Cecropiaceae: Coussapoa and Pourouma, with an ... - CNCFlora

Organization for Flora Neotropica 

Cecropiaceae: Coussapoa and Pourouma, with an Introduction to the Family 

Author(s): C. C. Berg, R. W. A. P. Akkermans, E. C. H. van Heusden 

Reviewed work(s): 

Source: Flora Neotropica, Vol. 51, Cecropiaceae: Coussapoa and Pourouma, with an Introduction 

to the Family (Apr. 11, 1990), pp. 1-208 

Published by: New York Botanical Garden Press on behalf of Organization for Flora Neotropica 

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FLORA NEOTROPIC 

MONOGRAPH 51 

CECROPIACEAE: COUSSAPOA AND 

POUROUMA, WITH AN INTRODUCTION 

TO THE FAMILY 

by 

C. C. Berg, R. W. A. P. Akkermans and E. C. H. van Heusden 

. i ., """ 

///A 

"^C^ - - -- - ------------------- 

TIOPIC 

OF CANCER 

NEOTROPICsAh 

Published for 

Organization for Flora Neotropica 

by 

The New York Botanical Garden 

New York 

Issued 11 April 1990

Copyright ? 1990 


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-dcl9 85-647083 

AACR 2 MARC-S 

Library of Congress [8508] 

ISBN 0-89327-352-x

CECROPIACEAE: COUSSAPOA AND POUROUMA, 

WITH AN INTRODUCTION TO THE FAMILY 

C. C. BERG,1 R. W. A. P. AKKERMANS,2 and E. C. H. van Heusden3 

CONTENTS 

Abstract .................................................................. 2 

Introduction to the Family (C. C. BERG) ....... ..................................... ............ 2 

History and Circumscription of the Family ..... ................. ............................ 2 

Description of the Family ......... ........................................ .. .............. 3 

Key to the Neotropical Genera ................................................ ............. 3 

Genus Description of Cecropia .... ......................................................... 3 

Coussapoa (C. C. BERG AND R. W. A. P. AKKER NS) ............................................ 4 

In tro d u ctio n ............................................................................. 4 

Taxonom ic H istory ....................................................................... 4 

M o rp h o logy ........... ........................ ........................................... 4 

A natom y (K . BoNSEN ) .................................................................... 8 

Distribution and Ecology ......................... ......................................... 10 

Systematic Position of the Genus ........................................................... 15 

Systematic Arrangement of the Species ...................................................... 15 

Taxonom y ........ ........................ .............................................. 16 

Generic D escription .................................................................. 16 

K eys to the Species .................. ........ ......................................... 17 

Treatments of the Species and Subspecies .... .................................... 28 

U nnam ed Collections .... ........................ .......... .......................... 108 

Names Not Included .................................................................. 109 

Pourouma (C. C. BERG AND E. C. H. VAN HEUSDEN) .............................................. 110 

Introduction ............................................................................. 110 

Taxonom ic H istory ....................................................................... 111 

M orphology .............................................................................. 111 

Phenology ............................................................................... 113 

Pollination .................. .... ......................................................... 113 

Dispersal ............................................................................... 114 

R elations w ith A nts ....................................................................... 114 

Distribution and Ecology ......... ...... ................................ ................... 114 

Systematic Position of the Genus .... ..................... .................................. 115 

Systematic Arrangement of the Species ...................................................... 115 

U ses .................................................................................... 116 

Taxonom y ............................................................................... 116 

Generic Description .... .. ........................................................ , ... 116 

Key to the Species and Subspecies ...... ................................................ 117 

Treatments of the Species and Subspecies ............................................. 122 

Unnamed Collections ............. . ............................................ 193 

Names Excluded ................................................................. .... 194 

A cknow ledgm ents ............................................................................ 194 

Literature Cited .............. .. ...... ................................... ..................... 194 

N um erical List of Taxa ......... .............................................................. 196 

L ist of E xsiccatae ............................................................................. 197 

Index of Vernacular Names . ................................................................... 205 

Index of Scientific Names ................................... .............................. .. .. 206 

Arboretum and Botanical Garden (ARBOHA), University of Bergen, N-5067 Store Milde, Norway. 

2 Institute for Systematic Botany, State University of Utrecht, Heidelberglaan 2, 3508 TC Utrecht, The 

Netherlands. 





1

2 Flora Neotropica 

ABSTRACT 

Berg, C. C. (Arboretum and Botanical Garden [ARBOHA], University of Bergen, N-5067 

Store Milde, Norway), R. W. A. P. Akkermans and E. C. H. van Heusden (Institute for 

Systematic Botany, State University of Utrecht, Heidelberglaan 2, 3508 TC Utrecht, The 

Netherlands). Cecropiaceae: Coussapoa and Pourouma. Flora Neotropica 51: 1-000. 1990. 

The present monograph comprises the revisions of two of the three Neotropical genera of 

the Cecropiaceae: Coussapoa and Pourouma. For Coussapoa 46 species are recognized at 

present; 15 of them have been described as new during the present study. All or most species 

are hemi-epiphytes. They have entire leaves and are dioecious. The small simple flowers 

are borne in uni- to pluricapitate inflorescences. The staminate flowers have one stamen or 

two or three connate stamens. The fruits are small. The genus ranges from southern Mexico 

to south-eastern Brazil and is absent in the West Indies. Most species are components of 

lowland rain forest habitats. Few species are wide-spread, most species have relatively small 

or even very small (known) ranges of distribution. The species are morphologically clearcut 

and most of them rather uniform. For several reasons, e.g., lack of staminate material, 

the species are (provisionally) arranged in alphabetical sequences. Little is known about the 

biology of the genus. For Pourouma 25 species are presently recognized. Four taxa have 

been described as new during the preparation of this revision. All species are terrestrial, 

usually small to medium-sized trees. They have entire to palmately lobed to parted leaves 

and are dioecious. The pistillate flowers are pedicellate and are arranged in repeatedly 

branched to subumbellate inflorescences. The small staminate flowers are pedicellate or 

sessile and occur + loosely glomerate or in globose heads. The stamens are free in all species 

but one. The fruits are relatively large. The genus ranges from Central America to southeastern 

Brazil and is absent in the West Indies. Most of them occur in + disturbed and/or 

+ open places in lowland rain forest. Besides a number of clear-cut and slightly variable 

species the genus comprises a number of (very) variable and + complex species: P. bicolor, 

P. cucura, P. guianensis, P. melinonii, P. mollis, and P. tomentosa. The former three and 

the latter three species are often difficult to separate from each other (and some other species). 

These six taxa and some others show a considerable plasticity, especially with regard to the 

leaf shape, leaf dimension, and composition of the indument, and most prominently so in 

the Upper Amazon Basin. Little is known about the biology of the genus. P. cecropiifolia 

is in cultivation as a fruit tree. 

INTRODUCTION TO 

THE FAMILY 

The present monograph is comprised of re- 

visions of two Neotropical genera of the Ce- 

cropiaceae, Coussapoa and Pourouma. To place 

these revisions in a proper setting, there is pro- 

vided first a history and circumscription of the 

family, along with a brief discussion of the char- 

acters that separate the family from the related 

Moraceae and Urticaceae, a description of the 

family and the type genus, Cecropia, and a key 

to the Neotropical genera. 

History and Circumscription 

of the Family 

The family was proposed by Berg (1978a) and 

accepted by Cronquist (1981). It comprises six 

genera: three (Cecropia, Coussapoa and Pourouma) 

that are Neotropical, two (Musanga and 

Myrianthus) African, and an Asian-Australasian 

genus (Poikilospermum). The majority of the 

members of the family constituted subfamily 

Conocephaloideae in Engler's Moraceae (Engler, 

1889), but Poikilospermum and some of the 

species presently included in it were included in 

the Urticaceae. Chew Wee-lek (1963) proposed 

ranking the four microspermous genera (Cecropia, 

Coussapoa, Poikilospermum and Musanga) 

under the Urticaceae and leave the two macrospermous 

genera (Myrianthus and Pourouma) in 

the Moraceae. Corer (1962), however, regarded 

all six genera as members of the Urticaceae. The 

creation of a new family appeared to be an appropriate 

resolution to an unsatisfactory situation. 

Several features suggest that the Cecropiaceae

Introduction 

are more closely related to the Urticaceae than attached, entire, palmately-incised, or peltate and 

to the Moraceae. Poikilospermum shows more radially-incised; venation pinnate, (sub)palmate, 

urticaceous features, including the typical urti- trinervate, or radial; stipules fully amplexicaul 

caceous cystoliths, than the other five genera (cf. and connate. Inflorescences in the leaf axils, usu- 

Berg, 1989c; Bonsen & ter Welle, 1983). All gen- ally in pairs, unisexual, branched, with the flowera 

match the Urticaceae in the (sub)basal at- ers ? solitary or clustered in heads or spikes, or 

tachment of the ovule and the absence of milky unbranched with a single head or spike, bracteate 

sap. They are, however, clearly different from or ebracteate. Staminate flowers with 2-4 free or 

the Urticaceae in the absence of urticaceous (i.e., connate tepals; stamens 1-4; pistillode absent. 

explosive) stamens. Such stamens are inflexed in Pistillate flowers with 2-4 connate tepals; pistil 

the bud and, at anthesis, bend suddenly and one, ovary free from the perianth, unilocular; 

elastically outward, throwing out the pollen. The ovule one, (sub)basally attached; stigma one, 

African and Neotropical genera have a truly penicillate to peltate. Fruit an achene or somewoody 

habit (cf. Bonsen & ter Welle, 1983). Oth- times (in Coussapoa) ? drupaceous, enveloped 

er features which characterize the family are strict by a ? fleshy perianth; seed small, with endodioecy, 

watery sap that turns black after exposure sperm, or large and without endosperm; embryo 

to the air, the common occurrence of palmately- straight, cotyledons equal and flat or thickened. 

incised leaves and, in certain species, radially- A pantropical family with six genera and apincised, 

peltate leaves, large stipules (connected proximately 180 species. 

with the protection of young inflorescences), and 

adventitious aerial roots. While these roots are 

manifest as stilt-roots in most genera, in Cous- Key to the Neotropical Genera 

sapoa they are associated with the hemi-epi- 1. Lamina peltate, radially-incised. ...... Cecropia. 

phytic habit and in Poikilospermum with the 1. Lamina basally attached, entire or palmately-inclimbing 

habit. 

cised. 

The inflorescences of the 2. Staminate flowers with 1 or with 2 or 3 con- 

Cecropiaceae vary nate 

from being branched, with a + loose 

stamens; pistillate flowers sessile, in gloarrange- 

bose (to ellipsoid) heads; fruits small; stipule 

ment of the flowers (as in some staminate inflo- scars usually ascending. .......... Coussapoa. 

rescences of Pourouma), to unbranched, with a 2. Staminate flowers with 2-4 free stamens (or, 

single globose flower head (as in some pistillate 

in P. napoensis, 3-4 connate stamens); pistilinflorescences 

of Coussapoa), to a subumbellate 

late flowers pedicellate (to subsessile), solitary 

or in non-capitate clusters; fruits 

inflorescence (as in 

large; stipule 

Pourouma minor), or clusters scars horizontal. ................ Pourouma. 

of spikes enveloped by a spathe (as in most Cecropia 

species). 

In all Cecropiaceae the fruits are enveloped by 

a ? fleshy perianth, and with regard to pollination 

and dispersal, the Cecropiaceae are more 

similar to the Moraceae than to the Urticaceae. 

The genus Cecropia shows the most derived 

characters in the family, not only with regard to 

leaves and inflorescences, but also in the features 

connected with the adaptation to a mutualistic 

relation with ants, as well as in the peculiar detachment 

(abscission) of the anthers (cf. Berg, 

1977b). 

Cecropia Loefling, Iter Hispan. 272. 1758, nom. 

conserv. Type. C. peltata Linnaeus. 

Trees, terrestrial, usually with stilt-roots; internodes 

usually hollow. Lamina peltate, radially 

incised, venation radial, petiole mostly with 

1-2 trichilia (patches of dense indument, usually 

with trichomes forming Miillerian bodies) at the 

base. Inflorescences digitate clusters of spikes (or 

a single spike), usually enveloped by a spathe 

until anthesis, interfloral bracts absent. Perianth 

tubular; stamens two; stigma penicillate to peltate. 

Fruit small, dry, enveloped by a greenish 

Description of the Family 

Cecropiaceae 

Trees or shrubs, terrestrial or hemi-epiphytic, 

with aerial roots, and watery sap turning black 

upon exposure. Leaves in spirals; lamina basally 

fruiting perianth. 

With approximately 80 species throughout the 

Neotropics, with a distinct concentration of 

species in the Andean region. 

Revisions of Coussapoa and Pourouma are 

presented sequentially below. 

3


COUSSAPOA 

Engler's system of the Moraceae (Engler, 1889). 

This subfamily was transferred to the Urticaceae 

Introduction 

by Corner (1962), but has recently been recognized 

as a 

Coussapoa, one of the three 

separate family Cecropiaceae (Berg, 

Neotropical genera 

of the Cecropiaceae, is a well-defined 

1978a; Cronquist, 1981). 

genus. 

The majority of its species are arboreous hemiepiphytes, 

a peculiar life-form shared with a few 

Morphology 

other genera like Ficus (subg. Urostigma), Clusia, Habit. - Coussapoa species are shrubs or more 

and Schefflera. The species are rather similar in 

their ecology and show little variation in 

commonly small to medium-sized trees (up to 

morca. 

20 m 

phological characters. Knowledge of their 

tall), but they can sometimes form 

biolhuge 

wide-crowned trees up to 40 m tall, 

ogy is very limited. 

e.g., C. curranii 

and C. trinervia. On labels the 

The poor state of botanical exploration of 

height is 

many 

regions of the Neotropics has adversely affected 

probably not always correctly stated, because for 

the present revision, as is evident from the fact 

epiphytes the host often appears to be included. 

Most 

that it has not proved possible to make a 

Coussapoa species are hemi-epiphytic or 

systematic 

arrangement of the species and from the 

secondarily terrestrial. Probably only C. trinervia 

is 

way in which the keys have had to be constructbasically 

terrestrial. As in Ficus, Coussapoa 

ed. In about one-third of the species only one of 

species can be epilithic. The (hemi-)epiphytic 

the sexes is known. Many species are known from 

growth habit is evident in the presence of aerial 

roots. These aerial roots can form baskets of roots 

only one or at most just a few specimens. More 

around the stem of the host tree. Sometimes, the 

than one-quarter of the species now known have 

aerial root 

been recognized as new during the 

system consists of a main (vertical) 

preparation axis with at more or less 

of this revision; many of them are based on reregularly 

distanced pairs 

of horizontal branches 

cent collections. The increase in the number of 

clasping the trunk of the 

host tree. In 

species suggests that further exploration will lead 

general, hemi-epiphytic Coussapoa 

to the discovery of additional 

species are not true (or vigorous) stranglers like 

species. 

hemi-epiphytic Ficus species. 

Most Coussapoa species have straight, stiff 

Taxonomic 

branches. The smaller branches can be hollow 

History 

(widely so in C. asperifolia, but narrowly in sev- 

Coussapoa was established by Aublet (1775), eral other species, e.g., C. villosa). When branchwho 

described two species from French Guiana. es are hollow they are often inhabited by biting 

Names were added by Trecul (1847), Klotzsch ants (Azteca?) or stinging ants (often in the case 

(1847), Miquel (1853), and later on by Standley of C. villosa). These insects bore openings to the 

(1919, 1924, 1930, 1937a, 1940), Mildbraed interior of the branches. Like (most?) other gen- 

(1928, 1942), and Cuatrecasas (1951, 1956b), era of the Cecropiaceae, when wounded Cousamong 

others, so that ca. ninety names were cre- sapoa species exude a (more or less mucilagiated 

for thirty-four of the taxa recognized in the nous) watery sap which turns black after exposure 

present paper. In the course of the preparation to the air. 

of present treatment 15 new species were de- Indumentum. -Four types of trichomes can 

scribed (Akkermans & Berg, 1982; Berg, 1983). be recognized: 

Taxonomic, mostly floristic, treatments of the 

genus are to be found in the work of Trecul (1847), 1. Unicellular arachnoid (cobwebby) hairs. These 

Flora brasiliensis (Miquel, 1853), Flora of Peru very thin, crinkled, interwoven hairs of dif- 

(Macbride, 1937), Flora of Panamai (Woodson & ferent lengths are mostly white, sometimes 

Schery, 1960), Flora of Suririame (Berg, 1975), brownish. 

and Flora Costaricensis (Burger, 1977). Thus, 2. Unicellular non-arachnoid, thicker and mostsince 

the treatments by Trecul and Miquel, the ly more or less straight hairs of different lengths 

genus has never been studied as a whole. and white, yellow, or brown. 

The genus belongs to a group of genera that 3. Pluricellular hairs-dark or pale brown, short 

constitute the subfamily Conocephaloideae in and subglobose to long and moniliform. These

Coussapoa 

hairs are very common on young leaves, stipules, 

inflorescences, etc. In dry material they 

are more or less granular (powdery) in appearance. 

4. Pearl bodies (or glands)-hyaline, succulent, 

more or less club-shaped, and confined almost 

entirely to the leaves. These trichomes 

are found in specimens of Coussapoa microcarpa 

and C. villosa cultivated in green houses. 

Pearl bodies are probably common in the genus, 

but are often not found in the field because 

they are easily detached; moreover, they 

are probably harvested by ants (cf. O'Dowd, 

1982). 

As with many other genera of the Cecropiaceae, 

these four types of hairs are also often 

encountered in Coussapoa species. Many species 

have a dense or rather dense indumentum. In 

several species arachnoid hairs are lacking and 

most of these species have sparse indumentum 

as well. 

common in elliptic leaves, and in obovate leaves 

normally there is none. Venation types in the 

genus can be seen in Figure 1. 

The ovate lamina with branched, basal, lateral 

veins and a relatively long petiole can be regarded 

as the least derived type of leaf in the genus, 

as it shows the closest resemblance to leaves in 

other genera of the family, especially Pourouma. 

The leaves of P. minor resemble those of C. contorta 

and the leaves of P. tomentosa and P. meliononii 

are reminiscent of those of C. villosa and 

C. nitida, respectively. The obovate, trinervate 

lamina with a short petiole can be regarded as a 

derived type of leaf. Another type of leaf, which 

can also be regarded as derived, is that found in, 

e.g., C. parvifolia and C. contorta. This type of 

leaf has an elliptic to subovate lamina with many 

unbranched lateral veins, a very regular venation, 

and a relatively short petiole. These features 

are characteristic more of the Moraceae than of 

the Cecropiaceae. 

The indumentum of the leaves also shows some 

Leaves. -The petiole in most species is rela- variation in denseness and occurrence of differtively 

long-up to half the length of the lamina. ent types of hairs. The occurrence of a dense and 

There is a tendency towards shortening of the varying indumentum appears to be weakly corpetiole 

(to one-tenth of the length of the lamina) related with primitiveness in other leaf characin 

species with derived (i.e., obovate, trinerved) ters. 

types of lamina. The relative length of the petiole C. contorta is distinct in having domatiumis 

not subject to great variation as is the case in like cavities in the axils of the lateral veins. 

related genera, such as Myrianthus and Pou- The stipules are connate, fully amplexicaul and 

rouma. 

caducous, usually leaving oblique scars, char- 

The length of the lamina varies from more acteristic for the genus. The length varies conthan 

50 cm to less than 10 cm and more or less siderably in the genus. One can roughly recognize 

parallels the change in the shape of the lamina three categories: long stipules (5-15 cm long or 

from cordiform, subreniform, or broadly ovate longer), medium-long stipules (1-5 cm long) and 

to elliptic or oblong to obovate or subobovate; short stipules (less than 1 cm long). Long stipules 

at the same time, this correlates with a gradation give protection for a longer time to young leaves 

in the leaf base from cordate to truncate to and developing inflorescences, but a correlation 

rounded to obtuse to subacute. 

between the type of inflorescence or habit pref- 

The venation is pinnate and brochidodromous erence and the length of the stipules is not apwith 

a more or less regular parallel tertiary ve- parent. 

nation in the intercostal area. The venation may Inflorescences. - Coussapoa, like all other gentend 

toward subpalmate (C. vannifolia), or may era of the Cecropiaceae, is dioecious. Monoebe 

more or less clearly trinervate (C. trinervia, cious specimens are rare. 

C. cinnamomifolia), with this development being The inflorescences occur in pairs in the leaf 

combined with a change towards a (sub)obvate axils, a feature common in the Cecropiaceae and 

lamina. In ovate leaves the basal pair of lateral related families. The ramification of the inflovein 

branches is mostly branched. Especially in rescence is basically similar to that usually found 

large leaves, the other lateral veins show little in Myrianthus (see: De Ruiter, 1976) and Poubranching 

(being mostly only furcate); however, rouma. The common peduncle bears at its apex 

in C. asperifolia the other lateral veins are mostly (at least) three branches, a pattern repeated in 

branched. Branching of the lateral veins is less secondary and further ramifications. However, 

5


Ni 

3 4 5 

FIG. 1. Schematic drawings of types of venation in Coussapoa: 1, most lateral veins branched; 2, 3, only 

the basal pair of lateral veins branched; 4, lateral veins unbranched, the basal pair reaching the margin above 

the middle; 5, three-nerved. 

2

Morphology 

T 

FIG. 2. Schematic drawings of types of pistillate inflorescences in Coussapoa. 

FIG. 2. Schematic drawings of types of pistillate inflorescences in Coussapoa. 

reduction of the number of branches to two, re- bracts can be subpeltate or more or less cucullate. 

sulting in a (sub)dichotomous ramification, is also Sometimes, these bracts are truly peltate--as in 

found, especially in the ultimate ramification, C. contorta. Bracts outside the flower heads are 

and may occur down to the primary ramification mostly scale-like and ovate to oblong. 

as well. Pistillate inflorescences are schematized Staminate Flowers.-The small staminate 

in Figure 2. 

flowers have a tubular perianth which is 3-lobed 

The sessile flowers are arranged in terminal at the apex. The number of stamens is 3, 2, or 

globose heads. Solitary (often more or less abort- 1. In 3- and 2-staminate flowers the stamens are 

ed) flowers may occur elsewhere in the inflores- fused, forming a stalk with, respectively, 6 (or 

cence, but their presence is mostly connected with occasionally 5) or 4 thecae at the apex. The numpoints 

of branching. Solitary flowers are rare in ber of stamens is very constant and may prove 

pistillate inflorescences. 

to be the (only?) basis for subdivision of the genus 

The staminate inflorescences are normally re- and arrangement of the species. The length of 

peatedly branched and bear small terminal heads. the filaments varies from about as long as the 

In pistillate inflorescences, however, the number perianth to more than twice as long as the periof 

flower heads is often reduced to one, due to anth, the differences being distinctive at the 

reduction of the ramification and/or to fusion of species level. 

the flower heads. In some species, e.g., C. bre- Pistillate Flowers. -The small pistillate flowvipes, 

C. duquei, the flower heads are ellipsoid ers have a tubular perianth with a narrow (alto 

clavate and combined with a very short pe- most) entire opening that allows the style through. 

duncle. 

The flowers are free, but in C. parviceps the mar- 

The inflorescences are bracteate in the major- gins of the flattened apices of the perianths of 

ity of the Coussapoa species. The bracts are mostly adjacent flowers cohere. The ovary is free and 

confined to the flower heads, but sometimes oc- the stigma is penicillate to subpeltate. 

cur elsewhere in the inflorescence, as in the sta- Staminate and pistillate flowers and their asminate 

inflorescences of C. tessmannii. In some sociated bracts are shown schematically in Figspecies, 

e.g., C. microcarpa and C. viridifolia, the ure 3. 

interfloral bracts are very small; they may also Fruits and Seeds.-The small fruits are free 

be few in number or sometimes even absent. In from the perianth. The pericarp consists of a 

C. parviceps, the only species in which the pis- rather thick and crustaceous endocarp and a very 

tillate flowers are connate (or cohering), inter- thin to rather thick, slightly fleshy to mucilagifloral 

bracts are absent in the pistillate inflores- nous layer. Thus, depending on the features of 

cence but present in the staminate. This lack of the exocarp, the ellipsoid to ovoid, 1-3 mm long 

interfloral bracts is apparently a derived state. fruitlets are achene-like or more or less drupa- 

The interfloral bracts are more or less distinct- ceous. Fruits are depicted schematically in Figly 

spathulate, often with the lower part very nar- ure 4. 

row (and stipe-like). Relatively large interfloral In C. asperifolia, while the exocarp is present 

7


1 2 3 4 5 6 7 a ' 9. 

FIG. 3. Schematic drawings of flowers and interfloral bracts in Coussapoa: 1-3, staminate flowers, 1, with 

three connate stamens, 2, with two connate stamens, 3, with one stamen; 4, pistillate flower; 5-9, various 

interfloral bracts. 

as a very thin (membranaceous) outer layer, the 

mesocarp is relatively thick and expands in such 

a way as to push the endocarp body out of the 

perianth, the apex of which becomes torn in the 

process. Whether this phenomenon is confined 

to C. asperifolia or also occurs in other species 

is not clear. At any rate, it is not general in Coussapoa, 

as previously assumed (Berg, 1975, 1977a). 

In many (or all?) species the fruiting perianth 

becomes more or less juicy, and it is also often 

colored-orange, yellow, or brown. Sometimes, 

as in C. villosa, the perianth remains greenish, 

at least the apical part. 

Coussapoa is one of the four microspermous 

genera of the Cecropiaceae. In the genus microspermy 

is related to the (hemi-)epiphytic habit, 

as well as to the occurrence of a mucilaginous 

layer (cf. Berg, 1983b). 

Phenology, Pollination, and Dispersal. - 

Available herbarium annotations suggest that in 

general Coussapoa species flower (and fruit) 

throughout the year. 

Nothing is known about pollination in Coussapoa. 

Neither the structure of the inflorescences 

nor the features of the stamens suggest the occurrence 

of wind pollination, however. 

In most cases dispersal is probably endozo- 

ochorous, often by frugiferous birds, but also by 

monkeys (Dr. M. van Roosmalen, pers. comm.). 

Exozoochorous dispersal may occur when the 

exocarp is more or less mucilaginous (and sticky), 

as in C. asperifolia. 

Anatomy 

By K. BONSEN 

Secondary Xylem. -General Properties. Col- 

or: light yellowish brown. Grain: straight, some- 

times slightly interlocked. Texture: medium. 

Odor and taste: not characteristic. Specific grav- 

ity: 0.50-0.75. 

Structure. Based on eight species, nine speci- 

mens, the growth rings are faint or absent. Ves- 

sels diffuse, round to oval, solitary (25-83%), and 

in radial multiples and irregular clusters of 2-8 

(-21), (0-1)1-6(-11) per sq mm, diameter (200-) 

220-300(-340) gm. Vessel member length (400) 

475-600(-725) Mm. Perforations simple. Inter- 

vascular pits alternate, round or polygonal, 10- 

15 Lm. Thin-walled tyloses are usually present. 

Fibers non-septate, diameter 18-25 ,m, walls 2- 

3.5 tm, L/W ratio 2-5, gelatinous fibers usually 

present. Pits simple, mainly on the radial walls. 

a ad tfg 

1 2 34 5 6 

FIG. 4. Schematic drawings of fruits in Coussapoa: 1-4, C. asperifolia type, 1, 2, fruit in perianth, 3, fruit 

in opened perianth, 4, endocarp body; 5, 6, C. villosa type, fruit in perianth. a, embryo; b, endosperm; c, testa; 

d, endocarp; e, mucilaginous mesocarp; f, exocarp; g, perianth.

Anatomy 

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Length (875-)1100-1800(-2175) .m, F/V ratio 

2.5-3.7. 

Rays heterogeneous, uniseriate (21-35%) and 

multiseriate (3-)4-7(-9) per mm, sheath cells 

present or absent. Uniseriate rays mainly composed 

of square to upright cells, ray height 

(200-)300-500(-980) gtm. Multiseriate rays 

composed of upright and procumbent cells, vertically 

compounded 1-10%, (450-)700-1100 

(-1600) ,m high, 3-6 cells in width, uniseriate 

parts (0-)1-2(-8) cells. 

Parenchyma paratracheal, banded, irregular, 

wavy, (0-)1-2(-3) per mm, (3-)5-9(-15) cells in 

width. Strands 5-8(-14) cells, length (530-)600- 

710(-870) Mm, containing some to many rhombic 

crystals. Radial latex tubules have been observed 

in Coussapoa latifolia (see Fig. 5). 

The wood of Coussapoa resembles that of Myrianthus 

and some species of Pourouma. The 

presence of confluent-banded parenchyma in the 

wood of Coussapoa distinguishes this genus from 

the wood of Cecropia and Musanga (Bonsen & 

ter Welle, 1983). 

' 

. . . . 

Leaf Anatomy. -Structure. Based on 11 

species, 18 specimens. 

In surface view. Indumentum of thin, inter- 

woven, unicellular arachnoid hairs, abaxial pres- 

ent or absent; unicellular needle-shaped, often 

wavy hairs (mostly on abaxial surface, rarely on 

adaxial surface); adaxial, glandular hairs with 

multicellular, globular heads on 3-5-celled, uni- 

seriate stalks, mostly in groups of 2-4, present 

or absent; abaxial, uniseriate, 6-10-celled, curved, 

glandular hairs with globular to elongated heads, 

mostly present; and conical papillae sometimes 

present. Epidermal cells polygonal; adaxial cells 

overlying large crystalliferous mesophyll cells 

forming a rosette. Stomata confined to abaxial 

surface, anomocytic, average length of guard cell 

pairs 15-20 gm, average width 12-18 um. Hy- 

dathodes formed by 10-15 water pores each, 

present or absent on adaxial surface. Minor veins 

usually very prominent in abaxial epidermis. 

In transverse section. Lamina bifacial. Epi- 

dermal cells small, especially abaxially between 

the veins. Adaxial epidermal cells sometimes with


silicified outer walls. Stomata sometimes raised higher altitudes (from ca. 800 m) in the Guiana 

above level of unspecialized cells. Adaxial hy- (=Guayana) Highland region, e.g., C. argentea 

podermis of two or three layers of parenchyma and C. viridifolia, while others are found in higher 

cells, including mucilage cells except for C. vil- altitudes (from ca. 900 m or from ca. 1400 m) 

losa. Mesophyll consisting of one layer of pali- in the Andean region, e.g., C. cinnamomifolia 

sade cells, sometimes subdivided loose spongy and C. duquei; C. crassivenosa is found in both 

tissue, with or without an intermediate layer. regions. Some species also occur in drier habitats, 

Veins with sclerenchymatous vertical bundle such as savanna (C. viridifolia) or semi-decidusheath 

extensions (touching adaxial hypodermis ous forest (C. purpusii). Most lowland species 

and abaxial epidermis). Midrib with a flat or appear to occur both in upland and riverine forraised 

adaxial surface and a prominently raised est. A few species, like C. nitida, appear to show 

abaxial surface; peripheral ground tissue paren- a preference for riverside (varzea) forest. C. trichymatous 

to collenchymatous, interspersed with nervia, the only truly terrestrial species of the 

mucilage cells; vascular system composed of a genus, is confined to riverbanks, apparently 

closed or variously interrupted cylinder, partly (only?) of black-water rivers. 

or wholly surrounded by sclerenchyma fibers, and Only two species, C. villosa and C. asperifolia, 

enclosing one or two rows of bundles which are have an extended distribution. Most of the other 

situated in the same direction as the most abaxial species are confined to, or at least distinctly asbundle 

of the cylinder. Vascular system of petiole sociated with, one of the phytogeographic resimilar. 

Crystals present as druses throughout the gions, such as eastern Brazil, Amazon Basin, 

mesophyll, in the petiole and midrib; rhombic eastern Guiana (=Guayana Lowland) region, 

crystals sometimes present. Distinguishment of western Guiana (=Guayana Highland) region, 

the inner structure of the leaves of Coussapoa Andean region, Pacific coastal region, and Cenfrom 

those of the other Neotropical genera, Ce- tral America. Many species are only known from 

cropia and Pourouma, can be accomplished on a very small area. 

the basis of the petiole vascular system type Several species with medium-sized areas show 

(Bonsen & ter Welle, 1983). Some differences in a discontinuous distribution. It is, however, not 

the inner structure of some species within the certain whether these discontinuities are real or 

genus Coussapoa are mentioned by Renner due to insufficient exploration. The Amazonian 

(1907). 

C. trinervia has a disjunct distribution, which 

appears to be related to its ecology (see above). 

Distribution and C. crassivenosa is known from 

Ecology 

four remote regions. 

C. latifolia and C. asperifolia ssp. asperi- 

The genus, being distinctly associated with the folia, common taxa of the eastern Guiana region 

rain forest habitat, is widespread in tropical to eastern Parf, show similar gaps in distribution 

America, but is absent in the West Indies; see and are found segregated in more central parts 

Figures 6-9. 

of the Amazon Basin as well. If these gaps are 

Most species are components of lowland rain real, the same factors may have caused the westforests. 

A few species, e.g., Coussapoa villosa and ern Parf gap in the distribution of C. nitida, as 

C. asperifolia, extend into submontane or mon- well as the disjunct distribution of the species 

tane forest. Some species are associated with pairs C. arachnoidea-C. orthoneura and C. le- 

FIG. 6. Distribution of Coussapoa angustifolia, C. arachnoidea, C. argentea, C. asperifolia ssp. asperifolia, 

C. asperifolia ssp. magnifolia, C. asperifolia ssp. rhamnoides, C. batavorum, C. brevipes, C. chocoensis, C. 

cinnamonea, C. cinnamomifolia, C. contorta, C. crassivenosa, and C. cupularis. 

FIG. 7. Distribution of Coussapoa curranii, C. duquei, C. echinata, C. ferruginea, C. floccosa, C. glaberrima, 

C. herthae, C. latifolia, C. leprieurii, C. macerrima, C. microcarpa, C. microcephala, C. napoensis, and C. 

parvifolia. 

FIG. 8. Distribution of Coussapoa longepedunculata, C. manuensis, C. nitida, C. nymphaeifolia, C. oligocephala, 

C. orthoneura, C. ovalifolia, C. pachyphylla, C. parviceps, C. purpusii, C. scabra, and C. sprucei. ? 

unnamed collection aff. C. ovalifolia.

Distribution and Ecology 

0 C.angustifolia 

Aublet 

OC.arachnoidea Akkermans 6 Berg 

^^-^^'.L_ n //i C ^ 4-Lc


v* r 

r'^S'9n 7 

J 

(^^'^^J^'^ 

- ^^Q^0 C.duquei Standley 

A C.echinata Akkermans Berg 

* C.curranli Blake i . e 

- 

T C.ferruginea Trecul 

| ? (% 'J/' / { , t 

__l 

|: 

A C.glaberrima 

Burger 

* C.herthee 

S Mildbred 

C.p_rvi fo ^g d C-latifolia Aublet 

C .floccosa i 

y 

Akkermans Berg . 

B 

C.cerricroc Akkerpa (SchQtt) Berg izzini ~ - 

0 C'microcephala Trdcul ' 

V' C.parvifolia Standley 

A C.napoensis Akkermmns & Berg 

,.


OC.nitida Mfquel /). ^~:t ,._ A *C.nymphaeifolia Standley 

~ 

.AC.oligo ala Donnael Smith 

C.orthoneura Stand ey 

*C.pachyphylla Akkermans & Berg 0 C.ovalifolia Trcul C . 

HC.pcyhl tcuatl Aknn Tra ~ t C.oval 1 

i fol ia 

C.parviceps Standey\ 

C.scabra Akkermans 6 Berg 

*C.sprucel 

Mildbraed 

\ 

% 

13

1.trinervia 

*C.villosa Poeppig 6 Endlicher 

AC.viridifolia 

Cuatrecasas 

I 

C. tessmannll M Ildbrwd 

A C. fulvescens C.C. Berg 

VC. tolimensis C.C. Berg 

Ce. voiaria C.C. Berg 

C 

Mil dbraed 

A C.vannifolia Cuatrecasas 

FIG. 9. Distribution of Coussapoa 

fulvescens, C. tessmannii, C. tolimensis, C. trinervia, C. vannifolia, C. 

villosa, C. viridifolia, and C. valaria (name is misspelled in legend).

Systematic Position 15 

prieurii-C. sprucei. This type of distribution 

may be connected with the distribution of Pleistocene 

rain forest refugia (cf. Prance, 1977). 

Another remarkable discontinuity in distribution 

occurs with C. cinnamomifolia, a species of 

medium altitudes in the Andean region. This 

discontinuity shows resemblances to that of two 

partial areas of C. crassivenosa. The Ecuador- 

Bolivia disjunction is probably not genuine and 

more likely to be due to inadequate exploration 

in Peru. 

The discontinuities in the distribution of C. 

microcarpa, especially the isolated localities in 

Perambuco and Paraiba, could be related to the 

more or less isolated occurrence of forest with 

rain forest elements on low mountain tops. 

sion of the diaspores is another feature which the 

two genera have in common; in Coussapoa it is 

(probably) confined to C. asperifolia and in Poi- 

kilospermum it is only found in subg. Liguli- 

stigma (cf. Chew Wee-Lek, 1963). This phenom- 

enon, in combination with microspermy, is 

probably associated with the hemi-epiphytic 

habit. In Coussapoa the mucilaginous layer that 

swells and pushes the endocarp out of the 

perianth is formed by the middle layer of the 

pericarp. In Poikilospermum, however, the ex- 

pulsion of the fruits is achieved by the swelling 

of the inner surface of the perianth, which be- 

comes mucilaginous during the ripening of the 

fruit (cf. Chew Wee-Lek, 1963). 

Judging from (the nature of) the similarities 

In the present treatment, several pairs of closely and differences, the two genera are ecological 

related species are recognized, e.g., C. crassive- equivalents rather than systematically closely renosa-C. 

ferruginea, C. latifolia-C. viridifolia. lated. Morphological as well as anatomical char- 

These pairs of species have allopatric distribu- acters (cf. Bonsen & ter Welle, 1983) suggest a 

tions. 

closer relationship of Coussapoa to the other five 

genera of the family, especially to the Neotrop- 

Systematic Position of the Genus ical ones. Coussapoa matches Cecropia in microspermy. 

The two genera show similarities in 

Coussapoa, together with the Neotropical gen- the flowers, but otherwise are quite different. 

era Cecropia and Pourouma, the African genera More similarities are found between Coussapoa 

Musanga and Myrianthus, and the Asiatic genus and Pourouma, although the latter genus is mac- 

Poikilospermum, constitute the family Cecropi- rospermous. The inflorescences of Coussapoa reaceae 

(Berg, 1978). 

semble the staminate ones of some Pourouma 

Poikilospermum, geographically isolated from species, e.g., P. tomentosa and P. mollis; some 

the other genera, has more characters, especially Pourouma species have leaves strongly reminisanatomical 

ones, in common with the Urticaceae cent of those common in Coussapoa (see above). 

than the other genera (cf. Bonsen & ter Welle, 

1983). The latter authors, indeed, suggest trans- 

Systematic 

ferring the 

Arrangement 

genus back to the Urticaceae. Neverof 

the 

theless, considering the distribution of 

Species 

characters 

throughout the (complex of) families Cecropi- In the present revision the species are treated 

aceae, Urticaceae, and Moraceae, placing Poi- in alphabetical order, as it is not possible to prokilospermum 

in the Cecropiaceae appears to be pose a satisfactory systematic arrangement. 

justified, although not beyond all doubt. For most characters clear discontinuities in 

Both Coussapoa and Poikilospermum are pre- their variation are lacking. Primitive and derived 

dominantly hemi-epiphytic (cf. Chew Wee-Lek, state (as the extremes of a morphological series) 

1963). However, the two genera differ in habit! can be inferred, like lamina broadly ovate-nar- 

Coussapoa species are shrubs or trees with straight rowly obovate, basal lateral veins branched-unand 

stiff branches, while Poikilospermum species branched, lateral veins numerous-few, petiole 

are more or less liana-like with long and weak long-short, stipules long-short, indumentum with 

(scrambling) branches (cf. Chew Wee-Lek, 1963). all types of hairs and ? dense-less diverse and 

The two genera resemble each other in the ? sparse, inflorescences repeatedly branchedbranched 

inflorescences with terminal globose unbranched, interfloral bracts present-absent, and 

heads, but the basic ramification of the inflores- stamens three-two. See Table I. 

cence appears to be different. Dichotomous ram- In some species (e.g., C. nymphaeifolia, C. vilification 

appears to be basic in Poikilospermum, losa, and C. tessmannii) a predominance of 

but is derived in Coussapoa (see above). Expul- "primitive" characters can be found or a pre-


Table I 

Extremes in a morphological character series in 

Coussapoa, inferred to be primitive and derived 

respectively. 

Primitive states Derived states 

Lamina broadly ovate Lamina narrowly obovate 

Basal lateral veins Basal lateral veins unbranched 

branched 

Lateral veins numerous Lateral veins few 

Petiole long 

Petiole short 

Stipules long Stipules short 

Indument with all types Indument less diverse 

of hairs and ? dense and + sparse 

Inflorescences repeatedly Inflorescences unbranched 

branched 

Interfloral bracts present Interfloral bracts absent 

Stamens 3 Stamen 1 

number of stamens in arranging the species, since 

for about one-third of the species recognized material 

belonging to only one sex is known. 

Acceptance of the number of stamens as the 

basis for subdividing the genus would imply the 

occurrence of parallel (or homologous) differentiation 

of many characters, e.g., in the differentiation 

of the leaf shape, dimensions, venation, 

indumentum, and length of the petiole, in three 

groups of species (judging from the comparison 

of characters in the taxa of which the number of 

stamens is known). 

Taxonomy 

1. Coussapoa Aublet, Hist. pl. Guiane 2: 955. 

1775; Trecul, Ann. Sci. Nat. Bot. Ser. 3, 8: 92. 

1847; Miquel, Fl. bras. 4(1): 131. 1853; Macbride, 

Publ. Field Mus. Bot. 13(2.2): 295. 1937; 

Woodson & Schery, Ann. Missouri Bot. Gard. 

47: 168. 

dominance of"derived" characters in others 

1960; Berg, Fl. Suriname 5(1): 279. 

(e.g., 

C. cinnamomifolia, C. trinervia, and C. viridi- 

1975; Burger, Fieldiana Bot. 40: 131. 1977; 

Akkermans & 

folia), with the majority of the species some- 

Berg, Proc. Kon. Ned. Akad. 

Wetensch. Ser. 

where in between. A few species have 

C, 85(4): 441. 1982. Lectotype 

exceptional 

characters: C. trinervia has a truly terrestrial hab- species (Berg, Fl. Suriname 5(1): 279. 1975): 

it, C. parviceps has fused pistillate flowers; C. 

Coussapoa latifolia Aublet. 

contorta has domatium-like cavities in the axils Hemi-epiphytic (or epiphytic) or terrestrial 

of the lateral veins; C. asperifolia has abruptly trees or shrubs with aerial roots or stilt-roots. 

thickened twigs with wide cavities and, more- Leaves in spirals, entire, venation pinnate to triover, 

a thick mucilaginous mesocarp, and C. pur- nervate, margin entire to subcrenate, stipules 

pusii is (often?) deciduous. 

fused, fully amplexicaul, usually leaving oblique 

The difference in the number of stamens ap- scars. Inflorescences mostly in pairs in the leaf 

pears to be a promising basis for the arrangement axils, branched or in pistillate ones often unof 

the species or the subdivision of the genus. branched, bracteate (mostly only with interfloral 

The significance of these differences lies not only bracts) or ebracteate. Flowers in globose (to elin 

the absence of transitions but also in the dis- lipsoid or clavate) terminal heads, free or pistiltribution 

ranges: the predominance of species with late ones sometimes connate, perianth tubular, 

3-staminate flowers in Central America and the in staminate flowers (2-)3(-4)-lobed, in pistillate 

Pacific coastal region of Colombia and Ecuador; ones entire; stamens two or three and connate, 

the predominance of species with 2-staminate or one; ovary free, stigma penicillate to subpelflowers 

in eastern South America, especially east- tate. Fruit a drupelet or almost an achene, enern 

Brazil; and the concentration of species with closed in the enlarged + fleshy perianth; seed 

uni-staminate flowers in the Upper Amazon Ba- with endosperm, embryo straight with flat and 

sin. It is not yet possible, however, to use the equal cotyledons and relatively short radicle.


General key 

Keys to the species in: 

1. Central America and Mexico 

2. Colombia 

3. Venezuela 

4. the Guianas2 

5. Ecuador 

Keys to the Species of Coussapoa 

6. Peru and Bolivia " \ . 7 / 

7. Amazonian Brazil 

8. Eastern Brazil 

General Key to the Species of Coussapoa 

1. Interfloral bracts absent. 

2. Basal lateral veins branched. 

3. Lamina beneath with dense indumentum of minute hairs, at least in the areoles and on the 

reticulum. 

4. Indumentum (unicellular hairs only!) predominantly white; pistillate inflorescences unicapitate; 

stamens 3. 

5. Intercostal venation prominent beneath; Costa Rica. .............. 29. C. nymphaeifolia. 

5. Intercostal venation plane beneath; Panama. ............................. 6. C. brevipes. 

4. Indumentum (unicellular hairs only!) at least partly brownish or yellow; pistillate inflorescences 

usually pluricapitate; stamen 1 or unknown; Amazon Basin or Colombia at ca. 1900 m. 

6. Lamina usually elliptic; lateral veins 5-9 pairs. ..................... See 12. C. cupularis. 

6. Lamina usually ovate; lateral veins 8-15 or 14-22 pairs. 

7. Lateral veins 8-15 pairs; peduncle of the pistillate inflorescence 1-2.5 mm thick. 

8. Intercostal venation (almost) plane; indumentum of petiole consisting of dense 

minute hairs and distinctly longer, yellowish hairs; Amazon Basin ..... 28. C. nitida. 

8. Intercostal venation prominent; indumentum of the petiole consisting of hairs not 

distinctly different in length; Colombia (ca. 1900 m). ............. 45. C. tolimensis. 

7. Lateral veins 14-22 pairs; peduncle of the pistillate inflorescence 6-10(-15) mm thick; 

Colombia (Amazonas). ......................................... 8. C. cinnamomea. 

3. Lamina beneath with sparse indumentum or glabrous, at least in the areoles and on the reticulum. 

9. Stipules 0.5-1 cm long; Venezuela (Bolivar and Amazonas), Guyana, and Brazil (Amazonas, 

R oraim a). ........................................ .................. 43. C. viridifolia. 

9. Stipules at least 1 cm long. 

10. Lateral veins 3-6 pairs, the basal pair reaching the margin at or above the middle of the 

lamina; pistillate flowers cohere; stamens 3; Central America and Pacific coastal region 

of Colombia and Ecuador. ...........................................34. C. parviceps. 

10. Lateral veins 7 or more pairs, the basal pair reaching the margin below the middle of the 

lamina; if less than 7 pairs of lateral veins then the pistillate flowers free and stamen 1. 

11. Lamina and leafy twigs glabrous; base of the lamina cordate; Colombia (Choco). 

.............................................................. 5. C. batavorum . 

2 

17


11. Lamina and leafy twigs at least sparsely puberulous; base of the lamina acute to 

truncate. 

12. Stipules (normally) 1-4 cm long. 

13. Margin of the lamina revolute at the base; Pacific coastal region (Colombia). 

......................................................... 46. C. valaria. 

13. Margin of the lamina not revolute at the base; Amazon Basin and Central 

America. 

14. Lamina coriaceous, mostly ovate; stamen 1; Amazon Basin (and Panama?). 

............................................. 32. C. ovalifolia. 

14. Lamina subcoriaceous, elliptic to subobovate; stamens 2; Costa Rica. 

........................................ 23. C. macerrima. 

12. Stipules 4-13 cm long; Amazonian Ecuador and Colombia. .... 27. C. napoensis. 

2. Basal lateral veins unbranched (or occasionally with a single branch). 

15. Stipules 0.5-1 cm long; stamens 2; common peduncle or branches of the pistillate inflorescence 

not distinctly broadened towards the flower heads; eastern Venezuela, Guyana, and Brazil 

(Am azonas, Roraim a). ................................................... 43. C. viridifolia. 

15. Stipules (normally) at least 1 cm long, if shorter, then stamen 1 and branches of the pistillate 

inflorescence more or less distinctly broadened towards the flower heads. 

16. Margin of the lamina distinctly revolute towards the base; pistillate inflorescences mostly 

unicapitate; stamens 2; eastern Brazil. ................................ 25. C. microcarpa. 

16. Margin of the lamina plane, or if more of less revolute, then the pistillate inflorescences 

pluricapitate and stamen 1; Amazon Basin or Central Brazil. 

17. Lateral veins 2-8; basal pair of lateral veins reaching the margin above to just below 

the middle of the lamina and/or (often) the lamina beneath with dense, minute hairs 

in the areoles. 

18. Intercostal venation prominent beneath; lamina beneath and inflorescences with 

dense, white, arachnoid indumentum; Brazil (Amapa) .......... 2. C. arachnoidea. 

18. Intercostal venation plane beneath; lamina and inflorescences without arachnoid 

indumentum, or if present, then mostly sparse and soon disappearing; Upper 

Amazon Basin. 

19. Basal lateral veins reaching the margin above to just below the middle of the 

lamina; stipules mostly without or with sparse, long, yellow hairs; branches 

of the pistillate inflorescence slightly broadened towards the flower heads; 

Upper Amazon Basin and Colombia (Santander). .......... 31. C. orthoneura. 

19. Basal lateral veins reaching the margin below to just above the middle of the 

lamina; stipules with dense, long, yellow hairs; branches of the pistillate inflorescence 

strongly broadened towards the flower heads; Brazil (R6ndonia). 

......................................................... 12. C. cupularis. 

17. Lateral veins (normally) 7-13 or 7-21 pairs, basal pair reaching the margin far below 

the middle of the lamina; lamina beneath glabrous or with sparse hairs in the areoles. 

20. Stipules (normally) 1-4 cm long. 

21. Lamina coriaceous, mostly ovate; stamen 1; Amazon Basin (and Panama?). 

......................................................... 32. C. ovalifolia. 

21. Lamina subcoriaceous, elliptic to subobovate; stamens 2; Costa Rica. ..... 

........................................ 23. C. macerrima. 

20. Stipules 4-13 cm long; Amazonian Ecuador and Colombia. ...... 27. C. napoensis. 

1. Interfloral bracts present (sometimes small and/or few!). 

22. Lamina trinervate or with only the basal pair of lateral veins prominent and the other lateral veins 

weak and inconspicuous. 

23. Lamina trinervate; intercostal venation prominent beneath; Amazon Basin .... 40. C. trinervia. 

23. Lamina with only the basal pair of lateral veins prominent and the other lateral veins weak 

and inconspicuous; intercostal venation beneath; Bolivia (La Paz), Ecuador (Pastaza), and 

Colombia (Antioquia). ..............................................9. C. cinnamomifolia. 

22. Lamina pinnately veined. 

24. Lamina above with white arachnoid hairs, at least (sub)persistent on the midrib; eastern Brazil. 

25. Stipules ca. 0.5 cm long ............................................... 13. C. curranii. 

25. Stipules 1-2.5 cm long ................................................ 17. C. floccosa. 

24. Lamina above glabrous (sometimes except for the base of the midrib) or scabrous to scabridulous 

with short, rigid hairs; not in eastern Brazil. 

26. Lamina scabrous to scabridulous above. 

27. Stipules ca. 6 cm long; lamina beneath with arachnoid hairs; basal lateral veins branched; 

Peru (Loreto) and Ecuador (Napo). ........................ 22. C. longepedunculata. 

27. Stipules up to 3.5 cm long, mostly less than 1 cm long; lamina beneath lacking 

arachnoid hairs, or if present then the basal lateral veins unbranched.

Coussapoa 19 

28. Leafy twigs (rather) abruptly thickened and distinctly hollow; widespread .... 

............................................................ 4. C asp erifolia. 

28. Leafy twigs gradually thickened and solid. 

29. Lateral veins 6-13 pairs; lamina subcoriaceous; southern Amazon Basin. ... 

............................... ................ ........... 37. C. scabra. 

29. Lateral veins 6-7 pairs; lamina coriaceous; French Guiana and Brazil (Para). 

........................................................ 21. C. leprieurii. 

26. Lamina smooth and glabrous above. 

30. Basal lateral veins unbranched (or sometimes with 1 or 2 branches). 

31. Lamina beneath with domatium-like cavities in the axils of the lateral veins; 

interfloral bracts large and peltate; western Ecuador and Colombia. .......... 

....................................... ...................... 10 . C . contorta. 

31. Lamina beneath without domatium-like cavities; interfloral bracts relatively small 

and spathulate to subpeltate. 

32. Stipules 0.5-1 cm long. 

33. Lateral veins 2-3(-4) pairs. 

34. Apex of the lamina acute; Central America. ...... 18. C. glaberrima. 

34. Apex of the lamina rounded to obtuse; Surinam, French Guiana, 

and Brazil (Amapa and Para). .................... 1. C. angustifolia. 

33. Lateral veins at least 4 pairs. 

35. Young leaves with arachnoid indumentum; lamina obovate to subobovate; 

lateral veins 7-10 pairs; terminal buds slender; eastern 

Brazil. .................... ..................... 13. C. curranii. 

35. Young leaves without arachnoid indumentum; lamina elliptic, or if 

(sub)obovate, then lateral veins 4-7 pairs and/or terminal buds 

swollen; not in eastern Brazil. 

36. Terminal buds swollen; Upper Amazon Basin and Colombia. 

........................................ 35. C. parvifolia. 

36. Terminal buds slender. 

37. Lateral veins 4-7 pairs. 

38. Hairs on the main veins beneath of equal length; Lower 

Amazon Basin, Surinam, and French Guiana. ... 

.............. ...................... 20. C. latifolia. 

38. Hairs on the main veins beneath of different lengths; 

eastern Venezuela, Guyana, and Brazil (Amazonas, 

Roraima). ......................... 43. C. viridifolia. 

37. Lateral veins 7-11 pairs; Upper Amazon Basin and Colombia. 

................................ 35. C. parvifolia. 



40. Lamina glabrous; Central America. .............. 18. C. glaberrima. 

40. Lamina beneath with (partly arachnoid) indumentum; Venezuela. 

........................................ 3. C. argentea. 


41. Lamina beneath with dense indumentum. 

42. Arachnoid hairs present on the lamina beneath, at least in young 

leaves. 

43. Arachnoid hairs brownish. 

44. Lateral veins 10-14 pairs; pistillate inflorescences unicapitate, 

head ellipsoid to clavate; staminate flower 

heads 4-8 mm in diameter; Andean Colombia and 

Ecuador (1400-2000 m). ............... 14. C. duquei. 

44. Lateral veins at most 10(-11); pistillate inflorescences 

pluricapitate, or if unicapitate then the heads globose; 

staminate flower heads 1-3 mm in diameter. 

45. Lateral veins 4-5 pairs; lamina oblong; French 

Guiana and Brazil (Amapa). ..... 16. C. ferruginea. 

45. Lateral veins (4-)5-8(-11) pairs; lamina mostly 

elliptic to ovate. 

46. Hairs on the smaller veins beneath appressed; 

Guianas ................ 26. C. microcephala. 

46. Hairs on the smaller veins beneath (at least 

partly) patent. 

47. Stamens slightly longer than the peri-


anth; pistillate flower heads 1-3; eastern 

Brazil. ............... 33. C. pachyphylla. 

47. Stamens distinctly longer than the perianth; 

pistillate flower heads 3-7; Venezuela 

(Bolivar), Brazil (Roraima), Amazonian 

Bolivia and Ecuador, and 

Panama. ............ 11. C. crassivenosa. 

43. Arachnoid hairs whitish. 

48. Lamina mostly ovate; lateral veins 9-16 pairs; stamen 

1; peduncle of the pistillate inflorescence 2-4 mm thick; 

Amazon Basin. .................... 39. C. tessmannii. 

48. Lamina mostly elliptic to oblong; lateral veins 5-12 

pairs; stamens 3; peduncle of the pistillate inflorescence 

ca. 1 mm thick. 

49. Stipules 0.5-1.5(-2) cm long; heads of staminate 

inflorescence many, 3-4 mm in diameter; heads 

of pistillate inflorescence 2-5; perianth glabrous; 

Amazon Basin. .................... 38. C. sprucei. 

49. Stipules 1-5 cm long; heads of staminate inflorescence 

3-10, 4-6 mm in diameter; heads of pistillate 

inflorescence 1(-3); perianth minutely puberulous; 

Central America to Mexico.......... 

............................. 30. C. oligocephala. 

42. Arachnoid hairs lacking on the lamina beneath, or if present 

then sparse and (soon) disappearing. 

50. Lateral veins 5-8 pairs; eastern Brazil. .. 33. C. pachyphylla. 

50. Lateral veins 8-12 pairs; not in eastern Brazil. 

51. Margin of the lamina revolute at the base; Upper Amazon 

Basin and Colombia. ........... 35. C. parvifolia. 

51. Margin of the lamina not revolute at the base. 

52. Petiole 1-3 cm long; stipules 1.5-2.5 cm long; 

pistillate inflorescences unicapitate, the peduncle 

0.5-1 cm long; Colombia (Choc6). 7. C. chocoensis. 

52. Petiole 3-9 cm long; stipules 2-6 or 3-20 cm long; 

pistillate inflorescences pluricapitate, or if unicapitate 

then the peduncle at least 1 cm long. 

53. Heads of pistillate inflorescence 6-10; stamens 

3; Pacific coastal region of Ecuador and 

Colombia. ................... 19. C. herthae. 

53. Heads of pistillate inflorescence 1(-2); stamens 

2 or unknown. 

54. Lamina broadly elliptic to obovate, apex 

shortly acuminate; pistillate flower heads 

ca. 4 mm, in fruit up to 12 mm in diameter; 

Amazonian Peru ........... 

...................... 24. C. manuensis. 

54. Lamina mostly ovate to subovate, apex 

acute to obtuse; pistillate flower heads 

ca. 5-10 mm, in fruit up to 40 mm in 

diameter; widespread....... 42. C. villosa. 

41. Lamina beneath with sparse indumentum. 

53. Margin of the lamina distinctly revolute towards the base. 

56. Stipules 1-7 cm long; terminal buds slender; eastern Brazil 

...................................... 25. C. m icrocarpa. 

56. Stipules up to 1.3 cm long; terminal buds usually swollen; 

Upper Amazon Basin and Colombia. ..... 35. C. parvifolia. 

55. Margin of the lamina plane or entirely revolute. 

57. Leafy twigs (rather) abruptly thickened and with wide cavities; 

stamen 1; widespread ............... 4. C. asperifolia. 

57. Leafy twigs gradually thickened and solid or with narrow 

cavities. 

58. Stipules glabrous or puberulous. 

59. Basal lateral veins reaching the margin above the


middle of the lamina; base of the lamina acute; 

pistillate inflorescences pluricapitate; Panama. .. 

............................... 15. C. echinata. 

59. Basal lateral veins mostly reaching the margin 

below the middle of the lamina; base of the lamina 

mostly obtuse to (sub)cordate. 

60. Stamens 3; pistillate inflorescences unicapitate; 

northern Central America to Mexico. 

............................. 36. C. purpusii. 

60. Stamens 2; pistillate inflorescences pluricapitate; 

South America. 

61. Hairs on the main veins beneath equal 

in length and arachnoid hairs lacking; 

Surinam, French Guiana, and Brazil 

(Amapa, Para, and Amazonas). ...... 

......................... 20. C. latifolia. 

61. Hairs on the main veins beneath of different 

lengths and/or on the main veins 

and margin subpersistent arachnoid 

hairs; Guianas....... 26. C. microcephala. 

58. Stipules (sub)sericeous, (sub)hirsute, (sub)villous, etc. 

62. Stamens 3; pistillate flower heads 6-10, 5-9 mm 

in diameter; Pacific coastal region of Ecuador and 

Colombia. 

63. Petiole 5-10 cm long, or if shorter, then stamens 

3, pistillate flower heads 6-10, 5-9 mm 

in diameter; Pacific coastal region of Colombia 

and Ecuador. 

64. Stipules 2-6 cm long. ..... 19. C. herthae. 

64. Stipules 1.5-2 cm long. .. 44. C. fulvescens. 

63. Petiole up to 5 cm long; stamens 2; flower 

heads 1-5; not in Pacific coastal region. 

62. Stamens 2; pistillate flower heads 1-3, or if more 

than 3 then 2-5 mm in diameter. 

64. Stamens slightly longerthan the perianth; pistillate 

flower heads 1-3; eastern Brazil ...... 

........................ 33. C. pachyphylla. 

64. Stamens distinctly longer than the perianth; 

pistillate flower heads (1-)3-15; Guianas. .. 

........................ 26. C. microcephala. 


65. Lamina beneath with (rather) dense, (rather) conspicuous and (sub)persistent 

arachnoid indumentum. 

66. Arachnoid hairs on the lamina brownish. 

67. Lateral veins 4-8(-11) pairs; pistillate inflorescences mostly pluricapitate; 

staminate flower heads ca. 2-3 mm in diameter. 

68. Hairs on the smaller veins beneath (at least partly) patent. 

69. Stamens slightly longer than the perianth; pistillate flower heads 

1-3; eastern Brazil. ....................... 33. C. pachyphylla. 

69. Stamens distinctly longer than the perianth; pistillate flower 

heads 3-7; Venezuela (Bolivar), Brazil (Roraima), Amazonian 

Bolivia and Ecuador, and Panamf. ......... 11. C. crassivenosa. 

68. Hairs on the smaller veins beneath appressed; Guianas.......... 

............................................ 26. C. m icrocephala. 

67. Lateral veins (7-)10-24 pairs; pistillate inflorescences usually unicapitate; 

staminate flower heads 4-10 mm in diameter. 

70. Common peduncle of the staminate inflorescence 1-2 cm long, that 

of the pistillate inflorescence 0.5-0.8 cm long; pistillate heads ellipsoid 

to clavate; Andean Ecuador and Colombia (1400-2000 m). .. 

.................................................. 14. C. duquei. 

70. Common peduncle of the staminate inflorescence 2-6 cm long, that 

of the pistillate inflorescence (1-)2-13 cm long; pistillate flower 

heads (sub)globose; widespread. ..................... 42. C. villosa.


66. Arachnoid hairs on the lamina whitish. 

71. Lateral veins ca. 10 or 9-24 pairs; lamina mostly longer than 15 cm; 

stipules mostly longer than 2 cm. 

72. Lamina beneath with dense brown hairs on the main veins; common 

peduncle of the staminate inflorescence ca. 9 cm long; Amazonian 

Peru and Ecuador. ....................... 22. C. longepedunculata. 

72. Lamina beneath with whitish hairs on the main veins; peduncle of 

the staminate inflorescence up to 6 cm long. 

73. Stamen 1; pistillate inflorescences pluricapitate and common 

peduncle 2-3 cm long; Amazon Basin. ....... 39. C. tessmannii. 

73. Stamens 2 or 3; pistillate inflorescences unicapitate, or if 2-3capitate 

then the common peduncle longer than 3 cm. 

74. Stamens 3; pistillate flower heads ellipsoid to clavate; peduncle 

of the pistillate inflorescence 1-2 cm long; Central 

America. .......................... 29. C. nymphaeifolia. 

74. Stamens 2; peduncle of the pistillate inflorescence (1-)2- 

13 cm long; pistillate flower heads (sub)globose; widespread. 

................................... 42. C. villosa. 

71. Lateral veins 4-10(-11) pairs; lamina mostly shorter than 15 cm. 

75. Hairs on the smaller veins beneath appressed; Guianas.......... 

............................................ 26. C. m icrocephala. 

75. Hairs on the smaller veins beneath (at least partly) patent. 

76. Stipules up to 2 cm long. 

77. Hairs on the stipules predominantly weak and crinkled; 

stamens 2; pistillate flower heads 1-3; eastern Brazil. ... 

..................................... 33. C. pachyphylla. 

77. Hairs on the stipules predominantly stiff and straight; stamens 

3; pistillate flower heads 2-6; Amazonian Brazil. .. 

.......................................... 34. C. sprucei. 

76. Stipules 3-20 cm long; widespread. .............. 42. C. villosa. 

65. Lamina beneath without arachnoid indumentum, or if present then sparse and 

soon disappearing. 

78. Leafy twigs (rather) abruptly thickened and with wide cavities; stamen 1; 

widespread. ............................................. 4. C. asperifolia. 

78. Leafy twigs gradually thickened, solid or with narrow cavities; stamens 2 

or 3. 


80. Lamina almost as long as broad; pair of main basal veins reaching 

the margin far above the middle of the lamina; most lateral veins 

branched; Pacific coastal region of Ecuador and Colombia. ...... 

............................................... 41. C. vannifolia. 

80. Lamina mostly distinctly longer than broad, or if almost as long as 

broad then only the basal pair of lateral veins branched. 

81. Hairs on the smaller veins beneath appressed; Guianas. ..... 

........................................ 26. C. m icrocephala. 

81. Hairs on the smaller veins beneath at least partly patent. 

82. Stamen 1; pistillate inflorescences pluricapitate and the 

common peduncle 2-3 cm long and 2-4 cm thick, perianth 

of the pistillate flower (almost) glabrous; Amazon Basin. 

...................................... 39. C. tessm annii. 

82. Stamens 2 or 3; pistillate inflorescences unicapitate, or if 

pluricapitate then the common peduncle longer than 3 cm, 

less than 2 mm thick; and/or the perianth of the pistillate 

flower densely puberulous. 

83. Stamens 3; pistillate inflorescence with 6-10 heads, 5- 

9 (in fruit up to 15) mm in diameter and the common 

peduncle 3-6 mm long; Pacific coastal region of Ecuador 

and Colombia. ................. 19. C. herthae. 

83. Stamens 2; pistillate inflorescences with 1-3 heads, or 

if more then the heads 3-5 mm in diameter and the 

common peduncle 1-3 cm long. 

84. Stamens slightly longer than the perianth; pistil-


late flower heads 1-3; perianth of the pistillate 

flower glabrous; eastern Brazil. .. 33. C. pachyphylla. 

84. Stamens distinctly longer than the perianth; pistillate 

flower heads 3-7, or if less then the perianth 

of the pistillate flower densely puberulous. 

85. Staminate flower heads 2-3 mm in diameter; 

pistillate flower heads 3-7, and 2-5 mm in 

diameter; Venezuela (Bolivar), Brazil (Roraima), 

Amazonian Bolivia and Ecuador, and 

Panama. ................. 1. C. crassivenosa. 

85. Staminate flower heads 5-10 mm in diameter; 

pistillate flower heads 1(-3) and 5-10 mm 

in diameter; widespread. ....... 42. C. villosa. 


86. Stamen 1; pistillate inflorescences pluricapitate and the common 

peduncle 2-3 cm long and 2-4 cm thick; perianth of the pistillate 

flower (almost) glabrous; Amazon Basin. ......... 39. C. tessmannii. 

86. Stamens 2 or 3; pistillate inflorescences unicapitate, or if pluricapitate 

then the common peduncle longer than 3 cm, less than 2 mm 

thick, and/or the perianth of the pistillate flower densely puberulous. 

87. Stamens 3; pistillate flower heads 6-10; Pacific coastal region 

of Ecuador and Colombia. ..................... 19. C. herthae. 

87. Stamens 2; pistillate flower heads 1(-3); widespread ......... 

........................................ 42. C. villosa. 

1. Key to the Coussapoa Species in Central America and Mexico 


2. Lamina beneath glabrous or with sparse indumentum. 

3. Lateral veins 3-6 pairs, the basal pair reaching the margin above or at the middle of the lamina. 

.. ............... ....................................................... 34. C. parviceps. 

3. Lateral veins 7-13 pairs, the basal pair reaching the margin below the middle of the lamina. 

4. Stipules 1-2 cm long; lamina 5-13 x 4-7 cm. ........................... 23. C. macerrima. 

4. Stipules 2-3 cm long; lamina 13-20 x 9-13 cm. ................. See under 32. C. ovalifolia. 


5. Intercostal venation prominent beneath; stipules with brownish indumentum (unicellular hairs 

only). ........................................................ .. .... 39. C. nym phaeifolia. 

5. Intercostal venation plane beneath; stipules with whitish indumentum (unicellular hairs only!). 

........................................................................... 6. C brevipes. 

1. Interfloral bracts present. 

6. Lamina beneath with dense indumentum, partly arachnoid indumentum. 

7. Arachnoid indumentum on the lamina brownish. .......................... 11. C. crassivenosa. 

7. Arachnoid indumentum on the lamina whitish. 

8. Basal pair of lateral veins unbranched. ................................. 30. C. oligocephala. 

8. Basal pair of lateral veins branched. 

9. Stamens 3; (common) peduncle of the pistillate inflorescence 1-2 cm long, pistillate flower 

heads ellipsoid to obovoid. ...................................... 39. C. nymphaeifolia. 

9. Stamens 2; (common) peduncle of the pistillate inflorescence (1-)2-13 cm long, pistillate 

flower heads (sub)globose. ............................................. 42. C. villosa. 

6. Lamina beneath glabrous or with sparse indumentum, arachnoid indumentum lacking. 

10. All or most lateral veins branched; leafy twigs more or less abruptly thickened, with wide cavities. C 

......... . ..................................................... 

asperifolia. 

10. All or most lateral veins unbranched; leafy twigs gradually thickened, solid or with narrow 

cavities. 

11. Lateral veins 2-3 pairs. ............................................ 18. C. glaberrima. 


12. Basal pair of lateral veins unbranched; pistillate inflorescences unicapitate. ......... 

...................................... ............................ 36. C. purpusii. 

12. Basal pair of lateral veins (faintly) branched; pistillate inflorescences pluricapitate. .. 

................................................................. 15. C. echinata.


2. Key to the Coussapoa Species in Colombia 


2. Lateral veins 2-8 pairs, basal pair reaching the margin above (or at) the middle of the lamina. 

3. Basal pair of lateral veins unbranched; stamen 1; pistillate flowers free. ........ 31. C. orthoneura. 

3. Basal pair of lateral veins branched; stamens 3; pistillate flowers cohere. .........34. C. parviceps. 

2. Lateral veins at least 8 pairs, or if less then basal pair reaching the margin below the middle of the 

lamina. 

4. Lamina in the areoles beneath glabrous or with sparse hairs. 

5. Stipules 2-3 cm long . .................................................... 46. C. valaria. 

5. Stipules 4-13 cm long. 

6. Base of the lamina cordate; basal pair of lateral veins reaching the margin above or at the 

middle of the lamina . .............................................. 5. C. batavorum. 

6. Base of the lamina (sub)obtuse or rounded to truncate; basal pair of lateral veins reaching 

the margin below the middle of the lamina ........................... 27. C. napoensis. 

4. Lamina in the areoles beneath with dense, minute hairs. 

7. Indumentum of the petiole consisting of dense, minute hairs and distinctly longer, yellowish 

hairs . .................................................................... C. 28. nitida. 

7. Indumentum of the petiole consisting of short hairs, not distinctly different in length. 

8. Indumentum of the leafy twigs and stipules (unicellular hairs only) brown(ish); petiole 5- 

7 m m thick. ..................................................... 8. C. cinnam omea. 

8. Indumentum of the leafy twigs and stipules (unicellular hairs only) white or yellowish; 

petiole 2-4 mm thick. 

9. Intercostal venation almost plane; pistillate inflorescences unbranched ... 6. C. brevipes. 

9. Intercostal venation prominent; pistillate inflorescences branched .... 45. C. tolimensis. 


10. Lamina trinervate ........................................................... 40. C. trinervia. 


11. Lamina beneath with arachnoid indumentum. 

12. Arachnoid indumentum on the lamina beneath brown(ish). 

13. Lateral veins 4-8(-11) pairs; pistillate inflorescences pluricapitate; staminate flower 

heads ca. 2-3 mm in diameter ................... .............. 11. C. crassivenosa. 

13. Lateral veins (7-)10-24 pairs; pistillate inflorescences (usually) unicapitate; staminate 

flower heads 4-10 mm in diameter. 

14. Common peduncle of the staminate inflorescence 1-2 cm long, that of the pistillate 

inflorescence 0.5-0.8 cm long; pistillate flower heads ellipsoid to clavate. ..... 

........................... ................................... 14. C . duquei. 


inflorescence (1-)2-13 cm long; pistillate flower heads (sub)globose. .. 42. C. villosa. 

12. Arachnoid indumentum on the lamina beneath whitish. 

15. Stamen 1; pistillate inflorescence pluricapitate and the common peduncle 2-3 cm long, 

2-4 mm thick; perianth of the pistillate flower almost glabrous ...... 39. C. tessmannii. 

15. Stamens 2 or 3; pistillate inflorescence unicapitate, or if pluricapitate then the common 

peduncle longer than 3 cm, less than 2 mm thick, and/or the perianth of the pistillate 

flower densely puberulous. 

16. Stamens 3; pistillate flower heads 6-10. ......................... 19. C. herthae. 

16. Stamens 2; pistillate flower heads 1(-3). ........................ 42. C. villosa. 

11. Lamina beneath without arachnoid indumentum. 

17. Lamina beneath with domatium-like cavities in the axils of the lateral veins; interfloral 

bracts large and peltate .................................. ............. 10. C. contorta. 

17. Lamina beneath without domatium-like cavities; interfloral bracts relatively small and 

spathulate to subpeltate. 

18. Stipules glabrous or sparsely puberulous ............................ 34. C. parviceps. 

18. Stipules with + dense indumentum. 

19. Basal lateral veins distinctly branched. 

20. Lamina about as long as broad; lateral veins 5-9 pairs. ..... 41. C. vannifolia. 

20. Lamina distinctly longer than broad; lateral veins 8-12 pairs.... 19. C. herthae. 

19. Basal lateral veins unbranched (or sometimes with 1-2 branches). 

21. Lateral veins 5-6 pairs; intercostal venation inconspicuous ............. 

................................................... 9. C. cinnam om ifolia. 

21. Lateral veins 7-12 pairs; intercostal venation conspicuous. 

22. Stipules 0.3-1.3 cm long. ............................ 35. C. parvifolia. 

22. Stipules 1.5-2.5 cm long.


23. Lateral veins 10-12 pairs; petiole 1-3 cm long; pistillate inflores- 

cences unbranched. .............................. 7. C. chocoensis. 

23. Lateral veins 5-10 pairs; petiole 5-10 cm long; pistillate inflores- 

cences branched. ............................... 44. C. fulvescens. 

3. Key to the Coussapoa Species in Venezuela 


2. Stipules 0.5-1 cm long; stamens 2; pistillate flower heads 1-3. .................... 

2. Stipules (0.5-)1-7 cm long; stamen 1; pistillate flower heads 2-10. ............... 


3. Lamina trinervate ............................................................. 

43. C. viridifolia. 

31. C. orthoneura. 

40. C. trinervia. 



5. Arachnoid indumentum on the lamina brownish . ...................... 


6. Lateral veins 2-3(-4) pairs ........ ..................................... 

6. Lateral veins 4-24 pairs . .................. ........................... 


11. C. crassivenosa. 

3. C. argentea. 

42. C. villosa. 

7. Leafy twigs (rather) abruptly thickened with wide cavities; stamen 1 ......... 4. C. asperifolia. 

7. Leafy twigs gradually thickened solid or with narrow cavities. 

8. Lamina with patent hairs on the venation beneath; stipules 1-5 cm long. ............. 

....................................................11. C. crassivenosa. 

8. Lamina with appressed hairs on the venation beneath; stipules 0.3-1(-1.3) cm long. 

9. Lateral veins 7-11 pairs; terminal buds usually swollen; margin of the lamina ? revolute 

at the base. ....................................... .... 35. C. parvifolia. 

9. Lateral veins 4-7 pairs; terminal buds slender; margin of the lamina not revolute at 

the base ........................................................ 43. C. viridifolia. 

4. Key to the Coussapoa Species in Guyana, Surinam, and French Guiana 

1. Lamina above scabrous. 

2. Basal lateral veins reaching the margin above the middle of the lamina. ............4. C. asperifolia. 

2. Basal lateral veins reaching the margin below the middle of the lamina ............. 21. C. leprieurii. 

1. Lamina above smooth. 


4. Arachnoid indumentum on the lamina brownish. 

5. Lamina beneath with patent hairs on the venation. 

6. Lateral veins 4-5 pairs; lamina oblong. ............................... 16. C. ferruginea. 

6. Lateral veins (4-)5-8 pairs; lamina mostly elliptic to ovate to obovate or to suborbicular. 

..........................1.............. ...... . . .. 1 . C. crassivenosa. 

5. Lamina beneath with appressed hairs on the venation .................. 26. C. microcephala. 


7. Lateral veins 2-3(-4)pairs. ......................... ... .... ................. 3.C. argentea. 

7. Lateral veins 4-11 pairs. .......................... .................. 26. C. microcephala. 


8. Lateral veins 2-4 pairs. .................. ..... ........ .................... 1. C. angustifolia. 


9. Leafy twigs (rather) abruptly thickened with wide cavities; stamen 1. .........4. C. asperifolia. 

9. Leafy twigs gradually thickened and solid or with narrow cavities. 

10. Lateral veins 7-11 pairs; terminal buds usually swollen. .............. 35. C. parvifolia. 

10. Lateral veins 4-7 pairs; terminal buds slender. 

11. Hairs on the lamina beneath of about equal length .................. 20. C. latifolia. 

11. Hairs on the lamina beneath of different lengths ... ................ 43. C. viridifolia. 

5. Key to the Coussapoa Species in Ecuador 


2. Basal pair of lateral veins unbranched (or occasionally a single branch). 

3. Lateral veins 2-7 pairs, the basal pair reaching the margin above to just below the middle of the 

lam ina. ................................................................. 3 1. C . orthoneura. 

25


3. Lateral veins (5-)7-21 pairs, the basal pair reaching the margin (far) below the middle of the 

lamina. 

4. Stipules 1-4 cm long; apex of the lamina mostly acute to shortly acuminate. . 32. C. ovalifolia. 

4. Stipules 4-13 cm long; apex of the lamina mostly obtuse. .................. 27. C. napoensis. 

2. Basal pair of lateral veins branched. 

5. Stipules glabrous or sparsely puberulous .....................................34. C. parviceps. 

5. Stipules subvelutinous, subhirsute, or subsericeous. 

6. Stipules 1-4 cm long; apex of the lamina mostly acute to shortly acuminate. .. 32. C. ovalifolia. 

6. Stipules 4-13 cm long; apex of the lamina mostly obtuse. .................. 27. C. napoensis. 


7. Lamina trinervate or only the basal pair of lateral veins strong and the other lateral veins weak and 

inconspicuous. 

8. Lamina trinervate; intercostal venation prominent beneath. .... ............ 40. C. trinervia. 

8. Lamina with only the basal pair of lateral veins strong and the other lateral veins weak and 

inconspicuous; intercostal venation plane beneath. ............... ........9. C. cinnamomifolia. 


9. Lamina beneath with arachnoid indument. 

10. Arachnoid indumentum on the lamina beneath brown(ish). 

11. Lateral veins 4-8(1 1) pairs; pistillate inflorescences pluricapitate; staminate flower heads 

ca. 2-3 mm in diameter. ........................................ 11. C. crassivenosa. 

11. Lateral veins (7-)10-24 pairs; pistillate inflorescences (usually unicapitate; staminate 

flower heads 4-10 mm in diameter. 


inflorescence 0.5-0.8 cm long; pistillate flower heads ellipsoid to clavate. ....... 

...................................... . ........................ 14. . C . duquei. 


inflorescence (1)2-13 cm long; pistillate flower heads (sub)globose .... 42. C. villosa. 

10. Arachnoid indumentum on the lamina beneath whitish. 

13. Stamens 2; pistillate flower heads 1(-3). ................................. 42. C. villosa 

13. Stamens 3; pistillate flower heads 3-10. 

14. Common peduncle 8-20 cm long; base of the lamina cordate to rounded ........ 

...................................................... 22. C longepedunculata. 

14. Common peduncle up to 7 cm long; base of the lamina usually obtuse to rounded. 

............................................................... 19. C. herthae. 


15. Lamina beneath with domatium-like cavities in the axils of the lateral veins; interfloral bracts 

large and peltate. ...................................................... 10. C. contorta. 

15. Lamina beneath without domatium-like cavities; interfloral bracts relatively small and spathulate 

to subpeltate. 

16. All lateral veins usually branched; pistillate inflorescences usually unicapitate; stamen 

1. ......... ..................................... . .. 1. .. C. asperifolia. 

16. Only the basal pair of lateral veins usually branched; pistillate inflorescences pluricapitate; 

stamens 3. 

17. Stipules glabrous or sparsely puberulous ............................... 34. C. parviceps. 

17. Stipules with dense indumentum. 

18. Lamina about as long as broad; lateral veins 5-9 pairs. ....... 41. C. vannifolia. 

18. Lamina distinctly longer than broad; lateral veins 8-12 pairs. 

19. Common peduncle 8-20 cm long; base of the lamina cordate to rounded. 

..................................... 22. C. longepedunculata. 

19. Common peduncle up to 7 cm long; base of the lamina usually obtuse to 

rounded. .............................................. 19. C. herthae. 

6. Key to the Coussapoa Species in Peru and Bolivia 


2. Basal lateral veins unbranched (or occasionally with a single branch). 

3. Lateral veins 7-21 pairs and the lamina beneath (almost) glabrous in the areoles. .. 32. C. ovalifolia. 

3. Lateral veins 2-9 pairs and/or the lamina beneath with dense, minute hairs in the areoles. 

4. Basal lateral veins reaching the margin above or just below the middle of the lamina; stipules 

mostly without or with sparse long yellow hairs; branches of the pistillate inflorescence slightly 

broadened towards the flower heads ..................................... 31. C. orthoneura. 

4. Basal lateral veins reaching the margin below to just above the middle of the lamina; stipules


with dense, long, yellow hairs; branches of the pistillate inflorescence strongly broadened 

towards the flower heads ................................................ 12. C. cupularis. 


5. Lateral veins 5-9 pairs; lamina usually elliptic. ........................... See 12. C. cupularis. 

5. Lateral veins 8-15 or 14-22; lamina usually ovate. 

6. Indumentum of the petiole consisting of dense minute hairs and distinctly longer, yellowish 

hairs; lateral veins 8-15 pairs; peduncle of the pistillate inflorescence 1-2 mm thick; Amazon 

Basin. ........................... .................................... 28. C. nitida. 

6. Indumentum of the petiole consisting of short hairs, not differing distinctly in length; lateral 

veins 14-22 pairs; peduncle of the pistillate inflorescence 6-10(-15) mm thick; Colombia 

(Am azonas). ........................................................ 8. C. cinnam om ea. 


7. Lamina trinervate or only the basal pair of lateral veins strong and the other lateral veins weak and 

inconspicuous. 

8. Lamina trinervate; intercostal venation prominent beneath. .................... 40. C. trinervia. 

8. Lamina with only the basal pair of lateral veins strong and the other lateral veins weak and 

inconspicuous; intercostal venation plane beneath. ....................... 9. C. cinnamomifolia. 


9. Lamina beneath glabrous or with sparse indumentum. 

10. Basal lateral veins unbranched; stipules up to 1(-1.3) cm long; margin of the lamina + 

revolute at the base. .................................................. 35. C. parvifolia. 

10. Basal and/or other lateral veins usually branched; stipules up to 3.5 cm long; margin of the 

lamina not revolute at the base. ......................................... 4. C. asperifolia. 


11. Lamina beneath without arachnoid indumentum and basal pair of veins unbranched. .... 

.................................................................... 24. C. manuensis. 

11. Lamina beneath with arachnoid indumentum and basal pair of lateral veins branched. 

12. Stamens 3; common peduncle 8-20 cm long; pistillate flower heads 3-10 ........... 

............... ........................................... 22. C. longepedunculata. 

12. Stamens 2 or 1; common peduncle up to 6 cm long, or if longer in pistillate inflorescences 

then a single flower head. 

13. Stamens 2; pistillate inflorescences unicapitate, or if pluricapitate then the perianth 

of the pistillate flower densely puberulous. ......................... 42. C. villosa. 

13. Stamen 1; pistillate inflorescences pluricapitate and perianth of the pistillate flower 

alm ost glabrous. ........................................ ... 39. C. tessmannii. 

7. Key to the Coussapoa Species in Amazonian Brazil 



3. Stipules 0.5-1 cm long; stamens 2; pistillate flower heads 1-3 and common peduncle or branches 

not thickened below these heads. ........................................... 43. C. viridifolia. 

3. Stipules usually longer than 2 cm, if shorter than 1 cm then stamen 1 or pistillate flower heads 

2-10 and the branches of the inflorescence more or less thickened below these heads. 

4. Intercostal venation prominent beneath; lamina beneath and inflorescences with dense, white 

arachnoid indumentum; Lower Amazon Basin. .......................... 2. C. arachnoidea. 

4. Intercostal venation plane beneath; lamina and inflorescences without arachnoid indumentum, 

or if present then mostly soon disappearing; Upper Amazon Basin. 

5. Basal lateral veins reaching the margin above to just below the middle of the lamina; 

stipules mostly without or with sparse, long, yellow hairs; branches of the pistillate inflorescence 

slightly thickened below the flower heads. .................... 31. C. orthoneura. 

5. Basal lateral veins reaching the margin below to just above the middle of the lamina; 

stipules with dense, long, yellow hairs; branches of the pistillate inflorescence strongly 

thickened below the flower heads. .................................... 12. C. cupularis. 

2. Lateral veins (7-)9-22 pairs. 

6. Basal lateral veins (mostly unbranched; lamina beneath in the areoles glabrous or sparsely puberulous. 

................................................................. 32. C. ovalifolia. 

6. Basal lateral veins branched; lamina beneath in the areoles with dense, minute hairs. 

7. Petiole 2-4 mm thick, with dense, minute hairs and sparse, much longer, yellowish hairs. 

......................................................................... 28. C . m itida. 

7. Petiole 5-7 mm thick, with short hairs about the same in length. ......... 8. C. cinnamomea. 

27



8. Lamina trinervate ............................................................. 40. C. trinervia. 


9. Lamina scabrous above. 

10. Lateral veins unbranched. ................................................ 37. C. scabra. 

10. All or most lateral veins branched .......................................4. C. asperifolia. 

9. Lamina smooth above. 

11. Lam ina trinervate. ..................................................... 40. C. trinervia. 


12. Lamina beneath with dense, partly arachnoid indumentum. 

13. Arachnoid indumentum on the lamina brownish. 

14. Lateral veins 4-5 pairs; lamina oblong. ..................... 16. C. ferruginea. 

14. Lateral veins (4-)5-8 pairs; lamina mostly elliptic to ovate to obovate or to 

suborbicular. ........................................... 11. C. crassivenosa. 


15. Stamens 2; pistillate inflorescences (usually) unicapitate; perianth of the pistillate 

flower densely puberulous. .............................. 42. C. villosa. 

15. Stamen 1; pistillate inflorescences pluricapitate; perianth of the pistillate flower 

almost glabrous. ........................................ 

39. C. tessmannii. 

12. Lamina beneath glabrous or with sparse indumentum and arachnoid indumentum lacking. 

16. Lateral veins 2-4 pairs. .......................................1. C. angustifolia. 


17. Lateral veins 7-11 pairs; terminal buds usually swollen. ...... 35. C. parvifolia. 

17. Lateral veins 4-7 pairs; terminal buds slender. 

18. Leafy twigs (rather) abruptly thickened and with wide cavities; stamen 1. 

...................................................... . C. asperifolia. 

18. Leafy twigs gradually thickened and solid or with narrow cavities. 

19. Hairs on the lamina beneath distinctly different in lengths. ........ 

................................................. 43. C. viridifolia. 

19. Hairs on the lamina beneath about equal in length ..... 20. C. latifolia. 

8. Key to the Coussapoa Species in Eastern Brazil 

(Rio Grande do Sul to Pernambuco) 

1. Stipules up to 1 cm long. ........................................... 13. C. curranii. 


2. Lamina above with subpersistent, white, arachnoid indumentum. ................... 17. C. floccosa. 

2. Lamina above glabrous. 

3. Stamens much longer than the perianth; pistillate inflorescences mostly unicapitate, (common) 

peduncle 1-2.5 cm long; stipules up to 7 cm long; leaf margin usually revolute towards the base. 

........................................................................ 25. C . m icrocarp a. 

3. Stamens as long as the perianth; pistillate inflorescences mostly pluricapitate, common peduncle 

2.5-4 cm long; stipules up to 2 cm long; leaf margin entirely revolute or plane ............... 

....................................................................... 33. C . p achyp hylla. 

1. Coussapoa angustifolia Aublet, Hist. pl. Gui- 

ane 2: 956, t. 363. 1775; Miquel, Fl. bras. 4(1): 

138. 1853; Berg. Fl. Suriname 5(1): 283. 1975. 

Type. French Guiana. Without locality, -, 

Aublet s.n. (BM, leaf fragment). Fig. 10. 

Shrub or tree, mostly hemi-epiphytic, up to 30 

m tall. Leafy twigs 3-8 mm thick, glabrous or 

sparsely, minutely puberulous, lenticels numer- 

ous and conspicuous. Lamina coriaceous ob- 

ovate or subobovate (to oblanceolate or ellip- 

tic to oblong), 2-11 x 1-6 cm, apex obtuse to 

rounded or emarginate, base acute to cuneate (or 

to subobtuse), margin entire; upper surface glabrous, 

lower surface glabrous or sparsely appressed-puberulous; 

lateral veins 2-4 pairs, 

straight or curved, basal pair unbranched, reaching 

the margin far above the middle of the lamina; 

intercostal venation mostly plane; petiole 1- 

5 cm long, glabrous or sparsely appressed-puberulous; 

stipules 0.5-1 cm long, glabrous or 

sparsely appressed-puberulous. Staminate inflorescences 

poorly branched, heads 3-5, sometimes 

partly fused, globose, 5-8 mm diam.; common 

peduncle 0.5-1.5 cm long, minutely


FIG. 10. Coussapoa angustifolia: 1, leafy twig with pistillate inflorescences (Schulz 10344a); 2, staminate 

inflorescences (Pires et al. 50631). 

puberulous; perianth 1 mm high, glabrous; sta- 

men one, as long as the perianth. Pistillate inflo- 

rescences unbranched or rarely branched; heads 

1(-2), globose, ca. 5 mm, in fruit up to 20 mm 

diam.; (common) peduncle 0.3-3 cm long, mi- 

nutely puberulous; perianth 1(-2) mm high, gla- 

brous; fruiting perianth yellow to orange. Inter- 

floral bracts spathulate to subpeltate, apex 

ciliolate. 

Distribution (Fig. 6). Surinam, French Guiana, 

and Brazil (Amapa and Para); in upland and riverine 

forest. 

Specimens studied. SURINAM. Brownsberg, 6 Sep 

1915 (e), BW 772 (A, K, NY, U); Brokopondo Lake,

30 

10 Sep 1965 (st), Van Donselaar 2569 (U); Wilhelmina 

Mts., Zuid: River, 45 km above confluence with Lucie 

River, 25 Sep 1963 (9), Irwin et al. 57574 (B, G, MO, 

NY, OXF, R, S, U); Coppename River, Bitagron, 28 

Jul 1954 (9), Lindeman 6362 (F, U); Zanderij Savanna, 

28 Aug 1954 (st), Lindeman 6545 (U); Zanderij, 15 

Apr 1954 (st), Lindeman 6600 (MO, U); Saramacca 

River, Jacobkondre, 19 Jun 1944 (6), Maguire 23867 

(A, F, K, NY, U, US); Saramacca River, nr. Kwattahede, 

22 Jun 1944 (2), Maguire 23942 (A, F, K, MO, 

NY, P, RB, U); Wilhelmina Mts., S of Julianatop, 7 

Aug 1963 (8), Schulz 10344a (NY, U); Suriname River, 

Gansee, 8 Jul 1908 (9), Tresling 55 (U). 

FRENCH GUIANA. Without locality, - (st), Aublet 

s.n. (BM, type collection); Itany River, island in 

rapids, 30 Sep 1961 (9), BAFOG (=Graimleux) 7938 

(CAY, U); Maroni River, 15 Sep 1903 (9), Geay 3285 

(P), Oyapock River, nr. Grand Roche Fall, 17 Jul 1960 

(9), Maguire et al. 47049 (G, GH, LE, NY, OXF, U, 

US); nr. Cayenne, - (9), Martin s.n. (BM, BR, K); 

Maroni River, 1854 (9), Melinon 172 (LE, P); Camopi 

River, below Grand Tamouri Creek, 2 Feb 1968 (2), 

Oldeman et al. 207 (CAY, P, U, US); Sinnamary River, 

1.5 km above Courcibo River, 7 Aug 1967 (9), Oldeman 

B1167 (CAY); Kourou River, 2 km above Cr6ole, 

18 Sep 1967 (9), Oldeman B1324 (CAY, U); Oropu 

River, 1.5 km above Degrad Lalande, 16 Oct 1965 (9), 

Oldeman 1619 (CAY); Approuague River, 5 km above 

Maripa Creek, 3 Feb 1967 (9), Oldeman 2476 (CAY, 

P, U); Oyapock River, above Moutouci Falls, 15 May 

1970 (6), Oldeman B3243 (CAY, U); Sinnamary, road 

to St. Elie, 18 Jul 1977 (6), Sastre 5500 (P, U). 

BRAZIL. AMAPA: Rio Araguari, between 1055'N, 

51?51'W and 2?5'N, 51056'W, 1 Sep 1961 (9), Pires et 

al. 50631 (NY, U, US). PARA: Mun. Oriximina, road 

to Cachoeira Porteira, 18 Jun 1980 (9), Cid et al. 1053 

(U); Rio Paru do Oeste, Cachoeira Chuvisco, 17 Sep 

1980 (9), Cid et al. 2223 (U), Ilhas de Breves, Furo 

Macujubim, 16 Nov 1922 (9), Ducke (HJBR) 18461 

(RB); Rio Trombetas, Cachoeira Porteira, 8 Jun 1978 

(9), N. T. Silva et al. 4757 (U). 

This species, with few lateral veins in a mostly 

subobovate lamina, appears to be related to C. 

trinervia. In its leaf characters C. angustifolia is 

transitional to the trinervate state of the latter 

species. It also shows many similarities to the 

Central American C. glaberrima. 

2. Coussapoa arachnoidea Akkermans & C. C. 

Berg, Proc. Kon. Ned. Akad. Wetensch. Ser. 

C, 85(4): 442. 1982. Type. Brazil. Amapa: Rio 

Araguari, between 1?55'N, 51?59'W and 2?5'N, 

51?56'W, 1 Sep 1961 (2), Pires et al. 50656 

(holotype, IAN; isotypes, M, MG, NY, OXF, 

US, VEN). Fig. 11. 

Tree. hemi-epiphytic, up to 35 m tall. Leafy 

twigs 3-8 mm thick, with sparse, arachnoid hairs 

and yellowish, straight hairs on the scars of the 

Flora Neotropica 

stipules. Lamina coriaceous, elliptic to ovate or 

to lanceolate, (6-)9-16 x (2-)3-7 cm, apex acute, 

base obtuse to acute, margin entire; upper surface 

glabrous, lower surface in the areoles rather 

densely, minutely puberulous, on the main veins 

sparse, rather long, yellowish hairs, initially the 

whole surface densely covered with white arachnoid 

hairs; lateral veins 5-8 pairs, almost straight, 

basal pair unbranched, reaching the margin at or 

just below the middle; intercostal venation rather 

prominent; petiole 3-6 cm long, villous with white 

arachnoid hairs, glabrescent; stipules 5-8 cm long, 

villous with white arachnoid hairs, mixed with 

rather long and straight appressed, yellowish hairs 

and with reddishbrown pluricellular hairs. Staminate 

inflorescences unknown. Pistillate inflorescences 

branched; heads 2-4, globose ca. 4-5 

mm diam.; common peduncle 1.5-2 cm long, 

densely villous with yellowish to white arachnoid 

hairs, mixed with straight appressed yellowish 

hairs and with reddish-brown pluricellular hairs, 

apices of the branches broadened towards the 

heads; perianth ca. 1 mm high, minutely puberulous. 

Interfloral bracts absent. 

Distribution (Fig. 6). Brazil (Amapa). 

Specimens studied. BRAZIL. AMAPA: Rio Araguari, 

between 1?55'N, 51059'W and 2?5'N, 51056'W, 1 Sep 

1961 (v), Pires et al. 50656 (IAN, M, MG, NY, OXF, 

US, VEN, type collection). 

Coussapoa arachnoidea appears to be closely 

related to C. orthoneura, from which it differs in 

the prominent intercostal venation and the dense 

covering of white arachnoid hairs on the leaves 

and inflorescences. Staminate inflorescences are 

needed to determine the validity or not of the 

presumed relationship. It may prove to be distinct 

from C. orthoneura only at the subspecific 

level. 

3. Coussapoa argentea Akkermans & C. C. Berg, 

Proc. Kon. Ned. Akad. Wetensch. Ser. C, 85(4): 

443. 1982. Type. Venezuela. Bolivar: 119 km 

S of El Dorado, 12 Jan 1964 (6), Steyermark 

et al. 92991 (holotype, VEN; isotypes, NY, U, 

US). Fig. 12. 

Tree, hemi-epiphytic or terrestrial, up to 30 m 

tall. Leafy twigs 3-5 mm thick, brownish pu- 

berulous to strigose or partly to hirsute and with 

brown to white arachnoid hairs. Lamina coria- 

ceous, obovate to elliptic, occasionally ovate or


FG. 11. Coussapoa1 ly tg wh p e is (s et 

FIG. 11. Coussapoa arachnoidea: 1, leafy twig with pistillate inflorescences 

(Pires et al. 50656). 

oblong, 2-12 x 1-6 cm, apex obtuse to acute or 

acuminate, base obtuse, margin entire; upper 

surface glabrous, lower surface sparsely to densely 

appressed-puberulous and with a dense cov- 

cm. 

31 

ering of silvery arachnoid hairs, deciduous with 

age; lateral veins 2-3(-4) pairs, slightly curved, 

basal pair unbranched, reaching the margin above 

the middle of the lamina; intercostal venation


c. 

FIG. 12. Coussapoa argentea: 1, leafy twig with pistillate inflorescences (Steyermark & Nilsson 487); 2, leafy 

twig with staminate inflorescences (Steyermark & Nilsson 388).


slightly prominent to almost plane; petiole 1-2 abruptly thickened just below the apex and widely 

cm long, brownish puberulous to strigose or part- hollow, (rather) sparsely puberulous to hispiduly 

to hirsute and with brown to white arachnoid lous, often also with brown pluricellular hairs, 

hairs; stipules 1-5 cm long, densely covered with sometimes partly hirtellous. Lamina coriaceous 

brownish curled to arachnoid hairs and whitish to subcoriaceous, obovate to subobovate to elstraight 

hairs. Staminate inflorescences repeat- liptic to oblong to suborbicular to ovate or to 

edly branched; heads many, globose, ca. 2-3 mm cordiform, 7-32 x 4-24 cm, apex rounded to 

diam.; common peduncle 2-3 mm long, puber- obtuse to emarginate, base obtuse to rounded to 

ulous to hirtellous; perianth ca. 1 mm high, truncate to cordate, margin entire to subcrenate; 

3-lobed, glabrous; stamens two, exceeding the upper surface scabrous with minute rigid hairs 

perianth. Pistillate inflorescences unbranched; to smooth and glabrous, lower surface densely 

head globose, ca. 4 mm diam.; peduncle 2-3.5 to sparsely puberulous with curved (to straight) 

cm long, puberulous, and with silvery arachnoid hairs or only minutely puberulous on the main 

hairs; perianth ca. 1 mm high, glabrous. Inter- veins, sometimes partly hirtellous, in juvenile 

floral bracts spathulate to subpeltate, sometimes leaves sometimes the whole surface hisfew. 

pid(ulous); lateral veins 4-9 pairs, usually all or 

Distribution (Fig. 6). Venezuela (Bolivar); most of them branched, sometimes the basal pair 

mostly in moist forest on table mountains, be- unbranched or faintly branched but some of the 

tween (450-)1000 and 1300 m. 

other lateral veins poorly branched, (main) basal 

pair reaching the margin at, below, or above the 

Specimens studied. VENEZUELA. BOLIVAR: Road middle of the 

El Dorado-Santa 

lamina; intercostal venation 

Elena, km 109, 23 

prom- 

Feb 1959 (6), Bernardi 

7238 

inent to 

(G); upper Rio Cuyuni, Cumbre de La Es- plane; petiole 1.5-15 cm long, sparsely 

calera, 1000 m, 20-21 Aug 1962 (6), Maguire et al. to densely minutely puberulous, sometimes part- 

46879 (U, US, VEN); nr. Cerro Uei, between Luepa ly hirtellous; stipules 0.5-3.5 cm long, puberuand 

Cerro Venamo, 1100 m, 20-22 Apr 1960 (6), Stey- lous to hispidulous, often also with brown pluriermark 

& Nilsson 388 (NY, US, VEN); NE of Luepa, cellular 

1300 m, 23 Apr 1960 (v), Steyermark & Nilsson 

hairs, sometimes 

487 

partly hirtellous. 

(NY, VEN); 119 km S of El Dorado, 12 Jan 1964 Staminate 

(6), 

inflorescences branched; heads many, 

Steyermark et al. 92991 (NY, U, US, VEN, type col- free or partly fused, (sub)globose, 3-10 mm diam.; 

lection). 

common peduncle 2-8 cm long, puberulous; 

perianth ca. 1 mm high, minutely puberulous or 

Coussapoa argentea is similar to C. micro- glabrous; stamen one, as long as or slightly longer 

cephala in many features. The species differs from than the perianth. Pistillate inflorescence unthe 

latter in the smaller number of lateral veins, branched or branched; heads 1-6, free or partly 

the basal pair of which reaches the margin below fused, 3-12 mm, in fruit up to 30 mm diam.; 

the middle of the lamina. Moreover, the pistillate (common) peduncle 1-9.5 cm long, puberulous 

inflorescence is unicapitate, being normally plur- to almost glabrous; perianth 1.2 mm high, miicapitate 

in C. microcephala. The two species nutely puberulous or glabrous. Interfloral bracts 

appear to differ ecologically (altitudinal condi- (sub)spathulate to (sub)peltate, minutely pubertions) 

and may prove to be distinct only at the ulous at the apex. 

subspecies level. 

C. asperifolia differs from the other Coussapoa 

species: (1) The endocarp body is pushed out of 

4. Coussapoa asperifolia Trecul, Ann. 

the 

Sci. Nat. perianth through the torn apex of the peri- 

Bot. Ser. 3, 8: 96. 1847; Miquel, Fl. bras. anth, due to the 

4(1): 

expansion of the mucilaginous 

135. 1853; Macbride, Publ. Field Mus. Bot. mesocarp (see p. 8). (2) Furthermore, the leafy 

13(2.2): 296. 1937; Berg, Fl. Suriname twigs widen rather abruptly 

5(1): 

just below the apex 

284. 1975. Lectotype. Suriname. 

and have 

Without 

wide 

lo- 

cavities, which are often occupied 

cality, 1843 ($), Hostmann 1189 by ants. 

(holotype, P; 

Within this 

isotypes, B, G, K, LE, U). 

widespread and polymorphic 

species three subspecies can be recognized. These 

Tree or shrub, hemi-epiphytic or terrestrial, up three subspecies are not well separated morphoto 

25 m tall. Leafy twigs 3-20 mm thick, (usually) logically or geographically.


Key to the Subspecies of Coussapoa asperifolia 

1. Lamina scabrous to scabridulous above, densely (to rather sparsely) puberulous with curved hairs on 

the smaller veins beneath; intercostal venation prominent; lamina mostly obovate. ....a. ssp. asperifolia. 

1. Lamina smooth (to scabridulous) above, glabrous or sparsely puberulous on the smaller veins, sparsely 

to rather densely minutely puberulous with straight hairs on the main veins beneath; intercostal venation 

plane to prominent. 

2. All or most lateral veins branched, sometimes poorly so and the basal pair unbranched; lamina 

mostly ovate to cordiform or elliptic; pistillate heads mostly 1, more than 10 mm, in fruit up to 35 

mm in diam. ........................................................... ..b. ssp. magnifolia. 

2. Few or none of the lateral veins (poorly) branched, the basal pair unbranched or faintly branched; 

lamina mostly obovate to subobovate; pistillate heads 3-5, ca. 3-6 mm, in fruit up to 15 mm in 

diam ........ ........................................................... c. ssp. rham noides. 

4a. Coussapoa asperifolia ssp. asperifolia 1921 (6), BW 5262 (A, U); without locality, 1843 (8), 

Fig. 13. Hostmann 1189 (B, G, K, LE, P, U, lectotype collection); 

Lely Mts., 19 Sep 1975 (st), Lindeman et al. 34 

Coussapoa ficina Standley, Publ. Field Mus. Bot. 17: (U); Concession Haenen, W of mouth of Coppename 

161. 1937. Type. Brazil. Amazonas: Nr. Manaus, River, 25 Mar 1954 (st), Lindeman 5709 (U); upper 

road to Aleixo, 12 Aug-1 Sep 1936 (9), Krukoff7966 Litani River, 9 Aug 1937 (a), Rombouts 803 (A, K, 

(holotype, NY; isotypes, A, BM, F, K, LE, MO, S, NY, U); Plantation Lustrijk, Dec 1837 (8), Splitgerber 

U). 

448 (=122) (L, P); Maratakka River, 20 Aug 1976 (2), 

Coussapoa cayennensis Hawkes, Phytologia 3: 30. 1948. Teunissen (LBB) 16031 (U). 

Type. French Guiana. Nr. Cayenne, 17 Jul 1921 (9), FRENCH GUIANA. Itany River, island in rapids, 

Broadway 880 (holotype, NY; isotypes, GH, NY, 30 Sep 1962 (9), BAFOG (=Graimleux) 7942 (CAY, 

US). 

U); nr. Cayenne, 17 Jul 1921 (9), Broadway 880 (GH, 

NY, US, type collection of C. cayennensis); Oyapock 

Lamina mostly obovate to subobovate or el- River, Les Trois Sauts, 27 Mar 1975 (9), Grenand 966 

liptic to oblong, mostly 10-17 cm long, base 

(CAY); Anse Bruyere, 15 Aug 1978 (a), Grenand 1596 

(U); nr. Cayenne, 

mostly obtuse to rounded; upper surface scabrous 

with a dense covering of minute rigid hairs, 

sometimes scabridulous, lower surface on the 

smaller veins mostly densely puberulous with 

curved (to straight) hairs; lateral veins 4-7 pairs, 

all or most of them branched, the (main) basal 

pair reaching the margin above the middle of the 

lamina; intercostal venation prominent; petiole 

1.5-9 cm long; stipules 0.5-1.5 cm long. Staminate 

inflorescences with heads up to 10 mm 

diam.; common peduncle up to 4.5 cm long. Pistillate 

inflorescences with mostly, sometimes two, 

(partly fused) heads, mostly more than 1 cm, in 

fruit up to 2.5 cm diam.; (common) peduncle 1- 

3 cm long. 

Distribution (Fig. 6). Eastern Guiana region 

(Surinam and French Guiana), in Brazil in Amapa 

and eastern Para and also, apparently segregated 

from the main area, in the central part of 

the Amazon Basin; in upland and riverine forest. 

This distribution shows striking similarities to 

that of C. latifolia. 

Specimens studied. SURINAM. Section 0, 11 Nov 

1915 (st), BW1322 (U); Corantijn River, Kabouri, 26 

Jun 1916 (st), BW2070 (U); Maratakka River, 25 Nov 

1917 (9), BW 3430 (F, NY, U, US); Section 0, 10 Aug 

- (9), Leman s.n. (P); nr. Cayenne, 

Mar 1900 (st), Lemee s.n. (P); without locality, - (9), 

Leprieur s. n. (P); Kuataka, Antecume Pata, confluence 

of Itany River and Marouini River, 14 Nov 1977 (9), 

Moretti 837 (CAY); Approuague River, between Sapokaye 

Creek and Grand Canori Falls, 23 Oct 1968 

(9), Oldeman T243 (CAY, GH, NY, P, U); Oyapock 

River, between Notaye Creek and Caiman Creek, 10 

Jul 1969 (st), Oldeman T366 (CAY, U); Oyapock River, 

Savane-Roche, Baton Pilon, 12 Jul 1969 (6), Oldeman 

B2538 (CAY, U); Oyapock River, above Oscar, 

18 Jun 1970 (9), Oldeman B3420 (CAY, P, U); without 

locality, 1820 (9), Perrotet s.n. (P); Sinnamary, road to 

St. Elie, km 8, 28 Aug 1979 (st), Provost 738 (U); 

Acarouany, 1856 (st), Sagot s.n. (P); nr. Kourou, 14 

Dec 1978 (9), Sastre 6417 (CAY, P, U); without locality, 

23 Jun 1921 (9), Wachenheim 426 (P). 

BRAZIL. AMAPA: Rio Araguari, between 1?2'N, 

51?15'W and 0?57'N, 51?29'W, 13 Sep 1961 (6), Pires 

et al. 50913 (NY, OXF, R, U, US). AMAZONAS: Nr. 

Manaus, BR. 17, ca. km 9, 5 Aug 1955 (9), D. Coelho 

(INPA) 1577 (U), 8 Jul 1955 (8), Chagas (INPA) 1364 

(U); nr. Manaus, Colonia Joao Alfredo, 21 Aug 1947 

(9), Ducke 2103 (IAN, R, U); nr. Manaus, Pensador, 

Aug 1910 ($), Ule 8836 (G, K, MG); Manaus-Caracarai 

road, ca. km 130, 13 Feb 1974 (9), Stewardet al. P.20255 

(MO, U). PARA: Res. Florest, Rio Aura, 18 Oct 1978 

(9), Carauta et al. 3011 (RB); Belem, 21 Oct 1915 (st), 

Ducke (HAMP) 15804 (BM); Mun. Oriximinf, Rio 

Mapuera, 19 Aug 1986 (a), Ferreira et al. 7831 (BG); 

Arama, 2 Nov 1900 (9), Huber (HAMP) 1873 (G); Rio 

Jan, rd. to Munguba, km 15, 13 Jul 1964 (9), N. T. 

Silva 2166 (U).

.". ~:.,, . 

.


4b. Coussapoa asperifolia ssp. magnifolia (Trecul) 

Akkermans & C. C. Berg, Proc. Kon. Ned. 

Akad. Wetensch. Ser. C, 85(4): 445. 1982. 

Fig. 14. 

Coussapoa magnifolia Trecul, Ann. Sci. Nat. Bot. Ser. 

3, 8: 98. 1847; Miquel, Fl. bras. 4(1): 139. 1853; 

Macbride, Publ. Field Mus. Bot. 13(2.2): 297. 1937; 

Woodson & Schery, Ann. Missouri Bot. Gard. 47: 

169. 1960. Type. Peru. Without locality, - (2), Ruiz 

& Pavon s.n. (or 4) (holotype, FI; isotypes, B, BM, 

F, US). 

Coussapoa ruizii Klotzsch, Linnaea 20:529. 1847. Type. 

Peru. Without locality, - Specimens studied. PANAMA. CANAL ZONE: Fort 

San Lorenzo, 13 Sep 1947 (a), Alien 5120 (BM, G, 

MO); Barro Colorado Island, 27 Apr 1968 (a), Croat 

5124 (DUKE, F, MO), (9), Croat 5125 (F), 1 Oct 1968 

(9), Croat 6588 (MO), 10 Feb 1969 (8), Croat 7859 

(MO), 13 Jul 1969 (6), Foster 1101 (DUKE, F), Chagres, 

22 Jun 1860 (9 + d), Hayes 354 (NY, syntypes of C. 

chagresiana); nr. Fort Sherman, 16 Nov 1955 (2), Johnston 

1709 (A); E of Indio Tower, 2 Apr 1956 (9), Johnston 

1768 (A, MO); Barro Colorado Island, 16 Feb 

1932 (a), Woodworth & Vestal 606 (A, F, MO). 

COLOMBIA. ANTIOQUIA: Rio Porce, 1000 m, Dec 

1962 (2), Espinal 882 (COL). BOYACA: El Humbo region, 

130 km N ofBogoti, 1200-1300m, 19 May 1933 

(9), Ruiz & Pavon s.n. (or (8), Lawrance 810 (A, F, G, K, MO, US, type collection 

4) (holotype, B; isotypes, BM, F, FI, US). 

of C. hypochlora). CAQUETA: La Tagua, 5 May 1953 

Coussapoa hypochlora Standley, Publ. Field Mus. Bot. (9), Romero-Castaneda 4211 (COL, MO). CHOCO: Nr. 

17: 162. 1937. Type. Colombia. Boyaca: El Humbo Helipad, Rio Truando, at junction with Quebrada Buche, 

region, 130 km N of Bogota, ca. 1150 m, 19 May 1 Apr 1968 (a), Duke 15742 (U); El Valle, beach, 5 

1933 (6), Lawrance 810 (holotype, F; isotypes, A, G, Aug 1976 (9), Gentry et al. 17217 (MO, NY, U); Rio 

K, MO, US). 

El Valle, between El Valle and village nr. mouth of Rio 

Coussapoa cardonae Pittier, Bol. Soc. Venez. Cienc. Mutatt, 10 Aug 1976 (8), Gentry et al. 17493 (U); 5 

Nat. 8: 257. 1943, as cardonaei. Type. Venezuela. km E of Istmo de San Pablo, ca. 12 km W of Las 

Bolivar: Rio Caura, sabanas de Canaracuni, Jan 1942 Animas, 12 Jan 1979 (2), Gentry et al. 24074 (MO, U). 

(9), Cardona 374 (holotype, VEN; isotypes, GH, S, CUNDINAMARCA: La Palma, road to Pacho, Rio Murca, 

US). 

1150-1400 m, 27 Jul 1947 (2), Garcia-Barriga 12408 

Coussapoa chagresiana Hawkes, Phytologia 3: 30. 1948. (US). SANTANDER: Rio Opon, Atlantico railway, 5 Jul 

Syntypes. Panama. Canal Zone, Chagres, 22 Jun 1860 1958 (6), Garcia-Barriga 16239 (NY). VALLE: Nr. 

(6 and 9), Sutton Hayes 354 (NY). 

Buenaventura, 15 Nov 1979 (8), Van Rooden et al. 258 

(U). VAUPES: Rio Vaup6s, Yurupari Falls, 12 Apr 1953 

Lamina mostly ovate to cordiform or elliptic (8), Schultes et al. 18996 (F), Nov 1951 (2), Schultes et 

al. 19758 

to suborbicular, sometimes obovate, mostly 17- 

(GH, MO, US). 

VENEZUELA. AMAZONAS: Between Yavita and 

25 cm long, base often subcordate to cordate; Maroa, 6-19 Jul 1969 (st), Bunting et al. 3959 (MY, 

upper surface smooth and glabrous, sometimes U); upper Rio Orinoco, Raudal de los Guaharibos, 24 

scabridulous, lower surface on the smaller veins Jul 1951 (2), Croizat 343 (F, US); Yavita, 10 Jul 1972 

glabrous or sparsely puberulous, on the main (9), Lizot (1972-) 1 (US, VEN); San Carlos de Rio 

Negro, 7 Mar 1942 (9), LI. Williams 14675 (A, F, G, 

veins sparsely to densely minutely puberulous K, NY, RB, US, VEN). BOLIVAR: Rio Caura, sabanas 

with straight hairs, sometimes on the main veins de Canaracuni, Jan 1942 (9), Cardona 374 (A, S, US, 

partly hirtellous; lateral veins 4-9 pairs, all or VEN, type collection of C. cardonae); Rio Cuyuni, El 

most of them branched, sometimes the basal pair Chibau, 2 km S of Rio Chibau, 1 Sep 1962 (9), Maguire 

et al. 53517 

not or faintly branched but some of the other 

(F, NY, OXF, US, VEN); Rio Cuyuni, NE 

of Luepa, 1300 m, 23 Apr 1960 (9), Steyermark & 

lateral veins poorly branched, basal pair reaching Nilsson 507 (NY, US, VEN); Ptari-tepui, 1600 m, 11 

the margin at, below, or above the middle of the Nov 1964 (st), Steyermark 60002 (F, VEN); between 

lamina; intercostal venation plane to prominent; Santa Teresita de Kavanayen and Rio Pacairao, Quebrada 

petiole 4-17 cm long; stipules 0.5-3.5 cm Oparu-ma, 1065-1220 m, 20 Nov 1944 (9), Steylong. 

ermark 60432 (F, VEN); between Rio Paramichi and 

Staminate inflorescences with the heads up to 6 Salto de Chalimano, NW of Serrania Pia-soi, 5 Jan 

mm diam.; common peduncle up to 8 cm long. 1962 (9), Steyermark 90583 (US, VEN); Rio Parami- 

Pistillate inflorescences mostly with one head, chi, between mouth of Rio Paramichi and Salto de 

more than 1 cm, in fruit up to 3 cm diam.; (com- Chalimano, 8-9 Jan 1962 (9), Steyermark 90736 (K, 

mon) peduncle 1.5-9.5 cm 

NY, US); Rio Cuyuni, 140-142 km S of El Dorado, 

long. 

1300-1380 m, 22-28 Dec 1970 (9), Steyermark et al. 

Distribution (Fig. 6). Amazon Basin (Bolivia, 104348 (MY, NY). DELTA AMACURO: Cariapo, 13 Dec 

Brazil, Peru, Ecuador, Colombia, and Venezue- 1953 (9), Little et al. 15990 (VEN); Dto. Antonio Diaz, 

la), western Guiana region (Guyana, Venezuela), 

Canfo Guiniquina, 5 Feb 1977 (6), Marcano-Berti et al. 

84-22-77 

and in the central and Pacific coastal part of Co- 

(U); Rio Cuyubini, above Casa Cuyubini, 12 

Nov 1960 (9), Steyermark 87480 (G, GH, NY, U, US, 

lombia, as far north as Panama; in upland and VEN); Dto. Antonio Diaz, Caio Htioba, tributary of 

riverine forests up to 2000 m. 

Rio Araquoa, 16 Oct 1977 (st), Steyermarket al. 114759


FIG. 14. Coussapoa asperifolia ssp. magnifolia: 1, leafy twig with pistillate inflorescences (Harling et al. 

14733); 2, staminate inflorescence (Garcia-Barriga 16239).


(MO); Dto. Antonio Diaz, upper part of Cafno Jobure, 4c. Coussapoa asperifolia ssp. rhamnoides 

7 Apr 1955 (2), Wurdack 286 (F, NY). 

(Standley) Akkermans & C. C. Berg, Proc. Kon. 

GUYANA. Upper Rupununi River, nr. Dadanawa, 

24-29 Jul 1922 (9), De la Cruz 1729 

Ned. Akad. Wetensch. Ser. 

(F, GH, MO, NY); 

C, 85(4): 445. 

Mazaruni station, 7 May 1943 (2), Fanshawe 1278 (FD 1982. Fig. 15. 

4014) (K, NY); Essequibo River, Tiger Creek, 3 Jun 

1943 (6), Fanshawe 1337 (FD 4073) (K, NY); Esse- Coussapoa rhamnoides Standley, Publ. Field Mus. Bot. 

quibo River, Bartica, 8 Jul 1952 (6), Fanshawe 3409 17: 166. 1937. Type. Brazil. Amazonas: Mun. Sao 

(FD 6973) (K, NY); Pomeroon River, Kabakaburi, 19 Paulo de Olivenca, 11 Sep-26 Oct 1936 (9), Krukoff 

Aug 1918 (2), Hohenkerk s.n. (FD 738) (K, OXF); Ber- 8406 (holotype, NY; isotypes, A, BM, F, G, K, LE, 

bice River, Apr 1889 (2), Jenman 4947A (K); Deme- MO, S, U). 

rara River, - (8), Parker s.n. (K); Ekaria, Rosedale, 

Aug 1924 (2), Persaud 116 (B, F, K, NY). 

Lamina obovate to subobovate or elliptic to 

ECUADOR. NAPO: 3 mi SW of Tena, 28 Aug 1960 oblong, 7-16 cm long, base obtuse to rounded 

(2), Grubb et al. 1498 (K, NY). PASTAZA: Nr. Mera, (to subcordate); upper surface smooth and glaca. 

1000 m, 10 Dec 1955 (9), Asplund 18780 (S); between 

Palora and Shell, 4 Mar 1980 (st), Berg & Akbrous; 

lower surface on the main veins sparsely 

kermans 1119 (U), (9), Berg & Akkermans 1133 (U); minutely puberulous with straight hairs; lateral 

nr. Mera, 17 Jan 1977 (9), Harling et al. 14733 (GB), veins 4-6 pairs, unbranched or some of the lat- 

18 Mar 1940 (8), Lugo 83 (S). 

eral veins (poorly) branched, the basal pair un- 

PERU. Without locality (2), Ruiz & Pavon s.n. (or branched or faintly branched, reaching the mar- 

4) (B, BM, F, FI, US, type collection of C. magnifolia 

and C. ruizii). JUNiN: La Merced, 10-24 Aug 1923 gin above the middle of the lamina; petiole 2-7 

(2), 

Macbride 5447 (F, C). LORETO: Prov. Maynas, Pefia cm long; stipules 0.5-2 cm long. Pistillate inflo- 

Negra, 25 km SW of Iquitos, 1 Aug 1972 (st), Croat rescences with (1-)3-5 (sometimes partly fused) 

18643 (MO, NA); Prov. Requena, Rio Ucayali, Jenaro heads, 3-6 mm, in fruit up to 15 mm diam.; 

Herrera, below Requena, 9 Dec 1977 (2), Gentry et al. 

21319 (MO, U); Prov. Maynas, nr. Quistococha, 26 

(common) peduncle 1-4 cm long. 

May 1978 (6), Gentry et al. 22280 (MO, U); Prov. 

Distribution (Fig. 6). Amazon Basin (Brazil, 

Maynas, Rio Nanay, below Bellavista, 31 May 1972 Peru); in upland and riverside (varzea) forest. 

(6), McDaniel 16092 (NA); Prov. Maynas, Dist. Iquitos, 

Puerto Almendra, Rio Nanay, 1 Nov 1976 (6), Specimens studied. PERU. LORETO: Rio Tacsha 

Revilla 1260 (MO, U). SAN MARTIN: Prov. Marichal Curaray, 18 Sep 1972 (2), Croat 20370 (F, GH, NA, 

Caceres, Dist. Tocache Nuevo, Fundo Gran Chaparral, NY, P, RB). 

11 Nov. 1975 (6), Schunke 8666 (U); Prov. Mo- BRAZIL. AMAZONAS: Rio Solimoes, Rio Bia, affluyobamba, 

1906 (st), Weberbauer 4472 (G). 

ent of Rio Jutai, 5 Nov 1975 (Q), L. Coelho et al. 332 

BRAZIL. ACRE: Cruzeiro do Sul, 2 Mar 1976 (2), (U); Barcelos, 7 Sep 1962 (Q), Duarte et al. 6970 (RB); 

Marinho 348 (IAN). AMAZONAS: Rio Negro, Sao Ga- Mun. Sao Paulo de Olivenca, nr. Palmares, 11 Sep-26 

briel de Cachoeira, 28 Feb 1975 (9), Cordeiro 367 (IAN); Oct 1936 (2), Krukoff8406 (A, BM, F, G, K, LE, MO, 

mouth of Rio Caiari, affluent of Rio Negro, 15 Sep NY, S, U, type collection of C. rhamnoides); Manaus- 

1952 (?), Fro6s et al. 28585 (IAN); mun. Humaiti, Itacoatiara road, Res. Flor. Ducke, 1 Sep 1966 (2), 

nr. Livramento on Rio Livramento, 12 Oct-6 Nov Prance et al. 2153 (INPA, NY, U). PARA: Rio Tapajos, 

1934 (2), Krukoff6662 (A, B, BM, F, G, K, LE, MO, Periquito, 27 Jul 1923 (2), Ducke (HJBR) 18460 (RB). 

NY, RB, S, U, US); nr. Manaus, 1906 (2) Lambroy s.n. 

(P); Rio Negro, Uanari, nr. Sao Gabriel, 31 Oct 1947 

(6), Pires 805 (IAN, NY), 2 Nov 1947 (9), Pires 830 5. Coussapoa batavorum Akkermans & C. C. 

(NY); Rio Negro, between Sao Gabriel and Sao Felipe, Berg, Proc. Kon. Ned. Akad. Wetensch. Ser. 

base of Serra Uanari, Oct 1947 (6), Schultes & Pires C, 85(4): 447. 1982. 

8976 (GH, U); Rio Negro, Sao 

Type. Colombia. Valle: 

Gabriel de Cachoeira, 

Jan-Aug 1852 (6), Spruce 2260 (BR, K, P). MATO 

Nr. Buenaventura, 5 Dec 1979 (9), Van Rood- 

GRosso: Rio Aripuafia, nr. Humboldt Center, 59?21'N, en et al. 700 (holotype, COL; isotypes, AAU, 

10?12'S, 19 Oct 1973 (st), Berg s.n. (U); Rio Juruena, K, NY, U, US). Fig. 16. 

airport, 17 Jul 1977 (9), M. G. Silva et al. 3368 (U). 

PARA: Almeirim, Barreira da Velha Pobre, 6 Jul 1919 Tree, terrestrial, ca. 15 m tall. Leafy twigs 10- 

(2), Ducke (HBJR) 13605 (RB); Rio Branco de Obidos, 19 mm 

21 Jul 1918 (6), Ducke (HAMP) 17132 thick, glabrous. Lamina coriaceous, 

(MG). 

RONDONIA: Nr. Porto Velho, 28 Jun 1952 (9), J. F. broadly ovate to cordiform, 36-40 x 34-37 cm, 

Silva 227 (IAN); Rio Madeira, Cachoeira Misericorde, apex obtuse to emarginate, base cordate, margin 

Riberao, 2 Aug 1968 (9), Prance et al. 6718 (NY, U). subcrenate; both surfaces glabrous; lateral veins 

RORAIMA: West facing slopes of Serra Tepequem, 1200 10-13 

m, 20 Feb 1967 (2), Prance et al. 4588 

pairs, 3-5 of which arise at the base of the 

(NY, U). 

BOLIVIA. LA PAZ: Prov. S. Yungas, basin of Rio midrib, straight, main basal pair branched, 

Bopi, San Bartolome, nr. Calisaya, 1-22 Jul 1939 (2), reaching the margin at or above the middle of 

Krukoff 10123 (A, F, G, K, MO, NY, S, U). the lamina; intercostal venation almost plane to


FIG. 15. Coussapoa asperifolia ssp. rhamnoides: 1, leafy twig with pistillate inflorescences (Krukoff 8406). 

slightly prominent; petiole 10-12 cm long, dis- 

tinctly ribbed and densely minutely puberulous; 

stipules 8-10 cm long, densely to sparsely red- 

dish-brown puberulous and with longer straight 

white hairs. Staminate inflorescences unknown. 

Pistillate inflorescences unbranched or poorly 

branched; heads 1-2(-3), free or partly fused, 

globose to ovoid, ca. 8-10 mm, in fruit up to 27 

mm diam.; common peduncle 0.5-2 cm long, 

rather densely puberulous to hirtellous; perianth 

1 cm 

1(-3) mm high, white puberulous; fruiting peri- 

anth brown. Interfloral bracts absent. 

Distribution (Fig. 6). Colombia (Choco); in 

riverside forest. Known only from the type col- 

lection. 

Knowledge of the staminate inflorescence is 

required in order to determine the relationships 

of this species. In the shape, size, and sparse 

indumentum of the lamina, it resembles C. as- 

perifolia ssp. magnifolia as well as forms of C.


FIG. 16. Coussapoa batavorum: 1, leafy twig with pistillate inflorescences 

(Van Rooden et al. 700).


vannifolia having leaves with a sparse indumen- NY); Cocle del Norte, 23 Aug 1978 (2), Hammel 4475 

tum. 

(U); ca. 11 km SW of Cerro Braja, 9?25'N, 79?35'W, 

2 May 1981 (8), Sytsma et al. 4225 (U). PANAMA: Cerro 

Jefe, 3 Mar 1979 (8), d'Arcy 12329 (U); road to Cerro 

6. Coussapoa brevipes Pittier, Contr. U.S. Natl. Campana, 3 km from Interamerican Highway, 18 Sep 

Herb. 18:225. 1917; Woodson & Schery, Ann. 1968 (6), Correa et al. 1031A (DUKE, F, MO); Cerro 

Missouri Bot. Gard. 47:171. 1960. Type. Pan- Campana, 9 Apr 1970 (9), Croat 14212 (GH, MO); 

amf. San Bias: Nr. Puerto Obaldia, hills of 

Cerro Jefe region, La Eneida, 25 Mar 1968 (2), Dressier 

3454 (DUKE, 

Sperdi, Sep 1911 

MO); 10 km N of 

(v), Pittier 4386 

Margarita, on road 

(holotype, to Madronio, 31 Jan 1979 (8), Hammel 6006 (U); 3 mi 

US; isotype, B). Fig. 17. NE of Altos de Pacora, Campo Tres, 10 Mar 1973 (6), 

Liesner 515 (K, MO, NY, US); Altos de 

Tree, terrestrial or 

Campana, 28 

hemi-epiphytic, up to 15 m May 1977 (8), Mendez 15 (MO); 6.5 km N of Lago 

tall. Leafy twigs 4-11 mm thick, more or less Azul, 13 Jan 1974 (9), Nee 9281 (MO, NY, U); El 

densely white (appressed-)puberulous and with Llano-Carti road, 8 mi from Pan American Hwy., 17 

reddish-brown pluricellular hairs, or also with Apr 1981 (2), Sytsma 4024 (U); Cerro Campana, 18 

sparse arachnoid hairs. 

Sep 1968 

Lamina 

(a), Weaver et al. 1677 

coriaceous, ovate 

(DUKE). SAN BLAS: 

Road El Llano-Carti-Tupile, 1 mile from continental 

to elliptic or occasionally obovate, 5-30 x 2.5- divide, 30 Mar 1973 (9), Liesner 1309 (DUKE, GH, 

18 cm, apex obtuse to acute or to subcordate, MO, NY, US); nr. Puerto Obaldia, hills of Sperdi, Sep 

base obtuse to rounded to subcordate, sometimes 1911 (9), Pittier 4386 (B, US, type collection); Rio Acla, 

acute, margin entire or subcrenate towards the 8048'38"N, 77040'30"W, 7 Feb 1979 (2), Sugden 415 

(K). 

apex; upper surface glabrous, lower surface appressed-puberulous, 

densely so in the areoles, Coussapoa brevipes shows morphological afand 

on the main veins often with white arach- finities especially with C. chocoensis but also with 

noid hairs which are soon deciduous; lateral veins C. contorta. 

6-18 pairs, straight or slightly curved, basal pair 

branched, reaching the margin below the middle 7. Coussapoa chocoensis Cuatrecasas, Caldasia 

of the lamina; intercostal venation almost plane; 7:287.1956. Type. Colombia. Choc6: Rio San 

petiole 1-7 cm long, 2-4 mm thick, appressed- Juan, Quebrada del Taparal, 30 May 1946 (2), 

puberulous and often also with white arachnoid Cuatrecasas 21446 (holotype, F). Fig. 18. 

hairs which are infrequently deciduous; stipules 

3-10 cm 

Tree, 

long, densely covered with 

hemi-epiphytic. Leafy twigs 3-6 mm 

appressed 

white 

thick, brownish subhirsute to hirtellous. 

puberulous hairs, mixed 

Lamina 

with reddishbrown 

pluricellular hairs and often also 

coriaceous, ovate to 

with 

elliptic, 8-16 x 4-12, apex 

sparse to dense white arachnoid hairs and/or 

shortly acuminate, base rounded to subcordate, 

long 

yellowish straight hairs. 

margin 

Staminate 

entire; upper surface glabrous, lower surinflorescences 

face 

branched; heads 4-20, globose, ca. 4-5 mm 

densely appressed-puberulous in the areoles, 

diam.; 

common 

sparsely puberulous to hirtellous on the smaller 

peduncle 0.5-3 cm long, densely covered 

with short 

veins, 

white puberulous and 

densely subhirsute, 

reddishsubvelutinous,hirtelbrown 

lous, 

pluricellular hairs; perianth ca. 1 

strigose or 

mm 

strigillose on the main veins; 

high, lateral veins 10-12 pairs, curved and 

densely minutely puberulous; stamens three, far 

distinctly 

exceeding the 

looping, basal 

perianth. Pistillate 

pair unbranched or 

inflorescences 

sparsely 

unbranched; heads globose to 

branched, 

ellipsoid, ca. 5-7 

reaching the margin below the middle 

of the 

mm, in fruit up to 14 mm diam.; peduncle 0.4lamina; 

intercostal venation rather prom- 

1.5(-2.5) cm long, densely white 

inent; petiole 1-3 cm 

puberulous and 

long, brown subhirsute to 

also with reddish-brown 

subvelutinous; stipules 1.5-2.5 cm 

pluricellular 

long, brown 

hairs; perianth 

ca. 1-2 mm 

(sub)hirsute. Staminate inflorescences unknown. 

high, densely minutely pub- Pistillate inflorescences 

erulous; fruiting perianth orange. Interfloral bracts 

unbranched; head (globose 

or 

absent. 

ellipsoid?), ca. 15 mm diam.; peduncle 

0.5-1 cm 

Distribution (Fig. 6). Panama, in forests up to 

long, hirtellous to subhirsute; perianth 

ca. 2 mm 

800 m. 

high, minutely brown puberulous. Interfloral 

bracts subspathulate, hirtellous at the 

Specimens studied. PANAMA: COCLE: Nr. La Mesa, apex. 

4 km from El Valle, 15 Sep 1968 (Y), Correa et al. 1002 

(DUKE, F); foothills of Cerro Pil6n, nr. El Valle, 5 Oct Distribution (Fig. 6). Colombia (Choc6); low- 

1967 (Q), Duke et al. 14712 (MO, US). COLON: Nr. land, riverside. Known only from the type col- 

Guasimo, 22 Apr 1970 (Q), Croat 9973A (F, GH, MO, lection. 

41

A' 

.. 

. 

FIG. 17. Coussapoa brevipes: 1, leafy twig with staminate inflorescences (Weaver & Foster 1677); 2, pistillate in 

11 

c1m 

I

'? 

' Cous': '.' .. . . . c e :1.. 

~~~~~~~~~~~~~~~~~~~~~~~~~~~?~ 

8~~~~~~~~~~~~~~~ 

\~~~~~~~~~~~~~~~~~~~~~? '' ;: 

?~. .: .. :7 

?::: 

.:; 0 .-; - :. . 

i'1'::" .. - . 

,~~ ~ j. 'i : 

' 

?'"'",...?: 

." :,,; ' .? 

. 

iI.' ;... ?. ? 

':I~~~ 

?.:.;?...; 

?~~~~~ 

~~~~'"' (.:?? 

"~~~~~~~~~~~~~~~~~~~' 

. . i,:~..~~~~~~~~~~~~~~~~~~ 

....: 

"' ~~~~~. ?'"....:',,: 

?.;t~' . 

'-~~~~ ' ~ ~~~~~~~I 

~~~~~~~~~~~~~~~~ 

. '? ~~~~~~~~~~~~~ 

::i.: ? 

' 

.,. ':." ::'"..?.. 

''(...'".1 ~~? '"!;:"i,~'i ~~~~~~~ ~ 

FIG. 18. Coussapoa chocoensis: 1, leafy twig with pistillate inflorescences (Cuatrecasas 214 

~ 

.. \ ..?. . . 

FIG. 18. Coussapoa chocoensis: , leafy twig with pistillate iniorescences (Cuatrecasas 

~ cm


This species resembles C. brevipes, from which 

it clearly differs in the brown indumentum and 

the presence of interfloral bracts. 

8. Coussapoa cinnamomea Cuatrecasas, Calda- 

sia 7: 288. 1956. Type. Colombia. Amazonas: 

Trapecio Amaz6nico, Rio Loretoyacu, 28 Sep 

1946 (9), Schultes & Black 8269 (=Black & 

Schultes 46-137) (holotype, F; isotypes, A, 

IAN, K, US). Fig. 19. 

C. cinnamomea has many features in common 

with C. tessmannii, but in most parts is larger 

and more robust than the latter species. The two 

species appear to be closely related. 

9. Coussapoa cinnamomifolia Mildbraed, No- 

tizbl. Bot. Gart. Berlin 15: 783. 1942. Type. 

Ecuador. Pastaza: Nr. Mera, ca. 1000 m, 28 

Oct 1938 (i), Schultze-Rhonhof 2943 (holo- 

type, B). Fig. 20. 

Tree, mostly terrestrial, up to 35 m tall. Leafy 

twigs 2-5 mm thick, densely brownish appressed-puberulous, 

glabrescent. Lamina coriaceous, 

oblong to subovate or subobovate, 3-13 

x 1-5 cm, apex (sub)acute to obtuse, base obtuse 

to acute, margin entire; upper surface glabrous, 

lower surface glabrous or appressed-puberulous 

in the lower part; lateral veins 3-6 pairs, the basal 

pairs strong, unbranched, reaching the margin 

far below the middle of the lamina, the other 

lateral veins weak and inconspicuous; intercostal 

venation plane; petiole 1-2 (or -3) cm long, 

densely brownish appressed-puberulous; stipules 

0.5-2.5 cm long, brownish subsericeous to subvelutinous. 

Staminate 


inflorescences poorly 

tall. Leafy twigs 9-22 mm thick, brownish 

branched; heads 2-7, partly fused, globose, ca. 

2-3 mm 

(sub)velutinous, glabrescent. Lamina coria- diam.; common peduncle 1.5-3 cm long, 

ceous, (broadly) ovate, 16-38 x 

brownish 

12-28 puberulous, perianth ca. 1 mm high, 

cm, apex 

obtuse to (acute), base rounded to truncate or to sparsely minutely puberulous; stamens three, just 

cordate, margin subcrenate; upper surface exceeding the perianth. Pistillate inflorescences 

glaunbranched 

or 

brous, lower surface densely puberulous in the 

poorly branched; heads 1-2, 

sometimes partly fused, globose, ca. 4-7 mm, in 

areoles, (rather) sparsely appressed puberulous 

fruit 

to substrigose on the veins; lateral veins 14-22 up to 9 mm diam.; (common) peduncle 2- 

5 cm 

pairs, (almost) straight, basal pair branched, 

long, brownish puberulous; perianth ca. 1 

mm 

reaching the margin below the middle of the lam- high, glabrous; fruiting perianth yellow. Inina; 

intercostal venation terfloral bracts spathulate to 

(almost) plane; petiole 

subpeltate or pel- 

5-15 cm long, 5-7 mm 

tate. 

thick; shortly subvelu- 

Distribution 

tinous, also brown pluricellular hairs. Staminate 

(Fig. 6). Known in Colombia 

inflorescences unknown. Pistillate (Antioquia), Ecuador (Pastaza), and Bolivia 


(La 

branched; heads 2-4, often partly fused, sub- Paz); in forest at altitudes between (400-?) 800 

and 

globose, in 

1200 m. 

fruit up to 40 mm in diam.; common 

peduncle 2-4 cm long, peduncle and branches 

Specimens studied. COLOMBIA. ANTIOQUIA: Mun. 

6-10(-15) mm thick, shortly velutinous; peri- San Luis, rd. Medellin-Bogota, quebrada La Tebaida, 

anth glabrous. Interfloral bracts absent. 1000-1100 m, 22 Jun 1987 (2), Callejas et al. 3997 

Distribution (Fig. 6). Colombia (Amazonas, (BG); nr. San Luis, 900-1000 m, 21 Jun 1982 (6), Her- 

Trapecio Amaz6nico); riverside forest. 

nandez et al. 369 (HUA). 

ECUADOR. PASTAZA: Nr. Mera, ca. 1000 m, 14 

Specimens studied. COLOMBIA. AMAZONAS: Tra- Feb 1955 (9), Asplund 18840 (S), 28 Oct 1938 (6), 

pecio Amaz6nico, Rio Loretoyacu, 22 Nov 1945 (2), Schultze-Rhonhof2943 (B, type collection). 

Duque-Jaramillo 2179 (COL), Oct 1945 (2), Schultes BOLIVIA. LA PAZ: Prov. Larejaca, Copacabana, 10 

6693 (F, GH, U, US), 28 Sep 1946 (2), Schultes & Black km S of Mapiri, 850-950 m, 8 Oct-15 Nov 1939 (6), 

8269 (=Black & Schultes 46-137) (F, GH, IAN, K, US, Krukoff 11275 (A, F, G, K, MO, NY, S, U, US), (9) 

type collection). 

Krukoff 11276 (A, F, G, MO, S, U, US). 

In many characters, and especially in the sub- 

trinervate lamina, C. cinnamomifolia is similar 

to C. trinervia. The 3-staminate flowers, how- 

ever, suggest relationships with other species, in 

particular with C. parviceps. 

The Colombian collection differs somewhat 

from the other collections in the longer petiole 

(up to 3 cm) and peltate and larger bracts. 

10. Coussapoa contorta Cuatrecasas, Caldasia 7: 

289. 1956. Type. Colombia. Valle: Pacific 

Coast, Rio Naja, braza Aji, Calle Larga, 28 

Feb 1943 (6), Cuatrecasas 14284 (holotype, 

F). Fig. 21.


"~ [ 

FIG. 19. Coussapoa cinnamomea: 1, leafy twig with pistillate inflorescences (Schultes 6693 (inflorescences) 

and Schultes & Black 8269 (leaf and stipules)). 

1 cm 

45


1 cm 

FIG. 20. Coussapoa cinnamomifolia: 1, leafy twig with pistillate inflorescences (Krukoff11276); 2, staminate 

inflorescences (Krukoff 11275). 

Shrub or tree, hemi-epiphytic or terrestrial, up 

to 20 m tall. Leafy twigs 3-6 mm thick, yellowish 

appressed-puberulous or sometimes partly hirtellous. 

Lamina chartaceous to coriaceous, ellip- 

tic or oblong to ovate or obovate, 5-26 x 1.5- 

13 cm, apex acute to acuminate, base acute to 

obtuse and occasionally to cordate, margin en- 

tire; upper surface glabrous, lower surface on the


X { cm 

FIG. 21. Coussapoa contorta: 1, leafy twig with pistillate inflorescences (Javita & Epling 1097); 2, leafy twig 

with staminate inflorescences (Cuatrecasas 14284). 

main veins rather sparsely appressed-puberulous 

and sometimes, only in the axils of the lateral 

veins on more or less distinct domatium-like 

cavities, distinctly longer stiff yellowish hairs, on 

the smaller veins rather sparsely puberulous to 

glabrous; lateral veins 6-15 pairs, slightly curved, 

basal pair unbranched, reaching the margin below 

the middle of the lamina; intercostal vena- 

47


tion plane; petiole 1-8 cm long, appressed-pu- small, ca. I cm long stipules, the basal pair of 

berulous; stipules 2-4 cm long, yellowish lateral veins reaching the margin of the lamina 

appressed-puberulous to (sub)sericeous. Stami- at about the middle, and the rather dense brownnate 

inflorescence poorly branched; heads 6-10, ish hairs on the perianth of the pistillate flower. 

globose, ca. 2-3 mm diam.; common peduncle It could represent a distinct species or subspecies. 

1-3 cm long, densely puberulous to hirtellous; 

perianth ca. 1 mm high, minutely puberulous; 

stamens three, far exceeding the perianth. Pis- 11. Coussapoa crassivenosa Mildbraed, Notizbl. 

tillate inflorescences mostly unbranched, occa- Bot. Gart. Berlin 15: 784. 1942. Type. Ecuasionally 

branched; heads 1-2, globose, 5-7 mm, dor. Pastaza: Nr. Mera, ca. 1000 m, 7 Sep 1938 

in fruit up to 10 mm diam.; (common) peduncle (a), Schultze-Rhonhof 2783 (holotype, B). 

1-2 cm long, densely puberulous to hirtellous; 

Fig. 22. 

perianth 1-2 mm high, minutely puberulous. Interfloral 

bracts peltate, relatively large, densely 

Coussapoa steyermarkii Cuatrecasas, Fieldiana 

Bot. 

puberulous. 

28(1): 212. 1951. Type. Venezuela. Bo- 

Distribution (Fig. 6). Colombia and 

livar: 

Ecuador, 

Ptari-tepui, 1600 m, 10 Nov 1944 (Y), 

Pacific coastal region, also in Costa Rica?; in Steyermark 60002a (holotype, F; isotype, 

forest up to 1500 m altitude. 

VEN). 

Specimens studied. COLOMBIA. CAUCA?: Tree, often 

Juntas, 

hemi-epiphytic, up to 20 m tall. 

- (2), Lehmann BT694 (A, GH, K, L, NY). CHocO: Leafy twigs 6-9 mm thick, densely (pale) brown 

Rio San Juan, Quebrada de Taparal, 30 May 1946 (6), puberulous to hirtellous or to subhirsute to stri- 

Cuatrecasas 21496 (F); Rio San Juan, between Que- gose. Lamina coriaceous, oblong to elliptic to 

brada La Sierpe and Quebrada El Quicharo, 27 Mar suborbicular to subobovate or to 

1979 (6), Forero et al. 4099 (COL, MO, U); Rio San 

(sub)ovate, 4- 

Juan, nr. Palestina, 26 Mar 1979 (6), Forero et al. 4110 24(-43) x 2-14(-20) cm, apex acuminate to ob- 

(COL, MO, U); Rio San Juan, Quebrada de Taparal, tuse and to rounded, base obtuse to rounded (or 

5 Dec 1979 (9), Van Rooden et al. 699 (U). NARIRO: to acute), margin entire; upper surface glabrous, 

Mun. Iscuand6, Quebrada La Berrera, 28 Nov 1955 lower surface 

(6), Romero-Castaieda 5512 strigose to subsericeous on the main 

(COL). VALLE: Pacific 

Coast, Rio Naja, Calle Larga, 28 Feb 1943 (6), Cua- veins, sparsely (along margin of the lamina) putrecasas 

14284 (F, type collection); Pacific Coast, Rio berulous to hirtellous, in addition covered 

Cajambre, 5-15 May 1944 (9), Cuatrecasas 17647 (F). with (rather) dense, sometimes very sparse, 

ECUADOR. BOLIVAR: Valley of Rio Tablas, 1300 (sub)persistent 

m, 1 

(pale) brown arachnoid hairs; lat- 

Oct 1943 (st), Acosta Solis 6008 (F). CARCHI: Rio 

eral veins 

Blanco, tributary of Rio San Juan, above Chical, 1300- 

(4-)5-8 pairs, slightly curved, basal 

1500 m, 25 Sep 1979 (Q), Gentry et al. 26525 (SEL, pair branched (or sometimes unbranched), 

U). ESMERALDAS: Rio Onzole, Estero Chontaduro, 14 reaching the margin at or above the middle of 

Jul 1966 (Q), Javita et al. 1097 (MO, NY, S, US). IM- the lamina; intercostal venation (very) promi- 

BABURA: Collapi, 4 Jul 1949 (2), Acosta Solis 12797 

(F). EL 

nent; 

ORO: Trail to 

petiole 1-4 cm 

Sambotambo, Rio Moro 

long, (puberulous to) sub- 

Moro, 

S of Buenaventura, highway to Portovele, 1035-1800 strigose to hirsute; stipules 1-5 cm long, brown 

m, 29 Aug 1943 (2), Steyermark 54214 (F). PICHINCHA: subsericeous to subhirsute. Staminate inflores- 

Nr. Santo Domingo de los Colorados, 30 Aug 1930 (6), cences branched; heads 4-20, globose, ca. 2-3 

Benoist 3010 (P); road Santo Domingo de los Colo- mm diam.; common peduncle 1-4.5 cm 

rados-Quininde (km 170-175), 2 

long, 

Sep 1949 (st), Acosta 

Solis 13687, 4 Sep 1949 (8), Acosta Solis 13713 (rather) densely puberulous to hirtellous to 

(F). 

strigillose; 

perianth ca. 1 mm high, minutely brown 

The taxonomic relationships of C. contorta are puberulous; stamens two, far exceeding the perinot 

clear. Overall similarities suggest affinity with anth. Pistillate inflorescences branched; heads 3- 

C. brevipes, but a connection with C. oligoceph- 7, globose, 2-5 mm diam.; common peduncle 1- 

ala is also possible. The species is distinguished 3 cm long, brownish puberulous to hirtellous to 

by the domatium-like cavities in the axils of the strigillose; perianth ca. 1-1.5 mm high, glabrous. 

lateral veins and by the large interfloral bracts. Interfloral bracts spathulate, often the lower part 

Collection Sanchez & Zamora 512, from Cos- very narrow, minutely puberulous at the apex. 

ta Rica, San Jose, below La Hondrua, Parque Distribution (Fig. 6). Presently known from 

Nacional Braulio Carrillo, 2 Jan 1984 (2), (AAU) the following disjunct areas: the eastern Guiana 

differs from the collections cited above in the region (Venezuela and northern Brazil), Ama-


I. 

FIG. 22. Coussapoa crassivenosa: 1, leafy twig with pistillate inflorescences (Steyermark et al. 104296); 2, 

leafy twig with staminate inflorescences (Krukoff 11085). 

zonian Bolivia, Amazonian Ecuador, and Pan- NY); Meseta del Jaua, Cerro Sarisarinama, 700 m, 12ama; 

in forests, mostly between 700 and 15 Feb 1974 (6), Steyermark et al. 108988 (F, K, NY); 

1700 m. 

6 km N of Mision de Santa Teresita de Kavanayen, 

1100 m, 21 Feb 1978 (st), Steyermark et al. 115549 

(U, VEN). 

ECUADOR. MORONA-SANTIAGO: Cord. de Cutucui, 

Specimens studied. PANAMA. PANAMA: Road El 5-10 km from Logrono, 1200-1500 m, 7-9 Oct 1975 

Llano-Carti, 9'20'N, 78?50'W, 7 Jan 1981 (d), Hahn (2), Little et al. 635 (COL, Q). PASTAZA: Road Puyo- 

347 (U). 

Tena, ca. 8 km, Tnte. Hugo Ortiz, ca. 950 m, 4 Mar 

VENEZUELA. AMAZONAS: Brazilian border, ca. 1980 (8), Berg & Akkermans 1120 (U); between Palora 

2?27'24'N, 63056'W, 22 May 1972 (a), Steyermark and Shell, ca. 900 m, 4 Mar 1980 (6), Berg & Akker- 

106118 (U, VEN). BoLnvAR: Meseta del Jaua, Cerro mans 1129 (st), Berg& Akkermans 1131 (U); nr. Mera, 

Sarisarinama, 700 m, 10 Feb 1976 (a), Brewer-Carias ca. 1000 m, 7 Sep 1938 (8), Schultze-Rhonhof2783 (B, 

s.n. (COL, MY); Ptari-tepui, 1600 m, 10 Nov 1944 (2), type collection), 20 Nov 1938 (9), Schultze-Rhonhof 

Steyermark 60002a (F, VEN, type collection ofC. stey- 3006 (B). 

ermarkil); Quebrada O-paru-ma, between Santa Ter- BRAZIL. RoRAIMA: Nr. Auaris, 4?3'N, 64?22'W, 

esita de Kavanayen and Rio Pacairao, 1065-1220 m, 760-800 m, 7 Feb 1969 (6), Prance et al 9679 (F, GH, 

20-21 Nov 1944 (st), Steyermark 60364 (F, VEN); K, NY, P, S, U, US); Serra Surucucu, 26 Jan 1975 (e), 

Ptari-tepui, 1220 m, 28 Nov 1944 (st), Steyermark Ribeiro 612 (IAN, MG, RB), 23 Jan 1975 (9), Rosa 

60673 (F); Chimanta-tepui (Torono-tepui), 1700 m, 296 (IAN). 

21 May 1953 (9), Steyermark 75522 (F, NY, VEN); BOLIVIA. LA PAZ: Prov. Larecaja, Copacabana, ca. 

Rio Cuyuni, ca. 142 km S of El Dorado, 1300-1380 10 km S of Mapiri, 850-950 m, 8 Sep-15 Nov 1939 

m, 22-28 Dec 1970 (2), Steyermark et al 104296 (MY, (8), Krukoff 11085 (A, F, G, K, MO, NY, S, US). 

1 cm


This species has a remarkable, apparently dis- 

continuous, distribution. It appears to be very 

closely related to C. ferruginea. The two taxa 

could prove to be conspecific, although probably 

distinct at the subspecific level. Several speci- 

mens from Venezuela and the Roraima Territory 

(Brazil) are distinct in having relatively broad (to 

suborbicular) leaves. 

12. Coussapoa cupularis Akkermans & C. C. 


C, 85(4): 448. 1982. Type. Brazil. Rond6nia: 

P6rto Velho, 5 Jun 1952 (9), J. F. Silva 69 

(holotype, IAN). Fig. 23. 

Trees, terrestrial. Leafy twigs 3-7 mm thick, 

brownish puberulous to hirtellous. Lamina coriaceous, 

elliptic to obovate, 5-10 x 3-7 cm, 

apex obtuse to shortly acuminate, base obtuse to 

(sub)cordate, margin entire to subcrenate; upper 

surface glabrous, lower surface on the midrib and 

lateral veins sparsely to rather densely appressedpuberulous 

to strigillose, for the rest densely (appressed-)puberulous; 

lateral veins 5-8 pairs, 

slightly curved, basal pair unbranched or occasionally 

with a single branch, reaching the margin 

below to just above the middle of the lamina; 

intercostal venation plane; petiole 2-5 cm long, 

densely puberulous to strigillose; stipules 2-5 cm 

long, yellow to brownish subsericeous to subhirsute. 

Staminate inflorescences unknown. Pistil- 

of which tends to be below the middle; in the 

strongly broadened, almost cupuliform apices of 

the head-bearing branches of the pistillate inflo- 

rescences; and in the relatively dense brown pa- 

pillate apex of the perianth of the pistillate flow- 

er. Moreover, the stipules are densely covered 

with long yellowish hairs, which is uncommon 

in C. orthoneura. Coussapoa cupularis also shows 

resemblances to C. nitida. 

Because of the occurrence of more or less conspicuously 

broadened branches below the pistillate 

flower heads and other similarities, as in the 

indumentum, C. cupularis, C. orthoneura, C. 

arachnoidea, and C. nitida appear to constitute 

a rather distinct group among the taxa with 

ebracteate inflorescences and unistaminate flowers. 

Collection Huashikat 1585, Peru, Amazonas, 

basin of Rio Santiago, 65 km N of Pinglo, near 

Caterpiza, 19 Dec 1979 (9) (MO, U), resembles 

C. cupularis in many characters. The main difference 

is the branching of the basal lateral veins. 

Less important are the greater number of lateral 

veins (8-9 pairs) and some differences in the indumentum. 

In the leaf venation the specimen 

approaches C. nitida. The identity of this collection 

is uncertain. 

13. Coussapoa curranii Blake, Contr. U.S. Natl. 

Herb. 20: 237. 1919. Type. Brazil. Bahia: Rio 

late inflorescences branched; heads 3-4, globose, Gongoji, 1 Oct-30 Nov 1915 (9), Curran 8 

ca. 4-6 mm, in fruit up to ca. 10-15 mm diam.; (holotype, US; isotype, RB). Fig. 24. 

common peduncle 2-4 cm long, puberulous to 

Coussapoa warburgiana Mildbraed, Notizbl. Bot. Gart. 

hirtellous, branches below the heads more or less Berlin 10: 416. 1928. Type. Brazil. Rio de Janeiro: 

strongly broadened (more or less cupuliform); Serra da Estrella, 21 May 1877 (3), Glaziou 8934 

perianth ca. 1-2 mm high, subpapillate with mi- (holotype, B; isotypes, BR, G, K, LE, MO, P, S, U, 

nute thick brown hairs. Interfloral bracts absent. US). 

Coussapoa incomitata Standley, Publ. Field Mus. Bot. 

Distribution (Fig. 6). Brazil (Rond6nia); in for- 22: 72. 1940. Type. Brazil. Minas Gerais: Mun. 

est of terra firme. 

Tombos, Fazenda da Cachoeira, 29 Jul 1935 (9), 

Mello Barreto 1795 (holotype, F). 

Specimens studied. BRAZIL. RONDONIA: Road Porto 

Velho-Cuiaba, km 117, Rio Jamari, 15 Aug 1968 Tree, (mostly?) terrestrial, up to 40 m tall. 

(2), Prance et al. 6969 (U); 8 km from Porto Velho, 5 Leafy twigs 4-6 mm thick, glabrous or sparsely 

Jun 1952 (2), J. F. Silva 69 (IAN, type collection), 26 

Jun 1952 (2), J. F. Silva 211 puberulous, sometimes initially with sparse 

(IAN). 

arachnoid hairs. Lamina (sub)coriaceous, ob- 

Coussapoa cupularis, represented by three col- ovate or subobovate, 4-19 x 1-9 cm, apex oblections 

from the same area, appears to be closely tuse to rounded to emarginate or shortly acurelated 

to C. orthoneura, and may even prove minate, base obtuse to acute, margin entire; upper 

not to be distinct at the species level. At present, surface glabrous or with sparse (to rather dense) 

the species differs from C. orthoneura in the basal white arachnoid hairs which soon disappear, 

lateral veins which mostly reach the margin be- lower surface sparsely puberulous, often also with 

low the middle of the lamina, the broadest part (mostly sparse) deciduous arachnoid hairs; lat-


1 cm 

FIG. 23. Coussapoa cupularis: 1, leafy twig with pistillate inflorescences (J. F. Silva 69). 

eral veins 7-10 pairs, basal pair unbranched or 

sometimes faintly branched, reaching the margin 

below the middle of the lamina; intercostal venation 

slightly prominent, the smaller veins plane; 

petioles 1-5 cm long, glabrous to sparsely pub- 

erulous or also with sparse to dense arachnoid 

hairs; stipules 0.4-0.6 cm long, appressed-pub- 

erulous to strigillose to subsericeous (to subhir- 

51


I 

1 cm 

FIG. 24. Coussapoa curranii: 1, leafy twig with staminate inflorescences 

(Glaziou 8934). 

sute). Staminate inflorescences branched; heads 

several to many, globose, ca. 2 mm diam.; com- 

mon peduncle 1-5 cm long, sparsely to densely 

puberulous or glabrous; perianth ca. 1 mm high, 

sparsely minutely puberulous; stamens two, ex- 

ceeding the perianth. Pistillate inflorescences 

branched; heads 3-6, globose, ca. 4-6 mm diam.; 

common peduncle 1-4 cm long, patent to ap- 

pressed puberulous; perianth ca. 1-2 mm high, 

minutely puberulous. Interfloral bracts sub- 

spathulate to subpeltate, puberulous at the apex. 

Distribution (Fig. 7). Eastern Brazil, from Rio 

de Janeiro to Bahia; in moist forests up to 

500 m. 

Specimens studied. BRAZIL. Without locality, 2 Dec 

1968 (8), Gomes et al. 2818 (RB). BAHIA: Rio Gongoji, 

1 Oct-20 Nov 1915 (9), Curran 8 (RB (HJBR 13070), 

US, type collection). EsPiarro SANTO: Res. Flor. de 

Linhares, 24 May 1978 (d), DAF 004 (GUA). MINAS 

GERAIS: Mun. Tombos, Fazenda da Cachoeira, 29 Jul 

1935 (9), Mello Barreto 1795 (F). Rio DE JANEIRO: 

Corcovado, 1857 (2), Casaretto 617 (G); Givea-Sao 

Conrado, Jun 1960 (2), Duarte 5239 (M, NY, RB, US); 

Serra da Estrella, Petr6polis, 21 May 1877 (a), Glaziou 

8934 (B, BR, G, K, LE, MO, P, U, US, type collection 

of C. warburgiana); Corcovado, 29 Jun 1876 (st), Glaziou 

8934a (P); Rio de Janeiro, Tijuca, Jan 1837 (2), 

Guillemin 1339 (P); Rio de Janeiro, Jardim Botanico, 

3 Sep 1926 (2), Kuhlmann s.n. (U); Cordeira, Fazenda 

Santa Clara, Feb 1970 (6), Lisboa s.n. (R); Rio de Janeiro, 

Jardim BotAnico, 29 Jul 1922 (8), Occhioni et 

al. (HJBR) 5864 (GUA, RB, U), 25 Jul 1922 (9), Occhioni 

et al. (HJBR) 5917 (RB). 

Coussapoa curranii is very closely related to 

C. floccosa. Affinities of these two species with 

other Coussapoa are not clear.


14. Coussapoa duquei Standley, Publ. Field Mus. PASTAZA: Rd. Hollin-Loreto-Coca, between Rio Pu- 

Bot. 22: 71. 1940. Type. Colombia. Caldas: cuno and Cacerio de Guamani, 1200 m, 12 Dec 1987 

Palestina, 1500 m, 15 Jun 1938 

(6), Cer6n M. 2955 

(d), Duque- 

(BG); nr. Mera, ca. 1400 m, 22 

Nov 1938 (9), Schultze-Rhonhof3020 (B, type collec- 

Jaramillo 1767 (holotype, F; isotypes, A, K, tion of C. apoda). PICHINCHA: Rd. Nanegalito-Pacto, 

NY, US). Fig. 25. 5 km W of Tulipe, 1300-1500 m, 22 Jul 1980 (6), 

Holm-Nielsen et al. 24519 (BG). 

Coussapoa apoda Mildbraed, Notizbl. Bot. Gart. Berlin 

15: 784. 1942. Type. Ecuador. Pastaza: Nr. Mera, C. duquei is closely related to C. villosa, esca. 

1400 m, 22 Nov 1938 (9), Schultze-Rhonhof3020 pecially to the form with brown arachnoid in- 

(holotype, B). 

dumentum (see p. 104). The species is distinct in 

the shortly pedunculate pistillate inflorescence 

with an ellipsoid to clavate head. This taxon could 

be regarded as distinct only at the subspecies 

level. 

Tree, (secondarily?) terrestrial, up to 30 m tall. 

Leafy twigs 5-10 mm thick, hirtellous to hirsute. 

Lamina coriaceous, elliptic to ovate, 8-18 x 5- 

11 cm, apex obtuse to acute to shortly acuminate, 

base subacute to obtuse (or to subcordate), margin 

entire to subcrenate; upper surface glabrous, 

lower surface puberulous to hirtellous on the 

veins, the whole surface densely covered with 

(sub)persistent pale yellow-brown arachnoid 

hairs; lateral veins 10-14 pairs, straight, main 

basal pair (sometimes faintly) branched, reaching 

the margin below the middle of the lamina; intercostal 

venation prominent; petiole 2-10 cm 

long, (rather) sparsely puberulous, with whitish 

to yellowish-brown arachnoid hairs which are 

also sometimes deciduous; stipules 2-7 cm long, 

white to brown strigose to hirsute and with reddish-brown 

arachnoid hairs. Staminate inflorescences 

branched; heads many, globose to oblate, 

ca. 4-8 mm diam.; common peduncle 1-2 cm 

long, puberulous to hirtellous, sometimes also 

with arachnoid hairs; perianth ca. 1 mm high, 

yellowish-brown puberulous; stamens two, far 

exceeding the perianth. Pistillate inflorescences 

unbranched; heads ellipsoid to clavate, ca. 1.5 

x 1 cm; peduncle 0.5-0.8 cm long, puberulous 

to hirtellous hairs, sometimes also with arachnoid 

hairs; perianth ca. 1-2 mm high, densely, 

minutely puberulous. Interfloral bracts (sub) 

spathulate, puberulous at the apex. 

Distribution (Fig. 7). Andean region (Colombia 

and Ecuador); in (sub)montane forest, between 

1200 and 2000 m. 

15. Coussapoa echinata Akkermans & C. C. Berg, 


449. 1982. Type. Panama. Panama: Cerro Jefe, 

Altos de Pacora, Rio Diablo, 15 km NE of 

Cerro Azul village, 5 Jan 1975 (9), Gentry & 

Mori 13408 (holotype, U; isotype, MO). 

Fig. 26. 

Tree, hemi-epiphytic. Leafy twigs 4-6 mm 

thick, glabrous or sparsely appressed-puberulous. 

Lamina coriaceous, oblong to elliptic to 

subobovate, 9-20 x 4-10 cm, apex acute to acuminate, 

base acute, margin entire; upper surface 

glabrous, lower surface (purplish or reddish), glabrous 

or on the veins sparsely puberulous; lateral 

veins 4-5 pairs, slightly curved, basal pair (faint- 

Specimens studied. COLOMBIA. ANTIOQUIA: Medellin, 

ca. 1500 m, - ly) branched, reaching the margin above the middle 

of the lamina; intercostal venation slightly 

prominent to plane; petiole 2-6 cm long, glabrous; 

stipules 1.5-2.5 cm long, sparsely appressed-puberulous. 

Staminate inflorescences 

branched; heads numerous, ca. 2 mm diam.; 

common peduncle ca. 1.5-2 cm long, puberulous; 

perianth ca. 1 mm high, glabrous; stamens 

three. Pistillate inflorescences branched; heads 3- 

8, globose, ca. 4-6 mm, in fruit up to 8 mm 

diam.; common peduncle 1-3.5 cm long, with 

sparse patent short hairs, occasionally mixed with 

some longer hairs; flowers free, perianth ca. 1-2 

mm high, (when dry) with an acute apex, gla- 

(st), Toro 120c 

brous; 

(US). CALDAS: 

fruiting perianth orange. Interfloral bracts 

Cord. Central, Palestina, 1500 m, 15 Jun 1938 (8), narrowly spathulate, short, sparsely, minutely 

Duque-Jaramillo 1767 (A, F, K, NY, US, type collec- puberulous. 

tion). NORTE DE SANTANDER: Ocafa, Jun 1845 (9), Pur- Distribution (Fig. 7). Panama (Panama and 

die s.n. (K). 

San 

ECUADOR. NAPO: 

Bias); in wet 

SE 

forest, 

of Borja, ca. 

up to 800 m. 

1900 m, 3 Aug 

1960 (d), Grubb et al. 1205 (K, NY); NE of Borja, ca. Specimens studied. PANAMA. PANAMA: Road El 

1800 m, 15 Aug 1960 (9), Grubb et al. 1283 (K, NY); Llano-Carti, km 17-20, 8 Mar 1974 (9), Dressier 4636 

nr. Borja, 1650 m, 16 Aug 1975 (9), Little et al. 212 (F); Cerro Jefe, Altos de Pacora region, Rio 

(Q), 1850 

Diablo, 

m, 16 Aug 1975 (8), Little et al. 213 (Q). 15 km E of Cerro Azul village, 5 Jan 1975 (2), Gentry

54 

2 ~ ? I 

wo t :1cm 


FIG. 25. Coussapoa duquei: 1, leafy twig with pistillate inflorescences (Grubb et al. 1283); 2, staminate 


(Grubb et al. 1205).

Coussapoa SS 

FIG. 26. Coussapoa 

'1 R~~~~~~~~~~~~~~~~~~~~~~~~? 

: e' 

r ]; e 

echinata: 1, leafy twig with pistillate inflorescences 

4636).i? ?. 

(Dressler


G. 27. oussapoaferruginea: 1, leafy twig with staminate inflorescences (Irwin am et 

FIG. 27. Coussapoa ferruginea: 1, leafy twig with staminate inflorescences (Irwin et al. 48631). 

& Mori 13408 (MO, U, type collection). SAN BLAS: 

Nusagandi, rd. to Carti, 18 Jul 1984 (a), McDonagh et 

al. 114 (MO). 

The species shows similarities to C. parviceps, 

from which it is distinct in the shape of the leaves, 

the free pistillate flowers, and the presence of 

interfloral bracts in the pistillate heads. On the 

other hand, C. echinata shows similarities to C. 

herthae, especially in the acute apex of the fruit- 

ing perianth (in dry material). 

16. Coussapoa ferruginea Trecul, Ann. Sci. Nat. 

Bot. Ser. 3, 8: 93. 1847; Miquel, Fl. bras. 4(1): 

137. 1853; Berg, Fl. Suriname 5(1): 286. 1975. 

Type. French Guiana. Without locality, 1838 

(6), Leprieurs.n. (holotype, P; isotypes, G, GH, 

L, U). Fig. 27. 

Tree, hemi-epiphytic. Leafy twigs 3-7 mm 

thick, brown(ish) (sub)hirsute. Lamina coria- 

ceous, lanceolate to oblong, 2-8 x 1-4 cm, apex 

acute to subobtuse, base acute, margin entire; 

upper surface glabrous, lower surface appressed- 

puberulous to strigose on the main veins, pu- 

berulous to hirtellous (to subhirsute) on the 

smaller veins, the whole surface covered with 

dense persistent brown arachnoid hairs; lateral 

veins 4-5(-6) pairs, straight, basal pair unbranched, 

reaching the margin below the middle 

of the lamina; intercostal venation prominent; 

petiole 0.5-1.5 cm long, densely brown hirtellous 

to subhirsute; stipules 1.5-3 cm long, brown 

(sub)hirsute to strigose or partly whitish subsericeous. 

Staminate inflorescences branched; heads 

several to many, ca. 2-3 mm diam.; common 

peduncle ca. 1 cm long, densely hirtellous; peri- 

anth ca. 1 mm high, densely puberulous; stamens 

two, far exceeding the perianth. Pistillate inflorescences 

unknown. Interfloral bracts spathulate, 

puberulous. 

Distribution (Fig. 6). French Guiana and the 

adjacent part of Amapa (Brazil); in riverine forest. 

Specimens studied. FRENCH GUIANA. Without 

locality, 1838 (d), Leprieur s.n. (F, G, GH, L, P, R, U, 

type collection), 1840 (8), Leprieur s.n. (FI, G, GH, K, 

P). 

BRAZIL. AMAPA: Rio Oiapoque, between mouth of 

Camopi River and Cachoeira Camaraua, 3 Oct 1960 

(d), Irwin et al. 48631 (F, K, MO, NY, U, US). 

The species is similar in many characters to 

C. crassivenosa. However, it is distinct from the


FIG. 28. Coussapoafloccosa: 1, leafy twig with pistillate inflorescences 

(Irwin 2058). 

latter in having a narrower lamina, a smaller 

number of lateral veins, and a relatively short 

petiole. Pistillate inflorescences are needed to decide 

whether this taxon is distinct from C. crassivenosa 

at the specific or only intraspecific level. 

17. Coussapoa floccosa Akkermans & C. C. Berg, 


451. 1982. Type. Brazil. Minas Gerais: Vi9osa, 

16 Nov 1935 (2), Kuhlmann 2074 (holotype, 

RB; isotypes, RB, U, US). Fig. 28. 

Tree or shrub, (mostly?) hemi-epiphytic. 

Leafy twigs 10-15 mm thick, (rather) densely 

57 

puberulous to (sub)hirsute. Lamina coriaceous, 

elliptic to ovate or obovate, 10-25 x 6-17 cm, 

apex rounded to obtuse or to emarginate, base 

obtuse to rounded to cordate, margin entire; upper 

surface often densely covered with white 

arachnoid hairs which later disappear, lower surface 

hirtellous to subtomentellous on the veins, 

the whole surface sparsely to densely covered 

with white arachnoid hairs, later deciduous; lateral 

veins 7-10 pairs, (main) basal pair distinctly 

branched, reaching the margin at or above the 

middle of the lamina; intercostal venation very 

prominent; petiole 1.5-5 cm long, tomentose to 

densely hirtellous; stipules 1-2.5 cm long, white 

to brownish appressed-puberulous to strig(ill)ose


to subsericeous (to subhirsute), sometimes also mm high, glabrous. Interfloral bracts (sub)peltate, 

with sparse arachnoid hairs. Staminate inflores- nearly glabrous. 

cences branched; heads numerous, globose, ca. 4 Distribution (Fig. 7). Central America: Costa 

mm diam.; common peduncle 1.5-4 cm long, Rica (Puntarenas), Nicaragua (Bluefields and Rio 

puberulous; perianth ca. 1 mm high, puberulous; San Juan), Panama (San Blas); in wet (riverside 

stamens two, exceeding the perianth. Pistillate or swamp) forest. 

inflorescences branched; heads 2-3, partly fused, 

(sub)globose, 5-7 mm diam.; common peduncle 

Specimens studied. NICARAGUA. BLUEFIELDS: Rio 

1-3 cm long, densely puberulous; perianth 1-2 

Kama, between Kama and Canio Valentin, 10 Mar 

1966 (2), Proctor et al. 27080 (F, US, type collection). 

mm high, glabrous. Interfloral bracts sub- Rio SAN JUAN: 1 km NW of Rio Santa Cruz, 22 Feb 

spathulate, puberulous at the apex. 

1984 (2), Moreno 23629 (BG). 

Distribution (Fig. 7). Brazil (Minas Gerais, Vi- COSTA RICA. PUNTARENAS: Peninsula de Osa, 8 

qosa) in 

km S of 

forest. 

Rinc6n, 28 Feb 1965 (9), Jimenez 3016 (F). 

PANAMA. SAN BLAS: El Llano-Carti rd., km 19.1, 

Specimens studied. BRAZIL. MINAS GERAIS: Est. 1 Feb 1985 (2), Nevers et al. 4804 (WIS), 3 Nov 1985 

Exp. de Agua Limpa, Feb 1968 (2), Gomes et al. 2819 (st), Nevers et al. 6186 (BG). SAN BLAS-PANAMA: El 

(RB); Vi9osa, 10 Nov 1958 (2), Irwin 2058 (F, NY, 

Llano-Carti rd., 28 Aug 1982 (d), Hamilton et al. 1062 

R, US), 31 Aug 1934 (st), Kuhlmann 1832 (US), 1935 (BG). 

(st), Kuhlmann 2073 (RB, US), 16 Nov 1935 (2), Kuhlmann 

2074 (RB, U, US, type collection), 31 Aug 1934 

This species appears to be one of a group which 

(st), Kuhlmann 2117 (RB), 1935 (9), Kuhlmann 2207 also includes C. angustifolia and C. trinervia and 

(GUA), 8 Oct 1976 (9), G. Rodrigues et al. 877 (RB); is characterized by shortly petiolate, more or less 

Capolivinba, nr. Rio Novo, Sep 1895 (6), Schwacke distinctly obovate leaves with few lateral veins, 

11895 (B, P). 

and sparse indumentum. 

This species, known only from the surroundings 

of Vicosa, is very closely related to C. cur- 19. Coussapoa herthae Mildbraed, Notizbl. Bot. 

ranii; but the two taxa are sufficiently different Gart. Berlin 14: 29. 1938. Type. Ecuador. Pifor 

them each to be regarded as species in their chincha: San Carlos de los Colorados, 5 Oct 

own right. 

1935 (d), Schultze-Rhonhof 953 (holotype, B). 

Fig. 30. 

18. Coussapoa glaberrima W. Burger, Phytolo- 

gia 26: 422. 1973; Burger, Fieldiana Bot. 40: 

133, t. 22. 1977. Type. Nicaragua. Bluefields: 

Rio Kama, between Kama and Canio Valentin, 

10 Mar 1966 (2), Proctor et al. 27080 (holo- 

type, F; isotype, US). Fig. 29. 

Tree, hemi-epiphytic or terrestrial, small. 

Leafy twigs 2-5 mm thick, glabrous. Lamina 

subcoriaceous, obovate to narrow elliptic, 2-14 

x 1-6 cm, apex acute, base acute to obtuse, margin 

entire; both surfaces glabrous; lateral veins 

2-3 pairs, slightly curved, basal pair indistinctly 

branched, reaching the margin far above the 

middle of the lamina; intercostal venation slightly 

prominent; petiole 0.5-2 cm long, glabrous; 

stipules 1.2 cm long, glabrous. Staminate inflorescences 

branched; heads ca. 10-12, ca. 2 mm 

diam.; common peduncle 1.5-3 cm long, gla- 

brous; perianth ca. 1 mm high, glabrous; stamens 

two, just exceeding the perianth. Pistillate inflorescences 

poorly branched; heads 2-3, globose, 

ca. 6 mm, in fruit up to 15 mm diam.; common 

peduncle 2-4 cm long, glabrous; perianth ca. 1 

Tree, mostly hemi-epiphytic, up to 25 m tall. 

Leafy twigs 5-13 mm thick densely, yellow to 

brown, strigose to hirsute to subvillous. Lamina 

coriaceous, elliptic to ovate to obovate or to ob- 

long, 16-32 x 10-20 cm, apex acuminate to 

acute, base obtuse to rounded or occasionally 

subcordate, margin entire or subcrenate; upper 

surface glabrous, sparsely to rather densely pub- 

erulous to hirtellous or on the main veins to 

subsericeous, sometimes with white arachnoid 

hairs which soon disappear; lateral veins (4-) 

8-11 pairs, curved, basal pair distinctly to faintly 

branched, reaching the margin usually below, 

sometimes at or just above the middle of the 

lamina, sometimes also other lateral veins 

branched, a distinct submarginal vein in the lower 

part of the lamina; intercostal venation (often 

very) prominent; petiole (1-)3-9 cm long, puberulous 

to hirtellous (to subhirsute); stipules 2- 

6 cm long, densely reddish- to yellowish-brown 

subsericeous to subhirsute to subvillous. Staminate 

inflorescences branched; heads many, 

globose, 2-3 mm in diam.; common peduncle


1 cm 

FIG. 29. Coussapoa glaberrima: 1, leafy twig with pistillate inflorescences 

(Proctor et al. 27080). 

(1-?)3-7 cm long, puberulous to hirtellous; peri- 

anth ca. 1 mm high, glabrous; stamens three, far 

exceeding the perianth. Pistillate inflorescences 

uador and southern Colombia); in wet forest, up 

to 1200 m. 

branched; heads 6-10, sometimes partly fused, 

Specimens studied. COLOMBIA. NARuio: Bay of 

Tumaco, Rio 

(sub)globose, 5-9 mm, in fruit up to 13 mm 

Yanaje, 6 Jul 1955 (9), Romero-Castadiam.; 

ineda 5283 (COL). 

common peduncle 3-6 cm long, 2-3 mm thick, ECUADOR. CARCHI: Rio San Juan, nr. Chical, 12 

puberulous to hirtellous; perianth ca. 2 mm high, km below Maldonado, 1200 m, May 1978 (2), Madison 

minutely puberulous; fruiting perianth 

et al. 4764 

orange. 

(AAU, F, QCA, SEL, U); El Pail6n, 45 km 

below 

Interfloral bracts spathulate to Maldonado, 28 Nov 1979 (d), Madison et al. 

subpeltate, pub- 7101 (SEL, U). ESMERALDAS: Nr. Lita, W of Rio Lita, 

erulous at the apex. 

9 Feb 1981 (2), Berg 1259 (QCA, U). GUAYAS: Nr. 

Distribution (Fig. 7). Pacific coastal region (Ec- Bucay (=Grl. Elizalda), 8-15 Jun 1945 (9), CampE3674

60 

cm1 

2 I! 

1 


FIG. 30. Coussapoa herthae: 1, leafy twig with pistillate inflorescences (Schulze-Rhonhof 1953); 2, staminate 

inflorescences (Dodson 5776).


(NY, U). IMBABURA: Parambas, 30 May 1949 (9), Acosta 

Solis 12661 (F). Los Rios: Rio Palenque Biol. St., 

km 56 on road Quevedo-Santo Domingo de los Colorados, 

27 Feb 1975 (6), Dodson 5776 (MO, QCA), 

29 May 1976 (2), Dodson 6082 (MO), 20 Mar 1980 

(a), Dodson et al. 9539 (SEL, U), 13 Feb 1974 (6), 

Gentry 9892 (MO, U). PICHINCHA: 35 km N of Santo 

Domingo de los Colorados, nr. bridge over Rio Blanco, 

3 Feb 1974 (2), Gentry 9606 (MO, U); San Carlos de 

los Colorados, 5 Oct 1935 (6), Schultze-Rhonhof 1953 

(B, type collection). 

Coussapoa herthae shows general similarity to 

C. nymphaeifolia and in the echinate (dry) fruiting 

head to C. echinata. 

The species is very variable in the shape and 

venation of the leaves. 

20. Coussapoa latifolia Aublet, Hist. pl. Guiane 

2: 955, t. 362. 1775; Trecul, Ann. Sci. Nat. 

Bot. Ser. 3, 8: 94. 1847; Miquel, Fl. bras. 4(1): 

135. 1853; Berg, Fl. Suriname 5(1): 280. 1975. 

Type. French Guiana. Without locality, - (9), 

Aublet s. n. (holotype, BM). Fig. 31. 

Coussapoa obovata Miquel, Het Instituut (1842): 200, 

t. 3. 1843. Type. Surinam. Plantation Bergendaal, - 

(2), Focke 9 (holotype, L). 

Coussapoa latifolia Aublet var. obovata (Miquel) Miquel, 

Fl. bras. 4(1): 135. 1853. 

Coussapoa froesii Standley, Publ. Field Mus. Bot. 17: 

162. 1937. Type. Brazil. Maranhao: Rio Maracassume 

region, 14 Sep 1932 (2), Fr6es 1907 (holotype, 

NY; isotypes, A, G, K, U). 

Shrub or tree, mostly hemi-epiphytic, up to 35 

m tall. Leafy twigs 4-9 mm thick, glabrous. Lamina 

coriaceous or subcoriaceous, ovate to elliptic, 

sometimes oblong to obovate, occasionally to 

suborbicular, 4-18(-25) x diam.; common peduncle 0.5-3 cm long, sparsely 

to densely appressed-puberulous; perianth ca. 

0.5-1 mm high, glabrous; stamens two, just exceeding 

the perianth. Pistillate inflorescences 

branched; heads 2-15, globose, ca. 3-4 mm, in 

fruit up to 8 mm diam.; common peduncle 1-3 

cm long, sparsely to densely appressed-puberulous; 

perianth ca. 1-1.5 mm high, glabrous. 

Interfloral bracts small, subpeltate to (sub) 

spathulate, minutely puberulous at the apex. 

Distribution (Fig. 7). Western Guiana region 

(Surinam and French Guiana), in Brazil from 

Amapa through eastern Para to Maranhao; further, 

apparently segregated from the main area 

in the central part of the Amazon Basin; forest 

of terra firme. 

Specimens studied. SURINAM. Brownsberg, 3 Sep 

1915 (9), BW(Stahel & Gonggrijp) 732 (K, MO, NY, 

P), 19 Sep 1931 (9), Van Emden s.n. (F, U, US), 23 

Sep 1931 (st), Van Emden LVI (U); Wilhelmina Mts., 

upper Gran Rio, Maup6 dam, 20 Feb 1926 (st), Exp. 

Wilhelmina Gebergte 196 (U); without locality, 

3-15 cm, apex acute 

to shortly acuminate to obtuse to rounded, base 

acute to obtuse or to subcordate, margin entire, 

plane or more or less revolute towards the base; 

upper surface glabrous, lower surface glabrous or 

sparsely appressed-puberulous with hairs of about 

equal length; lateral veins 4-7 pairs, (slightly) 

curved, basal pairs mostly branched, sometimes 

(especially) in narrow leaves unbranched, reaching 

the margin at to below or (especially in obovate 

leaves) above the middle of the lamina, 

the other lateral veins sometimes poorly branched 

(furcate); intercostal venation (almost) plane; 

petiole 1-7 cm long, sparsely to rather densely 

minutely appressed-puberulous; stipules 0.5- 

1.5(-2.5) cm long, sparsely to densely whitish to 

brownish appressed-puberulous, sometimes 

brownish subvelutinous. Staminate inflorescences 

branched; heads many, globose, ca. 1 mm 

- (2), 

Focke s.n. (GH); plantation Bergendaal, - (9), Focke 

9 (L, type collection of C. obovata = C. latifolia var. 

obovata); plantation Blauberg, - (9), Focke 418 (U); 

Distr. Nickerie, ca. 20 km SW of Avanavero dam site, 

14 Nov 1976 (9), Heyde et al. 78 (U); Wilhelmina Mts., 

Lucie River, 2-5 km below confluence with Oost River, 

8 Sep 1963 (9), Irwin et al. 55487 (MO, NY, OXF, R, 

S, U, US); Lely Mts., 19 Sep 1975 (st), Lindeman et 

al. 33 (U), 19 Sep 1975 (8), Lindeman et al. 73 (U), 4 

Oct 1975 (9), Lindeman et al. 749 (K, U), 5 Oct 1975 

(st), Lindeman et al. 799 (U); Jodensavanne-Mapane 

Creek area, camp 8, 21 Dec 1954 (st), Lindeman 6938 

(U); Coppename River, Doksi Creek, 11 Oct 1954 (6), 

Mennega 271 (C, NY, U); Tanjimama River, Kodji 

Creek, 25 Nov 1954 (6), Mennega 521 (A, C, NY, U); 

Voltzberg, 22 Sep 1956 (st), Schulz (LBB) 7787 (U); 

Jodensavanne-Mapane Creek area, 5 May 1967 (st), 

Vreden (LBB) 11747 (U). 

FRENCH GUIANA. Without locality, - (2), Aublet 

s.n. (BM, type collection), Feb 1868 (2), Aubry-Lecompte 

s.n. (FI); Sinnamary, road to St. Elie, 22 Oct 

1981 (9), Billiet et al. 1108 (U); Montagne de Kaw, 13 

Dec 1954 (9), Cowan 38774 (F, K, NY, P, US); Mont 

Belvedere, 18 Nov 1974 (2), De Granville et al. B5191 

(CAY, U); Oyapock River, Myuli Creek, Les Trois 

Sauts, 15 Sep 1977 (6), Grenand 1478 (CAY, U); without 

locality, - 

(9), Leman s.n. (P); Cayenne, - (9), 

Martin s.n. (or 112) (K); without locality, - (9), Melinon 

s.n. (A, P); Maroni River, 1864 (9), Melinon 18 

(GH, K, LE, P); Antecume Pata, at confluence of Itany 

River and Maroni River, 14 Nov 1977 (2), Moretti 836 

(CAY); Kourou River, 3 km above Couy Creek, 19 

Sep 1967 (6), Oldeman B1338 (CAY, U); Cayenne, - 

(st), Poiteau s.n. (FI); Acarouany, 1857 (6), Sagot 1136 

(B, BM, G, GH, K, S, U); Sinnamary, road to St. Elie, 

15 Sep 1978 (6), Sastre 6130 (P, U). 

BRAZIL. AMAPA: "Gafcho area," 21 Oct 1979 (9), 

Austin et al. 7149 (U); Rio Araguari, between 1?26'N, 

61

FIG. 31. Coussapoa latifolia: 1, leafy twig with pistillate inflorescences (Ducke (HJBR) 18456); 2, staminate inflor


51?58'W and 109'N, 51?52'W, 11 Sep 1961 (2), Pires 

et al. 50864 (NY, OXF, P, R, U, US); Rio Araguari, 

nr. 1?11'N, 52?8'W, 1 Oct 1964 (6), Pires et al. 51409 

(F, GH, NY, U); Col6nia Torrao, 2?25'N, 51015'W, 29 

Aug 1962 (2), Pires et al. 52656 (NY, OXF, U, US). 

AMAZONAS: Road Manaus-Caracarai, km 97, 30 Aug 

1979 (e), Cid et al. 942 (U); road Manaus-Porto Velho, 

km 235, Igarape Acu, 24 Nov 1973 (d), Lleras et al. 

P19661 (F, M, MO, NY, P, S, U); road Manaus-Caracarai, 

km 159, 20 Sep 1974 (6), Prance et al. 22720 

(K, MO, NY, S, U, US). MARANHAO: Rio Maracassume 

region, 14 Sep 1932 (2), Frogs 1907 (A, G, K, 

NY, U, type collection of C. froesii). PARA: Col6nia 

Benjamin Constant, Braganoa, 21 Nov 1908 (2), 

Anonymous (HAMP) 9783 (G); Sta. Izabel, 10 Sep 1922 

(2), Ducke (HJBR) 18456 (RB); Col6nia Augusto Montenegro, 

Igarape Pitor6, 18 Sep 1958 (2), Frogs 34651 

(IAN); road Belem-Brasilia, km 93, 19 Sep 1959 (2), 

Kuhlmann et al. 263 (NY, US); road Belem-Brasilia, 

km 301-306, 7 Aug 1960 (2), Oliveira 985 (U); Rio 

Jari, Monte Dourado, 16 Nov 1967 (2), Oliveira 3550 

(NY), 9 Nov 1967 (2), Oliveira 3599(NY), 8 Nov 1967 

(9), Oliveira 3682 (NY), 23 Jan 1968 (st), Oliveira 3954 

(NY), road Capanema-Maranhao, km 96,27 Oct 1965 

(2), Prance et al. 1720 (MO, NY, S, U, US), km 98, 

21 Aug 1964 (2), Prance et al. 58783 (U); Peixe Boi, 

26 Oct 1907 (2), Seguiera (HAMP) 8808 (BM, G, U); 

Rio Jari, Monte Dourado, 15 Oct 1968 (d), N. T. Silva 

1207 (NY), 28 Oct 1968 (2), N. T. Silva 1321 (NY, U, 

US); Rio Jari, between Pilao and Repartimento, 12 

Nov 1968 (2), N. T. Silva 1380 (IAN, NY, U, US). 

Coussapoa latifolia is very closely related to 

C. viridifolia. The species shows affinities with 

C. microcephala, C. microcarpa, C. pachyphylla, 

and also with the Central American C. macering 

the margin at or below the middle of the 

lamina, the other lateral veins sometimes 

branched; intercostal venation prominent; petiole 

1.5-3 cm long, puberulous to hirtellous; stipules 

0.5-1 cm long, yellowish to brown hirsute 

to subsericeous; terminal buds often more or less 

swollen. Staminate inflorescences branched; 

heads many, globose, ca. 6 mm diam.; common 

peduncle 4-7 cm long, puberulous to hirtellous; 

perianth ca. 1 mm high, glabrous; stamens three, 

far exceeding the perianth. Pistillate inflorescences 

unbranched; heads globose, ca. 5-8 mm 

diam.; peduncle 0.5-2.5 cm long, puberulous to 

hirtellous; perianth ca. 1-2 mm high, glabrous. 

Interfloral bracts spathulate to subpeltate, puberulous. 

Distribution (Fig. 7). In (eastern) French 

Guiana and in Brazil in eastern Para; in upland 

and riverine forest. 

Specimens studied. FRENCH GUIANA. Without 

locality (3), Leprieur s.n. (P, type collection); Approuague 

River, between Sapokaye Creek and Grand 

Canori Falls, 23 Oct 1968 (6), Oldeman T243a (CAY); 

Sapkaye Creek, 22 Oct 1968 (3), Oldeman B1946 (CAY, 

K, P, U); Approuague River, Mapaou Falls, 26 Jan 

1970 (6), Oldeman B2861 (CAY, U); Oyapock River, 

3 km above Camopi, 8 Apr 1970 (6), Oldeman 3062 

(CAY, P, U). 

BRAZIL. PARA: Ilhas de Breves, Furo Macujubim, 

16 Nov 1912 (6), Ducke (HJBR) 18462 (RB); Alto 

Quatipuir, Dec 1899 (6), Huber(HAMP) 1775 (F, MG). 

rima. Because of similarities in the vegetative 

parts, C. latifolia can be confused with C. orthoneura, 

especially specimens with elliptic leaves. 

Coussapoa leprieurii, one of the few species in 

which the lamina is scabrous above, appears to 

be rather closely related to C. sprucei. Specimens 

with an elliptic to oblong lamina having arachnoid 

hairs beneath are very similar to C. sprucei. 

Specimens with obovate leaves are reminiscent 

of (the unrelated) C. asperifolia ssp. asperifolia. 

21. Coussapoa leprieurii Benoist, Bull. Mus. Hist. 

Nat. Paris 30:103. 1924. Type. French Guiana. 

Without locality, - (6), Leprieur s.n. (holo- 

type, P). Fig. 32. 

Tree, hemi-epiphytic, up to 10 m tall. Leafy 

twigs 5-8 mm thick, brown(ish) puberulous to 

hirtellous to hirsute, solid. Lamina coriaceous, 

(broadly) ovate to (broadly) elliptic to (broadly) 

obovate to oblong, 5-18 x 1.5-12 cm, apex ob- 

tuse to subacuminate to acute, base subcordate 

to obtuse or acute, margin entire, often more or 

less revolute, especially towards the base; upper 

surface scabrous, lower surface puberulous, on 

the main veins to hirtellous, sometimes also white 

arachnoid hairs present; lateral veins 6-7 pairs, 

straight or curved, basal pair unbranched, reach- 

22. Coussapoa longepedunculata Akkermans & 

C. C. Berg, Proc. Kon. Ned. Akad. Wetensch. 

Ser. C. 85(4): 453. 1982. Type. Peru. Loreto: 

Prov. Maynas, Quebrada Sucusari, Llachapa 

camp of Explorama, N side of Rio Napo, below 

Mazan, 6 Nov 1979 (6), Gentry et al. 27566 

(holotype, U; isotype, MO). Fig. 33. 

Tree, up to 25 m tall, hemi-epiphytic or terrestrial. 

Leafy twigs 1-1.5 cm thick, brown hirsute. 

Lamina coriaceous, oblong to elliptic, 15- 

50 x 7-30 cm, apex obtuse, base cordate to 

rounded margin repand to subcrenate; upper sur-


FG2 3,si1e tg ts iore s l a 1 cm 

FIG. 32. Coussapoa leprieurii: 1, leafy twig with staminate inflorescences (Oldeman B1946); 2, pistillate 

inflorescences (Oldeman 3062). 

face scabridulous to smooth, sparsely hispidu- 

lous to hirtellous, often ? bullate, lower surface 

sparsely hirtellous on the veinlets, between the 

lateral veins white arachnoid hairs, rather dense 

or sparse and then concentrated at the margin; 

lateral veins 9-11 pairs, more or less curved, at 

least the lower ones branched, basal pair reaching 

the margin far below the middle of the lamina; 

intercostal venation prominent; petiole 3-10 cm 

long, brown hirsute; stipules 2-6 cm long, some- 

times subpersistent, hirsute to subsericeous, with 

brown hairs of different lengths. Staminate in- 

florescences rather poorly branched; common 

peduncle 9-16 cm long, the branches in two clusters 

and all or some of them very short, peduncle 

and branches brown hirtellous to pubescent; 

heads many, (sub)globose, 3-5 mm diam.; perianth 

ca. 1 mm high, sparsely minutely puberulous; 

stamens three, exceeding the perianth. Pistillate 

inflorescences branched; heads 3-10, 

(sub)globose, 5-8 mm, in fruit up to 15 mm diam.; 

common peduncle 8-20 cm long, peduncle and 

branches brown hirtellous to subtomentose; perianth 

ca. 1 mm long, glabrous; fruiting perianth 

red. Interfloral bracts ca. 1 mm long, spathulate, 

subpeltate, at the apex puberulous.


?~ 

?~~~~~~~~~~~~~~~~~~~~~~~~~~~L 

~ ' ~/ ~ 1o g . 

' ', 

_ i , ., 

'? 

... ~ ~ ~ .'. ~ ~ ~~~~~~~~~~. 

,... 

: ~. - 

"' 

-r . . ' - ..; 

???~ ~~~~~~~~~~' 

? 

.' 

,, - 

? 

?' 

. . 

.. ? . . ... . 

or ?- -.' . 

*~~ -, ~ ~~~~ -..s 

??.? 

... ,, 

~~~~~~~~~~~~~~~~~~... 

-~~~~~~~~~~~~ : 

.'? ~ ..- ~ ~ *-. .I 

~~ ,. .... 

. ''' =, '? IY~~~ 

~~~~~~ .~~~~~~~~~~~. 

I' 

r~~~~~~~~~~~~~~~~. o o- 

..,. 

I '~'~~~~~~~~~~~~~~~~~~~~~' 

~'C ?1 .'E":? . 

..? 

", 

'~~~~~~~~' 

.' 

-?~~~~1 

?~~~~~~~~~? 

FIG 33 

c. 

Cossaoa 

? 

onepeunclat: 1 lafytwi wih samiat iniorsceces(Gntr etal.2756) 

''. 

..


Distribution (Fig. 8). Peru (Loreto) and Ec- 

uador (Napo); in non-inundated or periodically 

inundated forest. 

Specimens studied. ECUADOR. NAPO: Parque Nacional 

Yasuni, 16-19 Jan 1988 (d), Ceron M. 3495 

(BG), 9-19 Jan 1988 (9), Neill et al. 8289 (BG). 

PERU. LORETO: Prov. Maynas, Quebrada Sucusari, 

Llachapa camp of Explorama, N side of Rio Napo, 

below Mazan, 6 Nov 1979 (a), Gentry et al. 27588 (MO, 

U, type collection); Prov. Maynas, Puerto Almendras, 

12 Jan 1982 (9), Vdsquez 2852 (BG); Iquitos, Rio Itaya, 

Buena Suerte, 16 Nov 1986 (9), Vdsquez et al. 8380 

(BG); Nauta, 12 Dec 1986 (9), Vdsquez et al. 8600 (BG). 

C. longepedunculata resembles C. nymphaeifolia 

in many features. It is distinct from the 

latter especially in the indumentum, the wider 

(looser) venation, and in the scabridulous upper 

surface of the lamina. The species also shows 

affinities with C. sprucei and C. leprieurii. 

and allied species. In the shape and venation of 

the leaves it resembles C. contorta, from which 

it clearly differs in the 2-staminate flower. 

24. Coussapoa manuensis C. C. Berg, Proc. Kon. 

Ned. Akad. Wetensch. Ser. C, 86(3): 305. 1983. 

Type. Peru. Madre de Dios: Parque Nacional 

del Manu, Rio Manu, Cocha Cashu Station, 

26 Oct 1980 (v), Foster 5641 (holotype, F). 

Fig. 35. 

Tree, hemi-epiphytic, 25 m tall. Lamina coriaceous, 

broadly elliptic to obovate, 10.5-19 x 

8-13 cm, apex shortly acuminate, base rounded 

to subcordate, margin entire to subcrenate; upper 

surface glabrous or sparsely puberulous at the 

base of the midrib, lower surface densely puberulous 

to hirtellous (to subtomentose); lateral 

veins 8-12 pairs, almost straight, the basal pair 

unbranched, reaching the margin below the mid- 

23. Coussapoa macerrima Akkermans & C. C. dle of the lamina; intercostal venation more or 

Berg, Proc. Kon. Ned. Akad. Wetensch. Ser. less prominent; petiole 3-5 cm long, puberulous 

C, 85(4): 455. 1982; Burger, Fieldiana Bot. 40: to hirtellous; stipules ca. 6 cm long, chartaceous, 

133, t. 22. 1977 (sub C. contorta). Type. Costa subsericeous (to subhirsute), with rather con- 

Rica. Puntarenas: Above Palmar Norte de Osa, spicuous (parallel) venation. Staminate inflores- 

22 Feb 1951 (d), Allen 5949 (holotype, F; iso- cences unknown. Pistillate inflorescences untypes, 

A, F). Fig. 34. branched; heads globose, ca. 4 mm, in fruit up 

Tree, up to 25 m tall. Leafy twigs 2-4 mm 

to 12 mm diam.; peduncle 2.2-2.7 cm long, puthick, 

sparsely appressed-puberulous, on the scars berulous; perianth ca. 1 mm high, sparsely miof 

the stipules distinctly longer hairs. Lamina nutely puberulous; fruiting perianth yellow. Insubcoriaceous, 

elliptic to oblong to subobovate, 

terfloral bracts spathulate, puberulous at the apex. 

5-13 x 4-7 cm, apex shortly acuminate, base 

Distribution (Fig. 8). Peru (Madre de Dios). 

acute to obtuse, margin entire; upper surface 

Known 

glaonly 

from the type collection. 

brous, lower surface on the main veins rather 

C. manuensis shows no distinct affinities with 

sparsely appressed-puberulous with hairs of dif- any of the other Coussapoa species. 

ferent lengths or partly strigillose; lateral veins 

25. 

7-13 pairs, straight to curved, basal pair un- 

Coussapoa microcarpa (Schott) Rizzini, Dusenia 

branched reaching the margin below the middle 

1(5): 295. 1950. Fig. 36. 

of the lamina; intercostal venation plane; petiole Brosimum microcarpon Schott in Sprengel, Syst. Veg. 

2-5 cm long, sparsely appressed-puberulous with 4 (Curr. Post., App.): 403. 1827. Type. Brazil. Withhairs 

of different 

out 

lengths; stipules 1-2 cm locality, 

long, (sub)sericeous. Staminate inflorescences 

branched; heads many, globose, ca. 1-2 mm 

diam.; common peduncle 2-3 cm long, appressed-puberulous; 

perianth ca. 1 mm high, glabrous; 

stamens two, far exceeding the perianth. 

Pistillate inflorescences unknown. Interfloral 

bracts absent. 

Distribution (Fig. 7). Costa Rica (Puntarenas); 

in forest at 450 m. Known only from the type 

collection. 

This species may be related to C. microcephala 

- (Y), Schott s.n. (holotype, W, destroyed). 

Coussapoa schottii Miquel, Fl. bras. 4(1): 137. 1853, 

as a synonym of Brosimum microcarpon. 

Coussapoa schottii Miquel var. lanceolata Miquel, Fl. 

bras. 4(1): 137. 1853. Type. Brazil. Without locality, 

- (Y), Pohl s.n. (holotype, U; isotypes, B, M (herb. 

Zuccarini 105 sub forma angustifolia)). 

Coussapoa schottii Miquel var. longifolia Miquel, Fl. 

bras. 4(1): 137. 1853. Type. Cult. bot. garden Minchen, 

1835 (2), herb. Zuccarini 103 (holotype, M). 

Coussapoa fontanesiana Trecul, Ann. Sci. Nat. Bot. 

Ser. 3, 8: 94. 1847. Syntypes. Brazil. Sao Paulo: 

Without locality, - (8 and 2), Gaudichaud 922 (lectotype 

(Q), P).


I 

1 cm 

FIG. 34. Coussapoa macerrima: 1, leafy twig with staminate inflorescences (Allen 5949) 

Shrub or tree, terrestrial or occasionally hemi- 

epiphytic, up to 20 m tall. Leafy twigs 2-5 mm 

thick, white to yellowish appressed-puberulous 

to hirtellous or to hirsute, sometimes also with 

white to brownish arachnoid hairs. Lamina co- 

riaceous, subovate to ovate oblong to elliptic to 

(sub)obovate or to lanceolate, 3-21 x 2-7 cm, 

apex shortly acuminate to acute or to obtuse, 

base acute to obtuse to rounded, margin entire, 

usually revolute towards the base; upper surface 

glabrous, lower surface glabrous or sparsely appressed-puberulous 

(to strigose) to (sub)hirsute

FIG.c3.m cousapoa anuenss 1 leves sipue n itlaeiforsecs(otr54) 

FIG. 35. Coussapoa manuensis: 1, leaves, stipules and pistillate inflorescences (Foster 5641


"-; ~~~~~~~~~~~~~~~~~~~~~~'? 

?: ' ~C .. ? 

~~~~~~~~~~~~~~.. U ? : 

?? 

: 7:I .i 

?~ ~~~~~~~~~~? 

.. 1~~~~~~~~~~~~~ 

?: ~~~~~~~~~~~~~~~. ~ ~ ~ ~ : .?? ,. 

~~~~~~~~~~~~~~~~.2 

~~?. ?'''" .:? 

.. ".' 

I- ".i ii{ 

' . .. ~ 

FIG. 36. Coussapoa microcarpa: l, leafy twig with pistillate inflorescences (Mexia 5115); 2, leafy twig with 

staminate inflorescence (Hage 31). 

on the main veins; lateral veins 6-11 pairs, 

straight, basal pair unbranched, reaching the 

margin below the middle of the lamina; intercostal 

venation plane to slightly prominent; petiole 

1-4 cm long, appressed-puberulous to hirtellous 

to (sub)hirsute; stipules 1-7 cm long, 

yellowish puberulous to subsericeous to hirsute 

or also with white to brownish arachnoid hairs. 

Staminate inflorescences branched; heads 5-9, 

globose, ca. 2-3 mm diam.; common peduncle 

1-2 cm long, puberulous to hirtellous; perianth 

ca. 1 mm high, minutely puberulous; stamens 

two, far exceeding the perianth. Pistillate inflo- 

rescences unbranched or rarely branched; heads 

1(-5), globose, ca. 3-5 mm, in fruit up to 10 mm 

diam.; (common) peduncle 2-5 cm long, pu-


berulous to hirtellous; perianth ca. 1 mm high, 

glabrous; fruiting perianth yellow and orange. Interfloral 

bracts small, narrowly spathulate, often 

only a few are sometimes absent. 

Distribution (Fig. 7). Brazil, from Rio Grande 

do Sul to Espirito Santo, common in forests up 

to 1800 m and in restinga vegetation; moreover, 

apparently in more or less isolated populations 

further north, in Bahia (near Ilheus) and in Paraiba 

and Pernambuco (on the top of low mountains?). 

Specimens studied. BRAZIL. Without (certain) locality, 

1814-1817 (9), Bowie & Cunningham 57 (BM), 

- (d and 2), Burchell 3148 (GH, K, P), - (6), Glaziou 

1942 (P), - (6), Glaziou 2016 (P), - (6), Glaziou 6009 

(C), - (9), Graham s.n. (K), - (2), Pohl s.n. (B, M 

(herb. Zuccarini 105), U, type collection of C. schottii 

var. lanceolata), 11 Nov 1822 (d), Riedel s.n. (LE), 5 

Oct 1863 (st), Warming s.n. (C), 1835 (d), (cult. in hort. 

bot. Munchen), herb. Zuccarini 103 (M, type collection 

of C. schottii var. longifolia). BAHIA: Nr. Ilheus, 1 Apr 

1965 (2), Belem et al. 644 (U); without locality, - Porto de Cima, 26 Jun 1914 (6), Jonsson 610a (F, GH, 

K, NY, S); Rio Sao Joao, W of road to Joinville, N of 

Garuva, 16 Jul 1966 (st), Lindeman et al. 1857 (U); 

Serra do Mar, nr. old road Curitiba-Morretes, Bella 

Vista, 25 Jul 1967 (st), Lindeman et al. 5682 (U); nr. 

Pontal do Sul, 20 Jun 1967 (st), Lindeman et al. 5748 

(U); Caioba, 35 km S of Paranagua, 10 Nov 1947 (9), 

Tessmann s.n. (MO); Mun. Guaraqueqaba, Serrinha, 

13 Apr 1967 (6), Tessmann s.n. (MO). PERNAMBUCO: 

"Gurgaf" (=Gurjao?), 14 May 1952 (8), Ducke & Lima 

104 (R). Rio DE JANEIRO: Rio de Janeiro, 

(9), 

Blanchet s.n. (BM, M, U); nr. Ilheus, 1836 (2), Blanchet 

2327 (G, M, P), 30 Nov 1970 (6), Emygdio 3018 (=Emmerich 

3556) (R); Itabuna, 16 Dec 1966 (9), Emygdio 

et al. 2447 (=Emmerich et al. 3002 = Andrade et al. 

2340) (R); nr. Ilheus, 23 Nov 1970 (6), Hage 31 (GUA, 

U), 24 Nov 1971 (2), Pinheiro 1716 (U); Castelo Novo, 

Oct 1821 (2), Riedel s.n. (LE), Mar 1822 (2), Riedel 664 

(LE). ESPiRITO SANTO: Mun. Linhares, Res. Biol. Sooretama, 

14 Mar 1972 (2), Sucre 8690 (U). MINAS GERMS: 

Lagoa Santa, - (6), Lund 139 (C); Viqosa, Fazenda de 

Aguada, 29 Sep 1930 (2), Mexia 5115 (A, BM, F, G, 

GB, GH, K, MO, NY, S, U); Lagoa Santa, - (?), 

Warming 1917 (C), 13 Oct 1863 (?), Warming 1918 

(C), 5 Oct 1863 (6), Warming 1919 (C), 29 Dec 1864 

(8), Warming 1942 (C). PARAiBA: Serro Araripe, Santa 

Ana, Aug 1921 (2), Luetzelburg 12471 (M). PARANA: 

Mun. Guaratuba, Garuva, 28 Jul 1960 (9), Duarte et 

al. 5328 (F, RB); Jacarehy, 27 Sep 1908 (st), Dusen 

6635 (GH, NY); Caita de Agua, 28 Aug 1910 (st), 

Dusen 10134 (NY); Morretes, 7 Sep 1910 (2), Dusen 

10186 (F, GH, K, L, MO, NY, P, S); Jacarehy, 25 Mar 

1911 (6), Dusen 11419 (F, GH, NY); Rio Cubatao, 28 

Dec 1911 (8), Dusen 13653 (F, GH, NY, S); Jacarehy, 

26 Mar 1914 (6), Dusen 14718 (F, GH, K, MO, NY, 

P), 11 May 1915 (st), Dusen 17030 (NY); Cerro Azul, 

2 Oct 1949 (6), Hatschbach 1475 (S); Mun. Paranagua, 

Rio Cachoeirinha, 25 Aug 1951 (6), Hatschbach 2458 

(RB); Mun. Paranagua, Porto D. Pedro 2?, 31 Jan 1961 

(8), Hatschbach 7807 (RB); Mun. Cerro Azul, Turvo, 

6 Feb 1961 (6), Hatschbach 7833 (RB); between Curitiba 

and Paranagua, 10 Nov 1948 (6), Hatschbach 

16304 (F, K); Mun. Guaraquecaba, Rio do Cedro, 8 

Nov 1968 (9), Hatschbach 18680 (C, MO, NY); Mun. 

Bocaiuva do Sul, Sesmaria, Rio Capivari, 11 Nov 1968 

(6), Hatschbach 20253 (C); Mun. Guaratuba, Rio Tupitinga, 

29 Apr 1972 (6), Hatschbach 29629 (NA); Paranagua, 

7 Mar 1914 (2), Jonnson 5a (F, GH, K, NY, S); 

- (2), Blanchet 

s.n. (NY), 1835 (8), Blanchet s.n. (FI); Itatiaia, 10 

Jan 1910 (st), Campos Porto 863 (RB); Rio de Janeiro, 

Collegio Anglo-Brasileiro, 22 Sep 1916 (2), Constantino 

(HJBR) 19689 (RB); Restinga de Itapeba, nr. Canal 

das Taxas, 22 May 1963 (2), Carauta 178 (GUA, 

K); Rio de Janeiro, Gavea, 11 Aug 1974 ($), Carauta 

1716 (F, GUA, K, U); Rio de Janeiro, Morro da Urca, 

17 Apr 1975 (st), Carauta 1780 (F), (2), Carauta 1781 

(GUA); Rio de Janeiro, Botanical Garden, 9 Nov 1978 

(2), Carauta 3050 (RB); Rio de Janeiro, Alto da Boa 

Vista, 7 Aug 1974 (2), Carauta et al. 1714 (F, GUA, 

RB); between Serra de Macah6 and Novo Friburgo, 

Sao Pedro, 18 Oct 1977 (2), Carauta et al. 2722 (GUA, 

U); Rio de Janeiro, Pao de Acucar, 26 Aug 1979 (2), 

Carauta et al. 3166 (GUA, U); Turin, 1857 (2), Casaretto 

643 (G); Joatinga Joa, 24 Mar 1959 ($), Duarte 

et al. 4650 (RB); Dois Irmaos, 19 Feb 1921 (2), Ducke 

et al. (HJBR) 16361 (RB); Meio Serra, old road to 

Petr6polis, 3 Nov 1928 (9), Ducke (HJBR) 21200 (RB); 

Serra de Itatiaia, Mont Serrat, 21 Oct 1903 (2), Dusen 

2160 (S); Rio de Janeiro, Prainha, 4 Dec 1978 (2), 

Ferreira 493 (RB); Serra dos Orgaos, Apr 1837 (6), 

Gardner 732 (BM, K); Rio de Janeiro, Gavea, Aug 

1837 (9), Gardner 5632 (BM, K); Teres6polis, 1100 m, 

31 Jan 1978 (9), Gentry et al. 918 (MO, U); Rio de 

Janeiro, Gavea, 19 Jan 1861 (st), Glaziou s.n. (P), 28 

Aug 1861 (st), Glaziou s.n. (P); Rio de Janeiro, Copacabana, 

4 May 1866 (9), Glaziou 1013 (C, NY, P); 

Rio de Janeiro, Gavea, 28 Apr 1867 (6), Glaziou 1138 

(C, P); Rio de Janeiro, - (st), Glaziou 1881 (G); Rio 

de Janeiro, Gavea, 28 Nov 1869 (6), Glaziou 4937 (P); 

Rio de Janeiro, Copacabana, 1871 (6), Glaziou 4938 

(C: K, P); Rio de Janeiro, Tijuca, 6 Feb 1871 (2), Glaziou 

6009 (K, P); Rio de Janeiro, 12 Aug 1876 (2), 

Glaziou 8936 (A, K, LE, P); Serra dos Orgaos, 23 Mar 

1880 (2), Glaziou 12166 (C, F, G, K, LE, P); Petr6polis, 

Bairro Amoeda, Dec 1943 (8), Goes et al. 815 (GUA, 

RB); Recreio dos Bandeirantes, 30 km W of Rio de 

Janeiro, 5 Mar 1964 (6), Lems s.n. (NY); Rio de Janeiro, 

1839 (6), Luschnath s.n. (LE); between Serra de 

Macah6 and Novo Friburgo, 1000 m, 18 Oct 1977 (2), 

Maas et al. 3325 (K, U); Petr6polis, Vale das Videiras, 

1800 m, 6 Jan 1973 (2), Martinelli 150 (U); Rio de 

Janeiro, 1867 (8), Miers s.n. (BM); Rio de Janeiro, 

Tijuca, 1879 (2), Miers s.n. (BM); Rio de Janeiro, Jul 

1878 (6), Miers 2714 (K, P); Rio de Janeiro, Tijuca, 

Jul 1878 (9), Miers 3792 (K); Rio de Janeiro, Pedreiro, 

Jul 1878 (6), Miers 3858 (K, P); Restinga de Jacarepagua, 

Res. Biol., 12 Apr 1967 (6), Moreira 46 (F, GH, 

GUA, US); Rio de Janeiro, Corcovado, 20 Sep 1974 

(8), Mosen 2614 (S); Rio de Janeiro, Sep 1862 (9), Nadeaud 

s.n. (P); Rio de Janeiro, Copacabana, Oct 1862


(a), Nadeaud s. n. (P); Rio de Janeiro, Botanical Garden, 

8 Mar 1936 (9), Occhioni (HJBR) 35333 (RB); Serra 

dos Orgaos, Rio Paquequer, 4 Mar 1949 (9), Pereira 

39 (RB); Rio de Janeiro, Botanical Garden, 23 May 

1961 (9), Pereira 5670 (B, M); Rio de Janeiro, Gavea, 

3 Oct 1927 (9), Pessoal do Horto Florestal 654 (RB); 

Rio de Janeiro, Tijuca, 14 Jun 1969 (st), Plowman 2919 

(=Sucre 5209) (GH, K, US); Rio de Janeiro, 1829 (a), 

Riedel 7 (LE), Sep-Dec 1831 (9), Riedel 75 (LE, NY, 

RB, US), 1832 (st), Riedel et al. s.n. (LE), 1832 (st), 

Riedel et al. I (LE); Teres6polis, Rio Imbui, 2 Oct 1952 

(9), Rizzini 144 (RB); Belem, Oct 1867 (9), Schwacke 

1067 (RB); Rio de Janeiro, Corcovado, 2 Nov 1883 

(9), Schwacke 6783 (RB), 11 Mar 1883 (2), Schwacke 

8418 (R); Rio de Janeiro, Copacabana, 25 Oct 1967 

(9), Sucre 1783 (F, GUA, K, RB); Cabo Frio, Restinga 

do Per6, 14 Sep 1968 (9), Sucre 3633 (GUA, U); Restinga 

de Jacarapagua, 8 May 1969 (9), Sucre 4989 (U); 

Parades da Subida, Pedra da Panela, 3 Nov 1971 (9), 

Sucre 7878 (U); Rio de Janeiro, Gavea, Nov 1899 (8), 

Ule s.n. (R); Rio de Janeiro, Copacabana, - (9), Ule 

1693 (P); Itatiaia, 1935 (a), Zikan (HJBR) 29279 (RB). 

Rio GRANDE DO SUL: Fazenda das Almas, nr. Palmares, 

Jan 1945 (8), Buck (in Rambo) 26425 (B); P6rto 

Alegre, 1 Oct 1957 (9), Camargo 1849 (B); Morro Teres6polis, 

Dec 1940 (8), Leite 413 (A), Dec 1942 (8), 

Leite 872 (A); P6rto Alegre, 2 Jun 1893 (9), Malme 

826 (S), 26 Feb 1902 (6), Malme 1428 (R, S), 16 Oct 

1932 (d and 9), Rambo 426 (F, MO, S), 10 Nov 1946 

(9), Rambo 27092 (FI, S), 2 Oct 1948 (9), Rambo 37782 

(C), 1 Dec 1948 (9), Rambo 38427 (B); Lami, nr. Viamao, 

3 Jan 1949 (6), Rambo 39413 (F); Estacao Pereci, 

14 Jan 1949 (9), Rambo 39780 (P); Ilha das Flores, 22 

Apr 1949 (9), Rambo 41173 (C, K, US); Lagoa dos 

Barros, nr. Osoria, 14 Feb 1949 (9), Rambo 44743 (P); 

Fazenda do Arroio, nr. Osoria, 4 Jan 1950 (a), Rambo 

45136 (K); Lagoa dos Quadros, nr. Torres, 21 Feb 1950 

(a), Rambo 45904 (K); nr. Tramandai, 5 Mar 1950 (a), 

Rambo 46145 (P); without locality, - NY, S, U), 20 Dec 1952 (6), Reitz et al. 5030 (F, G, 

NY, S, U); Mina Velha, Garuva, Sao Francisco do Sul, 

8 Nov 1957 (9), Reitz et al. 5611 (B, US); Serra do 

Matador, Rio do Sul, 26 Jan 1959 (2), Reitz et al. 8294 

(G, M); Ilha da Santa Catarina, 6 Mar 1962 (8), Sehnem 

7993 (B); P6rto Belo, Ilha Joao da Cunha, 31 Mar 1957 

(Q), L. B. Smith et al. 12307 (NY, R, US). SAO PAULO: 

Mun. Picinguaba, Picinguaba beach, 2 Oct 1975 (9), 

Araujo et al. 848 (RB); Mun. Iguape, Morro das Pedras, 

Aug 1915 (8), Brade 7887 (R), 1917 (9), Brade 7949 

(R, RB); Mun. Iguape, Iguape Island, 19 Feb 1965 (2), 

Eiten et al. 6214 (MO, US); without locality, 1917 (6), 

Frazas(HJBR) 13059 (RB); 1833 (2), Gaudichaud 992 

(P); Sao Paulo, Parque do Estado, 4 Sep 1933 (6), Hoehne 

30923 (F, GUA, NY, P); road Caraguatatuba-Ubatuba, 

10 Oct 1968 (9), Leitao Filho 685 and 686 (GUA); 

Santos, 1 Dec 1874 (2), Mosen 2941 (C, S). 

This species appears to be related to C. latifolia 

and C. microcephala. It is quite variable in the 

shape and dimensions of the lamina. A form with 

relatively long and narrow leaves is cultivated in 

several European greenhouses and (in The Netherlands) 

as an indoor ornamental. 

26. Coussapoa microcephala Tr6cul, Ann. Sci. 

Nat. Bot., Ser. 3, 8: 96. 1847; Miquel, Fl. bras. 

4(1): 136. 1853; Berg, Fl. Suriname 5(1): 282. 

1975. Type. Guyana. Pomeroon River, 

(9), Sello 241 

(B, P, US); Rio Tapuchy, 1837 (6), Tweedie 29 (K); 

Col6nia Sao Pedro Torres, 12 Nov 1968 ($), Vianna 

et al. 5465 (U). SANTA CATARINA: Rio Claro, Linha de 

Joinville, 30 Oct 1882 (6), Anonymus s.n. (RB); Ilha 

da Santa Catarina, 8 Dec 1950 (9), Duarte et al. 3410 

(F, K, NY, RB, U); Ibirama, 13 Dec 1953 (8), Gevieski 

85 (B, NY, S, US); Rio Pirai, 6 Jan 1950 (6), Hans 335 

(R, RB); Brusque, Mata Sao Pedro, 30 Dec 1949 (a), 

Klein 79B (= Veloso 97) (RB, US); Morro de Fazenda, 

Itajai, 25 Mar 1954 (6), Klein 772 (S, US); Morro dos 

Conventos, E of Ararangua, 5 Nov 1972 (6), Lima 

20795 (U), 15 Nov 1971 (9), Lindeman et al. 9105 and 

9110 (U); Sombrio, 3 Sep 1945 (8), Reitz 1912 (F, R, 

NY); Sao Francisco do Sul, 9 Jan 1951 (9), Reitz 3809 

(US); Campo do Massiambfi, Palhoca, 24 Sep 1953 

(a), Reitz et al. 975 (NY, S); Morro de Fazenda, Itajai, 

3 Mar 1954 ($), Reitz et al. 1688 (NY, S, US); Sombrio, 

8 Feb 1946 (8), Reitz et al. 2008 (US); Cunhas, Itajai, 

5 Aug 1954 (8), Reitz et al. 2027 (US); Morro de Ressacada, 

Itajai, 29 Mar 1957 (9), Reitz et al. 2913 (US); 

Morro do Bau, Itajai, 1 Nov 1951 (6), Reitz 4167 (NY, 

S); Trds Barras, Garuva, Sao Francisco do Sul, 24 Aug 

1957 (9), Reitz et al. 4692 (NY); Campo do Massiambi, 

Palhoca, 19 Dec 1952 (9), Reitz et al. 4965 (F, G, 

- (6), 

Schomburgk 876 (holotype, P; isotypes, BM, 

F, FI, NY). Fig. 37. 

Coussapoa fagifolia Klotzsch, Linnaea 20: 528. 1847; 

Miquel, Fl. bras. 4(1): 136. 1853. Type. Guyana. 

Pomeroon River, - (8), Schomburgk 1366 (=876?) 

(holotype, B; isotype, K). 

Coussapoa cuneata Miquel, Fl. bras. 4(1): 138. 1853. 

Type. Guyana. Berbice River, - (6), Schomburgk 

287 (holotype, U; isotypes, BM, G, GH, K, L, P, 

OXF, U). 

Tree, often hemi-epiphytic, up to 20 m tall. 

Leafy twigs 2-9 mm thick, sparsely to densely 

yellowish appressed-puberulous to hirtellous to 

subhirsute or also with white to brown arachnoid 

hairs. Lamina coriaceous (to subcoriaceous), 

ovate to subovate or elliptic to oblong, 3-27 x 

1.5-16 cm, apex acuminate to acute, base rounded 

to obtuse, margin entire, + revolute towards 

the base; upper surface glabrous, lower surface 

densely to very sparsely appressed-puberulous 

(on the main veins with hairs of different lengths) 

or also densely to sparsely covered with white to 

pale brown arachnoid hairs, disappearing or more 

or less persistent on the main veins and the margin; 

lateral veins 4-11 pairs, straight or slightly 

curved, basal pair branched or unbranched, in 

71

FIG. 37. Coussapoa microcephala: ssp. microcephala: 1, leafy twig with pistillate inflorescences (Maguire 24588); 2, lea 

(Gleason 405). 

I 

1cm


ovate to elliptic leaves reaching the margin below 

or sometimes at the middle of the lamina; intercostal 

venation more or less prominent to 

plane; petiole 0.5-5 cm long, appressed-puberulous 

to hirtellous or also with arachnoid hairs; 

(U); Pakaraima Mts., Kamarang, 8 Sep 1979 (6), Maas 

et al. 4126 (U); Kaieteur Gorge, 14 May 1944 (6), 

Maguire et al. 23449 (F, G, K, MO, NY, P, U, US); 

Demerara River, Comaka, Apr 1933 (6), Persaud 222 

(F); Essequibo River, Moraballi Creek, nr. Bartica, 18 

Sep 1929 (8), Sandwith 290 (K, NY, RB), 19 Sep 1929 

stipules 1.2-5 cm long, yellowish puberulous to (2), Sandwith 302 (K, U), 7 Oct 1929 (6), Sandwith 394 

subsericeous, often also with brownish arachnoid (K, P, US); without locality, 

hairs; terminal buds slender. Staminate inflorescences 

branched; heads six to many, globose, ca. 

2 mm diam.; common peduncle 1-4 cm long, 

puberulous; perianth ca. 1 mm high, glabrous; 

stamens two, far exceeding the perianth. Pistillate 

inflorescences branched, occasionally unbranched; 

heads (1-)3-15, sometimes fused, globose, 

2-5 mm, in fruit up to 10 mm diam.; 

(common) peduncle 1-10 cm long, puberulous; 

perianth ca. 1 mm high, glabrous. Interfloral 

bracts (sub)spathulate, minutely puberulous at 

the apex. 

Distribution (Fig. 7). Common in Guyana, apparently 

rare in Surinam and French Guiana; in 

forest, often along streams, at altitudes up to 

900 m. 

- (?), Schomburgk s.n. 

(F, G), - (6), Schomburgk 118 (B, K), - (9), Schomburgk 

167 (G, K, P); Berbice River, 1837 (6), Schomburgk 

287 (BM, G, GH, K, L, OXF, P, U, type collection 

of C. cuneata); Pomeroon River (?), - (6), 

Schomburgk 876 (by error 276) (BM, F, FI, G, NY, P, 

type collection of C. microcephala), Schomburgk 1366 

(=?876) (B, K, type collection of C. fagifolia); Potaro 

River, nr. Amatuk, 26 Aug 1959 (8), Whitton 179 (K). 

SURINAM. Without locality, - (9), Anderson s.n. 

(BM); Tafelberg, 1 Sep 1944 (9), Maguire 25488 (A, F, 

G, K, MO, NY, U, US). 

FRENCH GUIANA. Road to Brazil, km 6, at bridge 

over Compt6 River, ca. 51 km S of Cayenne, 1 Jan 

1977 (2), Mori 8864 (CAY, MO, NY, U). 

Specimens studied. GUYANA. Northwest District, 

Kaituma River, 27 Oct 1908 (a), Anderson 188 (K, 

NY); Cuyounia Creek, - (8), Appun 296 (K); Potaro 

River, nr. Kaieteur Falls, 18 Feb 1962 ($), Cowan et 

al. 1868 (F, K, US), 18 Feb 1962 (9), Cowan et al. 1880 

(F, K, US); Northwest District, Anabisi River, 15 Feb 

1922 (9), De la Cruz 1367 (GH, NY); upper Rupununi 

River, nr. Dadanawa, 14 Jun 1922 (a), De la Cruz 1501 

(F, GH, MO, NY); between Demerara River and Ber- 

bice River, 5?50'N, 15-19 Jul 1922 (6), De la Cruz 

1577 (F, GH, MO, NY); upper Mazaruni River, ca. 

60?10'W, 22 Sep-6 Oct 1922 (8), De la Cruz 2241 (F, 

GH, MO, NY, US), De la Cruz 2387 (F, GH, MO, 

NY, US); Demerara River, Malali, 30 Oct-5 Nov 1922 

(6), De la Cruz 2636 (F, GH, K, NY); upper Mazaruni 

River, Kamakusa, 23-29 Nov 1922 (9), De la Cruz 

2813 (F, GH, MO, NY); Demerara River, Malali, 30 

Oct-5 Nov 1922, De la Cruz 3070 (NY); Essequibo 

River, Winiperu Creek, 2 Nov 1939 (9), Fanshawe 295 

(FD 3031) (K, NY, U); Mazaruni River, Kurupung 

River, Sep 1925 (a), Gleason 405 (K, NY); Potaro Riv- 

er, Tumatumari, 4-6 Jul 1921 (9), Gleason 415 (GH, 

K, NY); Potaro River, nr. Kaieteur Falls, 17 Jan 1954 

(a), Irwin 183 (US); Mazaruni River, Sep 1880 (9), 

Jenman 652 (K); upper Demerara River, nr. Great 

Falls, Sep 1887 (2), Jenman 3995 (K, NY); Berbice 

River, May 1889 (2), Jenman 4947 (K), without lo- 

cality, Jun 1889 (e), Jenman 5315 (K); Demerara Riv- 

er, Mar 1898 (9), Jenman 7310 (K, S, U); Pomeroon 

River, nr. Macasuma, Feb 1904 (e), Jenman 7933 (K); 

Kupurung River, 22 Nov 1922 (9), Leng 177 (NY); 

Waine River, 10-15 km S of Kwabanna, 10 Aug 1977 

(9), Maas et al. 2478 (F, K, U); S of Timehri, Mr. 

Thompson's farm, 17 Oct 1979 (a), Maas et al. 3622 

73 

Coussapoa microcephala is closely related to 

C. argentea. Because of more or less distinct sim- 

ilarities in the dimensions, shape, margin, and 

venation of the leaves these two species together 

with C. latifolia, C. viridifolia, C. parvifolia, C. 

microcarpa, C. pachyphylla (and C. macerrima) 

constitute a more or less distinct group within 

the taxa having 2-staminate flowers. 

27. Coussapoa napoensis Akkermans & C. C. 

Berg, Proc. Kon. Ned. Akad. Wetensch., Ser. 

C. 85(4): 457. 1982. Type. Ecuador. Napo: 

Coca, 17 Jan 1973 (6), Lugo 2820 (holotype, 

GB). Fig. 38. 

Tree, up to 25 m tall, terrestrial or hemi-epiphytic. 

Leafy twigs 5-12 mm thick, sparsely puberulous, 

and mostly also with sparse, distinctly 

longer, straight to curved, stiff hairs. Lamina coriaceous, 

broadly ovate to broadly elliptic, 7-30 

x 6-22 cm, apex obtuse, occasionally acuminate, 

base subobtuse to rounded or to truncate, 

margin entire to subcrenate; upper surface glabrous, 

lower surface on the main veins with appressed 

straight hairs of different lengths and with 

sparse arachnoid hairs which later disappear; lateral 

veins (5-)7-12(-15) pairs, almost straight, 

basal pair (sometimes faintly) branched, reaching 

the margin below the middle of the lamina; intercostal 

venation slightly prominent to almost 

plane; petiole 3-10 cm long, with appressed 

straight hairs of different lengths, glabrescent; 

stipules 4-13 cm long, subvelutinous, also with 

distinctly longer straight stiff hairs. Staminate


~.I. 

FIG. 38. Coussapoa napoensis: 1, leafy twig with staminate inflorescences 

(Lugo 2857). 

I cm.


inflorescences repeatedly branched; heads many, 

globose, 2-3 mm diam.; common peduncle 2.5- 

6 cm long, puberulous to hirtellous; perianth ca. 

0.5-1 mm high, glabrous; stamen one, far exceeding 

the perianth. Pistillate inflorescences 

branched; heads 3-9, subglobose, 6-10 mm 

diam.; common peduncle 3-4 cm long, puberulous 

to subhirsute; perianth ca. 1 mm high, glabrous. 


Distribution (Fig. 7). Ecuador (Napo) and Colombia 

(Putumayo); Amazonian forest. 

Specimens studied. COLOMBIA. PUTUMAYO: Rio 

Putumayo, Pifiufia Negro, 20 Nov 1940 (6), Cuatrecasas 

10714 (F, US). 

ECUADOR. NAPO: Nr. Coca, 17 Jan 1973 (6), Lugo 

2820 (GB, type collection), 22 Jan 1973 (6), Lugo 2909 

(GB); Rio Payamino, nr. Payamino Capihuara, 19 Jan 

1973 (6), Lugo 2857 (GB); Shushufindi, 4 Aug 1975 

(e), Little et al. 43 (Q); 20 km W of Coca, 22-23 April 

1985 (v), Palacios et al. 6387 (BG). 

This species has many features in common 

with C. ovalifolia. It is distinct from the latter 

mainly in the longer stipules (of fertile twigs!), 

the smaller number of lateral veins, and the larger 

flower heads of the staminate inflorescence. The 

two species could prove to be distinct only at the 

subspecific level. 

28. Coussapoa nitida Miquel, Fl. bras. 4(1): 113, 

t. 44. 1853. Type. Brazil. Para: "Jaguary" 

(=Jaguarari?), - (e), Martius s.n. (lectotype, 

M; isolectotypes, B?, BR, U). Fig. 39. 

Coussapoa intermedia Miquel, Fl. bras. 4(1): 133, t. 

43. 1853. Type. Brazil. Without locality ("prov. Paraensi 

et Rio Negro"), - sparsely (to rather densely) covered with arachnoid 

hairs which are deciduous; lateral veins 8- 

15 pairs, mostly straight, basal pair (normally) 

branched, reaching the margin below the middle 

of the lamina; intercostal venation (almost) plane; 

petiole 3-13 cm long, 2-4 mm thick, densely 

brownish to white puberulous and with distinctly 

longer yellowish appressed to patent hairs; stipules 

2.5-10 cm long, yellowish to brownish subsericeous 

to hirsute or predominantly densely 

puberulous. Staminate inflorescence branched; 

heads numerous, free, globose, ca. 2 mm diam., 

often several solitary flowers on the branches of 

the inflorescence; common peduncle 2-5 cm long, 

rather densely puberulous to hirtellous to subhirsute; 

perianth ca. 1 mm high, glabrous or minutely 

puberulous; stamen one, far exceeding the 

perianth. Pistillate inflorescence branched; heads 

2-7, free, globose, ca. 4-6, in fruit up to 20 mm 

diam.; common peduncle 2-7 cm long, peduncle 

and branches 1-2 mm thick, rather densely puberulous 

to hirtellous, apices of the branches often 

more or less broadened towards the heads; perianth 

ca. 1 mm high, sparsely minutely white puberulous 

to glabrous. Interfloral bracts absent. 

Distribution (Fig. 8). Amazon Basin (Brazil 

and Peru), especially along the Amazon River; 

mostly in riverside (varzea) forest. 

Specimens studied. COLOMBIA. AMAZONAS: Rio 

Caqueta, Puerto Cordoba, 13 Nov 1912 (8), Ducke 

(HAMP) 12238 (MG); Trapecio Amazonico, nr. mouth 

of Rio Loretoyacu, 26 Nov 1945 (2), Duque-Jaramillo 

2207 (COL); Rio Loretoyacu, Oct 1945 (9), Schultes 

6726 (F, GH), Sep 1946 (2), Schultes et al. 8380 (F). 

(8), Martius s.n. 

PERU. LORETO: Rio 

(holotype, 

Itaya, ca. 5 km above Iquitos, 

6 

M; isotype, B). 

Aug 1972 (a), Croat 18850 (MO, U); Prov. Maynas, 

Coussapoa schunkei Standley, Publ. Field Mus. 

Rio 

Bot. Ampiyacu, between Pebas and mouth of Rio Ya- 

22: 72. 1940. Type. Peru. Loreto: Rio Mazan, Gam- guasyacu, 7 Nov 1977 (2), Gentry et al. 20381 (MO, 

itanacocha, 27 Jan 1935 (6), Schunke 130 (holotype, 

U); Rio Ampiyacu, above junction of Rio Yaguasyacu, 

9 Nov 

F; 

1977 

isotypes, A, NY, US). 

(2), Gentry et al. 20495 (MO, U); Rio 

Yavari, opposite Paumari, 23 Nov 1977 (a), Gentry et 

Tree, hemi-epiphytic or terrestrial, up to 30 m al. 20802 (U); Rio Nanay, Mishana, halfway between 

tall. Leafy twigs 5-12 mm thick, densely brown- Iquitos and Santa Maria de Nanay, 26 Feb 1979 (st), 

ish 

Gentry et al. 25124 

puberulous to 

(MO, U); lower Rio 

shortly velutinous, often 

Mom6n, nr. 

partly Iquitos, 8 Dec 1979 (9), J. Jones et al. 9712 (LAM, U), 

(sub)hirsute. Lamina (sub)coriaceous, ovate (to 9 Dec 1979 (8), J. Jones et al. 9769 (LAM, U); Rio 

elliptic), 8-30 x 4-20 cm, apex acute to obtuse, Ampiyacu, Puca Urquillo, 2 Apr 1977 (9), Plowman 

base obtuse to rounded to truncate to et al. 6555 

cordate, 

(F, GH, K, U); Rio Ampiyacu, Pebas, 16 

Jul 1976 

margin entire to subcrenate; upper surface (a), Revilla 805 (MO); Rio Mom6n, Mogla- 

noncillo, 17 Aug 1976 (9), Revilla 1099 

brous, lower 

(MO, 

surface 

U); Rio 

densely minutely puberu- Mom6n, above Bellavista, 19 Sep 1975 (9), Rimachi 

lous in the areoles and on the reticulum, usually 1943 (NA); Rio Mazan, Gamitanacocha, 27 Jan 1935 

sparsely puberulous to hirtellous on the (parallel) (8), Schunke 130 (A, F, NY, US, type collection of C. 

tertiary venation, (appressed-)puberulous to stri- schunkei). 

BRAZIL. Without 

gose or to sparsely hirsute on 

locality (Prov. Paraensi et Rio 

the midrib and Negro), 

lateral veins, sometimes the whole surface 

- (a), Martius s.n. (B, M, type collection of C. 

intermedia); - (a + 9, monoecious!), Poeppigs.n. (BR).


FIG. 39. Coussapoa nitida: 1, leafy twig with pistillate inflorescences 

(Schultes 6726).


ACRE: Rio Purus, nr. mouth of Rio Macaua, tributary 

of Rio Iaco, 21 Aug 1933 (6), Krukoff5596 (A, BM, 

F, G, K, LE, M, MO, NY, Q, S, U, US). AMAPA: Rio 

Jari, 2 km E of Arumanduba, 27 Jun 1961 (2), W. A. 

Egler et al. 46027A (IAN, MG, MO, NY, S, U). 

AMAZONAS: Rio Solim6es, B6ca de Manaquiri, 3 Oct 

1973 (6), Berg et al. P17591 (F, M, MO, P, S, U), 3 

Oct 1973 (2), Berg et al. P17592 (F, MO, K, P, S, U); 

Rio Japura, Mun. Maraa, Ilha dos Macacos, 30 Oct 

1977 (8), Damiao 2503 (INPA); Rio Japura, Costa do 

Jacitaria, 8 Dec 1977 (8), Damido 2817 (INPA); Rio 

Solimoes, Fonte Boa, 9 Oct 1945 (6), Ducke 1795 (A, 

F, K, MG, NY, R, RB, US); Rio Japura, 19 Sep 1904 

(9), Ducke (HAMP) 6782 (S, U); Rio Ia, 9 Sep 1906 

(9), Ducke (HAMP) 7715 (BM, G); Rio Tonantins, 26 

Oct 1949 (6), Frogs 25531 (IAN); Mun. Sao Paulo de 

Olivenca, nr. Palmares, 11 Sep-26 Oct 1936 (6), Krukoff 

8401 (A, B, BM, F, G, K, LE, NY, P, S, U, US); 

Rio Solimoes, Mamia, 20 Jan 1924 (8), Kuhlmann 1178 

(RB); Rio Javari, behind Palmeiras (army post), 6 Aug 

1973 (8), Lleras et al. P17197 (C, K, M, MO, NY, U, 

US). PARA: Rio Tajapurir, Ant6nia Lemos, Igarape Pixuna, 

19 Jul 1948 (2), Black 48-2949 (IAN); Rio Tajapuri, 

Ilha Sao Sebastiao, Nazar6, 31 Jul 1948 (6), 

Black 48-3017 (IAN); Gurupa, 27 Jan 1916 (6), Ducke 

(HJBR) 13066 (=HAMP 16003) (BM, RB); Rio Tajapurfi, 

- 21 cm, apex acute to obtuse to shortly acuminate, 

base cordate, margin entire or subcrenate; upper 

surface glabrous, lower surface densely minutely 

puberulous in the areoles and on the reticulum, 

(rather) sparsely hirtellous to puberulous on the 

other veins, on the main veins and margin also 

sparse white arachnoid hairs; lateral veins 9-11 

pairs, slightly curved, basal pairs branched, 

reaching the margin below the middle of the lamina, 

other lateral veins often poorly branched 

(furcate); intercostal venation very prominent; 

petiole 8-13 cm long, sparsely to densely minutely 

puberulous and sparsely covered with distinctly 

longer straight stiff hairs; stipules 0.5-3.5 

cm long, brownish subsericeous and with a dense 

covering of reddish-brown pluricellular hairs. 

Staminate inflorescences branched; heads many, 


1.5-3 cm long, puberulous to hirtellous, on the 

branches to tomentellous, mixed with sparse to 

dense reddish-brown pluricellular hairs; perianth 

(), Martius s.n. (FI); "Jaguary" (=?Jaguar- ca. 1 mm high, densely minutely puberulous; 

ari), Aug (9), Martius s.n. (B, BR, M, U, lectotype col- stamens three, exceeding the perianth. Pistillate 

lection), "Insulae Archipelagi Paraensis," Aug (d and inflorescences unbranched; heads ellipsoid to 

2), Martius s.n. (or 2673) (M, L, LE); Rio Jari, between 

Monte Dourado and Caracura, 18 Nov 1967 (9), Oliv- obovoid, ca. 20 x 10, in fruit up to 15 mm diam.; 

eira 3627 (IAN, NY); Rio Jari, between Monte Dour- peduncle 1-2 cm long, reddish-brown puberuado 

and Patricia, 27 Mar 1970 (6), N. T. Silva 303 lous to hirtellous; perianth ca. 2 mm high, dense- 

(IAN). 

ly reddish-brown puberulous. Interfloral bracts 

subpeltate, often only a few or sometimes absent. 

Distribution (Fig. 8). Costa Rica (Alajuela); in 

rain forest up to 800 m. 

Coussapoa nitida can easily be confused with 

C. tessmannii, due to the strong overall similar- 

ities. However, C. nitida is clearly distinct from 

the latter in the absence ofinterfloral bracts. Oth- 

er differences occur in the indumentum, espe- 

cially in that of the petiole, and in the diameter 

of the peduncle of the pistillate inflorescence. C. 

nitida appears to be related to C. cupularis and 

more remotely to C. orthoneura and C. arach- 

noidea. 

29. Coussapoa nymphaeifolia Standley, Proc. 

Biol. Soc. Wash. 37: 50. 1924; Burger, Field- 

iana Bot. 40: 134, t. 22. 1977. Type. Costa 

Rica. Alajuela: Road to San Carlos valley, 

Buena Vista, 16 Apr 1903 (2), Cook & Doyle 

157 (holotype, US). Fig. 40. 

Tree, hemi-epiphytic or terrestrial, ca. 20 m 

tall. Leafy twigs 5-17 mm thick, densely mi- 

nutely puberulous or also with dense reddish- 

brown pluricellular hairs. Lamina coriaceous, 

(broadly) elliptic to ovate, (9-)24-29 x (6-)18- 

Specimens studied. COSTA RICA. ALAJUELA: Road 

to San Carlos Valley, Buena Vista, 16 Apr 1903, (2), 

Cook & Doyle 157 (US, type collection); canton San 

Carlos, Villa Quesada, 21 Feb 1939 (8), A. Smith 1632 

(F, MO). HEREDIA: Nr. Puerto Viejo de Sarapiqui, 3 

Apr 1974 (2), Hartshorn 1436 (U). 

77 

Although C. nymphaeifolia is reminiscent of 

C. villosa in general appearance and also to C. 

duquei in the nature of the pistillate inflores- 

cence, these species are probably unrelated. The 

occurrence of 3-staminate flowers and other sim- 

ilarities in leaf characters and the staminate in- 

florescence suggest a relationship with C. lon- 

gepedunculata. 

30. Coussapoa oligocephala Donnell Smith, Bot. 

Gaz. 40: 11. 1905. Type. Guatemala. Alta Ve- 

rapaz: Cubilguitz, Apr 1904 (6), Von Tuerk- 

heim 8659 (=11.942) (holotype, US; isotype, 

B). Fig. 41.


?~ ~~~~~~~~~~~~~~~~~ 

.?' 

? 

:~~~~~~~~~~~~'- 

- ,??? I I 'I'~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~ 

'~~~~~~~~?: 

. 

P~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~ 

.I . ,. " 

r.~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~ 

?I ?::~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~ 

. . 

' ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~~~~ " 

~~~~~~ ''~ ~ ~ ~~~~~~~~~~~~~ 

.? ?~~~~~~~~~~~~ 

~. c , ,,. 

? ' 'y 

. 

~~~~~~~~~~~. . 

F~~~~~~~~~~~~~~~~~~~~...: 

?~~~~~~~~~~~~~~~~~~~~~~~~~~~ : 

?':. 

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

-L 

..T! 

?. jI .?~ 

~~~~~~~~~ .....?:;:"li?b 

i" 

~~~~~~~~~~~ 

.. .... ';: -:F: ..~,?; 

~~~~~~~~~~: ~ ?? .. . * 

"' 

~~;.? .~~~~~~~~~~~~~~' 

'~~~~~~~~~~~~~~ ? ;?r 

~~~~~~~~~~~~~~~~~~~~~~~~~~~'-?.- 

I.':! 

?-:' 

,:: 

, i' 

i; , : , ... 

?~~~~~~~~~~~~~~~~~~~~~~1 

"' 

" 

FIG. 40. Coussapoa nymphaeifolia: 1, leafy twig with staminate inflorescences, 2, leaf (A. Smith 1632). 

?:". 

'. .:i.- ; .


I 

1 cm 

FIG. 41. Coussapoa oligocephala: 1, leafy twig with staminate inflorescences (Schipp 999); 2, pistillate 


(Contreras 5787); 3, leaf (Contreras 5787). 

79

80 

Specimens studied. MEXICO. TABASCO: La Palma, 

Balancan, 6 Jun 1939 (6), Matuda 3300 (A, F, K, NA, 

NY). 

BELIZE. EL CAYO DISTRICT: Nr. El Cayo, 1933 (d), 


Shrub or tree, hemi-epiphytic or GUATEMALA. ALTA VERAPAZ: 

terrestrial, up 

Sebol, 21 Apr 1964 

(8), Contreras 4449 

to 20 m tall. Leafy twigs 4-7 mm 

(NY, S); El Cacao, Trece Aguas, 

thick, ap- - (9), Cooks.n. (F, US); Cerro Chinaja, between Finca 

pressed-puberulous to hirtellous to strigillose or Yalpemech and Chinnaja, 1-2 Apr 1942 (st), Steyerto 

hirsute. Lamina (sub)coriaceous, elliptic to mark 45663 (F); Cubilgultz, Apr 1904 (6), Von Tuerckoblong 

to (sub)ovate to (sub)obovate, 4-21 x 1- heim 8659 (=11.942) (B, US, type collection). IZABAL: 

7 cm, apex shortly acuminate to acute to 

Rio 

obtuse, 

Dulce, road Seja-Cienaga, km 5, 11 Jul 1970 (2), 

Contreras 10195 (BM, DUKE, MO, U); Rio Dulce, 21 

base obtuse to subcordate, margin entire; upper Jul 1936 (2), Hatch et al. s.n. (F); lower Rio Oscuro, 

surface glabrous, lower surface minutely puber- SW of Lake Izabal, 27 Apr 1966 (9), G. C. Jones et al. 

ulous to glabrous in the areoles sparsely hirtel- 3152 (NY, US); Lake Izabal, Jagua Creek, 31 May 1965 

lous on the smaller veins, the main veins gla- (9), Snedaker C39 (F), 28 May 1966 (9), Snedaker D88 

brous, the whole surface with (persistent) white (A, F); Rio Dulce, between Livingstone and 6 mi up 

river, 14 Apr 1940 (8), Steyermark 39460 (F); Rio Frio, 

arachnoid hairs; lateral veins (5-)8-12, straight, 17 Dec 1941 (st), Steyermark 39934 (F, US); Rio Dulce, 

basal pair unbranched, reaching the margin be- 21 May 1939 (8), Wilson 381 (F). PETEN: Vaxactun, 

low the middle of the lamina; intercostal vena- 30 May 1931 (2), Bartlett 12356 (GH, NY, S, US); 

tion prominent; petiole (1-)2-6 cm Tikal, 31 Jul 1959 (2), Contreras 61 (NY, 

long, sparsely 

S); Lake Itza, 

between Remate and San Andr6s, 18 Apr 1960 (9), 

to densely minute puberulous to hirtellous, or Contreras 845 (F, NY, S, US); old road to Machaquila, 

hairs mixed with longer patent to appressed, and Dolores, 27 Apr 1961 (8), Contreras 2220 (NY, S); nr. 

sometimes also arachnoid hairs; stipules 1-5 cm San Andr6s, Apr 1962 (2), Contreras 3559 (F, NY, S); 

long, (rather) sparsely appressed-puberulous (to Macanche, 18 May 1966 (9), Contreras 5787 (GH, US); 

Cadenas 

brownish subsericeous). Staminate road, km 138, La Cumbre, 25 Sep 1966 (9), 

inflores- Contreras 6231 (BM, MO, U); Lake Yaxha, archeocences 

branched; heads 3-10, globose, ca. 4-6 logical camp on north shore, 18 Jun 1973 (st), Croat 

diam.; common peduncle 1-5 cm long, with pu- 24663 (MO, NY); nr. Carmelita, 25 Jun 1942 (st), F. 

berulous to hirtellous to subtomentellous; peri- E. Egler 42-229 (F); 5 mi S of Tikal, 19 Jun 1973 (9), 

anth ca. 1 mm high, minutely puberulous; sta- Gentry 8348 (U); nr. San Andr6s, 2 May 1933 (9), 

Lundell 3170 

mens three, far 

(F); La Libertad, 31 

exceeding the perianth. Pistillate 

May 1933 (9), Lundell 

3535 (F, S); Tikal, 30 Apr 1959 (6), Lundell 15930 

inflorescences unbranched or occasionally poorly (S), 2 Jul 1959 (8), Lundell 16122 (F, S), 4 Jul 1959 

branched; heads 1(-3), globose, ca. 5-8 mm, in (9), Lundell 16162 (G, NY, S, US), 27 Jul 1969 (6), 

fruit up to 10 mm diam.; (common) peduncle 1- Ortiz 200 (BM, F); San Benito, nr. Playa Planca, 26 

5 cm long, ca. 1 mm thick, puberulous to hir- Apr 1970 (8), Ortiz 1030 (BM, F, MO, NY); Tikal, 29 

May 1971 (9), Ortiz 1810 (BM, F, US); Cerro Ceibal, 

tellous to subtomentellous; perianth ca. 1 mm between Rio Santa M6nica and mouth of Rio Martin, 

high, minutely puberulous; perianth orange or 30 Apr 1942 (st), Steyermark 46103 (F); Rio Machayellow. 

Interfloral bracts (sub)spathulate to sub- quila, N of El Cambio, 25 Apr 1942 (6), Steyermark 

peltate, minutely puberulous at the 45981 

apex. 

(F). 

Distribution (Fig. 8). Northern Central Amer- Coussapoa oligocephala is strongly reminisica 

(Guatemala and Belize) to southern Mexico cent of the Amazonian C. sprucei, especially when 

(Tabasco); in lowland (often riverine) forest. the specimens have an elliptic to subovate lamina 

with subcordate base. 

31. Coussapoa orthoneura Standley, Publ. Field 

Mus. Bot. 17: 165. 1937. 

Chanek 220 

Type. Brazil. Ama- 

(F, K); between Millionaro and Cueras, 

30 May 1973 (6), Dwyer 10799 (U); Vaca, 12 

zonas: Mun. Sao Paulo de 

May 

OlivenSa, nr. Pal- 

1938 (6), Gentle 2602 (A, F, K, MO); nr. Millionaro, mares, 26 Oct-11 Dec 1936 (2), Krukoff8518 

30 May 1973 (6), Gentry 7727 (MO, NY); Valentin, (holotype, NY; isotypes, A, B, BM, F, G, K, 

Jun-Jul 1936 (6), Lundell 6224 (F, GH, NY, S, US), LE, MO, SU, U, US). Fig. 42. 

Jun-Jul 1936 (2), Lundell 6350 (C, F, GH, MO, NY, 

S, US); Macaw Bank, 3 Apr 1969 (9), Proctor 30273 Coussapoa williamsii Cuatrecasas, Fieldiana Bot. 28(1): 

(BM, MO). TOLEDO DISTRICT: Bolo Camp, 13 Apr 212. 1951. Type. Venezuela. Terr. Fed. Amazonas: 

1944 (6), Gentle 4528 (DUKE); Honey Camp-Orange Isla Solitaria, between Tamatama and Esmeralda, 7 

Walk, Sep 1928 (st), Lundell 2 (BM, F, K), 24 Oct 1929 May 1942 (6), LI. Williams 15240 (holotype, F; iso- 

(6), Lundell 648 (F, K, MO, NY, S, US); Toledo, 1906- types, A, G, NY, RB, US, VEN). 

1907 (6), Peck 497 (GH, K, NY, U); Punta Gorda, 7 Coussapoa prancei C. C. Berg, Acta Bot. Neerl. 27(1): 

Aug 1932 (6), Schipp 999 (A, BM, F, G, GH, K, MO, 11. 1978. Type. Brazil. Amazonas: Rio Cuieras, nr. 

NY, S). 

Jarada, 17 Sep 1973 (9), Prance et al. 18032 (holo-


FIG. 42. Coussapoa orthoneura: 1, leafy twig with pistillate inflorescences (Prance 14092); 2, leafy twig with 

staminate inflorescences (Schultes et al. 13774). 

type, INPA; isotypes, C, F, K, M, MICH, MO, NY, 

P, SU, U, US). 


tall. Leafy twigs 2-6 mm thick, glabrous or 

sparsely, puberulous, sometimes hirtellous. 

Lamina (sub)coriaceous, obovate to (broadly) elliptic 

to oblong, 2-15(-24) x 1-8(-11) cm, apex 

1 cm 

81 

acute to shortly acuminate or obtuse, base obtuse 

to acute or truncate, margin entire or subcrenate; 

upper surface glabrous, lower surface glabrous or 

sparsely to densely appressed-puberulous, sometimes 

substrigose or with white arachnoid hairs 

later deciduous; lateral veins 2-7 pairs, straight 

or slightly curved, basal pairs unbranched reach-


ing the margin above to just below the middle PERU. AMAZONAS: Prov. Bagua, Dist. Cenepa, 

of the lamina; intercostal venation (almost) plane; 

Nueva Nazareth, nr. mouth of Rio Imaza, on Rio Mapetiole 

1-7 cm long, glabrous or raiion, 26 Jan-18 Feb 1967 (2), Tillett 671-33 (YEN). 

appressed-pu- LORETO: Prov. Maynas, Rio Nanay, nr. Moron Cocha, 

berulous to strigillose or to hirtellous; stipules N of Iquitos, 21 Mar 1977 (6), Gentry et al. 18528 

(0.5-)1-7 cm long, sparsely to densely appressed- (MO, U), Rio Nanay, nr. Caserio Santa Clara, nr. Iquipuberulous 

or to mostly sparsely yellowish sub- tos, 19 Nov 1976 (2), Revilla 1847 (U). 

sericeous (to subhirsute or to subvillous), occa- BRAZIL. AMAZONAS: Manaus, Igarap6 do Leao, 3 

sionally also with sparse to rather dense 

Sep 1957 (6), L. Coelho 

white 

(INPA) 5728 (U); Nogueira- 

Tef'e, 26 Oct 1972 (2), Danta 12383 (INPA); Tonanarachnoid 

hairs. Staminate inflorescences tins, 7 Feb 1944 (9), Ducke 1528 (NY, US); Rio Purus, 

branched; heads many, sometimes partly fused, Quinta de Sao Christovao, 18 Jan 1873 (6), Glaziou 

globose, ca. 1-6 mm diam.; common peduncle 10070a (P); Mun. Sao Paulo de Olivenca, nr. Palmares, 

1-3 cm long, with 

11 

(dense) appressed-puberulous, 

Sep-26 Oct 1936 (2), Krukoff8518 (A, BM, F, G, 

K, LE, NY, S, U, US, 

sometimes 

type collection), (6), Krukoff8587 

also with reddish-brown pluricellular (F); Mun. Sao Paulo de Olivenca, Belem Creek, 26 

hairs; perianth ca. 1 mm high, glabrous or min- Oct-11 Dec 1936 (6), Krukoff8967 (A, B, BM, F, G, 

utely puberulous; stamen one, just exceeding K, LE, MO, NY, S, U); Lago do Tefe, 10 km above 

the perianth. Pistillate inflorescences branched; 

Tefe, 26 Jul 1971 (8), McDaniel et al. 2666 (US); Rio 

heads 2-10, sometimes partly fused, 3-6 mm, in Ituxi, nr. B6ca do Curuquete, 9 Jul 1971 ((), Prance et 

al. 14029 (F, GH, K, M, S, U); Rio Cuieras, just below 

fruit up to 15 mm diam.; common peduncle 1- mouth of Rio Branchino, 29 Sep 1971 (6), Prance et 

3 cm long, (densely) appressed-puberulous, al. 15039 (F, GH, K, M, P, S, U); Rio Cuieras, nr. 

sometimes partly hirtellous, apices of the head- Jarada, 17 Sep 1973 (2), Prance et al. 18032 (C, F, K, 

bearing branches often (especially in fruit) more M, MICH, MO, NY, P, S, U, US, type collection of 

C. 

or less 

prancei); road Manaus-Itacoatiara, km 29, 14 Sep 

swollen; perianth ca. 1-2 mm high, gla- 1974 (2), Prance et al. 22729 (INPA, U); Mun. Purabrous 

or sparsely papillate at the apex; fruiting quequara, road to Manaus, Panama, 6 Jun 1963 (2), 

perianth yellow to red. Interfloral bracts absent. W. A. Rodriques et al. 5266 (U); Rio Castanho, road 

Distribution (Fig. 8). Upper Amazon Basin to Careiro, 11 Jul 1972 (8), M. Silva et al. 466 (U); Rio 

(Brazil, Peru, Ecuador, and Venezuela); in Uaup6s, Taracua, Rio Tikie, 28 Jan-9 Feb 1948 (6), 

up- Schultes et al. 9685 (GH, K); Rio Uaup6s, Panur6, Oct 

land and riverine (varzea) forest; also known from 1852-Jan 1853 (8), Spruce 2498 (BM, G, GH, K, LE, 

Central Columbia (Santander, at ca. 1000 m). NY, OXF, P). 

Specimens studied. COLOMBIA. AMAZONAS: Rio 

Igara-Parana, affluent of Rio Putumayo, La Chorrera, 

19 Dec 1973 (9), Gasche et al. 29 (U). GUAINiA: Rio 

Guainia, Manacal, 21 Oct 1977 (2), Espina et al. 219 

(COL). SANTANDER: Rio Negro, El Play6n, 1000 m, - 

Coussapoa orthoneura is a variable species, 

especially in the shape, dimensions, and secondary 

venation of the lamina and also in the indumentum 

of the stipules. More or less distinct 

(st), Franco & Garces 419 (COL). VAUPES: Upper Rio regional forms, like the small-leaved form found 

Vaup6s, La Jirisa, 9 Jan 1944 (a), Guttierrez et al. 562 near Manaus, at the time described as C. prancei, 

(COL, GH); upper Rio Vaup6s, between La Jarisa and have 

Las Bocas, 12 Jan 1944 (Q), Guttierrez et al. 595 

proved not to be sufficiently different to be 

(COL); 

Rio Apaporis, nr. Cachivera de Jirijirimo, 8 Jul 1951 recognized as distinct (infraspecific) taxa. Cous- 

(Y), Schultes et al. 12985 (F); Rio Apaporis, between sapoa arachnoidea appears to be closely related 

Rio Pacoa and Rio Kananari, 27 Aug 1951 (6), Schultes to C. orthoneura, which shows affinities espeet 

al. 13774 (COL, GH, U); Rio Apaporis, Raudal de 

Jirijirimo, below 

cially to C. cupularis and C. nitida. In the 

mouth of Rio Kananari, 14 

veg- 

Feb 1952 

(9), Schultes et al. 15324 

etative 

(GH, 

parts the 

US). 

species can easily be confused 

VENEZUELA. AMAZONAS: Rio Casiquiare, be- with C. latifolia, C. viridifolia and C. microcephtween 

Lago de Vasiva and Rio Pacimoni, 1853-1854 ala, but it is quite distinct from these three species 

(a), Spruce 3231 (B, C, G, K, P, S); Rio Ocama, above in the ebracteate inflorescence and the staminate 

confluence with Rio Orinoco, nr. Ocama Mission, 23- flower. 

24 May 1972 (8), Steyermark 106167 (U, VEN); Isla 

Solitaria, between Tamatama and Esmeralda, 7 May 

1942 (6), Ll. Williams 15240 (A, F, G, type collection 32. Coussapoa ovalifolia Trecul, Ann. Sci. Nat. 

of C. williamsii); Esmeralda, 14 May 1942 (a), LI. Wil- Bot. S6r. 3, 8: 95. 

liams 

1847; 

15363 

Miquel, Fl. bras. 4(1): 

(G, NY, US, VEN). 

139. 

ECUADOR. NAPO: Misahualli, mouth of Rio Misa- 

1853; Macbride, Publ. Field Mus. Bot. 

hualli, 3 Mar 1980 (st), Berg & Akkermans 1111 (U); 13(2.2): 298. 1937. Type. Peru. Huanuco: Maroad 

Coca-Lago Agrio, 9 km NE of Rio Coca, 20 Mar cora ?, 

1980 (st), Brandbyge et al. 30258 (AAU). 

- (2), Ruiz & Pavon s.n. (or 2) (holotype, 

FI; isotypes, F, US). Fig. 43.


FIG. 43. Coussapoa ovalifolia: 1, leafy twig with pistillate inflorescences 

(Krukoff5657); 2, staminate inflo- 

rescences (Ule 9316). 

83


Coussapoa hirsuta Tr6cul, Ann. Sci. Nat. Bot. Ser. 3, & Akkermans 1132 (U). ZAMORA-CHINCHIPE: Road 

8: 97. 1847; Macbride, Publ. Field Mus. Bot. 13(2.2): to Guayzimi, 27 km NNE of Zamora, 17 Sep 1975 

297. 1937. Type. Peru. Huanuco: Macora, - (9), (st), Little et al. 424 (COL, Q). 

Ruiz & Pavon s.n. (or 5) (holotype, FI; isotype, B). PERU. AMAZONAS: E ofHuampmai, 4 Jul 1974 (2), 

Coussapoa acutifolia Klotzsch, Linnaea 20: 529. 1847; Berlin 1531 (MO, U). HuANuco: Prov. Leoncio Prado, 

Macbride, Publ. Field Mus. Bot. 13(2.2): 296. 1937. Tingo Maria, 7 Dec 1981 (9), Plowman et al. 11179 (U); 

Type. Peru. Huanuco: Cochero (fide Macbride, 1937), Chinchoa-Pillao-Pozuzo-Mucuna, - (2), Ruiz & Pa- 

- (9), Ruiz & Pavon s. n. (or 7) (holotype, B; isotype, von s.n. (or 6), (B, F, lectotype collection of C. puber- 

FI). 

ula); Cochera, - (2), Ruiz & Pavon s.n. (or 7) (B, F, 

Coussapoa puberula Klotzsch, Linnaea 20: 529. 1847. FI, US, type collection of C. acutifolia); Macora, 

Type. Peru. Huinuco: "Chinchao-Pillao-Pozuzo- 

Mucona," - (9), Ruiz & Pavon s.n. (or 6) (holotype, 

B; isotype, F). 

Coussapoa setosa Klotzsch, Linnaea 20: 528. 1847. 

Type. Peru. Huinuco: Macora, - (9), Ruiz & Pavon 

s.n. (or 5) (holotype, B; isotype, FI). 

- (2), 

Ruiz & Pavon s.n. (or 2) (F, FI, US, type collection of 

C. ovalifolia), - (2), Ruiz & Pavon s.n. (or 7) (B, FI, 

type collection of C. hirsuta and C. setosa); Prov. Pachitea, 

Dtto. Honoria, Bosque Nac. Iparia, Schunke V. 

1162 (F, G, NY, US). JUNIN: Valley of Rio Paucartambo, 

nr. Perene bridge, 19 Jun 1929 (a), Killip et al. 

25341 (F, NY, S, US); Panchis trail, between San Nicolas 

and Azupizfi, 6 Jul 1929 (2), Killip et al. 26095 

(NY); La Merced, 10-24 Aug 1928 (a), Macbride 5594 

(F). LORETO: Prov. Maynas, Rio Nanay, Mishana, halfway 

between Iquitos and Santa Maria de Nanay, 26 

Jul 1979 (st), Gentry 25151 (MO, U); Rio Marafion, 

Yanache, 2 Mar 1924 (9), Kuhlmann 1540 (U); Quebrada 

Aucaya, 12 May 1973 (9), Rimachi 359 (NA); 

Rio Maranion, between Iquitos and Pongo de Manseriche, 

1924 (9), Tessmann 3922 (G); Matari Cocha, 3 

Oct 1968 (d), Torres M. 482 (K). MADRE DE DioS: 

Parque Nacional del Manu, Cocha Cashu, between 

Punagua and Tayakome, 17-24 Aug 1974 (2), Foster 

et al. 3412 (F). SAN MARTIN: Eslab6n, between Saposoa 

and Tinge de Saposoa, Rio Saposoa, 8 Sep 1948 

(9), Ferreyra 4813 (US); between Gramalote and Saposoa, 

29 Apr 1962 (a), Woytkowski 7306 (GH, US). 

BRAZIL. ACRE: Mouth of Rio Macaua, tributary of 

Rio laco, 26 Aug 1933 (9), Krukoff 5657 (A, BM, F, 

G, K, LE, M, S, U); nr. Tarauaca, 25 Sep 1970 (2), 

Prance et al. 7371 (U); Rio Acre, Seringal Sao Francisco, 

Aug 1911 (9), Ule 9315 (G, K, L, MG), May 

1911 (a), Ule 9316 (G, K, L, MG, US). 

BOLIVIA. SANTA CRUZ: Buena Vista, Jun 1915 (8), 

Steinbach 1484 (GH, U). 

Tree, hemi-epiphytic, up to 20 m tall. Leafy 

twigs 3-9 mm thick, glabrous or puberulous, often 

also with distinctly longer straight stiff hairs. 

Lamina coriaceous, ovate to subovate (or elliptic 

to obovate), 7-32 x 2-16 cm, apex shortly acu- 

minate to acute (to obtuse), base obtuse to round- 

ed or acute to truncate, margin entire or sub- 

crenate; upper surface glabrous, lower surface 

glabrous or with the main veins having appressed 

straight hairs of different lengths, often initially 

also with sparse or sometimes rather dense white 

arachnoid hairs on the main veins and along the 

margin; lateral veins 7-21 pairs, almost straight 

to curved, basal pair unbranched or in large leaves 

poorly branched, reaching the margin far below 

the middle of the lamina; intercostal venation 

almost plane to slightly prominent; petiole 2-9 

cm long, glabrous or with appressed hairs of dif- 

ferent lengths; stipules 1-4 (in rapidly growing 

shoots up to 13) cm long, subsericeous (to sub- 

hirsute). Staminate inflorescences repeatedly 

branched; heads numerous, globose, ca. 1(-2) mm 

diam.; common peduncle 1.5-3 cm long puberu- 

lous to hirtellous, occasionally also with sparse 

arachnoid hairs; perianth 0.5-1 mm high, gla- 

brous; stamen one, just exceeding the perianth. 

Pistillate inflorescences branched or sometimes 

unbranched; heads 1-5, subglobose, 5-8 mm, in 

fruit up to 15 mm diam.; (common) peduncle 2- 

6 cm long, puberulous to hirtellous; perianth ca. 

1-2 mm high, glabrous; fruiting perianth brown- 

ish. Interfloral bracts absent. 

Distribution (Fig. 8). Upper Amazon Basin 

(Bolivia, Peru, Brazil and Ecuador); in upland 

and riverine forest, up to 900 m. 

Specimens studied. ECUADOR. NAPO: Lago Agrio, 

3 Apr 1980 (9), Brandbyge et al. 30421 (AAU). PASTA- 

ZA: Between Palora and Shell, 4 Mar 1980 (st), Berg 

Coussapoa ovalifolia is closely related to C. 

napoensis. The collection Gentry & Mori 14011, 

Panama, Cerro Tacaruna, premontane wet for- 

est, 1300 m, 30 Jan 1975 (MO, U), probably 

represents a separate taxon, possibly distinct at 

the subspecific level from the Amazonian rep- 

resentatives of C. ovalifolia. The collection is 

similar in most characters to the other collections 

of C. ovalifolia, but is distinct mainly in the short 

(1-1.5 cm long) peduncle of the pistillate inflo- 

rescence andthe branching of the basal lateral 

veins-features which hardly justify recognition 

as a distinct species. Staminate material is need- 

ed to confirm the present supposition. 

33. Coussapoa pachyphylla Akkermans & C. C. 


C, 85(4): 459. 1982. Type. Brazil. Bahia: Mun.


lcm. 

FIG. 44. Coussapoa pachyphylla: 1, leafy twig with pistillate inflorescences 

(Mori & Santos 11655). 

Santa Cruz de Cabralia, old road to Santa Cruz, 

ca. 15 km NW of P6rto Seguro, 3 Apr 1979 

(9), Mori & dos Santos 11655 (holotype, CE- 

PEC; isotype, U). Fig. 44. 

Shrub, often (hemi-)epiphytic, up to 5 m tall 

(or taller?). Leafy twigs 4-8 mm thick, rather 

densely to sparsely appressed-puberulous, also 

with distinctly longer straight stiff hairs and brown 

pluricellular hairs, glabrescent. Lamina coriaceous, 

ovate to elliptic (or obovate), 6-16(-24) 

x 3.5-10(-19), apex acute to subacuminate, base 

acute, obtuse or sometimes subcordate, margin 

entire to subcrenate; upper surface glabrous, lower 

surface densely to sparsely puberulous to subtomentose, 

initially often with sparse brownish 

arachnoid hairs, especially along the margin; lateral 

veins 5-8 pairs, basal pair branched or sometimes 

(in small leaves) unbranched, reaching the 

margin below the middle of the lamina; inter- 

costal venation more or less prominent; petiole 

0.5-3(-5) cm long, densely to sparsely puberu- 

lous, often also with distinctly longer straight stiff 

hairs; stipules 1-2 cm long, densely covered with 

short reddish-brown arachnoid hairs. Staminate 

inflorescences branched; heads ca. 6-12, some- 

times fused, globose, ca. 1-2 mm diam.; com- 

mon peduncle 1-2 cm long, densely puberulous; 

perianth ca. 0.5 mm high, nearly glabrous; sta- 

mens two, as long as the perianth. Pistillate in- 

florescences unbranched or few-branched; heads 

1-3, sometimes partly fused, ca. 5-8 mm, in fruit 

up to 13 mm diam.; (common) peduncle 2.5-4 

cm long, 1-1.5 mm thick, densely puberulous; 

perianth 1(-2) mm high, glabrous. Interfloral 

bracts, few, subpeltate to spathulate. 

Distribution (Fig. 8). Brazil (Bahia); in humid 

forests in the coastal region.

86 

Specimens studied. BRAZIL. BAHIA: Road Una-Ilheus, 

ca. km 21, 2 Apr 1980 (st), Berg et al. 1153 

(CEPEC, K, NY, RB, U); Mun. Santa Cruz de Cabralia, 

old road to Santa Cruz, ca. 15 km NW of Porto Seguro, 

3 Apr 1979 (Q), Mori et al. 11655 (CEPEC, U, type 

collection); Ilh6us-Repartimento (Fortuna), 21 Nov 

1944 (8), Vellozo 737 (R). 


tall. Leafy twigs 4-8 mm thick, glabrous or pu- 

berulous. Lamina coriaceous, broadly ovate to 

elliptic or suborbicular, sometimes tending to 

obovate, 7-24 x 4-18 cm, apex shortly acumi- 

nate, sometimes almost rounded or obtuse, base 

rounded to truncate or subcordate, margin en- 

tire; upper surface glabrous, lower surface gla- 

brous, occasionally sparsely puberulous to to- 

mentellous; lateral veins 3-6 pairs, slightly 

curved, basal pair branched, reaching the margin 

at or above the middle of the lamina; intercostal 

venation plane or slightly prominent; petiole 

(0.5-)2-7 cm long, occasionally sparsely puber- 

ulous; stipules 1.5-5 cm long, glabrous or sparse- 

ly puberulous. Staminate inflorescences branched; 

heads many, globose, ca. 1 mm diam.; common 

peduncle 1-5 cm long, glabrous or sparsely pu- 

berulous; perianth ca. 0.5 mm high, glabrous; 

stamens three, just exceeding the perianth. Pis- 

tillate inflorescences branched; heads 3-12, glo- 

bose, ca. 2-3 mm, in fruit up to 6 mm diam.; 

common peduncle 1-4 cm long, glabrous or 

sparsely puberulous; flowers connate, perianth 

ca. 1 mm high, glabrous. Interfloral bracts lack- 

ing or in the staminate heads a few, more or less 

distinctly spathulate (more or less cucullate), gla- 

brous or ciliolate bracts. 


Distribution (Fig. 8). Costa Rica, Panama, and 

in the Pacific coastal region of Colombia and 

northern Ecuador; in forests up to 1250 m. 

Specimens studied. COSTA RICA. ALAJUELA: Nr. 

San Ram6n, Cataratas (Los Angeles) de San Ram6n, 

17 Apr 1935 (9), Brenes 20542 (F, NY, type collection 

This species appears to be related to C. latifolia of C. brenesii); San Carlos, Buena Vista, Finca La Conand 

allied taxa. 

stancia, 3 Mar 1963 (2), Jimenez 433 (F, MO); Rio San 

Rafael, 2 km W of La Marina, Llanura de San Carlos, 

21 Feb 1966 (6), Molina R. et al. 17694 (F); San Carlos, 

34. Coussapoa parviceps Standley, Proc. Biol. La Sucre, 1025 m, 2 Mar 1939 (6), A. Smith 1650 (A, 

Soc. Wash. 37: 51. 1924; Burger, Fieldiana F, MO), 1 Mar 1939 (2), A. Smith 1939 (6), A. Smith 

Bot. 40: 136, t. 22. 1977. Type. Costa Rica. 1650 (A, F, MO), 1 Mar 1939 (2), A. Smith 1663 (F). 

CARTAGO: 

Puntarenas: Agua Buena valley, nr. Cafnas 

Rio Pejibaye, between Rio Taus and Quebrada 

Azul, 28 May 1972 (st), Lent 2538 (COL, F); 

Gordas, 1100 m, Feb 1897 (9), Pittier 11166 Moraria de Turrialba, 21 Apr 1975 (st), Poveda 951 

(holotype, US; isotype, MO). Fig. 45. (F); nr. Orosi, 30 Mar 1924 (6), Standley 39792 (US). 

PUNTARENAS: Canias Gordas, Finca Loma Linda, 1140 

Coussapoa brenesii Standley, Publ. Field Mus. Bot. 18: m, 23 Feb 1973 (9), Busey 628 (F, GH, MO); Rio Java, 

383. 1937. Type. Costa Rica. Alajuela: nr. San Ra- Las Cruces Bot. Garden, 16 Mar 1978 (9), Hartshorn 

m6n, Cataratas (Los Angeles) de San Ram6n, 17 Apr 2157 (U); Agua Buena valley, nr. Canias Gordas, 1100 

1935 (Q), Brenes 20542 (holotype, F; isotype, NY). m, Feb 1897 (2), Pittier 11166 (MO, US, type collec- 

Coussapoa oligoneura Mildbraed, Notizbl. Bot. Gart. 

tion). 

Berlin 10: 415. 1928. Type. Colombia. Without lo- PANAMA. CHIRIQUi: Burica Peninsula, Mellize, 6 

cality, 19 Feb 1892 (9), Triana 866 (holotype, B; mi S of Puerto Armuelles, 5 Mar 1973 (st), Liesner 

isotype, P). 

417A (MO). COCLE: Road La Pineda-El Cope, nr. sawmill 

above El Cop6, 1000 m, 20 May 1978 (6), Hammel 

2576 (U). PANAMA: Road El Llano-Carti, km 8-12, 

13 May 1973 (st), Nee et al. 8822 (MO, NY, U). VERA- 

GUAS: NW of Santa F6, 2.7 km from Escuela Agricola 

Alto de Piedra, 30 Mar 1975 (2), Mori et al. 5360 (MO, 

NY). 

COLOMBIA. Without locality, 19 Mar - (2), Triana 

866 (B, P, type collection of C. oligoneura). NARIuO: 

Barbacoas, May 1853 (9), Triana 1866 (NY, photograph). 

VALLE: Rio Cajambre, Barco, 21-30 Apr 1944 

(2), Cuatrecasas 17351 (F); Rio Anchicaya, Alto Yunda, 

1000 m, May 1972 (2), Hilty M87 (MO). 

ECUADOR. ESMERALDAS: Alto Tambo, 23 Sep 1965 

(8), Little et al. 21136 (NY, Q, US). 

Coussapoa parviceps differs from other Cous- 

sapoa species in the connate pistillate flowers, 

which is probably the reason why interfloral 

bracts, present in staminate inflorescences, are 

lacking in the pistillate ones. 

Several characters, particularly as exhibited in 

the collection Little & Dixon 21136, suggest 

taxonomic relationship to C. cinnamomifolia. 

The collections Liesner 417A and Nee et al. 

8822, both from Panama, presumably represent 

juvenile specimens of the species. They are dis- 

tinct in having narrower (oblong to subovate) 

and larger (up to 50 cm long) laminae with up 

to 18 pairs of lateral veins, the basal pair of which 

reaches the margin below the middle of the lam- 

ina; petiole up to 20 cm long; the densely puberu-


Fa1to4 I" 

FIG. 4.osppc 11 w 

.cm 

FIG. 45. Coussapoa parviceps: 1, leafy twig with pistillate inflorescences 

(Jimenez 433). 

lous twigs, petioles, and stipules (both sides!); 

and in the presence of white arachnoid hairs on 

the margin and midrib on the lower surface of 

the lamina. The characters of these specimens 

are not included in the species description. Cous- 

sapoa parviceps appears to be the only Coussapoa 

species with pronounced dimorphism of leaves 

depending upon age. 

87 

35. Coussapoa parvifolia Standley, Publ. Field 

Mus. Bot. 17: 165. 1937. Type. Brazil. Ama- 

zonas: Mun. Sao Paulo de Olivenga, nr. Pal- 

mares, 11 Sep-26 Oct 1936 (Q), Krukoff8273 

(holotype, NY; isotypes, A, BM, F, G, K, LE, 

MO, S, U, US). Figs. 46, 47. 

Cousspoa cornifolia Standley, Publ. Field Mus. Bot. 

17: 160. 1937. Type. Brazil. Amazonas: Mun. Sao

FIG. 46. Coussapoa parvifolia: 1, leafy twig with pistillate inflorescences (Krukoff8539); 2, leafy twig with 

staminate inflorescences (Prance et al. 9054); 3, leaf (Prance et al. 22999); 4, stipules (Krukoff 8539).


F: 1, ly tg wh 1 cm 

FIG. 47. Coussapoa parvifolia: 1, leafy twig with pistillate inflorescences 

(Krukof 8273). 

Paulo de Oliven:a, Bel6m Creek, 26 Oct-11 Dec ger brownish patent hairs, soon disappearing; 

1936 (d), Krukoff8897 (holotype, NY; isotypes, A, stipules 0.4-1(-1.3) cm long, appressed-puber- 

F, G, K, LE, MO, P, S, U, US). 

ulous to 

Coussapoa microcephala subsp. cornifolia Akkermans subsericeous, terminal buds mostly 

& C. C. Berg, Proc. Kon. Ned. Akad. Wetensch., Ser. swollen (containing young inflorescences). Sta- 

C, 85(4): 457. 1982. 

minate inflorescences repeatedly branched; heads 

ca. 15-30, globose, ca. 2-3 mm diam.; common 

Tree, up to to 20 m tall, hemi-epiphytic. Leafy peduncle 1.5-2 cm long, rather sparsely pubertwigs 

2-5 mm thick, rather sparsely appressedulous 

or partly hirtellous; perianth ca. 1 mm high, 

puberulous, sometimes on the younger parts also minutely ciliolate; stamens two, filaments just to 

longer, brownish, patent (to appressed) hairs, soon far exceeding the perianth. Pistillate infloresdisappearing. 

Lamina coriaceous to subcoria- cences unbranched or branched heads (1-5, gloceous, 

lanceolate to narrowly ovate, sometimes bose, 4-8 mm diam.; (common) peduncle 1-2.5 

subobovate, 4-14(-21) x 1.5-5.5(-11) cm, apex 

cm long, puberulous perianth ca. 1 mm high, 

acute to acuminate, base acute to obtuse, margin glabrous. Interfloral bracts (sub)spathulate, mientire, 

towards the base + revolute; upper sur- nutely puberulous. 

face glabrous, lower surface ? densely ap- 

Distribution (Fig. 8). Upper Amazon Basin, in 

pressed-puberulous to glabrous; lateral veins Brazil (Amazonas), Peru (Loreto), Venezuela 

(4-)7-11 pairs, basal pair unbranched, reaching (Amazonas), also in Extra-Amazonian Colombia 

the margin below or in subobovate leaves above (Antioquia and Vichada); in non-inundated forthe 

middle of the lamina; intercostal venation est. 

slightly prominent; petiole 0.5-3(-4) cm long, Specimens studied. COLOMBIA. ANTIOQuIA: Rd. 

appressed-puberulous to glabrous also with lon- Mutat&-Pavarand6, between haciendas La Esperanza


and Mocari, 150 m, 6 Mar 1987 (6), Fonnegra et al. cm long, glabrous; perianth 1-2 mm high, gla- 

1798 (BG, HUA). VICHADA: Gaviotas, Caino Ariba, 19 

Feb 1973 (9), Cabrera 2668 brous; fruiting perianth yellow or orange. Inter- 

(COL). 

VENEZUELA. AMAZONAS: Rio Cucucucuma, be- floral bracts (sub)spathulate, sparsely puberulous 

low Raudal Pacure, 9 Nov 1950 (6), Maguire et al. at the apex. 

29408 (NY); Dpto. Atabapo, 15 km SE of San Fer- Distribution (Fig. 8). Southern Mexico and 

nando de Atabapo, 10-16 Feb 1988 (d), Stergios et al. north-western 

11530 

Guatemala; in evergreen forest in 

(BG). 

BRAZIL. AMAZONAS: Mun. Sao Paulo de 

moist 

Olivenoa, 

places (often along streams) or in seminr. 

Palmares, 11 Sep-26 Oct 1936 (2), Krukoff 8273 deciduous forest; up to 1700 m. 

(A, F, G, K, LE, MO, NY, P, S, U, US, type collection 

of C. parvifolia), (2), Krukoff8539 (A, BM, F, G, LE, Specimens studied. MEXICO. CHIAPAS: Ovando Mt., 

MO, NY, P, S, U, US); Mun. Sao Paulo de 17 Dec 1936 

Olivenca, 

(6), Matuda 446 (MO, NY, US); Santa 

Belem Creek, 26 Oct-11 Dec 1936 (6), Krukoff8658 

Rita, Mapastepec, Jan 1938 (6), Matuda 2020 (A, F, 

(A, BM, F, G, K, LE, MO, NY, P, S, U, US), (6), Krukoff 

K, NA, NY); between Guatimoc and Cacahuatan, 3 

8897 Dec 1941 

(A, BM, F, G, K, LE, MO, NY, P, S, U, US, type 

(9), Miranda 1737 (US); NW of San Fercollection 

of C. cornifolia); rd. Manaus-Itacoatiara, km nando, NW ofTuxtla, 1000 m, 2 Apr 1950 (9), Miranda 

65-70, 23 Oct 1963 (8), Oliveira 2764 (IAN); rd. Ma- 

6164 (US); nr. El Ocote, 30 km NW of Ocozocuantla, 

naus-Itacoatiara, km 61, Res. Flor. Walter Egler, 17 24 Mar 1951 (9), Miranda 6273 (US), GUERERO: Distr. 

Dec 1968 (6), Prance et al. 9054 Montes de 

(M, MO, NY, P, S, 

Oca, San Antonio, 18 Apr 1938 (9), Hinton 

U); rd. Manaus-Porto Velho, km 510, 5 km N of Rio 14018 (K, NY, P, US). JALISCO: Sierra Madre Occi- 

Purusinho, 17 Oct 1974 (6), Prance et al. 22999 (INPA, 

dental, NW of San Sebastian, Las Mesitas, Arroyo de 

NY, U, US). PARA: Rd. Cuiaba-Santar6m, km las 

1417, Caaitas, 1700 m, 15 Mar 1927 (6), Mexia 1872 (A, 

Rio Itapacura, 25 Nov 1977 (6), Prance et al. 25748 BM, F, GH, MO, NY, US). VERACRUZ: Paso del In- 

(MO, RB, U). 

genio, Jun 1971 (9), Calzada 325 (U); without locality, 

1930 (st), Purpus s.n. (B, F, K); Zacualpan, Mar 1908 

This species belongs to a group with two sta- (6), Purpus 5996 (BM, F, GH, MO, NY, US), Mar 1930 

mens and leaf margins which mostly are revolute (8), Purpus 11161 (K), Apr 1928 (6), Purpus 11162 (F, 

at the base, comprising e.g., C. 

type collection), Jun 1930 

microcarpa and 

(9), Purpus 11162 (A, K, 

MO), Sep 1930 (9), Purpus 11162 (C), Apr 1931 (9), 

C. microcephala. 

Purpus 11162 (C), May 1933 (6 and 9), Purpus 11162 

Prance et al. 9054 is aberrant in having (A), May 1934 (6 and 9), Purpus 11162 (K, US), Sep 

(sub)obovate leaves. 

1929 (8), Purpus 11182 (A). 

GUATEMALA. QUEZALTENANGO: Colomba, ca. 

1000 m, 28 Dec 1934 (6), Skutch 2023 (BM, F, GH, 

36. Coussapoa purpusii Standley, Publ. Field NY). SAN MARCOS: Finca El Porvenir, on Potrero Ma- 

Mus. Bot. 8: 6. 1930. Type. Mexico. Veracruz: tasan, Rio Cabuis, Volcan Tajumulco, 1000-1300 m, 

Zacualpan, Apr 1928 (6), Purpus 11162 12 Mar 

(ho- 

1940 (9), Steyermark 37602 (F), 25 Mar 1943 

lotype, F). Fig. 48. 

(st), Steyermark 52334 (F). 

The species appears to be related to C. parviceps. 

Tree, hemi-epiphytic or terrestrial, often (?) 

deciduous, up to 20 m tall. Leafy twigs 3-6 mm 

thick, glabrous or puberulous on the scars of the 

stipules. Lamina chartaceous, ovate to elliptic or 

to obtuse, occasionally obovate, 4-16 x 1-7 cm, 

apex acute to acuminate to obtuse, base obtuse 

sometimes subacute or subcordate, margin en- 

tire; both surfaces glabrous; lateral veins 4-6 pairs, 

curved, basal pair unbranched reaching the mar- 

gin below, at, or slightly above the middle of the 

lamina; intercostal venation plane; petiole 1-5 

cm long, glabrous; stipules 1-3 cm long, glabrous 

or sparsely to densely appressed-puberulous. 



2-4 cm long, glabrous; perianth ca. 1 mm high, 

glabrous or sparsely minutely puberulous; sta- 

mens three, far exceeding the perianth. Pistillate 

inflorescences unbranched; heads globose, 4-6 

mm, in fruit up to 10 mm diam.; peduncle 3-4 

37. Coussapoa scabra Akkermans & Berg, Proc. 

Kon. Ned. Akad. Wetensch. Ser. C 85(4): 460. 

1982. Type. Brazil. Rondonia: Rio Ouro Pre- 

to, affluent of Rio Pacaas Novos, 18 Sep 1923 

(9), Kuhlmann 470 (HJBR 19836) (holotype, 

RB; isotypes, RB, U). Fig. 49. 

Tree. Leafy twigs 2-5 mm thick, puberulous 

to subhirsute; internodes solid. Lamina charta- 

ceous, elliptic to ovate or obovate, 4-10 x 2.5- 

7 cm, apex acute to shortly acuminate, base ob- 

tuse to acute, margin entire; upper surface, sca- 

brous, lower surface, scabridulous, sparsely to 

rather densely puberulous to hirtellous; lateral 

veins 6-13 pairs, straight to curved, basal pair 

unbranched, reaching the margin far below the 

middle of the lamina; intercostal venation slight- 

ly prominent; petiole 1-3 cm long, densely pu-

FIG. 48. Coussapoa purpusii: 1, leafy twig with pistillate inflorescences (Hinton 14018); 2, leafy twig with 

staminate inflorescences (Skutch 2023).

92 

1cm 


FIG. 49. Coussapoa scabra: 1, leafy twig with pistillate inflorescences 

(Kuhlmann 470). 

berulous to hirtellous; stipules 0.5-0.7 cm long, 

subhirsute. Staminate inflorescences unknown. 

Pistillate inflorescences unbranched or poorly 

branched; heads 1-5, globose, 2-4 mm diam., in 

fruit up to 6 mm; (common) peduncle 1-3 cm 

long, puberulous; perianth ca. 1 mm high, glabrous. 

Interfloral bracts (sub)spathulate, minutely 

puberulous at the apex. 

Distribution (Fig. 8). Brazil, R6ndonia. Known 

only from the type collection. 

The position of this species, whose leaves have 

a characteristic scabrous and chartaceous lamina, 

is unclear. The shape and venation of the 

lamina show similarities to those found in C. 

parvifolia and C. macerrima. 

38. Coussapoa sprucei Mildbraed, Notizbl. Bot. 

Gart. Berlin 10:415.1928. Type. Brazil. Ama- 

zonas: "Rio Negra," probably Rio Taruma, 

Dec 1854 (9), Spruce 3782 (holotype, B; iso- 

types, BM, BR, C, G, GH, K, LE, NY, OXF, 

P). Fig. 50. 

Tree or shrub, mostly hemi-epiphytic, up to 

30 m tall. Leafy twigs 3-7 mm thick, puberulous 

to hirtellous to brownish hirsute, sometimes also


2 1cm 

FIG. 50. Coussapoa sprucei: 1, leafy twig with pistillate inflorescences (Spruce 3782); 2, leafy twig with 

staminate inflorescences 

(Prance et al. 14976). 

with white arachnoid hairs. Lamina coriaceous, 

oblong to elliptic, subovate or to subovate, 5-18 

x 3-8 cm, apex shortly acuminate to acute, base 

subcordate to rounded, margin entire; upper surface 

glabrous, lower surface puberulous in the 

areoles, hirtellous (to tomentellous) on the smaller 

veins, appressed-puberulous on the main veins, 

sometimes also with a few distinctly longer appressed 

hairs, the whole surface covered with 

rather dense to sparse (sub)persistent white 

arachnoid hairs; lateral veins 5-10 pairs, slightly 

curved, basal pair unbranched or inconspicuously 

branched, reaching the margin below the 

middle of the lamina; intercostal venation (very) 

prominent; petiole 1-4 cm long, puberulous, often 

also having distinctly longer straight hairs and 

white arachnoid hairs; stipules 0.5-1.5(-2) cm 

long, brownish sericeous to subhirsute, terminal 

buds often more or less swollen. Staminate inflorescences 

branched; heads many, globose, ca. 

3-4 mm diam.; common peduncle (2-)4-8 cm 

long, puberulous to hirtellous; perianth ca. 1 mm 

high, glabrous; stamens three, far exceeding the 

perianth. Pistillate inflorescences branched; heads 

2-5, globose, ca. 3-5 mm in fruit up to 8 mm 

diam.; common peduncle 2-8 cm long, ca. 1 mm 

thick, puberulous to hirtellous; perianth ca. 1 

mm high, glabrous. Interfloral bracts spathulate 

to subpeltate, puberulous, mostly conspicuous. 

Distribution (Fig. 8). Brazil, Amazon basin, 

near Manaus, and in the basin of the Rio Trombetas; 

in forest of terra firme. 

93 

Specimens studied. BRAZIL. AMAZONAS: Nr. Manaus, 

Rio Taruma, 27 Oct 1943 (6), Ducke 1354 (A, 

F, NY, R, RB, US); nr. Manaus, C6lonia Joao Alfredo, 

27 Sep 1946 (8), Ducke 1999 (A, NY, R, RB, U); nr. 

Manaus, Rio Taruma, Cachoeira Grande, 17 Oct 1929 

(9), Ducke (HJBR) 23609 (RB); Manaus, 17 Oct 1929 

(9), Killip & Smith 30179 (NY); Rio Cuieras, 2 km 

below mouth ofRio Brancinho, 27 Sep 1971 (6), Prance 

et al. 14976 (K, M, MO, NY, S, U), 12 Sep 1973 (8), 

Prance et al. 17847 (F, K, M, MO, NY, P, S, U); "Rio 

Negro," possibly Rio Taruma, Dec 1854 (9), Spruce 

3782 (B, BM, BR, C, G, GH, K, LE, NY, OXF, P, 

type collection). PAIA: Rio Mapuera, 11 Dec 1907 (2), 

Ducke (HAMP) 9096 (MG, US). 

Coussapoa sprucei appears to be closely related 

to C. leprieurii. It differs distinctly from the latter 

in the smooth and glabrous upper surface of the 

lamina. Other differences are of little signifi- 

cance. C. sprucei also resembles the Central 

American C. oligocephala, from which it differs 

mainly in the length of the stipules and to a lesser 

extent in the nature of the indumentum. This 

group of three species is probably related to the 

large-leaved taxa C. nymphaeifolia and C. lon- 

gepedunculata, judging from the similarities in 

indumentum, leaf venation, and number of sta- 

mens. 

39. Coussapoa tessmannii Mildbraed, Notizbl. 

Bot. Gart. Berlin 10: 413. 1928; Macbride, 

Publ. Field Mus. Bot. 13(2.2): 298. 1937. Type. 

Peru. Loreto: Mouth of Rio Santiago, 3 Dec 

1924 (p), Tessmann 4673 (holotype, B; iso- 

types, G, S). Fig. 51.


FIG. 51. Coussapoa tessmannii: 1, leafy twig with pistillate inflorescences (Krukoff 8994); 2, staminate 

inflorescences (Krukoff 1187).


Tree or shrub, (mostly) hemi-epiphytic, up to PERU. LORETO: Prov. Maynas, Rio Yaguasyuca, af- 

20 m tall. Leafy twigs 5-13 mm, brown 

fluent of Rio 

puber- 

Ampiyacu, nr. Brillo Nuevo, 2 Mar 1978 

(2), Balick et al. 1021 (MO, U); road Zungarochaulous 

to strigillose to hirtellous to strigose or to Puerto Almendra, 3 Dec 1964 (2), Dodson et al. 3017 

(sub) velutinous, often also with a dense covering (F); Rio Yavari, behind Angamo Garrison, 5 Oct 1973 

of (long) dark brown pluricellular hairs and (a), Lleras et al. P17178 (F, K, NY, P, S, U); Puerto 

sometimes whitish arachnoid hairs as well. Lam- Almendra, nr. Rio Nanay, 23 Feb 1977 (a), Revilla 

2378 

ina coriaceous, ovate to elliptic, 13-32 x 11-20 

(U); mouth of Rio Santiago, 3 Dec 1924 (2), 

Tessmann 4673 (B, G, S, type collection). 

cm, apex subacute to obtuse to rounded, base BRAZIL. AMAZONAS: Rio Negro, Uaup6s, Jauarite, 

obtuse to subcordate, margin entire to subcren- 23 Aug 1945 (9), Froes 21264 (NY); Mun. Sao Paulo 

ate; upper surface glabrous, lower surface densely 

de Olivenca, Belem Creek, 26 Oct-11 Dec 1936 (2), 

minutely puberulous in the areoles and on the Krukoff8994 (A, G, K, LE, NY, P, S, U); road Manaus-Porto 

Velho, km 379, Rio Jutai, 12 Oct 1974 

reticulum, to (sparsely) hirtellous on the (paral- (9), Prance et al. 22836 (MG, NY, U); Rio Javari, 7 

lel) tertiary venation, appressed-puberulous to hours above Paumari, 16 Oct 1976 (2), Prance et al. 

strig(ill)ose on the midrib and lateral veins, the 23846 (NY, U). PARA: Cupary, Sep 1931 (6), da Costa 

whole surface with rather sparse to dense sub- 90 (IAN); upper Rio Cupary, Sep 1931 (a), Krukoff 

1147 

persistent white to pale brown arachnoid 

(A, BM, G, K, NY, P, S, U), Krukoff 1187 (A, 

hairs; BM, G, K, MO, NY, S, U); road Santarem-Palhao, 

lateral veins 9-16 pairs, straight, main basal pair km 70, 15 Sep 1969 (8), M. Silva et al. 2591 (F, K, 

branched, sometimes faintly so (in small leaves MG, NY, S, U, US). RONDONIA: Porto Velho, 15 Sep 

sometimes unbranched), reaching the margin far 1975 (a), Mota et al. 154 (U). 

below or sometimes just below the middle of the In the vegetative parts C. tessmannii shows 

lamina; intercostal venation very to slightly similarities to C. villosa, but it is distinct from 

prominent; petiole 3.5-9 cm long, 2-5 mm thick, the latter in having a ramified, and thus pluribrown(ish) 

puberulous to hirtellous to subvelucapitate, 

pistillate inflorescence and a unistamitinous, 

often also with dark brown pluricellular nate flower. C. tessmannii shows distinct affinhairs 

and sometimes arachnoid hairs; stipules 2- ities with C. cinnamomea, e.g., in the relatively 

3(-9) cm long, yellowish to brown strigose to thick peduncle and branches of the pistillate insubhirsute 

or also with dark brown pluricellular florescence, and it can be easily confused with 

hairs. Staminate inflorescences branched; heads C. nitida. Several collections made in the Rio 

many, crowded at the end of the branches, Tapajos area (da Costa 90, Krukoff 1147 and 

(sub)globose, 2-4 mm diam., often fused; com- 1187, and M. Silva et al. 2591) differ from the 

mon peduncle 1-3 cm long, brown puberulous other collections in the presence of a dense covto 

hirtellous to short-velutinous or also (densely) ering of dark brown moniliform pluricellular (= 

covered with dark brown pluricellular hairs; peri- granular) hairs on the leafy twig, petiole, and 

anth ca. 1 mm high, puberulous; stamen one, inflorescence. Moreover, the lamina tends to be 

just exceeding the perianth. Pistillate inflores- elliptic rather than ovate (as is normal in the 

cences branched; heads 2-7, (sub)globose, 5-10 

species). In the vegetative parts, these collections 

mm, in fruit up to 15 mm diam.; common pe- show striking similarities to the aberrant pistilduncle 

2-3 cm long, peduncle and branches 2- late collection (Prance et al. 26402) assigned to 

4 mm thick, appressed puberulous to hirtellous C. villosa (see p. 104). 

to short-velutinous or also with dark brown pluricellular 

hairs; perianth ca. 1 mm high, (almost) 

glabrous, sometimes muriculate. 

40. 

Interfloralbracts 

Coussapoa trinervia Spruce ex Mildbraed, 

(sub)spathulate to subpeltate, puberulous at the 

Notizbl. Bot. Gart. Berlin 10:416. 1928. Type. 

apex; extrafloral bracts often present on the 

Venezuela. Terr. Fed. Amazonas: Rio Casibranches 

of staminate inflorescences. 

quiare, between Lago de Vasiva and Rio Pa- 

Distribution (Fig. 9). Amazon Basin cimoni, 1853-1854 

(Brazil, 

(2), Spruce 3464 (holotype, 

Peru, and Colombia); in upland and riverside B; isotypes, BM, BR, G, GH, K, LE, NY, OXF, 

forest. 

P). Fig. 52. 

Specimens examined. COLOMBIA. VAUPES: Rio 

Tree, terrestrial, up to 20 m tall, mostly with 

Papuri, nr. Montfort Mission, 3 Sep 1943 (s), P. H. stilt roots. Leafy twigs 3-8 mm thick, glabrous. 

Allen 3103 (COL). 

Lamina coriaceous, subobovate to obovate to 

95

2 

1cm 

FIG. 52. Coussapoa trinervia: 1, leafy twig with pistillate inflorescences (Berg et al. P18453); 2, leafy twig 

with staminate infiorescences (Schunke 371).


oblong to elliptic, 4-14 x 1.5-7 cm, apex obtuse 

to shortly acuminate, base (sub)acute to cuneate, 

margin entire; both surfaces glabrous; trinervate, 

the single pair of lateral veins, unbranched, 

reaching the margin near or at the apex of the 

lamina; intercostal venation prominant; petiole 

1-4 cm long, glabrous; stipules 0.5-1.5 cm long, 

glabrous or occasionally appressed-puberulous. 


often partly fused, globose, 2-5 mm diam.; com- 

mon peduncle 1-3 cm long, glabrous or sparsely 

puberulous; perianth ca. 1 mm high, glabrous; 

stamen one, just exceeding the perianth. Pistil- 

late inflorescences unbranched; heads globose, ca. 

5-7 mm, in fruit up to 10 mm diam.; peduncle 

1-3 cm long, glabrous or sparsely puberulous; 


bracts subspathulate to peltate, sparsely puber- 

ulous at the apex. 

Distribution (Fig. 9). Amazon Basin (Brazil, 

Peru, Ecuador, Colombia, and Venezuela); the 

main area apparently in the north-western part 

of Amazonia-elsewhere in smaller, more re- 

stricted, areas; on periodically inundated banks 

(always?) of black-water rivers. 

Rio Mazan, Gamitanococha, 14 Mar 1935 (6), Schunke 

371 (A, F, NA, NY). 

BRAZIL. AMAZONAS: Rio Negro Basin, Tapuruquara, 

21 Oct 1971 (9), Prance et al. 15764 (F, GH, 

K, NY, S, U); Rio Uaup6s, Panur6, Oct 1852-Jan 1853 

(Y), Spruce 2616 (B, BM, BR, C, G, GH, K, LE, NY, 

OXF, P). MARANHXO: Turiaga, Astonas, 6 Dec 1978 

(2), Rosa et al. 2874 (U). MATO GROSSO: Rio Aripuafia, 

nr. Humboldt Center, 10?12'S, 59?21'W, 10 Oct 1973 

(a), Berg et al. P18409 (F, K, MO, NY, P, U, US), 12 

Oct 1973 (2), Berg et al. P18453 (K, MO, NY, S, U, 

US); Rio Juruena, Cachoeira de Sao Joao da Barra, 3 

Jun 1977 (a), Rosa et al. 2053 (NY); Rio Juruena, 

Igarape Chuini, 15 Jul 1977 (2), M. G. Silva et al. 3345 

(U). PARA: Mun. Itapiranga, Rio Uamata, nr. Igarap6 

Santa Luzia, 16 Aug 1974 (2), Cid et al. 435 (U), nr. 

Cachoeira do Tucumari, 19 Aug 1974 (Y), Cid et al. 

511 (U); Belem, between Val de Caes and Sao Joaquin, 

7 Nov 1922 (2), Ducke (HJBR) 18455 (RB); Belem, 

Sep-Oct 1961 (st), Pires 51736 (NY). 

This remarkable species, probably the only 

truly terrestrial species, resembles terrestrial fig 

trees (like F. trigona L. f.) in habit and has me- 

Specimens studied. COLOMBIA. CAQUETA: Rio Caqueta, 

Solano, 8 km SE ofTres Esquinas, below mouth 

of Rio Orteguaza, 7 Mar 1945 (st), Little et al. 9629 

(COL); Rio Apaporis, between Rio Pacoa and Rio 

Kananari, Soratama, 16 Aug 1961 (d), Schultes et al. 

13589 (GH, U). 

VENEZUELA. AMAZONAS: Rio Manariche, Kalohi-teri, 

- lastomataceous leaves. It appears to be confined 

to periodically inundated banks of black-water 

rivers; this may explain the disjunct distribution. 

Coussapoa trinervia appears to be closely related 

to C. angustifolia, as the two species have many 

characters in common. The difference in the 

number of stamens suggests that C. trinervia and 

C. cinnamomifolia are not related, in spite of 

similarities in the leaf venation. 

41. Coussapoa vannifolia Cuatrecasas, Caldasia 

(2), Lizot 83 (VEN); Rio Yatua, just above 7: 296. 1956. 

Piedra Arauicaua, 28 Sep 1957 (2), Maguire et al. 

Type. Colombia. Valle: Rio An- 

41620 

(K), Esmeralda-Ocana, Mavaca, 23 Apr 1968 (2), Me- chicayf, between Sabuletas and Quebrada Tadina 

397 (VEN); Rio Casiquiare, between Lago de Va- tabro, 28-29 Sep 1946 (9), Cuatrecasas 22048 

siva and Rio Pacimoni, 1853-1854 (9), Spruce 3464 (holotype, F). Fig. 53. 

(B, BM, BR, G, GH, K, LE, NY, OXF, P, type collection); 

Rio Yatua, just above Piedra Arauicaua, 16 Jul Coussapoa rotunda Little, Phytologia 18: 196. 1969. 

1959 (9), Wurdack et al. 43485 (K). 

Type. Ecuador. Esmeraldas: Junction of Rio Hoja 

ECUADOR. NAPO: Rio Cuyabeno, ca. 0?10'S, Blanca and Rio Hualpi, 50 km S of Borb6n, 14 Sep 

76?55'W, 16 Feb 1980 (d), Berg & Akkermans 1038 1965 (6), Little & Dixon 21056 (holotype, US; iso- 

(U), 17 Feb 1980 (2), Berg & Akkermans 1048 (U), 18 types, NY, Q). 

Feb 1980 (2), Berg & Akkermans 1058 (U); Laguna 

Cuyabeno, 7 Aug 1980 (9), Jaramillo et al. 2888 (AAU), 

Tree, terrestrial or hemi-epiphytic, up to 20 m 

PASTAZA: Rio Capihuari, 23 Jul 1980 (st), Ollgaard et tall. Leafy twigs 5-9 mm thick, white to brown 

al. 35068 (AAU). 

hirtellous to hirsute. Lamina coriaceous, 

PERU. 

broadly 

LORETO: Quebrada Cuninico, 7 Jul 1972 (2), ovate to cordiform to subreniform 

Croat 

or to 

17784 

subor- 

(F, NY); Rio Tacsha Curary, 18 Sep 1972 

(a), Croat 20411 (C, F, GH, L, MO, NA, NY, P, 

bicular, 4-22 x 4-25 

S); 

cm, apex rounded to ob- 

Rio Yavari, Emilia, above Atalaia del Norte, 22 Nov tuse, base cordate, occasionally obtuse, margin 

1977 (2), Gentry et al. 20766 (MO, U); Prov. Maynas, entire or subcrenate; upper surface glabrous, low- 

Rio Mom6n, tributary of Rio Nanay, nr. Iquitos, 9 er surface rather 

Dec 1979 (2), J. Jones et 

densely white puberulous to hiral. 

9788 (LAM, U); Rio Na- tellous on the main 

nay, Quebrada Morropon, 26 Jun 1972 (2), McDaniel 

veins, otherwise glabrous or 

16344 (NA); Rio Nanay, Mapa Cocha, 17 Mar 1976 sparsely puberulous to hirtellous on the tertiary 

(a), McDaniel et al. 20555 (NA); Rio Nanay, Quebrada (parallel) venation; lateral veins 5-9 pairs, in- 

Morropon, 31 May 1976 (2), Rimachi 2297 (F, MO), cluding 2-4 pairs of basal veins, basal veins

FG53 Cosaovanfla1leftwgwtstmntifoecne Apud141' 

....... cm .... I 

FIG. 53. Coussapoa vannifolia: 1, leafy twig with staminate infiorescences (Asplund 1640


branched, the upper pair of basal veins reaching 

the margin at or above the middle of the lamina, 

the other lateral veins often poorly branched (furcate); 

intercostal venation almost plane; petioles 

1-6 cm long, (rather) densely white to brownish 

hirtellous to hirsute; stipules 2-8 cm long, densely 

puberulous to hirtellous to strigillose, mostly 

also with brown pluricellular hairs. Staminate 

inflorescences branched; heads many, globose, ca. 

1.2 mm diam.; common peduncle 1-2 cm long, 

hirtellous to subtomentose; perianth ca. 1 mm 

high, apex minutely puberulous; stamens three, 

just exceeding the perianth. Pistillate inflorescences 

branched; heads 3-5, free or sometimes 

partly fused, (sub)globose, 6-9 mm diam.; common 

peduncle 1.5-2.5 cm long, hirtellous to subtomentose; 

perianth ca. 1 mm high, papillate. 

Interfloral bracts present, subpeltate, at the apex 

minutely puberulous, in pistillate inflorescences 

often only a few. 

Distribution (Fig. 9). Ecuador and Colombia, 

Pacific coastal region; in lowland rain forest. 

Specimens studied. COLOMBIA. CAUCA: Rio Timbiqui, 

- Coussapoa vellerea Klotzsch, Linnaea 20: 527. 1847; 

Miquel, Fl. bras. 4(1): 139. 1853; Macbride, Publ. 

Field Mus. Bot. 13(2.2): 299. 1937. Syntypes. Peru, 

Huanuco: Macora, 

(8), Lehmann BT948 (B, GH, K, NY). VALLE: 

Rio Callima, between Pailon and El Coco, 23 May 

1946 (Q), Cuatrecasas 21256 (F); Rio Anchicaya, between 

Sabuletas and Quebrada de Tatabro, 28-29 Sep 

1946 (Q), Cuatrecasas 22048 (F, type collection). 

ECUADOR. ESMERALDAS: Junction of Rio Hoja 

Blanca and Rio Hualpi, 14 Sep 1965 (6), Little et al. 

21056 (NY, Q, US, type collection of C. rotunda). Los 

Rios: Rio Palenque, along road after crossing Rio Bimbe 

and Rio Waija, 7 Oct 1976 (Q), Dodson et al. 6513 

(MO, QCA). PICHINCHA: Nr. Santo Domingo de los 

Colorados, 18 May 1955 (a), Asplund 16401 (S), 20 

Aug 1920 (st), Benoist 3042 (P, U), 5 Feb 1979 (Q), 

Dodson 7416 (SEL, U), 5 Feb 1979 (6), Dodson et al. 

7616 (MO), 9 Apr 1892 (9), Sodiro 193/12 (B). 

In the shape and venation of the leaves, C. 

vannifolia is reminiscent of broad-leaved forms 

of C. asperifolia ssp. magnifolia but also of C. 

batavorum. Because of the difference in the number 

of stamens, a taxonomic relationship with C. 

asperifolia is unlikely. 

42. Coussapoa villosa Poeppig & Endlicher, Nov. 

gen. sp. pl. 2: 33, t. 147. 1838; Trecul, Ann. 

Sci. Nat. Bot. S6r. 3, 8: 99. 1847; Miquel, Fl. 

bras. 4(1): 138. 1853; Macbride, Publ. Field 

Mus. Bot. 13(2.2): 299. 1937. Type. Peru. 

Huanuco: Casapi (fide Macbride, 1937) ($ and 

2 ?) Poeppig s.n. (holotype, W, destroyed; isotypes 

(6), B, OXF). Fig. 54. 

- (6 and 2), Ruiz & Pavon s.n. 

(or 3) (holotype (2), B; isotype (2), FI; isotypes (8), 

BR, G, OXF, US). 

Coussapoa martiana (Miquel, Fl. bras. 4(1): 132, t. 42. 

1853. Type. Brazil. Without locality ("prov. Paraensi 

et Rio Negro"), - (2), Martius s.n. (holotype, M). 

Coussapoa subincana Miquel, Fl. bras. 4(1): 134, t. 45. 

1853. Type. Brazil. Without locality, - (6), Martius 

s.n. (holotype, M; isotypes, BR, M, U). 

Coussapoapanamensis Pittier, Contr. U.S. Natl. Herb. 

18: 226. 1917; Woodson & Schery, Ann. Missouri 

Bot. Gard. 47: 169, t. 59. 1960; Burger, Fieldiana 

Bot. 40:135, t. 22. 1977. Type. Panama. Col6n: Rio 

Fat6, Jul 1911 (9), Pittier 3892 (holotype, US; isotypes, 

B, GH, NY). 

Coussapoa donnell-smithii Mildbraed, Notizbl. Bot. 

Gart. Berlin 10: 414. 1928. Type. Costa Rica. Cartago: 

Rio Turrialba, Mar 1894 (2), Donnell Smith 

4826 (holotype, US; isotypes, GH, K, US). 

Coussapoa grandiceps Killip in Macbride, Publ. Field 

Mus. Bot. 13(2.2): 297. 1937. Type. Peru. Loreto: 

Lower Rio Huallaga, Yan6n, 5-11 Sep 1929 (2), Killip 

& Smith 29246 (holotype, US). 

Coussapoa standleyi Macbride, Publ. Field Mus. Bot. 

13(2.2): 298. 1937. Type. Peru. Huanuco: Huamalies, 

between Monzon and Rio Huallaga, - (9), Weberbauer 

3702 (holotype, B, destroyed; isotype, G). 

Coussapoa araneosa Standley, Publ. Field Mus. Bot. 

17: 158. 1937. Type. Brazil. Amazonas: Mun. Humayta, 

Tres Casas, 14 Sep-11 Oct 1934 (2), Krukoff 

6524 (holotype, F; isotypes, A, BM, K, LE, MO, NY, 

S, U, US). 

Coussapoa boliviana Standley, Publ. Field Mus. Bot. 

17: 159. 1937. Type. Bolivia. Santa Cruz: Prov. Ichilo, 

Rio Vibora, 5 Oct 1926 (2), Steinbach 7567 (holotype, 

F; isotypes, A, GH, K, MO, S, U). 

Coussapoa eggersii Standley, Publ. Field Mus. Bot. 17: 

160. 1937. Type. Ecuador. Manabi: El Recreo, Dec 

1891 (6), Eggers 14165 (holotype, F; isotypes, probably 

in A, BR, L, M, US). 

Coussapoa embirana Standley, Publ. Field Mus. Bot. 

17:161. 1937. Type. Brazil. Amazonas: Basin of Rio 

Jurua, nr. mouth of Rio Embira (tributary of Rio 

Tarauaca), 26 Jun 1933 (6), Krukoff 4994 (holotype, 

NY; isotypes, A, G, K, U, US). 

Coussapoa lawrancei Standley, Publ. Field Mus. Bot. 

17: 164. 1937. Type. Colombia. Boyaca: El Humbo, 

130 km N of Bogota, ca. 1000 m, 26 Apr 1933 (6), 

Lawrance 769 (holotype, F; isotypes, A, B, F, FI, G, 

K, MO, S, US). 

Coussapoa lehmannii Standley, Publ. Field Mus. Bot. 

17: 164. 1937. Type. Ecuador. Azuay: Western Andes 

of Cuenca, Chacayacu, - (2), Lehmann 5606 

(holotype, F; isotype, K). 

Coussapoa subcrenata Standley, Publ. Field Mus. Bot. 

17: 166. 1937. Type. Brazil. Amazonas: Mun. Tefe, 

Rio Solim6es, Paleta, 21 May 1933 (6), Krukoff4518 

(holotype, F; isotypes, A, BM, G, K, M, MO, NY, 

S, U, US). 

Coussapoa danielis Cuatrecasas, Caldasia 7: 290. 1956. 

99


FIG. 54. Coussapoa villosa: 1, leafy twig with pistillate inflorescences (Steyermark et al. 95170); 2, twig with 

stipules and staminate inflorescences (Elias 631); 3, pistillate inflorescences (Donnell Smith 4826). 

3 

1 cm


Type. Colombia. Antioquia: Jardin, 1800 m, Dec 

1941 (2), Bro. Daniel 2652 (holotype, US; isotypes, 

COL, F). 

Coussapoa macarenensis Cuatrecasas, Caldasia 7: 292. 

1956. Type. Colombia. Meta: Sierra de la Macarena, 

Caino Entrada, 24 Dec 1949 (5), Phillipson & Idrobo 

1914 (holotype, BM; isotypes, COL, US). 

Coussapoa macarenensis Cuatrecasas var. antioquiensis 

Cuatrecasas, Caldasia 7: 293. 1956. Type. Colombia, 

Antioquia: Rio Cauca, Puento Valdiva, 17- 

20 Feb 1942 (2), Metcalf& Cuatrecasas 30093 (holotype, 

F; isotypes, A, MO, US). 

Coussapoa mutisii Killip & Cuatrecasas, Caldasia 7: 

293. 1956. Type. Colombia. Without locality, - lutinous or to (sub)hirsute, often also with reddish-brown 

pluricellular hairs and sometimes 

arachnoid hairs; perianth ca. 1 mm high, densely, 

minutely puberulous; stamens two, (rather) far 

exceeding the perianth. Pistillate inflorescence 

mostly unbranched, sometimes poorly branched; 

heads 1(-4), free or occasionally partly fused, 

(sub)globose, ca. 5-10 mm, in fruit up to 40 mm 

diam.; (common) peduncle (1-)2-13 cm long, 1- 

4 mm thick, densely to sparsely puberulous to 

hirtellous to shortly velutinous or to (sub)hirsute, 

(2), 

Mutis 646 

often also with reddish-brown 

(holotype, US). 

pluricellular hairs 

Coussapoa planitiensis Cuatrecasas, Caldasia 7: 295. and sometimes arachnoid hairs; perianth ca. 1- 

1956. Type. Colombia. Vaupes: Rio Guayabero, 8 2 mm high, densely minutely white to brownish 

Nov 1939 (6), Cuatrecasas 7510 (holotype, US). puberulous; fruiting perianth at the apex green, 

below the apex orange. Interfloral bracts 

(sub)spathulate to subpeltate, white to brownish 


Distribution (Fig. 9). Central America (from 

Guatemala to Panama), the Andean region (Ecuador 

to Venezuela), extending to the coastal 

mountain range of Venezuela, and wide-spread 

(especially) in the (Upper) Amazon Basin; in upland, 

riverine, and mountain forest, up to 2400 

m; often in secondary vegetation. 

Tree or shrub, hemi-epiphytic or terrestrial, up 

to 35 m tall. Leafy twigs 5-15 mm thick, sparsely 

to densely white (to brown) puberulous to hirtellous 

to (sub)hirsute to (sub)villous. Lamina 

coriaceous, broadly ovate to subovate, sometimes 

to elliptic, occasionally to obovate, 6-60 

x 5-40 cm, apex obtuse to acute, sometimes 

rounded to emarginate, base obtuse to cordate 

(sometimes deeply cordate with overlapping 

lobes), margin entire to subcrenate; upper surface 

glabrous, lower surface sparsely to densely minutely 

puberulous in the areoles and mostly also 

on the reticulum, hirtellous to hispidulous on the 

(parallel) tertiary venation, sometimes puberu- 

Specimens studied. GUATEMALA. IZABAL: Peten 

road, 2 km from Fronteras, Rio Juan Vicente, 11 May 

1971 (8), Contreras 10763 (DUKE, MO, P, US); Quirigua, 

15-31 Mar 1922 (6), Standley 23766 (GH, NY), 

lous to (sub)hirsute, on the midrib and the lateral 26-27 Apr 1939 (9), Standley 72288 (F, NY); Lago 

veins sparsely, sometimes rather densely, puber- Izabal, between Punta dos Reales and Punta de Leulous 

to hirtellous or to (sub)hirsute, the whole 

chuga, 17 Apr 1940 (2), Steyermark 39618 (F). 

HONDURAS. ATLANTIDA: La Ceiba, 22 Sep 1916 

surface covered with sparse to dense whitish to (9), Dyer A85 (F, US); Lancetilla Valley, nr. Tela, 6 

brownish subpersistent arachnoid hairs; lateral Dec 1927-20 Mar 1928 (st), Standley 53230 (A, F, 

veins (7-)9-24, straight or slightly curved, main US), (9), Standley 53991 (A, F, US), (st), Standley 54325 

basal pair branched, reaching the margin below (A, F), (2), Standley 55155 (A, F, US), (st), Standley 

55445 (A, F, 

the middle of the lamina, in 

US), (6), Standley 56623 

large leaves 

(F). COMAYAGUA: 

some- Lake Yojoa, Pito Solo, nr. Pito Solo, 16 Apr 1951 (6), 

times also with other lateral veins poorly P. H. Alien 6186 (F, GH), 8 Aug 1932 (st), Edwards 

branched (furcate); intercostal venation more or 400 (A, F, US); 15 km E of Lake Yojoa, 20 May 1974 

less prominent; petiole (2-)3-20 cm long, white (6), Hazelett s.n. (MO); nr. Pito Solo, 14 Aug 1973 (2), 

Hazelett 664 

(to yellowish) puberulous to hirtellous to 

(MO); Lake Yojoa, 29 Jul-10 Aug 1951 

(6), Howard et al. 605 (A); nr. Pito Solo, 18 Apr 1945 

(sub)hirsute or to (sub)villous; stipules 3-20 cm (2), J. V. Rodriguez 2929 (F); Rio Tempemechin, ca. 

long, puberulous to hirtellous to strigose to hir- 6 km N of Pito Solo, 15 Apr 1951 (d), L. O. Williams 

sute, mostly also with dense (moniliform) red- et al. 18014 (F, GH). COPAN: Rio Copan, 2 mi E of 

dish-brown pluricellular hairs and sometimes Copan ruins, 27 Aug 1975 (8), Molina et al. 30747 (F, 

MO). CORTES: Naciemento del Rio 

arachnoid 

Lindo, nr. 

hairs. Staminate 

Jaral, 9 

inflorescence Apr 1947 (6), Standley et al. 7081 (F); Rio Londo, 8 

branched, occasionally unbranched; heads (1-)2- Apr 1945 (6), L. O. Williams et al. 12394 (F, GH); Rio 

30(-ca. 50), often partly fused, (sub)globose to Lindo, nr. Carrizal, 12 Apr 1951 (st), L. O. Williams 

ovoid to oblate, often more or less irregular in et al. 17817 (F), GRACIAS A Dios: Rio Platano, 0-4 

hours 

shape and dimensions, ca. 5-10(-20) mm upstream Ras, 23 May 1973 (9), Gentry et al. 

diam.; 7530 (MO, U). 

common peduncle 2-6 cm long, densely to NICARAGUA. BLUEFIELDS: Finca Santa Rosa, ca. 

sparsely puberulous to hirtellous to shortly ve- 2.5 km ENE of Rama, 5 Apr 1966 (6), Proctor et al. 

101


27336 (F, US). CHONTALES: Nr. La Libertad, 29 May- Cricamola Valley, 17 Feb-l Mar 1928 (8), Cooper 538 

1 Jun 1947 (st), Standley 8861 (F). JINOTEGA: Cord. (F, K); Changuinola Valley, 10 Jan 1924 (2), Dunlap 

Isabelia, Macizos de Penias Blancas, 7 May 1976 (9), 299 (F, US); nr. Chiriqui Lagoon, 9 Nov 1940 (st), Von 

Neill 7179 (MO). MATAGALPA: Rio Yasica, 20 km E Wedel 1548 (S), 22 Nov 1940 (6), Von Wedel 1733 

of Matagalpa, 25 May 1977 (2), Neill 1984 (U); 10-15 (GH, US), 26 Oct 1941 (9), Von Wedel 2879 (GH, MO, 

km NE of Matagalpa, 16 Jan 1963 (2), L. O. Williams US); Rio Cricamola, between Finca St. Louis and Konet 

al. 24005 (F). NUEVA SEGOVIA: Rio Solonli, 5 km kintoe, 12-16 Aug 1938 (6), Woodson et al. 1893 (F, 

N of Jalapa, 5 Apr 1977 (8), Neill 1653 (U). ZELAYA: MO, NA, NY). CANAL ZONE: Barro Colorado Island, 

Comarca El Hormiguero, W of Rio Uli, 6 May 1977 7 Feb 1969 (2), Croat 7768 (F, MO), 10 Feb 1969 (6), 

(6), Neill 1919 (U). 

Croat 7839 (MO), 21 Jun 1971 (2), Croat 15064 (DUKE, 

COSTA RICA. ALAJUELA: Between La Palma and F, GH, MO, NY), 25 Jun 1968 (2), Foster 631 (DUKE); 

Rio Platanillo, 19 Feb 1964 (6), Jimenez 1750 (US); Rio Providencia, 5 Dec 1973 (st), Gentry 8707 (MO, 

NE-base of Volcan Arenal, 5 km W of Fortuna San U); Gatun, 4 Mar 1860 (9), Hayes 1008 (NY); Mindi 

Carlos, 29 Apr 1972 (6), Lent 2520 (F, MO, U, US); Hills, 28 Mar 1956 (st), Johnston 1725 (A); Barro Col- 

N side of Volcan Arenal, Quebrada Guillermina, 21 orado Island, 19 Mar 1932 (9), Shattuck 832 (F, MO); 

Apr 1973 (st), Lent et al. 3388 (F). CARTAGO: Road nr. Fort Sherman, 15 Jan 1924 (st), Standley 30924 

Turrialba-Moravia de Chirrip6, between Tuis and Bajo (US); Barro Colorado Island, 18-24 Nov 1925 (2), 

Pacuare, 15 Nov 1975 (8), Burger et al. 10040 (F); Standley 41170 (US), 12 Mar 1931 (6), Wilson 127 (F, 

Turrialba, 10 Dec 1952 (2), Cordoba 295 (NA); Rio MO), 6 Feb 1932 (6), Woodworth et al. 445 (A, F, MO), 

Turrialba, Mar 1894 (2), Donnell Smith 4826 (GH, K, 22 Feb 1932 (9), Woodworth et al. 660 (A, F, MO). 

US, type collection of C. donnell-smithii); Atirro, Apr CHIRIQUI: La Fortuna hydroelectric project, 1200 m, 

1896 (2), Donnell Smith 6770 (BM, F, GH, K, US); 22 Mar 1978 (2), Hammel 2176 (U). COLON: Rio 

Rio Raventaz6n, nr. Turrialba, 13 Feb 1971 (6), Gilles Guache, 1 Oct 1972 (st), Gentry 6314 (MO); road to 

et al. 10191 (F, GH); Rio Pejibaye, between Rio Taus Portobelo, Villa Alondra, 13 Apr 1971 (6), Lao et al. 

and Quebrada Azul, 28 May 1972 (2), Lent 2535 (F, 21 (DUKE, MO); Rio Fat6, Jul 1911, (2), Pittier 3892 

MO, U); Turrialba, May 1847 (st), Oersted 14317 (C, (B, GH, NY, US, type collection of C. panamensis). 

part of the type collection of Urostigma intermargin- PERLAS ARCHIPELAGO: San Jose Island, 28 May 1945 

ale), May 1898 (2), Pittier 12288 (B, BM, GH, P, US), (2), Erlanson 240 (GH, NA, NY), 29 Mar 1945 (9), 

1 Sep 1920 (st), Rowle et al. 856 (NY, US); nr. Pejivalle, Johnston 580 (GH), 2 Apr 1945 (6), Johnston 606 (FI, 

Jan 1940 (6), Skutch 4612 (A, MO, NA, US), 7-8 Feb GH, MO), (st), Johnston 608 (GH), 5 Apr 1945 (2), 

1926 (6), Standley et al. 47791 (US). GUANACASTE: El Johnston 652 (BM, DUKE, FI, GH, MO, P, S), 10 Apr 

Arenal, 18-19 Jan 1926 (2), Standley et al. 45182 (F, 1945 (2), Johnston 678 (GH). 

US); nr. Tilerfn, 10-31 Jan 1926 (6), Standley et al. COLOMBIA. Without locality, 

44550 (US), Standley et al. 44968 (F, US), (2), Standley 

et al. 45661 (US), 3 Jan 1964 (2), L. O. Williams et al. 

26529 (F, US). LIMON: Road Bribri-Bratsi, Rio Sixaola, 

12 Feb 1977 (8), Burger et al. 10474 (DUKE, F, 

MO); Las Diamantas, Rubber Expt. Station, 5 May 

1943 (st), Dayton 3019 (F); between Portete and Moin, 

13 Feb 1965 (2), Jimenez 2887 (F); nr. mouth of Rio 

Blanco, N of Moin, 13 Feb 1965 (9), Lent 349 (COL); 

ca. 4.5 km N of El Carmen, 17 Mar 1973 (st), Lent 

3279 (F); Zent, Oct 1904 (6 and 2), Pittier s.n. (GH, 

US); affluence of Rio Parismina and Rio Reventaz6n, 

Suerre, 3 Oct 1951 (st), Shank et al. 4294 (F); nr. Guapiles, 

12-13 Mar 1924 (6), Standley 37259 (US); Rio 

Reventaz6n, Finca Montecristo, below Cairo, 18-19 

Feb 1926 (6), Standley et al. 48594 (F, US); Puerto 

Viejo, Apr 1895 (2), Tonduz 9520 (F, US); PUNTARE- 

NAS: Golfito, 27 Jul 1951 (8), P. H. Allen 6280 (F, GH, 

US); Burica Peninsula, Quebrada Palito, 6 Mar 1973 

(st), Croat 22635 (MO). SAN JOSt: Guayabo, 3 Feb 

1908 (st), Ferry s.n. (F); Rio Vargas, Tabarcia, Mora, 

22 Apr 1963 (2), Jimenez 657 (COL, F); San Isidro El 

General, 3 Mar 1966 (6), Molina et al. 18289 (F, US); 

El General, Mar 1939 (6), Skutch 4256 (A, K, MO, S, 

US), (2), Skutch 4262 (A, K, MO, S, US), (6), Skutch 

4267 (A, K, MO, S, US); Las Vueltas, Tucirrique, Feb 

1899 (8), Tonduz 13280 (BM, F, G, K, LE, MO, P, 

US). 

PANAMA. Without locality (Isthmus of Panama), 

1859-1860 (st) Hayes 414 (NY), Hayes 867 (NY), (2), 

Hayes 986 (NY), (2), Hayes s.n. (BM). BocAs DEL TORO: 

- (2), Mutis 646 

(US, type collection ofC. mutisii), (a), Mutis 4558 (US), 

- (2), Triana 867 (BM, K, P), - (6), Triana s.n. (COL). 

AMAZONAS: Nr. Puerto Narifio, 24 Jul 1965 (6), Lozano 

640 (COL), 28 Jan-7 Feb 1969 (6), Plowman et al. 

2362 (F, GH, K, NY, S), 13 Sep 1963 (2), Soejarto 856 

(A). ANTIOQUIA: Cocorna, Jun 1937 (st), Bro. Daniel 

1793 (US); Jardin, 1800 m, Dec 1941 (2), Bro. Daniel 

2652 (COL, F, US, type collection of C. danielis); Rio 

Porce, 1100 m, 19 Sep 1963 (2), Espinal 1314 (COL); 

Rio Cauca, Puerto Valdiva, 17-20 Feb 1942 (2), Metcalf& 

Cuatrecasas 30093 (A, MO, US, type collection 

of C. macarenensis var. antioquensis). BOLiVAR: 150 

km N of Barrancabermeja, Mico-Ahumado camp, ca. 

8?15N, 74?4'W, 25 Aug 1966 (6), De Bruijn 1123 (K, 

M, MO, NY, S, U, VEN). BOYACA: El Humbo, 130 

km N of Bogotf, ca. 1000 m, 26 Apr 1933 (6), Lawrance 

769 (A, B, F, FI, G, K, MO, S, US, type collection of 

C. lawrancei). CALDAS: Salamina, Aug 1943 (?), Bro. 

Tomas 1923 (US). CAQUETA: Rio Caquetf, Rio Orteguaza, 

nr. Potosi, 7 Mar 1945 (2), Little et al. 9641 

(NY, US), 15 Mar 1945 (8), Little et al. 9789 (NY, US). 

CHOC6: Between Alto Curiche and Camp Curiche, E 

od Boca Curiche, 20 May 1967 (2), Duke et al. 11273 

(US); road Medellin-Quibd6, 6 km W of Siete, 38 km 

W of Bolivar, 1400 m, 6 Jan 1979 (2), Gentry et al. 

23713 (MO, U). CUNDINAMARCA: 5 km W of El Salto 

de Tequendama, on road to El Colegio, 1800 m, 14 

Jul 1972 (8), Barclay et al. 3593 (NA); between El Salto 

and El Colegio, 1480 m, 10 Mar 1940 (st), Cuatrecasas 

8291 (F, US); nr. Albfn, Aug 1962 (2), Ferndndez-


Perez F3 (COL); 3 km N of Tena, nr. La Laguna de 

Pedro Palo, 2080 m, 9 May 1952 (8), Ferndndez-Perez 

et al. 1458 (US); Mun. La Mesa, road to Mosquera, 

1800 m, 5 Aug 1963 (8), Soejarto 327 (A, K); Rio 

Bogota, 1600 m, 1909 (a), Uribe-Uribe 364 (COL). 

META: Rio Negro, 20 km W of Villaviicencio, 24 Mar 

1972 (Q), Barclay et al. 3294 (NA, US); nr. Villavicen- 

cio, 1-15 Aug 1946 (8), Duque-Jaramillo 3957 (COL), 

- (a), Karsten s.n. (LE); Sierra de la Macarena, Calno 

Entrada, 24 Dec 1949 (6), Phillipson et al. 1914 (BM, 

COL, US, type collection of C. macarenensis), 13 Jan 

1950 (2), Phillipson et al. 2095 (BM); nr. Villavicencio, 

Llano de San Martin, Feb 1856 (9), Triana 1866 (COL, 

F, K, NY). VALLE: Rio Sanquinini, La Laguna, 1250- 

1400 m, 10-20 Dec 1943 (st), Cuatrecasas 15589 (F); 

Gibraltar, N of Las Brisas, 2100-2200 m, 25 Oct 1946 

(2), Cuatrecasas 22526 (F). VAUPES: Rio Guayabero, 

8 Nov 1939 (a), Cuatrecasas 7510 (US, type of C. 

planitensis). 

VENEZUELA. Without locality, 26 Jun 1956 (2), 

Bernardi s.n. (8), 4 Mar 1957 (6), Bernardi s.n. (NY), 

1865 (a), Ernst 324 (BM), 1930-1934 (a), Vogl 1379 

(BR). AMAZONAS: Raudal de los Guaharibos, 24 Jul 

103 

et al. 95170 (NY, U, US, VEN); nr. Canoabo, 5 Mar 

1964 (8), Trujillo 6141 (MY). Departamento Federal: 

Nr. Caracas, 1940 (6), Bro. Elias 130 (F), Jan 1931 (6), 

Bro. Elias 631, 1 Oct 1972 (6), Hoyos 2043 (VEN), 17 

Nov 1975 (2), Oliva s.n. (MY, VEN), 24 Nov 1975 (9), 

Oliva s.n. (VEN). FALCON: Distr. Acosta, Mun. Jacura, 

Cerra de la Mina, 13 Feb 1961 (st), Ruiz-Terdn 423 

(MER). FALCON-YACACUY: Res. For. Rio Tocuyu, 

Quebrada El Charalito, Aug 1970 (9), Blanco 958 (NY, 

VEN). MERIDA: Merida(?), 26 Jun 1956 (9), Bernardi 

s.n. (MY). MIRANDA: Cerros del Bachiller, between 

Quebrada Corozal and Quebrada Santa Cruz, S of Santa 

Cruz, 18-19 Mar 1978 (st), Steyermarket al. 116463 

(MO, U, VEN). YARACUY: Bosques de Yumare, 10 Feb 

1959 (8), Bernardi 7017 (FI, G, MO); 35 km N of San 

Felipe, 19 Feb 1954 (6), Little 16245 (VEN); El Guaremal, 

between Valencia and San Felipe, Feb 1919 (8), 

Pittier 8404 (GH, US, VEN); Rio Taria, La Llanada 

de Taria, 5 Mar 1973 (st), Romero 548 (MY). ZULIA: 

Quebrada Perayra, affluent of Rio Tokuku, SW of 

Machiques, 29 Aug 1967 (St), Steyermark 99829 (VEN). 

ECUADOR. AZUAY: Chacayuca, western Andes of 

Cuenca, 

1951 (st), Croizat 312 (US); Salto Salas, 20 Aug 1951 

(2), Croizat 541 (US). ARAGUA: Cerro de Avila, 1 May 

1859 (8), Crueger s.n. (K); Colonia Tovar, May 1854 

(2), Fendler 1237 (GH, K), Jul 1854 (a), Fendler 1237a 

(GH, MO); nr. Maracay, 30 Jan 1857 (6), Fendler 1237 

(G), 1860 (9), Fendler 1237a (G), Rancho Grande, 6 

Jun 1963 (st), Montaldo 3435 (MY); Rio Chuao, El 

Medio, 15 Mar 1926 (8), Pittier 12124 (US, VEN); nr. 

Maracay, 2 Apr 1926 (9), Pittier 12142 (US, VEN), 2 

Apr 1926 (2), Pittier 12144 (A, G, M, MO, NY, VEN); 

road Maracay-Choroni, nr. El Castafio, 18 Feb 1937 

(a), Pittier 13985 (F, K, NY, US, VEN); Cerros de 

Guamitas, 1100 m, 27 Feb 1948 (a), Pittier et al. 15733 

(US, VEN); Parque Nacional Henri Pittier, 31 Oct 1975 

(2), H. Rodriguez 399 (MY); nr. El Castaino, 23 Mar 

1953 (2), Schnee 1337 (MY); road Maracay-Choroni, 

20 Dec 1948 (6), Standen 20 (MY); nr. El Castaino, 21 

Feb 1968 (2), Trujillo 8765 (MY); Guamitas, 12 May 

1938 (a), Ll. Williams 10071 (B, F, VEN), between 

Guamitas and El Lim6n, 28 Jan 1940 (a), Ll. Williams 

12348 (F, S, US, VEN); nr. El Castafio, 18 Feb 1937 

(st), Ll. Williams 13895 (US). BARINAS: Road Barinitas-Barinas, 

Mar 1953 (9), Aristeguieta 1699 (MY, NY, 

US, VEN); nr. Barinitas, Quebrada Parangula, Mar 

1953 (9), Aristeguieta 1704 (MY, NY, VEN); confluence 

of Rio Curbacitico and Rio Madre del Monte, 10 

Dec 1954 (a), Bernardi 1724 (F, FI, K, VEN); Alto del 

Aguada, Pedreza, 19 Feb 1955 (9), Bernardi 1980 (F, 

NY); Barinitas, 26 Jun 1956 (st), Bernardi 3354 (NY); 

between Altamira and Filo de Mertiuez, nr. Calderas, 

4 Mar 1957 (6), Bernardi 6348 (G); Res. For. Ticoporo, 

Rio Bumbum, 3 May 1964 (8), Breteler 3889 (G, MO, 

NY, P, S, U, US, VEN); Rio Caparo, 2-5 km above 

dam, 12 Mar 1980 (9), Liesner et al. 9443 (MO, U); 

Cuidad Bolivia, 12 Jan 1968 (a), R. F. Smith 3313 

(VEN); Res. For. Caparo, E of El Canton, 10 Apr 1968 

(2), Steyermark et al. 102083 (U, VEN), 12 Apr 1968 

(2), Steyermark et al. 102211 (F, MY, VEN). CARABOBO: 

Bejuma, 18 Feb 1954 (8), Little 16230 (VEN); Rio San 

Gian, S of Borburata, 27-28 Mar 1966 (9), Steyermark 

- (9), Lehmann 5606 (F, K, type collection 

of C. lehmannii). ESMERALDAS: Borb6n, 11 Sep 1965 

(8), Little et al. 21047 (NY, Q, US); 10 km SE of Esmeraldas, 

29 Sep 1965 (9), Little et al. 21172 (NY, Q, 

US). GUAYAS: Nr. Naranjito, 6-7 Jul 1945 (st), Camp 

E3626 (NY, U); Baloa, - (9), Eggers 14165 (A, L, LE, 

M, US). IMBABURA: Nr. Lita, 7 Feb 1981 (8), Berg 

1241 (U). Los Rios: Pichilingue, 18 May 1943 (2), 

Little 6461 (F, US); Rio Palenque Biol. Station, km 56 

on road Quevedo-Santo Domingo de los Colorados, 

11 Feb 1973 (6), Dodson 5241 (MO, QCA, SEL, U), 

27 Feb 1975 (6), Dodson 5778 (QCA, US), 29 Jul 1972 

(6), Dwyer 10305 (MO), 15 Feb 1974 (9), Gentry 9934 

(MO, QCA, U), 22 Feb 1974 (6), Gentry 10127 (MO, 

QCA, U); canton Vinces, hacienda Santa Lucia, 18- 

28 Oct (8), Mexta 6573 (F, NA). MANABI: El Recreo 

Dec 1891 (8), Eggers 14165 (A, BR, L, M, US, type 

collection of C. eggersii), 1897 (9), Eggers 15615 (F, 

LE, P). MORONA-SANTIAGO: Macas, 18 Mar 1956 (2), 

Asplund 19835 (S); 5 km SW of Macas, 26 Jan 1981 

(9), Berg 1222 (U); Cord. de Cucutfi, 5-10 km E of 

Logronio, 1200-1500 m, 7-9 Oct 1975 (6), Little et al. 

602 (COL, Q). NAPO: Rio Cuyabeno, ca. 0?10'S, 

76?55'W, 16 Feb 1980 (2), Berg & Akkermans 1033 

and 1040 (U), 19 Feb 1980 (6), Berg & Akkermans 

1062 (U); Misahualli, mouth of Rio Misahualli, 3 Mar 

1980 (9), Berg & Akkermans 1113 (U); 1 km S of Lago 

Agrio, 5 Nov 1974 (6), Gentry 12474 (MO, U); Rio 

Putumayo, just above mouth of Rio Gueppi, 19 May 

1978 (9), Gentry 22095 (MO, U). PICHINCHA: Road 

Santo Domingo de los Colorados- Quinind6 (km 170- 

175), 1 Sep 1949 (st), Acosta Solis 13629 (F); Santo 

Domingo de los Colorados, 20 Aug 1930 (st), Benoist 

3044 (P, S, U, US); nr. Alluriquin, bridge over Rio 

Toachi, 23 Feb 1981 (6), Berg 1301 (U); road Santo 

Domingo de los Colorados-Chona, km 5, 3 Apr 1943 

(8), Little 6155 (K, Q); Santo Domingo dos Colorados, 

21 Apr 1943 (9), Little 6328 (F, K, NY, Q, US); nr. 

Alluriquin, at confluence of Rio Alluriquin and Rio 

Toachi, 14 Mar 1967 (8), Sparre 14822 (S). ZAMORA- 

CHINCHIPE: Quebrada Las Pavas Pavas, 14 km from


Loja, 1800 m, 21 Aug 1975 (st), Samaniego et al. 92 

(COL, Q). 

PERU. Without locality, - (9), Dombey s.n. (F, P, 

US), 1835 (6), Matthews 2068 (K, OXF, U), 1829 (6), 

Poeppig s.n. (B, OXF, type collection). AMAZONAS: Rio 

Cenepa, Aintami Creek, 24 Nov 1972 (6), Berlin 372 

(MO, U); Pongo de Manseriche, 1924 (9), Tessmann 

4696 (B, NY); Rio Santiago, Caterpiza, 20 Nov 1979 

(8), Tunqui 103 (U). Huanuco: Pampayacu, nr. Huanaco, 

20 Jan 1927 (6), Kanehira 99 (GH); Prov. Leoncio 

Prado, Tingo Maria, 7 Dec 1981 (6), Plowman et al. 

11180 (U); Macora, - (9), Ruiz & Pavon s.n. (or 3) (B, 

FI, type collection of C. vellera), - (5), Ruiz & Pavon 

s.n. (BR, G, OXF, US); Huamalies, between Monzon 

and Rio Huallaga, - (9), Weberbauer 3702 (G, type 

collection of C. standleyi). JUNIN: La Merced, Puente 

Quimiri, 24 Oct 1950 (6), Nunez 2928 (US); Sanabeni, 

4 Sep 1960 (8), Woytkowski 5956 (MO, U). LORETO: 

Prov. Maynas, Rio Itaya, above Puerto Belen, 16 Nov 

1978 (9), Diaz et al. 618 (U); Prov. Maynas, Rio Itaya, 

nr. Palo Seco, ca. 40 km above Iquitos, 20 Mar 1977 

(6), Gentry et al. 18458 (MO, U); Prov. Requena, Rio 

Tapiche, tributary of Rio Ucayali, 1 hour above Requena, 

8 Dec 1972 (,), Gentry et al. 21291 (MO, U); 

lower Rio Huallaga, Yan6n, 5-11 Sep 1929 (9), Killip 

et al. 29249 (NY, US, type collection of C. grandiceps); 

Rio Marafion, lower Rio Ampiyacu, 6 Mar 1977 (6), 

Prance et al. 24687 (U); Prov. Maynas, Rio Nanay, 

Mishana, 1978 (6), Ramirez 1084 (U); Rio Ucuyali, 

Yarinacocha, between 10?S and mouth, 1923 (6), Tessmann 

3461 (B, G, NY, S); lower Rio Huallaga, Yurimaguas, 

26 Oct 1929 (6), LI. Williams 4179 (F); Puerto 

Arturo, Yurimaguas, 22 Nov 1929 (2), LI. Williams 

5349 (B, F). (LORETO?): Rio Zungarococha, 28 Jan 

1978 (6), Ayala et al. 1436 (MO, U). MADRE DE Dios: 

Prov. Tambopata, affluence of Rio Tambopata and Rio 

Le Torre, Tambopata Nat. Res., ca. 30 km SSW of 

Puerto Maldonado, 16 May 1980 (9), Balbour 5372 

(U); Parque Nacional del Manu, Rio Manu, Pakitza 

station, 9 Nov 1980 (6), Foster 5721 (F). 

BRAZIL. Without locality, - (9), Martius s.n. (M, 

type collection of C. martiana), - (6), Martius s.n. (BR, 

M, U, type collection of C. subincana). ACRE: Rio Acre, 

Itui, 4 Nov 1923 (9), Kuhlmann 759 (RB, U); Tarauaca, 

1-2 km E of Rio Tarauaca, 24 Sep 1968 (2), Prance et 

al. 7530 (F, GH, K, NY, P, R, S, U, US); Rio Jurui- 

Mirim, Aug 1901 (6), Ule 5717 (B, G, K, L, MG). 

AMAZONAS: Rio Solimoes, Mun. Tefe, Paleta, 21 May 

1933 (6), Krukoff4518 (A, BM, F, G, K, M, MO, NY, 

S, U, US, type collection of C. subcrenata); nr. mouth 

of Rio Embira, tributary of Rio Tarauaca, 26 Jun 1933 

(6), Krukoff4994 (A, G, K, NY, U, US, type collection 

of C. embirana); Mun. Humaita, Tres Casas, 14 Sep- 

11 Oct 1934 (9), Krukoff 6165 (A, F, FI, K, MO, NY, 

S, U, US), Krukoff 6524 (A, BM, F, G, K, LE, MO, 

NY, S, U, US, type collection of C. araneosa); Rio 

Madeira, Boa Hora, 4 Sep 1923 (2), Kuhlmann 368 

(RB); Rio Japura, - K, NY, P, R, S, U, US); Rio Purus, between Lago Quati 

and Lago Arima, 20 Jun 1971 (6), Prance et al. 13438 

(K, M, MO, NY, S, U, US); Manaquiri, Jun 1851 (8), 

Spruce 1608 (BM, BR, C, G, GH, K, LE, NY, P, OXF). 

MATO GROSSO: Rio Aripuana, nr. Humboldt Center, 

ca. 10?12'S, 59021'W, 12 Oct 1973 (9), Berget al. P18455 

(F, K, MO, NY, S, U, US), 22 Oct 1973 (6), Berg et 

al. P19841 (F, K, MO, NY, P, S, U, US); Sarar6, 14? 

50'S, 59'45'10"W, 4 Aug 1972 (6), Pires et al. 16461 

(U). PARA: Alto Tapaj6s, Rio Curucii 1-5 km above 

Missao Cururui, 7?35'S, 57?31'W, 17 Feb 1974 (9), Anderson 

11033 (MO, U); Gleba Bacaja, below mouth of 

Rio Bacaja, 22 Nov 1980 (2), Prance et al. 26402 (U). 

RORAIMA: Rio Mucajai, Posto 17 Mar 1971 (6), Prance 

et al. 11066 (K, M, NY, P, S, U). 

BOLIVIA. SANTA CRUZ: Prov. Ichilo, Rio Vibora, 

5 Oct 1926 (2), Steinbach 7567 (A, F, GH, K, MO, S, 

U, type collection of C. boliviana). 

Coussapoa villosa is a widespread species (like 

C. asperifolia) and is extremely variable in many 

characters. Although the occurrence of some of 

the features is associated with certain parts of the 

species area, e.g., branched pistillate inflorescences 

in Central America and brown arachnoid 

hairs in Colombia and adjacent parts of Ecuador, 

infraspecific taxa cannot readily be recognized. 

Some specimens from the periphery of the species 

area and/or with features near the extreme of the 

variation must be mentioned. Barclay et al. 3593 

(Colombia) has staminate inflorescences with extremely 

numerous (ca. 50) heads; Gentry et al. 

23713 (Colombia) has pistillate inflorescences 

with very short (less than 2 cm long) peduncles 

(in this respect it forms a transition to C. duquei); 

Prance et al. 26402 (Brazil, Pari) is peculiar in 

that the head of the pistillate inflorescence is lobed 

and obliquely attached to the peduncle. This last 

collection matches some more or less aberrant 

collection of C. tessmannii (see p. 95) in the dense 

covering of dark brown moniliform pluricellular 

(=granular) hairs on the peduncle, leafy twig, and 

petiole. 

Typification of C. villosa: It is unknown 

whether the destroyed type material in W consisted 

of a pistillate and a staminate specimen as 

the drawing in Poeppig & Endlicher (1838) suggests. 

The specimens in B and OXF which could 

replace the holotype material are staminate. 

(6), Martius s.n. (G); Lago de Ma- Typification of C. eggersii: Eggers made sevnacapuru, 

10 Oct 1972 (2), 0. Pires 248 (U); Tefe, 23 eral collections under number 

Jul 1972 

14165- 

(9), PLK& Urbano 12296 

staminate 

(INPA); road Boca 

do Acre-Rio Branco, km 1-4, 19 Sep 1966 (6), Prance and pistillate ones from different localities in Ecet 

al. 2306 (F, GH, K, P, R, S, U); Rio Demini, To- uador (El Recreo and Baloa)-at several dates. 

totobi, 28 Feb 1969 (2), Prance et al. 10348 (F, GH, These collections have been mixed up. The spec-


imen in F on which C. eggersii is based is staminate 

and possibly from El Recreo. In the original 

description the specimen is wrongly described 

as pistillate. 

43. Coussapoa viridifolia Cuatrecasas, Fieldiana 

Bot. 28(1): 213. 1956. Type. Venezuela. Bolivar: 

0-8 km N of Santa Teresita de Kavanayen, 

23 Nov 1944 (6), Steyermark 60456 (holotype, 

F; isotype, VEN). Fig. 55. 

Coussapoa viridifolia Cuatrecasas var. tenuifolia Cuatrecasas, 

Fieldiana Bot. 28(1): 214. 1956. Type. Venezuela. 

Bolivar: Cerro Duida, Canio Negro, 25-26 

Aug 1944 (9), Steyermark 57955 (holotype, F; isotype, 

VEN). 

Tree or shrub, often hemi-epiphytic, up to 20 

m tall. Leafy twigs 2-5 mm thick, appressedpuberulous 

to substrigose to hirtellous, with hairs 

differing distinctly in length. Lamina coriaceous 

to subcoriaceous, obovate to elliptic to oblong, 

2.5-10 x 1-6 cm, apex obtuse to shortly acuminate 

or rounded to emarginate, base obtuse to 

acute to cuneate to rounded, margin entire to 

subcrenate; upper surface glabrous, lower surface 

sparsely puberulous to substrigose on the main 

veins, the hairs differing distinctly in length; lateral 

veins 4-7 pairs, basal pair unbranched or 

faintly branched reaching the margin at or above 

the middle of the lamina, sometimes some of the 

other lateral veins poorly branched (furcate); intercostal 

venation plane to more or less prominent; 

petiole 1-3 cm long, sparsely (to rather 

densely) appressed-puberulous to substrigose, 

with hairs differing distinctly in length; stipules 

0.5-1 cm long, densely appressed-puberulous to 

subsericeous; terminal buds slender. Staminate 

inflorescences branched; heads 3-14, globose, ca. 

2-4 mm diam.; common peduncle 1-3 cm long, 

sparsely puberulous; perianth ca. 1 mm high, 

glabrous; stamens two, just exceeding the perianth. 

Pistillate inflorescences unbranched or 

poorly branched; heads 1-3, globose, ca. 3-6 mm, 

in fruit up to 8 mm diam.; (common) peduncle 

0.5-2.5 cm long, sparsely (to rather densely) puberulous; 


bracts often only a few (sometimes lacking?), 

small, subspathulate to subpeltate, minutely 


Distribution (Fig. 9). Eastern Guiana (=Highland) 

region, in Venezuela (Bolivar and Ama- 

105 

zonas), Guyana (Upper Mazaruni), and Brazil 

(Roraima); in rain forest and savannas, up to 

1250 m. 

Specimens studied. VENEZUELA. AMAZONAS: 

"Alto Orinoco," 1951 (2), Croizat 959 (F, US); Cerro 

Duida, Canio Negro, 25-26 Aug 1944 (2), Steyermark 

57955 (F, VEN, type collection of C. viridifolia var. 

tenuifolia). BOLIVAR: Region of Rio Icabaru and Rio 

Hacha, 2 Jan 1956 (2), Bernardi 2744 (F, FI, NY, VEN); 

between Santa Teresita de Kavanayen and ravine ca. 

8 km NW of it, 23 Nov 1944 (8), Steyermark 60451 

(F, VEN, type collection). 

GUYANA. Kamarang River, below mouth of Utschi 

Creek, 30 Oct 1960 (8), Tillett et al. 45869 (F, NY, U). 

BRAZIL. AMAZONAS: Serra de Neblina, Rio Cauaburi-Rio 

Maturaca, 600-1300 m, 16 Dec 1965 (2), 

N. T. Silva et al. 60670 (U). RORAIMA: Serra Surucucu, 

2 Feb 1975 (9), Ribeiro 655 (IAN), 1 Feb 1975 (2), Rosa 

339 (IAN). 

This species appears to be closely related to C. 

latifolia. The two are allopatric and apparently 

ecologically distinct, but morphologically very 

close. The two taxa show (minor) differences in 

the leaf venation, the length of the stipules, and 

the indumentum of the leaves. The morphological 

differences suggest the possibility of distinction 

at the subspecific level. 

The following three species have been discov- 

ered and described after submission of the post- 

review manuscript of the revision and are, there- 

fore, not placed in the alphabetical sequence. 

44. Coussapoa fulvescens C. C. Berg, Brittonia 

42: 59-65. 1990. Type. Colombia. Choc6: Rd. 

Quibdo-Bolivar, ca. km 50, 9 Jul 1986 (2), 

Berg 1546 (holotype, COL; isotypes, AAU, 

BG, F, HUA, MO, NY). Fig. 56. 

Tall, hemi-epiphytic tree. Leafy twigs ca. 10 

mm thick, white-appressed-puberulous, partly 

also brownish-hirsute, lenticels conspicuous. 

Lamina coriaceous, oblong to subobovate, 18- 

26 x 9-15 cm, apex short-acuminate to obtuse, 

base obtuse, margin entire; upper surface glabrous, 

lower surface sparsely appressed-puberulous; 

lateral veins 7-9 pairs, basal pair unbranched 

(or very faintly branched) reaching the 

margin at or just below the middle of the lamina, 

usually the second and/or third lateral veins (from 

the base) with one or two strong branches, tertiary 

venation distinct, slightly prominent; petiole 

5-10 cm long, appressed-puberulous; stipules 1.5-


FXG. 56. Coussapoafillvescens: 1, leafy twig with Distillate in~lorescences (Berg 1546), x0.75. 

r;:~~~~~~~~~~~~~~~~~~' 

":~~~~~~~~~~~~~~~.:? 

1~~?. ~ . 

?i I?:i, 

?~~~~~~~~~~~~~~~~~~~~~~' 

"' 

FIG. 56. Coussaoafulvescens: 1 leafy twig wih pistillate inflorescence (Beg 1546), x0.75


2 cm long, yellowish- to brown-(sub)hirsute. Pistillate 

inflorescences branched; heads 10-20, 

sometimes partly fused, (sub)globose, 3-5 cm in 

diam.; common peduncle 3-6 cm long, ca. 2 mm 

thick, appressed-puberulous and partly also 

brownish-hirtellous; perianth ca. 2 mm high, glabrous, 

+ distinctly ribbed. Interfloral bracts subulate 

to narrowly spathulate, with a few hairs at 

the apex. 

Distribution (Fig. 9). Colombia (Choc6), only 

known from the type locality, in lowland secondary 

growth. 

Tree, 18 m tall. Leafy twigs 5-12 mm thick, 

yellowish- to whitish-hirtellous. Lamina coriaceous, 

ovate to elliptic, 10-21 x 8-16 cm, apex 

shortly acuminate (to acute), base rounded to 

subcordate, margin coarsely crenate to subentire; 

upper surface with sparse hairs on the midrib, 

lower surface hirtellous on the veins, in the reticulum 

minutely puberulous; lateral veins 12- 

14 pairs, basal pair branched, reaching the margin 

below the middle of the lamina, tertiary and 

quarterary venation and reticulum prominent; 

petiole 4-6 cm long, 2-4 mm thick, hirtellous, 

hairs not distinctly different in length; stipules 

4-12 cm long, densely yellowish-puberulous and 

partly also hirsute. Pistillate inflorescences 

branched; heads 4-5, globose (to subdiscoid), in 

fruit 0.7-1.3(-1.5) mm in diam.; common peduncle 

2.5-3 cm long, ca 2-2.5 mm thick, hirtellous; 

perianth ca. 1.5 mm high, at the apex 

puberulous, in fruit red. Interfloral bracts absent. 

Distribution (Fig. 9). Colombia (Tolima), only 

known from the type locality. 

Specimens studied. COLOMBIA. TOLIMA: 3.4 km 

from Las Juntas, 1940 m, 9 Apr 1984 (v), Escobar et 

al. 4219 (HUA, type collection). 

This species resembles C. nitida, known from 

the Amazon Basin and mostly occurring in riv- 

erside vegetation. It differs clearly from the latter 

in the prominent tertiary and quartemary ve- 

nation at the lower surface of the lamina and in 

the red-colored fruiting perianth. 

46. Coussapoa valaria C. C. Berg, Brittonia 42: 

59-65.1990. Type. Colombia. Valle: Bajo Cali- 

ma, rd. to Dindo, 25 Sep 1986 (2), Monsalve 

B. 1176 (holotype, CUVC; isotypes, BG, 

MO). Fig. 58. 

Specimens studied. COLOMBIA. CHOC6: Rd. 

Quibdo-Bolivar, ca. km 50, 9 Jul 1986 (9), Berg 1546 

(AAU, BG, COL, F, HUA, MO, NY, type collection). 

This species is closely related to C. herthae, Tree, hemi-epiphytic. Leafy twigs 5-10 mm 

known from western Ecuador and Narifio (Co- thick, brown strigose, at least at the scars of the 

lombia). It differs from the latter, e.g., in the stipules, glabrescent. Lamina coriaceous, ellipindument 

of the leaves, the venation, the shorter tic, 15-17 x 11-14 cm, apex rounded to shortly 

stipules, the more numerous and smaller flower acuminate, base rounded to subcordate, margin 

heads of the pistillate inflorescence. The two taxa subentire, at the base revolute; upper surface glamight 

prove to be distinct only at the subspecific brous, lower surface strigillose at the base of the 

level. C. fulvescens may also be related to C. midrib, on the midrib and lateral veins with a 

echinata. 

soon disappearing white arachnoid indument; 

lateral veins 8-10 pairs, basal pair + branched, 

45. Coussapoa tolimensis C. C. Berg, Brittonia reaching the margin at or just below the middle 

42: 59-65. 1990. Type. Colombia. Tolima: 3.4 of the lamina, tertiary venation almost plane; 

km from Las Juntas, 1940 m, 9 Apr 1984 (9), petiole 5-6.5 cm long, glabrous or brownish stri- 

Escobar et al. 4219 (holotype, HUA). gose, stipules 2-3 cm long, rather densely whit- 

Fig. 57. ish appressed-puberulous to -strigillose, or also 

with white arachnoid hairs. Pistillate inflorescences 

branched; heads 3-9, 3-4 (in fruit 7-9) 

mm diam.; common peduncle 2-4 cm long, 

densely minutely puberulous; perianth ca. 1.5 

mm high, minutely puberulous around the aperture. 


Distribution (Fig. 9). Colombia (Valle), in lowland 

forest. 

Specimens studied. COLOMBIA. VALLE: Bajo Calima, 

rd. to Juanchaco Palmeras, 10 Jul 1984 (9), Gentry 

et al. 47838 (BG); Bajo Calima, rd. to Dindo, 25 Sep 

1986 (9), Monsalve B. 1176 (BG, MO, type collection). 

This species resembles both C. ovalifolia and 

C. parviceps. It differs from these species in, e.g., 

the venation and the number and dimensions of 

the flower heads of the pistillate inflorescences. 

Unnamed Collections 

Gentry & Mori 14011 from Panama: Discussed 

under C. ovalifolia. 

Huashikat 1585 from Peru: Discussed under C. 

cupularis.


FIG. 57. Coussapoa tolimensis: 1, leafy twig with pistillate inflorescences 

(Albert de Escobar et al. 4219), x0.6. 

Names Not Included 

Coussapoa dealbata Andre, Ill. Hort. 17: 17. 

1870--nomen nudum. 

Coussapoa dolichandra Cuatrecasas Caldasia 7: 

287. 1956-based on collection Cuatrecasas 

17458 (Colombia, Valle: Rio Cajambre, 5-15 

May 1944 (holotype and isotype, F)-consists 

109 

of discordant elements: staminate inflores- 

cences of a Coussapoa species and leafy twigs 

of some species not belonging to the Urticales, 

but possibly to the Lauraceae. The latter parts 

constitute the lectotype, chosen here. 

Coussapoa emarginata Killip in Macbride, Publ. 

Field Mus. Bot. 13(2.2): 296. 1937 = Pourou- 

ma minor Benoist.


FIG. 58. Coussapoa valaria: 1, leafy twig with pistillate inflorescences 

(Gentry et al 47838), x0.55. 

Coussapoa krukovii Standley, Publ. Field Mus. 

Bot. 17:1937 = Pourouma tomentosa Miquel. 

Coussapoa laevigata Poeppig & Endlicher, Nov. 

gen. sp. pl. 2: 33. 1838-based on material 

from Brazil (Amazonas: Tefe). The holotype 

in W has been destroyed. Isotypes have not 

been traced. The identity is uncertain. The 

name possibly applies to C. villosa. 

Coussapoa leopoldii Micheli, Rev. Hort. (1894): 

251. 1894-nomen nudum. 

Coussapoa obovata Warburg ex Glaziou, Bull. 

Soc. Bot. France 59 (Mem. 3g): 645. 1913nomen 

nudum. 

Coussapoa pittieri Standley in Hoyos, Los Ar- 

= Ficus intermarginalis (Liebmann) Miquel, 

Ann Bot. Lugd. Bat. 3:297. 1867; see Standley, 

Publ. Field Mus. Bot. 18(1): 383. 1937. 

POUROUMA 

Introduction 

Pourouma is the smallest of the three Neotropical 

genera of the Cecropiaceae. It is a welldefined 

genus of small to medium-sized trees of 

the rain forest areas of South and Central America, 

and has not drawn much attention, except 

that the fruits of P. cecropiifolia are suitable for 

boles de Caracas, ed. 2 (Monogr. 24. Soc. Cienc. human consumption. The genus is rather uni- 

Nat. La Salle, Caracas, Venezuela) p. 242, tab. form in its ecology. Besides a number of clear- 

(of C. villosa)-nomen nudum. 

cut species, mostly of limited distribution, it 

Coussapoa plicata Warburg ex Ule, Bot. Jahrb. comprises several more wide-spread and rather 

40: 143. 1907- nomen nudum. 

complex groups of taxa with a confusing varia- 

Coussapoa rekoi Standley, Contr. U.S. Natl. Herb tion in leaf shapes and indument, and to which 

20: 211. 1919 = Poulsenia armata (Miquel) the following quotation applies: "The species of 

Standley: see Mildbraed, Notizbl. Bot. Gart. Pourouma are perhaps the most ambiguous and 

Berlin 10: 413. 1928; Standley, Trop. Woods vexatious of the American Moraceae, a dubious 

33: 4. 1933. 

distinction" (Woodson & Schery, 1960:166). The 

Urostigma intermarginale Liebmann, Kongel. lack of sufficient field work must be deemed a 

Danske Vidensk.-Selsk. Skr. Ser. 5,2:328.1851 great disadvantage in the preparation of the pres-

Morphology 

ent revision, especially with regard to the vari- 

ation in the leaves. 

111 

They are often very dense and form a powdery 

layer in dried material. The denseness of these 

hairs is often variable. 

Taxonomic History 

2. Unicellular arachnoid (cobwebby) hairs. These 

very thin, crinkled, interwoven hairs of dif- 

The genus Pourouma was established by Au- ferent length are mostly white, sometimes 

blet (1775) who described one species, P. gui- brownish (in P. ferruginea). Short arachnoid 

anensis. Names were added by Martius (in Spix hairs are always found in the areoles at the 

and Martius, 1831, 1843), Tr6cul (1847), Klotzsch lower leaf surface. Longer arachnoid hairs can 

(1847), and Miquel (1853), and in this century be found on the smaller veins, and in some 

by Rusby (1910), Benoist (1922, 1924), Mild- species (e.g., P. ferruginea and P. tomentosa) 

braed (1927, 1928), Macbride (1930), Ducke also on the main veins, from which they often 

(1932a, 1932b, 1947), Standley (1937a, 1937b, disappear with age. Long arachnoid hairs also 

1939), Cuatrecasas and Standley (in Cuatrecasas, occur on the petioles, stipules, leafy twigs and/ 

1951, 1956b), Cuatrecasas (1954, 1956a, 1956b, or the inflorescences in some species. 

1956c, 1967), Woodson and Schery (1960), and 3. Unicellular non-arachnoid, thicker and mostfinally 

by Berg and Kooy (1982), Berg and van ly more or less straight hairs. Short hairs are 

Heusden (1988), and Berg (1989, 1990). 

usually whitish, the longer ones mostly yel- 

Since the comprehensive treatments of Pou- low(ish). They can be found on most parts of 

rouma by Trecul (1847) and Miquel (1853) the the plant. In some species, forms with only 

genus has only been treated for regional floras: short, whitish hairs occur beside forms pos- 

Peru (Macbride, 1937), Panama (Woodson & sessing longer yellowish hairs. In several 

Schery, 1960), Costa Rica (Burger, 1977), and species, all or some specimens have short, 

Surinam (Berg, 1975). 

rigid hairs on the upper or sometimes on the 

lower leaf surface. 

Morphology 

Leaves. -The leaves are medium-sized to 

Habit.-Pourouma species are usually small 

large, 

entire or 

to medium-sized trees, often with stilt-roots from palmately incised. In taxa with incised 

leaves the first formed leaves are 

the lower part of the trunk, as in most of the 

entire; subseother 

terrestrial, non-climbing species of Cecro- quently leaves sooner or later, frequently or alpiaceae. 

In some species (e.g., P. ways, become incised. In adult specimens, eshirsutipetiolata) 

the trunk may become buttressed. The architecpecially 

on flowering branches, entire leaves can 

ture of the trees of P. minor is that of the model 

again be found, although different in shape and 

texture from the leaves in the 

of Aubreville (F. Halle, pers. comm.). The 

juvenile stage. Entire 

and incised leaves 

branches may be thick (as in species with incised 

may occur on the same 

branchlet. Leaves similar to those formed in the 

leaves) or rather thin (as in species with mediumsized, 

entire 

subjuvenile stage often appear again on sucker 

leaves). 

shoots of adult 

The wood (and other vegetative parts) of some 

specimens with entire leaves. 

Leaves are 

species (e.g., P. myrmecophila) give out a 

always incised in adult specimens of 

pecu- Pourouma 

liar odor, indicated as cecropiifolia, and probably also in P. 

'spearmint', 'winter-green', 

'balsam', or 'bengie'. For other species (e.g., P. 

myrmecophila, P. oraria, P. stipulacea, and P. 

villosa. Incised leaves 

quianensis) a 'balsam' odor is reported for the 

appear to occur constantly 

in some 

fruits. 

subspecies of species in which entire 

leaves are common. In 

When cut the plant parts exude a species in which 

watery sap 

only 

entire leaves are 

which is clear or sometimes reddish (e.g., in P. 

found, the leaves in the juvenile 

and 

bicolor); after exposure to the air it turns black. subjuvenile stages are basically similar to 

those of adult 

Indumentum. -Three types of trichomes can 

specimens. A number of species 

be 

appear to be constantly entire-leaved (e.g., P. borecognized: 

livarensis, P. minor, P. ovata, and P. phaeotri- 

1. Pluricellular hairs, often more or less globose, cha). In this group of species the leaves may vary 

pale brown to dark brown or purple. These from predominantly ovate to elliptic to oblong, 

hairs occur on most young parts of the plant. or even to obovate.


Leaf texture varies from chartaceous or subcoriaceous 

to coriaceous, and leafindument from 

flowers. A subumbellate pistillate inflorescence 

is characteristic for Pourouma minor. Staminate 

dense (with the three types of trichomes repre- inflorescences are often more or less dorsivensented) 

to sparse. 

trally flattened. 

The venation varies from pinnate in entire The flowers of the staminate inflorescences are 

leaves to subpalmate in incised leaves. In species more or less closely clustered at the end of the 

in which palmately incised leaves occur, the bas- branches. In several species the flowers form 

al lateral veins even of entire leaves are normally (sub)globose heads. 

branched, while in species in which incised leaves The inflorescences are mostly ebracteate, alare 

not formed, the basal lateral veins are often though minute bracts are found in staminate and 

unbranched or only faintly or sparsely branched. pistillate inflorescences of P. cucura, and some- 

The tertiary venation is scalariform. 

what larger (caducous) bracts at the base of the 

The leaf margin is entire or often more or less peduncle are sometimes found in other species. 

faintly crenate. In most species the petioles are Staminate Flowers. -Staminate flowers are 

relatively long, with a considerable variation in pedicellate or sessile and tetramerous, or somelength 

on the same twig. Relatively short peti- times trimerous. The tepals are free, basally conoles, 

only slightly varying in length, are found in nate, or almost entirely connate, and then they 

a few species (e.g., P. formicarum). In P. for- form an urceolate or infundibuliform perianth. 

micarum and P. myrmecophila the base of the The stamens are short in species with free or 

petiole is swollen and has an abaxial slit. These basally connate tepals, but in species with almost 

two species are myrmecophilous. 

entirely connate tepals the filaments are longer 

The mostly large stipules are connate and fully than the tepals. Probably in connection with the 

amplexicaul, and appear to be of importance for rather narrow opening of the urceolate perianth, 

the protection of young inflorescences-espe- the anthers are outside the perianth (long?) before 

cially staminate inflorescences with flowers with anthesis. The filaments are free, sometimes baa 

connate (more or less urceolate) perianth and sally connate, but in Pourouma melinonii subsp. 

the anthers already exserted before anthesis, as glabrata they can be entirely connate, thus largely 

in P. tomentosa. In some species the stipules are similar to the androecium characteristic for 

short, e.g., P. ovata. While in most species the Coussapoa. In the center of the flower one can 

stipules are caducous, they are (sub)persistent in usually find a tuft of hairs, which might represent 

P. myrmecophila, P. oraria, and P. stipulacea. a pistillode. 

The stipules leave a horizontal annular scar, not Pistillate Flowers. -Pistillate flowers are aloblique 

as in Coussapoa. This feature may help ways pedicellate. The perianth is tubular with an 

to distinguish sterile specimens of the two genera, entire or slightly lobed margin. The ovary is free 

of which the leaves can be very similar. from the perianth. The short style bears a stigma 

The leaf characters mentioned are more or less that is subpeltate or peltate to knob-like, in Poucorrelated. 

In taxa with incised leaves the leaves rouma minor. 

tend to be large, often subcoriaceous and densely Fruits and Seeds.- The fruit has a dry pericarp 

hairy, with relatively long petioles, often quite and is completely enclosed by the enlarged perivariable 

in length, and often with long stipules. anth. The inner layer of the fruiting perianth be- 

Contrariwise, in taxa with entire leaves, the leaves comes soft and whitish and the other layer (skin) 

tend to be medium-sized, usually coriaceous and ? leathery and at full maturity often purple to 

often rather sparsely hairy. They have relatively black or red-brown. The whole (fruit and perishort 

petioles, more or less constant in length, anth) can be indicated as a pseudodrupe (or funcand 

the stipules tend to be small. 

tional drupe). 

Inflorescences. -The inflorescences are nearly The pedicel under mature fruits may become 

always ramified, basically with three branches reddish (a contrasting color against black?). 

departing from the top of the peduncle. Further Whether this color is caused by a change of color 

ramification is usually (sub)dichotomous. The of tissues of the pedicel or by the presence of redprimary 

ramification can be dichotomous as well. brown pluricellular hairs is not clear. 

In pistillate inflorescences the ramification is often The fruiting perianth is reported to be aromore 

reduced than in the staminate ones, with matic for some species (e.g., Pourouma bicolor). 

a tendency to a subumbellate arrangement of the Pourouma and the African Myrianthus are

Morphology; Phenology; Pollination 113 

macrospermous, in contrast to the other four 

Phenology 

genera of the Cecropiaceae. 

General Remarks. -The greatest variation of From label data alone it is difficult if not imcharacters 

(mainly, but not only of the leaves) in possible to draw conclusions about the phenolthe 

genus is found among taxa in which incised ogy of a particular species. 

leaves do not occur constantly. Only part of this In an area explored (Montagne de Trinite, 

variation is connected with distribution. Taxa French Guiana, short rainy season, Jan-Feb 1984) 

with constantly incised leaves in adult specimens 

none of the numerous Pourouma specimens of 

and those with entire leaves in all stages of de- different species was bearing flowers or fruits. 

velopment tend to be rather uniform, even if This 

they suggested seasonal flowering. The combihave 

a relatively wide distribution. 

nation of label data of all Pourouma collections 

It is somewhat difficult to indicate which char- made in the Guianas gives a pattern indicating 

acters can be regarded as derived or as a main 

special- flowering period in August-October (in 

ized. In several of its characters (especially of the the long dry season) and a fruiting period which 

vegetative parts) Pourouma is (like the African may extend to February, thus from the end of 

genus Myrianthus) intermediate between Cecro- the long dry season into a short rainy season. 

pia (and the African genus Musanga) and Cous- According to Croat (1978) P. bicolor fruits in 

sapoa (and the Malesian Panama in the end of the 

genus Poikilospermum). 

dry season. According 

From about the middle of the spectrum of vari- to Martius (in Spix & Martius, 1831) P. cecroation 

in Pourouma one could indicate a pro- piifolia can bear (ripe?) fruits in May and Nogression 

towards Cecropia, ending in a form with vember. The occurrence of these two periods of 

thick branches and large, constantly incised leaves fruiting gets some support from the label data of 

with many incisions (but never peltate as in Ce- the examined material of this species. Falcao and 

cropia), represented, e.g., by P. cecropiifolia and Lleras (1980) found that P. cecropiifolia in the 

P. cuspidata; and another progression towards Manaus region was flowering at the height of the 

Coussapoa, ending in a form with rather slender rainy season (April-June), and producing a crop 

branches and medium-sized, constantly entire of ripe fruit at the end of the dry season to the 

leaves. As indicated (p. 5), there is an overlap beginning of the next rainy season. 

of the leaf characters of Pourouma and Cous- The period between flowering and bearing ripe 

sapoa, and sterile material of both genera can be fruit appears to be longer in P. cecropiifolia than 

easily confused. 

in (most?) other species of Pourouma. The dis- 

In addition to these characters one can also crepancy between the data supplied by Falcao 

regard as derived: sparse indument (e.g., Pourou- and Lleras and the observations of Martius might 

ma ovata), elliptic to obovate leaves (e.g., P. be caused by the presence of several strains. In 

phaeotricha and P. bolivarensis), unbranched lat- the publication "Underexploited Tropical Plants 

eral veins (e.g., P. acuminata and P. with 

ovata), peti- Promising Economic Value" (National 

olulate leaf segments (in taxa with incised leaves), Academy of Sciences, Washington, D.C., 1975) 

as in P. tomentosa subsp. persecta, short a 

petioles fruiting period of three months in the rainy 

of constant length (P. formicarum), swollen and season is mentioned for P. cecropiifolia. 

saccate base of the petiole (P. formicarum, P. 

myrmecophila), short stipules (e.g., P. ovata), sta- 

Pollination 

minate flowers in globose heads (P. melinonii 

and P. myrmecophila), subumbellate pistillate Little is known about pollination in Pourouinflorescences 

(P. minor), elongate peduncle of ma. Neither morphological features (of infloresthe 

pistillate inflorescence (P. ferruginea and P. cences and flowers) nor habit and habitat (small 

ovata), staminate flowers with fused tepals (P. to medium-sized trees in forests) suggest the ocmollis), 

stamens with partly or fully connate fil- currence of wind pollination. The herbarium maaments 

(P. melinonii and P. napoensis respec- terial indicates that most pistils develop into 

tively), and knob-like stigma (P. minor). fruits, which suggests an effective pollination sys- 

The above summation of derived characters tem. Falcao and Lleras (1980) found a number 

and specializations shows that they tend to be of bee species as pollinators of P. cecropiifolia. 

'at the Coussapoa side' of the variation spectrum The bees collect pollen in the staminate infloin 

Pourouma. 

rescences and fly to pistillate inflorescences. The

114 

authors do not give indications about the attractant(s) 

of the pistillate inflorescences. One 

may wonder whether the pluricellular ('glandular') 

hairs, which occur so abundantly in inflorescences 

of most Pourouma species (as a powdery 

layer, as in dry material?), may play a role 

(as attractant or a substitute for pollen?). 

Dispersal 

Again, little is known about dispersal of Pourouma 

seeds. One may assume that the 'fruits' 

with a fleshy mesocarp-like inner layer of the 

enlarged perianth, and a purple to black (or a 

red-brown) exocarp-like outer layer (skin) are 

eaten by several frugiverous arboreal animals; 

monkeys are mentioned in label data. 

The presence of very long peduncles of pistillate 

inflorescences in P. ovata and P. ferruginea 

could be an adaptation to dispersal by bats. 

Relations with Ants 

Pourouma formicarum and P. myrmecophila 

are myrmecophilous. These species have swollen 

and saccate bases of the petiole ("domatia"), usually 

occupied by small ants. According to Benson 

(1985) these ants belong to genus Allomerus and 

presumably eat food bodies, produced on the 

inner surface of the domatia. Sometimes, as in 

P. mollis subsp. triloba (collection Prance et al. 

7907) and most collections of P. napoensis, ants 

(of the genus Crematogaster) build tubular nests 

on the leafy twigs and on the lower leaf surface. 

The nest-building on the lower leaf surface (only 

if the main venation is subpalmate?) starts in the 

axils of the main veins; then the nests look like 

domatia. Later on these nests are often extended 

as tubes. A peculiar aspect is the use of the long, 

rather stiff hairs (occurring on various parts of 

the plant) in the nest building, perhaps as a protective 

cover of the nests. 


The distribution range of Pourouma is almost 

the same as that of Coussapoa. Pourouma is con- 

fined to rain forest areas, but is notably absent 

in the West Indies and southern Mexico. Most 

are lowland species, found at altitudes up to 1000 

m, occasionally up to ca. 1500 m, and excep- 

tionally (P. tomentosa in Bolivia and Peru) up 


to ca. 1800 or 1900 m. Some taxa (P. bolivarensis 

and P. guianensis subsp. venezuelensis) appear 

to be confined to altitudes of about 1000 m (600- 

1400 m). The unnamed collections Gentry et al. 

22866 and D. N. Smith et al. 4743 (see unnamed 

collections 1, p. 193) may represent a montane 

taxon. Most species are components of non-inundated 

forest, but P. acuminata is often found 

in (occasionally?) inundated forest. Pourouma 

(probably) needs light for germination and is 

commonly found in places with more or less distinct 

traces of previous disturbance of the forest 

(e.g., chablis) or in rather open forest (e.g., on 

rather poor ? sandy soils) and secondary forest. 

A number of species have a wide distribution. 

All these species are represented in the Amazon 

Basin and the Guianas. Two, P. bicolor and P. 

minor, extend to Central America, and P. melinonii 

to Panama. Pourouma mollis, P. velutina, 

and P. guianensis also occur in eastern Brazil, 

the latter in the coastal mountain range of Venezuela 

as well. Pourouma tomentosa, however, 

is confined to the Amazon Basin and the Guianas. 

Most of these wide-spread species are very 

variable in a confusing way. In five of these 

species, two or more infraspecific taxa are recognized. 

The other species are much more uniform and 

have much smaller or even very small distribution 

ranges. Only one of the this group of 

species (P. hirsutipetiolata, from northern and 

western Colombia) is subdivided into two subspecies 

(but in a more clear way than in the widespread 

species). This species and P. oraria (from 

western Colombia) are the only ones not occurring 

in the Amazon Basin or the Guianas. Of the 

remaining species, three (P. bolivarensis, P. saulensis, 

and P. stipulacea), are confined to the 

Guiana region, all probably with very small distribution 

ranges. The others are confined to the 

Amazon Basin, or almost so as P. acuminata, P. 

cucura, and P. ovata (found in the Guinean part 

of Venezuela). 

The present range of distribution of the very 

uniform P. cecropiifolia might be largely anthropogenous. 

The Amazon Basin and the Guianas in all respects 

represent the center of the genus. Three 

centers of distribution can be distinguished: 

(a) The Upper Amazon Basin (down to ca. 

60?W) with Pourouma acuminata, P. cecropiifolia, 

P. cucura, P. cuspidata, P. elliptica, P. fer-


ruginea, P. formicarum, P. herrerensis, P. myrmecophila, 

P. napoensis, P. ovata, P. phaeotricha, 

as well as P. bicolor subsp. tessmannii, P. mollis 

subsp. triloba, and P. tomentosa subspp. apiculata, 

persecta, and tomentosa. A concentration 

of taxa is found in the north-western part of this 

region. 

(b) The Guiana (or Guayana) region, the Lower 

Amazon Basin (Brazil: eastern Para and Amapa), 

including an extension to eastern Brazil, with 

the following taxa confined or distinctly connected 

to this area: Pourouma bolivarensis, P. 

saulensis, as well as P. bicolor subsp. digitata, P. 

melinonii subsp. melinonii, P. mollis subsp. mol- 

lis, P. tomentosa subsp. essequiboensis and subsp. Pourouma shows many morphological simimaroniensis, 

and P. velutina. 

larities to the African genus Myrianthus (with 7 

(c) The Pacific Coastal region (Colombia and species), revised by de Ruiter (1976). The sim- 

Ecuador) to Panama, with extensions to Gua- ilarities are found in the habit, the leaf shape and 

temala and through northern Colombia to north- dimension and their infraspecific variation, and 

western Venezuela, with Pourouma hirsutipetio- the (variable) length of the petiole. Furthermore, 

lata, P. oraria, P. bicolor subsp. chocoana and the staminate inflorescences show similarities, 

subsp. scobina, and P. melinonii subsp. glabrata. but the pistillate ones are different, as they are 

The features of the East-Brazilian populations capitate in Myrianthus, where the pistillate flowof 

P. guianensis, P. mollis, and P. velutina, re- ers are sessile and clustered in a single head. Both 

semble those of the material of the same species genera are macrospermous in contrast to the othin 

the Lower Amazon Basin and adjacent parts er genera of the Cecropiaceae. The two genera 

of the Guianas. 

also show strong ecological similarities and are 

Pourouma bicolor, P. melinonii, P. mollis and found in similar types of habitat. These two gen- 

P. tomentosa have one (or two) morphologically era may also be regarded not only as genetically 

distinct entities (subspecies) distinctly connected related but also as the most primitive among the 

with the Guiana (Lowland) region and the ad- Cecropiaceae. 

jacent part of the Amazon Basin. In P. bicolor The leaves of some taxa, Pourouma cecropiand 

P. melinonii one or two disjunct subspecies ifolia and P. aspera subsp. digitata, resemble those 

occur in the third center of the genus and in P. of Cecropia because of the great number of inguianensis 

a disjunct subspecies occurs in north- cisions, but the leaves are never peltate as they 

ern Venezuela. 

always are in Cecropia. Leaves of some other 

In the wide-spread species the material in the taxa resemble leaves of some Coussapoa species 

Upper Amazon Basin is morphologically more (see p. 5). 

variable than in the other parts of their distribution 

ranges. Pourouma cucura shares this trait. 

Systematic 

In 

Arrangement 

most parts of the distribution range of Pouof 

the 

rouma the taxa (even those at the 

Species 

subspecific 

level) can be recognized, and well-collected ma- On the basis of overall similarities, several 

terial of adult specimens can readily be identi- groups of species can be recognized. Not all 

fied. In contrast, the situation in the Upper Am- species can be placed with certainty into one of 

azon Basin, especially in its northwestern part, them, however, as the staminate inflorescences 

is rather unclear with regard to the delimitation are not known (as in P. bolivarensis, P. saulensis, 

of subspecies (even species in some cases). In and P. stipulacea). 

several species (e.g., P. bicolor, P. mollis, P. to- One group of species is characterized by havmentosa) 

morphologically aberrant collections ing the tepals of the staminate flower almost enhave 

been made, e.g., in the valley of the Rio tirely connate, forming an urceolate infundibu- 

Santiago (Peru, Amazonas). 

liform perianth with the filaments of the stamens 

115 

In some cases (P. cucura and P. tomentosa 

subsp. persecta) the present data suggest the oc- 

currence of a disjunction between populations in 

the southwestern part and those in the north- 

western part of the Amazon Basin. 

The genus appears to be underrepresented in 

a zone from southeastern to northwestern Para 

largely consisting of'mata de cip6' and north of 

the Rio Solim6es of patches of'campos de terra 

firme' (Pires, 1973). A similar disjunctive gap is 

found in Coussapoa (see p. 10). 

Systematic Position of the Genus


longer than the tepals. The staminate flowers are the fleshy (mesocarp-like) inner layer of the periusually 

clustered in distinct, terminal, globose anth strongly resembles that ofgrapes. P. cecropiheads. 

The arachnoid indument often occurs on ifolia (Amazon grape, uva, uvilla, imbauiba do 

the lateral veins of the lamina beneath or also vinho) is one of the taxa mentioned in "Underon 

other parts, such as the stipules. The lamina exploited Tropical Plants with Promising Ecois 

usually smooth above. The leaves are entire nomic Value" (National Academy of Sciences, 

or palmately incised. The basal part of the leaf Washington, D.C., 1975). P. cecropiifolia can start 

margin is formed by the basal part of the basal reproducing (thus forming fruits) a few years after 

lateral vein in all (or most?) species. Pourouma planting, as small trees. Other Pourouma species 

mollis, P. melinonii, P. hirsutipetiolata, P. to- may have fruits as tasty as those of P. cecropimentosa, 

P. herrerensis, and P. napoensis com- ifolia, but the fleshy mesocarp-like layer is usuprises 

this group, as may P. acuminata and P. ally thinner and many of them become mediumstipulacea, 

of which the staminate flowers are sized trees before fruiting. Pourouma cecropistill 

unknown. 

ifolia fruits are used to make a sweet wine. For 

About equally well-defined is a group formed a detailed study on the reproduction and propby 

Pourouma myrmecophila, P. formicarum, and erties of the fruits of P. cecropiifolia see Falcao 

P. phaeotricha, in which the laminae are mostly and Lleras (1980). 

scabrous above. The staminate flowers, with (almost) 

free tepals and short filaments, are clus- 

Taxonomy 

tered in distinct, terminal, globose heads. The 

hairs of the tepals and the stigma are long (com- Pourouma Aublet, Hist. pl. Guiane 2: 891. 1775; 

pared with other species). The lamina is entire Tr6cul, Ann. Sci. Nat. Bot., Ser. 3, 8: 100. 

or 3-lobed and the tendency toward entire leaves 1847; Miquel in Martius, Fl. bras. 4(1): 122. 

is strong. The base of the petiole of P. myrme- 1853; Bentham in Bentham & Hooker, Gen. 

cophila and P. formicarum is swollen and sac- pi. 3(1): 357. 1880; Baillon, Hist. pl. 6: 141. 

cate. These two species are myrmecophilous. 1875-1876; Engler in Engler & Prantl, Nat. 

Less well-defined is a group of species in which Pflanzenfam. 3(1): 95. 1889; Benoist, Arch. 

the lamina is usually scabrous and/or has more Bot. Mem. 5: 29. 1931; Woodson & Schery, 

than 7 incisions. The staminate flowers are more Ann. Missouri Bot. Gard. 47: 165. 1960. Berg 

or less clustered, but not in globose heads. In in Lanjouw & Stoffers, Fl. Suriname 5(1): 265. 

most of the species the lamina is incised. This 1975. Burger, Fieldiana Bot. 40: 200. 1977; 

group is formed by Pourouma guianensis, P. as- Berg & van Heusden, Proc. Kon. Ned. Akad. 

pera, P. velutina, P. cucura, P. cuspidata, and P. Wetensch., Ser. C., 91(2): 105. (1988). Type 

cecropiifolia. 

species. Pourouma guianensis Aublet. 

The rest of the species cannot be satisfactorily Trees, terrestrial, often with stilt-roots. Leaves 

grouped, except for pairs of species showing more in spirals, entire or 

or 

palmately incised, 

less distinct 

margin 

similarities, such as Pourouma entire to subcrenate; stipules fused, 

minor and P. bolivarensis or 

fully am- 

P. ovata and P. 

plexicaul. Inflorescences in the leaf axils, 

saulensis. 

mostly 

in pairs, branched (or to subumbellate in pistil- 

Pourouma ovata and P. ferruginea both have late inflorescences), 

pistillate inflorescences with 

mostly ebracteate, somevery 

long peduncles. times bracteate; staminate flowers sessile or 

But they are so different in other 

pedfeatures 

that 

icellate, 

this 

tepals (3-)4, free or 

similarity appears not 

connate; stamens 

to indicate common 

(3-)4, 

origin, but 

mostly free, sometimes connate; 

rather a similar 

pistillate 

adaptation (to dis- flowers pedicellate, perianth tubular; ovary free, 

persal by bats?). 

stigma (sub)peltate or sometimes-knob-like. Fruit 

an achene, large, surrounded by the enlarged, 

Uses 

more or less fleshy perianth with a purplish to 

blackish or red-brown outer 

Most, if not all, Pourouma species have edible 

layer at full matufruits. 

As far as known, only P. cecropiifolia is 

rity; seed without endosperm, embryo straight, 

cultivated as a fruit tree, however. The taste of 

cotyledons thick, radicle short.

Pourouma 117 

Key to the Species and Subspecies of Pourouma 

1. Base of the petiole swollen. 

2. Petiole 1-1.5 cm long ..................................................... 8. P. formicarum. 

2. Petiole 8-27 cm long ..................................................... 7. P. myremcophila. 

1. Base of the petiole not swollen. 

3. Stipules (sub)persistent. 


5. Stipules with dense, white, arachnoid hairs; Guyana. ...................... 14. P. stipulacea. 

5. Stipules without white, arachnoid hairs; Colombia .............................. 18. P. oraria. 

4. Lamina scabrous above ....................................... 3e. P. bicolor subsp. chocoana. 

3. Stipules caducous. 

6. Lamina more or less scabrous above. 

7. Lamina entire. 

8. Lateral veins 2 x 7-10 (in large leaves up to 13), the basal pair unbranched (or faintly 

branched); lamina mostly elliptic to oblong or to subobovate. 

9. Pluricellular hairs on the leafy twigs dense; staminate flowers in globose heads; pistillate 

inflorescence with 11-15 flowers. .......... ......................... 9. P. phaeotricha. 

9. Pluricellular hairs on the leafy twigs sparse or lacking; staminate flowers not in globose 

heads; pistillate inflorescence with 2-8 flowers. ........................ 2. P. velutina. 

8. Lateral veins 2 x 10-25, the basal pair branched; lamina mostly ovate. 

10. Lamina above yellowish-hirsute to -hirtellous over the whole surface. 

11. Staminate flower with connate tepals and filaments longer than the perianth; 

(fruiting) perianth of pistillate flower velutinous. . ............. 25. P. herrerensis. 

11. Staminate flower with (almost) free tepals and filaments shorter than the perianth; 

(fruiting) perianth of pistillate flower hispidulous to hirtellous ........ 5. P. cucura. 

10. Lamina above yellowish- to whitish-hirtellous only on the main veins. 

12. White, arachnoid hairs on the main veins of the lamina beneath and/or also on 

the stipules, petioles, leafy twigs, and peduncles ........................... 

.......................................... 13b. P. tomentosa subsp. apiculata. 

12. White, arachnoid hairs only in the areoles or also on the smaller veins of the 

lamina beneath. 

13. Hairs on the lamina beneath (more or less) appressed. .......... 3. P. bicolor. 

14. Stipules hairy inside. ...................... 3a. P. bicolor subsp. bicolor. 

14. Stipules glabrous inside. ............... 3b. P. bicolor subsp. tessmannii. 

13. Hairs on the lamina beneath at least partly patent. 

15. Stipules inside densely hairy; pluricellular hairs on the leafy twigs sparse 

or lacking; pistillate inflorescences with 2-8 flowers ........ 2. P. velutina. 

15. Stipules inside glabrous, or if with sparse hairs, then pluricellular hairs 

on the leafy twigs usually (rather) dense; pistillate inflorescences usually 

with at least 10 flowers. 

16. Pluricellular (brown) hairs sparse or lacking on the leafty twigs and 

petioles .. ............................... .......... 5. P. cucura. 

16. Pluricellular (brown) hairs (rather) dense on the leafy twigs and the 

petioles. 

17. Arachnoid hairs on the lamina beneath confined to the areoles; 

tepals of the staminate flowers (almost) free; apex of the pedicels 

at fruit not widened.. ....... la. P. guianensis subsp. guianensis. 

17. Arachnoid hairs on the lamina beneath also on the smaller 

veins; tepals of the staminate flowers connate; apex of the pedicels 

at fruit widened (aberrant form from Peru)............ 

.................................. 10. P. m ollis subsp. triloba. 

7. Lamina lobed to parted. 

18. Stipules hairy inside. 

19. Lamina with the hairs beneath on the veins more or less appressed and the smaller 

veins plane or only slightly prominent . ............................... 3. P. bicolor. 

20. Lamina (usually) 5-7(-9)-fid to -parted. 

21. Lamina with arachnoid indument beneath, covering also the almost plane, 

smaller veins. 

22. Stipules subpersistent; midsegment of the lamina oblong to elliptic; 

leafy twigs often hirsute; Panama and western Colombia .......... 

......... ..................... ...... 3e. P. bicolor subsp. chocoana.


22. Stipules caducous; midsegment of the lamina lanceolate; leafy twigs 

never hirsute; Guianas and Lower Amazon Basin (to southern Venezuela?). 

................................. 3c. P. bicolor subsp. digitata. 

21. Lamina with arachnoid indument beneath (almost) confined to the areoles, 

the smaller veins more or less prominent; Central America and the Pacific 

coastal region of Colombia, Ecuador, and northwest Venezuela (also Amazonian 

Peru?). ............................. 3d. P. bicolor subsp. scobina. 

20. Lamina (usually) 3-lobed to -fid. 

23. Lamina with arachnoid indument beneath, covering also the almost plane 

smaller veins or with arachnoid indument confined to the areoles, but then 

the spots of white indument distinctly separated by the thick smaller veins; 

pistillate inflorescences mostly with up to 25(-35) flowers; Guianas, Amazon 

Basin, to northern Colombia. ................. 3a. P. bicolor subsp. bicolor. 

23. Lamina with arachnoid indument beneath (almost) confined to the areoles, 

the smaller veins more or less prominent and thin; pistillate inflorescences 

with up to 50 flowers; Central America, the Pacific coastal region of Colombia, 

Ecuador, and to northwestern Venezuela (also Amazonian Peru?). 

........................................... 3d. P. bicolor subsp. scobina. 

19. Lamina with hairs beneath (partly) patent on the veins, the smaller veins prominent. 

24. Leafy twigs yellow-hirsute and/or the whole upper surface of the lamina hirtellous; 

fruiting perianth rarely scabrous; Upper Amazon Basin and northern Venezuela. 

..................................... .............. 1. P. guianensis. 

25. Lamina 3-5-fid to -parted; leafy twigs and petioles with long yellow hairs; 

stipules always hairy inside; fruiting perianth 2-2.5 cm long; northern Venezuela. 

............................ lb. P. guianensis subsp. venezuelensis. 

25. Lamina entire, 5-7-fid to -parted, or if 3-5-fid to -parted, then the leafy 

twigs and the petioles without long yellow hairs, the stipules glabrous inside 

and/or the fruiting perianth 1.2-2 cm long; Guiana region, eastern Brazil, 

Amazon Basin to eastern Colombia. ..... la. P. guianensis subsp. guianensis. 

24. Leafy twigs pale yellow-hirtellous (to subtomentose); upper surface of the lamina 

only hirtellous on the main veins; fruiting perianth usually scabrous; Central 

America, the Pacific coastal region of Colombia and Ecuador (also Amazonian 

Peru?). ................................ ...... 3d. P. bicolor subsp. scobina. 

18. Stipules glabrous inside. 

26. Lamina yellow-hirsute to -hirtellous above on the whole surface. 

27. Brown, pluricellular hairs on the leafy twigs and petioles sparse or lacking; 

arachnoid hairs on the lamina beneath also on the smaller veins. 

28. Lamina 7-fid to -parted, usually longer than 30 cm; perianth of pistillate 

flower hirtellous to hirsute; filaments connate. ............ 24. P. napoensis. 

28. Lamina at most 5-parted, usually up to 20 cm long; perianth of pistillate 

flower hispidulous; filaments free. ........................... 5. P. cucura. 

27. Brown, pluricellular hairs on the leafy twigs and petioles (rather) dense and/or 

the arachnoid hairs on the lamina beneath (almost) confined to the areoles. 

.......................................... la. P. guianensis subsp. guianensis. 

26. Lamina yellow-hirsute to -hirtellous or whitish strigose above only on the main 

veins. 

29. Pluricellular (brown) hairs sparse or lacking on the leafy twigs and petioles. 

30. Lamina 7-fid to -parted, usually longer than 30 cm; perianth of pistillate 

flower hirtellous to hirsute; filaments connate. ............ 24. P. napoensis. 

30. Lamina at most 5-parted, usually up to 20 cm long; perianth of pistillate 

flower hispidulous; filaments free. ........................... 5. P. cucura. 

29. Pluricellular (brown) hairs (rather) dense on the leafy twigs and petioles. 

31. Lamina with more or less appressed hairs on the veins beneath. ........ 

........................................ 3b. P. bicolor subsp. tessm annii. 

31. Lamina with (partly) patent hairs on the veins ....................... 

...................................... la. P. guianensis subsp. guianensis. 


32. Lamina scabrous beneath. 

33. Stipules glabrous inside. ..........................................6. P. cuspidata. 

33. Stipules hairy inside. ................................. 3a. P. bicolor subsp. bicolor. 

32. Lamina smooth beneath. 

34. Lamina 7-11-parted (to -fid).


35. Leafy twigs hirsute, or if glabrous or puberulous, then the incisions reaching 

down to the petiole and the segments often petiolulate. 

36. Lamina hirsute to hirtellous above, at least on the main veins ........... 

...................................................... 24. P. napoensis. 

36. Lamina (sub)glabrous above. ............. 13c. P. tomentosa subsp. persecta. 

35. Leafy twigs puberulous; incisions usually not entirely down to the petiole and 

leaf segments with a broader base. 

37. Base of the lamina usually deeply cordate; fruiting perianth 1.5-3.5 cm long; 

Upper Amazon Basin to eastern Colombia. .............. 4. P. cecropiifolia. 

37. Base of the lamina often truncate to shallowly cordate; fruiting perianth to 

ca. 1.5 cm long; Guianas and Lower Amazon Basin (to southern Venezuela?). 

............................................ 3c. P. bicolor subsp. digitata. 

34. Lamina entire or at most 5-parted. 

38. Petiole with a mixture of minute whitish hairs, long yellowish hairs, and sparse 

brown pluricellular hairs. 

39. Staminate flowers with connate tepals and filaments longerthan the perianth; 

(fruiting) perianth of pistillate flower velutinous. ......... 25. P. herrerensis. 

39. Staminate flowers with (almost) free tepals and filaments shorter than the 

perianth; (fruiting) perianth hispidulous to hirtellous. .......... 5. P. cucura. 

38. Petiole with indument otherwise. 

40. Lamina 3-lobed to 5-parted. 

41. Arachnoid hairs of indument confined to the lamina beneath in the 

areoles or also the smaller veins. 

42. Lamina beneath with the hairs on the veins appressed. 

43. Stipules hairy inside; tepals of the staminate flowers almost 

free. 

44. Leafy twigs with appressed hairs. ............ 3. P. bicolor. 

Repeat 20-20 for subspecies; see also unnamed collection 

#5. 

44. Leafy twigs with patent hairs. ............... 17. P. villosa. 

43. Stipules glabrous inside; tepals of the staminate flowers connate. 

45. Leafy twigs sparsely appressed-puberulous to strigose. 

...................................... 11. P. m elinonii. 

46. Base of the lamina truncate to subcordate (occasionally 

deeply cordate); fruiting perianth usually densely 

puberulous to subvelutinous; stamens with the filaments 

(almost) free; Guianas and Amazon Basin. .. 

.................. 1 la. P. melinonii subsp. melinonii. 

46. Base of the lamina deeply cordate; fruiting perianth 

almost glabrous; stamens with the filaments mostly 

connate, Panama, northern and western Colombia. 

1 lb. P. melinonii subsp. glabrata. 

45. Leafy twigs hirsute. . 12b. P. hirsutipetiolata subsp. hispida. 

42. Lamina beneath with the hairs on the veins patent. 

47. Leafy twigs with appressed hairs ........................ 

.......................... 1 la. P. melinonii subsp. melinonii. 

47. Leafy twigs with patent hairs. 

48. Basal part of the margin of the lamina formed by the basal 

lateral veins; tepals of the staminate flowers for the greater 

part connate; fruiting perianth velutinous. .... 10. P. mollis. 

49. Pistillate inflorescences with the flowers in two more 

or less distinct clusters; Guianas, eastern Brazil, Lower 

Amazon Basin. ........ 1 Oa. P. mollis subsp. mollis. 

49. Pistillate inflorescences with the flowers usually more 

or less diffusely distributed; Upper Amazon Basin and 

eastern Colombia. ....... 1Ob. P. mollis subsp. triloba. 

48. Basal part of the margin of the lamina separated from the 

basal lateral veins by mesophyll; tepals of the staminate 

flowers free or up to halfway connate; fruiting perianth 

sparsely puberulous ....................... 17. P. villosa. 

41. Arachnoid hairs also on the petiole, the main veins of the lamina be-


neath, the leafy twigs, the stipules, the peduncles and/or the perianth 

of the pistillate flower. 

50. Peduncle of the pistillate inflorescence 32-48 cm long; tepals of the 

staminate flower almsot free ..................... 16. P. ferruginea. 

50. Peduncle of the pistillate inflorescence 2-13 cm long; tepals of the 

staminate flower connate. 

51. Lower leaf surface tomentose to subvelutinous ........... 

................................ lOa. P. m ollis subsp. mollis. 

51. Lower leaf surface appressed-puberulous to strigose on the 

main veins. ............................... 13. P. tomentosa. 

52. Stipules hairy inside. ....13b. P. tomentosa subsp. apiculata. 


53. Leafy twigs yellow-hirsute (or glabrous). .......... 

.................. 13c. P. tomentosa subsp. persecta. 

53. Leafy twigs puberulous to hirtellous. 

54. Base of the lamina deeply cordate. ........... 

......... 13d. P. tomentosa subsp. essequiboensis. 

54. Base of the lamina truncate to subcordate. 

55. Apex of the lamina acuminate; heads of staminate 

flowers 2-3 mm in diameter; Guianas, 

Amapa, and near Manaus. ........... 

....... 13e. P. tomentosa subsp. maroniensis. 

55. Apex of the lamina usually rounded; heads 

of staminate flowers 4-6 mm in diameter; 

Upper Amazon Basin. .................. 

......... 13a. P. tomentosa subsp. tomentosa. 



57. Lamina broadest above or in the middle. 

58. Arachnoid hairs only on the lamina beneath in the areoles or 

also on the smaller veins. 

59. Leafy twigs and lamina beneath on the main veins with 

appressed hairs. ...................... 22. P. bolivarensis. 

59. Leafy twigs and lamina beneath on the main veins with 

patent hairs. 

60. Basal (most) part of the margin of the lamina formed 

by the basal lateral veins; tepals of the staminate flower 

for the greater part connate. .......... 10. P. mollis. 

Repeat 48-48 for subspecies. 

60. Basal (most) part of the margin of the lamina separated 

from the basal lateral veins by mesophyll; tepals 

of the staminate flower free or up to halfway connate. 

61. Stipules hairy inside. .............. 17. P. villosa. 

61. Stipules glabrous inside. ......... 21. P. elliptica. 

58. Arachnoid hairs also on the petiole, the main veins of the 

lamina beneath, the leafy twigs, the stipules, the peduncle and/ 

or the perianth of the pistillate flower ..................... 

........................ 13a. P. tomentosa subsp. tomentosa. 

57. Lamina broadest below the middle. 

62. Lamina with ? patent hairs on the main veins beneath. 

63. Basal part of the margin of the lamina formed by the basal 

lateral veins; tepals of the staminate flower for the greater 

part connate. 

64. Leafy twigs with patent hairs. .......... 10. P. mollis. 

Repeat 49-49 for subspecies; see also unnamed collection 

#2. 

64. Leafy twigs with appressed hairs. ................ 

.................. 1 a. P. melinonii subsp. melinonii. 

63. Basal part of the margin of the lamina separated from the 

basal lateral veins by mesophyll; tepals of the staminate 

flower free or up to halfway connate. ........ 17. P. villosa. 

See also unnamed collection #4. 

62. Lamina with appressed hairs on the main veins beneath.


65. Arachnoid hairs confined to the areoles or also on the 

smaller veins on the lamina beneath. 

66. Stipules densely or sparsely hairy inside. 

67. Staminate flowers with connate tepals and filaments 

longer than the perianth; (fruiting) perianth 

of pistillate flower velutinous; stipules 

sparsely hairy inside. .......... 25. P. herrerensis. 

67. Staminate flowers with (almost) free tepals, filaments 

shorter than the perianth; (fruiting) perianth 

hispidulous, scabrous; stipules densely hairy 

inside. .............. 3a. P. bicolor subsp. bicolor. 

See also unnamed collections #1, #3, and #4. 


68. Leafy twigs sparsely appressed-puberulous to 

strigose. ...................... 11. P. melinonii. 

Repeat 46-46 for subspecies. 

68. Leafy twigs hirsute ........................ 

..... 12a. P. hirsutipetiolata subsp. hirsutipetiolata. 

65. Arachnoid hairs also on the petiole, the main veins of the 

lamina beneath, the leafy twigs, the stipules, the peduncles 

and/or the perianth of the pistillate flower. 

69. Peduncle of the pistillate inflorescence 32-48 cm long; 

tepals of the staminate flower almost free. ........ 

.................................. 16. P. ferruginea. 

69. Peduncle of the pistillate inflorescence 2-13 cm long; 

tepals of the staminate flower connate. ........... 

.................................. 13. P. tom entosa. 

70. Stipules hairy inside........................ 

.............. 13b. P. tomentosa subsp. apiculata. 


71. Apex of the lamina usually rounded to emarginate, 

sometimes short-acuminate; heads 

of the staminate inflorescences 4-6 mm in 

diameter; Upper Amazon Basin .......... 

......... 13a. P. tomentosa subsp. tomentosa. 

71. Apex of the lamina acuminate; heads of the 

staminate inflorescences 2-3 mm in diameter; 

Guianas, Amapa, and near Manaus. 

....... 13e. P. tomentosa subsp. maroniensis. 

56. Basal lateral veins unbranched. 


73. Leafy twigs white-puberulous to subglabrous. 

74. Lateral veins 2 x 7-12; stipules glabrous inside; peduncle 

of the pistillate inflorescence 3-5 cm long. . 19. P. saulensis. 

74. Lateral veins 2 x 9-21; stipules hairy inside; peduncle of 

the pistillate inflorescence up to 32 cm long. .. 20. P. ovata. 

73. Leafy twigs yellow-sericeous to -hirtellous .... 9. P. phaeotricha. 


75. Lower surface of the lamina initially entirely or largely covered 

with white or brownish, arachnoid hairs; peduncle of the pistillate 

inflorescence up to ca. 50 cm long. ..... 16. P. ferruginea. 

75. Lower surface of the lamina with white arachnoid hairs in the 

areoles only (or sometimes extending to the main veins); peduncle 

of the pistillate inflorescence up to ca. 10 cm long. 

76. Lamina with dense yellow hairs on the main veins beneath. 

77. Leafy twigs with very short and much longer yellow 

hairs. ............................... 21. P. elliptica. 

77. Leafy twigs with hairs of about equal length. ...... 

...................................... 23. P. m inor. 

76. Lamina with sparse and inconspicuous hairs on the main 

veins beneath. 

78. Hairs on leafy twigs very short ..... 25. P. herrerensis. 

78. Hairs on leafy twigs long or absent. ... 15. P. acuminata.


1. Pourouma guianensis Aublet, Hist. pl. Guiane 

2: 892, t. 341. 1775; Miquel in Martius, Fl. 

bras. 4(1): 127. 1853; Berg, Fl. Suriname 5(1): 

267. 1975; Croat, Flora Barro Colorado Island 

360. 1978. Type. French Guiana. Sinnamary 

R., without date (Y), Aublet s.n. (holotype, BM, 

only photographs in NY and U seen; isotypes?, 

LE, S). 

(8-)10-25; pedicel 0.2-0.5 cm, in fruit 0.4-1 cm 

long, perianth 2-4 mm long, subvelutinous, 

sometimes sparsely puberulous, stigma subpel- 

tate, 1.5-2 mm diam., shortly pilose. Fruiting 

perianth purple, (sub)ovoid to ellipsoid, 1.2-2 or 

2-2.5 cm long, mostly (sub)velutinous, some- 

times sparsely puberulous, scabrous or almost 

glabrous. 

This taxon can be difficult to distinguish from 

P. bicolor. The two taxa have overlapping dis- 

tributions and are each very variable with, more- 


thick, white-puberulous, white- to yellow-hirtellous, 

yellow-(sub)velutinous or yellow-hirsute, over, quite an overlap of characters and their 

with dense to/or (very) sparse, brown, pluricel- variation. Considering the variation in both taxa, 

lular hairs; lenticels rather conspicuous. Lamina one would be inclined to combine the two. How- 

3-fid to 3-lobed, 3- or 5-fid (to -lobed or to -part- ever, as distinct morphological entities occur in 

ed), sometimes 5-7-fid to -parted ca. 10-25(-45) the same areas without having clear intermedix 

10-25(-45) cm, or entire (to 3-lobed), broadly ates it is likely that they are genetically separated; 

ovate to elliptic (to oblong), (4-)10-20 x (2-)7- only in a few cases do the collected specimens 

14 cm, subcoriaceous to chartaceous, apex acu- show ? intermediate features. 

minate, base cordate, the sinus rather shallow Pourouma guianensis can be distinguished 

and wide or deep and narrow, often with over- from P. bicolor in the following ways: 

lapping lobes, sometimes truncate, in entire leaves (a.) The indument of the inner surface of the 

to rounded (to subacute); upper surface scabrous, stipules. In subsp. guianensis the inner surface 

on the main veins hirtellous or hirsute or the of the stipules is usually glabrous or with sparse, 

whole surface hirtellous to subvelutinous, lower white hairs. However, in some forms of subsp. 

surface subtomentose to subvelutinous or hir- guianensis and in subsp. venezuelensis the inner 

tellous, or on the smaller veins to puberulous, surface of the stipules is generally covered with 

hairs on main veins often + appressed, white, (rather dense) yellow hairs, apparently correlated 

arachnoid hairs usually (almost) confined to the with the yellow-hirsute indument on the leafy 

areoles; tertiary venation and reticulum + prom- twigs, the outer surface of the stipules, and other 

inent (and conspicuous) beneath; lateral veins parts. Pourouma bicolor (except for subsp. tess- 

(7-)12-26 pairs, basal pair branched; petiole 4- mannii.) is likewise characterized by the occur- 

25(-40) cm long, whitish-yellow-puberulous to rence of dense, whitish to yellow hairs on the 

-hirtellous to -hirsute and with dense to (very) inner surface of the stipules, but in this species 

sparse, brown, pluricellular hairs; stipules (2-)4- (except for subsp. chocoana.) this indument co- 

15 cm long, caducous, outside whitish- (to pale occurs with an indument on the leafy twigs, petyellow-)subsericeous 

to -subhirsute to -subto- ioles, and outer surface of the stipules consisting 

mentose or to -subvelutinous or yellow-hirsute, of short, appressed hairs. 

and with dense to (very) sparse, brown, pluri- (b.) The indument on the smaller veins of the 

cellular hairs, inside glabrous or with sparse, white lamina beneath. In P. guianensis this consists of 

hairs or with rather dense, yellow hairs. Stami- relatively long, patent hairs. In P. bicolor the 

nate inflorescences up to 20 cm long and 12 cm hairs on both the main veins and the smaller 

wide; peduncle 4-6 cm long, peduncle and veins is mostly appressed, only occasionally pabranches 

puberulous to hirtellous to (sub)hirsute tent on the smaller veins and then mostly shorter 

or to subvelutinous, often with dense, brown, than in P. guianensis. Only in several specimens 

pluricellular hairs; flowers + clustered, but not of P. bicolor subsp. scobina does the indument 

in (sub)globose heads; tepals lanceolate, (almost) resemble that normally occurring in P. guianenfree; 

filaments shorter than the tepals. Pistillate sis. 

inflorescences in fruit up to 20 cm long and 12 (c.) The prominence of the reticulum and occm 

wide; peduncle (3-)6-15(-20) cm long, in- currence of the white-arachnoid indument on the 

dument of the peduncle and branches similar to lamina beneath. In P. guianensis the reticulum 

that of the staminate inflorescence; flowers is prominent and, moreover, conspicuous as the


white-arachnoid indument is usually confined to 

the areoles. In P. bicolor the reticulum is, in most 

of the subspecies, merely slightly prominent to 

plane and usually inconspicuous as the arachnoid 

indument usually also occurs on the smaller veins. 

However, in a form of P. bicolor subsp. bicolor, 

occurring in the western part of the range of the 

subspecies, the arachnoid indument is confined 

to the very small areoles surrounded by relatively 

thick veinlets, and in P. bicolor subsp. scobina 

the arachnoid indument is also more or less confined 

to the areoles, but in this subspecies this 

feature is combined with ? prominent smaller 

veins. 

In P. guianensis two allopatric subspecies can 

be recognized. 

Key to the Subspecies 

of Pourouma guianensis 

1. Lamina 3-5-fid to -parted; leafy twigs and petioles 

with long yellow hairs; stipules hairy inside; 

fruiting perianth 2-2.5 cm long; northern Venezuela 

................ b. subsp. venezuelensis. 

1. Lamina entire, 5-7-fid to -parted, or if 3-5-fid 

to -parted, then the leafy twigs and the petioles 

without long yellow hairs, the stipules glabrous 

inside, and/or the fruiting perianth 1.2-2 cm long; 

Guiana region, eastern Brazil, Amazon Basin to 

eastern Colombia. ........ a. subsp. guianensis. 

la. Pourouma guianensis Aublet subsp. 

guianensis. Fig. 59. 

Pourouma palmata Poeppig & Endlicher, Nov. gen. 

sp. pi. 2: 29, t. 141. 1838; Tr6cul, Ann. Sci. Nat. 

Bot., Ser. 3, 8: 104. 1847; Miquel in Martius, Fl. 

bras. 4(1): 126. 1853; Macbride, Publ. Field Mus. 

Nat. Hist., Bot. Ser., 13(2.2): 293. 1937. Type. Peru. 

San Martin: Tocache, Aug 1830 (6 and 9), Poeppig 

s.n. or 1881 (holotype or syntypes, W, destroyed; 

lectotype, G (Q specimen!), fragment of 6 specimen 

in F). 

Pourouma acutiflora Trecul, Ann. Sci. Nat. Bot., S&r. 

3, 8: 105. 1847; Miquel in Martius, Fl. bras. 4(1): 

126. 1853. Type. Brazil. Rio de Janeiro: Without 

locality, 1839 (6), Guillemin 1024 (holotype, P). 

Pourouma cinerascens Martius ex Miquel in Martius, 

Pourouma scabra Rusby, Bull. New York Bot. Gard. 

6: 498. 1910. Type. Bolivia. Pando: Santa Barbara, 

30 Aug 1902 (9), R. S. Williams 1560 (holotype, NY; 

isotype, US). 

Pourouma radula Benoist, Bull. Mus. Hist. Nat. (Paris) 

28: 320. 1922; Woodson & Schery, Ann. Missouri 

Bot. Gard. 47: 166, t. 58. 1960. Type. Colombia. 

Without locality, without date (a), Triana 860 (ho- 

lotype, P; isotypes, E, NY). 

Pourouma subtriloba Rusby, Mem. New York Bot. 

Gard. 7: 232. 1927. Type. Bolivia. La Paz: Nr. Tu- 

mapasa, 12 Oct 1921 (9), Cardenas 1990 (holotype, 

NY; isotypes, GH, US). 

Pourouma substrigosa Mildbraed, Notizbl. Bot. Gart. 

Berlin-Dahlem 10:192. 1927; Macbride, Publ. Field 

Mus. Nat. Hist., Bot. Ser., 13(2.2): 293. 1937. Type. 

Peru. Loreto: Pongo de Manseriche, 26 Nov 1924 

(8), Tessmann 4642 (holotype, B, isotypes, B, G, 

NY). 

Pourouma mildbraediana Standley, Publ. Field Mus. 

Nat. Hist., Bot. Ser., 17: 183. 1937. Type. Brazil. 

Amazonas: Mun. Sao Paulo de Olivenoa, nr. Pal- 

mares, 11 Sep-26 Oct 1936 (a), Krukoff 8386 (ho- 

lotype, NY; isotypes, A, BR, F, G, LE, MO, P, S, U, 

US). 

Leafy twigs pale to bright yellow-hirtellous to 

-subvelutinous to -hirsute, brown, pluricellular 

hairs dense to sparse. Lamina entire, 3-5-lobed 

to -parted, or sometimes 5-7-parted, base shal- 

lowly to deeply cordate or sometimes rounded; 

lateral veins mostly 10-16, sometimes up to 20, 

or occasionally up to 25 pairs; stipules outside 

pale to bright yellow-hirtellous, -subvelutinous, 

-subtomentose or -hirsute, inside glabrous or 

sometimes hairy. Fruitingperianth 1.2-2 cm long, 

mostly densely hairy, sometimes sparsely hairy 

or scabrous. 

Distribution (Fig. 60). Throughout the Ama- 

zon Basin, extending to eastern Colombia and 

the Guiana region; occurring disjunctly in east- 

ern Brazil (from Pernambuco to Santa Catarina); 

mostly in non-inundated forest, sometimes in 

periodically inundated (varzea) forest; mostly at 

low altitudes, in Venezuela (Bolivar) up to ca. 

1300 m, in Bolivia up to 1500 m. 

Specimens examined. Colombia. Without locality, 

without date (d), Triana 860 (E, NY, P, type collection 

Fl. bras. 4(1): 125, t. 37. 1853. Type. Brazil. Ama- of P. radula), without date (6 and 9), Triana s.n. (E, P). 

zonas: Rio Japura, Maripi, without date (6), Martius META: Puerto Lopez, 3 Aug 1944 (st), Liitle et al. 8423 

s.n. (holotype, M; isotypes, M, U). 

(F); Villavicencio, Llano de San Martin, without date 

Pourouma heterophylla Martius ex Miquel in Martius, (9), Karsten s.n. (GOET, LE). 

Fl. bras. 4(1): 125. 1853. Type. Brazil. Amazonas: VENEZUELA. AMAZONAS: Rio Manavicke, Indian 

Rio Japura, Dec 1819 (st, juv), Martius s.n. (holo- village "Kalchi-Teri," without date (juv.), Lizot s.n. 

type, M; isotypes, M, U). 

(VEN); Sierra Parima, 1300 m, 18-23 May 1972 (st), 

Pourouma fulginea Miquel in Martius, Fl. bras. 4(1): Steyermark 106086 (NY); Parima Mts., nr. Simara- 

129. 1853. Type. Brazil. Amazon Basin, without date wochi, Rio Matacuni, 18 Apr-23 May 1973 (d), Stey- 

(Q), Martius s.n. (holotype, M; isotype, U). ermark 107154 (F, U, VEN). BOLIVAR: Reserva Fo-


JII'd~~~~~~~i 

_ ,e 

4~, ,. \ 

as t af + 

POUROUMA einerascenx

Pourouma125 

restal "La Paragua," Rio Asa, Jun 1970 (st,juv), Blanco 

810 (VEN); slopes of Quebrada O-paru-ma, between 

Santa Teresita de Kavanayen and Rio Pacairao (tributary 

of Rio Mouak), alt. 1065-1220 m, 20-21 Nov 

1944 (2), Steyermark 60407 (F, VEN); 45 km N of 

Tumeremo, Sierra de Nuria, 5-8 Feb 1961 (st), Steyermark 

89134 (VEN); Rio Nichare (tributary of Rio 

Caura), nr. the confluence with Rio Cicuta, 25 Apr 

1966 (2), Steyermark et al. 95711 (VEN). DELTA 

AMACURO: E of Rio Grande, ENE of El Palmar, 29 

Nov-18 Dec 1964 (2), Marcano Berti 471 (NY, VEN), 

19 Jan 1965 (2), Marcano Berti 611 (F, NY, VEN); Rio 

(9), Huashikat 1216 (BG); Rio Cenepa, 30 Jan 1973 

(6), Kayap 268 (F, GH, MO), 21 Feb 1973 (9), Kayap 

393 (F, GH); Quebrada Huampami, 5 Jul 1974 (9), 

Kayap 1057 (F, GH, MO). HuANuco: Tingo Maria, 

13 Aug 1940 (9), Asplund 13020 (G, A, S, US); Pachita, 

Codo de Pozuzo, 24 Oct 1982 (6), Foster 9373 (BG); 

Distr. Churubamba, Rio Cayumba, 11 Sep 1936 (9), 

Mexia 8172 (F, G, GB, GH, MO, NA, NY, S, U, UC, 

US); between Monz6n and Rio Huallaga, without date 

(9), Weberbauer 3639 (G). LORETO: Prov. Coronel Portillo, 

Dist. Calleria, rd. Pucallpa-Huanuco, km 34, 23 

May 1968 (6), Castillo S. 11 (DUKE, F, P, US); Que- 

Amacuro, Venezuela-Guyana frontier, Imataca Mts., brada Shanuce, above Yurimaguas, 11 Jul 1972 (2), 

downstream from San Victor, 4 Nov 1960 (st), Stey- Croat 18016 (F, MO, NY); Rio Napo, Caserio Bella 

ermark 87307 (NY, VEN). 

Vista, 22 Sep 1972 (6), Croat 20161 (C, DUKE, F, GH, 

GUYANA. Kartabo region, Kartabo, 22 Jul 1920 NA, NY); Rio Ucayali, Jenaro Herrera, 7 Dec 1977 

(st, juv) Bailey 18 (GH), Bailey 19 (GH), Bailey 20 (st), Gentry et al. 21205 (MO, U); Prov. Maynas, Yana- 

(GH), 20 Aug 1920 (st), Bailey 146 (GH), Bailey 147 mono Explorama Tourist Camp, 15 Jul 1983 (st), Gen- 

(GH); Pomeroon district, Kamwatta, 21 Sep 1921 (st), try et al. 43066 (BG); Pongo de Manseriche, nr. mouth 

Cruz 1175 (NY); between Demarara R. and Berbice of Rio Santiago, 2 Dec 1931 (d), Mexia 6201 (F, G, 

R., 19 Jul 1922 (9), de la Cruz 1657 (F, MO, NY, US); GB, GH, MO, NY, S, U, UC, US), 26 Nov 1924 (8), 

Kurumaikabra Creek, Essequibo R., 22 Nov 1930 (2), Tessmann 4642 (B, G, NY, type collection of P. sub- 

FD 1011 (FHO); Takutu Creek to Puruni R., Mazaruni strigosa). MADRE DE DIos: Parque Nacional del Manu, 

R., 25 Oct 1944 (6), Fanshawe 2050 (=FD 4786) (A, Rio Manu, Cocha Cashu Station, Jul 1978 (st), Foster 

F, NY, U, US); Mazaruni station, forest nursery, 1945 et al. 6568 (F); Rio Tambopata, at mouth of Rio D'Or- 

(st, juv), FD 4970 (U); FD 5011 (U, S); Kanuka Mts., bigny, 1 and 2 Mar 1981 (st), Gentry et al. 31816 and 

Iramaipang, Nov 1948 (6), FD 5936 (NY); 11/2 mile 31933 (U). PASCO: Prov. Oxapampa, Vivero, Puerto 

along Bartica-Potaro rd., without date (juv), FD 6914 Bermudez, 16 Jan 1983 (st), Gentry et al. 42029 (BG). 

(U); Tumatumari, 18 Jun-8 Jul 1921 (st), Gleason 174 SAN MARTIN: Tocache, Aug 1830 (6 and 9), Poeppig 

(GH, NY); Pakaraima Mts., just N of Paruima Mis- s.n. (or 1881) (F, G, type collection of P. palmata); 

sion, 19 Oct 1981 (9), Maas et al. 5858 (U); basin of Maynas Alto, without date (9), Poeppig s.n. (B, OXF); 

Shodikar Creek, tributary of Essequibo R., 8-22 Jan without locality, without date (6), Poeppig s.n. (LE, P); 

1938 (2), A. C. Smith 2845 (A, F, G, MO, NY, P, S, Distr. Tocache, rd. to Almendras, 7 Aug 1969 (2), J. 

U, US); Marudi Mts., 02?15'N, 59?10'W, 12 Nov 1982 Schunke V. 3309 (F, G, US); rd. to Shunthe, E ofPuente 

(2), Stoffers et al. 311, 312 (U). 

de Palo Blanco, 14 Jul 1974 (9), J. Schunke V. 7415 

SURINAM. Raleigh falls, base of Voltzberg, Nature (MO, NY, U). 

Reserve on the Coppename R., 18 Nov 1979 (9), Mori BRAZIL. Without locality, 1839 (3), Guillemin 1024 

et al. 8662 (NY, U). 

(P, type collection of P. acutiflora); "Amazon Basin," 

FRENCH GUIANA. Without locality, without date without locality, without date (9), Martius s.n. (M, U, 

(2), Aublet s.n. (BM, LE, ?S, type collection of P. gui- type collection of P. fulginea). ACRE: Cruzeiro do Sul, 

anensis); nr. Saul 3 Oct 1973 (9), Granville et al. B.5071 21 Feb 1976 (9), Marinho 273 (IAN), 18 Feb 1976 (2), 

(U); Paul Isnard region, 5 km of Citron, 12 Nov 1982 Monteiro et al. 477 (INPA, MG); rd. Rio Branco-Porto 

(2), Granville 5284 (U); Oyapock R., Trois Sauts, 13 Acre, km 83, 10 Oct 1980 (9), Nelson 681 (BG) 

Jul 1774 (st), Grenand 445 (CAY); Zidock Ville, 26 

Aug 1976 (st), Grenand 1352 (CAY); Comte R., Cacao, 

25 Feb 1965 (2), Halle 1132 (U, P); nr. Sail, Jan 1975 

(2), Moretti 102 (CAY); Comte R., ca. 60 km S of 

Cayenne, 26 Feb 1965 (9), Oldeman 1179 (CAY); Saiil, 

19 Oct 1971 (2), Oldeman B.4108 (CAY, P, U); Kourou, 

without date (9), (herb.) Richard s.n. (P). 

ECUADOR. NAPO: Rio Pucino, first major tributary 

of Rio Aguarico, above bridge at Aguarico, 10 Feb 

1974 (2), Gentry 9824 (U); Rio Napo, 8 km below 

Puerto Mishualli, 25-27 Mar 1986 (2) Neill et al. 7082 

(BG). 

PERU. AMAZONAS: 6 Apr 1973 (2), Ancuash 178 (F, 

GH, MO); Rio Santiago, nr. Caterpiza, 9 Nov 1979 

= Cid 

et al. 2870 (U); rd. Sena Madureira-Rio Branco, nr. 

km 7, 30 Sep 1968 (6), Prance et al. 7670 (GH, INPA, 

M, NY, R, S, U, US). AMAZONAS: Rio Antimari 30 

Feb 1904 (st), Huber (MG) 4244 (MG); Rio Madeira, 

nr. Calama, 7 Nov 1931 (9), Krukoff 1297 (G, NY, P, 

S, U); nr. mouth of Rio Embira, 14 Jan 1933 (9), Krukoff4817 

(A, F, G, MO, NY, S, U, US); Mun. Humaita, 

Tres Casas, 14 Sep-11 Oct 1934 (o), Krukoff6238 (B, 

BR, F, LE, MO, NY, S, U, US); Sao Paulo de Olivenca, 

nr. Palmares 11 Sep-26 Oct 1936 (9), Krukoff8369 (A, 

B, BR, F, G, LE, NY, P, U), (6), Krukoff8386 (A, BR, 

F, G, LE, MO, NY, P, S, U, US, type collection of P. 

mildbraediana); Sao Paulo de Olivenca, Creek Belem, 

26 Nov-11 Dec 1936 (2), Krukoff8652 (A, BR, F, G, 

FIG. 59. Pourouma guianensis subsp. guianensis. From Martius, Flora Brasiliensis 4(1). 1853: tab. 37 (as 

P. cinerascens). Leafy twig with staminate inflorescence, leaf apex, part of staminate inflorescence (28), diagram 

of staminate flower (D), staminate flower (1).

~~~ II ~~~ I 

P. guiamnonsi 

^%^_ 1 ^vf'^^/9^^ ^^\?^^^/ v * \ subsp. quirnnisL 

---- ---------- - -- 

^l - 

i~subap. vwwwcLensnir, 

- '--- - - p l 

FIG. 60. Distribution of Pourouma guianensis subsp. guianensis, P. guianensis subsp. venezuelenyis', 

Pourou- 

ma velutina.


LE, MO, NY, P, S, U, US); Mun. Coari, Rio Coari, 

20 Apr 1976 (st), Magnago et al. (INPA) 58046 (INPA); 

Rio Japura, Dec 1819 (st, juv), Martius s.n. (M, U, 

type collection ofP. heterophylla); Rio Japura, Mapiri, 

without date (8), Martius s.n. (M, U, type collection of 

P. cinerascens); Manaus-Itacoatiara rd., km 27, Reserva 

Florestal Ducke, 30 Jun 1976 (st), Mello (INPA) 

57509 (INPA); km 26,6 Jan 1977 (st), Nascimento 303 

(INPA); Rio Canuma, Mun. Nova Olinda do Norte, 

Nov 1976 (st), Monteiro 1295 (INPA); Rod. BR-174, 

Reserva Biologica do INPA, 28 Sep 1976 (st), Mota 

740 (INPA); Reserva Florestal Ducke, 11 Jan 1976 

(st), Nascimento 387 (INPA), Nov 1972 (st), Rodrigues 

9219 (INPA); rd. Humaita to Labrea, km 80, between 

Rios Ipixuna and Itaparana, 29 Jan 1970 (9), Prance 

et al. 3260 (F, INPA, NY, R, S, U, US); basin of Rio 

Demeni, nr. Totobi, 25 Feb 1969 (9), Prance et al. 

10233 (C, G, INPA, MO, NY, R, S, U, US); Manaus- 

P6rto Velho rd. BR-319, km 175, between Rio Tupana 

and Rio Igapo-aqu, 11 Oct 1974 (9), Prance et al. 22803 

(INPA, MO, U). BAHIA: Agua Preta, 13 Oct 1937 (a), 

Bondar s.n. (GUA); Rio das Contas, 3 Oct 1919 (8), 

Curran 19 (C, F, MO, NY, US); rd. from Camaca- 

Canavieiro, Apr 1965 (9), Magalhaes 733 (M); nr. Urucuca, 

4 Nov 1978 (9), Mori et al. 11046 (RB, U); Ilh6us, 

Sep 1821 (9 and d), Riedel 2 or s.n. (LE, mixed with 

sterile specimen of P. mollis), 10 Nov 1944 (9), Velloso 

723 (R). CEARA: Without locality, without date (st), 

Allemao et al. 446 (R); Serra de Baturit6, Sep 1897 (st), 

Huber 238 (MG). EspiRITO SANTO: 9 Aug 1965 (9), 

Belem 1471 (MG, NY, U); rd. BR-5 (BR-101), Morro 

Dantos, 8 Aug 1965 (9), Lanna Sobrinho 1011 (FEE- 

MA, U); nr. Linhares, 1 Oct 1971 (6), T. S. Santos 

2035 (MG, NY, U). MATO GROSSO: Rd. to Fontanilha, 

15 Sep 1976 (9), Gomes et al. 326 (INPA); Sarar6, 10 

Aug 1978 (9), Pires et al. 16547 (MG); Nuicleo de Humboldt, 

(9), Roth 36 (INPA); Rio Juruena, road to Aripuana, 

km 5, 9 Jul 1977 (a), M. G. Silva et al. 3305 

(MG); Aripuana, rd. to airport, 2 Jun 1979 ($), M. G. 

Silva et al. 4750 (MG). MINAS GERAIS: Viosa, 16 May 

1935 (9), Kuhlmann 2075 (US), 1936 (9), Kuhlmann 

(RB) 2200 (RB); Mun. Caratinga, Faz. Montes Claros, 

23 Jan 1980 (9), Nsihimura 39 (FEEMA, GUA, U); 

Rio Novo, - da Divisa, 19 Dec 1964 (6), Hatschbach 12049 (F); 

above Antonia, 18 Jan 1966 (9), Hatschbach et al. 13537 

(F, NY, P, RB, U, US); Mun. Guaraquecaba, Rio Bananal, 

8 Dec 1970 (9), Hatschbach 25765 (C, MO, NA, 

S, UC); Rio do Costa, 9 Feb 1972 (9), Hatschbach 

29129 (NA). PERNAMBUCO: Quipapa, Eng. Brejino 22 

Mar 1967 (9), Andrade-Lima 67-4983 (F). Rio DE 

JANEIRO: Tingua, 1 Oct 1946 (6), Brade et al. 18619 

(GUA, RB, U); Palmeiras, 13 Jan 1877 (st), Glaziou 

8935 (E, P); Serra dos Orgaos, Tabuinha, 21 Mar 1980 

(st), Glaziou 12173 (P, mixed with Cecropia glaziovii); 

Serra da Estrella, nr. Mandioca, 15 May 1880 (6), Glaziou 

12174 (C, E, P), without date (6), Glaziou 20408 

(C, P); Matta do Teizeira Borges, nr. Gavea, 12 Dec 

1928 (9), Pessoal do Horto Florestal 650 (RB); Matta 

dos Tres Barros, nr. Gavea, 14 Mar 1927 (9), Pessoal 

do Horto Florestal 651 (RB). RONDONIA: Mineraao 

Campo Novo, 100 km SW of Ariquemes, 10 Oct 1979 

(9), Zarucchi 2744 (INPA, NY). RORAMA: Rio Catrimani, 

13 Feb 1975 (9), Pires 15088 (IAN); Serra Tepequem, 

west facing slopes, 1200 m, 17 Feb 1967 (9), 

Prance et al. 4438 (G, INPA, M, NY, P, R, S, U, US); 

Indian trail Surucucu-Uaica, nr. Uaica airstrip, Rio 

Uraricoeira, 2 Mar 1971 (9), Prance et al. 10836 (F, 

GH, INPA, M, NY, P, S, U); between Botamatatedi 

and Maita, 9 Feb 1971 (9), Prance et al. 13579 (F, 

INPA, NY, U, US). SANTA CATARINA: Brusque, 15 Feb 

1951 (9), Klein 89 (US); nr. Blumeneau, 18 Jan 1955 

(9), Klein 110 (NY, S, UC, US); Brusque, 15 Feb 1952 

(9), Veloso 89 (RB); nr. Blumeneau, Nov 1888 (6), Ule 

991 (F, HGB, P, US). SAO PAULO: Ubatuba, 8 Jan 

1985 (9), Gentry et al. 49346 (BG). 

BOLIVIA. LA PAZ: Nr. Tumapasa, 12 Oct 1921 

(9), Cardenas 1990 (GH, NY, US, type collection of P. 

subtriloba); 16 km N ofCarrasco, 1500 m, 31 Oct 1984 

(9), Solomon et al. 12655 (BG). PANDO: Mukden, Jun- 

Dec 1979 (9), Izawa 9 (U); Prov. N. Suarez, 20 Aug 

1978 (9), Mences 759 (MG); W bank of Rio Madeira, 

nr. Abufia, 22 Jul 1970 (9), Prance et al. 6253 (B, INPA, 

NY, S, U, UC, US), 19 Nov 1968 (9), Prance et al. 

8670 (F, INPA, MG, NY, S, U); Prov. N. Suarez, San 

Pedro 13 Oct 1977 (9), Terceros 1400 (INPA); Santa 

Barbara, 30 Aug 1902 (9), R. S. Williams 1560 (NY, 

(9), Schwacke 10393 (P). PARA: Serra US, type collection of P. scabra). 

dos Carajas, Serra Norte, ca. 20 km N of Amza Exploration 

camp, 17 Oct 1977 (9), Berg et al. 598 (MG, Vernacular names. Colombia. Meta: papa- 

U); Cuiaba-Santarem rd., km 1183, 2 Feb 1977 (9), quillo. Venezuela. Bolivar: amia-yek; Delta 

Berg et al. 772 (F, MG, MO, RB, S, U); Breves, Oct- Amacuro: chaparro de agua. Guyana. buruma, 

Nov 1957 (a), Pires et al. 6663 (U); Rio Itacaiunas, 

affluent of Rio Tocantins, Serra Buritirama, Jul 1970 sandpaper. Surinam. Manbospapaja. French 

(9), Pires et al. 12570 (IAN); Capanema-Maranhao rd. Guiana. Bois canon, kuluma, kalate, (Wayapi). 

BR-22, km 96, 29 Oct 1965 (9), Prance et al. 1777 (F, Peru. Amazonas: shuiya (Huambisa), sugkama(t) 

GH, NY, P, S, U, US); Gleba Bacaja, just below mouth shuiya, tsakap suiya (Huambisa); Huanuco: paof 

Rio Bacaja 21 Nov 1980 (st), Prance et al. 26375 

(U); Santar6m, km 70 Estrada do 

paya del monte; Loreto: uvilla; San Martin: uvil- 

Palhao, Ramal do 

Caetetu, 16 Sep 1969 (a), M. Silva et al. 2623 (G, NY, la, uvilla blanca. Brazil. Bahia: itararanga, tara- 

S, U, US). PARANA: P6rto de la Cima, 17 Jul 1911 (st), ranga blanca; Maranhao: kaymbe'y (Ka'apor); 

Dusen 11942 (S); Jacarehy (=Sao Joao), 8 Oct 1915 Mato Grosso; imbaubarana, imbauba-torem; 

(st), Duskn 17239 (F, NY, S), 22 Nov 1915 (9 and 6), Para: amapati or mapati; Parana: imbaubarana, 

Dusen' 17345 (F, GH, NY, S); Morretes, 1904 (st), 

Dusen s.n. (S); 14 Aug 1911 (st), Dusen s.n. pau de jacu; Pernambuco: embauba da mata. 

(F, GH, 

NY, S); Mun. Guaratuba, Pedra Branca do Araraquara, 

This subspecies is very variable in the density 

3 Nov 1960 (9), Hatschbach 7411 (R, U, US), 3 Nov and length of the indument. The material from 

1960 (6), Hatschbach 7466 (US); Mun. Guarataba, Rio eastern Brazil often has 5-fid leaves, the indu-


ment is rather short, and the leaf base is deeply tinct forms of the subspecies in the same area 

to shallowly cordate. Some collections from the and the absence of intermediates between these 

southern part of the Amazon Basin (Bolivia and forms. 

adjacent parts of Brazil) usually have 5-7-fid to 

-parted leaves, while elsewhere in the Amazon 

Basin the leaves are 3-lobed to -fid or entire. 

Most collections from the Guiana region and the 

Lower Amazon Basin have 3-fid leaves with a 

shallowly cordate to truncate leaf base or entire 

leaves; the indument is short, and the brown, 

pluricellular hairs on the twigs are usually dense. 

Material from the Upper Amazon Basin is 

much more variable. Some of the collections have 

almost elliptic (to oblong) entire leaves. Another 

group of collections have entire leaves (often with 

relatively short petioles) or 3-fid to -parted, 

sometimes 5-parted leaves with the lobes rather 

close together, due to a smaller angle in which 

the main basal lateral veins depart from the midrib. 

These collections often have sparse pluricellular 

hairs on the leafy twigs, and those with 

entire leaves resemble P. velutina, from which 

they differ in a greater number of lateral veins. 

A third group of specimens have 3-fid to -parted 

leaves with a deeply cordate leaf base, often with 

overlapping lobes. Several of these collections 

have relatively long yellow hairs on the leafy 

twigs and stipules, or have a rather densely hirtellous 

upper leaf surface. The collections with 

long, yellow hairs usually have stipules in which 

the inner surface is rather densely hairy, unlike 

the other collections of subsp. guianensis which 

have the inner surface of the stipules glabrous or 

sometimes with very sparse, white hairs. Three 

collections (Croat 20616, Krukoff4817 and 8369) 

have 3-fid, deeply cordate leaves and long, yellow 

hairs on various parts, including the inner surface 

of the stipules and more numerous lateral veins 

(up to 25 pairs), in medium-sized leaves less than 

1 cm from each other. Other collections ofsubsp. 

guianensis (and collections of subsp. venezuelensis) 

have the lateral veins more than 1 cm from 

each other. 

The material examined does not provide clear 

indications about the nature of the various forms 

within the subspecies no more than about the 

distribution of these forms. Material received 

from an inventory of a forest in the Upper Rio 

Moa region (Brazil, Acre) by Campbell and collaborators 

indicates the presence of several dis- 

In several specimens from the Upper Amazon 

Basin the leafy twigs, petiole, stipules, and lower 

surface of the lamina are covered by a white 

layer, actually the mycelium of some fungus which 

apparently uses the pluricellular hairs as a substrate. 

lb. Pourouma guianensis Aublet subsp. venezuelensis 

(Cuatrecasas) C. C. Berg & van Heusden, 

Proc. Kon. Ned. Akad. Wetensch., Ser. 

C. 91(2): 106. 1988. Fig. 61. 

Pourouma venezuelensis Cuatrecasas, Bol. Soc. Venez. 

Ci. Nat. 15: 107. 1954. Type. Venezuela. Aragua: 

Parque Nacional Henri Pittier, 20 Nov 1953 (6), 

Aristeguieta 2005 (holotype, F, not seen; isotype, 

VEN). 

Leafy twigs yellow-hirsute, brown, pluricellular 

hairs dense. Lamina 3-5-fid to -parted, base 

? deeply cordate; lateral veins 16-26 pairs; stipules 

outside yellow-hirsute, inside with yellow 

hairs. Fruiting perianth 2-2.5 cm long. 

Distribution (Fig. 60). Northern Venezuela 

(Aragua, Carabobo, Miranda, and Yaracuy), 

coastal mountains; in forest at altitudes between 

ca. 1000 and 1200 m. 

Specimens examined. VENEZUELA. ARAGUA: Parque 

Nacional "H. Pittier," 20 Nov 1953 (6), Aristeguieta 

2005 (VEN, type collection of P. venezuelensis), 

12 Mar 1964 (6), Ijjdsz-Madriz 28 (VEN), 24 Dec 1946 

(9), Lasser 2202 (VEN); Rancho Grande, 30 Dec 1946 

(6), Lasser 2267 (VEN); Guamitas, 5 Nov 1947 (6), 

Pittier et al. 15641 (US, VEN); Parque Nacional "H. 

Pittier," 6 Jan 1968 (st), R. F. Smith V-3321 (VEN); 

Chorni, 39 Apr-1 May 1972 (2), Steyermark et al. 

105863 (NY, VEN); Agua Amarilla, 12 Nov 1947 (6), 

Tamayo 3370 (VEN), Rancho Grande, 26 Mar 1938 

(9), LI. Williams 9983 (A, F, GH, NY, US, VEN). 

CARABOBO: Between Valencia and Campanero, 7 Mar 

1860 (2), Fendler 2420 (G); Cumbre de Valencia, Pt. 

Cabello, without date (6), Karsten s.n. (LE); E of Los 

Tangues, S of Borburata, 31 Mar 1966 (9). Steyermark 

et al. 95378 (NY, VEN, U). MIRANDA: Guatopo, 19 

Nov 1956 (9), Bernardi 5696 (VEN), Bernardi s.n. (NY); 

San Juan, 18 km SW of Cupira, 2-7 Sep 1977 (6), 

Gonzalez et al. 1319 (BG); S of Santa Cruz, Mar 1978 

(2), Steyermark et al. 116884 (BG); 11 km SSE of El 

Guapo, 27-28 Mar 1978 (2), Steyermark et al. 116966 

(U). YARACUY: N of Salom, 4 Mar 1982 (st), Liesner 

et al. 12394 (U).

Pourouma 

~ I I 

MINISTERIO DE AGRICULTUPA V CRIA 

IERDBAJiO NACIONAL DF VENEZUFLA 

FIG. 61. Pourouma guanenss Abl subsp. Leafy t wig with pistillate inflorescences (Steyermark et 

ial. 105863).sden tr: a 

D Ce.: C,C, Blrg 

E,C.I. van in- He usden 

Utrecht r'. 

...X'lr , A. R I%, ? %t 

,,,. J.&M 

a. 

Cm"t 

FIG. 61. Pourouma guianensis subsp. venezuelensis. Leafy twig with pistillate inflorescences (Steyermark et 

al. 105863). 

Vernacular names. Venezuela. Carabobo: tambor; 

Miranda: yagrumo negro. 

This subspecies is very uniform compared with 

subsp. guianensis. Some collections of the latter 

subspecies from the Upper Amazon Basin resemble 

subsp. venezuelensis, but are distin- 

129 

guished by smaller leaves and smaller fruiting 

perianths. 

2. Pourouma 

velutina Martius ex Miquel in Mar- 

tius, Fl. bras. 4(1): 130, t. 41. 1853; Berg, Fl. 

Suriname 5(1): 269. 1975. Type. Brazil. Para:

130 


Mouth of Rio Tocantins, Aug 1819 (6), Mar- widened apex; perianth 4-6 mm long, yellowishtius 

s.n. (lectotype, M, chosen here; isolecto- velutinous, at the apex with brown, pluricellular 

type, M). 

Fig. 62. hairs; stigma peltate, 1.5-3 mm diam., with 

sparse, yellow hairs and dense, brown, pluricel- 

Pourouma steyermarkii Standley & Cuatrecasas in lular hairs. Fruiting perianth black, ovoid to el- 

Cuatrecasas, Fieldiana Bot. 28(1): 210. 1951. Type. 

Venezuela. Bolivar: Ptari-tepui, between Rio 

lipsoid or sometimes to 

Karuai 

oblongoid, 1.2-1.8 x 

and first ridge above Rio Karuai, 28 Nov 1944 

0.6-1 

(v), 

cm, densely to sparsely hairy. 

Steyermark 60665 (holotype, F; isotype, VEN). Distribution (Fig. 60). Venezuela (Amazonas 

and Bolivar), Surinam, French Guiana, Ama- 

Tree, up to 15 m tall. Leafy twigs 2-6 mm zonian Brazil (Amapa, Amazonas, Mato Grosso, 

thick, brownish- to yellowish- to whitish-seri- and Para) and Peru (Loreto), and eastern Brazil 

ceous to -puberulous and sometimes with brown, (Bahia and Espirito Santo); in non-inundated 

arachnoid hairs. Lamina entire, elliptic to ovate, forest, mostly at low altitudes, in Venezuela up 

rarely to obovate, 6-30 x 2-20 cm, coriaceous to ca. 1200 m. 

to subcoriaceous or sometimes chartaceous, apex 

acuminate, base rounded to acute, sometimes Specimens examined. VENEZUELA. AMAZONAS: 

truncate or 

Atures, 20 

subcordate; upper surface 

Apr 1978 (9), Davidse et al. 15365 (MO, 

scabrous, U); Camani, 24 Apr 1971 (9), Foldats 114-1A (NY, 

the main veins brown-tomentose to -sericeous, VEN). BOLIVAR: Lower S-facing slopes of Ptari-tepui 

sometimes with orange or orange-brown, pluri- between Rio Karuai and first ridge above Rio Karuai, 

cellular hairs; lower surface whitish-sericeous to ca. 1200 m, 28 Nov 1944 (9) Steyermark 60665 (F, 

-velutinous, sometimes with VEN, type collection of P. steyermarkii). 

brown, pluricellular SURINAM. Sectie O, 11 Nov 1915 (st), BW 1368 

hairs, white arachnoid hairs in the areoles and (U); Tibiti savanna, 20 Jan 1949 (st), Lanjouw et al. 

on the smaller veins; lateral veins 7-13 pairs, 1761 (NY, U); Jodensavanne, Mapane Creek, 13 Jun 

basal pair branched or unbranched, tertiary ve- 1953 (st), Lindeman 4057 (U); 1 Oct 1953 (st), Lindenation 

prominent beneath; petiole 1.5-20 cm man 4794 (U); Upper Coppename R., 6 Aug 1954 (st), 

Lindeman 6418 

long, yellow- to brown-sericeous to 

(U). 

-tomentose, FRENCH GUIANA. St. Laurent, 10 Oct 1956 (9), 

and sometimes with brown, pluricellular hairs; BAFOG 7562 (CAY, NY, P, U); Gregoire, 26 Jan 1972 

stipules 2-12 cm long, caducous, outside yellow- (9), Deward 138 (CAY, P, U); Maroni R., 1891 (6), 

sericeous to -velutinous, and often with brown Gandoger 19 (P); Orapu, 26 Oct 1970 (9), Oldeman 

B.3786A 

to yellow, pluricellular hairs, inside (A, CAY, P, U); Acarouany, 1858 ($), 

yellow-se- 

Sagot 

1163 (BR, F, G, GH, GOET, NY, P, S, U); Godebert, 

riceous to -velutinous. Staminate inflorescences 20 Dec 1920 (6), Wachenheim 271 (P). 

up to 12 cm long and 8 cm wide; peduncle 2-7 PERU. LORETo: Rio Nanay, trail from Astoria to 

cm long, peduncle and branches yellowish-seri- Rio Mazan, 9 Jan 1976 (9), McDaniel et al. 20403 

ceous to -velutinous or to -tomentose, and often (NA). 

BRAZIL. AMAPA: Mun. 

with Mazagao, 75-80 km WSW 

dense, brown, pluricellular hairs; flowers of Macapa, 5-10 km SW of Rio Prato, 20 Dec 1984 

sessile, diffusely distributed along the ultimate (9), Daly et al. 3936 (BG); Clevelandia, 2 Aug 1960 (6), 

branches; tepals 1.3-1.8 mm long, free or basally Westra 47303 (FHO, GH, MG, NY, U, UC, US). 

connate, white-puberulous, filaments shorter than AMAZONAS: Manaus, 9 Aug 1962 (6), Chagas (INPA) 

the perianth, free. Pistillate inflorescences in fruit 

1655 (INPA, U); Rio Uatuma, Itapiranga, 27 Aug 1979 

(9), Cid 849 

up to 12 cm long and 7 

(U); Manaus, 8 Jul 1933 (9 and 6), Ducke 

cm wide; peduncle 2.5- 

(RB) 25247 (INPA, RB); Manaus-Itacoatiara rd., km 

7 cm long, peduncle and branches yellowish- to 26, 4 Jan 1977 (st), Nascimento 262 (INPA); track from 

brownish-sericeous to -velutinous, sometimes to Boca de Acre airstrip to Monte Verde, 21 Sep 1966 

-tomentose or to -puberulous, and with dense, (9), Prance et al. 2469 (INPA, MO, NY, R, S, U, UC, 

brown, pluricellular hairs; flowers 2-8, + dis- US); Manaus, Aleixo rd., km 11, 2 May 1974 (6), Prance 

et al. 21011 (INPA, MG, MO, NY, S, U, US); Manaus, 

tinctly in two clusters; pedicel up to 0.4 cm long, Binda Creek rd., 26 Jun 1961 (9 and 6), Rodrigues et 

in fruit up to 1 cm, already at anthesis with a al. 2892 (INPA as 2092, MG, RB, U); Manaus Pas- 

FIG. 62. Pourouma velutina. From Martius, Flora Brasiliensis 4(1). 1853: tab. 41. Leafy twig with staminate 

inflorescences, diagram of staminate flower (D), part of staminate inflorescence (28), staminate flower (1), tepals 

(4), stamens (7). 

--9


'~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~r 

'~i ' 

" 

,?il:~~~~~~~~~~ 

, ^ > 3 

\ 

POUROUMA vebl iainas.


sarinho rd., 14 Sep 1961 (9), Rodrigues et al. 3267 (U), 4(1): 129, t. 39. 1853. Type. Colombia. Ama- 

9 Aug 1962 (2), Rodrigues et al. 4585 (U). BAHIA: Road zonas: Rio 

from Rod. Sao Jose-Buerarena to Una, 7 Jul 1980 Caqueta, Puerto Miraia, Jan 1820 

(juv), 

Berg et al. 1143 (U); Una, 29 Aug 1978 (2), Mori et al. (Q), Martius s.n. (holotype, M; isotype, U). 

11026 (MG, RB, U); between Vit6ria and Salvador, 

without date (2), Sello s.n. (B). ESPIRITO SANTO: Rio Tree, up to 15 m tall. Leafy twigs 3-15 mm 

Doce, Velho-Pancas rd., 20 Sep 1930 (6), M. Kuhlmann thick, yellowish- to whitish-puberulous to 

383 (RB); Linhares, 11 Mar 1972 (st), Sucre 8684 (U). -(sub)sericeous, or yellow-hirsute, and with sparse 

MATO GROSSO: Aripuana, 28 Mar 1977 (st), Gomes et to dense, brown, pluricellular hairs. Lamina enal. 

1021 (INPA), 12 Apr 1977 (st), Gomes et al. 1196 

(INPA), without date (9), Rylands 15 (INPA); Mun. 

tire, ovate to subovate or elliptic to oblong, 5- 

Sinop, 6-9 km E of BR-163, on rd. to Fazenda Lon- 25 x 3-18 cm, 3-5-lobed to -parted, or 5-9drina, 

24 Sep 1985 (9), Thomas et al. 4018 (BG); Mun. parted, 13-30(-42) x 13-30(-40) cm, coriaceous 

Sinop-Colider, BR-080, ca. 91 km E of junction with to chartaceous, apex acuminate (to obtuse), base 

BR-163, Serra Formosa, 3 Oct 1985 (9), Thomas et al. (obtuse to) truncate to deeply cordate; upper sur- 

4179 (BG); Mun. Vila Bela Santissima Trinidade, 4 

km 

face 

S of border ofRond6nia, 3 

scabrous to 

Nov 1985 (9), Thomas 

scabridulous or sometimes 

et al. 4810 (BG). PARA: Belem, 2 Jul 1914 (8), Ducke smooth, hairy on the main veins, puberulous on 

(MG) 15350 (G, MG, P, US); Belem, 18 Jul 1899 (2), the smaller veins, lower surface smooth, some- 

Huber (MG) 1640 (G, MG, P, S, U); Gurupa (2), Ducke times scabridulous, yellowish-appressed-puber- 

(MG) 15935 (MG); Rio Xingfi, Vit6ria-Ponte Nova ulous on the main 

rd., 7 Aug 1918 (9), Ducke (MG) 17167 (MG); B6a 

veins, arachnoid hairs in the 

Vista, 15 Oct 1897 (2), Guedes (MG) 1235 areoles and on the 

(G, MG, 

smaller 

P, 

veins, or confined to 

S, U); Bel6m-Brasilia rd., km 93, 19 Aug 1959 (6), M. the areoles; lateral veins (6-)12-25(-40) pairs, 

Kuhlmann et al. 52 (GUA, MG, R, S, US); mouth of basal pair branched, tertiary venation almost 

Rio Tocantins, Aug 1819 (8), Martius s.n. (M, lectotype plane beneath; petiole 

collection of P. 

(2-)4-30(-46) cm 

velutina); Jacuarary (=Jaguarari), 21 

long, 

Aug 1819 (8), Martius s.n. (M); Bel6m, 12 Aug 1945 appressed-puberulous to -strigose, sometimes 

(9), Pires et al. 161 (RB); Pirelli, Jul 1958 (a), Pires yellowish-hirtellous to -(sub)tomentose or yel- 

7015 (U); Belem, Sep-Oct 1961 (9), Pires 51882 (NY, low-hirsute; stipules 2-13.5(-25) cm long, out- 

U, US); Ilha de Breu, 5 Oct 1965 (9), Prance et al. 1548 side yellowish- to whitish-appressed-puberulous 

(F, GH, NY, P, S, U, US); Cuiaba-Santarem rd., km to 

163, 10 Nov 1977 (2), Prance et al. 25167 

-(sub)sericeous or sometimes yellowish-hir- 

(F, INPA, 

MO, RB, U), km 941, 13 Nov 1977 (9), Prance et al. tellous to -(sub)tomentose or yellow-hirsute, in- 

25363 (MO, RB, S, U, US); Santar6m-Palhao rd., km side ? densely hairy, sometimes glabrous, ca- 

35, 28 Aug 1969 (9), M. Silva et al. 2423 (F, GH, MG, ducous. Staminate inflorescences up to 17 cm 

NY, S, U, US). 

long and 14 cm wide; peduncle 2.5-7 cm long, 

yellow-puberulous and with dense (most dense 

Vernacular names. Venezuela. Amazonas: cu- on the branches), dark-brown or red-brown, 

cura. Surinam. Boroma or boroma ibeberobana 

pluricellular hairs;flowers usually sessile, ? clus- 

(Arawak), bospapaja, yarayara (Carib). French 

tered, but not in (sub)globose heads; tepals 1-2 

Guiana. Bois canon, papaye apici (Paramaka). mm long, free or basally connate, (sparsely) pu- 

Brazil. Amazonas and Mato Grosso: imbafibaraberulous; 

filaments shorter than the tepals. Pisna; 

Para: mapatirana. 

tillate inflorescences in fruit up to 26 cm long and 

This species can sometimes be very difficult 14 cm wide; peduncle 1.5-14 cm long peduncle 

to distinguish from forms ofP. guianensis subsp. and branches with indument similar to that of 

guianensis with entire leaves and sparse pluri- the staminate inflorescences; flowers 6-40; pedcellular 

hairs on the leafy twigs. 

icel 0.2-0.5 cm long, in fruit 0.4-1 cm long; peri- 

The lectotype is chosen from the two (syntype) anth 2-4 mm long, scabrous; stigma peltate, 1-2 

collections of Martius cited above. 

mm in diam. Fruiting perianth purple to black, 

ovoid to ellipsoid, ca. 1.5 cm long, scabrous (or 

3. Pourouma bicolor Martius, Syst. mat. med. smooth), sparsely hairy. 

bras. 34. 1843; Miquel in Martius, Fl. bras. In P. bicolor five subspecies can be recognized. 

Key to the Subspecies of Pourouma bicolor 

1. Stipules glabrous inside. ........................................................ . subsp. tessm annii. 

1. Stipules hairy inside. 

2. Arachnoid hairs on the lower leaf surface confined to the areoles or almost so.


3. Smaller veins (reticulum) of the lamina beneath plane or almost so, the veinlets rather thick and 

the spots of arachnoid indument distinctly separated; lamina mostly entire or 3-lobed; Upper 

Amazon Basin to northern Colombia. ...................................... a. subsp. bicolor. 

3. Smaller veins (reticulum) of the lamina beneath more or less prominent, the spots of arachnoid 

indument mostly not distinctly separated; lamina mostly 5-7-parted; Central America to the 

Pacific coastal region of Colombia and Ecuador and to northwestern Venezuela (also in Amazonian 

Peru?). .................................... ....................... d. subsp. scobina. 

2. Arachnoid hairs on the lower leaf surface also covering the smaller veins, usually covering the area 

between the parallel tertiary veins. 

4. Lamina usually 5-7-parted; pistillate inflorescences often with up to 50 flowers. 

5. Stipules subpersistent; segments ofthe lamina relatively broad; leafy twigs often yellow-hirsute; 

Panama and western Colombia. ....................................... e. subsp. chocoana. 

5. Stipules caducous; segments of the lamina relatively narrow; leafy twigs never hirsute; Guianas 

and the Lower Amazon Basin (to southern Venezuela?). ................... c. subsp. digitata. 

4. Lamina usually entire or 3-lobed; pistillate inflorescences mostly with up to 25(-35) flowers; 

Guiana region, Amazon Basin to northern Colombia. ........................ a. subsp. bicolor. 

3a. Pourouma bicolor Martius subsp. bicolor. 

Fig. 63. 

Pourouma aspera Tr6cul, Ann. Sci. Nat. Bot., Ser. 3, 

8: 102. 1847; Standley, Publ. Field Mus. Nat. Hist., 

Bot. Ser., 18(1): 391. 1937; Standley & Steyermark, 

Fieldiana Bot. 24(4): 52.1946. Type. French Guiana. 

Without locality, without date (9), Poiteau s.n. (ho- 

lotype, P; isotypes, LE, P). 

Pourouma crassivenosa Mildbraed, Notizbl. Bot. Gart. 

Berlin-Dahlem 10:419.1928. Type. Bolivia. La Paz: 

Nr. Mapiri, Sep 1907 (2), Buchtien 2050 (holotype, 

US; isotypes, B, NY). 

Pourouma lawrancei Standley, Publ. Field. Mus. Nat. 

Hist., Bot. Ser., 17: 183. 1937. Type. Colombia. Bo- 

yaca: El Humbo, 31 Mar 1933 (2), Lawrance 727 

Specimens examined. COLOMBIA. Without locality, 

without date (2), Triana 861 (E, P). AMAZONAS: 

Rio Caqueta, Puerto Mirafia, 1820 (2), Martius s.n. (M, 

U type collections of P. bicolor) (M, U). 

AMAZONAS-VAUPES: Rio Apaporis, Jinogoj6, 25 Sep 

1952 (2), Schultes & Cabrera 17615 (F, type collection 

*of P. schultesii). ANTIOQUIA: Confluence of Rios Ite 

and Tamar into Rio Cimatarra, ca. 38 km W of Bar- 

rancabermeja, 24 Nov 1967 (6), Bruijn 1497 (F, M, 

MO, NY, S, U, US, VEN), 26 Nov 1967 (6), Bruijn 

1514 (F, M, MO, NY, S, U, US, VEN), (st), Bruijn 

1516 (F, M, MO, NY, S, U, US, VEN), (8), Bruijnl517 

(F, M, MO, NY, S, U, US, VEN), 27 Nov 1967 (6), 

Bruijn 1526 (F, MO, NY, S, U, US, VEN). BOYACA: 

El Humbo, 31 Mar 1933 (6), Lawrance 727 (A, E, F, 

G, MO, S, UC, type collection of P. lawrancei). META: 

(holotype, F; isotypes, A, E, G, MO, S, UC). Footpath between Rio Giiejar and Cana Guapayita, 

Pourouma schultesii Cuatrecasas, Caldasia 7: 303. 1956. Cafno Yerli, 20-28 Dec 1950 (2), Idrobo et al. 786 (F, 

Type. Colombia. Amazonas-Vaup6s: Rio Apaporis, MO, NY, US); Villavicencio (2), Karsten s.n. (LE); Sier- 

Jinogoje, 25 Sep 1952 (2), Schultes & Cabrera 17615 ra de la Macarena, Cafno Ciervo, 12 Jan 1950 (2), Phil- 

(holotype, F). 

ipson et al. 2086 (S, US). PUTUMAYO: Santa Rosa on 

Pourouma camaratana Cuatrecasas, Acta Bot. Venez. Rio Guamfies, 26 Nov 1966 (2), Pinkley 556 (S). 

2(5-8): 202.1967. Type. Venezuela. Bolivar: Auyan- SANTANDER: Region Carare-Op6n (Capote), nr. Rio 

tepui, above valley of Camaratana, 18 May 1964 (2), Magdalena, 6 Jul 1968 (9), Garcia et al. 9 (US). Vi- 

Steyermark 94810 (holotype, US; isotypes, NY, CHADA: 3 km S ofLinea Roja, Parque Nacional Naturel 

VEN). 

"El Tuparro," 14 Mar 1985 (2), Zarucchi et al. 3720 

(BG). 

Leafy twigs appressed-puberulous to strigose. VENEZUELA. AMAZONAS: Reserva Forestal "El 

Lamina entire to 3-lobed, sometimes 3-fid, oc- Sipapo," Rio Sipapo, May 1971 (2), Blanco 1149 (NY, 

casionally (in subjuvenile material?) 5-parted; US, VEN); Rio Marawinuma, nr. Cerro de La Neblina 

Base 

upper surface scabrous, sometimes smooth, low- Camp, 6 Feb 1984 (st), Liesner 15681 (BG), 28 

Feb 1984 (9), Liesner 16299 

er surface 

(BG); Depto. 

smooth, sometimes 

Atures, N 

scabridulous, the side of Rio Cataniapo, 45 km SE of Puerto Ayacucho, 

arachnoid hairs in the areoles and on the smaller 13 May 1980 (2), Steyermark et al. 122390 (U); Depto. 

veins or confined to the areoles, but then the Atabapo, Rio Cunucunuma, between Cerro Duida and 

smaller veins relatively thick and the spots of 

Cerro Huachamacahi, Jan-Feb 1982 (9), Steyermark 

et al. 12585 

arachnoid hairs 

(BG); Rio 

clearly separated; stipules inside 

Marawinuma, nr. Cerro de La 

Neblina Base Camp, 5 May 1984 (6), Thomas 3370 

glabrous, caducous. Pistillate inflorescences (BG). BOLIVAR: Rio Chizca, Uriman, 5-8 Sep 1953 

mostly with 3-25 flowers. 

(st), Bernardi 910 (NY, VEN); Rio Hacha, 28 Dec 1955 

Distribution (Fig. 68). Throughout the Ama- (2), Bernardi 2657 (G, NY,); slopes ofQuebrada O-pozon 

Basin and the Guiana region, extending to 

ru-ma, between Santa Teresita de Kavanayen and Rio 

Pacairao (tributary of Rio 

northern Colombia; in 

Mouak), 20-21 Nov 1944 

non-inundated forest at 

(st), Steyermark 60401 (F); lower SW slopes of Chilow 

altitudes, in Bolivia up to ca. 900 m. manta-tepui (Torono-tepui), nr. Rio Tirica, 24 May


!ie. 

~.~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~:' 

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r-? *~~~. .. 

z 

' ~ ~ ~ 

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i ~ 

~~~~~~~~~~~~~~~~~~~~~~~~~~~~~ 

.. . . : ,,- 

ii!:..-~~ 

~ 

-P~~~~~~~~~~~~~~~~~~,~ 

,..~ 

:,, , 

POURi~A biol. 

?f 

'. 

. ~~~~~~~~


1953 (st), Steyermark 75540 (F, NY, VEN); Sierra de Reserva Florestal Ducke, Nov 1972 (st), Rodrigues 9206 

Lema, lowland between Rio Chicanan and Rio Ay- (INPA). MATO GROSSO: Aripuana, 21 Feb-2 Aug 1977 

aiche, between Puerta Lema and base of the Sierra, 24 (2 and st), Gomes et al 676, 786, 843, 966, 979, 1052, 

Aug 1961 (9), Steyermark 89457 (NY, VEN); frontier 1072, 1079, 1088, 1123, 1176, 2399 (INPA); Ari- 

Venezuela-Brazil, NE ofSerrania Pia-soi, 5-6 Jan 1962 puana, nr. Humboldt Centre, rd. to Rio Juruena, 8 Oct 

(st), Steyermark 90644 (VEN); Auyan-tepui, above 1973 (6), Prance et al. 18250 (INPA, NY, U), 30 Nov 

valley of Camarata, alt. 1500-1530 m, 18 May 1964 1978 (2), Rylands 52 (INPA). PARA: Rios Mojfi and 

(9), Steyermark 94180 (NY, US, VEN, type collection Acara, S of Belem, ca. 7 km N of Mojfi, 3 Jun 1969 

of P. camaratana); Rio Caura, nr. Salto Para, 15-17 (6), Austin et al. 4140 (MO); Faro, 6 Jan 1920 (2), Ducke 

Jan 1977 (9), Steyermark et al. 113096 (U). 

(RB) 13057 (RB); between Rio Pacaja and Rio Muira- 

SURINAM. Forest Reserve Zanderij 1, 24 Oct 1917 piranga, SW of Ilha de Breu, 20 Sep 1965 (2), Prance 

(9), BW3376 (MO, U); Watramiri, 8 Oct 1918 (9), BW et al. 1397 (NY, US); BR-22, km 64, 25 Aug 1964 (6), 

4033 (NY, U), 13 Dec 1920 (9), BW 5036 (U); Sectie Prance et al. 58862 (F, GH, NY, S, U, US); Belem 

0, Feb 1943 (9), Stahel (Woodherb. Sur.) 166 (A, F, (IAN) 17 Oct 1963 (2), N. T. Silva 57832 (NY, U, US). 

NY, U, UC). 

RONDONIA: Forte Principe da Beira, Estrada do Igarape 

FRENCH GUIANA. S of Saul, 12 Feb (9), Leeu- da Viiva, 5 Jan 1962 (2), Rodrigues et a. 4237 (U). 

wenberg 11784 (CAY); without locality, (9), Poiteau RORAIMA: Indian trail from Surucucu W to Uaica bes.n. 

(LE, P, type collection of P. aspera). 

tween Botamatedi and Maitf, 10 Feb 1971 (2), Prance 

ECUADOR. NAPO: 6 km SE of Los Sachas, 14 Apr et al. 13582A (INPA, F, GH, M, NY, P, S, U). 

1985 (9), Baker 6017 (BG); 5 km N of Coca, on Coca- BOLIVIA. LA PAZ: Nr. Mapiri, Sep 1907 (2), Buch- 

Lago Agrio rd., 16 Mar 1980 (st), Brandbyge et al. tien 2050 (B, NY, US, type collection of P. crassiveno- 

30194 (AAU); El Chuncho, Estaci6n INIAP, 2 Oct sa), Sep 1907 (6), Buchtien 2122 (NY, US), 12-30 Sep 

1987 (st), Palacios 2059 (BG). 

1939 (9), Krukoff10867 (A, F, G, MO, NY, S, U, UC, 

PERU. AMAZONAS: Rio Santiago, nr. Caterpiza, 14 US), Sep 1939 (8), Krukoff 10874 (F, NY), 8 Oct-15 

Nov 1979 (9), Huashikat 1241 (U), 28 Nov 1979 (2), Nov 1939 (2), Krukoff 11254 (F, NY). 

Junqui 149 (U). LORETO: Rio Mom6n, 4 Sep 1972 (9), 

Croat 19997 (MO); Maynas, Yaguasyacu, tributary of Vernacular names. Colombia. Antioquia-Bo- 

Rio Ampiyacu, below Borro Indian village of Brilla livar: cirpe (or sirpe) hembra, 

Nueva, 7 

cirpe (or 

Nov 1977 (9), Gentry et al. 20356 

sirpe) 

(MO); Rio 

Ucayali, Jenaro Herrera, 7 Dec 1977 (9), Gentry et al. macho; Boyaca: cormi; Meta: caimar6n de mico. 

21193 (MO, U); Rio Javari, between Rio Curaqa R. Putumayo: otsepacho tsaha (Kofan). Venezuela. 

and Tambaqui, 28 Oct 1976 (9), Prance et al. 24188 Amazonas: cucura or cucure, sadha'fhi; Bolivar: 

(INPA, MG, U). 

cay-bari-cay, cay-i-wari-cay-yek, sarasara. Suri- 

BRAZIL. Without locality (9), Burchel 9792 (GH, nam. Boroma 

LE, P, under the same number P. mollis (Arawak), pourouma or 

subsp. 

puruma 

mollis). 

ACRE: Upper Rio Moa, Fazenda Arizona, 10-16 Oct (Carib). Peru. Amazonas: paui shuina, tsakap sui- 

1985 (st), Campbell et al. 6650 (BG); Rio Jurua, nr. ya (Huambisa). Brazil. Acre: imbafbarana. 

Cruzeiro do Sul, 22 Oct 1966 (9), Prance et al. 2754 Amazonas: imbafiba; Mato Grosso: imbafibar- 

(INPA, MG, U). AMAPA: Rio Oiapoque, 1-3 km N of 

ana, 

Cachoeira Tres Saltos, 12 Sep 1960 (6), Irwin 

tamaoquare; Para: 

et al. 

garguaba, mapatirana. 

48192 (GH, M, NY, P, U, US). AMAZONAS: 27 

Bolivia. La Paz: uva menueda. 

Sep 

1973 (8), Berg et al. P. 18152 (INPA, MO, NY, P, U); In and towards the eastern part of the species 

Manaus-Caracarai rd., km 148, 26 Sep 1973 (6), Bisby range (from Colombia to Bolivia) the arachnoid 

et al. P.18118 (F, INPA, MO, NY, S, U, US); Rio hairs on the lower leaf surface are confined to the 

Negro, Uaupes, 22 Jan 1960 (9), Cavalcante 777 (MG); areoles in 

Serra de Neblina, Rio Cauaburi, Camp Tucano, 5 

combination 

Dec 

with an increase of the 

1965 (9), Maguire et al. 60333 (F, MG, NY, P, U, US); 

thickness of the smaller veins, causing distinct 

Benjamin Constant, 16 Oct 1956 (9), Meyer Drees 9 separation of the "islets" of arachnoid indument. 

(INPA, U); Manaus, Reserva Florestal Ducke, 6 Dec The transition of the normal form of subsp. bi- 

1976 (2), Nascimento et al. (INPA) 66338) (INPA); Rio 

color, with the arachnoid indument 

Negro, Uaup6s, 1 May 1947 (9), Pires 502 (NY); Rio 

extending 

Negro, Sao Gabriel, 1 May 1947 (6), Pires 587 

over the 

(NY); relatively thin smaller veins to the form 

Tapuruquara, rd. to airport, 17 Oct 1917 (9), Prance de scribed above, is rather gradual. Both forms 

et al. 15366 (INPA, M, MO, NY, S, U, US); Manaus, may occur in the same region. In the eastern part 

FIG. 63. Pourouma bicolor subsp. bicolor. From Martius, Flora Brasiliensis, 4(1). 1853: tab. 39, Leafy twig 

with fruiting pistillate inflorescence, fruiting perianth (17), fruiting perianth and fruit (1711), pericarp (43), fruit 

(19), pericarp and seed (21), seed (19).


A 3 

?3X3l cm 

t. '- ? 

FG. 64. Pourouma bicolor subsp. tessmanni. Leafy twigs with pistillate 94).'ur inflorescence (Tunqui 

(! 

L 

..' 

i-" 

i. _ ....... 

' 

.':..


of the species range the upper leaf surface is often 

smooth. See unnamed collection #1 (p. 193). 

3b. Pourouma bicolor Martius subsp. tessmannii 

(Mildbraed) C. C. Berg & van Heusden, Proc. 

Kon. Ned. Akad. Wetensch., Ser. C 91(2): 106. 

1988. Fig. 64 

Pourouma tessmannii Mildbraed, Notzibl. Bot. Gart. 

Berlin-Dahlem 10: 192. 1927. Type. Peru. Loreto: 

Pongo de Manseriche, 8 Oct 1924 (6), Tessmann 

4236 (holotype, B; isotypes, NY, S, US). 

Leafy twigs appressed-puberulous to strigose. 

Lamina entire or occasionally 3-lobed to -fid; 

upper surface scabrous, lower surface smooth, 

appressed hairs on the main veins, arachnoid 

hairs in the areoles and on the smaller veins; 

smaller veins (almost) plane; stipules outside with 

appressed hairs, inside glabrous, caducous. Pistillate 

inflorescences mostly with 3-16 flowers. 

Distribution (Fig. 68). Peru (Amazonas and 

Loreto); in non-inundated forest. 

Specimens examined. PERU. AMAZONAS: Rio Santiago, 

65 km N of Teniente Pinglo, Quebrada Caterpiza, 

13 Oct 1979 (2), Huashikat 912 (U), 30 Jan 1980 

(9), Huashikat 1872 (U); Rio Santiago, nr. La Paz, 12 

Nov 1979 (2), Tunqui 7 (BG); Rio Santiago, Quebrada 

Caterpiza, 17 Nov 1979 (6 and 2), Tunqui 94 (U). 

HuANuco: Prov. Leoncio Prado, 80 km NE of Tingo 

Maria, nr. La Divisoria, 18 Jan 1976 (2), Gentry et al. 

1604 (DUKE, MO, NY). LORETO: 22 km S of km 86 

on Pucallpa-Tingo Maria hwy., 11 Feb 1981 (2), Gentry 

et al. 31197 (U); Pongo de Manseriche, nr. mouth of 

Rio Santiago, 8 Oct 1924 (d), Tessmann 4236 (B, NY, 

S, US, type collection of P. tessmannii). 

Vernacular names. Peru. Amazonas: shuiya 

(Huambisa), tanta shuiya, tentar shuina (Huambisa). 

This subspecies is similar to subsp. bicolor in 

most features but the stipules are glabrous inside. 

3c. Pourouma bicolor Martius subsp. digitata 

(Trecul) C. C. Berg & van Heusden, Proc. Kon. 

Ned. Akad. Wetensch., Ser. C 91(2): 106. 

1988. Fig. 65. 

Pourouma digitata Tr6cul, Ann. Sci. Nat. Bot., S6r. 3, 

8: 106. 1847; Miquel in Martius, Fl. bras. 4(1): 124. 

1853. Type. French Guiana. Without locality, with- 

out date (9), Poiteau s.n. (lectotype P, chosen here; 

isolectotype B). 

Leafy twigs appressed-puberulous to strigose. 

Lamina 5-7(-9)-parted, midsegment lanceolate 

to obovate, base truncate to shallowly cordate or 

sometimes deeply cordate; upper surface scabrous 

to scabridulous, sometimes smooth, lower 

surface with appressed hairs on the main veins, 

arachnoid hairs in the areoles and on the smaller 

veins; smaller veins (almost) plane: stipules outside 

with appressed hairs, inside hairy, caducous. 

Pistillate inflorescences mostly with 10-50 flowers. 

Distribution (Fig. 68). Guyana, Surinam, 

French Guiana, and Brazil (Amapa and Parf), 

and probably in southeastern Venezuela; in noninundated 

forest at low altitudes. 

Specimens exmained. VENEZUELA. AMAZONAS: 

Nr. Cerro de La Neblina Base Camp, 19 Apr 1984 (st), 

Gentry et al 46752 (BG). 

GUYANA. NW slopes of Kanuku Mts., drainage of 

Moku-Moku Creek, Takutu tributary, 31 Mar-16 Apr 

1938 (2), A. C. Smith 3564 (B, F, LE, MO, NY, P, S, 

U, US). 

SURINAM. Watramiri, 9 Jun 1916 (st), BW 2030 

(U); Kaboeri, 30 Oct 1920, B W 4866 (U); Brownsberg, 

4 Oct 1923 (S), BW 6292 (U); Distr. Brokopondo, 2 

km S of Gansee, 27 Apr 1964 (st), Donselaar 1262 (U); 

Nassau Mts., at km 0.2 on terrace of Marowijne R., 

16 Feb 1949 (st), Lanjouw et al. 2213 (F, NY, RB, U); 

Distr. Saramacca, nr. Groningen, 19 Feb 1954 (st), 

Lindeman 5468 (U); Rikanau, nr. Moengo, 6 Jun 1954 

(st), Lindeman 6123 (U); S of Moengotapoe, 14 Jun 

1954 (st), Lindeman 6166 (U); Jodensavanne-Mapane 

Creek area, 10 Dec 1954 (st), Lindeman 6765 (U); 

Distr. Nickerie, area of Kabalebo Dam project, 4 km 

NW of rd., km 39.5, 4 Nov 1981 (st), Lindeman & de 

Roon 750 (U); Lely Mts., E rd. on plateau 1, 29 Sep 

1976 (juv), Lindeman & Stoffers et al. 539 (U), 22 Jan 

1970 (2), Oldenburger et al. 1120 (U); Sipaliwini area, 

Brazilian frontier, 2.5 km S of Sipaliwini R., Feb 1970 

(st), Oldeburger et al. 1406 (U); Winnana Creek, Upper 

Kabouir Creek, Corantyne R., 20 May 1960 (st), Schulz 

(LBB) 8316 (U); Section 0, ca. 5?N, 55?W, Nov 1942 

(a), Stahel 123A (U), Dec 1942 (S), Stahel (Woodherb. 

Sur.) 166B (A, NY, U, UC). 

FRENCH GUIANA: Cayenne, 1 Dec 1955 (2), BAF- 

OG (=Bena) 1050 (CAY, F, U); upper Oyapock R., 

Zidock ville, 6 Sep 1977 (st), Grenand 1442 (CAY, U); 

Saul, 8 Dec. 1982 (2), Mori et al. 15350 (BG); deserted 

village of Eau Claire, about 7 km N of Belizon, 12 Feb 

1966 (9), Oldeman 2030 (CAY); Yaroupi R., 8 km 

downstream of Saut Tainoua, 18 Apr 1970 (st), Oldeman 

3115 (CAY, P, U); without locality, without date 

(9), Poiteau s.n. (B, P, type collection of P. digitata), 

without date (8), Poiteau s.n. (P); Oyapoque R., Camopi, 

8 Mar 1976 (9), Sastre 4350 (P, U); Mt. St. 

Marcel, 28 Mar 1976 (9), Sastre 4579 (P). 

BRAZIL. AMAPA: Serra do Navio, 1961 (2), Aubreville 

144 (P, U, US); rd. to Clevelandia, 14 Oct 1950 

(9), Fr6es 26631 (IAN), 13 Oct 1960 (a), Irwin 48682 

(GH, IAN, NY, U, US); Rio Oiapoque, Mt. Tipac, 16 

Oct 1960 (8), Irwin 48755 (IAN, NY); Rio Oiapoque, 

Mt. Tipac, 16 Oct 1961 (6), Pires et al. 51624 (B, F,


? ..J 

'" - i; . . . . . 

.ru 

_l 

* 

,| x. . . . | |.. | . . 

FIG. 65. Pourouma bicolor subsp. digitata. Leaf and pistillate inflorescences BAFOG 1050). 

FIG. 65. Pourouma bicolor subsp. digitata. Leaf and pistillate inflorescences (BAFOG 1050). 

NY, R, UC, US). PARA: Rio Jari, Monte Dourado, 4 

Feb 1969 (9), Cavalcante 2297 (MG); 25-35 and 65 

km SW of Tucurui, 4 and 18-20 Nov 1981 (9), Daly 

et al. 1208 and 1474 (BG); Rio Jari, Monte Dourado, 

15 Nov 1968 (9), N. T. Silva 1411 (NY, U, US), 15 

Nov 1968 (8), N. T. Silva 1412 (NY, U). 

Vernacular names. Surinam. boroma (Ara- 

wak), bospapaja, pourouma or puruma (Carib), 

yarayara (Carib). French Guiana. a'ilea (Waya- 

pi), male bois canon or bois canon, kulumatelelel 

(Wayapi). Brazil. Para: mapatirana.


This subspecies resembles P. cecropiifolia very 

much. It is therefore somewhat doubtful whether 

Gentry et al. 46752 (from Venezuela, Amazonas) 

belongs to subsp. digitata or represents a juvenile 

form of P. cecropiifolia. Subspecies digitata dif- 

30 Apr 1939 (st), Standley 72667 (F); between Virginia 

and Lago Izabal, 5 Apr 1940 (st), Steyermark 38916 

(F). 

BELIZE. Belmopan, 21 Jun 1973 (6), Croat 24831 

(MO); Stann Creek district, Middlesex, 5 May 1939 

(9), Gentle 2787 (A, F, NA, NY, US); Stann Creek 

valley, 18 Mar 1940 (9), Gentle 3265 (A, NY); Monkey 

R., 18 Oct 1942 (6), Gentle 4211 (DUKE); El Cayo 

district, Hummingbird Hwy., 16 Apr 1955 (9), Gentle 

8675 (F, S, US); Maya Mts., 18 Jun 1930 (9), Schipp 

fers from P. cecropiifolia in the smaller number 

of incisions of the lamina and usually a scabrous 

to scabridulous upper surface of the lamina. 

Nests of small ants are sometimes found in 127 (A, F, G, GH, MO, NY, S). 

between the bases of the main veins. 

HONDURAS. Rio Esperanza, 28 Feb 1903 (6), Wil- 

From the two syntype collections, Leprieurs.n. son 606 (NY, US). ATLANTIDA: San Alejo, 22-27 Apr 

and Poiteau s. n. (staminate specimens), the latter 1947 (st), Standley 7878 (F); Tela, 4 Dec 1927-20 Mar 

1928 

is chosen as lectoptype collection. 

(st), Standley 52900 (A, F, US), (9), Standley 

53951 (F, US). CORTES: Agua Azul, 9 Feb 1952 (9), 

Allen 6401 (F, GH, US). GRACIAS A Dios: Rio Platano, 

17-23 

3d. Pourouma bicolor Martius subsp. scobina May 1973 (9), Clewell et al. 4142 (MO). 

NICARAGUA. ZELAYA: Puerto Cabezas, 2 Dec 1927 

(Benoist) C. C. Berg & van Heusden, Proc. (9), Englesing 50 (F), (st), Englesing 52A (F); between 

Kon. Ned. Akad. Wetensch., Ser. C 91(2): 106. Bluefields and Ginney Point, 24 Mar 1949 (st), Molina 

1988. Fig. 66. R. 1963 (GH); El Recreo-El Pijibaye rd., 11 May 1949 

(st), Standley 19904 (F); rd. to Colonia Manantiales, S 

Pourouma scobina Benoist, Bull. Mus. Hist. Nat. (Paris) of ridge of Serranias de Yolania, 29-31 Oct 1977 (st), 

28: 320. 1922; Woodson & Schery, Ann. Missouri Stevens 4839 (BG); ca. 14.3 km N of El Enpalme, along 

Bot. Gard. 47: 167. 1960. Type: Costa Rica. San new rd. to Mina Nueva America, 27 Apr 1978 (9), 

Jose: Tucurrique, Dec 1898 (i), Tonduz 12930 (ho- Stevens 8323 (BG). 

lotype, P; isotypes, F, GH, M, NY, S, US). 

COSTA RICA. ALAJUELA: Cariblanco, Quebrada 

Pourouma crassivenia Cuatrecasas, Revista Acad. Co- Arrayanes, 1 Jul 1973 (6), Lent 3547 (DUKE, F, MO, 

lomb. Ci. Exact. 9(36/37): 340. 1956. Type. Colom- U). CARTAGO: Turrialba, 20 May 1951 (9), Le6n 3501 

bia. Valle: Rio Digua, Piedra de Moler, 24 Aug 1943 (NA). HEREDIA: Puerto Viejo, Rio Sarapiqui, 19 Jan 

(9), Cuatrecasas 15071 (holotype, F; isotypes, GH, 1974 (st), Hartshorn 1348 (F), 1 Oct 1974 (9), Hart- 

NY). 

shorn 1539 (F, U); Rio Sarapiqui, Tirimbina, 16 Jul 

1971 (6), Lent 2004 (F, G, MO, NY, U, US); Rio 

Leafy twigs appressed-puberulous to strigose Sarapiqui, La Virgen, 23 Jul 1979 (9), Stevens 13341 

or hirtellous. Lamina usually 5-7(-9)-parted, (U). LIMON: Madre de Dios, 26 Feb 1949 (9), Holdridge 

sometimes 3-lobed to -parted, midsegment 

2514 (US); Rio Hondo, Aug 1901 (9), Pittier 16163 

mostly lanceolate to oblong, sometimes (G, US). PUNTARENAS: Valley of Rio Diquis, between 

elliptic Union and San Pedro, 29 Mar 1898 (9), Pittier 12105 

to obovate, base + deeply cordate; upper surface (BR, P, US). SAN JOSE: San Pablo, 2 Mar 1966 (9), 

scabrous (to scabridulous), lower surface smooth, Jimenez 3816 (F); San Isidro, El General, 2 Mar 1966 

hairs on the main vein appressed to patent, ar- (9), Molina R. 18227 (F, NY, US); El General, Jan 

achnoid hairs (almost) confined to the areoles 

1939 (6), Skutch 4083 (A, MO, NY, S, US), Feb 1939 

but the spots of arachnoid 

(9), Skutch 4225 

hairs mostly not clear- 

(A, MO, NY, S); Rio de las Vueltas, 

Tucurrique, Dec 1898 (6), Tonduz 12930 (F, GH, M, 

ly separated by the smaller veins; smaller veins NY, P, S, US, type collection of P. scobina). 

more or less prominent; stipules outside with ap- PANAMA. COCLE: Valley of Rio Anton, 1000 m, 

pressed or patent hairs, inside hairy, caducous. 27 Sep 1946 (9), Allen 3742 (BR, E, F, G, MO, P). 

Pistillate 

COLON: Santa 

inflorescences with (10-)20-50 (or more 

Rita, Feb 1968 (9), Gomez-Pompa 3385 

(MO). DARIEN: Cerro Pirre, 14 Dec 1962 (9), Duke 

?) flowers. 

6588 (MO); Pidiaque, 28 Mar 1966 (9), Duke 8035 

Distribution (Fig. 68). Central America (from (MO); between Cerro Pierre and Piji Vasal, 15 Nov 

Guatemala to Panama) and the Pacific coastal 1977 (st), Folsom 6377 (MO); Cocalito, 8-9 Apr 1978 

region of Colombia and Ecuador, and to north- (st), Garwood 759 (F); Cerro Pirre, 1000-1400 m, 29 

Dec 1972 

western Venezuela, possibly also in 

(9), 

Peru 

Gentryet al. 6997 (F, MO); Rio Setigandi, 

(Loreto 19 Apr 1980 (9), Gentry et al. 28609 (U); Manene, 23 

and San Martin); in non-inundated forest at al- Dec 1980 (9), Hartman 12176 (U); between Paya and 

titudes up to ca. 1600 m. 

Pucro, 12 Jun 1959 (9), Steam et al. 423 (G, GH, LE, 

US); Pediaque Hill, nr. Rio Sabana and Rio Lara, 16 

Specimens examined. GUATEMALA. ALTA VERA- Jul 1966 (2), Tyson et al. 4734 (DUKE, GH, MO); 

PAZ: Finca Cubilguitz, Mar 1913 (Q), Tuerckheim 4082 Cana, 17 Apr-8 Jun 1908 (9), R. S. Williams 983 (NY, 

(US). IZABAL: Puerto Mendez, 12 Jun 1970 (2), Con- US). PANAMA: El Llano-Carti rd., 20 Feb 1973 (9), 

treras 10026 (DUKE, F, U, US); El Estor, 7 Mar 1972 Correa et al. 1854 (F, MO); Cerro Azul, 26 Jul 1970 

(2), Contreras 11195 (DUKE, P, S, US); Entre Rios, (st), Croat 11591 (F, MO); between Cerro Azul and


' I? 

UNITDO STATES NATIONAL li55UII5553 

inch ? 

. J e . td. . 

-N- 

FIG. 66. Pourouma bicolor subsp. scobina. Leafy twig with pistillate inflorescences 

(Allen 6401). 

Cerro Jefe, 6 Aug 1968 (Q), Dressier 3579 (DUKE, GH, 

MO); El Llano-Carti rd., 17 Apr 1981 (Q), Sytsma 4018 

(U). SAN BLAS: Cangandi, 10 Feb 1986 (st), Nevers et 

al. 7178 (BG), 27 Mar 1986 (st), Nevers et al. 7542 

(BG). 

COLOMBIA. CHOC6: 

Rio Curiche, camp Curiche, 

25 May 1967 (9), Duke 11599 (US). VALLE: Rio Digua, 

Piedra de Moler, 24 Aug 1943 (2), Cuatrecasas 15071 

(F, GH, NY, type collection of P. crassivenia); Rio 

Sanquinini, La Laguna, 14 Dec 1943 (d), Cuatrecasas


15520 (F, GH, NY), (9), Cuatrecasas 15534 (F, GH, subspecies and in that of the other subspecies, 

NY). 

except for subsp. chocoana. Such specimens are 

VENEZUELA. BARINAS: Pedreza, Feb 1955 (2), 

Bernardi 1986 (NY, VEN); rd. Barinitas-El Cacao, 21 hardly distinguishable from P. guianensis subsp. 

Sep 1982 (9), Marcano-Bert et al. 982-154 (U). MERI- guianensis with regard to the leaf indument (see 

DA: Caiio Amarillo, 5 Feb 1955 (2), Bernardi 1919 (F, p. 122). Fortunately these taxa can mostly be 

NY, VEN); El Vigia, 14 Jun 1979 (6), Marcano-Bert distinguished on the presence of hairs on the inet 

al. 238-979 (U). TACHIRA: La Fria, 22 Dec 1965 (2), ner surface of the 

Breteler 4920 (MO, U); 35 km SSE of San stipules and are, moreover, 

Cristobal, 

20-21 Mar 1981 (2), Liesner et al. 10944 (BG); San geographically separated. However, two recent 

Joaquino de Navay, 6 Nov 1979 (2), Stevermark et al. collections from Peru (San Martin; Gentry et al. 

119396 (U). ZULIA: Dtto. Col6n, nr. Casuigua El Cubo, 45160) and (Loreto, Gentry et al. 42168) may 

21 Jan 1979 (9), Bunting 6836 (BG); Sierra de Perija, also 

SW ofMachiques, 31 Aug 1967 (2), Steyermark 99945 

belong to subsp. scobina. The indument on 

the 

(U, VEN). 

petiole, the leafy twigs, and the main veins 

ECUADOR. CARCHI: Maldonado, 25 Sep 1979 (6), of the lamina beneath is denser than usual in the 

Gentry et al. 26530 (U). EL ORO: Piedras, 20 Jun 1943 collections from Central America and western 

(st), Little 6640 (F, US). ESMERALDAS: Santo Domingo Ecuador. The latter collection has, moreover, 

de los Colorados-Quininde rd., 14 Apr 1943, Acosta 

S. 13605 (F); Quininde, 12-14 Apr 1943 (2), Little 6241 bright-yellow hairs on the stipules and 9-parted 

(F, US), (st), Little 6258 (F, UC, US); San Lorenzo, 20 leaves with subpetiolulate segments. The identity 

Apr 1943 (2), Little 6301 (F, U, UC, US). IMBABURA: of these two collections is rather uncertain. 

Collapi, 4 Jun 1949 (st), Acosta S. 12884 (F). Los Rios: 

Rio Palenque Biol. Station, 16 Aug 1976 (st), Dodson 

6156 (MO), 2 Oct 1976 (st), Dodson et al. 6312 (MO), 3e. Pourouma bicolor Martius subsp. chocoana 

(6), Dodson et al. 6327 (GB, MO, U), 2 Feb 1974 (9), (Standley) C. C. Berg & van Heusden, Proc. 

Gentry 9552 (MO, U). PICHINCHA: Santo Domingo de Kon. Ned. Akad. Wetensch., Ser. C 91(2): 106. 

los Colorados, 30 Aug 1930 (6), Benoist (P, S, U), 17 

Oct 1961 (2), Cazalet et 

1988. 

al. 5037 (FHO, NY, UC, US), 

Fig. 67 

19 Oct 1961 (6), Cazalet et al. 5072 (FHO, NY, UC, Pourouma chocoana Standley, Publ. Field Mus. Nat. 

US). 

Hist., Bot. Ser., 22: 73. 1940. 

PERU. LORETO: Prov. Maynas, Yanamono 

Type: Colombia. Cho- 

Explora- c6: Junction of Rio Condoto and Rio San 

ma Tourist Camp, 26 Jun 1983 (st), Gentry et al. 

Juan, 20 

42168 

(BG). SAN 

Apr 1939 

MARTIN: 

(2), Killip & A. C. Smith 35095 

Rioja-Pomacochas rd., below 

(holotype, 

F; 

Venceremos, 1600 m, 8 Feb 

isotype, US). 

1984 (9), Gentry et al. Pourouma 

45160 

johnstonii Woodson, Ann. Missouri Bot. 

(BG). 

Gard. 47: 166. 1960; Burger, Fieldiana Bot. 40: 201. 

1977. Type. Panama. Canal Zone, W of Lim6n Bay, 

Vernacular names. Guatemala. Guarumo de Gatun Locks, Maru Towers, 27 Mar 1956 (Q), Johnston 

1714 

montafia. Belize. Trumpet tree. Honduras. 

(holotype, MO; isotype, A). 

Guarumo de montafia, mano de le6n. Nicaragua. 

Guarumo macho, pacica, yahal (Moskito Indians). 

Costa Rica. Chumico, guarumo de montafia, 

lija. Panama. Darien: guarumo macho, piraejo. 

Colombia. Choc6: uva. Venezuela. Barinas: 

tinajero. Merida: orumo de monte. Ecuador. Esmeraldas: 

guagay, guarumo de montafia, uva, uva 

del monte. 

The lamina is mostly 5-7-fid to -parted, but 

in Panama 3-lobed laminas occur. The number 

of flowers in the pistillate inflorescence is often 

50 or more. In Guatemala and Panama collections 

with (10-20) flowers in the pistillate inflorescence 

have been made. In several collections 

in the southern part of the subspecies range (Ec- 

Leafy twigs patent- to appressed-puberulous 

and at least on the scars of the stipules (sparsely) 

yellow-hirsute. Lamina mostly 5-fid to -parted, 

sometimes 3-lobed to -fid or entire, occasionally 

(in subjuvenile material?) 7-parted, midsegment 

broadly elliptic to oblong, base deeply or sometimes 

shallowly cordate; upper surface scabrous, 

lower surface smooth, appressed hairs on the main 

veins, arachnoid hairs in the areoles and on the 

smaller veins, mostly covering the areas between 

the parallel tertiary veins; smaller veins (almost) 

plane; stipules outside yellow-hirsute to -subsericeous, 

inside hairy, subpersistent. Pistillate inflorescences 

mostly with up to 50 (or more?) flowers. 

uador to Panama) the indument of the lamina 

beneath, the petioles, stipules, peduncles and leafy 

twigs is patent (hirtellous to (sub)tomentose) instead 

of appressed as in most collections of this 

Distribution (Fig. 68). Panama (Canal Zone 

and Col6n) to western Colombia (Choc6 and 

Valle); in non-inundated forest at altitudes up to 

ca. 1000 m. 

141


movririg-. o 

-.. 

Ievsed for YXora eotr ei'' 

Pouroum bicolar orths* subsp, chocoasa 

?Staodisy) C.C. krg & van Seusden 

Dec~L~E~I!t.: C.c. Barg Utrechtr 19L6 

OPPANABMt 

der*-o lno Tru4. . 5 

un~err 

t.e inch99 Jaa 

en 

~rp~ 

i 

~ ~ ~ r 

rortbO~h ofr trwil, tr--, : rtire tr,an~~~~~~~~~~~~~~~~~~~rt-in~f4etedm 

tkr~- ~-~:ht brow:1., ,('!'t wi'; 

~ag 

irr-f~'Lr hc~riz,.ntml rineg; c~r~. 1 

%N0 p 

C.RAVfJR~rok;%s~l 

6,FR pCv.bl Ftrol..' 

Il . Feb. r ; z, 1c60 

"'Cf.7 

FIG. 67. Pourouma bicolor subsp. chocoana. Leafy twig with staminate inflorescence 

(Croat 8097). 

Specimens examined. PANAMA. Without locality, 

1861 (st), Hayes 32 (E), without date (2), Hayes 348 

(NY). CANAL ZONE: Nr. Gamboa, 8 Feb 1966 (2), Blum 

2039 (MO), 19 Jun 1968 (st); Barro Colorado Island, 

22 Feb 1969 (d), Croat 8097 (F, MO, NY), (9), Croat 

8100 (F, MO), 14 Mar 1969 (st), Croat 8648 (MO), 23 

May 1970 (st), Croat 10343 (MO), 9 Jun 1970 (st), 

Croat 10830 (MO); nr. Gamboa, Foster et al. 584 

(DUKE, NY), 18 Dec 1972 (d), Gentry 6679 (MO); 

Gatun Lake, Gatun Locks, 27 Mar 1956 (2), Johnston 

1714 (A, MO, type collection of P. johnstonii); Barro 

Colorado Island, 16 Aug 1927 (st), Kenoyer 312 (US), 

27 Oct 1931 (st), Shattuck 260 (A, F, MO), 4 Dec 1931 

(d), Shattuck 522 (F, MO, US) and 525 (F, MO); N of 

Frijoles, 19 Dec 1923 (st), Standley 27479 and 27502 

(US). CoIkN: Donese district, Camp Batija, 9-14 May 

1968 (9), Holdridge 6254A (MO). 

COLOMBIA. CHOC6: Quibdo-Istmina rd., 48 km 

S of Quibdo, nr. Certegui, 8 Jan 1979 (9), Gentry et al 

23835 (BG); Quibdo-Tutunendo rd., 3 km W of Tutunendo, 

7 Jan 1981 (st), Gentry et al. 30278 (U); junction 

of Rio Condoto and Rio San Juan, 20 Apr 1939

_' "( 

O^^Jgy -' i biccoolr 

P. bicolor subsp. 

-r~ E^^ 

- 

_SAs. 

* 

L ? 

*?'7 ^ ^ 

[ 

*' 

"*^ 

^ ~ * nde 

s.cbina 

Aand 

p. biColor subsp. digitate 

- 

P. bicolor subsp. 

P. bicolor *P. ccrepifo. 

'- s 

FIG. 68. Distribution of Pourouma bicolor subsp. bicolor, P. bicolor subsp. chocoana, P. bicolor subsp. 

digitata, P. bicolor subsp. scobina, P. bicolor subsp. tessmannii, and P. cecropiifolia.


(9), Killip 35095 (F, US, type collection ofP. chocoana). 7: 231. 1927. Type. Bolivia. Beni: Rurrenabaque, 

NARIMO: Mun. Barbacoas, Barbacoas-Junin rd., 1050 Oct 1921 (9), Rusby 1599 (holotype, NY; isotypes, 

m, 7 Aug 1982 (9), Mora 2295 (US); Mun. Iscuande, F, GH, US). 

Rio Sequi6n, 22 Nov 1955 (d), Romero-Castaneda 5484 

(AAU). VALLE: Rio Yurumangui, Veneral, 28 Jan-10 Tree, up to 20 m tall. Leafy twigs 5-30 mm 

Feb 1944 (a), Cuatrecasas 15876 (F); Rio Calima, La 

Trojita, 19 Feb-10 Mar 1944 (2), Cuatrecasas 16315 

thick, (sparsely) white-pubescent and with sparse 

(F); La Esperanza, 6-7 Mar 1944 (d), Cuatrecasas 16755 

to dense, brown, pluricellular hairs, often gla- 

(F); San Isidro, 2-5 May 1944 (8), Cuatrecasas 17357 brescent. Lamina 7-1 1-parted, rarely 3-5-lobed, 

(F); Rio Cajambre, Silva, 5-15 May 1944 (st), Cuatre- ca. 10-40 x 13-60 cm, coriaceous, apex acucasas 

17451 and 17479 (F); Bajo Calima, ca. 15 km N minate to acute, base deeply cordate, with or 

of Buenaventura, 18 Feb 1983 (st), Gentry et al. 40479 without 

(BG); Bajo Calima, rd. to Juanchaco Palmares, 18 Jul overlapping lobes; upper surface smooth, 

1984 (2), Gentry et al. 48303 (BG). 

white puberulous on the main veins and sometimes 

with sparse, red-brown, pluricellular hairs 

Vernacular names: Panama. Mangabe. Colom- usually on the smaller veins, lower surface whitebia. 

Antioquia: sirpo; Nariiio: guagay. or sometimes yellow-, usually appressed-pubes- 

This subspecies can be easily distinguished cent to -puberulous on the veins, occasionally 

from subsp. scobina by the arachnoid indument brown, pluricellular hairs on the main veins, 

covering the smaller veins (between the parallel arachnoid hairs in the areoles, often also on the 

tertiary veins), and often also by the relatively smaller veins; lateral veins ca. 15-30 pairs, terbroad 

segments of the lamina. Some of the spec- tiary venation almost plane beneath; petiole 8- 

imens are yellow-hirsute on the leafy twigs, or 50 cm long, glabrous or occasionally with brown, 

also on the stipules, petioles, main veins of the pluricellular hairs; stipules 2-20 cm long, outside 

lamina above and/or the peduncles. Such spec- white-appressed-puberulous and with (dense) redimens 

resemble P. oraria very much. The two brown, pluricellular hairs, inside densely yellowtaxa 

differ in the smooth upper surface of the hirsute and sometimes with brown, pluricellular 

lamina, the smooth fruiting perianth, and the hairs, caducous. Staminate inflorescences up to 

concentration of hirsute indument at the adaxial 27 cm long and 17 cm wide; peduncle 2-11 cm 

side of the petiole. The resemblances between long, peduncle and branches yellow- or white-, 

these two taxa are so striking that we sometimes usually appressed-puberulous and usually with 

wondered whether they could be two forms of dense, (dark- or red-)brown, pluricellular hairs; 

the same species (distinct from P. bicolor). flowers sessile, ? diffusely distributed along the 

ultimate branches; tepals ca. 2.5 mm long, free 

4. Pourouma cecropiifolia Martius in Spix & or occasionally basally connate, ciliolate; fila- 

Martius, Reise Bras. 3:1130. 1831, as Puruma ments shorter than the tepals, free. Pistillate incecropiaefolia; 

Martius, Syst. mat. med. bras. florescences in fruit up to 24 cm long and 22 cm 

34. 1843; Miquel in Martius, Fl. bras. 4(1): wide; peduncle 2-13.5 cm long, peduncle and 

123, t. 36. 1843; Macbride, Publ. Field Mus. branches with indument similar to that of the 

Nat. Hist., Bot. Ser., 13(2.2): 291. 1937. Type. 

staminate inflorescence; flowers 25-170; pedicel 

Brazil. "In silvis prov. Paraensis et Rio Ne- up to 0.6 cm, in fruit up to 2 cm long; perianth 

gro," Dec 1819 (st), Martius s. n. (lectotype M, 

4-9 mm long; white- to yellow-puberulous and 

chosen here). Fig. 69. usually with (dark or red-) brown, pluricellular 

hairs; stigma peltate, ca. 1.5-2 mm diam., sparsely 

white-puberulous. Fruiting perianth dark purple 

to black, ovoid to subglobose, 1.5-3.5 x 0.7-2 

cm, densely to sparsely puberulous and sometimes 

with brown, pluricellular hairs. 

Pourouma multifida Trecul, Ann. Sci. Nat. Bot., Ser. 

3, 8: 107. 1847; Miquel in Martius, Fl. bras. 4(1): 

123, 1853. Type. Brazil? Without locality, without 

date (9), Anonymus (=Ferreira?) s.n. (holotype, P; 

isotype, P). 

Pourouma sapida Karsten, Fl. Columb. 2: 19, t. 110. 

1862. Type. Colombia. Meta: Villavicencio, nr. Jiramene, 

Oct 1872 (a), Karsten s.n. (holotype, GOET; 

isotype?, LE-with fragments of staminate inflorescences!). 

Pourouma edulis Dufresne, Ill. Hort. 20: 70. 1873. Based 

on (living?) material from Colombia, collected by 

Linden). 

Pourouma uvifera Rusby, Mem. New York Bot. Gard. 

Distribution (Fig. 68). Extending from Colombia 

(Meta) through Amazonian Ecuador, Peru, 

and Brazil (Acre and western Amazonas) to Amazonian 

Bolivia, also in Amazonian Venezuela; 

in non-inundated forest, at altitudes up to ca. 

1000 m. Often cultivated and their occurrence 

often indicating previous human inhabitation.


,J, 

? ^T \Sr~~u. I 

. . 

0f W/+ O.+ 14+ I+ 

P OUR 0 ITMA cecropiaefolia. 

FIG. 69. Pourouma cecropiifolia. From Martius, Flora Brasiliensis 4(1). 1853: tab. 36. Leaf, stipule (32), 

staminate inflorescence, diagram of staminate flower (D), part of staminate flower (28), staminate flower (1), 

tepals (4), stamens (7).


Also cultivated in some places outside the species 

range, e.g., Bahia (Brazil). 

al. 43137 (BG); 2-8 Aug 1929 (9), Killip et al. 27381 

(NY, US); Yurimaguas, 24-25 Aug 1929 (2), Killip et 

al. 27932 (F, NY, US); Iquitos, 26 Sept 1929 (2), Killip 

et al. 29839 (F, NY, US), Apr 1930 (6), Klug 1185 (F, 

Specimens examined. COLOMBIA. Without locality, 

without date (9), Triana 862 (E, NY, P), without NY, US), May 1930 (2), Klug 1326 (F, NY, US); Rio 

date (9), Triana 1892 (NY). AMAZONAS: Rio Boiauasu, Marafion, Perseveranca, 5 Feb 1924 (2), Kuhlman 1337 

28 Nov 1945 (st), Duque-Jaramillo 2261 (NY); Rio (F, RB); Pongo de Manseriche, 11 Dec 1931 (2), Mexia 

Loretoyacu, Barracon, 25 Dec 1945 (2), Duque-Jara- 6257 (F, G, GB, NY, S, U, UC, US); Prov. Requena, 

millo 2446 (NY); between Rio Kananari and Rio Pa- nr. Jenaro Herrera, 4 Nov 1986 (2), Peters 187 (BG); 

coa, 1-5 Dec 1951 (6), Garcia-Barriga 13871 (US); Rio Rio Yaguasyacu, Brillo Nuevo, 16 Apr 1977 (2), Plow- 

Igara-Parana, La Chorrera, 20 Sep 1973 (2), Sastre 2274 man et al. 6903 (F, GH); Rio Yucayali, 1923 (a), Tess- 

(G, P); Los Monos, 24 Sep 1978 (st), Pabon E. 573 mann 3054 (G, S); Rio Itaya, Paraiso, 9 Oct 1929 (2), 

(herb. Araracuara); Rio Caraparana, El Encanto, 22- Ll. Williams 3347 (F, US); Yurimaguas, 4 Nov 1929 

28 May 1942 (st), Schultes 3826 (NA), (2), Schultes (9), Ll. Williams 3984 (F), 10 Nov 1929 (2), LI. Wil- 

3862 (F); Rio Igara-Parana, La Chorrera, 4-10 1942 liams 4627 (F). MADRE DE Dios: Rio Manu, Cocha 

(2), Schultes 3901 (F); Rio Boiauasu, Nov 1945 (2), Cashu Station, 25 Nov 1980 (2), Foster 5905 (F), 15 

Schultes 6805 (F, US). CAQUETA: Las Guacamayas, 28 Sep 1979 (st), Foster 7001 (F); Tambopata, 19 Feb 

Apr 1944 (9), Little 7756 (US); Morelia, 7 Oct 1941 1984 (st), Gentry et al 45660 (BG). PASCO: Prov. Ox- 

(6), von Sneidern s.n. (S). META: Villavicencio, nr. Jira- apampa, Iscozacin, 24 Jun 1982 (2), D. Smith et al. 

mene, Oct 1872 (9), Karsten s.n. (GOET, LE), (6), Kar- 2089 (BG). SAN MARTIN: Puerto Pizana, 27 Jul 1973 

sten s.n. (GOET, LE, type collection of P. sapida). (9), J. Schunke V. 6485 (F, GH, MO, NY). UCAYALI: 

PUTUMAYO: Rio Putumayo, 28 Aug 1966 (9), Pinkley Prov. Coronel Portillo, Lago Yarinacocha, Povenir, 5 

405 (S); Puerto Lim6n, 27-28 Feb 1942 (st), Schultes Nov 1984 (2), Maas et al. 6201 (U); Prov. Coronel 

3343 (F). VAUPES: Mitui, 10 Sep 1939 (9), Cuatrecasas Portillo, km 13 of rd. from Campo Verde, km 34 of 

et al. 6737 (F); Upper Rio Guaviare, 9 Nov 1939 (6), rd. Pucallpa-Tingo Maria, 7 Nov 1984 (2), Maas et al. 

Cuatrecasas 7602 (F, US); Mitui, 7-8 Nov 1952 (2), 6225 (U). 

Humbert et al. 27221 (P, U, US); nr. Yavarate, 20 Nov BRAZIL. Without locality, without date (2), Anon- 

1952 (6), Romero-Castaneda 3647 (NY); Mitu, 26 Sep ymus (=Ferreira?) s.n. (P, type collection of P. multi- 

1976 (6), Zarucchi 2145 (INPA). 

fida). ACRE: Rd. Abuna-Rio Branco, km 242-246, 20 

VENEZUELA. AMAZONAS: Cerro Duida, base of Jul 1968 (2), Forero et al. P.6419 (INPA, NY, R, S, 

Duida, Jan-Feb 1939 (st), Farinas et al. 347 (NY, U, U); mouth of Rio Macauhan, 8 Aug 1933 (2), Krukoff 

VEN); Depto. Atures, 35 km SE of Puerto Ayacucho, 5327 (A, F, G, LE, M, MO, NY, S, U, UC, US), (a), 

25 Sep 1980 (9), Guanchez 239 (VEN); Rio Orinoco, Krukof 5332 (A, F, G, M, MO, NY, S, U, UC); Rio 

between Cafno Grulla and Grulla, 8 Apr 1978 (9), Mo- Branco, Parque Zoobotanico da Univ. Fed., 15 Oct 

rillo et al. 7384 (VEN); Depto. Atapabo, nr. Culebra, 1980 (2), Nelson 705 (BG); Tarauaca, 24 Sep 1968 (2), 

Rio Cunucunuma, 22 Mar-4 Apr 1983 (st), Steyer- Prance et al. 7503 (F, INPA, M, NY, P, R, S, U, US); 

mark et al. 129163 (BG, VEN). 

Sao Francisco, Jun 1911 (2), Ule 9314 (G, MG). 

ECUADOR. MORONA-SANTIAGO: 5 km SW of Ma- AMAZONAS: Rio Javari, Atalaia do Norte, without date 

cas, 26 Jan 1981 (6), Berg 1223 (BG, U). NAPO: Tena, (2), Braga et al. 3178 (INPA); Manaus (cult.!), 15 Apr 

20 Oct 1939 (9), Asplund 9462 (R); San Pablo de los 1955 (2), Chagas (INPA) 962 (INPA, U), 29 Jul 1978 

Secoyas, 19 Aug 1980 (9), Asanza C. 32929 (AAU); (6), Falcao 215 (INPA); Marco, 29 Jan 1969 (2), Croat 

Aiiangu, 30 May-21 Jun 1982 (st), SEF8764 and 8930 7633 (MO); Rio Japura, mouth of Rio Apaporis, 4 Dec 

(U); 3 km E of Atahualpa, 10 km E of Puerto Napo, 1912 (a), Ducke (MG) 12366 (MG); Rio Solimoes, Fon- 

13 Dec 1985 (9), Neill et al. 7044 (BG). PASTAZA: Rio teboa, 8 Jun 1945 (9), Fr6es 21048 (NY, US); Tefe, 19 

Curaray, Loricachi, 30 May 1980 (9), Jaramillo et al. Jul 1972 (9), Krieger et al. (INPA) 2268 (INPA); mouth 

31516 (AAU). 

of Rio Embira, 1 Jul 1933 (2), Krukoff 5109 (A, F, G, 

PERU. AMAZONAS: Rio Cenepa, 13 Dec 1972 (2), M, MO, NY, S, UC, US); Sao Paulo de Olivenca, 11 

Berlin 523 (F, GH, NY), 4 Aug 1974 (9), Berlin 1999 Sep-26 Oct 1936 (2), Krukoff8332 (BR, F, G, LE, MO, 

(NY). HUANUCO: Tingo Maria, 10 Aug 1940 (9), As- P, S, U, US), (2), Krukoff 8469 (A, B, BR, F, G, LE, 

plund 12933 (S, US), 10 Jul 1957 (st), Ellenberg 2288 MO, NY, P, S, U, US); Manaus, Estrada Roas de Maio, 

(U); Aucayacu, 27 Sep 1967 (9), Lao et al. s.n. (F, G, Armando (cult.!), 25 Sep 1973 (8), Lisboa 27 (INPA); 

US); Puerto Inca, 7 Dec 1968 (2), J. Schunke V. 2824 Rio Papori, Trindade, 13 Dec 1928 (st), Luetzelburg 

(F, G, GH, NY, US). JUNiN: La Merced, 10-24 Aug 23913 (R), 27 Dec 1928 (2), Luetzelburg23887 (M, R); 

1923 (9), Macbride 5446 (F, GH, US). LORETO: Iquitos, Rio Curicuriari, 8 Jul 1979 (2), Maia et al. 497 (INPA); 

29 Oct 1970 (2), Asplund 14124 (G, S); Zungarococha, various collections without precise indications of lo- 

28 Jan 1978 (st), Ayala et al. 1446 (U); Rio Ampiaco, calities (probably most of them along Rio Amazonas 

Puca Urquilla, 24 Feb 1972 (9), Croat 20662 (DUKE, and Rio Japurf, (Dec) 1819-(Feb) 1820 (6 or st), Mar- 

MO, NY, U); Iquitos, 8 Aug 1906 (9), Ducke (MG) tius s.n. (G, M, including lectotype of P. cecropiifolia); 

7581 (G, MG); El Sacramento, 89 km S of Pucallpa, Taparuquara, 22 Oct 1971 (2), Prance et al. 15791 

22 Jul 1957 (st), Ellenberg 2474 (U); halfway between (INPA, NY, U), Jul-Aug 1967 (9), Schultes 24547 

Iquitos and Santa Maria de Nanay, 30 May 1978 (st), (INPA); between Barcellos and Sao Gabriel, Dec 1851 

Gentry et al. 22460 (U); Prov. Maynas, Yanamono (2), Spruce 2023 (B, BR, CGE, P); Rio Jurua, Bom Fin, 

Explorama Tourist Camp, 15 Jul 1983 (st), Gentry et Oct 1900 (2), Ule 5265 (B, F, G, HBFG, MG, NY);


Rio Jurua, Marary, 8 Oct 1900 (c), Ule 5266 (HBG, 1951 (e), Garcia-Barriga 14002 (holotype, US; iso- 

MG). 

type, NY). 

BOLIVIA. BENI: Prov. Moxos, S of Moxos, 18 Sep 

1976 (9), Meneces et al. 338 (MG, MO); Rurrenabaque, Tree, up to 30 m tall. Leafy twigs 4-8 mm 

Oct 1921 (9), Rusby 1599 (F, GH, NY, US). PANDO: thick, densely to sparsely yellow-hirsute, these 

Prov. Nicolas Suarez, Mukden, Jun-Dec 1979 (2), Izawa 

21 long hairs often intermixed with much shorter, 

(U). 

whitish hairs, sometimes also with sparse, brown, 

Vernacular names. Colombia. bochoa tsaha pluricellular hairs, sometimes (sub)glabrous or 

(Kofan), caimaron, caimaron silvestre, uva, uva subvelutinous. Lamina entire, ovate to subovate 

del monte, uvilla, uvo; Amazonas: gurucana. or elliptic to oblong and 6-20 x 2.5-12 cm or 

Venezuela: cocura, sadajii. Peru. sacha uvilla, 3-lobed to -parted (or in subjuvenile material up 

uvilla, suia; Pasco: shewantaqui. Brazil: cucura, to 5-parted, sometimes with the segments almost 

cucuva, imbauba (or ambauva) mansa, imbafba petiolulate) and up to ca. 25 x 25 cm, coriaceous 

(or ambauva) do vinho, mapati, puruma, puru- to subcoriaceous, apex acuminate, acute, apicma-y, 

sucuuiba, uva. Bolivia. uva silvestre; Beni: ulate, or sometimes emarginate, base subcordate 

tanaribe, uva del monte. 

to truncate to rounded to subacute; upper surface 

This species shows strong similarities to P. bi- ? scabrous, occasionally smooth, yellow-hirsute 

color, especially to the subspecies digitata and to -hirtellous on the whole surface or only on the 

scobina. It mainly differs in the greater number main veins, lower surface yellow-hirsute (to -hirof 

segments of the leaves of adult specimens (nor- tellous) to -substrigose on the main veins, white 

mally 7-11 versus normally 5-7 in subsp. digi- arachnoid hairs in the areoles and on the smaller 

tata and subsp. scobina), the smooth upper sur- veins; lateral veins 10-20 pairs, the basal pair 

face, and the sparser indument on the leafy twigs. branched, tertiary venation prominent beneath; 

The lamina of subsp. digitata and subsp. scobina petiole 3-17 cm long, yellow-hirsute, intermixed 

is sometimes smooth and subjuvenile specimens with short, whitish hairs; stipules 2-12.5 cm long, 

may have up to 9 leaf segments, which makes outside yellow-hirsute to -strigose, inside gladistinction 

of the taxa on the basis of morpho- brous, caducous. Staminate inflorescences up to 

logical characters sometimes impossible. For- 12 cm long and 4.5 cm wide; peduncle 3-6.5 cm 

tunately, the taxa are allopatric (or almost so, see long, peduncle and branches yellow-hirsute; 

p. 139). 

flowers sessile, rarely pedicellate, clustered, but 

The lectotype ofP. cecropiifolia is selected from not in globose heads; tepals almost free, (sparsenine 

Martius syntype collections listed above, ly) puberulent; filaments free, shorter than the 

that of P. sapida from two Karsten (syntype) col- tepals. Pistillate inflorescences in fruit up to 19 

lections. 

cm long and 10.5 cm wide; peduncle 3-9 cm long, 

The fully mature fruiting perianth of most (if peduncle and branches yellow-hirsute;flowers 20not 

all) Pourouma species is suitable for human 25; pedicel up to 0.6 cm long, in fruit up to 1 

consumption, but P. cecropiifolia is the only cm; perianth 0.3-0.9 mm long, hispidulous (or 

species in cultivation (see p. 116). 

hirtellous); stigma peltate, ca. 2 mm in diam., 

The fact that the species is so often found in whitish puberulous. Fruiting perianth blue-black, 

(formerly) inhabited places suggests that the ellipsoid to oblongoid to subglobose, ca. 1-1.5 

present distribution is (at least) partly anthro- cm long, hispidulous (or hirtellous). 

pogenous. 

Distribution (Fig. 74). Colombia (Amazonas 

and Vaupes), Venezuela (Amazonas and Boli- 

5. Pourouma cucura Standley & Cuatrecasas in var), Ecuador (Napo and Morona-Santiago), Peru 

Cuatrecasas, Fieldiana Bot. 28(1): 211. 1951. (Loreto and Madre de Dios) and Brazil (Amapa, 

Type. Venezuela. Amazonas: Capihuara, Alto Amazonas, and Mato Grosso), extending to east- 

Casiquiare, 5 Jun 1942 (Q), LI. Williams 15812 ern Venezuela (Bolivar); in non-inundated or in 

(holotype, F; isotypes, G, NY, RB, US). periodically inundated (varzea or igap6) forest; 

Fig. 70. mostly at low altitudes, sometimes (in Ecuador) 

up to ca. 1500 m. 

Pourouma garciana Cuatrecasas, Caldasia 7:298. 1956. 

Type. Colombia. Amazonas-Vaupes: Rio Apaporis, Specimens examined. COLOMBIA. AMAZONAS: Rio 

between Rio Kananari and Rio Pacoa, 1-15 Dec Apaporis, above mouth of Rio Kananari, Soratama,


dl;i?~~~~~~~~~~~~~~~~~~~~~~~~~,T! 

~ ~ ~ ~ ~ ~ ~ ~ ~ ?~ 

::. 

IWITi NAIONAL to PiQtJLAS IA AMAI7AA4A 

5352 ew J Starl,l. /rurourm eurura 

.F2;?Z,3!: -~ '; T.'l 

^* 1 " -' 

FIG.~~~~~~~~~~~~~~~~~~1 

'v-~~~ 70. cucra 

^near 

- 

rlpu , Hufb,b't C t.r. - t 

Rio Jurw-. 59"'?1'N: 10 ':" 

disturbed f"rest r"n terra firrie. ;ie, . 

Nx hnc. lt . W*itbh tilt. 7.t. i.- ; - 

gla oiua to whitte benestb, 3 partim-l r *r 

fol~avt., p.-dofv pwerianthb, papper & "r " 

inftovescence brown. Puroua~ Exuding Lefwgwihpitllat 

nireenesl br iw (rnce -] et : 

t 'nning to blacsk. "UmbinublI 'A ,' L 

O.T.Pr nce, F.A.ilsby, C...Berg. 

W.C.St_fw d, E . Lleras, T . s.., 

-- 

ii____ _ _ . ctber A 

FIG. 70 Pourouma cu. atIg~~~~~ with pist~ilat ~ 

flrM.. a"n: s ( 

PA. r' ane, et. 

i.8 

l. 84) 

FIG. 70. Pourouma cucura. Leafy twig with pistillate inflorescences (Prance et al. 18240). 

15-19 Dec 1951 (9), Garcia-Barriga 14106 (NY, US), 

18 Mar 1952 (9), Schultes et al. 15973 (F), Jan 1952 

(6), Schultes et al. 19835 (US). AMAZONAS-VAUPES: Rio 

Apaporis, between Rio Kananari and Rio Pacoa, 1- 

15 Dec 1951 (9), Garcia-Barriga 14002 (GH, NY, US, 

type collection of P. garciana). 

VENEZUELA. AMAZONAS: Capihuara, Alto Casiquiare, 

5 Jun 1942 (9), Ll. Williams 15812 (G, NY, 

RB, VEN, US, type collection of P. cucura). BOUVAR: 

Rio Tirica, Oct 1947 (9), Cardona 2170 (MO, VEN). 

ECUADOR. NAPO: Rio Cuyabeno, 16 Feb 1980 (8), 

Berg & Akkermans 1041 (U), Berg & Akkermans 1042 

20.


(U), 17 Feb 1980 (6), Berg & Akkermans 1045 (U), (2), 3-lobed leaves (similar to the common leaf shape 

Berg & Akkermans 1053 (U), 20 Feb 1980 (st), Berg in P. bicolor 

& Akkermans 1070 (U); Rio 

subsp. bicolor). Subjuvenile material 

Aguarico, Shushufindi, 

26 Dec 1974 (2), Vickers 86 from these two countries often have 

(F). 

up to 7-parted 

PERU. LORETO: Rio Ucayali, Jenaro Herrera, Jul- leaves, with the segments often petiolulate. 

Sep 1976 (6), Bernardi s.n. (U); Rio Nanay, Puerto The species may prove to consist of two or 

Almendras, 22 Feb 1979 (st), Gentry et al. 24900 (U); three morphologically readily distinguishable 

Rio Yaguasyacu, tributary of Rio Ampiyacu, 9 Nov 

1977 (6), Gentry et al. 20515 (MO, U); Rio entities, 

Nanay, 

mainly based on differences in the shape 

halfway between Iquitos and Santa Maria de Nanay, 26 and incisions of the lamina. The relatively small 

Feb 1979 (st), Gentry et al. 25100, 25050 (U); E of number of collections prevent establishing new 

Tamshiyacu, below Serafin Filomeno, 19 Mar 1979 infra-specific taxa. 

(2), Gentry et al. 25854 (U); Rio Nanay, between Iqui- The 

tos and Santa Maria, de Nanay, 23 Feb 1979 identity of the two Rosa collections from 

(2), Gentry 

et al. 31655 (U); below Serafin Filomeno, 19 Mar 1979 Amapa (Serro do Navio), consisting only of (fal- 

(2), Gentry et al. 25854 (U); Rio Yaguasyacu, nr. Brillo len?) leaves, is not certain. The petioles and the 

Nuevo, 16 Apr 1977 (st), Plowman et al. 6906 (F, GH); main veins of the lamina beneath are densely 

Rio Nanay, nr. Iquitos, 14 Nov 1976 (st), Revilla 1818 

yellow-hirsute. 

(U); rd. to Pena Negra, 25 km from Iquitos, 30 Nov 

1976 (9), Rimachi 2724 (DUKE, F, MO, NY) Rio Nanay, 

above Bella Vista, 18 Jan 1977 (2), Ramachi 2765 6. Pourouma cuspidata Mildbraed, Notizbl. Bot. 

(F, MO); rd. from Orellana to Sta. Catalena, km 5, 9 

Gart. Berlin-Dahlem 10: 417. 

Dec 1982 (2), C. R. Rodrigues 944 (BG). MADRE 1926; 

DE 

Warburg 

Dios: Rd. Maldonado-Quincemil, 10 Jul 1968 (6), 

ex Ule, Bot. Jahrb. Syst. 40:150. 1907, nomen. 

Acosta M. 12 (BG, DUKE, F, NY, P); Tambopata, 22 Type. Brazil. Acre: Confluence of Rio Jurua- 

Feb 1984 (st), Gentry et al. 45922 (BG). 

Mirim and Rio Juruf, Aug 1901 (6), Ule 5719 

BRAZIL. ACRE: Reserva INCRA Santa Luzia, km 

40, BR-364, 5-19 Oct 1984 (st), Campbell et al. 6927 

(lectotype B, chosen here; isolectotypes, F, G, 

(BG). AMAPA: Serro do Navio, P6rto Platon, 10 Oct- GH, MG). 

15 Dec 1976 (st), Rosa 1091 (MG), 10 Oct-15 Dec Pourouma 

1976 (st), Rosa 1159 (MG). AMAZONAS: Rio tergoscabra Cuatrecasas, Caldasia 7: 304. 

Japura, 1956. 

Vila Bittancourt, 20 Nov 1982 (8), Amaral et al. 604 

Type. Colombia. Caqueta: Rio Caqueta, below 

mouth of Rio 

(BG); Manaus-Itacoatiara rd., km 26, 12 Aug 1976 

Orteguaza, Solano, 8 km SE of Tres 

(st), Oliveira (INPA) 60534 (INPA); Tototobi, Rio De- Esquinas, 4 Mar 1945 (v), Little et al. 9527 (holotype, 

mini, 25 Feb 1969 (9), Prance et al. 10231 

F; isotypes, NY, US). 

(INPA, MG, 

MO, NY, U, US); Manaus-Itacoatiara rd., km 26, 2 Tree, up to 30 m tall. Leafy twigs 8-12 mm 

Oct 1965 (st), Rodrigues 8066 (INPA). MATO GROSSO: 

Aripuana, 28 Jan 1977 (st), Gomes et al. 614 thick, white-hirtellous to -puberulous, and with 

(INPA), 

21 Feb 1977 (st), Gomes et al. 844 (INPA), 29 Mar usually rather dense, brown, pluricellular hairs. 

1977 (st), Gomes et al. 1040 (INPA); Aripuana, nr. Lamina 3-lobed or 3-5 parted, 10-35 x 18-45 

Humboldt Centre, rd. to Rio Juruena, 8 Oct 1973 (2), cm, coriaceous, apex long acuminate, base cor- 

Prance et al. 18240 (F, INPA, MG, MO, NY, P, S, U, date, with or without overlapping lobes; upper 

US). 

surface smooth, hirtellous or occasionally also 

Vernacular names. Venezuela. Amazonas: cu- with brown, pluricellular hairs on the main veins, 

cura; Bolivar: kaibarikei (Arekuna), sarasara lower surface strigose, main veins hispidulous 

(Camaracoto). Ecuador. Santiago-Zamora: guar- and sometimes with brown, pluricellular hairs, 

umo colorado. Peru. Loreto: dacha uvillos, uvil- arachnoid hairs confined to the areoles; lateral 

la; Brazil. Amazonas: imbaubarana branca; veins 13-30 pairs, tertiary venation + prominent 

Amapa: imbaubarana branca, inbaubarana folha beneath; petiole 11-40 cm long, white-hispidupeluda; 

Mato Grosso: imbaiuba or umbauiba, im- lous, and with brown, pluricellular hairs; stipules 

baubarana. 

7-13 cm long, outside white-appressed-puberu- 

The species is very variable. The long, yellow lous and with brown, pluricellular hairs, inside 

hairs on the leafy twigs, petioles and stipules vary sparsely white-puberulous to subglabrous, cafrom 

very dense to sparse. The leaves of (adult) ducous. Staminate inflorescences up to 11 cm 

specimens collected in Colombia, and Venezuela long and 4.5 cm wide; peduncle 4-6 cm long, 

and in Amapa and Amazonas (Brazil) normally (densely) white-puberulous and with (dense), 

have entire and elliptic leaves. The collections brown, pluricellular hairs; flowers sessile, + difmade 

in Mato Grosso (Brazil) have on the same fusely distributed along the ultimate branches; 

twig entire and 3-parted leaves. The material from tepals ca. 1 mm long, free or basally connate, 

Ecuador and Peru mostly have ovate and often sparsely hairy; filaments shorter than the tepals. 

149


Pistillate inflorescences in fruit up to 12.5 cm 

long and 8 cm wide; peduncle 4-6 cm long, peduncle 

and branches with indument similar to 

that of the staminate inflorescences; flowers 10- 

35; pedicel 0.2-0.4 cm, in fruit up to 0.8 cm long; 

perianth 2-6 mm long, hispidulous and with 

brown, pluricellular hairs; stigma peltate, ca. 1.5 

mm diam., white puberulous. Fruiting perianth 

ovoid to ellipsoid, 12-15 x 5-11 mm, scabrous. 

prominent beneath; petiole 8-27 cm long, whitish-hirsute 

to -hirtellous to -subhispid, at the base 

swollen and saccate; stipules 2-7.5 cm long, outside 

yellowish-hirsute, inside sparsely hairy, ca- 

Distribution (Fig. 74). Colombia (Caqueta) Ecuador 

(Napo), Peru (Pasco), and Brazil (Acre and 

Amazonas); in non-inundated forest. 

Specimens examined. COLOMBIA. CAQUETA: Solano, 

8 km SE of Tres Esquinas, Rio Caqueta, below 

mouth of Rio Orteguaza, 4 Mar 1945 (2), Little et al. 

9527 (F, GH, NY, US, type collection of P. tergoscabra). 

ECUADOR. NAPO: Afiangu, 30 May-21 Jun 1982 

(st), SEF 8532 (U). 

PERU. PASCO: Prov Oxapampa, Rio Iscozacin, Cabeza 

de Mono, 12 Jun 1983 (st), Gentry et al. 41890 

(BG). 

BRAZIL. ACRE: Reserva INCRA Santa Luzia, km 

40, BR-364, 5-19 Oct 1984 (st), Campbell et al. 6834 

(BG); confluence of Rio Jurua-Mirim and Rio Jurua, 

Aug 1901 (6), Ule 5719 (B, F, G, GH, MG, lectotype 

collection ofP. cuspidata), Aug 1901 (2), Ule 5719 (B, 

G). AMAZONAS: Sao Paulo de Olivenca, 11 Sep-26 Oct 

1936 (Q), Krukoff 8373 (A, F, NY, S, U). 

ducous. Staminate inflorescences up to 12 cm 

long and 6 cm wide; peduncle 1-3.5 cm long, 

peduncle and branches puberulous or partly hirtellous;flowers 

sessile, clustered in globose heads, 

these 2-3 mm diam.; tepals ca. 0.5-1 mm long, 

basally connate, hirtellous; filaments shorter than 

the tepals. Pistillate inflorescences branched, in 

fruit up to 8 cm long and 7 cm wide; peduncle 

3-4 cm long, indument or peduncle and branches 

similar to that of the staminate inflorescences; 

flowers 8-ca. 30; pedicel up to 0.3 cm long, in 

fruit up to 0.5 cm; perianth 0.3-0.4 cm long, 

(densely) hirsute-hirtellous, usually arachnoid 

hairs near the apex; stigma peltate, ca. 1.5 mm 

diam., hirtellous. Fruitingperianth purple, ovoid 

to ellipsoid, 1-1.5 cm long, puberulous, also 

sparsely yellow-hirsute, sometimes with brown, 

pluricellular hairs. 

Distribution (Fig. 74). Colombia (Amazonas), 

Peru (Loreto), and Brazil (Amazonas); in noninundated 

forest. 

This species, characterized by incised leaves 

and the smooth upper surface, but scabrous low- 

er surface of the lamina, may be related to P. 

bicolor. 

Ule collected staminate and pistillate speci- 

mens under the same number 5719. The sta- 

minate collection is selected as the lectotype col- 

lection. 

7. Pourouma myrmecophila Ducke, Bull. Mus. 

Hist. Nat. (Paris), Ser. 2, 4: 723. 1932. Type. 

Brazil. Amazonas: Manaus, Cachoeira do 

Mindui, 8 Jul 1929 (d), Ducke (RB) 23607 (lec- 

totype, RB, chosen here; isolectotypes, G, P, 

S, US). Fig. 71. 

Tree, up to 8 m tall, or shrub, myrmecophilous. 

Leafy twigs 0.6-1 cm thick, densely yellow(ish)-hirsute, 

glabrescent, and with reddishbrown, 

pluricellular hairs. Lamina 3-5-lobed, to 

-fid, 13-38 x 15-42 cm, subcoriaceous to chartaceous, 

rarely coriaceous, midsegment relatively 

long, apex acuminate, base subcordate to deeply 

cordate; upper surface scabrous, yellow-hirsute 

to -hirtellous on the main veins, elsewhere 

sparsely so or only hispidulous, lower surface 

pale yellow-hirtellous on the veins, white, arach- 

noid hairs in the areoles or sometimes also on 

the smaller veins; lateral veins 14-17 pairs, the 

basal pair usually branched, tertiary venation 

Specimens examined. COLOMBIA. AMAZONAS: 

Canto Aduche, 12 Nov 1981 (2), La Rotta 101 (herb. 

Araracuara), 4 Mar 1982 (2), La Rotta 139 (herb. Araracuara). 

PERU. LORETO: 13 Jan 1976 (2), Gentry et al. 15876 

(U); Puerto Almendras, 11 Nov 1984 (2), Maas et al. 

6249 (U), 13 Jan 1976 (2), McDaniel et al. 20436 (U); 

Rio Nanay, Puerto Almendras, 29 Nov 1976 (2), Revilla 

1966 (U). 

BRAZIL. AMAZONAS: Mun. Tefe, Lago de Tefe, 14 

Dec 1982 (8), Amaral et al. 76113 (BG); Rio Cuieras, 

mouth of Rio Branchinho, Sep 1973 (st), Berg 276 

(MO, U); upper Rio Alalau, 24 Feb 1968 (2), Bovan 

267 (INPA); BR-17, km 11, 4 Aug 1955 (6), Chagas 

(INPA) 1564 (INPA, U); Manaus-Mindfi rd., 3 Nov 

1955 (9), L. Coelho (INPA) 2265 (INPA); Mun. Jutai, 

Copessfi, nr. P6rto Alfonso, 24 Oct 1986 (2), Daly et 

al. 4129 (BG), 24 Oct 1986 (6), Daly et al. 4135 (BG); 

Manaus, Aleixo rd., 29 Dec 1942 (9), Ducke 1149 (MG, 

MO, NY, R, US), 23 Sep 1945 (6), Ducke 1764 (A, F, 

MG, NY, R, US), 19 Sep 1929 (2), Ducke (RB) 23606 

(RB, US); Manaus, Cachoeira do Mindu, 8 Jul 1929 

(8), Ducke (RB) 23607 (G, P, RB, S, lectotype collection 

of P. myrmecophila); Manaus, BR-17, km 3, without 

date (a), Francisco (INPA) 2186 (INPA); nr. P6rto Velho, 

8 Sep 1923 (a), Kuhlmann (RB) 19843 (B, U);


~~~~~~~~~~~~~~~~~~~~~~~~.. 

44deS A._ S 4 V 

?Cm Ira= BDT~OANWAL GARDENN 

VMVMW~T NACIONAL DE PESQUMA%A DA AMAZ6.%-I 

gftt. at A? .m 

-Gd , - v~~~~~~~~~~~~~T?e m e h. y tg 

' S Anto NAntni 

,24313 

x 22 d-.meterl de pAbonIariA ,M-.anaus-I 

Crcrintuits 220 aForest on terra 

Pouroum& 

? mateat 

S:nto Antonio de Abonari. 

brown. 

C rac&raj, Km 220. Forest 

I:anaur,an 

terra 

br&ct brown. 

Prne,RJ 

Negat've 

G.T. 

ime 

ol. vember 2619? 

iach 

L,F,~L.F Coelho- J.F. Ramosj_ ... _ .ln: 

.% '1??~ 1 1 1 55551 nr. C=ll 21 cr 31 | 51 wf *-ffi0st I.... 

rwr ar 

1w JtPAN n^ 

: 

21RK 3F1N41 

71i~ 

FIG. 71. Pourouma myrmecophila. LeafS twig with pistillate inflorescences (Prance et al. 24313). 

Manaus, Reserva Florestal Ducke, 6 Dec 1976 (st), 

Nascimento et al. (INPA) 66396 (INPA); Manaus-Itacoatiara 

rd., km 192, 13 Sep 1974 (6), Pennington et 

al. P.22647 (INPA, NY); Humaita-Labrea rd., km 80, 

27 Jan 1970 (9), Prance et al. 3253 (F, INPA, NY, R, 

151 

S, U, US); Rio Cuieras, mouth of Rio Brancinho, 24 

Sep 1971 (d), Prance et al. 14795 (F, INPA, NY, U); 

Rio Javari, Attalaia, 14 Oct 1976 (9), Prance et al. 

23776 (G, INPA, MG, NY, U); Manaus-Caracarai rd., 

km 220, Santa Antonio de Abonari, 26 Nov 1976 (2),


Prance et al. 24313 (INPA, MO, NY, S, U, US); Ma- perianth (sparsely) hirsute; stigma peltate, ca. 1.5 

naus-Itacoatiara rd., km 79, 1 Sep 1965 (8), Rodrigues cm diam., subhirsute. Fruiting perianth ovoid to 

et al. 7069 (U), km 74, 3 Sep 1965 (d), Rodrigues et 

al. 7076 (INPA). 

ellipsoid, ca. 1.5 cm long, blue-black, yellowhirsute. 

Vernacular names. Brazil. Amazonas: embau- Distribution (Fig. 74). Brazil (Amazonas); in 

barana or imbafbarana. 

non-inundated forest. 

From the syntype collections of P. myrmecophila, 

Ducke (RB) 23606 and 23607 and Kuhl- Specimens examined. BRAZIL. AMAZONAS: Mun. 

Sao Paulo de Olivenca, Rio Jandiatuba, 27 Nov 1986 

mann (RB) 19843, Ducke (RB) 23607, is chosen (9), Daly et al. 4456 (BG); nr. Sao Paulo de Olivenca, 

as the lectotype collection. 

5 Dec 1986 (8), Daly et al. 4485 (BG); Tonantins, Rio 

The collection Kuhlmann (RB) 150082, ac- Solim6es, 7 Feb 1944 (9), Ducke 1917 (F, NY, R, type 

cording to the label from Rio de Janeiro, is collection of P. 

prob- 

formicarum); Sao Antonia de Ia, 2 

ably either from Amazonas or was cultivated in 

May 1945 (8). Fr6es 20870 (NY, US), 19 Aug 1973 

(8). Lleras et al. P.17411 (F, INPA, MO, NY, S, U, 

Rio de Janeiro. 

US); Carauari, Poco Jurua, 7 Jul 1980 (8), A. S. Silva 

Vegetative parts (e.g., the wood) are (strongly) et al. 476 (BG, INPA). 

aromatic, smelling like spearmint or, according 

to other labels, like 

Vernacular names. Brazil. Amazonas: imbai- 

"bengue." 

binha, mapati. 

This species matches P. myrmecophila in the 

8. Pourouma formicarum Ducke, Trop. Woods swollen and saccate base of the petiole, a feature 

90: 9. 1947. Type. Brazil. Amazonas: Rio So- connected with myrmecophily. The two species 

limoes, Tonantins, 7 Feb 1944 (9), Ducke 1917 also show strong resemblances in the indument, 

(not 1916!) (holotype, R, not indicated by staminate inflorescences and flowers in the pis- 

Ducke, designated here; isotypes, F, NY). tillate flowers. They are quite distinct in the shape 

Fig. 72. of the lamina and in the length of the petiole. 

Tree, up to ca. 10 m 

"The 

tall, or 

branchlets are inhabited 

shrub, myrmeby 

stinging 'jicophilous. 

Leafy twigs ca. 6 mm quitaia' ants which nest 

thick, yellowamong 

the hairs" (Daly 

to 

et 

brownish-hirsute and 

al. 

with sparse, brown 4456). 

pluricellular 

hairs. Lamina entire, subovate, (14-) 18- 

32 x 6-12 

9. 

cm, chartaceous 


to subcoria- 

phaeotricha Mildbraed, Notizbl. 

Bot. Gart. Berlin-Dahlem 

ceous, apex abruptly short-acuminate, base 

10:193.1927. 

ob- 

Type. 

tuse to 

Peru. Loreto: 

subacute; upper surface scabrous to sca- 

Iquitos, 20 Aug 1925 (2), Tessmann 

5364 

bridulous, yellow-hirsute, lower surface + 

(holotype, B; isotypes, S, 

densely 

US). 

yellow-hirsute on the main veins to (very) sparse- 

Fig. 73. 

ly hirsute on the smaller veins, arachnoid hairs Tree, up to 10 m tall. Leafy twigs 4-6 mm 

in the areoles and on the smaller veins; lateral thick, yellow-sericeous to -hirtellous, and with 

veins 10-20 pairs, the basal pair unbranched, dense long, brown pluricellular hairs. Lamina 

tertiary venation slightly prominent beneath; entire, oblong to obovate, 8-23 x 3-9 cm, copetiole 

1-1.5 cm long, yellow-hirsute, at the base riaceous to subcoriaceous, apex acuminate, base 

swollen and saccate; stipules 3-5 cm long, outside acute to rounded; upper surface smooth or scayellow-hirsute, 

inside glabrous, caducous. Sta- brous, yellow-hirtellous on the main veins, lower 

minate inflorescences up to 12 cm long and to 6 surface yellow-hirsute on the veins and sparse or 

cm wide; peduncle 2-4 cm long, peduncle and sometimes dense, orange, pluricellular hairs, 

branches yellow-hirtellous to -hirsute; flowers arachnoid hairs confined to the areoles or almost 

sessile, clustered in globose heads, these 2-3 mm absent; lateral veins 8-10 pairs, basal pair unin 

diam.; tepals ca. 1 mm long, almost free, yel- branched, tertiary venation prominent beneath; 

lowish-hirtellous; filaments shorter than the te- petiole 2-6 cm long, indument similar to that of 

pals. Pistillate inflorescences branched, in fruit the twigs; stipules 1-4.5 cm long, outside yellowup 

to 7.5 cm long and 3.5 cm wide; peduncle ca. sericeous to -velutinous, inside sparsely yellow- 

4 cm long, peduncle and branches with indument hairy to glabrous, caducous. Staminate infloressimilar 

to that of the staminate inflorescences; cences up to 10 cm long and 6 cm wide; peduncle 

flowers 5-6; pedicel in fruit up to 0.5 cm long; 1.5-3 cm long, white- to yellow-sericeous to -ve-


FG 

R.: 

17. 411... 

. ur Uri? fr)i' -r : * 

If*,, r, , '*^>r ,, i /'.: 

X>.t 'i. E.< L: . ?J.1 W.C... 

. c O k ..y . 

penin * .l ei 

P Cul betLw. *. 

;i5 Behi m Ant r .i e J . 

72_.. Poru a'o1r1ma1m.11 1ea i 11g w h s i InfloFre"nce tiete. s e 

'__A 35 

212*. 

likl , 1 . -. 

i* ...... 

FIG. 72. Pouroumaformicarum. Leafy twig with staminate inflorescences (Lleras et al. P.17411). 

lutinous, and with dense (but on the ultimate 

branches sparse) brown, pluricellular hairs;flowers 

sessile, clustered in globose heads, these ca. 

2 mm diam.; tepals ca. 0.7 mm long, free or 

basally connate, hairy; filaments shorter than the 

petals. Pistillate inflorescences in fruit up to 9 cm 

long and 8 cm wide; peduncle 2-5 cm long, peduncle 

and branches yellow- to white-hirtellous 

to sericeous; flowers 11-15; pedicel 0.1-0.2 cm 

long, in fruit up to 0.9 cm; perianth ca. 2 mm


Pourouma phaeotricha 

HAtM OF PLRU 

mildbr. 

Maynas: Dtto. Iquitos. lio :lar.na, 

Caserio de 

IItjt 

Iishana trail to : 

|tg N ~ative i ?nch 1 2 de Yarana, uplands, I( :, 

"uvilla",. 

green. 

FIG. 73. Porio p ec fL. 

:- R~oX// r-5355S 1 I 21 31 41 5 Dececber 4, 1976 

/ . -,,,,,,,I,,,h,,,l.,,I,,,, II all C Manuel Rimachi Y. 2725 

FIG. 73. Pourouma phaeotricha. Leafy twigs with pistillate inflorescences 

(Rimachi Y. 2725).


long, yellow-velutinous, with long, brown, pluri- mostly also on the smaller veins, occasionally 

cellular hairs; stigma peltate, ca. 1.5 mm diam., also on the leaf margin; lateral veins (6-)10-25 

yellow-puberulous. Fruiting perianth ovoid to al- (-30) pairs, basal pair branched or sometimes (in 

most ellipsoid, 1.1-1.5 x 0.7-0.9 cm long, pu- obovate to elliptic leaves) unbranched, the basal 

berulous. 

part of these veins forming the basal part of the 

Distribution (Fig. 85). Peru (Loreto) and Brazil leaf margin (thus not separated from the margin 

(Amazonas, along Rio Javari); in non-inundated by mesophyll), reticulum and tertiary venation 

forest. 

(rather) plane beneath; petiole 3-22(-35) cm long, 

sparsely to densely white-puberulous (to -subve- 

Specimens examined. PERU. LORETO: Rio Nanay, 

Casario de Mishana 3 Dec 1976 (9), Davidson 5203 lutinous) to yellow-hirsute, occasionally also with 

(MO, U, US), 4 Dec 1976 (9), Rimachi 2725 (DUKE, long white, arachnoid hairs; stipules 3-16 cm long, 

F, NY); Iquitos, 3 Dec 1976 (9), Tessmann 5364 (B, outside yellow(ish)-(sub)velutinous to -hirsute to 

S, US, type collection of P. phaeotricha). -(sub)sericeous, inside glabrous or sometimes 

BRAZIL. AMAZONAS: Rio Javari, Estirao do Ecuador, 

8 Aug 1973 (8), Lleras et al. P.17242 (F, INPA, 

sparsely hairy, caducous. Staminate inflores- 

MO, NY, U, US), 21 Oct 1976 (v), Prance et al. 23966 cences up to 10 cm long and 5 cm wide; peduncle 

(G, INPA, NY, U). 

2-6 cm long, peduncle and branches whitish- to 

yellow-puberulous 

Vernacular names. Peru. Loreto: Sacha 

(to -subvelutionous) to -hiruvilla, 

tellous;flowers sessile or pedicellate, most of them 

uvilla. 

in 

The features of the staminate inflorescences 

globose heads, these 3-4 mm diam.; tepals 

0.5-1 mm 

and flowers and the pistillate flowers suggest close 

long, connate, forming a ? urceolate 

perianth, 

relationships to P. heterophylla and P. 

hairy; filaments longer than the periformicaanth, 

free. Pistillate 

rum. However, P. phaeotricha lacks the swollen 

inflorescences in fruit up to 

18 cm 

petiole base. 

long and 10 cm wide; peduncle 3-12 cm 

The two collections from the 

long, peduncle and branches with indument sim- 

Brazilian bank 

ilar to that of the staminate 

of Rio Javari have leaves with a scabrous 

inflorescence;flowers 

upper 

surface in 

10-30, in two clusters or more 

contrast to the other collections. 

diffusely distributed 

in the inflorescence; pedicel up to 1 cm, in 

fruit 

10. Pourouma mollis Trecul, Ann. Sci. Nat. 

up to 1.5 cm long, often already at anthesis 

Bot., with a 

Ser. 3, 8: 102, 1847; Miquel in Martius, Fl. 

strongly (+ capula-like) widened apex; 

bras. 4(1): 127. 1853; Berg, Fl. Suriname 

perianth 0.5-1 cm long, yellow(ish)-(sub)ve- 

5(1): 

272, t. 15b. 1975. Type. French Guiana. 

lutinous, 

Withapex 

? tuberculate; stigma peltate, ca. 

2 mm 

out locality, without date (2), Leprieur s.n. 

diam., white-puberulous. Fruiting peri- 

(lec- anth black or 

totype, P. chosen by Berg, Fl. Suriname 

purplish, ovoid to ellipsoid or to 

5(1): 

272. 

oblongoid, 1-2 x 0.7-1.5 cm, usually 

1975; 

densely 

isolectotype, G). 

hairy. 

This species is very closely related to P. melinonii. 

We had considered recognizing the two 

taxa only at the subspecific level, but the lack of 

intermediates in areas where the two taxa grow 

together indicates that they are probably good 

species. 

Two allopatric subspecies can be recognized. 


thick, rather sparsely to densely white-puberu- 

lous (to -subvelutinous) to yellow-hirsute, and 

with sparse to dense, purple to reddish-brown, 

pluricellular hairs, occasionally sparse, white, 

arachnoid hairs. Lamina entire, ovate to elliptic 

(or obovate), 6-25(-35) x 4-20(-30) cm or 3-5- 

lobed to -parted, ca. 10-25(-40) x 10-25(-40) 

cm, coriaceous (to subcoriaceous), apex acumi- 

nate, base acute, rounded to truncate, or sub- 

cordate; upper surface smooth, sometimes sca- 

brous, hairy on the main veins or subglabrous, 

occasionally yellow-hirsute, lower surface, 

sparsely to densely puberulous to hirtellous to 

subtomentose, often part of the hairs on the main 

veins appressed, arachnoid hairs in the areoles, 

Key to the Subspecies of 

Pourouma mollis 

1. Lamina usually entire; pistillate inflorescences 

with the flowers in two, more or less distinct 

clusters; Guianas, Lower Amazon Basin, and 

eastern Brazil. ................ a. subsp. mollis. 

1. Lamina usually 3-5 parted; pistillate inflorescences 

usually with the flowers diffusely distrib-


uted; Upper Amazon Basin to eastern Colombia. 

............................ b. subsp. triloba. 

lOa. Pourouma mollis Trecul subsp. mollis. 

Fig. 75b-f. 

Lamina usually entire, sometimes 3-lobed or 

to 5-parted, subjuvenile material usually 3-5- 

parted; upper surface smooth, with hairs on the 

main veins or subglabrous. Pistillateflowers ar- 

ranged in two + distinct clusters, due to the 

shortening of one (or two) of the primary branch- 

es and the short pedicels, being at fruit up to 0.7 

cm long. 

Distribution (Fig. 74). Eastern Guyana, Suri- 

nam, French Guiana, Amazonian Brazil 

(Amampa and Para), and eastern Brazil (Bahia 

and Espirito Santo); in non-inundated forest at 

low altitudes. 

1978 (6), DAF 027 (GUA); Reserva de Linhares, 30 

Oct 1930 (6), Kuhlmann 446 (U); Linhares, Corriogo 

de Durao, 2 Jan 1972 (9), Sucre 8303 (U). PARA: Faro, 

6 Jan 1920 (6), Ducke (RB) 13058 (RB); Obidos, 27 

Dec 1913 (9), Ducke (MG) 15261 (MG); Belem, 1 Feb 

1928 (6), Ducke (RB) 19455 (RB); Rio Guama; Sao 

Miguel, Oct 1906 (6), Goeldi (MG) 7731 (G, MG, S, 

U); Acarai Mts., Rio Taruini, 20 Nov 1952 (6), Guppy 

656 (=FD 7671) (NY, US); Belem, 19 Sep 1963 (6), 

Oliveira 2597 (IAN), 26 Sep 1963 (9), Oliveira 3071 

(IAN), 3 Sep 1966 (3), Pires 10123 (IAN), 24 Aug 1967 

(), Pires et al. 10578 (IAN), 2 Sep 1967 (6), Pires et 

al. 10809 (IAN), Sep-Oct 1961 (9), Pires 51676 (F, 

NY, US); Cuiaba-Santar6m rd., km 919,13 Nov 1977 

(9), Prance et al. 25337 (F, MG, RB, U); Mun. Almeirim, 

Monte Dourado, 2 Dec 1978 (6), Santos 455 

(U); Belem, 12 Jan 1966 (6), M. Silva 354 (MG), 4 Oct 

1942 (6), M. B. Silva 130 (NY, US). 

Vernacular names. Surinam. Boroma (Arowak), 

bospapaja, granboesipapaja, yarayara 

(Carib). French Guiana. Bois canon male, bois 

Specimens examined. GUYANA. Essequibo R., nr. 

canon sauvage. Brazil. Amapa: uva de macaco. 

Rockstone, 1 Aug 1921 (juv), Gleason 670 (GH, NY, Bahia: itararanga, tararanga, tararanga vermel- 

US), 4 Nov 1979 (2), Maas et al. 3938 (U). 

ha; Maranhao: ama'yrary (Ka'apor); Para: im- 

SURINAM. Peninica Creek, 20 Apr 1950 (st), BW bauiba, imbauibarana vermelha, mapatirana, ka- 

1128 (U); Sectie 0, 15 Sep 1916 (st), BW2410 (NY), 

U), 19 Dec 1919 (2), BW 4494 (NY, U), 25 Feb 1921 moyuwa (Wai-wai), sa-ouro (Mawayan). 

(2), BW5048 (U, UC), 15 Nov 1921 (2), BW 5440 In 

(A, Amapa and Para this subspecies can be con- 

RB, U, US), 28 Jan 1922 (9), BW 5581 (MO, U), 24 fused with a form of P. melinonii subsp. meli- 

Nov 1922 (2), BW 5992 (F, U); nr. Gansee, 26 May nonii with patent hairs on the lower leaf surface. 

1964 (juv), Donselaar 1373 (U); nr. Brownsberg, 14 

Oct 1964 (st), Donselaar 1681 (U), 4 

However, the latter has 

Jan 1966 

sparse, appressed hairs 

(juv), 

Donselaar 2924 (U); Gran Dam, 13 Oct 1966 (st), Donon 

the leafy twigs and stipules and the flowers in 

selaar 3688 (U); Mapane Creek, Feb 1964 (2), Elburg the pistillate inflorescence are diffusely distrib- 

(LBB) 9843 (MO, NY, U), 7 Mar 1963 (2), LBB 9424 uted, thus not in two clusters as in subsp. mollis. 

(U); Suriname R., without date (8 and 9), Hostmann Hostmann & Kappler 1272 

(& Kappler) 1272 

(6) belongs to P. 

(B, CGE, G, GH, LE, NY, OXF, P, 

S, U, mixed with P. 

melinonii 

melinonii subsp. melinonii); Nas- 

subsp. melinonii while Hostmann (& 

sau Mts., 16 Feb 1949 (9), Lanjouw et al. 2106 (NY, Kappler) 1272 (9 and 6) belongs to P. mollis subsp. 

U); Jodensavanne, 27 Mar 1953 (st), Lindeman 3631 mollis. 

(U), 16 May 1953 (st), Lindeman 3989 (U), 20 Dec The description of P. mollis was based on the 

1954 (st), Lindeman 6923 (U). 

collections Blanchet 2361 

FRENCH GUIANA. Massif des Emerillons, source 

(from Brazil, Bahia); 

of Approuage R., 19 Sep 1980 (8), Cremers 6726 

Hostmann 

(U); 

(from Suriname), and Leprieur 141. 

upper Crique Nouciri, 13 Dec 1983 (st), Cremers 8271 The latter was selected as the lectotype collection 

(U); without locality, without date (2), Leprieur s.n. (G, (Berg, 1975). 

P, lectotype collection of P. mollis); Orapu, 13 Oct 

1966 (9), Oldeman B.644 (CAY, P, U); piste de St. Elie, 

km 16, 19 Apr 1982 (st), Riera 573 (U). 

1Ob. Pourouma mollis Trecul subsp. triloba 

BRAZIL. Without locality, without date (a), Burchell (Trecul) C. C. Berg & van Heusden, Proc. Kon. 

9792 (GH, P, the same collection number is also used Ned. Akad. Wetensch., Ser. C, 91(2): 107. 1988. 

for a collection of P. guianensis subsp. guianensis). 

AMAPA: Serra do Navio, 1961 (st), Rodrigues 2984 Pourouma triloba Trecul, Ann. Sci. Nat. Bot., Ser. 3, 

(INPA); between P6rto Platon and Serra do Navio, 8:104. 1847 (Aug); Miquel in Martius, Fl. bras. 4(1): 

1976 (st), Rosa 1158 (MG). BAHIA: Without locality, 126. 1853; Macbride, Publ. Field Mus. Nat. Hist., 

without date (6), Blanchet 2361 (or 94) (FHO, G, LE, Bot. Ser., 13(2.2): 294. 1937. Type. Peru. Huanuco: 

P); P6rto Seguro, 20 Sep 1968 (6), Almeida et al. 71 Macora, without date (6!), Ruiz & Pavon s.n. (lec- 

(NY, U); Agua Preta, 23 Feb 1937 (9), Bondar 2169 totype P, chosen here; isolectotypes, G, OXF, US). 

(GUA); Rio das Contas, 3 Oct 1919 (8), Curran 21 (C, Pourouma jussiaeana Trecul, Ann. Sci. Nat. Bot., Ser. 

F, NY, US); Ilheus, 1848 (8), Luschnath s.n. (BR), Sep 3, 8: 106. 1847; Miquel in Martius, Fl. bras. 4(1): 

1821 (a), Riedel 2 (LE); Castelo Novo, 15 Sep 1944 (2), 124. 1853. Type. Peru. Buena Vista, without date 

Velloso 1083 (R). ESPiRITO SANTO: Linhares, 18 Aug (2), Anonymus s.n. (holotype, P).

.? 

P. cucura P. cuspidats 

GP. 7rorPcabrL P 

leo myrsoMaphile 

P. 

r~~~~~~~~~~~~~~~~~~~ 

subsp. 

tizab 

'' ? ~~~~ulSlsp. grlabrat 

~~~I 'I ~~~~Ubp. mebli inmii 

FIG. 74. Distribution of Pourouma cucura, P. cuspidata, P. formicarum, P. melinonii subsp. glabrata, P. 

melinonii subsp. melinonii, P. mollis subsp. mollis, P. mollis subsp. triloba, and P. myrmecophila.


~, 

a- 

e t~~~e7 f~~


Pourouma triloba Klotzsch, Linnaea 20: 526. 1847 

(Sep). Type. Peru. Huanuco: Macora, without date 

(9!), Ruiz & Pavon s.n. (holotype, B; isotypes, BR, 

G, OXF). 

Pourouma jaramilloi Cuatrecasas, Caldasia 7: 300. 

1956. Type. Colombia. Meta: Sierra de la Macarena, 

Cafno Ciervo, 19 Jan 1950 (2), Philipson & Jaramillo 

2133 (holotype, BM, not seen; isotypes, F, US). 

V. 10650 (BG). UYACALLI: Prov. Coronel Portillo, Pucallpa-Tingo 

Maria rd., 1980 (9), Trucios 9 (BG). 

BRAZIL. ACRE: Upper Rio Moa, nr. Fazenda Arizona, 

24-30 Sep 1984 (st), Campbell et al. 8383 (BG); 

mouth of Rio Macauhan, 5 Aug 1933 (d), Krukoff5309 

(F, G, GB, M, MO, NY, S, U, UC, US); Sena Madureira, 

Rio Caete, 7 Oct 1968 (d), Prance et al. 7907 (F, 

GH, NY, P, R, S, U, US). 

Lamina usually 3-5-parted, sometimes entire; Vernacular names. Peru. Amazonas: suia, suir 

upper surface sometimes scabrous; petiole oc- shuina (Huambisa), uhukamsuiya; San Martin: 

casionally with white-arachnoid hairs. Pistillate obija, uvilla. 

flowers usually more or less diffusely distributed This subspecies is more variable than subsp. 

in the inflorescence, sometimes in two ? distinct mollis and may prove to consist of several infraclusters; 

pedicels in fruit up to 1 cm long. specific taxa. In particular Huashikat 944, Hu- 

Distribution (Fig. 74). Colombia (Meta), Peru ashikat 1027, Tunqui 296 (all from Peru, Ama- 

(Amazonas, Loreto, Madre de Dios, Pasco and zonas, valley of Rio Santiago) and Gentry et al. 

San Martin), and Brazil (Acre); in non-inundated 15621 (from Loreto, Rio Nanay) are a distinct 


set of collections. They have entire leaves with 

a ? scabrous upper leaf surface, the basal lateral 

Specimens examined. COLOMBIA. META: San Juan veins are often 

de Arama, Rio Guejar, 26 Jan 1951 (9), Idrobo et al. 

unbranched, and the stipules are 

1316 (US); Sierra de la Macarena, Caino Ciervo, 19 Jan 

rather densely hairy inside. 

1950 (9), Philipson & Jaramillo 2133 (BM, P, US, type Prance et al. 7907 has relatively many lateral 

collection of P. jaramilloi). 

veins, up to ca. 30 pairs, whereas in other col- 

ECUDADOR. NAPO: Lumbaqui, 25 Nov 1987 (8), lections there are at most 25 pairs. 

Pennington et al. 12300 (BG). 

PERU. Without locality, without date 

Schunke V. 7729 and 10650, Foster 5791, D. 

(9), Anonymus 

s.n. (P, type collection of P. jussiaeana). AMAZONAS: N. Smith 5134, and Trucios 9 have entire leaves 

Rio Cenepa, 30 Dec 1972 (9), Berlin 746 (F, GH, MO); and the pistillate inflorescences tend to the type 

Rio Santiago, nr. Caterpiza, 15 Oct 1979 (9), Huashikat of inflorescence normal in subsp. mollis. 

944 (U), 24 Oct 1979 (2), Huashikat 1027 (BG); Rio The 

Huampi, 26 Jul 1974 (8), Kayap 1306 (MO, US); Rio 

description of P. triloba Tr6cul was based 

Santiago, nr. Caterpiza, 12 Dec 1979 (9), Tunqui 296 on both staminate and pistillate material collect- 

(BG). HUANUco: Tingo Maria, 13 Jul 1957 (st), Ellen- ed by Ruiz & Pavon, the staminate material is 

berg 2297 (U); Macora, without date (6), Ruiz & Pavon chosen as the lectotype collection, the pistillate 

s.n. (G, OXF, P, US, lectotype collection of P. triloba material has been used for the description of P. 

Tr6cul), without date (2), Ruiz & Pavon s.n. (B, BR, G, 

OXF, type collection of P. triloba 

triloba Klotzsch. 

Klotzsch); Pampayacu, 

5 Jun 1927 (d), Sawada 21 (F). JUNiN: Prov. Satipo, 

Gran Pajonal, S of Chequitaro, 22 Sep 1983 (8), D. N. 11. Pourouma melinonii Benoist, Bull. Mus. Hist. 

Smith 5134 (BG). LORETO: Prov. Maynas, Rio Nanay, Nat. (Paris) 28: 318. 1922; Berg, Fl. Suriname 

Puerto Almendras, 5 Jan 1976 (9), Gentry et al. 15621 

(BG); 22 km S of km 86 on Pucallpa-Tingo Maria 5(1): 274, t. 15a. 1975. Type. French Guiana. 

rd., 

11 Feb 1981 (2), Gentry et al. 31179 (BG); Prov. La 

May- Mana, Sep 1846 (9), Sagot 990 (lectotype, 

nas, Yanamono Explorama Tourist Camp, 27 Jun 1983 P. chosen by Berg, Fl. Suriname 5(1): 274.1975; 

(st), Gentry et al. 42232 (BG), 8 Jul 1983 (st), Gentry isolectotype, B). 

et al. 42793 (BG); Yurimaguas, Oct-Nov 1929 (6), LI. 

Williams 4688 (F). MADRE DE Dios: Rio Manu, Pak- Tree, up to 30 m tall. Leafy twigs 4-12 mm 

itza Station, 20 Nov 1980 (9), Foster 5781 (BG, F); thick, sparsely whitish, mostly appressed-puber- 

Rio Manu, Cocha Cashu Station, Jul 1978 (st), Foster ulous to 

et al. 6569 (F). SAN MARTIN: Prov. Mariscal 

strigose, and with purple to reddish- 

Caceres, 

Rio Uchiza, 24 Jul 1974 (a), J. Schunke V. 7729 (MO, 

brown, pluricellular hairs and occasionally sparse, 

NY, U); Prov. Mariscal Caceres, Dist. Tocache Nuevo, white, arachnoid hairs. Lamina entire, ovate (to 

Quebrada de Ischichini, 12 Oct 1978 (2), J. Schunke elliptic) or 3-lobed, 6-25(-35) x 4-16(-27) cm 

FIG. 75. a. Pourouma melinonii subsp. melinonii. Leafy twig with pistillate inflorescences (Irwin et al. 47832). 

b-f. P. mollis subsp. mollis. b. Leafy twig with pistillate inflorescences (Elburg (LBB) 9843). c. Staminate flower 

(Pires et al. 51625). d. Pistillate flower. e. Ovule f. Seed and embryo (BW 5581).


or 3-5-parted and ca. 10-27 x 10-27 cm, co- 


riaceous, apex acuminate, base acute, rounded 

Pourouma melinonii 

to truncate to subcordate or cordate; upper surface 

smooth, hairy on the main veins or subgla- 

1. Lamina usually entire, if 3-5-parted, then the 

base 

brous, lower surface usually truncate to subcordate; fruiting peri- 

(rather) sparsely appressed- anth sparsely or densely hairy; stamens usually 

puberulous to strigose or patent-puberulous to with free filaments; Amazon Basin and Guianas. 

hirtellous or subglabrous on the veins, arachnoid .......................... a. subsp. m elinonii. 

hairs in the areoles or also on the smaller veins; 1. Lamina 3-5-parted, base more or less deeply corlateral 

veins 12-20 pairs, basal pair branched, 

date; fruiting perianths subglabrous; stamens 

(often) with connate filaments; Panama and 

the basal part of these veins forming the basal northern and western Colombia .............. 

part of the leaf margin (thus not separated from ........................... b. subsp. glabrata. 

the margin by mesophyll), tertiary venation almost 

plane beneath; petiole 3-22 cm long, strilla. 

Pourouma melinonii Benoist 

gillose to strigose or puberulous to 

subsp. 

hirtellous; melinonii. 

stipules (2-)4-17 cm long, outside sparsely white- 

Fig. 75a. 

strigose, inside glabrous, caducous. Staminate Pourouma apaporiensis Cuatrecasas, Caldasia 7: 297. 

inflorescences up to 12 cm long and 12 cm wide; 1956. Type. Colombia. Amazonas-Vaupes: Rio 

peduncle 2-6 cm long, peduncle and branches Apaporis, Lagunas del Chucuro, 22 Nov 1951 (9), 

Garcia-Barriga 13643 (holotype, US; isotype, 

sparsely to rather 

NY). 

densely puberulous; flowers Pourouma apaporiensis Cuatrecasas forma macrosessile 

or pedicellate, most of them in globose phylla Cuatrecasas, Caldasia 7:298. 1956. Type. Coheads, 

these 3-4 mm diam.; tepals 0.5-1 mm lombia. Vaup6s: Rio Cubiyi, 8 Dec 1943 (9), Allen 

long, connate, forming a ? urceolate 3251 

perianth, (holotype, US, isotype, MO). 

sparsely to densely hairy; filaments longer than Leafy twigs hairy or subglabrous. Lamina 

the tepals, free or + connate. Pistillate inflores- mostly entire with a rounded to truncate base, 

cences in fruit up to 17 cm long and 11 cm wide; sometimes 3-lobed with a truncate to subcordate 

peduncle 3-12 cm long, peduncle and branches base, occasionally 3-parted with a deeply cordate 

with indument similar to that of the staminate base; lower surface on the veins ? sparsely hairy, 

inflorescence; flowers 10-30; pedicel up to 1 cm, mostly with appressed hairs, sometimes with 

in fruit up to 1.5 cm long; perianth 0.5-1 cm patent hairs. Stamens with (almost) free filalong, 

(rather) sparsely puberulous, apex tuber- ments. Fruiting perianth sparsely or + densely 

culate; stigma peltate, ca. 2 mm diam. Fruiting puberulous to subvelutinous. 

perianth dark violet or dark maroon, ovoid to Distribution (Fig. 74). Colombia (Amazonas), 

ellipsoid to oblongoid, 1-2 x 0.7-1.5 cm, sparse- Venezuela (Amazonas and Bolivar), Guyana, 

ly to densely puberulous to subvelutinous or Surinam, French Guiana, Peru (Amazonas and 

subglabrous. 

Loreto), and Brazil (Amapa, Amazonas, Mato 

This species is very closely related to P. mollis Grosso, and Para); in non-inundated forest at 

from which it is not always easy to separate. One low altitudes. 

of the main reasons for treating P. melinonii as 

distinct at the species level is the co-occurrence Specimens examined. COLOMBIA. AMAZONAS: Rio 

of P. mollis subsp. mollis and P. melinonii subsp. Apaporis, between Rio Pacoa and Rio Kananari, 15- 

19 Dec 1951 

melinonii in the Guianas without indications of 

(9), Garcia-Barriga 14116 (NY, US). 

AMAZONAS-VAUPES: Rio Apaporis, Lagunas del Chuinterbreeding. 

P. melinonii can be distinguished ruco, 22 Nov 1951 (9), Garcia-Barriga 13643 (NY, US, 

from P. mollis by the sparse indument in most type collection of P. apaporiensis). VAUPES: Cabeceras 

parts of the plant, but especially on the stipules; Rio Cubyyfi, 8 Dec 1943 (9), Allen 3251 (MO, type 

only the fruiting perianth is sometimes 

collection of P. 

densely 

apaporiensis Forma macrophylla). 

VENEZUELA. AMAZONAS: 

puberulous to subvelutinous. The sparse indu- 

Upper Rio Orinoco, 

1951 (9), Croizat 962 (US); base of Cerro Duida, Janment 

on the stipules and leafy twigs of P. meli- Feb 1969 (st), Farinas et al. 490 (U, VEN); Rio Cainonii 

is whitish and usually appressed, whereas quiare, Rio Vaciva and Rio Yatua, Apr 1853 (6), Spruce 

the dense indument on the stipules and 3176 

leafy (C, CGE, E, GH, GOET, MG, NY, OXF, P). 

twigs of P. mollis is yellow and 

BOLiVAR: Rio Chirea Uriman, 5 

usually + 

Aug 1953 

patent. 

(st), Bernardi 

902 (NY, VEN); Rio Icabaru and Rio Hacha, 7 

Within P. melinonii two allopatric subspecies Jan 1956 (9), Bernardi 2808 (G, NY); Mun. Urdaneta, 

can be recognized. 

Rio Chinarok, 10 Jan 1986 (9), Hernandez 317 (BG);


between Rio Chicanan and Rio Ayaiche, 24 Aug 1961 (Arawak), bospapaja, granboesipapaja, poeroe- 

(6), Steyermark 89452 (NY, VEN); Sierra Ichun, Salto ma or 

Maria Espuma, 29 Dec 1961 

puruma (Carib), 

(6), Steyermark 90408 

yarayara (Carib). French 

(NY); El Tigre, 2 May 1939 (9), LI. Williams 12008 

Guiana. Bois canon, papaye apici (Paramaka). 

(F, S, US, VEN). 

Peru. Amazonas: dakamtazshuiya, mimpashui- 

GUYANA. Pakaraima Mts., Kamana falls, 25 Aug ya washi shuina; Loreto: uvilla. Brazil. Amazo- 

1961 (6), Maguire et al. 45945 A (NY, US); upper nas: imbaibarana; Para: mapatirana. 

Mazaruni R., Kako R., 23 Sep 1960 (6), Tillett et al. The collection from Amazonian 

45495 Colombia, in- 

(U). 

SURINAM. Marowijne R., without date (6), (Host- cluding the type collection of P. apaporiensis, 

mann &) Kappler 1272 (LE, M, P); Grote Zwiebel- have deeply cordate leaves, like the material of 

zwamp, 22 Sep 1948 (2), Lanjouw et al. 399 (NY, U); subsp. glabrata, but in contrast to the other col- 

Jodensavanne, 4 Jul 1961 (st), Schulz (LBB) 8976 (F, lections of subsp. melinonii. Pistillate specimens 

U). 

FRENCH GUIANA. Without locality, without data 

from this area can be distinguished from those 

(2), Aublet s.n. (BM, S), 1986 (6), Melinon 457 (A, G, of subsp. glabrata in the dense indument of the 

P, U), 1863 (2), Melinon s.n. (P), 1863 (6), Melinon s.n. perianth of the pistillate flower. 

(E, LE, NY, P, US), (8), Poiteau s.n. (LE), 1865 (st) The description of P. melinonii was based on 

Sagot s.n. (P); Bolatee Creek, 2 Nov 1948 (st), BAFOG 

the collections 

4272 (CAY, U);'St. Pierre Creek, 28 Feb 1959 Kappler 1272 (from 

(st), 

Suriname) 

Cantonnement de Cayenne s.n. (CAY); Trois Sauts, 28 and three collections made in French Guiana: 

Oct 1974 (2), Lescure 343 (CAY); Cayenne, without Sagot 517 (p.p.), Sagot 990, and Melinon 457. 

date (2), Martin s.n. (BR, F, MO, U); Cacao, S of Cay- Sagot 990 has been selected as the lectotype colenne, 

21 Apr 1965 (6), Oldeman 1243 (CAY); piste de lection 

St. Elie, 1 Jun 1981 (st), Sabatier 51 (Berg, 1975). The (Hostmann &) Kappler 

(CAY); Acarouany, 

May 1858 (6), Sagot 517 (F, P; under the same number collection has the same number as a collection 

the lectotype collection of P. laevis); La Mana, Sep 1856 of Hostmann (& Kappler) of P. mollis subsp. mol- 

(2), Sagot 990 (B, P, lectotype collection of P. meli- lis. 

nonii), 1856 (2), Sagot 990bis (G); Godebert, 13 Jul 

1921 (2), Wachenheim 232 and s.n. (P). 

PERU. AMAZONAS: Monte Chichijam, 24 Apr 1973 lib. Pourouma melinonii Benoist subsp. glabra- 

(2), Ancuash 290 (F, GH, MO); Rio Santiago, nr.. Ca- ta C. C. Berg & van Heusden, Proc. Kon. Ned. 

terpiza, 7 Sep 1977 (2), Huashikat 431 (BG); Rio Cusu, Akad. Wetensch., Ser. C, 91(2): 108. 1988. 

Yucui, 12 Mar 1973 (2), Kayap 571 (GH). LORETO: 

Rio Tacsha, Curaray, 19 Sep 1972 (6), Croat 20447 

Type. Panama. Canal Zone: Pipeline Rd., 10 

(C, km NW of 

DUKE, F, GH, MO, NY). 

Gamboa, 14 Dec 1973 (9), Berg & 

BRAZIL. AMAPA: Rio Jaue, 3 km E of confluence Nee 355 (holotype, U; isotypes, BG, COL, K, 

with Rio Oiapoque, 26 Aug 1960 (2), Irwin et al. 47832 MO, NY, PMA). Fig. 76. 

(GH, NY, U, US); Rio Mutura, 22 Sep 1960 (2), Irwin 

et al. 48439 (NY, U, US); Mun. Mazagao, 81 km NW Leafy twigs subglabrous. Lamina usually 5- or 

of Macapa, 20 Dec 1984 (2), Mori et al. 17478 (BG). 3-parted with a distinctly cordate base, some- 

AMAZONAS: Mun. Tefe, Rio Solimoes, Lago de Tefe, times entire and then with a cordate to subacute 

15 Oct 1982 (2), Amaral et al. 101 (BG); Rio Japura, 

Vila Bittancourt, 19 Nov 1982 (2), Amaral et al. 595 base; lower surface on the main veins subgla- 

(BG); Manaus-Caracarai rd., km 148, 27 Sep 1973 (2), brous. Stamens often with + connate filaments. 

Berg et al. P. 18136 (F, INPA, MO, NY, P, S, U, US); Fruiting perianth (sub)glabrous. 

Mun. Anori, rd. Amori-Anama, 27 Nov 1975 (8), D. Distribution (Fig. 74). Panama and Colombia 

Coelho et al. 672 (INPA); Rio Solimoes, Belem, 16 

Dec 1948 (6), Fr6es 23725 (IAN), 25 Mar 1976 (Antioquia); in non-inundated forest at low al- 

(st), 

Mello s.n. (INPA); Manaus-Itacoatiara rd., km 26, 2 titudes. 

Sep 1966 (6), Prance et al. 2180 (F, GH, INPA, NY, 

P, R, S, U, US). MATO GROSSO: Brasilia-Acre Specimens examined. PANAMA. CANAL ZONE: 

hwy., 

215 km, west of Rio Juruena, 2 Sep 1963 Pipeline Rd., ca. 10 km NW of Gamboa, 14 Dec 1973 

(6), Maguire 

et al. 56511 (F, NY, RB, U, US). PARA: Rio (9), Berg & Nee 355 (BG, COL, K, MO, NY, PMA, 

Guama, 

U, 

Sao Miguel, Oct 1906 (2), Goeldi (MG) 7732 type collection of P. melinonii subsp. 

(G, 

glabrata), 12 Oct 

MG); 

Rio 

1971 

Jari, Monte Dourado, 16 Nov 1967 (2), Oliveira (9), Gentry 2046 (MO); Gamboa, 21 Nov 1971 

3736 (IAN), 22 Jan 1968 (2), Oliveira 3936 (NY), 30 

(9), Gentry 2657 (MO, NA). DARIEN: S of El Real, nr. 

Cana 

Jan 1961 (9), Oliveira 4047 (NY), 9 Sep 1968 (2), N. 

mine, 21 Aug 1987 (9), McPherson 11510 (BG). 

SAN BLAS: 

T. Silva 921 

Rio 

(NY, U), 11 Sep 1968 (6); N. T. Silva 949 

Cangandi, nr. Cangandi, 18 Feb 1984 

(NY, U), 16 Sep 1968 (2) N. T. Silva 995 (9), Nevers 4929 (BG). 

(IAN, NY). COLOMBIA. ANTIOQUIA: 38 km W of Barrancabermeja, 

24 Feb 1967 (8), Bruijn 1496 (F, M, MO, NY, 

Vernacular names. Venezuela. Bolivar: kai- S, U, US, VEN), 25 Feb 1967 (9), Bruijn 1502 (F, M, 

warikai, yagrumo-sunsun. Surinam. Boroma MO, NY, S, U, US, VEN), 1 Mar 1967 (st), Bruijn

162 

1546 (F, M, MO, NY, S, U, US, VEN), 2 Feb 1967 

(st), Bruijn 1555 (F, M, MO, NY, S, U, US, VEN); 

Anori, 6-12 Sep 1973 (9), Soejarto 4283 (MO). 

Vernacular names. Colombia. Antioquia: cirpe 

macho. 

12. Pourouma hirsutipetiolata Mildbraed, No- 

tizbl. Bot. Gart. Berlin-Dahlem 10: 420. 1928. 

Type. Colombia. Bolivar: Norosi-Tiquiso trail, 

lands of Loba, Apr-May 1916 (Q), Curran 116 

(holotype, US, fragment in B). 

Tree, up to ca. 30 m tall, with butresses. Leafy 

twigs 6-20 mm thick, yellow-hirsute and with 

minute, whitish, appressed or patent hairs and 

usually also with brown, pluricellular hairs. 

Lamina entire, ovate (to elliptic or suborbicular), 

10-25 x 7-18 cm, or 3-lobed to -parted and ca. 

10-35 x 10-35 cm, coriaceous, apex acuminate, 

base cordate or subcordate or sometimes rounded; 

upper surface smooth, hairy on the main veins 

or subglabrous, lower surface (rather) sparsely 

strigose to strigillose on the veins or puberulous 

to hirtellous on the smaller veins, arachnoid hairs 

in the areoles, usually extending to the lateral 

veins; lateral veins (5-)12-15 pairs, basal pair 

branched, tertiary and quarterary venation 

rather prominent beneath; petiole 3-20 cm long, 

yellow-hirsute, usually also with minute, whitish 

hairs; stipules 4-24 cm long, outside yellow-hirsute 

(to -subsericeous), and also with much 

shorter, whitish hairs, inside glabrous, caducous. 

Staminate inflorescences up to 10 cm long and 

8 cm wide; peduncle 3-4 cm long, densely pub- 


leafy twigs, and the more prominent smaller veins 

at the lamina beneath, and the broader segments 

of the incised leaves. It differs from P. mollis in 

the cordate leaf base of the incised leaves and 

the more prominent smaller veins on the lamina 

beneath. 

Two (probably) allopatric subspecies can be 

recognized. 


Pourouma hirsutipetiolata 

1. Lamina entire; northern Colombia. .......... 

.................... a. subsp. hirsutipetiolata. 

1. Lamina 3-lobed to -parted; western Colombia 

and Ecuador. ................ b. subsp. hispida. 

12a. Pourouma hirsutipetiolata Mildbraed subsp. 

hirsutipetiolata. Fig. 77. 

Lamina entire. 

Distribution (Fig. 81). Colombia (Antioquia, 

Bolivar, and Santander); in non-inundated, lowland 

forest. 

Specimens examined. COLOMBIA. ANTIOQUIA: 38 

km W ofBarrancabermeja, 1 Mar 1967 (9), Bruijn 1547 

(F, M, MO, NY, S, U, US, VEN). BOLIVAR: Norosi- 

Tiquisio trail, Lands of Loba, Apr-May 1916 (2), Curran 

116 (B, US, type collection of P. hirsutipetiolata). 

SANTANDER: Bucaramanga, 11 Dec 1977 (2), Renteria 

et al. 45 (MO); rd. to Citimarra, 9 Dec 1979 (2), Renteria 

et al. 2093 (HUA); 16 km SE of Barrancabermeja, 

26 Aug 1954 (9), Romero-Castaneda 4715 (MO, US). 

Vernacular names. Colombia. Santander: sirpe 

erulous or also hirtellous; flowers sessile, most of or sirpo, urumo blanco. 

them in subglobose heads, these ca. 3 mm diam.; 

tepals 1-1.5 mm long, connate, forming a 12b. Pourouma hirsutipetiolata Mildbraed subsp. 

(sub)infundibuliform perianth, often with an hispida (Standley & Cuatrecasas) C. C. Berg & 

oblique apex, sparsely hairy; filaments longer than van Heusden, Proc. Kon. Ned. Akad. Wethe 

tepals, free. Pistillate inflorescences in fruit tensch., Ser. C, 91(2): 108. 1988. Fig. 78. 

up to 22 cm long and 10 cm wide; peduncle 5- Pourouma hispida Cuatrecasas, Standley & Cuatreca- 

13 cm long; peduncle and branches minutely sas in Cuatrecasas, Caldasia 7: 299. 1956. Type. Copuberulous, 

sparsely so on the peduncle, more lombia. Valle: Rio Anchicaya, La Planta, 5 Aug 1943 

densely on the branches and very densely on the (9), Cuatrecasas 14881 (holotype, F). 

pedicels; flowers 7-17; pedicel 0.5-1 cm long, in Lamina 3-lobed to -parted. 

fruit up to 1.5 cm long; perianth 5-10 cm long, Distribution (Fig. 81). Colombia (Choc6, Valle) 

densely short-velutinous; stigma peltate, ca. 1.5 and Ecuador (Esmeraldas); in non-inundated, 

mm diam., white puberulous. Fruiting perianth lowland forest. 

dark violet or black, ellipsoid to ovoid, ca. 1.5- 

2 x 1-1.5 cm, densely minutely puberulous. Specimens examined. COLOMBIA. CHocO: La Mo- 

This species is closely related to P. jarra, 5 Nov 1983 (2), Juncosa 1253 (BG). VALLE: Rio 

melinonii, 

Anchicaya, La Planta, 5 Aug 1943 (2), Cuatrecasas 

from which it differs in the shape of the staminate 14881 (F, type collection of P. hispida); Rio Calima, 

flowers, the long yellow hairs on the petioles and La Trojita, 19 Feb-10 Mar 1944 (9), Cuatrecasas 16826


FIG. 

FLORA 76 Prom mn b gbith pitt inre PANAMENSIS Br 

b*. . . , ., 

A _l I - p , 10 . I w W 

rrrO p _ f-nTt. 

; 

- 

^ 

. r 

^I 

- ^ , O^ t a r r o l e . .,, ^ V4 

]..p 

2m 20 - x ? 30 * I with L n_5 ;il1t Sotwo MP 5 tul bl z 

' 

I.-it^^^;.;'' *-i FrtouM mal.nrt^.?, Bf??.ft [,..; C Berg i * , 

i^. ?: .- 

'Fu.: C.C. arit o llreah twig t 

i. 

*'*^ 'AM9 i^ fe tr ^"b-t* cc*B"^*t'a"*l*)^ - 

FIG. 76. Pourouma melinonii subsp. glabrata. Leafy twig with pistillate inflorescences (Berg et al. 355). 

(F); Rio Dagua, between Las Cascadas and Alto de 

Yundo, 7 Jul 1984 (9), Gentry et al. 48341 (BG). 

ECUADOR. ESMERALDAS: Rio Pambil, Estero Bravo, 

6 Jul 1966 (d), Jdtiva 303 (NY), Jdtiva et al. 1079 

(NY, UC, US); Panadero, 5 km from the river, 26 Jul 

1967 (d), Jdtiva et al. 2033 (MO, NY, US), 26 Jul 1967 

(9), Jdtiva et al. 2037 (NY, U, UC, US); Rio Hoja 

Blanca and Rio Hualpi, 15 Sep 1965 (d), Little et al. 

21070 (F, MO, NY, US); Rio Guayllabamba, Quinin- 

d6, 4 Oct 1965 (d), Little et al. 21225 (NY, US). 

Vernacular names. Ecuador. Esmeraldas: uva.


INSTIIWJO FORESAL ULATIMO AMERICANO 

1I|e1 D . att 1eS s U | I A 

h 

1g|nc i :.. 

.. 2 ..:. 

-. . 

11, 2 _34 5 ta 

. 

rir, .oula 3s t 

. . . . 

.... *.., -, . ? 

UNITED STATES ;' - ; 

Pouriou hirauriprt iolata Mtldbread subap. 

*!37'0h3 hr*,rup,,-tolat ... . -. 

NToAL Hn UI De. C . . Berg Utrech IatmD1l 1986 m1a 

ilo bm 

NATiONAL HERSA1IU E.C.K. van Keusden 

FIG. 77. Pourouma hirsutipetiolata subsp. hirsutipetiolata. Leafy twig with pistillate inflorescences (Bruijn 

1547). 

13. Pourouma tomentosa Martius ex Miquel in 

Martius, Fl. bras. 4(1): 128, t. 38. 1853. Type. 

Colombia. Amazonas or Caqueta: Araracuara, 

Dec 1819 (Q), Martius s.n. (lectotype, M, cho- 

sen here; isolectotype, U). 

Tree, up to ca. 30 m tall. Leafy twigs 4-15 mm 

thick, white-puberulous or partly also yellow- 

hirsute, often also with white, arachnoid hairs, 

pluricellular hairs lacking or very sparse. Lamina 

entire, ovate to elliptic (to oblong), (4-)10-20 

(-40) x (3-)5-15(-27) cm or 3-5-lobed to -parted 

or 5-9-parted and ca. 10-35 x 10-35 cm, coriaceous, 

apex acuminate, rounded or sometimes 

emarginate, base acute to obtuse, rounded, truncate, 

or cordate; upper surface smooth, sometimes 

scabrous, hairy on the main veins or only


m.m,--- 

* _ . 1 2'. A . .. . . ; 

FIG. 78. Pourouma hirsutipetiolata subsp. hispida. Leafy twig with staminate inflorescences (Little e a. 

21225). 

on the midrib, in scabrous leaves hispidulous 

over the whole surface, lower surface rather 

sparsely appressed-puberulous to strigose on the 

(main) veins, arachnoid hairs in the areoles, usually 

extending to the midrib, but on the main 

veins ? disappearing with age; lateral veins 7- 

25 pairs, the basal pair branched or sometimes 

(in elliptic leaves) unbranched, the basal part of 

these veins forming the basal part of the leaf 

margin (thus not separated from the margin by 

mesophyll), tertiary (and quarternary) veins often 

rather prominent beneath; petiole 2-26 cm long, 

white-puberulous or yellow-(sub)hirsute, and 

usually also white, arachnoid hairs; stipules (2-)6- 

18 cm long, outside whitish-strigose to yellowhirsute 

or -hirtellous, and with dense to sparse,


white, arachnoid hairs, usually also pluricellular This species is closely related to P. mollis and 

hairs, inside glabrous or with dense or sparse, P. melinonii. It is distinguishable by the occuryellow(ish) 

hairs, caducous. Staminate inflores- rence of ? dense, arachnoid hairs on the leafy 

cences up to 12 cm long and 12 cm wide; pe- twigs, stipules, petioles, main veins at the lower 

duncle 2-5 cm long, peduncle and branches + leaf surface and/or the inflorescences, and, moredensely 

puberulous, often also with white, arach- over, by the lack or scarcity of pluricellular hairs 

noid hairs; flowers sessile or pedicellate, most of on the leafy twigs. 

them in (sub)globose heads, these 3-5 mm diam.; Five subspecies, partly allopatric, can be rectepals 

0.5-1 mm long, connate, forming a more ognized. A few specimens from Peru (Amazonas) 

or less distinctly urceolate perianth, sparsely to cannot be satisfactoraly inserted in one of these 

densely hairy; filaments longer than the tepals, subspecies. They may represent another subspefree. 

Pistillate inflorescences in fruit up to 20 cm cies. These collections (Kayap 201, Tunqui 217, 

long and 13 cm wide; peduncle 2-13 cm long, Huashikat 966, 1432 and 1482) largely match 

peduncle and branched sparsely to densely pub- the collections under subsp. tomentosa, but the 

erulous or also yellow-(sub)hirsute, often with arachnoid indument is almost lacking on most 

white, arachnoid hairs; flowers pedicellate (or parts, except for the main veins of the lamina 

subsessile), pedicel up to 0.5 cm long, in fruit up beneath and the petioles. Moreover, the hairs on 

to 1.5 cm long; perianth 5-10 mm long, mostly the fruiting perianth are much shorter, and the 

densely, yellowish, short-velutinous, sometimes apex of the leaves is short-acuminate. They are 

sparsely puberulous, or also with white, arach- provisionally placed under subsp. tomentosa. 

noid hairs; stigma (sub)peltate, ca. 1.5 mm diam. From the two syntype collections made by 

Fruiting perianth black or purple, oblongoid to Martius, one is chosen as the lectotype collection. 

ellipsoid to ovoid, 1.5-2 x 0.8-1.5 cm, mostly 

densely, yellowish-puberulous to -velutinous or 

-subhirtellous. 

Key to the Subspecies of Pourouma tomentosa 

1. Stipules hairy inside. .................................. ... ................. b. subsp. apiculata. 


2. Leafy twigs yellow-hirsute (or glabrous); lamina 5-9-parted. ..................... c. subsp. persecta. 

2. Leafy twigs puberulous or hirtellous; lamina entire or 3-5-parted. 

3. Base of the lamina more or less deeply cordate; lamina 3-5-parted. ...... d. subsp. essequiboensis. 

3. Base of the lamina rounded to truncate or subcordate; lamina usually entire. 

4. Lamina 3-5-fid to -parted ............................................ a. subsp. tomentosa. 


5. Apex of the lamina rounded to emarginate, (or short-acuminate in aberrant specimens 

from Peru, Amazonas); heads of the staminate inflorescence 4-5 mm diam.; upper Amazon 

Basin. .......... ................................. a. subsp. tomentosa. 

5. Apex of the lamina acuminate; heads of the staminate inflorescence 2-3 mm diam.; 

Guianas, Amapa, and near Manaus. ............................. e. subsp. maroniensis. 

13a. Pourouma tomentosa Miquel subsp. 

tomentosa. Fig. 79. 

Pourouma albistipulata Cuatrecasas, Bol. Soc. Venez. 

Ci. Nat. 17: 92. 1956. Type. Venezuela. Amazonas: 

Rio Guainia, Victorino, 22 Mar 1942 (9), LI. Williams 

14833 (holotype, F; isotypes, F, G, NY, RB, 

S, US, VEN). 

Leafy twigs and petioles white-puberulous. 

Lamina entire, elliptic to ovate, sometimes (in 

Colombia and Ecuador) 3- or 5-parted, if entire, 

then often thickly coriaceous, + plicate, and apex 

rounded or emarginate (or sometimes short-acuminate), 

base rounded to truncate, or (in incised 

leaves) to subcordate; upper surface smooth; stipules 

outside with dense, white-arachnoid hairs, 

inside glabrous. Heads of the staminate inflorescences 

ca. 4-6 mm in diam. Pistillate flowers + 

diffusely distributed in the inflorescence, sometimes 

clustered in two groups; peduncle mostly 

up to 6(-7) cm long; perianth without dense, 

white, arachnoid indument, in fruit yellowish 

short-velutinous.

+ AJ ++ 

P O R O ULMA tomentosa. 

FIG. 79. Pourouma tomentosa subsp. tomentosa. From Martius, Flora Brasiliensis 4(1). 1853: tab. 38. Leafy twig with 

section of pistillate flower (17), stigma (13), pericarp (19), seed (21), cotyledons (24).

168 

Distribution (Fig. 81). The north-western part 

of the Upper Amazon Basin; in non-inundated 

forest, at low altitudes but in Peru (Amazonas) 

up to 2000 m. 

Specimens examined. COLOMBIA. AMAZO- 

NAS-CAQUETA: Araracuara, Dec 1819 (9), Martius s.n. 

(M, U, lectotype collection of P. tomentosa), Feb 1820 

(st), Martius s.n. (M, U). VAUPES?: Lagos del Paso, 25 

Dec 1975 (6), Roa 252 (INPA). 

VENEZUELA. AMAZONAS: Conuco, 2 Dec 1977 

(st), Liesner 4105 (U); Cerro Duida, 23 Aug 1944 (st), 

Steyermark 57872 (F); Cerro Yapacana, 3 May 1970 

(st), Steyermark et al. 103026 (U, Ven); Victorina, Rio 

Guainia, 22 Mar 1942 (9), LI. Williams 14833 (G, NY, 

RB, US, VEN, type collection of P. albistipulata). 

ECUADOR. NAPO: 8 km SE of Tena, 30 Sep 1960 

(9), Grubb et al. 1681 (NY); Parque Nacional Yasuni, 

9-19 Jan 1988 (9), Neill et al. 8158 (BG). 

Vernacular names. Colombia. Chiricaba (Kar- 

ijona), guaumoutognac (Tukano); Amazonas- 

Vaupes: ambauiba do vinho. Venezuela. Ama- 

zonas: cocora, cocora montanero, cucura. Peru. 

Amazonas: paiu shuiya (Huambisa), paui shuina; 

Loreto: sacha uvilla. Brazil. Amazonas: imbaiba 

(or ambauva) do vinho. 

Neill et al. 8158 (from Ecuador) and Rao 252 

(from Colombia) differ from the other collections 

in the 3- or 5-parted lamina, but match in other 

features subsp. tomentosa. Grubb et al. 1681 (from 

Ecuador) has 5-parted leaves and the postillate 

flowers clustered as in subsp. apiculata, but the 

yellow hairs normally occurring on the leaves of 

that subspecies are wanting. Gentry et al. 22865 

(from Peru, Amazonas, 1800-1950 m!) is another 

collection which could not be placed satisfacto- 

rily. It matches subsp. tomentosa, except for the 

yellow-hirsute leafy twigs and stipules. 


See also unnamed collections under number 2 

(p. 193). 

13b. Pourouma tomentosa Miquel subsp. api- 

culata (Benoist) C. C. Berg & van Heusden, 

Proc. Kon. Ned. Akad. Wetensch., Ser. C, 

91(2): 108. 1988. 

Pourouma apiculata Spruce ex Benoist, Bull. Mus. Hist. 

Nat. (Paris) 28: 321. 1922. Type. Brazil. Amazonas: 

Rio Uaupes, nr. Panur6 (=Ipanor6), Oct 1852-Jan 

1853 (Q), Spruce 2865 (holotype, P; isotypes, B, BR, 

C, CGE, E, F, G, GH, GOET, LE, MG, NY, OXF, 

P, US). 

Pourouma apiculata Spruce ex Mildbraed, Notzbl. Bot. 

Gart. Berlin-Dahlem 10: 419. 1928. Based on the 

specimen of Spruce 2865 in B. 

Coussapoa krukovii Standley, Publ. Field Mus. Nat. 

PERU. AMAZONAS: 12-18 km on trail E of La Peca Hist., Bot. Ser., 17: 163. 1937. Type. Brazil. Amain 

Serrania de Bagua, 1800-1950 m, 14 Jun 1978 (8), zonas: Mun. Sao Paulo de Olivenca, 11 Sep-26 Oct 

Gentry et al. 22865 (BG); Rio Santiago, Caterpiza, 17 1936 (a), Krukoff 8223 (holotype, NY; isotypes, A, 

Oct 1979 (9), Huashikat 966 (BG), 4 Dec 1979 (9), F, G, GH, LE, MO, P, S, U, US). 

Huashikat 1432 (U), 10 Dec 1979 (9), Huashikat 1482 

(BG); Rio Cenepa, Quebrack Yutai-entsa, 22 Jan 1973 Leafy twigs yellow-hirsute (sometimes only on 

(9), Kayap 201 (F, GH, NO); Rio Santiago, Quebrada the scars of the stipules), this indument often 

Caterpiza, 3 Dec 1979 (9), Tunqui 217 (BG). LORETO: extended to the petioles. Lamina usually entire 

Prov. Requena, Arboretum Jenaro Herrera, Aug-Sep and ovate, sometimes 3-lobed or (in 

1976 

subjuvenile 

(6), Bernardi s.n. (arbre 6/27) (U); Prov. Maynas, 

Dist. Iquitos, Zungaro Cocha, 13 Dec 1967 (9), Reyna material?) 3-5-parted, apex usually acuminate, 

R. 37 (BG, F). 

base broadly acute to truncate or (in subjuvenile 

BRAZIL. AMAZONAS: Tefe, 25 Sep 1940 (st), Black material?) occasionally shallowly cordate; upper 

47-1520 (IAN); Manaus-Itacoatiara road, km 26 (st), surface smooth or scabrous; stipules outside with 

L. Coelho (INPA) 5223 (INPA); 5 Jun 1976 (st), Haroldo 

(INPA) 57512 (INPA), 26 Oct 1965 (6), Loureiro (rather) dense, white arachnoid hairs, inside with 

(INPA) 16461 (INPA), 19 Mar 1976 (st), Mello (INPA) yellow hairs. Heads of the staminate inflores- 

55388 (INPA); Manaus, Dec 1850-Mar 1851 (9), Spruce cences 3-4 mm diam. Perianth of the staminate 

951 (M), Spruce 1219 (P), Spruce s.n. (B, CGE, E, F, flower densely puberulous. Peduncle of the pis- 

G, GOET, LE, NY, OXF, P). 

tillate inflorescences mostly up to 8 cm long. Pistillate 

flowers usually in two ? distinct clusters; 

perianth yellow-velutinous, without dense, white, 

arachnoid hairs. 

Distribution (Fig. 81).Venezuela (Bolivar), Peru 

(Loreto), and Brazil (Amazonas, Mato Grosso, 

and Roraima); in non-inundated forest. 

Specimens examined. VENEZUELA, AMAZONAS: 

Depto. Rio Negro, nr. Cerro de La Neblina Base Camp, 

22 Feb 1984 (9), Liesner 16175 (BG), 4 May 1984 (2), 

Thomas 3349 (BG). BOLiVAR: Rio Hacha, Rio Icabaru, 

2 Jan 1956 (8), Bernardi 2731 (F, NY), Rio Carfin, Apr 

1945 (a), Cardona 1222 (US, VEN). 

PERU. LORETO: Prov. Maynas, Rio Nanay, Mishana, 

6 Jan 1983 (st), Gentry et al. 39264 (BG). 

BRAZIL. AMAZONAS: Sao Paulo de Olivenca, 7 Oct 

1931 (8), Ducke s.n. (RB); Humaita, 12 Oct-6 Nov 

1934 (9), Krukoff 6887 (A, BR, F, G, LE, MO, NY, 

RB, S, U, US); Sao Paulo de Olivenca, 11 Sep-26 Oct 

1936 (8), Krukoff8223 (A, F, G, GH, LE, MO, NY, 

P, S, U, US, type collection of Coussapoa krukovii); 

Democracia, 31 Aug 1923 (2), Kuhlmann 255 (BG);


Varadouro do Morcego, 31 Aug 1923 (8), Kuhlmann PERU. LORETO: Prov. Alto Amazonas, Rio Pastaza, 

291 (RB); Manaus-Itacoatiara rd., km 57, 15 Sep 1976 Andoar, 21 Nov 1980 (2), Vdsquez et al. 809 (BG). 

(a), Mota 646 (INPA); km 159, 20 Sep 1974 (8), Prance PAsco: Prov. Oxapampa, Distr. Iscozacin, 22 Sep 1986 

et al. 22708 (INPA, MG, U), 291 (RB); Tef6, 28 Nov (8), Pariona et al. 954 (BG). 

1959 (st), Rodrigues et al. 1434 (INPA); Rio Solim6es, BRAZIL. AMAZONAS: Sao Paulo de Olivenca, 11 

Fonte Boa, 26 Oct 1968 (6), M. Silva 2199 (G, MG); Sep-26 Oct 1936 (9), Krukoff 8325 (A, BR, F, G, LE, 

Ipanor6 ("Panur6"), Oct 1852-Jan 1853 (9), Spruce NY, PS, U). 

2865 (B, BR, C, CGE, E, F, G, GH, GOET, LE, MG, BOLIVIA. COCHABAMBA: Torora, Jungas of Jana 

NY, OXF, P, US, type collection ofP. apiculata). MATO Maya, 1800 m, 24 Jun 1947 (2), Cardenas 3973 (US) 

GRosso: Rio Aripuana, Nucleo Pioneiro Humboldt, LA PAZ: Larecaja, Mapiri, 12-30 Sep 1939 (8), Krukoff 

Rio Juruena rd., 25 Oct 1973 (8), Berg et al. P.19878 10861 (A, F, G, MO, NY, S, U), 8 Oct-15 Nov 1939 

(F, INPA, MO, NY, P, S, U, US); Aripuana, 18 Feb- (9), Krukoff 11062 (A, F, G, MO, NY, S, U, UC, US, 

9 May 1977 (st), Gomes et al. 826, 1517, 1540, 1588, type collection of P. tomentosa subsp. persecta). 

1598, 1604, 1610, 1679 (INPA), 8 Nov 1978 (2), Rylands 

30 (INPA); Mun. Vila Bela da Santissima Trin- Vernacular names. Brazil. Amazonas: mapati 

idad, 5 km S of border of Rond6nia, 2 Nov 1985 (2), or 

Thomas et al. 4765 (BG). RONDONIA: Rd. to Abuna to 

mapaty; Mato Grosso: imbafibarana. Bolivia. 

Guajara-Mirim, 1 km N of Riberao, 25 Jul 1968 Cochabamba: uva de monte. 

(a), 

Prance et al. 6292 (G, INPA, NY, S, U, US). RORAIMA: Cardenas 3973 (from Bolivia) differs from the 

Anaris, 12 Feb 1969 (2), Prance et al. 9857 (F, GH, others in having the pistillate flowers in two clus- 

INPA, M, NY, P, K, S, U, US), Serra dos Surucucus, ters in the inflorescences. 

15 Feb 1969 (2), Prance 9952 (C, INPA, MO, MY, R, 

S, U). 

Vdsquez et al. 809 (from Peru) differs from the 

other collections in the absence of long yellow 

Vernacular names. Venezuela. Bolivar: sara- hairs on the leafy twigs. The same applies to 

sara (Arekuna). Brazil. Amazonas: imbauibarana, collection Korning et al. 47618 which has, morepurumai. 

over, distinctly petiolulate leaf segments. 

13c. Pourouma tomentosa Miquel subsp. per- 

secta C. C. Berg & van Heusden, Proc. Kon. 

Ned. Akad. Wetensch., Ser. C, 91(2): 108.1988. 

Type. Bolivia. La Paz: Prov. Larecaja, Copa- 

cabana, 10 km S ofMapiri, 8 Oct-15 Nov 1939 

(2), Krukoff 11062 (holotype, U; isotypes, A, 

F, G, MO, NY, S, UC, US). Fig. 80. 

13d. Pourouma tomentosa Miquel subsp. esse- 

quiboensis (Standley) C. C. Berg & van Heus- 

den, Proc. Kon. Ned. Akad. Wetensch., Ser. 

C 91(2): 109. 1988. 

Pourouma essequiboensis Standley, Lloydia 2: 175. 

1939; Berg, Fl. Suriname 5(1): 276. 1975. Type. 

Guyana. Essequibo R., nr. mouth of Onoro Creek, 

Leafy twigs and petioles yellow-hirsute, some- 15-24 Dec 1937 (d), A. C. Smith 2731 (holotype, F; 

times glabrous. Lamina 5-9-parted, with inci- isotypes, A, G, Mo, NY, P, S, U, US). 

sions down to (near) the petiole, sometimes with 

the segments petiolulate, occasionally entire, apex Leafy twigs and petioles without long yellow 

acuminate, base cordate to subcordate; upper hairs. Lamina 3-5-parted, apex acuminate, base 

surface smooth; stipules outside with sparse, ? deeply cordate; upper surface smooth; stipules 

white, arachnoid hairs, inside with sparse yellow with rather dense, white, arachnoid hairs, inside 

hairs. Heads of the staminate inflorescences ca. 

glabrous. Heads of the staminate inflorescences 

5 mm in diam. Peduncle of the pistillate inflo- ca. 3 mm diam. Peduncle of the pistillate inflorescences 

often longer than 8 cm. Pistillateflow- rescences mostly up to 6(-7) cm long. Pistillate 

ers + diffusely distributed in the inflorescence or 

flowers ? diffusely distributed in the infloressometimes 

clustered in two groups; perianth of cence. Perianth of the pistillate flower yellowthe 

pistillate flower yellow-velutinous, without 

velutinous, without dense, white, arachnoid hairs. 

dense, white, arachnoid hairs. 

Distribution (Fig. 81). Guyana and Brazil 

Distribution (Fig. 81). Brazil (Amazonas and 

(Amazonas and Para); in non-inundated forest. 

Mato Grosso), Peru (Loreto), and Bolivia (La Paz 

and Cochabamba); in non-inundated forest, at Specimens examined. GUYANA. Mouth of Onoro 

altitudes up to 1800 m. 

Creek, 15-24 Dec 1937 (6), A. C. Smith 2731 (A, F, 

G, GH, MO, NY, P, S, U, US, type collection of P. 

Specimens examined. ECUADOR. NAPO: Aiiangu, essequiboensis). 

30 May-21 Jun 1982 (st), SEF 8823 (U), 1-15 Feb BRAZIL. AMAZONAS: Itapiranga, Rio Uatuma, 27 

1986 (2), Korning et al. 47618 (BG). 

Aug 1979 (C), Cid et al. 847 (BG, INPA); Rio Jurua,


_ I 

Revised 

SAt L arsed forkFloa e d.rrovc e. e 

FIG. 80. Pourouma tomentosa subsp. persecta. Leafy twig with pistilate inflorescences (Krukoff 8325). 

Saracura, 22 Aug 1975 (9), D. Coelho et al. (INPA) 

53383 (INPA, U); Codajas, Rio Capitari, 3 Sep 1950 

(d), Fr6es 26520 (US). PARA: Cuiaba-Santarem rd., km 

1180, 17 Nov 1977 (2), A. S. Silva et al. 215 (BG, COL, 

K, MO, U). 

w 

L U - a SL I-. ITtA 

ora fm bro picn a T. 

13e. Pourouma tomentosa Miquel subsp. ma- 

roniensis (Benoist) C. C. Berg & van Heusden, 

Proc. Kon. Ned. Akad. Wetensch., Ser. C 91(2): 

109. 1988. Fig. 82a. 

FIG. 81. Distribution of Pourouma herrerensis, P. hirsutipetiolata subsp. hirsutipetiolata, P. hirsutipetiolata 

subsp. hispida, P. napoensis, P. tomentosa subsp. apiculata, P. tomentosa subsp. essequiboensis, P. tomentosa 

subsp. maroniensis, P. tomentosa subsp. persecta, and P. tomentosa subsp. tomentosa.


qlq, 

_ j PIF P. t osantosa stubsp. t lontosa 

__ / *~ _~ * 

C~ Osa iF~ P with entire lamina 

ab^P~ 0?^P "L^ I-~~S 

^fr/I Z 1^ ^ 

P..hir?utiptiolat. 

1A 

^-^ 

^ 0with incised 

Smrine 

,wrr@e ,3 

s 

^Sfti^"1~^PA ?- 'AP. naponsis 

i-r g^ -r ----- ---p.-' b-------AS"s"eioovsZ 

'l^^ ? C 

^P0 

. sqa ounu 

sulbOp. mflroni.nsis 

.h a,~_ 

,ns is 

171


FIG. 82. a. Pourouma tomentosa subsp. maroniensis. Leaf and pistillate inflorescences (Lanjouw et al. 432). 

b. Pourouma minor. Leafy twig with pistillate inflorescences (Prance et al. 10089). 

Pourouma maroniensis Benoist. Bull. Mus. Hist. Nat. 

(Paris) 28: 319. 1922; Berg, Fl. Suriname 5(1): 276, 

t. 14a. 1975. Type. French Guiana. Godebert, 23 

Jun 1921 (d), Wachenheim ser. 2: 392 (holotype, P). 

Leafy twigs and stipules without long yellow 

hairs. Lamina entire and ovate, only in subju- 

venile material to 3-parted, but then the base to 

truncate or at most subcordate, apex acuminate; 

upper surface smooth; stipules outside with dense, 

white, arachnoid hairs, inside glabrous. Heads of 

the staminate inflorescences normally 2-3 mm 

diam. Perianth of the staminate flower sparsely 

puberulous. Peduncle of the pistillate inflores- 

cences mostly up to 7 cm long. Pistillate flowers 

+ diffusely distributed in the inflorescence. Peri- 

anth of the pistillate flower sparsely white-pub- 

erulous or yellow-velutinous, often with dense, 

white, arachnoid hairs. 

Distribution (Fig. 81). Eastern Surinam, French 

Guiana, and Brazil (Amapa and Para); in non- 


Specimens examined. SURINAM. Moengo, 25 Aug 

1951 (st), BBS 298 (U); Gansee, 21 Apr 1964 (st), 

Donselaar 1226 (U); Grote Zwiebelzwamp, 23 Sep 1948 

(9), Lanjouw et al. 432 (F, NY, U); Lely Mts., 2 Oct 

1975 (st), Lindeman et al. 701 (U). 

FRENCH GUIANA. St. Laurent, Feb 1956 (9), 

BAFOG 7228 (NY, P, U); Mana, 3 Mar 1956 (9), BAF- 

OG 7330 (NY, P); Charvein, 29 Jun 1914 (d), Benoist


465 (P); Trois Sauts, Zidockville, 28 Jan 1975 (2), Gren- type collection of P. stipulacea); 1-6 km W of Kaand 

688 (CAY); Mana R., Saut Sabbat, 16 Jul 1981 marang, Mazaruni R., 21 Aug 1977 (st), Maas et al. 

(2), Granville 4531 (BG); Godebert, 23 Jun 1921 (6), 2592 (U). 

Wachenheim 392 (P, type collection of P. maroniensis). 

This species shares (sub)persistent stipules with 

BRAZIL. AMAPA: Serra do Navio, 4 Jan 1985 (6), P. oraria and P. bicolor subsp. chocoana. Most 

Mori et al. 17672 (BG), 25 Sep 1961 (6), Pires et al. 

51193 (B, FHO, INPA, NY, P, U, US); Rio 

features, e.g., the white, arachnoid indument on 

Araguari, 

9 Oct 1961 (6), Pires et al. 51625 (FHO, NY, U, US); 

most parts, suggest relationship to P. tomentosa. 

Serra do Navio, 1961 (st), Rodrigues 2986 and 2987 According to Boom (et al. 8102) the exudate 

(INPA). PARA: Rio Jari, Monte Dourado, 22 Jan 1968 is red and viscous. 

(2), Oliveira 3936 (NY). 

Vernacular names. Surinam. Boroma (Ara- 

wak), boesipapja, granboesipapaja. French 

Guiana. Bois canon, bouchi papaye (Paramaka), 

kukuma, wilaupiyua (Wayapi). 

The morphological differences between subsp. 

maroniensis, subsp. apiculata and subsp. tomen- 

tosa are not strong. 

14. Pourouma stipulacea C. C. Berg, Brittonia 

34(1): 37, t. 2. 1982. Type. Guyana. Bartica- 

Potaro rd., 25 Jan 1943 (2), Fanshawe 1105 = 

FD 3851 (holotype, K). Fig. 83. 

Tree, up to 6 m tall. Leafy twigs 1-2.5 cm thick, 

yellow-hirsute. Lamina 5-parted, 25-50 x 35- 

75 cm, coriaceous, apex acuminate, base deeply 

cordate, often with lobes overlapping; upper surface 

smooth, hirtellous on the main veins, lower 

surface white-puberulous and with sparse, orange, 

pluricellular hairs on the veins, arachnoid 

hairs ? confined to the areoles; lateral veins 20- 

30 pairs, tertiary (and quartemary) veins prominent 

beneath; petiole 23-65 cm long, sparsely 

yellow-hirsute; stipules 15-20 cm long, outside 

yellow-hirsute and with sparse to dense, white, 

arachnoid hairs, and with sparse, minute, orange 

pluricellular hairs, inside sparsely yellow-hirsute, 

subpersistent. Staminate inflorescences unknown. 

Pistillate inflorescences in fruit up to 15 

cm long and 8 cm wide; peduncles 10 cm long, 

peduncle and branches sparsely yellow-hirsute 

15. Pourouma acuminata Martius ex Miquel in 

Martius, Fl. bras 4(1): 130. t. 40. 1853; Mar- 

tius, Syst. mat. med. bras. 34. 1843, nomen. 

Type. Brazil. Amazonas: Rio Japura, between 

Maripi and Parana-Mirim, Dec 1819 (9), Mar- 

tius s.n. (holotype, M; isotype, U). Fig. 84. 

Pourouma populifolia Standley, Publ. Field Mus. Nat. 

Hist., Bot. Ser., 17: 184. 1937. Type. Brazil. Amazonas: 

Mun. Sao Paulo de Olivenca, nr. Palmares, 

1 Sep-26 Oct 1936 (9), Krukoff8427 (holotype, NY; 

isotypes, A, BR, F, G, LE, P, S, U, US). 


thick, (sparsely) yellow-hirsute. Lamina entire, 

(broadly) ovate to deltoid or to elliptic, 5-19 x 

3-10 cm, coriaceous, apex (long) acuminate, base 

rounded to acute; upper surface smooth, main 

veins sparsely hairy, lower surface white-, ap- 

pressed -puberulous on the main veins, minutely 

puberulous on the smaller veins, arachnoid hairs 

in the areoles, sometimes extending to the main 

veins; lateral veins 7-14 pairs, basal pair un- 

branched, tertiary venation slightly prominent 

beneath; petiole sparsely appressed-puberulous 

to glabrescent, and usually sparsely hirsute; stip- 

ules 5-10 cm long, outside yellow-hirsute and 

with dense, white, arachnoid hairs, occasionally 

with minute, appressed, white hairs, inside gla- 

brous, caducous. Staminate inflorescences unknown. 

Pistillate inflorescences in fruit up to 10 

cm long and 4 cm wide; peduncle 4.5-8 cm long, 

peduncle and branches puberulous (velutinous 

and with (dense), white, arachnoid hairs; flowers on the ultimate branches);flowers 4-17; pedicels 

ca. 15; pedicel in fruit 0.5-1 cm long; stigma 0.3-0.6 cm long, in fruit up to 1.5 cm long; peripeltate, 

2 mm diam. Fruitingperianth ovoid, ca. anth 2-5 mm 

2 x 

long, densely yellow-velutinous, 

1 cm, yellow-hirsute and with dense, white- apex papillose; stigma peltate, 1.5-2 mm diam., 

arachnoid hairs. 

sometimes puberulous. Fruiting perianth ovoid 

Distribution (Fig. 85). Guyana; in non-inun- to elliptic, ca. 1-1.6 x 1 cm, densely yellowdated 


velutinous. 

Specimens examined. GUYANA. Nr. Kamerang, 20 Distribution (Fig. 85). Brazil (Amazonas) and 

Jun 1987 (st), Boom et al. 8102 (BG); Bartica-Potaro Peru (Loreto); in periodically inundated (varzea) 

rd., 25 Jan 1943 (2), Fanshawe 1105 (=FD 3851) (K, forest or also in non-inundated forest.


- - 

g 

FIG. 83. Pourouma stipulacea. Leaf and pistillate inflorescence (Fanshawe 1105); twig with stipules (Maas 

et al 2592). 

FIG. 84. Pourouma acuminata. From Martius, Flora Brasiliensis 4(1). 1853: tab. 40. Leafy twig with pistillate 

inflorescences, fruiting perianth and fruit (17), fruit (19), fruiting perianth (4311), pericarp and seed (19, 21), 

seed (21), embryo (23), radicle (25).


I0711~~~~~~~~~~~~~~~~~ 41 al 

9- . . .. ,~ act ninat 

..4 

I'OU'R M A L aeminnata.

+~~~5 P. acuminata .--ID P. 

fei\`f `~ p ~rugines 

4~074' -~-3~ 

-P.


Specimens examined. VENEZUELA. BOLvAR: Mun. white-puberulous, densely so on the ultimate 

Foraneo Aripao, Carno Fatua, 2-5 May 1988 (st), Ay- branches; flowers sessile, not densely clustered; 

mard et al. 6778 (BG). 

PERU. LORETO: Rio Itaya, 5 km above Iquitos, 8 tepals ca. 1.5 mm long, almost free, densely pu- 

Aug 1972 (9), Croat 18888 (C, DUKE, F, GH, NA, berulous; filaments shorter than the tepals. Sta- 

NY, P); Rio Itaya, Nuevo Esperanza, 10 Oct 1975 (9), minate inflorescences unknown. Pistillate inflo- 

McDaniel et al. 20362 (NA); Rio Momon, 12 Oct 1977 rescences up to 70 cm long and 16 cm wide; 

(2), Rimachi 3256 (MO). 

peduncle 32-48 cm 

BRAZIL. AMAZONAS: Rio Japura, Vila 

long, peduncle and branches 

Bittancourt, 

11 Nov 1982 (9),Amaral etal. 473(BG); Rio ICa, 1877with 

indument similar to that of the staminate 

1878 (2), Jobert 682 (P); Mun. Sao Paulo de Olivena, inflorescences;flowers 7-13; pedicel 0.7-1.3 cm 

nr. Palmares, 11 Sep-26 Oct 1936 (9), Krukoff 8427 long, in fruit up to 5 cm; perianth ca. 1 cm long, 

(A, BR, F, G, LE, MO, NY, P, S, U, US, type collection densely white-puberulous; 

of P. 

stigma peltate, ca. 3 

populifolia); Rio Japura, Parana-Mirim, Dec 1819 mm 

(2), Martius s.n. (M, U, type collection of P. acumi- diam.; bilobate, densely puberulous. Fruitnata); 

Rio Ia, Oct 1877 (9), (herb.) Schwacke 560 (R, ing perianth globose, ca. 2 cm diam., apiculate, 

RB). 

yellow-short-velutinous. 

Distribution 

Vernacular names. Peru. Loreto: sacha 

(Fig. 85). Brazil (Amazonas), in 

uvilla, non-inundated forest. 

uvilla. Brazil. Amazonas: mapati or mapaty. 

This species is rather uniform. It resembles the Specimens examined. BRAZIL. AMAZONAS: Rio Soform 

ofP. herrerensis with oblong to elliptic leaves lim6es, mouth of Rio Jutai, 5 Sep 1975 (2), L. Coelho 

339 

and unbranched basal lateral veins, but can be 

(INPA, U); Sao Paulo de Olivenca, 10 Oct 1931 

($), Ducke (RB) 25244 (RB); Sao Paulo de Olivenca, 

distinguished from the latter by the indument of Creek Belem, 26 Oct-11 Dec 1936 (6), Krukoff8807 

the leafy twigs, consisting of only long hairs or (A, F, NY, type collection ofP.ferruginea); Tonantins, 

wanting. 

19 Feb 1944 (2), Ducke 1527 (A, F, IAN, MG, NY, R, 

Glaziou 10070 (according to the label from 

UC, US); Rio Solim6es, Sao Antonio de Ila, 2 Dec 

1948 (2), Fr6es 23693 (IAN); Manaus-Porto Velho 

Brazil, Espirito Santo, Itapemerim) is probably hwy., BR-319, km 380, 2 km S of Rio Jutai, 13 Oct 

the same collection as Schwacke 560. 

1974 (6), Prance et al. 22859 (INPA, MG, U). 

16. Pourouma ferruginea Standley, Publ. Field 

Mus. Nat. Hist., Bot. Sr., 17:181. 1937. Type. 

Brazil. Amazonas: Mun. Sao Paulo de Oliven- 

ca, Belem Creek, 26 Oct- 1 Dec 1936 (6), Kru- 

koff 8807 (holotype, NY; isotypes, A, F). 

Fig. 86. 

Tree, up to 20 m tall. Leafy twigs ca. 10-15 

mm thick, white-puberulous and with dense, 

purplish, pluricellular hairs; lenticels conspicuous. 

Lamina entire, ovate and 11-32 x 8-23 cm 

or 3-5-fid and up to ca. 45 x 45 cm, coriaceous, 

? plicate, apex rounded to acuminate, base truncate 

to cordate; upper surface smooth, hairy on 

the main veins, lower surface sparsely subsericeous 

to strigose on the main veins, the whole 

surface covered with white or brownish, arachnoid 

hairs, which disappear from the main veins; 

lateral veins 8-16 pairs, the basal pair branched, 

occasionally unbranched, tertiary venation plane 

to slightly prominent beneath; petiole 4-30 cm 

long, with dense whitish, arachnoid hairs; stipules 

3-13 cm long, outside hispid to substrigose, 

inside glabrous, caducous. Staminate inflorescences 

up to 30 cm long and up to 20 cm wide; 


Vernacular names. Brazil. Amazonas: mapatirana. 

This species matches P. ovata in the longly 

pedunculate, pendulous pistillate inflorescences 

with globose and apiculate fruiting perianths. The 

two species are not closely related. 

See unnamed collection number 3 (p. 193). 

17. Pourouma villosa Trecul, Ann. Sci. Nat. Bot., 

Ser. 3, 8: 103. 1847; Miquel in Martius, Fl. 

bras. 4(1): 127. 1853. Type. French Guiana. 

Without locality, 1840 (6), Leprieur s.n. (holotype, 

P). Fig. 87. 

Pourouma laevis Benoist, Bull. Mus. Hist. Nat. (Paris) 

28: 319. 1922; Berg, Fl. Suriname 5(1): 270. 1975. 

Type. French Guiana. Acarouany, 1858 (6), Sagot 

517 (lectotype, P, chosen here; isolectotypes, B, BR, 

F, G, GH, GOET, P, S, U, US). 


thick, yellow(ish)-puberulous to -subvelutinous 

to -subhirsute, or (at least on the scars of the 

stipules) pale yellow- to whitish-villous, with 

dense, brown, pluricellular hairs. Lamina 3-5lobed 

to -fid, sometimes entire and ovate, rarely 

obovate to oblong, 5-40 x 4-42 cm, coriaceous, 

rarely chartaceous, apex acuminate, base deeply


?~~~~~~~~~~~~~? 

;: - ~ 

? 

. 

' 

.. '". 

~ 

~~~L~~r~~AI ~~~~R~~~?? 

Ir ?i 

~ ..194 ~ . ~ ~ 

. . ' !':,"..: 

r~~~~~~~~':. 

. :.:.::.. . . ........... .... 

Neg|tive nch ~ ~ .. 

TO 2 

rr 

F; ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ,' ~- ,~ ...............~,':..:,,. 

? ' ':.-L,. . . ~ . ,> ~ ~ 

la"?fr ~~~~~~~ 

,~'~ .... ,i 4: ,.~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~ 

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

-..-.:. ~ 4~ ~~~~ 

-?. ._~ ... 

Jo - ......":.... . 

.....: . 

FIG. 86. Pourouma ferruginea. Leafy twig with pistillate inflorescence (Ducke 1527). 

.... 

. . ' . ':'.-,.C.~ . . . . . _ 

IG86Poruafruine. Laytwi ihpsilt nirsecDce12) 

cordate to subcordate, sometimes with overlap- 

ping lobes; upper surface smooth, hairy on the 

main veins, lower surface white-puberulous to 

yellow-hirsute on the veins, usually with brown, 

pluricellular hairs on the main veins, and white, 

arachnoid hairs in the areoles and on the smaller 

veins; lateral veins 6-20 pairs, the basal pair 

branched (and their basal parts separated from 

'


11 

'.ciLnnr'.mon-lik 

2 3 4 5 

THNE NEW YORK BOTANICAL GARDEN 

No". '.3 : j ", -'. 

Old G.D: car.p I!. ? ".. 

Faysetr AIrstri'.) 

kNut.v Wnch 1of 2fpmribo. 

stilt roots. Fril r'd 

odor:?'" 

BOTANICAf!' ? O iO 

FIG. 87. Pourouma villosa. Leafy twig with pistillate inflorescences (Mori et al. 8583). 

the margin by mesophyll), tertiary venation 

slightly prominent beneath; petiole 6-35 cm long, 

yellow-hirsute and with long and short, appres- 

sesd, white hairs, and with (dense), brown, arach- 

' 

179 

noid hairs; stipules 3-25 cm long, outside yellowhirsute 

to -velutinous or pale yellow- to whitishvillous, 

also with dense, brown, pluricellular hairs, 

inside yellow-hirsute to glabrous; caducous.


Staminate inflorescences up to 21 cm long and and Serro do Navio, 10 Oct-15 Dec 1976 (st), Rosa 

12 cm wide; peduncle 2-10 cm 1126 

long, yellow-pu- 

(MG). AMAZONAS: Manaus, 23 Oct 1969 (8), L. 

Coelho 15 (INPA, U), 17 Jan 1931 (2), Ducke 

berulous and with dense 

(RB) 

brown, pluricellular 25243 (RB), 2 Apr 1976 (st), Macedo s.n. (INPA), 24 

hairs; flowers sessile, mostly clustered at the ul- Oct 1978 (8), Rodrigues et al. 10082 (INPA), 1850timate 

branches; tepals 1-2 mm long, ovate to 1851 (8), Spruce s.n. (G), Mar 1912 (2), Ule 8839 (G, 

lanceolate, (almost) free or up to halfway con- MG). MARANHAO: Rod. Belem-Brasilia, km 380-375, 

28 

nate, sparsely hairy; filaments shorter than the Aug 1960 (2), Oliveira 1074 (IAN). PARA: Faro, 3 

Jan 1920 (9), Ducke (MG) 10736 = (RB) 13056 (MG, 

tepals. Pistillate inflorescences in fruit up to 19 RB); Rio Branco, Obidos, Serra da Boa Vista, 23 Dec 

cm long and 10 cm wide; peduncle 2-12 cm long, 1913 (9), Ducke (MG) 25225 (MG); upper Rio Cupary, 

peduncle and branches yellow-puberulous to between Rio Xingu and Rio Tapaj6s, Sep 1931 (2), 

-hirtellous and with dense, brown to red-brown Krukoff 1197 (A, G, NY, P, S, U); Rio Tapaj6s, Boa 

or dark 

Vista, 1931 (2), Monteiro da Costa 92 

brown, pluricellular hairs;flowers ca. 10- 

(F); Rio Jari, 

Monte Dourado, 14 Jan 1968 (a), Oliveira 3915 (NY). 

40; pedicels up to 0.6 cm long, in fruit up to 1.2 RONDONIA: P6rto Velho-Cuiaba hwy., Santa Barbara, 

cm; perianth 4-8 mm long, yellow-puberulous 16 Aug 1968 (8), Prance et al. 7160 (F, GH, INPA, 

and with dense, dark brown, pluricellular hairs; NY, P, S, U, US). 

stigma peltate, 1.5-2.5 mm diam. Fruiting peri- Vernacular names. Surinam. Boroma (Aroanth 

purple, (sub)ovoid to ellipsoid, ca. 1.5-2 x 

wak), granboesipapaja, kobe (Djoeka), manbos- 

0.7-1.5 cm, yellow-puberulous and with sparse papaja, pourouma or puruma (Carib), yarayara 

to dense, dark brown, pluricellular hairs. 

(Carib). French Guiana. Bois canon, 

Distribution 

(bouchi) pa- 

(Fig. 85). Surinam, French pate or papaye (Paramaka). Brazil. Amapa: im- 

Guiana, and Brazil (Amapa, Amazonas, Para, bafiba bengue; Para: imbafiba branca, imbafiba 

and Rond6nia); mostly in non-inundated forest, de cheiro uvilha. 

sometimes in riverine forest, at low alti- The relationships with other Pourouma species 

tudes. 

are not clear. The collections from Amazonas 

Specimens examined. SURINAM. Moengo, 16 and 

Aug Rond6nia, three collections from Para (the 

1951 (st), BBS 297 (U); Brownsberg, 15 Jul 1916 (st), two Ducke collections and Oliveira 3915) and 

BW2095 (U), 13 Sep 1917 (6), BW 3255 (NY, U), 7 the type collection of P. villosa, have long pale 

Sep 1917 (6), BW3302 (B, F, MO, P, U), 20 Sep 1918 

(), BW 4010 (U, UC), 17 Sep 1921 (9), BW 

yellow hairs on the 

5390 

petioles and leafy twigs, but 

(F, 

RB, U), 28 Jun 1924 (6), BW 6509 (U); Brownsweg, 

they are similar to the other collections in all 

16 Nov 1964 (st), Donselaar 1888 (U); Wilhelmina other features. Together with the overlap in dis- 

Mts., 10 Sep 1963 (8), Irwin et al. 55576 (B, FHO, GH, tribution this is a reason not to recognize the 

M, MO, NY, U, US), 20 Sep 1963 (6), Irwin et al. form with long hairs as a distinct 

55884 infraspecific 

(FHO, GH, NY, S, U, US); Fallawatra, 6 Nov 

1971 (2), Jimenez-Saa 1563 = taxon. 

LBB 14296 (NY, U); 

Lely Mts., 29 Sep 1975 (st), Lindeman et al. 540 Cut fruit 

(U), 

give out a cinnamon-like odor (Mori 

5 Oct 1975, Lindeman et al. 790 (U); Paris Jacob Creek, et al. 8583). 

28 Jun 1965 (st), Maas et al. (LBB) 11017 (U); Kayser The description of P. laevis was based on Ben- 

Mts., 29 Oct 1976 (9), Mori et al. 8535 (F, NY, U, US); oist 

Sectie O, Nov 1942 

341, 

(9), Stahel (Woodherb. Sur.) 123B 

Sagot 517, and Sagot 972. Sagot 517 

(U), Dec 1942 (9), Stahel (Woodherb. Sur.) 166A 

is chosen as the 

(U). 

lectotype collection. 

FRENCH GUIANA. Without locality (9), Aublet s. n. 

(G), 1840 (6), Leprieur s.n. (P, type collection of P. 18. Pourouma oraria Standley & Cuatrecasas in 

villosa); Charvein-Acarouany rd., km 1, 27 Oct 1953 Cuatrecasas, Caldasia 7: 301. 

(9), BAFOG 93-M(CAY, P, U); St. 

1956; 

Laurent, Dec 

Woodson, 

1955 

(9), BAFOG 7087 

Ann. Missouri 

(CAY, NY, P, U); Route 

Bot. 

de 

Gard. 47:167.1960. 

Cayenne, 

Type. 

km 14, 28 Oct 1957 (8), BAFOG 7745 (CAY, NY, P); 

Colombia. Valle: Rio Yurumagui, Veneral, 28 

Charvein, (9), Benoist 341 (P); Acarouany, 1858 (6), Jan 1944 (2), Cuatrecasas 15720 (holotype, F). 

Sagot 517 (F, G, GH, GOET, P, S, U, U S, lectotype 

collection of P. laevis; in F and P, material of P. mel- Tree, up to 20 m tall. Leafy twigs up to 15 mm 

inonii subsp. melinonii under the same number), 1856 thick, yellow-puberulous and with dense, brown, 

(9), Sagot 970 (P), (9), Sagot 972 (B, BR, F, G, P, S, pluricellular hairs. Lamina 7-fid, 28-32 x 40 

U), (9), Sagot s.n. (U), (st), Sagot s.n. (G). 

BRAZIL. AMAPA: 35 km ESE of Oiapoque, 2 Dec 

cm, coriaceous, apex acuminate, base deeply cor- 

1984 (9), Daly et al. 3790 (BG); Rio Amapari, Rancho date with overlapping lobes; upper surface 

Santa Ana, 30 Sep 1961 (6), Pires et al. 51390 (FHO, smooth, main veins hairy, lower surface puber- 

IAN, INPA, MG, MO, NY, US); between P6rto Platon ulous on the veins, arachnoid hairs confined to


the areoles; lateral veins up to ca. 30 pairs, ter- pressed-puberulous on the main veins, arachtiary 

and quarternary venation prominent be- noid hairs in the areoles and on the smaller veins; 

neath; petiole 30 cm long, yellow-hirsute at the lateral veins 7-12 pairs, basal pair unbranched, 

adaxial side, for the rest sparsely hairy; stipules tertiary venation plane beneath; petiole 2-9 cm 

10-12 cm long, outside yellow-hirsute, and with long, with some (very) sparse, white hairs; stipwhite 

appressed hairs, occasionally with brown, ules 1-5 cm long, outside yellow-appressed-pupluricellular 

hairs, inside yellow-hirsute, subper- berulous, inside glabrous, caducous. Staminate 

sistent. Staminate inflorescences about 25 cm long inflorescences unknown. Pistillate inflorescences 

and 15 cm wide; peduncle ca. 12 cm long, yellow- in fruit up to 8 cm long and 5 cm wide; peduncle 

puberulous; flowers in clusters of up to 10 flow- 2-3.5 cm long, peduncle and branches minutely 

ers; tepals free, ca. 2 mm long; filaments shorter yellow-puberulous, the ultimate branches more 

than the tepals. Pistillate inflorescences in fruit densely hairy, and with brown, pluricellular hairs; 

up to 22 cm long and 18 cm wide; peduncle 14- flowers 7-10; pedicel 0.1-0.5 cm long, in fruit up 

15 cm long, peduncle and branches yellow-pu- to 1 cm; perianth ca. 1.5-2 mm long, yellowberulous 

and with dense, dark-brown or red- puberulous; stigma bilobed, ca. 2.5 mm diam. 

brown or brown, pluricellular hairs; flowers ca. Fruitingperianth ovoid, ca. 2 x 0.8 cm, sparsely, 

100; pedicels ca. 5 mm long, in fruit up to 10 minutely appressed-puberulous. 

mm long; perianth 3-5 mm long, yellow-puber- Distribution (Fig. 85). French Guiana and Braulous 

and with dense, dark-brown, pluricel- zil (Amapa); in non-inundated forest. 

lular hairs; stigma peltate, 1.5-2 mm diam. 

Fruiting perianth ovoid to ellipsoid, ca. 15 x 10 

Specimens examined. FRENCH GUIANA. Saiil, 20 

Dec 1976 (Q), Granville 2744 (CAY), Saul, km 2.7 on 

mm, + sparsely hairy. 

trail on Montagne Belvedere Sud, 21 Oct 1971 (v), 

Distribution (Fig. 85). Colombia (Choc6 and Oldeman B.4132 (CAY, P, U, VEN type collection of 

Valle); in lowland forest. 

P. saulensis). 

BRAZIL. AMAPA: 109 km SSE of Oiapoque, 5 Dec 

Specimens examined. COLOMBIA. CHOC6: Be- 1984 (v), Mori et al. 17185 (BG). 

tween Certegui and Las Animas, 17 Aug 1976 (9), Gentry 

et al. 17804 (BG, U); N of Certegui, 13 Jun 1982 Vernacular name. French Guiana. Bois canon. 

(8), Gentry et al. 36797 (JUAM). VALLE: Costa del This species shows similarities to P. ovata in 

Pacifico, Rio Yurumangui, Veneral, 28 Jan 1944 (9), the indument, the short stipules, and the un- 

Cuatrecasas 15720 (F, type collection of P. oraria). branched, basal, lateral veins. The species are 

distinct in the number of lateral veins, the length 

of the peduncle of the pistillate inflorescence, and 

the shape of the fruiting perianth. 

This species resembles P. bicolor subsp. chocoana 

in many features (see p. 144). It differs 

from the latter in the smooth upper surface of 

the lamina, the smooth fruiting perianth, and the 

concentration of the long hairs at the adaxial side 

of the petiole. P. oraria may be much closer to 

P. bicolor (subsp. chocoana) than suggested by 

the arrangement oftaxa in the present treatment. 

19. Pourouma saulensis C. C. Berg, Brittonia 

34(1): 36, t. 1. 1982. Type. French Guiana. 

Sail, km 2.7 on trail of Montagne Belvedere 

Sud, 21 Oct 1971 (9), Oldeman B.4132 (ho- 

lotype, U; isotypes, CAY, P, VEN). Fig. 88. 


thick, minutely puberulous to glabrous. Lamina 

entire, (broadly) ovate to elliptic, 6-14 x 3-9.5 

cm, coriaceous, apex (sub)acuminate, base 

rounded to obtuse; upper surface smooth, some 

sparse, white hairs and brown, pluricellular hairs 

on the main veins, lower surface white-, ap- 

20. Pourouma ovata Trecul, Ann. Sci. Nat. Bot., 

Ser. 3, 8: 101. 1847, Miquel in Martius, Fl. 

bras. 4(1): 132.1853. Type. Brazil. Para: Without 

locality, without date (d), Ferreira s.n. (holotype, 

P). Fig. 89. 

Pourouma longipendula Ducke, Notizbl. Bot. Gart. 

Berlin-Dahlem 11: 581. 1932. Type. Brazil. Amazonas: 

Manaus, falls of Rio Taruma; 17 Oct 1929 

(v), Ducke (RB) 23608 (holotype, RB, not seen; isotypes, 

B, G, S, US). 


thick, sparsely white-appressed-puberulous, more 

densely haired at the nodes. Lamina entire, elliptic 

to ovate, 7-26 x 3-13 cm, coriaceous, apex 

short acuminate, base acute to obtuse to rounded 

or sometimes truncate; upper surface smooth, 

sparsely hairy and occasionally with some brown,


[-- 

i. " 

Q~~~~~~~~~~~," 

.' 

? " 

FIG. 88. Pourouma saulensis. Leafy twig, (schematic) pistillate inflorescence, and fruiting perianth (Granville~~~~~~ 

2744); pstillat flower OldemanB.413') 

FIG. 88. Pourouma saulensis. Leafy twig, (schematic) pistillate inflorescence, and fruiting perianth (Granville 

2744); pistillate flower (Oldeman B.4132). 

pluricellular hairs on the midrib, lower surface 

white-puberulous on the veins, arachnoid hairs 

confined to the areoles; lateral veins 9-21 pairs, 

basal pair unbranched or occasionally branched, 

tertiary venation slightly prominent beneath; 

petiole 2-7.5 cm long, subglabrous to sparsely 

white hairy; stipules 1.5-3 cm long, outside yel- 

low-velutinous, usually with brown, pluricellular 

hairs, inside yellow-(or orange-brown- or 

white-)sericeous, caducous. Staminate inflores- 

cences up to 10 cm long and 9.5 cm wide; pe- 

duncle 1-5.5 cm long, sparsely puberulous to 

glabrescent, ultimate branches usually more 

densely yellow- to orange-puberulous; flowers 

sessile or occasionally pedicellate, mostly in 

subglobose heads, these 3-7 mm diam.; tepals 

ca. 1 mm long, connate, rather densely hairy; 

filaments shorter than the tepals. Pistillate inflorescences 

in fruit up to 38 cm long and 11 cm 

wide; peduncle up to 32 cm long, peduncle and 

branches puberulous to almost glabous; flowers 

7-20; pedicel 0.2-2.4 cm long, in fruit 1-3 cm, 

obconical, conspicuously lenticellate; perianth 3- 

5 mm long, densely short-velutinous; stigma pel- 

.. 

- 

'


Ak~~~~~~~~~~~~?IIk 

i. 

| I 3 -2^ 2 A? :. 

T rM w Tons BOTANICAL GAIM'W 

liBlTIm n lA L AiASS PA MZI. 

ii~L m.---.-V---lmDI, 

_ r ,.l. _ l,-~v kd L 

of tioe rm ch liho. 

oree 

? 

wdll 

Frest om terra firme on cly. 

Tm 1in x 20cm; frvit green, maturing red, 

clew light bow . ri. oraa 

w 

' g- '*--- J 

. 

'l - 

.* 

ovd'sember 1-8.... 

FIG. 89. Pourouma ovata. Leafy twig with pistillate inflorescences 

(Steward et al. 107).


tate, bilobed, ca. 1.5-2 mm diam. Fruiting peri- 

anth red (mature?), (depressed-)globose, ca. 1.5 

cm diam., puberulous. 

Distribution (Fig. 85). Colombia (Vaup6s), 

Venezuela (Bolivar), Peru (Loreto), and Brazil 

(Acre, Amazonas, and western Para); in non- 


Vernacular names. Peru. Loreto: chullachaqui, 

chullachaqui blanco, chullachaqui caspi, sacha 

uvillos. Brazil. Amazonas: imbafbarana. 

This species resembles P. ferruginea in the longpedunculate 

and pendulous pistillate inflorescences 

and the globose and apiculate fruiting 

perianth, but is in other features quite distinct 

from the latter. In the 

Specimens examined. COLOMBIA. VAUPES: Mon- 

vegetative parts it shows 

fort, 30 Nov 1952 (9), Romero-Castaneda 3848 resemblances to P. saulensis in several characters 

(F, 

MBG, NY); Rio Apaporis, Soratama, 26 Mar 1952 (2), (seep. 181). 

Schultes et al. 16009 (C, U, US). 

VENEZUELA. BOLIVAR: Sierra Ichfin, Salto Maria 

Espuma, 28 Dec 1961 (6), Steyermark 90368 (NY, U, 21. Pourouma elliptica Standley, Publ. Field 

VEN). 

Mus. Nat. Hist, Bot. Ser., 17:181. 1937. Type. 

PERU. LORETO: Prov. Requena, Rio Ucayali, Ar- 

Brazil. Amazonas: Mun. Sao Paulo de Olivenboretum 

Jenaro Herrera, Aug-Sep 1976 (st), Bernardi 

5.16.5 (U), Aug-Sep 1976 (9), Bernardi s.n. (tree 4/8) 9a, nr. Palmares, 11 Sep-26 Oct 1936 (6), Kru- 

(U), 3 Jan 1974 (9), Diaz 22-A (G); Prov. Maynas, Rio koff8388 (holotype, NY; isotypes, A, BR, F, 

Nanay, Mishana, 23 Mar 1979 (st), Gentry et al. 26049 G, LE, MO, P, S, U). Fig. 90. 

(U), 24 Mar 1979 (st), Gentry et al. 26177 (U), 12 Jan 

1983 (st), Gentry et al. 39379 (BG); Prov. Requena, Tree, up to ca. 15 m tall. Leafy twigs ca. 10 

Arboretum Jenaro Herrera, 9 Dec 1980 (9), Vdsquez mm thick, densely yellow-hirsute and yellowet 

al. 1004 (BG). 

BRAZIL. ACRE: Cruzeiro do Sul, 11 Feb 1976 subhispidulous, thus a mixture of hairs 

(9), 

distinctly 

Monteiro et al. 309 (INPA), 1 Mar 1976 (2), Ramos et different in length, moreover, with sparse redal. 

204 (INPA). AMAZONAS: Manaus, Reserva Florestal dish-brown, pluricellular hairs and sparse, white, 

Ducke, 27 Nov 1957 (st), D. Coelho (INPA) 5979 arachnoid hairs. Lamina entire, elliptic to ob- 

(INPA), 13 Sep 1957 (6), L. Coelho (INPA) 5808 (U); long, 13-30 x 6.5-20 cm, coriaceous, apex 

Manaus, waterfalls of Rio Taruma, 17 Oct 1929 (9), 

Ducke (RB) 23608 (B, G, RB, S, US, type collection abruptly, short-acuminate, base acute to roundof 

P. longipendula); Manaus, Reserva Florestal Ducke, ed to truncate; upper surface smooth, hairy on 

17 Oct 1929 (9), Killip et al. 30139 (A, F, NY, US); the main veins, to puberulous on the smaller 

Humaita, 7-18 Nov 1934 (2), Krukoff 7073 (A, B, BR, veins, lower surface yellow-(sub)hirsute on the 

F, G, GB, LE, MO, NY, RB, S, U, US); Sao Paulo de main 

Olivenca, 26 Nov 1936 (9), Krukoff 8695 

veins, to puberulous on the smaller veins, 

(A, B, BR, 

F, G, MO, NY, P, S, U, US); Manaus-P6rto Velho rd., white, arachnoid hairs in the areoles and on the 

km 240, 21 Nov 1973 (9), Lleras et al. P.19586 (F, smaller veins, lateral veins 10-16 pairs, the basal 

INPA, MO, NY, P, S, U, US); Manaus, Reserva Flo- pair unbranched or sparsely and/or faintly 

restal Ducke, 10 Nov 1965 (9), Loureiro (INPA) 16968 branched, tertiary venation almost plane be- 

(U); Torquato-Tapajos rd., km 118, 28 Aug 1975 (9), 

Loureiro et al. (INPA) 50623 (INPA); Manaus-Cara- neath; petiole 5-13.5 cm long, yellow-hirsute and 

carai rd., km 57, 16 Sep 1976 (9), Mota 680 (INPA); with sparse to dense, white, arachnoid hairs; stip- 

Tapuruquara, 4 Jul 1972 (9), Prance et al. 15393 (F, ules 3-9 cm long, caducous, outside yellow-hir- 

GH, INPA, M, NY, P, S, U); km 245, 13 Mar 1974 sute, inside glabrous. Staminate inflorescences 

(9), Prance et al. 20454 (INPA, MO, NY, S, U, US), 

km 380, 13 Oct 1974 (9), Prance et al. 22864 up to 19 cm 

(INPA, 

long and up to 14 cm wide; peduncle 

MO, NY, P, S, U, US); Manaus, Reserva Florestal 4-7 cm long, yellow-hirsute on the peduncle to 

Ducke, 21 Aug 1965 (9), Rodrigues 5449 (NY, U); hispidulous on the ultimate branches, sometimes 

Manaus-Itacoatiara rd., 3 Sep 1965 (9), Rodrigues et also white, arachnoid hairs; flowers sessile, not 

al. 7081 (U), 26 Nov 1965 (9), Rodrigues et al. 7292 

(U); Manaus-Caracarai rd., km 60, 20 Aug 1976 

densely clustered; tepals ca. 1.5 mm long, almost 

(6), 

Shima et al. 14 (INPA); between Rio Castanho and free, densely hispidulous; filaments shorter than 

Rio Tupanan, 14 Jul 1972 (8), M. F. Silva et al. 742 the tepals. Pistillate inflorescences unknown. 

(U); 18 Jul 1972 (9), M. F. Silva et al. 869 (INPA); Distribution (Fig. 85). Brazil (Amazonas); in 

Manaus-P6rto Velho rd., km 250, 5 Jan 1974 (9), Stew- non-inundated forest. 

ard et al. P.20156 (INPA, U). PARA: Without locality, 

(6), Ferreira s.n. (P, type collection of P. ovata); Cuiaba- Specimens examined. BRAZIL. AMAZONAS: Espe- 

Santarem rd., km 1230, (9), Prance et al. 25530 (F, ranga, mouth of Rio Janvari, 21 Sep 1931 (6), Ducke 

MO, RB, S, U); Rio Trombetas (6), N. T. Silva et al. (RB) 25246 (RB); Sao Paulo de Olivenca, nr. Palmares, 

4740 (MG, U). RORAIMA: Manaus-Caracarai rd., km 11 Nov-26 Oct 1936 (6), Krukoff 8388 (A, BR, F, G, 

343, 18 Nov 1977 (9), Steward et al. 107 (U). LE, MO, NY, P, S, U, type collection of P. elliptica).


I I 

STPE 

,E Fn CL1 |SOP Y 

r. 

Tr e rn rt. 'mh; trunk 41 G 

Safe o AmSaonm: r u.odeO 

near Plmamres. Sept. Il-OCt. 26. 1M91 

. 

.di- tb.td .: thlu th. New YO3 k 

hlod= 6 tc .'bre 4937 

FIG. 90. Pourouma elliptica. Leafy twig with staminate inflorescence 

(Krukoff 

8388).


Vernacular names. Brazil. Amazonas: mapati 

or mapaty. 

22. Pourouma bolivarensis C. C. Berg, Brittonia 

34(1): 39, t. 3. 1982. Type. Venezuela. Bolivar: 

Cerro Venamo, SW part, upper Rio Venamo, 

10 Jan 1964 (9), Steyermark et al. 92936 (ho- 

lotype, VEN; isotypes, NY, U). Fig. 91. 

5(1): 278, t. 14b. 1975; Burger, Fieldiana Bot. 

40: 202, t. 22. 1977. Type. French Guiana. 

Saint Jean de Maroni, 17 Mar 1914 (9), Benoist 

960 (lectotype, P, chosen here, cf. Berg, Fl. 

Suriname 5(1): 278. 1975). Fig. 82b. 

Pourouma aurea Ule ex Mildbraed, Notizbl. Bot. Gart. 

Berlin-Dahlem 10: 418. 1928; Ule, Bot Jahrb. Syst. 

40:149. 1907, nomen. Type. Brazil. Acre: Rio Jurua- 

Mirim, Aug 1901 (s), Ule 5718 

Tree, up to 15 m tall. 

(holotype, B; iso- 

Leafy twigs 5-8 mm types, G, MG). 

thick, puberulous. Lamina entire, broadly ellip- Pourouma folleata Macbride, Publ. Field Mus. Nat. 

tic to obovate, sometimes broadly ovate or sub- Hist., Bot. Ser., 8(2): 114. 1930, l.c. 13(2.2): 292. 

orbicular, 5-19 x 3-14 1937. 

cm, coriaceous, apex Type. Peru. Junin; Chanchamayo valley, 1924- 

1927 

shortly acuminate to rounded to emarginate, base 

(8), C. Schunke 416 (holotype, F; isotype, B). 

Pourouma isophlebia Standley, Publ. Field Mus. Nat. 

acute to obtuse; upper surface smooth, sparsely Hist., Bot. Ser., 17: 182. 1937. Type. Brazil. Amawhite-hirtellous 

or glabrous on the midrib, lower zonas: Mun. Sao Paulo de Oliven9a, nr. Palmares, 

surface appressed whitish-pubescent on the main 11 Sep-26 Oct 1936 (9), Krukoff8037 (holotype, NY; 

veins, arachnoid hairs in the areoles and ? cov- isotypes, A, BR, G, LE, P, S, U, US). 


ering the smaller veins; lateral 

subplicata 

veins 6-10 

Standley, Publ. Field Mus. Nat. 

pairs, Hist., Bot. Ser., 17: 184. 1937. Type. Brazil. Acre: 

basal pair and sometimes the second pair Nr. mouth of Rio Macauhan, 4 Aug 1933 (9), Krukoff 

branched, tertiary venation plane or slightly 5282 (holotype, F; isotypes, A, G, LE, M, MO, NY, 

prominent beneath; petiole 1-6.5 cm long, pu- S, U, UC, US). 

berulous; stipules 2-6 


cm 

umbellata 

long, outside Standley, Publ. Field Mus. Nat. 

yellow- Hist., Bot. Ser., 17: 185. 1937. Type. Brazil. Amapuberulous, 

inside glabrous, caducous. Stami- zonas: Mun. Humaita, between Rio Livramento and 

nate inflorescences unknown. Pistillate inflores- Rio Ipixuna, 7-18 Nov 1934 (9), Krukoff 7071 (hocences 

in fruit up to 10.5 cm long and 2 cm wide; lotype, F; isotypes, A, BR, G, GB, LE, MO, NY, 

peduncle 3-7 cm long, peduncle and branches RB, S, U, US). 

Coussapoa emarginata Killip ex 

yellow-puberulous; flowers 7-12; pedicel 0.2-1 

Macbride, Publ. Field 

Mus. Nat. Hist., Bot. Ser., 13(2.2): 296. 1937. Type. 

cm long, in fruit up to 2 cm; perianth ca. 3 mm Peru. Loreto: Mishuyacu, nr. Iquitos, 24-28 Sep 1929 

long, puberulous; stigma peltate, more or less (9), Killip & A. C. Smith 29955 (holotype, US; isobilobed, 

2-4 mm diam. Fruiting perianth ma- types, B, F). 

roon-red, ca. 1.8 x 1 

Pourouma cuatrecacasii 

cm, puberulous. 

Standley in Cuatrecasas, Revista 

Acad. Colomb. Ci. Exact. 

Distribution 

9(36/37): 339. 1956. 

(Fig. 85). Venezuela (Bolivar); in Type. Colombia. Vaupes: Mandi, nr. Mitfi, 24 Oct 

humid, (sub)-montane forest, at altitudes be- 1939 (9), Cuatrecasas 7299 (holotype, US). 

tween 900 and 1350 m. 

Pourouma umbellifera Burger, Phytologia 26: 430. 1973. 

Type. Costa Rica. Heredia: Nr. Rio Sarapiqui, Ti- 

Specimens examined. VENEZUELA. BOLIVAR: Cer- rimbina, 21 Jan 1972 (9), Lent 2327 (holotype, F; 

ro Venamo, Rio Venamo, 8 Jan 1964 (9), Steyermark isotypes, B, US). 

et al. 92850 (U, VEN); Cerro Venamo, SW part, upper 

Rio Venamo, alt. 950-1000 m, 10 Jan 1964 (9), Stey- Tree, up to ca. 35 m tall. Leafy twigs 2-13 mm 

ermark et al. 92936 (NY, U, VEN, type collection of thick, yellow-sericeous to -hirsute, or appressed- 

P. bolivarensis); Rio Cuyuni, Rio Anawaray-parui, 1300- 

1350 

puberulous, 

m, 25 Dec 1970 occasionally almost 

(st), Steyermark et al. 

glabrous, some- 

104466 

times with 

(NY, VEN). 

sparse brown, pluricellular hairs. 

Lamina entire, narrowly to broadly obovate to 

Vernacular names. Venezuela. Bolivar: cay- (broadly) elliptic, rarely to oblong, 4-45 x 1-15 

wari-cay-yek (Arekuna). 

cm, coriaceous to chartaceous, apex (long) acu- 

This species resembles P. minor, from which minate to mucronate or rounded to emarginate, 

it differs in the sparser indument, branched pis- base acute to rounded, rarely to retuse; upper 

tillate inflorescences, and branched basal lateral surface smooth, yellow-sericeous, hirtellous, hirveins. 

sute, or puberulous on the midrib, occasionally 

brown, pluricellular hairs on the main veins, low- 

23. Pourouma minor Benoist, Bull. Mus. Nat. er surface yellow-sericeous to -velutinous or to 

Hist. (Paris) 30: 103. 1924; Berg, Fl. Suriname -hirsute on the main veins (minutely) puberulous


. . . 

FIG. 91. Pourouma bolivarensis. Leafy twig with pistillate inflorescences (Steyermark et al. 92936); sterile 

leafy twig (Steyermark et al. 104466). 

to hirtellous on the smaller veins, orange, pluri- 

cellular hairs present or not, arachnoid hairs in 

the areoles, usually ? extending to the smaller 

veins; lateral veins 6-26 pairs, basal pair un- 

branched, tertiary venation (almost) plane be- 

neath; petiole 0.7-15 cm long, yellow-velutinous 

to -sericeous or hirsute; stipules 1-15 cm long, 

outside yellow-sericeous to -velutinous, some-


times -hirtellous, sometimes with orange-brown, (F, VEN), (2), Steyermark 60414 (F, VEN); Rio Chipluricellular 

hairs, inside glabrous, caducous. canan, Puerta Lema, SE of El Dorado, 24 Aug 1961 

Staminate inflorescences up to 12 cm long and 

(2), Steyermark 89506 (NY, U, VEN); Venezuelan- 

Brazilian frontier, Serrania 

7.5 cm wide; peduncle 1.5-8.5 cm 

Pia-soi, 5-6 Jan 1962 (st), 

long, yellow- Steyermark 90620 (VEN). 

sericeous to -velutinous; flowers sessile or pedi- GUYANA. Kanuku Mts., Iramaipang, Nov 1948 (6), 

cellate, diffusely distributed along the ultimate Wilson-Browne 549 (=FD 5948) (NY). 

branches or occasionally + clustered; tepals 1.5- SURINAM. Kabalebo, 12 Nov 1976 (st), Heyde et 

al. 32 

3 mm long, connate, puberulous to almost (U), 13 Nov 1976 (9), Heyde et al. 39 

gla- 

(U); River 

Nassau, Mts, 16 Feb 1949 (2), Lanjouw et al. 2214 

brous; filaments shorter than the tepals. Pistillate (NY, U); Paris Jacob Creek, 29 Jun 1965 (st), Maas et 

inflorescences in fruit up to 12 cm long and 7 cm al. (LBB) 11027 (U); Tapanahony R., at mouth of 

wide, subumbellate; peduncle 1-9 cm long, yel- Paloemeu R., Aug 1959 (st), Schulz 8161 (U); Goddo, 

low-sericeous; flowers 2-11; pedicel 0.2-1.2 cm 

26 Jan 1926 (9), Stahel (Exp. Wilhelminageb.) 82 (F, 

long, in fruit up to 3.5 cm; perianth 2-6 mm 

U); Voltzberg, Troon (LBB) 16310 (U). 

FRENCH GUIANA. St. Laurent, 13 Jan 1950 (9), 

long, usually densely hairy; stigma knob-shaped, BAFOG 5042 (P), 2 Jan 1956 (2), BAFOG 7152 (CAY, 

variously lobed, 2.5-6 mm diam. Fruiting peri- NY, P, U), 16 Jan 1956 (2), BAFOG 7173 (CAY, NY, 

anth ovoid with an apiculate apex, ca. 1-2.5 x P, U), 8 Feb 1956 (2), BAFOG 7261 (NY, P), 10 Feb 

1-1.5 

1956 

cm, reddish, 

(2), BAFOG 7268 

hairy. 

(CAY, NY, P, U); Saint Jean 

de Maroni, 17 Mar 1914 

Distribution 

(2), Benoist 960 

(Fig. 85). From Nicaragua to the 

(P, lectotype 

collection of P. minor); 16 Mar 1914 (2), Benoist 978 

Amazon Basin and the Guiana regions; in non- (P); Aug 1973 (st), Garnier 112 (CAY); between Saut 

inundated forest at altitudes up to ca. 1500 m. Emerillon and Etats-Unis, 21 Aug 1970 (st), Granville 

581 (CAY, P, U); Trois Sauts, 3 Nov 1974 (9), Lescure 

Specimens examined. NICARAGUA. Rio San Juan, 422 (CAY); upper Approuage R., Creek Par6pou, 7 

nr. Caino Chotaleno, 20 km NW of El Costillo, 7-9 Feb 1967 (st), Oldeman 2504 (CAY, NY, P, U); Ap- 

Mar 1978 (st), Neill 3389 (BG). 

prouage R., Pierrette, 14 Sep 1968 (st), Oldeman 2821 

COSTA RICA. ALAJUELA: 15 Sep 1973 (6), Hart- (CAY, P); Godebert, without date (st), Wachenheim 

shorn 1295 (F); Corazon de Jesus, 29 Mar (6), Hart- 18 (P), 23 Jun 1921 (6), Wachenheim 467 (P), 13 Jul 

shorn 1431 (F, U). HEREDIA: Tirimbina, 12-15 Aug 1921 (st), Wachenheim s.n. (A, E, U, US). 

1971 (st), Burger et al. 8137 (F); Las Horquetas, 25 ECUADOR. NAPO: Rd. Coca-Lago Agrio, 9 km NE 

May 1973 (6), Hartshorn 1224 (F, MO, U); nr. Rio of Rio Coca, 20 Mar 1980 (st), Brandbyge et al. 30252 

Sarapiqui at Tirimbina, 21 Jan 1972 (Q), Lent 2327 (F, (AAU); Guayusa, 2 hr upstream Rio Coca from Coca, 

GH, MO, NY, U, US, type collection of P. umbelli- 24 Mar 1980 (st), Brandbyge et al. 30327 (AAU); Afnfera); 

La Virgen, 23 Jul 1979 (6), Stevens 13340 (U). angu, 30 May-21 Jun 1982 (st), SEF8617, 9033, 9159 

LIMON: Between Siquirres and Turrialba, 10 Oct 1972 (U); MORONA-SANTIAGO: nr. Bomboiza, 27 Sep 1985 

(st), Holdridge 6818 (F, MO, U); Guapiles, 12-13 Mar (st), Shakaim RBAE-26 (BG). 

1924 (st), Standley 37121 (US); Cairo, 18-19 Feb 1926 PERU. AMAZONAS: Rio Santiago, nr. Caterpiza, 2 

(juv), Standley et al. 48600 (US). 

Nov 1979 (6), Huashikat 1150 (U); mouth of Rio San- 

PANAMA. COCLE: El Cope, 1 Sep 1977 (6), Berg tiago, 8-13 Oct 1962 (6), Wurdack 2169 (F, G, GH, 

400 (U); 25 Nov 1977 (st), Folsom et al. 6469 (U). SAN NY, P, S, UC). HUANUCO: Rio Pachitea, Tournavista, 

BLAS: Cerro San Jose, Yar Birea, 5 Feb 1986 (st), Nevers 8 Aug 1967 (6), J. Schunke V. 2139 (F, G, GH, 

et al. 6997 

NY, 

(MO). 

US); Prov. Leoncia Prado, Tingo Maria, 17 Feb 1964 

COLOMBIA. AMAZONAS-VAUPES: Rio Apaporis, (st), Vasquez 17 (F). JUNiN: San Ram6n, Aug 1925 (6), 

between Rio Kananari and Rio Pacoa, 1-15 Dec 1951 C. Schunke A-95 (F, NY, US); Chanchamayo Valley, 

(2), Garcia-Barriga 13836 (NY), 28 Sep 1951, Schultes 1924-1927 (8), C. Schunke 416 (B, F, type collection 

et al. 14148 (U). ANTIOQUIA: Between Dos Bocas and ofP.folleata). LORETO: Puerto Almendras, 7 Aug 1973 

Anori, 24-31 May 1973 (6), Soejarto et al. 4063 (A, (st), Ayala 341 (U); Rio Nanay, halfway between Iqui- 

MO). CHOCO: Novita, Cerro Torra, 22 Feb 1977 (2), tos and Santa Maria de Nanay, 31 

Forero 

May 1975 

et al. 3145 

(st), Gen- 

(RB, U). META: Sierra de la Ma- try et al. 22381 (U), 23 Mar 1979 

carena, 19 

(st), 

Jan 1950 

Gentry et al. 

(6), Philipson et al. 2134 (US). 26049 (U), 24 Mar 1979 (st), Gentry et al. 26177 

VALLE: Rio 

(U); 

Anchicaya, Alto Yunda, 1000 m, May 1972 Prov. Maynas, Yanamono Explorama Tourist 

(Q), Hilty M-86 

Camp, 

(MO). VAUPES: Mandi, nr. Mitfi, 24 1 Jul 1983 (st), Gentry et al. 42568 

Oct 

(BG); 

1939 

Mishuyacu, 

(2), Cuatrecasas 7299 (US, type collection of nr. Iquitos, 24-28 Sep 1929 (2), 

P. 

Killip & Smith 29955 

cuatrecacasii); Rio Inirida, San Joaquin, 27 Jan 1955 (US, B, F, type collection of Coussapoa 

(6), Fernandez 2019 

emarginata); 

(US). 

Yurimaguas-Tarapoto rd., km 14, 8 Nov 1967 (2), 

VENEZUELA. AMAZONAS: El Dorado-Sta. Elena Soria S. 21 (F, G, US). MADRE DE DIos: Rio 

rd., S of El 

Manu, 

Dorado, km 79, 25 Feb 1959 (2), Bernardi Cocha Cashu station, 2 Sep 1976 

7250 

(9), Foster et al. 5019 

(F, G, NY, VEN); Sierra Parima, Simarawochi, (F), 23 Oct 1979 (6), Foster et al. 7199 

18 Apr-23 May 1973 

(F); Rio Tam- 

(Q), Steyermark 107064 (F, U). bopata, mouth of Rio 

BOLiVAR: 

D'Orbigny, 4 Mar 1981 

Between Santa 

(st), 

Teresita de Kavanayen and Gentry et al. 31938, 31976 

Rio Pacairao, 20-21 

(U); Rio 

Nov 1944 

Manu, Cocha Ca- 

(6), Steyermark 60399 shu Station, 8 Aug 1983 (st), Gentry et al. 43454 (BG);


upper Rio Madre de Dios, 17 Jun 1954 (6), Rauh 1609 Thomas et al. 4794 (BG). PARA: Cuiaba-Santarem rd., 

(F). UCAYALI: Pucallpa-Tingo Maria rd., San Alejan- km 919, 13 Nov 1977 (9), Prance et al. 25336 (F, RB, 

dro, Gentry et al. 16181 (F, MO, NY); Pucallpa-Tingo U). RONDONIA: Mun. Presidente Medici, rd. Cuiaba- 

Maria rd., km 88, 13 Mar 1982 (st), Gentry et al. 36309 P6rto Velho, km 300, 25 Jun 1984 (d), Cid et al. 4798 

(BG). 

(BG); Mun. Ouro Preto, rd. Cuiaba-P6rto Velho, km 

BRAZIL. ACRE: Upper Rio Moa, Fazenda Arizona, 353, 39 Jun 1984 (6), Cid et al. 4919 (BG); Mun. Pi- 

10-15 Oct 1985 (st), Campbell et al. 6250 (BG); Abuna- menta Bueno, Guapor6, 7 Nov 1964 (st), Vieiro et al. 

Rio Branco hwy., km 242-246, 18 Jul 1968 (6), Forero 985 (U). RORAIMA: Serra dos Surucucus, 19 Feb 1969 

et al. 6367 (C, INPA, MO, NY, S, UC, US); nr. mouth (2), Prance et al. 10089 (G, INPA, NY, R, S, U, US), 

of Rio Macauhan, tributary of Rio Yaco, 4 Aug 1933 20 Feb 1969 (2), Prance et al. 10124 (INPA, NY, S, 

(9), Krukoff 5282 (A, G, LE, M, MO, NY, S, U, UC, U, US); Rio Negro, left bank of Parana do Marara, 7 

US, type collection of P. subplicata); rd. Brasileia-As- Feb 1977 (9), M. R. Santos 172 (MG, U). 

sis, km 16, 3 Nov 1980 (9), Lowrie et al. 710 (U); Sena BOLIVIA. BENI: Rio Beni, Cachuela Esperanza, 29 

Madureira, 3 Oct 1968 (9), Prance et al. 7792 (GH, Nov 1922 (6), Meyer 403 (U). PANDO: Barrola, 17 km 

INPA, M, NY, P, R, S, U); Rio Jurua-Mirin, Aug 1901 from Cobija, 18 May 1977 (st), Meneces 613 (MG, 

(6), Ule 5718 (B, G, MG, type collection of P. aurea). MO); San Francisco, 50 km from Cobija to Puerto 

AMAPA: Confluence of Rio Oiapoque and Rio Ian6, 27 Rico, 1 Jun 1977 (6), Meneces 647 (MO); rd. Cobija 

Aug 1960 (6), Irwin et al. 47874 (F, FHO, GH, MG, to Extrema, Villa Marieta, 24 Jun 1978 (2), Meneces 

NY, RB, US); Rio Oiapoque, nr. mouth of Rio Ya- 725 (INPA); 3 km above Abuna, 13 km 1968 (a), Prance 

roupi, 24 Sep 1960 (6), Irwin et al. 48468 (M, MG, et al. 8378 (B, INPA, NY, S, U, UC, US); Rio Na- 

NY, P, RB, UC); Rio Oiapoque, nr. Cachoeria Santa reuda, Nicolas Suarrez, SW of Cobija, 31 Jul 1982 

Maria, 21 Aug 1961 (6), Pires et al. 50419 (FHO, G, (5), Sperling et al. 6430 (BG). 

MG, NY, S, US); 40 min up Rio Falcino, 14 Nov 1961 

(8), Pires et al. 50941 (F, MO, NY); Serra do Navio, Vernacular names. Colombia. Vaupes: uva sil- 

26 Sep 1961 (Q), Pires et al. 51247 (GH, MG, NY, MO, vestre. Venezuela. Bolivar: majagua, cay-wari- 

U, US); AMAZONAS: Manaus-Itacoatiara rd., Reserva 

Florestal Ducke, 6 Aug 1976 (st), Aluisio (INPA) 70809 

cay-yek, coiwaricoi-yek, kai-wa-rei-kei-yek. Su- 

(INPA); Uaup6s, Serra Sao Gabriel, 14 Feb 1959 

rinam. 

(9), Bospapaja, granboesipapaja. French 

Cavalcante 606 (MG); Rio Uatumi, Itapiranga, 24 Aug Guiana. Bois canon, bois canon male or male 

1979 (6), Cid et al. 721 (INPA, U); Rio Jurua, Sta. bois canon, bouchi papaie and papaye (Para- 

Rosa, 15 Aug 1975 (st), D. Coelho et al. (INPA) 51350 maka), kulumasi (Wayapi). Peru. Amazonas: 

(INPA), 22 Aug 1975 (st), D. Coelho et al. (INPA) sacha 

52381 (INPA); Manaus-Itacoatiara rd., Reserva Flo- uvila, shuvija (Aguaruna); Huanuco and 

restal Ducke, 28 Sep 1962 (8), Duarte 7194 (INPA, M); 

San Martin: uvilla or uvilla blanca; Loreto: uvilla 

Manicor6, nr. Santa F6, 8-11 Sep 1934 (9), Krukoff lanuda. Brazil. Amapa: mapatirana; Amzonas: 

6041 (A, B, BR, F, LE, MO, NY, S, U, US); Humaita, imbaibarana, purumai, tourem; Mato Grosso: 

plateau between Rio Livramento and Rio Ipixuna, 7- 

18 Nov 1934 (9), Krukoff 7071 

caramuri, imbaubarana, torena. Bolivia. Pando: 

(A, BR, G, GB, LE, 

MO, NY, RB, S, S, U, US, type collection of P. um- ambaibochi, ambaibillo. 

bellata); Sao Paulo de Olivenca, nr. Palmares, 11 Sep- The species is rather variable in the dimen- 

26 Oct 1936 (9), Krukoff 8073 (A, BR, G, LE, MO, sions and shape of the leaves. The indument 

NY, P, S, U, US, type collection ofP. isophlebia); upper mostly consists of appressed hairs but patent hairs 

Rio Negro Uaupes, 12 Feb 1959 (9), Rodrigues 884 

(U); Tefe, 24 Nov 1959 

occur in the 

(st), Rodrigues et al. 1424 

Upper Amazon Basin, e.g., in the 

(INPA); Manaus-Itacoatiora rd., Reserva Florestal type collection ofP. aurea. The number of lateral 

Ducke, 19 Oct 1965 (6), Rodrigues 7532 (INPA); Tefe, veins varies from 6 to ca. 25 pairs, but in some 

23 Aug 1979 (6), Rodrigues et al. 10182 (INPA); Ma- collections (of juvenile material?) probably benaus-P6rto 

Velho rd., between Rio Castanho and Rio 

Tapuna, 18 Jul 1972 (8), M. F. Silva et al. 903 

longing to P. minor, e.g., Ayala 341 and Gentry 

(INPA). et 

MATO 

al. 

GROSSO: Aripuana, 18 Jun 1975 (8), Cordeiro 22381, the number of lateral veins may rise 

140 (US), 25 Jan-14 May 1977 (st), Gomes et al. 547, to 40 pairs. 

593, 632, 635, 646, 655, 673, 690, 759, 789, 810, 813, The species is a clear-cut one, characterized 

838, 872, 891, 924, 926, 931, 933, 955, 1033, 1118, by the subumbellate pistillate inflorescence and 

1120, 1129, 1156, 1175, 1255, 1319, 1336, 1338, 1356, the knob-like 

1378, 1419, 1423, 1425, 

stigma. It shows resemblances of 

1444, 1446, 1534, 1568, 1570, 

1577, 1584, 1593, 1594, 1595, 1636, 1640, 1641, 1692, 

P. bolivarensis (see p. 186). 

1731, 1745, 1769, 1822, 1823, 1867, 1893, 1917, 1983 The fruits are eaten by the monkey Cebus apel- 

(INPA), 14 Jul 1977 (6), Gomes et al. 2170 (INPA), 19 la (Foster et al. 5019). 

Jan 1979 (9), 29 May 1976 (6), Monteiro et al. 1119 From the three syntype collections from French 

(INPA), M. G. Silva et al. 4326 (MG), 1 Jan 1979 (6), 

M. G. Silva et al. 4738 (MG), 8 Oct 1979 

Guiana 

(9), Rylands 

(Benoist 960, Benoist 978, and Wach- 

25 (INPA); Mun. Vila Bela da Santissima Trinidade, enheim 18) Benoist 960 has been selected as the 

4 km S of border with Rond6nia, 3 Nov 1985 (2), lectotype collections (Berg, 1975).


Pourouma napoensis and P. herrerensis have 

been included in this revision after submission 

of the (pre-review) manuscript and are, therefore, 

placed at the end of the systematic treatment, 

and not after P. hirsutipetiolata and P. mollis, 

respectively. 

24. Pourouma napoensis C. C. Berg, Brittonia 

42: 59-65. 1990. Type. Ecuador. Napo: Re- 

serva Biologia Jatun Sacha, 14 Aug 1987 (6), 

Palacios 1874 (holotype, QCNE; isotypes, 

AAU, BG, MO, NY, QAME). Fig. 92. 

Tree, up to 25 m tall. Leafy twigs 1-2.5 cm 

thick, yellow-hirsute, pluricellular hairs absent. 

Lamina 7-fid to -parted, ca. 25-45 x 25-45 cm, 

coriaceous, apex acuminate, base deeply cordate 

with overlapping lobes; upper surface smooth to 

slightly scabrous, yellow-hirsute to -hirtellous on 

the whole surface or only on the main veins, 

lower surface yellow-hirsute to -hirtellous on the 

veins, arachnoid hairs in the areoles and on the 

reticulum; lateral veins 16-25 pairs; petiole ca. 

25-40 cm long, yellow-hirsute; stipules 10-16 cm 

long, outside yellow-hirsute, inside glabrous, caducous. 

Staminate inflorescences ca. 12-22 cm 

long, ca. 8-14 cm wide; peduncle 4-8 cm long, 

peduncle and branches yellow-hirtellous to -pu- 

In the shape, dimensions, and indument of the 

lamina this species resembles P. oraria. It differs 

from the latter in the staminate flowers with their 

connate tepals and fully connate filaments. These 

features suggest a close relationship to P. hirsu- 

tipetiolata. 

25. Pourouma herrerensis C. C. Berg, Candollea 

44(2): 513. 1989. Type, Peru, Loreto, Prov. 

Requena, Jenaro Herrera, Reserva Forestal, 

tree 4/122, 2 Oct 1985 (d), Spichiger et al. 1995 

(holotype, G; isotype, BG). Fig. 93. 


thick, minutely puberulous to hispidulous (and 

+ scabrous) or also with much longer, yellow 

hairs, pluricellular hairs dark brown and rather 

sparse. Lamina entire, oblong to elliptic to ovate, 

8-20 x 3-11.5 cm, (sub)coriaceous, apex shortacuminate, 

base acute to rounded; upper surface 

sparsely appressed-puberulous on the midrib or 

the whole surface covered with long, yellow hairs, 

lower surface appressed-puberulous on the main 

veins or also with long yellow hairs, arachnoid 

hairs in the areoles and on the reticulum; lateral 

veins 9-14 pairs, basal pair unbranched or 

branched, the basal part of these veins forming 

the basal part of the leaf margin (thus not separated 

from the margin by mesophyll), tertiary 

venation slightly prominent; petiole 3-10 cm 

berulous; flowers (sub)sessile, in 

long, minutely puberulous or also with long, yelglobose 

heads, low 

these 7-12 mm diam.; perianth, tubular, 1-2 mm 

hairs; stipules 3.5-8 cm long, caducous, outside 

high, puberulous at the apex; stamens 3-4, filadensely 

puberulous and with brown, pluricellular 

ments completley connate, 4-6 mm long. Pistilhairs, 

or also with long, yellow hairs, 

sometimes 

late inflorescences up to 10 cm long and up to 

only a few appressed ones, inside 

12 cm wide, branched; peduncle 3-5 cm sparsely short-pubescent. Staminate infloreslong, 

cences 

peduncle and branches yellow-hirsute to -hirtel- up to 10 cm long and up to 9 cm wide; 

lous; flowers ca. 40-70; pedicel 0.3-0.5 cm, in 


fruit up to 1.2 cm long; perianth ca. 5 mm densely puberulous to short-velutinous or also 

long, with 

yellow-hirsute to -hirtellous, sparsely so in the long, yellow hairs; flowers sessile or pedicellate, 

most of them in globose heads, numerous 

uppermost part; stigma peltate. 

and 3-4 mm diam. or less 

Distribution (Fig. 81). Ecuador (Napo), in non- 

(5-12) and 5-8 mm 

inundated forest, at low altitudes. 

diam.; tepals ca. 1 mm long, connate, forming a 

+ urceolate perianth, densely hairy; filaments 

Specimens examined. ECUADOR. NAPO: Reserva exceeding the tepals, free. Pistillate inflorescences 

Biologica Jatun Sacha, Rio Napo, 8 km below Puerto up to 14 cm long and 9 cm wide; peduncle and 

Misahualli, 4 Sep 1987 (2), Cer6n M. 2019 (BG); 8 km branches short-velutinous; flowers 7-25, in 1-4 

SE of Tena, 30 Aug 1960 (2), Grubb et al. 1545 (K, 

NY); Reserva Biologica Jatun Sacha, Rio Napo, 8 km 

+ distinct clusters, pedicels up to 0.7 cm long; 

below Puerto Misahualli, 2 Oct 1986 (2), Palacios 1337 perianth yellow-velutinous, except for the apex; 

(BG), 14 Aug 1987 (a), Palacios 1874 (BG, type col- stigma subpeltate. 

lection of P. napoensis); Lumbaqui, 12 May 1987 (a), Distribution (Fig. 81). Peru (Loreto), in non- 

Pennington et al. 12244 (BG, K). 


Specimens examined. PERU. LORETO: Prov. Requena, 

Jenaro Herrera, Aug-Sep 1976 (6), Bernardi s. n.


bi 

f 

In 

i 

FIG. 92. Pourouma napoensis. a. Leaf(Grubb et a. 1545). b. Stipules and staminate inflorescence (Pennington 

et al. 12244). c. Pistillate inflorescence (Grubb et al. 1545). d. Staminate flower (Pennington et al 12244). e. 

Pistillate flower (Ceron M. et al 2019).

.At 

FIG. 93. Pourouma herrerensis. a. Leafy twig with staminate inflorescences (Spichiger et al. 1995). b. Pistillate 

inflorescence (Spichiger et al. 1999). c. Leaf (Vasquez et al. 2623).


(BG), 9 Dec 1977 (d), Gentry et al. 21346 (U); Prov. 

Maynas, Puerto Almendras, 11 Nov 1984 (9), Maas et 

al. 6267 (BG, U); Prov. Requena, Jenaro Herrera, Reserva 

Forestal, 2 Oct 1985 (d), Spichiger et al. 1995 

(BG, G, type collection), 17 Sep 1982 (6), Spichiger et 

al. 1996 (BG, G), 9 Oct 1982 (st), Spichiger et al. 1997 

(BG, G), 14 Sep 1982 (d), Spichiger et al. 1998 (BG, 

G), 10 Nov 1982 (9), Spichiger et al. 1999 (BG, G); 

Prov. Maynas, Puerto Almendras, 17 Apr 1980 (6), 

Vdsquez et al. 167 (BG), 21 Oct 1981 (9), Vdsquez et 

al. 2623 (BG); Prov. Maynas, Rio Nanay, Mishana, 24 

Apr 1986 (9), Vdsquez et al. 7535 (BG). 

This species has a strange mixture of features. 

The staminate inflorescences and flowers resemble 

those of P. mollis, the pistillate inflorescences 

and flowers those of P. mollis subsp. triloba and 

P. acuminata. In the indument it differs from 

both. One of the forms of this species has very 

short and stiff hairs, resulting in a slightly scabrous 

surface of the leafy twigs, petioles, stipules, 

and main veins of the lamina beneath. The other 

form has this short indument mixed with much 

longer yellow hairs, as in P. cucura, and bears, 

moreover, long yellow hairs on the upper leaf 

surface, like some forms of P. cucura. Some of 

the specimens have oblong to elliptic leaves with 

Unnamed Collections 

1. PERU. AMAZONAS: Serrania de Bagua, ca. 12- 

18 km on trail E of La Peca, 1800-1950 m, lower 

montane cloud forest, 14 Jun 1978 (6), Gentry et al. 

22866 (U). SAN MARTIN: Prov. Rioja, rd. Pedro Ruiz- 

Moyobamba, km 390, Venceremos, 2100 m, 7-9 Aug 

1983 (st), D. N. Smith et al. 4742 (MO). 

The indument of the leafy twigs, stipules, petioles, 

and lower surface of the lamina largely 

matches that of P. bicolor subsp. bicolor. The 

specimens differ from that taxon in the smooth 

and puberulous upper surface of the lamina, the 

broadly ovate lamina, the slender (ca. 15 x 10 

cm) staminate inflorescence, the oblong to elliptic 

193 

tepals of the staminteflower, and the sparse pluricellular 

hairs. 

These specimens may represent a distinct cloud 

forest species (see p. 114). 

2. PERU. LORETO: Rio Corrientes, between Tiente 

Lopez and Puesto Avanzado, 4 Apr 1977 (st), Gentry 

et al. 19048 (BG); Santa Rosa, 1-5 Sep 1929 (st), Killip 

& A. C. Smith 28800 (F, NY). 

BRAZIL. ACRE: Reserva INCRA Santa Luzia, BR- 

364, km 40, 5-19 Oct 1984 (st), Campbell et al. 6820 

(NY) and 7689 (BG). 

These have ovate to elliptic leaves (16-32 x 

12-22 cm) with a deeply cordate base. The leafy 

twigs, petioles, stipules, and main veins at the 

lower surface of the lamina have dense, long, 

patent, yellow hairs. The stipules are glabrous 

inside. The presence of arachnoid hairs on the 

petiole, main veins of the lamina beneath, and 

stipules suggest affinities to P. tomentosa. These 

collections may represent a new subspecies of 

this species. 

3. VENEZUELA. AMAZONAS: Nr. Yavita, 21 Apr 

1970 (st), Steyermark et al. 102874 (BG, MO, NY, 

VEN); Dept. Rio Negro, Cerro de la Neblina, Rio Mathe 

basal pair of lateral veins unbranched; such warinuma, 8-10 Jan 1984 (st), Steyermarket al. 129773 

specimens bear some resemblance to P. acumi- (MO). 

nata. Other specimens (with long hairs) have These collections have thickly coriaceous 

ovate leaves and the basal pairs of lateral veins leaves, thick leafy twigs with conspicuous lentibranched. 

However, both types of leaves may cels, and dense, almost black, pluricellular hairs 

occur on the same plant, or even on the same on the leafy twigs and stipules. In these features 

branch. 

they match P. ferruginea, but the characteristic, 

The material with a scabrous upper leaf sur- dense (white to brownish) villous-arachnoid inface, 

provisionally kept in P. mollis subsp. tri- dument on the lower surface of the lamina is 

loba, resembles P. herrerensis in the variation of wanting and the stipules are densely hairy inside. 

the leaves. 

These collections might represent an abberant 

form of P. ferruginea or perhaps of P. melinonii 

subsp. melinonii. 

4. COLOMBIA. VAUPES: Alto Vaupes, 25 Nov 1975 

(9), Roa 251 (INPA). 

This collection has leafy twigs with yellow- 

hirsute indument and conspicuous lenticels. The 

indument of the leafy twigs and the leaf surface 

resembles that of P. cucura, but the thickly co- 

riaceous, somewhat plicate lamina has the hairs 

on its upper surface confined to the main veins. 

Moreover, the stipules are hairy inside and the 

reticulum on the lamina beneath is very prom- 

inent. The fruiting perianth is scabrous. This 

specimens has a mixture of characters of P. cu- 

cura and P. bicolor subsp. bicolor.

194 

5. ECUADOR. NAPO: Aiiangu, 1-15 Feb 1986 (st), 

Korning & Thomson 47801 and 47807 (BG). 

The latter has most characters in common with 

the form of P. bicolor subsp. bicolor with an entire 

lamina, being smooth above. However, the 

indument of the leafy twigs, petiole, and lower 

leaf surface resembles that of P. cucura. The former 

collection with 5-parted leaves and incisions 

down to the petiole may represent the juvenile 

form of the latter collection. The mixture of characters 

suggests possible hybridization. 

Names Excluded 

Pourouma paraensis-nomen (in schedule) given 

by Huber; refers to Goeldi (MG) 7732 = P. 

melinonii subsp. melinonii. 

Pourouma retusa Bentham-nomen, see Miquel 

in Martius, Fl. bras. 4(1): 128. 1853; refers to 

Spruce s. n. from Brazil, Amazonas, Manaussyntype 

collection of P. tomentosa. 

Pourouma trianae A. Richter-nomen (in schedula); 

refers to Triana 862 = P. cecropiifolia. 

Pourouma ulei Warburg ex Ule-nomen; see Ule, 

Bot. Jahrb. Syst. 40: 132. 1907; refers to Ule 

9314 = P. cecropiifolia. 

ACKNOWLEDGMENTS 

The authors of the Coussapoa revision (C.C.B. 

and R.W.A.P.A.) are much indebted to Dr. N. 

G. Bisset (Chelsea College, London) for correct- 

ing the English text. They thank Mr. Bonsen (In- 

stitute for Systematic Botany, Utrecht) for con- 

tributing the chapter on the wood and leaf 

anatomy, and Drs. E. C. H. van Heusden (In- 

stitute for Systematic Botany, Utrecht) for pre- 

paring the distribution maps. They are grateful 

to the curators of the cited herbaria for putting 

their collections at the disposal of the authors. 

Grants from the Netherlands Foundation for Ad- 

vancement of Tropical Research (WOTRO) to 

C.C.B. enabled part of the collections and data 

on which the treatment is based to the gathered 

during several South American trips. They also 

thank Dr. L. B. Holm-Nielsen (Botanical Insti- 

tute, Aarhus) for his help during fieldwork in 

Ecuador. The drawings were prepared by Ms. Siri 

Herland (Bergen), Miss E. M. Hupkens van der 

Elst (Utrecht), and Mr. T. Schipper (Utrecht). 

The revision of Pourouma was started by Drs. 


E. Kieft (Utrecht), and some elements of that 

unfinished study have been used by the present 

authors (C.C.B. and E.C.H.vH.). The senior au- 

thor is indebted to the University of Bergen, Fac- 

ulty of Mathematics and Natural Sciences, for 

covering traveling expenses, and to the Botanical 

Institute at Bergen for their assistance. The il- 

lustrations were prepared by Ms. Siri Herland 

(Bergen), Mr. H. Rypkema (Utrecht), and Mr. T. 

Schipper (Utrecht). The authors are indebted to 

Ms. Kathleen Welling for processing part of the 

text, to Drs. W. H. A. Hekking (Utrecht), Drs. 

E. Simonis (Utrecht), Ms. Edit Fjaere (Bergen) 

and Ms. Francina Berg (Bergen) for their assis- 

tance in preparing the list of exsiccatae, and to 

Drs. E. Simonis for his contributions to the preparation 

of the lists of specimens examined and 

the distribution maps. 

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29. 

Cuatrecasas, J. 1951. In J. A. Steyermark, Botanical .1942. Moraceae II. In L. Diels, Neue Arten 

exploration of Venezuela--I: Moraceae. Fieldiana aus Ecuador. Notizbl. Bot. Gart. Berlin-Dahlem 

Bot. 28(1): 210-216. 

15: 783-785. 

.1954. Dos Moraceas y dos Compuestas nue- Miquel, F. A. W. 1843. Observations botanicae de 

vas de Venezuela. Bol. Soc. Venez. Ci. Nat. 15: quibusdam plantis, quas in colonia Surinamensi 

107-111. 

legit vir. graviss. H. C. Focke. Het Instituut 2(1842): 

. 956a. Notas a la Flora de Colombia, XIV. 185-205, t. 3. 1843. 

Revista Acad. Colomb. Ci. Exact. 9(36/37): 325- . 1853. Urticales. In C. F. P. von Martius, 

341, with photographs. 

Flora Brasiliensis 4(1): Coussapoa: 131-139, t. 42- 

.1956b. Morfceas nuevas de Colombia. Cal- 45. Pourouma: 122-131, t. 36-41. 

dasia 7: 287-304. 

O'Dowd, D. J. 1982. Pearl bodies as ant food: An 

1956c. Notas a la Flora de Venezuela. Bol. ecological role of some leaf emergences of tropical 

Soc. Venez. Ci. Nat. 17: 80-97. 

plants. Biotropica 14(1): 40-49. 

. 1967. Moraceae. In J. A. Steyermark, Flora Pires, J. Murqa. 1973. Tipos de vegetaco da Amadel 

Auyan-tepui. Acta Bot. Venez. 2(5-8): 202- z6nia. O Museu Goeldi no Ano do Sesquicentena- 

205. 

rio. Belem. 

Ducke, A. 1932a. Neue Arten aus der Hylea Brasi- Pittier, H. 1917. New and noteworthy plants from


Colombia and Central America-6. Contr. U.S. 

Natl. Herb. 18: 225-259. 

.1943. Notasdendrologicasde Venezuela. VIII. 

Bol. Soc. Venez. Ci. Nat. 8(56): 257-264. 

Poeppig, E. F. & S. Endlicher. 1838. Nova genera ac 

species plantarum 2. Leipzig. 

Prance, G. T. 1977. The phytogeographic subdivisions 

of Amazonia and their influence on the selection 

of biological reserves. In G. T. Prance & 

T. S. Ellias (eds.), Extinction is forever. The New 

York Botanical Garden, New York. 

Renner, 0. 1907. Beitrige zur anatomie und systematiek 

der Artocarpeen und Conocephaleen, inbesondere 

der Gattung Ficus. Bot. Jahrb. Syst. 39: 

319-448. 

Ruiter, G. de. 1976. Revision of the genera Myrianthus 

and Musanga (Moraceae). Bull. Jard. Bot. 

Etat. 46: 471-510. 

Rusby, H. H. 1910. New species from Bolivia, collected 

by R. S. Williams- 1. Bull. New York Bot. 

Gard. 6: 487-517. 

1927. Descriptions of new genera and species 

of plants collected on the Mulford biological exploration 

of the Amazon valley. Mem. New York 

Bot. Gard. 7: 205-384. 

Smith, J. Donnell. 1905. Undescribed plants of Guatemala 

and other Central American republics. Bot. 

Gaz. 40: 1-11. 

NUMERICAL LIST OF TAXA 

1. Coussapoa 

1-1. C. angustifolia Aublet 

1-2. C arachnoidea Akkermans & C. C. Berg 

1-3. C. argentea Akkermans & C. C. Berg 

1-4. C. asperifolia Tr6cul 

a. subsp. asperifolia 

b. subsp. magnifolia (Trecul) Akkermans & C. 

C. Berg 

c. subsp. rhamnoides (Standley) Akkermans & 

C. C. Berg 

1-5. C. batavorum Akkermans & C. C. Berg 

1-6. C. brevipes Pittier 

1-7. C. chocoensis Cuatrecasas 

1-8. C. cinnamomea Cuatrecasas 

1-9. C. cinnamomifolia Mildbraed 

1-10. C. contorta Cuatrecasas 

1-11. C. crassivenosa Mildbraed 

1-12. C. cupularis Akkermans & C. C. Berg 

1-13. C. curranii Blake 

1-14. C. duquei Standley 

1-15. C. echinata Akkermans & C. C. Berg 

1-16. C. ferruginea Trecul 

1-17. C. floccosa Akkermans & C. C. Berg 

1-18. C. glaberrima Burger 

1-19. C. herthae Mildbraed 

1-20. C. latifolia Aublet 

1-21. C. leprieurii Benoist 

1-22. C. longepedunculata Akkermans & C. C. Berg 

1-23. C. macerrima Akkermans & C. C. Berg 

1-24. C. manuensis C. C. Berg 

1-25. C. microcarpa (Schott) Rizzini 

1-26. C. microcephala Trecul 

1-27. C. napoensis Akkermans & C. C. Berg 

Spix, J. B. & C. F. P. von Martins. 1831. Reise in 

Brasiliens 3. Miinchen. 

Standley, P. C. 1919. Studies of tropical American 

phanerogams-No. 3. Contr. U.S. Natl. Herb 20(6): 

173-220. 

1924. Nine new species of plants from Central 

America. Proc. Biol. Soc. Wash. 37: 49-54. 

. 1930. Studies of American plants-III. Publ. 

Field Mus. Nat. Hist., Bot. Ser., 8(1): 3-73. 

. 1933. Poulsenia, a genus oftrees ofthe family 

Moraceae. Trop. Woods 33: 4-5. 

- . 1937a. StudiesofAmericanplants-VII. Publ. 


1937b. Flora of Costa Rica: Moraceae. Publ. 


1939. In A. C. Smith, Notes on a collection 

of plants from British Guiana. Lloydia 2:161-218. 

1940. Studies of American plants-X. Publ. 


Trecul, A. 1847. Sur la famille des Artocarp6es. Ann. 

Sci. Nat. Bot., Ser. 3, 8: 38-157. 

Ule, E. 1907. Die Pflanzenformationen des Amazonas-Gebietes. 

Bot. Jahrb. Syst. 40: 114-172. 

Woodson, R. E. & R. W. Schery. 1960. Flora of Panama: 

Moraceae. Ann. Missouri Bot. Gard. 47(2): 

114-178. 

1-28. C. nitida Miquel 

1-29. C. nymphaeifolia Standley 

1-30. C. oligocephala Donnell Smith 

1-31. C. orthoneura Standley 

1-32. C. ovalifolia Tr6cul 

1-33. C. pachyphylla Akkermans & C. C. Berg 

1-34. C. parviceps Standley 

1-35. C. parvifolia Standley 

1-36. C. purpusii Standley 

1-37. C. scabra Akkermans & C. C. Berg 

1-38. C. sprucei Mildbraed 

1-39. C. tessmannii Mildbraed 

1-40. C. trinervia Mildbraed 

1-41. C. vannifolia Cuatrecasas 

1-42. C. villosa Poeppig & Endlicher 

1-43. C. viridifolia Cuatrecasas 

1-44. C. fulvescens C. C. Berg 

1-45. C. tolimensis C. C. Berg 

1-46. C. valaria C. C. Berg 

2. Pourouma 

2-1. P. guianensis Aublet 

a. subsp. guianensis 

b. subsp. venezuelensis (Cuatrecasas) C. C. Berg 

& van Heusden 

2-2. P. velutina Miquel 

2-3. P. bicolor Martius 

a. subsp. bicolor 

b. subsp. tessmannii (Mildbraed) C. C. Berg & 

van Heusden 

c. subsp. digitata (Tr6cul) C. C. Berg & van 

Heusden 

d. subsp. scobina (Benoist) C. C. Berg & van 

Heusden

Exsiccatae 197 

e. subsp. chocoana (Standley) C. C. Berg & van 

Heusden 

2-4. P. cecropiifolia Martius 

2-5. P. cucura Standley & Cuatrecasas 

2-6. P. cuspidata Mildbraed 

2-7. P. myrmecophila Ducke 

2-8. P. formicarum Ducke 

2-9. P. phaeotricha Mildbraed 

2-10. P. mollis Tr6cul 

a. subsp. mollis 

b. subsp. triloba (Tr6cul) C. C. Berg & van Heusden 

2-11. P. melinonii Benoist 

a. subsp. melinonii 

b. subsp. glabrata C. C. Berg & van Heusden 

2-12. P. hirsutipetiolata Mildbraed 

a. subsp. hirsutipetiolata 

b. subsp. hispida (Standley & Cuatrecasas) C. C. 

Berg & van Heusden 

2-13. P. tomentosa Miquel 

LIST OF EXSICCATAE 

a. subsp. tomentosa 

b. subsp. apiculata (Benoist) C. C. Berg & van 

Heusden 

c. subsp. persecta C. C. Berg & van Heusden 

d. subsp. essequiboensis (Standley) C. C. Berg & 


3. subsp. maroniensis (Benoist) C. C. Berg & 


2-14. P. stipulacea C. C. Berg 

2-15. P. acuminata Miquel 

2-16. P. ferruginea Standley 

2-17. P. villosa Trecul 

2-18. P. oraria Standley & Cuatrecasas 

2-19. P. saulensis C. C. Berg & Kooy 

2-20. P. ovata Tr6cul 

2-21. P. elliptica Standley 

2-22. P. bolivarensis C. C. Berg 

2-23. P. minor Benoist 

2-24. P. napoensis C. C. Berg 

2-25. P. herrerensis C. C. Berg 

Some recent collections have been added to this list without being cited in the lists of specimens. 

Acosta M., A., 12 (2-5). 

Acosta Solis, M., 6008 (1-10); 12661 (1-19); 12797 (1- 

10); 12884, 13605 (2-3d); 13629(1-42); 13687, 13713 

(1-10). 

Alencar, L., 497 (2-4). 

Allemao, F. et al., 446 (2-la). 

Allen, P. H., 3103 (39); 3251 (2-1 a); 3742 (2-3d); 

5120 (1-4b); 5949 (1-23); 6186, 6280 (1-42); 6401 

(2-3d). 

Aluisio, J., 70809 (2-23). 

Amaral, I. L. et al., 101 (2-11a); 473 (2-15); 478 (1- 

40); 595 (2-1 la); 604 (2-5); 666 (2-23); 761 (2-7); 

IG2-10-328, IG2-16-509 (2-5). 

Ameida, J. et al., 71 (2-10a). 

Ancuash, E., 178 (2-la); 290 (2-1 la). 

Anderson, C. W., 188, s.n. (1-26). 

Andrade, A. et al., 2340 (1-25). 

Andrade Lima, D. de, see Lima. D. 

Anonymus, 9783 (1-20); s.n. (2-4, 1-25). 

Appun, C. F., 296 (1-26). 

Araujo, D. S. Dunn de et al., 848 (1-25). 

Aristeguieta, L., 1699, 1704 (1-42); 2005 (2-16). 

Asanza C., E., 32929 (2-4). 

Asplund, E., 9462, 12933 (2-4); 13020 (2-la); 14124 

(2-4); 16401 (1-41); 18780 (1-4b); 18840(1-9); 19835 

(1-42). 

Aublet, J. B. C. F., s.n. (1-20, 2-la, 2-1 la, 2-17). 

Aubreville, A., 144 (2-3c). 

Aubry-Lecompte, C. E., s.n. (1-20). 

Austin, D. F. et al., 4140 (2-3a); 7149 (1-20). 

Ayala, F. (et al.), 341 (2-spec.); 1436 (1-42); 1446 (2- 

4). 

Aymard, G. et al., 3990 (2-23); 6778 (2-15). 

BAFOG, 93-M (2-17); 1050 (2-3c); 4272 (2-1 la); 5042 

(2-23); 7087 (2-17), 7152, 7173 (2-23); 7228 (2-13e); 

7261, 7268 (2-23); 7330 (2-13e); 7562 (2-2); 7745 

(2-17); 7938 (1-1); 7942, s.n. (1-4a). 

Bailey, I. W., 18, 19, 20, 146, 147 (2-la). 

Baker, M. A., 6017 (2-3a). 

Balick, M. et al., 1021 (1-39). 

Barbour, P. J., 5372 (1-42). 

Barclay, A. S. et al., 3294, 3593 (1-42). 

Barrier, S., 5114 (2-13e). 

Bartlett, H. H., 12356 (1-30). 

Bastos, M. A., 2051 (2-la); 2220 (2-23). 

BBS, 3003 (1-20). 

BBS (=Bosbeheer Suriname), 297 (2-17); 298 (2-13a); 

1128 (2-la). 

Belem, R. P. (et al.), 644 (1-25); 1471 (2-la). 

Benoist, R., 341 (2-17); 465 (2-13e); 960, 978 (2-23); 

3003 (2-3d); 3010 (1-10); 3042 (1-41); 3044 (1-42). 

Berg, C. C. (et al.), 276 (2-7); 355 (2-1 lb); 400 (2-23); 

BG.598, BG.772 (2-la); 1033 (1-42); 1038 (1-40); 

1040 (1-42); 1041, 1042, 1045 (2-5); 1048 (1-40); 

1053 (2-5); 1058 (1-40); 1062 (1-42); 1070 (2-5); 

1111 (1-31); 1113 (1-42); 1119 (1-4b); 1120, 1129, 

1131 (1-11); 1132 (1-32); 1133 (1-4b); 1143 (2-2); 

1153 (1-33); 1222 (1-42); 1223 (2-4); 1241 (1-42); 

1259 (1-19); 1301, (1-42); 1223 (2-4); 1546 (1-44); 

P.17591, P.17592 (1-28), P.18136 (2-1 la); P.18152 

(2-3a); P.18409, P.18453 (1-40); P.18455, P.19841 

(1-42); P.19878 (2-13b), s.n. (1-4b, 1-42, 2-13a). 

Berlin, B., 372 (1-42); 523 (2-4); 746 (2-lOb); 1531 (1- 

32); 1999 (2-4). 

Bernardi, L., 902 (2-1 la); 910 (2-3a); 1724(1-42); 1919 

(2-3d); 1980 (1-42); 1986 (2-3d); 2657 (2-3a); 2731 

(2-13b); 2744 (1-43); 2808 (2-1 la); 3354 (1-42); 5696 

(2-lb); 6348, 7017 (1-42); 7238 (1-3); 7250 (2-23); 

5.16.5. (2-20); s.n. (tree 3/100) (2-25); s.n. (tree 4/8) 

(2-23); s.n. (tree 6/27) (2-13a); s.n. (1-42, 2-lb, 2-5, 

2-13a). 

Bisby, F. et al., P. 18118 (2-3a). 

Billiet, F. et al., 1108 (1-20). 

Black, G. A. (et al.), 46-137 (1-8); 47-1520 (2-13a); 48- 

2949, 48-3017 (1-28). 

Blanchet, J. S., 160, 2327 (1-25); 2361 (2-lOa); s.n. (1- 

25). 

Blanco, C., 810 (2-la); 958 (1-42); 1149 (2-3a).


Blum, J. L., 2039 (2-3e). 

Coelho, D. (et al.), 672 (2-1 la); 1577 (1-4a); 5979 (2- 

Boerboom, J., 9199 (2-10a). 

20); 51350, 52381 (2-23); 52383 (2-13d). 

Bondar, G., 2169 (2-10a); s.n. (2-la). 

Coelho, L. (et al.), 15 (2-17); 332 (1-4c); 339 (2-16); 

Boom, B. et al., 7812 (1-4b); 7870 (1-9); 8102 (2-14). 2265 (2-7); 5223 (2-13a); 5728 (1-31); 5808 (2-20). 

Bowie, J. et al., 57 (1-25). 

Constanino, D., 19689 (1-25). 

Boyan, R., 267 (2-7). 

Contreras, E., 61, 845, 2220, 3559, 4449, 5787, 6231 

Brade, A. C. (et al.), 7887, 7949 (1-25); 18619 (2-la). (1-30); 10026 (2-3d); 10195 (1-30); 10763 (1-42); 

Braga, P. L. S., 3178 (2-4). 

11195 (2-3d). 

Brandbyge, J. et al., 30194 (2-3a); 30252 (2-23); 30258 Cook, O. F. et al., 157 (1-29); s.n. (1-30). 

(1-31); 30327 (2-23); 30421 (1-32). 

Cooper, G. P., 538 (1-42). 

Brenes, A. M., 20542 (1-34). 

Cordeiro, M. R., 140 (2-23); 367 (1-4b). 

Breteler, F. J., 3889 (1-42); 4920 (2-3d). 

Cordoba, J. J., 295 (1-42). 

Brewer-Carias, C., s.n. (1-11). 

Correa, M. D. et al., 1002, 1031A (1-6); 1854 (2-3d). 

Broadway, W. E., 880 (1-4a). 

Costa, M. da, 90 (1-39). 

Bruijn, J. de, 1123 (1-42); 1496 (2-1 lb); 1497 (2-3a); Cowan, R. S. (et al.), 1868, 1880 (1-26); 38774 (1-20). 

1502 (2-llb); 1514, 1516, 1517, 1526 (2-3a); 1546 Cremers, G., 6726, 8271 (2-lOa). 

(2-1 lb); 1547 (2-12a); 1555 (2-1 lb). 

Croat, T. B., 5124,5125,6588 (1-4b); 7633 (2-4); 7768, 

Buchtien, 0., 2050, 2122 (2-3a). 

7839 (1-42); 7859 (1-4b); 8097, 8100, 8648 (2-3e); 

Buck, P., 26425 (1-25). 

9973A, 14212(1-6); 10343, 10830 (2-3e); 11591 (2- 

Bunting, G. S. (et al.), 3959 (1-4b); 6836 (2-3b). 3d); 15064 (1-6); 17784 (1-40); 18016 (2-la); 18643 

Burchell, W. J., 3148 (1-25); 9792 (2-3a and 2-lOa). (1-4b); 18850 (1-28); 18888 (2-15); 19997 (2-3a); 

Burger, W. C. et al., 8137 (2-23); 10040, 10474 (1-42). 20370 (1-4c); 20411 (1-40); 20447 (2-1 a); 20616 

Busey, P., 628 (1-34). 

(2-la); 20662 (2-4); 22635 (1-42); 24663 (1-30); 

BW (=Boschwezen) 732 (1-20); 772 (1-1); 1322 (1-4a); 24831 (2-30); 59353 (2-3a). 

1368 (2-2); 2030 (2-3c); 2070 (1-4a); 2095 (2-17); Croizat, L., 312 (1-42); 343 (1-4b); 541 (1-42); 959 (1- 

2410 (2-lOa); 3255, 3302 (2-17); 3376 (2-3a); 3430 43); 962 (2-1la). 

(1-4a); 4010 (2-17); 4033 (2-3a); 4494 (2-1Oa); 4866 Crueger, H., s.n. (1-42). 

(2-3c); 5036 (2-3a); 5048 (2-10a); 5262 (1-4a); 5390 Cruz, J. S. de la, 1175 (2-la); 1367 (1-26); 1501 (1- 

(2-17); 5440, 5581, 5992 (2-10a); 6292 (2-3c); 6509 26a); 1577 (1-26); 1657 (2-la); 1729 (1-4b); 2241, 

(2-17). 

2387, 2636, 2813, 3017 (1-26). 

Cabrera, J., 2668 (1-35). 

Cuatrecasas, J. (et al.), 6737 (2-4); 7299 (2-23); 7510 

Callejas, R. et al., 3997 (1-9). 

(1-42); 7602 (2-4); 8291 (1-42); 10714 (1-27); 14284 

Calzada, J. I., 325 (1-36). 

(1-10); 14881 (2-12b); 15071, 15520, 15534 (2-3d); 

Camargo, -, 1849 (1-25). 

15589 (1-42); 15720 (2-18); 15876, 16315, 16755 

Camp, W. H., E3626 (1-42); E3674 (1-19). 

(2-3e); 16826 (2-12b); 17251 (1-34); 17357 (2-3c); 

Campbell, D. G. et al., 6820 (2-unnamed); 6834 (2-6), 17451, 17479 (2-3e); 17647 (1-10); 21256 (1-41); 

6927 (2-5); 7001 (2-la); 7157 (2-3a); 7264 (2-23); 21446 (1-7); 21496 (1-10); 22048 (1-41); 22526 (1- 

7600 (2-3a); 7689 (2-unnamed); 8065, 8127, 8267 42). 

(2-la); 8344 (2-23); 8383 (2-lOb); 8463 (2-23); 8537 Curran, H. M., 8 (1-13); 19 (2-la); 21 (2-1Oa); 116 (2- 

(2-la); 8844 (2-4); 8997 (2-3a); 9120 (2-lOb); 9521 12a). 

(2-3a); 9610 (2-la). 

DAF, 004 (1-13); 027 (2-lOa). 

Campos Porto, P., 863 (1-25). 

Daly, D. C. et al., 1208, 1474 (2-3c); 3790 (2-17); 3936 

Cantonnement de Cayenne, s.n. (2-1 la). 

(2-2); 4129, 4135 (2-7); 4457 (2-16); 4456, 4485 (2- 

Carauta, J. P. P. (et al.), 178, 1714, 1716, 1780, 1781, 8), 5356 (1-11). 

2722; 3011 (1-4a); 3050, 3166 (1-25). 

Damazio, L. B., s.n. (1-25). 

Cardenas, M., 1990 (2-1a); 3973 (2-13c). 

Damiao, C., 2503, 2817 (1-28); 680 (2-20), see also 

Cardona, F., 374 (1-4b); 1222 (2-13b); 2170 (2-5). Mota, C. Damiao A. de. 

Casaretto, J., 617 (1-13); 643 (1-25). 

Damiao, J. P. L. et al., 1805 (2-3a); 1822 (2-13b). 

Castillo S., M., 11 (2-la). 

Daniel, H., 1793, 2652 (1-42). 

Cavalcante, P., 606 (2-23); 777 (2-3a); 2297 

Danta, M., 12383 (1-31). 

(2-3c). 

Cazolet, P. C. D. et al., 5037, 5072 D'Arcy, W. G., 12329 (1-6). 

(2-3d). 

Cer6n M., C. E. 

Davidse, G. et 

(et al.), 252 

al., 15365 

(2-4); 261 

(2-2). 

(2-la); 2019 (2- 

24); 2955 

Davidson, C., 5203 

(1-14); 2993 (1-9); 3389 

(2-9). 

(1-32); 3495 (1- Dayton, W. A., 3019 (1-42). 

22). 

Denslow, J., 2698 (2-3e). 

Chagas, J., 962 (2-4); 1364 (1-4a); 1564 (2-7); 1655 (2- Deward, G., 27 (2-17). 

2). 

Diaz, C. et al., 618 (1-32). 

Cid (Ferreira), C. A. et al., 435, 511 (1-40); 721 (2-23); Diaz, M., 22A (2-20). 

847 (2-13d); 849 (2-2); 942 (1-20); 1053, 2223 (1- Dodson, C. H. (et al.), 3017 (1-39); 5241 (1-42); 5776 

1); 2870 (2-la); 3297 (2-20); 6200 (2-2); 7031 (2- (1-19); 5778 (1-42); 6082 (1-19); 6156, 6312, 6327 

30); 7831 (1-4a). 

(2-3d); 6513, 7416, 7616 (1-41); 9539 (1-19); 15201 

Claes, F., 26 (2-4). 

(1-34). 

Clewell, A. et al., 4142 (2-3d). 

Doeve, -, 3003 (1-20).

Exsiccatae 

Dombey, J., s.n. (1-42). 

Focke, H. C., 9, 418, s.n. (1-20). 

Donnell Smith, J., 4826, 6770 (1-42). 

Foldats, E., 114-1A (2-2). 

Donselaar, J. van, 1226 (2-13e); 1262 (2-3c); 1373, Folsom, J. P. et al., 6377 (2-3d); 6469 (2-23). 

1681 (2-lOa); 1888 (2-17); 2569 (1-1); 2924 (2-lOa); Fonnegra, R. et al., 1798 (1-35). 

3333 (2-3c); 3788 (2-lOa). 

Forero, E. et al., 3145 (2-23); 4099, 4110(1-10); 6367 

Dressier, R. L., 3454 (1-6); 3579 (2-3d); 4636 (1-15). (2-23), P.6419 (2-4). 

Duarte, A. P. (et al.), 3410, 4650, 5328 (1-25); 6970 Forest Department British Guyana, see FD. 

(1-4a); 7194 (2-23); 9785 (2-1 la); s.n. (1-13). Foresta, H. de., 526 (2-13e); 676, 677, 4970, 5011 (2- 

Ducke, A. (et al.), 104 (1-25); 1149 (2-7); 1354 (1-38); la). 

1527 (2-16); 1528 (1-20); 1764 (2-7); 1795 (1-28); Foster, R. B. (et al.), 584 (2-3e); 631 (1-42); 1101 (1- 

1917 (2-8); 1999 (1-38); 2103 (1-4a); 6782, 7715 (1- 4b); 3412 (1-32); 5019 (2-23); 5641 (1-24); 5719 (2- 

28); 9096 (1-38); 10736 (2-17); 12238 (1-28); 12366 10b); 5721 (1-42); 5903 (2-4); 6568 (2-la); 6569 (2- 

(2-4); 13056 (2-17); 13057 (2-3a); 13058 (2-10a); 10b); 7001 (2-4); 7194 (2-10b); 7199 (2-23); 7200, 

13065 (1-4b); 13066 (1-28); 14499 (2-7); 15255 (2- 7201 (2-la); 7995 (2-lOb); 8023, 9373 (2-la). 

17); 15261 (2-lOa); 15350(2-2); 15804(1-4a); 15935 Francisco, 2186 (2-7). 

(2-2); 16003 (1-28); 16321 (1-25); 17132 (1-4b); Franco, J. et al., 419 (1-31). 

17164 (2-7); 17167 (2-2); 18455 (1-40); 18456 (1- Frazao, A., s.n. (1-25). 

20); 18460 (1-4c); 18461 (1-1); 18462 (1-21); 19455 

Froes, R. de Lemos (et al.), 1907 

(2-1Oa); 21200 (1-25); 23606, 23607 (2-7); 23608 

(1-20); 20870 (2-8); 

(2- 21048 (2-4); 21264 (1-39); 23693 

20); 23609 (1-38); 25243 (2-17); 25244 (2-16); 25245 

(2-16); 23725 (2lla); 

25531 (1-28); 26520 

(2-13b); 25246 (2-21); 25247 (2-2); 25248 

(2-13d); 26631 (2-3c); 

(1-38); 28585 

s.n. 

(1-4b); 34651 (1-20). 

(2-13b). 

Duke, J. A. (et al.), 6588, 8035 (2-3d); 11273 

Gandoger, M., 19 (2-2). 

(1-42); 

11599 (2-3d); 14712 (1-6); 15742 Garcia-Barriga, H., 12408 (1-4b); 13643 

(1-4b). 

(2-1 la); 13836 

Dunlap, V. C., 299 (2-23); 13871 (2-4); 14002, 14106 

(1-42). 

(2-5); 14116 (2- 

1 

Duque-Jaramillo, J. M., 1767 (1-4); 2179 (1-8); 2207 

la); 16239 (1-4b). 

(1-28); 2261, 2446 (2-4); 3957 Garcia, J. H. et 

(1-42). 

al., 9 (1-3a). 

Dus6n, P., 2160, 6635, 10134, 10186, 11419 (1-25); 

Gardner, G., 732, 5632, 5858 (1-25). 

11942 (2-la); 13653, 14718, 17030 (1-25); 17239, Gamier, Bro. A., 112 (2-23). 

17345 (2-la); s.n. (1-25, 2-la). 

Garwood, N. C., 759 (2-3d). 

Dwyer, J. D., 4734 (2-3d); 10305 (1-42); 10799 (1-30). Gasche, J. et al., 29 (1-31). 

Dyer, F. J., A-85 (1-42). 

Gaudichaud, C., 992 (1-25). 

Edwards, J. B., 400 (1-42). 

Geay, M. F., 3285 (1-1). 

Eggers, H. F. A. von, 14165, 15615 (1-42). 

Gentle, P. H., 2602 (1-30); 2787, 2917, 3265, 4211 (2- 

Egler, F. E., 42-229 (1-30). 

3d); 4528 (1-30); 6312, 6997, 8675 (2-3d). 

Egler, W. A. et al., 46027 (1-28). 

Gentry, A. H. (et al.), 918 (1-25); 2046, 2657 (2-1 lb); 

Eiten, G. et al., 6214 (1-25). 

6314 (1-42); 6679 (2-3e); 7530 (1-42); 7727, 8348 

Ellenberg, H., 2288 (2-4); 2297 (2-lOb); 2474 (2-4). (1-30); 8707 (1-42); 9552 (2-3d); 9606 (1-19); 9824 

Elias, Bro., 130, 631 (1-42). 

(2-la); 9892 (1-19) 9934, 10127, 12474 (1-42); 13408 

Emden, W. C. van, L-VI, s.n. (1-20). 

(1-15); 14011 (see 1-32); 15621 (2-lOb); 15876 (2- 

Emmerich, M. (et al.), 3003, 3556 (1-25). 

7); 16041 (2-3b); 16181 (2-23); 17217, 17493 (1-4b); 

Emygdio de Mello Filho, L. (et al.), 412, 2340, 2447, 17804 (2-18); 18458 (1-42); 18528 (1-31); 19048 (2- 

3018 (1-25). 

unnamed); 20381 (1-28); 20356 (2-3a); 20495 (1- 

Englesing, F. C., 50, 52a (2-3d). 

28); 20515 (2-5); 20766 (1-40); 20802 (1-28); 21167 

Erlanson, C. 0., 240 (1-42). 

(1-35); 21193 (2-3a); 21205 (2-la); 21291 (1-42); 

Ernst, W. R., 324 (1-42). 

21319 (1-4b); 21346 (2-25); 22095 (1-42); 22280 (1- 

Escobar, L. Albert de et al., 4219 (1-45). 

4b); 22381 (2-spec.); 22460 (2-4); 22865 (2-13a); 

Espina, J. et al., 219 (1-31). 

22866 (2-unnamed); 23713 (1-42); 23835 (2-3e); 

Espinal, T. S., 822 (1-4b); 1314 (1-42). 

24074 (1-4b); 24900, 25050, 25078, 25100 (2-5); 

Falcao, M., 215 (2-4); 1105 (2-14). 

25124 (1-28); 25151 (1-32); 25527 (2-23); 25854 (2- 

Fanshawe, D. B., 1105 (2-14); 2050 (2-la); 3031 (1- 5); 26049, 26117 (2-23); 26525 (1-10); 26530 (2-3d); 

26); 4970, 5011 (2-1a). 

27194 (2-23); 27566 (1-22); 27799 (2-spec.); 28609 

Farifias, J. et al., 347 (2-4); 490 (2-1 la). 

(2-3d); 30278 (2-3e); 31179 (2-10b); 31197 (2-3b); 

FD (=Forest Department British Guyana), 1011 (2- 31271 (2-23); 31655 (2-5); 31816, 31933 (2-la); 

la); 3031 (1-26); 3851 (2-14); 4014, 4073 (1-4b); 31938, 31976, 36231 (2-23); 36241 (2-4); 36309 (2- 

4090, 4970, 5011, 5936, 6914 (2-la); 6973 (1-4b). 23); 36797 (2-18); 39264 (2-13b); 39379 (2-20); 

Fendler, A., 1237, 1237a (1-42); 2420 (2-lb). 

40479 (2-3e); 41890 (2-6); 41894 (2-13); 42029 (2- 

Fernandez P., A. (et al.), F3, 1458 (1-42); 2019 (2-23). la); 42168 (2-3d); 42232 (2-1 Ob); 42568 (2-23); 42793 

Ferreira, C. A. Cid, see Cid (Ferreira), C. A. 

(2-lOb); 43137 (2-4); 43454 (2-23); 45160 (2-3d); 

Ferreira, V. F., 493 (1-25); s.n. (2-20). 

45660 (2-4); 45915 (2-la); 45922 (2-5); 46154 (1- 

Ferreyra, R., 4813 (1-32). 

40); 46752 (2-3c); 47838 (1-46); 48303 (2-3e); 49026 

Ferry, J. F., s.n. (1-42). 

(2-lOa); 49127 (2-1 la); 49346 (2-13); 51089, 51111 

Feuillet, C. P., 1070 (2-23). 

(2-23); 51154 (2-la); 51177 (2-5); 51207 (2-23); 

199


51355 (2-19); 51515 (2-23); 53622 (2-3e); 53829 (1- 

10); s.n. (2-5). 

Gevieski, A., 85 (1-25). 

Gillis, W. T. et al., 10191 (1-42). 

Glaziou, A. F. M., 81, 1013, 1138, 1942, 2016, 4937, 

6009 (1-25); 8934, 8934a (1-13); 8935 (2-la); 8936 

(1-25); 10070a (1-31); 10070 (2-15); 12166 (1-25); 

12173 (2-la); 16350, s.n. (1-25); 20408 (2-la). 

Gleason, H. A., 174 (2-la); 405, 415 (1-26); 670 (2- 

10a). 

Goeldi, A., 7731 (2-10a); 7732 (2-1 la); 7773 (2-lOa). 

Goes, O. C. et al., 815 (1-25). 

Gomes, M. (et al.), 326 (2-la); 547, 593 (2-23); 614 

(2-5); 632, 635 (2-23); 636 (2-13); 646, 655, 673 (2- 

23); 676 (2-3a); 690, 759 (2-23); 786 (2-3a); 789, 

810, 813 (2-23); 826 (2-13b); 838 (2-23); 843 (2-3a); 

844 (2-5); 872, 891, 924, 926, 931, 933, 955; 966, 

979 (2-3a); 1021 (2-2); 1033 (2-23); 1040 (2-5); 1048 

(2-23); 1052, 1072, 1079, 1088 (2-3a); 1118, 1120 

(2-23); 1123 (2-3a); 1129, 1156, 1175; 1176 (2-3a); 

1196 (2-2); 1255, 1319, 1336, 1338, 1356 (2-23); 

1357 (2-2); 1378, 1419, 1423, 1425, 1444, 1446 (2- 

23); 1517 (2-13b); 5134 (2-23); 1540 (2-13b); 1534, 

1568, 1570, 1577, 1584 (2-23); 1588 (2-13b); 1593 

(2-25); 1594, 1595(2-23); 1598, 1604, 1610(2-13b); 

1636, 1640, 1641 (2-23); 1679 (2-13b); 1692, 1731, 

1745, 1769, 1822, 1823, 1867, 1893, 1917, 1983, 

2170 (2-23); 2399 (2-3a). 

Gomes, V. et al., 2818 (1-13); 2819 (1-17). 

Gomez Pompa, A. et al., 3385 (2-3d). 

Gonzalez, A. et al., 1319 (2-1b). 

Graham, Mrs., s.n. (1-25). 

Graimleux, -, 7938 (1-1); 7942 (1-4a). 

Granville, J.-J. de (et al.), 581 (2-23); 2744 (2-19); 4531 

(2-13e); B.5071 (2-la); B.5191 (1-20); 5284 (2-la); 

5642 (2-23). 

Grenand, P., 445 (2-la); 688 (2-13e); 966 (1-4a); 1352 

(2-la); 1442 (2-3c); 1478 (1-20); 1596 (1-4a). 

Grubb, P. J. et al., 1545 (2-24); 1681 (2-13a). 

Grubbe, J. P. et al., 1205, 1283 (1-14); 1498 (1-4b). 

Guanchez, F., 239 (2-4). 

Guedes, M. or T., 1235 (2-2). 

Guillemin, J.-B. A., 1024 (2-la); 1339 (1-13). 

Guppy, N., 656 (2-lOa). 

Gutierrez, G. et al., 562 (1-31). 

Haber, W. A., 1545 (1-34). 

Hage, J. L., 31 (1-25). 

Hahn, W., 347 (1-11). 

Halle, F., 1132 (2-la). 

Hamilton, B. et al., 1062 (1-18). 

Hammel, B., 2176, 2576 (1-34); 4475, 6006 (1-6). 

Hans, D., 335 (1-25). 

Harling, G et al., 14733 (1-4b). 

Haroldo, 57512 (2-13a). 

Hartman, R. L., 12176 (2-3d). 

Hartshorn, G. S., 1224, 1295 (2-23); 1348 (2-3d); 1431 

(2-23); 1436 (1-29); 1539 (2-3d); 2157 (1-34). 

Hatch, W. R. et al., s.n. (1-30). 

Hatschbach, G., 1475, 2458 (1-25); 7411, 7466 (2-la); 

7807, 7833, 16304, 18680, 20253 (1-25); 25765, 

29129 (2-la); 29629, 41730 (1-25). 

Hayes, S., 32, 348 (2-3e); 354 (1-4b); 414, 867, 986, 

1008, s.n. (1-42). 

Hazlett, D., 664, s.n. (1-42). 

Hernandez, J. J. et al., 369 (1-9). 

Herrera Ch., G., 501, 503 (1-34); 505 (1-29). 

Heyde, N. M. et al., 32, 39 (2-23); 78 (1-20). 

Hilty, S., M-86 (2-23); M-87 (1-34). 

Hine, R., P-1609 (2-23). 

Hinton, G. B., 14018 (1-36). 

Hohenkerk, L. S., 738 (1-4b). 

Hoehne, F. C., 30923 (1-25). 

Holdridge, L. R., 2514 (2-3d); 6254a (2-3e); 6818 (2- 

23). 

Holm-Nielsen, L. et al., 24519 (1-14); 24856 (1-10); 

26331, 26610 (1-42). 

Hoist, B. K. et al., 2421 (2-1 la); 2719 (2-la); 3195 (2- 

3a). 

Hostmann, F. W. R., 1189 (1-4a). 

Hostmann, F. W. (& A. Kappler), 1272 (2-lOa and 

2-1 la). 

Howard, R. A. et al., 605 (1-42). 

Hoyos, J., 2043 (1-42). 

Huashikat, V., 431 (2-1 la); 912 (2-3b); 944 (2-lOb); 

966 (2-13a); 1027 (2-10b); 1150(2-23); 1216 (2-la); 

1241 (2-3a); 1432, 1482 (2-13a); 1585 (see 1-12); 

1872 (2-3b). 

Huber, J. E., 238 (2-la); 1640 (2-2), 1775 (1-21), 1873 

(1-4a), 4244 (2-la). 

Humbert, H. et al., 27221 (2-4). 

Idrobo, J. M. et al., 786 (2-3a); 1316 (2-lOb). 

Ijjasz-Madriz, -, 28 (2-1b). 

Irwin, H. S. (et al.), 183 (1-26); 2058 (1-17); 4783 (2- 

lla); 47874 (2-23); 48192 (2-3a); 48439 (2-1 la); 

48468 (2-23); 48631 (1-16); 48682, 48755 (2-3c); 

55487 (1-20); 55576, 55884 (2-17); 57574 (1-1). 

Izawa, K., 9 (2-la); 21 (2-4). 

Janse, C. 0., 276 (2-3d). 

Jaramillo, J. et al., 2088 (1-40); 4586, 31516 (2-4). 

Jativa, C. (etal.), 303, 1079 (2-12b); 1097 (1-10); 2033, 

2037 (2-12b). 

Jenman, G. S., 652, 3995, 4947 (1-26); 4947A (1-4b); 

5315 (1-26); 6310 (2-la); 7310, 7933 (1-26). 

Jimenez, M. A., 433 (1-35); 657, 1750, 2887 (1-42); 

3016 (1-18); 3816 (2-3d). 

Jim6nez Saa, H., 15631 (=LBB 14296) (2-17). 

Jobert, -, 682 (2-15). 

Johnston, I. M., 580, 606, 608, 652, 678 (1-42); 1709 

(1-4b); 1714 (2-3e), 1725, 1768 (1-42). 

Jones, G. C. et al., 3152 (1-30). 

Jones, J. et al., 9712, 9769 (1-28); 9788 (1-40). 

Jonsson, G., 5A, 610a (1-25). 

Juncosa, A., 1253 (2-12b). 

Kanehira, R., 99 (1-42). 

Kappler, A., 1272 (2-11 a). 

Karsten, H., s.n. (1-42, 2-la, 2-lb, 2-3a, 2-4). 

Kayap, R., 201 (2-13a); 268, 393 (2-la); 571 (2-1 la); 

1057 (2-la); 1306 (2-10b). 

Kenoyer, L. A., 312 (2-3e). 

Killip, E. P. (etal.), 25341, 26095 (1-32); 27381,27932 

(2-4), 28800 (2-unnamed); 29246 (1-42); 29839 (2- 

4); 29955 (2-23); 30139 (2-20); 30179 (1-38); 35095 

(2-3e). 

Kinloch, J. B., s.n. (1-30). 

Klein, R. M., 89 (2-la); 79B, 772 (1-25); 1110 (2-la). 

Klug,G., 1185, 1326 (2-4).


Korning, J. et al., 47618 (2-13c), 47641, 47645 (2-3a); Lindeman, J. C. (et al.), 1857 (1-25); 3631, 3689 (2- 

47801, 47807 (2-unnamed). 

10a); 4057 (2-2); 5468 (2-3c); 5682 (1-25); 5709 (1- 

Kosei Izawa (see also Izawa, K), 9 (2-la); 21 (2-4). 4a); 5748 (1-25); 6123,6166 (2-3c); 6362 (1-1); 6418 

Krieger, P(adre) L. (as PLK) (et al.), 12268 (2-4); 12296 (2-2); 6445, 6600 (1-1); 6765 (2-3c); 6923 (2-lOa); 

(1-42). 

6938 (1-20); 9105, 9110 (1-25). 

Krukoff, B. A., 1147, 1187 (1-39); 1197 (2-17); 1297 Lindeman, J. C. & A. C. de Roon, 747a (2-3c); 828a, 

(2-la); 4518 (1-42); 4817 (2-la); 4994 (1-42); 5109 842b (2-3a). 

(2-4); 5282 (2-23); 5309 (2-lOb); 5327, 5332 (2-4); Lindeman, J. C. & A. L. Stoffers et al., 33 (1-29); 34 

5596 (1-28); 5657 (1-32); 6041 (2-23); 6165 (1-42); (1-4a); 73 (1-20); 540 (2-17); 701 (2-13e); 749 (1- 

6238 (2-la); 6524 (1-42); 6662 (l-4b); 6887 (2-13b); 20); 790 (2-17); 799 (1-20). 

7071 (2-23); 7073 (2-20); 7966 (1-4a); 8073 (2-23); Lisb6a, B., s.n. (1-13, 1-25). 

8223 (2-13b); 8273 (1-35); 8325 (2-13c); 8332 (2- Lisb6a, P. L., 27 (2-4). 

4); 8369 (2-5); 8223 (2-13b); 8273 (1-35); 8325 (2- Little, E. L. (et al.), 43 (1-27); 212, 213 (1-14); 424 (1- 

13c); 8332 (2-4); 8369 (2-la); 8373 (2-6); 8386 (2- 32); 602 (1-42); 635 (1-11); 6155 (1-42); 6241, 6258, 

la); 8388 (2-21); 8401 (1-28); 8406 (1-4c); 8427 6301 (2-3d); 6328, 6461 (1-42); 6640 (2-3d); 7756 

(2-15); 8469 (2-4); 8518 (1-31); 8539 (1-35); 8587A (2-4); 8423 (2-la); 9527 (2-6); 9629 (1-40); 9641, 

(1-31); 8652 (2-la); 8658 (1-35); 8695 (2-20); 8755 9789 (1-42); 15990 (1-4b); 16230, 16245, 21047 (1- 

(1-35); 8807 (2-16); 8897 (1-35); 8967 (1-31); 8994 42); 21056 (1-41); 21070 (2-12b); 21136 (1-34); 

(1-39); 10123 (1-4b); 10861 (2-13c); 10867, 10874 21172 (1-42); 21225 (2-12b). 

(2-3a); 11062 (2-13c); 11085 (1-11); 11254 (2-3a); Lizot, J., 83 (1-40); 1972-1 (1-4b); s.n. ("Cl") (2-la). 

11275 (1-9). 

Lleras, E. et al., P.17178, P.17197 (1-39); P.17242 (2- 

Kuhlmann, J. G. (et al.), 255 (2-13b); 263 (1-20); 291 9); P.17411 (2-8); P.17484 (1-4a); P.19586 (2-20); 

(2-13b); 368 (1-42); 382 (2-2); 446 (2-10a); 470 (1- P. 19661 (1-20). 

37); 759 (1-42); 1178 (1-28); 1337 (2-4); 1540 (1- Loureiro, A. (et al.), 16461 (2-13a); 16568, 16968, 

32); 1832, 2073, 2074 (1-17); 2075 (2-la); 2177 (1- 50623 (2-20). 

17); 2200 (2-la); 2207 (1-17); 5004 (1-25); 19843, Lowrie, S. R. et al., 710 (2-23). 

150082 (2-7); s.n. (1-13). 

Lozano, C. G., 640 (1-42). 

Kuhlmann, M. et al., 52 (2-2). 

Luetzelburg, Ph. von, 12471 (1-25); 23887, 23913 (2- 

Lambroy, -, s.n. (1-4b). 

4). 

Lanjouw, J. et et al., 399 (2-1 la); 432 (2-13e); 1761 Lugo, M., 83 (1-4b); 2820, 2857, 2909, 3563 (1-27). 

(2-2); 2106 (2-10a); 2113 (2-3c); 2114 (2-23). Lund, P. W., 139 (1-25). 

Lanna Sobrinho J. de Paula et al., 376, 1011 (2-la). Lundell, C. L., 2, 648, 3170, 3535, 6224, 6350, 15930, 

Lao, E. A. et al., 21 (1-42). 

16122, 16162 (1-30). 

Lao, M. R. et al., s.n. (2-4). 

Luschnath, B., s.n. (1-25); 40, s.n. (2-lOa). 

Lasser, T., 2202, 2267 (2-lb). 

Maas, P. J. M. et al., 2478 (1-26); 2592 (2-14); 3325 

Lawrance, A. E., 727 (2-3a); 769 (1-42); 810 (1-4b). (1-25); 3622 (1-26), 3938 (2-1Oa); 4126 (1-26); 4679, 

LBB (='s Lands Bosbeheer Suriname), 8000 (2-17); 4740, 4746, (1-19); 5858 (2-la); 6201, 6225 (2-4); 

8161 (2-23); 8316 (2-3c); 8976 (2-1 la); 9199, 9424, 6249 (2-7); 6267 (2-25); 11017 (2-17); 11027 (2-23). 

9843 (2-10a), 11017(2-17); 11027 (2-23); 11747 (1- Macbride, J. F., 5446 (2-4); 5447 (1-4b); 5594 (1-32). 

20); 14296 (2-17); 16031, 16310 (1-4a). 

Macedo, R., 55753 (2-17). 

Leeuwenberg, A. J. M., 11784 (2-3a). 

Madison, M. T. et al., 4764, 7101 (1-19). 

Lehmann, F. C., BT-694 (1-10); 948 (1-41); 5606 (1- Magalhaes, G. Mendes, 733 (2-la). 

42). 

Magnago, 

Leitao Filho, H. P., 685, 686 (1-25). 

Leite, J. E., 413, 892 (1-25). 

Leman, D. S. (herb.), s.n. (1-4a, 1-20). 

Lemee, A. M. V., s.n. (1-4a); s.n. (2-3a, 2-3c). 

Lems, K., s.n. (1-25). 

Leng, H., 177 (1-26). 

Lent, R. W. (et al.), 349 (1-42); 2004 (2-3d); 2327 (2- 

23); 2520, 2535 (1-42); 2538 (1-34); 3279, 3388 (1- 

42); 3547 (2-3d); 5320 (1-42). 

Le6n, J., 3501 (2-3d). 

Leprieur, F. R., 141 (2-lOa); s.n. (1-4a, 1-16, 1-21, 

2-17). 

Lescure, J. P., 343 (2-1 la); 422 (2-23); 2075 (2-5). 

Liesner, R. L. (et al.), 417A (1-34); 515, 1309 (1-6), 

4105 (2-13a); 8750 (1-31); 9443 (1-42); 10944 (2- 

3d); 12394 (2-lb); 16175 (2-13b); 16299 (2-3a); 

22180 (1-11); 24462 (2-la). 

Lima, A., see Lima, D. de Andrade 

Lima, D. de Andrade (et al.), 20795 (1-25); 58-3185 

(2-2); 67-4983 (2-la). 

Lima, R., 21 (2-lOa). 

- et al., 58046 (2-la). 

Maguire, B. (et al.), 4945A (2-1 la); 23449 (1-26); 23867, 

23942 (1-1); 24588 (1-26); 29408 (1-35); 41620 (1- 

40); 46879 (1-3); 47049 (1-1); 53517 (1-4a); 56511 

(2-1 la); 60333A (2-3a). 

Maia, L. A. et al., 497 (2-4). 

Malme, G. O. A., 826, 1428 (1-25). 

Marcano-Berti, L. (et al.), 471, 611 (2-la); 238-979 (2- 

3d); 84-2-77 (1-4b); 982-154 (2-3d). 

Marinho, L. R., 273 (2-la); 348 (1-4b). 

Martin, J., s.n. (1-20, 2-la, 2-3c, 2-1 la). 

Martinelli, G., 150 (1-25). 

Martius, K. F. P. von, 2620 (2-2); 2673 (1-28); s.n. (1- 

28, 1-42, 2-la, 2-2, 2-3a, 2-4, 2-7, 2-13a, 2-15). 

Mathews, S., 2060 (1-42). 

Matuda, E., 446, 2020 (1-36); 3300 (1-30). 

McDaniel, S. (et al.), 2666 (1-31); 16092 (1-4b); 16344 

(1-40); 20362 (2-15); 20403 (2-2); 20438 (2-7); 20555 

(1-40). 

McDonagh, J. F. et al., 114 (1-15). 

McPherson, G., 7594 (1-6); 11510 (2-1 lb). 

Medina, E., 379 (1-40).


Melinon, E., 18 (1-20); 172 (1-11); 457 (2-11a); s.n. 

(1-20, 2-1 la). 

Mello Barreto, H., 1795 (1-13). 

Mello, F., 55.388 (2-13a); 55.444 (2-1 la); 57509 (2- 

la). 

Mendez, R., 15 (1-6). 

Meneces, E. (et al.), 338 (2-4); 613, 647, 725 (2-23); 

759 (2-la). 

Mennega, A. M. W., 271 (1-6); 521 (1-20). 

Metcalf, R. D. et al., 30093 (1-42). 

Mexia, Y., 1872 (1-36); 5115 (1-25); 6573 (1-42). 

Meyer Drees, E., 5595 (2-3a). 

Meyer, D. G., 403 (2-23). 

Miers, J., 2714, 3792, 3858 (1-25); 6201 (2-la); 6257 

(2-4); 8172 (2-la); s.n. (1-25). 

Miranda Bastos, A., 2051 (2-la); 2220 (2-23). 

Miranda, F., 1737, 6164, 6273, 7180 (1-36). 

Molina R., A. et al., 1963 (2-3d); 17694 (1-34); 18227 

(2-3d); 18289, 30747 (1-42). 

Monsalve B., M., 1176 (1-46); 1259 (1-4b). 

Montaldo, P., 3435 (1-42). 

Monteiro da Costa, R. C., 90 (1-42); 92 (2-17). 

Monteiro, O. P. (et al.), 309 (2-20); 477 (2-la); 1119 

(2-23); 1295 (2-la). 

Mora, L. E., 2295 (2-3e). 

Morales, -, 38 (1-41). 

Moreira, A. S., 46 (1-25). 

Moreno, P. P., 23629 (1-18). 

Moretti, C., 102 (2-la); 836 (1-20); 837 (1-4a). 

Mori, S. A. (et al.), 5360 (1-34); 8583 (2-17); 8662 (2- 

la); 8864 (1-26); 10408 (2-4); 11025 (2-lOa); 11026 

(2-2); 11046 (2-la); 11655 (1-33); 14940 (2-19); 

15127, 15342 (2-13e); 15350 (2-3c); 15776 (1-40); 

16046 (2-la); 17185 (2-19); 17478 (2-1la); 17672 

(2-13e). 

Morillo, G. et al., 7384 (2-4). 

Mosen, H., 2614, 2941 (1-25). 

Mota, C. Domiao A. de (et al.), 154 (1-39); 680 (2-20); 

740 (2-la); see also Domino, C. 

Mota, M. G. et al., 1040 (2-5). 

Mutis, J. C., 646, 649, 4558 (1-42). 

Nadeaud, J., s.n. (1-25). 

Nascimento, R. J. (et al.), 262 (2-2); 303, 387 (2-la); 

66338 (2-3a); 66396 (2-7). 

Nee, M. (et al.), 8822 (1-34); 9281 (1-6); 30877 (2- 

13a). 

Neill, D. (et al.), 1653, 1919, 1984, (1-42); 3389 (2- 

23); 7044 (2-4); 7082 (2-la); 7179 (1-42); 8158 (2- 

13a); 8289 (1-22). 

Nelson, B., 680, 681 (2-la); 705 (2-4). 

Nevers, G. de (et al.), 4320 (1-6); 4697, 4789 (1-11); 

4929 (2-1 lb); 4982 (1-15); 5062, 5502, 6152 (1-6); 

6186 (1-18); 6194 (1-6); 6997 (2-23); 7170, 7542 (2- 

3d). 

Nishimura, A., 39 (2-1a). 

Nufez, O. V., 2928 (1-42). 

Nuiiez, P., 1823, 5761, 5946 (2-23); 5949 (2-la). 

Occhioni, P. (et al.), 5864, 5917 (1-13); 8228, 35333 

(1-25). 

Oersted, A. S., 14317 (1-42). 

Oldeman, R. A. A. (et al.), 207 (1-1); T.243 (1-4a); 

T.243a (1-21); 366 (1-4a); B.644 (2-10a); B.1 176 (1- 

1); 1179 (2-la); 1243 (2-1 la); B.1324 (1-1); B.1338 

(1-20); 1619 (1-1); B. 1946 (1-21); 2030 (2-3c); 2476 

(1-1); 2504(2-23); B.2538(1-4a); 2821 (2-23); B.2861, 

3062 (1-21); 3115 (2-3c); B.3243 (1-1); B.3420 (1- 

4a); B.3786A (2-2); B.4108, B.4132 (2-la). 

Oldenburger, F. H. F. et al., 1120, 1406 (2-3c). 

Oliva, F., s.n. (1-42). 

Oliveira, E. de, 985 (1-20); 1974 (2-17); 2597 (2-10a); 

2764 (1-35); 3071 (2-1Oa); 3550, 3599 (1-20); 3627 

(1-28); 3682 (1-20); 3736 (2-1 la); 3915 (2-17); 3936 

(2-13e); 3954 (1-20); 4047 (2-1 la); 60534 (2-5). 

011gaard, B. et al., 35068 (1-49); 57061 (2-3a). 

Ortega, -, s.n. (1-34). 

Ortiz, R. T., 200, 1030, 1810 (1-30). 

Palacios, W. (et al.), 1874 (2-24), 2059 (2-3a); 6387 

(1-27). 

Pariona, W. et al., 954 (2-13c). 

Parker, C. S., s.n. (1-4b). 

Peck, M. E., 497 (1-30). 

Pennington, T. D. (et al.), 12244 (2-24), P.22647 (2- 

7). 

Pereira, A. B., 39 (1-25). 

Pereira, L., 5670 (1-25). 

Perrottet, G. S., s.n. (1-4a). 

Persaud, A. C., 116 (1-4b); 222 (1-26). 

Pessoal do Horto Florestal, 650, 651 (2-la); s.n. (1- 

25). 

Peters, C., 187 (2-4). 

Petilon, -, 154 (1-4a). 

Philipson, W. R. et al., 1914 (1-42); 2086 (2-3a); 2095 

(1-42); 2133 (2-lOb); 2134 (2-23). 

Pinheiro, G. S., 1716 (1-25). 

Pinsley, H. V., 405 (2-4); 556 (2-3A). 

Pires, J. Murqa (et al.), 502, (2-39); 507, 542, 587 (2- 

3a); 805, 830 (1-4b); 6663 (2-la); 7015 (2-2); 10123, 

10758, 10794, 10809 (2-lOa); 12570, 15088 (2-la); 

16461 (1-42); 16547 (2-1a); 50419 (2-23); 50631 (1- 

1); 50656 (1-2); 50864 (1-20); 50913 (1-4a); 51193 

(2-13e); 51247 (2-23); 51390 (2-17); 51409 (1-20); 

51624 (2-3c); 51625 (2-13e); 51676 (2-10a); 51736 

(1-40); 51882 (2-2); 52656 (1-20). 

Pires, O. et al., 248 (1-42). 

Pittier, H. (et al.), 2286 (2-3e); 3892 (1-42); 4386 (1- 

6); 8404 (1-42); 11166 (1-34); 12105 (2-3d); 12124, 

12142, 12144, 12288, 13985 (1-42); 15641 (2-1b); 

15733 (1-42); 16163 (2-3d); s.n. (1-42). 

Plowman, T. C. (etal.), 2362 (1-42); 2919 (1-25); 6555 

(1-28); 6903 (2-4); 6906 (2-5); 11179 (1-32); 11180 

(1-42). 

Poeppig, E., 1881 (2-la); s.n. (1-28, 1-42, 2-la, 2-3c). 

Pohl, J. E., s.n. (1-25, 1-28). 

Poiteau, P. A., s.n. (1-20, 2-3c, 2-3a, 2-1 la, 2-3a, 2-3c, 

2-1 la). 

Poveda, L. J., 951 (1-34). 

Prance, G. T. (et al.), 1379 (2-3a); 1548 (2-2); 1720 (1- 

20); 1777 (2-la); 2153 (1-4c); 2180 (2-1 la); 2306 (1- 

42); 2469 (2-2); 2754 (2-3a); 3253 (2-7); 3260, 4438 

(2-la); 4588 (1-4b); 6253 (2-la); 6292 (2-13b); 6419 

(2-4); 6718 (1-4b); 6969 (1-31); 7160 (2-17); 7371 

(1-32); 7503 (2-4); 7530 (1-42); 7670 (2-la); 7792 

(2-23); 7907 (2-lOb), 8378 (2-23); 8670 (2-la); 9054 

(1-35); 9679 (1-11); 9857, 9952 (2-13b); 10089,10124 

(2-23); 10231 (2-5), 10233 (2-la); 10348 (1-42); 

10836 (2-la); 11066, 13438 (1-42); 13579 (2-la); 

13582-A (2-3a); 14029, 14146 (1-31); 14795 (2-7); 

14976 (1-38); 15039 (1-34); 15366 (2-3a); 15393 (2-

Exsiccatae 

20); 15764 (1-40); 15791 (2-4); 17242 (2-9); 17847 Ruiz-Teran, L., 423 (1-42). 

(1-38); 18032 (1-31); 18204 (2-23), 18240 (2-5); Rusby, H. H., 1599 (2-4). 

18250 (2-3a); 19878 (2-13b); 20454 (2-20), 21011 Rylands, A. B., 15 (2-2); 25 (2-23); 30 (2-13b); 52 (2- 

(2-2); 22708 (2-13b); 22720 (1-20); 22729 (1-31); 3a); 59 (2-2). 

22740 (1-20); 22803 (2-la); 22836 (1-20); 22859 (2- Sabatier, D. R. L., 501 (2-1 la). 

16); 22864 (2-20); 22999 (1-35); 23776 (2-7); 23846 Sagot, P. A., 517, 970, 972 (2-17); 990, 990bis (2-1 la); 

(1-39); 23966 (2-9); 24188 (2-3a); 24313 (2-7); 24687 1136 (1-20); 1163 (2-2); s.n. (1-4a, 1-20, 2-17). 

(1-42); 25167, 25336 (2-2); 25337 (2-10a); 25363 (2- Samaniego, A. et al., 92 (1-42). 

2); 25530 (2-20); 25748 (1-35); 26375 (2-la); 26402 Sandwith, N. Y., 290, 302, 394 (1-26). 

(1-42); 58783 (1-20); s.n. (2-5). 

Santos, M. R., 172 (2-23); 455 (2-lOa). 

Pr6vost, M. F. et al., 738 (1-4a); 982, 983 (2-3c). Santos, T. S. dos, 2035 (2-la). 

Proctor, G. R., 27080 (1-18); 27336 (1-42); 30273 (1- Sastre, C., 2274 (2-4); 4350, 4579 (2-3c); 5500 (1-1); 

30). 

6130 (1-20); 6417 (1-4a). 

Pruski, J. et al., 3235 (2-13b); 3239 (2-20). 

Sawada, M., 21 (2-1 Ob). 

Purdie, W., s.n. (1-14). 

Schipp, W. A., 127 (2-3d); 999 (1-30). 

Purpus, C. A., 5996, 11161, 11162, 11182, s.n. (1-36). Schnee, L., 1337 (1-42). 

Rambo, B., 426, 27092, 37782, 38427, 39413, 39780, Schomburgk, R., 118, 167, 287, 876, 1366, s.n. (1-26). 

41173, 44743, 45136, 45904, 46145 (1-25). Schubert, B. G. et al., 748 (2-3d). 

Ramirez, C. R., 1084 (1-31). 

Schultes, R. E. (et al.), 3343, 3826, 3862, 3901 (2-4); 

Ramos, J. et al., 204 (2-20). 

6693 (1-8); 6726 (1-28); 6805, 8175 (2-4); 8269 (1- 

Rauh, W., P-1609 (2-23). 

8); 8380 (1-28); 8976 (1-4b); 9685, 12985 (1-31); 

Reineck, E. M. et al., 446 (1-25). 

13589(1-40); 13774(1-31); 14148(2-23); 15324(1- 

Reitz, R. (et al.), 975, 1688, 1912, 2008, 2027, 2913, 31); 15973(2-5); 16009(2-20); 16112(1-31); 16334, 

3809, 4176, 4692, 4965, 5030, 5611, 8294 (1-25). 16367 (1-42); 17615 (2-3a); 18969, 19758 (1-4b); 

Renteria, E. et al., 45, 2093 (2-12a). 

19835 (2-5); 24547 (2-4). 

Revilla, J., 805, 1099 (1-28); 1260 (1-4b); 1818 (2-5); Schultze-Rhonhof, H., 1953 (1-19); 2783 (1-11); 2943 

1847 (1-31); 1966 (2-7); 2387 (1-39); 3387 (1-28). (1-9); 3006 (1-11); 3020 (1-14). 

Reyno, R. N., 37 (2-13a). 

Schulz, J. P. (etal.), 7787 (1-20); 8000 (2-17); 8161 (2- 

Ribeiro, B. G. S., 612 (1-11); 665 (1-43). 

23); 8316 (2-3c); 8976 (2-1 la); 10344a (1-1). 

Richard L. C. (herb.)., s.n. (1-20, 2-la). 

Schunke, C., 416, A-95 (2-23). 

Riedel, L. (et al.), 1 (1-25); 2 (2-la and 2-lOa); 7 (1- Schunke, J. M., 130 (1-28); 371 (1-40). 

25); 37, 46 (2-la); 75 (1-25); 160 (2-la); 664; s.n. Schunke V., J., 1162 (1-32); 2139 (2-23); 2824 (2-4); 

(1-25, 2-la). 

3309, 3552 (2-la); 6485 (2-4); 7415 (2-la); 7729 (2- 

Riera, B. J. J.-L., 206 (2-3c); 301, 571 (2-17); 573 (2- 10b); 8487 (2-23); 8666 (1-4b); 10650 (2-lOb). 

10a); 714 (2-la); 723 (2-13e); 923 (2-la). 

Schwacke, W., 560 (2-15); 1072, 6783, 8418 (1-25); 

Rimachi, Y. M., 359 (1-32); 1943 (1-28); 2297 (1-40); 10393 (2-la); 11895 (1-17). 

2724 (2-5); 2725 (2-9); 2765 (2-5); 3256 (2-15). SEF (=Studies of Ecuadorian Forest), 8532 (2-6); 8617 

Rizzini, C. T., 415, 1114 (1-25). 

(2-23); 8764 (2-4); 8823 (2-13c); 8930 (2-4); 9033, 

Roa T., A., 251 (2-unnamed); 252 (2-13a). 

9159 (2-23); 10366, 10378 (2-spec.). 

Rodrigues, C. R., 944 (2-5). 

Sehnem, A., 7993 (1-25). 

Rodrigues, G. et al., 877 (1-17). 

Sello(w), F., 241 (1-25); s.n. (2-2). 

Rodrigues, W. A. (et al.), 884 (2-23); 1415 (2-7); 1424 Shakaim, S., RBAE-26 (2-23). 

(2-23); 1434 (2-13b); 2092 (2-2) = 2892 (2-2); 2984 Shank, P. J. et al., 4294 (1-42). 

(2-1Oa); 2986, 2987 (2-13e); 3267 (2-2); 4237 (2-3a); Shattuck, O. E., 260, 522, 525 (2-3e); 832 (1-42). 

4585 (2-2); 5266 (1-31); 5449 (2-20); 7069, 7076 (2- Shepherd, D., 364 (2-23); 376, 379 (2-3a). 

7); 7081,7292 (2-20); 7532 (2-23); 8065 (2-13); 8066 Shima, D., 14 (2-20). 

(2-5); 9204 (2-la); 9206 (2-3a); 9219 (2-la); 10082 Sigueira, R., 8808 (1-20). 

(2-17); 10182 (2-23); IG1-2A-70 (2-5). 

Silva, A. S. Lima de et al., 215 (2-13d); 476 (2-8). 

Rodriguez, H., 399 (1-42). 

Silva, J. F., 69, 211 (1-20); 227 (1-4b). 

Rodriguez, J. V., 2929 (1-42). 

Rohr, J. B. von, 151 

Silva, M., 354 (2-lOa). 

(1-4a). 

Rombouts, H. E., 803 

Silva, M. Barbosa da, 130 (2-lOa). 

(1-4a). 

Romero, F., 548 Silva, M. F. et al., 466 

(1-42). 

(1-31); 742 (2-20); 869 (2-lOa); 

Romero-Castaieda, R., 3647 (2-4); 3840 (2-5); 3848 

903 (2-23); 2166 (1-4a); 2199 (2-13b); 2423 (2-2); 

(2-20); 4211 (1-4b); 4715 (2-12a); 5283 (1-19); 5484 

2591 (1-39); 2623 (2-la); 2712 (1-4a). 

(2-3e); 5512 (1-10). 

Silva, M. G. et al., 3345 (1-40); 3368 (1-4b); 4326, 

Rooden, J. van et al., 258 (1-4b); 699 (1-10); 700 4738 

(1- (2-23); 4750 (2-la). 

5). 

Silva, N. T. (et al.), 921, 949, 995 (2-1 la); 1207, 1321, 

Rosa, N. A. (et al.), 296 (1-11); 339 (1-43); 1091 (2- 

1380 (1-20); 1411, 1412 (2-3c); 3013 (1-28); 4757 

5); 1126 (2-17); 1158 (2-lOa); 1159 (2-5); 1265 (2- (1-10); 60670 (1-43). 

23); 2053, 2874 (1-40). 

Skutch, A. F., 2023 (1-36); 4083 (2-3a); 4225 (2-3d); 

Rowlee, W. W. et al., 856 (1-42). 

4256, 4262, 4267, 4612 (1-42); 4740 (2-20); 57832, 

Ruiz, H. & J. Pavon, 2 (1-32); 3 (1-42); 4 (1-4b); 5, 6, 58862 (2-3a). 

7 (1-32); s.n. (1-4b, 1-32, 1-42, 2-lOb). 

Smith, A., 1632 (1-29); 1663, 1650 (1-34). 

203


Smith, A. C., 2731 (2-13d); 2753, 2799 (1-35); 2845 3d); 122390, 125857 (2-3a); 129163 (2-4); 129733 

(2-1a); 3564 (2-3c). 

(2-unnamed); 131962 (2-1 la). 

Smith, D. N. (et al.), 1090 (1-32); 2089 (2-4); 4743 (2- Stoffers, A. L. et al., 244 (2-17); 265, 311, 312 (2-la); 

unnamed); 5134 (2-10b); 5300 (2-4). 

319 (2-17). 

Smith, L. B. et al., 12307 (1-25). 

Sucre, D., 1783, 3633, 4989, 5209, 5330, 7878 (1-25); 

Smith, R. F., 3221 (2-lb); 3313 (1-42). 

8303 (2-lOa); 8684 (2-2); 8690 (1-25). 

Smith, S. F. et al., 695 (2-3a). 

Sugden, A., 415 (1-42). 

Snedaker, S. C., C-39, D-88 (1-30). 

Sytsma, K. J., 4018 (2-3d); 4024, 4225 (1-6). 

Sneidern, K. von., s.n. (2-4). 

Tamayo, F., 3370 (2-lb). 

Sodiro, A., 193/12 (1-41). 

Teixeira, L. O. A. (et al.), 61 (2-3a); 849 (1-4B). 

Soejarto, D. D., 327, 856 (1-42); 2917 (2-23); 3385 (2- Terceros, W., 1400 (2-la). 

3e); 3489, 4063 (2-23); 4283 (2-1 lb). 

Tessmann, G., 3054 (2-4); 3416 (1-42); 3922 (1-32); 

Solomon, J. C. (et al.), 9254 (2-23); 12655 (2-la). 4236 (2-3b); 4642 (2-la); 4673 (1-39); 4696 (1-42); 

Soria S., M. A., 21 (2-23). 

s.n. (1-25); 5364 (2-9). 

Sparre, B., 14822 (1-42). 

Teunissen, P., 16031 (1-4a). 

Sperling, C. R. et al., 6430 (2-23). 

Thomas, W. W., 3349 (2-13b); 3370 (2-3a). 

Spichiger, G. et al., 1995, 1996, 1997, 1998, 1999 (2- Tillett, S. S. (et al.), 45869 (1-43); 671-33 (1-3 1); 49495 

25). 

(2-1 la). 

Splitgerber, F. L., 448 (1-4a). 

Tomas, Bro., 1923 (1-42). 

Spruce, R., 951, 1219 (2-13a); 1509 (1-4a); 1608 (1- Tonduz, A., 9520 (1-42); 12930 (2-3d); 13280 (1-42). 

42); 2023 (2-4); 2260 (1-4b); 2498 (1-31); 2616 (1- Toro, R. A., 120C (1-14). 

40); 2865 (2-13b); 3176 (2-1 la); 3231, 3464 (1-40); Torres, M. J., 482 (1-32). 

3782 (1-38); s.n. (2-13a, 2-17). 

Toth, M. J., 482 (1-32). 

Stahel, G.et al., 732 (1-20). 

Tresling, J., 55 (1-1). 

Stahel, G., (Exp. Wilhelmina Gebergte) 196 (1-20). Triana, J. J., 860 (2-la); 861 (2-3a); 862 (2-4); 866 (1- 

Stahel, G., (Woodherb. Suriname) 82 (2-23); 123A (2- 34); 867 (1-42); 1846 (2-4); 1866 (1-34, 1-42); s.n. 

3c); 123B (2-17); 166 (2-3a); 166A (2-17); 166B (2- (1-42). 

3c). 

Troon, F. van et al., 16310 (2-23); s.n. (2-la). 

Standen, -, 20 (1-42). 

Trucios, T., 9 (2-1Ob). 

Standley, P. C. (et al.), 7081 (1-42); 7878 (2-3d); 8861, Trujillo, B., 6141, 8765 (1-42). 

12288 (1-42); 19904 (2-3d); 23766 (1-42); 27479, Tuerkheim, H. von, 4082 (2-3d); 8659 (1-30). 

27502 (2-3e); 30924 (1-42); 37121 (2-23); 37259 (1- Tunqui, S., 7, 94 (2-3b); 103 (1-42); 149 (2-3a); 217 

42); 39792 (1-34); 41170, 44550, 44968, 45182, (2-13a); 296 (2-10b). 

45661, 47791, 48594 (1-42); 48600 (2-23); 52900 Tweedie, J., 29 (1-25). 

(2-3d); 53230 (1-42); 53951 (2-3d); 53991, 53425, Tyson, E. L. et al., 4734 (2-3d). 

55155, 55445, 56623, 72288 (1-42); 72667 (2-3d). Ule, E. H. G., 991 (2-la); 1693 (1-25); 5265, 5266 (2- 

Steinbach, J., 1484 (1-32); 7567 (1-42). 

4); 5717 (1-42); 5718 (2-23); 5719 (2-6); 8836 (1- 

Stergios, B. et al., 11530 (1-35). 

4a); 8839 (2-17); 9314 (2-4); 9315, 9316 (1-32); s.n. 

Ster, W. L. et al., 423 (2-3d). 

(1-25). 

Stevens, W. D., 4839, 8323 (2-3d); 13340 (2-23); 13341 Uribe-Uribe, L., 364 (1-42). 

(2-3d). 

Vasquez A., R. et al., 17 (2-23); 167 (2-25); 809 (2- 

Steward, W. C. et al., 107, P.20156 (2-20); P.20255 13c); 2396 (2-la); 2491 (2-23); 2553, 2559 (1-42); 

(1-4a). 

2623 (2-25); 2630 (2-7); 2635 (2-5); 2539 (1-39); 

Steyermark, J. A. (et al.), 388, 487 (1-3); 507 (1-4b); 2719 (2-1 lb); 2852 (1-22); 2861 (2-23); 3965, 4201 

37602 (1-36); 38916 (2-3d); 39460 (1-30); 39618 (1- (2-7); 5111 (2-9); 5433 (1-31); 5745 (2-la); 5853 (2- 

42); 39934, 45663, 45981, 46103 (1-30); 52334 (1- 9); 6110 (2-23); 6156 (2-20); 6942 (1-42); 7104, 7522 

36); 54214(1-10); 54533 (1-42); 57872(2-13a); 57955 (1-31); 7535 (2-25); 7669 (1-4b); 8380, 8600 (1-22). 

(1-43); 60002 (1-4b); 60002a, 60364 (1-11); 60399 Veloso, H., 97 (1-25). 

(2-23); 60401 (2-3a); 60407 (2-1a); 60414 (2-23); Vellozo, -, 723 (2-la). 

60432 (1-4b); 60456 (1-43); 60665 (2-2); 60673 (1- Vellozo, H. P., 89, 723 (2-la); 1083 (2-lOa); 737 (1- 

11); 60800 (2-23); 75522 (1-11); 75540 (2-3a); 87307 33). 

(2-la); 87480 (1-4b); 89134 (2-la); 89452 (2-1 la); Vianna, E. C. et al., 5465 (1-25). 

89457 (2-3a); 89506 (2-23); 90368 (2-20); 90408 (2- Vickers, W. T., 86 (2-5). 

1 la); 90583 (1-4b); 90620 (2-23); 90644 (2-3a); 90736 Vieira, M. G. de et al., 985 (2-23); 1004 (2-20). 

(l-4b); 92850, 92936 (2-22); 92991 (1-3); 94180 (2- Villiers, J. F., 2088, 2111 (2-23). 

3a); 95170(1-42); 95378 (2-lb); 95711 (2-la); 99829 Vogl, C., 1379, s.n. (1-42). 

(1-42); 99945 (2-3d); 102082, 102211 (1-42); 102874 Vreden, J., 11717(1-20). 

(2-unnamed); 103026 (2-13a); 104296 (1-11); 104348 Wachenheim, H., 18 (2-23); 232 (2-1 la); 271 (2-2); 

(1-4b); 104466 (2-23), 105863 (2-lb); 106086 (2- 392 (2-13e); 426 (1-4a); 467 (2-23); s.n. (1-20, 2-la, 

la); 106118 (1-11); 106167 (1-31); 107064 (2-23); 2-23). 

113096 (2-3a); 114759 (1-4b); 115549(1-11); 116463 Warming, J. E. B., 1942, s.n. (1-25). 

(1-42); 116884 (1-4b); 116966 (2-1b); 119396 (2- Weaver, R. et al., 1677 (1-6).


Weberbauer, A., 3639 (2-la); 3702 (1-42); 4472 (1- 

44b). 

Wedel, H. von, 1548, 1733, 2879 (1-42). 

Westra, L. Y. T., 47303 (2-2). 

Whitton, B. A., 179 (1-25). 

Williams, LI. (et al.), 3347, 3984 (2-4); 4179 (1-42); 

4627 (2-4); 4688 (2-lOb); 5349 (1-42); 9983 (2-lb); 

10071 (1-42); 12008 (2-lla); 12348, 12394, 13895 

(1-42); 14675 (1-4b); 14833 (2-13a); 15240, 15363 

(1-31); 15812 (2-5). 

Williams, L. O. et al., 17817, 18014, 24005, 26529 (1- 

42). 

Williams, R. S., 501 (2-lb); 983 (1-3d); 1560 (2-la). 

Wilson-Browne, G., 549 (2-23). 

Wilson, C. L., 127 (1-42); 381 (1-30). 

Wilson, P., 606 (2-3d). 

Woodson, R. E. et al., 1893 (1-42). 

Woodworth, R. H. et al., 445 (1-42); 606 (1-4b); 660 

(1-42). 

Woytkowski, F., 5926 (1-42); 7306 (1-32). 

Wurdack, J. J. (et al.), 286 (1-4b); 2169 (2-23); 43485 

(1-40). 

Yele, -, DR-23 (1-26b). 

Zarucchi, J. L., 2145 (2-4); 2744 (2-la); 3720 (2-3a); 

3272 (2-3e). 

Zikan, J. F., s.n. (1-25). 

Zuccarini, J. G. (herb.), 103 (1-25). 

INDEX OF VERNACULAR NAMES OF POUROUMA 

amapati (la) 127 

cucuva (4) 147 

ama'yrary (1Oa) 156 

dacha uvillos (5) 149 

ambafba do vinho (13a) 168 

dakamtazshiuya (1 la) 161 

ambauva mansa (4) 147 

embauba da mata (la) 127 

ambauva do vinho (4, 13a) 147, 168 

embaubarana (7) 152 

a'ilea (3c) 138 

garguaba (3a) 135 

ambaibochi (23) 189 

granboesipapaja (10a, la, 13e, 17, 23) 156, 161, 173, 

ambaibillo (23) 189 

180, 189 

amia-yek (la) 127 

guagay (3d) 141 

bochoa tsaha (4) 147 

guarumo colorado (5) 149 

boesipapaja (13e) 173 

guarumo de montafia (3d) 141 

bois canon (la, 2, 3c, la, 13e, 17, 19, 23) 127, 132, guarumo macho (3d) 141 

138, 161, 173, 180, 181, 189 

guaumoutognac (13a) 168 

bois canon male (10a, 23) 156, 189 

gurucana (4) 147 

bois canon sauvage (10a) 156 

hembra (3a) 135 

boroma (2, 3a, 3c, 10a, Ila, 13e, 17) 132, 135, 138, imbafba (4, 5, 10a) 147, 149, 156 

156, 161, 173, 180 

imbauba bengue (17) 180 

boroma ibeberobana (2) 132 

imbauba branca (17) 180 

bospapaja(2, 3c, 10a, 1 la, 23) 132, 138, 156, 161,189 imbauba de cheirouvilha (17) 180 

bouchi papaie (23) 189 

imbauba do vinho (4, 13a) 147, 168 

bouchi papaye (13e, 17, 23) 173, 180, 189 

imbaiba mansa (4) 147 

buruma (la, 3c) 127, 138 

imbaibarana (la, 2, 3a, 5, 7, lla, 13b, 13c, 20, 23) 

caimar6n (4) 147 

127, 132, 135, 149, 152, 161, 169, 180, 189 

caimar6n de mico (3a) 135 

imbafbarana branca (5) 149 

caimar6n silvestre (4) 147 

imbaubarana folha peluda (5) 149 

caramuri (23) 189 

imbaubarana vermelha (10a) 156 

cay-bari-cay (3a) 135 

imbauba-tor6m (la) 127 

cay-wari-cay-yek (3a, 22, 23) 135, 186, 189 

imbaubinha (8) 152 

chaparro de agua (la) 127 

itararanga (la, 10a) 127, 156 

chiricaba (13a) 168 

kaibarikei (5) 149 

chullachaqui (20) 184 

kai-wa-rei-kei-yek (23) 189 

chullachaqui blanco (20) 184 

kaiwarikai (1 a) 161 

chullachaqui caspi (20) 184 

kalate (la) 127 

chumico (3d) 141 

kamoyuwa (10a) 156 

cirpe (3a) 135 

kaymbe'y (la) 127 

cirpe macho (3a, 1 b) 135, 162 

kukuma (13e) 173 

cocora(13a) 168 

kuluma (la) 127 

cocura (4) 147 

kulumasi (23) 189 

coiwaricoi-yek (23) 189 

kulumatelelel (3c) 138 

cormi (3a) 135 

lija (3d) 141 

cucura (2, 3a, 4, 5) 132, 135, 147, 149 

majagua (23) 189 

cucure (3a) 135 

male bois canon (3c, 23) 138, 189


manbospapaja (la, 17) 127, 180 

sirpe macho (3a) 135 

mangab6 (3e) 144 

sirpo (3e, 12a) 144, 162 

mano de leon (3d) 141 

sucufba (4) 147 

mapati (la, 4, 8, 13c, 15, 21) 127, 147, 152, 169, 177, sugkama(t) shuiya (la) 127 

186 

suia(4, lOb) 147, 159 

mapatirana (2, 3a, 3c, 10a, la, 16, 23) 132, 135, 138, suir shuina (lOb) 159 

156, 161, 177, 189 

tamaoquare (3a) 135 

mapaty (13c, 15, 21) 169, 177, 186 

tambor (lb) 129 

mimpa shuiya washi shuina ( la) 161 

tanaribe (4) 147 

obija (1Ob) 159 

tanta shuiya (3b) 137 

orumo de monte (3d) 141 

tararanga (lOa) 156 

otsepacho (3a) 135 

tararanga blanca (la) 127 

pacica (3d) 141 

tararanga vermelha (lOa) 156 

papaie (17) 180 

tentar shuina (3b) 137 

papaquillo (la) 127 

tinajero (3d) 141 

papaya del monte (la) 127 

torena (23) 189 

papaye (17) 180 

tourem (23) 189 

papaye apici (2, 1 la) 132, 161 

trumpet tree (3d) 141 

pau de jacu (la) 127 

tsaha (3a) 135 

pai shuina (3a, 13a) 135, 168 

tsakap suiya (la, 3a) 127, 135 

pai shuiya (13a) 168 

uhukamsuiya (10b) 159 

piraejo (3d) 141 

umbaouba (5) 149 

pourouma(3a, 3c, lla, 17) 135, 138, 161, 180 urumo blanco (12a) 162 

puruma(3a, 3c, lla, 17) 135, 138, 161, 180 uva (3d, 4, 12b) 141, 147, 163 

puruma (4) 147 

uva de macaco (lOa) 156 

purumai (13b, 23) 169, 189 

uva de(l) monte (3d, 4, 13c) 141, 147, 169 

puruma-y (4) 147 

uva silvestre (4, 23) 147, 189 

sacha uvilla (4, 9, 13a, 15, 23) 147, 155, 168, 177, 189 uva medueda (3a) 135 

sacha uvillos (20) 184 

uvilla(la,4, 5, 9, 10b, 1a, 15,23) 127, 147, 149, 155, 

sadajii (4) 147 

159, 161, 177, 189 

sadha'fhi (3a) 135 

uvilla blanca (la, 23) 127, 189 

sandpaper (la) 127 

uvilla lanuda (23) 189 

sa-ouro (10a) 156 

uvo (4) 147 

sarasara (3a, 5, 13b) 135, 149, 169 

wilaupiyua (13e) 173 

shewantoqui (4) 147 

yagrumo negro (lb) 129 

shuiya (la, 3b) 127, 137 

yagrumo-sunsun (10a) 156 

shuvija (23) 189 

yahal (3d) 141 

sirpe (3a, 12a) 135, 162 

yaryara (2, 3c, lOa, lla, 17) 132, 138, 156, 161, 180 

INDEX OF SCIENTIFIC NAMES 

Synonyms are in italics. Page numbers in boldface indicate primary page references. Page numbers 

with an asterisk (*) indicate pages with illustrations or maps. 

Allomeris 114 

Azteca 4 

Brosimum 66 

microcarpon 66 

Cebus apella 189 

Cecropia 2, 3, 9, 10, 15, 113, 115 

peltata 3 

Cecropiaceae 2, 3, 5, 15, 110, 113 

Clusia 4 

Coussapoa 2, 3, 4, 5, 6*, 7*, 8*, 9, 10, 15, 16, 17-28 

(keys), 112, 113, 114, 115, 194 

acutifolia 84 

angustifolia 11*, 28, 29*, 58, 97 

apoda 53 

arachnoidea 10, 11*, 30, 31*, 50, 77, 82 

araneosa 99 

argentea 10, 11*, 30, 32*, 73 

asperifolia 4, 5, 7, 8*, 10, 11*, 15, 16, 33, 34 (key), 

99, 104 

subsp. asperifolia 10, 11*, 34, 35*, 63 

subsp. magnifolia 11*, 36, 37*, 39, 99 

subsp. rhamnoides 11*, 38, 39* 

batavorum 11*, 38, 40*, 99 

boliviana 99 

brenesii 86 

brevipes 7, 11*, 41, 42* 

cardonaei 36 

cayennensis 34 

chagresiana 34 

chocoensis 11*, 41, 43* 

cinnamomea 11*, 44, 45*, 95 

cinnamomifolia 5, 10, 11*, 15, 16, 44, 46*, 86, 97

Index of Scientific Names 207 

contorta 5, 7, 11*, 41, 44, 47*, 48, 66 

cornifolia 87 

crassivenosa 10, 11*, 15, 48, 49*, 56 

cuneata 71 

cupularis 11*, 50, 51*, 77, 82 

curranii 4, 12*, 50, 52*, 58 

danielis 99 

dealbata 109 

dolichandra 109 

donnell-smithii 99 

duquei 7, 10, 12*, 53, 54*, 77, 104 

echinata 12*, 53, 55*, 61, 108 

eggersii 99, 104, 105 

emarginata 109, 186 

embirana 99 

fagifolia 71 

ferruginea 12*, 15, 56* 

ficina 34 

floccosa 12*, 52, 57* 

fontanesiana 66 

froesii 61 

fulvescens 14*, 105, 107*, 108 

glaberrima 12*, 58, 59* 

grandiceps 99 

herthae 12*, 56, 58, 60*, 61, 108 

hirsuta 84 

hypochlora 36 

incomitata 50 

intermedia 75 

krukovii 110, 168 

laevigata 110 

latifolia 9*, 10, 12*, 15, 16, 61, 62*, 63, 71, 73, 82, 

86, 105 

var. obovata 61 

lawrancei 99 

lehmannii 99 

leopoldii 110 

leprieurii 12*, 15, 63, 64*, 66, 93 

longepedunculata 13*, 63, 65*, 77, 93 

macarenensis 101 

var. antioquiensis 101 

macerrima 12*, 63, 66, 67*, 73, 92 

magnifolia 36 

manuensis 13*, 66, 68* 

martiana 99 

microcarpa 5, 7, 12*, 15, 63, 66, 69*, 73, 90 

microcephala 12*, 33, 63, 66, 71, 82, 90 

subsp. corifolia 89 

subsp. microcephala 72* 

mutisii 101 

napoensis 12*, 73, 74*, 84 

nitida 5, 10, 13*, 50, 75, 76*, 77, 82, 108 

nymphaeifolia 13*, 15, 61, 66, 77, 78*, 93 

obovata 61, 110 

oligoneura 86 

oligocephala 13*, 48, 77, 79*, 80, 93 

orthoneura 10, 13*, 30, 50, 63, 77, 80, 81*, 82 

ovalifolia 13*, 82, 83*, 84, 108 

pachyphylla 13*, 63, 73, 84, 85* 

panamensis 99 

parviceps 7, 13*, 16, 44, 56, 86, 87*, 90, 108 

parvifolia 5, 12*, 73, 87, 88*, 89*, 92 

pittieri 110 

planitiensis 101 

plicata 110 

prancei 80, 82 

puberula 84 

purpusii 10, 13*, 16, 90, 91* 

rekoi 110 

rhamnoides 38 

rotunda 97 

ruizii 36 

scabra 13*, 90, 92* 

schottii 66 

var. lanceolata 66 

var. longifolia 66 

schunkei 75 

setosa 84 

sprucei 13*, 15, 63, 66, 80, 92, 93* 

standleyi 99 

steyermarkii 48 

subcrenata 99 

subincana 99 

tessmannii 14*, 15, 44, 77, 93, 94*, 104 

tolimensis 14*, 108, 109* 

trinervia 4, 5, 10, 14*, 16, 44, 59, 95, 96*, 97 

valaria 108, 110* 

vanniifolia 5, 14*, 39, 97, 98* 

vellerea 99 

villosa 4, 5, 8*, 10, 14*, 15, 53, 77, 95, 99, 100*, 

110 

viridifolia 7, 10, 14*, 15, 16, 63, 82, 105, 106* 

var. tenuifolia 105 

volaria 14* 

warburgiana 50 

williamsii 80 

Crematogaster 114 

Ficus 4, 110 

intermarginalis 110 

Moraceae 2, 3, 5, 15, 110 

Musanga 2, 9, 15, 113 

Myrianthus 2, 5, 9, 15, 112, 113, 115 

Poikilospermum 2, 3, 15, 113 

subgen. Ligulistigma 15 

Poulsenia 110 

armata 110 

Pourouma 2, 3, 5, 9, 10, 15, 110, 111, 113, 114, 116, 

117-121 (key), 194 

acuminata 113, 114, 116, 173, 175*, 176*, 193 

acutiflora 123 

albistipulata 166 

apaporiensis 160, 161 

forma macrophylla 160 

apiculata Benoist 168 

apiculata Mildbraed 168 

aspera 116, 133 

subsp. digitata 115 

aurea 186, 189 

bicolor 111, 112, 114, 115, 122, 123, 132, 132-133 

(key) 

subsp. bicolor 123,133, 134*, 143*, 149, 193, 194 

subsp. chocoana 115, 141, 142*, 143*, 181 

subsp. digitata 115, 137, 138*, 139, 143* 

subsp. scobina 115, 122, 123,139, 140*, 141, 143*, 

144 

subsp. tessmannii 115, 136*, 137, 143*


bolivarensis 111, 113, 114, 115, 116, 176*, 186, 

187*, 189 

camaratana 133 

cecropiifolia 110, 111, 113, 114, 116,139, 143*, 144, 

145*, 147, 194 

chocoana 141 

cinerascens 123 

crassivenia 139 

crassivenosa 133 

cuatrecasasii 186 

cucura 112, 114, 115, 116, 147, 148*, 157*, 193, 

194 

cuspidata 113, 114, 116, 149, 157* 

digitata 137 

edulis 144 

elliptica 114, 176*, 184, 185* 

essequiboensis 169 

ferruginea 111, 113, 114, 116, 176*, 177, 178*, 184, 

193 

folleata 186 

formicarum 112, 113, 114,115,116,152,153*, 155, 

157* 

fulginea 123 

garciana 147 

guianensis 111, 114, 115, 116, 122, 123 

subsp. guianensis 122, 123, 124*, 126*, 128, 129, 

132 

subsp. venezuelensis 114, 122, 126*, 128, 129* 

herrerensis 115, 116, 171*, 177, 190, 192*, 193 

heterophylla 123, 155 

hirsutipetiolata 111, 114, 115, 116, 162 (key), 190 

subsp. hirsutipetiolata 162, 164* 

subsp. hispida 162, 165* 

hispida 162 

isophlebia 186 

jaramilloi 159 

johnstonii 141 

jussiaeana 156 

laevis 180 

lawrancei 133 

longipendula 181 

maroniensis 172 

melinonii 5, 113, 114, 115, 116, 155, 159, 160 (key) 

subsp. glabrata 112, 115, 157*, 161, 163* 

subsp. melinonii 115, 156, 157*, 158*, 160, 193, 

194 

mildbraediana 123 

minor 3, 5, 109, 111, 112, 113, 114, 116, 176*, 186, 

189 

mollis 15, 114, 115, 116, 155, 155-156 (key), 160, 

193 

subsp. mollis 115, 156, 157*, 158*, 159, 160 

subsp. triloba 114, 115, 156, 157*, 193 

multifida 144 

myrmecophila 111, 112, 113, 114, 115, 116, 150, 

151*, 152, 157* 

napoensis 113, 114, 115, 116, 171*, 190, 191* 

oraria 111, 112, 114, 115, 176*, 180, 190 

ovata 111, 112, 113, 114, 115, 116, 176*, 177,181, 

183* 

palmata 123 

paraensis 194 

phaeotricha 1 111,113, 115,116,152,154*, 155,176* 

populifolia 173 

radula 123 

retusa 194 

sapida 144, 147 

saulensis 114, 115, 116, 176, 181, 182*, 184 

scabra 123 

scobina 139 

schultesii 133 

steyermarkii 130 

stipulacea 111, 112, 114, 115, 116, 173, 174*, 176* 

subplicata 186 

substrigosa 123 

subtriloba 123 

tergoscabra 149 

tessmannii 137 

tomentosa5, 15, 110, 111, 112, 114, 115, 116,164, 

166 (key), 193, 194 

subsp. apiculata 115, 168, 171*, 173 

subsp. essequiboensis 115, 169, 171* 

subsp. maroniensis 115, 170, 171*, 172* 

subsp. persecta 113, 115, 169, 170*, 171* 

subsp. tomentosa 115, 166, 167*, 171*, 173 

trianae 194 

triloba Klotzsch 159 

triloba Trecul 156, 159 

ulei 194 

umbellata 186 

umbellifera 186 

uvifera 144 

velutina 114, 115, 116, 126*, 128, 129, 131* 

venezuelensis 128 

villosa 111, 176*, 177, 179* 

Schefflera 4 

Urostigma 10 

intermarginala 110 

Urticaceae 2, 3, 15

Cecropiaceae: Coussapoa and Pourouma, with an ... - CNCFlora

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