AUSTRALIAN BIOLOGICAL RESOURCES STUDY, CANBERRA SPECIES PLANTARUM FLORA OF THE WORLD. Part 10. CHRYSOBALANACEAE 2 Acioa to Magnistipula

Transcription

1 AUSTRALIAN BIOLOGICAL RESOURCES STUDY, CANBERRA SPECIES PLANTARUM FLORA OF THE WORLD Part 10. CHRYSOBALANACEAE 2 Acioa to Magnistipula by Ghillean T. Prance & Cynthia A. Sothers

2 Commonwealth of Australia 2003 This work is copyright. Apart from any use as permitted under the Copyright Act 1968, no part may be reproduced or distributed by any process or stored in any retrieval system or data base without prior written permission from the copyright holder. Requests and inquiries concerning reproduction and rights should be addressed to the Director, Australian Biological Resources Study, GPO Box 787, Canberra, ACT 2601, Australia. EDITORS Anthony E. Orchard Annette J.G. Wilson ASSISTANT EDITOR Brigitte Kuchlmayr This work may be cited as: G.T.Prance & C.A.Sothers, Chrysobalanaceae 2: Acioa to Magnistipula, Species Plantarum: Flora of the World Part 10: (2003). This book is available from: Australian Biological Resources Study GPO Box 787 Canberra ACT 2601 Australia National Library of Australia Cataloguing in Publication entry Prance, Ghillean. Chrysobalanaceae 2, Acioa to Magnistipula. Bibliography. Includes index. ISBN Chrysobalanaceae. I. Australian Biological Resources Study. II. Title. (Series : Species plantarum: flora of the world ; pt. 10). III. Sothers, Cynthia

3 CONTENTS Contributors iv Species Plantarum Committee v Introduction vii Acioa 1 Couepia 5 Maranthes 56 Atuna 66 Dactyladenia 76 Hirtella 95 sect. Myrmecophila 106 sect. Hirtella 110 Kostermanthus 172 Magnistipula 178 subg. Magnistipula 179 sect. Magnistipula 180 sect. Animalculum 183 sect. Peregrinator 184 subg. Pellegriniella 190 subg. Tolmiella 191 References 194 Appendix 197 Maps 198 Index 262 iii

4 CONTRIBUTORS TO THIS PART Text Ghillean T. Prance, The Herbarium, Royal Botanic Gardens, Kew, Richmond, Surrey TW9 3AB, England. Cynthia A. Sothers, The Herbarium, Royal Botanic Gardens, Kew, Richmond, Surrey TW9 3AB, England. Illustrations Illustrations prepared for this volume were drawn by Margaret Tebbs (Plates 61, 63, 65 67, 74 and 75) and J.M.Fothergill (plate 58), both of The Herbarium, Royal Botanic Gardens, Kew, Richmond, Surrey TW9 3AB, England. Other illustrations in this work have been reproduced, with permission, from a number of sources, which are acknowledged with each plate. iv

5 Species Plantarum Steering Committee (as at November 2003) G. Achoundong, Herbier National de Cameroon, Yaoundé, Cameroon C.E. Anderson, University of Michigan, Ann Arbor, USA A. Anton, Museo Botánico, Universidad Nacional de Córdoba, Argentina W. Berendsohn, Botanischer Garten und Botanisches Museum, Berlin, Germany D.E. Boufford, Harvard University Herbaria, Cambridge, Mass., USA L. Boulos, Dokki, Cairo, Egypt R.K. Brummitt, Herbarium, Royal Botanic Gardens, Kew, UK S. Castroviejo, Real Jardín Botánico, CSIC, Madrid, Spain A.C. Chikuni, National Herbarium and Botanic Gardens of Malawi, Zomba, Malawi M.D. Correa, Universidad de Panamà, Panama T.F. Daniel, California Academy of Sciences, San Francisco, Calif., USA D.Darnaedi, Kebun Raya Indonesia, Bogor, Indonesia P. Dávila, Escuela Nacional de Estudios Profesionales, UNAM, Tlalnepantla, Mexico T.S. Filgueiras, Reserva Ecológica do IBGE, Brasilia, Brazil E. Forero, Universidad Nacional de Colombia, Bogotá, Colombia I. Friis, Botanical Museum and Library, Copenhagen, Denmark V. Funk, Smithsonian Institution, Washington DC, USA D. Geltman, Komarov Botanical Institute, St Petersberg, Russia A.S. George, Kardinya, WA, Australia W. Greuter, Botanischer Garten und Botanisches Museum, Berlin, Germany C. Hilton-Taylor, I.U.C.N., Cambridge, UK Hong D.Y., Chinese Academy of Sciences, Beijing, China K. Iwatsuki, Yokohama, Japan B. Jonsell, Bergius Foundation, Stockholm, Sweden B. Khayota, National Museums of Kenya, Nairobi, Kenya J. Kirschner, Academy of Sciences, Průhonice, Czech Republic H.P. Linder, Insitut für Systematische Botanik, Zurich, Switzerland P.-A. Loizeau, Conservatoire et Jardin Botaniques, Genève, Switzerland J.L. Luteyn, New York Botanic Gardens, Bronx, New York, USA K. Marhold, Slovak Academy of Sciences, Bratislava, Slovakia M. Moraes, Universidad Mayor de San Andres, La Paz, Bolivia P. Morat, Muséum National d'histoire Naturelle, Paris, France N.R. Morin, The Arboretum at Flagstaff, Arizona, USA Nguyen T.H., Institute of Ecology & Biological Resources, Hanoi, Vietnam H. Ohashi, Tohoku University, Sendai, Japan A.E. Orchard, Department of Environment and Heritage, Canberra, ACT, Australia R.J. Pankhurst, Royal Botanic Garden, Edinburgh, UK M. Sanjappa, Botanical Survey of India, Calcutta, India T. Santisuk, Royal Forest Department, Bangkok, Thailand Sebsebe Demissew, Addis Ababa University, Addis Ababa, Ethiopia J.H. Seyani, Commonwealth Secretariat, London, UK G.F. Smith, National Botanical Institute, Pretoria, South Africa W.D. Stevens, Missouri Botanical Garden, Saint Louis, Missouri, USA v

6 W.L. Wagner, Smithsonian Institution, Washington DC, USA A.J.G. Wilson, Australian Biological Resources Study, Canberra, ACT, Australia K. Wilson, Royal Botanic Gardens, Sydney, NSW, Australia vi

7 INTRODUCTION Species Plantarum aims to provide in concise format, and with standardised data fields, basic taxonomic information on the vascular plants of the world, including accepted names and synonyms with bibliographic data, types of names, keys and descriptions from family to varietal levels, geographical distributions, ecological information and other related matters, and to publish it in both hard copy and electronic form. The format of the Species Plantarum is based on that of Flora of Australia, with some departures made necessary by the different scale of the project. Initially at least, the series is being edited and published for the Species Plantarum Project and IOPI by the Australian Biological Resources Study (ABRS), producers of Flora of Australia. Treatments are contributed on a voluntary basis. Each part of Species Plantarum is intended to provide a complete account of a family, subfamily, large genus or other related taxonomic group. While treatments of small families may be shorter, it is intended that contributions will, in general, cover at least 50 to 100 species. The taxonomy adopted is that of the author, although the family delimitations recommended are initially those of R.K.Brummitt, Vascular Plant Families and Genera (1992). The order of taxa within families, genera and species in the Species Plantarum is intended to reflect natural relationships, so far as this is possible in a linear sequence. Maps are provided for each species, or in those cases where infraspecific taxa are recognised, for each of the terminal taxa. Distribution maps are based on those in R.K.Brummitt, World Geographical Scheme for Recording Plant Distributions ed. 2 (2001), and the 'countries' adopted are those of Level 3 and 4 of that work. Description of distribution follows the same work, with a two-digit code for regions and a three-letter code for the 'country'. Upper case letters for the 'country' indicate native distribution; lower case letters indicate that the taxon is only present in that 'country' as an introduced and naturalised plant. If a taxon is extinct in a 'country', this is indicated by a dagger. Distribution of species as cultivated plants is not included. Misapplied and invalid names are, in general, omitted. Journal titles are abbreviated according to G.D.R.Bridson & E.R.Smith, Botanico-Periodicum-Huntianum / Supplementum (1991). Book titles are abbreviated according to F.A.Stafleu & R.S.Cowan, Taxonomic Literature (2nd edn) Vols 1 7, and Supplements (1976 ), except that upper case initial letters are used for proper names and significant words. Authors of plant names are abbreviated according to R.K.Brummitt & C.E.Powell, Authors of Plant Names (1992). A separate part, Introduction to the Series, provides a history of the project, a glossary, guide for contributors and key to the conventions used in describing distribution. These resources will also be available on the World Wide Web, initially through the ABRS site (currently with links from the IOPI site (currently and others. A.E.Orchard & A.J.G.Wilson Canberra November 2003 vii

8 viii

9 11. ACIOA Acioa Aubl., Hist. Pl. Guiane 2: 698, t. 280 (1775); from the vernacular name 'acioua' in French Guiana. Acia Schreb., Gen. Pl. ed. 8, 2: 458 (1791) nom. superfl. illeg. [pro parte excl. syn. Couepia]; Moquilea subg. Acioa (Aubl.) Blume, Mus. Bot. 2: 92 (1856)["1852"]. Type: Acioa guianensis Aubl. Dulacia Neck., Elem. Bot. 2: 414 (1790), nom. inval., non Dulacia Vell. (1823) (Olacaceae). Trees or shrubs. Leaves with stipules small, intrapetiolar; petiole eglandular; lamina coriaceous, with 1 or 2 pairs of conspicuous glands at base and several smaller discoid glands; lower surface glabrous. Inflorescence a rather lax thyrse with flattened axes. Bracteoles minute, persistent, eglandular. Flowers bisexual, strongly zygomorphic. Receptacle tube obconic, slightly curved (cyathiform in A. schultesii). Sepals 5, glabrous outside except for the ciliolate margins, suborbicular, deeply concave, very unequal in size, 3 large, 2 very small; 3 outer sepals with 1 or 2 large discoid glands towards the base. Petals 5, fimbriate. Stamens 10 20, ligulately connate (free in A. edulis), coiled in bud; staminodes absent or represented by minute denticulations on the faucal annulus. Ovary 1-carpellary, 1-locular, inserted laterally at mouth of receptacle tube, glabrous or tomentose; style hairy at base or throughout, very slightly 3-lobed at apex. Drupe (only known in A. edulis and A. guianensis) 8 cm or more long; pericarp poorly differentiated into mesocarp and endocarp, thick, hard and fibrous, glabrous outside and inside, on germination breaking up irregularly. Four species with restricted distributions in lowland tropical rainforest in northern South America. See also individual species below. The genus Dactyladenia from Africa, which also has a staminal ligule, has been included in Acioa by many authors, but it is quite distinct. Germination is phanerocotylar; cotyledons fleshy, without chlorophyll; eophylls with intrapetiolar stipules, the first 2 opposite, otherwise alternate. A.P. de Candolle, Prodr. 2: 526 (1825); H.E.Baillon, Adansonia, 7: 221 (1867) quoad A. guianensis tantum; W.O.Focke, in H.G.A.Engl. & K.A.E.Prantl, Nat. Pflanzenfam. 3(3): 60, t. 33, (1894) quoad relat. spec. Amer. tantum; J.Hutchinson, Gen. Fl. Pl. 1: 192 (1964) excl. syn. Dactyladenia Welw. & Griffonia Hook.f.; G.T.Prance, Fl. Neotrop. Monogr. 9: 357, t. 58 (1972) excl. syn. Dactyladenia & Griffonia; R.Letouzey & F.White, Fl. Cameroun 20 (Fl. Gabon 24): 6, quoad relat. spec. Amer. tantum; G.T.Prance & F.White, The genera of Chrysobalanaceae, Philos. Trans. B320: (1988). 1 Receptacletube obconic, much longer than the sepals, gradually narrowed into the pedicel; sepals tomentellous inside; filaments completely free, or united to form an unbranched staminal ligule 2 Leaves with inconspicuous venation; receptacle tube more than 5 times as long as broad, relatively thin-walled; faucal annulus relatively poorly developed, not blocking throat; filaments united to form a staminal ligule; ovary densely hairy; style hairy for most of length 3 Leaves shortly cuspidate at apex, cuneate or decurrent into petiole at base 2. A. guianensis 3: Leaves rounded or emarginate at apex, rounded at base 4. A. somnolens 2: Leaves with prominently reticulate venation; receptacle tube less than 3 times as long as broad, almost solid; faucal annulus well developed, very thick and fleshy, almost blocking the throat; filaments completely free; ovary glabrous; style hairy only at the base 1. A. edulis 1: Receptacle tube cyathiform, as long as the sepals; sepals glabrous inside; staminal ligule with several branches at base 3. A. schultesii 1

10 SPECIES PLANTARUM FLORA OF THE WORLD (2003) 1. Acioa edulis Prance, Acta Amazon. 2(1): 12 (1972) Couepia edulis (Prance) Prance, Acta Amazon. 5(2): 143 (1975). T: Brazil North: Amazonas, Rio Ituxi, Boca do Curuquetê, G.T.Prance et al ; holo: NY; iso: FHO, INPA, MG, RB, US. Illustrations: G.T.Prance, Acta Amazon. 2(1): 14, 15 (1973); G.T.Prance & F.White, Philos. Trans. B320: 114 (1988). Map: G.T.Prance, Fl. Neotrop. Monogr. 9S: 209 (1989). Tree to 25 m tall. Trunk slightly buttressed to 0.5 m. Branches glabrous when young. Leaves with stipules linear, membranous, 5 8 mm long, glabrous, caducous; petioles cm long, terete, glabrous; lamina oblong, thickly coriaceous, cm, rounded at base, acuminate at apex with acumen 2 6 mm long, glabrous on both surfaces, with 2 glands near to base of lower surface; midrib prominulous above, prominent beneath; primary veins 9 11 pairs, prominent on both surfaces; secondary venation prominent on both surfaces. Inflorescences much-branched slightly corymbose panicles, 5 10 cm long; rachis and branches glabrous. Receptacle conical, slightly curved near base, 6 7 mm long, glabrous on exterior, with pilose hairs on interior below insertion of style and glabrous on other side; interior lined by an extremely thick disc with only a small hollow. Calyx lobes 5, rounded, unequal, 3 5 mm long, glabrous with 2 glands on exterior, with margins ciliate, appressedpuberulous on interior. Petals 5, white, caducous, glabrous, with margins ciliate. Stamens 17 20, inserted in 2 rows around 1 /2 of the thick staminal ring, the other 1 /2 of the ring with staminodes, with a circle of deflexed hairs inserted on interior at base of staminal ring; filaments glabrous, free; anthers dorsifixed. Ovary inserted at mouth of receptacle on the same side as stamens, glabrous on exterior, glabrous within, unilocular; ovules 2. Style inserted at base of ovary towards interior, equalling stamens in length; swollen base pilose; filamentous portion glabrous. Fruit ellipsoid, cm long, 4 5 cm broad; exocarp glabrous, lenticellate; mesocarp mm thick; endocarp thin, bony, fragile, glabrous within; cotyledons completely filling central cavity. Fig. 50A E. Known only from a small area of Amazonas, Brazil, where, however, it is locally abundant. 84: BZN (AM). Forest of non-flooded ground (terra firme). Also cultivated outside its natural range, especially along the Rio Solimões in Amazonas, Brazil. Map BRAZIL NORTH: Amazonas, Coari, Km 6 on road to airport, J.F.Ramos P23251 (BR, INPA, NY, P, W, WAG); Amazonas, Coari, Km 6 on road to airport, D.G.Campbell P21130 (INPA, NY, P); Amazonas, Mun. de Tonantins, Rio Tonantins, Uitinga Paraná, D.C.Daly et al (INPA, K, NY); Amazonas, Rio Ituxi, Boca do Remanzinho, B.A.Krukoff 5822 (NY); Amazonas, Lago de Tefé, Manua, E.Lleras et al. P16655 (INPA, NY). Germination epigeal, first leaves opposite. An oil is extracted from the fruit for use in cooking and soap making. The fleshy cotyledons are edible and the fruit are gathered in large numbers in the Rio Ituxi region. The kernel is eaten raw or is crushed and added to tapioca cakes (beiju) (G.T.Prance, 1989: 97). 2. Acioa guianensis Aubl., Hist. Pl. Guiane 2: 698 (1775) Acia dulcis Willd., Sp. Pl., ed. 4, 3: 717 (1800), nom. superfl. illeg.; Acioa dulcis (Willd.) Steud., Nomencl. Bot., ed. 1, 9 (1821), nom. superfl. illeg.; Moquilea aubletiana Blume, Mus. Bot. 2: 92 (1856) ["1852"]; Licania aubletiana (Blume) Lemée, Fl. Guyane Franç. 2: 23 (1952). T: French Guiana: without locality, J.B.C.F.Aublet; holo: BM. Illustrations: J.B.C.F.Aublet, Hist. Pl. Guiane, Atlas t. 280 (1775); G.T.Prance, Fl. Neotrop. Monogr. 9: 360 (1972); G.T.Prance, Fl. Guianas 85: 4 (1986). Map: G.T.Prance, Fl. Neotrop. Monogr. 9S: 261 (1989). Tree, medium-sized. Leaves with stipules caducous (not seen); petioles 5 12 mm long, mm thick, glabrous, terete; lamina ovate to oblong, cm, cuneate or slightly tapered at base, abruptly short-acuminate at apex, with acumen 2 5 mm long, glabrous on both surfaces; primary veins 7 9 pairs, inconspicuous on both surfaces. Inflorescences terminal and axillary panicles, with rachis and branches glabrous. Receptacle obconictubular, mm long, glabrous on exterior, glabrous within except for deflexed hairs at throat. Calyx lobes 5, pubescent within. Petals 5, with minutely serrate margins. Stamens unilateral, 10 12; filaments ligulately connate. Ovary inserted at mouth of receptacle, 2

11 CHRYSOBALANACEAE (Acioa) Figure 50. Acioa. A E, A. edulis. A, habit; B, seedling; C, stipules; D, flower bud and ½ flower; E, fruit (A, D, J.F.Ramos P23251; B C, G.T.Prance BI13; E, G.T.Prance et al ). F G, A. schultesii. F, young and fully developed flower bud; G, ½ male flower and developing fruit (F G, H.Clark 6740). Scale bars: A, E = 5 cm; B = 4 cm; C, D, F, G = 5 mm. Drawn by Rosemary Wise. Previously published in G.T.Prance & F.White, Philos. Trans. B320: 114 (1988). 3

12 SPECIES PLANTARUM FLORA OF THE WORLD (2003) densely pilose. Style pilose on lower portion. Fruit ellipsoid; epicarp glabrous and sparsely lenticellate; mesocarp to 14 mm thick, fibrous; endocarp thick, hard, merging gradually into mesocarp, with thick fibres, lacking mechanism for seedling escape, glabrous within. French Guiana and Amazonian Brazil. 82: FRG. 84: BZN (AM, AP). Terra firme forest. Map FRENCH GUIANA: without locality, P.A.Poiteau s.n. (P); without locality, G.S.Perrottet s.n. (P). 84. BRAZIL NORTH: Amapá, Rio Araguari, J.M.Pires et al (IAN, NY, P). 3. Acioa schultesii Maguire, Brittonia, 7: 272 (1951) T: Brazil North: Amazonas, Rio Dimiti, Cerro Dimiti, R.E.Schultes & López 9958; holo: NY; iso: K. Illustrations: G.T.Prance, Fl. Neotrop. Monogr. 9: 360 (1972); G.T.Prance, Fl. Venez. Guayana 4: 203 (1998); G.T.Prance & F.White, Philos. Trans. B320: 114 (1988). Map: G.T.Prance, Fl. Neotrop. Monogr. 9S: 261 (1989). Tree, small. Leaves with stipules caducous (not seen); petioles 3 6 mm long, glabrous, thick; lamina broadly elliptic to oblong-elliptic, cm, obtusely cordate at base, rounded to short-acuminate at apex, glabrous on both surfaces; primary veins inconspicuous. Inflorescences axillary and terminal panicles; rachis and branches glabrous. Bracts and bracteoles small; margins ciliolate. Receptacle broadly campanulate, gibbous, glabrous on exterior, glabrous within except for deflexed hairs at throat. Calyx lobes 5, glabrous on both surfaces except for ciliate margins. Petals 5; margins ciliate. Stamens c. 15, unilateral; filaments ligulately connate, with ligules 5 mm thick. Ovary pilose, inserted at mouth of receptacle. Style glabrous except for hirsute base. Fruit ovoid, cm; exocarp glabrous, finely lenticellate. Fig. 50F G. Upper Rio Negro region of Venezuela, Colombia and Brazil. 82: VEN. 83: CLM. 84: BZN (AM). Abundant in forests on white sand (bana and caatinga). Map VENEZUELA: Amazonas, San Carlos de Río Negro, L.M.Berti & P.Salcedo (MO, NY); Amazonas, San Carlos de Río Negro, H.L.Clark & P.Maquirino 6737 (FHO, NY); Amazonas, San Carlos de Río Negro, H.L.Clark & P.Maquirino 6740 (FHO, NY); Amazonas, San Carlos de Río Negro, H.L.Clark & P.Maquirino 6661 (FHO, NY); Amazonas, San Carlos de Río Negro, H.L.Clark 8211 (NY). 4. Acioa somnolens Maguire, Brittonia 7: 272 (1951) T: French Guiana: Cayenne, M.Mélinon 230; holo: P. Map: G.T.Prance, Fl. Neotrop. Monogr. 9S: 261 (1989). Tree? Leaves with stipules caducous (not seen); petioles thick, 8 10 mm long, 4 5 mm broad; lamina broadly elliptic to oblong-elliptic, cm, rounded to slightly retuse at apex, rounded at base, glabrous on both surfaces; primary veins 6 8 pairs, inconspicuous on both surfaces. Inflorescences terminal and axillary panicles, glabrous. Bracts and bracteoles small, caducous. Receptacle obconic-tubular, mm long, glabrous on exterior, glabrous within except for deflexed hairs at throat. Calyx lobes 5, pubescent within. Petals 5, caducous. Stamens c. 15, unilateral; filaments ligulately connate; ligule mm long, 2 3 mm broad. Ovary pilose, inserted at mouth of receptacle. Style sparsely hirsute for 2 /3 its length. Fruit unknown. French Guiana and Brazil. 82: FRG. 84. BZN (AM). Terra firme forest. Map BRAZIL NORTH: Amazonas, Manicoré, C.D.A.Mota s.n. (INPA61659). Very closely related to A. guianensis and possibly not distinct. African species, other than A. goetzeana, which were formerly placed in Acioa, have been transferred to Dactyladenia, and Asiatic species to Kostermanthus. 4

13 CHRYSOBALANACEAE 12. COUEPIA Couepia Aubl., Hist. Pl. Guiane 1: 519 (1775); Acia Schreb., Gen. Pl., ed. 8, 2: 458 (1791), nom. superfl. illeg. The name comes from the vernacular name in French Guiana, 'couepi'. Type: Couepia guianensis Aubl. Couepia sect. Hemicouepia Benth., J. Bot. (Hooker) 2: 212 (1840). T: Not cited. Grymania C.Presl, Epim. Bot. 193 (1851), nom. nud. pro parte quoad G. polyandra tantum. Pleragina Arruda ex H.Kost., Trav. Brazil 499 (1816), nom. nud. pro parte quoad P. rufa tantum. Trees or shrubs. Leaves with stipules subulate or deltate, usually persistent or sub-persistent; petiole eglandular; lamina often with 1 or 2 pairs of glands at base and sometimes with several small marginal glands especially near apex; lower surface of lamina glabrous or with an arachnoid indumentum. Inflorescence usually a congested thyrse, often with a few ascending branches, less often a few-flowered spike or raceme, very rarely flowers solitary, or densely crowded in a long-pedunculate compound corymb. Pedicels usually shorter than (often much shorter than) receptacle tube. Bracts and bracteoles usually persistent, rarely enclosing flower-buds in small groups, eglandular. Flowers bisexual, slightly zygomorphic. Receptacle tube turbinate to narrowly cylindric, often slightly curved, ventricose, rarely longer than the calyx, hollow, usually glabrous inside except at throat, less often hairy within; faucal annulus well developed. Sepals 5, subequal, spreading or reflexed, acute or rounded, usually eglandular. Petals 5, ±equalling sepals, orbicular to lingulate, sometimes shortly unguiculate. Stamens usually or more, rarely less than 15; filaments undulate in bud with 3 or more bends; inserted on abaxial surface of faucal annulus, usually forming a complete circle, less frequently unilateral, far-exserted, but not much longer than the combined length of calyx and receptacle tube; staminodes absent or short and filiform. Ovary 1 (2 or 3)-carpellary, 1-locular, inserted laterally at mouth of receptacle tube; style filiform, far-exserted, indistinctly 3-lobed at apex, hairy for greater part of length. Drupe cm or more long; endocarp hard, granular, shortly hairy inside, on germination breaking up irregularly, surface without longitudinal channels, rough and irregular owing to fusiform anticlinal aggregations of stone cells and fibres which penetrate the mesocarp ±deeply. A genus of 71 species confined to the Neotropics, where they range from Mexico to southern Brazil, but only 4 species, C. hondurasensis, C. platycalyx, C. polyandra and C. scottmorii occur outside South America and the genus is unrepresented in the West Indies. Germination is cryptocotylar; cataphylls 8 20; eophylls alternate (D.B.Fanshawe 2232, C. guianensis). A.P. de Candolle, Prodr. 2: 526 (1825); G.Bentham, Contributions towards a flora of South America. XII. Enumeration of plants collected by Mr. Schomburgk in British Guiana, in J.D.Hooker, London. J. Bot. 2: 215 (1840); J.D.Hooker, in G.Bentham & J.D.Hooker, Gen. Pl. 1: 608 (1865) excl. syn. Acioa; J.D.Hooker, in C.F.P.Martius, Fl. Bras. 14(2): 40 (1867); W.O.Focke in H.G.A.Engler & K.A.E.Prantl, Nat. Pflanzenfam. 3(3): 59 (1894); J.Hutchinson, Gen. Fl. Pl. 1: 192 (1964) excl. syn. Dulacia; G.T.Prance, Fl. Neotrop. Monogr. 9: 202, t (1972). 1 Inflorescence a raceme or spike 2 Leaves prominently reticulate beneath, with conspicuous parallel primary veins; exocarp often tomentellous 3 Inflorescence with short silver-gray pubescence; receptacle turbinate; bracteoles caducous 3: Inflorescence with dense ferrugineous or grey-puberulous pubescence; receptacle cylindrical to obconical; bracteoles usually persistent 4 Inflorescence grey puberulous; leaf venation only prominulous; fruit exterior glabrous and smooth 4: Inflorescence densely ferrugineous; leaf venation prominent; fruit exterior verrucose or tomentose 33. C. elata 5. C. bernardii 5

14 SPECIES PLANTARUM FLORA OF THE WORLD (2003) 5 Leaf lower surface with distinct stomatal cavities; fruit exterior verrucose 11. C. foveolata 5: Leaf lower surface reticulate, but without stomatal cavities; fruit exterior usually pubescent, rarely verrucose 6 Leaf acumen mm long; receptacle long and slender, mm long 7. C. parillo 6: Leaf acumen 1 12 mm long; receptacle short and thick, 3 10 mm long 7 Inflorescence many-flowered; leaves ovate to oblong-elliptic cm long 10. C. canomensis 7: Inflorescence few-flowered; leaves elliptic, cm long 8 Leaves elliptic, cm broad, lower surface with pubescence not covering entire surface, and venation conspicuous 8. C. steyermarkii 8: Leaves oblong to oblong-lanceolate, cm broad, lower surface with lanate pubescence covering entire surface and obscuring reticulate venation 9. C. canescens 2: Leaves not prominently reticulate beneath; fruit exocarp always glabrous, smooth or verrucose 9 Bracteoles persistent, and at least 3 /4 the length of receptacle or more than 10 mm long 10 Flowers not more than 4 mm long 17. C. spicata 10: Flowers 6 mm long or more 11 Flowers mm long; stamens more than 50; leaves cm long 12 Primary leaf veins distinctly anastomosing at margins to form a marginal vein; leaf base rounded to subcordate; receptacle subcylindrical or tubular, mm long 13 Flowers mm long; receptacle subcylindrical, glabrous except for reflexed hairs at apex; stipules to 15 mm long, caducous 38. C. insignis 13: Flowers mm long; receptacle tubular, tomentose within almost to base; stipules mm long, persistent 39. C. cidiana 12: Primary leaf veins not anastomosing; leaf base cuneate to rounded; receptacle tubular, mm long 14 Leaves elliptic, cm broad; primary veins pairs; receptacle tube mm long 36. C. martinii 14: Leaves oblong-lanceolate, mm broad; primary veins pairs; receptacle tube mm long 37. C. bondarii 11: Flowers 6 12 mm long; stamens 15 28; leaves 6 18 cm long 15 Leaves cm long, caudate-acuminate; exterior of receptacle with red-brown pubescence; stamens 15 19; style pubescent at base only 13. C. exflexa 15: Leaves 6 13 cm long, apex acuminate; exterior of receptacle with light-brown sericeous pubescence; stamens c. 25; style pubescent for 3 /4 of length 14. C. habrantha 9: Bracteoles caducous or less than 1 /2 the length of receptacle, under 10 mm long 16 Receptacle pubescent inside to the base; filaments hirsute; leaves hirsute beneath 40. C. recurva 16: Receptacle glabrous inside except at throat; filaments glabrous; leaves with arachnoid indumentum or glabrous beneath 17 Stamens less than 40 6

15 CHRYSOBALANACEAE (Couepia) 18 Receptacle subcylindrical, densely ferrugineous-sericeous 19 Leaves cm long; primary veins 8 14 pairs, slightly impressed; petioles 2 5 mm long 15. C. scottmorii 19: Leaves 7 24 cm long; primary veins pairs, plane or slightly impressed; petioles 6 10 mm long 20 Leaves narrowly oblong, 7 10 cm long, cm broad, glabrous beneath; receptacle light-brown on exterior; stamens c C. carautae 20: Leaves broadly oblong, cm long, cm broad, ferrugineous-lanate-pubescent beneath; receptacle ferrugineous on exterior; stamens c C. magnoliifolia 18: Receptacle cylindrical, with sparse appressed pubescence only 21 Leaves obovate, cm long, bluntly acuminate; bracteoles persistent or subpersistent; stamens in a complete circle 41. C. obovata 21: Leaves oblong-lanceolate, 9 15 cm long, with a long acumen; bracteoles caducous; stamens unilateral 1. C. guianensis 17: Stamens numerous (more than 60) 22 Receptacle glabrous or with sparse appressed grey tomentum on exterior 23 Leaves 9 17 cm long, densely appressed-lanate pubescent beneath; primary veins pairs 42. C. williamsii 23: Leaves 5 8 cm long, sparsely pubescent beneath; primary veins 7 10 pairs 44. C. marleneae 22: Receptacle with dense brown pubescence on exterior 24 Leaves oblong-lanceolate 27. C. krukovii 24: Leaves oblong to oblong-elliptic 25 Primary veins pairs 26. C. macrophylla 25: Primary veins pairs 26 Leaf apex with prominent well-developed acumen 46. C. chrysocalyx 26: Leaf apex rounded to bluntly acuminate 28. C. latifolia 1: Inflorescence a panicle 27 Bracts and bracteoles persistent at flowering, at least 1 /2 as long as receptacle 28 Flowers mm long; exterior of receptacle and calyx lobes with a long ferrugineous sericeous pubescence; leaves glabrous (rarely glabrescent) beneath 47. C. eriantha 28: Flowers 7 20 mm long; exterior of receptacle shortly brown- to greytomentose; leaves arachnoid-pubescent beneath 29 Leaves thick and coriaceous; bracteoles always persisting through flowering; rachis of inflorescence and receptacle longitudinally striate, or if not, petioles mm long 30 Flowers 7 15 mm long; leaf bases subcordate, rarely rounded; rachis and receptacle with longitudinal striations 18. C. bracteosa 30: Flowers mm long; leaf bases rounded to subcuneate; rachis and receptacle not longitudinally striate 20. C. belemii 29: Leaves thin and membranous; bracteoles persistent only in bud; rachis and receptacle not longitudinally striate; petioles 4 8 mm long 31 Stamens connate at base for at least 1 mm; receptacle tapering to base, subturbinate 48. C. trapezioana 31: Stamens free almost to base; receptacle subcylindrical 19. C. subcordata 7

16 SPECIES PLANTARUM FLORA OF THE WORLD (2003) 27: Bracts and bracteoles not persistent at flowering, or small and inconspicuous 32 Interior of receptacle filled with hairs to base 33 Primary veins 10 15; exterior of receptacle and calyx lobes with sparse appressed pubescence not completely covering surface 2. C. paraensis 33: Primary veins 17 28; exterior of receptacle and calyx lobes densely pubescent, completely covering surface 34 Petioles canaliculate above; flowers mm long; exterior of receptacle and calyx lobes with short brown pubescence; stamens C. excelsa 34: Petioles not canaliculate; flowers 8 12 mm long; exterior of receptacle and calyx lobes with sparse short grey pubescence; stamens C. caryophylloides 32: Interior of receptacle glabrous except for deflexed hairs at throat 35 Leaf lower surface completely glabrous; exterior of receptacle usually almost glabrous 36 Peduncles elongated (30 80 cm long); exterior of calyx lobes with 2 sessile glands 37 Exterior of receptacle pubescent; calyx tube cylindrical-turbinate, mm long; stamens c. 32; petioles 4 8 mm long 51. C. longipendula 37: Exterior of receptacle glabrous or almost so; calyx tube campanulate, 4 6 mm long; stamens 16 21; petioles mm long 52. C. dolichopoda 36: Peduncles short (less than 10 cm long); exterior of calyx lobes eglandular 38 Stamens C. joaquinae 38: Stamens Leaves oblong to oblong-lanceolate, prominently acuminate; inflorescence little-branched, almost racemose; petioles 8 12 mm; stamens c C. glabra 39: Leaves ovate to elliptic, bluntly acuminate; inflorescence much branched; petioles 4 7 mm; stamens C. glabra 35: Leaf lower surface lanate; exterior of receptacle pubescent 40 Exterior of receptacle and calyx lobes sparsely appressedpuberulous, the pubescence not forming a complete covering 41 Stipules persistent, adnate to the base of petiole; rachis of inflorescence 3 5 mm thick; petal margins glabrous 49. C. stipularis 41: Stipules caducous, not adnate to base of petiole; rachis of inflorescence mm thick; petal margins ciliate 42 Leaf lower surface with distinct, prominulous, parallel secondary venation at 90 to primary veins 5. C. bernardii 42: Leaf lower surface densely lanate-pubescent and smooth, venation obscured beneath pubescence 43 Leaves bluntly or shortly acuminate to obtuse; receptacle subcampanulate, 3 5 mm thick at top; inflorescence a muchbranched panicle 44 Leaves ovate to oblong, cm broad 2. C. paraensis 44: Leaves oblong-lanceolate, cm broad 3. C. maguirei 43: Leaves prominently acuminate; receptacle cylindrical, c. 2 mm thick at top; inflorescence often with only short 2 3 flowered branches 8

17 CHRYSOBALANACEAE (Couepia) 45 Inflorescences erect, much-branched, predominantly terminal panicles 1. C. guianensis 45: Inflorescences little-branched racemose panicles, axillary or predominantly axillary, but with a small terminal branch 46 Inflorescences single racemose reflexed panicles; bracteoles persistent 50. C. reflexa 46: Inflorescences erect, terminal and in the upper 2 6 axils; bracteoles caducous 1. C. guianensis 40: Exterior of receptacle and calyx lobes with dense pubescence forming a complete covering 47 Leaf lower surface prominently reticulate 48 Leaves with deep well-defined stomatal cavities, cm long, bluntly acute at apex 35. C. amaraliae 48: Leaves without stomatal cavities, acuminate at apex, or with poorly defined shallow cavities, and then exceeding 6 cm in length 49 Receptacle gradually tapering to a long slender pedicel; exterior of receptacle rufous-pubescent; primary veins not impressed on upper leaf surface 53. C. cognata 49: Receptacle changing abruptly to a short thick pedicel; exterior of receptacle brown-pubescent; primary veins impressed on upper leaf surface 34. C. racemosa 47: Leaf lower surface with plane to prominent venation but not prominently reticulate 50 Receptacle slightly curved anteriorly in bud 51 Leaf with midrib deeply impressed above 53. C. cognata 51: Leaf with midrib plane above 52 Leaves 4 7 cm long; primary veins C. pernambucensis 52: Leaves cm long; primary veins Exterior of receptacle and calyx lobes grey-puberulous; primary leaf veins 18 23, slightly impressed above 64. C. impressa 53: Exterior of receptacle and calyx lobes brown-tomentose; primary leaf veins 12 14, plane above 54. C. multiflora 50: Receptacle erect, not curved in bud 54 Outer surface of petals distinctly pubescent 55 Leaves cm long; primary veins Leaves oblong-lanceolate to oblong; acumen 5 7 mm long; base cuneate; petioles 5 7 mm long 71. C. parvifolia 56: Leaves orbicular to elliptic; acumen 0 3 mm long; base rounded to cordate; petioles 2 mm long 57 Leaf lower surface rufous; stamens 25 26; exocarp smooth; petioles 2 5 mm 15. C. scottmorii 57: Leaf lower surface grey; stamens 30 60; exocarp verrucose; petioles 2 mm 24. C. uiti 55: Leaves (4 ) 6 16 cm long; primary veins Flowers 6 12 mm long; receptacle cylindrical, c. 2 mm thick at top below calyx lobes 9

18 SPECIES PLANTARUM FLORA OF THE WORLD (2003) 59 Inflorescence and flowers densely ferrugineous-sericeous; flowers 8 12 mm long; petioles 5 10 mm long; receptacle not striate on exterior 65. C. meridionalis 59: Inflorescence and flowers with short grey pubescence; flowers 6 8 mm long; petioles 3 7 mm long; exterior of receptacle longitudinally striate 55. C. ulei 58: Flowers mm long; receptacle subcampanulate, 4 6 mm thick 60 Pedicels 6 15 mm long; exterior of receptacle and calyx lobes short grey-sordid-puberulent 25. C. cataractae 60: Pedicels not exceeding 6 mm in length; exterior of receptacle and calyx lobes spreading brown-tomentose 61 Leaves cm long, cm broad, rufouspubescent beneath, greying with age 56. C. comosa 61: Leaves cm long, cm broad, grey-pubescent beneath 62 Leaves coriaceous, rounded to bluntly acuminate at apex 32. C. grandiflora 62: Leaves chartaceous, prominently acuminate 57. C. venosa 54: Petals glabrous except for ciliate margins 63 Receptacle cylindrical or subcylindrical 64 Inflorescence of densely clustered glomerules 65 Leaves oblong, cm broad with short abrupt acumen 2 3 mm long; petioles mm; primary veins impressed above 66. C. coarctata 65: Leaves elliptic, cm broad with long curved acumen mm long; petioles 5 7 mm; primary veins plane above 60. C. hondurasensis 64: Inflorescence a loosely branched panicle 66 Stamens c. 35; leaf base subcordate to rounded 19. C. subcordata 66: Stamens 15 28; leaf base rounded to cuneate 67 Primary leaf veins 8 14 pairs; petioles terete or canaliculate; stamens around partial or complete circle 68 Flowers borne on distinctly articulated pedicels with lower part of pedicel persistent after fall of some flowers 6. C. monteclarensis 68: Pedicels not conspicuously articulated and no part remaining after flower fall 69 Petioles terete, with 2 (often obscure) glands; stamens 11 19, around of circle 58. C. polyandra 69: Petioles canaliculate, eglandular; stamens 19 28, around 3 /4 or complete circle 70 Leaves cm; base cuneate, subcuneate, equal; petioles 4 6 mm 4. C. sandwithii 70: Leaves cm; base rounded, unequal; petioles 8 11 mm 30. C. leitaofilhoi 67: Primary leaf veins pairs; petioles canaliculate; stamens always around complete circle 71 Petioles with 2 glands; leaves cm long; stamens ± C. nutans 71: Petioles eglandular; leaves 7 15 cm long; stamens

19 CHRYSOBALANACEAE (Couepia) 72 Bracteoles 3 15 mm, enclosing young buds; leaf acumen 5 12 mm 22. C. morii 72: Bracteoles 1 35 mm, not enclosing young buds; leaf acumen 3 5 mm 48. C. trapezioana 63: Receptacle broadly campanulate or turbinate 73 Stamens Leaves ovate, 4 9 cm long; primary veins 11 15; flowers 8 12 mm long; petioles 4 8 mm long 29. C. ovalifolia 74: Leaves oblong, cm long; primary veins 16 20; flowers mm long; petioles mm long 31. C. schottii 73: Stamens more than Receptacle broadly turbinate, flattened, 3 mm long, almost solid 61. C. platycalyx 75: Receptacle subcampanulate, not flattened, 4 12 mm long, hollow 76 Petioles mm long; inflorescence densely crowded 67. C. longipetiolata 76: Petioles 2 12 mm long; inflorescence lax 77 Leaves cm 24. C. uiti 77: Leaves cm 78 Leaf margins undulate and revolute 62. C. rufa 78: Leaf margins plane 79 Leaf with short and blunt acumen; lower surface rufous-pubescent; calyx lobes rufous-pubescent on exterior and yellow-brown-pubescent within 63. C. robusta 79: Leaf with finely pointed acumen, mm long or abrupt acumen 3 10 mm long; lower surface grey or brown pubescent; calyx lobes grey-pubescent on both surfaces 80 Primary leaf veins pairs; close together; leaf acumen mm long; exocarp smooth 69. C. froesii 80: Primary leaf veins pairs, cm apart; leaf acumen 3 10 mm long abrupt; exocarp lenticellate 70. C. rankiniae 1. Couepia guianensis Aubl., Hist. Pl. Guiane 1: 519 (1775) Acia amara Willd., Sp. Pl. ed. 4, 3(1): 717 (1800), nom. superfl. illeg.; Acioa amara Steud., Nomencl. Bot., ed. 1, 9 (1821), nom. superfl. illeg.; Moquilea couepia Steud., Nomencl. Bot., ed. 2, 2: 159 (1841), nom. superfl. illeg. T: French Guiana: J.B.C.F.Aublet s.n.; lecto: BM, fide G.T.Prance, Fl. Neotrop. Monogr. 9S: 64 (1989); photo: BR. Illustration: J.B.C.F.Aublet, Hist. Pl. Guiane, Atlas t. 207 (1775). Tree to 25 m tall. Branches puberulous when young, soon becoming glabrous. Leaves with stipules 1 3 mm long, linear, early caducous; petioles 3 9 mm long, canaliculate above, pubescent when young, becoming glabrous and rugose with age; lamina oblong to oblonglanceolate, membranous to coriaceous, cm, rounded to subcuneate at base, acuminate at apex with acumen 5 18 mm long, glabrous above, densely grey- to brown-lanate, or glabrous, or with a sparse caducous pubescence beneath, frequently with 2 glands at junction with petiole; primary veins pairs, plane above, prominent beneath; midrib prominulous above, prominent beneath. Inflorescences terminal and axillary littlebranched panicles or racemes; rachis and branches sparsely puberulous to glabrous. Bracts and bracteoles minute, membranous, ovate, caducous. Receptacle cylindrical to subcampanulate, ( 12) mm long, mm broad below calyx, sparsely puberulous to glabrous externally, glabrous within except for deflexed hairs around throat; pedicels mm 11

20 SPECIES PLANTARUM FLORA OF THE WORLD (2003) long. Calyx lobes 5, rounded, mm long, puberulous or glabrous externally. Petals 5, white, ciliate. Stamens 14 30, unilateral, inserted around 1 /2 of a circle with short staminodes opposite them. Ovary villous. Style pubescent for at least 1 /2 its length. Fruit rounded to ovoid, 3 4 cm long, cm broad; exocarp smooth, glabrous; mesocarp thin, fleshy; endocarp thin, fragile, granular in texture, glabrous within. 1 Inflorescences of racemes (rarely with a few short branches bearing 2 flowers), usually in at least 3 axils below the apex of branch; leaf lamina cm long, chartaceous; petioles 6 9 mm long, receptacle always narrowly cylindrical; growing on terra firme 1: Inflorescences of panicles, usually only 1 or 2 axils below apex; leaf lamina cm long, coriaceous or chartaceous; petioles 3 6 mm long; receptacle cylindrical or subcampanulate; growing on terra firme or flooded river banks 2 Leaves chartaceous; lower side glabrous or sparsely pubescent; on flooded river banks 2: Leaves thickly coriaceous; lower side densely lanate-pubescent; on terra firme 1a. Couepia guianensis Aubl. subsp. guianensis 1a. subsp. guianensis 1b. subsp. glandulosa 1c. subsp. divaricata Couepia leptostachya Benth. ex Hook.f., in C.F.P. Martius, Fl. Bras. 14(2): 44 (1867). T: Brazil North: Amazonas, Manaus, R.Spruce 1536; lecto: K, fide G.T.Prance, Fl. Neotrop. Monogr. 9: 214 (1972); isolecto: BM, C, CGE, GOET, LE, M, NY, OXF, P, RB, W. Couepia versicolor Benoist, Bull. Mus. Natl. Hist. Nat. 29: 596 (1923). T: French Guiana: Maroni, M.Mélinon s.n.; holo: P. Couepia surinamensis Kleinhoonte, Recueil Trav. Bot. Néerl. 22: T: Suriname: Brownsberg, B.W.3080; lecto: U, fide G.T.Prance, Fl. Neotrop. Monogr. 9: 208 (1972); isolecto: K, NY. Map: G.T.Prance, Fl. Neotrop. Monogr. 9S: 212 (1989). Leaf lamina chartaceous, cm long; lower surface densely lanate-pubescent; petioles 6 9 mm long. Inflorescence mainly of terminal axillary racemes, usually in several axils below the apex, a few with short branches bearing 2 or 3 flowers. Receptacle 7 12 mm long, always narrowly cylindrical. Guianas, Venezuela, Colombia, Peru, Ecuador and Brazil. 82: FRG, GUY, SUR, VEN. 83: CLM, ECU, PER. 84: BZN (AM, AP, PA, RO); BZE (MA). Growing on terra firme. Map FRENCH GUIANA: Berges de la Mana, Saut Fracas, F.Hallé 662 (NY, P). GUYANA: U. Takutu-U. Region, Essequibo Kwitaro R., D.Clarke 6270 (K, US). SURINAME: Sectie O, G.Stahel 54 (MAD, U, WAG, WIS, Z). VENEZUELA: Amazonas, Dept. Atabapo, Caño Iguapo, 15 km SE of La Esmeralda, G.Aymard & L.Delgado 8209 (K, PORT). 83. COLOMBIA: Antióquia, Mun. de Turbo, road Tapón del Darién, Sector Río León, J.Brand & M.González 993 (COL, JAUM, MO). PERU: Loreto, Maynas, Iquitos, Puerto Almendras, R.Vásquez & N.Jaramillo (K, MO). 84. BRAZIL NORTH: Amapá, A.M.Bastos 60 (F, IAN, NY, US); Amazonas, Rio Curuquetê, Cachoeira Santo Antônio, G.T.Prance et al (INPA, K, NY, P); Pará, Cuiabá-Santarém km 1180, A.S.Silva et al. 220 (AAU, MG, NY); Rondônia, Mun. de Porto Velho, Samuel Dam area, W.W.Thomas 6316 (INPA, K, NY). 1b. Couepia guianensis subsp. glandulosa (Miq.) Prance, Brittonia 33: 350 (1981) Couepia glandulosa Miq., Stirp. Surinam. Select. 28 (1851); Moquilea glandulosa (Miq.) Walp., Ann. Bot. Syst. 2: 463 (1852). T: Suriname, W.R.Hostmann & A.Kappler 859; holo: U; iso: BM, GH, GOET, K, LE, M, P, S, W. Couepia myrtifolia Benth. ex Hook.f., in C.F.P.Martius, Fl. Bras. 14(2): 44 (1867). T: Venezuela: Amazonas, San Carlos, Río Negro, R.Spruce 3072; lecto: K, fide G.T.Prance, Fl. Neotrop. Monogr. 9S: 209 (1972); isolecto: BM, BR, C, CGE, F, NY, P, TCD, W. Couepia thyrsiflora Hook.f., in C.F.P.Martius, Fl. Bras. 14(2): 45 (1867). T: Venezuela: Amazonas, San Carlos, Río Negro, R.Spruce 3681; lecto: K, fide G.T.Prance, Fl. Neotrop. Monogr. 9: 209 (1972); isolecto: BM, BR, F, GH, GOET, LD, LE, NY, P, RB, W. Illustrations: G.T.Prance, Fl. Venez. Guayana 4: 210 (1998); G.T.Prance & F.White, Philos. Trans. B320: 119 (1988). 12

21 CHRYSOBALANACEAE (Couepia) Map: G.T.Prance, Fl. Neotrop. Monogr. 9S: 212 (1989). Leaf lamina chartaceous to thinly coriaceous, cm long; lower surface glabrous or sparsely lanate-pubescent; petioles 3 6 mm long. Inflorescences of terminal and axillary panicles with small few-flowered branches, in only 1 or 2 axils below the apex of branch. Receptacle 5 10 mm, subcampanulate to narrowly cylindrical. Guianas, Colombia and Brazil. 82: FRG, GUY, SUR, VEN. 83: CLM. 84: BZN (AM, AP, PA, RR), BZE (MA, PE). Growing on flooded river banks. Map FRENCH GUIANA: R. Oyapok, Saut Ako, P.Grenand 2878 (CAY, K). GUYANA: U. Takutu-U. Region, Essequibo, S Pakaraima Mts., Tipuru R., B.Hoffman 1099 (K, US). VENEZUELA: Amazonas, Dept. Atabapo, Río Asisa, L.Delgado 867 (K, PORT). 83. COLOMBIA: Guianía, San Felipe, A.Gentry & B.Stein (MO, NY). 84. BRAZIL NORTH: Amapá, Rio Oiapoque, at mouth of Rio Iauê, W.Egler & J.M.Pires (NY); Amazonas, Alto Rio Negro, P.Acevedo-Rodriguez et al (INPA, US); Pará, Campina de Arumatena, A.Ducke MG15632 (BM, MG, P); Roraima, Rio Branco, Rio Anauá, Campo de Três Ilhas, N.T. da Silva 4521 (FHO, MG, NY). BRAZIL NORTHEAST: Maranhão, Rio Maracassumé, R.L.Fróes 1945 (BM, IAN, NY, P, US); Pernambuco, Escada, Eng. Conceição, D.Andrade-Lima (IPA). 1c. Couepia guianensis subsp. divaricata (Huber) Prance, Brittonia 33: 351 (1981) Couepia divaricata Huber, Bol. Mus. Paraense Hist. Nat. 6: 75 (1910). T: Brazil North: Pará, Belém, J.Huber MG2030; holo: MG; iso: BM. Couepia divaricata var. strictiuscula Huber, Bol. Mus. Paraense Hist. Nat. 6: 76 (1910). T: Brazil North: Pará, Peixe-Boi, R.S. Rodrigues MG8274; lecto: MG, fide G.T.Prance, Fl. Neotrop. Monogr. 9S: 65 (1989); isolecto: BM, P, RB15104, U. Parinari krukovii Gleason, Bull. Torrey Bot. Club 60: 353 (1933), as Parinarium. T: Brazil North: Rondônia, near Tabajara, upper Rio Machado region, B.A.Krukoff 1362; holo: NY; iso: BM, K, MICH, P. Map: G.T.Prance, Fl. Neotrop. Monogr. 9S: 211 (1989). Leaf lamina coriaceous, cm long; lower surface densely lanate-pubescent; petioles 3 6 mm long. Inflorescences of terminal and axillary panicles with small few-flowered branches in only 1 or 2 axils below the apex of branch. Receptacle 5 7 mm long, cylindrical. Guianas and Brazil. 82: GUY, SUR, VEN. 84: BZE (MA), BZN (AM, AP, PA, RO). Growing on terra firme. Map GUYANA: Mazaruni Station, D.B.Fanshawe 242 (FD2978) (K, NY). SURINAME: Sectie O, B.A.Krukoff (NY, US). VENEZUELA: Delta Amacuro, E of Río Grande, ENE of El Plamar, C.Blanco 265 (NY, VEN); Amazonas, 5 km NE of San Carlos de Rio Negro, R.Liesner 4052 (NY, VEN). 84. BRAZIL NORTH: Amapá, Rio Araguari, J.M.Pires et al (IAN, NY); Amazonas, Manaus, Reserva Florestal Ducke, P.A.C.L.Assunção 600 (INPA, K); Pará, Santarém, Km 70 road to Palhão, M.G.Silva & R.Souza 2580 (MG, NY). BRAZIL NORTHEAST: Maranhão, Mun. de Carutapera, Gurupiuna, trib. Rio Gurupi, W.Balée 3018 (MG, NY). 2. Couepia paraensis (Mart. & Zucc.) Benth., J. Bot. (Hooker) 2: 216 (1840) Moquilea paraensis Mart. & Zucc., Abh. Math.-Phys. Cl. Königl. Bayer. Akad. Wiss., 1: 390 (1832). T: Brazil North: Pará, C.F.P.Martius s.n.; lecto: M, fide G.T.Prance, Fl. Neotrop. Monogr. 9: 212 (1972). Tree to 20 m tall, usually smaller. Leaves with stipules 1 2 mm long, caducous; petioles 4 7 mm long, pubescent, canaliculate above; lamina ovate to oblong, cm, rounded to cordate at base, blunt or short-acuminate at apex, with acumen 1 5 mm long, glabrous above, usually densely grey to brown lanate-arachnoid beneath, rarely glabrous or puberulous, with 2 glands at base of lower surface of lamina; primary veins pairs, prominent beneath; midrib glabrous or pubescent above. Inflorescences terminal panicles, sparsely appressedpubescent. Bracts and bracteoles mm long, ovate, caducous. Receptacle subcylindrical to subcampanulate, 8 12 mm long, sparsely appressed-pubescent on exterior, completely hairy within or glabrous except for deflexed hairs at throat. Calyx lobes rounded. Petals 5, white, ciliate on margins. Stamens 25 45, inserted in 1 /2 to 2 /3 of a circle, with short staminodes opposite. Ovary pilose. Style densely pubescent for 3 /4 of its length. Fruit ellipsoid, cm long 2 3 cm broad; epicarp smooth, glabrous; mesocarp thin, fleshy; endocarp thin, fragile, granular in texture, with interior glabrous. 13

22 SPECIES PLANTARUM FLORA OF THE WORLD (2003) 1 Interior of receptacle hairy throughout 2a. subsp. paraensis 1: Interior of receptacle glabrous except for deflexed hairs at throat 2 Bark of young stem thin; on forest and savanna margins 2b. subsp. glaucescens 2: Young stem with a thick corky bark; in cerrado 2c. subsp. cerradoana 2a. Couepia paraensis (Mart. & Zucc.) Benth. subsp. paraensis Couepia rivalis Spruce, Notes Bot. Amazon. 1: 149 (1908), nom. nud. Couepia vismioides Klotzsch ex Hook.f., in C.F.P.Martius, Fl. Bras. 14(2): 48 (1867), nom. nud. in synon. Map: G.T.Prance, Fl. Neotrop. Monogr. 9S: 219 (1989). Bark of young stem thin. Receptacle interior hairy throughout. Eastern and central Amazonian Brazil. 82: VEN. 84: BZN (AM, PA, RO, TO), BZC (GO, MT). Sandy river banks and beaches, and open riverine forest. Map VENEZUELA: Bolívar, Cedeño Distr., S of Caicara del Orinoco, G.Aymard & B.Stergios 3230 (NY, PORT). 84. BRAZIL NORTH: Amazonas, Mun. de Maués, Rio Apoquitaua, above São Sebastião, C.A. Cid Ferreira 4258 (INPA, NY); Amazonas, km 135 Humaitá-Itaituba, G.Vieira et al. 123 (INPA, NY); Pará, Mun. de Santarém, Alter do Chão, L.V.Ferreira 21 (INPA, K); Pará, Mun. de Oriximiná, Igarapé Jaramacará, Campos de Ariramba, G.Martinelli et al (INPA, K, NY); Rondônia, Cachoeira Samuel, B.Maguire et al (NY). BRAZIL WEST-CENTRAL: Goiás, Mun. de Araguatins, E.Mileski 350 (RB); Mato Grosso, Rio Arinos, J.G.Kuhlmann 306 (R). 2b. Couepia paraensis subsp. glaucescens (Spruce ex Hook.f.) Prance, Fl. Neotrop. Monogr. 9: 212 (1972) Couepia glaucescens Spruce ex Hook.f., in C.F.P.Martius, Fl. Bras. 14(2): 49 (1867). T: Brazil North: Amazonas, R.Spruce 1752; lecto: K. fide G.T.Prance, Fl. Neotrop. Monogr. 9: 212 (1972); isolecto: BM, C, CGE, F, GH, GOET, LE, M, NY, OXF, P, RB, TCD, W. Couepia duckei Huber, Bol. Mus. Paraense Hist. Nat. 5: 371 (1909). T: Brazil North: Pará, savannas E of Faro, fl., A.Ducke MG8536; holo: MG; iso: BM, RB Illustration: G.T.Prance, Fl. Venez. Guayana 4: 210 (1998). Map: G.T.Prance, Fl. Neotrop. Monogr. 9S: 219 (1989). Bark of young stem thin. Receptacle interior glabrous except for deflexed hairs at throat. Venezuela and C & W Amazonia. 82: GUY, VEN. 83: BOL, CLM, PER. 84: BZC (MT, TO), BZN (AC, AM, PA, RO, RR). Sandy river banks and beaches and open riverine forest. Map VENEZUELA: Bolívar, Cedeño Distr., Rio Parguaza, between El Carmen & Vuelta Larga, N.Cuello 643 (K, PORT); Amazonas, Dept. Atabapo, E of Caño Perro de Agua, 30 km SE of confluence of Ríos Orinoco and Ventuari, O.Huber & S.Tillett 2757 (MYF, NY, VEN). 83. BOLIVIA: Dept. Pando, Madre de Dios Prov., Río Manupari, San Francisco, T.Killeen 4490 (K, LPB, MO). COLOMBIA: Vichada, P. N. Natural El Tuparro, Caño Peinilla, trib Río Tomo, J.L.Zarucchi & C.E.Barbosa 3626 (MO, NY). PERU: Loreto, Caserío Gamitana, Río Mazán, C.Grández et al (K, MO). 84. BRAZIL NORTH: Acre, Mun. de Cruzeiro do Sul, Rio Juruá, 5 km from Porto Walter, C.A. Cid Ferreira et al (INPA, K, NY); Amazonas, Mun. de Tefé, Lago Tefé, NW shore, T.Plowman et al (INPA, NY); Pará, Mun. de Oriximiná, Rio Mapuera, Campina das 3 Ilhas, C.A. Cid Ferreira et al (INPA, K, NY); Rondônia, margin of Rio Preto, M.R.Cordeiro 695 (IAN, INPA, MG). BRAZIL WEST-CENTRAL: Mato Grosso, Rio Xingu, B.C. dos Passos 1082 (RB). 2c. Couepia paraensis subsp. cerradoana Prance, Fl. Neotrop. Monogr. 9: 213 (1972) T: Brazil North: Tocantins, Filadélfia, G.T.Prance & N.T.Silva 58519; holo: NY; iso: F, M, MO, UB. Map: G.T.Prance, Fl. Neotrop. Monogr. 9S: 219 (1989). Bark of young stem thick and corky. Receptacle interior glabrous, except for deflexed hairs at throat. In the north of the Planalto of central Brazil. 84: BZC (MT), BZE (MA), BZN (PA, TO). Growing in cerrado. Map