Vol. 2 part 1 - Species Plantarum Programme

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

© 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. 

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: 1–268 (2003). 

National Library of Australia 

Cataloguing in Publication entry 

EDITORS 

Anthony E. Orchard 

Annette J.G. Wilson 

ASSISTANT EDITOR 

Brigitte Kuchlmayr 

Prance, Ghillean. 

Chrysobalanaceae 2, Acioa to Magnistipula. 

Bibliography. 

Includes index. 

ISBN 0 642 56833 2. 

This book is available from: 

Australian Biological Resources Study 

GPO Box 787 

Canberra ACT 2601 

Australia 

1. Chrysobalanaceae. I. Australian Biological Resources Study. 

II. Title. (Series : Species plantarum: flora of the world ; pt. 10). 

III. Sothers, Cynthia. 

584.82

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

iv 

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.

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

vi 

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

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 http://www.deh.gov.au/biodiversity/abrs) with links from the IOPI site (currently 

http://life.csu.edu.au/iopi/iopihome) and others. 

A.E.Orchard & A.J.G.Wilson 

Canberra 

November 2003 

vii

viii

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: 112–115 (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

2 

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. 14015; 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 1.5–2.5 cm 

long, terete, glabrous; lamina oblong, thickly coriaceous, 7–17 × 4.5–12 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, 6–7.5 cm long, 4–5 cm broad; exocarp 

glabrous, lenticellate; mesocarp 12–14 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 1. 

84. 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. 4348 (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). 


Tree, medium-sized. Leaves with stipules caducous (not seen); petioles 5–12 mm long, 1.5–3 mm 

thick, glabrous, terete; lamina ovate to oblong, 7–17.5 × 3–8.3 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, 

10–30 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,

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. 

14015). 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

4 


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 2. 

82. 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. 50490 (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). 


Tree, small. Leaves with stipules caducous (not seen); petioles 3–6 mm long, glabrous, thick; 

lamina broadly elliptic to oblong-elliptic, 8–16 × 4.5–8.5 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, 3.8 × 3 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 3. 

82. VENEZUELA: Amazonas, San Carlos de Río Negro, L.M.Berti & P.Salcedo 86-979 (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. 


Tree? Leaves with stipules caducous (not seen); petioles thick, 8–10 mm long, 4–5 mm 

broad; lamina broadly elliptic to oblong-elliptic, 10–15 × 6–9 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, 18–20 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 10–12 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 4. 

84. 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.

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 20–100 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 2.5–12 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.34–41 (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 33. C. elata 

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 5. C. bernardii 

4: Inflorescence densely ferrugineous; leaf venation prominent; fruit 

exterior verrucose or tomentose 

5

6 


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 5.5–18 mm long; receptacle long and slender, 

11–22 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 

8.5–18 cm long 10. C. canomensis 

7: Inflorescence few-flowered; leaves elliptic, 2–6.5 cm long 

8 Leaves elliptic, 2.7–4 cm broad, lower surface with pubescence 

not covering entire surface, and venation conspicuous 8. C. steyermarkii 

8: Leaves oblong to oblong-lanceolate, 1.5–3.2 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 16–75 mm long; stamens more than 50; leaves 

14–26 cm long 

12 Primary leaf veins distinctly anastomosing at margins to form a 

marginal vein; leaf base rounded to subcordate; receptacle 

subcylindrical or tubular, 30–50 mm long 

13 Flowers 16–18 mm long; receptacle subcylindrical, glabrous except 

for reflexed hairs at apex; stipules to 15 mm long, caducous 38. C. insignis 

13: Flowers 60–75 mm long; receptacle tubular, tomentose within 

almost to base; stipules 20–35 mm long, persistent 39. C. cidiana 

12: Primary leaf veins not anastomosing; leaf base cuneate to 

rounded; receptacle tubular, 12–15 mm long 

14 Leaves elliptic, 6–10.5 cm broad; primary veins 10–12 pairs; 

receptacle tube 20–25 mm long 36. C. martinii 

14: Leaves oblong-lanceolate, 3.5–7 mm broad; primary veins 

17–25 pairs; receptacle tube 12–15 mm long 37. C. bondarii 

11: Flowers 6–12 mm long; stamens 15–28; leaves 6–18 cm long 

15 Leaves 10–18 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

CHRYSOBALANACEAE (Couepia) 

18 Receptacle subcylindrical, densely ferrugineous-sericeous 

19 Leaves 2.5–5 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 15–20 pairs, plane or 

slightly impressed; petioles 6–10 mm long 

20 Leaves narrowly oblong, 7–10 cm long, 2–3.3 cm broad, glabrous 

beneath; receptacle light-brown on exterior; stamens c. 35 16. C. carautae 

20: Leaves broadly oblong, 12–24 cm long, 4.5–8.5 cm broad, 

ferrugineous-lanate-pubescent beneath; receptacle 

ferrugineous on exterior; stamens c. 25 12. C. magnoliifolia 

18: Receptacle cylindrical, with sparse appressed pubescence only 

21 Leaves obovate, 2.5–10 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 12–15 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 24–30 pairs 26. C. macrophylla 

25: Primary veins 16–20 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 20–25 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 13–18 mm long 

30 Flowers 7–15 mm long; leaf bases subcordate, rarely rounded; 

rachis and receptacle with longitudinal striations 18. C. bracteosa 

30: Flowers 18–22 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

8 


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 12–17 mm long; exterior 

of receptacle and calyx lobes with short brown pubescence; 

stamens 38–40 23. C. excelsa 

34: Petioles not canaliculate; flowers 8–12 mm long; exterior of 

receptacle and calyx lobes with sparse short grey pubescence; 

stamens 20–35 21. 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, 

14–22 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 

10–12 mm long 52. C. dolichopoda 

36: Peduncles short (less than 10 cm long); exterior of calyx lobes 

eglandular 

38 Stamens 9–11 45. C. joaquinae 

38: Stamens 25–110 

39 Leaves oblong to oblong-lanceolate, prominently acuminate; 

inflorescence little-branched, almost racemose; petioles 

8–12 mm; stamens c. 110 43. C. glabra 

39: Leaves ovate to elliptic, bluntly acuminate; inflorescence much 

branched; petioles 4–7 mm; stamens 25–45 2. 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 1–2.5 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, 2.5–8.5 cm broad 2. C. paraensis 

44: Leaves oblong-lanceolate, 1.5–4 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


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, 2.5–6 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 7–10 68. C. pernambucensis 

52: Leaves 9–18.5 cm long; primary veins 12–23 

53 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 2.5–5.5 cm long; primary veins 5–9 

56 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 9–17 

58 Flowers 6–12 mm long; receptacle cylindrical, c. 2 mm 

thick at top below calyx lobes 

9

10 


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 12–20 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 4–8.5 cm long, 2–3.8 cm broad, rufouspubescent 

beneath, greying with age 56. C. comosa 

61: Leaves 7.5–18 cm long, 3.5–9 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, 4–4.5 cm broad with short abrupt acumen 

2–3 mm long; petioles 14–18 mm; primary veins 

impressed above 66. C. coarctata 

65: Leaves elliptic, 5.5–9.5 cm broad with long curved 

acumen 10–20 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 18–240° of circle 58. C. polyandra 

69: Petioles canaliculate, eglandular; stamens 19–28, 

around 3 /4 or complete circle 

70 Leaves 4.5–9.5 × 2–3.5 cm; base cuneate, 

subcuneate, equal; petioles 4–6 mm 4. C. sandwithii 

70: Leaves 9–17 × 3.5–7.5 cm; base rounded, unequal; 

petioles 8–11 mm 30. C. leitaofilhoi 

67: Primary leaf veins 14–19 pairs; petioles canaliculate; 

stamens always around complete circle 

71 Petioles with 2 glands; leaves 13–16 cm long; stamens ±16 59. C. nutans 

71: Petioles eglandular; leaves 7–15 cm long; stamens 20–24


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 14–20 

74 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, 10–18 cm long; primary veins 16–20; 

flowers 11–15 mm long; petioles 10–15 mm long 31. C. schottii 

73: Stamens more than 28 

75 Receptacle broadly turbinate, flattened, 3 mm long, 

almost solid 61. C. platycalyx 

75: Receptacle subcampanulate, not flattened, 4–12 mm long, 

hollow 

76 Petioles 14–18 mm long; inflorescence densely crowded 67. C. longipetiolata 

76: Petioles 2–12 mm long; inflorescence lax 

77 Leaves 2.5–5.5 × 1.5–3.5 cm 24. C. uiti 

77: Leaves 9–27 × 4–12 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, 10–14 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 20–24 pairs; close together; 

leaf acumen 10–14 mm long; exocarp smooth 69. C. froesii 

80: Primary leaf veins 12–16 pairs, 1–2.5 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, 4.5–16.5 × 2.5–5.5 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 10–15 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, 

4.5–10 (–12) mm long, 1–2.5 mm broad below calyx, sparsely puberulous to 

glabrous externally, glabrous within except for deflexed hairs around throat; pedicels 0.5–4 mm 

11

12 


long. Calyx lobes 5, rounded, 2–2.5 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, 2.5–3 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 

9–16.3 cm long, chartaceous; petioles 6–9 mm long, receptacle always 

narrowly cylindrical; growing on terra firme 1a. subsp. guianensis 

1: Inflorescences of panicles, usually only 1 or 2 axils below apex; leaf 

lamina 5–10.5 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 1b. subsp. glandulosa 

2: Leaves thickly coriaceous; lower side densely lanate-pubescent; on 

terra firme 1c. subsp. divaricata 

1a. Couepia guianensis Aubl. subsp. guianensis 

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: 390. 1925. T: Suriname: Brownsberg, 

B.W.3080; lecto: U, fide G.T.Prance, Fl. Neotrop. Monogr. 9: 208 (1972); isolecto: K, NY. 


Leaf lamina chartaceous, 9–16.3 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 5. 

82. 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 10144 (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. 14347 (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).



Leaf lamina chartaceous to thinly coriaceous, 5–10.5 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 6. 

82. 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 46457 

(MO, NY). 84. BRAZIL NORTH: Amapá, Rio Oiapoque, at mouth of Rio Iauê, W.Egler & J.M.Pires 47766 

(NY); Amazonas, Alto Rio Negro, P.Acevedo-Rodriguez et al. 8370 (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 67-5048 (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. 


Leaf lamina coriaceous, 6–10.5 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 7. 

82. GUYANA: Mazaruni Station, D.B.Fanshawe 242 (FD2978) (K, NY). SURINAME: Sectie O, B.A.Krukoff 

12293 (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. 51314 (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, 5.5–16 × 2.5–8.5 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 10–15 pairs, prominent beneath; 

midrib glabrous or pubescent above. Inflorescences terminal panicles, sparsely appressedpubescent. 

Bracts and bracteoles 0.5–2.5 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, 3.5–4 cm long × 2–3 cm broad; epicarp smooth, glabrous; mesocarp thin, fleshy; 

endocarp thin, fragile, granular in texture, with interior glabrous. 

13

14 


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. 


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 8. 

82. 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. 12267 (INPA, K, NY); Rondônia, Cachoeira Samuel, B.Maguire et al. 56726 (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, RB15109. 

Illustration: G.T.Prance, Fl. Venez. Guayana 4: 210 (1998). 


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 9. 

82. 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. 1678 (K, MO). 84. BRAZIL NORTH: Acre, Mun. de Cruzeiro do Sul, Rio 

Juruá, 5 km from Porto Walter, C.A. Cid Ferreira et al. 10527 (INPA, K, NY); Amazonas, Mun. de Tefé, Lago 

Tefé, NW shore, T.Plowman et al. 12458 (INPA, NY); Pará, Mun. de Oriximiná, Rio Mapuera, Campina das 3 

Ilhas, C.A. Cid Ferreira et al. 7795 (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. 


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 10.


84. BRAZIL NORTH: Pará, Conceição do Araguaia, E.Mileski 155 (RB); Barreira do Campo, E.Mileski 187 (RB). 

BRAZIL NORTHEAST: Maranhão, 17 km N of Carolina, Fazenda Madeira, J.A.Ratter et al. 6686 (E, K, UB). 

BRAZIL WEST-CENTRAL: Mato Grosso, Mun. de São Félix do Araguaia, Rio Araguaia, C.A. Cid Ferreira 

et al. 6368 (INPA, NY). 

3. Couepia maguirei Prance, Fl. Neotrop. Monogr. 9: 214 (1972) 

T: Venezuela: Bolívar; Cerro Guaiquinima, Río Paragua, Isla Cassabe, B.Maguire 32714; holo: NY. 

Illustration: G.T.Prance, Fl. Neotrop. Monogr. 9: 215 (1972). 


Shrub or small tree. Branches sparsely pubescent when young. Leaves with stipules linear, 

1–2 mm long, caducous; petioles 4–5.5 mm long, pubescent, canaliculate above; lamina 

oblong-lanceolate, coriaceous, 5–11 × 1.5–4 cm, acuminate at apex with acumen 3–6 mm 

long, rounded to cordate at base, glabrous and shiny above, densely arachnoid-tomentose 

beneath; primary veins 11–16 pairs, prominent beneath. Inflorescences of terminal panicles; 

rachis and branches sparsely grey-pubescent. Bracts and bracteoles c. 5 mm long, caducous. 

Receptacle subcampanulate, 8 mm long, sparsely grey-tomentose on exterior, glabrous 

within except for ring of deflexed hairs around throat; pedicels 2–3 mm long. Calyx lobes 

rounded. Petals 5, deciduous; margins shortly ciliate. Stamens 55–60, inserted around 

complete circle. Ovary densely pilose. Style pilose. Fruit ovoid, exocarp glabrous, smooth; 

endocarp granular thin, fragile, glabrous within. 

Venezuela and Brazil (upper Rio Negro). 82: VEN. 84: BZN (AM). A rheophyte species of 

river islands and rocky places. Map 11. 

82. VENEZUELA: Amazonas, Río Guainia, Maroa, Ll.Williams 14356 (VEN). 84. BRAZIL NORTH: 

Amazonas, Camanaus, Rio Negro, G.T.Prance et al. 16035 (INPA, NY); Amazonas, Rio Negro, R.L.Fróes 

21432 (IAN, NY). 

4. Couepia sandwithii Prance, Fl. Neotrop. Monogr. 9: 216 (1972) 

T: Guyana: Mazaruni Station, D.B.Fanshawe 708 (F.D.3444); holo: NY; iso: BRG, K, U, US. 



Tree, large. Branches glabrous when young. Leaves with stipules small, deciduous; petioles 

4–6 mm long, sparsely pubescent, canaliculate above; lamina oblong to elliptic, 4.5–9 × 

2–3.5 cm, acuminate at apex with acumen to 6 mm long, cuneate or subcuneate at base, 

glabrous above, densely tomentose beneath; primary veins 8–14 pairs, prominent beneath. 

Inflorescences of terminal and axillary panicles; rachis and branches shortly browntomentose. 

Bracts and bracteoles ovate, to 3 mm long. Receptacle cylindrical, 3–4.5 mm 

long, grey-brown tomentose on exterior, glabrous within except for ring of deflexed hairs 

around throat; pedicels c. 1 mm long. Calyx lobes rounded to acute. Petals 5, white, 

deciduous; margins shortly ciliate. Stamens 20–28, inserted around 2/3 or complete circle, 

slightly connate at base. Ovary pilose. Style pilose. Fruit ovoid; exocarp smooth, glabrous. 

Guyana and Venezuela. 82: GUY, VEN. Primary forest. Flowering June–Oct. Map 12. 

82. GUYANA: Region U. Takutu-U. Essequibo, Kaoma Mts., 5 km S of Kassikaityu R., D.Clarke 5015 (K, US); 

Moraballi Creek, D.B.Fanshawe 584 (F.D.3320) (BRG, F, K, NY). VENEZUELA: Bolívar, El Paraiso, 53 km 

NE of Caserío Las Rosas, San Felix, L.M.Berti 685 (NY, VEN); Delta Amacuro, Río Cuyubini, Cerro La 

Paloma, Sierra Imataca, J.A.Steyermark 87610 (NY, VEN). 

5. Couepia bernardii Prance, Brittonia 33: 347 (1981) 

T: Peru: Loreto, Prov. Requena, Río Ucayali, Arboretum Jenaro Herrera, L.Bernardi 6-6; holo: NY; iso: G. 

Illustration: G.T.Prance, Brittonia 33: 348 (1981). 


Tree. Branches puberulous when young, soon becoming glabrous. Leaves with stipules linearlanceolate, 

tomentellous, to 6 mm long, caducous; petioles 4–7 mm long, tomentellous, 

15

16 


Figure 51. Couepia bernardii. A, habit; B, leaf lower surface; C, flower; D, ½ flower; 

E, petal (A–E, L.Bernardi 6-6). Scale bars: A = 3 cm; B = 1 cm; C–E = 2 mm. Drawn by 

Bobbi Angell. Previously published in G.T.Prance, Brittonia 33: 348 (1981).


becoming glabrous with age, rugulose, canaliculate above, eglandular; lamina elliptic, 

chartaceous, 9–16 × 3.5–6.5 cm, rounded to subcuneate at base, apiculate at apex with 

acumen 2–8 mm long, glabrous above, with a short appressed-lanate caducous pubescence 

beneath; midrib prominent on both surfaces; primary veins 12–14 pairs, plane above, 

prominent beneath; secondary venation prominulous, more or less parallel and 90° to primary 

veins. Inflorescences of terminal and axillary little-branched panicles with central rachis and 

short 2–3 flowered branches, or of racemes; rachis and branches puberulous. Receptacle 

cylindrical 7–8 mm long, sparsely grey-puberulous externally, glabrous within except for 

deflexed hairs at throat; pedicels c. 1 mm long. Calyx lobes 5, rounded, 2.5 mm long, 

puberulous externally. Petals 5, white; margins ciliate. Stamens 2–23, unilateral, with arc of 

circle opposite to them toothed. Ovary tomentose. Style pubescent on lower portion only. Fruit 

ovoid, c. 5 cm long × 4 cm broad; exocarp hard and smooth or crustaceous when dry, glabrous; 

mesocarp woody, 6–10 mm thick when dry; endocarp thin, bony, glabrous within. Fig. 51. 

Western Amazonia. 83: PER. 84: BZN (AM). Growing in terra firme forest. Map 13. 

83. PERU: Loreto, Río Ucayali, Arboretum Jenaro Herrera, L.Bernardi 16219 (G, NY); Loreto, Río Ucayali, 

Arboretum Jenaro Herrera, R.Vásquez et al. 2085 (MO, NY); Maynas Prov., Puerto Almendras, Río Nanay, 

G.Grández & A.Sarmiento 1506 (K, MO). 84. BRAZIL NORTH: Amazonas, Mun. de Humaitá, near Três 

Casas, B.A.Krukoff 6381 (BM, BR, NY, US); Amazonas, near Livramento, B.A.Krukoff 6635 (BM, BR, NY, US). 

6. Couepia monteclarensis Prance, Fl. Neotrop. Monogr. 9S: 66 (1989) 

T: Brazil Southeast: Minas Gerais, Estação Biológica de Caratinga, M.A.Lopes 113; holo: BHCB; iso: NY; 

photo: K. 

Illustration: G.T.Prance, Fl. Neotrop. Monogr. 9S: 67 (1989). 

Tree. Branches glabrous when young, conspicuously lenticellate. Leaves with stipules 

caducous (not seen); petioles 8–10 mm long, glabrescent, canaliculate, with a single gland on 

1 side or eglandular; lamina oblong, chartaceous, 7.5–11 × 3.3–4.6 cm, subcuneate at base, 

apiculate at apex with apex 4–6 mm long, glabrous above, with a short brown-lanate 

pubescence beneath; midrib prominulous above, prominent beneath; primary veins 8–10 

pairs, plane above, prominent beneath. Inflorescences of terminal panicles, 6–8 cm long, 

with a central rachis and short dichotomous branches bearing 2 to 4 flowers; rachis and 

branches grey-puberulous. Receptacle cylindrical, 5–7 mm long, grey puberulous externally, 

glabrous within except for deflexed hairs at throat; pedicels 3–5 mm long, articulated at 

middle and with lower part persistent after flower dehiscence. Calyx lobes 5, rounded to 

slightly acute, 2–3 mm long, grey puberulous on exterior. Petals 5, white, glabrous except 

for ciliate margins. Stamens 18–20, inserted almost around complete circle. Ovary inserted 

near mouth of tube, lanate. Style hirsute for 3/4 of length. Fruit when young ovoid; exocarp 

smooth, glabrous; mesocarp granular; endocarp bony, glabrous within. Fig. 52. 

NE and SE Brazil. 84: BZE (BA), BZL (MG, RJ). Growing in Atlantic coastal forest. Map 14. 

84. BRAZIL NORTHEAST: Bahia, Mun. de Wenceslau Guimarães, 3 km W of Nova Esperança, W.W.Thomas 

et al. 9326 (K, NY). BRAZIL SOUTHEAST: Minas Gerais, Estação Biológica de Caratinga, I.R.Andrade 

13366 (BHCB, K); Minas Gerais, Estação Biológica de Caratinga, L.V.Costa 368 (BHCB, K); Minas Gerais, 

Fazenda Montes Claros, Caratinga, C.V.Mendonça Filho 390 (BHCB, SPF); Rio de Janeiro, Mun. de Santa 

Maria Madalena, P. E. Desengano, Serra da Furquilha, Fazenda da Furquilha, C.Farney 1451 (K, NY, RB); 

Rio de Janeiro, Mun. de Parati, Fazenda Taquari, H.C. de Lima 3632 (K, RB); Rio de Janeiro, Mun. de Angra 

dos Reis, Ilha Grande, RBEPS, curso médio do rio Capivari, R.Ribeiro 740 (RB). 

7. Couepia parillo DC., Prodr. 2: 526 (1825) 

Moquilea parillo (DC.) Steud., Nomencl. Bot., ed. 2, 2: 159 (1841), as parilla. T: French Guiana, Herb. 

Ventenat s.n.; holo: G. 

Couepia pauciflora Huber, Bol. Mus. Paraense Hist. Nat. 5: 372 (1909). T: Brazil North: Pará, Lago de 

Faro, Igarapé do Dedal, A.Ducke MG8630; holo: MG; iso: BM, RB. 

Couepia villosa Maguire, Bull. Torrey Bot. Club 75: 378 (1948). T: Suriname: Tafelberg, B.Maguire 24782; 

holo: NY; iso: BR, F, M, MO. 

Illustration: G.T.Prance, Atas do Simpósio sôbre a Biota Amazônica 4: 220–222 (1967). 


17

18 


Figure 52. Couepia monteclarensis. A, habit; B, flower bud; C, ½ flower; D, petal; E, floral 

diagram; F, young fruit; G, fruit cross-section (A–E, M.A.Lopes 113; F–G, BHCP 7147). 

Scale bars: A = 3 cm; B–D = 3 mm; G = 5 mm. Drawn by Bobbi Angell. Previously 

published in G.T.Prance, Fl. Neotrop. Monogr. 9S: 67 (1989).


Tree to 20 m tall or shrub. Leaves with stipules 1.5–5 mm long, linear; petioles 2–9 mm 

long; lamina oblong or oblong-elliptic, 5–15.5 × 1.7–5.8 cm, acuminate at apex with acumen 

5.5–18 mm long, rounded to cuneate at base, glabrous above, prominently reticulate beneath, 

with silver-grey pubescence between reticulations; midrib impressed, pubescent above; 

primary veins 10–15 pairs, prominent beneath, slightly impressed above. Inflorescences 

racemes of 5–21 flowers, or rarely flowers solitary; rachis with ferrugineous-brown pubescence. 

Bracts and bracteoles 2–6 mm long, linear, persistent. Receptacle slender, tubular, 11–22 mm 

long, with thick dense hirtellous pubescence and some longer hairs on exterior, glabrous 

within or with hairs nearly to base on proximal side beneath ovary only; pedicels 1–3 mm 

long. Calyx lobes acute. Petals 5, white, glabrous on margins. Stamens 45–62, inserted almost 

in a complete circle, with a short portion of toothed staminodes. Ovary densely villous. Style 

sparsely pubescent at base. Fruit globose, c. 2.5 cm diameter; epicarp soft yellow-brownvelutinous; 

endocarp thin, fragile, granular in texture, sparsely hirsute within. Fig. 53. 

Guianas and Amazonian Brazil, Ecuador, Peru and Colombia. 82: FRG, GUY, SUR, VEN. 

83: CLM, ECU, PER. 84: BZN (AC, AM, AP, PA, RR). In savannas or primary forest, and 

also common in secondary forest. Map 15. 

Figure 53. Couepia parillo. A, habit; B, detail of stipules and petiole; C, ½ flower and petal; 

D, young fruit (A–C, S.A.Mori et al. 20759; D, Oliveira 4111). Scale bars: A, D = 3 cm; 

C = 5 mm. Drawn by Bobbi Angell. Previously published in S.A.Mori et al., Guide Vasc. Pl. 

Central French Guiana p. 204 (2002). 

19

20 


82. GUYANA: Potaro-Siparuni Region, Iwokrama Reserve, D.Clarke 2598 (K, US). SURINAME: Sipaliwini, 

Blanche Marie Falls, Nickerie R., R.Evans et al. 2671 (K, MO). VENEZUELA: Amazonas, Dept. Río Negro, 

Río Mawarinuma, Neblina Massif, G.Davidse & J.S.Miller 27217 (MO, NY, US, VEN). 83. COLOMBIA: 

Vaupés, near Mitú, J.Betancur et al. 2865 (COL, K, US). ECUADOR: Napo, Aguarico Cantón, Res. Faunística 

Cuyabeno, Laguna Zancudo Cocha, W.Palacios 7751 (K, MO). PERU: Loreto, Prov. Loreto, Nauta, Río 

Marañón, R.Vásquez & N.Jaramillo 3463 (MO, NY). 84. BRAZIL NORTH: Acre, Mun. de Cruzeiro do Sul, 

44 km E of Cruzeiro do Sul, C.A. Cid Ferreira et al. 5336 (INPA, NY); Amazonas, Mun. de Barcelos, 3 km S 

of Central Serra of Serra Aracá, I.L.Amaral 1698 (INPA, NY); Amapá, Mun. de Oiapoque, 109 km SSE of 

Oiapoque road to Calçoene, S.A.Mori et al. 17186 (MG, NY); Roraima, Rio Catrimani, Igarapé Camogi, 

J.M.Pires et al. 14059 (IAN). 

8. Couepia steyermarkii Maguire, Fieldiana, Bot. 28: 252 (1952) 

T: Venezuela: Bolívar, SE slopes of Ptari-tepui, J.A.Steyermark 60025; holo: F; iso: NY, VEN. 


Tree, medium-sized. Leaves with stipules to 2 mm long, subulate, caducous; petioles 5–7 mm 

long, short-hirtellous; lamina elliptic to oblong-ovate 4–6 × 2.7–4 cm, rounded to cuneate at 

base, shortly acuminate at apex with acumen 1–3 mm long, glabrous above, prominently 

reticulate beneath, grey-lanate between reticulations; primary veins 9–12 pairs, prominent 

beneath, impressed above; midrib impressed above, pubescent at base. Inflorescences fewflowered 

axillary racemes; rachis and branches with dense ferrugineous pubescence. Bracts 

and bracteoles 1–3 mm long, ovate, persistent. Receptacle subsessile, cylindrical, 8–10 mm 

long, short-hirtellous on exterior, glabrous within except for deflexed hairs at throat. Calyx 

lobes acute, ovate. Petals 5, white, glabrous on margins. Stamens inserted on 2/3 of circle. 

Ovary densely villous. Style pubescent at base. Fruit unknown. 

Venezuela (highland region of Bolívar state). 82: VEN. In montane forest at 1000–1600 m. 

Map 16. 

82. VENEZUELA: Bolívar, Sifontes Dept., Mun. de Urdaneta, L.Hernández & N.Dezzeo 112 (MYF, NY); 

Bolívar, Distr. Roscio, 15 km ESE of San Ignacio de Yuruaní, O.Huber et al. 9152 (MYF). 

9. Couepia canescens (Gleason) Prance, Acta Bot. Venez. 9: 119 (1974) 

Parinari canescens Gleason, Bull. Torrey Bot. Club 59: 370 (1931), as Parinarium. T: Venezuela: 

Amazonas, Cerro Duida, Aguita Slope, G.Tate 870; holo: NY; iso: K. 



Tree to 10 m tall. Branches tomentellous when young. Leaves with stipules c. 1 mm long, 

linear, tomentose, caducous; petioles 2–3 mm long, densely tomentose, terete; lamina oblong 

to oblong-lanceolate, coriaceous, 2–6.5 × 1.5–3.2 cm, cuneate to rounded at base, acuminate 

at apex with acumen 2–5 mm long, glabrous above except on midrib, densely brown-lanate 

pubescent beneath with pubescence obscuring rather prominent venation, with parallel 

primary veins; midrib impressed and tomentellous towards base above, prominent and lanate 

pubescent beneath; primary veins 10–12 pairs, plane or slightly impressed above, prominent 

beneath. Inflorescences terminal and subterminal racemes 1.5–3 cm long; rachis densely 

brown tomentose. Bracts and bracteoles elliptic, c. 1 mm long, tomentose on exterior, 

caducous. Receptacle campanulate, straight, not curved, 3–4 mm long, brown-tomentose on 

exterior, glabrous within except for deflexed hairs at throat; pedicels 0.5–1 mm long. Calyx 

lobes 5, acute, tomentose on exterior. Petals 5, white, glabrous. Stamens 12–16, unilateral 

with a few short staminodes opposite. Ovary villous. Style glabrous except at extreme base. 

Fruit globose; exocarp glabrous, verrucose when mature; pericarp hard, thick, 

undifferentiated, densely lanate within. 

Venezuela and Guyana. 82: GUY, VEN. Guyana Highland region. Map 17. 

82. GUYANA: Demerara-Mahaica region, track to St. Cuthbert's Mission, T.McDowell 1741 (NY). 

VENEZUELA: Bolívar, SE facing bluffs of NW Chimantá-tepui, J.A.Steyermark 75526 (NY); Amazonas, 

Sierra Parima, J.A.Steyermark 105936 (NY, VEN).


10. Couepia canomensis (Mart.) Benth. ex Hook.f., in C.F.P.Martius, Fl. Bras. 14(2): 

42 (1867) 

Moquilea canomensis Mart., Nov. Gen. Sp. Pl. 2: 80 (1827); Hirtella canomensis (Mart.) Spreng., Syst. Veg. 4 

(Cur. Post.): 341 (1827). T: Brazil North: Amazonas, Canuma, C.F.P.Martius s.n.; holo: M; iso: BR, K. 

Parinari pilosa Standl., Field Mus. Nat. Hist., Bot. Ser. 17(3): 257 (1937). T: Brazil North: Amazonas, 

Mun. de Humaitá, plateau between Rios Livramento and Ipixuna, B.A.Krukoff 7109; holo: F; iso: A, BM, BR, 

IAN, K, LE, MICH, MO, NY, U. 


Shrub or small tree to 10 m tall. Leaves with stipules 3–8 mm long, linear, persistent; petioles 

6–11 mm long, with dense ferrugineous pubescence; lamina ovate to oblong elliptic, 8.5–18 × 

3–8 cm, rounded to subcuneate at base, short-acuminate at apex with acumen 2.5–12 mm 

long, glabrous above, prominently reticulate beneath with short grey-brown pubescence 

between reticulations; primary veins 10–15 pairs, prominent beneath, slightly impressed 

above; midrib impressed, usually pubescent above. Inflorescences densely-flowered axillary 

racemes; rachis and branches ferrugineous-hirtellous. Bracts and bracteoles 2.5–7 mm long, 

persistent. Receptacle subcampanulate-cylindrical, 5.5–10 mm long, dense ferrugineous-brown 

pubescent on exterior, glabrous within except for deflexed hairs at throat; pedicels c. 1 mm 

long. Calyx lobes acute, hirsute on exterior. Petals 5, white, glabrous on margins. Stamens 

40–51, inserted in a nearly complete circle, with a short row of toothed staminodes opposite. 

Ovary densely villous. Style pubescent at base only. Fruit round to oblong, 3.5–5 × 2.5–3.5 cm; 

epicarp short-velutinous; endocarp thin, fragile, granular, sparsely-pubescent within. 

W and C Amazonia and S Guyana. 82: GUY, VEN. 83: CLM, PER. 84: BZN (AM, PA, RO). 

In primary and secondary forest and clearings on non-flooded ground. Map 18. 

82. GUYANA: Rupununi Distr., Kanuku Mts., Two-Head Mt., M.J.Jansen-Jacobs et al. 3434 (K, U). 

VENEZUELA: Amazonas, Savannas of Santo Antonio, Río Orinoco, Ll.Williams 15055 (F, US, VEN). 

83. COLOMBIA: Vaupés, Mitú, lower Río Kubiyú, J.L.Zarucchi 1221 (ECON, NY); Caquetá, Araracuara, 

A.Gentry et al. 65297 (K, MO). 84. BRAZIL NORTH: Amazonas, Mun. de São Paulo de Olivença, Estrada 

Bonfim, D.C.Daly et al. 4408 (INPA, K, NY); Amazonas, Manaus, Reserva Florestal Ducke, M.A.D. de Souza 

467 (INPA, K); Pará, Alto Rio Tapajós, São Raimundo, N.A.Rosa & M.R.Santos 1926 (MG, NY); Rondônia, 

Mun. de Porto Velho, Samuel Dam area, C.A. Cid Ferreira 7427 (INPA, NY). 

11. Couepia foveolata Prance, Fl. Neotrop. Monogr. 9: 221 (1972) 

T: Venezuela: Bolívar, Río Caroní, below Urimán, J.A.Steyermark & J.J.Wurdack 62; holo: NY; iso: BM, F. 



Tree to 30 m tall. Branches hirtellous when young, soon becoming glabrous. Leaves with 

stipules c. 1 mm long, subulate, caducous; petioles 5–9 mm long, hirtellous-pubescent; 

lamina oblong, 3.5–9.5 × 1.5–3.7 cm, acuminate at apex with acumen 3–7 mm long, cuneate 

or subcuneate at base, glabrous and shiny above; prominently reticulate with hair-filled 

stomatal cavities beneath; primary veins 9–11 pairs, prominent beneath; midrib impressed 

above, pubescent towards base. Inflorescences of axillary racemes, few-flowered, densely 

ferrugineous-brown-pubescent. Bracts and bracteoles oblong, to 7 mm long, persistent. 

Receptacle subcylindrical, 8–10 mm long, hirtellous-pubescent on exterior, glabrous within 

except for deflexed hairs around throat. Calyx lobes acute. Petals 5, deciduous; margins 

glabrous. Stamens c. 20, unilateral with tooth-like staminodes opposite. Ovary villous, style 

pilose on basal portion. Fruit ovoid, 2.6 × 3.2 cm; exocarp verrucose; mesocarp thin, fleshy; 

endocarp thin, granular, sparsely hirsute within. 

Venezuela and Brazil. 82: VEN. 84: BZN (AM). Growing in riverine forest. Map 19. 

82. VENEZUELA: Bolívar, Salto Erepuchi, Alto Río Caroní, F.Cardona 2555 (VEN); Bolívar, Mun. Raul 

Leoni, S of Uriman, A.Fernández 2288 (K); Bolívar, Distr. Piar, Río Aparamán, rapids of Yuray-merú, SW 

base of Amaruay-tepui, B.Holst & R.L.Liesner 2669 (MO, NY); Río Paragua between mouth and Quebrada del 

Trueno, B.Stergios 10186 (NY, PORT). 84. BRAZIL NORTH: Amazonas, near Serra Aracá, N.A.Rosa & 

M.R.Cordeiro 1713 (IAN, NY); Amazonas, Rio Negro at confluence with Rio Uaupés, Serra Camaleão, 

D.W.Stevenson et al. 1011 (INPA, K, NY). 

21

22 


12. Couepia magnoliifolia Benth. ex Hook.f., in C.F.P.Martius, Fl. Bras. 14(2): 43 (1867) 

T: Brazil North: Amazonas, Manaus, R.Spruce 1414; lecto: K, fide G.T.Prance, Fl. Neotrop. Monogr. 9: 224 

(1972); iso: C, CGE, F, GH, GOET, LD, LE, NY, OXF, P, RB. (R.Spruce s.n.: BM, W, probably isotypes). 

Illustration: G.T.Prance & F.White, Philos. Trans. B320: 117 (1988). 


Tree, small to medium-sized. Leaves with stipules linear, to 1 mm long, membranous, 

caducous; petioles 7–8 mm long, tomentose, terete; lamina oblong, 14–24 × 5.5–8.5 cm, 

rounded to subcuneate at base, shortly acuminate at apex with acumen 3–5 mm long, 

glabrous above, with dense rufous-brown arachnoid pubescence beneath; primary veins 15–18 

pairs, prominent beneath, slightly impressed above. Inflorescences dense axillary or terminal 

racemose spikes; rachis ferrugineous-brown-sericeous. Bracts ovate, 5–7 mm long, caducous. 

Receptacle subcylindrical, 8–9 mm long, ferrugineous-sericeous on exterior, glabrous within 

except for deflexed hairs at throat; pedicels 1 mm long. Calyx lobes rounded. Petals 5, white, 

with ciliate margins. Stamens c. 25, inserted in a complete circle. Ovary pilose. Style 

densely pubescent for 3/4 its length. Fruit ellipsoid; epicarp smooth, glabrous; pericarp thick, 

fragile, glabrous within. 

French Guiana and Brazil. 82: FRG. 84: BZN (AM). In non-flooded forest. Flowering 

Feb.–June. Map 20. 

82. FRENCH GUIANA: R. Sinnamary, Crique Plomb, D.Loubry 1698 (CAY). 84. BRAZIL NORTH: 

Amazonas, Manaus, Reserva Florestal Ducke, R.C.Forzza 293A (INPA, K); Amazonas, Rio Negro, Araras, 

A.Loureiro et al. INPA39469 (INPA, NY); Amazonas, Manaus-Porto Velho road, km 253, G.T.Prance et al. 

20729 (INPA, NY). 

13. Couepia exflexa Fanshawe & Maguire, Bull. Torrey Bot. Club 75: 376 (1948) 

T: Guyana: Mazaruni Station, D.B.Fanshawe 722 (F.D. 3458); lecto: NY, fide G.T.Prance, Fl. Neotrop. 

Monogr. 9: 225 (1972); isolecto: BRG, K. 


Tree to 30 m tall. Leaves with stipules minute; petioles 10–12 mm long, sparsely pubescent, 

eglandular; lamina oblong, 10–18 × 5–8 cm; obtuse to cuneate at base, apiculate at apex, 

7–10 mm long, glabrous above (sparsely-ferrugineous-pubescent on young leaves), brown 

lanate-arachnoid beneath; midrib prominent above, glabrous except at base; primary veins 

14–17 pairs, prominent on lower surface. Inflorescences densely-flowered spikes 3–6 cm 

long with a reddish-brown pubescence. Bracts 5–8 mm long, ovate, equalling receptacle tube 

in length. Receptacle subsessile, 7–10 mm long, cylindrical but strongly recurved distally on 

anterior, dense red-brown pubescent on exterior, glabrous within except for deflexed hairs at 

throat. Calyx lobes acute. Petals 5, white, deciduous, ciliate on margins. Stamens 15–19, 

slightly connate at base, unilateral, with 5–6 short filamentous staminodes opposite. Ovary 

pilose. Style sparsely pubescent at base. Fruit oblong-oval; epicarp smooth, glabrous; 

pericarp thin, hard, sparsely pubescent within. 

Guianas. 82: FRG, GUY. In primary forest. Map 21. 

82. FRENCH GUIANA: Sinnamary R., above Petit Saut, S.A.Mori et al. 23464 (CAY, NY). GUYANA: 

Pakaraima Mts., Sukabi R., P.Mutchnick 117 (K, US); 15 km S of Mabura, M.Polak & H. ter Steege 186 (K, U); 

Mabura Hill, H. ter Steege 421 (K, U). 

14. Couepia habrantha Standl., Field Mus. Nat. Hist., Bot. Ser. 17: 249 (1937) 

T: Brazil North: Amazonas, Mun. de Humaitá, between Monte Cristo and Santa Vitória, Rio Ipixuna, 

B.A.Krukoff 7252; holo: F; iso: A, BM, BR, IAN, K, MO, NY, RB, U, US. 


Tree to 12 m tall. Leaves with stipules to 2 mm long, caducous; petioles 5–8 mm long, 

canaliculate above; lamina oblong-elliptic, 6–13 × 2.5–6 cm, cuneate to rounded at base, 

acuminate at apex with acumen 4–14 mm long, glabrous above, densely reddish-brown to 

grey-arachnoid beneath; midrib prominent and glabrous on upper surface; primary veins 11–15 

pairs, prominent beneath. Inflorescences short dense axillary and terminal spikes; rachis with


ferrugineous-brown pubescence. Bracts and bracteoles 7–9 mm long, ovate, acute, deciduous. 

Receptacle subsessile, subcylindrical, 8–10 mm long, densely ferrugineous-sericeous on 

exterior, glabrous within except for deflexed hairs at throat. Calyx lobes rounded. Petals 5, 

white, ciliate on margins. Stamens c. 25, inserted in a semicircle, with short filamentous 

staminodes opposite. Style densely pubescent for 3/4 of its length. Ovary pilose. Fruit ellipsoid; 

epicarp smooth, glabrous; mesocarp thin, fleshy; pericarp thin, fibrous, fragile, glabrous within. 

Guyana, Colombia, Venezuela and C Brazilian Amazonia. 82: FRG, GUY, VEN. 83: CLM. 

84: BZN (AM, PA). In riverine and non-flooded forest. Flowering Dec.–May. Map 22. 

82. FRENCH GUIANA: Saül, 10 km ESE of Eaux Claires, S.A.Mori et al. 23795 (K, NY). GUYANA: Potaro- 

Siparuni Region, Kaieteur Falls N. P., W.Hahn 4757 (K, US). VENEZUELA: Amazonas, Río Casiquiare above 

Chapezon, R.Liesner & H.Clark 8913 (MO, NY). 83. COLOMBIA: Amazonas, Rio Caquetá, Araracuara, 

H.Vester 48 (K). 84. BRAZIL NORTH: Amazonas, Manaus, Reserva Florestal Ducke, M.A.D. de Souza 393 

(INPA, K); Amazonas, Carauari, Rio Juruá, A.S.Silva et al. 534 (IAN, NY); Pará, Rio Cururú, alto Rio 

Tapajós, W.Egler 1258 (MG). 

15. Couepia scottmorii Prance, Fl. Neotrop. Monogr. 9S: 68 (1989) 

T: Panama: Prov. Panama, Cerro Jefe, S.A.Mori 7116; holo: NY; iso: BM, MO. 



Tree to 15 m tall. Branches sparsely puberulous when young. Leaves with stipules minute, 

lanceolate, caducous; petioles 2–5 mm long, canaliculate, rugulose, puberulous; lamina 

elliptic, thickly coriaceous, 2.5–5 × 1.6–3 cm, rounded to slightly subcordate at base, 

apiculate to bluntly acuminate at apex with acumen 0–5 mm long, glabrous above, densely 

brown-arachnoid pubescent beneath; midrib prominulous above, prominent beneath; primary 

veins 8–14 pairs, slightly impressed above, prominulous beneath. Inflorescences terminal 

and subterminal racemes 3.5–5.5 cm long; rachis densely yellow-brown tomentellous. Bracts 

and bracteoles caducous. Receptacle subcylindrical, 4–6 mm long, densely tomentellous on 

exterior, glabrous within except for deflexed hairs at throat. Calyx lobes 5, acute. Petals 5, 

sparsely pubescent on exterior, with ciliate margins. Stamens c. 26, inserted around complete 

circle, glabrous except for mass of hair around interior of united bases of filaments. Style 

densely pubescent for 3/4 of its length. Ovary densely lanate. Fruit ellipsoid, 5 × 3 cm; 

exocarp smooth, glabrous; mesocarp thin, fleshy; pericarp c. 3 mm thick. Fig. 54. 

Panama; known only from Cerro Jefe. 80: PAN. In cloud forest dominated by Clusia and 

Colpothrinax cookii at 1000 m. Map 23. 

80. PANAMA: Panamá, Cerro Jefe, G.McPherson 10588 (MO, NY); Panamá, Cerro Jefe, G.McPherson 12340 

(MO, NY); Panamá, Cerro Jefe, summit road trail to Chagras Valley, G.McPherson 15297 (MO, NY). 

16. Couepia carautae Prance, Fl. Neotrop. Monogr. 9S: 70 (1989) 

T: Brazil Southeast: Espírito Santo, Linhares, Reserva Florestal da CVRD, estrada 143A6, km 1.430, Talhão 

601, J.Spada 31/78; holo: INPA; iso: NY, RB; photo: K. 



Tree 22 m tall. Branches glabrous when young. Leaves with stipules linear, to 1 mm long, 

subpersistent; petioles 6–10 mm long, glabrous, rugulose, terete; lamina narrowly oblong, 

coriaceous, 7–10 × 2–3.3 cm, subcuneate at base, acuminate at apex with acumen 4–7 mm 

long, finely pointed, glabrous above, glabrous and waxy beneath; midrib prominulous above, 

prominent and almost glabrous beneath, with only a few short stiff hairs; primary veins 16–20 

pairs, plane above, prominent beneath. Inflorescences terminal and subterminal racemes; 

rachis densely yellow-brown sericeous-tomentose. Bracteoles ovate, 6–7 mm long, tomentose 

on exterior, caducous. Receptacle subcylindrical, 7–8 mm long, yellow-brown sericeous on 

exterior, glabrous within; pedicels 0.5 mm long. Calyx lobes 5, acute. Petals 5, glabrous on 

exterior; margins ciliate. Stamens c. 35, inserted around complete circle, filaments exserted. 

Ovary pilose. Style densely hirsute to apex. Fruit not seen. 

23

24 


Figure 54. Couepia scottmorii. A, habit; B, fruit; C, leaf lower surface; D, flower; E, ½ flower; 

F, petals (A–F, S.A.Mori 7116). Scale bars: A, B = 4 cm; D–F = 5 mm. Drawn by Bobbi 

Angell. Previously published in G.T.Prance, Fl. Neotrop. Monogr. 9S: 69 (1989).


SE Brazil. 84: BZL (ES). In rainforest on clay soil. Map 24. 

Known only from the type collection 

The wood is used for railroad ties (G.T.Prance, 1989: 70). 

17. Couepia spicata Ducke, Arq. Inst. Biol. Veg. 2: 36 (1935) 

T: Brazil North: Amazonas, Manaus, A.Ducke RB 25002; holo: RB; iso: IAN, K, NY, P, U. 


Tree, medium-sized. Leaves with stipules to 4 mm long, lanceolate, caducous; petioles 3–6 mm 

long, pubescent, shallowly canaliculate; lamina oblong-lanceolate, 4.6–9 × 1.4–2.8 cm, cuneate 

at base, acuminate at apex with acumen 6–11 mm long, glabrous above, densely reddishbrown-arachnoid 

beneath; midrib prominent above with lower portion pubescent; primary veins 

9–12 pairs, prominent on lower surface. Inflorescences dense-flowered terminal and axillary 

spikes; rachis with ferrugineous-brown sericeous pubescence. Bracts and bracteoles ovate, 

3–4 mm long, equalling the flowers, persistent. Receptacle sessile, turbinate, 3–4 mm long, 

ferrugineous-brown pubescent on exterior, glabrous within except for deflexed hairs at throat. 

Calyx lobes rounded. Petals 5, white, ciliate on margins. Stamens 14–18, inserted in a semicircle, 

with a sterile portion opposite. Ovary pilose. Style glabrous except at base. Fruit unknown. 

Brazil (Amazonas). 84: BZN (AM). Growing in terra firme forest. Map 25. 

84. BRAZIL NORTH: Amazonas, Manaus-Itacoatiara Km 135, O.P.Monteiro s.n. (INPA50017); Amazonas, 

Mun. de Novo Aripuanã, Nova Prainha, C.D.A.Mota & O.P.Monteiro s.n. (INPA60724); Amazonas, Carauari, 

Poço Juruá 1, A.S.L. da Silva et al. 511 (MG, NY). 

18. Couepia bracteosa Benth., J. Bot. (Hooker) 2: 215 (1840) 

Moquilea bracteosa (Benth.) Walp., Repert. Bot. Syst. 2: 7 (1843).T: Guyana, R. & R.Schomburgk 485; holo: 

K; iso: BM, CGE, GH, L, OXF, TCD, W. 

Moquilea rufa Barb.Rodr., Hort. Flumin. 165 (1895). T: Not traced. 

Couepia bracteosa Benth. var. grandifolia Benoist, Bull. Mus. Hist. Nat. (Paris), sér. 2, 29: 597 (1923). 

T: French Guiana, M.Mélinon s.n.; holo: P. 

Couepia bracteosa Benth. var. minor Ducke, Arch. Jard. Bot. Rio de Janeiro 5: 117 (1930). T: Brazil: 

Amazonas, Manaus, upper rio Tarumã, A.Ducke RB19756; holo: RB; iso: K, US. 

Illustrations: J.D.Hooker, in C.F.P.Martius, Fl. Bras. 14(2), t.15 (1867); G.T.Prance, Fl. Guianas 85: 11 

(1986); G.T.Prance & F.White, Philos. Trans. B320: 119 (1988). 


Tree to 20 m tall, usually smaller. Leaves with stipules 1.5–3.5 mm long, subulate, 

deciduous; petioles 8–18 mm long, sparsely pubescent, shallowly canaliculate; lamina ovateelliptic, 

8–22 × 4–9.5 cm, subcordate at base, short-acuminate at apex, glabrous above, grey 

to brown-lanate beneath; primary veins 12–25 pairs, prominent beneath. Inflorescences 

terminal panicles with short silver-grey pubescence. Bracts ovate, often exceeding the 

receptacle tube, enclosing small groups of buds, 7–12 mm long, persistent. Receptacle 

subsessile, cylindrical, 7–12.5 mm long, densely brown-pubescent on exterior, glabrous 

within except for deflexed hairs at throat. Calyx lobes acute. Petals 5, white, sparsely ciliate 

on margins. Stamens 25–40, connate at base of filaments, inserted in an arc of 270°, with a 

few short filamentous staminodes opposite. Ovary densely villous. Style pubescent. Fruit 

globose, 8 × 7 cm; epicarp verrucose or smooth; mesocarp thick, fleshy; endocarp thin, 

rather fragile, granular, glabrous within. 

Guianas and Amazonia. 80: cos. 82: FRG, GUY, SUR. 83: CLM, PER. 84: BZN (AM, AP, 

PA, RO). In primary and secondary forest on non-flooded ground. Map 26. 

80. COSTA RICA: Cartago, Turrialba (cultivated), K.S.Bawa 383 (MO). 82. FRENCH GUIANA: Piste de 

Saint Elie, B.Riera 869 (CAY, K). GUYANA: Potaro-Siparuni Region, Iwokrama Rainforest Reserve, Burro- 

Burro R., D.Clarke 198 (K, US). SURINAME: Tumuc Humac Mts., W of Talouakem Peak on Brazil border, 

P.Acevedo-Rodriguez 6120 (K, US). 83. COLOMBIA: Vaupés, Estac. Biológica Caparú, A.Rodriguez & 

E.Palacios 95 (K). PERU: Loreto, Prov. Requena, Río Ucayali, Arboretum Jenaro Herrera, L.Bernardi 1-163 

(G, NY). 84. BRAZIL NORTH: Amapá, Mun. de Oiapoque, BR-156, 109 km SSE of Oiapoque towards 

25

26 


Calçoene, S.A.Mori et al. 17219 (MG, NY); Amazonas, Manaus-Porto Velho Highway km 235, G.T.Prance 

et al. 20768 (INPA, NY, P); Pará, Mun. de Oriximiná, Rio Trombetas, 5 km below Cachoeira Porteira, 

C.A. Cid Ferreira et al. 1281 (INPA, NY); Rondônia, Mun. de Porto Velho, Samuel Dam area, C.A. Cid Ferreira 

7470 (INPA, K, NY). 

Widely cultivated for its edible fruit. 

19. Couepia subcordata Benth. ex Hook.f., in C.F.P.Martius, Fl. Bras. 14(2): 46 (1867) 

T: Brazil North: Amazonas, Manaus, R.Spruce 1423; lecto: K, fide G.T.Prance, Fl. Neotrop. Monogr. 9: 228 

(1972); isolecto: CGE, LE, M, NY, OXF, P. (R.Spruce s.n.: BM, W, probably isotypes). 

Couepia amazonica Fritsch, Ann. K. K. Naturhist. Hofmus. 5: 2 (1890). T: Brazil North: Amazonas, Tefé 

(cultivated), E.Poeppig 2814; holo: W; iso: F, GOET, K, L, LE, NY, P. 

Moquilea inaequalis Poepp. ex Fritsch, Ann. K. K. Naturhist. Hofmus. 5: 12 (1890), nom. nud. in synon. 

Parinari polystachya Poepp. ex Fritsch, Ann. K. K. Naturhist. Hofmus. 5: 12 (1890), as Parinarium polystachyum, 

nom. nud. in synon. 

Illustration: J.D.Hooker, in C.F.P.Martius, Fl. Bras. 14(2), t. 16 (1867). 


Tree, medium-sized. Leaves with stipules c. 2 mm long, deciduous; petioles 6–7 mm long, 

pubescent, terete, eglandular; lamina oblong-elliptic, 12–17 × 5.5–7.5 cm, membranous, 

cordate to rounded at base, acuminate at apex with acumen 3–8 mm long, glabrous above, 

with short, thick, silver-grey to brown-arachnoid pubescence beneath; primary veins 13–17 

pairs, prominent beneath. Inflorescences terminal and axillary panicles; rachis and branches 

with short dense light brown pubescence. Bracts and bracteoles ovate, 6–11 mm long, often 

as long as the receptacle tube, persistent up to early flowering. Receptacle subcylindrical, 

9–12 mm long, with short dense pubescence on exterior, glabrous within except for deflexed 

hairs at throat; pedicels 1–2 mm long. Calyx lobes acute. Petals 5, white, ciliate on margins. 

Stamens c. 35, inserted in a complete circle. Ovary villous. Style pubescent for 2/3 its length. 

Fruit ellipsoid, 4.5–9 × 3–5 cm; epicarp smooth, glabrous; mesocarp thick, fleshy; endocarp 

thin, fragile, fibrous, glabrous within. 

Amazonia. 83: CLM, ECU, PER. 84: BZN (AM, PA). Forest. The natural distribution is hard 

to determine since this species is so widely cultivated in Amazonia both by indigenous 

peoples and in towns. Map 27. 

83. COLOMBIA: Amazonas, Leticia, cultivated, E.Acero & R.Rodriguéz 894 (COL). ECUADOR: Los Rios, 

Picilingue, INIAP Experimental Station, T.D.Pennington & G.Tenorio 10687 (K, NY). PERU: Loreto, Prov. 

Requena, Sahua Cocha near Jenaro Herrera, C.Peters 188 (NY); Loreto, Maynas, Río Marañón 120 km from 

Iquitos, J.Torres 85 (ECON). 84. BRAZIL NORTH: Amazonas, Manaus, Colônia Oliveira Machado, 

A.Castellanos s.n. (INPA27546); Amazonas, Rio Solimões, São Paulo de Olivença, E.Lleras et al. P17392 

(INPA, NY); Pará, Belém, cultivated, A.Ducke RB15120 (RB). 

Cultivated in the vicinity of Manaus as a shade plant and in Peru for the edible fruit. The wood 

is used in carpentry and for carving. 

20. Couepia belemii Prance, Fl. Neotrop. Monogr. 9: 228 (1972) 

T: Brazil Northeast: Bahia, Maraú, R.P.Belém & R.S.Pinheiro 3146; holo: NY; iso: UB. 



Tree, medium sized. Branches glabrous when young, rugose. Leaves with stipules caducous 

(not seen); petioles 13–20 mm long, glabrescent, rugose, weakly canaliculate; lamina oblong 

to oblong-elliptic, 11–18 × 4–7 cm, acuminate at apex with acumen 4–8 mm long, rounded to 

subcuneate at base, glabrous above, densely lanate-arachnoid pubescent beneath; midrib 

prominent and glabrous above, prominent beneath; primary veins 18–24 pairs, plane above, 

prominent beneath. Inflorescences of terminal panicles; rachis and branches ferrugineoustomentose. 

Receptacle subsessile, campanulate, 8–14 mm long, densely tomentose on exterior, 

glabrous within except for ring of deflexed hairs around throat. Calyx lobes acute, 7–10 mm


long, tomentose on exterior. Petals 5, white, with ciliate margins. Stamens c. 30, inserted 

around complete circle. Ovary lanate-pilose. Style tomentose. Fruit not seen. 

E and SE Brazil. 84: BZE (BA), BZL (ES). Common in the remnants of the southern Bahia 

wet forest and in coastal forest. Flowering in January. Map 28. 

84. BRAZIL NORTHEAST: Bahia, Mun. de Una, 20 km NW of Una, S.A.Mori et al. 9336 (BISH, CEPEC, K, 

NY); Bahia, Maraú, R.P.Belém 3169 (BR, IAN, NY, UB). BRAZIL SOUTHEAST: Bahia, Reserva Florestal de 

Porto Seguro, G.L.Farias 217 (K); Espírito Santo, Reserva Florestal da CVRD, Linhares, J.S. 007/77 (INPA). 

21. Couepia caryophylloides Benoist, Bull. Mus. Natl. Hist. Nat. 28: 253 (1922) 

T: French Guiana: Godebert, H.Wachenheim 3-11; lecto: P, fide G.T.Prance, Fl. Neotrop. Monogr. 9: 229 

(1972). 


Tree to 25 m tall. Leaves with stipules c. 2 mm long, caducous; petioles 7–9 mm long, 

canaliculate above, glabrous or sparsely pubescent; lamina oblong-elliptic to oblong-lanceolate, 

10–21 × 4–7.5 cm, rounded at base, acuminate at apex with acumen 4–10 mm long, glabrous 

above, densely grey-brown-arachnoid beneath; primary veins 17–25 pairs, prominent beneath; 

midrib glabrous above. Inflorescences dense terminal or axillary panicles 4–6 cm long; 

rachis and branches densely grey-pubescent. Bracts and bracteoles ovate, to 8 mm long, early 

caducous. Receptacle subcylindrical 5–7 mm long, longitudinally ridged when dry, with 

short grey pubescence on exterior; interior lined by dense reflexed hairs to base, or confined 

to ring at throat; pedicels 1–3 mm long. Calyx lobes rounded, ovate, 3–4 mm long. Petals 5, 

white, ciliate on margins. Stamens 22–35, inserted in a semicircle, with a toothed ridge of 

staminodes opposite. Ovary villous. Style pubescent almost to apex. Fruit round to oval; 

epicarp smooth, glabrous; mesocarp thin, fleshy; endocarp granular, fragile, glabrous within. 

Guianas and Brazil. 82: FRG, SUR. 84: BZN (AM, AP, PA). Primary forest on high ground. 

Flowering June to December. 

Interior of receptacle hairy to base 21a. subsp. caryophylloides 

Interior of receptacle glabrous except for ring of hairs around throat 21b. subsp. glabra 

21a. Couepia caryophylloides Benoist subsp. caryophylloides 

Receptacle interior hairy to base. 

Guianas and Northern Brazil. 82. FRG, SUR. 84. BZN (AM, AP, PA). Map 29. 

82. FRENCH GUIANA: Comté R., Etienne, J.J. de Granville [B.]5263 (BR, CAY, K, P); Oiapok, St. Paul, 

R.Oldeman B2514 (P). SURINAME: Mapane Creek, J.Elburg LBB9373 (LBB, US). 84. BRAZIL NORTH: 

Amapá, Rio Oiapoque, Mt. Tipac, H.S.Irwin 48695 (F, GH, IAN, MO, NY, US); Amazonas, Distr. 

Agropecuário, Reserva 1501, N of Manaus, S.A.Mori et al. 20044 (INPA, K, NY); Pará, Cuantã dos Anajás, 

Rio Anajás, H.T.Beck et al. 282 (INPA, K, NY). 

21b. Couepia caryophylloides subsp. glabra Prance, Fl. Neotrop. Monogr. 9S: 72 (1989) 

T: Brazil North: Pará, Santarém-Cuiabá Hwy., km 82, Medeiros & Marinho 58; holo: IAN. 

Receptacle interior glabrous except for ring of hairs around throat. 

Brazil (Pará); known only from the type collection. 84: BZN (PA). Map 30. 

22. Couepia morii Prance, Kew Bull. 47: 254 (1992) 

T: Brazil North: Amazonas, N of Manaus, Distrito Agropecuário, Reserva 1501, S.A.Mori & P.A.C.L.Assunção 

21398; holo: INPA; iso: K, NY. 

Illustration: G.T.Prance, Kew Bull. 47: 255 (1992). 

Tree to 30 m tall. Branches sparsely puberulous when young soon becoming glabrous. 

Leaves with stipules caducous (not seen); petioles 6–10 mm long, sparsely puberulous soon 

becoming glabrous, eglandular, rugose, canaliculate above; lamina oblong, coriaceous, 

27

28 


7–13.5 × 3–5.8 cm; base rounded to subcuneate; apex abruptly acuminate with acumen 

5–12 mm long; glabrous above, densely appressed lanate pubescent beneath; midrib glabrous 

and prominent above, prominent beneath; primary veins 16–19 pairs, plane above, prominent 

beneath. Inflorescences of terminal and axillary little-branched panicles, 2–5 cm long; rachis 

and branches tomentellous; flowers borne on rachis or on small branches in groups of 2 or 3. 

Bracts and bracteoles large and enclosing groups of buds but very early caducous and only 

visible in young inflorescences, 3–15 mm long, membranous tomentellous on exterior, 

puberulous within. Receptacle cylindrical, tomentellous on exterior, glabrous within except 

at throat. Calyx lobes 5, tomentellous on exterior. Petals 5, glabrous except for ciliate 

margins. Stamens c. 20, inserted around complete circle, with a mass of deflexed hairs 

around base. Ovary tomentellous, inserted at mouth of receptacle. Style hirsute for 1/2 of 

length. Fruit not seen. 

Central Amazonia, Brazil. 84: BZN (AM). Forest on terra firme. Map 31. 

84. BRAZIL NORTH: Amazonas, N of Manaus, Distrito Agropecuário, Reserva 1501, M.A. de Freitas et al. 

838 (INPA, K, NY); Amazonas, N of Manaus, Distrito Agropecuário, Reserva 1501, S.A.Mori & E. da Costa 

Pereira 20517 (K, NY, US); Amazonas, N of Manaus, Distrito Agropecuário, Reserva 1501, S.A.Mori et al. 

21411 (K, NY); Amazonas, 90 km N of Manaus, Distrito Agropecuário, Fazenda Dimona, Reserve 2303, 

S.S. da Silva INPA/WWF 3304.2044.2 (K, NY). 

23. Couepia excelsa Ducke, Arch. Jard. Bot. Rio de Janeiro 5:116 (1930) 

T: Brazil North: Pará, Rio Trombetas, Lago Salgado, A.Ducke RB19758; holo: RB; iso: IAN, K, P, U. 


Tree to 25 m tall. Leaves with stipules to 3 mm long, caducous; petioles 8–10.5 mm long, 

terete, pubescent; lamina elliptic to oblong-lanceolate, 13–25 × 4–7 cm, rounded to subcordate 

at base, acuminate at apex with acumen 4–9 mm long, sparsely appressed-pubescent 

above, becoming glabrous with age, densely grey-arachnoid beneath; midrib sparsely 

pubescent above; primary veins 23–38 pairs, prominent on lower surface. Inflorescences 

short-branched panicles or racemes, axillary or terminal, 3–6 cm long; rachis and branches 

brown tomentellous. Bracts and bracteoles ovate, to 8 mm long, caducous. Receptacle 

subcampanulate, 9 mm long, densely brown-pubescent on exterior; interior lined to base by 

dense reflexed hairs; pedicels 2–3 mm long. Calyx lobes rounded, ovate, 4.5–6 mm long. 

Petals 5, white, ciliate on margins. Stamens 38–40, inserted in an arc of 270°, sometimes 

with a short sterile toothed portion of circle opposite. Ovary villous. Style pubescent almost 

to apex. Fruit oval, 6–7 × 4 cm; epicarp smooth, glabrous; endocarp granular, fragile, 

glabrous within. 

Eastern Amazonian Brazil and in French Guiana. 82: FRG. 84: BZN (AP, PA, RR). Nonflooded 

forest. Flowering July–Oct. Map 32. 

82. FRENCH GUIANA: Anapaïké, M.Fleury 1348 [sterile] (CAY, K). 84. BRAZIL NORTH: Amapá, 

Camaipi, S.A.Mori et al. 15859 (K, NY); Amapá, Serra do Navio, Rio Araguari, J.M.Pires et al. 51248 (F, GH, 

K, MG, NY, UB, US); Pará, Mun. de Oriximiná, Rio Cachorro, C.A. Cid Ferreira et al. 8054 (INPA, K, NY); 

Roraima, Manaus-Caracaraí, km 329, W.C.Steward et al. 54 (INPA, NY). 

24. Couepia uiti (Mart. & Zucc.) Benth. ex Hook.f., in C.F.P.Martius, Fl. Bras. 14(2): 

47 (1867) 

Moquilea uiti Mart. & Zucc., Abh. Math.-Phys. Cl. Königl. Bayer. Akad. Wiss., 1: 390 (1832); Moquilea eliti 

(Mart. & Zucc.) Walp., Repert. Bot. Syst. 2: 6 (1843). T: Brazil Northeast: Bahia, banks of Rio São 

Francisco, C.F.P.Martius s.n.; lecto: M, designated here. 

Couepia martiana Hook.f., in C.F.P.Martius, Fl. Bras. 14(2): 47 (1867). T: Brazil Northeast: Piauí, 

C.F.P.Martius s.n.; lecto: M, designated here. 

Couepia paraguariensis Hassl., Repert. Spec. Nov. Regni Veg. 7: 375 (1909). T: Paraguay: 'Alto-P. Chaco', 

K.Fiebrig 1379; holo: G; iso: K. 

Couepia dahlgrenii Standl., Field Mus. Nat. Hist., Bot. Ser. 17(3): 249 (1937). T: Brazil Northeast: Piauí, 

'Campo major', B.E.Dahlgren 986; holo: F. 

Map: G.T.Prance, Fl. Neotrop. Monogr. 9S: 224 (1989).


Tree to 4 m tall or shrub. Leaves with stipules to 6 mm long, linear, membranous; petioles 

2 mm long, densely pubescent, often with 2 conspicuous glands at lamina base, shallowly 

canaliculate; lamina orbicular to elliptic, 2.5–5.5 × 1.5–3.5 cm, rounded to cordate at base, 

short-acuminate or apiculate at apex with acumen 1–3 mm long, sparsely tomentose above 

when young, becoming glabrous with age, densely grey-arachnoid beneath; midrib pubescent 

above; primary veins 5–9 pairs, prominent beneath. Inflorescences terminal panicles; rachis 

and branches with short grey-brown tomentum. Bracts and bracteoles ovate to lanceolate, to 

8 mm long, deciduous. Receptacle subcampanulate, 5–8 mm long, with short light brown 

pubescence on exterior, glabrous within except for deflexed hairs at throat; pedicels 3–5 mm 

long. Calyx lobes rounded. Petals 5, white, sparsely pubescent, ciliate on margins. Stamens 

30–60, inserted in a complete circle. Ovary densely pilose. Style pubescent for 2/3 its length. 

Fruit ovoid, 3–3.5 × 1.5–2 cm; epicarp verrucose, glabrous; mesocarp thick, fleshy; endocarp 

thin, fragile, of a granular texture, glabrous within. 

Bolivia, Brazil and Paraguay. 83: BOL. 84: BZC (GO, MS, MT), BZE (AL, BA, PE, PI), 

BZL (MG, SP), BZN (RO). 85: PAR. Savannas, cerrados, and rocky places beside streams 

and sandy riverbanks. Map 33. 

83. BOLIVIA: Dept. Santa Cruz, Angel Sandoval, San Matias, R.Guillén 2221 (K, MO). 84. BRAZIL 

NORTH: Rondônia, Rio Madeira, Cachoeira Misericórdia, Ribeirão, G.T.Prance et al. 6722 (INPA, NY, US). 

BRAZIL NORTHEAST: Bahia, Mun. de Bom Jesus da Lapa, Rio das Rãs, G. & M.Hatschbach 55186 

(K, MBM); Piauí, Mun. de São Raimundo Nonato, 1 km SW of Fundação Ruralista, 8–10 km NNE of Curral 

Novo, G.P.Lewis & H.P.N.Pearson 1142 (K, NY). BRAZIL SOUTHEAST: Minas Gerais, Turmalina, Peixe 

Cru, S.T.Silva 636 (BHCB, K). BRAZIL WEST-CENTRAL: Goiás, Rio Corumbá, Mun. de Ipameri, 

G.Hatschbach 38936 (MBM, NY); Mato Grosso, Rio Negro, Fazenda Rio Negro, G.T.Prance & G.B.Schaller 

26302 (CEN, NY); Mato Grosso do Sul, Mun. de Corumbá, Fazenda Santa Catarina, A.Allem 2324 (CEN). 

85. PARAGUAY: Concepción, Paso Horqueta del Río Aquidabán, 48 km N of Concepción, D.R.Brunner et al. 

987 (K, MO, PY); Estancia 3 Hermanas, Potrero Aquidabán, I.Basualdo 3560 (MO). 

25. Couepia cataractae Ducke, Arq. Serv. Florest. 1(1): 31 (1939) 

T: Brazil North: Pará, Rio Tapajós, Maranhãozinho Cachoeira, A.Ducke RB 15136; holo: RB; iso: IAN, K, U, US. 


Shrub or small tree. Leaves with stipules linear, to 1.5 mm long, membranous, deciduous; 

petioles 3–5 mm long, sparsely pubescent, canaliculate above; lamina oblong, 4–10 × 1.5–5 cm, 

rounded to subcordate at base, rounded to obtuse at apex, glabrous above, densely greyarachnoid 

beneath, usually becoming glabrous with age; primary veins 10–15 pairs, rather 

inconspicuous beneath; midrib prominent beneath, glabrous above except at base. 

Inflorescences terminal and axillary panicles; rachis and branches with short dense grey 

pubescence. Bracts and bracteoles 1.5–5 mm long, ovate, deciduous. Receptacle 5–7 mm 

long, subcampanulate-turbinate, often slightly curved anteriorly, short grey-pubescent on 

exterior, glabrous within except for deflexed hairs at throat; pedicels slender, 6–13 mm long. 

Calyx lobes rounded. Petals 5, white, pubescent, ciliate on margins. Stamens 60–90, inserted 

in a complete circle; filaments very far exserted. Ovary pilose. Style pubescent for 3/4 its 

length. Fruit ovoid, with a verrucose epicarp; mesocarp thin; endocarp thin, fragile, granular, 

sparsely pubescent within. 

Amazonian Brazil. 84: BZN (PA, RO). Rocky places beside rivers. Map 34. 

84. BRAZIL NORTH: Pará, Itapiranga, Rio Tocantins, C.C.Berg & A.J.Henderson 684 (AAU, MG, NY); Pará, 

Rio Xingu, Praia de Parati below Altamira, G.T.Prance et al. 26343 (MG, NY); Pará, Tucuruí, Breu Branco, 

Rio Tocantins, J.Revilla et al. 8666 (INPA); Pará, Rio Tapajós, Cachoeira do Acará, B.G.S.Ribeiro 277 (IAN); 

Rondônia, Rio Urupá, M.R.Cordeiro 468 (IAN, INPA). 

26. Couepia macrophylla Spruce ex Hook.f., in C.F.P.Martius, Fl. Bras. 14(2): 41 (1867) 

T: Peru: Loreto (in Monte Guayrapurina), R.Spruce 4049; lecto: K, fide G.T.Prance, Fl. Neotrop. Monogr. 

9: 233 (1972); isolecto: OXF. 

Couepia speciosa Pilg., Bot. Jahrb. Syst. 37:539 (1906). T: Peru: A.Weberbauer 4649; holo: B, destroyed; 

photo: F. 

Illustration: J.D.Hooker, in C.F.P.Martius, Fl. Bras. 14(2), t. 13 (1867). 

29

30 



Tree, small. Leaves with stipules 15–20 mm long, linear, caducous; petioles thick, 8–15 mm long, 

terete, sparsely pubescent, with 2 medial sessile glands; lamina oblong to oblong-elliptic, 

19–22 × 6–12 cm on flowering branches (much larger on sterile branches, up to 40 × 14 cm), 

glabrous above, densely rufous-to-grey arachnoid beneath, rounded to cordate at base, 

abruptly obtuse to abruptly acuminate at apex with acumen 4–8 mm long; primary veins 26–30 

pairs, prominent beneath, slightly impressed above. Inflorescences dense-flowered racemes; 

rachis with short reddish-brown pubescence. Bracts and bracteoles ovate, to 15 mm long, 

persistent. Receptacle 20–25 mm long, cylindrical, with short reddish-brown pubescence on 

exterior, glabrous within except for deflexed hairs at throat; pedicels 2–3.2 mm long. Calyx 

lobes rounded. Petals 5, persistent, sparsely ciliate at margins. Stamens numerous (c. 280), 

inserted in a complete circle, in many rows. Ovary villous. Style densely pubescent 

throughout. Fruit (when young) ellipsoid, 4 × 1.5 cm; exocarp smooth, glabrous. 

W and C Amazonia, Panama. 80: PAN. 83: ECU, PER. 84: BZN (AC, AM). Non-flooded 

forest. Map 35. 

80. PANAMA: Darien, Río Ucurganti, N.Bristan 1121 (MO, NY). 83. ECUADOR: Napo, P. N. Yasuní, 

A.Dik 881 (K, MO); Sucumbios, Gonzalo Pizarro Cantón, Campo Bermejo 6 Norte, C.E.Cerón et al. 9193 

(K, MO, QCNE). PERU: Amazonas, Valle del Río Santiago, 65 km N of Pinglo, V.Huashikat 1591 (MO); 

Loreto, Maynas, road Iquitos-Nauta, R.Vásquez & N.Jaramillo 10579 (K, MO). 84. BRAZIL NORTH: Acre, 

Tarauacá, Rio Muru, Seringal Vitória Velha, M.Silveira et al. 934 (K, NY). 

27. Couepia krukovii Standl., Field Mus.Nat Hist., Bot. Ser. 17(3): 250 (1937) 

T: Brazil North: Amazonas, Mun. de Manicoré, near Bella Vista, B.A.Krukoff 6208; holo: F; iso: A, BM, BR, 

K, MO, NY, RB, U. 


Tree, small to medium-sized. Leaves with stipules linear, to 1.5 cm long, caducous; petioles 

thick, 3–6 mm long, glabrous, eglandular, terete; lamina oblong-oblanceolate, 22–33 × 6–7.5 cm, 

rounded to cordate at base, acuminate at apex with acumen 7–10 mm long, glabrous on both 

surfaces except when very young; midrib prominent above, glabrous when mature; primary 

veins 17–23 pairs, prominent beneath, slightly impressed above. Inflorescences denselyflowered 

terminal and axillary spicate racemes; rachis with short tomentellous brown 

pubescence. Bracts and bracteoles 6–10 mm long, ovate, acute, persistent. Receptacle 

subsessile, cylindrical, 2.5 cm long, slightly curved, with short dense brown pubescence on 

exterior, glabrous within except for deflexed hairs at throat. Calyx lobes rounded, 1 cm long. 

Petals 5, ciliate on margins. Stamens numerous (c. 255), inserted in a complete circle. Ovary 

villous. Style pubescent for 2/3 its length. Fruit unknown. 

Central parts of the basins of the Rios Madeira and Purus, extending into Bolivia and 

Colombia. 83: BOL, CLM. 84: BZN (AM, RO). Confined to várzea forests. Map 36. 

83. BOLIVIA: Dept. Beni, Cachuela Esperanza, Río Beni, G.Meyer 46 (Z). COLOMBIA: Amazonas, 

Araracuara, Villazal, Río Caquetá, C.Londoño et al. 881 (NY). 84. BRAZIL NORTH: Amazonas, Rio Purus 

between Redenção and Itaboca, G.T.Prance et al. 16315 (INPA, K, NY, U, US); Rondônia, Rio Madeira, 

Calama, M.Goulding 183 (MG). 

28. Couepia latifolia Standl., Field Mus. Nat. Hist., Bot. Ser. 17(3): 251 (1937) 

T: Brazil North: Amazonas, Mun. de Manicoré, near Bella Vista, B.A.Krukoff 6005; holo: F; iso: A, BM, BR, 

K, MO, NY, RB, US. 


Tree. Leaves with stipules lanceolate, c. 3 mm long, subpersistent; petioles 4–9 mm long, 

pubescent, with 2 medial glands, terete; lamina oblong-ovate, 17–24 × 5–12 cm, glabrous 

above, sparsely grey-arachnoid or glabrous beneath, rounded at base, rounded or with short 

and blunt acumen at apex; primary veins 16–19 pairs, prominent beneath, slightly impressed 

above. Inflorescences axillary racemes; rachis tomentellous. Bracts and bracteoles 4–8 mm long, 

ovate, acute, persistent, tomentellous on exterior. Receptacle cylindrical, sessile, 1.5–2.5 cm 

long, with dense light-brown pubescence on exterior, glabrous within except for deflexed


hairs at throat. Calyx lobes rounded. Petals 5, persistent, ciliate on margins. Stamens 

numerous, inserted in a complete circle. Ovary villous. Style densely pubescent for 3/4 its 

length. Fruit ovoid, 5–9 cm long, 1.5–5.5 cm broad; epicarp smooth, glabrous; mesocarp 

thick, fleshy; endocarp thin, of a granular texture, fragile, glabrous within. 

Basins of the Rios Purus and Madeira. 83: BOL, CLM, PER. 84: BZN (AC, AM). Forest in 

periodically flooded areas. Map 37. 

83. BOLIVIA: Dept. Beni, Prov. Vaca Diez, 17 km S of Riberalta, D.C.Daly et al. 6254 (K, NY). 

COLOMBIA: Vaupés, Est. Biol. Caparú, S.Defler 103 (MO). PERU: Madre de Dios, Cocha Cashu Biol. 

Station, A.Gentry 43576 (MO, NY). 84. BRAZIL NORTH: Acre, Sena Madureira, Rio Macauã, Provenir, 

D.C.Daly et al. 8118 (K, NY); Amazonas, Boca do Acre, G.T.Prance et al. 2339 (INPA, NY). 

29. Couepia ovalifolia (Schott) Benth. ex Hook.f., in C.F.P.Martius, Fl. Bras. 14(2): (1867), 

as ovatifolia 

Chrysobalanus ovalifolius Schott, in C.P.J.Sprengel, Syst. Veg. 4 (Cur. Post.): 406 (1827). T: Brazil 

Southeast: Rio de Janeiro, Tijuró, H.W.Schott s.n.; holo: W; iso: K, NY. 

Couepia hypoleuca Miq., Linnaea 19: 438 (1847), cited as Moquilea hypoleuca in Index Kewensis. T: Brazil 

Southeast: Rio de Janeiro, P.Claussen 2023; holo: L; iso: C, G, MO, P, U, W, WU, Z. 


Shrub, dwarf to medium-sized tree. Leaves with stipules to 2 mm long, caducous; petioles 

4–7 (–10) mm long, pubescent when young, shallowly canaliculate; lamina ovate, 4–10.5 × 

2.5–7 cm, rounded to cordate at base, short-acuminate at apex with acumen 1–7 mm long, 

glabrous above, densely silver-grey-arachnoid beneath, sometimes becoming glabrous with 

age; midrib prominent beneath, pubescent above when young; primary veins 11–15 pairs, 

prominent beneath, impressed above. Inflorescences terminal panicles; rachis and branches 

with short light brown pubescence. Bracts and bracteoles oblong-ovate, 3–6.5 mm long, 

caducous. Receptacle subcampanulate, 4–5 mm long, densely light-brown-pubescent on 

exterior, glabrous within except for deflexed hairs at throat; pedicels 1.5–4 mm long. Calyx 

lobes rounded, 3–4 mm long. Petals 5, white, ciliate on margins. Stamens 16–19, inserted in 

an arc of 240°, with a sterile portion opposite. Ovary densely pilose. Style pubescent for 1/2 

its length. Fruit elliptic; epicarp smooth, glabrous or softly pubescent when young; mesocarp 

thin, fleshy; endocarp thin, fragile, granular, glabrous within. 

On the coast of Brazil from Pernambuco to Rio de Janeiro. 84: BZE (BA, PE), BZL (ES, MG, 

RJ). Restingas, littoral forest, sand plains, and sand bars. Flowering Aug.–Nov. Map 38. 

84. BRAZIL NORTHEAST: Bahia, Mun. de Ilhéus, 2 km NE of Banco da Vitória, W.W.Thomas et al. 10716 

(K, NY); Bahia, Mun. de Maraú, 6 km on road to Ibaitaba, A.M de Carvalho et al. 3280 (CEPEC, K); 

Pernambuco, Mun. de Olinda, Rio Doce near Recife, G.T.Prance & D.A.Lima 4810 (NY). BRAZIL 

SOUTHEAST: Espírito Santo, Reserva Florestal da CVRD, Linhares, G.L.Farias 73 (K); Rio de Janeiro, 

Pedra de Itaúna, M.B.Casari 277 (GUA); Rio de Janeiro, restinga da Barra da Tijuca, G.Martinelli 80 (RB). 

30. Couepia leitaofilhoi Prance, Kew Bull. 54: 108 (1999) 

T: Brazil Southeast: São Paulo, Mun. de Ubatuba, Picinguaba, A.Furlan 762; holo: HRCB; iso: K, NY. 


Map: G.T.Prance loc.cit.: 106. 

Tree. Branches glabrous when young, conspicuously lenticellate. Leaves with stipules minute, 

interpetiolar, caducous; petioles 8–11 mm, glabrous, weakly canaliculate above, eglandular; 

lamina oblong, chartaceous, 9–17 × 3.5–7.5 cm, acuminate at 7–12 mm long apex, rounded 

and slightly unequal at base, glabrous above, with a short brown, appressed-lanate 

pubescence beneath; midrib prominulous above, prominent and almost glabrous beneath; 

primary veins 11–13 pairs, plane above, prominulous beneath. Inflorescences of terminal and 

axillary panicles; rachis and branches brown-tomentellous. Receptacle cylindrical, 6–7 mm 

long, brown-tomentellous on exterior, glabrous within except for deflexed hairs on throat; 

pedicels 0.5 mm long. Calyx lobes 5, acute, brown-tomentellous on exterior. Petals 5, white; 

margins ciliate. Stamens 19–21, inserted around complete circle. Ovary lanate. Style hirsute 

for 2/3 of length. Fruit ovoid, 3 cm long, 2.5 cm wide; exocarp smooth, glabrous. 

31

32 


Brazil (São Paulo). 84: BZL (SP). Forest. Map 39. 

84. BRAZIL SOUTHEAST: São Paulo, Mun. de Ubatuba, Picinguaba, F.C.P.Garcia et al. 411 (HRCB, K); 

São Paulo, Mun. de Ubatuba, Picinguaba, F.C.P.Garcia et al. 621 (HRCB, K); São Paulo, Mun. de Ubatuba, 

Picinguaba, J.E.L.S.Ribeiro et al. 639 (HRCB, K). 

31. Couepia schottii Fritsch, Ann. K. K. Naturhist. Hofmus. 5:13 (1890) 

Chrysobalanus macrophyllus Schott, in C.P.J.Sprengel, Syst. Veg. 4(2): 407 (1827), non Couepia macrophylla 

Spruce ex Hook.f. (1867). T: Brazil Southeast: Rio de Janeiro, H.W.Schott s.n.; holo: W; iso: NY. 


Tree to 25 m tall. Leaves with stipules c. 2 mm long, caducous; petioles 12–15 mm long, 

sparsely pubescent, shallowly canaliculate; lamina oblong, 10–18 × 4–7 cm, subcordate at 

base, rounded to short-acuminate at apex with acumen 1–3 mm long, glabrous above, densely 

grey-brown-arachnoid beneath; primary veins 16–20 pairs, prominent beneath, impressed 

above. Inflorescences terminal panicles; rachis and branches with short light brown 

pubescence. Bracts and bracteoles ovate, acute, to 4 mm long, caducous. Receptacle 

subcampanulate, 5–6 mm long, with short light brown pubescence on exterior, glabrous 

within except for deflexed hairs at throat; pedicels 2 mm long. Calyx lobes rounded. Petals 5, 

ciliate on margins. Stamens 15–16, inserted in a semicircle, with a few small filamentous 

staminodes opposite. Ovary densely villous. Style pubescent for 1/2 its length. Fruit spherical, 

c. 3.5 cm in diam.; epicarp smooth, glabrous; mesocarp thin, fleshy; endocarp thin, granular, 

fragile, glabrous within. 

Brazil (Bahia to Rio de Janeiro). 84: BZE (BA), BZL (ES, RJ). Coastal restingas. Flowering 

Aug.–Nov. Map 40. 

84. BRAZIL NORTHEAST: Bahia, Mun. de Uruçuca, 28 km NE of Uruçuca, S.A.Mori et al. 13065 (CEPEC, 

K, NY). BRAZIL SOUTHEAST: Espírito Santo, Res. Florestal Linhares, J.Spada 148 (INPA, RB); Rio de 

Janeiro, restinga de Grumari, D.Araújo 5261 (GUA); Rio de Janeiro, Mun. de Saquarema, Res. Ecol. Estadual 

Jacarepiá, D.Araújo 9650 (K, GUA); Rio de Janeiro, Mun. de Saquarema, restinga de Massambaba, C.Farney 

1101 (NY, RB). 

32. Couepia grandiflora (Mart. & Zucc.) Benth. ex Hook.f., in C.F.P.Martius, Fl. Bras. 

14(2): 46 (1867) 

Moquilea grandiflora Mart. & Zucc., Abh. Math.-Phys. Cl. Königl. Bayer. Akad. Wiss., 1: 388 (1832). T: Brazil 

West-central: Goiás, C.F.P.Martius s.n.; lecto: M, fide G.T.Prance, Fl. Neotrop. Monogr. 9: 236 (1972). 

Couepia formosana Taub., Bot. Jahrb. Syst. 21: 429 (1896). T: Brazil West-central: Goiás, Formosa, E.Ule 

2881; holo: B, destroyed; iso: HBG. 

Couepia suberosa Pilg., Notizbl. Bot. Gart. Berlin-Dahlem 8: 538 (1923). T: Brazil Southeast: Minas 

Gerais, Rio Pardo, L.Riedel 504; holo: B, destroyed; iso: OXF. 


Tree to 8 m tall. Branches pubescent when young; older branches often with a conspicuous 

thick cork covering c. 4 mm thick. Leaves with stipules to 2 mm long, subulate, caducous; 

petioles 3–13 mm long, pubescent, terete; lamina oblong to ovate, 7.5–18 × 3.5–9 cm, 

cordate to cuneate at base, rounded to bluntly acuminate at apex, glabrous above, or with 

sparse ferrugineous pubescence when young, shortly dense-arachnoid beneath; primary veins 

10–16 pairs, prominent beneath, prominulous above; midrib prominent on both surfaces. 

Inflorescences terminal and axillary panicles; rachis and branches with dense yellow-brown 

pubescence. Bracts and bracteoles 1.5–5 mm long, ovate, membranous, caducous. Receptacle 

campanulate, 5–11.5 cm long, densely cinereous-pubescent, glabrous within except for 

deflexed hairs at throat; pedicels 2–5 mm long. Calyx lobes rounded. Petals 5, white, 

pubescent, ciliate on margins, caducous. Stamens 50–125, inserted in a complete circle. 

Ovary densely villous. Style pubescent for most of its length. Fruit oblong; epicarp smooth, 

glabrous, drying black; pericarp thick, fleshy, glabrous within. 

Brazil, Bolivia and Paraguay. 83: BOL. 84: BZC (DF, GO, MS, MT), BZE (BA, MA, PI), 

BZL (MG, SP), BZN (RO, TO), BZS (PR). 85: PAR. Cerrados and savannas. Flowering 

July–Dec. Map 41.


84. BRAZIL NORTH: Rondônia, Km 16 Vilhena-Colorado, B.W.Nelson 344 (INPA, NY). BRAZIL 

NORTHEAST: Bahia, Mun. de Formosa do Rio Preto, near Rio Richão, Fazenda Lagoa de Fora, B.J.Dias et 

al. 107 (K); Maranhão, Grajahú, A.Lisboa MG2494 (BM, MG, RB); Piauí, banks of Rio Gurguéia, G.Gardner 

2563 (BM, F, GH, K, OXF, P, US, W). BRAZIL SOUTHEAST: Minas Gerais, Lagoa Santa, A.E.Brina & 

L.V.Costa BHCB32757 (BHCB); São Paulo, Mun. de Botucatu, km 27–29 road S Manuel-Sta. Maria da Serra, 

A.Amaral Jr. et al. 3 (BOTU, HRCB, K). BRAZIL WEST-CENTRAL: Distrito Federal, Res. Ecológica IBGE, 

M.L.M.Azevedo et al. 922 (IBGE); Goiás, Mun. de Posse, Serra Geral de Goiás, I. & G.Gottsberger 152-25771 

(BOTU, NY); Mato Grosso do Sul, Mun. de Iguatemi, 5 km W of Iguatemi, R.Callejas et al. 1969 (K, MBM, NY). 

85. PARAGUAY: Dept. Canindeyu, 60 km E of Mbutuy, Estancia Río Corrientes, D.R.Brunner et al. 946 (K, MO). 

33. Couepia elata Ducke, Arq. Inst. Biol. Veg. 2: 35 (1935) 

T: Brazil North: Amazonas, Manaus, A.Ducke RB25001; holo: RB; iso: K, P, U. 


Tree, large. Leaves with stipules minute; petioles 4–6 mm long, shortly pubescent; lamina 

ovate 4–7 × 2.5–3 cm, obtuse to subcuneate at base, caudately acuminate at apex with 

acumen 6–9 mm long, glabrous above, prominently reticulate beneath with silver-grey 

pubescence between reticulations; midrib impressed above, pubescent on lower portion; 

primary veins 6–8 pairs, prominent beneath. Inflorescences terminal and axillary fewflowered 

lax racemes, with short grey-pubescence. Bracts and bracteoles minute, caducous. 

Receptacle obconical-turbinate, 4–5 mm long, sparsely pubescent on exterior, glabrous 

within except for deflexed hairs at throat; pedicels 2.5–3.5 mm long. Calyx lobes acute. 

Petals 5, white, glabrous on margins, deciduous. Stamens 35–40, inserted in 1/2 a circle, with 

a row of short-toothed staminodes opposite. Ovary densely villous. Style glabrous except at 

base. Fruit globose, 5–6 cm diameter; epicarp densely verrucose; mesocarp thick; endocarp 

fragile, thick, granular. 

Guianas, Colombia, W and C Amazonian Brazil. 82: FRG, GUY. 83: CLM. 84: BZN (AC, 

AM). Non-flooded forest. Flowering Mar.–June. Map 42. 

82. FRENCH GUIANA: Sinnamary R., above Petit Saut, S.A.Mori et al. 23651 (K, NY). GUYANA: Cuyuni- 

Mazaruni Region, Pakaraima Mts., Karowrieng R., 1 km SE of Maipuri Falls, B.Hoffman 3036 (K, US). 

83. COLOMBIA: Amazonas, Río Cahuina, M.Sanchez S. et al. 980 (K). 84. BRAZIL NORTH: Amazonas, 

Manaus, Reserva Florestal Ducke, P.A.C.L.Assunção 506 (INPA, K); Amazonas, Rio Camanaú, Mun. de Novo 

Airão, R.Miller 173 (INPA, K); Acre, Cruzeiro do Sul, Estrada Alemanha, G.T.Prance et al. 11943 (INPA, NY). 

34. Couepia racemosa Benth. ex Hook.f., in C.F.P.Martius, Fl. Bras. 14(2): 43(1867) 

T: Brazil North: Amazonas, Falls of São Gabriel, R.Spruce 1776; lecto: K, fide G.T.Prance, Fl. Neotrop. 

Monogr. 9: 238 (1972); isolecto: OXF, P. 

Couepia racemosa var. reticulata Pilg., Verh. Bot. Vereins Prov. Brandenburg 47: 150 (1905). T: Brazil: 

Amazonas, Manaus, E.Ule 6149 (K, MG). 

Couepia calophlebia Standl., Field Mus. Nat. Hist., Bot. Ser. 17(3): 248 (1937). T: Brazil North: 

Amazonas, Rio Embira, B.A.Krukoff 4913; holo: F; iso: A, BM, F, K, M, MO, NY, RB, U, US. 

Illustration: J.D.Hooker, in C.F.P.Martius, Fl. Bras. 14(2), t. 14(1) (1867). 


Tree to 30 m tall. Leaves with stipules c. 1.5 mm long, caducous; petioles 8–12 mm long, 

with 2 sessile glands near to lamina base; lamina elliptic to oblong, 5–18.5 × 2–7.5 cm, 

rounded to cuneate at base, acuminate at apex with acumen 4.5–20 mm long, glabrous above, 

prominently reticulate beneath, or with stomatal cavities, with short appressed grey pubescence; 

midrib prominent beneath, slightly impressed above; primary veins 8–12 pairs, prominent 

beneath, slightly impressed above. Inflorescences little branched densely-flowered panicles; 

rachis and branches with short brown tomentellous pubescence. Bracts and bracteoles ovate to 

oblong, 2–6 mm long, caducous. Receptacle cylindrical-turbinate, 5–7.5 mm long, with a short 

brown pubescence on exterior, glabrous within except for deflexed hairs at throat. Calyx 

lobes rounded to acute. Petals 5, white, exceeding calyx lobes, glabrous on margins. Stamens 

24–30, unilateral, with a smooth glabrous untoothed portion of the circle opposite them. 

Ovary densely villous. Style pubescent at base only. Fruit elliptic, pointed at apex, 3–4.5 × 

2–3.5 cm; epicarp verrucose; pericarp undifferentiated, rather fragile, sparsely pubescent within. 

33

34 


Figure 55. Couepia amaraliae. A, habit; B, leaf lower surface; C, flower bud; D, flower; 

E, ½ flower; F, petal; G, fruit (A–F, G.T.Prance et al. 29261; G, N.A.Rosa & I.Cordeiro 1713). 

Scale bars: A, G = 3 cm; B = 5 mm; C–F = 4 mm. Drawn by Bobbi Angell. Previously 

published in G.T.Prance, Fl. Neotrop. Monogr. 9S: 204 (1989).


C and W Amazonia in Venezuela, Colombia, Peru and Brazil. 82: VEN. 83: CLM, PER. 

84: BZN (AM, PA). Non-flooded forest or on periodically flooded ground, riverine forest 

and river banks, and secondary forest. Flowering Feb.–Aug. Map 43. 

82. VENEZUELA: Amazonas, Dept. Atures, Río Cataniapo, 2 km from San Pedro de Cataniapo, F.Guanchez 

396 (MYF). 83. COLOMBIA: Amazonas, Resguardo Aduche, Correg. de Pto. Santander, D.Cárdenas & 

F.Andoque 4282 (COAH, K). PERU: Loreto, Maynas, Llachapa, Río Napo, R.Vásquez & N.Jaramillo 3762 

(MO, NY). 84. BRAZIL NORTH: Amazonas, Mun. de Tefé, Lago Tefé, Vila Nogueira, I.L.Amaral et al. 56 

(INPA, NY, US); Amazonas, Manaus, Estrada Mauá, G.T.Prance et al. 11631 (INPA, NY); Pará, 25 km up 

Rio Mapuera, D.G.Campbell et al. P22563 (INPA, NY, US). 

35. Couepia amaraliae Prance, Fl. Neotrop. Monogr. 9S: 72 (1989), as amaralae 

T: Brazil North: Amazonas, base of Serra Aracá, 3 km E of Rio Jauari, G.T.Prance et al. 29261; holo: INPA; 

iso: NY. 



Shrub 1–4 m tall. Branches appressed lanate pubescent when young, becoming glabrous with 

age. Leaves with stipules caducous; petioles 4–5 mm long, lanate when young, rugulose, 

terete, eglandular; lamina oblong to oblong-elliptic, coriaceous, 2.5–6 × 1.5–3.2 cm, rounded 

at base, acute or bluntly acuminate at apex, glabrous above, prominently reticulate beneath 

with conspicuous deep stomatal cavities filled with short grey pubescence; midrib prominent 

and sparsely lanate beneath, slightly impressed above; primary veins 9–10 pairs, prominent 

and lanate beneath, plane to slightly impressed above. Inflorescences of few-branched 

panicles or racemes, axillary and terminal; rachis and branches brown-tomentellous. Bracts 

and bracteoles ovate, caducous, tomentellous on both surfaces. Receptacle campanulateturbinate, 

3–4 mm long, tomentellous on exterior, glabrous within except for deflexed hairs 

at throat. Calyx lobes 5, acute, 2 mm long, tomentellous on both surfaces. Petals 5, white, 

glabrous. Stamens 35–40, inserted around 1/2 of circle. Ovary densely villous. Style glabrous 

except for deflexed hairs at base. Fruit globose, 5 cm diam.; exocarp glabrous, lenticellate; 

pericarp 2–2.5 mm thick, hard, woody, tomentose within. Fig. 55. 

Brazil (Amazonas). 84: BZN (AM). Sandy soil forests (campina and campinarana). Map 44. 

84. BRAZIL NORTH: Amazonas, 3 km S of central part of Serra Aracá, M.G.Silva et al. 7121 (INPA, NY); 

Amazonas, 3 km S of central part of Serra Aracá, W.A.Rodrigues et al. 10509 (INPA, NY); Amazonas, Rio 

Aracá near mouth of Rio Jauari, I.Cordeiro 126 (INPA, SP); Amazonas, Rio Aracá, R.L.Fróes & G.Addison 

29277 (IAN, NY); Amazonas, Rio Jauari near Ipsilon junction, G.T.Prance et al. 29420 (INPA, NY). 

36. Couepia martinii Prance, Fl. Neotrop. Monogr. 9: 238 (1972) 

T: French Guiana: Cayenne, J.Martin s.n.; holo: NY; iso: BM, E-GL, F, K. 



Tree. Branches glabrous when young, lenticellate. Leaves with stipules small, caducous; 

petioles 10–12 mm long, canaliculate; lamina elliptic, 10.5–25 × 5–10.5 cm, acuminate at 

apex with acumen 4–8 mm long, cuneate at base, glabrous above, appressed grey-arachnoidtomentose 

beneath; midrib prominulous above, prominent beneath; primary veins 10–13 pairs, 

slightly impressed above, prominent beneath. Inflorescences of terminal spikes, denseflowered; 

rachis ferrugineous-tomentose. Bracts and bracteoles large, ovate, to 17 mm long, 

acute at apex. Flowers 25–35 mm long. Receptacle cylindrical, 20–25 mm long, browntomentose 

on exterior, glabrous within except for ring of deflexed hairs around throat. Calyx 

lobes acute. Petals 5, glabrous, deciduous. Stamens c. 65, inserted around complete circle. 

Ovary villous. Style densely pilose. Fruit not seen. 

French Guiana. 82: FRG. Forest on well drained soil. Map 45. 

82. FRENCH GUIANA: Monte Chauve, G.Cremers & F.Crozier 15093 (CAY, ULM, US). 

35

36 


37. Couepia bondarii Prance, Fl. Neotrop. Monogr. 9: 240 (1972) 

T: Brazil Northeast: Bahia, without precise locality, G.Bondar 1863; holo: SP. 



Tree, small. Branches glabrous when young. Leaves with stipules lanceolate, membranous, to 

30 mm long, caducous; petioles 6–10 mm long, terete, tomentellous; lamina oblong-lanceolate, 

15–25 × 3.5–7 cm, acuminate at apex with acumen c. 5 mm long, subcuneate at base, 

glabrous above, densely rufous-brown-arachnoid pubescent beneath; midrib slightly prominent 

and glabrous above; primary veins 17–25 pairs, plane above, prominent beneath. 

Inflorescences of terminal and axillary fasciculate racemes; rachis ferrugineous-tomentellous. 

Bracts and bracteoles large, ovate, persistent, to 25 mm long, densely brown-puberulous on 

exterior, sparsely puberulous within. Flowers 20–30 mm long. Receptacle tubular, sessile, 

12–15 mm long, rufous-tomentellous on exterior, glabrous within except for ring of deflexed 

hairs around throat. Calyx lobes acute, rufous-tomentellous, 10–16 mm long. Petals 5, 

sericeous on outer surfaces. Stamens c. 65, inserted around complete circle, connate at base 

for 1.5 mm. Ovary pilose. Style tomentose. Fruit ellipsoid, 7 × 4.5 cm; exocarp smooth, 

glabrous; mesocarp c. 4 mm thick when dry; endocarp thin, bony, glabrous within. 

Known only from 2 collections, one from the state of Bahia, Brazil, and the other without 

any locality. 84: BZE (BA). Habitat unknown. Map 46. 

84. BRAZIL NORTHEAST: Bahia, Mun. de Uruçuca, Res. Biológica Gregório Bondar, A.M. de Carvalho et al. 

3249 (CEPEC, K). 

38. Couepia insignis Fritsch, Ann. K. K. Naturhist. Hofmus. 5: 11 (1890) 

T: Brazil Northeast: Bahia, J.S.Blanchet 3209; holo: W; iso: G, P. 


Tree. Leaves with stipules linear, to 15 mm long, membranous, caducous; petioles 8–11 mm 

long, sparsely pubescent, terete; lamina oblong, 15–24 × 8–9.5 cm, rounded at apex, rounded 

to cordate at base, glabrous above, densely grey-arachnoid beneath; primary veins 21–25 

pairs, impressed above, prominent beneath, arcuate and anastomosing near the lamina 

margin. Inflorescences few-flowered racemes; rachis with short rufous pubescence. Bracts 

and bracteoles persistent, 10–28 mm long, oblong, acuminate, membranous. Receptacle 

subcylindrical, 8–9 mm long, with short brown pubescence on exterior, glabrous within 

except for deflexed hairs at throat; pedicels 1–2 mm long. Calyx lobes oblong-lanceolate, 

10–12 mm long, acute. Petals 5, ciliate on margins. Stamens numerous (over 50). Ovary 

villous. Style pubescent for 3/4 its length. Fruit (section only seen) 7.5 cm diam.; exocarp 

glabrous, wrinkled when dry; mesocarp fleshy, 4–10 mm thick when dry; endocarp c. 1 mm 

thick, fibrous, glabrous within. 

Brazil (Bahia). 84: BZE (BA). Coastal forest. Map 47. 

84. BRAZIL NORTHEAST: Bahia, Mun. de Una, km 27 Ilhéus-Una, A.M. de Carvalho & G.P.Lewis 858 

(CEPEC, K, NY); Bahia, Mun. de Ilhéus-Olivença, Km 6 Olivença-Maruim, A.Amorim et al. 481 (CEPEC, K, NY); 

Bahia, restinga de Olivença, L.E. de Mello Filho 3061 (CEPEC). 

39. Couepia cidiana Prance, Acta Amazon. 13(1): 21 (1983) 

T: Brazil North: Pará, Mun. de Oriximiná, Rio Parú do oeste, left margin of Cachoeira Chuvisco, C.A. Cid Ferreira 

et al. 2261; holo: INPA96602; iso: FHO, NY. 


Treelet 8–10 m tall. Branches tomentose when young. Leaves with stipules linear, 

membranous, 2–2.5 cm long, persistent; petioles 5–7 mm long, terete, eglandular, densely 

tomentose; lamina oblong-lanceolate, coriaceous, 15–26 × 7–9.5 cm broad, subcordate at base, 

acuminate at apex with acumen 7–10 mm long, glabrous above, densely grey short-lanate 

beneath with contrasting brown-tomentose venation; midrib prominulous above, tomentellous 

when young; primary veins 26–35 pairs, lightly impressed above, prominent and tomentose


beneath, anastomosing c. 3 mm from margin to form a conspicuous marginal vein; secondary 

veins ±parallel, prominulous. Inflorescences few-flowered racemes; rachis tomentose. Bracts 

and bracteoles persistent, 2–3.6 cm long, tomentose on exterior, glabrous within, ovate, 

acuminate. Flowers 60–75 mm long. Receptacle cylindrical, 3–4 cm long, densely velutinoustomentellous 

on exterior, tomentose within to base, sessile. Calyx lobes 5, triangular, acute, 

tomentose on exterior, tomentose within except for glabrous lower portion, 10–13 mm long. 

Petals 5, white, glabrous on exterior; margins ciliate. Stamens c. 165, inserted around a 

complete circle, fused at base to form a ring c. 5 mm tall with stamens inserted in several 

rows on exterior of ring; interior of ring densely tomentose. Ovary inserted near mouth of 

tube, densely villous. Style sparsely hirsute pubescent on lower 1/3. Fruit not seen. 

Brazil (western Pará). 84: BZN (PA). Forest on terra firme. Map 48. 

84. BRAZIL NORTH: Pará, Rio Cachorro, Serra do Cachorro, Mun. de Oriximiná, 16 km NW of Cachoeira 

Porteira, G.Martinelli 7052 (INPA, NY, RB). 

40. Couepia recurva Spruce ex Prance, Fl. Neotrop. Monogr. 9: 242 (1972) 

T: Ecuador: 'In Andibus Ecuadorensibus', R.Spruce 5067; lecto: NY, fide G.T.Prance, Fl. Neotrop. Monogr. 9: 

242 (1972); isolecto: BM, CGE, K, LE, OXF, W. 

Illustrations: G.T.Prance, Fl. Neotrop. Monogr. 9: 243 (1972); Fl. Ecuador 10: 13 (1979). 


Tree 10–20 m. Branches rufous-pubescent when young, lenticellate. Leaves with stipules to 

3 mm long, caducous; petioles c. 2 mm long, hirsute, terete; lamina oblong-ovate, 5.5–7.5 × 

3–4 cm, acuminate at apex with acumen to 9 mm long, rounded at base, sparsely hirsute on 

both surfaces; primary veins 10–12 pairs, prominent beneath. Inflorescences of axillary 

racemes; rachis rufous-brown-hirsute. Bracts and bracteoles caducous (not seen). Receptacle 

cylindrical, 20–25 mm long, rufous-brown-hirsute on exterior, densely tomentose to base 

within; pedicels 5–7 mm long. Calyx lobes rounded. Petals 5, obtuse, deciduous; margins 

glabrous. Stamens c. 17, unilateral; filaments hirsute. Ovary densely pubescent. Style pilose. 

Fruit not seen. Fig. 56. 

Ecuador and Peru. 83: ECU, PER. Andean primary forest, to 1700 m. Map 49. 

83. ECUADOR: Tungurahua, La Victoria, valley of Río Pastaza, E.Asplund 10055 (S); Napo, El Charo 

Cantón, Río Quijos, at Coca Hydroelectric Project, W.Palacios 5956 (K, MO, QCNE); Zamora-Chinchipe, 

Zamora, P. N. Podocarpus, Guardería Río Bombuscaro, W.Palacios & M.Tirado 13264 (K, MO, QCNE); 

Zamora-Chinchipe, Zamora, P. N. Podocarpus, Guardería Río Bombuscaro, W.Palacios & M.Tirado 13304 (K, MO, 

QCNE). PERU: Cajamarca, San Ignacio Prov., Huarango, Nuevo Mundo, J.Campos & S.Nuñez 4605 (K, MO). 

Recently collected in Peru but previously known only from the type collection and from 

Tungurahua and Zamora in the high Andes of Ecuador. 

41. Couepia obovata Ducke, Arq. Inst. Biol. Veg. 2: 35 (1935) 

T: Brazil North: Amazonas, Manaus, Estrada do Aleixo, A.Ducke RB24996; holo: RB; iso: IAN, K, P, U, US. 


Tree, small to medium-sized. Leaves with stipules 1–2 mm long, lanceolate, caducous; petioles 

3–7 mm long; lamina obovate, 4–10 × 2–5 cm, subcuneate at base, bluntly acuminate 

at apex with acumen 2–5 mm long, glabrous above, with a sparse covering of appressed grey 

hairs beneath; primary veins 8–10 pairs, prominent beneath. Inflorescences terminal and 

axillary racemes, 3–6 cm long; rachis with sparse silver-grey pubescence. Bracts and bracteoles 

1.5–3.5 mm long, ovate, persistent. Receptacle cylindrical, 7–9 mm long curved, with a few 

short appressed hairs on exterior, glabrous within except for deflexed hairs at throat, 

subsessile. Calyx lobes rounded. Petals 5, white, ciliate on margins. Stamens 16–21, inserted in 

a complete circle. Ovary lanate. Style glabrous except at base. Fruit ovoid, 2–3 × 1.5–2.5 cm; 

epicarp smooth, glabrous; endocarp thick, hard, of a granular texture, glabrous within. 

Guianas, Venezuela and central, Colombian, Peruvian and Ecuadorean Amazonia. 82: FRG, 

GUY, SUR, VEN. 83: CLM, ECU, PER. 84: BZN (AM, PA). Non-flooded forest. Map 50. 

37

38 


Figure 56. Couepia recurva. A, habit; B, leaf lower surface; C, flower; D, flower section; 

E, petal; F, ovary and style; G, ovary section; H, diagrammatic view of stamen and style 

base (A–H, R.Spruce 5067). Scale bars: A = 5 cm; B–E = 1 cm; F, H = 3 mm; G = 1 mm. 

Drawn by R.A.J. after Charlie Clare. Previously published in G.T.Prance, Fl. Neotrop. Monogr. 

9: 243 (1972).


82. FRENCH GUIANA: Saül, La Fumée Mountain Trail, S.A.Mori & A.E.Hartley 18552 (CAY, NY, US). 

GUYANA: Potaro-Siparuni Region, Kaieteur Falls N. P., C.L.Kelloff 926 (K, US). SURINAME: lower slopes 

of Juliana Top, Wilhelmina Gebergte, H.S.Irwin et al. 54802 (F, MO, NY, P, US). VENEZUELA: Amazonas, 

Dept. Atures, Puerto Ayacucho, Río Cataniapo, A.Castillo 2208 (NY). 83. COLOMBIA: Vaupés, Rio 

Apaporis, Jinogoje, R.E.Schultes & I.Cabrera 15662 (NY, US). ECUADOR: Napo, Cantón Tena, Est. Biol. 

Jatun Sacha, 8 km E of Misahualli, W.Palacios 4164 (K, MO). PERU: Huánuco, Prov. Pachitea, Sira Mts. near 

Pucallpa, W.Morawetz & B.Wallnöfer 15-24888 (K, W); Loreto, Maynas, Mishana, R.Vásquez et al. 3997 

(MO, NY). 84. BRAZIL NORTH: Amazonas, Rio Ituxi, W of Boca do Curuquetê, G.T.Prance et al. 14095 

(INPA, K, MG, NY, S, U, US); Amazonas, Manaus, Reserva Florestal Ducke, M.A.S.Costa 350 (INPA, K). 

42. Couepia williamsii J.F.Macbr., Candollea 5: 366 (1934) 

T: Peru: Loreto, Río Amazonas, La Victoria, Ll.Williams 2975; holo: F. 


Tree, small. Leaves with stipules to 3 mm long, linear, caducous; petioles 5–7 mm long, with 

a few stiff erect hairs, shallowly canaliculate above, with 2 inconspicuous glands at lamina 

base; lamina oblong to oblong-lanceolate, 9–17 × 2.5–5 cm, cuneate to subcuneate at base, 

abruptly acuminate at apex with acumen 12–22 mm long, glabrous above, densely appressedlanate 

beneath; primary veins 12–15 pairs, prominent beneath; midrib prominent above. 

Inflorescences terminal and axillary few-flowered racemes; rachis sparsely puberulous. 

Bracts and bracteoles to 3 mm long, early caducous. Receptacle slender, tubular, 14–19 mm 

long, curved, glabrous on exterior or with a few appressed hairs, glabrous within except for 

deflexed hairs at throat; pedicels 3–5 mm long. Calyx lobes rounded. Petals 5, white, ciliate 

on margins. Stamens c. 70, inserted in a complete circle. Ovary villous. Style pubescent for 

2/3 its length. Fruit globose, c. 6.5 cm in diameter; epicarp glabrous, verrucose; mesocarp 

thick, fleshy; endocarp thin, fragile, granular, glabrous within. 

Western Amazonia. 83: CLM, PER. 84: BZN (AC, AM). Forest on non-flooded and 

periodically flooded ground. Flowering Aug.–Nov. Map 51. 

83. COLOMBIA: Amazonas, Río Igara-Parana, trib. of Río Putumayo, C.Sastre 3273 (COL, P). PERU: 

Loreto, Maynas, Río Nanay, between Iquitos and Santa Maria de Nanay, A.Gentry et al. 36440 (MO, NY); 

Loreto, Caserío Mishana, Río Nanay, J.Ruíz & H.Murphy 107 (K, MO). 84. BRAZIL NORTH: Acre, 

Cruzeiro do Sul, Rio Juruá, Igarapé Viseu, C.A. Cid Ferreira et al. 10599 (INPA, K, NY); Amazonas, Rio 

Javari, above Paumari, G.T.Prance et al. 23816 (INPA, NY, US); Amazonas, Boca do Acre, G.T.Prance et al. 

2484 (INPA, NY, US). 

43. Couepia glabra Prance, Acta Amazon. 2(1): 10 (1972) 

T: Brazil North: Amazonas, Rio Cuieiras, just below mouth of Rio Branquinho, G.T.Prance, D.Coelho & 

O.P.Monteiro 1494; holo: NY; iso: FHO, INPA. 


Tree 10 m tall. Branches glabrous when young. Leaves with stipules lanceolate, membranous, 

c. 2 mm long, caducous; petioles 8–12 mm long, glabrous, canaliculate, rugose, eglandular; 

lamina oblong to oblong-lanceolate, coriaceous, 11–21 × 4–7 cm; base subcuneate; apex 

acuminate with acumen 5–9 mm long, usually curved; glabrous on both surfaces; midrib 

prominent on both surfaces; primary veins 16–17 pairs, prominulous or plane above, 

prominent beneath. Inflorescences of terminal and axillary panicles, 4.5–9 cm long, or rarely 

of unbranched racemes; rachis and branches with minute sparse pubescence. Bracts and 

bracteoles caducous. Receptacle cylindrical-turbinate, 15–20 mm long, sparsely puberulous 

in bud, soon becoming glabrous, glabrous within except for dense deflexed hairs around 

throat. Calyx lobes 5, sparsely puberulous when young, soon becoming glabrous; margins 

ciliate. Petals 5, glabrous, caducous. Stamens c. 110, inserted around a complete circle; 

filaments tomentose around base, glabrous for most of length. Ovary pilose. Style pilose at 

base, glabrous above. Fruit unknown. 

Brazil (lower Rio Negro region). 84: BZN (AM). Forest on terra firme. Map 52. 

84. BRAZIL NORTH: Amazonas, Manaus-Caracaraí Road Km 115, Rio Urubu, G.T.Prance et al. 21636 

(INPA, NY). 

39

40 


44. Couepia marleneae Prance, Acta Amazon. 4(1): 17 (1974) 

T: Brazil North: Amazonas, Manaus-Porto Velho Rd., between Rios Castanho and Tupana, M.F. da Silva et al. 

822; holo: INPA; iso: NY, RB. 


Tree to 8 m tall. Branches sparsely puberulous when young, soon becoming glabrous. Leaves 

with stipules triangular, 1 mm long; petioles 4–7 mm long, sparsely puberulous or glabrous, 

canaliculate above; lamina oblong-lanceolate, coriaceous, 5–8 × 1.5–2.4 cm, subcuneate at base, 

acuminate at apex with acumen 5–15 mm long, with 2 glands towards base at junction with 

petiole, glabrous above, with sparsely lanate pubescence beneath; midrib glabrous and 

prominulous above, prominent beneath; primary veins 7–10 pairs, prominent beneath. 

Inflorescences of terminal racemes; rachis grey-puberulous. Bracts small and caducous. 

Receptacle cylindrical, swollen towards base, 15–18 mm long, sparsely puberulous on exterior, 

±glabrous within but densely pilose around base of stamens. Calyx lobes 5, rounded, 

sparsely puberulous on exterior; margins ciliate. Petals 5, white; margins ciliate. Stamens 

numerous, inserted around a complete circle. Ovary densely pilose. Style pilose for 1/2 of length. 

Fruit when young ovoid; exocarp smooth, glabrous; mesocarp fleshy; endocarp hard, fragile. 

Brazil, known only from south of Manaus. 84: BZN (AM). Forest on terra firme. Map 53. 

84. BRAZIL NORTH: Amazonas, between Rio Araçá, and Lago Castanho, Manaus-Porto Velho road, M.F. da 

Silva 467 (INPA, NY); Amazonas, between Rio Araçá, and Lago Castanho, Manaus-Porto Velho road, M.F. da 

Silva 490 (INPA, NY); Amazonas, Mun. de Nova Olinda, Rio Paca, trib. of Rio Mari Mari, C.Todzia 2295 

(INPA, NY); Amazonas, Mun. de Maués, Rio Urupadi, 4 km from mouth of Igarapé Quininha, J.L.Zarucchi 

3079 (INPA, NY). 

45. Couepia joaquinae Prance, Kew Bull. 46: 107 (1991) 

T: Brazil North: Amapá, Mun. de Mazagão, Morros do Felipe, R. F. Felipe, M.A.Conceição 176; holo: MG; iso: K. 


Tree 20–30 m tall. Branches glabrous when young. Leaves with stipules triangular, 

membranous, 1.5–2 mm long, early caducous; petioles 7–12 mm long, terete, eglandular, 

glabrous; lamina oblong to oblong-lanceolate, chartaceous, 10–15 × 3–5.5 cm broad, cuneate 

at base, acuminate at apex with acumen 8–15 mm long, often slightly curved, glabrous on 

both surfaces; midrib plane above, prominent and glabrous beneath; primary veins 10–14 

pairs, plane above, prominulous beneath. Inflorescence of terminal and axillary panicles or 

rarely racemes; flowers borne in small groups of 2–4 along rachis; rachis and branches 

glabrous or sparsely puberulous. Bracts and bracteoles minute, lanceolate, membranous, 

caducous, sparsely puberulous. Receptacle narrowly cylindrical, almost glabrous and striate 

when dry on exterior, glabrous within except for few hairs at base. Calyx lobes 5, acute, 

sparsely puberulous on exterior; margins ciliate. Petals 5, white, glabrous. Stamens 9–11, 

unilateral; filaments exserted, glabrous. Ovary lanate. Style glabrous except for few hairs at 

base. Fruit ovoid, 3.5–4.5 × 1.9–2.4 cm; exocarp sparsely and inconspicuously lenticellate; 

mesocarp 1.5 mm thick, fleshy; endocarp 0.5 mm thick, hard, fibrous, glabrous within. 

French Guiana and Brazil (Amapá). 82: FRG. 84: BZN (AP). Forest on terra firme on a 

table-top hill on clay oxisol. Map 54. 

82. FRENCH GUIANA: Piste de Saint Elie, D.Sabatier & M.F.Prévost 3764 (CAY, K). 84. BRAZIL 

NORTH: Amapá, Mun. de Mazagão, Morros do Felipe, H.T.Beck et al. 37 (K, MG, NY); Amapá, Mun. de 

Mazagão, Morros do Felipe, M.A.Conceição 175 (K, MG). 

46. Couepia chrysocalyx (Poepp. & Endl.) Benth. ex Hook.f., in C.F.P.Martius, Fl. Bras. 

14(2): 42 (1867) 

Moquilea chrysocalyx Poepp., in E.F.Poeppig & S.F.Endlicher, Nov. Gen. Sp. Pl. 3: 75 (1845). T: Peru: 

Amazonas, Maynas, E.Poeppig 2306; lecto: W, fide G.T.Prance, Fl. Neotrop. Monogr. 9: 245 (1972); isolecto: 

F, NY. 


Map: G.T.Prance, Fl. Neotrop. Monogr. 9S: 207 (1989).


Tree to 10 m tall. Leaves with stipules to 7 mm long, linear, caducous; petioles 4–7 mm long 

sparsely pubescent, terete, with 2 sessile glands near base of lamina; lamina oblong, 10–22 × 

4–8.5 cm, rounded to cuneate at base, acuminate at apex with acumen 2–12 mm long, 

glabrous above (or with sparse appressed pubescence when very young), densely silver-greyarachnoid 

beneath; primary veins 15–20 pairs, prominent beneath. Inflorescences short 

densely-flowered axillary corymbose racemes; rachis with short reddish-brown pubescence. 

Bracts and bracteoles 1.5–3.5 mm long; deciduous. Receptacle cylindrical, slightly curved, 

9–14 mm long, with short reddish-brown pubescence on exterior, glabrous within except for 

deflexed hairs at throat, with thick walls leaving the cavity small; pedicels 2–3 mm long. 

Calyx lobes rounded. Petals 5, white, sparsely ciliate or glabrous on margins. Ovary densely 

villous. Style densely villous almost to apex. Stamens c. 100, inserted in a complete circle; 

filaments far exserted, forming a dense tangled mass. Fruit globose to oblong, to 9 cm long; 

epicarp verrucose or smooth, glabrous; mesocarp thick, fleshy; endocarp thick, fragile, 

granular-fibrous, glabrous within. 

Widespread in western Amazonia and cultivated elsewhere. 82: VEN. 83: CLM, ECU, PER. 

84: BZN (AC, AM, PA). Non-flooded forest and secondary forest. Map 55. 

82. VENEZUELA: Amazonas, San Carlos de Rio Negro, 20 km S of confluence of Rio Negro and Brazo 

Casiquiare, R.L.Liesner 6492 (MO, NY). 83. COLOMBIA: Vaupés, Mitu (cultivated), J.L.Zarucchi et al. 

1112 (COL); Amazonas, Mun. de La Pedrera, Inspección Santa Isabel, P. N. Natural Cahuinarí, A.Rudas et al. 

1040 (MO). ECUADOR: Napo, P. N. Yasuní, M.Aulestia 2760 (K, MO); Napo, Estación Biológica Jatun 

Sacha, Río Napo, C.E.Cerón 3754 (K, MO); Pastaza, Pastaza Cantón, Moretecocha Petroleum Well, 

S.Espinoza et al. 741 (K, MO); Zamora-Chinchipe, trail Shaime to Hito, J.Jaramillo 14444 (NY). PERU: 

Loreto, Pampa Hermosa, Río Corrientes, W.H.Lewis et al. 10282 (K, MO). 84. BRAZIL NORTH: Acre, 

Cruzeiro do Sul, O.P.Monteiro & C.Damião 454 (INPA, NY); Amazonas, Mun. de Borba, Rio Abacaxis, Vila 

Uaxiní, C.A. Cid Ferreira 4088 (INPA, NY). 

Cultivated in the Amazon basin for its edible fruit and as an ornamental. The Pará collections 

and some others are probably from cultivated trees since it appears to be native only in 

western Amazonia. 

47. Couepia eriantha Spruce ex Hook.f., in C.F.P.Martius, Fl. Bras. 14(2): 45 (1867) 

T: Brazil North: Pará, Santarém, by Rio Tapajós, R.Spruce 313; lecto: K, fide G.T.Prance, Fl. Neotrop. 

Monogr. 9: 246 (1972); isolecto: CGE, GH, OXF, (spec. s.n. BM, W, probably isotypes). 

Illustration: J.D.Hooker, in C.F.P.Martius, Fl. Bras. 14(2), t. 14(2) (1867). 


Shrub or small tree. Leaves with stipules to 3 mm long, linear, caducous; petioles 3.5–6 mm 

long, pubescent; lamina oblong to oblong-lanceolate, 10–14 × 3–5.5 cm, cordate to cuneate 

at base, apiculate or acuminate at apex with acumen to 5 mm long, glabrous on both 

surfaces; primary veins 15–20 pairs, prominent beneath. Inflorescences terminal and axillary 

panicles; rachis and branches densely brown-villous. Bracts and bracteoles ovate, to 15 mm 

long, persistent. Receptacle cylindrical, 15–18 mm long, densely ferrugineous-villous on 

exterior, glabrous within except for deflexed hairs at throat. Calyx lobes rounded, sericeous. 

Petals 5, white, pubescent, ciliate on margins. Stamens c. 40, inserted in a complete circle. 

Ovary densely villous. Style pubescent for 3/4 its length. Fruit (immature) long and narrow, 

with smooth glabrous epicarp. 

Central Amazonian Brazil. 84: BZN (AM, PA). Primary forest on periodically flooded 

ground. Flowering Nov.–Mar. Map 56. 

84. BRAZIL NORTH: Amazonas, Rio Solimões, Lago do Castanho, M.Honda & F.Mello INPA35428 (INPA, 

NY); Amazonas, Rio Purus, between Bom Intento and Boa Fé, G.T.Prance et al. 16368 (INPA, NY); 

Amazonas, Rio Negro, Ilha da Samaúma, L.Coelho INPA7259 (IAN, INPA, MG, NY); Pará, Lago Salgado, 

beira do Castanhal, A.Ducke MG8879 (BM, MG, RB); Pará, Santarém, Rio Maicú, Serra de Taperinha, 

M.B.Silva 1408 (MG, SP). 

48. Couepia trapezioana Cuatrec., Brittonia 8: 197 (1956) 

T: Colombia: Amazonas, Trapecio Amazónico, R. Loretoyacu, R.E.Schultes 8266; holo: F; iso: IAN, U. 


41

42 


Tree, medium-sized to large. Leaves with stipules to 2 mm long, linear, caducous; petioles 

5–10 mm long, sparsely pubescent, shallowly canaliculate; lamina ovate to oblong-elliptic, 

10–15 × 4–10 cm, rounded at base, acuminate at apex with acumen 3–5 mm long, glabrous 

above, dense grey or brown-appressed-arachnoid beneath; primary veins 14–18 pairs, 

prominent beneath. Inflorescences terminal panicles; rachis and branches with short light 

brown pubescence. Bracts and bracteoles oblong-ovate, acute, 6–8 mm long, caducous to 

subpersistent. Receptacle cylindrical, 7–8 mm long, densely light-brown-pubescent on exterior, 

glabrous within except for deflexed hairs at throat; pedicels 1 mm long. Calyx lobes narrow, 

rounded to acute. Petals 5; margins ciliate. Stamens c. 24, inserted in a nearly complete 

circle; filaments connate at base. Ovary villous. Style pubescent for 1/2 its length. Fruit 

(immature) ellipsoid; epicarp smooth, glabrous; mesocarp thin, fleshy; endocarp thin, fragile, 

granular, glabrous within. 

Western Amazonia. 83: CLM, PER. 84: BZN (AM). Forest. Map 57. 

83. COLOMBIA: Amazonas, Loretoyacu, R.E.Schultes & G.A.Black 46-153 (IAN, NY). PERU: Loreto, Río 

Nanay, Puerto Almendras, 20 km WSW of Iquitos, J.Chota 6 (K, NY); Loreto, Río Ampiacu, Puca Urquillo, 

T.B.Croat 20663 (MO, NY, P, WAG). 84. BRAZIL NORTH: Amazonas, Santa Isabel, Rio Negro, J.M.Pires 

315 (IAN, NY). 

49. Couepia stipularis Ducke, Arq. Inst. Biol. Veg. 4: 3 (1938) 

T: Brazil North: Amazonas, Manaus, A.Ducke RB 35583; holo: RB; iso: INPA, K, NY, P, U, US. 


Tree to 30 m tall. Leaves with stipules lanceolate, to 15 mm long, persistent and adnate to 

base of petiole, silvery-grey-pubescent; petioles 9–15 mm long, silvery-grey-pubescent, 

terete, with 2 glands near lamina base; lamina oblong to oblong-lanceolate, 10–18 × 3.7–7 

cm, rounded to subcordate at base, mucronate at apex with acumen 3–4 mm long, glabrous 

above, with a few short grey hairs beneath; primary veins 14–20 pairs, prominent beneath. 

Inflorescences terminal and axillary panicles 4.5–9 cm long, densely-flowered, with a thick 

erect striated rachis; rachis and branches with sparse appressed grey pubescence. Bracts and 

bracteoles 1–5 mm long, lanceolate, caducous. Receptacle c. 7 mm long, subcylindrical and 

curved anteriorly, grey-pubescent on exterior, glabrous within except for deflexed hairs at 

throat; pedicel 3 mm long. Calyx lobes rounded. Petals 5, white, caducous, glabrous or with 

a few isolated hairs on margins. Stamens 27–35, inserted in a nearly complete circle. Ovary 

pilose. Style glabrous except for base. Fruit ovoid, 3.5–5 × 2.5–3.5 cm; pericarp thin, fragile, 

composed of coarse fibres, glabrous within. 

Brazil (vicinity of Manaus); known only from 2 collections. 84: BZN (AM). Non-flooded 


84. BRAZIL NORTH: Amazonas, Manaus, A.Ducke 1755 (A, F, IAN, K, MG, NY, R, US). 

50. Couepia reflexa Ducke, Arch. Jard. Bot. Rio de Janeiro 5: 118 (1930) 

T: Brazil North: Pará, Jurití Velho, A.Ducke RB19757; holo: RB; iso: IAN, K, NY, P, U. 


Tree, medium-sized. Leaves with stipules c. 2 mm long, caducous; petioles 5–8 mm long, 

glabrous, terete; lamina membranous, oblong-elliptic, 10–17 × 4–8 cm, subcuneate at base, 

acuminate at apex with acumen 9–10 mm long, glabrous on both surfaces; primary veins 

10–13 pairs, prominent on both surfaces. Inflorescences axillary reflexed racemose panicles 

to 8 cm long; rachis and branches sparsely silver-grey pubescent. Bracts and bracteoles 1–3 mm 

long, lanceolate, persistent. Receptacle cylindrical, c. 7 mm long, sparsely silver-greypubescent 

on exterior, glabrous within except for deflexed hairs at throat; pedicels 3 mm 

long. Calyx lobes oblong, rounded. Petals 5, white, ciliate on margins. Stamens c. 35, 

inserted in a complete circle. Ovary pilose. Style glabrous except at base. Fruit unknown. 

Brazil; known only from the type collection. 84: BZN (PA). Primary forest on high ground. 

Map 59.


51. Couepia longipendula Pilg., Notizbl. Königl. Bot. Gart. Berlin 6: 141 (1914) 

T: Brazil North: Amazonas, Manaus, Bosossa, E.Ule 8854; holo: B, destroyed; lecto: K, fide G.T.Prance, 

Fl. Neotrop. Monogr. 9: 248 (1972); iso: L, LA, MG. 


Tree 5–30 m tall. Leaves with stipules lanceolate, to 3 mm long, persistent; petioles 4–8 mm 

long, shallowly weakly canaliculate, glabrous; lamina oblong-elliptic, 6–16 × 3–7.5 cm, 

rounded to cuneate at base, caudately acuminate at apex with acumen 1–1.5 cm long, 

glabrous on both surfaces; primary veins 5–8 pairs, almost plane. Inflorescences pendulous 

panicles on long (to 60 cm) peduncles, mostly glabrous but sparsely pubescent towards apex. 

Bracts and bracteoles 1–3 mm long, apiculate, caducous. Receptacle turbinate, 10–15 mm 

long, with short light brown pubescence on exterior, completely glabrous within. Calyx lobes 

rounded, with 2 sessile glands on exterior. Petals 5, rose, caducous, ciliate on margins. 

Stamens c. 32, inserted in a nearly complete circle, with a few toothed staminodes 

interspersed on distal side opposite ovary. Ovary densely villous. Style glabrous, flattened at 

base. Fruit round to ellipsoid, 4–6 × 4 cm; epicarp with short sparse brown pubescence; 

pericarp hard, thick, fibrous, not differentiated, densely lanate within. 

Brazil (lower Rio Negro region). 84: BZN (AM, PA). Non-flooded forest. Flowering Aug.–Nov. 

Map 60. 

84. BRAZIL NORTH: Amazonas, foothills of Serra Curicuriari, G.T.Prance et al. 16143 (INPA, NY, US); 

Amazonas, Distrito Agropecuário, Reserva 1501, N of Manaus, M.A.Freitas et al. 327 (INPA, K, NY); 

Amazonas, Santo Antônio de Abonari, Manaus-Caracaraí km 220, G.T.Prance et al. 24269 (INPA, NY, U, US); 

Pará, Rio Trombetas, Planalto Saracá, N.T. da Silva & M.R.Santos 4612 (MG, NY); Pará, Mun. de Oriximiná, 

Rio Trombetas near Cachoeira Porteira, C.A. Cid Ferreira & J.Ramos 1059 (INPA, MG, NY, US). 

The fruit contains oil and is cultivated around Manaus for its edible cotyledons. 

52. Couepia dolichopoda Prance, Brittonia 26: 302 (1974) 

T: Peru: Loreto, Varadura de Mazán, from Río Amazonas to Río Napo, T.B.Croat 19382; holo: NY; iso: MO, RB. 

Illustrations: G.T.Prance, Brittonia 26: 303 (1974); G.T.Prance, Fl. Colomb. Monogr. 19: 26 (2001). 


Tree to 30 m tall. Branches completely glabrous when young. Leaves with stipules unknown; 

petioles 10–12 mm long, glabrous, shallowly canaliculate; lamina elliptic, coriaceous, 9–11.5 × 

3.8–5.3 cm, cuneate at base, acuminate at apex with acumen 10–15 mm long, glabrous on 

both surfaces; midrib prominulous on both surfaces, glabrous; primary veins 6–8 pairs, 

almost plane on both surfaces. Inflorescences of pendulous panicles on long peduncles up to 

75 cm; rachis and branches glabrous; branches small, numerous. Bracts and bracteoles 

0.5–1.5 mm long, acute, glabrous except for the sparsely ciliate margins, subpersistent. 

Receptacle campanulate; tube 4–6 mm long when mature, contracted into pedicels 10–15 mm 

long, glabrous within and without. Calyx lobes 5, rounded, unequal, with 2 considerably 

longer than the other 3, sparsely puberulous when young; margins ciliate; exterior with 

2 sessile conspicuous glands. Petals 5, glabrous, caducous, red. Stamens 16–21, unilateral, 

with c. 8 tooth-like staminodes opposite. Ovary sparsely villous. Style glabrous. Fruit 

obpyriform, 4.5 cm long × 3.5 cm wide at broadest point which is well below mid-point; 

exocarp smooth, glabrous; mesocarp thick, hard, fibrous; endocarp thin, bony. Fig. 57. 

Western Amazonia. 83: CLM, PER. 84: BZN (AM). Forest on terra firme. Map 61. 

83. COLOMBIA: Amazonas, Correg. de Puerto Santander, La Chorrera, 7 km SW of Araracuara, D.Cárdenas 

& M.V.Arbelaez 4220 (COAH, K); Caquetá, Río Messay, A.Roa 365 (COL, INPA). PERU: Loreto, Maynas, 

Caserío Mishana, Río Nanay, J.Ruíz & H.Murphy 178 (K, MO); Loreto, Maynas Prov., Estac. Biol. Callicebus, 

Río Nanay-Mishana, R.Vasquez et al. 2788 (MO, NY); Loreto, Maynas Prov., near Brillo Nuevo, Río 

Yaguayaca, A.Gentry & J.Revilla 20385 (MO, NY). 84. BRAZIL NORTH: Amazonas, Mun. de Amaturá, 

São Domingos, Rio Solimões, D.C.Daly et al. 4387 (INPA, NY); Amazonas, Rio Javari, behind Estirão do 

Equador, E.Lleras P17294 (INPA, NY); Amazonas, Carauari, Rio Juruá, A.S.L. da Silva et al. 452 (MG, NY). 

The fruit (cotyledons) is edible, and is used as a source of cooking oil. 

43

44 


Figure 57. Couepia dolichopoda. A, habit; B, flower bud; C, ½ flower; D, petal; 

E, diagrammatic view of staminal arrangement (A–E, T.B.Croat 19382). Scale bars: A = 5 cm; 

B, C = 1 cm; D = 3 mm. Drawn by Charlie Clare. Previously published in G.T.Prance, 

Brittonia 26: 303 (1974).


53. Couepia cognata (Steud.) Fritsch, Ann. K. K. Naturhist. Hofmus. 4: 60 (1889) 

Hirtella cognata Steud., Flora 26: 761 (1843); Couepia steudeliana Miq., Stirp. Surinam. Select. 28 (1851); 

Moquilea steudeliana (Miq.) Walp., Ann. Bot. Syst. 2: 463 (1852), nom. superfl. illeg.; Parinari hostmannii 

Fritsch, Ann. K. K. Naturhist. Hofmus. 5: 13 (1890), as Parinarium hostmanni, nom. superfl. illeg. T: Suriname, 

W.R.Hostmann 795; holo: P; iso: CGE, GH, K, LE, NY, TCD, U, W. 


Shrub or small tree 1–10 m tall (rarely large trees). Leaves with stipules to 9 mm long, 

linear-filamentous, deciduous; petioles 2 mm long, densely pubescent; lamina oblong to 

oblong-lanceolate, 2.5–6 (–15) × 1–2 (–4.5) cm, cuneate to rounded at base, acuminate at 

apex with acumen 2–12 mm long, glabrous or sparsely appressed-pubescent above, with 

grey- to brown-lanate pubescence beneath which usually obscures the prominently reticulate 

parallel venation; midrib prominent beneath, impressed above, pubescent on both surfaces; 

primary veins 7–10 (–15) pairs, prominent beneath. Inflorescences terminal panicles; rachis 

and branches brown-villous. Bracts and bracteoles lanceolate, to 6 mm long, caducous. 

Receptacle cylindrical, curved anteriorly, 5–8 mm long, reddish-brown and often villous on 

exterior, glabrous within except for deflexed hairs at throat; pedicels 1–3 mm long. Calyx 

lobes acute. Petals 5, white or pale pink, glabrous. Stamens 12–25, unilateral, with a few 

teeth-like staminodes opposite. Ovary villous. Style pubescent. Fruit globose; epicarp 

pubescent when young, verrucose and glabrous when mature; pericarp hard and thick, 

undifferentiated, densely lanate within. 

1 Leaf lamina thin and membranous; flowers with short ferrugineous 

pubescence on exterior 53c. var. membranacea 

1: Leaf lamina thick and coriaceous; flowers shaggy-villous on exterior 

2 Leaf lamina 2.5–6.5 × 1–2 cm 53a. var. cognata 

2: Leaf lamina 7–15 × 3–4.5 cm 53b. var. major 

53a. Couepia cognata (Steud.) Fritsch var. cognata 

Licania exiguifolia Standl., Bull. Torrey Bot. Club 67: 285 (1940). T: Guyana, A.S.Pinkus 245; holo: F; iso: 

MO, NY, S. 


Shrub or small tree. Leaf lamina thick and coriaceous, 2.5–6.5 × 1–2 cm; flowers shaggyvillous 

on exterior; calyx lobes acute. 

Guianas and adjacent Venezuela and Brazil. 82: GUY, SUR,VEN. 84: BZN (AM). Savanna, 

savanna forest, scrubland, and secondary forest. Map 62. 

82. GUYANA: Upper Demerara-Berbice Region, E of Linden, junction of road to Mabura Hill, B.M.Boom 

7145 (NY, P); Demerara-Mahaica Distr., Linden Highway, M.J.Jansen-Jacobs & B.J.H. ter Welle 5616 (K, U). 

SURINAME: Jodensavanna, B.W.5229 (A, K, MO, NY). VENEZUELA: Bolívar, Mun. Raúl Leoni, 5 km from 

base of Cerro Camarón, 42.5 km SE of Entre Rios, G.Aymard & A.Fernández 7173 (K, MO, PORT). 

84. BRAZIL NORTH: Amazonas, Rio Içana, near Tunui, R.L.Fróes 28080 (IAN, UB). 

53b. Couepia cognata var. major Prance, Fl. Neotrop. Monogr. 9: 249 (1972) 

T: Guyana: Bartica to Potaro road, mile 107, D.B.Fanshawe 1000 (F.D. 3736); holo: NY; iso: BRG, K. 


Tree to 35 m tall. Leaf lamina coriaceous, 7–15 × 3–4.5 cm. Flowers villous on exterior. 

Calyx lobes acute. 

Guyana (Potaro River region). 82: GUY. Primary forest. Map 63. 

82. GUYANA: 3 km above Kaieteur Falls, Potaro R., R.S.Cowan & T.Soderstrom 1789 (NY, P, US); 3 km 

above Kaieteur Falls, Potaro R., R.S.Cowan & T.Soderstrom 1992 (NY, US). 

45

46 


53c. Couepia cognata var. membranacea Prance, Fl. Neotrop. Monogr. 9: 249 (1972) 

T: Guyana: Upper Mazaruni River, D.B.Fanshawe & B.Maguire 32566; holo: NY; iso: F, K, US. 


Shrub or small tree. Leaf lamina membranous, thin, 4.2–6.5 × 1.6–3.0 cm. Flowers shortly 

rufous-tomentellous. Calyx lobes rounded. 

Guyana (Upper Mazaruni region). 82: GUY. Forest and savanna woodlands. Map 64. 

82. GUYANA: Membaru-Kurupuny Trail, B.Maguire & D.B.Fanshawe 32436 (F, NY, US). 

54. Couepia multiflora Benth., J. Bot. (Hooker) 2: 215 (1840) 

Moquilea multiflora (Benth.) Walp., Repert. Bot. Syst. 2: 7 (1843). T: Guyana, R. & R.Schomburgk 112; 

holo: K; iso: BM, BR, CGE, F, GH, L, LE, NY, OXF, P, W. 


Tree to 6 m tall. Leaves with stipules linear, to 9 mm long, caducous, membranous; petioles 

4–7 mm long, pubescent, terete, eglandular; lamina oblong to oblong-elliptic, 9–15 × 4–9 cm, 

rounded to cordate at base, rounded to short-acuminate at apex with acumen 1–5 mm long, 

glabrous above, densely grey-arachnoid beneath; midrib prominent beneath, pubescent above; 

primary veins 12–14 pairs, prominent beneath, slightly impressed above. Inflorescences 

terminal panicles; rachis and branches densely brown-tomentose. Bracts and bracteoles 2–5 mm 

long, oblong to lanceolate, caducous. Receptacle c. 7 mm long, cylindrical, curved anteriorly 

in bud, with short thick brown tomentum on exterior, glabrous within except for deflexed 

hairs at throat; pedicels 1 mm long. Calyx lobes rounded. Petals 5, white; margins glabrous 

or with a few isolated hairs. Stamens c. 28, inserted in an arc of c. 240°, with a sterile portion 

of circle opposite. Ovary pilose. Style pubescent for 1/2 its length. Fruit ellipsoid, 6.5 × 3.5 cm; 

exocarp smooth, glabrous; mesocarp c. 3 mm thick when dry, fleshy; endocarp thin, fibrous. 

Guyana and Brazil (Roraima). 82: GUY. 84: BZN (RR). Savannas and savanna scrublands. 

Flowering Oct.–Apr. Map 65. 

82. GUYANA: North of Rupununi, D.H.Davis 84 (NY); U. Takutu-U. Essequibo Region, S Rupununi, 

Bakparwan Creek, SE of Aishalton, T.W.Henkel 3696 (K, US). 84. BRAZIL NORTH: Roraima, Boa Vista, 

I.L.Amaral et al. 1448 (INPA, NY, UB, US); Roraima, 20 km from Ponte dos Macuxis, N.A.Rosa & 

M.R.Cordeiro 1423 (MG, NY); Roraima, Road Boa Vista-Venezuela, Rio Caomé, J.A.Silva et al. 643 (K, NY). 

55. Couepia ulei Pilg., Verh. Bot. Vereins Prov. Brandenburg 47: 149 (1905) 

T: Brazil North: Amazonas, Rio Juruá, Juruá-mirim [Juruá-Miry], E.Ule 5547; holo: B, destroyed; lecto: K, 

fide G.T.Prance, Fl. Neotrop. Monogr. 9: 251 (1972); isolecto: INPA, L, MG, RB. 


Tree to 15 m tall, or shrub. Leaves with stipules to 3 mm long, linear, caducous; petioles 3–7 mm 

long, pubescent, shallowly canaliculate, eglandular; lamina elliptic, 5–10.5 × 2.5–4.5 cm, 

rounded at base, acuminate at apex with acumen 3–11 mm long, glabrous above, densely greyto 

rufous-pubescent beneath; primary veins 10–13 pairs, prominent beneath. Inflorescences 

terminal few-branched panicles; rachis and branches with short grey-brown pubescence. 

Bracts and bracteoles 1–2 mm long, ovate, caducous. Receptacle subcylindrical, 5–7 mm long, 

shortly grey pubescent on the longitudinally striate exterior, glabrous within except for deflexed 

hairs at throat; pedicels 1–2 mm long. Calyx lobes rounded to acute. Petals 5, white, 

pubescent on exterior, ciliate on margins. Stamens c. 35, inserted in a complete circle. Ovary 

villous. Style pubescent for 1/2 its length. Fruit elongate ellipsoid, 5 × 2 cm; epicarp glabrous, 

verrucose; mesocarp fleshy, thin; endocarp thin, fragile, granular, densely pubescent within. 

Western Amazonia. 82: VEN. 83: CLM, ECU, PER. 84: BZN (AC, AM). Riverine forest and 

non-flooded forest. Flowering continuously. Map 66. 

82. VENEZUELA: Apure, R. F. San Camilo, 4–5 km N of San Camilo, Río Nulita, J.A.Steyermark et al. 

101788 (NY). 83. COLOMBIA: Amazonas, 2 km N of Leticia, C.Sastre & A.Gómez-Pompa 514 (P). 

ECUADOR: Napo, Río Lagarto Cocha, L.Holm-Nielsen et al. 20051 (AAU, QCA). PERU: Loreto, Maynas, 

Río Ampiyacu, J.Revilla 816 (MO, NY); Quebrada Paparo, R.Vásquez & N.Jaramillo 11633 (K, MO).


84. BRAZIL NORTH: Acre, Cruzeiro do Sul, Igarapé Preto, near confluence with Rio Moa, G.T.Prance et al. 

2968 (INPA, NY, US); Amazonas, Reserva Florestal Ducke, C.A.Sothers 992 (INPA, K); Amazonas, Mun. de 

Limoeiro, Est. Ecológica Juami-Japurá, Rio Japurá below mouth of Rio Puruê, C.A. Cid Ferreira et al. 7248 

(INPA, K, NY, US). 

56. Couepia comosa Benth., J. Bot. (Hooker) 2: 215 (1840) 

Moquilea comosa (Benth.) Walp., Repert. Bot. Syst. 2: 6 (1843). T: Guyana: Falls of Essequibo R., R. & 

R.Schomburgk 28; holo: K; iso: BM, BR, CGE, F, L, LE, NY, P, W. 


Tree to 8 m tall or shrub. Leaves with stipules linear, 1–2 mm long, caducous; petioles 2–4 mm 

long, short-pubescent, terete; lamina ovate-elliptic, 4–8.5 × 2–3.8 cm, shortly acuminate at 

apex with acumen 1–5 mm long, rounded to cordate at base, glabrous when young above, 

densely reddish-brown-pubescent beneath; primary veins 9–12 pairs, prominent beneath; 

midrib pubescent on both surfaces. Inflorescences compact little-branched terminal panicles; 

rachis and branches with short brown pubescence. Bracts and bracteoles to 3 mm long, ovate, 

caducous. Receptacle subcylindrical, 8–10 mm long, with short brown pubescence on 

exterior, glabrous within except for deflexed hairs at throat; pedicels 3 mm long. Calyx lobes 

rounded. Petals 5, white, sparsely pubescent on exterior, ciliate on margins. Stamens c. 65, 

inserted in a complete circle; filaments far exserted. Ovary pilose. Style densely pubescent 

for 2/3 its length. Fruit (in single specimen examined) ellipsoid, 4 × 1.8 cm; epicarp 

verrucose; pericarp undifferentiated, hard, thick. 

Guyana and Venezuela (Bolívar). 82: GUY, VEN. Rocky places beside rivers and on river 

islands in Guyana, and in rainforest in Venezuela. Flowering Sept.–Apr. Map 67. 

82. GUYANA: Potaro-Siparuni Region, Iwokrama Rainforest Reserve, Kurupukari base camp, D.Clarke 688 

(BM, K, US); W Demerara, Mabura Hill, Essequibo R., Waraputa Falls, H. ter Steege & P. de Jager 311 (K, U); 

Cuyuni R., Akaio Falls, N.Y.Sandwith 687 (K, NY, P, RB, U, US). VENEZUELA: Bolívar, Mun. Raul Leoni, 

Río Caroni, S of Plomo, G.Picón & J.Rosales 1505 (K, MYF); Bolívar, Reserva Forestal Imataca, Río Cuyuní, 

Isla Anacocco, B.Stergios et al. 4991 (FLAS, MO, NY, PORT). 

57. Couepia venosa Prance, Fl. Neotrop. Monogr. 9: 251 (1972) 

T: Brazil Southeast: São Paulo, Alto da Serra, F.C.Hoehne 17197; holo: NY; iso: K, SP, SPF. 



Tree or shrub. Leaves with stipules minute, caducous; petioles 7–13.5 mm long, terete; 

lamina oblong-elliptic, 8.5–15 × 3.5–6 cm, rounded to subcuneate at base, acuminate at apex 

with acumen 7–10 mm long, glabrous and shining above, densely tomentose beneath; 

primary veins 11–14 pairs. Inflorescence of terminal panicles, shortly brown-tomentose. 

Bracts and bracteoles small, oblong, caducous. Receptacle subcampanulate, 6–7 mm long, 

densely brown-tomentose on exterior; pedicels 2–3 mm long. Calyx lobes rounded. Petals 

pubescent on exterior. Stamens 37–50, inserted around complete circle. Ovary densely 

tomentose; style pilose almost to apex. Fruit ellipsoid, exocarp smooth, glabrous. 

Southern-central Brazil. 84: BZL (MG, RJ, SP). Mountain slopes. Map 68. 

84. BRAZIL SOUTHEAST: Minas Gerais, Serra de Antônio Pereira, C.A.Schwacke 13682 (RB); Rio de 

Janeiro, Nova Friburgo, Res. Ecol. Municipal de Macaé de Cima, C.M.B.Correia et al. 212 (K, RB); Rio de 

Janeiro, Macaé de Cima, Rio das Flores, G.Martinelli 11935 (NY, RB); São Paulo, Ubatuba, Núcleo 

Picinguaba, M.A. de Assis 409 (HRCB, K, SP); São Paulo, Iguape, Est. Ecol. Juréia-Itatins, I.Cordeiro et al. 

1494 (K, SP). 

58. Couepia polyandra (Kunth) Rose, Contr. U. S. Natl. Herb. 5: 196 (1899) 

Hirtella polyandra Kunth, Nov. Gen. Sp. Pl. 6: 246 (1823); Moquilea kunthiana Mart. & Zucc., Abh. Math.- 

Phys. Cl. Königl. Bayer. Akad. Wiss. München 1: 390 (1832); Couepia kunthiana (Mart. & Zucc.) Benth., 

J. Bot. (Hooker) 2: 216 (1840); Grymania polyandra (Kunth) Presl, Epimel. Bot. 193 (1849). T: Mexico, 

A. von Humboldt & Bonpland s.n.; holo: P. 

47

48 


Hirtella dodecandra Sessé & Moç. ex DC., Prodr. 2: 529 (1825); Couepia dodecandra (DC.) Hemsl., 

Hooker's Icon. Pl. 27. 6: 2620, 2621 (1901). T: J.M.Moçiño & M.Sessé t.302 (accession number 6331.0103 

in the Hunt Institute for Botanical Documentation); neotype: A.L.P.P. de Candolle, Calques Fl. Mexique, t.302 

(1874), designated here. 

Couepia floccosa Fritsch, Ann. K. K. Naturhist. Hofmus. 5: 12 (1890). T: Costa Rica Guanacaste, E.R. von 

Friedrichsthal 256; holo: W. 


Tree to 15 m tall. Leaves with stipules 2–4 mm long, linear, membranous, caducous; petioles 

4–7 mm long, with 2 inconspicuous medial glands, terete; lamina oblong to oblong-elliptic, 

6–13 cm long on flowering branches, longer on sterile and fruiting branches, 2.5–5.5 cm 

wide, rounded to cuneate at base, acuminate at apex with acumen 2–10 mm long, glabrous 

above when mature, with a caducous pubescence when young, densely arachnoid beneath; 

midrib prominent above, pubescent when young; primary veins 8–15 pairs, prominent on 

both surfaces. Inflorescences terminal and axillary panicles; rachis and branches with short 

light-brown pubescence. Bracts and bracteoles 1–3.5 mm long, ovate, caducous. Receptacle 

subcylindrical, c. 4 mm long, with short appressed pubescence on exterior, glabrous within 

except for deflexed hairs at throat; calyx lobes rounded. Petals 5, white, glabrous but with 

ciliate margins. Stamens 11–21, inserted in an arc of 180–240°, with a few staminodes 

opposite. Ovary villous. Style pubescent for 1/2 its length. Fruit ellipsoid to obovoid, 5–7 cm 

long × 3–4.5 cm broad when mature; epicarp smooth, glabrous; mesocarp thick, fleshy; 

endocarp thin, fragile, of a fibrous texture, glabrous within. 

Central America from Sinaloa in Mexico to Panama. 79: MXC, MXG, MXN, MXS, MXT. 

80: BLZ, COS, ELS, GUA, HON, NIC, PAN. 81: cub. Riverine forest and low woodland. 

Map 69. 

79. MEXICO GULF: Veracruz, Distr. Papuntla, I.Kelly 339 (US). MEXICO NORTHWEST: Sinaloa, Villa 

Union, J.G.Ortega 6198 (US). MEXICO SOUTHWEST: Jalisco, road to Barra de Navidad to Tequezquitlán, 

R.McVaugh 11879 (MICH, NY). 80. COSTA RICA: Heredia, P. N. Braulio Carrillo, Magsasay, Sarapiqui, 

R.Aguilar 130 (CR, K, MO). EL SALVADOR: Morazan, Sensemora, M.L.Reyna 1433 (F, K, LAGU). 

GUATEMALA: Peten, La Libertad, M.Aguilar 36 (A). HONDURAS: Atlantida, between La Masica & La 

Ceiba, D.Hazlett 3270 (NY). NICARAGUA: Rivas, near Moyogalpa, M.Nee & W.Robleto Téllez 28191 (MO). 

PANAMA: Cocle, 7 km N of El Cope on road to Summit, J.P.Folsom et al. 5672 (MO, NY). 81. CUBA: 

Alacranes, cultivated, G.M.Fortún & Menacal 8565 (NY). 

59. Couepia nutans Prance, Brittonia 31: 248 (1979) 

T: Colombia: Valle, Alto Yunda, Río Anchicaya, S.Hilty A-1; holo: US; iso: NY, UC. 

Illustrations: G.T.Prance, Brittonia 31: 249 (1979); G.T.Prance, Fl. Colomb. Monogr. 19: 38 (2001). 


Tree to 25 m. Branches sparsely puberulous when young, soon becoming glabrous. Leaves 

with stipules caducous, not seen; petioles 5–8 mm long, weakly canaliculate above, 

puberulous, with 2 glands; lamina elliptic, coriaceous, 13–16 × 5–7.5 cm, rounded at the 

base, acuminate at the apex with acumen 4–5 mm long, glabrous above, densely arachnoidpubescent 

beneath; midrib prominulous above, prominent beneath; primary veins 17–19 

pairs, plane above, prominent beneath. Inflorescence of terminal and axillary panicles to 7 cm 

long; rachis and branches with short light brown tomentellous pubescence. Bracts ovate, 

membranous, c. 5 mm long, caducous; bracteoles oblong, 2–3 mm long, puberulous on exterior, 

membranous, subpersistent. Receptacle subcylindrical, c. 5 mm long, shortly tomentellous on 

the exterior, glabrous within except for deflexed hairs at the throat. Calyx lobes 5, rounded, 

tomentellous. Petals 5, white, glabrous except ciliate margins. Stamens c. 16, inserted in a 

complete circle. Ovary villous. Style tomentose on lower 1/3 only. Fruit not seen. 

Colombia; known only from the type collection. 83: CLM. Collected at an altitude of 1000 m. 

Map 70. 

60. Couepia hondurasensis Prance, Kew Bull. 54: 110 (1999) 

T: Honduras: Yoro, Río Guán Guán, Km 1 road to San José Texíguat, R.Aguilar 4084; holo: K; iso: MO. 

Illustration: G.T.Prance, loc.cit.: 111.

Map: G.T.Prance loc.cit.: 106. 


Shrub 4 m tall. Branches glabrescent when young, conspicuously lenticellate. Leaves with 

stipules axillary, lanceolate, tomentellous, caducous, c. 4 mm long; petioles 5–7 mm long, 

weakly canaliculate, appressed pubescent, glabrescent; lamina broadly elliptic, chartaceous, 

12–19 × 5.5–9.5 cm, finely acuminate at apex with acumen 10–20 mm long, curved, 

subcuneate at base, glabrous above, lanate-pubescent beneath; midrib prominulous above, 

prominent beneath; primary veins 12–15 pairs, plane above, prominulous beneath. 

Inflorescences of densely clustered panicles with only short lateral branches; rachis and 

branches brown-tomentellous; bracteoles ovate, 4–5 mm long, caducous. Flowers 10–15 mm 

long on short articulated 2–4 mm pedicels. Receptacle cylindrical, lanate-pubescent on 

exterior, glabrous within except for reflexed hairs around throat. Calyx lobes lanate on 

exterior, glabrous except for puberulous margins. Petals 5, caducous. Stamens 25–30 inserted 

around complete circle. Ovary inserted on mouth of receptacle, villous. Style hirsute to apex. 

Fruit not seen. 

Honduras; known only from the type collection. 80: HON. Habitat unknown. Map 71. 

61. Couepia platycalyx Cuatrec., Fieldiana, Bot. 27: 66 (1950) 

T: Colombia: Cundinamarca, J.Cuatrecasas & E.Pérez-Arbeláez 15336; holo: F; iso: COL, US. 


Tree. Leaves with stipules caducous; petioles 7–8 mm long, glabrous, terete; lamina obovate 

to obovate-orbicular, 8.5–12 × 7–9 cm, rounded at base and apex, glabrous above, with very 

sparse short appressed pubescence beneath; primary veins 10–12 pairs, prominent beneath. 

Inflorescences terminal little-branched panicles; rachis and branches with a few minute 

appressed hairs only. Bracts and bracteoles to 1.5 mm long, caducous. Receptacle broadly 

turbinate, flattened, almost solid, oblique, 3 mm long, appressed pubescent on exterior, with 

dense deflexed hairs at throat within filling the very small cavity; pedicels 1.5 mm long. 

Calyx lobes rounded. Petals glabrous, white. Stamens c. 54, inserted in a complete circle. 

Ovary villous. Style pubescent at base only. Fruit ovoid, 8 × 5.5 cm; exocarp glabrous, 

smooth or with a few large scabs; mesocarp thin, fleshy; endocarp thin, bony, glabrous 

within. Fig. 58. 

Costa Rica, Venezuela, Colombia and Ecuador. 80: COS. 82: VEN. 83: CLM, ECU. High 

altitude Andean and Central American forests. Map 72. 

80. COSTA RICA: Limón, Cantón de Talamanca, Bratsi, Alto Lari, Kivut, R.Aguilar 1133 (INB, K, MO); 

Heredia, Cantón de Sarapiqui, Rara Avis, 15 km SW of Horquetas, O.Vargas 175 (CR, K, MO). 

82. VENEZUELA: Lara, 8 km from Sanare, W.Meijer et al. VEN92 (NO, NY). 83. COLOMBIA: Antióquia, 

Mun. de El Retiro, 4 km WNW of Cabecera of Quebrada La Agudelo, R.Bernal & G.Galeano 142 (HUA); 

Mun. de Yarumal, 1-5 km on road to San Fermin de Briceño, J.L.Luteyn & O.Escobar 13275 (NY); Valle, Res. 

Nat. El Refugio Torremdinos, Mun. de Dagua, E.Calderón-Sáenz 143C (K), 144C (K). ECUADOR: 

Esmeraldas, Río Lita, Sector El Cristal, W.Palacios 4340 (K, MO). 

62. Couepia rufa Ducke, Arch. Mus. Rio de Janeiro 22: 66 (1919) 

T: Brazil Northeast: Pernambuco, Recife, M. do Amaral Mello (comm. A.Lutz) MG3575, lectotype collection 

fide G.T.Prance, Fl. Neotrp. Monogr. 9: 254 (1972); lecto: RB, designated here; isolecto: BM, F, MG, P, R, US. 

Pleragina rufa Arruda ex H.Kost., Trav. Brazil 499 (1816), nom. inval., fide G.T.Prance, Mem. N.Y. Bot. 

Gard. 15(1): 133 (1966). 

Couepia glazioviana Pilg., Notizbl. Bot. Gart. Berlin-Dahlem 8: 537 (1923). T: Brazil Southeast: Rio de 

Janeiro, cultivated, A.F.M.Glaziou 18217; holo: B, destroyed; lecto: NY, designated here; iso: BR, LE, PR. 


Tree, large. Leaves with stipules linear, to 17 mm long, caducous; petioles 5–10 mm long, 

thick; lamina elliptic, 10–27 × 6.5–11.5 cm, subcordate at base, rounded to shortly acuminate 

at apex, glabrous above, densely rufous-arachnoid beneath; margins undulate, revolute; primary 

veins 15–25 pairs, prominent beneath, impressed above. Inflorescences terminal many-flowered 

panicles to 15 cm long; rachis and branches striate, thick, shortly grey-brown-pubescent. 

49

50 


Figure 58. Couepia platycalyx. A, habit (G.Herrera 5375); B, fruit (O.Vargas 175); C, ovary 

and style; D, ½ flower; E, flower; F, petal (C–F, G.Herrera 5375). Scale bars: A, B = 3 cm; 

C, D, F = 2 mm; E = 3 mm. Drawn by J.M.Fothergill.


Bracts and bracteoles oblong to oblong-lanceolate, to 7 cm long, caducous. Receptacle 

turbinate, thick-walled, c. 12 mm long, shortly light-brown-pubescent on exterior, glabrous 

within except for deflexed hairs at throat; pedicels 1–3.5 mm long. Calyx lobes rounded, 5–7 mm 

long. Petals 5, white, with ciliate margins. Stamens c. 60, inserted in a complete circle. 

Ovary pilose. Style pubescent for 3/4 of its length. Fruit ovoid, to 11 × 6 cm; epicarp 

verrucose; mesocarp thin, fleshy; endocarp thick, fragile. 

NE and SE Brazil. 84: BZE (BA, PE), BZL (MG). Primary forests. Flowering Sept.–Dec. 

Map 73. 

84. BRAZIL NORTHEAST: Bahia, Mun. de Una, Res. Biol. Mico-Leão, Km 46 Ilhéus-Una, A.M. de Carvalho 

et al. 6477 (CEPEC, K, NY); Pernambuco, Recife, M.O.Cavalcanti s.n. (RB256783); Pernambuco, Chã de 

Peroba, Quinta Georgelândia, F.Richardson s.n. (IPA30570). BRAZIL SOUTHEAST: Minas Gerais, Parque 

Estadual do Rio Doce, L.V.Costa s.n. (BHCB28653, K), E.P.Heringer 18549 (NY). 

Cultivated for its edible fruit which is sold in markets in Recife. 

Barbosa Rodrigues (Hort. Flum.: 165, 1894) described a species Moquilea rufa. This was 

based mainly on material of Couepia bracteosa Benth., but he wrote in his notes that 

material from Northeast Brazil with the local name 'oity-coroya' fell into his concept of 

M. rufa. Since this was a description that included two species recognised here, Ducke was 

correct to describe the material from Northeast Brazil as a new species rather than make a 

new combination. Moquilea rufa Barb.Rodr. is thus placed under synonymy of C. bracteosa. 

63. Couepia robusta Huber, Bol. Mus. Paraense Hist. Nat. 6: 76 (1910) 

T: Brazil North: Pará, Timbotena, Bragança railroad, R.S.Rodrigues MG9651; holo: MG; iso: BM, NY, P, RB, 

U, US. 


Tree to 30 m tall. Leaves with stipules to 5 mm long, lanceolate, caducous; petioles 5–8 mm 

long, with short brown pubescence; lamina oblong to elliptic, 9–23 × 4–11 cm, rounded to 

cordate at base, rounded or shortly acuminate at apex with acumen 1–5 mm long, glabrous 

above, densely rufous-brown-arachnoid beneath; primary veins 15–21 pairs, prominent 

beneath. Inflorescences terminal little-branched panicles; rachis and branches with reddishbrown 

tomentum. Bracts and bracteoles 3–7 mm long, ovate, caducous. Receptacle 

subcampanulate, 7–9 mm long, densely reddish-brown-pubescent on exterior, glabrous 

within except for deflexed hairs at throat; pedicels 2 mm long. Calyx lobes rounded, 6–7 mm 

long, densely reddish-brown-pubescent on exterior, densely light-brown-pubescent within. 

Petals 5, white, ciliate on margins. Stamens c. 43, inserted in an arc of c. 240°, with a 

toothed ridge of staminodes opposite. Ovary densely villous, often with 2 carpels developing. 

Style pubescent for 1/2 its length. Fruit ovoid, 5–6 × 3.5–4 cm; epicarp smooth, glabrous; 

mesocarp thick, fleshy, fibrous; endocarp thin, fragile, granular, sparsely pubescent within. 

Brazilian Amazonia. 84: BZE (MA), BZN (AM, AP, PA). Non-flooded forest. Flowering 

Aug.–Dec. Map 74. 

84. BRAZIL NORTH: Amapá, Mun. de Mazagão, Km 80 Macapá-Monte Dourado, B.V.Rabelo et al. 3020 

(MG, NY, US); Amazonas, Manaus, Rio Tarumã, A.Ducke 1267 (A, F, IAN, MG, NY, R, RB, US); Pará, 

30 km W of Tomé-Açu-Paragominas road, 30 km S of Tomé-Açu, D.C.Daly et al. 831 (MG, NY, US). 

BRAZIL NORTHEAST: Maranhão, Turiaçu, km 153, BR-316, N.A.Rosa & H.Vilar 3155 (MG). 

64. Couepia impressa Prance, Fl. Neotrop. Monogr. 9: 255 (1972) 

T: Brazil Northeast: Pernambuco, Mata Dois Irmãos, D.A.Lima 54-1897; holo: IPA; iso: IPA. 



Tree. Branches glabrous when young. Leaves with stipules c. 2 mm long, subulate, caducous; 

petiole 6–12 mm long, glabrous, canaliculate, eglandular; lamina oblong to oblong-lanceolate or 

ovate, 9–22 × 4–8.5 cm, acuminate at apex with acumen 3–7 mm long, rounded at base, 

glabrous above, sparsely pubescent beneath; midrib glabrous above; primary veins 14–22 

pairs, impressed above, prominent beneath. Inflorescences of terminal panicles; rachis and 

51

52 


branches densely grey-pubescent. Bracts and bracteoles ovate, 2–3.5 mm long, caducous. 

Receptacle subcampanulate, 6–7 mm long, slightly curved, grey-puberulous on exterior, 

glabrous within except for ring of deflexed hairs around throat. Calyx lobes acute. Petals 5, 

caducous; margins shortly ciliate. Stamens 15–17 inserted around 2/3 of circle; filaments 

slightly connate at base. Ovary villous. Style pubescent to middle. Fruit globose to ellipsoid; 

4.5–6 × 3–3.5 cm; exocarp smooth, glabrous; mesocarp fleshy, 3–5 mm thick when dry; 

endocarp c. 1 mm thick, fibrous, glabrous within. 

NE Brazil. 84: BZE (AL, BA, PE). Primary forest. 

Leaves oblong-ovate, 19–22 × 4–8.5 cm; primary veins 19–22 pairs 64a. subsp. impressa 

Leaves ovate, 9–13 × 4–7.5 cm; primary veins 14–19 pairs 64b. subsp. cabraliae 

64a. Couepia impressa Prance subsp. impressa 

Map: Prance, Fl. Neotrop. Monogr. 9S: 213 (1989). 

Leaves oblong-ovate, 19–22 × 4–8.5 cm; primary veins 19–22 pairs. 

NE Brazil. 84: BZE (AL, PE). Map 75. 

84. BRAZIL NORTHEAST: Alagoas, União dos Palmares, Fazenda Santo Antônio, I.Pontual 66-280 (IPA, 

PEUFR); Pernambuco, Caruaru, Res. Municipal de Brejos dos Cavalos, M.Borges et al. 50 (NY, US). 

64b. Couepia impressa subsp. cabraliae Prance, Revista Brasil. Bot. 2: 31 (1979) 

T: Brazil Northeast: Bahia, Santa Cruz de Cabrália, R.P.Belém & R.S.Pinheiro 2837; holo: CEPEC; iso: MG, NY. 


Leaves ovate, 9–13 × 4–7.5 cm; primary veins 14–19 pairs. Fruit globose, 2.2 ×1.9 cm, glabrous. 

NE Brazil. 84: BZE (BA). Common in restinga. Map 76. 

84. BRAZIL NORTHEAST: Bahia, Mun. de Una, 9 km E of Una, A.M. de Carvalho & G.P.Lewis 8777 

(CEPEC, K, NY); Bahia, Mun. de Maraú, Km 11 Porto de Campinhos-Maraú, A.M. de Carvalho et al. 216 

(CEPEC, NY); Bahia, Mun. de Ilhéus, Mata da Esperança, A.M. de Carvalho et al. 4693 (CEPEC, NY). 

65. Couepia meridionalis Prance, Fl. Neotrop. Monogr. 9: 256 (1972) 

T: Brazil Southeast: São Paulo, Santos, H.Mosén 3377; holo: S; iso: K, LE, P. 



Tree. Leaves with stipules small, caducous; petioles 5–10 mm long, canaliculate; lamina elliptic 

to oblong-elliptic, 6.5–11.5 × 3–6 cm, rounded at base, acuminate at apex with acumen 3–5 mm 

long, glabrous above, densely lanate tomentose beneath; primary veins 10–13 pairs. 

Inflorescence of little-branched terminal panicles, sericeous-tomentose. Receptacle 

cylindrical, 8–9 mm long, densely sericeous-tomentose on exterior; pedicels 1.5–2 mm long; 

petals pubescent on exterior. Stamens c. 25 inserted around complete circle. Ovary densely 

pilose; style pilose almost to apex. Fruit not seen. 

Brazil (São Paulo); known only from the type collection. 84: BZL (SP). Map 77. 

66. Couepia coarctata Prance, Fl. Neotrop. Monogr. 9S: 77 (1989) 

T: Brazil Northeast: Bahia, Mun. de Maraú, 10 km S of Porto de Campinhos on road to Ubaitaba, 

A.M.Carvalho & G.P.Lewis 1116; holo: CEPEC; iso: NY. 



Tree to 6 m tall. Branches sparsely puberulous when young, soon becoming glabrous and 

lenticellate. Leaves with stipules 1–2 mm long, ovate, membranous, caducous; petioles 6–10 mm 

long, lanate, with 2 glands at base (visible in young petioles only), becoming rugulose with 

age; lamina oblong to oblong-elliptic, thickly coriaceous, 6–13 × 3.5–5.5 cm, subcordate to


Figure 59. Couepia coarctata. A, habit; B, fruit; C, leaf lower surface, D, flower bud; 

E, margin of sepal showing glands; F, flower section; G, petal (A–G, A.Carvalho 1116). 

Scale bars: A = 2 cm; D, F, G = 5 mm. Drawn by Bobbi Angell. Previously published in 

G.T.Prance, Fl. Neotrop. Monogr. 9S: 78 (1989). 

53

54 


rounded at base, rounded at apex, acute or bluntly acuminate, glabrous and lustrous above, 

densely velutinous-arachnoid ferrugineous beneath; midrib slightly impressed above, 

prominent and lanate beneath; primary veins 9–13 pairs, impressed above, prominent 

beneath. Inflorescences of terminal densely crowded panicles with many short branches 

only; rachis and branches puberulous. Bracts and bracteoles small, membranous, very early 

caducous. Receptacle cylindrical, 7–9 mm long, with short dense puberulous pubescence on 

exterior, glabrous within except for deflexed hairs at throat and dense lanate pubescence 

around stamen bases; pedicels 2–6 mm long, thin. Calyx lobes 5, acute, with sessile glands 

around margins. Petals 5, glabrous except for ciliate margins. Stamens c. 17, inserted around 

a complete circle. Ovary densely lanate. Style pubescent for 3/4 of its length. Fruit ellipsoidpyriform, 

tapering towards apex, 4.5 cm long × 3 cm broad; exocarp smooth, glabrous; 

pericarp hard, fibrous, 3 mm thick, glabrous within. Fig. 59. 

Brazil (Bahia, Mun. de Maraú). 84: BZE (BA). Restinga. Map 78. 

84. BRAZIL NORTHEAST: Bahia, Mun. de Maraú, road to Porto de Campinhos, A.M. de Carvalho & 

G.P.Lewis 1106 (CEPEC, NY, RB); Bahia, km 11 Porto de Campinhos-Maraú, A.M. de Carvalho et al. 194 

(CEPEC, NY). 

67. Couepia longipetiolata Prance, Revista Brasil. Bot. 2: 31 (1979) 

T: Brazil Northeast: Bahia, Itacaré, Estrada do aeroporto, T.S. dos Santos 2935; holo: CEPEC; iso: FHO, NY. 

Illustration: G.T.Prance, Revista Brasil. Bot. 2: 32 (1979). 


Tree 8 m tall. Branches glabrous when young, conspicuously lenticellate. Leaves with stipules 

small, ovate, membranous, caducous; petioles 14–18 mm long, canaliculate on upper surface, 

lanate pubescent, transversely rugose, 2–3 mm thick; lamina oblong, coriaceous, 12–15 × 

4–4.5 cm, cuneate at base, shortly and abruptly acuminate at apex with acumen 2–3 mm 

long, glabrous above except on midrib, lanate brown-pubescent beneath, appearing smooth 

but pubescence covering a deeply reticulate venation with stomatal crypts; midrib prominent 

beneath, slightly impressed and lanate on basal 1/2 above; primary veins 11–13 pairs, slightly 

prominent beneath, slightly impressed to plane above. Inflorescences of terminal and 

subterminal panicles, densely crowded, short branched; rachis and branches rufoustomentellous. 

Bracts caducous, to 5 mm long, membranous, glabrous; bracteoles caducous, 

c. 1 mm long, tomentellous on exterior. Receptacle campanulate-turbinate, 4–5 mm long, 

rufous tomentellous on exterior, glabrous within except for deflexed hairs at throat. Calyx 

lobes acute, tomentellous on both surfaces. Petals 5, cream, glabrous with ciliate margins. 

Stamens c. 30, inserted around a complete circle, far exserted. Ovary villous-pubescent. 

Style pilose-tomentose almost to apex, slightly exceeding the filaments. Fruit not seen. 

Brazil (Bahia); known only from two collections. 84: BZE (BA). Littoral forest. Map 79. 

84. BRAZIL NORTHEAST: Bahia, Mun. de Maraú, Fazenda Água Boa, 22 km E of Ubaitaba, S.A.Mori 12749 

(CEPEC, NY). 

68. Couepia pernambucensis Prance, Fl. Neotrop. Monogr. 9: 256 (1972) 

T: Brazil Northeast: Pernambuco, Recife, Mata Dois Irmãos, Córrego de Sabiá, D.A.Lima 53-1206; holo: IAN; 

iso: IPA, PEUFR. 



Tree, large. Branches puberulous when young, soon becoming glabrous and lenticellate. 

Leaves with stipules small, caducous; petioles 4–5 mm long, lanate-pubescent, eglandular, 

canaliculate above; lamina ovate-elliptic to oblong, 5–6.5 × 2–2.8 cm, acuminate at apex 

with acumen 2–6 mm long, rounded at base, glabrous above, sparsely arachnoid pubescent 

beneath; midrib pubescent towards base; primary veins 8–10 pairs, prominent beneath. 

Inflorescences of terminal and axillary panicles; rachis and branches densely tomentellous. 

Bracts and bracteoles to 2.5 mm long, caducous. Receptacle cylindrical, 5–6 mm long, 

slightly curved, densely tomentellous on exterior, glabrous within except for ring of hairs


around throat; pedicels c. 1 mm long. Calyx lobes acute. Petals 5, caducous, white; margins 

ciliate. Stamens 14–17, inserted around complete circle. Ovary pilose. Style pilose. Fruit 

ellipsoid, exocarp smooth, glabrous; endocarp thin, granular, fragile, glabrous within. 

Brazil (Pernambuco and Bahia). 84: BZE (BA, PE). Primary forest. Map 80. 

84. BRAZIL NORTHEAST: Bahia, Una, Fazenda São Rafael, T.S.Santos 322 (NY). 

69. Couepia froesii Prance, Fl. Neotrop. Monogr. 9: 258 (1972) 

T: Brazil North: Pará, Porto do Móz, Jaraucá, R.L.Fróes 32164; holo: IAN; iso: NY. 



Tree, small. Branches glabrous when young, lenticellate. Leaves with stipules small, caducous; 

petiole 6–8 mm long, glabrous, eglandular, canaliculate above; lamina oblong, 11–23 × 

4.5–8.5 cm, caudate-acuminate at apex with acumen 8–14 mm long, rounded at base, 

glabrous above, densely short-grey-lanate pubescent beneath; midrib glabrous; primary veins 

20–24 pairs, impressed above, prominent beneath. Inflorescences little-branched terminal 

and axillary panicles; rachis and branches densely grey-pubescent. Bracts and bracteoles 

small, caducous. Receptacle subcampanulate, 6 mm long, grey-pubescent on exterior, glabrous 

within except around throat below stamens; pedicels c. 1 mm long. Calyx lobes acute. Petals 

5, white, caducous; margins shortly ciliate. Stamens c. 30, inserted around complete circle. 

Ovary pilose. Style pubescent to middle. Fruit when young ovoid; exocarp smooth, 

pubescent soon becoming glabrous; endocarp granular, thin, fragile. 

Central Brazilian Amazonia. 84: BZN (AM, PA). Non-flooded forest. Map 81. 

84. BRAZIL NORTH: Amazonas, Rio Demini, Cuiufite, R.L.Fróes & G.Addison 28965 (IAN); Pará, Porto de 

Moz, R.L.Fróes 32195 (IAN, NY). 

Tree used extensively for construction by Yanomami indians. 

70. Couepia rankiniae Prance, Kew Bull. 47: 640 (1992), as rankinae 

T: Brazil North: Amazonas, N of Manaus, Distrito Agropecuário, Fazenda Porto Alegre, L.Mars et al. 

3304.1149; holo: INPA; iso: K. 


Tree to 20 m tall. Branches puberulous when young, soon becoming glabrous. Leaves with 

stipules axillary, lanceolate, c. 3 mm long, membranous, caducous; petioles 7–12 mm long, 

slightly canaliculate above, usually with 1 or 2 glands on lower surface and glands 

sometimes extending to midrib, sparsely puberulous, rugose; lamina oblong to oblongelliptic, 

chartaceous, 7.5–18.5 × 3.5–5.0 cm, rounded at base, abruptly acuminate at apex 

with acumen 3–10 mm long, glabrous above, with a dense appressed lanate pubescence 

beneath; midrib prominent and sparsely puberulous, glabrescent beneath, prominulous and 

glabrous above; primary veins 12–16 pairs, prominent beneath, plane or slightly impressed 

above. Inflorescence (only 1 partial inflorescence seen) paniculate, with flowers borne in 

small groups on a single rachis 6–8 cm long; rachis and branches tomentellous. Bracts and 

bracteoles caducous. Flower bud 12 mm long. Receptacle campanulate, tomentellous on 

exterior, glabrous within except for pilose mass around throat. Calyx lobes 5, acute, 

tomentellous on exterior. Petals 5. Stamens numerous, inserted around complete circle. 

Ovary inserted at mouth of receptacle. Style hirsute at base. Fruit when young globose, 

glabrous, densely lenticellate on exterior. 

Central Amazonia (N of Manaus) and recently collected in central French Guiana. 82: FRG. 

84: BZN (AM). Forest on terra firme. Map 82. 

82. FRENCH GUIANA: Base camp at Pic Matécho, c. 22.5 km NE of Les Eaux Claires, S.A.Mori & N.P.Smith 

25174 (K, NY). 84. BRAZIL NORTH: Amazonas, Manaus, Distrito Agropecuário, 90 km NNE of Manaus, 

Reserve 1501, A.A.Oliveira et al. 433 (INPA, K, NY, SPF); Amazonas, Fazenda Dimona, Reserve 2206, 

C.A.Mackenzie 2206.1521 (INPA, K); Amazonas, Fazenda Esteio, Reserve 1302, J.R.M.Nascimento 1302.2786 

(INPA, K). 

55

56 


71. Couepia parvifolia Prance, Fl. Neotrop. Monogr. 9: 258 (1972) 

T: Brazil Southeast: Rio de Janeiro, Serra de Petrópolis, 60 km E of Rio de Janeiro, A.P.Duarte 1508; holo: 

NY; iso: RB65362. 


Tree, small. Branches tomentellous when young, soon becoming glabrous. Leaves with 

stipules to 2 mm long, caducous; petioles 5–7 mm long, lanate-pubescent, glabrescent, 

eglandular, canaliculate above; lamina oblong to oblong lanceolate, 2.5–6.5 × 0.8–2.5 cm, 

acuminate at apex with acumen 5–7 mm long, cuneate at base, glabrous above, densely 

lanate-tomentose beneath; midrib glabrous; primary veins 7–9 pairs, prominent beneath. 

Inflorescences little-branched terminal panicles; rachis and branches brown-tomentose. 

Bracts and bracteoles 1–2.5 mm long, ovate, caducous. Receptacle subcampanulate, 5–6 mm 

long, brown-tomentose on exterior, glabrous within except for around throat beneath 

stamens; pedicels short, 0.25 mm long. Calyx lobes rounded. Petals 5, densely pubescent; 

margins ciliate. Stamens c. 28, inserted around complete circle. Ovary densely lanatepubescent. 

Fruit ovoid, 3.5 × 2.2 cm; exocarp smooth, glabrous; endocarp thin, granular on 

exterior, fragile, glabrous within. 

Brazil; known only from Rio de Janeiro. 84: BZL (RJ). Forests. Map 83. 

84. BRAZIL SOUTHEAST: Rio de Janeiro, Rio de Janeiro, Mata do Pai Ricardo, near Horto Florestal, 

Pessoal do Horto Florestal 1012 (NY, RB139762); Rio de Janeiro, Rio de Janeiro, Alto da Boa Vista, Estrada 

Vista Chinesa, near curve of km 1, C.A.L. de Oliveira 1713 (GUA, K, NY). 

Species Incertae Sedis 

Chrysobalanus subundulatus Brign., Mem. Mat. Fis. Soc. Ital. Sci. Modena, Pt. Mem. Fis., 

Ser. II 1: 71 (1862) 

Probably a Couepia sp. 

13. MARANTHES 

Maranthes Blume, Bijdr. Fl. Ned. Ind. 89 (1825); Greek for “withering flowers”. 

Exitelia Blume, Fl. Javae 1: vii (1828) nom. superfl. illeg.; Parinari subg. Exitelia Blume, Mélanges Bot. 

2: 10 (1855); Parinari sect. Exitelia (Blume) Müll.Hal., Ann. Bot. Syst. 4: 645 (1857). T: Maranthes 

corymbosa Blume. 

Grymania C.Presl, Epim. Bot. 193 (1851)["1849"]. T: Grymania salicifolia Presl. 

Parinari sect. Sarcostegia Benth., in W.J.Hooker, Niger Fl. 335 (1849); Parinari subg. Sarcostegia (Benth.) 

Miq., Fl. Ned. Ind. 1(1): 355 (1855). T: not designated. 

Trees. Leaves with a pair of glands at the junction of the lamina and eglandular petiole, 

rarely with marginal glands as well; stipules intrapetiolar, deltate, free or variously united, 

mostly caducous; lamina lower surface glabrous or with an arachnoid indumentum. 

Inflorescence usually a many-flowered corymbose panicle, rarely a lax few-flowered thyrse 

or a raceme of few-flowered monochasial cymes. Bracts and bracteoles eglandular, nearly 

always caducous, but bracteoles persisting up to anthesis in 2 species, not enclosing flowerbuds 

in small groups. Flowers bisexual, slightly zygomorphic; receptacle tube obconic, often 

slightly to strongly curved, always gradually narrowed into the pedicel, nearly always solid 

and almost completely filled with nectariferous tissue, only rarely hollow and then elongate 

and curved, glabrous inside on one side and hairy on the other or completely glabrous; faucal 

annulus well developed. Sepals 5, suborbicular, deeply concave. Petals 5, suborbicular to 

broadly lingulate. Stamens 20–60; filaments white, tightly undulate in bud with 2 or more 

undulations, inserted in 2 rows on the margin of the faucal annulus, forming a more or less 

complete circle, much longer than the sepals, usually occurring in a tangled mass; 

staminodes few, vestigial or absent. Ovary 2–3-carpellary, 2-locular, inserted laterally at

CHRYSOBALANACEAE (Maranthes) 

mouth of receptacle tube; style curved upwards, much longer than sepals, glabrous except at 

base. Drupe fleshy; endocarp very hard, fibrous, with a rough exterior; densely tomentose 

inside, with 2 lateral plates which break away on germination and allow the seedlings to 

escape. 2n = 20 (M. glabra ('Parinari glabra'), S.Mangenot & G.Mangenot, 1957 & 1962). 

12 species, of which 10 occur in tropical Africa and one each in tropical America and Asia. 

Of the African species, 6 occur in the forests of the Guineo-Congolian region, one in riparian 

forest slightly further north, one in rainforest on the East African mountains, one in the 

woodlands of the Sudanian region and another in the woodlands of the Zambezian region. 

Germination is phanerocotylar; cotyledons fleshy, pale green, soon shrivelling; cataphylls 

absent; first 2 eophylls opposite, the others opposite or alternate, with narrowly lanceolate, 

persistent stipules (Mensbruge, 1966: 124–131). 

A.J.G.H.Kostermans, A monograph of the genus Maranthes Bl. and Cyclandrophora Hassk. 

(Chrysobalanaceae) of the Asiatic and Pacific area, Candollea 20: 106 (1965); G.T.Prance, 

Notes on Chrysobalanaceae of Angola, Bol. Soc. Brot. sér. 2 40: 183 (1966); G.T.Prance, 

Maranthes, a new generic record for America, Brittonia 20: 203 (1968); G.T.Prance, 

Fl. Neotrop. Monogr. 9: 201, t. 33 (1972); F.White, The taxonomy, ecology and chorology of 

African Chrysobalanaceae (excluding Acioa), Bull. Jard. Bot. Belg. 46: 294 (1976); F.White, 

Distrib. Pl. African. 10: 313–325 (1976); F.White, Fl. Zamb. 4: 41, t. 11 (1978); R.Letouzey 

& F.White, Fl. Cameroun 20 (Fl. Gabon 24): 97, t. 29–33 (1978); G.T.Prance & F.White, 

The genera of Chrysobalanaceae, Philos. Trans. B320: 116-120 (1988). 

1 Receptacle tube elongate-turbinate, much longer than wide, usually 

longer than the sepals, gradually narrowed into the pedicel 

2 Flowers 3.5–5.5 cm long 

3 Leaves caudate-acuminate, cuneate or rounded at base; flowers 

5–5.5 cm long, glabrous outside; filaments c. 4 cm long; fruit angular 

in transverse section 5. M. gabunensis 

3: Leaves shortly cuspidate, subcordate at base; flowers c. 3.5 cm long, 

grey-velutinous-tomentellous outside; filaments c. 6 cm long; fruit terete 12. M. sanagensis 

2: Flowers up to 2 cm long 

4 Pedicel distinctly shorter than the receptacle tube 

5 Lower leaf surface persistently tomentose; indumentum fulvous, at 

least at first 2. M. chrysophylla 

5: Lower leaf surface glabrous 

6 Receptacle tube glabrous outside 8. M. kerstingii 

6: Receptacle tube tomentellous outside 7. M goetzeniana 

4: Pedicel at least as long as the receptacle tube 

7 Receptacle tube + sepals c. 1 cm long 

8 Leaf-apex rounded or subacute 4. M. floribunda 

8: Leaf-apex acuminate 8. M. kerstingii 

7: Receptacle tube + sepals c. 0.7 cm long 

9 Venation prominent on both surfaces; inflorescence few-flowered 

(c. 20), a raceme of cymes; drupe narrowly obovoid, c. 5 × 2 cm 9. M. panamensis 

9: Venation obscure; inflorescence many-flowered (>50), a much 

branched panicle of cymes; drupe ellipsoid, c. 4 × 3 cm 6. M. glabra 

1: Receptacle tube shortly turbinate, no longer than wide, ±equalling or 

shorter than the sepals, sharply differentiated from the pedicel 

10 Leaves cordate at base, denticulate towards apex; tertiary nerves subscalariform 

and conspicuous; bracteoles persistent up to or after anthesis 1. M. aubrevillei 

10: Leaves not cordate, entire; tertiary nerves not subscalariform, 

inconspicuous; bracteoles caducous 

57

58 


11 Leaves obtuse or apiculate at apex; receptacle tube + calyx c. 0.8 cm 

long 10. M. polyandra 

11: Leaves acuminate or subacuminate; receptacle tube + calyx up to 

0.7 cm long 

12 Leaves subacuminate, with lower surface arachnoid-tomentellous; 

receptacle tube cinnamoneous-tomentellous 11. M. robusta 

12: Leaves acuminate, glabrous; receptacle tube grey-tomentellous 3. M. corymbosa 

1. Maranthes aubrevillei (Pellegr.) Prance, Bull. Jard. Bot. Belg. 46: 295 (1976) 

Parinari aubrevillei Pellegr., Bull. Soc. Bot. France 78: 440 (1931), as Parinarium. T: Ivory Coast: 

Abidjan, A.Aubréville 185; holo: P. 

Illustrations: R.Letouzey & F.White, Fl. Cameroun 20 (Fl. Gabon 24): 101 (1978). 

Map: F.White, Distrib. Pl. African. 10: 314 (1976). 

Tree up to 30 m. Leaf lamina up to 12 × 5.5 cm; margin denticulate towards apex; apex 

shortly cuspidate; base cordate; tertiary nerves subscalariform; lower surface persistently 

grey-arachnoid-tomentellous. Bracteoles c. 0.4 × 0.25 cm, persistent up to or after anthesis. 

Upper pedicel c. 0.25 cm long. Receptacle tube c. 0.5 × 0.5 cm. Sepals c. 0.5–0.6 cm long. 

Flower exterior cinnamoneous-tomentose. Fruit ellipsoid, c. 4.5–5 × 2.5 cm; exocarp 

lenticellate. 

Sierra Leone to Cameroon. 22: BEN, BKN, GHA, IVO, LBR, NGA, SIE, TOG. 23: CMN. 

Evergreen rainforest. Map 84. 

22. GHANA: Ankasa F. R., A.A.Enti & Hall GC39137 (K). IVORY COAST: near Azaguié, A.Chevalier 22355 

(K, P); Dakpadou, A.Aubréville 847 (BR, K, P). SIERRA LEONE: Kambui, S of Kenema, H.C.King 283 (K). 

23. CAMEROON: margin of Cañon du Ntem, Oveng, 60 km E of Campo, R.Letouzey 10310 (P). 

2. Maranthes chrysophylla (Oliv.) Prance, Bull. Jard. Bot. Belg. 46: 295 (1976) 

Parinari chrysophylla Oliv., Fl. Trop. Afr. 2: 369 (1871), as Parinarium chrysophyllum; Ferolia chrysophylla 

(Oliv.) Kuntze, Revis. Gen. Pl. 1: 216 (1891). T: Gabon (Upper Guinea, Gabon River): G.Mann 978; holo: K; 

iso: P. 

Illustrations: A.Chevalier, Veg. Ut. Afr. Trop. Franç. 9, t. 17 (1917); R.Letouzey & F.White, Fl. Cameroun 20 

(Fl. Gabon 24): 103 (1978). 

Map: F.White, Distrib. Pl. African. 10: 315–317 (1976). 

Tree up to 30 m. Leaf lamina up to 25 × 13.5 cm; apex shortly acuminate or shortly 

cuspidate; base cuneate to subcordate; lower surface tomentellous, at first fulvous, ultimately 

fulvous or cinereous. Upper pedicel 0.2–0.3 cm long. Receptacle tube 0.4–0.6 cm long, 

fulvous-tomentellous outside. Sepals 0.4–0.6 cm long, fulvous-tomentellous outside. 

Maranthes chrysophylla subsp. chrysophylla occurs in evergreen rainforest in an interrupted 

band from Liberia to Gabon. The parapatric M. chrysophylla subsp. coriacea F.White occurs 

further inland, from Cameroon to SW Zaire. 

Leaves chartaceous, lanceolate-elliptic, gradually narrowed to a shortly 

acuminate apex; indumentum of lower leaf surface persistently fulvous; 

receptacle + pedicel c. 1 mm long 2a. subsp. chrysophylla 

Leaves coriaceous, oblong-elliptic or ovate-elliptic, suddenly contracted to 

the very short cuspidate apex; indumentum of lower leaf surface fulvous at 

first, soon becoming silvery; receptacle + pedicel c. 0.8 cm long 2b. subsp. coriacea 

2a. Maranthes chrysophylla (Oliv.) Prance subsp. chrysophylla 

Leaves chartaceous, lanceolate-elliptic, gradually narrowed to a shortly acuminate apex; 

indumentum of lower leaf surface persistently fulvous; receptacle plus pedicel c. 1 mm long. 

Liberia to Gabon. 22: GHA, IVO, LBR, NGA. 23: CMN, GAB. Coastal evergreen rainforest. 

Map 85.


22. GHANA: W Region, Tankwa Distr., Tano Nimiri F. R., A.A.Enti FH7543 (K). IVORY COAST: Tabou, 

A.Aubréville 1678 (BR, K, P). LIBERIA: Sinoe Co., Sangwin, J.T.Baldwin Jr. 11331 (K, NY, WAG). 

NIGERIA: Oban, P.A.Talbot s.n. (K). 23. CAMEROON: Londji, 45 km NNE of Kribi, J.J.Bos 4735 (BR, K, 

WAG); between Babong and Ogurany, 45 km SW of Mamfe, R.Letouzey 13587 (BR, K, P, WAG). GABON: 

Libreville, R.P.Klaine 1938 (K, P); Atsié near Lambaréné, F.Fleury in A.Chevalier 26564 (BR, K, P). 

2b. Maranthes chrysophylla subsp. coriacea F.White, Bull. Jard. Bot. Belg. 46: 296 (1976) 

T: Gabon: Oyem, G.Le Testu 9397; holo: P; iso: BR, WAG. 

?Parinari iodocalyx Mildbr., Notizbl. Bot. Gart. Berlin-Dahlem 8: 57 (1921), as Parinarium. T: Cameroon: 

near Dengdeng, 250 km NE of Yaoundé, G.W.J.Mildbraed 8670; holo: B, destroyed. 

Leaves coriaceous, oblong-elliptic or ovate-elliptic, suddenly contracted to the very short 

cuspidate apex; indumentum of lower leaf surface fulvous at first, soon becoming silvery. 

Receptacle plus pedicel c. 0.8 cm long. 

Cameroon, Gabon and SW Zaire. 23. CMN, GAB, ZAI. Inland rainforest and semi-deciduous 


23. CAMEROON: Meyos Mela, 32 km ENE of Djoum, R.Letouzey 8315 (K, P); 5 km SSW of Mayos, 

R.Letouzey 3963 (P); Yokadouma-Batouri, km 32, R.Letouzey 5232 (BR, P). GABON: Lastoursville, G. Le Testu 

7409 (BR, P); Mbighou, G.Le Testu 5969 (BR, P). ZAIRE: Gimbi, J.Wagemans 2209 (BR, FHO). 

3. Maranthes corymbosa Blume, Bijdr. Fl. Ned. Ind.: 89 (1825) 

Exitelia corymbosa (Blume) Span., Linnaea 15: 174 (1841); Parinari maranthes Blume, Ann. Mus. Bot. 2: 99 

(1856), nom. superfl. illeg., as Parinarium; Parinari corymbosa (Blume) Miq., Fl. Ned. Ind. 1(1): 356 (1855) 

et: 1084 (1858), as Parinarium corymbosum; Petrocarya maranthes (Blume) Miers, J. Linn. Soc., Bot. 17: 336 

(1879), nom. superfl. illeg.; Ferolia corymbosa (Blume) Kuntze, Revis. Gen. Pl. 1: 216 (1891). T: Jawa: 

Prov. Krawang, near Tjiradja, Kitnah, C.L.Blume 1516; holo: L; iso: BO. 

Maranthes multiflora Korth., Verh. Nat. Gesch. Ned. Bezitt., Bot. 259, tab. 70 (1844); Exitelia multiflora 

(Korth.) Walp., Repert. Bot. Syst. 5: 115 (1845–46); Parinari multiflora (Korth.) Miq., Fl. Ned. Ind. 1(1): 356 

(1855) et 1084 (1858), as Parinarium multiflorum. T: Borneo, P.W.Korthals s.n.; holo: L. 

Parinari griffithiana Benth., in W.J.Hooker, Niger Fl. 334 (1849), as Parinarium griffithianum; Ferolia 

griffithiana (Benth.) Kuntze, Rev. Gen. Pl. 1: 216 (1891). T: Malaya: Malacca, W.Griffith s.n.[20472]; 

holo: K; iso: L. 

Grymania salicifolia C.Presl, Epim. Bot. 193 (1851)["1849"]; Parinari salicifolia (C.Presl) Miq., Fl. Ned. Ind. 

1(1): 357 (1855), as Parinarium salicifolium; Ferolia salicifolia (C.Presl) Kuntze, Revis. Gen. Pl. 1: 216 (1891). 

T: Philippines: Luzon, Prov. of Alvay, Cuming 1057I; syn: C, K, L, MO, P, PR?, PRC? 

Maranthes speciosa Korth. ex Miq., Fl. Ned. Ind. 1(1): 357 (1855). T: Borneo: Martapura, P.W.Korthals 

s.n.; holo: L. 

Parinari racemosa S.Vidal, Cat. Met. Pl. Leños. 29 (1880), as Parinarium racemosum. T: not designated. 

Polyalthia pulchrinervia Boerl., Cat. Pl. Phan. 20 (1899). T: Jawa: Cult. Hort. Bogor IV G 42; holo: BO; iso: L. 

Parinari petiolata Malm, Notizbl. Bot. Gart. Berlin 11: 630 (1932), as Parinarium petiolatum. T: Lesser 

Sunda Is.: Flores, Mbaedjae, I.Rensch 4; holo: B? 

Parinari palauensis Kaneh., J. Dept. Agric. Kyushu Imp. Univ. 4: 325 (1934), as Parinarium palauense. 

T: Caroline Is.: Palau, R.Kanehira 469; holo: FU; iso: BISH, GH, US. 

Chrysobalanus ciliatus Korth. ex Miq., Fl. Ned. Ind. 1(1): 357 (1855), nom. inval. in synon. 

Ferolia minutiflorum Kuntze, Revis. Gen. Pl. 1: 216 (1891), nom. nud. 

Illustrations: A.J.G.H.Kostermans, Candollea 20: 143, 144, 145 (1965); G.T.Prance, Fl. Males. Ser. 1, 10(4): 

672 (1989). 

Map: G.T. Prance, Fl. Males. Ser. 1, 10(4): 674 (1989). 

59

60 


Figure 60. Maranthes. A–D, M. floribunda. A, habit; B, ½ flower; C, developing fruits; 

D, endocarp (A, Astle 753; B, Whellan 1553; C–D, F.White 3484). E, M. glabra. Stipules 

(J.D.Chapman 5244). Scale bars: A, C = 5 cm; B = 5 mm; D = 2 cm. Drawn by Janet Dyer 

& Rosemary Wise. Previously published in G.T.Prance & F.White, Philos. Trans. B320: 122 

(1988).


Tree up to 40 m tall, but sometimes flowering when only a few metres high. Leaf lamina up to 

14 × 6.5 cm; apex acuminate; base cuneate; lower surface glabrous. Upper pedicel 0.1–0.3 cm 

long. Receptacle tube 0.25 × 0.25 cm to 0.4 × 0.4 cm. Sepals 0.25–0.4 cm long. Flower 

exterior grey-tomentellous with denser and paler indumentum on 'protected' part of sepals. 

Extending from S Thailand through the Malay Peninsula and the Indonesian Archipelago to 

the Caroline Islands, tropical Australia and the Solomon Islands. 41: THA. 42: BOR, JAW, 

LSI, MLY, MOL, PHI, SUL, SUM. 43: BIS, NWG, SOL. 50: NTA, QLD. 62: CRL. 

Throughout its range it is rarely found very far from the coast. In Malaysia it is common in 

coastal areas, especially on rocky and sandy hills by the sea; exceptionally it can occur 

inland up to 600 m. In Australia, besides occurring as a 30 m tall tree in fringing forest, it is 

also found as a shrub on sand dunes and immediately behind mangrove swamps. Map 87. 

42. JAWA: Sukabuni Distr., Pelabuanratu, S.H.Koorders 34244 (BO, L). MALAYA: Johore, Sg. Rhu Reba, 

Jason Bay, E.J.H.Corner SFN28495 (BO, GH, K, SING). MALUKU: Kairatu Gemba, Kuswata & Soepadmo 

117 (A, BH, BISH, BM, BO, BRI, CAL, CANB, G, K, KEP, L, LAE, NX, P, PNH, SING, SXD, US). 

PHILIPPINES: Mt. Darna, G.E.Edaño PNH10847 [as 18047 in BR] (BO, BR, L, PNH). SULAWESI: 

Minhassa, Kajuwata, S.H.Koorders 18548 (BO, L). SUMATERA: S Bangka, Lobok Besar, 

A.J.G.H.Kostermans & Anta 1200 (A, BISH, BM, BO, BRI, CANB, K, KEP, L, LAE, NY, P, PNH, SYD, 

SING). 43. NEW GUINEA: Papua New Guinea, Lower Fly R., opposite Sturtevant Is., L.J.Brass 8014 (BO, 

GH, K, L). SOLOMON IS.: Vanikoro, near Laminu R., Walker BSIP219 (GH, K). 50. NORTHERN 

TERRITORY: Katherine Gorge, G.T.Prance 26603 (K, L, NY). 62. CAROLINE IS.: Palau, Babeldaob, 

F.R.Fosberg 32552 (BISH, US). 

Despite its very wide distribution, M. corymbosa is not very variable, other than in habit. For 

a full bibliography see A.J.G.H.Kostermans (1965). 

The wood is used for house-building and for posts. The fruit is edible. 

4. Maranthes floribunda (Baker) F.White, Bull. Jard. Bot. Belg. 46: 297 (1976) 

Parinari floribunda Baker, Bull. Misc. Inform. Kew 1897: 265 (1897), as Parinarium floribundum; Parinari 

polyandra subsp. floribunda (Bak.) R.A.Graham, Kew Bull. 1957: 230 (1957); Maranthes polyandra subsp. 

floribunda (Bak.) Prance, Bol. Soc. Brot., ser. 2, 40: 184 (1966). T: Malawi: Fort Hill, Whyte s.n.; holo: K; 

iso: BR (fragment). 

Parinari bequaertii De Wild., Feddes Repert. 13: 108 (1914), as Parinarium bequaerti. T: Zaire: 

Lumbumbashi, J.C.C.Bequaert s.n.; holo: BR. 

Parinari bequaertii var. longistaminea Hauman, Bull. Jard. Bot. État 21: 188 (1951). T: Zambia: Mufulira, 

F.Eyles 8201; holo: K; iso: BM, BR, SRGH. 

Illustrations: F.White, Fl. Zamb. 4: 42 (1978); G.T.Prance & F.White, Philos. Trans. B320: 122 (1988). 


Tree up to 10 m tall. Leaf lamina up to 15 × 8 cm; apex usually rounded, sometimes subacute; 

base rounded or subcordate, rarely cuneate; lower surface densely arachnoid-tomentose 

at first, glabrescent but with some hairs always persisting. Upper pedicel 0.7–1.2 cm long. 

Receptacle tube 0.7–1 × 0.4 cm. Sepals 0.7–0.8 cm long. Flower exterior usually greyvelutinous-tomentellous, 

less often coarsely fulvous-tomentose with spreading hairs or completely 

glabrous. Fruit ellipsoid to ovoid, c. 2.3 × 1.5 cm; exocarp glabrescent. Fig. 60A–D. 

Angola to Malawi and Tanzania. 23: ZAI. 25: TAN. 26: ANG, MLW, ZAM. Wetter types of 

Brachystegia, Julbernardia, Isoberlinia woodland (miombo). Map 88. 

23. ZAIRE: Kahemba, P.Duvigneaud 976 (BR); N of Lukafu, G.F. de Witte 6605 (BR, K). 25. TANZANIA: 

Tabora Distr., Ugalla R., P.Greenway & A.C.Hoyle 1034 (FHO); Mpanda Distr., Kabungu FR, S.R.Semsei 

2583 (BR, FHO, K). 26. ZAMBIA: Mbala Distr., L. Chila, H.M.Richards 13122 (EA, K); Solwezi-Nchangu, 

km 30, J.P.M.Brenan & P.Greenway 7998 (BM, BR, FHO, K); near Lundazi, R.G.Robertson 146 (BM, FHO, 

K). ANGOLA: Moxico Texeira de Souza, J.Gossweiler 12229 (BM, LISC); Lunda, Luma-Cassai, A.W.Exell & 

F.A.Mendonça 317 (BM, COI). 

5. Maranthes gabunensis (Engl.) Prance, Bol. Soc. Brot. ser. 2, 40: 184 (1966) 

Parinari gabunensis Engl., Bot. Jahrb. Syst.17: 87 (1893), as Parinarium gabunense. T: Gabon, H.Soyaux 

45; holo: B, destroyed; lecto: P, fide G.T.Prance & F.White, Philos. Trans. B320: 128 (1988); isolecto: K, Z. 

61

62 


Parinari gabunensis var. mayumbensis De Wild., Miss. Em. Laurent: 89 (1905). T: Zaire: Mayumbe, 

Temvo, E.Laurent s.n.; lecto: BR, designated here. 

Parinari klaineana Pierre, in A.Chevalier, Veg. Ut. Afr. Trop. Franç. 9: 202 (1917), as Parinarium 

klaineanum, nom. inval. in synon. 

Illustration: R.Letouzey & F.White, Fl. Cameroun 20 (Fl. Gabon 24): 107 (1978). 


Tree up to 20 m tall. Leaf lamina up to 26 × 12 cm; apex caudate-acuminate; base rounded or 

cuneate; lower surface glabrous. Inflorescence a relatively few-flowered (c. 20) raceme of 

few-flowered cymes. Upper pedicel c. 1.5 cm long. Receptacle tube c. 2.5 × 0.6 cm, often 

curved, glabrous outside, hollow. Sepals about 1.2 cm long, glabrous outside except for 

sparsely and minutely puberulous 'protected' parts. Filaments about 4 cm long. Fruit ellipsoid, 

c. 4 × 2 cm; exocarp glabrescent. 

West Africa in a narrow coastal belt from SE Nigeria to the extreme SW corner of Zaire, just N 

of the River Zaire. 22: NGA. 23: CAB, CMN, CON, EQG, GAB, ZAI. Evergreen rainforest. 

Map 89. 

22. NIGERIA: Calabar-Mamfe, km 117, Onyeachusim & Latilo FHI54211 (K). 23. CABINDA: Buco Zau, 

J.Gossweiler 6889 (BM, BR, COI, K). CAMEROON: Bipinde, G.Zenker 2296 (BM, BR, K, P, W, WAG, WU, 

Z); J.P.M.Brenan & C.F.A.Onochie 9470 (BM, FHO, K, P). CONGO: Kouilou R., 4 km above Kakamoeka, 

C.Farron 4982 (P). GABON: Libreville, R.P.Klaine 2060 (P); Tchibanga, G. Le Testu 1123 (BM, P); Km 12 

Doussala-Tchibanga, J.M.Reitsma 2810 (NY, WAG). ZAIRE: De Brieg 13 (BR, syntype of var. mayumbensis). 

M. gabunensis is the only species of the genus with a truly hollow receptacle. 

6. Maranthes glabra (Oliv.) Prance, Bol. Soc. Brot., sér 2, 40: 184 (1966) 

Parinari glabra Oliv., Fl. Trop. Afr. 2: 370 (1871), as Parinarium glabrum; Ferolia glabra (Oliv.) Kuntze, 

Revis. Gen. Pl. 1: 216 (1891). T: Equatorial Guinea: River Muni, G.Mann 1832; holo: K; iso: P. 

Parinari gilletii De Wild., Ann. Mus. Congo, Bot., Sér. 5, 1: 245 (1906), as Parinarium; Parinari glabra var. 

gilletii (De Wild.) Hauman, Bull. Jard. Bot. État 21: 186 (1951); Maranthes glabra var. gilletii (De Wild.) 

Mendes, Consp. Fl. Angol. 4: 16 (1970). T: Zaire: Kinshasa, J.Gillet 2529; holo: BR. 

Parinari montana Engl., in G.W.J.Mildbraed, Wiss. Erg. Deut. Zeutr-Afr. Exped., Bot. 2: 228 (1911), as 

Parinarium montanum, nom. superfl. illeg. T: Zaire: Beni-Irumu, G.W.J.Mildbraed 2804; holo: B, 

destroyed; lecto: BR, designated here; isolecto: P. 

Parinari tibatensis Engl., Bot. Jahrb. Syst. 46: 141 (1911), as Parinarium tibatense. T: Cameroon: Tibati, 

C.L.Ledermann 2367; holo: B, destroyed. 

Parinari vassonii A.Chev., Explor. Bot. Afrique Occ. Franç. 1: 252 (1920), nom. nud., as Parinarium vassoni. 

Parinari klainei Aubrév., Fl. For. Soud.-Guin. 203 (1950), nom. inval. (without latin description); Parinari 

robusta var. klainei Aubrév. & Pellegr., Not. Syst. (Paris) 14: 59 (1950), nom. inval. (without latin description). 



Tree up to 20 (–40) m tall, but sometimes flowering when only 5 m tall. Leaf lamina up to 14 × 

5.5 cm; apex caudate-acuminate; base cuneate; lower surface usually glabrous, rarely with a 

sparse whitish caducous arachnoid indumentum. Upper pedicel c. 0.5 cm long. Receptacle 

tube c. 0.4 × 0.25 cm. Sepals c. 0.3 cm long. Flower exterior usually glabrous except for the 

whitish-tomentellous 'protected' parts of the sepals. Drupe ellipsoid, c. 4.5 × 3.5 cm; exocarp 

glabrescent. Fig. 60E. 

Widely distributed throughout the greater part of the Guineo-Congolian region from Sierra 

Leone to Ituri in E Zaire and in Cabinda and Angola. 22: GHA, IVO, LBR, NGA, SIE. 23: 

CAB, CAF, CMN, CON, EQG, GAB, ZAI. 26: ANG. Evergreen and semi-evergreen 

rainforest; absent only from the drier types. Map 90. 

22. IVORY COAST: Abidjan, A.Aubréville 127 (BR, K, P). NIGERIA: Sapoba, J.D.Kennedy 2129 (FHO, K). 

LIBERIA: Dukwia R., G.P.Cooper 454 (FHO, K). SIERRA LEONE: Kambui Reserve, King 68 (K). 

23. CAMEROON: 5 km SSW of Mayos, R.Letouzey 3969 (BR, K, P, WAG). CENTRAL AFRICAN 

REPUBLIC: Boukoko near Mbaiki, R.P.C.Tisserant 1779 (P). CONGO: 25 km W of Sibiti, C.Farron 4376 

(BR, P, WAG). GABON: Tchibanga, G. Le Testu 1341 (BM, BR, P); between Cap Esterias and Cap Santa 

Clara, J.M.Reitsma 1455 (NY, WAG). ZAIRE: Lac Tumba, Mpotia, J.Léonard 606 (BM, BR, FHO, K, P). 

The seeds are edible and wood is used for carpentry or canoe-building.


7. Maranthes goetzeniana (Engl.) Prance, Bull. Jard. Bot. Belg. 46: 303 (1976) 

Parinari goetzeniana Engl., Bot. Jahrb. Syst. 34: 153 (1904), as Parinarium goetzenianum. T: Tanzania: 

E Usumbara Mts., H.G.A.Engler 496a; holo: B, destroyed. 


Tree, 20–35 m tall. Leaf lamina up to 15 × 8 cm; apex suddenly and acutely acuminate; base 

rounded to cuneate; lower surface glabrous. Upper pedicel c. 0.2 cm long. Receptacle tube 

0.4 × 0.25 cm to 0.6 × 0.3 cm, tomentellous outside. Sepals 0.3–0.5 cm long, tomentellous 

outside. Fruit ovoid, 2.3 × 1.5 cm; exocarp glabrous. 

Northern Tanzania to Zimbabwe and Mozambique. 25: TAN. 26: MOZ, ZIM. Lowland and 

transitional rainforest on mountains. Map 91. 

25. TANZANIA: E Usumbara Mts., P.J.Greenway 4044 (FHO); E Nguru foothills, Mahonda-Manyangu area, 

Parry 251 (K). 26. MOZAMBIQUE: Serra do Gúruè, R. Licungo, A.R. da Torre & Correia 14844 (LISC); 

Dombe, Maronga, J.S.Simão 449 (LISC). ZIMBABWE: E Vumba Mts., N.C.Chase 6639 (BR, COI, LISC, P, 

SRGH); Chimanimani Mts., Haroni-Makurupini Forest, H.Wild et al. 6669 (BR, FHO, K, P). 

8. Maranthes kerstingii (Engl.) Prance, Bull. Jard. Bot. Belg. 46: 303 (1976) 

Parinari kerstingii Engl., Bot. Jahrb. Syst. 46: 140 (1911), as Parinarium. T: Togo, O.Kersting 547; syn: B, 

destroyed; lecto: K, fide G.T.Prance & F.White, Philos. Trans. B320: 128 (1988); isolecto: BM, P. 



Tree up to 20 m tall. Leaf lamina up to 14 × 6 cm; apex caudate-acuminate; base cuneate or 

rounded; lower surface glabrous. Upper pedicel c. 0.3 cm long. Receptacle tube c. 1 × 0.4 cm, 

glabrous outside. Sepals c. 0.5 cm long, sparsely and minutely puberulous outside on exposed 

parts, whitish-tomentellous on 'protected' parts. Fruit ellipsoid, c. 2.5–5 × 1 cm; exocarp 

glabrescent. 

Sudanian Region from Togo to the Central African Republic. 22: GHA, NGA, TOG. 

23: CAF, CMN, ZAI. Fringing forest in the wetter parts of the Sudanian region. In West 

Africa occurs mainly in upland areas. Map 92. 

22. GHANA: Togo Hills, Shiare, J.Jenik & J.B.Hall cc1173 (K). NIGERIA: Milliken Hill, Engu headwater 

area, F.N.Hepper 2228 (BR, K, P); Adamawa, Mai Idanu, R. Mbam, M.G.Latilo & B.O.Daramola FHI34473 

(FHO, K, P). TOGO: Sokodé-Basari, O.Kersting 320 (K); Atakpame, Doering 297 (K). 23. CAMEROON: 

Bayama, R.Letouzey 3465bis (BR, K, P); 10 km S of Ngaoundéré, A.J.M.Leeuwenberg 7686 (BR, K, P, WAG). 

CENTRAL AFRICAN REPUBLIC: between Nanu and Buar, G.W.J.Mildbraed 9307 (K). 

Several local medicinal properties. 

Parinari kerstingii sensu Aubrév. refers to Maranthes glabra (Oliv.) Prance. 

9. Maranthes panamensis (Standl.) Prance & F.White, Brittonia 37: 76 (1985) 

Couepia panamensis Standl., Trop. Woods 42: 22 (1935). T: Panama: San Blas Distr., Perme, G.P.Cooper 

279; holo: F; iso: GH, NY, US. 

Maranthes corymbosa non Blume (1825): G.T.Prance, Fl. Neotrop. Monogr. 9: 203 (1972) 

Illustration: G.T.Prance, Fl. Neotrop. Monogr. 9: 203 (1972), as M. corymbosa. 


Tree, up to 25 m. Leaf lamina up to 13 × 4.5 cm; apex caudate-acuminate; base cuneate; 

venation prominent on both surfaces; lower surface glabrous. Inflorescence a few-flowered 

(c. 20) raceme of cymes. Upper pedicel c. 0.7 cm long. Receptacle tube c. 0.5 × 0.3 cm. 

Sepals c. 0.3 cm long. Flower exterior sparsely and minutely puberulous except for the 

whitish-tomentellous 'protected' parts of the sepals. Drupe narrowly obovoid, c. 5 × 2 cm. 

Fig. 61. 

Panama (Isthmus), and recently collected in Nicaragua and Costa Rica. 80: COS, NIC, PAN. 

Dense rainforest. Map 93. 

63

64 


Figure 61. Maranthes panamensis. A, habit; B, flower bud; C, flower; D, fruit (A–C, 

Aguilar & Quesada 3795; D, Robles et al. 2060). Scale bars: A, D = 3 cm; B, C = 5 mm. 

Drawn by Margaret Tebbs.


80. COSTA RICA: Puntarenas, Cantón de Osa, RF Golfo Dulce, R.Aguilar 1272 (K, MO); Heredia, P. N. 

Braulio Carrillo, Magsasay, G.Carballo 141 (K, MO); Limón, Cordillera de Talamanca, Matina, G.Herrera 

2187 (K, MO). NICARAGUA: Zelaya, Río Barberena, 5 km from Nueva Guinea, M.Araquistain 142 (MO). 

PANAMA: Prov. Panamá, Cerro Jefe, near Eneida, M.D.Correa & R.L.Dressler 958 (NY, MO); Canal Zone, 

9 km NW of Gamboa, M.Nee 7675 (MO, NY); Cerro Azure, J.P.Folsom, A.Gentry & D.Daly 1949 (MO, NY). 

10. Maranthes polyandra (Benth.) Prance, Bol. Soc. Brot., sér. 2, 40: 184 (1966) 

Parinari polyandra Benth., in J.D.Hooker, Niger Fl. 333 (1849), as Parinarium polyandrum; Ferolia 

polyandra (Benth.) Kuntze, Revis. Gen. Pl. 1: 216 (1891). T: Nigeria, T.Vogel 3; holo: K. 

Parinari polyandra var. cinerea Engl., Bot. Jahrb. Syst. 17: 87 (1893), as Parinarium polyandrum var. cinereum. 

T: Sudan: SW of Tondi, G.A.Schweinfurth 4067; syn: B, destroyed; lecto: K, designated here; isolecto: P. 

Parinari polyandra var. pleiocarpa Engl., Bot. Jahrb. Syst. 46: 140 (1911), as Parinarium polyandrum var. 

pleiocarpum. T: Togo: Sokode-Basari, O.Kersting 84; syn: B, destroyed; lecto: K, designated here. 

Parinari baoulensis A.Chev., Explor. Bot. Afrique Occ. Franç. 1: 250 (1920), as P. baoulense, nom. nud. 

Parinari polyandra var. villosa Aubrév., Fl. For. Soud.-Guin. 207 (1950), nom. inval. 

Parinari polyandra var. argentea Aubrév., loc. cit., nom. inval. 



Tree, rarely more than 7 m tall. Leaf lamina up to 15 × 8.5 cm; apex obtuse or apiculate; base 

rounded or obtuse; lower surface persistently grey-arachnoid-tomentellous or almost glabrous. 

Upper pedicel 0.3–0.4 cm long. Receptacle tube c. 0.4 × 0.4 cm. Sepals c. 0.4 cm long. 

Flower exterior grey-tomentellous. Fruit ellipsoid, 2.5 × 2 × 1.5 cm; exocarp glabrescent. 

Widespread in the Sudanian region from the Ivory Coast and eastern Mali to the Sudan 

Republic. 22: BEN, BKN, GHA, IVO, MLI, NGA, TOG. 23: CAF. 24: SUD. Woodland and 

wooded grassland. Map 94. 

22. BENIN: Parakou-Djougou, G.Cremers 881 (BR, P). BURKINA: Bobo, Dioulasso-Banfora, km 32, 

A.J.M.Leeuwenberg 4410 (K, P, WAG). GHANA: Yendi, N.C.McLeod 814 (K). IVORY COAST: P. N. Bouna, 

Téhini, 40 km E of Ouangofétini, W.J.J.O. de Wilde 784a (K, P, WAG). MALI: Sikasso, J.G.Adam 15082 (P). 

NIGERIA: Enugu Onitsha, km 38, F.White 8418 (FHO). TOGO: Hohoe, J.A.Beveridge 2932 (FHO). 

23. CENTRAL AFRICAN REPUBLIC: Kourou, A.Chevalier 8286 (BR, K, P). 24. SUDAN: Abu Satta Hills, 

N of Tambura, J.G.Myers 7107 (K). 

11. Maranthes robusta (Oliv.) Prance, Bull. Jard. Bot. Belg. 46: 306 (1976) 

Parinari robusta Oliv., Fl. Trop. Afr. 2: 370 (1871), as Parinarium robustum; Ferolia robusta (Oliv.) Kuntze, 

Revis. Gen. Pl. 1: 216 (1891). T: Nigeria: Nun R., G.Mann 481; holo: K; iso: P. 


Tree, up to 12 m tall. Leaf lamina up to 12.5 × 6 cm; apex subacuminate; base cuneate; lower 

surface persistently pale-fulvous-arachnoid-tomentellous. Upper pedicel 0.3–0.4 cm long. 

Receptacle tube c. 0.3 × 0.3 cm, tomentellous with cinnamoneous hairs. Sepals c. 0.4 cm long, 

densely puberulous with greyish hairs. Fruit ellipsoid, 3–3.4 × 2–2.5 cm, sometimes only one 

seed developing; exocarp glabrescent, occasionally with few large round lenticels on exterior. 

West Africa from the Ivory Coast to Nigeria. 22: BEN, GHA, IVO, NGA, TOG. Swamp 

forest and semi-evergreen rainforest. Map 95. 

22. BENIN: Dogba, banks of R. Ouénié, G. Le Testu 239 (BM, BR, P). GHANA: Ankaful, J.B.Hall 1669 (K); 

Volta R., Senchi, F.R.Irvine 2625 (FHO). IVORY COAST: Dabou, A.Chevalier 16205 (BR, P); 6 km NE of 

Béréby, R.A.A.Oldeman 465 (BR, K, P, WAG). NIGERIA: Onitscha, A.H.Unwin 72 (K); Kabba, Kotun Karafi 

F. R., A.T.Johnstone K22 (FHO). 

12. Maranthes sanagensis F.White, Adansonia, sér. 2, 16: 232 (1976) 

T: Cameroon: 7 km N of Olembé, Sanaga R., A.J.M.Leeuwenberg 5476; holo: P; iso: BR, K, WAG. 

Illustrations: F.White, Adansonia, sér. 2, 16: 233 (1976); R.Letouzey & F.White, Fl. Cameroun 20 (Fl. Gabon 

24): 121 (1978). 


65

66 


Tree, 6–8 m tall. Bole short, but up to 1 m in diameter, often twisted and prostrate. Leaf 

lamina up to 15 × 9.5 cm; apex abruptly cuspidate; base subcordate; lower surface glabrous. 

Inflorescence a relatively few-flowered (c. 20) simple or branched raceme of few-flowered 

cymes. Upper pedicel c. 1.2 cm long. Receptacle tube c. 1.5 × 0.5 cm., grey-velutinoustomentellous 

outside. Sepals c. 1 cm long, grey-velutinous-tomentellous outside. Filaments 

c. 6 cm long. Fruit ellipsoid, 6.3 × 2.5 × 1.8 cm; exocarp lenticellate, glabrescent. 

Cameroon (banks of the Sanaga River and the lower courses of its tributaries). 23: CMN. 

Gallery forest. Map 96. 

23. CAMEROON: Sanaga R., 70 km SSW of Bafia, R.Letouzey 9809 (BR, K, P); Nachtigal, Sanaga R., 

J.Raynal 10531 (P); Yangafok, 25 km ENE of Bafia, R.Letouzey 9606bis (BR, K, P, WAG); 7 km N of 

Olembé, 60 km N of Yaoundé, Sanaga R., A.J.M.Leeuwenberg 5478 (BR, K, P, WAG). 

Unplaced Names 

Parinari poggei Engl., Bot. Jahrb. Syst. 26: 378 (1899), as Parinarium. 

T: ?Mukenge, Pogge 914; holo: B, lost. 

Probably a Maranthes. 

14. ATUNA 

Atuna Raf., Sylva Tellur. 153 (1838); derived from the Moluccan vernacular name 'Atun'. 

Type: Atuna racemosa Raf. 

Cyclandrophora Hassk., Flora 25(2), Beibl. 1: 47 (1842); Moquilea sect. Cyclandrophora (Hassk.) Endl., 

Gen. Pl., Suppl. 3: 103 (1843); Parinari subg. Cyclandrophora (Hassk.) Blume, Mélanges Bot. 2: 10 (1855); 

Parinari sect. Cyclandrophora (Hassk.) Müll.Hal., Ann. Bot. Syst. 4(5): 645 (1863). T: Cyclandrophora 

glaberrima Hassk. 

Parinari subg. Macrocarya Miq., Fl. Ned. Ind. 1: 354 (1855). T: not designated. 

Entosiphon Bedd., Madras J. Lit. Sci., Ser. 3, 1: 44 (1864). T: Entosiphon indicum Bedd. 

Atunus Lam., Encycl. 1: 329 (1783), nom. inval. (see discussion below). 

Trees up to 45 m tall; ultimate shoots with a complicated system of divaricate branching. 

Leaves with stipules large, prominently keeled, lateral, persistent or subpersistent; petiole 

eglandular; lamina with a pair of glands on the midrib at or near its base on lower surface; 

lower surface almost glabrous, with minute papillae on veins giving a beaded appearance. 

Inflorescence a raceme or sparsely branched contracted panicle. Bracts and bracteoles 

conspicuous, persistent, eglandular, not enclosing the flower-buds in groups. Flowers 

bisexual, zygomorphic. Receptacle tube obconic, as long as to much longer than the sepals, 

hollow, hairy inside throughout; throat blocked by long retrorse hairs. Sepals 5, broadly 

ovate to lanceolate, apiculate, keeled, tomentellous on both surfaces. Petals 5, glabrous on 

both surfaces. Stamens 10–25, posterior; filaments inserted on margin of faucal annulus, free 

to base, exserted, up to twice as long as sepals; staminodes forming a barely visible 

denticulate margin to the faucal annulus. Ovary monocarpellary, 2-locular, becoming 

unilocular in fruit owing to development of a single ovule, inserted at mouth of receptacle 

tube. Drupe large; epicarp glabrous, densely verrucose; mesocarp transversely fibrous; 

endocarp hard, thick, shortly and sparsely hairy inside; breaking up irregularly at 

germination. Cotyledons large and strongly ruminate. 

Southern India eastwards to Samoa. Kostermans recognised 11 species but the differences 

between some are slight and only 8 are recognised here. The species now placed in Atuna 

were formerly in Parinari, but are quite distinct (see G.T.Prance & F.White, 1988: 130–135). 

Germination is cryptocotylar, eophylls alternate.

CHRYSOBALANACEAE (Atuna) 

The generic name is derived from the Moluccan vernacular name 'Atun'. Rumphius, in his 

Herbarium Amboinense, latinized this to 'Atunus', but used this in two quite different senses. 

In vol. 1: 171, t. 66 (1741) his 'Atunus alba' applied to the plant in Chrysobalanaceae here 

referred to Atuna racemosa Raf., whereas in vol. 3: 96, t. 63 (1743) his 'Atunus litorea' was 

what is now Heritiera littoralis Lam. in Sterculiaceae. Being published before 1753, neither 

of these names is valid. However, both were taken up separately by Lamarck in his 

Encyclopedia, and much confusion has resulted. 'Atunus' of Lam., Encycl. 1: 329 (1783) is 

the Chrysobalanaceous plant, while 'Atunus' of Lam., Encycl., Suppl. 1: 536 (1811) is the 

Heritiera. The nomenclatural arguments by Panigrahi & Purohit (1983) are open to serious 

doubt. Lamarck's intentions over some of his apparent new generic names is questionable, 

especially when the list of generic names at the end of each volume is taken into account. 

The name 'Atunus' is not listed as a generic name on p. 750 of vol. 1 of the Encyclopedia, 

although it might appear to be validated by a 'descriptio generico-specifica' on p. 329, and 

we prefer to treat it as not accepted by the author and so not validly published. Some other 

comparable names in the same work, however, are currently treated as valid names. If Atunus 

Lam. 1783 were regarded as valid, it would be homotypic with Atuna Raf. 1838 which would 

then be an orthographic variant of it. 'Atunus' Lam. 1811, said by the author to be a 'nom 

malais', seems to be not accepted and probably is to be treated as accompanied by only a 

specific and not generic description, and so is also not validly published. The name Atuna 

Raf., which has been in common use for the last 35 years, is confirmed as the earliest generic 

name available for this genus and should continue to be used. 

A.J.G.H.Kostermans, A monograph of the genera Maranthes Bl. and Cyclandophora Hassk. 

(Chrysobalanaceae) of the Asiatic and Pacific area, Candollea 20: 103–158 (1965); Atuna 

Raf. versus Cyclandophora Hassk. (Rosaceae-Chrysobalanoideae), Reinwardtia 7: 421–422 

(1969); G.T.Prance, Chrysobalanaceae, Fl. Males., Ser. 1, 10(4): 635–678 (1989); G.T.Prance 

& F.White, The genera of Chrysobalanaceae, Philos. Trans., B, 320: 130–135 (1988). 

1 Leaf apex rounded; primary veins 6–8 pairs [Fiji] 2. A. elliptica 

1: Leaf apex acuminate or acute; primary veins usually more than 10 pairs 

[India to Pacific, including Fiji] 

2 Receptacle tube cylindrical and narrow 

3 Leaves broadly elliptic, 8–10 cm broad; rounded at base; apex shortly 

acuminate with acumen 2–3 mm long 3. A. latifrons 

3: Leaves oblong, 2.5–6 cm broad; subcuneate to rounded at base; apex 

with long thin acumen 4–22 mm long 

4 Receptacle 8–13 mm long; leaf apex long acuminate; base rounded 4. A. nannodes 

4: Receptacle 5–7 mm long; leaf apex short acuminate with acumen 

3–10 mm long; base cuneate 5. A. penangiana 

2: Receptacle tube funnel-shaped to campanulate 

5 Leaves broadly ovate, thickly coriaceous, cordate at base, 4.5–12 cm 

long 6. A. cordata 

5: Leaves usually elliptic, chartaceous to thinly coriaceous, usually 

rounded at base (if cordate then exceeding 10 cm in length) 

6 Fertile stamens 15–20; inflorescence tomentellous [not in India] 1. A. racemosa 

6: Fertile stamens 12–14; inflorescence sericeous or sparsely pilose 

[India] 

7 Inflorescence sparsely pilose; leaves elliptic-lanceolate, with 8–10 

pairs of primary veins 7. A. indica 

7: Inflorescence densely sericeous; leaves lanceolate, with 12–16 

pairs of primary veins 8. A. travancorica 

67

68 


1. Atuna racemosa Raf., Sylva Tellur. 153 (1838) 

T: Indonesia: Amboina; holo: plate 66 in G.E.Rumphius, Herb. Amboin. 1 (1741). 

Bole fluted or not. Branches glabrous to strigose when young. Leaves with stipules 8–20 mm 

long, lanceolate, acute, subpersistent; petioles slender or thick, 3–7 mm long, glabrous, 

glabrescent or puberulous; lamina chartaceous to coriaceous, 4.5–25 (–35) × 2–11 cm, ovate, 

oblong-ovate, elliptic, oblong or even lanceolate, acuminate at apex, rounded subcordate or 

subcuneate at base, glabrous on both surfaces when mature; midrib prominent on both 

surfaces; primary veins 9–13 pairs, straight or arcuate, prominulous above, prominent 

beneath. Inflorescences of axillary racemes or little branched, to 15 cm long; rachis 

tomentellous to sericeous. Bracts and bracteoles c. 3–8 mm long, ovate to oblong, caducous 

or persistent. Flowers 8–17 mm long. Receptacle turbinate-campanulate, 4–10 mm long, 

tomentose to sericeous on exterior. Calyx lobes ovate to ovate-oblong, 4–7 mm long, 

sericeous or tomentellous on exterior, tomentellous within. Petals ovate-oblong to oblong, 

blue or white, 5–10 mm long. Stamens 13–20, 8–15 mm long, with tooth-like staminodes 

opposite. Style equalling filaments; stigma small. Ovary pilose or densely villous. Fruit 

ellipsoid to subglobose to slightly pyriform; epicarp crustaceous verrucose; endocarp thin, 

densely pilose within. 

Leaf lamina 10–25 (–35) cm long, usually elliptic, oblong or lanceolate but 

sometimes ovate, chartaceous or thickly coriaceous; apex long, finely 

acuminate, 6–25 mm long; petioles thick; flowers 10–17 mm long; medium 

to large trees often with fluted bole 1a. subsp. racemosa 

Leaf lamina 4.5–12 cm long, usually ovate or oblong-ovate, subcoriaceous 

or coriaceous; apex bluntly acuminate, 3–10 mm long; petioles thin; flowers 

8–11 mm long; large trees with cylindrical bole 1b. subsp. excelsa 

1a. Atuna racemosa Raf. subsp. racemosa 

Chrysobalanus racemosus Roxb., Fl. Indica, ed. 2, 2: 506 (1832). T: Indonesia: Mollucas, cult. in Calcutta 

Botanical Garden, W.Roxburgh s.n.; holo: BM. 

Cyclandrophora glaberrima Hassk., Flora 25(2), Beibl. 1: 47 (1842); Parinari glaberrima (Hassk.) Hassk., 

Tijdschr. Natuurl. Gesch. Physiol. 10: 147 (1843), as Parinarium glaberrimum; Petrocarya glaberrima 

(Hassk.) Miers, J. Linn. Soc. Bot. 17: 336 (1879); Ferolia glaberrima (Hassk.) Kuntze, Revis. Gen. Pl. 1: 216 

(1891). T: Indonesia: Jawa, cult. in Hort. Bogor IV H6, A.J.G.H.Kostermans 11105; neo: BO, designated 

here; isoneo: A, BISH, BRI, CAL, CANB, K, KEP, L, LAE, NY, P, PNH, US. 

Parinari scabra Hassk., Tijdschr. Natuurl. Gesch. Physiol. 10: 147 (1843), as Parinarium scabrum, nom. nud.; 

Hassk., Cat. Hort. Bot. Bogor. 269 (1854), ined.; Hassk., Flora 27: 585 (1844), diagnosis; Ferolia scabra 

(Hassk.) Kuntze, Revis. Gen. Pl. 1: 216 (1891); Petrocarya scabra (Hassk.) Miers, J. Linn. Soc., Bot. 17: 336 

(1879); Cyclandrophora scabra (Hassk.) Kosterm., Candollea 20: 126 (1965); Atuna scabra (Hassk.) Kosterm., 

Reinwardtia 7: 422 (1969). T: Jawa, A.J.G.H.Kostermans & Anta 1136; neo: BO, designated here. 

Parinari lanceolata Teijsm. & Binn., Cat. Hort. Bot. Bogor. 253, 255 (1866), nom. nud., as Parinarium 

lanceolatum; Parinari scabra var. lanceolata Koord. & Valeton, Bijdr. Boomsoort. Java 5: 338 (1900), as 

Parinarium scabrum var. lanceolatum. T: Jawa: without collector's name, Hort. Bogor IV H9; lecto: BO, 

fide A.J.G.H.Kostermans, Candollea 20: 127 (1965). 

Parinari margarata A.Gray, U.S. Expl. Exped., Phan. 1: 489 (1854), as Parinarium margaratum. T: Fiji: 

Sandalwood Bay, C.Wilkes s.n.; holo: US; iso: GH, fragment. 

Parinari laurina A.Gray, U.S. Expl. Exped., Phan. 1: 490 (1854), as Parinarium laurinum; Cyclandrophora 

laurina (A.Gray) Kosterm., Candollea 20: 135 (1965). T: Samoa: US Pacific Exploring Expedition s.n.; 

holo: GH; iso: NY, P. 

Parinari elata King, J. Asiat. Soc. Bengal, Pt. 2, Nat.Hist. 66: 280 (1897), as Parinarium elatum; 

Cyclandrophora elata (King) Kosterm., Candollea 20: 122 (1965); Atuna elata (King) Kosterm., Reinwardtia 

7: 421 (1969). T: Malaya: Peninsular Malaysia, Perak, Larut, King's Collector 3436; holo: K; iso: DC, L, 

LE, P, SING. 

Parinari macrophylla Teijsm. & Binn., Cat. Hort. Bot. Bogor. 256 (1866), nom. nud.; Parinari scabra var. 

macrophylla Teijsm. & Binn. ex Koord. & Valeton, Bijdr. Boomsoort. Java 5: 338 (1900). T: Jawa. 

Parinari hahlii Warb., Tropenpflanzer 6: 370 (1902), as Parinarium. T: Caroline Is., Ponape, O.Warburg 

s.n.; holo: B, destroyed.


Parinari mindanaensis Perkins, Fragm. Fl. Philipp. 119 (1904), as Parinarium mindanaense. T: Philippines: 

Mindanao, O.Warburg 14484; holo: B, destroyed. 

Parinari racemosa Merr., Publ. Bur. Sci. Gov. Lab. 17: 19 (1904), nom. illeg., non Vidal (1880); Parinari 

curranii Merr., Philipp. J. Sci., Bot. 4: 264 (1909), as Parinarium. T: Philippines: Masbate Is., R. Marintoc, 

E.D.Merrill 2614; holo: not designated. 

Parinari warburgii Perkins ex Merr., J. Straits Branch Roy. Asiat. Soc. 76: 82 (1917), as Parinarium. 

T: not designated. 

Atunus alba Rumph., Herb. Amboin. 1: 171, t. 66 (1741), nom. inval. 

Parinari amboinensis Teijsm. & Binn., Cat. Hort. Bot. Bogor. 254 (1866), as Parinarium amboinense, nom. nud. 

Illustrations: A.J.G.H.Kostermans, Candollea 20: 149, 155, 157 (1965). 

Map: G.T.Prance, Fl. Males., Ser. 1, 10(4): 670 (1989). 

Branches glabrous or appressed strigose when young. Leaves with stipules stiff, to 20 mm 

long, glabrous to strigose; petioles thick, 3–7 mm long, glabrous or pilose glabrescent; 

lamina usually chartaceous, more rarely stiffly coriaceous, broadly ovate, elliptic, oblong or 

even lanceolate, 10–25 (–35) × 3.5–11 cm, acuminate with acumen 6–25 mm long, rounded 

to subcordate at base, sometimes sparsely strigose beneath on lower portion when young; 

primary veins 10–13 pairs, straight or arcuate; venation conspicuously papillose and often 

giving leaf a scabrous appearance. Inflorescences of axillary racemes or little branched with 

up to 3 racemose branches on short main peduncle, 5–15 cm long; rachis tomentellous to 

sericeous. Bracts and bracteoles ovate, acute, to 8 mm long, caducous. Receptacle 5–10 mm 

long, tomentose to sericeous on exterior; pedicels 0.5–1 mm long. Calyx lobes 4–7 mm long, 

ovate to ovate-oblong, densely tomentellous on both surfaces. Petals ovate-oblong, to 10 mm 

long, blue or white. Stamens 15–20, pale blue, to 15 mm long. Ovary densely villous. Fruit 

ellipsoid to subglobose, to 7.5 cm diam.; mesocarp to 11 mm thick, endocarp 1–3 mm. 

Thailand to the Pacific, including Malay Peninsula (Perak, Singapore), Sumatera, Borneo 

(Sarawak, Brunei), Sulawesi, Philippines, Maluku (Amboin, Seram, Ternate), New Guinea, 

New Britain, Solomon Is., Caroline Is., Fiji, Samoa, Tonga. 41: THA. 42: BOR, jaw, MLY, 

MOL, PHI, SUL, SUM. 43: BIS, NWG, SOL. 60: FIJ, SAM, TON. 62: CRL. Well-drained 

lowland or hill forest up to 600 m, above riverbeds, freshwater or brackish swamp. 

Map 97. 

41. THAILAND: Pen, Narathiwat, Waeng, S.Phusomsaeng 413 (L). 42. BORNEO: Sabah, G.H.S.Wood 

SAN-A3492 (K, L). MALAYA: S Kedah, E of summit of A Gn. Boungsn F. R., L.H.Shing FRI6958 (K, L). 

PHILIPPINES: Elphinstone Prov., Tawao, A.D.E.Elmer 20714 (BISH, BO, BRI, C, D, F, GH, K, L, MO, PNH, 

S, SING, Z). 43. BISMARCK ARCHIPELAGO: New Britain, W Nakanai, Malalia near Cape Hoskins, 

A.Floyd NGF6567 (BO, GH, K, L). NEW GUINEA: W Irian, Fakfak, Anahasi, bb32693 (A, BO, L); Papua 

New Guinea, W Division, Oriemo R., K.J.White & E.Gray NGF10373 (BO, K, L). SOLOMON IS.: Santa 

Ysabel, T.C.Whitmore BSIP2324 (K, L). 60. FIJI: Viti Levu, Mbua, A.C.Smith 4723 (BISH, GH, K, L, LE, 

S, US). 62. CAROLINE IS.: Truk, C.Y.Wong 273 (BISH, K, US). 

The fruit (cotyledon) is grated and used for caulking canoes, widely used in the Pacific 

Islands. An oil is extracted from the seeds and used variously in different areas. The leaves 

are used to thatch the outside walls in Fiji. The wood is used for posts and poles, but is not of 

good quality. 

For a note about the typification of the name Cyclandrophora glaberrima Hassk., see 

A.J.G.H.Kostermans (1965). Since he did not choose a type one is chosen here following the 

reasoning in Kostermans' discussion, that the name was based on a specimen growing in 

Bogor Botanic Garden. A widely distributed specimen is chosen as neotype. 

Kostermans (1965a: 141) dismissed Chrysobalanus racemosus Roxb. as a 'mixtum 

compositu' and stated that he had not seen a type specimen. While the description of the fruit 

does not fit Chrysobalanaceae, there is a type specimen at BM which is certainly this 

subspecies of Atuna racemosa. Chrysobalanus racemosus Roxb. (1832) is an older name 

than Atuna racemosa Raf. (1838), but since this epithet is already occupied in Atuna by the 

Rafinesque name this must stand and a new combination to acknowledge the Roxburgh name 

cannot be made. 

69

70 


1b. Atuna racemosa subsp. excelsa (Jack) Prance, Fl. Males. Ser. 1, 10(4): 670 (1989) 

Petrocarya excelsa Jack, Malayan Misc. 2(7): 66 (1822); Parinari jackiana Benth., in W.J.Hooker, Niger Fl. 

335 (1849), as Parinarium jackianum; Ferolia jackiana (Benth.) Kuntze, Revis. Gen. Pl. 1: 216 (1891); 

Cyclandrophora excelsa (Jack) Kosterm., Candollea 20: 128 (1965); Atuna excelsa (Jack) Kosterm., 

Reinwardtia 7: 422 (1969). T: Sumatera, A.J.G.H.Kostermans & Anta 1136; lecto: BO, fide 

A.J.G.H.Kostermans, Candollea 20: 129 (1965); isolecto: A, K, L, PNH, SING. 

Parinari asperula Miq., Fl. Ned. Ind., Eerste Bijv. 115 (1860), as Parinarium asperulum, nom. nud; ibid. 307 

(1861), descr.; Ferolia asperula (Miq.) Kuntze, Revis. Gen. Pl. 1: 216 (1891); Cyclandrophora asperula 

(Miq.) Prance ex Kosterm., Candollea 20: 130 (1965). T: Indonesia: Sumatera, Lubu-Alang, J.E.Teijsmann 

s.n.; holo: U; iso: K. 

Parinari spicata King, J. Asiat. Soc. Bengal, Pt. 2, Nat. Hist. 66: 279 (1897), as Parinarium spicatum. 

T: Malaysia: Perak, Ulu Bubong, King's Collector 10326; lecto: SING, designated here; isolecto: CAL, K, 

L, P, Z. 

Parinari maingayi King, J. Asiat. Soc. Bengal 66: 280 (1897), as Parinarium. T: Malaysia: Malay 

Peninsula, A.C.Maingay 618; lecto: L, designated here. 

Parinari villamilii Merr., Philipp. J. Sci., Bot. 10: 308 (1915), as Parinarium; Cyclandrophora villamilii 

(Merr.) Prance ex Kosterm., Candollea 20: 126 (1965); Atuna villamilii (Merr.) Kosterm., Reinwardtia 7: 422 

(1969). T: Philippines: Mindanao, Zamboanga Distr., Margosatubig, Villamil 21863; holo: UC; iso: BO, 

fragment. 

Parinari elliptica Teijsm. & Binn., Cat. Hort. Bogor.: 254 (1854), as Parinarium ellipticum, ined. (not 

effectively published). 

Illustrations: A.J.G.H.Kostermans, Candollea 20: 156 (1965); G.T.Prance & F.White, Philos. Trans. B320: 

131 (1988); G.T.Prance, Fl. Males. Ser. 1, 10(4): 668 (1989). 

Map: G.T.Prance, Fl. Males. Ser. 1, 10(4): 670 (1989). 

Branches sparsely strigose, glabrescent when young, obscurely lenticellate. Leaves with 

stipules 8–15 mm long, sparsely strigose; petioles slender, 3–6 mm long, puberulous, 

glabrescent or glabrous; lamina rigidly chartaceous to coriaceous, ovate to oblong-ovate or 

less frequently oblong, 4.5–12 × 2–5 cm, with acumen 3–10 mm long, subcordate, rounded 

or subcuneate at base; primary veins 9–13 pairs, arcuate with venation papillose giving a 

beaded appearance. Inflorescence of axillary racemes to 7.5 cm long, or little branched with 

2 or more racemose branches on short main peduncle; rachis and branches densely short 

sericeous. Bracts and bracteoles oblong, c. 3 mm long, persistent. Receptacle 4–7 mm long, 

sericeous on exterior. Calyx lobes ovate, equal, to 4 mm long, sericeous on exterior, 

tomentellous within. Petals white to bluish white, oblong, to 5 mm long, caducous. Stamens 

13–18, to 8 mm long. Ovary pilose. Fruit subglobose to slightly pyriform, 5–7 × 3.5–4.5 cm; 

mesocarp fibrous, 5–8 mm thick. Fig. 62. 

Malay Peninsula (Kedah and Trengganu southward), Sumatera, Java, Borneo (Brunei, Sabah, 

Sarawak, Kalimantan), N Sulawesi. 42: BOR, JAW, MLY, PHI, SUL, SUM. 43: NWG. 

Lowland forests on well-drained soils on ridges and hillsides to 750 m alt. Map 98. 

42. BORNEO: Brunei, Ulu Supon, Tutong, P.S.Ashton BRUN854 (BO, FHO, K, L); Sabah, Elphinstone Prov., 

Tawao, A.D.E.Elmer 21501 (BR, K, L, NY, P, PNH, SING, U, UC, Z); W Kalimantan, Nunukan Is., 

A.J.G.H.Kostermans 9166 (A, BM, BO, K, L, NY, P, SING); E Kalimantan, lower Mahakam R., Loa Djanan, 

near Samarinda, A.J.G.H.Kostermans 13127 (A, BM, BO, BRI, CAL, CANB, K, L, LAE, NY, P, PNH, SING). 

MALAYA: Johore, Bukit Naga Mengulor, Sungai Sedili, B.H.Ngadiman SF36898 (K). SULAWESI: Manado, 

Anon. bb17159 (A, BO, L). SUMATERA: Res. Palembang, Banjoeasin, L.J.W.Dorst 13R 1 P 911 (K); 

Keritang, Anon. bb28663 (A, BISH, BO, K, L, NY, SING). 43. NEW GUINEA: Papua New Guinea, 5 km N 

of Kiunga on road to Rumingue, D.B.Foreman & P.Katik NGF48461 (BISH, K, L). 

2. Atuna elliptica (Kosterm.) Kosterm., Reinwardtia 7: 421 (1969) 

Parinari elliptica Kosterm., Reinwardtia 7: 48 (1965); Cyclandrophora elliptica (Kosterm.) Kosterm. & 

Prance, Candollea 20: 120 (1965). T: Fiji: Vitu Levu, Sinzatoka District, Vatukarasa Valley, W.Parham 

s.n.; holo: K. 

Illustration: A.J.G.H.Kostermans, Candollea 20: 146 (1965). 

Tree 7–8 m tall. Branches appressed strigose when young, with numerous small lenticels. 

Leaves with stipules aciculate, to 10 mm long, caducous; petiole 5 mm long, terete, 

eglandular; lamina chartaceous to thinly coriaceous, elliptic, 9.5–15 × 4–6 cm, rounded or


Figure 62. Atuna racemosa subsp. excelsa. A, habit (Shing FRI 6960); B, ½ flower and petal 

(Hamid 11594); C, fruit and emerging seedling; D, fruit section showing ruminate 

cotyledons (C–D, Starzecha s.n.). Scale bars: A, C, D = 5 cm; B = 3 mm. Drawn by 

Rosemary Wise. Previously published in G.T.Prance & F.White, Philos. Trans. B320: 131 

(1988). 

71

72 


rarely acute at apex, rounded at base, glabrous and shiny above, with venation prominulous, 

with adpressed strigose hairs on basal portion beneath, glabrescent; midrib prominent beneath; 

primary veins 6–8 pairs, prominent beneath. Inflorescences of few-flowered panicles; rachis 

and small branches densely appressed grey-strigose pubescent. Bracts and bracteoles 

caducous. Receptacle infundibuliform, c. 5 mm long, densely appressed-strigose on exterior; 

pedicel 2–5 mm long. Calyx lobes ovate-elliptic, obtuse, to 5 mm long, densely sericeous on 

exterior, tomentellous within. Petals early caducous. Stamens c. 20, unilateral with tooth-like 

staminodes opposite; filaments c. 9 mm long. Ovary densely appressed-strigose. Style 

glabrous; stigma truncate. Fruit unknown. 

Fiji (Viti Levu). 60: FIJ. Habitat unknown. Map 99. 

60. FIJI: without locality, J.Horne s.n. (K). 

An oil is extracted from the cotyledons for use in local cosmetics. 

3. Atuna latifrons (Kosterm.) Prance & F.White, Philos. Trans., B320: 132 (1987) 

Parinari latifolia M.R.Hend., Gard. Bull. Straits. Settlem., Ser. 3, 7: 102 (1933), as Parinarium latifolium, 

nom. illeg. non P. latifolium (Oliv.) Exell (1928); Parinari latifrons Kosterm., Reinwardtia 7: 54 (1963); 

Cyclandrophora latifolia Prance ex Kosterm., Candollea 20: 121 (1965); Atuna latifolia Kosterm., 

Reinwardtia 7: 421 (1969). T: Malaya: Kedah-Perak Boundary, Md.Haniff S.F.N.21119; holo: K; iso: BO, 

GH, SING. 


Map: G.T.Prance, Fl. Malesiana, Ser. 1, 10(4): 666 (1989). 

Tree to 5 m tall. Branches densely lanate-tomentellous when young becoming glabrous, 

obscurely lenticellate. Leaves with stipules lanceolate, to 11 mm long, acute, keeled, 

sparsely appressed pubescent; petioles thick, 5–7 mm long, terete, densely brown lanate 

when young; lamina chartaceous, broadly elliptic, 11–13 × 8–12 cm, glabrous and shiny 

above, slightly bullate, glabrous beneath except for sparsely pilose venation, very shortly 

abrupt acuminate at apex with acumen 2–3 mm long, rounded at base then contracted into 

petiole; midrib prominent on both surfaces, slightly pilose towards base above, pilose 

beneath; primary veins 12–14 pairs, prominulous, inset in a groove above, prominent and 

pilose beneath; other venation prominulous. Inflorescences of axillary little-branched panicles 

or spikes, to 5 cm long, densely brown sericeous. Bracts and bracteoles to 15 mm long, 

ovate-lanceolate, acute, densely sericeous on exterior, appressed puberulous within. Receptacle 

tube narrowly cylindrical, 7–11 mm long, sericeous on exterior, sessile. Calyx lobes 

lanceolate to oblong-ovate, 5–10 mm long, unequal, densely sericeous on exterior, tomentellous 

within. Petals obovate narrowed at base, 10–11 mm long. Stamens c. 20, inserted on faucal 

annulus, 2 mm high with tooth-like staminodes opposite; filaments 10–12 mm long. Ovary 

densely strigose. Style slender, glabrous; stigma truncate. Fruit unknown. 

Peninsular Malaysia, on the Kedah-Perak border; known only from the type collection. 42: MLY. 

Habitat unknown. Map 100. 

4. Atuna nannodes (Kosterm.) Kosterm., Reinwardtia 7: 422 (1969) 

Parinari nannodes Kosterm., Reinwardtia 7: 50 (1965); Cyclandrophora nannodes (Kosterm.) Kosterm. & 

Prance, Candollea 20: 122 (1965). T: Borneo: Sarawak, O.Beccari, P.B.2955; holo: BO. 

Illustrations: A.J.G.H.Kostermans, Reinwardtia 7: 58 (1965); Candollea 20: 150 (1965). 


Tree to 20 m, usually smaller, unbuttressed. Branches sparsely appressed hirsutulous-strigose 

when young, soon glabrous, obscurely lenticellate. Leaves with stipules narrowly lanceolate, 

acute, 6–12 mm long, strigose to glabrous, subpersistent; petioles 2–4 mm long, glabrescent, 

eglandular, with lower part swollen, usually curved; lamina thinly coriaceous, oblonglanceolate, 

6.7–19 × 2.5–5.5 cm, glabrous on both surfaces, sometimes slightly bullate above, 

long slender acuminate at apex with acumen 7–22 mm long, rounded at base; midrib 

prominulous above, prominent beneath; primary veins 10–12 pairs, arcuate, prominulous on 

both surfaces or sometimes prominent beneath. Inflorescences axillary racemes 3–7 cm long;


Figure 63. Atuna nannodes. A, habit; B, flower; C, fruit (A–B, Everett 104906A; 

C, Y.C.Chan FRI 13314). Scale bars: A, C = 2 cm; B = 2 mm. Drawn by Margaret Tebbs. 

73

74 


rachis densely sericeous-tomentellous. Bracts and bracteoles lanceolate, 3–7 (–13) mm long, 

persistent, sericeous. Receptacle cylindrical, 8–13 mm long, densely sericeous on exterior, 

sessile; calyx lobes to 6 mm long, unequal, acute, sericeous on exterior. Petals white, 

spathulate to ovate, 8–12 mm long, narrowed to base. Stamens 18–20, black to purple; 

filaments 10–15 mm long, slightly unilateral with tooth-like staminodes opposite. Style to 15 mm 

long, glabrous; stigma capitate. Ovary pilose. Fruit ellipsoid, 3–4 × 1.5 cm, slightly tapered 

to base, crustaceous verrucose on exterior; mesocarp 2–2.5 mm, fibrous, hard; endocarp thin. 

Fig. 63. 

Malay Peninsula and Borneo (Sarawak and Sabah). 42: BOR, MLY. Lowland forest to 500 m, 

on well drained soil. Map 101. 

42. BORNEO: Sabah, Sandakan Distr., SSE of Telupid, D.Sundaling SAN92406 (L, SAN); Sabah, Tuinbun 

Mata Is., H.G.Keith BNBFP6225 (K, L); Sarawak, First Division, Mount Matang, M.Jacobs 5543 (B, BO, BH, 

CANB, G, K, L, PNH, SAR, US); 12 th mile Pennissen Road, Kuching Distr., Sibat ak Luang & Jugah ak Kudi 

S 25250 (K, L). MALAYA: Ulu Selangor, Semangkok F. R., Y.C.Chan FRI13314 (K, L); Johore, Sungai Kaya 

Ara, Mawai-Jemaluang road, E.J.H.Corner SF29329 (BO, K, SING); Negri Sembilan, Gunong Angri, 

R.E.Holttum 9893 (BO, K, SING, UC). 

5. Atuna penangiana (Kosterm. & Prance) Kosterm., Reinwardtia 7: 422 (1969) 

Cyclandrophora penangiana Kosterm. & Prance, Candollea 20: 124 (1965). T: Malaysia: Penang, C.Curtis 

203; holo: SING; iso: K, US. 



Tree to 20 m tall, unbuttressed. Branchlets glabrescent when young, obscurely lenticellate. 

Leaves with stipules lanceolate, acute, to 7 mm long, glabrous, stiff, subpersistent; petioles 

3–5 mm long, eglandular, glabrescent, smooth, not swollen or curved; lamina thinly 

subcoriaceous, oblong to oblong lanceolate, 3.7–13 × 2–5.5 cm, glabrous on both surfaces, 

acuminate at apex with acumen 3–10 mm long, cuneate at base; midrib flattened prominulous 

above, prominent beneath; primary veins 10–13 pairs, arcuate, prominulous on both surfaces. 

Inflorescences axillary racemes 3–7 cm long; rachis densely appressed pilose. Bracts and 

bracteoles sericeous to 10 mm long, persistent. Receptacle cylindrical, 5–7 mm long, 

sericeous pubescent on exterior; calyx lobes acute, 4–5 mm long, slightly unequal. Stamens 

c. 20; filaments to 8 mm long with tooth-like staminodes opposite. Style to 10 mm long, 

stigma capitate. Ovary pilose. Fruit (immature) ellipsoid; epicarp crustaceous, verrucose. 

Malay Peninsula. 42: MLY. Lowland forest on well-drained soil to 600 m especially on 

hillside and ridges. Map 102. 

42. MALAY PENINSULA: Penang Is., Government Hill, M.Haniff SF9108 (BO, K, SING); S Kelantan, 

Sungai Galas, P.F.Cockburn FRI7504 (K, L); Perak, Larut, King's Coll. 3537 (BO, K, L, P, US, SING); 

Compt. 24, Bukit Beruk F. R., K.M.Kochummen FRI2404 (K, L); Pahang, B. Gabing Bentong, F.G.Salleto 

C.F.3920 (K); Johore, Sungli Ayer Hiltum Besas, Gunag Pulai, J.Sinclair 7828 (US); W Kelantan, S Perias, 

near K. May, T.C.Whitmore FRI4125 (K, L). 

6. Atuna cordata Cockburn ex Prance, Brittonia 39: 364 (1987) 

T: Borneo: Sabah, Lahad Datu, mile 2, Silam, Ahmad Talip SAN47687; holo: L; iso: K, SAN. 

Illustration: G.T.Prance, Brittonia 39: 365 (1987). 

Map: G.T.Prance, Fl. Malesiana, Ser. 1, 10(4): 666 (1989). 

Tree to 40 m tall; trunk often with thick buttresses. Branches glabrescent when young, 

inconspicuously lenticellate. Leaves with stipules to 1.7 cm long, very early caducous; petioles 

1–3 mm long, short, thick, glabrous; lamina coriaceous, broadly ovate, 4.5–12 × 3–9.5 cm, 

abruptly acuminate at apex with acumen 1–3 mm long, cordate at base, glabrous and shiny 

above, glabrous beneath; midrib prominulous above, prominent beneath; primary veins 9–12 

pairs, lightly prominulous above, prominulous and glabrous beneath. Inflorescences of 

terminal and subterminal racemes 4–8 cm long, borne in single or more often in paired 

branches, densely tomentellous on exterior, puberulous within. Bracts and bracteoles ovate, 

tomentellous, early caducous. Receptacle 5–7 mm long, conical to campanulate, tomentellous


on exterior, sessile; calyx lobes slightly unequal, tomentellous on both surfaces. Petals c. 7 mm 

long, obovate, glabrous. Stamens c. 10, inserted on 1 side of ring; filaments 10–12 mm long. 

Ovary densely pilose. Style slender, hirsutulous on lower portion. Fruit 6 × 5 cm, ovoid; 

epicarp crustaceous, verrucose; mesocarp 5 mm thick, fibrous, hard; endocarp thin. 

Borneo (Sabah). 42: BOR. Hill forests on ultrabasic rock. Map 103. 

42. BORNEO: Sabah, Lahad Datu, Mt. Silam, Ahmad & Sabirin S4863 (SAN), S49043 (SAN); Sabah, Lahad 

Datu, Mt. Silam, W.Meijer S37935 (SAN); Sabah, Lahad Datu, Mt. Silam, J.Proctor SAN100859 (SAN); 

Sandakan, Tavai Plateau, W.Meijer 39330 (K, L). 

7. Atuna indica (Bedd.) Kosterm., Reinwardtia 7: 422 (1969) 

Entosiphon indicum Bedd., Madras J. Lit., Sci. Ser. 3, 1: 45 (1864); Parinari indica (Bedd.) Bedd., Icon. Pl. 

Ind. Or. 22, tab. 109 (1874), as Parinarium indicum; Ferolia indica (Bedd.) Kuntze, Revis. Gen. Pl. 1: 216 

(1891); Cyclandrophora indica (Bedd.) Prance ex Kosterm., Candollea 20: 124 (1965). 

R.H.Beddome s.n.; holo: K; probable isotype (R.H.Beddome 2686): BM. 

T: India: Wynaad, 

Illustrations: R.H.Beddome, Fl. Sylv. Tab. 191 (1872); Icon. Pl. Ind. Or. tab. 109 (1874); 

A.J.G.H.Kostermans, Candollea 20: 152 (1965). 

Tree, medium sized. Branches glabrous when young. Leaves with stipules lanceolate, 5–6 mm 

long, caducous; petioles 5–7 mm long; lamina elliptic-lanceolate, 18–24.5 × 4–7 cm, slightly 

undulate, glabrous on both surfaces, with 2 glands at junction with petiole beneath, 

acuminate at apex with acumen 7–10 mm long, cuneate at base; midrib prominulous above; 

prominent beneath; primary veins 16–20 pairs, prominulous above, prominent beneath; 

secondary venation conspicuously reticulate on both surfaces, papillate beneath. Inflorescences 

terminal and axillary racemes; rachis sparsely appressed-tomentose. Receptacle campanulatecylindrical, 

sericeous-tomentose on exterior; pedicels 1.25–1.7 mm long; calyx lobes ovate, 

acute, sericeous on exterior, tomentellous within. Petals narrowly ovate. Stamens 12–15, 

unilateral with staminodes opposite; filaments far exserted. Ovary pilose. Style pilose on 

basal portion only. Fruit not seen. 

Confined to the Western Ghats in India. 40: IND. Forests. Map 104. 

40. INDIA: Carcoon Ghat, Wynaad, R.H.Beddome s.n. (K); without locality, J.S.Law s.n. (K); without locality, 

Herb. Schinz s.n. (Z). 

8. Atuna travancorica (Bedd.) Kosterm., Reinwardtia 7: 422 (1969) 

Parinari travancorica Bedd., Icon. Pl. Ind. Or. 1: 43, tab. 189 (1874); Ferolia travancorica (Bedd.) Kuntze, 

Revis. Gen. Pl. 1: 216 (1891); Cyclandrophora travancorica (Bedd.) Prance ex Kosterm., Candollea 20: 125 

(1965). T: India: Travancore, R.H.Beddome 292; holo: K; iso: CAL. 

Illustrations: R.H.Beddome, Icon. Pl. Ind. Or. 1: tab. 189 (1874); A.J.G.H.Kostermans, Candollea 20: 153 (1965). 

Tree to 25 m. Branches pubescent when young. Leaves with stipules lanceolate-linear, acute, 

to 1 cm long, enveloping axillary bud; petiole slender, to 4 mm long, eglandular; lamina 

chartaceous, lanceolate, 7–12 × 1.8–3.5 cm, bluntly acuminate at apex with acumen 3–7 mm 

long, subcuneate at base, glabrous and densely reticulate on both surfaces; midrib 

prominulous above, prominent beneath; primary veins 12–16 pairs, prominulous on both 

surfaces, arcuate, pilose beneath when young; secondary venation papillate beneath. 

Inflorescences racemes to 5 cm long, few-flowered; rachis densely sericeous. Bracts and 

bracteoles caducous. Receptacle funnel-shaped, 5–6 mm long, densely villous-pubescent on 

exterior. Calyx lobes ovate, acute, silky-pubescent, 4 mm long. Petals white or pinkish, 

oblong-ovate, acute. Stamens c. 12; filaments exserted, glabrous. Ovary pilose; style 

glabrous; stigma truncate. Fruit ellipsoid, 5 × 4 cm; exocarp smooth, crustaceous, densely 

tomentose when young, glabrescent; pericarp c. 2 mm thick, densely tomentose within. 

Southern India (Travancore region). 40: IND. Confined to forests. Map 105. 

40. INDIA: Merchiston Estate, Colatoorpolay, T.F.Bowdillon s.n. (CAL); Merchiston Estate, Colatoorpolay, 

T.F.Bowdillon 584 (K). 

75

76 


15. DACTYLADENIA 

Dactyladenia Welw., Apont. 1: 572 (1859) [1858]; from the Greek, 'glandular fingers', 

referring to the stalked glands in some species. 

Type: Dactyladenia floribunda Welw. 

Griffonia Hook.f., in G.Bentham & J.D.Hooker, Gen. Pl. 1: 608 (c. 19 Oct. 1865) non Griffonia Baill., 

Adansonia 6: 188 (7 Oct. 1865, Leguminosae: Caesalpinioideae). T: Griffonia barteri Hook.f. ex Oliv. 

Acioa subg. Afracioa Letouzey, Fl. Cameroun 20 (Fl. Gabon 24): 7 (1978). T: Acioa pallescens Baill. 

Shrubs or small trees, sometimes scandent. Leaves with stipules lateral, often persistent; 

petiole eglandular; lamina usually with 2 glands on lower surface near base, often also with 

smaller glands near apex or scattered over surface; lower surface of lamina glabrous or with 

stiff ferruginous hairs, more rarely with whitish arachnoid or lanate indumentum. 

Inflorescence a simple raceme or of zig-zag (usually lax) false racemes, more rarely a 

subcapitate spike (D. whytei). Bracts and bracteoles not enclosing the flower-buds in small 

groups, often with stalked or sessile glands. Pedicels articulate. Flowers bisexual, strongly 

zygomorphic. Receptacle tube usually elongate and narrowly obconic-tubular, usually longer 

than the upper pedicel, often slightly gibbous at the base, glabrous inside except near the 

throat. Sepals 5, rounded or acute, usually spreading or reflexed, often with stalked or sessile 

marginal glands; adaxial surface whitish-tomentellous. Petals 5, caducous. Stamens 10–75, 

coiled in bud; filaments ligulately connate for most of length, inserted on posterior rim of 

faucal annulus, far-exserted, much longer than combined length of calyx and receptacle tube. 

Staminodes short, filiform or denticulate. Ovary monocarpellary, l-locular, inserted at mouth 

of receptacle tube; style filiform, slightly longer than stamens, slightly 3-lobed at apex. 

Drupe usually more than 3 cm long, ±ovoid, but usually markedly tapered towards the 

pointed apex; epicarp often ferruginous-tomentose and with longer hispid hairs, rarely 

glabrous; mesocarp thin; endocarp thin, hard, with a roughish surface, hairy inside, with no 

special mechanism allowing seedlings to escape. 2n = 22 (D. barteri, as 'Acioa berteri' 

(S.Mangenot & G.Mangenot, 1958; 1962). 

A genus of 30 species in tropical Africa, virtually confined to the Guineo-Congolian region. 

Most species occur in old secondary forest or primary forest, more rarely in coastal thicket. 

From the paucity of herbarium material it seems that several species are very rare. An 

important exception is D. barteri, which plays a significant role in agricultural practice in 

southeast Nigeria. Here it has been found that even on soils inherently low in mineral 

reserves, under a rainfall as high as 2250 mm per year, it is possible to follow 11/2 years of 

cropping with only 21/2 years of fallow of D. barteri (P.H.Nye & P.J.Greenland, 1960). In the 

same region this species is also planted with Anacardium occidentale and Erythrophleum 

suaveolens on badly eroded sites to restore forest conditions. The germination of those 

species examined is cryptocotylar, with opposite eophylls. 

E.A.J. de Wildeman, Contribution a l'étude des espèces africaines du genre Acioa Aubl., 

Bull. Jard. Bot. État 7: 191–216 (1920); R.Letouzey & F.White, Chrysobalanacées nouvelles 

du Cameroun et du Gabon, Adansonia Sér. 2, 16: 238–243 (1976); R.Letouzey & F.White, 

Fl. Cameroun 20 (Fl. Gabon 24): 6–60 (1978); G.T.Prance & F.White, The resurrection of 

the genus Dactyladenia (Chrysobalanaceae), Brittonia 31: 438–487 (1979); G.T.Prance & 

F.White, The genera of Chrysobalanaceae, Philos. Trans. B320: 133–145 (1988). 

1 Inflorescence a simple raceme (but sometimes subsimple in D. jongkindii) 

2 Receptacle tube usually glabrous (or often sparsely puberulous in 

species 2, 11, 12 and 14 or sparsely hispidulous in species 5), exterior 

of sepals glabrous (or sparsely puberulous in species 11 and 12) 

3 Receptacle tube ±equalling upper pedicel (above the articulation) 

4 Branchlets and lower leaf-surface ±hispid; bracteoles 5–7-palmatisect; 

bracts with glandular margin and 1–2 pairs of basal stipitate glands 1. D. smeathmannii 

4: Branchlets and lower leaf-surface glabrous or with sparse hairs on 

midrib and primary veins but not hispid

CHRYSOBALANACEAE (Dactyladenia) 

5 Branchlets, stipules, petioles, primary veins and inflorescence 

puberulous-tomentose; receptacle tube glabrous or sparsely hairy; 

stamens c. 25, 1.5–2.5 cm long 2. D. ndjoleensis 

5: Branchlets, stipules, petioles, primary veins, inflorescence and 

receptacle tube glabrous; stamens c. 15-20, 3–3.5 cm long 

6 Bracteoles ovate-lanceolate, denticulate-glandular; leaves with 

5–8 pairs of lateral nerves 3. D. scabrifolia 

6: Bracteoles lanceolate, 5–7-dentate, with substipitate to stipitate 

glands near the base; leaves with 3–5 pairs of lateral nerves 4. D. letestui 

3: Receptacle tube longer than upper pedicel (above the articulation) 

7 Bracteoles ovate, eglandular (receptacle tube sometimes hispidulose) 5. D. whytei 

7: Bracteoles palmatilobed to palmatisect 

8 Branchlets not densely covered with persistent golden tomentum; 

but variously pubescent, hispidulose or glabrescent 

9 Branchlets with hairs less than 1 mm long 

10 Stipules linear, 0.5–0.6 cm long; leaves with 6–8 pairs of lateral 

nerves; bracteoles 3 (–5)-palmatisect; receptacle tube 1.2–2 cm 

long; sepals 0.6–0.8 cm long; stamens c. 25 

10: Stipules lanceolate, 0.3–0.4 cm long; leaves with 4–5 pairs 

of lateral nerves; bracteoles 5-palmatilobed to palmatipartite; 

receptacle tube 0.7–1 cm long; sepals 0.4–0.5 cm long; 

6. D. gilletii 

stamens c. 15 7. D. librevillensis 

9: Branchlets with hairs more than 1mm long 

11 Stipules ±flat, linear or lanceolate, without inrolled margins; 

lower pedicel and receptacle tube glabrous 

12 Stipules lanceolate, 1–1.6 × 0.3–0.8 cm; lower leaf-surface 

glabrous; lateral nerves in 8–16 pairs 

12: Stipules linear, 0.6–0.8 cm long; lower leaf-surface hispid; 

lateral nerves in 6–10 pairs 

13 Bracteoles 3–5-palmatipartite; receptacle tube 0.8–1.2 cm long; 

10. D. campestris 

sepals 0.6 cm long; androecium 2.5–3 cm long; stamens c. 30 

13: Bracteoles transversely ovate but furnished with (3–) 5 

triangular teeth bearing a stipitate gland; receptacle tube 

1.6–2.2 cm long; sepals 0.6–1 cm long; androecium 

8. D. bellayana 

3.5–4.5 cm long; stamens c. 20 

11: Stipules saccate with inrolled margins; lower pedicel and 

receptacle tube sparsely puberulous 

14 Stipules with distinct, densely pubescent margins, without 

stalked glands at base; lower pedicel 2–7 mm long; bracteoles 

9. D. hirsuta 

3-lobed, 1–2 mm long 

14: Stipules without densely pubescent margins, ±evenly hairy, 

with stalked basal glands; lower pedicel 10–16 mm long; 

11. D. floretii 

bracteoles 5-partite, 4–6 mm long 12. D. jongkindii 

8: Branchlets densely covered with persistent golden tomentum 

15 Raceme 12–30 cm long; leaf-lamina cordate at base; lateral 

nerves in 10–16 pairs; bracteoles 5-palmatipartite to palmatisect 

15: Raceme 2–6 cm long; leaf-lamina rounded at base; lateral nerves 

in 7–10 pairs; bracteoles 5-palmatilobed to palmatipartite 

13. D. icondere 

(receptacle tube sometimes very sparsely pubescent) 14. D. floribunda 

2: Receptacle tube and exterior of sepals densely hairy 

77

78 


16 Receptacle tube ±as long as upper pedicel; leaf lamina glabrous, with 

base acute and lateral nerves in 4–6 pairs; bracteoles lanceolate, 

sometimes glandular 15. D. barteri 

16: Receptacle tube longer than upper pedicel; leaf lamina glabrous to 

densely hairy 

17 Bracteoles subreniform; leaf lamina cordate at base, with lower 

surface densely hairy; lateral nerves in c. 12 pairs 16. D. sapinii 

17: Bracteoles not subreniform; leaf lamina usually rounded to 

subcuneate at base (cordate only in D. pallescens), with lower 

surface glabrous to slightly scabrous; lateral nerves 6–12 pairs 

18 Receptacle tube and exterior of sepals densely silvery-tomentose; 

bracteoles ovate, entire or denticulate-glandular 

19 Receptacle tube c. 1.8 cm long; stamens 60–75; midrib densely 

puberulous to glabrescent below; lateral nerves in 8–12 pairs 17. D. pallescens 

19: Receptacle tube c. 1.5 cm long; stamens 35 or fewer; midrib 

glabrous or glabrescent; lateral nerves in 5–8 pairs 18. D. buchneri 

18: Receptacle tube and exterior of sepals sparsely pubescent to 

densely villous, not densely silvery-tomentose 

20 Branchlets and flowers densely fulvous-villous; bracteoles deltate 

with glandular appendages near base; racemes 3–4.5 cm long, 

few-flowered; lower pedicel c. 0.5 cm long; receptacle tube 

1.2–2 cm long; sepals 0.6–0.8 cm long; stamens 3–5 cm long 19. D. staudtii 

20: Branchlets and flowers sparsely pubescent, sometimes 

hispidulose; bracteoles 5 (–10)-palmatisect; racemes 10–15 cm 

long; lower pedicels 1 cm long; receptacle tube 0.6–1 cm long, 

distinctly gibbous at base; sepals 0.4–0.5 cm long, often with 

marginal glands; stamens 2–2.5 cm long 20. D. dewevrei 

1: Inflorescence compound (but sometimes subsimple in D. jongkindii) 

21 Receptacle tube and exterior of sepals glabrous 

22 Bracteoles subreniform, 5-denticulate, inserted at middle of pedicel 

below the articulation 21. D. laevis 

22: Bracteoles 5–7-palmatifid to palmatipartite; inflorescence sometimes a 

subsimple raceme 22. D. lehmbachii 

21: Receptacle tube and exterior of sepals not glabrous 

23 Bracteoles ovate, without conspicuous glandular denticulation 

(sometimes denticulate in D. mannii) 

24 Inflorescence puberulous; bracteoles inserted near the middle of 

lower pedicel, sometimes denticulate 23. D. mannii 

24: Inflorescence tomentellous to tomentose; bracteoles inserted near 

apex of lower pedicel 

25 Stipules ovate, 0.4–0.8 cm long 24. D. pierrei 

25: Stipules linear to lanceolate 

26 Receptacle tube c. 1.8 cm long; branchlets and inflorescence 

golden tomentose 27. D. dinklagei 

26: Receptacle tube up to 1.2 cm long 

27 Branchlets very sparsely puberulous or glabrescent; lamina 

subcordate or cordate at base 25. D. eketensis 

27: Branchlets tomentose at first, glabrescent; lamina rounded to 

subcordate 26. D. dichotoma 

23: Bracteoles ovate to deltate, with stipitate glands


28 Stipules subdeltate with flabellate nerves; bracts and bracteoles with 

a clavate apical gland 28. D. cinerea 

28: Stipules linear or lanceolate 

29 Branchlets and lower leaf-surface hispid; stipules linear, with longstipitate 

glands; bracts and bracteoles deltate, with long-stipitate glands 29. D. johnstonei 

29: Branchlets and lower leaf-surface arachnoid-tomentose; stipules 

falcate-lanceolate; bracts and bracteoles ovate, with stipitate glands 30. D. chevalieri 

1. Dactyladenia smeathmannii (Baill.) Prance & F.White, Brittonia 31: 486 (1979) 

Acioa smeathmannii Baill., Adansonia 10: 244 (1872). T: Sierra Leone: without precise locality, 

H.Smeathmann s.n.; holo: P, in Herb. Juss. no. 15895. 

Acioa parvifolia Engl., Bot. Jahrb. Syst. 26: 380 (1899). T: Sierra Leone: without precise locality, Afzelius 

s.n.; holo: UPS; photo: K. 

Acioa unwinii De Wild., Bull. Jard. Bot. État 7: 198 (1920). T: Sierra Leone: [Buri Town] Lea Beach, 

Unwin & Smythe 36; holo: K; iso: P; photo: BR. 

Shrub. Branches hispid when young, lenticellate. Leaves with stipules linear, early caducous, 

with stipitate glands when very young; petioles 2–4 mm long; lamina ovate, 6–11 × 3–4.5 cm, 

glabrous beneath except for dense-ferrugineous pubescence on venation, rounded at base, 

shortly acuminate at apex; midrib densely ferrugineous hirsute beneath; primary veins 6–8 

pairs, densely hirsute beneath. Inflorescence terminal racemes, 10–12 cm long; rachis 

hispidulous. Bracteoles 3 mm long, palmatisect, with 5–7 stipitate glands. Upper pedicel 8 mm 

long. Receptacle tube 0.8 cm long, glabrous, gibbous. Sepals 4–6 mm long, glabrous on 

exterior, with marginal stipitate glands, tomentellous within. Petals oval, glabrous, equalling 

sepals. Stamens c. 15, 2–3 cm long. Fruit obovoid, 4 × 2.5 × 2 cm; exocarp with short-densefulvous 

pubescence. 

Sierra Leone and Liberia. 22: LBR, SIE. Old secondary forest and on almost bare rock. 

Map 106. 

22. LIBERIA: Cola-Yoma National Forest, on bank of Maher R., P.P.C. van Meer 273 (BR, WAG). SIERRA 

LEONE: Ronietta, N.W.Thomas 5621 (K), 5502 (K, P). 

Differs from D. johnstonei in its simple inflorescence and glabrous receptacle and sepals. 

2. Dactyladenia ndjoleensis Breteler, Syst. Geogr. Pl. 69: 111 (1999) 

T: Gabon: c. 5–30 km NNW of Ndjolé, F.J.Breteler, C.C.H.Jongkind & J.J.Wieringa 11030; holo: WAG; iso: 

BR, C, G, K, LBV, MA, MO, NY, P, PRE. 

Illustration: F.J.Breteler, Syst. Geogr. Pl. 69: 112 (1999). 

Map: F.J.Breteler, Syst. Geogr. Pl. 69: 113 (1999). 

Tree 8–12 m high. Branches puberulous to tomentellous when young, often also with early 

deciduous arachnoid indumentum. Leaves with stipules narrowly triangular, 4–5 mm long, 

inserted on base of petiole, puberulous to tomentellous; petiole 3–4 mm long, puberulous to 

tomentellous; lamina ovate-elliptic, (6–) 7–9 × 2.5–3.5 (–4) cm, sparsely hairy on midrib and 

primary veins beneath, with margin and lower veins also arachnoid, with small flat glands 

beneath near base and acumen; cuneate to rounded at base, acuminate at apex with acumen 

0.5–1 cm long; midrib hairy beneath; primary veins 6–7 pairs. Inflorescences axillary or 

subterminal racemes; rachis puberulous; bracteoles 1–3 mm long, puberulous, those on 

peduncle 3–fid and others narrowly triangular, with sessile or short-stalked glands; lower 

pedicels 4–7 mm long; upper pedicel 4–6 mm long. Receptacle tube 5–6 mm long, glabrous 

at base adaxially, glabrous to hairy outside. Sepals 5–6 × 2.5–3.5 mm, glabrous or very 

sparsely hairy outside, densely silky-hairy within. Petals 6–9 mm long, glabrous. Stamens 

c. 25, 3–3.5 cm long. Fruit 2.5 cm long × 2 cm diam.; exocarp velutinous. 

Gabon. 23: GAB. Rainforest. Map 107. 

23. GABON: Rabi-Kounga, Echira Rd., F.J.Breteler & C.C.H.Jongkind 10177 (K, LBV, MO, P, WAG). 

This recently described species is close to D. letestui, but clearly distinct. 

79

80 


3. Dactyladenia scabrifolia (Hua) Prance & F.White, Brittonia 31: 486 (1979) 

Acioa scabrifolia Hua, Bull. Mus. Hist. Nat. (Paris) 3: 328 (1897). T: Guinea: Timbo, Caremonga, 

F.A.W.Miquel 24; holo: P. 

Tree to 20 m tall. Branches glabrous when young. Leaves with stipules linear, acute; petioles 

3–5 mm long, glabrous, rugose; lamina oblong, 6–13 (–15) × 3.5–5.5 cm, scabrous above, 

glabrous beneath, cuneate at base, finely acuminate at apex; midrib glabrous beneath; primary 

veins 5–8 pairs, glabrous beneath. Inflorescences terminal and axillary racemes 6–10 cm 

long; rachis shortly hispidulous, glabrescent. Bracts ovate-acute, glabrous; bracteoles 1–5 mm 

long, ovate-lanceolate, glandular-denticulate. Upper pedicel 6–14 mm long. Receptacle tube 

0.4–0.8 cm long, narrowly obconic, gibbous, glabrous on exterior. Sepals 4–6 mm long, 

glabrous. Petals oblong, equalling sepals, white to pink. Stamens c. 20, 1.5–2.5 cm long. 

Fruit ovoid, 4 × 3 × 2 cm; exocarp glabrous. 

Guinea to Mali and Ghana. 22: BKN, GHA, GUI, IVO, LBR, MLI, SIE. 23: CAF. Riparian, 

hill and secondary forest. Map 108. 

22. GUINEA: Mamou, Bouloukountou, S.Lisowski 51583 (BR, K); Kourio, Bilima, A.Chevalier 14889 (BR, K, P); 

Fouta Djalon, J.Langdale-Brown 2627 (BR, K). LIBERIA: Loffa County, Gbarnga-Zorzor road, St. Paul R., 

J.J.Bos 2498 (BR, K, WAG); W Prov., Boporo Distr., Beiden, J.T.Baldwin Jr. 10280A (K); Peáhtah, 

D.H.Linder 1114 (AA, K, P). SIERRA LEONE: Musaia, F.C.Deighton 5405 (K); Mamboma, F.C.Deighton 

3779 (K). 23. CENTRAL AFRICAN REPUBLIC: Sangha-Mbaere, E bank of Sangha River, c. 30 km S of 

Lidjombo, D.Harris 2718 (K). 

Closely related to D. barteri but differing in its stouter peduncle and pedicels, glabrous 

receptacle tube and scabrid leaves. 

4. Dactyladenia letestui (Letouzey) Prance & F.White, Brittonia 31: 485 (1979) 

Acioa letestui Letouzey, Adansonia, sér. 2, 16: 240 (1976). T: Gabon: La Lara, 30 km S of Mitzic, 

G. Le Testu 9352; holo: P; iso: BM, WAG. 

Acioa tessmannii Engl., Pflanzenw. Afr. 3: 311 (1915), nom. nud., based on G.Tessmann 221. 

Illustrations: R.Letouzey, Adansonia, Sér. 2, 16: 239 (1976); R.Letouzey & F.White, Fl. Cameroun 20 

(Fl. Gabon 24): 45 (1978). 

Tree to 25 m tall. Branches glabrous when young, lenticellate. Leaves with stipules linear, 

acute, 3–6 mm long, appearing intrapetiolar, but attached to extreme base of petiole; petioles 

3–5 mm long, transversely rugose, glabrous; lamina elliptic, 6–11.5 × 3.5–6 cm, sometimes 

slightly oblong, glabrous beneath, with 2–3 large circular glands at base and other scattered 

glands near midrib and 2 small glands at acumen, rounded at base, acuminate at apex with 

acumen c. 6 mm long. Inflorescences terminal and axillary racemes, 3–8 cm long, manyflowered, 

entirely glabrous. Bracts ovate to lanceolate, 2–3 mm long, often subglandular at 

margin and glandular at base and sometimes at end of nerves; bracteoles 1–1.5 mm long, 

lanceolate, 5–7-dentate with substipitate glands. Upper pedicel 6–10 mm long. Receptacle 

tube 0.5–0.6 cm long, slightly gibbous, glabrous on exterior. Sepals 4–5 mm long, sometimes 

with glandular margins. Petals white, obovate, obtuse at base, 4–5 mm long. Stamens c. 15, 

2–2.5 cm long. Fruit subglobose, 17 × 20–25 × 20–30 mm; exocarp tomentellous with sparse 

hispid hairs, knobbled when dry. 

Cameroon, Gabon and Equatorial Guinea. 23: CMN, EQG, GAB. Swamp forest. Map 109. 

23. CAMEROON: Doumé R., 40 km SW of Batouri, near Bimba, F.J.Breteler 2795 (BR, K, P, WAG); 30 km 

S of Mbalmayo, W.J.J.O. de Wilde & B.E.E. de Wilde-Duyfjes 1919 (BR, K, P, WAG, Z); Nyamtimbi near 

Doumé, Bertoua region, R.Letouzey 3143 (BR, P, WAG); confluence of Lom-Djérem, J.Mildbraed 8482 (K). 

EQUATORIAL GUINEA: G.Tessmann 221 (K, photo P). GABON: Mbigou, G. Le Testu 6537 (BM, BR, P, WAG). 

Closely related to D. barteri and D. scabrifolia, differing in its globose fruit. 

5. Dactyladenia whytei (Stapf) Prance & F.White, Brittonia 31: 486 (1979) 

Acioa whytei Stapf, J. Linn. Soc., Bot. 37: 97 (1905). T: Liberia: near Kakatown [Monrovia], A.Whyte s.n.; 

holo: K; iso: fragment and photo BR, photo P. 

Acioa stapfiana De Wild., Bull. Jard. Bot. État 7: 204 (1920). T: Sierra Leone: Sherbro, G.H.Garret s.n.; 

holo: K; photo: BR, P.


Shrub (lianescent) or small tree to 10 m tall. Branches hispid when young, soon glabrescent. 

Leaves with stipules ovate-lanceolate, toothed at base; petioles 3 mm long, thick; lamina 

ovate-lanceolate or oblong-lanceolate, 6–15 × 2.5–6 cm, ±hispid on venation beneath; 

rounded at base, acuminate at apex; midrib hispid beneath; primary veins 5–6 pairs, hispid 

beneath. Inflorescence terminal and axillary racemes, 1–3 cm long, densely flowered; rachis 

hispidulous; bracts ovate-lanceolate, 5–6 mm long; bracteoles 1.5 mm long, ovate, acute. 

Upper pedicel 2–8 cm long. Receptacle tube 1.6–2.2 cm long, gibbous, glabrous or 

hispidulous. Sepals 4–6 mm long, glabrous or sparsely hispidulous. Petals oblong-elliptic, 

2-lobed, equalling sepals, white, shortly unguiculate. Stamens c. 30, 3–5 cm long. Fruit 

broadly ovoid, 5 × 3 × 2.5 cm; exocarp tomentose. 

Guinea, Sierra Leone and Liberia. 22: GUI, LBR, SIE. Riparian forest and secondary areas. 

Map 110. 

22. LIBERIA: Devilbush, near Paynesville, 16 km E of Monrovia, A.J.M.Leeuwenberg & A.G.Voorhoeve 4891 

(BR, K, P, WAG); Careysburgh, 35 km from Monrovia, A.M. Van Marten 77 (K, WAG); Grand Bassa Co., 

Fortsville, J.T.Baldwin Jr. 11164 (K). SIERRA LEONE: Baiima, F.C.Deighton 6104 (K, P); Njala, 

F.C.Deighton 1319 (BM, BR, K); Bo F. R., D.Small 191 (K, P). 

This species is distinct in its short, congested, subcapitate racemes with the pedicels 

concealed by foliaceous bracts. 

6. Dactyladenia gilletii (De Wild.) Prance & F.White, Brittonia 31: 485 (1979) 

Acioa gilletii De Wild., Ann. Mus. Congo, Bot., Sér. 5, 1: 47 (1903). T: Zaire: Kinshasa, J.Gillet s.n.; holo: BR. 

Tree to 15 m tall. Branches hispid when young, glabrescent. Leaves with stipules linear, c. 4 mm 

long; petioles 2–3 mm long; lamina oblong-ovate, 8–16 × 3–6 cm, glabrous beneath except 

for short hairs on venation, rounded to subcuneate at base, long acuminate at apex; midrib 

with sparse short hairs beneath; primary veins c. 8, sparsely hairy. Inflorescences axillary and 

terminal racemes 3–5 cm long, glabrous. Bracts ovate-oblong, 3–4 mm long, with stipitate 

glands on margins; bracteoles c. 1.5 mm long, inserted near middle of lower pedicel, palmatisect 

with 3–5 stipitate glands. Upper pedicel 4–7 mm long. Receptacle tube 1.2–2 cm long, 

glabrous on exterior, slightly gibbous at base. Sepals 6–8 mm long, glabrous. Petals 8–9 mm 

long. Stamens c. 25, 2.5–3.5 cm long. Fruit ovoid, 5 × 3 × 2 cm; exocarp yellow, tomentellous. 

Gabon, Lower Congo, Zaire (Bas Zaire, Kasai, Forestier central) and Angola. 23: CON, 

GAB, ZAI. 26: ANG. Forests, especially in humid places along rivers. Map 111. 

23. GABON: Moyen-Ogooué, N of Ndjolé, G.McPherson 15811 (BR, K, MO, NY). ZAIRE: Yangambi, 

J.Louis 5903 (BR, K, P, W); near Kisangani, margin of Tshopo, S.Lisowski 17341 (BR, K); Leopoldville Prov., 

Leo-Kalina, L.Dubois 1544 (BR, K); between Liléko and Basoho, J.Louis 14582 (BR, K). 26. ANGOLA: 

NE of Lunda, J.Gossweiler 13820 (BM, K, P, US); Lunda, Dundo, R.G.N.Young 524 (BM, BR). 

Differs from D. dewevrei in its glabrous inflorescence, longer receptacle tube and (usually) 

3-lobed bracteoles. 

7. Dactyladenia librevillensis (Letouzey) Prance & F.White, Brittonia 31: 486 (1979) 

Acioa librevillensis Letouzey, Adansonia, sér. 2, 16: 238 (1976). T: Gabon: Cap Esterias, H.Klein 55; holo: 

P; iso: B, BR, K, WAG. 

Illustrations: R.Letouzey, Adansonia, Sér. 2, 16: 237 (1976); R.Letouzey & F.White, Fl. Cameroun 20 (Fl. 

Gabon 24): 49 (1978). 

Shrub, scandent. Branches sparsely hispid when young, glabrescent. Leaves with stipules 

attached to base of petiole, lanceolate, 3–4 mm long; petioles 3–5 mm long, hispidulous; 

lamina elliptic, 8–10 × 4–5 cm, sparsely hispid on venation beneath, with 2 large circular 

glands at base beneath and small glands below acumen, rounded to subcuneate at base, 

acuminate at apex with acumen 3–5 mm long; midrib hispid beneath; primary veins 4 or 5 

pairs, hispid beneath. Inflorescences terminal and axillary racemes, 3–5 cm long; rachis 

sparsely hispid; bracts deeply tricuspidate, densely hispid; bracteoles 1–2 mm long, 

palmatilobed to palmatipartite, with 5 stipitate glands. Upper pedicel 2–6 mm long. 

Receptacle tube 0.7–1 cm long, glabrous, gibbous. Sepals 4–5 mm long, glabrous on exterior. 

81

82 


Petals 4–5 mm long. Stamens c. 15, 2–2.5 cm long. Fruit ovoid, 2.7 × 2.6 × 3 cm; exocarp 

knobbled, with short grey pubescence. 


23. Gabon: Rabi, Divangui Rd., F.J.Breteler et al. 9765 (WAG); Rabi, Divangui Rd., F.J.Breteler et al. 9719 (WAG). 

Closely related to D. gilletii, and differing chiefly in its less deeply divided bracteoles and 

smaller flowers with few stamens. 

8. Dactyladenia bellayana (Baill.) Prance & F.White, Brittonia 31: 484 (1979) 

Acioa bellayana Baill., Adansonia 7: 224 (1867); Griffonia bellayana (Baill.) Oliv., Fl. Trop. Afr. 2: 371 (1871). 

T: Gabon: without locality, Griffon du Bellay 602; holo: P; iso: WAG. 


Shrub, climbing. Branches hispid when young. Leaves with stipules linear, 6–8 mm long, 

persistent; petiole 3–4 mm long; lamina oblong-elliptic, 10–18 × 4–7 cm, sparsely hispid 

beneath, with 2 glands at base, rounded to subcordate at base, acuminate at apex; midrib and 

primary veins hispid beneath; primary veins 6–8 (–10) pairs. Racemes axillary and terminal, 

6–12 (–20) cm long; rachis sparsely hispidulous, glabrescent; bracteoles 3–5-palmatipartite, 

with stipitate glands 5 mm long; upper pedicel 3–5 mm long. Receptacle tube 15–22 mm 

long, glabrous, gibbous at base. Sepals 6–9 mm long, glabrous. Petals white, broadly obovate 

with subcordate base, 6–8 mm long. Stamens c. 30, 4–6 cm long. Fruit narrowly ovoid, 

pointed at apex, 25 × 25 × 30–40 mm; exocarp tomentellous, with hispid hairs. 

Cameroon, Gabon and perhaps Zaïre. 23: CMN, GAB. Lowland forests. Map 113. 

23. CAMEROON: Douala-Edéa, km 14, Mpom 328 (SRFCam 2471) (P). GABON: near Libreville, R.P.Klaine 

101 (K, P); Loango, near Bango, H.Soyaux 123 (BR, K, P-photo, W). 

Closely related to D. campestris, D. floribunda and D. icondere. 

9. Dactyladenia hirsuta (A.Chev. ex De Wild.) Prance & F.White, Brittonia 31: 485 (1979) 

Acioa hirsuta A.Chev. ex De Wild., Bull. Jard. Bot. État 7: 200 (1920). T: Ivory Coast: Cavally Basin, 

Grabo, A.Chevalier 19738; holo: P; iso: WAG; photo: BR. 

Shrub? Branches densely velutinous pubescent when young. Leaves with stipules linear, hispid, 

to 8 mm long, caducous; petiole 3–4 mm long, hirsute; lamina elliptic, 5–12 × 2.5–4.5 cm, 

hirsute-pubescent beneath, rounded to subcordate at base, acuminate at apex; midrib villouspubescent 

beneath; primary veins 8–9 pairs. Inflorescence axillary racemes 2–4 cm long; 

rachis hispidulous; bracts entire, oval, 4 mm long; bracteoles 1.5 mm long, with 3–5 

triangular teeth bearing stipitate glands 1.5 mm long; upper pedicels 1–5 mm long. 

Receptacle tube 1.6–2.2 cm long, glabrous, gibbous at base. Sepals 6–10 mm long, glabrous. 

Petals to 15 mm long, caducous. Stamens c. 20, 3.5–4.5 cm long. Fruit unknown. 

Ivory Coast. 22: IVO. Forest. Map 114. 

22. IVORY COAST: Cavally Basin, A.Chevalier 19818 (K, P). 

10. Dactyladenia campestris (Engl.) Prance & F.White, Brittonia 31: 484 (1979) 

Acioa campestris Engl., Bot. Jahrb. Syst. 17: 87 (1893). T: Gabon: 'im savannenwald', H.Soyaux 362; holo: 

B, destroyed; lecto: P, fide G.T.Prance & F.White, Brittonia 31: 484 (1979); isolecto: K, Z; photo: BR. 

Acioa klaineana Pierre ex De Wild., Bull. Jard. Bot. État 7: 196 (1920). T: Gabon: Munda, Libange Farm, 

R.P.Klaine 111; holo: BR; iso: K, P. 

Acioa dawei Mendes, Mem. Junta Invest. Ultramar. Ser. 2. 19: 11 (1960). T: Cabinda: Belize, Mayombe, 

M.T.Dawe 225; holo: K. 

Illustration: R.Letouzey & F.White, Fl. Cameroun 20 (Fl. Gabon 24): 17 (1978) 

Shrub, sprawling, 5 m high. Branches hispid when young, soon glabrescent. Leaves with 

stipules lanceolate, 10–16 × 3–8 mm, hispid, often with circular glands towards base and on 

margins; petiole (3–) 5–6 (–8) mm long, hispid or hispidulous, glabrescent; lamina oblongelliptic 

(14–) 20–32 × (5–) 6–12 cm, glabrous beneath, with 2 large circular glands at base


near midrib and often with other glands especially on acumen, rounded to subcordate at base, 

bluntly acuminate at apex; midrib often with rapidly caducous hispid hairs beneath; primary 

veins (8–) 12–16 pairs. Racemes terminal and axillary, 6–12 (–20) cm long; rachis sparsely 

hispidulous, glabrescent; upper pedicel 3–5 mm long. Receptacle tube 1.5–2.2 cm long, 

glabrous; base gibbous. Sepals 6–9 mm long, glabrous. Petals white, subquadrangular, 

rounded to subcordate at base, 6–10 mm long. Stamens c. 30, 4–6 cm long. Fruit ovoid, 25 × 

30 × 70 mm; exocarp tomentellous, with longitudinal lines. 

SE Nigeria to Gabon and Cabinda. 22: NGA. 23: CAB, CMN, CON, EQG, GAB. Riverine 

forest and secondary forest. Map 115. 

22. NIGERIA: Eket Distr., R.A.Talbot 3123 (BM, K, Z); Calabar Prov., Oban, E.Ujor FHI31793 (K). 

23. CAMEROON: Kribi, P.R.Preuss 263 (BM, K); Douala Edea Reserve, Limbe, D.W.Thomas 306 (K, WAG); 

Bipinde, G.Zenker 2565 (BM, BR, K, P, Z). GABON: Essone, 40 km WSW of Mitzic, G. Le Testu 9363 (BR, K, P, 

WAG); SE of Sindora, bank of R. Ngounié, A.J.M.Leeuwenberg & J.G.M. Persoon 13602 (BR, K, LBV, WAG). 

Closely related to, and frequently confused with, D. bellayana and D. icondere. 

11. Dactyladenia floretii Breteler, Adansonia 22(1): 117 (2000) 

T: Gabon: near Gamba, F.J.Breteler, J.J.Dibata, J.Mayombo & J.M.Moussavou 14490; holo: WAG; iso: G, 

LBV, MO, P. 

Illustration: F.J.Breteler, Adansonia 22(1): 119 (2000). 

Map: F.J.Breteler, Adansonia 22(1): 120 (2000). 

Shrub, lianescent shrub or liana. Leaves with stipules saccate and margins inrolled, shortly 

stipitate, slightly curved or straight, ovoid-ellipsoid in outline, (5–) 8–20 × (2–) 6–8 mm, 

glabrous or with some arachnoid hairs sometimes mixed with hispid hairs; petioles ±terete, 

2–4 (–5) mm long; lamina coriaceous to papyraceous, oblong-elliptic, ovate-elliptic to 

oblong-obovate, (6–) 10–25 (–27) × (2–) 4–7 (–8) cm, glabrous above usually with some 

erect hairs or an arachnoid remnant on impressed midrib, glabrous or hispid beneath, 

(8–) 10–14 (15) pairs of main lateral nerves; narrowly and usually deeply cordate at base, 

acute to shortly and acutely acuminate at apex with acumen to 0.5 (–1.5) cm long; glands 

±flat, present only below. Inflorescences simple ±erect racemes, to 7 cm long, usually 

axillary, sometimes terminal, few- to many-flowered, puberulous; bracts ovate-triangular, 

soon reflexed, 4–5 mm long; bracteoles deeply 3-palmatilobate, 1–2 mm long, puberulous 

with lobes gland-tipped. Receptacle tube (11–) 16–20 mm long, gibbous at base, puberulous. 

Sepals ovate-oblong to elliptic, 5–8 × 2.5–4 mm, with acute to emarginate apex. Petals 

white, early caducous, thin, oblong-elliptic, 10–15 × 5–8 mm, with a short glabrous claw up 

to 1 mm long. Stamens 20–28, 4–4.5 cm long, glabrous; filaments united for c. 2/3 of their 

length; anthers c. 0.6 mm long. Style slightly longer than stamens, glabrous, with a small 

stigma; ovary long-velutinous at apex. Fruit ovoid, narrowly so or not, rounded at both ends, 

3.5–6 × 2–2.5 cm, velutinous and with some longer hispid hairs. 

Gabon. 23: GAB. Rainforest, forest edge next to savanna, coastal thicket and secondary 

vegetation. Map 116. 

23. GABON: Rabi, near camp, F.J.Breteler et al. 9457 (K, WAG); Ogooué-Maritime, Gamba, near airport, 

J.J.Wieringa 1130 (WAG); Ogooué-Maritime, Gamba, F.J.Breteler et al. 14435 (K, WAG). 

12. Dactyladenia jongkindii Breteler, Adansonia 22(1): 118 (2000) 

T: Gabon: 5–15 km NNW of Ndjolé, F.J.Breteler & C.C.H.Jongkind 10412; holo: WAG; iso: BR, G, K, LBV, 

MO, P. 

Illustration: F.J.Breteler, Adansonia 22(1): 121 (2000). 

Map: F.J.Breteler, Adansonia 22(1): 120 (2000). 

Shrub to treelet, 3–6 m tall. Leaves with stipules saccate and margins inrolled, ±ovoid in 

outline, (6–) 10–15 (–18) mm long, 4–5 mm in diam., rather sharply narrowed at base, 

gradually tapering at apex, with 1–4 long-stalked glands from petiole on distal side; petioles 

subterete, 3–5 mm long; lamina oblong to obovate-oblong, 14–25 (–30) × 5–10 cm, glabrous 

above except for slightly impressed hairy midrib, sparsely short to long-hairy beneath; 

83

84 


(10–) 12–15 pairs of main laterals; narrowly cordate at base, acutely acuminate at apex with 

acumen 0.5–1.5 cm long; glands flat to slightly impressed, small, beneath only. Inflorescences 

suberect simple racemes 10–15 cm long, many-flowered, puberulous. Bracteoles subopposite 

to alternate, inserted 2–4 mm below apex of lower pedicel, deeply 5-palmatilobate, 4–6 mm 

long; lobes gland-tipped, puberulous in basal part. Receptacle tube 15–17 mm long, gibbous 

at base, puberulous outside, glabrous inside except for some stiff hairs at mouth. Sepals 

spreading to reflexed, ovate-elliptic to oblong-obovate, 5–8 × 3–4 mm, tomentose inside and 

on those outside parts covered in bud, puberulous with some hispid hairs outside. Petals 

early caducous, thin, white, oblong-elliptic to obovate, 11–15 × 6–7 mm, glabrous, shortly 

clawed. Stamens 30–33, 5–6 cm long, glabrous, white; filaments united for c. 3/4 of their 

length; anthers ellipsoid, c. 0.6 mm long. Style slightly longer than stamens, glabrous; stigma 

small; ovary velutinous, c. 1 mm long. Fruit unknown. 


23. Gabon: near Libreville, F.J.Breteler & R.H.M.J.Lemmens 8410 (WAG). 

13. Dactyladenia icondere (Baill.) Prance & F.White, Brittonia 31: 485 (1979) 

Acioa icondere Baill., Adansonia 7: 223 (1867); Griffonia icondere (Baill.) Oliv., Fl. Trop. Afr. 2: 371 (1871). 

T: Gabon: without precise locality, Griffon du Bellay 313; lecto: P, fide G.T.Prance & F.White, Brittonia 31: 

485 (1979); isolecto: K. 

Illustrations: A.Chevalier, Veg. Ut. Afr. Trop. Franç. 9, t. 18 (1917); R.Letouzey & F.White, Fl. Cameroun 20 

(Fl. Gabon 24): 35 (1978). 

Shrub, climbing, 5–8 m tall. Branches densely-velutinous-tomentose when young. Leaves 

with stipules broadly lanceolate, 12 × 5 mm, velutinous-tomentose, sometimes with circular 

glands towards base; petioles 5–8 mm long, velutinous-tomentose; lamina broadly elliptic, 

15–25 × 8–13 cm, sometimes oblong to slightly obovate, sparsely lanate-pubescent beneath, 

with large circular glands near to midrib and at base, cordate to subcordate at base, 

acuminate at apex; midrib velutinous-tomentose beneath, glabrescent; primary veins 10–16 

pairs, lanate-pubescent beneath. Racemes terminal and axillary, 12–24 (–30) cm long; rachis 

sparsely pubescent; bracteoles 2–5 mm long, palmatipartite to palmatisect with 5 ±stipitate 

glands. Upper pedicel 2–6 mm long. Receptacle tube 1.6–2.2 cm long, gibbous at base, 

glabrous on exterior. Sepals 6–8 mm long, glabrous on exterior, sometimes with a single 

basal gland. Petals white, obovate to quadrangular, cordate at base, c. 8 mm long. Stamens 

30–40, 4–4.5 cm long. Fruit amygdaliform, 25 × 30 × 50 mm; exocarp dark yellow 

tomentose and with hispid hairs, with longitudinal ridges when dry. 

Cameroon to Cabinda and Zaire (Mayumbe). 23: CAB, CMN, CON, EQG, GAB, ZAI. 

Mainly in coastal thicket. Map 118. 

23. CAMEROON: near Grand Batanga, J.J.Bos 6064 (BR, K, WAG); Grand Batanga, 12 km S of Kribi, 

J.J.Bos 5279 (BR, K, WAG); km 45 Campo-Kribi, W.J.J.O. de Wilde 7812 (BR, P, WAG); Bipinde, G.Zenker 

1949 (BM, K, P, WAG, WU, Z). GABON: Libreville, A.Aubréville 99 (P); Cap Esterias, near Libreville, 

N.Hallé & J.-F.Villiers 5510 (P). 

Closely related to D. floribunda. 

14. Dactyladenia floribunda Welw., Apont. 1: 572 (1859) [1858] 

Acioa icondere var. welwitschii De Wild., Bull. Jard. Bot. État 7: 195 (1920); Acioa floribunda (Welw.) Exell, 

J. Bot. 66 (Suppl. Polypet.) 161 (1928). T: Angola: Serra do Alto Queta, F.M.J.Welwitsch 1289; lecto: 

LISU, fide Mendes, Consp. Fl. Angola 4: 20 (1970); isolecto: BM, COI, K, P; photo: BR. 


Shrub or tree up to 20 m high. Branches densely persistently velutinous-pubescent when 

young. Leaves with stipules linear-lanceolate, 5–7 mm long, with dense appressed pubescence; 

petioles 5–8 mm long, velutinous-tomentose; lamina elliptic to oblong, 11–15 (–18) × 

(4.5–) 6–9 cm, tomentose beneath, with 2 inconspicuous circular glands towards base, 

cuneate to rounded at base, acuminate at apex with acumen 3–5 mm long; midrib 

tomentellous beneath; primary veins 7–10 pairs, with sparse yellow pubescence beneath. 

Racemes terminal or axillary, 2–6 cm long, congested; rachis sparsely pubescent; bracteoles


c. 2 mm long, subterminal, 5-palmatilobed to palmatipartite, with elongate triangular 

glandular lobes, appressed pubescent; upper pedicel 3 mm long. Receptacle tube 1.2–1.8 cm 

long, distinctly gibbous at base, sometimes sparsely pubescent. Sepals 6–8 mm long, sparsely 

pubescent on exterior. Petals obovate, obtuse to subcordate at base, white, 15 × 6 mm. 

Stamens (15–) 20–25 (–30), 3–6 cm long. Fruit narrowly ellipsoid, 20–25 × 40 mm, 

terminating in 10 mm point when dry; exocarp dark yellow tomentellous. 

Cameroon to Angola. 23: CAB, CMN, CON, EQG, GAB, ZAI. 26: ANG. Forest on slopes to 

800 m. Map 119. 

23. CABINDA: Buco Zau, between Buco Xibaba and Chiongo, R.Monteiro et al. 253 (BM). CAMEROON: 

Ebolowa-Ambam, km 18, W.J.J.O. de Wilde 8174 (BR, P, WAG); Colline Zingui, 20 km WSW Ebolowa, 

R.Letouzey 10285 (BR, K, P). GABON: Abam, 25 km WNW of Oyem, G. Le Testu 9130 (BR, K, P, WAG). 

ZAIRE: Leopoldville Prov., Pandji near Tshela, A.Flamigni 10451 (BM, BR, K, P). 26. ANGOLA: Cuanza 

Norte, Cazengo, Monte Belo, J.Gossweiler 4405 (BR, K, LISC); Cuanza Norte, Cazengo, Monte Belo, 

J.Gossweiler 4383 (K). 

Closely related to D. icondere, which, however, is a species of coastal thicket. 

15. Dactyladenia barteri (Hook.f. ex Oliv.) Prance & F.White, Brittonia 31: 484 (1979) 

Griffonia barteri Hook.f. ex Oliv., Fl. Trop. Afr. 2: 373 (1871); Acioa barteri (Hook.f. ex Oliv.) Engl., Bot. 

Jahrb. Syst. 26: 382 (1899). T: Nigeria: Lagos, C.Barter 2183; lecto: K, fide G.T.Prance & F.White, 

Brittonia 31: 484 (1979); isolecto: P, W. 

Acioa tenuiflora Dinkl. & Engl., Bot. Jahrb. Syst. 26: 382 (1899). T: Liberia, A.Dinklage 1675; holo: B, 

destroyed; lecto: K, designated here; isolecto: CAS, P, Z. 

Acioa lanceolata Engl., Bot. Jahrb. Syst. 26: 382 (1899). T: Cameroon: Grand Batanga, near Kribi, 

A.Dinklage 925; lecto: HBG, designated here. 

Acioa trillesiana Pierre ex De Wild., Bull. Jard. Bot. État. 7: 206 (1920), nom inval. in synon. 

Illustrations: R. Letouzey & F.White, Fl. Cameroun 20 (Fl. Gabon 24): 11 (1978); G.T.Prance & F.White, 

Philos. Trans. B320: 134 (1988). 

Tree up to 12 m tall or climbing shrub. Branches hispidulose when young, rapidly glabrescent, 

lenticellate. Leaves with stipules usually attached to base of petiole, linear, 4–6 mm long; petiole 

3–4 mm long, hispid, glabrescent; lamina oblong-elliptic, 3.7–13 (–15) × 3.5–5 (–7) cm, 

glabrous, or sometimes scabrous beneath except on midrib, with various glands near to 

midrib, subcuneate at base, acuminate at apex; midrib sparsely hispid beneath, glabrescent; 

primary veins 4–6 pairs. Racemes terminal or subterminal, 3–4 (–12) cm long, puberulous; 

bracteoles 0.1–0.15 cm long, lanceolate, sometimes with stipitate glands; upper pedicel 

(0.5–) 0.8–1.2 (–1.5) cm long. Receptacle tube 4–6 mm long, ±gibbous at base, puberulous 

on exterior. Sepals 4–5 mm long, puberulous, often with stipitate glands on margin. 

Petals white, oblong-obovate with acute base or obtuse, 4–5 mm long. Stamens 15–20, 

(1.5–) 2.5 (–3) cm long. Fruit narrowly ovoid, 25 × 35 × 50 mm, acute at summit; exocarp 

glabrescent. Fig. 64. 

Widespread from Sierra Leone to E Zaire and Gabon. 22: BEN, GHA, IVO, LBR, NGA, SIE, 

TOG. 23: CMN, GAB, ZAI. Lowland rainforest. Map 120. 

22. GHANA: Benso, Colong Prov., J.E.Andoh FH5388 (BR, K, P). IVORY COAST: 17 km W of Abidjan, 

W.J.J.O. de Wilde 898 (BR, K, P, WAG). LIBERIA: Gbarnga, 3 mi NE of Suacoco, Z.D.Traub 264 (BM, BR, 

K, MO); E Province, Tappita, J.T.Baldwin 9108 (K, US). NIGERIA: Milliken Hill, Enugu, F.N.Hepper 2231 

(K, P); Onitsha Prov., Onitsha, Akpaka Forest Reserve, J.K.Adebusuyi FHI43903 (K, WAG). SIERRA 

LEONE: Makeni, F.C.Deighton 4259 (K, P). 23. CAMEROON: near Nyong, 35 km WSW of Eséka, 

W.J.J.O. de Wilde 2698 (BR, K, P, WAG, Z). GABON: R. Mondah, nr. Libreville, R.P.Klaine 3448 (P). 

ZAIRE: Kivu, A.Léonard 5162 (BM, BR, WAG). 

Closely related to D. scabrifolia. 

16. Dactyladenia sapinii (De Wild.) Prance & F.White, Brittonia 31: 486 (1979) 

Acioa sapinii De Wild., Bull. Jard. Bot. État. 4: 80 (1914). T: Zaire: Kasai, Bienge, A.Sapin s.n.; holo: BR; 

photo: P. 

Tree. Branches densely brown velutinous-pubescent when young. Leaves with stipules 

subulate-falcate, to 10 mm long, tomentellous, caducous; petiole to 10 mm long, densely 

85

86 


Figure 64. Dactyladenia barteri. A, habit; B, ½ flower; C, petals; D, anthers; E, fruit (A–D, 

B.O.Daramola FHI 45684; E, W.MacGregor 373). Scale bars: A, E = 3 cm; B, C = 5 mm; 

D = 2 mm. Drawn by Janet Dyer. Previously published in G.T.Prance & F.White, Philos. 

Trans. B320: 134 (1988).


brown velutinous-pubescent; lamina elliptic to ovate-elliptic, 10–17 × 4.5–11 cm, densely 

tomentose beneath, cordate at base, abruptly acuminate at apex; midrib densely tomentose 

beneath; primary veins c. 12 pairs. Inflorescences terminal or axillary racemes, densely 

tomentose, to 10 cm long; bracts oval-acute, c. 6 mm long, tomentose, eglandular; bracteoles 

6 × 4 mm, subreniform, opposite, eglandular; upper pedicel 10–14 mm long. Receptacle tube 

1.1–1.5 cm long, densely tomentose on exterior. Sepals broadly ovate, densely tomentose. 

Petals oblong, early caducous. Stamens c. 48, c. 5 cm long. Fruit unknown. 

Known only from the type collection. Zaire. 23: ZAI. Riparian forest. Map 121. 

Closely related to D. pallescens, and differing chiefly in its hairy leaf lower surface, longer 

pedicels, smaller flowers and subreniform bracteoles. 

17. Dactyladenia pallescens (Baill.) Prance & F.White, Brittonia 31: 486 (1979) 

Acioa pallescens Baill., Adansonia 7: 224 (1867); Griffonia pallescens (Baill.) Oliv., Fl. Trop. Afr. 2: 372 (1871). 

T: Gabon: without precise locality, Griffon du Bellay 261; holo: P. 

Griffonia pallescens var. scandens Oliv., Fl. Trop. Afr. 2: 372 (1871). T: Equatorial [Upper] Guinea: River 

Muni syn: G.Mann 1762; lecto: K. 

Griffonia pallescens var. arborescens Oliv., Fl. Trop. Afr. 2: 372 (1871). T: Equatorial [Upper] Guinea: Old 

Calabar; syn: G.Mann 2282; lecto: K. 

Acioa thollonii De Wild., Bull. Jard. Bot. État 7: 208 (1920). T: Gabon: R. Ogooué, Booué, F.-R.Thollon 

794; holo: BR; iso: P. 

Illustrations: R.W.J.Keay, Fl. Trop. W. Afr., ed. 2, 1: 432 (1958); R.Letouzey & F.White, Fl. Cameroun 20 

(Fl. Gabon 24): 53 (1978). 

Shrub 5–8 m tall, sometimes multiple-stemmed. Branches pubescent when young, soon 

glabrescent, lenticellate. Leaves with stipules linear, c. 8 mm long, pubescent, sometimes 

with a basal circular gland; petiole 3–8 mm long, densely pubescent, rarely glabrescent; 

lamina elliptic to oblong-elliptic or oblong, 8–25 × 4–11 cm, glabrous beneath, slightly 

rugose, with some large circular basal glands and smaller glands near acumen, cordate at 

base, abruptly acuminate at apex; midrib pubescent, soon glabrescent beneath; primary veins 

6–12 pairs. Inflorescence terminal and axillary racemes, 8–15 (–40) cm long; rachis densely 

yellow-grey tomentose; bracts 6 mm long, tricuspidate, often with 2 circular glands on 

exterior; bracteoles 2–3 mm long, ovate, denticulate-glandular or tricuspidate. Upper pedicel 

2–4 mm long. Receptacle tube 1.8–3.6 cm long. Sepals 8–10 mm long, often with a lateral 

basal gland. Petals white, broadly obovate, subcordate at base, 6–9 mm long. Stamens 60–75, 

4–6 cm long. Fruit borne on a robust peduncle, subovoid, 25 × 30 × 40 mm; exocarp 

pubescent with short tomentum and hispid hairs. Fig. 65. 

Nigeria to Cabinda and Zaire. 22: NGA. 23: CAB, CMN, CON, EQG, GAB, GGI, ZAI. 

Riparian and secondary forest. Map 122. 

22. NIGERIA: Itu, Cross River, J.H.Holland 35 (K); Calabar Prov., Akamkpa Rubber Estate, Dukwe, Calabar R., 

M.G.Latilo FHI41310 (K); Oban, P.A.Talbot 1599 (BM, K, Z). 23. CAMEROON: 15 km N of Kribi, 

D.Thomas 3443 (BR, K, MO); Lobe R., above Grand Batanga, J.J.Bos 3073 (BR, K, WAG). EQUATORIAL 

GUINEA: R. Muni, G.Mann 1762 (K, P). GABON: km 23, Moanda-Franceville, F.J.Breteler 7001 (BR, K, 

WAG); Lastoursville, G. Le Testu 7532 (BR, K, P, WAG). GULF OF GUINEA IS.: Bioko, km 31 Malabo- 

Riaba, Carvalho 2872 (BR, K, WAG); km 21-22, Carvalho 3798 (BR, K, WAG). 

Closely related to D. buchneri and D. sapinii. 

18. Dactyladenia buchneri (De Wild.) Prance & Sothers, Kew Bull. 57: 995 (2002) 

Acioa buchneri Engl., Bot. Jahrb. Syst. 17: 88 (1893). T: Angola: Lunda, Chitato, M.Buchner 10; holo: B, 

destroyed. 

Acioa lujae De Wild., Bull. Jard. Bot. État 7: 209 (1920); Dactyladenia lujae (De Wild.) Prance & F.White, 

Brittonia 31: 486 (1979). T: Zaire: Sankuru, fl., E.P.Luja 38; lecto: BR, fide G.T.Prance & F.White, 

Brittonia 31: 486 (1979); photo: K, P. 

Acioa gossweileri Cavaco, Bull. Mus. Natl. Hist. Nat., sér. 2, 26: 638 (1954). T: Angola: Chitato, Dundo, 

J.Gossweiler 14039; holo: P; iso: BM, COI, K, LUA. 

87

88 


Figure 65. Dactyladenia pallescens. A, habit; B, flower; C, petals; D, fruit (A–C, 

G. Le Testu 7508; D, R.Letouzey 13689). Scale bars: A, D = 3 cm; B, C = 1 cm. Drawn by 

Margaret Tebbs.


Shrub or small tree 6–8 m high. Branches shortly yellow-brown pubescent when young. 

Leaves with stipules linear, 5–6 mm long, hirsutulous; petioles 3–5 mm long, sparsely 

tomentellous, rugose; lamina elliptic-lanceolate, 10–13 × 4–5.5 cm, glabrous beneath except 

on midrib, rounded at base, shortly acuminate at apex; midrib pilose beneath; primary veins 

7–9 pairs. Inflorescence terminal and axillary racemes, 5–10 cm long, pale grey-tomentose. 

Bracteoles 2–3 mm long, entire or denticulate-glandular, sometimes tricuspidate. Upper 

pedicels 1–3 mm long. Receptacle tube 0.8–1.5 cm long, pale-grey tomentose on exterior. 

Sepals 5–8 mm long, grey tomentose. Petals white, 8–10 mm long, caducous. Stamens 25–35, 

2–3 cm long. Fruit ovoid, 5 × 3.5 × 2.5 cm; exocarp brown-pubescent. 

Central African Republic, Congo, Zaire and Angola. 23: CAF, CMN, CON, ZAI. 26: ANG. 

Riverine forests. Map 123. 

23. CAMEROON: Prov. Southwest, Takamanda Forest Reserve, near the Makone River, C.Doumenge 434 (K). 

ZAIRE: Kasai Distr., S of Pania Mutombo, Pangula, J.Bequaert 27 (BR, K, US); Babade, F.Matagne 250 (BR, K); 

Kwango, source of Jonquebi R., trib. of Mamba, R.Germain 2447 (BR, K); Lovanium, J.F.Carrington 17 (K); 

Kasai Prov., Kazumba, R. Miao, L.Liben 3409 (BR, K). 

Very closely related to D. pallescens and perhaps only subspecifically distinct. 

19. Dactyladenia staudtii (Engl.) Prance & F.White, Brittonia 31: 486 (1979) 

Acioa staudtii Engl., Bot. Jahrb. Syst. 26: 379 (1899). T: Cameroon: Lolodorf, A.Staudt 263; holo: B, 

destroyed; lecto: P, fide G.T.Prance & F.White, Brittonia 31: 486 (1979); isolecto: K, Z; photo: BR. 

Acioa talbotii Baker f., Cat. Pl. Oban : 29 (1913). T: Nigeria: Oban, P.A.Talbot 1533; holo: K; iso: BM, P, 

Z; sketch and fragment, BR. 


Shrub or small tree to 15 m tall. Branches densely grey to yellow villous when young, slowly 

glabrescent. Leaves with stipules narrowly triangular, 5–6 mm long, slightly falciform; 

petioles 2–5 mm long, villous; lamina elliptic 8–14 × 4–7 cm, slightly scabrous beneath, with 

circular glands towards base and under acumen beneath, cuneate to subcuneate at base, 

acuminate at apex with acumen 5–10 mm long; midrib sparsely villous, glabrescent beneath; 

primary veins 8–12 pairs, sparsely villous, glabrescent beneath. Inflorescence terminal and 

axillary racemes, 3–4.5 cm long, few-flowered, rachis and pedicels with yellowish villous 

hairs. Bracts lanceolate, 4 mm long, with margins denticulate; bracteoles 1–1.5 mm long, 

deltate-acuminate, with 2–4 stipitate glands. Upper pedicel 2–4 mm long. Receptacle tube 

1.2–2 cm long, villous on exterior. Sepals 6–8 mm long, villous on exterior. Petals white, 

obovate, 5–6 mm long. Stamens 15–20, 4–5 cm long. Fruit ellipsoid, 4.5 × 2.5 cm; exocarp 

fulvous-tomentose. 

SE Nigeria to Gabon. 22: NGA. 23: CMN, GAB. Primary forest and banks of river, to 200 m. 

Map 124. 

22. NIGERIA: Cross River, 5 km E of Obudu Cattle Ranch, G.E.Pilz 1973 (K, WAG). 23. CAMEROON: 

SW Prov., Korup N. P., S end, C.Doumenge 367 (K, MO). GABON: Ngongolane in Cristal Mts, N.Hallé 842 

(P); Tsango, 60 km towards SE of Sindara, G. Le Testu 6043 (BR, P); Moucoundangoy, 60 km ENE of 

Mbigou, G. Le Testu 8425 (BM, BR, K, P, WAG); Mbé R., Kinglé Power Station, F.J.Breteler & J.J.F.E. de 

Wilde 339 (BR, K, WAG). 

20. Dactyladenia dewevrei (De Wild. & T.Durand) Prance & F.White, Brittonia 31: 484 

(1979) 

Acioa dewevrei De Wild. & T.Durand, Ann. Mus. Congo, Bot., sér. 2, 1(2): 19 (1900). T: Zaire: Lukolela, 

A.Dewèvre 743; holo: BR. 

Acioa seretii De Wild., Ann. Mus. Congo, Bot., Sér. 5, 2: 254 (1908); Acioa dewevrei var. seretii (De Wild.) 

Hauman, Fl. Congo Belg. 3: 50 (1952). T: Zaire: Gombari, F.Seret 807; holo: BR; photo: K. 

Acioa vanhouttei De Wild., Ann. Mus. Congo, Bot., Sér. 5, 2: 255 (1908); Acioa dewevrei var. vanhouttei (De 

Wild.) Hauman, Fl. Congo Belg. 3: 50 (1952). T: Zaire: Kibula, J.Gillet 3964; holo: BR; photo: K, P. 

Acioa reygaertii De Wild., Bull. Jard. Bot. État 7: 201 (1920); Acioa dewevrei var. reygaertii (De Wild.) 

Hauman, Fl. Congo Belg. 3: 49 (1952). T: Zaire: Mobwasa, F.J.Reygaert 12029; holo: BR; photo: K, P. 

89

90 


Acioa brazzae De Wild., Bull. Jard. Bot. État 7: 202 (1920). T: Congo: Brazzaville, M.Thollon 565; lecto: 

BR, designated here; isolecto: P; photo: BR. 


Shrub or small tree, sometimes scandent. Branches yellowish villous-pubescent when young. 

Leaves with stipules linear, 3 mm long; petiole 3–4 mm, hispidulous, glabrescent; lamina 

elliptic, oblong or rarely lanceolate, (8–) 10–13 × (4–) 5–7 cm, with sparse hairs beneath, 

glabrescent, with circular glands scattered over lower surface especially near margin, with 2 

large glands at rounded base, acuminate at apex with acumen c. 5 mm long; midrib 

tomentellous, glabrescent beneath; primary veins 6 pairs, with sparse hairs beneath. Racemes 

terminal and subterminal, 10–15 cm long, with numerous flowers crowded near apex; rachis 

sparsely puberulous; bracteoles oval, palmatisect, c. 2 mm long, with c. 5–10 stipitate glands 

0.5–1.5 mm long; upper pedicel 2–4 mm long. Receptacle tube 0.7–1.1 cm long, narrowly 

obconic, sparsely puberulous on exterior. Sepals 4–5 mm long, sparsely puberulous. Petals 

white, c. 5 mm long. Stamens 30–35, 3–3.5 (–5) cm long. Fruit ellipsoid, 18 × 25 × 50 cm, 

with short 2 mm stipe; exocarp with short yellowish velutinous pubescence and hispid hairs. 

Nigeria to Zaire. 22: NGA. 23: CAF, CMN, CON, ZAI. Riverine habitats to 750 m. Map 125. 

22. NIGERIA: Jameson R., Sapoba Reserve, A.F.Ross 210 (K); S Nigeria, J.D. Kennedy 2058 (BR, FHO, K, P). 

23. CAMEROON: near Banga, 65 km SE of Batouri, R.Letouzey 4917 (BR, K, P, WAG). 

21. Dactyladenia laevis (Pierre ex De Wild.) Prance & F.White, Brittonia 31: 485 (1979) 

Acioa laevis Pierre ex De Wild., Bull. Jard. Bot. État. 7: 205 (1920). T: Gabon: Libreville, R.P.Klaine 

1890; holo: P; photo: BR. 


Shrub or small tree to 20 m tall. Branches with whitish arachnoid caducous indumentum 

when young. Leaves with stipules attached to base of petiole, lanceolate, 4–5 mm long; 

petioles 3–6 mm long, tomentellous, soon glabrous, transversely rugose; lamina elliptic to 

slightly oblong, 6–13 × 3–5.5 cm, glabrous beneath, with 2 circular glands towards base and 

others scattered and below the acumen, cuneate at base, often decurrent onto petiole, 

acuminate at apex with acumen curved; midrib glabrous beneath; primary veins 5–7 pairs, 

glabrous. Inflorescences terminal and axillary cymes, 4–8 cm long with rachis glabrous; 

bracts oval, 3–4 mm long, persistent, reflexed onto inflorescence axis; bracteoles 0.5–1.5 mm 

long, subreniform, with c. 5 marginal denticulations, very sparsely villous. Upper pedicel 

4–8 mm long. Receptacle tube (1.2–) 1.8–2.6 cm long, glabrous on exterior. Sepals 6–9 mm 

long. Petals obovate, obtuse at base, 6–7 mm long. Stamens 20–25, 3–4 cm long. Drupe 

subglobose, 18 × 20 × 25 mm; exocarp yellow-brown tomentose, densely colliculate. 

Gabon, known only from the vicinity of Libreville. 23: GAB. Map 126. 

23. GABON: Libreville, R.P.Klaine 2981 (P); Libreville, R.P.Klaine 3538 (BR fragments, P). 

This species differs from all others in its subreniform bracteoles. 

22. Dactyladenia lehmbachii (Engl.) Prance & F.White, Brittonia 31: 485 (1979) 

Acioa lehmbachii Engl., Bot. Jahrb. Syst. 26: 379 (1899). T: Cameroon: Buea, Lehmbach 115; holo: B, not 

destroyed. 

Acioa rudatisii Engl. ex De Wild., Bull. Jard. Bot. État 7: 215 (1920). T: Cameroon: Abonando, 

A.G.H.Rudatis 30; lecto: K, designated here; isolecto: BM; photo: BR, P. 


Tree to 20 m tall. Branches with golden-brown pubescence when young. Leaves with stipules 

linear, 5–8 mm long; petioles 3–6 mm long; lamina oblong-elliptic, 7–17 × 3–7 cm, glabrous 

and slightly scabrous beneath, with 2 glands at base and under acumen, rounded to cuneate at 

base, acuminate at apex; midrib slightly pubescent, glabrescent beneath; primary veins 5–9 

pairs, sparsely hispid, glabrescent. Inflorescences racemes or cymes, terminal and axillary, 

3–5 cm long; bracts lanceolate, 3–5 mm long; bracteoles 1–1.5 mm long, ovate-deltate, 

palmatifid to palmatipartite, with 5–7 substipitate to stipitate glands each 1–2.5 mm long. 

Upper pedicel 3–6 mm long. Receptacle tube 1.2–1.8 cm long, glabrous on exterior, gibbous


at base. Sepals 6–8 mm long, glabrous. Petals oblong-subrectangular, acute at base, 6–10 mm 

long. Stamens 15–20, 2.5–3 (–6) cm long. Fruit ovoid, 2–4 × 2.5–4.5 × 3.5–6 cm; exocarp 

tomentose and with hispid hairs. 

SE Nigeria and Cameroon. 22: NGA. 23: CMN. Forests. Map 127. 

22. NIGERIA: Calabar Prov., Itu, Eki, O.Akpata FHI3939 (K). 23. CAMEROON: Buea, J.B.Maitland 545 

(K, P); Kumba, J.Olorunfemi FHI30628 (K, P); 50 km ENE of Sangmélima, G.L.Bates 855 (BM); 40 km W of 

Mamfe, R.Letouzey 13563 (BR, K, P, WAG); between Tinta and Atolo-Boka, 15 km S of Akwaya, R.Letouzey 

14101 (BR, K, P); near Mamfe, M.G.Latilo & Oguntayo FHI67773 (K, WAG). 

Close to and easily confused with D. gilletii. 

23. Dactyladenia mannii (Oliv.) Prance & F.White, Brittonia 31: 486 (1979) 

Griffonia mannii Oliv., Fl. Trop. Afr. 2: 382 (1871); Acioa mannii (Oliv.) Engl., Bot. Jahrb. Syst. 26: 381 

(1899). T: Bioko [Fernando Po], G.Mann 1427; holo: K; iso: P; photo: BR. 


Shrub, climbing, 4–6 m tall. Branches arachnoid-tomentose when young, soon glabrescent, 

lenticellate. Leaves with stipules 6 × 3–4 mm with finely denticulate margins; petioles 8 mm 

long, tomentellous, glabrescent; lamina elliptic to oblong, 12–20 × 5–9 cm, glabrous beneath, 

with 2 large circular glands at base and small glands near base and apex, rounded to 

subcuneate at base, acuminate at apex with acumen c. 1 cm long; midrib glabrous beneath; 

primary veins 6–7 (–10) pairs. Inflorescences terminal and axillary cymes, 8–12 cm long; 

rachis and branches puberulent; bracts 2–3 mm long, slightly denticulate at margins; 

bracteoles 1–3 mm long, ovate, sometimes denticulate. Upper pedicel 2–8 mm long. 

Receptacle tube 1.4–2 cm long, gibbous. Sepals 6–7 mm long, sometimes with a lateral basal 

gland. Petals 4 mm long, broadly obovate. Stamens c. 30, 3–4 (–5) cm long. Fruit sub-ovoid, 

3 × 1.8 cm; exocarp tomentose. 

Known only from Bioko and Cameroon (vicinity of Mount Cameroon). 23: CMN, GGI. 

Secondary forest. Map 128. 

23. CAMEROON: near Buea, J.B.Maitland 467 (K, P); near Kumba, A.Staudt 513 (K, P, Z); Bay Manya, 15 km 

SW of Kumba, Babute 9 (FHO). 

24. Dactyladenia pierrei (De Wild.) Prance & F.White, Brittonia 31: 486 (1979) 

Acioa pierrei De Wild., Bull. Jard. Bot. État. 7: 212 (1920). T: Gabon: near Libreville, R.P.Klaine 196 in 

L.Pierre 6395; lecto: P, fide G.T.Prance & F.White, Brittonia 31: 486 (1979); isolecto: E, K, P; photo: BR. 


Shrub to 8 m tall. Branches arachnoid pubescent when young, sparsely lenticellate. Leaves 

with stipules oval, 8 × 4 mm, acute at apex; petiole 3–5 mm long, densely tomentose; lamina 

elliptic to obovate, 9–18 × 4.5–8 cm, glabrous beneath with 6–8 large circular glands 

towards base near midrib and small glands beneath acumen, rounded at base, acuminate at 

apex with acumen c. 5 mm long; midrib tomentellous, glabrescent beneath; primary veins 

7–10 pairs. Inflorescence dichotomous cymes, terminal and axillary, 5–20 cm long, somewhat 

corymbose; bracts tricuspidate, c. 5 mm long; bracteoles 2 mm long, ovate, acute, sometimes 

with lateral glands; upper pedicel 1–1.5 cm long. Receptacle tube 1.2–1.8 cm long, densely 

tomentellous. Sepals 6–8 mm long, with lateral glands. Petals obovate, 6–8 mm long, white 

to cream. Stamens 20–30, 2–4 cm long. Fruit amygdaliform, 30 × 40 × 65 (–75) mm; exocarp 

sparsely yellow-brown tomentellous. 

Known only from Gabon. 23: GAB. Habitat not known. Map 129. 

23. GABON: near Libreville, R.P.Klaine 510 (BM, P, US), 2033 (BR, P); Nyanga, G. Le Testu 1779 (BM, BR, 

K, P, Z), 2114 (BM, BR, K, P). 

25. Dactyladenia eketensis (De Wild.) Prance & F.White, Brittonia 31: 485 (1979) 

Acioa eketensis De Wild., Bull. Jard. Bot. État 7: 214 (1920). T: Nigeria: Eket Distr., P.A.Talbot s.n.; holo: 

BM; iso: BR. 


91

92 


Shrub or tree to 20 m tall. Branches sparsely puberulous when young, glabrescent, 

lenticellate. Leaves with stipules linear, 6 mm long, appressed-pubescent; petioles 3–4 mm 

long, glabrous; lamina oblong to elliptic, 10–14.5 (–17) × 4.5–5 (–8.5) cm, glabrous beneath 

except for midrib, with 2 large circular glands at base and 1 or 2 pairs higher up and 

alongside midrib and at acumen; rounded to subcordate at base; acuminate at apex with 

acumen c. 1 cm long, slightly curved towards base; midrib with a few scattered hairs only 

beneath; primary veins 7–8 pairs. Inflorescence terminal and axillary cymes, to 10 cm long; 

rachis puberulous; bracteoles 2 mm long, ovate, tomentellous on both surfaces; upper pedicel 

5–8 mm long. Receptacle tube 0.6–0.7 cm long, narrowly obconic, tomentellous on exterior. 

Sepals 5–6 mm long, tomentose, sometimes with lateral glands. Petals oblong, subcordate at 

base, 5 mm long. Stamens c. 20, 4–4.5 cm long. Fruit unknown. 

S Nigeria and Gabon; known from 3 collections. 22: NGA. 23: GAB. Map 130. 

22. NIGERIA: Eket District, P.A.Talbot 3239 (BM). 23. GABON: Haut Ramboué, A.Chevalier 27127 (P). 

26. Dactyladenia dichotoma (De Wild.) Prance & F.White, Brittonia 31: 485 (1979) 

Acioa dichotoma De Wild., Bull. Jard. Bot. État 7: 216 (1920). T: Nigeria: Eket Distr., P.A.Talbot 3048; 

holo: BM; iso: BR; photo: BR, K, P. 

Tree? Branches shortly tomentose when young, glabrescent. Leaves with stipules linear, c. 4 mm 

long, caducous; petioles 3–5 mm long, glabrous; lamina elliptic, oval-elliptic or obovateelliptic, 

7–16 × 3.5–9 cm, glabrous beneath, subcordate at base, acuminate at apex; midrib 

glabrous beneath; primary veins 7–9 pairs. Inflorescences terminal and axillary racemes, 

7–12 cm long, entirely covered with greyish tomentum; bracteoles 2–3 mm long, terminal, 

ovate; upper pedicels 2–4 mm long. Receptacle tube 0.7–1.1 cm long, narrowly obconic, 

grey-tomentose on exterior. Sepals 4–5 mm long. Petals caducous. Stamens 15–20, 2–3 cm 

long. Fruit unknown. 

Southern Nigeria; known only from the type collection. 22: NGA. Along rivers. Map 131. 

27. Dactyladenia dinklagei (Engl.) Prance & F.White, Brittonia 31: 485 (1979) 

Acioa dinklagei Engl., Bot. Jahrb. Syst. 26: 381 (1899). T: Liberia: Grand Bassa, M.Dinklage 1748; holo: 

B, not destroyed; iso: BM, E, WU. 

Tree, spreading, to 9 m high, with drooping branches. Branches tomentose when young. 

Leaves with stipules lanceolate, 5 × 2 mm; petioles densely ferruginous-pilose; lamina 

oblong, 8–15 × 3–6.5 cm, glabrous and shiny beneath except for pilose venation, rounded at 

base, acuminate at apex; midrib ferrugineous-pilose, glabrescent beneath; primary veins c. 8 

pairs, pilose beneath. Inflorescences terminal and axillary racemes, 5–10 cm long, densely 

golden-tomentose; bracts 5–6 mm long; bracteoles 2 mm long, terminal, ovate; upper pedicel 

2–3 mm long. Receptacle tube 1.8–2.8 cm long. Sepals 6–8 mm long, with lateral glands 

linear-oblong. Petals white, exceeding sepals. Stamens 30–35, 3–3.5 cm long. Fruit ovoid, 

4.5 × 4 × 2 cm; exocarp tomentellous to glabrescent. 

Liberia, Ivory Coast and Ghana. 22: GHA, IVO, LBR. Riversides. Map 132. 

22. GHANA: Asuandi, K.O.Darko 1038 (K, P); Bonsa R., Tarquah Distr., C.Vigne 973 (K). LIBERIA: Grand 

Bassa, M.Dinklage 2023 (BM, K, P, W); E Prov., Tchien Distr., Zeahtown, J.T.Baldwin 6980 (K). 

Some small-flowered specimens from Ghana resemble D. dichotoma and D. pierrei. 

28. Dactyladenia cinerea (Engl. ex De Wild.) Prance & F.White, Brittonia 31: 484 (1979) 

Acioa cinerea Engl. ex De Wild., Bull. Jard. Bot. État 7: 211 (1920). T: Cameroon: Bipinde, G.Zenker 

2903; holo: B, not destroyed; iso: BM, BR, K, P, W, WAG, WU, Z. 


Shrub to 15 m tall. Branches lanate-arachnoid pubescent when young, slowly caducous. 

Leaves with stipules triangular-subdeltoid, 6–8 mm long; petiole 6 mm long, canaliculate, 

glabrous; lamina elliptic, 12–14 (–18) × 6–7 (–9) cm, glabrous beneath, with 2 large circular 

glands near base and other glands c. 5 mm from margins; cuneate at base, acuminate at apex


with acumen c. 1 cm long; midrib glabrous; primary veins 6–9 pairs. Inflorescences terminal 

and axillary cymes, 8–10 cm long, slender, few-flowered, arachnoid pubescent when young; 

bracteoles c. 0.5 mm long, ovate-deltate, with a clavate apical and c. 6 marginal stipitate 

glands; upper pedicel 2–3 mm long. Receptacle tube 1–1.5 cm long, white lanate-pubescent 

on exterior. Sepals 4–5 cm long, with dense white lanate pubescence on exterior. Petals c. 5 mm 

long. Stamens c. 20, 2–3 cm long. Fruit unknown. 

Cameroon; known only from the Bipinde area. 23: CMN. Habitat unknown. Map 133. 

23. CAMEROON: Bipinde, G.Zenker 3210 (BR, K, P, US). 

Closely related to D. chevalieri. 

29. Dactyladenia johnstonei (Hoyle) Prance & F.White, Brittonia 31: 485 (1979) 

Acioa johnstonei Hoyle, Bull. Misc. Inform. Kew 1932: 258 (1932). T: Cameroon: Bamenda Division, Bum, 

A.T.Johnstone 74/31; holo: K; iso: FHO. 


Shrub or small tree. Branches hispid when young, with scattered hairs 2–3 mm long. Leaves 

with stipules triangular-elongate, c. 4 mm long, with stipitate glands on the margins; petioles 

5–6 mm long, rugose; lamina oval-elliptic, 6–12 × 2–5 cm, sometimes oblong, scabrous 

beneath, very sparsely hispid soon glabrescent, with small circular glands c. 5 mm from 

margin and at base, obtuse to rounded at base, bluntly acuminate at apex; midrib sparsely 

hispid, glabrescent beneath; primary veins c. 6 pairs. Inflorescence terminal and axillary 

cymes, 1or 2-branched, 10 cm long; rachis hispid, puberulous; bracts deltate, with c. 3–5 

stipitate proximal glands 2 mm long and shorter glands distally. Bracteoles c. 1 mm long, 

subterminal, broadly deltate, with 5–7 marginal stipitate glands 1.5 mm long alternating with 

shorter ones. Upper pedicel 4–6 mm long. Receptacle tube 1–1.2 cm long, gibbous at base. 

Sepals c. 5 mm long. Petals broadly obovate, white, obtuse at base, 5–6 mm long. Stamens c. 20, 

2.5 cm long. Fruit ovoid, 25 × 40 × 35–40 mm; exocarp olive-brown tomentose. Fig. 66. 

Nigeria and adjacent Cameroon. 22: NGA. 23: CMN. Riverine forest and edges of upland 


22. NIGERIA: Ogoja Prov., Obudu Cattle Ranch, J.B.Hall 3021 (K). 23. CAMEROON: R. Katsina, between 

Wum & Nkambe, F.N.Hepper & J.R.Charter 1927 (BR, K, P); Gepka, 5 km N of Nkambe, R.Letouzey 13219 (P); 

Fonfuka, nr Bamenda, J.B.Maitland 1401 (K, paratype). 

30. Dactyladenia chevalieri (De Wild.) Prance & F.White, Brittonia 31: 484 (1979) 

Acioa chevalieri De Wild., Bull. Jard. Bot. État 7: 213 (1920). T: Gabon: lac Oghémoué, F.Fleury in 

A.Chevalier 26665; lecto: P, fide G.T.Prance & F.White, Brittonia 31: 484 (1979); isolecto: BR, WAG; photo: BR. 


Shrub or tree to 30–35 m tall. Branches arachnoid-tomentose when young, glabrescent, 

lenticellate. Leaves with stipules lanceolate, 8 × 2 mm, arachnoid-tomentose when young; 

petiole 3–8 mm long, tomentose, soon glabrescent; lamina elliptic, 6–18 × 3–8.5 cm, 

caducous arachnoid-tomentose beneath, rounded to subcordate at base, acuminate at apex; 

midrib arachnoid pubescent when young; primary veins 6–8 (–10) pairs. Inflorescences 

terminal and axillary dichotomous cymes, 8–12 cm long; rachis tomentose; bracts and 

bracteoles 3 mm long, broadly ovate, with 7–9 stipitate glands 0.5 mm long; upper pedicels 

(2–) 4–6 mm long. Receptacle tube 1–1.5 cm long. Sepals 4–5 mm long, tomentose on 

exterior, with 7–9 stipitate glands on margins. Petals obovate, cuneate at base, 3–4 mm long. 

Stamens c. 20, 2–3 cm long. Fruit shortly stipitate, subovoid, 25 × 30 × 45 mm; exocarp 

tomentose. 

Known only from Bas-Ogooué, in Gabon. 23: GAB. River and lake margins. Map 135. 

23. GABON: Lac Awounga, 50 km WSW of Lambaréné, J.Dybowski 118 (syntypes BR, P); Lac Ezanga, 25 km S 

of Lambaréné, N.Hallé 2083 (BR, P); Île Njigoné, Lac Onangé, 40 km SW of Lambaréné, C.H.Pobéguin 28 (P); 

Île Njigoné, Lac Onangé, 40 km SW of Lambaréné, C.H.Pobéguin 108 (P). 

93

94 


Figure 66. Dactyladenia johnstonei. A, habit; B, flower; C, petals; D, fruit (A–C, F.N.Hepper 

& J.R.Charter 1927; D, J.B.Maitland 1401). Scale bars: A, D = 2 cm; B, C = 5 mm. Drawn by 

Margaret Tebbs.

CHRYSOBALANACEAE 

16. HIRTELLA 

Hirtella L., Sp. Pl. 34 (1753); Gen. Pl. ed. 5: 20 (1754); from the Latin, hirtus (hairy). 

Type: H. americana L. 

Trees and shrubs. Leaves with stipules lateral, often subulate or filiform, subpersistent; 

petiole eglandular; lower surface of lamina glabrous or with a few strigose or strigulose 

hairs; lamina sometimes with 2 large bulbous myrmecophilous inflations at base of lamina, 

otherwise lacking large basal glands but often with many small submarginal or scattered 

discoid glands. Inflorescence usually a lax raceme with patent flowers, or an elongate narrow 

lax thyrse with patent lateral branches, the latter often bearing several sterile bracts and 

ending in a single flower or a few cymosely arranged flowers, more rarely a corymb, or very 

short raceme, or dense thyrse, or complex cyme. Pedicels usually longer than and often much 

longer than receptacle tube. Bracts and bracteoles not enclosing flower-buds in small groups, 

often with stalked or sessile glands. Flowers bisexual, slightly zygomorphic. Receptacle tube 

subcampanulate to narrowly cylindric, slightly gibbous, usually shorter than sepals, usually 

glabrous inside except near throat. Sepals 5, subequal, usually spreading or reflexed, acute, 

often with sessile or shortly stalked glands on margin. Petals 5, shorter than sepals. Stamens 

3–9; filaments usually coiled in bud, rarely laxly undulate with a single undulation, inserted 

on posterior rim of faucal annulus, far-exserted, usually much longer than combined length 

of calyx and receptacle tube. Staminodes short, filiform. Ovary monocarpellary, 1-locular, 

usually inserted at mouth of receptacle tube; style filiform, far-exserted, very shortly 3-lobed 

at apex. Drupe usually less than 2.2 cm long, with exiguous mesocarp and smooth thin hard 

non-granular endocarp with 4–7 longitudinal shallow channels representing the lines of 

weakness that permit the seedling to escape. 

107 species, of which 105 are confined to tropical and subtropical America while H. zanzibarica 

and H. thouarsiana occur in eastern Africa and Madagascar, respectively. 

Germination is cryptocotylar; cataphylls c. 5, minute; eophylls alternate, with small, 

subulate, persistent stipules (J.A.Duke 1965: 320, t. 4 fig. 36; G.T.Prance Blastogeny colln. 

no. 9 = H. duckei). 

A.P. de Candolle, Prodr. 2: 528 (1825); C.F.P.Martius & J.G.Zuccarini, Chrysobalaneae, in 

J.G.Zuccarini, Plantarum novarum vel minus cognitarum descripta, Abh. Math.-Phys. Cl. Königl. 

Bayer. Akad. Wiss. München 1: 372 (1832); J.G.Zuccarini, Flora, 15 (Beibl. 2): 78 (1832); 

J.D.Hooker, in G.Bentham & J.D.Hooker, Gen. Pl. 1: 608 (1865); W.O.Focke, in A.Engler & 

K.A.E.Prantl, Nat. Pflanzenfam. 3 (3): 59, t. 31–32 (1894); J.Hutchinson, Gen. Fl. Pl. 1: 193 

(1954); G.T.Prance, Fl. Neotrop. Monogr. 9: 259, t. 43–57 (1972); F.White, The taxonomy, 

ecology and chorology of African Chrysobalanaceae (excluding Acioa), Bull. Jard. Bot. 

Belg. 46: 276 (1976); F.White, Distrib. Pl. African. 10: 287–292 (1976); F.White, Fl. Zamb. 

4: 44, t. 9 fig. b (1978); G.T.Prance & F.White, The genera of Chrysobalanaceae, Philos 

Trans. B320: 145–149 (1988). 

Key to Groups of Hirtella 

1 Leaf base with myrmecophilous swellings [sect. Myrmecophila] Group A 

1: Leaf base without myrmecophilous swellings [sect. Hirtella] 

2 Inflorescence a panicle (branched) Group B 

2: Inflorescence a raceme (unbranched) Group C 

95

96 


Key to Group A [Hirtella sect. Myrmecophila] 

1 Inflorescence fasciculate (bunched racemes), sometimes cauliflorous 

2 Stamens 4–5; ovary inserted at middle of receptacle; leaves 9–19 cm 

long, coriaceous with lower surface bullate 1. H. myrmecophila 

2: Stamens 6–7; ovary inserted at mouth of receptacle; leaves 17–30 cm 

long, membranous, with lower surface not bullate 2. H. physophora 

1: Inflorescence an elongate raceme or panicle, never cauliflorous 

3 Inflorescence a panicle, sometimes only slightly branched at base 

4 Inflorescence only slightly branched at base; leaves oblonglanceolate; 

bracts eglandular 3. H. vesiculosa 

4: Inflorescence with many small lateral branches; leaves ovate; bracts 

glandular 4. H. dorvalii 

3: Inflorescence an elongate raceme, never branched 

5 Exterior of receptacle and calyx lobes hirsutulous; inflorescence 

15–30 cm long 5. H. guainiae 

5: Exterior of receptacle and calyx lobes hispid only; inflorescence 

2–12 cm long 

6 Pedicels 4–6 mm long, with sparse hirsute pubescence 6. H. duckei 

6: Pedicels 8–15 mm long, with dense hirsute pubescence 7. H. revillae 

Key to Group B [Hirtella sect. Hirtella, species with a paniculate inflorescence] 

1 Bracts and/or pedicels glandular (with stalked or sessile glands) 

2 Leaves glabrous beneath when mature, or rarely with a few short stiff 

appressed hairs 

3 Stamens 6 or 7 

4 Bracts with a few sessile glands only; inflorescence rachis and 

branches and flower exterior glabrous or only sparsely hirsutulous 

5 Inflorescence a spreading branched panicle; rachis and branches 

glabrous 8. H. macrosepala 

5: Flowers borne in clusters of 2–3 along a central rachis rather than a 

branched inflorescence, giving a racemose appearance; rachis and 

branches hirsutulous 

6 Leaf lamina 6–9.5 × 2.5–3.8 cm; acumen 9–14 mm long; petioles 

2–2.5 mm long 14. H. barnebyi 

6: Leaf lamina 21–27 × 6–9 cm; acumen 1–5.5 mm long; petioles 

8–10 mm long 15. H. margae 

4: Bracts with numerous stalked glands; inflorescence rachis and 

branches and flower exterior pubescent 

7 Leaves 3.5–8.5 × 1.4–4.5 cm; stipules linear, to 6 mm long [Africa] 

8 Receptacle tube distinctly longer than the sepals [East Africa] 87. H. zanzibarica 

8: Receptacle tube shorter than the sepals [Madagascar] 88. H. thouarsiana 

7: Leaves (6.4–) 8–13 × 4–7 cm; stipules deltoid, to 2 mm long 

[Neotropics] 

9 Calyx lobes glandular; bract glands small, c. 0.1 mm in diam.; 

rachis and branches of inflorescence sparsely puberulous; leaves 

glabrous beneath 9. H. ulei

CHRYSOBALANACEAE (Hirtella) 

9: Calyx lobes eglandular; bract glands large, c. 0.5 mm in diameter; 

rachis and branches of inflorescence tomentellous; leaves 

sparsely appressed pubescent beneath 10. H. glabrata 

3: Stamens 3–5 

10 Glands large and solitary on a long stalk, arising from pedicel or 

junction of inflorescence; inflorescence distinctly thyrsoid 12. H. carbonaria 

10: Glands usually numerous, long or short stalked, arising from 

margins of bracts and bracteoles; inflorescence not thyrsoid 

11 Inflorescence a long rachis bearing clusters of 1–3 flowers on short 

branches giving a racemose appearance, bracts and bracteoles with 

few sessile glands only 

12 Flowers densely clustered in small groups; leaves oblonglanceolate, 

10–17 cm long; primary veins 10–13 pairs 13. H. araguariensis 

12: Flowers not clustered, only 1–2 per branch; leaves lanceolate or 

oblong, 9–12 cm long; primary veins 7–9 pairs 

13 Leaves lanceolate; most flowers solitary 35. H. castilloana 

13: Leaves oblong; most flowers in pairs 32. H. papillata 

11: Inflorescence spreading, with many distinct lateral branches, 

flowers not in distinct clusters; leaves oblong to elliptic 

14 Leaves 3–5.5 cm long, cordate at base; inflorescence scarcely 

branched, hirsutulous 17. H. cordifolia 

14: Leaves usually cuneate or subcuneate at base (if subcordate then 

7–17 cm long and inflorescence glabrescent); inflorescences 

usually much branched, glabrescent to pilose 

15 Bracteolar glands few, usually either sessile on margins or a 

single gland terminating apex; inflorescence usually pilose or if 

glabrescent then corymbose; leaf apex acuminate 

16 Leaves 12–17 cm long, rounded to subcordate at base; fertile 

stamens 5 18. H. insignis 

16: Leaves 3.5–15 cm long, cuneate at base; fertile stamens 3 

17 Leaves 10–15 cm long; bracts membranous, with several 

glands borne on margins; inflorescence 7–18 cm long, 

terminal 19. H. tocantina 

17: Leaves 4–9 cm long; bracts coriaceous, with single reflexed 

apical gland only; inflorescence 3–11 cm long, axillary or 

terminal 20. H. piresii 

15: Bracteolar glands numerous, borne on the margins, usually 

short-stalked; inflorescence spreading but not corymbose, 

usually glabrescent, if tomentellous then leaf apex rounded to 

acute 

18 Leaves 1.8–4.5 cm long, rounded to acuminate at apex, thickly 

coriaceous; primary veins 4–7 pairs 48. H. bahiensis 

18: Leaves 6–13.5 cm long, acuminate at apex, thinly coriaceous; 

primary veins 9–10 pairs 

19 Stamens 3; bracteoles coriaceous, ovate; petioles 3–4 mm 

long 21. H. davisii 

19: Stamens 4; bracteoles membranous, lanceolate; petioles 

1–2 mm long 22. H. subglanduligera 

2: Leaves hirsute beneath at least on principal venation, usually more 

generally so 

20 Calyx lobes with small stipitate glands 

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98 


21 Leaves orbicular to ovate-elliptic, 3–6.5 cm long, retuse to 

mucronate at apex (rarely acuminate); trunk with corky bark; 

receptacle pubescent within to base 23. H. ciliata 

21: Leaves oblong to ovate, 4.5–23 cm long, acute to acuminate at apex; 

trunk with thin bark; receptacle glabrous within except at throat 

22 Venation of leaves prominulous above; inflorescence 

subcorymbose, bearing numerous tightly clustered flowers; 

stamens 7 24. H. hoehnei 

22: Venation of leaves impressed or prominulous above; inflorescence 

lax, spreading, not subcorymbose, with flowers loosely arranged; 

stamens 3–5 

23 Leaves subcordate at base, subconduplicate; lower surface merely 

sparsely appressed-pubescent 39. H. adderleyi 

23: Leaves rounded to cuneate at base, rarely weakly subcordate but 

never subconduplicate; lower surface hirsute 25. H. glandulosa 

20: Calyx lobes eglandular 

24 Venation of leaves impressed above, or the leaf surface bullate 

25 Inflorescence and flowers rufous-tomentose; bracts bearing very 

few sessile glands, these often obscured by pubescence 54. H. obidensis 

25: Inflorescence and flowers tomentellous to puberulous, not rufous; 

bracts bearing many obvious stalked or sessile glands 

26 Leaf surface usually distinctly bullate; base usually subcordate 26. H. bullata 

26: Leaf upper surface plane, but with impressed venation; base 

rounded to subcuneate 

27 Bracteolar glands c. 0.5 mm in diameter, sessile or slightly 

stipitate; bracteoles distinct, ovate 

28 Leaves 2–7 cm broad; pedicels 0.5–2 mm 27. H. americana 

28: Leaves 8–9.5 cm broad; pedicels 3–5 mm 29. H. aequatoriensis 

27: Bracteolar glands c. 0.1 mm in diameter, on slender stalks; 

bracteoles reduced to a mass of glands with a slightly connate 

base 37. H. tentaculata 

24: Venation of leaves prominulous or plane above, the leaf surface 

never bullate 

29 Stamens 3 

30 Flowers borne in groups of 3 31. H. trichotoma 

30: Flowers not as above 

31 Bracteolar glands large, numerous, apparent; inflorescence 

tomentellous, many-flowered 27. H. americana 

31: Bracteolar glands small, few; inflorescence hispid or tomentose, 

few-flowered 

32 Bracteolar glands stipitate; inflorescence rachis hispid; 

flowers 6–8 mm long 30. H. guatemalensis 

32: Bracteolar glands sessile; inflorescence rachis tomentose; 

flowers c. 5 mm long 45. H. beckii 


33 Exocarp tomentose; endocarp thick and fibrous; bracts with few 

inconspicuous stipitate or sessile glands 

34 Inflorescence hispid, with flowers lax; bracts with small shortstipitate 

glands; stamens 4 30. H. guatemalensis


34: Inflorescence tomentellous, bearing small clusters of flowers on 

a long thick rachis; bracts with a few large sessile glands or a 

single terminal gland; stamens 6 33. H. eriandra 

33: Exocarp glabrescent; endocarp thin and bony; bracts with many 

conspicuous stipitate glands, except in H. liesneri 

35 Leaves lanceolate; inflorescence with well spaced flowers; 

bracteoles 6–9 mm long 35. H. castilloana 

35: Leaves oblong to ovate; inflorescence with flowers crowded; 

bracteoles 2–4 mm long 

36 Inflorescence hispid-hirsute; bracts with short-stalked glands 

arising from their margins; gland secretory tip small, ellipsoid 

37 Inflorescences predominantly terminal; leaf bases rounded to 

weakly subcordate 34. H. paniculata 

37: Inflorescences predominantly axillary; leaf bases deeply cordate 36. H. deflexa 

36: Inflorescence puberulous to tomentellous; bracts either reduced 

to numerous, long-stalked clavate glands arising from the 

junction of the pedicels with the stem, or ovate with glands 

borne on their margins; gland secretory tip large, flattened, 

forming an expanded head 

38 Glands arising from junction of pedicel and stem, with a long 

slender stalk and a small secretory tip, clavate; young stems 

puberulous or sparsely tomentellous 38. H. macrophylla 

38: Glands arising from the bracts, each gland with a thick stalk 

and a large secretory tip, flattened to form an expanded head; 

young stems with short dense compact puberulence 

39 Leaves 4–7 cm long, conduplicate; inflorescence 3.5–8 cm 

long; stipules lanceolate 39. H. adderleyi 

39: Leaves 6.5–15 cm long, not folded; inflorescence 12–19 cm 

long; stipules deltoid or linear-lanceolate 

40 Inflorescences c. 19 cm long, with very short lateral 

branches giving a racemose appearance; glands scarce, 

many bracts with single stalked gland at apex; 

inflorescence rachis sparsely hispid at base 16. H. liesneri 

40: Inflorescences 12–15 cm long, not appearing racemose; 

glands numerous on bracteoles; inflorescence rachis 

tomentellous, never hispid 

41 Leaves cordate at base; petioles 1–3 mm long; stipules 

linear, 5 mm long; flowers densely crowded 11. H. confertiflora 

41: Leaves rounded to subcordate at base; petioles 5–7 mm long; 

stipules deltoid, to 2 mm long; flowers laxly arranged 10. H. glabrata 

1: Bracts and pedicels eglandular 

42 Leaf apex retuse, rounded, or acute, never acuminate 

43 Stamens 5–6; leaf apex mostly rounded to retuse (or mucronulate) 40. H. punctillata 

43: Stamens 3–4; leaf apex acute 

44 Leaves elliptic, thick-coriaceous; inflorescence compact, to 3 cm long 41. H. corymbosa 

44: Leaves oblong, chartaceous; inflorescence lax and spreading, 

3.5–17 cm long 47. H. triandra 

42: Leaf apex distinctly acuminate, sometimes abruptly so and mucronate 

or cuspidate 

45 Stamens 3 

46 Leaf base subcordate; inflorescence 25–55 cm long, pendulous 42. H. pendula 

99

100 


46: Leaf base rounded to cuneate; inflorescence 1–15 cm long, erect 

47 Leaves lanceolate, at least 3× longer than broad; inflorescence and 

exterior of flowers glabrescent 43. H. barrosoi 

47: Leaves oblong to elliptic, not exceeding 2.5× longer than broad; 

inflorescence and exterior of flowers usually pubescent, rarely 

glabrous 

48 Inflorescence and exterior of flowers glabrous or glabrescent, or 

leaf apex distinctly cuspidate with a long fine acumen; 

inflorescences 1–4 cm long, predominantly axillary 62. H. bicornis 

48: Inflorescence and exterior of flowers pubescent or tomentose; 

leaf apex acuminate but never cuspidate; inflorescences 2–17 cm 

long, predominantly terminal 

49 Inflorescence and young stem hispid; calyx tube c. 6 mm long 

with lobes c. 5 mm long; corolla lobes c. 8 mm long; stipules 

6–10 mm long 44. H. leonotis 

49: Inflorescence usually puberulous, rarely weakly hirsute; young 

stem glabrous to tomentellous; calyx tube to 4 mm long with lobes 

to 3 mm long; corolla lobes to 5 mm long; stipules 2–6 mm long 

50 Inflorescence little-branched, almost racemose, but with a few 

branches bearing 2 flowers, others 1 flower and a single pair 

of bracts at junction with pedicel 46. H. mutisii 

50: Inflorescence much-branched, with branches usually bearing 

many bracts 

51 Leaves 4–14.5 × 2–5.5 cm; pedicels 1–3 mm long 47. H. triandra 

51: Leaves 15–17 × 8–10 cm; pedicels 0–0.5 mm long 49. H. latifolia 


52 Inflorescence appearing racemose, but in fact either a long central 

rachis bearing groups of flowers on short branches, or with a few 

branches with only 3 flowers each; leaf base sometimes cordate 

53 Inflorescence almost a raceme, but with a few branches bearing 2 

or 3 flowers; flowers laxly arranged, not borne in distinct groups 104. H. hebeclada 

53: Inflorescence a long central rachis with groups of flowers on short 

branches 

54 Bracteoles exceeding receptacle in length, persistent, enclosing 

buds; leaf base subcuneate; exocarp glabrous 50. H. suffulta 

54: Bracteoles much shorter than receptacle, not persistent, not 

enclosing buds; leaf bases subcordate to subcuneate; exocarp 

tomentose or glabrous 

55 Leaves 18–40 × 8–16.5 cm, with 14–18 pairs of primary veins; 

2 glands at junction of upper surface of lamina and petiole; 

inflorescence rufous-tomentellous 52. H. magnifolia 

55: Leaves 8–23 × 3.3–8 cm, with 10–13 pairs of primary veins; 

lamina base eglandular; inflorescence grey-puberulous or 

brown-tomentellous 

56 Leaves subcordate at base; lower lamina surface glabrescent; 

exocarp glabrous 51. H. elongata 

56: Leaves rounded to subcuneate at base; lower lamina surface 

sparsely pilose-hirsute; exocarp tomentose 33. H. eriandra 

52: Inflorescence a much-branched panicle without a long central rachis; 

leaf base rounded to cuneate 

57 Inflorescence and exterior of flowers tomentose, hirsute, or hispid, 

usually rufous to golden-brown


58 Inflorescence and young branches sparsely hispid; leaves 

membranous 53. H. rodriguesii 

58: Inflorescence and young branches tomentose or hirsute; leaves 

chartaceous to coriaceous 

59 Venation of leaves slightly impressed above; inflorescence 

rufous-tomentose 54. H. obidensis 

59: Venation of leaves plane or prominulous above; inflorescence 

brown-tomentose or hirsute 

60 Leaves 2.5–6 cm long; inflorescence 0.8–4 cm long 

61 Inflorescence 0.8–1.5 cm long, clustered; leaves ovate, 

rounded at base; stamens 7 55. H. cowanii 

61: Inflorescence 2–4 cm long, lax; leaves orbicular to oblongorbicular, 

usually subcordate at base; stamens 5 56. H. orbicularis 

60: Leaves 5.5–17 cm long; inflorescence 4–11 cm long 

62 Bracts 4–6 mm long, ovate, up to 1 /2 the length of receptacle; 

leaves ovate, thick-coriaceous, shortly mucronate at apex; 

inflorescence tomentose 57. H. guyanensis 

62: Bracts 1–3 mm long, oblong, to base of receptacle only; 

leaves elliptic, subcoriaceous, acuminate (never mucronate) 

at apex; inflorescence hirsutulous-tomentose 58. H. lightioides 

57: Inflorescence and exterior of flowers grey-puberulous to 

glabrescent 

63 Inflorescence a subcorymbose panicle 4–6.5 cm long; petioles 

7–8 mm long 59. H. aramangensis 

63: Inflorescence a spreading panicle (not subcorymbose); petioles 

0.5–5 mm long 

64 Lamina 9–15 cm long, with distinct basal glands; inflorescence 

lax and spreading, 7–19 cm long 60. H. rasa 

64: Lamina 2.5–8 (–9.5) cm long, with base eglandular or with 

indistinct glands; inflorescence short and compact, 1–8 cm long 

65 Inflorescence and exterior of flowers densely puberulous; 

leaves thick and coriaceous, with prominent venation; apex 

acuminate but never cuspidate 61. H. scabra 

65: Inflorescence and exterior of flowers glabrescent to 

puberulous; leaves chartaceous, with plane to prominulous 

venation; apex acuminate to cuspidate 62. H. bicornis 

Key to Group C [Hirtella sect. Hirtella, species with a racemose inflorescence] 

1 Glands present on bracts and bracteoles or pedicels (either stipitate or 

sessile) 

2 Leaves narrowly linear-lanceolate, 5.5–18 × 0.5–2.2 cm; rheophytic 63. H. angustissima 

2: Leaves ovate to oblong-lanceolate, 3–30 × 1.5–10 cm; not rheophytic 

3 Bracts and bracteoles bearing sessile glands only or with translucent 

glandular excretions, with glands often covering entire surface, most 

frequently paired toward bases of bracts and bracteoles; stipitate 

glands absent from bracts and pedicels, but bract apex occasionally 

glandular 

4 Leaf apex with curved acumen 25–35 mm long; stamens 9 or 10, 

young branches pilose 67. H. maguirei 

101

102 


4: Leaf apex with acumen to 18 mm long; stamens 4–7; young branches 

rarely pilose 

5 Leaves oblong-lanceolate; young stems hispid 

6 Leaves 9–14 cm long; pedicels 5–10 mm long; stipules eglandular 64. H. tenuifolia 

6: Leaves 5–8 cm long; pedicels 12–16 mm long; stipules with 

numerous stipitate glands 65. H. radamii 

5: Leaves oblong to ovate; young stems usually puberulous, less 

frequently hispid 

7 Young stem hispid; inflorescence rachis sparsely puberulous 72. H. racemosa 

7: Young stem and inflorescence rachis pilose or puberulous (hispid 

in H. pilosissima) 

8 Young stem and inflorescence thick-pilose or pilose-tomentose, 

the stem with persistent hairs 2–3 mm long; stamens 3–6 

9 Leaf margins hispid-ciliate; stamens 5–6; pedicels 1.5–3 mm 

long; bracts axillary 66. H. pilosissima 

9: Leaf margins ciliate; stamens 3; pedicels 3–6 mm long; bracts 

inserted on pedicels 44. H. leonotis 

8: Young stem and inflorescence usually puberulous, sometimes 

tomentellous; stamens 4–6 

10 Leaves thin, membranous; stamens 4 or 5; ovary inserted 

midway up receptacle 

10: Leaves thick, coriaceous; stamens 5 or 6; ovary inserted at 

mouth of receptacle 

11 Bracts and bracteoles caducous, with minute translucent 

secretions, sessile glands present or absent; pedicels 

6–25 mm long 

12 Bracts with sessile glands in addition to translucent 

secretion; pedicels 6–16 mm long, thick; inflorescence 

4.5–14 cm long; leaves sparsely appressed-pubescent on 

53. H. rodriguesii 

venation or glabrescent beneath 

12: Bracts with translucent glandular secretions only, other 

glands absent; pedicels 12–25 mm long, very slender; 

68. H. gracilipes 

inflorescence 1.5–5 cm long; leaves glabrous beneath 

11: Bracts and bracteoles persistent, with sessile glands only; 

pedicels 1.5–10.5 mm long 

13 Primary veins 6–10 pairs; leaves 3.5–18.5 (–19.5) cm long, 

glabrous or sparsely appressed-pubescent beneath; mature 

69. H. brachystachya 

leaves plane, not bullate 

13: Primary veins 11–15 pairs; leaves 19.5–25 cm long or if 

smaller (7–13 cm long) then hirsutulous beneath; mature 

leaves slightly bullate 

14 Leaves 19.5–25 cm long; primary veins 13–15 pairs; 

72. H. racemosa 

lower surface with few sparse appressed hairs 

14: Leaves 7–13 cm long; primary veins 11–13 pairs; lower 

73. H. juruensis 

surface hirsutulous 74. H. kuhlmannii 

3: Either bracts and bracteoles bearing stipitate glands, or with 1 or more 

solitary stipitate glands arising from some pedicels, and bracts often 

glandular at apex only 

15 Bracts and bracteoles eglandular, or with sessile glands and a 

glandular apex only; some pedicels with 1–3 stipitate glands 

16 Leaves ovate, cordate at base; bracts and bracteoles with a few 

sessile glands 75. H. standleyi


16: Leaves oblong to oblong-lanceolate, usually rounded to subcuneate 

at base, rarely cordate; bracts and bracteoles eglandular or with 

sessile glands 

17 Primary veins 17–19 pairs; leaves 19–26 cm long; pedicels 

8–12 mm long 76. H. longifolia 

17: Primary veins 8–15 pairs; leaves 8–18 cm long (to 24 cm in 1 species 

with short pedicels 0.5–2 mm long); pedicels 0.5–9 mm long 

18 Pedicels slender, 4.5–9 mm long (or if 3.5–5 mm then leaf 

constricted above base); leaves glabrous beneath except for 

pubescence on midrib and margin of extreme base of lamina; 

bracteoles eglandular or glandular 

19 Fertile stamens 3; leaves oblong-elliptic; bracts with apical 

glands 77. H. lemsii 

19: Fertile stamens 5–7; leaves elliptic to oblong-lanceolate; 

bracts glandular or eglandular 

20 Leaves elliptic to oblong-elliptic, 3.5–8 cm long; bracts 

usually with sessile and apical glands 72. H. racemosa 

20: Leaves oblong-lanceolate, 8–18 cm long; bracts eglandular 

21 Leaves subauriculate because of slight constriction above 

base; primary veins 11–13 pairs; ovary inserted near mouth 

of receptacle; fertile stamens 5 78. H. schultesii 

21: Leaves not constricted above base; primary veins 8–15 

pairs; ovary inserted at mouth or middle of receptacle; 

fertile stamens 4 or 6 

22 Primary veins 12–15 pairs; secondary venation slightly 

impressed; ovary inserted midway up receptacle; fertile 

stamens 4 79. H. paraensis 

22: Primary veins 8–10 pairs; tertiary venation prominulous; 

ovary inserted at mouth of receptacle; fertile stamens 6 80. H. sprucei 

18: Pedicels 1–3 mm long, thick; leaves hirsute beneath or glabrous 

with hirsute margin and midrib; bracteoles with glandular apex 

and often sessile glands 

23 Leaves bullate above, with distinctly impressed venation, 

secondary venation and reticulations extremely prominent 

beneath 81. H. lancifolia 

23: Leaves not bullate above, with venation prominent or plane, 

secondary venation and reticulations prominulous only beneath 

24 Leaves acuminate at apex; lower surface glabrous to sparsely 

hirsute 82. H. burchellii 

24: Leaves usually mucronate at apex; lower surface densely 

hirsute or glabrous 

25 Inflorescence densely tomentose; leaves mucronate at apex, 

hirsute beneath 83. H. mucronata 

25: Inflorescence sparsely puberulous; leaves acuminate at 

apex, glabrescent beneath 72. H. racemosa 

15: Bracts and bracteoles with numerous stipitate glands; pedicels 

eglandular 

26 Receptacle cylindrical, elongate 98. H. couepiiflora 

26: Receptacle campanulate 

27 Leaves bullate above, with impressed venation 26. H. bullata 

27: Leaves plane, not bullate above, with plane or prominulous 

venation 

103

104 


28 Leaves orbicular, rounded to retuse at apex; pedicels 7–20 mm 

long 84. H. longipedicellata 

28: Leaves oblong to elliptic, acuminate at apex; pedicels 

4–11 (–15) mm long 

29 Stipules bearing numerous stipitate glands 85. H. glandistipula 

29: Stipules eglandular 

30 Inflorescence short, compact, densely flowered, 3–7 cm 

long; calyx bearing numerous glands; leaves drying grey 86. H. martiana 

30: Inflorescence lax, 3–28 cm long; calyx eglandular or the 

glands scarce; leaves drying green to brown 

31 Calyx lobes 5–6 mm long; stamens 7 or 8; flowers 8–9 mm 

long 101. H. angustifolia 

31: Calyx lobes 1.5–4 mm long; stamens 3–6; flowers usually 

4–6 mm long (7–9 mm in H. santosii) 

32 Young branches and lower part of inflorescence rachis 

hispid-setose 

33 Leaves 2.5–6.5 cm long; inflorescence 3–5 cm long 90. H. pimichina 

33: Leaves 6–15 cm long; inflorescence 6–28 cm long 

34 Leaves entirely glabrous beneath, dried material with 

silver metallic sheen 91. H. subscandens 

34: Leaves hirsute beneath especially on principal 

venation, dried material without silver metallic sheen 

35 Bracteoles oblong, broad toward base, chartaceous, 

with short-stalked glands; apices terminating in a 

short-stalked gland 34. H. paniculata 

35: Bracteoles linear-lanceolate, coriaceous, with longstalked 

glands; apices always terminating in a longstalked 

gland 92. H. hispidula 

32: Young branches tomentellous to puberulous; lower part of 

inflorescence glabrous to puberulous or tomentellous 

36 Inflorescence tomentose to tomentellous 

37 Inflorescence tomentellous, 4.5–27 cm long; leaves 

broadest above middle; primary veins 8–12 pairs; 

pedicels 4–9 mm long; bract glands predominantly 

stipitate; stipules 2–3 mm long 93. H. silicea 

37: Inflorescence tomentose, 6–10 cm long; leaves 

broadest at middle; primary veins 15–19 pairs; 

pedicels 5–6 mm long; bract glands predominantly 

sessile; stipules 4–5 mm long 28. H. santosii 

36: Inflorescence glabrescent or very sparsely setose 

38 Stamens 3; calyx glandular 94. H. excelsa 

38: Stamens 5–6; calyx eglandular 

39 Leaf gradually tapered to apex from near base; 

pedicels 3–5 mm long 95. H. adenophora 

39: Leaf apex cuspidate; pedicels 8–15 mm long 96. H. caduca 

1: Bracts and pedicels eglandular 

40 Inflorescence clustered, fasciculate; leaves slightly bullate, 8–12.5 cm 

long; ovary inserted near base of receptacle 97. H. fasciculata 

40: Inflorescence of elongate racemes; leaves usually plane, if bullate 

then small; ovary inserted at or near mouth of receptacle


41 Receptacle elongate-cylindrical; calyx lobes glandular 

42 Inflorescences densely tomentose; leaves 10–15 cm long; flowers 

17–20 mm long 98. H. couepiiflora 

42: Inflorescences sparsely puberulous; leaves to 8.5 cm long; flowers 

7–9 mm long 99. H. tubiflora 

41: Receptacle campanulate; calyx lobes eglandular 

43 Leaves orbicular, predominantly retuse to rounded at apex; 

pedicels 7–20 mm long 84. H. longipedicellata 

43: Leaves oblong-lanceolate to ovate, acuminate at apex; pedicels to 

15 mm long, usually shorter 

44 Lower surface of leaf hirsute, pilose or hispid at least on primary 

and secondary venation 

45 Leaves oblong, oblong-lanceolate, or lanceolate; venation 

sometimes deeply impressed above; fertile stamens 7–9 

46 Leaves oblong, 3.5–8 cm long; venation deeply impressed 

above; flowers 4 mm long 100. H. floribunda 

46: Leaves lanceolate to oblong-lanceolate, 5–15 cm long; 

venation plane above; flowers c. 6 mm long 101. H. angustifolia 

45: Leaves ovate to elliptic; venation usually prominent above, 

rarely slightly impressed but then leaves broadly ovate; fertile 

stamens 3–9 

47 Leaf venation slightly impressed above; leaves broadest near 

base, 2.5–8 cm long, subcordate at base; inflorescence densely 

crowded, with short rachis 102. H. rugosa 

47: Leaf venation level to slightly prominent above; leaves 

broadest at or above middle, subcuneate to rounded at base; 

inflorescence lax with longer rachis 

48 Stamens 3 46. H. mutisii 


49 Leaf apex mucronate to abruptly acuminate; pedicels 

2–3 mm long 103. H. scaberula 

49: Leaf apex acute to acuminate but never mucronate or 

abruptly acuminate; pedicels 5–8 mm long 

50 Inflorescence and exterior of flowers densely tomentose; 

flowers 5–8 mm long 104. H. hebeclada 

50: Inflorescence and exterior of flowers puberulous; flowers 

4–5 mm long 72. H. racemosa 

44: Lower surface of leaf glabrous or with few sparse appressed hairs 

on midrib and primary veins 

51 Leaves narrowly lanceolate, 5.5–18 cm long, 0.5–2.2 cm broad 63. H. angustissima 

51: Leaves ovate to oblong 

52 Fertile stamens 7–9 

53 Petals slightly clawed, leaf base subcuneate; young branches 

glabrescent; inflorescence of densely crowded racemes 89. H. parviunguis 

53: Petals not clawed; leaf base subcordate to subcuneate; young 

branches puberulous to hispid; inflorescence of single racemes 

54 Leaf base subcordate; calyx lobes 4–6 mm long; young 

branches sparsely hispid 101. H. angustifolia 

54: Leaf base subcuneate to rounded; calyx lobes 1.5–2.5 mm 

long; young branches sparsely puberulous 105. H. enneandra 

105

106 


52: Fertile stamens 3–6 

55 Fertile stamens 3; inflorescence either 2–3-flowered or 

densely many-flowered and tomentose; filaments only 

slightly exceeding calyx lobes 

56 Leaves 7–13 cm long, 3–5 cm broad, chartaceous; 

inflorescence 2–3-flowered 106. H. pauciflora 

56: Leaves 3.5–5.5 cm long, 1.5–2.7 cm broad, thickcoriaceous; 

inflorescence many flowered 107. H. glaziovii 

55: Fertile stamens 4–6; inflorescence puberulous to tomentellous, 

many-flowered; filaments far exceeding calyx lobes 

57 Leaves thick coriaceous; inflorescence 4–15 cm long 72. H. racemosa 

57: Leaves chartaceous; inflorescence 2–4 cm long 

58 Leaf apex bluntly acuminate or acute; petioles 0.5–1 mm; 

flowers c. 3 mm long 70. H. arenosa 

58: Leaf apex caudate; petioles 1.5–2.5 mm; flowers c. 6 mm 

long 71. H. conduplicata 

Hirtella sect. Myrmecophila 

Hirtella sect. Myrmecophila Prance, Fl. Neotrop. Monogr. 9: 261 (1972) 

T: Hirtella myrmecophila Pilg. 

Leaf bases with 2 swollen ant-cavities. 

A section of 7 species. Guianas and Amazonia. 

1. Hirtella myrmecophila Pilg., Verh. Bot. Vereins Prov. Brandenburg 47: 148 (1905) 

T: Brazil North: Amazonas, Rio Negro, Manaus, Flores, E.Ule 6916; holo: B, destroyed; lecto: HBG, fide 

G.T.Prance, Fl. Neotrop. Monogr. 9: 271 (1972). 

Hirtella myrmecophila var. tetrandra Huber, Bol. Mus. Paraense Hist. Nat. 5: 369 (1909). T: Brazil North: 

Pará, Faro, Ajurua, A.Ducke MG8557; holo: MG; iso: BM, P, US. 


Tree to 7 m tall. Branches sparsely hispid when young. Leaves with stipules filamentous, to 

12 mm long, subpersistent, hispid; petioles 4–6 mm long, sparsely pilose to hispid, enveloped 

by paired ant-cavities; lamina coriaceous, oblong to oblong-lanceolate, 9–19 × 4–7 (–8.5) cm, 

acuminate at apex with acumen 6–17 mm long, subcordate at base, slightly bullate beneath, 

with sparse hispid pubescence on principal venation; primary veins 12–14 pairs, prominent 

above; midrib prominent above. Inflorescences fasciculate racemes, 1–3.5 cm long; rachis 

glabrescent with a few hispid hairs. Bracts and bracteoles linear, to 3.5 mm long, usually 

with paired sessile basal glands, hispid. Flowers c. 4 mm long. Receptacle campanulate, sparsely 

hispid on exterior, glabrous within except for deflexed hairs at throat; pedicels 4–5 mm long. 

Calyx lobes acute, sparsely hispid on exterior, puberulous within. Petals 5, glabrous. 

Stamens 4–5, unilateral; filaments far exceeding calyx lobes. Style glabrous except for 

hirsute base. Ovary inserted midway up receptacle tube, pilose. Fruit not seen. Fig. 67. 

Brazil (Central Amazonia). 84: BZN (AM, PA). Non-flooded forest. Map 136. 

84. BRAZIL NORTH: Amazonas, Manaus, Reserva Florestal Ducke, M.A.S.Costa 344 (INPA, K); Amazonas, 

Manaus, Tarumã-Ponta Negra road, E.Lleras P17561 (INPA, NY); Amazonas, Manaus, Uipiranga, J.G.Kuhlmann 

RB17977 (K, RB, S); Pará, Mun. de Oriximiná, Rio Trombetas, Porto Trombetas, C.A. Cid Ferreira et al. 1899 

(INPA, NY).


Figure 67. Hirtella myrmecophila. A, habit; B, leaf base showing ant pouches; C, ½ flower; 

D, fruit (A–B, C.A. Cid Ferreira et al. 8214; C, A.Ducke 25008; D, E.Lleras et al. P17561). 

Scale bars: A = 3 cm; B, D = 1 cm; C = 5 mm. Drawn by Margaret Tebbs. 

107

108 


2. Hirtella physophora Mart. & Zucc., Abh. Math.-Phys. Cl. Königl. Bayer. Akad. Wiss. 

1: 374 (1832) 

T: Brazil North: Amazonas, C.F.P.Martius s.n.; holo: M. 

Hirtella cauliflora Huber, Bol. Mus. Paraense Hist. Nat. 6: 73 (1910). T: Brazil: Amapá, Rio Oiapoque, 

A.Ducke MG4745; holo: MG; iso: BM, K. 

Hirtella cotticaensis Kleinhoonte, Recueil Trav. Bot. Néerl. 22: 386 (1925). T: Suriname: Cottica River, 

J.W.Gonggrijp BW5333; holo: U; iso: NY. 



Tree to 6 m tall. Stems sparsely hispid when young, lenticellate. Leaves with stipules 

filamentous, to 1.5 cm long, subpersistent, hispid; petioles 5–8 mm long, hispid, enveloped by 

swollen ant-cavities which fuse with leaf base; lamina membranous, oblong-elliptic to oblong, 

17–30 × 6–11 cm, acuminate to mucronate at apex with acumen 5–20 mm long, subcordate at 

base, hirsute on venation beneath; midrib prominent above; primary veins 13–16 pairs, 

prominent beneath. Inflorescences fasciculate racemes, 2–4 cm long, hispid. Bracts and 

bracteoles linear to filamentous, 2–8 mm long, hispid, persistent. Flowers 5–7 mm long. 

Receptacle campanulate, hispid on exterior, glabrous within except for deflexed hairs at 

throat; pedicels 3–4 mm long. Calyx lobes oblong-lanceolate, hispid on exterior, puberulous 

within near apex, glabrous beneath. Petals 5, glabrous, twice as long as calyx lobes. Stamens 

6, unilateral; filaments far exceeding calyx lobes. Style hirsute on lower portion up to 1/2 of 

its length. Ovary inserted at mouth of receptacle, pilose. Fruit ellipsoid; epicarp smooth, 

glabrous; mesocarp thin, fleshy; endocarp thin, hard, bony. 

Guianas and Amazonia. 82: FRG, GUY, SUR, VEN. 83: CLM, PER. 84: BZN (AC, AM, AP, 

PA, RO). Non-flooded forest. Map 137. 

82. FRENCH GUIANA: Saül, La Fumée, M.F.Prévost 1806 (CAY, P). GUYANA: U. Takutu-U. Essequibo 

Region, Kuyuwini R., 2–4 km S of Marudi Mt., T.W.Henkel 3026 (K, US). SURINAME: Confluence of Lucie 

and Oost Rivers, H.S.Irwin et al. 55520 (F, GH, NY, US). VENEZUELA: Amazonas, Dept. Atabapo, Sector 

Norte de la Sierra Parima, O.Huber & M.Colchester 8399 (MYF, US). 83. COLOMBIA: Caquetá, 

Araracuara, J.Duivenvoorden 2656 (K). PERU: Loreto, Maynas, Río Nanay, 30 km SW of Iquitos, R.B.Foster 

4299 (F). 84. BRAZIL NORTH: Amapá, Mun. de Oiapoque, 109 km SSE of Oiapoque road to Calçoene, 

S.A.Mori et al. 17201 (MG, NY); Amazonas, Manaus, Reserva Florestal Ducke, C.A.Sothers 999 (INPA, K); 

Pará, Mun. de Oriximiná, Rio Trombetas, 23 km along road from Cachoeira Porteira, C.A. Cid Ferreira & 

C.D.A.Mota 1131 (INPA, NY); Rondônia, 15 km N of Ariquemes, M.Nee 34361 (K, NY). 

3. Hirtella vesiculosa Suess., Repert. Spec. Nov. Regni Veg. 42: 43 (1937) 

T: Colombia: Río Negro, San Felipe, P.von Luetzelburg 22245; holo: M. 


Tree to 8 m tall. Branches hirsute when young. Leaves with stipules filamentous, to 8 mm 

long, hispid, caducous; petioles hispid, enveloped by swollen ant-cavities; lamina coriaceous, 

oblong-lanceolate, 13.5–24 × 2.5–6 cm, acuminate at apex with acumen 5–10 mm long, 

subcordate at base, hirsute on venation beneath; midrib prominulous above; primary veins 

15–20 pairs, prominent beneath. Inflorescences elongate racemose panicles 8–16 cm long, 

branched slightly at base only; rachis and branches hirsute. Bracts and bracteoles linear, to 8 mm 

long, persistent, hirsute. Flowers 5–6 mm long. Receptacle campanulate, hispid on exterior, 

glabrous within except for deflexed hairs at throat; pedicels 6–7 mm long. Calyx lobes ovate, 

hispid on exterior, puberulous within near apex. Petals 5, exceeding calyx lobes, glabrous. 

Stamens 6; filaments far exceeding calyx lobes. Style hirsute at base only. Ovary inserted at 

mouth of receptacle, pilose. Fruit not seen. 

Colombia (San Felipe on the Río Negro); known only from the original collections. 83: CLM. 

No habitat information available. Map 138. 

83. COLOMBIA: Guainía, San Felipe, Río Negro, P.von Luetzelburg 22226 (M); Guainía, San Felipe, Río 

Negro, J.T.Baldwin 3200 (US).


4. Hirtella dorvalii Prance, Fl. Neotrop. Monogr. 9: 273 (1972) 

T: Brazil North: Roraima, Caracaraí, Rio Branco, A.Ducke 1330; holo: NY; iso: A, IAN, MG, R, RB, US. 



Shrub. Branches hirsute when young. Leaves with stipules filamentous, c. 6 mm long, hirsute, 

caducous, eglandular; petioles 2–4 mm long, eglandular, hispid, terete; lamina coriaceous, 

ovate to oblong-ovate, slightly bullate, 8.5–14 × 5.5–7 cm, bearing swollen ant-cavities at 

subcordate base, hirsute beneath, acuminate at apex with acumen 1–3 mm long, subcordate at 

base; midrib prominent on both surfaces, hirsute beneath; primary veins 9–12 pairs, 

prominent and hirsute beneath. Inflorescences panicles, 14–25 cm long; rachis and branches 

hispid-pilose. Bracts and bracteoles oblong, 2–7 mm long, persistent, with stipitate glands. 

Flowers 4.5–6 mm long. Receptacle campanulate, hispid on exterior, glabrous within except 

for deflexed hairs at throat. Calyx lobes acute, pilose on exterior, puberulous within. Petals 5, 

glabrous, purple, exceeding calyx lobes. Stamens 6 or 7, unilateral; filaments far exserted. 

Ovary pilose. Style hirsute at base only. Fruit not seen. 

Northern Brazil. 84: BZN (AM, RR). Low caatinga forest on sandy soils. Map 139. 

84. BRAZIL NORTH: Amazonas, Rio Cuiuini, P.Acevedo-Rodriguez 821 (INPA, K, US); Roraima, Caracaraí, 

J.M.Pires & P.F.Leite 14849 (IAN, INPA). 

5. Hirtella guainiae Spruce ex Hook.f., in C.F.P.Martius, Fl. Bras. 14(2): 31 (1867) 

T: Venezuela: Amazonas, Río Guainia, San Miguel, R.Spruce 3523; lecto: K, fide G.T.Prance, Fl. Neotrop. 

Monogr. 9: 275 (1972); isolecto: BM, BR, P, TCD, W. 

Illustration: J.D.Hooker, in C.F.P.Martius, loc.cit., t. 10. 


Shrub or small tree to 6 m tall. Branches hispid-hirsute when young. Leaves with stipules 

filamentous, to 2 cm long, hispid, subpersistent; petioles 5–9 mm long, hispid, enveloped by 

swollen ant-cavities; lamina thin-coriaceous, oblong, 14–23 × 5.5–9.5 cm, acuminate at apex 

with acumen 2–18 cm long, subcordate at base, hirsute on venation beneath; midrib 

prominulous above; primary veins 11–16 pairs, prominent and hirsute beneath. Inflorescences 

elongate racemes, 15–30 cm long; rachis hirsutulous pubescent. Bracts and bracteoles 

lanceolate, 4–6 mm long, with a broad usually glandular base. Flowers 5–6 mm long. 

Receptacle campanulate, hirsutulous-puberulous on exterior, glabrous within except for deflexed 

hairs at throat; pedicels 3–8 mm long. Calyx lobes oblong-ovate, hirsutulous on exterior, 

puberulous within. Petals 5, purple, exceeding calyx lobes. Stamens 4 or 5, unilateral; 

filaments far exceeding calyx lobes. Style sparsely hirsute up to 1/2 its length. Ovary inserted 

near mouth of receptacle. Fruit not seen. 

Western Amazonia. 82: VEN. 83: CLM, ECU, PER. 84: BZN (AC, AM, RO). Non-flooded 


82. VENEZUELA: Amazonas, Río Casiquiare, El Porvenir, M.Colella et al. 2191 (K, NY). 83. COLOMBIA: 

Amazonas, Araracuara, Río Caquetá, A.C.Londoño et al. 1618 (NY); Vaupés, Río Kananari, trib. of Río 

Apaporis, R.E.Schultes & I.Cabrera 14522 (BM, GH). PERU: Loreto, Prov. Maynas, Iquitos, Buena Suerte, 

R.Vásquez & N.Jaramillo 8406 (K, MO). 84. BRAZIL NORTH: Acre, Palmeira airport, C.D.A.Mota 246 

(INPA); Amazonas, mouth of Rio Marié, Rio Negro, M.T.Madison et al. 6303 (INPA, NY); Rondônia, Rio 

Madeira, 2 km S of Rio Abunã, G.T.Prance et al. 6008 (F, INPA, K, P, U, US). 

6. Hirtella duckei Huber, Bol. Mus. Paraense Hist. Nat. 6: 74 (1910) 

T: Brazil North: Amazonas, lower Rio Japurá, A.Ducke MG6756; holo: MG; iso: BM, P, RB, US. 

Hirtella formicaria Suess., Repert. Spec. Nov. Regni Veg. 39: 14 (1935). T: Brazil North: Amazonas, Mun. de 

Tefé, Paranagua, B.A.Krukoff 4557; holo: B, lost; lecto: K, designated here; iso: A, BM, F, LE, M, MICH, S, US. 


Shrub. Branches hispid when young, lenticellate. Leaves with stipules filamentous, to 1 mm 

long, hispid, persistent; petiole 5–8 mm long, sparsely pilose, with 2 swollen ant-cavities; 

109

110 


lamina oblong to elliptic, thin-coriaceous, (15–) 20–30 × 6–11 cm, acuminate at apex with 

acumen 5–20 mm long, cordate at base, hirsute on venation beneath; midrib prominulous 

above; primary veins 11–18 pairs, prominent and hirsute beneath. Inflorescences elongate 

racemes, 2–12 cm long; rachis hispid. Bracts and bracteoles 2–3 mm long, linear, hirsute, 

subpersistent, eglandular. Flowers 5–6 mm long. Receptacle campanulate, hispid on exterior, 

glabrous within except for deflexed hairs at throat; pedicels 4–6 mm long. Calyx lobes 

oblong, hispid on exterior, puberulous within. Petals 5, pale purple, exceeding calyx lobes. 

Stamens 5–8, unilateral; filaments far exceeding calyx lobes. Style hirsute at base only. 

Ovary pilose, inserted at or near mouth of receptacle. Fruit not seen. 

Guyana and C and W Amazonia. 82: GUY, VEN. 83: BOL, CLM, PER. 84: BZN (AC, AM, PA). 

Non-flooded forest. Map 141. 

82. GUYANA: Akarai Mts., between R. Mapuera and Shodikar Creek, A.C.Smith 2999 (NY, P, US). 

VENEZUELA: Amazonas, 5 km NW of San Carlos de Río Negro, R.Liesner 3970 (MO, NY). 83. BOLIVIA: 

Dept. Pando, Río Abunã, 3 km above Río Negro, G.T.Prance et al. 8521 (INPA, NY, P). COLOMBIA: 

Vaupés, Intend Amazonas, L.E.Mora & T. van der Hammen APA95 (COL). PERU: Loreto, Río Ucayali, 

Arboreto Jenaro Herrera, L.Bernardi 7-2 (G, NY). 84. BRAZIL NORTH: Acre, Mun. de Mâncio Lima, 7 km 

from town, C.A. Cid Ferreira et al. 10916 (INPA, K, NY); Amazonas, Lábrea, G.T.Prance et al. 13882 (INPA, 

F, K, NY, US); Pará, Km 120 Itaituba-Humaitá, P.Bamps 5378 (NY). 

7. Hirtella revillae Prance, Acta Amazon. 8: 587, 589 (1979) 

T: Peru: Loreto, Maynas, Río Nanay, 4 km from Mishana, A.Gentry, F.Ayala & J.Revilla 15807; holo: NY; 

iso: K, MO. 

Illustration: G.T.Prance, Acta Amazon. 8: 588 (1979). 


Tree 8 m tall. Branches hispid when young. Leaves with stipules linear, persistent, hispid; 

petioles 5–6 mm long, hispid, with 2 swollen ant-cavities; lamina oblong, chartaceousmembranous, 

19–22 × 8–10 cm, rounded at base, abruptly acuminate at apex with acumen 

7–10 mm long, hirsute on venation beneath, with sparse appressed hairs on upper surface; 

midrib prominent beneath, prominulous above, hirsute on both surfaces; primary veins 13–17 

pairs, prominent beneath, prominulous above. Inflorescences of axillary racemes 5–6 cm 

long; rachis light brown hispid; lower pedicels longer than upper ones giving a slightly 

corymbose appearance. Bracts and bracteoles linear, persistent, hispid. Flowers 8–10 mm 

long (excluding pedicels). Receptacle campanulate, light brown hispid-hirsute on exterior, 

glabrous within (even at throat) except around base of ovary; pedicels 8–15 mm long, 

hirsute. Calyx lobes 5, lanceolate, hispid on exterior. Petals 5, white, glabrous. Stamens 6; 

filaments far exceeding calyx lobes. Style hirsute on lower portion only. Ovary inserted at 

mouth of receptacle, glabrous around base. Fruit not seen. 

Peru; known only from the Río Nanay. 83: PER. Upland forest on white sand, poorly 

drained, swampy. Collected in flower Nov.–Mar. Map 142. 

83. PERU: Loreto, Iquitos, Puerto Almendras, F.Ayala 3072 (K, MO); Loreto, Mishana, Río Nanay, A.Gentry 

et al. 36438 (MO, NY), R.Vásquez & G.Criollo 1750 (K, MO); Loreto, road Nauta-Iquitos Km 5, R.Vásquez & 

N.Arevalo 8975 (K, MO); Loreto, Maynas, Río Nanay, between Iquitos and Santa Maria de Nanay, A.Gentry et al. 

20678 (NY), 31725 (MO, NY), 36438 (MO, NY). 

Hirtella L. sect. Hirtella 

Hirtella sect. Hirtella 

Type: Hirtella americana L. 

Tachibota Aubl., Hist. Pl. Guiane 1: 287, t. 112 (1775). T: Tachibota guianensis Aubl. (= Hirtella 

racemosa Lam. var. racemosa). 

Causea Scop., Intr. Hist. Nat., 210 (1777). T: not designated. 

Salmasia Schreb., Gen. Pl. 1: 201 (1789), nom. illeg., non Salmasia Bubani (1873). T: not designated.


Cosmibuena Ruiz & Pav., Fl. Peruv. Prodr. 10, t. 2 (1794), nom. rejic., non Cosmibuena Ruiz & Pav., Fl. 

Peruv. 3: 2 (1802), t. 198 (1789), nom. cons. T: not traced. 

Thelira Thouars, Gen. Nov. Madagasc. 21 (1806); Thelyra DC., Prodr. 2: 527 (1825), orth. var. T: not 

designated. 

Brya Vell., Fl. Flumin. 146 (1829) and 4: t. 1 (1831), nom. illeg., non Brya P. Browne (1756, Leguminosae). 

T: B. racemosa Vell. 

Sphenista Raf., Sylva Tellur. 90 (1838). T: Sphenista peruviana (Ruiz & Pavon) Raf. (= Hirtella triandra Sw. 

subsp. triandra). 

Zamzela Raf., Sylva Tellur. 90 (1838). T: Zamzela racemosa Raf. (= Hirtella triandra Sw. subsp. triandra). 

Leaf bases without swollen ant cavities. 

A section of 100 species in the neotropics and 2 in Africa and Madagascar. 

8. Hirtella macrosepala Sandwith, Bull. Misc. Inform. Kew 1939: 549 (1940) 

T: Guyana: Berbice River, Dukalikuru Creek, T.A.W.Davis 590 (F.D.2631); holo: K. 



Tree to 18 m tall. Branches glabrous when young. Leaves with stipules lanceolate, 2–5 mm 

long, glabrous when mature, eglandular, persistent; petioles 3–5 mm long, glabrous, terete, 

eglandular; lamina oblong, coriaceous, 6.5–17 × 3–8 cm, cuspidate at apex with acumen 2–6 mm 

long, rounded at base, glabrous beneath; primary veins 7–10 pairs, prominulous on both 

surfaces; midrib prominulous above, prominent beneath. Inflorescences axillary and terminal 

panicles 2.5–5 cm long, usually subcorymbose; rachis and branches glabrous or glabrescent. 

Bracts and bracteoles c. 2 mm long, ovate, glabrescent, persistent; apex with 1 or 2 sessile 

glands. Flowers 6–8 mm long. Receptacle campanulate, glabrous or glabrescent on exterior, 

glabrous within except at throat; pedicels 2–8 mm long. Calyx lobes rounded, glabrous or 

glabrescent on exterior, tomentellous within; margins eglandular. Petals 5, white, glabrous. 

Stamens 6 or 7, unilateral; filaments glabrous, connate at base, far exceeding calyx lobes. 

Style hirsute up to 1/2 its length. Ovary inserted near mouth of receptacle, villous. Fruit not 

seen. 

Eastern Venezuela and the Guianas. 82: GUY, SUR, VEN. Primary forest. Flowering Sept.–Apr. 

Map 143. 

82. FRENCH GUIANA: Mana, For. Serv. 7259 (NY, P). GUYANA: Mazaruni Station, CAP35 (FD5344) (F, 

NY, P, US). VENEZUELA: Delta Amacuro, E of Río Grande, ENE of El Palmar, L.M.Berti 375 (F, NY, SP, US); 

Bolívar, between El Dorado and Caserío del Km 88, C.Blanco 591 (NY). 

9. Hirtella ulei Pilg., Verh. Bot. Vereins Prov. Brandenburg 47: 148 (1905) 

T: Brazil North: Amazonas, Manaus, Ponta Negra, E.Ule 6150; holo: B, destroyed; lecto: HBG, fide 

G.T.Prance, Fl. Neotrop. Monogr. 9: 277 (1972); isolecto: MG. 


Shrub to 4 m tall. Branches glabrous when young, becoming lenticellate with age. Leaves with 

stipules 1–2 mm long, deltoid, intrapetiolar, puberulous, eglandular, persistent; petioles 3–7 mm 

long, puberulous, becoming glabrous with age, eglandular, terete; lamina coriaceous, ovateelliptic, 

6.5–13 × 4–7 cm, acuminate at apex with acumen 2–6 mm long, rounded to 

subcordate at base, glabrous beneath, with palisade glands frequent towards base; primary 

veins 8–12 pairs, prominulous on both surfaces; midrib prominulous above, glabrous and 

prominent beneath. Inflorescences terminal panicles 10–45 cm long; rachis and branches 

puberulous to glabrescent. Bracts and bracteoles 1.5–4 mm long, ovate, persistent, puberulous, 

with numerous stipitate glands. Flowers 5–7 mm long. Receptacle campanulate, puberulous on 

exterior, glabrous within except for reflexed hairs at throat; pedicels 1–2 mm long. Calyx 

lobes acute, with small stipitate glands on margins, glabrescent on exterior, pubescent within. 

Petals 4, pink, glabrous. Stamens 6–7, unilateral; filaments far exceeding calyx lobes. Style 

hirsute on lower portion up to 1/3 its length. Ovary inserted at mouth of receptacle, pilose. 

Fruit oblong; epicarp glabrous; mesocarp very thin, fleshy; endocarp thin, hard, bony. 

111

112 


Venezuela (upper Orinoco region) and Brazil (Rio Negro region). 82: VEN. 83: CLM. 

84: BZN (AM). Sandy margins and islands of rivers, and savanna margins. Flowering 

throughout the year. Map 144. 

82. VENEZUELA: Amazonas, Dept. Casiquiare, Yavita, G.Bunting et al. 3746 (U); San Carlos de Río 

Negro, H.L.Clark 7789 (MO, NY). 83. COLOMBIA: Vaupés, Mitú, J.Betancur et al. 2880 (COL, K, NY). 

84. BRAZIL NORTH: Amazonas, Serra Aracá, J.M. de S. Miralha 63 (INPA, NY); Amazonas, Manaus- 

Caracaraí Km 130, Igarapé Lajes, G.T.Prance et al. 21032 (INPA, NY); Amazonas, Rio Xie, Vila Inamui, 

N.T.Silva 3897 (IAN). 

10. Hirtella glabrata Pilg., Notizbl. Königl. Bot. Gart. Berlin 6: 138 (1914) 

T: Brazil North: Amazonas, Manaus, Pensador, E.Ule 8984; holo: B, destroyed; lecto: K, fide G.T.Prance, 

Fl. Neotrop. Monogr. 9: 277 (1972); isolecto: L, US. 


Shrub or small tree to 8 m tall. Branches tomentellous when young, soon becoming glabrous 

and lenticellate. Leaves with stipules to 2 mm long, deltoid, persistent, intrapetiolar, 

eglandular; petioles 4–6 mm long, tomentellous when young, terete, eglandular; lamina 

coriaceous, oblong-elliptic to elliptic, 6.5–13 × 4–6.2 cm, acute to acuminate at apex with 

acumen 4–5 mm long, rounded to subcordate at base; lower surface with few sparse 

appressed hairs, with palisade glands frequent on lower portion; primary veins 9–12 pairs, 

prominulous on both surfaces; midrib prominulous above, prominent and puberulous 

beneath. Inflorescences terminal panicles 10–25 cm long; rachis and branches velutinoustomentellous. 

Bracts and bracteoles 2.5–4.5 mm long, ovate, persistent, with numerous to 

few large stipitate glands, or rarely glands mostly sessile. Flowers 5–7 mm long. Receptacle 

campanulate, sparsely tomentellous on exterior, glabrous within except for deflexed hairs at 

throat; pedicels 0.5–2 mm long. Calyx lobes acute, eglandular, sparsely tomentellous on 

exterior, grey-pubescent within. Petals 5, glabrous. Stamens 6 or 7, unilateral with short 

staminodes opposite them; filaments far exceeding calyx lobes. Style hirsute for 1/2 its length. 

Ovary inserted at mouth of receptacle, tomentellous. Fruit ellipsoid; epicarp smooth, 

glabrous; mesocarp thin, fleshy; endocarp thin, hard, bony, hirsute within. 

Venezuela and C Brazilian Amazonia. 82: VEN. 84: BZN (AM, PA). Open forest, savannas, 

and secondary forest. Flowering Apr.–Sept. Map 145. 

82. VENEZUELA: Amazonas, Dept. Atures, Caño Cabeza de Manteco, trib. of Río Autana, F.Guanchéz & 

E.Melgueiro 3454 (MYF, NY); Bolívar, Mun. de Cedeño, Picachos de Sabana Nueva, S.Elcoro 246 (K, PORT). 

84. BRAZIL NORTH: Amazonas, Rio dos Pombos, trib. of Rio Yuma, 73 km E of Rio Aripuanã, 

C.E.Calderón et al. 2634 (INPA, NY, US); Amazonas, Mun. de Borba, Rio Madeira, 8 km from Borba, C.A. 

Cid Ferreira 3869 (INPA, NY); Amazonas, Manaus-Manacapuru, 5 km from Cacau Pirêra, G.T.Prance et al. 

23508 (INPA, NY). 

11. Hirtella confertiflora Prance, Brittonia 33: 354 (1981) 

T: Venezuela: Amazonas, Río Coro-coro, J.A.Steyermark, M.G.Casanova & P.Redmond 117921; holo: NY; 

iso: MO, VEN. 

Illustration: G.T. Prance, Brittonia 33: 355 (1981). 


Tree 4 m tall. Branches densely ferrugineous tomentose when young. Leaves with stipules 

lanceolate, to 5 mm long, caducous, eglandular; petioles 1–3 mm long, terete, tomentellous; 

lamina oblong-elliptic, coriaceous, 10.5–12 × 4.8–5.7 cm, cordate at base, shortly and 

abruptly acuminate at apex with acumen 1–3 mm long, glabrous and shiny above, with sparse 

appressed hirsute pubescence on lower surface; midrib prominulous above, prominent and 

appressed beneath; primary veins 10–13 pairs, prominulous above, prominent beneath; 

secondary venation prominulous on both surfaces. Inflorescences of terminal and axillary 

panicles with central rachis and short densely clustered lateral branches bearing 2–4 flowers, 

12–15 cm long; rachis and branches tomentellous. Bracts and bracteoles 1–3 mm long, 

triangular to lanceolate, persistent, with numerous stipitate tack-like glands. Flowers 5–7 mm 

long. Receptacle campanulate, tomentellous externally, glabrous within except for reflexed 

hairs at throat; pedicels 3–7 mm long. Calyx lobes 5, acute, eglandular, tomentellous


Figure 68. Hirtella confertiflora. A, habit; B, leaf lower surface; C, flower and bracteole; 

D, petal; E, ½ flower (A–E, J.A.Steyermark et al. 117921). Scale bars: A = 3 cm; B = 1 cm; 

C–E = 2 mm. Drawn by Bobbi Angell. Previously published in G.T.Prance, Brittonia 33: 355 

(1981). 

113

114 


externally, puberulous within. Petals 5. Stamens 6, unilateral; filaments far exceeding calyx 

lobes. Style sparsely hirsute on lower ¼. Ovary inserted at mouth of receptacle, pilose. Fruit 

not seen. Fig. 68. 

Venezuela; known only from the type collection. 82: VEN. Gallery forest bordering tree 

savanna. Map 146. 

12. Hirtella carbonaria Little, J. Wash. Acad. Sci. 38: 88 (1948) 

T: Ecuador: Esmeraldas, San Lorenzo, near Río Nadadero, E.L.Little 6275; holo: US; iso: A, F, K, MAD, NY. 


Tree to 10 m tall. Branches sparsely hirsutulous when young, soon becoming glabrous and 

lenticellate. Leaves with stipules linear, puberulous, eglandular, subpersistent, 2–4 mm long; 

lamina coriaceous, elliptic to ovate-elliptic, 4.5–9.5 × 2.5–4.5 cm, acuminate at apex with 

acumen 4–10 mm long, rounded at base, glabrous beneath except for a few appressed hairs; 

primary veins 6–9 pairs, prominulous on both surfaces; midrib prominent on both surfaces, 

sparsely hirsutulous beneath. Inflorescences terminal and axillary panicles, 3–9 cm long; rachis 

and branches glabrescent, with a few stiff appressed hairs. Bracts and bracteoles 1–3 mm 

long, ovate, hirsutulous, persistent; solitary glands on a long stalk, 1–2 mm long, arising 

from the junction of the pedicel and branch or from pedicel. Flowers 4–5 mm long. 

Receptacle campanulate, glabrous on exterior, grey-pubescent within. Petals 5, white, glabrous. 

Stamens 3, unilateral, with 3 short filamentous staminodes opposite them; filaments far 

exceeding calyx lobes. Style hirsute on lower portion. Ovary inserted near mouth of 

receptacle, tomentose. Fruit ellipsoid; epicarp smooth, glabrous when mature; mesocarp very 

thin, fleshy; endocarp thin, bony, densely hirsutulous within. 

Pacific coast of Ecuador and southern Colombia. 83: CLM, ECU. Coastal thickets and 

margins of mangrove swamps in areas periodically flooded by sea water. Flowering 

throughout the year. Map 147. 

83. COLOMBIA: Chocó, Río San Juan, below Docordó, E.Forero et al. 4410 (COL, MO, NY); Nariño, 

Tumaco, Estera de Aguaclara, J.M.Acevedo & Pinilla C65 (COL); Nariño, Mun. de Tumaco, Río Mira, H.León 

et al. 1561 (NY, U); Valle, El Forye near Buenaventura, E.P.Killip & J.Cuatrecasas 38965 (COL, F, GH, NY, 

S, US). ECUADOR: Esmeraldas, San Lorenzo, Río San Antonio, El Chorro, B.Sparre 18306 (S). 

The wood is used for making charcoal. 

13. Hirtella araguariensis Prance, Fl. Neotrop. Monogr. 9: 278 (1972) 

T: Brazil North: Amapá, Rio Araguari, J.M.Pires, W.A.Rodrigues & G.C.Irvine 51537; holo: NY; iso: IAN, 

INPA, MG, US. 



Tree to 8 m tall. Branches hispid when young, soon glabrous. Leaves with stipules linear, to 

8 mm long, pubescent, eglandular, persistent; petioles 2–3 mm long, terete, eglandular, hispid 

when young; lamina subcoriaceous, oblong-lanceolate, 10–17 × 3.5–5.3 cm, acuminate at 

apex with acumen 5–12 mm long, subcordate at base; primary vein prominent on both 

surfaces; secondary veins 10–13 pairs, slightly prominent on both surfaces. Inflorescences of 

panicles with long central rachis and small clusters of 1–3 flowers giving a racemose 

appearance, 10–25 cm long; rachis and branches light-brown pilose-tomentose. Bracts and 

bracteoles oblong, 3–6 mm long, acute, tomentellous, persistent; margins eglandular or with 

few sessile glands; pedicels 0.5–1 mm long. Flowers 5–7 mm long. Receptacle campanulate, 

tomentellous on exterior, glabrous within at base. Calyx lobes acute, tomentose outside, 

puberulous within; margins eglandular. Petals 5, white, glabrous. Stamens 5, unilateral, with 

base of filaments hirsute, far exceeding calyx lobes. Style hirsute. Fruit not seen. 

Brazil, from north of Manaus eastward to Amapá and French Guiana. 82: FRG. 84: BZN 

(AM, AP). Upland forest. Flowering in October. Map 148. 

82. FRENCH GUIANA: Basin of R. Approuague, Crique Nouragues, J.J. de Granville 10377 (CAY, NY). 

84. BRAZIL NORTH: Amapá, Porto Grande, Vila Nova, N.A.Rosa 1052 (MG, NY); Amazonas, Mun. de


Presidente Figueiredo, Rio Uatumã, I.L.Amaral et al. 1757 (INPA); Amazonas, Rio Uatumã, Mun. de Itapiranga, 

Igarapé Catitu, C.A. Cid Ferreira et al. 640 (INPA, NY); Amazonas, Mun. de Presidente Figueiredo, Rio Uatumã, 

Nazaré, near mouth of Rio Pitinga, C.A. Cid Ferreira et al. 6946 (INPA, K, NY). 

14. Hirtella barnebyi Prance, Brittonia 33: 352 (1981) 

T: Brazil North: Rondônia, Point 22 of RADAM SC-20-XD, margin of Rio Preto, near rapids, M. dos R. 

Cordeiro 703; holo: INPA57032; iso: IAN, INPA54595. 

Illustration: G.T.Prance, loc.cit. (1981). 


Shrub 2 m tall. Branches sparsely hirsutulous when young, soon becoming glabrous. Leaves 

with stipules minute, persistent, c. 1 mm long, tomentellous; petioles 2–2.5 mm long, sparsely 

tomentellous, terete, eglandular; lamina oblong, chartaceous, 6–9.5 × 2.5–3.8 cm, rounded at 

base, acuminate at apex with acumen 9–14 mm long, glabrous beneath except for a few stiff 

appressed hairs on venation; primary veins 8–11 pairs, prominulous on both surfaces; midrib 

prominulous above, prominent beneath. Inflorescences 8–12 cm long, of little-branched 

panicles with a central rachis and small lateral branches bearing 2–3 flowers only; rachis and 

branches very sparsely hirsutulous. Bracts and bracteoles 1–1.5 mm long, lanceolate, 

persistent, with sessile glands on some; caducous bracteoles with large terminal glands present 

only in bud. Flowers 6–7 mm long. Receptacle campanulate, with a few sparse appressed 

hairs on exterior, glabrous within except at throat; pedicels 5–7 mm long, sparsely appressed 

hirsute. Calyx lobes 5, reflexed in open flowers, acute, hirsutulous externally, densely greytomentellous 

within; margins eglandular. Petals 5, glabrous. Stamens 6, unilateral; filaments 

glabrous, or with a few hairs on lower 10 mm, free to base, far exceeding lobes. Style hirsute 

up to 1/3 of length. Ovary inserted near mouth of receptacle, villous. Fruit not seen. Fig. 69. 

Brazil; known only from the type. 84: BZN (RO). Near rapids. Map 149. 

15. Hirtella margae Prance, Proc. Kon. Ned. Akad. Wetensch. C. 89: 111 (1986) 

T: Suriname: Brokopondo Distr., Brownsberg Nature Park, L.Roberts LBB16303; holo: NY. 

Illustration: G.T.Prance, Proc. Kon. Ned. Akad. Wetensch. C. 89: 112 (1986). 


Tree to 3 m tall. Branches glabrous when young, becoming lenticellate with age. Leaves with 

stipules linear-lanceolate, persistent, 7–9 mm long, eglandular, appressed pubescent; petioles 

8–10 mm long, rugulose, with appressed pubescence of stiff long hairs, terete, eglandular; 

lamina oblong to oblong-lanceolate, coriaceous, 21–27 × 6–9 cm, rounded to subcordate at 

base, acuminate at apex with acumen 1–5.5 cm long, glabrous beneath except for a few stiff 

appressed hairs on venation; midrib prominulous and glabrous above, prominent and with 

sparse appressed pubescence beneath; primary veins 15–18 pairs, prominulous above, 

prominent beneath. Inflorescences of little-branched panicles with a central rachis and small 

groups of flowers on short branches and solitary flowers; rachis dark brown-tomentellous. 

Bracts and bracteoles 3–6 mm long, lanceolate, persistent, appressed tomentellous, with few 

sessile glands on margins of most bracteoles. Flowers 5–7 mm long, solitary or in small 

clusters on central rachis. Receptacle campanulate, appressed hirsutulous on exterior, 

glabrous within except for pilose deflexed hairs around throat. Calyx lobes 5, acute, 

appressed hirsutulous on exterior, grey-tomentellous within, eglandular. Petals 5, pale 

purple, glabrous. Stamens 6, unilateral; filaments far exceeding calyx lobes. Style pilose on 

lower portion, glabrous above. Ovary inserted near mouth of receptacle, pilose. Fruit ovoid, 

c. 3 mm long, 1.8–2.5 cm broad, appressed-hirsutulous on exterior; mesocarp thin, fleshy; 

endocarp thin, hard, bony, densely hirsute within. 

Guianas. 82: FRG, GUY, SUR. Primary forest. Map 150. 

82. FRENCH GUIANA: Piste de Saint Elie, M.F.Prévost & D.Sabatier 2994 (CAY, K). GUYANA: Puruni, 

J.Boyan 65 (NY). SURINAME: Bakhuis Mts., W bank of Coppename R., P.A.Florschütz & P.J.M.Maas 2810 (NY). 

115

116 


Figure 69. Hirtella barnebyi. A, habit; B, leaf lower surface; C, flower bud; D, ½ flower; 

E, petal (A–E, M.R.Cordeiro 703). Scale bars: A = 3 cm; B = 1 cm; C–E = 2 mm. Drawn by 

Bobbi Angell. Previously published in G.T.Prance, Brittonia 33: 352 (1981).


16. Hirtella liesneri Prance, Fl. Neotrop. Monogr. 9S: 89 (1989) 

T: Venezuela: Táchira, Río San Buena, 10 km W of La Fundación, R.Liesner, A.González & R.F.Smith 9633; 

holo: NY; iso: MO, VEN. 

Hirtella racemosa var. glandipedicellata Prance, Fl. Neotrop. Monogr. 9: 330 (1972). T: Colombia: Caquetá, 

Morelia, A. von Sneidern A1190; holo: S; iso: BM, COL, NY. 



Tree 7 m tall. Branches sparsely hispid when young, becoming glabrous with age. Leaves with 

stipules linear, persistent, c. 3 mm long, eglandular; petioles c. 2 mm long, terete, sparsely 

hispid when young; lamina oblong, chartaceous, 10–15 × 3.5–5 cm, subcordate at base, 

acuminate at apex with acumen 10–15 mm long; lower surface with sparse appressed hairs, 

especially on venation; midrib prominulous above, prominent and appressed pubescent beneath; 

primary veins 12–16 pairs, prominulous on both surfaces. Inflorescences of terminal and 

axillary panicles, with a long central rachis and very short 3–5 flowered lateral branches 

giving a racemose appearance, 20–22 cm long; rachis puberulous with lower part sparsely 

hispid. Bracteoles triangular, membranous; apex terminating in a stalked gland. Flowers 6–7 mm 

long. Receptacle campanulate, sparsely hispid on exterior, glabrous within except for ring of 

deflexed hairs at throat; pedicels 2–4 mm long. Calyx lobes 5, acute, eglandular, sparsely 

hirsute on exterior, puberulous within. Petals 5, glabrous. Stamens 6, unilateral; filaments far 

exceeding calyx lobes. Style hirsute on lower 1/3. Ovary inserted at mouth of receptacle, 

pilose. Fruit when young ellipsoid; exocarp sparsely appressed pubescent, becoming glabrous 

with age. 

Venezuela and Colombia. 82: VEN. 83: CLM. Primary forested area on sandy soil, 700–1000 m. 

Map 151. 

82. VENEZUELA: Bolívar, SE of Santa Elena de Uairén, A.L.Bernardi 6760 (NY). 83. COLOMBIA: Guaviare, 

Mun. de San José del Guaviare, km 3 San José-Puerto Arturo, R.López et al. 973 (COAH, K). 

17. Hirtella cordifolia Prance, Fl. Neotrop. Monogr. 9: 279 (1972) 

T: Venezuela: Amazonas, Río Hechimoni, Sabana Hechimoni, J.J.Wurdack & L.S.Adderley 436211; holo: NY. 



Shrub, small. Branches puberulous when young, glabrescent, lenticellate. Leaves with stipules 

linear, to 3 mm long, sparsely pubescent, eglandular, persistent; petioles c. 1 mm long, terete, 

eglandular, hirsute when young; lamina cordate-ovate, coriaceous, 3.5–5.5 × 1.8–2.9 cm, 

acuminate at apex with acumen 2–7 mm long, cordate at base, glabrous on both surfaces 

except on venation beneath; midrib prominent and hirsute above; primary veins 7–9 pairs 

slightly prominent on both surfaces. Inflorescences of little-branched panicles or racemes, 

2.5–5.5 cm long; rachis and branches hirsute-puberulous. Bracts and bracteoles 1.5–3 mm 

long, acute, persistent, puberulous, with stipitate glands on margins. Flowers c. 6 mm long. 

Receptacle campanulate, hirsute on exterior, glabrous within except for hairs around throat; 

pedicels 3–6 mm long. Calyx lobes acute, sparsely hirsute on exterior, grey-pubescent 

within. Petals 5, glabrous, pink. Stamens 4 or 5, unilateral, with toothed staminodes opposite; 

filaments glabrous, far exserted, connate at base. Ovary inserted midway up receptacle, 

sparsely pilose. Style hirsute. Fruit ellipsoid; exocarp smooth, glabrous; mesocarp thin, 

fleshy; endocarp thin, hard, bony, hirsute within. 

Venezuela (Río Hechimoni). 82: VEN. Savannas. Flowering in July. Map 152. 

82. VENEZUELA: Amazonas, Río Hechimoni, Sabana Hechimoni, B.Maguire, J.J.Wurdack & Bunting 37665 (NY). 

18. Hirtella insignis Briq. ex Prance, Fl. Neotrop. Monogr. 9: 279 (1972) 

T: Brazil Northeast: Bahia, without precise locality, J.S.Blanchet 1508; holo: BM; iso: G. 



117

118 


Tree to 15 m tall. Branches sparsely hispid when young, soon becoming glabrous and 

lenticellate. Leaves with stipules linear, small; petioles 3–5 mm long, terete, eglandular, 

glabrous when mature; lamina oblong, coriaceous 7–17 × 3.6–6 cm, acuminate at apex with 

acumen 2–7 mm long, rounded or subcuneate at base, with few sparse appressed hairs 

beneath, otherwise glabrous, with glands towards base; midrib slightly prominent above, 

prominent and sparsely hispid beneath; primary veins 10–15 pairs, slightly prominent on 

both surfaces. Inflorescences of terminal much-branched panicles 6–7 cm long; rachis and 

branches glabrous. Bracts and bracteoles oblong, acute, glabrescent, persistent; apex with a 

long stipitate gland. Flowers 5–6 mm long. Receptacle campanulate, almost glabrous on 

exterior but with few appressed hairs, glabrous towards base within; pedicels 7–16 mm long, 

eglandular. Calyx lobes rounded or acute, glabrous on exterior, brown-pubescent within; 

margins ciliate. Petals 5, glabrous. Stamens 5, unilateral; filaments glabrous, far exserted. 

Ovary pilose. Style hirsute at base. Fruit not seen. 

Eastern Brazil. 84: BZE (BA), BZL (ES). Atlantic coastal forests. Collected in flower in 

February. Map 153. 

84. BRAZIL NORTHEAST: Bahia, Mun. de Una, 4 km N of Comandatuba, W.W.Thomas et al. 6024 (K, NY); 

Bahia, Valença, Km 10 Valença–Guaibim, T.S. dos Santos 2898 (CEPEC, NY); Bahia, Ilhéus, road Olivença 

to Maruim, 5.4 km SW of Olivença, W.W.Thomas et al. 8923 (CEPEC, K, NY). BRAZIL SOUTHEAST: 

Espírito Santo, Res. Biol. de Comboio, G.L.Farias 270 (K); Espírito Santo, Mun. de Itapemirim, 3 km N of 

Marataízes, J.R.Pirani et al. 3585 (K, NY, SP); Espírito Santo, R. F. Linhares, J.Spada 210 (RB). 

19. Hirtella tocantina Ducke, Arch. Jard. Bot. Rio de Janeiro 3: 266 (1922) 

T: Brazil North: Pará, Rio Tocantins, Alcobaça, A.Ducke MG16274; holo: MG; iso: P, RB, US. 


Tree, medium-sized. Branches sparsely pubescent when young, soon glabrous and lenticellate. 

Leaves with stipules linear, to 5 mm long, glabrescent, subpersistent, eglandular; petioles 

4–5 mm long, glabrescent, terete, eglandular; lamina coriaceous, oblong 10–15 × 3–5.5 cm, 

acuminate at apex with acumen 8–12 mm long, cuneate at base; lower surface with several 

palisade glands towards base, glabrous; primary veins 9–12 pairs, prominulous on both 

surfaces; midrib prominulous and glabrous on both surfaces. Inflorescences terminal and 

axillary panicles 7–12 cm long; rachis and branches with sparse appressed pubescence. 

Bracts and bracteoles oblong-lanceolate, 2.5–3.5 mm long, membranous, persistent, sparsely 

puberulous; margins with both sessile and short-stipitate glands. Flowers 5–6 mm long. 

Receptacle campanulate, glabrescent on exterior, glabrous within except for hairs at throat; 

pedicels 1–3 mm long. Calyx lobes acute, glabrescent on exterior, grey-pubescent within; 

margins glandular. Petals 5, white, glabrous. Stamens 3, unilateral, with 3 short filamentous 

staminodes opposite; filaments far exceeding calyx lobes. Style hirsute up to 1/3 its length. 

Ovary inserted at mouth of receptacle, pilose. Fruit not seen. 

Brazil (collected from the banks of the Rios Tocantins, Tapajós and Madeira) and Peru. 

83: BOL, PER. 84: BZN (AC, AM, PA). Non-flooded forest. Map 154. 

83. PERU: Huánuco, Dept. Llullapichis, Prov. Puerto Inca, B.K.Saldaña 311 (G, K). 84. BRAZIL NORTH: 

Acre, 33 km NNE of Rio Branco on road to Porto Acre, D.C.Daly et al. 6103 (K, NY); Acre, Mun. de 

Brasiléia, Seringal Porangaba, D.C.Daly 6804 (K, NY); Amazonas, Mun de Humaitá, 150 km from Humaitá on 

Estrada do Estanho, A.Henderson et al. 278 (INPA, NY); Pará, Rio Tapajós, near Pimental, A.Ducke RB18236 

(P, RB, S, US). 

20. Hirtella piresii Prance, Fl. Neotrop. Monogr. 9: 283 (1972) 

T: Brazil North: Rondônia, Porto Velho, Estrada de Rodagem, km 8, G.A.Black, E.Cordeiro & J.Francisco 

52-14653; holo: NY; iso: IAN. 



Tree, small to medium-sized. Branches glabrescent when young. Leaves with stipules linear, 

1 mm long, eglandular, caducous; petioles 1–2 mm long, terete, eglandular, tomentellous 

when young; lamina coriaceous, oblong, 3.5–9 × 1.4–3.2 cm, glabrous beneath, acuminate at


apex with acumen 2–10 mm long, cuneate at base; primary vein prominulous above, prominent 

beneath, glabrescent; secondary veins 8–11 pairs, slightly prominent on both surfaces. 

Inflorescences terminal and axillary panicles, 3–11 cm long; rachis and branches glabrescent. 

Bracts and bracteoles oblong to lanceolate, 2–5.5 mm long, puberulous or glabrescent, 

persistent; bracteoles with a single reflexed apical gland; pedicels 1.5–3 mm long. Flowers 

4–5 mm long; receptacle campanulate, glabrous on exterior, glabrous within towards base. 

Calyx lobes acute, glabrous on exterior, glabrescent within; margins ciliate, eglandular. 

Petals 5, glabrous. Stamens 3, unilateral, with dentiform staminodes opposite; filaments 

glabrous, far exserted; ovary pilose. Style hirsute. Fruit pyriform when young; epicarp 

smooth, glabrous; mesocarp thin, fleshy; endocarp hard, thin, bony, hirsute within. 

Brazilian Amazonia, Guianas and Bolivia. 82: FRG, SUR. 83: BOL. 84: BZC (MT), BZN 

(AC, AM, PA, RO). Non-flooded forest. Flowering May–Sept. Map 155. 

82. FRENCH GUIANA: R. Arataye, Saut Pararé, J.F.Villiers 2729 (CAY). SURINAME: Nickerie R., 3 km NE 

of Kamisa Falls, LBB11049 (US). 83. BOLIVIA: Dept. Pando, N Suarez Prov., 14 km towards Campo Ana, 

B.Rodriguez 1419 (NY). 84. BRAZIL NORTH: Acre, Mun. de Brasiléia, Km 26 road to Assis Brasil, 

C.A. Cid Ferreira et al. 10203 (INPA, K, NY); Amazonas, N of Manaus, Distrito Agropecuário, Reserva 1501, 

B.Boom et al. 8646 (INPA, K, NY); Pará, Rio Tocantins, Jacundá, M.G.Silva & R.Bahia 3128 (MG, NY); 

Rondônia, Mun. de Porto Velho, Vila Mutumparaná, C.A. Cid Ferreira 8851 (INPA, K, NY). BRAZIL WEST- 

CENTRAL: Mato Grosso, Mun. de Sinop, 142 km from Peixoto de Azevedo, C.A. Cid Ferreira 6341 (INPA, NY). 

21. Hirtella davisii Sandwith, Bull. Misc. Inform. Kew 1935: 125 (1935) 

T: Guyana: Cuyuni R., Oko Creek, T.A.W.Davis 352 (FD2345); holo: K; iso: FHO. 

Illustration: G.T.Prance, Fl. Guianas 85: 33 (1986). 


Tree to 30 m tall. Branches sparsely pilose when young, soon becoming glabrous and lenticellate. 

Leaves with stipules linear, 3–4 mm long, subpersistent, eglandular, glabrous; petioles 3–4 mm 

long, puberulous to glabrous, terete, eglandular; lamina oblong to oblong-elliptic, coriaceous, 

6–13.5 × 2.3–5 cm, glabrous beneath, acuminate at apex with acumen 5–10 mm long, 

subcuneate at base, with several palisade glands towards base beneath; primary veins 9 or 10 

pairs, prominulous on both surfaces; midrib prominulous above, prominent and with sparsely 

appressed pubescence beneath. Inflorescences terminal and axillary panicles 5–14 cm long; 

rachis and branches sparsely pilose. Bracts and bracteoles 1–3.5 mm long, ovate to lanceolate, 

puberulous, persistent, coriaceous, with numerous stipitate glands. Flowers 5–6 mm long. 

Receptacle campanulate, puberulous on exterior, glabrous within except at throat; pedicels 

0.5–2 mm long. Calyx lobes rounded to acute, puberulous on exterior, grey-pubescent 

within; margins eglandular. Petals 5, white, glabrous. Stamens 3, unilateral; filaments far 

exceeding calyx lobes. Style hirsute up to 1/3 its length. Ovary inserted at mouth of 

receptacle, pilose. Fruit ellipsoid; epicarp glabrous, smooth; mesocarp thin, fleshy; endocarp 

thin, bony, densely pilose-lanate within. 

Nicaragua and Costa Rica, Eastern Venezuela, Guianas, and eastern Amazonian Brazil. 80: COS, 

NIC. 82: FRG, GUY, VEN. 84: BZE (MA), BZN (PA). Primary forest on high ground. 

Flowering Mar.–May in the Guianas, Sept.–Oct. in Brazil. Map 156. 

80. NICARAGUA: Río San Juan, mouth of Río Santa Cruz, tributary of Río San Juan, P.P.Moreno 25541 & 

25614 (MO). 82. FRENCH GUIANA: Saül, Grand Boeuf Mort trail, S.A.Mori & C.Gracie 18929 (K, NY); 

Saut Pararé, Arataye, D.Sabatier 939 (CAY). GUYANA: Kamuni Creek, Groete Creek, Essequibo R., B.Maguire 

& D.B.Fanshawe 22912 (A, F, K, NY, US). VENEZUELA: Bolívar, Magdalena, El Palmar, 15 km NE of Río 

Grande, A.L.Bernardi 7111 (NY); Delta Amacuro, E of Río Grande, ENE of El Palmar, A.Zabala 80 (VEN). 

84. BRAZIL NORTHEAST: Maranhão, Fazenda São Francisco, 11 km N of Km 337 of BR 316, D.C.Daly et 

al. 259 (MG, NY). 

22. Hirtella subglanduligera Pilg., Notizbl. Königl. Bot. Gart. Berlin 6: 140 (1914) 

T: Brazil North: Acre, Seringal Auristella, E.Ule 9414; holo: B, destroyed; lecto: MG14263, fide G.T.Prance, 

Fl. Neotrop. Monogr. 9: 284 (1972); isolecto: L, US. 


119

120 


Tree to 25 m tall. Branches sparsely pilose when young, becoming glabrous and lenticellate 

with age. Leaves with stipules linear, to 6 mm long, glabrous, subpersistent, eglandular; petioles 

1–2 mm long, glabrous, terete, eglandular; lamina ovate-elliptic to elliptic, subcoriaceous, 

6–9 × 2.5–4 cm, acuminate at apex with acumen 4–5 mm long, subcuneate at base, glabrous 

beneath or bearing a few stiff appressed hairs, with a few palisade glands towards base; 

primary veins 9 or 10 pairs, prominulous on both surfaces; midrib prominulous above, 

prominent and sparsely pubescent beneath. Inflorescences terminal and axillary panicles 

8–22 mm long; rachis and branches sparsely pilose. Bracts and bracteoles oblong-lanceolate, 

membranous, puberulous, persistent; margins bearing stipitate glands. Flowers c. 7 mm long. 

Receptacle campanulate, puberulous on exterior, glabrous within except for throat; pedicels 

1.5–3 mm long. Calyx lobes rounded to acute, puberulous on both surfaces; margins sparsely 

glandular. Petals 5, white, glabrous. Stamens 4, unilateral, with 4 short filamentous 

staminodes opposite; filaments glabrous, far exceeding calyx lobes. Style hirsute up to 1/3 its 

length. Ovary inserted at mouth of receptacle, pilose. Fruit not seen. 

Brazil (upper Rio Acre); known only from the type collection. 84: BZN (AC). Terra firme 

forest. Flowering in July. Map 157. 

23. Hirtella ciliata Mart. & Zucc., Abh. Math.-Phys. Cl. Königl. Bayer. Akad. Wiss. 

1: 378 (1832) 

T: Brazil Northeast: Bahia, 'in carasco et tabuleiro', C.F.P.Martius 1722; holo: K; iso: BR. 

Hirtella rubra Benth., J. Bot. (Hooker) 2: 217 (1840). T: Guyana: Essequibo, Rob.Schomburgk 113; holo: 

K; iso: BM, CGE, GH, LE, NY, OXF, P, W. 

Hirtella rotundata Pilg., Notizbl. Königl. Bot. Gart. Berlin 6: 140 (1914). T: Brazil Northeast: Ceará, 

Restinga Bemfica, near Fortaleza, E.Ule 9043; holo: B, destroyed; iso: K. 

Grangeria brasiliensis Hoffmanns. ex Mart. & Zucc., Abh. Math.-Phys. Cl. Königl. Bayer. Akad. Wiss. 1: 378 

(1832), nom. nud., in synon. 


Tree to 12 m tall, with thick corky bark on trunk. Branches tomentellous when young, 

becoming glabrous with age. Leaves with stipules c. 1 mm long, oblong, tomentellous, 

eglandular, persistent; petioles 0.5–2 mm long, tomentellous, terete, eglandular; lamina 

orbicular to ovate-elliptic, coriaceous, 3–6.5 × 1.5–3.6 cm, retuse to acuminate at apex, most 

frequently mucronate, subcordate to rounded at base, hirsute beneath or with sparse 

appressed pubescence; palisade glands present; primary veins 6–9 pairs, plane above, 

prominulous beneath; midrib prominent on both surfaces. Inflorescences terminal and 

subterminal panicles 5–22 cm long; rachis and branches sparsely tomentellous. Bracts and 

bracteoles 1–3 mm long, ovate to oblong, pubescent, persistent, with numerous stipitate 

glands. Flowers 3.5–5 mm long. Receptacle campanulate, puberulous on exterior, pubescent 

to base within; pedicels 2–6 mm long. Calyx lobes acute, puberulous on exterior, greypubescent 

within; margins glandular. Petals 5, glabrous, greenish-white. Stamens 6–9, 

unilateral with a few short filamentous staminodes opposite; filaments slightly exceeding 

calyx lobes. Style hirsute at base only. Ovary inserted at mouth of receptacle, pilose. Fruit 

round to ellipsoid; epicarp smooth and glabrous when mature; mesocarp thin, fleshy; 

endocarp thin, bony, hirsute within. Fig. 70E. 

Guianas and Amazonia, planalto of C Brazil, and NE Brazil. 82: FRG, GUY. 84: BZC (GO, 

TO), BZE (BA, CE, MA, SE, PE, PI), BZL (MG), BZN (AP, PA). Savannas, cerrados and 

taboleiros. Map 158. 

82. FRENCH GUIANA: 2 km from km 86 of road NI beyond Sinnamary, F.Billiet & B.Jadin 1458 (BM, BR, 

CAY). GUYANA: Potaro-Siparuni Region, 2–3 km N of Kato, L.J.Gillespie 853 (K, US). 84. BRAZIL 

NORTH: Pará, Mun. de Óbidos, Campos de Ariramba, Igarapé Mutum, C.Farney 2053 (INPA, K, NY); 

Tocantins, fazenda Bragança, near Darcinópolis, J.A.Ratter et al. 6776 (E, K). BRAZIL NORTHEAST: Bahia, 

Mun. de Tucano, km 7–10 Tucano-Ribeira do Pombal, A.M. de Carvalho et al. 3917 (CEPEC, K, NY); Ceará, 

Tabuleiro-Itaperi, Parangaba, A.Ducke HPB1550 (HPB, NY); Piauí, 17 km N of Fortaleza dos Nogueiras road 

to Serra Negra, J.A.Ratter et al. 7799 (E, K); Sergipe, Mun. de Estância, km 11 Estância-Abais, A.Mattos Silva 

et al. 3004 (CEPEC, K). BRAZIL SOUTHEAST: Minas Gerais, Mun. de Formoso, Parque Nacional Grande 

Sertão Veredas, T.S.Filgueiras et al. 3577 (IBGE, K). BRAZIL WEST-CENTRAL: Goiás, Guarani de Goiás, 

road BR020, G.Hatschbach 50541 (K, MBM).


Figure 70. Hirtella. A–C, H. glandulosa. A, habit; B, flower and glands; C, ½ flower 

(A–C, H.S.Irwin et al. 16435, 16745). D, H. martiana. Flower bud (H.S.Irwin et al. 18039). 

E, H. ciliata. Fruit (G.T.Prance & N.T.Silva 58456). Scale bars: A = 5 cm; B, D = 1 cm; 

C = 2 mm; E = 2 cm. Drawn by Janet Dyer. Previously published in G.T.Prance & F.White, 

Philos. Trans. B320: 147 (1988). 

121

122 


24. Hirtella hoehnei Pilg., Revista Mus. Paul. Univ. São Paulo 13: 1251 (1922) 

T: Brazil West-central: Mato Grosso, Ponta de Pedra, Chapadão dos Parecís, margin of Rio Sacurina, 

F.C.Hoehne 2141; holo: B, destroyed; photos: F, NY. 

Illustration: R.K.F.Pilger, Revista Mus. Paul. Univ. São Paulo 13: 1251, t. 2 (1922). 


Shrub to 5 m tall. Branches tomentellous when young. Leaves with stipules lanceolate, 

tomentose, to 6 mm long, persistent, eglandular; petioles 2–4 mm long, villous-tomentellous, 

terete, eglandular; lamina ovate to ovate-elliptic, coriaceous, 4.5–15 × 2.5–7 cm, acute to 

acuminate at apex with acumen 0–4 mm long, subcordate at base, hirsute along venation 

beneath, with palisade glands towards base; primary veins 7–11 pairs, plane to prominulous 

above, prominent beneath; midrib prominulous above, prominent and hirsute beneath. 

Inflorescences densely clustered terminal and subterminal panicles 8–20 cm long; rachis and 

branches hirsutulous. Bracts and bracteoles 2–5 mm long, acute, oblong, membranous, 

hirsutulous, persistent; margins with numerous stipitate glands. Flowers 7–8 mm long. 

Receptacle campanulate, hirsutulous on exterior, glabrous within except for deflexed hairs at 

throat; pedicels 2–4 mm long. Calyx lobes acute, hirsutulous on exterior, grey-pubescent 

within; margins with stipitate glands. Petals 5, white, glabrous. Stamens 7, unilateral, with 

short filamentous staminodes opposite; filaments far exceeding calyx lobes. Style hirsute up 

to 1/3 its length. Ovary inserted at mouth of receptacle, pilose. Fruit ellipsoid; epicarp 

smooth, glabrous; mesocarp thin, fleshy; endocarp thin, bony, densely hirsute within. 

Planalto of C Brazil. 84: BZC (GO, MS, MT), BZN (PA). Beside rivers and in wet open 

marshy places (brejos). Flowering May–Oct. Map 159. 

84. BRAZIL NORTH: Pará, Mun. de Itaituba, Km 794 Santarém-Cuiabá, Serra do Cachimbo airport, 

I.L.Amaral et al. 1006 (INPA, NY). BRAZIL WEST-CENTRAL: Goiás, Mun. de Rio Verde, road Rio Verde- 

Jutaí, G.J.Shepherd et al. 7443 (MG); Mato Grosso, Road Cuiabá-Chapada dos Guimarães, Ribeirão 

Salgadeira, A.Maciel et al. 326 (INPA); Mato Grosso, Ponta de Pedra, Chapadão dos Parecís, margin of Rio 

Sacurina, F.C.Hoehne 2154 (SP). 

25. Hirtella glandulosa Spreng., Neue Entdeck. Pflanzenk. 1: 303 (1820) 

T: Brazil: probably Espírito Santo, F.Sellow 44; holo: B, destroyed; lecto: G, fide G.T.Prance, Fl. Neotrop. 

Monogr. 9: 287 (1972); isolecto: fragment F; F.Sellow s.n. (probably isotype) LE. 

Hirtella damaziana Beauverd, Bull. Herb. Boissier, Sér. 2, 7: 706 (1907). T: Brazil Southeast: Minas 

Gerais, Belo Horizonte, L.B.Damazio 1594; holo: G. 

Hirtella hookeri Pilg., Notizbl. Königl. Bot. Gart. Berlin 6: 139 (1914). T: Brazil Northeast: Ceará, without 

locality, G.Gardner 1947; holo: B, destroyed; lecto: K, designated here; iso: BM, F, NY, P, SP, US. 

Hirtella velutina Pilg., Notizbl. Königl. Bot. Gart. Berlin 6: 141 (1914). T: Guyana: without locality, R. & 

R.Schomburgk 1051; holo: B, destroyed; iso: K. 

Hirtella wachenheimii Benoist, Bull. Mus. Natl. Hist. Nat. 29: 595 (1923). T: French Guiana: Godebert, 

H.Wachenheim 372; holo: P; iso: K. 



Tree to 25 m tall, usually much smaller. Branches tomentose when young, becoming glabrous 

and lenticellate with age. Leaves with stipules linear, 4–8 mm long, tomentellous, eglandular, 

persistent; petioles 1.5–5 (–12) mm long, tomentellous, terete, eglandular; lamina oblong to 

ovate, thick-coriaceous, 4.5–23 × 2.5–11.5 cm, acuminate at apex with acumen 2–5 mm long, 

rounded to cuneate at base, rarely weakly subcordate, hirsute beneath with palisade glands 

rare; primary veins 8–15 pairs, impressed above, prominent beneath; midrib prominulous above, 

prominent and hirsute beneath. Inflorescences terminal and subterminal panicles 9–26 cm long; 

rachis and branches hirsute-tomentellous. Bracts and bracteoles ovate, coriaceous, persistent, 

pubescent, with numerous stipitate glands. Flowers 6–8 mm long. Receptacle campanulate, 

puberulous on exterior, glabrous within except for deflexed hairs at throat; pedicels 1–3 mm 

long. Calyx lobes acute, puberulous on exterior, grey-pubescent within; margins with stipitate 

glands. Petals 5, white, glabrous. Stamens 3–5, unilateral, with a few filamentous staminodes 

opposite; filaments far exceeding calyx lobes. Style hirsute on lower portion only. Ovary 

inserted at mouth of receptacle, pilose. Fruit round to pyriform; epicarp smooth, glabrous 

when mature; mesocarp thin, fleshy; endocarp thin, bony, densely hirsute within. Fig. 70A–C.


Commonest in the planalto of C Brazil, but extending into the savannas of Amazonia and the 

Guianas. 82: FRG, GUY, SUR, VEN. 83: BOL. 84: BZC (DF, GO, MT, MS), BZE (BA, CE, 

MA, PI), BZL (MG), BZN (AM, AP, PA). Damp forest in open places, mainly gallery forest, 

river banks and savanna margins. Map 160. 

82. GUYANA: Arawai Creek, D.B.Fanshawe 313 (FD304a) (NY, P). VENEZUELA: Bolívar, Distr. Cedeño, km 6 

Maniapure-Caicara, B.Boom & M.Grillo 6542 (MYF, NY). 83. BOLIVIA: Dept. Santa Cruz, Velasco Prov., 

San José de Campamento, R.Guillen & Mormañas 4621 (K, MO, USZ). 84. BRAZIL NORTH: Amapá, Mun. 

de Macapá, Riozinho, 122 km NW of Porto Grande, S.A.Mori & R.Cardoso 17579 (MG, NY, SP); Amazonas, 

Humaitá, 15 km on road to Manaus, K.Kubitzki & H.-H.Poppendieck 79-45 (NY); Pará, Curuçá, road to 

Matucal, P.B.Cavalcante 3373 (FHO, MG, NY). BRAZIL NORTHEAST: Bahia, Mun. de Utinga, 9 km NW of 

Utinga road to Morro do Chapéu, L.P. de Queiroz & N.S.Nascimento 4217 (HUEFS, K). BRAZIL 

SOUTHEAST: Minas Gerais, Mun. de Diamantina, Cons. Mata, G.Hatschbach 30202 (MBM, NY, Z). BRAZIL 

WEST-CENTRAL: Distrito Federal, APA de Cafuringa, Fazenda Dois Irmãos, B.A.S.Pereira & D.Alvarenga 

2227 (IBGE, K); Mato Grosso, Chapada dos Guimarães, Véu de Noiva, G.T.Prance et al. 19078 (INPA, NY). 

26. Hirtella bullata Benth., J. Bot. (Hooker) 2: 216 (1840) 

T: Guyana: near Mt. Arawagang, R. & R.Schomburgk 132; holo: K; iso: BM. 

Hirtella aureohirsuta Pilg., Bot. Jahrb. Syst. 37: 538 (1906). T: Peru: Loreto, Mogobamba, A.Weberbauer 

4487; holo: B, destroyed; iso: G; iso fragment F. 

Hirtella amplexicaulis Pilg., Notizbl. Königl. Bot. Gart. Berlin 6: 138 (1914). T: Brazil North: Roraima, 

Surumi, E.Ule 7981; holo: B, destroyed; iso: L, K, MG. 

Hirtella stipitadenia Rusby, Mem. New York Bot. Gard. 7: 249 (1927). T: Bolivia: Rurrenabaque, 

H.H.Rusby 768; holo: NY; iso: US. 

Illustrations: J.D.Hooker, in C.F.P.Martius, Fl. Bras. 14(2), t. 11(1) (1867); G.T.Prance, Fl. Venez. Guayana 

4: 219 (1998). 


Shrub or small to medium-sized tree, with long horizontal branches often sprawling over 

vegetation. Branchlets tomentellous when young, becoming glabrous and lenticellate with age. 

Leaves with stipules 3–6 mm long, linear-lanceolate, tomentellous, eglandular, persistent; 

petioles 1.5–4 mm long, tomentose, terete, eglandular; lamina oblong to ovate, thickcoriaceous, 

6–22 × 3.5–10 cm, acute to short-acuminate at apex with acumen 0–12 mm long, 

subcordate or rarely rounded at base, bullate above with impressed venation, hirsute beneath; 

primary veins 11–17 pairs, impressed above, prominent beneath; midrib prominulous above, 

prominent and hirsute-tomentose beneath. Inflorescences terminal and axillary panicles 9–35 cm 

long, with long central rachis bearing short lateral branches with 1–3 flowers, or rarely reduced 

to racemes, 8–33 cm long; rachis and branches tomentose. Bracts and bracteoles 1–4 mm 

long, oblong-lanceolate, bearing large sessile glands or with numerous stipitate glands arising 

from the region of pedicel articulation. Flowers 4.5–8 mm long. Receptacle campanulate, 

tomentellous on exterior, glabrous within except at throat; pedicels 1–3 mm long. Calyx 

lobes acute, tomentellous on exterior, sparsely grey-puberulous within; margins eglandular. 

Petals 5, white, glabrous. Stamens 4–6, unilateral with small toothed staminodes opposite; 

filaments glabrous, purple, far exceeding calyx lobes. Style pilose for up to 1/2 its length. 

Ovary inserted at mouth of receptacle, pilose. Fruit ellipsoid; epicarp smooth and glabrous; 

mesocarp thin, fleshy; endocarp very thin and bony, hirsute within. 

Guyana and W Amazonia to Bolivia. 82: GUY, VEN. 83: BOL, CLM, PER. 84: BZN (AC, 

AM, RR). Open habitats at river and savanna margins and savannas. Map 161. 

82. GUYANA: Cuyuni-Mazaruni region, Pakaraima Mts., 4 km E of Chinowieng, T.W.Henkel 251 (K, US). 

VENEZUELA: Amazonas, San Carlos de Río Negro, J.G.Wessels-Boer 2334 (NY, U); Dept. Atabapo, slope of 

Huachamacari, R.L.Liesner 18344 (MO, NY); Bolívar, Distr. Piar, between Auyan-tepui and Uaipán-tepui, Río 

Urimán, O.Huber et al. 9899 (MYF, NY). 83. BOLIVIA: Dept. Beni, Moxos Province, km 27 San Borja- 

Trinidad, D.N.Smith et al. 14189 (K, LPB, MO). COLOMBIA: Guainía, Río Inírida, Caranocoa, A.Fernández- 

Perez & R.Jaramillo 7005 (COL). PERU: Madre de Dios, Tambopata Prov., Tambopata Reserve, C.Reynel & 

E.Meneses 5100 (K, MO); San Martín, Mariscal Caceres Prov., 12 km W of Tocache Nuevo, A.Gentry et al. 

25613 (MO, NY). 84. BRAZIL NORTH: Amazonas, Rio Negro, Marabitanas, O.C.Nascimento et al. 279 

(INPA, NY); Roraima, Rio Uraricoera, Serra Curacinha, J.M.Pires et al. 16939 (MG). 

123

124 


27. Hirtella americana L., Sp. Pl. 34 (1753) 

Chrysobalanus americanus (L.) Morales, Anales Acad. Ci. Med. Habana 23: 391 (1887). T: Tropical 

America: without locality, Linnaeus, Hortus Cliffortianus, pl. 17; holo: BM. 

Hirtella mollicoma Kunth, Nov. Gen. Sp. 7: 263 (1825); Hirtella mollissima Hemsl., Biol. Centr. Am. Bot. 

1: 366 (1880), sphalm; Chrysobalanus mollicoma (Kunth) M.Gómez, Fl. Cuba 39 (1887). T: Colombia: 

Magdalena, banks of Río Magdalena, El Espiñal, A. von Humboldt & A.J.A.Bonpland 1798; holo: P; fragment F. 


Tree to 20 m tall, usually smaller, or shrub. Branches tomentellous when young, becoming 

glabrous with age. Leaves with stipules lanceolate, 2–8 mm long, tomentellous, eglandular, 

persistent; petioles 2–3 mm long, tomentellous, terete, eglandular; lamina elliptic to oblong, 

coriaceous, 5.5–19 (–23) × 2–7 cm, acuminate at apex with acumen 2–8 mm long, rounded to 

subcuneate at base; lower surface hirsute; primary veins 9–13 pairs, plane or slightly 

impressed above, prominent beneath; midrib prominulous above, prominent and tomentellous 

beneath. Inflorescences terminal and axillary panicles 5–20 mm long; rachis and branches 

tomentellous. Bracts and bracteoles 1–5 mm long, ovate, persistent, tomentellous, bearing 

numerous large sessile or slightly stipitate glands. Flowers 4–6 mm long. Receptacle 

campanulate, tomentellous on exterior, glabrous within except for throat; pedicels 0.5–2 mm 

long. Calyx lobes rounded, tomentellous on exterior, grey-pubescent within; margins eglandular. 

Petals 5, glabrous, white. Stamens 3, unilateral with small teeth opposite; filaments white to 

purple, far exceeding calyx lobes. Style hirsute up to 1/2 its length. Ovary inserted at mouth 

of receptacle, pilose. Fruit ellipsoid; epicarp sparsely pubescent to glabrous; mesocarp thin, 

fleshy; endocarp thin, hard, bony, hirsute within. 

Cuba, through Central America, Mexico (Tabasco), to Panama, and in N Colombia and 

Venezuela. 79: MXT. 80: BLZ, COS, GUA, HON, NIC, PAN. 81: CUB, DOM. 82: VEN. 83: 

CLM, ECU. Dry forest on elevated ground. Map 162. 

79. MEXICO TROPICAL: Tabasco, La Palma, Balancan, E.Matuda 3270 (A, F, K, MEXU, MICH, MO, NY). 

80. BELIZE: Toledo, Big Fall Estate, E of Southern Hwy., G.Davidse et al. 35585 (K, MO). COSTA RICA: 

Guanacaste, Cantón de La Cruz, P. N. Guanacaste, Cerro El Hacha, R.Espinoza 278 (K, MO). NICARAGUA: 

Zelaya, Mun. de Siuna, La Pimienta, F.Ortiz 1974 (MO, NY). PANAMA: San José I., Pearl Archipelago, 

C.O.Erlanson 121 (GFI, MICH, NY, US). 81. CUBA: Isle of Pines, S of Nueva Jerona, O.E.Jennings 208 

(BM, GH, NY, US). 82. VENEZUELA: Zúlia, Dept. Perijá, 15–17 km SE of San Felipe, G.S.Bunting 5655 (NY). 

83. COLOMBIA: Meta, R. N. de La Macarena, Quebrada Honda below confluence of Río Zanza, J.Thomas et al. 

1536 (COL, P). ECUADOR: Napo, Río Cuyabeno, 6 km above Puerto Bolívar, J.Brandbyge et al. 36177 

(AAU, QCA). 

28. Hirtella santosii Prance, Revista Brasil. Bot. 2: 34 (1979) 

T: Brazil Northeast: Bahia, Una, Fazenda São Rafael, T.S. dos Santos 300; holo: CEPEC; iso: NY, RB. 

Illustration: G.T.Prance, Revista Brasil. Bot. 2: 36 (1979). 


Tree 8 m tall. Branches tomentellous when young, becoming glabrous with age, not conspicuously 

lenticellate. Leaves with stipules linear, 4–5 mm long, subpersistent, tomentellous; 

petioles 3–4 mm long, terete, tomentellous, eglandular; lamina elliptic, coriaceous, 7–12.5 × 

2–5.5 cm, rounded to subcuneate at base, acuminate at apex with acumen 4–10 mm long, 

hirsute beneath on venation; midrib prominulous above but in impressed channel, prominent 

and densely hirsute beneath; primary veins 15–19 pairs, prominulous above but in impressed 

channels, prominent and densely hirsute beneath; secondary venation slightly impressed 

above giving lightly bullate appearance, prominulous beneath. Inflorescences of terminal racemes 

to 6–10 cm long; rachis and branches densely rufous-tomentose. Bracts small, persistent, 

lanceolate, tomentose; bracteoles ovate, persistent, tomentose with numerous sessile or shortly 

stipitate glands around margins. Flowers 7–9 mm long. Receptacle campanulate, tomentellous 

on exterior, glabrous within except for deflexed hairs at throat; pedicels 5–6 mm long, 

tomentose, eglandular except on paired bracteoles. Calyx lobes 5, acute, tomentose on 

exterior, densely tomentose on interior. Petals 5, glabrous. Stamens 6, unilateral. Ovary inserted 

in middle of receptacle, pilose. Style pilose at base, glabrous above. Fruit not seen. 

Brazil; known only from the type collection. 84: BZE (BA). Forest. Map 163.


29. Hirtella aequatoriensis Prance, Kew Bull. 54: 995 ("1999") [2000] 

T: Ecuador: Pastaza, Pastaza Cantón, 35 km S of Curaray, E.Gudiño 728; holo: QCNE; iso: K, MO. 

Illustration: G.T. Prance, Kew Bull. 54: 996 (1999). 

Tree 12 m tall. Branches tomentellous when young. Leaves with stipules lanceolate, c. 3 mm 

long, tomentose, persistent; petioles 4–5 mm long, terete, eglandular, tomentose; lamina elliptic, 

coriaceous, 14–18 × 8–9.5 cm, acuminate at apex with acumen 5–9 mm long, subcuneate at 

base, pubescent above on midrib and primary veins, sparsely hirsute beneath; midrib plane 

above, prominent and tomentellous beneath; primary veins 10–13 pairs, slightly impressed 

above in mature leaves; secondary venation slightly impressed. Inflorescences sparsely-branched 

panicles with a central rachis 7–25 cm long and short branches bearing 1–3 flowers; rachis and 

branches hispid-tomentose; bracteoles persistent, tomentose, bearing numerous stalked tacklike 

glands; bracts eglandular. Flowers c. 4 mm long. Receptacle campanulate, hispidtomentose 

on exterior, glabrous within except for deflexed hairs in throat; pedicels 3–5 mm 

long, with stalked glands. Calyx lobes rounded, hispid-tomentose on exterior, densely tomentose 

within; margins eglandular. Petals 5, white, glabrous. Stamens 3, unilateral; filaments far 

exceeding calyx lobes. Style hirsute on lower 1/2. Ovary inserted at mouth of receptacle, 

pilose. Fruit not seen. 

Ecuador. 83: ECU. Primary forest. Map 164. 

83. ECUADOR: Zamora-Chinchipe, Zamora Cantón, Jamboe Bajo, E border of Podocarpus N. P., J.L.Clark et 

al. 3160 (K, MO). 

30. Hirtella guatemalensis Standl., Trop. Woods 11: 19 (1927) 

T: Guatemala: Livingston Dept., Livingston, H. von Tuerckheim II-1141 (Herb. J.D.Smith 8680); holo: US. 

Illustration: P.C.Standley, Field Mus. Nat. Hist., Bot. Ser. 10, fig. 38 (1931). 


Tree, small. Branches hispid when young, becoming glabrous and lenticellate with age. 

Leaves with stipules c. 5 mm long, linear, pilose, persistent, eglandular; petioles 1.5–4 mm 

long, hispid when young, becoming glabrous with age, terete, eglandular; lamina oblong, 

subcoriaceous, 11–22 × 4–7.5 cm, acuminate at apex with acumen 8–20 mm long, rounded to 

weakly subcordate at base, hirsute on primary venation beneath; primary veins 11–16 pairs, 

almost plane above, prominent beneath; midrib prominulous above, prominent and hirsute 

beneath. Inflorescences terminal and axillary little-branched panicles, with a long central 

rachis 9–18 cm long, bearing short few-flowered lateral branches; rachis and branches 

hispid. Bracts and bracteoles 3–5 mm long, oblong to oblong-lanceolate, hispid, persistent; 

margins of bracts bearing small stipitate glands. Flowers 6–8 mm long. Receptacle 

campanulate, hispid on exterior, glabrous within except at throat; pedicels 4–10 mm long. 

Calyx lobes rounded, hispid on exterior, puberulous within; margins eglandular. Petals 5, 

glabrous, white. Stamens 3–4, unilateral with short filamentous staminodes opposite; 

filaments glabrous, far exceeding calyx lobes. Style hirsute up to 1/3 its length. Ovary 

inserted at mouth of receptacle, pilose. Fruit round to ellipsoid; epicarp tomentose; mesocarp 

very thin, fibrous; endocarp 1.5–2 mm thick, hard, fibrous, hirsute within. 

Central America. 80: BLZ, COS, GUA, HON, NIC, PAN. Forest on well drained soil. 

Flowering Dec.–Feb. Map 165. 

80. BELIZE: Toledo, San Antonio road, near Westmorland, N.S.Stevenson 10697 (US); Forest Home, 

W.A.Schipp 1098 (A, BM, F, FI, GH, K, MICH, MO, NY, S, Z). COSTA RICA: Guanacaste, Cantón de 

Liberia, Finca La Pasmompa, R.Robles 2831 (K, MO). GUATEMALA: Dept. Izabal, between Cienaya and 

Seja, E.Contreras 10716 (US). HONDURAS: Gracios a Dios, 2 km NW of Bulebar, J.Saunders 1197 (NY). 

NICARAGUA: Río San Juan, Santa Eduviges, P.P.Moreno 23014 (MO, NY); Zelaya, Km 8.1–12.2 Kururia– 

San Jerónimo, W.D.Stevens 7555 (MO, NY). 

31. Hirtella trichotoma Prance, Kew Bull. 47: 642 (1992) 

T: Costa Rica: Limón, Cerro Coronel, E of Río Zapote, W.D.Stevens, G.Herrera & O.M.Montiel 24969; holo: 

K; iso: MO. 

Illustration: G.T.Prance, loc.cit.: 643. 

Map: G.T.Prance, loc.cit.: 645. 

125

126 


Tree 15 m tall. Branches puberulous when young, becoming glabrous with age. Leaves with 

stipules caducous (not seen); petioles 2–3 mm long, terete, eglandular; lamina elliptic, 

chartaceous, 14–21.5 × 6.7–10 cm, rounded at base, acuminate at apex with acumen 5–10 mm 

long, glabrous above, hirsute beneath on venation; midrib prominulous and glabrous above, 

prominent and hirsute beneath; primary veins 11–13 pairs, slightly impressed above, prominent 

and sparsely hirsute beneath; venation sparsely hirsute and papillate. Inflorescence a panicle 

with a central rachis 8–27 cm long and short branches 4–7 mm long bearing 3 flowers each; 

rachis and branches rufous-tomentose. Bracts and bracteoles lanceolate, 2–5 mm long, 

tomentellous, with few sessile glands and occasionally an apical gland. Flowers 7–10 mm 

long. Receptacle campanulate, tomentose on exterior, glabrous within except at throat and at 

base of ovary; pedicels 6–8 mm long, bracteolate. Calyx lobes 5, acute, tomentose on exterior, 

tomentellous within. Petals 5, white, caducous. Stamens 3, inserted on one side of ovary, 

with short staminodes opposite; filaments glabrous, far exceeding calyx lobes. Ovary 

inserted at mouth of receptacle, pilose. Style hirsute for 3/4 of length. Fruit not seen. 

Costa Rica. 80: COS. Tall evergreen forest. Flowering in March. Map 166. 

80. COSTA RICA: Limon, Cantón de POCOCI, R. N. F. S. Barra del Colorado, Llanura de Tortuguero, 

F.Araya & J.González 520 (K, INB). 

32. Hirtella papillata Prance, Kew Bull. 49: 359 (1994) 

T: Costa Rica: Puntarenas, R. F. Golfo Dulce, Quebrada Agua Buena, B.Hammel 18150; holo: MO; iso: CR. 

Illustration: G.T.Prance, loc.cit.: 360. 

Shrub 4 m tall. Branches puberulous when young, inconspicuously lenticellate. Leaves with 

stipules linear, 5–6 mm long, axillary, caducous; petioles 2–4 mm long, terete, puberulous, 

eglandular; lamina oblong, coriaceous, 14–19 × 6–7.5 cm, rounded at base, bluntly acuminate at 

apex with acumen 2–5 mm long, sparsely hirsutulous beneath on principal venation with 

minor venation papillate, glabrous above; midrib prominulous above, prominent beneath; 

primary veins 13–16 pairs, prominulous above, prominent beneath. Inflorescences of terminal 

much-branched panicles; rachis and branches grey-puberulous. Bracts and bracteoles 1–2 mm 

long, acute, puberulous, with sessile glands on about 1/3 and others eglandular. Flowers c. 4 mm 

long. Receptacle campanulate, tomentellous on exterior, glabrous within except for deflexed 

hairs at throat; pedicels 1–2 mm long, tomentellous, eglandular. Calyx lobes 5, acute, 

tomentellous on exterior, puberulous within, with minute glands on margins. Petals 5, white. 

Stamens 3, unilateral with minute staminodes opposite. Ovary inserted at mouth of receptacle, 

pilose. Style pilose on lower 1/3 only. Fruit not seen. 

Central America. 80: COS, HON. Rainforest on slopes 250–300 m. Map 167. 

80. COSTA RICA: Puntarenas, R. F. Golfo Dulce, between Rancho Quemado and Drake, R.Aguilar 113 (CR, 

K, MO); Golfito, Refugio Nacional de Fauna Silvestre, Cerro Nicuesa, G.Rivera 2256 (CR, K, MO). 

HONDURAS: Atlantida, base of N slope of Pico Bonito, 10 km SW of La Ceiba, R.Evans 1654 (K, MO). 

The type of the species name was erroneously cited as B.Hammel 12150 in the protologue. 

33. Hirtella eriandra Benth., J. Bot. (Hooker) 2: 217 (1840) 

T: Brazil North: Amazonas, Pedrero, Rio Negro, R. & R.Schomburgk 886; holo: K; iso: BM, CGE, GH, NY, P, W. 


Shrub or small tree. Branches tomentellous when young, becoming glabrous with age. Leaves 

with stipules linear, to 5 mm long, tomentellous, persistent, eglandular; petioles 2–4 mm long, 

tomentose, terete, eglandular; lamina oblong, chartaceous to subcoriaceous 8–19 × 3.3–8 cm, 

acuminate at apex with acumen 3–15 mm long, rounded at base, hirsute beneath at least on 

principal venation, usually on other veins as well, with palisade glands frequent towards 

base; primary veins 10–13 pairs, plane or nearly so above, prominent beneath; midrib 

prominent above, prominent and hirsute beneath. Inflorescences terminal and subterminal 

panicles, with a thick central rachis 12–50 cm long bearing short branches or clusters of 

flowers; rachis and branches tomentellous. Bracts and bracteoles 2–6 mm long, oblong to 

lanceolate, tomentose, with a few large sessile glands or a large terminal gland on some


bracts only. Flowers 6–7 mm long. Receptacle campanulate, tomentellous on exterior, glabrous 

within except at throat; pedicels 0.5–6 mm long. Calyx lobes acute, sparsely tomentellous on 

exterior, sparsely puberulous to glabrescent within; margins eglandular. Petals 5, greenishwhite, 

glabrous. Stamens 6, unilateral; filaments pink, far exceeding calyx lobes, hirsutulous on 

lower portion. Style hirsute up to 1/2 its length. Ovary inserted at mouth of receptacle, 

tomentose. Fruit rotund; epicarp tomentose, longitudinally striate when mature; mesocarp thin; 

endocarp thin, fibrous, hard, hirsutulous within. 

Amazonia and C Brazil. 83: BOL, CLM, PER. 84: BZE (MA), BZN (AM, AP, PA). Riverine 

forest, savanna margins or cerrado and secondary forest. Flowering Aug.–Dec. Map 168. 

83. BOLIVIA: Dept. Pando, 7–10 km SW of Guayaramirim, road to Riberalta, W.R.Anderson 11921 (INPA, NY). 

COLOMBIA: Tolima, Mun. de Mariquita, Mariquita F.R., A.Gentry et al. 71863 (K, MO). PERU: Amazonas, 

Río Santiago, 2 km from Caterpiza, V.Huashikat 783 (MO, NY); Loreto, Maynas, Prov., Sargento Lores, Río 

Tuhuayo, R.Vásquez & N.Jaramillo 13209 (K, MO). 84. BRAZIL NORTH: Amapá, Parque Florestal do 

Amapá, Fazendinha, D.F.Austin et al. 6994 (MG, NY); Amazonas, Rio Negro, between Rio Caurés and Barcelos, 

G.T.Prance et al. 15134 (FHO, INPA, NY, U, US); Pará, Tomé–Açu, O.C.Nascimento 352 (FHO, MG, NY). 

BRAZIL NORTHEAST: Maranhão, km 180 of BR-316, Zé Doca, N.A.Rosa & H.Vilar 2923 (MG, NY). 

34. Hirtella paniculata Sw., Prodr. 51 (1788) 

Hirtella hirsuta Lam., Tabl. Encycl. 4(2): 114 (1797) ["1793"], nom. superfl. illeg.; Hirtella aggregata Poir., 

Encycl., Suppl. 3: 53 (1813), nom. superfl. illeg.; Hirtella indecora Schott ex Spreng., Syst. Veg. 4(2): 404 (1827), 

nom. superfl. illeg. T: French Guiana: without locality, J.P.B. von Rohr s.n.; holo: BM, Banks Herb.; iso: C, LE. 

Hirtella bracteosa Steud., Flora 26: 761 (1843). T: Suriname: without locality, W.R.Hostmann 601; holo: P; 

iso: BM, C, CGE, F, GH, GOET, L, LE, M, NY, S, W. 

Illustration: W.Fawcett & A.B.Rendle, Fl. Jamaica 3(1): 255 (1914). 


Shrub to 4 m tall or small tree. Branches pilose when young, becoming glabrous with age. 

Leaves with stipules 3–12 mm long, linear, persistent, sparsely hirsute; petioles 1–2 mm 

long, terete, eglandular, sparsely pilose; lamina oblong-lanceolate to ovate-orbicular, most 

frequently ovate, coriaceous, 2.5–13.5 × 1.3–5.5 cm, acuminate at apex with acumen 1–7 mm 

long, rounded to subcordate at base, hirsute beneath at least on venation, with palisade 

glands present towards base; primary veins 7–9 pairs, plane or nearly so above, prominent 

beneath; midrib prominulous above, prominent and hirsute beneath. Inflorescences terminal 

and axillary little-branched panicles, 3–26 cm long, or rarely reduced to racemes; rachis and 

branches sparsely hirsute-hispid. Bracts and bracteoles 2–6 mm long, oblong to lanceolate, 

membranous, sparsely hirsute, persistent; margins bearing small stipitate glands or sessile 

rather inconspicuous glands. Flowers 5–7 mm long. Receptacle campanulate, hirsute on 

exterior, glabrous within except for deflexed hairs at throat; pedicels 1–3 mm long. Calyx 

lobes acute, hirsute on exterior, grey-puberulous within; margins eglandular. Petals 5, white 

to purple, glabrous. Stamens 5–6, unilateral with a toothed portion of the circle opposite; 

filaments purple, far exceeding calyx lobes. Style hirsute up to 1/2 its length. Ovary inserted 

at or near mouth of receptacle, pilose. Fruit ellipsoid; epicarp smooth, glabrous; mesocarp 

thin, fleshy; endocarp thin, hard, bony, hirsute within. 

St. Vincent, Trinidad, Venezuela, Guianas, Colombia and N Brazil. 81: TRT, WIN. 82: FRG, 

GUY, SUR, VEN. 83: CLM. 84: BZN (AM, AP, PA, RR). Open sandy places, savannas, 

savanna margins, sandy river banks and islands. Flowering throughout the year. Map 169. 

81. TRINIDAD-TOBAGO: Trinidad, Piarco Savanna, S of Arouca, N.L.Britton et al. 89 (GH, K, NY, US). 

82. FRENCH GUIANA: Mt. Des Trois Pitons, J.J. de Granville 4321 (CAY, NY, P). GUYANA: E Berbice- 

Corentyne Region, Awara Savanna, 85 mi [135 km] S of Canje R., J.J.Pipoly 9334 (NY, P, US). SURINAME: 

Sipaliwini, Blanche Marie Waterfall, Nickerie R., R.Evans et al. 2413 (K, MO). VENEZUELA: Bolívar, 8 km 

from Ciudad Piar, E.Briceño et al. 4 (K, MYF); Amazonas, Dept. Atabapo, between Culebra airport and Cerro 

Duida, R.L.Liesner 17576 (MO, NY). 83. COLOMBIA: Meta, Mun. La Macarena, Río Cristales, R.Callejas 

& O.Marulanda 7160 (K, MO). 84. BRAZIL NORTH: Amazonas, Rio Negro, Rio Téa, 40 km above mouth, 

K.Kubitzki et al. 79-241 (INPA, US); Pará, P. N. Tucumaque, Rio Paru do Oeste, P.B.Cavalcante 2477 (MG); 

Roraima, Mun. de Caracaraí, Manaus-Caracararaí km 522–524, C.A. Cid Ferreira 9151 (INPA, K, NY). 

127

128 


35. Hirtella castilloana Prance, Kew Bull. 54: 110 (1999), as castillana 

T: Venezuela: Amazonas, Río Cataniapo, A.Castillo 3335; holo: UCV; iso: K. 


Map: G.T.Prance, Kew Bull. 54: 106 (1999). 

Tree to 6 m tall. Branches with few stiff appressed hairs when young, soon becoming 

glabrous and shiny. Leaves with stipules linear, 5–6 mm long, 0.5 mm wide, early caducous; 

petioles 2–3 mm long, with a few stiff appressed hairs, eglandular, terete; lamina lanceolate, 

chartaceous, 10–14 × 1.8–3.5 cm, acuminate at apex with acumen 5–12 mm long, slightly 

subcordate at base, glabrous above, with a few stiff appressed hairs beneath on midrib and 

principal venation, with paired sessile glands towards base of lamina and several others 

along lamina near to midrib; midrib prominulous above, prominent beneath; primary veins 

10–13 pairs, prominulous on both surfaces. Inflorescences racemes or occasionally a few 

branches bearing 2 flowers, 6–11 cm long; rachis sparsely appressed-hispid. Bracts and 

bracteoles 6–9 mm long, persistent; margins of most with few sessile glands; tip glandular. 

Flowers 4–5 mm long. Receptacle campanulate, sparsely appressed hispid on exterior, 

glabrous within except for deflexed hairs at throat; pedicels 3–5 mm long. Calyx lobes acute, 

with sparse appressed hairs on exterior, grey-tomentellous within; margins eglandular. Petals 

5, white, glabrous. Stamens 6, unilateral; filaments purple, far exceeding calyx lobes. Style 

hirsute on lower ¼. Ovary inserted near to mouth of receptacle, pilose. Fruit not seen. 

Amazonian Venezuela. 82: VEN. 83: CLM. Savannas. Map 170. 

82. VENEZUELA: Amazonas, Dept. Atures, Puerto Ayacucho, Samanapo, Río Cataniapo, A.Castillo 1340 

(K, UCV). 83. COLOMBIA: Vaupés, Mitú, lower Río Paraná-pichuna, J.L.Zarucchi 1983 (K, NY). 

36. Hirtella deflexa Maguire, Fieldiana, Bot. 28: 255 (1952) 

T: Venezuela: Bolívar, Mt. Roraima district, Arabupu, A.S.Pinkus 46; holo: NY; iso: F, S, US. 


Shrub or small tree. Branches pilose when young, becoming glabrous with age. Leaves with 

stipules to 6 mm long, linear, hispid, subpersistent, eglandular; petioles 1–2 mm long, terete, 

eglandular, hispid; lamina ovate, subcoriaceous, 4–8 × 2–4 cm, acute to acuminate at apex 

with acumen 0–5 mm long, cordate at base, sparsely hirsute beneath; primary veins 9–12 pairs, 

plane or almost so above, prominent beneath; midrib prominulous above, prominent and hirsute 

beneath. Inflorescences terminal and predominantly axillary little-branched panicles, 4.5–10 cm 

long; rachis and branches sparsely hirsute. Bracts and bracteoles 2–3 mm long, oblong, 

membranous, persistent; margins bearing slender stipitate glands. Flowers 5–6 mm long. 

Receptacle campanulate, sparsely hirsutulous on exterior, glabrous within except at throat; 

pedicels 2–3.5 mm long. Calyx lobes rounded to acute, hirsutulous on exterior, greypubescent 

within; margins eglandular. Petals 5, glabrous, white. Stamens 4–6, unilateral; 

filaments reddish purple, far exceeding calyx lobes. Style hirsute on lower portion only. 

Ovary inserted midway up receptacle, villous. Fruit not seen. 

Venezuela (Amazonas and Bolívar). 82: VEN. River banks. Flowering Nov.–Jan. Map 171. 

82. VENEZUELA: Bolívar, Distr. Sifontes, Río Yuruaní, 5 km S of Wadakapiapué-tepui, O.Huber 11774 

(MYF, NY); Bolívar, 146 km S of El Dorado, J.A.Steyermark et al. 104223 (NY, VEN); Amazonas, Dept. 

Casiquiare, 12 km from San Fernando de Atabapo, G.Aymard et al. 6458 (K, MO, PORT); Amazonas, Dept. 

Río Negro, N of Cumaquita, 90 km ENE of San Carlos de Río Negro, R.Liesner & F.Delascio 21878A (MO, NY). 

37. Hirtella tentaculata Poepp., in E.F.Poeppig & S.F.Endlicher, Nov. Gen. Sp. Pl. 3: 75 

(1845) 

T: Brazil North: Pará, Breves, E.Poeppig s.n.; holo: W; iso: LE; iso fragment: F. 



Tree to 20 m tall. Branches tomentellous when young, becoming glabrous and lenticellate 

with age. Leaves with stipules 2–3 mm long, ovate, tomentose, persistent; petioles 3–8 mm


long, tomentellous, caducous, eglandular; lamina elliptic to oblong-elliptic, coriaceous, 7–24 × 

3.5–10.5 cm, acuminate at apex with acumen 2–10.5 mm long, rounded at base, with 

impressed venation above, hirsute beneath, with a few palisade glands towards base; primary 

veins 10–13 pairs, impressed above, prominent beneath; midrib plane above, prominent and 

hirsute beneath. Inflorescences densely-flowered terminal and axillary panicles 6–25 cm 

long; rachis and branches tomentellous to puberulous. Bracts and bracteoles 1–6 mm long, 

oblong to lanceolate; glands numerous, stipitate on stalks up to 8 mm long, arising from 

pedicels and axils of flowers. Flowers 4–6 mm long. Receptacle campanulate, puberulous on 

exterior, glabrous within except at throat. Calyx lobes rounded to acute, puberulous on 

exterior, grey-pubescent within; margins eglandular. Petals 5, glabrous. Stamens 3–5, unilateral, 

with short filamentous staminodes opposite; filaments glabrous, far exceeding calyx lobes. 

Style hirsute up to 1/3 its length. Ovary inserted at mouth of receptacle, pilose. Fruit ellipsoid; 

epicarp smooth, glabrous; mesocarp thin, fleshy; endocarp thin, hard, bony, hirsute within. 

Eastern Venezuela, Guyana, and NE Brazil. 82: GUY, VEN. 84: BZE (MA), BZN (PA). 

Non-flooded or periodically flooded forest. Flowering May–Nov. Map 172. 

82. GUYANA: Mt. Ayanganna, NE side, S.S. & C.Tillett 44969 (NY). VENEZUELA: Bolívar, lower Quebrada 

O-paru-má, trib of Río Pacairo, J.A.Steyermark 60561 (F, NY, US); Urimán, Río Chirea, L.Bernardi 769 (K, 

NY, VEN). 84. BRAZIL NORTH: Pará, Rio Jarí, Estrada Planalto A., N.T.Silva 2170 (BISH, IAN, NY); 

Pará, Igarapé Cuçari, Planalto de Santarém, R.L.Fróes 31771 (IAN). BRAZIL NORTHEAST: Maranhão, Mun. 

de Monção, P. I. Guajá, Rio Turiaçu, W.Balée 3275 (MG, NY). 

38. Hirtella macrophylla Benth. ex Hook.f., in C.F.P.Martius, Fl. Bras. 14(2): 36 (1867) 

T: Venezuela: Amazonas, San Carlos, Rio Negro, R.Spruce 3095; lecto: K, fide G.T.Prance, Fl. Neotrop. 

Monogr. 9: 295 (1972); isolecto: BM, BR, CGE, LE, NY, OXF, P, TCD, W. 

Illustrations: J.D.Hooker, in C.F.P.Martius, Fl. Bras. 14(2), t. 12 (1867); G.T.Prance, Fl. Ecuador 10: 17 (1979). 


Tree, small. Branches tomentellous when young, soon becoming glabrous and lenticellate. 

Leaves with stipules 2–3 mm long, subulate, persistent, tomentose; petioles 3–10.5 mm long, 

tomentellous, terete, eglandular; lamina elliptic, coriaceous, 10–18 × 5–9 cm, acuminate at 

apex with acumen 3–10 mm long, rounded to subcuneate at base, sparsely hirsute beneath on 

primary venation, with palisade glands frequent towards base; primary veins 7–10 pairs, 

prominulous above, prominent beneath; midrib plane above, prominent and hirsute beneath. 

Inflorescences terminal and axillary panicles 10–30 cm long; rachis and branches 

tomentellous to sparsely puberulous. Bracts and bracteoles to 8 mm long, oblong, puberulous, 

persistent, with numerous long-stalked glands arising from pedicels and axils of flowers. 

Flowers 5–6 mm long. Receptacle campanulate, puberulous on exterior, glabrous within 

except at throat. Calyx lobes rounded, sparsely puberulous on exterior, grey-pubescent 

within; margins eglandular. Petals 5, glabrous. Stamens 4–6, unilateral with a toothed 

portion of arc opposite them; filaments glabrous, far exceeding calyx lobes. Style hirsute up 

to 1/2 its length. Ovary inserted at mouth of receptacle, pilose. Fruit ellipsoid; epicarp 

smooth, glabrous; mesocarp thin, fleshy; endocarp hard, thin, bony, hirsute within. Fig. 71. 

Amazonian Colombia, Guianas, Peru, Ecuador, Venezuela and Brazil. 82: FRG, VEN. 83: CLM, 

ECU, PER. 84: BZC (MT), BZN (AM, PA). River banks and secondary forest. Map 173. 

82. FRENCH GUIANA: Piste de St. Elie, D.Sabatier & M.F.Prevost 3656 (CAY, K). VENEZUELA: 

Amazonas, Dept. Casiquiare, Yavita, G.S.Bunting et al. 3825 (U); Bolívar, Mun. de Raul Leoni, 53 km SW of 

Uriman, A.Fernández 3339 (MYF, NY). 83. COLOMBIA: Vaupés, Rio Apaporis, Soratama, R.E.Schultes & 

I.Cabrera 13622 (A, BM, COL, GH, NY, US). ECUADOR: Napo, Tena Cantón, Jatun Sacha Biol. Sta. 8 km E 

of Misahuallí, D.Neill & S.Espinoza 9829 (K, MO, QCNE). PERU: Pasco, Prov. Oxapampa, Distr. Isocozacín, 

W.Pariona et al. 867 (K, MO); Amazonas, Bagua Prov., Yamayakat, R.Vásquez & N.Jaramillo 20306 (K, 

MO). 84. BRAZIL NORTH: Amazonas, Manaus, Rio Negro, Igarapé Buião, R.E.Schultes & W.A.Rodrigues 

26184A (ECON, INPA); Pará, Marajó, Campos de Joanes, N.A.Rosa 3607 (MG, NY). BRAZIL WEST- 

CENTRAL: Mato Grosso, Mun. de Novo Aripuanã, Km 120 towards São Francisco Mine, C.A. Cid Ferreira 

5668 (INPA, NY). 

129

130 


Figure 71. Hirtella macrophylla. A, habit; B, flower and buds; C, flower section; D, petal 

(A–D, H.S.Lugo 4286 and D.Coelho INPA3935). Scale bars: A = 5 cm; B = 2 mm; C, D = 1 mm. 

Drawn by H.M.Fukuda. Previously published in G.T.Prance, Fl. Ecuador 10: 17 (1979).

Vol. 2 part 1 - Species Plantarum Programme

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