flora neotropica - CNCFlora

FLORA NEOTROPICA 

MONOGRAPH 9S 

CHRYSOBALANACEAE 

by 

Ghillean T. Prance 

C ^ \ [ ??TROPIC OP CANCER 

|tROPIC CO CAPIICO IN 

Published for 

Organization for Flora Neotropica 

by 

The New York Botanical Garden 

New York 

Issued 8 March 1989


by 



Board Members 

Paulo G. Windisch, President, Universidade Estadual Paulista (UNESP), Depto. de 

Botanica, Caixa Postal 136, 15001 S. Jose do Rio Preto, Sao Paulo, Brazil (1990). 

Stephan Beck, Vice-President, Herbario Nacional de Bolivia, Casilla 20127, La Paz, 

Bolivia (1990). 

Alwyn Gentry, Secretary, Missouri Botanical Garden, P.O. Box 299, St. Louis, Mis- 

souri 63166, U.S.A. (1993). 

C. C. Berg, Treasurer, The Norwegian Arboretum and the Botanical Garden, Uni- 

versity of Bergen, N-5067 Store Milde, Norway (1993). 

Enrique Forero, Missouri Botanical Garden, P.O. Box 299, St. Louis, Missouri 63166, 

U.S.A. (1990). 

Alcides R. Teixeira, Chacara Botanica, Caixa Postal 85, 13300 Itui, Sao Paulo, Brazil 

(1990). 

Ghillean T. Prance, Royal Botanic Gardens, Kew, Richmond, Surrey TW9 3AB, 

United Kingdom (1990). 

Paul J. M. Maas, Institute of Systematic Botany, Heidelberglaan 2, P.O. Box 80.102, 

3508 TC Utrecht, Netherlands (1993). 

Nanuza Menezes, Departamento de Botanica, Instituto de Biociencias-USP, Caixa 

Postal 11461, 05421 Sao Paulo, Sao Paulo, Brazil (1993). 

Scott A. Mori, Ex Officio, Herbarium, The New York Botanical Garden, Bronx, 

New York 10458. 

Staff Committee 

Ghillean T. Prance, Chairman 

Enrique Forero, Ex-Officio 

Editorial Committee 

James L. Luteyn, Editor 

Scott A. Mori, Editor 

Maria L. Lebr6n-Luteyn, Managing Editor 

H. David Hammond, Copy Editor 

Leif Ryvarden (1987-1992) 

John Wurdack (1984-1989) 

William R. Anderson (1984-1989) C. C. Berg (1984-1989) 

William Burger (1986-1991) Mireya Correa (1986-1991) 

Monograph 9S was edited by H. David Hammond and Maria L. Lebr6n-Luteyn 

ORGANIZATION FOR FLORA NEOTROPICA 

Founded and Conducted under the Auspices of UNESCO 

GEOGRAPHIC CONSULTANTS 

Central America and Mexico ........................... .......... Jerzy Rzedowski 

West Indies ..................................................... Richard A. Howard 

Colombia- Ecuador .................................................. Enrique Forero 

Peru-Bolivia ... ................................................. Ram6n H. Ferreyra 

Venezuela-Guiana .............................................. Leandro Aristeguieta 

Brazil ................................................... Luiz Guimaraes de Azavedo 

THIS PUBLICATION IS PRINTED ON ACID-FREE PAPER.

FLORA NEOTROPIC 

MONOGRAPH 9S 


by 

Ghillean T. Prance 

l\ -I- 

TROPIC OF CANCER 

FLORA 

N E OTROPICA~ 

TROPIC OF CAPRICORN 

Published 

for 


by 


New York 

Issued 8 March 1989

Copyright ? 1989 


Published by 


Bronx, New York 10458 

International Standard Serial Number 0071-5794 

Library of Congress Cataloging in Publication Data 

Flora neotropica. - Monograph no. 1 - New York: Published 

for Organization for Flora Neotropica by the New York 

Botanical Garden, 1968- 

v.: ill.; 26 cm. 

Irregular. 

Each issue has distinctive title. 

Separately cataloged and classified in LC before monograph no. 40. 

ISSN 0071-5794 = Flora neotropica. 

1. Botany-Latin America-Classification-Collected works. 2. Botany- 

Tropics-Classification-Collected works. 3. Botany-Classification-Col- 

lected works. I. Organization for Flora Neotropica. II. New York Botanical 

Garden. 

QK205.F58 581.98'012-dc19 85-647083 

AACR 2 MARC-S 

Library of Congress [8508] 

ISBN 0-89327-338-4


GHILLEAN T. PRANCE1 

TABLE OF CONTENTS 

A b stra ct . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 

Resumo ..................................................................... ......... 1 

In tro d u ctio n .................................................................................. 2 

E m bryology .............................. .................................................... 3 

R are Species ........................................................................ .......... 3 

System atic Treatm ent .......................................................................... 4 

C hrysobalanus ............................................................................ 4 

L ica n ia . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6 

P a rin a ri ....... .... ......... ...................... ............. .. .................... ... .. 5 6 

Exellodendron .................................................................. .......... 59 

M ara n thes ................... ................................................ .......... 59 

C ou ep ia .................................................................................. 5 9 

Hirtella .................................................................................. 79 

Acioa ................................................................................. .. 96 

N eocary a ................................................................................. 

9 7 

A cknow ledgm ents ............................................................................. 

97 

L iterature C ited ............................................................................... 

97 

Numerical List of Taxa, Including Notes on Rarity and Endangerment, 

with Index to D istribution M aps .............................................................. 99 

Supplemental List of Exsiccatae Studied Since 1972 Monograph .................................... 107 

Second Supplemental List of Exsiccatae .......................................................... 

124 

Distribution Maps ................................................................ .... 127 

Index of Local N am es .......................................................................... 263 

Index of Scientific N am es ...................................................................... 263 

ABSTRACT 

Since the Flora Neotropica account of the Chrysobalanaceae was published in 1972, much 

new material has been collected. This supplement to the earlier monograph has studied 

6170 new collections and includes the descriptions of 67 new taxa that have been described 

since the monograph, 64 of which are published as new species in this work. Full descriptions 

are given of all new species and they are numbered next to their closest relatives in the 

original monograph. New generic keys are provided to incorporate all the new species and 

other minor changes in taxonomy that were made in the light of the better material of many 

species. 

RESUMO 

Depois a publicacao da familia Chrysobalanaceae (1972) em Flora Neotropica muito 

material botanica foi coletada. Foram estudadas 6170 colecoes novas e descritas 67 taxa 

novas, incluido 64 apresentadas nesta obra. Describces de tudos esp6cies descritas desde 

1972 e chaves novas para cada genero sao foricidas. 

1New York Botanical Garden, Bronx, New York 10458, U.S.A. Present address: Director, Royal Botanic 

Gardens, Kew, Richmond, Surrey TW9 3AB, United Kingdom. 

1

2 Flora Neotropica 

INTRODUCTION 

This supplement to my "Flora Neotropica" 

monograph of the Chrysobalanaceae (Prance, 

1972) has become necessary because of the 

amount of collecting activity over the last decade 

and a half. The 6170 additional specimens studied 

have included 67 new taxa, numerous range 

extensions and other missing data for species 

which were described from incomplete material 

in 1972. The addition of the new taxa, as well 

as the increasingly recognized morphological 

variation of some previously poorly known 

species, means that the 1972 keys are out of date 

and do not function for some of the new material. 

I have, therefore, brought together all this new 

information into a supplement, which should be 

used in conjunction with the 1972 monograph. 

In 1972, 7762 specimens were studied, so a 

79.48% increase of the collections has occurred 

in a 16 year period. This has produced 64 new 

species, or one for every 96.4 collections, indicating 

the still poorly collected status of the neotropics. 

Most of the new taxa have already been published 

in a series of mainly regional papers (Berlin 

& Prance, 1978; Prance, 1973, 1974a, 1974b, 

1974c, 1976,1977a, 1979a, 1979b, 1979c, 1979d, 

1981, 1984, 1986a). Additional information has 

also been given in Prance (1975), and the Surinam 

species were treated in Prance and G6rts 

van Rijn (1976). The other purpose of this work 

is to publish the hitherto unpublished distribution 

maps of Chrysobalanaceae, most of which 

were not included in the original monograph, and 

from which extensive data for several biogeographical 

papers (for example, Mori et al., 1981; 

Prance, 1974d, 1977b, 1979e, 1982a; Prance & 

Mori, 1983) have been taken. Since the distribution 

maps are, together with distributional data 

from other families such as the Caryocaraceae 

tion about the family, and then presents data 

about each of the species for which significant 

new information has been gathered. These data 

include significant range extensions, description 

of organs not previously described, especially the 

fruit of several species, and any changes in cir- 

cumscription based on the new material. The 

new species are placed and numbered next to the 

species to which they are most closely related 

using a decimal point after the number. It is grat- 

ifying to note that while many new species have 

been found, they all fit extremely well into the 

genera as previously circumscribed. The only ge- 

neric change is in the circumscription of Acioa. 

This genus originally included the African species 

with ligular stamens, which we have now concluded 

(Prance & White, 1979) should be separated 

into the genus Dactyladenia. At the same 

time, in the Americas the circumscription of the 

genus cannot be confined to the three species 

with ligular stamens. We also concluded that in 

spite of its free stamens, Couepia edulis (Prance) 

Prance should be placed where I originally placed 

it before the flowers were known, in the genus 

Acioa. 

The amount of new information collected about 

Chrysobalanaceae in the last 15 years is a tribute 

to the collecting efforts of many people throughout 

the neotropics. It has enabled us to bring 

"Flora Neotropica" from a series based on very 

preliminary information to much more definitive 

treatments, which will certainly be reflected 

in the future monographs of the series. 

Keys are provided in the systematic section 

only for those genera in which there have been 

changes since 1972. 

Apart from an important contribution on the 

embryology of Chrysobalanaceae (Tobe & Raven, 

1984, see separate section), studies of the 

pollen (Demchenko, 1973; Patel et al., 1983) and 

various papers by this author, there have been 

and Dichapetalaceae, the basis for much of my few important papers on the family since those 

work on centers of endemism, it is important to reviewed in Prance (1972). However, some represent 

the maps here. The dots represent pres- gional treatments of the family, based on the 

ence on a degree square of latitude, rather than original monograph [Ecuador (Prance, 1979d), 

the exact locality of a collection. 

Venezuela (Prance, 1982b), Surinam (Prance & 

Although there are a large number of neotrop- Gorts van Rijn, 1976), and the Guianas (Prance, 

ical species of Chrysobalanaceae that are still in- 1986b)], also included considerably more maadequately 

collected, the family is much better terial than the 1972 monograph. Spichiger and 

known today than it was in 1970 when work on Masson's (1984) review of most of the Peruvian 

the first monograph was completed. This sup- species of Chrysobalanaceae, in an account of 

plement discusses briefly new general informa- the species in the Arboretum Jenaro Herrera in

Rare Species 3 

Loreto Department, was also largely based on thick; inner epidermis of ii developing into the 

the "Flora Neotropica" account. 

endothelium which directly borders the embryo 

The monographs of the 61 African species of sac and accumulates starch grains; micropyle 

Chrysobalanaceae have also allowed an inter- formed by both integuments. Fertilization poesting 

comparison between the two continents rogamous; endosperm formation of the Nuclear 

(Letouzey & White, 1976, 1978a, 1978b; White, type; seed exalbuminous. Young seed coat com- 

1976). This also led to the removal of the African posed of both testa and tegmen; exotegmen fispecies 

with a staminal ligule from the neotrop- brous; exotesta one-layered, composed of cuboid 

ical Acioa to a distinct genus Dactyladenia Welw. tanniniferous cells; endotesta a few layers thick 

(Prance & White, 1979). Likewise the species of composed of unspecialized small cells; micro- 

Malesian Chrysobalanaceae have recently been testa 30-50 cells thick in C. icaco, composed 

revised and their relationships to neotropical taxa mostly of vessels, 3-4 layered and lacking vasare 

now better understood (Prance, 1979b, 1987, cular tissue in L. michauxii. Mature seed mein 

press). Finally, a detailed worldwide treatment sotestal. 

of the genera of Chrysobalanaceae (Prance & The embryological data given above show that 

White, 1988) has been completed. 

there is little difference between the species studied. 

The Chrysobalanaceae differ from the Ro- 

EMBRYOLOGY 

saceae, to which it has frequently been related, 

in the tenuinucellate ovule, the very small nu- 

At the time of the 1972 monograph little de- cellus that disintegrates early, and the presence 

tailed work had been done on the embryology of of an endothelium. Tobe and Raven conducted 

the Chrysobalanaceae. Recently Tobe and Raven their investigation to consider the possible re- 

(1984) studied the embryology of three neotrop- lationship of the Chrysobalanaceae to the Myrical 

species-Chrysobalanus icaco L., Licania tales, but concluded that it is embryologically 

apetala (E. Mey.) Fritsch, and L. michauxii quite different from that well-defined order. They 

Prance. This has at least provided some basic proposed a relationship to the Theales, which 

data for the tribe Chrysobalaneae. The most im- share with Chrysobalanaceae a tenuinucellate 

portant embryological features are given below. ovule, a small disintegrating nucellus, an endo- 

Anthers tetrasporangiate, the walls five layers thelium, Nuclear-type endosperm formation, and 

thick, formation of the Basic type; epidermis an exalbuminous seed. However, the Theales difpersistent 

and often tanniniferous, endothecium fer from Chrysobalanaceae in their one-celled 

fibrous, the two middle layers ephemeral, the ovule, archesporium, an Allium-type embryo sac, 

tapetum glandular with two-nucleate cells. Cy- and persistent antipodal cells. In their argument 

tokinesis in the microspore mother cell simul- Tobe and Raven misrepresent the Chrysobalanataneous; 

microspore tetrad usually tetrahedral ceae as lacking stipules. Since there are major 

and occasionally decussate; pollen grains two- embryological differences between the Chrysocelled 

when shed. Ovule anatropous and tenui- balanaceae and both the Rosales and Theales, it 

nucellate, the nucellus very small; all archespo- seems unlikely that the family is any better placed 

rial cells developing directly into megaspore in the Theales than in the Rosales and a more 

mother cells, undergoing meiosis; megaspore definite placement can only be made after contetrads 

linear; chalazal megaspore functional, de- siderable data are obtained from other fields, as 

veloping into a modified type of the Polygonum- well as embryological information from a much 

type embryo sac; mature embryo sac four-nu- wider range of Chrysobalanaceae, instead ofjust 

cleate, comprising an egg, two synergids and a three species in the two most closely related gencentral 

nucleus which is probably triploid; an- era. 

tipodal cells absent from the beginning, hypostase 

differentiating in Chrysobalanus but not Li- 

RARE SPECIES 

cania; the nucellar tissue soon disintegrating 

except the megaspore and embryo sac. Ovule At the 1986 meeting of the Organization for 

bitegmic, the inner integument (ii) and outer in- Flora Neotropica it was recommended that 

tegument (oi) initiated desmally, the ii up to five monographers comment on and point out rare 

or eight cells thick and the oi more than five cells and endangered species in their groups. Although

4 

Flora Neotropica 

our knowledge of tropical groups such as Chryso- collecting activity. The endangerment status used 

balanaceae is far from adequate, the collection in the list is based on the collection record and 

data are now certainly sufficient to pick out a few on the current condition of the habitat in which 

of the rarest species. All rare species are marked they grow. Poorly known species from relatively 

in the species list preceding the list of exsiccatae, undisturbed areas such as the Guianas are not 

and it gives the rarest and possibly most endan- generally listed unless they are known only from 

gered. The species listed as rare include all those a single collection. 

known from a single collection and not re-col- I consider as endangered 31 of the 100 rarest 

lected since 1972 as well as others that appear species. These are mainly from areas that are foci 

to be particularly threatened by virtue of their of forest destruction such as Central America, 

habitat. Species described since 1972 and known Choc6, Rond6nia, and eastern Brazil. The fact 

from a single collection are only listed if they are that there are at least 100 rare species, 31 of 

from particularly threatened areas such as At- which are endangered, shows the desperate need 

lantic coastal forests of Brazil or from Rond6nia, for a greater conservation effort of the neotropas 

it is too early to judge their frequency from ical forests. 

Revised Key to Species of Chrysobalanus 

1. Leaves orbicular to ovate-elliptic, the apex retuse 

to rounded or minutely and bluntly acuminate; 

midrib glabrous beneath. 1. C. icaco. 

1. Leaves elliptic to oblong, with a distinct acumen 

5-15 mm long; midrib sparsely hirsute beneath. 

2. Leaves chartaceous, acuminate; branches 

conspicuously lenticellate; inflorescence graypuberulous. 

2. C. cuspidatus. 

2. Leaves thickly coriaceous, caudate; branches 

not conspicuously lenticellate; inflorescence 

ferrugineous. 

3. C. venezuelanus. 

1-1. Chrysobalanus icaco Linnaeus, Sp. pl. 1:513. 

1753. 

Chrysobalanus interior Small, Man. s. e. fl. 645. 1933. 

Type. U.S.A. Florida: Hammocks, Long Key, Everglades, 

Small & Carter 3166 (lectotype, here designated, 

NY). 

The synonym C. interior was overlooked in 

Prance (1972) and in the treatment for the ge- 

neric flora of the southeastern U.S. (Prance, 1970). 

This name, although validly published, was not 

picked up in either the "Index Kewensis" or the 

"Gray Herbarium Card Index." There are no 

significant differences and C. interior falls well 

within the range of variation of C. icaco as cir- 

cumscribed in Prance (1972). 

The distribution of C. icaco will be found in 

Figure 20.2 

2 Figures 20 to 155 are grouped at the end of the 

monograph. 

SYSTEMATIC TREATMENT 

1. Chrysobalanus Linnaeus 

1-3. Chrysobalanus venezuelanus Prance, sp. 

nov. Type. Venezuela. Bolivar: Icabarui River 

region, headwater of Rio Hacha, 450-850 m, 

3 Jan 1956 (fl), A. L. Bernardi 2777 (holotype, 

NY; isotype, NY). Fig. 1. 

Species a C. cuspidato foliis coriaceis, latioribus, 

ramulis juvenilibus haud conspicue lenticellatis, 

inflorescentiis ferrugineo-pubescentibus 

differt. 

Tree to 10 m tall, the young branches sparsely 

puberulous to glabrescent. Leaf lamina oblongelliptic, 

coriaceous, 5.5-11 x 2.2-4.2 cm, cuneate 

at base, caudate at apex, the acumen 10- 

17 mm long, glabrous and shiny above, glabrous 

beneath except for appressed hairs on midrib and 

primary veins of lower surface of very young 

leaves; midrib prominulous above, prominent 

beneath; primary veins 6-8 pairs, widely spaced, 

prominulous on both surfaces; petioles 2-4 mm 

long, terete, rugulose, glabrous. Stipules, caducous, 

membranous, axillary. Inflorescences of 

few-flowered cymules inserted on rachis to 5 cm 

long, the rachis and branches ferrugineous-pubescent; 

bracts and bracteoles ovate, membranous, 

puberulous on exterior, ca. 1 mm long, 

caducous. Flowers 3 mm long, inserted in small 

cymules; receptacle campanulate, tomentellous 

on exterior, tomentose within; calyx lobes rounded, 

tomentellous; petals five, ovate, glabrous; stamens 

14-15, inserted around complete circle, the 

filaments shortly exserted, pubescent; ovary in-

Systematic Treatment 5 

Gfwysobarlana 

,C~ 0 

r ?I 

a( cmj 

i ~~~~~~~~~~~~~~~~~v10 

I~~~~~~~~~~~~~~~~~ 

/Th II h4.?-" 

'A ~ ~ ~ A 

)vc : . 

-V~~~~ 

FIG. 1. Chrysobalanus venezuelanus (Bernardi 2777). A, habit; B, leaf undersurface; C, flower buds; D, 

flower; E, flower section; F, ovary and style; G, petal; H, young fruit. 

serted at base of receptacle, lanate; style equalling 

filaments in length. Fruit (only seen young) oblong, 

deeply costate; exocarp glabrous except 

when very young. 

Distribution (Fig. 21). Forest on terra firme in 

eastern Venezuela. 

Additional 

specimens examined. VENEZUELA. 

BOLiVAR: W base of Amaruay-tepui, 5?56'N, 62?17'W, 

5 May 1986 (fl), Hoist & Liesner 2788 (MO, NY); 4 

km W of El Pauji, Rio Chaberu, 750-900 m, 12 Nov 

1985 (y fr), Liesner 19919 (MO, NY); Distr. Piar, Rio 

Aparaman, SW corer of Amaruay-tepui, 5?54'N, 

62?15'W, 22 Apr 1986 (fl), Liesner & Hoist 20192 (MO, 

NY); Quebrada Los Brasileros, 4.5 km SW of Icabaru, 

4?20'N, 61?48'W, 480 m, 16 Dec 1978 (fl), Steyermark 

et al. 117696 (NY, VEN). 

This species differs from the closely related 

.

6 

Chrysobalanus cuspidatus (Fig. 21) of the Lesser 

Antilles in the more robust inflorescence branches 

with a ferrugineous, not gray pubescence, the 

broader, more coriaceous leaves, and the less 

conspicuously lenticellate young branches. This 

is a most interesting occurrence of the genus 

Chrysobalanus since it is from inland continental 

2. Licania Aublet 

Revised Key to Species of Licania3 


South America. The genus is predominantly 

coastal (C. icaco) and from cloud forest in the 

Lesser Antilles (C. cuspidatus). The new species, 

C. venezuelanus, is found at elevations between 

450 and 850 m and is thus in a habitat very 

similar to that of C. cuspidatus in the montane 

forests of Guadeloupe and Martinique. 

Since the genus is large, now containing 190 species, the key is divided for convenience into three 

parts. Note: Species 1-152 are fully described in Prance (1972) and the 38 species described since 

1972 or in this work have complete descriptions following these keys. 

1. Stamens longer than calyx lobes. Key A: Subgenus Moquilea sections Moquilea and Leptobalanus. 

1. Stamens equal to or shorter than calyx lobes. 

2. Stamens 10-25. Key B: Subgenus Moquilea sections Microdesmia and 

Leptobalanus pro parte; subgenus Parinariopsis. 

2. Stamens 3-9. Key C: Subgenus Licania. 

Key A 

Species with 8-60 stamens, exserted beyond calyx lobes. 

1. Petals present; stamens 11-60. (For contrasting statement see p. 9.) Subg. Moquilea sect. Moquilea. 

2. Leaf underside with an appressed lanate-arachnoid pubescence. 

3. Inflorescence of fasciculate, short, dense-flowered racemes; primary veins impressed above (Panama). 

5.3. L. fasciculata. 

3. Inflorescence of lax, branched, racemose panicles or cymose panicles; primary veins usually 

prominulous or plane above. 

4. Primary leaf veins 8-15 pairs, the lamina oblong-lanceolate. 

5. Petioles 1-3 mm long; fruit 2.5-3 mm long, with smooth exterior, pericarp thin; inflorescence 

of small cymose panicles (SE U.S.A.). 

1. L. michauxii. 

5. Petioles 5-7 mm long; fruit 4-7 mm long, with a crustaceous, verrucose exterior, pericarp 

thick; inflorescence of racemose panicles (Bolivia, La Paz). 

2. L. boliviensis. 

4. Primary leaf veins 16-29 pairs, the lamina oblong to elliptic. 

6. Inflorescence ramiflorous, borne on young woody twigs well below shoot apex. 

7. Flowers (calyx and receptacle) 6-7 mm long, young leafundersurface with shaggy dense 

ferrugineous pubescence; leaves 27-47 cm long (Colombia, Valle, 0-50 m). 

5.1. L. gentryi. 

7. Flowers 2-5 mm long, leaf undersurface with compact appressed pubescence or with 

caducous pubescence, becoming glabrous when mature; leaves 16-33 cm long. 

8. Leaf undersurface glabrous when mature, with a lanate caducous pubescence on 

youngest leaves; flowers ca. 2 mm long (Colombia, Valle, 0-50 m). 46. L. velata. 

8. Leafundersurface with a compact persistent gray-lanate or brown pubescence; flowers 

3-5 mm long. 

9. Leaf undersurface ferrugineous to brown-tomentose, secondary venation not 

conspicuously reticulate (Colombia; Ecuador; Peru). 5. L. macrocarpa. 

9. Leaf undersurface gray-tomentellous, conspicuously reticulate with parallel secondary 

venation; leaves thickly coriaceous. 

10. Inflorescence 15-30 cm long, much-branched, flowers densely packed, bracteoles 

caducous (Colombia; Ecuador). 

4. L. durifolia. 

10. Inflorescence 8-15 cm long, unbranched or with a few branches; bracteoles 

persistent, membranous (Ecuador). 

4.2. L. grandibracteata. 

3 Note that Licania cecidiophora, L. mexicana, and L. tepuiensis are not included in these keys since they 

were described from incomplete material.


6. Inflorescence terminal or axillary, but on leafy shoot apex. 

11. Flowers 1.5-2 mm long; leaves 12-19 x 3-6 cm, brown-pubescent beneath, without 

conspicuously reticulate venation (Colombia). 

148. L. veneralensis. 

11. Flowers 3-5 mm long; leaves 15-35 x 6-13 cm, gray-pubescent beneath, often with 

conspicuously reticulate parallel tertiary venation. 

12. Inflorescence and flowers with light-brown-tomentellous pubescence, leaves gradually 

tapering from mid-point to acuminate apex; center primary veins far apart 

(18-22 mm), the secondary venation obscure (Colombia, Choc6). 3. L. maritima. 

12. Inflorescence and flowers with ferrugineous pubescence; leaves tapering from well 

above midpoint, center primary veins 5-13 mm apart, with conspicuous parallel 

secondary venation. 

13. Leaves 6-9.5 x 2-3 cm; petioles 4-5 mm long (Venezuela, Lara). 

5.4. L. montana. 

13. Leaves 12-46 x 3-26 cm; petioles 6-12 mm long. 

14. Inflorescences of racemose panicles, to 18 cm long; petioles 10-12 mm 

long; primary veins 16-19 pairs, pedicels 5 mm long (Colombia, Antioquia). 

5.2. L. cabrerae. 

14. Inflorescences unbranched racemes, or little-branched panicles, 5-13 cm 

long. Petioles 6-8 mm long, primary veins 21-24 pairs; flowers sessile 

(Peru, San Martin). 

4.1. L. filomenoi. 

2. Leaf underside glabrous when mature (sometimes lanate when young). 

15. Inflorescence and exterior of flowers glabrous or sparsely puberulous. 

16. Leaf broadly acuminate, the acumen 4-8 mm long; flowers 2.5-3 mm long; pedicels 2-5 

mm long (Brazil, Bahia). 

6. L. salzmannii. 

16. Leaf with well-developed finely pointed acumen 8-12 mm long; flowers 4-4.5 mm long; 

pedicels 1-3 mm long. 

17. Inflorescence and exterior of flowers puberulous; petioles 10-12 mm long; leaves elliptic, 

5.5-9 cm broad (Peru, Loreto). 

7. L. klugii. 

17. Inflorescence and exterior of flowers glabrescent; petioles 5-7(-9) mm long; leaves 

oblong to oblong-elliptic, 2-5.5 cm broad (Guianas; Peruvian Amazonia). 

8. L. guianensis. 

15. Exterior of flowers and usually inflorescence densely tomentose. 

18. Inflorescence either of dense pyramidal or cymose panicles with a glabrous to puberulous 

rachis, or a much branched panicle with the primary branches branched into small branches 

bearing 2 or more flowers. 

19. Inflorescence a much branched panicle 10-25 cm long, with the primary branches 

branched into small branches; leaves 5.5-10.5 cm broad (Ecuador). 13.1. L. dodsonii. 

19. Inflorescence of dense pyramidal or rather lax cymose panicles 1-5 cm long; leaves 1- 

4.5 cm broad. 

20. Inflorescence short and dense, the rachis glabrous; petioles 4-6 mm long (Mexico). 

9. L. retifolia. 

20. Inflorescence lax, the rachis puberulous; petioles 1-3 mm long (SE U.S.A.). 

1. L. michauxii. 

18. Inflorescence of lax racemes or racemose panicles, the rachis pubescent. 

21. Pedicels 3-7 mm long; leaves with finely pointed acumen 1.5-2.5 cm long (Brazil, 

Amazonas). 

10. L. longipedicellata. 

21. Pedicels 0.5-2.5 mm long; leaf acumen rather blunt, less than 10 mm long. 

22. Inflorescence of predominantly axillary racemes or few-branched panicles (NE 

Brazil). 

11. L. tomentosa. 

22. Inflorescence of predominantly terminal much-branched panicles or racemose panicles, 

rarely axillary, then much branched. 

23. Leaves with a lanate pubescence when young; petioles 8-16 mm long and very 

slender, not more than 1.2 mm thick (W Indies; Venezuela). 12. L. pyrifolia. 

23. Leaves glabrous even when young; petioles 3-12 mm long, stout, more than 

1.5 mm thick. 

24. Leaf bases cuneate; stamens ca. 30; petioles canaliculate (W Indies; Venezuela). 

13. L. leucosepala. 

24. Leaf bases cordate, rounded or subcuneate; stamens 12-23; petioles usually 

terete, rarely canaliculate. 

25. Leaves oblong-lanceolate, primary veins slightly impressed above. 

26. Petioles 10-13 mm long; leaves 10-14.5 x 2.5-5.2 cm; primary 

veins 11-17 pairs; stamens 17 (Peru; Brazil, W Amazonia). 

14. L. angusataa.


26. Petioles 5-7 mm long; leaves 14-25 cm long x 4.5-8 cm; primary 

veins 18-22 pairs; stamens 12-14 (Brazil, Para). 

14.1. L. anneae. 

25. Leaves ovate to oblong (to oblong-lanceolate in L. platypus); primary 

veins plane or prominulous above. 

27. Leaves 13-30 cm long; primary veins 13-22 pairs, leaf bases 

sometimes subcordate. 

28. Flowers borne in small cymules on short tertiary branches; 

petioles 12-16 mm long, canaliculate (Venezuela). 

23.1. L. tachirensis. 

28. Flowers borne on primary inflorescence branches; petioles 

8-14 mm long, terete. 

29. Leaves oblong to oblong-lanceolate, 3.5-8 cm broad; 

primary veins 15-22 pairs; petioles 9-14 mm long; leaf 

base rounded to subcuneate (Central America). 

15. L. platypus. 

29. Leaves elliptic, 7.5-9.5 cm broad; primary veins 14- 

15 pairs; petioles ca. 8 mm long; leaf base subcordate 

(Brazil, Maranhao). 

19. L. maranhensis. 

27. Leaves 5-15.5 cm long (to 21 cm in L. guatemalensis); primary 

veins 8-13 pairs (to 15 in L. britteniana); leaf bases subcuneate 

to rounded (to subcordate in L. britteniana). 

30. Petals not unguiculate, equalling or only shortly exceeding 

calyx lobes, filaments far exceeding petals. 

31. Flowers 4-5 mm long; stipules linear-filamentous, 5-7 

mm long (Mexico). 

16. L. gonzalezii. 

31. Flowers 2-3.5 mm long; stipules short-subulate, to 2 

mm long. 

32. Petioles 2-5 mm long; leaves obovate or elliptic, 

with cuspidate, mucronate, or abrupt apex. 

33. Inflorescence once-branched; leaves obovate 

to elliptic, 4.5-8.5 cm long (Brazil; Colombia; 

Peru, Amazonia). 

17. L. egleri. 

33. Inflorescence twice-branched; the flowers borne 

in small groups on short secondary branches; 

leaves elliptic, 10-21 cm long (Guatemala). 

18.3. L. guatemalensis. 

32. Petioles 6-14 mm long; leaves oblong to oblongelliptic, 

apex gradually acuminate (cuspidate in L. 

kallunkiae). 

34. Inflorescencerachis and branches sparselygraypubescent. 

35. Leaf apex cuspidate, the base subcuneate; 

stamens 11-12 (Panama). 

18.2. L. kallunkiae. 

35. Leaf apex acuminate, the base rounded; 

stamens 16-25 (Brazil, Amazonia; Peru). 

20. L. fritschii. 

34. Inflorescence rachis and branches tomentose. 

36. Inflorescence small and little-branched; 

flowers 3-4 mm long; ovary glabrous; petioles 

glabrous (Panama). 

18.1. L. chiriquiensis. 

36. Inflorescence large and much-branched; 

flowers 2-3 mm long; ovary pubescent; 

petioles tomentose when young. 

37. Flower buds globose; flowers 2 mm 

long; leaves 5.5-11 x 2-6 cm 

(Guianas; Brazil; Venezuela). 

18. L. minutiflora. 

37. Flower buds ovoid; flowers 2.5-3 mm 

long; leaves 8-16 x 3-7.5 cm (Colombia; 

Peru; Brazil, Amazonia). 

21. L. britteniana.


30. Petals unguiculate, much exceeding calyx lobes; filaments 

scarcely exceeding petals. 

38. Flowers 5-6 mm long; pedicels 2-2.5 mm long; ovary 

pilose (Brazil, Amazonia). 

22. L. unguiculata. 

38. Flowers 3 mm long; pedicels 0-0.25 mm long; ovary 

glabrescent (Peru; Brazil, Amazonia). 23. L. longipetala. 

1. Petals absent; stamens 8-14. Subg. Moquilea sect. Leptobalanus. 

39. Leaf underside glabrous or lanate, the venation not prominent, the stomatal cavities absent. 

40. Most flowers borne in small groups or cymules on distinct secondary branches (peduncles) of 

inflorescence; peduncles over 2 mm long. 

41. Rachis of inflorescence densely yellow-villous-tomentose (Venezuela; Brazil, Amazonia). 

24. L. wurdackii. 

41. Rachis of inflorescence gray-puberulous. 

42. Inflorescence short and compact, 3-5 cm long; flowers 4-5 mm long; leaves ovate 

(NE Brazil). 

25. L. turbinata. 

42. Inflorescence longer and spreading, 6-20 cm long; flowers 2-3.5 mm long; leaves 

oblong to elliptic. 

43. Flowers 3.5-5 mm long; petioles deeply canaliculate; with two prominent glands 

at lamina base; rachis arachnoid-pubescent when young (Amazonia). 26. L. lata. 

43. Flowers 2-3 mm long; petioles terete to shallowly canaliculate; lamina base eglandular; 

rachis puberulous (Guianas; Brazil to S Amazonia). 27. L. apetala. 

40. Flowers sessile or nearly so on primary branches of inflorescence. 

44. Leaf apex predominantly obtuse to rounded. 

45. Rachis of inflorescence densely villous-tomentose; flowers ca. 4 mm long (Venezuela; 

Brazil, NW Amazonia). 

24. L. wurdackii. 

45. Rachis of inflorescence gray-puberulous; flowers 3 mm long (Guianas; Brazil, Amazonia). 

28. L. parvifolia. 

44. Leaf apex distinctly acuminate (rarely only bluntly acuminate, L. jefensis). 

46. Flowers 4-5 mm long; receptacle cupuliform. 

47. Leaves glabrous beneath, inflorescence gray-villous-tomentose (Brazil, Mato 

Grosso). 

29. L. maguirei. 

47. Leaves with short persistent waxy lanate-pulverulent pubescence beneath; inflorescence 

brown-puberulous to tomentellous when mature (Amazonian Colombia; 

Peru; Brazil). 

26. L. lata. 

46. Flowers 2-3.5 mm long; receptacle urceolate or campanulate. 

48. Stipules persistent on young branches; petioles densely tomentose when young, 

2.5-3.5 mm thick; upper portion of ovary glabrous (central Brazil). 30. L. gardneri. 

48. Stipules usually caducous; petioles sparsely lanate-pubescent, puberulous, or glabrous, 

to 2 mm thick; upper portion of ovary pubescent. 

49. Inflorescence sparsely hirsutulous or yellow-arachnoid-tomentose; flower exterior 

yellow-brown-tomentose; petioles 2-3 mm long, canaliculate. 

50. Inflorescence sparsely hirsutulous; leaf apex finely acuminate, the acumen 

5-10 mm long (N Colombia). 

31. L. cuspidata. 

50. Inflorescence arachnoid-tomentose; leaf apex acute or bluntly acuminate, 

the acumen 3-5 mm long (Panama). 

31.1. L. jefensis. 

49. Inflorescence puberulous to tomentose; flower exterior gray-tomentose; petioles 

3-11 mm long, terete or shallowly canaliculate towards base. 

51. Leaf bases subcordate (Panama). 

31.2. L. morii. 

51. Leaf bases cuneate or subcuneate to rounded. 

52. Leaf undersurface rufous-brown-pubescent, with deeply reticulate 

venation; petioles 8-11 mm long; leaf apex cuspidate; inflorescence 

brown-tomentellous (Colombia, Valle). 31.3. L. cuatrecasasii. 

52. Leaf undersurface glabrous or rarely white-arachnoid-pubescent, venation 

smooth, not deeply reticulate; petioles 3-8 mm long; leaf 

apex acuminate; inflorescence brown or gray-puberulous to tomentose. 

53. Rachis and branches of inflorescence tomentose; petioles usually 

bearing two distinct glands (Central America). 

32. L. sparsipilis. 

53. Rachis and branches of inflorescence puberulous; petioles 

eglandular. 

54. Inflorescence and flower exterior sparsely puberulous; leaves 

thickly coriaceous, the apex finely attenuate with acumen 

10-25 mm long (Guianas; N Amazonian Brazil). 

27.1. L. granvillei.


54. Inflorescence usually densely gray-puberulous; leaves chartaceous, 

the apex acuminate with acumen 3-15 mm long 

(Colombia; Peru; Venezuela to S Brazil). 27. L. apetala. 

39. Leafundersurface with stomatal cavities or very deeply cut reticulation resembling stomatal cavities, 

lanate-pubescent in mouth of cavities. 

55. Inflorescence with secondary branches; flowers on distinct secondary branches of inflorescence 

that are at least 2 mm long. 

56. Flowers in small distinct groups or in cymules on short secondary branches (peduncles); 

leaf upper surface smooth, the base rounded to cuneate. 

57. Inflorescence 15-22 cm long, ferrugineous-arachnoid; leaf underside brown when dry 

(Guyana). 

35. L. persaudii. 

57. Inflorescence 6-8 cm long, gray-arachnoid or puberulous; leaf underside gray when 

dry (Guianas; Brazil, N Amazonia). 

36. L. sprucei. 

56. Flowers more or less clustered in large groups on primary and secondary branches of 

inflorescence, but without distinct peduncles; leaf upper surface usually papillose, the base 

often cordate (Brazil, Amazonia and south). 

37. L. sclerophylla. 

55. Inflorescence with primary branches only; flowers on primary branches of inflorescence, sessile 

or rarely with short peduncles not exceeding 1 mm in length. 

58. Pedicels 2 mm long (in bud), to 4 mm in flower (Guianas). 

38. L. albiflora. 

58. Pedicels not exceeding 2 mm, often shorter. 

59. Leaf reticulation extremely prominent on upper surface; fruit exterior appressedsordid-yellow-velutinous. 

60. Petioles 5-13 mm long; flowers ca. 3 mm long, borne on slender branchlets 0.5- 

1 mm thick; receptacle campanulate; flowers and inflorescence gray-tomentose 

(Panama; Venezuela; Guianas; Brazil, Amazonia). 

39. L. longistyla. 

60. Petioles 15-20 mm long; flowers ca. 4 mm long, borne on branchlets ca. 2 mm 

thick; receptacle cupuliform; inflorescence and flowers brown-tomentose (Colombia, 

Pacific). 

40. L. fuchsii. 

59. Leaf reticulation prominulous on upper surface; fruit exterior glabrous or browntomentose. 

61. Young branches densely tomentellous, with a thick corky bark (central Brazil). 

41. L. humilis. 

61. Young branches glabrous or nearly so, the bark thin. 

62. Flowers 3.5-5 mm long; leaf apex mucronate (Guyana). 42. L. foveolata. 

62. Flowers 2-3 mm long; leaf apex rounded to acuminate (Venezuela; Guianas; 

central and Amazonian Brazil). 

43. L. octandra. 

Key B 

Species with 10-25 stamens, equalling or shorter than calyx lobes. 

1. Petals present. 

2. Ovary inserted at base of receptacle; bracts and bracteoles small and not enclosing groups of flower 

buds. Subg. Moquilea sect. Microdesmia. 

3. Leaf underside with stomatal cavities or deeply cut reticulation; filaments connate over half of 

length in groups, densely pubescent throughout. 

4. Pubescence of inflorescence light gray; primary leaf veins prominulous above (NE Brazil). 

44. L. rigida. 

4. Pubescence of inflorescence ferrugineous; primary leaf veins slightly impressed above (Central 

America; W South America to Peru). 45. L. arborea. 

3. Leaf underside with inconspicuous venation, lacking stomatal cavities; filaments free almost to 

base, glabrous on upper portion. 

5. Leaves 15-36 cm long; stipules 12-22 mm long. 

6. Leaf undersurface with caducous, lanate pubescence; anthers reniform; inflorescence borne 

on woody defoliated stalks (Colombia, Valle). 46. L. velata. 

6. Leaf undersurface hirsute; anthers deltoid; inflorescence terminal (French Guiana; Brazil, 

Amapa). 149. L. amapaensis. 

5. Leaves less than 14 cm long; stipules 2-10 mm long; flowers in terminal or axillary inflorescences 

on young branchlets. 

7. Inflorescence of unbranched subspikes or racemes. 

8. Flowers ca. 1.5 mm long, borne in clusters along rachis, the rachis, 4-15 mm long 

(Colombia, Boyaca). 47. L. subarachnophylla. 

8. Flowers ca. 2.5 mm long, not clustered, the rachis 2-4 mm long (Peru, Madre dos 

Dios). 47.1. L. tambopatensis.


7. Inflorescence of panicles, or a panicle of cymules. 

9. Leaf underside densely lanate-pubescent. 

10. Leaf apex with a well-developed acumen 3-5 mm long; stamens ca. 25; exterior 

of receptacle ferrugineous (Colombia, Antioquia). 

48. L. salicifolia. 

10. Leaf apex obtuse to broadly acuminate, the acumen not exceeding 1 mm; stamens 

ca. 15; exterior of receptacle brown (Brazil, Goifs). 

49. L. araneosa. 

9. Leaf underside glabrous. 

11. Inflorescence a panicle of small cymules along the rachis and short secondary 

branches; petioles 2-5 mm long; stamens 8-10 (Old World). 

Sect. Hymenopus. 147. L. splendens, L. palawanensis. 

11. Inflorescence of panicles, flowers not grouped in cymules; petioles 6-11 mm long; 

stamens 14-25 (New World). 

12. Leaf acumen 3-7 mm long; leaves 5-10 cm long; stamens ca. 25; flowers 

sessile on primary branches of inflorescence, densely brown-tomentose (eastern 

Brazil). 

50. L. silvatica. 

12. Leaf acumen 10-15 mm long; leaves 7.5-14 cm long; stamens 12-14; flowers 

predominantly on short secondary branches of inflorescence, short gray-to- 

mentose (Colombia, Valle). 

51. L. chocoensis. 

2. Ovary inserted laterally; bracts and bracteoles large and enclosing small groups of flower buds 

(Venezuela; Guianas; Brazil, Amazonia). Subg. Parinariopsis. 52. L. licaniiflora. 

1. Petals absent. 

13. Leaf underside glabrous, or with an easily removed lanate pubescence, the reticulations shallow, 

stomatal cavities absent. Subg. Moquilea sect. Leptobalanus. 

14. Leaf undersurface glabrous; flowers in groups on distinct secondary branches of inflorescence; 

stamens 10-19 (Amazonia). 

15. Stamens ca. 19; leaves 15-20 cm long, with long acumen 11-19 mm long; flowers ca. 5 

mm long (Brazil, N Amazonia). 

33.1. L. joseramosii. 

15. Stamens 10-11; leaves 3-8.5 cm long, with acumen 0-6 mm long; flowers 2-3 mm long 

(Venezuela; Brazil, Amazonia). 

33. L. emarginata. 

14. Leaf underside lanate; flowers sessile on primary branches of inflorescence; stamens ca. 22 

(Colombia, Valle). 

34. L. calvescens. 

13. Leaf underside with stomatal cavities or with deep reticulations, pubescence dense and filling the 

hollows between venation. Subg. Licania sect. Licania (pro parte). 

16. Receptacle conical; petioles usually canaliculate, 8-13 mm long; midrib impressed on upper 

surface towards lamina base; leaves drying light brown (Guianas; Brazil, N Amazonia). 

108. L. majuscula. 

16. Receptacle barrel-shaped; petioles terete, 3-6 mm long; midrib prominulous at base above, 

but becoming impressed apically; leaves drying dark brown (Colombia; Peru; Venezuela; Brazil, 

Amazonia). 

140. L. mollis. 

Key C 

Species with 3-9 stamens, equalling or shorter than calyx lobes. 

1. Leaf underside glabrous or sparsely hirsute or hispid along venation; petals present or absent. (For 

contrasting statement see p. 13.) 

2. Petals present. (For contrasting statement see p. 13.) 

3. Leaves sparsely hirsute along venation of lower surface or hispid at least on midrib. Sect. Hirsuta. 

4. Inflorescence and flowers sparsely hirsutulous; leaf lamina appearing deeply rugose above; 

primary veins and venation deeply impressed on upper surface. 

5. Leaf oblong, apex acuminate (Brazil, Amazonia). 

53. L. hirsuta. 

5. Leaf ovate to ovate-elliptic, apex rounded to obtuse (Central America). 54. L. costaricensis. 

4. Inflorescence and flowers densely pubescent; leaf lamina not rugose, primary veins plane or 

impressed and venation more or less plane on upper surface. 

6. Young stems, leaf midrib, and lower inflorescence branches hispid (Venezuela). 

57.1. L. hispida. 

6. Young stems puberulous; leaf midrib hirsute; inflorescence and exterior of flowers with 

short gray pubescence or densely ferrugineous-tomentose. 

7. Inflorescence and exterior of flowers with a short gray pubescence (Trinidad; Venezuela; 

Guianas; Brazil, Amazonia). 

69. L. heteromorpha. 

7. Inflorescence and exterior of flowers densely ferrugineous-tomentose. 

8. Leaf chartaceous, apex acuminate (Bolivia, Amazonia). 

55. L. krukovii. 

8. Leaf thick-coriaceous, apex rounded to apiculate.


9. Primary leaf veins 6-9, prominulous above; petioles 3-4 mm long (Guyana; 

Venezuela). 

56. L. lasseri. 

9. Primary leaf veins 12-20, slightly impressed above; petioles 7-15 mm long 

(Guyana; Brazil, Amazonia). 

57. L. latifolia. 

3. Leaf underside glabrous. 

Sect. Hymenopus. 

10. Flowers in small cymules on short distinct secondary branches of inflorescence at least 1 mm long. 

11. Leaves with a finely pointed acumen 15-30 mm long; flowers ca. 1 mm long; ovary and style 

pubescent (Colombia, Valle). 

58. L. minuscula. 

11. Leaves rounded or acute to broadly acuminate, the acumen 2-8.8 mm long; flowers 2-5 mm 

long; ovary and style glabrous or pubescent. 

12. Inflorescence compact and triangular, to 8 cm long; rachis and branches glabrous, the 

rachis not lenticellate; peduncles 2-4 mm long; exterior of flowers glabrous (Central America). 

59. L. operculipetala. 

12. Inflorescence lax and spreading, over 8 cm long, or short, lax and of small cymules on 

central rachis; rachis and branches puberulous, the rachis often lenticellate; peduncles 0.5- 

2 mm long; exterior of flowers puberulous. 

13. Inflorescence of small cymules on a central rachis 1.5-10 cm long; stamens 8-10; 

receptacle oblique in bud; mature fruit to 1.5 cm long, usually smaller, not costate 

(Old World, Asia and Pacific). 

147. L. splendens, L. palawanensis.4 

13. Inflorescence lax and spreading, over 8 cm long; stamens 5-7; receptacle symmetrical 

in bud; mature fruit 1.5-4 cm long, costate (New World). 

14. Flowers 4-5 mm long; inflorescence rachis 2-3 mm thick; leaves thickly coriaceous 

(Venezuela). 

60.1. L. pakaraimensis. 

14. Flowers 1.5-3 mm long; inflorescence rachis 1 mm thick; leaves chartaceous or 

thinly coriaceous. 

15. Leaf apex rounded; reticulation intricate (Trinidad; Venezuela; Guianas; Brazil, 

Amazonia). 


15. Leaf apex acuminate; reticulation lax (Amazonia). 60. L. reticulata. 

10. Flowers borne mainly on primary branches of inflorescence or on secondary branches, but not in 

pedunculate cymules. 

16. Leaves narrowly oblong with nearly parallel sides; stamens usually slightly exceeding calyx 

lobes; mouth of receptacle filled by a dense lanate mass; petals slightly unguiculate. 

17. Receptacle narrowly urceolate, densely arachnoid-pubescent; flowers in dense glomerules 

on primary branches (Guyana; Brazil, Amazonia; Peru). 

61. L. arachnoidea. 

17. Receptacle campanulate, puberulous; flowers not densely glomerulate. 

18. Flowers 1.5-2 mm long; leaves to 17 cm long, usually smaller; stipules to 5 mm, 

caducous (Brazil, Amazonia). 

62. L. oblongifolia. 

18. Flowers 2.5-3 mm long; leaves usually exceeding 16 cm in length (to 40 cm); stipules 

to 15 mm, subpersistent (Guianas; Brazil, E Amazonia). 63. L. macrophylla. 

16. Leaves ovate to oblong-lanceolate, but sides converging; stamens shorter than calyx lobes; 

mouth of receptacle with short deflexed hairs only; petals with broad simple bases. 

19. Exterior of flowers and rachis and branches of inflorescence glabrous or sparsely hirsutulous. 

20. Leaf apex caudate to cuspidate. 

21. Leaves elliptic, coriaceous, the apex caudate; petioles 5-7 mm long (Fr. Guiana; 

Brazil, Amazonia; Colombia; Peru). 

64. L. caudata. 

21. Leaves narrowly oblong, chartaceous, the apex cuspidate; petioles 1-3 mm long 

(Brazil, Mato Grosso). 

64.1. L. miltonii. 

20. Leaf apex acute to acuminate. 

22. Stipules large and foliaceous, caducous; exterior of flowers glabrous (Venezuela). 

65. L. latistipula. 

22. Stipules small, lanceolate, persistent; exterior of flowers hirsutulous. 

23. Leaves 9-27 cm long, thick-coriaceous, the apex abruptly short-acuminate; 

primary veins plane above (Guianas; Brazil, Parf). 66. L. divaricata. 

23. Leaves 7-11 cm long, membranous, the apex with a well-developed acumen 

5-9 mm long; primary veins slightly impressed above (Costa Rica; Venezuela; 

Brazil, Para). 67. L. glabriflora. 

19. Exterior of flowers and usually rachis and branches of inflorescence densely puberulous 

to tomentose; pubescence completely covering exterior of calyx. 

4 A third Malesian species of Licania, L. fusicarpa (Kosterm.) Prance, probably belongs here, but is known 

only from fruiting material.


24. Midrib broad towards base, 2-3.5 mm thick; leaves very thick-coriaceous; stipules to 

15 mm long, subpersistent; anthers deltoid or nearly so (Venezuela; Guyana; Brazil, 

Amazon). 

68. L. intrapetiolaris. 

24. Midrib narrower towards base, 1-2 mm thick; leaves membranous to coriaceous; 

stipules to 8 mm long, persistent to caducous; anthers reniform. 

25. Leaf apex long acuminate, base cuneate, petioles eglandular, stipules small, axillary, 

leaf base confluent into petiole. 

26. Leaves 9-18 cm long x 4.2-7 cm broad; flowers brown pubescent (Surinam; 

Brazil, Amazonia). 

69.1. L. laevigata. 

26. Leaves 4-5 cm long x 2.5-3.8 cm broad; flowers gray pubescent (Brazil, 

Amazonia). 

69.2. L. occultans. 

25. Leaf apex bluntly acuminate, base subcordate, rounded, subcuneate; petioles usually 

glandular, leaf base not confluent (Trinidad; Venezuela; Guianas; Amazonia). 


2. Petals absent. Sect. Hymenopus. 

27. Leaves thin and membranous, the base cuneate, the venation equally prominent on both 

surfaces; receptacle urceolate (E-central Brazil). 

70. L. glazioviana. 

27. Leaves thick and coriaceous, the base usually rounded to cordate, rarely rounded-subcuneate 

(to cuneate in L. marleneae), the venation obscure on upper surface; receptacle conical to 

globose-cupuliform. 

28. Leaves rounded to obtuse at apex, rarely exceeding 9 cm in length. 

29. Flowers ca. 3 mm long; petioles terete (E Brazil). 

71. L. littoralis. 

29. Flowers ca. 2 mm long; petioles usually canaliculate. 

30. Inflorescence and flowers with brown pubescence; stamens 3; venation of lower 

surface of leaf often papillose; stipules intrapetiolar (Venezuela; Guyana). 

72. L. fanshawei. 

30. Inflorescence and flowers gray-puberulous or glabrescent; stamens 5; venation of 

leaf lower surface smooth-papillose; stipules adnate to extreme base of petiole. 

31. Leaf 2.5-6.5 cm long, the apex and base rounded (Guianas). 73. L. irwinii. 

31. Leaf 5.5-9.5 cm long, the apex acute, the base cuneate (Brazil, central Amazonia). 

73.1. L. marleneae. 

28. Leaves distinctly acuminate at apex, usually exceeding 8 cm in length. 

32. Midrib slightly impressed above; petioles canaliculate (French Guiana). 74. L. cyathodes. 

32. Midrib prominulous above; petioles terete. 

33. Flowers 2.5-3.5 mm long, ferrugineous-brown-pubescent; stipules 4-7 mm long, 

persistent (Venezuela; Guianas; Brazil, Amazonia). 

75. L. polita. 

33. Flowers 1.5 mm long, gray-brown-pubescent; stipules small, caducous (Colombia; 

Venezuela; Brazil, Para). 

76. L. silvae. 

1. Leaf underside pubescent, densely arachnoid-lanate or pulverulent-farinaceous-pubescent (not hirsute); 

petals absent. 

34. Flowers in small cymules, on long slender secondary branches (peduncles) less than 0.5 mm thick 

and attached to primary inflorescence branches; pedicels usually 0.25-3 mm long, rarely absent; 

fruit often very small, usually not exceeding 2 cm in length. 

Sect. Cymosa. 

35. Leaf underside with stomatal cavities or thick, coarse, deeply cut venation, lanate-arachnoid. 

36. Bracteoles large and enclosing groups of flower buds; flowers subsessile; peduncles of 

cymules short and rather thick, 1-3 mm long (Venezuela; Guianas). 7. L. densiflora. 

36. Bracteoles small and not enclosing groups of flower buds; flowers pedicellate; peduncles 

of cymules long and slender. 

37. Inflorescence and exterior of flowers ferrugineous-tomentose; young stems ferrugineous-tomentellous 

(fruit 3-5 mm long, stipitate, densely tomentose, in L. cuprea). 

38. Flowers ca. 2 mm long; stipules ca. 5 mm long; stamens 3 (Guyana). 78. L. cuprea. 

38. Flowers ca. 6 mm long; stipules 7-10 mm long; stamens 6-7 (Brazil, Espirito 

Santo). 

78.1. L. arianeae. 

37. Inflorescence and exterior of flowers gray-puberulous or gray- or brown-tomentose; 

young stems glabrous or puberulous (fruit 1-2 mm long, not stipitate). 

39. Midrib impressed for entire length; petioles 7-14 mm long; leaves oblong-lanceolate 

(Brazil, Amazonia). 

79. L. impressa. 

39. Midrib plane throughout or impressed at base only; petioles 2-10 mm long; leaves 

ovate to oblong. 

40. Leaf apex rounded; leaves orbicular-ovate, petioles 8-10 mm long; stamens 

7 (E Brazil). 

80.1. L. santosii. 

40. Leaf apex acute to acuminate; leaves elliptic; petioles 2-7 mm long; stamens 

3-7.


41. Leaf underside with lanate pubescence obscuring venation; stamens 6- 

7 (central Brazil). 

80. L. dealbata. 

41. Leaf underside pubescent only in the mouth of stomatal cavities, the 

reticulations glabrous; stamens 3-5. 

42. Leaves ovate, broadest near base, 3-8.5 cm long; stipules axillary 

(widespread). 

88. L. hypoleuca. 

42. Leaves oblong-elliptic, broadest about middle, 5-13 cm long; stipules 

adnate to petiole base (Venezuela; Guianas; Brazil, Amazonia). 

81. L. pallida. 

35. Leaf underside with a very fine, plane or prominulous venation, usually pulverulent-farinaceous. 

43. Exterior of flowers and branches of inflorescence entirely glabrous or glabrescent; fruit exterior 

often drying purple. 

44. Leaves oblong-lanceolate to oblong, the lower surface white-lanate (Brazil, Amazonia). 

82. L. gracilipes. 

44. Leaves ovate to oblong, the lower surface sparsely gray-pulverulent-farinaceous (Venezuela; 

Guianas; Brazil, Amazonia). 

83. L. parvifructa. 

43. Exterior of flowers and usually branches of inflorescence puberulous to tomentose; fruit exterior 

gray to brown. 

45. Leaf apex rounded, the margins revolute; inflorescence glabrous (Brazil, Para, Bahia). 

84. L. cymosa. 

45. Leaf apex acuminate, the margins not revolute; inflorescence usually puberulous or tomentose. 

46. Fruit elongate-pyriform, to 2.5 cm long, the exterior with short rufous-velutinous 

pubescence; either inflorescence and exterior of flowers brown-tomentellous or petioles 

7-12 mm long. 

47. Petioles 3-6 mm long, terete; inflorescence and flowers brown-tomentellous; leaves 

triangular or nearly so (W Indies). 

85. L. ternatensis. 

47. Petioles 7-12 mm long, canaliculate; inflorescence and flowers gray-puberulous; 

leaves oblong (Trinidad; Guianas; Amazonia). 

86. L. membranacea. 

46. Fruit ovoid to pyriform, rarely exceeding 1.2 cm long; exterior of flowers gray-puberulous 

or glabrescent; petioles 3-6 mm long. 

48. Leaves 6.5-16 cm long; midrib and primary veins prominent on upper surface; 

rachis and branches of inflorescence glabrescent (Brazil, Amapa). 87. L. piresii. 

48. Leaves 2.5-10 cm long; midrib plane or prominulous on upper surface; rachis 

and branches of inflorescence puberulous. 

49. Inflorescence spreading, the rachis 1-15 mm thick; lower leaf surface pulverulent 

(Venezuela). 87.1. L. furfuracea. 

49. Inflorescence compact, the rachis 0.5 mm thick; lower surface tomentellous 

(widespread). 

88. L. hypoleuca. 

34. Flowers sessile or subsessile on primary branches of inflorescence or on short (less than 0.5 mm long) 

thick peduncles only; fruit rarely less than 2 cm long. 

50. Leaf underside pulverulent-farinaceous. 

Sect. Pulverulenta. 

51. Leaf with rounded to acute apex, the margins usually revolute. 

52. Flowers 1.5-2 mm long; receptacle campanulate-cupuliform. 

53. Young branches and inflorescence densely tomentose; inflorescence to 4 cm long, 

recurved (Venezuela; Guyana). 

89. L. boyanii. 

53. Young branches and inflorescence branches glabrous; inflorescence usually exceeding 

4 cm long, erect (Guyana). 

90. L. buxifolia. 

52. Flowers ca. 3 mm long; receptacle urceolate. 

54. Leaves predominantly orbicular, occasionally oblong-elliptic, the apex rounded to 

retuse (NW Amazonian Venezuela; Brazil). 

91. L. orbicularis. 

54. Leaves elliptic, the apex acute. 

55. Receptacle narrowly urceolate-cylindrical; calyx lobes lanceolate (Brazil, Rond6nia). 

92. L. niloi. 

55. Receptacle broadly urceolate; calyx lobes deltoid (Venezuela; Guianas; Brazil, 

Amazonia). 93. L. coriacea. 

51. Leaf with well-developed acumen, the margins not revolute. 

56. Flowers 3-4 mm long; receptacle urceolate. 

57. Leaf apex caudate; leaves 4-5.5 cm long, chartaceous; exterior of receptacle redbrown-pubescent 

contrasting with white pubescence on interior of calyx lobes (Brazil, 

Rondonia). 95.1. L. teixeirae. 

57. Leaf apex acuminate or acute; leaves 4-15 cm long, usually coriaceous; exterior of 

receptacle and calyx lobes gray-pubescent.


58. Venation of leaf underside minutely reticulate, forming a network with less than 

0.25 mm between reticulations; reticulation apparent because of absence of 

pubescence on veins; leaves thin-chartaceous; stipules usually caducous (Peru; 

Colombia; Brazil, Amazonia). 

94. L. urceolaris. 

58. Venation and reticulation coarse, with 1-2 mm between reticulations; pubescence 

obscuring much of veins; leaves thick-coriaceous; stipules persistent. 

59. Leaf apex with finely pointed, well-developed acumen; rachis of inflorescence 

glabrous (Costa Rica; Panama; Guianas; Brazil, Para). 95. L. affinis. 

59. Leafapex acute or with short blunt acumen; rachis ofinflorescence pubescent 

(Venezuela; Guyana; Brazil, Amazonia). 

93. L. coriacea. 

56. Flowers 1.5-2 mm long; receptacle campanulate. 

60. Primary veins slightly impressed on upper surface; fruit exterior sordid-rufous-pubescent; 

branches of inflorescence densely tomentose to puberulous. 

61. Leaves thick-coriaceous; stipules caducous; flowers in clusters on short thick peduncles; 

stamens 3 (Colombia, Pacific Coast). 

96. L. glauca. 

61. Leaves thin, chartaceous-membranous; stipules persistent; flowers on primary and secondary 

branches of inflorescence; stamens 6-7 (Guianas; Brazil, Amapa, Parf). 

97. L. davillifolia. 

60. Primary veins plane or prominent on upper surface; fruit exterior glabrous, drying yellow; 

branches of inflorescence glabrous to puberulous. 

62. Leaves (11-)13-18 cm long, 4-8 cm broad (Guianas; Brazil, Amazonia). 98. L. elliptica. 

62. Leaves 4-10(-12) cm long, 2-5.5 cm broad (Venezuela; Guianas; Brazil, Amazonia; 

Bolivia). 

99. L. canescens. 

50. Leaf underside densely lanate-arachnoid or with stomatal cavities, never pulverulent. 

Subg. Licania sect. Licania. 

63. Flowers 6-7.5 mm long, stamens often connate for half their length or free. 

64. Stamens connate for half their length; leaves prominently reticulate but without stomatal 

cavities; petioles eglandular (Guyana; Surinam). 

100. L. couepiifolia. 

64. Stamens free to base; leaves with conspicuous stomatal cavities; petioles with 2 glands near 

base of lamina (Brazil, Amapf). 

101.1. L. naviculistipula. 

63. Flowers not exceeding 5.5 mm in length; stamens free almost to base. 

65. Leaf base distinctly cordate or subcordate; leaves usually ovate-orbicular. 

66. Leaves triangular-ovate, 10-16 cm long, membranous (Peru, Loreto). 101. L. trigonioides. 

66. Leaves orbicular to ovate, 3-9 cm long, usually coriaceous. 

67. Young stems hispid; lower surface of leaves with hirsutulous-hispid venation, the 

apex with well-developed acumen (Venezuela, Amazonas). 102. L. cordata. 

67. Young stems puberulous to tomentose; lower surface of leaves glabrous or lanate on 

venation, the apex acute or bluntly acuminate. 

68. Flowers 4.5-5.5 mm long; receptacle urceolate; stipules 5-6 mm long (Venezuela, 

Amazonas). 

103. L. foldatsii. 

68. Flowers 1.5-3.5 mm long; receptacle campanulate; stipules 1-3 mm long. 

69. Leaves submembranous, the lower surface with deeply cut venation, the 

pubescence occurring in cavities and hard to remove; fertile stamens 5-11. 

70. Stamens 8-11; petioles ca. 5 mm long; stipules 3-6 mm long (Amazonian 

Colombia; Venezuela; Brazil). 


70. Stamens 5-6; petioles 1.5-3 mm long; stipules 2-2.5 mm long (Amazonian 

Venezuela; Colombia). 

104. L. hebantha. 

69. Leaves thick-coriaceous, the lower surface with shallow venation, not forming 

cavities, the lanate pubescence easily rubbed off; fertile stamens 3. 

71. Pubescence obscuring venation; stipules adnate to extreme base of petiole 

(Venezuela, Bolivar). 

105. L. steyermarkii. 

71. Pubescence not obscuring venation; stipules axillary (Venezuela). 

106. L. subrotundata. 

65. Leaf base rounded to cuneate; leaves only rarely ovate-orbicular. 

72. Midrib and primary veins distinctly impressed on leaf upper surface. 

73. Leaf underside with well-developed stomatal cavities filled with lanate pubescence; 

petioles 8-17 mm long. 

74. Stomatal cavities conspicuous because of glabrous nerves and veins; leaves 4- 

8.5 cm long (Venezuela; Brazil, Amazonas). 

107. L. crassivenia. 

74. Stomatal cavities less conspicuous because of puberulous nerves and venation; 

leaves (8-)10-25 cm long. 

75. Flowers ca. 1.5-2.5 mm long; inflorescence much branched, predominantly 

axillary; fertile stamens 3 with 4 sterile staminodes; petioles conspicuously 

2-4-glandular (Surinam). 

108.1. L. jimenezii.


75. Flowers 3-5 mm long; inflorescence little branched, terminal; fertile stamens 

6-11, staminodes absent; petiole glands inconspicuous. 

76. Receptacle conical, 4-5 mm long; fruit tomentellous, the stipe 2-6 mm 

long; leaf underside brown-lanate (Guianas; Brazil, Amazonas). 

108. L. majuscula. 

76. Receptacle campanulate, 2.5-3 mm long; fruit pulverulent, the stipe 

8-15 mm long; leaf underside white-lanate (Venezuela; Guianas; Brazil, 

Amazonas). 

109. L. alba. 

73. Leaf underside without stomatal cavities but often prominently reticulate; petioles 2-6(-8) mm 

long. 

77. Exterior of receptacle sparsely puberulous, the pubescence not completely covering the surface; 

leaves oblong-lanceolate (widespread). 

114. L. kunthiana. 

77. Exterior of receptacle densely tomentose to tomentellous, the pubescence completely covering 

the surface; leaves oblong to elliptic. 

78. Leaves 2.5-5.5 cm long, not prominently reticulate beneath; petioles 1.5-3 mm long 

(Venezuela, Bolivar). 

110. L. hitchcockii. 

78. Leaves 5-22 cm long, prominently reticulate beneath; petioles 4-8 mm long. 

79. Flowers ca. 3.5-5 mm long; receptacle broadly cupuliform; leaves orbicular with 

rounded apex, the underside hirsute along venation (Guyana). 111. L. sandwithii. 

79. Flowers 2-3 mm long; receptacle campanulate; leaves oblong to elliptic, acute to 

acuminate at apex, the underside usually tomentellous, rarely hirsute on venation. 

80. Leaf undersurface with parallel secondary veins, giving a prominently reticulate 

appearance; pubescence brown; primary veins 7-10 (Venezuela; Guianas; Brazil, 

Amazonia). 

112. L. laxiflora. 

80. Leaf undersurface with diffuse secondary veins, less prominent; pubescence rufous; 

primary veins 5-6 (Venezuela; Guianas). 

113. L. rufescens. 

72. Primary veins and usually midrib plane or prominent on upper surface. 

81. Exterior of flowers and inflorescence branches gray-puberulous, the pubescence not completely 

covering surfaces. 

82. Receptacle campanulate; leaves without stomatal cavities (widespread). 114. L. kunthiana. 

82. Receptacle urceolate; leaves with shallow stomatal cavities (Brazil, Rond6nia). 

115. L. bellingtonii. 

81. Exterior of flowers densely tomentellous or tomentose, the pubescence completely covering surfaces. 

83. Leaf apex rounded, obtuse or bluntly acute, or rarely short-apiculate (never acuminate). 

84. Lanate pubescence of leaf underside hard to remove, covering deep stomatal cavities; 

receptacle urceolate (Guyana). 

116. L. compacta. 

84. Lanate pubescence of leaf underside easily removed, revealing little protruding venation, 

stomatal cavities absent; receptacle usually campanulate, urceolate in L. ovalifolia and 

L. savannarum only. 

85. Receptacle urceolate; stipules 3-5 mm long, adnate to petiole well above base. 

86. Stamens 3; leaves thickly coriaceous; petioles 4-7 mm long (Guianas; Brazil, 

Amapa). 117. L. ovalifolia. 

86. Stamens 9-11; leaves chartaceous; petioles 1.5-2.5 mm long (Brazil, Amazonas). 

119.1. L. stewardii. 

85. Receptacle usually campanulate, rarely urceolate; stipules 1-3 mm long, axillary or 

adnate to extreme base of petiole. 

87. Leaves ovate-orbicular, the apex often shortly apiculate. 

88. Lanate pubescence obscuring venation; stipules adnate to extreme base of 

petiole (Venezuela, Bolivar). 

105. L. steyermarkii. 

88. Lanate pubescence not obscuring venation; stipules axillary (Venezuela). 

106. L. subrotundata. 

87. Leaves oblong-lanceolate to elliptic, the apex rounded, acute or acuminate. 

89. Stipules adnate to petiole base; petioles 8-12 mm long; inflorescence branches 

very sparsely puberulous; stamens 5 (Brazil, Bahia). 138. L. bahiensis. 

89. Stipules axillary, caducous; petioles 2-7 mm long; inflorescence branches 

densely puberulous or tomentellous; stamens 3. 

90. Leaves oblong-elliptic, sparsely lanate, the underside minutely reticulate; 

primary veins 9-11; petioles 4-7 mm long; receptacle campanulate 

(Colombia). 118. L. caldasiana. 

90. Leaves oblong to oblong-lanceolate, the underside densely farinaceouslanate 

with obscured venation; primary veins 5-8; petioles 2-3 mm 

long; receptacle campanulate-urceolate (Brazil; Venezuela, Amazonas). 

119. L. savannarum.


83. Leaf apex distinctly acuminate or sharply acute. 

91. Leaves 3-4 cm long, with two conspicuous glands at junction of petiole with upper surface of 

lamina (Guyana). 

120. L. microphylla. 

91. Leaves predominantly large, exceeding 5 cm, lacking conspicuous glands. 

92. Stamens 3. 

93. Leaf undersurface with stomatal cavities. 

94. Leaf venation glabrous or glabrescent, hence conspicuous, with glands present at 

base of lower surface; leaves coriaceous; stipules less than 1.5 mm broad at base 

(Peru; Brazil; Colombian Amazonia). 

121. L. triandra. 

94. Leaf venation pubescent, lanate pubescence covering entire leaf and obscuring stomatal 

cavities of lower surface, lacking glands at leaf base; leaves chartaceous; stipules 

2.5 mm broad at base (Brazil, Para). 

121.1. L. tocantina. 

93. Leaf undersurface with prominent venation but no stomatal cavities, the venation pubescent 

and hence less conspicuous. 

95. Stipules caducous, petioles glabrous or tomentose; receptacle campanulate. 

96. Petioles tomentose when young; leaves elliptic to oblong, with well-developed 

acumen; primary veins 7-9 pairs (Venezuela; Guianas; Brazil). 122. L. discolor. 

96. Petioles glabrous; leaves oblong-lanceolate, finely apiculate; primary veins 10- 

12 pairs (Brazil, Amazonia). 

123. L. apiculata. 

95. Stipules persistent, petioles glabrous or puberulous when young; receptacle campanulate 

or urceolate. 

97. Inflorescence 8-15 cm long, spreading, with numerous primary branches; lower 

leaf surface brown-lanate; receptacle campanulate (Colombia; Venezuela; Guianas; 

Brazil, Amazonia, Bahia). 

124. L. micrantha. 

97. Inflorescence 5-6 cm long, with a few primary branches; lower leaf surface 

whitish-gray-pubescent; receptacle urceolate. 

98. Leaf apex with a finely pointed acumen; leaves thin-membranous (French 

Guiana; Brazil, Amapa). 

125. L. pruinosa. 

98. Leaf apex acute to bluntly acuminate; leaves coriaceous. 

99. Leaves thinly coriaceous, 4.5-10.5 x 2-5.5 cm; flowers 4-5 mm long; 

interior of receptacle pubescent on upper portion (E-central Brazil). 

126. L. nitida. 

99. Leaves thickly coriaceous, 3-5.5 x 2-3 cm; flowers 2 mm long; interior 

of receptacle glabrous on upper portion (Brazil, NW Amazonia). 

126.1 L. aracaensis. 

92. Stamens 4-8(-10). 

100. Leaf underside with deep, extremely conspicuous stomatal cavities, the pubescence 

confined to cavities made obvious by almost glabrous venation. 

101. Inflorescence and flowers ferrugineous-pubescent; stipules 1-2 mm long x 1-1.5 

mm broad at base, persistent, ferrugineous; leaf apex acute to bluntly acuminate 

(E-central Brazil). 

127. L. riedelii. 

101. Inflorescence and flowers gray-puberulous; stipules 2-4 mm long, 0.2-0.5 mm 

broad at base, persistent or caducous, pubescent but not ferrugineous; leaf with 

well-developed acumen. 

102. Bracteoles persistent, lanceolate; upper surface of midrib impressed; stipules 

caducous (Brazil, Amazonia). 

128. L. bracteata. 

102. Bracteoles caducous, triangular; upper surface of midrib plane; stipules small 

but persistent. 

103. Inflorescence branches thick; flowers subsessile; fruit exterior ferrugineous-velutinous 

(Venezuela; Colombia; Guianas; Brazil, Amazonia). 

129. L. parviflora. 

103. Inflorescence branches slender; flowers distinctly pedicellate; fruit exterior 

reddish-brown, short-pulverulent (Venezuela; Brazil, Amazonas). 

81. L. pallida. 

100. Leaf underside with poorly developed stomatal cavities or none, venation pubescent. 

104. Petioles 1.5-2 cm long; inflorescence rachis and branches glabrous or glabrescent 

(Guianas; Brazil, Para). 

130. L. robusta. 

104. Petioles to 1 cm long (to 1.5 cm in L. bahiensis and L. lamentanda); inflorescence 

rachis and branches usually densely tomentose or tomentellous. 

105. Inflorescence predominantly of axillary and terminal spikes, terminal inflorescences 

rarely little branched or with minute spikes along rachis. 

106. Leaves lanceolate, the lower surface deeply reticulate, with poorly 

developed stomatal cavities (Venezuela, Amazonas; Brazil, Roraima). 

131. L. lanceolata.


106. Leaves oblong to ovate-elliptic, the lower surface not deeply reticulate, 

lacking stomatal cavities. 

107. Flowers 2.5 mm long; receptacle broadly cupuliform; inflorescence 

puberulous; reticulate venation of leaf underside conspicuous 

(E-central Brazil). 

132. L. spicata. 

107. Flowers 1.5-2 mm long; receptacle campanulate; inflorescence 

tomentose; venation of leaf underside inconspicuous except in 

L. nelsonii. 

108. Flowers in short, dense, minute spikes attached to long 

rachis (Surinam). 

133. L. stricta. 

108. Flowers solitary along rachis or in dense glomerules, but 

not in minute spikes. 

109. Leaf undersurface conspicuously reticulate; stipules 

adnate to base of petiole; receptacle rufous-pubescent; 

flowers densely and evenly clustered along inflorescence 

rachis; large tree. 135.1. L. nelsonii. 

109. Leaf undersurface lanate, not conspicuously reticulate; 

stipules axillary; receptacle gray-brown pubescent; 

flowers in dense glomerules or small groups; 

small shrub to medium sized tree (15 m). 

110. Leaves thin and membranous, the acumen 

finely pointed; flowers in dense glomerules; inflorescences 

largely axillary (Venezuela; 

Guianas; Brazil, Bahia, Parf). 

134. L. leptostachya. 

110. Leaves thick and coriaceous, the acumen usually 

blunt; flowers in small groups or solitary 

but not glomerulate; inflorescences largely terminal 

(Venezuela; Guianas; Brazil, Amazonia). 

135. L. incana. 

105. Inflorescence of terminal and sub-terminal racemose panicles. 

111. Leaves lanceolate; low shrub or subshrub (Venezuela, Amazonas; Brazil, Roraima). 

131. L. lanceolata. 

111. Leaves ovate to oblong-lanceolate; trees or tall shrubs. 

112. Stipules 3-10 mm long and at least 1 mm broad at base, distinctly adnate to petiole or 

intrapetiolar, persistent and obvious. 

113. Midrib distinctly impressed above; bracteoles 0.2-1.5 mm long. 

114. Inflorescence ferrugineous-pubescent; petioles canaliculate; stipules adnate to 

petiole well away from axil (Bolivia; Brazil, Para). 136. L. paraensis. 

114. Inflorescence gray-puberulous; petioles terete; stipules intrapetiolar or adnate 

to inside of extreme base of petiole. 

115. Leaves with a finely pointed acumen; primary veins 11-12 pairs, the 

lower surface glandular at base (Colombia, Amazonas; Brazil, W Amazonia). 

137. L. vaupesiana. 

115. Leaves with blunt acumen; primary veins 6-8 pairs, the lower surface 

eglandular at base (E-central Brazil). 

138. L. bahiensis. 

113. Midrib plane or prominulous above; bracteoles 1.5-3 mm long. 

116. Leaves membranous; flowers ca. 2 mm long, gray-tomentellous; inflorescence 

gray-puberulous; stamens 5 (Brazil, Amapa). 

139. L. maxima. 

116. Leaves coriaceous; flowers 3-3.5 mm long, brown-tomentose; inflorescence 

brown-tomentose; stamens 8-11 (Amazonian Colombia; Venezuela; Brazil). 


112. Stipules usually less than (rarely exceeding) 2.5 mm long and very narrow to base, on 

outside of axils or adnate to extreme base of petiole, caducous or persistent, often inconspicuous. 

117. Stipules adnate to extreme base of petiole, persistent or subpersistent. 

118. Petioles 7-12 mm long, remaining tomentellous even with age; laminae 7- 

16 x 3-7.5 cm; stipules sub-persistent (Brazil, Para and central). 

141. L. blackii. 

118. Petioles 2-6 mm long, becoming glabrous with age; lamina 3-12 x 2.2-5.5 

cm; stipules persistent.


119. Inflorescence lax and spreading; leaf with a finely pointed acumen 8-20 mm 

long, the lower surface farinaceous-lanate, shallowly reticulate (Colombia; 

Venezuela; Guianas; Brazil, Amazonia). 

129. L. parviflora. 

119. Inflorescence and flowers densely crowded; leaf with blunt acumen 2- 

13 mm long, the lower surface lanate, deeply reticulate (Venezuela; 

Guianas to E-central Brazil). 

114. L. kunthiana. 

117. Stipules axillary, caducous or persistent. 

120. Petioles 11-15 mm long; flowers 3.5-4 mm long (Brazil, Bahia). 130.1. L. lamentanda. 

120. Petioles 2-10 mm long; flowers 2-3 mm long. 

121. Leaf undersurface with smooth inconspicuous reticulation, the pubescence easily removed; 

primary veins widely spaced, 1.2-2.5 cm apart; exterior of receptacle velutinouspubescent 

(Brazil, Amazonas). 

142. L. rodriguesii. 

121. Leaf underside with deeply cut reticulation, and hence pubescence hard to remove; primary 

veins not more than 1 cm apart; exterior of receptacle tomentellous. 

122. Receptacle globose; upper surface of youngest leaves appressed-strigose, soon becoming 

glabrous; inflorescence of axillary spikes and terminal panicles (E-central 

Brazil). 

132. L. spicata. 

122. Receptacle usually campanulate; upper surface of youngest leaves glabrous; inflorescence 

of terminal and subterminal racemose panicles. 

123. Leaves oblong-lanceolate; midrib slightly impressed on upper surface (E-central 

Brazil). 

143. L. indurata. 

123. Leaves ovate-elliptic to oblong; midrib usually plane on upper surface. 

124. Inflorescence much branched, spreading, lax; lower leaf surface rufouspubescent 

(Venezuela; Guianas; Brazil). 

113. L. rufescens. 

124. Inflorescence densely crowded, compact, little-branched; lower leaf surface 

gray-brown-pubescent. 

125. Petioles 4-7 mm long, terete; receptacle and calyx lobes tomentose 

on exterior. 

126. Leaves finely acuminate, the acumen 8-12 mm long; stipules 

caducous, ca. 1 mm long (Ecuador). 144.1. L. harlingii. 

126. Leaves bluntly acuminate to obtuse; stipules persistent, 1- 

3 mm long. 

127. Petioles soon becoming glabrous; leaf apex bluntly 

acuminate; young fruit pyriform (E-central Brazil). 

144. L. hoehnei. 

127. Petioles tomentose, becoming less so with age; leaf 

apex obtuse to acute; young fruit cylindrical when 

young, becoming pyriform (Panama; Trinidad; Venezuela). 

145. L. cruegeriana. 

125. Petioles 10-12 mm long, canaliculate; receptacle and calyx lobes 

velutinous on exterior (Brazil, Bahia). 

146. L. belemii. 

Additional Notes and Descriptions 

of Species of Licania 

The notes on the species that follow refer only 

to those species for which interesting and useful 

new data have been collected since the mono- 

graph (Prance, 1972). 

Subgenus Moquilea Section Moquilea 

2-3. Licania maritima Prance, Fl. Neotrop. 

Monogr. 9: 44. 1972. 

Fruit globose with warty surface, 9-10 cm in 

diam.; exocarp glabrous, crustaceous; mesocarp 

1.5-1.8 cm thick, fibrous; endocarp ca. 3 mm 

thick, hard and woody, glabrous within. 

Distribution (Fig. 59). Endemic to coastal forests 

of Choc6, Colombia. 

Additional specimen examined. COLOMBIA. CHOCO: 

Mun. El Valle, Jan 1985 (fr), P. Perez s.n. (MEDEL). 

This species was described from a single flow- 

ering specimen. The fruits are large and rather 

similar to those of L. macrocarpa, and are said 

to be edible.


a & 

s o~ , 

E I X D 

dI 

I X 

el 

e l ^ e: ^ e r X < ,,. vE , 

0 

0 

t 

CO i O O 

i. 

C ' 

A. 

FIG. 2. Licaniafilomenoi (Gentry et al 25665). A, habit; B, bract; C, flower bud; D, section of flower bud; 

E, ovary and style; F, stamen; G, H, petals; I, young fruit. 

21 

r, ;'"~ 

:~ . ~,;, ..'.';:~~i??;'.. 

,, ~ i ~ : - . ~~:.:;::.'::.-. '


puberulous beneath; primary veins 21-24 pairs, conspicuous parallel reticulate pattern; petioles 

slightly impressed above, prominent beneath; 8-12 mm long, puberulous, terete, eglandular. 

secondary veins more or less parallel on lower Stipules intrapetiolar, ca. 2 mm long, persistent. 

surface, forming a conspicuous reticulate pat- Inflorescences ramiflorous, little-branched racetern; 

petioles 6-8 mm long, terete, eglandular, mose panicles, to 15 cm long, the rachis and 

sparsely puberulous. Stipules not seen. Inflores- branches brown-shaggy-tomentose. Bracts 4-4.5 

cences of unbranched racemes or little-branched cm long, membranous, glabrous except for marpanicles, 

5-13 cm long, axillary or subterminal, gins. Flowers 5-6 mm long, solitary and not 

the rachis and branches ferrugineous tomentose. densely crowded on inflorescence branches. Re- 

Bracts at base of inflorescence large and persis- ceptacle cupuliform, sessile, sparsely puberulous 

tent, 25-35 cm long, chartaceous; bracteoles 5- on upper part of exterior, tomentose only around 

15 mm long, shortly tomentellous on exterior, base, densely tomentose within. Calyx lobes five, 

glabrescent within. Flowers ca. 5 mm long, sol- acute, puberulous on exterior, tomentellous 

itary along rachis and primary branches of inflo- within. Petals five, white. Stamens ca. 40, inrescence. 

Receptacle cupuliform, sessile, ferru- serted around complete circle, exserted, connate 

gineous-brown tomentose on exterior, densely at extreme base, with dense villous mass at base 

tomentose within. Calyx lobes five, acute, toof filaments. Ovary inserted at base of receptacle, 

mentose on exterior. Petals five, white, tomen- lanate. Style lanate for two-thirds of length, extellous 

on exterior, glabrous within. Stamens ca. ceeding filaments. Fruit not seen. 

35, inserted around complete circle, the filaments Distribution (Fig. 42). Known only from Anexserted 

(seen only in buds). Ovary inserted at dean foothills of Ecuador. 

base of receptacle, villous-pubescent. Style hir- Habitat. Primary forest. 

sute on lower portion. Youngfruit globose, exo- 

Additional 

carp lenticellate. 

specimens examined. ECUADOR. 

AZUAY: Between Rio Blanco and Rio Norcay on rd. 

Distribution (Fig. 38). Known only from the Chancanceo to Molleturo, 1520 m, 4 Jun 1943 (st), 

type collection from San Martin, Peru. Steyermark 52829 (F, NY). PICHINCHA: Reserva Fo- 

Habitat. Mature flatland forest on lateritic soil. restal Endesa, Rio Silanche, 0?5'N, 79002'W, 650-700 

This species is closely related to Licania duri- m, 10 Jun 1984 (st), Jaramillo 6672 (NY, QCA), 14 

folia, with which it shares a similar parallel re- 

Aug 1984 (st), Jaramillo 6960 (NY, QCA), 18 Mar 

1985 (st), Jaramillo 7527 (NY, QCA). 

ticulate venation pattern of the leafundersurface, 

but from which it differs in the much smaller Closely related to Licania durifolia and L. fiinflorescence, 

less densely crowded flower, and lomenoi, this species shares with them leaves with 

persistent bracts and bracteoles. It is named for a gray pubescence and conspicuous parallel sec- 

Filomeno Encarnaci6n, botanist of Iquitos, Peru. ondary venation. It differs from L. durifolia in 

the smaller unbranched inflorescence with less 

24.2. Licania grandibracteata Prance, sp. nov. 

Type. Ecuador. Pichincha: Reserva Florestal 

Endesa, Rio Silanche, 0?5'N, 79?02'W, 650- 

700 m, 7 Dec 1984 (fl), J. Jaramillo 7413 (holotype, 

NY; isotype, QCA). Fig. 3. 

A L. durifolio inflorescentiis minoribus, ramifloris, 

floribus haud agglomeratis, receptaculo extus 

sparse puberulo, bracteolis persistentibus differt. 

dense flowers, the persistent bracteoles and the 

sparse pubescence of the receptacle. It differs from 

both species in the ramiflorous inflorescence and 

from L. filomenoi in the much sparser pubescence 

of the inflorescence and flowers, and the 

larger membranous bracteoles. 

2-5. Licania macrocarpa Cuatrecasas, Fieldiana, 

Bot. 27: 107. 1951. 

Tree to 30 m tall, the young branches glabrous, 

conspicuously lenticellate. Leaf lamina oblong, 

coriaceous, 31-33 x 7-13.5 cm, rounded at base, 

bluntly acute at apex, with short appressed graybrown 

pubescence beneath; midrib prominulous 

above, prominent beneath; primary veins 21-24 

pairs, impressed above, prominent beneath; secondary 

veins more or less parallel, forming a 

Distribution (Fig. 59). Lowland forests to 600 

m in Colombia, Ecuador, and Peru. 

Additionalspecimensexamined. COLOMBIA. CHocO: 

3 km W of Tutunend6, 7 Jan 1981 (st), Gentry et al. 

30324 (MO, NY). 

ECUADOR. LOS RIOS-PICHINCHA: El Centinela, 

Montaiias de Lla, 6 Feb 1979 (fl), Dodson et al. 7516 

(MO), 2 Oct 1979 (fr), Dodson et al. 8672 (MO, NY);


t~~~~~~~~~~~? 

/\I YA*A ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ 4 

r~~yr ~~ ~r- I I ~ ~ ~ ";.i~~Tki~~e~~!'-?~~;~;. 

? ~ ~ no 

FIG 3 Lcaiagradirateta(Jramll 713. , hbi; , ea uderurac; , rac; . loer E 

flwr etin F vayan tye G etl


Cerro Antisana, NE of Borja (fl), Grubb et al. 1045 a very small ovary at the base, and the greater 

(NY). 

number of stamens. It is the 

PERU: PASCO: Prov. Oxapampa, Palcazu Valley, Is- 

largest-leaved species 

cozacin, 22 Jan 1984 (fl), Foster 7961 (MO, NY). 

of Licania so far described. It is related to the 

other large-leaved, lanate pubescent species of 

2-5.1. Licania gentryi Prance, sp. nov. Type. Co- Licania subgenus Moquilea, but is quite distinct 

lombia. Valle: Bajo Calima, 15 km N of Bue- from them by the leaf size and shape and the 

naventura, 3?56'N, 77?08'W, 50 m, 16 Feb ramiflorous nature of the inflorescence. 

1983 (fl), A. Gentry et al. 40355 (holotype, 

MO; isotype, NY). Fig. 4. 2-5.2. Licania cabrerae Prance, Brittonia 28: 

210-212, fig. 1. 1976. Type. Colombia. An- 

Species a L. cabrerae affinis, foliis 27-47 cm tioquia: Estaci6n Experimental Forestal Pielongis 

x 22-26 cm latis, inflorescentiis rami- dras Blancas, 13 Jul 1957 (fl), I. Cabrera R. 

floris, receptaculo crasso, staminibus circa 60 dif- 94 (holotype, COL). 

fert. 

Tree 10 m tall. Leaf lamina elliptic, charta- Small tree, the young branches ferrugineousceous, 

27-47 cm x 22-26 cm, rounded at base, 

lanate pubescent, soon glabrate. Leaflamina obabruptly 

mucronate at apex, the mucro 5-7 mm long to oblong-lanceolate, coriaceous, 12-17 x 

long, densely ferrugineous-lanate-pubescent be- 3-4.5 cm, subcuneate at base, acuminate at apex, 

neath, lanate above when young, becoming glathe 

acumen 8-15 mm long, lanate-tomentose on 

brous with age, the pubescence persisting only upper surface when young, soon becoming glaalong 

veins; midrib prominent beneath, plane brous, ferrugineous-lanate-tomentose beneath 

above, with a ferrugineous-lanate-pubescence 

when young, becoming gray-tomentellous with 

contrasting with the caducous lighter pubescence age; midrib prominulous above, lanate when 

of upper surface and lateral veins; primary veins young, prominent beneath, gray-tomentellous; 

25-29 pairs, prominent beneath, slightly im- primary veins 16-19 pairs, plane above, prompressed 

above; petioles 20-25 mm long, ferru- inent beneath; petioles 10-12 mm long, terete, 

gineous-lanate-pubescent, terete, eglandular. 

lanate-tomentose when young. Stipules mem- 

Stipules axillary, ca. 12 mm long, membranous, branous, triangular, caducous. Inflorescences of 

sparsely lanate on exterior, glabrous on inner sur- racemose panicles to 18 cm long, the rachis and 

face. Inflorescences of panicles borne on the branches densely ferrugineous-tomentose; bracts 

woody branches, the rachis and branches to- and bracteoles triangular, membranous, ferrumentellous; 

bracts large and membranous to 20 gineous-pubescent, glabrous on inner surface, remm 

long, persisting at base of inflorescence; flexed, caducous. Flower buds enclosed by large 

bracteoles membranous, to 20 mm long, tomen- navicular, amplexicaul bracts. Flowers 6-7 mm 

tellous on exterior, glabrous within. Flowers 6- long. Receptacle globose, ferrugineous-lanate on 

7 mm long, sessile. Receptacle cupuliform, fer- exterior, with a thick row ofdeflexed hairs around 

rugineous-lanate-pubescent on exterior, the walls faucal annulus within; pedicels ca. 0.5 mm long. 

thick, restricting the inner cavity, tomentose Calyx lobes five, triangular, acute, ferrugineouswithin. 

Calyx lobes five, triangular, lanate on ex- lanate on exterior, brown-tomentose within. Petterior, 

tomentellous within. Petals five, white. als five, white, tomentellous on exterior, glabrous 

Stamens ca. 60, inserted in several rows around within. Stamens 35-40, inserted in a complete 

a complete circle, the filaments glabrous, ex- circle, the filaments shortly exserted, free to base, 

ceeding calyx lobes, free to base, with a dense with a few scattered hairs on lower portion, pliring 

of reflexed hairs around base on interior of cate in bud. Ovary pilose. Style pilose almost to 

receptacle. Ovary inserted at base of receptacle, apex. Fruit unknown. 

small, tomentose. Style longer than filaments, Distribution (Fig. 30). Known only from the 

villous for three-fourths of length. Fruit not seen. type gathering collected in flower in July. A high 

This species (distribution in Fig. 40), most altitude species, the type from 2550 m. 

closely related to the highland Licania cabrerae, 

differs in the very much larger and broader leaves, 2-5.3. Licania fasciculata Prance, Acta Amain 

the inflorescence borne on woody branches, z6nica 8: 579-581, fig. 2. 1978. Type. Panama. 

the much thicker walls of the receptacle with only Colon: Zona de Santa Rita, 31 Aug 1972 (fl),


"-~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~ 

IVI]' 

D.~~~~~~~~~~~~~~?: 

o~.:: ... .. 

'~~~~~~~~~~~~~~~~~,?' 

E.? 

3 

enmI 

: 

' 

.. . .... . 

'i~~~~~~~~~~~~~~~~~~~~~~~~~~~' 

!~!"~'!~x ~~~~~~~~~~~~~~~~~~~~~~ .... 

.i 

'~.i[11.'I .ri}~! 

i i'i 

i 

X C. 

: 

' 

FIG. 4. Licania gentryi (Gentry 40355). A, leaf and leaf undersurface, B, inflorescence; C, flower bud; D), 

flower; E, petals, left inner surface, right outer surface; F, flower section. 

rr


Correa A. & Dressier 1815 (holotype, MO; 

isotype, NY). 

2-5.4. Licania montana Prance, Brittonia 28:212, 

fig. 2. 1976. Type. Venezuela. Lara: Vic. La- 

guna Negra, Loma de Los Naranjos, 24 Mar 

1975 (fl), Steyermark, R. Smith & C. Espinoza 

111541 (holotype, NY; isotype, VEN). 


tomentose soon becoming glabrous and lenti- 

cellate. Leaf lamina, narrowly elliptic, charta- 

ceous, 6-9.5 x 2-3 cm, subcuneate at base, acu- 

minate at apex, the acumen 3-5 mm long, 

glabrous above, shortly appressed-tomentellous 

beneath; midrib prominulous above, pubescent 

Tree 12 m tall, the young branches tomentellous, 

becoming glabrous and prominently lenticellate 

with age. Leaf lamina oblong to oblong 

lanceolate, thickly coriaceous, 9-13.5 x 2.8-4 

cm, cuneate at base, acuminate at apex, the acumen 

4-7 mm long, abrupt, glabrous above, with 

a compact appressed gray-lanate pubescence beneath; 

midrib prominulous and tomentellous towards base only, prominent beneath, sparsely 

above, prominent beneath; primary veins 15-19 lanate-arachnoid-pubescent; primary veins 15pairs, 

prominent beneath, slightly impressed 17 pairs, prominulous on both surfaces; petioles 

above; petioles 5-10 mm long, tomentellous, te- 4-5 mm long, terete, tomentellous, eglandular. 

rete. Stipules axillary, 3 mm long, persistent, Stipules caducous (not seen). Inflorescences of 

membranous, puberulous. Inflorescences of short racemose terminal and subterminal panicles, 8- 

fasciculate racemes 1-2.5 cm long with flowers 12 cm long; the rachis and branches ferrugineous 

densely clustered, completely obscuring the ra- tomentose. Bracts and bracteoles membranous, 

chis. Bracts and bracteoles 4-7 mm long, ovate, caducous, tomentose on exterior, sparsely putomentellous, 

subpersistent. Flowers 6-7 mm bescent within. Flowers 4-5 mm long. Receptacle 

long. Receptacle cupuliform, tomentose on ex- globose, sessile, ferrugineous-tomentose on exterior, 

tomentose within. Calyx lobes five, acute, terior, tomentose within, with a row of deflexed 

tomentellous on both surfaces. Petals five, white, hairs around the faucal annulus. Calyx lobes five, 

puberulous on exterior. Stamens ca. 60, inserted acute, triangular, ferrugineous-tomentose on exin 

complete circle; filaments far exceeding the terior, brown-tomentellous within. Petals five, 

calyx lobes, glabrous, united at base, the basal tomentellous on exterior, tomentellous towards 

fused portion bent inwards, villous. Ovary in- apex and glabrous towards base on inner surface, 

serted at base of receptacle, pilose. Style villous the margins ciliate. Stamens 30-35, inserted 

for two-thirds of length. Fruit not seen. around a complete circle, the filaments shortly 

Distribution. Figure 38. 

exserted, free to base. Ovary lanate. Style lanate- 

Additional specimen examined. PANAMA. PANAMA: pilose equalling filaments in length. Fruit un- 

El Llano-Carti Rd., 7.5 km N of Panamerican known. 

Hwy., 

23 Jan 1977 (fl), Folsom 1435 (NY). 

Distribution (Fig. 65). A cloud forest species 

collected at 1300-1500 m; still known only from 

Licaniafasciculata differs from all other species the 

in the 

type 

section by the 

gathering, collected in flower in March. 

fasciculate, densely flowered, 

racemose inflorescence. It is most closely 

related to the species group of L. maritima, L. 2-7. Licania klugii Prance, Fl. Neotrop. Monocabrerae, 

L. durifolia, L. montana, L. macro- gr. 9: 47. 1972. 

carpa, and L. veneralensis, and has the same pu- Additional material of this species, originally 

bescence and venation pattern of the leaf under- based on a single collection, shows it to be quite 

surface. It differs in the inflorescence and in the distinct. 

smaller leaves with fewer primary veins, which 

are impressed on the upper surface. Licaniafas- 

Additional specimens examined. PERU. LORETO: 

ciculata also has a greater number of stamens Maynas, Rio Mom6n, 24 Nov 1977 (y fr), Rimachi Y. 

3274 (NY). SAN MARTIN: Quebrada Hicte, Rio Huathan 

the other species listed above. It is a most llaga, 26 May 1964 (fl), Schunke V. 6502 (F). 

distinct species, not easily confused with any other 

in the genus. 

2-9. Licania retifolia Blake, Contr. Gray Herb. 

52: 66. 1917. 

This poorly known species was represented by 

the type from Guerrero, Mexico and one other

Systematic Treatment27 

collection in Prance (1972). Recent collections panulate, tomentellous on exterior, densely tohave 

extended the range to El Salvador. mentose within; pedicels 1-2 mm long, articu- 


late. Calyx lobes five, acute, tomentellous on both 

surfaces. Petals 

Additional specimens examined. MEXICO. 

five, white, glabrous. Stamens 

GUERRERO: Rinc6n Viejo, 15 Apr 1960 (fl), Kruse 261 30-35, inserted around complete circle, the fil- 

(ENCB), 12 Apr 1964 (fl), Kruse 1256 (ENCB). aments exserted. Ovary inserted at base of re- 

EL SALVADOR. AHUACHAPAN: Cerro La Piedra del ceptacle, tomentose. Style hirsute on lower half. 

Filo, 13?51'N, 89?56'W, 30 Jan 1980 (fl), Witsberger Fruit not seen. 

813 (NY). 

Habitat. Tropical rain forest. 

This can be a large tree. 

Additional specimen examined. ECUDAOR. ESME- 

Local name. El Salvador: mulo. 

RALDAS: Lita, 600 m, 20 May 1987 (fl), Van der Werff, 

Dodson & Palacios 9540 (NY, MO). 

2-10. Licania longipedicellata Ducke, Bull. Mus. 

Hist. Nat. (Paris), s6r. 2, 4: 725. 1932. This species belongs to section Moquilea and 

is most closely related to Licania leucosepala, 


but differs in the much broader leaves with widely 

Additional specimens examined. PERU. AMAZONAS: spaced secondary veins and in the loosely 

Rio Santiago, Caterpiza, 17 Sep 1979 (y fr), Huashikat branched inflorescence with articulations at each 

662 (MO, NY). 

branch quite unlike most species of Licania, 

BRAZIL. AMAZONAS: Tabatinga, Sao Leopoldo, 13 which have racemose panicles. L. dodsonii is also 

Oct 1976 (fl), Braga et al. 3127 (INPA). 

close to L. minutiflora but differs in the inflo- 

Local name. Peru: yukuku. 

rescence, the broader leaves that are subcuneate 

rather than rounded at the base, and acuminate 

2-13.1. Licania dodsonii Prance, sp. nov. Type. rather than cuspidate at the apex, and in the 

Ecuador. Esmeraldas: Mun. de Lita, 19 km N greater number of stamens. 

of Lita, road to San Lorenzo, ca. 650 m, 10 

May 1987 (fl), Acevedo & Daly 1657 (holotype, 2-14.1. Licania anneae Prance, Brittonia 31: 250, 

NY). Fig. 155. fig. 2. 1979. Type. Brazil. Para: Cuiaba-Santarem 

Species a L. leucosepala affinis, foliis latiori- 

Hwy., km 1305, vie. Igarap6 Jose Preto, 

22 Nov 1977 

bus, nervis secundariis 8-19 mm distantibus, in- 

(fl), Prance et al. 25652 (holoflorescentiis 

laxis multiramosis, ramis articulatis 

type, MG; isotypes, AAU, NY). 

differt. 

Tree 7 m tall, the young branches sparsely pu- 

Tree 10-15 m tall, the young branches gla- berulous, soon becoming glabrous. Leaf lamina 

brous, conspicuously lenticellate. Leaf lamina oblong-lanceolate, coriaceous, 14-25 x 4.5-8 cm, 

elliptic, chartaceous to thinly coriaceous, 9.5-17 rounded at the base, shortly acuminate or acute 

x 5.5-10.5 cm, rounded to subcuneate at base, at the apex (acumen 2-6 mm long), glabrous 

abruptly acuminate at apex, the acumen 3-5 mm above, pilose with a few sparse appressed hairs 

long, glabrous on both surfaces; midrib plane to beneath, especially on the primary veins; midrib 

prominulous above, prominent and appressed glabrous and prominulous above, prominent and 

villous pubescent beneath; primary veins 13-15 puberulous beneath; primary veins 18-22 pairs, 

pairs, plane to slightly impressed above, prom- slightly impressed above, prominent and puinent 

beneath; petioles 5-7 mm long, terete, berulous beneath, arcuate and conspicuously 

eglandular, sparsely puberulous, glabrescent. anastomosing ca. 2-3 mm from the margin; pet- 

Stipules not seen. Inflorescences of much iole 5-7 mm long, rugulose, slightly canaliculate, 

branched panicles, the primary branches with puberulous when young. Stipules caducous (not 

short thick secondary branches bearing groups seen). Inflorescences of terminal and axillary raof 

2-3 flowers, the branches with articulations at cemose panicles, the rachis and branches light 

junctions, the rachis and branches appressed to- brown-tomentellous. Bracts 5-18 mm long, obmentellous. 

Bracts and bracteoles minute, tri- long to lanceolate, membranous, persistent, toangular, 

puberulous, caducous while in early bud. mentellous on the exterior, sparsely puberulous 

Flowers 5-6 mm long. Receptacle broadly cam- within; bracteoles ovate, persistent, membra-


nous, glabrous or with a few sparse hairs and 

ciliate margins. Flowers 3-3.5 mm long, inserted 

on primary branches of the inflorescence. Receptacle 

cupuliform, extremely regular and round 

at the apex, short brown-tomentellous on exterior, 

tomentellous within. Calyx lobes five, acute, 

triangular, small, tomentellous on both surfaces, 

but more sparsely on the inner surface. Petals 

five, oblong, white, glabrous with ciliate margins, 

caducous. Stamens 12-14, inserted around a 

complete circle, filaments slightly exceeding the 

calyx lobes, glabrous, connate at base forming a 

ring 1 mm tall that is hirsute on both surfaces. 

Ovary inserted at the base of the receptacle, tomentose-pilose. 

Style lanate almost to the apex. 

Fruit not seen. 


Habitat. Disturbed forest beside road on marginally 

flooded and non-flooded areas. 

This species belongs to subgenus Moquilea section 

Moquilea, and is quite distinct within the 

section. It is closest to Licania angustata, a species 

of western Amazonia, but differs in the much 

larger leaves, the shorter petioles, the higher 

number of primary veins, the larger bracts, the 

fewer stamens, and the distinctive appressed pubescence 

of the lower leaf surface. 

2-16. Licania gonzalezii Miranda, Bol. Soc. Bot. 

Mexico 29: 36. 1965. 

This poorly known but distinct species was 

described from two collections from the States 

of Nayarit and Jalisco. A collection from Guer- 

rero has now been added. 


Additional specimen examined. MEXICO. GUERRERO: 

Rincon Viejo, 700 m, 2 Apr 1961 (fl), Kruse 625 

(ENCB). 

The field notes state that the flowers are white 

and very aromatic and are "pollinated" by Lep- 

idoptera, Hymenoptera, and Diptera. 

2-18.1. Licania chiriquiensis Prance, Brittonia 

29: 154, fig. 1. 1977. Type. Panama. Chiriqui: 

Cerro Colorado, 50 km N of San Felix, cloud 

forest, 1200-1500 m, 17 Aug 1975 (fl), Mori 

& Dressier 7778 (holotype, NY; isotype, MO). 


puberulous, soon becoming glabrous and con- 

spicuously lenticellate. Leaf lamina oblong-el- 

liptic, subcoriaceous, 5-8.5 x 2.8-3.8 cm, sub- 

cuneate at base, acuminate at apex, the acumen 

5-6 mm long, glabrous, midrib prominulous 


pairs, prominulous above, prominent beneath; 

petioles 5-7 mm long, glabrous, terete, eglan- 

dular, with very distinct articulation at junction 

with branch. Stipules caducous (not seen). Inflo- 

rescences (not fully developed in specimen ex- 

amined) of terminal and axillary panicles, the 

rachis and branches yellow-brown tomentellous. 

Bracts and bracteoles triangular, caducous, puberulous 

on exterior. Flowers 3-3.5 mm long. Receptacle 

broadly campanulate, tomentellous on 

exterior, hirsute within; pedicels ca. 0.5 mm long. 

Calyx lobes five, acute, tomentellous on exterior, 

puberulous within. Petals five, glabrous with ciliate 

margin. Stamens 13-15, inserted around a 

complete circle; filaments slightly exserted, free 

to base. Ovary inserted at base of receptacle, glabrous 

or sparsely hispid. Style sparsely hirsute. 



Habitat. Cloud forest over 1200 m. 

Additional specimen examined. PANAMA. PANAMA: 

Cerro Jefe, 30 Sep 1978 (fl), Hammel 4823 (MO, NY). 

Licania chiriquiensis belongs to subgenus Moquilea 

section Moquilea and is most closely related 

to L. minutiflora, a species of the Guianas 

and Amazonia. It differs in the smaller, less 

branched inflorescence with much larger flowers, 

the smaller leaves with longer petioles, the almost 

glabrous ovary and in the more tapered, 

less caudate leaf apex. 

2-18.2. Licania kallunkiae Prance, Acta Ama- 

z6nica 8: 583, fig. 4. 1978. Type. Panama. Co- 

16n: Santa Rita Rd., 17 km from Boyd-Roo- 

sevelt Hwy., 450 m, 14 Mar 1975 (fl), Mori & 

Kallunki 5052 (holotype, NY; isotype, MO). 

Tree 14 m tall, the young branches very sparsely 

puberulous, soon becoming glabrous, not conspicuously 

lenticellate. Leaf lamina oblong, coriaceous, 

5.5-10.5 x 2.2-4.0 cm, subcuneate at 

base, cuspidate at apex, acumen 6-10 mm long, 

glabrous on both surfaces; midrib plane above, 

prominent beneath, glabrous; primary veins 9- 

12 pairs, almost plane and inconspicuous on both


surfaces, glabrous; petioles 6-7 mm long, glabrous, 

canaliculate, eglandular. Stipules small, 

lanceolate, puberulous, caducous. Inflorescences 

terminal and axillary panicles 5-11 cm long, 

3-branched, the rachis and branches sparsely gray- 

1971 (fl), Contreras 10742 (holotype, LL; iso- 

type, US). 

Tree ca. 30 m, the young branches glabrous. 

Leaf lamina elliptic, 10-21 x 5.5-9.5 cm, the 

base rounded to 

puberulous. Bracts and bracteoles caducous 

subcuneate, the apex with a short 

(not 

acumen 4-7 mm 

seen). Flowers 2.5-3 mm long, borne in 2-3-flow- 

long, glabrous on both surfaces; 

ered cymules attached to primary branches primary veins 8-10 pairs, prominulous on both 

by 

short secondary branches or surfaces, secondary venation prominulous and 

peduncles. Receptacle 

campanulate, gray-puberulous on conspicuously reticulate on both surfaces; midrib 

exterior, 

tomentose within; pedicels ca. 1 mm prominulous and flattened on both surfaces, glalong,graypuberulous. 

Calyx lobes 

brous; petioles 3-5 mm long, slightly canalicufive, 

acute, gray-puberulous 

on both surfaces. Petals five, white. late, glabrous. Stipules small, triangular, axillary. 

Stamens 11-12, inserted in a Inflorescences terminal panicles to 13 cm long, 

complete circle; 

the flowers borne in small 

filaments exceeding calyx lobes, free to base. 

groups on short sec- 

Ovary inserted at base of receptacle, almost ondary branches, the rachis puberulous, soon 

glabrous 

with only a few hairs. Style lanate on lower glabrescent, the branches gray-tomentellous. 

Bracts and bracteoles 

portion, equalling filaments in length. Fruit not 

small, ovate, tomentellous, 

seen. 

persistent. Flowers ca. 2.5 mm long. Receptacle 

campanulate, tomentellous on exterior. Calyx 

Distribution (Fig. 51). Known from only one 

lobes small, to 1 mm long, triangular, tomentelcollection 

from Costa Rica and one from Panlous 

on exterior, puberulous within. Petals white, 

ama. 

tomentellous on exterior, sticking together and 

Habitat. Wet forest. 

dehiscing in a calyptra-like mass, 1.2 mm long. 

Additional specimen examined. COSTA RICA. Stamens 13-15, inserted in a complete circle; 

LIMON: Hone Creek, 8 km S of Cahuita, 30 m, 17 Jul filaments far exceeding calyx lobes, free to base, 

1976 (fl), J. & K. Utley 5488 (MO). 

glabrous. Ovary inserted at base of receptacle, 

Licania kallunkiae belongs to subgenus Mo- tomentose on exterior. Style glabrous. Fruit not 

quilea section Moquilea. It is most closely related seen. 

to L. minutiflora from the Guianas and Northern Distribution (Fig. 43). Known only from the 

Amazonia, but it differs in the inflorescence with type gathering from high forest. Flowering in May. 

the flowers borne in small groups on short ter- Licania guatemalensis belongs to subgenus 

tiary inflorescence branches, in the smaller leaves Moquilea section Moquilea, but is not easily conwith 

a more cuspidate acumen, and in the fewer fused with any other species in the section. The 

stamens. Licania kallunkiae has only 11-12 sta- leaves appear quite different from other species 

mens, fewer than any other species of the section, 

in the section in their venation and flattened 

but its exserted stamens and presence of petals, 

midrib with short, thick petiole. It is probably 

as well as its similarity to L. minutiflora, all place 

closest to L. kallunkiae from Panama, but differs 

it in section Moquilea. It differs from another in the much longer leaves with abrupt acumen, 

related and recently described species from Pan- the shorter, thicker petioles, the smaller petals, 

ama in the same section, L. chiriquiensis, in the and the glabrous style. 

smaller flowers with a gray-puberulous not yellow-tomentellous 

indumentum, the spreading, 

2-21.1. Licania cecidiophora Prance, Biotropica 

much longer inflorescences, and the thinner, 

10: 85, fig. 2A-E. 1978. Type. Peru. Amazochartaceous 

leaves with a more cuspidate atten- nas: East of military post Chavez Valdivia, 2- 

uate 12 

apex. 

Feb 1974 (fl bud), Ancuash 752 (holotype, 

NY; isotypes, F, MO). 

2-18.3. Licania guatemalensis Lundell, Wrightia Large tree to 30 m tall, the young branches 

5: 39. 1974; Prance, Acta Amaz6nica 8: 583. 

puberulous, becoming glabrous with age. Leaf 

1978. Type. Guatemala. Izabal: Between Seja lamina oblong, coriaceous, 9-12 x 3-5 cm, 

and Fronteras on Peten-Guatemala Rd., 6 May rounded at base and slightly decurrent into the


petiole, acuminate at apex, the acumen 5-8 mm 

long, glabrous and shining above, with a caducous 

lanate pubescence beneath, becoming 

glabrous with age; midrib prominent on both 

surfaces; primary veins 14-19 pairs, prominulous 

above, prominent beneath; petioles 16-22 

mm long, canaliculate, slightly winged on upper 

portion, tomentellous when young, eglandular. 

trada Belmonte, 12 Sep 1975 (fr), Cordeiro 755 (INPA), 

12 Sep 1975 (fl), Mota & Coelho 86 (INPA). 

2-23. Licania longipetala Prance, Fl. Neotrop. 

Monogr. 9: 62. 1972. 

Fruit narrowly ellipsoid, 6-7 x 1-1.5 cm; exocarp 

smooth and glabrous; mesocarp thin and 

fleshy; endocarp thin, hard and bony, sparsely 

Stipules interpetiolar, small, triangular. Inflorespuberulous 

within. 

cences of racemose panicles (only young ones Distribution. Figure 57. 

seen), the rachis and branches ferrugineous-tomentose. 

Bracts and bracteoles triangular, pu- Additional specimens examined. PERU. LORETO: Rio 

berulous on exterior, with ciliate margins. Flow- Nanay, Quebrada de Mapa Cocha, 23 Feb 1977 (fr), 

Rimachi Y. 2838 

ers buds only seen. 

(NY); Rio Amazonas, Quebrada de 

Receptacle campanulate, Yanayacu, 3 Mar 1977 (fr), Rimachi Y. 2866 (NY). 

brown-tomentose on exterior, lanate-hirsute BRAZIL. AMAZONAS: Tefe, Vila Nogueira, 17 Oct 

within. Calyx lobes five, acute, tomentose on ex- 1975 (fl), D. Coelho & Damiao s.n. (INPA 53295). 

terior. Petals five, puberulous on exterior, with PARA: Rio Itacaiunas, Serra Buritirana, 7 Oct 1970 (fl), 

Pires & Belem 12676 

ciliate margin. Stamens ca. 14, inserted around 

(IAN, NY). RONDONIA: Rio Machado, 

20 Oct 1978 (fr), Goulding 29 (INPA). 

a complete circle; filaments free (short and included 

in buds). Ovary inserted at base of recep- The fruits of this species are eaten by fish 

tacle, sparsely lanate-pubescent. Style glabrous. (Goulding, 1980). 



2-23.1. Licania tachirensis Prance, sp. nov. Type. 

Venezuela. Tachira: Rio San Buena, 10 km W 

Additional specimens examined. PERU. AMAZONAS: 

of La 

N of Cenepa, above Chinkan Entse Creek, 26 Feb 1973 Fundacion, 7?47'N, 71?46'W, 13-15 Mar 

(st), Berlin 902 (MO, NY), Mar 1973 (st), Berlin 976 1980 (fl bud), R. Liesner et al. 9606 (holotype, 

(MO, NY); N of Cenepa along Chinkan Entse Creek, NY; isotypes, MO, VEN). Fig. 5. 

25 Jul 1974 (st), Berlin 1799 (MO, NY). 

L. sectio Moquilea ab omnibus speciebus in- 

Local name. Aguaruna, Jivaro: duship. florescentiis terminalibus, paniculatis e cymulis 

Uses. The small round galls which form on the pluribus 2-4-floris breviter pedunculatis conleaf 

undersurface are used by the Aguaruana In- structis differt. Petiolus 12-16 mm longus, cadians 

as beads for their ceremonial capes (see naliculatus. 

Berlin & Prance, 1978). 

Tree 8 m tall, the young branches glabrous. 

This species belongs either to Licania subge- Leaf lamina oblong, coriaceous, 18-21 x 6.5-8 

nus Moquilea section Moquilea or section Mi- cm, rounded at base, abruptly acuminate at apex, 

crodesmia. Open flowers where the stamen length the acumen ca. 5 mm long, glabrous on both 

is apparent are needed to determine to which of surfaces; midrib prominulous above, prominent 

these two sections it belongs. Licania cecidi- and glabrous beneath; primary veins 13-16 pairs, 

ophora is most distinct, and easily recognized by prominulous above, prominent beneath; petioles 

the long petioles with slightly swollen base and 12-16 mm long, glabrous, rugulose, weakly canby 

the lamina decurrent into its upper portion. aliculate. Stipules not seen. Inflorescences of terminal 

and subterminal much branched panicles, 

2-22. Licania unguiculata Prance, Fl. Neotrop. the flowers in small cymules on short tertiary 

Monogr. 9: 60. 1972. 

branches, the rachis and branches puberulous. 


Bracts and bracteoles triangular, membranous, 

caducous. Receptacle campanulate, tomentel- 

Additional specimens examined. BRAZIL. lous on exterior, tomentose within, sessile. Calyx 

AMAZONAS: Manaus-Caracarai Rd., km 57, 13 Sep 1976 lobes 

(fl), Mota 594 (INPA), km 159,20 Sep 1974 

five, acute, tomentellous on exterior. Petals 

(fl), Prance 

et al. 22721 (INPA, NY), km 27, 19 Mar 1970 (fr), 

five, glabrous. Stamens 12-15, inserted in a com- 

Rodrigues 8770 (INPA); Rio Curicuriari, 3 Nov 1971 plete circle (seen only in bud). Ovary inserted at 

(fl), Prance et al. 16055 (INPA, NY). RONDONIA: Es- base of receptacle, pilose. Fruit not seen.

Systematic Treatment 

B. 

young buds); D, petal. 

31

32 


Habitat. Primary forested area on sandy soil, 

700-1000 m. 

This species is known also from specimens in 

bud. It is quite distinct from any other congener. 

It belongs to section Moquilea, but differs from 

all other species in the large inflorescence with 

the flowers borne in small cymules. The large 

leaves with long canaliculate petioles distinguish 

it from most species. 

Subgenus Moquilea Section Leptobalanus 

2-27.1. Licania granvillei Prance, Proc. Kon. 

Ned. Akad. Wetensch. Ser. C. 89: 114-116. 

1986. Type. French Guiana. Saul, Monts La 

Fumee, 3?37'N, 53012'W, 200-400 m, 21 Aug 

1982 (fl), Mori & Boom 14764 (holotype, NY; 

isotype, CAY). 


puberulous, soon becoming glabrous. Leaf lam- 

ina oblong, thickly coriaceous, 6-13 x 2.3-6 cm, 

cuneate to subcuneate at base, finely acuminate 

at apex, the acumen 10-25 mm long, glabrous 

on both surfaces, without stomatal cavities; mid- 

rib prominent on both surfaces, glabrous; pri- 

mary veins 12-16 pairs, prominulous on both 

surfaces, glabrous; petioles 5-8 mm long, can- 

aliculate above, glabrous, eglandular, rugulose. 

Stipules caducous (not seen). Inflorescences of 

racemose panicles 6-12 cm long, the rachis and 

branches sparsely appressed pubescent, appear- 

ing dark because pubescence does not form a 

complete covering. Bracts and bracteoles mi- 

nute, 0.25 mm long, membranous. Flowers 2-3 

mm long, sessile along primary branches of in- 

florescence. Receptacle campanulate, sparsely 

puberulous to glabrous on exterior, densely to- 

mentose within, sessile. Calyx lobes five, acute, 

puberulous on both surfaces, with tomentellous 

margins. Petals absent. Stamens 12, inserted 

around complete circle, the filaments exserted 

beyond calyx lobes, glabrous, but densely to- 

mentose around base, free to base. Ovary in- 

serted at base of receptacle, sparsely lanate. Style 

exserted, sparsely tomentose for half of length. 

Fruit globose to ellipsoid, 5.5 cm long x 3.5-4 

cm broad, glabrous and lenticellate on exterior; 

mesocarp fleshy, 3-4 mm thick; endocarp hard, 

bony, 1 mm thick, glabrous within. 


Distribution (Fig. 42). Upland forest on terra 

firme in hills from Colombia to French Guiana. 

Specimens examined. COLOMBIA. AMAZONAS: Rio 

Apaporis, Raudal Yayacopi, 0?5'S, 70?30'W, 800', 18 

Feb 1982 (fr), Schultes & Cabrera 15456 (NY); Jinogoje, 

700', 27 Feb 1982 (fr), Schultes & Cabrera 15660 

(NY). 

VENEZUELA. AMAZONAS: Trail south from Cerro 

Neblina base camp on Rio Mawarinuma from clear 

water stream to top of first small hills, 0?50'N, 66? 1 'W, 

3 May 1984 (fl, fr), Gentry & Stein 47121 (NY). 

FRENCH GUIANA. Saul, Monts La Fum6e, 200- 

400 m, 26 Aug 1982 (fl), Mori & Boom 14772 (CAY, 

NY), 28 Aug 1982 (fl), Mori & Boom 14790 (CAY, 

NY), 1 Sep 1982 (fl), Mori & Boom 14828 (CAY, NY), 

28 Mar 1983 (fr), Mori & Pipoly 15428 (CAY, NY), 1 

Apr 1983 (fr), Mori & Pipoly 15476 (CAY, NY), 16 

Apr 1983 (fr), Mori & Pipoly 15523 (CAY, NY). 

BRAZIL. AMAZONAS: Serra da Neblina, between Palmito 

and Tatu Camp, 400-600 m, 21 Dec 1965 (fr), 

Silva & Brazao 60697 (NY). 

This species is close to Licania apetala, es- 

pecially to its var. aperta, but differs in the larger 

flowers, the larger, thicker leaves with a much 

more attenuate apex, and the minute bracteoles. 

The Brazilian and Colombian material in fruit 

was kept aside as a possible new species at the 

time of my monograph (Prance, 1972), and it is 

now possible to describe it with the new material 

from French Guiana. This species is known in 

both localities from the hill slopes of mountains. 

It is named for J-J. de Granville who has done 

much to interpret the flora of French Guiana. 

2-31.1. Licania jefensis Prance, Brittonia 28: 

215-216, fig. 5.1976. Type. Panama. Panama: 

Summit of Cerro Jefe, 2 Apr 1969 (fl), Dwyer 

et al. 5047 (holotype, NY; isotype, MO). 

Shrub to 2 m tall, the young branches tomentose, 

soon becoming glabrous. Leaf lamina elliptic, 

coriaceous, 3-6 x 1.8-3.5 cm, rounded at 

base, acute to acuminate at apex, the acumen 3- 

5 mm long, glabrous on both surfaces when mature, 

when young with a lanate, caducous indumentum 

which is more persistent beneath, with 

scattered glands on lower surface, without stomatal 

cavities, midrib prominent on both surfaces; 

primary veins 8-11 pairs, prominulous 

above, prominent beneath; secondary venation 

conspicuously prominulous and reticulate on both 

surfaces; petioles 2.5-4 mm long, tomentose, 

eglandular, rugulose, slightly canaliculate. Stip-


ules small, axillary, caducous. Inflorescences of 

terminal and subterminal racemose panicles, 6- 

11 cm long, the rachis and branches yellowbrown-tomentose. 

Bracts and bracteoles ca. 0.5 

mm long, tomentose on exterior, persistent. 

Flowers ca. 2 mm long. Receptacle campanulate, 

sessile, tomentose on exterior, pilose within. Calyx 

lobes five, tomentose on exterior, puberulous 

within. Petals absent. Stamens ca. 13, inserted 

around complete circle; filaments far exserted, 

free to base. Ovary pilose. Style glabrous, equalling 

filaments in length. Fruit globose, ca. 25 cm 

in diam. epicarp smooth and glabrous; mesocarp 

thin and fleshy; endocarp hard, bony, thin. 

Distribution (Fig. 50). Known only from the 

cloud forests of Panama, flowering in March and 

April. 

Ovary pilose. Style glabrous except around base, 

equalling filaments in length. Fruit unknown. 


type gathering from moist forest at 350 m, col- 

lected in flower in February. 

2-32. Licania sparsipilis Blake, Contr. Gray 

Herb. 52: 67. 1917. 


Additional specimens examined. MEXICO. OAXACA: 

30 km S ofrd. Palomares to Uxpanapa, between arroyo 

Humaca and Rio Verde, 300 m, 20 Apr 1985 (fl), W. 

Thomas et al. 3575 (NY). 

EL SALVADOR. AHUACHAPAN: Rio El Venado, Finca 

San Benito, 6 Mar 1979 (y fr), Castro s.n. (NY). 

PANAMA. COLON: Santa Rita Ridge, 2 Mar 1975 

(fl), Mori & Kallunki 4914 (MO, NY). 

Additional specimens examined. PANAMA. PANAMA: 2-32.1. Licania cuatrecasasii Prance, Acta Ama- 

9.4 km N of Goofy Lake, 900 m, 11 Mar 1977 (fl), zonica 8: 577. 1978. Type. Colombia. Valle: 

Folsom et al. 1996 (MO, NY); Cerro Jefe, 1000 m, 11 

Alto 

Jun 1975 (y fr), Mori 6532 (NY), 29 Aug 1975 Yunda, Rio Anchicaya, 1000 m, Oct 1972 

(fr), 

Mori 7990 (MO, NY). 

(fl), Hilty 0-1 (holotype, US; isotype, NY). 

Tree to 30 m tall, the young branches puber- 

2-31.2. Licania morii Prance, Brittonia 28: 215, 

ulous, soon glabrate. Leaf lamina elliptic, corifig. 

4.1976. Type. Panama. Panama: El Llano- aceous, 8-12.5 x 3.5-8 cm, subcuneate at base, 

Carti Rd., 12 km from Inter American Hwy., 

cuspidate at apex, the acumen 10-15 mm long, 

15 Feb 1975 (fl), Mori & Kallunki 4665 (hoslightly 

curved, glabrous above, with a compact 

brown-lanate 

lotype, NY; isotype, MO). 

pubescence beneath, without stomatal 

cavities; primary veins 10-12 pairs, prom- 

Tree to 15 m tall, the young branches tomen- inent beneath, prominulous above; midrib 

tellous, soon becoming glabrous. Leaf lamina prominent on both surfaces; petioles 8-11 mm 

elliptic, coriaceous, 7-11 x 2.7-5.5 cm, subcor- long, tomentellous when young, terete or slightly 

date at base, acuminate at apex, the acumen 8- canaliculate, eglandular, transversely rugulose. 

12 mm long, glabrous and shiny above, with a Stipules caducous (not seen). Inflorescences of 

short glaucous appressed pubescence beneath and racemose panicles usually once-branched, ocwith 

conspicuous palisade glands, without sto- casionally with secondary branches up to 10 cm 

matal cavities; midrib prominulous above, long, the rachis and branches brown-tomentelprominent 

beneath; primary veins 7-9 pairs, lous. Bracts and bracteoles ovate, ca. 1 mm long, 

prominulous above, prominent beneath; petioles persistent, tomentellous on exterior, entire, 

3-6 mm long, sparsely tomentellous, weakly can- eglandular. Flowers ca. 2.5 mm long, sessile on 

aliculate. Stipules caducous, not seen. Inflores- primary and secondary branches of inflorescences 

of terminal or subterminal racemose pan- cence. Receptacle broadly campanulate, tomenicles, 

7-12 cm long, the rachis and branches tose on exterior, pilose within. Calyx lobes five, 

sericeous-tomentellous. Bracts and bracteoles acute, tomentellous on exterior, puberulous 

triangular, eglandular, ca. 1 mm long, caducous. within. Petals absent. Stamens 10-12, inserted 

Flowers ca. 2 mm long. Receptacle campanulate, in a complete circle; filaments far exceeding calyx 

sessile or with short pedicel to 1.5 mm long, lobes, free to base, glabrous. Ovary inserted at 

tomentose on exterior, pilose within. Calyx lobes base of receptacle, villous around base, but glafive, 

acute, tomentellous on both surfaces. Petals brous above. Style glabrous, equalling filaments 

absent. Stamens ca. 13, inserted around com- in length. Fruit not seen (25-30 mm long acplete 

circle, the filaments far-exserted, free to base. cording to field notes).


Distribution (Fig. 34). Known only from up- Licania mexicana was described from poor 

land forest of El Valle, Colombia. 

material with only old flowers present, making 

it difficult to relate to other 

Additional specimen examined. COLOMBIA. VALLE: 

species. It belongs to 

Bajo Calima, 15 km N of Buenaventura, 18 Feb 1983 subgenus Moquilea, either section Moquilea or 

(fl), Gentry & Juncosa 40467 (MO, NY). 

section Leptobalanus, depending on the presence 

or absence of 

This species from the 

petals, which cannot be observed 

highlands of Valle comes 

in the old flowers. It 

from an area in need of 

probably 

further 

belongs to section 

exploration. It is 

Leptobalanus and seems to be most 

most closely related to Licania 

closely reapetala 

and L. 

lated to the Central American 

sparsipilis, but differs from both in the 

species L. spardense 

sipilis. It differs in the short inflorescence branchbrown-lanate 

pubescence of the leaf undersurface, 

which is set in 

es, the less acuminate leaves with shorter 

the 

petioles, 

deeply reticulate secand 

the 

ondary and tertiary venation, as well as in 

greater number of stamens. 

the 

leaf's long cuspidate acumen, the larger petioles 

without the two glands ofL. sparsipilis, the brown- 2-33.1. Licania joseramosii Prance, Acta Amatomentellous 

pubescence of the inflorescences, z6nica 8: 581, fig. 3. 1978. Type. Brazil. Amaand 

the very small, broadly campanulate recep- zonas: Manaus-Caracarai Rd., km 130, 6 Jan 

tacle. 

1976 (fl), Monteiro & Ramos 29 (holotype, 

INPA 54340; isotype, NY). 

2-32.2. Licania mexicana Lundell, Wrightia 5: 

40. 1974; Prance, Acta Amazonica 8: 585. 

1978. Type. Mexico. Sinaloa: Between Rancho 

Del Pinio and Chele, 22 May 1943 (fl), Lundell 

13023 (holotype, LL; isotype, MICH). 

Tree 10 m tall, the young branches lanate to 

puberulous and soon glabrous. Leaf lamina nar- 

rowly oblong to lanceolate, coriaceous, 6.5-13 x 

2.5-4 cm, cuneate at base, gradually attenuate to 

acute apex, glabrous above, with a caducous lan- 

ate pubescence beneath when young, stomatal 

cavities absent; midrib prominulous above, 

prominent beneath; primary veins 6-8 pairs, 

prominulous on both surfaces, secondary vena- 

tion prominulous and conspicuously reticulate 

on both surfaces; petioles 1.5-3.5 mm long, ru- 

gose, terete, lanate when very young, soon be- 

coming glabrous. Stipules axillary, triangular, 

persistent, 2 mm long, lanate when young. In- 

florescence of terminal panicles 2-3.5 cm long, 

with short primary branches (only 2 old inflo- 

rescences seen) gray-brown, the rachis and 

branches tomentellous. Bracts and bracteoles 

small, ca. 1 mm long, persistent, tomentellous, 

ovate-triangular. Receptacle campanulate, to- 

mentellous on exterior, pilose within. Calyx lobes 

triangular, 1 mm long, acute, reflexed, tomen- 

tellous. Petals? (old flowers only present). Sta- 

mens 14-15, inserted around complete circle, 

exserted beyond calyx lobes; filaments glabrous, 

inserted at base of receptacle. Style glabrous. Fruit 

not seen. 

Distribution (Fig. 60). This species is known 

only from the type. 

Small tree 5 m tall, the young branches glabrous. 

Leaf lamina oblong to oblong-lanceolate, 

coriaceous, 13-20 x 4-6.5 cm, cuneate at base, 

finely acuminate at apex, the acumen 15-20 mm 

long, glabrous on both surfaces, without stomatal 

cavities; primary veins 9-14 pairs, prominulous 

on both surfaces; midrib prominent on both surfaces; 

petioles 4-5 mm long, rugulose tomentellous 

when young, terete, with two glands near 

junction with lamina. Stipules linear, ca. 6 mm 

long, hispidulous, caducous. Inflorescences of 

panicles with long thick central rachis and short, 

thin, lateral branches bearing 1-3 flowers, the 

rachis and branches tomentellous. Bracts and 

bracteoles lanceolate, subpersistent, tomentellous 

on exterior, glabrous within, entire, with 

long, thin acumen, eglandular. Flowers ca. 5 mm 

long. Receptacle campanulate, gray-tomentose 

on exterior, tomentose within. Calyx lobes five, 

acute, tomentose on exterior. Petals absent. Stamens 

ca. 19, inserted around complete circle; 

filaments slightly exceeding calyx lobes, free almost 

to base, glabrous except for pilose annular 

ring. Ovary inserted at base of receptacle, lanate. 

Style glabrous, equalling filaments in length. Fruit 

not seen. 

Licania joseramosii (distribution in Fig. 51), 

is a most distinct species that cannot be easily 

confused with any other in the genus. It is related 

to L. emarginata and L. calvescens but differs in 

a large number of characters such as the much 

longer leaves, the larger flowers, and the distinctive 

inflorescence. It differs from L. emarginata 

in the greater number of stamens, and from L.


calvescens in the flowers borne in small groups 

on secondary inflorescence branches and in the 

glabrous leaves. Superficially L. joseramosii also 

resembles L. longipedicellata in subgenus Moquilea 

section Moquilea but differs in the smaller, 

thicker leaves, the much less branched inflorescence, 

the smaller flowers, and the absence of 

petals. 

2-34. Licania calvescens Cuatrecasas, Fieldiana, 

Bot. 27: 64. 1950. 

This distinct species was known only from the 

type from El Valle in Prance (1972). Since then 

I have seen two further sterile inventory collections 

that may be referred to this Pacific coastal 

forest species (Fig. 30). 

2-39. Licania longistyla (Hooker f.) Fritsch, Ann. 

K. K. Naturhist. Hofmus. 4: 56. 1889. 

This species, common in Venezuela, the 

Guianas, and western and central Amazonia, has 

recently been collected in Panama; yet another 

Panama-Amazon disjunct. See Figure 58. 

Additional specimen examined. PANAMA. SAN BLAS: 

El Llano-Carti Rd., 9?19'N, 78?55'W, 300 m, 8 Jan 

1985 (fl), de Nevers et al. 4446 (MO, NY). 

2-43. Licania octandra (Hoffmannsegg ex Roemer 

& Schultes) Kuntze, Revis. gen. pl. 217. 

1891. Fig. 68. 

A new subspecies was described in Prance 

(1974a). The three subspecies may be distin- 

guished by the following key. 

1. Leaves 3-12 x 2-4 cm, the apex obtuse to acu- 

minate, the acumen 1-13 mm long. 

2. Leaves broadly ovate to oblong, obtuse to 

bluntly acuminate, the acumen 1-5 mm long; 

upper surface of leaf drying brown; young in- 

florescence with sparse gray-brown tomen- 

tum. a. subsp. octandra. 

2. Leaves oblong-lanceolate with a well-devel- 

oped finely pointed acumen 5-13 mm long; 

upper surface of leaf drying gray or green; 

young inflorescence usually with a rufous- 

brown arachnoid indumentum. 

b. subsp. pallida. 

1. Leaves 14-29 x 4.5-7 cm, long-acuminate at 

apex, the acumen 12-28 mm long. 

c. subsp. grandifolia. 

2-43c. L. octandra subsp. grandifolia Prance, 

Acta Amazonica 4(1): 18. 1974. Type. Brazil. 

Amazonas: Rio Javari, behind Estirao de 

Equador, 9 Aug 1973 (fl), Lleras et al. P17270 

(holotype, INPA; isotype, NY). 

Additional specimens examined. COLOMBIA. 

CHoc6: Trail Tubad6 to Quibd6-Tutunendo Rd., 17 Specimens examined. COLOMBIA. AMAZONAS: Rio 

Jan 1979 (st), Gentry & Renteria A. 24331 (MO, NY). Loreto-Yacu (st), Glenboski 206 (NY). PERU. LORETO: 

VALLE: Bajo Calima, N of Buenaventura, lower Rio Rio Tacha, Curaray, 18 Sep 1972 (fl), Croat 20372 

San Juan, 8 Dec 1981 (st), Gentry 35478 (MO, NY). (AAU, MO, NY); Maynas, Mishana, Rio Nanay halfway 

between Iquitos and Santa Maria de Nanay, 31 

2-38. Licania albiflora Fanshawe & Maguire, May 1978 (st), Gentry et al. 22387 (NY); Maynas, Dtto. 

Fernando 

Bull. Torrey Bot. Club 75: 318. 1948. 

Lores, Quebrada Tamshiyacu, 10 Jan 1977 

(fr), McDaniel & Rimachi 21133 (NY); Maynas, Iqui- 

This species was known from two collections tos, Carretera de Pefia Negra, 10 Feb 1977 (fr), Rimachi 

Y 2797 

from Guyana and Surinam in Prance (1972). A (NY); Prov. Loreto: Nauta, Rio Marafion, 9 

Nov 1982 (fl buds), Vdsquez & Jaramillo 3444 (NY); 

third collection from French Guiana has recently Maynas, Pto. Almendras (Rio Nanay), 19 Feb 1985 (fl 

been added, showing it to be rare but widespread buds), Vasquez & Jaramillo 6243 (NY). 

in the Guianas. See Figure 23. 

Additional specimen examined. FRENCH GUIANA. Subgenus Moquilea Section Microdesmia 

Saiil, La Fumee Trail, 27 Mar 1983 (fr), Mori & Pipoly 2-45. Licania arborea Seemann, Bot. 

15411 

voy. Her- 

(NY). 

ald 3: 118,t. 25. 1853. 

This species, common from Mexico through 

Central America, was known by only three col- 

lections from South America at the time of Prance 

(1972). It is apparently more widespread in South 

America, although it may well have been intro- 

duced there by indigenous traders because of its 

use as an oilseed. 



CHOCO: Mun. Riosucio, Parque Nacional Los Katios, 

1 Dec 1976 (fr), Le6n 632 (MO). 

VENEZUELA. ZULIA: Aricuiza, 19 Dec 1972 (fl), 

Veillon 131 (US, VEN) [possibly cultivated]. 

PERU. HUANUCO: Prov. Pachitea, Rio Pachitea nr. 

Miel de Abeja, 9 Mar 1967 (fr), Schunke V 1737 (COL, 

US); Dist. Honoria, Caserio Leoncio Prado, 29 Nov 

1963 (fl), Lao Magin 103 (F). LORETO: Maynas, trail 

from Indiana on Rio Amazonas to Rio Napo, 24 May 

1978 (fl), Gentry et al. 22199 (MO, NY). 

BRAZIL. ACRE: Rio Macaua, Mun. Sena Madureira 

(fl), Lima & Souza 231 (INPA, NY).


2-47. Licania subarachnophylla Cuatrecasas, 

Fieldiana, Bot. 27: 110. 1951. 

Distribution (Fig. 80). This little-collected 

species, known only from the type and one other 

collection, from Boyaca and Meta in Colombia, 

has now been found in Venezuela. It seems to 

be endemic to the gallery forests along the rivers 

of the savannas. 

subarachnophylla in the subgenus Moquilea section 

Microdesmia. These four are a closely related 

group of species each well separated from 

one another geographically or ecologically. Licania 

salicifolia is a montane species of Antioquia, 

Colombia, which differs in its much longer 

inflorescences with flowers in clusters and in the 

longer lanceolate leaves; L. subarachnophylla is 

a species of the gallery forests of the Llanos which 

Additional specimen examined. VENEZUELA. also differs in the much longer inflorescence with 

APURE: Distr. Pedro Camejo, along Quebrada El Porve- the flowers in clusters and in its smaller flowers; 

nir, 6?39'N, 67017'W, 21 Feb 1979 (fl), Davidse & Gon- L. araneosa is a species of the gallery forests of 

zalez 15541 (MO, NY). 

a restricted area of the planalto of Central Brazil, 

which differs in its smaller flowers, larger 

2-47.1. Licania tambopatensis Prance, sp. nov. branched inflorescences, and more coriaceous 

Type. Peru. Madre de Dios: Tambopata Na- leaves. 

ture Reserve, Laguna Coco Cocha, 5.2 km east This species is named for the Tambopata Naof 

lodge, 3 Jun 1986 (fl), V. A. Funk, B. Kahn ture Reserve, a remarkable sanctuary for the 

& S. Wiser 8415 (holotype, NY; isotype, species diversity of the upper Amazon region 

US). Fig. 6. where the highest known level of species diver- 

Ab L. araneosa floribus 2 mm longis, inflo- sity has been recorded for many organisms. 

rescentiis racemosis, foliis chartaceis differt. 

Shrub, the young branches tomentose becom- 

2-48. Licania salicifolia Cuatrecasas, Fieldiana, 

ing glabrous with age. Leaf lamina oblong, char- 

Bot. 27: 111. 1951. 

taceous, 3.8-6 x 1.5-2.4 cm, cuneate at base, This species was known from a single rather 

mucronate at apex, the mucro 1-2 mm long, gla- poor type specimen from Antioquia, Colombia. 

brous above except on midrib, appressed lanate- It is a high altitude endemic still only known 

pubescent beneath; midrib prominent beneath, from the type locality in the vicinity of Rionegro 

prominulous and tomentose above when young; (not the Rio Negro as erroneously stated in 

primary veins 11-14 pairs, prominulous be- Prance, 1972!). Espinal T. (1981) wrote a comneath, 

plane above; petioles 2-4 mm long, to- mentary on the rarity of this species and pointed 

mentose, terete, eglandular. Stipules linear, ax- out that residents of Comfama know this as "the 

illary, caducous, membranous. Inflorescences rare tree." See Figure 76. 

of terminal and axillary racemes, 2-4 cm, the 

rachis tomentose. Bracts and bracteoles mem- Additional specimens examined. COLOMBIA. 

ANTIOQUIA: Rionegro, Hacienda 

branous, caducous, triangular, tomentellous on 

Comfama, Jul 1981 

(st), Espinal T. et al. 4537 (MEDEL), 3 Dec 1981 (fl), 

exterior, glabrous within. Flowers 2-2.5 mm long, Espinal T. & A. Lenna T. 4608 (MEDEL), 3 July 1981 

sessile, densely packed along rachis but not clus- (fl), R. Jaramillo s.n. (COL, NY). 

tered. Receptacle cupuliform, sessile, brown-tomentose 

and lanate on exterior, densely tomen- Subgenus Licania Section Hirsuta 

tose within. Calyx lobes acute, triangular, lanate 

on both surfaces. Petals small, caducous, the 2-53. Licania hirsuta Prance, Fl. Neotrop. 

margins ciliate. Stamens ca. 15, inserted around Monogr. 9: 93. 1972. 

complete circle, equalling or only slightly ex- This species was described from two collecceeding 

the calyx lobes, glabrous, free to base. tions in Central Amazonas, Brazil, and has now 

Ovary inserted at base of receptacle. Style gla- also been found in Amapa. See Figure 46. 

brous except at base. Fruit not seen. 

Distribution (Fig. 62). Known only from the Additional specimens examined. BRAZIL. AMAPA: 

type collected 

Road 

along the edge of the lake. 

Tinguilim to BraSo, km 21, 2 Jun 1969 (fl), N. 

T. Silva 2096 

This 

(IAN, NY). AMAZONAS: Mun. 

new 

Humaita, 

species belongs close to the species rd. Humaita-Porto Velho, km 70, 5 May 1982 (fl), 

group of Licania araneosa, L. salicifolia, and L. Teixeira et al. 253 (INPA, NY).


ii~i""~~~~mm 

A. 

i K 

N.N 

FIG. 6. Licania tambopatensis (Funk et al. 8415). A, habit; B, leaf undersurface; C, inflorescence; D, bract 

and bracteole; E, flower; F, flower section; G, ovary. 

2-57.1. Licania hispida Prance, sp. nov. Type. 

Venezuela. Amazonas: Dept. Rio Negro, Ce- 

rro Aratitiyope, 2?10'N, 65?34'W, SSW of 

Ocamo, 990-1670 m, 24-28 Feb 1984 (fl, fr), 

J. A. Steyermark et al. 130185 (holotype, 

NY). Fig. 7. 

Species sectione Hirsutae pertinens, in qua ra- 

mulis juvenilibus, costis petiolis que hispidis, 

rachidibus inflorescentiarum basim versus his- 

pidis unica est. 

Tree 6 m tall, the young branches densely his- 

pid. Leaf lamina oblong to oblong-elliptic, co- 

riaceous, 7-11 x 2.5-5.2 cm, rounded and un- 

equal at base, apiculate or bluntly acuminate at 

apex, sparsely hispid on primary venation be- 

neath, otherwise glabrous on both surfaces; mid- 

rib prominulous above, prominent beneath, his-


,. '~~ 1cm.Y~gBB~~ 

12cm. 1 

cmEE;cm. 

G~~. D . "" B. ,,~~0 

FIG. 7. Licania hispida (Steyermark 130185). A, habit; B, stem and petiole; C, leaf undersurface; D, flower; 

E, flower section; F, young infructescence; G, young fruit. 

pid beneath especially on lower portion; primary 

veins 9-12 pairs, plane above, prominent be- 

neath, the lower ones sparsely hispid; petioles 4- 

7 mm long, terete, eglandular, hispid. Stipules 

linear, 5-6 mm long, subpersistent. Inflores- 

cences of little-branched panicles, the lower por- 

tion of rachis hispid, sparsely tomentellous on 

upper portion and on branches. Bracts and brac- 

teoles 1 mm long, lanceolate, persistent, tomen- 

tellous. Flowers in bud ca. 1.5 mm long, sessile 

on primary branches of inflorescence. Receptacle 

campanulate, sessile, gray-tomentose on exteri-


or, tomentose within. Calyx lobes five, acute, 

tomentose on both surfaces. Petals five, minute. 

Stamens five, inserted around complete circle; 

filaments shorter than calyx lobes, free, glabrous. 

Ovary inserted at base of receptacle, lanate. Style 

glabrous except at extreme base. Fruit (immaposite 

to three calyx lobes, shorter than calyx 

lobes, the filaments connate for lower third. Ovary 

pilose. Style pilose. Fruit unknown. 


type collection from 1400 m, collected in flower 

in May. 

ture) globose, exocarp ferrugineous-pubescent; 

endocarp thin, hard. 

2-61. Licania arachnoidea Fanshawe & Ma- 


type from forest at base of high bluffs of an igneous 

rock mountain. 

This new species belongs to section Hirsuta 

and is closely related to the other species in that 

section and in section Hymenopus. It is quite 

distinct nonetheless, and easily distinguished by 

the hispid pubescence of the stems, petioles, midrib 

and lower part of the inflorescence rachis. 

This type of hispid pubescence, common in the 

genus Hirtella, does not occur in other species 

of Licania. Licania hispida, with acute or only 

bluntly acuminate leaves is also distinguished 

from related species by the leaf shape. 

guire, Bull. Torrey Bot. Club 75: 318. 1948. 

Distribution (Fig. 25). A recent collection of 

this species was made in Loreto Dept., Peru. Previously 

it was known only from the Guianas and 

from a single collection from western Brazil. It 

adds to the growing list of species disjunct in 

eastern and western Amazonia that are not found 

in Central Amazonia, such as Licania guianensis 

and Couepia parillo. 

Additional specimen examined. PERU. LORETO: 

Maynas, Hondococha nr. Santa Cecilia, Rio Maniti, 

11 Nov 1976 (fl), Encarnaci6n 970 (NY, US). 

2-64. Licania caudata Prance, Fl. Neotrop. 

Monogr. 9: 103. 1972. 

Subgenus Licania Section Hymenopus 

2-60.1. Licania pakaraimensis Prance, Brittonia 

28:218, fig. 6. 1976. Type. Venezuela. Bolivar: 

Sierra Pakaraima, headwaters of Rio Paragua, 

3?40'N, 63?0'W, May 1973 (fl), Steyermark 

107357 (holotype, NY; isotype, VEN). 

Fruit oblong, 25 x 8 mm; exocarp smooth and 

glabrous, often slightly rugulose, wrinkled when 

dry; mesocarp thin, 0.25 mm thick, fleshy; en- 

docarp thin 0.5 mm thick, bony, slightly pu- 

berulous within. 


Tree to 10 m tall, the young branches VAUPES: 

sparsely Mitfi, 25 Apr 1975 (fr), Zarucchi 1278 (COL). 

FRENCH GUIANA. 

tomentellous, soon glabrate and lenticellate. Saul, Monts La Fumee, 12 Oct 

Leaf 1982 (st), Boom & Mori 1997 (NY). 

lamina narrowly oblong, coriaceous, 7.5-11.5 x PERU. LORETO: Rio Nanay above Bellavista, 1 Jun 

2.5-4.5 cm, cuneate at base, acuminate at apex, 1976 (fl), Rimachi Y. 2326 (NY); Dept. Sapuena, Arthe 

acumen 3-12 mm long, glabrous on both boretum Jenaro Herrera, 4 Jun 1974 (fl), Diaz 51-A 

surfaces; primary veins 11-17 pairs, plane above, (IAN); Pucallpa, 1 Aug 1980 (fr), Salazar 662 (MO, 

prominulous and glabrous beneath; petioles 3-5 

NY), 6 Jun 1960 (fl), Woytkowski 5773 (S, US). UCAYALI: 

Km 86 Pucallpa-Tingo Maria Rd., Jul 1978 (fr), Froehmm 

long, terete, rugulose, glabrous or with a few ner 362 (MO, NY). 

appressed hairs, eglandular. Stipules axillary, to- BRAZIL. ACRE: Cruzeiro do Sul, 12 Feb 1976 (fr), 

mentellous, caducous. Inflorescences of terminal Marinho 134 (IAN). AMAZONAS: Rio Cuieiras, nr. Lago 

de 

panicles with flowers grouped in small Peixe-Boi, Sep 1972 (fr), O. Pires & Honda 185 

cymules 

(INPA, NY); Manaus-Porto Velho Rd., between Rio 

on peduncles 1 mm long, the rachis and branches Castanho and Araca, 10 Oct 1974 (st), Prance et al. 

tomentellous, the rachis ca. 2 mm thick at base. 22775 (INPA, NY), 12 Jul 1972 (fl), M. F. da Silva 

Bracts and bracteoles ca. 0.5 mm long, triangular, 486 (INPA, NY). PARA: Rd. Oriximina to Obidos, km 

persistent. Flowers ca. 2.5 mm 70 at Rio 

long. Receptacle 

Cumina-Mirim, 14 Sep 1980 (fr), Cid et al. 

2545 (INPA, NY). 

campanulate, gray-tomentellous on exterior, tomentellous 

within; pedicels 0.25 mm long. Calyx This species was described from three colleclobes 

five, acute, gray-tomentellous on exterior, tions from the vicinity of Manaus. It has turned 

glabrous within. Petals five, caducous, remaining out to be a widespread and common Amazonian 

attached to one another and caducous in a single species, occurring in both inundated and terra 

calyptra-like unit. Stamens five, unilateral, op- firme forest. See Figure 31.


2-64.1. Licania miltonii Prance, Acta Amazo- 

nica 13: 24. 1983. Type. Brazil. Mato Grosso: 

Aripuana, km 238 ofrd. BR 174, Nuicleo Juina, 

area urbana, 17 Jan 1979 (fl), M. G. Silva & 

A. Pinheiro 4296 (holotype, MG; isotype, NY). 

Additional 

specimen examined. BRAZIL. MATO 

GROSSO: Aripuana, Nuicleo Juina, 28 May 1978 (fr), 

M. G. Silva & Rosario 4684 (MG). 

This species, most closely related to Licania 

caudata, differs in the much smaller, narrower, 

chartaceous leaves with cuspidate not caudate 

apices and shorter petioles, the smaller flowers 

with fewer stamens with free filaments, and 

smaller inflorescences. 

2-65. Licania latistipula Prance, Fl. Neotrop. 

Monogr. 9: 103. 1972. 

This species, with distinctively large stipules, 

was described from two collections in the Ori- 

noco Delta of Venezuela but also occurs in French 

Guiana. See Figure 54. 

Additional specimen examined. FRENCH GUIANA. 

Piste de St. Elie, 19 Oct 1977 (st), Lescure 758 (NY). 

Tree 5 m tall, the young branches sparsely puberulous, 

soon becoming glabrous. Leaf lamina 

narrowly oblong, chartaceous, 4-10.2 x 1.3-3.8 

cm, cuneate at base, cuspidate-acuminate at apex, 

the acumen 6-12 mm long, glabrous on both 

surfaces; midrib slightly prominulous above, 

prominent beneath, with a few stiff appressed 

hairs on both surfaces; primary veins 6-8 pairs, 

plane above, slightly prominulous beneath; petioles 

1-3 mm long, terete, eglandular, with few 

sparse appressed hairs. Stipules intrapetiolar, linear, 

persistent, ca. 2 mm long. Inflorescences terminal 

and axillary little-branched panicles 1-3 

cm, the rachis and branches very sparsely hirsutulous. 

Bracts and bracteoles minute, membranous, 

sparsely hirsutulous on exterior, persistent, 

entire. Flowers minute, 1-1.5 mm long, 

borne solitary on short primary branches of inflorescence. 

Receptacle campanulate, glabrous or 

sparsely hirsutulous on exterior, tomentose on 

exterior; pedicels 0.2 mm long. Calyx lobes five, 

acute, glabrous except for a few stiff appressed 

hairs on exterior, the margins ciliate. Petals five, 

tomentellous within towards apex, glabrous beneath, 

puberulous on outer surface. Stamens five, 

fertile, inserted on short thick filaments, connate 

at base. Ovary inserted at base of receptacle, 

sparsely hirsute. Fruit ellipsoid, ca. 2 cm long x 

2-67. Licania glabriflora Prance, Fl. Neotrop. 

Monogr. 9: 104. 1972. 

Distribution (Fig. 41). This species, described 

from Venezuela and French Guiana in Prance 

(1972), has now been collected in adjacent Brazil 

and Surinam, but also in Costa Rica, giving it a 

disjunct distribution similar to that of L. affinis. 

The Central American material differs only in 

having three rather than five stamens and often 

four rather than five petals. In all other respects 

it resembles the South American specimens studied. 

Additional specimens examined. COSTA RICA. 

HEREDIA: Finca La Selva, Rio Puerto Viejo, 14 Feb 

1982 (fl), Hammel 11149 (NY). 

SURINAM. Lely Mountains, SW plateau, 29 Sep 

1975 (fl), Lindeman & Stoffers 525 (S). 

FRENCH GUIANA. Riv. Grand Inini, Saint Batardeau, 

11 Sep 1970 (fl), Granville B3787 (CAY). 

BRAZIL. AMAPA: Between Porto Platon and Serra 

do Navio, Oct-Dec 1976 (st), Rosa 1165 (MG). 

Note that in Prance (1972: 105) this species 

was erroneously cited as Licania glabrifolia. 

2-69a. Licania heteromorpha Bentham, J. Bot. 

(Hooker) 2: 221. 1840. var. heteromorpha. 

1 cm broad; exocarp smooth, glabrous; mesocarp 

thin; endocarp 0.5 mm thick, glabrous within. 

Distribution (Fig. 64). Forest on terra firme, 

clay soil in Mato Grosso. 

This is the commonest and most collected 

Amazonian species of Licania. In Prance (1972) 

I also cited material from Rio de Janeiro for 

which I was uncertain if it was of cultivated or- 

igin or for the Glaziou specimens pirated. A new 

collection shows that this species is a native of 

the eastern Brazilian forests and is another Am- 

azon-eastern Brazil disjunct (Fig. 44). 

Additional specimen examined. BRAZIL. ESPIRITO 

SANTO: Mun. Linhares, Res. Florestal da Cia Vale do 

Rio Doce, 14 Dec 1981 (fl), H. C. de Lima 1660 (NY, 

RB). 

2-69e. Licania heteromorpha Bentham var. re- 

voluta Prance, Acta Amazonica 13: 24. 1983. 

Type. Brazil. Amazonas: 20 km NW of Ma- 

naus, Taruma development area, 21 Mar 1981 

(fl), B. W. & S. P. Nelson 1058 (holotype, INPA; 

isotype, NY). 

Leaves 2-4.5 x 1.1-2.3 cm, the margins revo- 

lute; anthers deltoid.


, _A. . . _ . 

=:z- ~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~l-i 

.. 

0-1~~~~~~~ 

E. C~~~~~~~~i .~ 

FIG. 8. Licania laevigata (Irwin et al. 54990; M. Silva 80). A, habit; B, flower; C, flower section; D, petal; 

E, ovary; F, floral diagram; G, young fruit. 

Habitat. Forest on terra firme, clay soil. 

This variety is quite distinct from var. heteromorpha 

and the other varieties of the species 

by its much smaller, thickly coriaceous leaves 

with revolute margins and a rounded to retuse 

apex. It is possibly a distinct species, but I hesitate 

to describe it as such on the basis of a single 

collection and in this complex, because there is 

a great deal of morphological variation in var. 

heteromorpha. It also differs from other varieties 

ofL. heteromorpha in the deltoid anthers, which 

also occur in the closely related L. intrapetiolaris. 

The anthers are distinctive because in the dried 

material the pointed apex is much darker in color 

than the broad basal portion. 

2-69.1. Licania laevigata Prance, sp. nov. Type. 

Brazil. Amazonas: Manaus-Caracarai Rd., km 

45, Reserva Biologica do INPA, 4 Apr 1972 

(fl), M. F. Silva & L. Coelho 80 (holotype, INPA 

35442; isotype, NY). Fig. 8.


Species sectione Hymenopus pertinens, a L. 

reticulata fructibus globosis exocarpio tomentellis, 

foliis laevigatis haud reticulatis, floribus 

extus brunneo tomentellis sessilibus differt. 

Tree to 20 m tall, the young branches glabrous. 

Leaf lamina oblong, coriaceous, 9-18 x 4.2-7 

cm, cuneate at base, acuminate at apex, the acumen 

4-10 mm long, glabrous on both surfaces, 

shiny above; midrib prominulous above, prominent 

beneath; primary veins 7-10 pairs, prominulous 

on both surfaces, widely spaced with 

1.2-1.8 mm between veins; petioles 5-8 mm long, 

with confluent leaf base, glabrous, eglandular. 

Stipules axillary, caducous. Inflorescences of terminal 

and subterminal racemose panicles, the 

rachis puberulous. Bracts and bracteoles minute, 

This species and the next (Licania occultans) 

are both close to Licania heteromorpha and I 

have previously placed some of the material 

within that species. The description of these 

species begins the dismemberment of that large 

and morphologically diverse species. The sepa- 

ration of L. laevigata and L. occultans is made 

possible by the collection of large population 

samples from the Reserva Ducke and the Re- 

serves of the INPA/WWF Minimum Critical Reserve 

Size Project near to Manaus. It is now apparent 

that they are different species that are easily 

distinguished in the field. 

Licania laevigata is closest to L. reticulata but 

differs in the smooth, not reticulate leaves that 

are more cuneate at the base, the round tomentose 

fruit rather than an ellipsoid, glabrous and 

costate one, the flowers borne sessile on the primary 

inflorescence branches and the brown tomentellous 

pubescence of the exterior of the 

flowers. 

2-69.2. Licania occultans Prance, sp. nov. Type. 

Brazil. Amazonas: Manaus-Itacoatiara Rd., 

km 31, CEPLAC Research Station, 6 Nov 1973 

(fl), Steward & Ramos P17669 (holotype, 

INPA; isotype, NY). Fig. 9. 

triangular, tomentose, subpersistent. Flowers ca. 

2 mm long, inserted on primary inflorescence 

branches. Receptacle urceolate, brown-tomen- 

A L. heteromorpha foliis 

tellous on 

longe acuminatis, peexterior, 

tomentose within; pedicels tiolis 

0.5-1 mm 

eglandulosis, laminis confluentibus differt. 

long. Calyx lobes five, triangular, to- 


mentellous on both surfaces. Petals five, pubes- with large conspicuous lenticels. Leaf lamina obcent 

on exterior, with ciliate margins. Stamens 

long-elliptic, chartaceous, 4.5-8 x 2.5-4 cm, cu- 

6-7, inserted around three-fourths of circle with 

neate at base, acuminate at apex, the acumen 4- 

tooth-like staminodes opposite; filaments short- 

9 mm 

er 

long, cuspidate, 

than 

glabrous on both surfaces; 

calyx lobes, free, glabrous; anthers delmidrib 

prominulous above, prominent beneath; 

toid. Ovary inserted at base of receptacle, toprimary 

veins 5-7 pairs, prominulous on both 

mentellous on exterior; style pubescent on lower 

surfaces, widely spaced; petioles 2-5 mm long, 

portion. Young fruit globose, exocarp densely with confluent leaf base, glabrous, eglandular. 

short-ferrugineous-tomentellous. 

Stipules 

Distribution (Fig. 62). Terra firme 

axillary, small, caducous. 

forest 

Inflorescences 

of 

of terminal and subterminal racemose panicles, 

Central Amazonia and Surinam. 

the rachis sparsely gray-puberulous. Bracts and 

Additional material. SURINAM. Frederik Top, 3 bracteoles triangular, minute, 0.5 mm long, perkm 

SSE of Juliana Top, 23 Aug 1963 (young fr), Irwin sistent, puberulous. Flowers ca. 1.5 mm long, 

et al. 54990 (NY). 

inserted on 

BRAZIL. AMAZONAS: Sao Gabriel, 27 May 1948 

primary inflorescence branches. Re- 

(y 

fr), Black 48-2902 (IAN, NY); Reserva Florestal Ducke, ceptacle campanulate, sparsely puberulous on 

Manaus, 6 Apr 1967 (fl), Byron & Elias 67-20 (INPA, exterior, tomentellous within; pedicels ca. 0.5 mm 

NY), 27 Mar 1952 (fl), L. Coelho s.n. (INPA 5209, long. Calyx lobes five, triangular, puberulous on 

NY), 29 Sep 1976 (st), Mello s.n. (INPA 60170); 24 both surfaces. Petals five, sparsely pubescent on 

Sep 1976 (st), Adair de Oliveira s.n. (INPA 74791), 19 

Aug 1976 (st), Adair de Oliveira s. n. (INPA 60162), 16 exterior, the margins ciliate. Stamens 5-6, in- 

Jan 1976 (st), Adair de Oliveira s.n. (INPA 59936), 14 serted around three-fourths of circle; filaments 

Sep 1971 (y fr), Prance et al. 14735 (INPA, NY). shorter than calyx lobes, free, glabrous. Ovary 

inserted at base of receptacle, puberulous on exterior; 

style villous on lower half. Fruit unknown. 

Distribution (Fig. 62). Forest on terra firme of 

Central Amazonia. 

Additional specimen examined. BRAZIL. AMAZONAS: 

Maues, 57?43'W, 3?23'S, 23 Jul 1983 (fl), S. R. Hill 

13163 (INPA, NY). 

This species differs from Licania heteromor- 

pha in the long acuminate leaves which are char-


3cm.[ 

.. ' .-. . . . . . 

.- . 

~,a',,,,,? W' .,',,'4g,,i..~';/~ / . 

I ,4< (


taceous and conduplicate, and in the eglandular 

petioles with the leaf base confluent into them. 

2-72. Licania fanshawei Prance, Fl. Neotrop. 

Monogr. 9: 112. 1972. 

2-73.1. Licania marleneae Prance, Brittonia 28: 

218, fig. 7. 1976. Type. Brazil. Amazonas: 

Manaus-Porto Velho Hwy. between Rio Cas- 

tanho and Rio Tupana, 18 Jul 1972 (fl), M. F. 

da Silva 873 (holotype, INPA). 

actually much more widespread, but only infre- 

quently collected. It has been collected in several 

places in Colombia and Venezuela. See Figure 

78. 

Additional specimens examined. COLOMBIA. META: 

Sierra de la 

Distribution (Fig. 38). This species was de- 

Macarena, Central Mountains, north ridge, 

30 Dec 1949 (fr), Philipson & Idrobo 2005 (BM, NY). 

scribed from material collected in Venezuela and VENEZUELA. TACHIRA: Rio San Buena, 10 km W 

Guyana. Recent collections show it to be dis- of La Fundaci6n, 15 Mar 1980 (fr), Liesner et al. 9584 

tributed throughout the Guianas. 

(MO, NY, VEN). ZULIA: Km 45 of Lara-Zulia Rd., 31 

Mar 1979 (fl, fr), Bunting & Fucci 7124 (NY). 

Additional specimens examined. SURINAM. No locality, 

13 Nov 1971 (fr), Jimenez-Sda 1608 (LBB, NY); 

Fallawatra, 22 Nov 1971 (st), Jimenez-Sda 1642 (LBB, Subgenus Licania Section Cymosa 

NY). 

FRENCH GUIANA. Saiil, Monts La Fum6e, 27 2-78.1. Licania arianeae 

Aug 

Prance, sp. nov. Type. 

1982 (fl), Mori & Boom 14780 (NY). 

Brazil. Espirito Santo: Reserva Florestal 

CVRD, Linhares, 19 May 1980 (fl), D. A. Foli 

228 (holotype, RB; isotype, NY). 

Figs. 10, 27. 

Species sectione Cymosae pertinens; a L. cu- 

prea staminibus 6-7, stipulis 6-10 mm longis, 

floribus 5-6 mm longis, a L. riedelii floribus in 

cymulis 2-3-floris dispositis, stipulis floribusque 

maioribus, foliis oblongo-ellipticis differt. 


lenticellate. Leaf lamina, oblong-elliptic, coriaceous, 

5.5-9.5 x 3-4.7 cm, cuneate at base, acute Tree to 20 m tall, the young branches densely 

at apex, glabrous on both surfaces; midrib plane ferrugineous-tomentose. Leaf lamina oblong to 

above, prominulous beneath; primary veins 7- oblong-elliptic, coriaceous, 7-11 x 3.5-5 cm, 

11 pairs, plane or rounded to 

prominulous above, 

subcuneate at 

promin- 

base, bluntly acumiulous 

beneath; petioles 2-3 mm nate or acute at 

long, glabrous, 

apex, the acumen 0-5 mm long, 

rugulose, eglandular. Stipules acute, ca. 0.75 mm glabrous above, with conspicuous deep-set stolong, 

persistent, sparsely puberulous, adnate to matal cavities beneath, filled with a lanate white 

base of petiole. Inflorescences of terminal and pubescence; midrib impressed for entire length 

subterminal branched panicles; rachis sparsely above, prominent and ferrugineous-tomentose 

puberulous, becoming glabrous with age, the beneath; primary veins 10-15 pairs, plane above, 

branches puberulous. Bracts and bracteoles ca. prominent and with a few sparse hairs beneath; 

0.5 mm long, persistent, triangular, puberulous petioles 6-8 mm long, densely ferrugineous-puon 

exterior. Flowers 2-2.5 mm long. Receptacle bescent, canaliculate. Stipules 6-10 mm long, 1.5campanulate, 

sessile, sordid-tomentellous on ex- 

2 mm broad at base, lanceolate, persistent, adterior, 

tomentose within. Calyx lobes nate to extreme base of 

five, acute, 

petiole. Inflorescences of 

tomentellous on exterior, puberulous within. terminal and subterminal panicles, the rachis and 

Petals absent. Stamens five, unilateral, inserted branches densely ferrugineous-tomentose. Bracts 

opposite to three sepals; filaments 

and bracteoles 

glabrous, 

minute, ca. 1 mm long, persistent. 

shorter than calyx lobes. Flowers 5-6 mm 

Ovary pilose. Style gla- 

long, inserted in small groups 

brous. Fruit unknown. 

ofcymules attached to primary branches by short 

Distribution (Fig. 60). Known only from the thick peduncles; receptacle urceolate, slightly 

type, from rain forest on terra firme, collected in gibbous, tomentose on exterior, lanate within. 

flower in July. 

Calyx lobes acute, tomentose on both surfaces. 

Petals absent. Stamens 6-7, unilateral with large 

2-76. Licania silvae Prance, Fl. 

lanate 

Neotrop. Mono- ridge opposite bearing short staminodes, 

gr. 9: 115. 1972. 

filaments shorter than calyx lobes, unequal in 

length. Ovary inserted at base of receptacle, vil- 

This species, described from material from lous; style exceeding filaments in length, lanate 

Brazilian Amazonia in Para and Amazonas, is for entire length. Fruit not seen.


3 cm. 

FIG. 10. Licania arianeae (Foli 228). A, habit; B, leaf undersurface; C, flower; D, flower section; E, stamen; 

F, ovary. 

F, ovary. 

Additional specimen examined. BRAZIL. ESPIRITO 

SANTO: Reserva Florestal de Linhares, 22 May 1972 

(fl), Lino 45 (NY, RB). 

This most distinct species belongs to section 

Cymosa, but has thicker peduncles bearing the 

cymules than in other species of that section. It 

11 

is closest to the Guianan species Licania cuprea, 

but differs in the larger flowers, larger stipules, 

gibbous receptacle, leaf shape, and greater number 

of stamens. It is one of the few species of 

Licania with extremely conspicuous, exposed 

stomatal cavities on the leafundersurface; in this


it superficially resembles L. riedelii, but differs 

in the inflorescence of cymules and the much 

larger stipules. 

2-79. Licania impressa Prance, Fl. Neotrop. 

Monogr.9: 118. 1972. 


Additional specimens examined. BRAZIL. 

AMAZONAS: Manaus-Itacoatiara Rd., km 69-70, 5 Sep 

1973 (fl), Prance et al. 17520 (INPA, NY); Rio Cuieiras, 

2 km below Rio Brancinho, 12 Sep 1973 (fl), Prance 

et al. 17820 (INPA, NY). PARA: Serra do Santar6m, 18 

Aug 1916 (fl), Ducke MG 16355 (MG); Tome Aqiu, 29 

Dec 1977 (fl), Nascimento 340 (MG, NY), 30 Dec 1977 

(fr), Nascimento 363 (MG, NY). 

2-80.1. Licania santosii Prance, Revista Brasil. 

Bot. 2: 28, fig. 1. 1979. Type. Brazil. Bahia: 

Itacare, 15 Oct 1968 (fl), J. Almeida 150 & T. 

S. dos Santos 150 (holotype, CEPEC; isotypes, 

AAU, FHO, MO, NY, U, US). 

Small tree 4 m tall, the young branches glabrous, 

conspicuously lenticellate. Leaf lamina 

ovate-orbicular, coriaceous, becoming thickly 

coriaceous with age, 6-10 x 4-6.5 cm, rounded 

at base, rounded to slightly acute at apex, glabrous 

on upper surface, the lower surface with 

stomatal crypts filled by dense lanate pubescence, 

the venation reticulate and conspicuous 

since it is not covered by the pubescence; midrib 

plane above, prominent beneath; primary veins 

7-8 pairs, plane above, prominent beneath; petioles 

8-10 mm long, terete, sparsely puberulous 

when young, eglandular, slightly rugulose. Stipules 

adnate to extreme base of petioles, persis- 

base of filaments, tomentose on lower half. Fruit 

not seen. 


type collection, from the seaside. 

Licania santosii belongs to section Cymosa on 

account of the cymose inflorescence branches. It 

is closely related to L. dealbata, from which it 

differs in the broader leaves with blunter apices, 

the longer petioles, the pubescence of the lower 

surface not obscuring the venation, and the grayish-brown 

pubescence of the inflorescence. It differs 

from L. hypoleuca, another species of the 

same section occurring in the same region, in the 

stomatal crypts and conspicuously reticulate ve- 

nation, the rounder leaves with blunt apices, and 

the less branched inflorescence with larger flowers 

and more stamens. 

2-81. Licania pallida Spruce ex Sagot, Ann. Sci. 

Nat. Bot., ser. 6, 15: 306. 1883. 

The first Peruvian collection of this Guianan 

and eastern Amazonian species has been col- 

lected (Fig. 69); see also Licania arachnoidea. 

Additional specimen examined. PERU. AMAZONAS: 

Rio Santiago, 65 km N of Pinglo, 4 Dec 1979 (fl), 

Huashikat 1425 (MO, NY). 

2-84. Licania cymosa Fritsch, Ann. K. K. Na- 

turhist. Hofmus. 4: 47. 1889. 

This little known species was studied from three 

highly disjunct collections in 1972. Several more 

from Bahia now have been collected (Fig. 35). 

Additional specimens examined. BRAZIL. BAHIA: 

Mun. Ilheus, Fazenda Barra do Manguinho, 29 Sep 

tent, lanceolate, ca. 2 mm long. Inflorescences of 1980 (fl), Mattos Silva et al. 1136 (CEPEC, NY), 5 Feb 

little-branched panicles 3-4 cm, the rachis and 1982 (fr), Mattos Silva et al. 1401 (CEPEC, NY); Una, 

branches brown-tomentellous. Bracts and brac- 

Oliven9a, 2 Sep 1971 (fr), Pinheiro 1558 (NY). 

teoles 1-2 mm long, ovate, persistent, tomen- Local name. oiti. 

tellous on exterior. Flowers ca. 2.5 mm long, in 

few-flowered cymules attached to rachis and pri- 2-87. Licania piresii Prance, Fl. Neotrop. Monomary 

branches by long slender peduncles. Re- gr. 9: 124. 1972. 

ceptacle campanulate, brown-tomentellous on 

exterior, tomentose within; pedicels 0.2 mm long. 


Calyx lobes five, acute, tomentellous-puberulous 

scribed from three collections from Amapa, Braon 

both surfaces. Petals absent. Stamens seven, 

zil. It has recently been collected in Guyana since 

inserted around complete circle, anthers and fil- my account for the flora of the Guianas was pubaments 

of variable size; filaments shorter than lished (Prance, 1986b). 

calyx lobes, glabrous, free to base. Ovary inserted Additional specimens examined. GUYANA. WEST 

at base of receptacle, villous. Style extending to DEMERARA REGION: Mabura Hill area, 5?20'N, 58?40'W,


3 Jun 1986, Pipoly 7513 (NY, US), 4 Jun 1986, Pipoly 

7524 (NY, US), Pipoly 7537 (NY, US). 

2-87.1. Licania furfuracea Prance, Brittonia 28: 

221, fig. 8. 1976. Type. Venezuela. Bolivar: El 

Dorado-Santa Elena Rd., km 251-253, 4 Jan 

1975 (fl), Steyermark 113900 (holotype, NY; 

isotype, VEN). 

2-90. Licania buxifolia Sandwith, Bull. Misc. In- 

form. 1931: 369. 1931. 

Distribution 

(Fig. 29). This species, only known 

from three collections in Prance (1972), is still 

known only from a restricted area of Guyana. 

Additional specimens examined. GUYANA. Mazaruni-Potaro 

District, Bartica-Potaro Rd., 24 mi S of 

Bartica, 15 Aug 1976 (fl), Mori et al. 8088 (NY); 19 

mi SW of Bartica, 17 Aug 1976 (fl), Mori et al. 8131 

(NY). 

2-94. Licania urceolaris Hooker f., Fl. bras. 14(2): 

15. 1867. 

This species has now been collected over a 

much wider area including Peru (Fig. 84). 


puberulous, becoming glabrous and lenticellate 

with age. Leaf lamina oblong, chartaceous, 3.5- 

5.5 x 1.5-2.5 cm, subcuneate at base, acuminate 

at apex, the acumen 3-7 mm long, glabrous above, 

with a waxy pulverulent-furfuraceous pubescence 

beneath; midrib plane or prominulous 



CAQUETA: Florencia, Rio Orteguaza, 21 Mar 1965 (fr), 

Garcia-Barriga 18202 (COL), Garcia-Barriga 18209 

(COL, US). 

PERU. LORETO: Requena, Rio Ucayali, Arboretum 

Jenaro Herrera, Jul-Sep 1976 (fl), Bernardi 5-36 (G, 

NY); Yanomono, Explorama Tourist Camp, trail to 

Rio Napo, 19 Feb 1981 (fr), Gentry et al. 31502 (MO, 

NY); Maynas, Puerto Almendras, 14 Feb 1977 (fr), 

Revilla 2345 (MO, NY). 

2-95. Licania affinis Fritsch, Ann. K. K. Naturhist. 

Hofmus. 4: 50. 1889. 

pairs, plane above, prominulous beneath; petioles 

2.5-5 mm long, sparsely puberulous, canaliculate. 

Stipules axillary, 1-2 mm long, 

lanceolate, sparsely puberulous, persistent. Inflorescences 

of terminal and axillary compound 

panicles, the flowers borne in small 2-3-flowered 

cymules on short peduncles; rachis and branches 

puberulous. Bracts and bracteoles ca. 5 mm long, 

oblong, persistent, tomentellous on exterior. 

Flowers ca. 2 mm long. Receptacle urceolate, tomentellous 

on exterior, appressed-puberulous on 

interior; pedicels 0.25 mm long. Calyx lobes five, 

acute, tomentellous on exterior, puberulous 

within. Petals absent. Stamens five, unilateral, 

inserted opposite three sepals; filaments glabrous, 

free to base, shorter than calyx lobes. Ovary 

tomentose. Style puberulous. Fruit unknown. 


from the type gathering, collected in flower in 

January in tall forest, beside a small stream. 

Subgenus Licania Section Pulverulenta 

Distribution (Fig. 22). This species was described 

from Guyana. Quite common in the 

Guianas, it was reported only from that region 

and adjacent Brazil in Prance (1972). Later it 

was collected in Panama and added to the increasing 

list of Panama-Guiana disjuncts, as 

pointed out in Prance (1974a). Its Central American 

range has recently been expanded into Costa 

Rica, showing that it exists in suitable forest sites 

well into Central America. 


HEREDIA: Rio Peje, SW sector of La Selva, 24 May 

1982 (fl), Hammel 12476 (NY). 

PANAMA. PANAMA: El Llano-Carti Rd., 10 km N 

of Pan American Hwy., 12 Dec 1973 (fl), Gentry 8884 

(MO, NY). COLON: Santa Rita, Feb 1968 (fl), G6mez- 

Pompa et al. 3347 (MEXU), Gomez-Pompa et al. 3354 

(MEXU), 10 Aug 1971 (fr), Lao et al. 10 (MO, NY). 

2-95.1. Licania teixeirae Prance, sp. nov. Type. 

Brazil. Rondonia: Minera~ao Santa Barbara, 

25 May 1982 (fl), L. O. A. Teixeira 728 (holotype, 

INPA 104465; isotypes, FHO, NY). 

Fig. 11. 

Species sectione Pulverulentae pertinens, L. 

urceolaria affinis, foliis chartaceis 4-5.5 mm longis, 

apice caudato, receptaculo urceolato, extus 

brunneo-tomentoso differt. 

Tree, the young branches glabrous, lenticellate. 

Leaf lamina narrowly ovate, chartaceous, 4-5.5 

x 1.7-2.3 cm, subcuneate and slightly unequal 

at base, caudate at apex, the acumen 1.2-1.8 mm 

long, glabrous above, with a waxy pulverulentfarinaceous 

pubescence beneath; midrib gla-


D. 

?..-n . .. 2mm t r 

FIG. 11. Licania teixeirae (Teixeira 728). A, habit; B, flower; C, flower section; D, apical view of flower; 

E, leaf undersurface 

showing pulverulent pubescence. 

brous and plane above, prominent beneath; primary 

veins 7-9 pairs, inconspicuous on both 

surfaces; petioles 4-6 mm long, glabrous, canaliculate. 

Stipules linear, 1 mm long, adnate to 

base of petiole. Inflorescences of terminal and 

axillary little-branched racemose panicles or racemes, 

the rachis and branches tomentellous, the 

apical panicles 3-4 cm long, the axillary racemes 

1-2 cm long. Bracts and bracteoles lanceolate, 

persistent, ca. 0.5 mm long, tomentellous. Flowers 

ca. 3 mm long, sessile on primary branches 

of inflorescence. Receptacle urceolate, sessile, rufous-brown-tomentellous 

on exterior, densely 

tomentellous within. Calyx lobes five, acute, rufous-brown-tomentellous 

on exterior contrasting 

sharply with gray-white-tomentellous pubescence 

of inner surface. Petals absent. Stamens 

five, inserted around 2/3 of circle, the filaments 

shorter than calyx lobes, tomentellous. Ovary 

inserted at base of receptacle, sparsely tomen- 

tellous on exterior. Style equalling filaments, la- 

nate-pubescent. Fruit unknown. 


type collection. 

This species is quite distinct from other species 

of Licania and is easily recognized in the dry 

condition by the contrasting colors of the recep- 

tacle and interior of the calyx lobes. It is a mem- 

ber of section Pulverulenta and differs from all 

other species in the leaf shape and the thin char- 

taceous leaves. This species comes from the same 

locality as another recently described species of 

Licania, L. bellingtonii. 

Subgenus Licania Section Licania 

2-100. Licania couepiifolia Prance, Fl. Neotrop. 

Monogr. 9: 134. 1972.



from a single collection from Guyana. A 

second collection from Surinam extends its range. 

Additional specimen examined. SURINAM. Sipaliwini 

area (fr), Oldenburger et al. 1256 (LBB). 

2-104. Licania hebantha Martius ex Hooker f., 

Fl. bras. 14(2): 17. 1867. 



AMAZONAS: Monochoa, Sabana de Mosco, 30 Dec 1976 

(fl), Sastre & Reaichel D. 5060 (P); Rio Caqueta, Araracuara, 

7 Jan 1977 (fl), Sastre & Reaichel D. 5178 (NY, 

P). 

2-105. Licania steyermarkii Maguire, Fieldiana, 

Bot. 28: 254. 1952. 


Additional specimens examined. VENEZUELA. 

AMAZONAS: Dept. Rio Negro, lower Rio Pacimoni, 8 

Feb 1981 (fl), Huber & Medina 5847 (NY, VEN). 

BOLiVAR: Between km 251.5 and 253 road El Dorado, 

Santa Elena, 4 Jan 1975 (st), Steyermark 111391 (NY); 

4.5 km SW of Icabaru, Quebrada Los Brasileros, 16 

Dec 1978 (fl), Steyermark et al. 117776 (NY). 

2-107. Licania crassivenia Spruce ex Hooker f., 

Fl. bras. 14(2): 14. 1867; Prance, Fl. Neotrop. 

Monogr. 9: 137. 1972. Fig. 32. 

Until recently this species was known only from 

the Spruce type. The material cited below keys 

to L. crassivenia and is included there for the 

present, but has much larger leaves than the type 

(10-13 x 4-5.5 cm as Distr., Falawatra, Nov 1971 (fl), Jimenez-Sda 

1549 (holotype, NY; isotype, LBB 14282). 

Tree to 26 m tall, the young branches tomentellous 

soon becoming glabrous. Leaf lamina oblong-elliptic, 

chartaceous, 14-19 x 6-7.5 cm; base 

subcuneate, apex acuminate, the acumen 10-18 

mm long, glabrous above, densely lanate beneath, 

deeply reticulate venation beneath; midrib 

impressed above, tomentellous towards base, 

prominent beneath; secondary veins 9-11 pairs, 

plane above, prominent beneath; petioles 15-20 

mm long, weakly canaliculate, shortly tomentellous, 

with two to many glands towards base. 

Stipules not seen. Inflorescences of terminal and 

axillary panicles, ca. 2.5 cm long; rachis and 

branches tomentellous. Bracts and bracteoles 

ovate-lanceolate, 0.6 mm long, persistent, pubescent. 

Flowers 1.5 mm long. Receptacle campanulate, 

tomentellous on exterior, tomentose 

within; pedicels 0.4 mm long. Calyx lobes five, 

tomentose. Petals absent. Stamens seven, unilateral, 

included, three large and fertile, four 

smaller and sterile; filaments glabrous, free to 

base. Ovary lanate-pubescent. Style lanate. Fruits 

pyriform, to 5 cm long including stipe of 1.5-2 

cm; exocarp velutinous, ferrugineous-pubescent; 

mesocarp thin; endocarp 1 mm thick, hard and 

fibrous, sparsely hirsute within. 


rain forests of Guyana and Surinam. 

Additional specimens examined. GUYANA. Murudi 

Mts., Mazoa Hill, 2?15'N, 59?10'W, 12 Nov 1982 

(fl), Stoffers et al. 30143 (NY, U). 

SURINAM. Sipaliwini Savanna area, forest W of 

compared to 5-8 x 1.5- Meyers farm, Jan 1970 (fr), Oldenburger et al. 1213 

3.8) and a yellow-brown-tomentellous inflores- (NY, U), Oldenburger et al. 1225 (NY, U). 

cence rather than gray. The leaves of the Her- Local name. rode kwepi. 

ndndez & Dezzeo material are exactly the same This species is most distinct and not easily 

shape as those of the type, gradually tapering confused with any other species. The numerous 

towards the apex from near to the base; they also axillary inflorescences, and the two types of stahave 

the extremely conspicuous stomatal cavi- mens distinguish this species. The larger stamens 

ties that are made obvious by the surrounding are fertile, the smaller ones have well-developed 

glabrous venation. This could be two different anthers but abortive pollen. Licaniajimenezii is 

species but I hesitate to describe another one probably closest to L. alba and L. robusta. It 

until further material is collected to show the differs from L. alba in the much smaller bracmorphological 

variation. 

teoles and flowers, the less conspicuous leaf re- 

Additional specimen examined. VENEZUELA. ticulation, the longer petioles, and the stamens, 

BOLiVAR: Sorochoroyen, Sifontes Dept., 5?13'N, and from L. robusta in the smaller flowers, the 

611 1'W, 20 Apr 1985 (fl), Herndndez & Dezzeo 128 pubescent inflorescence, the glandular petioles, 

(MYF, NY). 

the caducous stipules, and the impressed midrib. 

2-108.1. Licania jimenezii Prance, Acta Ama- 

zonica 2(1): 7. 1972. Type. Surinam. Nickerie 

2-117. Licania ovalifolia Kleinhoonte, Recueil 

Trav. Bot. Neerl. 30: 180. 1933. Fig. 69.


Additional specimens examined. SURINAM. Fal- 

lawatra, 7 Jan 1972 (st), Jimenez-Sda 1688 (LBB, NY). 

BRAZIL. AMAPA: Contagem, Oct-Dec 1976 (st), Rosa 

1061 (MG). 

2-119.1. Licania stewardii Prance, Brittonia 28: 


Manaus-Caracarai Rd., km 130, 13 Feb 1974 

(fl), Steward et al. P20251 (holotype, INPA; 

isotypes, FHO, NY, US). 

Small tree 3-5 m tall, the young branches puberulous, 

soon glabrate. Leaf lamina oblong, 

chartaceous, 4-8 x 2.2-4 cm, rounded or subcordate 

at base, rounded, acute or retuse at apex, 

glabrous above with a waxy lanate appressed pubescence 

beneath; midrib slightly impressed 


pairs, plane above, prominent beneath, the secondary 

venation conspicuously reticulate; petioles 

1.5-2.5 mm long, terete, eglandular, tomen- 

tellous when young. Stipules linear, 2-3 mm long, 

persistent, tomentellous, adnate to petioles. Inflorescences 

of terminal or subterminal little 

branched racemose panicles or simple racemes 

2-7 cm long; rachis and branches gray-tomentellous. 

Bracts and bracteoles narrowly lanceolate, 

1-2 mm long, tomentellous on exterior, persistent. 

Flowers 3-3.5 mm long. Receptacle 

campanulate, gray-tomentellous on exterior, lanate 

within; pedicels 0.25-0.5 mm long. Calyx 

lobes five, acute, gray-tomentellous on exterior, 

puberulous within. Petals absent. Stamens 9-11, 

inserted around complete circle, shorter than calyx 

lobes; filaments connate at base, the united 

part lanate. Ovary densely lanate. Style equalling 

filaments. Fruit unknown. 


only from the vicinity of the type locality where 

it was very common in low campina forest around 

a sandstone rock outcrop in an area where the 

original forest has now been destroyed. 


AMAZONAS: Manaus-Caracarai Rd., km 130, 16 Dec 

1975 (fl), D. Coelho et al. 708 (INPA 54246, NY), 25 

Aug 1976 (fl), Davis & D. Coelho 60303 (UEC), 13 Feb 

1974 (fl), A. Loureiro et al. s.n. (INPA 47911, NY), 15 

Feb 1974 (fl), INPA 47955 (INPA), 6 Jan 1976 (fl), 

Monteiro & Ramos 33 (INPA 54344, NY), 25 May 

1974 Rodrigues et al. 9284 (INPA), 22 Mar 1978 (y 

fr), N. T. Silva 4578 (MG, NY); km 350, 18 Nov 1977 

(fl), Steward 88 (INPA, NY), 14 Feb 1974 (fl), Steward 

P20326 (FHO, INPA, NY), 16 Feb 1974 (fl), Steward 

P20401 (FHO, INPA, NY), 4 Oct 1977 (fl), Ramos & 

D. Coelho 755 (INPA). 

2-121.1. Licania tocantina Prance, Acta Ama- 

zonica 13: 28. 1983. Type. Brazil. Para: Rio 

Tocantins, Tucurui, Breu Branco, 11 May 1978 

(fl), M. G. Silva & R. Bahia 3508 (holotype, 

MG; isotypes, INPA, NY). Fig. 82. 

Tree 20 m tall, the young branches sparsely 

puberulous, soon glabrate. Leaf lamina elliptic, 

chartaceous, 8-15 x 3.5-7.2 cm, subcuneate at 

base, acuminate at apex, the acumen 7-12 mm 

long, glabrous above, with stomatal cavities filled 

by dense lanate pubescence beneath; midrib plane 

or slightly impressed above; petioles 4-6 mm 

long, with two large conspicuous glands when 

young, canaliculate, tomentellous when young, 

becoming glabrous and rugulose with age. Stip- 

ules triangular, ca. 5 mm long, ca. 2.5 mm broad 

at base, inserted on base of petioles, persistent. 

Inflorescences of terminal and subterminal racemose 

panicles, the rachis and branches puberulous. 

Bracts and bracteoles 1-2 mm long, 

triangular, persistent, puberulous on both surfaces. 

Flowers ca. 1.5 mm long, sessile on primary 

branches of inflorescence. Receptacle campanulate, 

sessile, tomentose on exterior and 

within. Calyx lobes five, acute, puberulous on 

both surfaces. Petals absent. Stamens three, unilateral; 

filaments shorter than calyx lobes, glabrous. 

Ovary inserted at base of receptacle, pilose. 

Style equalling filaments, hirsute threefourths 

of length. Fruit pyriform ca. 2 cm long 

(5 cm stipe); exocarp ferrugineous-pubescent; 

mesocarp thin; endocarp hard, granular, 1 mm 

thick. 

Additional specimens examined. BRAZIL. PARA: Rio 

Tocantins, Tucurui, Igarap6 Cagancho, 2 Feb 1980 (fl), 

Lisboa et al. 1382 (MG, NY), 21 Aug 1980 (fr), Rodrigues 

et al. 10259 (INPA). 

This species is most closely related to Licania 

triandra, from which it differs in the thinner, 

chartaceous leaves with deep stomatal cavities 

covered by a lanate pubescence, which also ex- 

tends over the venation, making the cavities much 

less conspicuous than in L. triandra. It also dif- 

fers in the broader stipules adnate to the petiole 

and in the petiole rather than leaf bases glan- 

dular.


2-124. Licania micrantha Miquel, Stirp. suri- 

nam. select. 20. 1850. 

Distribution (Fig. 63). This species is wide- 

spread in Venezuela, Amazonia, and the Guia- 

nas. Its known range has now been extended west 

of the Andes from recent collections from Choc6 

and Valle in Colombia, and also to Atlantic 

coastal Brazil in the forests of Bahia. 

2-126.1. Licania aracaensis Prance, Brittonia 28: 


Serra Araca, 1?N, 63?W, 10 Feb 1975 (fl), Pires 

15027 (holotype, IAN; isotypes, INPA, NY). 

Tree 2 m tall, the young branches puberulous, 

soon becoming glabrous and lenticellate. Leaf 

lamina oblong, conduplicate, coriaceous, 3-5.5 

x 2-2.8 cm, subcuneate at base, acute to bluntly 

acuminate at apex, the acumen 2-6 mm long, 

glabrous above, densely lanate-tomentose beneath; 

midrib plane above, prominulous beneath; 

primary veins 6-8 pairs, plane above, 

prominulous beneath; petioles 3-4 mm long, terete, 

sparsely puberulous, eglandular. Stipules 

persistent, lanceolate, 1-1.5 mm long, adnate to 

base of petiole. Inflorescences terminal and subterminal 

racemose panicles, 4-7.5 cm long; rachis 

and branches puberulous. Bracts and bracteoles 

triangular, persistent, tomentose on 

exterior, ca. 0.75 mm long. Flowers ca. 2 mm 

long. Receptacle urceolate, sessile, yellow-tomentellous 

on exterior, tomentose towards base 

and glabrous above on interior. Calyx lobes five, 

acute, tomentellous on exterior, puberulous 

within. Petals absent. Stamens three, unilateral, 

opposite three calyx lobes; filaments glabrous, 

shorter than calyx lobes. Ovary tomentose. Style 

puberulous, shorter than calyx lobes. Fruit pyr- 

iform; exocarp glabrous, wrinkled when dry, the 

stipe 5-10 mm in dry fruit. 

Distribution (Fig. 25). This species is appar- 

ently endemic to the summit of the sandstone 

mountain Serra Araca at about 1000 m, flow- 

ering in February. 


AMAZONAS: Serra Araca, 1 Feb 1978 (fr), Rosa & Lira 

2337 (IAN, NY), 22 Feb 1984 (fr), Tavares et al. 114 

(INPA, NY). 


CHOCO: Rio Fujiad6, afluente del Rio San Juan, 4?36'N, 2-128. Licania bracteata 

76?54'W, 7 Apr 1979 (fl), Forero et al. 4788 Prance, Fl. Neotrop. 

(COL, 

MO). VALLE: Mun. Buenaventura, Concesi6n Cart6n Monogr. 9:155. 1972. 

de Colombia, 15 Nov 1979 (fl), van Rooden et al. 358 

Distribution 

(NY, U), 6 Dec 1979 (fl), van Rooden et al. 553 (Fig. 29). This species was de- 

(NY, 

U). 

scribed from considerable material from the vi- 

BRAZIL. BAHIA: Mun. Una, Maruim, 28 Apr 1981 cinity of Manaus, Brazil, where it is quite com- 

(st), Mori et al. 13774 (CEPEC, NY), 1 May 1981 (st), mon in the forests on terra firme. Recently it has 

Mori et al. 13884 (NY), 13 May 1981 (st), Mori et al. also been collected in Peru. 

14000 (NY); Esta;ao Experimental Lemos Maia, 24 

Oct 1980 (fr), Rylands 42/80 (NY), 11 Nov 1980 (st), Additional specimens examined. PERU. LORETO: Rio 

Rylands 56/80 (NY). 

Nanay, Santa Maria, 30 May 1963 (fl, fr), Ar6stegui V. 

105 (US); Rio Nanay, 28 Oct 1965 (fl), A. G. Ruiz 224 

(NY, US). 

Local name. Peru: parinari. 

2-130.1. Licania lamentanda Prance, sp. nov. 

Type. Brazil. Bahia: Municipality of Ilheus, 4 

km N of Oliven9a on road to Ilheus, 19 Apr 

1981 (fl), S. A. Mori et al. 13673 (holotype, 

CEPEC; isotype, NY). Fig. 12. 

Species sectione Licania pertinens. Folia 12- 

16.5 cm longa, petiolo 1.1-1.5 cm longo, velu- 

tino-tomentello. Flores 4.5-5.5 mm longi, sta- 

minibus 6-7, ovario densissime tomentoso. 

Tree, the young branches shortly tomentellous, 

becoming glabrous with age. Leaf lamina ob- 

long-elliptic, coriaceous, 12-16.5 x 6-8.5 cm, 

subcuneate to rounded at base, rounded to apic- 

ulate at apex, glabrous above, with a short ap- 

pressed-lanate brown pubescence beneath; mid- 

rib prominulous above, prominent beneath; 

primary veins 10-12 pairs, prominulous above, 

prominent beneath, secondary venation promi- 

nulous, ca. more or less parallel at 90? to primary 

veins; petioles 1.1-1.5 cm long, velutinous-to- 

mentellous, terete or weakly canaliculate above, 

eglandular. Stipules caducous (not seen). Inflo- 

rescences of racemose panicles, 10-14 cm long, 

the rachis and branches shortly tomentellous. 

Bracts and bracteoles membranous, persistent,


F.in(t3A, ;Clwr,foeeo. ai;B efudr 

3 cnr.l \I ~~~s~~llj Illit 'CV 

I 111 

I , . . . . . . . . 

FIG. 12. Licania lamentanda (Mori et al. 13673). A, habit; B, leaf undersurface; C, flower; D, flower section.


ovate, 1.5-3 mm long, tomentellous on exterior, Distribution (Fig. 56). This species, common 

puberulous within. Flowers 4.5-5.5 mm long. along river margins in the Guianas and northern 

Receptacle campanulate, 3.5 mm long, tomen- Para, has recently been found to be disjunct in 

tellous on exterior, tomentose within, subsessile. Atlantic Coastal Brazil, where it also grows on 

Calyx lobes five, acute, triangular, ca. 1.5 mm river margins. This adds yet another Amazolong, 

tomentellous on both surfaces. Petals ab- nian-Bahian disjunct, similar in distribution to 

sent. Stamens 6-7, shorter than calyx lobes, in- Licania cymosa which, however, is rarer in Amaserted 

around three-fourths of a circle. Ovary zonia. 

extremely densely tomentose, thick-walled. Style Additional 

pubescent on lower portion, glabrous above. Fruit 

specimen examined. BRAZIL. BAHIA: Rio 

Sto. Antonio, Mun. Andarai, 19 Jun 1984 (fl), Hatschellipsoid, 

5-6.5 mm long x 4-4.5 cm broad; bach & Kummrow 48068 (MBM, NY). 

exocarp densely velutinous tomentellous; pericarp 

hard, woody, ca. 6 mm thick, lanate within. 

Distribution (Fig. 53). Restinga on 2-135.1. Licania nelsonii 

sandy soil, 

Prance, sp. nov. Type. 

25 m altitude, growing with many individuals of Brazil. Amazonas: 3 km south of Serra Araca, 

piaCaba palm in Bahia. 

0?49'N, 63?19'W, 29 Feb 1984 (fl), W. A. Rodrigues 

et al. 10501 (holotype, INPA; isotype, 

Additional specimens examined. BRAZIL. BAHIA: NY). Figs. 13, 62. 

Mun. Ilheus, Fazenda Barra do Manquinho, km 10 

road Pontal-Olivenca, 5 Feb 1982 (fl, fr), L. A. Mattos Species a sectio Licania pertinens, a L. incana 

Silva et al. 1440 (CEPEC, NY); Mun. Uruguca, 28 km et L. leptostachya foliis subtus conspicae reti- 

NE of Urucuca on road to Serra Grande, 2 Dec 1979 

(fr), Mori et al. 13063 culatis, stipulis a base peciolarum adnatis, re- 

(CEPEC, NY). 

ceptaculo rufo-pubescente, floris dense fascicu- 

This most distinct new species is not easily latis, arboribus grandis differt. 

confused with others. It differs from related Tree to 30 m tall, the young branches brownspecies 

in section Licania in the large flowers and tomentose. Leaf lamina oblong-elliptic, chartaleaves. 

The thick-walled ovary with a dense al- ceous to coriaceous, 3-6.5 x 1.8-3.2 cm, roundmost 

wall-like covering of hair and the seven ed to slightly subcordate at base, acuminate at 

stamens also distinguish it from most species of apex, the acumen 1-3 mm long, densely whitethe 

section. The long petioles differentiate it from lanate pubescent beneath; midrib slightly imall 

except L. robusta, which has glabrous inflo- pressed above, prominent beneath; primary veins 

rescence branches and many other differences. 5-7 pairs, plane to lightly impressed above, 

The specific epithet, meaning "fit to be prominent beneath; venation conspicuously remourned," 

is derived from the pitiful state of the ticulate beneath in mature leaves; petioles 1-2 

forests of eastern Brazil where most of the ex- mm long, terete, eglandular, tomentose when 

citing novelties like this species are lamentably young. Stipules adnate to base of petioles, linear, 

on the verge of extinction. 

persistent, to 3 mm long. Inflorescences of terminal 

and axillary densely crowded spikes, 1- 

2-131. Licania lanceolata Prance, Fl. Neotrop. 4.5 cm long, the rachis densely tomentose. Bracts 

Monogr. 9:158. 1972. 

and bracteoles ovate, persistent, rufous-pubes- 

Distribution (Fig. 53). This species, common cent, ca. 1 mm long. Flowers ca. 2 mm long, 

in the Venezuelan Amazonian savannas, is acsessile 

and densely clustered along inflorescence 

tually more widespread. It has now been colrachis. 

Receptacle campanulate, sessile, rufous 

lected in the savannas of Roraima, Brazil, an 

tomentose on exterior, tomentose within. Calyx 

interesting distribution that is seen for a number 

lobes five, acute, cream when fresh, rufous-toof 

different species, for example Barcella odora 

mentose on exterior when dry, pale brown-to- 

(Trail) Drude 

mentose on interior. Petals absent. Stamens five, 

(Arecaceae). 

inserted opposite three calyx lobes; filaments 

Additional specimen examined. BRAZIL. RORAIMA: shorter than calyx lobes, free, glabrous. Ovary 

Manaus-Caracarai Rd., km 530, in basin of Rio Ana- inserted at base of receptacle, densely pilose. Fruit 

ua, 12 Feb 1979 (fl), Rodrigues et al. 10129 (INPA). unknown. 

Habitat. Igap6 forest in sandy soil. 

2-134. Licania leptostachya Bentham, J. Bot. 

(Hooker) 2: 220. 1840. 

Additional specimen examined. BRAZIL. AMAZONAS:


~i c, B. n 

FIG. 13. Licania nelsonii (Rodrigues 10501). A, habit; B, leaf undersurface; C, inflorescence; D, flower and 

bracteole; E, flower section; F, ovary; G, floral diagram. 

3 km south of Serra Araca, 0?49'N, 63?19'W, 16 Mar 

1984 (fl), Miralha 67 (INPA, NY). 

This species belongs to section Licania and is 

most closely related to L. incana and L. leptostachya. 

It is a much larger tree than either of 

those species and differs in the densely crowded 

inflorescences with a rufous pubescence when dry, 

in the stipules which are adnate to the petioles 

rather than axillary, in the leaves which are 

broadest quite near to their base, and in the conspicuously 

reticulate leaf undersurface. 

This species is named for Bruce Walker Nel- 

son, whose work in Brazil has made possible the 

Projeto Flora Amazonica series of expeditions, 

one of which collected the material of this new 

species. 

2-136. Licania paraensis Prance, Fl. Neotrop. 

Monogr. 9: 163. 1972. 

This poorly known species was described from 

material collected near Santarem, Brazil (Ducke


RB 18818) and in Pando, Bolivia (Prance et al. 

6532) (Fig. 70). One collection from Manaus, 

originally assigned to this species, belongs to Li- 

cania impressa Prance. Rodrigues & Loureiro 

7236 (INPA 15798) is actually the latter species. 

Additional specimens examined. BRAZIL. PARA: Rio 

Jari, Monte Dourado, 30 Nov 1978 (fl), Santos 444 

(MG, NY); Estrada Caracaru to Munguba, 17 Oct 1969 

(fl), N. T. Silva 2826 (NY). 

2-137. Licania vaupesiana Killip & Cuatrecasas, 

Fieldiana, Bot. 27: 105. 1951. 

2-144.1. Licania harlingii Prance, Fl. Ecuador 

10: 9-10, fig. 2. 1979. Type. Ecuador. Napo: 

Ca. 6 km S of Puerto Napo, 8 Apr 1969 (fl), 

H. Lugo S. 1054 (holotype, GB; isotype, NY). 

Tree to 26 m tall, the young branches puberulous, 

soon becoming glabrous and lenticellate 

with age. Leaf lamina ovate-elliptic, chartaceous, 

7-10 x 3-5 cm, subcuneate at base, acuminate 

at apex, the acumen 8-12 mm long, glabrous 

above, densely lanate-pubescent below with 

puberulous conspicuous venation; midrib slightly 

impressed above, prominent beneath; secondary 

veins 8-11 pairs, plane above, impressed be- 

plorama Camp, Rio Amazonas, 14 Jul 1983 (st), Gentry 

et al. 43029 (MO, NY). SAN MARTIN: Distrito Tocache 

Nuevo, Quebrada de Cachiyacu, 500-600 m, 9 

May 1975 (fl), Schunke V. 8439 (MO, NY). 

This species is close to Licania blackii of east- 

ern Amazonia, it differs in the small axillary, 

caducous stipules, the greater number of primary 

veins, and the shorter, darker pubescence of the 

flowers. It is also close to the Venezuelan L. crue- 

geriana, but differs in the larger leaves with a 

more acuminate apex, the smaller stipules, and 

the laxer, less branched inflorescence. 

Additional specimen examined. BRAZIL. AMAZONAS: 

Mun. Serrinha, igarap6 de Serrinha, 12 Nov 1977 (fl), 2-145. Licania cruegeriana Urban, Symb. An- 

Damido 2637 (INPA). 

till. 5: 352. 1908. 

Distribution (Fig. 34). Common in Trinidad 

and northern Venezuela, this species is also dis- 

junct in Panama. 

Additional specimens examined. PANAMA. COLON: 

23 Feb 1968 (fr), Duke 15256 (MO, NY). PANAMA: 5- 

6 mi N of El Llano, 8 Sep 1972 (fl), Gentry 5800 (MO, 

NY). 

2-146. Licania belemii Prance, Fl. Neotrop. 

Monogr. 9: 172. 1972. 


from a single specimen from Belmonte 

in Bahia. It appears still to be quite common in 

neath; petioles 5-6 mm long, tomentellous, terete. the forests of eastern Brazil since several new 

Stipules small, ca. 1 mm, axillary, caducous. In- collections have been made there. 

florescences of terminal racemose panicles, the Additional specimens examined. BRAZIL. BAHIA: 

rachis and branches yellow-brown, puberulous. Between Uruguca and Serra Grande, 16 Jul 1978 (y 

Bracts and bracteoles small, triangular-hastate, fr), Mori et al. 10252 (CEPEC, NY); Niicleo Colonial 

persistent, tomentellous on exterior. Flowers ca. de Una, between BR 101 Sao Jose and BA 265, 29 Oct 

1978 (fr), Mori & Thompson 11033 (CEPEC, NY); 

1.5 mm long, borne solitary along primary Una, Fazenda Sao Rafael, 16 Dec 1968 (fl), Santos 320 

branches of inflorescence. Receptacle globose, (NY); Camaca, estrada Rio Branco, 28 Jan 1971 (fl), 

sessile, brown-tomentellous on exterior, lanate Santos 1444 (NY). ESPIRITO SANTO: Linares, Reserva 

within. Calyx lobes acute, tomentellous on both CVRD, 11 Jan 1979 (fl), Foli 61/79 (INPA); Rio Doce, 

surfaces. Petals absent. Stamens 5-6, slightly uni- Lag6a do Durao, 14 May 1934 (fr), Kuhlmann 208 

(RB 35360). 

lateral; filaments free, shorter than calyx lobes, 

glabrous. Ovary inserted at base of receptacle, Local names. oiti, milho-torrado-amarelo. 

tomentellous. Style equalling filaments, glabrous 

or with a few hairs only. Fruit not seen. 2-148. Licania veneralensis Cuatrecasas, Field- 

Distribution (Fig. 43). Western Amazonia, in iana, Bot. 27: 109. 1951; Prance, Brittonia 28: 

Ecuador and Peru. 

212-215. 1976. 

Additional specimens examined. ECUADOR. NAPO: This species, poorly known at the time of the 

Apuya, ca. 6 km S of Puerto Napo, 6 Apr 1969 (fl), monograph, has been described in detail (Prance, 

Lugo S. 1039 (GB, NY). PERU. MADRE DE DIOS: Prov. 

Manu, Cerro de Pantiacolla, Rio Palatoa, 13 Dec 1985 

1976), and several additional collections have 

(fl), Foster 10996 (F, NY); Tambopata, Rio Tambonow 

been studied. It has now been shown to 

pata, 12?49'S, 18 Feb 1984 (fl), Gentry et al. 45582 belong to the L. durifolia complex, see p. 20. 

(MO, NY). LORETO: Maynas Prov. Yanamono Ex- Distribution. Figure 85.

56 

2-149. Licania amapaensis Prance, Fl. Neotrop. 

Monogr. 9: 174. 1972. Fig. 23. 

Inflorescence paniculate, the flowers borne in 

small cymules on short secondary branches. 

Young fruit ovoid, densely ferrugineous-veluti- 

nous-tomentose, longitudinally striate when dry; 


Additional specimens examined. COLOMBIA. mesocarp thin and fleshy; endocarp hard, 1.5 

CHOCO: Cabo Corrientes, Rio Parguera, 27 May 1974 mm thick in young fruit, glabrous within. 

(st), Warner 299 (COL). NARI/O: Tumaco, vic. of Sa- This 

lahonda, 5 Jul 1955 (fr), Romero C. 5270 (COL). vALLE: 

very distinct species with large stipules 

Bajo Calima, 9 May 1961 (fl bud), Cabrera R. 553 was described from a single specimen with young 

(COL), 18 Jul 1961 (fl), Cabrera R. 600 (COL). flower buds only. A recent collection, while also 

only in bud, does have fruit and also adds to 

inflorescence characters. 

Additional specimens examined. FRENCH 

GUIANA. Sail, Monts La Fum6e, 4 Nov 1982 (st), 

Mori & Boom 15161 (NY). 

BRAZIL. TERR. AMAPA: Agua Fria, 23 Oct 1979 (fl, 

fr), Austin et al. 7180 (MG, NY). PARA: Belem, 2 Mar 

1955 (st), T. N. Guedes 311 (IAN). 

3. Parinari Aublet 

Key to the American Species of Parinari 

1. Stipules (5-)10-40 mm long, persistent, semiamplexicaul. 

2. Leaf base subcordate to cordate, the apex blunt; inflorescence densely crowded. 3.1. P. alvimii. 

2. Leaf base rounded to acute, the apex acuminate, inflorescence laxer. 

3. Young branches with short soft pubescence; leaf base rounded to cordate. 1. P. campestris. 

3. Young branches with dense stiff ferrugineous-brown hairs; leaf base cuneate to subcuneate. 

3. P. rodolphii. 

1. Stipules small, 1-4 mm long, caducous, axillary and not clasping stem. 

4. Leaf undersurface white-lanate, the pubescence obscuring the stomatal cavities; subshrub or small 

shrub. 16. P. obtusifolia. 

4. Leaf undersurface with gray-brown pubescence not entirely obscuring the stomatal cavities; large 

shrubs or trees. 

5. Midrib distinctly impressed on leaf upper surface for entire length. 

6. Leaf apex rounded to apiculate or bluntly acuminate on some leaves, with acumen to 3 mm 

long; primary leaf veins 9-20 pairs. 

7. Leaves orbicular to broadly elliptic, 2.5-6 cm long, 1.5-4.5 cm broad, the primary veins 

9-14 pairs. 

10. P. maguirei. 

7. Leaves elliptic, 6.5-7.5 cm long, 3-4 cm broad, the primary veins 15-20 pairs. 

11. P. littoralis. 

6. Leaf apex with a well-developed acumen 7-15 mm long; primary leaf veins 21-30 pairs. 

8. Leaves 9-17 cm long; petioles usually with two pairs of glands; flowers 6-9 mm long; 

primary leaf veins more than 3 mm apart. 

2. P. montana. 

8. Leaves 2-7.5 cm long; petioles without distinct glands; flowers 5-6 mm long; primary leaf 

veins 1-2 mm apart. 

12. P. parvifolia. 

5. Midrib impressed only on lower portion of upper surface, or not impressed. 

9. Leaf base distinctly cordate or subcordate. 

10. Leaves ovate, the acumen 10-12 mm long; petioles 6-12 mm long, glandular; inflorescences 

much-branched, 5-6 cm long. 

13. P. cardiophylla. 

10. Leaves oblong, the acumen 1-3 mm long; petioles 3-4 mm long, eglandular; inflorescences 

little-branched, 2-4 cm long. 

17. P. romeroi. 

9. Leaf base rounded to cuneate. 

11. Primary leaf veins 28-35. 14. P. parilis. 

11. Primary leaf veins 12-26. 

12. Inflorescence a dense many-flowered corymbose panicle. 

13. Inflorescence and flowers with yellow-brown pubescence; leaf apex acute or 

short-acuminate, the acumen 0-4 mm long; young branches with short pubescence 

only. 5. P. occidentalis. 

13. Inflorescence and flowers with silver-gray pubescence; leaf apex with acumen 

8-13 mm long; young branches with both short pubescence and long stiff hairs. 

9. P. klugii. 

12. Inflorescence a lax panicle, not corymbose. 

14. Leaf ovate, with a well-developed slender acumen 9-16 mm long. 6. P. sprucei. 

14. Leaf elliptic to oblong-lanate, the acumen 0-12 mm long.



of Species of Parinari 

15. Primary veins 22-26 pairs; leaves 7.5-16 cm long. 

16. Upper surface of midrib slightly prominent; petioles terete. 

8. P. brasiliensis. 

16. Upper surface of midrib slightly impressed on lower portion; petioles 

canaliculate. 15. P. chocoensis. 

15. Primary veins 16-21 pairs; leaves 3-9.5 cm long. 

17. Young petioles canaliculate; leaf reticulation prominent on upper surface. 

7. P. pachyphylla. 

17. Young petioles terete; leaf reticulation inconspicuous on upper surface. 

4. P. excelsa. 

The distribution of Parinari as a whole is shown 

in Figure 86. 

3-3.1. Parinari alvimii Prance, Revista Brasil. 

Bot. 2: 37, fig. 6. 1979. Type. Brazil. Bahia: 

Una, Fazenda Sao Rafael, 29 Oct 1969 (fl), T. 

S. dos Santos 457 (holotype, CEPEC; isotypes, 

FHO, NY). 

Additional specimens examined. BRAZIL: BAHIA: 8 

km S of Itacare, 16 Oct 1968 (fl), Almeida & T. S. dos 

Santos 164 (CEPEC, NY); 2 km S of Itacare, 14 Jun 

1972 (fl), T. S. dos Santos 2305 (CEPEC, NY); 28 km 

from Urucuca on rd. to Serra Grande, 15 Jul 1978 (fl), 

Mori et al. 10243 (CEPEC, NY); Bom Gosto, Ilheus, 

Mar 1943 (st), Fr6es 12711/80 (A). 

Local name. oiti-mirim. 

Parinari alvimii differs from all species of Par- 

inari, except P. rodolphii and P. campestris, in 

its large, amplexicaul stipules. It is quite different 

from these two species in the inflorescence, which 

Tree to 20 m tall, the young branches densely is compact and smaller with densely crowded 

ferrugineous-tomentose, becoming glabrous and flowers, in the larger flowers, and the leaf shape, 

conspicuously lenticellate with age. Leaf lamina cordate at base and blunt at apex. Parinari aloblong 

to ovate, coriaceous, 5-12 x 4-10 cm, vimii differs from P. littoralis in the stipules, the 

subcordate to cordate at base, rounded to very larger leaves, the thicker, rufous petioles, the 

bluntly acute at apex glabrous above except on larger bracts and flowers, and the denser inflomidrib, 

tomentose and with stomatal cavities rescences. It differs from two eastern Brazilian 

obscured by dense pubescence beneath, fre- species, P. excelsa and P. brasiliensis, in the stipquently 

with two glands at junction with petiole, ules and bracts, the leaf shape, and the compact 

midrib prominent beneath, slightly impressed and inflorescence. 

tomentose above; primary veins 17-25 pairs, extremely 

prominent beneath, impressed above; 3-9. Parinari klugii Prance, Fl. Neotrop. Monopetioles 

3-8 mm long, tomentose, terete. Stipules gr. 9: 190. 1972. 

20-35 mm long, semiamplexicaul, persistent, 

membranous, exterior pilose-tomentose at cen- Fruit narrowly oblong, 4-6 x 2.2-2.5 cm; exoter 

of base, gray-lanate over rest of surface, but carp minutely lenticellate; mesocarp thin, fleshy; 

with a few pilose appressed hairs, glabrous with- endocarp hard and fibrous, pubescent within. 

in. Inflorescences of terminal and subterminal Distribution (Fig. 88). This species was dedense-flowered 

panicles, 3-5 cm long, the rachis scribed from a single collection from the Deand 

branches light brown tomentose. Bracts to partment of San Martin, Peru. Recent collections 

1 cm long, persistent, ferrugineous, tomentose indicate that it is quite common in Loreto Deon 

exterior, enclosing flower buds in small groups. partment. 

Receptacle subcampanulate-turbinate, brown- Habitat. It occurs on riverside temporarily 

tomentose on exterior, tomentose within. Petals flooded forests and upland terra firme. 

five, white, slightly shorter than calyx lobes. Stamens 

5-7, unilateral, with short tooth-like stam- Representative additional specimens examined. 

inodes opposite them. Ovary and lower two- PERU. AMAZONAS: Huambisha, Rio Santiago, below 

thirds of style densely pilose. Fruit unknown. Caterpiza, 20 Nov 1979 (fr), Tunqui 105 (MO, NY). 

LORETO: Prov. Requena, Rio Ucayali, Arboretum Je- 

Distribution (Fig. 87). Known only from coast- naro Herrera, Jul-Sep 1976 (fl), Bernardi 16334 (G, 

al Bahia. 

NY); Rio Tacsha, Curaray, 19 Sep 1972 (fl), Croat 

Habitat. Littoral forest. 

20443 (MO, NY); Rio Itaya nr. Palo Seca, 20 Mar


1977 (fr), Gentry et al. 18466 (MO, NY). MADRE DE 

DIOS: Rio Manfi, Pakitza Station, 19 Nov 1980 (fr), 

Foster 5774 (F). 

Local name. yakuku. 

3-10. Parinari maguirei Prance, Fl. Neotrop. 

Monogr. 9: 190. 1972, descr. emend. Type. 

Guyana. Kaieteur Savannas, 11 May 1944 (fr), 

Maguire & Fanshawe 23390 (holotype, NY; 

isotypes, F, M, US). 

(Steyermark et al. 117792). However, there is no 

reason to believe that this represents any diversification 

since neither collection has a complete, 

intact inflorescence. The flowers of this species 

are typical for any Parinari, the only notable feature 

being the extremely long pubescence of the 

style. 

3-11. Parinari littoralis Prance, Fl. Neotrop. 

Mongr. 9: 191. 1972. 

Tree to 15 m tall, the young branches tomen- Distribution (Fig. 88). This distinct species was 

tellous, becoming glabrous with age. Leaf lamina described from a single collection from Marau, 

orbicular to elliptic, coriaceous, 2.5-6 cm x 1.5- Bahia. It appears to be confined to a small area 

4.5 cm, rounded to subcordate at base, rounded of the coastal restinga forests of Bahia. 

to apiculate at apex, glabrous above, tomentose 

Additional 

and with stomatal cavities beneath; midrib 

specimens examined. BRAZIL. BAHIA: 

Belmonte to Itapebi Rd., km 21, 18 May 1979 (fl), 

slightly impressed on upper surface, prominent Mattos Silva et al. 386 (CEPEC, NY); Marau, 3 Aug 

beneath; primary veins 9-14 pairs, plane above, 1967 (fl), da Vinha 12 (NY); 5 km S of Marai (fl), Mori 

prominent beneath; petioles 2-6 mm long, terete, 

& Carvalho 12001 (CEPEC, NY); Ilh6us, 21 km from 

tomentose. Stipules axillary, lanceolate, 1-3 mm Olivenqa, 25 Oct 1972 (fl), Pinheiro 1948 (NY). 

long, caducous. Inflorescences of terminal and Local name. oiti. 

subterminal panicles to 8 cm long, the rachis and The field notes of Mori & Carvalho 12001 note 

branches gray-brown-tomentellous. Receptacle that "the fruits are covered by billions of black 

subcampanulate-turbinate, tomentose on exte- bees (Melipona) which remove the pericarp." 

rior, densely pilose within and around throat and 

sparsely puberulous at base; pedicels 0.5 mm 3-14. Parinari parilis Macbride, Candollea 5: 

long. Petals five, equalling calyx lobes. Stamens 367. 1934. 

6-7, unilateral, with short tooth-like staminodes 

opposite them. Ovary inserted at mouth of re- 

This species, known in 1972 only by two colceptacle, 

pilose; style long-hirsute. Fruit elliplections, 

the holotype and a paratype, has now 

soid, 4-5 cm x 3-3.5 

been re-collected several times 

cm; exocarp verrucose; 

(Fig. 91). 

mesocarp thin, fleshy; endocarp hard, thick, ex- 

Habitat. It appears to be confined to the seaternally 

fibrous and granular, densely lanate sonally flooded tahuampa forest. 

within. 

Representative additional specimens examined. 

PERU. LORETO: Rio Mazan above La Libertad, 10 Jul 

Distribution (Fig. 90). Venezuela and Guyana 1976 (fl), Gentry & Revilla 16638 (MO, NY); Rio Naon 

the plateau shared by these two countries. nay opposite Santa Clara, 7 Apr 1977 (fr), Gentry 19094 

Habitat. Collected in savanna margin and gal- (MO, NY); Rio Napo, Quebrada de Zucusari, 5 Apr 

1979 

lery forest. 

(fr), Rimachi Y. 4378 (NY); Rio Loreto-Yacu, 10 

km from Colombian frontier, 6 Feb 1969 (fl), Sastre 

Additional specimens examined. VENEZUELA. & Echeverry 641 (P); Rio Tacsha, Curaray, 12 Sep 1972 

BOLiVAR: E of Icabaru, 18 Dec 1978 (fl), Steyermark (fl), Croat 20367 (AAU, MO); Rio Yaguasyaca, tribet 

al. 117792 (NY, VEN), 18 Dec 1978 (fl, fr), Stey- utary of Rio Ampiyacu, 7 Nov 1977 (fr), Gentry & 

ermark et al. 117824 Revilla 20371 

(NY, VEN). 

(MO, NY). 

This species, described from two fruiting col- 

Local names. parinari, parinari blanco. 

lections from Guyana, has now been collected in 

flower in nearby Venezuela. The new collections 

have characteristic small orbicular leaves with a 

3-15. Parinari chocoensis Prance, Fl. Neotrop. 

Monogr. 9: 194. 1972. 

blunt apex which distinguish it from all other 

neotropical species of Parinari. The inflorescence 

is rather lax in one collection (Steyermark et al. 

117824) and compact and clustered in the other 


from a single collection from the Rio Baudo in 

Choc6, Colombia, has now been collected in the 

Rio San Juan region of the same department.



CHOCO: Hoya del Rio San Juan, Quebrada Cunperro, 

below Noanama, 8 Apr 1979 (fr), Forero et al. 4862 

(COL, MO, NY); basin of Rio San Juan, Rio Taparal, 

Nov-Dec 1979 (fr), van Rooden et al. 586 (COL, MO, 

NY). 

3-17. Parinari romeroi Prance, Fl. Neotrop. 

Monogr. 9: 400. 1972. 


4. Exellodendron Prance 

from two collections from the Department of 

Narifto, Colombia, is now also known from the 

nearby forests of Esmeraldas, Ecuador. 

Additional specimen examined. ECUADOR. ESME- 

RALDAS: Borb6n, 8 Sep 1965 (fr), Little & Dixon 21014 

(NY, US). 

Local name. cuero de sapo. 

The distribution of the genus is shown in Figure 93, that of Exellodendron barbatum and E. 

cordatum in Figure 94, and E. coriaceum, E. gardneri, and E. gracile in Figure 95. 

4-5. Exellodendron 

gracile (Kuhlmann) Prance, 

Fl. Neotrop. Monogr. 9: 200. 1972. 

This most distinctive species of Exellodendron 

was known only by the fruiting type in Prance 

(1972). It is obviously now very rare, since it 

occurs only in the much destroyed forest of Es- 

pirito Santo, Brazil. A new collection shows that 

E. gracile has flowers typical of the genus. 

5-1. Maranthes panamensis (Standley) Prance & 

White, Brittonia 37: 76. 1985. Fig. 153. 

5. Maranthes Blume 

Distribution. This species was placed in synonymy 

under the Malesian Maranthes corymbosa 

by Prance (1972) and appeared to be restricted 

to Cerro Jefe in Panama in this 

hemisphere. It has now been collected in the 

province of Heredia in Costa Rica and in Nicaragua, 

and so, like Licania affinis, occurs in these 

moist forested areas of Central America. This 

confirms that M. panamensis is native to Central 

America and not introduced, as originally suggested 

by Prance (1968). The good material now 

available confirms that the single neotropical 

6. Couepia Aublet 

Revised Key to Species of Couepia 

Additional specimen examined. BRAZIL. ESPiRITO 

SANTO: Reserva Florestal Linhares, Estrada 161, 22 

Jan 1973 (fl), Spada 151 (INPA, RB 162365). 

Local name. guaiti-mirim. 

This is noted as a tree 34 m tall of the "mata 

de taboleiro." 

species of this otherwise African and Malesian 

genus is quite distinct. 

Additional specimens examined. NICARAGUA. 

ZELAYA: Rio Barbereba, 5 km from Nueva Guinea, 

16?47'N, 84?29'W, 27 Aug 1982 (fr), Araquistain 3149 

(MO). 

COSTA RICA. HEREDIA: Finca la Selva, Rio Puerto 

Viejo, 25 Feb 1982 (fl), Hammel 11260 (NY), 5 Apr 

1982 (fr), Hammel & Schatz 11577 (NY). 

PANAMA. PANAMA: Cerro Jefe, 8 Aug 1968 (fr), 

Correa & Dressier 958 (MO, NY), 29 Jul 1967 (fr), 

Dwyer & Gauger 7324 (COL); Cerro Azure, 11 Mar 

1977 (fl), Folsom et al. 1949 (MO, NY). CANAL ZONE: 

Pipeline Rd., 9 km NW of Gamboa, 29 Oct 1973 (fr), 

Nee 7675 (MO, NY). 

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. 

28. C. elata. 

3. Inflorescence with dense ferrugineous or gray-puberulous pubescence; receptacle cylindrical to 

obconical; bracteoles usually persistent.


4. Inflorescence gray puberulous; leaf venation only prominulous; fruit exterior glabrous and 

smooth. 6.1. C. bernardii. 

4. Inflorescence densely ferrugineous; leaf venation prominent; fruit exterior verrucose or tomentose. 

5. Leaf undersurface with distinct stomatal cavities; fruit exterior verrucose. 10. C. foveolata. 

5. Leafundersurface 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. 

9. 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, the 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. 

8.1. 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. 

14. 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; 

leaves with rounded to subcordate bases; 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. 32. C. insignis. 

13. Flowers 60-75 mm long; receptacle tubular, tomentose within almost to base; 

stipules 20-35 mm long, persistent. 

32.1. 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 20-25 

mm long. 

30. C. martinii. 

14. Leaves oblong-lanceolate, 3.5-7 cm broad; primary veins 17-25 pairs; receptacle 

12-15 mm long. 

31. 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. 

12. C. exflexa. 

15. Leaves 6-13 cm long, apex acuminate; exterior of receptacle with light-brown sericeous 

pubescence; stamens ca. 25; style pubescent for /4 of length. 13. C. habrantha. 

9. Bracteoles caducous or less than half the length of receptacle, under 10 mm long. 

16. Receptacle pubescent inside to the base; filaments hirsute; leaves hirsute beneath. 

33. C. recurva. 

16. Receptacle glabrous inside except at throat; filaments glabrous; leaves with arachnoid indumentum 

or glabrous beneath. 

17. Stamens less than 40. 

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. 13.1. 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 ca. 35. 13.2. C. carautae. 

20. Leaves broadly oblong, 12-24 cm long, 4.5-8.5 cm broad, ferrugineouslanate-pubescent 

beneath; receptacle ferrugineous on exterior; stamens ca. 

25. 11. 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 complete circle. 34. C. obovata. 

21. Leaves oblong-lanceolate, 9-15 cm long, with long acumen; bracteoles caducous; 

stamens unilateral. 1. C. guianensis. 

17. Stamens numerous (more than 60). 

22. Receptacle glabrous or with sparse appressed gray tomentum on exterior.


23. Leaves 9-17 cm long, densely appressed-lanate pubescent beneath; primary 

veins 12-15 pairs. 

35. C. williamsii. 

23. Leaves 5-8 cm long, sparsely pubescent beneath; primary veins 7-10 pairs. 

35.2. C. marleneae. 

22. Receptacle with dense brown pubescence on exterior. 

24. Leaves oblong-lanceolate. 

23. C. krukovii. 

24. Leaves oblong to oblong-elliptic. 

25. Primary veins 24-30 pairs. 

22. C. macrophylla. 

25. Primary veins 16-20 pairs. 

26. Leaf apex with prominent well-developed acumen. 36. C. chrysocalyx. 

26. Leaf apex rounded to bluntly acuminate. 24. C. latifolia. 

1. Inflorescence a panicle. 

27. Bracts and bracteoles persistent at flowering, at least half 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. 37. C. eriantha. 

28. Flowers 7-20 mm long; exterior of receptacle shortly brown- to gray-tomentose; 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. 15. C. bracteosa. 

30. Flowers 18-22 mm long; leaf bases rounded to subcuneate; rachis and receptacle not 

longitudinally striate. 17. 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. 

38. C. trapezioana. 

31. Stamens free almost to base; receptacle subcylindrical. 

16. C. subcordata. 

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. 3. 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. 19. C. excelsa. 

34. Petioles not canaliculate; flowers 8-12 mm long; exterior of receptacle and calyx lobes 

with sparse short gray pubescence; stamens 20-35. 18. C. caryophylloides. 

32. Interior of receptacle glabrous except for deflexed hairs at throat. 

35. Leaf underside 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 14-22 mm long, cylindrical-turbinate; 

stamens ca. 32; petioles 4-8 mm long. 

41. 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. 

41.1. C. dolichopoda. 

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

38. Inflorescence a much-branched corymbose panicle, rachis and branches glabrous; 

petioles 1.5-2.5 cm long; stamens 17-20; disc very thick with only small central 

cavity. 

8-4. Acioa edulis. 

38. Inflorescence little or much-branched, not corymbose; rachis and branches sparsely 

puberulous; petioles 4-12 mm long; stamens 25-120; disc thin with large central 

cavity. 

39. Leaves oblong to oblong-lanceolate, prominently acuminate; inflorescence 

little-branched, almost racemose; petioles 8-12 mm; stamens ca. 110. 

35.1. C. glabra. 

39. Leaves ovate to elliptic, bluntly acuminate; inflorescence much-branched; 

petioles 4-7 mm; stamens 25-45. 3. C. paraensis. 

35. Leaf undersurface lanate; exterior of receptacle pubescent. 

40. Exterior of receptacle and calyx lobes sparsely appressed-puberulous, 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. 

39. C. stipularis.


41. Stipules caducous, not adnate to base of petiole; rachis of inflorescence 1-2.5 mm 

thick; petal margins ciliate. 

42. Leaf undersurface with distinct, prominulous, parallel secondary venation, 

90? to primary veins. 6.1. C. bernardii. 

42. Leaf undersurface 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 much-branched panicle. 

44. Leaves ovate to oblong, 2.5-8.5 cm broad. 3. C. paraensis. 

44. Leaves oblong-lanceolate, 1.5-4 cm broad. 5. C. maguirei. 

43. Leaves prominently acuminate; receptacle cylindrical, ca. 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. 

40. 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 undersurface prominently reticulate. 

48. Leaves with deep, well-defined stomatal cavities, 2.5-6 cm long, bluntly acute 

at apex. 

29.1. C. amaralae. 

48. Leaves without stomatal cavities, acuminate at apex, or with poorly defined 

shallow cavities, 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. 42. C. cognata. 

49. Receptacle changing abruptly to a short thick pedicel; exterior of receptacle 

brown-pubescent; primary veins impressed on upper leaf surface. 

29. C. racemosa. 

47. Leaf undersurface with plane to prominent venation but not prominently reticulate. 

50. Receptacle slightly curved anteriorly in bud. 

51. Leaf with midrib much impressed above. 42. C. cognata. 

51. Leaf with midrib plane above. 

52. Leaves 4-7 cm long; primary veins 7-10. 53. C. pernambucensis. 

52. Leaves 9-18.5 cm long; primary veins 12-23. 

53. Exterior of receptacle and calyx lobes gray-puberulous; primary 

leaf veins 18-23, slightly impressed above. 51. C. impressa. 

53. Exterior of receptacle and calyx lobes brown-tomentose; primary 

leaf veins 12-14, plane above. 43. 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. 

55. C. parvifolia. 

56. Leaves orbicular to elliptic, acumen 0-3 mm long, base rounded 

to cordate; petioles 2 mm long. 

20. C. uiti. 

55. Leaves (4-)6-16 cm long; primary veins 9-17. 

57. Flowers 6-12 mm long; receptacle cylindrical, ca. 2 mm thick 

at top below calyx lobes. 

58. Inflorescence and flowers densely ferrugineous-sericeous; 

flowers 8-12 mm long; petioles 5-10 mm long; receptacle 

not striate on exterior. 52. C. meridionalis. 

58. Inflorescence and flowers with short gray pubescence; flowers 

6-8 mm long; petioles 3-7 mm long; exterior of receptacle 

longitudinally striate. 44. C. ulei. 

57. Flowers 12-20 mm long; receptacle subcampanulate, 4-6 mm 

thick.


59. Pedicels 6-15 mm long; exterior of receptacle and calyx 

lobes short gray-sordid-I erulent. 21. C. cataractae. 

59. Pedicels not exceeding 6 mm in length; exterior of receptacle 

and calyx lobes spreading brown-tomentose. 

60. Leaves 4-8.5 cm long, 2-3.8 cm broad, rufous-pubescent 

beneath, graying with age. 45. C. comosa. 

60. Leaves 7.5-18 cm long, 3.5-9 cm broad, gray-pubescent 

beneath. 

61. Leaves coriaceous, rounded to bluntly acuminate 

at apex. 

27. C. grandiflora. 

61. Leaves chartaceous, prominently acuminate. 

46. C. venosa. 

54. Petals glabrous except for ciliate margins. 

62. Receptacle cylindrical or subcylindrical. 

63. Inflorescence of densely clustered glomerules; primary leaf veins 

impressed above. 52.1. C. coarctata. 

63. Inflorescence a loosely branched panicle; primary leaf veins 

plane to prominent above. 

64. Stamens ca. 35; leaf base subcordate to rounded. 

16. C. subcordata. 

64. Stamens 15-28; leaf base rounded to cuneate. 

65. Flowers borne on distinctly articulated pedicels with 

lower part of pedicel persistent after fall of some flowers. 

6.2. C. monteclarensis. 

65. Pedicels not conspicuously articulated and no part remaining 

after flower fall. 

66. Leaves 10-16 cm long, rounded at base; primary 

veins 14-19 pairs. 

67. Receptacle cylindrical, usually glabrous at base 

within; stamens ca. 24. 38. C. trapezioana. 

67. Receptacle subcylindrical, pubescent within 

to base; stamens ca. 16. 47.1. C. nutans. 

66. Leaves 4.5-13 cm long, cuneate to rounded at 

base; primary veins 8-15 pairs. 

68. Receptacle cylindrical; stamens 20-28. 

6. C. sandwithii. 

68. Receptacle subcylindrical; stamens 11-21. 

47. C. polyandra. 

62. Receptacle broadly campanulate or turbinate. 

69. Stamens 14-20. 

70. Leaves ovate, 4-9 cm long; primary veins 11-15; flowers 

8-12 mm long; petioles 4-8 mm long. 25. C. ovalifolia. 

70. Leaves oblong, 10-18 cm long; primary veins 16-20; flowers 

11-15 mm long; petioles 10-15 mm long. 26. C. schottii. 

69. Stamens more than 28. 

71. Receptacle broadly turbinate, flattened, 3 mm long, almost 

solid. 48. C. platycalyx. 

71. Receptacle subcampanulate, not flattened, 4-12 mm long, 

hollow. 

72. Petioles 14-18 mm long; inflorescence densely crowded. 

52.2. C. longipetiolata. 

72. Petioles 2-10 mm long; inflorescence lax. 

73. Leaves 2.5-5.5 cm long, 1.5-3.5 cm broad. 

20. C. uiti. 

73. Leaves 9-27 cm long, 4-12 cm broad. 

74. Leaf margins undulate and revolute. 

49. C. rufa. 

74. Leaf margins plane. 

75. Leaf with short and blunt acumen, the 

lower surface rufous-pubescent; calyx 

lobes rufous-pubescent on exterior and



of Species of Couepia 

Distribution of Couepia as a whole is shown 

in Figure 96. 

yellow-brown-pubescent within. 

50. C. robusta. 

75. Leaf with finely pointed acumen, 8-14 

mm long, the lower surface gray-pubescent; 

calyx lobes gray-pubescent on both 

surfaces. 54. C. froesii. 

In Prance (1981) the circumscription of this 

species was changed considerably from that in 

Prance (1972). Three species recognized in 1972 

were merged and are treated as subspecies of this 

now rather well defined but variable species. 

6-1. Couepia guianensis Aublet, Hist. pl. Guiane 

1: 519, t. 207. 1775. Type. French Guiana. Key ta Subspecies of Couepia guianensis 

Aublet s.n. (lectotype, BM). 

1. Infloresnces of racemes (rarely with a few short 

Acia amara Willdenow, Linn, Sp. pi. ed. 4. 3(1): 7117. branctsB bearing 2 flowers), usually in at least 3 

1800. 

axils bulw the apex of branch; leaf lamina 9- 

Acioa amara Steudel, NomencLf bot. ed. 1. 9. 1821.. 16.3 crmlksng, chartaceous; petioles 6-9 mm long, 

Moquilea couepia Steudel, Nomeid. bot. ed. 2.2: 15. receptadleatways narrowly cylindrical. Plants of 

1849. 

terra firnrm. a. subsp. guianensis. 

1. Infloresemmes of panicles, usually only 1 or 2 

Tree to 25 m tall, the young branches puber- axils beliw apex; leaf lamina 5-10.5 cm long, 

ulous, soon becoming glabrous. Leaf lamina ob- coriaceoussa chartaceous; petioles 3-6 mm long; 

receptacle 

long to oblong-lanceolate, membranous to coc%lindrical 

or subcampanulate. Plants 

of terra firme or flooded river banks. 

riaceous, 4.5-16.5 x 2.5-5.5 cm, rounded to 2. Leaves chartaceous, the underside glabrous 

subcuneate at base, acuminate at apex, the acu- or sparsely pubescent, plants of flooded river 

men 5-18 mm long, glabrous above, densely 

banks. 

grayb. 

subsp. glandulosm. 

2. Leaves 

to brown-lanate, or glabrous, or with a thickLy coriaceous, the undersidte 

sparse 

densely lanate-lgpuescent, plant ofterra firme:. 

caducous pubescence beneath, frequently with 

c.. subsp. divaricatm. 

two glands atjunction with petioae; primary veiins 

10-15 pairs, plane above, prominent beneath; 

6-la. 

midrib prominulous above, prominent beneath; 

Couepia g_i ensis AutIlf subsp. _punensis. 

petioles 3-9 mm long, canaliculate 

Fig.. isM. 

above, pubescent 

when young, becoming glabrous and ru- Couepia leptostachyi Bentham ex Hooker f., FR. lmas. 

gose with age. Stipules 1-3 mm long, linear, early 14(2): 44.. 1867. Type. Brazil. Amazonas: Maumus 

caducous. Inflorescences terminal and (fl), Spruce 15(hiolotype, K; isotypes, BM, Q,GGE, 

axillary GOET, LE, M1 NY, OXF, P)) 

little-branched panicles or racemes, the rachis Couepia IvtolorBenoist, Bull Mus. Hist. NatL (aris) 

and branches sparsely puberulous to glabrous. 29: 596. 1923. Type. French Guiana (fl), M&linon 

Bracts and bracteoles minute, membranous, s.n. (holotype, P). 

ovate, caducous. Receptacle cylindrical to sub- Couepiasurinamensis Kleinhoonte, Recueil Trav. Bot. 

Needr 22: 390. 1925. 

campanulate, 4.5-10(12) mm long, 1-2.5 mm 

Type. Surinam. Sectie O (fl), 

B. W. 3080 (lectotype, U; isolectotypes, K, NY). 

broad below calyx, sparsely puberulous to glabrous 

externally, glabrous within except for de- Leaf lamina chartaceous, 9-16.3 cm long, the 

flexed hairs around throat; pedicels 0.5-4 mm lower surface densely lanate-pubescent; petioles 

long. Calyx lobes five, rounded, 2-2.5 mm long, 6-9 mm long. Inflorescence mainly of terminal 

puberulous or glabrous externally. Petals five, axillary racemes, usually in several axils below 

white, ciliate. Stamens 14-30, unilateral, insert- the apex, a few with short branches bearing two 

ed around half of a circle with short staminodes or three flowers. Receptacle 7-12 mm long, alopposite 

them. Ovary villous. Style pubescent ways narrowly cylindrical. Plants of terra firme. 

for at least half its length. Fruit rounded to ovoid, 

3-4 cm long, 2.5-3 cm broad; exocarp smooth, 6-lb. Couepia guianensis Aublet subsp. glanglabrous; 

mesocarp thin, fleshy; endocarp thin, dulosa (Miquel) Prance, Brittonia 33: 350. 

fragile, granular in texture, glabrous within. 1981. Fig. 105.


Couepia glandulosa Miquel, Stirp. surinam. select. 28. 

1851. Type. Suriname (fl), Hostmann & Kappler 859 

(holotype, U; isotypes, BM, GH, GOET, K, LE, M, 

P, S). 

Moquilea glandulosa (Miquel) Walpers, Ann. 2: 463. 

1852. 

Couepia myrtifolia Bentham ex Hooker f., Fl. bras. 

14(2): 44. 1867. Type. Venezuela. Amazonas: San 

Carlos (fl), Spruce 3681 (holotype, K; isotypes, BM, 

BR, F, GH, GOET, LD, LE, NY, P). 

Leaf lamina chartaceous to thinly coriaceous, 

5-10.5 cm long, the 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 one or 

two axils below the apex of branch. Receptacle 

5-10 mm, subcampanulate to narrowly cylindrical. 

Plants of flooded river banks. 

6-1c. Couepia guianensis Aublet subsp. divaricata 

(Huber) Prance, Brittonia 33: 351. 

1981. Fig. 104. 

Couepia divaricata Huber, Bol. Mus. Paraense Hist. 

Nat. 6: 75. 1910. Type. Brazil. Para: Belem, 14 May 

1901 (fl), Huber MG 2030 (holotype, MG; isotype, 

BM). 

Couepia divaricata Huber var. strictiuscula Huber, Bol. 

Mus. Paraense Hist. Nat. 6: 76. 1910. Type. Brazil. 

Para: Peixe-Boi, Sep 1908 (fl), R. S. Rodrigues MG 

8274 (lectotype, MG; isolectotypes, BM, P, RB 15104, 

neate at base, apiculate at apex, the 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; petioles 4-7 

mm long, tomentellous, becoming glabrous with 

age, rugulose, canaliculate above, eglandular. 

Stipules linear-lanceolate, tomentellous, to 6 mm 

long, caducous. Inflorescences of terminal and 

axillary little-branched panicles with central 

rachis and short 2-3 flowered branches, or of 

racemes, the rachis and branches puberulous. 

Receptacle cylindrical, 7-8 mm long, sparsely 

gray-puberulous externally, glabrous within except 

for deflexed hairs at throat; pedicels ca. 1 

mm long. Calyx lobes five, rounded, 2.5 mm 

long, puberulous externally. Petals five, white, 

the margins ciliate. Stamens 20-23, unilateral, 

the arc of the circle opposite to them toothed. 

Ovary tomentose. Style pubescent on lower portion 

only. Fruit ovoid, ca. 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 and bony, glabrous 

within. 

Distribution (Fig. 97). Western Amazonia. 

U). 

Parinari krukovii Gleason, Bull. Torrey Bot. Club 55: 

353. 1933. Type. Brazil. Terr. Rondania: Rio Machado, 

Tabajara, Nov-Dec 1931 (fl), Krukoff 1362 

(holotype, NY; isotypes, K, MICH). 

Representative specimens examined. PERU. LORETO: 

Prov. Requena, Rio Ucayali, Arboretum Jenaro Herrera, 

Jul-Sep 1974 (fr), Bernardi 7-84 (G, NY), 27 Aug 

1976 (fl bud), Bernardi 16219 (G, NY). 

BRAZIL. AMAZONAS: Mun. Humaita, Livramento, 

Oct-Nov 1934 (fr), Krukoff6635 (NY, US); Tres Casas, 

Sep-Oct 1934 (fr), Krukoff6381 (NY, US). 

Leaf lamina coriaceous, 6-10.5 cm long, the 

lower surface densely lanate-pubescent; petioles 

3-6 mm long. Inflorescences of terminal and ax- 

illary panicles with small few-flowered branches 

in only one or two axils below the apex of branch. 

Receptacle 5-7 mm long, cylindrical. Plants of 

terra firme. 

Specimen citations for these three subspecies 

were given in Prance (1981). 

6-6.1. Couepia bernardii Prance, Brittonia 33: 

347. 1981. Type. Peru. Loreto: Prov. Requena, 

Rio Ucayali, Arboretum Jenaro Herrera, 

4?55'S, 73?45'W, Tree Number 6-6, Jul-Sep 

1974 (fl), Bernardi 6-6 (holotype, NY; isotype, 

G). 

Tree, the young branches puberulous, soon be- 

coming glabrous. Leaf lamina elliptic, charta- 

ceous, 9-16 x 3.5-6.5 cm, rounded to subcu- 

Local names. Brazil: uchirana. Peru: parinari 

blanco. 

The closely related species Couepia bernardii, 

C. obovata, C. reflexa, and C. sandwithii form a 

superspecies (see Prance, 1972, for other similar 

groups in Couepia). The greater amount of material 

available now enables me to group these 

species together more clearly than was possible 

in 1972. They are characterized by the almost 

racemose inflorescence with small groups of 

flowers on short branches, thus technically panicles, 

the long tubular receptacle that is curved 

in bud at the apex, and the reticulate leaf undersurface 

with parallel secondary venation in 

all species except C. reflexa. I am maintaining 

all four species at present since they do appear 

to be distinct. Since C. reflexa is only known by


Table II 

Differences between species of the Couepia obovata species group 

C. bernardii C. obovata C. reflexa C. sandwithii 

Leaf laminae (cm) 9-16 

Apex acuminate 

Acumen length 2-8 

4-10 

apiculate or acuminate 

2-5 

11-14 

acuminate 

8-10 

4.5-9 

acuminate, rounded or 

apiculate 

5-6 

(mm) 

Flower 

exterior 

Leaves 

sparsely puberulous 

coriaceous 

glabrous or sparsely 

puberulous 

chartaceous to coriaceous 

sparsely puberulous 

chartaceous 

densely tomentellous 

coriaceous 

a single type collection it is still hard to evaluate. 6-6.2. Couepia monteclarensis Prance, sp. nov. 

The other species appear to be separate. They Type. Brazil. Minas Gerais: Estagao Biologica 

have distinct geographical distributions, al- Caratinga, 19 Feb 1984 (fl), M. A. Lopes 113 

though C. obovata and C. bernardii are sympatric (holotype, BHCB; isotype, NY). Fig. 14. 

in Amazonian Peru. C. obovata is much more 

widespread, in Central Amazonia and the South 

Species C. sandwithii et C. bernardii affinis, 

inflorescentiis 

of the Guianas, however. C. sandwithii is allolaxioribus, 

floribus, cum pedicellis 

articulatis munitis 

patric, found in the Orinoco Delta region and 

(1-5 mm longis), petiolis 8- 

10 mm 

Guyana. These species can be differentiated 

longis diversa. 

by 

the characters given in Table II and in the 

Tree, the young branches glabrous, conspicukey 

which follows. In the original description of C. 

ously lenticellate. Leaf lamina oblong, chartabernardii 

(Prance, 1981) I referred two collecceous, 

7.5-11 x 3.3-4.6 cm, subcuneate at base, 

tions from Venezuela to that species. These two 

apiculate at apex, the apex 4-6 mm long, glabrous 

collections 

above, with a short-brown-lanate 

(Steyermark 87610, 

pubes- 

88108) actually cence 

belong to C. sandwithii. 

beneath; midrib prominulous above, 

The species Couepia monteclarensis is also 

prominent beneath; primary veins 8-10 pairs, 

closely related to this group of species, but is 

plane above, prominent beneath; petioles 8-10 

mm 

more distinct. For differences see under that long, glabrescent, canaliculate, with a single 

gland on one side or eglandular. Stipules caspecies. 

ducous (not seen). Inflorescences of terminal 

panicles, 6-8 cm long, with a central rachis and 

short dichotomous branches bearing 2 to 4 flow- 

Key to Species of Couepia obovata ers, the rachis and branches gray-puberulous. Re- 

Species Group 

ceptacle cylindrical, 5-7 mm long, gray puberulous 

externally, glabrous within except for 

1. Flower exterior densely tomentellous, forming a deflexed hairs at 

complete covering; leaf underside usually ferru- 

throat; pedicels 3-5 mm long, 

gineous-brown-tomentose C. sandwithii. articulated at middle and with lower part per- 

1. Flower exterior glabrous or sparsely puberulous, sistent after flower dehiscence. Calyx lobes five, 

not forming a complete covering; leaf underside rounded to slightly acute, 2-3 mm long, gray 

gray-puberulous. 

2. 

puberulous on exterior. Petals 

Leaves membranous, venation smooth and 

five, white, glanot 

reticulate beneath, the apex with long acubrous 

except for ciliate margins. Stamens 18-20, 

men 8-10 mm long 

C. reflexa. inserted almost around complete circle. Ovary 

2. Leaves chartaceous to thinly coriaceous, re- inserted near mouth of tube, lanate. Style hirsute 

ticulate beneath with parallel secondary ve- for three-fourths of 

nation, the 

length. Young fruit ovoid; 

apex bluntly acute to acuminate. 

3. Leaf lamina 4-10 cm long, oblong to el- exocarp smooth, glabrous; mesocarp granular; 

liptic, the apex acuminate C. obovata. endocarp bony, glabrous within. 

3. Leaf lamina 9-16 cm long, obovate, the Distribution. Atlantic coast forest of Minas 

apex usually apiculate C. bernardii. Gerais.


\I I 3cm. '1'1~~~~~~~~~~~~~~ 

PA ]-~~~~~~~~~~~~~~~~~~. 

.-t I 

t GNllp^ 15 

a 

nwated;eclarem"' I 

mm !l t 

FIG. 14. Couepia monteclarensis (Lopes 113). A, habit; B, flower; C, flower section; D, petal; E, floral 

diagram; F, young fruit; G, fruit cross-section. 

E.


Additional specimen studied. BRAZIL. MINAS GERAIS: 

Estacao Biol6gica de Caratinga, 22 May 1984 (y fr), P. 

M. Andrade 252 (BHCB, NY). 

This species is named for the Monteclaro Reserve 

where it occurs in a vestige of Atlantic 

coastal forest that harbors this and many other 

severely endangered species. 

This species is most closely related to Couepia 

sandwithii from the Guianas and C. bernardii 

from western Amazonia. It differs primarily in 

the larger flowers and in the inflorescence which 

is much more open and with the flowers borne 

in pairs on long articulated pedicels. In addition, 

the petioles of C. monteclarensis are much longer 

and the secondary veins wider apart. This is a 

most distinct and easily recognized species. The 

two collections come from a small remnant of 

the Brazilian Atlantic coastal forest on the Fazenda 

Montes Claros in eastern Minas Gerais. 

The fact that new species are still being collected 

in what is left of those forests emphasizes the 

importance of their conservation. It is fortunate 

that World Wildlife Fund and the Fundaqao 

Brasileira para a Conservaqao da Natureza have 

a project at Montes Claros. It is to be hoped that 

this important tract of land can be permanently 

preserved. 

6-8.1. Couepia canescens (Gleason) Prance, Acta 

Bot. Venez. 9: 119-120. 1974. Type. Vene- 

zuela. Amazonas: Cerro Duida (fl), Tate 870 

(holotype, NY; isotype, K). 

Parinari canescens Gleason, Bull. Torrey Bot. Club 59: 

370. 1931. 

Couepia cognata (Steudel) Fritsch var. cognata pro parte 

sensu Prance, Fl. Neotrop. Monogr. 9: 149. 1972. 

Tree to 10 m tall, the young branches tomentellous. 

Leaf lamina oblong to oblong-lanceolate, 

coriaceous, 2-6.5 x 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, the 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; petioles 2-3 mm long, 

densely tomentose, terete. Stipules ca. 1 mm long, 

linear, tomentose, caducous. Inflorescences terminal 

and subterminal racemes 1.5-3 cm long, 

the rachis densely brown tomentose. Bracts and 

bracteoles elliptic, ca. 1 mm long, tomentose on 

exterior, caducous. Receptacle campanulate, 

straight, not curved, 3-4 mm long, brown-to- 

mentose on exterior, glabrous within except for 

deflexed hairs at throat; pedicels 0.5-1 mm long. 

Calyx lobes five, acute, tomentose on exterior. 

Petals five, white, glabrous. Stamens 12-16, uni- 

lateral with a few short staminodes opposite them. 

Ovary villous. Style glabrous except at extreme 

base. Fruit globose; exocarp glabrous and ver- 

rucose when mature; pericarp hard and thick, 

undifferentiated, densely lanate within. 

Distribution (Fig. 98). Guayana Highland region 

of Venezuela. 


AMAZONAS: Cerro Duida, Serra Parima, 54 km NW of 

Rio Orinoco, 2?27'N, 63?56'W, 18-23 May 1972 (fr), 

Steyermark 106117 (NY), (fl), Steyermark 105936 

(NY). BOLIVAR: Km 119 S of El Dorado, 12 Jan 1964 

(fr), Steyermark & Dunsterville 93035 (NY, VEN); SE 

bluffs of Chimanta-tepui, 21 May 1953 (fr), Steyermark 

75526 (NY); Abucapa-tepui, NW of Chimanta, 

18 Apr 1955 (st), Steyermark 75127 (NY). 

Couepia canescens is closest to C. cognata, but 

differs in the shorter racemose inflorescence, the 

shorter thicker pedicels, the calyx tube that is 

campanulate and erect, not curved, the shorter 

and caducous bracteoles, and the shorter pubescence 

of the inflorescence and flowers. In Couepia 

cognata the inflorescence is nearly always a 

slightly branched panicle; only rarely is it reduced 

to a raceme and then it is longer than that 

of C. canescens. These two species are ecologically 

distinct. Couepia cognata is a small tree of 

the forest on higher mountain slopes. 

6-13.1. Couepia scottmorii Prance, sp. nov. Type. 

Panama. Prov. Panama: Cerro Jefe, 1000 m, 

14 Jul 1975 (fl), Mori 7116 (holotype, NY; 

isotype, MO). Fig. 15. 

Species C. magnoliifoliae affinis, sed foliis mi- 

noribus, 2.5-5 cm longis, nervis primariis 8-14 

jugis, petiolis 2-5 mm diversa; a C. habrantha 

foliis minoribus, bracteolis minoribus caducis, 

nervis primariis impressis differt. 


puberulous. Leaf lamina elliptic, thickly coria- 

ceous, 2.5-5 x 1.6-3 cm, rounded to slightly 

subcordate at base, apiculate to bluntly acumin- 

ate at apex, the acumen 0-5 mm long, glabrous


Fi 

71 ?':~~~~~~~~~~~~~~~ m 

scodmor 

FI.1.Cupasctmri(oi716.A ai;B rut ,la nesufc;D lwr E lwrscin 

F, petals.~~~~~~~~~~~~~~~~~~~~~~~~~~'~3 

FIG. 15. Couepia scottmorii (Mori 7116). A, habit; B, fruit; C, leaf undersurface; D, flower; E, flower section; 

F, petals.


above, densely brown-arachnoid pubescent be- midrib prominulous above, prominent and alneath; 

midrib prominulous above, prominent most glabrous beneath, with only a few short stiff 

beneath; primary veins 8-14 pairs, slightly im- hairs; primary veins 16-20 pairs, plane above, 

pressed above, prominulous beneath; petioles 2- prominent beneath; petioles 6-10 mm long, gla- 

5 mm long, canaliculate, rugulose, puberulous. brous, rugulose, terete. Stipules linear, to 1 mm 

Stipules minute, lanceolate, caducous. Inflores- long, subpersistent. Inflorescences terminal and 

cences terminal and subterminal racemes 3.5- subterminal racemes, the rachis densely yellow- 

5.5 cm long, the rachis densely yellow-brown- brown-sericeous-tomentose. Bracteoles ovate, 6- 

tomentellous. Bracts and bracteoles caducous. 7 mm long, tomentose on exterior, caducous. 

Receptacle subcylindrical, 4-6 mm long, densely Receptacle subcylindrical, 7-8 mm long, yellowtomentellous 

on exterior, glabrous within except brown-sericeous on exterior, glabrous within; 

for deflexed hairs at throat. Calyx lobes five, acute. pedicels 0.5 mm long. Calyx lobes five, acute. 

Petals five, sparsely pubescent on exterior, with Petals five, glabrous on exterior, the margins cilciliate 

margins. Stamens ca. 26, inserted around iate. Stamens ca. 35, inserted around complete 

complete circle, glabrous except for mass of hair circle, filaments exserted. Ovary pilose. Style 

around interior of united bases of filaments. Style densely hirsute to apex. Fruit not seen. 

densely pubescent for three-fourths of its length. Habitat. Rain forest on clay soil. 

Ovary densely lanate. Fruit ellipsoid, 5 x 3 cm; Local name. milho torrado. 

exocarp smooth, glabrous; mesocarp thin, fleshy; Uses. The wood is used for railroad ties. 

pericarp ca. 3 mm thick. 

This species belongs to the Couepia magno- 

Distribution and habitat (Fig. 115). Cloud for- liifolia superspecies and is the first of that group 

est dominated by Clusia and Colpothrinax cookii to be found in eastern Brazil. The four related 

at 1000 m, on Cerro Jefe, Panama. 

species occur in Amazonia and the Guianas. It 

differs from C. 

Additional specimens examined. PANAMA. PANAMA: 

spicata and C. habrantha in the 

Cerro Jefe, 1000 m, Sep 1972 (fr), Gentry 6172 smaller and 

(MO, 

caducous bracteoles, the narrow 

NY); Cerro Jefe trail on ridge NE from summit, 1000 leaves and the greater number of stamens. It difm, 

18 Dec 1974 (fr), Mori & Nee 3741 (MO, NY). fers from C. magnoliifolia and C. reflexa in the 

This species also belongs to the smaller, 

Couepia magnarrowly 

oblong leaves, the glabrous, 

not 

noliifolia superspecies and extends that ferrugineous-pubescent, lower surface of the 

group 

from the Guianas to Panama. It is related to, but leaves, and in the greater number of stamens. It 

quite distinct from, C. habrantha and 

is 

C. 

most 

magclosely 

related to the Central Amazonian 

noliifolia, differing in the much smaller 

C. 

leaves 

habrantha. Couepia carautae is completely 

with fewer primary veins and shorter petioles. It 

different from the only two other species of eastdiffers 

from C. carautae in the small and 

ern Brazil with racemose 

elliptic 

inflorescences, C. inleaves 

with a densely pubescent undersurface. signis and C. bondarii, both of which have much 

larger leaves and flowers. Couepia carautae is one 

6-13.2. Couepia carautae 

of 

Prance, sp. nov. 

the few species of Couepia without a lanate 

Type. 

Brazil. Espirito Santo: Linhares, Reserva Flo- pubescence on the lower leaf surfaces and in this 

restal de CVRD, Estrada 143A6, km 

character it differs from all other eastern Brazil- 

1.430, 

Talhao 601, 24 

ian 

Jan 1978 (fl), J. Spada 31/78 species. 

It is with 

(holotype, INPA; isotype, NY). Figs. 16, 99. 

pleasure that I name this species for 

Dr. Pedro Carauta of Rio de Janeiro, who has 

Species C. magnoliifoliae affinis, sed foliis sub- done much to provide interesting material for 

tus glabris minoribus oblongo-lanceolatis, stami- my studies of Chrysobalanaceae. 

nibus 35, receptaculo extus brunneo-tomentoso 

differt; a C. habranthae foliis subtus glabris, brac- 

6-14. 

teolis minoribus caducis, staminibus 35 differt. 

Couepia spicata Ducke, Arq. Inst. Biol. Veg. 

2: 

Tree 22 m 

36. 1935. 

tall, the young branches glabrous. 

Leaf lamina narrowly oblong, coriaceous, 7-10 Distribution (Fig. 115). This species, known in 

x 2-3.3 cm, subcuneate at base, acuminate at 1972 only from the type found near Manaus, 

apex, the acumen 4-7 mm long, finely pointed, appears to be rather widely distributed. 

glabrous above, glabrous and waxy beneath; Habitat. It is a species of forest on terra firme.


'" 

GouAO,P 

caraute 

3cm. 

. 

A 

5% mm. 

FIG. 16. Couepia carautae Prance (Spada 31/78). A, habit; B, leaf undersurface; C, flower; D, flower section; 

E, petal. 


AMAZONAS: Manaus-Itacoatiara Rd., km 135, 11 Jul 

1975 (fl), Monteiro s.n. (INPA 50017); Novo Aripauna, 

Nova Prainha, 23 Jul 1976 (fr), Mota & Monteiro s.n. 

(INPA 60724), 31 Jul 1976 (fl), Mota & Monteiro s.n. 

(INPA 60898), 10 Aug 1976 (fl), Mota & Monteiro s.n. 

(INPA 61341). 

6-17. Couepia belemii Prance, Fl. Neotrop. 

Monogr. 9: 228. 1972.

72 


from two collections from coastal Bahia, appears 

to be quite common in the remnants of the south- 

ern Bahia wet forest. 

6-23. Couepia krukovii Standley, Publ. Field 

Mus. Nat. Hist., Bot. Ser. 17: 250. 1937. 

Distribution and habitat (Fig. 107). In 1972 

this species was still known only from the two 

type collections, the type and a paratype, from 

the Rio Madeira region. It appears to be confined 

to varzea forests of the central part of the basins 

of the Rios Madeira and Purus, extending into 

Bolivia. 


AMAZONAS: Rio Purus betwen Redencao and Itaboca, 

22 Nov 1971 (fl), Prance et al. 16315 (FHO, INPA, 

MG, NY, US). 

BOLIVIA. BENI: Rio Beni, Cachuela Esperanza, 28 

Feb 1924 (fl), Meyer 46 (Z). 


6-29.1. Couepia amaralae Prance, sp. nov. Type. 

Brazil. Amazonas: Base of Serra Araca, 3 km 

E of Rio Jauari, 0?49'N, 63?19'W, 27 Feb 1984 

(fl), Prance et al. 29261 (holotype, INPA; iso- 

type, NY). Figs. 17, 97. 


Marau, 18 Jan 1967 (fl), Belem 3169 (IAN, NY, UB); Species C. racemosae affinis, foliis minoribus 

20 km NW of Una, 27 Feb 1978 (fl), Mori et al. 9336 2.5-6 cm 

(CEPEC, NY). ESPIRITO SANTO: Reserva Florestal 

longis, subtus cavis stomatalis profun- 

CVRD, Linhares, 8 Nov 1977 (fl), Spada 007/77 dis, petiolis 4-5 mm longis differt. 

(INPA). 

Shrub 1-4 m tall, the young branches appressed 

lanate pubescent, becoming glabrous with 

6-18b. Couepia caryophylloides Benoist subsp. age. Leaf lamina oblong to oblong-elliptic, coglabra 

Prance, subsp. nov. Type. Brazil. Para: riaceous, 2.5-6 x 1.5-3.2 cm, rounded at base, 

Santarem-Cuiabfa Hwy., km 82, 30 Mar 1984 acute or bluntly acuminate at apex, glabrous 

(fl), Medeiros & Marinho 58 (holotype, IAN). above, prominently reticulate beneath with con- 

Fig. 99. spicuous deep stomatal cavities filled with short 

gray pubescence; midrib prominent and sparsely 

A subsp. caryophylloide receptaculo intus gla- lanate beneath, slightly impressed above; pribro 

differt. 

mary veins 9-10 pairs, prominent and lanate be- 

This material is identical to subsp. caryophyl- neath, plane to slightly impressed above; petioles 

loides except for the glabrous interior of the re- 

4-5 mm long, lanate when young, rugulose, terete, 

ceptacle, a consistent character in Couepia; I have 

eglandular. Stipules caducous. Inflorescences of 

thus recognized the new taxon, which is geo- little branched panicles or racemes, axillary and 

graphically distant from subsp. caryophylloides, terminal, the rachis and branches brown-tomenat 

the subspecific level. 

tose. Bracts and bracteoles ovate, caducous, tomentellous 

on both surfaces. Receptacle cam- 

6-22. Couepia macrophylla Spruce ex Hooker f., 

panulate-turbinate, 3-4 mm long, tomentellous 

Fl. bras. 14(2): 43. 1867. 

on exterior, glabrous within except for deflexed 

Distribution (Fig. 108). The range of this species hairs at throat. Calyx lobes five, acute, 2 mm 

has been considerably extended by recent col- long, tomentellous on both surfaces. Petals five, 

lections. 

white, glabrous. Stamens 35-40, inserted around 

half of circle. Ovary densely villous. Style gla- 

Additional specimens examined. PANAMA. DARIEN: 

Rio Ucurganti, 7 Jul 1967 (fl), Bristan 1121 

brous except for deflexed hairs at base. Fruit 

(MO, NY). 

glo- 

BRAZIL. ACRE: Rio Caete, tributary of Rio laco, 25 bose, 5 cm diam.; exocarp glabrous, lenticellate; 

Sep 1978 (fl), J. Ramos et al. 651 (INPA). 

pericarp 2-2.5 mm thick, hard and woody, tomentose 

within. 


AMAZONAS: Margin of Rio Jauari, Araca, 1 Mar 1977 

(fr), Rosa & Cordeiro 1713 (MG, NY); Rio Araca, 4 

Nov 1952 (fr), Fr6es 29277; 3 km S of Serra Araca, 

0?49'N, 63?19'W (fl), Rodrigues et al. 10509 (INPA, 

NY), Rodrigues et al. 10528 (INPA, NY). 

This distinctive new species belongs to the 

Couepia species group with deeply reticulate 

leaves. The lower leaf surface has deep stomatal 

cavities more similar to those that occur in Par- 

inari and a few species of Licania than to those 

of other species of Couepia. It differs from C. 

foveolata, which also has stomatal cavities, in the 

branched inflorescence and in the short blunt 

leaves, and from C. steyermarkii and C. canes-


W,.


cens in the branched inflorescence, the stomatal 

cavities and the leaf shape. It differs from C. 

racemosa in the much smaller, blunt leaves, the 

deep well-defined stomatal cavities, the shorter 

petioles and the lanate pubescence of the leaf 

venation below. 

It is with pleasure that this species is dedicated 

to Ieda L. do Amaral, the Brazilian botanist who 

was leader of the Araca expedition on which the 

type was collected. 

6-32. Couepia insignis Fritsch, Ann. K. K. Na- 

turhist. Hofmus. 5: 11. 1890. Fig. 106. 

This distinctive species was described from 

scant material, all from Bahia, Brazil but without 

precise locality, in Prance (1972). The nineteenth 

century material is now complemented by two 

recent collections indicating that this species still 

survives in the coastal forest. 


Mun. de Una, 27 km S of Olivenqa, 2 Dec 1981 (fl), 

Carvalho & Lewis 858 (CEPEC, NY); Restinga de Olivenca, 

1 Dec 1979 (fl), Mello Filho 3061 (CEPEC). 

6-32.1. Couepia cidiana Prance, Acta Amaz6n- 

ica 13: 21. 1983. Type. Brazil. Para: Mun. de 

Oriximina, Rio Paru do Oeste, left margin of 

Cachoeira Chuvisco, 7 Sep 1980 (fl), C. A. Cid 

et al. 2261 (holotype, INPA 96602; isotypes, 

FHO, NY). 

Treelet 8-10 m tall, the young branches tomentose. 

Leaf lamina oblong-lanceolate, coriaceous, 

15-26 x 7-9.5 cm broad, subcordate at 

base, acuminate at apex, the acumen 7-10 mm 

long, glabrous above, densely gray 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 ca. 3 mm from margin to 

form a conspicuous marginal vein; secondary 

veins more or less parallel, prominulous; petioles 

5-7 mm long, terete, eglandular, densely tomentose. 

Stipules linear, membranous, 2-2.5 cm long, 

persistent. Inflorescences few-flowered racemes, 

the 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 velutinous-tomentellous on exterior, tomentose 

within to base, sessile. Calyx lobes five, 

triangular, acute, tomentose on exterior, tomentose 

within except for glabrous lower portion, 

10-13 mm long. petals five, white, glabrous on 

exterior, the margins ciliate. Stamens ca. 165, 

inserted around a complete circle, fused at base 

to form a ring ca. 5 mm tall, the stamens inserted 

in several rows on exterior of ring, the interior 

of ring densely tomentose. Ovary inserted near 

mouth of tube, densely villous. Style sparsely 

hirsute pubescent on lower third. Fruit not seen. 

Distribution and habitat (Fig. 99). Forest on 

terra firme of western Para, Brazil. 

Additional specimen examined. BRAZIL. PARA: Rio 

Cachorro, Serra do Cachorro, 16 km NW of Cachoeira 

Porteira, 22 Jun 1980 (fl), Martinelli 7052 (INPA, NY). 

This species is closest to Couepia insignis, C. 

martinii, and C. bondarii. It differs from all three 

in the tomentose interior of the receptacle, an 

uncommon feature in the genus, occurring in only 

one other species with a racemose inflorescence, 

the Andean C. recurva, which is otherwise very 

different. Couepia cidiana also differs from its 

three closest relatives in the much longer flowers, 

the persistent and longer stipules and the large 

number (26-35) of primary leaf veins. 

6-33. Couepia recurva Spruce ex Prance, Fl. 

Neotrop. Monogr. 9: 242. 1972. 


scribed from a single Spruce collection from Ec- 

uador without locality. A second collection from 

the Ecuadorean Andes has now been examined. 

Additional specimen examined. ECUADOR. 

TUNGURAHUA: La Victoria, valley of Rio Pastaza, 1300 

m, 1 Dec 1939, Asplund 10055 (S). 

6-35.1. Couepia glabra Prance, Acta Amazonica 

2(1): 10. 1973. Type. Brazil. Amazonas: Rio 

Cuieiras, just below mouth of Rio Branquin- 

ho, Sep 1971 (fl), Prance, D. Coelho & Mon- 

teiro 14942 (holotype, NY; isotypes, FHO, 

INPA). 

Tree 10 m tall, the young branches glabrous. 

Leaf lamina oblong to oblong-lanceolate, coria- 

ceous, 11-21 x 4-7 cm, base subcuneate, apex 

acuminate, the 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; 

petioles 8-12 mm long, glabrous, canalic-


ulate, rugose, eglandular. Stipules lanceolate, 

membranous, ca. 2 mm long, caducous. 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 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 five, sparsely puberulous 

when young, soon becoming glabrous, 

the margins ciliate. Petals five, glabrous, caducous. 

Stamens ca. 110, inserted around a complete 

circle, the filaments tomentose around base, 

glabrous for most of length. Ovary pilose. Style 

pilose at base, glabrous above. Fruit unknown. 


lower Rio Negro region of Brazil. 

Habitat. Forest on terra firme. 

Small tree to 8 m tall, the young branches 

sparsely puberulous soon becoming glabrous. 

Leaf lamina oblong-lanceolate, coriaceous, 5-8 

x 1.5-2.4 cm, base subcuneate, apex acuminate, 

the acumen 5-15 mm long, with two 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; petioles 4-7 mm long, sparsely 

puberulous or glabrous, canaliculate above. Stipules 

triangular, 1 mm long. Inflorescences of ter- 

minal racemes, the rachis gray-puberulous. Bracts 

small and caducous. Receptacle cylindrical, 

swollen towards base, 15-18 mm long, sparsely 

puberulous on exterior, glabrous within except 

for dense pilosity around base of stamens. Calyx 

lobes five, rounded, sparsely puberulous on ex- 

terior, the margins ciliate. Petals five, white, the 

margins ciliate. Stamens numerous, inserted 

around a complete circle. Ovary densely pilose. 

Style pilose for half of length. Youngfruit ovoid, 

exocarp smooth, glabrous; mesocarp fleshy; en- 

docarp hard, fragile. 

Distribution (Fig. 108). Known only from south 

of Manaus. 



AMAZONAS: Manaus-Porto Velho Hwy., between Rios 

Castanho and Araca, 12 Jul 1972 (st), M. F. da Silva 

et al. 490 (INPA, NY), 11 Jul 1972 (fl), M. F. da Silva 

467 (INPA, NY); km 113, 12 Jul 1972 (fl), M. F. da 

Silva et al. 267 (INPA); Furo de Castanho, estrada 

Araca, 22 Jul 1972 (fl), M. F. da Silva 1027 (NY); Nova 

Olinda, Rio Paca, tributary of Rio Marimari, 2 Jul 

1983 (fl), Todzia 2295 (INPA, NY). 

Additional specimen examined. BRAZIL. AMAZONAS: Couepia marleneae is most closely related to 

Manaus-Caracarai Rd., km 115, Rio Urubu, 13 Aug C. williamsii, 

1974 (fl), Prance et al. 

differing in the much smaller leaves 

21636 (INPA, NY). and few primary nerves, the sparse pubescence 

Couepia glabra is quite unlike any of the species and more reticulate venation of the leaf underalready 

described, but it is probably closest to C. surface, and the more prominent conspicuous 

williamsii from which it differs in the usually glands at the junction of the lamina and the petpaniculate 

inflorescence, the larger leaves, the iole of the leaves. In C. williamsii the leaves are 

almost glabrous style, the glabrous underside of 9-18 cm long, and 2.5-5.5 cm broad, and have 

the leaves, as well as in a number of additional 12-15 pairs of primary veins. Couepia marlenesmall 

characters. 

ae is also rather similar to C. glabra, but it is not 

so closely related, differing in the unbranched 

6-35.2. Couepia marleneae Prance, Acta Ama- racemose inflorescence, the much smaller leaves, 

z6nica 4(1): 17. 1974. Type. Brazil. Amazonas: the very sparse but definite pubescence of the 

Manaus-Porto Velho Rd., between Rios Cas- exterior of the flowers, and the glands at the lamtanho 

and Tupana, 17 Jul 1972 (fl), M. F. da ina base. 

Silva et al. 822 (holotype, INPA; isotype, NY). 

6-38. Couepia trapezioana Cuatrecasas, Brittonia8: 

197. 1956. Fig. 118. 

Additional specimens examined. PERU. LORETO: Rio 

Ampiacu, Puca Urquillo, 24 Sep 1972 (fl), Croat 20663 

(MO, NY); Rio Nanay, Mishana, 16 Aug 1978 (fl), 

Ramirez C. 87 (MO, NY). 

6-41.1. Couepia dolichopoda Prance, Brittonia 

26: 302. 1974. Type. Peru. Loreto: Varadura 

de Mazan, from Rio Amazonas to Rio Napo 

(fl), Croat 19382 (holotype, NY; isotype, MO). 

Tree to 30 m tall, the young branches com- 

pletely glabrous. Leaf lamina elliptic, coria- 

ceous, 9-11.5 x 3.8-5.3 cm, cuneate at base, 


glabrous on both surfaces; midrib prominulous

76 

on both surfaces, glabrous; primary veins 6-8 

pairs, almost plane on both surfaces; petioles 10- 

12 mm long, glabrous, shallowly canaliculate. 

Stipules not seen. Inflorescences of pendulous 

panicles on long peduncles up to 75 cm, the rachis 

and branches glabrous, branches small and 

numerous. Bracts and bracteoles 0.5-1.5 mm 

long, acute, glabrous except for the sparsely ciliate 

margins, subpersistent. Receptacle campanulate, 

the tube 4-6 mm long when mature, contracted 

into pedicels 10-15 mm long, glabrous 

within and without. Calyx lobes five, rounded, 



BOLIVAR: Reserva Forestal Imataca, Rio Cuyuni Sector 

Caiio Negro, 15 Jan 1983 (fl), Stergios et al. 4991 (MO, 

NY); El Dorado, 3 May 1957 (fl), Couret 243 (US). 

647.1. Couepia nutans Prance, Brittonia 31: 248, 

fig. 1. 1979. Type. Colombia. El Valle: Alto 

Yunda, Rio Anchicaya, 1000 m, May 1973 

(fl), HiltyA-1 (holotype, US; isotypes, NY, UC). 

Tree to 25 m, the young branches sparsely pu- 

berulous, soon becoming glabrous. Leaf lamina 

elliptic, coriaceous, 13-16 x 5-7.5 cm, rounded 

at the base, acuminate at the apex, the acumen 

4-5 mm long, glabrous above, densely arach- 

noid-pubescent beneath; midrib prominulous 


pairs, plane above, prominent beneath; petioles 

5-8 mm long, weakly canaliculate above, pu- 

unequal, two considerably longer than the other 

three, sparsely puberulous when young, the margins 

ciliate, the exterior with two sessile conspicuous 

glands. Petals five, glabrous, caducous, 

red. Stamens 16-21, unilateral, with ca. eight 

tooth-like staminodes opposite to them. Ovary 

sparsely villous. Style glabrous. Fruit obpyri- berulous, with two glands. Stipules caducous, not 

form, 4.5 cm long, 3.5 cm wide at broadest point seen. Inflorescences of terminal and axillary panwhich 

is well below mid-point, exocarp smooth, icles to 7 cm long, the rachis and branches with 

glabrous; mesocarp thick, hard, fibrous; endo- short light brown tomentellous pubescence. Bracts 

carp thin and bony. 

ovate, membranous, ca. 5 mm long, caducous; 

Distribution (Fig. 101). Western Amazonia in bracteoles oblong, 2-3 mm long, puberulous on 

the Peru-Brazil frontier region. 

exterior, membranous, subpersistent. Receptacle 


subcylindrical, ca. 5 mm long, shortly tomentellous 

on the exterior, glabrous within except for 


Maynas, nr. Brilla Nueva, Rio Yaguasyacu, 8 Nov 1977 

deflexed hairs at the throat. Calyx lobes five, 

(fr), Gentry & Revilla 20385 (MO, NY); Quebrada Su- rounded, tomentellous. Petals five, white, glacusari, 

N side of Rio Napo below Mazan, 6 Nov 1979 brous except for the ciliate margins. Stamens ca. 

(fr), Gentry et al. 27591 (MO, NY). 

16, inserted in a complete circle. Ovary villous. 

BRAZIL. AMAZONAS: Rio Javari, 10 Aug 1973 (fl), 

Lleras et al. P17294 (INPA, NY), 17 Dec 1975 (fr), N. 

Style tomentose on lower third only. Fruit not 

T. Silva 4028 

seen. 

(IAN). 


Uses. The fruit (cotyledons) is edible. type gathering. 

Couepia dolichopoda is closest to C. longipen- Habitat. Collected at an altitude of 1000 m, 

dula but differs in the larger petioles, the reduced where it is an uncommon upland species. 

number of stamens, the glabrous peduncles, the Couepia nutans is closely related to C. polyanshorter 

campanulate calyx tube and the mark- dra, a Mexican and Central American species, 

edly unequal calyx lobes. 

the range of which extends south into Costa Rica. 

Couepia nutans differs from C. polyandra in the 

6-45. Couepia comosa Bentham, J. Bot. (Hook- larger leaves with a great number of primary 

er) 2: 215. 1840. 

veins, the slightly canaliculate petioles, the slightly 

Distribution and habitat (Fig. 100). This larger flowers with a darker brown pubescence, 

species 

and the 

was known until recently only from Guyana. It pendulous leaves. 

has now been collected in Venezuela and is not 

cited for that country in 

6-48. 

my account for the Flora Couepia platycalyx Cuatrecasas, Fieldde 

Venezuela (Prance, 1982b). The Guyana ma- iana, Bot. 27: 66. 1950. 

terial is from rocky places beside rivers and one This is one of the most distinctive species of 

of the Venezuelan collections (Stergios et al. 4991) Couepia because of the flattened, almost solid 

is from rain forest. There is no doubt that the receptacle. A third and fourth collection add to 

material of this distinct species is conspecific. the two studied in 1972.


Distribution and habitat (Fig. 111). It is con- lenticellate. Leaflamina oblong to oblong-ellipfined 

to the high altitude Andean forests of Co- tic, thickly coriaceous, 6-13 x 3.5-5.5 cm, sublombia 

and Venezuela. 

cordate to rounded at base, apex rounded, 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; 

petioles 6-10 mm long, lanate, with two glands 

at base 

6-51. Cnmepia imipessa Prance, Fl. 

(visible in young petioles only) becoming 

Neotrop. 

Mongr. 9: 255. 1972. Fig. 106. rugulose with age. Stipules 1-2 mm long, ovate, 

membranous, caducous. Inflorescences of ter- 

This species is now divided into two subspe- minal densely crowded panicles with many short 

cies, diiinguished in the following key. branches only, the rachis and branches puberulous. 

Bracts and bracteoles 

1. Leaves oblong-ovate, 19-22 cm long, 4-8.5 cm 

small, membranous, 

broad- primary veins 19-22 pairs. 

very early caducous. Receptacle cylindrical, 7-9 

a. subsp. impressa. mm long, with short dense puberulous pubes- 

1. Leaves ovate, 9-13 cm long, 4-7.5 cm broad; cence on exterior, glabrous within except for deprimary 

veins 14-19 pairs. b. subsp. cabraliae. flexed hairs at throat and dense lanate pubescence 

around stamen bases; pedicels 2-6 mm 

6-51b. C. impressa Prance subsp. cabraliae 

long, thin. Calyx lobes five, acute, with sessile 

Prance, Revista Brasil. Bot. 2: 31. 1979. Type. glands around margins. Petals five, glabrous ex- 

Brazil. Bahia: Santa Cruz de Cabralia, 5 Nov 

cept for ciliate margins. Stamens ca. 17, inserted 

1966 (fl), Belkn & Pinheiro 2837 (holotype, around a complete circle. Ovary densely lanate. 

CEPEC; isotypes, MG, NY). 

Style pubescent for three-fourths of its length. 

Leaves ovate, 9-13 cm long, 4-7.5 cm broad; Fruit elipsoid-pyriform, tapering towards apex, 

primary veins 14-19 pairs; petioles 7-10 cm long 4.5 cm long, 3 cm broad; exocarp smooth, glabrous; 

pericarp hard, fibrous 3 mm thick, gla- 


ANTIOQIA. Mun El Retiro, nr. quebrada La Aguadelo, 

2200-2300 m, 12 Apr 1980 (fr), Bernal & Galeano 142 

(HUA). 

VENEZUELA. LAA 8 km from Sanare, 1500-1800 

m, 29 Jul 1979 (f), Meijer et al. VEN92 (NY). 

Representative specimens examined. BRAZIL. BA- brous within. 

HIA: Santa Crz de Cabralia, 4 Nov 1966 (fl), BeIMm 

& Pinheiro 2846 (CEPEC, NY), 8 Feb 1967 (y fr), 

Habitat. Restinga. 

Belem & PineMro 3315 (CEPEC, NY), 21 Oct 1978 

Additional 

(fl), MFri et al. 10934 (CEPEC, NY); 9 km E of specimens examined. BRAZIL. BAHIA: 

Una, Mun. 

3 Dec 1981 (t Carwvao & Lewis 877 Marau, rd. to Porto de Campinhos, 17 km from 

(CEPEC, NY); 

Porto de Campinhos-Marai Rd., km 11, 26 Feb 1980 

Marau, 7 Jan 1982 (fl), Carvalho & Lewis 1106 (CE- 

(fr), Carmlko et al. 216 (NY); km 12, rd. Porto PEC, NY); km 11, Porto de Campinhos-Marai, 26 

Seguro Feb 1980 

to EunlpoEis, 9 Nov 1972 fr), Euponino 309 

(fl), Carvalho et al. 194 (CEPEC, NY). 

(CEPEC, 

NY), 7-9 km W of Porto Seguro, 1 Dec 1978 (fl), 

Eupoino 405 (CEPEC, NY); 4 km N of Uruguca, 4 

Nov 1978 (fl), Mori 11037 (CEPEC, NY); Ilheus, 12 

Nov 1979 (fl), T. S. dos Santos 1279 (CEPEC, NY). 

This is a common subspecies of restinga. 

Local name: oiti mirim. 

6-52.1. Couepia coarctata Prance, sp. nov. Type. 

Brazil. Bahia: Mun. de Marafu, 10 km S of 

Porto de Campinhos on rd. to Ubaitaba, 7 Jan 

1982 (fl, fr), Carvalho & Lewis 1116 (holotype, 

CEPEC; isotype, NY). Figs. 18, 100. 

Species a C. meridionali affinis, petalis extus 

glabris, nervis primariis supra impressis, calycis 

lobis glandulis sessilibus munitis differt. 

Small tree to 6 m tall, the young branches 

sparsely puberulous, soon becoming glabrous and 

This species is best distinguished by the dense- 

ly clustered inflorescence with numerous short 

branches, also by the small sessile glands around 

the margin of the calyx lobes, and the extremely 

lustrous leaf upper surfaces. It is probably most 

closely related to the more southerly C. merid- 

ionalis, from which it also differs in the im- 

pressed venation of the leaf upper surface and 

the glabrous petals. Couepia coarctata is named 

for the crowded, clustered inflorescence. It is not 

easily confused with any other species of the genus. 

It was collected in two places on coastal 

restinga on the small peninsula that runs north 

from Marau. 

6-52.2. Couepia longipetiolata Prance, Revista 

Brasil. Bot. 2: 31, fig. 3. 1979. Type. Brazil.

.~? 

'" 

?. 

'~~~~~~~~~~~~~~~~~~~~.'; .. 

Y~~~~~~~~~~~~~X 

l 

?? I /? I ?1 ----I , ,?=, E 

"r;~~~~~~~~~~~~~ 

?:~ ~~~~Cup.. . ~ ~ .- 

FIG. 18. Coucpia coarctata Carvalho & Zwwi$1 ] ] d). A habit; B, fruit; C, leaf udersurface; ]D, flower bud 

E, glandular margin of calyx lobe; F, flower section; G, petal.~~~~~? 

eoa 

CO4C&t,,1 

FIG. 18. Couepia coarctata (Carvaho & Lewis 16). A, habit; B, fruit; C, leaf undersurface; D, flower bud; 

E, glandular margin of calyx lobe; F, flower section; G, petal. 

78 

. 

78~~~~~~~~:


Bahia: Itacare, Estrada do Aeroporto, 28 Feb 

1975 (fl), T. S. dos Santos 2935 (holotype, CE- 

PEC; isotypes, FHO, NY). 

Tree 8 m tall, the young branches glabrous, 

conspicuously lenticellate. Leaf lamina oblong, 

coriaceous, 12-15 x 4-4.5 cm, cuneate at base, 

shortly and abruptly acuminate at apex, the 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 half above; primary veins 11-13 

pairs, slightly prominent beneath, slightly impressed 

to plane above; petioles 14-18 mm long, 

canaliculate on upper surface, lanate pubescent, 

transversely rugose, 2-3 mm thick. Stipules small, 

ovate, membranous, caducous. Inflorescences of 

terminal and subterminal panicles, densely 

crowded, short branched, the rachis and branches 

rufous-tomentellous. Bracts caducous, to 5 mm 

long, membranous, glabrous; bracteoles caducous, 

ca. 1 mm long tomentellous on e : rior. 

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 five, 

cream, glabrous with ciliate margins. Stamens ca. 

30, inserted around a complete circle, far exserted. 

Ovary villous-pubescent. Style pilose-tomen- 

7. Hirtella L. 

Distribution of Hirtella as a whole is shown in Figure 119. 

Key to Groups of Hirtella 

tose almost to apex, slightly exceeding the filaments. 


Distribution (Fig. 107). Known only from two 

collections from the forests of Bahia. 

Habitat. Littoral forest. 

Additional specimen examined. BRAZIL. BAHIA: 22 

km E of Ubaitaba, 25 Aug 1979 (st), Mori 12749 (NY). 

Local name. oiti. 

Couepia longipetiolata is related to the more 

southern species C. meridionalis, differing in the 

narrower leaves, the much denser inflorescence, 

the longer petioles and the glabrous exterior of 

the petals. It differs from C. schottii in the longer 

petioles, the fewer primary leaf veins, the dense 

inflorescence, and the greater number of stamens 

inserted around a complete circle. Couepia longipetiolata 

is not easily confused with any other 

eastern Brazilian species because of its long petioles 

and distinctive, densely crowded, shortbranched 

inflorescence. 

6-53. Couepia pernambucensis Prance, Fl. Neotrop. 

Monogr. 9: 256. 1972. 

Distribution 

(Fig. 111). This species, described 

from a single collection from Pernambuco, has 

now also been collected in Bahia. 

Additional specimen examined. BRAZIL. BAHIA: Una, 

Fazenda Sao Rafael, 16 Dec 1968 (fl), T. S. dos Santos 

322 (CEPEC, NY). 

1. Leaf base with myrmecophilous swellings. Group A: Section Myrmecophila. 

1. Leaf base without myrmecophilous swellings. 

Section Hirtella. 

2. Inflorescence a panicle (branched). Group B. 

2. Inflorescence a raceme (unbranched). Group C. 

Key to Group A: Hirtella Section Myrmecophila 

1. Inflorescence fasciculate (bunched racemes) sometimes cauliflorous. 

2. Stamens 4-5; ovary inserted at middle of receptacle; leaves 9-19 cm long, coriaceous, the lower 

surface bullate. 1. H. myrmecophila. 

2. Stamens 6-7; ovary inserted at mouth of receptacle; leaves 17-30 cm long, membranous, the lower 

surface not bullate. 2. H. physophora. 

1. Inflorescence an elongate raceme or panicle, never cauline. 

3. Inflorescence a panicle, sometimes only slightly branched at base. 

4. Inflorescence only slightly branched at base; leaves oblong-lanceolate; 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, with sparse hirsute pubescence. 

6. H. duckei. 

6. Pedicels 8-15 mm with dense hirsute pubescence. 6.1. H. revillae. 

Key to Hirtella Group B: Species without myrmecophilous swellings at 

leaf base and 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 6-7. 

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, the rachis and branches glabrous. 

7. H. macrosepala. 

5. Flowers borne in clusters of 2-3 along a central rachis rather than a branched inflorescence, 

giving a racemose appearance, the rachis and branches hirsutulous. 

6. Leaf lamina 6-9.5 cm long, 2.5-3.8 cm broad, the acumen 9-14 mm long; petioles 2- 

2.5 mm long. 

11.1. H. barnebyi. 

6. Leaf lamina 21-27 cm long, 6-9 cm broad, the acumen 1-5.5 mm long; petioles 8-10 

mm long. 

11.2. H. margae. 

4. Bracts with numerous stalked glands; inflorescence rachis and branches, and flower exterior 

pubescent. 

7. Leaves 3.5-8.5 cm long, 1.4-4.5 cm broad; stipules linear, to 6 mm long (Africa). 

90. H. zanzibarica. 

7. Leaves (6.4-)8-13 cm long, 4-7 cm broad; stipules deltoid, to 2 mm long (neotropics). 

8. Calyx lobes glandular; bract glands small, ca. 0.1 mm in diameter; rachis and branches 

of inflorescence sparsely puberulous; leaves glabrous beneath. 8. H. ulei. 

8. Calyx lobes eglandular; bract glands large, ca. 0.5 mm in diam.; rachis and branches 

of inflorescence tomentellous; leaves sparsely appressed pubescent beneath. 

9. H. glabrata. 


9. Glands large and solitary on a long stalk, arising from pedicel or junction of inflorescence; 

inflorescence distinctly thyrsoid. 

10. H. carbonaria. 

9. Glands usually numerous, long or short stalked, arising from the margins of bracts and 

bracteoles; inflorescence not thyrsoid. 

10. Inflorescence a long rachis bearing small clusters of flowers on very short branches; leaves 

oblong-lanceolate. 

11. H. araguariensis. 

10. Inflorescence spreading, with many distinct lateral branches, flowers not in distinct clusters; 

leaves oblong to elliptic. 

11. Leaves 3-5.5 cm long, cordate at base; inflorescence scarcely branched, hirsutulous. 

12. H. cordifolia. 

11. Leaves usually cuneate or subcuneate at base (if subcordate then 7-17 cm long and 

inflorescence glabrescent); inflorescences usually much branched. 

12. 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. 

13. Leaves 12-17 cm long, rounded to subcordate at base; fertile stamens 5. 

13. H. insignis. 

13. Leaves 3.5-15 cm long, cuneate at base; fertile stamens 3. 

14. Leaves 10-15 cm long; bracts membranous, the glands several, borne 

on margins; inflorescence 7-18 cm long, terminal. 14. H. tocantina. 

14. Leaves 4-9 cm long; bracts coriaceous, with single reflexed apical gland 

only; inflorescence 3-11 cm long, axillary or terminal. 15. H. piresii. 

12. 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. 

15. Leaves 1.8-4.5 cm long, rounded to acuminate at apex, thickly coriaceous; 

primary veins 4-7 pairs. 37. H. bahiensis. 

15. Leaves 6-13.5 cm long, acuminate at apex, thinly coriaceous; primary veins 

9-10 pairs.


16. Stamens 3; bracteoles coriaceous, ovate; petioles 3-4 mm long. 

16. H. davisii. 

16. Stamens 4; bracteoles membranous, lanceolate; petioles 1-2 mm long. 

17. H. subglanduligera. 

2. Leaves hirsute beneath at least on principal venation, usually more generally so. 

17. Calyx lobes with small stipitate glands. 

18. 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. 18. H. ciliata. 

18. Leaves oblong to ovate, 4.5-23 cm long, acute to acuminate at apex; trunk with thin bark; 

receptacle glabrous within except at throat. 

19. Venation of leaves prominulous above; inflorescence subcorymbose, bearing numerous 

tightly clustered flowers; stamens 7. 19. H. hoehnei. 

19. Venation of leaves impressed or prominulous above; inflorescence lax, spreading, not 

subcorymbose, the flowers loosely arranged; stamens 3-5. 

20. Leaves subcordate at base, subconduplicate, the lower surface merely sparsely 

appressed-pubescent. 

29. H. adderleyi. 

20. Leaves rounded to cuneate at base, rarely weakly subcordate but never subconduplicate, 

the lower surface hirsute. 20. H. glandulosa. 

17. Calyx lobes eglandular. 

21. Venation of leaves impressed above, or the leaf surface bullate. 

22. Inflorescence and flowers rufous-tomentose; bracts bearing very few sessile glands, these 

often obscured by pubescence. 

42. H. obidensis. 

22. Inflorescence and flowers tomentellous to puberulous, not rufous; bracts bearing many 

obvious stalked or sessile glands. 

23. Leaf surface usually distinctly bullate, the base usually subcordate. 21. H. bullata. 

23. Leaf upper surface plane, but with impressed venation, the base rounded to subcuneate. 

24. Bracteolar glands ca. 0.5 mm in diam., sessile or slightly stipitate; bracteoles 

distinct, ovate. 22. H. americana. 

24. Bracteolar glands ca. 0.1 mm in diam., on slender stalks; bracteoles reduced 

to a mass of glands with a slightly connate base. 27. H. tentaculata. 

21. Venation of leaves prominulous or plane above, the leaf surface never bullate. 


26. Bracteolar glands large, numerous, apparent; inflorescence tomentellous, manyflowered; 

exocarp glabrescent; endocarp thin and bony. 22. H. americana. 

26. Bracteolar glands small, few; inflorescence hispid, few-flowered; exocarp tomentose; 

endocarp thick and fibrous. 23. H. guatemalensis. 


27. Exocarp tomentose; endocarp thick and fibrous; bracts with few inconspicuous 

stipitate or sessile glands. 

28. Inflorescence hispid, the flowers lax; bracts with small short-stipitate glands; 

stamens 4. 23. H. guatemalensis. 

28. 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. 24. H. eriandra. 

27. Exocarp glabrescent; endocarp thin and bony; bracts with many conspicuous stipitate 

glands, except in H. liesneri. 

29. Inflorescence hispid-hirsute; bracts with short-stalked glands arising from their 

margins, the secretory tip small, ellipsoid. 

30. Inflorescences predominantly terminal; leaf bases rounded to weakly subcordate. 

25. H. paniculata. 

30. Inflorescences predominantly axillary; leaf bases deeply cordate. 

26. H. deflexa. 

29. 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, the secretory tip large, 

flattened, forming an expanded head. 

31. Glands arising from junction of pedicel and stem, with a long slender 

stalk and a small secretory tip, the whole appearing clavate; young stems 

puberulous or sparsely tomentellous. 28. H. macrophylla. 

31. 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.


32. Leaves 4-7 cm long, conduplicate; inflorescence 3.5-8 cm long; stipules 

lanceolate. 29. H. adderleyi. 

32. Leaves 6.5-15 cm long, not folded; inflorescence 12-19 cm long; 

stipules deltoid or linear-lanceolate. 

33. Inflorescences ca. 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. 11.3. H. liesneri. 

33. Inflorescences 12-15 cm long, not appearing racemose; glands 

numerous on bracteoles; inflorescence rachis tomentellous, never 

hispid. 

34. Leaves cordate at base; petioles 1-3 mm long; stipules linear, 

5 mm long; flowers densely crowded. 9.1. H. confertiflora. 

34. Leaves rounded to subcordate at base; petioles 5-7 mm long; 

stipules deltoid, to 2 mm long; flowers laxly arranged. 

9. H. glabrata. 

1. Bracts and pedicels eglandular. 

35. Leaf apex retuse, rounded, or acute, never acuminate. 

36. Stamens 5-6; leaf apex mostly rounded to retuse (or mucronulate). 30. H. punctillata. 

36. Stamens 3-4; leaf apex acute. 

37. Leaves elliptic, thick-coriaceous; inflorescence compact, to 3 cm long. 31. H. corymbosa. 

37. Leaves oblong, chartaceous; inflorescence lax and spreading, 3.5-17 cm long. 

36. H. triandra. 

35. Leaf apex distinctly acuminate, sometimes abruptly so and mucronate or cuspidate. 


39. Leaf base subcordate; inflorescence 25-55 cm long, pendulous. 

33. H. pendula. 

39. Leaf base rounded to cuneate; inflorescence 1-15 cm long, erect. 

40. Leaves lanceolate, at least three times longer than broad; inflorescence and exterior 

of flowers glabrescent. 

32. H. barrosoi. 

40. Leaves oblong to elliptic, not exceeding 2.5 times longer than broad; inflorescence 

and exterior of flowers usually pubescent, rarely glabrous. 

41. 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. 

50. H. bicornis. 

41. Inflorescence and exterior of flowers pubescent or tomentose; leaf apex acuminate 

but never cuspidate; inflorescences 2-17 cm long, predominantly terminal. 

42. Inflorescence and young stem hispid; calyx tube ca. 6 mm long, the lobes ca. 

5 mm long; corolla lobes ca. 8 mm long; stipules 6-10 mm long. 

34. H. leonotis. 

42. Inflorescence usually puberulous, rarely weakly hirsute; young stem glabrous 

to tomentellous; calyx tube to 4 mm long, the lobes to 3 mm long; corolla 

lobes to 5 mm long; stipules 2-6 mm long. 

43. Inflorescence little-branched, almost racemose, but with a few branches 

bearing two flowers, others one flower and a single pair of bracts at 

junction with pedicel. 

35. H. mutisii. 

43. Inflorescence much-branched, the branches usually bearing many bracts. 

44. Leaves 4-14.5 cm long, 2-5.5 cm broad; pedicels 1-3 mm long. 

36. H. triandra. 

44. Leaves 15-17 cm long, 8-10 cm broad; pedicels 0-0.5 mm long. 

38. H. latifolia. 


45. 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 three flowers each; leaf 

base sometimes cordate. 

46. Inflorescence almost a raceme, but with a few branches bearing two or three flowers; 

flowers laxly arranged, not borne in distinct groups. 

85. H. hebeclada. 

46. Inflorescence a long central rachis with groups of flowers on short branches. 

47. Bracteoles exceeding receptacle in length, persistent, enclosing buds; leaf base 

subcuneate; exocarp glabrous. 

39. H. suffulta. 

47. Bracteoles much shorter than receptacle, not persistent, not enclosing buds, leaf 

bases subcordate to subcuneate; exocarp tomentose or glabrous. 

48. Leaves 18-40 cm long, 8-16.5 cm broad, with 14-18 pairs of primary veins, 

and 2 glands at junction of upper surface of lamina and petiole; inflorescence 

rufous-tomentellous. 40.1. H. magnifolia.


48. Leaves 8-23 cm long, 3.3-8 cm broad, with 10-13 pairs of primary veins, 

the lamina base eglandular; inflorescence gray-puberulous or brown-tomentellous. 

49. Leaves subcordate at base, the lower surface glabrescent; exocarp glabrous. 

40. H. elongata. 

49. Leaves rounded to subcuneate at base, the lower surface sparsely pilosehirsute; 

exocarp tomentose. 24. H. eriandra. 

45. Inflorescence a much-branched panicle without a long central rachis; leaf base rounded to 

cuneate. 

50. Inflorescence and exterior of flowers tomentose, hirsute, or hispid, usually rufous to 

golden-brown. 

51. Inflorescence and young branches sparsely hispid; leaves membranous. 

41. H. rodriguesii. 

51. Inflorescence and young branches tomentose or hirsute; leaves chartaceous to 

coriaceous. 

52. Venation of leaves slightly impressed above; inflorescence rufous-tomentose. 

42. H. obidensis. 

52. Venation of leaves plane or prominulous above; inflorescence brown-tomentose 

or hirsute. 

53. Leaves 2.5-6 cm long; inflorescence 0.8-4 cm long. 

54. Inflorescence 0.8-1.5 cm long, clustered; leaves ovate, rounded at 

base; stamens 7. 43. H. cowanii. 

54. Inflorescence 2-4 cm long, lax; leaves orbicular to oblong-orbicular, 

usually subcordate at base; stamens 5. 44. H. orbicularis. 

53. Leaves 5.5-17 cm long; inflorescence 4-11 cm long. 

55. Bracts 4-6 mm long, ovate, up to half the length of receptacle; leaves 

ovate, thick-coriaceous, shortly mucronate at apex; inflorescence 

tomentose. 45. H. guyanensis. 

55. Bracts 1-3 mm long, oblong, to base of receptacle only; leaves elliptic, 

subcoriaceous, acuminate (never mucronate) at apex; inflorescence 

hirsutulous-tomentose. 46. H. lightioides. 

50. Inflorescence and exterior of flowers gray-puberulous to glabrescent. 

56. Inflorescence a subcorymbose panicle 4-6.5 cm long; petioles 7-8 mm long. 

47. H. aramangensis. 

56. Inflorescence a spreading panicle (not subcorymbose); petioles 0.5-5 mm long. 

57. Lamina 9-15 cm long, with distinct basal glands; inflorescence lax and spreading, 

7-19 cm long. 

48. H. rasa. 

57. Lamina 2.5-8(-9.5) cm long, the base eglandular or with indistinct glands; 

inflorescence short and compact, 1-8 cm long. 

58. Inflorescence and exterior of flowers densely puberulous; leaves thick 

and coriaceous, with prominent venation, the apex acuminate but never 

cuspidate. 

49. H. scabra. 

58. Inflorescence and exterior of flowers glabrescent to puberulous; leaves 

chartaceous, with plane to prominulous venation, the apex acuminate 

to cuspidate. 

50. H. bicornis. 

Key to Hirtella Group C: Species without myrmecophilous swellings at 

leaf base and 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 cm long, 0.5-2.2 cm broad; rheophytic. 

51. H. angustissima. 

2. Leaves ovate to oblong-lanceolate, 3-30 cm long, 1.5-10 cm broad; not rheophytic. 

3. Bracts and bracteoles bearing sessile glands only or with translucent glandular excretions, the 

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. Leaves oblong-lanceolate. 

5. Leaves 9-14 cm long; young stems puberulous; pedicels 5-10 mm long; stipules eglandular. 

52. H. tenuifolia. 

5. Leaves 5-8 cm long; young stems hispid; pedicels 12-16 mm long; stipules with numerous 

stipitate glands. 

52.1. H. radamii. 

4. Leaves oblong to ovate. 

6. Young stem hispid; inflorescence rachis sparsely puberulous. 

56. H. racemosa.


6. Young stem and inflorescence rachis pilose or puberulous, never hispid. 

7. Young stem and inflorescence thick-pilose or pilose-tomentose, the stem with persistent 

hairs 2-3 mm long; stamens 3-6. 

8. Leaf margins hispid-ciliate; stamens 5-6; pedicels 1.5-3 mm long; bracts axillary. 

53. H. pilosissima. 

8. Leaf margins eciliate; stamens 3; pedicels 3-6 mm long; bracts inserted on pedicels. 

34. H. leonotis. 

7. Young stem and inflorescence usually puberulous, sometimes tomentellous; stamens 

4-6. 

9. Leaves thin, membranous; stamens 4-5; ovary inserted midway up receptacle. 

41. H. rodriguesii. 

9. Leaves thick, coriaceous; stamens 5-6; ovary inserted at mouth of receptacle. 

10. Bracts and bracteoles caducous, with minute translucent secretions, sessile glands 

present or absent; pedicels 6-25 mm long. 

11. Bracts with sessile glands in addition to translucent secretion; pedicels 6- 

16 mm long, thick; inflorescence 4.5-14 cm long; leaves sparsely appressedpubescent 

on venation or glabrescent beneath. 54. H. gracilipes. 

11. Bracts with translucent glandular secretions only, other glands absent; pedicels 

12-25 mm long, very slender; inflorescence 1.5-5 cm long; leaves 

glabrous beneath. 55. H. brachystachya. 

10. Bracts and bracteoles persistent, with sessile glands only; pedicels 1.5-10.5 mm 

long. 

12. Primary veins 6-10 pairs; leaves 3.5-18.5(-19.5) cm long, glabrous or 

sparsely appressed-pubescent beneath; mature leaves plane, not bullate. 

56. H. racemosa. 

12. 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. 

13. Leaves 19.5-25 cm long; primary veins 13-15 pairs; lower surface 

with few sparse appressed hairs. 57. H. juruensis. 

13. Leaves 7-13 cm long; primary veins 11-13 pairs; lower surface hirsutulous. 

58. H. kuhlmannii. 

3. Either bracts and bracteoles bearing stipitate glands, or with one or more solitary stipitate glands 

arising from some pedicels, and bracts often glandular at apex only. 

14. Bracts and bracteoles eglandular, or with sessile glands and a glandular apex only; some 

pedicels with 1-3 stipitate glands. 

15. Leaves ovate, cordate at base; bracts and bracteoles with a few sessile glands. 

59. H. standleyi. 

15. Leaves oblong to oblong-lanceolate, usually rounded to subcuneate at base, rarely subcordate; 

bracts and bracteoles eglandular or with sessile glands. 

16. Primary veins 17-19 pairs; leaves 19-26 cm long; pedicels 8-12 mm long. 

60. H. longifolia. 

16. Primary veins 8-15 pairs; leaves 8-18 cm long (to 24 cm in one species with short 

pedicels 0.5-2 mm long); pedicels 0.5-9 mm long. 

17. 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. 

18. Fertile stamens 3; leaves oblong-elliptic; bracts with apical glands. 

61. H. lemsii. 

18. Fertile stamens 5-7; leaves elliptic to oblong-lanceolate; bracts glandular 

or eglandular. 

19. Leaves elliptic to oblong-elliptic, 3.5-8 cm long; bracts usually with 

sessile and apical glands. 


19. Leaves oblong-lanceolate, 8-18 cm long; bracts eglandular. 

20. Leaves subauriculate because of slight constriction above base; 

primary veins 11-13 pairs; ovary inserted near mouth of receptacle; 

fertile stamens 5. 62. H. schultesii. 

20. Leaves not constricted above base; primary veins 8-15 pairs; 

ovary inserted at mouth or middle of receptacle; fertile stamens 

4 or 6. 

21. Primary veins 12-15 pairs; secondary venation slightly impressed; 

ovary inserted midway up receptacle; fertile stamens 

4. 63. H. paraensis.


21. Primary veins 8-10 pairs; tertiary venation prominulous; 

ovary inserted at mouth of receptacle; fertile stamens 6. 

64. H. sprucei. 

17. Pedicels 1-3 mm long, thick; leaves hirsute beneath or glabrous with hirsute 

margin and midrib; bracteoles with glandular apex and often sessile glands. 

22. Leaves bullate above, with distinctly impressed venation, secondary venation 

and reticulations extremely prominent beneath. 65. H. lancifolia. 

22. Leaves not bullate above, the venation prominent or plane, secondary 

venation and reticulations prominulous only beneath. 

23. Leaves acuminate at apex, the lower surface glabrous to sparsely hirsute. 

66. H. burchellii. 

23. Leaves usually mucronate at apex, the lower surface densely hirsute 

or glabrous. 

24. Inflorescence densely tomentose; leaves mucronate at apex, hirsute 

beneath. 67. H. mucronata. 

24. Inflorescence sparsely puberulous; leaves acuminate at apex, glabrescent 

beneath. 56. H. racemosa. 

14. Bracts and bracteoles with numerous stipitate glands; pedicels eglandular. 

25. Receptacle cylindrical, elongate. 

79. H. couepiiflora. 

25. Receptacle campanulate. 

26. Leaves bullate above, with impressed venation. 21. H. bullata. 

26. Leaves plane, not bullate above, with plane or prominulous venation. 

27. Leaves orbicular, rounded to retuse at apex; pedicels 7-20 mm long. 

68. H. longipedicellata. 

27. Leaves oblong to elliptic, acuminate at apex; pedicels 4-11(-15) mm long. 

28. Stipules bearing numerous stipitate glands. 

69. H. glandistipula. 

28. Stipules eglandular. 

29. Inflorescence short, compact, densely flowered, 3-7 cm long; calyx 

bearing numerous glands; leaves drying gray. 70. H. martiana. 

29. Inflorescence lax, 3-28 cm long; calyx eglandular or the glands scarce; 

leaves drying green to brown. 

30. Calyx lobes 5-6 mm long; stamens 7-8; flowers 8-9 mm long. 

82. H. angustifolia. 

30. Calyx lobes 1.5-4 mm long; stamens 3-6; flowers usually 4-6 mm 

long (7-9 mm in H. santosii). 

31. Young branches and lower part ofinflorescence rachis hispidsetose. 

32. Leaves 2.5-6.5 cm long; inflorescence 3-5 cm long. 

71. H. pimichina. 

32. Leaves 6-15 cm long; inflorescence 6-28 cm long. 

33. Leaves entirely glabrous beneath, dried material with 

silver metallic sheen. 72. H. subscandens. 

33. Leaves hirsute beneath, especially on principal venation, 

dried material without silver metallic sheen. 

34. Bracteoles oblong, broad toward base, chartaceous, 

with short-stalked glands, the apices terminating 

in a short-stalked gland. 

25. H. paniculata. 

34. Bracteoles linear-lanceolate, coriaceous, with 

long-stalked glands, the apices always terminating 

in a long-stalked gland. 73. H. hispidula. 

31. Young branches tomentellous to puberulous; lower part of 

inflorescence glabrous to puberulous or tomentellous. 

35. Inflorescence tomentose or tomentellous. 

36. Inflorescence tomentellous, 4.5-27 cm long, leaves 

broadest above middle, primary veins 8-12 pairs; 

bract glands predominantly stipitate; stipules 2-3 

mm long. 

74. H. silicea. 

36. 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. 22.1. H. santosii.


35. Inflorescence glabrescent or very sparsely setose. 

37. Stamens 3; calyx glandular. 75. H. excelsa. 

37. Stamens 5-6; calyx eglandular. 

38. Leaf gradually tapered to apex from near base; 

pedicels 3-5 mm long. 76. H. adenophora. 

38. Leaf apex cuspidate; pedicels 8-15 mm long. 

77. H. caduca. 

1. Bracts and pedicels eglandular. 

39. Inflorescence clustered, fasciculate; leaves slightly bullate, 8-12.5 cm long; ovary inserted near base 

of receptacle. 

78. H. fasciculata. 

39. Inflorescence of elongate racemes; leaves usually plane, if bullate then small; ovary inserted at or 

near mouth of receptacle. 

40. Receptacle elongate-cylindrical; calyx lobes glandular. 

41. Inflorescences densely tomentose; leaves 10-15 cm long; flowers 17-20 mm long. 

79. H. couepiiflora. 

41. Inflorescences sparsely puberulous; leaves to 8.5 cm long; flowers 7-9 mm long. 

80. H. tubiflora. 

40. Receptacle campanulate; calyx lobes eglandular. 

42. Leaves orbicular, predominantly retuse to rounded at apex; pedicels 7-20 mm long. 

68. H. longipedicellata. 

42. Leaves oblong-lanceolate to ovate, acuminate at apex; pedicels to 15 mm long, usually 

shorter. 

43. Lower surface of leaf hirsute, pilose or hispid at least on primary and secondary 

venation. 

44. Leaves oblong, oblong-lanceolate, or lanceolate, the venation sometimes deeply 

impressed above; fertile stamens 7-9. 

45. Leaves oblong, 3.5-8 cm long, venation deeply impressed above; flowers 4 

mm long. 

81. H. floribunda. 

45. Leaves lanceolate to oblong-lanceolate, 5-15 cm long; venation plane above; 

flowers ca. 6 mm long. 

82. H. angustifolia. 

44. Leaves ovate to elliptic, the venation usually prominent above, rarely slightly 

impressed but then leaves broadly ovate; fertile stamens 3-9. 

46. Leaf venation slightly impressed above; leaves broadest near base, 2.5-8 cm 

long, subcordate at base; inflorescence densely crowded, with short rachis. 

83. H. rugosa. 

46. Leaf venation level to slightly prominent above; leaves broadest at or above 

middle, subcuneate to rounded at base; inflorescence lax with longer rachis. 

47. Stamens 3. 35. H. mutisii. 


48. Leaf apex mucronate to abruptly acuminate; pedicels 2-3 mm long. 

84. H. scaberula. 

48. Leaf apex acute to acuminate but never mucronate or abruptly 

acuminate; pedicels 5-8 mm long. 

49. Inflorescence and exterior of flowers densely tomentose; flowers 

5-8 mm long. 

85. H. hebeclada. 

49. Inflorescence and exterior of flowers puberulous, flowers 4-5 

mm long. 


43. Lower surface of leaf glabrous or with few sparse appressed hairs on midrib and 

primary veins. 

50. Leaves narrowly lanceolate, 5.5-18 cm long, 0.5-2.2 cm broad. 

51. H. angustissima. 

50. Leaves ovate to oblong. 

51. Fertile stamens 7-9. 

52. Petals slightly clawed, leaf base subcuneate; young branches glabrescent; 

inflorescence of densely crowded racemes. 70.1. H. parviunguis. 

52. Petals not clawed; leaf base subcordate to subcuneate; young branches 

puberulous to hispid; inflorescence of single racemes. 

53. Leaf base subcordate; calyx lobes 4-6 mm long; young branches 

sparsely hispid. 82. H. angustifolia. 

53. Leaf base subcuneate to rounded; calyx lobes 1.5-2.5 mm long; 

young branches sparsely puberulous. 

86. H. enneandra. 

51. Fertile stamens 3-6. 

54. Fertile stamens 3; inflorescence either 2-3-flowered or densely manyflowered 

and tomentose; filaments only slightly exceeding calyx lobes.



of Species of Hirtella 

7-4. Hirtella dorvalii Prance, Fl. Neotrop. 

Monogr. 9: 273. 1972. 

55. Leaves 7-13 cm long, 3-5 cm broad, chartaceous; inflorescence 2- 

3-flowered. 87. H. pauciflora. 

55. Leaves 3.5-5.5 cm long, 1.5-2.7 cm broad, thick-coriaceous; inflorescence 

many-flowered. 

88. H. glaziovii. 

54. Fertile stamens 4-6; inflorescence puberulous to tomentellous, manyflowered; 

filaments far exceeding calyx lobes. 

56. Leaves thick coriaceous; inflorescence 4-15 cm long. 56. H. racemosa. 

56. Leaves chartaceous; inflorescence 2-4 cm long. 

57. Leaf apex bluntly acuminate or acute; petioles 0.5-1 mm; flow- 

ers ca. 3 mm long. 

Distribution and habitat (Fig. 128). This little 

known myrmecophilous species was described 

from a single specimen by Prance (1972). Further 

collections, all from the same area, near Cara- 

carai, indicate that this is a locally abundant 

species occurring only in low caatinga forest on 

sandy soils. 

Additional specimens examined. BRAZIL. TERR. 

RORAIMA: Rio Ajarani, 28 May 1974 (fl), Pires & Cavalcante 

14375 (IAN, INPA), 1 Jul 1974 (fl), Pires & Leite 

14849 (IAN, INPA); Caracarai, 27 Apr 1974 (fl), Pires 

& Cavalcante 14360 (IAN, NY). 

7-6.1. Hirtella revillae Prance, Acta Amazonica 

8: 587, 89, fig. 6. 1979. Type. Peru. Loreto: 

Maynas, Rio Nanay, 4 km from Mishana, 150 

m, 19 Jan 1975 (fl), Gentry, Ayala & Revilla 

15807 (holotype, NY; isotype, MO). 

55.1. H. arenosa. 

57. Leaf apex caudate; petioles 1.5-2.5 mm; flowers ca. 6 mm long. 

55.2. H. conduplicata. 

in even at throat except around base of ovary; 

pedicels 8-15 mm long, hirsute. Calyx lobes five, 

lanceolate, hispid on exterior. Petals five, white, 

glabrous. Stamens six, the filaments far exceeding 

calyx lobes. Style hirsute on lower portion 

only. Ovary inserted at mouth of receptacle, glabrous 

around base. Fruit not seen. 

Distribution (Fig. 128) and phenology. Known 

only from the Rio Nanay in Peru, collected in 

flower from November through March. 

Habitat. Upland forest on white sand, poorly 

drained, swampy. 


Maynas, Mishana, Rio Nanay between Iquitos and 

Santa Maria de Nanay, 13 Nov 1977 (fl), Gentry 20678 

(NY), 25 Feb 1981 (fl), Gentry et al 31725 (MO, NY), 

19 Mar 1982 (fl), Gentry et al. 36438 (MO, NY). 

Hirtella revillae belongs to the section Myrmecophila, 

in which the swollen ant cavities at 

the leaf bases, and hispid pubescence are characteristic 

of all members. It differs from the other 

species in the section in the very long pedicels 

and in the distinctive dense hirsute pubescence 

of the pedicels and flowers, and the glabrous ovary 

and mouth of the 

Trees 8 m tall, the young branches 

receptacle. It is most closely 

hispid. Leaf 

related to H. 

lamina oblong, chartaceous-membranous, 19-22 

physophora. 

x 

The inflorescence is of rather intermediate 

8-10 cm, rounded at base and bearing two 

swollen ant cavities, abruptly acuminate at length in comparison to other species of section 

apex, 

the acumen 7-10 mm long, hirsute on venation Myrmecophila, which have either much more 

beneath, with sparse appressed hairs on compact fasciculate inflorescences or elongate raupper 

cemes or 

surface; midrib 

panicles. Only Hirtella duckei has raprominent 

beneath, prominucemose 

inflorescences as short as H. revillae but 

lous above, hirsute on both surfaces; primary 

H. duckei differs in 

veins 13-17 

many other ways. 

pairs, prominent beneath, prominulous 

above. Stipules linear, persistent, hispid. 

7-9.1. Hirtella confertiflora 

Inflorescences of axillary racemes 5-6 cm 

Prance, Brittonia 33: 

long, 354. 1981. 

the rachis light brown hispid, the lower Type. Venezuela. Amazonas: Rio 

pedicels 

longer than the upper ones giving a slightly cor- 

Coro-Coro, 5?35'N, 50?10'W, 22 Feb 1979 (fl), 

ymbose appearance. Bracts and bracteoles 

Steyermark et al. 117921 (holotype, NY; isolinear, 

persistent, hispid. Flowers 8-10 mm types, MO, VEN). 

long (excluding 

pedicels). Receptacle campanulate, light Tree 4 m tall, the young branches densely ferbrown 

hispid-hirsute on exterior, glabrous with- rugineous tomentose. Leaf lamina oblong-ellip-


tic, coriaceous, 10.5-12 x 4.8-5.7 cm, cordate 7-11.1. Hirtella barnebyi Prance, Brittonia 33: 

at base, shortly and abruptly acuminate at apex, 352. 1981. Type. Brazil. Rond6nia: Point 22 

the acumen 1-3 mm long, glabrous and shiny of RADAM SC-20-XD, margin of Rio Preto 

above, with sparse appressed hirsute pubescence nr. rapids, 30 Aug 1975 (fl), M. dos R. Cordeiro 

on lower surface; midrib prominulous above, 703 (holotype, INPA 57032; isotypes, IAN, 

prominent and appressed beneath; primary veins INPA 54595). Fig. 122. 

10-13 pairs, prominulous above, prominent be- 

Shrub 2 m 

neath, secondary venation prominulous on both 

tall, the young branches sparsely 

surfaces; petioles 1-3 mm long, terete, tomenhirsutulous, 

soon becoming glabrous. Leaf lamina 

tellous. Stipules lanceolate, to 5 cm long, ca- oblong, chartaceous, 6-9.5 x 2.5-3.8 cm, 

rounded at 

ducous, eglandular. Inflorescences of terminal and 

base, acuminate at apex, the acumen 

9-14 mm 

axillary panicles with central rachis and short long, glabrous beneath except for a few 

stiff 

densely clustered lateral branches bearing 2-4 appressed hairs on venation; primary veins 

8-11 

flowers, 12-15 cm long, the rachis and branches 

pairs, prominulous on both surfaces; midrib 

tomentellous. Bracts and bracteoles 1-3 mm prominulous above, prominent beneath; petlong, 

ioles 2-2.5 mm 

triangular to lanceolate, persistent, with numerlong, 

sparsely tomentellous, teous 

stipitate tack-like glands. Flowers 5-7 mm 

rete, eglandular. Stipules minute, persistent, ca. 

1 mm 

long. Receptacle campanulate, tomentellous ex- long, tomentellous. Inflorescences 8-12 

cm 

ternally, glabrous within except for reflexed hairs long, of little-branched panicles with a central 

rachis and small lateral branches 

at throat; pedicels 3-7 mm long. Calyx lobes 

bearing 2-3 

five, flowers 

acute, eglandular, tomentellous 

only, the rachis and branches very sparseexternally,puberulous 

within. Petals five. Stamens six, unily 

hirsutulous. Bracts and bracteoles 1-1.5 mm 

lateral, the filaments far exceeding calyx lobes. 

long, lanceolate, persistent, with sessile glands 

on 

Style sparsely hirsute on lower fourth. Ovary insome, 

and caducous bracteoles with large terminal 

serted at mouth of receptacle, pilose. Fruit not 

gland present only in bud. Flowers 6-7 

mm 

seen. 

long. Receptacle campanulate, with a few 


sparse appressed hairs on exterior, glabrous within 

type collection, from gallery forest bordering tree 

except at throat; pedicels 5-7 mm long, sparsesavanna. 

ly appressed hirsute. Calyx lobes five, reflexed in 

This species is closest to Hirtella glabrata, but 

open flowers, acute, hirsutulous externally, 

differs in the shorter, more compact inflores- densely gray-tomentellous within, the margins 

cence, the oblong leaves, the distinctly cordate eglandular. Petals five, glabrous. Stamens six, 

leaf base, the shorter petioles and the long linear 

unilateral, the filaments glabrous, or with a few 

hairs on lower 10 

stipules. H. confertiflora is also close to H. admm, 

free to base, far exceeding 

lobes. 

derleyi, but differs in the longer leaves that are Style hirsute up to one third of length. 

not conduplicate, in the compact inflorescence 

Ovary inserted near mouth of receptacle, villous. 


with shorter, weaker branches, and in the longer This is most 

secretory tip of the bracts. 

closely related to Hirtella araguariensis 

which has a similar subracemose inflorescence 

with a central rachis and short 

7-11. Hirtella araguariensis Prance, Fl. Neo- branches bearing 2-3 flowers. This type of inflotrop. 

Monogr. 9: 278. 1972. 

rescence occurs in only a few species such as H. 

eriandra and H. elongata, which are otherwise 

Distribution (Fig. 121). This species, described quite distinct from H. barnebyi. This species diffrom 

a type collection from Amapa, is much fers from H. araguariensis in the smaller, less 

more widely distributed, ranging from north of lanceolate leaves with rounded, not subcordate, 

Manaus, east to Amapa. 

bases, in the sparsely hirsutulous inflorescence 

and flowers, and in the much smaller stipules 

Additional specimens examined. BRAZIL. AMAPA: and bracteoles. 

Porto Grande, 29 Nov 1976 (fl), Rosa 1052 (MG, NY). 

AMAZONAS: Rio Uatuma, Mun. Itapiranga, Igarap6 Catita, 

21 Aug 1979 (fl), Cid et al. 640 (INPA, NY); Rio 7-11.2. Hirtella margae Prance, Proc. Kon. Ned. 

Pitinga, 27 Aug 1979 (fl), Cid et al. 860 (INPA, NY). Akad. Wetensch. Ser. C. 89: 111-113. 1986.


Type. Suriname. Brokopondo Distr.: Browns- 

berg Nature Park, 21 Jan 1978 (fl, fr), Roberts 

LBB16303 (holotype, NY). Fig. 138. 

Local name. Guyana: bokobokotokon. 

This species, closest to Hirtella araguariensis 

is distinguished by the larger, more coriaceous 

leaves, the hirsutulous dark pubescence of the 

inflorescence, and the less grouped flowers of the 

inflorescence. 

7-11.3. Hirtella liesneri Prance, sp. nov. Type. 

Venezuela. Tachira: Rio San Buena, 10 km W 

of La Fundaci6n, 7?47'N, 71046'W, 13-15 Mar 

1980 (fl), R. Liesner et al. 9633 (holotype, NY; 

isotypes, MO, VEN). Figs. 19, 136. 

Species H. araguariensi affinis, inflorescentiis 

Small trees to 3 m tall, the young branches puberulis versus basim hispidis, bracteolis glanglabrous, 

becoming lenticellate with age. Leaf duligeris, foliisque chartaceis diversa. 

lamina oblong to oblong-lanceolate, coriaceous, Tree 7 m tall, the young branches sparsely his- 

21-27 x 6-9 cm, rounded to subcordate at base, pid, becoming glabrous with age. Leaf lamina 

acuminate at apex, the acumen 1-5.5 cm long, oblong, chartaceous, 10-15 x 3.5-5 cm, subglabrous 

beneath except for a few stiff appressed cordate at base, acuminate at apex, the acumen 

hairs on venation; midrib prominulous and gla- 10-15 mm long, the lower surface with sparse 

brous above; prominent and with sparse ap- appressed hairs, especially on venation; midrib 

pressed pubescence beneath; primary veins 15- prominulous above, prominent and appressed 

18 pairs, prominulous above, prominent be- pubescent beneath; primary veins 12-16 pairs, 

neath; petioles 8-10 mm long, rugulose, with ap- prominulous on both surfaces; petioles ca. 2 mm 

pressed pubescence of stiff long hairs, terete, long, terete, sparsely hispid when young. Stipules 

eglandular. Stipules linear-lanceolate, persistent, linear, persistent, ca. 3 mm long, eglandular. In- 

7-9 mm long, eglandular, appressed pubescent. florescences ofterminal and axillary panicles, with 

Inflorescences of little-branched panicles with a a long central rachis and very short 3-5 flowered 

central rachis and small groups of flowers on lateral branches, giving a racemose appearance, 

short branches and solitary flowers; the rachis 20-22 cm long, the rachis puberulous, the lower 

dark brown-tomentellous. Bracts and bracteoles part sparsely hispid. Bracteoles triangular, mem- 

3-6 mm long, lanceolate, persistent, appressed branous, the apex terminating in a stalked gland. 

tomentellous, with few sessile glands on margins Flowers 6-7 mm long. Receptacle campanulate, 

of most bracteoles. Flowers 5-7 mm long, soli- sparsely hispid on exterior, glabrous within extary 

or in small clusters on central rachis. Re- cept for ring of deflexed hairs at throat; pedicels 

ceptacle campanulate, appressed hirsutulous on 2-4 mm long. Calyx lobes five, acute, eglandular, 

exterior, glabrous within except for pilose de- sparsely hirsute on exterior, puberulous within. 

flexed hairs around throat. Calyx lobes five, acute, Petals five, glabrous. Stamens six, unilateral, the 

appressed hirsutulous on exterior, gray-tomen- filaments far exceeding calyx lobes. Style hirsute 

tellous within, eglandular. Petals five, pale pur- on lower third. Ovary inserted at mouth of reple, 

glabrous. Stamens six, unilateral; filaments ceptacle, pilose. Young fruit ellipsoid; exocarp 

far exceeding calyx lobes. Style pilose on lower 

sparsely appressed pubescent, becoming glaportion, 

glabrous above. Ovary inserted near to brous with age. 

mouth of receptacle, pilose. Fruit ovoid, ca. 3 Habitat. Primary forested area on sandy soil, 

mm long, 1.8-2.5 cm broad, appressed-hirsu- 700-1000 m. 

tulous on exterior; mesocarp thin, fleshy; endo- This species is quite distinct in its inflorescarp 

thin, hard-bony, densely hirsute within. cence, and is one of the few hirtellas with a shortbranched 

Additional specimen examined. GUYANA. Puruni, 

elongated paniculate inflorescence that 

2 Apr 1953 (fr), J. Boyan 75 (FD 7749) (NY). 

resembles a raceme. It is probably closest to Hirtella 

araguariensis from which it differs in the 

sparsely hispid to puberulous (not tomentellous) 

inflorescence, the larger branches and pedicels, 

the fewer glands on the bracteoles, and the larger 

chartaceous leaves. 

7-13. Hirtella insignis Briquet ex Prance, Fl. 

Neotrop. Monogr. 9: 279. 1972. 

Distribution 

(Fig. 135). This species, described 

from just two collections, appears to be quite


G. 19. Hirtella liesneri (Liesner et a 9633). A, habit; B, bracteole; C, flower and petal; , flower section. 

FIG. 19. Hirtella liesneri (Liesner et al. 9633). A, habit; B, bracteole; C, flower and petal; D, flower section.


common in Atlantic coastal forests of Bahia and 

Espirito Santo. 

teoles ovate, persistent, tomentose with numerous 

sessile or shortly stipitate glands around margins. 

Flowers 7-9 mm long. Receptacle 

Additional specimens examined. BRAZIL. BAHIA: Km 

9, rd. Porto Seguro to Eunopolis, 8 Feb 1972 (fl), Eu- campanulate, tomentellous on exterior, glabrous 

ponino 212 (NY); 5 km N ofComandatuba, SE of Una, within except for deflexed hairs at throat; pedi- 

25 Jan 1977 (fl), Harley 18256 (K); rd. Porto Seguro cels 5-6 mm long, tomentose, eglandular except 

to Santa Cruz de Cabralia, 20 May 1971 (fl), T. S. dos on paired bracteoles. Calyx lobes five, acute, to- 

Santos 1677 (NY); Camaca, 24 May 1971 (fl), T. S. 

dos Santos 1695 (NY); Km 10, rd. Valenqa to 

mentose on 

Guabim, 

exterior, densely tomentose on in- 

22 Feb 1975 (fl), T. S. dos Santos 2898 (CEPEC, NY). 

terior. Petals five, glabrous. Stamens six, unilat- 

ESPiRITO SANTO: Reserva Florestal da CVRD, Lin- eral. Ovary inserted in middle of receptacle, 

hares, 8 May 1979 (fl), Foli 76/79 (INPA). pilose. Style pilose at base glabrous above. Fruit 

not seen. 

7-16. Hirtella davisii Sandwith, Bull. Misc. In- Distribution (Fig. 147). Known only from the 

form. 1935: 125. 1935. Fig. 128. type collection. 

This species, previously known from Vene- 

Habitat. Forest. 

zuela, Guyana, and Brazil, has recently been col- 

This species is close to the Caribbean Hirtella 

lected in Nicaragua and is another example of a 

americana but differs in the unbranched inflo- 

Guiana-Central America disjunction. Perhaps it rescence, the less impressed leaf venation, and 

has been overlooked, because it is a 

in 

large tree, 

many other minor characters. It is not easily 

30 m tall, unlike most other species of Hirtella 

confused with other species of Hirtella. 

which are shrubs and small understory trees. This 

species is noted as 30 m tall in both the Nicara- 7-29. Hirtella adderleyi Prance, Fl. Neotrop. 

guan and Guyanan collections. 

Monogr. 9: 296. 1972. Fig. 120. 

Additional specimens examined. NICARAGUA. RIO Representative additional specimens examined. 

SAN JUAN: Rio Santa Cruz, 1101 'N, 84?24'W, 22 Mar VENEZUELA. AMAZONAS: Depto. Atures, E of Sa- 

1985 (fl), Moreno 25541 (MO); Rio Sabalo, 11?02'N, nariapo, Apr 1979 (fl), Davidse et al. 6753 (NY); Cas- 

84?28'W, 23 Mar 1985 (fl), Moreno 25614 erio de 

(MO). 

Canaripo, 22 Aug 1978 (fl), Huber 2439 (NY, 

VEN); SE foot of Cerro Moriche, 19 Feb 1979 (fl), 

Huber 3205 (NY, VEN); Rio Ventuari, 10 km E of 

7-22.1. Hirtella santosii Prance, Revista Brasil. Carmelitas, 20 Feb 1979 (fl), Huber 3253 (NY); 22 km 

Bot. 2: 34, fig. 5. 1979. Type. Brazil. Bahia: S of confluence of Rios Manapiare and Ventuari, 27 

Una, Fazenda Sao Rafael, 10 Dec 1968 (fl), T. Feb 1979 (fl), Huber 3469 (NY); Serrania del Parfi, N 

S. dos Santos 300 

of 

(holotype, CEPEC; isotype, 

upper Rio Pari, 2 Mar 1979 (fl), Huber 3589 (NY); 

4 km N of Rio Sipapo, 28 Jul 1980 (fl), Huber 5577 

NY). 

(NY, VEN); E slopes of Cerro Calentura, 5?56'N, 

Tree 8 m tall, the young branches tomentel- 65?40'W, 11 Apr 1974 (fl), Jangoux 10111 (NY). 

lous, becoming glabrous with age, not conspicuously 

lenticellate. Leaf lamina elliptic, coria- 7-30. Hirtella punctillata Ducke, Arch. Jard. Bot. 

ceous, 7-12.5 x 2-5.5 cm, rounded to subcuneate Rio de Janeiro 3: 268. 1922; Prance, Fl. Neoat 

base, acuminate at apex, the acumen 4-10 mm trop. Monogr. 9: 296. 1972. Fig. 143. 

long, hirsute beneath on venation; midrib pro- This rather common species of the Guayana 

minulous above but in impressed channel, prom- Highland and the sandy caatingas of the upper 

inent and densely hirsute beneath; primary veins Rio Negro is interesting because in several col- 

15-19 pairs, prominulous above but in im- lections (for example Huber 10313 and 10875) 

pressed channel, prominent and densely hirsute most of the leaves are opposite. It is the only 

beneath; secondary venation slightly impressed species of Chrysobalanaceae known to have opabove 

giving lightly bullate appearance, promin- posite leaves. 

ulous beneath; petioles 3-4 mm long, terete, tomentellous, 

eglandular. Stipules linear, 4-5 mm 

7-31. Hirtella 

long, subpersistent, tomentellous. 

corymbosa Chamisso & Schlech- 

Inflorescences 

of terminal racemes to 6-10 cm long, the rachis 

tendal, Linnaea 2: 545. 1827. Fig. 127. 

and branches densely rufous-tomentose. Bracts Habitat. Growing on open restinga with sasmall, 

persistent, lanceolate, tomentose; brac- vannas.


Additional specimen examined. BRAZIL. BAHIA: 

Mun. de Mucuri, 7 km NW of Mucuri, 14 Sep 1978 

(fl), Mori et al. 10470 (CEPEC, NY). 

7-34. Hirtella leonotis Pittier, Arb. Arbust. Ve- 

nez. 2: 23. 1923. 

Fruit ovoid, ca. 2 cm in diam.; exocarp densely 

tomentellous. 

Distribution (Fig. 136). Forests of northern 

Colombia and Venezuela; in Venezuela only in 

the Federal District and State of Miranda. 

This species was known only from the type in 

1972. Recent collections from Cerros de Ba- 

chiller and Guatopo show that it is a most dis- 

tinct species. The fruit can now be described. It 

is one of the few species of Hirtella that has a 

densely tomentose exocarp. 

7-37. Hirtella bahiensis Prance, Fl. Neotrop. 

Monogr. 9: 307. 1972. Fig. 122. 

Additional specimens examined. BRAZIL. BAHIA: Nr. 

Santa Cruz de Cabralia, 21 Oct 1978 (fl), Mori 10936 

(CEPEC, NY); Ilh6us Rd. to Olivenga, 12 Nov 1970 

(fl), T. S. dos Santos 1284 (NY). ESPIRITO SANTO: Reserva 

Florestal CVRD, Linhares, 24 Nov 1978 (fl), I. 

A. Silva 30 (INPA). 

7-40.1 Hirtella magnifolia Prance, Acta Ama- 

zonica 8: 585, 587, fig. 5. 1979. Type. Brazil. 

Amazonas: Rio Javari, Estirao do Equador, 

21 Oct 1976 (fl), Prance et al. 23974 (holotype, 

INPA; isotypes, FHO, MG, MO, NY, US). 

Tree to 10 m tall, the young branches shortly 

tomentellous becoming glabrous and conspicu- 

ously lenticellate with age. Leaf lamina oblong- 

elliptic, chartaceous, 18-40 x 8-16 cm, the base 

rounded, abruptly acuminate at apex, the acu- 

men 7-13 mm long, curved, glabrous above, with 

a few stiff appressed hairs beneath on venation, 

with two glands at junction of upper surface of 

lamina and the petioles; primary veins 14-18 

pairs, prominent beneath, prominulous above; 

midrib prominent beneath, prominulous above, 

tomentellous on both surfaces; petioles 5-9 mm 

long, 3.5-6 mm thick, tomentellous, eglandular, 

terete. Stipules early caducous (not seen). Inflorescences 

terminal panicles with a long central 

rachis 12-18 cm long and many short few-flowered 

lateral primary branches, the rachis and 

branches rufous-tomentellous. Bracts and bracteoles 

ovate, persistent, gray-brown-tomentellous 

on both surfaces, eglandular. Flowers 5-6 

mm long. Receptacle campanulate, tomentellous 

Additional specimens examined. COLOMBIA. on 

ANTIOQUIA: Buenos Aires forest above rd. to Anori, 26 exterior, glabrous within except for sparsely 

Apr 1973 (fl), Soejarto et al. 4025 (MO); Autopista pilose area around throat; pedicels 1-2 mm long, 

Medellin-Bogoti, Setor Rio Samara-Rio Claro, 400- tomentellous. Calyx lobes five, acute, gray-to- 

1000 m, 19 Mar 1982 (fl), Herndndez & Albert de E. mentellous on both surfaces. Petals five, white, 

244 (HUA). 

glabrous. Stamens 5-7, unilateral with toothed 

VENEZUELA. MIRANDA: Parque Nacional de Guatopo 

(fr), Aristeguieta 7096 (VEN); Cerros del Ba- portion of ring opposite to them, filaments far 

chiller, 25 Mar 1978 (st), Steyermark & Davidse 116868 exceeding calyx lobes. Ovary inserted at mouth 

(MO, NY, VEN), 25 Mar 1978 (fl, fr), Steyermark & of receptacle, pilose. Style glabrous. Fruit not 

Davidse 116933 (MO, NY, VEN). 

seen. 

Distribution (Fig. 138). Known only from Loreto, 

Peru, and adjacent Brazil. 

Habitat. Upland forest on terra firme, understory 

in open clearings. 


Prov. Requena, Arboretum Jenaro Herrera, Jul-Sep 

1976 (fl), Bernardi 1-56 (G, NY), 15 Nov 1974 (fl), 

Diaz s.n. (G), 7 Dec 1977 (st), Gentry et al. 21213 

(MO, NY). 

Hirtella magnifolia has the largest leaves of 

any described species of the genus, often attain- 

ing 40 cm in length on the fertile branches. It is 

most closely related to H. elongata and H. er- 

iandra, but differs from both in the larger leaves 

with a greater number of primary veins. It differs 

further from H. elongata in the rufous-tomen- 

tellous pubescence of the inflorescence, the two 

glands at the junction of the upper surface of the 

leaf lamina and the petioles, the rounded not 

subcordate leaf bases, and the longer inflores- 

cence branches; and further from H. eriandra in 

the inflorescence branching and the laminar 

glands. 

7-48. Hirtella rasa Standley, Publ. Field Mus. 

Nat. Hist., Bot. Ser. 17: 252. 1937. Fig. 146. 

Additional specimen examined. PERU. LORETO: Pu- 

callpa, km 75, 9 Sep 1980 (fl), Angelo 11 (MO, NY). 

Local name. lobo apacharama.


7-52.1. Hirtella radamii Prance, Acta Amaz6- 

nica 12: 22. 1983. Type. Brazil. Rond6nia: Iga- 

rape Preto, RADAM SC-20-XA-Ponto 27, 

62?14'W, 8?58'S, 30 Jun 1975 (fl),J. C. daSilva 

100 (holotype, MG). Fig. 143. 

Shrub 3 m tall, the young branches hispid. Leaf 

lamina oblong-lanceolate, chartaceous, 5-8 x 

1.5-2.3 cm, subcordate at base, acuminate at 

apex, the acumen 3-5 mm long, glabrous on both 

surfaces; midrib prominulous above, prominent 


and glabrous on both surfaces; petioles 1- 

1.5 mm long terete, sparsely hispid, eglandular. 

Stipules lanceolate, to 5 mm long, membranous, 

caducous, with numerous long-stipitate glands. 


glabrous above, glabrous beneath except for 

appressed pubescence on midrib; midrib prom- 

inulous above, prominent beneath; primary veins 

8-10 pairs prominulous on both surfaces; peti- 

oles 0.5-1 mm long, terete, eglandular, glabrous 

or with a few adpressed hairs. Stipules 2 mm 

long, linear, subpersistent. Inflorescences ter- 

minal and subterminal racemes, 3-4 cm long, 

the rachis sparsely hispid. Bracts and bracteoles 

narrowly triangular, persistent, eglandular, ca. 1 

mm long, sparsely hispid on exterior. Flowers 3 

mm long. Receptacle campanulate, sparsely hispid 

on exterior, almost glabrous within except 

for a few appressed hairs with a ring of long 

deflexed hairs around base of staminal 

Inflorescences of terminal racemes to 12 cm ring; pedlong, 

icels 5-7 mm 

the rachis glabrous. Bracts and bracteoles 1.5-3 

long, eglandular, sparsely hispid. 

mm long, oblong, with two sessile glands toward 

Calyx lobes five, acute, reflexed, sparsely hispid 

on 

apex, a few of the youngest bracteoles with one 

exterior, densely gray-puberulous on interior. 

Petals 

or two stipitate glands in addition to the sessile 

five, purple, glabrous. Stamens 5-6, unipaired 

glands. Flowers ca. 5 mm 

lateral, far exserted. Ovary tomentose, inserted 

long. Receptacle at mouth of 

campanulate, glabrous on 

receptacle. Style pilose. Fruit unexterior, 

glabrous known. 

within except for deflexed hairs at throat; pedi- Habitat and 

cels 12-16 mm 

phenology. This species is known 

long, extremely slender, eglan- from two white sand 

dular. Calyx lobes five, acute, glabrous on extecampinas 

of Central Amazonia. 

Collected in flower in 

rior, puberulous within, eglandular. Petals 

July. 

five, 

glabrous. Stamens six, unilateral, the filaments Additional specimens examined. BRAZIL. 

far exceeding the calyx lobes, glabrous. Style hir- AMAZONAS: Manacapuru, Igarap6 Branco, 6 Jul 1958 

sute for half of length. Ovary inserted at mouth (fl), Ferreira 58-320 (INPA, NY); Serra dos 6 Lagos, 

18 Jul 1979 

of receptacle, pilose. Fruit not seen. 

(fl), Maia et al. 663 (INPA). 

Local name. caripe torrado. 

7-55.2. Hirtella conduplicata Prance, Brittonia 

This species is closest to Hirtella tenuifolia, 28: 227, fig. 12. 1976. Type. Brazil. Amazonas: 

from which it differs in the smaller leaves, the 

Lago de Castanho-Mirim, 25 Jun 1973 (fl), B. 

hispid pubescence of the young stem, and the 

Albuquerque et al. 887 (holotype, INPA). 

much longer pedicels. It differs from H. racemosa 

Fig. 125. 

in the narrow, oblong-lanceolate leaves, the long 

pedicels and the hispid young stems. It differs Tree to 20 m tall, the young branches sparsely 

from H. sprucei in the smaller leaves, the long hispid. Leaf lamina conduplicate, oblong-ellippedicels, 

the bracteolar glands and the absence tic, chartaceous, 3.5-5.5 x 2-2.5 cm, subcordate 

of pedicel glands. 

at base, caudate at apex, the acumen 8-15 mm 

long, glabrous above, glabrous or with sparse ap- 

7-55.1. Hirtella arenosa Prance, Brittonia 28: 

pressed pubescence on veins beneath, midrib 


prominulous on both surfaces; primary veins 5- 

7 

Manaus-Itacoatiara Rd., km 180, 7 Jul 1980 

pairs, prominulous on both surfaces; petioles 

1.5-2.5 mm 

(fl), W. A. Rodrigues et al. 8257 

long, terete, eglandular, sparsely his- 

(holotype, 

INPA). Fig. 122. 

pid. Stipules ca. 1 mm long, caducous, hispid. 

Inflorescences of terminal and axillary racemes 

Shrub to 3 m tall, the young branches puber- 2-3.5 cm long, the rachis sparsely hispid. Bracts 

ulous, soon becoming glabrous. Leaf lamina and bracteoles ca. 1 mm long, oblong, acute, 

oblong-elliptic, chartaceous, 3-5.5 x 1.2-2 cm, eglandular, sparsely hirsute. Flowers 5-6 mm 

cuneate at base, acute, rounded, or shortly long. Receptacle campanulate, sparsely hirsute


on exterior, glabrous within except for circle of Amazonia. Although it is quite distinct by its 

deflexed pilose hairs around staminal ring; pedlong, bracteate inflorescence, it is little collected 

icels 6-10 mm long, eglandular, sparsely hirsute, and apparently rare. One recent collection has 

thick. Calyx lobes five, rounded, reflexed, sparse- been made. 

ly hirsute on exterior, densely gray-puberulous 

Additional 

within. Petals five, glabrous, white. Stamens 

specimen studied: BRAZIL. PARA: Mun. 

six, de Itaituba, Santar6m-Cuiaba Hwy., km 780, Serra do 

unilateral, far exserted, purple, connate and hir- Cachimbo, 9?22'S, 54?54'W, 29 Apr 1983 (fl), Amaral 

sute towards base. Ovary pilose, inserted near et al. 1041 (INPA, NY). 

mouth of receptacle. Style hirsute. Fruit unknown. 

7-61. Hirtella lemsii L. O. Williams & Prance, 

Habitat and phenology. This species was col- Fl. Neotrop. Monogr. 9: 334. 1972. Fig. 136. 

lected only at the type locality in rain forest on 

terra firme, flowering in June. 

This species, described from a single collection 

from Costa Rica, appears to be quite common 

Additional specimen examined. BRAZIL. AMAZONAS: in that 

Lago de Castanho-Mirim, 25 Jun 1973 country, and the fruit is described for the 

(fl), Albuquerque 

et al. 886 first time. 

(INPA, NY). 

Fruit oblong, ridged, and with two small protrusions 

when dry, slightly tapered towards base, 

2.5-3 cm long; exocarp sparsely tomentose when 

young; mesocarp thin and fleshy; endocarp hard, 

thin. 

7-56d. Hirtella racemosa Lam. var. hispida 

Prance, Proc. Kon. Ned. Akad. Wetenschapp. 

Ser. C. 89: 113. 1986. Type. Brazil. Amapa: 

Rio Falsino, 10 km above Rio Araguari, 26 

Aug 1983 (fl), B. V. Rabelo et al. 2381 (ho- 

lotype, MG; isotype, NY). Fig. 146. 

The young branches hispid. Leaf lamina 15- 

22 x 6-7.5 cm; pedicels thin, 8-12 mm long. 

Habitat. Variety hispida is a small shrub of 

1.5 m of the understory of forest on terra firme. 


GUIANA. Haut Crique Baboune, 2 Aug 1981 (fl), de 

Granville 4723 (NY). 

BRAZIL. AMAZONAS: Mun. de Axinim, lower Rio 

Paca, 4?07'S, 58?58'W, 1 Jul 1983 (fl), Zarucchi et al. 

2920 (INPA, NY). AMAPA: Rio Oiapoque, Mt. Alikene, 

nr. Riv. Camopi, 30 Sep 1960 (fl), Irwin 48589a (NY). 

This variety was described to accommodate 

specimens of Hirtella racemosa that are distinctly 

hispid on the young stems. It has the large 

leaves characteristic of var. racemosa but the 

long thin pedicels characteristic of var. hexandra. 

The latter character is almost constant to separate 

the two varieties, although a few collections 

placed in var. racemosa have long thin pedicels 

(e.g., Chagas 1252, INPA, NY). The above material 

is obviously closest to H. racemosa and is 

best regarded as a variety until further collections 

and field studies are made. 

7-58. Hirtella kuhlmannii Pilger, Notizbl. Bot. 

Gart. Berlin-Dahlem 8: 538. 1923. Fig. 135. 

This species was known in the monograph from 

the type and one other collection from southern 


HEREDIA: Rio Puerto Viejo, 2 km above Rio Sarapiqui, 

14 Jun 1968 (fl), Burger & Stolze 5784 (F); Finca La 

Selva, Puerto Viejo, 17 Sep 1969 (fl), Frankie 398C 

(F), 7 Mar 1970 (fl), Hartshorn 809 (NY), Sep 1974 

(fl), Hartshorn 1523 (NY), 13 Aug 1961 (fl), Rosbach 

3719 (GH). PUNTARENAS: Ridge between Banegas and 

Rio Riyito, 7 km W of Rinc6n de Osa, 8 Oct 1984 (fr), 

Grayum 4090 (MO, NY); Llorona trail to San Pedrillo, 

Corcovado National Park, 21 Jul 1977 (fr), Hartshorn 

1878 (NY). 

7-67. Hirtella mucronata Prance, Fl. Neotrop. 

Monogr. 9: 339. 1972. 

Distribution (Fig. 139). This species was based 

on a type from southern Guyana and two col- 

lections from the Rio Negro region of Brazil. 

New material adds to this range. 

Additional specimens examined. SURINAME. Lely 

Mountains, 175 km SSE of Paramaribo, 12 Oct 1976 

(fl), Mori & Bolten 8459 (NY). 

BRAZIL. AMAZONAS: Manaus-Itacoatiara Rd., km 

185, 15 Dec 1966 (fl), Prance et al. 3653 (MG); Manaus-Caracarai 

Rd., km 220, Santo Ant6nio de Abonari, 

24 Nov 1976 (fl), Prance et al. 24257 (INPA, 

NY). 

7-70.1. Hirtella parviunguis Prance, Revista 

Brasil. Bot. 2: 34, fig. 4. 1979. Type. Brazil. 

Espirito Santo: Linhares, estrada da Povoa-ao 

ao Linhares, 30 Mar 1971 (fl), T. S. dos Santos 

1512 (holotype, CEPEC; isotypes, FHO, NY).



appressed puberulous, glabrescent, lenticellate. 

Leaf lamina narrowly oblong, coriaceous, 4-7 x 

1.8-2.5 cm, cuneate to subcuneate at base, acuminate 

at apex, with acumen 2-5 mm long, glabrous 


on both surfaces; midrib prominulous 

above, prominent beneath, venation papillose on 

upper surface; petioles 3-4 mm long, terete, 

eglandular, sparsely puberulous when young, rugulose. 

Stipules lanceolate 2-4 mm long, puberulous 

when young, subpersistent, adnate to extreme 

base of petiole, eglandular. Inflorescences 

of terminal and axillary densely crowded racemes 

3-5.5 cm long, the rachis sparsely tomentellous. 

Bracts and bracteoles ovate, 1.5-2.5 mm 

long, persistent membranous, eglandular. Flowers 

ca. 7 mm long. Receptacle campanulate, 

sparsely puberulous on exterior, glabrous within 

except for deflexed hairs at throat; pedicels 3-6 

mm long. Calyx lobes five, acute, sparsely puberulous 

on exterior, tomentellous within, the 

margins eglandular. Petals five, glabrous, with 

minute claw at base. Stamens seven, unilateral, 

the filaments glabrous, far exceeding the calyx 

lobes. Style pilose, with a few hairs on lower half. 

Ovary inserted at mouth of receptacle, pilose. 


Distribution (Fig. 141). Atlantic coastal forest 

and cacao plantations. 

7-75. Hirtella excelsa Standley ex Prance, Fl. 

Neotrop. Monogr. 9: 345. 1972. Fig. 130. 

Additional specimens examined. PERU. HUANUCO: 

Prov. Pachitea, Bosque Nacional de Iparia, Rio Pachi- 

tea, 8 Aug 1967 (fr), Schunke V. 2137 (NY); Dist. 

Honoria, Carretera Miel de Abejas, km 1.5, 8 Aug 1967 

(fr), Schunke V. 10 (F). SAN MARTIN: Santa Margarita, 

W of Nueva Aspusana, 6 Aug 1962 (fr), Mathias & 

Taylor 6108 (F). 

BRAZIL. ACRE: Between Porangaba and Papagaio, 

Rio Jurua-Mirim, 18 May 1981 (fl), Maas et al. P13134 

(INPA, NY). MATO GROSSO: Rio Tucunazinho, BR 174, 

km 330, 8 Jun 1979 (fr), M. G. Silva & Rosdrio 4826 

(MG), 10 Jun 1979 (fr), M. G. Silva & Rosdrio 4845 

(MG, NY). 

Local name. Peru: apacharama amarillo. 

7-79. Hirtella couepiiflora Prance, Fl. Neotrop. 

Monogr. 9: 350. 1972. Fig. 127. 


GUIANA. Riv. Oyapock, Petit Toucouchi, Lagon Trois 

Pitons, 6 Aug 1969 (fl), Oldeman T447 (CAY). 

BRAZIL. AMAPA: Clevelandia, Aug 1960 (st), Pires 

7700 (IAN). 

Local name. Fr. Guiana: gaulette. 

7-80. Hirtella tubiflora Cuatrecasas, Fieldiana, 

Bot. 27: 59. 1950. 

Distribution (Fig. 150). This species was known 

only from the type collection from the coastal 

lowlands of Valle in Colombia at 30-50 m, in 

the Rio Calima region. Recently, four collections 

Additional specimen examined. BRAZIL. ESPiRITO 

SANTO: CVRD Linhares, 10 Jul 1979 (fl), Foli 80 have been made from the wet forests of Panama 

(INPA). 

at about 800 m altitude, showing an interesting 

Local name. macucurana. 

distribution pattern. 

Hirtella parviunguis is distinct from all related 

species by the small slightly clawed petals. It is 

most closely related to H. martiana, a Planalto 

species, but, it differs not only in the petals but 

also in the eglandular bracteoles and calyx lobes, 

and the less pubescent inflorescence and style. It 

differs from H. angustifolia, another eastern Brazilian 

species, in the inflorescences of densely 

Additional specimens examined. PANAMA. PANAMA: 

Cerro Jefe, 26 Sep 1975 (fl), J. T. & F. Witherspoon 

8543 (MO, NY), 15 Feb 1982 (fl), Knapp 3525 (MO, 

NY). SAN BLAS: El Llano-Carti Rd., km 19.1, 11 Mar 

1985 (fl), Nevers & Herrera 5106 (MO, NY). VERAGUAS: 

3-4 km W of Santa F6, 2500' (fl), Nee 11288 (MO, 

NY). 

COLOMBIA. CHOCO: Colombia-Panama frontier, 

24 Sep 1979 (fl), Barbosa 1213 (COL). 

crowded short racemes, the subcuneate leaf bases, 

and the puberulous (not hispid), young 

branches and inflorescences. This species also 

resembles H. glaziovii from the same region, but 

differs in the petals, the greater number of stamens, 

the much less tomentose inflorescence and 

flowers, the larger flowers, and the more crowded 

inflorescences. 

7-87. Hirtella pauciflora Little, J. Wash. Acad. 

Sci. 38: 88. 1948. Fig. 141. 

Additional specimens examined. ECUADOR. LOS 

RIOS: Jauneche forest, Canton Vinces, 1 Oct 1979 (fl), 

Dodson et al. 8655 (NY); Rio Palenque Biological Station, 

km 56, Quevedo to Santo Domingo, 6 Mar 1974 

(fl), Dodson 5464 (US). 

Local name. coquito.



of Species of Acioa 

Distribution of Acioa as a whole is seen in 

Figure 153. 

8-1. Acioa guianensis Aublet, Hist. pl. Guiane 

2: 698, t. 280. 1775. 

Acioa remains the most poorly known genus 

of neotropical Chrysobalanaceae. Very few col- 

lections have been made of this apparently very 

rare species. 

A recent collection of A. guianensis from the 

Municipality of Manicore on the Madeira River 

is the first reliable collection from Central Ama- 

zonia and confirms the likelihood that seeds col- 

lected by Ducke and grown in the Botanical Gar- 

den of Rio de Janeiro are really from the Rio 

Purus region. All previous collections of this 

species had been made in French Guiana and 

adjacent Amapa. See Figure 154. 

8. Acioa Aublet 

Tree 25 m tall, the trunk slightly buttressed to 

0.5 m, the young branches glabrous. Leaf lamina 

oblong, thickly coriaceous, 7-17 x 4.5-12 cm, 

rounded at base, acuminate at apex, the acumen 

2-6 mm long, glabrous on both surfaces, with 

two 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. Stipules linear, membranous, 5-8 mm 

long, glabrous, caducous. Inflorescences muchbranched, 

slightly corymbose panicles, 5-10 cm 

long, the rachis and branches glabrous. Receptacle 

conical and slightly curved near base, 6-7 

mm long, glabrous on exterior, the interior lined 

by an extremely thick disc with only a small 

hollow, the interior with pilose hairs below insertion 

of style, glabrous on other side. Calyx 

lobes five, rounded, unequal, 3-5 mm long, glabrous 

with two glands on exterior, the margins 

ciliate, appressed-puberulous on interior. Petals 

five, white, caducous, glabrous, the margins cil- 

Additional specimen examined. BRAZIL. AMAZONAS: iate. Stamens 

Mun. of Manicore: RADAM SB-20-XA, Point 

17-20, inserted in two rows around 

9, 

60?53'W, 6?50'S (fl), C. D. A. Mota s.n. half of the thick staminal 

(INPA 61659). 

ring, the other half of 

the ring with staminodes, with a circle ofdeflexed 

8-3. Acioa schultesii Maguire, Brittonia 7: 272. hairs inserted on interior at base of staminal ring, 

1951. 

the filaments glabrous, the anthers dorsifixed. 

Ovary inserted at mouth of receptacle on the 

This species was known until recently only from same side as stamens, glabrous on exterior, glathe 

type collection from the Rio Dimite in Brazil. brous within, unilocular; ovules two. Style in- 

It has now been collected from the region of San serted at base of ovary towards interior, the swol- 

Carlos de Rio Negro in Venezuela, where it is len base pilose, the filamentous portion glabrous, 

apparently quite common, occurring frequently equalling stamens in length. Fruit ellipsoid, 6- 

in the IVIC ecological study sites in the region 7.5 cm long, 4-5 cm broad, exocarp glabrous, 

(see Fig. 154). It is a tree 20-40 m tall, flowering lenticellate; mesocarp 12-14 mm thick; endoin 

June-August and known locally as pasista. carp thin and bony, fragile, glabrous within, the 

cotyledons completely filling the central cavity. 

Additional specimens examined. VENEZUELA. Germination 

AMAZONAS: 4.3 km NE of San Carlos de Rio 

epigeal, first leaves opposite. 

Negro, 

1?56'N, 67?3'W, 19 Jun 1978 (fl), Clark & 

Distribution. 

Maquirino 

Mainly in the basin of the Rio 

6661 (FHO, NY), 20 Jul 1978 (fl), Clark & Maquirino 

Purus and the central Solimoes. 

6737 (FHO, NY), 3 Aug 1978 (fl), Clark & Maquirino Habitat. Forest on terra firme. 

6740 (FHO, NY), 25 Jul 1978 (fl), Clark & Maquirino 

6753 (fl) (NY), 25 Aug 1981 (fl), Clark 8211(NY), 5 Additional 

km N 

specimens examined. BRAZIL. 

of Boca de Casiquiare, 1?57'N, 67008'W, 5 Feb AMAZONAS: Coari, 11 

1980 (fr), Liesner & Clark 9107 

May 1974 (fr), CampbellP21130 

(MO, NY). 

(INPA, NY); 10 Feb 1975 (fl), Ramos P23251 (FHO, 

INPA, MG, NY); Lago de Tefe, 9 May 1974 (st), 

8-4. Acioa edulis Prance, Acta Amaz6nica 

Campbell P21129 (INPA, NY), 20 Nov 1959 (fr), Ro- 

2(1): 

drigues & L. Coelho 1408 (INPA); Rio Ituxi, Boca do 

12-16. 1971. Couepia edulis (Prance) Prance, Remanzinho, 1933 (fl bud), Krukoff5822A (NY); Ma- 

Acta Amazonica 5: 143. 1975. 

nua, Lago de Tefe, 13 Jul 1973 (fr), Lleras et al. P16655


(INPA, NY); Santo Ant6nio de Iga, 19 Aug 1973 (fr), 

Lleras et al. P17397 (INPA, NY). 

Local name. castanha de cotia. 

Uses. The fleshy cotyledons are edible and the 

fruits are gathered in large numbers by Brazilians 

in the Rio Ituxi region. The kernel is eaten raw 

or is crushed and added to their tapioca cakes 

(beiju). Its oil is also extracted and used for cooking 

and soap making. 

The fact that I have already placed this most 

distinctive new species in two genera indicates 

9-1. Neocarya macrophylla (Sabine) Prance in 

F. White, Bull. Jard. Bot. Etat. 46: 308. 1976. 

The eight genera treated above are those in- 

digenous to the neotropics. The monotypic West 

African genus Neocarya is grown at the Summit 

Botanical Garden in Panama and so it is men- 

tioned briefly here. Neocarya is described in full 

in Prance and White (1988), and is a segregate 

ACKNOWLEDGMENTS 

its problematic nature. It is quite unlike any others. 

It resembles Acioa vegetatively and in many 

floral features, but does not have what was previously 

considered to be the central, uniting character 

of the genus: the stamens fused into a ligule. 

A recent review of the genera (Prance & White, 

1988) showed that it is best placed inAcioa rather 

than Couepia even though it does not have the 

fused stamens. This species forms a link with 

Couepia, a genus in which fused stamens never 

occur. 

9. Neocarya Prance 

I am especially grateful to Mrs. Edith Topfler 

for the preparation of most of the distribution 

maps and for many hours of painstaking vol- 

unteer work for this project. I also particularly 

thank Carol Gracie for the preparation of the 

final version of the maps, and Bobbi Angell for 

drawings of the new species. I am grateful to 

Rosemary Lawlor and Mickey Maroncelli for the 

typing and word processing of various drafts of 

this manuscript and to H. David Hammond, 

William R. Anderson, and two anonymous re- 

viewers for reviewing an earlier draft. Most of 

from the closely related genus Parinari. It differs 

in the large gibbous receptacle, the greater num- 

ber of stamens and the very different fruit struc- 

ture. 

Specimen examined. PANAMA. CANAL AREA: Sum- 

mit Garden, 6 May 1986 (fl), de Nevers et al. 7726 

(MO, NY). 

LITERATURE CITED 

Berlin, B. & G. T. Prance. 1978. Insect galls and 

human ornamentation. The ethnobotanical significance 

of a new species of Licania from Amazonas, 

Peru. Biotropica 10: 81-86. 

Demchenko, N. I. 1973. The pollen morphology of 

the family Chrysobalanaceae. Pages 69-73 in Pollen 

and spore morphology of recent plants (in Russian). 

Proc. 3rd Int. Palynol. Conf., Acad. Sci. 

USSR. 

Espinal T., S. 1981. El frbol raro de Comfama en 

Rionegro. Flora Antioquenia 1: 1-3. 

Goulding, M. 1980. The fishes and the forest: Explorations 

in Amazonian natural history. University 

of California Press, Berkeley. 

my fieldwork, which has been essential for an Letouzey, R. & F. White. 1976. Chrysobalanacees 

understanding for the family, has been supported nouvelles du Cameroun et du Gabon. Adansonia, 

by a series of grants from the National Science Ser. 2, 16: 229-243. 

-- 

Foundation, lately by grant BSR-8409536 which 

& . 1978a. Chrysobalanacees. In Flore 

du Cameroun 20: 1-128, 237-247. Museum Nais 

gratefully acknowledged. I am grateful to col- tional d'Histoire Naturelle, Paris. 

laborators in many neotropical herbaria and bo- & . 1978b. Chrysobalanacees. In Flore 

tanical institutions, especially to the directors and du Gabon 24: 138, 194-201. Museum National 

staffs of the Instituto Nacional de Pesquisas da d'Histoire Naturelle, Paris. 

Amaz6nia (INPA) in Manaus, Brazil, and of the Mori, S. A., B. M. Boom & G. T. Prance. 1981. Distribution 

patterns and conservation of eastern Bra- 

Museu Paraense Emilio Goeldi (MG) in Belem, zilian coast forest tree species. Brittonia 33: 233- 

Brazil. I am especially grateful to David Johnson 245. 

for much editorial assistance. 

Patel, V., J. J. Skvarla & P. H. Raven. 1983. Pollen


ultrastructure of Chrysobalanaceae. Vidya 26: 1- tropical species with relation to history, dispersal 

10. 

and ecology, with special reference to Chrysobala- 

Prance, G. T. 1968. Maranthes (Chrysobalanaceae), naceae, Caryocaraceae and Lecythidaceae. Pages 

a new generic record for America. Brittonia 20: 59-87 in K. Larsen & L. B. Holm-Nielsen (eds.), 

203-204. 

Tropical botany. Academic Press, New York. 

. 1970. The genera of Chrysobalanaceae in the . 1981. Notes on Couepia and Hirtella (Chryssoutheastern 

United States. J. Arold Arbor. 51: obalanaceae). Brittonia 33: 347-356. 

521-528. 

. 1982a. Forest refuges: Evidences from woody 

. 1972. Monograph of Chrysobalanaceae. Fl. angiosperms. Pages 137-151 in G. T. Prance (ed.), 

Neotrop. Monogr. 9: 1-406. 

Biological diversification in the tropics. Columbia 

. 1973. New and interesting Chrysobalanaceae University Press, New York. 

from Amazonia. Acta Amaz6nica 2(1): 7-16. 

. 1982b. Chrysobalanaceae. Flora de Vene- 

. 1974a. Supplementary studies of American zuela 14(2): 325-487. 

Chrysobalanaceae. Acta Amaz6nica 4(1): 17-23. 

1984. New taxa of Amazonian Chrysobala- 

1974b. A new Peruvian species of chirop- naceae. Acta Amaz6nica 13: 21-30. 

terophilous Couepia (Chrysobalanaceae). Britto- 1986a. Studies on the flora of the Guianas. 

nia 26: 302-304. 

19: New taxa of Chrysobalanaceae for the flora of 

1974c. A note on Couepia cognata (Steud.) the Guianas. Proc. Kon. Ned. Akad. Wetensch., 

Fritsch and related species (Chrysobalanaceae). Ser. C. 89: 111-116. 

Acta Bot. Venez. 9: 119-122. 

. 986b. Flora of the Guianas Series A: Pha- 

1974d. Phytogeographic support for the the- nerogams. Family 85. Chrysobalanaceae. Koeltz 

ory of Pleistocene forest refuges in the Amazon Scientific Books, Koenigstein, Federal Republic of 

Basin based on evidence for distribution patterns Germany. 

in Caryocaraceae, Chrysobalanaceae, Dichapeta- . 1987. Notulae de Chrysobalanaceis Malelaceae 

and Lecythidaceae. Acta Amaz6nica 3(3): sianis Praecursoriae. Brittonia 39: 364-370. 

5-28. 

. In press. Chrysobalanaceae. In C. Kalkman 

1975. The correct name for castanha de cutia (ed.), Flora Malesiana. 

(Couepia edulis (Prance) Prance-Chrysobalana- - & A. R. A. Gorts van Rijn. 1976. Chrysobalceae). 

Acta Amaz6nica 5(2): 39-41. 

anaceae. Pages 524-555 in J. Lanjouw & A. L. 

. 1976. Additions to neotropical Chrysobal- Stoffers. Additions and corrections to the Flora of 

anaceae. Brittonia 28: 209-230. 

Suriname 2(2). 

.1977a. Two new species for the flora of Pan- -& S. A. Mori. 1983. Dispersal and distribuama. 

Brittonia 29: 154-158. 

tion of Lecythidaceae and Chrysobalanaceae. Son- 

. 977b. The phytogeographic subdivisions of derb. Naturwiss. Ver. Hamburg 7: 163-186. 

Amazonia and their influence on the selection of & F. White. 1979. Resurrection of the genus 

biological reserves. Pages 195-213 in G. T. Prance Dactyladenia (Chrysobalanaceae). Brittonia 31: 

& T. S. Elias (eds.), Extinction is forever. The New 483-487. 

York Botanical Garden, New York. 

& . 1988. A generic monograph of the 

.1979a. New genera and species of Chryso- Chrysobalanaceae and its relevance to practical 

balanaceae from Malesia and Oceania. Brittonia and theoretical taxonomy and evolutionary biol- 

31: 79-95. 

ogy. Phil. Trans. Roy. Soc. B. 320: 1-184. 

. 1979b. Two new species of neotropical Spichiger, R. & D. Masson. 1984. The Chrysobalana- 

Chrysobalanaceae. Brittonia 31: 248-252. 

ceae of the Arboretum Jenaro Herrera. 5th Con- 

.1979c. New and interesting species of Chrys- tribution to the study of the flora and vegetation 

obalanaceae. Acta Amaz6nica 8: 577-589. 

of the Peruvian Amazon. Candollea 39: 13-44. 

. 1979d. Chrysobalanaceae. In G. Harling & Tobe, H. & P. H. Raven. 1984. An embryological 

B. Sparre (eds.), Flora of Ecuador 10: 1-24. Stock- contribution to systematics of the Chrysobalanaholm, 

Sweden. 

ceae I. Tribe Chrysobalaneae. Bot. Mag. (Tokyo) 

1979e. The taxonomy and phytogeography 97:397-411. 

of the Chrysobalanaceae of the Atlantic coastal White, F. 1976. The taxonomy, ecology and chorolforests 

of Brazil. Revis. Brasil. Bot. 2: 19-39. ogy of the African Chrysobalanaceae (excluding 

1979f. Distribution patterns of lowland neo- Acioa). Bull. Jard. Bot. Etat. 46: 265-350.

Numerical List of Taxa 99 

Numerical List of Taxa, Collecting History, Status of Endangerment and Figure 

Numbers. Taxa in boldface described since 1972 

Map Number of Exsiccatae 

Fig. 

No. 1972 May 1987 Status' 

1. Chrysobalanus 

1. 

2. 

3. 

C. icaco L. ............. ................................................... 20 

C. cuspidatus Griseb. ex. Duss............................................ . 21 

C. venezuelanus Prance .....................................................21 

>10 

>10 

0 

217 

3 

5 R 

2. Licania 

1. L. michauxii Prance .......-............ ..... ..... 61 >10 

2. L. boliviensis Prance ................................. ....... ... 28 3 

3. L. m aritim a Prance .................................................................... 59 3 

4. L. durifolia Cuatrec. ............-....................... ....... 

36 4 

4.1. L. filomenoi Prance ............................................ . 38 - 

4.2. L. grandibracteata Prance ................................................. 38 - 

5. L. macrocarpa Cuatrec ......................... .......... .........42 2 

5.1. L. gentryi Prance ........................................ ....................40 - 

5.2. L. cabrerae Prance .............-...................... 30 - 

5.3. L. fasciculata Prance ..........................-.......... 38 - 

5.4. L. montana Prance .................................................... 65 -1 

6. L. salzmannii (Hook. f.) Fritsch .- ............................. 77 5 

7. L. klugii Prance .............................................. 

-51 1 

8. L. guianensis (Aubl.) Griseb .....................-............. .. 43 >10 

9. L. retifolia Blake .... ................. ..........................73 ... 

2 

10. L. longipedicellata Ducke ..........................-........... 57 2 

11. L. tomentosa (Benth.) Fritsch ....................-................ 82 >10 

12. L. pyrifolia Griseb ............. -73 ............................ . 

>10 

13. L. leucosepala Griseb. .......................................................55 10 

13.1. L. dodsonii Prance ........................................ ..................155 - 

14. L. angustata Prance .............................................. . 23 2 

14.1. L. anneae Prance ........................................... . 23 - 

15. L. platypus (Hemsl.) Fritsch ............-....................... 72 >10 

16. L. gonzalezii Miranda .................-......................... 42 2 

17. L. egleri Prance ...................-............... ... 37 >10 

18. L. minutiflora (Sagot) Fritsch ...................-................... 64 >10 

18.1. L. chiriquiensis Prance .......................................... . 33 - 

18.2. L. kallunkiae Prance .....................-.........-.. ......................51 - 

18.3. L. guatemalensis Lundell .........................-................... 43 - 

19. L. maranhensis Prance ...................................................... 60 1 

20. L. fritschii Prance ............................................... . 39 4 

21. L. britteniana Fritsch ....-...................... ... 

.. 29 >10 

21.1. L. cecidiophora Prance ..................................................30 - 

22. L. unguiculata Prance .-......................... ....... . 

83 8 

23. L. longipetala Prance ....................-.......-.....-..-- . 57 3 

23.1. L. tachirensis Prance .....................-........................ . 81 - 

24. L. wurdackii Prance .........................-........... .. 85 - 

25. L. turbinata Benth. ....................................... ...................84 9 

26. L. lata Macbr .................... ............................. 53 >10 

46 

0 

1 

6 

1 

4 

10 

1 

1 

3 

3 

2 

14 

1 

2 

11 

24 

14 

2 

0 

1 

31 

2 

45 

16 

2 

2 

2 

0 

0 

15 

4 

15 

7 

1 

12 

2 

21 

R 

R 

R 

R 

R 

R 

R, E 

R 

R, E 

R 

R 

R 

R 

R 

R, E 

R 

R 

27. L. apetala (E. Mey.) Fritsch 

a. var. apetala ..--.........................--....... 24 

b. var. aperta (Benth.) Prance .......................-............ 24 

27.1. L. granvillei Prance ....................................................... . 42 

28. L. parvifolia Huber ............-.............................. 71 

29. L. maguirei Prance .......................................... 59 

30. L. gardneri (Hook. f.) Fritsch ....................................... 40 

31. L. cuspidata (Rusby) Prance ....................................... 34 

31.1. L. jefensis Prance .......................................-...... . 50 

31.2. L. morii Prance ..............-........-............... 65 

>10 

>10 

-13 

-45 

3 

>10 

1 

- 

- 

132 

65 

0 

22 

0 

6 

1 

R 

R, E 

R


Continued 

Map 

Fig. 

No. 

Number of Exsiccatae 

1972 May 1987 Status' 

32. L. sparsipilis Blake ................................................................................. 

79 7 

32.1. L. cuatrecasasii Prance ....................................................................... 34- 

6 

2 

32.2. L. mexicana Lundell ............................................................................. 

60- 1 R 

33. L. emarginata Hook. f. ...................................................................... 389 2 

33.1. L. joseramosii Prance .............................. ................ .. .............. ... 51- 1 

34. L. calvescens Cuatrec. ..-.--.............-....-...... ......... 30 2 2 

35. L. persaudii Fanshawe & Maguire . ............................................ 71 9 1 

36. L. sprucei (Hook. f.) Fritsch . ............. ..... 79 >10 21 

37. L. sclerophylla (Mart. ex Hook. f.) Fritsch ......................... 78 >10 30 

38. L. albiflora Fanshawe & Maguire .................................... 23 2 0 

39. L. longistyla (Hook. f.) Fritsch . .................................. 58 >10 47 

40. L. fuchsii Prance ....................................... ....................................... 39 1 0 

41. L. humilis Cham. & Schlecht. ............................ ... 47 >10 34 

42. L. foveolata Prance ............................................................................ 39 2 0 

43. L. octandra (Hoffmgg. ex Roem. & Schult.) Kuntze 

a. subsp. octandra ................................................................................. 

67 >10 63 

b. subsp. pallida (Hook. f.) Prance ........................................... 68 >10 76 

c. subsp. grandifolia Prance ......................................................... 68 0 8 

44. L. rigida Benth. ....................................................................................... 

74 >10 9 

45. L. arborea Seem. ..................................................................................... 

26 >10 60 

46. L. velata Cuatrec . .................................................................................... 

85 1 3 

47. L. subarachnophylla Cuatrec. ........................................................ 80 2 2 

47.1. L. tambopatensis Prance .................................................................... 62- 1 

48. L. salicifolia Cuatrec ........................................................................... 

76 1 4 

49. L. araneosa Taub. ....................... ................................................ 25 4 0 

50. L. silvatica Glaziou ex Prance . ..................................................... 

79 1 0 

51. L. chocoensis Cuatrec. ......................................................................... 33 3 4 

52. L. licaniiflora (Sagot) Blake . ........................................ 56 >10 27 

53. L. hirsuta Prance . ................................... .. 46 2 7 

54. L. costaricensis Standl. & Steyerm . .............................. 32 1 0 

55. L. krukovii Standl. .................. ............................................................... 

51 4 4 

56. L. lasseri Maguire ....................... ............................. ......... 54 8 14 

57. L. latifolia Benth. ex Hook. f. ....................................................... 54 >10 31 

57.1. L. hispida Prance .................................................................................... 

46 -1 

58. L. minuscula Cuatrec .......................................................... 63 1 1 

59. L. operculipetala Standl. & L. 0. Wms. 68 2 0 

60. L. reticulata Prance ............................................................................... 

75 >10 21 

R 

R 

R 

R 

R 

R 

R 

R 

R, E 

R, E 

R 

R 

R, E 

60.1. L. pakaraimensis Prance ............................................................... 70- 

61. L. arachnoidea Fanshawe & Maguire . ...................-- ...... 25 2 

62. L. oblongifolia Standl ....i.... .......................................... 66 >10 

63. L. macrophylla Benth. .................... ................................. 58 >10 

64. L. caudata Prance .................................................................................. 

31 3 

5 

1 

20 

27 

24 

R 

64.1. L. miltonii Prance ................................................................................ 64- 

65. L. latistipula Prance .............................................................................. 

54 4 

66. L. divaricata Benth. .............................................................................. 

36 >10 

2 

2 

5 

67. L. glabrifora Prance ............................................................................. 

41 5 

68. L. intrapetiolaris Spruce ex Hook. f. ........................................ 50 8 

69. L. heteromorpha Benth. 

a. var. heteromorpha ........................................................................ 44 >10 

b. var. glabra (Mart. ex Hook. f.) Prance ........................... 45 >10 

c. var. subcordata Fritsch ................................................................ 45 4 

d. var. perplexans Sandw. ............................................................... 45 >10 

e. var. revoluta Prance .......................................................................- 

69.1. L. laevigata Prance -..............................-......... - 

62-9 

69.2. L. occultans Prance .......................................................................... 62- 

70. L. glazioviana Warm .......................................................................... 

41 2 

4 

31 

213 

93 

1 

0 

1 

2 

1 

R 

R


Continued 

Map 

Fig. 

No. 


1972 May 1987 Status' 

71. L. littoralis Warm. 

a. var. littoralis .................................................................................... 

57 8 

b. var. cuneata Kuhlm ..................................................................... 

72. L. fanshawei Prance ............................................................................. 

38 5 

73. L. irwinii Prance ...................................................................................... 

503 

73.1. L. marleneae Prance .............................................................................. 

60- 

74. L. cyathodes R. Ben .................................................... 35 4 

75. L. polita Spruce ex Hook. f. ..................-............................ 72 >10 

76. L. silvae Prance ........................................................................................ 

78 3 

77. L. densiflora Kleinh .............................................................................. 

36 >10 

78. L. cuprea Sandw .................................................................................... 

34 8 

78.1. L. arianeae ................................................................................................... 

27 -2 

79. L. impressa Prance ............................................................................... 

49 4 

80. L. dealbata Hook. f ........................................................................... 35 >10 

80.1. L. santosii Prance ........................................................................... 77- 

81. L. pallida Spruce ex Sagot ............................................................... 69 >10 

82. L. gracilipes Taub ................................................................................. 

42 >10 

83. L. parvifructa Fanshawe & M aguire .. .................................... 71 7 

84. L. cymosa Fritsch -............. - .............. .......... 35 3 

85. L. ternatensis Hook. f. ex Duss ..-............... ............ 82 >10 

86. L. membranacea Sagot ex Lanes .............................................. 61 >10 

87. L. piresii Prance ....................................................................................... 

73 3 

87.1. L.furfuracea Prance ............................................................................. 

40- 

88. L. hypoleuca Benth. 

a. var. hypoleuca .........-.............................. 48 >10 

b. var. foveolata Prance ................................................................2 

89. L. boyanii Tutin ..-........................... ......... 28 5 

90. L. buxifolia Sandw ............................................................................... 

29 3 

91. L. orbicularis Spruce ex Hook. f. ................................................ 69 7 

92. L. niloi Prance ...................................................................................... 

66 1 

93. L. coriacea Benth ................................................................................ 

32 >10 

94. L. urceolaris Hook. f. .......................................................................... 

84 8 

95. L. affinis Fritsch ....................................................................................... 

22 >10 

95.1. L. teixeirae Prance ................................................................................. 

81 -1 

96. L. glauca Cuatrec .................................................................................. 

41 3 

97. L. davillifolia Benoist ....................................... 35 >10 

98. L. elliptica Standl. .................................................................................. 

37 7 

99. L. canescens Benoist ............................................................................. 

31 >10 

100. L. couepiifolia Prance .......................................................................... 

32 1 

100.1. L. naviculistipula Prance ...................................... 65 2 

101. L. trigonioides M acbr. ....................................... 82 1 

102. L. cordata Prance ............................................................................... 33 5 

103. L. foldatsii Prance .................................................................................. 

39 8 

104. L. hebantha M art. ex Hook. f. ........................................ 46 3 

105. L. steyermarkii M aguire ..............-........................ 80 5 

106. L. subrotundata M aguire ................................................................ 81 7 

107. L. crassivenia Spruce ex Hook. f. ............................................... 32 1 

108. L. majuscula Sagot ................................................................................ 

59 >10 

108.1. L. jimenezii Prance ................................................................................ 

50 - 

109. L. alba (Bern.) Cuatrec ..................................................................... 22 >10 

110. L. hitchcockii M aguire ....................................................................... 46 2 

111. L. sandwithii Prance ......................................................................... 77 1 

112. L. laxiflora Fritsch .............................................................................. 55 >10 

113. L. rufescens Klotzsch ex Fritsch ................................... 76 >10 

114. L. kunthiana Hook. f. ......................................................................... 52 >10 

115. L. bellingtonii Prance ........................................................................... 

27 1 

6 

0 

3 

2 

1 

0 

13 

10 

35 

1 

6 

22 

4 

46 

8 

14 

5 

3 

20 

3 

1 

103 

0 

4 

4 

3 

1 

8 

17 

5 

2 

5 

9 

89 

1 

0 

0 

3 

3 

2 

4 

2 

8 

15 

4 

27 

0 

0 

11 

5 

60 

0 

R, E 

R 

R 

R 

R 

R 

R, E 

R, E 

R 

R 

R, E 

R, E 

R 

R 

R, E


Continued 


Fig. 


116. L. compacta Fritsch ....-.......................... ..... 33 1 

117. L. ovalifolia Kleinh ......................................................... . 69 9 

118. L. caldasiana Cuatrec. ......................................................30 3 

119. L. savannarum Prance .......................................................77 >10 

119.1. L. stewardii Prance ...........................................................80 - 

120. L. microphylla Fanshawe & Maguire ..................................... 61 1 

121. L. triandra Mart. ex Hook. f. ................................ 83 7 

121.1. L. tocantina Prance ............................................................................... 

82 - 

122. L. discolor Pilg............................................................... 36 >10 

123. L. apiculata Prance ............................................. . 25 3 

124. L. micrantha Miq ................................................................................. 

63 0 

125. L. pruinosa Benoist ............................................................................... 

73 5 

126. L. nitida Hook. f. .................. ................. ........... 66 9 

126.1. L. aracaensis Prance .................................................. 25 - 

127. L. riedelii Prance ..................................................................................... 

75 4 

128. L. bracteata Prance ......................................... .......................... 29 8 

129. L. parviflora Benth ......................................................... 70 >10 

130. L. robusta Sagot .................................................... ... 75 8 

130.1. L. lamentanda Prance ................................................ 53 - 

131. L. lanceolata Prance ............................................... 53 2 

132. L. spicata Hook. f. ........................................................... 79 >10 

133. L. stricta Kleinh ........................................................... 80 1 

134. L. leptostachya Benth ...................................................... . 56 >10 

135. L. incana Aubl. ..................................................................................... 

49 >10 

135.1. L. nelsonii Prance ...........................................................62 - 

136. L. paraensis Prance ...........................................................70 4 

137. L. vaupesiana Killip & Cuatrec. ....................................... 84 3 

138. L. bahiensis Prance ....................... ............ ...... ... 27 1 

139. L. maxima Prance .. .................. 60 1 

140. L. mollis Benth. ....................................................................................... 

65 >10 

141. L. blackii Prance .................................................. 28 >10 

142. L. rodriguesii Prance ........................................ . 76 8 

143. L. indurata Pilg .............................................. 49 2 

144. L. hoehneiPilg. ...............................................................47 >10 

144.1. L. harlingii Prance ...........................................................43 - 

145. L. cruegeriana Urb ......................................................... 34 >10 

146. L. belemii Prance .................................................................................. 

27 1 

147. L. splendens (Korth.) Prance ........................-............ 

.......-- Asiatic 

147.1. L. palawanensis Prance ................................................................. - Asiatic 

147.2. L. fusicarpa (Kosterm.) Prance ....................................... - Asiatic 

148. L. veneralensis Cuatrec. ........................................................... 85 1 

149. L. amapaensis Prance ...................................................... . 23 1 

150. L. tepuiensis Prance ..........................................................81 1 

151. L. obtusifolia Fritsch -1.....................................- 

152. L. roraimensis Standl ..................................................... . 76 1 

3. Parinari 

0 

3 

0 

21 

16 

0 

7 

3 

12 

0 

64 

0 

5 

4 

2 

8 

33 

3 

3 

23 

1 

0 

46 

21 

2 

6 

1 

1 

1 

43 

19 

6 

1 

15 

8 

10 

6 

11 

4 

0 

0 

0 

R 

R 

R 

R 

R, E 

E 

R 

R 

R, E 

R 

R 

R 

R 

R 

1. P. campestris Aubl. .................................. .................. . 87 >10 

2. P. montana Aubl ....................................... 90 >10 

3. P. rodolphii Huber .................................................. 92 >10 

3.1. P. alvimii Prance ..................................................... . 87- 

4. P. excelsa Sabine ................................................. .... 89 >10 

5. P. occidentalis Prance ........................................................91 4 

6. P. sprucei Hook. f. ................................................................................ 

92 7 

7. P. pachyphylla Rusby .......................................................................... 

91 >10 

8. P. brasiliensis (Schott) Hook. f. .......................................... 87 2 

9. P. klugii Prance ...............................................................88 1 

29 

14 

15 

2 

126 

8 

21 

35 

3 

11 

R


Continued 

Map 

Fig. 



10. 

11. 

P. maguirei Prance ........................................................... 

P. littoralis Prance ........................................................ 

90 

88 

2 

1 

3 

6 

12. 

13. 

14. 

15. 

P. parvifolia Sandw. .................................... .................. 91 

P. cardiophylla Ducke .................. ................. ................... 88 

P. parilis Macbr ...................................................................................... 91 

P. chocoensis Prance ................................................... ............. ......... ... 88 

4 

3 

2 

1 

1 

0 

12 

4 

R 

16. 

17. 

P. obtusifolia Hook. f. ....................-................ 

P. romeroi Prance ....................................................... 

90 

92 

>10 

2 

28 

3 

4. Exellodendron 

1. 

2. 

3. 

4. 

5. 

E. coriaceum (Benth.) Prance ............................................... 95 

E. cordatum (Hook. f.) Prance .............................................. 94 

E. barbatum (Ducke) Prance ........................................................ 94 

E. gardneri (Hook. f.) Prance .................................................. 95 

E. gracile (Kuhlm.) Prance .............................................................. 95 

>10 

5 

>10 

6 

1 

24 

18 

30 

10 

2 R, E 

5. Maranthes 

1. M. panamensis (Standl.) Prance & White 153 4 12 

6. Couepia 

1. 

(2.) 

C. guianensis Aubl. 

a. var. guianensis 1>.......10................................................ 105 >10 

b. var. glandulosa (Miq.) Prance ............................................... 105 >10 

c. var. divaricata (Hub.) Prance ............................................ 104 >10 

C. glandulosa Miq. 

66 

18 

14 

= 3. 

(4.) 

C. guianensis var. glandulosa .. - 

C. paraensis (Mart. & Zucc.) Benth. 

a. subsp. paraensis .............................................................................. 112 >10 

b. subsp. glaucescens (Spruce ex Hook. f.) Prance ....... 112 4 

c. subsp. cerradoana Prance ................................................ 112 4 

C. leptostachya Benth. ex Hook. f. = 

C. guianensis var. guianensis ............................................. 

36 

114 

4 

- 

5. 

6. 

C. maguirei Prance .................................... . 

....... 108 4 

C. sandwithii Prance 1.. 115 4 

1 

4 

6.1. C. bernardii Prance ........................................................ 97 - 14 

6.2. C. monteclarensis Prance .......2................................... - 2 

7. C. parillo DC ................................................................. 115 

8. C. steyermarkii Maguire ................................... ......... 116 

8.1. C. canescens (Gleason) Prance . ................. .................. 98 

9. C. canomensis (Mart.) Benth. ex Hook. f. .......................... 98 

10. C. foveolata Prance ......................................................... . 103 

11. C. magnoliifolia Benth. ex Hook. f. ........................................ 108 

12. C. exflexa Fanshawe & Maguire ........................................... 103 

13. C. habrantha Standl . ....................................................... 106 

>10 

1 

1 

>10 

7 

>10 

3 

>10 

32 

3 

21 

3 

21 

0 

7 

R 

13.1. C. scottmorii Prance .......................................................... 115 - I R, E 

13.2. C. carautae Prance ................................ ...... . 99 

14. C. spicata Ducke ................................................... 115 

15. C. bracteosa Benth. ........................... ................... . 98 

16. C. subcordata Benth. ex Hook. f. .................................... . 116 

17. C. belemii Prance .................................................... . 97 

- 

1 

>10 

>10 

3 

1 

7 

40 

11 

6 

R, E 

18. 

19. 

C. caryophylloides Benoist ................................................. . 

a. subsp. caryophylloides ............................... 

b. subsp. glabra Prance ....................................... ............ .- 

C. excelsa Ducke ...............................................102 

99 

>10 

4 

>10 

1 

2 

20. 

21. 

22. 

23. 

C. uiti (Mart. & Zucc.) Benth. ex Hook. f. ......................... 117 

C. cataractae Ducke ................................................ 99 

C. macrophylla Spruce ex Hook. f. ........................................ 108 

C. krukovii Standl ................................. ........ ............ .... 107 

>10 

8 

9 

2 

10 

16 

3 

2


Continued 

Map 


Fig. 


24. C. latifolia Standl. ............................................................... 107 

25. C. ovalifolia (Schott) Benth .......................................1..... 111 

26. C. schottii Fritsch ................................................................. . 115 

27. C. grandiflora (Mart. & Zucc.) Benth. ex Hook. f........ 104 

28. C. elata Ducke ..................................................... 102 

29. C. racemosa Benth. ex Hook. f. ............................. ........... . 113 

29.1. C. amaralae Prance ............................................... . . 97 

2 

>10 

>10 

>10 

>10 

- 

1 

23 

8 

27 

7 

34 

12 

30. C. martinii Prance .......................................................... . 109 

31. C. bondarii Prance ................................... 97 

32. C. insignis Fritsch ..........................................................106 

32.1. C. cidiana Prance .................................................. 99 

33. C. recurva Spruce ex Prance ................................................. 114 

34. C. obovata Ducke ............................................. .. 110 

35. C. williamsii M acbr ......................................................... . 118 

35.1. C. glabra Prance .......................................................103 

35.2. C. marleneae Prance .......................................... 108 

36. C. chrysocalyx (Poepp. & Endl.) Benth. ex Hook. f. ...100 

37. C. eriantha Spruce ex Hook. f............................................ 102 

38. C. trapezioana Cuatrec ......................... ............. 118 

39. C. stipularis Ducke ................................ ... 116 

40. C. reflexa Ducke ...................................................114 

41. C. longipendula Pilg .............................................. ... 107 

41.1. C. dolichopoda Prance ....................... 

. . 101 

42. C. cognata (Steud.) Fritsch 

a. var. cognata ............................................................101 

b. var. major Prance ................................... 101 

c. var. membranacea Prance .................................. .. . 101 

43. C. multiflora Benth. ......... ...................................... . 109 

44. C. uleiPilg ..................................................... 117 

45. C. comosa Benth .................. ........... . .. . 

100 

46. C. venosa Prance ...................................................... 118 

47. C. polyandra (Kunth) Rose ................................... 113 

47.1. C. nutans Prance ........................................... . 

109 

48. C. platycalyx Cuatrec ............................... ...... 111 

49. C. rufa Ducke ......................................................... 114 

50. C. robustaHuber .................................................114 

51. C. impressa Prance 

a. subsp. impressa ........................................ ................ . 106 

b. subsp. cabraliae Prance . ....................................... 100 

52. C. meridionalis Prance ..............................................109 

52.1. C. coarctata Prance ......................................... 100 

52.2. C. longipetiolata Prance .................................... 107 

53. C. pernambucensis Prance ................ .................... 111 

54. C. froesii Prance ........................................................... 103 

55. C. parvifolia Prance .........................................................111 

7. Hirtella 

1 

2 

5 

- 

1 

>10 

>10 

-2 

- 

>10 

>10 

4 

2 

1 

>10 

- 

>10 

3 

2 

>10 

>10 

>10 

9 

>10 

- 

2 

>10 

>10 

4 

- 

1 

- 

- 

1 

3 

2 

0 

0 

2 

2 

1 

15 

22 

8 

42 

5 

4 

0 

0 

12 

11 

3 

0 

0 

5 

52 

3 

1 

21 

1 

4 

1 

22 

3 

10 

0 

3 

2 

1 

0 

0 

R, E 

R, E 

R, E 

R 

R 

R 

R, E 

E 

R, E 

R, E 

R 

R, E 

1. H. myrmecophila Pilg .......................... ....... 139 

2. H. physophora Mart. & Zucc ............................................. . 142 

3. H. vesiculosa Suesseng. ................................................... . 152 

4. H. dorvalii Prance ............................... ..... 128 

5. H. guainiae Spruce ex Hook. f. ............................................. 132 

6. H. duckei Huber . ......................................... .................. . 128 

6.1. H. revillae Prance ............................................146 

7. H. macrosepala Sandw ........ ........................... 137 

8. H. ulei Pilg. .................................................... . 152 

9. H. glabrata Pilg ............................................................. 132 

>10 

>10 

2 

13 

>10 

>10 

- 

6 

10 

>10 

17 

51 

0 

35 

28 

2 

3 

20 

11 

R


Continued 


Fig. 


9.1. H. confertiflora Prance ........................................ 126 - 1 R 

10. H. carbonaria Little . ...................... ................ 125 >10 3 

11. H. araguariensis Prance ...................................................... 121 

11.1. H. barnebyi Prance .......................................................... . 122 

11.2. H. m argae Prance .................................................................................. 138 

11.3. H. liesneri Prance ............................................. 136 

1 

- 

- 

- 

4 

2 

3 

1 

12. 

13. 

14. 

H. cordifolia Prance . ................ ............................................................. 

126 2 

H. insignis Briq. ex Prance 15-----.......................................... 135 2 

H. tocantina Ducke 103-----------.. ...................................150 3 

0 

7 

3 

R 

15. H. piresii Prance ----....................................... 130-- 

143 0 8 

16. H. davisii Sandw ................................8... ........................1280 9 

17. 

18. 

H. subglanduligera Pilg ..................410................... 149 

H. ciliata Mart. & Zucc. ..................................................... 126 

1 

>10 

0 

45 

R, E 

19. 

20. 

21. 

H. hoehneiPilg. .............................................................134 

H. glandulosa Spreng. ..........................11 >0......... 131 

H. bullata Benth ............................................................124 

8 

>10 

>10 

6 

99 

54 

22. H. americanaL ............................................... . 120 

22.1. H. santosii Prance ........ ............................................... 147 

23. H. guatemalensis Standl ................................. 133 

24. H. eriandra Benth .................. ............................ ........... 129 

>10 

- 

>10 

>10 

65 

1 

17 

49 

R, E 

25. 

26. 

27. 

28. 

29. 

30. 

31. 

32. 

33. 

34. 

35. 

H. paniculata Sw ... ........................... 140 >10 

H. deflexa Maguire ...................................... .127 1 

H. tentaculata Poepp. & Endl .................................................... 150 >10 

H. macrophylla Benth. ex Hook. f. ..................................... 138 >10 

H. adderleyi Prance .... .......................................... . 

120 2 

H. punctillata Ducke ............................ ............. ... 143 >10 

H. corymbosa Cham. & Schlecht ........................................ 127 4 

H. barrosoi Prance .. ........... . 122 >10 

H. pendula Soland. ex Lam .................... ................... 141 2 

H. leonotis Pittier . . .... ......... 136 1 

H. mutisii Cuatrec ....................................... 139 8 

78 

4 

1 

8 

15 

18 

0 

0 

1 

7 

5 

R 

36. H. triandra Sw. 

37. 

a. subsp. triandra ................................. .... 151 

b. subsp. punctulata (Miq.) Prance ........................... 152 

c. subsp. media (Standl.) Prance ........................................... 152 

H. bahiensis Prance ............................................... 122 

>10 

>10 

>10 

3 

187 

8 

15 

3 

38. H. latifolia Prance .. .1.......................................136 . 

2 14 

39. 

40. 

40.1 

41. 

42. 

43. 

44. 

45. 

46. 

47. 

48. 

H. suffulta Prance . .................................................................................. 

149 6 

H. elongata Mart. & Zucc ................ .................... 129 >10 

H. magnifolia Prance . .......................................138 . 

- 

H. rodriguesii Prance . ......................................147 5 

H. obidensis Ducke ............................................... 140 >10 

H. cowanii Prance & Maguire ..-...-........................... . 127 5 

H. orbicularis Prance . ......................................140 3 

H. guyanensis (Fritsch) Sandw............................................ .... 133 8 

H. lightioides Rusby . ............................................................................ 

137 3 

H. aramangensis Prance .-. ...................... .. 121 1 

H. rasa Standl ........ ...................... 146 2 

15 

64 

6 

12 

14 

2 

0 

0 

8 

0 

1 

R 

R, E 

49. H. scabra Benth. .................................................... 148 >10 26 

50. H. bicornis Mart. & Zucc. 

a. var. bicornis ......................................................123 >10 29 

b. var. pubescens Ducke ............................................ . 123 >10 45 

51. H. angustissima Sandw ............................ ....... 121 8 

52. H. tenuifolia Prance .... . .. 150 3 

52.1. H. radamii Prance . ............................................................................... 

143 - 

1 

17 

1 

53. H. pilosissima Mart. & Zucc ............... ..................... 142 >10 22


Continued 


Fig. 


54. H. gracilipes (Hook. f.) Prance ............................................. . 143 

55. H. brachystachya Spruce ex Benth........................................ 125 

55.1. H. arenosa Prance ...................... ...... .. 122 

55.2. H. conduplicata Prance ............................................................. 125- 

56. H. racemosa Lam. 

>10 

>10 

- 

67 

10 

4 

2 

57. 

58. 

59. 

60. 

61. 

62. 

63. 

64. 

65. 

66. 

67. 

a. var. racemosa ........................................................ ... 145 >10 

b. var. hexandra (Willd. ex Roem. & Schult.) Prance 144 >10 

c. var. glandipedicellata Prance .................................................. 146 2 

d. var. hispida Prance ....................................................................... 146 - 

H. juruensis Pilg ................................................................................ 

135 3 

H. kuhlmannii Pilg ......................................................... .... 135 2 

H. standleyi Baehni & Macbr ..................... ......................... 149 2 

H. longifolia Benth. ex Hook. f. .-......................................... 137 1 

H. lemsii L. O. Wms. & Prance ........................................... 136 1 

H. schultesii Prance .........................-...-......................... 148 7 

H. paraensis Prance ..-.............-..-..................... . 141 >10 

H. sprucei Benth .................................................................................... 

148 >10 

H. lancifolia Ducke ...........................-................. .. 135 5 

H. burchellii Britton .....-....................-......................... 124 >10 

H. mucronata Prance ..................-............................. ..... 139 3 

282 

340 

0 

3 

1 

1 

7 

0 

10 

31 

10 

6 

2 

24 

3 

R, E 

R 

R 

R, E 

68. H. longipedicellata Prance .-.................................. ............... 137 6 

69. H. glandistipula Ducke ................ ................. 132 8 

70. H. martiana Hook. f. ......................................................................... 

139 >10 

70.1. H. parviunguis Prance ................. ................ ........... 141 - 

71. H. pimichina Lasser & Maguire .................................................. 143 4 

72. H. subscandens Spruce ex Hook. f .............................-.......... 149 4 

73. H. hispidula Miq ............................................................. 134 >10 

74. H. silicea Griseb .....................-............... . ....... ................ 148 >10 

75. H. excelsa Standl. ex Prance ............................ ....... 130 4 

5 

2 

11 

2 

1 

1 

78 

19 

10 

76. 

77. 

78. 

79. 

80. 

81. 

82. 

83. 

84. 

85. 

86. 

H. adenophora Cuatrec. ................. ............... ............ 120 3 

H. caduca Fanshawe & Maguire ................................................. 125 6 

H. fasciculata Prance . -................................................ .. 130 2 

H. couepiiflora Prance .........-............................................. 127 2 

H. tubiflora Cuatrec .............................. ........................... 150 1 

H. floribunda Cham. & Schlecht ......................................... 130 >10 

H. angustifolia Schott ....................................................... 121 >10 

H. rugosa Thuill. ex Pers ...........................-...................... 147 >10 

H. scaberula Spruce ex Hook. f. .................................-.......... 147 1 

H. hebeclada Moric. ex DC. .......................-..-............... . 134 >10 

H. enneandra Cuatrec ....................... .......-....... .. ...... ...... 130 1 

0 

1 

1 

2 

6 

0 

8 

10 

0 

11 

0 

R 

R 

R 

87. 

88. 

89. 

90. 

91. 

92. 

8. Acioa 

H. pauciflora Little .............................. .............. ........... 141 1 

H. glaziovii Taub ...................................-....................... 132 7 

H. megacarpa R. A. Grah. ............................. ........ .. - African 

H. zanzibarica Oliv ............................ ...........................African 

H. cliffortiana Veil .......................................................... . - I 

H. pohlii Hook. f. ......................... .....-..... 1 

2 

0 

0 

0 

R 

R 

1. 

2. 

3. 

4. 

A. guianensis Aubl .......................................................... . 154 

A. somnolens Maguire ...................................................... 154 

A. schultesii Maguire ......................................................... 154 

A. edulis Prance ..................................................... ........ 102 

7 

1 

2 

- 

1 

0 

10 

8 

R 

9. Neocarya 

1. N. macrophylla (Sabine) Prance ..............................-......... . - 1 

(African 

sp.) 

'R = rare; E = endangered.

Supplemental List of Exsiccatae 107 

SUPPLEMENTAL LIST OF EXSICCATAE 

This list includes material not cited in Prance (1972), largely recent collections made since that 

date and a few older ones not seen previously. 

Abadie, B., 50T (6-44) 

Acero, E. et al., 57 (7-50b); 195 (2-27b); 729 (2-121) 

Acevedo R., P. et al., 1657 (20-13.1) 

Ackerly, D. 0., 07 (7-2); 152 (7-50a); 161 (7-56b) 

Acosta M., A., 1 (2-17) 

Adair de Oliveira, INPA59936, INPA60162 (2-69.1); 

INPA73467 (6-41); INPA73471, INPA73472 (2-62); 

INPA73476, INPA73477 (2-69a); INPA73483 (2- 

64); INPA73484 (6-la); INPA74791 (2-69.1); INPA 

s.n. (2-69a) 

Adams, C. D. A., 14572 (1-1); YSB 86 (7-74) 

Agostini, G., 305 (2-77); 1104 (7-36a) 

Albert de Escobar, L. et al., 1414 (7-36a); 3282 (2-51); 

3525 (1-1) 

Albuquerque, B. W. P. de et al., 646 (7-56a); 671 (7- 

6); 672 (2-69b); 798 (2-81); 841 (2-36); 876 (7-2); 

886 (7-55.2); 67/20 (2-69.1); 887 (7-55.2); 919 (2- 

17); 950 (2-69b); 954 (2-81); 956 (2-69b); 998 (2- 

81); 1010 (2-39); 1048 (2-81) 

Alencar, L., 26 (7-56a); 89 (7-24); 112 (2-69b); 161 (7- 

56b); 238 (2-129); 249 (2-88); 251 (6-lb); 260 (2- 

88); 390 (7-5); 486, 499 (3-6); 570 (7-55); 616 (3- 

41); 725 (2-129) 

Allem, A. et al., 7, 2324, 2391 (6-20); 2489 (2-28) 

Allen, P. H., 3019 (2-68) 

Almeida, E. de F. et al., 275 (2-80.1); 283 (4-2) 

Almeida, J. et al., 150 (2-80.1); 164 (3-1.3); 185 (7- 

36a); 1308, 1791 (6-25) 

Aluisio de Sousa, J., INPA59853 (2-99); s.n. 

INPA61445 (7-1); INPA59853 (2-99); INPA61868 

(6-4); INPA61951 (7-6); 70666 (2-62); 70678 (6-11); 

70679 (7-1); 70682 (6-34); INPA-s.n. (2-43b) 

Amaral, D. L., 728 (7-20) 

Amaral, I. L. et al., 11 (6-3b); 39 (7-40); 40 (2-75); 43 

(2-129); 102 (2-27a); 56 (6-29); 131 (2-69a); 176 (7- 

73); 186 (2-124); 216 (2-75); 333 (2-114); 353 (3-4; 

390 (2-69b); 466 (2-26); 535 (2-129); 588 (2-26) 596 

(2-129); 602 (2-114); 703 (7-56a); 729 (7-40); 864 

(7-20); 909 (7-56a); 1006 (7-19); 1022 (7-50a); 1041 

(7-58); 1050 (7-36a); 1105(2-17); 1119(7-20); 1193 

(7-56b); 1271 (7-73); 1368 (2-43b); 1396 (2-57); 1435 

(7-25); 1448 (6-43); 1498 (6-15); 1695, 1697, 1698 

(6-7); 1708 (2-119.1); 1757 (7-11) 

Ancuash, E., 4, 354 (6-36); 752 (2-21.1); 1282 (7-21) 

Anderson, A. B., 207 (7-9); 354 (6-29) 

Anderson, C. W., 14 (2-109); 57 (2-77); 325 (3-1) 

Anderson, W. R., 6248, 6259 (2-80); 6873 (4-4); 7281 

(7-54); 7420 (2-30); 7543 (2-54); 7898, 8085 (2-80); 

8702 (7-20); 10071 (3-16); 10653 (7-24); 10771 (6- 

9); 11037 (2-27a); 11065 (6-9); 11037 (2-27a); 11065 

(6-9); 11921 (7-24); 12064 (2-27b); 12079 (2-69b); 

12208 (2-43b); 12303 (7-54) 

Andrade, P. M. et al., 252 (6-6.2); 420 (6260) (2-114); 

621 (6541) (7-64) 

Andrews, L. M., 911 (2-1); 3-103 (2-40) 

Angelo, L., 11 (7-48); 11 (2-7) 

Antonio, T., 2384 (7-56b); 3763 (7-56a); 4324 (7-56b); 

4599, 4665 (7-36a) 

Aranda, A., 60 (2-15); 73 (7-36a) 

Araquistain, M., 34 (1-1); 2181 (7-22); 3128 (7-36a); 

3149 (5-1) 

Arafijo, D., 1425 (7-85); 2151 (1-1); 4041 (7-36b); 4146 

(1-1); 4536 (7-36b); 4719 (6-26); 4727 (6-25); 4736 

(1-1); 4932, 5261 (6-26); 5834, 6309, 6337 (1-1); 

6400 (6-25); 6408, 7517 (1-1) 

Araijo et al., s.n. INPA94326 (7-73) 

Araijo, J. da S., 72 (2-6); 104 (7-56b) 

Argent, G. C., 6479 (2-41) 

Aristeguieta, L. et al., 4493 (7-56b); 4990 (2-27); 5131 

(2-27b); 6433 (2-12); 6477 (7-56b); 6493 (6-3); 6867 

(1-1); 6908 (7-36a); 6977 (3-7); 7096 (7-34); 12529 

(7-22) 

Armond, 175 (3-4) 

Ar6stegui V., A., 105 (2-128) 

Asplund, E., 10055 (6-33); 12538 (2-4) 

Assis, J. S., 52 (4-2); 188 (2-80); s.n. (4-2) 

Atwood, J. T., 4526, 4723, 5261 (1-1) 

Austin, D. F. et al., 6994 (7-24); 6995 (2-43a); 7058 

(7-50a); 7180 (2-149); 7185 (2-109); 7251 (2-134); 

7315 (2-69a); 7368 (2-109); 7394 (2-43a); 7396 

(3-4) 

Avalone, C. L. et al., 26 (2-41) 

Ayala, F., 615 (2-39); 2888 (6-16) 

Aymard, G. et al., 371 (2-77); 2536, 2637 (7-36a); 3230 

(6-3b); 3343 (7-40); 3483 (7-56b); 3900 (2-109); 3911 

(7-74); 4074 (2-145); 4076 (2-109); 4142 (7-56) 

Badillo, V. M., 1559 (7-56b) 

BAFOG, 119 (3-1) 

Bagazo, N., 166 (7-56a) 

Bahia, R. P., 54 (7-56b) 

Bahia, T. R., 129 (2-124); 130 (7-53); 145 (6-3b); 191 

(2-62); 198 (2-124); 215 (6-41) 

Baker, M. A., 6056, 6112 (6-36) 

Baldwin, A. A., s.n. (2-1) 

Balee, W. L. et al., 17 (6-la); 44 (2-99); 46 (2-86); 125 

(2-99); 174 (2-114); 175, 182 (2-69a); 218 (2-99); 

228 (2-86); 243, 256, 286 (2-69a); 308 (6-la); 325, 

326 (4-3); 336 (2-69a); 339, 354 (2-86); 421 (2-57); 

426 (2-99); 501 (2-69a); 508 (2-99); 581 (2-114); 599 

(2-69a); 664 (6-la); 948 (2-27a); 975 (2-99); 1053 

(2-69a); 1058 (7-56a); 1087 (6-lc); 1089 (6-la); 1107 

(2-99); 1116 (2-63); 1124 (6-la); 1157 (3-3); 1170, 

1179 (2-99); 1186 (6-la); 1191 (2-63); 1273 (3-3); 

1276 (2-99); 1296, 1316 (6-lc); 1359 (2-114); 1375, 

1378, 1429 (6-la); 1439 (2-99); 1451, 1455 (6-la); 

1484 (2-99); 1501, 1509, 1516, 1518, 1520, 1521, 

1522, 1524 (6-la); 1550 (7-56b); 1656, 1685, 1747 

(7-63); 1786 (7-14); 1804 (2-69a); 1806 (2-8); 1861 

(2-27a); 1934 (7-66); 3808 (2-17); 4063 (2-124) 

Bamps, P., 5378 (7-6) 

Barbosa, C., 1213 (7-80) 

Barbour, P. J., 5478 (7-56b); 5505 (7-36a); 5764 (7- 

56a) 

Barclay, A. S. et al., 3701 (7-22) 

Barlow, F. D., 30/62B (7-56a)

108 

Barrier, S. et al., 2618 (7-56a); 3863 (7-39); 3989, 4259 

(2-86) 

Barros, S., 79 (6-3c) 

Barroso, G. M. et al., 181, 290 (7-18); 355 (7-56b) 

Bastos, N. C. et al., 79 (7-20); 87, 117 (2-43a); 127 (7- 

56b) 

Bautista, H. P., 34 (3-2); 48 (2-135); 52, 55 (7-56b) 

Bawa, K. S., 383 (6-15) 

Beaman, J. H., 6092 (6-47); 6263, 6333 (7-36c); 6423 

(6-47) 

Beck, C. H., s.n. (2-1) 

Beck, S. G., 1480 (7-56a); 4049, 6915, 7158, 8319 (7- 

36a); 10158, 10172 (2-28); 12247 (7-26a) 

Becker, R., 54 (7-54) 

Begazo, N., 38 (2-64); 88 (2-27a); 144 (7-50b); 159, 

166, 201 (7-56a) 

Belem, R. P. et al., 2837, 2846 (6-5lb); 3169 (6-17); 

3315 (6-5 lb) 

Bellido, L. C., 2, 35 (2-27a) 

Bena, P., 1317 (2-27a); 1328 (2-52); 1757 (7-36a) 

Benitez de Rojas, C. E., 2578 (2-89); 2579 (2-81) 

Benson, W. W., 5671 (7-50); INPA92145 (7-50a) 

Berg, C. C., 221 (6-25); 617 (2-17); 684 (6-21); 686 (4- 

3); 697 (2-81); 712 (2-69a); 719 (2-81); P18449 (7- 

54); P18452, P18565 (2-28); P18654 (2-75); P18678 

(3-4); P18693 (2-28); P19462 (2-60); P19751 (2-28); 

P19793 (6-3b); P19796 (7-73); P19922 (2-99); 

P19928 (2-sp.); P19940 (2-28) 

Berlin, B., 354 (6-36); 902, 976, 1799 (2-21.1); 3529 

(7-56b) 

Bermudez, B., 6 (1-1) 

Bernal, C. et al., 142 (6-48) 

Bernardi, A. L., 1-56 (7-40.1); 1-95 (2-64); 1-163 (6- 

15); 2-109 (6-44); 2-162, 2-163 (2-17); 3-51 (2-55); 

3-99 (6-44); 3-124 (6-34); 4-142 (6-15); 5-36 (2-94); 

5-120 (2-124); 6-6 (6-6.1); 6-74 (6-44); 6-123 (7-41); 

6-148 (2-141); 6-189 (2-27a); 6-192 (2-27); 7-2 (7- 

6); 7-84 (6-6.1); 7-89 (7-41); 8-30 (2-94); 8-128 (6- 

15); 8-139 (2-60); 8-151 (2-55); 9-78 (6-15); 9-157 

(2-141); 1624 (3-4); 2030 (2-13); 2127 (3-3); 2777 

(1-3); 3069 (7-21); 3081 (7-56d); 6628, 6682 (3-4); 

6760 (7-25); 7111 (7-16); 7356 (2-122); 7394 (7- 

56b); 7416, 7672 (7-36a); 7824 (6-3b); 7825, 7828 

(2-77); 8011 (2-122); 16219 (6-6.1); 16233 (7-6); 

16334 (3-9) 

Berry, P. E., 600 (2-69b); 633 (7-62); 662 (7-56); 737 

(7-62); 1352 (7-50b); 1519 (2-69a); 2066 (2-27a); 

2073 (2-69a); 2149 (7-49); 2231 (3-4); 3383 (3-7) 

Bethancourt, A. et al., 91 (7-56b) 

Bhorai, M., 8781 (2-44) 

Billiet, F. et al., 1023 (7-25); 1180 (2-69a); 1201 (2- 

108); 1234 (2-124); 1265 (1-1); 1298 (7-25) 

Bisby, F. etal., P18071 (2-69a); P18077 (2-82); P18085, 

P18102 (4-3); P18107 (7-56a); P18123 (2-56a) 

Bitaillon, C., 64 (3 sp.) 

Black, G. A., 48-2902 (2-69.1); 48-3610 (2-63); 49- 

8473 (3-3) 

Blanco C., C. A., 55 (2-113); 67 (7-20); 194, 254 (2- 

77); 421 (2-109); 533 (7-20); 673 (7-16); 764 (7-56b); 

766 (2-27b); 885 (7-36a); 1105 (2-17); 1112 (7-56); 

1141 (2-17); 1157 (6-4); 1166a(2-77); 1166b(2-93); 

1204 (3-6); 1221 (6-4); 1255 (2-140) 

Bokermann, W., 5732 (7-82) 

Boom, B. et al., 1997 (2-64); 4087 (7-53); 4167 (7-46); 


4178 (7-56a); 4245 (7-46); 4335, 4338, 4343, 4346, 

4386,4387,4395 (2-43b); 4424 (7-46); 4426 (2-43b); 

4428 (2-21); 4461 (2-43b); 4464 (7-46); 4473 (2- 

43b); 4497 (7-46); 4630 (7-53); 4700, 4729 (7-56a); 

4940 (6-14); 5084 (2-43b); 5367 (7-62); 5868 (2- 

27.1); 5919, 5952 (2-26); 6032 (7-56b); 6089 (2-88); 

6263 (3-4); 6359 (2-145); 6490 (2-13); 6492 (2-99); 

6514 (3-4); 6534 (2-88); 6536 (2-43a); 6542 (7-20); 

6551 (3-4); 6574 (7-56b); 6587 (3-4); 6592 (2-13); 

6607 (2-99); 6623 (2-13); 6633 (2-145); 6636 (2-99); 

6641 (2-43a); 6660 (2-145); 6676 (2-99); 6677 (2- 

88); 6685 (2-99); 6838 (7-83) 

Borjas M., G., 175 (7-27) 

Bossio, H., 34 (2-69b) 

Boucas, P. R. P. et al., 13 (2-52); 17 (2-114); 148 (7- 

56b) 

Boyan, J., 75 FD7749 (7-11.2) 

Braga, M. M. N. et al., 145 (6-3a) 

Braga, P. I. S. et al., 3127 (2-10); 3347 (7-56a) 

Brand, J., 971 (3-7); 992 (2-45) 

Brandbyge, J. et al., 30575 (7-40); 31228 (7-56b); 32659 

(6-36); 35032 (2-8); 36119 (7-53); 36177 (7-22) 

Breedlove, D. E. et al., 20898 (1-1); 34086 (7-22); 34130 

(7-56b); 58466 (7-22) 

Breteler, F. J., 4606 (3-7); 4799 (7-50b); 4881 (7-40); 

4883 (7-56b); 4959, 4961 (2-77); 4994 (2-109); 5043 

(2-27a); 5078 (2-77); 5123 (2-13); 5133 (3-4); 5134 

(7-16) 

Bristan, N., 1121 (6-22); 1131 (7-36a) 

Brown, S. et al., 1639 (1-1) 

Bruijn, J. de, 1252 (3-7); 1626 (3-3); 1635 (2-77); 1638 

(2-88); 1664 (7-36a); 1688 (3-4); 1694(2-109); 1696, 

1698 (3-4); 1716 (2-77); 1729 (2-114); 1735, 1736 

(2-88) 

Brumbach, W. C., 8332 (2-1); 9263, 9327, 9655, 9739 

(1-1) 

Briinig, E. et al., 160 (2-68) 

Bunting, G. S. et al., 3505 (7-25); 3744 (2-69a); 3746, 

3825 (7-28); 4002 (1-68); 4032 (3-1); 4084 (2-88a); 

4091 (2-119); 4231 (2-83); 5521 (3-7); 5655 (7-22); 

5889 (3-7); 5947 (2-12); 6224 (3-7); 6225 (7-22); 

6543 (3-7); 6682 (1-1); 6854 (2-43a); 7055 (3-7); 

7124 (2-76); 7152 (7-22); 7219 (2-43a); 7321 (7-20); 

8457 (3-7); 8725 (7-36a); 8750 (2-76); 8758 (7-22); 

8763 (2-76); 8764 (3-7); 8797 (7-36a); 8836 (7-56b); 

8949 (7-36a); 8954, 9192 (3-7); 9832 (2-114); 10151, 

10216 (7-36a); 10587 (7-56b); 10759 (2-43a); 10800 

(3-7); 10929 (7-56a); 11036 (2-43a); 11274 (3-7); 

11507 (2-114); 11529 (2-43a); 11992 (7-36a) 

Burch, D., 3409 (1-1) 

Burger, W. C. et al., 5784 (7-61) 

Buschbacher, R. J., 39 (21) 

Busey, P., 354 (7-56a) 

B. W., 3371 (6-1a); 3404 (7-50b); 3827 (2-130); 4818 

(7-25) 

Byron et al. (=Albuquerque, B. W. P. de), 67-20 (2- 

69.1); 67-34 (6-15); 67-59 (2-39); 67-70 (2-69a); 264 

(6-3); 370 (3-6); 372 (7-50a); 377 (2-27a); 509 (6- 

3b); 577 (2-1); 592 (2-28); 594 (6-3b); 633 (7-40); 

886 (7-55b); 924 (2-57); 927 (2-17); 946 (2-28) 

Cabrera, E., 3138 (1-1) 

Cabrera R., I., 94 (2-5.2); 553, 600 (2-148); 2590 (3- 

4); 2597 (7-56b); 3217 (7-56a); 3359 (2-69a) 

Cain, S. A., 12 (7-36a)


Caldas, T. S. P., 21 (6-27) 

Calder6n, C., 79-105 (2-88a); 2521 (7-53); 2534 (7- 

56a); 2542, 2591 (6-29); 2634 (7-9); 2806 (7-56a); 

2820 (2-37); 2936 (2-17); 2938 (2-88); 2944 (2-60) 

Calder6n, M., 34 (1-1) 

Calzada, J. I. et al., 868 (6-47); 7166 (1-1); 8542 (2- 

45 

Cambar, I., 111 (1-1) 

Campbell, D. G. et al., 6254 (2-99); 6341 (2-43b); 6540 

(7-21); 6476 (2-114); 6503 (2-64); 6543 (2-99); 6573 

(2-43b); 7020 (7-50b); P20826 (7-55); P20859 (6- 

11); P20868 (2-69b); P20872 (2-140); P20887 (7-6); 

P20896 (2-99); P20897 (7-2); P21129, P21130 (8- 

4); P21210 (7-36a); P21834 (2-140); P21918 (2-14); 

P21963 (2-57); P21976 (2-27a); P22052 (6-3); 

P22126 (6-9); P22271 (2-99); P22302 (2-36); P22305 

(7-56a); P22313 (2-134); P22314 (2-13); P22315 (6- 

3); P22336 (7-56a); P22370 (7-52); P22389 (2-134); 

P22404 (2-36); P22457 (7-56b); P22470 (2-36); 

P22563 (6-29) 

Capus, F., 147 (2-114) 

Carauta, J. P. P., 973 (7-56b); 2319 (7-85); 3391 (2- 

11) 

Cardona, F., 2489, 2555 (6-10) 

Carnevali, G. et al., 1480 (7-9) 

Carrasquilla, L., 363 (1-1) 

Carreira, L. et al., 724 (7-66); 850 (7-73) 

Carvalho, A.M. de et al., 193 (6-25); 194(6-52.1); 216 

(6-51b); 353 (2-80a); 469 (7-20); 609 (1-1); 647 (7- 

82); 858 (6-32); 877 (6-51b); 1046 (7-20); 1077 (4- 

4); 1106, 1116 (6-52.1); 1366 (2-11) 

Casari, M. B. et al., 277 (6-25); 1073 (2-18) 

Castellanos, A., s.n. INPA27546 (6-16); 24054 (7-54); 

24110, 24150, 24180 (6-27); 24185, 24280 (2-41); 

25263 (2-43a) 

Castilla, A., 152 (2-27a) 

Castillo, A., 31 (6-6a); 422 (2-27b); 692, 705 (7-36a); 

736, 742 (7-56a); 1311 (3-4); 1407, 1417(2-77); 1526 

(7-25); 1535 (6-3b); 1592 (2-83); 1798, 1848 (7-25) 

Castro, J. H., s.n. (2-32) 

Castro, M., 10 (2-45) 

Cavalcante, P. B., 441 (7-56a); 1789 (6-3b); 2309 (2- 

81); 2441 (7-56b); 2065, 2276 (6-15); 2477 (7-25); 

2764 (6-15); 2768 (6-41); 2809, 2889, 2951 (7-56a); 

3041 (6-36); 3088 (2-69b); 3202 (6 sp.); 3276 (2- 

27b); 3308 (3-6); 3371 (7-56b); 3373 (7-20); 3378 

(2-43a) 

Cedillo T., R., 577 (2-45) 

Cerrato, B. C. A., 141 (1-1) 

Cesar, A., 1 (7-73) 

Cezario & M. G. Vieira, s.n. INPA 20606 (6-15) 

Champagne, H., 49-1 (2-130); 49-2 (2-86), 49-3 (2- 

109); 49-4 (2-86) 

Chan, C. et al., 813, 1483, 1615 (1-1) 

Chavelas P., J. et al., ES2059, ES2124, ES2144, 2600 

(7-56b); ES2818 (7-36); ES2960, 3343 (2-15) 

Chiang, F. et al., 510 (2-45) 

Christenson, G. M. et al., 1155 (2-114); 1161 (2-27); 

1399 (2-69b); 1430 (2-140) 

Chrostowski, M. S., 70-398 (7-73) 

Churchill, H. W. et al., 4345 (7-56b) 

Cid, C. A. et al., 59 (7-52); 228 (7-56a); 278 (2-88); 

317 (7-56a); 365 (3-4); 428 (2-75); 431 (7-52); 487 

(7-8); 489 (7-56a); 516 (3-4); 640 (7-11); 704 (7-56a); 

860 (7-11); 1059 (6-41); 1131 (7-2); 1139 (2-93); 

1153 (2-134); 1201 (6-29); 1230 (2-134); 1281 (6- 

15); 1433, 1549 (7-56b); 1579 (6-3a); 1627 (4-3); 

1644 (7-50b); 1645 (7-56a); 1671 (7-56b); 1769 (6- 

3a); 1797 (7-56a); 1899(7-1); 1912 (7-56a); 1916(2- 

43b); 1971 (7-15); 2089 (2-134); 2207 (6-3a); 2217 

(7-56a); 2261 (6-32.1); 2333 (6-3a); 2437 (7-50a); 

2534 (2-64); 2542 (7-56a); 2545 (7-66); 2665, 2792, 

2798 (7-56a); 2946 (7-66); 3048 (2-69a); 3184 (6- 

29); 3211 (6-36); 3226 (2-69a); 3296 (6-29); 3349 

(2-75); 3427 (7-62); 3509 (2-52); 3514 (2-69b); 3555 

(6-3b); 3556 (2-52); 3575 (2-27a); 3633 (2-129); 3650 

(2-27a); 3688 (2-129); 3696 (6-3b); 3728 (2-124); 

3747 (2-129); 3778 (6-lb); 3810 (6-3b); 3856 (6-9); 

3869 (7-9); 3948 (2-69b); 3895 (7-6); 3949 (7-56b); 

3953 (2-69a); 3993 (7-2); 4003 (7-56b); 4088 (6-36); 

4162 (7-52); 4181 (7-56b); 4252 (2-69b); 4258 (6- 

3a); 4340, 4434, 4474 (2-41); 4500 (7-56b); 4530 (7- 

73); 4536, 4574 (7-36a); 4600, 4656 (7-56a); 4661 

(7-36a); 4682 (7-73); 4685 (7-56a); 4724, 4729 (7- 

56b); 4768, 4831, 4956 (7-36a); 4995 (7-56a); 5065 

(6-la); 5259 (7-5); 5336 (6-7); 5343 (2-121); 5589 

(2-83); 5662, 5860 (2-43b); 5956 (2-16); 5982 (7- 

50b); 6052 (3-16); 6093 (6-27); 6103 (3-16); 6135 

(7-20); 6137 (6-27); 6170 (2-124); 6218 (7-56a); 6228 

(2-30); 6232 (7-54); 6253 (2-37); 6318 (2-124); 6341 

(7-15); 6368 (6-3c); 6380 (2-37); 6434, 6477 (2-114); 

6501 (2-141); 6503 (2-27a); 6505 (2-141); 6521 (2- 

27a); 6571 (7-66) 

Clark, H. L., 6491 (3-4); 6661, 6737, 6740, 6753 (8- 

3); 6963 (2-69a); 7033 (2-82); 7113 (2-27a); 7165 

(2-39); 7250 (2-69a); 7290 (7-8); 7337 (2-69a); 7342 

(2-82); 7362 (2-69a); 7406 (2-62); 7473 (7-21); 7508 

(2-69a); 7527, 7629 (2-36); 7669 (3-6); 7778 (2-69a); 

7779 (7-8); 7783 (2-57); 7789 (7-8); 7811 (2-39); 

7817 (2-69b); 7846 (2-57); 7924 (2-39); 7926 (2-76); 

7926 (2-94); 7928 (2-24); 7981 (2-27a); 8019 (2- 

129); 8032 (2-27a); 8057, 8063 (8-3); 8082 (3-la); 

8157 (6-la); 8211 (8-3); 8232 (2-24); 8327 (7-8) 

Clausen, R. T. et al., 6238 (2-1) 

Coelho, D. et al., 356, 380 (2-52); 395 (6-3a); 406 (2- 

27a); 411 (6-3a); 708 (2-119.1); 817 (7-8); 843 (2- 

36); 861 (2-64); INPA5332 (2-124); s.n. INPA51497 

(7-73); INPA53263 (7-40); INPA53289 (7-56); 

INPA53292 (6-3b); INPA53295 (2-23); INPA53339 

(2-43b); INPA92490 (7-6) 

Coelho, L. et al., 17 (7-6); 92 (2-43b); 134 (7-73); 206 

(2-109); 308 (2-94); 361 (7-6); 382 (6-41); 432 (7- 

6); 479 (2-69b); 492 (2-57); 496 (3-6); 522 (2-60a); 

533 (2-69b); 561, 572, 650 (2-81); 651 (6-29); 708 

(2-119a); 835 (2-69a); 1837 (7-54); 1955 (2-43b); 

1975 (7-56b); 1979 (7-56a); 1986 (2-79); 5209 (2- 

69.1); INPA25903 (2-112); s.n. INPA42005 (6-21); 

s.n. INPA42095 (7-25); s.n. INPA42141 (7-20); s.n. 

INPA42161, s.n. INPA42171 (6-3a); INPA53292 

(6-3b) 

Coker, W. C., s.n. (2-1) 

Collares, J. E. R. et al., 110 (7-18); 121 (2-80) 

Cominote, J., 84 (3-4) 

Conant, D. S., 993 (7-56a); 1100 (7-1) 

Conejos, J., 24 (2-77); 28 (3-3); 43 (3-4) 

Contreras, E., 10715, 10716 (7-23); 10719 (7-56b); 

10729 (7-23); 10742 (2-18.3) 

Coradin, L. et al., 75 (7-20); 689 (7-25); 781, 784, 785


(7-56b); 1051 (7-25); 1080 (7-73); 4876 (7-25); 4880, 

5022,5023, 5068 (7-56b); 6434 (2-80); 6471 (7-56b); 

7425 (2-80) 

Cordeiro, I. et al., 89 (4-1); 98 (2-91); 119 (2-69a); 147 

(2-99); 175 (6-la); 285 (7-49); 289 (2-165); 300 (2- 

99); 306 (3-6); CFSC6544 (2-144); CFSC6782, 

CFSC7551 (7-54) 

Cordeiro, M. dos R., 14 (2-99); 82 (2-43b); 100 (6-20); 

105 (2-43b); 153 (3-4); 159 (2-43b); 169 (7-73); 263 

(2-98); 468 (6-21); 497 (7-56b); 535 (2-43b); 626 (2- 

62); 695 (6-3b); 703 (7-11.1); 755 (2-22); 759 (2-62); 

814(2-37); 1027 (2-69a); 1156(2-144); 1162(2-17); 

1270 (2-27b); 4679 (2-15) 

Correa A., M.D. et al., 689 (7-38); 958 (5-1); 1815 (2- 

5.3); 4679 (2-15) 

Correll, D. S. & H. B., 35873, 42500 (2-1); 45500, 

49992 (1-1); 51946, 52557 (2-1) 

Costich, D. et al., 809 (7-2); 847 (7-1); 927 (2-114); 

961 (6-9) 

Couret, 18 (7-73); 243 (6-45); 263 (2-18); 264 (2-77) 

Cowan, C. et al., 2253 (1-1) 

Cox, D. K., 3304 (7-56b) 

Cremers, G., 4628 (7-73); 5091 (7-69a); 5338 (7-25); 

5367 (2-134); 6498, 6538 (7-56a); 6546 (7-74); 7011 

(2-114); 7711 (1-1) 

Croat, T. B., 8129 (7-36a); 10018 (1-1); 13246 (7-36a); 

14648 (2-88); 16751 (7-36a); 17723 (6-44); 17750 

(2-69b); 17867 (6-44); 18574 (7-56a); 19382 (6-41.1); 

19883 (2-39); 20014 (2-17); 20218 (2-69b); 20332 

(6-22); 20367 (3-14); 20372 (2-43c); 20410 (7-56a); 

20443 (3-9); 20663 (6-38); 20673 (2-69b); 23256, 

24187, 24211 (1-1); 24289 (7-36a); 24492, 24977 

(1-1); 33627 (7-36a); 39057 (1-1); 45746 (2-45); 

54099 (7-21); 54147 (7-49); 51405 (7-36a); 57982 

(6-36); 59199 (7-56b); 61008 (1-1) 

Cuadros V., H., 2133, 2181 (7-22) 

Cruz, A. et al., 21 (7-36a); 151 (7-21); 170 (7-36a) 

Cumana, L. J. et al., 779 (1-1) 

Curran, H. M., 2M57, 2M74 (1-1) 

Daly, D. C. et al., 141 (7-56a); 149 (4-3); 158 (7-56a); 

181 (2-114); 259 (7-16); 369 (2-56b); 509 (2-141); 

639 (2-56b); 679 (2-69a); 681 (7-52); 703 (7-20); 709 

(7-18); 814 (2-108); 820 (2-69a); 831 (6-50); 833 (2- 

81); 839 (2-56b); 862 (2-63); 875 (2-56b); 882 (2- 

63); 898 (2-27a); 992 (7-56b); 1028 (7-24); 1056 (7- 

56b); 1115 (6-1); 1117 (2-114); 1342 (2-69a); 1373 

(2-27a); 1374 (7-56a); 1413 (7-24); 1511 (2-69a); 

1516, 1552 (2-24); 2035 (7-56a); 2069 (7-36a); 2928 

(2-99); 3825, 3850, 3927 (2-69a); 3931 (7-24); 3943 

(6-la); 3968 (2-39); 4037 (2-108) 

Dambros, L. A., 82, 86 (6-27); 87, 119 (2-41); 138 (7- 

54) 

Damiao, C., 593 (6-34); 594 (2-22); 600 (6-15); 616 

(2-99); 619 (6-44); 621 (6-3b); 633 (7-50b); 656 (2- 

128); 674 (2-114); 695 (2-124); 702 (6-44); 718 (7- 

50b) 2491 (2-69b); 2513 (2-124); 2527 (2-69b); 2531, 

2579 (6-3b); 2587, 2599 (2-27a); 2637 (2-137); 2712 

(3-4); 2784 (2-69b); 2884, 2898 (2-27a); 2904 (6- 

3b); 2908 (7-56a); 2924 (6-15); 3034 (6-36); 3067 

(2-124); 3071 (6-lb); 3079 (2-27a); 3081 (2-69b) 

Danin, A., 76-35-5 (2-45); 77-3-6 (7-56b) 

Dantas, M., 12374 (6-3b); 12382 (7-40); 12391 (2- 

124); 12392 (7-29) 

D'Arcy, W. G. et al., 13662 (7-56a); 14273, 14297, 

14329, 14548 (7-56b); 14559 (1-1); 14596, 14653, 

15015 (7-56b) 

Daubenmire, R., 95 (2-15); 392 (7-56b); 489 (2-45); 

716 (7-56b) 

Davidse, G. et al., 2327 (1-1); 4306 (7-40); 4358 (2- 

12); 4446 (7-56a); 4466 (3-7); 5415 (2-69b); 6753, 

6953 (7-29); 12132A, 12208A (2-27b); 12260A (2- 

69b); 12354 (7-56b); 12361 (2-18); 12385A (3-7); 

12402 (2-69b); 12423 (7-56a); 12429, 12435 (6-3b); 

12517 (7-56b); 12531, 12535 (2-27b); 12538 (2-69); 

12540 (2-27b); 12553 (6-3b); 12625 (2-69b); 12637, 

12714 (2-27b); 12737 (6-3b); 12807, 12855 (2-27b); 

13015 (6-3b); 13055 (7-56b); 13109 (2-69); 13148 

(2-27); 13243,13414 (2-27b); 13838 (7-56b); 13852, 

13944 (2-88a); 13953 (2-27b); 13979 (2-134); 13981 

(7-56b); 14090 (2-69b); 14180(7-40); 14305 (2-88a); 

14327 (2-60b); 14366, 14393 (2-27b); 14536 (2-88a); 

14546 (2-69b); 14654 (2-88a); 14708 (2-27b); 14711 

(6-3b); 14765, 14842(2-27b); 15107 (2-56b); 15118, 

15212 (6-3b); 15292 (7-56b); 15451 (6-3b); 15490 

(2-88a); 15491 (7-56b); 15541 (2-47); 15550 (7-40); 

15619 (2-140); 15662 (7-56); 15665 (2-69b); 15756 

(2-140); 15761 (7-56a); 15768(2-140); 15769 (6-3b); 

15788, 15894 (2-69b); 15907, 16005 (6-3b); 16022 

(3-7); 16025 (7-56b); 16027 (2-88a); 16030 (2-134); 

16093 (7-40); 16150 (2-69b); 16176 (3-7); 16184 (2- 

27b); 16265, 16308 (2-122); 16276 (7-56b); 16291 

(3-1); 16363 (2-43a); 16420 (7-20); 16426 (2-13); 

16432, 17476 (2-77); 16753 (7-29); 16889 (2-121); 

16979 (7-62); 17010 (2-27b); 17032 (2-119); 17088 

(2-28); 17098 (2-131); 17144 (2-119); 17219 (6-3b); 

17329 (2-69b); 17387 (2-28); 17389 (2-119); 17409 

(2-131); 17666 (2-141); 17721 (2-39); 17837 (1-1); 

18056 (2-52); 18304 (7-50a); 18372 (7-22); 18575 

(2-114); 19155, 19188, 19482 (7-36a); 20227 (1-1); 

20999 (7-36a); 22578a, 22747 (7-21); 22903 (7-25); 

22962 (7-56a); 23267 (2-45); 23705 (2-88); 27217 

(6-7); 27627 (2-27a); 27671 (2-140); 27730 (2-27a); 

27735 (2-102); 27740 (2-88); 27852 (7-8); 27883 (2- 

27b); 27886 (2-88); 27888 (2-69b); 27951 (7-40); 

30157 (2-45) 

Davidson, C. et al., 5206 (7-5); 5357 (7-53); 5437 (2- 

140); 8516 (7-36a); 9774 (7-40); 9848 (6-44); 10581 

(7-56a); 10604 (6-29) 

Davila, H. R. et al., 2 (7-56b) 

Davis, D. H., 180 (7-25); 204 (2-39); 814 (6-43); 890 

(7-56b); 1657 (6-43) 

Davis, E. W. et al., 1080 (7-56b) 

Davis, H. A., 15609 (1-1) 

Davis, P. H. et al., 2259 (7-20); 60303 (2-119.1); 60326 

(7-56a); 60337 (6-28) 

De La Cruz, J. S., 3406 (2-66) 

Delascio, F. C. et al., 5534 (2-27b); 7361 (4-3); 9662 

(2-140); 11135 (6-3b); 11187 (7-56b); 11190 (6-3b); 

11349 (7-56b); 11350, 11409 (6-3b) 

Denslow, J. et al., 76 (6-57); 79-25 (7-61) 

Dept. de Botanica USP, 3 (7-56a) 

Deward, G. 155 (2-134) 

Dezzeo, N., 293 (7-56a) 

Dias, A. A., 46 (2-30) 

Diaz, C. et al., 68 (7-5); 288 (6-3b); 397 (6-35); 438 

(2-140); 638 (7-40); 916 (2-68); 944 (7-40); 1527 (6- 

44); 1751-18 (7-75); 1739-118 (2-43c); 1758-32, 

1758-22 (2-27a)


Diaz, M., 2-A (6-44); 51-A (2-64); 87A (2-141); 127A 

(6-44); 162A (7-41); 6-52 (7-40.1) 

Dieckman, L., 318 (2-88a) 

Dionisio et al., 120 (6-46) 

Dios Holmquist, J. de, 60 (6-3b) 

Dodson, C. H. et al., 1170 (7-35); 2945 (2-69a); 5464 

(7-87); 7516 (2-5); 8655 (7-87); 8672, 12407 (2-5) 

Dressier, R. L., 3322 (7-38); 3416 (7-36a); 3736 (7- 

38); 4622 (2-88) 

Duarte, A. P., 5420 (2-4); 5607 (2-71a) 

Ducke, A., HPB1550 (7-18); MG7367 (2-129); 

MG7876 (7-63); MG10563 (2-57); MG10966 (2- 

43a); MG12221 (2-129); MG15289 (2-76); 

MG16153, MG16260 (2-43a); MG16355 (2-79); 

MG16430 (2-60); MG16525 (2-43a); RB15137 (6- 

18) 

Duke, J. A., 5682 (7-56); 13513 (7-36a); 15256 (2-145) 

Dunbar, P. L., s.n. (1-1) 

Duran, A. R. Camacho, 147 (7-56b) 

Dwyer, J. D. et al., 511, 592 (1-1); 1047 (2-31.1); 4298 

(7-56b); 5047 (2-31.1); 7324 (5-1); 10986 (1-1); 

12252, 15138 (7-56b) 

Echeverry E., R., 2078 (2-56) 

Egler, W. A., 577 (2-36) 

Ehrendorfer, F., 74108-25 (2-122) 

Eiten, G. & L., 3253 (2-41); 4665 (4-2); 8389 (6-27); 

8461 (7-19); 8503, 8734 (7-20); 8904 (2-27a); 9764 

Fernmndez C., J., 3844 (6-27) 

Fernandez, M. M. et al., 5 (2-80) 

Fernmndez P., A. et al., 6124 (2-24); 6975, 7005 (7-21) 

Ferrari, G., 1231 (2-27a); 1272 (2-12) 

Ferreira, E., 58-320 (7-55.1) 

Ferreyra, R., 4603, 4636 (7-56b) 

Feucht, S. P., 743 (1-1) 

Feuillet, C., 487 (7-74); 985 (7-52); 1089 (2-124); 2303 

(2-86); 2314 (2-52) 

Filho, L. C. de O., 27 (7-18); 28 (7-20) 

Figueroa, V., 122 (7-56b) 

Flores, J. S., 8256, 8322, 8800, 9323, 9867 (1-1) 

Florschiitz, P. A. et al., 2810 (7-1 lb) 

Foli, D. A., 49 (3-4); 61/79 (2-146); 76/79 (7-13); 80 

(7-70.1); 89 (7-70a); 120 (2-6); 228 (2-78.1) 

Folsom, J. P. et al., 1435 (2-5.3); 1932 (7-36a); 1949 

(5-1); 1996 (2-31.1); 2774 (1-1); 3508 (2-88); 3539 

(7-36a); 5672 (6-47); 6781 (7-36a) 

Fonseca, S., 350 (2-30) 

Fonseca, W. N., 129 (4-2) 

Fontella, J., 2275 (1-1) 

Forero, E. et al., 4005 (2-58); 4410, 4424 (7-10); 4684 

(7-35); 4788 (2-124); 4862 (3-15) 

Forero P., L., s.n. (1-1) 

Forest Service, French Guiana, 3511 (2-134); 3513 (6- 

18); 3758 (7-25); 4237 (7-56a); 4503 (7-25); 6097 

(2-52); 6231, 6440 (2-69a); 7259 (7-7); 7786, 7793 

(6-27) 

Elburg, J., LBB9373 (6-18); LBB9387 (3-1); LBB9440 

(2-134); LBB9815 (7-56a) 

Elcoro, S. et al., 210 (7-56a); 223 (2-56); 233 bis (2- 

69a) 

Emmerich, M., 4579 (2-41); 4601 (7-54); 4638 (7-20) 

Emygdio, L. (see Mello Filho, L. E.) 

Encarnaci6n, F., 970 (2-61); E1060 (2-69a); 1243 (2- 

23); 1254 (6-44); 25033 (2-27a); 25037 (6-3b); 25041 

(2-27a); 26124(7-41); 26183 (7-36a); 26201 (2-121); 

26303 (6-38); 26369 (2-27a); 26392 (2-69a); 26409 

(2-98) 

Erlanson, C. O., 78 (1-1) 

Espejel, I., 5-3, 404 (1-1) 

Espina, J. et al., 205 (6-36); 207 (3-6) 

Espinal, M., 163 (7-56b) 

Espinal T., S. et al., 4110 (2-4); 4537, 4608 (2-48) 

Estrada, U. et al., 11 (2-45) 

Euponino, A., 195 (7-64); 212 (7-13); 309 (6-5 lb); 361 

(7-50); 405 (6-51b) 

Faircloth, W. R., 1077 (2-1) 

Falcao, M. et al., 143 (2-43b); 192 (6-15) 

Fanshawe, D. B., 621, 638 (6-6) 

Farifias, M. et al., 299 (2-88); 330 (7-21); 471 (7-8); 

658 (2-69b); 661 (2-27a) 

Farney, C., 1101 (6-26) 

Fennell, J. L. et al., 955 (1-1) 

Fernandez, A. et al., 1021 (7-74); 1618 (7-25); 1706 

(2-27a); 1738 (7-56b); 1779 (2-27a); 1783 (6-3b); 

1784 (2-27a); 2855, 2872 (7-56b); 2873 (7-21); 3022 

(7-40); 3531 (2-69a); 3561 (3-1); HPB2517 (7-18); 

HPB3075 (1-1); HPB3453 (2-44); HPB3782 (2-43a); 

HPB6908 (6-51a); HPB8066 (7-56b); HPB7221, 

HPB8811, HPB9032 (7-18); HPB9373 (2-37); 

HPB 10015 (4-2); HPB 10016 (7-18); HPB 10051 (2- 

30); HPB 10723 (6-27); HPB 10724 (7-20); HPB 10595 

(2-30); HPB10928 (2-27a); s.n. (1985) (2-28) 

(6-2) 

Foresta, H. de, 190 (2-109); 202 (2-69a); 277 (2-109); 

555 (7-2); 630 (7-74); 642 (7-25); 692 (2-149); 746 

(7-56a); 750 (2-134); 751 (2-63) 

Fortin et al., 8565 (6-47) 

Fosberg, F. R. et al., 29203 (6-44); 42591 (7-56b); 

53830, 54087 (1-1) 

Foster, R. B. et al., 2252 (2-15); 3091 (7-36a); 3725 

(6-36); 4043 (7-5); 4074 (6-35); 4299 (7-2); 4385 (7- 

53); 4568 (6-36); 4570 (3-14); 4683 (6-35); 4709, 

4788 (7-56a); 5623 (2-21); 5774 (3-9); 5909 (2-21); 

7961 (2-5); 8021 (6-36); 8733, 8926 (7-73); 10942 

(2-88); 10996 (2-144.1) 

Fournet, A., 445 (2-56b) 

Frame, D., 201 (7-56a) 

Frankie, G. W., 259C, 292C (7-56b); 398C (7-61) 

Freeland, J. et al., 112 (6-47) 

Freeman, O. M., s.n. (1-1); s.n. (2-1) 

Froehner, C., 361 (2-39); 362 (2-64) 

Fr6es, R. L., 41 (2-88); 12711/80 (3-3.1); 21321 (6- 

35); 22431 (6-lb); 22645 (6-la); 24434 (7-73); 25483 

(6-9); 25505 (4-1); 25928 (2-63); 27517 (3-4); 29044 

(6-3b); 29277 (6-29.1); 29562 (7-24); 30435, 30654, 

30688 (6-lb); 31898 (2-66); 32159 (2-97); 32311 (2- 

63); 33653 (6-la); 34168 (2-97); 34888 (6-lb) 

Funk, V. A. et al., 8415 (2-47.1) 

Furlan, A. et al., CFSC7513 (6-27); CFCR7138 (7-20) 

Garcia, C., 30 (2-45) 

Garcia-Barriga, H. et al., 3431 (2-27b); 18202, 18207, 

18209 (2-94); 18457 (2-129); 18528 (2-12); 18924 

(7-36a); 20323 (7-56b) 

Gamier, 76 (7-56b); 79 (6-7) 

Gasche, J. et al., 28 (7-24); 103 (2-69b); 154 (7-5); 211 

(2-37); 216 (6-29) 

Genelle, P. et al., 261 (2-1) 

Gentry, A. H. et al., 385 (2-45); 388 (3-45); 5800 (2- 

145); 6172 (6-13.1); 7605 (2-88); 7723, 8364 (7-22);


8884 (2-95); 10418 (2-27a); 10780 (7-35); 10831 (6- 

3b); 10835 (7-21); 10863 (2-68); 10936 (7-40); 10966 

(2-24); 12905 (2-27a); 13306 (7-56b); 13439 (7-38); 

14554 (7-56b); 14570 (7-25); 14624 (2-140);14641 

(7-56b); 14943 (7-74); 15536 (6-3b); 15767 (2-27); 

15807 (7-6.1); 16638 (3-14); 16701 (7-40); 17224 

(1-1); 17482 (7-10); 18140, 18148 (7-22); 18318, 

18455 (6-44); 18466 (3-9); 18527 (2-52); 18543 (7- 

40); 19094 (3-14); 19098 (2-21); 20371 (3-14); 20385 

(6-41.1); 20678 (7-6.1); 20679 (7-5); 20834 (2-43b); 

20842 (6-3b); 21045 (7-5); 21130 (6-15); 21209 (2- 

43b); 21213 (7-40.1); 21452 (3-16); 22199 (2-45); 

23387 (2-43c); 24116 (3-14); 24331 (2-34); 24962 

(6-7); 25012 (2 sp.); 25057 (6-35); 25064 (3-14); 

25069 (6-35); 25073 (2-124); 25193 (7-53); 25203 

(7-5); 25310 (6-35); 25613 (7-21); 25665 (2-4.1); 

25845 (7-56b); 25866A (2-140); 25913 (3-14); 26030 

(6-44); 26035 (2-140); 26072, 26173 (6-7); 26202 

(2-69b); 26866 (7-36a); 27591 (6-41.1); 27642 (7- 

62); 29704 (3-9); 30147, 30201 (2-51); 30324 (2-5); 

31284 (7-56a); 31502 (2-94); 31657 (2-69a); 31701 

(6-35); 31725 (7-6.1); 31733 (3 sp.); 31824 (7-50b); 

31936 (7-73); 32085 (6-44); 32617 (6-47); 35478 (2- 

34); 36188 (7-56a); 36227 (2-43b); 36238 (7-62); 

36269 (2-43b); 36275 (7-62); 36291, 36304 (2-99); 

36311 (7-50b); 36438 (7-6.1); 36440 (6-35); 36626 

(7-73); 37115 (6-36); 37252 (7-56a); 38047, 38059 

(7-36a); 38077 (3-4); 39243, 39318, 39329 (2-26); 

39350 (2-69a); 39408 (2-128); 39412 (2-99); 39471 

(2-124); 40274 (2-46); 40355 (2-5.1); 40416 (2-124); 

40467 (2-32.1); 40474 (7-56a); 41138 (3-7); 41332 

(7-56b); 41569 (7-53); 41600 (2-69a); 41608 (2-124; 

41682 (2-123); 41712 (3 sp.); 41731 (2 sp.); 41895 

(7-53); 41939 (3 sp.); 42074 (7-56a); 42805 (2-5); 

43029 (2-144.1); 43178 (6-44); 43195 (2-21); 43351 

(7-36a); 43403 (3-5); 43576 (6-24); 43653 (3-5); 

43687 (7-46); 43754 (2 sp.); 43777 (2-21); 43811 (6- 

36); 44539 (3-5); 45582 (2-144.1); 45669 (7-75); 

46170 (2-99); 46213 (2-43b); 46275 (6-36); 46277, 

46278 (2-27b); 46290 (2-83); 46377 (2-69a); 46457 

(6-lb); 46471 (7-21); 47062 (2-63); 47121 (2-27.1); 

47379 (2-69b); 47385 (3-1); 47551 (7-22); 47821 (3- 

17); 47961 (2-148); 48461 (2-15); 48513 (7-61); 48953 

(2-124); 48978 (6-la); 48988 (2-114); 49015 (6-la); 

49045 (7-63); 49103 (2-63); 50319 (7-56a); 51140 

(2-69a); 51214 (7-56a); 51396 (2-94); 51556 (2-21); 

52175 (2-158) 

Gibbs, P. et al., 2708 (7-54); 2738 (3-16); 2839 (7-20); 

3363 (6-27); 3522 (7-82); 3524 (2-126); 3837 (2- 

114); 5087 (2-80); 5165 (7-20) 

Giovanni, F. di, 2 (2-69a); 77 (7-40); 80 (2-140) 

Giulietti, A. M. et al., CFCR2254 (2-144); CFCR3552, 

CFCR6760 (7-20) 

Glaziou, A. F. M. et al., 5689, 7602, 9391, 13796 

(3-4) 

Glenboski, L., 206 (2-43c) 

Godfrey, R. K. et al., 52402 (2-1); 53779, 58121 (1-1) 

G6es, O. C. et al., 152 (2-143); 943, 1046 (3-4) 

Goesnner, F., s.n. (6-3b) 

Gomes, M. et al., 8, 92 (7-56a); 106 (7-56b); 145 (2- 

37); 156 (3-4); 195 (2-37); 226 (7-56a); 381 (2-37); 

432 (3-4); 447 (6-3b); 448, 476 (2-28); 477 (2-27a); 

554, 867, 1015 (7-50b); 1114 (6-la); 1252, 1479, 

1484 (7-50b) 

G6mez, L. D. etal., 18927 (7-56b); 20461 (2-95); 21178 

(7-56b); 23057 (6-47); 23356 (2-15); 24100 (7-36a) 

G6mez-Pompa, A. et al., 1283 (2-15); 3347, 3354 (2- 

95); s.n. (2-88) 

Gonggrijp, J. W., 284 (2-69a) 

Gongora, E., 438, 518 (1-1) 

Gonzales, A., 1159(2-21) 

Gonzales L., L. A., 4752, 4757 (7-56b) 

Gonziles-Quintero, L., 3435 (7-56b); 3505 (7-36a) 

Goodland, R., 289 (7-56b); 712 (2-41); 755 (7-56b); 

894 (6-27); 927 (3-16); 949 (7-18); 963 (7-54); 1074 

(2-135) 

Goodrum, P., 9 (2-1) 

Gordon, B. L., 9 (2-15) 

Gottsberger, G., 16-28782 (7-18); 16-29782, 18-29782 

(4-2); 22-81182 (7-18); 15-15183 (6-21) 

Gottsberger, I. S. et al., 225 (2-41); 368, 152-25771 

(6-27) 

Goulding, M., 29, 54 (2-23); 153 (6-44); 183 (6-23) 

G6mez, L. D. et al., 23356 (2-15) 

Grant, G. B., s.n. (2-1) 

Granville, J. J. de, C-2 (7-74); C-17 (7-56a); 21 (2-60); 

C-51 (7-56a); 115 (7-2); C-143 (7-74); 488 (2-27b); 

509 (7-2); 582 (2-134); 599 (2-50b); 951, 1090 (7- 

56a); T1203 (7-52); 1250 (7-2); 1436, 1739 (7-73); 

1740, 1791 (7-56a); 1802 (7-25); 1883 (2-88); 1946 

(7-56a); 2682 (6-7); 3134 (7-52); 3140 (7-73); 3249, 

3673 (7-2); 3726 (2-88a); 4028, 4136 (7-2); 4321 (7- 

25); 4606 (7-52); 4684 (2-134); 4687 (2-69a); 4700 

(7-56a); 4723 (7-56d); 4731 (2-88); 4838 (7-73); 4867 

(2-69a); 4875 (2-99); 4979 (2-93); 5051 (7-2); 5234 

(2-88); 5242, 5429 (7-2); 5537 (7-56a); 5652 (7-50b); 

5719, 5722 (2-124); 6121 (4-3); 6124, 6171 (7-74); 

6181 (7-2); 6183 (2-114); 6259 (7-2); 6271, 6571 (2- 

64); 6727, 6954 (7-56a); 6993 (2-88); 7177 (7-52) 

B3637 (2-77); B3704 (7-2); B3732 (6-lc); B3785 (7- 

56a); B3787 (2-67); B4499 (7-73); B4507 (7-2); B4611 

(7-56a); B4673 (2-8); 4684 (2-134); B4797 (7-2); 

B4849 (2-88); B4856 (7-56a); B4891 (6-2); B4906 

(7-56a); B5061 (7-2); B5263 (6-18); B5376 (7-73); 

B6326 (2-135); BC101 (2-109) 

Grayum, M. et al., 4090 (7-61); 4971 (7-36a) 

Greenman, J. M. et al., 5739, 5833 (1-1) 

Grenand, P., 38 (2-99); 139, 227 (2-69a); 547 (7-20); 

548, 614 (2-124); 642 (2-69b); 967 (3-2); 1506 (3- 

3); 1582 (7-52); 1718 (2-134); 1774 (6-7); 1818 (2- 

25); 1905 (2-69a); 2037 (7-50b); 2083 (7-73); 2138 

(7-51); 2459 (2-140) 

Grijalva, A., s.n. (1-1); 2308 (2-15); 3443, 3546 (2-45); 

3760 (7-36a); 3890 (2-15) 

Grubb, P. J. et al., 1045 (2-5) 

Guanchez, F. et al., 108 (7-25); 251 (2-88); 252 (2-12); 

396 (6-29); 409 (7-40); 423 (7-25); 510 (7-62); 720 

(2-81); 734, 793 (2-69a); 804 (7-30); 943 (2-83); 965 

(2-103); 989 (2-81); 1098 (7-49); 1228 (2-69b); 1249 

(3-6); 3258 (7-49); 3375 (3-1); 3454 (7-9); 3480 (2- 

121); 3501 (2-140); 3536 (7-49); 3647 (2-99) 

Guarim Neto, G. et al., 38 (7-6); 312 (2-43a) 

Guedes, T. N., 311 (2-149) 

Guevara, L. C. de, 1400, 2125 (2-12)


Guimarbes, J. G., 17 (7-66); 1150, 1462 (2-41) 

Gunn, C. R., 3357 (2-1) 

Gutierrez V., G., 638 (7-40); 2784 (2-17) 

Guzman & Castro, 60 (1-1); 156 (2-45); 559 (1-1) 

Guzman B., R., s.n. (2-15) 

Hage, J. L. et al., 285 (7-20); 564 (7-36a); 1896 (2-11) 

Hahn, W. et al., 444 (1-1) 

Halle, F., 494 (7-52); 2807 (7-73) 

Hammel, B. et al., 975, B1768 (7-56b); 4823 (2-18.1); 

11149 (2-67); 11260, 11577 (5-1); 12476 (2-95); 

14137 (5-1) 

Hamilton, C. et al., 2965 (7-56b); 3281 (7-36a) 

Hansen, B. et al., 9268 (7-83) 

Hanke, W., 73 (6-4) 

Harley, R. M. etal., 15752 (6-25); 16097 (2-114); 17120 

(1-1); 17370(7-20); 17401 (2-88): 18067 (1-1); 18256 

(7-13); 18581 (7-20); 18585 (7-56b); 19058 (7-18); 

19083 (6-20); 20074 (7-20); 21645 (2-30); 21649 (7- 

54); 21735, 21751 (2-80); 21779 (7-18); 22240 (7- 

56b); CFCR7352 (4-4); CFCR7515 (7-18) 

Harling, G. et al., 19409 (7-35) 

Harriman, N. A. 16126 (7-56b) 

Harschberger, J. N., s.n. (2-1) 

Hartmann, R. L., 12005 (7-36a); 12178 (7-56a) 

Hartshorn, G., 809 (7-61); 886 (2-15); 1287 (6-47); 

1523 (7-61); 1625 (2-88a); 1803 (7-56b); 1850 (6- 

47); 1878 (7-61); 2414 (7-56a); 2659 (7-41); 2662, 

2670 (2-27a); 2742 (7-50b) 

de Hass Sr. et al., 136 (7-54); 152 (2-126); 154 (2-43a); 

229 (2-126); 232 (7-70) 

Hatschbach, G., 17886, 20216 (7-85); 21790 (2-41); 

24241 (7-54); 24595 (2-30); 24632 (6-27); 25298 (7- 

66); 29890 (7-54); 30202 (7-20); 31917 (3-16); 32418 

(2-30); 32424 (7-20); 33215 (2-30); 34633 (2-41); 

34674 (2-30); 34686 (7-19); 35405 (7-20); 38466 (7- 

56b); 38936 (6-20); 40137 (2-37); 41243 (7-20); 

41689 (7-54); 43158 (2-114); 43760, 43796 (2-41); 

44093 (7-20); 44112 (7-18); 46750 (1-1); 47027 (7- 

31); 47033 (1-1); 47303 (3-16); 47791 (7-31); 48068 

(2-134); s.n. (2-144) 

Hazlett, D., 3270 (6-47) 

Heithaus, E. R., 48 (7-56b) 

Henderson, A. et al., 267 (7-53); 278 (7-14); 343 (7- 

52); 359 (7-78); 412 (3-4); 448 (7-56a); 477 (6-3b). 

Hendrix, L., 196 (7-5) 

Hensold, N. et al., SPF20850 (6-27) 

Herb. Forestal del Peru, 125 (2-17) 

Heringer, E. P. et al., 1247, 1430, 1492, 1498 (7-54); 

1626 (7-20); 1631, 1687 (7-70); 1761 (2-43a); 1770 

(7-54); 1782 (7-70); 1815 (2-41); 1894 (7-20); 1902, 

1918 (2-43a); 1923, 1934 (7-54); 1957 (7-70); 2067 

(7-54); 2243 (7-20); 2279 (7-66); 2419 (7-70); 2439 

(2-126); 2760 (3-4); 2813 (2-132); 3200 (2-27a); 4527, 

4584, 4805 (7-54); 5021, 5114 (7-20); 5197 (7-70); 

5324 (2-43a); 5399 (7-70); 5409 (2-114); 5445 (6- 

27); 5598, 5604 (2-27a); 5625 (2-41); 5628 (2-27a); 

5696 (2-41); 6258, 6367 (3-16); 6980 (2-43); 7079 

(3-16); 7121 (7-20); 14074 (2-114); 14098, 15109 

(7-54); 15181 (2-43a); 16852, 17520 (7-20); 17533 

(6-27); 17658, 17792 (7-54); 17837 (2-27a); 17856 

(7-54); 18075 (3-16); 18514 (7-54); 18549 (6-49) 

Heringer Salles, A. E., 170 (7-56b) 

Hermann, F. J., 11001 (7-40) 

Hemrndez, H. S., s.n. (1-1) 

Hernmndez, J. J. et al., 244 (7-34) 

Hernndez, L. et al., 112 (6-8); 128 (2-107); 160 (3-1); 

187 (2-69a); 188 (2-43a); 304 (2-69a); 337 (7-25); 

405 (2-113); 442 (3-4) 

Hemnandez, R., 83176 (2-45) 

Heyde, N. M. et al., 179 (7-20) 

Hilario, L., 32 (7-56b) 

Hill, S. R. et al., 2421 (1-1); 12784 (7-9); 12945 (7- 

56a); 13163 (2-69.2) 

Hilty, S., A-1 (6-47.1); 0-1 (2-32.1) 

Hoehne, W., 6171 (2-144) 

Holdridge, L. R. etal., 2517 (7-36a); 6457 (2-15); 6526 

(7-38) 

Holm-Nielsen, L. et al., 19678 (7-56b); 20051 (6-44) 

Hoist, B. et al., 2220 (7-49); 2235 (2-56); 2403 (2-68); 

2592 (7-56b); 2665 (2-91); 2669 (6-10); 2788 (1-3); 

2813 (2-124); 2817 (2-91); 2839 (2-135) 

Honda, M. et al., s.n. (2-69a); s.n. (6-37) 

Hoock, D. K., s.n. (7-18); s.n. (7-25) 

Hopkins, M. J. G. et al., 508 (7-56a); 733 (3-4) 

Howard, R. A. et al., 18133 (7-36a); 18538(1-1); 18675 

(1-2); 18881, 19004, 19389 (7-36a); 19406, 20001 

(1-2) 

Huashikat, V., 622 (2-10); 663 (7 sp.); 783, 946 (7-24); 

1425 (2-81) 

Huber, O. et al., 442, 443 (2-24); 450 (7-68); 524 (2- 

69b); 578 (6-56b); 598 (7-56b); 692 (3-4); 1021 (1- 

1); 1217, 1243 (7-68); 1345, 1425 (7-56b); 1470 (7- 

9); 1553 (2-119); 1561 (6-3b); 1574 (2-119); 1580 

(2-27a); 1628 (2-131); 1669 (2-69b); 1788, 1805 (2- 

119); 1868 (7-71); 2035 (2-131); 2355 (7-49); 2439 

(7-29); 2481 (2-68); 2744 (2-88a); 2747 (6-3b); 2755 

(2-119); 2757 (6-3b); 2763 (7-49); 2821 (2-119); 2870 

(2-39); 3037 (7-49); 3038 (6-3b); 3048 (7-49); 3078 

(7-56b); 3083 (2-140); 3165 (2-131); 3205 (7-29); 

3229 (7-59); 3253 (7-29); 3263 (7-59); 3274 (2-131); 

3276 (2-119); 3280 (6-3b); 3288 (7-59); 3298, 3333 

(2-131); 3337 (7-25); 3344 (2-27b); 3366 (2-27a); 

3397 (2-119); 3441, 3449 (7-29); 3466 (7-21); 3469 

(7-29); 3476 (7-59); 3589 (7-29); 3663 (2-131); 3678 

(7-29); 3685 (2-28); 3690, 3740 (2-119); 3853 (7- 

59); 3886 (2-68); 3914 (2-131); 3932 (2-68); 3960 

(2-131); 4093 (2-68); 4112 (6-3b); 4114 (7-211); 4735 

(2-88); 4744 (7-56b); 4760 (6-3b); 4792 (7-56b); 4795 

(2-83); 4871 (7-59); 4899 (2-88); 4921 (2-131); 5011 

(7-8); 5079 (7-59); 5086, 5151 (2-68); 5162 (2-131); 

5163 (7-29); 5179 (7-56b); 5185 (2-83); 5191 (2- 

27b); 5314, 5343 (2-68); 5360 (7-8); 5412 (2-68); 

5523 (2-131); 5577 (7-29); 5578 (2-68); 5820 (2- 

27a); 5821 (2-119); 5834 (2-88a); 5847 (2-105); 5870 

(2-119); 5944,6076,6088 (2-131); 6155 (7-29); 6210 

(7-21); 6287 (7-56b); 6288 (3-7); 6326 (6-36a); 6670, 

7277 (7-49); 8231 (2-56); 8288 (7-20); 8399 (7-2); 

8433 (2-105); 8448 (7-56b); 8554 (2-56); 9138 (2- 

69a); 9152 (6-8); 9238 (2-69a); 9396 (7-21); 9655 

(3-10); 9696 (7-49); 9899 (7-21); 10253 (7-49); 10286 

(7-21); 10313 (7-30); 10680 (7-8); 10682 (2-119a); 

10852 (2-56); 10875 (7-30) 

Huft, M. et al., 1954, 1982 (7-36a) 

Hume, H. H., s.n. (2-1)


Hunt, D. R., 5506 (3-16) 

Ibarra M., G., 660, 857 (6-47) 

Irwin, H. S. et al., 6070 (2-80); 18231 (7-20); 21229 

(7-56b); 25543, 25651 (3-16); 26068 (7-54); 26586 

(2-27); 27405 (2-126); 28029 (2-144); 30786 (4-4); 

30895 (2-80); 30916 (4-4); 30917 (2-80); 31037 (7- 

56b); 31065 (4-4); 31363, 31376, 31563 (4-2); 47423 

(2-60); 48589a (7-56d); 54990 (2-69.1) 

Isern, J., 1179 (6-44) 

Jacquemin, H., 1403 (7-56a); 1408 (2-109) 

Jangoux, J. et al., 6 (2-134); 30 (7-24); 40 (4-3); 52 (2- 

81); 80 (2-27a); 105 (6-8a); 152 (6-lc); 273 (7-56a); 

296 (7-24); 372 (7-56b); 375, 479, 495 (2-27a); 644 

(7-24); 656 (7-56a); 687 (2-134); 900 (7-56b); 973, 

977 (7-56a); 1008 (7-56b); 1011 (6-lc); 1159 (7-56a); 

1519(2-134); 1604,1629 (7-56a); 1691 (6-3a); 10111 

(7-29) 

Janssen, A. et al., 464 (2-37); 469 (7-73); 478 (7-53); 

523, 545 (7-66); 546 (7-24); s.n. on 17 VI 1980 (2- 

43b) 

Jaramillo, J., 6672 (2-4.2); 6861 (2 sp.); 6960, 7413, 

7527 (2-4.2); s.n. (2-48) 

Jaramillo M., R. et al., 8257 (2-48) 

Jenssen, J., E., 41fr, 54fr (2-69a) 

Jesus, J. A. de, 505 (6-51b); 594 (7-20) 

Jim6nez-Saa, H., 1207 (7-50b); 1246 (7-36a); 1260 (2- 

27a); 1293 (7-22); 1294 (2-27a); 1518 (3-4); 1526 

(2-134); 1543 (3-1); 1540(2-112); 1547 (2-88); 1549 

(2-108.1); 1556 (6-1); 1572 (7-42); 1577, 1586 (2- 

77); 1592 (2-88); 1599 (4-3); 1608 (2-72); 1612 (2- 

88); 1642 (2-72); 1643 (2-43a); 1652 (2-69a); 1656 

(3-11); 1673(2-108); 1683 (2-98); 1688 (2-117); 1691 

(2-112); 1692 (2-134); 1693 (2-108); LBB14332 (4- 

3) 

Judziewicz, E. J., 4481 (7-56b) 

Juncosa, A., 1394 (2-4) 

Jones, G. C. et al., 3418 (2-32) 

de Jong et al., 15784 (2-77) 

Judd, W. S., 1330 (1-1) 

Kalloo, M. B., B.240 (2-86) 

Kayap, R., 49 (7-56a); 674 (6-36) 

Keel, S., 218 (2-27a); 222 (6-3b); 304 (7-24) 

King, S. R., 426 (7-40) 

Kirkbride, J. H. et al., 1000 (2-45); 1133 (7-36a); 1365 

(7-36); 1447, 1448, 2527 (7-36a); 2915 (7-50a) 

Klug, G., 2503 (6-3b) 

Knapp, S. et al., 1255, 1896, 2271, 2785 (7-56b); 2786 

(7-18); 2876 (7-25); 2974, 3017, 3270, 3368 (7-56b); 

3525 (7-80); 3935 (7-36a) 

Kosei, I., 10 (7-56a) 

Koyama, T. et al., 7292 (7-56b) 

Kral, R., 31346, 35228, 35683, 35691 (2-1) 

Krieger, P. L. et al., 12609 (2-27a); 12766 (6-3b); 12795 

(6-29); 12228 (6-3b); 12256 (7-73); 12816 (2-129) 

Krukoff, B. A., 5822A (8-4); 6227, 6361 (2-81); 6381, 

6635 (6-6.1); 6709 (6-3b); 7088 (2-17) 

Kruse, H., 104 (2-45); 261 (2-9); 625 (2-16); 1256 (2- 

9); s.n. (6-47) 

Kubitzki, K., 75-53 (2-96b); 75-54 (6-44); 75-85 (6- 

3b); 79-241 (7-25); 21726 (2-140); 85-30 (3-3) 

Kuhlmann, J. G., 154 (3-4); 208 (2-146). 

Kuhlmann, M. et al., 58 (6-4); 92 (6-la); 161 (6-lc) 

Labroy, s.n. (6-2); s.n. (6-9) 

Lacerda, P. et al., 95 (7-56b); 137 (2-52) 

Laguna, A., 16, 41 (7-36a) 

Lamonica Freire, E. M. de, 15 (7-66) 

Landrum, L. R., 4114 (7-85) 

Lane, F., s.n. (7-54) 

Lanna Sobrinho, J. P., 280 (6-25); 346 (6-16); 385 (7- 

54); 642 (2-70); 686, 890 (3-4) 

Lao, E. et al., 10 (2-95) 

Lao Magin, R., 61, 62 (2-39); 72 (7-75); 103 (2-45); 

5059 (2-43c) 

Laughlin, R. M., 292 (2-45) 

Lawesson, J. E., 43446 (6-36) 

L.B.B. (Lands Bosbeheer Suriname), 11049 (7-15) 

Leeds, A. N., 432 (1-1) 

Leeuwenberg, A. J. M., 11837 (2-69a) 

Leitao-Filho, H. F., 2074, 2091, 2161, 5742 (2-41); 

7980 (7-20); 8281 (2-41); 12476 (7-54); 13118 (7- 

85) 

Leite, I. et al., 14 (6-44) 

Lems, K., 5250 (2-130); 5272 (2-69a); 5373 (3-1); 

64021702 (2-135); 650127 (7-61) 

Lent, R. W., 2258 (2-88); 2319 (7-36c) 

Le6n, H., 632 (2-45) 

Lescure, J. P., 307 (7-56a); 390 (7-50b); 708 (7-18); 

758 (2-65) 

Lewis, G. P. et al., 1142 (6-20); CFCR6988 (4-4) 

Lewton, F. L., s.n. (2-1) 

Liesner, R. L. et al., 370 (7-36a); 3016 (2-15); 3374 (7- 

8); 3580 (6-lc); 3591 (7-21); 3654 (2-69c); 3678 (7- 

62); 3897 (6-36); 3940 (7-6); 3948 (7-40); 3970 (7- 

5); 4052 (6-lc); 4104 (2-69a); 4191 (2-140); 5223 (7- 

36a); 5572 (2-27b); 5780 (7-25); 5876 (2-77); 5885 

(7-56a); 5921 (2-77); 5957 (7-25); 6087 (7-21); 6130 

(7-62); 6378 (7-5); 6492 (6-36); 6530 (2-140); 6716 

(7-62); 6903 (2-68); 6917, 7014 (2-69b); 7069 (6- 

lb); 7136 (2-69b); 7209 (6-lb); 7236 (7-49); 7350 

(2-39); 7451 (7-5); 7581 (7-21); 8466 (7-62); 8530 

(7-5); 8550 (7-21); 8615 (2-69a); 8621, 8638 (2-24); 

8715 (7-40); 8742 (7-21); 8851 (2-69b); 8875 (2- 

112); 8907 (2-27b); 8913 (6-13); 8917 (7-40); 9050 

(7-55); 9107 (8-3); 9114 (2-68); 9143 (2-24); 9177 

(7-36a); 9361 (2-43b); 9392 (7-22); 9434 (7-36a); 

9507 (2-68); 9538 (7-56a); 9563 (6-3b); 9580 (7-56); 

9584 (2-76); 9606 (2-23.1); 9633 (7-102); 9887 (2- 

114); 10404 (7-56a); 10424 (7-56b); 11009 (2-43a); 

11027 (7-56b); 11085 (2-122); 11095 (7-36a); 11170 

(7-56a); 11195 (2-13); 11219 (7-25); 11293 (7-25); 

11343 (2-122); 11345 (7-20); 11442, 11455 (7-56a); 

12437 (3-7); 12471 (7-36a); 13063, 13354 (7-22); 

13339 (2-43a); 13660 (7-36a); 13990 (2-68); 14051 

(2-129); 14060 (2-77); 14070 (7-73); 14445 (7-36a); 

16615 (7-30); 16624 (2-68); 17107 (3-1); 17231 (7- 

56b); 17576 (7-25); 17847 (7-73); 17927 (2-83); 

18344 (7-21); 18626 (7-43); 18666 (7-25); 19484 (2- 

69a); 19507 (2-13); 19612 (7-56a); 19724 (7-49); 

19796 (7-56a); 19852 (7-49); 19919 (1-3); 20100 (7- 

56a); 20133 (7-21); 20192 (1-3); 20517 (7-56a); 20527 

(2-56); 20562, 20627 (2-81); 20648 (2-56); 20675 

(7-30); 20744 (2-135); 20811 (7-25); 20888 (2-56); 

20943 (2-124); 20952 (2-81) 

Lima, A., 390-68 (2-141)


Lima, D. A., 65-4367 (7-50); 67-5059 (7-54); 68-5458 

(7-18); 69-5595 (7-50) 

Lima, D. P., 13083 (3-11) 

Lima, E. et al., 45 (2-81) 

Lima, H. C. de et al., 415 (2-144); 1019 (7-20); 1039 

(7-54); 1040 (7-20); 1600 (2-69a); 2178 (2-127); 2275 

(2-114); 2683 (3-4) 

Lima, J. et al., 147, 160, 208 (7-56a); 231 (2-45); 269 

(7-56a); 493 (2-18); 550 (7-56a) 

Lima, J. P. de S., 89 (2-41); 101 (2-28); 104 (7-54) 

Lima, R., 105 (2-114) 

Lindeman, J. C., 64 (7-7); 162 (3-1); 223 (6-7); 235 (7- 

73); 257 (2-88); 329 (7-73); 384 (7-56a); 479 (6-4); 

503 (7-56a); 525 (2-67); 531 (7-73); 677 (7-56a); 

7947 (2-18); LBB15315 (2-63) 

Lino, A. M., 9 (6-26); 45 (2-78.1); 71 (2-71) 

Liogier, A. H. et al., 19025, 28174 (7-36a); 28322 (7- 

83); 29196, 29296 (1-1); 31856 (7-36a); 32784 (7- 

83); 34476 (7-36a); 35577 (2-44); s.n. (1-1); 

Lisboa, P. et al., 235 (7-73); 481 (2-75); 522 (7-73); 

595 (2-114); 621 (2-141); 630 (6-3b); 712 (7-54); 

1132(3-4); 1205 (7-56b); 1377(2-24); 1382(2-121.1); 

1485 (2-27a); INPA52980, INPA53023 (7-56a); 

INPA53045 (2-43b); INPA53135 (7-56a); 

INPA53161 (2-99); INPA53215 (3-4); INPA53233 

(2-37); INPA53235 (2-30); INPA53238 (2-28) 

Little, E. L. et al., 443 (7-36); 13775, 20668 (1-1); 

21014 (3-17); 21659, 21660, 23787, 25013 (1-1); 

25020 (7-22); 25052 (2-88); 25053 (7-56b); 25058 

(7-22); 25070 (7-36a); 25101 (7-22); 25194 (7-36c); 

25307 (7-36a); 25313 (7-88); 25314 (3-36c); 25326 

(7-56b); 25386 (7-23) 

Lizot, J., s.n. (2-81); 66 (2-68); 75/46 (2-43b) 

Lleras, E. etal., P16597 (7-56b); P16639 (6-34); P16655 

(8-4); P16665 (6-3b); P16893 (6-36); P16962 (6-35); 

P16977 (2-62); P17025 (7-6); P17058, P17142 (7- 

62); P17196 (3-5); P17216 (7-95); P17270 (2-43c); 

P17294 (6-41.1); P17392 (6-16); P17397 (8-4); 

P17483 (6-29); P17488 (7-40); P17501 (2-124); 

P17508 (7-73); P17561 (7-1); P17564 (7-56a); 

P17569 (7-8); P19571 (2-57); P19665 (2-69b); 

P19676 (7-6) 

Lobo, M. G. A. et al., 3, 38, 70, 120, 163 (7-56b); 174 

(6-15); 184 (2-114); 270 (4-3); 314 (7-24); 1102 (2- 

43a) 

L6pes-Forment, W., 1356 (6-47) 

Lopes, M. A. et al., 113 (6-6.2); 176 (6261) (2-114) 

Lopez-Palacios, S. et al., 4467 (6-3b); 4479 (2-12) 

Loureiro, A. et al., s.n. (2-28); s.n. (2-29); s.n. (2-43); 

s.n. (2-69a); s.n. (2-81); s.n. (3-4); s.n. (6-3a); s.n. (6- 

4); s.n. (6-9); s.n. (6-11); s.n., s.n. (6-15); s.n. (6-28); 

s.n. (6-29); s.n., s.n. (7-73); INPA37544 (7-56b); 

INPA37629 (7-2); INPA37826 (2-27a); INPA37848 

(7-56a); INPA38099 (3-2); INPA38929 (2-28); 

INPA38985 (7-56b); INPA39510 (7-56a); INPA- 

47911, INPA47955 (2-119.1); INPA47967 (6-4); 

INPA47973 (2-27a); INPA48112 (2-57); INPA48136 

(3-4); INPA48139 (2-43b); INPA48149 (2-69b); 

INPA48160, INPA48165 (2-43b); INPA48206 (2- 

57); INPA48600 (2-140) 

Loureiro, R. L., 11 (2-18) 

Lourteig, A., 2340 (7-85) 

Lowrie, S. R. et al., 115 (7-24); 284, 302 (7-56a); 483 

(2-141); 545, 567 (7-66); 568, 706 (7-56a) 

Luetzelburg, P. von, 28349 (2-44) 

Lugo, H. S., 1039, 1054 (2-144.1); 2144, 2160, 2202, 

3230 (7-56a); 3491, 3576 (6-36); 3681, 3715, 3775 

(7-56b); 4286 (7-28) 

Lundell, C. L., 13023 (2-32.2) 

Luteyn, J. L., 8622, 8979, 8997 (6-36); 9234 (7-36a); 

11506 (7-83) 

Maas, P. J. M. et al., 386 (2-98); 391, 1757 (7-56a); 

3517, 3576 (6-15); 3656 (7-56b); 3799 (7-73); 3822 

(2-77); 3817, 3836, 3872 (7-25); 4527 (7-21); 5187 

(2-106); 5171 (7-68); 5490, 5544 (7-25) 5838 (2-56); 

P12658 (2-114); P12696 (7-73); P12762 (3-2); 

P12846 (2-22); P13041 (7-56a); P13076 (7-40); 

P13107 (7-5); P13134 (7-75); LBB10811 (2-134) 

Maasola, J., 9 (7-22) 

MacBryde, B. et al., 1376 (6-36) 

MacDougall, T., s.n., s.n., s.n. (7-56b) 

Macedo, M. et al., 150 (7-54); 235 (2-28); 282 (2-41); 

480 (2-56a); 495 (6-21); 553 (7-20); 609 (2-37); 846 

(3-15); 1169 (2-11); 1297, 1395 (7-54); 1629, 1630 

(2-28) 

Macedo, R., s.n. INPA55750 (6-50) 

Maciel, A., 124 (2-28); 128 (7-54); 326 (7-19) 

Maciel, G., 5 (1-1) 

Maciel, U. N. et al., 72 (7-56b); 83 (6-3a); 88 (7-24); 

127 (7-50a); 133 (2-27a); 166 (6-3a); 260, 303 (7- 

56b); 321 (4-1); 477 (2-69a); 717 (2-8) 

Madison, M. T. et al., 135 (7-56a); 175 (2-69b); 195 

(2-88); PFE211 (6-3b); PFE303 (7-2); 358, 650 (7- 

62); 6175 (2-69b); 6195 (2-88); 6211 (3-3b); 6303 

(7-5) 

Magnago, H., 107 (2-124); 112,230 (3-16); 272 (2-41); 

s.n. INPA58151 (2-129) 

Maguire, B. et al., 30975 (6-3b); 37707 (7-56b); 47080 

(2-36) 

Maia, L. A. et al., 26 (7-56a); 112 (2-69b); 161 (7-56b); 

189 (7-24); 238 (2-129); 458 (2-35); 486 (3-6); 570 

(7-55); 663 (7-55.1); 725 (2-129) 

Makrinius, E., 691 (7-56b) 

Malpica, A. A., 1 (2-17) 

Mantovani, W., 993 (6-27) 

Marcano Berti, L. A. et al., 119 (3-4); 211 (2-99); 284 

(2-77); 575 (2-12); 652 (2-88); 682, 687, 792 (2-77); 

861 (7-36a); 896 (7-22); 918 (7-36a); 1523 (3-7); 

2603 (3-4); 2521 (6-3b); 2617 (2-73); 2620 (7-56b); 

2858 (2-27); 2951 (7-22); 3029 (7-36a); 21-1-77 (3- 

1); 67-1-77, 52-2-77 (2-39); 56-2-77 (7-25); 67-2-77 

(1-1); 77-2-77 (3-4); 13-3-77 (7-25); 6-10-78 (2-114); 

64-979 (2-69a); 86-979 (8-3); 164-979 (3-7); 468- 

979 (7-36a); 19-980 (6-la); 41-980 (2-27b); 54-980 

(7-56b); 58-980 (3-7); 60-980 (7-56b); 43-981 (2-68); 

47-981 (3-4); 89-981, 99-981 (2-69a); 135-981 (7- 

49); 178-981 (3-7); 200-981, 216-981 (2-43a); 290- 

981 (2-12); 982-157 (7-36a); 983-018 (2-114); 983- 

022 (3-7); 983-075 (2-43a) 

Marinho, L. R., 125 (2-27); 134 (2-64); 158 (2-26); 248 

(6-35); 267 (2-99); 382 (2-129); 395 (2-69b); 405 (2- 

57); 447 (3-6); 472 (2-69b); 473 (2-57); 502 (2-69a); 

561 (7-5) 

Marshall, S. A. et al., 6562 (1-1)


Martin, R. T. et al., 1329 (7-56a); 1380, 1672 (2-1); 

1676 (6-9); 5387 (7-56b) 

Martinelli, G. et al., 80 (6-25); 372 (7-29); 2189 (4-5); 

4114 (6-25); 6121 (2-71a); 6789 (7-42); 6859 (2- 

135); 6904 (7-25); 7052 (6-32.1); 7095 (6-3b); 7156, 

7166 (2-134); 7177 (3-4); 7226 (2-99); 7231 (2-134); 

7234 (2-99); 7244 (2-129); 7374 (6-3a); 7375 (7- 

56b); 7470 (7-54); 8466 (3-4); 8739 (7-54); 9649 (7- 

20); 9671 (6-51b) 

Martinez S., E., 1539 (1-1); 5798 (7-56b) 

Martins, H. F., 336 (2-11); 354, 414 (6-25) 

Martins, O., MG8155 (2-99) 

Martins, T., 18 (2-41) 

Martius, K. P. F. von, s.n. (3-4); 1840 (3-16) 

Mathias, M. E. et al., 6108 (7-75) 

Mattos Filho, A., 345 (7-54); 348 (7-20) 

Mattos Silva, L. A. et al., 386 (3-11); 390 (7-20); 1072 

(2-?); 1073 (6-?); 1136 (2-84); 1176 (1-1); 1196 (2- 

71); 1211 (2-80a); 1401,1402(2-84); 1440(2-130.1); 

1586 (7-20); 1700 (6-51b); 1839 (2-84); 1870 (2- 

71a); 1920 (2-88); 1984 (2-114) 

Maury, C., 277 (3-16) 

Mautone, L. et al., 118 (6-25) 

Mazzeo, P. M., 199 (2-1) 

McDaniel, S., 2660, 2496 (6-3b); 16280 (7-40); 17187 

(2-8); 20530 (6-44); 21133 (2-43c); 22346 (6-44) 

McPherson, G., 6979, 8017 (7-36a); 8213 (7-56a); 8231 

(7-36a); 8457 (7-56a); 8479 (2-15); 8577 (7-38); 8771 

(7-36a); 9015 (7-38); 9498 (2-31.1); 9612 (7-80) 

Medeiros et al., 58 (6-18b) 

Medina, E., 291 (7-40); 322 (2-27a); 533 (2-119); 542 

(2-103) 

Meehan, s.n. (1-1) 

Meijer, W. et al., VEN92 (6-48) 

Meijeraan, J. W., 19 (7-7) 

Mejia, M. et al., 6254, 6581, 10197 (1-1); 11174 (7- 

36a); 23689 (1-1) 

Melin, D., 41 (6-3a); 173 (2-27) 

Mello, F. et al., 2 (6-11); 16 (2-69a); 24 (7-50b); 25 (3- 

4); 43, 45 (2-64); 48 (2-69a); 55 (7-41); 71 (7-24); 

114 (7-56b); s.n. INPA51803 (6-3b); INPA55359 (2- 

69a); INPA53361, INPA57834 (6-50); INPA57842 

(2-39); INPA57898 (6-34); INPA57962 (6-35); 

INPA57963 (7-24); INPA57971 (2-129); INPA60170 

(2-69.1) 

Mello Filho, L. E., 3061 (6-32); 3618 (7-54) 

Mello Silva, R. et al., CFCR7548 (7-18); CFCR8313 

(7-54); CFCR8397 (7-70) 

Mendonsa, S., 2 (7-73) 

Meneces, E. et al., 340 (3-5); 669 (7-50b); 706 (2-99); 

2061 (2 sp.) 

Meneces, S., 360 (2-21); 669 (7-50a) 

Mennega, A. M. W. et al., 882 (2-135) 

Mercado, R., 9 (6-47) 

Meredith, H. B., s.n. (2-1); s.n. (1-1) 

Meyer, G., 46 (6-23); 78 (2-43b); 105 (7-66); 202 (7- 

56a); 294 (2-43b) 

Miers, H., RB179792 (7-85) 

Mileski, E., 66 (7-66); 155 (6-3c); 168 (7-54); 171 (2- 

27a); 187 (6-3c); 350 (6-3a) 

Miralha, J. M. S., 49 (7-2); 63 (7-8); 67 (2-135.1); 74, 

84, 87, 92 (2-43a) 

Miranda, C. A., 168 (2-80); 199 (4-2) 

Miranda, F. E. et al., 299 (2-134); 309 (7-56a); 378 (2- 

136); 407 (7-30); 491 (2-69a); 539 (7-56b); 595 (6- 

lb); 614 (7-24); 700 (7-56b) 

Mocquerys, 1021 (1-1) 

Molina, D., 139 (7-22) 

Molina, R., B-3 (2-18); 31 (3-6) 

Molino, I., G416 (1-1) 

Monsalve B., M., 152 (2-124); 481 (2-148); 445, 467 

(2-46); 523 (3-15); 651 (2-148); 758 (2 sp.); 805 (2- 

148) 

Montalvo, E A., 4450 (7-56b) 

Monteiro, O. P. et al., 19 (2-124); 29 (2-33.1); 32, 33 

(2-119.1); 45 (1-1); 127 (6-3b); 133 (2-39); 162 (2- 

88); 187 (7-56a); 375 (2-26); 412 (7-56b); 454 (6- 

36); 516 (6-35); 532 (7-2); 779 (2-43b); 781 (7-56a); 

803 (7-24); 804 (2-75); 877 (2-43b); 895 (7-56a); 922 

(6-34); 925 (6-9); 931 (2-43b); 936 (2-62); 945 (2- 

142); 1032 (7-50b); 1059, 1067 (2-43b); 1071 (3-4); 

1245 (7-1); 1270 (6-44); 1291 (2-27); 1363 (2-62); 

1419 (2-69b); 1458 (6-3b); s.n. INPA50017 (6-14); 

s.n. INPA50852 (2-43b); INPA50875 (2-69a); s.n. 

INPA53419 (7-56a); INPA53534 (7-1); s.n. INPA- 

50017 (6-14); s.n. (6-34) 

Montouchet, P., 2202 (7-56b) 

Mora, L. E., 2515 (1-1); 4424 (6-36) 

Moreno, P. P. etal., 7263, 12059b, 12072 (1-1); 12173, 

14570, 14612 (7-56b); 15205 (1-1); 16078B (7-22); 

22530, 22830, 22876, 22957 (2-45); 23014, 23072 

(7-23); 23130 (7-36c); 23166 (2-88); 23233 (7-36c); 

23258 (7-23); 23449 (2-45); 23567 (7-22); 23748 (7- 

36a); 23994 (7-22); 24061 (7-36a); 24627, 24980 (7- 

56b); 25541 (7-16); 25576 (7-56a); 25614 (7-16) 

Moretti, C., 1264 (3-4); 4510 (6-7) 

Mori, S. A. et al., 2075, 2370 (7-38); 2876 (7-56a); 

3653 (7-38); 3741 (6-13.1); 4091 (7-56a); 4188 (2- 

95); 4665 (2-31.2); 4914 (2-31.2); 5052 (2-18.2); 5148 

(7-38); 5171 (2-88); 5306 (7-36a); 5543 (7-56a); 6532 

(2-31.1); 7116 (6-13.1); 7778 (2-18.1); 7990 (2-31.1); 

8015 (2-66); 8048 (2-135); 8064 (6-15); 8081 (2-89); 

8088 (2-90) 8107 (2-109); 8131 (2-90); 8140 (7-25); 

8158 (3-1); 8181 (2-39); 8217 (2-109); 8218 (2-113); 

8224 (7-56a); 8231 (2-77); 8306, 8330 (2-135); 8401 

(2-108); 8459 (7-67); 8532 (6-7); 8536 (7-56a); 8573 

(6-2); 8590 (7-2); 8672 (7-56a); 8678 (4-3); 8862 (2- 

112); 8880 (2-69a); 9002 (7-24); 9003 (2-129); 9008 

(6-29); 9009 (2-129); 9010 (7-40); 9015 (6-3b); 9065 

(6-44); 9115 (7-40); 9234 (6-44); 9336 (6-17); 9598 

(6-25); 9757 (1-1); 10241 (2-146); 10243 (3-3.1); 

10252 (2-146); 10470 (7-31); 10871, 10934 (6-51b); 

10936 (7-37); 11033 (2-146); 11037 (6-51b); 12001 

(3-11); 12246 (7-20); 12354 (7-70); 12749 (6-52.2); 

13063, 13673 (2-130.1); 13774 (2-124); 13819 (2- 

144); 13884(2-124); 13989(2-88); 13995(2-6); 14000 

(2-124); 14017 (6-17); 14575 (1-1); 14718 (2-109); 

14763 (2-69a); 14764, 14772(2-27.1); 14780(2-72); 

14782 (2-43a); 14783 (7-39); 14790 (2-27.1); 14791 

(2-43a); 14814 (2-99); 14821 (6-7); 14828 (2-27.1); 

14830 (2-99); 14835 (7-73); 14870 (7-56a); 14897 

(7-74); 14963 (7-39); 14978 (2-69a); 15160 (3-2); 

15161 (2-149); 15201 (2-69a); 15230 (7-39); 15232 

(2-112); 15244 (2-86); 15254(2-112); 15285 (7-20);


15299 (2-69a); 15403 (1-112); 15411 (2-38); 15428 

(2-27.1); 15435 (2-27a); 15461 (2-108); 15476 (2- 

27.1); 15501 (2-99); 15523 (2-27.1); 15650 (2-8); 

15796 (7-42); 15802 (2-99); 15822 (2-112); 15840 

(2-76); 15855 (7-50b); 15859 (6-19); 15879 (7-24); 

15902 (2-124); 15944 (2-69a); 16006 (2-17); 15975, 

16007 (6-50); 16038, 16089 (7-42); 16120 (2-88a); 

16143 (7-42); 16172 (2-39); 16177, 16180 (7-42); 

16181 (2-108); 16214 (7-42); 16228 (2-99); 16234 

(2-88a); 16290 (6-50); 16313 (2-99); 16312 (2-124); 

16316 (7-24); 16323 (6-50); 16349 (2-69a); 16398 

(6-50); 16402 (7-24); 16414 (2-43a); 16424 (7-42); 

16444 (7-24); 16468 (7-42); 16476 (2-99); 16516 (2- 

27a); 16528(2-63); 16762(7-20); 16792 (3-16); 16842 

(7-20); 16843 (7-51); 17186 (6-7); 17193 (2-141); 

17201 (7-2); 17216 (2-64); 17219 (6-15); 17224 (2- 

69a); 17238 (7-2); 17244 (3-12); 17245 (7-50b); 

17290, 17332, 17380, 17422 (2-69a); 17476 (2-39); 

17500 (7-24); 17547 (2-109); 17553 (2-99); 17578 

(2-69a); 17579 (7-20); 17606 (2-39); 17658 (7-20); 

17707 (2-109); 17713 (7-20); 17731 (2-86); 17961 

(7-39); 17977 (2-27c); 17980 (2-109); 18012(2-27.1); 

18049 (2-99); 18080 (1-13); 18116 (7-50a); 18186 

(2-69a) 

Morillo, G. et al., 3460 (7-55); 3650 (2-99); 5121 (2- 

69a); 5135 (7-21); 5314 (7-5); 5436 (7-21); 6948 (6- 

3b) 

Mosier, C. A. et al. 49 (1-1) 

Mota, C. D. et al., 86 (2-22); 89 (2-62); 138 (6-3b); 

191 (2-37); 246 (7-5); 739 (2-124); s.n. INPA60383 

(2-141); INPA s.n., INPA60396 (2-69a); INPA60444, 

INPA60480 (7-56b); INPA60611 (7-56a); INPA- 

60614, INPA60635 (6-29); INPA60656 (6-13); 


(6-14); INPA60732 (7-50a); INPA60869 (7-56a); 


(6-29); INPA61336 (7-30); INPA61341 (6-14); 

INPA61558 (7-25); INPA61609 (2-62); INPA61613 

(2-36); INPA61659 (8-1); INPA61672 (6-3a); s.n. 

(6-29); s.n. (7-56a) 

Museu Goeldi, Belem, 9628 (2-57) 

Narvaez, E., 633 (1-1) 

Nascimento, O. C. et al., 20 (6-44); 29 (6-28); 191 (2- 

69b); 250 (6-2); 259 (2-69b); 279 (7-21); 340 (2-79); 

352 (7-24); 363 (2-79); 453 (7-56a); 560 (6-3b); 571 

(7-56b); 604 (2-69b); 611 (2-88a); 811 (6-lb); 867 

(7-20); 875 (2-43a) 

Nee, M. et al., 7675 (5-1); 11288 (7-80); 17773 (2-81); 

24631 (2-15); 27960 (2-45); 28191 (6-47); 28329 (2- 

15); 28850 (1-1); 30848, 30999 (2-26); 31863 (6-3b) 

Nehlin, S. 0., s.n. (7-33) 

Neill, D., 1794 (7-36a); 1896 (7-2); 2829, 3103, 3136 

(2-45); 3755 (7-23); 3795 (7-22); 4044 (7-23); 4104 

(7-22); 4124 (7-56b); 4174 (7-36a); 4175, 4318 (7- 

22); 4338 (7-23); 4508 (7-56b); 4566 (1-1) 

Nelson, B. W. et al., 344 (6-27); 630 (3-5); 804 (2-27b); 

807 (2-69a); 862 (7-73); 871 (6-la); 913 (2-140); 931 

(2-27a); 933, 939 (2-140); 956 (2-27a); 1058 (2-69e); 

1075 (7-56b); 1185 (6-3a); 1401 (7-56b); 2617 (2- 

1); P21071 (2-57) 

Nelson, C. et al., 4253 (7-56b) 

Nelson, E. B., 4481, 4533 (1-1) 

Nepomuceno, V., HPB2489 (2-43a) 

Nevers, G. de et al., 3816 (7-38); 4446 (2-39); 4618 

(1-1); 4773 (7-56b); 5106 (7-80); 5440 (2-5); 5767, 

5840 (2-95); 6446, 6558 (7-56b); 7112 (2-145); 7124 

(2-88a); 7553, 7581, 7603 (5-1); 7726 (9-1) 

Nevling, L. R. et al., 2604 (7-56b) 

Noblick, L. R., 1622, 2164 (7-18); 2864 (7-20); 3088 

(7-18) 

Nunes, E., HPB5947 (4-2) 

Obando, R., 17 (1-1) 

Oldeman, R. A. A. et al., 11 (2-4); 198 (2-27a); 291 

(7-2); 1650, 1672 (7-56a); 1839 (6-18); 1871 (2-97); 

1954, 2166 (2-99); 2237 (2-69a); 2773 (7-2); 2868 

(7-56a); 3097 (2-134); 3223 (7-2); B492 (7-73); B551 

(2-52); B898 (7-2); B1298 (2-69a); B1712 (2-97); 

B1734A(7-50b); B1771 (7-56a); B1875 (7-74); B2031 

(2-69a); B2514 (6-18); B2524 (7-50b); B2676 (6-7); 

B2668 (2-129); B2688 (2-26); B3389 (6-2); B3458 

(7-56a); B3464 (2-134); B3507 (7-56b); B3538 (2- 

98); B3993 (7-2); B3999 (7-73); B4065 (2-86); B4353 

(2-8); T226 (2-97); T280 (2-69a); T351 (2-97); T370 

(7-50b); T447 (7-79); T486 (2-27a); T593 (2-27b); 

T597, T648 (2-27c); T726 (2-124); T741 (6-2); T761 

(2-69a); T919 (7-2); T933 (6-7) 

Oldenburger, F. H. F. et al., 99 (7-56); 232 (7-20); 479 

(2-122); 483 (2-108); 620 (7-56b); 961 (2-39); 1179 

(2-112); 1213, 1225 (2-108.1); 1244 (7-56a); 1256 

(2-100); 1416 (7-50a) 

Oliveira, A. R. de, INPA58658 (6-4); INPA58700 (2- 

99); INPA59593 (6-9); INPA59622 (2-27); 

INPA59668 (2-43b); INPA59761, INPA59774 (2- 

62); INPA59825 (6-15); INPA59877 (2-124); 

INPA59908 (6-9); INPA59936 (2-69.1); INPA59967 

(2-27); INPA59901 (2-69b); INPA60162, INPA- 

73491 (2-69.1) 

Oliveira, E. de, 302 (6-lc); 455 (4-3); 2130 (6-1b); 2655 

(6-la); 3586, 4054 (2-69a); 4343 (2-63); 4471 (2- 

69a); 5111 (7-56b); 5666 (3-1); 5617 (6-la); 5639 

(6-1c); 5709 (2-124); 5721 (6-lb); 5840 (6-50); 5962 

(2-52); 6217, 6230 (7-56b); 6283 (2-69a); 6379 (7- 

56b); 6494 (2-52) 

Oliveira, H. B., 9(2-82) 

Oliveira, M., 3017 (2-69); 3792 (2-43); 3922 (2-99); 

3973 (2-136); 4909 (7-20); 5325 (7-56b); 5378 (2- 

52); 6210 (2-43) 

Oliveira, M. C. C., 2 (6-25) 

Oliveira, P. E., 3 (7-54) 

Oliveira, P. I., 508 (4-2) 

Oliveira, R. F. de, 485 (1-1) 

Ongley, J. C., P21739 (2-60) 

O'Neill, H., 8625, 8626, 8627 (7-56b) 

Oren, D. C., 3 (7-30) 

Orlandi, R. P., 66 (2-80); 599 (4-2) 

Ortega U., A., 33 (2 sp.); 73, 74 (2-88); 76 (2-27a); 110 

(2-144.1); 146 (6-36); 154 (2-27a) 

Ortega, F., 565 (1-1); 768 (2-12); 1691, 2517 (7-36a) 

Ortiz, F., 1383 (7-23); 1974 (7-22) 

Ortiz, R. T., 592, 1024 (7-56b); 2638 (7-22) 

Osmarino (P. Monteiro) et al., s.n. (6-4); s.n. (6-44) 

Pab6n E., M., s.n. (7-30) 

Pabst, G., 5219 (3-4); 9559 (2-44) 

Palacios, W. et al., 827 (6-36)


Paray, L., 523, 1817 (7-56b); 1944 (7-36a); 2017 (6- 

47) 

Passos, B. C. dos, 1073 (7-20); 1082 (6-3b) 

Paula, J. Elias de, 684 (7-20); 1180 (1-1); 1497 (2-11); 

1725 (6-20) 

Pedrosa, J. S. et al., 1107 (1-1) 

Pena, B. S., 73 (7-56b); 82 (3-4); 99 (2-52); 184 (7-24); 

242 (7-50b); 262 (7-50a); 324 (2-69a); 338 (7-25); 

409 (6-3a); 464 (2-27a); 484 (2-37); 486 (7-54); 542 

(6-la); 688 (2-62); 2002 (7-20) 

Pennington, T. D. et al., 10152 (2-45); 10687 (6-16); 

21675 (7-8) 

Pereira, B. A. S., 59, 64 (6-27); 318 (2-43a) 

Perez, J. L. A. et al., 4837 (7-56a) 

Perez, P., s.n. (2-3) 

Peters, C., 39 (6-44); 188 (6-16); 60-84 (6-41.1); 84- 

11 (6-44) 

Phelps, K. D., 511 (7-43) 

Philcox, D., 3573 (2-17); 7774 (1-1); 7966 (7-25) 

Philipson, W. R. et al., 2005 (2-76) 

Pic6n, G. et al., 914 (7-25); 1045 (7-49) 

Pinheiro, R. S., 448 (2-11); 1016 (2-36a); 1177 (3-36a); 

1492 (2-11); 1558 (2-84); 1948 (3-11); 2078 (2-138); 

2120 (7-36b) 

Pinto, G. C. P., 176/81 (4-2); 54/82 (7-54) 

Pinto, P. et al., 828 (7-40); 1172 (2-69b); 1244 (7-56b); 

1256 (2-69b); 1269, 1317, 1344 (7-56b); 1345 (2- 

69b); 1472 (7-56b); 1574 (2-69b); 1674 (7-22); 1902 

(1-1) 

Piniate, P. et al., s.n. (6-36); 1017 (3-6) 

Pio, L. C., 13 (6-27) 

Pipoly, J. J., 4535, 4692, 4795 (7-22); 4992 (7-56b); 

6724 (2-39); 6730 (6-15); 6743 (4-3); 6751 (2-122); 

6758, 6778 (2-43a); 6780 (2-102); 6788 (6-29.1); 

6798 (6-7); 6823 (2-57); 6828 (2-17); 6892 (2-122); 

7372 (2-135); 7356 (6-15); 7360 (6-42a); 7374 (6- 

15); 7426 (6-42a); 7440 (2-90); 7446 (7-56a); 7497 

(2-93); 7507 (2-109); 7513, 7524, 7537 (2-87); 7701, 

7760 (2-135); 7820 (7-21); 7828, 7927 (2-135); 7941 

(7-21) 

Pirani, J. R. et al., 1216 (7-20); 1217 (2-37); 1250 (2- 

114); 1292 (2-37); 1341 (3-16); 1352 (2-27a); 

CFCR461, CFCR864 (7-20); CFCR877, CFSC7971 

(7-54); CFCR8277 (6-27); CFCR8282 (7-20) 

Pires, J. M. et al., 74 (3-4); 108 (7-1); 184 (2-69a); 185 

(2-64); 197 (6-3a); 210 (6-lc); 290 (6-34); 1622 (6- 

3c); 2914 (2-144); 3195, 3544 (7-40); 4146, 4923 (6- 

la); 5293 (2-98); 5409 (2-63); 5864 (2-36); 5978 (2- 

98); 6326 (2-17); 7124 (3-3); 7184 (2-69a); 7700 (7- 

79); 7925 (6-9); 10302 (6-la); 10543 (2-99); 10695 

(4-3); 10755 (2-69a); 10781 (4-3); 10844 (2-69a); 

10938 (2-63); 11005 (2-69a); 11012 (2-63); 11014 

(6-la); 11029 (2-69a); 11055, 11065, 11066, 11137 

(2-63); 11265 (2-69a); 11296 (2-52); 11359 (2-8); 

11484(4-3); 11786(2-8); 11903 (3-2); 11931 (6-la); 

12189 (2-69a); 12196 (2-18); 12214, 12245 (7-36a); 

12394 (7-56a); 12556 (2-81); 12573 (7-36a); 12581 

(7-56a); 12676 (2-23); 13077 (6-36); 13120 (6-50); 

13149 (3-1); 13291 (2-86); 13295 (6-4); 13342 (7- 

18); 13649 (6-4); 13737 (2-8); 13739 (7-36a); 13868 

(2-99); 13915 (4-1); 13921 (2-27a); 13952 (6-7); 

13957 (2-140); 13970A (4-1); 13972 (2-69b); 13977 

(6-3); 13991 (2-140); 14008 (3-6); 14022 (4-1); 14044 

(2-27a); 14059 (6-7); 14064 (2-141); 14072 (2-57); 

14088 (2-75); 14157 (2-88); 14172 (2-93); 14192 (7- 

49); 14206, 14217 (2-88); 14239 (6-7); 14249 (3-4); 

14272 (2-93); 14360, 14375 (7-4); 14440 (6-3b); 

14508 (3-1); 14542 (2-69a); 14548 (2-88); 14597 (3- 

1); 14658 (6-3); 14755 (2-27a); 14760 (4-1); 14761 

(7-25); 14738 (2-69b); 14783 (2-69a); 14849 (7-4); 

14910(2-114); 15027 (2-126.1); 15939(7-20); 16135 

(7-18); 16215 (6-3c); 16414 (7-66); 16560 (3-4); 

16641 (6-27); 16713 (2-27b); 16811 (2-18); 16832 

(3-4); 16930 (2-18); 16939 (7-21); 16967 (7-54); 

17066 (7-66); 17135 (2-30); 50346 (6-7); 50930 (6- 

10) 

Plowman, T. et al., 7003 (2-69b); 7016 (7-40); 7699 

(3-7); 7736 (2-27b); 7739 (7-56a); 7784 (7-56b); 9591 

(7-63); 9674 (7-56b); 9837 (7-63); 10001 (3-16); 

11371 (7-54); 11452, 11466 (7-56a); 12224 (7-5); 

12390 (2-69a); 12397 (2-129); 12414(2-124); 12448 

(7-73); 12457 (2-129); 12458 (6-3b); 12466 (3-4); 

12489 (7-40); 12525 (2-99); 12533 (7-56); 12725 (7- 

20) 

Poole, J. M., 1679 (2-69b); 1797 (2-88); 1798 (6-lb); 

1804 (2-88); 1855 (7-56a); 1995 (2-27a); 2092 (3-6); 

2094 (2-129); 2118 (2-124) 

Popenoe, J., 103 (6-47); 161, 242 (1-1); 718 (2-1) 

Porter, D. M., 4078 (7-36a) 

Poterima, L. B., 6385 (3-4) 

Prance, G. T. et al., 3653 (7-67); 8821 (3-4); 10492 (2- 

77); 10614(7-73); 10675, 10694(3-4); 10883 (2-18); 

11144, 11323 (7-56a); 11460 (2-18); 11471 (4-1); 

11480 (2-27a); 11492 (7-56a); 11506 (6-3); 11513 

(2-27a); 11585 (6-3a); 11631 (6-29); 11793 (3-2); 

11842 (7-5); 11883 (2-43); 11939 (2-94); 11943 (6- 

28); 12058 (6-35); 12079 (6-36); 12151 (7-56b); 

12227 (2-99); 12228, 12344 (7-73); 12345 (7-56b); 

12560 (6-36); 13336 (7-56a); 13348 (2-88); 13349 

(6-44); 13353 (2-36); 13404 (3-4); 13419 (7-56a); 

13462 (7-56b); 13507 (2-27a); 13638 (2-77); 13693 

(3-4); 13732 (7-56a); 13849 (7-6); 13868 (7-40); 

13882 (7-6); 13883 (7-56a); 13925 (6-29); 13963, 

13975 (2-69b); 13990 (7-56a); 14015 (8-4); 14024 

(2-43b); 14070 (2-81); 14077 (7-6); 14079 (7-40); 

14092 (2-17); 14095 (6-34); 14155 (7-40); 14158 (2- 

94); 14159 (2-62); 14169 (2-17); 14172 (2-99); 14178 

(2-43b); 14194 (2-88); 14252 (2-17); 14276 (2-43b); 

14320 (2-99); 14324 (7-56b); 14374 (6-4); 14388 (2- 

56b); 14644 (2-69a); 14719, 14720 (1-1); 14735 (2- 

69.1); 14755 (7-36a); 14787 (6-3a); 14839 (6-29); 

14924 (6-3b); 14931 (2-27a); 14942 (6-35.1); 14981 

(7-2); 14987 (2-27a); 14995 (2-36); 15001 (2-88); 

15012 (6-15); 15013 (2-69a); 15018 (2-88); 15026 

(2-98); 15030 (2-69a); 15043 (2-99); 15053 (6-3a); 

15150 (2-69a); 15116 (6-3b); 15134 (7-24); 15166 

(7-56a); 15265 (2-77); 15271 (7-56b); 15274 (6-2); 

15336 (2-69a); 15388 (7-62); 15397 (2-77); 15426 

(2-24); 15433 (6-3b); 15463 (2-88); 15503 (2-93); 

15505 (4-1); 15604 (7-72); 15745 (2-52); 15820 (7- 

62); 15935 (6-34); 16014 (6-2); 16035 (6-5); 16040 

(2-99); 16048 (7-62); 16055 (2-22); 16141 (7-41); 

16143 (6-41); 16167 (2-88); 16196 (4-1) 16215 (2- 

27a); 16229 (2-69b); 16315 (6-23); 16368 (6-37);


16401 (7-56); 16408 (2-39); 16452 (7-62); 16518 (6- 

44); 16521 (6-41); 16524 (2-43b); 16526(7-2); 16529 

(6-29); 16540, 16542 (1-1); 16545 (2-1); 16548 (7- 

21); 16562(6-42a); 16563(6-15); 16564(2-78); 16772 

(7-36a); 17519 (6-4); 17520 (2-79); 17541 (2-69a); 

17575 (7-2); 17750 (7-1); 17751, 17800 (2-69a); 

17816 (2-119); 17820 (2-79) 17828 (7-73); 17897 

(2-124); 17948 (2-82); 17950(2-60); 17985 (2-27a); 

18000, 18001 (2-69a); 18051 (6-29); 18237 (7-56a); 

18301 (2-69a); 18304 (6-3b); 18310 (2-17); 18727 

(7-1); 18739(6-4); 18747 (6-9); 18759 (6-15); 18840 

(6-27); 18920 (3-16); 19040 (2-114); 19061 (7-54); 

19078 (7-20); 19161 (2-144); 19329 (2-37); 19361 

(7-66); 20012 (2-68); 20199 (7-56a); 20555 (7-6); 

20591 (2-69b); 20592 (2-28); 20611 (2-69a); 20626 

(6-3b); 20633 (2-88); 20729 (6-11); 20768 (6-15); 

21006 (6-16); 21007 (2-85); 21032 (7-8); 21629 (7- 

6); 21636 (6-35.1); 21639 (7-8); 21652 (6-29); 21653 

(7-56a); 21661 (7-2); 21670 (2-114); 22611 (2-64); 

22622 (6-41); 22658 (7-56a); 22682 (7-73); 22693 

(2-39); 22721 (2-22); 22775 (2-64); 22807 (2-121); 

22822 (2-17); 22832 (2-99); 22843 (2-21); 22850 (2- 

124); 22858 (2-43b); 22939 (2-28); 22972 (2-33); 

22977 (7-56b); 22986 (2-43b); 23028 (2-64); 29997 

(2-17); 23131 (4-1); 23253 (2-26); 23366 (7-2); 23379 

(6-11); 23395 (6-9); 23481 (6-29); 23505 (2-26); 

23508 (7-9); 23548 (7-56b); 23595 (6-9); 23601 (7- 

1); 23762 (6-35); 23789 (7-62); 23816 (6-35); 23826 

(7-41); 23860 (7-56a); 23938 (6-7); 23971 (6-29); 

23974 (7-40.1); 24123 (7-56a); 24163 (7-62); 24166 

(7-40); 24177 (7-62); 24215 (2-27a); 24216 (4-1); 

24238 (7-56b); 24251 (2-124); 24257 (7-67); 24268 

(2-36); 24269 (6-41); 24287 (6-4); 24297 (4-3); 24539 

(2-129); 24574 (7-2); 24731 (7-63); 24759 (2-28); 

24800 (2-141); 24875 (2-75); 24895 (7-20); 25001 

(7-50a); 25065 (2-57); 25087 (2-81); 25140 (2-75); 

25190 (7-20); 25264 (2-75); 25190 (7-20); 25264 (2- 

17); 25531 (2-69a); 25574 (4-3); 25575 (2-27b); 25642 

(2-39); 25652 (2-14.1); 25677 (2-99); 25789 (7-2); 

25843 (2-69a); 25885 (4-3); 26127 (2-43b); 26130, 

26190 (2-28); 26219 (2-43b); 26302 (6-20); 26318 

(6-3b); 26319, 26323 (4-1); 26324 (6-3a); 26327 (4- 

1); 26343 (6-21); 26352,26359 (2-27a); 26372,26509 

(7-36a); 26564 (2-8); 26630 (6-16); 27992 (7-56b); 

28053 (2-51); 28076 (1-1); 28134 (2-11); 28154 (1- 

1); 28078 (7-2); 28100 (7-74); 28154 (1-1); 28510 

(7-56b); 28171 (2-145); 28373 (7-21); 28376 (7-25); 

28399 (7-21); 28448 (2-56); 28453 (2-69b); 28472 

(7-56b); 28693 (2-23); 28762 (2-114); 28868 (6-29); 

28869 (7-49); 28875 (2-93); 28903 (2-57); 28908 (4- 

3); 28919 (2-43a); 28941,29000 (7-30); 29037,29091 

(2-135); 29098 (2-69a); 29137 (2-135); 29163 (7-49); 

29181 (2-69a); 29261 (6-29.1); 29331 (6-57); 29357, 

29389 (7-83); 29420 (6-29.1); 29443 (6-la); 29469 

(7-62); 29513 (6-9); 29663 (2-69a); 29675 (6-29a); 

29715 (6-9); 29721 (7-49); 29738 (2-37); 29741 (2- 

36); 29749 (6-29.1); 29766 (7-49); 29768 (6-29a); 

29809, 29815 (2-119a); 29834 (2-131); 29843 (6- 

29.1); 29856 (4-1); 29877 (2-39); 29913 (2-18); 29931 

(2-140); 29947 (7-56a); 29941 (2-88); 30011 (2-140); 

30053 (7-25); 30110 (7-36a) 

Prevost, M. F. et al., 311 (2-56a); 343 (7-25); 355 (2- 

109); 358 (2-86); 378 (7-56a); 393 (2-114); 437 (2- 

36); 448 (2-56a); 932 (3 sp.); 1059 (2-134); 1168 (7- 

52); 1201 (2-114); 1495 (6-7); 1518 (7-52); 1574 (2- 

69a) 

Procter, J., 4737 (7-56a); 4738 (2-135) 

Pruski, J., 1511 (1-1) 

Pulle, A. A., 182 (3-1) 

Pulle-Lutz, 1130 (7-36b) 

Pyron, J. H. et al., 3145 (2-1) 

Quintero, A., 2204 (7-36a) 

Rabelo, B., 152 (2-43a); 164 (3-4); 230 (2-36); 605 (7- 

18); 1528 (7-25); 1827 (3-4); 1839 (2-39); 2181 (7- 

56a); 2193 (2-39); 2249 (3-4); 2275 (2-43a); 2285 

(7-52); 2380 (7-50b); 2381 (7-56d); 2475 (2-18); 2477 

(7-24); 2478,2480,2482 (2-43a); 2486 (7-50b); 2491 

(2-114); 2660 (2-39); 2695 (3-4); 2712 (2-8); 2774 

(3-4); 2792 (6 sp.); 2885 (2-114); 2922 (7-18); 2925 

(2-69a); 2928 (7-56a); 2931 (2-69); 2938 (2-69a) 2976 

(2-43a); 2990 (2-69a); 3018 (3-2); 3020 (6-50); 3029 

(2-69a); 3038 (3-2); 3058 (2-39); 3072 (2-69a); 3128 

(2-109); 3135 (6-15); 3147 (2-69a); 3197 (6-la); 3223 

(3-2) 

Ramcharan, E. K., 438 (1-1) 

Ramia, M., 7176 (2-27b) 

Ramirez C., R., 4 (6-34); 18 (7-40); 87 (6-38); 104 (2- 

5); 1083 (7-40); 1094 (2-140) 

Ramos, J. F. et al., 90 (6-41); 138 (7-56a); 157 (2-26); 

378 (2-69a); 382 (6-11); 429, 433 (7-5); 637 (7-56a); 

646 (7-66); 651 (6-22); 755 (2-119.1); 792 (7-65); 

851 (2-17); 865 (7-56a); 877 (7-30); 914 (2-63); 940 

(3-3); 942 (2-136); 945 (2-114); 1048 (2-63); 1087 

(2-27a); 1090 (2-17); 1154 (7-56b); 1165, 1196 (2- 

69a); 1507 (7-63); 1589 (2-81); 1668 (7-63); P21800 

(6-11); P23251 (8-4); INPA62116 (7-50a); INPA2157 

(6-9); INPA62177 (7-56b); INPA62187 (7-56a); 

INPA62246 (7-56b); INPA62279, INPA62293 (7- 

24); INPA62319 (6-34) 

Ramos, R., 174 (7-22) 

Ramsey, G. W. et al., 137 (2-1) 

Rankin, J., 27 (2-64); 29 (2-69a); 137 (2-108) 

Ratter, J. A., 2074, 2163 (2-41); 3232 (2-30); 3557 (7- 

66); 3619 (6-27); 3739 (3-16); 3845 (2-4); 3941 (2- 

30); 3956 (2-28); 4439 (2-27a); 4441 (2-30); 5076 

(2-18) 

Rau, E. A., s.n. (2-1) 

Ravelo, O., 33 (2-113) 

Reeder et al., LBB12309 (2-69a) 

Reis, G., 149 (2-43a) 

Reis, L. Q., s.n. INPA57832, s.n. INPA57833 (6-50) 

Reitz, R. et al., 18086 (3-4) 

Renteria A., H., 1460 (1-1) 

Renteria, E. et al., 2186 (2-56a) 

Revilla, J., 171 (7-40); 359 (2-27a); 383 (2-27); 402 

(6-3b); 406 (2-37); 479, 580, 676 (6-44); 690 (2-69b); 

698 (6-3b); 816 (6-14); 1120 (7-56b); 1167 (6-6a); 

1483 (7-5); 1666 (2-17); 1814 (7-40); 1843 (7-56b); 

1903 (2-140); 2138 (6-la); 2218, 2247 (2-140); 2345 

(2-94); 2421 (6-44); 2465, 4071, 4075 (6-3b); 4156 

(4-1); 4537 (7-56a); 7062 (2-22); 7086 (2-128); 7106 

(2-57); 8372 (7-24); 8380 (2-69a); 8414 (2-17); 8416 

(2-67); 8421 (2-136); 8433 (2-63); 8434 (2-17); 8529, 

8531 (7-56b); 8666 (6-21)


Reyna R., N., 31, 132-I (2-128) 

Reynel R., C., 366 (2-21); 422 (7-73); 444 (7-36a); 498 

(3-9); 558 (2-27a); 566 (7-41); 643 (2-43b); 653 (7- 

41); 695 (2-45); 850a (7-56a); 967 (7-36a) 

Riba, R. et al., 292 (2-15) 

Ribanov, J. et al., 151 (2-43b); 169, 205 (2-108); 218, 

245 (2-88); 268 (7-56a); 277 (7-50b); 281 (2-108); 

295, 303 (3-4); 319 (2-81); 325 (2-124); 345 (2-81); 

347 (7-50b); 353 (7-1); 390 (2-36); 391 (2-69a) 

Ribeiro, B. G. S., 142 (6-13); 151 (6-la); 183 (2-52); 

273 (7-56b); 277, 299 (6-21); 307 (7-56a); 447 (6- 

3a); 484 (6-37); 752 (2-119); 844, 1013 (7-8); 1090 

(7-50a); 1105, 1107 (2-52); 1307 (3-1); 1511 (7-18); 

1548 (7-24); 1584 (7-56a); 1608 (7-24); s.n. (7-30) 

Richardson, W. D., 744 (7-25) 

Rico Gray, V., 124, 444 (1-1) 

Riedel, L., s.n. (3-16); 405 (2-25); 1111 (2-30); 1521 

(2-27b); 1869 (7-85); 2319 (3-16); 2841 (6-27) 

Riedel, W. D., 1371 (6-16); 1667 (3-4) 

Rimachi Y., M., 1803 (2-52); 1807 (7-40); 1854 (2- 

69b); 1861 (2-60); 1881 (2-140); 1884 (7-40); 2264 

(6-35); 2302 (7-40); 2324 (7-53); 2326 (2-64); 2424 

(6-3b); 2740 (7-53); 2789 (6-35); 2797 (2-43c); 2809 

(6-44); 2838, 2866 (2-23); 3136 (2-69b); 3165 (2- 

52); 3250 (6-36); 3260 (6-26); 3274 (2-7); 3277 (2- 

69b); 3357 (6-44); 3374 (7-56b); 3391 (7-5); 3465 

(6-lb); 3472 (2-26); 3493 (3-9); 3643 (7-56a); 3681 

(2-69a); 3685 (3-4); 4208 (7-24); 4257 (2 sp.); 4267 

(6-3b); 4378 (3-14); 4622 (2-26); 4681 (7-40) 

Rizzini, G. M., 158 (7-36a) 

Rizzo, J. A. et al., 2990 (2-114); 4312 (7-54) 

Roa, A., 365 (6-41.1); 435 (2-82) 

Robbins, S. B., 5793 (1-1) 

Robert, A., 567b (6-27) 

Roberts, L., LBB14763 (2-117); LBB16303 (7-11.2) 

Robertson, K. R. et al., 243 (2-77) 

Robleto, W., 8 (1-1); 194 (2-15); 372, 408 (2-45); 641 

(7-22); 739 (6-47) 

Rodrigues, I. A., 234 (7-54) 

Rodrigues, R. S., MG8813 (2-86) 

Rodrigues, W. A. et al., 887f(2-124); 1408 (8-4); 5402 

(6-la); 7263 (2-136); 8257 (7-55.1); 8353 (2-27a); 

8770 (2-22); 8815 (2-27a); 8832 (4-1); 9064 (6-28); 

9097 (2-53); 9278 (2-88); 9284 (2-119.1); 9317 (7- 

8); 9405 (7-56a); 9408 (7-54); 9411 (7-50a); 9621 

(2-69a); 10064 (2-43); 10129 (2-131); 10224 (2-57); 

10230 (7-56b); 10259 (2-121.1); 10497 (2-57); 10501 

(2-135.1); 10505 (6-15); 10509, 10528 (6-29.1); 

10542 (2-37); 10556 (2-69a); 10559 (6-7); 10609, 

10616 (2-57); 10630 (6-15); 10635, 10642 (2-57) 

Rodriguez A., A. A. et al., 57 (7-56b) 

Rodriguez, B., 1419 (7-15) 

Rogers, G., 27 (7-5) 

Rojac, C., 4117 (3-16) 

Rojas, M., 50 (2-15) 

Romero Castafieda, R., 1212 (7-21); 5270 (2-148); 5570 

(3-17); 8445 (7-56a) 

Rombouts, H. E., 321 (7-20) 

Rooden, J. van et al., 358 (2-124); 536 (2-32); 553 (2- 

124); 586 (3-15) 

Rosa, M., 74 (7-36b) 

Rosa, N. A. et al., 33 (2-134); 76 (7-56a); 170 (2-28); 

242 (2-37); 373 (2-140); 378 (2-24); 381 (2-27a); 384 

(3-6); 436 (6-21); 513(2-94); 540 (6-41); 672 (2-141); 

732 (2-69a); 847 (7-56a); 882 (7-50a); 888 (2-37); 

914 (7-20); 916 (2-43b); 961, 988 (7-18); 1006 (2- 

63); 1052 (7-11); 1061 (2-117); 1133 (7-50b); 1134 

(2-109); 1163 (7-20); 1165 (2-67); 1353 (7-69); 1423 

(6-43); 1452 (7-56b); 1467 (2-27b); 1498 (7-25); 1564 

(2-119.1); 1622(2-28); 1689 (2-36); 1697 (4-3); 1707 

(7-8); 1713(6-29.1); 1795 (2-112); 1799 (2-109); 1926 

(3-9); 1931 (3-4); 1967 (2-75); 1988 (2-28); 2025 (7- 

54); 2046 (2-43b); 2098 (7-73); 2210 (3-4); 2308 (2- 

114); 2310 (2-69a); 2331 (7-56a); 2337 (2-126.1); 

2405 (6-50); 2568 (2-56b); 2708 (2-86); 2724 (7- 

56a); 2727 (2-79a); 2753 (2-99); 2848 (2-86); 2891 

(2-37); 2923 (7-24); 2924 (2-99); 2930 (6-12); 2942 

(2-27a); 2954 (2-134); 3097 (2-27b); 3120 (2-122); 

3124 (2-18); 3148 (3-3); 3155 (6-50); 3157 (2-81); 

3310, 3586 (7-36a); 3651 (7-53); 3655 (7-14); 3607 

(7-28) 

Rosario, C. da S., 9 (7-73) 

Rosario T., A. J., 82 (2-45) 

Rosbach, G. B., 3719 (7-61) 

Ruiz, A. G. et al., 224 (2-128) 

Ruiz C., J., 30 (2-43c); 500 (2-22) 

Ruiz, D., 182 (7-22) 

Ruiz, T. et al., 3949 (2-69b) 

Rutkis, E., 55 (2-26); 285 (6-3b); 712 (2-27b) 

Rylands, A., 42/80 (2-124); 53 (2-99); 56/80 (2-124) 

Rzedowski, J., 22516 (2-15) 

Sabatier, D., 50 (2-63); 75 (2-69a); 71 (2-65); 110 (2- 

99); 119 (6-7); 130 (2-35); 205 (6-la) 

Safford, W. E., s.n. (2-1) 

Sagastegui A., A., 550, 6866 (7-56b) 

Salazar, A., 2, 662 (2-64) 

Saldana, 655 (3-4) 

Sanchez, E., 8 (1-1) 

Sanchez V., P. et al., 424 (7-36a) 

Sandino, J. C. et al., 797 (1-1); 1822 (2-45); 2239 (1- 

1); 2678 (7-22); 3012 (1-1); 3630 (2-15); 3949, 4046 

(1-1); 4312 (6-47); 4395, 4407 (2-15); 4447 (2-45); 

4746 (7-22); 4772 (7-36c); 4963 (2-45) 

Santino, 284 (7-54) 

Santos, A., 84 (7-36a) 

Santos, F. S., 193 (6-49); 409 (2-lOOa) 

Santos, J. L. dos et al., 681 (2-119.1); 717 (2-131); 758 

(2-165) 

Santos, J. U., 255 (7-73) 

Santos, M. R., 11 (2-69b); 57 (2-22); 120 (2-69b); 162 

(2-129); 166 (3-4); 281 (2-69a); 286 (7-24); 309 (7- 

56a); 377 (2-39); 396 (2-27a); 444 (2-136); 466 (7- 

50b); 472 (2-69a); 485 (2-27a); 507 (2-99); 532 (7- 

63); 569 (7-56a); 643 (2-69a); 652 (3-4); 686 (2-39) 

Santos, R. R., 49 (4-3) 

Santos, T. S. dos et al., 164 (3-18); 300 (7-22.1); 309 

(6-26); 311 (3-18); 320 (2-146); 322 (6-53); 326 (2- 

114); 457 (3-3.1); 526 (7-36a); 570 (7-64); 945 (2- 

43a); 1279 (6-51b); 1284 (7-37); 1402 (7-20); 1441 

(7-36a); 1444 (2-146); 1512 (7-70.1); 1677, 1695 (7- 

13); 1837 (2-43a); 1995 (7-36a); 2287 (7-20); 2305 

(3-3.1); 2349, 2468 (2-43a); 2693 (7-36a); 2898 (7- 

13); 2935 (6-52.2); 2984 (7-20) 

Sarmiento, A. C., 604 (4-2); 614/80, 651 (7-18) 

Sastre, C. et al., 514 (6-44); 641 (3-14); 671 (7-56a); 

839, 854 (7-40); 872 (7-56b); 1027 (7-40); 1311 (2-


69a); 1363 (7-56a); 1397, 1415 (7-73); 1617, 1816 

(7-56a); 2357 (7-62); 2427 (2-69b); 2436 (7-62); 2437 

(2-68); 3127 (7-5); 3268 (7-56b); 3273 (7-35); 3350 

(2-69b); 3358 (7-55); 3370 (7-56b); 3443 (3-1); 3486 

(2-68); 4107 (7-73); 4156(7-18); 4348 (2-134); 4426, 

4536 (2-112); 5060 (2-104); 5070 (7-5); 5106 (7-2); 

5178 (2-104); 5225 (7-40); 5587, 5674 (7-73); 5688 

(7-56a); 5990 (7-18); 5991 (7-56a); 6110 (3-1) 

Saunders, J., 153, 223 (1-1); J 233 (7-22); 417 (1-1); 

530, 574 (7-56b); 743 (1-1); 1197 (7-23); 1304 (7- 

56b) 

Sauvain, M., 156 (2-88) 

Schatz, G. E. et al., 779 (7-56b) 

Scheiner, P., 50 (2-114); 58 (7-54) 

Schmidt, E., 26 (3-5) 

Schnell, R., 11129 (7-25); 11134 (1-1); 11425 (7-56); 

11728 bis (7-56a); 11751 (7-2); 11849 (7-56); 11899 

(7-73); 12040 (3-1); 12069 (6-lb); 12089 (6-1c); 

12114(3-1) 

Schomburgk, R. H., 318 (2-134) 

Schubert, B. G., 42 (1-1) 

Schultes, R. E. et al., 6520 (6-3b); 12979 (3-6); 13416 

(2-69b); 14913(2-39); 15456, 15660(2-27.1); 16856 

(2-69b); 19161 (7-21); 19398 (7-56b); 19421 (7-5); 

19602 (2-69b); 26038 (7-56b); 26106 (6-9); 26139a 

(6-13); 26184a (7-28) 

Schunke V., J., 10 (7-75); 16 (2-68); 1737 (2-45); 2008 

(7-36a); 2050 (2-43b); 2063 (7-56a); 2137 (7-75); 

2187 (2-69a); 2194 (7-56a); 2587 (2-43b); 3344 (6- 

36); 3525 (7-56a); 3794 (7-28); 4103 (2-141); 4128 

(7-56a); 4389, 4610 (7-36a); 4767 (6-36); 4900 (2- 

129); 5622, 6210 (7-21); 6247 (6-16); 6314 (2-129); 

6447 (7-73); 6502 (2-7); 6518 (7-28); 6637 (7-56a); 

6671, 7222 (7-21); 8439 (2-144.1); 11926 (6-34) 

Schwacke, C. A. W., 256, 4256 (3-4) 

SEF (Studies ofEcuadorean Forests), 9269 (2-60); 10321 

(6-36) 

Seymour, F. C., 3787 (7-56b); 4593, 5957 (1-1) 

Shepherd, G. J., 7443 (7-19) 

Shepherd, J. D., 197 (7-34) 

Shepherd, W. O., 62, 145 (2-1) 

Sidney, (Fonseca) et al., 199 (7-54); 1302, 424 (2-30) 

Silva, A. F., s.n. INPA68839 (6-29) 

Silva, A. S. L. da et al., 8 (2-99); 65 (2-141); 132 (2- 

129); 134 (2-81); 144 (2-56b); 220 (6-4); 426 (3-2); 

452 (6-57); 511 (6-14); 534 (6-13); 540 (7-5); 569 

(6-9); 1980 (2-134) 

Silva, F. C. F. da, 35 (7-54); 82 (2-144); 237 (2-44) 

Silva, I. A., 8 (2-69a); 30 (7-37); 94 (2-114); 131 (7- 

85); 136 (7-64); 267, 268 (3-4); 283 (6-17) 

Silva, J. A., 105 (7-56b); 178 (2-124); 193 (2-99); 202 

(2-27b); 242 (2-81); 278 (3-4); 290 (2-17); 301 (6- 

3b); 338 (2-27a); 356 (2-129) 

Silva, J. C. da, 40 (7-19); 100 (7-52.1) 

Silva, M. F. F. etal., 1102 (2-43a); 1382 (2-134); 1404, 

1446 (7-56a) 

Silva, Manoel, s.n. INPA27688 (2-112) 

Silva, Marlene F. da et al., 45 (2-57); 46 (7-56a); 50 

(2-62); 80 (2-69.1); 127 (4-1); 135 (6-3b); 146 (4-1); 

177 (7-73); 193 (7-2); 223 (6-15); 224 (2-124); 228, 

243 (2-69a); 248 (6-29); 262 (2-83); 267 (6-35.2); 

337 (2-43b); 348 (2-17); 360 (2-142); 391 (2-69b); 

465 (6-15); 467 (6-35.2); 486 (2-64); 490 (6-69.1); 

516 (2-69a); 527 (7-1); 535 (7-56b); 562 (2-75); 569 

(7-6); 705 (2-99); 732 (7-1); 782 (7-20); 788 (7-1); 

791 (7-6); 822 (6-35.2); 824 (2-83); 828 (2-142); 873 

(2-73.1); 874 (2-21); 901 (2-83); 914, 922 (3-2); 930 

(2-99); 942 (3-2); 967 (6-34) 981 (2-88); 993 (6-29); 

1027 (6-35.2); 1151 (7-56a); 1186 (2-28); 1207 (2- 

69b); 1228 (3-6); 1339 (2-27a); 1365 (2-140); 1466 

(3-6); 1474 (7-62); 1510 (7-21); 1532 (2-140); 1538 

(2-27a); 1618 (7-55); 1620 (7-56a); 1630(2-28); 1671 

(7-55); 1701 (2-69a); 1815, 1866(2-69b); 1918, 1932 

(3-4); 2043 (2-27b); 2230 (6-15) 

Silva, Milton G. da et al., 716 (2-86); 892 (6-3a); 936, 

945 (2-27a); 1147 (2-60); 1181 (6-37); 1220 (2-81); 

1304 (6-3a); 1408 (6-37); 1914 (2-69a); 2190 (6-21); 

2345, 2471,2686 (7-56a); 2716 (7-24); 2718 (7-56a); 

2759 (7-18); 2818 (1-1); 2837 (7-56b); 3122 (3-4); 

3128 (7-15); 3217 (2-17); 3230 (2-37); 3269 (2-30); 

3327 (2-37); 3408 (2-63); 3496 (7-56b); 3508 (2- 

121.1); 3527 (2-37); 3554 (2-69a); 3570, 3599 (7- 

53); 3620 (6-3a); 3630 (2-129); 3705 (7-56a); 3754 

(7-24); 3792 (2-27a); 3793 (6-21); 3805 (2-22); 3825 

(6-3a); 3873 (6-21); 3962 (2-52); 4281 (2-27); 4296 

(2-64.1); 4586 (2-41); 4662 (2-57); 4684 (2-64.1); 

4744 (7-56a); 4746 (3-4); 4753 (7-20); 4826, 4845 

(7-75); 4892 (7-20); 4906 (2-41); 4945 (7-54); 5008 

(2-41); 5020 (6-27); 5026 (2-41); 5207 (7-30); 5237 

(6-1c); 5279 (7-30); 5324 (2-81); 5348 (6-lc); 5402 

(2-43b); 5442 (2-81); 5455 (7-56a); 5489 (2-69a); 

5498 (6-50); 5500 (2-57); 5513 (7-56a); 5522 (2-81); 

5524 (7-69); 5528 (7-42); 5534, 5535 (6-50); 5537 

(6-la); 6574 (2-92); 6589 (2-53); 7121 (6-29.1); 7122 

(2-119.1) 

Silva, M. N., 271 (7-20); 390 (3-4); 403 (7-56b) 

Silva, N. T. da, 593 (6-lc); 838 (2-8); 1045 (6-50); 1756 

(3-4); 1796 (4-3); 1825 (2-114); 1831 (7-24); 1841 

(6-la); 1857 (3-4); 1881 (6-50); 1901 (7-36a); 1969 

(7-50b); 2093 (2-39); 2096 (2-53); 2170 (7-27); 2203 

(2-53); 2215 (7-42); 2294 (6-la); 2329 (2-83); 2380 

(3-4); 2385 (7-15); 2386 (7-42); 2408 (6-la); 2420 

(7-56a); 2512 (7-42); 2517 (6-1a); 2547 (7-56a); 2621 

(6-7); 2683 (6-la); 2768, 2782 (2-69a); 2826 (2-136); 

2833 (6-la); 2879, 2911, 2925 (2-124); 2950 (7-24); 

3170 (6-lc); 3203 (3-4); 3223 (6-la); 3241 (3-4); 

3292 (7-50b); 3296 (6-la); 3310 (3-2); 3311 (3-4); 

3316 (6-la); 3342 (6-50); 3391 (7-36a); 3424 (6-la); 

3556 (2-17); 3615 (2-141); 3631 (2-17); 3727 (2-75); 

3742 (2-129); 3874 (2-140); 3894 (7-21); 3897 (7- 

8); 3926 (2-43a); 3950 (6-la); 3978 (2-99); 4028 (6- 

41.1); 4174 (7-62); 4456 (7-56b); 4459 (2-17); 4490 

(2-27a); 4492 (2-129); 4521 (6-lb); 4578 (2-119.1); 

4609 (2-43b); 4612 (2-36); 4753 (2-69a); 4755 (2- 

142); 4764 (6-21); 4796, 4805 (6-3a); 4810 (2-30); 

4817 (7-56b); 4821 (2-27a); 4827 (6-20); 4837 (7- 

54); 5064 (2-37); 5086 (7-24); 57170 (3-16); 60697 

(2-27.1) 

Silva, S. B. da et al., 383 (7-18) 

Silva Costa, J. da, 1221 (2-28) 

Simpson, D. R. et al., 51 (2-43b); 76 (7-40); 732 (7- 

53); 761 (7-40) 

Skog, L. et al., 5643 (7-56b) 

Slane, V., 12 (1-1) 

Smith, C. E., Jr., 6034 (6-3b); 6054 (2-12)


Smith, D. et al., 1173 (2-98); 8409 (7-56b) 

Smith, E., 8 (2-21) 

Smith, F. D., Jr., s.n. (2-12) 

Smith, R. F., V1592 (2-76); 4301, 4302 (2-12) 

Smith, S. F. et al., 121, 190, 234, 297, 340, 381 (7- 

56a); 590 (7-53); 7856 (2-69b) 

Snow, D. W., 8 (7-25); 21 (7-26); 37 (7-25) 

Soares, S. et al., s.n. on 18 II 1985 (2-122) 

Sobrinho, J. F. et al., 322 (7-18); 890 (3-4) 

Soejarto, D. D. et al., 360 (1-1); 449 (2-12); 694 (2- 

69a); 2396 (6-la); 4025 (7-34) 

Solomon, J. C., 6111 (7-56a); 6205 (7-66); 6509, 7605 

(7-36a); 7659 (7-56b); 7788 (7-53); 7856 (2-69b); 

7944 (7-66); 8502 (7-46); 8784 (7-36a); 9393 (7-46) 

Sonkin, L., 340 (3-4) 

Soria S., M. A., 20 (2-22) 

Soto Nuiiez, J. C. et al., 32 (2-45) 

Souza, A. B. de, 78 (6-25); 83 (4-2) 

Souza, D. S., 242 (1-1) 

Souza, J. L., 69 (7-56a) 

Souza, H. M. de, IAC21452 (7-54) 

Spada, J., 007/77 (6-17); 148 (6-26); 151 (4-5); 193 (2- 

43a); 210 (7-13); 329 (2-114); 31/78 (6-13.2); 67/78 

(2-43a); 79/78 (2-71) 

Sperling, C. et al., 5792 (7-56a); 5794 (7-36a); 5940 

(2-43a); 5942 (3-4); 5945, 5977 (2-99); 6029 (7-36a); 

6028 (7-53); 6048 (7-36a); 6072 (7-73); 6102 (7-15); 

6124 (2-134); 6176 (7-73); 6178 (7-15); 6251 (2-17); 

6327 (7-65); 6334 (2-43b); 6370 (7-36a); 6437,6443, 

6640 (7-56a) 

Spetzman, L. A., 539 (1-1) 

Spichiger et al., 1004 (7-41); 1170 (7-40) 

Spongberg, S. A. et al., 17197 (2-1) 

Stahel, G., 322 (2-88); 353 (2-18) 

Stannard, B. et al., CFCR5960 (2-144) 

Stein, B A. et al., 1329 (7-56b); 1369 (7-36a); 1471 (7- 

55); 1486 (7-62) 

Stergios, B. et al., 2683 (3-1); 2787 (7-56a); 3179 (2- 

69b); 3464 (1-1); 4112 (2-88); 4208 (7-5); 4420 (7- 

21); 4685, 4881 (7-36a); 4991 (6-45); 5535 (2-27b); 

5718 (7-22); 5793 (7-36a); 6175 (6-45); 7944 (3 sp.); 

8455, 8460 (7-40); 8470 (2-27); 8619, 8623 (7-56b) 

Stevens, W. D. et al., 5453 (2-45); 7204, 7478 (7-22); 

7555, 7634 (7-23); 7642 (7-56b); 7714 (1-1); 7802 

(7-22); 7818 (1-1); 7853 (7-56b); 7866 (2-88); 7902 

(1-1); 8811 (2-22); 8152 (7-56b); 8283 (7-36c); 8469 

(7-56b); 9669 (2-45); 10473 (1-1); 10475 (7-56b); 

10478, 10686 (1-1); 12104, 12769 (7-56b); 17147 

(2-45); 17560 (2-15); 17746 (1-1); 18644 (7-56b); 

19456, 19563, 19584, 19606 (1-1); 19826 (2-88); 

19828 (7-56b); 19993 (1-1); 20058 (2-88); 20075 (7- 

56b); 20635 (2-88); 20636 (7-56b); 20875 (1-1); 

21460 (7-22); 23357 (2-15) 

Stevenson, N. S., YALE10697 (7-23) 

Steward, W. C. et al., 36 (2-69a); 38 (2-36); 54 (6-19); 

61 (7-73); 88 (2-119.1); 110(2-33); 130(2-134); 134 

(6-3b); 174 (7-21); 289 (2-27); 341 (2-27a); 390 (2- 

69b); 511 (7-56a); P17669 (2-69.2); P17680 (3-4); 

P17697 (7-8); P20128 (6-29); P20241 (2-69a); 

P20251 (2-119.1); P20304 (6-4); P20310 (2-27); 

P20326 (2-119.1); P20328 (2-88); P20369 (2-36); 

P20401 (2-119.1) 

Steyermark, J. A. et al., 52829 (2-4.2); 75127, 75526 

(6-8.1); 86613 (3-3); 87137 (7-74); 87610, 88108 (6- 

6); 93035 (6-8.1); 101826 (3-7); 102626 (3-1); 102951 

(2-27a); 102953 (2-119); 102974 (2-27a); 103013 (7- 

21); 103225 (6-3b); 103247 (2-131); 103260(2-119); 

104154 (2-56); 104223 (7-26); 104352 (2-124); 

104492 (7-26); 104547 (2-69a); 104777 (7-22); 

105503 (2-56); 105936 (6-8.1); 106087 (2-69a); 

106117 (6-8.1); 106359 (7-49); 106411 (2-26); 

106412(2-28); 106645 (7-49); 106847 (7-22); 107132 

(7-50b); 107357 (2-60.1); 107456 (2-68); 108611 (7- 

56b); 108939,109173(2-135); 109900(2-12); 111391 

(2-105); 111507 (2-69a); 111512 (2-145); 111541 (2- 

5.4); 111609(7-22); 112386 (7-21); 113206, 113240 

(2-89); 113875 (6-3b); 113900 (2-87.1); 114424 (3- 

1); 114811 (2-39); 115119 (7-73); 115131 (2-8); 

115141(2-39); 115545(3-7); 116339 (7-36a); 116506 

(3-7); 116808 (2-86); 116841 (2-114); 116868, 

116931, 116933 (7-34); 117028 (2-39); 117588 (2- 

56); 117617 (2-68); 117651 (7-50b); 117696 (1-3); 

117776 (2-105); 117792, 117824 (3-10); 117839 (7- 

25); 117921 (7-9.1); 118135 (2 sp.); 119155 (6-3b); 

119388 (2-43a); 119397 (2-57); 119454, 119576 (6la); 

119749 (7-36a); 120004 (7-22); 120609 (2-69a); 

120732 (7-74); 121338 (7-36a); 121636 (2-69a); 

121703 (7-74); 121791 (7-36a); 121800 (7-22); 

121858 (7-36a); 122046 (2 sp.); 122072 (2-106); 

122405 (6 sp.); 122715, 122853, 122913, 123127, 

123184, 123300 (7-36a); 123392 (3-7); 123659 (7- 

68); 124598, 124638 (7-36a); 124748 (6 sp.); 124924 

(7-36a); 125670 (2-88); 125859, 126219 (2-68); 

129177 (3-4); 129367 (2-124); 130185 (2-57.1); 

130873 (2-124); 130906A (2-99); 131173 (2-27b); 

131203 (3-4); 131406 (6-3b); 131527 (7-25); 131653 

(3-4); 131676 (7-56b); 131888 (2-68); 131889 (7- 

56a); 131957 (2-81); 132163 (2-135) 

Stoffers, A. L. et al., 114 (2-108); 140 (2-90); 178 (7- 

56b); 222 (7-36a); 254 (7-73); 300 (2-108a); 404, 

492, 513 (7-25); 517 (6-43); 3659 (7-56a); 30143 (2- 

108.1) 

Strang, H. E., 208 (6-25) 

Strudwick, J. J. et al., 3042 (7-56b); 3347 (2-134); 3587, 

3589, 3614, 3689 (7-56b); 3766 (2-134); 3789 (7- 

20); 4047 (7-56b); 4245 (7-50a); 4352 (7-20); 4395, 

4450 (7-56b) 

Sucre, D., 1053 (6-25); 3525 (7-36b); 3949 (1-1); 4275 

(7-85); 5730 (7-36b); 6384 (7-82); 7939 (6-25); 9350, 

9384 (7-56b); 9579 (6-25); 10248 (1-1); 10299 (2- 

44); 11352 (7-36b) 

Svensson, B. et al., 685 (1-1) 

Sytsma, K. J. et al., 1551 (7-38); 3183, 3258, 3325, 

3335, 3557, (7-56b) 

Takeuchi, s.n. INPA7809 (2-22) 

Tamashiro, J. Y., 6554 (2-30); 8765 (7-82) 

Tamayo, F., 3541 (2-27a) 

Tate, G., 870 (6-8.1) 

Tavares, A. S., 114 (2-126.1) 

Tawjoeran, J. A., LBB14443 (4-3) 

Taylor, E. L., 1298 (7-18); 1301 (7-56a) 

Teixeira, L. O. A., 43 (7-56b); 122 (2-36); 157 (6-34); 

253 (2-53); 563 (7-56a); 728 (2-95.1); 798 (7-55a); 

804 (7-56b); 927 (6-la); 975 (7-6); 1201 (7-50b); 

1203 (6-9); 1222 (2 sp.); 1238 (2-69a); 1246 (3-4); 

1365 (7-66); 1511 (7-56a); 1581 (2-60)


Tellez, O., 2002, 2098 (7-22) 

Terceros, W. et al., 21 (7-36a) 

Terezo, E. F., s.n. INPA139843 (7-25) 

Thomas, W. W., 3195 (7-30); 3575 (2-32); 3598, 3672 

(7-56b); 3747 (6-47); 3808 (2-17); 3810 (2-124); 3836 

(2-37); 3876 (2-144); 4091 (2-56a); 4095 (2-27a); 

4154 (7-20); 4309 (2-30); 4324 (2-37); 4329 (7-20); 

4404 (2-114); 4457 (7-20); 4506, 4523, 4525 (2-27a); 

4572 (2-28); 4656 (7-66) 

Thore, R. F. et al., 57938, 57955 (2-1) 

Tidestrom, I., 4182 (1-1) 

Tjon Lim San, R., LBB14813 (6-1) 

Todzia, C. et al., 2212 (6-9); 2226 (7-9); 2283 (7-6); 

2295 (6-35.2); 2328 (2-129) 

Torres, A.M., 1804 (1-1) 

Torres C., R., 639 (6-47) 

Torres, J., 85 (6-16); 93 (6-35); 290 (6-3b); 830 (2- 

140); 919 (6-36) 

Trigos, R. C. (see Cedillo T., R.) 

Trinidade, L., 30070 (2-127) 

Troth, R. G., 1117 (2-12) 

Trujillo, B. et al., 3663 (7-74); 3765 (7-56a); 3789 (2- 

12); 4497 (7-56a); 4534 (7-25); 4655, 5323 (2-12); 

5518 (7-20); 5774 (6-3b); 5816, 5847 (7-56b); 5940 

(7-25); 6058 (2-26); 14980 (7-40); 15043, 15168, 

15243 (7-25); 15262 (7-56b); 15316a (6-3b); 15338 

(2-114); 15361a (2-88a); 15381 (7-56b); 15458 (3- 

4); 15473 (2-69a); 16224 (2-39); 17354 (7-25) 

Tunqui, S., 105 (3-9) 

Tyson, E. L., 7367 (7-56b) 

Ucan ek, E et al., 627, 1084 (1-1) 

Uhl, C. F., 458 (7-56a) 

Ule, E., 4214 (6-50) 

Univ. Brasilia, Taxonomy Class, 189 (2-114); 519 (3- 

16) 

Uribe U., L., 127 (7-25) 

Utley, J. et al., 5488 (2-18.2) 

Utrera, A., 81 (7-49) 

Valle, M. A., 46 (2-69b); 149 (7-56b) 

Van der Werff, H. et al., 6998 (2-88); 9540 (2-13.1) 

Van Hall, C. J. B., 47 (2-66); 49a (6-1); 49b (2-135); 

51 (2-27a) 

Varela, J. R. C., s.n. (1-1) 

Vasconcelos, H. L., C2-7, H13 (2-69a); H16 (2-81); 

H19 (6-28); J7 (2-76); J38 (7-50b); J53 (2-76); J59 

(2-142); S20 (2-69a) 

Vasquez, R. et al., 110 (6-44); 182 (2-27); 184 (6-3b); 

1008 (6-6.1); 1102 (6-44); 1220 (7-56b); 1547 (2-99); 

1255 (6-44); 1309 (6-6a); 1392 (2-99); 2085 (6-6a); 

3387 (7-56a); 3463 (6-7); 3444 (2-43c); 3472 (7-5); 

3479 (7-40); 3488 (2-69a); 3572 (7-56b); 3599, 3640 

(2-27b); 3696 (6-44); 3759 (2-52); 3762 (6-29); 3860 

(2-36a); 3976 (2-26); 3995 (2-69a); 3997 (6-34); 4000, 

4011, 4023, 4043 (2-69a); 4263 (6-3a); 4312 (2-21); 

4460 (2-39); 4492 (7-5); 4681 (6-36); 4788 (2-43b); 

4973 (7-36a); 5204 (6-36); 5255 (6-3b); 5263 (2-52); 

5267 (2-60); 5271, 5497 (7-53); 5637 (7-24); 5650 

(7-56b); 5655, 5656 (6-35); 5698 (2-62); 5782 (6- 

35); 5812 (2-39); 6124 (2-128); 6127 (7-5); 6135 (2- 

88a); 6147 (2-69a); 6152 (2-94); 6166 (7-53); 6197 

(6-62); 6220 (2-88a); 6243 (2-43c); 6411 (3-4); 6435 

(2-23); 6467 (6-44); 6622 (2-22) 

Vaz, A. M. S. F., 158 (7-18) 

Veillon, J. P., 2/v (2-27); 57 (7-36a); 131 (2-45) 

Velasquez, N., 28 (2-45) 

Vellow, 299 (3-4) 

Ventura A., F., 976 (7-36); 19786 (7-36c); 20011 (7- 

36a); 20203 (7-36c) 

Viera, M. G. et al., 8, 13 (7-56a); 72 (2-124); 123 (6- 

3a); 134 (7-50a); 151 (7-20); 281 (7-56a); 633 (2- 

41); 750 (2-57); 877 (2-114); 929 (2-82); 949 (6-27); 

977 (2-17); 995 (4-4) 

Vilhena, R., 46 (7-56a); 143 (7-50a); 144 (6-3a); 147 

(4-1); 211, 266 (7-50a); 289 (4-1); 304 (2-27b); 305 

(6-3a) 

Vincelli, P. C., 532 (2-88); 586 (1-1) 966 (7-25); 1012 

(2-124); 1055 (2-69a); 1066 (2-39) 

Vinha, S. G. da, 12 (3-11) 

Vital, D. M. et al., s.n. (1981) (7-54) 

Voeks, R., 28 (2-130.1); 53 (6-51b); 72 (7-37) 

Vogl, C., s.n. (3-7) 

Wachenheim, 115 (2-69a) 

Wagner, R. J., 1747 (7-83) 

Warer, R. H., 299 (2-148) 

Webster, G. L. et al., 9806 (2-69a) 

Wendt, T. et al., 3302 (7-36c); 3325 (2-15); 3724 (7- 

36c); 3899 (7-56b) 

Wessels Boer, J. G., 2073 (2-109); 2074 (7-56a); 2317 

(6-13); 2334 (7-21); 2405 (7-62) 

West, E., s.n. (2-44) 

Wherren, L., 16 (7-73); 105 (2-98) 

Whetstone, R. D., 13350, 14376 (2-1); 14501 (1-1); 

14514 (2-1) 

White, S. et al., 460 (7-22) 

Whitefoord, C., 2761 (2-88a); 2828 (7-56b); 3084 (1- 

1); 3151 (7-56b); 3290 (7-36a) 

Widgren, J. F., s.n. (3-4) 

Wilbur, R. L. et al., 10847 (2-14) 

Williams, LI., 3173 (6-16); 12510 (2-27); 14914 (4-1); 

16049 (2-69b) 

Williams, L. O. et al., 15962 (6-3b); 26513 (2-45) 

Williamson, C. S., s.n. (2-1) 

Witherspoon, J. T. & F., 8543 (7-80) 

Witsberger, D., 813 (2-9) 

Wolfe, F., 12181 (7-36a) 

Wood, C. W., 292, 410 (2-145) 

Woodbury, R. O., s.n., 30464 (7-83) 

Worthington, R. D., 13561 (1-1) 

Woytkowski, F., 5773 (2-64); 6318 (6-44) 

Young, K. et al., 146 (7-56a); 1037 (7-75); 1047 (7- 

56a) 

Zaandam, C., 6764 (6-la) 

Zabala, A., 80, 102, 162 (7-16) 

Zanoni, T. et al., 12668, 15904, 17737, 21206, 25782 

(1-1) 

Zappi, D. C. et al., CFCR8476 (2-43a) 

Zarucchi, J. L. et al., 1221 (6-9); 1278 (2-64); 1944 (3- 

6); 1983 (7-25); 2038 (7-25); 2548 (7-8); 2574 (2- 

57); 2580 (2-124); 2585 (7-20); 2858 (7-1); 2862 (7- 

9); 2920 (7-56d); 3079, 3100 (35.2); 3162 (2-129); 

3174, 3186 (2-140); 3396A (2-69b); 3401 (2-27b); 

3452 (2-83); 3491 (2-69b); 3533 (2-39); 3529 (7-40); 

3583 (3-4); 3626 (6-36); 3630 (2-24); 3634 (7-56b); 

3666 (6-3b); 3667 (2-69b); 3689 (2-140); 3698 (2- 

69a); 3773 (3-4)


SECOND SUPPLEMENTAL LIST OF EXSICCATAE 

Acevedo, P. et al., 1607 (7-2); 1657 (2-13.1) 

Acosta P., R. et al., 1653 (7-56b) 

Alvarez, H. J. et al., 34 (1-1) 

Amaral, I. L. et al., IG2-6-173, TF-2-15 (2-57) 

Aumeeruddy, 73 (2-114); 92 (2-109) 

Axelrod, F., 610 (7-83) 

Ayala, F. et al., 2888 (6-16); 3023 (2-43b); 3572 (7- 

56b) 

Aymard C., G. et al., 4825 (2-68); 4872 (7-49); 5355, 

5425 (7-25); 5552 (7-36a) 

Bal6e, W. et al., 2658 (7-24); 2737, 2829, 2836, 2836, 

2882 (6-la); 2894 (2-69a); 2902, 2925, 2928, 2929 

(6-la); 2943 (2-67); 2982, 3004 (6-la); 3018 (6-lc); 

3031 (7-56b); 3056 (6-lc) 

Bamps, P., 5478 (6-27) 

Barrier, S., 3863 (7-16); 3989,4259, 4814 (2-77); 4976 

(7-20); 5003 (7-50b); 5022 (2-114); 5183 (7-16); 5191, 

5058 (2-77) 

Beck, S. G., 10013 (2-69b); 10160 (2-69b) 

Billiet, F. et al., 1457 (2-135); 1458 (7-18); 1809 (2- 

109) 

Boom, B. M., 6838, 6881 (7-83); 7119 (2-135); 7127 

(6-7); 7136 (2-78); 7142,7145 (6-42a); 7363 (2-108); 

7529 (7-74) 

Brant, A. E. et al., 1055 (7-56b) 

Calzada, J. I., 5997 (7-56b) 

Castillo, A., 1340, 1518 (7-25); 1653 (2-81); 1859 (2- 

137); 2094, 2173 (7-25); 2208 (6-34); 2305 (2-83); 

2306 (2-121); 2317 (2-83); 2414 (7-56b); 2438 (3-4) 

Cid Ferreira, C. A., 5463 (7-50a); 5490 (2-43b); 5668 

(7-28); 5706, 5762 (2-43b); 5847, 5848 (7-56b); 5981 

(7-1); 6660 (2-62); 6674 (2-124); 6689 (2-129); 6696 

(2-52); 6737 (2-97); 6833 (6-7); 6844 (2-75); 6850 

(2-81); 6894 (2-39); 6897 (3-4); 6903 (2-27); 6911, 

6916 (3-4); 6917 (2-8); 6946 (7-11); 7033 (6-la); 

7064 (6-lb); 7070 (2-27b); 7078 (3-4); 7105 (2-140); 

7143 (2-119); 7145 (7-55); 7160 (2-27b); 7185 (6- 

44); 7209 (3-9); 7248 (6-44); 7254 (2-69b); 7262 (2- 

124); 7291 (3-9); 7326 (6-3b); 7650 (2-134); 7663 

(2-93); 7681 (3-4); 7703 (2-93); 7709 (7-56a); 7719 

(7-39); 7750, 7766 (2-36); 7770 (2-83); 7783 (3-4); 

7795 (6-3b); 7812 (2-36); 7820 (2-88); 7856 (7-50b); 

7857 (6-la); 7863 (7-56b); 7917 (4-3); 7949 (6-la); 

7957 (7-56a); 7968 (2-134); 7978 (6-3a); 8039 (2- 

134); 8041 (2-52); 8054 (6-19); 8073 (2-97); 8143 

(3-4); 8167 (2-75); 8180 (7-52); 8183 (3-4); 8190 (2- 

69a); 8194 (7-9); 8214 (7-1) 

Coradin, L. et al., 5809 (7-18); 6132 (7-56b) 

Cremers, G., 4664 (2-63); 9945 (7-2) 

Croat, T. B. et al., 64503 (7-56b) 

Daly, D. C. et al., 4348 (8-4); 4387 (6-44.1); 4390 (2- 

27b); 4408 (6-9); 4461 (2-69a); 5210 (2-4.2); 5426 

(2-69a); 5447 (2-22); 5565 (7-55); 5568 (4-1); 5635 

(6-34); 5670 (2-46b) 

Davidse, G. et al., 30899, 30941 (1-1); 31993 (7-56b); 

32033, 32393 (7-36a); 32745, 32934, 33165 (1-1) 

Descoings, B. et al., 20068 (6-7) 

Diaz S., C. et al., 2483 (6-3a) 

Dodson, C. et al., 14621 (6-la) 

Dubs, B., 44 (6-27); 237 (2-28); 351 (6-20); 358, 398 

(2-18); 458 (2-28) 

Faber-Langendoen, D. et al., 287 (2-124); 352 (2-96); 

439 (7-56a); 572 (2-148); 660 (2-144.1); 663 (3-17); 

689 (2-46); 693 (3-15); 867 (2-148); 883 (2-124); 979 

(2-148); 1084 (2-51) 

Farney, C. et al., 1218 (6-46); 1451 (6-13.2) 

Fernandez, A., 2942 (7-25); 3311 (2-68); 3339 (7-28) 

Fernmndez Casas et al., 4065 (2-41) 

Feuillet, C., 1089 (2-77); 1374 (2-135); 1388 (3-1); 

1419 (2-124); 1445 (2-69a); 2262 (7-56b); 2303 (2- 

124); 3579 (7-25); 3809 (7-2); 3866 (7-52); 4034 (7- 

25); 4297 (7-2) 

Filho, A. C. et al., 4672 (3-8) 

Fleury, M., 327 (2-124) 

Foresta, H. de, 228 (2-149); 618 (7-52); 733 (2-69a); 

750 (2-134) 

Forget, 279 (7-25); 323, 324, 327 (2-77); 377 (7-74) 

Fournet, A., 211 (7-52) 

Foster, R. B. et al., 11562 (7-21) 

Garcia, R. et al., 335 (1-1) 

Garcia-Barriga H., 20911 (7-8) 

Garwood, N. et al., 231 (7-36a) 

Gentry, A. etal., 25069 (6-35); 53381 (2-32.1); 53676 

(3-15); 53738 (2-5.1); 53780 (2-148); 54188 (2-64); 

54361 (2-5); 54458 (6-22); 54965 (2-43b); 56813 (2- 

144.1); 56842 (7-35); 56884 (2-148); 56892 (2-40); 

56919 (2-46); 56947 (7-35); 56973 (2-124); 57015 

(7-10); 57018 (2-46); 57062 (2-148); 57063 (2-124); 

57516 (7-56a); 57675 (7-75); 57677 (7-56a); 57951 

(7-75) 

G6mez, L. D. et al., 23350 (7-22) 

G6mez, S., 178 (7-22) 

Granville, J. J. de et al., 604 (7-73); B.3787 (2-67); 

5484 (2-56a); 5719, 5722 (2-77); 6121 (4-3); 6183 

(2-77); 6423 (2-67); 6641 (2-27a); 7366 (7-73); 7407 

(7-2); 7467 (2-88); 7651 (7-52); 7786, 8004 (7-74); 

8019 (2-88); 8035 (7-52); 8087 (2-77); 8144 (2-88); 

8201 (2-77); 8678 (7-74); 8820 (7-2); 9072 (6-7); 

9184 (7-52); 9254 (7-56b); 9638 (6-lb); 10243 (7- 

25) 

Grayum, M. H. et al., 5731 (7-36a); 6252 (7-56b) 

Guanchez, F. et al., 4575 (2-83); 4606 (2-43a); 4796 

(2-69a); 4799 (6-3b) 

Haase, R., 634 (2-69b) 

Hammel, B. et al., 7972 (7-61); 11048 (2-88); 14427 

(2-13.1); 14490 (5-1); 14586 (7-36c) 

Hartshorn, G. et al., 2919 (7-50a) 

Henderson, A. et al., 461 (2-115) 

Heringer, E. P., 17167 (3-16) 

Hernmndez, L. et al., 128 (2-107); 175 (2-68); 479 (2- 

107); 550 (7-20) 

Hoff, M., 5002 (1-1) 

Hoist, B. et al., 2640 (7-74); 3046 (7-68); 3052 (7-56b); 

3199 (2-83); 3260 (7-25); 3436 (2-69a) 

Huber, O. et al., 11774 (7-26); 11778 (2-57); 11798 

(3-10); 11973 (6-7); 12062 (2-69b) 

Ibarra M., G. et al., 2284 (7-36c) 

Jacquemin, H., 2016 (7-50b) 

Jansen-Jacobs, M. J. et al., 75 (7-56b); 136 (6-43); 210 

(2-134); 231 (7-56a); 238 (2-135); 347 (7-36a); 495 

(2-27b); 496 (6-15) 

Jaramillo, J., 7457 (2-4.2); 9036 (2-27a); 9066 (7-22)

Second Supplemental List of Exsiccatae 125 

Keel, S. et al., 264 (6-3b) 

587 (6-7); 589 (2-35); 624 (2-69a); 671, 731, 735, 

Kral, R., 72077 (7-49) 

766, 767, 768, 769, 770, 804 (2-77); 830 (1-1); 841 

Kubitzki, K. et al., 79-105 (2-88); 79-43 (7-20); 84- (3-3); 872 (6-la); 890 (2-77); 1090 (2-109) 

293 (6-3b) 

Rivas, R. M., 122 (1-1) 

Kvist, L. P. et al., 187A (7-51) 

Rivero, R., 883 (7-36a) 

Lane, C. & R. Gieschen, 50 (1-1) 

Rodrigues, W. A., IG1-8A-589 (2-57); IG1-10-464 (6- 

Le6n, H. et al., 1347 (7-35); 1561 (7-10) 

15); IL8-22 (2-119.1) 

Liesner, R. et al., 21204 (2-83); 21216 (7-21); 21384 Roosmalen, M. van, 12 (7-73) 

(7-68); 21511 (7-21); 21836 (2-83); 21839 (7-25); Rosa, N. A. et al., 4287 (7-24); 4392 (3-4) 

21873 (2-105) 

Rosales, J. et al., 15, 17 (6-3b); 19 (2-27a); 33 (7-25); 

Lima, H. C. de, 2689 (6-41); 3155 (4-5) 

52 (7-56b); 84, 109 (6-3b) 

Lindeman, J. C. et al., 784 (2-108) 

Rosario, C. S., 96 (7-24); 101 (7-56a); 137 (7-18) 

Maguire, B. et al., 37515 (2-119); 56834 (6-27) Rutkis, E. & K. Udris, 634 (3-7) 

Maciel, U. N. et al., 720 (7-36a); 810 (7-75); 823 (7- Sabatier, D., 88 (2-27.1); 165 (2-86); 484 (2-77); 509, 

20); 829 (7-18) 

572 (7-15); 644, 676 (7-15); 725 (2-77); 830 (7-74); 

Martinelli, G. et al., 11935 (6-46) 

849 (2-134); 866 (2-95); 893 (2-86); 925 (2-77); 939 

McPherson, G., 7449 (2-31.1); 10024 (2-18.2); 10222 (7-16); 1003(2-63); 1022(2-124); 1028 (2-27.1); 1066 

(2-88); 10272 (7-56b); 10278 (2-88); 10588 (6-13.1); (2-97); 1072 (7-73); 1104 (2-99) 

10610 (2-5.3); 10647 (2-31.1); 10804 (2-88) Santos, J. L. dos et al., 681 (2-119.1); 717 (2-131); 732 

Miralha, J. M. S. et al., BO-1-81, BO-1-114, BO-2- (4-1); 758 (2-119.1) 

190 (2-43a) 

Sastre, C. et al., 8016 (7-25); 8129 (7-56b) 

MonsalveB.,M., 1128 (2-5.1); 1278 (2-40); 1297, 1308 Sauvein, M., 218 (2-8); 401 (2-77); 589 (7-56a) 

(2-124); 1503 (2-148) 

Serv. Forestier (Fr. Guiana), 85M (2-124) 

Moraes, M., 513 (7-66); 524 (7-73) 

Sherman, C., 151 (7-22) 

Moretti, C., 872 (6-7); 925 (2-86); 985 (2-69a) Silva, M. G. et al., 3122 (3-4); 4684 (2-64); 5827 (2- 

Mori, S. A. et al., 15152 (2-64); 18484 (7-74); 18534 88); 6216 (7-56a); 6301 (2-30) 

(7-50b); 18522 (6-34) 

Silva, N. T. et al., 4594 (2-128) 

Nee, M., 31393 (7-56a) 

Skog, L. et al., 7476 (7-18) 

Neill, D. et al., 7269 (7-53); 7217 (2-45); 7462 (7-50b); Smith, Damon, 193 (7-61) 

7471 (7-56a); 7667 (2-5) 

Smith, S. F. et al., 1074 (7-66) 

Nelson, B. W. et al., 301 (7-9, 7-56a); 445, 454, 467 Sobel, G. L. et al., 4568 (2-69a); 4569 (2-39); 4643 (2- 

(6-3b) 

141); 4671A (2-69a); 4673A (2-124); 4704 (2-52); 

Nevers, G. de et al., 7519 (2-31.2); 7589 (2-88) 4759 (2-39); 4764 (6-36); 4818 (2-39); 4836 (2-27a); 

Nunez, P. et al., 5370 (7-36a); 5828 (7-66); 6181 (7- 4860 (3-4) 

75) 

Solabarrieta, S., 155 (7-22) 

Oldeman, R. A. A., 1535 (2-134); 1825 (2-63); 2433 Stein, B. A. et al., 3936 (6-44); 3945A (7-5) 

(2-69a); B.2819 (2-27a); B.2278 (2-86); B3119 (2- Stergios, B. et al., 5073 (2-69a); 7319 (7-40); 7419 (2- 

87); B3127 (2-27.1); B3559, T655 (6-lb) 

102); 7472 (2-69b); 7473 (3-6); 7549 (2-140); 7550 

Padilla, F., 191 (7-22) 

(7-56a); 7580 (2-24); 7609 (6-36); 8066 (2-24); 8084 

Palacios, W. et al., 1122 (2-64); 1159 (7-53); 1269 (6- (7-40); 8157 (2-27a); 8175 (2-69b); 9863 (2-68) 

36); 1484 (7-50b) 

Stevens, W. D., 24575 (2-88); 24627 (7-36c); 24628 

Pennington, T. D. et al., 12230 (6-36) 

(2-95) 

Pimentel, J. & R. Garcia, 97-A (1-1) 

Steyermark, J. A. et al., 125689 (2-77); 125863 (2-27a); 

Pipoly, J. J. et al., 8024 (2-77); 8146 (7-56a); 8179 (2- 126228 (2-77) 

69b); 8181, 8241 (2-108); 8337 (2-86); 8428 (6-42a); Taylor, C. M., 7633 (1-1) 

8429 (6-15); 8432 (2-135); 8438 (6-42a); 8444 (2- Thomas, W. W. et al., 4926 (2-43b); 4951 (7-56a); 4969 

73); 8445 (2-135); 8462 (6-42a); 8583 (2-90); 8589 (7-53); 5005 (2-43b); 5050 (6-15); 5070 (7-50b); 5163 

(3-12); 8597 (2-99); 8815 (2-69.1); 8704 (2-90); 8849 (2-17); 5192 (2-99); 5194 (7-56a); 5275, 5258 (2- 

(2-68); 8850 (2-69a); 8853 (2-109); 8929 (2-69a); 99); 5340 (7-9); 5343 (2-69a); 5414 (7-56b); 5439 

8930 (2-68); 8934 (2-112); 8940 (2-124); 8942 (2- (7-42); 5442 (2-99); 5456 (7-56a); 5458 (2-88); 5612 

109); 8945 (2-124); 8955 (2-108); 8959 (6-15); 8962 (6-27) 

(2-108); 9140 (2-135); 9159 (6-15); 9334, 9394 (7- Thorne, R. F. et al., 48111 (1-1); 48566 (2-1) 

25); 9432 (3-1); 9536 (2-93); 9562,9581 (2-78); 9596 Torres, G. A., 117(1-1) 

(6-15); 10741 (2-69a); 10842 (7-21) 

Torres C., R. et al., 86 (7-56b) 

Prevost, M. F., 1201 (2-114); 1301 (2-134); 1574 (2- Valdespino, I. A. et al., 254, 377 (2-88a) 

69a); 1698 (3-1); 1806 (7-2) 

Vasquez, R. et al., 2411 (7-53); 2412 (2-10); 2729 (2- 

Proctor, G. R., 41577 (7-83) 

5); 2788 (6-41a); 2808 (6-7); 2821 (6-35); 2898 (2- 

Pruski, J. et al., 3205 (7-1); 3352 (2-63); 3381 (2-18); 26); 2949 (2-52); 3168 (7-40); 3213, 6050 (7-36a); 

3354 (7-56a) 

6648, 6694 (2-8); 6769 (2-124); 6780 (6-6.1); 6832, 

Puig, H., 10256 (2-35) 

6856 (7-5); 7150 (2-144.1); 7192 (2-62); 7446 (2- 

Queiroz, L. P. de, 1825 (7-56b); 1827 (7-20) 

69a); 7457 (2-39); 7517 (2-69a); 7610 (2-140); 7615 

Ramirez, J. O. et al., 28 (2-114) 

(2-88); 7622 (7-41); 7628 (7-40); 7660 (2-140); 7679 

Riera, 49 (2-77); 450 (6-15); 514 (6-la); 586 (7-50b); (7-5); 7868 (7-56); 7909 (2-27a); 7916 (7-40); 7926


(2-69b); 7963, 8031 (2-69a); 8098 (6-36); 8112 (6- 

41.1); 8177 (2-5); 8188, 8203 (2-43b) 

Ventura, E. & E. Lopez, 781, 811, 838 (7-56b) 

Ventura A., F., 20795, 21298, 21541 (7-56b) 

Vera, S. E. N., 215 (2-68) 

Viellescazes, A., 447,471, 502 (2-77); 513 (7-50b); 534 

(7-56b) 

Villiers, J. J., 2246 (2-114); 2729 (7-15); 3719, 3780, 

3781, 3786, 3787, 3788, 3797, 3830, 3843, 3859 (2- 

77); 3884 (7-16); 3947, 6064 (2-77); 6603 (7-16) 

Wendt, T. et al., 3373 (2-32) 

Werff, H. van der et al., 540 (2-13.1) 

Woodbury, R. O. et al., 1-71 (1-1) 

Zanoni, T. et al., 11795 (1-1); 29406 (6-36a); 32064, 

32446 (1-1); 32568 (7-36a) 

Zarucchi, J. L. et al., 3593 (2-69b); 4889 (2-88) 

Zaruma, J., 558 (7-53); 753 (6-36); 809 (7-50b) 

Zona, S., 136 (2-1)

90 80 70 60 50 

0.b o o s 

50m --------


_? ^^*60 . ._____ __60_ 

V-' cs' C. cuspidatus C. venezuelanus 

IS 

FIG. 2.DtitooCrsa 

?C4 

""lau u C e0 

_ _ _ _ _ _ _ _ _ _ _0 0 

0 HABITAT 0 HABITAT 

Slope forest _ ______Terra firme forest _ | 

JFIM1 AI - stibuJti AO fN o C a ui JF MAIMnd C j nAISIOn D 

FRUIT I I I III I IIIIFRU IT r 

FLOWER_I I I I 1 I I I - I/ IFLOW W ER I 

_60 

1 60 

FIG. 21. Distribution of Chrysobalanus cuspidatus and C. venezuelanus.

Distribution Maps 

80 70 60 50 

0 

/^\ ^ L. affinis 

0 

1 o 

FRUIT I 

80 70 60 

80 70 60 50 

I\ ~L. alba 

0 

'" 

HABI.itAT.o~ -y -^ 

Terra irme forest 

FRUITM*ii I /.. I 

. . .. 

80 70 60 b 

FIG. 22. Distribution of Licania affinis and L. alba. 

0 

10 

129


10 

60 50 60 50 

0o L. albiflora 0 1 L. amapaensis 

0 

00 ~ ~ ~ ~ ~ 0 

1S- 

0 

--------------- ~------ 

RUIT aifra L. 

70 60 50 

FIG. 23 Dstbution of Licania albilora, L. amapaensis, L. angustata, and L. anneae 

HABITAT 

Fi-- 

~ 

.\ HABITAT . ... 

. 

Terra firme forest - Seasonally floode orest 

J FIM A MIIJIJIAIS INDJ 

'1J 1~~FRUIT~ ~ ~ l ~~ ~FRUIT ~ 

CFLOWER:::^: JY~ 10 

IFIMIAMIJ IJ IAIS OINID 

I I I I I I I I / 

FLOWERi i I II i il 

Iig L_ 

i 

70 60 60 __ 50 

FIG. 23. Distribution of Licania albiflora, L. amapaensis, L. angustata, and L. anneae.


Ic 

ol 

70 60 50 40 

L ^ 

o- 

CC^""'^ 

L. apetala var aperta 

Savanna margins 

Seasonally flooded 

_____ 

forest \' 

+ v qf \ 

v^- |/ 

/'1 5 \\\/) 

/ \ /} ( ; /- \^ 

\ /^ 

10 

JF \FM /A/ J/J A 

FRUIT d 

FLOWER I ************ X | 

IO II I 

s\< | / 

I 

\ N^ 

: 

v 

\ 

Savanna 0 

10 TT^ 

70 60 50 40 

~--L. 

i 

apetala var. apetala 

( 'u0 

FFABITAT 

Beaches 

24. Distribution of.Licania.apet 

Gallery forest" 


}\ 

Seasonally flooded forest 

"JFMIAIM 

J JJ IAIS O0 N Q 

FLOWER;: 

70 

- 

60 

.... . .. 

50 40 

FIG. 24. Distribution of Licania apetala. 

131


I' 

60O 50 __ 

' L. apiculata 

.----- 1 0 

60 

L. aracaensis 

lC?--- , 

10 

_ _ _ _ _ _ I o . _ 

_ 

i 

ABITAT I BITAT I 

Beaches ( , Montane & Cloud forest (. 

FRUIT 

7JF_MAjJ J A S CN1Q _ ( 

I FRUII I I 

J FIMAIMIJIJIAS IOI NIDI 

I I ) 1 

FIG. 2 D , a .. . ar 

"I iI 

50w )50 

Terra firme forest __ 

L ~ JIFIMIAiM.JoJIA 

" _ S_ND. \ 

0FRUIT T 

' ^\ 

1111~1 1 1 1 1 1 

FLOWERR 

/\ \^ 

& 

S ( 

.../ 2. JF.MAIMJTJ IJAIS.N 

/ I LIFRUIT :l 

11 i 

10 1 1I " 

L I I/ ____ 1 

1 

70 60 50 

FIG. 25. Distribution of Licania apiculata, L. aracaensis, L. arachnoidea, and L. araneosa. 

50


100 90 80 

-.^r^^'f "VL. *arborea 

0 / 

0 0aOo: 

Dry forest, semideciduous FIG. 26. Distribution \ of Licania arborea. 

IJIFIMINJIJIAIS 

?~~~~~~~~R)rj~~~~~~~~~~~~ 

? ? 

SIOND A 

100 90 80 70 

o 

h 

133


50 40 50 40 

arianeae L. bahiensis 

./-~. ..- 

0r 

J 

-------- - -- 

-^-.."'"^^' ' l 

IF_IMIAIMJIJIAISIOINIDI 

T r forest 

Terra firme 

FRUIT | | | | i 

| 

I I I 

i i i | 'O HABITAT \ :|/ 

IVJ IJJASS 

F 

IND I 

2 .FLOWER I I I I II! I 

I 2: 1 

E 

1Te firm 

2C FLOWER * 

? ...MIA~JIJAI SIOINlol ,.f......I\.SIN 

I I I JFABWJJAT 

forest___...... ~~';Terra - firme forest - 

~"~"~ZIIg'0 

n ^ 

--H--444--^---J>:FW I 11 1 

\ V - I0 _- // I FRUIT I 

: I 

40 50 

FIG. 27. Distribution of Licania Licania arianeae, L. bahiensis, L. belemii, and L. bellingtonii. 

.


V 

o o 70 ITX 

Gallery forest 

70 60 50 

?_60 ' / 

5 h_L. blackii 

Seasonally flooded forest \ ) 

'" 

JFMAMJJ M I J......?/.. ASOND .. JK s -? / 

FRUIT 0--^ | g l | | | | | 0 1@1@1 1 ) /\ s X -. |. . . 

FLOWER :::ggi^ ,. 

m., 

ylUUl | , J 

r 

- 70 ___60 

/ \ 

i 

50 

( 

J 

> 

. 

( 

70 60 60 50 

",^L. boliviensiso 

L. boyanii 

Terra firme forest White sand forest or campina > ( 

FRUIT .AT 2 D it i a FR IT,. b nJ L. b n 

FLOWER MN I II :II 1 g _ 

>s FLOWER I _ :................. * 

770 

r~~~~~~~~~~~~~c~',- 

FIG. 28 sr D i o f bto Li c a b i cii L.blvess , n.byni 

~ 

bU -50 

FIG. 28. Distribution of Licania blackii, L. boliviensis, and L. boyanii. 

135


80 70 60 50 

T 

IIw ' 

?" 

- -- 

^ 

L. bracteata 

"JIF AIMIJIJIAISIOIN 

_ - D Y 0\ ( S ? 

FRUIT I I /I I I , I ,\ 

FLOWER, 

* 

*- 1T1z1 I _ y 

80 70 60 50 

L. britteniana 

Terra firme forest \ / /// 

Seasonally flooded forest \ 

80 70 60 50 

80 70 60 50 

cli S^X -' _. -L. buxifolia 0 

White sand forest or campina _ -' / /- .'- \ 

FLOWER 

- I _ 

" __ \;\( > ---n 

_- 8U - 70 i60 U 0 

FIG. 29. Distribution of Licania bracteata, L. britteniana, and L. buxifolia. 

|


H 

L. cabrerae L. caldasiana 

o0 o 

0 

Montane & Cloud forest T o te ( 

JF MAM,J A S C5ND 

J 

FRUIT | _ | X X IR T \ 

0 

''''^^ 71 

. __ 

70 80 70 

F L OWE-R I I I FLOWER~~~ 

8070807 

80 

__0 

70__ 

80 70 

FIG. 30. Distribution of Licania cabrerae, L. caldasiana (exact locality in Colombia unknown), L. calvescens, 

and L. cecidiophora. 

137


Sersonally 

70 60 50 

L..-. aL. canescens 

f looded forest X / 

) 1 / x \ f 

\ 

~\\ZZZL , S^MiÂsonD/ I- / 

..................... ^.^ , 

Licaiaan1e 

FI.3.Dstiuino 

FRUIT _*_* ____** .. l 

\ . / 

FLOWER ___W * __ '* 

/ \ 

FIG. FI.3.Dsrbto 3 1. Distribution of fLcni Licania canescens aecn and n L. .cuaa0 caudata. 

' 

p '


60 

____80 70 

^10 ^L. coriacea 0 

;yL. 

_ 60 0 

_ 800 

o _ __ - _ __ _ __ _ 6 _ 

_0__ 

oTerra 3 rine forest ain A 

c i 

costaricensis 

6L0ueioi0 

L 

FIG I. IIoI o e I . I I I I a L a i 

FIG. 32. Distribution of Licania co uepiifolia, and L. 

crassivenia. 

6 

139


80 70 80 70 

L. chiriquiensis L. chocoensis 

6^ 

0^', 

v 

) " 

0 

HABITAT 

Montane & Cloud forest __ Terra firme forest'- 

U /. "1"1 

I I I I I I FRUIT F I U I- 

60II 6070 

0 i- L. compacta I o 

. -.10.. 

0 0 6 : 

FIG. 33 Distbution of Liania hiriquiensis, L. hooensis, L. ompata, and L. ordata. 

F~~IG~~',. 3.D b f a u L. chocesavannas,Loata. 

FIG. 33. Distribution of Licania chiriquiensis, L. chocoensis, L. 

compacta, 

L. and cordata. 

o

Distribution Maps 141 

_70 60 70 60 

L. cruegeriana L. cuatrecasasii 

Terra trine forest M & Clu fores 

|FLOWEFI| I II I 101E _ 1 1 

I A 

60 5 70 6 

00 

I 

? 

o o~~~~~~ 

o o~~~~~~~ 

B I ^ 

7"I 

^ " 

^0 

FI.3. itibto f iaiacugein,L.careaaii .cpra ndL usiaa 

FIG. 34. Distribution of Licania cuatrecasas,i, cruegeriana, L. 

L. cuprea, and L. cuspidata. 

6 0 

_^g_


L. cyathodes L. cymosa 

HABITAT 

/~J/ ~~I~-~ 'THABITAT " 

f 

Seasonally flooded forest Terra firme forest 

IJIFIMIAIMIJIJIAISIOINIDI 

) I | IJIFIMAIMIJIJIAISOINI DI 

FRUIT I I I i1 I 1 I- 1- / - IFRUIT II I I II 

[FLOWERI 1 Il I I I I I I 

L. davillifolia L. dealbata 

Terra firme forest HABIT 

Seasonally flooded forest Cerrado 

FI I3 I AIMIJJ I AISIO N D i 

FRUIT I A 

thJoFdL.ALMsJaJ IAaSIOA NID L 

J- 

1 I / 

FLIWERFLOWER :: :g lI 

i FRUIT 

!_1FLOWER : 

I 

g 

I 

I I 

FIG. 35. Distribution of Licania cyathodes, L. cymosa, L. davillifolia, and L. dealbata.


_60 _60 50 

Terra firme forest __ ( \ 

JIFIMIATJIJIAIS IONID / /) 

I E * 

\0 

I 

on ntTerra M fnrme forestudforest 

IJIFIMIAI^^J IIAISIOINDI /-^1 

FRUIT I I I I I 

I1 1 

10 FLOWER I1Ig g11 1 , 1 X,^ i . FLOWER _ 

60 60 '*60 50 

FIG.^^ L. divarica ra . ds durifolia 

1 I1ABIT AT 

F 60 50 80 70or Ld_ c L d l 

Savanna_______________ It ) / I ABITAT 

Terra firme forest- rrMontane e Cloudf & forest _ 

JIFIMIA - 0JJ JASND-- JAFIM - NJIMAAs, -\ 

, AM , J 

D / 

FRUIT 

/y / 

- 

j 

IFRUIT 

----I I 

--I-I/I 

FLOWERE *gg g l WWI ....... I FLOWER gI I mm I gI 

60 50 80 70 

Lica0ia FIG. 36. Distribution of densiflora,L. discolor, L. divaricata, and L. durifolia.


's" 

- 

80 70 60_50 

--0 

f o; 

i . 

' 

' 

L egleri 

HLABITAT \ (''"*/ ^ ...... 

i^r-^Q......... . . . . 

Secondary forest t , : , 

g ? , -) - ) 

U Terra firme forest _ < - - / . 

,_ W / .\- - 1 

Seasonally flooded forest / ^ > I \ I t / 

JFMAM _ /A _ J J AS * N D 1D 

v \ . f| 

_ 

- 

FRU I I I 

o 

^ 

Ter im oet __^ j ^- U \ \ \ 

- - JFAM -- jA'O - D *'I / /\ N i^K/ ,f) ^ V- 

FR T 

o . A 

-g 

^ 

/--< \\ '\ I L^?/ 

I I I g |^, ( I \ ,) )_ 

80 70 60 50 

FIG. 37. Distribution of Licania egleri and L. elliptica.


/^P^; 

0 

7______0 

? 

60 70 

Terra|irmeforetTera firme forest 

10^^^-__ 1 

L. fasciculata fir L. lomenoi 

_LEFIMAIJ I I J lNIDI 0 

80 7IF0MI 

HABITAT MAMî^SQNDHABITAT 

8070IJ 

HABIT AT 11\ 

r H~ I \ 

I.ABIT 

JATf 

- 

rTÂ J )\\ ( 

| 

FIG.38. Distribution of Liania L.. emargnata, fanshaW ::i ::L.fa :ilata,andL. .......... 

70 60 60 --0 

80 70 80 70 

L. fasciculata L. filomenoi. 

UV'^ / ~ ":;t'RUI ~ ~ ~ ~ ~X 

' 

FRUIT | \ |Terra | | firme | forest / 

.1 

~~~T 

Terra firme forest- 

HABIT AT =2JIF5yAJAZ1A 

Terra firme forestFRUIT 

I 

FLlOW iER 

SIOINID 

FLOWER I - \ ' " 

7 ^ 0 

8 0 

____7 0 80 70 

FIG. 38. Distribution of Licania emarginata, L. fanshawei, L. fasciculata, and L. filomenoi. 

5 

145 

p


70 60 70 60 

10 o 2 

L. foldatsii L. foveolata 

'-^-^-^, ,~ .....-'-^-f i,'^' (e^^^--^J--------- - 

Savanna ____ _ .. HABITAT . 

Savanna margins _/ I Slope forest 

JIFIMAIMIJIJIAISONIDI J IFIMAIMJIJIASONrISI N 

FRUIT |Ai le | | [ 1 I 

\ / FRUIT 

. 

. 

10 0o 

70 -0 70 60 

60 50 80 7 

o o L. fritschii L. fuchsii 

? !U8_______7 

0IG 3 Dtuof fot,fvaLftiaLf 

HABIT 


FRUIT I9 ID I I I 

FLOW ER 101 I I I I1 

oI I 

I 

I f dI 

i i!1I 

f a I .fr 

W LO ER 

I I 

I I I 

80 70 

FIG. 39. Distribution of Licania foldatsii, L. foveolata, L. fritschii, and L. fuchsii.


7!F- 

70 60 50 0 07 

~.0. 

1 0 

Seasonally flooded forest ( ' I , y Terra firme forest / / / 


FRUIT 

FLO W ERI Jll 

II 

.I..!.i. ! 

6 0 50 5 80 70 

IlFRIT f |I | | | | ||II T _ ___ 

FIG. 40. Distbuton of Lcanfurfuracea L. gentryiand L. gardneri. 

147


10 

I 

. 70 ...7 

Q 

eL. 

4 

glabriflora 

FRUIT * 

ÛIT 

-


11Q___ i_m ______ ^ B M60 

L. gonzalezii .. L. gracilipes 

FRUIT _ _RUIT T__i__ / I 

FLOWER _i ll I _ 1 1i| II |..OWER..... ! - 

110 1(0 _ 

_60 

_ 

0 ~ 

80__________70 70 60 

L. grandibracteata d-- L. granvillei 

011 

FLOWERI 

I 14 I I! I 1 FOWER:: 

JIFMI 1MJ JIISO II D h(--l----- f 

ol77ArS~ i 

FRUIT I I I1 I II I I I I~j I-IIRU IT I j~ I I IJr-' ~cr 0 

FLWE J II 1 I1 I1~ "~~..:: ??Lnl IIL0 E 

~ 0 1 7 

0I.4.Dsrbto fL ozlzi .gaiie,L rnivcet,adL rnili 

80 ' 70 670 7' 

_ 

FIG. 42. Distribution ofL. gonzalezii, L. gracilipes, L. grandibracteata, and L. granvillei. 

149


0 . 

0 

80 70 60 50 

90 0 

'^ 

% ,? 

, ? L. guianensis 

0 z /t X 

=60(09(0000 6 

L. guatemalensis, and L. harlingii 

Terra firmme forest / \ 

HABITAT 

HABITAT 

Terra firme forest Terra firme forest 

I FRUIT I - _ T/ / / IFRUIT I l I el l I i le!l Io I ( 

I FLOWER| l JIFiMAuIMI J aJAISOINDI n o Ies L a LA.MJA i an lng 

FRUIT I V 

FRUIT I I I I 1 I I / 

FLOWER ? 1 _ _ _ - ?/ 0 

IFLOWERJOIJ,I FOWEIIl?, J. Sl... '... -?" J} l mole O _ _^ 

FIG. 43. Distribution of Licania guianensis, L. guatemalensis, and L. harlingii.


700 

50 

0 

- ^ o?L. heteromorpha 

-v ? N ^ var.y heteromorpha 

.,-'Y .... 0'-^S?,< 4^^B^ ' . I ^ 

..... 

l. '^. 

FRUIT,;,,** ******* ^J0. . \ \ 

FLOWER .. . 

.. 

FIG. 44. Distribution of Licania heteromorpha var. heteromorpha. 

FRUITA T - 1 1 1 O y I


80 70 60 50 

o;-N , =_ U^-"L. heteromorpha O 

:'^ ' 

y //o I 

var. glabra 

60 50 0 

Seasona Il flooded forest . A Sao ly \\ forest ( \ \ 

10 FIMIAIMJ JJALS OINID ^\ 

--- 

^ --aij^ 7 p'-- 

1|FRUIT I I - 1 F I (0 

I I I IFLOWERIaS I \ I 

_60_ 8 

50 

HABITAT 

G var. perplexans var. subcordata 0 

Terraf foirm o e forest Sd 

60FRUTT * 

*"; . ..... 1RUT 4 k 

FLOWER u OW E.... 

FIG. 45. Distribution of Licania heteromorpha vars. glabra, perplexans, and subcordata. 

AI


70 60 650 

L. hebantha L. hirsuta 

I HA ITAT ^/y-Terra Iirme forest 

0 / 

Amazonian caatinga Seasonally flooded forest /i 

FLOWERI I e I I I I EI I 

E 

1 - I 

_ 70 60: 70 60 

FL. L. hispida , L. hitchcockii 

?1 

Montane & Cloud forest Slope forest 

0 

0 0 0 

FRUIT 1 IS I IFRUIT 1 * 

A 

760 7- g..70 60 

FIG. 46. Distribution of Licania hebantha, L. hirsuta, L. hispida, and L. hitchcockii. 

153


70 60 50 40 

^ ^ ^ ^FMAM ^ 

< *i (\^ L. humilis 

Terra firme forest I ) t 1 '- 

'FLOWERI _ 1 _ 111 g _ _111,_ I â 

I __/ _ 

2CiS~~~~~~~~~~~~ 

ITAT 

ASOND ****-*** 

...../ 

!!ss^^ j^/... 

70 60 50 40 

0 

i h 

FIG. 47. Distrbution ofLicania hoehnei and L. humilis.

90 80 70 60 

0 

00 0 . 

_jS^--- T^ t^l00 * 

HABITAT~~~~~~~~~ 

var.. hypoleuca (^(/ 

~JSavanna margins HABITAToveola var foveolatata 

1FRUIT IOI*IIOI@I@I@I*I' 

/{ ^ ^10 

FIG. 48. Distribution of Licania hypoleu.a. 

I 

HA BITGAT var. hypoleuca o h 

Savanna I 

Savanna margins FIABITAT, v ar. foveolai 

Terra firme forest ____ Terra firme forest 

____ V^l / 

JF JIFIMAMIJIJ AMJJA 1AIs AS NID \ F M 

***** FRUIT I(J A J A AS IONI 

FRUIT 101 10 FRUIT IJIFI--I ..,\I.IAIsMNIDI 

A0 

IFLOWERgS0 FLIII -1010 

ggg*0 

Z~LLI Jg* _FLOWER:^: \ FOWER 

^gI 0 

""J':^-~\ 

9080 50 ~~~80 70 60 

/ 

" 

FIG. 48. Distribution of Licania hypoleuca.


60 50 

0 0 L. impressa 0 0 L. incana 

Iv ,' 

0 40 

FRUIT I I~ I I I I I^ it I0 I In R 

p e 

FLOWER IG. 1D :: i I I Ln.... FLOWERi 

and 

50 40 __ 60 

4 D L. indurata( a L. inpae 

.. A . ........ . 

l~ ~~oy~r ~ ~ r 

^ 

y 

~-~. Gallery forest 

Montane & Cloud forest 

: 

JIFIMIAIMIJ 

I 

- - - IJIAIS - IOR NID 

RI T i H-11- FRUIT 

FL OWER - 


i L. intrapetiolaris o ^ 

\ J 

^.\ 

A y r / r 

1uTerra firme forest ____ ( 

J FMALMJJASOIN D 

FRUIT * * 

FLOWER R _i8"""_ _"_ I 

' 

T \ 

( \ 

\ ;^ 

I I I _ 

0 Soefrest 

JF_IA 

1 FRUIT _* 

I LFLOWER __I _ 

______ 

IAJjAISiO5ND 

fI 

\ 

A) 

1__ 

70 70 1J1 

60 

FIG. 5 Dsi t iaL n . f je i fe n ns it is p 

L . jim and e ne z ii 

FiABIT AT 

0~~~~~~~ 

Montane& Clou forest Terra fire forest 

JIFIMIAI14JIJIAIS lo(IN 

FRUIT 

FLOWER 

I i I I FIJEI 

FRUIT IrJIRU n I J 

14 

J IIJAIS IOIN D 

I I I 

OWER 

?T II I~l I 1.E I I I I 

bu 1_ 0 bu 

FI.5.Ditiuin fLcna nrptilrs rii,L. jefensisanL.jm . 

0 b 

FI.5.Dsrbto f iai nrptoais .iwni .jfnss n .jmnzi 

ez. 

157


0 

" 

7-O-? 

L. joseramAosii r I L. kallunkia 

_ I 

60 _ _ 

. 

^ 70 

FIA I Ftbi o iABliTATe L kru 

0 D 

FIG. 51. 51 Distribution of Licania joseramosii, L. kallunkiae, kallunkiae L. L. kiug klugii, and L L. krukovii. 

10 

700 

10


,M..,J 

C|: 

FLAITA 

70 60 50 40 

--- -----__ L. 

J7 

kunthiana 

........ 

FL 1 O00111 

7060504 

Seasonally flooded forest O 

0 

0 

F. . sbo fc a na 0 

i \ ~~ L1 I Y?\ 

`?-~ 

.i. 

... .. 

A'ON 

??,I~~~~~~~~~~~~~~~~ 

/ f 

o?/ 

........ 

?- ~ ~~ ~~ ~ ~~~ ~~~~~~~~~~~~~~~~~~~~~~~~~..... 

c.c? ~ ~ FG.5.Dsrbuino ianaknhaa 

70 60 50 40 

FIG. 52. Distribution of Licania kunthiana.


50 40_ 70 

0 L. lamentanda L. lanceolata 

805 70 0- 0 0 

Terrda trme forest . 4~~


op-AI 

70 60 50 

L. latifolia 

Terra firme forest ....... . 7. . 

J_FIMIAMjJSO J\ N D J 

FRUIT I1 

FLOWER," _;g I 1 Wv 1 1 1W |F V 

60 

m 

A / 

5O 

II 

60 50 60 50 

0 o 

/3^^- 

--lo --10 ^^ 

Slope forest HABITAT , 

Gallery forest___________ / 

IJFIG. 54. FasAIMIJ,adL D stbuin o Lcni 

FRUIT ji ~i I I 

I 

I I 

I 

OI 

FLOWER I__. J. ___ J J 

' , 

'-6-0.............. 

50 ' 

Terra firme forest. 

a . J 

FRUIT 111 III 

FLOWER _ J 

60 

J Ai SOINpDl 

I ! ! !!/! 

50 

FIG. 54. Distribution of Licania latifolia, L. lasseri, and L. latistipula. 

1 0


80 70 60 50 

.Tea firme forest :i . / i' 

/ 

J 

F 

FRUIT M.......... 

80 70 60 50 

FIG.L laoL. I leucosepala 

i 

FLO WER I I- L**'Jol 

I 

.,jyl 

Terra firme forest --- ~ : 

~. 

I 

..80 . ...70 60 50 

FIG. 55. Distribution of Licania laxifora and L. leucosepala. 

0 

I 

f


70 60 5 0 40 

0aN1 1 FLOER10 HABrI t TAT { 


)\ 7 

70 60 50 

_____)40 _ 

.0 r70 60 50 

Seasonally flooded forest i1 I \ 

. 0 

Gallety forest 

FIG. 56. Distribution of Licania leptostachya and L. icanifor a 

Seasonally flooded forest / p/ 

FRUIT 

80 70 60 50 

FIG. 56. Distribution of Licania leptostachya and L. licaniiflora. 

163


50 40 

| L. littoralis L. longipedicellata 

4 

010. / .6.0 L 

I. 5 

HABIT AT : o 

Restinga 

'-,J 

HABITA 

Sesn---al flo de foSeasonally I 

flooded forest 

JIF1MAIM J JJ IA S OINID IJ IA ISIO N D 

FLOWER _ 1!l I I I 

50 40 

iJ 

WER - 

"Y \))6 ?^ ^ ^ r'~l L.Jongipetala 

FRUIT II! ! !el F I | 


80 70 60 50 

FIG. 57. Distribution of Licania littoralis, L. longipedicellata, and L. longipetala. 

70


80 70 60 50 

L '. 

10~~,,,? ~0 



FRUI 1 

M I- I I I I I 

,k ^^- ~ L. longistyla 

/* ^ 

a u 70 60 50 

80 70 60 50 

FIG. 

/... "'. O 58. o Distribution of 


" 

L. 

Licania longistyla and 

\\ 

macrophylla 

uu.. 73 60 50 

FIG. 58. Distribution of Licania longistyla and L. macrophylla. 

0


60 70 560 50 

^ ?L. macrocarpa 0- L maguirei 

...................1 

1 o -~- 


50 40 60 50 

o L. maranhensis -JL. marleneae 

0 

... 

..........It I I i' 7~7"t~-' .. 

0 0 

20 HABITAT 


TR\fJFI 

Terra firme forest 

TAFIMIAMIJ( 

JIA IS / 

| 

50 40 _ ~60 50 

60 5 110 

FIG. 6 D L. maxiamaar\ L.m- L aa mexicana. 

FIG. 6~~~~0 

HABIT AT 

Terra firme forest H10 ABITAT 

IIIJIFIM "IASÔND J 

~Z ZZ l J MIAIMIJIJIA ISIO NID 

FRUIT 

iFRUIT 

i 

J 1 

JFLOWERi.! ,I!!I_I I I IFLOWER i/ 

60 

_ 50 110 100 

FIG. 60. Distribution of Licania maranhensis, L. marleneae, L. maxima, and L. mexicana. 

0


70 60 50 

i fo oI L. membranacea 

Terra firme forest \ ) 

10 

FLOWER ___ _ *_ I \ ** I A /I / * * : 

...........10 I 1 

60 50 

90_______8 _____s60 50 

r - L. michauxii L. microphyllal10 

3;0 3 00I 

oABITA HABITI 

FRUIT T-II I I 1 

.WE 066 g I fFLOWAERg Ig 

FRUIT 

I 

: 

/ / 

YU 80 60 bU 

FIG. 61. Distribution of Licania membranacea, L. michauxii, and L. microphylla.


60 60 

0. L. laevigata LL. nelsonii 

1 

0 

HABITAT HABITA 

Terra firme forest /\ Swamps j \ 

IFRUIT 

FLOWER 

NDA-| I/ M 

I: 

\ 

\ I 

FRUIT I I |I I 

FOWER' I\ L I 

I 

I 

I 

- 

I 

60 

G. . Ditibton o c 50 

l a . o L c 

70 n . 

0 

-- - ------ 

L. occultanso L. tambopatensis 

? 

' 

1 0 

"" 

FRUIT I::I: ::: 

60 50 70 

FIG. 62. Distribution of Licania laevigata, L. nelsonii, L. occultans, and L. tambopatensis. 

:


70 O 

508 

0 ! 


L. micrantha 

FRUIT 99 i? 9 

FLowERgs I j4N-1 0 '0 

70 

-^ ?/ /'^ 

60 

^ 

/"**..-...-.o... :' j 

50 40 

I 90 r _ 

LI 

^ 

HABITAT0~~ o 0 

^ 

-)A - 

FLOWER 

0 80 70 60 

FIG. 63. Distribution of Licania micrantha and L. minuscula. 

0


70 6050 40 

- 

~'-ye 

" ei::, O-L. .... 

minutiflora 

TERr I r f t 1 1 . 1 - t.---E-l - 

I 

yj~ -..... ':....L ....1/l' " ' 

0 Terra firme forest 

.. .... . 

"JIFIMAMJIJ'IISIOI'I 1 

_ 60 50 40 

FIG 64 Dstrbuionof icniamiutilor ad L mltoii 

miltonii 

0 _ 

F I G. 64. Distribution minutifora of-Licania 

and L. miltonii. 

10 

171


70__ 

L. montana , o 

f ) 

70o_ 

HABITAT 

FHABITAT 

Beaches 

Montane & Cloud forest / Seasonally flooded forest . 

oxw L. mollis 

JRFIMIAIMIJIJ oAISINIDI1 1 FA J IAIS 

I 

FLOWERUI i I I I I I 

FLOWER g "I 

Il 

FLOWER 

- 

70 

U 

D 

W 

80 _ 50 40 

L. morii y L. naviculistipula 

: 

f 10 

FIG. 65. Distribution of Licania montana, L. mollis, L. morEi, and L~~~~~~~~~~~~~~~......... 

....n u l 

HABITAT BITATI 

Terra firme forest, Terra firme forest 

J. FMA F MJJA,M JJ ,A S ON, J IFIMIAI J IJ A - IS FND 

FRUIT FRUIT11- 

FLOWER FLOWER ' 

I I I 

? 

7 0 o 4U 

FIG. 65. Distribution of Licania montana, L. mollis, L. morii, and L. naviculistipula.


70 50 40 

L._niloi 10 L. nitida 

_' 0 

HABITAT 

T Cerra fr fIrme forest Or 

HABITAT 

rrado 

IJ FMAMIJIJASI JIFIMIAIMIJI-J IAISIOINIDI 

FL I OWERI +...gg.. L______ 

10 70 

40 

80 70 60 50 

?/,){j^l ?~"-'v-~} 

L. oblongifolia 

o --~ . , . ,,.: e ,. 

.......... .. ..... 


FRUIT 

J FMAIMIJJIASOI NIDU 

I 

iee 

illlle ;.0-" 

,, ',:, . 

o)-.... 

...o 

......... 

(' 

......./ ' 

80 70 60 50 

FIG. 66. Distribution of Licania nilo, L. nitida, and L. oblongifolia. 

173


70 60 50 40 

, I. 

,C L. octandra subsp. octandra 

.. . . 

.._.J~-':MAMJ /' D :: < / . ........ 

70 60 50 40 

.......... 

HAB T ... ...... ? 

Gallery-|@/,O 

IGallery f, 1 O fI 

forest . / "


80 70 60 50 


Jz 

r L . o ctandra subsp. pallidal 

D,.I > \


0~~ 

70 60 _ 60 50 

0 

0 

670 60 50 

)Jo^ 

o ,.^~~~ . . 

%L. 

pallida 


lFI 

FRUIT 

FLOWER 

AMJ IJIAISI 

M 

F) w i 

D \i \ 

\ 

a 

i.) ~-/( 

--? 

\ 

. 

V 

< ; 7I 

/ 

_ -770 60 50 

FIG. 69. Distribution of Licania orbicularis, L. ovalifolia, and L. pal/ida. 

pallida.


70 606 _ 0 

, L. pakaraimensis L. paraensis 

12 C 

?TFRUIT 

HABI'TAT /J^-r ^ ,/4 ^,._ally 

flood d f r s _ 

:3!MAMliASOND^-^-'.î/ 

?r o l *** 

[ijgsissftg-^ 

8U 

7 3 

* - 

.^^^\k)/ 7 

^ ^^ 

X . . 1 

^ _ 

Jff/\77z^^:-7t^^^~~~,? 

( A 

/ iO^w /A 

' ^ ^ -^ ^' / 'M 

/ 

/^ 

M S onalne FIG 70 flooded itiuino forest iaapkriess .preni,adL avfoa 

\FRUIT/ of |L1n a L aap/ / 

FIG. 70. Distribution of Licania pakaraimensis, L. paraensis, L. and parviflora. 

177


0 

o 

'r1 

Iz 2 " 

v v 


70 60 50 

~ 

F A J A [ N 

I~~~~ ~~FRUIT J|F|M|A*MJ|J|A|SI y........ 

}\ ~... 

FRUIT 

x, L. parvifolia 

r.' '""' .... 

. 

. 

X ' L 

Gallery forest ~ ~ 0 1 

) / 

7O, ~~~ 70 

- 

60~ 60 0 ' 5 50 0 60 

,IG 7 parvifructa L. 

L. a persaudi 

FIG. 71. Distribution of Licania parvfolia, L. parvifructa, and L. persaudi. 

e1 

HABITAT~ 

/ / 

/~ A 

~ 

r :erra firme i.Terra forest ___ > 

firme forest _ 

FIG.__ 6 

7iboiaiapf,L________6 

60


.. 

A 0ITT 

100 90 80 

_ - ----- 


IFIMJ A|MIJ IJIASlO NID I 

. .. ..L. platypus 

_8_?__10 0 9 0 70 

__ 80_ _____70 60 50 

FIG. 72. Distribution of Licania platypus and L. polita 

Seasonally flooded forest 0/ / ? >\ 1 j *\ 7 ] / / 1 

JFMA JJASO ND // / 

_ Y t00 . ) ( 

(


50 50 

Terra firme foreso t :; ___ *'"' ........5"y " " ^ S ...easonally. . flooded : forest 

FRUI____T_I!i 

M 

FRUIT 

iM 

FLOWER i _lll _ 

_FMAIMJlAIS ASN N 

I _ ! 

) 

/ 

I 

I _ 

FRUIT 

IFLOWER /i _ 

50 lOiND 

/ i 

0 

,ro^ 

..5050 

70 60 100 90 

L. pyrifolia . .--.- . 

7_Bo~~ 

- ..---, 

HABITAT 'ABITAT 

Gallery forest Gallery forest 


0 

.' 

L. retifolia 

J IFA 

FRUIT I i 

IMJ JASOND jNID 

FRUIT 

J AIS NID 

FLOWER FLOWE 

m7 

o 

'_ 60-uo u90 

FIG. 73. Distribution of Licania piresii, L. pruinosa, L. pyrifolia, and L. retifolia.


70 60 50 40 

^ ^ .or.r ?. .L. rigida2o 

"? 0 

o 50 4 

Arid formations, caatinQa ai .... 

/ 

FRUIT 

I / ' ~ o L. rigida (cultivated) 

0 80 70 60--- 

FIG. 74. Distribution of Licania rigida. 

~'" 

" 

181


80 70 60 

.0*^ ^^ o Y L. reticulata 

0 

Terra firme forest - -- 

Seasonally flooded forest - - 10 

J A S D I 

;FRUIT * 7 ) \I 

,,__70 ? 8^ 

-60 50 

50 40 50 

L. riedelii L. robusta 

10 

0 

'HAAHATT 0 10 

""**....^ ^ ~"~\~"~~'~---^Terra firme forest 

FIG. 75. Distribution of Licania reticulata, L. riedelii, and L. robusta.


60 50 

I 

60 50 

.: L. rodriguesii L. roraimensis 

10'~~~~~~~~~~~~~~\ I~?c-1 

HA^^BITAT1 V A ( ILHABITAT F 

10 

Terra firm st ontane Cloud C d forest f 

J F1AMJJASIOND )/.ONDMJ F J S ND 

IFRUIT i /e / I FRUIT _ ../ _ 

FLOWER * 1"i I *le* : I I I * . .J '" IFLOWER ! J 

60 560 b 

^_ _ _ _ _ _ _ __60 _ 

80 70 

FIG -. . L. rufescens o 

L. salicifolia 

Slope forest 

' 

) \ Gallery forest /, 

/ 

/ 

' 

JFMA'ND -"-- /[\ f ' I JIFMIAIMIJIJ 

--- ! IAI S NID./ 

10 FRUI T_ -------^ I1- -1 FRUIT I i I- /,;) I ~ 

PLOWgR~":::^îj|_j~~7r----lo IFLOWERI II I i i II - 

bo o L 7o 80 

FIG. 76. Distribution of Licania rodriguesii,L. roraimensis, L. rufescens, and L. salicifolia. 

o 

183


50 40 __60 

L. salzmannii L. sandwithii 

FIG. 77. Distribution of Licania sazmannii, L. sandwithii, L. santsii, and L. savannarum. 

Restinga Terra firme forest 

FRUIT IFRUIT II 

! I I i 

FLOWERI I i I i?iI 

L 

IW .. I I .... I 

50 40 60 

Rest5nga 4020 

Terra 

7mr0 

s 

50 40 70 | 60 A \ 

FIGL 7.DsrbtooLiaiLsadih- L. santos. . . L. . savannarum and 

b) 4070 (0 

i 

L. santosii -? L. savannarum 

FIG. 77. Distribution of Licania salzmannii, L. sandwithii, L. santosii, and L. savannarum.


Cerrado : ' 

70'Gallery forest 

,'~ ~l:~)~, 

f.,/~ ,t 

/ 

L."sclerophylla 




./ . 

. 

1\) 

\ ) 

, 

\ 

* 

I // 

_//j 

/ 

//// 

/ 

) . 

;: 

/ 

/ 

/ / 

JFiMAJJJASOND /I J VJjA: N 

; / 

/', I 

FLOWER F 

| 

________1 

'' 

--.( 

. 

: 

70 _____ 60 40-- 

50 

70 60 50 

.T) \^ / ? 

I~~L. silvae 

70 60 b 

FIG. 78. Distribution of Licania sclerophylla and L. silvae. 

50 

0 

0


5Q 40 . ... _90_ _ 80 

L. silvatica 

10,; , 0; V6610


60 70 60 

L. stewardii.. L. steyermarkii 

0 0 

' % . .... 1.o ... ... 

i I I I 

A I jFM wi MA/ 1 ]MIJIJAIAISO 

IStjr 101 ND NDJ \ 

- 

iFLOWER 

FLOWER I iJ 

S ND 

FRUIT _ _!1 I *(I ITI I I _ * 

60 

, L. stricta L. subarachnophylla 

....... HABITA 

60bU 

10 HABIT Gallery forest 

Seasonally floodd flooded forest Terra firme forest 

IJ IFIMIAMJIJIAISONID Ja 

FRUIT 

FLWR IFLOWER ": 

FRUIT 

I 

IFIMA MJIJIAISO 

I 1 l 1 

1H 

FIG. 80. Distribution of Licania stewardii, L. steyermarkii, L. stricta, and L. subarachnophylla. 

60 

.. 

187


o 

70 60 __70 

70 7 

-,... ..... 

^ 

HABITAT^ \ \^ J) 

HABITAT 

( '^^ 

Montane & Cloud forest N / Slope forest|M|AIM|J rS I 

- ~ 

- 

FRUIT --FLOWER Ig 1-----l-1 \ I FLOWERI II-- 

- -- I I I I I I I I I I 1 1FRUIT I I I I I I \/ 

:FL:OWER I I I: 1 1 ,I I ,FLOWER _ I I, : I I 

70_0 0 

_ /v^ L. teixeirae L. tepuiensis 

Terra Cfirmest Soe forest \ 

FIG. 0 81. Distribution of Licania subrotundata, teixeirae, L. tachirensis, L. and tepuiensis L. 

Terra firme forest ?, Montane-& Cloud forest 

JIFMIAIMIJ AISIOND / J F MAM JJAS III [JF ! nD 

AM IJIFIMIAIMIJJAIJSIÎSOIFNID 

S!qlNID ! . f I 

FRUIT I III 

I 

WER I I I MMI I 

FLOWER 

FIG. 81. Distribution of Licania subrotundata, L. tachirensis, L. teixeirae, and L. tepuiensis. 

o 

.


I?, 

60 so 

L. ternatensis L. tocantina 

1 00 

Slope forest 

\ ^^'Terra 

firme forest- 0 

JIFIMIAM J J AIS ONID 5J FlMAWMAJSJNA1r 1 

~ 

FRUIT i~*~ _ 

S_FRUIT 

FLOWER _I :ggg_ ff:I /t FLOWE _ _ _ _----H/ 

50 40 I 80 70 

I 

tomentosa, ,, L. 

HAFITAT BI / \ \ \ 

Restinga 

FRUIT- 

10 HAB ITAT 

LOWER! I IIi$11Ig$1 . I Terra firme forest 

/ 

FIG8.i 

4U 4U80 

s o tiu 

JIFIMIAIMJIJIAISlOtNID 

FRUIT 

L 

Linatnts,LtcnnFLOWER igid 

70I 11FLOWERI I I I 1N1 1 1 1 1:1 1 1


'^ 

80 70 60 50 

* 

.... . ... . L. triandra 

..,. ~ 

~ ~ ~ ~ ~ ~ ~ .. 

Terra BITAT~~~~~~~~~~~~~~~~~~~~~ 

firme forest _ \ \ I i \ ~ 

A 

J FMAltJ J AISIOND - 

I ) 

^ 

rl \ \ > ( 

FLOWER_III_* ! , ^ /^ S^\ V / L / //I I 

?U 70 60 50 

80 

___ 70 ____60 _______50 

~~ I~~~~~~~~~~~~~~~~~ 

"~~~~'~/~ ooj 

0!?-.:~- 

L.unguiculata 

FLW FR i-g o 

" " " _ (.; /^ v In \? /y// 

HABITAT " '""T 

~ 

FI Terra firme foresta . 

Seasonally flooded forest "\ r 

. 

. 

.. ... 

y 

^ 

( 

.... 

> / 

JFIMAIMJ J A SND^- )^ / V M t ( 

8U 60 50_ 

FIG. 83. Distribution of Licania triandra and L. unguiculata.


/L. turbinata ] L. urceolaris 

1 O 

u


80 70 80 70 

L. velata L. veneralensis 

10..-/ ~",~c~~,o11~....-10 /' ol10 

~ 

Io ____ J o . 

? 

...%~. ~ 

? 

-- 

o 

'-.J.=,--- 0, 

~, ~ :: 

HABITA^ - - 

^r^^V 


/^ 

'- 

^ HABITAT- 

Terra firme forest -- I 

l JIFIMIAIMIJ J O .'I 'JFI 

I F!1.FRUITII I I I '~ I 

!FLOWER Ill i I!1 lel a I ...... 1 

I 

:LFLOWER 

I 

I I FRUIT 

I I I 

I 

I PI!!".. 

_1 U 80 70 60 

0', 0 - 

L. wurdackii 

10~1 0 

10 0 

'0 o"'I 

i--L---4T.--0 

o o 

FL {, O A (X1 

JIFMAIMIJIJASND A\ .H , I I 

L ggE 11 11 1II!I I . ) t. I I f . (\ /I // )f v1 

8U ~ ~~70 60 

FIG. 85. Distribution of Licania velata, L. veneralensis, and L. wurdackii. 

0- 

- 

"- I


8 0 60 50 

CParinari Composite Map 

.0 0 

o 

0 

F W-EROO." *@ @-@'@ 

u.. I 

f 

8 un... D .fh 

0 

0 

O 

^^ Fs IV \ ^^ I\ ~ 

" ; ?~~~ 

................. e */-^^ 7^^ r*^>^ . .. 

h~~~~~~~~~~~~~~~~~~~~~~~..1 

30 FRUIT **** 0 10 ** ** , 

rY- / 

/


50 40 50 40 

P. alvimii (7|.) 

P. brasiliensis 

l J ~F s sJ A J S~.. i .. ..... 

Terra firm forest e 

_ _ _ 

FRUIT _ _--- 

FLOWER _r _ _ _ _ _;____ I I 

50 40 

201 HABITAT I 


4 

j 

70 60 50 

8 Dr on of Prri al P. campestris 

Gallery forest'- 1- 

10 FLOWER 

gigBigSgtigg^ /JTTJ7\)-"-7'-----^^ .....-^-) 

~70 60~ 0 

FIG. 87. Distribution of Parinari alvimii, P. brasiliensis, and P. campestris.


"''HEITAD FHA 

60 50 80 70 

P. cardiophylla 

0;~~~i o 

M 7 

;T 

0 

P. chocoensis 

10 10 

10* 

AISIOINIDI 

Terra firme forest y Terra firme forest 2__. 

JI J IAISIAOINIQI 

FRUIT II I I I I I I I I IFRU 

[FLOWER I:II 

60 50 

\ 

I I 

I 

! 

IT 

8" 

IJIFIMIA8J IJ|AISIO NI 

! ~... I I I i 

0 70 

80 8 b 70 

50 40 

70 

FIG. /.Dsrbtonof p. klugiio a P. i P. littoralis 

\' 0 10 

V^T \0 AT ;\ 

* HABITAT 

HABITAT, Restinga 

Terra s 

firme 

firme 

.t 

^ AF MI - J-J AISFOND IS 

FRUIT i 

v ' 

\o ov 

FLOWER Eir J 

\ 

Terra forest 

1 J J'FM A J J A S OINID 

FRUIT 

FLOWER 

2 

, 70 50 40 

FIG. 88. Distribution of Parinari cardiophylla, P. chocoensis, P. klugii, and P. littoralis.


10 

1~0 

o 

HABITAT 

0 

3C Terra firme forest 

______ 

70 605 

o 

P. excelsa 

. 

....... ___1_ _ _ 

FG89Distri b u i o of Pa rn a e xcelsa..?? r - 

JAFMAMJJASOND \ 

70 60 50 40 

FIG. 89. Distribution of Parinari excelsa. 

10


HABITAT 

....... 

P. maguirei P. obtusifolia 

............................ 

HABITAT 

JIFIMIA J JJIA SCN 

FRUIT _-- -- 

I I I - 

FLOWER = I I 

60 

D IXA 

I 

JIFI MIA_ ,71J,A S C N DI 

FRUIT * _"i * g _ 

FLOWERgg ogiggg1gS 

/ 

/ 

/ 

40 

7 ,o0 50 

FIG.'^ 90 Dsrbto P iP. montana 

! / ~ c . ..... 

~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~c~~ 

oJ 

F A J'" . N D .-' / 

...........-'t 

FLOWER" : . ... . . . . . . ... . 

Terra firme forest I 

FRUIT 9 

" 

* * ( ,71 f \ 7 A / 

0 

70 60 50 

FIG. 90. Distribution of 

Parinari Parinari maguirei, P. obtusifolia, and P. montana. 

montana


Semideciduous forest ___ . Seasonally flooded forest J< < 

IFRU 

0P'~~~60 

JF MAMJJASOIND/ 50 

TA_re____/'_ 

.- 

HA_ FRUI 

7 F 0 EAMJJAs . 

oND 

_i~_ _ _ ~//_ 

O )/ 

\11 

70 60 _ 

OWI. R ^P p/ l rv |If^li 

,~ 0 

P. par la P. P occidentalis 

^ ^ 

~~HABITAT-- ~7Z-:--74-JL_': /{ -^ HABITATY' ^ ^ '^^^ J 

FIG. 91. Distribution of Parinari pachyphya, P. parvifoia, P. pariis, 

P. and o cciden talis 

Terra firme forest Tera a e for 

FRUIT 9 i o P1 o a ay ---c FRUIT A a P id ai 

~ ~^\


FRU 

- 


' / / )/ ( 

P. rodolphii 

80 70 70 60 

FIG 92 Distribution of Parinari rodolphii, P. romeroi, and P. sprucei.ucei 

HABITAT HABITT 

Terra firme forest Seasonally flooded forest 

J,FMAM,1 JI1AISOD JF A J AS D 

FRUIT 

FLOWER 

' 

8U 70 70 6' 

FIG. 92. Distribution of Parinari rodolphii, P. romeroi, and P. sprucei. 

199


0. 

70 650 40' 

_ 70 0 60 460 

50 

Exellodendron Composite Map 

FIG 93 Exellodendron.-. 

Distbution of the genus 

; 

JFIM:iA J/ ]AND/ i 

FIG. 93. Distribution of the genus Exellodendron.


0 

70 60 .0 

"Y~ 

...:: _... E. barbatum 

1E I 

60 50 40 

7 70 5 40 

' '7 0 60 5 0 40 

FIG. 94. Distribution of Exellodendron Exellodendron barbatum and E. cordatum. 

201


70 60 50 

.o gr E. grE. coriaceum 

Gallery 

forest... . .' 

Savanna mar ins 


- 

' ...... 

.. . 

FRUIT _ i 

FRUIT - 

60 50 

50 40 50 40 

E. gardneri E. gracile 

FLOWER_I __ __ - 

-_ LOWER I I i I I I 

HABITAT^ ) 

? 

^--j~l HABITAT 2 

Cerrado_______________ Gallery forest 

JFIMAMJ IJA SIOINIDI cr JFM AMIJ J A S gOiND 

FRUIT 

FLOWER 

gm I 11 I I [- 

I8I 

--IFRU RUI IT ? 

FLOWER 

* I I 

II:II 

- 

40 4' 

FIG. 95. Distribution of Exellodendron coriaceum, E. gardneri, and E. gracile.

0 

___ 100_____ 90 80 ___ 70 __ _ __ 60 

\ -oniancaîtin aCouepia / 

r 

O0 

0~~~~~~~~~~~~~~~~~~~~~~ 

o~~~~~~~~~~~~~~~~~~~eF 

|Seondary&Clu forest ___ ______ - Slop ire forest 

\ - - --V iX - 3 .s^9. X / T ap |\^ X7. " 

|Becodr 

Terr fims fres 

Montane & Cloud forest~~~~~~~~~~~~~~~~~~~~~~I 

White sand fo /rest or campina 

forest 

____ \ 

SeaIsonally tlooded forest j- 

y 

^' 

^ 

\ ^^- . ^ 'f a ; I/ W 

j?| 

4 l &4. 

,/


60 504n 

10o --' C. amaralae 1 C. belemii 

10 HABITAT 

"" Ar 

(\ A \ i ) f Z"^?~HAI I / TATA j 

J| Jii J S- 

Savanna margins ) OI FRUIT _I ___ I 

Amazonian caatinga__ __ 

60 

4F5' 

70 50 40 

10 . 

C. bernardii 10 

C. bondarii 

' '7 0 ' 

'~~~~~~~~~~ 

FIG. 9 . Dsrbto fCupaaaaa,C eei,C enri,adC odri 

T1erra firme forest .. ......... 

IG. 97 J I Is o C m C b . / . . 

FRUIT .':" " '""""'4 

FLOWER. I I I II II 'M I


80 70 60 50 

^.~3 .: - 

C. bracteosa 

,,t_J___f^.f'y 

[FLOWER X - _ -. I 

*.jo .,, 

Secondary forest 


iA)l 

80 70 7 

60 

50 

60 70 60 

C. canescens 

" 

o? 

C. canomensis 

p' ^ "o o 0 /IiS ^ C 

./ '] 


. 

.\ 

Slope 

HABITA r i o o C b , c e adCcnoes. 

orest 

60 70 60 

' 

/ f 

FIG. 98. Distribution of Couepia bracteosa, C. canescens, and C. canomensis. 

205


50 40 60 50 

1 / C. carautae C. caryophylloides 

HABIT HABITAT--7 

Terra firme forest Terra frme forest 0 

FRUIT J IF IMIAMI 

( 

J IJ N I I J JIA S OIND 

-FRUIT I Ik II FRUIT I Im 1 /A 

IFLOWER M I I I I I 1Im1[1@1 1 I.L|| IFLOWERI I I I I I i11 1 I ' 

40 60 50 

60 50 60 

F 9. is C. cataractae andC. cidiana 

Beaches 

Rheophyte 


H/A "0 A 


J F M AMJAJIJoINAI 

FRUIT 

Li /I 

FLOWER_ IM01g 

/I 

I 1_J 

yJ 

....' FRUIT 

FLOWER 

FIMIAIMJIJIAISIOMNIDT 

660 

50 

FIG. 99. Distribution of Couepia carautae, C. caryophylloides, C. cataractae, and C. cidiana.


FRUBITAT';., 1 r / C. chrysocalyx 

' 

0 0" ' ' '- 

0 

' 

FLoWERI" -'--. z 1E1zzE1 ..... J ' :.. ? -e .v 



I J, F'A'J I I I 

FRUIT l 

FLOWER '"" iil l 

? 

AI 

,- 

, . 

. 

( 3 

l '".'''** 

/~~ /,. 

J 

o- 

''''_ 

- 

~ 

- 

' / 

. 

'1 

) 

: 

80 _ 70 W0 50 

540 4060 50 

.^ 

..0~~ 

FI.. C. 

Ducoarctatoua c , C. C. comosa 

\ 

^ 

- - ^ *K 

. .......... ...... 

..... 

..':' 

HA 

B T T 

) 

HABITATi 7^?P^~^--^ 

Beaches 

Restinga 

heophyte 

JIF^M2 J 

AFR|)|JJST-- 

J A S I--- 

FRUIT 

IF A IMI J IJA SO ND 

I 

--- 

40 60 50 

FIG. 100. Distribution of Couepia chrysocalyx, C. coarctata, and C. comosa. 

_ 

207

208 

60 60 

loi, C. cognata C. cognata 


-"-... ~-var. cognata10 7' (, var. major 

/: . 

?"- 

'.i> M. 

Savanna margins ( 1 0 HABITAT 

Secondary forest/ \ Terra firme forest 0 

J|F|M|A|M|JJ|A|S|O|NP \ 

JFMAMJ JJASO D D\ 

60 80 70 

0 C. cognata / C. dolichopoda 

var. membranacea 

Savanna ___ \) ) / \ Terra firme forest , 

JIFIM|AiJ|JA|S|O|ND /iD 

0'FRUIT -I 

I 

' 

1 i.........F 

|IFLOWER 

_IIII l I I 

) 

I 

I 

IJIFIMAIMIJIJIAISIONID 

I \I I 

FLOWER 

O6080 70 

FIG. 101. Distribution of Couepia cognata and C. dolichopoda. 

o 

(


60 60 

C. eriantha Acioa edulis 

XHABIToAT.......0 

HABITAT 


FRUIT 

JFIMAMJIJLAISoD 

I I 

\ 

\ 

FRUIT 

JFMMJ 

* 

J A S 0 N 

** 

^1 

FLOWER gg___ 

Oi_0 

I // //1 FLOWER _g__ _ __ 2 \)/J 

1 0^ 

70 60 60_50 

^0 

HABITAT, HABITAT 


.... .... 

JFIMA^ JIJIAISONID ASN0JFMA'J || 

FRUIT 1 1 hdIII 1 1 I T 

FLOWERI 1 1 1 1 1E\\) 

1 

I 

IAISIONID 

|IFLOWER 

/ ( 

60___ 70 

60 50 

FIG. 102. Distribution of Couepia eriantha, Acioa edulis, C. elata, and C. excelsa. 

0 

0


60 60 

00 0 . exflexa C. foveolata 

0 1o0o 

HABITAT 3 D o ui e ABITAT 

. i a C 

Terra firme forest \ TSeasonally flooded forest 

JIF MAMAIS.ND D i C. FfeaAIMI JJIAISd Cl 

FRUIT I I IIIIcIIII_I 

FLOWER 

I 6N 

-^ 60 . _ 

I0sioaa.oC. 

FLOWWER I(^ 

froesii C. glabra 

FHABITAT/........... 

HABITAT 


FRUIT _ n [f FRUIT 

FLOWER FLOW II I:I is/ 

60 

^^^^^^^^^__^^___60 

FIG. 103. Distribution of Couepia exflexa, C. foveolata, C. froesii, and C glabra.


70 60 50 40 

..:erra...~... ...fi...foe..::: 70 .. ;.~_.. ......f .^- ..... . .. 

,?-. . 

HABIT I I I I 

FRU 

7 ?60 50t?4 

ISO 

............... 

JIFMAM IJIJI A IN 

FIG. 104. Distribution of Couepia grandiflora an 

F ER W 

............ : 

I 

7060 60 5 0 

FIG. 104. Distribution of Couepia 

grandifora 

and C 

guianensis subsp. divaricata. 

..... 

211


70 60 50 

I o. . --. - C. guianensis subsp. glandulosa 

Seasonally floded forest 

o\3 

yo 

.,,.,- 

- 

FG 1C. guianensis subsp. guianensis 

0 0 

\ 

70 60 50 0 

FG105 DisrbtoofCupaginnisus.gadlsansus.uaess 

70 60 5O 

FIG. 105. Distribution of Couepia guianensis subsp. glandulosa and 

subsp. guianensis. 

. 

10


I:::' 

01 

60 50 40 

C. habrantha \ C. impressa : 

cabraliae 

f ' 0 ?subsp. 

' 

. . . . ..I.C 1 

HABITAT S V/HATT 

0 Terra firme forest TerrHABITAT for-e- 

Seasonally flooded forest / Restinga - 

J FIMAIM JIJ AISIOINID JIF1\M AMJIJ IAIS N D 

FLOWERgggJ_I ~ 

- 

_ I I Ii /j / TFLOWERUI R 

50 40 50 40 

C. impressa | .'.i , C. insignis 

subsp. impressa 

/ :.............. 

HiABITAt, Restinga 


2f-g ITerra firme forest 20 

IE 

FRUIT 

IJFIMIAIJ IJIAISIOINIDI 

I I I I I I IFRUIT 

I I 

IJFIMIAMJIJIAISIOINID 

-- -- 

FLOWERI |I I I I I I IMll-E25 

4U 

--- 

4U 

FIG. 106. Distribution of Couepia habrantha, C. impressa, and C. insignis.


~""~'~ ~.. , I///' ~. ~ 

............ ' ' 

~0'.. I-.~'~ latifolia 

C . 

HABLTAT C, HABITAT 10 

Seasonall flooded forest Terra R firme forest 

FT-MTA 

JO.1 I JIFIMIAIM J J J A SNIIND 

FRUIT , 

*__****,* I ! FRUIT I 

:::WER : 

__=:; _S_ _ \YVFLOWER 

E- -B 

IG C. 

l.ongipendulao 

60 50 40 

ICo longipetiolat C. 

Terra firme 

forestJ s 

T esringa f 

1 JFMAMJJ AI 

10 FRUIT 

IongIpenIa 

FLOWER 

60 

.....i.......... 2J MA 

0/ 10 FRUIT IM 

F 

J J AIS0 ND - 

I -I 

I: H-H 

40 

FIG. 107. Distribution of Couepia krukovii, C. latifolia, C. longipendula, and C. longipetiolata.


70 60 60 50 

1^^^C. macrophylla 0 1 C. magnolifolia 0 

.......... ... ......... 

i 0 

Terra firme forest..^ Terra firme forest ) 

20 . JFMA__ JASJOJ N D , _-2C 10 IJ FIfAIMIJIJIAISOND ,/ | 

FRUIT I _ _FRUIT 

I 110 _ _ 

I.____________________60 6_ 

d' 

_ 60 

0 

^C. maguirei marl. C. 'eneae 

0 HABI 

0I 

C magnoliifolia, C. magiei, and C maleneae. 

Rheophyte ________ _ ) ) / Terra firme forest // 

I JIFIA>JIJIAIO " 

1C IFRUIT 1 1 I L 

FLOWER 

X 

// 

./. z 

10 

I IJIFIMIAIJJASNDI 

IFRUIT I I IIII1 11111 

FLOWER I I I I I I 

f .1) \ 

// 

/ 

/A 

- I60 60 O 

________060 

FIG. 108. Distribution of Couepia macrophylla, C. magnoliifolia, C. maguirei, and C. marleneae. 

215


60 50 50 

C. martinii / LC. meridionalis 

. Terra firme forest 

JF|MAIMIJIJIAISE NID1 

FRUIT 

FLOWER 

XI 

JFIAIM 

FRUIT_ 

JJAS ND 

/ 

~ - FLOWER 

60 -50 50 40 

60 80 70 

F-IG. 

1 C. multiflora C. nutans 

0 

........-------- 

0 ^ ^ ^^ --\~~~~~~~~~; 

6O\ 5^-^/''- 

.0 

'0 5 O 40^ - 

HABITAT 

savanna 


IFCABITAT. 


F JG1 M0 D sibIAISoOIN uiD JIFa ji MAli J iSDloND 

FLOWERJ 1.J 1? -WE-7R "i IE 

I_,1_60 

800 

FIG. 109. Distribution ofCouepia martinii, C. meridionalis, C multiflora, and C. nutans.


.0 

80 70 60 50 

^ .0 S.^ o,o&tA c. obovata 

0A-- 


0 0 

....; 

' 

N ,,,/ ,,/ 

- 

1. 

)r 

\ / 

- 

FRUIT 

CRg 

- * : vf . A /1 \ / / / 

FLOWi ./ 1__/K //' j .J \ 

______ 80___ 70 60 50 

8F 1 tf701O i oCu60aObat ad50 

1: 

2 I 

Savanna__________-'-"I 

FIG.1 10. DistributionofCouepiaobovata npa rillo,l 

IFNIAIMIJIJIAISIOINID 

l 

L 

........... 

FRUITA Terra firme forest .-/ 

Secondary forest ________ -- x-^ -j .. . ..,.. 

80 70 60 50 

FIG. 110. Distribution of Couepia obovata and C. parillo.


~ 

C. ovalifolias C. parvifolia 

HABITATk P I IHABITAT 

~~FLOWERI g"g"~SMg" ......E| *FLOWERg"""" ,' 

.__........... 

T ) C. pernambucensis . C. platycalyx 

FIG 11 .. b fo ..... f C o . b s . 

firme Terra forest Montane& 

_ 

FRUIT 

JIFMIAIMIJ 

I I JIAISIO NIDI 

I -p- I 1 I 1111|1 

-II 

FIRUIT 

IJ IFIMIAIMIJIAISIOI NI -/ 

FIG. IFLOWER 11. Distribution I of Couepia :I ovalifolia, I I.1-. I C. II parvifolia, FLOWER C. pernambucensis, and plat70calyx. C. 

40 

FIG. 111. Distribution of Couepia ovalifolia, C. parvifolia, C. pernambucensis, and C. platycalyx.


_ 0 70 60 

?k?^Vy 

0 Bec 

/.es~ 

C 

.t^C. 

.,/. 

paraensi1 


V 

300 ( 

^] 

HABITAT 

100 90 80 

C. polyandra 

Gallery forest _ 'o 5 1 

Terra firme forest \ / 

IJFMAMJJAS I i I ND 

f 

FRUIT gi _0I , 

/ 

110 100 90 80 

80 70 6030 

HABITAT \ 

10 Terra firme forest - 

C. racemosa 



'/ \ L 

White sand forest or campina / \ / /// 

| tJ]FMMIAIMIJM|J|ASS ) IQI A ) \ 

L1I 

\ 

L 

FG80 1.Dsrb60 70 

50 

FIG. 113. Distribution of Couepia polyandra and C. racemosa.


80 70 5 

C. recurva C. reflexa 

'1 O0 

0 HABIT^__AT RinHABITAT 

Montane & Cloud forest Terra firme forest 

JIFIM_AMJ J JASOND JA 

FRUIT ____I _i_ 

FLOWER _ * 

_ 

/ 

/ 

_ / _ =_ _ 

IJ JFIMIAIMIJIJIAISO ND 

IFRUIT I I 

IFLOWER _ I Il G! 

/ 

/ 

I u 

70u 

60 50 50 40 

FIG. 14. Distribution of Couepia recurva, C. robusta, C. refexa, and C. rufa. 

^V^ V HABITAT 

HABITAT 


Restinga 

^ 

__ 

~ 


JFIMIAIMIJIJ IAISIOINIDI JJ'' '''t JFMAMJJASJND A - 1 - - 

FRUIT I I I I I /I IFRUIT I 

LFLOWE^ 4 : 1 i"i:iii ll FLowER-g I --: 

bUG 14Dsiu 50 ru 40 

FIG. 114. Distribution of Couepia recurva, C. reflexa, C. robusta, and C. rufa. 

g


~_______0__________ 50 40 

C. sandwithii C. schottii 

10 

0 

c. scottmorii 00 c.?i~~~~~~~ ..... s 

\ 

HABITAT J , 


./ . . .HABITAT 

I Terra firme forest 

F 

JIG IM 

AtJIAISION D 

\s IhIFI IJIFIMAIMIJ JIAISId NID 

FRUIT I i I I I 1 I I 

FLOWERI i I ll I i 

60 

FRUI I I T 

ll I FLOWERS 

1_ 

40 

.__ 80 70 60 

FI 1 C. scottmorii and spicatai C. 

HABITAT HABITAT .. 

Montane & Cloud forest 10 Terra firme forest. . ......... 

FRUIT I II 

FLOWER ::::::g:FSII:: 

II 

ler 

I I FRUIT HIT 1/ 

-LOWER 

80 70 go 

FIG. 115. Distribution of Couepia sandwithii, C. schottii, C. scottmorii, and C spicata.


10 1o 

60 , 

C. steyermarkii C. stipularis 

I - 

( 

Montane oud forest\ Terra firme forest 

FRUIT 

_- 

--- IFIMAIM JAso 

I I I 

_ 

ND 

_ 

/) 

- - 

I 

_ 

/ FRUIT 

___ 

JIFIMAMJIJ IAI S 

Il /I) I 

I 

N \ 

60 

60 

80 70 60 50 

{err/ ,;M:jAO: 

JFR MA! l i IAI 

-: 

/y'. 

I 

' 

\-'. sub/rd C. 

80 . .o 

o 7 

50 .. 

FIG. 116. Distribution of Couepia steyermarkii, C. stipularis, and C subcordata. 

60 

223


50 

70 60 504 

HABITAT 

C. ulti 

^ ilFMIMIJIJ (A)S\ N 

--- -; A 

2 

FRUIT | \TELw R: BiiSjai5i^.)- 

|."._ 

|N|D| . 'I 

-.-._ 

t '-- 

_ FLO\VER_ : g B 

-__ 

_ ___ 

-> 

/ 

FIG. 117. Distribution of Couepia uiti and C. ulei. 

^ -~^ 

80 70 60 50 

FIG. 117. Distbution of Couepia uti and C. ule.


60 50 40 

C. trapezioana f /jC. venosa 

1O ' 0 

10.HABITAT 

- 

--$--hOl I HABITAT 

0^nffi^flS^ 

t030 ^ 

FLOWERI I I I I I I I * I I I L\ FLOWER g I 111 _ 

FI.180 Do 70no 6CtCvna adCwlimi 

i 0 

60 4U 50 

80 70 60 50 

225

100 90 80 70 60 

----------- 

HABITAT 

10 Rheophyte 


0 

------------- H 

A 0;S 0 

10 White sand orest or campina Atazonian caatinga 

avanna margins 

Savanna 

Cerrado....... 

. 


Coastal thickets ____ 

Arid formations. caatinaa 

2 White sand forest or campina 

Terra firma forest 

\ - - 

Gallery forest Distributionoe 

margenus 

Slope forest 

Beaches 

Restinga 


FIG. 119. 

100 90 80 70 050 

FIG. 119. Distribution of the genus


in 

70 60 80 

H. adderleyi H. adenophora 

7.\^-^^-^g^ ______^ 10-- r,, - ----- T. - - - 

0. 

JIFIMAIJM,A SOND . ? ^ VIFIMIA JJ AS N ID 

FRUIT | IVE (I|FRUIT |II ; 

II 

! 

I I 

- FRUIT 

1, 1.|,/ I! 

, 

FLOWER 

^^ 

- 

ggggssass8-- 

US^^ 

\ 

- 

--O 

FL l 

IcT OWE0 T I 

0 

I- 

10 

90 800 60 

10 

90 80 70 

G. 120. Distribution of Hirtella adderley, H. adenophora, and H. americana. 

FRUIT.i 

FLOWER 0i 

O ,6 o ~ ~ 80706 

FIG.120 Ditriutin o Hitela aderlyi,H. denphoa, nd . aeriana 

FIG. Distribution 120. of Hirtella adderleyi, H. adenophora, and ame H. ricana. 

227 

o r o~~~~~~~~~~~~~~~~~~~~~ 

lkz v


I 

. -- -.? , 50 -- ___4__50 40 1 60 50 

/ H. angustifolia H. angustissima 

~(-'-I)~I 

HABITAT / 


1 HABITAT 

TRhophyte 

- 

FRUI 

1~FRUIT 

FLOWER I I I 11 

fL I I I IFIAIMIJ 

I I 

JAIS 

I I I 

ER 

ND 

60 

5^^0 ________4_____508 ,50 0___________0_0______ 80 70 

: H. araguariensis H. aramangensis 

FiBA 


F7-- "'^:' T^~ 10 

/ yTerra 

HABITAT 

firme forest 

JIFI MAIMJJI|J IAS 

FRUIT I 

FLOWER . 

_FRUI 1--1/ 

X 

I1I I 

JAISJOINIDI II IJIF MIAM J 

IFRU I J 

LOWER i 

( oo 

i 

60 50 

I I I i I 

80 70 

FIG. 121. Distribution of Hirtella angustifolia, H. angustissima, H. araguariensis, and H. aramangensis. 

o


60 50 40 

.-.-'H. arenosa H. bahiensis 

10 

HABITAT F(AB / ( ( ( 

White sand forest or campina 

J1 

HABITAT 


FRI JIFIMMAI|J IJIAIS oIOIN 

'f---j 

FRUIT I I I I I I-I IF I I I II 

WE'_- 

RII 

IFLOWER - I I 

14/ T-l-I-I ell ,I I I 

604U 

1170fiAIMIJJ AISIOINIDI 

70 _ 50 40 

H. barnebyi, H.H. and barrosoi 

ID 

HABITAT _ ----- >- ---I- F ABITAT 


FLOWERIG 12 isrbt o Hrel a , HFL.OahEnRs 

FIG. 122. Distribution of Hirtella arenosa, H. bahiensis, H. barnebyi, and H. barrosoi. 

2 

~ 

229


O 

70 60 50 40 

ELOWER I/^^ IH. \ 

bicornis 

var. bicornis 


Restinga 

Beaches 


FRUIT 

FLOWER 

H * 

* 

f 

) 

________0 

40 

80 70 60 50 

^V^,^?^J S"^H. bicornis 

ST (/ ^var. 

pubescens 

oO^ 

^l^ ? 

1 Terra firme forest 

FIG. 123. Distribution of Hirtella bicornis. 

0


__?^^"*^^70 80 60 70 60 50 

.. 

........... ......;.'.....~ . 

~ 

JF"IG1Dsb o o( H 70 l 

60 

1 .1 

Gallery forest _ 

Gallery -^~~` forest M.^^ '^?2 

____________ 

- T^^S.""^ : ^ 

'erra~~~~~ 

. ~ .. . 

--T^ ^ - 

-/- _ _ " _j V 

"v *^- -^ 

fim oet ____ / //^Y( / 

"^^- ^ ^ ^ 

Terra firme forest .. 

HG 24 itrbton of HIrelblatan fbucl 

Savanna G. 124. Distribution of Hirtella bullata and H. burcheli. 

0 

231 

^~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~


70 70 00 

Gallery forestGaller 

Seasonally flooded forest / -Savanna 

forest 

margins / 

\ 

) f \ 

( 

70 . _I6 60 

,,60 

I 

, m 

/ H. carbonaria io _H. conduplicata10 

0 0 

\I 0 

JIFIMIAMJIJjAISIOINIDI-_ JI JFMA J /F/.' SN 

FRUIT FRUIT 

FLOWER 15 isl 

tggri FbOWER H 

c 

80 

70_60 50 

FIG. 125. Distribution of Hirtella brachystachya, H. caduca, H. carbonaria, and conduplicata. H. 

FCoastal thicketsI ,_\ 

HABITAT 

Mangallove margins I TGallerra frme forest 

FIG. 125. Distribution of Hirtella brachystachya, H. caduca, H. carbonaria, and H. conduplicata. 

1 

"


70 60 50 4 

^'^X^ -H. ciliata 

0 S r 'i 0 

HABIT ATi |giii * ^ V 

706060 60 

H. confertiflora O C H. cordifolia 

FHABITATG 126. 


Dtbi HiABITAT 

&\\''- Savanna 

fr a H orflia. 

W 

0 JIFIMIAMJ IJiAIS OFN D 

FRUIT I II A | 

FLOFLWERI I I 

0 FRUIT 

ELWE I I 

J F M A JJAS 

I 

I I :I I / 

ND 

I > - 

FIG. 126. Distribution of Hirtella ciliata, H. confertiflora, and H. cordifolia.


s40 50 

'?.i i 

B 

H. corymbosa H. couepiiflora 

60 

60_____________________ 

40 50 iS-0 

_50 

o, o H. cowanii o H. deflexa 

FHABITATG. 12. D b /fi . "ym HABIT AT 

Montane & Clou forestGaller foreste : 

JFMAMJJASNP J FIMIAIJ IJASON \[ 

FRUIT 

1) I 

FRUIT __I___ I 

\ 

FLOWERI li I /(1 lI_ 

( FLOWER _~_~__ _ _ ~_ I ( \ 

60 _ 60 

FIG. 127. Distribution of Hirtella corymbosa, H. couepiiflora, H. cowanii, and H. deflexa. 

FIG. 127. Distribution of Hirtella corymbosa, H. couepiifiora, H. cowantii, and H. defiexa.


i0m~~~~70 

00, .. o H. davisii 

Terra firme forest / / / / 

80 7 0 60 5 0 

"^^H^^'^MB^^"B^^B^^50 0 

70 60 

rf 

1 01 

? H. dorvali H. duckei 

IW: : F/LOWEg g I I I \ 

F 

FIG. 128. Distribution of Hirtella davisii, H. dorvalii, and H. duckei. 

G. 128. Distribution of Hirtella. ds, H. dra, and H. ducke. 

235

236 

80 70 

70 


'- H. elongata 

Gallery forest Y* 3*3i R 

7(, ,2i0 * | 

FLO 

| ER 80 80 7.0 60 60 50 50 0 

80 70 60 50 

01 ~ ~ FG 129 Ditiuino itlaeogtn. ........... 

Secondary forestI 

' 

D st riuto1 o l e t an H 

Seasonally flooded forest \ 

JIFIMAIM Jl IAIS OINIDI O N D \ \ I 

FRUIT *Ig~ I "~i"' H 1 14 ^\ 

r 

Y- 

1 

^ 

\ 

V 

w ^ 

8U 70 60 50 

FIG. 129. Distribution of Hirtella elongata and H. eriandra.


80 70 70 

?t AH. enneandra ; .. " 

H. excels 

FRI Y~3 I v I'-v/ _ I I II *y_ 

y I R 

Montane & Cloud forest Terra firme forest- 

FRUIT ! I1 I 11 ! FRUIT ? --:7-"'- -- 

IFLOWER 8 _/__ 0 

1 1 I 1 

. 

i 

7 -J 

i FLOWER I 

0.70 

_ 60 , 50 50 40 

o H. fasiculata H floribunda 

1 FIG.O~~~~~~~~~~~~! 


. / 

\ ' 

FJI3FIMIAAMIj IJ ÎS ONIDI30IN 

J 

FRUIT 

- 

I I -'1 FRUIT I I' I I I 

FLOWER_ I I i I)' _ II I :I _ 

::1 FgLOWER ! I 

U 5 74U 

FIG. 130. Distribution of Hirtella enneandra, H. excelsa, H. fasciculata, and H. floribunda. 

! 

237


70 60 50 

^^^ l-ô^^ ~~~~~~~~~~ 

1~~~~~~ 

0~~~~~~~~ 

0~~~~~~~~~~~ 

0 

1 

-- 

?e 

0 

O 

io '^^^^ 

3C Savan,a 

f 

"^ -- 

FRUIT ~ ~~ ~ 

0 

/ 

H. glandulosa 

o 0 

. ... . .. .. .. ........:: 

BITAT~~~~~~~~~~~~~~~ 

J F A J JA 

'p ^ 7 - ~ Y ~ - V -T^ - 

:;_____________\ ^--( .... .. 

---- 

1 y Y * 

I I 

!Savanna margins _____ iip~ Y7 ^ - ~ - - ~ ^ 

i 

,FLOWERlell!iillell!l~ele!tel iele!ll elel30 llllel ...._F l RUIT 

70 60 50 40 

7- 

1 

.... 

.......... 

------^ '^Z // ~-- - 

FIG. 131. Distribution of Hirtella glandulosa.


60 50 

10. btH. glabrata,1 H. glandistipula 

HABITAT 

avanna 

econdary forest HABITA 

hite sand forest or campina 

1 JFMAF M A Ml J AIS 0 NIDI J-I 

FRUIT 

FLOWER B 1 /I E 1 

6050 

0 

Terra firme 

FRUIT 

I FLOWER 

forest 

MAIMIJIJ IA SkND I ) 

10 

H. glaziovii 

5I .-^^ ^^ 8^T^0 ^^0 ^ 

0 

H. guainiae 

HABITAT 

BITAT 


JF AMJJ AS ONDJ 

FRUIT I I I I I I I I I 

FLOWER ________lgft 

____I-I 

L_^^^^^^^ ^^40 ^ 

_ 

|pFRU ITI 

FLOWER 

_80 

fM J 

III /I 

M"[ gbbI 

SN 

1-SSl 

-U 

0 

| 

FIG. 132. Distribution of Hirtella glabrata, H. glandistipula, H. glaziovii, and H. guainiae. 

0


70 60 50 

A/ / -/ ',,\H. . gracilipes 

1~_ ____ 90_________ 80 _ _ __ 60__ 0 

1 

F iraH. guatemalensis H. a guyanensis 

L 

WER f:g 

FIG0 I 

. :: 

.. :' 

.......... . 

FIG.90 133. D n of0 l 

60 

FIG. 133. Distribution of Hirtella H. guatemalensis, gracilipes, 

and H. guyanensis.


80 70 60 50 

^^^^:^? 

4 

0 

H. hispidula 10 

FRU IT 

aiFLOWER 

50__.o 80 

iFLOWER 

70 

FLOWER 

60 

50 4 40 

_ 

60 50 

0 

FIG. 134. Distribution of Hirtella hispidula, H. hebeclada, and H hoeh nei 

H.hoHAeIThAT- 

50 40 1 60 50 

0 

241


50 40 60 %/ ir50u 

H. insignis 

H. juruensis 

10~ ~ ~ ~ ~ ~~ ~10 

AT.T.-.. 

.. . . .... 

1 0 

BITAT 

Restinga - J/ I 

FRUIT 

FLOWERI Ii- 1I1111 ____ 

FRUIT 

'FLOWER 

I 

40__ 60 50 

60 50 50 40 

H. kuhimannii H. lancifolia 

0 ABITAT 10 

HABSecondary 

forest 


IJFMIAIMIJJIAISOINID 

FRUIT 

/n\ 

!FRUIT I II 

FLOWER _I _ / 

60 50 

( 

I_ 

I 

T erra firme forest 

JFMA^JJA -/,IS A s 

I! 

)FLOWER . 

_ 50 

^ 

o 

40 

/ 

FIG. 135. Distribution of Hirtella insignis, H. juruensis, H. kuhlmannii, and H. lancifolia. 

0 

0


80 70 80 

{FLOWER-- -H. = latifolia 

H. lemsii 

10 -M 010 0 

70 

0 

V 

0/D~~ 0 


H. leonotis H. liesneri 

Terra firme forest Slope forest / 

FRI 

J3FM. AJDJAoND I IJFM JAI SH N sn . 

II 

|FLOWER _I- 1Fl 

70^ 

I I I I I LOWE II 

7 - I 

O I I I I FRUIT I-I I I [:L - 

FIG. 136. Distribution of Hirtella latifolia, H. lemsii, H. leonotis, and H. liesneri. 

0 

243


70 70 

- H. lightioides H. longifolia 

10, 7 6-- 10 

Terra firme forestfHtligod,H Amazonian caatinga t,a . 

FRUIT I FRUIT 

70 70__________ 

____7_______60 _60_______________ 50 

HABIT AT L ' 

H. longipedicellata 0 H. macrosepala 0 

Savanna margins F _ITAT 

Slope forest Terra firme forest 

J(FIMMAIMIJJAS OND J JASON D ..... 

FRUIT 

FLOWER 

L:::I: 

I_^ I ^ .w 

" 

\ 

TR-U RUT- --------R 10I I 

FLOWER 

_60 70 60 

50 

FIG. 137. Distribution of Hirtella lightioides, H. longifolia, H. longipedicellata, and H. macrosepala. 

0 

3


80 70 60 50 

_0 _Secondary , 

forest : 

........,, 

H. magnifoliaS:7- 

. 

H. macrophylla H. 

880 70 60 50 

. 

1. H. magnifolia a nd H. margae 

Seasona firme foodrest Terra fe fos ' 0 

I IJIFFIMIAIMIJI JIAISO\ NII 

/ 

~ 

rFLOWERFI I I = = =II _\ = il !.'?" 

F 

b o 

G80 70 

HABITAT \ 

F0A 


FRUIT 

FLOWER 

____________ 

_ _ 

^\ ) 

-- 

FRUIT *_ I 

FLOWER_ 

80 70 6U 56 

FIG. 138. Distribution of Hirtella macroPhylla, H. magnifolia, and H. margae. 

. 

245


?10F 

50 40 60 50 

- H-. martiana > H. mucronata 

0 

.. 

........ 

HABITAT- - ..... 

Swamps 

Gallery forest . 

I HABITAT 


10 

|J|FIM|A|MIJJ IJIAISIA |A|S|O|N|D| NID I 

FRUIT 1 1 1 I I 

FLOWERI 0g;ggg I lO 1 1 

I I 

J F_ AIQJ 

FRUIT _ 

LFLOWER 

F 

_ 

----- 

40 

g 

50 

J 

10 

80 70 60 50 

.0 

H. mutisii H. myrmecophila 

10 

'erra firme forest __[erra firme forest 1 j 

0 

"~/U 

b'HABIT_T U 

....o 

FIG. 139. Distribution of Hirtella martiana, H. mucronata, H. mutis, and 

H. myrmecophla. 

FIG. 139. Distribution of Hirtella martiana, H. mucronata, H. mutisii, and H. myrmecophila.


(,. 

60 50 60 


H. obidensis H. orbicularis 

% 

o , 


HG. 10 itbtnbor s H. paniculata 

1 03 

FRU II F L O,... 

80 7060 50 

FIG. 140. Distribution of Hirtella obidensis, H. orbicularis, and H. paniculata. 

FIG. 140. Distribution of Hirtella obidensis, H. orbicularis, and H. paniculata. 

247


0. 

50 50 40 

,) 

: 

H. paraensis 

FHABITAT / /I HABTij 

50 50 

-- 

0 

H. parviunguis 

402 

90 80 70 60 

G 1 H. pauciflora C 

' H. pendula 

0 (. 

0 0 

1j^^V^-\J^ Id. HABII-AT t-II0HABITAT ^ -^'0 0 

Terra firme forest . Terra firme forest 

ZZZZ J AIJ I J I A NID0 0 "^ 

/


0 

80 70 60 50 

Terrafirme for t,,. 

F 2 i7 

I IJIFIG. JIJ IoI S li 

6. 

.. 

. 5 

FIG 142. Distribution of Hirtella ilosissima and H. physophora. 

FG14.Dsrbto n of Hit piloism el n Hhspoa


70 60 60 50 

H. pimichina H. piresi 

l .. 

HABITAT? // /J -HABITA.. /r^ 


/ _ _,__.~_T .......f^ 


1FRUI 

JFIF AM AJ T J AS N ND) 

1FMAJ F= 

FRUI T 

AI 

I I II I I 

1 ~i10 ^s:^. 

0~~~O7 

--- 

FIGOW 143. . Distributn of H . a p 

pii H. p til , ad H. r mii 

'C 

60 50 70 

FIG. 143. Distribution of Hirtella pimichina, H. piresii, H. punctillata, and H. radamii. 

0

100 90 

l2( C^^^^^--^^ .~ C: 

10 

10: BI TATI 

80, 

70 60 

o , H. r 

....' ....... 

White sand forest or campina( / 

Terra JSav '7nna firme forest'-' " =l ) 

/---i- -J 

\ 

Savanna margins~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~^ 

FRUIT -I i ***__* **, ,\ . ; 

FLOWER 

100 90 80 70 60 

FIG. 144. Distribution of Hirtella racemosa var. hexandra.


10 10 

^^^P^^^ ?^ 

10 '[ 

80 70 60 50 

*


80 70 50 

/ H. racemosa H. racemosa 

var. glandipedicellata var. hispida 

80 - 10 .. .... 

l , (e 

20 HABITAT- 

- 

20 

FRUIT II11I 

FLOWER 

II I I 

~ 

I 

F 

80 70 ...... 50 

80 70 80 70 

? ^ 

. 

.0 

rasa 

' 0 

Ho e H -H. rasa revillae 

0 

| | | | /: H. revillae 

20 HABITAT 

Terra SlopTerra firme fiorest forest 

.. 

( ,\ ' 

BITAT 

White White sand forest or campina campi'"' 

JIFIMAMJ IJIAS - JFMAJJASND" 

FIG. 146. Distribution of Hirtella racemosa var. glandipedicellata, H. racemosa var. hispida, H. rasa, and 

H. revillae. 

' 

/U 

sIa'!j;l~ 7h'te 

; o0 

IJ;ANi.,' 

FRUIT UIT I 

FLOWER! i I IFLOWRI I I I I, 

70 80


70 60 70 60 

H. rodriguesii 

''I Wl" I I0I 

HAB17AT-^4' /Bi\ 'TTHABITAT ~ 

> 

f 

Terra firme forest \Montane & Cloud forest - 

JIFIMIAIMIJIJIAISND\ J IIAIMJ IAISoNI 

H. rugosa 

50 40 I 80 70 

G 1... H. santosii r a H. a scaberula 

^0 0 

HABITAT^k \ --:????./ 

0.ABITAT 

Terra firme forest Amazonian caatinga 

J IFIMIAI JI As A oN NJIFIMIA1J I I 

I 

IJIAISND 

FRUIT IFRUIT I I I I I_ I I I 

FLOWER __I I I I I I I I ^- FLOWER I_______I I I 

4U80 Tu70 

FIG. 147. Distribution of Hirtella rodriguesii, H. rugosa, H. santosii, and H. scaberula. 

0 

I" 

.


10 

g H. scabra H. schultesii 

10I 120^ .. I I I . y 

/ 

HABITAT 

v 

^/ ^ 

Savanna IJ F M J JWSTerra \ 

firme foresi t 

HABITHAT 

FRUIT I J i 

0 

10C0 

AS 

FLOWER FOlO_WE_ll 

I 

iT J( Is I 

FIG. 148. Distribution of Hirtella scabra,H. H. silicea, schultesi, and H. sprucei 

vTerra time forest .Terra firme forest 

_~60 _ - bU _ _ 4u 

G. 148. Distribution oHirtlla scabra, H. schultesii, silicea, H. and H. sprucei. 

FIG. 148. Distribution of Hirtella scabra, H. schultesii, H. silicea, and H. sprucei. 

A 

.e_ 

70 

M^ 

'/2 

.


80 70 70 

r H. standleyi H. subglanduligera 

~~>FRUIT . LFRUIT- I I- I I _ 

01r 1 -?c'HxIAS0 

__ 8_ 7 _ 

750_= 

71 0 0 -606 

0 

60 60 

HABIT 11\IAT 1 

/I I \T-A I L I I I I 

NIFLOlWER ___g__ I __I |I 

1FLWE T 

IE I S I 

FIG. 149. Distribution of Hirtella standleyi, H. subglanduligera, H. subscandens, and H. suffulta. 

50


60 50 60 50 

H. tentaculata H. tocantina 



JIFIMIAIMI JIJ IAISIOIN eJ 

tubifBITAT H. 


FMAIMIJ IJIAIS I 

X 

FRUIT _ _ _ _ 6___ 

FLOWER _ _:_I '*ggS_rl 

_ 

I @1 1I 

FRUIT 

I =IFL I 

// 

J 

60 50 50 

60 50 80 

r'oLH. tenuifolia H. tubiflora 

Secondary forest .----^ 1 HABITAT^ -_ /== s 

Terra firme forest / . . Terra firme forest >1 

IFRUIT I|I I| I IIJ||IN1 1 I l Y JM X 2/ 1 ^^ 1 

FIG. 150. Distribution of Hirtella tentaculata, H. tocantina, H. tenuifolia, and H. tubiflora.

0 

100 

/ "I ^ ~ 

90_ 80 70 


90 50 40 

-H. triandra H triandra 

subsp. media20 1 subsp. punctulata 

.. ............. 

HABITAT 

Terra firme forestSlope 

HABITAT 


lope forest 

forest - 

JIFIMIAIMI JJ IAIS OIND 

FRUIT 

- 

_ -- - 

L -FRUIT 

FLOWER FL~OWERI __ 1 I I@ T1 1l I 1 ___ I I 1 ___ I @ 

90 

I 

IJ IFIMAIMIJIJ IAIOI N ID 

I I I I I I I I 

rILOWERI FLOWERi:::I I I r Bs*g 1 L 

40 

__ 

U'-60 70 60 

' .. - JH. ulei H. vesiculosa 

- 0 

-o- -- 0 

HABITAT .. .. 

Savanna ..... 

White sand forest or campina /0 

Beaches 

FRUIT= 

FLOWER : 

'! 

:FLOWER 

FR! 

70 

1 I I I 

. I II 

I I I 

I I / 

FIG. 152. Distribution of Hirtella triandra subsp. media, H. triandra subsp. punctulata, H. ulei, and H. 

vesiculosa. 

, 

259


ol F RUI 

o 1^\-_ ^^ 

90 8070 

-JMaranthes 

0 ~~ ~ I-rr 

FLOWER I _ _ _ 

o 

_ 

panamensis 0 

Slope forest 

Amazonian caatinga 


__I-80____-70_ _ 60 

1 

50 

FIG. 153. Distribution of Maranthes panamensis and the genus Acioa. 

_. 

_


60 0 o , I50 _7 60_I 

-


HABITA 

'Ir\ 1 )'II i JI)I MJIJ 

1' II( ) EL1 

\~~~~~~~~~~~~~~~~7 

FI.15 

sectin; E petl; Lcnadosni(Aeeo& F,ovar andstyl. Inst: dstriutionof ay159.A L ait dodonii 

,oenn lwe u;C,foe; ,foe 

FIG. 155. Licania dodsonii (Acevedo & Daly 1659). A, habit; B, opening flower bud; C, flower; D, flower 

section; E, petal; F, ovary and style. Inset: distribution of L. dodsonii.

Index of Scientific Names 263 

apacharama amarillo (Peru) 95 

bokobokotokon 89 

carip6 torrado 93 

castanha de cotia 97 

coquito 95 

cuero de sapo 59 

duship 30 

gaulette (Fr. Guiana) 95 

guaiti-mirim 59 

INDEX OF LOCAL NAMES 

lobo apacharama 92 

macucurana 95 

maeneyowaie (Auka Indian) 20 

milho torrado 70 

milho-torrado-amarelo 55 

mulo (El Salvador) 27 

oiti 46, 55, 58 

oiti-mirim 57, 77 

INDEX OF SCIENTIFIC NAMES 

parinari 58 

parinari (Peru) 51 

parinari blanco 58, 65 

pasista 96 

rode kwepi 49 

uchirana (Brazil) 65 

yakuku 58 

yukuku (Peru) 27 

New names and combinations are in bold face and synonyms are in italic. Page numbers in bold 

face indicate primary page references. Page numbers with an asterisk (*) indicate pages with illus- 

trations or maps. 

Acia amara 64 

Acioa 2, 3, 96, 97, 260* 

amara 64 

edulis 96, 106, 209* 

guianensis 96, 106, 261* 

schultesii 96, 106, 261* 

somnolens 106, 261* 

Arecaceae 53 

Barcella odora 53 

Caryocaraceae 2 

Chrysobalanaceae 2, 3, 4, 91 

Chrysobalaneae 3 

Chrysobalanus 3, 4 

cuspidatus 6, 99, 128* 

icaco 3, 4, 6, 99, 127* 

interior 4 

venezuelanus 4, 5*, 6, 99, 128* 

Clusia 70 

Colpothrinax cookii 70 

Couepia 59, 64, 65, 72, 97, 203* 

amaralae 72, 73*, 104, 204* 

belemii 71, 103, 204* 

bernardii 65, 66, 68, 103, 204* 

bondarii 70, 74, 104, 204* 

bracteosa 103, 205* 

canescens 68, 72, 103, 205* 

canomensis 103, 205* 

carautae 70, 71*, 103, 206* 

caryophylloides 72, 103, 206* 

subsp. caryophylloides 72, 103 

subsp. glabra 72, 103 

cataractae 103, 206* 

chrysocalyx 104, 207* 

cidiana 74, 104, 206* 

coarctata 77, 78*, 104, 207* 

cognata 68, 104, 208* 

var. cognata 68, 104, 208* 

var. major 104, 208* 

var. membranacea 104, 208* 

comosa 76, 104, 207* 

divaricata 65 

divaricata var. strictiuscula 65 

dolichopoda 75, 76, 104, 208* 

edulis 2, 96 

elata 104, 209* 

eriantha 104, 209* 

excelsa 103, 209* 

exflexa 103, 210* 

foveolata 72, 103, 210* 

froesii 104, 210* 

glabra 74, 75, 104, 210* 

glandulosa 65, 103 

grandiflora 104, 211* 

guianensis 64 

subsp. divaricata 65, 103, 211* 

subsp. glandulosa 64, 103, 212* 

subsp. guianensis 64, 103, 212* 

habrantha 70, 103, 213* 

impressa 77, 104, 213* 

subsp. cabraliae 77, 104, 213* 

subsp. impressa 104, 213* 

insignis 70, 74, 104, 213* 

krukovii 72, 103, 214* 

latifolia 104, 214* 

leptostachya 64, 103 

longipendula 76, 104, 214* 

longipetiolata 77, 79, 104, 214* 

macrophylla 72, 103, 215* 

magnoliifolia 70, 103, 215* 

maguirei 103, 215* 

marleneae 75, 104, 215* 

martinii 74, 104, 216* 

meridionalis 77, 79, 104, 216* 

monteclarensis 66, 67*, 68, 103 

multiflora 104, 216* 

myrtifolia 65 

nutans 76, 104, 216* 

obovata 65, 66, 104, 217* 

ovalifolia 104, 218* 

paraensis 219*

264 

subsp. cerradoanda 103, 219* 

subsp. glaucescens 103, 219 

subsp. paraensis 103, 219* 

parillo 39, 103, 217* 

parvifolia 104, 218* 

perambucensis 79, 104, 218* 

platycalyx 76, 104, 218* 

polyandra 76, 104, 220* 

racemosa 104, 220* 

recurva 74, 104, 221* 

reflexa 65, 70, 104, 221* 

robusta 104, 221* 

rufa 104, 221* 

sandwithii 65, 66, 68, 103, 222* 

schottii 79, 104, 222* 

scottmorii 68, 69*, 103, 222* 

spicata 70, 103, 222* 

steyermarkii 72, 103, 223* 

stipularis 104, 223* 

subcordata 103, 223* 

surinamensis 64 

trapezioana 75, 104, 225* 

uiti 103, 224* 

ulei 104, 224* 

venosa 104, 225* 

versicolor 64 

williamsii 75, 104, 225* 

Dactyladenia 2, 3 

Dichapetalaceae 2 

Exellodendron 59, 200* 

barbatum 59, 103, 201* 

cordatum 59, 103, 201* 

coriaceum 59, 103, 202* 

gardneri 59, 103, 202* 

gracile 59, 103, 202* 

Hirtella 79, 91, 92, 226* 

adderleyi 88, 91, 105, 227* 

adenophora 106, 227* 

americana 91, 105, 227* 

angustifolia 95, 106, 228* 

angustissima 105, 228* 

araguariensis 88, 89, 105, 228* 

aramangensis 105, 228* 

arenosa 93, 106, 229* 

bahiensis 92, 105, 229* 

bamebyi 88, 105, 229* 

barrosoi 105, 229* 

bicoris 105, 230* 

var. bicomis 105, 230* 

var. pubescens 105, 230* 

brachystachya 106, 232* 

bullata 105, 231* 

burchellii 106, 231* 

caduca 106, 232* 

carbonaria 105, 232* 

ciliata 105, 223* 

cliffortiana 106 

condifolia 105, 233* 

conduplicata 93, 106, 232* 

confertiflora 87, 88, 105, 233* 

cordifolia 105, 233* 

corymbosa 91, 105, 234* 

couepiiflora 95, 106, 234* 

cowanii 105, 234* 

davisii 91, 105, 235* 

deflexa 105, 234* 

dorvalii 87, 104, 235* 

duckei 87, 104, 235* 

elongata 88, 92, 105, 236* 

enneandra 106, 237* 

eriandra 88, 92, 105, 236* 

excelsa 95, 106, 237* 

fasciculata 106, 237* 

floribunda 106, 237* 

glabrata 88, 104, 239* 

glandistipula 106, 239* 

glandulosa 105, 238* 

glaziovii 95, 106, 239* 

gracilipes 106, 240* 

guainiae 104, 239* 

guatemalensis 105, 240* 

guyanensis 105, 240* 

hebeclada 106, 241* 

hispidula 106, 241* 

hoehnei 105, 241* 

insignis 89, 105, 242* 

juruensis 106, 242* 

kuhlmannii 94, 106, 242* 

lancifolia 106, 242* 


lemsii 94, 106, 243* 

leonotis 92, 105, 243* 

liesneri 89, 90*, 105, 243* 

lightioides 105, 244* 

longifolia 106, 244* 

longipedicellata 106, 244* 

macrophylla 105, 245* 

macrosepala 104, 244* 

magnifolia 92, 105, 245* 

margae 88, 105, 245* 

martiana 95, 106, 246* 

megacarpa 106 

mucronata 94, 106, 246* 

mutisii 105, 246* 

myrmecophila 104, 246* 

obidensis 105, 247* 

orbicularis 105, 247* 

paniculata 105, 247* 

paraensis 106, 248* 

parviunguis 94, 95, 106, 248* 

pauciflora 95, 106, 248* 

pendula 105, 248* 

physophora 87, 104, 249* 

pilosissima 105, 249* 

pimichina 106, 250* 

piresii 105, 250* 

pohlii 106 

punctillata 91, 105, 250* 

racemosa 93, 106 

var. glandipedicellata 106, 253* 

var. hexandra 94, 106, 251* 

var. hispida 94, 106, 253* 

var. racemosa 94, 106, 252* 

radamii 93, 105, 250* 

rasa 92, 105, 253* 

revillae 87, 104, 253* 

rodriguesii 105, 254* 

rugosa 106, 254* 

santosii 91, 105, 254* 

scaberula 106, 254* 

Flora Neotropica


scabra 105, 255* 

schultesii 106, 255* 

sect. Myrmecophila 79, 87 

silicea 106, 255* 

sprucei 106, 255* 

standleyi 106, 256* 

subglanduligera 105, 256* 

subscandens 106, 256* 

suffulta 105, 256* 

tentaculata 105, 257* 

tenuifolia 93, 105, 257* 

tocantina 105, 257* 

triandra 105 

subsp. media 105, 259* 

subsp. punctulata 105, 259* 

subsp. triandra 105, 258* 

tubiflora 95, 106, 257* 

ulei 104, 259* 

vesiculosa 104, 259* 

zanzibarica 106 

Licania 3, 6, 39, 72 

subg. Licania 36, 39, 44, 47, 48 

sect. Cymosa 44, 45, 46 

sect. Hirsuta 36, 39 

sect. Hymenopus 39 

sect. Licania 48, 53, 54 

sect. Pulverulenta 47, 48 

subg. Moquilea 19, 24, 28, 29, 30, 32, 34, 35, 36 

sect. Leptobalanus 32, 34 

sect. Microdesmia 30, 35, 36 

sect. Moquilea 19, 20, 27, 28, 29, 30, 32, 34, 35 

subg. Parinariopsis 6, 11 

affinis 47, 59, 101, 129* 

alba 101, 129* 

albiflora 35, 100, 130* 

amapaensis 56, 102, 130* 

angustata 28, 99, 130* 

anneae 27, 99, 130* 

apetala 3, 32, 34, 99 

var. aperta 32, 99, 131* 

var. apetala 99, 131* 

apiculata 102, 132* 

aracaensis 51, 102, 132* 

arachnoidea 39, 46, 100, 132* 

araneosa 36, 100, 132* 

arborea 35, 100, 133* 

arianeae 44, 45*, 101, 134* 

bahiensis 102, 134* 

belemii 55, 102, 134* 

bellingtonii 48, 101, 134* 

blackii 55, 102, 135* 

boliviensis 99, 135* 

boyanii 101, 135* 

bracteata 51, 102, 136* 

britteniana 99, 136* 

buxifolia 47, 101, 136* 

cabrerae 20, 24, 26, 99, 137* 

caldasiana 102, 137* 

calvescens 35, 100, 137* 

canescens 101, 138* 

caudata 39, 40, 100, 138* 

cecidiophora 29, 30, 99, 137* 

chiriquiensis 28, 29, 99, 140* 

chocoensis 100, 140* 

compacta 102, 140* 

cordata 101, 140* 

coriacea 101, 139* 

costaricensis 100, 139* 

couepiifolia 48, 101, 139* 

crassivenia 49, 101, 139* 

cruegeriana 55, 102, 141* 

cuatrecasasii 33, 100, 141* 

cuprea 45, 101, 141* 

cuspidata 99, 141* 

cyathodes 101, 142* 

cymosa 46, 53, 101, 142* 

davillifolia 101, 142* 

dealbata 46, 101, 142* 

densiflora 101, 143* 

discolor 102, 143* 

divaricata 100, 143* 

dodsonii 27, 99, 262* 

durifolia 20, 22, 26, 55, 99, 143* 

egleri 99, 144* 

elliptica 101, 144* 

emarginata 100, 145* 

fanshawei 44, 101, 145* 

fasciculata 24, 26, 99, 145* 

filomenoi 20, 21*, 22, 99, 145* 

foldatsii 101, 146* 

foveolata 100, 146* 

fritschii 99, 146* 

fuchsii 100, 146* 

furfuracea 47, 101, 147* 

fusicarpa 102 

gardneri 99, 147* 

gentryi 20, 24, 99, 147* 

glabriflora 40, 100, 148* 

glabrifolia (sphalm.) 40 

glauca 101, 148* 

glazioviana 100, 148* 

gonzalezii 28, 99, 149* 

gracilipes 101, 149* 

grandibracteata 20, 22, 23*, 99, 149* 

granvillei 32, 99, 149* 

guatemalensis 29, 99, 150* 

guianensis 39, 99, 150* 

harlingii 55, 102, 150* 

hebantha 49, 101, 153* 

heteromorpha 40, 41, 42 

var. glabra 100, 152* 

var. heteromorpha 100, 151* 

var. perplexans 100, 152* 

var. revoluta 40, 100 

var. subcordata 100, 152* 

hirsuta 36, 100, 153* 

hispida 37, 38*, 100, 153* 

hitchcockii 101, 153* 

hoehnei 102, 154* 

humilis 100, 154* 

hypoleuca 46, 101, 155* 

var. foveolata 101, 155* 

var. hypoleuca 101, 155* 

impressa 46, 55, 101, 156* 

incana 54, 102, 156* 

indurata 102, 156* 

inpae 156* 

intrapetiolaris 41, 100, 157* 

irwinii 101, 157* 

jefensis 32, 99, 157*


jimenezii 49, 101, 157* 

joseramosii 34, 100, 158* 

kallunkiae 28, 29, 99, 158* 

klugii 26, 99, 158* 

krukovii 100, 158* 

kunthiana 101, 159* 

laevigata 41*, 42, 100, 169* 

lamentanda 51, 52*, 102, 160* 

lanceolata 53, 102, 160* 

lasseri 100, 161* 

lata 99, 160* 


latistipula 40, 100, 161* 

laxiflora 101, 162* 

leptostachya 53, 54, 102, 163* 

leucosepala 27, 99, 162* 

licaniiflora 100, 163* 

littoralis 101, 164* 

var. cuneata 101 

var. littoralis 101 

longipedicellata 27, 35, 99, 164* 

longipetala 30, 99, 164* 

longistyla 35, 100, 165* 

macrocarpa 19, 20, 22, 26, 99, 166* 

macrophylla 100, 165* 

maguirei 99, 166* 

majuscula 101, 166* 

maranhensis 99*, 167* 

maritima 19, 20, 26, 99, 166* 

marleneae 44, 101, 167* 

maxima 102, 167* 

membranacea 101, 168* 

mexicana 34, 100, 167* 

michauxii 3, 99, 168* 

micrantha 51, 102, 170* 

microphylla 102, 168* 

miltonii 40, 100, 171* 

minuscula 100, 170* 

minutiflora 27, 28, 29, 99, 171* 

mollis 102, 172* 

montana 20, 26, 99, 172* 

morii 33, 99, 172* 

naviculistipula 101, 172* 

nelsonii 53, 54*, 102, 169* 

niloi 101, 173* 

nitida 102, 173* 

oblongifolia 100, 173* 

obtusifolia 102 

occultans 42, 100, 169* 

octandra 35 

subsp. grandifolia 35, 100, 175* 

subsp. octandra 35, 100, 174* 

subsp. pallida 35, 100, 175* 

operculipetala 100, 175 

orbicularis 101, 176* 

ovalifolia 49, 102, 176* 

pakaraimensis 39, 100, 177* 

palawanensis 102 

pallida 46, 101, 176* 

paraensis 54, 102, 177* 

parviflora 102, 177* 


parvifructa 101, 178* 

persaudii 100, 178* 

piresii 46, 101, 180* 

platypus 99, 179* 

polita 101, 179* 

pruinosa 102, 180* 

pyrifolia 99, 180* 

reticulata 26, 42, 100, 182* 

retifolia 99, 180* 

riedelii 46, 102, 182* 

rigida 100, 181* 

robusta 53, 102, 182* 

rodriguesii 102, 183* 

roraimensis 102, 183* 

rufescens 101, 183* 

salicifolia 36, 100, 183* 

salzmannii 99, 184* 

sandwithii 101, 184* 

santosii 46, 101, 184* 

savannarum 102, 184* 

sclerophylla 100, 185* 

silvae 44, 101, 185* 

silvatica 100, 186* 

sparsipilis 33, 34, 100, 186* 

spicata 102, 186* 

splendens 102 

sprucei 100, 186* 

stewardii 50, 102, 187* 

steyermarkii 49, 101, 187* 

stricta 102, 187* 

subarachnophylla 36, 100, 187* 

subrotundata 101, 188* 

tachirensis 30, 31*, 99, 188* 

tambopatensis 36, 37*, 100, 169* 

teixeirae 47, 48*, 101, 188* 

tepuiensis 102, 188* 

teratensis 101, 189* 

tocantina 50, 102, 189* 

tomentosa 99, 189* 

triandra 50, 102, 190* 

trigonioides 101, 189* 

turbinata 99, 191* 

unguiculata 30, 99, 190* 

urceolaris 47, 101, 191* 

vaupesiana 55, 102, 191* 

velata 20, 100, 192* 

veneralensis 20, 26, 55, 102, 192* 

wurdackii 99, 192* 

Maranthes 59 

corymbosa 59 

panamensis 59, 103, 260* 

Moquilea 64 

couepia 64 

glandulosa 65 

Myrtales 3 

Neocarya 97 

macrophylla 97, 106 

Parinari 56, 57, 58, 72, 97, 193* 

alvimii 57, 102, 194* 

brasiliensis 57, 102, 194* 

campestris 57, 102, 194* 

canescens 68 

cardiophylla 103, 195* 

chocoensis 58, 103, 195*


excelsa 57, 102, 196* 

klugii 57, 102, 195* 

krukovii 65 

littoralis 57, 58, 103, 195* 

maguirei 58, 103, 197* 

montana 102, 197* 

obtusifolia 103, 197* 

occidentalis 102, 198* 

pachyphylla 102, 198* 

parilis 58, 103, 198* 


rodolphii 57, 102, 199* 

romeroi 59, 103, 199* 

sprucei 102, 199* 

Rosaceae 3 

Theales 3

GHILLEAN T. PRANCE 

Ghillean T. Prance was born in Suffolk, England. He was educated at Malvern 

College in Worcestershire and holds a B.A., M.A., and D.Phil. from Oxford 

University. 

From 1963 to 1988 he was a staff member of the New York Botanical Garden 

and during that time conducted a series of botanical expeditions in Brazilian 

Amazonia. He has published 8 books and over 180 scientific and general articles 

on plant systematics, plant ecology, ethnobotany and conservation. He is author 

of four previous monographs in Flora Neotropica: Chrysobalanaceae (1972), Dichapetalaceae 

and Rhabdodendraceae (1972) and Caryocaraceae (1973). He is coauthor 

of Lecythidaceae with Scott A. Mori (Part I, 1979 and Part II, in press). 

He holds a Fil.Dr. (honoris causa) from Goteborg University and is a Fellow of 

the Linnean Society of London and the Explorers Club, a corresponding member 

of the Brazilian Academy of Sciences and a foreign member of the Royal Danish 

Academy of Sciences and Letters. He was Executive Director of Flora Neotropica 

from 1975 to 1988 and Director of the New York Botanical Garden Institute of 

Economic Botany from 1981 to 1988. He is currently Director of the Royal Botanic 

Gardens, Kew, Great Britain.

FLORA NEOTROPICA 

Flora Neotropica is designed to present in monographic form taxonomic 

accounts of all plants growing spontaneously within the Western Hemisphere 

tropics. Geographic, ecologic, cytologic, anatomic, morphologic, chemical, and 

economic data will provide complementary information for each contribution. 

Bibliography, citation of specimens, and indexes are intended to facilitate con- 

sultation. 

Monographs of Flora Neotropica will be issued separately without regard to 

taxonomic sequence. Each author will be wholly responsible for his or her own 

contribution, being restricted only by the general style and form of the work. 

Those interested in preparing a monograph for Flora Neotropica must consult 

the Staff Committee: 

Cryptogams: 

Dr. S. Rob Gradstein 

Institute of Systematic Botany 

Heidelberglaan 2 

P.O. Box 80.102 

3508 TC Utrecht 

The Netherlands 

Phanerogams: 

Dr. Enrique Forero 

Herbarium 

Missouri Botanical Garden 

P.O. Box 299 

St. Louis, Missouri 63166 

For further information write to Scott A. Mori, Executive Director, Orga- 

nization for Flora Neotropica, The New York Botanical Garden, Bronx, N.Y. 

10458, U.S.A. 


Monograph 

1 Cowan, B. Swartzia (Leguminosae, Caesalpinioideae, $15.00 

Swartzieae) (Reprinted 1987) 

2 Cuatrecasas, J. Brunelliaceae (Reprinted 1984) $15.00 

2 sup. Cuatrecasas, J. Brunelliaceae (Supplement) $21.00 

3-5 Singer, R. Omphalinae-Phaeocollybia- $13.00 

Strobilomycetaceae 

6 Lowy, B. Tremellales (Reprinted 1987) $13.00 

6 sup. Lowy, B. Tremellales (Supplement) $ 4.50 

7 Berg, C. C. Moraceae, Olmedieae & Brosimeae $18.00 

(Reprinted 1985) 

8 Maas, P. J. M. Zingiberaceae: Costoideae $13.95 

9 Prance, G. T. Chrysobalanaceae $27.95 

9S Prance, G. T. Chrysobalanaceae $ 

10-11 Prance, G. T. Dichapetalaceae & Rhabdodendraceae $13.95 

12 Prance, G. T. & Caryocaraceae $ 9.95 

Silva, M. F. 

13 Rogers, D. J. & Manihot, Manihotoides (Euphorbiaceae) $23.95 

Appun, S. G. 

14(1) Smith, L. B. & Pitcairioideae (Bromeliaceae) $45.00 

Downs, R. J. 

14(2) Smith, L. B. & Tillandsioideae (Bromeliaceae) $49.50 

Downs, R. J. 

14(3) Smith, L. B. & Bromelioideae (Bromeliaceae) $65.00 

Downs, R. J. (Reprinted 1983) 

15 Morley, T. Memecyleae (Melastomataceae) $22.00 

16 Farr, M. L. Myxomycetes $22.50 

17 Singer, R. Marasmieae (Tricholomataceae) $25.00 

18 Maas, P. J. M. Renealmia (Zingiberaceae- $21.00 

Zingiberoideae), Costoideae (Additions) 

19 Lleras, E. Trigoniaceae $ 7.25 

20 Johnston, M. C. & Rhamnus $10.00 

Johnston, L. A.

Continued from Cover 3 

21 Prance, G. T. & Lecythidaceae-Part I (Asteranthos, $28.00 

Mori, S. A. Gustavia, Grias, Allantoma & 

Cariniana) 

22 Sleumer, H. 0. Flacourtiaceae $47.50 

23 Hansen, B. Balanophoraceae $10.50 

24 Buritica, P. & Pucciniosireae (Uredinales, Pucciniaceae) $ 7.75 

Hennen, J. F. 

25 Gentry, A. H. Bignoniaceae-Part I (Crescentieae & $15.75 

Tourretieae) 

26 Mesa, A. Nolanaceae $12.00 

27 Poppendieck, H-H. Cochlospermaceae $ 6.50 

28 Pennington, T. D. Meliaceae $65.00 

29 Landrum, L. R. Myrceugenia (Myrtaceae) $20.00 

30 Gates, B. Banisteriopsis, Diplopterys (Malpighiaceae) $35.00 

31 Kubitzki, K. & Lauraceae I (Aniba and Aiouea) $22.50 

Renner, S. 

32 Singer, R. Hydropus (Basidiomycetes) $25.00 

33 Kaastra, R. C. Pilocarpineae (Rutaceae) $31.50 

34 Daniel, T. Carlowrightia (Acanthaceae) $20.50 

35 Luteyn, J. L. Cavendishia (Ericaceae) $41.00 

36 Forero, E. Connaraceae $37.00 

37 Paden, J. W. Sarcosomataceae (Pezizales $ 5.50 

Sarcoscyphineae) 

38 Sleumer, H. 0. Olacaceae $26.00 

39 Rogers, G. K. Henriquezieae (Rubiaceae) $25.00 

40-42 Maas, P. J. M. et al. Saprophytes, pro parte $49.50 

43, 44 Hopkins, H. C. F. Parkia (Leguminosae: Mimosoideae) $44.75 

(one vol.) da Silva, M. F. Dimorphandra (Caesalpiniaceae) 

45 Landrum, L. R. Campomanesia, etc. (Myrtaceae) $35.50 

46 Hekking, W. H. A. Violaceae. Part I-Rinorea and $40.50 

Rinoreocarpus 

47 Molau, U. Scrophulariaceae-Part I. Calceolarieae $59.00 

48 Todzia, C. A. Chloranthaceae: Hedyosmum $29.00 

49 Simpson, B. B. Krameriaceae $ 

Postage and Handling: 

U.S. ORDERS: $2.00 + 3% of sub-total. 

NON-U.S. ORDERS: $3.00 + 4% of sub-total. Payment in U.S. currency drawn on a 

U.S. bank. Thank you. 

Order from: Scientific Publications Department, The New York Botanical Garden, Bronx, 

NY 10458 U.S.A.

flora neotropica - CNCFlora

You also want an ePaper? Increase the reach of your titles

Delete template?

Save as template?