely yellow. 30 spp., 29 in South America, some up to Central America (one endemic to Mexico); only two species in Brazil, none endemics.

37. Leucheria Lag. Annual or perennial herbs; leaves alternate, rosulate, lamina linear, narrowly lanceolate, spathulate, oblong or ovate, entire, dentate, coarsely lobed, pinnatisect or pinnatifid; capitula solitary or in few- to many-headed corymbs or panicles, pedicellate, appearing radiate; florets several to many, homomorphic, hermaphrodite, fertile, outer lip white, pink, lilac or wine-coloured. 40 spp. from Argentina and Chile, one up to Bolivia and Peru.

38. Leunisia Phil. Viscid low ‘subshrub’ or perennial herb; leaves alternate to loosely spiralled, irregularly toothed or sometimes entire; capitula terminal, solitary, homogamous, discoid; florets numerous, hermaphrodite, fertile; corollas yellow. Only one sp., L. laeta Phil., endemic to Chile.

39. Lophopappus Rusby. Well-branched shrubs; stems often viscous; leaves alternate or in dense axillary clusters, simple, often viscous, capitula solitary or in tight clusters at apices of branches, sessile or short-pedicellate, homogamous; florets usually few, yellowish white to white, usually sweet-smelling, hermaphrodite, fertile. 4 spp., Argentina, Bolivia, Chile, Peru.

Lophopappus is rather similar to Proustia but differs in its solitary or few, grouped capitula, the lack of terminal spines on short branches, to some degree corolla colour (white in Lophopappus, pink or purple in Proustia). The recent treatment of genera for the Flora of Peru (Ferreyra 1995) has suggested that Lophopappus be treated as congeneric with Proustia. They are treated as separate genera here.

40. Macrachaenium Hook. f. Perennial herb; leaves mostly in loose basal rosette, alternate, lamina oblong, ovate or broadly ovate, coarsely and deeply, often irregularly, lobed, sometimes pinnatifid to almost runcinate-pinnatifid, often irregularly subdentate; capitula solitary on scapiform peduncles, homogamous, usually discoid, sometimes conspicuously radiate, apparently nodding; florets bilabiate, all hermaphrodite and fertile; corollas white. Only one sp., M. gracile Hook. f., Argentina and Chile.

41. Marticorenia Crisci. Shrub with short woody caudex; leaves alternate, lamina ovate, lobulate, becoming lanceolate above; capitula in a lax many-headed corymb, homogamous; florets hermaphrodite; corollas violet-pink. Only one sp., M. foliosa (Phil.) Crisci, endemic to Chile.

42. Moscharia Ruiz & Pav. Annual odiferous herbs; leaves alternate, simple, lamina elliptic, entire, coarsely dentate or lobed to pinnatisect; capitula in lax, terminal corymbs, discoid, homogamous; florets few, hermaphrodite, bilabiate; corollas pink or violet. Two spp., endemic to Chile.

43. Nassauvia Comm. ex Juss. Perennial herbs, subshrubs or shrubs, often compact and caespitose cushions; leaves alternate, sessile, usually densely crowded, capitula generally in complex, terminal, often dense, sometimes globular synflorescences, rarely solitary or in few-headed dichasia, rarely with solitary shortpedicellate axillary capitula forming a terminal ‘spike’, capitula sessile or subsessile, homogamous; involucre cylindrical; florets few, (2–4–)5, hermaphrodite; corollas white, rarely violet-pinkish or yellowish. 39 spp. from Argentina (inc. Falkland Islands) and Chile, one also in Bolivia.

44. Oxyphyllum Phil. Erect shrub, each leaf axil with a dense cluster of simple, linear, spine-tipped immature leaves; stem leaves pinnatifid, rarely entire, segments and apices spiny; capitula in a terminal, dense, few- to many-headed corymb, homogamous, discoid; florets few, dimorphic, outer florets apparently sterile and each subtended by an internal palea, inner florets fertile; corollas pinkish-white. Only one p., O. ulicinum Phil., endemic to Chile (Atacama Desert).

45. Panphalea Lag. Slender annual or perennial rhizomatous herbs, sometimes with xylopodium; leaves mostly basally rosulate, alternate, entire and linear-lanceolate to orbicular, coarsely lobed or lyrate-pinnatifid; capitula in lax, relatively few-headed corymbs, small, appearing radiate (although all florets identical), homogamous; florets hermaphrodite, many; corollas white. 7 spp., Argentina, Brazil (6, 3 endemic), Paraguay and Uruguay. P. ramboi Cabrera from highlands of Rio Grande do Sul state is considered a rare species in Brazil by Plantas Raras do Brasil’s book.

46. Perezia Lag. Perennial, usually strongly rosettiform, sometimes caespitose herbs, rarely tall leafy-stemmed herbs; leaves simple, radical or alternate, entire or lyrate, capitula on 1–(2)-headed scapes arising from basal rosette, or in few- to many-headed dense or lax and spreading panicles; capitula appearing radiate but being discoid, homogamous, usually erect, rarely nodding; florets usually several to many (8–40); corollas bilabiate, yellow, blue, purple, violet, red, or crimson, rarely white or cream, occasionally with outer lip of one colour and inner of another (usually yellow). 38 spp., 37 in Argentina, Bolivia, Brazil (4, 2 endemic), Chile, Colombia, Ecuador, Paraguay, Peru and Uruguay, and 1 endemic to Mexico.

47. Pleocarphus D. Don. Shrub; leaves sessile, linear, entire, margins conspicuously revolute; capitula many to numerous in an elongated panicle, pedicellate, homogamous; florets homomorphic, hermaphrodite, fertile; corollas yellow. Only one sp., P. revolutus D. Don, endemic to N Chile.

48. Polyachyrus Lag. Decumbent, scandent or prostrate subshrubs or shrubs, rarely herbs; leaves alternate, lamina pinnate-lobed or pinnatisect, entire or coarsely dentate; capitula in solitary apical glomerules or glomerules in pseudocorymbs, each subtended by one bract; capitula numerous, sessile, 2- or rarely 3-flowered; florets hermaphrodite, fertile, sweet-smelling. 7 spp., all in Chile, 3 up to Peru.

49. Proustia Lag. Scandent or erect shrubs or rarely small trees, stems unarmed or spiny; leaves simple, alternate, capitula racemose or paniculate, erect or pendent, pedunculate or sessile, florets few, hermaphrodite, sweet-smelling; corollas bilabiate, pink or purple. 4 spp., Peru (3, 2 endemics), Bolivia, Chile and Argentina.

50. Quelchia N.E.Br. Poorly branched shrubs or small trees; leaves simple, alternate, sometimes densely clustered towards branch apices; Capitula in dense terminal or subterminal cymes of glomerules, single-flowered, homogamous; Florets hermaphrodite, usually actinomorphic; corollas red or white to cream. 4 spp., 3 endemic to Pantepui Life Zone, Auyán-tepui and Chimantá, in Guiana Shield of southern Venezuela, at 1,600 – 2,800m elevation range, and Q. conferta also in mountains of Guyana.

51. Spinoliva G.Sancho, Luebert & Katinas. Shrubs or small trees, young branches unarmed, subglabrous, glandular with minute glands, older branches glabrous; leaves alternate, sessile, blades obovate to elliptic, coriaceous, margin spiny, glabrous or tomentose below; capitula in thyrses, those of secondary axes in spiciform arrangements; capitula sessile, homogamous, discoid. Involucre cylindrical, phyllaries 4-seriate, imbricate, coriaceous. Only one sp., S. ilicifolia (Hook. & Arn.) G. Sancho, endemic to C Chile.

52. Triptilion Ruiz & Pav. Annual or perennial herbs or subshrubs; leaves alternate, in basal rosettes in some herbaceous plants, lamina entire or pinnatisect with denticulate spiny margins; capitula densely aggregated into paniculate glomerules, sessile to shortpedicellate, few-flowered, discoid, homogamous; florets hermaphrodite; corollas glabrous, white or blue. 7 spp., Argentina and C Chile.

53. Trixis P. Browne. Perennial herbs, subshrubs, shrubs, scandent/trailing shrubs or small trees, sometimes with stem tubers; leaves alternate, lamina simple, narrowly lanceolate, elliptical, oblanceolate or obovate to oblong, entire or often denticulate; capitula in usually terminal, sometimes axillary, lax cymes, corymbs or panicles, occasionally aggregated into pseudoglomerules, homogamous; florets few to many (5–c. 70), hermaphrodite, all fertile; corollas yellow to orange, rarely white. 45 spp., 26 in South America, in Argentina, Bolivia, Brazil (16, 9 endemics), Chile, Colombia, Costa Rica, Cuba, Ecuador, El Salvador, Guatemala, Honduras, Mexico, Nicaragua, Panamá, Paraguay, Peru, Uruguay, U.S.A., Venezuela, Caribbean.

4. SUBFAMILY STIFFTIOIDEAE (9/41) ▸ outsiders Salcedoa (1; Hispaniola).

54. Achnopogon Maguire, Steyermark &Wurdack. Shrubs or small trees; leaves densely spiralled or rosulate; capitula solitary, sessile, axillary or in few-headed cymes or in axillary cymes at apices of long flowering branches, homogamous, bilabiate; involucres narrow-cylindrical; phyllaries c. 4-seriate, imbricate, gradate; receptacle small, epaleaceous, naked; florets few (2–6), hermaphrodite; corollas white, zygomorphic. Two spp., endemic to Auyan-tepui and Chimantá-tepui, southern Venezuela (Guiana Shield), 1,800 – 2,500m elevation range.

55. Dinoseris Griseb. Only one sp., D. salicifolia Griseb., from Bolivia and N Argentina.

56. Duidaea S.F. Blake. Shrubs (with extremely woody bases) or dwarf trees; leaves alternate or densely spiralled, simple; Capitula usually solitary, axillary or subterminal, or on medium-length pedicels, homogamous; involucre hemispherical or cylindrical to campanulate; Florets few to many (8–24), bilabiate, hermaphrodite, fertile; corollas white, red or reddish purple. 4 spp., endemic to the Guiana Shield of Venezuela (Venezuelan Guyana), at 1,000 – 2,800m elevation range.

57. Eurydochus Maguire &Wurdack. Trees or treelets; leaves simple, alternate, usually in terminal clusters at stem apices; Capitula solitary, subterminal on long sparsely bracteolate pedicels, homogamous; involucre hemispherical to campanulate; phyllaries imbricate, few-seriate (c. 6–8), subequal; receptacle broad, naked, convex; florets numerous (40–50), hermaphrodite; corollas bilabiate, red. Only one sp., E. bracteatus Maguire & Wurdack, Amazonas state in N Brazil and Venezuela (Guiana Shield).

58. Glossarion Maguire &Wurdack. Shrubs or small trees; leaves alternate, short petiolate; capitula solitary, axillary, homogamous, discoid or ligulate; involucre turbinate to narrowly campanulate or cylindrical; florets bilabiate or ligulate, bilabiate corollas, ligulate corollas usually with limb tightly rolled in apical portion; corollas rose-coloured to orange-red. Two spp., endemic to Mount Neblina, in Amazonas state in northern Brazil (only G. bilabiatum (Maguire) Pruski) and southern Venezuela (both) in the Guiana Shield, 1,600 – 2,800m elevation range.

59. Gongylolepis R.H. Schomb. Small to large shrubs or trees; leaves simple, alternate or densely spiraled; Capitula solitary, sessile or pedicellate, or few to many corymbose to subumbellate, homogamous, few to many-flowered (6–150), large; Florets hermaphrodite; corollas bilabiate, white or pale yellow, sometimes yellowish or reddish. 14 spp., N Brazil (2, none endemic), Colombia, Guyana and Venezuela (9 endemics). G. colombiana Maguire is a Andean sp. of another otherwise enterily Guyana group of mutisioid composites, occurring on sandstone in Venezuela´s Tachira state and adjacent areas across the Colombian border.

60. Hyaloseris Griseb. Shrubs or small trees; leaves opposite, simple; capitula two to several subsessile in dense terminal or axillary clusters or terminal, homogamous, few- or many-flowered, usually appearing ligulate. 6 spp., Argentina and Bolivia.

61. Neblinaea Maguire &Wurdack. Poorly branched shrubs or trees/treelets; leaves alternate to ± densely spiralled, pseudopetiolate, oblanceolate, entire; capitula solitary or few to several in cymes, homogamous, bilabiate; florets few (2–5), hermaphrodite, fertile; corollas white, glabrous. Only one sp., N. promontorium Maguire & Wurdack, endemic to the Guiana Shield of Venezuela and neighbouring Amazonas state in N Brazil, 1,200 – 2,100m elevation range.

62. Stifftia J. C. Mikan. Shrubs to small trees, sometimes vines, large imbricate involucres, long actinomorphic corollas with strongly coiled lobes. 6 spp., 5 spp. found in the tropical forest of E Brazil with two additional species found in the rainforests of northern Brazil and one restricted of French Guiana

5. SUBFAMILY WUNDERLICHIOIDEAE (8/44) ▸ two tribes, both in South America.

5.1 WUNDERLICHIOIDEAE ▸ TRIBE WUNDERLICHIAE (4/38) - all genera in South America.

63. Chimantaea Maguire, Steyerm. &Wurdack. Small shrubs or rather low trees/treelets (to c. 9m); leaves sessile or pseudopetiolate, spiraled; Capitula solitary, sessile, terminal, discoid, homogamous; Florets hermaphrodite, few to many (7– 35, rarely to 100); corollas actinomorphic, 5-lobed, yellowish or yellowish-green, lobes stiff, very long, erect; basal anther appendages caudate, entire or scarcely ‘erose’/pilose. 9 spp., endemic to in tepuis of Aprada, Auyan, Cerro El Sol, Chimantá, Murisipan in Guianan Shield in Venezuela, at 1,600 – 2,800m elevation range.

64. Stenopadus S.F. Blake. Trees or shrubs; leaves alternate or loosely spiralled, simple, oblanceolate or round, entire; capitula solitary, terminal or rarely in few-headed cymes, homogamous, discoid; florets few to many (5–100), actinomorphic, hermaphrodite; corolla magenta. 15 spp., endemic to the Guiana Shield of Brazil (2, one endemic), Colombia, Venezuela (7 endemics), except by one from southern Ecuador.

65. Stomatochaeta (S.F. Blake) Maguire &Wurdack. Trees, treelets (or possibly shrubs); leaves alternate or sometimes pseudowhorled, simple, sessile or pseudopetiolate; Capitula solitary, terminal, often surrounded by a pseudowhorl of leaves, homogamous, discoid; Florets few to many, hermaphrodite, all fertile; corollas actinomorphic, cream-coloured, corolla lobes stiff, erect, long. 6 spp., endemic to the Guiana Shield of Venezuela (5 endemics), Roraima state in N Brazil and Guyana (only S. condensata (Baker) Maguire & Wurdack), 700 – 2,800m elevation range.

66. Wunderlichia Riedel ex Benth. & Hook. f. Shrubs or small trees very stout, virgate, pyrrophytic, with large, homogamous capitula with actinomorphic corollas produced at the end of the dry season on leafless stems. 5 spp., endemic to highlands of central Brazil (states of Mato Grosso and Tocantins to São Paulo and Espírito Santo), the gross morphology of Wunderlichia (tree-like habit, coriaceous, caducous, densely pubescent leaves) is distinctive among genera included in classical Mutisieae, and it appears to be the result of adaptation to the seasonally dry conditions of the Campo Rupestre of central Brazil; three species from Bahia, Rio de Janeiro and Mnas Gerais states are considered rare species in Brazil by Plantas Raras do Brasil’s book. Phytomelanins are only found in Asparagales seeds and in Asteraceae fruits of Heterocoma DC., Wunderlichia Riedel ex Benth, and some Asteroideae.

5.2 WUNDERLICHIOIDEAE ▸ TRIBE HYALIDAE (4/6) - outsiders Leucomeris (2; Himalayas, Yunnan, Burma, Thailand, Vietnam), Nouelia (1; SW China)

67. Hyalis D. Don ex Hook. & Arn. Rhizomatous shrubs; leaves alternate, simple, capitula few in corymbs, radiate, pedicellate; florets hermaphrodite, 5–6, fragrant; corollas glabrous, usually pink, sometimes white or purplish, marginal florets 4–5, central floret solitary, actinomorphic, deeply divided. Two spp., Bolivia, Paraguay, Argentina.

68. Ianthopappus Roque & D.J.N. Hind. Subshrub; leaves alternate; Capitula few to many in lax corymbs, radiate, heterogamous; Ray florets female, fertile; corolla glabrous, disc florets hermaphrodite, fertile, actinomorphic; corollas purplish, lobes revolute; basal anther appendages caudate, long-attenuate, margins pilose. Only one sp., I. corymbosus (Less.) Roque & D.J.N. Hind, northern Argentina, Rio Grande do Sul state in S Brazil, and Uruguay.

6. SUBFAMILY GOCHNATIOIDEAE (9/90) ▸ outsiders Nahuatlea (7; S Arizona and Texas, Mexico), Anastraphia (33; Mexico, Cuba, Hispaniola, the Bahamas), Tehuasca (1, Mexico).

69. Cnicothamnus Griseb. Shrubs, florets red; Two spp. from Bolivia to Paraguay and NW Argentina.

70. Cyclolepis D. Don. Gynodioecious spiny shrubs; leaves alternate, simple; capitula few to several on short side shoots, sessile, spicate, discoid, heterogamous; female florets few, fertile; corollas yellowish; Hermaphrodite florets few, fertile; corolla yellowish. Only one sp., C. genistoides D. Don, Argentina and Paraguay.

71. Gochnatia D. Don. 14 spp., one in Ecuador, remaining from Peru and Bolivia to N Argentina.

72. Moquiniastrum Kunth. Shrubs, sub-shrubs or trees, sometimes with xylopodium; leaves alternate, petiolate to sub-sessile, limb discolor, elliptic or rarely ovate or cordate, pubescent usually on abaxial face or less commonly on both faces, margin entire or serrate; capitula isomorphic or sub-dimorphic, arranged in usually leafy paniculiform or less commonly corymbiform synflorescences; cypselas cylindrical to cuneate, costate, sericeous. 22 spp., E Bolivia (2, one endemic), Argentina (3), E Brazil (20, 13 endemics), Paraguay (6), and Uruguay (1), and one up to Venezuela.

73. Richterago Kuntze. Subshrubs or shrubs, often single-stemmed, or small rosetes with long, scapose inflorescences; leaves alternate, rosulate, lamina linear, obovate or spathulate, entire or denticulate; capitula solitary on scapes or in few-headed panicles, radiate or discoid. 16 spp. endemics to E & C Brazil, mainly endemic to the Espinhaco Range of the mountains in Minas Gerais, but R. radiata (Vell.) N. Roque is the most widely distributed species, occurring in Tocantins, Mato Grosso, Goiás, Distrito Federal, Minas Gerais, São Paulo, and Paraná states.

74. Vickia N. Roque & G. Sancho. & Hook. f. Shrubs, 30–50 cm tall, branches cylindrical, sulcate, glabrous; leaves alternate, congested, petiole 2–4 mm long, lamina coriaceous, concolorous, 2.5–4.5(–5.0) × 2.5–3.5 cm, orbiculate to widely elliptic, base rounded to slightly cordate, margins denticulate, apex rounded to obtuse, glabrous on both surfaces, leaf venation actinodromous basal. Only one sp., V. rotundifolia (Less.) N. Roque & G. Sancho, endemic to E Brazil from Rio de Janeiro to Paraná state, possibly extinct.

6. SUBFAMILY CARDUOIDEAE - (89/3.160–3.285) ▸ four tribes, Dicomeae (8/c 113, Tropical and southern Africa, Madagascar, Socotra, Arabian Peninsula, Pakistan, India), Oldenburgieae (1/4, southern parts of W and E Cape) and Tarchonantheae (2/20, Tropical and southern Africa, Madagascar, Mascarene Islands, Arabian Peninsula) do not occur in South America. Among Cardueae (78/3.020–3.145), all outsiders are Macaronesia and Mediterranean to Eurasian and Africa except Cirsium (c 465; temperate regions on the Northern Hemisphere, Mediterranean), Arctium (14; temperate regions in the Old World, Rhaponticum (27; Tenerife, Mediterranean, temperate and subtropical Asia, E and SE Australia).

Herbs or rarely shrubs. Leaves alternate, often thistle-like; capitula homogamous, discoid or radiant; phyllaries imbricate, multiseriate, often spiny or appendaged; receptacle usually densely bristly or fimbriate; florets purple, white or yellow; corollas regular; corolla-lobes long; anther-bases tailed; style arms usually obtuse, short and with a thickened often hairy zone below them; achenes not black; pappus usually of hairs, sometimes plumose, or of scales.

75. Plectocephalus D. Don. (inc. Centaurodendron, Yunquea) Unarmed annual or perennial herbs, densely glandular, sometimes pachycaul tree with soft wood; leaves terminal on the stems; leaves with minutely denticulate, cartilaginous margins and veins, or very large (to 30 cm), obovate, with broad (alate) semi-amplexicaul petiole; margins serrate; capitula terminal, solitary or laxly corymbose, heterogamous. 15 spp., two in North America, one in Ethiopia and 12 in South America, 11 endemics to Chile and P. tweediei (Hook. & Arn.) N.Garcia & Susanna in Brazil (Rio Grande do Sul), Paraguay (Guairá), Uruguay, Argentina (Buenos Aires, Chaco, Misiones, Corrientes, Entre Ríos, Formosa).

7. SUBFAMILY CICHORIOIDEAE - (280/11,270-16,300) ▸ Cichorieae + [Eremothamneae + [Arctotideae + [Platycarpheae + [Liabeae + [Distephanus + Moquinieae + Vernonieae]]]]]]. Heterolepis (3), shrubs distributed mainly in the Cape provinces, has sometimes been assigned to Arctotideae, but its systematic affiliation is not resolved. 4 of 8 tribes absents in South America: Eremothamneae (2/3; W South Africa, southern Namibia), Arctotideae (14/195–210; tropical and S Africa, SE Australia, Tasmania, with their highest diversity in the Cape Provinces), Platycarpheae (2/3; southern Africa) and Distephaninae (1/c. 42, tropical and S Africa, Socotra, Madagascar, Mauritius, China).

7.1 CICHORIOIDEAE ▸ TRIBE CICHORIEAE (107/9,500 – 14,650) - 4 unplaced genera, outsiders of then are Ixeridium (16; E and SE Asia, Malesia to New Guinea) and Spiroseris (1; S. phyllocephala; Pakistan). 11 subtribes, 6 absents in South America: Warioniinae (1/1; NW Sahara), Scorzonerinae (10/290-310; North Africa, Europe, Mediterranean to W and C Asia, Armenia and Iran to Central Asia, Pakistan, Mongolia, Arabian Peninsula, NW China, southern Canada, U.S.A.), Scolyminae (4/c. 20; Mediterranean to Türkiye and Afghanistan, North Africa), Cichoriinae (6/c. 33; southern Europe, Macaronesia, Mediterranean to Yemen, North Africa, Ethiopia to South Africa and Madagascar, Zambia and Malawi, Sierra Nevada in California), Lactucinae (16/169-199; nearly cosmopolitan except tropical America) and Chondrillinae (3/c. 30; Europe, temperate Asia).

Several genera in Cichorieae are agamospermous, i.a. Taraxacum, with about 2,400 species, and Hieracium, with at least 2,900–5,500 and probably far more than 10,000 species. In many groups (e.g. in Pilosella) the species number is constantly in a state of flux, since the numerous species are not fixed apomicts, but mainly amphiapomictic and hybridize frequently.

■ SUBTRIBE HYOSERIDINAE ▸ 5/c. 210; Macaronesia, Mediterranean to Syria, Asia to SE Asia, New Zealand, Caribbean

76. Sonchus L. (inc. Thamnoseris) Rosette trees and shrubs with leaf rosettes at apex of stems, involucral bracts in several rows, receptacle naked or bristly, capitula variously arranged, sometimes solitary, florets whitish or orange-yellow. 97 spp., 85 in over over Old World from South Africa to Siberia, Ireland to Tasmania, and 12 spp., endemics to Juan Fernandez Islands (Chile).

■ SUBTRIBE HIERACIINAE ▸ outsiders Schlagintweitia (3; Pyrenees, Central Europe), Andryala (30; Mediterranean, Macaronesia, North Africa, the Middle East), Hispidella (1; the Iberian Peninsula).

77. Hieracium L. Perennial herbs with branched stocks, but without stolons, leaves and stems with branched and unbranched hairs, involucral bracts in several rows, receptacle naked, florets yellow or rarely reddish, style branches long, achenes cylindric, 10-ribbed, ribs apically confluent into an obscure ring, pappus of scabrid-barbellate fragile bristles in two rows. x = 9, diploids, triploids, tetraploids, pentaploids. 4,566 spp. (largest genus worldwide), from Eurasia, 130 in New World, South America (90); 5 spp. in Brazil, 4 endemics; numerous clones described as species. Hieracium belongs three subgenera:

§ subg. Chionoracium ▸ only of entirely sexual, diploid species, 129 spp., 39 spp. from North America to Panamá and 90 spp. in South America.

§ subg. Hieracium ▸ c. 4,300 spp., Eurasia, with the diploid species H. umbellatum L. up to North America; few sexual diploids (c. 20).

§ subg. Pilosella ▸ mainly distributed in Europe and West Asia with about 150 spp. and comprises a few native species occurring in NW Africa; few sexual diploids (also c. 20).

■ SUBTRIBE MICROSERIDINAE – outsiders Krigia (9; C and S Canada, U.S.A., N Mexico), Pinaropappus (7–10; S U.S.A., Mexico, Guatemala), Marshalljohnstonia (1; Coahuila in N Mexico), Shinnersoseris (1; S Canada, C and S U.S.A., N Mexico), Chaetadelpha (1; SW U.S.A.), Lygodesmia (9; SW Canada, U.S.A., Mexico), Pyrrhopappus (1–5; U.S.A., N Mexico); Glyptopleura (2; W U.S.A.), Nothocalais (4; SW Canada, W U.S.A.), Stebbinsoseris (2; California, Arizona, NW Mexico), Uropappus (3; SW Canada, W U.S.A., N Mexico), Atrichoseris (1; SW U.S.A., NW Mexico), Munzothamnus (1; San Clemente Island off California), Stephanomeria (18; SW Canada, W U.S.A., NW Mexico), Rafinesquia (2; SW U.S.A., NW Mexico), Pleiacanthus (1; W U.S.A.), Prenanthella (1; W U.S.A., N Mexico), Anisocoma (1; SW U.S.A., NW Mexico), Calycoseris (2; SW U.S.A., NW Mexico).

78. Agoseris Rafin. Perennial to annual herb, scapose, involucral bracts in 2–4 series, receptacle naked, florets yellow to redorange, achenes fusiform, tapered to beak, pappus of scabrid-barbellate bristles. x = 9, diploids and tetraploids. 11 spp., 9 in W North America (only one in Mexico), 2 restricted in Chile and Argentina.

79. Malacothrix DC. Perennial to annual herbs, involucral bracts in 3–6 series, receptacle naked or with fragile bristles, ligules yellow or white, achenes fusiform, truncate, pappus of slender rays. x = 9, 8, 7, diploids and tetraploids. 20 spp., W North America and Mexico, M. clevelandii A. Gray disjunct in Mexico, U.S.A. and southern South America in Argentina and Chile.

80. Microseris D. Don. Perennial to annual herbs, leaves mostly basal, involucral bracts in 2 to several series, receptacle naked, florets white to orange, achenes cylindric to fusiform, not beaked, with c. 10 ribs, pappus of 5 to many narrowly lanceolate, bristle-tipped, awn-like scales. x = 9, diploids and tetraploids. 13 spp., W North America, M. pygmaea D. Don in Chile and S Peru, and one in Australia, and New Zealand.

81. Picrosia D. Don. Perennial herbs, involucral bracts in a single series, receptacle naked, flowers white, pink or violet, achenes fusiform, with long beak, pappus of many scabrid bristles. x = 7, diploids. Two spp., Peru to Chile, Argentina and S Brazil (both, none endemics).

■ SUBTRIBE HYPOCHAERIDINAE – outsiders are Urospermum (2; Canary Islands, Mediterranean to Pakistan), Prenanthes (1; Europe, Middle East), Scorzoneroides (c 25; Europe, Canary Islands, Mediterranean, North Africa to the Middle East, temperate Asia), Helminthotheca (5; SE Europe, Mediterranean to Iran, North Africa), Picris (c 60; Europe, Mediterranean, African mountains, temperate Asia to N Australia), Hedypnois (4; Macaronesia, Mediterranean to Iran), Leontodon (c 80; Europe, Mediterranean, North Africa, temperate Asia, SW Asia to Iran).

82. Hypochaeris L. Perennial to annual herbs with coarse, multicellular, unbranched hairs, involucral bracts in several rows, receptacle scaly, florets yellow, white or pink, achenes beaked, sometimes dimorphic, pappus of plumose bristles with stiff pinnulae. x = 6, 5, 4, 3, diploids and tetraploids. Eurasia, northern Africa and South America (60), from Colombia and W Venezuela to Chile and Argentina and S Brazil (9, 2 endemic).

■ SUBTRIBE CREPIDINAE – outsiders are Acanthocephalus (2; Altai, Uzbekistan), Ixeris (25–28; E and SE Asia), Youngia (c 35; Asia to India and Japan), Crepidiastrum (c 25; E Asia to Japan and Taiwan in China), Askellia (12; Iran, Central Asia to Pakistan and Mongolia, N India, Himalayas, Tibet, Siberia to Russian Far East, China, temperate North America), Crepis (c 200; the Northern Hemisphere, tropical and southern Africa, South America), Dianthoseris (1; E and NE African highlands), Lagoseris (2; Crimea, Türkiye), Rhagadiolus (2; Mediterranean to Iran, Canary Isles, British Isles to Mediterranean, Türkiye, the Caucasus), Lapsana (1; Europe, W and SW Asia); Syncalathium (5; Tibet, China), Hololeion (3; southern Korean Peninsula, Japan, China, Korean Peninsula, Russian Far East), Soroseris (7; Himalayas, Tibet, W and SW China), Dubyaea (18; Himalayas, Tibet, W China), Nabalus (c 25; China, Korean Peninsula, Japan, Russian Far East, Canada, U.S.A.); Garhadiolus (4; Cyprus, Egypt, Türkiye, the Caucasus, the Middle East and Arabian Peninsula to Central Asia and China), Lagoseriopsis (1; Central Asia), Heteracia (1; Balkan Peninsula, Crimea, the Caucasus, SW and Central Asia to China), Heteroderis (1; Egypt, Arabian Peninsula, SW to Central Asia, Pakistan).

83. Taraxacum Weber. Perennial scapose herbs with taproot, leaves in basal rosette, scape leafless, hollow, involucral bracts in two unequal series, the outer shorter, often reflexed at tips, receptacle naked, capitula with very many flowers, florets yellow. 2,421 spp. (3^th largest worldwide), spp., North Hemisphere, South America and New Zealand. 32 spp. in New World, 13 from Mexico southwards, in Mexico (3, two endemics, one up to Panama); two in Caribbean to Colombia, one up to Costa Rica; T. craspedotoides A.J.Richards from Colombia to Venezuela; six from Peru (3, two endemics) to S Chile and Argentina, and T. fernandezianum Dahlst. from Juan Fernández Is., C. Chile, S. Brazil to Argentina (Neuquén, Salta).

7.2 CICHORIOIDEAE ▸ TRIBE LIABEAE (17/ 190) - all genera in South America. Mexico, Central America, the Caribbean, tropical South America, with their largest diversity in NW South America. Small trees, shrubs or annual or perennial herbs. Latex usually present. The Liabeae are distributed throughout much of the Neotropics, but they exhibit their greatest generic and specific concentration in W South America and the most likely place of origin of the modern day tribe is in the Andean Cordillera; The center of generic diversity is in Peru where 13 of the 18 genera are found, followed by Ecuador (8), Colombia (7), Bolivia (6), Costa Rica (5), Panamá (5), Venezuela (4), Mexico (4), Argentina (5), Guatemala (2), El Salvador (2), Honduras (2), Brazil, Nicaragua, and Caribbean (Cuba, Hispaniola, Jamaica) with one genus each. Only one genus absent in South America; 4 endemic genera from Peru.

84. Austroliabum Cabrera. Annual to perennial herbs or subshrubs, with latex. 3 spp., endemic to NW Argentina.

85. Bishopanthus H. Rob. Shrub; with latex; leaf bases fused into sheath; blades 3-nervate, bullate; heads solitary, pedunculate, campanulate; involucral bracts c. 25, in 2 subequal series; epaleaceous; ray florets c. 20; limbs linear; disc florets c. 25; bases of anther thecae short-tailed, short-fringed. Only one sp., B. soliceps H. Rob., endemic to Peru.

86. Cacosmia Kunth. Shrub; usuallywith latex; stems densely pubescent; leaf bases fused into sheath; blades 3–5-nervate, bullate; inflorescence densely corymbiform; heads cylindrical; involucral bracts 20–25, 5–6-seriate, outer bracts ranked, inner bracts not in ranks; receptacle epaleaceous; ray florets 5, limbs broad; disc florets 5–6; bases of anther thecae without tails, minutely digitate; cypselae 3–5-angled, glabrous, raphids elongate; pappus lacking. 3 spp. in Ecuador, one into Peru.

87. Chionopappus Benth. Shrubs; latex not noted; stems arachnoidtomentose; leaf bases fused into sheath, blades 3-nervate; inflorescences simple dichasia; heads campanulate; involucral bracts 50–55, c. 5-seriate; paleae strap-shaped; ray florets c. 40; disc florets 75–125; corollas red, glabrous; bases of anther thecae short-tailed; cypsela 8–10-ribbed, setulae minute, raphids elongate; pappus bristles 8–10, plumose. Only one sp., C. benthamii S.F. Blake, endemic to Peru.

88. Chrysactinium (Kunth)Wedd. Perennial rhizomatous herbs, rosettiformor shortstemmed; stems evanescent-tomentose; leaf bases cuneate or petioliform; blade 3-nervate, tomentose; heads solitary, long-pedunculate, broadly campanulate; involucral bracts 40–60, 4–5-seriate; receptacle squamulose; ray florets 30–60; disc florets 30–60; anther collar cells annulate; cypselae 8–10- ribbed; pappus bristles 30–60, white, no squamellae. 6 spp., Ecuador, northern Peru.

89. Dillandia V.A. Funk & H. Rob. Moderate-sized to small herbs less than 60 cm tall; latex not noted; stems arachnoid-tomentose; leaf bases sessile, subpetiolate or perfoliate; surface bullate; blades pinnately veined; inflorescences of 1–2 heads or, more frequently, a 3–7-headed subumbel; ray florets 15–40, limbs oblong to narrowly oblong; disc florets 10–30, corollas yellow to yellow and purple, tubes pilose, anther thecae pale, bases rounded; cypselae (immature) 7–10-ribbed, densely setulose, with a few subquadrate raphids; pappus bristles 10–30, sometimes shorter outer bristles. Three spp., Colombia, Ecuador and Peru.

90. Erato DC. Coarse perennial herbs; stems and leaves pilose or strigose; leaves with oblong pseudostipules; blades palmately 5–9-veined, surfaces green; inflorescence cymose to subumbellate; heads broadly campanulate; involucral bracts 40–100, c. 4-seriate, tips sometimes tomentose; receptacle foveolate and ridged; ray florets 75–230; disc florets 20–150; cypselae 4-ribbed, glabrous or hispidulous; pappus of 25–50 bristles or short awns, no outer series. 5 spp., 1 Costa Rica and Panamá, 4 in Andes from Venezuela to Bolivia.

91. Ferreyranthus H. Rob. & Brettell. Shrubs or weak trees; without latex; stems arachnoid-tomentose; leaf bases fused into sheath; blades pinnately veined; inflorescences densely corymbiform; heads broadly campanulate; involucral bracts 45–55, c. 5-seriate; receptacle squamuliferous; ray florets 8–12, limbs short; disc florets 12–25, corollas glandular-dotted; bases of anther thecae short-tailed, strongly fringed; cypselae c. 10-ribbed, with setulae and glands, raphids elongate; pappus bristles 10–15, squamellae narrow. 8 spp., Peru, one into Ecuador.

92. Liabum Adans. Perennial herbs; without latex; stems arachnoid-tomentose; leaf bases connected across node, often forming nodal disc; blades 3-nervate; inflorescence partly subumbellate. 41 spp., Mexico, Central America, Greater Antilles, and Andes of South America, including here Venezuela and W Brazil, in forests of Acre state; 30 spp. in South America. L. acuminatum Rusby was identified (by H. Robinson) from a 1968 collection from Acre, Brazil making it the first confirmed record of the tribe from that country; subsequently, L. amplexicaule Poepp. was recorded from the region.

93. Microliabum Cabrera. Annual to perennial herbs or subshrubs;with latex; stems white-tomentose and glandular-hairy; leaf bases broadened or with pseudostipules or nodal discs; blades 3-nervate; heads solitary or in cymes, broadly campanulate; involucral bracts 30–75, 2– 4-seriate, subequal to gradate; receptacle epaleaceous; rays 10–30; limbs narrowly elliptical to linear; disc florets 15–175; bases of anther thecae with few or no teeth; cypselae 8–10-ribbed, setuliferous, raphids elongate. 3 spp., southern Bolivia, N Argentina.

94. Munnozia Ruiz & Pav. Annual or perennial herbs or subshrubs. Petioles sometimes winged, bases often pseudostipulate; blades 3-nervate to pinnately veined, usually tomentose abaxially; inflorescence terminal, more or less corymbose; heads broadly campanulate; involucral bracts 17–70, 2–4-seriate, subequal to gradate; receptacle often squamulose; florets yellow, rarelywhitish to lavender; rayflorets 6–70; disc florets 9–85; cypselae 6–10-ribbed, setuliferous; pappus bristles sordid to reddish, 5–55, with squamellae. 47 spp., mostly from the Andes from Venezuela to Argentina (46 in South America), two in Costa Rica, Panamá.

95. Oligactis (Kunth) Cass. (exc. Sampera p.p.) Shrubs and vines, without latex; leaf bases sometimes confluent across nodes; blades pinnately veined; inflorescence densely corymbiform; heads narrowly to broadly campanulate; involucral bracts 16–55, 4–5-seriate; receptacle ridged and squamuliferous; ray florets 3–18, limbs short; disc florets 3–34; style branches rather long; anther thecae bases digitate; cypselae 5–8-ribbed, with glands and contorted setulae, raphids subquadrate; pappus bristles 20–35, tips often broad, squamellae 7–10. 5 spp. from Costa Rica to Peru and Venezuela.

96. Paranephelius Poepp. Acaulescent perennial herbs; leaf bases not sheathing; blades trinervate to pinnately veined; heads sessile, broadly campanulate; involucral bracts 40–50, c. 4-seriate; receptacle ridged; ray florets 20–35; style branches spiralled; disc florets 20–33; cypsela c. 10-ribbed, glabrous or with some arachnoid hairs; pappus bristles 45–80, outer series indistinct. 6 spp., Ecuador, Peru, Bolivia, one to northern Argentina.

97. Philoglossa DC. Small erect to creeping herbs; stems and leaves pilose to strigose; leaf bases with oblong pseudostipules; blades 3-nervate, surfaces green; involucres broadly campanulate; bracts 20–30, 3–4- seriate, subequal to gradate; ray florets 21–70.Disc florets 30–60, yellow, rarely deep purple or brown; cypselae compressed, 2-ribbed, mostly glabrous; pappus deciduous squamellae or awns, or lacking. 5 spp., southern Colombia to Bolivia, all in Peru (3 endemics).

98. Pseudonoseris H. Rob. & Brettell. Small perennial herbs; leaf bases sessile, subauriculate; blades pinnately veined; inflorescence scapose or subscapose; heads pedunculate, broadly campanulate; involucral bracts c. 40, c. 4-seriate; receptacle subglabrous; ray florets yellow or yellow-orange, 15–20; disc florets orange-yellow or red, 25–55. Three spp. from Peru, one of them up to Bolivia.

99. Sampera V. A. Funk & H. Rob. (inc. Oligactis p.p.) Scrambling shrubs, moderately branching, stems terete to strongly hexagonal, mostly tomentose, without latex; nodes with or without disks; leaves opposite; petioles with or without wings, sometimes included in perfoliate leaf pairs; blades sharply delimited and rounded to slightly cordate at base or confluent with petiole, ovate to oblong-ovate, margins subentire to serrate, never angulate, upper surface flat to slightly bullate, densely tomentose below, pinnately veined; inflorescence terminal on branches, corymbiform, peduncles less than 5 cm long, thinly to densely tomentose. 8 spp. from Colombia to to N Peru.

100. Sinclairia Hook. & Arn. Subshrubs, shrubs or vines, 2 spp. epiphytes; with latex; stems arachnoid-tomentose; leaves opposite or ternate, sometimes seasonally deciduous, bases without sheaths or pseudostipules; blades 3-nervate; inflorescences laxly to densely corymbiform or pyramidal; heads narrowly to broadly campanulate; involucral bracts 18–45, 3–5-seriate; receptacle glabrous, spinulose or puberulous; ray florets absent or 4–25; disc florets 5–30; anther thecae bases minutely crenulate; cypselae c. 8–10-ribbed, glabrous or setuliferous, raphids elongate; pappus bristles 30–40, squamellae 15–20. 18 spp., Mexico, Central America, one up to W Colombia.

101. Stephanbeckia H. Rob. & V. A.Funk. Small, short-stemmed, basally branched, annual or short-lived perennial herbs with dense white tomentum on parts of petioles, undersurfaces of leaf blades, peduncles and margins of involucral bracts; latex lacking. Only one sp., S. plumosa H. Rob. & V. A. Funk, restricted Tarija departament, Andes from S Bolivia.

7.3 CICHORIOIDEAE ▸ TRIBE MOQUINIEAE (2/2) - both species endemic to Brazil.

102. Moquinia DC. Trees or shrubs gynodioecious; stem hairs white, arachnoid; leaf lower surface whitish or pale yellowish-tomentose; inflorescences with densely racemiform branches; involucral bracts c. 25, c. 4–5-seriate; florets 4 or 5. Only one sp., M. racemosa (Spreng) DC., endemic to the states of Bahia and Minas Gerais in Brazil.

103. Pseudostifftia H. Rob. Shrubs or trees; monoecious; stem and leaf hairs appressed, symmetrically T-shaped, cap-cells broadly fusiform, stalks slender; inflorescences with corymbiform branches; involucral bracts c.18, c. 5-seriate; floret 1. Only one sp., P. kingii H. Rob, Bahia, Brazil.

7.4 CICHORIOIDEAE ▸ TRIBE VERNONIEAE (134/1.335–1.350) - outsiders are 31 genera only in tropical Africa, 6 from Mexico to Central America, 5 endemics to Madagascar, 3 only in Cuba and Jamaica, 16 from India to China and New Guinea, two endemics to Australia, 3 shared from continental Africa and Madagascar, 6 from tropical and arid areas in Africa to Asia, inc. Madagascar and Sri Lanka; and two at U.S.A. assignation: Stokesia (1; SE U.S.A.) and Hesperomannia (3; Hawaii); subtribes annuncied includes only the tribes with South American genera, without details.

■ SUBTRIBE VERNONIINAE

104. Allocephalus J. Bringel, J. N. Nakaj. and H. Rob. Annual herb ca. 70 cm tall; stems cylindrical or flattened and sulcate when dried; leaves alternate; capitula discoid, sessile, in axillary glomerules or small spikes; inflorescence cymose; florets 6 per capitulum, the corolla purple, ca. 7.7–8.0 mm long, actinomorphic. Only one sp., A. gamolepis J. Bringel, J. N. Nakaj. and H. Rob., restricted to one location in the Paranan River Valley area (small area of NE Goiás state) in the understory of the dry forest on limestone outcrops covered by a thin layer of soil and all the specimens belong to the same collection site.

105. Aynia H. Rob. Perennial herbs; hairs simple; peduncles usually long; heads subtended by large foliose bracts, involucral bracts c. 100 in 4–5 series, 10–25mm long; florets c. 50; anthers tails short, truncate; style with node; sweeping hairs pointed; achene 10-ribbed, raphids subquadrate; pappus capillary, with squamellae. Pollen echinolophate, with intercolpus-aligned polar lacunae. Only one sp., A. pseudascaricida H. Rob., endemic to Peru.

106. Chrysolaena H. Rob. Perennial, usually xylopodial herbs, sometimes with rhizophores, sericeous or lanate with simple yellowish hairs; inflorescence seriate-cymose, heads sessile; involucral bracts/florets 1–2/1; florets 10–65; anther bases obtuse; apical appendage usually with glands; style without prominent node; sweeping hairs broadacicular. 19 spp., Brazil (16, only 5 endemic) to Argentina and Bolivia. Two spp. from Paraná state are considered rare species in Brazil by Plantas Raras do Brasil’s book.

107. Cololobus H. Rob. Subshrubs; hairs simple or asymmetrically T - shaped; inflorescence thyrsoid; involucral bracts, c. 6-seriate, c. 35, in outer 4 series pubescent, inner mostly glabrous; florets 20–30; corollas glabrous, lobes very short; anther base obtuse; style with node; sweeping hairs blunt, often septate. 5 spp. from Espírito Santo, Rio de Janeiro state, in SE Brazil.

108. Cyrtocymura H. Rob. Perennial herbs; hairs simple; inflorescences scorpioid-cymose with crowded sessile heads, deciduous with age; involucral bracts 20–30, in c. 3 series; florets 14–30; corolla lobes sericeous; anther bases rounded; style with node; sweeping hairs broadly acicular; achenes 10-costate, setuliferous, idioblasts bulging, raphids elongate; pappus capillary, outer squamellae persistent. 6 spp., C. scorpioides (Lam.) H. Rob. widely distribuited in tropical America, one in Caribbean, and 4 in Brazil and adjacent Argentina.

109. Dasyanthina H. Rob. Perennial herbs 2–4m high; stem hairs T-shaped; inflorescences rounded-thyrsoid; peduncles slender; involucral bracts c. 60 in 5–6 series; florets c. 25; corolla insidewith unicellular hairs; anther tails unsclerified, connective with glands; stylar node annular; sweeping hairs pointed; achenes 8–10- ribbed, setuliferous, raphids elongate; pappus capillary, squamellae persistent. Two spp., E Brazil (Espírito Santo to São Paulo and Minas Gerais states) and adjacent Pantanal in Paraguay.

110. Dipterocypsela S.F. Blake. Fleshy herbs; hairs symmetrically T-shaped. Seriate cymes with crowded sessile heads; involucral bracts c. 12 in c. 2 series, subequal, winged; florets c. 12; corollas zygomorphic, inner lobes longer; anther bases rounded; apical appendage with gland; style with node; sweeping hairs fusiform, septate. Only one sp., D. succulenta S.F. Blake, endemic to Colombia.

111. Echinocoryne H. Rob. Perennial herbs; sericeous with straight hairs; heads pedunculate; involucral bracts c. 110–500 in 6–9 series, linear, straight, pungent; florets 15–60; anther bases rounded; style with node; sweeping hairs acicular; achenes 5-costate, sericeous, raphids elongate; pappus capillary, with squamellae. Pollen echinolophate, no polar lacunae. Six spp., endemics to Brazil; one sp, from Goiás state, is considered a rare species in Brazil by Plantas Raras do Brasil’s book (assignated as Vernonia).

112. Eirmocephala H. Rob. Perennial herbs; hairs simple; inflorescence seriate- or densely scorpioid-cymose; heads sessile, persistent; involucral bracts 24–65, in c. 4 series; florets 7–35; anthers tailed; apical appendage often glanduliferous; style with node; sweeping hairs acicular; achenes 10-veined, setuliferous, without idioblasts, raphids elongate; pappus capillary, with squamellae. Three spp., Central America (1) to Venezuela and Bolivia, one of them in Brazil.

113. Heterocypsela H. Rob. Perennial herbs; hairs symmetrically T-shaped. Peduncles 5–30mm long; involucral bracts c. 70 in c. 6 series, caudate-acuminate; florets c. 60–70; anther bases blunt; style with node; sweeping hairs sharp; outer achenes obcompressed, margins winged, raphids subquadrate, pappus bristles deciduous; inner achenes prismatic, setuliferous, pappus more persistent; pollen subtriporate. Two spp., endemics to Minas Gerais, Bahia and Tocantins states in C Brazil.

114. Lepidaploa (Cass) Cass. Herbs or shrubs, rarely annual; sericeous or tomentose, hairs simple or T-shaped; leaves usually alternate; inflorescences seriately cymose; heads usually sessile; involucral bracts 20–70, in 3–6 series; florets (8–)10–35; anther bases obtuse; apical appendages rarely with glands; style with node; sweeping hairs broadly acicular; achenes 8–10-ribbed, raphids elongate. 145 spp., tropical America, 106 in South America, 58 in Brazil, 43 endemics; one sp., from Minas Gerais state, is considered a rare species in Brazil by Plantas Raras do Brasil’s book (assignated as Vernonia).

115. Lessingianthus H. Rob. Perennial herbs, sometimes xylopodial, with stem tubers and roots crown; hairs simple; inflorescences simple or seriate-cymose; heads sessile or pedunculate; involucral bracts 45–100 in 4–6 series; florets 15–50; anther bases rounded; style usually without node; sweeping hairs sharp; achenes 5-costate, without glands, raphids subquadrate or elongate; pappus capillary, with squamellae. 138 spp., 134 from Ecuador to Argentina and Uruguay, mostly Brazil (119, 82 endemics) and Argentina, two endemics to Colombia, one endemic to Venezuela, and L. rubricaulis (Bonpl.) H. Rob. in Central America to Brazil and Cono Sur. Nine spp., in several states, are considered rare species in Brazil by Plantas Raras do Brasil’s book (all assignated as Vernonia).

116. Mattfeldanthus H. Rob. & R. M. King. Shrubs; hairs simple; inflorescence base pseudotrichotomous, branches seriate-cymose, bracts foliiform; heads sessile; involucral bracts c. 100, c. 7-seriate; florets 14–16; outer corollas zygomorphic, innermost lobe longer; anther tails lobed; style with node; sweeping hairs acicular; achenes c. 10-costate, sericeous, raphids elongate; pappus capillary, with squamellae. Two spp., endemics to Bahia, Minas Gerais states in E Brazil.

117. Pseudopiptocarpha H. Rob. Shrubs or subshrubs; hairs appressed, symmetrically T-shaped; heads axillary, clustered, sessile; involucral bracts 25–30 in c. 5 series; florets 8–10; anther tails short; style with node; sweeping hairs sharp; achenes weakly 10-costate, setulae, glands, and idioblasts scattered, raphids subquadrate; pappus bristles capillary, with squamellae. Three spp., all endemic to Colombia.

118. Quechualia H. Rob. Erect or scrambling shrubs; hairs asymmetrically T-shaped, often proliferated; inflorescences thyrsoid; heads pedunculate; involucral bracts 60–90 in 5–6 series; florets 30–55; corolla limb often with multicellular hairs inside; anther connectives with glands, tails denticulate; style with node; sweeping hairs subacicular; achenes 8–10-ribbed, setuliferous, raphids elongate. 4 spp. in Bolivia (3 endemics), one up to Cono Sur and Peru.

119. Stenocephalum Sch. Bip. Perennial herbs, sometimes with xylopodium; hairs simple, often arachnoid; leaves pale tomentose below; heads axillary or in panicles; involucral bracts 15–22 in 3–4 series; florets 4–7(–10); anther bases rounded; style with node; sweeping hairs short, sharp; achenes 10-ribbed, setuliferous, without glands, idioblasts, or raphids; pappus capillary, squamellae present. 6 spp., Central America and tropical South America, 5 in Brazil, two endemics.

120. Stilpnopappus Mart. ex DC. Perennial herbs or shrubs; hairs simple; inflorescences axillary or seriate-cymose; heads sessile, 1 to few per node; involucral bracts 20–50 in 3–4 series; florets 6–50; anther bases obtuse; style with node; sweeping hairs acicular; achenes 8-costate, projecting rim at top, setuliferous, idioblasts scattered, raphids subquadrate; pappus of lanceolate awns. 9 spp., all endemics to Brazil, one of them, from Piauí state, is considered a rare species in Brazil by Plantas Raras do Brasil’s book.

121. Strophopappus DC. 9 spp., 8 in Brazil (6 endemics, 2 up to Bolivia) and Bolivia, and one endemc to Venezuela.

122. Struchium P. Browne. Decumbent annuals; hairs sparse, simple; heads axillary, clustered; involucral bracts 20–25 in c. 2 subequal series; florets 50–70; corolla lobes 3 or 4; anther bases rounded, connective with glands; style with node; sweeping hairs sharp; achenes 3–5-angled, without setulae, idioblasts numerous, raphids elongate; pappus a cartilaginous sleeve. Only one sp., S. sparganophora (L.) O. Kuntze, pantropical, widely adventive in Belize, Bolivia, Brazil, Cuba, India, Jamaica, Panamá, Peru. Pantropical, in open woodland, wet areas in riverine forests, margins of seasonally flooded areas. 0–1,200m.

123. Trepadonia H. Rob. Vines; hairs mostly symmetrically T-shaped; leaves alternate or opposite; inflorescence branching often at 90°, branchlets often subracemose; involucral bracts c. 25 in c. 5 series; florets 8–10; corollas glabrous; anther bases obtuse; style with node; sweeping hairs acicular; achenes 10-costate, setulae scattered, raphids subquadrate; pappus capillary, squamellae persistent. Two spp., endemic to Peru.

124. Vernonanthura H. Rob. Subshrubs to small trees, sometimes xylopodial; hairs simple or T-shaped; inflorescences thyrsoid; heads sessile to long-pedunculate; involucral bracts 6–30(–60) in 4–10 series; florets 4–30; corollas without hairs; anther bases obtuse or tailed; appendages often with glands or hairs; style with node; sweeping hairs short-acute; achenes 8–10-costate, setuliferous or with idioblasts. 68 spp., tropical America, 54 in South America, 44 in Brazil, 26 endemics; two of them, in Distrito Federal and Minas Gerais state, are considered rare species in Brazil by Plantas Raras do Brasil’s book (both assignaded as Vernonia).

125. Vernonia Schreb. (exc. Mesanthophora) Perennial herbs, bases decumbent; hairs simpleor symmetrically T-shaped; inflorescences cymose with branches longer than central axis; heads usually pedunculate; involucral bracts c. 50 in 5–6 series; florets 8–120; anthers without tails; appendages often with glands; style with node; sweeping hairs acute, sometimes septate; achenes 5–10-costate, with setulae, glands and/or idioblasts. 22 spp., SE U.S.A., Bahamas (1), to C Mexico (3), and two distinctive species disjunct in South America: V. echioides Less. and V. incana Less., in wetlands and marshes of Paraguay, Uruguay, Brazil and Argentina.

Nine species previously belonging to Vernonia, from several states are all considered rare species in Brazil by Plantas Raras do Brasil’s book, but non detected in VPA circumscription.

126. Xiphochaeta Poepp. Aquatic short-lived herbs; hairs simple, appressed; heads sessile, 1(–3) in axils; involucral bracts 70– 80 in 3–4 series; florets c. 30; anther bases not calcarate; appendages with glands; style with node; sweeping hairs acicular; achenes 5-costate, setulae and idioblasts scattered, raphids elongate; pappus segments c. 10, canaliculate, non-costate; pollen echinolophate. Only one sp., X. aquatica Poepp., Amazon and Orinoco basins, in Brazil, Peru, Venezuela and Guianas.

■ SUBTRIBE PIPTOCARPHINAE

127. Critoniopsis Sch.Bip. (inc. Oliganthes) Shrubs or trees; hairs often stellate; leaves alternate or opposite, petiole often lobed or winged; inflorescences terminal; involucral bracts 18–25(–35) in 4–6 series; florets 1–11(–15 or 20); corolla throat present, lobes often recurved, with small glands; anthers often with unsclerified tails; style usually with node; sweeping hairs obtuse; achenes 5–10-costate; pappus capillary. 72 spp., Mexico, Central America, Andes to Brazil; 71 spp. in South America, only 5 in Brazil, all endemics.

128. Cuatrecasanthus H. Rob. Weak shrubs; hairs with apical cell enlarged at base; inflorescence terminal; heads sessile in glomerules; involucral bracts c. 15 in 5–6 series; florets 1; corolla throat lacking; anthers with fimbriate tails; style with node; sweeping hair tips rounded; achenes 10-costate, glanduliferous and spiculiferous, raphids lacking; pappus capillary, outer setae short. 6 spp., restricted from Ecuador and Peru.

129. Dasyandantha H. Rob. Small trees; stems sublanate, hairs simple; leaves to 30 cm long; inflorescence thyrsoid-paniculate; heads sessile in glomerules; involucral bracts c. 30, in c. 4 series; florets c. 12; corolla throat enlarged, pilosulous inside; anthers with unsclerified tails; style with node; sweeping hairs blunt-tipped; achenes 8-ribbed, setuliferous; pappus capillary, fragile. Only one sp., D. cuatrecasasiana (Aristeg.) H. Rob., endemic to Andes of Venezuela.

130. Huberopappus Pruski. Shrubs; hairs stellate; inflorescences of few shortpedunculate heads and small foliiform bracts; involucral bracts 14–17 in 3–4 series; florets 19–22; anther bases rounded; style without node; sweeping hairs obtuse, often septate; achenes 3–5- angled, c. 10-striate, glabrous; pappus collarform, some with single awn. Only one sp., H. maigualidae Pruski, endemic to Pantepui Life Zone in Sierra de Maigualida, S Venezuela (Bolivar-Amazonas border), 2,000 – 2,100m elevation range.

131. Joseanthus H. Rob. Shrubs or trees; hairs sometimes T-formed; leaves opposite; inflorescences densely corymbose.Heads shortly pedunculate; involucral bracts 20–30 in c. 4–5 series; florets 9–12; corolla densely pilosulous, without throat; anthers with fimbriate tails; style with node; sweeping hairs obtuse; achenes 3–8-costate, with glands and small setulae; pappus capillary, with squamellae. 4 spp. from Colombia and Ecuador.

132. Piptocarpha R. Br. Scandent scrambling shrubs or trees, sometimes with roots crown; hairs stellate or lepidote; leaves alternate or opposite; heads in axillary or terminal clusters; involucral bracts 18–30 in 3–4 series; florets 3–20; corolla lobes often recurved; anther tails sclerified; style with node; sweeping hairs blunt, often septate; achenes 3–5-costate, glabrous, with idioblasts, raphids short-oblong; pappus capillary. 49 spp. of South America, all endemic but one into Mexico and two in Caribbean; 32 in Brazil, 23 endemics.

133. Piptocoma Cass. Trees and shrubs; hairs stellate to lepidote; inflorescences corymbiform; florets 1–12; corolla throat very short; anther bases rounded; style without node; sweeping hairs mostly blunt, often septate; achenes weakly 7–10-costate, idioblasts present; pappus with deciduous linear segments, persistent outer scales. 18 spp., 4 only in Caribbean, 14 in northern South America (Guyana to Peru), highly centered in Venezuela and Colombia, only two of them outside this area, both in Brazil, none endemic.

134. Yariguianthus S Díaz & B.V. Rodríguez C. Shrubs or treelets, scandent; cylindric stems. Only one sp., Y. glomerulatus S Díaz & B.V. Rodríguez C, known only Santader departament in Colombia.

■ SUBTRIBE CHRESTINAE

135. Chresta Vell. ex DC. Perennial herbs and subshrubs, sometimes xylopodial; hairs symmetrically or asymmetrically T-shaped or forming dense felt; inflorescences long-peduculate, clusters globose, subumbellate or spicate; involucral bracts 10–20 in 5–6 series; florets 2–12; corolla tube long; anther bases rounded; appendage sometimes with glands; sweeping hairs fusiform. 17 spp., 16 endemic to Brazil, C. exsucca DC. also in Bolivia; within the dry seasonal scrubland of NE Brazil (caatinga), Chresta is one of the Asteraceae genera that is found outside the rocky grasslands (campos rupestres), although it is strongly associated with rocky environments; of the four species of Chresta found in the dry seasonal scrubland of NE Brazil (caatinga), three are associated with quartzitic outcrops and one is usually found on granitic inselbergs, all growing directly on top of the rocks or in pockets of plant debris found in small cracks and crevices.

136. Soaresia Sch. Bip. Subshrubs, with xylopodia; stems felted with fusiform hairs; leaves imbricate, silvery velvety, veins nearly longitudinal. Rows of sessile heads in upper axils; involucral bracts c. 12 in c. 3 series; florets 4; corollas sericeous; anthers tailed; sweeping hairs acicular; achene 5-ribbed, sericeous, raphids elongate; pappus awns c. 15, subulate, basally winged. Only one sp., S. velutina Sch. Bip., endemic to Rondonia to Minas Gerais in center Brazil.

■ SUBTRIBE PACOURININAE

137. Pacourina Aubl. Hairs simple, small; leaf bases clasping, margins spinose-toothed; heads single and sessile in axils; involucral bracts c. 50 in c. 3 series, broad, green, margins whitish; florets c. 50; corolla lobes sclerified distally; anther tails toothed; achene c. 10- costate, idioblasts in furrows, raphids none; pappus bristles short, deciduous, squamellae persistent. Only one sp., P. edulis Aubl., Central and South America in Argentina, Belize, Bolivia, Brazil, Colombia, Ecuador, Dominican Republic, Guatemala, Nicaragua, Paraguay, Peru, Suriname, in riverine or lake margins as an aquatic or semiaquatic, seasonally flooded grassland.

■ SUBTRIBE ERLANGEINAE

138. Acilepidopsis H. Rob. Perennial herbs decumbent from thickened rhizome; with numerous glandular dots, hairs forming felt on stems; inflorescence branches seriately cymose; heads sessile; involucral bracts c. 30; florets 8–13, corolla lobes reflexed, sericeo-pilosulous; anther bases and appendages sclerified; style without node; achene 10-costate; raphids lacking; pappus capillary. Only one sp., A. echitifolia (Mart. ex DC.) H. Rob., Argentina, Bolivia, S Brazil and Paraguay.

139. Mesanthophora H. Rob. (off Vernonia) Glabrous perennial herbs; leaves sessile, shortly decurrent, auriculate, glandular-dotted below. Racemiform leafy cymes with peduncles from middle of internodes; involucral bracts c. 100 in c. 5 series, linear-lanceolate, erect-patent; florets 90–100; anther bases rounded; style with node; achenes 8–10-costate, costae scabrid; pappus bristles deciduous, capillary, squamellae persistent. Only one sp., M. brunneri H.Rob., SE Bolivia, Paraguay and Brazil (Mato Grosso do Sul, disjunct in Minas Gerais).

140. Telmatophila Mart. ex Baker. Perennials; sericeous with simple hairs; inflorescences axillary, secondary heads surrounded by foliaceous bracts; heads cylindrical; subequal involucral bracts and florets 4; corollas with long hairs distally; style base without node; achenes oblong-ovoid, c. 8-costate, with idioblasts, long setulae deeply divided, raphids elongate; pappus of c. 8 short, laciniate scales. Only one sp., T. scolymastrum Mart. ex Baker, from E Piauí to Pernambuco state in NE Brazil.

■ SUBTRIBE TRICHOSPIRINAE

141. Trichospira Kunth. Perennials; hairs simple, arachnoid; leaves sessile, tomentose below.Heads axillary, sessile; involucral bracts c. 12, subequal, receptacle with few pales; florets c. 10; corollas 4-lobed; anthers with spurs short; style without node; sweeping hairs acute, often septate; achenes with 2 marginal ribs and 2 or 3 weaker ones on surfaces, raphids lacking. Only one sp., T. verticillata (L.) S.F. Blake, tropical America in Bolivia, Brazil, Colombia, Cuba, Dominican Republic, Guatemala, Guyana, Haiti, Mexico, Panamá, Peru, Venezuela; disturbed areas, riverbanks, damp roadsides, wet savannas of C Brazil (cerrado).

■ SUBTRIBE ROLANDRINAE

142. Rolandra Rottb. Shrubs; hairs simple; leaves white-tomentose below; inflorescences axillary, sessile; involucral bracts 2, awned; corolla 4-lobed, glabrous; anther bases rounded, with sterile margin; style without node; sweeping hairs acicular; achenes 5-veined, without setulae, with glands, many poorly differentiated idioblasts; pappus a persistent ring of jagged scales. Only one sp., R. argentea Rottb., tropical America in Bolivia (Pando), Brazil, Colombia, Costa Rica, French Guiana, Guyana, Honduras, Panamá, Puerto Rico, Surinam, Venezuela.

143. Spiracantha Kunth. Weak herbs; hairs simple; leaves white-tomentose below; inflorescence of pedunculate, 3–4-bracted glomerules; involucral bracts c. 6, inner awned.Corolla with filaments near sinuses, lobes 4 or 5; anther bases obtuse; style without node; sweeping hairs acicular; achenes fusiform, weakly 5–6- veined, few glands and setulae above; pappus of short bristles. Only one sp., S. cornifolia Kunth, from Mexico to Colombia and Venezuela and Caribbean.

■ SUBTRIBE GYMNANTHEMINAE

144. Gymnanthemum Cass. Shrubs or trees; stems often felted, hairs rarely asymmetrically T-shaped; inflorescences densely corymbiform; c. 43 spp. from Africa and Asia, with G. amygdalinum (Delile) Sch. Bip. ex Walp. also in dry areas of Brazil and Bolivia.

■ SUBTRIBE ELEPHANTOPINAE

145. Caatinganthus H. Rob. Short-lived herbs, tomentum arachnoid; leaves linear. Branches spiciform, 3–20 adaxial heads, biseriate; outer involucral bracts 4–6, foliiform distally, inner 4, scariose. Anther bases rounded; style with node; sweeping hairs acicular; achene 10-ribbed, long-setuliferous, raphids subquadrate; pappus setae c. 10 and scales 10, large. Pollen lophate, single polar lacuna. Two spp., endemics to NE Brazil.

146. Elephantopus L. Perennials, sometimes with xylopodium; pilose, hairs simple, stiff; leaves mostly basal; heads in bracteate glomerules; involucral bracts 8; florets 2–4; corollas zygomorphic, with deep inner sinus; anthers not calcarate, not tailed; style without node; sweeping hairs acicular; achenes 10-costate, setulose, raphids elongate; pappus bristles 5–15(40–81), straight, basally winged. 20 or more species, pantropical, 17 in New World, 11 in South America, 9 in Brazil, six endemics.

147. Orthopappus Gleason. Perennials, hairs simple, stiff, yellow, with rhizophores; leavesmostly basal; inflorescence a spiciform cluster of narrow heads; involucral bracts 8; corollas zygomorphic, inner sinus deepest; anthers not long-calcarate; style with node; sweeping hairs acicular; achenes 5-costate, setuliferous, idioblasts along costae, raphids elongate; pappus bristles 2–3-seriate, inner basally winged. Only one sp., O. angustifolius (Sw.) Gleason, tropical America (Argentina, Bolivia, Brazil, Honduras, Colombia, Ecuador, El Salvador, Guatemala, Honduras, Mexico, Panamá, Paraguay, Peru, Surinam, Uruguay, Venezuela, Caribbean.

148. Pseudelephantopus Rohr. Perennial herbs; hairs stiff, simple; leaves mostly basal; inflorescences spiciform with clustered sessile heads; involucral bracts 8; corollas zygomorphic with deep inner sinus; anther shortly calcarate; style without node; sweeping hairs acicular; achenes 10-costate, setuliferous, with glands, idioblasts along costae, raphids short-oblong; pappus bristles 5, contorted, base broadened. Two spp., tropical America, P. spicatus (Juss. ex Aubl.) C.F. Baker in Bolivia (La Paz), British Honduras, Chile, Colombia, Costa Rica, Ecuador, El Salvador, Guatemala, Honduras, Mexico, Nicaragua, Panamá, Peru, Venezuela, Caribbean; and P. spiralis (Less.) Cronquist, from Argentina, Bolivia, Brazil, Colombia, Costa Rica, Ecuador, Peru, Venezuela and Caribbean.

■ SUBTRIBE LYNCHNOPHORINAE ▸ before the present study, little attention has been paid to the habit, leaf sheath and capitulescence position in the group. Lychnophorinae are shrubs, treelets, trees, or caulirosulas, the latter being a highly specialized habit to alpine habitat (Andine paramos (Espeletia s.l.) or Chimantá Tepui (Chimantea). Caulirosulate plants evolved independently three times in Brazilian campo rupestre lineages (Prestelia, Proteopsis and Minasia). They possess several peculiar structural features, such as a pseudotrunk covered with leafsheath remains, a dense indumentum, tightly furled leaf rosettes, water-storage strategies, epidermal cell wall and cuticle thickenings, and sclereids. These features are usually found also in other taxa adapted to live in the extreme conditions of the rocky grasslands (campos rupestres, intense solar radiation, great fluctuations in daily temperature, water scarcity during autumn and winter, and the occurrence of fires). Almost all species of Lychnophorinae are endemic to Brazil, with the exception of Eremanthus mattogrossensis Kuntze and E. rondoniensis MacLeish & H.Schumach., occurring in Brazil and Bolivia, Centratherum cardenasii H.Rob. that occurs only in Bolivia and C. punctatum, with a large distribution in several tropical countries.

149. Albertinia Spreng. Branching shrubs; Inflorescence loosely corymbiform, heads pedunculate; involucral bracts 55–60 in c. 3 series; deep receptacular pits enclosing achenes; florets 45–50; anther bases rounded; style with node; sweeping hairs broad-acicular; achene c. 10-costate, raphids subquadrate; pappus capillary, tawny. Pollen non-lophate. Only one sp., A. brasiliensis Spreng, endemic to E Brazil (Sergipe to Rio de Janeiro states).

150. Anteremanthus H. Rob. Shrubs; hairs mostly T-formed; inflorescence thyrsoid, with gradate foliaceous bracts; involucral bracts c. 60 in 5–6 series; florets c. 60; corolla tube short, lobes with contorted hairs; anther bases rounded; sweeping hairs pointed, often septate; achene 8–10-ribbed, densely long-setulose; pappus capillary, with squamellae. Two spp. endemic to Brazil, A. hatschbachii H. Rob. from northern Espinhaço Range in Minas Gerais (a rare species in Brazil by Plantas Raras do Brasil’s book), and A. piranii Roque & F. A. Santana, endemic to southern Bahia.

151. Blanchetia DC. Shrubs, with intermixed long dark multicelular hairs and pale stellate hairs; inflorescences corymbiform with foliose bracts; involucral bracts 25–30 in c. 4 series; receptacle with thin partitions; florets 8–10; filaments inserted near sinuses; anthers not tailed; style without node; sweeping hairs pointed; achenes 10-costate, hairless; pappus deciduous, c. 20 flattened bristles. Two spp., B. heterotricha DC. from NE Brazil (Paraíba to Bahia states), and B. coronata (G.M. Barroso) Loeuille & Pirani, endemic to Bahia state, Brazil.

152. Centratherum Cass. Perennial herbs or shrubs branching near base; hairs T-shaped and simple multicellular; involucral bracts 30–40, in c. 4 series, with distal margins scarious; florets 40–100+; anthers not tailed; sweeping hairs acicular; achenes 8–10-ribbed; pappus of short deciduous awns or lacking. Three spp., C. cardenasii H. Rob. endemic to Bolivia, and Cono Sur, C. repens (Spreng.) Loeuille & Pirani endemic to Brazil, and C. punctatum Cass. widely distribuited; includes the unique herbs in this subtribe.

153. Chronopappus DC. Shrubs; stemfelted with thick-walled fusiformand stellate hairs; leaves bullate, abaxial tomentum of contorted stellate-based hairs; heads sessile in upper axils, c. 8 pubescent subinvolucral bracts; involucral bracts c. 8–10, rather deciduous; florets 8–10; anther bases rounded; sweeping hairs pointed; achenes 10-angled, glabrous; inner pappus of flattened bristles. Two spp., endemic to northern Minas Gerais state in SE Brazil.

154. Eremanthus Less. Shrubs and trees; stems felted or woolly or stellate-lepidote; heads usually in 1 to many dense, globular clusters; involucral bracts 10–40 in 4–7 usually gradate series, innermost deciduous; florets 1–9, 24–26; corolla tube short; anther bases rounded; sweeping hairs pointed, often septate; achenes 3–5-angled, or 10(–20)-costate; pappus bristles 3–5-seriate, broadened to filiform. n = 15, 18. 24 spp. from SE to NE Brazil, two of them (E. mattogrossensis Kuntze and E. rondoniensis MacLeish & T. Schumach.) reaching into E Bolivia; two spp. form Minas Gerais and Bahia states are rare species in Brazil by Plantas Raras do Brasil’s book.

155. Gorceixia Baker. Trees; stems partially winged; hairs stellate; inflorescence corymbiform, of secondary heads containing many sessile heads; secondary involucre of canescent bracts; involucre cylindrical, of 5 or 6 lanceolate subequal bracts; florets 5; corollas glabrous; anther tails small; style without node; sweeping hairs sharp; achenes tetragonous, glabrous, raphids elongate; pappus a collar. Only one sp., G. decurrens Baker, endemic to E Brazil (Bahia, Minas Gerais and Espírito Santo states), the tallest species of this subtribe.

156. Heterocoma DC. Perennial herbs or shrubs, or trees, sometimes acaulescent; hairs appressed; leaves crowded, bases vaginate, lanate of leafbases clasping; large head in each upper axil; corolla tube short; anthers not tailed; sweeping hairs pointed, often septate; achenes 10-costate, with phytomelanin; pappus elements twisted, strap-shaped, deciduous. 6 spp., 4 endemic to Minas Gerais, one endemic to Bahia and one shared in Goiás and Minas Gerais states. Phytomelanins are only found in Asparagales seeds and in Asteraceae fruits of Heterocoma DC., Wunderlichia Riedel ex Benth, and some Asteroideae.

157. Hololepis DC. Shrubs or trees; hairs symmetrically T-shaped. Petioles long; heads axillary, solitary, longpedunculate; subinvolucral bracts c. 8, foliiform, trinervate; involucral bracts c. 30 in c. 4 series; receptacular spines linear; florets 25–35; anther tails lobed; sweeping hairs short-pointed to obtuse; achenes 4-angled, glabrous, raphids subquadrate; pappus of flattened persistent bristles, 2–3-seriate. Two spp. endemic to Espírito Santo and Minas Gerais states in SE Brazil.

158. Lychnocephalus Mart. ex Candolle. Treelets, sometimes candelabriform, rarely subshrubs to shrubs; stems poorly branched; leaves alternate, sessile or petiolate, with semi-amplexicaul or amplexicaul sheath, discolorous, venation eucamptodromous or sometimes parallelodromous or actinodromous; inflorescence an axillary or terminal, pedunculate or sessile, solitary 2–3 syncephalia (third-order); capitulum sessile; florets 4–15; corolla purple, rarely white; corolla lobes glabrous or pubescent. 8 spp., endemic of the central portion of the Espinhaço Range of mountains in Minas Gerais, SE Brazil; one of them is considered a rare species in Brazil by Plantas Raras do Brasil’s book (as Lychnophora).

159. Lychnophora Mart. Usually candelabriform or rosettiform shrubs or small trees; stem felted with fusiform hairs; leaves often crowded, with stellate or asymmetrical T-shaped hairs below; heads sessile in lobular or hemispherical clusters; involucral bracts c. 12–25 in 4–6 series, innermost deciduous; florets 1–12; corollas glabrous; anther bases rounded; sweeping hairs sharp, mostly septate; achenes 4–5-angled and 8–10-veined, glabrous; pappus of flattened, usually twisted bristles or straps, often with squamae. n = 17. 32 spp., SE to NE Brazil, in Bahia to São Paulo and Goiás states; 6 spp. from Minas Gerais and Bahia states are considered rare species in Brazil by Plantas Raras do Brasil’s book (two of then as Lychnophoriopsis).

160. Lychnophorella Loeuille, Semir & Pirani. Shrubs, rarely treelets, sometimes candelabriform; stems densely branched; leaves alternate, sessile, rarely shortly petiolate, with a pad-like sheath, blade usually ericoid, coriaceous, discolorous, margin entire, flat or revolute; inflorescence a terminal, sessile or rarely pedunculate, solitary syncephalium (second-order) or rarely a congested dichasium of glomerules of capitula (L. leucodendron); florets 1–5; corolla purple, tube longer or the same size as limb. 8 spp., endemic to center Minas Gerais and Bahia states, with two of them, one in each state, are considered rare species in Brazil by Plantas Raras do Brasil’s book (as Lychnophora in the work).

161. Maschalostachys Loeuille & Roque. Treelets; stems monopodial, initially leafy, later becoming leafless, with transverse linear leaf-scars following leaf shedding; leaves alternate or in pseudo-rosette at the apex of stems, simple, sessile or petiolate, with semi-amplexicaul leaf sheath, venation eucamptodromous, margin flat; capitula fused in a syncephalium (secondary order inflorescence) surrounded by secondary leafy bracts and organized in loose axillary spikes or frequently in a panicle of spikes (rarely a cyme); cypsela cylindrical or obconic, angled, 10-ribbed, carpopodium annuliform, inconspicuous. Two spp., endemic to Bahia and Minas Gerais state in SE Brazil.

162. Minasia H. Rob. Perennial rosettiform silvery herbs, sometimes caulirosulate; hairsT-shaped; inflorescences scapose, branched, with crowded or pedunculate heads; involucral bracts 50–60 in c. 5 series, with blunt tomentose tips; florets 20–30; anther with minute tails; sweeping hairs pointed, septa few; achene c. 8-veined, at least some setulae; pappus capillary, with squamellae. 7 spp., all endemic to Minas Gerais state in SE Brazil; 4 of 7 are considered rare species in Brazil by Plantas Raras do Brasil’s book.

163. Paralychnophora Sch. Bip. 6 spp. from NE to E Brazil, Pernambuco to Minas Gerais state; two spp. from Bahia state are considered rare species in Brazil by Plantas Raras do Brasil’s book.

164. Piptolepis Sch. Bip. Ericoid shrubs; hairs appearing granular, stellate, sometimes also thick-walled, straight; heads single, sessile or pedunculate; involucral bracts 8–18(–25) in 3–4 series; florets 8–18; anther tails short, lobed; sweeping hairs short-acute; achenes strongly 10–12-costate, furrows with setulae and glandular dots; pappus bristles flattened, bases often broad, sometimes with shorter setae. 17 spp., 14 in Espinhaço Range of Minas Gerais state and 3 in Goiás state.

165. Prestelia Sch. Bip. Perennial acaulous subshrubs, sometimes caulirosulate, rootstock broad; axils lanulose; hairs stellate-based; leaves linear; inflorescences 1 to many pedunculate compound heads, secondary involucre of c. 5 subequal bracts. Individual heads 5–10 in cluster, sessile; involucral bracts c. 12 in c. 2 series; florets 5–6; anthers not tailed; sweeping hairs acute; achenes c. 8-costate, scattered setulae, glands and idioblasts; pappus bristles in 2–3 series, no squamellae. Three spp., endemic to Minas Gerais state in SE Brazil.

166. Proteopsis Mart. & Zucc. ex Sch.Bip. Coarse perennial herbs, sometimes caulirosulate; silvery with appressed elliptic-stellate hairs; leaves smaller above, bases clasping; inflorescences a dense cluster of large heads; involucral bracts c. 60 in c. 6 series, strongly pungent-acuminate; florets c. 80; anthers with broad tails; sweeping hairs acute; achenes 10-ribbed, glabrous; pappus of few deciduous awns. Two spp., SE Brazil, in Minas Gerais and Bahia state.

167. Vinicia Dematt. Erect suffrutex, similar to Lychnophora and Chronopappus. Only one sp., V. tomentosa Dematt, endemic to Serra do Cabral in Minas Gerais state.

8. SUBFAMILY ASTEROIDEAE (1.181/17.360-17.620) ▸ probable topology is [Abrotanella + Doronicum + Senecioneae + [[Calenduleae + [Gnaphalieae + [Astereae + Anthemideae]]] + [Inuleae + [Athroismeae + [Feddeeae + [Helenieae + [Coreopsideae + [[Neurolaeneae + [Tageteae + [Bahieae + Chaenactideae]]] + [[Polymnieae + [Heliantheae + [Millerieae + [Madieae + [Eupatorieae + Perityleae]]]]]]]]]]]]]]]]. 22 tribes.

MEGA CLADE SENECIONICUM 1/3

Three tribes; outsiders tribes: Doronicum Clade (1/c 40; Europe except northern parts, Mediterranean, temperate Asia).

8.1 ASTEROIDEA ▸ TRIBE ABROTANELLOIDEAE (1/20) - a single genus.

168. Abrotanella Cass. Dwarf or prostrate perennial herbs, often forming mats or cushions; leaves alternate, sessile with sheathing base, entire, coriaceous or subcarnose; capitula mostly sessile, small, disciform; florets few, whitish, greenish yellow or red to purplish; marginal florets female, tubular; corolla lobes with central vascular strand and without lateral strands. 20 spp., New Guinea, Australasia, New Zealand, 7 in southern South America from Chile to Tierra del Fuego, Falklands and J. Fernandes.

8.2 ASTEROIDEA ▸ TRIBE SENECIODEAE (140/3,200–3,235) - outsiders unvailable. Senecioneae has na almost cosmopolitan distribution with its primary centers of diversity in southern Africa (ca. 700) and South America (ca. 1,300). Its species occur in almost all environments. They grow in aquatic to desert habitats, from low altitudes to alpine communities, and from arctic regions to tropical areas.

169. Acrisione B. Nord. Shrubs or small trees with large soft pith; leaves alternate, cauline, petiolate, entire with denticulate margins, elliptic-ovate to obovate; capitula several to many, corymbose, radiate, ecalyculate, yellow flowered. Two spp., endemics to central Chile and adjacent Argentina.

170. Aequatorium B. Nord. Erect shrubs or trees, stellate-tomentose in young parts; leaves alternate, petiolate, entire, lanceolate to elliptic-ovate, with margins often dentate or denticulate, coriaceous; Capitula several to numerous, paniculate-corymbose, radiate, calyculate; ray florets white or cream; disc florets perfect; corolla deeply lobed, white or pale yellowish. 17 spp., 16 in Peru, Ecuador and Colombia, and one endemic to south Venezuela.

171. Aetheolaena Cass. 21 spp. from Venezuela to Peru, one up to Bolivia.

172. Angeldiazia M.O. Dillon & Zapata. Annual, dichotomously branched delicate climbing stems; conspicuously pinnatisect leaves; solitary disciform capitula, peduncles 5-10 cm long; involucres lacking calyculi; 12-16 florets with yellow corollas. Only one sp., A. weigendii M.O. Dillon & Zapata, endemic to department of Lambeyeque, Peru

173. Anticona E. Linares, J. Campos & A. Galán. Herbs to 12 cm tall, glabrous. Only one sp., A. glareophila (Cuatrec.) E. Linares, J. Campos & A. Galán, endemic to Department of Huancavelica and Lima, Peru.

174. Arbelaezaster Cuatrec. Erect perennial herb with rhizome; leaves cauline, shortly petiolate, ovate, entire with denticulate or serrate margins; capitula several, paniculatecorymbose, discoid with greenish-yellow florets, calyculate. Only one sp., A. ellesworthii (Cuatrec.) Cuatrec., restrcted of Colombia and Venezuela.

175. Blennosperma Less. Small annual herbs; leaves alternate, sessile, mostly pinnatifid; capitula solitary on naked peduncles, radiate, yellow- or purplish-flowered; receptacle conical; ray florets female, tube reduced; disc florets 5-lobed, functionally male. 3 spp., B. chilense Less. in Chile and two remaining in California.

176. Cabreriella Cuatrec. Scandent glabrous shrubs; leaves opposite, sessile or subsessile, ovate-cordate, entire with dentate margins; capitula several, corymbose, radiate or discoid, yellow-flowered, calyculate; disc florets perfect. Two spp. from Colombia, one up to Venezuela.

177. Caxamarca Dillon & Sagástegui. Perennial herb with foetid tuberous fasciculate roots and unbranched erect stem. Basal leaves rosulate, large, dissected; cauline leaves smaller, sessile, entire with serrate to dentate margins, with decurrent base forming wings; capitula several, corymbose-cymose, radiate, yellow-flowered; disc florets numerous, with deeply lobed corolla. Two spp. endemics to Peru.

178. Chaetacalia Pruski. Glabrous perennial herbs 0.4–1.5 m tall; stems erect, subterete, exalate, 6+-striate, glabrous, sometimes deflected at nodes, leafy into capitulescence, few-branched distally, internodes mostly shorter than leaves, pith solid; neither caudex nor roots seen. Only one sp., C. stylotricha (Cabrera) Pruski, known only from S & C Bolivia in the department of Santa Cruz, where it occurs in openings in semideciduous forests from 525–1,500 m elevation.

179. Chersodoma Phil. Dioecious shrubs, subshrubs or perennial herbs; leaves petiolate or subsessile, often tomentose beneath, entire; margins sometimes dentate, often revolute; capitula solitary, pedunculate, or few and laxly paniculate, homogamous; florets either pistillate with aborted anthers or functionally male with sterile ovary. 11 spp., Andean South America from Argentina, Bolivia, Chile and Peru.

180. Culcitium Bonpl. 15 spp. from Colombia to S Argentina and Chile.

181. Dalairea Lemaire. Two spp., D. odorata Lemaire from South Africa (but has been introduced to most continents, having become naturalised and even a noxious invasive species in Australia, New Zealand and many countries in Asia, Europe and the Americas) and D. aparadensis Funez & Hassemer, endemic to Morro da Igreja, at Urubici municipality, Santa Catarina state, S Brazil.

182. Dendrophorbium (Cuatrec.) C. Jeffrey. Erect suffrutescent herbs (‘dendrophorbs’), shrubs or trees; leaves large, cauline, often assembled apically on branches, entire, petiolate, margins dentate or denticulate; capitula numerous in dense terminal corymbose or paniculate synflorescences. 70 spp., 69 in South America (and one restricted of Caribbean) 62 in Andes of South America from Bolivia to Venezuela, Argentina, Brazil (12, 10 endemic), Paraguay. One sp. from São Paulo state is considered a rare species in Brazil by Plantas Raras do Brasil’s book.

183. Dorobaea Cass. Perennial glabrous herbs with fibrous roots; leaves rosulate, petiolate, lanceolate to ovate, pinnatipartite or laciniate; capitula solitary on long, bracteate peduncles, large, radiate, conspicuously calyculate, yellow- or orange-flowered; corolla lobes lanceolate with a median resin duct. Two spp. from Colombia, Peru and Ecuador.

184. Dresslerothamnus H. Rob. Scandent shrubs with T-shaped or substellate trichomes; leaves cauline, petiolate, elliptic-ovate, entire; capitula several, corymbose-paniculate, radiate, calyculate; rays flagelliform, twisted and recurved, reddish; disc florets perfect, yellow. 5 spp., 4 in Panamá and Costa Rica (one up to Colombia) a one fully endemic to Colombia.

185. Emilia (Cass.) Cass. Annual or perennial herbs with fibrous roots, mostly glabrous; leaves cauline, sessile or petiolate, sometimes amplexicaul; capitula solitary or few to several, corymbose, radiate or discoid, ecalyculate; florets white, pink, red, purple, orange or yellow; involucral bracts uniseriate. 117 spp., tropical regions, mostly in Africa (some tropical weeds); only one sp. in New World, E. fosbergii Nicolson, in over tropical America, restricted for Western Hemisphere.

186. Erechtites Raf. Annual or perennial herbs; leaves cauline, sessile, pinnatilobate or entire with serrate margins; capitula several, corymbose, disciform; involucre cylindrical, calyculate; marginal florets female, filiform; disc florets perfect, white to yellowish. 7 spp., North and South America, Caribbean up to Brazil (6, none endemics), Bolivia and adjacent Cono Sur.

187. Garcibarrigoa Cuatrec. Perennial herbs; leaves petiolate, entire with serrulate margins, elliptic-lanceolate, basally auriculate and stem-clasping, strongly pinnately veined; capitula solitary or few, corymbose, radiate, orange- or yellow-flowered. Two spp. from Colombia, one up to Ecuador.

188. Graphistylis B. Nord. Erect subshrubs or somewhat lignescent herbs, usually pubescent and glabrescent; leaves shortly petiolate or subsessile, entire, elliptic to lanceolate, with serrate or denticulate margins; capitula several to numerous, corymbose-paniculate, radiate, yellow- or rarely white-flowered, calyculate; disc florets perfect, corolla lobes triangular-ovate with a median resin duct. Nine spp., endemic to mountains regions of Minas Gerais, São Paulo, Paraná and Rio de Janeiro states, Brazil; three of them (Rio de Janeiro, Minas Gerais and São Paulo states one each) are considered rare species in Brazil by Plantas Raras do Brasil’s book.

189. Gynoxys Cass. (inc. Nordenstamia) Shrubs or trees; leaves opposite, petiolate or subsessile, entire, elliptic-oblong to ovate or obovate, coriaceous, densely tomentose beneath; capitula few tomany, corymbose-paniculate, radiate or discoid, yellow-flowered. 130 spp., Andes from Venezuela to Bolivia (60 in Peru); the largest genus endemic to South America absent in Brazil.

190. Haplosticha Phil. Three spp. from Argentina and Chile.

191. Hoehnephytum Cabrera. Suffrutescent herbs or shrubs; leaves cauline, petiolate, elliptic, entire; capitula several, corymbose, discoid, few-flowered, yellow, calyculate. Two spp. endemic to Brazil, both collected in Bahia state, with one of then in this state considered a rare species in Brazil by Plantas Raras do Brasil’s book.

192. Iocenes B.Nord. Only one sp., I. virens (Phil.) Pruski., endemic to Patagonian Argentina and Chile, where it is found from 200–1,500 m elevation.

193. Lasiocephalus Willd. ex Schltdl. 4 spp., Colombia, Ecuador and Peru.

194. Lomanthus B.Nord. & Pelser. 20 spp., from Ecuador to NW Argentina.

195. Misbrookea V.A. Funk. Perennial rhizomatous herb formingmats or growing solitary, densely hairy with long strigose trichomes; leaves closely set and appressed, sessile, oblong-obovate, greyish-green; capitula solitary, sessile or subsessile, radiate, ecalyculate; involucral bracts connate; ray florets white; disc florets perfect, yellow. Only one sp., M. strigosissima (A. Gray) V.A. Funk, high Andes of Bolivia and Peru.

196. Monticalia C. Jeffrey. (inc. Pentacalia p.p., Senecio p.p.) Erect shrubs or shrublets; leaves closely set, shortly petiolate to subsessile, often small, ericoid or linear-lanceolate to elliptic-ovate; capitula several to numerous, corymbose, radiate or discoid, mostly yellow-flowered. 15 spp., one only in Central America and remaining from Venezuela to Peru, 7-10 of them restricted of Ecuador.

197. Paracalia Cuatrec. Scandent shrublets; leaves cauline, petiolate, elliptic-ovate, entire; capitula several, corymbose, small and few-flowered, discoid, ecalyculate; corolla white, deeply lobed. Three spp., two in Peru and one in Bolivia.

198. Paragynoxys (Cuatrec.) Cuatrec. Erect single-stemmed or little-branched trees or treelets; leaves alternate, crowded towards branch ends, petiolate, large, entire, elliptic-ovate or obovate, coriaceous; capitula numerous, fewflowered, paniculate, discoid, calyculate; corolla white, deeply lobed. 12 spp. in Andes, 11 in Colombia and Venezuela, and one endemic to Ecuador.

199. Pentacalia Cass. Scandent shrubs or epiphytes, glabrous or pubescent with simple hairs, sometimes cushions in northern Andes; leaves petiolate, oblong to elliptic-ovate, entire with margins often dentate to serrate, often coriaceous; capitula several to many, corymbosely paniculate in terminal or lateral synflorescences, radiate or disciform, yellow- or rarely white-flowered; anthers sagittate to caudate. 224 spp., 15 in Mexico to South America and Caribbean, 2 in Caribbean and South America, and 208 from Venezuela to Bolivia, P. epiphytica (Kuntze) Cuatrec. in Peru to Cono Sur, and only two in E South America, both endemic to Atlantic Forest of SE Brazil.

Brazilian spp. are P. desiderabilis (Vell.) Cuatrec. and P. tropicalis (Cabrera) C.Jeffrey. P. desiderabilis occurs from Bahia to Rio Grande do Sul states, while P. tropicalis is only found in the Bahia, Espírito Santo and Rio de Janeiro states. P. desiderabilis is common in the edges of Atlantic forests, in altitudes ranging from 400 to 2,040m. P. desiderabilis is an attractive liane with leaves often fleshy to subfleshy, showy radiate heads due yours yellow florets in thyrsoid to corymbose paniculate capitulescences.

200. Pseudogynoxys (Greenm.) Cabrera. Scandent herbs or subshrubs; leaves petiolate, entire or rarely lobate, margins often serrate; capitula few to several, occasionally solitary, corymbose, radiate, orange-red- or orange- to yellowflowered, fragrant, calyculate. 15 spp., South America (13) from Bolivia and Brazil (two, none endemic) to Central America and Mexico. P. chenopodioides (Kunth) Cabrera is cultivated as an ornamental and occurs as a garden escapee.

201. Rockhausenia D.J.N.Hind Hind. (inc. Werneria p.p.) Perennial rosulate herbs growing solitary or in small clumps, sometimes cushions; leaves sessile, closely set at rhizome tips or below capitula, entire, glabrous; capitula solitary, sessile or pedunculate, ecalyculate; involucral bracts connate to various degrees; ray florets white or rarely yellow; disc florets perfect, white or yellow. 27 spp., essentially restricted to South America in high Andes of Argentina, Bolivia, Chile, Colombia, Ecuador, Peru and Venezuela, with the exception of R. nubigena (Kunth) D.J.N.Hind, which is found from S Mexico and Guatemala south to S Bolivia.

202. Roldana La Llave & Lex. Herbs, subshrubs or treelets, mostly with a hairy tuberous caudex and fleshy roots; stems with large pith; leaves alternate, petiolate, often peltate, palmately or sometimes pinnately veined, rounded-ovate, entire and dentate to lobate or pinnatisect. Capitula several to numerous, paniculate to corymbose, radiate, disciform or discoid, yellow-, white- or greenish-flowered. 58 spp., SW U.S.A. (Arizona), Mexico to Panamá, with R. jurgensenii (Hemsl.) H. Rob. & Brettell disjunct in Mexico, Central America and Cono Sur.

203. Scrobicaria Cass. Small erect shrubs; leaves cauline, opposite, shortly petiolate or subsessile, entire with dentate margins; capitula few to several, rather densely corymbose, discoid, calyculate, yellow-flowered. Three spp., two in Colombia and one in Venezuela.

204. Senecio L. Herbs, subshrubs, shrubs, or small trees with alternate (sometimes rosulate) leaves, sometimes cushions; involucre campanulate or cup-shaped; capitula radiate, disciform, or discoid; florets often yellow, sometimes white, green, pink, purple, or rarely blue; cypselas homomorphic, 8–12-ribbed, with papillate surface; carpopodium present; pappus bristles numerous, slender, barbellate, white. 1,450 spp., cosmopolitan, 752 in New World, except Caribbean, many in South America (647) and Southern Africa; 387 in Cono Sur, 61 in Brazil (42 endemics), 224 in Chile (the largest Chilean genus of angiosperms); in Brazil is centered of sandy soils of Rio Grande do Sul state. Nine Senecio species from Bahia, Minas Gerais, Espírito Santo, Santa Catarina and Rio Grande do Sul state considered a rare species in Brazil by Plantas Raras do Brasil’s book.

205. Talamancalia H. Rob. & Cuatrec. Herbs, subshrubs or shrubs, erect or ascending, hirsute or lanate, glabrescent; leaves cauline, petiolate with a winged petiole and half-clasping base, pinnatilobate at least proximally, otherwise entire with serrate margins; capitula few to several, laxly corymbose, radiate, orange- or yellow-flowered; disc florets numerous; corolla lobes narrowly oblong. 4 spp., two in Ecuador and two in Panamá and Costa Rica.

206. Werneria Kunth. (exc. Rockhausenia) Perennial herbs forming hummocks or mats, sometimes cushions; rhizomes covered with leaves or leaf bases; leaves closely spirally set, entire and thick or small with lobed tips, bright green at least distally, becoming brown or blackish; capitula solitary, sessile, ecalyculate, radiate; involucral bracts connate at least basally; ray florets white or purplish; disc florets perfect, white or yellow. 24 spp., high Andes from Argentina and Chile to Colombia.

MEGA CLADE ASTEROIDEAE 2/3

Four tribes, three in South America and one, Calenduleae (4/c 119; Central and South Europe, Macaronesia, Mediterranean to Iran), absent.

8.3 ASTEROIDEA ▸ TRIBE GNAPHALIEAE (175/1.995–2.020) - outsiders unvailable.

207. Achyrocline (Less.) DC. Perennial herbs; leaves alternate, flat with entire margins, tomentose on both surfaces; capitula many in corymbs, rarely solitary; involucral bracts papery, coloured, stereome divided; receptacle flat, epaleate; outer florets filiform, yellow; central florets perfect, yellow. 41 spp. in Central and South America (36) up to Brazil (19, 13 endemics), few in Africa, one in Madagascar and Comores. Plantas Raras do Brasil’s book asignated Stenocline heringeri H. Rob, rare from Distrito Federal, but Stenocline is a genus endemic to Madagascar and Mauritius; this account places this spp. as A. heringeri (H.Rob.) Deble & Marchiori.

208. Anaphalis DC. Subdioecious perennial herbs or shrubs; leaves alternate, flat or revolutewith entire margins, tomentose on both surfaces; capitula many in corymbs; involucral bracts papery, white, stereome divided; receptacle flat, epaleate; outer florets filiform, yellow; central florets functionally male, yellow. 110 spp., largest genus in Asian Gnaphalieae; one common sp., A. margaritacea Reiche is dusjunctly distributed in Asia and North America; one rarely collected species, A. chilensis Reiche, is endemic to Chilean Andes in South America; the remaining species of the genus occupy a wide and largely continuous distribution in the mountains of Central, temperate E,and SE Asia.

209. Antennaria J. Gaertn. Dioecious perennial herbs or subshrubs; leaves alternate, flatwith entire margins, tomentose on both surfaces, abaxially only, or rarely glabrous; capitula solitary, only a few together, or many in corymbs; involucral bracts papery, coloured, stereome undivided; receptacle flat, epaleate; female florets filiform, white or purplish; male florets tubular, white or purplish. c. 40 spp., Asia, Europe, 38 in North and South America (3 in continent, one in tropical Andes, two in Cono Sur).

210. Berroa Beauverd. Perennial herb; leaves alternate, flat with entire margins, tomentose on both surfaces; capitula solitary; involucral bracts papery, brownish, stereome undivided; receptacle flat, epaleate; outer florets filiform, purple; central florets perfect, purple. Only one sp., B. gnaphalioides (Less.) Beauverd, Colombia, S Brazil (only Rio Grande do Sul state), Uruguay and Argentina.

211. Chevreulia Cass. Perennial herbs; leaves opposite, flat with entire margins, tomentose abaxially; capitula solitary; involucral bracts papery, brownish, stereome undivided; receptacle flat, epaleate; outer florets filiform, purple; central florets perfect, purple. 6 spp., 5 in southern South America at S Brazil (3, 1 endemic), Bolivia, Paraguay, Uruguay, Chile, N and C Argentina, Falkland Islands, and C. acuminata Less. in over South America.

212. Chionolaena DC. Shrubs; leaves alternate, revolute with entire margins, tomentose on both surfaces; capitula solitary or many in corymbs or dense clusters; involucral bracts papery, coloured, stereome undivided; receptacle flat, epaleate; outer florets filiform, purple; central florets functionally male, purple. 22 spp., 8 in Mexico, Guatemala up to Costa Rica, 2 only in N Colombia/Venezuela, and 12 in S Brazil, 11 endemics (4, from Minas Gerais, Bahia and Rio de Janeiro states, are considered rare species in Brazil by Plantas Raras do Brasil’s book) and one shared with Venezuela.

213. Chryselium Urtubey & S. E. Freire. Rosulate herbs, monoecious, 10–30 cm tall, stoloniferous, stolons 3–11cm long, stems erect, unbranched, leafy, whitish woolly; cauline leaves sessile, linear to narrowly oblanceolate, margin flat, apex acute-mucronate; basal leaves narrowly oblanceolate to oblanceolate; discolorous, upper surface slightly lanuginose to glabrescent, and few biseriate glandular trichomes hidden under the wool. Only one sp., C. gnaphalioides (Kunth) Urtubey & S. E. Freire, known from the tropical Andes of Venezuela, Colombia, Ecuador, Peru, and Bolivia.

214. Cuatrecasasiella H. Rob. Dioecious perennial herbs; leaves opposite, flat with entire margins, tomentose on both surfaces; capitula solitary, sessile; involucral bracts papery, brownish, stereome undivided; receptacle flat, epaleate; female florets filiform, purple; male florets purple. Two spp., one in S Ecuador, Peru, another in Bolivia, Chile and Argentina.

215. Facelis Cass. Annual herbs; leaves alternate, flat with entire margins, tomentose on both surfaces; capitula solitary or only a few together; involucral bracts papery, brownish, stereome undivided; receptacle flat, epaleate; outer florets filiform, purple; central florets perfect, purple. Three spp., S Brazil (only F. retusa (Lam.) Sch. Bip.), Peru, Bolivia, Paraguay, Chile, Uruguay, Argentina, F. retusa (Lam.) Sch. Bip. disjunt also in North America.

216. Gamochaeta Wedd. Annual or perennial herbs, sometimes cushions; leaves alternate, flat with entire margins, tomentose on both surfaces; capitula many in head-like or spike-like clusters; involucral bracts papery, brownish, stereome undivided; receptacle flat, epaleate; outer florets filiform, purple; central florets perfect, purple. 53 spp., North to South America; 47 in South America, 17 in Brazil, 8 endemics.

217. Gamochaetopsis Anderb. & Freire. Perennial herb; leaves alternate, flat with entire margins, tomentose on both surfaces; capitula few together in small, flat-topped glomerules; involucral bracts papery, brownish, stereome divided; receptacle flat; outer florets filiform; central florets perfect. Only one sp., G. alpina (Poepp. & Endl.) Anderb. & S.E. Freire, restricted of Chile and Argentina.

218. Gnaphalium L. Annual or perennial herbs; leaves alternate, flat with entire margins, tomentose on both surfaces; capitula solitary or only a few together; involucral bracts papery, brownish, stereome undivided; receptacle flat, epaleate; outer florets filiform, purple; central florets perfect, purple. About 80 species, cosmopolitan, 26 in New World, 17 in South America, two in Brazil, none endemic.

219. Jalcophila M.O. Dillon & Sagást. Perennial, alpine cushion plants; leaves alternate, slightly revolute with entire margins, tomentose adaxially only; capitula solitary; involucral bracts papery, brownish, stereome undivided; receptacle flat, epaleate; outer florets filiform, purple; central florets perfect, purple. 4 spp., two in Colombia (one up to Ecuador), Peru e Bolivia one species each.

220. Loricaria Wedd. Dioecious shrubs; leaves alternate, concave to involute with entire margins, tomentose adaxially only; capitula solitary or only a few together; involucral bracts papery, brownish, stereome undivided; receptacle flat, often paleate; female florets filiform, yellow; male florets yellow. 20 spp., mountains of Colombia to Bolivia.

221. Lucilia Cass. Perennial herbs, sometimes cushions; leaves alternate, flat with entire margins, tomentose onboth surfaces; capitula solitary or only a few together; involucral bracts papery, brownish, stereome usually undivided; receptacle flat, epaleate; outer florets filiform, purple; central florets perfect, purple. 7 spp., one in northern Andes up to Cono Sur, remaining 6 in southern continent, in S Brazil (5, 1 endemic), Paraguay, Uruguay, N and C Argentina, Chile and Andes of Bolivia.

222. Micropsis DC. Annual herbs; leaves alternate, flatwith entiremargins, tomentose on both surfaces; capitula only a few together; involucral bracts papery, hyaline, stereome undivided; receptacle flat, paleate; outer florets filiform, purple; central florets perfect, purple. 5 spp., S Brazil (3, only Rio Grande do Sul state, none endemics), Paraguay, Uruguay, NE Argentina, and C Chile

223. Mniodes (A. Gray) Benth. (inc. Belloa) Dioecious, perennial, many as alpine cushions; leaves alternate, concave to involute with entire margins, tomentose distally with an adaxial hair tuft; capitula solitary; involucral bracts papery, brownish, stereome undivided; receptacle flat, epaleate; female florets filiform, purple; male florets purple. 25 spp., Venezuela, Colombia, Ecuador, Peru (2 endemics), Bolivia to Cono Sur.

224. Pseudognaphalium Kirp. Perennial, biennial or annual herbs; leaves alternate, flat with entire margins, tomentose on both surfaces; capitula many in corymbs; involucral bracts papery, coloured, stereome divided; receptacle flat, epaleate; outer florets filiform, yellow; central florets perfect, yellow. 103 spp., Africa, Asia, 55 in New World, Central, North and South America (15), New Zealand; two spp. in Brazil, none endemic.

225. Psilocarphus Nutt. Annual herbs; leaves opposite, flat with entire margins, tomentose on both surfaces; capitula only a few together; involucral bracts cartilaginous, hyaline, stereome undivided; receptacle peg-like, epaleate; outer florets filiform, purple; central florets functionally male, purple. 4 spp., all in North America and Mexico but two disjuncts also in Argentina and Chile in South America.

226. Raouliopsis S.F. Blake. Perennial, dense cushions; leaves alternate, concave to involute with entire margins, tomentose distally with an adaxial hair tuft; capitula solitary, sessile; involucral bracts papery, brownish, stereome undivided; receptacle flat, epaleate; outer florets filiform, purple; central florets perfect, yellow. Only one sp., R. seifrizii S.F. Blake, endemic to Sierra Nevada de Santa Marta, N Colombia.

227. Stenophalium Anderb. Perennial herbs; leaves alternate, flat with entire margins, tomentose onboth surfaces; capitula only a few together, in terminal corymbs; involucral bracts papery, white, stereome divided; receptacle flat, epaleate; all florets perfect, yellow. 4 spp., endemics to Brazil.

228. Stuckertiella Beauverd. Annual herbs; leaves alternate, flat with entire margins, tomentose on both surfaces; capitula only a few together, in terminal clusters; involucral bracts papery, brownish, stereome undivided; receptacle flat, epaleate; outer florets filiform, purple; central florets functionally male, purple. Two spp. from Cono Sur, one up to Ecuador.

8.4 ASTEROIDEA ▸ TRIBE ANTHEMIDAE (105/1.860–1.880) - outsiders unvailable.

229. Artemisia L. Annual or perennial herbs, subshrubs or shrubs. Indumentum absent or of basifixed, medifixed or stellate hairs; leaves alternate, variously lobed or dissected, rarely entire; capitula discoid or disciform, usually arranged in a long panicle but this sometimes much reduced and racemose, spiciform or subglobose; receptacle flat to conical, sometimes pilose, epaleate or occasionally paleate. Central florets hermaphrodite and fertile or functionally male; corolla 5-lobed, yellow or sometimes reddish-violet or purple. 472 spp., Northern Hemisphere, South America, Southern Africa, Pacific Islands; 4 spp. occur in South America, in Argentina and Chile: A. copa Phil., A. echegarayi Hieron., A. magellanica and A. mendozana DC.

230. Leptinella Cass. Perennial or facultative annual herbs; indumentum of basifixed hairs; leaves alternate or opposite, pectinate or 1–2-pinnatisect; capitula solitary, discoid or disciform, pedunculate; outer florets female, fertile, tubular, yellow, apically 4-lobed, inflated to form a hollow space between the outer and inner epidermis; inner florets functionally male; corolla yellow, apically 4-lobed. 33 spp., New Guinea, Australia, New Zealand, L. scariosa Cass. in South America (Chile, Argentina), also in Falkland Islands.

231. Soliva Ruiz & Pav. (inc. Cotula) Annual herbs; indumentum of basifixed hairs; leaves alternate, 2–3-pinnatisect; capitula solitary, sessile in leaf axils, disciform; outer florets female, fertile; tube and limb absent; style spinescent at maturity; inner florets functionally male; corolla 3–4-lobed, yellow; style branches fused. 10 spp. in South America, North America and Australia; only one species widely distributed as weed, S. stolonifera (Brot.) Sweet, which occurs in Argentina, Bolivia, Chile; in Peru and Colombia apparently introduced, Uruguay, U.S.A.; also known from France, Portugal, Madeira and Spain (Canary Islands). Brazil has 3 spp., none endemics.

8.5 ASTEROIDEA ▸ TRIBE ASTEREAE (218/3,400–3,440) - outsiders unvailable; subtribes annuncied includes only the tribes with South American genera, without details.

■ SUBTRIBE HINTERHUBERINAE

232. Aylacophora Cabrera. Intricately branching, subaphyllous shrubs; branches glabrous with tomentose grooves; leaves sparse, linear, entire, acute, with evanescente tomentum; heads terminal on branchlets, discoid; receptacular pales oblong-lanceolate, conduplicate, distally ciliate, acuminate; florets 20–25, perfect, yellow, funnelform, lobes lanceolate, erect-spreading. Only one sp., A. deserticola Cabrera, endemic to central Argentina in Neuquen province.

233. Blakiella Cuatrec. Perennial subshrubs, hirsutulous and densely stipitate-glandular; leaves sessile, base subauriculate, margins crenulate, revolute; inflorescences of few clustered long-pedunculate heads; Peripheral florets multiseriate, 200–270, pistillate, yellow, tube subcapillary; disc florets 55–85, functionally male, yellow, tubenarrow, limb campanulate, lobes triangular. Only one sp., B. bartsiifolia (S.F. Blake) Cuatrec., restricted for Colombia and Venezuela.

234. Cabreraea Bonif. Shrubs subglobose, leaves dendsely arranged. Only one sp., C. andina (Cabrera) Bonif., endemic to W & C Argentina, at 2,000-3,000m elevation range.

235. Chiliophyllum Phil. Small ericoid shrubs; stems, leaf undersides and involucres tomentose or glutinous; leaves sessile, linear-oblong to obovate, margins entire to revolute; heads solitary on branchlets, short-pedunculate; involucres broadly campanulate; bracts c. 15–25, persistent, broadly lanceolate to linear-lanceolate, acute, margins lacerate-ciliate; receptacular pales rather conduplicate. Only one sp., C. densifolium Phil., endemic to Argentina.

236. Chiliotrichiopsis Cabrera. Perennial, glutinous, ericoid shrubs; stems white-tomentose, glabrescent, brownish-lined, blackening; leaves sessile, linear to oblanceolate, margins entire, revolute, glabrous above, tomentose beneath except on midvein; heads solitary on short branches, sessile; bracts 4–5-seriate, bases pale, thickened, median nerve green, margins broadly scarious, lacerate-ciliate. Three spp., one in Peru and two in Bolivia and Argentina.

237. Chiliotrichum Cass. Ericoid shrubs; stems, leaf undersides and outer involucral bracts pale tomentose; leaves subcoriaceous, sessile, linear to oblanceolate, entire, glabrous adaxially; heads on clustered ebracteate peduncles; bracts 3–4-seriate, gradate, caducous, mostly indurate, oblong-lanceolate, subentire, narrowly scarious, green patches on outer bracts. Two spp. from Andes of Argentina and Chile.

238. Diplostephium Kunth. Shrubs or small trees; stems, leaf undersides and involucres puberulous to lanate or gland-dotted; leaves usually petiolate, coriaceous, margins plane or revolute; inflorescences 1-headed to corymbose or thyrsoid; pistillate florets 1–3-seriate, white to bluish or purplish; disc florets functionally male or rarely perfect, yellow, green or violaceous, tubular-campanulate, lobes triangular. 55 spp., mainly from Venezuela to Bolivia, two up to Cono Sur, distributed on high mountains zones from Venezuela (1) to Chile (3) with the exception 12 in the Sierra Nevada de Santa Marta in Colombia; it makes part of the high andean forest and the paramo ecosystems. Colombia has the most species with 63, the majority of them found in the Oriental Cordillera, which has 33 registered species until this moment. Diplostephium is the third most diverse genus on the paramos with 70 species after Pentacalia and Senecio.

239. Floscaldasia Cuatrec. Minute herbaceous subshrubs; stems branching, repent, rooting, glabrous; densely foliose branches decumbent or erect; leaves sessile, oblong or trilobed, base vaginate, margins ciliate; heads solitary on erect hirsutulous scapes; bracts 2–3-seriate, subequal, oblong-lanceolate to oblong, herbaceous; receptacles epaleate. Two spp. from Colombia and Ecuador.

240. Flosmutisia Cuatrec. Rhizomatous, rosulate perennial herbs, pilose and minutely stipitate-glandular; leaves sessile, elliptic-oblong to obovate; heads solitary; involucres broadly campanulate; Pistillate florets 2–3-seriate, white, subbilabiate, 5-lobed, outer lobes linear, the two inner lobes filiform; disc florets functionally male, white, funnelform with expanded limb, lobes triangular, erectspreading, pilosulous. Only one sp., F. paramicola Cuatrec., endemic to Colombian paramo.

241. Guynesomia Bonifacino & Sancho. Shrubs, stems glandular; leaves sparse, sessile, linear, clasping, entire, glandular on both surfaces; inflorescence racemiform or thyrsiform; heads c. 15–40, pedunculate, disciform; pistillate florets 10–11, 1-seriate, yellow, limb 2–3mmlong, tridentate, lateral lobes reduced; disc florets 13–16, perfect, yellow, tubular, lobes triangular, spreading. Only one sp., G. scoparia (Phil.) Bonifacino & Sancho, endemic to Chile.

242. Haroldia Bonif. Shrubs more or less globbose. Only one sp., H. mendoncina (Cabrera) Bonif., endemic to N Mendoza, Argentina.

243. Hinterhubera Sch. Bip. Ex Wedd. Small ericoid shrubs, with sessile or stipitate glands and non-glandular hairs; stems defoliated below, densely covered with leaf bases or scars; leaves linear or oblong, margins revolute; heads disciform, sessile or pedunculate on branch tips; Peripheral florets 90–170, pistillate, white or yellow, tubular; disc florets 9–60, functionally male, white or yellow, tube narrow. 8 spp., restricted for Colombia and Venezuela.

244. Katinasia Bonif. Shrubs hemisphaeric; branches more or less dendely disposed. Only one sp., K. cabrerae (Bonif.) Bonif., endemic to Mendoza and Neuquen, Argentina.

245. Kieslingia Faúndez, Saldivia & A.E.Martic. Shrubs with persistent leaves, alternate, tripartite, hirsute, resinous; capitula discoid, homogamous, hemispheric, solitary, shortly pedunculate, terminal on little branches with few leaves, these gradually reduced in size towards the capitulum; corollas tubular, infundibuliform, actinomorphic, with five lobes, reflexed to upcurved in maturity, with thickened margins; cypsela densely pubescent. Only one sp., K. chilensis, restricted to the Andean Pre-mountain Range along the Huasco river basin in the southern portion of the Atacama region, Chile, between elevations of 1,600 and 2,500m.

246. Laestadia Kunth ex Less. Matted herbs, mostly procumbent, gland-dotted, glutinous, sometimes hirtellous; leaves crowded, sessile, sometimes subclasping, oblong to spathulate, entire, 1-nerved. Peripheral florets 2–4-seriate, pistillate, regular with 4 or 5 deeply cut, triangular; disc florets 15–25, functionally male, white to purple, abruptly expanded from short tube, limb cut to near base, lobes triangular. 6 spp., one in paramos of Costa Rica, one endemic to Hispaniola, remaining 4 in Andes from Venezuela to Bolivia, high elevations.

247. Lepidophyllum Cass. Cupressiform, densely branched shrubs; glabrous, glutinous; stems subfrondiform with many short spreading branches; leaves decussate, scale-like, closely abutting, as wide as long, rhombic, with slightly rounded keel; pistillate florets 3 or 4, yellow, radiate or bilabiate with 3 outer lobes short, coiling; disc florets few, perfect, yellow, narrowly funnelform; style appendages linear. Only one sp., L. cupressiforme (Lam.) Cass., in Patagonia of Chile and Argentina.

248. Linochilus Benth. (~ Piofontia) Small trees, shrubs or subshrubs 0.1–10m tall, woody, branching sympodial by substitution with branches terminated by capitulescences; branches cylindrical, minutely ribbed, tomentose or glabrous, glandular or eglandular, striate when old; terminal shoots often tomentose; leaves linear, lanceolate, ellipsoid, oblong, ovate, or obovate; margins entire, denticulate, or serrate, membranous to coriaceous, usually revolute, sometimes flat. 60 spp., and is a major component of the flora of the Northern Andes, the Sierra Nevada of Santa Marta, and the Talamanca Cordillera; the southern boundary of Piofontia is central Ecuador, near the Huancabamba depression, which defines the southern boundary of the northern Andes; most of the species inhabit the paramo ecosystem, but a clade of approximately 16 spp., which contains the species of serie Denticulata sensu Cuatrecasas (1969), occurs in the cloud forest.

249. Llerasia Triana. Shrubs, small trees or vines, often gland-dotted; stems and leaf undersides pale-tomentose; leaves short-petiolate, coriaceous, entire or dentate, pinnately veined; inflorescences corymbose; heads homogamous; Florets 3–15, perfect, yellow, tubular, limb slightly broader, lobes deeply cut, linear-oblong, spreading; style base bulbous, appendages triangular, hairy. 14 spp. from Colombian to Bolivian Andes.

250. Nardophyllum (Hook. & Arn.) Hook. & Arn. (exc. Paleaepappus) Small spreading shrubs, sometimes cushions; stems, leaf undersides, and involucres often tomentose; branchlets sometimes spiniform, internodes often angled or with dark lines; leaves sessile, subcoriaceous, oblong to linear, entire; heads 1–5, terminal or axillary; rays lacking; florets few, perfect, yellow, narrowly funnelform, lobes triangular to lanceolate, often recurved. 5 spp. from Argentina and Chile.

251. Novenia S.E. Freire. Acaulescent, caespitose, perennial herbs; leaves in rosettes, bases involute, densely villous; blades linear-lanceolate, coriaceous, with 3 grooves adaxially, glabrous; inflorescences sessile, glomerulate, 1–4-headed; Pistillate florets 3–9, whitish, filiform, eradiate; disc florets functionally male, whitish, narrowly tubular, lobes oblong-triangular, erect; style branches linear-lanceolate. Two spp. in Andes from N Argentina to S Peru.

252. Ocyroe Phil. Shrubs irregularly branched, branches densely to loosely arranged, divaricate, tomentose, thorny, internodes short; leaves arranged in macro– and brachyblasts, narrowly obovate; capitula solitary, terminal, homogamous, discoid, sessile to shortly pedunculate. Only one sp., O. armata (Wedd.) Bonif., from NE Chile, NW Argentina, SW Bolivia, 2,800 – 4,600m elevation range.

253. Oritrophium (Kunth) Cuatrec. Perennial herbs from thick erect to horizontal rhizome; leaves mostly in basal rosettes, bases usually densely hairy, blades oblanceolate to linear, glabrous to sericeous; inflorescences scapose, 1-headed, bracteoles linear-subulate; bracts 2–3-seriate, gradate to subequal, linear to oblong-lanceolate, obscurely veined; receptacles epaleate. 24 spp., 22 in Andean from Venezuela to Bolivia, and two endemics to Mexico.

254. Paleaepappus Cabrera. (off Nardophyllum) Small, divaricately branched shrubs, Young branchlets thorn-like; young branches, Young leaves and involucres tomentose; leaves small, subcoriaceous, sessile, oblong to spathulate, entire, caducous; heads solitary on bracteose branches, discoid; involucres campanulate; bracts 4–5-seriate, gradate, broadly lanceolate, shortacute, entire, indurate, caducous. Only one sp., P. patagonicus Cabrera, endemic to Patagonia of Argentina.

255. Parastrephia Nutt. Ericoid to cupressiform shrubs, resinous to partly tomentose; leaves alternate, sessile, coriaceous, midvein depressed and tomentose abaxially, remainder of leaf glabrous; heads solitary or in small clusters, disciform; pistillate florets 1-seriate, yellow, tubular or with small limb; disc florets perfect, yellow, narrowly funnelform; style appendages lanceolate, papillose to base of branches. 5 spp. from Argentina, Chile, Bolivia and Peru.

■ SUBTRIBE GRANGEINAE

256. Egletes Cass. Erect to procumbent, taprooted, annual herbs; stems much-branched, viscid with usually stipitate glands; leaf bases petiolate to sessile, subclasping, blades obovate to spathulate, lobed or toothed to pinnatifid or bipinnatifid; heads usually solitary and axillary or few in loosely corymbose inflorescences, short-pedunculate; receptacles conical. 7 spp., 6 from SW U.S.A. (southern Texas) to Guianas and Colombia (all absents in Brazil), and E. viscosa (L.) Less. in over tropical and subtropical New World.

257. Plagiocheilus Arn. ex DC. Erect to creeping perennial herbs; pilose to hirsute on leaf-base margins, leaf surfaces or sometimes on stems; petioles winged with broadened bases, leaf blades pinnately to bipinnately dissected; heads solitary on axillary peduncles or short-pedunculate in small terminal clusters. 6 spp. from Venezuela to Bolivia and Argentina.

■ SUBTRIBE LAGENOPHORINAE

258. Lagenophora Cass. Perennial stoloniferous herbs or subshrubs, often matted; usually puberulous; stems erect or procumbent; leaves cauline or mostly in rosette, sessile to petiolate, linear to spathulate, entire or dentate; heads solitary, long-pedunculate; bracts 2–3-seriate, subequal, more than 10, narrowly oblong, acute or obtuse, herbaceous, entire or denticulate, margins scarious. 15 spp., one widely distributed in Australia, New Zealand (six endemics), Guatemala (one endemic), one from Japan to New Zealand, one endemic to Japan, one endemic to Borneu, one only Australia and New Zealand, and three restricted of Argentina and Chile in South America.

259. Talamancaster Pruski. (inc. Myriactis) Small perennial herbs (rarely collected in first year and appearing as annuals), commonly with rhizomes, subscapose, remotely bracteate-leaved, or leafy to near capitula, never truly scapose; stems single or more commonly few-branched from base, often brownish red, leaves commonly basal and cauline, present at flowering, proximal ones spreading laterally, cauline leaves increasingly appressed distally toward capitula; roots fibrous; herbage with mostly patent simple non-colored trichomes. 6 spp., almost all in Panamá and Costa Rica except T. andinus (V.M.Badillo) Pruski also in Venezuela.

■ SUBTRIBE BACCHARIDINAE

260. Archibaccharis Heering. Perennial herbs, shrubs or vines, functionally dioecious, with stipitate or sessile glands, not glutinous; stems straight, twining or zigzag; leaves petiolate or sessile; inflorescences terminal or axillary, strongly cymose or corymbose, unisexual or vestigially gynomonoecious. Involucral bracts 3–5-seriate, gradate. 36 spp. from Mexico and Central America, one up to N Colombia.

261. Baccharis L. Small trees, shrubs or perennial herbs, dioecious or rarely monoecious, sometimes xylopodial, cushions or with roots crown; stems sometimes winged; leaves rarely opposite, scale-like or absent, lamina linear to ovate, obovate or pinnatisect, glabrous to glandular-dotted and glutinous, rarely tomentose; inflorescence corymbose or thyrsoid, spicate, racemose or 1-headed; heads unisexual, discoid; bracts 3–8- seriate, gradate; receptacles plano-convex, usually epaleate. 402 spp., distributed from southern Canada to southern South America (350), inc. Chile; the most important centers of species richness are the Andes from Colombia to central Chile and central Argentina, and the mountains of SE Brazil (168 in country, 110 endemics), Uruguay, and E Paraguay. 8 spp. from Rio de Janeiro, Minas Gerais, Paraná, Santa Catarina, Goiás and Espírito Santo states are considered rare species in Brazil by Plantas Raras do Brasil’s book.

In Brazil by species common in secondary vegetation and with a wide distribution to narrow and critically endangered endemics from mountain summit. Most Baccharis species are concentrated in the central and eastern regions, growing mainly in savannas of C Brazil (cerrado) and grasslands in high altitud grasslands (campos de altitude), rocky grasslands (campos rupestres) and pampa.

Baccharis L. sect. Caulopterae DC. is represented by c. 30 spp. and is restricted to South America, and it’s characterized by presence of winged stems, epaleaceous clinanthia densely covered with biseriate glandular hairs, papillose glabrous achenes, and pappus bristles of female flowers enlarged basally and fused into a ring. This section occurs mainly in the Andes, from Colombia to the centre of Argentina and in SE South America, encompassing E Brazil, northern Argentina, Paraguay and Uruguay. Brazil is the country with the highest diversity of species for the section, which increases with altitude. Diesel cites 15 spp. of the Trimera group for the state of Rio Grande do Sul.

■ SUBTRIBE PODOCOMINAE

262. Asteropsis Less. Perennial herbs; stems, leaves and involucres loosely tomentose-villous; leaves densely spirally inserted, sessile, non-clasping, 1-nerved, linear, entire; inflorescences terminal, 1- to few-headed; involucres hemispheric, 20–30mm wide; bracts 4–5-seriate, slightly gradate, linear-lanceolate, with long-attenuate, reddish tips. Only one sp., A. megapotamica (Spreng.) Marchesi, Bonifacino & Sancho, Rio Grande do Sul state in S Brazil, and Uruguay, at grasslands.

263. Exostigma Sancho. (off Podocoma) Perennial glabrate to glabrous, somewhat rhizomatous herbs, the roots fibrous; leaves congested at the base, alternate, sessile; capitula in terminal, congested corymbose arrays; ray florets (2–)3-seriate, pistillate, the corolla white; disc florets hermaphroditic, the corolla yellowish, cypselae elliptic. Two spp., E. notobellidiastrum (Griseb.) Sancho occurs in N Argentina, S Bolivia, SE Brazil, E Paraguay, and Uruguay; 0–2,000m, in semi-shade of forest margins, in shade of forest understory, or on river banks among rocks in wet soil; it is apparently common; and E. rivulare (Gardner) Sancho is known from NE Argentina, S & SE Brazil, E Paraguay, and Uruguay; 0–1,200m, in shade, in forest understory, or on riverbanks on rocks and wet soils; it has been indicated as common.

264. Inulopsis (DC.) O. Hoffm. Perennial herbs from rhizomes; leaves and stems glabrate to hirsute-villous, sometimes gland-dotted, resinous; leaves all basal or some cauline, obovate to linear, with 1 or 2 pairs of longitudinal veins from base; heads solitary on scapes; involucre hemispheric to turbinate; bracts 2–4-seriate, outer 1/2–3/4 as long as inner, lanceolate to lanceolate-oblong, herbaceous, 1-nerved. 4 spp. from southern Brazil, two up to Paraguay and E Bolivia.

265. Laennecia Cass. Annual to short-lived perennial, taprooted herbs, white-tomentose or coarsely hairy, often glanddotted; leaves sessile, lanceolate or oblanceolate to oblong, toothed to pinnately lobed, rarely entire; inflorescences spicate or racemose to loosely thyrsoid or corymbose; involucres turbinate; pistillate florets several-seriate, white, filiform-tubular, eradiate and apically fimbriate or short-radiate. 16 spp., SW U.S.A., Mexico to northern South America (5, Andean from Venezuela to Chile and Argentina).

266. Microgyne Grau. Perennial herbs to subshrubs; caudex woody; stems, leaves and involucres gland-dotted, resinous, sparingly hispid and with arachnoid hairs; leaves densely spirally inserted, linear, mostly apically trifid with linear lobes; heads solitary, on long peduncles; bracts 2–3-seriate, subequal, narrowly oblong to linear, herbaceous with hyaline margins, inner bracts white-indurate at base. Two spp., M. trifurcata (Less.) Grau and M. marchesiana Bonifacio & Sancho, both Rio Grande do Sul state in S Brazil, also in Argentina and Uruguay.

267. Podocoma Cass. (exc. Exostigma) Perennial herbs, rhizomatous; usually coarsely pubescent, eglandular. Basal leaves often persistent, cauline leaves clasping; inflorescences 1-headed or loosely corymbose; involucres campanulate, (5–)7–15(–17) mm wide; bracts 3–5-seriate, strongly gradate, stiffly indurate, narrowly lanceolate or oblong-lanceolate, greenish along middle, without orange resin ducts. 7 spp. from Brazil (but only one endemic), six up to Cono Sur, also in Bolivia.

268. Sommerfeltia Less. Perennial herbs or subshrubs, fromwoody caudex; stems, leaves, and involucres with dense sessile or short-stipitate glands; leaves densely spirally arranged, stiff, entire to pinnately dissected, tips spinose; inflorescence 1-headed to loosely corymbose; bracts 3–4-seriate, gradate, lanceolate to linear, thick and indurate, greenish in middle, carinate, without orange veins, sides hyaline. Two spp. from Argentina an Uruguay, one up to southern Brazil.

■ SUBTRIBE SOLIDAGININAE

269. Gundlachia A. Gray. Evergreen, resinous, punctate shrubs; stems non- to many-branched; leaves usually evenly spaced, spreading to appressed, sessile to shortpetiolate, linear to obovate or spathulate, flat to involute-terete; inflorescences terminal, racemose to corymbose or thyrsoid; heads 1–5 in clusters or hidden by leaves. 11 spp., Caribbean, Mexico and U.S.A. (Texas), G. corymbosa (Urb.) Britton ex Bold. also in Venezuela.

270. Gutierrezia Lag. Annual or perennial, taprooted herbs or small shrubs; leaves decurrent, linear to lanceolate or spathulate, punctate and glutinous, glabrous, entire; heads solitary or 3–6 in clusters; involucres cylindric to campanulate; bracts 2–4-seriate, gradate, stramineous, 1- or 2-nerved, bases white-indurate; receptacles flat to columnar, with uncinate hairs. 30 spp., 16 mostly Mexico, also SW U.S.A., 14 endemics to SW South America (Chile, Argentina and Bolivia).

271. Solidago L. Perennial herbs from rhizome or caudex, rarely thickly taprooted; leaves linear or linear-lanceolate to ovate, some 3-nerved, entire to toothed, some gland-dotted; inflorescence cylindric to pyramidal or racemose with secund or arcuate branches; bracts 3–4-seriate, gradate, with an orange-glandular midrib, often with green patch apically; receptacles sometimes with caducous pales. c. 110 spp., 93 mostly North America, 8 in Mexico, 6 in South America, from Bolivia, Chile, Argentina, Brazil (only S. chilensis Meyen), Uruguay, Paraguay, and 10–20 in Eurasia.

■ SUBTRIBE MACHAERANTHERINAE

272. Grindelia Willd. Annual or biennial herbs to subshrubs, taprooted to stoloniferous; with punctate or stipitate glands or eglandular, often glutinous; leaves mostly oblong-lanceolate, entire to spinulose, glandular-toothed or pinnatifid; inflorescences usually 1-headed (corymbose in some Mexican species); involucres campanulate; bracts 4–8-seriate, subequal or gradate, bases usually sclerified, tips erect to spreading or reflexed. 70 spp., U.S.A., Mexico, 20 in southern Andes (Peru southwards) and Rio Grande do Sul state in S Brazil (6, 2 endemics), 3 in Chile.

273. Haplopappus Cass. Perennial herbs, subshrubs or shrubs; usually glabrous and glutinous to stipitate-glandular; stems nearly acaulescent to caulescent and branching; leaves coriaceous, sessile, entire or dentate to pinnately lobed; inflorescences on scapiform peduncles with scale-like bracteoles and 1 head, or corymbose to thyrsoid. 72 spp., one in Ecuador, two in Peru, one in Bolivia and Cono Sur, and remaining restricteds for Cono Sur.

South American Haplopappus sect. Diplostephioides (Benth. & Hook. f.) Blake has been treated as the genus Llerasia Triana; all North American and Central American species identified as Haplopappus have been transferred to other genera; of the 4 sections of South American Haplopappus, sects. Haplopappus and Gymnocoma Nutt. are closely related to each other; sects. Polyphylla Hall and Xylolepis Hall may be congeneric with Hazardia Greene, based on morphological and chemical evidence. Only a single South American species - H. glutinosus Cass., the generic type - has so far been included in molecular-based phylogenetic analyses.

■ SUBTRIBE SYMPHYOTRICHINAE

274. Psilactis A. Gray. Annual or perennial herbs, usually taprooted; stems, involucral bracts and often leaves stipitateglandular, sometimes tomentose. Lower leaves obovate to linear-lanceolate, early deciduous, upper leaves entire, subclasping; inflorescence loosely thyrsoid to corymbose or with branches 1-headed; involucre broad, bracts 2–4-seriate, subequal to gradate, outer bracts indurate along basal margins. 6 spp., mostly U.S.A. (Texas) and Mexico, P. brevilingulata Sch. Bip. ex Hemsl. up to Andes of Peru in northern South America.

275. Symphyotrichum Nees. Mostly perennial herbs, with rhizomes or short caudex, fibrous-rooted; leaves petiolate to sessile and clasping, linear to cordate, entire or toothed; inflorescence cylindrical to diffusely thyrsoid or corymbose; involucres narrowly campanulate to hemispheric; bracts 3–7-seriate, gradate to subequal, mostly with pale indurated bases and herbaceous tips. 93 spp., mostly North America, some Mexico and South America (7), in Peru southwards, inc. Chile and Brazil (3, one endemic), few widely naturalized. 2 subgenera: subg. Symphyotrichum and subg. Virgulus.

■ SUBTRIBE CHRYSOPSIDINAE

276. Noticastrum DC. Perennial herbs or subshrubs, scapiform with basal rosettes or with stolons, decumbent to erect; pubescence non-glandular, stipitate-glandular and/or woolly; leaves often copiously silky near base; inflorescence 1- to few-headed; involucre campanulate to turbinate; bracts 3–6-seriate, gradate, linear-lanceolate, often reddish-margined. 20 spp., Andean from Ecuador to Chile and Argentina, and S Brazil (8, three endemics), one up to Venezuela.

■ SUBTRIBE CONYZINAE

277. Apopyros G.L. Nesom. Erect perennial herbs, from rhizomes or tubers, stiffly pilose-hirsute to nearly glabrous, eglandular; leaves cauline, stiffly erect, shiny-indurate, sessile, entire, 3(–5) -nerved, basal leaves scale-like; inflorescence 1-headed or thyrsoid-corymbose; involucres broadly campanulate; bracts 3–4- seriate, weakly gradate, with resinous midvein, margins narrowly hyaline. Two spp. from Brazil, one up to Paraguay and Argentina.

278. Conyza Less. Annual or perennial herbs, nearly glabrous to coarsely hispid-pilose; leaves linear to oblanceolate, entire to pinnatifid; inflorescences broadly ellipsoid to columnar or corymbose; involucres cylindric to hemispheric; bracts 2–4-seriate, subequal to weakly gradate, outer bractswith 3 resinous nerves; receptacle sometimes a hypanthium-like cup. 60–100 spp., primarily tropical and subtropical, some introduced pantropically; 55 spp. in New World, 5 in South America, 14 in Brazil, two endemics.

Species of Conyza native to Africa apparently are more similar and perhaps more closely related to genera of subtribe Grangeinae; molecular data indicate that American Conyza, as currently viewed, is polyphyletic, having arisen several times from within the nexus of Erigeron L.; the taxonomy of subtribe Conyzinae is currently under review.

279. Darwiniothamnus Harling. Shrubs, glabrous to strigose, sometimes minutely glandular; leaves clustered at ends of branches, narrowly linear to oblanceolate or lanceolate, sessile or short-petiolate, entire to mucronate; inflorescences leafy-bracteate corymbose or nearly 1-headed; buds erect; involucres cupulate to hemispheric. bracts 4–6-seriate, strongly gradate, 1-nerved. Two spp. endemic to Galapagos Islands, Ecuador.

280. Erigeron L. Annual to perennial herbs, rarely shrubs, with caudex or rhizomes, sometimes cushions; stems, leaves, involucres glabrous to variously pubescent, often with stipitate glands. Basal rosettes sometimes persistent, cauline leaves linear to lanceolate or spathulate, entire to pinnatifid; inflorescences 1-headed or corymbose to thyrsoid; involucres turbinate to hemispheric; bracts 2–4(–7)-seriate, equal to strongly gradate, with 1(–3) resinous veins; receptacles flat to conical. 457 spp., Eurasia (from Alpes to SE Asia), 298 in New Wolrd, 234 in North America, 165 confined to North America north of Mexico, 5 confined to Central America, 91 in Mexico (64 endemic, 2 up to Central America and Caribbean, and 25 have part of their range in the U.S.A.); 36 in South America (from Colombia to Argentina, a half in Chile, six of then endemic to Juan Fernandez Archipelago), Caribbean (20), Galapagos, Eurasia.

281. Hysterionica Willd. Annual or perennial taprooted herbs; caudex simple; stems sometimes branched at base; leaves closely inserted, little reduced above, blades oblanceolate, glandular-pubescent; heads 1 to several, peduncles short or long; involucres hemispheric; bracts 1–2-seriate, subequal, lanceolate, acute, glandular-pubescent and hirsute outside. 13 spp. restricted to southern Brazil (6, 5 endemics), mainly in mountains, vertical rockies, canions, highly inaccessible, sometimes above 1,000m elevation range, also Bolivia, Uruguay and north and center of Argentina; two spp. from highlands of Rio Grande do Sul state are considered rare species in Brazil by Plantas Raras do Brasil’s book.

282. Leptostelma D. Don. Coarse perennial herbs, erect from decumbent bases; stems, leaves and involucres often puberulous to hispid; stems often broad with broad pith; leaves cauline, herbaceous, broadly inserted, narrowed to base, rarely pseudopetiolate, blades ovate to oblanceolate, venation pinnate, margins serrate; inflorescences corymbose to rather subumbellate; heads large; peduncles short to long. 6 spp., SE South America in Bolivia to Uruguay and Brazil (4, 2 endemics); one sp. from Itatiaia Massif in Rio de Janeiro state is considered a rare species in Brazil by Plantas Raras do Brasil’s book.

283. Neja D.Don. Perennial herbs, usually from taproot or branched caudex; leaves mostly basal, filiform to linearoblanceolate, crinkly-pilose with long cilia; heads solitary, long-pedunculate or on sparsely bracteolate stems; involucre broadly turbinate to hemispheric; bracts 2–4-seriate, gradate, narrowly triangular. Three spp., two in SE Brazil (both) and adjacent Argentina and Uruguay, one disjunct, endemic to Cuba.

MEGA CLADE HELIANTHICUM 3/3

15 tribes, outsider are Feddeeae (1/1; E Cuba), Chaenactideae (3/21; W North America, Mexico) and Polymnieae (1/5; Canada, C and E U.S.A.)

8.6 ASTEROIDEA ▸ TRIBE INULEAE (58/580–585) - outasiders unvailable.

284. Epaltes Cass. Herbs; leaves alternate or subopposite, dentate, sparsely hairy or glabrous; capitula heterogamous, solitary or few in terminal clusters; receptacle epaleate; florets white or purple. Marginal florets female; corolla filiform; disc florets functionally male; Epaltes is heterogeneous and likely to be polyphyletic; the genus is diagnosed only by reduction or loss of pappus. 10 spp., 3 in Asia, 3 in Australia, Madagascar, Cuba, Mexico and Brazil one endemic each; Brazilian species in E. brasiliensis DC.

285. Pluchea Cass. Shrubs or herbs; leaves alternate or subopposite, dentate to serrate or entire, generally notdecurrent, hairy; capitula heterogamous, disciform, solitary or few to many in corymbs; receptacle epaleate; florets purple; marginal florets female; corolla filiform; disc florets functionally male. 80 spp., pantropical, 20 in New World, 10 in South America, only three in Brazil, two endemics.

286. Pterocaulon Ell. Herbs, sometimes with xylopodium; leaves alternate, dentate to serrate, decurrent into long wings; capitula heterogamous, disciform, terminal, forming dense glomerules or long, more or less dense spikes or racemes of capitula; receptacle epaleate; florets purplish. Marginal florets female; corolla filiform; disc florets perfect, purple. 18 spp., 12 in North to South America (11, all in Brazil, none endemic), Australia and adjacent areas.

287. Pseudoconyza Cuatrec. Shrubs or herbs; leaves alternate, simple, dentate, serrate or lobed, hairy; capitula heterogamous, disciform, in loose or dense corymbs or panicles, or more or less solitary; marginal florets female, in several rows; corolla yellow or purple, filiform; corolla yellow, white or purple; pappus of barbellate, capillary bristles in one row, with patent (type) or adpressed teeth. Only one sp., P. viscosa (Mill.) DÁrcy, from Mexico, Central America, Caribbean, Venezuela, Ecuador, Peru, Bolivia and tropical Old World.

288. Stenachaenium Benth. Herbs; leaves alternate, dentate to serrate, decurrent, tomentose; capitula heterogamous, disciform, terminal, solitary or few; receptacle epaleate; involucral bracts in several rows; florets white, yellow or purple. Marginal florets female; corolla filiform; disc florets perfect. 5 spp., South America in Brazil (all 5, two endemic), Uruguay, Paraguay and Argentina.

289. Tessaria Ruiz & Pavon. Trees; leaves alternate, entire, glabrous or sparsely hairy; capitula heterogamous, disciform, few in loose corymbs; receptacle epaleate; florets pink; marginal florets female; corolla filiform; disc florets functionally male, usually one; corolla lobes longer than tube. 5 spp. from Central America to Argentina, inc. Bolivia (4), Brazil (2, none endemics), Colombia, Paraguay, Peru, Venezuela, in yungas, sandy river margins (often forming large pure stands, often as a pioneer species on exposed substrate), and forests.

8.7 ASTEROIDEA ▸ TRIBE ATROISMEAE (9/80–85) - outsiders Lowryanthus (1; SE Madagascar), Athroisma (12; tropical regions in the Old World), Blepharispermum (16; Africa, Arabian Peninsula to Sri Lanka), Leucoblepharis (1; India), Anisochaeta (1; E Cape, KwaZulu-Natal), Artemisiopsis (1; tropical and S Africa), Symphyllocarpus (1; E Siberia, Manchuria), Anisopappus (35–40; tropical and S Africa, Madagascar, India, S China, Burma and N Thailand).

290. Centipeda Lour. Annual or perennial herbs; leaves alternate, sessile, blades linear to obovate, variously toothed, rarely entire; capitula sessile to shortly pedunculate, terminal, appearing axillary, solitary or in simple monochasial cymes, disciform or radiate. 10 spp., Australia, New Zealand, SE Asia, Madagascar, Mascarenes, Tahyti, Papua New Guinea, and C. elatinoides (Less) Benht & Hook endemic to southern South America, in S Chile and Neuquen province of Argentina.

8.8 ASTEROIDEA ▸ TRIBE HELENIEAE (13/130) - outsiders are Balduina (3; SE U.S.A.), Marshallia (10; U.S.A.), Pelucha (1; Mexico), Plateilema (1; Texas, Mexico), Psathyrotes (3; SW U.S.A., Mexico), Trichoptilium (1; SW U.S.A., NW Mexico), Amblyolepis (1; Texas, Mexico), Baileya (3; SW U.S.A., Mexico), Psilostrophe (7; W U.S.A., Mexico), Tetraneuris (9; North America, Mexico).

291. Gaillardia Foug. Annual or perennial herbs, rarely caespitose, low shrubs with xylopodia, sometimes forming rosettes, rarely strongly aromatic; leaves petiolate or sessile, entire to pinnatifid sometimes semisucculent. 21 spp., 19 in SE Canada, northern Mexico, SW U.S.A., 3 in temperate South America in Brazil (only G. megapotamica (Spreng.) Baker), Argentina, Uruguay and Paraguay.

292. Helenium L. Annual or perennial herbs; leaves alternate, usually sessile, mostly decurrent, blades linear to lanceolate, elliptic; capitula terminal, solitary or in open paniculiform cymes, sometimes on swollen peduncles, radiate or discoid. 33 spp., North (center of diversity) and South America, 9 in continent, all restricted of Cono Sur except H. radiatum (Less.) Bierner, which reaches into S Brazil.

293. Hymenoxys Cass. Annual or perennial herbs, sometimeswith a thickened caudex; leaves sessile or petiolate, entire or mostly trilobed or bipinnate, blades mostly linear or lanceolate, rarely ovate. 25 spp., E and C Mexico, W U.S.A., temperate South America (4), from Peru to Cono Sur, two of then in SE Brazil, none endemic.

8.9 ASTEROIDEA ▸ TRIBE COREOPSIDAE (28/465–475) - outsiders are Diodontium (1; N Australia), Glossocardia (12; SE Asia, Malesia to islands in the Pacific), Trioncinia (1; Queensland), Coreocarpus (6; SW U.S.A., Mexico), Dicranocarpus (1; SW U.S.A., Mexico), Fitchia (7; Polynesia), Goldmanella (1; Central America), Henricksonia (1; Mexico), Leptosyne (3; SW U.S.A., NW Mexico), Moonia (6; S India, Sri Lanka), Narvalina (1; Hispaniola), Oparanthus (3; Rapa Island, Marquesas Islands), Petrobium (1; St. Helena), Koehneola (1; E Cuba), Pinillosia (1; Cuba, Hispaniola), Tetraperone (1; Cuba).

294. Bidens L. Annual or perennial herbs, vines or shrubs, sometimes aquatic; leaves opposite, sometimes alternate towards inflorescence, rarely whorled, simple to pinnately compound, blades deltate to ovate in outline; capitula in terminal, open to congested corymbiform to paniculiform cymes, radiate or discoid. 280 spp., worldwide, 116 in New World, 38 in South America, 18 in Brazil, 8 endemics. B. campanulata Bringel & T. B. Cavalc., endemic to center Brazil, is the unique species of this genus in South America with campanulate flowers.

295. Chrysanthellum Rich. Annual or perennial herbs, sometimes with a woody caudex; leaves basal in a loose rosette, cauline mostly alternate, pinnatifid; capitula terminal, solitary or in open paniculiform cymes, radiate. c. 12 spp., pantropical, apparently not in Oceania, most species in Mexico, only 3 in South America, two only in Ecuador and C. indicum DC. widely distributed, also in Brazil.

296. Coreopsis L; leaves opposite to alternate towards inflorescence, blades simple or 1–3 pinnately compound, triplinerved, pinnate; capitula solitary or in open paniculiform or corymbiform cymes, discoid or radiate. 85 spp., America, 43 in South America, absent in Brazil.

297. Cosmos Cav. Annual or tuberous perennial herbs; leaves opposite, blades deltate, sagittate or ovate in outline, 1–3-pinnatifid or pinnate, rarely simple; capitula terminal, solitary or in very open paniculiform cymes, discoidor radiate. 37 spp., Mexico, Central America and Andean South America (7), with C. caudatus Kunth up to Brazil.

298. Cyathomone S.F. Blake. Shrubs (?); leaves opposite, ternate, biternate or pinnate-ternate; capitula in terminal, paniculiform cymes. Only one sp., C. sodiroi (Hieron.) S.F. Blake, endemic to Ecuador.

299. Dahlia Cav. Tuberous perennial herbs or shrubs, one species epiphytic, sometimes rupicolous; leaves opposite, sometimes whorled, sometimes semisucculent, blades simple to 1–3-pinnatifid, ovate to deltate in outline, rarely cordate; capitula solitary or loosely aggregated in paniculiform cymes, radiate, nodding. 42 spp. from Mexico and Central America, two up to Colombia, widely cultivated.

300. Ericentrodea S.F. Blake. Weak shrubs or vines; leaves opposite, blades ovate to deltate in outline, 2–3-pinnate, segments linear to lanceolate; capitula in terminal, corymbiform cymes, discoid, rarely radiate. 6 spp., Andes, 5 from Colombia to Ecuador and one endemic to Bolivia.

301. Heterosperma Cav. Annual or perennial herbs, weak shrubs; leaves opposite, blades simple to 1–2-pinnatifid, segments linear; capitula terminal, solitary, radiate. 11 spp., North America to Argentina, Caribbean, and Venezuela, slightly centered in Peru, 11 in South America.

302. Hidalgoa La Llave & Lex. Vines or lianas. Leaves opposite, petioles twining at their bases, blades suborbicular in outline, pedate or pinnate with 3–5 or more leaflets; capitula terminal or axillary, solitary or in simple cymes on long peduncles, radiate; involucres campanulate, phyllaries dimorphic, outer fleshy, herbaceous, spreading, inner membranaceous. 4 spp., Mexico and Central America up to Venezuela and Peru, two in South America.

303. Isostigma Less. Perennial herbs, sometimes forming rosettes with well-developed xylopodia; leaves alternate, petiolate or sessile, blades linear to obovate in outline, entire or variously dissected, deeply dentate to twice trifoliolate, segments linear to filiform; capitula terminal, solitary, scapose, radiate, rarely discoid. 12 spp. from Argentina, Bolivia, Brazil (6, 5 endemics), Paraguay and Uruguay.

304. Staurochlamys Baker. Annual herbs; leaves opposite, blades entire, lanceolate; pappus absent. Only one sp., S. burchelii Baker, endemic to north central Brazil, in Maranhão, Piauí, Tocantins and Goiás states.

305. Thelesperma Less. Perennial herbs or weak shrubs; leaves opposite, blades linear or ovate to trullate in outline, 1–2- pinnatifid, segments linear; capitula terminal, solitary on long peduncles or in open paniculiform cymes, radiate or discoid. 14 spp. in SW U.S.A., NE Mexico (all endemic but one), and one also in Argentina and Uruguay.

8.10 ASTEROIDEA ▸ TRIBE NEUROLAENEAE (5/170–175) - outsiders Heptanthus (7; Cuba), Greenmaniella (1; Mexico).

306. Calea L. (inc. Tyleropappus) Perennial herbs sometimes with woody xylopodia, shrubs, sometimes scandent to vine-like or small trees; leaves opposite, rarely alternate, whorled or basal, blades linear to ovate; capitula solitary or in variously thyrsoid, paniculiform or corymbiform cymes, discoid or radiate. 151 spp., neotropical, 142 in South America, 90 in Brazil, 69 endemics; three species from Goiás and Minas Gerais states are considered rare species in Brazil by Plantas Raras do Brasil’s book.

307. Enydra Lour. Perennial herbs, aquatic or of very wet areas, with fistulose stems, rooting at nodes; leaves opposite, blades lanceolate to ovate or obovate, entire to broadly serrate; capitula axillary, solitary, sessile, radiate or disciform. 10 spp., pantropical, 6 spp. in New World, all in South America, two in Brazil, none endemic.

308. Neurolaena R. Br. Annual or perennial herbs, shrubs or small trees; leaves alternate, blades lanceolate to ovate, trullate, sometimes shallowly trilobed; capitula in terminal, paniculiform, monochasial cymes, discoid or radiate. 11 spp. from North America to Central America, except by N. lobata (L.) Cass. from Mexico, Central America, Caribbean, Trinidad & Tobago, Venezuela, Guianas, Colombia, Ecuador, Peru and Bolivia.

8.11 ASTEROIDEA ▸ TRIBE TAGETEAE (30/270–285) - outsiders unvailable.

309. Dyssodia Cav. Annual or perennial herbs; leaves opposite or alternate above, blades pinnatifid, segments linear to oblanceolate with scattered, pellucid glands; capitula terminal, solitary, or in open paniculiform cymes, sometimes syncephalous and the syncephalia resembling single, radiate capitula, discoid or radiate. Involucres cylindric to hemispheric, phyllaries dimorphic in 1–3 series, outer herbaceous (calyculus), inner herbaceous to membranaceous, with pellucid oval or narrowly oblong glands. 8 spp., 4 in North America to Central America, and 4 endemics to Peru.

310. Flaveria Juss. Annual or perennial herbs, shrubs, rarely small trees; leaf blades linear to elliptic; capitula in terminal or axillary congested corymbiform cymes, tightly aggregated into glomerules with peripheral heads having one ray floret, radiate or discoid. 21 spp., New World, only two South America, both widely distributed (both also occur in Brazil); 1 sp. in Australia; introduced elsewhere.

311. Jaumea Pers. Perennial herbs or prostrate shrubs of estuarine sandy areas or salty mud flats; leaves opposite, blades linear to terete, succulent; capitula terminal, appearing axillary, solitary, radiate or discoid. Two spp., J. carnosa (Less.) A. Gray in coastal W U.S.A., NW Mexico, and J. linearifolia (Juss.) DC. in coastal, central Argentina, Uruguay and Rio Grande do Sul state in S Brazil.

312. Pectis L. Annual or perennial herbs, sometimes stoloniferous; leaves opposite, with pairs of setaceous bristles, blades linear to narrowly ovate, abaxial surfaces with large glands. 95 spp., tropical and subtropical America, 37 in South America, 15 in Brazil, 9 endemics.

313. Porophyllum Adans. Annual or perennial herbs or densely branched shrubs, sometimes weak stoloniferous shrubs; leaves opposite or alternate, blades linear to ovate, with embedded, pellucid glands, herbage strongly aromatic. 29 spp., tropical and subtropical New World, 12 in South America, 8 in Brazil, two endemics.

314. Schizotrichia Benth. in Benth. & Hook. f. Shrubs; leaves opposite, blades narrowly ovate to ovate or elliptic, entire to deeply serrate, with scattered orange glands. Only one sp., S. eupatoioides Benth., endemic to Peru.

315. Tagetes L. Annual or perennial herbs, sometimes shrubs; leaves opposite, simple or pinnately dissected, blades linear to lanceolate or ovate in outline, segments sometimes reduced to bristles, glands scattered throughout blade. 51 spp., tropical and subtropical America, a few species adventive and widely distributed in temperate and tropical regions of the Old World; all spp. in New World, 29 in South America, only T. ostenii Hicken in Brazil, also in Cono Sur and Bolivia.

316. Thymophylla Lag. Annual or perennial herbs, sometimes persisting as weak shrubs; leaves opposite below or alternate throughout, blades linear or filiform, spathulate, entire or pinnatisect, segments linear or filiform, essentially without setae, glands scattered. 13 spp. from Mexico and U.S.A., with T. pentachaeta (DC.) Small disjunct in W Argentina in South America.

8.12 ASTEROIDEA ▸ TRIBE BAHIEAE (23/c. 92) - outsiders unvailable; tropical and southern Africa, Rapa Island, tropical to warm-temperate America, with their highest diversity in Mexico; the monotypic genus Apostates is endemic to Rapa Island, French Polynesia whereas Hypericophyllum is found in tropical Africa.

317. Bahia Lag. Annual, biennial or perennial herbs, sometimes rhizomatous; leaves opposite, blades linear to ovate, entire to 1–3-pinnate, segments filiform to linear; capitula terminal, solitary or in open paniculiform cymes, radiate. 7 spp., W U.S.A., Mexico, and B. ambrosioides Lag. endemic to Chile.

318. Holoschkuhria H. Rob. Annual or perennial herbs, weak shrubs; leaves opposite, blades simple, ovate to elliptic; capitula in terminal, open paniculiform cymes, sometimes decussate, capitula discoid. Only one sp., H. tetramera H. Rob., from northern Peru.

319. Nothoschkuhria B.G. Baldwin. Only one sp., N. degenerica (Kuntze) B.G.Baldwin, from Bolivia to NW Argentina.

320. Schkuhria Roth. Annual herbs; leaves alternate, filiform or linear, ovate to trullate in outline, 1–3-pinnate, segments linear; capitula terminal, solitary or in open paniculiformcymes, discoid or radiate. 3 spp., one endemic to Mexico and S. schkuhrioides Thell. from S U.S.A to Argentina and Brazil.

8.13 ASTEROIDEA ▸ TRIBE HELIANTHEAE (124/1,650-1,685) - outsiders unvailable; subtribes annuncied includes only the tribes with South American genera, without details. Herbs, or shrubs, rarely tree -like; leaves often opposite, sometimes alternate, pubescent often rather rough and scabrid, rarely glabrous or glabrescent; capitula often radiate, sometimes disciform or discoid rays usually broad; phyllaries 1-2-multi-seriate, when uniseriate often conspicuously gland-dotted; receptacle often scaly, sometimes naked; corolla-lobes short; flower colour usually yellow, sometimes purple, disc sometimes dark-coloured; anther-bases more or less obtuse; style arms truncate or appendiculate. Achenes usually black with phytomelanin in walls; pappus usually of awns or scales, sometimes absent, rarely of hairs, sometimes plumose.

■ SUBTRIBE AMBROSIINAE

321. Ambrosia L. Annual or perennial herbs, shrubs; leaves opposite or upper alternate, blades lobed or dissected, deltate, ovate to subrotundiformin outline, triplinerved; capitula discoid, unisexual, sessile, pistillate capitula solitary or in clusters at base of spiciform cymose inflorescences of functionally staminate capitula. 45 spp., Canada to Chile, introduced elsewhere; 11 in South America, 6 in Brazil, one endemic.

322. Parthenium L. Annual or perennial herbs, shrubs, rarely small trees; leaves alternate, petiolate, blades unlobed or lobed sometimes 2 times pinnate, ovate to subsagittate; capitula in terminal, simple, open or congested paniculiform cymes, radiate. 16 spp., over America, 3 in South America, 1 endemic to Bolivia, 1 in Bolivia and Cono Sur, and P. histeriophorus L. widely distribuited, inc. Brazil.

323. Xanthium L. Annual herbs, sometimes spiny; leaves alternate, petiolate, blades ovate to deltate, simple or lobed, triplinerved; capitula discoid, unisexual, sessile, pistillate solitary or in clusters at base of terminal or axillary spiciform cymose inflorescences composed of functionally staminate heads; phyllaries of functionally staminate capitula free. 5 species of disturbed habitats, warm and temperate regions of the world; 4 in South America, two in Brazil, none endemics.

■ SUBTRIBE ECLIPTINAE ▸ currently there are two distinct concepts for Aspilia Thouars and Wedelia Jacq. The first concept separates Aspilia and Wedelia by neutral ray flowers vs. fertile ray flowers, respectively; this concept was adopted by Santos (2001) who added the presence of a scar at the base of cypselae in Aspilia of Brazil (vs. its absence in Wedelia), a taxonomic position followed by the Brazilian researchers. The second concept considers only Wedelia as a valid genus encompassing all Aspilia species. Studies of molecular phylogeny are being conducted in the Aspilia and Wedelia complex in order to recover monophyletic groups (Alves in prep) and for this, new nomenclatural arrangements are required as follow below.

324. Aspilia L. Shrubs, sometimes xylopodial; 70 spp., from Brazil (65, 52 endemics), 9 in Brazil and adjacent Cono Sur, 4 only in Cono Sur, and A. jelskii (Hieron.) S.F. Blake endemic to Peru; 7 spp., all in center states, are considered rare species in Brazil by Plantas Raras do Brasil’s book.

325. Baltimora L. Annual erect herbs; leaves opposite, petiolate, blades ovate, triplinerved; capitula in terminal, open paniculiform cymes, radiate; involucres cylindrical to campanulate, phyllaries in 2 series, subequal to shallowly gradate; receptacles flat to convex. Two spp., Neotropics, both in South America, only the widely distribuited B. recta (Brandegee) Stuessy in Brazil.

326. Blainvillea Cass. Annual or perennial erect herbs, weak shrubs; leaves opposite or alternate, petiolate, blades lanceolate to broadly ovate, triplinerved; capitula in terminal, open paniculiform cymes, radiate; involucres subcylindrical, campanulate to oblong, sometimes hemispheric, phyllaries in 2–4 series, chartaceous with green longitudinal striae; receptacles flat to minutely convex. 10 species, pantropical, only one in New World, B. dichotoma (Murray) Cass. ex Hemsl., scattered, in Venezuela, Guianas, Ecuador, Bolivia and Brazil.

327. Calyptocarpus Less. Prostrate to weakly erect perennial herbs or weak shrubs; leaves opposite, petiolate, blades ovate to lanceolate, triplinerved; capitula terminal and axillary, solitary or in simple cymes, radiate; pappus of 2 divergent or reflexed awns. x = 12. Three spp., two in southern U.S.A., Mexico, Central America, Cuba up to Venezuela, and C. brasiliensis (Nees & Mart.) B. Turner in Brazil, Argentina, Paraguay and Uruguay.

328. Clibadium L. Shrubs or trees; leaves opposite, sessile topetiolate, blades lanceolate to broadly cordiform, triplinerved; capitula in terminal open or congested paniculiform to corymbiform, sometimes glomerule like cymes, disciform, rarely epaleate; involucres cylindric, campanulate, hemispherical, subequal, membranous, scarious, the inner enclosing the marginal cypselae, greenish to purple, sometimes white. 40 spp., neotropics, 34 in over South America, 3 in Brazil, none endemic.

329. Delilia Spreng. Erect or decumbent annual herbs; leaves opposite, blades ovate to lanceolate, triplinerved; capitula in terminal and axillary umbelliform cymes, radiate; involucres flattened, disc-like, plano-compressed, phyllaries 2–4, herbaceous, one orbicular the others suborbicular, ovate to obovate. Three spp., D. biflora (L.) Kuntze widely distribuited in tropical America, the other two are endemic to the Galápagos Islands.

330. Dimerostemma Cass. Erect annual or perennial herbs, shrubs, sometimes with xylopodium; leaves opposite or alternate, petiolate or subsessile, blades narrowly lanceolate to ovate, sometimes subcordate, triplinerved; capitula solitary or in paniculiform cymes, discoid or radiate; involucres broadly campanulate to hemispheric, phyllaries in 3–4 series; receptacles convex. 26 spp. from Bolivia, Paraguay (secondary centre), Argentina, Uruguay and Brazil (22, 11 endemics); two spp. from Goiás state are considered rare species in Brazil by Plantas Raras do Brasil’s book.

331. Eclipta L. Erect or decumbent annual or perennial herbs; leaves opposite, blades lanceolate, narrowly ovate or oval, pinnately veined or triplinerved; capitula axillary or terminal, solitary or in simple cymes, radiate; involucres campanulate to hemispherical, phyllaries in 2–3 series, subequal; receptacles shallowly convex. 4 spp., E. prostrata (L.) L. widely distribuited, one only Colombia, and two in Brazil and adjacent Cono Sur.

332. Elaphandra Strother. Perennial herbs or shrubs; leaves opposite, petiolate, blades lanceolate to ovate, triplinerved; capitula in terminal, simple dichasial or open paniculiform cymes, radiate, rarely discoid; involucres hemispherical, phyllaries in 2–3 series, subequal to gradate; receptacles flat to convex. 14 spp., 1 in Panamá, 13 in tropical Andes of South America and Trinidad & Tobago, only one in Brazil, E. ulei (Hieron.) H. Rob., also in Bolivia.

333. Eleutheranthera Poit. ex Bosc. Erect annual herbs; leaves opposite, petiolate, blades ovate to trullate, triplinerved; capitula axillary, simple, nodding, radiate or discoid; involucres campanulate, phyllaries in 1 series, herbaceous; receptacles flat. Two spp., E. ruderalis (Sw.) Sch. Bip. over neotropical (absent in Brazil, however), adventive in tropical regions of the Old World, and E. tenella (Kunth) H. Rob. endemic to Colombia.

334. Idiopappus H. Rob. & Panero. Small trees; leaves opposite, petiolate, broadly ovate to subcordate, triplinerved; capitula in terminal, open paniculiform to thyrsoid cymes, radiate; involucres hemispherical to shallowly patent with age, phyllaries in 3 series, subequal; receptacles conical to subglobose with age. Only one sp., I. saloyensis (Domke) H. Rob. & Panero, endemic to Ecuador.

335. Kingianthus H. Rob. Shrubs; leaves alternate, petiolate, blades ovate, triplinerved; capitula in terminal, paniculiform cymes of congested corymbiform cymes, radiate; involucres campanulate, phyllaries in 1–2 series, subequal; receptacles convex. Two spp., endemics to Ecuador.

336. Lasianthaea DC. Perennial herbs or shrubs, rarely small trees; leaves opposite, petiolate or subsessile, blades lanceolate to ovate or elliptic, triplinerved; capitula terminal, solitary or in open to congested corymbiform or paniculiform cymes, radiate; involucres cylindric to hemispheric, phyllaries in 3–5 series, subequal to imbricate, distal apices sometimes chartaceous, red, purple or yellow; receptacles convex. 12 spp., mainly from SW U.S.A., Mexico, Central America, L. fruticosa (L.) K.M. Becker also in Venezuela.

337. Leptocarpha DC. Perennial herbs or weak shrubs; leaves opposite or alternate, blades ovate, triplinerved; capitula in terminal, simple dichasia or open paniculiform cymes, radiate; involucres hemispherical to patente with age, phyllaries in 2 series; receptacles convex with linear paleae. Only one sp., L. rivularis DC., endemic to Chile.

338. Melanthera Rohr. Annual or perennial herbs, scandent or erect shrubs; leaves opposite, blades narrowly obovate to linear-lanceolate, ovate, triplinerved, unlobed, trilobed or pinnately lobed; capitula in terminal, open paniculiform cymes, rarely solitary, discoid or radiate; involucres campanulate to hemispherical, phyllaries in 1–2 series, subequal; receptacles convex. 5 spp., North America, Mexico, Caribbean, Central America, South America (3), also in Africa; two spp. in Brazil, none endemics.

339. Monactis Kunth. Shrubs or small trees; leaves alternate, petiolate, blades lanceolate to broadly ovate or trullate, pinnately veined or triplinerved; capitula in terminal, congested to open corymbiform cymes, radiate or discoid, only one or two ray florets per head, ray florets towards the outside of congested corymbiform cymes; involucres cylindrical to campanulate, phyllaries in 1–2 series, gradate. 12 spp. from Colombia, Ecuador and Peru.

340. Oblivia Strother. Scandent shrubs; leaves opposite, petiolate, blades lanceolate-elliptic to ovate, 3–5-plinerved; capitula in terminal, congested corymbiform cymes, radiate; involucres campanulate, phyllaries in 2– 3 series, subequal; receptacles convex. Three spp. from Panamá and Venezuela to Bolivia.

341. Otopappus Benth. Shrubs or small trees; leaves opposite, petiolate, blades lanceolate to ovate, 2–7-plinerved; capitula terminal, solitary or in small paniculiform to corymbiform cymes, radiate or discoid; involucres campanulate to hemispherical, phyllaries in 3–7 series, gradate; receptacles convex to conic. 17 spp., 16 in Mexico and Central America, and one endemic to Colombia.

342. Oyedaea DC. Shrubs or small trees; leaves opposite, petiolate, blades elliptic to ovate, pinnately veined or triplinerved; capitula terminal, in simple or open paniculiform cymes, radiate; involucres campanulate to hemispherical, phyllaries in 2–5 series; receptacles shallowly convex to convex. 19 spp. from Guianas and Venezuela to Bolivia, one reaching into Central America.

343. Pascalia Ortega. Stoloniferous perennial herbs; leaves opposite, sessile to shortly petiolate, blades lanceolate to broadly ovate or oval, triplinerved; capitula terminal, solitary or in simple dichasia, radiate; involucres hemispherical, phyllaries in 2–3 series; receptacles convex to hemispherical. Only one sp., P. glauca Gómez-Ortega is said to be native to southern South America, where it is currently widely distributed from northern Chile and Paraguay through the Argentinian ‘pampas’ to southern Brazil and Uruguay; it was introduced in North America, SW Europe, India, SE Australia, New Zealand and South Africa, and it is regarded as an invasive weed, very dangerous for grazing livestock, which causes acute lethal hepatotoxicosis when ingested.

344. Perymenium Schrad. Perennial herbs, shrubs or small trees; leaves opposite, blades lanceolate to ovate sometimes cordiform, triplinerved, rarely pinnately veined; capitula in terminal, simple dichasia or open paniculiform cymes, radiate; involucres hemispherical, phyllaries in 2–4 series; receptacles convex. 64 spp., Mexico (centre of diversity) to Peru; 10 spp. in South America.

345. Podanthus Lag. Dioecious shrubs, sometimes with purplish stems, forming lignotubers; leaves opposite, blades ovate to trullate; capitula terminal, solitary, or in simple or leafy paniculiformcymes, discoid; involucres patent or reflexed, phyllaries in 1–2 series, subequal; receptacles convex to globose; corollas yellow or green-yellow. Two spp., endemics to Chile.

346. Riencourtia Cass. Annual or perennial herbs, sometimes with xylopodium; leaves opposite, petiolate or sessile, blades linear to elliptic or ovate, 1–5-veined; capitula in terminal, open paniculiform cymes of tightly clustered glomerule-like cymes, disciform; involucres cylindrical, phyllaries in 3–4 series, subequal; receptacles flat, epaleate. 4 spp. from Brazil, one endemic, two into adjacent countries of northern South America and one up to Central America.

347. Schizoptera Turcz. Erect annual herbs; leaves opposite, petiolate, blades ovate to oval, apices acuminate, triplinerved; capitula in paniculiform cymes with subumbellate paracladia, on pedicels of various lengths, radiate; involucres campanulate, phyllaries in 2 series, subequal; receptacles flat to shallowly convex. Only one sp., S. peduncularis S.F. Blake, restricted of Ecuador and Peru.

348. Sphagneticola O. Hoffm. Perennial herbs, prostrate, rooting at nodes; leaves opposite, blades ovate to trullate, trilobed, triplinerved; capitula terminal, appearing axillary, solitary, radiate; involucres turbinate, phyllaries in 2 series, subequal, apices foliaceous, expanded; receptacles convex to conical. 4 species, pantropical, three in New World, two in South America, both widely distribuited, both in Brazil.

349. Steiractinia S.F. Blake. Shrubs or small trees; leaves opposite, blades lanceolate to ovate, pinnate or triplinerved; capitula in terminal, simple or open paniculiform cymes, radiate; involucres hemispherical, phyllaries in 2–4 series; receptacles convex. 12 spp. from Colombia, one up to Ecuador another up to Venezuela.

350. Synedrella Gaertn. Erect to slightly decumbent annual or perennial herbs; leaves opposite, shortly petiolate, blades ovate to elliptic, triplinerved; capitula in axillary, simple cymes, radiate; involucres cylindrical, phyllaries in 2 series. Only one sp., S. nodiflora (L.) Gaertn., Mexico, Central America, Caribbean, Trinidad & Tobago, Venezuela, Guianas, Brazil, Ecuador, Peru, Bolivia, Cono Sur, adventive in other tropical regions.

351. Synedrellopsis Hieron. & Kuntze. Prostrate to decumbent annual or perennial herbs rooting at the nodes; leaves opposite, petiolate, blades ovate to trullate, triplinerved; capitula axillary, solitary, discoid; florets 4, those subtended by the opposing phyllaries tubular, pistillate, corollas 3- or 4-lobed, the internal bisexual, corollas 4-lobed, yellow and uniformly thickened with sclerified cells. Only one sp., S. grisebachii Hieron. & Kuntze, from Argentina, Bolivia and Paraguay.

352. Tilesia G. Mey. Perennial herbs, mostly scandent shrubs, stems striate; leaves opposite, blades lanceolate, ovate or oval, pinnately veined or triplinerved; capitula in terminal, simple or paniculiform cymes, discoid or radiate; involucres hemispherical, phyllaries in 2–3 series; receptacles shallowly convex. Three spp., T. baccata (L.) Pruski over range of genus (Central and South America), T. rubens (Alexander) Pruski only in Guianas, and T. macrocephala (H. Rob.) Pruski in Venezuela, Colombia and Ecuador.

353. Trigonopterum Hook. f. Shrubs; leaves opposite, blades linear and slightly involute; capitula terminal, solitary, radiate or discoid; involucres campanulate to hemispherical, phyllaries in 2 series; receptacles convex. Only one sp., T. laricifolia (Hook. f.) W.L. Wagner & H. Rob., endemic to Galápagos Islands in Ecuador.

354. Tuberculocarpus Pruski. Perennial herbs or subshrubs; leaves opposite, blades lanceolate, pinnately veined; capitula terminal, solitary or in simple cymes, radiate; involucres campanulate to hemispherical, phyllaries in 2–3 series; receptacles flat to shallowly convex. Only one sp., T. ruber (Aristeg.) J.F. Pruski, endemic to the Guiana Shield of southern Venezuela, 50 - 200m elevation range.

355. Wedelia Jacq. Erect to prostrate annual or perennial herbs, shrubs, sometimes with xylopodium; leaves opposite, blades lanceolate to ovate, oval or elliptic, subcordate, usually triplinerved, sometimes trilobed, rarely deeply serrate; capitula terminal, solitary or in open paniculiform cymes, radiate, rarely discoid; involucres campanulate to hemispherical, phyllaries in 2–4 series; receptacles convex. 83 spp. of New World, 55 spp. in South America, 29 in Brazil, 22 endemics. W. souzae H.Rob. from Goiás state is considered a rare species in Brazil by Plantas Raras do Brasil’s book.

356. Zexmenia La Llave & Lex. Scandent to erect perennial herbs, shrubs or lianas; leaves opposite, blades ovate-elliptic to lanceolate, pinnately nerved or triplinerved; capitula in terminal, simple dichasia or umbelliform cymes, radiate. Involucres broadly campanulate to hemispheric, phyllaries in 2–3 series; ray florets pistillate, corollas yellow to yelloworange; disc florets bisexual, corollas yellow to orange with or without a few fibres embedding the vascular strands. Three spp., two in Mexico and Central America, and Z. foliosa Rusby ex W.W. Jones from Bolivia to N Argentina and Paraguay.

■ SUBTRIBE ENCELIINAE

357. Encelia Adans. Annual or perennial herbs, shrubs; leaves alternate, blades linear to ovate sometimes laciniate, triplinerved or with a single vein; capitula terminal, solitary or in open paniculiform cymes, rarely in a corymbiformcyme, radiate or discoid; involucres campanulate to hemispherical, phyllaries in 2–4 series, subequal; receptacles shallowly convex. 20 spp., 16 in Mexico and W U.S.A., 1 in Galápagos Islands, 3 from Peru (two endemics), Chile and Bolivia.

358. Flourensia DC. Shrubs or small trees; leaves alternate, blades lanceolate to ovate, pinnately nerved, usually with resinous exudates, entire to broadly dentate; capitula terminal, solitary or in paniculiform or corymbiform cymes, radiate or discoid; involucres turbinate, campanulate to hemispheric, phyllaries in 2–5 series, subequal to gradate; receptacles flat to convex. 28 spp., 13 spp. in Mexico, SW U.S.A., 15 spp. in south-central Andes and interior of central Argentina (Peru southwards) up to Chile.

■ SUBTRIBE ENGELMANNIINAE

359. Borrichia Adans. Stoloniferous shrubs of muddy or sandy coastal marshes; leaves opposite, blades obovate, succulent, perfoliate, entire to broadly serrate; capitula terminal, solitary, radiate; involucres hemispheric, phyllaries in 2–4 series, subequal; receptacles convex, paleae lignified, apices somewhat acicular, capitulum echinate. Three spp., two only in tropical and subtropical coasts of North America and Caribbean, and the B. peruviana (Lam.) DC. endemic to Peru.

■ SUBTRIBE HELIANTHINAE

360. Aldama La Llave. (inc. Viguiera p.p., Garcilassa, Rhysolepis) Erect rarely decumbent annual or perennial herbs, shrubs, rarely trees, sometimes xylopodial; leaves opposite or alternate, linear to ovate, sometimes cordate or deltate; capitula solitary or incongested to open paniculiform cymes, radiate, rarely discoid; involucres cylindrical to hemispherical, phyllaries in 2–7 series, subequal to prominently gradate; receptacles flat to convex, rarely conical. 118 spp., over America, inc. Chile, 77 in South America, 45 in Brazil, 36 endemics; one sp. from Mato Grosso do Sul state is considered a rare species in Brazil by Plantas Raras do Brasil’s book (as in Viguiera). Mexico has 40 spp., 39 endemics.

361. Heiseria E.E.Schill. & Panero. (off Viguiera) Annuals, 1.1–6 dm tall; stems striate, glabrate to sparsely hispid-pilose; leaves opposite, blades ovate to lanceolate, 15–70mm long, 4–27 mm wide. Two spp., endemic to Andean Peru.

362. Lagascea Cav. Erect annual or perennial herbs, erect or scandent shrubs; leaves opposite, blades lanceolate to ovate to oblanceolate or elliptic; capitula of 1 or 2 (rarely 8) florets, aggregated into tight glomerule-like clusters and subtended by leafy bracts producing a compound capitulum, discoid; corollas white, pink or purple-pink, yellow or orange-yellow; anthers black, pink or yellow. 8 spp., over neotropical, most species in southern Mexico, introduced elsewhere; two spp. in South America, one endemic to Colombia and L. mollis Cav. widely distributed in tropical America.

363. Pappobolus S.F. Blake. Annual or perennial herbs, shrubs, small trees; leaves opposite or alternate, blades linear to broadly ovate; capitula solitary or in simple dichasial or monochasial to paniculiform or corymbiform cymes, sometimes subaxillary, radiate; involucres campanulate to hemispherical, phyllaries in 3–6 series, subequal to gradate, sometimes outermost spreading and leaf-like. 38 spp. from mountains of Peru and Ecuador, one also in Colombia.

364. Scalesia Arn. Shrubs, trees; leaves alternate, blades lanceolate to ovate, rarely cordate, unlobed to deeply lobed or 2-pinnate; capitula solitary or in corymbiform cymes, discoid, disciform or radiate; involucres campanulate to hemispherical, phyllaries in 2–4 series, subequal to gradate, sometimes outermost spreading and leaf-like; receptacles flat to shallowly convex, paleae sometimes with trilobed apices. 15 spp. endemics to Galápagos Islands in Ecuador.

365. Sclerocarpus Jacq. Erect annual or perennial herbs; leaves alternate, blades lanceolate to ovate, unlobed to deeply lobed; capitula solitary or in open paniculiform cymes, radiate; involucres campanulate to hemispherical, phyllaries 0 to few in 1–2 series, subequal; receptacles conical, paleae indurate and tightly wrapping cypselae (perigynia), appearing tubular, shed with the cypsela. 8 spp., 7 from SW U.S.A., Mexico, Central America, coasts of Colombia and Venezuela, one endemic to Colombia, some reposts also in Africa.

366. Simsia Pers. Erect annual or perennial herbs, shrubs; leaves opposite, auriculate, blades lanceolate, ovate to subcordate, sometimes deltate, trilobate, unlobed to shallowly lobed; capitula solitary or in open paniculiform cymes, radiate or discoid; involucres broadly cylindrical, turbinate, rarely hemispherical, phyllaries in 2–4 series, gradate; receptacles convex. 27 spp., New World, 5 in South America (slightly centered in Colombia), only the widely distributed S. dombeyana DC. in Brazil.

367. Syncretocarpus S.F. Blake. Erect annual or perennial herbs, shrubs; leaves alternate, blades narrowly lanceolate to ovate; capitula in open paniculiform cymes, radiate; involucres campanulate to hemispherical, phyllaries in 2–3 series, subequal; receptacles shallowly convex, paleae deciduous. Three spp., endemics to S & C Peru.

368. Tithonia Desf. Erect annual or perennial herbs, shrubs, rarely trees; leaves alternate, blades narrowly ovate to ovate, sometimes auriculate, bases attenuate; capitula solitary or in open paniculiform cymes, in most species on fistulose peduncles, radiate; involucres hemispherical, phyllaries in 2–5 series, subequal, sometimes outermost foliaceous; receptacles shallowly convex. 13 spp., SW U.S.A. to northern South America (only T. rotundifolia (Mill.) S.F. Blake, collected in Venezuela), one or 2 spp. introduced in most other tropical regions of the world.

369. Viguiera Kunth. (exc. Aldama p.p., Heiseria). 19 spp. from U.S.A. to Argentina, 8 in South America, all endemics in Venezuela (2), Peru (2), Paraguay (2) except from V. pazensis Rusby native from Peru to Chile (Tarapacá) and Argentina (Jujuy, Salta).

■ SUBTRIBE MONTANOINAE

370. Montanoa Cerv. in La Llave & Lex. Erect perennial herbs (?), shrubs, trees; leaves opposite, blades mostly ovate, entire or pinnatifid, triplinerved, rarely 5–7-plinerved; capitula terminal, mostly in paniculiform to corymbiform or thyrsoid cymes, radiate, rarely discoid; involucres mostly hemispherical, phyllaries 3–7 in 1–2 series, subequal; receptacles convex, paleae accrescent after anthesis and enfolding cypselae. 26 spp., most species in Mexico, Central America, northern South America (5) south to northern Peru, with two endemics to Venezuela.

■ SUBTRIBE SPILANTHINAE

371. Acmella Rich. ex Pers. Erect or decumbent annual or perennial herbs, sometimes rooting at the nodes; leaf-blades filiform to mostly ovate, triplinerved; capitula terminal or axillary, solitary or in open paniculiform cymes, discoid or radiate; involucres hemispherical, sometimes patent with age, phyllaries in 1–3 series, subequal; receptacles conic, paleate. c. 30 spp., pantropical, 29 species in the New World, 21 in South America, 12 in Brazil, 3 endemics.

372. Oxycarpha S.F. Blake. Stoloniferous, succulent, perennial herbs or weak low shrubs; leaf-blades filiform, succulent; capitula terminal, solitary, discoid; involucres campanulate, phyllaries in 3–5 series, gradate, coriaceous; receptacles conic, paleae coriaceous, apices aristate to acicular; corollas white, yellow. Only one sp., O. suaedifolia S.F. Blake, restricted from Venezuela and Colombia.

373. Salmea DC. Erect or scandent shrubs; leaf-blades lanceolate to ovate, coriaceous, triplinerved; capitula in open paniculiform cymes of variously congested corymbiform cymes, discoid; involucres campanulate to hemispherical, sometimes patent with age, phyllaries in 2–5 series, subequal or gradate; receptacles conical, paleate; corollas white or creamy white, sometimes purplish. 11 spp., Mexico, Central America, Caribbean, and only one in South America, S. scandens (L.) DC., from Venezuela, Guianas, Colombia, Ecuador, Peru, Bolivia, Brazil and Cono Sur.

374. Spilanthes Jacq. Perennial herbs, some prostrate and becoming woody at base; leaf-blades linear to elliptic, weakly triplinerved; capitula terminal, rarely axillary, solitary or in simple cymes, discoid; involucres hemispherical, phyllaries in 2–3 series, subequal; receptacles convex to conic, paleate; corollas white or purplish white; anthers black, appendages without glands. 6 spp., pantropical, 4 in New World from Mexico to Chile and Brazil, all in South America, two in Brazil, none endemic.

■ SUBTRIBE VERBESININAE

375. Verbesina L. Annual or perennial herbs, erect, rarely vine-like or straggly shrubs, or trees sometimes up to 25m tall; leaves alternate or opposite, sometimes with petiolar wings decurrent, sometimes forming dainty rosettes; capitula terminal, solitary, sometimes scapose, or in paniculiform or corymbiform cymes, discoid or radiate; involucres cylindrical, turbinate, campanulate, hemispheric. 338 spp., over New World, highest number of species in Mexico and tropical Andes, also in Chile, 141 in South America, only 10 in Brazil, 8 endemics.

■ SUBTRIBE ZINNIINAE

376. Heliopsis Pers. Erect to decumbent annual or perennial herbs, weak shrubs; leaves opposite, sometimes alternate distally, blades filiform to suborbicular, mostly ovate, pinnately veined or triplinerved, entire to deeply serrate or lobed; capitula terminal, solitary, sometimes on long peduncles or in open paniculiform cymes on sometimes fistulose peduncles, radiate or discoid. 15 spp., Central and northern South America with 3 spp. in Venezuela, Colombia, Ecuador, Peru, Bolivia, Argentina and Brazil (only H. buphthalmoides (Jacq.) Dunal, non endemic).

377. Sanvitalia Lam. Erect or prostrate annual or perennial herbs, rarely shrubs; leaves opposite, blades linear to ovate, mostly triplinerved, entire to shallowly serrate; capitula terminal, solitary or in open paniculiform cymes, radiate; involucres hemispherical, phyllaries in 2–4 series, subequal. 6 species, 5 in North America, Mexico, Central America, and S. versicolor Griseb. in Bolivia and Argentina.

378. Zinnia L. Erect annual or perennial herbs or shrubs; leaves opposite, linear to ovate, entire, rarely serrate; capitula terminal, solitary or in open paniculiform cymes on sometimes fistulose peduncles, radiate, rarely discoid; involucres turbinate, campanulate or hemispherical, phyllaries in 2–5 series, subequal to highly gradate; receptacles convex to conic, paleate, paleae with rounded, flat tips. 21 species, SW U.S.A., neotropical region, only two in South America (scattred from Central America to Andes), none in Brazil.

8.14 ASTEROIDEA ▸ TRIBE MILLERIEAE (32/c 430) - outsiders are Bebbia (2; SW U.S.A., NW Mexico), Dyscritothamnus (2; Mexico), Tetragonotheca (2; SE U.S.A.), Alepidocline (4; Mexico, Central America), Cuchumatanea (1; Guatemala), Faxonia (1; Baja California), Oteiza (4; Mexico, Guatemala), Selloa (6; Mexico, Central America), Guardiola (14; Mexico), Axiniphyllum (6; Mexico), Guizotia (6; tropical and subtropical Africa), Micractis (3; tropical Africa, Madagascar), Rumfordia (7; Mexico, Central America), Trigonospermum (6; Mexico).

379. Acanthospermum Schrank. Annual herbs; leaves petiolate to subsessile, blades elliptic, trullate or ovate, entire to lobed, triplinerved; capitula terminal or axillary, solitary. 6 spp., Central America to Cono Sur and Caribbean, 2 in Brazil, both widely distributed, adventive in other tropical regions of the world; 5 spp. in South America.

380. Alepidocline S.F. Blake. Annual herbs; leaves petiolate, blades lanceolate to broadly ovate, obscurely to strongly triplinerved; capitula terminal, solitary or in open paniculiform cymes, radiate. 5 spp. from Mexico and Central America, one also in Venezuela.

381. Alloispermum Willd. Perennial herbs or shrubs; leaves subsessile to petiolate, blades linear-lanceolate to ovate, triplinerved; capitula in terminal, simple, open to congested paniculiform or corymbiform cymes, radiate, rarely discoid. 15 species, Mexico and Central America to Peru and Venezuela, 7 in South America.

382. Aphanactis Wedd. (inc. Selloa) Annual or perennial herbs, variously persisting as caespitose, matted, cushion-like shrubs, creeping among forbs or small rosettes; leaves sessile, shallowly perfoliate, blades oval to ovate or oblong, glabrous to densely silvery-pilose, sometimes forming rosettes. 14 spp., one endemic to Mexico and 11 from Venezuela to Bolivia in South America.

383. Carramboa Cuatrec. Small trees with most leaves concentrated on apices of the branches; leaves alternate, ovate to pandurate, petiolate, very large, bright green, margins undulate, adaxial surfaces shallowly bullate, petioles and abaxial surface covered in a dense lanate, ferrugineous golden indumentum, leaf bases ampliated and forming a cupule around stem. 4 spp., endemics to Venezuela.

384. Desmanthodium Benth. Perennial herbs, shrubs or treelets; leaves opposite, sometimes perfoliate, blades ovate to lanceolate, triplinerved or pentanerved. 7 spp., 6 in Mexico and Central America, and D. blepharodon S.F. Blake endemic to Venezuela.

385. Espeletia Mutis ex Humb. & Bonpl. (inc. Coespeletia, Espeletiopsis, Libanothamnus, Paramiflos, Ruilopezia, Tamania) Acauli- or caulirosulate shrubs with marcescente leaves, sometimes tree-like with most leaves concentrated on apices of branches (mainly monocaulous), a few species monocarpic; leaves alternate, with very short internodes and ampliated bases, sometimes forming a large rosette, mostly with leaf bases wrapping around stem forming a cupule, blades lanceolate to fusiform, densely lanate on abaxial surfaces; capitula in terminal or axillary, dichasial or monochasial, corymbiform or paniculiform cymes, radiate or discoid.

140 spp., 54 in Venezuela radiation (48 entirely endemic to the Venezuelan Cordillera de Mérida, 2 broadly distributed in this range and nearby areas in Colombia and Venezuela, and 4 endemic to the northern section of the Colombian Cordillera Oriental and Sierra de Perijá) and 86 in Colombia radiation (74 endemic to the Colombian Cordillera Oriental (including four extending into the Venezuelan side of Páramo de Tamá), six are found in the Colombian Cordillera Central (four endemic to this range, one extending into Ecuador, and one shared with the Colombian Cordillera Occidental), three are endemic to the Colombian Cordillera Occidental, and two are endemic to Sierra de Perijá (including one yet to be found within Colombian borders).

386. Galinsoga Ruiz & Pav. Annual or perennial herbs, rarely shrubs; leaves petiolate or sessile, blades linear-lanceolate to ovate, triplinerved; capitula terminal, solitary or in open simple to paniculiform cymes, radiate, rarely discoid. 16 spp., neotropical, 2 species adventives in the Old World, 9 in South America, highly centered in Peru to Cono Sur, but two widely distributed in neotropics, both in Brazil.

387. Ichthyothere Mart. Perennial herbs or shrubs, aromatic; leaves petiolate or sessile, blades lanceolate to ovate, glabrous or pubescent, membranaceous to semisucculent, triplinerved; capitula in terminal, commonly congested simple, rarely open paniculiform cymes, disciform. 26 spp., Panamá, tropical South America (all species); 19 in Brazil, 13 endemics; four spp., from Goiás, Mato Grosso do Sul and Rondonia states, are considered rare species in Brazil by Plantas Raras do Brasil’s book.

388. Jaegeria Kunth. Annual or perennial herbs, some aquatic, sometimes stems fistulose and rooting at nodes; leaves opposite, sessile or petiolate, blades lanceolate to ovate, 3–5-nerved; capitula terminal or axillary, solitary or in paniculiform cymes, radiate. 10 spp., neotropical, most species in Mexico, three in South America – Ecuador and Colombia one endemic each, and the widely distributed J. hirta (Lag.) Less., present in Brazil.

389. Lecocarpus Decne. Shrubs; leaves petiolate, blades ovate to oblong in outline, deeply dissected to pectinate, semisucculent; capitula terminal, solitary. Three spp., endemics to Galápagos Islands.

390. Melampodium L. Annual or perennial herbs some developing a woody caudex, erect or prostrate; leaves usually petiolate, blades linear to ovate-rhombic to shallowly trullate, entire or lobed, uninerved or triplinerved, sometimes pinnately veined; capitula terminal, solitary or in simple, paniculiform or corymbiform cymes. 41 spp., neotropical, only two in South America, both widely distributed, both in Brazil.

391. Milleria L. Annual herbs; leaves perfoliate or auriculate, blades broadly lanceolate to suborbicular, sometimes trullate, triplinerved to pentanerved; capitula in terminal, open paniculiform cymes with distinctive morphology, composed of several branching dichasia which end in multiple, diverging cincinni subtended by a small bract, radiate. Three spp. from Mexico to Bolivia and Caribbean; two in South America.

392. Sabazia Cass. (inc. Freya) Perennial herbs, rarely weak shrubs; leaves petiolate or subsessile, blades lanceolate to ovate, sometimes margins deeply dentate, triplinerved; capitula terminal, solitary or in open paniculiform cymes, radiate. 15 spp., 12 from Mexico to Central America, two endemics to Colombia and one endemic to Venezuela.

393. Schistocarpha Less. Perennial herbs or subshrubs; leaves petiolate, petioles variously winged, blades ovate to broadly ovate, triplinerved, margins sometimes deeply serrate; capitula in terminal, congested paniculiform cymes, radiate or disciform; involucres campanulate to hemispheric, phyllaries in 3–4 series, subequal to shallowly gradate, scarious. 13 spp., neotropical, most species in Mexico and Central America; 2 spp. in South America, one of them restricted of northern Andes.

394. Sigesbeckia L. Annual or perennial herbs, sometimes forming a woody caudex with thickened roots and/or with fistulose stems; leaves with winged petioles, sometimes perfoliate, blades ovate to trullate, rarely broadly ovate, triplinerved; capitula in terminal, open paniculiform cymes, radiate.; involucres turbinate to campanulate, hemispheric, phyllaries dimorphic, outer lanceolate to shallowly oblanceolate, herbaceous, sometimes densely glandular, inner erect, cucullate. 10 spp., pantropical, Mexico to Cono Sur and Caribbean, absent in Brazil; 7 spp. in South America.

395. Smallanthus Mack. Annual or perennial herbs, shrubs or small trees, soemtimes with rhizophores; leaves petiolate, petioles variously winged, blades lanceolate to ovate, sometimes suborbicular or broadly deltate, entire to strongly dentate, triplinerved; capitula in terminal, corymbiform or open paniculiform cymes, radiate, sometimes nodding after anthesis. 21 spp., neotropical, 14 in South America, 5 in Brazil, 2 endemics; S. araucariophilus Mondin from Rio Grande do Sul state considered a rare species in Brazil by Plantas Raras do Brasil’s book.

396. Stachycephalum Sch. Bip. ex Benth. Perennial herbs or weak, straggly, short-lived shrubs; capitula in terminal, congested corymbiform or scorpioid cymes, radiate, sometimes the female floret somewhat separated from the male disc florets on a stipitate, lateral receptacle. Three spp., one in Mexico and two in Ecuador, one of then reaching into Argentina.

397. Tamananthus V.M. Badillo. Perennial herbs or shrubs; leaves alternate, sessile to shortly petiolate, bases expanded but not forming a cupule around stem, blades lanceolate to oval, sometimes narrowly obovate, with a single main vein; capitula in terminal, open paniculiform cymes, radiate. Only one sp., T. crinitus V.M. Badillo, endemic to Venezuela.

398. Tridax L. Annual or perennial herbs, shrubs; leaves opposite, blades linear to ovate, simple to deeply lobed, 1-veined or triplinerved; capitula terminal, solitary or in open paniculiform cymes, discoid, disciform or radiate. 35 spp., neotropical, 8 in South America, only two in Brazil, none endemics, one species a pantropical weed.

8.15 ASTEROIDEA ▸ TRIBE MADIEAE (34/205-220) - outsiders unvailable.

399. Amblyopappus Hook. & Arn. Annual herbs; leaves mostly alternate, blades linear to ovate in outline, pinnatifid or the distal entire; capitula congested-corymbose or congested-paniculate, radiate or disciform. Only one sp., A. pusillus Hook. & Arn., W U.S.A., W Mexico and Chile.

400. Lasthenia Cass. Annual or perennial herbs, sometimes semiaquatic; leaves opposite, blades linear to lanceolate, entire or toothed to pinnatifid, sometimes ciliate, glabrous to sparsely hairy; capitula laxly paniculate or solitary, radiate. 18 spp., 17 in W U.S.A., SW Canada, NW Mexico (Baja California), and L. kunthii (Less.) Hook. & Arn., in Chile and Argentina.

401. Madia Molina. Annuals, often aromatic; leaves proximally in rosettes or opposite (often crowded), distally alternate, blades lanceolate or oblong-linear to linear, usually entire, hirsute to strigose, often also glandular-pubescent; capitula corymbose, paniculate, racemose or spicate, usually radiate; peduncular bract tips each without a pit-gland, tack-gland or spine. 11 spp., U.S.A., Canada, and M. sativa Molina in Argentina, Chile and Hawaii, and M. chilensis Reiche only in Cono Sur.

8.16 ASTEROIDEA ▸ TRIBE PERITYLEAE (8/82–87) - outsiders Amauria (3; SW U.S.A., Mexico), Eutetras (2; Mexico), Pericome (3; SW U.S.A., Mexico).

402. Galeana La Llave & Lex. Leaves opposite, viscid, petiolate, blades deltate to ovate, triplinerved; capitula in terminal, open corymbiform cymes. Involucres ovoid to cylindric, phyllaries 5, biseriate; receptacles flat to shallowly convex; ray florets 3, fertile, corollas creamy white. Only one sp., G. pratensis Rydb., Mexico, Central America and N Colombia.

403. Lycapsus Phil. Perennial herbs or weak shrubs; leaves alternate, semisucculent, 1–2-pinnate, lobes linear; capitula terminal in open paniculiform cymes, radiate. Only one sp., L. tenuifolius Phil., San Felix and San Ambrosio Islands off the coast of northern Chile.

404. Perityle Benth. Annual or perennial herbs but mostly rupicolous shrubs, some with brittle branches; leaves opposite or alternate, petiolate, sometimes densely glandular and viscid, blades deltate to ovate, cordate or rarely linear to spathulate, entire to deeply dissected. 70 spp., 69 in W U.S.A., Mexico, Central America, one disjunct in Peru and Chile, and another endemic to Peru.

405. Unxia L. f. Annual or perennial herbs, shrubs; leaves opposite, blades elliptic to lanceolate, triplinerved, entire to shallowly serrate; capitula tightly clustered in terminal, simple cymes, radiate. Three spp., one widely distributed in this range, one in Brazil and Venezuela, and a third endemic to Brazil.

406. Villanova Lag; leaves opposite or alternate, viscid, petiolate, blades dissected, once pinnate, ovate in outline; capitula in terminal, open, simple or corymbiform cymes. 6 spp., one in Mexico and Central America, and 5 from Venezuela to Cono Sur in South America.

8.17 ASTEROIDEA ▸ TRIBE EUPATORIEAE (168/2.530–2.575) - outsiders unvailable; subtribes annuncied includes only the tribes with South American genera, without details. Herbs, shrubs, climbers, rarely trees; leaves usually opposite, sometimes whorled, sometimes alternate; capitula homogamous, discoid; phyllaries imbricate, 2-multi-seriate; receptacle usually naked; corollas regular; corolla-lobes short; flower colour purple, blue or white, never yellow; anther-bases obtuse; style arms obtuse, more or less club-shaped, often very conspicuous and long-exserted; achenes black with phytomelanin in walls; pappus usually of hairs.

■ SUBTRIBE CRITONIINAE

407. Amboroa Cabrera. Small erect subshrubs or shrubs; leaves opposite, lamina narrowly elliptical, remotely serrulate, sharply acute; inflorescences on slender peduncles, with solitary capitula or a pair of sessile capitula; florets 50–≥80; corollas white, narrowly funnelform, with long cylindrical basal tube and rather cylindrical throat, glabrous. Two spp., Bolivia, Peru one each.

408. Aristeguietia R.M. King & H. Rob. Erect to procumbent shrubs to small trees; leaves opposite; capitula few to many, corymbose, pedicellate; florets 13–100; corollas bluish, lavender, purple or pink, narrowly funnelform, inner and usually outer surfaces glabrous, some species with few hairs or few to many small glands on lobes. 21 spp., 20 in Colombia, Ecuador, Peru, and one endemic to Chile.

409. Asplundianthus R.M. King & H. Rob. Erect to scandent shrubs or trees; leaves opposite, lamina ovate to lanceolate, subserrate to serrate; inflorescence usually corymbose-paniculate, capitula sessile in glomerules; florets 6–10; corollas lilac, lavender or purple, narrowly funnelform, usually with glands on outer surface. 11 spp., Venezuela to Peru.

410. Austrocritonia R.M. King & H. Rob. Erect shrubs or small trees; leaves opposite, lamina ovate to elliptical, entire or remotely serrulate to closely serrate; inflorescence broadly corymbose; capitula sessile in clusters; florets 5 or c. 10; corollas white, narrowly funnelform, glabrous on inner and outer surfaces. 4 spp., endemics to Brazil, from Bahia to Paraná states.

411. Badilloa R.M. King & H. Rob. Erect shrubs, pubescent; leaves opposite, lamina oblong to lanceolate, serrate to remotely subserrulate; inflorescence corymbose; florets usually 4–10(–23); corollas white, lavender, pink or violet, narrowly funnelform, glabrous on inner surfasse and on outer surface of throat. 11 spp., Venezuela to Peru.

412. Bishovia R.M. King & H. Rob. Erect perennial herbs or subshrubs; leaves alternate above, opposite near base, lamina ovate to broadly ovate, serrate to sublobate, acute; inflorescence a diffuse somewhat leafy cyme; florets 30–60, fragrant; corollas lavender, narrowly funnelform, with cylindrical basal tub. Two spp., Argentina and Bolivia one each.

413. Castanedia R.M. King & H. Rob. Erect shrubs; leaves opposite, lamina elliptical or elliptical-oblong to slightly obovate, remotely serrulate, short-acute to obtuse; inflorescence a dense corymbose panicle; involucre cylindrical; Florets 6–7; corollaswhite, narrowly funnelform, sparsely glanduliferous on narrow tube and on throat, more glands on lobes. Only one sp., C. santamartensis R.M. King & H. Rob., endemic to Colombia.

414. Chacoa R.M. King & H. Rob. Erect flexuous shrubs; leaves opposite, lamina ovate to deltoid, serrate; inflorescences lax terminal pyramidal panicles; florets c. 20; corollas white, with slender basal tubes and narrowly campanulate limbs, glanduliferous on outer surface. Only one sp., C. pseudoprasiifolia (Hassl.) R.M. King & H. Rob., restricted from Argentina, S Brazil and Paraguay.

415. Corethamnium R.M. King & H. Rob. Erect shrubs; leaves opposite, lamina ovate to suborbicular, crenate-serrulate, short-obtuse to rounded; inflorescence of dense small corymbose panicles on leafy branches; florets c. 6; corollas white, with narrowly cylindrical tube and throat without external differentiation, with only lobes spreading, outer surface very sparsely glanduliferous, inner surfasse glabrous. Only one sp., C. chocoensis R.M. King & H. Rob., endemic to Colombia.

416. Critonia P. Browne. Coarse subshrubs to small trees or woody vines or climbers; leaves opposite, lamina elliptical to broadly ovate, entire to serrate; inflorescences usually thyrsoid-paniculate, with branches opposite and usually spreading at 90° angles; involucres usually cylindrical to fusiform; florets 4–12; corollas whitish, tubular or narrowly funnelform, glabrous outside or rarely a few glands on lobes. 36 spp., Central and South America (6) up to Cono Sur, Greater Antilles; two spp. in Brazil, none endemic.

417. Critoniella R.M. King & H. Rob. Erect herbs or shrubs; leaves opposite, lamina ovate to broadly ovate, serrulate to serrate, acute to acuminate; inflorescence a broadly corymbose to cymose panicle; capitula sessile on congested glomerulate branchlets; involucres narrowly cylindrical; Florets 6–25; corollas white, lavender, bluish or purple, narrowly funnelform, glabrous on inner surface and outside below lobes. 6 spp., Venezuela, Colombia, Ecuador and Peru.

418. Cronquistianthus R.M. King & H. Rob. Erect or flexuous shrubs; leaves opposite, lamina ovate or lanceolate to linear, entire to serrate; inflorescence a terminal corymbose panicle; florets 8–18, fragrant; corollas white, lavender or bluish, narrowly funnelform with broadly cylindrical basal tube, outer surface glabrous or with few glands or scattered hairs, inner usually glabrous, rarely with small hairs. 23 spp. from Colombia, Ecuador and Peru (high diversity).

419. Grosvenoria R.M. King & H. Rob. Erect shrubs or small trees; leaves opposite, lamina ovate to narrowly elliptical, entire to remotely serrate, sharply acute to short-acuminate; inflorescence broadly corymbose paniculate; florets 5–10, fragrant; corollas pink to white, narrowly funnelform, with tube glabrous, without hairs on inner surface; cells of throat oblong with sinuous lateral walls. 6 spp. from Colombia, Ecuador and Peru.

420. Hughesia R.M. King & H. Rob. Woody vines; leaves opposite, lamina ovate, subentire, remotely and minutely serrulate, shortly acute, minutely acuminate; inflorescences terminal on lateral branches, distinctly thyrsoidpaniculate with spreading branches; capitula sessile or subsessile in small clusters; florets c. 9; corollas purple in distal half when dry, narrowly funnelform with broadly cylindrical basal tubes, glabrous. Only one sp., H. reginae R.M. King & H. Rob., endemic to Peru.

421. Idiothamnus R.M. King & H. Rob. Erect spreading shrubs or small trees; leaves opposite, lamina elliptical to ovate, narrowly acuminate, serrate to remotely subserrulate; inflorescences terminal, corymbose; florets c. 12–20; corollas whitish to lavender, narrowly funnelform, with broadly cylindrical basal tube, glabrous on outer surface below lobes. 4 spp., one in Bolivia and Argentina, one endemic to Brazil, one in Peru, and one in Venezuela.

422. Imeria R.M. King & H. Rob. Erect slender shrubs or small trees; leaves opposite; inflorescences terminal, corymbose, capitula sessile in dense clusters; florets 8– 10; corollas rose-coloured or pink, narrowly funnelform, glabrous, lobes slightly longer than wide, smooth on inner surface, densely mamillose to papillose outside on the tips forming sclerified cap. Only one sp., I. memorabilis (Maguire & Wurd.) R.M. King & H. Rob., endemic to the Guiana Shield of Brazil, Venezuela, 600 – 1,200m elevation range.

423. Koanophyllon Arruda. Shrubs or small trees, rarely vines; leaves opposite, rarely alternate, lamina broadly lanceolate to elliptical, entire to serrate, rarely irregularly lobed; inflorescences pyramidally paniculate to corymbose; florets 5– 20; corollas whitish to greenish yellow, rarely violet, funnelformwith broadly cylindrical basal tube. 123 species, neotropics (68 endemics to Caribbean); 23 spp. in South America, 14 in Brazil, 10 endemics

424. Lorentzianthus R.M. King & H. Rob. Erect shrubs; leaves opposite, lamina ovate, often rather large, serrate, narrowly acuminate; inflorescence a pyramidal panicle; florets c. 10– 12; corollaswhitish to purple, narrowly funnelform with narrowly cylindrical basal tube, mostly glabrous outside below lobes, glabrous on inner surface. Only one sp., L. viscidus (Hook. & Arn.) R.M. King & H. Rob., restricted for Argentina and Bolivia.

425. Malmeanthus R.M. King & H. Rob. Erect shrubs; leaves opposite, lamina ovate, serrulate to subentire, scarcely acuminate; inflorescences terminal, corymbose-paniculate, with ascending mostly alternate branching; involucres campanulate; florets 5–22; corollas whitish, narrowly funnelform, with broad basal tube, glabrous. Three spp. from Brazil, one also in Argentina, Paraguay and Uruguay.

426. Neocabreria R.M. King & H. Rob. Erect subshrubs; leaves opposite, lamina narrowly elliptical, closely serrulate to crenate-serrulate; inflorescence a corymbose panicle; florets 6–25; corollas white to rose-purple, narrowly funnelform, glabrous on outer surface, with numerous hairs on inner. 5 spp. from Brazil, two up to Argentina and Paraguay.

427. Nothobaccharis R.M. King & H. Rob. Erect shrubs; leaves densely spirally inserted, lamina small, suborbicular to elliptical, dentate to crenate; inflorescence a dense thyrsoid panicle, with branches usually spiciform; capitula crowded; florets 6–8; corollas whitish, narrowly funnelform, glabrous on inner and most of outer surfaces. Only one sp., N. candolleana (Steud.) R.M. King & H. Rob., endemic to Peru.

428. Ophryosporus Meyen. Erect herbs or subshrubs, rarely scandent, sometimes with xylopodium; leaves usually opposite; inflorescence corymbose or thyrsoid, with corymbose branches; lateral capitula sometimes from axils of lower phyllaries of central capitula; florets 3–12; corollas white, with constricted basal tube and narrowly funnelform or campanulate limb, usually glanduliferous outside, glabrous or rarely puberulous inside. 41 spp., 37 in Argentina, Bolivia, Chile, Colombia, Ecuador, Peru, and 4 remaining are endemics to Brazil.

429. Santosia R.M. King & H. Rob. Woody vines; leaves opposite, lamina ovate to ovate-elliptical, subentire; inflorescences elongate, terminal, thyrsoid panicles; florets 1–10; corollas white, narrowly funnelform, with broadly cylindrical base, essentially glabrous on outer surface. Only one sp., S. talmonii R.M. King & H. Rob., endemic to Brazil, in Bahia state.

430. Sphaereupatorium (O. Hoffm.) Kuntze ex B.L. Rob. Erect perennial herbs or shrubs; leaves opposite; inflorescence laxly thyrsoidpaniculate, with branches and branchlets at right angles; capitula sessile in terminal globose clusters; florets c. 11; corollas white, narrowly funnelform, with broadly cylindrical basal tube, outer surface sparsely glandularpunctate, inner surface glabrous. Only one sp., S. scandens (Gardner) R.M. King & H. Rob., restricted from Bolivia and Brazil.

431. Steyermarkina R.M. King & H. Rob. Vines or flexuous shrubs; leaves opposite, lamina ovate, entire, obtuse to acute; inflorescence a lax thyrsoid panicle; florets 3–5; corollas white, narrowly funnelform, outer surface glabrous or with minute glands or large hairs on base of throat and lobes, inner densely pilose on throat. 4 spp., three endemics to Brazil (S. dispalata (Gardner) R.M.King & H.Rob. in Paraná, Minas Gerais, São Paulo, Rio de Janeiro; S. dusenii (Malme) R.M.King & H.Rob. in Paraná, Santa Catarina; and S. pyrifolia (DC.) R.M.King & H.Rob. in Bahia, Espírito Santo, Paraná, Santa Catarina, Minas Gerais, São Paulo, Rio de Janeiro) and one endemic to W Venezuela (S. triflora R.M.King & H.Rob., Trujillo)

432. Uleophytum Hieron. Woody vines; leaves opposite, lamina broadly oblong-ovate, minutely denticulate, acuminate; inflorescence of numerous capitula clustered in axils of leaves; florets c. 55–60; corollas whitish (?), narrowly funnelform, glabrous on inner and lower outside surfaces. Only one sp., U. scandens Hieron., endemic to Peru.

433. Tuberostylis Steetz. Creeping to scandent herbs or shrubs, full epiphytic; leaves opposite; inflorescences terminal on lateral branches or sessile in axils of leaves; capitula sessile in small panicles or axillary fascicles; florets c. 10–20; corollas white, narrowly tubular with slightly thickened base, glabrous on inner surfasse and lower outside surface. Two spp. from Panamá, Colombia (both, one endemic) and Ecuador.

■ SUBTRIBE GYPTIDINAE

434. Agrianthus Mart. ex DC. Erect many-branched shrubs; leaves usually densely spirally inserted, lamina elliptical to oblong-lanceolate or subulate; inflorescence terminal on branches, abrupt, a dense cluster of sessile or subsessile capitula; florets 20–45; corollas usually pink or white, rarely purplish, narrowly funnelform, with resin ducts along veins in throat. 9 spp., endmeics to Brazil.

435. Arrojadocharis Mattf. Annual or short-lived perennial herbs or shrubs; leaves spirally inserted, sessile; lamina linear; inflorescence of solitary capitula or lax corymbs terminating leafy branches; florets c. 50–60; corollas pink, funnelform, with small glands on outer surface. Two spp., endemics to Brazil.

436. Bahianthus R.M. King & H. Rob. Erect shrubs or small trees, viscid; leaves densely spirally inserted, with distinct narrow petioles, lamina obovate, remotely serrate, obtuse or truncate; inflorescence a corymbose panicle; capitula hemispherical; florets 15–22; corollas pink or white, narrowly funnelform to subcylindrical, tubes scarcely narrower than throat, resin ducts narrow and solitary along veins of throat. Only one sp., B. viscosus (Spreng.) R.M. King & H. Rob., in Bahia and Espírito Santo states, E Brazil.

437. Barrosoa R.M. King & H. Rob. Erect perennial herbs; leaves opposite, sometimes alternate above, lamina lanceolate to broadly ovate, serrate to crenulate; inflorescence densely corymbose; florets 20–55; corollas funnelform, white, pink, blue or purple; throat smooth. 12 spp., 3 in Venezuela and Colombia, remaining in Bolivia, Brazil (7, 5 endemics) and Cono Sur.

438. Bejaranoa R.M. King & H. Rob. Erect subshrubs or shrubs; leaves alternate or sometimes opposite, lamina ovate or ovatelanceolate, serrate to doubly serrate; inflorescence terminal with densely corymbose branches; florets 4–10; corollas white to pale lavender, narrowly funnelform, glabrous inside, outside with glands above. Two spp., B. balansae (Hieron.) R.M. King & H. Rob. from Bolivia and Paraguay, and B. semistriata (Sch. Bip. ex Baker) R.M. King & H. Rob. endemic to Brazil.

439. Bishopiella R.M. King & H. Rob. Acaulescent, scapose, annual or short-lived perennial herbs or subshrubs; leaves forming rosette, lamina fleshy, oblanceolate, entire, narrowly obtuse; inflorescence scapiform, a small, few-branched cyme; florets c. 40–50; corollas white, shortly funnelform from a broad, tapering, scarcely narrowed base, outer surfasse with a few short-stalked glands. Only one sp., B. elegans R.M. King & H. Rob., endemic to Bahia state, NE Brazil.

440. Campuloclinium DC. Erect coarse herbs or subshrubs; leaves opposite or alternate, lamina ovate to narrowly oblong; inflorescences corymbose, capitula few to many, moderate-sized or often large; florets 30–100; corollas pink, lavender or purple, narrowly funnelform, basal tube somewhat constricted above nectary. 12 spp. from Brazil, 7 endemics and 5 also in Argentina, Bolivia, Colombia, Guatemala, Honduras, Mexico and Paraguay.

441. Catolesia D.J.N. Hind. Multi branched shrubs; leaves densely spiralled, sessile, longest surrounding inflorescences and equalling or overtopping capitula, entire, slightly involute, apices acute, hooked upwards; inflorescence terminal, corymbose; involucre campanulate, phyllaries biseriate, distant, outer persistent, inner deciduous or easily falling; Corolla pink; tube glabrous, expanding gradually from base to lobes. Three spp., all endemic to rocky grasslands (campos rupestres) of Diamantina highs in center Bahia state, Brazil.

442. Conocliniopsis R.M. King &H. Rob. Erect subshrubs or shrubs; leaves opposite, few alternate above, lamina ovate, strongly crenate; inflorescence a dense corymbose panicle or cyme; Florets 20–30; corollas blue or lavender, narrowly funnelform, outer surface glanduliferous, inside glabrous. Only one sp., C. grossedentata (DC.) Hind, scattered, in Caribbean, Brazil, Colombia and Venezuela.

443. Dasycondylus R.M. King & H. Rob. Erect or spreading subshrubs or shrubs, sometimes subscandent; leaves opposite, lamina ovate to oblong, entire to serrate; inflorescence a corymbose panicle; florets 20–60; corollas white, narrowly funnelform. 8 spp., 7 endemics to Brazil and one restricted of Bolivia and Peru.

444. Diacranthera R.M. King & H. Rob. Perennial herbs or subshrubs; leaves opposite, lamina ovate to elliptical, crenulate to serrulate; inflorescence slightly to strongly cymose with many branches; florets 50–65; corollas pinkish, narrowly funnelform. Three spp., endemics to dry areas in Ceará to Bahia states in NE Brazil.

445. Gyptidium R.M. King & H. Rob. Erect annual herbs; leaves opposite, sometimes alternate above, lamina ovate to lanceolate, crenulate; inflorescence cymose to subcymose, capitula sessile to long-pedicellate; florets 50–80; corollas white or pale lilac, with very narrow basal tube; throat narrowly campanulate, smooth on both surfaces. Two spp., Argentina and Brazil one endemic each.

446. Gyptis (Cass.) Cass. Perennial herbs with sometimes with xylopodium, rhizophores or roots crown; leaves opposite, often becoming alternate above, lamina ovate to bipinnatifid, serrulate to deeply dissected; inflorescence subscapose, usually densely corymbose or cymose above. Phyllaries 2–3-seriate, 16–25, weakly subimbricate, subequal, with apical pubescence; receptacle flat, glabrous; florets 4–26; corollas narrowly funnelform, white, pink or violet. 6 spp., Bolivia, Argentina, Brazil (4, one endemic), Paraguay and Uruguay.

447. Lapidia Roque & S.C. Ferreira. Robust shrubs, weakly ramified at base and well branched in the apex; leaves only in the apex of the branches, opposite-decussate, somewhat fleshy, tomentose, glabrescent; capitulescence corymbiform with capitula cylindrical turbinate; cypselae 5-ribbed, carpopodium annuliform, pappus of bristles, setae fringed, distinct in size, purplish or white with purple apex. Only one sp., L. apicifolia Roque & S.C. Ferreira, endemic to rocky fields areas in Morro de Chapéu municipality, Diamantina highs, Bahia state, Brazil.

448. Lasiolaena R.M. King & H. Rob. Erect shrubs; leaves inserted in a dense spiral, short-petiolate, lamina narrowly to broadly obovate, serrulate above, obtuse or shortly acute; inflorescence densely corymbose on tips of leafy branches; florets 18–45; corollas narrowly funnelform, with scattered small glands on outer surface. 7 spp. endemics to Bahia state, NE Brazil.

449. Litothamnus R.M. King & H. Rob. Erect glabrous shrubs or small trees with subfleshy stems and leaves; leaves opposite, lamina elliptical to slightly obovate, entire; inflorescences corymbose, with opposite branches; florets (5–)15–25; corollas white, narrowly funnelform, sparsely glandular-puberulous on outer surface. Two spp., endemics to Bahia state, NE Brazil.

450. Lourteigia R.M. King & H. Rob. Small to medium-sized subshrubs or shrubs, sometimes procumbent; leaves opposite, lamina ovate to narrowly elliptical, crenulate to serrate, lower surface often white-tomentose; inflorescence terminal, densely corymbose; florets 20(–40); corollas lilac, blue, purple or greenish white, funnelform, tube narrowed above nectary. 10 spp. confined to Colombia and Venezuela.

451. Macropodina R.M. King & H. Rob. Erect subshrubs or shrubs; leaves usually opposite, becoming alternate above, lamina ovate, serrulate; inflorescence a lax cyme; florets (20–) 25–30(–50); corollas pale blue, narrowly funnelform, basal tubes elongate. Three spp. in Brazil, one up to Paraguay.

452. Morithamnus R.M. King, H. Rob. Erect shrubs or trees, with candelabra-like branching, viscid; leaves opposite or alternate, lamina obovate to oblanceolate, entire (very rarely crenulate), obtuse or acute; inflorescence abruptly terminal on branches; florets c. 25–100; corollas pink or white, narrowly funnelform; tubes broad, gradually broadened into cylindrical throat; resin ducts of throat paired along veins. Two spp., endemic to Bahia state in NE Brazil.

453. Neocuatrecasia R.M. King & H. Rob. Erect to procumbent perennial herbs; leaves usually opposite, lamina ovate to deltoid or narrowly oblong, entire or dentate to deeply lobed; inflorescence a lax to rather dense corymbose or cymose panicle; florets 17–50; corollas white, with narrow basal tube, with abruptly expanded usually elongate and campanulate throat; cells of limb oblong with sinuous lateral walls. 13 spp. confined to mountains of Bolivia and Peru.

454. Prolobus R.M. King & H. Rob. Erect shrubs; leaves mostly opposite, becoming alternate above, lamina ovate, coarsely serrate; inflorescence cymose, with ascending branches, often extra-axillary; florets c. 12–14; corollas pale violet, glanduliferous above on outer surface, tube broadly cylindrical; throat narrowly funnelform. Only one sp., P. nitidulus (Baker) R.M. King & H. Rob., endemic to Bahia state, NE Brazil.

455. Semiria D.J.N. Hind. Moderately branched small tree; leaves sessile, spirally inserted; inflorescence terminal, of few to several capitula; involucres campanulate; Florets c. 40, hermaphrodite; corollas pink, corolla tube cylindrical narrowing slightly towards base, glandular-punctate throughout and stipitate-glandular towards base. Only one sp., S. viscosa D.J.N. Hind, endemic to Bahia state, NE Brazil.

456. Stylotrichium Mattf. Erect shrubs; leaves densely spirally inserted, lamina narrowly obovate to orbicular, with prominent reticulate venation beneath; inflorescence abruptly corymbose to subumbellate on tips of leafy branches; florets c. 25–50; corollas white, short-funnelform, with stalked glands below lobes, outer surfaces of lobes glandular-punctate. 6 spp., endemic of Bahia state, Brazil.

457. Urolepis (DC.) R.M. King & H. Rob. Coarse, erect annual herbs or subshrubs; leaves opposite, lamina broadly deltoid, dentate or denticulate; inflorescence a corymbose or subcymose panicle; florets 100–150; corollas pink, with long narrow basal tube; throat funnelform below, becoming cylindrical above, with outer surface of tube and throat glabrous. Only one sp., U. hecatantha (DC.) R.M. King & H. Rob., Argentina, Bolivia, S Brazil, Paraguay and Uruguay.

458. Vittetia R.M. King & H. Rob. Erect shrubs or subshrubs; leaves opposite or alternate, subsessile, lamina orbicular to broadly ovate or oblong, entire to crenate-serrate; inflorescence an ascending multi-branched corymbose panicle; florets 10–12; corollas white to pink, narrowly funnelform or with narrow basal tube and rather campanulate limb. Two spp., from Minas Gerais to Santa Catarina states, SE Brazil.

■ SUBTRIBE OXYLOBINAE

459. Ageratina Spach. Perennial, usually erect herbs or shrubs; leaves usually opposite, lamina narrowly elliptical to deltoid, mostly toothed, lobed, serrate or crenate; capitula laxly to densely corymbose; florets 10–60, often sweetly scented; corollas white or lavender, usually with slender basal tube and campanulate limb in subgenera Ageratina and Klattiella, others narrowly funnelform. 307 spp., tropics and subtropics of the New World, one in Chile, 98 in South America, absent in Brazil. King and Robinson (1970) first provided an infrageneric division recognizing four subgenera, later raising subg. Pachythamnus to generic status.

460. Kaunia R.M. King & H. Rob. Erect shrubs or small trees; leaves opposite, lamina usually ovate, entire to serrate, broadly to narrowly acute; inflorescences terminal on branches, thyrsoid to corymbose with densely corymbose branches; florets (10–)16–50; corollas usually white or violet, narrowly funnelform, glabrous inside and on lower outer surface. 10 spp., 9 in Andes from Ecuador to NW Argentina, and K. rufescens (Lund ex DC.) R.M. King & H. Rob. in Bolivia, Paraguay, NE Argentina and C & S Brazil.

461. Jaramilloa R.M. King & H. Rob. Erect shrubs or small trees, to 3m tall; leaves opposite, lamina often large, broadly oblong, serrulate to scarcely undulate, apex short-acute; inflorescences terminal on branches, broadly corymbose, capitula short-pedicellate or sessile in glomerules; florets 14–20; corollas whitish, with narrow cylindrical basal tube and narrowly campanulate limb, glabrous on inner and lower outside surface. Two spp., endemics to Colombia.

462. Oxylobus (Moçino ex DC.) A. Gray. Decumbent herbs to low shrubs; leaves opposite, lamina small, ovate to oblong, crenate to subentire; capitula laxly to densely corymbose to subcymose; florets 20–75, slightly scented; corollas white or pink, with a long narrow basal tube and a narrowly campanulate limb. 6 spp., Mexico, Guatemala, O. glandulifer (Sch. Bip. ex Hemsl.) A. Gray up to Colombia and Venezuela.

■ SUBTRIBE MIKANIINAE

463. Mikania Willd. Usually woody vines, sometimes erect perennial herbs or shrubs, sometimes with xylopodium, moderately branched, never rosulate; leaves opposite or whorled, sessile to long-petiolate, lamina linear to broadly ovate, base narrow to cordate, membranous to coriaceous; inflorescence terminal on stems or lateral branches, cymose to corymbose or thyrsoid, capitula clustered, sessile to pedicellate, with subinvolucral bract; florets 4; corollas white or pink, funnelform or with variously campanulate limb. 443 spp., pantropical, 440 in New World, although principally neotropical with a few apparent natives in the Old World tropics; the report of dioecy in the genus is apparently restricted only to species from the Greater Antilles; 385 spp. in South America, 207 in Brazil, 143 endemics.

■ SUBTRIBE ADENOSTEMMATINAE

464. Adenostemma J.R. Forst. & J.G. Forst. Perennial herbs; leaf lamina narrowly elliptical to broadly ovate or hastate, crenate to strongly serrate, acute to slightly acuminate; inflorescence very laxly cymose; florets 10–60; corollas usuallywhite, narrowly funnelform or with narrow basal tube and broadly campanulate limb, usually with hairs or glands on outer surface, hairs often moniliform. Circa 26 spp., pantropical, 14 in New World, 12 in South America, 7 spp. in Brazil, two endemics.

465. Gymnocoronis DC. Annual to perennial erect herbs, submersed aquatic; leaf lamina lanceolate to ovate or deltoid; inflorescence strongly cymose. Phyllaries c. 20–50, biseriate, equal to subequal; receptacle with discrete oval scars and with soft tissue between; florets 50–200; corollas white, narrowly funnelform, with short-stalked glands on outer surface. Three spp., Guatema and Mexico one endemic each, and G. spilanthoides (D. Don ex Hook. & Arn.) DC. from Argentina, Bolivia, Brazil, Paraguay, Peru and Uruguay.

G. spilanthoides is becoming na invasive alien species in several regions of the world. The species is problematic in Australia, New Zealand, Japan and China (inc. Taiwan) and has recently naturalized in Italy. The species was reported from India (reputedly thesource of introduction to Australia through the aquarium trade) by Parsons & Cuthbertson (2001). There are relatively recent records of naturalization in Japan (in 1995), Taiwan in China (in 2001) and mainland China (in 2007). Kodono (2004) reports this species as rapidly naturalizing, occurring from Kyushu to central Japan. G. spilanthoides was reported as casual in 1988 in Hungary, occurring in the thermal waters of Lake Haeviz and ditches near. Ardenghi et al. (2016) report two naturalized occurrences in NW Italy (Lombardia region). The population in Italy stretches along the water body for 519 m, and occupies the whole canal width (1–4m)(Ardenghi et al., 2016). G. spilanthoides was ﬁrst reported as a naturalized species in Australia, reported from Taree in New South Wales (NSW) in 1980. It has since spread in NSW and also naturalized inthe states of Victoria and Queensland. It has been eradicated from an ornamental pond site in Perth and Margaret River in W Australia, the only known sites in thatstate. In New Zealand, G. spilanthoides was ﬁrst recorded as naturalized on the Papakura Stream in South Auckland in 1990 and has since been found through muchof lowland North Island and two South Island sites, the furthest south being in Canterbury.

466. Sciadocephala Mattf. Perennial herbs; leaf lamina narrowly ovate to elliptical or slightly obovate, entire to serrate; capitula solitary or laxly subcymose. Phyllaries c. 6– 14, persistent, 1–2-seriate, subequal to equal, separate to base; receptacle with discrete oval scars separated by soft tissue; florets c. 9–15; corollas white, narrowly funnelform, with sparse hairs on outer surface. 6 spp., 4 from Panamá to Peru, S. pakaraimae (Maguire & Wurdack) R. M. King & H. Rob. endemic to Guyana, and S. gracieliae N. Biggs & D. J. N. Hind, from dense forest of Amazon of N Mato Grosso state in Brazil.

Sciadocephala presently appears to be restricted to the relatively dense shade of forest floors or on mossy banks; this is an uncommon environment to find many Compositae. One other species preferring a similar heavily shaded environment is Cephalopappus sonchifolius Nees & Mart. (a rosulate stoloniferous herb with similar scapiform inflorescences and few capitula in the Nassauviinae) which is found in the dense shade of the Atlantic Forest of E Brazil (in the states of Bahia and Rio de Janeiro).

■ SUBTRIBE FLEISCHMANNIINAE

467. Fleischmannia Sch. Bip. Erect annual or perennial herbs or subshrubs; leaves usually opposite, lamina elliptical to rhomboidal or broadly cordate-ovate, upper part serrate or crenulate, or lamina dissected into long narrow segments; inflorescence with laxly cymose to densely corymbose branches; florets (10–) 20–50; corollas white, lavender, bluish or purple, with rather short basal tube. 100 spp., neotropical, 59 in South America, 7 in Brazil, 4 endemics.

■ SUBTRIBE AGERATINAE ▸ possibly biphyletic, with only Ageratum and Conoclinium (of Gyptidinae)

468. Acritopappus R.M. King & H. Rob. Shrubs or trees; leaves opposite, lamina ovate, ovate-elliptical, lanceolate or linear, glabrous or pubescent, often viscid, serrate to subserrulate or nearly entire, obtuse, short-acute to longacuminate; inflorescence on leafy side branches, branches densely subcymose to appearing subverticillate; florets 5–100; corollas usually pale lavender or pink, narrowly funnelform. 19 species, all endemics to Brazil.

469. Ageratum L. Annual to perennial herbs or subshrubs; leaves opposite or sometimes alternate, lamina elliptical or lanceolate to deltoid or ovate, entire to dentate. In florescence cymose to subcymose, sometimes subumbellate; florets 20–125; corollas white, blue or lavender, funnelform or with distinct basal tube. 36 spp., Central and South America (8), none in Chile; one species (A. houstonianum Mill.) widely cultivated, and another (A. conyzoides L.), although sometimes cultivated, is a widely distributed weed throughout the tropics in both the Old and New Worlds; 4 spp. in Brazil, 2 endemics.

470. Ascidiogyne Cuatrec. Prostrate, somewhat fleshy, stoloniferous herbs with erect rosulate branches; leaves of prostrate stems opposite, congested on erect stems, lamina ovate or obovate to narrowly elliptical, entire; inflorescence of clustered short 1-headed scapes; florets 5–7; corollas white, tube strongly constricted and pilose in upper part, limb broadly campanulate and glabrous. Two spp., endemics to Peru; the peculiar fluid-filled sac, formed from the outer wall of the achene, is present only in the type, and evident only in fresh material.

471. Cavalcantia R.M. King & H. Rob. Erect annual or short-lived perennial herbs. Lower leaves opposite, alternate above, lamina ovate to deltoid, distinctly shallowly lobed; capitula strongly divaricately cymose or aggregated in glomerules; florets c. 6–30; corollas white; basal tube short, broad below and constricted above, densely pubescent; limb funnelform. Two spp. endemics to Carajas massif in Pará state, N Brazil.

472. Centenaria P.Gonzáles, A.Cano & H.Rob. Small, erect, annual herbs, to 30 cm tall; leaves opposite, blade ovate to broadly elliptical, coarsely to finely serrate; inflorescence a diffuse corymbose cyme; phyllaries 5, distant, biseriate, subequal to equal, persistent, oblong-elliptical to obovate-elliptical, with shortly apiculate apices; receptacle flat, foveolate, glabrous, epaleaceous. Only one sp., C. rupacquiana P. Gonzáles, A. Cano & H. Rob. Known only from the type locality in Rupac, northeast from Lima Department.

473. Ellenbergia Cuatrec. Erect annual herb; leaves opposite, a few alternate above, lamina ovate to ovate-elliptical, crenate-dentate, acute; inflorescence a lax panicle with subcymose branches; florets 10–12; corollas white (?), with distinct constricted basal tube, glandular-punctate outside; throat broadly campanulate, papillate on inner surface. Only one sp., E. glandulata Cuatrec., endemic to Peru.

474. Ferreyrella S.F. Blake. Small, erect, annual herbs; leaves opposite below, alternate above, lamina ovate to broadly elliptical, coarsely to finely serrate; inflorescence a diffuse corymbose cyme; florets c. 30; corollas white, with short constricted basal tube bearing glandular or eglandular hairs; throat short and broad-campanulate. Two spp. endemics to Peru.

475. Gardnerina R.M. King & H. Rob. Annual or short-lived perennial herbs, erect from decumbent bases; leaves opposite below, alternate above, lamina ovate to rhomboid, repand-dentate to pinnatifid; inflorescence a few-headed cyme; florets 12–15; corollas white, funnelform, with stipitate glands outside in lower part, throat with short non-glandular hairs inside on and near bases of filaments. Only one sp., G. angustata (Gardner) R.M. King & H. Rob., endemic to Brazil, in Goiás state.

476. Guevaria R.M. King & H. Rob. Small perennial herbs, decumbent or erect with decumbent bases; leaves opposite or alternate, lamina ovate, crenulate to serrulate; inflorescence laxly paniculate with cymose branches; florets 15–40; corollas white, with distinct constricted basal tube bearing many hairs; throat shortly and broadly campanulate. 5 spp. from Ecuador and Peru.

477. Phalacraea DC. Perennial herbs, procumbent or erect from decumbent bases; leaves opposite, lamina ovate to broadly triangular, crenate to serrate; inflorescence rather laxly cymose, with denser ultimate branching; florets 10–18; corollas white, with constricted basal tube and abruptly expanding campanulate limb, many scattered minutely gland-tipped hairs on tube, more sparse on limb. 4 spp. from Colombia, Ecuador and Peru.

478. Piqueriella R.M. King & H. Rob. Small, annual or short-lived perennial herbs; leaves usually opposite, lamina ovate, with many large teeth, shortly and narrowly acuminate; inflorescence a lax cyme; florets c. 8; corollas whitish, with distinct constricted basal tube, glabrous on outer surface; throat broadly and shortly campanulate. Only one sp., P. brasiliensis R.M. King & H. Rob., Brazil, endemic to Ceará state.

479. Radlkoferotoma Kuntze. Shrubs or small trees; leaves opposite, lamina ovate to lanceolate, serrate; inflorescence corymbose; florets c. 35–70; corollas white or rosaceous, funnelform, with only minute glands on outer surface. Three spp. from S Brazil, two up to Uruguay.

480. Scherya R.M. King & H. Rob. Erect perennial herbs; leaves opposite, lamina linear, entire; inflorescence terminal, subscapose, cymose to subcymose; florets c. 25; corolla pale, funnelform, with glands on outer surface. Only one sp., S. bahiensis R.M. King & H. Rob., endemic to Bahia state, Brazil.

481. Stevia Cav. Mostly erect, annual or perennial herbs or shrubs; leaves opposite or in some species alternate, lamina linear to orbicular, entire to serrate or dentate, rarely deeply lobed; inflorescence diffuse or dense corymbose clusters on tips of branches; involucres cylindrical, rarely funnelform, narrow at base; florets 5; corollas white or lavender to purple, basal and distal parts sometimes of different colour, narrowly funnelform below lobes or with somewhat expanded throat, usually with hairs or glands on outer surface, with erect hairs on inner surface of throat. 259 spp., from U.S.A. and Mexico south to Central (excluding the Caribbean) and South America (152) up to Chile (2), 33 in Brazil (25 endemics); a difficult genus with an inadequate infrageneric division. Robinson’s original treatments cover much of the distribution of the genus, with the exception of the Brazilian species; a number of more recent regional treatments have been supplemented by the addition of many new species; a modern revision is certainly needed to adequately document the problems of hybridization and apomixis in the genus. S. rabaudiana (Bertoni) Bertoni from SW Brazil and E Paraguay is possibly the most sweet plant worldwide. Six species from Minas Gerais, Santa Catarina, Paraná and Rio de Janeiro states are considered rare species in Brazil by Plantas Raras do Brasil’s book.

482. Teixeiranthus R.M. King & H. Rob. Erect or decumbent, annual or short-lived perennial herbs; leaves opposite, lamina elliptical to linear; inflorescence a corymbose cyme; florets c. 30; corollas pale reddish, cylindrical with extreme base campanulate, base fused directly to top of achene, sometimes with poorly developed abscission zone. Two spp. in Bahia and Minas Gerais states, Brazil.

■ SUBTRIBE EUPATORIINAE ▸ as currently circumscribed, Eupatorium is composed of 48 species of annual or perennial herbs from E North America, the Caribbean in Cuba, S and E Asia (8 in Philippines), with one species (E. cannabinum L.) reaching North Africa and Europe.

483. Austroeupatorium R.M. King & H. Rob. Erect herbs or subshrubs; leaves opposite below, often subopposite or alternate above, lamina ovate to narrowly oblong, usually crenulate to serrulate; capitula in a flattened corymbose panicle; florets 9–23, fragrant; corollas white, rarely lilac, narrowly funnelform with rather narrow tube, glands on outer surface. 13 spp. in Brazil (11, 3 endemics), Bolivia and Cono Sur, except by two reaching in Caribbean and Central America, Caribbean, one species adventive in palaeotropics.

484. Eupatorium L. 60 spp., 25 from Algeria and Norway to Sakhalin and Vietnam, 28 from Canadá to Mexico and Cuba, and 7 spp. in South America, in Paraguay (E. areniscophilum Cabrera and E. gnaphalioides Cabrera endemics), Brazil (E. hagelundii Matzenb., E. lineatum Sch.Bip. ex Baker, E. rosengurttii Cabrera and E. semiamplexifolium G.S.S.Almeida & Carv.-Okano endemics); E. maracayuense Chodat occur in both countries.

485. Hatschbachiella R.M. King & H. Rob. Erect herbs or subshrubs; leaves opposite or alternate, lamina elliptical to narrowly elliptical, entire to remotely serrulate; inflorescence diffuse, with corymbose panicles at tips of leafy branches; florets 10–12; corollas white, narrowly funnelform with a narrow basal tube, glanduliferous on outer surface. Two spp. in Brazil, one of them up to Argentina, Paraguay and Uruguay.

486. Stomatanthes R.M. King & H. Rob. Perennial herbs or subshrubs; leaves alternate, opposite or ternate, lamina elliptical or oblanceolate to ovate or orbicular, entire to markedly dentate; inflorescence usually pyramidal to thyrsoidparticulate, sometimes corymbose; florets 4–11; corollas white, funnelform or nearly tubular, glabrous or glanduliferous with few to many hairs outside. 16 spp., 3 in E, C and S Africa, and 13 in Brazil, 4 of them up to Bolivia, Paraguay and Uruguay.

The results of nuclear ITS sequencing of Eupatorium s.l. by Schmidt and Schilling (2000) suggest that ‘Stomatanthes appeared to be more closely related to Praxelinae (Chromolaena) than it is to any member of the Liatrinae–Eupatorium clade’. This is not supported, however, by its morphology. It is retained here in the Eupatoriinae.

■ SUBTRIBE PRAXELINAE

487. Chromolaena DC. Erect to somewhat scandent perennial herbs, subshrubs or shrubs, sometimes xylopodial; leaves usually opposite, lamina mostly ovate or triangular to elliptical, sometimes linear, subentire to lobed; capitula usually thyrsoid to candelabriform or on laxly to densely corymbose branches, rarely solitary on long erect peduncles; florets 6–75; corollas white, blue, lavender or purple, rather cylindrical with scarcely narrower base, outer surface smooth below lobes, with few to many short-stalked glands, often with rather stiff hairs. 153 spp., New World tropics and subtropics, one species a pantropical weed, 121 in South America, 64 in Brazil (46 endemics). King and Robinson (1987) noted two subgenera and provided characters in their generic description to separate species of subgenus Osmiella.

488. Eitenia R.M. King & H. Rob. Erect annual or short-lived perennial herbs; leaves opposite, lamina ovate, coarsely serrate to sublobate, acute; inflorescence a rather lax broadly cymose panicle; florets c. 40–50; corollas violet or white, narrowly funnelform, with long cylindrical throat. Two spp., endemics to Goiás state and Distrito Federal in C Brazil.

489. Eupatoriopsis Hieron. Erect annual herbs; leaves opposite, lamina ovate to rather elliptical, serrate, short-acute; inflorescence laxly cymose; florets c. 30; corollas lilac, short-funnelform, with very short basal tube, glabrous on outer surface belowlobes; corolla lilac. Only one sp., E. hoffmanniana Hieron., endemic to Brazil, in Minas Gerais state.

490. Lomatozona Baker. Erect perennial herbs or subshrubs, covered with minute stipitate glands; leaves opposite, lobed to deeply dissected, lower surface with very large sessile globular glands; inflorescence laxly cymose; florets 10–27; corollas white or bluish, narrowly funnelform, with ocasional glands on outer surface. 4 spp. endemic to Brazil, in Mato Grosso and Goiás states.

491. Praxeliopsis G.M. Barroso. Erect annual (or possibly short-lived perennial) herbs, essentially glabrous; leaves alternate, sessile, linear, entire; inflorescence laxly cymose, pedicels very long; florets c. 16; corollas lilac, hypocrateriform, with elongate narrowly funnelform basal tube, glabrous on both surfaces; throat very short and spreading. Only one sp., P. mattogrossensis G.M. Barroso, from Brazil (Mato Grosso state) and Bolivia.

492. Praxelis Cass. Erect to decumbent annual or perennial herbs or subshrubs; leaves opposite or whorled, lamina ovate to elliptical or filiform, subentire to sharply serrate; capitula solitary on long erect peduncles to laxly thyrsoid or rather densely corymbose, usually campanulate; florets 25–30; corollas white, blue or lavender, narrowly funnelform or with cylindrical throat and slightly narrower basal tube, outer surface mostly smooth, with a few glands. 18 spp., South America, one species adventive in Asia and Australia; 16 spp. in Brazil, 8 endemics.

■ SUBTRIBE DISYNAPHIINAE

493. Acanthostyles R.M. King & H. Rob. Erect shrubs; leaves opposite, lamina narrowly lobed to pinnately dissected; inflorescence a long pyramidal panicle; corollas narrowly funnelform, purple-lilac. Only one sp., A. buniifolius (Hook. & Arn.) R.M. King & H. Rob., Argentina, Bolivia, Rio Grande do Sul state in S Brazil, Paraguay and Uruguay.

494. Campovassouria R.M. King & H. Rob. Erect shrubs or subshrubs; leaves opposite to alternate, usually closely spaced, lamina narrowly lanceolate to narrowly oblong or linear, entire to serrulate; inflorescence densely corymbose- paniculate. Corollas narrowly funnelform, lavender to purple. Two spp. from S Brazil, C. cruciata (Vell.) R.M. King & H. Rob. also in Argentina, Bolivia, Paraguay and Uruguay.

495. Disynaphia Hook. & Arn. ex DC. Erect shrubs or subshrubs, sometimes with xylopodium; leaves alternate, usually densely spirally inserted, lamina linear to oblong or oblanceolate, entire tominutely serrulate; inflorescence corymbose-paniculate; corollas broadly tubular below, slightly broadening above, purple, pink or white. 14 spp. from Brazil, 6 of them also in Argentina, Paraguay and Uruguay.

496. Grazielia R.M. King & H. Rob. Coarse, mostly erect, herbs, subshrubs or shrubs; leaves opposite, lamina ovate to lanceolate, sometimes pinnately to bipinnately dissected into narrow segments, often serrate; inflorescence densely corymbose. Corollas with broadly tubular bases, slightly broadening above, white, rose, lilac or purple. 10 spp. from Brazil, two also in Argentina, Paraguay and Uruguay.

497. Raulinoreitzia R.M. King & H. Rob. Erect shrubs; leaves opposite, lamina elliptical to linear, serrulate; inflorescence a pyramidal, often pendulous, panicle; corolla white, narrowly funnelform. Three spp. in Brazil (all spp., one endemic), Bolivia, Peru, Argentina, Paraguay and Uruguay.

498. Symphyopappus Turcz. Erect shrubs or small trees, often viscid; leaves opposite, sometimes becoming alternate near inflorescence, lamina ovate-lanceolate, serrate; inflorescence rather densely corymbose; corollas with broadly tubular base, slightly broadening above, rarely narrowly funnelform throughout, white, rose or purple-pink. 14 spp., 13 endemics to Brazil (Bahia to Rio Grande do Sul state), and S. apurimacensis H. Rob. endemic to Peru.

■ SUBTRIBE AYAPANINAE

499. Alomiella R.M. King & H. Rob. Erect to decumbent perennial herbs; leaves opposite to subopposite, lamina broadly ovate, serrate, acute; inflorescence laxly cymose. Phyllaries 20–30, subimbricate, 3-seriate, unequal, elliptical to oblong, receptacle flat, glabrous; florets c. 40; corollas white, narrowly funnelform, glabrous below onboth surfaces, veins greatly thickened in tube and throat. Two spp., endemics to Mato Grosso state in C Brazil.

500. Ayapana Spach. Erect perennial herbs; leaves mostly opposite, very rarely spiralled, lamina narrowly ovate to elliptical, entire to serrulate; inflorescence laxly paniculate, with laxly or densely corymbose to subcymose branches; florets 5–40; corollas white or pink, narrowly funnelform to nearly tubular, glabrous on inner surface, with glands on outer surface of lobes. 17 spp., neotropics up to Argentina and Brazil (4, none endemic).

501. Ayapanopsis R.M. King & H. Rob. Erect perennial herbs or subshrubs; leaves opposite, lamina elliptical or ovate to deltoid, acute to acuminate, serrate to nearly entire; inflorescence a corymbose panicle with corymbose to weakly cymose branches; florets 35–150; corollas pink to violet, narrowly funnelform. 17 spp., over South America, only one in Brazil, A. oblongifolia (Gardner) R.M. King & H. Rob., endemic.

502. Condylidium R.M. King & H. Rob. Erect to decumbent perennial herbs or subshrubs; leaves opposite, lamina ovate to ovate-lanceolate, bluntly serrate to subentire, short-acuminate; inflorescence thyrsoid-paniculate, with laxly and divaricately cymose branches; florets 5–6; corollas white, with a short constricted basal tube, with abruptly and rather narrowly campanulate limb. Two spp., C. iresinoides (Kunth) R.M. King & H. Rob. in Central America, Caribbean, Trinidad & Tobago, Venezuela, Guyana, Colombia, Ecuador, Peru, Bolivia, and C. cuatrecasii R.M. King & H. Rob. endemic to Colombia.

503. Gongrostylus R.M. King & H. Rob. Slender epiphytic vines, sparingly branched; leaves opposite, lamina ovate, remotely serrate; inflorescences mostly axillary, corymbose with cymose branches; florets c. 20; corollas white, very narrowly funnelform, mostly glabrous with glands on outer surface of lobes. Two spp., Colombia (both, one endmeic), Costa Rica, Ecuador, Panamá.

504. Gymnocondylus R.M. King & H. Rob. Erect perennial herbs; leaves opposite, lamina ovate, crenulate, scarcely acuminate; inflorescence a laxly corymbose cyme. Phyllaries c. 50, distant, 2–3-seriate, unequal, narrowly lanceolate to linear; receptacle slightly convex, glabrous; florets 60–80; corollas white, narrowly funnelform, basal tube very narrow below. Only one sp., G. galeopsifolius (Gardner) R.M. King & H. Rob., endemic to center Brazil in Goiás, Distrito Federal, Minas Gerais states.

505. Heterocondylus R.M. King & H. Rob. Erect to subscandent perennial herbs or subshrubs. At least lower leaves opposite, upper alternate in some species, lamina ovate to narrowly oblong or panduriform, entire to serrate; inflorescence pyramidal to distinctly cymose; capitula large, sometimes nodding; florets 20–80; corollas white to pink or reddish-purple, narrowly funnelform; cells of limb elongate with mostly sinuous lateral walls. 8 spp., Central America, Venezuela, Guianas, Colombia, Brazil (7, 3 endemics), Ecuador, Peru, Bolivia, Cono Sur.

506. Isocarpha R. Br. Erect annual or perennial herbs; leaves opposite or alternate, lamina narrowly elliptical, entire to serrulate, narrowly acute; inflorescence a lax, sometimes leafy panicle; florets more than 100; corollas white to pink, usually narrowly funnelform with a distinct short basal tube, rarely cylindrical, glands on outer surfasse mostly on tube and lobes. 5 spp., southern U.S.A. to Bolivia and Brazil (3, one endemic); all species occor in South America.

507. Lepidesmia Klatt. Erect perennial herbs; leaves opposite, lamina rather fleshy, lanceolate to linear-lanceolate, entire to subentire, blunt; inflorescence terminal, with elongate lower internodes, ending in dense cymes; florets 3–7; corollas white, narrowly funnelform, glabrous below, with a few glands on lobes. Only one sp., L. squarrosa Klatt, restricted for Colombia and Venezuela.

508. Monogereion G.M. Barroso & R M. King. Erect, short-lived perennial herbs or subshrubs; leaves mostly alternate, basal leaves opposite; inflorescence diffuse, with capitula laxly cymosely disposed on ends of leafy branches; florets c. 25–30; corollas white, narrowly funnelform, with hairs near base of tube, near tips of lobes and on entire upper two-thirdsof inner surface of lobes. Only one sp., M. carajensis G.M. Barroso & R.M. King, Carajas massif in Pará state, N Brazil.

509. Parapiqueria R.M. King & H. Rob. Small, erect, annual or short-lived perennial herbs; leaves opposite, upper subopposite to alternate, lamina linear; inflorescence very diffuse with many capitula, branches rather thyrsoid paniculate; florets c. 12; corollas white, with a short glabrous basal tube, limb broadly campanulate, with a short throat. Only one sp., P. cavalcantei R.M. King & H. Rob., Carajas massif in Pará state, N Brazil.

510. Polyanthina R.M. King & H. Rob. Erect perennial herbs; leaves opposite, sometimes alternate above, serrate; inflorescence a lax thyrsoid or pyramidal panicle with denser cymose branches. Phyllaries c. 40–50, subimbricate, 2–3-seriate, lanceolate; receptacle slightly convex, shortly puberulous; florets c. 200–300; corollas white, very narrowly tubular, glabrous on inner and outer surface. Only one sp., P. nemorosa (Klatt) R.M. King & H. Rob., Costa Rica to Bolivia and Venezuela.

511. Siapaea Pruski. Herbs. Stems repent, usually rooting at nodes; leaves opposite, simple, lamina elliptic-lanceolate, serrate, acute; inflorescence terminal, cymose, few-headed; phyllaries 1–2-seriate, few, distant, subequal; receptacle convex to conical, glabrous, epaleaceous; florets several, hermaphrodite; corollas cream, corolla tube glabrous, corolla lobes papillose; anther cylinder included within throat. Only one sp., S. liesneri Pruski, endemic to the Guiana Shield of Venezuela, 200 – 300m elevation range.

■ SUBTRIBE ALOMIINAE▸ possibly monophyletic if Dissothrix removed.

512. Austrobrickellia R.M. King & H. Rob. Erect or spreading to arching subshrubs or shrubs; leaves opposite, lamina ovate, entire to sharply dentate; inflorescence a lax leafy thyrsoid panicle, branches densely corymbose at tips; florets 3–12; corollas greenish white to purple, tubular, sometimes with slight constrictions above and near base, glabrous on outer surfasse or with few minute glands on lobes. Three spp. from Brazil (2, one endemic), Bolvia and Argentina.

513. Brickellia Elliott. Erect annual or perennial herbs, subshrubs or shrubs; leaves opposite or alternate, lamina linear, lanceolate, ovate, deltoid or lobate; capitula usually clustered in leafy thyrsoid panicle, sometimes corymbose or cymose, rarely solitary and nodding on long peduncles; florets c. 4–100; corollas usually white to cream-coloured, sometimes purplish, tubular or rarely narrowly funnelform. 98 species, New World, highly centered in Mexico (86, 59 endemics), and only the widely distribuited B. diffusa (Vahl) A. Gray in South America, inc. Brazil

514. Condylopodium R.M. King & H. Rob. Erect to subscandent shrubs; leaves opposite, lamina broadly elliptical, entire to remotely serrulate; inflorescence broadly pyramidally paniculate; florets c. 10–12; corollas greenish white, minimally narrowly funnelform, with glands above on outer surface. Six spp. from Colombia, one up to Ecuador.

515. Crossothamnus R.M. King & H. Rob. Erect shrubs; leaves opposite to alternate, lamina ovate, serrulate to subserrulate; inflorescence thyrsoid-paniculate, with branches rather densely corymbose; florets c. 10; corollas white, slightly funnelform, slightly narrowed above, glanduliferous on outer surface. Three spp., two in Peru and one in Ecuador.

516. Dissothrix A. Gray. Erect annual herbs; leaves opposite, lamina ovatelanceolate, serrate; inflorescence a loose leafy thyrsoid panicle with cymose branches; florets 6–8; corollas whitish, tubular, somewhat constricted above, glands dense at tips of lobes, very sparse elsewhere on outer surface, veins distinctly thickened below. Only one sp., D. imbricata (Gardner) B.L. Rob., Brazil, endemic to Ceará state; known only from the type collections.

517. Goyazianthus R.M. King & H. Rob. Erect cinereo-puberulous and glandular-punctate subshrubs. Primary leaves usually alternate, branch leaves and bracts of inflorescence usually opposite, lamina narrowly oblong-oblanceolate, entire; inflorescence thyrsoid-paniculate, with subcymose branches; florets 4; corollas creamy white, tubular, narrower above, with numerous glands on outer surface. Only one sp., G. tetrastichus (B.L. Rob.) R.M. King & H. Rob., endemic to Brazil, in Goiás state and Distrito Federal.

518. Helogyne Nutt. Erect subshrubs or shrubs; leaves alternate, lamina small, elliptical to lanceolate, entire; inflorescence a dense rather pyramidal or thyrsoid panicle or sometimes diffuse and leafy; florets 5–18; corollas white, pink, purple, or according to some descriptions, yellow, tubular and somewhat constricted above or funnelform, with many minute glands on outer surface, at least on lobes. 8 spp. from Argentina, Bolivia, Chile and Peru.

519. Leptoclinium Benth. & Hook. f. Erect shrubs; leaves alternate, imbricate, lamina broadly lanceolate, entire; inflorescence of small terminal corymbose panicles; florets 5; corollas creamy white, tubular, glabrous. Only one sp., L. trichotomum (Gardner) Benth. ex Baker, Brazil, endemic to Goiás state.

520. Planaltoa Taub. Erect, densely hirtellous subshrubs or shrubs; leaves alternate, imbricate, minutely serrulate; inflorescence terminal, densely corymbose or thyrsoid; capitula sessile in small bracteolate clusters; florets 3–5; corollas pink, tubular or minimally funnelform, outer surface densely pubescent with eglandular hairs, stipitate glandular hairs, or short-stalked glands. Two spp. endemics to Goiás state in C Brazil.

521. Pseudobrickellia R.M. King & H. Rob. Small trees or erect, often somewhat fasciculated shrubs; leaves densely spirally inserted, lamina narrowly linear, glabrous; inflorescence terminal on leafy branches, densely corymbose to somewhat pyramidal; florets 2–4(–8?); corollas greenish white, tubular or minimally funnelform, glabrous on outer surface. Two spp. endemics to Brazil.

■ SUBTRIBE HEBECLIINAE

522. Bartlettina R.M. King & H. Rob. Erect shrubs or small trees; leaves opposite, lamina lanceolate to broadly ovate; inflorescence usually corymbose-paniculate; involucres broadly campanulate; florets 8–150; corollas white, lavender, blue or purple, inner surface glabrous. 39 spp., tropical Central and South America (8, all single country range), one of them, B. hemisphaerica (DC.) R.M. King & H. Rob., endemic to Rio de Janeiro state in SE Brazil.

523. Guayania R.M. King & H. Rob. Erect perennial herbs or shrubs; leaves opposite, lamina elliptical to broadly ovate, serrate; inflorescences strongly cymose, ultimate branchlets with clusters of sessile or subsessile capitula; involucres campanulate; Florets 5–25; corollas bluish-white, lavender or white, with cylindrical basal tube, outer surface glabrous below lobes, inner glabrous. 5 species, endemic to the Guiana Shield of Brazil (4, none endemic), Colombia, Guyana, Venezuela, 100 – 2,800m elevation range.

524. Hebeclinium DC. Large herbs or subshrubs; leaves opposite, lamina broadly ovate to deltoid or lanceolate, usually crenate or serrate-pinnate; inflorescence a lax cyme with widely spreading branches; involucres broadly campanulate; Florets (12–) 20–80; corollas white or pink, outer surfasse glabrous below, inner surface of throat in some species with numerous hairs. 29 spp., neotropics, 25 in South America, only one in Brazil, the widely distributed H. macrophyllum (L.) DC.

■ SUBTRIBE TRICHOGONIINAE ▸ only three genera, near endemic to Brazil.

525. Platypodanthera R.M. King & H. Rob. Erect or ascending, rarely procumbent, annual or perennial herbs or subshrubs; stems slightly hexagonal; leaves usually alternate, lamina ovate to lanceolate, serrate; inflorescence a lax cymose or subcorymbose panicle with elongatebasal internodes; florets 40–130; corollas pink or lavender, rarely white, narrowly funnelform, glabrous. Only one sp., P. melissifolia (DC.) R.M. King & H. Rob., endemic to NE Brazil in the states of Alagoas, Bahia, Paraíba and Pernambuco in the dry seasonal scrubland of NE Brazil (caatinga) and savannas of C Brazil (cerrado).

526. Trichogonia (DC.) Gardner. Erect perennial herbs or subshrubs, sometimes with xylopodium; leaves usually alternate, opposite in some species at least below, lamina linear to broadly cordate, usually crenulate to crenate; inflorescence a lax to dense cymose or corymbose panicle; florets 10–60; corollas pink, purple or white, narrowly funnelform, basal tube sometimes narrow and elongate, limb with dense pubescence on upper throat and lobes. 21 spp., 17 in Brazil (14 endemic), and Bolivia, Paraguay, the northern Andes of Colombia and Venezuela, and the lowlands of NE Venezuela.

527. Trichogoniopsis R.M. King & H. Rob. Erect perennial herbs or subshrubs; leaves alternate or opposite, lamina ovate, serrate; inflorescence a lax cymose or corymbose panicle; florets 25–50; corollas white, narrowly funnelform, outer surface with few or no hairs above. Three spp., in the Atlantic Forest and savannas of C Brazil (cerrado) in the states of Bahia, Ceará, Espírito Santo, Mato Grosso, Minas Gerais, Paraná, Rio de Janeiro and São Paulo states in Brazil.

■ SUBTRIBE NEOMIRANDEINAE

528. Neomirandea R.M. King & H. Rob. Large herbs or shrubs to small trees, epiphytic or humicolous; leaves opposite or whorled, lamina deltoid or aceriform to elliptical or oblong, often slightly fleshy, entire to coarsely lobed and dentate; inflorescence a broadly cymose or corymbose panicle, with clustered capitula; florets 2–28; corollas white to reddish purple, narrowly funnelform, with or without hairs inside throat. 26 spp., Mexico to Colombia and Ecuador, centered in Costa Rica and Panamá; 8 spp. in South America, absent in Brazil.

62. ESCALLONIALES

A SINGLE FAMILY, PRESENT IN SOUTH AMERICA.

ESCALLONIACEAE

§ FULLY CHLOROTROPHIC

Genera/species 8/c 125 Distribution mainly southern hemisphere. Habit Usually bisexual (rarely unisexual), usually evergreen trees and shrubs (Valdivia gayana is a small shrub; Tribeles australis is a low procumbent and creeping shrub; Eremosyne pectinata is an annual herb); three genera in New World.

SYSTEMATIC outsiders Eremosyne (1; SW W Australia), Anopterus (2; Australia), Polyosma (c 60; E Himalayas and S China to E Queensland, E New South Wales and New Caledonia), Forgesia (1; Réunion).

1. Escallonia Mutis ex L.f. Shrubs or trees, terrestrial, rarely epiphytic; stems often with exfoliating bark; stipules usually absent; leaves alternate, simple, margins entire or often with glandular teeth; inflorescences mostly terminal racemes or panicles, or axillary and of solitary flowers; flowers actinomophic, bisexual; calyx campanulate, 5-lobed; petals 5, free, imbricate, often spathulate; stamens 5, free, anthers dithecal, longitudinally dehiscent, basifixed; intrastaminal disc present, surrounding the base of the style; ovary inferior, 2-3-locular, placentation axile, ovules many, style 1, stigmas 1-3; fruits septicidal capsules; seeds many, small, striate. 45 spp., all in South America from Venezuela to S Argentina up to mountains areas in SE Brazil (9, 7 endemics, Espirito Santo southwards) and Costa Rica (2); two in Uruguay and one in Paraguay; only six in equatorial Andes; locally the wood of Escallonia is used for fuel, charcoal, and timber; cultivated throughout the world for ornamental purposes; it is often dominant in Andean forests at high elevations; the highest species diversity of Escallonia is found in Chile and Argentina, with 15 endemics in to region.

2. Rayenia Menegoz & A.E. Villarroel. Perennial creeping subshrub, usually forming cushions of variable size, sometimes with unclear limits between neighbouring individuals, 5–20 cm tall, 20–180 cm wide, 50–1050 cm long. Only one sp., R. malalcurensis Menegoz & A.E. Villarroel, endemic to the eastern Andean ranges in the locality of San Fabián de Alico, Punilla Province, Ñuble Region, Chile.

3. Tribeles Phil. Small, prostrate shrub with spirally-arranged leaves that have broad bases and three small teeth at the apex; the small, terminal flowers have a contorted corolla, extrorse anthers, pollen with interrupted muri, and a style with a three-lobed, subclavate stigma; the shiny seeds long remain attached to the columella of the capsule. Only one sp., T. australis Phil., temperate Chile and S Argentina.

4. Valdivia Gay ex J. Remy. Small subshrub with indehiscent dry fruits and fused styles. numerous seeds are small and have long, almost unbranched ridges, and are borne on deeply intruding, bilobed and pitted placentas. Only one sp., V. gayana J. Rémy, known only three known localities in the province of Valdivia, south-central Chile, where it grows in sheltered and calcareous habitats (caves) close to the coast.

63. DESFONTAINIALES

A SINGLE FAMILY, PRESENT IN SOUTH AMERICA.

COLUMELIACEAE

Genera/species 2/5 Distribution tropical America. Habit bisexual, evergreen shrubs or small trees. Bud scales absent. Use Medicinal plants, carpentry, dyeing substances. Two genera.

Columelliaceae is currently unassigned in the Asterids. It has previously been placed in various orders: Rosales (Cronquist 1981), Saxifragales (Takhatajan 1983), Hydrangeales (Thorne 1992) and Dipsacales (Backlund 1996); its closest relative genera, Collumelia and Desfontainia are synonymised within the Columelliaceae due to similarities in wood anatomy, distribution, habit, large, showy flowers and the many seeds.

SYSTEMATIC both genera occur in South America.

1. Columellia Ruiz & Pav. Evergreen shrubs or trees; leaves simple, opposite, decussate; inflorescences few-flowered, terminal and/or axillary cymes or panicles or flowers solitary; flowers slightly zygomorphic, (4-)5(-8)-merous; corolla yellow, sympetalous, the lobes imbricate in bud; fruit septicidal, imperfectly 4-locular capsule with persistent calyx; seeds many, minute, laterally compressed, oblong, smooth. 4 spp., the Andes from S Colombia to Bolivia (all in Peru, one endemic), from about 1600 to 3600 metres above sea level.

Columellia have an epigynous flowers, zygomorphic corolla, 2 stamens and capsular fruit; Gelsemiaceae with leaf blade margins do not have spine like teeth; Fagraea Thunb. and Potalia Aubl.in in Gentianaceae have capsular fruit, an inferior ovary and no spine-like teeth around leaf blade.

2. Desfontainia Ruiz & Pav. Erect or sprawling evergreen shrubs, or small trees, 1-4m tall, with stiff branchlets, sometimes decumbent; leaves simple, opposite; flowers hermaphroditic, mainly solitary but also aggregated in inflorescences, terminal and in leaf axils, pentamerous; fruit ovoidal to globose, white berries (green when mature); seeds many, dark brown, shiny. Only one sp., D. spinosa Ruiz & Pavon, from Andes from Costa Rica (in 2000-4000 m) to Cape Horn in Chile (0-120 m), and Venezuela; Desfontainia is by some scientists considered to have only one very variable species D. spinosa Ruiz & Pav.; however at least three species have been described.

§ D. spinosa Ruiz & Pavon s.s. - corolla with acuminate lobes, 1.5-2(-3) times as long as calyx lobes, these are always pubescent on back and margin; Costa Rica to Tierra del Fuego.

§ D. fulgens D.Don - corolla gradually dilated towards the porrect or half-spreading limb, tube cylindrical, calyx lobes widely oblong, apex rounded, without prominent midrib; Chile and Argentina.

§ D. splendens Bonpl. - corolla abruptly dilated into half spreading or spreading limb, tube cylindrical, calyx lobes widely oblong or oblong acuminate, apex rounded to acute; Costa Rica to Bolivia.

64. BRUNIALES

BRUNIALES DOES NOT OCCUR IN SOUTH AMERICA, AND IS COMPOSED OF A SINGLE FAMILLY, BRUNIACEAE (6/81).

65. PARACRYPHYALES

PARACRYPHIALES DOES NOT OCCUR IN SOUTH AMERICA, AND IS COMPOSED OF A SINGLE FAMILLY, PARACRYPHIACEAE (3/35).

66. DIPSACALES

TWO FAMILIES, BOTH IN SOUTH AMERICA.

CAPRIFOLIACEAE

§ FULLY CHLOROTROPHIC, CUSHIONS

Genera/species 34/837–987 Distribution mainly temperate regions on the Northern Hemisphere; some species in subtropical regions, few species on tropical mountains, SE Australia and Tasmania, with their largest diversity in Himalayas and China Habit trees or herbs that can be recognized by their opposite leaves and often rather weakly monosymmetric flowers with a more or less radially symmetric calyx; the overy is inferior and the fruits are often few or one seeded. The bark in the woody taxa often comes off in thin flakes.

SYSTEMATIC Diervilloideae (1-2/16, E Asia, S.E. U.S.A.), Caprifolioideae (5/220, mostly N. temperate, esp. E Asia and E. North America), Heptacodium Clade (1/1, E. China), Linnaeoideae (6/32, Circumboreal, Mexico, China to Japan), Zabelia (1/4-6, Afganistan, Tian Shan to Japan), Morinoideae (2-3/13, Balkans to China) and Dipsacoideae (11/290, Eurasia, Africa, esp. Mediterranean region, to Malesia) not occur in South America; among Valerianoideae subfamily, outsiders are Patrinia (c 17; Ural and W Asia to Himalayas and E Asia); Nardostachys (1; Himalayas, W China); Centranthus (9; S Europe, Mediterranean), Valerianella (c 70; temperate regions on the Northern Hemisphere south to North Africa).

1. Valeriana L. (inc. Aretiastrum, Phyllactis, Stangea) Terrestrial annual, biennial, or perennial herbs sometimes subshrubs or cushion, or shrublets, or climbers or scandent plants, sometimes rosettes, lax and succulentd, usually with characteristic odour, especially when dry (associated with valeric acid), sometimes connate leaves and rather small flowers; the inflorescence is cymose and bracteolate; the sympetalous corolla is often spurred but other than that may be only weakly monosymmetric; the calyx often develops greatly in fruit. c. 270 spp., mainly in the Northern Hemisphere, 221 in South America, mainly restricted to the high altitude areas of the Andes, Peru (92, 55 endemics), Argentina (48, 15 endemics), Chile (44, 14 endemics), Ecuador (38, 8 endemics), Brazil (18, 15 endemics), Venezuela (2, none endemics), and SE South Africa. One or two weedy species found in the Neotropics.

Valeriana is the most morphological diverse genus in South America. Although its center of origin may be Asia, the present-day center of Valerianaceae species diversity is in South America, where many different morphological forms - from rosette plants (V. rigida Ruiz & Pav.) to microphyllous shrubs (V. microphylla Kunth) to annual vine-like species (V. chaerophylloides Sm.) - occupy a wide range of habitats; Valeriana are abundant and diverse in the Andes, especially in the paramo, scattered along the crests of the highest ranges in the northern Andes or on isolated mountaintops from 3,000 to 5,000 m; many of the high Andean species of Valeriana are growing as semirosulate or pulvinate plants in the Puna at altitudes of up to nearly 5,000 m; here, they have to survive recurrent night frosts of sometimes 1 to 4.5 ºC, with the risk of frost drought because the upper layers of soil often remain frozen for many hours; in the paramos of Colombia, Ecuador and northern Peru, associations of ligneous, shrubby or even more or less arborescent species occur; on the other hand, especially in Chile, a considerable number of annual species is found; some species, e.g. V. polystachya Sm. and V. salicariifolia Vahl, grow in swampy areas.

In Brazil almost all of them consisting of herbs restricted to the southern and southeastern states of the country and generally collected in montane or submontane habitats; only one spp. arborescent, a strikingly distinct, V. tajuvensis Sobral, endemic of a montane massif in the southernmost Brazilian state of Rio Grande do Sul and Santa Catarina states, occurs in E South America; arborescent species of Valeriana are quite common in the Andean highlands of Colombia, Peru, and Venezuela.

VIBURNACEAE

§ FULLY CHLOROTROPHIC

Genera/species 5/160–165 Distribution mainly temperate regions in the Northern Hemisphere; some species in subtropical regions, few species in tropical mountains, SE Australia and Tasmania, with their largest diversity in Himalaya and China. Habit usually bisexual (in Viburnum rarely polygamomonoecious), perennial herbs (Adoxa, Sinadoxa, Tetradoxa, some species of Sambucus), evergreen or deciduous trees or shrubs (Sambucus, Viburnum). Viburnum (c 150). Northern temperate, tropical mountains (absent from Africa). Usually shrubs (rarely trees).

Mostly northern temperate, with their highest diversity in China, tropical and subtropical mountains; they are characterised by opposite toothed leaves, small five- or, more rarely, four-petalled flowers in cymose inflorescences, and the fruit being a drupe. Some Cornaceae look similar to Viburnum, but the former often have 4-merous flowers; they lack stellate indumentum, and their lateral veins ascend towards the apex of the blade. Useful tips for generic identification: Sambucus has compound leaves, and Viburnum, simple leaves.

SYSTEMATIC two tribes, both in South America.

1. TRIBE ADOXEAE (3-4/29-30) ‣ outsiders Adoxa (2; temperate regions on the Northern Hemisphere south to Himalayas and C U.S.A.), Tetradoxa (1; China), Sinadoxa (1; China).

1. Sambucus L. Perennial herbs, shrubs or small trees up to ca. 7 m high, deciduous or semi-evergreen; young stems and branches with a prominent pith with tannin ducts, and in woody taxa with conspicuous lenticels; winter buds perulate; flowers mainly hermaphroditic, some dimorphic flowers or functionally unisexual individuals recorded. 10 spp. in temperate to subtropical regions and on tropical mountains, mainly of the northern hemisphere, with 4 spp. in South America: S. australis Cham. & Schltdl. in S Brazil and Cono Sur, S. mexicana C. Presl ex DC. in Ecuador and Peru (possibly a mistake), S. nigra L. disjunct in Colombia, also Mexico (possibly a mistake), and S. peruviana Kunth from Ecuador to Bolivia; two spp. in E Australia and Tasmania, S. ebulus ssp. africana (Engl.) Bolli in E Africa.

2. TRIBE VIBURNEAE (1/175) ‣ a single genus.

2. Viburnum L. Usually shrubs (rarely trees); hairs often stellate or peltate; foliaceous to scale-like appendages or nectariferous glands sometimes present on petiole or leaf base; nectary epigynous disc-like epithelial structure on ovary; flowers sometimes strongly fragrant. 150-175 spp., 78 in New World, 28 in South America, all species are restricted tropical Andes except V. tinoides L., which occur in Guiana Shield, in Venezuela, Guyana and Amazonas state in Brazil (only in Mount Aracá), also in Colombia and E Venezuela. 10 sections:

§ sect. Lentago ‣ 7 spp., E North America, except V. elatum Benth. in Mexico.

§ sect. Megalotinus ‣ 18 spp., SE Asia, extending west to India and south to Indonesia.

§ sect. Odontotinus ‣ 37 spp., temperate Asia and E North America, except V. orientale Pall. in the Caucasus Mountains.

§ sect. Opulus ‣ 5 spp., circumboreal.

§ sect. Oreinotinus ‣ 38 spp., Mexico, Caribbean, Central and South America; includes all species of continent.

§ sect. Pseudotinus ‣ 4 spp., Asia, except V. lantanoides Michx. in E North America.

§ sect. Solenotinus ‣ 26 spp., Asia, extending west to India and south to Indonesia.

§ sect. Tinus ‣ 7 spp., Asia, except V. tinus L. in Europe.

§ sect. Tomentosa ‣ 2 spp., China, Japan.

§ sect. Viburnum ‣ 14 spp., Asia, except V. lantana L. in Europe.

67. APIALES

FAMILIES ABSENTS IN SOUTH AMERICA: MYODOCARPACEAE (2/15), PENNANTIACEAE (1/3), PITTOSPORACEAE (9/293) AND TORRICELLIACEAE (3/11).

GRISELINIACEAE

§ FULLY CHLOROTROPHIC

Genera/species 1/7 Distribution New Zealand, Stewart Island, SE Brazil, Paraguay, Chile. Habit dioecious, evergreen trees or shrubs (sometimes lianas or epiphytes). Only one genus.

Araliaceae is closest to this family but Griseliniaceae lack resin ducts, umbels, and stipules found in that family.

SYSTEMATIC a single genus.

1. Griselinia Forst. & Forst. Shrubs, rarely exceeding 2m in height, sometimes epiphytic in coniferous Araucaria and Podocarpus species; leaves alternate, simple, margins entire, dentate or spinose, apex acute, obtuse or minutely 3-fid; inflorescences axillary racemes or panicles; flowers unisexual and plants dioecious; petals 5 in staminate flowers, greenish yellow, very small, imbricate, free; petals absent in pistillate flowers; fruits drupaceous, blackish; seed 1. 7 spp., two in New Zealand and 5 in South America, smaller shrubs, 1–5 m tall: three endemics of Chile; G. racemosa (Phil.) Taub. from S Chile and adjacent Argentina; and G. ruscifolia (Gay) Ball. in S Brazil, Araucaria forest and Atlantic coastal forest (the only true Neotropical taxa G. ruscifolia (Gay) Ball var. itatiaiae ((Wawra) Taub.), also in loma vegetation (Chile).

ARALIACEAE

§ FULLY CHLOROTROPHIC

Genera/species 46/c. 2,000 Distribution mainly tropical and subtropical regions in the Southern and Northern Hemispheres; a few genera in temperate areas. Habit usually bisexual (sometimes monoecious, andromonoecious, gynomonoecious, polygamomonoecious, or dioecious), evergreen or deciduous trees, shrubs or lianas (sometimes perennial herbs or suffrutices).

Some species are aquatic or helophytes. Leaf scars large and distinct. Small shrubs to large trees, less commonly lianas or herbs, glabrous or pubescent. Plants terrestrial, hemi-epiphytic, or climbing. 415 spp. in South America. The woody plants are often sparsely branched with pachycaulous stems; the leaves are often clustered toward apex of branches, with petioles of rather unequal lengths (especially in Dendropanax), and bases sometimes sheathing. Three other genera of the family, Oplopanax (Torr. & A. Gray) Miq., Panax L. and Pseudopanax K. Koch, are native to the Americas, but all are found exclusively in temperate areas. Additional generic names have commonly been applied to some Neotropical taxa of Araliaceae, such as Gilibertia Ruiz & Pav. (under Dendropanax) and Sciadodendron Griseb. (under Aralia).

No characters are always present in Araliaceae. However, the presence of flowers usually grouped in umbels or capitula and nectariferous discs is useful for recognizing members of the family.

SYSTEMATIC two subfamilies, both in South America.

1. SUBFAMILY HYDROCOTYLOIDEAE (3–4/c 260) ‣ outsiders Trachymene (c 60; SE Asia, Malesia, New Guinea, Australia, New Caledonia, Fiji), Uldinia (1; S Australia), Neosciadium (1; W Australia).

1. Hydrocotyle L. Herbs; leaf blades simple (often peltate and orbicular) to palmately lobed or compound; membranaceous, the margins entire to crenate or dentate; inflorescences axillary, often simple umbellate but sometimes compound umbellate or forming continuous to interrupted spikes; flowers with 5 free, valvate petals; stamens 5; carpels 2, ovary inferior, styles free or basally connate; fruits dry schizocarps, usually 5-ribbed, flattened laterally. 177 spp. worldwide, 79 in New World, 71 in South America, 21 in Brazil (12 endemics), widely distributed in moist habitats, especially in the Andes.

2. SUBFAMILY ARALIOIDEAE (c 34/1,380–1,400) ‣ outsiders Harmsiopanax (3; Malesia to New Guinea), Osmoxylon (c 60; Taiwan in China, Malesia to islands in Pacific); Astrotricha (c 20; Australia); Cheirodendron (6; Hawaii; Marquesas Islands), Motherwellia (1; Queensland), Cephalaralia (1; Queensland, New South Wales); Cussonia (c 20; tropical and southern Africa, Madagascar, Mascarene Islands), Seemannaralia (1; South Africa); Polyscias (c 160; tropical regions in the Old World east to New Caledonia, with their largest diversity in Madagascar), Meryta (28–30; E Australia, New Caledonia, Vanuatu, New Zealand, Three Kings Islands and Hen and Chickens Islands off New Zealand, Norfolk Island, Pitcairn Island, Society Islands, Marquesas Islands and other islands in SW Pacific), Pseudopanax (11–12; Tasmania, New Zealand), Plerandra (33; New Guinea, Solomon Islands, Vanuatu, Fiji); Heteropanax (8; S Asia, China), Fatsia (3; Japan, Korean Peninsula, Taiwan in China), Oplopanax (3; Japan, NW North America), Metapanax (2; S China, N Vietnam), Macropanax (17; Himalayas, China, SE Asia, W Malesia), Kalopanax (1; Siberia, China, Korean Peninsula, Japan), Tetrapanax (1; S China inc. Taiwan), Heteropanax (9; South and SE Asia, China), Schefflera (13; New Zealand to Pacific), Astropanax (18–19; Africa, Madagascar, Seychelles), Neocussonia (31; Tanzania to S Africa, Madagascar), Sinopanax (1; Taiwan in China), Merrilliopanax (3; Himalayas, northern Burma, Yunnan), Hedera (12–15; Europe, Macaronesia, Mediterranean, NW Africa, central and southern Asia to Japan and Taiwan in China), Trevesia (7; India, SE Asia, Malesia), Brassaiopsis (40–45; Himalayas, China, Thailand, Indochina, W and C Malesia), Eleutherococcus (c 40; Himalayas, SE Siberia, China, Korean Peninsula, Japan, SE Asia, Malesia to Philippines), Anakasia (1; New Guinea), Woodburnia (1; Burma).

2. Aralia L. Armed or unarmed, terrestrial, mostly deciduous shrubs or trees; leaves stipulate, base adnate to clasping; blades 1-4-pinnately compound; leaflets with margin entire to variously toothed; inflorescence terminal or lateral; paniculate, corymbose, or compound umbellate, ultimate units umbellules or sometimes capitula or racemules; pedicels articulated (rarely unarticulated); flowers with 5--10(--12) free, imbricate petals; fruits berry -like drupes. 70 spp. from E and SE Asia, Malesia, North America, 14 spp. in New World, up to Argentina, most species in seasonally dry forests, usually below 500 m; as recently circumscribed, Aralia has six sections:

§ sect. Aralia ‣ three North American and 11 Asian species.

§ sect. Dimorphanthus (Miq.) Miq. ‣ two E North American and 27 Asian species.

§ sect. Humiles Harms ‣ three North and Central American species.

§ sect. Nanae Harms ‣ one North American species.

§ sect. Pentapanax (Seem.) J. Wen ‣ 19 Asian species.

§ sect. Sciadodendron (Griseb.) J. Wen ‣ 5 spp., one in Cuba, four in South America, A. bahiana J. Wen endemic to Bahia state, Brazil; A. warmingiana (Marchal) J. Wen in Brazil, Bolivia and Cono Sur, A. excelsa (Griseb.) J.Wen from North America to Venezuela and Guianas, and A.a soratensis Marchal in Peru, Bolivia and Cono Sur.

3. Cephalopanax G.M. Plunkett, Lowry & D.A.Neill (inc. Schefflera p.p.). Hermaphroditic, unarmed, terrestrial, ever-green trees, often with ferruginous orreddish-brown (less commonly gray)indument, especially on young parts; stems monocaulous to moderately branched, pachycaulous. Two described species and an estimated five to ten new species that remain to be de-scribed, from W Venezuela to Pasco region in Peru.

4. Crepinella Marchal. Trees, leaves with coriaceous leaflets and small stipular ligules, compoundumbellate inflorescences, and 2- to 5-carpellate ovaries. 32 spp., largely restricted to montane vegetation on the sandstone tepuis of the Guiana Shield (30 in Venezuela, 18 endemics), with just three species occurring on sandstone substrates elsewhere in N Brazil (11, none endemics), Colombia, Ecuador, and Peru.

5. Dendropanax Decne. & Planch. Glabrous plants with simple or palmatilobed leaves, which are often 3-nerved at the base and have schizogenous glands and small intrapetiolar stipules; the inflorescences are simple or more or less compoundly arranged umbels of unjointed, pedicellate 5-9-merous flowers with valvate petals, glandular and versatile anthers, and fruits that sometimes have a distinct stylar column. 92 spp. from tropical and subtropical Asia and Neotropics, 69 in New World, 41 in South America, 22 in Brazil (16 endemics, 4 of then in E country are rare plants, by Plantas Raras do Brasil’s book), up to Argentina, moist forests at lowland to mid-elevation areas, usually below 1,500 m.

Most of the Brazilian species of Dendropanax are endemic to the Atlantic rain forests of the eastern coast, where more than 10 species occur, some of which are, as yet, undescribed. D. bahiensis Fiaschi, together with D. exilis (Toledo) Jung and other as yet undescribed species from Espírito Santo state, these species form a group characterized by a shrubby habit, leaves with schizogenous dots visible on the abaxial surface, reduced inflorescences (simple or branched), and small and shortpedicellate greenish flowers; this group is endemic to the interior of the Atlantic rain forests of E Brazil, ranging from Bahia to Santa Catarina states.

6. Didymopanax Decne. & Planch. 37 spp., 28 endemics to Brazil (11 of then are rares, all in SE region but one in Mato Grosso, another in Amazonas state, by Plantas Raras do Brasil’s book), six in Brazil and adjacent counties, two endemics to Venezuela, and D. morototoni (Aubl.) Decne. & Planch. from Mexico and the Caribbean to southern Brazil and NE Argentina.

Most species of the genus are endemic in mid- to high-elevation areas of the Atlantic Forest (12), the rocky grasslands (campos rupestres), highlands of savannas of C Brazil (cerrado, 7), and the northwestern Amazon rainforests in the border areas among Brazil, Venezuela, and Colombia (5); about two-thirds of the species of genus are forest-dwellers, whereas the remaining occur mostly in savannas; the genus is poorly represented in seasonally dry tropical forests, where only a single widely distributed species is found. D. confusus (Marchal) Fiaschi & G.M.Plunkett (Colombia, Peru, N Brazil) apparently has the largest fruits and the largest number of leaflets for any New World Araliaceae.

7. Oreopanax Decne. & Planch. Pubescent or glabrous, terrestrial or epiphytic shrubs or trees; leaves with blade simple or palmately lobed to compound, highly variable in shape, the margins entire or variously toothed; inflorescences terminal, paniculate, the ultimate units capitula; bracts well-developed; pedicels lacking; flowers with 5--7 free, valvate petals; stamens 5--7; carpels 2-12; ovary inferior, styles free or connate basally; fruits drupes. 141 spp. over mountainous areas of Neotropics, 114 in South America, especially above 1,500 m in the Andes; two spp. in E Brazil, O. fulvus (Jacq.) Decne. & Planch. endemic and O. capitatus Marchal, ranging from S Mexico to Brazil.

8. Raukaua Seem. Scandents shrubs, composed leaves. 12–20 spp., mainly in Tasmania and New Zealand, and two in South America, both in Chile and Argentina.

9. Sciodaphyllum P.Browne. 145 spp. (more than 200 remain to be described), 19 restricted of Costa Rica and Panamá, 4 only Caribbean, 2 in both South and Central America (or Caribbean), and remaining 120 only in South America, with high diversities in Colombia (58), Ecuador (33), Venezuela (25), Panamá (21), Peru (23) and Bolivia (12); wide range of humid or seasonally moist habitats, from sea level to nearly 3,500 m elevation. 6 spp. in Brazil, none endemics and shared with only Venezuela except S. sprucei Seem. shared with Peru, Ecuador and Colombia.

APIACEAE

§ FULLY CHLOROTROPHIC

Genera/species 447/3,460-3,580 Distribution cosmopolitan except polar areas. Habit usually bisexual (sometimes andromonoecious or polygamomonoecious, rarely dioecious), usually perennial, biennial or annual herbs (sometimes suffrutices or shrubs, rarely trees). Some representatives are xerophytic, aquatic or helophytic; internodes usually hollow.

Myrrhidendron is the most tree-like genus in the Neotropics, but species of Arracacia, Coaxana, Coulterophytum, Enantiophylla, Neonelsonia, Prionosciadium, Mathiasella and Dahliaphyllum also contain secondary (woody) tissue. Azorella species form densely tufted cushion plants. Conium maculatum L. is deadly poisonous and widely naturalized. 776 spp. in New World, 278 in South America; predominantly herbs with pinnately or palmately dissected leaves; inflorescences usually in umbels, less often in heads; flowers with 5 sepals, petals and stamens; inferior ovary with two carpels; fruits are schizocarps with 2 mericarps.

SYSTEMATIC eight high lineages: Platysace (Mackinlayoideae (Klotzschia (Azorelloideae (Hermas (Phlyctidocarpa + Saniculoideae) (Apioideae)))))); clades Platysace Megacore (1/c 25), Hermas clade (1/9, W and E Cape), Phlyctidocarpeae (1/1, Outjo and Kaokoveld in Namibia) does not occur in South America.

1. SUBFAMILY MACKINLAYOIDEAE (10/c 95) ‣ outsiders Actinotus (c 20; Australia, New Zealand), Apiopetalum (2; New Caledonia), Mackinlaya (5; Central Malesia to islands in Pacific); Chlaenosciadium (1; W Australia), Brachyscias (1; W Australia), Xanthosia (c 25; Australia, with their highest diversity in Western Australia); Pentapeltis (1; W Australia), Schoenolaena (1; W Australia).

1. Centella L. c. 50 spp., southern Africa to Zimbabwe and Malawi, with C. asiatica (L.) Urb., pantropical, cited in New World from North America, Caribbean, Venezuela, Colombia, Peru, Bolivia, Brazil and Cono Sur; one variety of this spp. is often treated as a independent spp., C. erecta (L. f.) Fernald., from North America to N South America and Antilles.

2. Micropleura Lag. Herbs. Two spp., one in Sierra Nevada de Santa Marta, Colombia and another from Mexico to Panamá.

2. KLOTZSCHIA CLADE (1/3) ‣ a single genus.

3. Klotzschia Cham. Herbs with peltate leaves, acaulis to tall, sometimes with taproot tubers; fruit without wings or ‘pseudo-wings’; ribs without distinct oil ducts. Single fused ventral bundle replacing carpophore. Three spp., K. brasiliensis Cham. and K. rhizophylla Urb. from Minas Gerais and Bahia states, very small acaulis herbs from dry mountains, and K. glaziovii Urb., a single erect few branched shrub, endemic to Goiás state.

3. SUBFAMILY AZORELLOIDEAE (16/113 - 118) ‣ four clades, all in South America.

3.1 AZORELLOIDEAE ▸ DIPOSIS CLADE - a single genus.

4. Diposis DC. Geophyte herbs. Three spp. in southern Chile and Argentina up to Uruguay.

3.2 AZORELLOIDEAE ▸ BOLAX CLADE ‣ outsiders Dichosciadium (1; Australia), Drusa (1; Canary Islands, Somalia).

5. Bolax Comm. ex Juss. Small cushions. Two spp. from temperate regions in Chile and Argentina.

6. Bowlesia Ruiz & Pav. Ephemeral annuals. 16 spp., all South America (only two up to North America and Mexico), mainly from Peru to Patagonia, and one, B. incana Ruiz & Pav. reaching into southern Brazil (also in Chile, Peru and Mexico).

7. Homalocarpus Hook. & Arn. 6 spp. endemics to Chile.

3.3 AZORELLOIDEAE ▸ ASTERISCIUM CLADE ‣ outsider Oschatzia (2; SE Australia, Tasmania)

8. Asteriscium Cham. & Schltdl. Succulent. 9 spp., Chile (4 endemics) and Argentina (4 endemics).

9. Domeykoa Phil. Ephemeral annuals. 5 spp., two in S Peru and three endemics to N Chile.

10. Eremocharis Phil. 9 spp. from Peru (8), Ecuador (1) and Chile (1).

11. Gymnophyton Clos. Leafless shrubs. 6 spp., the Andes in Bolivia, Chile and Argentina.

12. Pozoa Lag. Perennial short herbs, succulent; leaves ovate–orbicular to reniform, slightly or doubly dentate, usually with 13–30 triangular teeth; umbels has 25–45 flowers, some staminate; flowers are usually greenish-yellow; fruits are oblong–ovate, the mature carpels being strongly compressed. Two spp. widely distributeds at elevations between 1,000 and 4,000 m and distributed along the southern Andes in Chile and Argentina.

3.4 AZORELLOIDEAE ▸ AZORELLA CLADE ‣ outsiders Dickinsia (1; SW China), Diplaspis (3; SE Australia, Tasmania).

13. Azorella Lam. (inc. Huanuca, Mulinum, Schizeilema) Compiscuous, cushion or mat-like, membranaceous herbs to 2 cm to 2m high, some spinescent. 58 spp., New Zealand (14), Macquarie Island (2), Australia (1 endemic), and 42 in South America from Venezuela to Patagonia, highly centered in Argetina and Chile, A. biloba (Schltdl.) Wedd. up to Costa Rica, and A. selago Hook. f. up to Kerguelen, Heard Island and McDonald, Crozet, and Prince Edward Islands.

14. Spananthe Jacq. Herbs with small witish flowers. Only one sp., S. paniculata Jacq., Caribbean, Mexico to Venezuela, Peru, Bolivia, Ecuador and Brazil.

4. SUBFAMILY APIOIDEAE (395–435/2.850–3.010) ‣ 30 high and complex lineages, mostly African, temperate Asia or Australasian, only eight in New World, five of then up to South America.

APIOIDEAE ▸ SOUTH AMERICAN UNPLACED APIOIDEAE

15. Austropeucedanum Mathias & Constance. Only one sp., A. oreopansii (Griseb.) Mathias & Constance, endemic to NW Argentina.

16. Paraselinum H. Wolff. Only one sp., P. weberbaueri H. Wolff, from Peru (Ayacucho region and Cordillera de Raura, above 3,500 m altitudinal range, known only 2 collects) and Bolivia.

4.1 APIOIDEAE ▸ TRIBE OENANTHEAE (c 20/c 175) ‣ outsiders mainly North America from Canada to Mexico and Cuba (10) except Sium (9; the Northern Hemisphere, Africa), Berula (5; Northern Hemisphere, Africa, St. Helena), Cryptotaenia (5; Northern Hemisphere, E African mountains), Helosciadium (c 45; temperate regions in the Old World), Naufraga (1; Majorca), Cicuta (5; Northern Hemisphere), Oenanthe (c 28; Northern Hemisphere, tropical African mountains, India to Malesia and Australia), Trocdaris (1; SW Europe, Morocco).

17. Lilaeopsis Greene. 14-20 spp., 5-11 scattered in Mauritius, New Zealand and Australia, and 9 in New World, six in South America from Venezuela to Cono Sur, 4 spp. in Brazil (mainly up to Cono Sur, L. tenuis A.W. Hill endemic), also Argentina, Colombia, Chile; this species are morphologically similar and occur in aquatic sites of the Americas, New Zealand and Australia; simple umbel is extremely rare in Apioideae, appearing consistently only in Oreomyrrhis, Lilaeopsis, and Neogoezia Hemsl.

4.2 APIOIDEAE ▸ TRIBE SCANDICEAE (c 40/c 335) ‣ mainly in temperate Old World, but endemic genera in North America; three genera in in South America.

18. Daucus L. Annual, biannual or perennial herbs, erect or decumbent; leaves alternate, leaves strongly pinnatifid, umbels terminal or (mainly) axllary, flowers incompiscuos. 20 spp., almost worldwide, centrered in Mediterranean (mainly Algeria) except fro D. glochidiatus (Labill.) Fisch., C.A.Mey. & Ave-Lall. from Australia, New Zealand and Tasmania, and the two spp. of New World, D. montanus Humb. & Bonpl. ex Spreng. from Mexico to Argentina continuous in western coast, and D. pusillus Michx. from Chile to S Brazil, disjunct in North America.

19. Oreomyrrhis Endl. Herbs with inflorescence consisting of a simple umbel borne terminally on peduncles that arise from the bases of sheathing and often rosetted leaves. 23 spp., three in Taiwan in China, 1 in Borneu, 4 in New Guinea, 10 in Australia and New Zealand, two in Mexico, 1 in Guatemala, O. andicola (Kunth) Hook. f. from Costa Rica and Venezuela to Argentina, and O. hookeri Mathias & Constance in southern Chile and Argentina. Simple umbel is extremely rare in Apioideae, appearing consistently only in Oreomyrrhis, Lilaeopsis Greene, and Neogoezia Hemsl.

20. Osmorhiza Raf. Plants andromonoecious, slender to robust, perennial, herbaceous, aromatic, caulescent, dying back to a basal rosette of leaves; stems erect to spreading-ascending or decumbent, solitary to densely clustered, branching, fistulose, pubescent to glabrous. 10 spp., 4 only in North America, 1 only in Asia and 5 in South America: three occur in North America disjunct Chile/Argentina, O. mexicana Griseb. occur from Mexico to Costa Rica, isolated in Colombia, S Peru and N Bolivia, and from N Argentina; and O. glabrata Phil. only in Chile and Argentina.

4.3 APIOIDEAE ▸ TRIBE APIEAE (12–13/34–38) ‣ cosmopolitan, with their highest diversity in warm-temperate regions in Africa to SW Asia; one genus in South America.

21. Apium L. Herbs. 12 spp., amphipolar disjunction pattern; A. graveolens L. occur in Europe, W Asia or N and E Africa up to Kazakhstan, Oman and Lybia; two are endemics to S Australia, A. prostratum Labill. ex Vent. from Australia, New Zealand, Namibia, South Africa, NE Argentina, S Brazil and Uruguay, and remaining 8 restricteds for Cono Sur except A. sellowianum H.Wolff up to Bolivia and S Brazil.

4.4 APIOIDEAE ▸ TRIBE PYRAMIDOPTEREAE (c 30/175–180) ‣ temperate to subtropical regions on the Northern Hemisphere and one genus (and a single) in Argentina, with the largest diversity in Mediterranean to Central Asia.

22. Cyclospermum Lag. Three spp., C. leptophyllum (Pers.) Sprague. widely from U.S.A. to Argentina, Brazil and Caribbean, and two remaining scattered from S Ecuador to Chile and Uruguay.

23. Notiosciadium Speg. Only one sp., N. pampicola Speg., native from E Argentina to Uruguay.

4.5 APIOIDEAE ▸ TRIBE SELINEAE (c 65/c 730–785) ‣ very diverse in Old World; Arracacia clade is distributed in Mexico, Central and South America; perennial endemic North American clade is centered in the Rocky Mountains.

► SOUTH AMERICAN NON ARRACACIA CLADE ‣ three genera, all in South America.

24. Ammoselinum Torr. & A. Gray. Herbs, annual, odorless or "faintly Pastinaca-scented" in some species; leaves all alternate; basal 3-ternately compound, cauline 2–3-ternately compound; umbels compound, loosely convex, axillary, pedunculate or sessile (rays appearing to arise from leaf axils), peripheral flowers not different; flowers bisexual; petals white, margins entire; schizocarps ovoid-oblong to urceolate-ovoid or broadly ellipsoid. Three spp., A. rosengurtii Mathias & Constance from Uruguay and Rio Grande do Sul state in S Brazil (Santana do Livramento and Quaraí municipalities), often reported by Argentina, and two in North America.

25. Oligocladus Chodat & Wilczek. Only one sp., O. patagonicus (Speg.) Pérez-Mor., endemic to Argentina.

26. Spermolepis Raf. Annual herbs, slender taprooted, glabrous, not aromatic or sometimes (S. lateriflora) with a "carrot" odor; stems erect, 5–80 cm, simple or few-branched from basal to medial nodes; leaves all alternate; flowers bisexual; petals white, oblong or elliptic to ovate, apex not inflexed, margins entire; schizocarps broadly ovoid to ellipsoid or elliptic-ovoid, sometimes slightly beaked. 11 spp., one in Hawaii, S. castellanosii Pérez-Mor endemic to Rio Negro, Neuquen, Mendoza in Argentina, and remainign nine native to North America and Mexico.

27. Tauschia Sclecht. 34 spp., 31 in North America and Mexico, two in Guatemala, and T. nudicaulis Schltdl. disjunt in Mexico and Ecuador.

► ARRACACIA CLADE ‣ outsiders Coaxana (2; Mexico), Mathiasella (1; Mexico), Prionosciadium (11; Mexico), Dahliaphyllum (1; Mexico), Coulterophytum (4; Mexico), Enantiophylla (1; Central America), Rhodosciadium (9; Mexico).

28. Arracacia Bancr. Perennial herbs with taproots or tubers. 35 spp., 11 spp. in South America, from Venezuela to Bolivia, 6 in Peru and Colombia, some locally endemics. A. xanthorrhiza Bancr. is only native umbellifer domesticated in South America, is widely distributed through Bolivia, Peru and Ecuador from 2,000 to 4,000 m, cultivated for its tuberous storage roots, which are prepared in a way similar to potatoes and estimated to be a major food staple for approximately 80–100 million people.

29. Cotopaxia Mathias & Constance. Two spp., C. whitei Constance & W.S. Alverson endemic to small area in Sierra Nevada de Santa Marta, Colombia, and C. asplundii Mathias & Constance endemic to Ecuador.

30. Donnellsmithia J.M.Coult. & Rose. 20 spp., Mexico to Panamá with two up to Colombia and Venezuela.

31. Myrrhidendron J.M.Coult. & Rose. Shrubs to small trees. 5 spp., three in Central America and two in Colombia (one endemic) and Ecuador.

32. Neonelsonia J.M.Coult. & Rose. Only one sp., N. acuminate (Benth.) J.M.Coult. & Rose, from Mexico, Central and northern South America to Venezuela and Peru.

33. Niphogeton Schltdl. 18 spp., high mountains herbs, all in northern Andes, from Costa Rica (only two) to Bolivia and Venezuela.

34. Ottoa Kunth. Only one sp., high mountain O. oenanthoides Kunth, from Mexico to Ecuador and Venezuela.

35. Perissocoeleum Mathias & Constance. 4 spp., one in Andes from Colombia and adjacent Venezuela, and remaining three endemic to Sierra Nevada de Santa Marta in Colombia (above 3260 m high range).

5. SUBFAMILY SANICULOIDEAE (9/330–350) ‣ two tribes, Steganotaenieae (2/3, Ethiopia to South Africa) do not occur in South America; among Saniculeae, outsiders are Alepidea (c 40; tropical Africa), Actinolema (2; Mediterranean), Arctopus (3; Cape), Astrantia (11; Central and South Europe, W Asia), Petagnaea (1; Sicily).

36. Eryngium L. Prostrate herbs annual or perennial, few woody, only a few centimeters to erect and up to 3 m tall; leaves may have long petiolated leaves or sessile ones, with entire to partite blades, entire, setose or spiny margins, and first order venation either pinnate, palmate or even parallel-veinedcapitate inflorescences and single bract per flower. c. 250 spp., temperate and tropical worldwide, except Africa, 169 in New World; 97 in South America, and 59 spp. in Brazil (34 endemics); two clades:

§ subg. Eryngium ‣ ‘Old World’ clade automatically established as. It includes E. maritimum L., the type species of the genus, and all species from Africa, Europe, and Asia, except four western Mediteranean species.

§ subg. Monocotyloidea ‣ ‘New World’ clade, characterized by the possession of sessile, generally linear, parallelveined leaves and a well-developed cauline axis (erect and with several internodes); includes all species from New World plus some species from Mediterranean region. E. pandanifolium Cham. & Schltdl., from S Brazil, NE Argentina, Paraguay and Uruguay, is the tallest Apiaceae from South America, up to 4m tall.

37. Sanicula L. Perennial herbs. 41 spp., almost cosmopolitan, absent from New Guinea and Australia, 21 in New World; 4 sections:

§ sect. Pseudopetagnia ‣ 6 spp., Asia.

§ sect. Sanicoria ‣ 18 spp. W North America, 3 in Hawaii.

§ sect. Sanicula ‣ 13 spp., cosmopolitan, only one in Asia, includind the three South American spp., S. crassicaulis Poepp. ex DC. and S. graveolens Poepp. ex DC. disjuncts from North America and Cono Sur, and S. liberta Cham. & Schltdl. from Mexico to Bolivia and Venezuela.

§ sect. Tuberculatae ‣ 3 spp., China, Korea and Japan.

ANGIOSPERMS: TABLES

Here is a list of all the APG families and orders in 2023, with the number of Brazilian, unbrazilian and Brazilian endemic genera in each one, as part of project Synopsis of South American Angiosperms.

WHITE LINES: families/orders that occur native in Brazil.

ORCHID PINK LINES: exxofamilies/exxorders.

GRAY LINES: groups totally absent in South America.

BY FAMILY

FAMILY	BRAZILIAN GENERA	UNBRAZILIAN GENERA	BRAZILIAN ENDEMIC GENERA
AMBORELLACEAE
HYDATELLACEAE
CABOMBACEAE	1	1
NYMPHAEACEAE	2
AUSTROBAILEYACEAE
TRIMENIACEAE
SCHISANDRACEAE
CANELLACEAE	1	1
WINTERACEAE	1
SAURURACEAE
PIPERACEAE	3
ARISTOLOCHIACEAE	2	1
MYRISTICACEAE	5
MAGNOLIACEAE	1
DEGENERIACEAE
HIMANTANDRACEAE
EUPOMATIACEAE
ANNONACEAE	29	3	3
CALYCANTHACEAE
SIPARUNACEAE	1
GOMORTEGACEAE		1
ATHEROSPERMATACEAE		2
HERNANDIACEAE	2	1
MONIMIACEAE	5	1	3
LAURACEAE	25	3	3
CHLORANTHACEAE	1
ARACEAE	38	8	7
TOFIELDIACEAE	1
ALISMATACEAE	5
BUTOMACEAE
HYDROCHARITACEAE	5	1
SCHEUCHZERIACEAE
MAUNDIACEAE
APONOGETONACEAE
JUNCAGINACEAE	1	1
ZOSTERACEAE		1
POTAMOGETONACEAE	3
POSIDONIACEAE
RUPPIACEAE	1
CYMODOCEACEAE	1	1
ACORACEAE
PETROSAVIACEAE
NARTHECIACEAE	1
BURMANNIACEAE	7
THISMIACEAE	1	1
AFROTHISMIACEAE
TACCACEAE	1
DIOSCOREACEAE	1
TRIURIDACEAE	5	1
VELLOZIACEAE	2	1
STEMONACEAE
CYCLANTHACEAE	8	3
PANDANACEAE
CAMPYNEMATACEAE
MELANTHIACEAE		1
PETERMANNIACEAE
ALSTROEMERIACEAE	2	1
COLCHICACEAE
PHILESIACEAE		2
RIPOGONACEAE
SMILACACEAE	1
CORSIACEAE		1
LILIACEAE
ORCHIDACEAE	200	95	24
BORYACEAE
BLANDFORDIACEAE
ASTELIACEAE		1
LANARIACEAE
HYPOXIDACEAE	2
TECOPHILAEACEAE		3
DORYANTHACEAE
IXIOLIRIACEAE
IRIDACEAE	20	8	4
XERONEMATACEAE
ASPHODELACEAE	1	1
AMARYLLIDACEAE	11	27	3
ASPARAGACEAE	6	7
ARECACEAE	36	16	1
DASYPOGONACEAE
HANGUANACEAE
COMMELINACEAE	14	2	1
PHILYDRACEAE
PONTEDERIACEAE	2
HAEMODORACEAE	2	1
TYPHACEAE	1	1
BROMELIACEAE	55	13	26
RAPATEACEAE	9	7
XYRIDACEAE	4	1
ERIOCAULACEAE	15		3
MAYACACEAE	1
THURNIACEAE	1
JUNCACEAE	2	5
CYPERACEAE	32	8
RESTIONACEAE		2
FLAGELLARIACEAE
JOINVILLEACEAE
ECDEIOCOLEACEAE
POACEAE	174	60	23
STRELITZIACEAE	1
LOWIACEAE
HELICONIACEAE	1
MUSACEAE
CANNACEAE	1
MARANTACEAE	10	1
COSTACEAE	3	1
ZINGIBERACEAE	1
CERATOPHYLLACEAE	1
EUPTELEACEAE
PAPAVERACEAE		2
CIRCAEASTERACEAE
LARDIZABALACEAE		2
MENISPERMACEAE	16		1
BERBERIDACEAE	1
RANUNCULACEAE	3	8
SABIACEAE	2
NELUMBONACEAE		1
PLATANACEAE
PROTEACEAE	3	5
TROCHODENDRACEAE
BUXACEAE	1	1
MYROTHAMNACEAE
GUNNERACEAE	1
DILLENIACEAE	6
PERIDISCACEAE	1	1
PAEONIACEAE
ALTINGIACEAE
HAMAMELIDACEAE		1
CERCIDIPHYLLACEAE
DAPHNIPHYLLACEAE
ITEACEAE
GROSSULARIACEAE		1
SAXIFRAGACEAE		5
CRASSULACEAE	1	1
APHANOPETALACEAE
TETRACARPAEACEAE
PENTHORACEAE
HALORAGACEAE	3	1
CYNOMORIACEAE
VITACEAE	2	1
KRAMERIACEAE	1
ZYGOPHYLLEACEAE	2	9
QUILLAJACEAE	1
FABACEAE	217	56	23
SURIANACEAE	1
POLYGALACEAE	11	4	1
ROSACEAE	6	5
BARBEYACEAE
DIRACHMACEAE
ELAEAGNACEAE
RHAMNACEAE	13	9	2
ULMACEAE	2
CANNABACEAE	2	1
MORACEAE	17	1
URTICACEAE	13	1
NOTHOFAGACEAE		3
FAGACEAE		2
MYRICACEAE		1
JUGLANDACEAE		3
CASUARINACEAE
TICODENDRACEAE
BETULACEAE		1
APODANTHACEAE	2
ANISOPHYLLEACEAE	2		1
CORYNOCARPACEAE
CORIARIACEAE		1
CUCURBITACEAE	19	7	2
TETRAMELACEAE
DATISCACEAE
BEGONIACEAE	1
LEPIDOBOTRYACEAE	1
CELASTRACEAE	22	4	2
HUACEAE
CONNARACEAE	5		1
OXALIDACEAE	2
CUNONIACEAE	2	2
ELAEOCARPACEAE	2	2
CEPHALOTACEAE
BRUNELLIACEAE		1
PANDACEAE
RHIZOPHORACEAE	4
ERYTHROXYLACEAE	1
PERACEAE	3
RAFFLESIACEAE
EUPHORBIACEAE	65	10	4
CENTROPLACACEAE
CTENOLOPHONACEAE
OCHNACEAE	17	2	1
PICRODENDRACEAE	2	1
PHYLLANTHACEAE	16	4	2
ELATINACEAE	2
MALPIGHIACEAE	46	7	4
BALANOPACEAE
TRIGONIACEAE	2	1	1
DICHAPETALACEAE	3
EUPHRONIACEAE	1
CHRYSOBALANACEAE	13	1
LOPHOPYXIDACEAE
PUTRANJIVACEAE	1
PASSIFLORACEAE	7	2
LACISTEMATACEAE	2
SALICACEAE	18	3	1
VIOLACEAE	13	3	2
GOUPIACEAE	1
ACHARIACEAE	4		1
CARYOCARACEAE	2
HUMIRIACEAE	8		1
IRVINGIACEAE
LINACEAE	4
IXONANTHACEAE	2
CALOPHYLLACEAE	8	1
CLUSIACEAE	10	2	1
BONNETIACEAE	2
PODOSTEMACEAE	14	1	3
HYPERICACEAE	2
GERANIACEAE	2	1
FRANCOACEAE	1	4
COMBRETACEAE	4
LYTHRACEAE	11	2	1
ONAGRACEAE	4	3
VOCHYSIACEAE	5	1
MYRTACEAE	20	9	4
MELASTOMATACEAE	57	24	10
CRYPTERONIACEAE
ALZATEACEAE		1
PENAEACEAE
APHLOIACEAE
GEISSOLOMATACEAE
STRASBURGERIACEAE
STAPHYLEACEAE	1
GUAMETALACEAE
STACHYURACEAE
CROSSOSOMATACEAE
PICRAMNIACEAE	3	1
BIEBERSTEINIACEAE
NITRARIACEAE
KIRKIACEAE
BURSERACEAE	5
ANACARDIACEAE	14	9	2
SAPINDACEAE	26	7	2
RUTACEAE	31	7	4
SIMAROUBACEAE	7
MELIACEAE	6	2
GERRARDINACEAE
PETENAEACEAE
TAPISCIACEAE		1
DIPENTODONTACEAE		1
CYTINACEAE		1
MUNTINGIACEAE	1	2
NEURADACEAE
MALVACEAE	74	34	7
SPHAEROSEPALACEAE
THYMELAEACEAE	7	2
BIXACEAE	2	1
SARCOLAENACEAE
CISTACEAE	1	1
DIPTEROCARPACEAE		1
AKANIACEAE
TROPAEOLACEAE	1
TIGANOPHYTACEAE
MORINGACEAE
CARICACEAE	2
LIMNANTHACEAE
SETCHELLANTHACEAE
KOEBERLINIACEAE		1
BATACEAE	1
SALVADORACEAE
EMBLINGIACEAE
TOVARIACEAE		1
PENTADIPLANDRACEAE
GYROSTEMONACEAE
RESEDACEAE
CAPPARACEAE	11	8	1
CLEOMACEAE	6	4	1
BRASSICACEAE	4	45
AEXTOXICACEAE		1
BERBERIDOPSIDACEAE		1
BALANOPHORACEAE	6	1	1
OLACACEAE	2
APTANDRACEAE	3
COULACEAE	1
ERYTHROPALACEAE	2	1
STROMBOSIACEAE	1
XIMENIACEAE	3		1
MYSTROPETALACEAE
OCTOKNEMACEAE
OPILIACEAE	1
SANTALACEAE	7	5
LORANTHACEAE	12	5
MISODENDRACEAE		1
SCHOEPFIACEAE	2	1
FRANKENIACEAE		1
TAMARICACEAE
PLUMBAGINACEAE	2	2
POLYGONACEAE	10	5
DROSERACEAE	1
NEPENTHACEAE
MACARTHURIACEAE
KEWACEAE
DROSOPHYLLACEAE
DIONCOPHYLLACEAE
ANCISTROCLADACEAE
RHABDODENDRACEAE	1
SIMMONDSIACEAE
PHYSENACEAE
ASTEROPEIACEAE
CARYOPHYLLACEAE	10	12
ACHATOCARPACEAE	1
AMARANTHACEAE	20	11	4
STEGNOSPERMATACEAE
LIMEACEAE
LOPHIOCARPACEAE
BARBEUIACEAE
GISEKIACEAE
AIZOACEAE	2	2
PETIVERIACEAE	6	2
MICROTEACEAE	1
PHYTOLACCACEAE	1	2
SARCOBATACEAE
NYCTAGINACEAE	9	8	4
MOLLUGINACEAE	2
MONTIACEAE		7
DIDIEREACEAE
BASELLACEAE	1	2
HALOPHYTACEAE		1
TALINACEAE	1	1
PORTULACACEAE	1
ANACAMPSEROTACEAE		2
CACTACEAE	42	59	16
HYDROSTACHYACEAE
NYSSACEAE
CURTISIACEAE
GRUBBIACEAE
CORNACEAE		1
HYDRANGEACEAE		1
LOASACEAE	5	9
BALSAMINACEAE
MARCGRAVIACEAE	6	1
TETRAMERISTACEAE	1	1
FOUQUIERIACEAE
POLEMONIACEAE		12
LECYTHIDACEAE	10	1
SLADENIACEAE
PENTAPHYLACACEAE	2	1
SAPOTACEAE	21	2	1
EBENACEAE	2
PRIMULACEAE	12	5
THEACEAE	1
SYMPLOCACEAE	1
DIAPENSIACEAE
STYRACACEAE	1
SARRACENIACEAE	1
RORIDULACEAE
ACTINIDIACEAE		1
CLETHRACEAE	1	1
CYRILLACEAE	1
MITRASTEMONACEAE		1
ERICACEAE	13	19
ONCOTHECACEAE
METTENIUSACEAE	4	1
ICACINACEAE	3
EUCOMMIACEAE
GARRYACEAE
RUBIACEAE	121	47	16
GENTIANACEAE	30	14	7
LOGANIACEAE	5	1
GELSEMIACEAE	1
APOCYNACEAE	82	19	7
VAHLIACEAE
EHRETIACEAE	3	5	1
BORAGINACEAE	3	11
HYDROPHYLLACEAE		1
NAMACEAE		2
HELIOTROPIACEAE	3	1
CORDIACEAE	2
CODONACEAE
WELLSTEDIACEAE
CONVOLVULACEAE	20		1
SOLANACEAE	32	29	4
MONTINIACEAE
SPHENOCLEACEAE
HYDROLEACEAE	1
PLOCOSPERMATACEAE
CARLEMANNIACEAE
OLEACEAE	3	2
TETRACHONDRACEAE		2
CALCEOLARIACEAE	1	1
PELTANTHERACEAE		1
GESNERIACEAE	26	33	6
PLANTAGINACEAE	21	13	7
SCROPHULARIACEAE	3	1
STILBACEAE
LINDERNIACEAE	4		2
PEDALIACEAE
LAMIACEAE/LABIATAE	39	7	7
PHRYMACEAE		2
PAULOWNIACEAE
MAZACEAE
OROBANCHACEAE	10	5	3
WIGHTIACEAEA
LENTIBULARIACEAE	2	1
ACANTHACEAE	33	12	5
BIGNONIACEAE	29	10	1
THOMANDERSIACEAE
SCHLEGELIACEAE	1	2
VERBENACEAE	15	12
BYBLIDACEAE
MARTYNIACEAE	3	1	1
STEMONURACEAE	1
CARDIOPTERIDACEAE	1
PHYLLONOMACEAE		1
HELWINGIACEAE
AQUIFOLIACEAE	1
ROUSSEACEAE
CAMPANULACEAE	6	5
PENTAPHRAGMATACEAE
STYLIDIACEAE		2
ALSEUOSMIACEAE
PHELLINACEAE
ARGOPHYLLACEAE
MENYANTHACEAE	1
GOODENIACEAE	1	1
CALYCERACEAE	1	7
ASTERACEAE	271	257	73
ESCALLONIACEAE	1	3
COLUMELLIACEAE		2
BRUNIACEAE
PARACRYPHIACEAE
ADOXACEAE	2
CAPRIFOLIACEAE	1
PENNANTIACEAE
TORRICELLIACEAE
GRISELINIACEAE	1
PITTOSPORACEAE
ARALIACEAE	7	2
MYODOCARPACEAE
APIACEAE	10	27	1
	2767	1384	356
TOTAL	4151

BY ORDER

FAMILY	BRAZILIAN GENERA	UNBRAZILIAN GENERA	BRAZILIAN ENDEMIC GENERA
Amborellales
Nymphaeales	3	1
Austrobayleales
Canellales	2	1
Piperales	5	1
Magnoliales	35	3	3
Laurales	33	8	6
Chloranthales	1
Alismatales	55	12	7
Acorales
Petrosaviales
Dioscoreales	11	1
Pandanales	15	5
Liliales	3	5
Asparagales	240	142	31
Arecales	36	16	1
Commelinales	18	3	1
Poales	294	97	52
Zingiberales	17	2
Ceratophyllales	1
Ranunculales	20	12	1
Proteales	5	6
Tetrachondrales
Buxales	1	1
Gunnerales	1
Dilleniales	6
Saxifragales	5	10
Vitales	2	1
Zygophyllales	3	9
Fabales	230	60	24
Rosales	53	17	2
Fagales		10
Cucurbitales	24	8	3
Celastrales	23	4	2
Oxalidales	11	5	1
Malpighiales	273	38	21
Geraniales	3	5
Myrtales	101	40	15
Crossossomatales	1
Picramniales	3	1
Sapindales	89	25	8
Huertales		2
Malvales	85	42	7
Brassicales	25	59	2
Berberidopsidales		2
Santalales	40	14	2
Caryophyllales	111	117	24
Cornales	5	11
Ericales	73	45	1
Metteniusales	4	1
Oncothecales
Icacinales	3
Garryales
Gentianales	239	81	30
Vahliales
Boraginales	11	20	1
Solanales	53	29	5
Lamiales	190	105	32
Aquifoliales	1	1
Cardiopteridales	2
Asterales	280	272	73
Escalloniales	1	3
Desfontainiales		2
Bruniales
Paracryphyales
Dipsacales	3
Apiales	18	29	1
	2767	1384	356
TOTAL	4151

NEWS AND UPDATES

OCTOBER 31, 2023

∎ remarkable notes: Karremans rename Tubella (Luer) Archila as Karma Karremans (Harvard Papers in Botany, 2023), however, here the change is not considered due to our taking Tubella synonymized under Trichosalpinx Luer; here we take Brunelliaceae absent in Brazil. The only accurate information about the family in the country is the description of B. neblinensis Steyerm. & Cuatrec. (Steyermark, Annals of the Missouri Botanical Garden, Annals of the Missouri Botanical Garden, 1987), which cites a collection on Mount Neblina, at 1,500m altitude, without specifying which side of the mountain. Due to the extreme possibility that the species only occurs on the Venezuelan side, we maintain the exclusion. POWO does not recognize the species in Brazil (SEE).

∎ addition of the following genera (new or revalidated) missing in VPA: Condilorachia (Poaceae), the Brazilian species of Trisetum, with some from the Southern Cone, thus excluding this name from the flora of the country; Boldrinia, Calotheca, Erianthecium, Lombardochloa, Microbriza, Poidium, Rhombolytrum, Rosengurttia, all dismembered by Chascolythrum (Poaceae); Tumultivenia, Uniostium (Anacardiaceae), new names for South American members of Cyrtocarpa Kunth, the former for NE Colombia to NE Roraima state in Brazil, including Guyana and Venezuela, and the latter endemic to C & NE Brazil.

∎ full requalification of the data of the following genera/tribes, for which we will use a circumscription - linked - other than the VPA: Eperua (Fabaceae, Fortes et al., Phytotaxa, 2023, with 19 spp.), Erythrina (only for Brazil, Fabaceae, Guedes-Oliveira et al., PhytoKeys, 2023, same number as VPA), Begonia (only for Peru, Begoniaceae, Moonlight et al., European Journal of Taxonomy, 2023, -9 Sspp. if compared with VPA), Acalypha (only for South America, Euphorbiaceae, Cardiel et al., European Journal of Taxonomy, 2023, -40 Sspp. if compared with VPA), Bougainvillea (Nyctaginaceae, Bautista et al., PLANTS, 2022, with 11 spp.), Cuspidaria (Bignoniaceae, Francisco et al., Taxon, 2023, with 19 spp., excluding C. bracteata), Burmeistera (only for Colombia, Campanulaceae, González, Phytotaxa, 2023, with 57 spp. in country).

∎ acceptance of synonymizations at generic level, but not present in the VPA: Aspidogyne Garay, Kreodanthus Garay, Ligeophila Garay, Platythelys Garay, Rhamphorhynchus Garay and Stephanothelys Garay under Microchilus C.Presl (Orchidacae, via POWO); Gallardoa Hicken and Cordobia Nied. under Mionandra Griseb. (Malpighiaceae, Almeida et al., Plant Ecology and Evolution, 2023); Catimbaua L.P.Felix, Christenh. & E.M.Almeida, Isabelcristinia L.P.Felix, Christenh. & E.M.Almeida under Ameroglossum Eb. Fisch., S. Vogel & A.V. Lopes (Linderniaceae, Santos et al., Molecular Phylogenetics and Evolution, 2023); Nordenstaimia Rich. under Gynoxys L. (Escobari, Borsch & Kilian, PhytoKeys, 2018).

∎ inclusion of new records for Brazilian states: NONE.

∎ new records at the genus level for other South American countries: NONE.

∎ new generic records for Brazil apart VPA, for genera yet in VPA: Burmeistera Karst. & Triana (Campanulaceae), collected in the Monte Neblina region (Catalogo UCS).

∎ rejection of the Brazilian occurrence of the following valid species: NONE.

∎ simple synonymizations, at species level: Lorostemon negrense under L. colombianum (Clusiaceae, BR/CL/VZ, 2022), Bacopa llanorum under Lindernia brachyphylla (Plantaginaceae up to Linderniaceae, CL/VZ, 2023), Stachytarpheta viscidula under S. commutata (Verbenaceae, BR, 2023).

∎ inclusion of new national records at species level (by data revision, or by new discoveries): Ocotea fistulosa, O. pautensis, Andea homeieri (Lauraceae, new in CL, 2023, formerly in EC), Mimosa asperoides (Fabaceae, new in BR, 2023, formerly in Cono Sur), Ditaxis [Argythamnia] salina (Euphorbiaceae, new in BR, 2023, formerly in AR, PAR).

∎ distribution reduction, apart the next topic: Senna splendida (Fabaceae, excluded in GU's/BL, now only BR/PAR, 2023).

∎ emancipations, rank elevations or re-establishments at species level: Acianthera kautskyi (elevation rank, Orchidaceae, BR, 2023), Celtis alnifolia (re-established, Cannabaceae, BR, 2023), Begonia corallina (re-established, outside B. maculata, Begoniaceae, BR, 2022), Macropsychanthus nitidus (re-established, ex-Dioclea rostrata var. nitida, Fabaceae, BR, 2022), Senna gloriosa (elevation rank, Fabaceae, BR/BL, 2023), Dolichandra coccinea (re-established, Bignoniaceae, BR, 2023).

∎ consolidated species recognized here, absent in the VPA: Cleistes castaneoides (Orchidaceae, BR, sometimes synonimized under Cleistes rosea), Nasa hastata (Loasaceae, EC/PR, 2023).

∎ 18⋅10⋅23 ‣ rediscovered species, presents in VPA: Thismia caudata (Thismiaceae, BR, 2023), Prosthechea sessiliflora (Orchidaceae, BR, 2023), Aristida ekmaniana, A. macrophylla (Poaceae, BR, 2023), Nasa colanii, N. ferox, N. hastata, N. humboldtiana, N. ramirezii, N. solaria (Loasaceae, EC/PR, 2023).

∎ simple-species renames: Catasetum lendarium (Orchidaceae, ex-C. meeae, BR, 2023), Epidendrum vulcanitungurahuae (Orchidaceae, ex-E. tungurahuae , PR, 2023), Mucuna pungens (Fabacae, ex-Dolichos pungens, BR, 2023), Lucuma kossmanniae sp. (Sapotaceae, ex-Pouteria kossmanniae, BR, 2023), Scyphostelma erikseniae (Apocynaceae, ex-Cynanchum erikseniae, EC, 2023), S. fasciculiﬂorum (Apocynaceae, ex-C. fasciculiﬂorum, EC, 2023), S. jaramilloi (Apocynaceae, ex-C. jaramilloi , EC, 2023), S. quitensis (Apocynaceae, ex-C. quitensis, EC, 2023), S. stenospira (Apocynaceae, ex-C. stenospira, EC, 2023), S. unguiculatum (Apocynaceae, ex-C. unguiculatum, PR, 2023), S. purpurascens (Apocynaceae, ex-Metastelma purpurascens, EC, 2023).

∎ addition to our list of 81 species absents in VPA South America list in April 3, 2023, being 77 ENDEMICS among countries: Brazil (38), Colombia (15), Ecuador (11), Peru (6), Bolivia (5), Guianas (1), Uruguay (1); and 4 NON ENDEMICS: as BR/VZ(1)⋅ BR/GU´s (1) ⋅ CL/PR(1) ⋅ EC/PR (1): Aristolochia franzii (Aristolochiaceae, BR/FG, 2023), Persea basiobtusa, P. dryadum (Lauraceae, EC, 2023), Aiouea chicaque (Lauraceae, CL, 2023), Aniba brochidodroma, A. glabra, A. palaciosii, A. tomentella, A. verticillata (Lauraceae, CL[2], EC[3,5], PR[1,4], 2023), Mollinedia pignalii (Monimiaceae, BR, 2023), Anthurium sp. (Araceae, CL, 2023), Dieffenbachia simoneae (Araceae, BR, 2022), Philodendron madalenense (sect. Macrobelium, Araceae, BR, 2023), Thismia violacea (Thismiaceae, BR, 2023), Barbacenia glaucescens, B. mellosilvae (Velloziaceae, BR, 2023), Vanilla rupicola (Orchidaceae, BR, 2023), Vanilla cameroniana (Orchidaceae, FG, 2023), Pelexia [Pachygenium] muyscarum (Orchidaceae, CL, 2023), Cyrtopodium izaguirreae (Orchidaceae, UR, 2023), Bulbophyllum parex (Orchidaceae, CL, 2023), Catasetum krahlii (Orchidaceae, BR, 2023), Eltroplectris paranaënsis (Orchidaceae, BR, 2023), Epidendrum edquenii (Orchidaceae, PR, 2023), Lepanthes bachue (Orchidaceae, CL, 2023), Masdevallia leonor-baeziana (Orchidaceae, CL, 2023), Masdevallia emieliana (Orchidaceae, PR, 2023), Octomeria giordanii (Orchidaceae, BR, 2023), Pleurothallis celsia (Orchidaceae, CL, 2023), Pleurothallis franciana, P. petroana (Orchidaceae, CL, 2023), Pleurothallis kashi-menkakarai, P. lapoi, P. marioandresavilae, P. sabanillae, P. tinajillensis (Orchidaceae, EC, 2023), Pleurothallis inaudita (Orchidaceae, EC, 2023), Telipogon mariae-luisae (Orchidaceae, PR, 2023), Dichorisandra pulcherrima (Commelinacae, BR, 2023), Syagrus aristeae (Arecaceae, BR, 2023), Tillandsia alcatrazensis (Bromeliaceae, BR, 2023), Tillandsia chingacensis (Bromeliaceae, CL, 2023), Forzzaea bahiana (Bromeliaceae, BR, 2023), Krenakanthus ribeiranus (Bromeliaceae, BR, 2023), Merostachys nigricans (Poaceae, BL, 2023), Macropsychanthus ruschii (Fabaceae, BR, 2023), Eriosema parvifolium (Fabaceae, BR/VZ, 2023), Senna dryadica, S. franciscana (Fabaceae, BR, 2023), Inga micronectarium (Fabaceae, BR, 2023), Celtis flumeniana (Cannabaceae, BR, 2023), Begonia littoralis (Begoniaceae, BR, 2023), Begonia piraquara (Begoniaceae, BR, 2023), Begonia sp. (Begoniaceae, CL, 2023), Passiflora yolandae (Passifloraceae, EC, 2023), Protium inversum (Burseraceae, EC/PR, 2023), Microlicia diamantinensis, M. sublaevis, M. reptans (Melastomataceae, BR, 2023), Microlicia indurata (Melastomataceae, BR, 2023), Pleroma curucutuense (Melastomataceae, BR, 2023), Poteranthera inopinata (Melastomataceae, BL, 2023), Merianthera calyptrata (Melastomataceae, BR, 2023), Eugenia campininha (Myrtaceae, BR, 2023), Pachira inaequalivalvis, P. deflexifolia (Malvaceae, BR, 2023), Neea bahiensis, N. manaosensis (Nyctagiaceae, BR, 2023), Guettarda [Tournefortiopsis] triflora (Rubiaceae, CL, 2023), Allamanda alagoana (Apocynaceae, BR, 2023), Scyphostelma bolivianum, S. gracile, S. rotorum, S. solomonii (Apocynaceae, PR[3], BL[1,2,4], 2023), Petunia toropiensis (Solanaceae, BR, 2023), Iochroma orozcoae (Solanacae, CL, 2023), Solanum confertiflorum (Solanaceae, BR, 2018), Solanum kollastrum (Solanaceae, BR, 2018), Schlegelia longirachis (Schlegeliaceae, CL, 2023), Mendoncia amabilis (Acanthaceae, CL/PR, 2023), Stachytarpheta lajedicola (Verbenaceae, BR, 2023).

∎ exclusion from our list of occurrence in the New World of these species or genera: NONE.

AUGUST 06, 2023

∎ although important works have emancipated Hydnoraceae, Lactoridaceae (Jost et al., Frontiers in Plant Science, 2021) and Parnassiaceae (Simmons, Lombardi & Biral, Systematic Botany, 2023) from Aristolochiaceae and Celastraceae, here we still consider the unions, in accordance with the APG Website (SEE).

∎ important changes at the generic level accepted: the emancipation of Acanthinophyllum Allemão from Clarisia in Moraceae (Gardner, Molecular Phylogenetics and Evolution, 2023); the displacement of all New World Polygala L. species to Senega Spach. in Polygalaceae (Pastore et al., Annals of the Missouri Botanical Garden, 2023); the immersion of Afrocalliandra E.R.Souza & L.P.Queiroz (endemic of Africa) in Calliandra Benth., in the Fabaceae (Thulin, Phytotaxa, 2023); the consolidation of Abarema Pittier as a endemic genus in Brazil, in Fabaceae (Guerra et al., Phytotaxa, 2023); the expansion of Euxylophora Huber to Colombia and its exclusion from the list of endemic genera in Brazil in Rutaceae (Echeverri et al, Phytotaxa, 2023); the renaming of Acantholippia Griseb. to Troncosoa N. O’Leary & P. Moroni, and the emancipation of Salimenaea N. O’Leary & P. Moroni from Lippia L., in Verbenaceae (O'Leary et al., Novon, 2023); the recognition of Ancashia Al-Shehbaz, Salariato, A. Cano & Zuloaga, Borealandea Al-Shehbaz, Salariato, A. Cano & Zuloaga, and Stenodrabopsis Al-Shehbaz, Salariato, A. Cano & Zuloaga, and the imersion of Brayopsis Gilg & Muschl. within Eudema Bonpl. in Brassicaceae (Al-Shehbaz, Annals of the Missouri Botanical Garden, 2023); the re-estabilishment of Aphanandrium Lindau from Panamá to Venezuela and Peru, in Acanthaceae (Cornejo et al., Harvard Papers in Botany, 2023); addition of the following changes at the generic level: addition of Achrouteria Eyma., Chloroluma Baill., Cornuella Pierre, Englerella Pierre, Labatia Sw., Lucuma Molina, Martiusella Pierre, Nemaluma Baill., Peteniodendron Lundell, Prieurella Pierre, Ragala Pierre, all these in Sapotaceae and present in Brazil except Cornuella; Stapfochloa (Poaceae, tropical New World inc. Brazil); Adolphoduckea Paudyal & Delprete, Motleyothamnus Paudyal & Delprete, and Coutareopsis Paudyal & Delprete in Rubiaceae, the first in Brazil; Cipurospsis (Bromeliaceae) and Maguireothamnus (Rubiaceae) as are new genera for Brazil; exclusion of Philippiamra Kuntze (Montiaceae, included in Cistanthe) for South America and Exostema (Rubiaceae) in Brazil.

∎ emancipations, renames, rank elevations and re-establishments accepted: Nymphaea pedersenii (former N. amazonum subsp. pedersenii, Nymphaeaceae, BR/AR/PAR/UR, 2023), Dyckia tomentosa (former name unknown, Bromeliaceae, BR, 2023); Ficus llanensis, F. llewelynii, F. mitrophora (emancipated species, former names unvailable, Moraceae, range unknown, 2023), Eugenia pisonis (Myrtaceae, BR, 2023).

∎ simple renames: Chromolaena amambayensis (Asteraceae, ex-Eupatorium amambayense, PAR, 2023).

∎ here we consolidate the occurrence of Ennealophus N.E.Br in Brazil via POWO (Iridaceae, SEE), despite the denial of Reflora (SEE); in addition, we consider here the non-occurrence of two contoversial species: Cantua megapotamica Spreng. (Polemoniacae, SEE) and Juglans brasiliensis Dode (Juglandacaee, SEE).

∎ species no cited in VPA accepted here: Rubus pendulus (Rosaceae, CL/EC/PR).

∎ deletions and synonimizations: Catasetum sanguineum under C. naso (Orchidaceae, BR, 2023), Dyckia polyclada under unknown species (Bromeliaceae, BR, 2023), Bredemeyera altissima, B. lucida under B. divaricata, B. myrtifolia, B. densiflora under B. bracteata, B. disperma under B. laurifolia. (Polygalaceae, tropical South America, 2023; Brazil loss 4 of these names), Arapatiella emarginata under A. psylophylla (Fabaceae, BR, 2023), Borreria catolensis under Psyllocarpus intermedius (Rubiaceae, BR, 2023).

∎ new records accepted: Peperomia cacaophila (Piperaceae, new in PR, 2023; formerly only in EC), Parkia barnebyana (Fabaceae, new in BR, 2023; former only in VZ), Monnina salicifolia, M. smithii (Polygalaceae, new in VZ, 2023; formerly in CL/EC/PR/BL), Macrolobium machaerioides (Fabaceae, new in EC, 2023; formerly in CL/PR), Cedrela kuelapensis (Meliaceae, new in EC, 2023; formerly in PR).

∎ addition to our list of 129 species absents in last update, being 117 ENDEMICS among countries: Brazil (61), Colombia (19), Ecuador (13), Peru (17), Bolivia (4), Argentina (2), Paraguay (1); and 12 NON ENDEMICS: as GU/SR (+AMC); CL/EC/PR; CL/EC/PR; VZ/CL/EC/PR; CL (+ AMC); CL/EC; CL/EC; CL/EC; BR/CL; BR/PR; BR/PR; BR/BL: Nymphaea caatingae, N. francae, N. harleyi, N. paganuccii, N. rapinii (Nymphaeaceae, BR, 2023), Magnolia buenaventurensis (Magnoliaceae, EC, 2023), Compsoneura nallarettiana (Myristicaceae, PR, 2023), Aristolochia brachylimba (Aristolochiaceae, PR, 2023), Piper oteguanum, P. rubrifolium, P. caguanense, P. laperdizense (Piperaceae, CL; PR; CL; EC, 2023), Peperomia pilocarpa, P. riosaniensis, P. sagasteguii, P. symmankii, P. vivipara (Piperaceae, PR, 2023), Piper quinchasense (Piperaceae, CL, 2023), Piper svenningii (Piperaceae, CL/EC/PR, 2023); Mollinedia arianeae (Monimiaceae, BR[RJ], 2023), Anthurium pluricarinatum (Araceae, BR, 2023), Philodendron agudeloi, P. arevaloi, P. crystallum, P. dalyi (Araceae, CL, 2023), Philodendron josephii (Araceae, BR, 2023), Thismia pseudomelanomitra (Thismiaceae, BR, 2023), T. paradisiaca (Thismiaceae, CL, 2023), Hippeastrum curupira, H. laklano (Amaryllidaceae, BR, 2023), Vanilla rupicola (Orchidaceae, BR, 2023), Vanilla calamitosa (Orchidaceae, BR, 2023), Bulbophyllum parex (Orchidaceae, CL, 2023), Liparis altomayoënsis (Orchidaceae, PR, 2023), Epidendrum centralense (Orchidaceae, PR, 2023), Lepanthes carrizosana (Orchidaceae, CL, 2023), Lepanthes cordillerana (Orchidaceae, CL, 2023), Restrepia santanderensis (Orchidaceae, CL, 2023), Syagrus carvalhoi (Areaceae, BR[MG], 2023), Dyckia magnifica (Bromeliaceae, BR, 2023), Stigmatodon lemeanus (Bromeliaceae, BR, 2023), Chusquea riparia (Poaceae, BR, 2023), Guadua leonardoana (Poaceae, BR, 2023), Chamaecostus manausensis, Costus alfredoi, C. antioquiensis, C. callosus, C. cochabambae, C. convexus, C. douglasdalyi, C. gibbosus; C. obscurus; C. oreophilus, C. pitalito, C. prancei, C. pseudospiralis, C. rubineus, C. whiskeycola (Costaceae, BR; BL; VZ/CL/EC/PR; CL (+CR, PN); BL; CL/EC; BR/PR; EC; PR; EC; CL; BR/PR; BR/BL; PR; CL/EC/PR, 2023), Oxalis jacobinensis (Oxalidaceae, BR, 2023), Erythroxylum confertifolium (Erythroxylaceae, BR, 2023), Amanoa condorensis (Phyllanthaceae, EC, 2023), Phyllanthus megastylus (Phyllanthaceae, BR, 2023), Phyllanthus capixaba (Phyllanthaceae, BR, 2023), Turnera carolina (Passifloraceae, BR, 2023), Passiflora rosacea (Passifloraceae, BL, 2023), Croton restingae (Euphorbiaceae, BR[RJ], 2023), Dalechampia seccoi (Euphorbiaceae, BR, 2023), Asemeia coracoralinae, A. minensis, A. nana (Polygalaceae, BR, 2023), Bredemeyera ericifolia (Polygalaceae, BR, 2023), Schnella madeleinae (Fabaceae, CL, 2023), Eperua cerradoensis, E. manausensis (Fabaceae, BR, 2023), Macrolobium ceriferum, M. longistipitatum (Fabaceae, CL, 2023), Macrolobium paulobocae (Fabaceae, BR, 2023), Indigofera morroensis (Fabaceae, BR[BA], 2023), Mimosa crassifolia, M. bahiana, M. confusa, M. melosa (Fabaceae, BR, 2023), Protium herisonii (Burseraceae, BR, 2023), Cedrela angusticarpa (Meliaceae, EC, 2023), Zanthoxylum palustre (Rutaceae, BR/CL, 2023), Podandrogyne flammea, P. websteri (Cleomaceae, CL/EC, 2023), Cardamine peruviana (Brassicaceae, PR, 2023), Vochysia wilsonii (Vochysiaceae, CL, 2023), Myrcia [Calyptranthes] magna (Myrtaceae, EC, 2020), Myrciaria caerulescens (Myrtaceae, BR, 2023), Myrcia capixaba, M. forzzae (Myrtaceae, BR, 2023), Microlicia rosanae, M. septentrionalis (Melastomataceae, BR, 2023), Microlicia delicata, M. pumila (Melastomataceae, BR, 2023), Tibouchina falcifolia, T. longisquamata (Melastomataceae, BR, 2023), Pterolepis xaxa (Melastomataceae, BR, 2023), Catostemma lanceolatum (Malvaceae, BR, 2023), Heisteria austroecuadorica (Erythropalaceae, EC, 2023), Coccoloba efigeniana (Polygonaceae, GUY/SR, also AMC), Neea rubescens (Nyctaginaceae, BR, 2023), Cumulopuntia mollispina (Cactaceae, PR, 2023), Parodia hofackeriana (Cactaceae, BR, 2023), Arrojadoa flava (Cactaceae, BR, 2023), Eschweilera foetulenta, E. magnifica (Lecythidaceae, EC, 2023), Myriopus gleissonii (Heliotropiaceae, CL, 2023), Varronia elsieae (Cordiaceae, BR, 2023), Petaloselma garrapatense, Philibertia escoipensis (Apocynaceae, AR, 2023), Matelea anomala (Apocynaceae, BL, 2023), Chionanthus monteazulensis (Oleaceae, BR, 2023), Besleria naquenensis (Gesneriacae, CL, 2023), Monopyle chocoensis, M. connata (Gesneriaceae, CL/EC, 2023), Hyptidendron dorothyanum (Lamiaceae, BR, 2023), Marsypianthes dunensis (Lamiaceae, BR, 2023), Salvia cajamarcana (Lamiaceae, PR, 2023), Aphanandrium narupayacuensis (Acanthaceae, EC, 2023), Stachytarpheta longibracetata, S. rizzoi (Verbenaceae, BR, 2023), Pinguicula jimburensis, P. ombrophila (Lentibulariaceae, EC, 2023), Nolana hoxeyi, N. samaensis (Solanaceae, PR, 2023), Solanum helix (Solanaceae, BR, 2023), Lepidaploa estevesiana (Asteraceae, BR, 2023), Eryngium cerradense (Apiaceae, PAR, 2023).

MAY 20, 2023

∎ inclusion of data among the family Afrothismiaceae, dismembered of Thismiaceae (Bio rXiv).

APRIL 15, 2023

∎ addition of the following genera (new or revalidated) missing in VPA: Thaumatophyllum (Araceae, dismembered from Philodendron, tropical America, 2018), Krenakanthus, Siqueiranthus, Orthocryptanthus (Bromeliaceae, BR, 2022), Actinocephalus, Cora, Cryptanthella, Floralia, Giulettia, Gnomus, Hydriade, Monosperma (Eriocaulaceae, tropical New World, 2023; the number of species in the genus Paepalanthus Mart. has been reduced), Aenigmanu (Picramniaceae, BR/PR, 2021), Gyrosphragma (Lythraceae, BR, 2022), Mahechadendron (Vochysiaceae, CL, 2022), Diadorimia (Rubiaceae, BR, dismembered of Psyllocarpus, 2021).

∎ full requalification of the data of the following genera/tribes, for which we will use a circumscription - linked - other than the VPA: Lemnoideae (Araceae, Bog, Appenroot & Sree, Nordic Journal of Botany, 2020, Hagenbachia (Asparagaceae, POWO), Phytelephas (Arecaceae, Escobar et al., Molecular Phylogenetics and Evolution, 2022, with 6 spp.), Canna (Cannaceae, H.Maas-van de Kamer & P.J.M. Maas, Blumea, 2008), Sedum (Crassulaceae, Messerschmid & al., Taxon, 2020; version expanded, including almost all New World Crassluaceae except Crassula and some small genera), Quinchamalium (Schoepfiaceae, POWO, with only a single species; many sources points up to 23 spp.), Petunia (Solanaceae, VER, rejecting Mexican records), Capsicum (Solanaceae, Gloria E. Barboza et al., PhytoKeys, 2022), Sclerophyllax (Solanaceae, Chiarini et al., Taxon, 2022).

∎ acceptance of synonymizations at generic level, but not present in the VPA: Limnobium Rich. under Hydrocharis L. (Byng & Christenh., Global Fl., 2018), Egeria Planch. and Apalanthe Planch. under Elodea L. (Byng & Christenh., Global Fl., 2018), Lillaea Bonpl. under Triglochin L. (Mering & Kadereit, Molecular Phylogenetics and Evolution, 2015), Eucharis, Caliphruria under Urceolina (Amaryllidaceae, Byng, Christenhusz & Fay, Flora Global, 2018; Urceolina is not endemic to Peru now), Habranthus, Sprekelia, Haylockia, Rhdophiala p.p. under Zephyranthes (Amaryllidaceae, Taxon, 2019), Leucothauma under Pyrolirion (Amaryllidaceae, POWO), Crocopsis under Clinanthus (Amaryllidaceae, POWO), Lytocaryum Toledo under Syagrus Mart. (Arecaceae, Noblic & Meerow, Palms, 2015), Huanaca Cav., Mulinum Pers., Schizeilema Hook. f. under Azorella (Plunkett e Nicolas, Brittonia, 2016).

∎ inclusion of new records for Brazilian states: Peridiscus in Amapá (Aymard & Arellano, Harvard Papers in Botany, 2018).

∎ new records at the genus level for other South American countries: Peridiscus in CL (Aymard & Arellano, Harvard Papers in Botany, 2018).

∎ new generic records for Brazil apart VPA, for genera yet in VPA: Cressa (Convolvulaceae, Reflora 2020), Darcyanthus (Solanaceae, Phytotaxa, 2020), Ammoselinum (Apiaceae, REFLORA).

∎ rejection of the Brazilian occurrence of the following valid species: Crassula longipes (Crassulaceae, asignated as Brazilian by VPA, however rejected here by Giuffre (Dissertation, 2019) and POWO), Kallstroemia maxima (Periódicos UEFS, 2014; all Kalstroemia recors in Brazil belongs K. tribuloides); Deprea (Solanaceae, cited in Brazil by Reflora, but rejected in Brazil by VPA).

∎ simple synonymizations, at species level: Barbacenia burlemarxii under B. pabstiana (Velloziaceae, BR, 2022), Barbacenia gaveensis, B. foliosa, B. seubertiana under B. squamata (Velloziaceae, BR, 2022), Catasetum ciliatum under C. roseo-album (Orchidaceae, South America, 2021), Clowesia amazonica under unknown species (Orchidaceae, N South America, 2022), Microchilus longicornu under unknown species (Orchidaceae, BR, 2022), Epidendrum amarajiense, E. campaccii, E. thiagoi under unknown species (Orchidaceae, BR, 2021), Vriesea vexata under Stigmatodon sp. (Bromeliacae, BR, 2022), Vriesea sp.1, V.sp.2, V.sp.3, V.sp.4, V.sp.5, V.sp.6, V.sp.7, V.sp.8 under Stigmatodon (Bromeliaceae, BR, 2022), Paepalanthus bongardii under Actinocephalus camptophyllus (Eriocaulaceae, BR, 2021), Paepalanthus macrocephalus under P. argyropus (Eriocaulaceae, BR, 2021), Paepalanthus melanolepis, P. pseudoelongata under Actinocehalus macrocephalus (Eriocaulaceae, BR, 2021), Paepalanthus petraeus under P. uai (Eriocaulaceae, BR, 2021), Paepalanthus decorus under P. flaccidus (Eriocaulaceae, BR, 2021), Carex obtusisquama under C. crassiflora (Cyperaceae, AR/BL, 2021), C. tessellata under C. acutata (Cyperaceae, AR/BL, 2021), Agrostis meridensis under Podagrostis meridensis (Poaceae, VZ, 2021), Maranta anderssoniana, M. hatschbachiana under unknown species (Marantaceae, BR, 2021), Celtis morifolia under Celtis flavovenarum (Cannabaceae, BR, 2023), Oxalis irreperta under O. riparia (Oxalidaceae, BR, 2021), Macroptilium longepedunculatum under under unknown species (Fabaceae, Mexico to Uruguay except Brazil, 2022), Senega [Polygala] coridifolia under Senega [P.] aspalatha (Polygalaceae, BR, 2022), Passiflora tolimana, P. gleasonii, P. metae under P. acuminata (Passifloraceae, BR/VZ/GU/CL, 2022), Passiflora rupestris under P. pohlii (Passifloraceae, BR, 2022), Phyllanthus atalaiensis under P. heteradenius (Phyllanthaceae, BR, 2022), Stillingia scutellifera under S. salpingadenia (Euphorbiaceae, AR/PAR, 2019), Dalechampia brevipedunculata under D. magnoliifolia (Euphorbiaceae, BR/VZ/BL/SR, 2019), Dalechampia uleana under D. cujabensis (Euphorbiaceae, BR/VZ/BL/SR, 2019), Croton rufo-argenteus under C. tricolor (Euphorbiacae, BR, 2021), Croton subcompressus under C. compressus (Euphorbiacae, BR, 2021), Croton guildingii under C. suavis (Euphorbiaceae, VZ/CL/CRB, 2021), Croton allemii under C. triqueter (Euphorbiaceae, BR, 2021), Croton stenotrichus under C. hilarii (Euphorbiaceae, BR, 2021), C. apurensis, C. glauca, C. ramulosa, C. sessilifolia under C. antisyphilitica (Euphorbiaceae, BR/VZ, 2023), Croton brachiata under C. flava (Euphorbiaceae, BR, 2023), Cardiospermum urvilleoides under Serjania urvilleoides (Sapindaceae, BR, 2022), Conchocarpus limae, C. lilacinus under C. ruber (Rutaceae, BR, 2022; all as Almeidea in VPA), Plinia martinellii under Eugenia sp. 1 (Myrtaceae, BR, 2021), Plinia sebastianopolitana under Eugenia sp.2 (Myrtaceae, BR, 2021), Microlicia fasciculata, M. variolosa var. hirsuta under M. hirsuta (Melastomataceae, BR, 2022), Microlicia nortecipoana, M. petiolulata under M. hirtoferruginea (Melastomataceae, BR, 2021), Microlicia souza-limae under M. helvola (Melastomataceae, BR/BL, 2020), Sida pseudorubifolia under S. rubifolia (Malvaceae, BR, 2022), Pachira dolichocalyx under P. macrocalyx (Malvaceae, GU/GF, 2021), Pachira nitida under P. minor (Malvaceae, tropical America, 2021), Sabulina altoandina under Arenaria pycnophylloides (Caryophyllaceae, AR, 2022), Arenaria palustris under A. serpens (Caryophyllaceae, PR, 2022), Alternanthera piptantha under A. altacruzensis (Amaranthaceae, Cono Sur, 2022), Ternstroemia borbensis under T. dentata (Pentaphyllacaceae, BR, 2021), Evolvulus linarioides under Evolvulus saxifragus (Convolvulaceae, BR, 2022), Cuscuta orbiculata under Cuscuta tinctoria (Solanaceae, BR, 2021), Jacquemontia heterantha under J. cumanensis (Convolvulaceae, BL/PR, 2021), Jacquemontia prominens under J. unilateralis (Convolvulaceae, BL/PR, 2021), Carapichea altsonii, C. nivea, C. sandwithiana under Notopleura (Rubiaceae, GF, 2022), Carapichea pacimonica under C. ligularis (Rubiaceae, GF, 2022), Psychotria squamelligera under Carapichea squamelligera (Rubiaceae, GF, 2022), Appunia brachycalyx under A. morindoides (Rubiaceae, BR/GU/GF, 2021), Borreria quadrifaria under B. rubrostipulata (Rubiaceae, BR/BL/CS, 2022), Sanchezia stenomacra under S. aurea (Acanthaceae, BR, 2021), S. cyathibractea under S. macrocnemis (Acanthaceae, BR, 2021), S. bicolor, S. flava, S. megalia, S. speciosa under S. oblonga (Acanthaceae, BR, 2021), S. capitata, S. stenantha under S. ovata (Acanthaceae, BR, 2021), S. lutea under S. parviflora (Acanthaceae, BR, 2021), S. skutchii under S. putumayensis (Acanthaceae, BR, 2021), S. arborea, S. decora under S. scandens (Acanthaceae, BR, 2021), Leucheria cerberoana, L. cumingii, L. daucifolia, L. glabriuscula, L. glandulosa, L. menana, L. mille-folium, L. multiflora, L. tenuis under L. tomentosa (Asteraceae, CH, 2021), Pentacalia viburnifolia, P. brittoniana, P. miguelii under P. psidiifolia (Asteraceae, BL, 2021), P. inquisiviensis under P. cardenasii (Asteraceae, BL, 2021), P. sailapatensis under P. urubambensis (Asteraceae, BL, 2021), Pentacalia hitchcockii, P. huamaliensis under P. theifolia (Asteraceae, PR, 2019), P. lanceolifolia under P. andrei (Asteraceae, PR, 2019), P. tillettii under P. riotintis (Asteraceae, PR, 2019), Stevia breviaristata under S. vaga (Asteraceae, AR, 2021), Senecio phylloleptus under S. reicheanus (Asteraceae, CH, 2022), S. behnii under S. eriocladus (Asteraceae, CH, 2022), Praxelis macrocarpa under P. capillaris (Asteraceae, BR, 2021), Lucilia lycopodioides under L. adpressa (Asteraceae, BR/AR/BL, 2022).

∎ inclusion of new national records at species level (by data revision, or by new discoveries): Aristolochia goudotii (Aristolochiaceae, new in BR, 2019, formerly in CL), Limnocharis laforestii (Alismataceae, new in BR; former widely in tropical New World), Halodule beaudettei (Cymodoceaceae, new in BR, 2017, formerly in AMC, CRBN), Maxillaria aureoglobula (Orchidaceae, new in BR and VZ, 2016, formerly in CL), Scaphyglottis punctulata (Orchidaceae, new in BR, 2022, formerly from AMC do CL and CRBN), Trithrinax schizophylla (Arecaceae, new in BR, 2015, formerly BL, AR, PAR), Cyperus oxylepis (Cyperaceae, new in BR, 2022, formerly in CAM to Cono Sur), Monophyllanthe araracuarensis (Marantaceae, new in BR; former only in Colombia), Begonia cinnabarina (Begoniaceae, new in AR, 2022, formerly un PR, BL), Tristicha trifaria (Podostemaceae, new in CL/EC, 2023, formerly in BR/AR/PAR/BL/GU/CAM), Lophogyne aeruginosa (Podostemaceae, new in EC, 2023, formerly in VZ/SR), Passiflora garckei (FAMILY, new in BR, 2014, formerly in CL/VZ/GU), Ancistrothyrsus scopae (Passifloraceae, new in BR, 2020, formerly in GU'S), Dalechampia brownsbergensis (FAMILY, new in BR, 2019, formerly in VZ/SR), Dalechampia attenuistylus (FAMILY, new in BR, 2019, formerly in VZ/BL), Euphorbia riinae (Euphorbiaceae, new in BR, 2022, formerly in BL), Croton lombardianus (Euphorbiaceae, new in BR, 2021, formerly in Cono Sur), Sida hassleri (Malvaceae, new in BR, 2022, formerly in Cono Sur), Microphyes minima (Caryophyllaceae, new in AR, 2022, formerly in CH), Cordia weddellii (Cordiacae, new in BR, 2021, formerly in BL), Varronia urticifolia (Cordiacae, new in AR, 2022, formerly in BR), Euploca krapovickasii (Heliotropiaceae, new in UR, 2021, formerly in BR), Bonamia rosiewiseae (Convolvulaceae, new in Brazil, former in topical America, 2017), Voyria alvesiana (Gentianaceae, new in SR/GF, 2022, formerly in Brazil), Stemodia lanceolata (Plantaginaceae, new in BR, 2018, formerly in BL/Cono Sur), Aphelandra martiusii (Acanthaceae, new in CL/PR, 2022, formerly in BR).

∎ distribution reduction, apart the next topic: U. amesthytina (Lentibulariacae, excluded in MX/BLZ, now only BR/VZ, 2021).

∎ emancipations, rank elevations or re-establishments at species level: Paepalanthus politus (elevation-rank, Eriocaulaceae, BR, 2022), Chusquea simplicissima, C. weberbaueri (elevation-rank, outside C. spicata, Poaceae, PR, 2019), Sorghastrum canescens (reestablishment, Poaceae, BR, 2021), Cenostigma diversifolium (re-established, Fabaceae, BR, 2021), Tetrathylacium nutans (re-established, Salicacae, EC, 2019), Cnidoscolus neglectus (re-established, Euphorbiaceae, BR, 2021), Croton lagunillae (elevation-rank, outside C. guildingii, Euphorbiaceae, VZ, 2021), Campomanesia repanda, C. rhombea (re-established, Myrtaceae, BR, 2021), Microlicia acerosa (elevation-rank, ex-Microlicia linifolia var. naudiniana, Melastomataceae, BR, 2021), Pleroma imperatore (re-established, Melastomataceae, BR, 2021), Pachira manausensis (re-established, ex-P. aquatica var. manausensis, Malvaceae, BR, 2022), Schinus minutiflora, S. subtridentata (elevation rank, Anacardiaceae, BL, 2022), Portulaca ferricola (former P. mucronulata var. microphylla, Portulacaceae, BR/BL, 2022), Opuntia canterae (re-estabilished, Cactaceae, UR, 2020), Rudgea megalocarpa (emancipation, Rubiaceae, CL/EQ, 2021), Palicourea villipila (re-established, outside P. axillaria, Rubiaceae, VZ, 2021), Schizanthus fallax (re-estabilished, Solanaceae, CH, 2021), Lippia rodriguezii (re-established, outside L. asperrima, Verbenaceae, AR, 2022), Stachytarpheta chapadensis (re-established, Verbenaceae, BR, 2020), U. bicolor, U. damazioi, U. hirtella, U. lindmanii, U. roraimensis, U. trinervia, U. velascoensis (re-estabilisheds, Lentibulariaceae, BR[2,3,4,5,6,7], VZ[exact 4], BL[exact 4], CAM [exact 1], 2022), Utricularia trinervia (elevation-rank, Lentibulariaceae, ex-U. amethystina, BLZ/MX/VZ/BR, 2021), Piptolepis pseudomyrtus (re-estabilished outside P. buxoides, Asteraceae, BR, 2022), Hydrocotyle alpina, H. palacea, H. quinqueradiata (elevation-rank, Araliaceae, BR, 2022).

∎ rediscovered species: Bauhinia conceptionis (Fabaceae, CL, 2021), Ayenia glabrescens (Malvaceae, BR, 2021), Ternstroemia killipiana (Pentaphyllacaceae, CL, 2021), Varronia neowediana (Cordiaceae, BR, 2022).

∎ simple-species renames: Mespilodaphne sp. (Lauraceae, ex-Ocotea ligulata, BR, 2021), Guettarda [Tournefortiopsis] sp. (Rubiaceae, ex-Chomelia torrana, GF, 2022), Benzingia chocoensis (Orchidaceae, ex-Chondrorhyncha chocoensis, CL, 2019), Ixyophora velastiguii (Orchidaceae, ex-Chondrorhyncha velastiguii, CL/EC, 2019), Madisonia sp. (Orchidaceae, ex-Specklinia ordinata, EC, 2023), Clematicissus pruinata (Vitaceae, ex-C. simsiana, C South America, 2021), Pseudosamanea carbonaria (Fabaceae, ex-Albizia carbonaria, CAM to PR, 2022), Cerradicola praeandina (Fabaceae, ex-Camptosema praeandinum, BL/AR, 2021), Inga inundata (Fabaceae, ex-Zygia inundata, BR, 2018), Oxalis cipoensis (Oxalidaceae, ex-Oxalis calcicola, BR, 2021), Eugenia rogersiana (Myrtaceae, ex-Calycorectes rogersianus, BR, 2021), Eriotheca crenulata (Malvaceae, BR, ex-Bombax crenulata, 2021), Lisymachia neolongipedicellata (renamed, Primulaceae, ex-Lysimachia longipedicellata, BR, 2022), Arenaria weberbaueri (Caryophyllaceae, ex-Alsine weberbaueri, PR, 2023), Arenaria humboldtiana (Caryophyllaceae, ex-Scleranthus peruvianus, PR, 2023), Oxybasis frigida (Amaranthaceae, ex-Chenopodium frigidum, Cono Sur, 2022), Oxybasis parodii (Amaranthaceae, ex-Chenopodium parodii, Cono Sur, 2022), Gentianella gynophora (rename, Gentianaceae, ex-G. fiebrigii, AR, 2022), Symphyllophyton sp. 5 (rename, Gentianaceae, ex-Schultesia sucreana, BR, 2021), Carapichea galbaoensis (Rubiaceae, ex Psychotria galbaoensis, BR/GF, 2022), Fabiana patagonica (Solanaceae, ex-Petunia patagonica, AR, 2022), Justicia comata (Acanthaceae, ex-Spigelia beccabungoides, UR, 2001), Thyrsacanthus angustissimus (Acanthaceae, ex-Justicia angustissima, BR, 2022), Rhabdocaulon grandiflorum (Lamiaceae, ex-Hoehnea grandiflora, BR, 2022), Thyrsacanthus ramosus (Acanthaceae, ex-Anisacanthus pohlii, BR, 2022), Justicia triloba (Acanthaceae, ex-Anisacanthus trilobus, BR, ?), Hyptis piranii (Lamiaceae, ex-Oocephalus piranii, BR, 2022), Hypenia sp. (Lamiaceae, ex-Eriope machrisae, BR, 2021), Cantinoa lythroides (Lamiaceae, ex-Oocephalus lythroides, BR, 2022), Stachytarpheta mollis (Verbenaceae, ex-Stachytarpheta dawsonii, BR, 2021), Erechtites albiflorus (Asteraceae, tropical South America, ex-Erechtites petiolatus, 2021), Wedelia monantha (Asteraceae, ex-Aster monanthus, BR, 2022), Conocliniopsis grossedentata (Asteraceae, ex-C. prasiifolia, BR, 2022), Monticalia sp. 1 (Asteraceae, ex-Pentacalia chimborazensis, Central Andes, 2021), Monticalia sp. 2 (Asteraceae, ex-Pentacalia aclydiphylla, Central Andes, 2021), Monticalia sp. 3 (Asteraceae, ex-Pentacalia mini-aurita, Central Andes, 2021), Monticalia sp. 4 (Asteraceae, ex-Senecio pleniauritus, Central Andes, 2021).

∎ addition to our list of 867 species absents in VPA South America list in April 3, 2023, being 828 ENDEMICS among countries: Brazil (502), Colombia (118), Peru (72), Ecuador (60), Bolivia (27), Venezuela (17), Chile (12), Guianas (11), Argentina (8), Paraguay (1); and 39 NON ENDEMICS: as BR/VZ(2)⋅ BR/EC(1) ⋅ BR/AR(1) ⋅ BR/BL(1) ⋅ BR/CL (2) ⋅ BR/GU´s (3) ⋅ AR/BL (3) ⋅ AR/UR(1) ⋅ AR/PAR(1) ⋅ CL/PAN(1) ⋅ CL/VZ(1) ⋅ CL/EC(10) ⋅ CL/PR(1) ⋅ PR/BL(1) ⋅ EC/PR (5) ⋅ PR/CH(1) ⋅ CL/EC/PR(1) ⋅ AR/PAR/CH(1) ⋅ BR/BL/PAR(1) ⋅ CL/EC/PR/BR(1): Victoria boliviana (Nymphaeaceae, BL, 2022), Cinnamodendron brasiliense, C. catarinense (Canellacae, BR, 2020), Aristolochia pyriflora (Aristolochiaceae, BR, 2022), Peperomia renzopalmae, P. ricardofernandezii (Piperaceae, PR, 2020), P. arianeae, P. forzzae, P. jaramilloae, P. marquetii, P. martinellii, P. michelineae, P. moreiranum, P. rizzinii, P. sucreanum (Piperaceae, BR, 2022), P. brumadinense, P. cariacicaense, P. moringanum (Piperaceae, BR, 2022), Piper callejasii (Piperaceae, CL/PR, 2021), Piper hoyoscardozii, P. indiwasii, P. nokaidoyitau, P. velae (Piperaceae, CL[1,2,3,4], EC[1,2], PR[2], 2022), Piper pseudopeculiare, P. resinaense (Piperaceae, CL, 2021), Aiouea albopunctata (Lauraceae, BR, 2022), Licaria monsalveae, Ocotea sacculifera (Lauraceae, CL[1], BL[2], 2021), Ocotea mello-silvae (Lauraceae, BR, 2021), Mezilaurus conceicionensis (Lauraceae, BR, 2022), Ocotea bilocellata (Lauraceae, BR, 2022), Persea psammophila (Lauraceae, BR, 2021), Mollinedia ruschii (Monimiacae, BR, 2021), Duguetia leucotricha (Annonaceae, BR, 2023), Xylopia maasiana (Annonaceae, BR, 2021), Xylopia sp. (Annonaceae, PR, 2021), Magnolia clementinana, M. manuensis (Magnoliaceae, PR, 2020), Otoba scottmorii, O. squamosa (Myristicaceae, CL, 2021), Virola calimensis, V. cogolloi, V. excisa, V. tuckerae, V. alvaroperezii, V. bombuscaroensis, V. yasuniana, V. aguarunana, V. cumala, V. parkeri (Myristicaceae, CL[1,2,3,4], EC[1,3,5,6,7], PR[3,8,9,10], BR[3,7], 2022), Anthurium atrovinosum, A. brigadeiroense (Araceae, BR, 2020), Anthurium sp.1 (Araceae, CL, 2022), Anthurium sp.2 (Araceae, CL, 2022), Anthurium alegrense (Araceae, BR, 2021), Anthurium bovinii (Araceae, BR, 2022), Anthurium huaytae (Araceae, PR, 2023), Anthurium caldasii (Araceae, CL, 2022), Anthurium mayoanum (Araceae, BR, 2021), Anthurium sterilispadix (Araceae, BR, 2022), Anthurium alfcardozoi, A. coltovarense, A. galipanense, A. georgetatei, A. guaicaipurense, A. hansonianum, A. ronliesneri (Araceae, VZ, 2021), Chlorospatha minima, C. silverstonei (Araceae, CL, 2019), Monstera sp.1, Monstera sp. 2 (Araceae, CL, 2021), Philodendron rio-pretense (Araceae, BR, 2022), Xanthosoma gratieae (Araceae, EC, 2022), Dioscorea ayardei (Dioscoreaeae, AR, 2021), Dioscorea chusqueifolia (Dioscoreaeae, PR, 2019), Dioscorea magnibracteata (Dioscoreaeae, EC, 2022), Thismia petasiformes (Thismiaceae, BR, 2022), Thismia cordata (Thismiaceae, BR, 2022), Thismia mantiqueirensis (Thismiaceae, BR, 2022), Thismia andicola (Thismiaceae, BR, 2023), Thismia calcarata, T. variabilis (Thismiaceae, BR, 2023) Barbacenia amphirupia, B. maritima (Velloziaceae, BR, 2022), Vellozia inselbergae (Velloziaceae, BR, 2021), Vellozia albohexandra, V. mellosilvae (Velloziaceae, BR, 2022), Hippeastrum lara-ricoi L. (Amaryllidaceae, BL, 2022), Hippeastrum carassense, H. velloziflorum (Amaryllidaceae, BR, 2022), Hippeastrum abatinguara (Amaryllidaceae, BR, 2022), Hippeastrum escoipense (Amaryllidaceae, AR, 2022), Miersia putaendensis (Amaryllidaceae, CH, 2021), Miersia stellata (Amaryllidaceae, CH, 2022), Furcraea abisaii (Asparagaceae, CL, 2020), Alophia graniticola (Iridaceae, BR, 2021), Herbertia guyunusae (Iridaceae, BR, 2022), Sisyrinchium caratuvense, S. iguazuanum, S. usneoides (Iridaceae, BR; BR; BR/AR, 2022), Vanilla andina (Orchidaceae, EC, 2022), Andinia barba-caprina, A. crassipetala (Orchidaceae, PR, 2021), Andinia peruviana (Orchidaceae, PR, 2022), Campylocentrum alvesii (Orchidaceae, CL, 2020), Cattleya mireileana (Orchidaceae, BR, 2022), Cattleya porphyrascens (Orchidaceae, BR, 2023), Clowesia arevaloi (Orchidaceae, EC, 2022), Coryanthes charlesiana (Orchidaceae, BR, 2022), Crossoglossa dapaensis, C. elvirae (Orchidaceae, CL, 2022), Cranichis beckii, C. maldonadoana (Orchidaceae, BL, 2021), Cyrtochilum pollex (Orchidaceae, PR, 2022), Cyrtochilum bockemuehlae (Orchidaceae, CL, 2022), Cyrtochilum guavianum (Orchidaceae, CL, 2022), Cyrtopodium valebellae (Orchidaceae, BL, 2023), Dichaea andina (Orchidaceae, CL, 2021), Epidendrum alejandrinae (Orchidaceae, PR, 2022), Epidendrum calimaense (Orchidaceae, CL, 2022), Epidendrum churubambense, E. unchogense (Orchidaceae, PR, 2022), Epidendrum claustralis (Orchidaceae, PR, 2022), Epidendrum itacolomiensis (Orchidaceae, BR, 2022), Epidendrum sonsonense (Orchidaceae, CL, 2022), Epidendrum olorteguii (Orchidaceae, PR, 2022), Epidendrum dayseae (Orchidaceae, BR, 2022), Epidendrum lufinorum (Orchidaceae, PR, 2021), Epidendrum chrisii-sharoniae (Orchidaceae, PR, 2022), Habenaria longissima (Orchidaceae, BR, 2021), Habenaria abscondita (Orchidaceae, BR, 2022), Habenaria aranifera (Orchidaceae, BR, 2022), Habenaria bryophila, H. hygrophila, H. subrepens, H. compluviosa (Orchidaceae, BR, 2022), Habenaria karstica (Orchidaceae, BR, 2022), Laelia [Schomburkgia] vandenbergiana (Orchidaceae, BR, 2022), Lepanthes attenboroughii (Orchidaceae, CL, 2022), Lepanthes wakemaniae (Orchidaceae, CL, 2022), Lepanthes sylvilagus (Orchidaceae, CL, 2022), Lepanthes florenciana (Orchidaceae, CL, 2022), Lepanthes anchicayae, L. microcalodyction (Orchidaceae, CL, 2020), Lepanthes suelipinii (Orchidaceae, BR, 2019), Lepanthes vere-aurum (Orchidaceae, EC, 2020), Lepanthes kokonuko, L. jucas (Orchidaceae, CL, 2020), Lepanthes oro-lojaensis, L. microprosartima, L. caranqui (Orchidaceae, EC, 2021), Lepanthes cardenasii, L. davidii, L. dorae, L. morae (Orchidaceae, CL, 2022) Lepanthes irmae, L. fimbriatilabia, L. pogonochila (Orchidaceae, CL, 2021), Lepanthes pembertonii, L. hwangiae (Orchidaceae, CL, 2022), Masdevallia bastantei (Orchidaceae, PR, 2021), Masdevallia britoi (Orchidaceae, BR, 2022), Masdevallia purocafeana (Orchidaceae, EC, 2022), Maxillaria anacatalinaportillae (Orchidaceae, EC, 2021), Maxillaria bicentenaria (Orchidaceae, PR, 2021), Maxillaria luizotavioi (Orchidaceae, BR, 2023), Microchilus dasilvae (Orchidaceae, BR, 2022), Mormodes matogrossensis (Orchidaceae, BR, 2020), Myoxanthus oliviae (Orchidaceae, PR, 2019), Octomeria pacii, O. panguiensis (Orchidaceae, EC, 2021), Octomeria imigiae (Orchidaceae, BR, 2022), Opilionanthe magdalenae (Orchidaceae, PR, 2021), Pelexia [Pachygenium] laurense (Orchidaceae, AR, 2022), Palmorchis triquilhada (Orchidaceae, BR, 2020), Phragmipedium cabrejosii (Orchidaceae, PR, 2019), Pityphyllum [Maxillaria] mercedes-abarcae (Orchidaceae, EC, 2021), Platystele finleyae (Orchidaceae, EC, 2022), Platystele peruviana (Orchidaceae, PR, 2022), Pleurothallis sp. (Orchidaceae, EC, 2021), Pleurothallis villahermosae (Orchidaceae, CL, 2022), Pleurothallis ariana-dayanae (Orchidaceae, EC, 2022), Pleurothallis mark-wilsonii (Orchidaceae, CL, 2022), Pseudolepanthes bihuae (Orchidaceae, EC, 2021), Stelis excentrica (Orchidaceae, CL, 2020), Telipogon sp. (Orchidaceae, EC, 2021), Telipogon crisariasae (Orchidaceae, EC, 2022), Trisetella pachycaudata (Orchidaceae, EC, 2021), Dichorisandra rhizantha (Commelinaceae, BR, 2022), Dichorisandra rigidiflora (Commelinaceae, BR, 2022), Mauritiella disticha (Arecaceae, BR, 2021), Butia buenopolensis (Arecaceae, BR, 2021), Alcantarea alegrensis(Bromeliaceae, BR, 2021), Alcantarea chimera(Bromeliaceae, BR, 2022), Cryptanthus apiculatantherus, C. brevibracteatus (Bromeliaceae, BR, 2021), Cryptanthus flesheri, C. lutandensis, C. santosii, C. solidadeanus (Bromeliaceae, BR, 2021), Cryptanthus euglossinii (Bromeliaceae, BR, 2022), Cryptanthus pirambuensis, C. vinosibracteatus (Bromeliaceae, BR, 2022), Dyckia avacanoeira, D. oreadica (Bromeliaceae, BR, 2022), Dyckia pseudodelicata (Bromeliaceae, BR, 2022), Hohenbergia amargosensis (Bromeliaceae, BR, 2022, unrecognized in WCSPF), Hohenbergia ymboreorum (Bromeliaceae, BR, 2022), Hohenbergia nidularioides (Bromeliaceae, BR, 2021), Pitcairnia mineira (Bromeliaceae, BR, 2022), Puya pendula (Bromeliaceae, CL, 2021), Ronnbergia igneosepala, R. robusta, R. veitchioides (Bromeliaceae, CL, 2021), Stigmatodon enigmaticus (Bromeliaceae, BR, 2023), Tillandsia mantiqueirae (Bromeliaceae, BR, 2021), Tillandsia oliveirae (Bromeliaceae, BR, 2021), Tillandsia nathanii, T. ertonii (Bromeliaceae, BR, 2022), Waltilia itambana (Bromeliaceae, BR, 2021), Wittmackia conduruensis, W. guedesiae (Bromeliaceae, BR, 2023), Bulbostylis litoreamazonicola (Cyperaceae, BR, 2021), Carex phylloscirpoides (Cyperaceae, CH, 2021), Carex pedicularis (Cyperaceae, AR/PAR/CH, 2021), Carex giovanniana (Cyperaceae, AR/BL, 2021), Cephalocarpus insolitus, C. neblinensis (Cyperaceae, VZ/BR, 2021), Cyperus prophyllatus (Cyperaceae, BR, 2021), Cyperus zikae (Cyperaceae, VZ, 2022), Actinocephalus brevifolius (Eriocaulaceae, BR, ?), Eriocaulon benedictum, E. naviculum (Eriocaulaceae, BR, 2022), Paepalanthus irwinii (Eriocaulaceae, BR, 2022), Paepalalanthus oreodoxus (Eriocaulaceae, BR, 2021), Paepalanthus paganuccii (Eriocaulaceae, BR, 2022), Paepalanthus sinuosus, Sygonanthus culcitosus (Eriocaulaceae, BR, 2021), Xyris serrana (Xyridaceae, BR, 2021), Arthrostylidium cachimboense, Merostachys cachimboensis (Poaceae, BR, 2022), Aristida surperuanensis (Poaceae, PR, 2019), Aristida diego-santiagoii (Poaceae, PR, 2022), Chascolytrum neobulbosum (Poaceae, BR, 2019), Chascolytrum serranum (Poaceae, BR, 2020), Chusquea cordata (Poaceae, BR, 2023), Chusquea recurvata, C. acutigluma (Poaceae, VZ[1], CL[2], 2022), Chusquea calderoniae (Poaceae, BR, 2023), Chusquea gamarrae, C. intipaqariy (Poaceae, PR, 2019), Eremitis aemula (Poaceae, BR, 2022), Eremitis clarkiae, E. vinacea (Poaceae, BR, 2021), Eremitis grandiflora, E. paucifolia, E. victoriae (Poaceae, BR, 2021), Eremitis jardimii (Poaceae, BR, 2020), Merostachys judziewiczii, M. lage-vianae (Poaceae, BR, 2021), Merostachys soderstromii (Poaceae, BR, 2021), Poa bricenoi (Poaceae, VZ, 2021), Pariana caxiuanensis (Poaceae, BR, 2021), Parianella capixaba (Poaceae, BR, 2022), Paspalum plurinerve, P. vacarianum, P. flavescens (Poaceae, BR[1,2], AR/UR[3], 2022), Maranta bahiensis, M. villosovagina, M. chrysogina, M. vieirae, M. lorifolia (Marantaceae, BR, 2021), Maranta pilosissima (Marantaceae, BR, 2022), Meliosma chanchamayensis, M. dazae (Sabiaceae, PR, 2022), Abuta alto-macahensis (Menispermaceae, BR, 2022), Ranunculus oblitus (Ranunculaceae, PR/CH, 2022), Davilla coriacea, D. undulata (Dilleniaceae, BR, 2018), Davilla pygmaea (Dilleniaceae, BR, 2022), Doliocarpus heterophyllus, D. serrulatus (Dilleniaceae, BR, 2022), Connarus foreroi, C. revolutus, C. pedicellatus (Connaraceae, BR[2, 3], PR[1], 2021), Rourea diamantina (Connaraceae, BR, 2020), Sloanea pilosa (Elaeocarpaceae, EC, 2022), Oxalis lourteigiana, O. pampeana (Oxalidaceae, BR, 2021), Oxalis pardoensis (Oxalidaceae, BR, 2022), Celtis serratissima, C. spinosissima,C. clausseniana (Cannabaceae, BR[1, 2, 3], BL[1], PAR[1], 2020), Rhamnidium riograndens (Rhamnaceae, BR, 2021), Lachemilla rothmaleriana, L. argentea, L. cyanea (Rosaceae, CL[1, 2], PR[3], 2019), Margyricarpus lanatus, M. microcarpus (Rosaceae, BR, 2021), Rubus longistipularis, R. maquipucunensis (Rosaceae, EC, 2021), Pilea bradei (Urticaceae, BR, 2023), Adesmia subtropicalis (Fabaceae, BR, 2021), Bauhinia arleneae (Fabaceae, BR, 2022), Bauhinia andrade-limae (Fabaceae, BR, 2022), Bauhinia orbiculata (Fabaceae, BR, 2023), Chamaecrista oppositifolia, C. longistyla (Fabaceae, BR;BR/BL, 2019), Chamaecrista almanegra (Fabaceae, CL, 2022), Chamaecrista acicularis (Fabaceae, BR, 2021), Chamaecrista forzzae (Fabaceae, BR, 2022), Chamaecrista sempreviva (Fabaceae, BR, 2022), Copaifera appendiculata (Fabaceae, BR, 2022), Collaea insignis (Fabaceae, BR, 2022), Deguelia tenuiflora (Fabaceae, BR, 2021), Dipteryx hermetopascoaliana (Fabaceae, BR, 2022), Heterostemon amoris (Fabaceae, CL, 2021), Inga luschnathiana (Fabaceae, BR, 2022), Inga coleyana (Fabaceae, EC, 2022), Inga teresensis, I. tripa (Fabaceae, BR, 2021), Macroptilium albidum (Fabaceae, BR, 2022), Mimosa pseudoracemosa, M. detonsa (Fabaceae, BR, 2021), Mimosa afranioi, M. emaensis, M. robsonii, M. sevilhae (Fabaceae, BR, 2021), Mimosa cavalcantina, M. gustavoi, M. venosa (Fabaceae, BR, 2022), Mimosa brevicalyx (Fabaceae, BR, 2021), Nissolia rondoniensis (Fabaceae, BR, 2021), Nissolia bracteosa (Fabaceae, BR, 2021), Ormosia corcovada (Fabaceae, CL, 2022), Poepiggia densiflora (Fabaceae, BR, 2021), Stryphnodendron flavotomentosum (Fabaceae, BR, 2021), Senna bahiensis (Fabaceae, BR, 2021), Senna pluribracteata (Fabaceae, BR, 2021), Stylosanthes acicularis (Fabaceae, BR, 2023), Asemeia aguiariana, A. campestris, A. eglandulosa, A. subaphylla (Polygalaceae, BR, 2021), Senega [Polygala] bringelii, Senega [P.] tocantinensis (Polygalaceae, BR, 2021), Senega [Polygala capitolensis], Senega [P.] jardimii, Senega [P.] paganuccii (Polygalaceae, BR, 2021), Senega [Polygala] michelliana (Polygalaceae, BR, 2022), Senega [Polygala] petricola (Polygalaceae, BR, 2022), Senega [Polygala] payuniensis, Senega [P.] nevadensis (Polygalaceae, AR, 2022), Erythroxylum sp. 1 (Erythroxylaceae, PAN/CL, 2021), Erythroxylum savannarum (Erythroxylaceae, VZ/CL, 2022), Clusia nascimentojuniorii (Clusiaceae, BR, 2021), Garcinia fluviatilis (Clusicaeae, BR/CL, 2022), Garcinia apostoloi (Clusiaceae, BR, 2023), Tovomita cornuta (Clusiaceae, BR, 2021), Tovomita manauara (Clusiaceae, BR, 2022), Tovomita maxima, T. saulensis (Clusiaceae, BR[1], GU[1], GF[1,2], 2022), Ouratea yamamotoana (Ochnaceae, BR, 2022), Lacistema ligiae (Lacistemaceae, BR, 2021), Bunchosia andina, B. parrae, B. phaeocarpa, Byrsonima anisophylla, B. cardenasii, B. goiana, B. nana, Tetrapterys catarinensis (Malpighiaceae, BR[1,5,9], CL[2,6,7,8], EC[2,4], PR[3,4], 2020), Heteropterys rosmarinifolia, H. tocantinensis, H. walteri, H. veadeirensis (Malpighiaceae, BR, 2021), Phyllanthus dardanoi (Phyllanthaceae, BR, 2021), Phyllanthus dracaenoides (Phyllanthaceae, BR, 2021), Phyllanthus chapadensis (Phyllanthaceae, BR, 2022), Phyllanthus lilliputianus, P. sobralii (Phyllanthaceae, BR, 2022), Phyllanthus pterocaulis (Phyllanthaceae, BR, 2022), Phyllanthus longipetiolatus (Phyllanthaceae, BR, 2022), Casearia valenciana (Salicaceae, BR, 2022), Tetrathylacium vraem (Salicaceae, PR, 2022), Pombalia cristalina, P. insignis (Violaceae, BR, 2022), Viola ornata, Viola longibracteolata (Violaceae, PR, 2022), Viola marcelaferreyrae (Violaceae, AR, 2022), Passiflora acreana (Passifloraceae, BR, 2023), Passiflora bacabensis (Passifloraceae, BR, 2021), Passiflora calicicalyx (Passifloraceae, BL, 2022), Passiflora caparaoensis (Passifloraceae, BR, 2022), Passiflora carajasensis (Passifloraceae, BR, 2021), Passiflora coelestis (Passifloraceae, BR, 2022), Passiflora jorgeana (Passifloraceae, BR, 2021), Passiflora guayaquilensis (Passifloraceae, EC, 2022), Passiflora ketura (Passifloraceae, PR, 2022), Passiflora purii (Passifloraceae, BR, 2022), Passiflora mistratensis (Passifloraceae, CL, 2022), Turnera macrosperma (Passifloraceae, BR, 2020), Bahiana pyriformis (Euphorbiaceae, BR, 2022), Bahiana occidentalis (Euphorbiaceae, PR, 2023), Bernardia erecta, B. monoica, B. stylosa (Euphorbiaceae, BR, 2021), Caperonia amarumayu, C. maracaibensis (Euphorbiaceae, BL[1], VZ[2], 2021), Cnidoscolus mcvaughii (Euphorbiaceae, BR, 2021), Croton almadinensis, C. amaraliae, C. cuadrosii, C. curculiospermus, C. huamaliensis, C. kallunkiae, C. pluriglandulosus (Euphorbiaceae, BR[1,2,6,7], CL[3], BL[4], PR[5], 2021), Croton bacupariensis (Euphorbiaceae, BR, 2021), Croton guaritensis (Euphorbiaceae, BR, 2022), Croton sertanejus (Euphorbiaceae, BR, 2022), Croton stellatorotatus, C. hatschbachii (Euphorbiaceae, BR, 2022), Croton seccoi (Euphorbiaceae, BR, 2021), Euphorbia berryi V.W.Steinm (Euphorbiaceae, BR, 2022), Euphorbia blepharadena, E. longipedunculata, E. sobolifera (Euphorbiaceae, BR, 2021), Mabea dalyana (Euphorbiaceae, BR, 2022), Begonia altimontana, B. inumbrata, B. pedrabrancencis (Begoniaceae, BR, 2022), Begonia atlantica, Begonia umbrosa (Begoniaceae, BR, 2022), Begonia castrosouzae (Begoniaceae, BR, 2022), Begonia diegoi, B. galeanoi, B. mamapachensis, B. perijaensis, B. vinagrera (Begoniaceae, CL, 2021), Begonia embera (Begoniaceae, CL, 2022), Begonia galea (Begoniaceae, BL, 2022), Begonia piranga (Begoniaceae, BR, 2021), Begonia pseudodendron (Begoniaceae, CL/EC, 2022), Bixa atlantica (Bixaceae, BR, 2022), Apeiba trombetensis (Malvaceae, BR, 2012), Ayenia albiflora (Malvaceae, BR, 2022), Callianthe capixabae (Malvaceae, BR, 2022), Eriotheca alversonii (Malvaceae, BR, 2020), Hibiscus marioniae (Malvaceae, GU, 2022), Pseudobombax furadense (Malvaceae, BR, 2022), Quararibea alversonii (Malvaceae, BR, 2022), Quararibea bovinii (Malvaceae, BR, 2021), Sida nordestinensis (Malvaceae, BR, 2020), Triumfetta decaglandulata (Malvaceae, BR, 2022), Waltheria marielleae (Malvaceae, BR, 2022), Cuphea araguaiaensis, C. auriflora, C. praetermissa (Lythraceae, BR, 2023), Cuphea dryadica, C. venosa (Lythraceae, BR, 2022), Vochysia carol-scottii (Vochysiaceae, CL, 2021), Vochysia pongo-qonecensis (Vochysiaceae, PR, 2022), Vochysia sobralii (Vochysiaceae, BR, 2022), Eugenia aeterna (Myrtaceae, BR, 2023), Eugenia bragae (Myrtaceae, BR, 2022), Eugenia cabofriana, E. longimitra, E. pachypoda (Myrtaceae, BR, 2022), Eugenia canumana, E. humaitana, E. jutai, E. multilocellata, E. seislagoana, E. xinguana (Myrtaceae, BR, 2022), Eugenia quilombola (Myrtaceae, BR, 2022), Eugenia buenaventurensis, E. cherimolioides, E. gloriae, E. linaresii, E. melocactoides, E. vallecaucana, Myrcia cabrerae, M. chocoensis, M. glaucocarpa, M. ramirezii, M. samanensis, M. sancarlosiana, M. toledoana (Myrtaceae, CL, 2021), Eugenia nordestina (Myrtaceae, BR, 2022), Eugenia paranapanemensis (Myrtaceae, BR, 2022), Eugenia reperta (Myrtaceae, BR, 2022), Eugenia saxatilis (Myrtaceae, BR, 2021), Eugenia delicata, E. superba (Myrtaceae, BR, 2023), Myrcia longipetiolata (Myrtaceae, BR, 2021), Myrcia chrysotrichoma, M. ribeirensis (Myrtaceae, BR, 2021), Myrcia garuvana (Myrtaceae, BR, 2022), Myrcia stellaris (Myrtaceae, BR, 2021), Myrcia psamophila (Myrtaceae, BR, 2019), Plinia vilabela (Myrtaceae, BR, 2021), Acisanthera saxatilis (Melastomataceae, BR, 2022), Allomaieta pterocalycina (Melastomataceae, CL, 2021), Alloneuron trinervium (Melastomataceae, CL, 2022), Boyania kenwurdackii (Melastomataceae, GU, 2022), Chaetogastra cogniauxiana (Melastomataceae, BR, 2021), Comolia abaetensis (Melastomataceae, BR, 2021), Macrocentrum aurimontium (Melastomataceae, BR, 2021), Marcetia auricularia, M. santosiae, M. unguiculata (Melastomataceae, BR, ?), Meriania bicentenaria, M. vasquezii, M. bongarana, M. callosa, M. juanjil, M. hirsuta, M. megaphylla, M. sumatika, M. escalerensis (Melastomataceae, PR, 2022), Miconia florbella, M. valenzuelana (Melastomataceae, PR, 2021), Miconia waimiri-atroari (Melastomataceae, BR, 2021), Miconia lucenae (Melastomataceae, BR, 2020), Miconia quartzicola, M. spiritusanctensis (Melastomataceae, BR, 2022), Microlicia ascendens, M. barbata, M. daneui, M. piatensis, M. prostrata, M. tetramera (Melastomataceae, BR, 2022), Microlicia barretoana, M. longifolia (Melastomataceae, BR, 2021), Microlicia bicolor, M. pataroi (Melastomataceae, BR, 2022), Microlicia coronata (Melastomataceae, BR, 2022), Microlicia decumbens, M. macaubensis (Melastomataceae, BR, 2022), Microlicia johnwurdackiana, M.deflexa (Melastomataceae, BR, 2021), Microlicia gracilis, M. xylopodifera (Melastomataceae, BR, 2020), Microlicia gertii, M. purpurata, M. trianae (Melastomataceae, BR, 2021), Microlicia angelana, M. debilis (Melastomataceae, BR, 2022), Microlicia piauiensis (Melastomataceae, BR, 2022), Microlicia schwackeana (Melastomataceae, BR, 2022), Microlicia woodgyeriana (Melastomataceae, BR, 2021), Microlicia windschii (Melastomataceae, BR, 2020), Microlicia joaosemiriana, M. longiglandulosa, M. jolyana (Melastomataceae, BR, 2021), Pleroma brevicomosum, P. caetanoi, P. miconiifolium, P. petrophylax (Melastomataceae, BR, 2022), Pleroma joelsilvae (Melastomataceae, BR, 2022), Pleroma piranii (Melastomataceae, BR, 2022), Pleroma barbellatum (Melastomataceae, BR, 2022), Pleroma congestifolium, P. martinellii (Melastomataceae, BR, 2023), Tryssophyton quadrifolius (Melastomataceae, GU, 2019), Astronium woodii (Sapindaceae, BL, 2021), Schinus congestiflora, S. obliqua, S. tarijensis, S. villosa (Anacardiaceae, BL, 2022), Trattinnickia zickeliana, T. nebulae, T. dalyana (Meliaceae, BR, 2021), Ruagea obovata, R. beckii, R. parvifructa (Meliaceae, EC[3], PR[1,3], BL[1,2], 2021), Amyris pacis (Rutaceae, CL, 2022), Conchocarpus kallunkiae (Rutaceae, CL, 2021), Helietta hirsuta (Rutaceae, CL, 2023), Lubaria heterophylla (Rutaceae, CL, 2021), Spiranthera sp. (Rutaceae, PR, 2022), Zanthoxylum complexum, Z. kallunkiae, Z. pluvimontanum (Rutaceae, EC, 2022), Thinouia cazumbensis (Sapindaceae, BR, 2020), Capparidastrum alboannulatum (Capparaceae, CL, 2020), Eudema [Brayopsis] arequipa (Brassicaceae, PR, 2022), Vascocellea badilloi (Caricaceae, PR, 2020), Guapira fundacionensis, G. guasarensis (Nyctaginaceae, VZ, 2022), Neea campanulata (Nyctaginaceae, BR, 2021), Neea gustaviaefolia (Nyctaginaceae, CL, 2021), Coccoloba gigantifolia (Polygonaceae, BR, 2019), Arthrocereus grandiflorus (Cactaceae, BR, 2021), Cumulopuntia flexibilispina (Cactaceae, BL, 2021), Melocactus pachycephalus, M. alex-bragae (Cactaceae, BR, 2021), Caiophora vallegrandensis (Loasaceae, AR, 2022), Xylopodia laurensis (Loasaceae, AR/BL, 2022), Loasa chrysantha (Loasaceae, CH, 2022), Cathedra rupestris (Aptandraceae, BR, 2021), Heisteria longipedicellata (Erythropalaceae, BR, 2021), Aetanthus alternifolius (Loranthaceae, CL, 2021), Dendrophthora kuijtiana (Santalaceae, VZ, 2022), Lissocarpa bracki (Ebenacaee, PR, 2020), Eschweilera podoaquilae (Lecythidacae, EC, 2023), Eschweilera brevipetiolata (Lecythidacae, CL, 2023), Ternstroemia bahiensis, T. rupestris, T. megaphylla, T. longipetiolata (Pentaphyllacaceae, BR, 2021), Lysimachia abscondita, L. campestris, L. catharinensis, L. longipedicellata, L. parvula (Primulaceae, BR, 2021), Chromolucuma brevipedicellata (Sapotaceae, BR, 2021), Chrysophyllum lancisepalum (Sapotaceae, BR, 2021), Laplacea plicata (Theaceae, EC, 2022), Agarista revolutissima (Ericaceae, BR, 2022), Plutarchia dolos (Ericaceae, CL, 2019), Psammisia murriensis (Ericaceae, CL, 2022), Themistoclesia diminuta (Ericaceae, CL, 2022), Johnstonella punensis (Boraginaceae, CH, 2022), Keraunea bullata , K. confusa , K. velutina (Ehretiaceae, BR, 2023), Cordia obtusiloba (Cordiaceae, BR, 2021), Varronia minensis (Cordiaceae, BR, 2022), Euploca decorticans (Heliotropiaceae, BR, 2022), Euploca riochiquensis (Heliotropiaceae, BL, 2022), Myriopus elinae (Heliotropiaceae, BR, 2022), Borreria kelleri (as Spermacocce here, Rubiaceae, AR/PAR, 2022), Borreria savannicola (Rubiaceae, BR, 2022), Galianthe holmneielsenii (Rubiaceae, CL/EQ, 2021), Hippotis antioquiana, H. ecuatoriana, H. elegantula, H. vasqueziana, Schradera cernua, S. francoae, S. condorica, S. morindoides (Rubiaceae, CL[1,5,6], EC[2,3,7,8], PR[4,8], 2019), Mitracarpus semirianus (Rubiaceae, BR, 2022), Paganuccia icatuensis (Rubiaceae, BR, 2021), Palicourea aristata, P. quibdoana, P. santanderiana, P. winfriedii(Rubiaceae, VZ[1,5], EC[2], CL[3,4], 2021), Psyllocarpus itakangapyra, P. vianae (Rubiaceae, BR, 2022), Rudega approuaguensis, R. glomerulata, R. graniticola, R. itoupensis, R. jadinii, R. leucocarpa (Rubiceae, GF[1, 2, 3, 4, 5, 6], SR[3, 6], BR[6], 2022), Rudgea barbosae, R. campanana, R. cardenasii, R. chocoana, R. elegans, R. homeieri, R. inflata, R. retiniphylloides, R. sanluisensis, R. suberosa, R. zappiae (Rubiaceae, CL[1,3,4,7,8,9], EC[4,6,10,11], BR[7], PR[5,10], 2021), Rudgea billietiae (Rubiaceae, GF, 2022), Rudgea infundibuliformis (Rubiaceae, BR, 2022), Staelia schumannii (Rubiaceae, BR, 2021), Tournefortiopsis [G.] crassifolia, T. [G.] deviana, T. [G] robusta , T. [G] sopkinii, T. [G] tamboana (Rubiaceae, CL, 2021), Aspidosperma dardanoanum (Apocynaceae, BR, 2021), Forsteronia manausana, F. nitida, F. paraensis, F. prancei (Apocynaceae, BR;BR;BR/SR;BR, 2019), Gonolobus rumihuilcanus (Apocynaceae, EC, 2022), Hemipogon trilobatus (Apocynaceae, BR, 2020), Mandevilla bullata (Apocynaceae, BR, 2020), Mandevilla declinata, M. fornograndensis, M. mysteriosa, M. obovata (Apocynaceae, BR, 2022), Oxypetalum kassneri (Apocynaceae, BR, 2021), Oxypetalum timbense (Apocynaceae, BR, 2022), Petalostelma atlanticum (Apocynaceae, BR, 2021), Philibertia woodii (Apocynaceae, BL, 2021), Ruehssia quirinopolensis (Apocynaceae, BR, 2021), Symphyllophyton sp.1, S. sp.2, S. sp.3, S. sp.4 (Gentianaceae, BR, 2021), Voyria bicolor (Gentianaceae, GF, 2022), Bonamia eustachioi (Convolvulaceae, BR, 2021), Dicranostyles yrypoana (Convolvulaceae, BR, 2023), Evolvulus aureus (Convolvulaceae, BR, 2023), Evolvulus longipedicellatus (Convolvulaceae, BR, 2022), Evolvulus saxatilis (Convolvulaceae, BR, 2020), Evolvulus veadeirensis (Convolvulaceae, BR, 2022), Cuscuta mantiqueirana (Convolvulaceae, BR, 2021), Ipomoea bonsai (Convolvulaceae, BR, 2020), Ipomoea lanifolia (Convolvulaceae, BR, 2021), Jacquemontia atlantica (Convolvulaceae, BR, 2022), Jacquemontia boliviana, J. chuquisacensis, J. cuspidata, J. longipedunculata, J. mairae (Lauraceae, BL, 2022), Athenaea altoserranae, A. hunzikeriana (Solanaceae, BR, 2021), Calibrachoa atropurpurea, C. synanthera (Solanaceae, BR, 2022), Doselia galilensis (Solanaceae, CL, 2022), Lycianthes amazonica (Solanaceae, BR, 2021), Nicotiana rupicola (Solanaceae, CH, 2022), Nicotiana gandarela (Solanaceae, BR, 2022), Schizanthus nutantiflorus (Solanaceae, CH, 2021), Solanum bohsii (Solanaceae, CL, 2022), Anemopaegma kawense (Bignoniaceae, GF, 2021), Handroanthus abayoy (Bignoniaceae, BL, 2022), Calceolaria flavida Lavandero (Calceolariaceae, CH, 2021), Besleria naquenensis (Gesneriaceae, CL, ?), Besleria azulensis, B. vanderwerffii (Gesneriaceae, PR, 2021), Besleria sp. (Gesneriaceae, CL, 2021), Columnea angulata, C. floribunda, C. tecta (Gesneriaceae, EC[1, 2, 3], CL[3], 2021), Columnea fluidifolia, C. pendens (Gesneriaceae, EC/CL, 2022), Drymonia intermedia, D. longiflora (Gesneriaceae, EC[1,2], CL[2], 2022), Drymonia peponifera (Gesneriaceae, EC, 2022), Glossoloma wiehleri (Gesneriaceae, EC, 2021), Glossoloma magenticristatum (Gesneriaceae, CL, 2023), Monopyle glutinosa (Gesneriaceae, EC, 2022), Paradrymonia vivianensis (Gesneriaceae, PR, 2021), Pearcea lutea (Gesneriaceae, EC, 2022), Sinningia ganevii, S. sulphurea (Gesneriaceae, BR, 2022), Cyanocephalus glocimarii (Lamiaceae, BR, 2021), Eriope orlandoi (Lamiaceae, BR, 2023), Hyptidendron cerradoense (Lamiaceae, BR, 2022), Hyptidendron pulcherrimum (Lamiaceae, BR, 2021), Hyptis melittiflora (Lamiaceae, BR, 2021), Hyptis spathulata Harley, H. cachimboensis (Lamiaceae, BR, 2021), Hyptis pastorei (Lamiaceae, BR, 2019), Oocephalus rhodocalyx (Lamiaceae, BR, 2021), Oocephalus viscaria (Lamiaceae, BR, 2019), Oocephalus griseus, O. campestris (Lamiaceae, BR, 2022), Salvia celendina (Lamiaceae, PR, 2021), Vitex pomerana (Lamiaceae, BR, 2022), Ameroglossum xukuruorum, A. asperifolium, A. intermedium, A. bicolor, A. fulniorum, A. alatum, A. genaroanum (Linderniaceae, BR, 2021), Agalinis marianae (Orobanchaceae, BR, 2022), Bacopa llanorum (Plantaginaceae, VZ, 2020), Plantago campestris (Plantaginaceae, BR, 2021), Lippia crucifera (Verbenaceae, BR, 2021), Lippia raoniana (Verbenaceae, BR, 2021), Stachytarpheta brevibracteata, S. longipedicellata, S. minasensis, S. ratteri (Verbenaceae, BR, 2022), Stachytarpheta flavovirescens, S. salimenae (Verbenaceae, BR, 2021), Stachytarpheta olearyana, S. vianae (Verbenaceae, BR, 2022), Dyschoriste eulinae, D. vinacea (Acanthaceae, BR, 2022), Harpochilus corrugatus (Acanthaceae, BR, 2022), Justicia montealegrensis, J. multiglandulosa, J. paraensis (Acanthaceae, BR, 2021), Justicia pusilla (Acanthaceae, BR, 2021), Justicia espiritosantensis (Acanthaceae, BR, 2022), Odontonema peruvianum (Acanthaceae, PR, 2019), Ruellia curupira, R. fulozinha, R. insurrecta, R. jiboia, R. taboleirana (Acanthaceae, BR, 2020), Ruellia umbrosa (Acanthaceae, BR, 2022), Ruellia whitneyana (Acanthaceae, BL, 2022), Sanchezia dubia (Acanthaceae, PR, 2021), Stenostephanus atrocalyx, S. densiflorus (Acanthaceae, PR, 2019), Stenostephanus borarum, S. brevistamineus(Acanthaceae, PR, 2022), Pinguicula chuquisacensis (Lentibulariaceae, BL, 2008), Utricularia amotape-huancabambensis (Lentibulariaceae, PR, 2021), Utricularia ariramba, U. jaramacaru (Lentibulariaceae, BR, 2020), Utricularia chapadensis, U. lunaris, U. pantaneira (Lentibulariaceae, BR, 2022), Burmeistera chrysothrix, B. crocodila, B. erosa, B. lingulata, B. sierrazulensis, B. valdiviana (Campanulaceae, EC, 2021), Siphocampylus flavescens (Campanulaceae, BR, 2022), Calea arachnoidea (Asteraceae, BR, 2021), Calea funkiana (Asteraceae, BR, 2021), Calea pruskiana, C. subintegerrima (Asteraceae, BR, 2022), Calea repanda (Asteraceae, BR, 2022), Calea sessilifolia (Asteraceae, BR, 2022), Calea grandiflora (Asteraceae, BR, 2023), Cololobus ruschianus (Asteraceae, BR, 2021), Diplostephium paposanum (Asteraceae, CH, 2022), Eremanthus tomentosus, Lychnophora goiana, L. planaltina (Asteraceae, BR, 2022), Espeletia ocetana (Asteraceae, CL, 2021), Espeletia saboyana (Asteraceae, CL, 2022), Heterocondylus penninervius (Asteraceae, BR, 2021), Heterocypsela brachylepis (Asteraceae, BR, 2020), Leucheria cantillanensis (Asteraceae, CH, 2020), Lepidaploa restingae (Asteraceae, BR, 2021), Lepidaploa campirupestris, Lessingianthus petraeus (Asteraceae, BR, 2021), Lychnocephalus cipoensis, L. grazielae, L. jolyanus (Asteraceae, BR, 2021), Lychnophora osanyiniana (Asteraceae, BR, 2021), Lychnophora pseudovillosissima (Asteraceae, BR, 2022), Lychnocephalus canus (Asteraceae, BR, 2022), Mikania mellosilvae (Asteraceae, BR, 2021), Mikania semirii, M. funkiae (Asteraceae, BR 2022), Pectis loiolae (Asteraceae, BR, 2021), Pentacalia atrovinosa (Asteraceae, EC, 2023), Piptolepis rosmariniifolia (Asteraceae, BR, 2019), Piptolepis corymbosa, P. pilosa, P. procumbens (Asteraceae, BR, 2021), Porophyllum woodii (Asteraceae, BL, 2020), Praxelis scaturicola (Asteraceae, BR, 2021), Rockhausenia [Werneria] huascarana, Rockhausenia [W.] rockhauseniana (Asteraceae, PR, 2020), Rockhausenia [Werneria] praetermissa (Asteraceae, AR/BL, 2021), Senecio sp. 1, Senecio sp. 2 (Asteraceae, PR, 2021), Senecio ephemerus (Asteraceae, CH, 2022), Senecio scapioides (Asteraceae, CL, 2020), Valeriana praecipitis (Caprifoliaceae, CH, 2022), Valeriana plateadensis, V. yacuriensis, V. xenophylloides (Caprifoliaceae, EC, 2023), Sciodaphyllum basiorevolutum, S. chachapoyense, S. rufilanceolatum (Araliaceae, EC; PR; EC/CL, 2021), S. montanum, S. undulatum (Araliaceae, CL, 2021), S. merinoi, S. purocafeanum, S. recaldiorum, S. zunacense (Araliaceae, EC, 2021), Eryngium absconditum (Apiaceae, PAR, 2022).

∎ exclusion from our list of occurrence in the New World of these species or genera: Typhonium blumei (by POWO), Cocus nucifera (by POWO), Tribulus (Zygophyllaceae, T. longipetalus by Flora of Pakistan) and T. zeyheri by Flora of Zimbabue), Hydrolea zeylanica (cited for Peru, excluded by Flora of China).