Cecropiaceae: Coussapoa and Pourouma, with an ... - CNCFlora
Cecropiaceae: Coussapoa and Pourouma, with an ... - CNCFlora
Cecropiaceae: Coussapoa and Pourouma, with an ... - CNCFlora
You also want an ePaper? Increase the reach of your titles
YUMPU automatically turns print PDFs into web optimized ePapers that Google loves.
Org<strong>an</strong>ization for Flora Neotropica<br />
<strong>Cecropiaceae</strong>: <strong>Coussapoa</strong> <strong><strong>an</strong>d</strong> <strong>Pourouma</strong>, <strong>with</strong> <strong>an</strong> Introduction to the Family<br />
Author(s): C. C. Berg, R. W. A. P. Akkerm<strong>an</strong>s, E. C. H. v<strong>an</strong> Heusden<br />
Reviewed work(s):<br />
Source: Flora Neotropica, Vol. 51, <strong>Cecropiaceae</strong>: <strong>Coussapoa</strong> <strong><strong>an</strong>d</strong> <strong>Pourouma</strong>, <strong>with</strong> <strong>an</strong> Introduction<br />
to the Family (Apr. 11, 1990), pp. 1-208<br />
Published by: New York Bot<strong>an</strong>ical Garden Press on behalf of Org<strong>an</strong>ization for Flora Neotropica<br />
Stable URL: http://www.jstor.org/stable/4393813 .<br />
Accessed: 23/03/2012 09:29<br />
Your use of the JSTOR archive indicates your accept<strong>an</strong>ce of the Terms & Conditions of Use, available at .<br />
http://www.jstor.org/page/info/about/policies/terms.jsp<br />
JSTOR is a not-for-profit service that helps scholars, researchers, <strong><strong>an</strong>d</strong> students discover, use, <strong><strong>an</strong>d</strong> build upon a wide r<strong>an</strong>ge of<br />
content in a trusted digital archive. We use information technology <strong><strong>an</strong>d</strong> tools to increase productivity <strong><strong>an</strong>d</strong> facilitate new forms<br />
of scholarship. For more information about JSTOR, please contact support@jstor.org.<br />
http://www.jstor.org<br />
New York Bot<strong>an</strong>ical Garden Press <strong><strong>an</strong>d</strong> Org<strong>an</strong>ization for Flora Neotropica are collaborating <strong>with</strong> JSTOR to<br />
digitize, preserve <strong><strong>an</strong>d</strong> extend access to Flora Neotropica.
FLORA NEOTROPIC<br />
MONOGRAPH 51<br />
CECROPIACEAE: COUSSAPOA AND<br />
POUROUMA, WITH AN INTRODUCTION<br />
TO THE FAMILY<br />
by<br />
C. C. Berg, R. W. A. P. Akkerm<strong>an</strong>s <strong><strong>an</strong>d</strong> E. C. H. v<strong>an</strong> Heusden<br />
. i ., """<br />
///A<br />
"^C^ - - -- - -------------------<br />
TIOPIC<br />
OF CANCER<br />
NEOTROPICsAh<br />
Published for<br />
Org<strong>an</strong>ization for Flora Neotropica<br />
by<br />
The New York Bot<strong>an</strong>ical Garden<br />
New York<br />
Issued 11 April 1990
Copyright ? 1990<br />
The New York Bot<strong>an</strong>ical Garden<br />
Published by<br />
The New York Bot<strong>an</strong>ical Garden<br />
Bronx, New York 10458<br />
International St<strong><strong>an</strong>d</strong>ard Serial Number 0071-5794<br />
Library of Congress Cataloging in Publication Data<br />
Flora neotropica. - Monograph no. 1 - New York: Published<br />
for Org<strong>an</strong>ization for Flora Neotropica by the New York<br />
Bot<strong>an</strong>ical Garden, 1968-<br />
v.: ill.; 26 cm.<br />
Irregular.<br />
Each issue has distinctive title.<br />
Separately cataloged <strong><strong>an</strong>d</strong> classified in LC before monograph no. 40.<br />
ISSN 0071-5794 = Flora neotropica.<br />
1. Bot<strong>an</strong>y-Latin America-Classification-Collected works. 2. Bot<strong>an</strong>y-<br />
Tropics-Classification-Collected works. 3. Bot<strong>an</strong>y-Classification-Col-<br />
lected works. I. Org<strong>an</strong>ization for Flora Neotropica. II. New York Bot<strong>an</strong>ical<br />
Garden.<br />
QK205.F58 581.98'012-dcl9 85-647083<br />
AACR 2 MARC-S<br />
Library of Congress [8508]<br />
ISBN 0-89327-352-x
CECROPIACEAE: COUSSAPOA AND POUROUMA,<br />
WITH AN INTRODUCTION TO THE FAMILY<br />
C. C. BERG,1 R. W. A. P. AKKERMANS,2 <strong><strong>an</strong>d</strong> E. C. H. v<strong>an</strong> Heusden3<br />
CONTENTS<br />
Abstract .................................................................. 2<br />
Introduction to the Family (C. C. BERG) ....... ..................................... ............ 2<br />
History <strong><strong>an</strong>d</strong> Circumscription of the Family ..... ................. ............................ 2<br />
Description of the Family ......... ........................................ .. .............. 3<br />
Key to the Neotropical Genera ................................................ ............. 3<br />
Genus Description of Cecropia .... ......................................................... 3<br />
<strong>Coussapoa</strong> (C. C. BERG AND R. W. A. P. AKKER NS) ............................................ 4<br />
In tro d u ctio n ............................................................................. 4<br />
Taxonom ic H istory ....................................................................... 4<br />
M o rp h o logy ........... ........................ ........................................... 4<br />
A natom y (K . BoNSEN ) .................................................................... 8<br />
Distribution <strong><strong>an</strong>d</strong> Ecology ......................... ......................................... 10<br />
Systematic Position of the Genus ........................................................... 15<br />
Systematic Arr<strong>an</strong>gement of the Species ...................................................... 15<br />
Taxonom y ........ ........................ .............................................. 16<br />
Generic D escription .................................................................. 16<br />
K eys to the Species .................. ........ ......................................... 17<br />
Treatments of the Species <strong><strong>an</strong>d</strong> Subspecies .... .................................... 28<br />
U nnam ed Collections .... ........................ .......... .......................... 108<br />
Names Not Included .................................................................. 109<br />
<strong>Pourouma</strong> (C. C. BERG AND E. C. H. VAN HEUSDEN) .............................................. 110<br />
Introduction ............................................................................. 110<br />
Taxonom ic H istory ....................................................................... 111<br />
M orphology .............................................................................. 111<br />
Phenology ............................................................................... 113<br />
Pollination .................. .... ......................................................... 113<br />
Dispersal ............................................................................... 114<br />
R elations w ith A nts ....................................................................... 114<br />
Distribution <strong><strong>an</strong>d</strong> Ecology ......... ...... ................................ ................... 114<br />
Systematic Position of the Genus .... ..................... .................................. 115<br />
Systematic Arr<strong>an</strong>gement of the Species ...................................................... 115<br />
U ses .................................................................................... 116<br />
Taxonom y ............................................................................... 116<br />
Generic Description .... .. ........................................................ , ... 116<br />
Key to the Species <strong><strong>an</strong>d</strong> Subspecies ...... ................................................ 117<br />
Treatments of the Species <strong><strong>an</strong>d</strong> Subspecies ............................................. 122<br />
Unnamed Collections ............. . ............................................ 193<br />
Names Excluded ................................................................. .... 194<br />
A cknow ledgm ents ............................................................................ 194<br />
Literature Cited .............. .. ...... ................................... ..................... 194<br />
N um erical List of Taxa ......... .............................................................. 196<br />
L ist of E xsiccatae ............................................................................. 197<br />
Index of Vernacular Names . ................................................................... 205<br />
Index of Scientific Names ................................... .............................. .. .. 206<br />
Arboretum <strong><strong>an</strong>d</strong> Bot<strong>an</strong>ical Garden (ARBOHA), University of Bergen, N-5067 Store Milde, Norway.<br />
2 Institute for Systematic Bot<strong>an</strong>y, State University of Utrecht, Heidelbergla<strong>an</strong> 2, 3508 TC Utrecht, The<br />
Netherl<strong><strong>an</strong>d</strong>s.<br />
3 Institute for Systematic Bot<strong>an</strong>y, State University of Utrecht, Heidelbergla<strong>an</strong> 2, 3508 TC Utrecht, The<br />
Netherl<strong><strong>an</strong>d</strong>s.<br />
4 Institute for Systematic Bot<strong>an</strong>y, State University of Utrecht, Heidelbergla<strong>an</strong> 2, 3508 TC Utrecht, The<br />
Netherl<strong><strong>an</strong>d</strong>s.<br />
1
2 Flora Neotropica<br />
ABSTRACT<br />
Berg, C. C. (Arboretum <strong><strong>an</strong>d</strong> Bot<strong>an</strong>ical Garden [ARBOHA], University of Bergen, N-5067<br />
Store Milde, Norway), R. W. A. P. Akkerm<strong>an</strong>s <strong><strong>an</strong>d</strong> E. C. H. v<strong>an</strong> Heusden (Institute for<br />
Systematic Bot<strong>an</strong>y, State University of Utrecht, Heidelbergla<strong>an</strong> 2, 3508 TC Utrecht, The<br />
Netherl<strong><strong>an</strong>d</strong>s). <strong>Cecropiaceae</strong>: <strong>Coussapoa</strong> <strong><strong>an</strong>d</strong> <strong>Pourouma</strong>. Flora Neotropica 51: 1-000. 1990.<br />
The present monograph comprises the revisions of two of the three Neotropical genera of<br />
the <strong>Cecropiaceae</strong>: <strong>Coussapoa</strong> <strong><strong>an</strong>d</strong> <strong>Pourouma</strong>. For <strong>Coussapoa</strong> 46 species are recognized at<br />
present; 15 of them have been described as new during the present study. All or most species<br />
are hemi-epiphytes. They have entire leaves <strong><strong>an</strong>d</strong> are dioecious. The small simple flowers<br />
are borne in uni- to pluricapitate inflorescences. The staminate flowers have one stamen or<br />
two or three connate stamens. The fruits are small. The genus r<strong>an</strong>ges from southern Mexico<br />
to south-eastern Brazil <strong><strong>an</strong>d</strong> is absent in the West Indies. Most species are components of<br />
lowl<strong><strong>an</strong>d</strong> rain forest habitats. Few species are wide-spread, most species have relatively small<br />
or even very small (known) r<strong>an</strong>ges of distribution. The species are morphologically clearcut<br />
<strong><strong>an</strong>d</strong> most of them rather uniform. For several reasons, e.g., lack of staminate material,<br />
the species are (provisionally) arr<strong>an</strong>ged in alphabetical sequences. Little is known about the<br />
biology of the genus. For <strong>Pourouma</strong> 25 species are presently recognized. Four taxa have<br />
been described as new during the preparation of this revision. All species are terrestrial,<br />
usually small to medium-sized trees. They have entire to palmately lobed to parted leaves<br />
<strong><strong>an</strong>d</strong> are dioecious. The pistillate flowers are pedicellate <strong><strong>an</strong>d</strong> are arr<strong>an</strong>ged in repeatedly<br />
br<strong>an</strong>ched to subumbellate inflorescences. The small staminate flowers are pedicellate or<br />
sessile <strong><strong>an</strong>d</strong> occur + loosely glomerate or in globose heads. The stamens are free in all species<br />
but one. The fruits are relatively large. The genus r<strong>an</strong>ges from Central America to southeastern<br />
Brazil <strong><strong>an</strong>d</strong> is absent in the West Indies. Most of them occur in + disturbed <strong><strong>an</strong>d</strong>/or<br />
+ open places in lowl<strong><strong>an</strong>d</strong> rain forest. Besides a number of clear-cut <strong><strong>an</strong>d</strong> slightly variable<br />
species the genus comprises a number of (very) variable <strong><strong>an</strong>d</strong> + complex species: P. bicolor,<br />
P. cucura, P. gui<strong>an</strong>ensis, P. melinonii, P. mollis, <strong><strong>an</strong>d</strong> P. tomentosa. The former three <strong><strong>an</strong>d</strong><br />
the latter three species are often difficult to separate from each other (<strong><strong>an</strong>d</strong> some other species).<br />
These six taxa <strong><strong>an</strong>d</strong> some others show a considerable plasticity, especially <strong>with</strong> regard to the<br />
leaf shape, leaf dimension, <strong><strong>an</strong>d</strong> composition of the indument, <strong><strong>an</strong>d</strong> most prominently so in<br />
the Upper Amazon Basin. Little is known about the biology of the genus. P. cecropiifolia<br />
is in cultivation as a fruit tree.<br />
INTRODUCTION TO<br />
THE FAMILY<br />
The present monograph is comprised of re-<br />
visions of two Neotropical genera of the Ce-<br />
cropiaceae, <strong>Coussapoa</strong> <strong><strong>an</strong>d</strong> <strong>Pourouma</strong>. To place<br />
these revisions in a proper setting, there is pro-<br />
vided first a history <strong><strong>an</strong>d</strong> circumscription of the<br />
family, along <strong>with</strong> a brief discussion of the char-<br />
acters that separate the family from the related<br />
Moraceae <strong><strong>an</strong>d</strong> Urticaceae, a description of the<br />
family <strong><strong>an</strong>d</strong> the type genus, Cecropia, <strong><strong>an</strong>d</strong> a key<br />
to the Neotropical genera.<br />
History <strong><strong>an</strong>d</strong> Circumscription<br />
of the Family<br />
The family was proposed by Berg (1978a) <strong><strong>an</strong>d</strong><br />
accepted by Cronquist (1981). It comprises six<br />
genera: three (Cecropia, <strong>Coussapoa</strong> <strong><strong>an</strong>d</strong> <strong>Pourouma</strong>)<br />
that are Neotropical, two (Mus<strong>an</strong>ga <strong><strong>an</strong>d</strong><br />
Myri<strong>an</strong>thus) Afric<strong>an</strong>, <strong><strong>an</strong>d</strong> <strong>an</strong> Asi<strong>an</strong>-Australasi<strong>an</strong><br />
genus (Poikilospermum). The majority of the<br />
members of the family constituted subfamily<br />
Conocephaloideae in Engler's Moraceae (Engler,<br />
1889), but Poikilospermum <strong><strong>an</strong>d</strong> some of the<br />
species presently included in it were included in<br />
the Urticaceae. Chew Wee-lek (1963) proposed<br />
r<strong>an</strong>king the four microspermous genera (Cecropia,<br />
<strong>Coussapoa</strong>, Poikilospermum <strong><strong>an</strong>d</strong> Mus<strong>an</strong>ga)<br />
under the Urticaceae <strong><strong>an</strong>d</strong> leave the two macrospermous<br />
genera (Myri<strong>an</strong>thus <strong><strong>an</strong>d</strong> <strong>Pourouma</strong>) in<br />
the Moraceae. Corer (1962), however, regarded<br />
all six genera as members of the Urticaceae. The<br />
creation of a new family appeared to be <strong>an</strong> appropriate<br />
resolution to <strong>an</strong> unsatisfactory situation.<br />
Several features suggest that the <strong>Cecropiaceae</strong>
Introduction<br />
are more closely related to the Urticaceae th<strong>an</strong> attached, entire, palmately-incised, or peltate <strong><strong>an</strong>d</strong><br />
to the Moraceae. Poikilospermum shows more radially-incised; venation pinnate, (sub)palmate,<br />
urticaceous features, including the typical urti- trinervate, or radial; stipules fully amplexicaul<br />
caceous cystoliths, th<strong>an</strong> the other five genera (cf. <strong><strong>an</strong>d</strong> connate. Inflorescences in the leaf axils, usu-<br />
Berg, 1989c; Bonsen & ter Welle, 1983). All gen- ally in pairs, unisexual, br<strong>an</strong>ched, <strong>with</strong> the flowera<br />
match the Urticaceae in the (sub)basal at- ers ? solitary or clustered in heads or spikes, or<br />
tachment of the ovule <strong><strong>an</strong>d</strong> the absence of milky unbr<strong>an</strong>ched <strong>with</strong> a single head or spike, bracteate<br />
sap. They are, however, clearly different from or ebracteate. Staminate flowers <strong>with</strong> 2-4 free or<br />
the Urticaceae in the absence of urticaceous (i.e., connate tepals; stamens 1-4; pistillode absent.<br />
explosive) stamens. Such stamens are inflexed in Pistillate flowers <strong>with</strong> 2-4 connate tepals; pistil<br />
the bud <strong><strong>an</strong>d</strong>, at <strong>an</strong>thesis, bend suddenly <strong><strong>an</strong>d</strong> one, ovary free from the peri<strong>an</strong>th, unilocular;<br />
elastically outward, throwing out the pollen. The ovule one, (sub)basally attached; stigma one,<br />
Afric<strong>an</strong> <strong><strong>an</strong>d</strong> Neotropical genera have a truly penicillate to peltate. Fruit <strong>an</strong> achene or somewoody<br />
habit (cf. Bonsen & ter Welle, 1983). Oth- times (in <strong>Coussapoa</strong>) ? drupaceous, enveloped<br />
er features which characterize the family are strict by a ? fleshy peri<strong>an</strong>th; seed small, <strong>with</strong> endodioecy,<br />
watery sap that turns black after exposure sperm, or large <strong><strong>an</strong>d</strong> <strong>with</strong>out endosperm; embryo<br />
to the air, the common occurrence of palmately- straight, cotyledons equal <strong><strong>an</strong>d</strong> flat or thickened.<br />
incised leaves <strong><strong>an</strong>d</strong>, in certain species, radially- A p<strong>an</strong>tropical family <strong>with</strong> six genera <strong><strong>an</strong>d</strong> apincised,<br />
peltate leaves, large stipules (connected proximately 180 species.<br />
<strong>with</strong> the protection of young inflorescences), <strong><strong>an</strong>d</strong><br />
adventitious aerial roots. While these roots are<br />
m<strong>an</strong>ifest as stilt-roots in most genera, in Cous- Key to the Neotropical Genera<br />
sapoa they are associated <strong>with</strong> the hemi-epi- 1. Lamina peltate, radially-incised. ...... Cecropia.<br />
phytic habit <strong><strong>an</strong>d</strong> in Poikilospermum <strong>with</strong> the 1. Lamina basally attached, entire or palmately-inclimbing<br />
habit.<br />
cised.<br />
The inflorescences of the 2. Staminate flowers <strong>with</strong> 1 or <strong>with</strong> 2 or 3 con-<br />
<strong>Cecropiaceae</strong> vary nate<br />
from being br<strong>an</strong>ched, <strong>with</strong> a + loose<br />
stamens; pistillate flowers sessile, in gloarr<strong>an</strong>ge-<br />
bose (to ellipsoid) heads; fruits small; stipule<br />
ment of the flowers (as in some staminate inflo- scars usually ascending. .......... <strong>Coussapoa</strong>.<br />
rescences of <strong>Pourouma</strong>), to unbr<strong>an</strong>ched, <strong>with</strong> a 2. Staminate flowers <strong>with</strong> 2-4 free stamens (or,<br />
single globose flower head (as in some pistillate<br />
in P. napoensis, 3-4 connate stamens); pistilinflorescences<br />
of <strong>Coussapoa</strong>), to a subumbellate<br />
late flowers pedicellate (to subsessile), solitary<br />
or in non-capitate clusters; fruits<br />
inflorescence (as in<br />
large; stipule<br />
<strong>Pourouma</strong> minor), or clusters scars horizontal. ................ <strong>Pourouma</strong>.<br />
of spikes enveloped by a spathe (as in most Cecropia<br />
species).<br />
In all <strong>Cecropiaceae</strong> the fruits are enveloped by<br />
a ? fleshy peri<strong>an</strong>th, <strong><strong>an</strong>d</strong> <strong>with</strong> regard to pollination<br />
<strong><strong>an</strong>d</strong> dispersal, the <strong>Cecropiaceae</strong> are more<br />
similar to the Moraceae th<strong>an</strong> to the Urticaceae.<br />
The genus Cecropia shows the most derived<br />
characters in the family, not only <strong>with</strong> regard to<br />
leaves <strong><strong>an</strong>d</strong> inflorescences, but also in the features<br />
connected <strong>with</strong> the adaptation to a mutualistic<br />
relation <strong>with</strong> <strong>an</strong>ts, as well as in the peculiar detachment<br />
(abscission) of the <strong>an</strong>thers (cf. Berg,<br />
1977b).<br />
Cecropia Loefling, Iter Hisp<strong>an</strong>. 272. 1758, nom.<br />
conserv. Type. C. peltata Linnaeus.<br />
Trees, terrestrial, usually <strong>with</strong> stilt-roots; internodes<br />
usually hollow. Lamina peltate, radially<br />
incised, venation radial, petiole mostly <strong>with</strong><br />
1-2 trichilia (patches of dense indument, usually<br />
<strong>with</strong> trichomes forming Miilleri<strong>an</strong> bodies) at the<br />
base. Inflorescences digitate clusters of spikes (or<br />
a single spike), usually enveloped by a spathe<br />
until <strong>an</strong>thesis, interfloral bracts absent. Peri<strong>an</strong>th<br />
tubular; stamens two; stigma penicillate to peltate.<br />
Fruit small, dry, enveloped by a greenish<br />
Description of the Family<br />
<strong>Cecropiaceae</strong><br />
Trees or shrubs, terrestrial or hemi-epiphytic,<br />
<strong>with</strong> aerial roots, <strong><strong>an</strong>d</strong> watery sap turning black<br />
upon exposure. Leaves in spirals; lamina basally<br />
fruiting peri<strong>an</strong>th.<br />
With approximately 80 species throughout the<br />
Neotropics, <strong>with</strong> a distinct concentration of<br />
species in the Ande<strong>an</strong> region.<br />
Revisions of <strong>Coussapoa</strong> <strong><strong>an</strong>d</strong> <strong>Pourouma</strong> are<br />
presented sequentially below.<br />
3
4 Flora Neotropica<br />
COUSSAPOA<br />
Engler's system of the Moraceae (Engler, 1889).<br />
This subfamily was tr<strong>an</strong>sferred to the Urticaceae<br />
Introduction<br />
by Corner (1962), but has recently been recognized<br />
as a<br />
<strong>Coussapoa</strong>, one of the three<br />
separate family <strong>Cecropiaceae</strong> (Berg,<br />
Neotropical genera<br />
of the <strong>Cecropiaceae</strong>, is a well-defined<br />
1978a; Cronquist, 1981).<br />
genus.<br />
The majority of its species are arboreous hemiepiphytes,<br />
a peculiar life-form shared <strong>with</strong> a few<br />
Morphology<br />
other genera like Ficus (subg. Urostigma), Clusia, Habit. - <strong>Coussapoa</strong> species are shrubs or more<br />
<strong><strong>an</strong>d</strong> Schefflera. The species are rather similar in<br />
their ecology <strong><strong>an</strong>d</strong> show little variation in<br />
commonly small to medium-sized trees (up to<br />
morca.<br />
20 m<br />
phological characters. Knowledge of their<br />
tall), but they c<strong>an</strong> sometimes form<br />
biolhuge<br />
wide-crowned trees up to 40 m tall,<br />
ogy is very limited.<br />
e.g., C. curr<strong>an</strong>ii<br />
<strong><strong>an</strong>d</strong> C. trinervia. On labels the<br />
The poor state of bot<strong>an</strong>ical exploration of<br />
height is<br />
m<strong>an</strong>y<br />
regions of the Neotropics has adversely affected<br />
probably not always correctly stated, because for<br />
the present revision, as is evident from the fact<br />
epiphytes the host often appears to be included.<br />
Most<br />
that it has not proved possible to make a<br />
<strong>Coussapoa</strong> species are hemi-epiphytic or<br />
systematic<br />
arr<strong>an</strong>gement of the species <strong><strong>an</strong>d</strong> from the<br />
secondarily terrestrial. Probably only C. trinervia<br />
is<br />
way in which the keys have had to be constructbasically<br />
terrestrial. As in Ficus, <strong>Coussapoa</strong><br />
ed. In about one-third of the species only one of<br />
species c<strong>an</strong> be epilithic. The (hemi-)epiphytic<br />
the sexes is known. M<strong>an</strong>y species are known from<br />
growth habit is evident in the presence of aerial<br />
roots. These aerial roots c<strong>an</strong> form baskets of roots<br />
only one or at most just a few specimens. More<br />
around the stem of the host tree. Sometimes, the<br />
th<strong>an</strong> one-quarter of the species now known have<br />
aerial root<br />
been recognized as new during the<br />
system consists of a main (vertical)<br />
preparation axis <strong>with</strong> at more or less<br />
of this revision; m<strong>an</strong>y of them are based on reregularly<br />
dist<strong>an</strong>ced pairs<br />
of horizontal br<strong>an</strong>ches<br />
cent collections. The increase in the number of<br />
clasping the trunk of the<br />
host tree. In<br />
species suggests that further exploration will lead<br />
general, hemi-epiphytic <strong>Coussapoa</strong><br />
to the discovery of additional<br />
species are not true (or vigorous) str<strong>an</strong>glers like<br />
species.<br />
hemi-epiphytic Ficus species.<br />
Most <strong>Coussapoa</strong> species have straight, stiff<br />
Taxonomic<br />
br<strong>an</strong>ches. The smaller br<strong>an</strong>ches c<strong>an</strong> be hollow<br />
History<br />
(widely so in C. asperifolia, but narrowly in sev-<br />
<strong>Coussapoa</strong> was established by Aublet (1775), eral other species, e.g., C. villosa). When br<strong>an</strong>chwho<br />
described two species from French Gui<strong>an</strong>a. es are hollow they are often inhabited by biting<br />
Names were added by Trecul (1847), Klotzsch <strong>an</strong>ts (Azteca?) or stinging <strong>an</strong>ts (often in the case<br />
(1847), Miquel (1853), <strong><strong>an</strong>d</strong> later on by St<strong><strong>an</strong>d</strong>ley of C. villosa). These insects bore openings to the<br />
(1919, 1924, 1930, 1937a, 1940), Mildbraed interior of the br<strong>an</strong>ches. Like (most?) other gen-<br />
(1928, 1942), <strong><strong>an</strong>d</strong> Cuatrecasas (1951, 1956b), era of the <strong>Cecropiaceae</strong>, when wounded Cousamong<br />
others, so that ca. ninety names were cre- sapoa species exude a (more or less mucilagiated<br />
for thirty-four of the taxa recognized in the nous) watery sap which turns black after exposure<br />
present paper. In the course of the preparation to the air.<br />
of present treatment 15 new species were de- Indumentum. -Four types of trichomes c<strong>an</strong><br />
scribed (Akkerm<strong>an</strong>s & Berg, 1982; Berg, 1983). be recognized:<br />
Taxonomic, mostly floristic, treatments of the<br />
genus are to be found in the work of Trecul (1847), 1. Unicellular arachnoid (cobwebby) hairs. These<br />
Flora brasiliensis (Miquel, 1853), Flora of Peru very thin, crinkled, interwoven hairs of dif-<br />
(Macbride, 1937), Flora of P<strong>an</strong>amai (Woodson & ferent lengths are mostly white, sometimes<br />
Schery, 1960), Flora of Suririame (Berg, 1975), brownish.<br />
<strong><strong>an</strong>d</strong> Flora Costaricensis (Burger, 1977). Thus, 2. Unicellular non-arachnoid, thicker <strong><strong>an</strong>d</strong> mostsince<br />
the treatments by Trecul <strong><strong>an</strong>d</strong> Miquel, the ly more or less straight hairs of different lengths<br />
genus has never been studied as a whole. <strong><strong>an</strong>d</strong> white, yellow, or brown.<br />
The genus belongs to a group of genera that 3. Pluricellular hairs-dark or pale brown, short<br />
constitute the subfamily Conocephaloideae in <strong><strong>an</strong>d</strong> subglobose to long <strong><strong>an</strong>d</strong> moniliform. These
<strong>Coussapoa</strong><br />
hairs are very common on young leaves, stipules,<br />
inflorescences, etc. In dry material they<br />
are more or less gr<strong>an</strong>ular (powdery) in appear<strong>an</strong>ce.<br />
4. Pearl bodies (or gl<strong><strong>an</strong>d</strong>s)-hyaline, succulent,<br />
more or less club-shaped, <strong><strong>an</strong>d</strong> confined almost<br />
entirely to the leaves. These trichomes<br />
are found in specimens of <strong>Coussapoa</strong> microcarpa<br />
<strong><strong>an</strong>d</strong> C. villosa cultivated in green houses.<br />
Pearl bodies are probably common in the genus,<br />
but are often not found in the field because<br />
they are easily detached; moreover, they<br />
are probably harvested by <strong>an</strong>ts (cf. O'Dowd,<br />
1982).<br />
As <strong>with</strong> m<strong>an</strong>y other genera of the <strong>Cecropiaceae</strong>,<br />
these four types of hairs are also often<br />
encountered in <strong>Coussapoa</strong> species. M<strong>an</strong>y species<br />
have a dense or rather dense indumentum. In<br />
several species arachnoid hairs are lacking <strong><strong>an</strong>d</strong><br />
most of these species have sparse indumentum<br />
as well.<br />
common in elliptic leaves, <strong><strong>an</strong>d</strong> in obovate leaves<br />
normally there is none. Venation types in the<br />
genus c<strong>an</strong> be seen in Figure 1.<br />
The ovate lamina <strong>with</strong> br<strong>an</strong>ched, basal, lateral<br />
veins <strong><strong>an</strong>d</strong> a relatively long petiole c<strong>an</strong> be regarded<br />
as the least derived type of leaf in the genus,<br />
as it shows the closest resembl<strong>an</strong>ce to leaves in<br />
other genera of the family, especially <strong>Pourouma</strong>.<br />
The leaves of P. minor resemble those of C. contorta<br />
<strong><strong>an</strong>d</strong> the leaves of P. tomentosa <strong><strong>an</strong>d</strong> P. meliononii<br />
are reminiscent of those of C. villosa <strong><strong>an</strong>d</strong><br />
C. nitida, respectively. The obovate, trinervate<br />
lamina <strong>with</strong> a short petiole c<strong>an</strong> be regarded as a<br />
derived type of leaf. Another type of leaf, which<br />
c<strong>an</strong> also be regarded as derived, is that found in,<br />
e.g., C. parvifolia <strong><strong>an</strong>d</strong> C. contorta. This type of<br />
leaf has <strong>an</strong> elliptic to subovate lamina <strong>with</strong> m<strong>an</strong>y<br />
unbr<strong>an</strong>ched lateral veins, a very regular venation,<br />
<strong><strong>an</strong>d</strong> a relatively short petiole. These features<br />
are characteristic more of the Moraceae th<strong>an</strong> of<br />
the <strong>Cecropiaceae</strong>.<br />
The indumentum of the leaves also shows some<br />
Leaves. -The petiole in most species is rela- variation in denseness <strong><strong>an</strong>d</strong> occurrence of differtively<br />
long-up to half the length of the lamina. ent types of hairs. The occurrence of a dense <strong><strong>an</strong>d</strong><br />
There is a tendency towards shortening of the varying indumentum appears to be weakly corpetiole<br />
(to one-tenth of the length of the lamina) related <strong>with</strong> primitiveness in other leaf characin<br />
species <strong>with</strong> derived (i.e., obovate, trinerved) ters.<br />
types of lamina. The relative length of the petiole C. contorta is distinct in having domatiumis<br />
not subject to great variation as is the case in like cavities in the axils of the lateral veins.<br />
related genera, such as Myri<strong>an</strong>thus <strong><strong>an</strong>d</strong> Pou- The stipules are connate, fully amplexicaul <strong><strong>an</strong>d</strong><br />
rouma.<br />
caducous, usually leaving oblique scars, char-<br />
The length of the lamina varies from more acteristic for the genus. The length varies conth<strong>an</strong><br />
50 cm to less th<strong>an</strong> 10 cm <strong><strong>an</strong>d</strong> more or less siderably in the genus. One c<strong>an</strong> roughly recognize<br />
parallels the ch<strong>an</strong>ge in the shape of the lamina three categories: long stipules (5-15 cm long or<br />
from cordiform, subreniform, or broadly ovate longer), medium-long stipules (1-5 cm long) <strong><strong>an</strong>d</strong><br />
to elliptic or oblong to obovate or subobovate; short stipules (less th<strong>an</strong> 1 cm long). Long stipules<br />
at the same time, this correlates <strong>with</strong> a gradation give protection for a longer time to young leaves<br />
in the leaf base from cordate to truncate to <strong><strong>an</strong>d</strong> developing inflorescences, but a correlation<br />
rounded to obtuse to subacute.<br />
between the type of inflorescence or habit pref-<br />
The venation is pinnate <strong><strong>an</strong>d</strong> brochidodromous erence <strong><strong>an</strong>d</strong> the length of the stipules is not ap<strong>with</strong><br />
a more or less regular parallel tertiary ve- parent.<br />
nation in the intercostal area. The venation may Inflorescences. - <strong>Coussapoa</strong>, like all other gentend<br />
toward subpalmate (C. v<strong>an</strong>nifolia), or may era of the <strong>Cecropiaceae</strong>, is dioecious. Monoebe<br />
more or less clearly trinervate (C. trinervia, cious specimens are rare.<br />
C. cinnamomifolia), <strong>with</strong> this development being The inflorescences occur in pairs in the leaf<br />
combined <strong>with</strong> a ch<strong>an</strong>ge towards a (sub)obvate axils, a feature common in the <strong>Cecropiaceae</strong> <strong><strong>an</strong>d</strong><br />
lamina. In ovate leaves the basal pair of lateral related families. The ramification of the inflovein<br />
br<strong>an</strong>ches is mostly br<strong>an</strong>ched. Especially in rescence is basically similar to that usually found<br />
large leaves, the other lateral veins show little in Myri<strong>an</strong>thus (see: De Ruiter, 1976) <strong><strong>an</strong>d</strong> Poubr<strong>an</strong>ching<br />
(being mostly only furcate); however, rouma. The common peduncle bears at its apex<br />
in C. asperifolia the other lateral veins are mostly (at least) three br<strong>an</strong>ches, a pattern repeated in<br />
br<strong>an</strong>ched. Br<strong>an</strong>ching of the lateral veins is less secondary <strong><strong>an</strong>d</strong> further ramifications. However,<br />
5
6 Flora Neotropica<br />
Ni<br />
3 4 5<br />
FIG. 1. Schematic drawings of types of venation in <strong>Coussapoa</strong>: 1, most lateral veins br<strong>an</strong>ched; 2, 3, only<br />
the basal pair of lateral veins br<strong>an</strong>ched; 4, lateral veins unbr<strong>an</strong>ched, the basal pair reaching the margin above<br />
the middle; 5, three-nerved.<br />
2
Morphology<br />
T<br />
FIG. 2. Schematic drawings of types of pistillate inflorescences in <strong>Coussapoa</strong>.<br />
FIG. 2. Schematic drawings of types of pistillate inflorescences in <strong>Coussapoa</strong>.<br />
reduction of the number of br<strong>an</strong>ches to two, re- bracts c<strong>an</strong> be subpeltate or more or less cucullate.<br />
sulting in a (sub)dichotomous ramification, is also Sometimes, these bracts are truly peltate--as in<br />
found, especially in the ultimate ramification, C. contorta. Bracts outside the flower heads are<br />
<strong><strong>an</strong>d</strong> may occur down to the primary ramification mostly scale-like <strong><strong>an</strong>d</strong> ovate to oblong.<br />
as well. Pistillate inflorescences are schematized Staminate Flowers.-The small staminate<br />
in Figure 2.<br />
flowers have a tubular peri<strong>an</strong>th which is 3-lobed<br />
The sessile flowers are arr<strong>an</strong>ged in terminal at the apex. The number of stamens is 3, 2, or<br />
globose heads. Solitary (often more or less abort- 1. In 3- <strong><strong>an</strong>d</strong> 2-staminate flowers the stamens are<br />
ed) flowers may occur elsewhere in the inflores- fused, forming a stalk <strong>with</strong>, respectively, 6 (or<br />
cence, but their presence is mostly connected <strong>with</strong> occasionally 5) or 4 thecae at the apex. The numpoints<br />
of br<strong>an</strong>ching. Solitary flowers are rare in ber of stamens is very const<strong>an</strong>t <strong><strong>an</strong>d</strong> may prove<br />
pistillate inflorescences.<br />
to be the (only?) basis for subdivision of the genus<br />
The staminate inflorescences are normally re- <strong><strong>an</strong>d</strong> arr<strong>an</strong>gement of the species. The length of<br />
peatedly br<strong>an</strong>ched <strong><strong>an</strong>d</strong> bear small terminal heads. the filaments varies from about as long as the<br />
In pistillate inflorescences, however, the number peri<strong>an</strong>th to more th<strong>an</strong> twice as long as the periof<br />
flower heads is often reduced to one, due to <strong>an</strong>th, the differences being distinctive at the<br />
reduction of the ramification <strong><strong>an</strong>d</strong>/or to fusion of species level.<br />
the flower heads. In some species, e.g., C. bre- Pistillate Flowers. -The small pistillate flowvipes,<br />
C. duquei, the flower heads are ellipsoid ers have a tubular peri<strong>an</strong>th <strong>with</strong> a narrow (alto<br />
clavate <strong><strong>an</strong>d</strong> combined <strong>with</strong> a very short pe- most) entire opening that allows the style through.<br />
duncle.<br />
The flowers are free, but in C. parviceps the mar-<br />
The inflorescences are bracteate in the major- gins of the flattened apices of the peri<strong>an</strong>ths of<br />
ity of the <strong>Coussapoa</strong> species. The bracts are mostly adjacent flowers cohere. The ovary is free <strong><strong>an</strong>d</strong><br />
confined to the flower heads, but sometimes oc- the stigma is penicillate to subpeltate.<br />
cur elsewhere in the inflorescence, as in the sta- Staminate <strong><strong>an</strong>d</strong> pistillate flowers <strong><strong>an</strong>d</strong> their asminate<br />
inflorescences of C. tessm<strong>an</strong>nii. In some sociated bracts are shown schematically in Figspecies,<br />
e.g., C. microcarpa <strong><strong>an</strong>d</strong> C. viridifolia, the ure 3.<br />
interfloral bracts are very small; they may also Fruits <strong><strong>an</strong>d</strong> Seeds.-The small fruits are free<br />
be few in number or sometimes even absent. In from the peri<strong>an</strong>th. The pericarp consists of a<br />
C. parviceps, the only species in which the pis- rather thick <strong><strong>an</strong>d</strong> crustaceous endocarp <strong><strong>an</strong>d</strong> a very<br />
tillate flowers are connate (or cohering), inter- thin to rather thick, slightly fleshy to mucilagifloral<br />
bracts are absent in the pistillate inflores- nous layer. Thus, depending on the features of<br />
cence but present in the staminate. This lack of the exocarp, the ellipsoid to ovoid, 1-3 mm long<br />
interfloral bracts is apparently a derived state. fruitlets are achene-like or more or less drupa-<br />
The interfloral bracts are more or less distinct- ceous. Fruits are depicted schematically in Figly<br />
spathulate, often <strong>with</strong> the lower part very nar- ure 4.<br />
row (<strong><strong>an</strong>d</strong> stipe-like). Relatively large interfloral In C. asperifolia, while the exocarp is present<br />
7
8 Flora Neotropica<br />
1 2 3 4 5 6 7 a ' 9.<br />
FIG. 3. Schematic drawings of flowers <strong><strong>an</strong>d</strong> interfloral bracts in <strong>Coussapoa</strong>: 1-3, staminate flowers, 1, <strong>with</strong><br />
three connate stamens, 2, <strong>with</strong> two connate stamens, 3, <strong>with</strong> one stamen; 4, pistillate flower; 5-9, various<br />
interfloral bracts.<br />
as a very thin (membr<strong>an</strong>aceous) outer layer, the<br />
mesocarp is relatively thick <strong><strong>an</strong>d</strong> exp<strong><strong>an</strong>d</strong>s in such<br />
a way as to push the endocarp body out of the<br />
peri<strong>an</strong>th, the apex of which becomes torn in the<br />
process. Whether this phenomenon is confined<br />
to C. asperifolia or also occurs in other species<br />
is not clear. At <strong>an</strong>y rate, it is not general in <strong>Coussapoa</strong>,<br />
as previously assumed (Berg, 1975, 1977a).<br />
In m<strong>an</strong>y (or all?) species the fruiting peri<strong>an</strong>th<br />
becomes more or less juicy, <strong><strong>an</strong>d</strong> it is also often<br />
colored-or<strong>an</strong>ge, yellow, or brown. Sometimes,<br />
as in C. villosa, the peri<strong>an</strong>th remains greenish,<br />
at least the apical part.<br />
<strong>Coussapoa</strong> is one of the four microspermous<br />
genera of the <strong>Cecropiaceae</strong>. In the genus microspermy<br />
is related to the (hemi-)epiphytic habit,<br />
as well as to the occurrence of a mucilaginous<br />
layer (cf. Berg, 1983b).<br />
Phenology, Pollination, <strong><strong>an</strong>d</strong> Dispersal. -<br />
Available herbarium <strong>an</strong>notations suggest that in<br />
general <strong>Coussapoa</strong> species flower (<strong><strong>an</strong>d</strong> fruit)<br />
throughout the year.<br />
Nothing is known about pollination in <strong>Coussapoa</strong>.<br />
Neither the structure of the inflorescences<br />
nor the features of the stamens suggest the occurrence<br />
of wind pollination, however.<br />
In most cases dispersal is probably endozo-<br />
ochorous, often by frugiferous birds, but also by<br />
monkeys (Dr. M. v<strong>an</strong> Roosmalen, pers. comm.).<br />
Exozoochorous dispersal may occur when the<br />
exocarp is more or less mucilaginous (<strong><strong>an</strong>d</strong> sticky),<br />
as in C. asperifolia.<br />
Anatomy<br />
By K. BONSEN<br />
Secondary Xylem. -General Properties. Col-<br />
or: light yellowish brown. Grain: straight, some-<br />
times slightly interlocked. Texture: medium.<br />
Odor <strong><strong>an</strong>d</strong> taste: not characteristic. Specific grav-<br />
ity: 0.50-0.75.<br />
Structure. Based on eight species, nine speci-<br />
mens, the growth rings are faint or absent. Ves-<br />
sels diffuse, round to oval, solitary (25-83%), <strong><strong>an</strong>d</strong><br />
in radial multiples <strong><strong>an</strong>d</strong> irregular clusters of 2-8<br />
(-21), (0-1)1-6(-11) per sq mm, diameter (200-)<br />
220-300(-340) gm. Vessel member length (400)<br />
475-600(-725) Mm. Perforations simple. Inter-<br />
vascular pits alternate, round or polygonal, 10-<br />
15 Lm. Thin-walled tyloses are usually present.<br />
Fibers non-septate, diameter 18-25 ,m, walls 2-<br />
3.5 tm, L/W ratio 2-5, gelatinous fibers usually<br />
present. Pits simple, mainly on the radial walls.<br />
a ad tfg<br />
1 2 34 5 6<br />
FIG. 4. Schematic drawings of fruits in <strong>Coussapoa</strong>: 1-4, C. asperifolia type, 1, 2, fruit in peri<strong>an</strong>th, 3, fruit<br />
in opened peri<strong>an</strong>th, 4, endocarp body; 5, 6, C. villosa type, fruit in peri<strong>an</strong>th. a, embryo; b, endosperm; c, testa;<br />
d, endocarp; e, mucilaginous mesocarp; f, exocarp; g, peri<strong>an</strong>th.
Anatomy<br />
, ,. : ?<br />
? ~ ~ ~ i<br />
I i ~ ~.: II~C ... ?<br />
~~~~:':-<br />
%,-: :<br />
;~ ~ ~<br />
' '" 2.'' ? .. ' , ' ? "'."' ..<br />
:i...<br />
t<br />
. ...-I. : .,<br />
:.<br />
' '<br />
%;.: ,,<br />
I,~~~~~.<br />
F?<br />
I<br />
'<br />
- i. " ' ~, c .?.h<br />
1;<br />
? t*l..:.<br />
.::.:.;.<br />
... ~ ,,~x ~ ~ .....~<br />
. "~~~-i<br />
FiG. 5. Woo of Co p~ltf'l9 , ,cos ci~ ~ lsci~(.22<br />
Length (875-)1100-1800(-2175) .m, F/V ratio<br />
2.5-3.7.<br />
Rays heterogeneous, uniseriate (21-35%) <strong><strong>an</strong>d</strong><br />
multiseriate (3-)4-7(-9) per mm, sheath cells<br />
present or absent. Uniseriate rays mainly composed<br />
of square to upright cells, ray height<br />
(200-)300-500(-980) gtm. Multiseriate rays<br />
composed of upright <strong><strong>an</strong>d</strong> procumbent cells, vertically<br />
compounded 1-10%, (450-)700-1100<br />
(-1600) ,m high, 3-6 cells in width, uniseriate<br />
parts (0-)1-2(-8) cells.<br />
Parenchyma paratracheal, b<strong><strong>an</strong>d</strong>ed, irregular,<br />
wavy, (0-)1-2(-3) per mm, (3-)5-9(-15) cells in<br />
width. Str<strong><strong>an</strong>d</strong>s 5-8(-14) cells, length (530-)600-<br />
710(-870) Mm, containing some to m<strong>an</strong>y rhombic<br />
crystals. Radial latex tubules have been observed<br />
in <strong>Coussapoa</strong> latifolia (see Fig. 5).<br />
The wood of <strong>Coussapoa</strong> resembles that of Myri<strong>an</strong>thus<br />
<strong><strong>an</strong>d</strong> some species of <strong>Pourouma</strong>. The<br />
presence of confluent-b<strong><strong>an</strong>d</strong>ed parenchyma in the<br />
wood of <strong>Coussapoa</strong> distinguishes this genus from<br />
the wood of Cecropia <strong><strong>an</strong>d</strong> Mus<strong>an</strong>ga (Bonsen &<br />
ter Welle, 1983).<br />
'<br />
. . . .<br />
Leaf Anatomy. -Structure. Based on 11<br />
species, 18 specimens.<br />
In surface view. Indumentum of thin, inter-<br />
woven, unicellular arachnoid hairs, abaxial pres-<br />
ent or absent; unicellular needle-shaped, often<br />
wavy hairs (mostly on abaxial surface, rarely on<br />
adaxial surface); adaxial, gl<strong><strong>an</strong>d</strong>ular hairs <strong>with</strong><br />
multicellular, globular heads on 3-5-celled, uni-<br />
seriate stalks, mostly in groups of 2-4, present<br />
or absent; abaxial, uniseriate, 6-10-celled, curved,<br />
gl<strong><strong>an</strong>d</strong>ular hairs <strong>with</strong> globular to elongated heads,<br />
mostly present; <strong><strong>an</strong>d</strong> conical papillae sometimes<br />
present. Epidermal cells polygonal; adaxial cells<br />
overlying large crystalliferous mesophyll cells<br />
forming a rosette. Stomata confined to abaxial<br />
surface, <strong>an</strong>omocytic, average length of guard cell<br />
pairs 15-20 gm, average width 12-18 um. Hy-<br />
dathodes formed by 10-15 water pores each,<br />
present or absent on adaxial surface. Minor veins<br />
usually very prominent in abaxial epidermis.<br />
In tr<strong>an</strong>sverse section. Lamina bifacial. Epi-<br />
dermal cells small, especially abaxially between<br />
the veins. Adaxial epidermal cells sometimes <strong>with</strong>
10 Flora Neotropica<br />
silicified outer walls. Stomata sometimes raised higher altitudes (from ca. 800 m) in the Gui<strong>an</strong>a<br />
above level of unspecialized cells. Adaxial hy- (=Guay<strong>an</strong>a) Highl<strong><strong>an</strong>d</strong> region, e.g., C. argentea<br />
podermis of two or three layers of parenchyma <strong><strong>an</strong>d</strong> C. viridifolia, while others are found in higher<br />
cells, including mucilage cells except for C. vil- altitudes (from ca. 900 m or from ca. 1400 m)<br />
losa. Mesophyll consisting of one layer of pali- in the Ande<strong>an</strong> region, e.g., C. cinnamomifolia<br />
sade cells, sometimes subdivided loose spongy <strong><strong>an</strong>d</strong> C. duquei; C. crassivenosa is found in both<br />
tissue, <strong>with</strong> or <strong>with</strong>out <strong>an</strong> intermediate layer. regions. Some species also occur in drier habitats,<br />
Veins <strong>with</strong> sclerenchymatous vertical bundle such as sav<strong>an</strong>na (C. viridifolia) or semi-decidusheath<br />
extensions (touching adaxial hypodermis ous forest (C. purpusii). Most lowl<strong><strong>an</strong>d</strong> species<br />
<strong><strong>an</strong>d</strong> abaxial epidermis). Midrib <strong>with</strong> a flat or appear to occur both in upl<strong><strong>an</strong>d</strong> <strong><strong>an</strong>d</strong> riverine forraised<br />
adaxial surface <strong><strong>an</strong>d</strong> a prominently raised est. A few species, like C. nitida, appear to show<br />
abaxial surface; peripheral ground tissue paren- a preference for riverside (varzea) forest. C. trichymatous<br />
to collenchymatous, interspersed <strong>with</strong> nervia, the only truly terrestrial species of the<br />
mucilage cells; vascular system composed of a genus, is confined to riverb<strong>an</strong>ks, apparently<br />
closed or variously interrupted cylinder, partly (only?) of black-water rivers.<br />
or wholly surrounded by sclerenchyma fibers, <strong><strong>an</strong>d</strong> Only two species, C. villosa <strong><strong>an</strong>d</strong> C. asperifolia,<br />
enclosing one or two rows of bundles which are have <strong>an</strong> extended distribution. Most of the other<br />
situated in the same direction as the most abaxial species are confined to, or at least distinctly asbundle<br />
of the cylinder. Vascular system of petiole sociated <strong>with</strong>, one of the phytogeographic resimilar.<br />
Crystals present as druses throughout the gions, such as eastern Brazil, Amazon Basin,<br />
mesophyll, in the petiole <strong><strong>an</strong>d</strong> midrib; rhombic eastern Gui<strong>an</strong>a (=Guay<strong>an</strong>a Lowl<strong><strong>an</strong>d</strong>) region,<br />
crystals sometimes present. Distinguishment of western Gui<strong>an</strong>a (=Guay<strong>an</strong>a Highl<strong><strong>an</strong>d</strong>) region,<br />
the inner structure of the leaves of <strong>Coussapoa</strong> Ande<strong>an</strong> region, Pacific coastal region, <strong><strong>an</strong>d</strong> Cenfrom<br />
those of the other Neotropical genera, Ce- tral America. M<strong>an</strong>y species are only known from<br />
cropia <strong><strong>an</strong>d</strong> <strong>Pourouma</strong>, c<strong>an</strong> be accomplished on a very small area.<br />
the basis of the petiole vascular system type Several species <strong>with</strong> medium-sized areas show<br />
(Bonsen & ter Welle, 1983). Some differences in a discontinuous distribution. It is, however, not<br />
the inner structure of some species <strong>with</strong>in the certain whether these discontinuities are real or<br />
genus <strong>Coussapoa</strong> are mentioned by Renner due to insufficient exploration. The Amazoni<strong>an</strong><br />
(1907).<br />
C. trinervia has a disjunct distribution, which<br />
appears to be related to its ecology (see above).<br />
Distribution <strong><strong>an</strong>d</strong> C. crassivenosa is known from<br />
Ecology<br />
four remote regions.<br />
C. latifolia <strong><strong>an</strong>d</strong> C. asperifolia ssp. asperi-<br />
The genus, being distinctly associated <strong>with</strong> the folia, common taxa of the eastern Gui<strong>an</strong>a region<br />
rain forest habitat, is widespread in tropical to eastern Parf, show similar gaps in distribution<br />
America, but is absent in the West Indies; see <strong><strong>an</strong>d</strong> are found segregated in more central parts<br />
Figures 6-9.<br />
of the Amazon Basin as well. If these gaps are<br />
Most species are components of lowl<strong><strong>an</strong>d</strong> rain real, the same factors may have caused the westforests.<br />
A few species, e.g., <strong>Coussapoa</strong> villosa <strong><strong>an</strong>d</strong> ern Parf gap in the distribution of C. nitida, as<br />
C. asperifolia, extend into submont<strong>an</strong>e or mon- well as the disjunct distribution of the species<br />
t<strong>an</strong>e forest. Some species are associated <strong>with</strong> pairs C. arachnoidea-C. orthoneura <strong><strong>an</strong>d</strong> C. le-<br />
FIG. 6. Distribution of <strong>Coussapoa</strong> <strong>an</strong>gustifolia, C. arachnoidea, C. argentea, C. asperifolia ssp. asperifolia,<br />
C. asperifolia ssp. magnifolia, C. asperifolia ssp. rhamnoides, C. batavorum, C. brevipes, C. chocoensis, C.<br />
cinnamonea, C. cinnamomifolia, C. contorta, C. crassivenosa, <strong><strong>an</strong>d</strong> C. cupularis.<br />
FIG. 7. Distribution of <strong>Coussapoa</strong> curr<strong>an</strong>ii, C. duquei, C. echinata, C. ferruginea, C. floccosa, C. glaberrima,<br />
C. herthae, C. latifolia, C. leprieurii, C. macerrima, C. microcarpa, C. microcephala, C. napoensis, <strong><strong>an</strong>d</strong> C.<br />
parvifolia.<br />
FIG. 8. Distribution of <strong>Coussapoa</strong> longepedunculata, C. m<strong>an</strong>uensis, C. nitida, C. nymphaeifolia, C. oligocephala,<br />
C. orthoneura, C. ovalifolia, C. pachyphylla, C. parviceps, C. purpusii, C. scabra, <strong><strong>an</strong>d</strong> C. sprucei. ?<br />
unnamed collection aff. C. ovalifolia.
Distribution <strong><strong>an</strong>d</strong> Ecology<br />
0 C.<strong>an</strong>gustifolia<br />
Aublet<br />
OC.arachnoidea Akkerm<strong>an</strong>s 6 Berg<br />
^^-^^'.L_ n //i C ^ 4-Lc
12 Flora Neotropica<br />
v* r<br />
r'^S'9n 7<br />
J<br />
(^^'^^J^'^<br />
- ^^Q^0 C.duquei St<strong><strong>an</strong>d</strong>ley<br />
A C.echinata Akkerm<strong>an</strong>s Berg<br />
* C.curr<strong>an</strong>li Blake i . e<br />
-<br />
T C.ferruginea Trecul<br />
| ? (% 'J/' / { , t<br />
__l<br />
|:<br />
A C.glaberrima<br />
Burger<br />
* C.herthee<br />
S Mildbred<br />
C.p_rvi fo ^g d C-latifolia Aublet<br />
C .floccosa i<br />
y<br />
Akkerm<strong>an</strong>s Berg .<br />
B<br />
C.cerricroc Akkerpa (SchQtt) Berg izzini ~ -<br />
0 C'microcephala Trdcul '<br />
V' C.parvifolia St<strong><strong>an</strong>d</strong>ley<br />
A C.napoensis Akkermmns & Berg<br />
,.
Distribution <strong><strong>an</strong>d</strong> Ecology<br />
OC.nitida Mfquel /). ^~:t ,._ A *C.nymphaeifolia St<strong><strong>an</strong>d</strong>ley<br />
~<br />
.AC.oligo ala Donnael Smith<br />
C.orthoneura St<strong><strong>an</strong>d</strong> ey<br />
*C.pachyphylla Akkerm<strong>an</strong>s & Berg 0 C.ovalifolia Trcul C .<br />
HC.pcyhl tcuatl Aknn Tra ~ t C.oval 1<br />
i fol ia<br />
C.parviceps St<strong><strong>an</strong>d</strong>ey\<br />
C.scabra Akkerm<strong>an</strong>s 6 Berg<br />
*C.sprucel<br />
Mildbraed<br />
\<br />
%<br />
13
1.trinervia<br />
*C.villosa Poeppig 6 Endlicher<br />
AC.viridifolia<br />
Cuatrecasas<br />
I<br />
C. tessm<strong>an</strong>nll M Ildbrwd<br />
A C. fulvescens C.C. Berg<br />
VC. tolimensis C.C. Berg<br />
Ce. voiaria C.C. Berg<br />
C<br />
Mil dbraed<br />
A C.v<strong>an</strong>nifolia Cuatrecasas<br />
FIG. 9. Distribution of <strong>Coussapoa</strong><br />
fulvescens, C. tessm<strong>an</strong>nii, C. tolimensis, C. trinervia, C. v<strong>an</strong>nifolia, C.<br />
villosa, C. viridifolia, <strong><strong>an</strong>d</strong> C. valaria (name is misspelled in legend).
Systematic Position 15<br />
prieurii-C. sprucei. This type of distribution<br />
may be connected <strong>with</strong> the distribution of Pleistocene<br />
rain forest refugia (cf. Pr<strong>an</strong>ce, 1977).<br />
Another remarkable discontinuity in distribution<br />
occurs <strong>with</strong> C. cinnamomifolia, a species of<br />
medium altitudes in the Ande<strong>an</strong> region. This<br />
discontinuity shows resembl<strong>an</strong>ces to that of two<br />
partial areas of C. crassivenosa. The Ecuador-<br />
Bolivia disjunction is probably not genuine <strong><strong>an</strong>d</strong><br />
more likely to be due to inadequate exploration<br />
in Peru.<br />
The discontinuities in the distribution of C.<br />
microcarpa, especially the isolated localities in<br />
Perambuco <strong><strong>an</strong>d</strong> Paraiba, could be related to the<br />
more or less isolated occurrence of forest <strong>with</strong><br />
rain forest elements on low mountain tops.<br />
sion of the diaspores is <strong>an</strong>other feature which the<br />
two genera have in common; in <strong>Coussapoa</strong> it is<br />
(probably) confined to C. asperifolia <strong><strong>an</strong>d</strong> in Poi-<br />
kilospermum it is only found in subg. Liguli-<br />
stigma (cf. Chew Wee-Lek, 1963). This phenom-<br />
enon, in combination <strong>with</strong> microspermy, is<br />
probably associated <strong>with</strong> the hemi-epiphytic<br />
habit. In <strong>Coussapoa</strong> the mucilaginous layer that<br />
swells <strong><strong>an</strong>d</strong> pushes the endocarp out of the<br />
peri<strong>an</strong>th is formed by the middle layer of the<br />
pericarp. In Poikilospermum, however, the ex-<br />
pulsion of the fruits is achieved by the swelling<br />
of the inner surface of the peri<strong>an</strong>th, which be-<br />
comes mucilaginous during the ripening of the<br />
fruit (cf. Chew Wee-Lek, 1963).<br />
Judging from (the nature of) the similarities<br />
In the present treatment, several pairs of closely <strong><strong>an</strong>d</strong> differences, the two genera are ecological<br />
related species are recognized, e.g., C. crassive- equivalents rather th<strong>an</strong> systematically closely renosa-C.<br />
ferruginea, C. latifolia-C. viridifolia. lated. Morphological as well as <strong>an</strong>atomical char-<br />
These pairs of species have allopatric distribu- acters (cf. Bonsen & ter Welle, 1983) suggest a<br />
tions.<br />
closer relationship of <strong>Coussapoa</strong> to the other five<br />
genera of the family, especially to the Neotrop-<br />
Systematic Position of the Genus ical ones. <strong>Coussapoa</strong> matches Cecropia in microspermy.<br />
The two genera show similarities in<br />
<strong>Coussapoa</strong>, together <strong>with</strong> the Neotropical gen- the flowers, but otherwise are quite different.<br />
era Cecropia <strong><strong>an</strong>d</strong> <strong>Pourouma</strong>, the Afric<strong>an</strong> genera More similarities are found between <strong>Coussapoa</strong><br />
Mus<strong>an</strong>ga <strong><strong>an</strong>d</strong> Myri<strong>an</strong>thus, <strong><strong>an</strong>d</strong> the Asiatic genus <strong><strong>an</strong>d</strong> <strong>Pourouma</strong>, although the latter genus is mac-<br />
Poikilospermum, constitute the family Cecropi- rospermous. The inflorescences of <strong>Coussapoa</strong> reaceae<br />
(Berg, 1978).<br />
semble the staminate ones of some <strong>Pourouma</strong><br />
Poikilospermum, geographically isolated from species, e.g., P. tomentosa <strong><strong>an</strong>d</strong> P. mollis; some<br />
the other genera, has more characters, especially <strong>Pourouma</strong> species have leaves strongly reminis<strong>an</strong>atomical<br />
ones, in common <strong>with</strong> the Urticaceae cent of those common in <strong>Coussapoa</strong> (see above).<br />
th<strong>an</strong> the other genera (cf. Bonsen & ter Welle,<br />
1983). The latter authors, indeed, suggest tr<strong>an</strong>s-<br />
Systematic<br />
ferring the<br />
Arr<strong>an</strong>gement<br />
genus back to the Urticaceae. Neverof<br />
the<br />
theless, considering the distribution of<br />
Species<br />
characters<br />
throughout the (complex of) families Cecropi- In the present revision the species are treated<br />
aceae, Urticaceae, <strong><strong>an</strong>d</strong> Moraceae, placing Poi- in alphabetical order, as it is not possible to prokilospermum<br />
in the <strong>Cecropiaceae</strong> appears to be pose a satisfactory systematic arr<strong>an</strong>gement.<br />
justified, although not beyond all doubt. For most characters clear discontinuities in<br />
Both <strong>Coussapoa</strong> <strong><strong>an</strong>d</strong> Poikilospermum are pre- their variation are lacking. Primitive <strong><strong>an</strong>d</strong> derived<br />
domin<strong>an</strong>tly hemi-epiphytic (cf. Chew Wee-Lek, state (as the extremes of a morphological series)<br />
1963). However, the two genera differ in habit! c<strong>an</strong> be inferred, like lamina broadly ovate-nar-<br />
<strong>Coussapoa</strong> species are shrubs or trees <strong>with</strong> straight rowly obovate, basal lateral veins br<strong>an</strong>ched-un<strong><strong>an</strong>d</strong><br />
stiff br<strong>an</strong>ches, while Poikilospermum species br<strong>an</strong>ched, lateral veins numerous-few, petiole<br />
are more or less li<strong>an</strong>a-like <strong>with</strong> long <strong><strong>an</strong>d</strong> weak long-short, stipules long-short, indumentum <strong>with</strong><br />
(scrambling) br<strong>an</strong>ches (cf. Chew Wee-Lek, 1963). all types of hairs <strong><strong>an</strong>d</strong> ? dense-less diverse <strong><strong>an</strong>d</strong><br />
The two genera resemble each other in the ? sparse, inflorescences repeatedly br<strong>an</strong>chedbr<strong>an</strong>ched<br />
inflorescences <strong>with</strong> terminal globose unbr<strong>an</strong>ched, interfloral bracts present-absent, <strong><strong>an</strong>d</strong><br />
heads, but the basic ramification of the inflores- stamens three-two. See Table I.<br />
cence appears to be different. Dichotomous ram- In some species (e.g., C. nymphaeifolia, C. vilification<br />
appears to be basic in Poikilospermum, losa, <strong><strong>an</strong>d</strong> C. tessm<strong>an</strong>nii) a predomin<strong>an</strong>ce of<br />
but is derived in <strong>Coussapoa</strong> (see above). Expul- "primitive" characters c<strong>an</strong> be found or a pre-
16 Flora Neotropica<br />
Table I<br />
Extremes in a morphological character series in<br />
<strong>Coussapoa</strong>, inferred to be primitive <strong><strong>an</strong>d</strong> derived<br />
respectively.<br />
Primitive states Derived states<br />
Lamina broadly ovate Lamina narrowly obovate<br />
Basal lateral veins Basal lateral veins unbr<strong>an</strong>ched<br />
br<strong>an</strong>ched<br />
Lateral veins numerous Lateral veins few<br />
Petiole long<br />
Petiole short<br />
Stipules long Stipules short<br />
Indument <strong>with</strong> all types Indument less diverse<br />
of hairs <strong><strong>an</strong>d</strong> ? dense <strong><strong>an</strong>d</strong> + sparse<br />
Inflorescences repeatedly Inflorescences unbr<strong>an</strong>ched<br />
br<strong>an</strong>ched<br />
Interfloral bracts present Interfloral bracts absent<br />
Stamens 3 Stamen 1<br />
number of stamens in arr<strong>an</strong>ging the species, since<br />
for about one-third of the species recognized material<br />
belonging to only one sex is known.<br />
Accept<strong>an</strong>ce of the number of stamens as the<br />
basis for subdividing the genus would imply the<br />
occurrence of parallel (or homologous) differentiation<br />
of m<strong>an</strong>y characters, e.g., in the differentiation<br />
of the leaf shape, dimensions, venation,<br />
indumentum, <strong><strong>an</strong>d</strong> length of the petiole, in three<br />
groups of species (judging from the comparison<br />
of characters in the taxa of which the number of<br />
stamens is known).<br />
Taxonomy<br />
1. <strong>Coussapoa</strong> Aublet, Hist. pl. Gui<strong>an</strong>e 2: 955.<br />
1775; Trecul, Ann. Sci. Nat. Bot. Ser. 3, 8: 92.<br />
1847; Miquel, Fl. bras. 4(1): 131. 1853; Macbride,<br />
Publ. Field Mus. Bot. 13(2.2): 295. 1937;<br />
Woodson & Schery, Ann. Missouri Bot. Gard.<br />
47: 168.<br />
domin<strong>an</strong>ce of"derived" characters in others<br />
1960; Berg, Fl. Suriname 5(1): 279.<br />
(e.g.,<br />
C. cinnamomifolia, C. trinervia, <strong><strong>an</strong>d</strong> C. viridi-<br />
1975; Burger, Fieldi<strong>an</strong>a Bot. 40: 131. 1977;<br />
Akkerm<strong>an</strong>s &<br />
folia), <strong>with</strong> the majority of the species some-<br />
Berg, Proc. Kon. Ned. Akad.<br />
Wetensch. Ser.<br />
where in between. A few species have<br />
C, 85(4): 441. 1982. Lectotype<br />
exceptional<br />
characters: C. trinervia has a truly terrestrial hab- species (Berg, Fl. Suriname 5(1): 279. 1975):<br />
it, C. parviceps has fused pistillate flowers; C.<br />
<strong>Coussapoa</strong> latifolia Aublet.<br />
contorta has domatium-like cavities in the axils Hemi-epiphytic (or epiphytic) or terrestrial<br />
of the lateral veins; C. asperifolia has abruptly trees or shrubs <strong>with</strong> aerial roots or stilt-roots.<br />
thickened twigs <strong>with</strong> wide cavities <strong><strong>an</strong>d</strong>, more- Leaves in spirals, entire, venation pinnate to triover,<br />
a thick mucilaginous mesocarp, <strong><strong>an</strong>d</strong> C. pur- nervate, margin entire to subcrenate, stipules<br />
pusii is (often?) deciduous.<br />
fused, fully amplexicaul, usually leaving oblique<br />
The difference in the number of stamens ap- scars. Inflorescences mostly in pairs in the leaf<br />
pears to be a promising basis for the arr<strong>an</strong>gement axils, br<strong>an</strong>ched or in pistillate ones often unof<br />
the species or the subdivision of the genus. br<strong>an</strong>ched, bracteate (mostly only <strong>with</strong> interfloral<br />
The signific<strong>an</strong>ce of these differences lies not only bracts) or ebracteate. Flowers in globose (to elin<br />
the absence of tr<strong>an</strong>sitions but also in the dis- lipsoid or clavate) terminal heads, free or pistiltribution<br />
r<strong>an</strong>ges: the predomin<strong>an</strong>ce of species <strong>with</strong> late ones sometimes connate, peri<strong>an</strong>th tubular,<br />
3-staminate flowers in Central America <strong><strong>an</strong>d</strong> the in staminate flowers (2-)3(-4)-lobed, in pistillate<br />
Pacific coastal region of Colombia <strong><strong>an</strong>d</strong> Ecuador; ones entire; stamens two or three <strong><strong>an</strong>d</strong> connate,<br />
the predomin<strong>an</strong>ce of species <strong>with</strong> 2-staminate or one; ovary free, stigma penicillate to subpelflowers<br />
in eastern South America, especially east- tate. Fruit a drupelet or almost <strong>an</strong> achene, enern<br />
Brazil; <strong><strong>an</strong>d</strong> the concentration of species <strong>with</strong> closed in the enlarged + fleshy peri<strong>an</strong>th; seed<br />
uni-staminate flowers in the Upper Amazon Ba- <strong>with</strong> endosperm, embryo straight <strong>with</strong> flat <strong><strong>an</strong>d</strong><br />
sin. It is not yet possible, however, to use the equal cotyledons <strong><strong>an</strong>d</strong> relatively short radicle.
<strong>Coussapoa</strong><br />
General key<br />
Keys to the species in:<br />
1. Central America <strong><strong>an</strong>d</strong> Mexico<br />
2. Colombia<br />
3. Venezuela<br />
4. the Gui<strong>an</strong>as2<br />
5. Ecuador<br />
Keys to the Species of <strong>Coussapoa</strong><br />
6. Peru <strong><strong>an</strong>d</strong> Bolivia " \ . 7 /<br />
7. Amazoni<strong>an</strong> Brazil<br />
8. Eastern Brazil<br />
General Key to the Species of <strong>Coussapoa</strong><br />
1. Interfloral bracts absent.<br />
2. Basal lateral veins br<strong>an</strong>ched.<br />
3. Lamina beneath <strong>with</strong> dense indumentum of minute hairs, at least in the areoles <strong><strong>an</strong>d</strong> on the<br />
reticulum.<br />
4. Indumentum (unicellular hairs only!) predomin<strong>an</strong>tly white; pistillate inflorescences unicapitate;<br />
stamens 3.<br />
5. Intercostal venation prominent beneath; Costa Rica. .............. 29. C. nymphaeifolia.<br />
5. Intercostal venation pl<strong>an</strong>e beneath; P<strong>an</strong>ama. ............................. 6. C. brevipes.<br />
4. Indumentum (unicellular hairs only!) at least partly brownish or yellow; pistillate inflorescences<br />
usually pluricapitate; stamen 1 or unknown; Amazon Basin or Colombia at ca. 1900 m.<br />
6. Lamina usually elliptic; lateral veins 5-9 pairs. ..................... See 12. C. cupularis.<br />
6. Lamina usually ovate; lateral veins 8-15 or 14-22 pairs.<br />
7. Lateral veins 8-15 pairs; peduncle of the pistillate inflorescence 1-2.5 mm thick.<br />
8. Intercostal venation (almost) pl<strong>an</strong>e; indumentum of petiole consisting of dense<br />
minute hairs <strong><strong>an</strong>d</strong> distinctly longer, yellowish hairs; Amazon Basin ..... 28. C. nitida.<br />
8. Intercostal venation prominent; indumentum of the petiole consisting of hairs not<br />
distinctly different in length; Colombia (ca. 1900 m). ............. 45. C. tolimensis.<br />
7. Lateral veins 14-22 pairs; peduncle of the pistillate inflorescence 6-10(-15) mm thick;<br />
Colombia (Amazonas). ......................................... 8. C. cinnamomea.<br />
3. Lamina beneath <strong>with</strong> sparse indumentum or glabrous, at least in the areoles <strong><strong>an</strong>d</strong> on the reticulum.<br />
9. Stipules 0.5-1 cm long; Venezuela (Bolivar <strong><strong>an</strong>d</strong> Amazonas), Guy<strong>an</strong>a, <strong><strong>an</strong>d</strong> Brazil (Amazonas,<br />
R oraim a). ........................................ .................. 43. C. viridifolia.<br />
9. Stipules at least 1 cm long.<br />
10. Lateral veins 3-6 pairs, the basal pair reaching the margin at or above the middle of the<br />
lamina; pistillate flowers cohere; stamens 3; Central America <strong><strong>an</strong>d</strong> Pacific coastal region<br />
of Colombia <strong><strong>an</strong>d</strong> Ecuador. ...........................................34. C. parviceps.<br />
10. Lateral veins 7 or more pairs, the basal pair reaching the margin below the middle of the<br />
lamina; if less th<strong>an</strong> 7 pairs of lateral veins then the pistillate flowers free <strong><strong>an</strong>d</strong> stamen 1.<br />
11. Lamina <strong><strong>an</strong>d</strong> leafy twigs glabrous; base of the lamina cordate; Colombia (Choco).<br />
.............................................................. 5. C. batavorum .<br />
2<br />
17
18 Flora Neotropica<br />
11. Lamina <strong><strong>an</strong>d</strong> leafy twigs at least sparsely puberulous; base of the lamina acute to<br />
truncate.<br />
12. Stipules (normally) 1-4 cm long.<br />
13. Margin of the lamina revolute at the base; Pacific coastal region (Colombia).<br />
......................................................... 46. C. valaria.<br />
13. Margin of the lamina not revolute at the base; Amazon Basin <strong><strong>an</strong>d</strong> Central<br />
America.<br />
14. Lamina coriaceous, mostly ovate; stamen 1; Amazon Basin (<strong><strong>an</strong>d</strong> P<strong>an</strong>ama?).<br />
............................................. 32. C. ovalifolia.<br />
14. Lamina subcoriaceous, elliptic to subobovate; stamens 2; Costa Rica.<br />
........................................ 23. C. macerrima.<br />
12. Stipules 4-13 cm long; Amazoni<strong>an</strong> Ecuador <strong><strong>an</strong>d</strong> Colombia. .... 27. C. napoensis.<br />
2. Basal lateral veins unbr<strong>an</strong>ched (or occasionally <strong>with</strong> a single br<strong>an</strong>ch).<br />
15. Stipules 0.5-1 cm long; stamens 2; common peduncle or br<strong>an</strong>ches of the pistillate inflorescence<br />
not distinctly broadened towards the flower heads; eastern Venezuela, Guy<strong>an</strong>a, <strong><strong>an</strong>d</strong> Brazil<br />
(Am azonas, Roraim a). ................................................... 43. C. viridifolia.<br />
15. Stipules (normally) at least 1 cm long, if shorter, then stamen 1 <strong><strong>an</strong>d</strong> br<strong>an</strong>ches of the pistillate<br />
inflorescence more or less distinctly broadened towards the flower heads.<br />
16. Margin of the lamina distinctly revolute towards the base; pistillate inflorescences mostly<br />
unicapitate; stamens 2; eastern Brazil. ................................ 25. C. microcarpa.<br />
16. Margin of the lamina pl<strong>an</strong>e, or if more of less revolute, then the pistillate inflorescences<br />
pluricapitate <strong><strong>an</strong>d</strong> stamen 1; Amazon Basin or Central Brazil.<br />
17. Lateral veins 2-8; basal pair of lateral veins reaching the margin above to just below<br />
the middle of the lamina <strong><strong>an</strong>d</strong>/or (often) the lamina beneath <strong>with</strong> dense, minute hairs<br />
in the areoles.<br />
18. Intercostal venation prominent beneath; lamina beneath <strong><strong>an</strong>d</strong> inflorescences <strong>with</strong><br />
dense, white, arachnoid indumentum; Brazil (Amapa) .......... 2. C. arachnoidea.<br />
18. Intercostal venation pl<strong>an</strong>e beneath; lamina <strong><strong>an</strong>d</strong> inflorescences <strong>with</strong>out arachnoid<br />
indumentum, or if present, then mostly sparse <strong><strong>an</strong>d</strong> soon disappearing; Upper<br />
Amazon Basin.<br />
19. Basal lateral veins reaching the margin above to just below the middle of the<br />
lamina; stipules mostly <strong>with</strong>out or <strong>with</strong> sparse, long, yellow hairs; br<strong>an</strong>ches<br />
of the pistillate inflorescence slightly broadened towards the flower heads;<br />
Upper Amazon Basin <strong><strong>an</strong>d</strong> Colombia (S<strong>an</strong>t<strong><strong>an</strong>d</strong>er). .......... 31. C. orthoneura.<br />
19. Basal lateral veins reaching the margin below to just above the middle of the<br />
lamina; stipules <strong>with</strong> dense, long, yellow hairs; br<strong>an</strong>ches of the pistillate inflorescence<br />
strongly broadened towards the flower heads; Brazil (R6ndonia).<br />
......................................................... 12. C. cupularis.<br />
17. Lateral veins (normally) 7-13 or 7-21 pairs, basal pair reaching the margin far below<br />
the middle of the lamina; lamina beneath glabrous or <strong>with</strong> sparse hairs in the areoles.<br />
20. Stipules (normally) 1-4 cm long.<br />
21. Lamina coriaceous, mostly ovate; stamen 1; Amazon Basin (<strong><strong>an</strong>d</strong> P<strong>an</strong>ama?).<br />
......................................................... 32. C. ovalifolia.<br />
21. Lamina subcoriaceous, elliptic to subobovate; stamens 2; Costa Rica. .....<br />
........................................ 23. C. macerrima.<br />
20. Stipules 4-13 cm long; Amazoni<strong>an</strong> Ecuador <strong><strong>an</strong>d</strong> Colombia. ...... 27. C. napoensis.<br />
1. Interfloral bracts present (sometimes small <strong><strong>an</strong>d</strong>/or few!).<br />
22. Lamina trinervate or <strong>with</strong> only the basal pair of lateral veins prominent <strong><strong>an</strong>d</strong> the other lateral veins<br />
weak <strong><strong>an</strong>d</strong> inconspicuous.<br />
23. Lamina trinervate; intercostal venation prominent beneath; Amazon Basin .... 40. C. trinervia.<br />
23. Lamina <strong>with</strong> only the basal pair of lateral veins prominent <strong><strong>an</strong>d</strong> the other lateral veins weak<br />
<strong><strong>an</strong>d</strong> inconspicuous; intercostal venation beneath; Bolivia (La Paz), Ecuador (Pastaza), <strong><strong>an</strong>d</strong><br />
Colombia (Antioquia). ..............................................9. C. cinnamomifolia.<br />
22. Lamina pinnately veined.<br />
24. Lamina above <strong>with</strong> white arachnoid hairs, at least (sub)persistent on the midrib; eastern Brazil.<br />
25. Stipules ca. 0.5 cm long ............................................... 13. C. curr<strong>an</strong>ii.<br />
25. Stipules 1-2.5 cm long ................................................ 17. C. floccosa.<br />
24. Lamina above glabrous (sometimes except for the base of the midrib) or scabrous to scabridulous<br />
<strong>with</strong> short, rigid hairs; not in eastern Brazil.<br />
26. Lamina scabrous to scabridulous above.<br />
27. Stipules ca. 6 cm long; lamina beneath <strong>with</strong> arachnoid hairs; basal lateral veins br<strong>an</strong>ched;<br />
Peru (Loreto) <strong><strong>an</strong>d</strong> Ecuador (Napo). ........................ 22. C. longepedunculata.<br />
27. Stipules up to 3.5 cm long, mostly less th<strong>an</strong> 1 cm long; lamina beneath lacking<br />
arachnoid hairs, or if present then the basal lateral veins unbr<strong>an</strong>ched.
<strong>Coussapoa</strong> 19<br />
28. Leafy twigs (rather) abruptly thickened <strong><strong>an</strong>d</strong> distinctly hollow; widespread ....<br />
............................................................ 4. C asp erifolia.<br />
28. Leafy twigs gradually thickened <strong><strong>an</strong>d</strong> solid.<br />
29. Lateral veins 6-13 pairs; lamina subcoriaceous; southern Amazon Basin. ...<br />
............................... ................ ........... 37. C. scabra.<br />
29. Lateral veins 6-7 pairs; lamina coriaceous; French Gui<strong>an</strong>a <strong><strong>an</strong>d</strong> Brazil (Para).<br />
........................................................ 21. C. leprieurii.<br />
26. Lamina smooth <strong><strong>an</strong>d</strong> glabrous above.<br />
30. Basal lateral veins unbr<strong>an</strong>ched (or sometimes <strong>with</strong> 1 or 2 br<strong>an</strong>ches).<br />
31. Lamina beneath <strong>with</strong> domatium-like cavities in the axils of the lateral veins;<br />
interfloral bracts large <strong><strong>an</strong>d</strong> peltate; western Ecuador <strong><strong>an</strong>d</strong> Colombia. ..........<br />
....................................... ...................... 10 . C . contorta.<br />
31. Lamina beneath <strong>with</strong>out domatium-like cavities; interfloral bracts relatively small<br />
<strong><strong>an</strong>d</strong> spathulate to subpeltate.<br />
32. Stipules 0.5-1 cm long.<br />
33. Lateral veins 2-3(-4) pairs.<br />
34. Apex of the lamina acute; Central America. ...... 18. C. glaberrima.<br />
34. Apex of the lamina rounded to obtuse; Surinam, French Gui<strong>an</strong>a,<br />
<strong><strong>an</strong>d</strong> Brazil (Amapa <strong><strong>an</strong>d</strong> Para). .................... 1. C. <strong>an</strong>gustifolia.<br />
33. Lateral veins at least 4 pairs.<br />
35. Young leaves <strong>with</strong> arachnoid indumentum; lamina obovate to subobovate;<br />
lateral veins 7-10 pairs; terminal buds slender; eastern<br />
Brazil. .................... ..................... 13. C. curr<strong>an</strong>ii.<br />
35. Young leaves <strong>with</strong>out arachnoid indumentum; lamina elliptic, or if<br />
(sub)obovate, then lateral veins 4-7 pairs <strong><strong>an</strong>d</strong>/or terminal buds<br />
swollen; not in eastern Brazil.<br />
36. Terminal buds swollen; Upper Amazon Basin <strong><strong>an</strong>d</strong> Colombia.<br />
........................................ 35. C. parvifolia.<br />
36. Terminal buds slender.<br />
37. Lateral veins 4-7 pairs.<br />
38. Hairs on the main veins beneath of equal length; Lower<br />
Amazon Basin, Surinam, <strong><strong>an</strong>d</strong> French Gui<strong>an</strong>a. ...<br />
.............. ...................... 20. C. latifolia.<br />
38. Hairs on the main veins beneath of different lengths;<br />
eastern Venezuela, Guy<strong>an</strong>a, <strong><strong>an</strong>d</strong> Brazil (Amazonas,<br />
Roraima). ......................... 43. C. viridifolia.<br />
37. Lateral veins 7-11 pairs; Upper Amazon Basin <strong><strong>an</strong>d</strong> Colombia.<br />
................................ 35. C. parvifolia.<br />
32. Stipules at least 1 cm long.<br />
39. Lateral veins 2-3(-4) pairs.<br />
40. Lamina glabrous; Central America. .............. 18. C. glaberrima.<br />
40. Lamina beneath <strong>with</strong> (partly arachnoid) indumentum; Venezuela.<br />
........................................ 3. C. argentea.<br />
39. Lateral veins at least 4 pairs.<br />
41. Lamina beneath <strong>with</strong> dense indumentum.<br />
42. Arachnoid hairs present on the lamina beneath, at least in young<br />
leaves.<br />
43. Arachnoid hairs brownish.<br />
44. Lateral veins 10-14 pairs; pistillate inflorescences unicapitate,<br />
head ellipsoid to clavate; staminate flower<br />
heads 4-8 mm in diameter; Ande<strong>an</strong> Colombia <strong><strong>an</strong>d</strong><br />
Ecuador (1400-2000 m). ............... 14. C. duquei.<br />
44. Lateral veins at most 10(-11); pistillate inflorescences<br />
pluricapitate, or if unicapitate then the heads globose;<br />
staminate flower heads 1-3 mm in diameter.<br />
45. Lateral veins 4-5 pairs; lamina oblong; French<br />
Gui<strong>an</strong>a <strong><strong>an</strong>d</strong> Brazil (Amapa). ..... 16. C. ferruginea.<br />
45. Lateral veins (4-)5-8(-11) pairs; lamina mostly<br />
elliptic to ovate.<br />
46. Hairs on the smaller veins beneath appressed;<br />
Gui<strong>an</strong>as ................ 26. C. microcephala.<br />
46. Hairs on the smaller veins beneath (at least<br />
partly) patent.<br />
47. Stamens slightly longer th<strong>an</strong> the peri-
20 Flora Neotropica<br />
<strong>an</strong>th; pistillate flower heads 1-3; eastern<br />
Brazil. ............... 33. C. pachyphylla.<br />
47. Stamens distinctly longer th<strong>an</strong> the peri<strong>an</strong>th;<br />
pistillate flower heads 3-7; Venezuela<br />
(Bolivar), Brazil (Roraima), Amazoni<strong>an</strong><br />
Bolivia <strong><strong>an</strong>d</strong> Ecuador, <strong><strong>an</strong>d</strong><br />
P<strong>an</strong>ama. ............ 11. C. crassivenosa.<br />
43. Arachnoid hairs whitish.<br />
48. Lamina mostly ovate; lateral veins 9-16 pairs; stamen<br />
1; peduncle of the pistillate inflorescence 2-4 mm thick;<br />
Amazon Basin. .................... 39. C. tessm<strong>an</strong>nii.<br />
48. Lamina mostly elliptic to oblong; lateral veins 5-12<br />
pairs; stamens 3; peduncle of the pistillate inflorescence<br />
ca. 1 mm thick.<br />
49. Stipules 0.5-1.5(-2) cm long; heads of staminate<br />
inflorescence m<strong>an</strong>y, 3-4 mm in diameter; heads<br />
of pistillate inflorescence 2-5; peri<strong>an</strong>th glabrous;<br />
Amazon Basin. .................... 38. C. sprucei.<br />
49. Stipules 1-5 cm long; heads of staminate inflorescence<br />
3-10, 4-6 mm in diameter; heads of pistillate<br />
inflorescence 1(-3); peri<strong>an</strong>th minutely puberulous;<br />
Central America to Mexico..........<br />
............................. 30. C. oligocephala.<br />
42. Arachnoid hairs lacking on the lamina beneath, or if present<br />
then sparse <strong><strong>an</strong>d</strong> (soon) disappearing.<br />
50. Lateral veins 5-8 pairs; eastern Brazil. .. 33. C. pachyphylla.<br />
50. Lateral veins 8-12 pairs; not in eastern Brazil.<br />
51. Margin of the lamina revolute at the base; Upper Amazon<br />
Basin <strong><strong>an</strong>d</strong> Colombia. ........... 35. C. parvifolia.<br />
51. Margin of the lamina not revolute at the base.<br />
52. Petiole 1-3 cm long; stipules 1.5-2.5 cm long;<br />
pistillate inflorescences unicapitate, the peduncle<br />
0.5-1 cm long; Colombia (Choc6). 7. C. chocoensis.<br />
52. Petiole 3-9 cm long; stipules 2-6 or 3-20 cm long;<br />
pistillate inflorescences pluricapitate, or if unicapitate<br />
then the peduncle at least 1 cm long.<br />
53. Heads of pistillate inflorescence 6-10; stamens<br />
3; Pacific coastal region of Ecuador <strong><strong>an</strong>d</strong><br />
Colombia. ................... 19. C. herthae.<br />
53. Heads of pistillate inflorescence 1(-2); stamens<br />
2 or unknown.<br />
54. Lamina broadly elliptic to obovate, apex<br />
shortly acuminate; pistillate flower heads<br />
ca. 4 mm, in fruit up to 12 mm in diameter;<br />
Amazoni<strong>an</strong> Peru ...........<br />
...................... 24. C. m<strong>an</strong>uensis.<br />
54. Lamina mostly ovate to subovate, apex<br />
acute to obtuse; pistillate flower heads<br />
ca. 5-10 mm, in fruit up to 40 mm in<br />
diameter; widespread....... 42. C. villosa.<br />
41. Lamina beneath <strong>with</strong> sparse indumentum.<br />
53. Margin of the lamina distinctly revolute towards the base.<br />
56. Stipules 1-7 cm long; terminal buds slender; eastern Brazil<br />
...................................... 25. C. m icrocarpa.<br />
56. Stipules up to 1.3 cm long; terminal buds usually swollen;<br />
Upper Amazon Basin <strong><strong>an</strong>d</strong> Colombia. ..... 35. C. parvifolia.<br />
55. Margin of the lamina pl<strong>an</strong>e or entirely revolute.<br />
57. Leafy twigs (rather) abruptly thickened <strong><strong>an</strong>d</strong> <strong>with</strong> wide cavities;<br />
stamen 1; widespread ............... 4. C. asperifolia.<br />
57. Leafy twigs gradually thickened <strong><strong>an</strong>d</strong> solid or <strong>with</strong> narrow<br />
cavities.<br />
58. Stipules glabrous or puberulous.<br />
59. Basal lateral veins reaching the margin above the
<strong>Coussapoa</strong> 21<br />
middle of the lamina; base of the lamina acute;<br />
pistillate inflorescences pluricapitate; P<strong>an</strong>ama. ..<br />
............................... 15. C. echinata.<br />
59. Basal lateral veins mostly reaching the margin<br />
below the middle of the lamina; base of the lamina<br />
mostly obtuse to (sub)cordate.<br />
60. Stamens 3; pistillate inflorescences unicapitate;<br />
northern Central America to Mexico.<br />
............................. 36. C. purpusii.<br />
60. Stamens 2; pistillate inflorescences pluricapitate;<br />
South America.<br />
61. Hairs on the main veins beneath equal<br />
in length <strong><strong>an</strong>d</strong> arachnoid hairs lacking;<br />
Surinam, French Gui<strong>an</strong>a, <strong><strong>an</strong>d</strong> Brazil<br />
(Amapa, Para, <strong><strong>an</strong>d</strong> Amazonas). ......<br />
......................... 20. C. latifolia.<br />
61. Hairs on the main veins beneath of different<br />
lengths <strong><strong>an</strong>d</strong>/or on the main veins<br />
<strong><strong>an</strong>d</strong> margin subpersistent arachnoid<br />
hairs; Gui<strong>an</strong>as....... 26. C. microcephala.<br />
58. Stipules (sub)sericeous, (sub)hirsute, (sub)villous, etc.<br />
62. Stamens 3; pistillate flower heads 6-10, 5-9 mm<br />
in diameter; Pacific coastal region of Ecuador <strong><strong>an</strong>d</strong><br />
Colombia.<br />
63. Petiole 5-10 cm long, or if shorter, then stamens<br />
3, pistillate flower heads 6-10, 5-9 mm<br />
in diameter; Pacific coastal region of Colombia<br />
<strong><strong>an</strong>d</strong> Ecuador.<br />
64. Stipules 2-6 cm long. ..... 19. C. herthae.<br />
64. Stipules 1.5-2 cm long. .. 44. C. fulvescens.<br />
63. Petiole up to 5 cm long; stamens 2; flower<br />
heads 1-5; not in Pacific coastal region.<br />
62. Stamens 2; pistillate flower heads 1-3, or if more<br />
th<strong>an</strong> 3 then 2-5 mm in diameter.<br />
64. Stamens slightly longerth<strong>an</strong> the peri<strong>an</strong>th; pistillate<br />
flower heads 1-3; eastern Brazil ......<br />
........................ 33. C. pachyphylla.<br />
64. Stamens distinctly longer th<strong>an</strong> the peri<strong>an</strong>th;<br />
pistillate flower heads (1-)3-15; Gui<strong>an</strong>as. ..<br />
........................ 26. C. microcephala.<br />
30. Basal lateral veins br<strong>an</strong>ched.<br />
65. Lamina beneath <strong>with</strong> (rather) dense, (rather) conspicuous <strong><strong>an</strong>d</strong> (sub)persistent<br />
arachnoid indumentum.<br />
66. Arachnoid hairs on the lamina brownish.<br />
67. Lateral veins 4-8(-11) pairs; pistillate inflorescences mostly pluricapitate;<br />
staminate flower heads ca. 2-3 mm in diameter.<br />
68. Hairs on the smaller veins beneath (at least partly) patent.<br />
69. Stamens slightly longer th<strong>an</strong> the peri<strong>an</strong>th; pistillate flower heads<br />
1-3; eastern Brazil. ....................... 33. C. pachyphylla.<br />
69. Stamens distinctly longer th<strong>an</strong> the peri<strong>an</strong>th; pistillate flower<br />
heads 3-7; Venezuela (Bolivar), Brazil (Roraima), Amazoni<strong>an</strong><br />
Bolivia <strong><strong>an</strong>d</strong> Ecuador, <strong><strong>an</strong>d</strong> P<strong>an</strong>amf. ......... 11. C. crassivenosa.<br />
68. Hairs on the smaller veins beneath appressed; Gui<strong>an</strong>as..........<br />
............................................ 26. C. m icrocephala.<br />
67. Lateral veins (7-)10-24 pairs; pistillate inflorescences usually unicapitate;<br />
staminate flower heads 4-10 mm in diameter.<br />
70. Common peduncle of the staminate inflorescence 1-2 cm long, that<br />
of the pistillate inflorescence 0.5-0.8 cm long; pistillate heads ellipsoid<br />
to clavate; Ande<strong>an</strong> Ecuador <strong><strong>an</strong>d</strong> Colombia (1400-2000 m). ..<br />
.................................................. 14. C. duquei.<br />
70. Common peduncle of the staminate inflorescence 2-6 cm long, that<br />
of the pistillate inflorescence (1-)2-13 cm long; pistillate flower<br />
heads (sub)globose; widespread. ..................... 42. C. villosa.
22 Flora Neotropica<br />
66. Arachnoid hairs on the lamina whitish.<br />
71. Lateral veins ca. 10 or 9-24 pairs; lamina mostly longer th<strong>an</strong> 15 cm;<br />
stipules mostly longer th<strong>an</strong> 2 cm.<br />
72. Lamina beneath <strong>with</strong> dense brown hairs on the main veins; common<br />
peduncle of the staminate inflorescence ca. 9 cm long; Amazoni<strong>an</strong><br />
Peru <strong><strong>an</strong>d</strong> Ecuador. ....................... 22. C. longepedunculata.<br />
72. Lamina beneath <strong>with</strong> whitish hairs on the main veins; peduncle of<br />
the staminate inflorescence up to 6 cm long.<br />
73. Stamen 1; pistillate inflorescences pluricapitate <strong><strong>an</strong>d</strong> common<br />
peduncle 2-3 cm long; Amazon Basin. ....... 39. C. tessm<strong>an</strong>nii.<br />
73. Stamens 2 or 3; pistillate inflorescences unicapitate, or if 2-3capitate<br />
then the common peduncle longer th<strong>an</strong> 3 cm.<br />
74. Stamens 3; pistillate flower heads ellipsoid to clavate; peduncle<br />
of the pistillate inflorescence 1-2 cm long; Central<br />
America. .......................... 29. C. nymphaeifolia.<br />
74. Stamens 2; peduncle of the pistillate inflorescence (1-)2-<br />
13 cm long; pistillate flower heads (sub)globose; widespread.<br />
................................... 42. C. villosa.<br />
71. Lateral veins 4-10(-11) pairs; lamina mostly shorter th<strong>an</strong> 15 cm.<br />
75. Hairs on the smaller veins beneath appressed; Gui<strong>an</strong>as..........<br />
............................................ 26. C. m icrocephala.<br />
75. Hairs on the smaller veins beneath (at least partly) patent.<br />
76. Stipules up to 2 cm long.<br />
77. Hairs on the stipules predomin<strong>an</strong>tly weak <strong><strong>an</strong>d</strong> crinkled;<br />
stamens 2; pistillate flower heads 1-3; eastern Brazil. ...<br />
..................................... 33. C. pachyphylla.<br />
77. Hairs on the stipules predomin<strong>an</strong>tly stiff <strong><strong>an</strong>d</strong> straight; stamens<br />
3; pistillate flower heads 2-6; Amazoni<strong>an</strong> Brazil. ..<br />
.......................................... 34. C. sprucei.<br />
76. Stipules 3-20 cm long; widespread. .............. 42. C. villosa.<br />
65. Lamina beneath <strong>with</strong>out arachnoid indumentum, or if present then sparse <strong><strong>an</strong>d</strong><br />
soon disappearing.<br />
78. Leafy twigs (rather) abruptly thickened <strong><strong>an</strong>d</strong> <strong>with</strong> wide cavities; stamen 1;<br />
widespread. ............................................. 4. C. asperifolia.<br />
78. Leafy twigs gradually thickened, solid or <strong>with</strong> narrow cavities; stamens 2<br />
or 3.<br />
79. Lateral veins 4-10(-11) pairs.<br />
80. Lamina almost as long as broad; pair of main basal veins reaching<br />
the margin far above the middle of the lamina; most lateral veins<br />
br<strong>an</strong>ched; Pacific coastal region of Ecuador <strong><strong>an</strong>d</strong> Colombia. ......<br />
............................................... 41. C. v<strong>an</strong>nifolia.<br />
80. Lamina mostly distinctly longer th<strong>an</strong> broad, or if almost as long as<br />
broad then only the basal pair of lateral veins br<strong>an</strong>ched.<br />
81. Hairs on the smaller veins beneath appressed; Gui<strong>an</strong>as. .....<br />
........................................ 26. C. m icrocephala.<br />
81. Hairs on the smaller veins beneath at least partly patent.<br />
82. Stamen 1; pistillate inflorescences pluricapitate <strong><strong>an</strong>d</strong> the<br />
common peduncle 2-3 cm long <strong><strong>an</strong>d</strong> 2-4 cm thick, peri<strong>an</strong>th<br />
of the pistillate flower (almost) glabrous; Amazon Basin.<br />
...................................... 39. C. tessm <strong>an</strong>nii.<br />
82. Stamens 2 or 3; pistillate inflorescences unicapitate, or if<br />
pluricapitate then the common peduncle longer th<strong>an</strong> 3 cm,<br />
less th<strong>an</strong> 2 mm thick; <strong><strong>an</strong>d</strong>/or the peri<strong>an</strong>th of the pistillate<br />
flower densely puberulous.<br />
83. Stamens 3; pistillate inflorescence <strong>with</strong> 6-10 heads, 5-<br />
9 (in fruit up to 15) mm in diameter <strong><strong>an</strong>d</strong> the common<br />
peduncle 3-6 mm long; Pacific coastal region of Ecuador<br />
<strong><strong>an</strong>d</strong> Colombia. ................. 19. C. herthae.<br />
83. Stamens 2; pistillate inflorescences <strong>with</strong> 1-3 heads, or<br />
if more then the heads 3-5 mm in diameter <strong><strong>an</strong>d</strong> the<br />
common peduncle 1-3 cm long.<br />
84. Stamens slightly longer th<strong>an</strong> the peri<strong>an</strong>th; pistil-
<strong>Coussapoa</strong> 23<br />
late flower heads 1-3; peri<strong>an</strong>th of the pistillate<br />
flower glabrous; eastern Brazil. .. 33. C. pachyphylla.<br />
84. Stamens distinctly longer th<strong>an</strong> the peri<strong>an</strong>th; pistillate<br />
flower heads 3-7, or if less then the peri<strong>an</strong>th<br />
of the pistillate flower densely puberulous.<br />
85. Staminate flower heads 2-3 mm in diameter;<br />
pistillate flower heads 3-7, <strong><strong>an</strong>d</strong> 2-5 mm in<br />
diameter; Venezuela (Bolivar), Brazil (Roraima),<br />
Amazoni<strong>an</strong> Bolivia <strong><strong>an</strong>d</strong> Ecuador, <strong><strong>an</strong>d</strong><br />
P<strong>an</strong>ama. ................. 1. C. crassivenosa.<br />
85. Staminate flower heads 5-10 mm in diameter;<br />
pistillate flower heads 1(-3) <strong><strong>an</strong>d</strong> 5-10 mm<br />
in diameter; widespread. ....... 42. C. villosa.<br />
79. Lateral veins at least 10 pairs.<br />
86. Stamen 1; pistillate inflorescences pluricapitate <strong><strong>an</strong>d</strong> the common<br />
peduncle 2-3 cm long <strong><strong>an</strong>d</strong> 2-4 cm thick; peri<strong>an</strong>th of the pistillate<br />
flower (almost) glabrous; Amazon Basin. ......... 39. C. tessm<strong>an</strong>nii.<br />
86. Stamens 2 or 3; pistillate inflorescences unicapitate, or if pluricapitate<br />
then the common peduncle longer th<strong>an</strong> 3 cm, less th<strong>an</strong> 2 mm<br />
thick, <strong><strong>an</strong>d</strong>/or the peri<strong>an</strong>th of the pistillate flower densely puberulous.<br />
87. Stamens 3; pistillate flower heads 6-10; Pacific coastal region<br />
of Ecuador <strong><strong>an</strong>d</strong> Colombia. ..................... 19. C. herthae.<br />
87. Stamens 2; pistillate flower heads 1(-3); widespread .........<br />
........................................ 42. C. villosa.<br />
1. Key to the <strong>Coussapoa</strong> Species in Central America <strong><strong>an</strong>d</strong> Mexico<br />
1. Interfloral bracts absent.<br />
2. Lamina beneath glabrous or <strong>with</strong> sparse indumentum.<br />
3. Lateral veins 3-6 pairs, the basal pair reaching the margin above or at the middle of the lamina.<br />
.. ............... ....................................................... 34. C. parviceps.<br />
3. Lateral veins 7-13 pairs, the basal pair reaching the margin below the middle of the lamina.<br />
4. Stipules 1-2 cm long; lamina 5-13 x 4-7 cm. ........................... 23. C. macerrima.<br />
4. Stipules 2-3 cm long; lamina 13-20 x 9-13 cm. ................. See under 32. C. ovalifolia.<br />
2. Lamina beneath <strong>with</strong> dense indumentum.<br />
5. Intercostal venation prominent beneath; stipules <strong>with</strong> brownish indumentum (unicellular hairs<br />
only). ........................................................ .. .... 39. C. nym phaeifolia.<br />
5. Intercostal venation pl<strong>an</strong>e beneath; stipules <strong>with</strong> whitish indumentum (unicellular hairs only!).<br />
........................................................................... 6. C brevipes.<br />
1. Interfloral bracts present.<br />
6. Lamina beneath <strong>with</strong> dense indumentum, partly arachnoid indumentum.<br />
7. Arachnoid indumentum on the lamina brownish. .......................... 11. C. crassivenosa.<br />
7. Arachnoid indumentum on the lamina whitish.<br />
8. Basal pair of lateral veins unbr<strong>an</strong>ched. ................................. 30. C. oligocephala.<br />
8. Basal pair of lateral veins br<strong>an</strong>ched.<br />
9. Stamens 3; (common) peduncle of the pistillate inflorescence 1-2 cm long, pistillate flower<br />
heads ellipsoid to obovoid. ...................................... 39. C. nymphaeifolia.<br />
9. Stamens 2; (common) peduncle of the pistillate inflorescence (1-)2-13 cm long, pistillate<br />
flower heads (sub)globose. ............................................. 42. C. villosa.<br />
6. Lamina beneath glabrous or <strong>with</strong> sparse indumentum, arachnoid indumentum lacking.<br />
10. All or most lateral veins br<strong>an</strong>ched; leafy twigs more or less abruptly thickened, <strong>with</strong> wide cavities. C<br />
......... . .....................................................<br />
asperifolia.<br />
10. All or most lateral veins unbr<strong>an</strong>ched; leafy twigs gradually thickened, solid or <strong>with</strong> narrow<br />
cavities.<br />
11. Lateral veins 2-3 pairs. ............................................ 18. C. glaberrima.<br />
11. Lateral veins 4-6 pairs.<br />
12. Basal pair of lateral veins unbr<strong>an</strong>ched; pistillate inflorescences unicapitate. .........<br />
...................................... ............................ 36. C. purpusii.<br />
12. Basal pair of lateral veins (faintly) br<strong>an</strong>ched; pistillate inflorescences pluricapitate. ..<br />
................................................................. 15. C. echinata.
24 Flora Neotropica<br />
2. Key to the <strong>Coussapoa</strong> Species in Colombia<br />
1. Interfloral bracts absent.<br />
2. Lateral veins 2-8 pairs, basal pair reaching the margin above (or at) the middle of the lamina.<br />
3. Basal pair of lateral veins unbr<strong>an</strong>ched; stamen 1; pistillate flowers free. ........ 31. C. orthoneura.<br />
3. Basal pair of lateral veins br<strong>an</strong>ched; stamens 3; pistillate flowers cohere. .........34. C. parviceps.<br />
2. Lateral veins at least 8 pairs, or if less then basal pair reaching the margin below the middle of the<br />
lamina.<br />
4. Lamina in the areoles beneath glabrous or <strong>with</strong> sparse hairs.<br />
5. Stipules 2-3 cm long . .................................................... 46. C. valaria.<br />
5. Stipules 4-13 cm long.<br />
6. Base of the lamina cordate; basal pair of lateral veins reaching the margin above or at the<br />
middle of the lamina . .............................................. 5. C. batavorum.<br />
6. Base of the lamina (sub)obtuse or rounded to truncate; basal pair of lateral veins reaching<br />
the margin below the middle of the lamina ........................... 27. C. napoensis.<br />
4. Lamina in the areoles beneath <strong>with</strong> dense, minute hairs.<br />
7. Indumentum of the petiole consisting of dense, minute hairs <strong><strong>an</strong>d</strong> distinctly longer, yellowish<br />
hairs . .................................................................... C. 28. nitida.<br />
7. Indumentum of the petiole consisting of short hairs, not distinctly different in length.<br />
8. Indumentum of the leafy twigs <strong><strong>an</strong>d</strong> stipules (unicellular hairs only) brown(ish); petiole 5-<br />
7 m m thick. ..................................................... 8. C. cinnam omea.<br />
8. Indumentum of the leafy twigs <strong><strong>an</strong>d</strong> stipules (unicellular hairs only) white or yellowish;<br />
petiole 2-4 mm thick.<br />
9. Intercostal venation almost pl<strong>an</strong>e; pistillate inflorescences unbr<strong>an</strong>ched ... 6. C. brevipes.<br />
9. Intercostal venation prominent; pistillate inflorescences br<strong>an</strong>ched .... 45. C. tolimensis.<br />
1. Interfloral bracts present.<br />
10. Lamina trinervate ........................................................... 40. C. trinervia.<br />
10. Lamina pinnately veined.<br />
11. Lamina beneath <strong>with</strong> arachnoid indumentum.<br />
12. Arachnoid indumentum on the lamina beneath brown(ish).<br />
13. Lateral veins 4-8(-11) pairs; pistillate inflorescences pluricapitate; staminate flower<br />
heads ca. 2-3 mm in diameter ................... .............. 11. C. crassivenosa.<br />
13. Lateral veins (7-)10-24 pairs; pistillate inflorescences (usually) unicapitate; staminate<br />
flower heads 4-10 mm in diameter.<br />
14. Common peduncle of the staminate inflorescence 1-2 cm long, that of the pistillate<br />
inflorescence 0.5-0.8 cm long; pistillate flower heads ellipsoid to clavate. .....<br />
........................... ................................... 14. C . duquei.<br />
14. Common peduncle of the staminate inflorescence 2-6 cm long, that of the pistillate<br />
inflorescence (1-)2-13 cm long; pistillate flower heads (sub)globose. .. 42. C. villosa.<br />
12. Arachnoid indumentum on the lamina beneath whitish.<br />
15. Stamen 1; pistillate inflorescence pluricapitate <strong><strong>an</strong>d</strong> the common peduncle 2-3 cm long,<br />
2-4 mm thick; peri<strong>an</strong>th of the pistillate flower almost glabrous ...... 39. C. tessm<strong>an</strong>nii.<br />
15. Stamens 2 or 3; pistillate inflorescence unicapitate, or if pluricapitate then the common<br />
peduncle longer th<strong>an</strong> 3 cm, less th<strong>an</strong> 2 mm thick, <strong><strong>an</strong>d</strong>/or the peri<strong>an</strong>th of the pistillate<br />
flower densely puberulous.<br />
16. Stamens 3; pistillate flower heads 6-10. ......................... 19. C. herthae.<br />
16. Stamens 2; pistillate flower heads 1(-3). ........................ 42. C. villosa.<br />
11. Lamina beneath <strong>with</strong>out arachnoid indumentum.<br />
17. Lamina beneath <strong>with</strong> domatium-like cavities in the axils of the lateral veins; interfloral<br />
bracts large <strong><strong>an</strong>d</strong> peltate .................................. ............. 10. C. contorta.<br />
17. Lamina beneath <strong>with</strong>out domatium-like cavities; interfloral bracts relatively small <strong><strong>an</strong>d</strong><br />
spathulate to subpeltate.<br />
18. Stipules glabrous or sparsely puberulous ............................ 34. C. parviceps.<br />
18. Stipules <strong>with</strong> + dense indumentum.<br />
19. Basal lateral veins distinctly br<strong>an</strong>ched.<br />
20. Lamina about as long as broad; lateral veins 5-9 pairs. ..... 41. C. v<strong>an</strong>nifolia.<br />
20. Lamina distinctly longer th<strong>an</strong> broad; lateral veins 8-12 pairs.... 19. C. herthae.<br />
19. Basal lateral veins unbr<strong>an</strong>ched (or sometimes <strong>with</strong> 1-2 br<strong>an</strong>ches).<br />
21. Lateral veins 5-6 pairs; intercostal venation inconspicuous .............<br />
................................................... 9. C. cinnam om ifolia.<br />
21. Lateral veins 7-12 pairs; intercostal venation conspicuous.<br />
22. Stipules 0.3-1.3 cm long. ............................ 35. C. parvifolia.<br />
22. Stipules 1.5-2.5 cm long.
<strong>Coussapoa</strong><br />
23. Lateral veins 10-12 pairs; petiole 1-3 cm long; pistillate inflores-<br />
cences unbr<strong>an</strong>ched. .............................. 7. C. chocoensis.<br />
23. Lateral veins 5-10 pairs; petiole 5-10 cm long; pistillate inflores-<br />
cences br<strong>an</strong>ched. ............................... 44. C. fulvescens.<br />
3. Key to the <strong>Coussapoa</strong> Species in Venezuela<br />
1. Interfloral bracts absent.<br />
2. Stipules 0.5-1 cm long; stamens 2; pistillate flower heads 1-3. ....................<br />
2. Stipules (0.5-)1-7 cm long; stamen 1; pistillate flower heads 2-10. ...............<br />
1. Interfloral bracts present.<br />
3. Lamina trinervate .............................................................<br />
43. C. viridifolia.<br />
31. C. orthoneura.<br />
40. C. trinervia.<br />
3. Lamina pinnately veined.<br />
4. Lamina beneath <strong>with</strong> arachnoid indumentum.<br />
5. Arachnoid indumentum on the lamina brownish . ......................<br />
5. Arachnoid indumentum on the lamina whitish.<br />
6. Lateral veins 2-3(-4) pairs ........ .....................................<br />
6. Lateral veins 4-24 pairs . .................. ...........................<br />
4. Lamina beneath <strong>with</strong>out arachnoid indumentum.<br />
11. C. crassivenosa.<br />
3. C. argentea.<br />
42. C. villosa.<br />
7. Leafy twigs (rather) abruptly thickened <strong>with</strong> wide cavities; stamen 1 ......... 4. C. asperifolia.<br />
7. Leafy twigs gradually thickened solid or <strong>with</strong> narrow cavities.<br />
8. Lamina <strong>with</strong> patent hairs on the venation beneath; stipules 1-5 cm long. .............<br />
....................................................11. C. crassivenosa.<br />
8. Lamina <strong>with</strong> appressed hairs on the venation beneath; stipules 0.3-1(-1.3) cm long.<br />
9. Lateral veins 7-11 pairs; terminal buds usually swollen; margin of the lamina ? revolute<br />
at the base. ....................................... .... 35. C. parvifolia.<br />
9. Lateral veins 4-7 pairs; terminal buds slender; margin of the lamina not revolute at<br />
the base ........................................................ 43. C. viridifolia.<br />
4. Key to the <strong>Coussapoa</strong> Species in Guy<strong>an</strong>a, Surinam, <strong><strong>an</strong>d</strong> French Gui<strong>an</strong>a<br />
1. Lamina above scabrous.<br />
2. Basal lateral veins reaching the margin above the middle of the lamina. ............4. C. asperifolia.<br />
2. Basal lateral veins reaching the margin below the middle of the lamina ............. 21. C. leprieurii.<br />
1. Lamina above smooth.<br />
3. Lamina beneath <strong>with</strong> arachnoid indumentum.<br />
4. Arachnoid indumentum on the lamina brownish.<br />
5. Lamina beneath <strong>with</strong> patent hairs on the venation.<br />
6. Lateral veins 4-5 pairs; lamina oblong. ............................... 16. C. ferruginea.<br />
6. Lateral veins (4-)5-8 pairs; lamina mostly elliptic to ovate to obovate or to suborbicular.<br />
..........................1.............. ...... . . .. 1 . C. crassivenosa.<br />
5. Lamina beneath <strong>with</strong> appressed hairs on the venation .................. 26. C. microcephala.<br />
4. Arachnoid indumentum on the lamina whitish.<br />
7. Lateral veins 2-3(-4)pairs. ......................... ... .... ................. 3.C. argentea.<br />
7. Lateral veins 4-11 pairs. .......................... .................. 26. C. microcephala.<br />
3. Lamina beneath <strong>with</strong>out arachnoid indumentum.<br />
8. Lateral veins 2-4 pairs. .................. ..... ........ .................... 1. C. <strong>an</strong>gustifolia.<br />
8. Lateral veins 4-11 pairs.<br />
9. Leafy twigs (rather) abruptly thickened <strong>with</strong> wide cavities; stamen 1. .........4. C. asperifolia.<br />
9. Leafy twigs gradually thickened <strong><strong>an</strong>d</strong> solid or <strong>with</strong> narrow cavities.<br />
10. Lateral veins 7-11 pairs; terminal buds usually swollen. .............. 35. C. parvifolia.<br />
10. Lateral veins 4-7 pairs; terminal buds slender.<br />
11. Hairs on the lamina beneath of about equal length .................. 20. C. latifolia.<br />
11. Hairs on the lamina beneath of different lengths ... ................ 43. C. viridifolia.<br />
5. Key to the <strong>Coussapoa</strong> Species in Ecuador<br />
1. Interfloral bracts absent.<br />
2. Basal pair of lateral veins unbr<strong>an</strong>ched (or occasionally a single br<strong>an</strong>ch).<br />
3. Lateral veins 2-7 pairs, the basal pair reaching the margin above to just below the middle of the<br />
lam ina. ................................................................. 3 1. C . orthoneura.<br />
25
26 Flora Neotropica<br />
3. Lateral veins (5-)7-21 pairs, the basal pair reaching the margin (far) below the middle of the<br />
lamina.<br />
4. Stipules 1-4 cm long; apex of the lamina mostly acute to shortly acuminate. . 32. C. ovalifolia.<br />
4. Stipules 4-13 cm long; apex of the lamina mostly obtuse. .................. 27. C. napoensis.<br />
2. Basal pair of lateral veins br<strong>an</strong>ched.<br />
5. Stipules glabrous or sparsely puberulous .....................................34. C. parviceps.<br />
5. Stipules subvelutinous, subhirsute, or subsericeous.<br />
6. Stipules 1-4 cm long; apex of the lamina mostly acute to shortly acuminate. .. 32. C. ovalifolia.<br />
6. Stipules 4-13 cm long; apex of the lamina mostly obtuse. .................. 27. C. napoensis.<br />
1. Interfloral bracts present.<br />
7. Lamina trinervate or only the basal pair of lateral veins strong <strong><strong>an</strong>d</strong> the other lateral veins weak <strong><strong>an</strong>d</strong><br />
inconspicuous.<br />
8. Lamina trinervate; intercostal venation prominent beneath. .... ............ 40. C. trinervia.<br />
8. Lamina <strong>with</strong> only the basal pair of lateral veins strong <strong><strong>an</strong>d</strong> the other lateral veins weak <strong><strong>an</strong>d</strong><br />
inconspicuous; intercostal venation pl<strong>an</strong>e beneath. ............... ........9. C. cinnamomifolia.<br />
7. Lamina pinnately veined.<br />
9. Lamina beneath <strong>with</strong> arachnoid indument.<br />
10. Arachnoid indumentum on the lamina beneath brown(ish).<br />
11. Lateral veins 4-8(1 1) pairs; pistillate inflorescences pluricapitate; staminate flower heads<br />
ca. 2-3 mm in diameter. ........................................ 11. C. crassivenosa.<br />
11. Lateral veins (7-)10-24 pairs; pistillate inflorescences (usually unicapitate; staminate<br />
flower heads 4-10 mm in diameter.<br />
12. Common peduncle of the staminate inflorescence 1-2 cm long, that of the pistillate<br />
inflorescence 0.5-0.8 cm long; pistillate flower heads ellipsoid to clavate. .......<br />
...................................... . ........................ 14. . C . duquei.<br />
12. Common peduncle of the staminate inflorescence 2-6 cm long, that of the pistillate<br />
inflorescence (1)2-13 cm long; pistillate flower heads (sub)globose .... 42. C. villosa.<br />
10. Arachnoid indumentum on the lamina beneath whitish.<br />
13. Stamens 2; pistillate flower heads 1(-3). ................................. 42. C. villosa<br />
13. Stamens 3; pistillate flower heads 3-10.<br />
14. Common peduncle 8-20 cm long; base of the lamina cordate to rounded ........<br />
...................................................... 22. C longepedunculata.<br />
14. Common peduncle up to 7 cm long; base of the lamina usually obtuse to rounded.<br />
............................................................... 19. C. herthae.<br />
9. Lamina beneath <strong>with</strong>out arachnoid indumentum.<br />
15. Lamina beneath <strong>with</strong> domatium-like cavities in the axils of the lateral veins; interfloral bracts<br />
large <strong><strong>an</strong>d</strong> peltate. ...................................................... 10. C. contorta.<br />
15. Lamina beneath <strong>with</strong>out domatium-like cavities; interfloral bracts relatively small <strong><strong>an</strong>d</strong> spathulate<br />
to subpeltate.<br />
16. All lateral veins usually br<strong>an</strong>ched; pistillate inflorescences usually unicapitate; stamen<br />
1. ......... ..................................... . .. 1. .. C. asperifolia.<br />
16. Only the basal pair of lateral veins usually br<strong>an</strong>ched; pistillate inflorescences pluricapitate;<br />
stamens 3.<br />
17. Stipules glabrous or sparsely puberulous ............................... 34. C. parviceps.<br />
17. Stipules <strong>with</strong> dense indumentum.<br />
18. Lamina about as long as broad; lateral veins 5-9 pairs. ....... 41. C. v<strong>an</strong>nifolia.<br />
18. Lamina distinctly longer th<strong>an</strong> broad; lateral veins 8-12 pairs.<br />
19. Common peduncle 8-20 cm long; base of the lamina cordate to rounded.<br />
..................................... 22. C. longepedunculata.<br />
19. Common peduncle up to 7 cm long; base of the lamina usually obtuse to<br />
rounded. .............................................. 19. C. herthae.<br />
6. Key to the <strong>Coussapoa</strong> Species in Peru <strong><strong>an</strong>d</strong> Bolivia<br />
1. Interfloral bracts absent.<br />
2. Basal lateral veins unbr<strong>an</strong>ched (or occasionally <strong>with</strong> a single br<strong>an</strong>ch).<br />
3. Lateral veins 7-21 pairs <strong><strong>an</strong>d</strong> the lamina beneath (almost) glabrous in the areoles. .. 32. C. ovalifolia.<br />
3. Lateral veins 2-9 pairs <strong><strong>an</strong>d</strong>/or the lamina beneath <strong>with</strong> dense, minute hairs in the areoles.<br />
4. Basal lateral veins reaching the margin above or just below the middle of the lamina; stipules<br />
mostly <strong>with</strong>out or <strong>with</strong> sparse long yellow hairs; br<strong>an</strong>ches of the pistillate inflorescence slightly<br />
broadened towards the flower heads ..................................... 31. C. orthoneura.<br />
4. Basal lateral veins reaching the margin below to just above the middle of the lamina; stipules
<strong>Coussapoa</strong><br />
<strong>with</strong> dense, long, yellow hairs; br<strong>an</strong>ches of the pistillate inflorescence strongly broadened<br />
towards the flower heads ................................................ 12. C. cupularis.<br />
2. Basal lateral veins br<strong>an</strong>ched.<br />
5. Lateral veins 5-9 pairs; lamina usually elliptic. ........................... See 12. C. cupularis.<br />
5. Lateral veins 8-15 or 14-22; lamina usually ovate.<br />
6. Indumentum of the petiole consisting of dense minute hairs <strong><strong>an</strong>d</strong> distinctly longer, yellowish<br />
hairs; lateral veins 8-15 pairs; peduncle of the pistillate inflorescence 1-2 mm thick; Amazon<br />
Basin. ........................... .................................... 28. C. nitida.<br />
6. Indumentum of the petiole consisting of short hairs, not differing distinctly in length; lateral<br />
veins 14-22 pairs; peduncle of the pistillate inflorescence 6-10(-15) mm thick; Colombia<br />
(Am azonas). ........................................................ 8. C. cinnam om ea.<br />
1. Interfloral bracts present.<br />
7. Lamina trinervate or only the basal pair of lateral veins strong <strong><strong>an</strong>d</strong> the other lateral veins weak <strong><strong>an</strong>d</strong><br />
inconspicuous.<br />
8. Lamina trinervate; intercostal venation prominent beneath. .................... 40. C. trinervia.<br />
8. Lamina <strong>with</strong> only the basal pair of lateral veins strong <strong><strong>an</strong>d</strong> the other lateral veins weak <strong><strong>an</strong>d</strong><br />
inconspicuous; intercostal venation pl<strong>an</strong>e beneath. ....................... 9. C. cinnamomifolia.<br />
7. Lamina pinnately veined.<br />
9. Lamina beneath glabrous or <strong>with</strong> sparse indumentum.<br />
10. Basal lateral veins unbr<strong>an</strong>ched; stipules up to 1(-1.3) cm long; margin of the lamina +<br />
revolute at the base. .................................................. 35. C. parvifolia.<br />
10. Basal <strong><strong>an</strong>d</strong>/or other lateral veins usually br<strong>an</strong>ched; stipules up to 3.5 cm long; margin of the<br />
lamina not revolute at the base. ......................................... 4. C. asperifolia.<br />
9. Lamina beneath <strong>with</strong> dense indumentum.<br />
11. Lamina beneath <strong>with</strong>out arachnoid indumentum <strong><strong>an</strong>d</strong> basal pair of veins unbr<strong>an</strong>ched. ....<br />
.................................................................... 24. C. m<strong>an</strong>uensis.<br />
11. Lamina beneath <strong>with</strong> arachnoid indumentum <strong><strong>an</strong>d</strong> basal pair of lateral veins br<strong>an</strong>ched.<br />
12. Stamens 3; common peduncle 8-20 cm long; pistillate flower heads 3-10 ...........<br />
............... ........................................... 22. C. longepedunculata.<br />
12. Stamens 2 or 1; common peduncle up to 6 cm long, or if longer in pistillate inflorescences<br />
then a single flower head.<br />
13. Stamens 2; pistillate inflorescences unicapitate, or if pluricapitate then the peri<strong>an</strong>th<br />
of the pistillate flower densely puberulous. ......................... 42. C. villosa.<br />
13. Stamen 1; pistillate inflorescences pluricapitate <strong><strong>an</strong>d</strong> peri<strong>an</strong>th of the pistillate flower<br />
alm ost glabrous. ........................................ ... 39. C. tessm<strong>an</strong>nii.<br />
7. Key to the <strong>Coussapoa</strong> Species in Amazoni<strong>an</strong> Brazil<br />
1. Interfloral bracts absent.<br />
2. Lateral veins 2-8 pairs.<br />
3. Stipules 0.5-1 cm long; stamens 2; pistillate flower heads 1-3 <strong><strong>an</strong>d</strong> common peduncle or br<strong>an</strong>ches<br />
not thickened below these heads. ........................................... 43. C. viridifolia.<br />
3. Stipules usually longer th<strong>an</strong> 2 cm, if shorter th<strong>an</strong> 1 cm then stamen 1 or pistillate flower heads<br />
2-10 <strong><strong>an</strong>d</strong> the br<strong>an</strong>ches of the inflorescence more or less thickened below these heads.<br />
4. Intercostal venation prominent beneath; lamina beneath <strong><strong>an</strong>d</strong> inflorescences <strong>with</strong> dense, white<br />
arachnoid indumentum; Lower Amazon Basin. .......................... 2. C. arachnoidea.<br />
4. Intercostal venation pl<strong>an</strong>e beneath; lamina <strong><strong>an</strong>d</strong> inflorescences <strong>with</strong>out arachnoid indumentum,<br />
or if present then mostly soon disappearing; Upper Amazon Basin.<br />
5. Basal lateral veins reaching the margin above to just below the middle of the lamina;<br />
stipules mostly <strong>with</strong>out or <strong>with</strong> sparse, long, yellow hairs; br<strong>an</strong>ches of the pistillate inflorescence<br />
slightly thickened below the flower heads. .................... 31. C. orthoneura.<br />
5. Basal lateral veins reaching the margin below to just above the middle of the lamina;<br />
stipules <strong>with</strong> dense, long, yellow hairs; br<strong>an</strong>ches of the pistillate inflorescence strongly<br />
thickened below the flower heads. .................................... 12. C. cupularis.<br />
2. Lateral veins (7-)9-22 pairs.<br />
6. Basal lateral veins (mostly unbr<strong>an</strong>ched; lamina beneath in the areoles glabrous or sparsely puberulous.<br />
................................................................. 32. C. ovalifolia.<br />
6. Basal lateral veins br<strong>an</strong>ched; lamina beneath in the areoles <strong>with</strong> dense, minute hairs.<br />
7. Petiole 2-4 mm thick, <strong>with</strong> dense, minute hairs <strong><strong>an</strong>d</strong> sparse, much longer, yellowish hairs.<br />
......................................................................... 28. C . m itida.<br />
7. Petiole 5-7 mm thick, <strong>with</strong> short hairs about the same in length. ......... 8. C. cinnamomea.<br />
27
28 Flora Neotropica<br />
1. Interfloral bracts present.<br />
8. Lamina trinervate ............................................................. 40. C. trinervia.<br />
8. Lamina pinnately veined.<br />
9. Lamina scabrous above.<br />
10. Lateral veins unbr<strong>an</strong>ched. ................................................ 37. C. scabra.<br />
10. All or most lateral veins br<strong>an</strong>ched .......................................4. C. asperifolia.<br />
9. Lamina smooth above.<br />
11. Lam ina trinervate. ..................................................... 40. C. trinervia.<br />
11. Lamina pinnately veined.<br />
12. Lamina beneath <strong>with</strong> dense, partly arachnoid indumentum.<br />
13. Arachnoid indumentum on the lamina brownish.<br />
14. Lateral veins 4-5 pairs; lamina oblong. ..................... 16. C. ferruginea.<br />
14. Lateral veins (4-)5-8 pairs; lamina mostly elliptic to ovate to obovate or to<br />
suborbicular. ........................................... 11. C. crassivenosa.<br />
13. Arachnoid indumentum on the lamina whitish.<br />
15. Stamens 2; pistillate inflorescences (usually) unicapitate; peri<strong>an</strong>th of the pistillate<br />
flower densely puberulous. .............................. 42. C. villosa.<br />
15. Stamen 1; pistillate inflorescences pluricapitate; peri<strong>an</strong>th of the pistillate flower<br />
almost glabrous. ........................................<br />
39. C. tessm<strong>an</strong>nii.<br />
12. Lamina beneath glabrous or <strong>with</strong> sparse indumentum <strong><strong>an</strong>d</strong> arachnoid indumentum lacking.<br />
16. Lateral veins 2-4 pairs. .......................................1. C. <strong>an</strong>gustifolia.<br />
16. Lateral veins 4-11 pairs.<br />
17. Lateral veins 7-11 pairs; terminal buds usually swollen. ...... 35. C. parvifolia.<br />
17. Lateral veins 4-7 pairs; terminal buds slender.<br />
18. Leafy twigs (rather) abruptly thickened <strong><strong>an</strong>d</strong> <strong>with</strong> wide cavities; stamen 1.<br />
...................................................... . C. asperifolia.<br />
18. Leafy twigs gradually thickened <strong><strong>an</strong>d</strong> solid or <strong>with</strong> narrow cavities.<br />
19. Hairs on the lamina beneath distinctly different in lengths. ........<br />
................................................. 43. C. viridifolia.<br />
19. Hairs on the lamina beneath about equal in length ..... 20. C. latifolia.<br />
8. Key to the <strong>Coussapoa</strong> Species in Eastern Brazil<br />
(Rio Gr<strong><strong>an</strong>d</strong>e do Sul to Pernambuco)<br />
1. Stipules up to 1 cm long. ........................................... 13. C. curr<strong>an</strong>ii.<br />
1. Stipules at least 1 cm long.<br />
2. Lamina above <strong>with</strong> subpersistent, white, arachnoid indumentum. ................... 17. C. floccosa.<br />
2. Lamina above glabrous.<br />
3. Stamens much longer th<strong>an</strong> the peri<strong>an</strong>th; pistillate inflorescences mostly unicapitate, (common)<br />
peduncle 1-2.5 cm long; stipules up to 7 cm long; leaf margin usually revolute towards the base.<br />
........................................................................ 25. C . m icrocarp a.<br />
3. Stamens as long as the peri<strong>an</strong>th; pistillate inflorescences mostly pluricapitate, common peduncle<br />
2.5-4 cm long; stipules up to 2 cm long; leaf margin entirely revolute or pl<strong>an</strong>e ...............<br />
....................................................................... 33. C . p achyp hylla.<br />
1. <strong>Coussapoa</strong> <strong>an</strong>gustifolia Aublet, Hist. pl. Gui-<br />
<strong>an</strong>e 2: 956, t. 363. 1775; Miquel, Fl. bras. 4(1):<br />
138. 1853; Berg. Fl. Suriname 5(1): 283. 1975.<br />
Type. French Gui<strong>an</strong>a. Without locality, -,<br />
Aublet s.n. (BM, leaf fragment). Fig. 10.<br />
Shrub or tree, mostly hemi-epiphytic, up to 30<br />
m tall. Leafy twigs 3-8 mm thick, glabrous or<br />
sparsely, minutely puberulous, lenticels numer-<br />
ous <strong><strong>an</strong>d</strong> conspicuous. Lamina coriaceous ob-<br />
ovate or subobovate (to obl<strong>an</strong>ceolate or ellip-<br />
tic to oblong), 2-11 x 1-6 cm, apex obtuse to<br />
rounded or emarginate, base acute to cuneate (or<br />
to subobtuse), margin entire; upper surface glabrous,<br />
lower surface glabrous or sparsely appressed-puberulous;<br />
lateral veins 2-4 pairs,<br />
straight or curved, basal pair unbr<strong>an</strong>ched, reaching<br />
the margin far above the middle of the lamina;<br />
intercostal venation mostly pl<strong>an</strong>e; petiole 1-<br />
5 cm long, glabrous or sparsely appressed-puberulous;<br />
stipules 0.5-1 cm long, glabrous or<br />
sparsely appressed-puberulous. Staminate inflorescences<br />
poorly br<strong>an</strong>ched, heads 3-5, sometimes<br />
partly fused, globose, 5-8 mm diam.; common<br />
peduncle 0.5-1.5 cm long, minutely
<strong>Coussapoa</strong> 29<br />
FIG. 10. <strong>Coussapoa</strong> <strong>an</strong>gustifolia: 1, leafy twig <strong>with</strong> pistillate inflorescences (Schulz 10344a); 2, staminate<br />
inflorescences (Pires et al. 50631).<br />
puberulous; peri<strong>an</strong>th 1 mm high, glabrous; sta-<br />
men one, as long as the peri<strong>an</strong>th. Pistillate inflo-<br />
rescences unbr<strong>an</strong>ched or rarely br<strong>an</strong>ched; heads<br />
1(-2), globose, ca. 5 mm, in fruit up to 20 mm<br />
diam.; (common) peduncle 0.3-3 cm long, mi-<br />
nutely puberulous; peri<strong>an</strong>th 1(-2) mm high, gla-<br />
brous; fruiting peri<strong>an</strong>th yellow to or<strong>an</strong>ge. Inter-<br />
floral bracts spathulate to subpeltate, apex<br />
ciliolate.<br />
Distribution (Fig. 6). Surinam, French Gui<strong>an</strong>a,<br />
<strong><strong>an</strong>d</strong> Brazil (Amapa <strong><strong>an</strong>d</strong> Para); in upl<strong><strong>an</strong>d</strong> <strong><strong>an</strong>d</strong> riverine<br />
forest.<br />
Specimens studied. SURINAM. Brownsberg, 6 Sep<br />
1915 (e), BW 772 (A, K, NY, U); Brokopondo Lake,
30<br />
10 Sep 1965 (st), V<strong>an</strong> Donselaar 2569 (U); Wilhelmina<br />
Mts., Zuid: River, 45 km above confluence <strong>with</strong> Lucie<br />
River, 25 Sep 1963 (9), Irwin et al. 57574 (B, G, MO,<br />
NY, OXF, R, S, U); Coppename River, Bitagron, 28<br />
Jul 1954 (9), Lindem<strong>an</strong> 6362 (F, U); Z<strong><strong>an</strong>d</strong>erij Sav<strong>an</strong>na,<br />
28 Aug 1954 (st), Lindem<strong>an</strong> 6545 (U); Z<strong><strong>an</strong>d</strong>erij, 15<br />
Apr 1954 (st), Lindem<strong>an</strong> 6600 (MO, U); Saramacca<br />
River, Jacobkondre, 19 Jun 1944 (6), Maguire 23867<br />
(A, F, K, NY, U, US); Saramacca River, nr. Kwattahede,<br />
22 Jun 1944 (2), Maguire 23942 (A, F, K, MO,<br />
NY, P, RB, U); Wilhelmina Mts., S of Juli<strong>an</strong>atop, 7<br />
Aug 1963 (8), Schulz 10344a (NY, U); Suriname River,<br />
G<strong>an</strong>see, 8 Jul 1908 (9), Tresling 55 (U).<br />
FRENCH GUIANA. Without locality, - (st), Aublet<br />
s.n. (BM, type collection); It<strong>an</strong>y River, isl<strong><strong>an</strong>d</strong> in<br />
rapids, 30 Sep 1961 (9), BAFOG (=Graimleux) 7938<br />
(CAY, U); Maroni River, 15 Sep 1903 (9), Geay 3285<br />
(P), Oyapock River, nr. Gr<strong><strong>an</strong>d</strong> Roche Fall, 17 Jul 1960<br />
(9), Maguire et al. 47049 (G, GH, LE, NY, OXF, U,<br />
US); nr. Cayenne, - (9), Martin s.n. (BM, BR, K);<br />
Maroni River, 1854 (9), Melinon 172 (LE, P); Camopi<br />
River, below Gr<strong><strong>an</strong>d</strong> Tamouri Creek, 2 Feb 1968 (2),<br />
Oldem<strong>an</strong> et al. 207 (CAY, P, U, US); Sinnamary River,<br />
1.5 km above Courcibo River, 7 Aug 1967 (9), Oldem<strong>an</strong><br />
B1167 (CAY); Kourou River, 2 km above Cr6ole,<br />
18 Sep 1967 (9), Oldem<strong>an</strong> B1324 (CAY, U); Oropu<br />
River, 1.5 km above Degrad Lal<strong><strong>an</strong>d</strong>e, 16 Oct 1965 (9),<br />
Oldem<strong>an</strong> 1619 (CAY); Approuague River, 5 km above<br />
Maripa Creek, 3 Feb 1967 (9), Oldem<strong>an</strong> 2476 (CAY,<br />
P, U); Oyapock River, above Moutouci Falls, 15 May<br />
1970 (6), Oldem<strong>an</strong> B3243 (CAY, U); Sinnamary, road<br />
to St. Elie, 18 Jul 1977 (6), Sastre 5500 (P, U).<br />
BRAZIL. AMAPA: Rio Araguari, between 1055'N,<br />
51?51'W <strong><strong>an</strong>d</strong> 2?5'N, 51056'W, 1 Sep 1961 (9), Pires et<br />
al. 50631 (NY, U, US). PARA: Mun. Oriximina, road<br />
to Cachoeira Porteira, 18 Jun 1980 (9), Cid et al. 1053<br />
(U); Rio Paru do Oeste, Cachoeira Chuvisco, 17 Sep<br />
1980 (9), Cid et al. 2223 (U), Ilhas de Breves, Furo<br />
Macujubim, 16 Nov 1922 (9), Ducke (HJBR) 18461<br />
(RB); Rio Trombetas, Cachoeira Porteira, 8 Jun 1978<br />
(9), N. T. Silva et al. 4757 (U).<br />
This species, <strong>with</strong> few lateral veins in a mostly<br />
subobovate lamina, appears to be related to C.<br />
trinervia. In its leaf characters C. <strong>an</strong>gustifolia is<br />
tr<strong>an</strong>sitional to the trinervate state of the latter<br />
species. It also shows m<strong>an</strong>y similarities to the<br />
Central Americ<strong>an</strong> C. glaberrima.<br />
2. <strong>Coussapoa</strong> arachnoidea Akkerm<strong>an</strong>s & C. C.<br />
Berg, Proc. Kon. Ned. Akad. Wetensch. Ser.<br />
C, 85(4): 442. 1982. Type. Brazil. Amapa: Rio<br />
Araguari, between 1?55'N, 51?59'W <strong><strong>an</strong>d</strong> 2?5'N,<br />
51?56'W, 1 Sep 1961 (2), Pires et al. 50656<br />
(holotype, IAN; isotypes, M, MG, NY, OXF,<br />
US, VEN). Fig. 11.<br />
Tree. hemi-epiphytic, up to 35 m tall. Leafy<br />
twigs 3-8 mm thick, <strong>with</strong> sparse, arachnoid hairs<br />
<strong><strong>an</strong>d</strong> yellowish, straight hairs on the scars of the<br />
Flora Neotropica<br />
stipules. Lamina coriaceous, elliptic to ovate or<br />
to l<strong>an</strong>ceolate, (6-)9-16 x (2-)3-7 cm, apex acute,<br />
base obtuse to acute, margin entire; upper surface<br />
glabrous, lower surface in the areoles rather<br />
densely, minutely puberulous, on the main veins<br />
sparse, rather long, yellowish hairs, initially the<br />
whole surface densely covered <strong>with</strong> white arachnoid<br />
hairs; lateral veins 5-8 pairs, almost straight,<br />
basal pair unbr<strong>an</strong>ched, reaching the margin at or<br />
just below the middle; intercostal venation rather<br />
prominent; petiole 3-6 cm long, villous <strong>with</strong> white<br />
arachnoid hairs, glabrescent; stipules 5-8 cm long,<br />
villous <strong>with</strong> white arachnoid hairs, mixed <strong>with</strong><br />
rather long <strong><strong>an</strong>d</strong> straight appressed, yellowish hairs<br />
<strong><strong>an</strong>d</strong> <strong>with</strong> reddishbrown pluricellular hairs. Staminate<br />
inflorescences unknown. Pistillate inflorescences<br />
br<strong>an</strong>ched; heads 2-4, globose ca. 4-5<br />
mm diam.; common peduncle 1.5-2 cm long,<br />
densely villous <strong>with</strong> yellowish to white arachnoid<br />
hairs, mixed <strong>with</strong> straight appressed yellowish<br />
hairs <strong><strong>an</strong>d</strong> <strong>with</strong> reddish-brown pluricellular hairs,<br />
apices of the br<strong>an</strong>ches broadened towards the<br />
heads; peri<strong>an</strong>th ca. 1 mm high, minutely puberulous.<br />
Interfloral bracts absent.<br />
Distribution (Fig. 6). Brazil (Amapa).<br />
Specimens studied. BRAZIL. AMAPA: Rio Araguari,<br />
between 1?55'N, 51059'W <strong><strong>an</strong>d</strong> 2?5'N, 51056'W, 1 Sep<br />
1961 (v), Pires et al. 50656 (IAN, M, MG, NY, OXF,<br />
US, VEN, type collection).<br />
<strong>Coussapoa</strong> arachnoidea appears to be closely<br />
related to C. orthoneura, from which it differs in<br />
the prominent intercostal venation <strong><strong>an</strong>d</strong> the dense<br />
covering of white arachnoid hairs on the leaves<br />
<strong><strong>an</strong>d</strong> inflorescences. Staminate inflorescences are<br />
needed to determine the validity or not of the<br />
presumed relationship. It may prove to be distinct<br />
from C. orthoneura only at the subspecific<br />
level.<br />
3. <strong>Coussapoa</strong> argentea Akkerm<strong>an</strong>s & C. C. Berg,<br />
Proc. Kon. Ned. Akad. Wetensch. Ser. C, 85(4):<br />
443. 1982. Type. Venezuela. Bolivar: 119 km<br />
S of El Dorado, 12 J<strong>an</strong> 1964 (6), Steyermark<br />
et al. 92991 (holotype, VEN; isotypes, NY, U,<br />
US). Fig. 12.<br />
Tree, hemi-epiphytic or terrestrial, up to 30 m<br />
tall. Leafy twigs 3-5 mm thick, brownish pu-<br />
berulous to strigose or partly to hirsute <strong><strong>an</strong>d</strong> <strong>with</strong><br />
brown to white arachnoid hairs. Lamina coria-<br />
ceous, obovate to elliptic, occasionally ovate or
<strong>Coussapoa</strong><br />
FG. 11. <strong>Coussapoa</strong>1 ly tg wh p e is (s et<br />
FIG. 11. <strong>Coussapoa</strong> arachnoidea: 1, leafy twig <strong>with</strong> pistillate inflorescences<br />
(Pires et al. 50656).<br />
oblong, 2-12 x 1-6 cm, apex obtuse to acute or<br />
acuminate, base obtuse, margin entire; upper<br />
surface glabrous, lower surface sparsely to densely<br />
appressed-puberulous <strong><strong>an</strong>d</strong> <strong>with</strong> a dense cov-<br />
cm.<br />
31<br />
ering of silvery arachnoid hairs, deciduous <strong>with</strong><br />
age; lateral veins 2-3(-4) pairs, slightly curved,<br />
basal pair unbr<strong>an</strong>ched, reaching the margin above<br />
the middle of the lamina; intercostal venation
32 Flora Neotropica<br />
c.<br />
FIG. 12. <strong>Coussapoa</strong> argentea: 1, leafy twig <strong>with</strong> pistillate inflorescences (Steyermark & Nilsson 487); 2, leafy<br />
twig <strong>with</strong> staminate inflorescences (Steyermark & Nilsson 388).
<strong>Coussapoa</strong> 33<br />
slightly prominent to almost pl<strong>an</strong>e; petiole 1-2 abruptly thickened just below the apex <strong><strong>an</strong>d</strong> widely<br />
cm long, brownish puberulous to strigose or part- hollow, (rather) sparsely puberulous to hispiduly<br />
to hirsute <strong><strong>an</strong>d</strong> <strong>with</strong> brown to white arachnoid lous, often also <strong>with</strong> brown pluricellular hairs,<br />
hairs; stipules 1-5 cm long, densely covered <strong>with</strong> sometimes partly hirtellous. Lamina coriaceous<br />
brownish curled to arachnoid hairs <strong><strong>an</strong>d</strong> whitish to subcoriaceous, obovate to subobovate to elstraight<br />
hairs. Staminate inflorescences repeat- liptic to oblong to suborbicular to ovate or to<br />
edly br<strong>an</strong>ched; heads m<strong>an</strong>y, globose, ca. 2-3 mm cordiform, 7-32 x 4-24 cm, apex rounded to<br />
diam.; common peduncle 2-3 mm long, puber- obtuse to emarginate, base obtuse to rounded to<br />
ulous to hirtellous; peri<strong>an</strong>th ca. 1 mm high, truncate to cordate, margin entire to subcrenate;<br />
3-lobed, glabrous; stamens two, exceeding the upper surface scabrous <strong>with</strong> minute rigid hairs<br />
peri<strong>an</strong>th. Pistillate inflorescences unbr<strong>an</strong>ched; to smooth <strong><strong>an</strong>d</strong> glabrous, lower surface densely<br />
head globose, ca. 4 mm diam.; peduncle 2-3.5 to sparsely puberulous <strong>with</strong> curved (to straight)<br />
cm long, puberulous, <strong><strong>an</strong>d</strong> <strong>with</strong> silvery arachnoid hairs or only minutely puberulous on the main<br />
hairs; peri<strong>an</strong>th ca. 1 mm high, glabrous. Inter- veins, sometimes partly hirtellous, in juvenile<br />
floral bracts spathulate to subpeltate, sometimes leaves sometimes the whole surface hisfew.<br />
pid(ulous); lateral veins 4-9 pairs, usually all or<br />
Distribution (Fig. 6). Venezuela (Bolivar); most of them br<strong>an</strong>ched, sometimes the basal pair<br />
mostly in moist forest on table mountains, be- unbr<strong>an</strong>ched or faintly br<strong>an</strong>ched but some of the<br />
tween (450-)1000 <strong><strong>an</strong>d</strong> 1300 m.<br />
other lateral veins poorly br<strong>an</strong>ched, (main) basal<br />
pair reaching the margin at, below, or above the<br />
Specimens studied. VENEZUELA. BOLIVAR: Road middle of the<br />
El Dorado-S<strong>an</strong>ta<br />
lamina; intercostal venation<br />
Elena, km 109, 23<br />
prom-<br />
Feb 1959 (6), Bernardi<br />
7238<br />
inent to<br />
(G); upper Rio Cuyuni, Cumbre de La Es- pl<strong>an</strong>e; petiole 1.5-15 cm long, sparsely<br />
calera, 1000 m, 20-21 Aug 1962 (6), Maguire et al. to densely minutely puberulous, sometimes part-<br />
46879 (U, US, VEN); nr. Cerro Uei, between Luepa ly hirtellous; stipules 0.5-3.5 cm long, puberu<strong><strong>an</strong>d</strong><br />
Cerro Venamo, 1100 m, 20-22 Apr 1960 (6), Stey- lous to hispidulous, often also <strong>with</strong> brown pluriermark<br />
& Nilsson 388 (NY, US, VEN); NE of Luepa, cellular<br />
1300 m, 23 Apr 1960 (v), Steyermark & Nilsson<br />
hairs, sometimes<br />
487<br />
partly hirtellous.<br />
(NY, VEN); 119 km S of El Dorado, 12 J<strong>an</strong> 1964 Staminate<br />
(6),<br />
inflorescences br<strong>an</strong>ched; heads m<strong>an</strong>y,<br />
Steyermark et al. 92991 (NY, U, US, VEN, type col- free or partly fused, (sub)globose, 3-10 mm diam.;<br />
lection).<br />
common peduncle 2-8 cm long, puberulous;<br />
peri<strong>an</strong>th ca. 1 mm high, minutely puberulous or<br />
<strong>Coussapoa</strong> argentea is similar to C. micro- glabrous; stamen one, as long as or slightly longer<br />
cephala in m<strong>an</strong>y features. The species differs from th<strong>an</strong> the peri<strong>an</strong>th. Pistillate inflorescence unthe<br />
latter in the smaller number of lateral veins, br<strong>an</strong>ched or br<strong>an</strong>ched; heads 1-6, free or partly<br />
the basal pair of which reaches the margin below fused, 3-12 mm, in fruit up to 30 mm diam.;<br />
the middle of the lamina. Moreover, the pistillate (common) peduncle 1-9.5 cm long, puberulous<br />
inflorescence is unicapitate, being normally plur- to almost glabrous; peri<strong>an</strong>th 1.2 mm high, miicapitate<br />
in C. microcephala. The two species nutely puberulous or glabrous. Interfloral bracts<br />
appear to differ ecologically (altitudinal condi- (sub)spathulate to (sub)peltate, minutely pubertions)<br />
<strong><strong>an</strong>d</strong> may prove to be distinct only at the ulous at the apex.<br />
subspecies level.<br />
C. asperifolia differs from the other <strong>Coussapoa</strong><br />
species: (1) The endocarp body is pushed out of<br />
4. <strong>Coussapoa</strong> asperifolia Trecul, Ann.<br />
the<br />
Sci. Nat. peri<strong>an</strong>th through the torn apex of the peri-<br />
Bot. Ser. 3, 8: 96. 1847; Miquel, Fl. bras. <strong>an</strong>th, due to the<br />
4(1):<br />
exp<strong>an</strong>sion of the mucilaginous<br />
135. 1853; Macbride, Publ. Field Mus. Bot. mesocarp (see p. 8). (2) Furthermore, the leafy<br />
13(2.2): 296. 1937; Berg, Fl. Suriname twigs widen rather abruptly<br />
5(1):<br />
just below the apex<br />
284. 1975. Lectotype. Suriname.<br />
<strong><strong>an</strong>d</strong> have<br />
Without<br />
wide<br />
lo-<br />
cavities, which are often occupied<br />
cality, 1843 ($), Hostm<strong>an</strong>n 1189 by <strong>an</strong>ts.<br />
(holotype, P;<br />
Within this<br />
isotypes, B, G, K, LE, U).<br />
widespread <strong><strong>an</strong>d</strong> polymorphic<br />
species three subspecies c<strong>an</strong> be recognized. These<br />
Tree or shrub, hemi-epiphytic or terrestrial, up three subspecies are not well separated morphoto<br />
25 m tall. Leafy twigs 3-20 mm thick, (usually) logically or geographically.
34 Flora Neotropica<br />
Key to the Subspecies of <strong>Coussapoa</strong> asperifolia<br />
1. Lamina scabrous to scabridulous above, densely (to rather sparsely) puberulous <strong>with</strong> curved hairs on<br />
the smaller veins beneath; intercostal venation prominent; lamina mostly obovate. ....a. ssp. asperifolia.<br />
1. Lamina smooth (to scabridulous) above, glabrous or sparsely puberulous on the smaller veins, sparsely<br />
to rather densely minutely puberulous <strong>with</strong> straight hairs on the main veins beneath; intercostal venation<br />
pl<strong>an</strong>e to prominent.<br />
2. All or most lateral veins br<strong>an</strong>ched, sometimes poorly so <strong><strong>an</strong>d</strong> the basal pair unbr<strong>an</strong>ched; lamina<br />
mostly ovate to cordiform or elliptic; pistillate heads mostly 1, more th<strong>an</strong> 10 mm, in fruit up to 35<br />
mm in diam. ........................................................... ..b. ssp. magnifolia.<br />
2. Few or none of the lateral veins (poorly) br<strong>an</strong>ched, the basal pair unbr<strong>an</strong>ched or faintly br<strong>an</strong>ched;<br />
lamina mostly obovate to subobovate; pistillate heads 3-5, ca. 3-6 mm, in fruit up to 15 mm in<br />
diam ........ ........................................................... c. ssp. rham noides.<br />
4a. <strong>Coussapoa</strong> asperifolia ssp. asperifolia 1921 (6), BW 5262 (A, U); <strong>with</strong>out locality, 1843 (8),<br />
Fig. 13. Hostm<strong>an</strong>n 1189 (B, G, K, LE, P, U, lectotype collection);<br />
Lely Mts., 19 Sep 1975 (st), Lindem<strong>an</strong> et al. 34<br />
<strong>Coussapoa</strong> ficina St<strong><strong>an</strong>d</strong>ley, Publ. Field Mus. Bot. 17: (U); Concession Haenen, W of mouth of Coppename<br />
161. 1937. Type. Brazil. Amazonas: Nr. M<strong>an</strong>aus, River, 25 Mar 1954 (st), Lindem<strong>an</strong> 5709 (U); upper<br />
road to Aleixo, 12 Aug-1 Sep 1936 (9), Krukoff7966 Lit<strong>an</strong>i River, 9 Aug 1937 (a), Rombouts 803 (A, K,<br />
(holotype, NY; isotypes, A, BM, F, K, LE, MO, S, NY, U); Pl<strong>an</strong>tation Lustrijk, Dec 1837 (8), Splitgerber<br />
U).<br />
448 (=122) (L, P); Maratakka River, 20 Aug 1976 (2),<br />
<strong>Coussapoa</strong> cayennensis Hawkes, Phytologia 3: 30. 1948. Teunissen (LBB) 16031 (U).<br />
Type. French Gui<strong>an</strong>a. Nr. Cayenne, 17 Jul 1921 (9), FRENCH GUIANA. It<strong>an</strong>y River, isl<strong><strong>an</strong>d</strong> in rapids,<br />
Broadway 880 (holotype, NY; isotypes, GH, NY, 30 Sep 1962 (9), BAFOG (=Graimleux) 7942 (CAY,<br />
US).<br />
U); nr. Cayenne, 17 Jul 1921 (9), Broadway 880 (GH,<br />
NY, US, type collection of C. cayennensis); Oyapock<br />
Lamina mostly obovate to subobovate or el- River, Les Trois Sauts, 27 Mar 1975 (9), Gren<strong><strong>an</strong>d</strong> 966<br />
liptic to oblong, mostly 10-17 cm long, base<br />
(CAY); Anse Bruyere, 15 Aug 1978 (a), Gren<strong><strong>an</strong>d</strong> 1596<br />
(U); nr. Cayenne,<br />
mostly obtuse to rounded; upper surface scabrous<br />
<strong>with</strong> a dense covering of minute rigid hairs,<br />
sometimes scabridulous, lower surface on the<br />
smaller veins mostly densely puberulous <strong>with</strong><br />
curved (to straight) hairs; lateral veins 4-7 pairs,<br />
all or most of them br<strong>an</strong>ched, the (main) basal<br />
pair reaching the margin above the middle of the<br />
lamina; intercostal venation prominent; petiole<br />
1.5-9 cm long; stipules 0.5-1.5 cm long. Staminate<br />
inflorescences <strong>with</strong> heads up to 10 mm<br />
diam.; common peduncle up to 4.5 cm long. Pistillate<br />
inflorescences <strong>with</strong> mostly, sometimes two,<br />
(partly fused) heads, mostly more th<strong>an</strong> 1 cm, in<br />
fruit up to 2.5 cm diam.; (common) peduncle 1-<br />
3 cm long.<br />
Distribution (Fig. 6). Eastern Gui<strong>an</strong>a region<br />
(Surinam <strong><strong>an</strong>d</strong> French Gui<strong>an</strong>a), in Brazil in Amapa<br />
<strong><strong>an</strong>d</strong> eastern Para <strong><strong>an</strong>d</strong> also, apparently segregated<br />
from the main area, in the central part of<br />
the Amazon Basin; in upl<strong><strong>an</strong>d</strong> <strong><strong>an</strong>d</strong> riverine forest.<br />
This distribution shows striking similarities to<br />
that of C. latifolia.<br />
Specimens studied. SURINAM. Section 0, 11 Nov<br />
1915 (st), BW1322 (U); Cor<strong>an</strong>tijn River, Kabouri, 26<br />
Jun 1916 (st), BW2070 (U); Maratakka River, 25 Nov<br />
1917 (9), BW 3430 (F, NY, U, US); Section 0, 10 Aug<br />
- (9), Lem<strong>an</strong> s.n. (P); nr. Cayenne,<br />
Mar 1900 (st), Lemee s.n. (P); <strong>with</strong>out locality, - (9),<br />
Leprieur s. n. (P); Kuataka, Antecume Pata, confluence<br />
of It<strong>an</strong>y River <strong><strong>an</strong>d</strong> Marouini River, 14 Nov 1977 (9),<br />
Moretti 837 (CAY); Approuague River, between Sapokaye<br />
Creek <strong><strong>an</strong>d</strong> Gr<strong><strong>an</strong>d</strong> C<strong>an</strong>ori Falls, 23 Oct 1968<br />
(9), Oldem<strong>an</strong> T243 (CAY, GH, NY, P, U); Oyapock<br />
River, between Notaye Creek <strong><strong>an</strong>d</strong> Caim<strong>an</strong> Creek, 10<br />
Jul 1969 (st), Oldem<strong>an</strong> T366 (CAY, U); Oyapock River,<br />
Sav<strong>an</strong>e-Roche, Baton Pilon, 12 Jul 1969 (6), Oldem<strong>an</strong><br />
B2538 (CAY, U); Oyapock River, above Oscar,<br />
18 Jun 1970 (9), Oldem<strong>an</strong> B3420 (CAY, P, U); <strong>with</strong>out<br />
locality, 1820 (9), Perrotet s.n. (P); Sinnamary, road to<br />
St. Elie, km 8, 28 Aug 1979 (st), Provost 738 (U);<br />
Acarou<strong>an</strong>y, 1856 (st), Sagot s.n. (P); nr. Kourou, 14<br />
Dec 1978 (9), Sastre 6417 (CAY, P, U); <strong>with</strong>out locality,<br />
23 Jun 1921 (9), Wachenheim 426 (P).<br />
BRAZIL. AMAPA: Rio Araguari, between 1?2'N,<br />
51?15'W <strong><strong>an</strong>d</strong> 0?57'N, 51?29'W, 13 Sep 1961 (6), Pires<br />
et al. 50913 (NY, OXF, R, U, US). AMAZONAS: Nr.<br />
M<strong>an</strong>aus, BR. 17, ca. km 9, 5 Aug 1955 (9), D. Coelho<br />
(INPA) 1577 (U), 8 Jul 1955 (8), Chagas (INPA) 1364<br />
(U); nr. M<strong>an</strong>aus, Colonia Joao Alfredo, 21 Aug 1947<br />
(9), Ducke 2103 (IAN, R, U); nr. M<strong>an</strong>aus, Pensador,<br />
Aug 1910 ($), Ule 8836 (G, K, MG); M<strong>an</strong>aus-Caracarai<br />
road, ca. km 130, 13 Feb 1974 (9), Stewardet al. P.20255<br />
(MO, U). PARA: Res. Florest, Rio Aura, 18 Oct 1978<br />
(9), Carauta et al. 3011 (RB); Belem, 21 Oct 1915 (st),<br />
Ducke (HAMP) 15804 (BM); Mun. Oriximinf, Rio<br />
Mapuera, 19 Aug 1986 (a), Ferreira et al. 7831 (BG);<br />
Arama, 2 Nov 1900 (9), Huber (HAMP) 1873 (G); Rio<br />
J<strong>an</strong>, rd. to Munguba, km 15, 13 Jul 1964 (9), N. T.<br />
Silva 2166 (U).
.". ~:.,, .<br />
.
36 Flora Neotropica<br />
4b. <strong>Coussapoa</strong> asperifolia ssp. magnifolia (Trecul)<br />
Akkerm<strong>an</strong>s & C. C. Berg, Proc. Kon. Ned.<br />
Akad. Wetensch. Ser. C, 85(4): 445. 1982.<br />
Fig. 14.<br />
<strong>Coussapoa</strong> magnifolia Trecul, Ann. Sci. Nat. Bot. Ser.<br />
3, 8: 98. 1847; Miquel, Fl. bras. 4(1): 139. 1853;<br />
Macbride, Publ. Field Mus. Bot. 13(2.2): 297. 1937;<br />
Woodson & Schery, Ann. Missouri Bot. Gard. 47:<br />
169. 1960. Type. Peru. Without locality, - (2), Ruiz<br />
& Pavon s.n. (or 4) (holotype, FI; isotypes, B, BM,<br />
F, US).<br />
<strong>Coussapoa</strong> ruizii Klotzsch, Linnaea 20:529. 1847. Type.<br />
Peru. Without locality, - Specimens studied. PANAMA. CANAL ZONE: Fort<br />
S<strong>an</strong> Lorenzo, 13 Sep 1947 (a), Alien 5120 (BM, G,<br />
MO); Barro Colorado Isl<strong><strong>an</strong>d</strong>, 27 Apr 1968 (a), Croat<br />
5124 (DUKE, F, MO), (9), Croat 5125 (F), 1 Oct 1968<br />
(9), Croat 6588 (MO), 10 Feb 1969 (8), Croat 7859<br />
(MO), 13 Jul 1969 (6), Foster 1101 (DUKE, F), Chagres,<br />
22 Jun 1860 (9 + d), Hayes 354 (NY, syntypes of C.<br />
chagresi<strong>an</strong>a); nr. Fort Sherm<strong>an</strong>, 16 Nov 1955 (2), Johnston<br />
1709 (A); E of Indio Tower, 2 Apr 1956 (9), Johnston<br />
1768 (A, MO); Barro Colorado Isl<strong><strong>an</strong>d</strong>, 16 Feb<br />
1932 (a), Woodworth & Vestal 606 (A, F, MO).<br />
COLOMBIA. ANTIOQUIA: Rio Porce, 1000 m, Dec<br />
1962 (2), Espinal 882 (COL). BOYACA: El Humbo region,<br />
130 km N ofBogoti, 1200-1300m, 19 May 1933<br />
(9), Ruiz & Pavon s.n. (or (8), Lawr<strong>an</strong>ce 810 (A, F, G, K, MO, US, type collection<br />
4) (holotype, B; isotypes, BM, F, FI, US).<br />
of C. hypochlora). CAQUETA: La Tagua, 5 May 1953<br />
<strong>Coussapoa</strong> hypochlora St<strong><strong>an</strong>d</strong>ley, Publ. Field Mus. Bot. (9), Romero-Cast<strong>an</strong>eda 4211 (COL, MO). CHOCO: Nr.<br />
17: 162. 1937. Type. Colombia. Boyaca: El Humbo Helipad, Rio Tru<strong><strong>an</strong>d</strong>o, at junction <strong>with</strong> Quebrada Buche,<br />
region, 130 km N of Bogota, ca. 1150 m, 19 May 1 Apr 1968 (a), Duke 15742 (U); El Valle, beach, 5<br />
1933 (6), Lawr<strong>an</strong>ce 810 (holotype, F; isotypes, A, G, Aug 1976 (9), Gentry et al. 17217 (MO, NY, U); Rio<br />
K, MO, US).<br />
El Valle, between El Valle <strong><strong>an</strong>d</strong> village nr. mouth of Rio<br />
<strong>Coussapoa</strong> cardonae Pittier, Bol. Soc. Venez. Cienc. Mutatt, 10 Aug 1976 (8), Gentry et al. 17493 (U); 5<br />
Nat. 8: 257. 1943, as cardonaei. Type. Venezuela. km E of Istmo de S<strong>an</strong> Pablo, ca. 12 km W of Las<br />
Bolivar: Rio Caura, sab<strong>an</strong>as de C<strong>an</strong>aracuni, J<strong>an</strong> 1942 Animas, 12 J<strong>an</strong> 1979 (2), Gentry et al. 24074 (MO, U).<br />
(9), Cardona 374 (holotype, VEN; isotypes, GH, S, CUNDINAMARCA: La Palma, road to Pacho, Rio Murca,<br />
US).<br />
1150-1400 m, 27 Jul 1947 (2), Garcia-Barriga 12408<br />
<strong>Coussapoa</strong> chagresi<strong>an</strong>a Hawkes, Phytologia 3: 30. 1948. (US). SANTANDER: Rio Opon, Atl<strong>an</strong>tico railway, 5 Jul<br />
Syntypes. P<strong>an</strong>ama. C<strong>an</strong>al Zone, Chagres, 22 Jun 1860 1958 (6), Garcia-Barriga 16239 (NY). VALLE: Nr.<br />
(6 <strong><strong>an</strong>d</strong> 9), Sutton Hayes 354 (NY).<br />
Buenaventura, 15 Nov 1979 (8), V<strong>an</strong> Rooden et al. 258<br />
(U). VAUPES: Rio Vaup6s, Yurupari Falls, 12 Apr 1953<br />
Lamina mostly ovate to cordiform or elliptic (8), Schultes et al. 18996 (F), Nov 1951 (2), Schultes et<br />
al. 19758<br />
to suborbicular, sometimes obovate, mostly 17-<br />
(GH, MO, US).<br />
VENEZUELA. AMAZONAS: Between Yavita <strong><strong>an</strong>d</strong><br />
25 cm long, base often subcordate to cordate; Maroa, 6-19 Jul 1969 (st), Bunting et al. 3959 (MY,<br />
upper surface smooth <strong><strong>an</strong>d</strong> glabrous, sometimes U); upper Rio Orinoco, Raudal de los Guaharibos, 24<br />
scabridulous, lower surface on the smaller veins Jul 1951 (2), Croizat 343 (F, US); Yavita, 10 Jul 1972<br />
glabrous or sparsely puberulous, on the main (9), Lizot (1972-) 1 (US, VEN); S<strong>an</strong> Carlos de Rio<br />
Negro, 7 Mar 1942 (9), LI. Williams 14675 (A, F, G,<br />
veins sparsely to densely minutely puberulous K, NY, RB, US, VEN). BOLIVAR: Rio Caura, sab<strong>an</strong>as<br />
<strong>with</strong> straight hairs, sometimes on the main veins de C<strong>an</strong>aracuni, J<strong>an</strong> 1942 (9), Cardona 374 (A, S, US,<br />
partly hirtellous; lateral veins 4-9 pairs, all or VEN, type collection of C. cardonae); Rio Cuyuni, El<br />
most of them br<strong>an</strong>ched, sometimes the basal pair Chibau, 2 km S of Rio Chibau, 1 Sep 1962 (9), Maguire<br />
et al. 53517<br />
not or faintly br<strong>an</strong>ched but some of the other<br />
(F, NY, OXF, US, VEN); Rio Cuyuni, NE<br />
of Luepa, 1300 m, 23 Apr 1960 (9), Steyermark &<br />
lateral veins poorly br<strong>an</strong>ched, basal pair reaching Nilsson 507 (NY, US, VEN); Ptari-tepui, 1600 m, 11<br />
the margin at, below, or above the middle of the Nov 1964 (st), Steyermark 60002 (F, VEN); between<br />
lamina; intercostal venation pl<strong>an</strong>e to prominent; S<strong>an</strong>ta Teresita de Kav<strong>an</strong>ayen <strong><strong>an</strong>d</strong> Rio Pacairao, Quebrada<br />
petiole 4-17 cm long; stipules 0.5-3.5 cm Oparu-ma, 1065-1220 m, 20 Nov 1944 (9), Steylong.<br />
ermark 60432 (F, VEN); between Rio Paramichi <strong><strong>an</strong>d</strong><br />
Staminate inflorescences <strong>with</strong> the heads up to 6 Salto de Chalim<strong>an</strong>o, NW of Serr<strong>an</strong>ia Pia-soi, 5 J<strong>an</strong><br />
mm diam.; common peduncle up to 8 cm long. 1962 (9), Steyermark 90583 (US, VEN); Rio Parami-<br />
Pistillate inflorescences mostly <strong>with</strong> one head, chi, between mouth of Rio Paramichi <strong><strong>an</strong>d</strong> Salto de<br />
more th<strong>an</strong> 1 cm, in fruit up to 3 cm diam.; (com- Chalim<strong>an</strong>o, 8-9 J<strong>an</strong> 1962 (9), Steyermark 90736 (K,<br />
mon) peduncle 1.5-9.5 cm<br />
NY, US); Rio Cuyuni, 140-142 km S of El Dorado,<br />
long.<br />
1300-1380 m, 22-28 Dec 1970 (9), Steyermark et al.<br />
Distribution (Fig. 6). Amazon Basin (Bolivia, 104348 (MY, NY). DELTA AMACURO: Cariapo, 13 Dec<br />
Brazil, Peru, Ecuador, Colombia, <strong><strong>an</strong>d</strong> Venezue- 1953 (9), Little et al. 15990 (VEN); Dto. Antonio Diaz,<br />
la), western Gui<strong>an</strong>a region (Guy<strong>an</strong>a, Venezuela),<br />
C<strong>an</strong>fo Guiniquina, 5 Feb 1977 (6), Marc<strong>an</strong>o-Berti et al.<br />
84-22-77<br />
<strong><strong>an</strong>d</strong> in the central <strong><strong>an</strong>d</strong> Pacific coastal part of Co-<br />
(U); Rio Cuyubini, above Casa Cuyubini, 12<br />
Nov 1960 (9), Steyermark 87480 (G, GH, NY, U, US,<br />
lombia, as far north as P<strong>an</strong>ama; in upl<strong><strong>an</strong>d</strong> <strong><strong>an</strong>d</strong> VEN); Dto. Antonio Diaz, Caio Htioba, tributary of<br />
riverine forests up to 2000 m.<br />
Rio Araquoa, 16 Oct 1977 (st), Steyermarket al. 114759
<strong>Coussapoa</strong> 37<br />
FIG. 14. <strong>Coussapoa</strong> asperifolia ssp. magnifolia: 1, leafy twig <strong>with</strong> pistillate inflorescences (Harling et al.<br />
14733); 2, staminate inflorescence (Garcia-Barriga 16239).
38 Flora Neotropica<br />
(MO); Dto. Antonio Diaz, upper part of Cafno Jobure, 4c. <strong>Coussapoa</strong> asperifolia ssp. rhamnoides<br />
7 Apr 1955 (2), Wurdack 286 (F, NY).<br />
(St<strong><strong>an</strong>d</strong>ley) Akkerm<strong>an</strong>s & C. C. Berg, Proc. Kon.<br />
GUYANA. Upper Rupununi River, nr. Dad<strong>an</strong>awa,<br />
24-29 Jul 1922 (9), De la Cruz 1729<br />
Ned. Akad. Wetensch. Ser.<br />
(F, GH, MO, NY);<br />
C, 85(4): 445.<br />
Mazaruni station, 7 May 1943 (2), F<strong>an</strong>shawe 1278 (FD 1982. Fig. 15.<br />
4014) (K, NY); Essequibo River, Tiger Creek, 3 Jun<br />
1943 (6), F<strong>an</strong>shawe 1337 (FD 4073) (K, NY); Esse- <strong>Coussapoa</strong> rhamnoides St<strong><strong>an</strong>d</strong>ley, Publ. Field Mus. Bot.<br />
quibo River, Bartica, 8 Jul 1952 (6), F<strong>an</strong>shawe 3409 17: 166. 1937. Type. Brazil. Amazonas: Mun. Sao<br />
(FD 6973) (K, NY); Pomeroon River, Kabakaburi, 19 Paulo de Olivenca, 11 Sep-26 Oct 1936 (9), Krukoff<br />
Aug 1918 (2), Hohenkerk s.n. (FD 738) (K, OXF); Ber- 8406 (holotype, NY; isotypes, A, BM, F, G, K, LE,<br />
bice River, Apr 1889 (2), Jenm<strong>an</strong> 4947A (K); Deme- MO, S, U).<br />
rara River, - (8), Parker s.n. (K); Ekaria, Rosedale,<br />
Aug 1924 (2), Persaud 116 (B, F, K, NY).<br />
Lamina obovate to subobovate or elliptic to<br />
ECUADOR. NAPO: 3 mi SW of Tena, 28 Aug 1960 oblong, 7-16 cm long, base obtuse to rounded<br />
(2), Grubb et al. 1498 (K, NY). PASTAZA: Nr. Mera, (to subcordate); upper surface smooth <strong><strong>an</strong>d</strong> glaca.<br />
1000 m, 10 Dec 1955 (9), Asplund 18780 (S); between<br />
Palora <strong><strong>an</strong>d</strong> Shell, 4 Mar 1980 (st), Berg & Akbrous;<br />
lower surface on the main veins sparsely<br />
kerm<strong>an</strong>s 1119 (U), (9), Berg & Akkerm<strong>an</strong>s 1133 (U); minutely puberulous <strong>with</strong> straight hairs; lateral<br />
nr. Mera, 17 J<strong>an</strong> 1977 (9), Harling et al. 14733 (GB), veins 4-6 pairs, unbr<strong>an</strong>ched or some of the lat-<br />
18 Mar 1940 (8), Lugo 83 (S).<br />
eral veins (poorly) br<strong>an</strong>ched, the basal pair un-<br />
PERU. Without locality (2), Ruiz & Pavon s.n. (or br<strong>an</strong>ched or faintly br<strong>an</strong>ched, reaching the mar-<br />
4) (B, BM, F, FI, US, type collection of C. magnifolia<br />
<strong><strong>an</strong>d</strong> C. ruizii). JUNiN: La Merced, 10-24 Aug 1923 gin above the middle of the lamina; petiole 2-7<br />
(2),<br />
Macbride 5447 (F, C). LORETO: Prov. Maynas, Pefia cm long; stipules 0.5-2 cm long. Pistillate inflo-<br />
Negra, 25 km SW of Iquitos, 1 Aug 1972 (st), Croat rescences <strong>with</strong> (1-)3-5 (sometimes partly fused)<br />
18643 (MO, NA); Prov. Requena, Rio Ucayali, Jenaro heads, 3-6 mm, in fruit up to 15 mm diam.;<br />
Herrera, below Requena, 9 Dec 1977 (2), Gentry et al.<br />
21319 (MO, U); Prov. Maynas, nr. Quistococha, 26<br />
(common) peduncle 1-4 cm long.<br />
May 1978 (6), Gentry et al. 22280 (MO, U); Prov.<br />
Distribution (Fig. 6). Amazon Basin (Brazil,<br />
Maynas, Rio N<strong>an</strong>ay, below Bellavista, 31 May 1972 Peru); in upl<strong><strong>an</strong>d</strong> <strong><strong>an</strong>d</strong> riverside (varzea) forest.<br />
(6), McD<strong>an</strong>iel 16092 (NA); Prov. Maynas, Dist. Iquitos,<br />
Puerto Almendra, Rio N<strong>an</strong>ay, 1 Nov 1976 (6), Specimens studied. PERU. LORETO: Rio Tacsha<br />
Revilla 1260 (MO, U). SAN MARTIN: Prov. Marichal Curaray, 18 Sep 1972 (2), Croat 20370 (F, GH, NA,<br />
Caceres, Dist. Tocache Nuevo, Fundo Gr<strong>an</strong> Chaparral, NY, P, RB).<br />
11 Nov. 1975 (6), Schunke 8666 (U); Prov. Mo- BRAZIL. AMAZONAS: Rio Solimoes, Rio Bia, affluyobamba,<br />
1906 (st), Weberbauer 4472 (G).<br />
ent of Rio Jutai, 5 Nov 1975 (Q), L. Coelho et al. 332<br />
BRAZIL. ACRE: Cruzeiro do Sul, 2 Mar 1976 (2), (U); Barcelos, 7 Sep 1962 (Q), Duarte et al. 6970 (RB);<br />
Marinho 348 (IAN). AMAZONAS: Rio Negro, Sao Ga- Mun. Sao Paulo de Olivenca, nr. Palmares, 11 Sep-26<br />
briel de Cachoeira, 28 Feb 1975 (9), Cordeiro 367 (IAN); Oct 1936 (2), Krukoff8406 (A, BM, F, G, K, LE, MO,<br />
mouth of Rio Caiari, affluent of Rio Negro, 15 Sep NY, S, U, type collection of C. rhamnoides); M<strong>an</strong>aus-<br />
1952 (?), Fro6s et al. 28585 (IAN); mun. Humaiti, Itacoatiara road, Res. Flor. Ducke, 1 Sep 1966 (2),<br />
nr. Livramento on Rio Livramento, 12 Oct-6 Nov Pr<strong>an</strong>ce et al. 2153 (INPA, NY, U). PARA: Rio Tapajos,<br />
1934 (2), Krukoff6662 (A, B, BM, F, G, K, LE, MO, Periquito, 27 Jul 1923 (2), Ducke (HJBR) 18460 (RB).<br />
NY, RB, S, U, US); nr. M<strong>an</strong>aus, 1906 (2) Lambroy s.n.<br />
(P); Rio Negro, U<strong>an</strong>ari, nr. Sao Gabriel, 31 Oct 1947<br />
(6), Pires 805 (IAN, NY), 2 Nov 1947 (9), Pires 830 5. <strong>Coussapoa</strong> batavorum Akkerm<strong>an</strong>s & C. C.<br />
(NY); Rio Negro, between Sao Gabriel <strong><strong>an</strong>d</strong> Sao Felipe, Berg, Proc. Kon. Ned. Akad. Wetensch. Ser.<br />
base of Serra U<strong>an</strong>ari, Oct 1947 (6), Schultes & Pires C, 85(4): 447. 1982.<br />
8976 (GH, U); Rio Negro, Sao<br />
Type. Colombia. Valle:<br />
Gabriel de Cachoeira,<br />
J<strong>an</strong>-Aug 1852 (6), Spruce 2260 (BR, K, P). MATO<br />
Nr. Buenaventura, 5 Dec 1979 (9), V<strong>an</strong> Rood-<br />
GRosso: Rio Aripuafia, nr. Humboldt Center, 59?21'N, en et al. 700 (holotype, COL; isotypes, AAU,<br />
10?12'S, 19 Oct 1973 (st), Berg s.n. (U); Rio Juruena, K, NY, U, US). Fig. 16.<br />
airport, 17 Jul 1977 (9), M. G. Silva et al. 3368 (U).<br />
PARA: Almeirim, Barreira da Velha Pobre, 6 Jul 1919 Tree, terrestrial, ca. 15 m tall. Leafy twigs 10-<br />
(2), Ducke (HBJR) 13605 (RB); Rio Br<strong>an</strong>co de Obidos, 19 mm<br />
21 Jul 1918 (6), Ducke (HAMP) 17132 thick, glabrous. Lamina coriaceous,<br />
(MG).<br />
RONDONIA: Nr. Porto Velho, 28 Jun 1952 (9), J. F. broadly ovate to cordiform, 36-40 x 34-37 cm,<br />
Silva 227 (IAN); Rio Madeira, Cachoeira Misericorde, apex obtuse to emarginate, base cordate, margin<br />
Riberao, 2 Aug 1968 (9), Pr<strong>an</strong>ce et al. 6718 (NY, U). subcrenate; both surfaces glabrous; lateral veins<br />
RORAIMA: West facing slopes of Serra Tepequem, 1200 10-13<br />
m, 20 Feb 1967 (2), Pr<strong>an</strong>ce et al. 4588<br />
pairs, 3-5 of which arise at the base of the<br />
(NY, U).<br />
BOLIVIA. LA PAZ: Prov. S. Yungas, basin of Rio midrib, straight, main basal pair br<strong>an</strong>ched,<br />
Bopi, S<strong>an</strong> Bartolome, nr. Calisaya, 1-22 Jul 1939 (2), reaching the margin at or above the middle of<br />
Krukoff 10123 (A, F, G, K, MO, NY, S, U). the lamina; intercostal venation almost pl<strong>an</strong>e to
<strong>Coussapoa</strong> 39<br />
FIG. 15. <strong>Coussapoa</strong> asperifolia ssp. rhamnoides: 1, leafy twig <strong>with</strong> pistillate inflorescences (Krukoff 8406).<br />
slightly prominent; petiole 10-12 cm long, dis-<br />
tinctly ribbed <strong><strong>an</strong>d</strong> densely minutely puberulous;<br />
stipules 8-10 cm long, densely to sparsely red-<br />
dish-brown puberulous <strong><strong>an</strong>d</strong> <strong>with</strong> longer straight<br />
white hairs. Staminate inflorescences unknown.<br />
Pistillate inflorescences unbr<strong>an</strong>ched or poorly<br />
br<strong>an</strong>ched; heads 1-2(-3), free or partly fused,<br />
globose to ovoid, ca. 8-10 mm, in fruit up to 27<br />
mm diam.; common peduncle 0.5-2 cm long,<br />
rather densely puberulous to hirtellous; peri<strong>an</strong>th<br />
1 cm<br />
1(-3) mm high, white puberulous; fruiting peri-<br />
<strong>an</strong>th brown. Interfloral bracts absent.<br />
Distribution (Fig. 6). Colombia (Choco); in<br />
riverside forest. Known only from the type col-<br />
lection.<br />
Knowledge of the staminate inflorescence is<br />
required in order to determine the relationships<br />
of this species. In the shape, size, <strong><strong>an</strong>d</strong> sparse<br />
indumentum of the lamina, it resembles C. as-<br />
perifolia ssp. magnifolia as well as forms of C.
40 Flora Neotropica<br />
FIG. 16. <strong>Coussapoa</strong> batavorum: 1, leafy twig <strong>with</strong> pistillate inflorescences<br />
(V<strong>an</strong> Rooden et al. 700).
<strong>Coussapoa</strong><br />
v<strong>an</strong>nifolia having leaves <strong>with</strong> a sparse indumen- NY); Cocle del Norte, 23 Aug 1978 (2), Hammel 4475<br />
tum.<br />
(U); ca. 11 km SW of Cerro Braja, 9?25'N, 79?35'W,<br />
2 May 1981 (8), Sytsma et al. 4225 (U). PANAMA: Cerro<br />
Jefe, 3 Mar 1979 (8), d'Arcy 12329 (U); road to Cerro<br />
6. <strong>Coussapoa</strong> brevipes Pittier, Contr. U.S. Natl. Camp<strong>an</strong>a, 3 km from Interameric<strong>an</strong> Highway, 18 Sep<br />
Herb. 18:225. 1917; Woodson & Schery, Ann. 1968 (6), Correa et al. 1031A (DUKE, F, MO); Cerro<br />
Missouri Bot. Gard. 47:171. 1960. Type. P<strong>an</strong>- Camp<strong>an</strong>a, 9 Apr 1970 (9), Croat 14212 (GH, MO);<br />
amf. S<strong>an</strong> Bias: Nr. Puerto Obaldia, hills of<br />
Cerro Jefe region, La Eneida, 25 Mar 1968 (2), Dressier<br />
3454 (DUKE,<br />
Sperdi, Sep 1911<br />
MO); 10 km N of<br />
(v), Pittier 4386<br />
Margarita, on road<br />
(holotype, to Madronio, 31 J<strong>an</strong> 1979 (8), Hammel 6006 (U); 3 mi<br />
US; isotype, B). Fig. 17. NE of Altos de Pacora, Campo Tres, 10 Mar 1973 (6),<br />
Liesner 515 (K, MO, NY, US); Altos de<br />
Tree, terrestrial or<br />
Camp<strong>an</strong>a, 28<br />
hemi-epiphytic, up to 15 m May 1977 (8), Mendez 15 (MO); 6.5 km N of Lago<br />
tall. Leafy twigs 4-11 mm thick, more or less Azul, 13 J<strong>an</strong> 1974 (9), Nee 9281 (MO, NY, U); El<br />
densely white (appressed-)puberulous <strong><strong>an</strong>d</strong> <strong>with</strong> Ll<strong>an</strong>o-Carti road, 8 mi from P<strong>an</strong> Americ<strong>an</strong> Hwy., 17<br />
reddish-brown pluricellular hairs, or also <strong>with</strong> Apr 1981 (2), Sytsma 4024 (U); Cerro Camp<strong>an</strong>a, 18<br />
sparse arachnoid hairs.<br />
Sep 1968<br />
Lamina<br />
(a), Weaver et al. 1677<br />
coriaceous, ovate<br />
(DUKE). SAN BLAS:<br />
Road El Ll<strong>an</strong>o-Carti-Tupile, 1 mile from continental<br />
to elliptic or occasionally obovate, 5-30 x 2.5- divide, 30 Mar 1973 (9), Liesner 1309 (DUKE, GH,<br />
18 cm, apex obtuse to acute or to subcordate, MO, NY, US); nr. Puerto Obaldia, hills of Sperdi, Sep<br />
base obtuse to rounded to subcordate, sometimes 1911 (9), Pittier 4386 (B, US, type collection); Rio Acla,<br />
acute, margin entire or subcrenate towards the 8048'38"N, 77040'30"W, 7 Feb 1979 (2), Sugden 415<br />
(K).<br />
apex; upper surface glabrous, lower surface appressed-puberulous,<br />
densely so in the areoles, <strong>Coussapoa</strong> brevipes shows morphological af<strong><strong>an</strong>d</strong><br />
on the main veins often <strong>with</strong> white arach- finities especially <strong>with</strong> C. chocoensis but also <strong>with</strong><br />
noid hairs which are soon deciduous; lateral veins C. contorta.<br />
6-18 pairs, straight or slightly curved, basal pair<br />
br<strong>an</strong>ched, reaching the margin below the middle 7. <strong>Coussapoa</strong> chocoensis Cuatrecasas, Caldasia<br />
of the lamina; intercostal venation almost pl<strong>an</strong>e; 7:287.1956. Type. Colombia. Choc6: Rio S<strong>an</strong><br />
petiole 1-7 cm long, 2-4 mm thick, appressed- Ju<strong>an</strong>, Quebrada del Taparal, 30 May 1946 (2),<br />
puberulous <strong><strong>an</strong>d</strong> often also <strong>with</strong> white arachnoid Cuatrecasas 21446 (holotype, F). Fig. 18.<br />
hairs which are infrequently deciduous; stipules<br />
3-10 cm<br />
Tree,<br />
long, densely covered <strong>with</strong><br />
hemi-epiphytic. Leafy twigs 3-6 mm<br />
appressed<br />
white<br />
thick, brownish subhirsute to hirtellous.<br />
puberulous hairs, mixed<br />
Lamina<br />
<strong>with</strong> reddishbrown<br />
pluricellular hairs <strong><strong>an</strong>d</strong> often also<br />
coriaceous, ovate to<br />
<strong>with</strong><br />
elliptic, 8-16 x 4-12, apex<br />
sparse to dense white arachnoid hairs <strong><strong>an</strong>d</strong>/or<br />
shortly acuminate, base rounded to subcordate,<br />
long<br />
yellowish straight hairs.<br />
margin<br />
Staminate<br />
entire; upper surface glabrous, lower surinflorescences<br />
face<br />
br<strong>an</strong>ched; heads 4-20, globose, ca. 4-5 mm<br />
densely appressed-puberulous in the areoles,<br />
diam.;<br />
common<br />
sparsely puberulous to hirtellous on the smaller<br />
peduncle 0.5-3 cm long, densely covered<br />
<strong>with</strong> short<br />
veins,<br />
white puberulous <strong><strong>an</strong>d</strong><br />
densely subhirsute,<br />
reddishsubvelutinous,hirtelbrown<br />
lous,<br />
pluricellular hairs; peri<strong>an</strong>th ca. 1<br />
strigose or<br />
mm<br />
strigillose on the main veins;<br />
high, lateral veins 10-12 pairs, curved <strong><strong>an</strong>d</strong><br />
densely minutely puberulous; stamens three, far<br />
distinctly<br />
exceeding the<br />
looping, basal<br />
peri<strong>an</strong>th. Pistillate<br />
pair unbr<strong>an</strong>ched or<br />
inflorescences<br />
sparsely<br />
unbr<strong>an</strong>ched; heads globose to<br />
br<strong>an</strong>ched,<br />
ellipsoid, ca. 5-7<br />
reaching the margin below the middle<br />
of the<br />
mm, in fruit up to 14 mm diam.; peduncle 0.4lamina;<br />
intercostal venation rather prom-<br />
1.5(-2.5) cm long, densely white<br />
inent; petiole 1-3 cm<br />
puberulous <strong><strong>an</strong>d</strong><br />
long, brown subhirsute to<br />
also <strong>with</strong> reddish-brown<br />
subvelutinous; stipules 1.5-2.5 cm<br />
pluricellular<br />
long, brown<br />
hairs; peri<strong>an</strong>th<br />
ca. 1-2 mm<br />
(sub)hirsute. Staminate inflorescences unknown.<br />
high, densely minutely pub- Pistillate inflorescences<br />
erulous; fruiting peri<strong>an</strong>th or<strong>an</strong>ge. Interfloral bracts<br />
unbr<strong>an</strong>ched; head (globose<br />
or<br />
absent.<br />
ellipsoid?), ca. 15 mm diam.; peduncle<br />
0.5-1 cm<br />
Distribution (Fig. 6). P<strong>an</strong>ama, in forests up to<br />
long, hirtellous to subhirsute; peri<strong>an</strong>th<br />
ca. 2 mm<br />
800 m.<br />
high, minutely brown puberulous. Interfloral<br />
bracts subspathulate, hirtellous at the<br />
Specimens studied. PANAMA: COCLE: Nr. La Mesa, apex.<br />
4 km from El Valle, 15 Sep 1968 (Y), Correa et al. 1002<br />
(DUKE, F); foothills of Cerro Pil6n, nr. El Valle, 5 Oct Distribution (Fig. 6). Colombia (Choc6); low-<br />
1967 (Q), Duke et al. 14712 (MO, US). COLON: Nr. l<strong><strong>an</strong>d</strong>, riverside. Known only from the type col-<br />
Guasimo, 22 Apr 1970 (Q), Croat 9973A (F, GH, MO, lection.<br />
41
A'<br />
..<br />
.<br />
FIG. 17. <strong>Coussapoa</strong> brevipes: 1, leafy twig <strong>with</strong> staminate inflorescences (Weaver & Foster 1677); 2, pistillate in<br />
11<br />
c1m<br />
I
'?<br />
' Cous': '.' .. . . . c e :1..<br />
~~~~~~~~~~~~~~~~~~~~~~~~~~~?~<br />
8~~~~~~~~~~~~~~~<br />
\~~~~~~~~~~~~~~~~~~~~~? '' ;:<br />
?~. .: .. :7<br />
?:::<br />
.:; 0 .-; - :. .<br />
i'1'::" .. - .<br />
,~~ ~ j. 'i :<br />
'<br />
?'"'",...?:<br />
." :,,; ' .?<br />
.<br />
iI.' ;... ?. ?<br />
':I~~~<br />
?.:.;?...;<br />
?~~~~~<br />
~~~~'"' (.:??<br />
"~~~~~~~~~~~~~~~~~~~'<br />
. . i,:~..~~~~~~~~~~~~~~~~~~<br />
....:<br />
"' ~~~~~. ?'"....:',,:<br />
?.;t~' .<br />
'-~~~~ ' ~ ~~~~~~~I<br />
~~~~~~~~~~~~~~~~<br />
. '? ~~~~~~~~~~~~~<br />
::i.: ?<br />
'<br />
.,. ':." ::'"..?..<br />
''(...'".1 ~~? '"!;:"i,~'i ~~~~~~~ ~<br />
FIG. 18. <strong>Coussapoa</strong> chocoensis: 1, leafy twig <strong>with</strong> pistillate inflorescences (Cuatrecasas 214<br />
~<br />
.. \ ..?. . .<br />
FIG. 18. <strong>Coussapoa</strong> chocoensis: , leafy twig <strong>with</strong> pistillate iniorescences (Cuatrecasas<br />
~ cm
44 Flora Neotropica<br />
This species resembles C. brevipes, from which<br />
it clearly differs in the brown indumentum <strong><strong>an</strong>d</strong><br />
the presence of interfloral bracts.<br />
8. <strong>Coussapoa</strong> cinnamomea Cuatrecasas, Calda-<br />
sia 7: 288. 1956. Type. Colombia. Amazonas:<br />
Trapecio Amaz6nico, Rio Loretoyacu, 28 Sep<br />
1946 (9), Schultes & Black 8269 (=Black &<br />
Schultes 46-137) (holotype, F; isotypes, A,<br />
IAN, K, US). Fig. 19.<br />
C. cinnamomea has m<strong>an</strong>y features in common<br />
<strong>with</strong> C. tessm<strong>an</strong>nii, but in most parts is larger<br />
<strong><strong>an</strong>d</strong> more robust th<strong>an</strong> the latter species. The two<br />
species appear to be closely related.<br />
9. <strong>Coussapoa</strong> cinnamomifolia Mildbraed, No-<br />
tizbl. Bot. Gart. Berlin 15: 783. 1942. Type.<br />
Ecuador. Pastaza: Nr. Mera, ca. 1000 m, 28<br />
Oct 1938 (i), Schultze-Rhonhof 2943 (holo-<br />
type, B). Fig. 20.<br />
Tree, mostly terrestrial, up to 35 m tall. Leafy<br />
twigs 2-5 mm thick, densely brownish appressed-puberulous,<br />
glabrescent. Lamina coriaceous,<br />
oblong to subovate or subobovate, 3-13<br />
x 1-5 cm, apex (sub)acute to obtuse, base obtuse<br />
to acute, margin entire; upper surface glabrous,<br />
lower surface glabrous or appressed-puberulous<br />
in the lower part; lateral veins 3-6 pairs, the basal<br />
pairs strong, unbr<strong>an</strong>ched, reaching the margin<br />
far below the middle of the lamina, the other<br />
lateral veins weak <strong><strong>an</strong>d</strong> inconspicuous; intercostal<br />
venation pl<strong>an</strong>e; petiole 1-2 (or -3) cm long,<br />
densely brownish appressed-puberulous; stipules<br />
0.5-2.5 cm long, brownish subsericeous to subvelutinous.<br />
Staminate<br />
Tree, hemi-epiphytic or terrestrial, up to 8 m<br />
inflorescences poorly<br />
tall. Leafy twigs 9-22 mm thick, brownish<br />
br<strong>an</strong>ched; heads 2-7, partly fused, globose, ca.<br />
2-3 mm<br />
(sub)velutinous, glabrescent. Lamina coria- diam.; common peduncle 1.5-3 cm long,<br />
ceous, (broadly) ovate, 16-38 x<br />
brownish<br />
12-28 puberulous, peri<strong>an</strong>th ca. 1 mm high,<br />
cm, apex<br />
obtuse to (acute), base rounded to truncate or to sparsely minutely puberulous; stamens three, just<br />
cordate, margin subcrenate; upper surface exceeding the peri<strong>an</strong>th. Pistillate inflorescences<br />
glaunbr<strong>an</strong>ched<br />
or<br />
brous, lower surface densely puberulous in the<br />
poorly br<strong>an</strong>ched; heads 1-2,<br />
sometimes partly fused, globose, ca. 4-7 mm, in<br />
areoles, (rather) sparsely appressed puberulous<br />
fruit<br />
to substrigose on the veins; lateral veins 14-22 up to 9 mm diam.; (common) peduncle 2-<br />
5 cm<br />
pairs, (almost) straight, basal pair br<strong>an</strong>ched,<br />
long, brownish puberulous; peri<strong>an</strong>th ca. 1<br />
mm<br />
reaching the margin below the middle of the lam- high, glabrous; fruiting peri<strong>an</strong>th yellow. Inina;<br />
intercostal venation terfloral bracts spathulate to<br />
(almost) pl<strong>an</strong>e; petiole<br />
subpeltate or pel-<br />
5-15 cm long, 5-7 mm<br />
tate.<br />
thick; shortly subvelu-<br />
Distribution<br />
tinous, also brown pluricellular hairs. Staminate<br />
(Fig. 6). Known in Colombia<br />
inflorescences unknown. Pistillate (Antioquia), Ecuador (Pastaza), <strong><strong>an</strong>d</strong> Bolivia<br />
inflorescences<br />
(La<br />
br<strong>an</strong>ched; heads 2-4, often partly fused, sub- Paz); in forest at altitudes between (400-?) 800<br />
<strong><strong>an</strong>d</strong><br />
globose, in<br />
1200 m.<br />
fruit up to 40 mm in diam.; common<br />
peduncle 2-4 cm long, peduncle <strong><strong>an</strong>d</strong> br<strong>an</strong>ches<br />
Specimens studied. COLOMBIA. ANTIOQUIA: Mun.<br />
6-10(-15) mm thick, shortly velutinous; peri- S<strong>an</strong> Luis, rd. Medellin-Bogota, quebrada La Tebaida,<br />
<strong>an</strong>th glabrous. Interfloral bracts absent. 1000-1100 m, 22 Jun 1987 (2), Callejas et al. 3997<br />
Distribution (Fig. 6). Colombia (Amazonas, (BG); nr. S<strong>an</strong> Luis, 900-1000 m, 21 Jun 1982 (6), Her-<br />
Trapecio Amaz6nico); riverside forest.<br />
n<strong><strong>an</strong>d</strong>ez et al. 369 (HUA).<br />
ECUADOR. PASTAZA: Nr. Mera, ca. 1000 m, 14<br />
Specimens studied. COLOMBIA. AMAZONAS: Tra- Feb 1955 (9), Asplund 18840 (S), 28 Oct 1938 (6),<br />
pecio Amaz6nico, Rio Loretoyacu, 22 Nov 1945 (2), Schultze-Rhonhof2943 (B, type collection).<br />
Duque-Jaramillo 2179 (COL), Oct 1945 (2), Schultes BOLIVIA. LA PAZ: Prov. Larejaca, Copacab<strong>an</strong>a, 10<br />
6693 (F, GH, U, US), 28 Sep 1946 (2), Schultes & Black km S of Mapiri, 850-950 m, 8 Oct-15 Nov 1939 (6),<br />
8269 (=Black & Schultes 46-137) (F, GH, IAN, K, US, Krukoff 11275 (A, F, G, K, MO, NY, S, U, US), (9)<br />
type collection).<br />
Krukoff 11276 (A, F, G, MO, S, U, US).<br />
In m<strong>an</strong>y characters, <strong><strong>an</strong>d</strong> especially in the sub-<br />
trinervate lamina, C. cinnamomifolia is similar<br />
to C. trinervia. The 3-staminate flowers, how-<br />
ever, suggest relationships <strong>with</strong> other species, in<br />
particular <strong>with</strong> C. parviceps.<br />
The Colombi<strong>an</strong> collection differs somewhat<br />
from the other collections in the longer petiole<br />
(up to 3 cm) <strong><strong>an</strong>d</strong> peltate <strong><strong>an</strong>d</strong> larger bracts.<br />
10. <strong>Coussapoa</strong> contorta Cuatrecasas, Caldasia 7:<br />
289. 1956. Type. Colombia. Valle: Pacific<br />
Coast, Rio Naja, braza Aji, Calle Larga, 28<br />
Feb 1943 (6), Cuatrecasas 14284 (holotype,<br />
F). Fig. 21.
<strong>Coussapoa</strong><br />
"~ [<br />
FIG. 19. <strong>Coussapoa</strong> cinnamomea: 1, leafy twig <strong>with</strong> pistillate inflorescences (Schultes 6693 (inflorescences)<br />
<strong><strong>an</strong>d</strong> Schultes & Black 8269 (leaf <strong><strong>an</strong>d</strong> stipules)).<br />
1 cm<br />
45
46 Flora Neotropica<br />
1 cm<br />
FIG. 20. <strong>Coussapoa</strong> cinnamomifolia: 1, leafy twig <strong>with</strong> pistillate inflorescences (Krukoff11276); 2, staminate<br />
inflorescences (Krukoff 11275).<br />
Shrub or tree, hemi-epiphytic or terrestrial, up<br />
to 20 m tall. Leafy twigs 3-6 mm thick, yellowish<br />
appressed-puberulous or sometimes partly hirtellous.<br />
Lamina chartaceous to coriaceous, ellip-<br />
tic or oblong to ovate or obovate, 5-26 x 1.5-<br />
13 cm, apex acute to acuminate, base acute to<br />
obtuse <strong><strong>an</strong>d</strong> occasionally to cordate, margin en-<br />
tire; upper surface glabrous, lower surface on the
<strong>Coussapoa</strong><br />
X { cm<br />
FIG. 21. <strong>Coussapoa</strong> contorta: 1, leafy twig <strong>with</strong> pistillate inflorescences (Javita & Epling 1097); 2, leafy twig<br />
<strong>with</strong> staminate inflorescences (Cuatrecasas 14284).<br />
main veins rather sparsely appressed-puberulous<br />
<strong><strong>an</strong>d</strong> sometimes, only in the axils of the lateral<br />
veins on more or less distinct domatium-like<br />
cavities, distinctly longer stiff yellowish hairs, on<br />
the smaller veins rather sparsely puberulous to<br />
glabrous; lateral veins 6-15 pairs, slightly curved,<br />
basal pair unbr<strong>an</strong>ched, reaching the margin below<br />
the middle of the lamina; intercostal vena-<br />
47
48 Flora Neotropica<br />
tion pl<strong>an</strong>e; petiole 1-8 cm long, appressed-pu- small, ca. I cm long stipules, the basal pair of<br />
berulous; stipules 2-4 cm long, yellowish lateral veins reaching the margin of the lamina<br />
appressed-puberulous to (sub)sericeous. Stami- at about the middle, <strong><strong>an</strong>d</strong> the rather dense brownnate<br />
inflorescence poorly br<strong>an</strong>ched; heads 6-10, ish hairs on the peri<strong>an</strong>th of the pistillate flower.<br />
globose, ca. 2-3 mm diam.; common peduncle It could represent a distinct species or subspecies.<br />
1-3 cm long, densely puberulous to hirtellous;<br />
peri<strong>an</strong>th ca. 1 mm high, minutely puberulous;<br />
stamens three, far exceeding the peri<strong>an</strong>th. Pis- 11. <strong>Coussapoa</strong> crassivenosa Mildbraed, Notizbl.<br />
tillate inflorescences mostly unbr<strong>an</strong>ched, occa- Bot. Gart. Berlin 15: 784. 1942. Type. Ecuasionally<br />
br<strong>an</strong>ched; heads 1-2, globose, 5-7 mm, dor. Pastaza: Nr. Mera, ca. 1000 m, 7 Sep 1938<br />
in fruit up to 10 mm diam.; (common) peduncle (a), Schultze-Rhonhof 2783 (holotype, B).<br />
1-2 cm long, densely puberulous to hirtellous;<br />
Fig. 22.<br />
peri<strong>an</strong>th 1-2 mm high, minutely puberulous. Interfloral<br />
bracts peltate, relatively large, densely<br />
<strong>Coussapoa</strong> steyermarkii Cuatrecasas, Fieldi<strong>an</strong>a<br />
Bot.<br />
puberulous.<br />
28(1): 212. 1951. Type. Venezuela. Bo-<br />
Distribution (Fig. 6). Colombia <strong><strong>an</strong>d</strong><br />
livar:<br />
Ecuador,<br />
Ptari-tepui, 1600 m, 10 Nov 1944 (Y),<br />
Pacific coastal region, also in Costa Rica?; in Steyermark 60002a (holotype, F; isotype,<br />
forest up to 1500 m altitude.<br />
VEN).<br />
Specimens studied. COLOMBIA. CAUCA?: Tree, often<br />
Juntas,<br />
hemi-epiphytic, up to 20 m tall.<br />
- (2), Lehm<strong>an</strong>n BT694 (A, GH, K, L, NY). CHocO: Leafy twigs 6-9 mm thick, densely (pale) brown<br />
Rio S<strong>an</strong> Ju<strong>an</strong>, Quebrada de Taparal, 30 May 1946 (6), puberulous to hirtellous or to subhirsute to stri-<br />
Cuatrecasas 21496 (F); Rio S<strong>an</strong> Ju<strong>an</strong>, between Que- gose. Lamina coriaceous, oblong to elliptic to<br />
brada La Sierpe <strong><strong>an</strong>d</strong> Quebrada El Quicharo, 27 Mar suborbicular to subobovate or to<br />
1979 (6), Forero et al. 4099 (COL, MO, U); Rio S<strong>an</strong><br />
(sub)ovate, 4-<br />
Ju<strong>an</strong>, nr. Palestina, 26 Mar 1979 (6), Forero et al. 4110 24(-43) x 2-14(-20) cm, apex acuminate to ob-<br />
(COL, MO, U); Rio S<strong>an</strong> Ju<strong>an</strong>, Quebrada de Taparal, tuse <strong><strong>an</strong>d</strong> to rounded, base obtuse to rounded (or<br />
5 Dec 1979 (9), V<strong>an</strong> Rooden et al. 699 (U). NARIRO: to acute), margin entire; upper surface glabrous,<br />
Mun. Iscu<strong><strong>an</strong>d</strong>6, Quebrada La Berrera, 28 Nov 1955 lower surface<br />
(6), Romero-Castaieda 5512 strigose to subsericeous on the main<br />
(COL). VALLE: Pacific<br />
Coast, Rio Naja, Calle Larga, 28 Feb 1943 (6), Cua- veins, sparsely (along margin of the lamina) putrecasas<br />
14284 (F, type collection); Pacific Coast, Rio berulous to hirtellous, in addition covered<br />
Cajambre, 5-15 May 1944 (9), Cuatrecasas 17647 (F). <strong>with</strong> (rather) dense, sometimes very sparse,<br />
ECUADOR. BOLIVAR: Valley of Rio Tablas, 1300 (sub)persistent<br />
m, 1<br />
(pale) brown arachnoid hairs; lat-<br />
Oct 1943 (st), Acosta Solis 6008 (F). CARCHI: Rio<br />
eral veins<br />
Bl<strong>an</strong>co, tributary of Rio S<strong>an</strong> Ju<strong>an</strong>, above Chical, 1300-<br />
(4-)5-8 pairs, slightly curved, basal<br />
1500 m, 25 Sep 1979 (Q), Gentry et al. 26525 (SEL, pair br<strong>an</strong>ched (or sometimes unbr<strong>an</strong>ched),<br />
U). ESMERALDAS: Rio Onzole, Estero Chontaduro, 14 reaching the margin at or above the middle of<br />
Jul 1966 (Q), Javita et al. 1097 (MO, NY, S, US). IM- the lamina; intercostal venation (very) promi-<br />
BABURA: Collapi, 4 Jul 1949 (2), Acosta Solis 12797<br />
(F). EL<br />
nent;<br />
ORO: Trail to<br />
petiole 1-4 cm<br />
Sambotambo, Rio Moro<br />
long, (puberulous to) sub-<br />
Moro,<br />
S of Buenaventura, highway to Portovele, 1035-1800 strigose to hirsute; stipules 1-5 cm long, brown<br />
m, 29 Aug 1943 (2), Steyermark 54214 (F). PICHINCHA: subsericeous to subhirsute. Staminate inflores-<br />
Nr. S<strong>an</strong>to Domingo de los Colorados, 30 Aug 1930 (6), cences br<strong>an</strong>ched; heads 4-20, globose, ca. 2-3<br />
Benoist 3010 (P); road S<strong>an</strong>to Domingo de los Colo- mm diam.; common peduncle 1-4.5 cm<br />
rados-Quininde (km 170-175), 2<br />
long,<br />
Sep 1949 (st), Acosta<br />
Solis 13687, 4 Sep 1949 (8), Acosta Solis 13713 (rather) densely puberulous to hirtellous to<br />
(F).<br />
strigillose;<br />
peri<strong>an</strong>th ca. 1 mm high, minutely brown<br />
The taxonomic relationships of C. contorta are puberulous; stamens two, far exceeding the perinot<br />
clear. Overall similarities suggest affinity <strong>with</strong> <strong>an</strong>th. Pistillate inflorescences br<strong>an</strong>ched; heads 3-<br />
C. brevipes, but a connection <strong>with</strong> C. oligoceph- 7, globose, 2-5 mm diam.; common peduncle 1-<br />
ala is also possible. The species is distinguished 3 cm long, brownish puberulous to hirtellous to<br />
by the domatium-like cavities in the axils of the strigillose; peri<strong>an</strong>th ca. 1-1.5 mm high, glabrous.<br />
lateral veins <strong><strong>an</strong>d</strong> by the large interfloral bracts. Interfloral bracts spathulate, often the lower part<br />
Collection S<strong>an</strong>chez & Zamora 512, from Cos- very narrow, minutely puberulous at the apex.<br />
ta Rica, S<strong>an</strong> Jose, below La Hondrua, Parque Distribution (Fig. 6). Presently known from<br />
Nacional Braulio Carrillo, 2 J<strong>an</strong> 1984 (2), (AAU) the following disjunct areas: the eastern Gui<strong>an</strong>a<br />
differs from the collections cited above in the region (Venezuela <strong><strong>an</strong>d</strong> northern Brazil), Ama-
<strong>Coussapoa</strong> 49<br />
I.<br />
FIG. 22. <strong>Coussapoa</strong> crassivenosa: 1, leafy twig <strong>with</strong> pistillate inflorescences (Steyermark et al. 104296); 2,<br />
leafy twig <strong>with</strong> staminate inflorescences (Krukoff 11085).<br />
zoni<strong>an</strong> Bolivia, Amazoni<strong>an</strong> Ecuador, <strong><strong>an</strong>d</strong> P<strong>an</strong>- NY); Meseta del Jaua, Cerro Sarisarinama, 700 m, 12ama;<br />
in forests, mostly between 700 <strong><strong>an</strong>d</strong> 15 Feb 1974 (6), Steyermark et al. 108988 (F, K, NY);<br />
1700 m.<br />
6 km N of Mision de S<strong>an</strong>ta Teresita de Kav<strong>an</strong>ayen,<br />
1100 m, 21 Feb 1978 (st), Steyermark et al. 115549<br />
(U, VEN).<br />
ECUADOR. MORONA-SANTIAGO: Cord. de Cutucui,<br />
Specimens studied. PANAMA. PANAMA: Road El 5-10 km from Logrono, 1200-1500 m, 7-9 Oct 1975<br />
Ll<strong>an</strong>o-Carti, 9'20'N, 78?50'W, 7 J<strong>an</strong> 1981 (d), Hahn (2), Little et al. 635 (COL, Q). PASTAZA: Road Puyo-<br />
347 (U).<br />
Tena, ca. 8 km, Tnte. Hugo Ortiz, ca. 950 m, 4 Mar<br />
VENEZUELA. AMAZONAS: Brazili<strong>an</strong> border, ca. 1980 (8), Berg & Akkerm<strong>an</strong>s 1120 (U); between Palora<br />
2?27'24'N, 63056'W, 22 May 1972 (a), Steyermark <strong><strong>an</strong>d</strong> Shell, ca. 900 m, 4 Mar 1980 (6), Berg & Akker-<br />
106118 (U, VEN). BoLnvAR: Meseta del Jaua, Cerro m<strong>an</strong>s 1129 (st), Berg& Akkerm<strong>an</strong>s 1131 (U); nr. Mera,<br />
Sarisarinama, 700 m, 10 Feb 1976 (a), Brewer-Carias ca. 1000 m, 7 Sep 1938 (8), Schultze-Rhonhof2783 (B,<br />
s.n. (COL, MY); Ptari-tepui, 1600 m, 10 Nov 1944 (2), type collection), 20 Nov 1938 (9), Schultze-Rhonhof<br />
Steyermark 60002a (F, VEN, type collection ofC. stey- 3006 (B).<br />
ermarkil); Quebrada O-paru-ma, between S<strong>an</strong>ta Ter- BRAZIL. RoRAIMA: Nr. Auaris, 4?3'N, 64?22'W,<br />
esita de Kav<strong>an</strong>ayen <strong><strong>an</strong>d</strong> Rio Pacairao, 1065-1220 m, 760-800 m, 7 Feb 1969 (6), Pr<strong>an</strong>ce et al 9679 (F, GH,<br />
20-21 Nov 1944 (st), Steyermark 60364 (F, VEN); K, NY, P, S, U, US); Serra Surucucu, 26 J<strong>an</strong> 1975 (e),<br />
Ptari-tepui, 1220 m, 28 Nov 1944 (st), Steyermark Ribeiro 612 (IAN, MG, RB), 23 J<strong>an</strong> 1975 (9), Rosa<br />
60673 (F); Chim<strong>an</strong>ta-tepui (Torono-tepui), 1700 m, 296 (IAN).<br />
21 May 1953 (9), Steyermark 75522 (F, NY, VEN); BOLIVIA. LA PAZ: Prov. Larecaja, Copacab<strong>an</strong>a, ca.<br />
Rio Cuyuni, ca. 142 km S of El Dorado, 1300-1380 10 km S of Mapiri, 850-950 m, 8 Sep-15 Nov 1939<br />
m, 22-28 Dec 1970 (2), Steyermark et al 104296 (MY, (8), Krukoff 11085 (A, F, G, K, MO, NY, S, US).<br />
1 cm
50 Flora Neotropica<br />
This species has a remarkable, apparently dis-<br />
continuous, distribution. It appears to be very<br />
closely related to C. ferruginea. The two taxa<br />
could prove to be conspecific, although probably<br />
distinct at the subspecific level. Several speci-<br />
mens from Venezuela <strong><strong>an</strong>d</strong> the Roraima Territory<br />
(Brazil) are distinct in having relatively broad (to<br />
suborbicular) leaves.<br />
12. <strong>Coussapoa</strong> cupularis Akkerm<strong>an</strong>s & C. C.<br />
Berg, Proc. Kon. Ned. Akad. Wetensch. Ser.<br />
C, 85(4): 448. 1982. Type. Brazil. Rond6nia:<br />
P6rto Velho, 5 Jun 1952 (9), J. F. Silva 69<br />
(holotype, IAN). Fig. 23.<br />
Trees, terrestrial. Leafy twigs 3-7 mm thick,<br />
brownish puberulous to hirtellous. Lamina coriaceous,<br />
elliptic to obovate, 5-10 x 3-7 cm,<br />
apex obtuse to shortly acuminate, base obtuse to<br />
(sub)cordate, margin entire to subcrenate; upper<br />
surface glabrous, lower surface on the midrib <strong><strong>an</strong>d</strong><br />
lateral veins sparsely to rather densely appressedpuberulous<br />
to strigillose, for the rest densely (appressed-)puberulous;<br />
lateral veins 5-8 pairs,<br />
slightly curved, basal pair unbr<strong>an</strong>ched or occasionally<br />
<strong>with</strong> a single br<strong>an</strong>ch, reaching the margin<br />
below to just above the middle of the lamina;<br />
intercostal venation pl<strong>an</strong>e; petiole 2-5 cm long,<br />
densely puberulous to strigillose; stipules 2-5 cm<br />
long, yellow to brownish subsericeous to subhirsute.<br />
Staminate inflorescences unknown. Pistil-<br />
of which tends to be below the middle; in the<br />
strongly broadened, almost cupuliform apices of<br />
the head-bearing br<strong>an</strong>ches of the pistillate inflo-<br />
rescences; <strong><strong>an</strong>d</strong> in the relatively dense brown pa-<br />
pillate apex of the peri<strong>an</strong>th of the pistillate flow-<br />
er. Moreover, the stipules are densely covered<br />
<strong>with</strong> long yellowish hairs, which is uncommon<br />
in C. orthoneura. <strong>Coussapoa</strong> cupularis also shows<br />
resembl<strong>an</strong>ces to C. nitida.<br />
Because of the occurrence of more or less conspicuously<br />
broadened br<strong>an</strong>ches below the pistillate<br />
flower heads <strong><strong>an</strong>d</strong> other similarities, as in the<br />
indumentum, C. cupularis, C. orthoneura, C.<br />
arachnoidea, <strong><strong>an</strong>d</strong> C. nitida appear to constitute<br />
a rather distinct group among the taxa <strong>with</strong><br />
ebracteate inflorescences <strong><strong>an</strong>d</strong> unistaminate flowers.<br />
Collection Huashikat 1585, Peru, Amazonas,<br />
basin of Rio S<strong>an</strong>tiago, 65 km N of Pinglo, near<br />
Caterpiza, 19 Dec 1979 (9) (MO, U), resembles<br />
C. cupularis in m<strong>an</strong>y characters. The main difference<br />
is the br<strong>an</strong>ching of the basal lateral veins.<br />
Less import<strong>an</strong>t are the greater number of lateral<br />
veins (8-9 pairs) <strong><strong>an</strong>d</strong> some differences in the indumentum.<br />
In the leaf venation the specimen<br />
approaches C. nitida. The identity of this collection<br />
is uncertain.<br />
13. <strong>Coussapoa</strong> curr<strong>an</strong>ii Blake, Contr. U.S. Natl.<br />
Herb. 20: 237. 1919. Type. Brazil. Bahia: Rio<br />
late inflorescences br<strong>an</strong>ched; heads 3-4, globose, Gongoji, 1 Oct-30 Nov 1915 (9), Curr<strong>an</strong> 8<br />
ca. 4-6 mm, in fruit up to ca. 10-15 mm diam.; (holotype, US; isotype, RB). Fig. 24.<br />
common peduncle 2-4 cm long, puberulous to<br />
<strong>Coussapoa</strong> warburgi<strong>an</strong>a Mildbraed, Notizbl. Bot. Gart.<br />
hirtellous, br<strong>an</strong>ches below the heads more or less Berlin 10: 416. 1928. Type. Brazil. Rio de J<strong>an</strong>eiro:<br />
strongly broadened (more or less cupuliform); Serra da Estrella, 21 May 1877 (3), Glaziou 8934<br />
peri<strong>an</strong>th ca. 1-2 mm high, subpapillate <strong>with</strong> mi- (holotype, B; isotypes, BR, G, K, LE, MO, P, S, U,<br />
nute thick brown hairs. Interfloral bracts absent. US).<br />
<strong>Coussapoa</strong> incomitata St<strong><strong>an</strong>d</strong>ley, Publ. Field Mus. Bot.<br />
Distribution (Fig. 6). Brazil (Rond6nia); in for- 22: 72. 1940. Type. Brazil. Minas Gerais: Mun.<br />
est of terra firme.<br />
Tombos, Fazenda da Cachoeira, 29 Jul 1935 (9),<br />
Mello Barreto 1795 (holotype, F).<br />
Specimens studied. BRAZIL. RONDONIA: Road Porto<br />
Velho-Cuiaba, km 117, Rio Jamari, 15 Aug 1968 Tree, (mostly?) terrestrial, up to 40 m tall.<br />
(2), Pr<strong>an</strong>ce et al. 6969 (U); 8 km from Porto Velho, 5 Leafy twigs 4-6 mm thick, glabrous or sparsely<br />
Jun 1952 (2), J. F. Silva 69 (IAN, type collection), 26<br />
Jun 1952 (2), J. F. Silva 211 puberulous, sometimes initially <strong>with</strong> sparse<br />
(IAN).<br />
arachnoid hairs. Lamina (sub)coriaceous, ob-<br />
<strong>Coussapoa</strong> cupularis, represented by three col- ovate or subobovate, 4-19 x 1-9 cm, apex oblections<br />
from the same area, appears to be closely tuse to rounded to emarginate or shortly acurelated<br />
to C. orthoneura, <strong><strong>an</strong>d</strong> may even prove minate, base obtuse to acute, margin entire; upper<br />
not to be distinct at the species level. At present, surface glabrous or <strong>with</strong> sparse (to rather dense)<br />
the species differs from C. orthoneura in the basal white arachnoid hairs which soon disappear,<br />
lateral veins which mostly reach the margin be- lower surface sparsely puberulous, often also <strong>with</strong><br />
low the middle of the lamina, the broadest part (mostly sparse) deciduous arachnoid hairs; lat-
<strong>Coussapoa</strong><br />
1 cm<br />
FIG. 23. <strong>Coussapoa</strong> cupularis: 1, leafy twig <strong>with</strong> pistillate inflorescences (J. F. Silva 69).<br />
eral veins 7-10 pairs, basal pair unbr<strong>an</strong>ched or<br />
sometimes faintly br<strong>an</strong>ched, reaching the margin<br />
below the middle of the lamina; intercostal venation<br />
slightly prominent, the smaller veins pl<strong>an</strong>e;<br />
petioles 1-5 cm long, glabrous to sparsely pub-<br />
erulous or also <strong>with</strong> sparse to dense arachnoid<br />
hairs; stipules 0.4-0.6 cm long, appressed-pub-<br />
erulous to strigillose to subsericeous (to subhir-<br />
51
52 Flora Neotropica<br />
I<br />
1 cm<br />
FIG. 24. <strong>Coussapoa</strong> curr<strong>an</strong>ii: 1, leafy twig <strong>with</strong> staminate inflorescences<br />
(Glaziou 8934).<br />
sute). Staminate inflorescences br<strong>an</strong>ched; heads<br />
several to m<strong>an</strong>y, globose, ca. 2 mm diam.; com-<br />
mon peduncle 1-5 cm long, sparsely to densely<br />
puberulous or glabrous; peri<strong>an</strong>th ca. 1 mm high,<br />
sparsely minutely puberulous; stamens two, ex-<br />
ceeding the peri<strong>an</strong>th. Pistillate inflorescences<br />
br<strong>an</strong>ched; heads 3-6, globose, ca. 4-6 mm diam.;<br />
common peduncle 1-4 cm long, patent to ap-<br />
pressed puberulous; peri<strong>an</strong>th ca. 1-2 mm high,<br />
minutely puberulous. Interfloral bracts sub-<br />
spathulate to subpeltate, puberulous at the apex.<br />
Distribution (Fig. 7). Eastern Brazil, from Rio<br />
de J<strong>an</strong>eiro to Bahia; in moist forests up to<br />
500 m.<br />
Specimens studied. BRAZIL. Without locality, 2 Dec<br />
1968 (8), Gomes et al. 2818 (RB). BAHIA: Rio Gongoji,<br />
1 Oct-20 Nov 1915 (9), Curr<strong>an</strong> 8 (RB (HJBR 13070),<br />
US, type collection). EsPiarro SANTO: Res. Flor. de<br />
Linhares, 24 May 1978 (d), DAF 004 (GUA). MINAS<br />
GERAIS: Mun. Tombos, Fazenda da Cachoeira, 29 Jul<br />
1935 (9), Mello Barreto 1795 (F). Rio DE JANEIRO:<br />
Corcovado, 1857 (2), Casaretto 617 (G); Givea-Sao<br />
Conrado, Jun 1960 (2), Duarte 5239 (M, NY, RB, US);<br />
Serra da Estrella, Petr6polis, 21 May 1877 (a), Glaziou<br />
8934 (B, BR, G, K, LE, MO, P, U, US, type collection<br />
of C. warburgi<strong>an</strong>a); Corcovado, 29 Jun 1876 (st), Glaziou<br />
8934a (P); Rio de J<strong>an</strong>eiro, Tijuca, J<strong>an</strong> 1837 (2),<br />
Guillemin 1339 (P); Rio de J<strong>an</strong>eiro, Jardim Bot<strong>an</strong>ico,<br />
3 Sep 1926 (2), Kuhlm<strong>an</strong>n s.n. (U); Cordeira, Fazenda<br />
S<strong>an</strong>ta Clara, Feb 1970 (6), Lisboa s.n. (R); Rio de J<strong>an</strong>eiro,<br />
Jardim BotAnico, 29 Jul 1922 (8), Occhioni et<br />
al. (HJBR) 5864 (GUA, RB, U), 25 Jul 1922 (9), Occhioni<br />
et al. (HJBR) 5917 (RB).<br />
<strong>Coussapoa</strong> curr<strong>an</strong>ii is very closely related to<br />
C. floccosa. Affinities of these two species <strong>with</strong><br />
other <strong>Coussapoa</strong> are not clear.
<strong>Coussapoa</strong> 53<br />
14. <strong>Coussapoa</strong> duquei St<strong><strong>an</strong>d</strong>ley, Publ. Field Mus. PASTAZA: Rd. Hollin-Loreto-Coca, between Rio Pu-<br />
Bot. 22: 71. 1940. Type. Colombia. Caldas: cuno <strong><strong>an</strong>d</strong> Cacerio de Guam<strong>an</strong>i, 1200 m, 12 Dec 1987<br />
Palestina, 1500 m, 15 Jun 1938<br />
(6), Cer6n M. 2955<br />
(d), Duque-<br />
(BG); nr. Mera, ca. 1400 m, 22<br />
Nov 1938 (9), Schultze-Rhonhof3020 (B, type collec-<br />
Jaramillo 1767 (holotype, F; isotypes, A, K, tion of C. apoda). PICHINCHA: Rd. N<strong>an</strong>egalito-Pacto,<br />
NY, US). Fig. 25. 5 km W of Tulipe, 1300-1500 m, 22 Jul 1980 (6),<br />
Holm-Nielsen et al. 24519 (BG).<br />
<strong>Coussapoa</strong> apoda Mildbraed, Notizbl. Bot. Gart. Berlin<br />
15: 784. 1942. Type. Ecuador. Pastaza: Nr. Mera, C. duquei is closely related to C. villosa, esca.<br />
1400 m, 22 Nov 1938 (9), Schultze-Rhonhof3020 pecially to the form <strong>with</strong> brown arachnoid in-<br />
(holotype, B).<br />
dumentum (see p. 104). The species is distinct in<br />
the shortly pedunculate pistillate inflorescence<br />
<strong>with</strong> <strong>an</strong> ellipsoid to clavate head. This taxon could<br />
be regarded as distinct only at the subspecies<br />
level.<br />
Tree, (secondarily?) terrestrial, up to 30 m tall.<br />
Leafy twigs 5-10 mm thick, hirtellous to hirsute.<br />
Lamina coriaceous, elliptic to ovate, 8-18 x 5-<br />
11 cm, apex obtuse to acute to shortly acuminate,<br />
base subacute to obtuse (or to subcordate), margin<br />
entire to subcrenate; upper surface glabrous,<br />
lower surface puberulous to hirtellous on the<br />
veins, the whole surface densely covered <strong>with</strong><br />
(sub)persistent pale yellow-brown arachnoid<br />
hairs; lateral veins 10-14 pairs, straight, main<br />
basal pair (sometimes faintly) br<strong>an</strong>ched, reaching<br />
the margin below the middle of the lamina; intercostal<br />
venation prominent; petiole 2-10 cm<br />
long, (rather) sparsely puberulous, <strong>with</strong> whitish<br />
to yellowish-brown arachnoid hairs which are<br />
also sometimes deciduous; stipules 2-7 cm long,<br />
white to brown strigose to hirsute <strong><strong>an</strong>d</strong> <strong>with</strong> reddish-brown<br />
arachnoid hairs. Staminate inflorescences<br />
br<strong>an</strong>ched; heads m<strong>an</strong>y, globose to oblate,<br />
ca. 4-8 mm diam.; common peduncle 1-2 cm<br />
long, puberulous to hirtellous, sometimes also<br />
<strong>with</strong> arachnoid hairs; peri<strong>an</strong>th ca. 1 mm high,<br />
yellowish-brown puberulous; stamens two, far<br />
exceeding the peri<strong>an</strong>th. Pistillate inflorescences<br />
unbr<strong>an</strong>ched; heads ellipsoid to clavate, ca. 1.5<br />
x 1 cm; peduncle 0.5-0.8 cm long, puberulous<br />
to hirtellous hairs, sometimes also <strong>with</strong> arachnoid<br />
hairs; peri<strong>an</strong>th ca. 1-2 mm high, densely,<br />
minutely puberulous. Interfloral bracts (sub)<br />
spathulate, puberulous at the apex.<br />
Distribution (Fig. 7). Ande<strong>an</strong> region (Colombia<br />
<strong><strong>an</strong>d</strong> Ecuador); in (sub)mont<strong>an</strong>e forest, between<br />
1200 <strong><strong>an</strong>d</strong> 2000 m.<br />
15. <strong>Coussapoa</strong> echinata Akkerm<strong>an</strong>s & C. C. Berg,<br />
Proc. Kon. Ned. Akad. Wetensch. Ser. C, 85(4):<br />
449. 1982. Type. P<strong>an</strong>ama. P<strong>an</strong>ama: Cerro Jefe,<br />
Altos de Pacora, Rio Diablo, 15 km NE of<br />
Cerro Azul village, 5 J<strong>an</strong> 1975 (9), Gentry &<br />
Mori 13408 (holotype, U; isotype, MO).<br />
Fig. 26.<br />
Tree, hemi-epiphytic. Leafy twigs 4-6 mm<br />
thick, glabrous or sparsely appressed-puberulous.<br />
Lamina coriaceous, oblong to elliptic to<br />
subobovate, 9-20 x 4-10 cm, apex acute to acuminate,<br />
base acute, margin entire; upper surface<br />
glabrous, lower surface (purplish or reddish), glabrous<br />
or on the veins sparsely puberulous; lateral<br />
veins 4-5 pairs, slightly curved, basal pair (faint-<br />
Specimens studied. COLOMBIA. ANTIOQUIA: Medellin,<br />
ca. 1500 m, - ly) br<strong>an</strong>ched, reaching the margin above the middle<br />
of the lamina; intercostal venation slightly<br />
prominent to pl<strong>an</strong>e; petiole 2-6 cm long, glabrous;<br />
stipules 1.5-2.5 cm long, sparsely appressed-puberulous.<br />
Staminate inflorescences<br />
br<strong>an</strong>ched; heads numerous, ca. 2 mm diam.;<br />
common peduncle ca. 1.5-2 cm long, puberulous;<br />
peri<strong>an</strong>th ca. 1 mm high, glabrous; stamens<br />
three. Pistillate inflorescences br<strong>an</strong>ched; heads 3-<br />
8, globose, ca. 4-6 mm, in fruit up to 8 mm<br />
diam.; common peduncle 1-3.5 cm long, <strong>with</strong><br />
sparse patent short hairs, occasionally mixed <strong>with</strong><br />
some longer hairs; flowers free, peri<strong>an</strong>th ca. 1-2<br />
mm high, (when dry) <strong>with</strong> <strong>an</strong> acute apex, gla-<br />
(st), Toro 120c<br />
brous;<br />
(US). CALDAS:<br />
fruiting peri<strong>an</strong>th or<strong>an</strong>ge. Interfloral bracts<br />
Cord. Central, Palestina, 1500 m, 15 Jun 1938 (8), narrowly spathulate, short, sparsely, minutely<br />
Duque-Jaramillo 1767 (A, F, K, NY, US, type collec- puberulous.<br />
tion). NORTE DE SANTANDER: Ocafa, Jun 1845 (9), Pur- Distribution (Fig. 7). P<strong>an</strong>ama (P<strong>an</strong>ama <strong><strong>an</strong>d</strong><br />
die s.n. (K).<br />
S<strong>an</strong><br />
ECUADOR. NAPO:<br />
Bias); in wet<br />
SE<br />
forest,<br />
of Borja, ca.<br />
up to 800 m.<br />
1900 m, 3 Aug<br />
1960 (d), Grubb et al. 1205 (K, NY); NE of Borja, ca. Specimens studied. PANAMA. PANAMA: Road El<br />
1800 m, 15 Aug 1960 (9), Grubb et al. 1283 (K, NY); Ll<strong>an</strong>o-Carti, km 17-20, 8 Mar 1974 (9), Dressier 4636<br />
nr. Borja, 1650 m, 16 Aug 1975 (9), Little et al. 212 (F); Cerro Jefe, Altos de Pacora region, Rio<br />
(Q), 1850<br />
Diablo,<br />
m, 16 Aug 1975 (8), Little et al. 213 (Q). 15 km E of Cerro Azul village, 5 J<strong>an</strong> 1975 (2), Gentry
54<br />
2 ~ ? I<br />
wo t :1cm<br />
Flora Neotropica<br />
FIG. 25. <strong>Coussapoa</strong> duquei: 1, leafy twig <strong>with</strong> pistillate inflorescences (Grubb et al. 1283); 2, staminate<br />
inflorescences<br />
(Grubb et al. 1205).
<strong>Coussapoa</strong> SS<br />
FIG. 26. <strong>Coussapoa</strong><br />
'1 R~~~~~~~~~~~~~~~~~~~~~~~~?<br />
: e'<br />
r ]; e<br />
echinata: 1, leafy twig <strong>with</strong> pistillate inflorescences<br />
4636).i? ?.<br />
(Dressler
56 Flora Neotropica<br />
G. 27. oussapoaferruginea: 1, leafy twig <strong>with</strong> staminate inflorescences (Irwin am et<br />
FIG. 27. <strong>Coussapoa</strong> ferruginea: 1, leafy twig <strong>with</strong> staminate inflorescences (Irwin et al. 48631).<br />
& Mori 13408 (MO, U, type collection). SAN BLAS:<br />
Nusag<strong><strong>an</strong>d</strong>i, rd. to Carti, 18 Jul 1984 (a), McDonagh et<br />
al. 114 (MO).<br />
The species shows similarities to C. parviceps,<br />
from which it is distinct in the shape of the leaves,<br />
the free pistillate flowers, <strong><strong>an</strong>d</strong> the presence of<br />
interfloral bracts in the pistillate heads. On the<br />
other h<strong><strong>an</strong>d</strong>, C. echinata shows similarities to C.<br />
herthae, especially in the acute apex of the fruit-<br />
ing peri<strong>an</strong>th (in dry material).<br />
16. <strong>Coussapoa</strong> ferruginea Trecul, Ann. Sci. Nat.<br />
Bot. Ser. 3, 8: 93. 1847; Miquel, Fl. bras. 4(1):<br />
137. 1853; Berg, Fl. Suriname 5(1): 286. 1975.<br />
Type. French Gui<strong>an</strong>a. Without locality, 1838<br />
(6), Leprieurs.n. (holotype, P; isotypes, G, GH,<br />
L, U). Fig. 27.<br />
Tree, hemi-epiphytic. Leafy twigs 3-7 mm<br />
thick, brown(ish) (sub)hirsute. Lamina coria-<br />
ceous, l<strong>an</strong>ceolate to oblong, 2-8 x 1-4 cm, apex<br />
acute to subobtuse, base acute, margin entire;<br />
upper surface glabrous, lower surface appressed-<br />
puberulous to strigose on the main veins, pu-<br />
berulous to hirtellous (to subhirsute) on the<br />
smaller veins, the whole surface covered <strong>with</strong><br />
dense persistent brown arachnoid hairs; lateral<br />
veins 4-5(-6) pairs, straight, basal pair unbr<strong>an</strong>ched,<br />
reaching the margin below the middle<br />
of the lamina; intercostal venation prominent;<br />
petiole 0.5-1.5 cm long, densely brown hirtellous<br />
to subhirsute; stipules 1.5-3 cm long, brown<br />
(sub)hirsute to strigose or partly whitish subsericeous.<br />
Staminate inflorescences br<strong>an</strong>ched; heads<br />
several to m<strong>an</strong>y, ca. 2-3 mm diam.; common<br />
peduncle ca. 1 cm long, densely hirtellous; peri-<br />
<strong>an</strong>th ca. 1 mm high, densely puberulous; stamens<br />
two, far exceeding the peri<strong>an</strong>th. Pistillate inflorescences<br />
unknown. Interfloral bracts spathulate,<br />
puberulous.<br />
Distribution (Fig. 6). French Gui<strong>an</strong>a <strong><strong>an</strong>d</strong> the<br />
adjacent part of Amapa (Brazil); in riverine forest.<br />
Specimens studied. FRENCH GUIANA. Without<br />
locality, 1838 (d), Leprieur s.n. (F, G, GH, L, P, R, U,<br />
type collection), 1840 (8), Leprieur s.n. (FI, G, GH, K,<br />
P).<br />
BRAZIL. AMAPA: Rio Oiapoque, between mouth of<br />
Camopi River <strong><strong>an</strong>d</strong> Cachoeira Camaraua, 3 Oct 1960<br />
(d), Irwin et al. 48631 (F, K, MO, NY, U, US).<br />
The species is similar in m<strong>an</strong>y characters to<br />
C. crassivenosa. However, it is distinct from the
<strong>Coussapoa</strong><br />
FIG. 28. <strong>Coussapoa</strong>floccosa: 1, leafy twig <strong>with</strong> pistillate inflorescences<br />
(Irwin 2058).<br />
latter in having a narrower lamina, a smaller<br />
number of lateral veins, <strong><strong>an</strong>d</strong> a relatively short<br />
petiole. Pistillate inflorescences are needed to decide<br />
whether this taxon is distinct from C. crassivenosa<br />
at the specific or only intraspecific level.<br />
17. <strong>Coussapoa</strong> floccosa Akkerm<strong>an</strong>s & C. C. Berg,<br />
Proc. Kon. Ned. Akad. Wetensch. Ser. C, 85(4):<br />
451. 1982. Type. Brazil. Minas Gerais: Vi9osa,<br />
16 Nov 1935 (2), Kuhlm<strong>an</strong>n 2074 (holotype,<br />
RB; isotypes, RB, U, US). Fig. 28.<br />
Tree or shrub, (mostly?) hemi-epiphytic.<br />
Leafy twigs 10-15 mm thick, (rather) densely<br />
57<br />
puberulous to (sub)hirsute. Lamina coriaceous,<br />
elliptic to ovate or obovate, 10-25 x 6-17 cm,<br />
apex rounded to obtuse or to emarginate, base<br />
obtuse to rounded to cordate, margin entire; upper<br />
surface often densely covered <strong>with</strong> white<br />
arachnoid hairs which later disappear, lower surface<br />
hirtellous to subtomentellous on the veins,<br />
the whole surface sparsely to densely covered<br />
<strong>with</strong> white arachnoid hairs, later deciduous; lateral<br />
veins 7-10 pairs, (main) basal pair distinctly<br />
br<strong>an</strong>ched, reaching the margin at or above the<br />
middle of the lamina; intercostal venation very<br />
prominent; petiole 1.5-5 cm long, tomentose to<br />
densely hirtellous; stipules 1-2.5 cm long, white<br />
to brownish appressed-puberulous to strig(ill)ose
58 Flora Neotropica<br />
to subsericeous (to subhirsute), sometimes also mm high, glabrous. Interfloral bracts (sub)peltate,<br />
<strong>with</strong> sparse arachnoid hairs. Staminate inflores- nearly glabrous.<br />
cences br<strong>an</strong>ched; heads numerous, globose, ca. 4 Distribution (Fig. 7). Central America: Costa<br />
mm diam.; common peduncle 1.5-4 cm long, Rica (Puntarenas), Nicaragua (Bluefields <strong><strong>an</strong>d</strong> Rio<br />
puberulous; peri<strong>an</strong>th ca. 1 mm high, puberulous; S<strong>an</strong> Ju<strong>an</strong>), P<strong>an</strong>ama (S<strong>an</strong> Blas); in wet (riverside<br />
stamens two, exceeding the peri<strong>an</strong>th. Pistillate or swamp) forest.<br />
inflorescences br<strong>an</strong>ched; heads 2-3, partly fused,<br />
(sub)globose, 5-7 mm diam.; common peduncle<br />
Specimens studied. NICARAGUA. BLUEFIELDS: Rio<br />
1-3 cm long, densely puberulous; peri<strong>an</strong>th 1-2<br />
Kama, between Kama <strong><strong>an</strong>d</strong> C<strong>an</strong>io Valentin, 10 Mar<br />
1966 (2), Proctor et al. 27080 (F, US, type collection).<br />
mm high, glabrous. Interfloral bracts sub- Rio SAN JUAN: 1 km NW of Rio S<strong>an</strong>ta Cruz, 22 Feb<br />
spathulate, puberulous at the apex.<br />
1984 (2), Moreno 23629 (BG).<br />
Distribution (Fig. 7). Brazil (Minas Gerais, Vi- COSTA RICA. PUNTARENAS: Peninsula de Osa, 8<br />
qosa) in<br />
km S of<br />
forest.<br />
Rinc6n, 28 Feb 1965 (9), Jimenez 3016 (F).<br />
PANAMA. SAN BLAS: El Ll<strong>an</strong>o-Carti rd., km 19.1,<br />
Specimens studied. BRAZIL. MINAS GERAIS: Est. 1 Feb 1985 (2), Nevers et al. 4804 (WIS), 3 Nov 1985<br />
Exp. de Agua Limpa, Feb 1968 (2), Gomes et al. 2819 (st), Nevers et al. 6186 (BG). SAN BLAS-PANAMA: El<br />
(RB); Vi9osa, 10 Nov 1958 (2), Irwin 2058 (F, NY,<br />
Ll<strong>an</strong>o-Carti rd., 28 Aug 1982 (d), Hamilton et al. 1062<br />
R, US), 31 Aug 1934 (st), Kuhlm<strong>an</strong>n 1832 (US), 1935 (BG).<br />
(st), Kuhlm<strong>an</strong>n 2073 (RB, US), 16 Nov 1935 (2), Kuhlm<strong>an</strong>n<br />
2074 (RB, U, US, type collection), 31 Aug 1934<br />
This species appears to be one of a group which<br />
(st), Kuhlm<strong>an</strong>n 2117 (RB), 1935 (9), Kuhlm<strong>an</strong>n 2207 also includes C. <strong>an</strong>gustifolia <strong><strong>an</strong>d</strong> C. trinervia <strong><strong>an</strong>d</strong><br />
(GUA), 8 Oct 1976 (9), G. Rodrigues et al. 877 (RB); is characterized by shortly petiolate, more or less<br />
Capolivinba, nr. Rio Novo, Sep 1895 (6), Schwacke distinctly obovate leaves <strong>with</strong> few lateral veins,<br />
11895 (B, P).<br />
<strong><strong>an</strong>d</strong> sparse indumentum.<br />
This species, known only from the surroundings<br />
of Vicosa, is very closely related to C. cur- 19. <strong>Coussapoa</strong> herthae Mildbraed, Notizbl. Bot.<br />
r<strong>an</strong>ii; but the two taxa are sufficiently different Gart. Berlin 14: 29. 1938. Type. Ecuador. Pifor<br />
them each to be regarded as species in their chincha: S<strong>an</strong> Carlos de los Colorados, 5 Oct<br />
own right.<br />
1935 (d), Schultze-Rhonhof 953 (holotype, B).<br />
Fig. 30.<br />
18. <strong>Coussapoa</strong> glaberrima W. Burger, Phytolo-<br />
gia 26: 422. 1973; Burger, Fieldi<strong>an</strong>a Bot. 40:<br />
133, t. 22. 1977. Type. Nicaragua. Bluefields:<br />
Rio Kama, between Kama <strong><strong>an</strong>d</strong> C<strong>an</strong>io Valentin,<br />
10 Mar 1966 (2), Proctor et al. 27080 (holo-<br />
type, F; isotype, US). Fig. 29.<br />
Tree, hemi-epiphytic or terrestrial, small.<br />
Leafy twigs 2-5 mm thick, glabrous. Lamina<br />
subcoriaceous, obovate to narrow elliptic, 2-14<br />
x 1-6 cm, apex acute, base acute to obtuse, margin<br />
entire; both surfaces glabrous; lateral veins<br />
2-3 pairs, slightly curved, basal pair indistinctly<br />
br<strong>an</strong>ched, reaching the margin far above the<br />
middle of the lamina; intercostal venation slightly<br />
prominent; petiole 0.5-2 cm long, glabrous;<br />
stipules 1.2 cm long, glabrous. Staminate inflorescences<br />
br<strong>an</strong>ched; heads ca. 10-12, ca. 2 mm<br />
diam.; common peduncle 1.5-3 cm long, gla-<br />
brous; peri<strong>an</strong>th ca. 1 mm high, glabrous; stamens<br />
two, just exceeding the peri<strong>an</strong>th. Pistillate inflorescences<br />
poorly br<strong>an</strong>ched; heads 2-3, globose,<br />
ca. 6 mm, in fruit up to 15 mm diam.; common<br />
peduncle 2-4 cm long, glabrous; peri<strong>an</strong>th ca. 1<br />
Tree, mostly hemi-epiphytic, up to 25 m tall.<br />
Leafy twigs 5-13 mm thick densely, yellow to<br />
brown, strigose to hirsute to subvillous. Lamina<br />
coriaceous, elliptic to ovate to obovate or to ob-<br />
long, 16-32 x 10-20 cm, apex acuminate to<br />
acute, base obtuse to rounded or occasionally<br />
subcordate, margin entire or subcrenate; upper<br />
surface glabrous, sparsely to rather densely pub-<br />
erulous to hirtellous or on the main veins to<br />
subsericeous, sometimes <strong>with</strong> white arachnoid<br />
hairs which soon disappear; lateral veins (4-)<br />
8-11 pairs, curved, basal pair distinctly to faintly<br />
br<strong>an</strong>ched, reaching the margin usually below,<br />
sometimes at or just above the middle of the<br />
lamina, sometimes also other lateral veins<br />
br<strong>an</strong>ched, a distinct submarginal vein in the lower<br />
part of the lamina; intercostal venation (often<br />
very) prominent; petiole (1-)3-9 cm long, puberulous<br />
to hirtellous (to subhirsute); stipules 2-<br />
6 cm long, densely reddish- to yellowish-brown<br />
subsericeous to subhirsute to subvillous. Staminate<br />
inflorescences br<strong>an</strong>ched; heads m<strong>an</strong>y,<br />
globose, 2-3 mm in diam.; common peduncle
<strong>Coussapoa</strong> 59<br />
1 cm<br />
FIG. 29. <strong>Coussapoa</strong> glaberrima: 1, leafy twig <strong>with</strong> pistillate inflorescences<br />
(Proctor et al. 27080).<br />
(1-?)3-7 cm long, puberulous to hirtellous; peri-<br />
<strong>an</strong>th ca. 1 mm high, glabrous; stamens three, far<br />
exceeding the peri<strong>an</strong>th. Pistillate inflorescences<br />
uador <strong><strong>an</strong>d</strong> southern Colombia); in wet forest, up<br />
to 1200 m.<br />
br<strong>an</strong>ched; heads 6-10, sometimes partly fused,<br />
Specimens studied. COLOMBIA. NARuio: Bay of<br />
Tumaco, Rio<br />
(sub)globose, 5-9 mm, in fruit up to 13 mm<br />
Y<strong>an</strong>aje, 6 Jul 1955 (9), Romero-Castadiam.;<br />
ineda 5283 (COL).<br />
common peduncle 3-6 cm long, 2-3 mm thick, ECUADOR. CARCHI: Rio S<strong>an</strong> Ju<strong>an</strong>, nr. Chical, 12<br />
puberulous to hirtellous; peri<strong>an</strong>th ca. 2 mm high, km below Maldonado, 1200 m, May 1978 (2), Madison<br />
minutely puberulous; fruiting peri<strong>an</strong>th<br />
et al. 4764<br />
or<strong>an</strong>ge.<br />
(AAU, F, QCA, SEL, U); El Pail6n, 45 km<br />
below<br />
Interfloral bracts spathulate to Maldonado, 28 Nov 1979 (d), Madison et al.<br />
subpeltate, pub- 7101 (SEL, U). ESMERALDAS: Nr. Lita, W of Rio Lita,<br />
erulous at the apex.<br />
9 Feb 1981 (2), Berg 1259 (QCA, U). GUAYAS: Nr.<br />
Distribution (Fig. 7). Pacific coastal region (Ec- Bucay (=Grl. Elizalda), 8-15 Jun 1945 (9), CampE3674
60<br />
cm1<br />
2 I!<br />
1<br />
Flora Neotropica<br />
FIG. 30. <strong>Coussapoa</strong> herthae: 1, leafy twig <strong>with</strong> pistillate inflorescences (Schulze-Rhonhof 1953); 2, staminate<br />
inflorescences (Dodson 5776).
<strong>Coussapoa</strong><br />
(NY, U). IMBABURA: Parambas, 30 May 1949 (9), Acosta<br />
Solis 12661 (F). Los Rios: Rio Palenque Biol. St.,<br />
km 56 on road Quevedo-S<strong>an</strong>to Domingo de los Colorados,<br />
27 Feb 1975 (6), Dodson 5776 (MO, QCA),<br />
29 May 1976 (2), Dodson 6082 (MO), 20 Mar 1980<br />
(a), Dodson et al. 9539 (SEL, U), 13 Feb 1974 (6),<br />
Gentry 9892 (MO, U). PICHINCHA: 35 km N of S<strong>an</strong>to<br />
Domingo de los Colorados, nr. bridge over Rio Bl<strong>an</strong>co,<br />
3 Feb 1974 (2), Gentry 9606 (MO, U); S<strong>an</strong> Carlos de<br />
los Colorados, 5 Oct 1935 (6), Schultze-Rhonhof 1953<br />
(B, type collection).<br />
<strong>Coussapoa</strong> herthae shows general similarity to<br />
C. nymphaeifolia <strong><strong>an</strong>d</strong> in the echinate (dry) fruiting<br />
head to C. echinata.<br />
The species is very variable in the shape <strong><strong>an</strong>d</strong><br />
venation of the leaves.<br />
20. <strong>Coussapoa</strong> latifolia Aublet, Hist. pl. Gui<strong>an</strong>e<br />
2: 955, t. 362. 1775; Trecul, Ann. Sci. Nat.<br />
Bot. Ser. 3, 8: 94. 1847; Miquel, Fl. bras. 4(1):<br />
135. 1853; Berg, Fl. Suriname 5(1): 280. 1975.<br />
Type. French Gui<strong>an</strong>a. Without locality, - (9),<br />
Aublet s. n. (holotype, BM). Fig. 31.<br />
<strong>Coussapoa</strong> obovata Miquel, Het Instituut (1842): 200,<br />
t. 3. 1843. Type. Surinam. Pl<strong>an</strong>tation Bergendaal, -<br />
(2), Focke 9 (holotype, L).<br />
<strong>Coussapoa</strong> latifolia Aublet var. obovata (Miquel) Miquel,<br />
Fl. bras. 4(1): 135. 1853.<br />
<strong>Coussapoa</strong> froesii St<strong><strong>an</strong>d</strong>ley, Publ. Field Mus. Bot. 17:<br />
162. 1937. Type. Brazil. Mar<strong>an</strong>hao: Rio Maracassume<br />
region, 14 Sep 1932 (2), Fr6es 1907 (holotype,<br />
NY; isotypes, A, G, K, U).<br />
Shrub or tree, mostly hemi-epiphytic, up to 35<br />
m tall. Leafy twigs 4-9 mm thick, glabrous. Lamina<br />
coriaceous or subcoriaceous, ovate to elliptic,<br />
sometimes oblong to obovate, occasionally to<br />
suborbicular, 4-18(-25) x diam.; common peduncle 0.5-3 cm long, sparsely<br />
to densely appressed-puberulous; peri<strong>an</strong>th ca.<br />
0.5-1 mm high, glabrous; stamens two, just exceeding<br />
the peri<strong>an</strong>th. Pistillate inflorescences<br />
br<strong>an</strong>ched; heads 2-15, globose, ca. 3-4 mm, in<br />
fruit up to 8 mm diam.; common peduncle 1-3<br />
cm long, sparsely to densely appressed-puberulous;<br />
peri<strong>an</strong>th ca. 1-1.5 mm high, glabrous.<br />
Interfloral bracts small, subpeltate to (sub)<br />
spathulate, minutely puberulous at the apex.<br />
Distribution (Fig. 7). Western Gui<strong>an</strong>a region<br />
(Surinam <strong><strong>an</strong>d</strong> French Gui<strong>an</strong>a), in Brazil from<br />
Amapa through eastern Para to Mar<strong>an</strong>hao; further,<br />
apparently segregated from the main area<br />
in the central part of the Amazon Basin; forest<br />
of terra firme.<br />
Specimens studied. SURINAM. Brownsberg, 3 Sep<br />
1915 (9), BW(Stahel & Gonggrijp) 732 (K, MO, NY,<br />
P), 19 Sep 1931 (9), V<strong>an</strong> Emden s.n. (F, U, US), 23<br />
Sep 1931 (st), V<strong>an</strong> Emden LVI (U); Wilhelmina Mts.,<br />
upper Gr<strong>an</strong> Rio, Maup6 dam, 20 Feb 1926 (st), Exp.<br />
Wilhelmina Gebergte 196 (U); <strong>with</strong>out locality,<br />
3-15 cm, apex acute<br />
to shortly acuminate to obtuse to rounded, base<br />
acute to obtuse or to subcordate, margin entire,<br />
pl<strong>an</strong>e or more or less revolute towards the base;<br />
upper surface glabrous, lower surface glabrous or<br />
sparsely appressed-puberulous <strong>with</strong> hairs of about<br />
equal length; lateral veins 4-7 pairs, (slightly)<br />
curved, basal pairs mostly br<strong>an</strong>ched, sometimes<br />
(especially) in narrow leaves unbr<strong>an</strong>ched, reaching<br />
the margin at to below or (especially in obovate<br />
leaves) above the middle of the lamina,<br />
the other lateral veins sometimes poorly br<strong>an</strong>ched<br />
(furcate); intercostal venation (almost) pl<strong>an</strong>e;<br />
petiole 1-7 cm long, sparsely to rather densely<br />
minutely appressed-puberulous; stipules 0.5-<br />
1.5(-2.5) cm long, sparsely to densely whitish to<br />
brownish appressed-puberulous, sometimes<br />
brownish subvelutinous. Staminate inflorescences<br />
br<strong>an</strong>ched; heads m<strong>an</strong>y, globose, ca. 1 mm<br />
- (2),<br />
Focke s.n. (GH); pl<strong>an</strong>tation Bergendaal, - (9), Focke<br />
9 (L, type collection of C. obovata = C. latifolia var.<br />
obovata); pl<strong>an</strong>tation Blauberg, - (9), Focke 418 (U);<br />
Distr. Nickerie, ca. 20 km SW of Av<strong>an</strong>avero dam site,<br />
14 Nov 1976 (9), Heyde et al. 78 (U); Wilhelmina Mts.,<br />
Lucie River, 2-5 km below confluence <strong>with</strong> Oost River,<br />
8 Sep 1963 (9), Irwin et al. 55487 (MO, NY, OXF, R,<br />
S, U, US); Lely Mts., 19 Sep 1975 (st), Lindem<strong>an</strong> et<br />
al. 33 (U), 19 Sep 1975 (8), Lindem<strong>an</strong> et al. 73 (U), 4<br />
Oct 1975 (9), Lindem<strong>an</strong> et al. 749 (K, U), 5 Oct 1975<br />
(st), Lindem<strong>an</strong> et al. 799 (U); Jodensav<strong>an</strong>ne-Map<strong>an</strong>e<br />
Creek area, camp 8, 21 Dec 1954 (st), Lindem<strong>an</strong> 6938<br />
(U); Coppename River, Doksi Creek, 11 Oct 1954 (6),<br />
Mennega 271 (C, NY, U); T<strong>an</strong>jimama River, Kodji<br />
Creek, 25 Nov 1954 (6), Mennega 521 (A, C, NY, U);<br />
Voltzberg, 22 Sep 1956 (st), Schulz (LBB) 7787 (U);<br />
Jodensav<strong>an</strong>ne-Map<strong>an</strong>e Creek area, 5 May 1967 (st),<br />
Vreden (LBB) 11747 (U).<br />
FRENCH GUIANA. Without locality, - (2), Aublet<br />
s.n. (BM, type collection), Feb 1868 (2), Aubry-Lecompte<br />
s.n. (FI); Sinnamary, road to St. Elie, 22 Oct<br />
1981 (9), Billiet et al. 1108 (U); Montagne de Kaw, 13<br />
Dec 1954 (9), Cow<strong>an</strong> 38774 (F, K, NY, P, US); Mont<br />
Belvedere, 18 Nov 1974 (2), De Gr<strong>an</strong>ville et al. B5191<br />
(CAY, U); Oyapock River, Myuli Creek, Les Trois<br />
Sauts, 15 Sep 1977 (6), Gren<strong><strong>an</strong>d</strong> 1478 (CAY, U); <strong>with</strong>out<br />
locality, -<br />
(9), Lem<strong>an</strong> s.n. (P); Cayenne, - (9),<br />
Martin s.n. (or 112) (K); <strong>with</strong>out locality, - (9), Melinon<br />
s.n. (A, P); Maroni River, 1864 (9), Melinon 18<br />
(GH, K, LE, P); Antecume Pata, at confluence of It<strong>an</strong>y<br />
River <strong><strong>an</strong>d</strong> Maroni River, 14 Nov 1977 (2), Moretti 836<br />
(CAY); Kourou River, 3 km above Couy Creek, 19<br />
Sep 1967 (6), Oldem<strong>an</strong> B1338 (CAY, U); Cayenne, -<br />
(st), Poiteau s.n. (FI); Acarou<strong>an</strong>y, 1857 (6), Sagot 1136<br />
(B, BM, G, GH, K, S, U); Sinnamary, road to St. Elie,<br />
15 Sep 1978 (6), Sastre 6130 (P, U).<br />
BRAZIL. AMAPA: "Gafcho area," 21 Oct 1979 (9),<br />
Austin et al. 7149 (U); Rio Araguari, between 1?26'N,<br />
61
FIG. 31. <strong>Coussapoa</strong> latifolia: 1, leafy twig <strong>with</strong> pistillate inflorescences (Ducke (HJBR) 18456); 2, staminate inflor
<strong>Coussapoa</strong> 63<br />
51?58'W <strong><strong>an</strong>d</strong> 109'N, 51?52'W, 11 Sep 1961 (2), Pires<br />
et al. 50864 (NY, OXF, P, R, U, US); Rio Araguari,<br />
nr. 1?11'N, 52?8'W, 1 Oct 1964 (6), Pires et al. 51409<br />
(F, GH, NY, U); Col6nia Torrao, 2?25'N, 51015'W, 29<br />
Aug 1962 (2), Pires et al. 52656 (NY, OXF, U, US).<br />
AMAZONAS: Road M<strong>an</strong>aus-Caracarai, km 97, 30 Aug<br />
1979 (e), Cid et al. 942 (U); road M<strong>an</strong>aus-Porto Velho,<br />
km 235, Igarape Acu, 24 Nov 1973 (d), Lleras et al.<br />
P19661 (F, M, MO, NY, P, S, U); road M<strong>an</strong>aus-Caracarai,<br />
km 159, 20 Sep 1974 (6), Pr<strong>an</strong>ce et al. 22720<br />
(K, MO, NY, S, U, US). MARANHAO: Rio Maracassume<br />
region, 14 Sep 1932 (2), Frogs 1907 (A, G, K,<br />
NY, U, type collection of C. froesii). PARA: Col6nia<br />
Benjamin Const<strong>an</strong>t, Brag<strong>an</strong>oa, 21 Nov 1908 (2),<br />
Anonymous (HAMP) 9783 (G); Sta. Izabel, 10 Sep 1922<br />
(2), Ducke (HJBR) 18456 (RB); Col6nia Augusto Montenegro,<br />
Igarape Pitor6, 18 Sep 1958 (2), Frogs 34651<br />
(IAN); road Belem-Brasilia, km 93, 19 Sep 1959 (2),<br />
Kuhlm<strong>an</strong>n et al. 263 (NY, US); road Belem-Brasilia,<br />
km 301-306, 7 Aug 1960 (2), Oliveira 985 (U); Rio<br />
Jari, Monte Dourado, 16 Nov 1967 (2), Oliveira 3550<br />
(NY), 9 Nov 1967 (2), Oliveira 3599(NY), 8 Nov 1967<br />
(9), Oliveira 3682 (NY), 23 J<strong>an</strong> 1968 (st), Oliveira 3954<br />
(NY), road Cap<strong>an</strong>ema-Mar<strong>an</strong>hao, km 96,27 Oct 1965<br />
(2), Pr<strong>an</strong>ce et al. 1720 (MO, NY, S, U, US), km 98,<br />
21 Aug 1964 (2), Pr<strong>an</strong>ce et al. 58783 (U); Peixe Boi,<br />
26 Oct 1907 (2), Seguiera (HAMP) 8808 (BM, G, U);<br />
Rio Jari, Monte Dourado, 15 Oct 1968 (d), N. T. Silva<br />
1207 (NY), 28 Oct 1968 (2), N. T. Silva 1321 (NY, U,<br />
US); Rio Jari, between Pilao <strong><strong>an</strong>d</strong> Repartimento, 12<br />
Nov 1968 (2), N. T. Silva 1380 (IAN, NY, U, US).<br />
<strong>Coussapoa</strong> latifolia is very closely related to<br />
C. viridifolia. The species shows affinities <strong>with</strong><br />
C. microcephala, C. microcarpa, C. pachyphylla,<br />
<strong><strong>an</strong>d</strong> also <strong>with</strong> the Central Americ<strong>an</strong> C. macering<br />
the margin at or below the middle of the<br />
lamina, the other lateral veins sometimes<br />
br<strong>an</strong>ched; intercostal venation prominent; petiole<br />
1.5-3 cm long, puberulous to hirtellous; stipules<br />
0.5-1 cm long, yellowish to brown hirsute<br />
to subsericeous; terminal buds often more or less<br />
swollen. Staminate inflorescences br<strong>an</strong>ched;<br />
heads m<strong>an</strong>y, globose, ca. 6 mm diam.; common<br />
peduncle 4-7 cm long, puberulous to hirtellous;<br />
peri<strong>an</strong>th ca. 1 mm high, glabrous; stamens three,<br />
far exceeding the peri<strong>an</strong>th. Pistillate inflorescences<br />
unbr<strong>an</strong>ched; heads globose, ca. 5-8 mm<br />
diam.; peduncle 0.5-2.5 cm long, puberulous to<br />
hirtellous; peri<strong>an</strong>th ca. 1-2 mm high, glabrous.<br />
Interfloral bracts spathulate to subpeltate, puberulous.<br />
Distribution (Fig. 7). In (eastern) French<br />
Gui<strong>an</strong>a <strong><strong>an</strong>d</strong> in Brazil in eastern Para; in upl<strong><strong>an</strong>d</strong><br />
<strong><strong>an</strong>d</strong> riverine forest.<br />
Specimens studied. FRENCH GUIANA. Without<br />
locality (3), Leprieur s.n. (P, type collection); Approuague<br />
River, between Sapokaye Creek <strong><strong>an</strong>d</strong> Gr<strong><strong>an</strong>d</strong><br />
C<strong>an</strong>ori Falls, 23 Oct 1968 (6), Oldem<strong>an</strong> T243a (CAY);<br />
Sapkaye Creek, 22 Oct 1968 (3), Oldem<strong>an</strong> B1946 (CAY,<br />
K, P, U); Approuague River, Mapaou Falls, 26 J<strong>an</strong><br />
1970 (6), Oldem<strong>an</strong> B2861 (CAY, U); Oyapock River,<br />
3 km above Camopi, 8 Apr 1970 (6), Oldem<strong>an</strong> 3062<br />
(CAY, P, U).<br />
BRAZIL. PARA: Ilhas de Breves, Furo Macujubim,<br />
16 Nov 1912 (6), Ducke (HJBR) 18462 (RB); Alto<br />
Quatipuir, Dec 1899 (6), Huber(HAMP) 1775 (F, MG).<br />
rima. Because of similarities in the vegetative<br />
parts, C. latifolia c<strong>an</strong> be confused <strong>with</strong> C. orthoneura,<br />
especially specimens <strong>with</strong> elliptic leaves.<br />
<strong>Coussapoa</strong> leprieurii, one of the few species in<br />
which the lamina is scabrous above, appears to<br />
be rather closely related to C. sprucei. Specimens<br />
<strong>with</strong> <strong>an</strong> elliptic to oblong lamina having arachnoid<br />
hairs beneath are very similar to C. sprucei.<br />
Specimens <strong>with</strong> obovate leaves are reminiscent<br />
of (the unrelated) C. asperifolia ssp. asperifolia.<br />
21. <strong>Coussapoa</strong> leprieurii Benoist, Bull. Mus. Hist.<br />
Nat. Paris 30:103. 1924. Type. French Gui<strong>an</strong>a.<br />
Without locality, - (6), Leprieur s.n. (holo-<br />
type, P). Fig. 32.<br />
Tree, hemi-epiphytic, up to 10 m tall. Leafy<br />
twigs 5-8 mm thick, brown(ish) puberulous to<br />
hirtellous to hirsute, solid. Lamina coriaceous,<br />
(broadly) ovate to (broadly) elliptic to (broadly)<br />
obovate to oblong, 5-18 x 1.5-12 cm, apex ob-<br />
tuse to subacuminate to acute, base subcordate<br />
to obtuse or acute, margin entire, often more or<br />
less revolute, especially towards the base; upper<br />
surface scabrous, lower surface puberulous, on<br />
the main veins to hirtellous, sometimes also white<br />
arachnoid hairs present; lateral veins 6-7 pairs,<br />
straight or curved, basal pair unbr<strong>an</strong>ched, reach-<br />
22. <strong>Coussapoa</strong> longepedunculata Akkerm<strong>an</strong>s &<br />
C. C. Berg, Proc. Kon. Ned. Akad. Wetensch.<br />
Ser. C. 85(4): 453. 1982. Type. Peru. Loreto:<br />
Prov. Maynas, Quebrada Sucusari, Llachapa<br />
camp of Explorama, N side of Rio Napo, below<br />
Maz<strong>an</strong>, 6 Nov 1979 (6), Gentry et al. 27566<br />
(holotype, U; isotype, MO). Fig. 33.<br />
Tree, up to 25 m tall, hemi-epiphytic or terrestrial.<br />
Leafy twigs 1-1.5 cm thick, brown hirsute.<br />
Lamina coriaceous, oblong to elliptic, 15-<br />
50 x 7-30 cm, apex obtuse, base cordate to<br />
rounded margin rep<strong><strong>an</strong>d</strong> to subcrenate; upper sur-
64 Flora Neotropica<br />
FG2 3,si1e tg ts iore s l a 1 cm<br />
FIG. 32. <strong>Coussapoa</strong> leprieurii: 1, leafy twig <strong>with</strong> staminate inflorescences (Oldem<strong>an</strong> B1946); 2, pistillate<br />
inflorescences (Oldem<strong>an</strong> 3062).<br />
face scabridulous to smooth, sparsely hispidu-<br />
lous to hirtellous, often ? bullate, lower surface<br />
sparsely hirtellous on the veinlets, between the<br />
lateral veins white arachnoid hairs, rather dense<br />
or sparse <strong><strong>an</strong>d</strong> then concentrated at the margin;<br />
lateral veins 9-11 pairs, more or less curved, at<br />
least the lower ones br<strong>an</strong>ched, basal pair reaching<br />
the margin far below the middle of the lamina;<br />
intercostal venation prominent; petiole 3-10 cm<br />
long, brown hirsute; stipules 2-6 cm long, some-<br />
times subpersistent, hirsute to subsericeous, <strong>with</strong><br />
brown hairs of different lengths. Staminate in-<br />
florescences rather poorly br<strong>an</strong>ched; common<br />
peduncle 9-16 cm long, the br<strong>an</strong>ches in two clusters<br />
<strong><strong>an</strong>d</strong> all or some of them very short, peduncle<br />
<strong><strong>an</strong>d</strong> br<strong>an</strong>ches brown hirtellous to pubescent;<br />
heads m<strong>an</strong>y, (sub)globose, 3-5 mm diam.; peri<strong>an</strong>th<br />
ca. 1 mm high, sparsely minutely puberulous;<br />
stamens three, exceeding the peri<strong>an</strong>th. Pistillate<br />
inflorescences br<strong>an</strong>ched; heads 3-10,<br />
(sub)globose, 5-8 mm, in fruit up to 15 mm diam.;<br />
common peduncle 8-20 cm long, peduncle <strong><strong>an</strong>d</strong><br />
br<strong>an</strong>ches brown hirtellous to subtomentose; peri<strong>an</strong>th<br />
ca. 1 mm long, glabrous; fruiting peri<strong>an</strong>th<br />
red. Interfloral bracts ca. 1 mm long, spathulate,<br />
subpeltate, at the apex puberulous.
<strong>Coussapoa</strong> 65<br />
?~<br />
?~~~~~~~~~~~~~~~~~~~~~~~~~~~L<br />
~ ' ~/ ~ 1o g .<br />
' ',<br />
_ i , .,<br />
'?<br />
... ~ ~ ~ .'. ~ ~ ~~~~~~~~~~.<br />
,...<br />
: ~. -<br />
"'<br />
-r . . ' - ..;<br />
???~ ~~~~~~~~~~'<br />
?<br />
.'<br />
,, -<br />
?<br />
?'<br />
. .<br />
.. ? . . ... .<br />
or ?- -.' .<br />
*~~ -, ~ ~~~~ -..s<br />
??.?<br />
... ,,<br />
~~~~~~~~~~~~~~~~~~...<br />
-~~~~~~~~~~~~ :<br />
.'? ~ ..- ~ ~ *-. .I<br />
~~ ,. ....<br />
. ''' =, '? IY~~~<br />
~~~~~~ .~~~~~~~~~~~.<br />
I'<br />
r~~~~~~~~~~~~~~~~. o o-<br />
..,.<br />
I '~'~~~~~~~~~~~~~~~~~~~~~'<br />
~'C ?1 .'E":? .<br />
..?<br />
",<br />
'~~~~~~~~'<br />
.'<br />
-?~~~~1<br />
?~~~~~~~~~?<br />
FIG 33<br />
c.<br />
Cossaoa<br />
?<br />
onepeunclat: 1 lafytwi wih samiat iniorsceces(Gntr etal.2756)<br />
''.<br />
..
66 Flora Neotropica<br />
Distribution (Fig. 8). Peru (Loreto) <strong><strong>an</strong>d</strong> Ec-<br />
uador (Napo); in non-inundated or periodically<br />
inundated forest.<br />
Specimens studied. ECUADOR. NAPO: Parque Nacional<br />
Yasuni, 16-19 J<strong>an</strong> 1988 (d), Ceron M. 3495<br />
(BG), 9-19 J<strong>an</strong> 1988 (9), Neill et al. 8289 (BG).<br />
PERU. LORETO: Prov. Maynas, Quebrada Sucusari,<br />
Llachapa camp of Explorama, N side of Rio Napo,<br />
below Maz<strong>an</strong>, 6 Nov 1979 (a), Gentry et al. 27588 (MO,<br />
U, type collection); Prov. Maynas, Puerto Almendras,<br />
12 J<strong>an</strong> 1982 (9), Vdsquez 2852 (BG); Iquitos, Rio Itaya,<br />
Buena Suerte, 16 Nov 1986 (9), Vdsquez et al. 8380<br />
(BG); Nauta, 12 Dec 1986 (9), Vdsquez et al. 8600 (BG).<br />
C. longepedunculata resembles C. nymphaeifolia<br />
in m<strong>an</strong>y features. It is distinct from the<br />
latter especially in the indumentum, the wider<br />
(looser) venation, <strong><strong>an</strong>d</strong> in the scabridulous upper<br />
surface of the lamina. The species also shows<br />
affinities <strong>with</strong> C. sprucei <strong><strong>an</strong>d</strong> C. leprieurii.<br />
<strong><strong>an</strong>d</strong> allied species. In the shape <strong><strong>an</strong>d</strong> venation of<br />
the leaves it resembles C. contorta, from which<br />
it clearly differs in the 2-staminate flower.<br />
24. <strong>Coussapoa</strong> m<strong>an</strong>uensis C. C. Berg, Proc. Kon.<br />
Ned. Akad. Wetensch. Ser. C, 86(3): 305. 1983.<br />
Type. Peru. Madre de Dios: Parque Nacional<br />
del M<strong>an</strong>u, Rio M<strong>an</strong>u, Cocha Cashu Station,<br />
26 Oct 1980 (v), Foster 5641 (holotype, F).<br />
Fig. 35.<br />
Tree, hemi-epiphytic, 25 m tall. Lamina coriaceous,<br />
broadly elliptic to obovate, 10.5-19 x<br />
8-13 cm, apex shortly acuminate, base rounded<br />
to subcordate, margin entire to subcrenate; upper<br />
surface glabrous or sparsely puberulous at the<br />
base of the midrib, lower surface densely puberulous<br />
to hirtellous (to subtomentose); lateral<br />
veins 8-12 pairs, almost straight, the basal pair<br />
unbr<strong>an</strong>ched, reaching the margin below the mid-<br />
23. <strong>Coussapoa</strong> macerrima Akkerm<strong>an</strong>s & C. C. dle of the lamina; intercostal venation more or<br />
Berg, Proc. Kon. Ned. Akad. Wetensch. Ser. less prominent; petiole 3-5 cm long, puberulous<br />
C, 85(4): 455. 1982; Burger, Fieldi<strong>an</strong>a Bot. 40: to hirtellous; stipules ca. 6 cm long, chartaceous,<br />
133, t. 22. 1977 (sub C. contorta). Type. Costa subsericeous (to subhirsute), <strong>with</strong> rather con-<br />
Rica. Puntarenas: Above Palmar Norte de Osa, spicuous (parallel) venation. Staminate inflores-<br />
22 Feb 1951 (d), Allen 5949 (holotype, F; iso- cences unknown. Pistillate inflorescences untypes,<br />
A, F). Fig. 34. br<strong>an</strong>ched; heads globose, ca. 4 mm, in fruit up<br />
Tree, up to 25 m tall. Leafy twigs 2-4 mm<br />
to 12 mm diam.; peduncle 2.2-2.7 cm long, puthick,<br />
sparsely appressed-puberulous, on the scars berulous; peri<strong>an</strong>th ca. 1 mm high, sparsely miof<br />
the stipules distinctly longer hairs. Lamina nutely puberulous; fruiting peri<strong>an</strong>th yellow. Insubcoriaceous,<br />
elliptic to oblong to subobovate,<br />
terfloral bracts spathulate, puberulous at the apex.<br />
5-13 x 4-7 cm, apex shortly acuminate, base<br />
Distribution (Fig. 8). Peru (Madre de Dios).<br />
acute to obtuse, margin entire; upper surface<br />
Known<br />
glaonly<br />
from the type collection.<br />
brous, lower surface on the main veins rather<br />
C. m<strong>an</strong>uensis shows no distinct affinities <strong>with</strong><br />
sparsely appressed-puberulous <strong>with</strong> hairs of dif- <strong>an</strong>y of the other <strong>Coussapoa</strong> species.<br />
ferent lengths or partly strigillose; lateral veins<br />
25.<br />
7-13 pairs, straight to curved, basal pair un-<br />
<strong>Coussapoa</strong> microcarpa (Schott) Rizzini, Dusenia<br />
br<strong>an</strong>ched reaching the margin below the middle<br />
1(5): 295. 1950. Fig. 36.<br />
of the lamina; intercostal venation pl<strong>an</strong>e; petiole Brosimum microcarpon Schott in Sprengel, Syst. Veg.<br />
2-5 cm long, sparsely appressed-puberulous <strong>with</strong> 4 (Curr. Post., App.): 403. 1827. Type. Brazil. Withhairs<br />
of different<br />
out<br />
lengths; stipules 1-2 cm locality,<br />
long, (sub)sericeous. Staminate inflorescences<br />
br<strong>an</strong>ched; heads m<strong>an</strong>y, globose, ca. 1-2 mm<br />
diam.; common peduncle 2-3 cm long, appressed-puberulous;<br />
peri<strong>an</strong>th ca. 1 mm high, glabrous;<br />
stamens two, far exceeding the peri<strong>an</strong>th.<br />
Pistillate inflorescences unknown. Interfloral<br />
bracts absent.<br />
Distribution (Fig. 7). Costa Rica (Puntarenas);<br />
in forest at 450 m. Known only from the type<br />
collection.<br />
This species may be related to C. microcephala<br />
- (Y), Schott s.n. (holotype, W, destroyed).<br />
<strong>Coussapoa</strong> schottii Miquel, Fl. bras. 4(1): 137. 1853,<br />
as a synonym of Brosimum microcarpon.<br />
<strong>Coussapoa</strong> schottii Miquel var. l<strong>an</strong>ceolata Miquel, Fl.<br />
bras. 4(1): 137. 1853. Type. Brazil. Without locality,<br />
- (Y), Pohl s.n. (holotype, U; isotypes, B, M (herb.<br />
Zuccarini 105 sub forma <strong>an</strong>gustifolia)).<br />
<strong>Coussapoa</strong> schottii Miquel var. longifolia Miquel, Fl.<br />
bras. 4(1): 137. 1853. Type. Cult. bot. garden Minchen,<br />
1835 (2), herb. Zuccarini 103 (holotype, M).<br />
<strong>Coussapoa</strong> font<strong>an</strong>esi<strong>an</strong>a Trecul, Ann. Sci. Nat. Bot.<br />
Ser. 3, 8: 94. 1847. Syntypes. Brazil. Sao Paulo:<br />
Without locality, - (8 <strong><strong>an</strong>d</strong> 2), Gaudichaud 922 (lectotype<br />
(Q), P).
<strong>Coussapoa</strong> 67<br />
I<br />
1 cm<br />
FIG. 34. <strong>Coussapoa</strong> macerrima: 1, leafy twig <strong>with</strong> staminate inflorescences (Allen 5949)<br />
Shrub or tree, terrestrial or occasionally hemi-<br />
epiphytic, up to 20 m tall. Leafy twigs 2-5 mm<br />
thick, white to yellowish appressed-puberulous<br />
to hirtellous or to hirsute, sometimes also <strong>with</strong><br />
white to brownish arachnoid hairs. Lamina co-<br />
riaceous, subovate to ovate oblong to elliptic to<br />
(sub)obovate or to l<strong>an</strong>ceolate, 3-21 x 2-7 cm,<br />
apex shortly acuminate to acute or to obtuse,<br />
base acute to obtuse to rounded, margin entire,<br />
usually revolute towards the base; upper surface<br />
glabrous, lower surface glabrous or sparsely appressed-puberulous<br />
(to strigose) to (sub)hirsute
FIG.c3.m cousapoa <strong>an</strong>uenss 1 leves sipue n itlaeiforsecs(otr54)<br />
FIG. 35. <strong>Coussapoa</strong> m<strong>an</strong>uensis: 1, leaves, stipules <strong><strong>an</strong>d</strong> pistillate inflorescences (Foster 5641
<strong>Coussapoa</strong> 69<br />
"-; ~~~~~~~~~~~~~~~~~~~~~~'?<br />
?: ' ~C .. ?<br />
~~~~~~~~~~~~~~.. U ? :<br />
??<br />
: 7:I .i<br />
?~ ~~~~~~~~~~?<br />
.. 1~~~~~~~~~~~~~<br />
?: ~~~~~~~~~~~~~~~. ~ ~ ~ ~ : .?? ,.<br />
~~~~~~~~~~~~~~~~.2<br />
~~?. ?'''" .:?<br />
.. ".'<br />
I- ".i ii{<br />
' . .. ~<br />
FIG. 36. <strong>Coussapoa</strong> microcarpa: l, leafy twig <strong>with</strong> pistillate inflorescences (Mexia 5115); 2, leafy twig <strong>with</strong><br />
staminate inflorescence (Hage 31).<br />
on the main veins; lateral veins 6-11 pairs,<br />
straight, basal pair unbr<strong>an</strong>ched, reaching the<br />
margin below the middle of the lamina; intercostal<br />
venation pl<strong>an</strong>e to slightly prominent; petiole<br />
1-4 cm long, appressed-puberulous to hirtellous<br />
to (sub)hirsute; stipules 1-7 cm long,<br />
yellowish puberulous to subsericeous to hirsute<br />
or also <strong>with</strong> white to brownish arachnoid hairs.<br />
Staminate inflorescences br<strong>an</strong>ched; heads 5-9,<br />
globose, ca. 2-3 mm diam.; common peduncle<br />
1-2 cm long, puberulous to hirtellous; peri<strong>an</strong>th<br />
ca. 1 mm high, minutely puberulous; stamens<br />
two, far exceeding the peri<strong>an</strong>th. Pistillate inflo-<br />
rescences unbr<strong>an</strong>ched or rarely br<strong>an</strong>ched; heads<br />
1(-5), globose, ca. 3-5 mm, in fruit up to 10 mm<br />
diam.; (common) peduncle 2-5 cm long, pu-
70 Flora Neotropica<br />
berulous to hirtellous; peri<strong>an</strong>th ca. 1 mm high,<br />
glabrous; fruiting peri<strong>an</strong>th yellow <strong><strong>an</strong>d</strong> or<strong>an</strong>ge. Interfloral<br />
bracts small, narrowly spathulate, often<br />
only a few are sometimes absent.<br />
Distribution (Fig. 7). Brazil, from Rio Gr<strong><strong>an</strong>d</strong>e<br />
do Sul to Espirito S<strong>an</strong>to, common in forests up<br />
to 1800 m <strong><strong>an</strong>d</strong> in restinga vegetation; moreover,<br />
apparently in more or less isolated populations<br />
further north, in Bahia (near Ilheus) <strong><strong>an</strong>d</strong> in Paraiba<br />
<strong><strong>an</strong>d</strong> Pernambuco (on the top of low mountains?).<br />
Specimens studied. BRAZIL. Without (certain) locality,<br />
1814-1817 (9), Bowie & Cunningham 57 (BM),<br />
- (d <strong><strong>an</strong>d</strong> 2), Burchell 3148 (GH, K, P), - (6), Glaziou<br />
1942 (P), - (6), Glaziou 2016 (P), - (6), Glaziou 6009<br />
(C), - (9), Graham s.n. (K), - (2), Pohl s.n. (B, M<br />
(herb. Zuccarini 105), U, type collection of C. schottii<br />
var. l<strong>an</strong>ceolata), 11 Nov 1822 (d), Riedel s.n. (LE), 5<br />
Oct 1863 (st), Warming s.n. (C), 1835 (d), (cult. in hort.<br />
bot. Munchen), herb. Zuccarini 103 (M, type collection<br />
of C. schottii var. longifolia). BAHIA: Nr. Ilheus, 1 Apr<br />
1965 (2), Belem et al. 644 (U); <strong>with</strong>out locality, - Porto de Cima, 26 Jun 1914 (6), Jonsson 610a (F, GH,<br />
K, NY, S); Rio Sao Joao, W of road to Joinville, N of<br />
Garuva, 16 Jul 1966 (st), Lindem<strong>an</strong> et al. 1857 (U);<br />
Serra do Mar, nr. old road Curitiba-Morretes, Bella<br />
Vista, 25 Jul 1967 (st), Lindem<strong>an</strong> et al. 5682 (U); nr.<br />
Pontal do Sul, 20 Jun 1967 (st), Lindem<strong>an</strong> et al. 5748<br />
(U); Caioba, 35 km S of Par<strong>an</strong>agua, 10 Nov 1947 (9),<br />
Tessm<strong>an</strong>n s.n. (MO); Mun. Guaraqueqaba, Serrinha,<br />
13 Apr 1967 (6), Tessm<strong>an</strong>n s.n. (MO). PERNAMBUCO:<br />
"Gurgaf" (=Gurjao?), 14 May 1952 (8), Ducke & Lima<br />
104 (R). Rio DE JANEIRO: Rio de J<strong>an</strong>eiro,<br />
(9),<br />
Bl<strong>an</strong>chet s.n. (BM, M, U); nr. Ilheus, 1836 (2), Bl<strong>an</strong>chet<br />
2327 (G, M, P), 30 Nov 1970 (6), Emygdio 3018 (=Emmerich<br />
3556) (R); Itabuna, 16 Dec 1966 (9), Emygdio<br />
et al. 2447 (=Emmerich et al. 3002 = Andrade et al.<br />
2340) (R); nr. Ilheus, 23 Nov 1970 (6), Hage 31 (GUA,<br />
U), 24 Nov 1971 (2), Pinheiro 1716 (U); Castelo Novo,<br />
Oct 1821 (2), Riedel s.n. (LE), Mar 1822 (2), Riedel 664<br />
(LE). ESPiRITO SANTO: Mun. Linhares, Res. Biol. Sooretama,<br />
14 Mar 1972 (2), Sucre 8690 (U). MINAS GERMS:<br />
Lagoa S<strong>an</strong>ta, - (6), Lund 139 (C); Viqosa, Fazenda de<br />
Aguada, 29 Sep 1930 (2), Mexia 5115 (A, BM, F, G,<br />
GB, GH, K, MO, NY, S, U); Lagoa S<strong>an</strong>ta, - (?),<br />
Warming 1917 (C), 13 Oct 1863 (?), Warming 1918<br />
(C), 5 Oct 1863 (6), Warming 1919 (C), 29 Dec 1864<br />
(8), Warming 1942 (C). PARAiBA: Serro Araripe, S<strong>an</strong>ta<br />
Ana, Aug 1921 (2), Luetzelburg 12471 (M). PARANA:<br />
Mun. Guaratuba, Garuva, 28 Jul 1960 (9), Duarte et<br />
al. 5328 (F, RB); Jacarehy, 27 Sep 1908 (st), Dusen<br />
6635 (GH, NY); Caita de Agua, 28 Aug 1910 (st),<br />
Dusen 10134 (NY); Morretes, 7 Sep 1910 (2), Dusen<br />
10186 (F, GH, K, L, MO, NY, P, S); Jacarehy, 25 Mar<br />
1911 (6), Dusen 11419 (F, GH, NY); Rio Cubatao, 28<br />
Dec 1911 (8), Dusen 13653 (F, GH, NY, S); Jacarehy,<br />
26 Mar 1914 (6), Dusen 14718 (F, GH, K, MO, NY,<br />
P), 11 May 1915 (st), Dusen 17030 (NY); Cerro Azul,<br />
2 Oct 1949 (6), Hatschbach 1475 (S); Mun. Par<strong>an</strong>agua,<br />
Rio Cachoeirinha, 25 Aug 1951 (6), Hatschbach 2458<br />
(RB); Mun. Par<strong>an</strong>agua, Porto D. Pedro 2?, 31 J<strong>an</strong> 1961<br />
(8), Hatschbach 7807 (RB); Mun. Cerro Azul, Turvo,<br />
6 Feb 1961 (6), Hatschbach 7833 (RB); between Curitiba<br />
<strong><strong>an</strong>d</strong> Par<strong>an</strong>agua, 10 Nov 1948 (6), Hatschbach<br />
16304 (F, K); Mun. Guaraquecaba, Rio do Cedro, 8<br />
Nov 1968 (9), Hatschbach 18680 (C, MO, NY); Mun.<br />
Bocaiuva do Sul, Sesmaria, Rio Capivari, 11 Nov 1968<br />
(6), Hatschbach 20253 (C); Mun. Guaratuba, Rio Tupitinga,<br />
29 Apr 1972 (6), Hatschbach 29629 (NA); Par<strong>an</strong>agua,<br />
7 Mar 1914 (2), Jonnson 5a (F, GH, K, NY, S);<br />
- (2), Bl<strong>an</strong>chet<br />
s.n. (NY), 1835 (8), Bl<strong>an</strong>chet s.n. (FI); Itatiaia, 10<br />
J<strong>an</strong> 1910 (st), Campos Porto 863 (RB); Rio de J<strong>an</strong>eiro,<br />
Collegio Anglo-Brasileiro, 22 Sep 1916 (2), Const<strong>an</strong>tino<br />
(HJBR) 19689 (RB); Restinga de Itapeba, nr. C<strong>an</strong>al<br />
das Taxas, 22 May 1963 (2), Carauta 178 (GUA,<br />
K); Rio de J<strong>an</strong>eiro, Gavea, 11 Aug 1974 ($), Carauta<br />
1716 (F, GUA, K, U); Rio de J<strong>an</strong>eiro, Morro da Urca,<br />
17 Apr 1975 (st), Carauta 1780 (F), (2), Carauta 1781<br />
(GUA); Rio de J<strong>an</strong>eiro, Bot<strong>an</strong>ical Garden, 9 Nov 1978<br />
(2), Carauta 3050 (RB); Rio de J<strong>an</strong>eiro, Alto da Boa<br />
Vista, 7 Aug 1974 (2), Carauta et al. 1714 (F, GUA,<br />
RB); between Serra de Macah6 <strong><strong>an</strong>d</strong> Novo Friburgo,<br />
Sao Pedro, 18 Oct 1977 (2), Carauta et al. 2722 (GUA,<br />
U); Rio de J<strong>an</strong>eiro, Pao de Acucar, 26 Aug 1979 (2),<br />
Carauta et al. 3166 (GUA, U); Turin, 1857 (2), Casaretto<br />
643 (G); Joatinga Joa, 24 Mar 1959 ($), Duarte<br />
et al. 4650 (RB); Dois Irmaos, 19 Feb 1921 (2), Ducke<br />
et al. (HJBR) 16361 (RB); Meio Serra, old road to<br />
Petr6polis, 3 Nov 1928 (9), Ducke (HJBR) 21200 (RB);<br />
Serra de Itatiaia, Mont Serrat, 21 Oct 1903 (2), Dusen<br />
2160 (S); Rio de J<strong>an</strong>eiro, Prainha, 4 Dec 1978 (2),<br />
Ferreira 493 (RB); Serra dos Orgaos, Apr 1837 (6),<br />
Gardner 732 (BM, K); Rio de J<strong>an</strong>eiro, Gavea, Aug<br />
1837 (9), Gardner 5632 (BM, K); Teres6polis, 1100 m,<br />
31 J<strong>an</strong> 1978 (9), Gentry et al. 918 (MO, U); Rio de<br />
J<strong>an</strong>eiro, Gavea, 19 J<strong>an</strong> 1861 (st), Glaziou s.n. (P), 28<br />
Aug 1861 (st), Glaziou s.n. (P); Rio de J<strong>an</strong>eiro, Copacab<strong>an</strong>a,<br />
4 May 1866 (9), Glaziou 1013 (C, NY, P);<br />
Rio de J<strong>an</strong>eiro, Gavea, 28 Apr 1867 (6), Glaziou 1138<br />
(C, P); Rio de J<strong>an</strong>eiro, - (st), Glaziou 1881 (G); Rio<br />
de J<strong>an</strong>eiro, Gavea, 28 Nov 1869 (6), Glaziou 4937 (P);<br />
Rio de J<strong>an</strong>eiro, Copacab<strong>an</strong>a, 1871 (6), Glaziou 4938<br />
(C: K, P); Rio de J<strong>an</strong>eiro, Tijuca, 6 Feb 1871 (2), Glaziou<br />
6009 (K, P); Rio de J<strong>an</strong>eiro, 12 Aug 1876 (2),<br />
Glaziou 8936 (A, K, LE, P); Serra dos Orgaos, 23 Mar<br />
1880 (2), Glaziou 12166 (C, F, G, K, LE, P); Petr6polis,<br />
Bairro Amoeda, Dec 1943 (8), Goes et al. 815 (GUA,<br />
RB); Recreio dos B<strong><strong>an</strong>d</strong>eir<strong>an</strong>tes, 30 km W of Rio de<br />
J<strong>an</strong>eiro, 5 Mar 1964 (6), Lems s.n. (NY); Rio de J<strong>an</strong>eiro,<br />
1839 (6), Luschnath s.n. (LE); between Serra de<br />
Macah6 <strong><strong>an</strong>d</strong> Novo Friburgo, 1000 m, 18 Oct 1977 (2),<br />
Maas et al. 3325 (K, U); Petr6polis, Vale das Videiras,<br />
1800 m, 6 J<strong>an</strong> 1973 (2), Martinelli 150 (U); Rio de<br />
J<strong>an</strong>eiro, 1867 (8), Miers s.n. (BM); Rio de J<strong>an</strong>eiro,<br />
Tijuca, 1879 (2), Miers s.n. (BM); Rio de J<strong>an</strong>eiro, Jul<br />
1878 (6), Miers 2714 (K, P); Rio de J<strong>an</strong>eiro, Tijuca,<br />
Jul 1878 (9), Miers 3792 (K); Rio de J<strong>an</strong>eiro, Pedreiro,<br />
Jul 1878 (6), Miers 3858 (K, P); Restinga de Jacarepagua,<br />
Res. Biol., 12 Apr 1967 (6), Moreira 46 (F, GH,<br />
GUA, US); Rio de J<strong>an</strong>eiro, Corcovado, 20 Sep 1974<br />
(8), Mosen 2614 (S); Rio de J<strong>an</strong>eiro, Sep 1862 (9), Nadeaud<br />
s.n. (P); Rio de J<strong>an</strong>eiro, Copacab<strong>an</strong>a, Oct 1862
<strong>Coussapoa</strong><br />
(a), Nadeaud s. n. (P); Rio de J<strong>an</strong>eiro, Bot<strong>an</strong>ical Garden,<br />
8 Mar 1936 (9), Occhioni (HJBR) 35333 (RB); Serra<br />
dos Orgaos, Rio Paquequer, 4 Mar 1949 (9), Pereira<br />
39 (RB); Rio de J<strong>an</strong>eiro, Bot<strong>an</strong>ical Garden, 23 May<br />
1961 (9), Pereira 5670 (B, M); Rio de J<strong>an</strong>eiro, Gavea,<br />
3 Oct 1927 (9), Pessoal do Horto Florestal 654 (RB);<br />
Rio de J<strong>an</strong>eiro, Tijuca, 14 Jun 1969 (st), Plowm<strong>an</strong> 2919<br />
(=Sucre 5209) (GH, K, US); Rio de J<strong>an</strong>eiro, 1829 (a),<br />
Riedel 7 (LE), Sep-Dec 1831 (9), Riedel 75 (LE, NY,<br />
RB, US), 1832 (st), Riedel et al. s.n. (LE), 1832 (st),<br />
Riedel et al. I (LE); Teres6polis, Rio Imbui, 2 Oct 1952<br />
(9), Rizzini 144 (RB); Belem, Oct 1867 (9), Schwacke<br />
1067 (RB); Rio de J<strong>an</strong>eiro, Corcovado, 2 Nov 1883<br />
(9), Schwacke 6783 (RB), 11 Mar 1883 (2), Schwacke<br />
8418 (R); Rio de J<strong>an</strong>eiro, Copacab<strong>an</strong>a, 25 Oct 1967<br />
(9), Sucre 1783 (F, GUA, K, RB); Cabo Frio, Restinga<br />
do Per6, 14 Sep 1968 (9), Sucre 3633 (GUA, U); Restinga<br />
de Jacarapagua, 8 May 1969 (9), Sucre 4989 (U);<br />
Parades da Subida, Pedra da P<strong>an</strong>ela, 3 Nov 1971 (9),<br />
Sucre 7878 (U); Rio de J<strong>an</strong>eiro, Gavea, Nov 1899 (8),<br />
Ule s.n. (R); Rio de J<strong>an</strong>eiro, Copacab<strong>an</strong>a, - (9), Ule<br />
1693 (P); Itatiaia, 1935 (a), Zik<strong>an</strong> (HJBR) 29279 (RB).<br />
Rio GRANDE DO SUL: Fazenda das Almas, nr. Palmares,<br />
J<strong>an</strong> 1945 (8), Buck (in Rambo) 26425 (B); P6rto<br />
Alegre, 1 Oct 1957 (9), Camargo 1849 (B); Morro Teres6polis,<br />
Dec 1940 (8), Leite 413 (A), Dec 1942 (8),<br />
Leite 872 (A); P6rto Alegre, 2 Jun 1893 (9), Malme<br />
826 (S), 26 Feb 1902 (6), Malme 1428 (R, S), 16 Oct<br />
1932 (d <strong><strong>an</strong>d</strong> 9), Rambo 426 (F, MO, S), 10 Nov 1946<br />
(9), Rambo 27092 (FI, S), 2 Oct 1948 (9), Rambo 37782<br />
(C), 1 Dec 1948 (9), Rambo 38427 (B); Lami, nr. Viamao,<br />
3 J<strong>an</strong> 1949 (6), Rambo 39413 (F); Estacao Pereci,<br />
14 J<strong>an</strong> 1949 (9), Rambo 39780 (P); Ilha das Flores, 22<br />
Apr 1949 (9), Rambo 41173 (C, K, US); Lagoa dos<br />
Barros, nr. Osoria, 14 Feb 1949 (9), Rambo 44743 (P);<br />
Fazenda do Arroio, nr. Osoria, 4 J<strong>an</strong> 1950 (a), Rambo<br />
45136 (K); Lagoa dos Quadros, nr. Torres, 21 Feb 1950<br />
(a), Rambo 45904 (K); nr. Tram<strong><strong>an</strong>d</strong>ai, 5 Mar 1950 (a),<br />
Rambo 46145 (P); <strong>with</strong>out locality, - NY, S, U), 20 Dec 1952 (6), Reitz et al. 5030 (F, G,<br />
NY, S, U); Mina Velha, Garuva, Sao Fr<strong>an</strong>cisco do Sul,<br />
8 Nov 1957 (9), Reitz et al. 5611 (B, US); Serra do<br />
Matador, Rio do Sul, 26 J<strong>an</strong> 1959 (2), Reitz et al. 8294<br />
(G, M); Ilha da S<strong>an</strong>ta Catarina, 6 Mar 1962 (8), Sehnem<br />
7993 (B); P6rto Belo, Ilha Joao da Cunha, 31 Mar 1957<br />
(Q), L. B. Smith et al. 12307 (NY, R, US). SAO PAULO:<br />
Mun. Picinguaba, Picinguaba beach, 2 Oct 1975 (9),<br />
Araujo et al. 848 (RB); Mun. Iguape, Morro das Pedras,<br />
Aug 1915 (8), Brade 7887 (R), 1917 (9), Brade 7949<br />
(R, RB); Mun. Iguape, Iguape Isl<strong><strong>an</strong>d</strong>, 19 Feb 1965 (2),<br />
Eiten et al. 6214 (MO, US); <strong>with</strong>out locality, 1917 (6),<br />
Frazas(HJBR) 13059 (RB); 1833 (2), Gaudichaud 992<br />
(P); Sao Paulo, Parque do Estado, 4 Sep 1933 (6), Hoehne<br />
30923 (F, GUA, NY, P); road Caraguatatuba-Ubatuba,<br />
10 Oct 1968 (9), Leitao Filho 685 <strong><strong>an</strong>d</strong> 686 (GUA);<br />
S<strong>an</strong>tos, 1 Dec 1874 (2), Mosen 2941 (C, S).<br />
This species appears to be related to C. latifolia<br />
<strong><strong>an</strong>d</strong> C. microcephala. It is quite variable in the<br />
shape <strong><strong>an</strong>d</strong> dimensions of the lamina. A form <strong>with</strong><br />
relatively long <strong><strong>an</strong>d</strong> narrow leaves is cultivated in<br />
several Europe<strong>an</strong> greenhouses <strong><strong>an</strong>d</strong> (in The Netherl<strong><strong>an</strong>d</strong>s)<br />
as <strong>an</strong> indoor ornamental.<br />
26. <strong>Coussapoa</strong> microcephala Tr6cul, Ann. Sci.<br />
Nat. Bot., Ser. 3, 8: 96. 1847; Miquel, Fl. bras.<br />
4(1): 136. 1853; Berg, Fl. Suriname 5(1): 282.<br />
1975. Type. Guy<strong>an</strong>a. Pomeroon River,<br />
(9), Sello 241<br />
(B, P, US); Rio Tapuchy, 1837 (6), Tweedie 29 (K);<br />
Col6nia Sao Pedro Torres, 12 Nov 1968 ($), Vi<strong>an</strong>na<br />
et al. 5465 (U). SANTA CATARINA: Rio Claro, Linha de<br />
Joinville, 30 Oct 1882 (6), Anonymus s.n. (RB); Ilha<br />
da S<strong>an</strong>ta Catarina, 8 Dec 1950 (9), Duarte et al. 3410<br />
(F, K, NY, RB, U); Ibirama, 13 Dec 1953 (8), Gevieski<br />
85 (B, NY, S, US); Rio Pirai, 6 J<strong>an</strong> 1950 (6), H<strong>an</strong>s 335<br />
(R, RB); Brusque, Mata Sao Pedro, 30 Dec 1949 (a),<br />
Klein 79B (= Veloso 97) (RB, US); Morro de Fazenda,<br />
Itajai, 25 Mar 1954 (6), Klein 772 (S, US); Morro dos<br />
Conventos, E of Arar<strong>an</strong>gua, 5 Nov 1972 (6), Lima<br />
20795 (U), 15 Nov 1971 (9), Lindem<strong>an</strong> et al. 9105 <strong><strong>an</strong>d</strong><br />
9110 (U); Sombrio, 3 Sep 1945 (8), Reitz 1912 (F, R,<br />
NY); Sao Fr<strong>an</strong>cisco do Sul, 9 J<strong>an</strong> 1951 (9), Reitz 3809<br />
(US); Campo do Massiambfi, Palhoca, 24 Sep 1953<br />
(a), Reitz et al. 975 (NY, S); Morro de Fazenda, Itajai,<br />
3 Mar 1954 ($), Reitz et al. 1688 (NY, S, US); Sombrio,<br />
8 Feb 1946 (8), Reitz et al. 2008 (US); Cunhas, Itajai,<br />
5 Aug 1954 (8), Reitz et al. 2027 (US); Morro de Ressacada,<br />
Itajai, 29 Mar 1957 (9), Reitz et al. 2913 (US);<br />
Morro do Bau, Itajai, 1 Nov 1951 (6), Reitz 4167 (NY,<br />
S); Trds Barras, Garuva, Sao Fr<strong>an</strong>cisco do Sul, 24 Aug<br />
1957 (9), Reitz et al. 4692 (NY); Campo do Massiambi,<br />
Palhoca, 19 Dec 1952 (9), Reitz et al. 4965 (F, G,<br />
- (6),<br />
Schomburgk 876 (holotype, P; isotypes, BM,<br />
F, FI, NY). Fig. 37.<br />
<strong>Coussapoa</strong> fagifolia Klotzsch, Linnaea 20: 528. 1847;<br />
Miquel, Fl. bras. 4(1): 136. 1853. Type. Guy<strong>an</strong>a.<br />
Pomeroon River, - (8), Schomburgk 1366 (=876?)<br />
(holotype, B; isotype, K).<br />
<strong>Coussapoa</strong> cuneata Miquel, Fl. bras. 4(1): 138. 1853.<br />
Type. Guy<strong>an</strong>a. Berbice River, - (6), Schomburgk<br />
287 (holotype, U; isotypes, BM, G, GH, K, L, P,<br />
OXF, U).<br />
Tree, often hemi-epiphytic, up to 20 m tall.<br />
Leafy twigs 2-9 mm thick, sparsely to densely<br />
yellowish appressed-puberulous to hirtellous to<br />
subhirsute or also <strong>with</strong> white to brown arachnoid<br />
hairs. Lamina coriaceous (to subcoriaceous),<br />
ovate to subovate or elliptic to oblong, 3-27 x<br />
1.5-16 cm, apex acuminate to acute, base rounded<br />
to obtuse, margin entire, + revolute towards<br />
the base; upper surface glabrous, lower surface<br />
densely to very sparsely appressed-puberulous<br />
(on the main veins <strong>with</strong> hairs of different lengths)<br />
or also densely to sparsely covered <strong>with</strong> white to<br />
pale brown arachnoid hairs, disappearing or more<br />
or less persistent on the main veins <strong><strong>an</strong>d</strong> the margin;<br />
lateral veins 4-11 pairs, straight or slightly<br />
curved, basal pair br<strong>an</strong>ched or unbr<strong>an</strong>ched, in<br />
71
FIG. 37. <strong>Coussapoa</strong> microcephala: ssp. microcephala: 1, leafy twig <strong>with</strong> pistillate inflorescences (Maguire 24588); 2, lea<br />
(Gleason 405).<br />
I<br />
1cm
<strong>Coussapoa</strong><br />
ovate to elliptic leaves reaching the margin below<br />
or sometimes at the middle of the lamina; intercostal<br />
venation more or less prominent to<br />
pl<strong>an</strong>e; petiole 0.5-5 cm long, appressed-puberulous<br />
to hirtellous or also <strong>with</strong> arachnoid hairs;<br />
(U); Pakaraima Mts., Kamar<strong>an</strong>g, 8 Sep 1979 (6), Maas<br />
et al. 4126 (U); Kaieteur Gorge, 14 May 1944 (6),<br />
Maguire et al. 23449 (F, G, K, MO, NY, P, U, US);<br />
Demerara River, Comaka, Apr 1933 (6), Persaud 222<br />
(F); Essequibo River, Moraballi Creek, nr. Bartica, 18<br />
Sep 1929 (8), S<strong><strong>an</strong>d</strong><strong>with</strong> 290 (K, NY, RB), 19 Sep 1929<br />
stipules 1.2-5 cm long, yellowish puberulous to (2), S<strong><strong>an</strong>d</strong><strong>with</strong> 302 (K, U), 7 Oct 1929 (6), S<strong><strong>an</strong>d</strong><strong>with</strong> 394<br />
subsericeous, often also <strong>with</strong> brownish arachnoid (K, P, US); <strong>with</strong>out locality,<br />
hairs; terminal buds slender. Staminate inflorescences<br />
br<strong>an</strong>ched; heads six to m<strong>an</strong>y, globose, ca.<br />
2 mm diam.; common peduncle 1-4 cm long,<br />
puberulous; peri<strong>an</strong>th ca. 1 mm high, glabrous;<br />
stamens two, far exceeding the peri<strong>an</strong>th. Pistillate<br />
inflorescences br<strong>an</strong>ched, occasionally unbr<strong>an</strong>ched;<br />
heads (1-)3-15, sometimes fused, globose,<br />
2-5 mm, in fruit up to 10 mm diam.;<br />
(common) peduncle 1-10 cm long, puberulous;<br />
peri<strong>an</strong>th ca. 1 mm high, glabrous. Interfloral<br />
bracts (sub)spathulate, minutely puberulous at<br />
the apex.<br />
Distribution (Fig. 7). Common in Guy<strong>an</strong>a, apparently<br />
rare in Surinam <strong><strong>an</strong>d</strong> French Gui<strong>an</strong>a; in<br />
forest, often along streams, at altitudes up to<br />
900 m.<br />
- (?), Schomburgk s.n.<br />
(F, G), - (6), Schomburgk 118 (B, K), - (9), Schomburgk<br />
167 (G, K, P); Berbice River, 1837 (6), Schomburgk<br />
287 (BM, G, GH, K, L, OXF, P, U, type collection<br />
of C. cuneata); Pomeroon River (?), - (6),<br />
Schomburgk 876 (by error 276) (BM, F, FI, G, NY, P,<br />
type collection of C. microcephala), Schomburgk 1366<br />
(=?876) (B, K, type collection of C. fagifolia); Potaro<br />
River, nr. Amatuk, 26 Aug 1959 (8), Whitton 179 (K).<br />
SURINAM. Without locality, - (9), Anderson s.n.<br />
(BM); Tafelberg, 1 Sep 1944 (9), Maguire 25488 (A, F,<br />
G, K, MO, NY, U, US).<br />
FRENCH GUIANA. Road to Brazil, km 6, at bridge<br />
over Compt6 River, ca. 51 km S of Cayenne, 1 J<strong>an</strong><br />
1977 (2), Mori 8864 (CAY, MO, NY, U).<br />
Specimens studied. GUYANA. Northwest District,<br />
Kaituma River, 27 Oct 1908 (a), Anderson 188 (K,<br />
NY); Cuyounia Creek, - (8), Appun 296 (K); Potaro<br />
River, nr. Kaieteur Falls, 18 Feb 1962 ($), Cow<strong>an</strong> et<br />
al. 1868 (F, K, US), 18 Feb 1962 (9), Cow<strong>an</strong> et al. 1880<br />
(F, K, US); Northwest District, Anabisi River, 15 Feb<br />
1922 (9), De la Cruz 1367 (GH, NY); upper Rupununi<br />
River, nr. Dad<strong>an</strong>awa, 14 Jun 1922 (a), De la Cruz 1501<br />
(F, GH, MO, NY); between Demerara River <strong><strong>an</strong>d</strong> Ber-<br />
bice River, 5?50'N, 15-19 Jul 1922 (6), De la Cruz<br />
1577 (F, GH, MO, NY); upper Mazaruni River, ca.<br />
60?10'W, 22 Sep-6 Oct 1922 (8), De la Cruz 2241 (F,<br />
GH, MO, NY, US), De la Cruz 2387 (F, GH, MO,<br />
NY, US); Demerara River, Malali, 30 Oct-5 Nov 1922<br />
(6), De la Cruz 2636 (F, GH, K, NY); upper Mazaruni<br />
River, Kamakusa, 23-29 Nov 1922 (9), De la Cruz<br />
2813 (F, GH, MO, NY); Demerara River, Malali, 30<br />
Oct-5 Nov 1922, De la Cruz 3070 (NY); Essequibo<br />
River, Winiperu Creek, 2 Nov 1939 (9), F<strong>an</strong>shawe 295<br />
(FD 3031) (K, NY, U); Mazaruni River, Kurupung<br />
River, Sep 1925 (a), Gleason 405 (K, NY); Potaro Riv-<br />
er, Tumatumari, 4-6 Jul 1921 (9), Gleason 415 (GH,<br />
K, NY); Potaro River, nr. Kaieteur Falls, 17 J<strong>an</strong> 1954<br />
(a), Irwin 183 (US); Mazaruni River, Sep 1880 (9),<br />
Jenm<strong>an</strong> 652 (K); upper Demerara River, nr. Great<br />
Falls, Sep 1887 (2), Jenm<strong>an</strong> 3995 (K, NY); Berbice<br />
River, May 1889 (2), Jenm<strong>an</strong> 4947 (K), <strong>with</strong>out lo-<br />
cality, Jun 1889 (e), Jenm<strong>an</strong> 5315 (K); Demerara Riv-<br />
er, Mar 1898 (9), Jenm<strong>an</strong> 7310 (K, S, U); Pomeroon<br />
River, nr. Macasuma, Feb 1904 (e), Jenm<strong>an</strong> 7933 (K);<br />
Kupurung River, 22 Nov 1922 (9), Leng 177 (NY);<br />
Waine River, 10-15 km S of Kwab<strong>an</strong>na, 10 Aug 1977<br />
(9), Maas et al. 2478 (F, K, U); S of Timehri, Mr.<br />
Thompson's farm, 17 Oct 1979 (a), Maas et al. 3622<br />
73<br />
<strong>Coussapoa</strong> microcephala is closely related to<br />
C. argentea. Because of more or less distinct sim-<br />
ilarities in the dimensions, shape, margin, <strong><strong>an</strong>d</strong><br />
venation of the leaves these two species together<br />
<strong>with</strong> C. latifolia, C. viridifolia, C. parvifolia, C.<br />
microcarpa, C. pachyphylla (<strong><strong>an</strong>d</strong> C. macerrima)<br />
constitute a more or less distinct group <strong>with</strong>in<br />
the taxa having 2-staminate flowers.<br />
27. <strong>Coussapoa</strong> napoensis Akkerm<strong>an</strong>s & C. C.<br />
Berg, Proc. Kon. Ned. Akad. Wetensch., Ser.<br />
C. 85(4): 457. 1982. Type. Ecuador. Napo:<br />
Coca, 17 J<strong>an</strong> 1973 (6), Lugo 2820 (holotype,<br />
GB). Fig. 38.<br />
Tree, up to 25 m tall, terrestrial or hemi-epiphytic.<br />
Leafy twigs 5-12 mm thick, sparsely puberulous,<br />
<strong><strong>an</strong>d</strong> mostly also <strong>with</strong> sparse, distinctly<br />
longer, straight to curved, stiff hairs. Lamina coriaceous,<br />
broadly ovate to broadly elliptic, 7-30<br />
x 6-22 cm, apex obtuse, occasionally acuminate,<br />
base subobtuse to rounded or to truncate,<br />
margin entire to subcrenate; upper surface glabrous,<br />
lower surface on the main veins <strong>with</strong> appressed<br />
straight hairs of different lengths <strong><strong>an</strong>d</strong> <strong>with</strong><br />
sparse arachnoid hairs which later disappear; lateral<br />
veins (5-)7-12(-15) pairs, almost straight,<br />
basal pair (sometimes faintly) br<strong>an</strong>ched, reaching<br />
the margin below the middle of the lamina; intercostal<br />
venation slightly prominent to almost<br />
pl<strong>an</strong>e; petiole 3-10 cm long, <strong>with</strong> appressed<br />
straight hairs of different lengths, glabrescent;<br />
stipules 4-13 cm long, subvelutinous, also <strong>with</strong><br />
distinctly longer straight stiff hairs. Staminate
74 Flora Neotropica<br />
~.I.<br />
FIG. 38. <strong>Coussapoa</strong> napoensis: 1, leafy twig <strong>with</strong> staminate inflorescences<br />
(Lugo 2857).<br />
I cm.
<strong>Coussapoa</strong> 75<br />
inflorescences repeatedly br<strong>an</strong>ched; heads m<strong>an</strong>y,<br />
globose, 2-3 mm diam.; common peduncle 2.5-<br />
6 cm long, puberulous to hirtellous; peri<strong>an</strong>th ca.<br />
0.5-1 mm high, glabrous; stamen one, far exceeding<br />
the peri<strong>an</strong>th. Pistillate inflorescences<br />
br<strong>an</strong>ched; heads 3-9, subglobose, 6-10 mm<br />
diam.; common peduncle 3-4 cm long, puberulous<br />
to subhirsute; peri<strong>an</strong>th ca. 1 mm high, glabrous.<br />
Interfloral bracts absent.<br />
Distribution (Fig. 7). Ecuador (Napo) <strong><strong>an</strong>d</strong> Colombia<br />
(Putumayo); Amazoni<strong>an</strong> forest.<br />
Specimens studied. COLOMBIA. PUTUMAYO: Rio<br />
Putumayo, Pifiufia Negro, 20 Nov 1940 (6), Cuatrecasas<br />
10714 (F, US).<br />
ECUADOR. NAPO: Nr. Coca, 17 J<strong>an</strong> 1973 (6), Lugo<br />
2820 (GB, type collection), 22 J<strong>an</strong> 1973 (6), Lugo 2909<br />
(GB); Rio Payamino, nr. Payamino Capihuara, 19 J<strong>an</strong><br />
1973 (6), Lugo 2857 (GB); Shushufindi, 4 Aug 1975<br />
(e), Little et al. 43 (Q); 20 km W of Coca, 22-23 April<br />
1985 (v), Palacios et al. 6387 (BG).<br />
This species has m<strong>an</strong>y features in common<br />
<strong>with</strong> C. ovalifolia. It is distinct from the latter<br />
mainly in the longer stipules (of fertile twigs!),<br />
the smaller number of lateral veins, <strong><strong>an</strong>d</strong> the larger<br />
flower heads of the staminate inflorescence. The<br />
two species could prove to be distinct only at the<br />
subspecific level.<br />
28. <strong>Coussapoa</strong> nitida Miquel, Fl. bras. 4(1): 113,<br />
t. 44. 1853. Type. Brazil. Para: "Jaguary"<br />
(=Jaguarari?), - (e), Martius s.n. (lectotype,<br />
M; isolectotypes, B?, BR, U). Fig. 39.<br />
<strong>Coussapoa</strong> intermedia Miquel, Fl. bras. 4(1): 133, t.<br />
43. 1853. Type. Brazil. Without locality ("prov. Paraensi<br />
et Rio Negro"), - sparsely (to rather densely) covered <strong>with</strong> arachnoid<br />
hairs which are deciduous; lateral veins 8-<br />
15 pairs, mostly straight, basal pair (normally)<br />
br<strong>an</strong>ched, reaching the margin below the middle<br />
of the lamina; intercostal venation (almost) pl<strong>an</strong>e;<br />
petiole 3-13 cm long, 2-4 mm thick, densely<br />
brownish to white puberulous <strong><strong>an</strong>d</strong> <strong>with</strong> distinctly<br />
longer yellowish appressed to patent hairs; stipules<br />
2.5-10 cm long, yellowish to brownish subsericeous<br />
to hirsute or predomin<strong>an</strong>tly densely<br />
puberulous. Staminate inflorescence br<strong>an</strong>ched;<br />
heads numerous, free, globose, ca. 2 mm diam.,<br />
often several solitary flowers on the br<strong>an</strong>ches of<br />
the inflorescence; common peduncle 2-5 cm long,<br />
rather densely puberulous to hirtellous to subhirsute;<br />
peri<strong>an</strong>th ca. 1 mm high, glabrous or minutely<br />
puberulous; stamen one, far exceeding the<br />
peri<strong>an</strong>th. Pistillate inflorescence br<strong>an</strong>ched; heads<br />
2-7, free, globose, ca. 4-6, in fruit up to 20 mm<br />
diam.; common peduncle 2-7 cm long, peduncle<br />
<strong><strong>an</strong>d</strong> br<strong>an</strong>ches 1-2 mm thick, rather densely puberulous<br />
to hirtellous, apices of the br<strong>an</strong>ches often<br />
more or less broadened towards the heads; peri<strong>an</strong>th<br />
ca. 1 mm high, sparsely minutely white puberulous<br />
to glabrous. Interfloral bracts absent.<br />
Distribution (Fig. 8). Amazon Basin (Brazil<br />
<strong><strong>an</strong>d</strong> Peru), especially along the Amazon River;<br />
mostly in riverside (varzea) forest.<br />
Specimens studied. COLOMBIA. AMAZONAS: Rio<br />
Caqueta, Puerto Cordoba, 13 Nov 1912 (8), Ducke<br />
(HAMP) 12238 (MG); Trapecio Amazonico, nr. mouth<br />
of Rio Loretoyacu, 26 Nov 1945 (2), Duque-Jaramillo<br />
2207 (COL); Rio Loretoyacu, Oct 1945 (9), Schultes<br />
6726 (F, GH), Sep 1946 (2), Schultes et al. 8380 (F).<br />
(8), Martius s.n.<br />
PERU. LORETO: Rio<br />
(holotype,<br />
Itaya, ca. 5 km above Iquitos,<br />
6<br />
M; isotype, B).<br />
Aug 1972 (a), Croat 18850 (MO, U); Prov. Maynas,<br />
<strong>Coussapoa</strong> schunkei St<strong><strong>an</strong>d</strong>ley, Publ. Field Mus.<br />
Rio<br />
Bot. Ampiyacu, between Pebas <strong><strong>an</strong>d</strong> mouth of Rio Ya-<br />
22: 72. 1940. Type. Peru. Loreto: Rio Maz<strong>an</strong>, Gam- guasyacu, 7 Nov 1977 (2), Gentry et al. 20381 (MO,<br />
it<strong>an</strong>acocha, 27 J<strong>an</strong> 1935 (6), Schunke 130 (holotype,<br />
U); Rio Ampiyacu, above junction of Rio Yaguasyacu,<br />
9 Nov<br />
F;<br />
1977<br />
isotypes, A, NY, US).<br />
(2), Gentry et al. 20495 (MO, U); Rio<br />
Yavari, opposite Paumari, 23 Nov 1977 (a), Gentry et<br />
Tree, hemi-epiphytic or terrestrial, up to 30 m al. 20802 (U); Rio N<strong>an</strong>ay, Mish<strong>an</strong>a, halfway between<br />
tall. Leafy twigs 5-12 mm thick, densely brown- Iquitos <strong><strong>an</strong>d</strong> S<strong>an</strong>ta Maria de N<strong>an</strong>ay, 26 Feb 1979 (st),<br />
ish<br />
Gentry et al. 25124<br />
puberulous to<br />
(MO, U); lower Rio<br />
shortly velutinous, often<br />
Mom6n, nr.<br />
partly Iquitos, 8 Dec 1979 (9), J. Jones et al. 9712 (LAM, U),<br />
(sub)hirsute. Lamina (sub)coriaceous, ovate (to 9 Dec 1979 (8), J. Jones et al. 9769 (LAM, U); Rio<br />
elliptic), 8-30 x 4-20 cm, apex acute to obtuse, Ampiyacu, Puca Urquillo, 2 Apr 1977 (9), Plowm<strong>an</strong><br />
base obtuse to rounded to truncate to et al. 6555<br />
cordate,<br />
(F, GH, K, U); Rio Ampiyacu, Pebas, 16<br />
Jul 1976<br />
margin entire to subcrenate; upper surface (a), Revilla 805 (MO); Rio Mom6n, Mogla-<br />
noncillo, 17 Aug 1976 (9), Revilla 1099<br />
brous, lower<br />
(MO,<br />
surface<br />
U); Rio<br />
densely minutely puberu- Mom6n, above Bellavista, 19 Sep 1975 (9), Rimachi<br />
lous in the areoles <strong><strong>an</strong>d</strong> on the reticulum, usually 1943 (NA); Rio Maz<strong>an</strong>, Gamit<strong>an</strong>acocha, 27 J<strong>an</strong> 1935<br />
sparsely puberulous to hirtellous on the (parallel) (8), Schunke 130 (A, F, NY, US, type collection of C.<br />
tertiary venation, (appressed-)puberulous to stri- schunkei).<br />
BRAZIL. Without<br />
gose or to sparsely hirsute on<br />
locality (Prov. Paraensi et Rio<br />
the midrib <strong><strong>an</strong>d</strong> Negro),<br />
lateral veins, sometimes the whole surface<br />
- (a), Martius s.n. (B, M, type collection of C.<br />
intermedia); - (a + 9, monoecious!), Poeppigs.n. (BR).
76 Flora Neotropica<br />
FIG. 39. <strong>Coussapoa</strong> nitida: 1, leafy twig <strong>with</strong> pistillate inflorescences<br />
(Schultes 6726).
<strong>Coussapoa</strong><br />
ACRE: Rio Purus, nr. mouth of Rio Macaua, tributary<br />
of Rio Iaco, 21 Aug 1933 (6), Krukoff5596 (A, BM,<br />
F, G, K, LE, M, MO, NY, Q, S, U, US). AMAPA: Rio<br />
Jari, 2 km E of Arum<strong><strong>an</strong>d</strong>uba, 27 Jun 1961 (2), W. A.<br />
Egler et al. 46027A (IAN, MG, MO, NY, S, U).<br />
AMAZONAS: Rio Solim6es, B6ca de M<strong>an</strong>aquiri, 3 Oct<br />
1973 (6), Berg et al. P17591 (F, M, MO, P, S, U), 3<br />
Oct 1973 (2), Berg et al. P17592 (F, MO, K, P, S, U);<br />
Rio Japura, Mun. Maraa, Ilha dos Macacos, 30 Oct<br />
1977 (8), Damiao 2503 (INPA); Rio Japura, Costa do<br />
Jacitaria, 8 Dec 1977 (8), Damido 2817 (INPA); Rio<br />
Solimoes, Fonte Boa, 9 Oct 1945 (6), Ducke 1795 (A,<br />
F, K, MG, NY, R, RB, US); Rio Japura, 19 Sep 1904<br />
(9), Ducke (HAMP) 6782 (S, U); Rio Ia, 9 Sep 1906<br />
(9), Ducke (HAMP) 7715 (BM, G); Rio Ton<strong>an</strong>tins, 26<br />
Oct 1949 (6), Frogs 25531 (IAN); Mun. Sao Paulo de<br />
Olivenca, nr. Palmares, 11 Sep-26 Oct 1936 (6), Krukoff<br />
8401 (A, B, BM, F, G, K, LE, NY, P, S, U, US);<br />
Rio Solimoes, Mamia, 20 J<strong>an</strong> 1924 (8), Kuhlm<strong>an</strong>n 1178<br />
(RB); Rio Javari, behind Palmeiras (army post), 6 Aug<br />
1973 (8), Lleras et al. P17197 (C, K, M, MO, NY, U,<br />
US). PARA: Rio Tajapurir, Ant6nia Lemos, Igarape Pixuna,<br />
19 Jul 1948 (2), Black 48-2949 (IAN); Rio Tajapuri,<br />
Ilha Sao Sebastiao, Nazar6, 31 Jul 1948 (6),<br />
Black 48-3017 (IAN); Gurupa, 27 J<strong>an</strong> 1916 (6), Ducke<br />
(HJBR) 13066 (=HAMP 16003) (BM, RB); Rio Tajapurfi,<br />
- 21 cm, apex acute to obtuse to shortly acuminate,<br />
base cordate, margin entire or subcrenate; upper<br />
surface glabrous, lower surface densely minutely<br />
puberulous in the areoles <strong><strong>an</strong>d</strong> on the reticulum,<br />
(rather) sparsely hirtellous to puberulous on the<br />
other veins, on the main veins <strong><strong>an</strong>d</strong> margin also<br />
sparse white arachnoid hairs; lateral veins 9-11<br />
pairs, slightly curved, basal pairs br<strong>an</strong>ched,<br />
reaching the margin below the middle of the lamina,<br />
other lateral veins often poorly br<strong>an</strong>ched<br />
(furcate); intercostal venation very prominent;<br />
petiole 8-13 cm long, sparsely to densely minutely<br />
puberulous <strong><strong>an</strong>d</strong> sparsely covered <strong>with</strong> distinctly<br />
longer straight stiff hairs; stipules 0.5-3.5<br />
cm long, brownish subsericeous <strong><strong>an</strong>d</strong> <strong>with</strong> a dense<br />
covering of reddish-brown pluricellular hairs.<br />
Staminate inflorescences br<strong>an</strong>ched; heads m<strong>an</strong>y,<br />
globose, ca. 4-6 mm diam.; common peduncle<br />
1.5-3 cm long, puberulous to hirtellous, on the<br />
br<strong>an</strong>ches to tomentellous, mixed <strong>with</strong> sparse to<br />
dense reddish-brown pluricellular hairs; peri<strong>an</strong>th<br />
(), Martius s.n. (FI); "Jaguary" (=?Jaguar- ca. 1 mm high, densely minutely puberulous;<br />
ari), Aug (9), Martius s.n. (B, BR, M, U, lectotype col- stamens three, exceeding the peri<strong>an</strong>th. Pistillate<br />
lection), "Insulae Archipelagi Paraensis," Aug (d <strong><strong>an</strong>d</strong> inflorescences unbr<strong>an</strong>ched; heads ellipsoid to<br />
2), Martius s.n. (or 2673) (M, L, LE); Rio Jari, between<br />
Monte Dourado <strong><strong>an</strong>d</strong> Caracura, 18 Nov 1967 (9), Oliv- obovoid, ca. 20 x 10, in fruit up to 15 mm diam.;<br />
eira 3627 (IAN, NY); Rio Jari, between Monte Dour- peduncle 1-2 cm long, reddish-brown puberuado<br />
<strong><strong>an</strong>d</strong> Patricia, 27 Mar 1970 (6), N. T. Silva 303 lous to hirtellous; peri<strong>an</strong>th ca. 2 mm high, dense-<br />
(IAN).<br />
ly reddish-brown puberulous. Interfloral bracts<br />
subpeltate, often only a few or sometimes absent.<br />
Distribution (Fig. 8). Costa Rica (Alajuela); in<br />
rain forest up to 800 m.<br />
<strong>Coussapoa</strong> nitida c<strong>an</strong> easily be confused <strong>with</strong><br />
C. tessm<strong>an</strong>nii, due to the strong overall similar-<br />
ities. However, C. nitida is clearly distinct from<br />
the latter in the absence ofinterfloral bracts. Oth-<br />
er differences occur in the indumentum, espe-<br />
cially in that of the petiole, <strong><strong>an</strong>d</strong> in the diameter<br />
of the peduncle of the pistillate inflorescence. C.<br />
nitida appears to be related to C. cupularis <strong><strong>an</strong>d</strong><br />
more remotely to C. orthoneura <strong><strong>an</strong>d</strong> C. arach-<br />
noidea.<br />
29. <strong>Coussapoa</strong> nymphaeifolia St<strong><strong>an</strong>d</strong>ley, Proc.<br />
Biol. Soc. Wash. 37: 50. 1924; Burger, Field-<br />
i<strong>an</strong>a Bot. 40: 134, t. 22. 1977. Type. Costa<br />
Rica. Alajuela: Road to S<strong>an</strong> Carlos valley,<br />
Buena Vista, 16 Apr 1903 (2), Cook & Doyle<br />
157 (holotype, US). Fig. 40.<br />
Tree, hemi-epiphytic or terrestrial, ca. 20 m<br />
tall. Leafy twigs 5-17 mm thick, densely mi-<br />
nutely puberulous or also <strong>with</strong> dense reddish-<br />
brown pluricellular hairs. Lamina coriaceous,<br />
(broadly) elliptic to ovate, (9-)24-29 x (6-)18-<br />
Specimens studied. COSTA RICA. ALAJUELA: Road<br />
to S<strong>an</strong> Carlos Valley, Buena Vista, 16 Apr 1903, (2),<br />
Cook & Doyle 157 (US, type collection); c<strong>an</strong>ton S<strong>an</strong><br />
Carlos, Villa Quesada, 21 Feb 1939 (8), A. Smith 1632<br />
(F, MO). HEREDIA: Nr. Puerto Viejo de Sarapiqui, 3<br />
Apr 1974 (2), Hartshorn 1436 (U).<br />
77<br />
Although C. nymphaeifolia is reminiscent of<br />
C. villosa in general appear<strong>an</strong>ce <strong><strong>an</strong>d</strong> also to C.<br />
duquei in the nature of the pistillate inflores-<br />
cence, these species are probably unrelated. The<br />
occurrence of 3-staminate flowers <strong><strong>an</strong>d</strong> other sim-<br />
ilarities in leaf characters <strong><strong>an</strong>d</strong> the staminate in-<br />
florescence suggest a relationship <strong>with</strong> C. lon-<br />
gepedunculata.<br />
30. <strong>Coussapoa</strong> oligocephala Donnell Smith, Bot.<br />
Gaz. 40: 11. 1905. Type. Guatemala. Alta Ve-<br />
rapaz: Cubilguitz, Apr 1904 (6), Von Tuerk-<br />
heim 8659 (=11.942) (holotype, US; isotype,<br />
B). Fig. 41.
78 Flora Neotropica<br />
?~ ~~~~~~~~~~~~~~~~~<br />
.?'<br />
?<br />
:~~~~~~~~~~~~'-<br />
- ,??? I I 'I'~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~<br />
'~~~~~~~~?:<br />
.<br />
P~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~<br />
.I . ,. "<br />
r.~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~<br />
?I ?::~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~<br />
. .<br />
' ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~~~~ "<br />
~~~~~~ ''~ ~ ~ ~~~~~~~~~~~~~<br />
.? ?~~~~~~~~~~~~<br />
~. c , ,,.<br />
? ' 'y<br />
.<br />
~~~~~~~~~~~. .<br />
F~~~~~~~~~~~~~~~~~~~~...:<br />
?~~~~~~~~~~~~~~~~~~~~~~~~~~~ :<br />
?':.<br />
I~~~~~~~~~~~~~~~~~~~~~<br />
-L<br />
..T!<br />
?. jI .?~<br />
~~~~~~~~~ .....?:;:"li?b<br />
i"<br />
~~~~~~~~~~~<br />
.. .... ';: -:F: ..~,?;<br />
~~~~~~~~~~: ~ ?? .. . *<br />
"'<br />
~~;.? .~~~~~~~~~~~~~~'<br />
'~~~~~~~~~~~~~~ ? ;?r<br />
~~~~~~~~~~~~~~~~~~~~~~~~~~~'-?.-<br />
I.':!<br />
?-:'<br />
,::<br />
, i'<br />
i; , : , ...<br />
?~~~~~~~~~~~~~~~~~~~~~~1<br />
"'<br />
"<br />
FIG. 40. <strong>Coussapoa</strong> nymphaeifolia: 1, leafy twig <strong>with</strong> staminate inflorescences, 2, leaf (A. Smith 1632).<br />
?:".<br />
'. .:i.- ; .
<strong>Coussapoa</strong><br />
I<br />
1 cm<br />
FIG. 41. <strong>Coussapoa</strong> oligocephala: 1, leafy twig <strong>with</strong> staminate inflorescences (Schipp 999); 2, pistillate<br />
inflorescences<br />
(Contreras 5787); 3, leaf (Contreras 5787).<br />
79
80<br />
Specimens studied. MEXICO. TABASCO: La Palma,<br />
Bal<strong>an</strong>c<strong>an</strong>, 6 Jun 1939 (6), Matuda 3300 (A, F, K, NA,<br />
NY).<br />
BELIZE. EL CAYO DISTRICT: Nr. El Cayo, 1933 (d),<br />
Flora Neotropica<br />
Shrub or tree, hemi-epiphytic or GUATEMALA. ALTA VERAPAZ:<br />
terrestrial, up<br />
Sebol, 21 Apr 1964<br />
(8), Contreras 4449<br />
to 20 m tall. Leafy twigs 4-7 mm<br />
(NY, S); El Cacao, Trece Aguas,<br />
thick, ap- - (9), Cooks.n. (F, US); Cerro Chinaja, between Finca<br />
pressed-puberulous to hirtellous to strigillose or Yalpemech <strong><strong>an</strong>d</strong> Chinnaja, 1-2 Apr 1942 (st), Steyerto<br />
hirsute. Lamina (sub)coriaceous, elliptic to mark 45663 (F); Cubilgultz, Apr 1904 (6), Von Tuerckoblong<br />
to (sub)ovate to (sub)obovate, 4-21 x 1- heim 8659 (=11.942) (B, US, type collection). IZABAL:<br />
7 cm, apex shortly acuminate to acute to<br />
Rio<br />
obtuse,<br />
Dulce, road Seja-Cienaga, km 5, 11 Jul 1970 (2),<br />
Contreras 10195 (BM, DUKE, MO, U); Rio Dulce, 21<br />
base obtuse to subcordate, margin entire; upper Jul 1936 (2), Hatch et al. s.n. (F); lower Rio Oscuro,<br />
surface glabrous, lower surface minutely puber- SW of Lake Izabal, 27 Apr 1966 (9), G. C. Jones et al.<br />
ulous to glabrous in the areoles sparsely hirtel- 3152 (NY, US); Lake Izabal, Jagua Creek, 31 May 1965<br />
lous on the smaller veins, the main veins gla- (9), Snedaker C39 (F), 28 May 1966 (9), Snedaker D88<br />
brous, the whole surface <strong>with</strong> (persistent) white (A, F); Rio Dulce, between Livingstone <strong><strong>an</strong>d</strong> 6 mi up<br />
river, 14 Apr 1940 (8), Steyermark 39460 (F); Rio Frio,<br />
arachnoid hairs; lateral veins (5-)8-12, straight, 17 Dec 1941 (st), Steyermark 39934 (F, US); Rio Dulce,<br />
basal pair unbr<strong>an</strong>ched, reaching the margin be- 21 May 1939 (8), Wilson 381 (F). PETEN: Vaxactun,<br />
low the middle of the lamina; intercostal vena- 30 May 1931 (2), Bartlett 12356 (GH, NY, S, US);<br />
tion prominent; petiole (1-)2-6 cm Tikal, 31 Jul 1959 (2), Contreras 61 (NY,<br />
long, sparsely<br />
S); Lake Itza,<br />
between Remate <strong><strong>an</strong>d</strong> S<strong>an</strong> Andr6s, 18 Apr 1960 (9),<br />
to densely minute puberulous to hirtellous, or Contreras 845 (F, NY, S, US); old road to Machaquila,<br />
hairs mixed <strong>with</strong> longer patent to appressed, <strong><strong>an</strong>d</strong> Dolores, 27 Apr 1961 (8), Contreras 2220 (NY, S); nr.<br />
sometimes also arachnoid hairs; stipules 1-5 cm S<strong>an</strong> Andr6s, Apr 1962 (2), Contreras 3559 (F, NY, S);<br />
long, (rather) sparsely appressed-puberulous (to Mac<strong>an</strong>che, 18 May 1966 (9), Contreras 5787 (GH, US);<br />
Cadenas<br />
brownish subsericeous). Staminate road, km 138, La Cumbre, 25 Sep 1966 (9),<br />
inflores- Contreras 6231 (BM, MO, U); Lake Yaxha, archeocences<br />
br<strong>an</strong>ched; heads 3-10, globose, ca. 4-6 logical camp on north shore, 18 Jun 1973 (st), Croat<br />
diam.; common peduncle 1-5 cm long, <strong>with</strong> pu- 24663 (MO, NY); nr. Carmelita, 25 Jun 1942 (st), F.<br />
berulous to hirtellous to subtomentellous; peri- E. Egler 42-229 (F); 5 mi S of Tikal, 19 Jun 1973 (9),<br />
<strong>an</strong>th ca. 1 mm high, minutely puberulous; sta- Gentry 8348 (U); nr. S<strong>an</strong> Andr6s, 2 May 1933 (9),<br />
Lundell 3170<br />
mens three, far<br />
(F); La Libertad, 31<br />
exceeding the peri<strong>an</strong>th. Pistillate<br />
May 1933 (9), Lundell<br />
3535 (F, S); Tikal, 30 Apr 1959 (6), Lundell 15930<br />
inflorescences unbr<strong>an</strong>ched or occasionally poorly (S), 2 Jul 1959 (8), Lundell 16122 (F, S), 4 Jul 1959<br />
br<strong>an</strong>ched; heads 1(-3), globose, ca. 5-8 mm, in (9), Lundell 16162 (G, NY, S, US), 27 Jul 1969 (6),<br />
fruit up to 10 mm diam.; (common) peduncle 1- Ortiz 200 (BM, F); S<strong>an</strong> Benito, nr. Playa Pl<strong>an</strong>ca, 26<br />
5 cm long, ca. 1 mm thick, puberulous to hir- Apr 1970 (8), Ortiz 1030 (BM, F, MO, NY); Tikal, 29<br />
May 1971 (9), Ortiz 1810 (BM, F, US); Cerro Ceibal,<br />
tellous to subtomentellous; peri<strong>an</strong>th ca. 1 mm between Rio S<strong>an</strong>ta M6nica <strong><strong>an</strong>d</strong> mouth of Rio Martin,<br />
high, minutely puberulous; peri<strong>an</strong>th or<strong>an</strong>ge or 30 Apr 1942 (st), Steyermark 46103 (F); Rio Machayellow.<br />
Interfloral bracts (sub)spathulate to sub- quila, N of El Cambio, 25 Apr 1942 (6), Steyermark<br />
peltate, minutely puberulous at the 45981<br />
apex.<br />
(F).<br />
Distribution (Fig. 8). Northern Central Amer- <strong>Coussapoa</strong> oligocephala is strongly reminisica<br />
(Guatemala <strong><strong>an</strong>d</strong> Belize) to southern Mexico cent of the Amazoni<strong>an</strong> C. sprucei, especially when<br />
(Tabasco); in lowl<strong><strong>an</strong>d</strong> (often riverine) forest. the specimens have <strong>an</strong> elliptic to subovate lamina<br />
<strong>with</strong> subcordate base.<br />
31. <strong>Coussapoa</strong> orthoneura St<strong><strong>an</strong>d</strong>ley, Publ. Field<br />
Mus. Bot. 17: 165. 1937.<br />
Ch<strong>an</strong>ek 220<br />
Type. Brazil. Ama-<br />
(F, K); between Millionaro <strong><strong>an</strong>d</strong> Cueras,<br />
30 May 1973 (6), Dwyer 10799 (U); Vaca, 12<br />
zonas: Mun. Sao Paulo de<br />
May<br />
OlivenSa, nr. Pal-<br />
1938 (6), Gentle 2602 (A, F, K, MO); nr. Millionaro, mares, 26 Oct-11 Dec 1936 (2), Krukoff8518<br />
30 May 1973 (6), Gentry 7727 (MO, NY); Valentin, (holotype, NY; isotypes, A, B, BM, F, G, K,<br />
Jun-Jul 1936 (6), Lundell 6224 (F, GH, NY, S, US), LE, MO, SU, U, US). Fig. 42.<br />
Jun-Jul 1936 (2), Lundell 6350 (C, F, GH, MO, NY,<br />
S, US); Macaw B<strong>an</strong>k, 3 Apr 1969 (9), Proctor 30273 <strong>Coussapoa</strong> williamsii Cuatrecasas, Fieldi<strong>an</strong>a Bot. 28(1):<br />
(BM, MO). TOLEDO DISTRICT: Bolo Camp, 13 Apr 212. 1951. Type. Venezuela. Terr. Fed. Amazonas:<br />
1944 (6), Gentle 4528 (DUKE); Honey Camp-Or<strong>an</strong>ge Isla Solitaria, between Tamatama <strong><strong>an</strong>d</strong> Esmeralda, 7<br />
Walk, Sep 1928 (st), Lundell 2 (BM, F, K), 24 Oct 1929 May 1942 (6), LI. Williams 15240 (holotype, F; iso-<br />
(6), Lundell 648 (F, K, MO, NY, S, US); Toledo, 1906- types, A, G, NY, RB, US, VEN).<br />
1907 (6), Peck 497 (GH, K, NY, U); Punta Gorda, 7 <strong>Coussapoa</strong> pr<strong>an</strong>cei C. C. Berg, Acta Bot. Neerl. 27(1):<br />
Aug 1932 (6), Schipp 999 (A, BM, F, G, GH, K, MO, 11. 1978. Type. Brazil. Amazonas: Rio Cuieras, nr.<br />
NY, S).<br />
Jarada, 17 Sep 1973 (9), Pr<strong>an</strong>ce et al. 18032 (holo-
<strong>Coussapoa</strong><br />
FIG. 42. <strong>Coussapoa</strong> orthoneura: 1, leafy twig <strong>with</strong> pistillate inflorescences (Pr<strong>an</strong>ce 14092); 2, leafy twig <strong>with</strong><br />
staminate inflorescences (Schultes et al. 13774).<br />
type, INPA; isotypes, C, F, K, M, MICH, MO, NY,<br />
P, SU, U, US).<br />
Tree, hemi-epiphytic or terrestrial, up to 35 m<br />
tall. Leafy twigs 2-6 mm thick, glabrous or<br />
sparsely, puberulous, sometimes hirtellous.<br />
Lamina (sub)coriaceous, obovate to (broadly) elliptic<br />
to oblong, 2-15(-24) x 1-8(-11) cm, apex<br />
1 cm<br />
81<br />
acute to shortly acuminate or obtuse, base obtuse<br />
to acute or truncate, margin entire or subcrenate;<br />
upper surface glabrous, lower surface glabrous or<br />
sparsely to densely appressed-puberulous, sometimes<br />
substrigose or <strong>with</strong> white arachnoid hairs<br />
later deciduous; lateral veins 2-7 pairs, straight<br />
or slightly curved, basal pairs unbr<strong>an</strong>ched reach-
82 Flora Neotropica<br />
ing the margin above to just below the middle PERU. AMAZONAS: Prov. Bagua, Dist. Cenepa,<br />
of the lamina; intercostal venation (almost) pl<strong>an</strong>e;<br />
Nueva Nazareth, nr. mouth of Rio Imaza, on Rio Mapetiole<br />
1-7 cm long, glabrous or raiion, 26 J<strong>an</strong>-18 Feb 1967 (2), Tillett 671-33 (YEN).<br />
appressed-pu- LORETO: Prov. Maynas, Rio N<strong>an</strong>ay, nr. Moron Cocha,<br />
berulous to strigillose or to hirtellous; stipules N of Iquitos, 21 Mar 1977 (6), Gentry et al. 18528<br />
(0.5-)1-7 cm long, sparsely to densely appressed- (MO, U), Rio N<strong>an</strong>ay, nr. Caserio S<strong>an</strong>ta Clara, nr. Iquipuberulous<br />
or to mostly sparsely yellowish sub- tos, 19 Nov 1976 (2), Revilla 1847 (U).<br />
sericeous (to subhirsute or to subvillous), occa- BRAZIL. AMAZONAS: M<strong>an</strong>aus, Igarap6 do Leao, 3<br />
sionally also <strong>with</strong> sparse to rather dense<br />
Sep 1957 (6), L. Coelho<br />
white<br />
(INPA) 5728 (U); Nogueira-<br />
Tef'e, 26 Oct 1972 (2), D<strong>an</strong>ta 12383 (INPA); Ton<strong>an</strong>arachnoid<br />
hairs. Staminate inflorescences tins, 7 Feb 1944 (9), Ducke 1528 (NY, US); Rio Purus,<br />
br<strong>an</strong>ched; heads m<strong>an</strong>y, sometimes partly fused, Quinta de Sao Christovao, 18 J<strong>an</strong> 1873 (6), Glaziou<br />
globose, ca. 1-6 mm diam.; common peduncle 10070a (P); Mun. Sao Paulo de Olivenca, nr. Palmares,<br />
1-3 cm long, <strong>with</strong><br />
11<br />
(dense) appressed-puberulous,<br />
Sep-26 Oct 1936 (2), Krukoff8518 (A, BM, F, G,<br />
K, LE, NY, S, U, US,<br />
sometimes<br />
type collection), (6), Krukoff8587<br />
also <strong>with</strong> reddish-brown pluricellular (F); Mun. Sao Paulo de Olivenca, Belem Creek, 26<br />
hairs; peri<strong>an</strong>th ca. 1 mm high, glabrous or min- Oct-11 Dec 1936 (6), Krukoff8967 (A, B, BM, F, G,<br />
utely puberulous; stamen one, just exceeding K, LE, MO, NY, S, U); Lago do Tefe, 10 km above<br />
the peri<strong>an</strong>th. Pistillate inflorescences br<strong>an</strong>ched;<br />
Tefe, 26 Jul 1971 (8), McD<strong>an</strong>iel et al. 2666 (US); Rio<br />
heads 2-10, sometimes partly fused, 3-6 mm, in Ituxi, nr. B6ca do Curuquete, 9 Jul 1971 ((), Pr<strong>an</strong>ce et<br />
al. 14029 (F, GH, K, M, S, U); Rio Cuieras, just below<br />
fruit up to 15 mm diam.; common peduncle 1- mouth of Rio Br<strong>an</strong>chino, 29 Sep 1971 (6), Pr<strong>an</strong>ce et<br />
3 cm long, (densely) appressed-puberulous, al. 15039 (F, GH, K, M, P, S, U); Rio Cuieras, nr.<br />
sometimes partly hirtellous, apices of the head- Jarada, 17 Sep 1973 (2), Pr<strong>an</strong>ce et al. 18032 (C, F, K,<br />
bearing br<strong>an</strong>ches often (especially in fruit) more M, MICH, MO, NY, P, S, U, US, type collection of<br />
C.<br />
or less<br />
pr<strong>an</strong>cei); road M<strong>an</strong>aus-Itacoatiara, km 29, 14 Sep<br />
swollen; peri<strong>an</strong>th ca. 1-2 mm high, gla- 1974 (2), Pr<strong>an</strong>ce et al. 22729 (INPA, U); Mun. Purabrous<br />
or sparsely papillate at the apex; fruiting quequara, road to M<strong>an</strong>aus, P<strong>an</strong>ama, 6 Jun 1963 (2),<br />
peri<strong>an</strong>th yellow to red. Interfloral bracts absent. W. A. Rodriques et al. 5266 (U); Rio Cast<strong>an</strong>ho, road<br />
Distribution (Fig. 8). Upper Amazon Basin to Careiro, 11 Jul 1972 (8), M. Silva et al. 466 (U); Rio<br />
(Brazil, Peru, Ecuador, <strong><strong>an</strong>d</strong> Venezuela); in Uaup6s, Taracua, Rio Tikie, 28 J<strong>an</strong>-9 Feb 1948 (6),<br />
up- Schultes et al. 9685 (GH, K); Rio Uaup6s, P<strong>an</strong>ur6, Oct<br />
l<strong><strong>an</strong>d</strong> <strong><strong>an</strong>d</strong> riverine (varzea) forest; also known from 1852-J<strong>an</strong> 1853 (8), Spruce 2498 (BM, G, GH, K, LE,<br />
Central Columbia (S<strong>an</strong>t<strong><strong>an</strong>d</strong>er, at ca. 1000 m). NY, OXF, P).<br />
Specimens studied. COLOMBIA. AMAZONAS: Rio<br />
Igara-Par<strong>an</strong>a, affluent of Rio Putumayo, La Chorrera,<br />
19 Dec 1973 (9), Gasche et al. 29 (U). GUAINiA: Rio<br />
Guainia, M<strong>an</strong>acal, 21 Oct 1977 (2), Espina et al. 219<br />
(COL). SANTANDER: Rio Negro, El Play6n, 1000 m, -<br />
<strong>Coussapoa</strong> orthoneura is a variable species,<br />
especially in the shape, dimensions, <strong><strong>an</strong>d</strong> secondary<br />
venation of the lamina <strong><strong>an</strong>d</strong> also in the indumentum<br />
of the stipules. More or less distinct<br />
(st), Fr<strong>an</strong>co & Garces 419 (COL). VAUPES: Upper Rio regional forms, like the small-leaved form found<br />
Vaup6s, La Jirisa, 9 J<strong>an</strong> 1944 (a), Guttierrez et al. 562 near M<strong>an</strong>aus, at the time described as C. pr<strong>an</strong>cei,<br />
(COL, GH); upper Rio Vaup6s, between La Jarisa <strong><strong>an</strong>d</strong> have<br />
Las Bocas, 12 J<strong>an</strong> 1944 (Q), Guttierrez et al. 595<br />
proved not to be sufficiently different to be<br />
(COL);<br />
Rio Apaporis, nr. Cachivera de Jirijirimo, 8 Jul 1951 recognized as distinct (infraspecific) taxa. Cous-<br />
(Y), Schultes et al. 12985 (F); Rio Apaporis, between sapoa arachnoidea appears to be closely related<br />
Rio Pacoa <strong><strong>an</strong>d</strong> Rio K<strong>an</strong><strong>an</strong>ari, 27 Aug 1951 (6), Schultes to C. orthoneura, which shows affinities espeet<br />
al. 13774 (COL, GH, U); Rio Apaporis, Raudal de<br />
Jirijirimo, below<br />
cially to C. cupularis <strong><strong>an</strong>d</strong> C. nitida. In the<br />
mouth of Rio K<strong>an</strong><strong>an</strong>ari, 14<br />
veg-<br />
Feb 1952<br />
(9), Schultes et al. 15324<br />
etative<br />
(GH,<br />
parts the<br />
US).<br />
species c<strong>an</strong> easily be confused<br />
VENEZUELA. AMAZONAS: Rio Casiquiare, be- <strong>with</strong> C. latifolia, C. viridifolia <strong><strong>an</strong>d</strong> C. microcephtween<br />
Lago de Vasiva <strong><strong>an</strong>d</strong> Rio Pacimoni, 1853-1854 ala, but it is quite distinct from these three species<br />
(a), Spruce 3231 (B, C, G, K, P, S); Rio Ocama, above in the ebracteate inflorescence <strong><strong>an</strong>d</strong> the staminate<br />
confluence <strong>with</strong> Rio Orinoco, nr. Ocama Mission, 23- flower.<br />
24 May 1972 (8), Steyermark 106167 (U, VEN); Isla<br />
Solitaria, between Tamatama <strong><strong>an</strong>d</strong> Esmeralda, 7 May<br />
1942 (6), Ll. Williams 15240 (A, F, G, type collection 32. <strong>Coussapoa</strong> ovalifolia Trecul, Ann. Sci. Nat.<br />
of C. williamsii); Esmeralda, 14 May 1942 (a), LI. Wil- Bot. S6r. 3, 8: 95.<br />
liams<br />
1847;<br />
15363<br />
Miquel, Fl. bras. 4(1):<br />
(G, NY, US, VEN).<br />
139.<br />
ECUADOR. NAPO: Misahualli, mouth of Rio Misa-<br />
1853; Macbride, Publ. Field Mus. Bot.<br />
hualli, 3 Mar 1980 (st), Berg & Akkerm<strong>an</strong>s 1111 (U); 13(2.2): 298. 1937. Type. Peru. Hu<strong>an</strong>uco: Maroad<br />
Coca-Lago Agrio, 9 km NE of Rio Coca, 20 Mar cora ?,<br />
1980 (st), Br<strong><strong>an</strong>d</strong>byge et al. 30258 (AAU).<br />
- (2), Ruiz & Pavon s.n. (or 2) (holotype,<br />
FI; isotypes, F, US). Fig. 43.
<strong>Coussapoa</strong><br />
FIG. 43. <strong>Coussapoa</strong> ovalifolia: 1, leafy twig <strong>with</strong> pistillate inflorescences<br />
(Krukoff5657); 2, staminate inflo-<br />
rescences (Ule 9316).<br />
83
84 Flora Neotropica<br />
<strong>Coussapoa</strong> hirsuta Tr6cul, Ann. Sci. Nat. Bot. Ser. 3, & Akkerm<strong>an</strong>s 1132 (U). ZAMORA-CHINCHIPE: Road<br />
8: 97. 1847; Macbride, Publ. Field Mus. Bot. 13(2.2): to Guayzimi, 27 km NNE of Zamora, 17 Sep 1975<br />
297. 1937. Type. Peru. Hu<strong>an</strong>uco: Macora, - (9), (st), Little et al. 424 (COL, Q).<br />
Ruiz & Pavon s.n. (or 5) (holotype, FI; isotype, B). PERU. AMAZONAS: E ofHuampmai, 4 Jul 1974 (2),<br />
<strong>Coussapoa</strong> acutifolia Klotzsch, Linnaea 20: 529. 1847; Berlin 1531 (MO, U). HuANuco: Prov. Leoncio Prado,<br />
Macbride, Publ. Field Mus. Bot. 13(2.2): 296. 1937. Tingo Maria, 7 Dec 1981 (9), Plowm<strong>an</strong> et al. 11179 (U);<br />
Type. Peru. Hu<strong>an</strong>uco: Cochero (fide Macbride, 1937), Chinchoa-Pillao-Pozuzo-Mucuna, - (2), Ruiz & Pa-<br />
- (9), Ruiz & Pavon s. n. (or 7) (holotype, B; isotype, von s.n. (or 6), (B, F, lectotype collection of C. puber-<br />
FI).<br />
ula); Cochera, - (2), Ruiz & Pavon s.n. (or 7) (B, F,<br />
<strong>Coussapoa</strong> puberula Klotzsch, Linnaea 20: 529. 1847. FI, US, type collection of C. acutifolia); Macora,<br />
Type. Peru. Huinuco: "Chinchao-Pillao-Pozuzo-<br />
Mucona," - (9), Ruiz & Pavon s.n. (or 6) (holotype,<br />
B; isotype, F).<br />
<strong>Coussapoa</strong> setosa Klotzsch, Linnaea 20: 528. 1847.<br />
Type. Peru. Huinuco: Macora, - (9), Ruiz & Pavon<br />
s.n. (or 5) (holotype, B; isotype, FI).<br />
- (2),<br />
Ruiz & Pavon s.n. (or 2) (F, FI, US, type collection of<br />
C. ovalifolia), - (2), Ruiz & Pavon s.n. (or 7) (B, FI,<br />
type collection of C. hirsuta <strong><strong>an</strong>d</strong> C. setosa); Prov. Pachitea,<br />
Dtto. Honoria, Bosque Nac. Iparia, Schunke V.<br />
1162 (F, G, NY, US). JUNIN: Valley of Rio Paucartambo,<br />
nr. Perene bridge, 19 Jun 1929 (a), Killip et al.<br />
25341 (F, NY, S, US); P<strong>an</strong>chis trail, between S<strong>an</strong> Nicolas<br />
<strong><strong>an</strong>d</strong> Azupizfi, 6 Jul 1929 (2), Killip et al. 26095<br />
(NY); La Merced, 10-24 Aug 1928 (a), Macbride 5594<br />
(F). LORETO: Prov. Maynas, Rio N<strong>an</strong>ay, Mish<strong>an</strong>a, halfway<br />
between Iquitos <strong><strong>an</strong>d</strong> S<strong>an</strong>ta Maria de N<strong>an</strong>ay, 26<br />
Jul 1979 (st), Gentry 25151 (MO, U); Rio Marafion,<br />
Y<strong>an</strong>ache, 2 Mar 1924 (9), Kuhlm<strong>an</strong>n 1540 (U); Quebrada<br />
Aucaya, 12 May 1973 (9), Rimachi 359 (NA);<br />
Rio Mar<strong>an</strong>ion, between Iquitos <strong><strong>an</strong>d</strong> Pongo de M<strong>an</strong>seriche,<br />
1924 (9), Tessm<strong>an</strong>n 3922 (G); Matari Cocha, 3<br />
Oct 1968 (d), Torres M. 482 (K). MADRE DE DioS:<br />
Parque Nacional del M<strong>an</strong>u, Cocha Cashu, between<br />
Punagua <strong><strong>an</strong>d</strong> Tayakome, 17-24 Aug 1974 (2), Foster<br />
et al. 3412 (F). SAN MARTIN: Eslab6n, between Saposoa<br />
<strong><strong>an</strong>d</strong> Tinge de Saposoa, Rio Saposoa, 8 Sep 1948<br />
(9), Ferreyra 4813 (US); between Gramalote <strong><strong>an</strong>d</strong> Saposoa,<br />
29 Apr 1962 (a), Woytkowski 7306 (GH, US).<br />
BRAZIL. ACRE: Mouth of Rio Macaua, tributary of<br />
Rio laco, 26 Aug 1933 (9), Krukoff 5657 (A, BM, F,<br />
G, K, LE, M, S, U); nr. Tarauaca, 25 Sep 1970 (2),<br />
Pr<strong>an</strong>ce et al. 7371 (U); Rio Acre, Seringal Sao Fr<strong>an</strong>cisco,<br />
Aug 1911 (9), Ule 9315 (G, K, L, MG), May<br />
1911 (a), Ule 9316 (G, K, L, MG, US).<br />
BOLIVIA. SANTA CRUZ: Buena Vista, Jun 1915 (8),<br />
Steinbach 1484 (GH, U).<br />
Tree, hemi-epiphytic, up to 20 m tall. Leafy<br />
twigs 3-9 mm thick, glabrous or puberulous, often<br />
also <strong>with</strong> distinctly longer straight stiff hairs.<br />
Lamina coriaceous, ovate to subovate (or elliptic<br />
to obovate), 7-32 x 2-16 cm, apex shortly acu-<br />
minate to acute (to obtuse), base obtuse to round-<br />
ed or acute to truncate, margin entire or sub-<br />
crenate; upper surface glabrous, lower surface<br />
glabrous or <strong>with</strong> the main veins having appressed<br />
straight hairs of different lengths, often initially<br />
also <strong>with</strong> sparse or sometimes rather dense white<br />
arachnoid hairs on the main veins <strong><strong>an</strong>d</strong> along the<br />
margin; lateral veins 7-21 pairs, almost straight<br />
to curved, basal pair unbr<strong>an</strong>ched or in large leaves<br />
poorly br<strong>an</strong>ched, reaching the margin far below<br />
the middle of the lamina; intercostal venation<br />
almost pl<strong>an</strong>e to slightly prominent; petiole 2-9<br />
cm long, glabrous or <strong>with</strong> appressed hairs of dif-<br />
ferent lengths; stipules 1-4 (in rapidly growing<br />
shoots up to 13) cm long, subsericeous (to sub-<br />
hirsute). Staminate inflorescences repeatedly<br />
br<strong>an</strong>ched; heads numerous, globose, ca. 1(-2) mm<br />
diam.; common peduncle 1.5-3 cm long puberu-<br />
lous to hirtellous, occasionally also <strong>with</strong> sparse<br />
arachnoid hairs; peri<strong>an</strong>th 0.5-1 mm high, gla-<br />
brous; stamen one, just exceeding the peri<strong>an</strong>th.<br />
Pistillate inflorescences br<strong>an</strong>ched or sometimes<br />
unbr<strong>an</strong>ched; heads 1-5, subglobose, 5-8 mm, in<br />
fruit up to 15 mm diam.; (common) peduncle 2-<br />
6 cm long, puberulous to hirtellous; peri<strong>an</strong>th ca.<br />
1-2 mm high, glabrous; fruiting peri<strong>an</strong>th brown-<br />
ish. Interfloral bracts absent.<br />
Distribution (Fig. 8). Upper Amazon Basin<br />
(Bolivia, Peru, Brazil <strong><strong>an</strong>d</strong> Ecuador); in upl<strong><strong>an</strong>d</strong><br />
<strong><strong>an</strong>d</strong> riverine forest, up to 900 m.<br />
Specimens studied. ECUADOR. NAPO: Lago Agrio,<br />
3 Apr 1980 (9), Br<strong><strong>an</strong>d</strong>byge et al. 30421 (AAU). PASTA-<br />
ZA: Between Palora <strong><strong>an</strong>d</strong> Shell, 4 Mar 1980 (st), Berg<br />
<strong>Coussapoa</strong> ovalifolia is closely related to C.<br />
napoensis. The collection Gentry & Mori 14011,<br />
P<strong>an</strong>ama, Cerro Tacaruna, premont<strong>an</strong>e wet for-<br />
est, 1300 m, 30 J<strong>an</strong> 1975 (MO, U), probably<br />
represents a separate taxon, possibly distinct at<br />
the subspecific level from the Amazoni<strong>an</strong> rep-<br />
resentatives of C. ovalifolia. The collection is<br />
similar in most characters to the other collections<br />
of C. ovalifolia, but is distinct mainly in the short<br />
(1-1.5 cm long) peduncle of the pistillate inflo-<br />
rescence <strong><strong>an</strong>d</strong>the br<strong>an</strong>ching of the basal lateral<br />
veins-features which hardly justify recognition<br />
as a distinct species. Staminate material is need-<br />
ed to confirm the present supposition.<br />
33. <strong>Coussapoa</strong> pachyphylla Akkerm<strong>an</strong>s & C. C.<br />
Berg, Proc. Kon. Ned. Akad. Wetensch. Ser.<br />
C, 85(4): 459. 1982. Type. Brazil. Bahia: Mun.
<strong>Coussapoa</strong> 85<br />
lcm.<br />
FIG. 44. <strong>Coussapoa</strong> pachyphylla: 1, leafy twig <strong>with</strong> pistillate inflorescences<br />
(Mori & S<strong>an</strong>tos 11655).<br />
S<strong>an</strong>ta Cruz de Cabralia, old road to S<strong>an</strong>ta Cruz,<br />
ca. 15 km NW of P6rto Seguro, 3 Apr 1979<br />
(9), Mori & dos S<strong>an</strong>tos 11655 (holotype, CE-<br />
PEC; isotype, U). Fig. 44.<br />
Shrub, often (hemi-)epiphytic, up to 5 m tall<br />
(or taller?). Leafy twigs 4-8 mm thick, rather<br />
densely to sparsely appressed-puberulous, also<br />
<strong>with</strong> distinctly longer straight stiff hairs <strong><strong>an</strong>d</strong> brown<br />
pluricellular hairs, glabrescent. Lamina coriaceous,<br />
ovate to elliptic (or obovate), 6-16(-24)<br />
x 3.5-10(-19), apex acute to subacuminate, base<br />
acute, obtuse or sometimes subcordate, margin<br />
entire to subcrenate; upper surface glabrous, lower<br />
surface densely to sparsely puberulous to subtomentose,<br />
initially often <strong>with</strong> sparse brownish<br />
arachnoid hairs, especially along the margin; lateral<br />
veins 5-8 pairs, basal pair br<strong>an</strong>ched or sometimes<br />
(in small leaves) unbr<strong>an</strong>ched, reaching the<br />
margin below the middle of the lamina; inter-<br />
costal venation more or less prominent; petiole<br />
0.5-3(-5) cm long, densely to sparsely puberu-<br />
lous, often also <strong>with</strong> distinctly longer straight stiff<br />
hairs; stipules 1-2 cm long, densely covered <strong>with</strong><br />
short reddish-brown arachnoid hairs. Staminate<br />
inflorescences br<strong>an</strong>ched; heads ca. 6-12, some-<br />
times fused, globose, ca. 1-2 mm diam.; com-<br />
mon peduncle 1-2 cm long, densely puberulous;<br />
peri<strong>an</strong>th ca. 0.5 mm high, nearly glabrous; sta-<br />
mens two, as long as the peri<strong>an</strong>th. Pistillate in-<br />
florescences unbr<strong>an</strong>ched or few-br<strong>an</strong>ched; heads<br />
1-3, sometimes partly fused, ca. 5-8 mm, in fruit<br />
up to 13 mm diam.; (common) peduncle 2.5-4<br />
cm long, 1-1.5 mm thick, densely puberulous;<br />
peri<strong>an</strong>th 1(-2) mm high, glabrous. Interfloral<br />
bracts, few, subpeltate to spathulate.<br />
Distribution (Fig. 8). Brazil (Bahia); in humid<br />
forests in the coastal region.
86<br />
Specimens studied. BRAZIL. BAHIA: Road Una-Ilheus,<br />
ca. km 21, 2 Apr 1980 (st), Berg et al. 1153<br />
(CEPEC, K, NY, RB, U); Mun. S<strong>an</strong>ta Cruz de Cabralia,<br />
old road to S<strong>an</strong>ta Cruz, ca. 15 km NW of Porto Seguro,<br />
3 Apr 1979 (Q), Mori et al. 11655 (CEPEC, U, type<br />
collection); Ilh6us-Repartimento (Fortuna), 21 Nov<br />
1944 (8), Vellozo 737 (R).<br />
Tree, hemi-epiphytic or terrestrial, up to 30 m<br />
tall. Leafy twigs 4-8 mm thick, glabrous or pu-<br />
berulous. Lamina coriaceous, broadly ovate to<br />
elliptic or suborbicular, sometimes tending to<br />
obovate, 7-24 x 4-18 cm, apex shortly acumi-<br />
nate, sometimes almost rounded or obtuse, base<br />
rounded to truncate or subcordate, margin en-<br />
tire; upper surface glabrous, lower surface gla-<br />
brous, occasionally sparsely puberulous to to-<br />
mentellous; lateral veins 3-6 pairs, slightly<br />
curved, basal pair br<strong>an</strong>ched, reaching the margin<br />
at or above the middle of the lamina; intercostal<br />
venation pl<strong>an</strong>e or slightly prominent; petiole<br />
(0.5-)2-7 cm long, occasionally sparsely puber-<br />
ulous; stipules 1.5-5 cm long, glabrous or sparse-<br />
ly puberulous. Staminate inflorescences br<strong>an</strong>ched;<br />
heads m<strong>an</strong>y, globose, ca. 1 mm diam.; common<br />
peduncle 1-5 cm long, glabrous or sparsely pu-<br />
berulous; peri<strong>an</strong>th ca. 0.5 mm high, glabrous;<br />
stamens three, just exceeding the peri<strong>an</strong>th. Pis-<br />
tillate inflorescences br<strong>an</strong>ched; heads 3-12, glo-<br />
bose, ca. 2-3 mm, in fruit up to 6 mm diam.;<br />
common peduncle 1-4 cm long, glabrous or<br />
sparsely puberulous; flowers connate, peri<strong>an</strong>th<br />
ca. 1 mm high, glabrous. Interfloral bracts lack-<br />
ing or in the staminate heads a few, more or less<br />
distinctly spathulate (more or less cucullate), gla-<br />
brous or ciliolate bracts.<br />
Flora Neotropica<br />
Distribution (Fig. 8). Costa Rica, P<strong>an</strong>ama, <strong><strong>an</strong>d</strong><br />
in the Pacific coastal region of Colombia <strong><strong>an</strong>d</strong><br />
northern Ecuador; in forests up to 1250 m.<br />
Specimens studied. COSTA RICA. ALAJUELA: Nr.<br />
S<strong>an</strong> Ram6n, Cataratas (Los Angeles) de S<strong>an</strong> Ram6n,<br />
17 Apr 1935 (9), Brenes 20542 (F, NY, type collection<br />
This species appears to be related to C. latifolia of C. brenesii); S<strong>an</strong> Carlos, Buena Vista, Finca La Con<strong><strong>an</strong>d</strong><br />
allied taxa.<br />
st<strong>an</strong>cia, 3 Mar 1963 (2), Jimenez 433 (F, MO); Rio S<strong>an</strong><br />
Rafael, 2 km W of La Marina, Ll<strong>an</strong>ura de S<strong>an</strong> Carlos,<br />
21 Feb 1966 (6), Molina R. et al. 17694 (F); S<strong>an</strong> Carlos,<br />
34. <strong>Coussapoa</strong> parviceps St<strong><strong>an</strong>d</strong>ley, Proc. Biol. La Sucre, 1025 m, 2 Mar 1939 (6), A. Smith 1650 (A,<br />
Soc. Wash. 37: 51. 1924; Burger, Fieldi<strong>an</strong>a F, MO), 1 Mar 1939 (2), A. Smith 1939 (6), A. Smith<br />
Bot. 40: 136, t. 22. 1977. Type. Costa Rica. 1650 (A, F, MO), 1 Mar 1939 (2), A. Smith 1663 (F).<br />
CARTAGO:<br />
Puntarenas: Agua Buena valley, nr. Cafnas<br />
Rio Pejibaye, between Rio Taus <strong><strong>an</strong>d</strong> Quebrada<br />
Azul, 28 May 1972 (st), Lent 2538 (COL, F);<br />
Gordas, 1100 m, Feb 1897 (9), Pittier 11166 Moraria de Turrialba, 21 Apr 1975 (st), Poveda 951<br />
(holotype, US; isotype, MO). Fig. 45. (F); nr. Orosi, 30 Mar 1924 (6), St<strong><strong>an</strong>d</strong>ley 39792 (US).<br />
PUNTARENAS: C<strong>an</strong>ias Gordas, Finca Loma Linda, 1140<br />
<strong>Coussapoa</strong> brenesii St<strong><strong>an</strong>d</strong>ley, Publ. Field Mus. Bot. 18: m, 23 Feb 1973 (9), Busey 628 (F, GH, MO); Rio Java,<br />
383. 1937. Type. Costa Rica. Alajuela: nr. S<strong>an</strong> Ra- Las Cruces Bot. Garden, 16 Mar 1978 (9), Hartshorn<br />
m6n, Cataratas (Los Angeles) de S<strong>an</strong> Ram6n, 17 Apr 2157 (U); Agua Buena valley, nr. C<strong>an</strong>ias Gordas, 1100<br />
1935 (Q), Brenes 20542 (holotype, F; isotype, NY). m, Feb 1897 (2), Pittier 11166 (MO, US, type collec-<br />
<strong>Coussapoa</strong> oligoneura Mildbraed, Notizbl. Bot. Gart.<br />
tion).<br />
Berlin 10: 415. 1928. Type. Colombia. Without lo- PANAMA. CHIRIQUi: Burica Peninsula, Mellize, 6<br />
cality, 19 Feb 1892 (9), Tri<strong>an</strong>a 866 (holotype, B; mi S of Puerto Armuelles, 5 Mar 1973 (st), Liesner<br />
isotype, P).<br />
417A (MO). COCLE: Road La Pineda-El Cope, nr. sawmill<br />
above El Cop6, 1000 m, 20 May 1978 (6), Hammel<br />
2576 (U). PANAMA: Road El Ll<strong>an</strong>o-Carti, km 8-12,<br />
13 May 1973 (st), Nee et al. 8822 (MO, NY, U). VERA-<br />
GUAS: NW of S<strong>an</strong>ta F6, 2.7 km from Escuela Agricola<br />
Alto de Piedra, 30 Mar 1975 (2), Mori et al. 5360 (MO,<br />
NY).<br />
COLOMBIA. Without locality, 19 Mar - (2), Tri<strong>an</strong>a<br />
866 (B, P, type collection of C. oligoneura). NARIuO:<br />
Barbacoas, May 1853 (9), Tri<strong>an</strong>a 1866 (NY, photograph).<br />
VALLE: Rio Cajambre, Barco, 21-30 Apr 1944<br />
(2), Cuatrecasas 17351 (F); Rio Anchicaya, Alto Yunda,<br />
1000 m, May 1972 (2), Hilty M87 (MO).<br />
ECUADOR. ESMERALDAS: Alto Tambo, 23 Sep 1965<br />
(8), Little et al. 21136 (NY, Q, US).<br />
<strong>Coussapoa</strong> parviceps differs from other Cous-<br />
sapoa species in the connate pistillate flowers,<br />
which is probably the reason why interfloral<br />
bracts, present in staminate inflorescences, are<br />
lacking in the pistillate ones.<br />
Several characters, particularly as exhibited in<br />
the collection Little & Dixon 21136, suggest<br />
taxonomic relationship to C. cinnamomifolia.<br />
The collections Liesner 417A <strong><strong>an</strong>d</strong> Nee et al.<br />
8822, both from P<strong>an</strong>ama, presumably represent<br />
juvenile specimens of the species. They are dis-<br />
tinct in having narrower (oblong to subovate)<br />
<strong><strong>an</strong>d</strong> larger (up to 50 cm long) laminae <strong>with</strong> up<br />
to 18 pairs of lateral veins, the basal pair of which<br />
reaches the margin below the middle of the lam-<br />
ina; petiole up to 20 cm long; the densely puberu-
<strong>Coussapoa</strong><br />
Fa1to4 I"<br />
FIG. 4.osppc 11 w<br />
.cm<br />
FIG. 45. <strong>Coussapoa</strong> parviceps: 1, leafy twig <strong>with</strong> pistillate inflorescences<br />
(Jimenez 433).<br />
lous twigs, petioles, <strong><strong>an</strong>d</strong> stipules (both sides!);<br />
<strong><strong>an</strong>d</strong> in the presence of white arachnoid hairs on<br />
the margin <strong><strong>an</strong>d</strong> midrib on the lower surface of<br />
the lamina. The characters of these specimens<br />
are not included in the species description. Cous-<br />
sapoa parviceps appears to be the only <strong>Coussapoa</strong><br />
species <strong>with</strong> pronounced dimorphism of leaves<br />
depending upon age.<br />
87<br />
35. <strong>Coussapoa</strong> parvifolia St<strong><strong>an</strong>d</strong>ley, Publ. Field<br />
Mus. Bot. 17: 165. 1937. Type. Brazil. Ama-<br />
zonas: Mun. Sao Paulo de Olivenga, nr. Pal-<br />
mares, 11 Sep-26 Oct 1936 (Q), Krukoff8273<br />
(holotype, NY; isotypes, A, BM, F, G, K, LE,<br />
MO, S, U, US). Figs. 46, 47.<br />
Cousspoa cornifolia St<strong><strong>an</strong>d</strong>ley, Publ. Field Mus. Bot.<br />
17: 160. 1937. Type. Brazil. Amazonas: Mun. Sao
FIG. 46. <strong>Coussapoa</strong> parvifolia: 1, leafy twig <strong>with</strong> pistillate inflorescences (Krukoff8539); 2, leafy twig <strong>with</strong><br />
staminate inflorescences (Pr<strong>an</strong>ce et al. 9054); 3, leaf (Pr<strong>an</strong>ce et al. 22999); 4, stipules (Krukoff 8539).
<strong>Coussapoa</strong> 89<br />
F: 1, ly tg wh 1 cm<br />
FIG. 47. <strong>Coussapoa</strong> parvifolia: 1, leafy twig <strong>with</strong> pistillate inflorescences<br />
(Krukof 8273).<br />
Paulo de Oliven:a, Bel6m Creek, 26 Oct-11 Dec ger brownish patent hairs, soon disappearing;<br />
1936 (d), Krukoff8897 (holotype, NY; isotypes, A, stipules 0.4-1(-1.3) cm long, appressed-puber-<br />
F, G, K, LE, MO, P, S, U, US).<br />
ulous to<br />
<strong>Coussapoa</strong> microcephala subsp. cornifolia Akkerm<strong>an</strong>s subsericeous, terminal buds mostly<br />
& C. C. Berg, Proc. Kon. Ned. Akad. Wetensch., Ser. swollen (containing young inflorescences). Sta-<br />
C, 85(4): 457. 1982.<br />
minate inflorescences repeatedly br<strong>an</strong>ched; heads<br />
ca. 15-30, globose, ca. 2-3 mm diam.; common<br />
Tree, up to to 20 m tall, hemi-epiphytic. Leafy peduncle 1.5-2 cm long, rather sparsely pubertwigs<br />
2-5 mm thick, rather sparsely appressedulous<br />
or partly hirtellous; peri<strong>an</strong>th ca. 1 mm high,<br />
puberulous, sometimes on the younger parts also minutely ciliolate; stamens two, filaments just to<br />
longer, brownish, patent (to appressed) hairs, soon far exceeding the peri<strong>an</strong>th. Pistillate infloresdisappearing.<br />
Lamina coriaceous to subcoria- cences unbr<strong>an</strong>ched or br<strong>an</strong>ched heads (1-5, gloceous,<br />
l<strong>an</strong>ceolate to narrowly ovate, sometimes bose, 4-8 mm diam.; (common) peduncle 1-2.5<br />
subobovate, 4-14(-21) x 1.5-5.5(-11) cm, apex<br />
cm long, puberulous peri<strong>an</strong>th ca. 1 mm high,<br />
acute to acuminate, base acute to obtuse, margin glabrous. Interfloral bracts (sub)spathulate, mientire,<br />
towards the base + revolute; upper sur- nutely puberulous.<br />
face glabrous, lower surface ? densely ap-<br />
Distribution (Fig. 8). Upper Amazon Basin, in<br />
pressed-puberulous to glabrous; lateral veins Brazil (Amazonas), Peru (Loreto), Venezuela<br />
(4-)7-11 pairs, basal pair unbr<strong>an</strong>ched, reaching (Amazonas), also in Extra-Amazoni<strong>an</strong> Colombia<br />
the margin below or in subobovate leaves above (Antioquia <strong><strong>an</strong>d</strong> Vichada); in non-inundated forthe<br />
middle of the lamina; intercostal venation est.<br />
slightly prominent; petiole 0.5-3(-4) cm long, Specimens studied. COLOMBIA. ANTIOQuIA: Rd.<br />
appressed-puberulous to glabrous also <strong>with</strong> lon- Mutat&-Pavar<strong><strong>an</strong>d</strong>6, between haciendas La Esper<strong>an</strong>za
90 Flora Neotropica<br />
<strong><strong>an</strong>d</strong> Mocari, 150 m, 6 Mar 1987 (6), Fonnegra et al. cm long, glabrous; peri<strong>an</strong>th 1-2 mm high, gla-<br />
1798 (BG, HUA). VICHADA: Gaviotas, Caino Ariba, 19<br />
Feb 1973 (9), Cabrera 2668 brous; fruiting peri<strong>an</strong>th yellow or or<strong>an</strong>ge. Inter-<br />
(COL).<br />
VENEZUELA. AMAZONAS: Rio Cucucucuma, be- floral bracts (sub)spathulate, sparsely puberulous<br />
low Raudal Pacure, 9 Nov 1950 (6), Maguire et al. at the apex.<br />
29408 (NY); Dpto. Atabapo, 15 km SE of S<strong>an</strong> Fer- Distribution (Fig. 8). Southern Mexico <strong><strong>an</strong>d</strong><br />
n<strong><strong>an</strong>d</strong>o de Atabapo, 10-16 Feb 1988 (d), Stergios et al. north-western<br />
11530<br />
Guatemala; in evergreen forest in<br />
(BG).<br />
BRAZIL. AMAZONAS: Mun. Sao Paulo de<br />
moist<br />
Olivenoa,<br />
places (often along streams) or in seminr.<br />
Palmares, 11 Sep-26 Oct 1936 (2), Krukoff 8273 deciduous forest; up to 1700 m.<br />
(A, F, G, K, LE, MO, NY, P, S, U, US, type collection<br />
of C. parvifolia), (2), Krukoff8539 (A, BM, F, G, LE, Specimens studied. MEXICO. CHIAPAS: Ov<strong><strong>an</strong>d</strong>o Mt.,<br />
MO, NY, P, S, U, US); Mun. Sao Paulo de 17 Dec 1936<br />
Olivenca,<br />
(6), Matuda 446 (MO, NY, US); S<strong>an</strong>ta<br />
Belem Creek, 26 Oct-11 Dec 1936 (6), Krukoff8658<br />
Rita, Mapastepec, J<strong>an</strong> 1938 (6), Matuda 2020 (A, F,<br />
(A, BM, F, G, K, LE, MO, NY, P, S, U, US), (6), Krukoff<br />
K, NA, NY); between Guatimoc <strong><strong>an</strong>d</strong> Cacahuat<strong>an</strong>, 3<br />
8897 Dec 1941<br />
(A, BM, F, G, K, LE, MO, NY, P, S, U, US, type<br />
(9), Mir<strong><strong>an</strong>d</strong>a 1737 (US); NW of S<strong>an</strong> Fercollection<br />
of C. cornifolia); rd. M<strong>an</strong>aus-Itacoatiara, km n<strong><strong>an</strong>d</strong>o, NW ofTuxtla, 1000 m, 2 Apr 1950 (9), Mir<strong><strong>an</strong>d</strong>a<br />
65-70, 23 Oct 1963 (8), Oliveira 2764 (IAN); rd. Ma-<br />
6164 (US); nr. El Ocote, 30 km NW of Ocozocu<strong>an</strong>tla,<br />
naus-Itacoatiara, km 61, Res. Flor. Walter Egler, 17 24 Mar 1951 (9), Mir<strong><strong>an</strong>d</strong>a 6273 (US), GUERERO: Distr.<br />
Dec 1968 (6), Pr<strong>an</strong>ce et al. 9054 Montes de<br />
(M, MO, NY, P, S,<br />
Oca, S<strong>an</strong> Antonio, 18 Apr 1938 (9), Hinton<br />
U); rd. M<strong>an</strong>aus-Porto Velho, km 510, 5 km N of Rio 14018 (K, NY, P, US). JALISCO: Sierra Madre Occi-<br />
Purusinho, 17 Oct 1974 (6), Pr<strong>an</strong>ce et al. 22999 (INPA,<br />
dental, NW of S<strong>an</strong> Sebasti<strong>an</strong>, Las Mesitas, Arroyo de<br />
NY, U, US). PARA: Rd. Cuiaba-S<strong>an</strong>tar6m, km las<br />
1417, Caaitas, 1700 m, 15 Mar 1927 (6), Mexia 1872 (A,<br />
Rio Itapacura, 25 Nov 1977 (6), Pr<strong>an</strong>ce et al. 25748 BM, F, GH, MO, NY, US). VERACRUZ: Paso del In-<br />
(MO, RB, U).<br />
genio, Jun 1971 (9), Calzada 325 (U); <strong>with</strong>out locality,<br />
1930 (st), Purpus s.n. (B, F, K); Zacualp<strong>an</strong>, Mar 1908<br />
This species belongs to a group <strong>with</strong> two sta- (6), Purpus 5996 (BM, F, GH, MO, NY, US), Mar 1930<br />
mens <strong><strong>an</strong>d</strong> leaf margins which mostly are revolute (8), Purpus 11161 (K), Apr 1928 (6), Purpus 11162 (F,<br />
at the base, comprising e.g., C.<br />
type collection), Jun 1930<br />
microcarpa <strong><strong>an</strong>d</strong><br />
(9), Purpus 11162 (A, K,<br />
MO), Sep 1930 (9), Purpus 11162 (C), Apr 1931 (9),<br />
C. microcephala.<br />
Purpus 11162 (C), May 1933 (6 <strong><strong>an</strong>d</strong> 9), Purpus 11162<br />
Pr<strong>an</strong>ce et al. 9054 is aberr<strong>an</strong>t in having (A), May 1934 (6 <strong><strong>an</strong>d</strong> 9), Purpus 11162 (K, US), Sep<br />
(sub)obovate leaves.<br />
1929 (8), Purpus 11182 (A).<br />
GUATEMALA. QUEZALTENANGO: Colomba, ca.<br />
1000 m, 28 Dec 1934 (6), Skutch 2023 (BM, F, GH,<br />
36. <strong>Coussapoa</strong> purpusii St<strong><strong>an</strong>d</strong>ley, Publ. Field NY). SAN MARCOS: Finca El Porvenir, on Potrero Ma-<br />
Mus. Bot. 8: 6. 1930. Type. Mexico. Veracruz: tas<strong>an</strong>, Rio Cabuis, Volc<strong>an</strong> Tajumulco, 1000-1300 m,<br />
Zacualp<strong>an</strong>, Apr 1928 (6), Purpus 11162 12 Mar<br />
(ho-<br />
1940 (9), Steyermark 37602 (F), 25 Mar 1943<br />
lotype, F). Fig. 48.<br />
(st), Steyermark 52334 (F).<br />
The species appears to be related to C. parviceps.<br />
Tree, hemi-epiphytic or terrestrial, often (?)<br />
deciduous, up to 20 m tall. Leafy twigs 3-6 mm<br />
thick, glabrous or puberulous on the scars of the<br />
stipules. Lamina chartaceous, ovate to elliptic or<br />
to obtuse, occasionally obovate, 4-16 x 1-7 cm,<br />
apex acute to acuminate to obtuse, base obtuse<br />
sometimes subacute or subcordate, margin en-<br />
tire; both surfaces glabrous; lateral veins 4-6 pairs,<br />
curved, basal pair unbr<strong>an</strong>ched reaching the mar-<br />
gin below, at, or slightly above the middle of the<br />
lamina; intercostal venation pl<strong>an</strong>e; petiole 1-5<br />
cm long, glabrous; stipules 1-3 cm long, glabrous<br />
or sparsely to densely appressed-puberulous.<br />
Staminate inflorescences br<strong>an</strong>ched; heads 2-6,<br />
globose, ca. 5-8 mm diam.; common peduncle<br />
2-4 cm long, glabrous; peri<strong>an</strong>th ca. 1 mm high,<br />
glabrous or sparsely minutely puberulous; sta-<br />
mens three, far exceeding the peri<strong>an</strong>th. Pistillate<br />
inflorescences unbr<strong>an</strong>ched; heads globose, 4-6<br />
mm, in fruit up to 10 mm diam.; peduncle 3-4<br />
37. <strong>Coussapoa</strong> scabra Akkerm<strong>an</strong>s & Berg, Proc.<br />
Kon. Ned. Akad. Wetensch. Ser. C 85(4): 460.<br />
1982. Type. Brazil. Rondonia: Rio Ouro Pre-<br />
to, affluent of Rio Pacaas Novos, 18 Sep 1923<br />
(9), Kuhlm<strong>an</strong>n 470 (HJBR 19836) (holotype,<br />
RB; isotypes, RB, U). Fig. 49.<br />
Tree. Leafy twigs 2-5 mm thick, puberulous<br />
to subhirsute; internodes solid. Lamina charta-<br />
ceous, elliptic to ovate or obovate, 4-10 x 2.5-<br />
7 cm, apex acute to shortly acuminate, base ob-<br />
tuse to acute, margin entire; upper surface, sca-<br />
brous, lower surface, scabridulous, sparsely to<br />
rather densely puberulous to hirtellous; lateral<br />
veins 6-13 pairs, straight to curved, basal pair<br />
unbr<strong>an</strong>ched, reaching the margin far below the<br />
middle of the lamina; intercostal venation slight-<br />
ly prominent; petiole 1-3 cm long, densely pu-
FIG. 48. <strong>Coussapoa</strong> purpusii: 1, leafy twig <strong>with</strong> pistillate inflorescences (Hinton 14018); 2, leafy twig <strong>with</strong><br />
staminate inflorescences (Skutch 2023).
92<br />
1cm<br />
Flora Neotropica<br />
FIG. 49. <strong>Coussapoa</strong> scabra: 1, leafy twig <strong>with</strong> pistillate inflorescences<br />
(Kuhlm<strong>an</strong>n 470).<br />
berulous to hirtellous; stipules 0.5-0.7 cm long,<br />
subhirsute. Staminate inflorescences unknown.<br />
Pistillate inflorescences unbr<strong>an</strong>ched or poorly<br />
br<strong>an</strong>ched; heads 1-5, globose, 2-4 mm diam., in<br />
fruit up to 6 mm; (common) peduncle 1-3 cm<br />
long, puberulous; peri<strong>an</strong>th ca. 1 mm high, glabrous.<br />
Interfloral bracts (sub)spathulate, minutely<br />
puberulous at the apex.<br />
Distribution (Fig. 8). Brazil, R6ndonia. Known<br />
only from the type collection.<br />
The position of this species, whose leaves have<br />
a characteristic scabrous <strong><strong>an</strong>d</strong> chartaceous lamina,<br />
is unclear. The shape <strong><strong>an</strong>d</strong> venation of the<br />
lamina show similarities to those found in C.<br />
parvifolia <strong><strong>an</strong>d</strong> C. macerrima.<br />
38. <strong>Coussapoa</strong> sprucei Mildbraed, Notizbl. Bot.<br />
Gart. Berlin 10:415.1928. Type. Brazil. Ama-<br />
zonas: "Rio Negra," probably Rio Taruma,<br />
Dec 1854 (9), Spruce 3782 (holotype, B; iso-<br />
types, BM, BR, C, G, GH, K, LE, NY, OXF,<br />
P). Fig. 50.<br />
Tree or shrub, mostly hemi-epiphytic, up to<br />
30 m tall. Leafy twigs 3-7 mm thick, puberulous<br />
to hirtellous to brownish hirsute, sometimes also
<strong>Coussapoa</strong><br />
2 1cm<br />
FIG. 50. <strong>Coussapoa</strong> sprucei: 1, leafy twig <strong>with</strong> pistillate inflorescences (Spruce 3782); 2, leafy twig <strong>with</strong><br />
staminate inflorescences<br />
(Pr<strong>an</strong>ce et al. 14976).<br />
<strong>with</strong> white arachnoid hairs. Lamina coriaceous,<br />
oblong to elliptic, subovate or to subovate, 5-18<br />
x 3-8 cm, apex shortly acuminate to acute, base<br />
subcordate to rounded, margin entire; upper surface<br />
glabrous, lower surface puberulous in the<br />
areoles, hirtellous (to tomentellous) on the smaller<br />
veins, appressed-puberulous on the main veins,<br />
sometimes also <strong>with</strong> a few distinctly longer appressed<br />
hairs, the whole surface covered <strong>with</strong><br />
rather dense to sparse (sub)persistent white<br />
arachnoid hairs; lateral veins 5-10 pairs, slightly<br />
curved, basal pair unbr<strong>an</strong>ched or inconspicuously<br />
br<strong>an</strong>ched, reaching the margin below the<br />
middle of the lamina; intercostal venation (very)<br />
prominent; petiole 1-4 cm long, puberulous, often<br />
also having distinctly longer straight hairs <strong><strong>an</strong>d</strong><br />
white arachnoid hairs; stipules 0.5-1.5(-2) cm<br />
long, brownish sericeous to subhirsute, terminal<br />
buds often more or less swollen. Staminate inflorescences<br />
br<strong>an</strong>ched; heads m<strong>an</strong>y, globose, ca.<br />
3-4 mm diam.; common peduncle (2-)4-8 cm<br />
long, puberulous to hirtellous; peri<strong>an</strong>th ca. 1 mm<br />
high, glabrous; stamens three, far exceeding the<br />
peri<strong>an</strong>th. Pistillate inflorescences br<strong>an</strong>ched; heads<br />
2-5, globose, ca. 3-5 mm in fruit up to 8 mm<br />
diam.; common peduncle 2-8 cm long, ca. 1 mm<br />
thick, puberulous to hirtellous; peri<strong>an</strong>th ca. 1<br />
mm high, glabrous. Interfloral bracts spathulate<br />
to subpeltate, puberulous, mostly conspicuous.<br />
Distribution (Fig. 8). Brazil, Amazon basin,<br />
near M<strong>an</strong>aus, <strong><strong>an</strong>d</strong> in the basin of the Rio Trombetas;<br />
in forest of terra firme.<br />
93<br />
Specimens studied. BRAZIL. AMAZONAS: Nr. M<strong>an</strong>aus,<br />
Rio Taruma, 27 Oct 1943 (6), Ducke 1354 (A,<br />
F, NY, R, RB, US); nr. M<strong>an</strong>aus, C6lonia Joao Alfredo,<br />
27 Sep 1946 (8), Ducke 1999 (A, NY, R, RB, U); nr.<br />
M<strong>an</strong>aus, Rio Taruma, Cachoeira Gr<strong><strong>an</strong>d</strong>e, 17 Oct 1929<br />
(9), Ducke (HJBR) 23609 (RB); M<strong>an</strong>aus, 17 Oct 1929<br />
(9), Killip & Smith 30179 (NY); Rio Cuieras, 2 km<br />
below mouth ofRio Br<strong>an</strong>cinho, 27 Sep 1971 (6), Pr<strong>an</strong>ce<br />
et al. 14976 (K, M, MO, NY, S, U), 12 Sep 1973 (8),<br />
Pr<strong>an</strong>ce et al. 17847 (F, K, M, MO, NY, P, S, U); "Rio<br />
Negro," possibly Rio Taruma, Dec 1854 (9), Spruce<br />
3782 (B, BM, BR, C, G, GH, K, LE, NY, OXF, P,<br />
type collection). PAIA: Rio Mapuera, 11 Dec 1907 (2),<br />
Ducke (HAMP) 9096 (MG, US).<br />
<strong>Coussapoa</strong> sprucei appears to be closely related<br />
to C. leprieurii. It differs distinctly from the latter<br />
in the smooth <strong><strong>an</strong>d</strong> glabrous upper surface of the<br />
lamina. Other differences are of little signifi-<br />
c<strong>an</strong>ce. C. sprucei also resembles the Central<br />
Americ<strong>an</strong> C. oligocephala, from which it differs<br />
mainly in the length of the stipules <strong><strong>an</strong>d</strong> to a lesser<br />
extent in the nature of the indumentum. This<br />
group of three species is probably related to the<br />
large-leaved taxa C. nymphaeifolia <strong><strong>an</strong>d</strong> C. lon-<br />
gepedunculata, judging from the similarities in<br />
indumentum, leaf venation, <strong><strong>an</strong>d</strong> number of sta-<br />
mens.<br />
39. <strong>Coussapoa</strong> tessm<strong>an</strong>nii Mildbraed, Notizbl.<br />
Bot. Gart. Berlin 10: 413. 1928; Macbride,<br />
Publ. Field Mus. Bot. 13(2.2): 298. 1937. Type.<br />
Peru. Loreto: Mouth of Rio S<strong>an</strong>tiago, 3 Dec<br />
1924 (p), Tessm<strong>an</strong>n 4673 (holotype, B; iso-<br />
types, G, S). Fig. 51.
94 Flora Neotropica<br />
FIG. 51. <strong>Coussapoa</strong> tessm<strong>an</strong>nii: 1, leafy twig <strong>with</strong> pistillate inflorescences (Krukoff 8994); 2, staminate<br />
inflorescences (Krukoff 1187).
<strong>Coussapoa</strong><br />
Tree or shrub, (mostly) hemi-epiphytic, up to PERU. LORETO: Prov. Maynas, Rio Yaguasyuca, af-<br />
20 m tall. Leafy twigs 5-13 mm, brown<br />
fluent of Rio<br />
puber-<br />
Ampiyacu, nr. Brillo Nuevo, 2 Mar 1978<br />
(2), Balick et al. 1021 (MO, U); road Zungarochaulous<br />
to strigillose to hirtellous to strigose or to Puerto Almendra, 3 Dec 1964 (2), Dodson et al. 3017<br />
(sub) velutinous, often also <strong>with</strong> a dense covering (F); Rio Yavari, behind Angamo Garrison, 5 Oct 1973<br />
of (long) dark brown pluricellular hairs <strong><strong>an</strong>d</strong> (a), Lleras et al. P17178 (F, K, NY, P, S, U); Puerto<br />
sometimes whitish arachnoid hairs as well. Lam- Almendra, nr. Rio N<strong>an</strong>ay, 23 Feb 1977 (a), Revilla<br />
2378<br />
ina coriaceous, ovate to elliptic, 13-32 x 11-20<br />
(U); mouth of Rio S<strong>an</strong>tiago, 3 Dec 1924 (2),<br />
Tessm<strong>an</strong>n 4673 (B, G, S, type collection).<br />
cm, apex subacute to obtuse to rounded, base BRAZIL. AMAZONAS: Rio Negro, Uaup6s, Jauarite,<br />
obtuse to subcordate, margin entire to subcren- 23 Aug 1945 (9), Froes 21264 (NY); Mun. Sao Paulo<br />
ate; upper surface glabrous, lower surface densely<br />
de Olivenca, Belem Creek, 26 Oct-11 Dec 1936 (2),<br />
minutely puberulous in the areoles <strong><strong>an</strong>d</strong> on the Krukoff8994 (A, G, K, LE, NY, P, S, U); road M<strong>an</strong>aus-Porto<br />
Velho, km 379, Rio Jutai, 12 Oct 1974<br />
reticulum, to (sparsely) hirtellous on the (paral- (9), Pr<strong>an</strong>ce et al. 22836 (MG, NY, U); Rio Javari, 7<br />
lel) tertiary venation, appressed-puberulous to hours above Paumari, 16 Oct 1976 (2), Pr<strong>an</strong>ce et al.<br />
strig(ill)ose on the midrib <strong><strong>an</strong>d</strong> lateral veins, the 23846 (NY, U). PARA: Cupary, Sep 1931 (6), da Costa<br />
whole surface <strong>with</strong> rather sparse to dense sub- 90 (IAN); upper Rio Cupary, Sep 1931 (a), Krukoff<br />
1147<br />
persistent white to pale brown arachnoid<br />
(A, BM, G, K, NY, P, S, U), Krukoff 1187 (A,<br />
hairs; BM, G, K, MO, NY, S, U); road S<strong>an</strong>tarem-Palhao,<br />
lateral veins 9-16 pairs, straight, main basal pair km 70, 15 Sep 1969 (8), M. Silva et al. 2591 (F, K,<br />
br<strong>an</strong>ched, sometimes faintly so (in small leaves MG, NY, S, U, US). RONDONIA: Porto Velho, 15 Sep<br />
sometimes unbr<strong>an</strong>ched), reaching the margin far 1975 (a), Mota et al. 154 (U).<br />
below or sometimes just below the middle of the In the vegetative parts C. tessm<strong>an</strong>nii shows<br />
lamina; intercostal venation very to slightly similarities to C. villosa, but it is distinct from<br />
prominent; petiole 3.5-9 cm long, 2-5 mm thick, the latter in having a ramified, <strong><strong>an</strong>d</strong> thus pluribrown(ish)<br />
puberulous to hirtellous to subvelucapitate,<br />
pistillate inflorescence <strong><strong>an</strong>d</strong> a unistamitinous,<br />
often also <strong>with</strong> dark brown pluricellular nate flower. C. tessm<strong>an</strong>nii shows distinct affinhairs<br />
<strong><strong>an</strong>d</strong> sometimes arachnoid hairs; stipules 2- ities <strong>with</strong> C. cinnamomea, e.g., in the relatively<br />
3(-9) cm long, yellowish to brown strigose to thick peduncle <strong><strong>an</strong>d</strong> br<strong>an</strong>ches of the pistillate insubhirsute<br />
or also <strong>with</strong> dark brown pluricellular florescence, <strong><strong>an</strong>d</strong> it c<strong>an</strong> be easily confused <strong>with</strong><br />
hairs. Staminate inflorescences br<strong>an</strong>ched; heads C. nitida. Several collections made in the Rio<br />
m<strong>an</strong>y, crowded at the end of the br<strong>an</strong>ches, Tapajos area (da Costa 90, Krukoff 1147 <strong><strong>an</strong>d</strong><br />
(sub)globose, 2-4 mm diam., often fused; com- 1187, <strong><strong>an</strong>d</strong> M. Silva et al. 2591) differ from the<br />
mon peduncle 1-3 cm long, brown puberulous other collections in the presence of a dense covto<br />
hirtellous to short-velutinous or also (densely) ering of dark brown moniliform pluricellular (=<br />
covered <strong>with</strong> dark brown pluricellular hairs; peri- gr<strong>an</strong>ular) hairs on the leafy twig, petiole, <strong><strong>an</strong>d</strong><br />
<strong>an</strong>th ca. 1 mm high, puberulous; stamen one, inflorescence. Moreover, the lamina tends to be<br />
just exceeding the peri<strong>an</strong>th. Pistillate inflores- elliptic rather th<strong>an</strong> ovate (as is normal in the<br />
cences br<strong>an</strong>ched; heads 2-7, (sub)globose, 5-10<br />
species). In the vegetative parts, these collections<br />
mm, in fruit up to 15 mm diam.; common pe- show striking similarities to the aberr<strong>an</strong>t pistilduncle<br />
2-3 cm long, peduncle <strong><strong>an</strong>d</strong> br<strong>an</strong>ches 2- late collection (Pr<strong>an</strong>ce et al. 26402) assigned to<br />
4 mm thick, appressed puberulous to hirtellous C. villosa (see p. 104).<br />
to short-velutinous or also <strong>with</strong> dark brown pluricellular<br />
hairs; peri<strong>an</strong>th ca. 1 mm high, (almost)<br />
glabrous, sometimes muriculate.<br />
40.<br />
Interfloralbracts<br />
<strong>Coussapoa</strong> trinervia Spruce ex Mildbraed,<br />
(sub)spathulate to subpeltate, puberulous at the<br />
Notizbl. Bot. Gart. Berlin 10:416. 1928. Type.<br />
apex; extrafloral bracts often present on the<br />
Venezuela. Terr. Fed. Amazonas: Rio Casibr<strong>an</strong>ches<br />
of staminate inflorescences.<br />
quiare, between Lago de Vasiva <strong><strong>an</strong>d</strong> Rio Pa-<br />
Distribution (Fig. 9). Amazon Basin cimoni, 1853-1854<br />
(Brazil,<br />
(2), Spruce 3464 (holotype,<br />
Peru, <strong><strong>an</strong>d</strong> Colombia); in upl<strong><strong>an</strong>d</strong> <strong><strong>an</strong>d</strong> riverside B; isotypes, BM, BR, G, GH, K, LE, NY, OXF,<br />
forest.<br />
P). Fig. 52.<br />
Specimens examined. COLOMBIA. VAUPES: Rio<br />
Tree, terrestrial, up to 20 m tall, mostly <strong>with</strong><br />
Papuri, nr. Montfort Mission, 3 Sep 1943 (s), P. H. stilt roots. Leafy twigs 3-8 mm thick, glabrous.<br />
Allen 3103 (COL).<br />
Lamina coriaceous, subobovate to obovate to<br />
95
2<br />
1cm<br />
FIG. 52. <strong>Coussapoa</strong> trinervia: 1, leafy twig <strong>with</strong> pistillate inflorescences (Berg et al. P18453); 2, leafy twig<br />
<strong>with</strong> staminate infiorescences (Schunke 371).
<strong>Coussapoa</strong> 97<br />
oblong to elliptic, 4-14 x 1.5-7 cm, apex obtuse<br />
to shortly acuminate, base (sub)acute to cuneate,<br />
margin entire; both surfaces glabrous; trinervate,<br />
the single pair of lateral veins, unbr<strong>an</strong>ched,<br />
reaching the margin near or at the apex of the<br />
lamina; intercostal venation promin<strong>an</strong>t; petiole<br />
1-4 cm long, glabrous; stipules 0.5-1.5 cm long,<br />
glabrous or occasionally appressed-puberulous.<br />
Staminate inflorescences br<strong>an</strong>ched; heads 3-9,<br />
often partly fused, globose, 2-5 mm diam.; com-<br />
mon peduncle 1-3 cm long, glabrous or sparsely<br />
puberulous; peri<strong>an</strong>th ca. 1 mm high, glabrous;<br />
stamen one, just exceeding the peri<strong>an</strong>th. Pistil-<br />
late inflorescences unbr<strong>an</strong>ched; heads globose, ca.<br />
5-7 mm, in fruit up to 10 mm diam.; peduncle<br />
1-3 cm long, glabrous or sparsely puberulous;<br />
peri<strong>an</strong>th ca. 1 mm high, glabrous. Interfloral<br />
bracts subspathulate to peltate, sparsely puber-<br />
ulous at the apex.<br />
Distribution (Fig. 9). Amazon Basin (Brazil,<br />
Peru, Ecuador, Colombia, <strong><strong>an</strong>d</strong> Venezuela); the<br />
main area apparently in the north-western part<br />
of Amazonia-elsewhere in smaller, more re-<br />
stricted, areas; on periodically inundated b<strong>an</strong>ks<br />
(always?) of black-water rivers.<br />
Rio Maz<strong>an</strong>, Gamit<strong>an</strong>ococha, 14 Mar 1935 (6), Schunke<br />
371 (A, F, NA, NY).<br />
BRAZIL. AMAZONAS: Rio Negro Basin, Tapuruquara,<br />
21 Oct 1971 (9), Pr<strong>an</strong>ce et al. 15764 (F, GH,<br />
K, NY, S, U); Rio Uaup6s, P<strong>an</strong>ur6, Oct 1852-J<strong>an</strong> 1853<br />
(Y), Spruce 2616 (B, BM, BR, C, G, GH, K, LE, NY,<br />
OXF, P). MARANHXO: Turiaga, Astonas, 6 Dec 1978<br />
(2), Rosa et al. 2874 (U). MATO GROSSO: Rio Aripuafia,<br />
nr. Humboldt Center, 10?12'S, 59?21'W, 10 Oct 1973<br />
(a), Berg et al. P18409 (F, K, MO, NY, P, U, US), 12<br />
Oct 1973 (2), Berg et al. P18453 (K, MO, NY, S, U,<br />
US); Rio Juruena, Cachoeira de Sao Joao da Barra, 3<br />
Jun 1977 (a), Rosa et al. 2053 (NY); Rio Juruena,<br />
Igarape Chuini, 15 Jul 1977 (2), M. G. Silva et al. 3345<br />
(U). PARA: Mun. Itapir<strong>an</strong>ga, Rio Uamata, nr. Igarap6<br />
S<strong>an</strong>ta Luzia, 16 Aug 1974 (2), Cid et al. 435 (U), nr.<br />
Cachoeira do Tucumari, 19 Aug 1974 (Y), Cid et al.<br />
511 (U); Belem, between Val de Caes <strong><strong>an</strong>d</strong> Sao Joaquin,<br />
7 Nov 1922 (2), Ducke (HJBR) 18455 (RB); Belem,<br />
Sep-Oct 1961 (st), Pires 51736 (NY).<br />
This remarkable species, probably the only<br />
truly terrestrial species, resembles terrestrial fig<br />
trees (like F. trigona L. f.) in habit <strong><strong>an</strong>d</strong> has me-<br />
Specimens studied. COLOMBIA. CAQUETA: Rio Caqueta,<br />
Sol<strong>an</strong>o, 8 km SE ofTres Esquinas, below mouth<br />
of Rio Orteguaza, 7 Mar 1945 (st), Little et al. 9629<br />
(COL); Rio Apaporis, between Rio Pacoa <strong><strong>an</strong>d</strong> Rio<br />
K<strong>an</strong><strong>an</strong>ari, Soratama, 16 Aug 1961 (d), Schultes et al.<br />
13589 (GH, U).<br />
VENEZUELA. AMAZONAS: Rio M<strong>an</strong>ariche, Kalohi-teri,<br />
- lastomataceous leaves. It appears to be confined<br />
to periodically inundated b<strong>an</strong>ks of black-water<br />
rivers; this may explain the disjunct distribution.<br />
<strong>Coussapoa</strong> trinervia appears to be closely related<br />
to C. <strong>an</strong>gustifolia, as the two species have m<strong>an</strong>y<br />
characters in common. The difference in the<br />
number of stamens suggests that C. trinervia <strong><strong>an</strong>d</strong><br />
C. cinnamomifolia are not related, in spite of<br />
similarities in the leaf venation.<br />
41. <strong>Coussapoa</strong> v<strong>an</strong>nifolia Cuatrecasas, Caldasia<br />
(2), Lizot 83 (VEN); Rio Yatua, just above 7: 296. 1956.<br />
Piedra Arauicaua, 28 Sep 1957 (2), Maguire et al.<br />
Type. Colombia. Valle: Rio An-<br />
41620<br />
(K), Esmeralda-Oc<strong>an</strong>a, Mavaca, 23 Apr 1968 (2), Me- chicayf, between Sabuletas <strong><strong>an</strong>d</strong> Quebrada Tadina<br />
397 (VEN); Rio Casiquiare, between Lago de Va- tabro, 28-29 Sep 1946 (9), Cuatrecasas 22048<br />
siva <strong><strong>an</strong>d</strong> Rio Pacimoni, 1853-1854 (9), Spruce 3464 (holotype, F). Fig. 53.<br />
(B, BM, BR, G, GH, K, LE, NY, OXF, P, type collection);<br />
Rio Yatua, just above Piedra Arauicaua, 16 Jul <strong>Coussapoa</strong> rotunda Little, Phytologia 18: 196. 1969.<br />
1959 (9), Wurdack et al. 43485 (K).<br />
Type. Ecuador. Esmeraldas: Junction of Rio Hoja<br />
ECUADOR. NAPO: Rio Cuyabeno, ca. 0?10'S, Bl<strong>an</strong>ca <strong><strong>an</strong>d</strong> Rio Hualpi, 50 km S of Borb6n, 14 Sep<br />
76?55'W, 16 Feb 1980 (d), Berg & Akkerm<strong>an</strong>s 1038 1965 (6), Little & Dixon 21056 (holotype, US; iso-<br />
(U), 17 Feb 1980 (2), Berg & Akkerm<strong>an</strong>s 1048 (U), 18 types, NY, Q).<br />
Feb 1980 (2), Berg & Akkerm<strong>an</strong>s 1058 (U); Laguna<br />
Cuyabeno, 7 Aug 1980 (9), Jaramillo et al. 2888 (AAU),<br />
Tree, terrestrial or hemi-epiphytic, up to 20 m<br />
PASTAZA: Rio Capihuari, 23 Jul 1980 (st), Ollgaard et tall. Leafy twigs 5-9 mm thick, white to brown<br />
al. 35068 (AAU).<br />
hirtellous to hirsute. Lamina coriaceous,<br />
PERU.<br />
broadly<br />
LORETO: Quebrada Cuninico, 7 Jul 1972 (2), ovate to cordiform to subreniform<br />
Croat<br />
or to<br />
17784<br />
subor-<br />
(F, NY); Rio Tacsha Curary, 18 Sep 1972<br />
(a), Croat 20411 (C, F, GH, L, MO, NA, NY, P,<br />
bicular, 4-22 x 4-25<br />
S);<br />
cm, apex rounded to ob-<br />
Rio Yavari, Emilia, above Atalaia del Norte, 22 Nov tuse, base cordate, occasionally obtuse, margin<br />
1977 (2), Gentry et al. 20766 (MO, U); Prov. Maynas, entire or subcrenate; upper surface glabrous, low-<br />
Rio Mom6n, tributary of Rio N<strong>an</strong>ay, nr. Iquitos, 9 er surface rather<br />
Dec 1979 (2), J. Jones et<br />
densely white puberulous to hiral.<br />
9788 (LAM, U); Rio Na- tellous on the main<br />
nay, Quebrada Morropon, 26 Jun 1972 (2), McD<strong>an</strong>iel<br />
veins, otherwise glabrous or<br />
16344 (NA); Rio N<strong>an</strong>ay, Mapa Cocha, 17 Mar 1976 sparsely puberulous to hirtellous on the tertiary<br />
(a), McD<strong>an</strong>iel et al. 20555 (NA); Rio N<strong>an</strong>ay, Quebrada (parallel) venation; lateral veins 5-9 pairs, in-<br />
Morropon, 31 May 1976 (2), Rimachi 2297 (F, MO), cluding 2-4 pairs of basal veins, basal veins
FG53 Cosaov<strong>an</strong>fla1leftwgwtstmntifoecne Apud141'<br />
....... cm .... I<br />
FIG. 53. <strong>Coussapoa</strong> v<strong>an</strong>nifolia: 1, leafy twig <strong>with</strong> staminate infiorescences (Asplund 1640
<strong>Coussapoa</strong><br />
br<strong>an</strong>ched, the upper pair of basal veins reaching<br />
the margin at or above the middle of the lamina,<br />
the other lateral veins often poorly br<strong>an</strong>ched (furcate);<br />
intercostal venation almost pl<strong>an</strong>e; petioles<br />
1-6 cm long, (rather) densely white to brownish<br />
hirtellous to hirsute; stipules 2-8 cm long, densely<br />
puberulous to hirtellous to strigillose, mostly<br />
also <strong>with</strong> brown pluricellular hairs. Staminate<br />
inflorescences br<strong>an</strong>ched; heads m<strong>an</strong>y, globose, ca.<br />
1.2 mm diam.; common peduncle 1-2 cm long,<br />
hirtellous to subtomentose; peri<strong>an</strong>th ca. 1 mm<br />
high, apex minutely puberulous; stamens three,<br />
just exceeding the peri<strong>an</strong>th. Pistillate inflorescences<br />
br<strong>an</strong>ched; heads 3-5, free or sometimes<br />
partly fused, (sub)globose, 6-9 mm diam.; common<br />
peduncle 1.5-2.5 cm long, hirtellous to subtomentose;<br />
peri<strong>an</strong>th ca. 1 mm high, papillate.<br />
Interfloral bracts present, subpeltate, at the apex<br />
minutely puberulous, in pistillate inflorescences<br />
often only a few.<br />
Distribution (Fig. 9). Ecuador <strong><strong>an</strong>d</strong> Colombia,<br />
Pacific coastal region; in lowl<strong><strong>an</strong>d</strong> rain forest.<br />
Specimens studied. COLOMBIA. CAUCA: Rio Timbiqui,<br />
- <strong>Coussapoa</strong> vellerea Klotzsch, Linnaea 20: 527. 1847;<br />
Miquel, Fl. bras. 4(1): 139. 1853; Macbride, Publ.<br />
Field Mus. Bot. 13(2.2): 299. 1937. Syntypes. Peru,<br />
Hu<strong>an</strong>uco: Macora,<br />
(8), Lehm<strong>an</strong>n BT948 (B, GH, K, NY). VALLE:<br />
Rio Callima, between Pailon <strong><strong>an</strong>d</strong> El Coco, 23 May<br />
1946 (Q), Cuatrecasas 21256 (F); Rio Anchicaya, between<br />
Sabuletas <strong><strong>an</strong>d</strong> Quebrada de Tatabro, 28-29 Sep<br />
1946 (Q), Cuatrecasas 22048 (F, type collection).<br />
ECUADOR. ESMERALDAS: Junction of Rio Hoja<br />
Bl<strong>an</strong>ca <strong><strong>an</strong>d</strong> Rio Hualpi, 14 Sep 1965 (6), Little et al.<br />
21056 (NY, Q, US, type collection of C. rotunda). Los<br />
Rios: Rio Palenque, along road after crossing Rio Bimbe<br />
<strong><strong>an</strong>d</strong> Rio Waija, 7 Oct 1976 (Q), Dodson et al. 6513<br />
(MO, QCA). PICHINCHA: Nr. S<strong>an</strong>to Domingo de los<br />
Colorados, 18 May 1955 (a), Asplund 16401 (S), 20<br />
Aug 1920 (st), Benoist 3042 (P, U), 5 Feb 1979 (Q),<br />
Dodson 7416 (SEL, U), 5 Feb 1979 (6), Dodson et al.<br />
7616 (MO), 9 Apr 1892 (9), Sodiro 193/12 (B).<br />
In the shape <strong><strong>an</strong>d</strong> venation of the leaves, C.<br />
v<strong>an</strong>nifolia is reminiscent of broad-leaved forms<br />
of C. asperifolia ssp. magnifolia but also of C.<br />
batavorum. Because of the difference in the number<br />
of stamens, a taxonomic relationship <strong>with</strong> C.<br />
asperifolia is unlikely.<br />
42. <strong>Coussapoa</strong> villosa Poeppig & Endlicher, Nov.<br />
gen. sp. pl. 2: 33, t. 147. 1838; Trecul, Ann.<br />
Sci. Nat. Bot. S6r. 3, 8: 99. 1847; Miquel, Fl.<br />
bras. 4(1): 138. 1853; Macbride, Publ. Field<br />
Mus. Bot. 13(2.2): 299. 1937. Type. Peru.<br />
Hu<strong>an</strong>uco: Casapi (fide Macbride, 1937) ($ <strong><strong>an</strong>d</strong><br />
2 ?) Poeppig s.n. (holotype, W, destroyed; isotypes<br />
(6), B, OXF). Fig. 54.<br />
- (6 <strong><strong>an</strong>d</strong> 2), Ruiz & Pavon s.n.<br />
(or 3) (holotype (2), B; isotype (2), FI; isotypes (8),<br />
BR, G, OXF, US).<br />
<strong>Coussapoa</strong> marti<strong>an</strong>a (Miquel, Fl. bras. 4(1): 132, t. 42.<br />
1853. Type. Brazil. Without locality ("prov. Paraensi<br />
et Rio Negro"), - (2), Martius s.n. (holotype, M).<br />
<strong>Coussapoa</strong> subinc<strong>an</strong>a Miquel, Fl. bras. 4(1): 134, t. 45.<br />
1853. Type. Brazil. Without locality, - (6), Martius<br />
s.n. (holotype, M; isotypes, BR, M, U).<br />
<strong>Coussapoa</strong>p<strong>an</strong>amensis Pittier, Contr. U.S. Natl. Herb.<br />
18: 226. 1917; Woodson & Schery, Ann. Missouri<br />
Bot. Gard. 47: 169, t. 59. 1960; Burger, Fieldi<strong>an</strong>a<br />
Bot. 40:135, t. 22. 1977. Type. P<strong>an</strong>ama. Col6n: Rio<br />
Fat6, Jul 1911 (9), Pittier 3892 (holotype, US; isotypes,<br />
B, GH, NY).<br />
<strong>Coussapoa</strong> donnell-smithii Mildbraed, Notizbl. Bot.<br />
Gart. Berlin 10: 414. 1928. Type. Costa Rica. Cartago:<br />
Rio Turrialba, Mar 1894 (2), Donnell Smith<br />
4826 (holotype, US; isotypes, GH, K, US).<br />
<strong>Coussapoa</strong> gr<strong><strong>an</strong>d</strong>iceps Killip in Macbride, Publ. Field<br />
Mus. Bot. 13(2.2): 297. 1937. Type. Peru. Loreto:<br />
Lower Rio Huallaga, Y<strong>an</strong>6n, 5-11 Sep 1929 (2), Killip<br />
& Smith 29246 (holotype, US).<br />
<strong>Coussapoa</strong> st<strong><strong>an</strong>d</strong>leyi Macbride, Publ. Field Mus. Bot.<br />
13(2.2): 298. 1937. Type. Peru. Hu<strong>an</strong>uco: Huamalies,<br />
between Monzon <strong><strong>an</strong>d</strong> Rio Huallaga, - (9), Weberbauer<br />
3702 (holotype, B, destroyed; isotype, G).<br />
<strong>Coussapoa</strong> ar<strong>an</strong>eosa St<strong><strong>an</strong>d</strong>ley, Publ. Field Mus. Bot.<br />
17: 158. 1937. Type. Brazil. Amazonas: Mun. Humayta,<br />
Tres Casas, 14 Sep-11 Oct 1934 (2), Krukoff<br />
6524 (holotype, F; isotypes, A, BM, K, LE, MO, NY,<br />
S, U, US).<br />
<strong>Coussapoa</strong> bolivi<strong>an</strong>a St<strong><strong>an</strong>d</strong>ley, Publ. Field Mus. Bot.<br />
17: 159. 1937. Type. Bolivia. S<strong>an</strong>ta Cruz: Prov. Ichilo,<br />
Rio Vibora, 5 Oct 1926 (2), Steinbach 7567 (holotype,<br />
F; isotypes, A, GH, K, MO, S, U).<br />
<strong>Coussapoa</strong> eggersii St<strong><strong>an</strong>d</strong>ley, Publ. Field Mus. Bot. 17:<br />
160. 1937. Type. Ecuador. M<strong>an</strong>abi: El Recreo, Dec<br />
1891 (6), Eggers 14165 (holotype, F; isotypes, probably<br />
in A, BR, L, M, US).<br />
<strong>Coussapoa</strong> embir<strong>an</strong>a St<strong><strong>an</strong>d</strong>ley, Publ. Field Mus. Bot.<br />
17:161. 1937. Type. Brazil. Amazonas: Basin of Rio<br />
Jurua, nr. mouth of Rio Embira (tributary of Rio<br />
Tarauaca), 26 Jun 1933 (6), Krukoff 4994 (holotype,<br />
NY; isotypes, A, G, K, U, US).<br />
<strong>Coussapoa</strong> lawr<strong>an</strong>cei St<strong><strong>an</strong>d</strong>ley, Publ. Field Mus. Bot.<br />
17: 164. 1937. Type. Colombia. Boyaca: El Humbo,<br />
130 km N of Bogota, ca. 1000 m, 26 Apr 1933 (6),<br />
Lawr<strong>an</strong>ce 769 (holotype, F; isotypes, A, B, F, FI, G,<br />
K, MO, S, US).<br />
<strong>Coussapoa</strong> lehm<strong>an</strong>nii St<strong><strong>an</strong>d</strong>ley, Publ. Field Mus. Bot.<br />
17: 164. 1937. Type. Ecuador. Azuay: Western Andes<br />
of Cuenca, Chacayacu, - (2), Lehm<strong>an</strong>n 5606<br />
(holotype, F; isotype, K).<br />
<strong>Coussapoa</strong> subcrenata St<strong><strong>an</strong>d</strong>ley, Publ. Field Mus. Bot.<br />
17: 166. 1937. Type. Brazil. Amazonas: Mun. Tefe,<br />
Rio Solim6es, Paleta, 21 May 1933 (6), Krukoff4518<br />
(holotype, F; isotypes, A, BM, G, K, M, MO, NY,<br />
S, U, US).<br />
<strong>Coussapoa</strong> d<strong>an</strong>ielis Cuatrecasas, Caldasia 7: 290. 1956.<br />
99
100 Flora Neotropica<br />
FIG. 54. <strong>Coussapoa</strong> villosa: 1, leafy twig <strong>with</strong> pistillate inflorescences (Steyermark et al. 95170); 2, twig <strong>with</strong><br />
stipules <strong><strong>an</strong>d</strong> staminate inflorescences (Elias 631); 3, pistillate inflorescences (Donnell Smith 4826).<br />
3<br />
1 cm
<strong>Coussapoa</strong><br />
Type. Colombia. Antioquia: Jardin, 1800 m, Dec<br />
1941 (2), Bro. D<strong>an</strong>iel 2652 (holotype, US; isotypes,<br />
COL, F).<br />
<strong>Coussapoa</strong> macarenensis Cuatrecasas, Caldasia 7: 292.<br />
1956. Type. Colombia. Meta: Sierra de la Macarena,<br />
Caino Entrada, 24 Dec 1949 (5), Phillipson & Idrobo<br />
1914 (holotype, BM; isotypes, COL, US).<br />
<strong>Coussapoa</strong> macarenensis Cuatrecasas var. <strong>an</strong>tioquiensis<br />
Cuatrecasas, Caldasia 7: 293. 1956. Type. Colombia,<br />
Antioquia: Rio Cauca, Puento Valdiva, 17-<br />
20 Feb 1942 (2), Metcalf& Cuatrecasas 30093 (holotype,<br />
F; isotypes, A, MO, US).<br />
<strong>Coussapoa</strong> mutisii Killip & Cuatrecasas, Caldasia 7:<br />
293. 1956. Type. Colombia. Without locality, - lutinous or to (sub)hirsute, often also <strong>with</strong> reddish-brown<br />
pluricellular hairs <strong><strong>an</strong>d</strong> sometimes<br />
arachnoid hairs; peri<strong>an</strong>th ca. 1 mm high, densely,<br />
minutely puberulous; stamens two, (rather) far<br />
exceeding the peri<strong>an</strong>th. Pistillate inflorescence<br />
mostly unbr<strong>an</strong>ched, sometimes poorly br<strong>an</strong>ched;<br />
heads 1(-4), free or occasionally partly fused,<br />
(sub)globose, ca. 5-10 mm, in fruit up to 40 mm<br />
diam.; (common) peduncle (1-)2-13 cm long, 1-<br />
4 mm thick, densely to sparsely puberulous to<br />
hirtellous to shortly velutinous or to (sub)hirsute,<br />
(2),<br />
Mutis 646<br />
often also <strong>with</strong> reddish-brown<br />
(holotype, US).<br />
pluricellular hairs<br />
<strong>Coussapoa</strong> pl<strong>an</strong>itiensis Cuatrecasas, Caldasia 7: 295. <strong><strong>an</strong>d</strong> sometimes arachnoid hairs; peri<strong>an</strong>th ca. 1-<br />
1956. Type. Colombia. Vaupes: Rio Guayabero, 8 2 mm high, densely minutely white to brownish<br />
Nov 1939 (6), Cuatrecasas 7510 (holotype, US). puberulous; fruiting peri<strong>an</strong>th at the apex green,<br />
below the apex or<strong>an</strong>ge. Interfloral bracts<br />
(sub)spathulate to subpeltate, white to brownish<br />
puberulous at the apex.<br />
Distribution (Fig. 9). Central America (from<br />
Guatemala to P<strong>an</strong>ama), the Ande<strong>an</strong> region (Ecuador<br />
to Venezuela), extending to the coastal<br />
mountain r<strong>an</strong>ge of Venezuela, <strong><strong>an</strong>d</strong> wide-spread<br />
(especially) in the (Upper) Amazon Basin; in upl<strong><strong>an</strong>d</strong>,<br />
riverine, <strong><strong>an</strong>d</strong> mountain forest, up to 2400<br />
m; often in secondary vegetation.<br />
Tree or shrub, hemi-epiphytic or terrestrial, up<br />
to 35 m tall. Leafy twigs 5-15 mm thick, sparsely<br />
to densely white (to brown) puberulous to hirtellous<br />
to (sub)hirsute to (sub)villous. Lamina<br />
coriaceous, broadly ovate to subovate, sometimes<br />
to elliptic, occasionally to obovate, 6-60<br />
x 5-40 cm, apex obtuse to acute, sometimes<br />
rounded to emarginate, base obtuse to cordate<br />
(sometimes deeply cordate <strong>with</strong> overlapping<br />
lobes), margin entire to subcrenate; upper surface<br />
glabrous, lower surface sparsely to densely minutely<br />
puberulous in the areoles <strong><strong>an</strong>d</strong> mostly also<br />
on the reticulum, hirtellous to hispidulous on the<br />
(parallel) tertiary venation, sometimes puberu-<br />
Specimens studied. GUATEMALA. IZABAL: Peten<br />
road, 2 km from Fronteras, Rio Ju<strong>an</strong> Vicente, 11 May<br />
1971 (8), Contreras 10763 (DUKE, MO, P, US); Quirigua,<br />
15-31 Mar 1922 (6), St<strong><strong>an</strong>d</strong>ley 23766 (GH, NY),<br />
lous to (sub)hirsute, on the midrib <strong><strong>an</strong>d</strong> the lateral 26-27 Apr 1939 (9), St<strong><strong>an</strong>d</strong>ley 72288 (F, NY); Lago<br />
veins sparsely, sometimes rather densely, puber- Izabal, between Punta dos Reales <strong><strong>an</strong>d</strong> Punta de Leulous<br />
to hirtellous or to (sub)hirsute, the whole<br />
chuga, 17 Apr 1940 (2), Steyermark 39618 (F).<br />
HONDURAS. ATLANTIDA: La Ceiba, 22 Sep 1916<br />
surface covered <strong>with</strong> sparse to dense whitish to (9), Dyer A85 (F, US); L<strong>an</strong>cetilla Valley, nr. Tela, 6<br />
brownish subpersistent arachnoid hairs; lateral Dec 1927-20 Mar 1928 (st), St<strong><strong>an</strong>d</strong>ley 53230 (A, F,<br />
veins (7-)9-24, straight or slightly curved, main US), (9), St<strong><strong>an</strong>d</strong>ley 53991 (A, F, US), (st), St<strong><strong>an</strong>d</strong>ley 54325<br />
basal pair br<strong>an</strong>ched, reaching the margin below (A, F), (2), St<strong><strong>an</strong>d</strong>ley 55155 (A, F, US), (st), St<strong><strong>an</strong>d</strong>ley<br />
55445 (A, F,<br />
the middle of the lamina, in<br />
US), (6), St<strong><strong>an</strong>d</strong>ley 56623<br />
large leaves<br />
(F). COMAYAGUA:<br />
some- Lake Yojoa, Pito Solo, nr. Pito Solo, 16 Apr 1951 (6),<br />
times also <strong>with</strong> other lateral veins poorly P. H. Alien 6186 (F, GH), 8 Aug 1932 (st), Edwards<br />
br<strong>an</strong>ched (furcate); intercostal venation more or 400 (A, F, US); 15 km E of Lake Yojoa, 20 May 1974<br />
less prominent; petiole (2-)3-20 cm long, white (6), Hazelett s.n. (MO); nr. Pito Solo, 14 Aug 1973 (2),<br />
Hazelett 664<br />
(to yellowish) puberulous to hirtellous to<br />
(MO); Lake Yojoa, 29 Jul-10 Aug 1951<br />
(6), Howard et al. 605 (A); nr. Pito Solo, 18 Apr 1945<br />
(sub)hirsute or to (sub)villous; stipules 3-20 cm (2), J. V. Rodriguez 2929 (F); Rio Tempemechin, ca.<br />
long, puberulous to hirtellous to strigose to hir- 6 km N of Pito Solo, 15 Apr 1951 (d), L. O. Williams<br />
sute, mostly also <strong>with</strong> dense (moniliform) red- et al. 18014 (F, GH). COPAN: Rio Cop<strong>an</strong>, 2 mi E of<br />
dish-brown pluricellular hairs <strong><strong>an</strong>d</strong> sometimes Cop<strong>an</strong> ruins, 27 Aug 1975 (8), Molina et al. 30747 (F,<br />
MO). CORTES: Naciemento del Rio<br />
arachnoid<br />
Lindo, nr.<br />
hairs. Staminate<br />
Jaral, 9<br />
inflorescence Apr 1947 (6), St<strong><strong>an</strong>d</strong>ley et al. 7081 (F); Rio Londo, 8<br />
br<strong>an</strong>ched, occasionally unbr<strong>an</strong>ched; heads (1-)2- Apr 1945 (6), L. O. Williams et al. 12394 (F, GH); Rio<br />
30(-ca. 50), often partly fused, (sub)globose to Lindo, nr. Carrizal, 12 Apr 1951 (st), L. O. Williams<br />
ovoid to oblate, often more or less irregular in et al. 17817 (F), GRACIAS A Dios: Rio Plat<strong>an</strong>o, 0-4<br />
hours<br />
shape <strong><strong>an</strong>d</strong> dimensions, ca. 5-10(-20) mm upstream Ras, 23 May 1973 (9), Gentry et al.<br />
diam.; 7530 (MO, U).<br />
common peduncle 2-6 cm long, densely to NICARAGUA. BLUEFIELDS: Finca S<strong>an</strong>ta Rosa, ca.<br />
sparsely puberulous to hirtellous to shortly ve- 2.5 km ENE of Rama, 5 Apr 1966 (6), Proctor et al.<br />
101
102 Flora Neotropica<br />
27336 (F, US). CHONTALES: Nr. La Libertad, 29 May- Cricamola Valley, 17 Feb-l Mar 1928 (8), Cooper 538<br />
1 Jun 1947 (st), St<strong><strong>an</strong>d</strong>ley 8861 (F). JINOTEGA: Cord. (F, K); Ch<strong>an</strong>guinola Valley, 10 J<strong>an</strong> 1924 (2), Dunlap<br />
Isabelia, Macizos de Penias Bl<strong>an</strong>cas, 7 May 1976 (9), 299 (F, US); nr. Chiriqui Lagoon, 9 Nov 1940 (st), Von<br />
Neill 7179 (MO). MATAGALPA: Rio Yasica, 20 km E Wedel 1548 (S), 22 Nov 1940 (6), Von Wedel 1733<br />
of Matagalpa, 25 May 1977 (2), Neill 1984 (U); 10-15 (GH, US), 26 Oct 1941 (9), Von Wedel 2879 (GH, MO,<br />
km NE of Matagalpa, 16 J<strong>an</strong> 1963 (2), L. O. Williams US); Rio Cricamola, between Finca St. Louis <strong><strong>an</strong>d</strong> Konet<br />
al. 24005 (F). NUEVA SEGOVIA: Rio Solonli, 5 km kintoe, 12-16 Aug 1938 (6), Woodson et al. 1893 (F,<br />
N of Jalapa, 5 Apr 1977 (8), Neill 1653 (U). ZELAYA: MO, NA, NY). CANAL ZONE: Barro Colorado Isl<strong><strong>an</strong>d</strong>,<br />
Comarca El Hormiguero, W of Rio Uli, 6 May 1977 7 Feb 1969 (2), Croat 7768 (F, MO), 10 Feb 1969 (6),<br />
(6), Neill 1919 (U).<br />
Croat 7839 (MO), 21 Jun 1971 (2), Croat 15064 (DUKE,<br />
COSTA RICA. ALAJUELA: Between La Palma <strong><strong>an</strong>d</strong> F, GH, MO, NY), 25 Jun 1968 (2), Foster 631 (DUKE);<br />
Rio Plat<strong>an</strong>illo, 19 Feb 1964 (6), Jimenez 1750 (US); Rio Providencia, 5 Dec 1973 (st), Gentry 8707 (MO,<br />
NE-base of Volc<strong>an</strong> Arenal, 5 km W of Fortuna S<strong>an</strong> U); Gatun, 4 Mar 1860 (9), Hayes 1008 (NY); Mindi<br />
Carlos, 29 Apr 1972 (6), Lent 2520 (F, MO, U, US); Hills, 28 Mar 1956 (st), Johnston 1725 (A); Barro Col-<br />
N side of Volc<strong>an</strong> Arenal, Quebrada Guillermina, 21 orado Isl<strong><strong>an</strong>d</strong>, 19 Mar 1932 (9), Shattuck 832 (F, MO);<br />
Apr 1973 (st), Lent et al. 3388 (F). CARTAGO: Road nr. Fort Sherm<strong>an</strong>, 15 J<strong>an</strong> 1924 (st), St<strong><strong>an</strong>d</strong>ley 30924<br />
Turrialba-Moravia de Chirrip6, between Tuis <strong><strong>an</strong>d</strong> Bajo (US); Barro Colorado Isl<strong><strong>an</strong>d</strong>, 18-24 Nov 1925 (2),<br />
Pacuare, 15 Nov 1975 (8), Burger et al. 10040 (F); St<strong><strong>an</strong>d</strong>ley 41170 (US), 12 Mar 1931 (6), Wilson 127 (F,<br />
Turrialba, 10 Dec 1952 (2), Cordoba 295 (NA); Rio MO), 6 Feb 1932 (6), Woodworth et al. 445 (A, F, MO),<br />
Turrialba, Mar 1894 (2), Donnell Smith 4826 (GH, K, 22 Feb 1932 (9), Woodworth et al. 660 (A, F, MO).<br />
US, type collection of C. donnell-smithii); Atirro, Apr CHIRIQUI: La Fortuna hydroelectric project, 1200 m,<br />
1896 (2), Donnell Smith 6770 (BM, F, GH, K, US); 22 Mar 1978 (2), Hammel 2176 (U). COLON: Rio<br />
Rio Raventaz6n, nr. Turrialba, 13 Feb 1971 (6), Gilles Guache, 1 Oct 1972 (st), Gentry 6314 (MO); road to<br />
et al. 10191 (F, GH); Rio Pejibaye, between Rio Taus Portobelo, Villa Alondra, 13 Apr 1971 (6), Lao et al.<br />
<strong><strong>an</strong>d</strong> Quebrada Azul, 28 May 1972 (2), Lent 2535 (F, 21 (DUKE, MO); Rio Fat6, Jul 1911, (2), Pittier 3892<br />
MO, U); Turrialba, May 1847 (st), Oersted 14317 (C, (B, GH, NY, US, type collection of C. p<strong>an</strong>amensis).<br />
part of the type collection of Urostigma intermargin- PERLAS ARCHIPELAGO: S<strong>an</strong> Jose Isl<strong><strong>an</strong>d</strong>, 28 May 1945<br />
ale), May 1898 (2), Pittier 12288 (B, BM, GH, P, US), (2), Erl<strong>an</strong>son 240 (GH, NA, NY), 29 Mar 1945 (9),<br />
1 Sep 1920 (st), Rowle et al. 856 (NY, US); nr. Pejivalle, Johnston 580 (GH), 2 Apr 1945 (6), Johnston 606 (FI,<br />
J<strong>an</strong> 1940 (6), Skutch 4612 (A, MO, NA, US), 7-8 Feb GH, MO), (st), Johnston 608 (GH), 5 Apr 1945 (2),<br />
1926 (6), St<strong><strong>an</strong>d</strong>ley et al. 47791 (US). GUANACASTE: El Johnston 652 (BM, DUKE, FI, GH, MO, P, S), 10 Apr<br />
Arenal, 18-19 J<strong>an</strong> 1926 (2), St<strong><strong>an</strong>d</strong>ley et al. 45182 (F, 1945 (2), Johnston 678 (GH).<br />
US); nr. Tilerfn, 10-31 J<strong>an</strong> 1926 (6), St<strong><strong>an</strong>d</strong>ley et al. COLOMBIA. Without locality,<br />
44550 (US), St<strong><strong>an</strong>d</strong>ley et al. 44968 (F, US), (2), St<strong><strong>an</strong>d</strong>ley<br />
et al. 45661 (US), 3 J<strong>an</strong> 1964 (2), L. O. Williams et al.<br />
26529 (F, US). LIMON: Road Bribri-Bratsi, Rio Sixaola,<br />
12 Feb 1977 (8), Burger et al. 10474 (DUKE, F,<br />
MO); Las Diam<strong>an</strong>tas, Rubber Expt. Station, 5 May<br />
1943 (st), Dayton 3019 (F); between Portete <strong><strong>an</strong>d</strong> Moin,<br />
13 Feb 1965 (2), Jimenez 2887 (F); nr. mouth of Rio<br />
Bl<strong>an</strong>co, N of Moin, 13 Feb 1965 (9), Lent 349 (COL);<br />
ca. 4.5 km N of El Carmen, 17 Mar 1973 (st), Lent<br />
3279 (F); Zent, Oct 1904 (6 <strong><strong>an</strong>d</strong> 2), Pittier s.n. (GH,<br />
US); affluence of Rio Parismina <strong><strong>an</strong>d</strong> Rio Reventaz6n,<br />
Suerre, 3 Oct 1951 (st), Sh<strong>an</strong>k et al. 4294 (F); nr. Guapiles,<br />
12-13 Mar 1924 (6), St<strong><strong>an</strong>d</strong>ley 37259 (US); Rio<br />
Reventaz6n, Finca Montecristo, below Cairo, 18-19<br />
Feb 1926 (6), St<strong><strong>an</strong>d</strong>ley et al. 48594 (F, US); Puerto<br />
Viejo, Apr 1895 (2), Tonduz 9520 (F, US); PUNTARE-<br />
NAS: Golfito, 27 Jul 1951 (8), P. H. Allen 6280 (F, GH,<br />
US); Burica Peninsula, Quebrada Palito, 6 Mar 1973<br />
(st), Croat 22635 (MO). SAN JOSt: Guayabo, 3 Feb<br />
1908 (st), Ferry s.n. (F); Rio Vargas, Tabarcia, Mora,<br />
22 Apr 1963 (2), Jimenez 657 (COL, F); S<strong>an</strong> Isidro El<br />
General, 3 Mar 1966 (6), Molina et al. 18289 (F, US);<br />
El General, Mar 1939 (6), Skutch 4256 (A, K, MO, S,<br />
US), (2), Skutch 4262 (A, K, MO, S, US), (6), Skutch<br />
4267 (A, K, MO, S, US); Las Vueltas, Tucirrique, Feb<br />
1899 (8), Tonduz 13280 (BM, F, G, K, LE, MO, P,<br />
US).<br />
PANAMA. Without locality (Isthmus of P<strong>an</strong>ama),<br />
1859-1860 (st) Hayes 414 (NY), Hayes 867 (NY), (2),<br />
Hayes 986 (NY), (2), Hayes s.n. (BM). BocAs DEL TORO:<br />
- (2), Mutis 646<br />
(US, type collection ofC. mutisii), (a), Mutis 4558 (US),<br />
- (2), Tri<strong>an</strong>a 867 (BM, K, P), - (6), Tri<strong>an</strong>a s.n. (COL).<br />
AMAZONAS: Nr. Puerto Narifio, 24 Jul 1965 (6), Loz<strong>an</strong>o<br />
640 (COL), 28 J<strong>an</strong>-7 Feb 1969 (6), Plowm<strong>an</strong> et al.<br />
2362 (F, GH, K, NY, S), 13 Sep 1963 (2), Soejarto 856<br />
(A). ANTIOQUIA: Cocorna, Jun 1937 (st), Bro. D<strong>an</strong>iel<br />
1793 (US); Jardin, 1800 m, Dec 1941 (2), Bro. D<strong>an</strong>iel<br />
2652 (COL, F, US, type collection of C. d<strong>an</strong>ielis); Rio<br />
Porce, 1100 m, 19 Sep 1963 (2), Espinal 1314 (COL);<br />
Rio Cauca, Puerto Valdiva, 17-20 Feb 1942 (2), Metcalf&<br />
Cuatrecasas 30093 (A, MO, US, type collection<br />
of C. macarenensis var. <strong>an</strong>tioquensis). BOLiVAR: 150<br />
km N of Barr<strong>an</strong>cabermeja, Mico-Ahumado camp, ca.<br />
8?15N, 74?4'W, 25 Aug 1966 (6), De Bruijn 1123 (K,<br />
M, MO, NY, S, U, VEN). BOYACA: El Humbo, 130<br />
km N of Bogotf, ca. 1000 m, 26 Apr 1933 (6), Lawr<strong>an</strong>ce<br />
769 (A, B, F, FI, G, K, MO, S, US, type collection of<br />
C. lawr<strong>an</strong>cei). CALDAS: Salamina, Aug 1943 (?), Bro.<br />
Tomas 1923 (US). CAQUETA: Rio Caquetf, Rio Orteguaza,<br />
nr. Potosi, 7 Mar 1945 (2), Little et al. 9641<br />
(NY, US), 15 Mar 1945 (8), Little et al. 9789 (NY, US).<br />
CHOC6: Between Alto Curiche <strong><strong>an</strong>d</strong> Camp Curiche, E<br />
od Boca Curiche, 20 May 1967 (2), Duke et al. 11273<br />
(US); road Medellin-Quibd6, 6 km W of Siete, 38 km<br />
W of Bolivar, 1400 m, 6 J<strong>an</strong> 1979 (2), Gentry et al.<br />
23713 (MO, U). CUNDINAMARCA: 5 km W of El Salto<br />
de Tequendama, on road to El Colegio, 1800 m, 14<br />
Jul 1972 (8), Barclay et al. 3593 (NA); between El Salto<br />
<strong><strong>an</strong>d</strong> El Colegio, 1480 m, 10 Mar 1940 (st), Cuatrecasas<br />
8291 (F, US); nr. Albfn, Aug 1962 (2), Ferndndez-
<strong>Coussapoa</strong><br />
Perez F3 (COL); 3 km N of Tena, nr. La Laguna de<br />
Pedro Palo, 2080 m, 9 May 1952 (8), Ferndndez-Perez<br />
et al. 1458 (US); Mun. La Mesa, road to Mosquera,<br />
1800 m, 5 Aug 1963 (8), Soejarto 327 (A, K); Rio<br />
Bogota, 1600 m, 1909 (a), Uribe-Uribe 364 (COL).<br />
META: Rio Negro, 20 km W of Villaviicencio, 24 Mar<br />
1972 (Q), Barclay et al. 3294 (NA, US); nr. Villavicen-<br />
cio, 1-15 Aug 1946 (8), Duque-Jaramillo 3957 (COL),<br />
- (a), Karsten s.n. (LE); Sierra de la Macarena, Calno<br />
Entrada, 24 Dec 1949 (6), Phillipson et al. 1914 (BM,<br />
COL, US, type collection of C. macarenensis), 13 J<strong>an</strong><br />
1950 (2), Phillipson et al. 2095 (BM); nr. Villavicencio,<br />
Ll<strong>an</strong>o de S<strong>an</strong> Martin, Feb 1856 (9), Tri<strong>an</strong>a 1866 (COL,<br />
F, K, NY). VALLE: Rio S<strong>an</strong>quinini, La Laguna, 1250-<br />
1400 m, 10-20 Dec 1943 (st), Cuatrecasas 15589 (F);<br />
Gibraltar, N of Las Brisas, 2100-2200 m, 25 Oct 1946<br />
(2), Cuatrecasas 22526 (F). VAUPES: Rio Guayabero,<br />
8 Nov 1939 (a), Cuatrecasas 7510 (US, type of C.<br />
pl<strong>an</strong>itensis).<br />
VENEZUELA. Without locality, 26 Jun 1956 (2),<br />
Bernardi s.n. (8), 4 Mar 1957 (6), Bernardi s.n. (NY),<br />
1865 (a), Ernst 324 (BM), 1930-1934 (a), Vogl 1379<br />
(BR). AMAZONAS: Raudal de los Guaharibos, 24 Jul<br />
103<br />
et al. 95170 (NY, U, US, VEN); nr. C<strong>an</strong>oabo, 5 Mar<br />
1964 (8), Trujillo 6141 (MY). Departamento Federal:<br />
Nr. Caracas, 1940 (6), Bro. Elias 130 (F), J<strong>an</strong> 1931 (6),<br />
Bro. Elias 631, 1 Oct 1972 (6), Hoyos 2043 (VEN), 17<br />
Nov 1975 (2), Oliva s.n. (MY, VEN), 24 Nov 1975 (9),<br />
Oliva s.n. (VEN). FALCON: Distr. Acosta, Mun. Jacura,<br />
Cerra de la Mina, 13 Feb 1961 (st), Ruiz-Terdn 423<br />
(MER). FALCON-YACACUY: Res. For. Rio Tocuyu,<br />
Quebrada El Charalito, Aug 1970 (9), Bl<strong>an</strong>co 958 (NY,<br />
VEN). MERIDA: Merida(?), 26 Jun 1956 (9), Bernardi<br />
s.n. (MY). MIRANDA: Cerros del Bachiller, between<br />
Quebrada Corozal <strong><strong>an</strong>d</strong> Quebrada S<strong>an</strong>ta Cruz, S of S<strong>an</strong>ta<br />
Cruz, 18-19 Mar 1978 (st), Steyermarket al. 116463<br />
(MO, U, VEN). YARACUY: Bosques de Yumare, 10 Feb<br />
1959 (8), Bernardi 7017 (FI, G, MO); 35 km N of S<strong>an</strong><br />
Felipe, 19 Feb 1954 (6), Little 16245 (VEN); El Guaremal,<br />
between Valencia <strong><strong>an</strong>d</strong> S<strong>an</strong> Felipe, Feb 1919 (8),<br />
Pittier 8404 (GH, US, VEN); Rio Taria, La Ll<strong>an</strong>ada<br />
de Taria, 5 Mar 1973 (st), Romero 548 (MY). ZULIA:<br />
Quebrada Perayra, affluent of Rio Tokuku, SW of<br />
Machiques, 29 Aug 1967 (St), Steyermark 99829 (VEN).<br />
ECUADOR. AZUAY: Chacayuca, western Andes of<br />
Cuenca,<br />
1951 (st), Croizat 312 (US); Salto Salas, 20 Aug 1951<br />
(2), Croizat 541 (US). ARAGUA: Cerro de Avila, 1 May<br />
1859 (8), Crueger s.n. (K); Colonia Tovar, May 1854<br />
(2), Fendler 1237 (GH, K), Jul 1854 (a), Fendler 1237a<br />
(GH, MO); nr. Maracay, 30 J<strong>an</strong> 1857 (6), Fendler 1237<br />
(G), 1860 (9), Fendler 1237a (G), R<strong>an</strong>cho Gr<strong><strong>an</strong>d</strong>e, 6<br />
Jun 1963 (st), Montaldo 3435 (MY); Rio Chuao, El<br />
Medio, 15 Mar 1926 (8), Pittier 12124 (US, VEN); nr.<br />
Maracay, 2 Apr 1926 (9), Pittier 12142 (US, VEN), 2<br />
Apr 1926 (2), Pittier 12144 (A, G, M, MO, NY, VEN);<br />
road Maracay-Choroni, nr. El Castafio, 18 Feb 1937<br />
(a), Pittier 13985 (F, K, NY, US, VEN); Cerros de<br />
Guamitas, 1100 m, 27 Feb 1948 (a), Pittier et al. 15733<br />
(US, VEN); Parque Nacional Henri Pittier, 31 Oct 1975<br />
(2), H. Rodriguez 399 (MY); nr. El Castaino, 23 Mar<br />
1953 (2), Schnee 1337 (MY); road Maracay-Choroni,<br />
20 Dec 1948 (6), St<strong><strong>an</strong>d</strong>en 20 (MY); nr. El Castaino, 21<br />
Feb 1968 (2), Trujillo 8765 (MY); Guamitas, 12 May<br />
1938 (a), Ll. Williams 10071 (B, F, VEN), between<br />
Guamitas <strong><strong>an</strong>d</strong> El Lim6n, 28 J<strong>an</strong> 1940 (a), Ll. Williams<br />
12348 (F, S, US, VEN); nr. El Castafio, 18 Feb 1937<br />
(st), Ll. Williams 13895 (US). BARINAS: Road Barinitas-Barinas,<br />
Mar 1953 (9), Aristeguieta 1699 (MY, NY,<br />
US, VEN); nr. Barinitas, Quebrada Par<strong>an</strong>gula, Mar<br />
1953 (9), Aristeguieta 1704 (MY, NY, VEN); confluence<br />
of Rio Curbacitico <strong><strong>an</strong>d</strong> Rio Madre del Monte, 10<br />
Dec 1954 (a), Bernardi 1724 (F, FI, K, VEN); Alto del<br />
Aguada, Pedreza, 19 Feb 1955 (9), Bernardi 1980 (F,<br />
NY); Barinitas, 26 Jun 1956 (st), Bernardi 3354 (NY);<br />
between Altamira <strong><strong>an</strong>d</strong> Filo de Mertiuez, nr. Calderas,<br />
4 Mar 1957 (6), Bernardi 6348 (G); Res. For. Ticoporo,<br />
Rio Bumbum, 3 May 1964 (8), Breteler 3889 (G, MO,<br />
NY, P, S, U, US, VEN); Rio Caparo, 2-5 km above<br />
dam, 12 Mar 1980 (9), Liesner et al. 9443 (MO, U);<br />
Cuidad Bolivia, 12 J<strong>an</strong> 1968 (a), R. F. Smith 3313<br />
(VEN); Res. For. Caparo, E of El C<strong>an</strong>ton, 10 Apr 1968<br />
(2), Steyermark et al. 102083 (U, VEN), 12 Apr 1968<br />
(2), Steyermark et al. 102211 (F, MY, VEN). CARABOBO:<br />
Bejuma, 18 Feb 1954 (8), Little 16230 (VEN); Rio S<strong>an</strong><br />
Gi<strong>an</strong>, S of Borburata, 27-28 Mar 1966 (9), Steyermark<br />
- (9), Lehm<strong>an</strong>n 5606 (F, K, type collection<br />
of C. lehm<strong>an</strong>nii). ESMERALDAS: Borb6n, 11 Sep 1965<br />
(8), Little et al. 21047 (NY, Q, US); 10 km SE of Esmeraldas,<br />
29 Sep 1965 (9), Little et al. 21172 (NY, Q,<br />
US). GUAYAS: Nr. Nar<strong>an</strong>jito, 6-7 Jul 1945 (st), Camp<br />
E3626 (NY, U); Baloa, - (9), Eggers 14165 (A, L, LE,<br />
M, US). IMBABURA: Nr. Lita, 7 Feb 1981 (8), Berg<br />
1241 (U). Los Rios: Pichilingue, 18 May 1943 (2),<br />
Little 6461 (F, US); Rio Palenque Biol. Station, km 56<br />
on road Quevedo-S<strong>an</strong>to Domingo de los Colorados,<br />
11 Feb 1973 (6), Dodson 5241 (MO, QCA, SEL, U),<br />
27 Feb 1975 (6), Dodson 5778 (QCA, US), 29 Jul 1972<br />
(6), Dwyer 10305 (MO), 15 Feb 1974 (9), Gentry 9934<br />
(MO, QCA, U), 22 Feb 1974 (6), Gentry 10127 (MO,<br />
QCA, U); c<strong>an</strong>ton Vinces, hacienda S<strong>an</strong>ta Lucia, 18-<br />
28 Oct (8), Mexta 6573 (F, NA). MANABI: El Recreo<br />
Dec 1891 (8), Eggers 14165 (A, BR, L, M, US, type<br />
collection of C. eggersii), 1897 (9), Eggers 15615 (F,<br />
LE, P). MORONA-SANTIAGO: Macas, 18 Mar 1956 (2),<br />
Asplund 19835 (S); 5 km SW of Macas, 26 J<strong>an</strong> 1981<br />
(9), Berg 1222 (U); Cord. de Cucutfi, 5-10 km E of<br />
Logronio, 1200-1500 m, 7-9 Oct 1975 (6), Little et al.<br />
602 (COL, Q). NAPO: Rio Cuyabeno, ca. 0?10'S,<br />
76?55'W, 16 Feb 1980 (2), Berg & Akkerm<strong>an</strong>s 1033<br />
<strong><strong>an</strong>d</strong> 1040 (U), 19 Feb 1980 (6), Berg & Akkerm<strong>an</strong>s<br />
1062 (U); Misahualli, mouth of Rio Misahualli, 3 Mar<br />
1980 (9), Berg & Akkerm<strong>an</strong>s 1113 (U); 1 km S of Lago<br />
Agrio, 5 Nov 1974 (6), Gentry 12474 (MO, U); Rio<br />
Putumayo, just above mouth of Rio Gueppi, 19 May<br />
1978 (9), Gentry 22095 (MO, U). PICHINCHA: Road<br />
S<strong>an</strong>to Domingo de los Colorados- Quinind6 (km 170-<br />
175), 1 Sep 1949 (st), Acosta Solis 13629 (F); S<strong>an</strong>to<br />
Domingo de los Colorados, 20 Aug 1930 (st), Benoist<br />
3044 (P, S, U, US); nr. Alluriquin, bridge over Rio<br />
Toachi, 23 Feb 1981 (6), Berg 1301 (U); road S<strong>an</strong>to<br />
Domingo de los Colorados-Chona, km 5, 3 Apr 1943<br />
(8), Little 6155 (K, Q); S<strong>an</strong>to Domingo dos Colorados,<br />
21 Apr 1943 (9), Little 6328 (F, K, NY, Q, US); nr.<br />
Alluriquin, at confluence of Rio Alluriquin <strong><strong>an</strong>d</strong> Rio<br />
Toachi, 14 Mar 1967 (8), Sparre 14822 (S). ZAMORA-<br />
CHINCHIPE: Quebrada Las Pavas Pavas, 14 km from
104 Flora Neotropica<br />
Loja, 1800 m, 21 Aug 1975 (st), Sam<strong>an</strong>iego et al. 92<br />
(COL, Q).<br />
PERU. Without locality, - (9), Dombey s.n. (F, P,<br />
US), 1835 (6), Matthews 2068 (K, OXF, U), 1829 (6),<br />
Poeppig s.n. (B, OXF, type collection). AMAZONAS: Rio<br />
Cenepa, Aintami Creek, 24 Nov 1972 (6), Berlin 372<br />
(MO, U); Pongo de M<strong>an</strong>seriche, 1924 (9), Tessm<strong>an</strong>n<br />
4696 (B, NY); Rio S<strong>an</strong>tiago, Caterpiza, 20 Nov 1979<br />
(8), Tunqui 103 (U). Hu<strong>an</strong>uco: Pampayacu, nr. Hu<strong>an</strong>aco,<br />
20 J<strong>an</strong> 1927 (6), K<strong>an</strong>ehira 99 (GH); Prov. Leoncio<br />
Prado, Tingo Maria, 7 Dec 1981 (6), Plowm<strong>an</strong> et al.<br />
11180 (U); Macora, - (9), Ruiz & Pavon s.n. (or 3) (B,<br />
FI, type collection of C. vellera), - (5), Ruiz & Pavon<br />
s.n. (BR, G, OXF, US); Huamalies, between Monzon<br />
<strong><strong>an</strong>d</strong> Rio Huallaga, - (9), Weberbauer 3702 (G, type<br />
collection of C. st<strong><strong>an</strong>d</strong>leyi). JUNIN: La Merced, Puente<br />
Quimiri, 24 Oct 1950 (6), Nunez 2928 (US); S<strong>an</strong>abeni,<br />
4 Sep 1960 (8), Woytkowski 5956 (MO, U). LORETO:<br />
Prov. Maynas, Rio Itaya, above Puerto Belen, 16 Nov<br />
1978 (9), Diaz et al. 618 (U); Prov. Maynas, Rio Itaya,<br />
nr. Palo Seco, ca. 40 km above Iquitos, 20 Mar 1977<br />
(6), Gentry et al. 18458 (MO, U); Prov. Requena, Rio<br />
Tapiche, tributary of Rio Ucayali, 1 hour above Requena,<br />
8 Dec 1972 (,), Gentry et al. 21291 (MO, U);<br />
lower Rio Huallaga, Y<strong>an</strong>6n, 5-11 Sep 1929 (9), Killip<br />
et al. 29249 (NY, US, type collection of C. gr<strong><strong>an</strong>d</strong>iceps);<br />
Rio Marafion, lower Rio Ampiyacu, 6 Mar 1977 (6),<br />
Pr<strong>an</strong>ce et al. 24687 (U); Prov. Maynas, Rio N<strong>an</strong>ay,<br />
Mish<strong>an</strong>a, 1978 (6), Ramirez 1084 (U); Rio Ucuyali,<br />
Yarinacocha, between 10?S <strong><strong>an</strong>d</strong> mouth, 1923 (6), Tessm<strong>an</strong>n<br />
3461 (B, G, NY, S); lower Rio Huallaga, Yurimaguas,<br />
26 Oct 1929 (6), LI. Williams 4179 (F); Puerto<br />
Arturo, Yurimaguas, 22 Nov 1929 (2), LI. Williams<br />
5349 (B, F). (LORETO?): Rio Zungarococha, 28 J<strong>an</strong><br />
1978 (6), Ayala et al. 1436 (MO, U). MADRE DE Dios:<br />
Prov. Tambopata, affluence of Rio Tambopata <strong><strong>an</strong>d</strong> Rio<br />
Le Torre, Tambopata Nat. Res., ca. 30 km SSW of<br />
Puerto Maldonado, 16 May 1980 (9), Balbour 5372<br />
(U); Parque Nacional del M<strong>an</strong>u, Rio M<strong>an</strong>u, Pakitza<br />
station, 9 Nov 1980 (6), Foster 5721 (F).<br />
BRAZIL. Without locality, - (9), Martius s.n. (M,<br />
type collection of C. marti<strong>an</strong>a), - (6), Martius s.n. (BR,<br />
M, U, type collection of C. subinc<strong>an</strong>a). ACRE: Rio Acre,<br />
Itui, 4 Nov 1923 (9), Kuhlm<strong>an</strong>n 759 (RB, U); Tarauaca,<br />
1-2 km E of Rio Tarauaca, 24 Sep 1968 (2), Pr<strong>an</strong>ce et<br />
al. 7530 (F, GH, K, NY, P, R, S, U, US); Rio Jurui-<br />
Mirim, Aug 1901 (6), Ule 5717 (B, G, K, L, MG).<br />
AMAZONAS: Rio Solimoes, Mun. Tefe, Paleta, 21 May<br />
1933 (6), Krukoff4518 (A, BM, F, G, K, M, MO, NY,<br />
S, U, US, type collection of C. subcrenata); nr. mouth<br />
of Rio Embira, tributary of Rio Tarauaca, 26 Jun 1933<br />
(6), Krukoff4994 (A, G, K, NY, U, US, type collection<br />
of C. embir<strong>an</strong>a); Mun. Humaita, Tres Casas, 14 Sep-<br />
11 Oct 1934 (9), Krukoff 6165 (A, F, FI, K, MO, NY,<br />
S, U, US), Krukoff 6524 (A, BM, F, G, K, LE, MO,<br />
NY, S, U, US, type collection of C. ar<strong>an</strong>eosa); Rio<br />
Madeira, Boa Hora, 4 Sep 1923 (2), Kuhlm<strong>an</strong>n 368<br />
(RB); Rio Japura, - K, NY, P, R, S, U, US); Rio Purus, between Lago Quati<br />
<strong><strong>an</strong>d</strong> Lago Arima, 20 Jun 1971 (6), Pr<strong>an</strong>ce et al. 13438<br />
(K, M, MO, NY, S, U, US); M<strong>an</strong>aquiri, Jun 1851 (8),<br />
Spruce 1608 (BM, BR, C, G, GH, K, LE, NY, P, OXF).<br />
MATO GROSSO: Rio Aripu<strong>an</strong>a, nr. Humboldt Center,<br />
ca. 10?12'S, 59021'W, 12 Oct 1973 (9), Berget al. P18455<br />
(F, K, MO, NY, S, U, US), 22 Oct 1973 (6), Berg et<br />
al. P19841 (F, K, MO, NY, P, S, U, US); Sarar6, 14?<br />
50'S, 59'45'10"W, 4 Aug 1972 (6), Pires et al. 16461<br />
(U). PARA: Alto Tapaj6s, Rio Curucii 1-5 km above<br />
Missao Cururui, 7?35'S, 57?31'W, 17 Feb 1974 (9), Anderson<br />
11033 (MO, U); Gleba Bacaja, below mouth of<br />
Rio Bacaja, 22 Nov 1980 (2), Pr<strong>an</strong>ce et al. 26402 (U).<br />
RORAIMA: Rio Mucajai, Posto 17 Mar 1971 (6), Pr<strong>an</strong>ce<br />
et al. 11066 (K, M, NY, P, S, U).<br />
BOLIVIA. SANTA CRUZ: Prov. Ichilo, Rio Vibora,<br />
5 Oct 1926 (2), Steinbach 7567 (A, F, GH, K, MO, S,<br />
U, type collection of C. bolivi<strong>an</strong>a).<br />
<strong>Coussapoa</strong> villosa is a widespread species (like<br />
C. asperifolia) <strong><strong>an</strong>d</strong> is extremely variable in m<strong>an</strong>y<br />
characters. Although the occurrence of some of<br />
the features is associated <strong>with</strong> certain parts of the<br />
species area, e.g., br<strong>an</strong>ched pistillate inflorescences<br />
in Central America <strong><strong>an</strong>d</strong> brown arachnoid<br />
hairs in Colombia <strong><strong>an</strong>d</strong> adjacent parts of Ecuador,<br />
infraspecific taxa c<strong>an</strong>not readily be recognized.<br />
Some specimens from the periphery of the species<br />
area <strong><strong>an</strong>d</strong>/or <strong>with</strong> features near the extreme of the<br />
variation must be mentioned. Barclay et al. 3593<br />
(Colombia) has staminate inflorescences <strong>with</strong> extremely<br />
numerous (ca. 50) heads; Gentry et al.<br />
23713 (Colombia) has pistillate inflorescences<br />
<strong>with</strong> very short (less th<strong>an</strong> 2 cm long) peduncles<br />
(in this respect it forms a tr<strong>an</strong>sition to C. duquei);<br />
Pr<strong>an</strong>ce et al. 26402 (Brazil, Pari) is peculiar in<br />
that the head of the pistillate inflorescence is lobed<br />
<strong><strong>an</strong>d</strong> obliquely attached to the peduncle. This last<br />
collection matches some more or less aberr<strong>an</strong>t<br />
collection of C. tessm<strong>an</strong>nii (see p. 95) in the dense<br />
covering of dark brown moniliform pluricellular<br />
(=gr<strong>an</strong>ular) hairs on the peduncle, leafy twig, <strong><strong>an</strong>d</strong><br />
petiole.<br />
Typification of C. villosa: It is unknown<br />
whether the destroyed type material in W consisted<br />
of a pistillate <strong><strong>an</strong>d</strong> a staminate specimen as<br />
the drawing in Poeppig & Endlicher (1838) suggests.<br />
The specimens in B <strong><strong>an</strong>d</strong> OXF which could<br />
replace the holotype material are staminate.<br />
(6), Martius s.n. (G); Lago de Ma- Typification of C. eggersii: Eggers made sevnacapuru,<br />
10 Oct 1972 (2), 0. Pires 248 (U); Tefe, 23 eral collections under number<br />
Jul 1972<br />
14165-<br />
(9), PLK& Urb<strong>an</strong>o 12296<br />
staminate<br />
(INPA); road Boca<br />
do Acre-Rio Br<strong>an</strong>co, km 1-4, 19 Sep 1966 (6), Pr<strong>an</strong>ce <strong><strong>an</strong>d</strong> pistillate ones from different localities in Ecet<br />
al. 2306 (F, GH, K, P, R, S, U); Rio Demini, To- uador (El Recreo <strong><strong>an</strong>d</strong> Baloa)-at several dates.<br />
totobi, 28 Feb 1969 (2), Pr<strong>an</strong>ce et al. 10348 (F, GH, These collections have been mixed up. The spec-
<strong>Coussapoa</strong><br />
imen in F on which C. eggersii is based is staminate<br />
<strong><strong>an</strong>d</strong> possibly from El Recreo. In the original<br />
description the specimen is wrongly described<br />
as pistillate.<br />
43. <strong>Coussapoa</strong> viridifolia Cuatrecasas, Fieldi<strong>an</strong>a<br />
Bot. 28(1): 213. 1956. Type. Venezuela. Bolivar:<br />
0-8 km N of S<strong>an</strong>ta Teresita de Kav<strong>an</strong>ayen,<br />
23 Nov 1944 (6), Steyermark 60456 (holotype,<br />
F; isotype, VEN). Fig. 55.<br />
<strong>Coussapoa</strong> viridifolia Cuatrecasas var. tenuifolia Cuatrecasas,<br />
Fieldi<strong>an</strong>a Bot. 28(1): 214. 1956. Type. Venezuela.<br />
Bolivar: Cerro Duida, C<strong>an</strong>io Negro, 25-26<br />
Aug 1944 (9), Steyermark 57955 (holotype, F; isotype,<br />
VEN).<br />
Tree or shrub, often hemi-epiphytic, up to 20<br />
m tall. Leafy twigs 2-5 mm thick, appressedpuberulous<br />
to substrigose to hirtellous, <strong>with</strong> hairs<br />
differing distinctly in length. Lamina coriaceous<br />
to subcoriaceous, obovate to elliptic to oblong,<br />
2.5-10 x 1-6 cm, apex obtuse to shortly acuminate<br />
or rounded to emarginate, base obtuse to<br />
acute to cuneate to rounded, margin entire to<br />
subcrenate; upper surface glabrous, lower surface<br />
sparsely puberulous to substrigose on the main<br />
veins, the hairs differing distinctly in length; lateral<br />
veins 4-7 pairs, basal pair unbr<strong>an</strong>ched or<br />
faintly br<strong>an</strong>ched reaching the margin at or above<br />
the middle of the lamina, sometimes some of the<br />
other lateral veins poorly br<strong>an</strong>ched (furcate); intercostal<br />
venation pl<strong>an</strong>e to more or less prominent;<br />
petiole 1-3 cm long, sparsely (to rather<br />
densely) appressed-puberulous to substrigose,<br />
<strong>with</strong> hairs differing distinctly in length; stipules<br />
0.5-1 cm long, densely appressed-puberulous to<br />
subsericeous; terminal buds slender. Staminate<br />
inflorescences br<strong>an</strong>ched; heads 3-14, globose, ca.<br />
2-4 mm diam.; common peduncle 1-3 cm long,<br />
sparsely puberulous; peri<strong>an</strong>th ca. 1 mm high,<br />
glabrous; stamens two, just exceeding the peri<strong>an</strong>th.<br />
Pistillate inflorescences unbr<strong>an</strong>ched or<br />
poorly br<strong>an</strong>ched; heads 1-3, globose, ca. 3-6 mm,<br />
in fruit up to 8 mm diam.; (common) peduncle<br />
0.5-2.5 cm long, sparsely (to rather densely) puberulous;<br />
peri<strong>an</strong>th ca. 1 mm high, glabrous. Interfloral<br />
bracts often only a few (sometimes lacking?),<br />
small, subspathulate to subpeltate, minutely<br />
puberulous at the apex.<br />
Distribution (Fig. 9). Eastern Gui<strong>an</strong>a (=Highl<strong><strong>an</strong>d</strong>)<br />
region, in Venezuela (Bolivar <strong><strong>an</strong>d</strong> Ama-<br />
105<br />
zonas), Guy<strong>an</strong>a (Upper Mazaruni), <strong><strong>an</strong>d</strong> Brazil<br />
(Roraima); in rain forest <strong><strong>an</strong>d</strong> sav<strong>an</strong>nas, up to<br />
1250 m.<br />
Specimens studied. VENEZUELA. AMAZONAS:<br />
"Alto Orinoco," 1951 (2), Croizat 959 (F, US); Cerro<br />
Duida, C<strong>an</strong>io Negro, 25-26 Aug 1944 (2), Steyermark<br />
57955 (F, VEN, type collection of C. viridifolia var.<br />
tenuifolia). BOLIVAR: Region of Rio Icabaru <strong><strong>an</strong>d</strong> Rio<br />
Hacha, 2 J<strong>an</strong> 1956 (2), Bernardi 2744 (F, FI, NY, VEN);<br />
between S<strong>an</strong>ta Teresita de Kav<strong>an</strong>ayen <strong><strong>an</strong>d</strong> ravine ca.<br />
8 km NW of it, 23 Nov 1944 (8), Steyermark 60451<br />
(F, VEN, type collection).<br />
GUYANA. Kamar<strong>an</strong>g River, below mouth of Utschi<br />
Creek, 30 Oct 1960 (8), Tillett et al. 45869 (F, NY, U).<br />
BRAZIL. AMAZONAS: Serra de Neblina, Rio Cauaburi-Rio<br />
Maturaca, 600-1300 m, 16 Dec 1965 (2),<br />
N. T. Silva et al. 60670 (U). RORAIMA: Serra Surucucu,<br />
2 Feb 1975 (9), Ribeiro 655 (IAN), 1 Feb 1975 (2), Rosa<br />
339 (IAN).<br />
This species appears to be closely related to C.<br />
latifolia. The two are allopatric <strong><strong>an</strong>d</strong> apparently<br />
ecologically distinct, but morphologically very<br />
close. The two taxa show (minor) differences in<br />
the leaf venation, the length of the stipules, <strong><strong>an</strong>d</strong><br />
the indumentum of the leaves. The morphological<br />
differences suggest the possibility of distinction<br />
at the subspecific level.<br />
The following three species have been discov-<br />
ered <strong><strong>an</strong>d</strong> described after submission of the post-<br />
review m<strong>an</strong>uscript of the revision <strong><strong>an</strong>d</strong> are, there-<br />
fore, not placed in the alphabetical sequence.<br />
44. <strong>Coussapoa</strong> fulvescens C. C. Berg, Brittonia<br />
42: 59-65. 1990. Type. Colombia. Choc6: Rd.<br />
Quibdo-Bolivar, ca. km 50, 9 Jul 1986 (2),<br />
Berg 1546 (holotype, COL; isotypes, AAU,<br />
BG, F, HUA, MO, NY). Fig. 56.<br />
Tall, hemi-epiphytic tree. Leafy twigs ca. 10<br />
mm thick, white-appressed-puberulous, partly<br />
also brownish-hirsute, lenticels conspicuous.<br />
Lamina coriaceous, oblong to subobovate, 18-<br />
26 x 9-15 cm, apex short-acuminate to obtuse,<br />
base obtuse, margin entire; upper surface glabrous,<br />
lower surface sparsely appressed-puberulous;<br />
lateral veins 7-9 pairs, basal pair unbr<strong>an</strong>ched<br />
(or very faintly br<strong>an</strong>ched) reaching the<br />
margin at or just below the middle of the lamina,<br />
usually the second <strong><strong>an</strong>d</strong>/or third lateral veins (from<br />
the base) <strong>with</strong> one or two strong br<strong>an</strong>ches, tertiary<br />
venation distinct, slightly prominent; petiole<br />
5-10 cm long, appressed-puberulous; stipules 1.5-
<strong>Coussapoa</strong> 107<br />
FXG. 56. <strong>Coussapoa</strong>fillvescens: 1, leafy twig <strong>with</strong> Distillate in~lorescences (Berg 1546), x0.75.<br />
r;:~~~~~~~~~~~~~~~~~~'<br />
":~~~~~~~~~~~~~~~.:?<br />
1~~?. ~ .<br />
?i I?:i,<br />
?~~~~~~~~~~~~~~~~~~~~~~'<br />
"'<br />
FIG. 56. Coussaoafulvescens: 1 leafy twig wih pistillate inflorescence (Beg 1546), x0.75
108 Flora Neotropica<br />
2 cm long, yellowish- to brown-(sub)hirsute. Pistillate<br />
inflorescences br<strong>an</strong>ched; heads 10-20,<br />
sometimes partly fused, (sub)globose, 3-5 cm in<br />
diam.; common peduncle 3-6 cm long, ca. 2 mm<br />
thick, appressed-puberulous <strong><strong>an</strong>d</strong> partly also<br />
brownish-hirtellous; peri<strong>an</strong>th ca. 2 mm high, glabrous,<br />
+ distinctly ribbed. Interfloral bracts subulate<br />
to narrowly spathulate, <strong>with</strong> a few hairs at<br />
the apex.<br />
Distribution (Fig. 9). Colombia (Choc6), only<br />
known from the type locality, in lowl<strong><strong>an</strong>d</strong> secondary<br />
growth.<br />
Tree, 18 m tall. Leafy twigs 5-12 mm thick,<br />
yellowish- to whitish-hirtellous. Lamina coriaceous,<br />
ovate to elliptic, 10-21 x 8-16 cm, apex<br />
shortly acuminate (to acute), base rounded to<br />
subcordate, margin coarsely crenate to subentire;<br />
upper surface <strong>with</strong> sparse hairs on the midrib,<br />
lower surface hirtellous on the veins, in the reticulum<br />
minutely puberulous; lateral veins 12-<br />
14 pairs, basal pair br<strong>an</strong>ched, reaching the margin<br />
below the middle of the lamina, tertiary <strong><strong>an</strong>d</strong><br />
quarterary venation <strong><strong>an</strong>d</strong> reticulum prominent;<br />
petiole 4-6 cm long, 2-4 mm thick, hirtellous,<br />
hairs not distinctly different in length; stipules<br />
4-12 cm long, densely yellowish-puberulous <strong><strong>an</strong>d</strong><br />
partly also hirsute. Pistillate inflorescences<br />
br<strong>an</strong>ched; heads 4-5, globose (to subdiscoid), in<br />
fruit 0.7-1.3(-1.5) mm in diam.; common peduncle<br />
2.5-3 cm long, ca 2-2.5 mm thick, hirtellous;<br />
peri<strong>an</strong>th ca. 1.5 mm high, at the apex<br />
puberulous, in fruit red. Interfloral bracts absent.<br />
Distribution (Fig. 9). Colombia (Tolima), only<br />
known from the type locality.<br />
Specimens studied. COLOMBIA. TOLIMA: 3.4 km<br />
from Las Juntas, 1940 m, 9 Apr 1984 (v), Escobar et<br />
al. 4219 (HUA, type collection).<br />
This species resembles C. nitida, known from<br />
the Amazon Basin <strong><strong>an</strong>d</strong> mostly occurring in riv-<br />
erside vegetation. It differs clearly from the latter<br />
in the prominent tertiary <strong><strong>an</strong>d</strong> quartemary ve-<br />
nation at the lower surface of the lamina <strong><strong>an</strong>d</strong> in<br />
the red-colored fruiting peri<strong>an</strong>th.<br />
46. <strong>Coussapoa</strong> valaria C. C. Berg, Brittonia 42:<br />
59-65.1990. Type. Colombia. Valle: Bajo Cali-<br />
ma, rd. to Dindo, 25 Sep 1986 (2), Monsalve<br />
B. 1176 (holotype, CUVC; isotypes, BG,<br />
MO). Fig. 58.<br />
Specimens studied. COLOMBIA. CHOC6: Rd.<br />
Quibdo-Bolivar, ca. km 50, 9 Jul 1986 (9), Berg 1546<br />
(AAU, BG, COL, F, HUA, MO, NY, type collection).<br />
This species is closely related to C. herthae, Tree, hemi-epiphytic. Leafy twigs 5-10 mm<br />
known from western Ecuador <strong><strong>an</strong>d</strong> Narifio (Co- thick, brown strigose, at least at the scars of the<br />
lombia). It differs from the latter, e.g., in the stipules, glabrescent. Lamina coriaceous, ellipindument<br />
of the leaves, the venation, the shorter tic, 15-17 x 11-14 cm, apex rounded to shortly<br />
stipules, the more numerous <strong><strong>an</strong>d</strong> smaller flower acuminate, base rounded to subcordate, margin<br />
heads of the pistillate inflorescence. The two taxa subentire, at the base revolute; upper surface glamight<br />
prove to be distinct only at the subspecific brous, lower surface strigillose at the base of the<br />
level. C. fulvescens may also be related to C. midrib, on the midrib <strong><strong>an</strong>d</strong> lateral veins <strong>with</strong> a<br />
echinata.<br />
soon disappearing white arachnoid indument;<br />
lateral veins 8-10 pairs, basal pair + br<strong>an</strong>ched,<br />
45. <strong>Coussapoa</strong> tolimensis C. C. Berg, Brittonia reaching the margin at or just below the middle<br />
42: 59-65. 1990. Type. Colombia. Tolima: 3.4 of the lamina, tertiary venation almost pl<strong>an</strong>e;<br />
km from Las Juntas, 1940 m, 9 Apr 1984 (9), petiole 5-6.5 cm long, glabrous or brownish stri-<br />
Escobar et al. 4219 (holotype, HUA). gose, stipules 2-3 cm long, rather densely whit-<br />
Fig. 57. ish appressed-puberulous to -strigillose, or also<br />
<strong>with</strong> white arachnoid hairs. Pistillate inflorescences<br />
br<strong>an</strong>ched; heads 3-9, 3-4 (in fruit 7-9)<br />
mm diam.; common peduncle 2-4 cm long,<br />
densely minutely puberulous; peri<strong>an</strong>th ca. 1.5<br />
mm high, minutely puberulous around the aperture.<br />
Interfloral bracts absent.<br />
Distribution (Fig. 9). Colombia (Valle), in lowl<strong><strong>an</strong>d</strong><br />
forest.<br />
Specimens studied. COLOMBIA. VALLE: Bajo Calima,<br />
rd. to Ju<strong>an</strong>chaco Palmeras, 10 Jul 1984 (9), Gentry<br />
et al. 47838 (BG); Bajo Calima, rd. to Dindo, 25 Sep<br />
1986 (9), Monsalve B. 1176 (BG, MO, type collection).<br />
This species resembles both C. ovalifolia <strong><strong>an</strong>d</strong><br />
C. parviceps. It differs from these species in, e.g.,<br />
the venation <strong><strong>an</strong>d</strong> the number <strong><strong>an</strong>d</strong> dimensions of<br />
the flower heads of the pistillate inflorescences.<br />
Unnamed Collections<br />
Gentry & Mori 14011 from P<strong>an</strong>ama: Discussed<br />
under C. ovalifolia.<br />
Huashikat 1585 from Peru: Discussed under C.<br />
cupularis.
<strong>Coussapoa</strong><br />
FIG. 57. <strong>Coussapoa</strong> tolimensis: 1, leafy twig <strong>with</strong> pistillate inflorescences<br />
(Albert de Escobar et al. 4219), x0.6.<br />
Names Not Included<br />
<strong>Coussapoa</strong> dealbata Andre, Ill. Hort. 17: 17.<br />
1870--nomen nudum.<br />
<strong>Coussapoa</strong> dolich<strong><strong>an</strong>d</strong>ra Cuatrecasas Caldasia 7:<br />
287. 1956-based on collection Cuatrecasas<br />
17458 (Colombia, Valle: Rio Cajambre, 5-15<br />
May 1944 (holotype <strong><strong>an</strong>d</strong> isotype, F)-consists<br />
109<br />
of discord<strong>an</strong>t elements: staminate inflores-<br />
cences of a <strong>Coussapoa</strong> species <strong><strong>an</strong>d</strong> leafy twigs<br />
of some species not belonging to the Urticales,<br />
but possibly to the Lauraceae. The latter parts<br />
constitute the lectotype, chosen here.<br />
<strong>Coussapoa</strong> emarginata Killip in Macbride, Publ.<br />
Field Mus. Bot. 13(2.2): 296. 1937 = Pourou-<br />
ma minor Benoist.
110 Flora Neotropica<br />
FIG. 58. <strong>Coussapoa</strong> valaria: 1, leafy twig <strong>with</strong> pistillate inflorescences<br />
(Gentry et al 47838), x0.55.<br />
<strong>Coussapoa</strong> krukovii St<strong><strong>an</strong>d</strong>ley, Publ. Field Mus.<br />
Bot. 17:1937 = <strong>Pourouma</strong> tomentosa Miquel.<br />
<strong>Coussapoa</strong> laevigata Poeppig & Endlicher, Nov.<br />
gen. sp. pl. 2: 33. 1838-based on material<br />
from Brazil (Amazonas: Tefe). The holotype<br />
in W has been destroyed. Isotypes have not<br />
been traced. The identity is uncertain. The<br />
name possibly applies to C. villosa.<br />
<strong>Coussapoa</strong> leopoldii Micheli, Rev. Hort. (1894):<br />
251. 1894-nomen nudum.<br />
<strong>Coussapoa</strong> obovata Warburg ex Glaziou, Bull.<br />
Soc. Bot. Fr<strong>an</strong>ce 59 (Mem. 3g): 645. 1913nomen<br />
nudum.<br />
<strong>Coussapoa</strong> pittieri St<strong><strong>an</strong>d</strong>ley in Hoyos, Los Ar-<br />
= Ficus intermarginalis (Liebm<strong>an</strong>n) Miquel,<br />
Ann Bot. Lugd. Bat. 3:297. 1867; see St<strong><strong>an</strong>d</strong>ley,<br />
Publ. Field Mus. Bot. 18(1): 383. 1937.<br />
POUROUMA<br />
Introduction<br />
<strong>Pourouma</strong> is the smallest of the three Neotropical<br />
genera of the <strong>Cecropiaceae</strong>. It is a welldefined<br />
genus of small to medium-sized trees of<br />
the rain forest areas of South <strong><strong>an</strong>d</strong> Central America,<br />
<strong><strong>an</strong>d</strong> has not drawn much attention, except<br />
that the fruits of P. cecropiifolia are suitable for<br />
boles de Caracas, ed. 2 (Monogr. 24. Soc. Cienc. hum<strong>an</strong> consumption. The genus is rather uni-<br />
Nat. La Salle, Caracas, Venezuela) p. 242, tab. form in its ecology. Besides a number of clear-<br />
(of C. villosa)-nomen nudum.<br />
cut species, mostly of limited distribution, it<br />
<strong>Coussapoa</strong> plicata Warburg ex Ule, Bot. Jahrb. comprises several more wide-spread <strong><strong>an</strong>d</strong> rather<br />
40: 143. 1907- nomen nudum.<br />
complex groups of taxa <strong>with</strong> a confusing varia-<br />
<strong>Coussapoa</strong> rekoi St<strong><strong>an</strong>d</strong>ley, Contr. U.S. Natl. Herb tion in leaf shapes <strong><strong>an</strong>d</strong> indument, <strong><strong>an</strong>d</strong> to which<br />
20: 211. 1919 = Poulsenia armata (Miquel) the following quotation applies: "The species of<br />
St<strong><strong>an</strong>d</strong>ley: see Mildbraed, Notizbl. Bot. Gart. <strong>Pourouma</strong> are perhaps the most ambiguous <strong><strong>an</strong>d</strong><br />
Berlin 10: 413. 1928; St<strong><strong>an</strong>d</strong>ley, Trop. Woods vexatious of the Americ<strong>an</strong> Moraceae, a dubious<br />
33: 4. 1933.<br />
distinction" (Woodson & Schery, 1960:166). The<br />
Urostigma intermarginale Liebm<strong>an</strong>n, Kongel. lack of sufficient field work must be deemed a<br />
D<strong>an</strong>ske Vidensk.-Selsk. Skr. Ser. 5,2:328.1851 great disadv<strong>an</strong>tage in the preparation of the pres-
Morphology<br />
ent revision, especially <strong>with</strong> regard to the vari-<br />
ation in the leaves.<br />
111<br />
They are often very dense <strong><strong>an</strong>d</strong> form a powdery<br />
layer in dried material. The denseness of these<br />
hairs is often variable.<br />
Taxonomic History<br />
2. Unicellular arachnoid (cobwebby) hairs. These<br />
very thin, crinkled, interwoven hairs of dif-<br />
The genus <strong>Pourouma</strong> was established by Au- ferent length are mostly white, sometimes<br />
blet (1775) who described one species, P. gui- brownish (in P. ferruginea). Short arachnoid<br />
<strong>an</strong>ensis. Names were added by Martius (in Spix hairs are always found in the areoles at the<br />
<strong><strong>an</strong>d</strong> Martius, 1831, 1843), Tr6cul (1847), Klotzsch lower leaf surface. Longer arachnoid hairs c<strong>an</strong><br />
(1847), <strong><strong>an</strong>d</strong> Miquel (1853), <strong><strong>an</strong>d</strong> in this century be found on the smaller veins, <strong><strong>an</strong>d</strong> in some<br />
by Rusby (1910), Benoist (1922, 1924), Mild- species (e.g., P. ferruginea <strong><strong>an</strong>d</strong> P. tomentosa)<br />
braed (1927, 1928), Macbride (1930), Ducke also on the main veins, from which they often<br />
(1932a, 1932b, 1947), St<strong><strong>an</strong>d</strong>ley (1937a, 1937b, disappear <strong>with</strong> age. Long arachnoid hairs also<br />
1939), Cuatrecasas <strong><strong>an</strong>d</strong> St<strong><strong>an</strong>d</strong>ley (in Cuatrecasas, occur on the petioles, stipules, leafy twigs <strong><strong>an</strong>d</strong>/<br />
1951, 1956b), Cuatrecasas (1954, 1956a, 1956b, or the inflorescences in some species.<br />
1956c, 1967), Woodson <strong><strong>an</strong>d</strong> Schery (1960), <strong><strong>an</strong>d</strong> 3. Unicellular non-arachnoid, thicker <strong><strong>an</strong>d</strong> mostfinally<br />
by Berg <strong><strong>an</strong>d</strong> Kooy (1982), Berg <strong><strong>an</strong>d</strong> v<strong>an</strong> ly more or less straight hairs. Short hairs are<br />
Heusden (1988), <strong><strong>an</strong>d</strong> Berg (1989, 1990).<br />
usually whitish, the longer ones mostly yel-<br />
Since the comprehensive treatments of Pou- low(ish). They c<strong>an</strong> be found on most parts of<br />
rouma by Trecul (1847) <strong><strong>an</strong>d</strong> Miquel (1853) the the pl<strong>an</strong>t. In some species, forms <strong>with</strong> only<br />
genus has only been treated for regional floras: short, whitish hairs occur beside forms pos-<br />
Peru (Macbride, 1937), P<strong>an</strong>ama (Woodson & sessing longer yellowish hairs. In several<br />
Schery, 1960), Costa Rica (Burger, 1977), <strong><strong>an</strong>d</strong> species, all or some specimens have short,<br />
Surinam (Berg, 1975).<br />
rigid hairs on the upper or sometimes on the<br />
lower leaf surface.<br />
Morphology<br />
Leaves. -The leaves are medium-sized to<br />
Habit.-<strong>Pourouma</strong> species are usually small<br />
large,<br />
entire or<br />
to medium-sized trees, often <strong>with</strong> stilt-roots from palmately incised. In taxa <strong>with</strong> incised<br />
leaves the first formed leaves are<br />
the lower part of the trunk, as in most of the<br />
entire; subseother<br />
terrestrial, non-climbing species of Cecro- quently leaves sooner or later, frequently or alpiaceae.<br />
In some species (e.g., P. ways, become incised. In adult specimens, eshirsutipetiolata)<br />
the trunk may become buttressed. The architecpecially<br />
on flowering br<strong>an</strong>ches, entire leaves c<strong>an</strong><br />
ture of the trees of P. minor is that of the model<br />
again be found, although different in shape <strong><strong>an</strong>d</strong><br />
texture from the leaves in the<br />
of Aubreville (F. Halle, pers. comm.). The<br />
juvenile stage. Entire<br />
<strong><strong>an</strong>d</strong> incised leaves<br />
br<strong>an</strong>ches may be thick (as in species <strong>with</strong> incised<br />
may occur on the same<br />
br<strong>an</strong>chlet. Leaves similar to those formed in the<br />
leaves) or rather thin (as in species <strong>with</strong> mediumsized,<br />
entire<br />
subjuvenile stage often appear again on sucker<br />
leaves).<br />
shoots of adult<br />
The wood (<strong><strong>an</strong>d</strong> other vegetative parts) of some<br />
specimens <strong>with</strong> entire leaves.<br />
Leaves are<br />
species (e.g., P. myrmecophila) give out a<br />
always incised in adult specimens of<br />
pecu- <strong>Pourouma</strong><br />
liar odor, indicated as cecropiifolia, <strong><strong>an</strong>d</strong> probably also in P.<br />
'spearmint', 'winter-green',<br />
'balsam', or 'bengie'. For other species (e.g., P.<br />
myrmecophila, P. oraria, P. stipulacea, <strong><strong>an</strong>d</strong> P.<br />
villosa. Incised leaves<br />
qui<strong>an</strong>ensis) a 'balsam' odor is reported for the<br />
appear to occur const<strong>an</strong>tly<br />
in some<br />
fruits.<br />
subspecies of species in which entire<br />
leaves are common. In<br />
When cut the pl<strong>an</strong>t parts exude a species in which<br />
watery sap<br />
only<br />
entire leaves are<br />
which is clear or sometimes reddish (e.g., in P.<br />
found, the leaves in the juvenile<br />
<strong><strong>an</strong>d</strong><br />
bicolor); after exposure to the air it turns black. subjuvenile stages are basically similar to<br />
those of adult<br />
Indumentum. -Three types of trichomes c<strong>an</strong><br />
specimens. A number of species<br />
be<br />
appear to be const<strong>an</strong>tly entire-leaved (e.g., P. borecognized:<br />
livarensis, P. minor, P. ovata, <strong><strong>an</strong>d</strong> P. phaeotri-<br />
1. Pluricellular hairs, often more or less globose, cha). In this group of species the leaves may vary<br />
pale brown to dark brown or purple. These from predomin<strong>an</strong>tly ovate to elliptic to oblong,<br />
hairs occur on most young parts of the pl<strong>an</strong>t. or even to obovate.
112 Flora Neotropica<br />
Leaf texture varies from chartaceous or subcoriaceous<br />
to coriaceous, <strong><strong>an</strong>d</strong> leafindument from<br />
flowers. A subumbellate pistillate inflorescence<br />
is characteristic for <strong>Pourouma</strong> minor. Staminate<br />
dense (<strong>with</strong> the three types of trichomes repre- inflorescences are often more or less dorsivensented)<br />
to sparse.<br />
trally flattened.<br />
The venation varies from pinnate in entire The flowers of the staminate inflorescences are<br />
leaves to subpalmate in incised leaves. In species more or less closely clustered at the end of the<br />
in which palmately incised leaves occur, the bas- br<strong>an</strong>ches. In several species the flowers form<br />
al lateral veins even of entire leaves are normally (sub)globose heads.<br />
br<strong>an</strong>ched, while in species in which incised leaves The inflorescences are mostly ebracteate, alare<br />
not formed, the basal lateral veins are often though minute bracts are found in staminate <strong><strong>an</strong>d</strong><br />
unbr<strong>an</strong>ched or only faintly or sparsely br<strong>an</strong>ched. pistillate inflorescences of P. cucura, <strong><strong>an</strong>d</strong> some-<br />
The tertiary venation is scalariform.<br />
what larger (caducous) bracts at the base of the<br />
The leaf margin is entire or often more or less peduncle are sometimes found in other species.<br />
faintly crenate. In most species the petioles are Staminate Flowers. -Staminate flowers are<br />
relatively long, <strong>with</strong> a considerable variation in pedicellate or sessile <strong><strong>an</strong>d</strong> tetramerous, or somelength<br />
on the same twig. Relatively short peti- times trimerous. The tepals are free, basally conoles,<br />
only slightly varying in length, are found in nate, or almost entirely connate, <strong><strong>an</strong>d</strong> then they<br />
a few species (e.g., P. formicarum). In P. for- form <strong>an</strong> urceolate or infundibuliform peri<strong>an</strong>th.<br />
micarum <strong><strong>an</strong>d</strong> P. myrmecophila the base of the The stamens are short in species <strong>with</strong> free or<br />
petiole is swollen <strong><strong>an</strong>d</strong> has <strong>an</strong> abaxial slit. These basally connate tepals, but in species <strong>with</strong> almost<br />
two species are myrmecophilous.<br />
entirely connate tepals the filaments are longer<br />
The mostly large stipules are connate <strong><strong>an</strong>d</strong> fully th<strong>an</strong> the tepals. Probably in connection <strong>with</strong> the<br />
amplexicaul, <strong><strong>an</strong>d</strong> appear to be of import<strong>an</strong>ce for rather narrow opening of the urceolate peri<strong>an</strong>th,<br />
the protection of young inflorescences-espe- the <strong>an</strong>thers are outside the peri<strong>an</strong>th (long?) before<br />
cially staminate inflorescences <strong>with</strong> flowers <strong>with</strong> <strong>an</strong>thesis. The filaments are free, sometimes baa<br />
connate (more or less urceolate) peri<strong>an</strong>th <strong><strong>an</strong>d</strong> sally connate, but in <strong>Pourouma</strong> melinonii subsp.<br />
the <strong>an</strong>thers already exserted before <strong>an</strong>thesis, as glabrata they c<strong>an</strong> be entirely connate, thus largely<br />
in P. tomentosa. In some species the stipules are similar to the <strong><strong>an</strong>d</strong>roecium characteristic for<br />
short, e.g., P. ovata. While in most species the <strong>Coussapoa</strong>. In the center of the flower one c<strong>an</strong><br />
stipules are caducous, they are (sub)persistent in usually find a tuft of hairs, which might represent<br />
P. myrmecophila, P. oraria, <strong><strong>an</strong>d</strong> P. stipulacea. a pistillode.<br />
The stipules leave a horizontal <strong>an</strong>nular scar, not Pistillate Flowers. -Pistillate flowers are aloblique<br />
as in <strong>Coussapoa</strong>. This feature may help ways pedicellate. The peri<strong>an</strong>th is tubular <strong>with</strong> <strong>an</strong><br />
to distinguish sterile specimens of the two genera, entire or slightly lobed margin. The ovary is free<br />
of which the leaves c<strong>an</strong> be very similar. from the peri<strong>an</strong>th. The short style bears a stigma<br />
The leaf characters mentioned are more or less that is subpeltate or peltate to knob-like, in Poucorrelated.<br />
In taxa <strong>with</strong> incised leaves the leaves rouma minor.<br />
tend to be large, often subcoriaceous <strong><strong>an</strong>d</strong> densely Fruits <strong><strong>an</strong>d</strong> Seeds.- The fruit has a dry pericarp<br />
hairy, <strong>with</strong> relatively long petioles, often quite <strong><strong>an</strong>d</strong> is completely enclosed by the enlarged perivariable<br />
in length, <strong><strong>an</strong>d</strong> often <strong>with</strong> long stipules. <strong>an</strong>th. The inner layer of the fruiting peri<strong>an</strong>th be-<br />
Contrariwise, in taxa <strong>with</strong> entire leaves, the leaves comes soft <strong><strong>an</strong>d</strong> whitish <strong><strong>an</strong>d</strong> the other layer (skin)<br />
tend to be medium-sized, usually coriaceous <strong><strong>an</strong>d</strong> ? leathery <strong><strong>an</strong>d</strong> at full maturity often purple to<br />
often rather sparsely hairy. They have relatively black or red-brown. The whole (fruit <strong><strong>an</strong>d</strong> perishort<br />
petioles, more or less const<strong>an</strong>t in length, <strong>an</strong>th) c<strong>an</strong> be indicated as a pseudodrupe (or func<strong><strong>an</strong>d</strong><br />
the stipules tend to be small.<br />
tional drupe).<br />
Inflorescences. -The inflorescences are nearly The pedicel under mature fruits may become<br />
always ramified, basically <strong>with</strong> three br<strong>an</strong>ches reddish (a contrasting color against black?).<br />
departing from the top of the peduncle. Further Whether this color is caused by a ch<strong>an</strong>ge of color<br />
ramification is usually (sub)dichotomous. The of tissues of the pedicel or by the presence of redprimary<br />
ramification c<strong>an</strong> be dichotomous as well. brown pluricellular hairs is not clear.<br />
In pistillate inflorescences the ramification is often The fruiting peri<strong>an</strong>th is reported to be aromore<br />
reduced th<strong>an</strong> in the staminate ones, <strong>with</strong> matic for some species (e.g., <strong>Pourouma</strong> bicolor).<br />
a tendency to a subumbellate arr<strong>an</strong>gement of the <strong>Pourouma</strong> <strong><strong>an</strong>d</strong> the Afric<strong>an</strong> Myri<strong>an</strong>thus are
Morphology; Phenology; Pollination 113<br />
macrospermous, in contrast to the other four<br />
Phenology<br />
genera of the <strong>Cecropiaceae</strong>.<br />
General Remarks. -The greatest variation of From label data alone it is difficult if not imcharacters<br />
(mainly, but not only of the leaves) in possible to draw conclusions about the phenolthe<br />
genus is found among taxa in which incised ogy of a particular species.<br />
leaves do not occur const<strong>an</strong>tly. Only part of this In <strong>an</strong> area explored (Montagne de Trinite,<br />
variation is connected <strong>with</strong> distribution. Taxa French Gui<strong>an</strong>a, short rainy season, J<strong>an</strong>-Feb 1984)<br />
<strong>with</strong> const<strong>an</strong>tly incised leaves in adult specimens<br />
none of the numerous <strong>Pourouma</strong> specimens of<br />
<strong><strong>an</strong>d</strong> those <strong>with</strong> entire leaves in all stages of de- different species was bearing flowers or fruits.<br />
velopment tend to be rather uniform, even if This<br />
they suggested seasonal flowering. The combihave<br />
a relatively wide distribution.<br />
nation of label data of all <strong>Pourouma</strong> collections<br />
It is somewhat difficult to indicate which char- made in the Gui<strong>an</strong>as gives a pattern indicating<br />
acters c<strong>an</strong> be regarded as derived or as a main<br />
special- flowering period in August-October (in<br />
ized. In several of its characters (especially of the the long dry season) <strong><strong>an</strong>d</strong> a fruiting period which<br />
vegetative parts) <strong>Pourouma</strong> is (like the Afric<strong>an</strong> may extend to February, thus from the end of<br />
genus Myri<strong>an</strong>thus) intermediate between Cecro- the long dry season into a short rainy season.<br />
pia (<strong><strong>an</strong>d</strong> the Afric<strong>an</strong> genus Mus<strong>an</strong>ga) <strong><strong>an</strong>d</strong> Cous- According to Croat (1978) P. bicolor fruits in<br />
sapoa (<strong><strong>an</strong>d</strong> the Malesi<strong>an</strong> P<strong>an</strong>ama in the end of the<br />
genus Poikilospermum).<br />
dry season. According<br />
From about the middle of the spectrum of vari- to Martius (in Spix & Martius, 1831) P. cecroation<br />
in <strong>Pourouma</strong> one could indicate a pro- piifolia c<strong>an</strong> bear (ripe?) fruits in May <strong><strong>an</strong>d</strong> Nogression<br />
towards Cecropia, ending in a form <strong>with</strong> vember. The occurrence of these two periods of<br />
thick br<strong>an</strong>ches <strong><strong>an</strong>d</strong> large, const<strong>an</strong>tly incised leaves fruiting gets some support from the label data of<br />
<strong>with</strong> m<strong>an</strong>y incisions (but never peltate as in Ce- the examined material of this species. Falcao <strong><strong>an</strong>d</strong><br />
cropia), represented, e.g., by P. cecropiifolia <strong><strong>an</strong>d</strong> Lleras (1980) found that P. cecropiifolia in the<br />
P. cuspidata; <strong><strong>an</strong>d</strong> <strong>an</strong>other progression towards M<strong>an</strong>aus region was flowering at the height of the<br />
<strong>Coussapoa</strong>, ending in a form <strong>with</strong> rather slender rainy season (April-June), <strong><strong>an</strong>d</strong> producing a crop<br />
br<strong>an</strong>ches <strong><strong>an</strong>d</strong> medium-sized, const<strong>an</strong>tly entire of ripe fruit at the end of the dry season to the<br />
leaves. As indicated (p. 5), there is <strong>an</strong> overlap beginning of the next rainy season.<br />
of the leaf characters of <strong>Pourouma</strong> <strong><strong>an</strong>d</strong> Cous- The period between flowering <strong><strong>an</strong>d</strong> bearing ripe<br />
sapoa, <strong><strong>an</strong>d</strong> sterile material of both genera c<strong>an</strong> be fruit appears to be longer in P. cecropiifolia th<strong>an</strong><br />
easily confused.<br />
in (most?) other species of <strong>Pourouma</strong>. The dis-<br />
In addition to these characters one c<strong>an</strong> also crep<strong>an</strong>cy between the data supplied by Falcao<br />
regard as derived: sparse indument (e.g., Pourou- <strong><strong>an</strong>d</strong> Lleras <strong><strong>an</strong>d</strong> the observations of Martius might<br />
ma ovata), elliptic to obovate leaves (e.g., P. be caused by the presence of several strains. In<br />
phaeotricha <strong><strong>an</strong>d</strong> P. bolivarensis), unbr<strong>an</strong>ched lat- the publication "Underexploited Tropical Pl<strong>an</strong>ts<br />
eral veins (e.g., P. acuminata <strong><strong>an</strong>d</strong> P. <strong>with</strong><br />
ovata), peti- Promising Economic Value" (National<br />
olulate leaf segments (in taxa <strong>with</strong> incised leaves), Academy of Sciences, Washington, D.C., 1975)<br />
as in P. tomentosa subsp. persecta, short a<br />
petioles fruiting period of three months in the rainy<br />
of const<strong>an</strong>t length (P. formicarum), swollen <strong><strong>an</strong>d</strong> season is mentioned for P. cecropiifolia.<br />
saccate base of the petiole (P. formicarum, P.<br />
myrmecophila), short stipules (e.g., P. ovata), sta-<br />
Pollination<br />
minate flowers in globose heads (P. melinonii<br />
<strong><strong>an</strong>d</strong> P. myrmecophila), subumbellate pistillate Little is known about pollination in Pourouinflorescences<br />
(P. minor), elongate peduncle of ma. Neither morphological features (of infloresthe<br />
pistillate inflorescence (P. ferruginea <strong><strong>an</strong>d</strong> P. cences <strong><strong>an</strong>d</strong> flowers) nor habit <strong><strong>an</strong>d</strong> habitat (small<br />
ovata), staminate flowers <strong>with</strong> fused tepals (P. to medium-sized trees in forests) suggest the ocmollis),<br />
stamens <strong>with</strong> partly or fully connate fil- currence of wind pollination. The herbarium maaments<br />
(P. melinonii <strong><strong>an</strong>d</strong> P. napoensis respec- terial indicates that most pistils develop into<br />
tively), <strong><strong>an</strong>d</strong> knob-like stigma (P. minor). fruits, which suggests <strong>an</strong> effective pollination sys-<br />
The above summation of derived characters tem. Falcao <strong><strong>an</strong>d</strong> Lleras (1980) found a number<br />
<strong><strong>an</strong>d</strong> specializations shows that they tend to be of bee species as pollinators of P. cecropiifolia.<br />
'at the <strong>Coussapoa</strong> side' of the variation spectrum The bees collect pollen in the staminate infloin<br />
<strong>Pourouma</strong>.<br />
rescences <strong><strong>an</strong>d</strong> fly to pistillate inflorescences. The
114<br />
authors do not give indications about the attract<strong>an</strong>t(s)<br />
of the pistillate inflorescences. One<br />
may wonder whether the pluricellular ('gl<strong><strong>an</strong>d</strong>ular')<br />
hairs, which occur so abund<strong>an</strong>tly in inflorescences<br />
of most <strong>Pourouma</strong> species (as a powdery<br />
layer, as in dry material?), may play a role<br />
(as attract<strong>an</strong>t or a substitute for pollen?).<br />
Dispersal<br />
Again, little is known about dispersal of <strong>Pourouma</strong><br />
seeds. One may assume that the 'fruits'<br />
<strong>with</strong> a fleshy mesocarp-like inner layer of the<br />
enlarged peri<strong>an</strong>th, <strong><strong>an</strong>d</strong> a purple to black (or a<br />
red-brown) exocarp-like outer layer (skin) are<br />
eaten by several frugiverous arboreal <strong>an</strong>imals;<br />
monkeys are mentioned in label data.<br />
The presence of very long peduncles of pistillate<br />
inflorescences in P. ovata <strong><strong>an</strong>d</strong> P. ferruginea<br />
could be <strong>an</strong> adaptation to dispersal by bats.<br />
Relations <strong>with</strong> Ants<br />
<strong>Pourouma</strong> formicarum <strong><strong>an</strong>d</strong> P. myrmecophila<br />
are myrmecophilous. These species have swollen<br />
<strong><strong>an</strong>d</strong> saccate bases of the petiole ("domatia"), usually<br />
occupied by small <strong>an</strong>ts. According to Benson<br />
(1985) these <strong>an</strong>ts belong to genus Allomerus <strong><strong>an</strong>d</strong><br />
presumably eat food bodies, produced on the<br />
inner surface of the domatia. Sometimes, as in<br />
P. mollis subsp. triloba (collection Pr<strong>an</strong>ce et al.<br />
7907) <strong><strong>an</strong>d</strong> most collections of P. napoensis, <strong>an</strong>ts<br />
(of the genus Crematogaster) build tubular nests<br />
on the leafy twigs <strong><strong>an</strong>d</strong> on the lower leaf surface.<br />
The nest-building on the lower leaf surface (only<br />
if the main venation is subpalmate?) starts in the<br />
axils of the main veins; then the nests look like<br />
domatia. Later on these nests are often extended<br />
as tubes. A peculiar aspect is the use of the long,<br />
rather stiff hairs (occurring on various parts of<br />
the pl<strong>an</strong>t) in the nest building, perhaps as a protective<br />
cover of the nests.<br />
Distribution <strong><strong>an</strong>d</strong> Ecology<br />
The distribution r<strong>an</strong>ge of <strong>Pourouma</strong> is almost<br />
the same as that of <strong>Coussapoa</strong>. <strong>Pourouma</strong> is con-<br />
fined to rain forest areas, but is notably absent<br />
in the West Indies <strong><strong>an</strong>d</strong> southern Mexico. Most<br />
are lowl<strong><strong>an</strong>d</strong> species, found at altitudes up to 1000<br />
m, occasionally up to ca. 1500 m, <strong><strong>an</strong>d</strong> excep-<br />
tionally (P. tomentosa in Bolivia <strong><strong>an</strong>d</strong> Peru) up<br />
Flora Neotropica<br />
to ca. 1800 or 1900 m. Some taxa (P. bolivarensis<br />
<strong><strong>an</strong>d</strong> P. gui<strong>an</strong>ensis subsp. venezuelensis) appear<br />
to be confined to altitudes of about 1000 m (600-<br />
1400 m). The unnamed collections Gentry et al.<br />
22866 <strong><strong>an</strong>d</strong> D. N. Smith et al. 4743 (see unnamed<br />
collections 1, p. 193) may represent a mont<strong>an</strong>e<br />
taxon. Most species are components of non-inundated<br />
forest, but P. acuminata is often found<br />
in (occasionally?) inundated forest. <strong>Pourouma</strong><br />
(probably) needs light for germination <strong><strong>an</strong>d</strong> is<br />
commonly found in places <strong>with</strong> more or less distinct<br />
traces of previous disturb<strong>an</strong>ce of the forest<br />
(e.g., chablis) or in rather open forest (e.g., on<br />
rather poor ? s<strong><strong>an</strong>d</strong>y soils) <strong><strong>an</strong>d</strong> secondary forest.<br />
A number of species have a wide distribution.<br />
All these species are represented in the Amazon<br />
Basin <strong><strong>an</strong>d</strong> the Gui<strong>an</strong>as. Two, P. bicolor <strong><strong>an</strong>d</strong> P.<br />
minor, extend to Central America, <strong><strong>an</strong>d</strong> P. melinonii<br />
to P<strong>an</strong>ama. <strong>Pourouma</strong> mollis, P. velutina,<br />
<strong><strong>an</strong>d</strong> P. gui<strong>an</strong>ensis also occur in eastern Brazil,<br />
the latter in the coastal mountain r<strong>an</strong>ge of Venezuela<br />
as well. <strong>Pourouma</strong> tomentosa, however,<br />
is confined to the Amazon Basin <strong><strong>an</strong>d</strong> the Gui<strong>an</strong>as.<br />
Most of these wide-spread species are very<br />
variable in a confusing way. In five of these<br />
species, two or more infraspecific taxa are recognized.<br />
The other species are much more uniform <strong><strong>an</strong>d</strong><br />
have much smaller or even very small distribution<br />
r<strong>an</strong>ges. Only one of the this group of<br />
species (P. hirsutipetiolata, from northern <strong><strong>an</strong>d</strong><br />
western Colombia) is subdivided into two subspecies<br />
(but in a more clear way th<strong>an</strong> in the widespread<br />
species). This species <strong><strong>an</strong>d</strong> P. oraria (from<br />
western Colombia) are the only ones not occurring<br />
in the Amazon Basin or the Gui<strong>an</strong>as. Of the<br />
remaining species, three (P. bolivarensis, P. saulensis,<br />
<strong><strong>an</strong>d</strong> P. stipulacea), are confined to the<br />
Gui<strong>an</strong>a region, all probably <strong>with</strong> very small distribution<br />
r<strong>an</strong>ges. The others are confined to the<br />
Amazon Basin, or almost so as P. acuminata, P.<br />
cucura, <strong><strong>an</strong>d</strong> P. ovata (found in the Guine<strong>an</strong> part<br />
of Venezuela).<br />
The present r<strong>an</strong>ge of distribution of the very<br />
uniform P. cecropiifolia might be largely <strong>an</strong>thropogenous.<br />
The Amazon Basin <strong><strong>an</strong>d</strong> the Gui<strong>an</strong>as in all respects<br />
represent the center of the genus. Three<br />
centers of distribution c<strong>an</strong> be distinguished:<br />
(a) The Upper Amazon Basin (down to ca.<br />
60?W) <strong>with</strong> <strong>Pourouma</strong> acuminata, P. cecropiifolia,<br />
P. cucura, P. cuspidata, P. elliptica, P. fer-
Distribution <strong><strong>an</strong>d</strong> Ecology<br />
ruginea, P. formicarum, P. herrerensis, P. myrmecophila,<br />
P. napoensis, P. ovata, P. phaeotricha,<br />
as well as P. bicolor subsp. tessm<strong>an</strong>nii, P. mollis<br />
subsp. triloba, <strong><strong>an</strong>d</strong> P. tomentosa subspp. apiculata,<br />
persecta, <strong><strong>an</strong>d</strong> tomentosa. A concentration<br />
of taxa is found in the north-western part of this<br />
region.<br />
(b) The Gui<strong>an</strong>a (or Guay<strong>an</strong>a) region, the Lower<br />
Amazon Basin (Brazil: eastern Para <strong><strong>an</strong>d</strong> Amapa),<br />
including <strong>an</strong> extension to eastern Brazil, <strong>with</strong><br />
the following taxa confined or distinctly connected<br />
to this area: <strong>Pourouma</strong> bolivarensis, P.<br />
saulensis, as well as P. bicolor subsp. digitata, P.<br />
melinonii subsp. melinonii, P. mollis subsp. mol-<br />
lis, P. tomentosa subsp. essequiboensis <strong><strong>an</strong>d</strong> subsp. <strong>Pourouma</strong> shows m<strong>an</strong>y morphological simimaroniensis,<br />
<strong><strong>an</strong>d</strong> P. velutina.<br />
larities to the Afric<strong>an</strong> genus Myri<strong>an</strong>thus (<strong>with</strong> 7<br />
(c) The Pacific Coastal region (Colombia <strong><strong>an</strong>d</strong> species), revised by de Ruiter (1976). The sim-<br />
Ecuador) to P<strong>an</strong>ama, <strong>with</strong> extensions to Gua- ilarities are found in the habit, the leaf shape <strong><strong>an</strong>d</strong><br />
temala <strong><strong>an</strong>d</strong> through northern Colombia to north- dimension <strong><strong>an</strong>d</strong> their infraspecific variation, <strong><strong>an</strong>d</strong><br />
western Venezuela, <strong>with</strong> <strong>Pourouma</strong> hirsutipetio- the (variable) length of the petiole. Furthermore,<br />
lata, P. oraria, P. bicolor subsp. choco<strong>an</strong>a <strong><strong>an</strong>d</strong> the staminate inflorescences show similarities,<br />
subsp. scobina, <strong><strong>an</strong>d</strong> P. melinonii subsp. glabrata. but the pistillate ones are different, as they are<br />
The features of the East-Brazili<strong>an</strong> populations capitate in Myri<strong>an</strong>thus, where the pistillate flowof<br />
P. gui<strong>an</strong>ensis, P. mollis, <strong><strong>an</strong>d</strong> P. velutina, re- ers are sessile <strong><strong>an</strong>d</strong> clustered in a single head. Both<br />
semble those of the material of the same species genera are macrospermous in contrast to the othin<br />
the Lower Amazon Basin <strong><strong>an</strong>d</strong> adjacent parts er genera of the <strong>Cecropiaceae</strong>. The two genera<br />
of the Gui<strong>an</strong>as.<br />
also show strong ecological similarities <strong><strong>an</strong>d</strong> are<br />
<strong>Pourouma</strong> bicolor, P. melinonii, P. mollis <strong><strong>an</strong>d</strong> found in similar types of habitat. These two gen-<br />
P. tomentosa have one (or two) morphologically era may also be regarded not only as genetically<br />
distinct entities (subspecies) distinctly connected related but also as the most primitive among the<br />
<strong>with</strong> the Gui<strong>an</strong>a (Lowl<strong><strong>an</strong>d</strong>) region <strong><strong>an</strong>d</strong> the ad- <strong>Cecropiaceae</strong>.<br />
jacent part of the Amazon Basin. In P. bicolor The leaves of some taxa, <strong>Pourouma</strong> cecropi<strong><strong>an</strong>d</strong><br />
P. melinonii one or two disjunct subspecies ifolia <strong><strong>an</strong>d</strong> P. aspera subsp. digitata, resemble those<br />
occur in the third center of the genus <strong><strong>an</strong>d</strong> in P. of Cecropia because of the great number of ingui<strong>an</strong>ensis<br />
a disjunct subspecies occurs in north- cisions, but the leaves are never peltate as they<br />
ern Venezuela.<br />
always are in Cecropia. Leaves of some other<br />
In the wide-spread species the material in the taxa resemble leaves of some <strong>Coussapoa</strong> species<br />
Upper Amazon Basin is morphologically more (see p. 5).<br />
variable th<strong>an</strong> in the other parts of their distribution<br />
r<strong>an</strong>ges. <strong>Pourouma</strong> cucura shares this trait.<br />
Systematic<br />
In<br />
Arr<strong>an</strong>gement<br />
most parts of the distribution r<strong>an</strong>ge of Pouof<br />
the<br />
rouma the taxa (even those at the<br />
Species<br />
subspecific<br />
level) c<strong>an</strong> be recognized, <strong><strong>an</strong>d</strong> well-collected ma- On the basis of overall similarities, several<br />
terial of adult specimens c<strong>an</strong> readily be identi- groups of species c<strong>an</strong> be recognized. Not all<br />
fied. In contrast, the situation in the Upper Am- species c<strong>an</strong> be placed <strong>with</strong> certainty into one of<br />
azon Basin, especially in its northwestern part, them, however, as the staminate inflorescences<br />
is rather unclear <strong>with</strong> regard to the delimitation are not known (as in P. bolivarensis, P. saulensis,<br />
of subspecies (even species in some cases). In <strong><strong>an</strong>d</strong> P. stipulacea).<br />
several species (e.g., P. bicolor, P. mollis, P. to- One group of species is characterized by havmentosa)<br />
morphologically aberr<strong>an</strong>t collections ing the tepals of the staminate flower almost enhave<br />
been made, e.g., in the valley of the Rio tirely connate, forming <strong>an</strong> urceolate infundibu-<br />
S<strong>an</strong>tiago (Peru, Amazonas).<br />
liform peri<strong>an</strong>th <strong>with</strong> the filaments of the stamens<br />
115<br />
In some cases (P. cucura <strong><strong>an</strong>d</strong> P. tomentosa<br />
subsp. persecta) the present data suggest the oc-<br />
currence of a disjunction between populations in<br />
the southwestern part <strong><strong>an</strong>d</strong> those in the north-<br />
western part of the Amazon Basin.<br />
The genus appears to be underrepresented in<br />
a zone from southeastern to northwestern Para<br />
largely consisting of'mata de cip6' <strong><strong>an</strong>d</strong> north of<br />
the Rio Solim6es of patches of'campos de terra<br />
firme' (Pires, 1973). A similar disjunctive gap is<br />
found in <strong>Coussapoa</strong> (see p. 10).<br />
Systematic Position of the Genus
116 Flora Neotropica<br />
longer th<strong>an</strong> the tepals. The staminate flowers are the fleshy (mesocarp-like) inner layer of the periusually<br />
clustered in distinct, terminal, globose <strong>an</strong>th strongly resembles that ofgrapes. P. cecropiheads.<br />
The arachnoid indument often occurs on ifolia (Amazon grape, uva, uvilla, imbauiba do<br />
the lateral veins of the lamina beneath or also vinho) is one of the taxa mentioned in "Underon<br />
other parts, such as the stipules. The lamina exploited Tropical Pl<strong>an</strong>ts <strong>with</strong> Promising Ecois<br />
usually smooth above. The leaves are entire nomic Value" (National Academy of Sciences,<br />
or palmately incised. The basal part of the leaf Washington, D.C., 1975). P. cecropiifolia c<strong>an</strong> start<br />
margin is formed by the basal part of the basal reproducing (thus forming fruits) a few years after<br />
lateral vein in all (or most?) species. <strong>Pourouma</strong> pl<strong>an</strong>ting, as small trees. Other <strong>Pourouma</strong> species<br />
mollis, P. melinonii, P. hirsutipetiolata, P. to- may have fruits as tasty as those of P. cecropimentosa,<br />
P. herrerensis, <strong><strong>an</strong>d</strong> P. napoensis com- ifolia, but the fleshy mesocarp-like layer is usuprises<br />
this group, as may P. acuminata <strong><strong>an</strong>d</strong> P. ally thinner <strong><strong>an</strong>d</strong> m<strong>an</strong>y of them become mediumstipulacea,<br />
of which the staminate flowers are sized trees before fruiting. <strong>Pourouma</strong> cecropistill<br />
unknown.<br />
ifolia fruits are used to make a sweet wine. For<br />
About equally well-defined is a group formed a detailed study on the reproduction <strong><strong>an</strong>d</strong> propby<br />
<strong>Pourouma</strong> myrmecophila, P. formicarum, <strong><strong>an</strong>d</strong> erties of the fruits of P. cecropiifolia see Falcao<br />
P. phaeotricha, in which the laminae are mostly <strong><strong>an</strong>d</strong> Lleras (1980).<br />
scabrous above. The staminate flowers, <strong>with</strong> (almost)<br />
free tepals <strong><strong>an</strong>d</strong> short filaments, are clus-<br />
Taxonomy<br />
tered in distinct, terminal, globose heads. The<br />
hairs of the tepals <strong><strong>an</strong>d</strong> the stigma are long (com- <strong>Pourouma</strong> Aublet, Hist. pl. Gui<strong>an</strong>e 2: 891. 1775;<br />
pared <strong>with</strong> other species). The lamina is entire Tr6cul, Ann. Sci. Nat. Bot., Ser. 3, 8: 100.<br />
or 3-lobed <strong><strong>an</strong>d</strong> the tendency toward entire leaves 1847; Miquel in Martius, Fl. bras. 4(1): 122.<br />
is strong. The base of the petiole of P. myrme- 1853; Bentham in Bentham & Hooker, Gen.<br />
cophila <strong><strong>an</strong>d</strong> P. formicarum is swollen <strong><strong>an</strong>d</strong> sac- pi. 3(1): 357. 1880; Baillon, Hist. pl. 6: 141.<br />
cate. These two species are myrmecophilous. 1875-1876; Engler in Engler & Pr<strong>an</strong>tl, Nat.<br />
Less well-defined is a group of species in which Pfl<strong>an</strong>zenfam. 3(1): 95. 1889; Benoist, Arch.<br />
the lamina is usually scabrous <strong><strong>an</strong>d</strong>/or has more Bot. Mem. 5: 29. 1931; Woodson & Schery,<br />
th<strong>an</strong> 7 incisions. The staminate flowers are more Ann. Missouri Bot. Gard. 47: 165. 1960. Berg<br />
or less clustered, but not in globose heads. In in L<strong>an</strong>jouw & Stoffers, Fl. Suriname 5(1): 265.<br />
most of the species the lamina is incised. This 1975. Burger, Fieldi<strong>an</strong>a Bot. 40: 200. 1977;<br />
group is formed by <strong>Pourouma</strong> gui<strong>an</strong>ensis, P. as- Berg & v<strong>an</strong> Heusden, Proc. Kon. Ned. Akad.<br />
pera, P. velutina, P. cucura, P. cuspidata, <strong><strong>an</strong>d</strong> P. Wetensch., Ser. C., 91(2): 105. (1988). Type<br />
cecropiifolia.<br />
species. <strong>Pourouma</strong> gui<strong>an</strong>ensis Aublet.<br />
The rest of the species c<strong>an</strong>not be satisfactorily Trees, terrestrial, often <strong>with</strong> stilt-roots. Leaves<br />
grouped, except for pairs of species showing more in spirals, entire or<br />
or<br />
palmately incised,<br />
less distinct<br />
margin<br />
similarities, such as <strong>Pourouma</strong> entire to subcrenate; stipules fused,<br />
minor <strong><strong>an</strong>d</strong> P. bolivarensis or<br />
fully am-<br />
P. ovata <strong><strong>an</strong>d</strong> P.<br />
plexicaul. Inflorescences in the leaf axils,<br />
saulensis.<br />
mostly<br />
in pairs, br<strong>an</strong>ched (or to subumbellate in pistil-<br />
<strong>Pourouma</strong> ovata <strong><strong>an</strong>d</strong> P. ferruginea both have late inflorescences),<br />
pistillate inflorescences <strong>with</strong><br />
mostly ebracteate, somevery<br />
long peduncles. times bracteate; staminate flowers sessile or<br />
But they are so different in other<br />
pedfeatures<br />
that<br />
icellate,<br />
this<br />
tepals (3-)4, free or<br />
similarity appears not<br />
connate; stamens<br />
to indicate common<br />
(3-)4,<br />
origin, but<br />
mostly free, sometimes connate;<br />
rather a similar<br />
pistillate<br />
adaptation (to dis- flowers pedicellate, peri<strong>an</strong>th tubular; ovary free,<br />
persal by bats?).<br />
stigma (sub)peltate or sometimes-knob-like. Fruit<br />
<strong>an</strong> achene, large, surrounded by the enlarged,<br />
Uses<br />
more or less fleshy peri<strong>an</strong>th <strong>with</strong> a purplish to<br />
blackish or red-brown outer<br />
Most, if not all, <strong>Pourouma</strong> species have edible<br />
layer at full matufruits.<br />
As far as known, only P. cecropiifolia is<br />
rity; seed <strong>with</strong>out endosperm, embryo straight,<br />
cultivated as a fruit tree, however. The taste of<br />
cotyledons thick, radicle short.
<strong>Pourouma</strong> 117<br />
Key to the Species <strong><strong>an</strong>d</strong> Subspecies of <strong>Pourouma</strong><br />
1. Base of the petiole swollen.<br />
2. Petiole 1-1.5 cm long ..................................................... 8. P. formicarum.<br />
2. Petiole 8-27 cm long ..................................................... 7. P. myremcophila.<br />
1. Base of the petiole not swollen.<br />
3. Stipules (sub)persistent.<br />
4. Lamina smooth above.<br />
5. Stipules <strong>with</strong> dense, white, arachnoid hairs; Guy<strong>an</strong>a. ...................... 14. P. stipulacea.<br />
5. Stipules <strong>with</strong>out white, arachnoid hairs; Colombia .............................. 18. P. oraria.<br />
4. Lamina scabrous above ....................................... 3e. P. bicolor subsp. choco<strong>an</strong>a.<br />
3. Stipules caducous.<br />
6. Lamina more or less scabrous above.<br />
7. Lamina entire.<br />
8. Lateral veins 2 x 7-10 (in large leaves up to 13), the basal pair unbr<strong>an</strong>ched (or faintly<br />
br<strong>an</strong>ched); lamina mostly elliptic to oblong or to subobovate.<br />
9. Pluricellular hairs on the leafy twigs dense; staminate flowers in globose heads; pistillate<br />
inflorescence <strong>with</strong> 11-15 flowers. .......... ......................... 9. P. phaeotricha.<br />
9. Pluricellular hairs on the leafy twigs sparse or lacking; staminate flowers not in globose<br />
heads; pistillate inflorescence <strong>with</strong> 2-8 flowers. ........................ 2. P. velutina.<br />
8. Lateral veins 2 x 10-25, the basal pair br<strong>an</strong>ched; lamina mostly ovate.<br />
10. Lamina above yellowish-hirsute to -hirtellous over the whole surface.<br />
11. Staminate flower <strong>with</strong> connate tepals <strong><strong>an</strong>d</strong> filaments longer th<strong>an</strong> the peri<strong>an</strong>th;<br />
(fruiting) peri<strong>an</strong>th of pistillate flower velutinous. . ............. 25. P. herrerensis.<br />
11. Staminate flower <strong>with</strong> (almost) free tepals <strong><strong>an</strong>d</strong> filaments shorter th<strong>an</strong> the peri<strong>an</strong>th;<br />
(fruiting) peri<strong>an</strong>th of pistillate flower hispidulous to hirtellous ........ 5. P. cucura.<br />
10. Lamina above yellowish- to whitish-hirtellous only on the main veins.<br />
12. White, arachnoid hairs on the main veins of the lamina beneath <strong><strong>an</strong>d</strong>/or also on<br />
the stipules, petioles, leafy twigs, <strong><strong>an</strong>d</strong> peduncles ...........................<br />
.......................................... 13b. P. tomentosa subsp. apiculata.<br />
12. White, arachnoid hairs only in the areoles or also on the smaller veins of the<br />
lamina beneath.<br />
13. Hairs on the lamina beneath (more or less) appressed. .......... 3. P. bicolor.<br />
14. Stipules hairy inside. ...................... 3a. P. bicolor subsp. bicolor.<br />
14. Stipules glabrous inside. ............... 3b. P. bicolor subsp. tessm<strong>an</strong>nii.<br />
13. Hairs on the lamina beneath at least partly patent.<br />
15. Stipules inside densely hairy; pluricellular hairs on the leafy twigs sparse<br />
or lacking; pistillate inflorescences <strong>with</strong> 2-8 flowers ........ 2. P. velutina.<br />
15. Stipules inside glabrous, or if <strong>with</strong> sparse hairs, then pluricellular hairs<br />
on the leafy twigs usually (rather) dense; pistillate inflorescences usually<br />
<strong>with</strong> at least 10 flowers.<br />
16. Pluricellular (brown) hairs sparse or lacking on the leafty twigs <strong><strong>an</strong>d</strong><br />
petioles .. ............................... .......... 5. P. cucura.<br />
16. Pluricellular (brown) hairs (rather) dense on the leafy twigs <strong><strong>an</strong>d</strong> the<br />
petioles.<br />
17. Arachnoid hairs on the lamina beneath confined to the areoles;<br />
tepals of the staminate flowers (almost) free; apex of the pedicels<br />
at fruit not widened.. ....... la. P. gui<strong>an</strong>ensis subsp. gui<strong>an</strong>ensis.<br />
17. Arachnoid hairs on the lamina beneath also on the smaller<br />
veins; tepals of the staminate flowers connate; apex of the pedicels<br />
at fruit widened (aberr<strong>an</strong>t form from Peru)............<br />
.................................. 10. P. m ollis subsp. triloba.<br />
7. Lamina lobed to parted.<br />
18. Stipules hairy inside.<br />
19. Lamina <strong>with</strong> the hairs beneath on the veins more or less appressed <strong><strong>an</strong>d</strong> the smaller<br />
veins pl<strong>an</strong>e or only slightly prominent . ............................... 3. P. bicolor.<br />
20. Lamina (usually) 5-7(-9)-fid to -parted.<br />
21. Lamina <strong>with</strong> arachnoid indument beneath, covering also the almost pl<strong>an</strong>e,<br />
smaller veins.<br />
22. Stipules subpersistent; midsegment of the lamina oblong to elliptic;<br />
leafy twigs often hirsute; P<strong>an</strong>ama <strong><strong>an</strong>d</strong> western Colombia ..........<br />
......... ..................... ...... 3e. P. bicolor subsp. choco<strong>an</strong>a.
118 Flora Neotropica<br />
22. Stipules caducous; midsegment of the lamina l<strong>an</strong>ceolate; leafy twigs<br />
never hirsute; Gui<strong>an</strong>as <strong><strong>an</strong>d</strong> Lower Amazon Basin (to southern Venezuela?).<br />
................................. 3c. P. bicolor subsp. digitata.<br />
21. Lamina <strong>with</strong> arachnoid indument beneath (almost) confined to the areoles,<br />
the smaller veins more or less prominent; Central America <strong><strong>an</strong>d</strong> the Pacific<br />
coastal region of Colombia, Ecuador, <strong><strong>an</strong>d</strong> northwest Venezuela (also Amazoni<strong>an</strong><br />
Peru?). ............................. 3d. P. bicolor subsp. scobina.<br />
20. Lamina (usually) 3-lobed to -fid.<br />
23. Lamina <strong>with</strong> arachnoid indument beneath, covering also the almost pl<strong>an</strong>e<br />
smaller veins or <strong>with</strong> arachnoid indument confined to the areoles, but then<br />
the spots of white indument distinctly separated by the thick smaller veins;<br />
pistillate inflorescences mostly <strong>with</strong> up to 25(-35) flowers; Gui<strong>an</strong>as, Amazon<br />
Basin, to northern Colombia. ................. 3a. P. bicolor subsp. bicolor.<br />
23. Lamina <strong>with</strong> arachnoid indument beneath (almost) confined to the areoles,<br />
the smaller veins more or less prominent <strong><strong>an</strong>d</strong> thin; pistillate inflorescences<br />
<strong>with</strong> up to 50 flowers; Central America, the Pacific coastal region of Colombia,<br />
Ecuador, <strong><strong>an</strong>d</strong> to northwestern Venezuela (also Amazoni<strong>an</strong> Peru?).<br />
........................................... 3d. P. bicolor subsp. scobina.<br />
19. Lamina <strong>with</strong> hairs beneath (partly) patent on the veins, the smaller veins prominent.<br />
24. Leafy twigs yellow-hirsute <strong><strong>an</strong>d</strong>/or the whole upper surface of the lamina hirtellous;<br />
fruiting peri<strong>an</strong>th rarely scabrous; Upper Amazon Basin <strong><strong>an</strong>d</strong> northern Venezuela.<br />
..................................... .............. 1. P. gui<strong>an</strong>ensis.<br />
25. Lamina 3-5-fid to -parted; leafy twigs <strong><strong>an</strong>d</strong> petioles <strong>with</strong> long yellow hairs;<br />
stipules always hairy inside; fruiting peri<strong>an</strong>th 2-2.5 cm long; northern Venezuela.<br />
............................ lb. P. gui<strong>an</strong>ensis subsp. venezuelensis.<br />
25. Lamina entire, 5-7-fid to -parted, or if 3-5-fid to -parted, then the leafy<br />
twigs <strong><strong>an</strong>d</strong> the petioles <strong>with</strong>out long yellow hairs, the stipules glabrous inside<br />
<strong><strong>an</strong>d</strong>/or the fruiting peri<strong>an</strong>th 1.2-2 cm long; Gui<strong>an</strong>a region, eastern Brazil,<br />
Amazon Basin to eastern Colombia. ..... la. P. gui<strong>an</strong>ensis subsp. gui<strong>an</strong>ensis.<br />
24. Leafy twigs pale yellow-hirtellous (to subtomentose); upper surface of the lamina<br />
only hirtellous on the main veins; fruiting peri<strong>an</strong>th usually scabrous; Central<br />
America, the Pacific coastal region of Colombia <strong><strong>an</strong>d</strong> Ecuador (also Amazoni<strong>an</strong><br />
Peru?). ................................ ...... 3d. P. bicolor subsp. scobina.<br />
18. Stipules glabrous inside.<br />
26. Lamina yellow-hirsute to -hirtellous above on the whole surface.<br />
27. Brown, pluricellular hairs on the leafy twigs <strong><strong>an</strong>d</strong> petioles sparse or lacking;<br />
arachnoid hairs on the lamina beneath also on the smaller veins.<br />
28. Lamina 7-fid to -parted, usually longer th<strong>an</strong> 30 cm; peri<strong>an</strong>th of pistillate<br />
flower hirtellous to hirsute; filaments connate. ............ 24. P. napoensis.<br />
28. Lamina at most 5-parted, usually up to 20 cm long; peri<strong>an</strong>th of pistillate<br />
flower hispidulous; filaments free. ........................... 5. P. cucura.<br />
27. Brown, pluricellular hairs on the leafy twigs <strong><strong>an</strong>d</strong> petioles (rather) dense <strong><strong>an</strong>d</strong>/or<br />
the arachnoid hairs on the lamina beneath (almost) confined to the areoles.<br />
.......................................... la. P. gui<strong>an</strong>ensis subsp. gui<strong>an</strong>ensis.<br />
26. Lamina yellow-hirsute to -hirtellous or whitish strigose above only on the main<br />
veins.<br />
29. Pluricellular (brown) hairs sparse or lacking on the leafy twigs <strong><strong>an</strong>d</strong> petioles.<br />
30. Lamina 7-fid to -parted, usually longer th<strong>an</strong> 30 cm; peri<strong>an</strong>th of pistillate<br />
flower hirtellous to hirsute; filaments connate. ............ 24. P. napoensis.<br />
30. Lamina at most 5-parted, usually up to 20 cm long; peri<strong>an</strong>th of pistillate<br />
flower hispidulous; filaments free. ........................... 5. P. cucura.<br />
29. Pluricellular (brown) hairs (rather) dense on the leafy twigs <strong><strong>an</strong>d</strong> petioles.<br />
31. Lamina <strong>with</strong> more or less appressed hairs on the veins beneath. ........<br />
........................................ 3b. P. bicolor subsp. tessm <strong>an</strong>nii.<br />
31. Lamina <strong>with</strong> (partly) patent hairs on the veins .......................<br />
...................................... la. P. gui<strong>an</strong>ensis subsp. gui<strong>an</strong>ensis.<br />
6. Lamina smooth above.<br />
32. Lamina scabrous beneath.<br />
33. Stipules glabrous inside. ..........................................6. P. cuspidata.<br />
33. Stipules hairy inside. ................................. 3a. P. bicolor subsp. bicolor.<br />
32. Lamina smooth beneath.<br />
34. Lamina 7-11-parted (to -fid).
<strong>Pourouma</strong> 119<br />
35. Leafy twigs hirsute, or if glabrous or puberulous, then the incisions reaching<br />
down to the petiole <strong><strong>an</strong>d</strong> the segments often petiolulate.<br />
36. Lamina hirsute to hirtellous above, at least on the main veins ...........<br />
...................................................... 24. P. napoensis.<br />
36. Lamina (sub)glabrous above. ............. 13c. P. tomentosa subsp. persecta.<br />
35. Leafy twigs puberulous; incisions usually not entirely down to the petiole <strong><strong>an</strong>d</strong><br />
leaf segments <strong>with</strong> a broader base.<br />
37. Base of the lamina usually deeply cordate; fruiting peri<strong>an</strong>th 1.5-3.5 cm long;<br />
Upper Amazon Basin to eastern Colombia. .............. 4. P. cecropiifolia.<br />
37. Base of the lamina often truncate to shallowly cordate; fruiting peri<strong>an</strong>th to<br />
ca. 1.5 cm long; Gui<strong>an</strong>as <strong><strong>an</strong>d</strong> Lower Amazon Basin (to southern Venezuela?).<br />
............................................ 3c. P. bicolor subsp. digitata.<br />
34. Lamina entire or at most 5-parted.<br />
38. Petiole <strong>with</strong> a mixture of minute whitish hairs, long yellowish hairs, <strong><strong>an</strong>d</strong> sparse<br />
brown pluricellular hairs.<br />
39. Staminate flowers <strong>with</strong> connate tepals <strong><strong>an</strong>d</strong> filaments longerth<strong>an</strong> the peri<strong>an</strong>th;<br />
(fruiting) peri<strong>an</strong>th of pistillate flower velutinous. ......... 25. P. herrerensis.<br />
39. Staminate flowers <strong>with</strong> (almost) free tepals <strong><strong>an</strong>d</strong> filaments shorter th<strong>an</strong> the<br />
peri<strong>an</strong>th; (fruiting) peri<strong>an</strong>th hispidulous to hirtellous. .......... 5. P. cucura.<br />
38. Petiole <strong>with</strong> indument otherwise.<br />
40. Lamina 3-lobed to 5-parted.<br />
41. Arachnoid hairs of indument confined to the lamina beneath in the<br />
areoles or also the smaller veins.<br />
42. Lamina beneath <strong>with</strong> the hairs on the veins appressed.<br />
43. Stipules hairy inside; tepals of the staminate flowers almost<br />
free.<br />
44. Leafy twigs <strong>with</strong> appressed hairs. ............ 3. P. bicolor.<br />
Repeat 20-20 for subspecies; see also unnamed collection<br />
#5.<br />
44. Leafy twigs <strong>with</strong> patent hairs. ............... 17. P. villosa.<br />
43. Stipules glabrous inside; tepals of the staminate flowers connate.<br />
45. Leafy twigs sparsely appressed-puberulous to strigose.<br />
...................................... 11. P. m elinonii.<br />
46. Base of the lamina truncate to subcordate (occasionally<br />
deeply cordate); fruiting peri<strong>an</strong>th usually densely<br />
puberulous to subvelutinous; stamens <strong>with</strong> the filaments<br />
(almost) free; Gui<strong>an</strong>as <strong><strong>an</strong>d</strong> Amazon Basin. ..<br />
.................. 1 la. P. melinonii subsp. melinonii.<br />
46. Base of the lamina deeply cordate; fruiting peri<strong>an</strong>th<br />
almost glabrous; stamens <strong>with</strong> the filaments mostly<br />
connate, P<strong>an</strong>ama, northern <strong><strong>an</strong>d</strong> western Colombia.<br />
1 lb. P. melinonii subsp. glabrata.<br />
45. Leafy twigs hirsute. . 12b. P. hirsutipetiolata subsp. hispida.<br />
42. Lamina beneath <strong>with</strong> the hairs on the veins patent.<br />
47. Leafy twigs <strong>with</strong> appressed hairs ........................<br />
.......................... 1 la. P. melinonii subsp. melinonii.<br />
47. Leafy twigs <strong>with</strong> patent hairs.<br />
48. Basal part of the margin of the lamina formed by the basal<br />
lateral veins; tepals of the staminate flowers for the greater<br />
part connate; fruiting peri<strong>an</strong>th velutinous. .... 10. P. mollis.<br />
49. Pistillate inflorescences <strong>with</strong> the flowers in two more<br />
or less distinct clusters; Gui<strong>an</strong>as, eastern Brazil, Lower<br />
Amazon Basin. ........ 1 Oa. P. mollis subsp. mollis.<br />
49. Pistillate inflorescences <strong>with</strong> the flowers usually more<br />
or less diffusely distributed; Upper Amazon Basin <strong><strong>an</strong>d</strong><br />
eastern Colombia. ....... 1Ob. P. mollis subsp. triloba.<br />
48. Basal part of the margin of the lamina separated from the<br />
basal lateral veins by mesophyll; tepals of the staminate<br />
flowers free or up to halfway connate; fruiting peri<strong>an</strong>th<br />
sparsely puberulous ....................... 17. P. villosa.<br />
41. Arachnoid hairs also on the petiole, the main veins of the lamina be-
120 Flora Neotropica<br />
neath, the leafy twigs, the stipules, the peduncles <strong><strong>an</strong>d</strong>/or the peri<strong>an</strong>th<br />
of the pistillate flower.<br />
50. Peduncle of the pistillate inflorescence 32-48 cm long; tepals of the<br />
staminate flower almsot free ..................... 16. P. ferruginea.<br />
50. Peduncle of the pistillate inflorescence 2-13 cm long; tepals of the<br />
staminate flower connate.<br />
51. Lower leaf surface tomentose to subvelutinous ...........<br />
................................ lOa. P. m ollis subsp. mollis.<br />
51. Lower leaf surface appressed-puberulous to strigose on the<br />
main veins. ............................... 13. P. tomentosa.<br />
52. Stipules hairy inside. ....13b. P. tomentosa subsp. apiculata.<br />
52. Stipules glabrous inside.<br />
53. Leafy twigs yellow-hirsute (or glabrous). ..........<br />
.................. 13c. P. tomentosa subsp. persecta.<br />
53. Leafy twigs puberulous to hirtellous.<br />
54. Base of the lamina deeply cordate. ...........<br />
......... 13d. P. tomentosa subsp. essequiboensis.<br />
54. Base of the lamina truncate to subcordate.<br />
55. Apex of the lamina acuminate; heads of staminate<br />
flowers 2-3 mm in diameter; Gui<strong>an</strong>as,<br />
Amapa, <strong><strong>an</strong>d</strong> near M<strong>an</strong>aus. ...........<br />
....... 13e. P. tomentosa subsp. maroniensis.<br />
55. Apex of the lamina usually rounded; heads<br />
of staminate flowers 4-6 mm in diameter;<br />
Upper Amazon Basin. ..................<br />
......... 13a. P. tomentosa subsp. tomentosa.<br />
40. Lamina entire.<br />
56. Basal lateral veins br<strong>an</strong>ched.<br />
57. Lamina broadest above or in the middle.<br />
58. Arachnoid hairs only on the lamina beneath in the areoles or<br />
also on the smaller veins.<br />
59. Leafy twigs <strong><strong>an</strong>d</strong> lamina beneath on the main veins <strong>with</strong><br />
appressed hairs. ...................... 22. P. bolivarensis.<br />
59. Leafy twigs <strong><strong>an</strong>d</strong> lamina beneath on the main veins <strong>with</strong><br />
patent hairs.<br />
60. Basal (most) part of the margin of the lamina formed<br />
by the basal lateral veins; tepals of the staminate flower<br />
for the greater part connate. .......... 10. P. mollis.<br />
Repeat 48-48 for subspecies.<br />
60. Basal (most) part of the margin of the lamina separated<br />
from the basal lateral veins by mesophyll; tepals<br />
of the staminate flower free or up to halfway connate.<br />
61. Stipules hairy inside. .............. 17. P. villosa.<br />
61. Stipules glabrous inside. ......... 21. P. elliptica.<br />
58. Arachnoid hairs also on the petiole, the main veins of the<br />
lamina beneath, the leafy twigs, the stipules, the peduncle <strong><strong>an</strong>d</strong>/<br />
or the peri<strong>an</strong>th of the pistillate flower .....................<br />
........................ 13a. P. tomentosa subsp. tomentosa.<br />
57. Lamina broadest below the middle.<br />
62. Lamina <strong>with</strong> ? patent hairs on the main veins beneath.<br />
63. Basal part of the margin of the lamina formed by the basal<br />
lateral veins; tepals of the staminate flower for the greater<br />
part connate.<br />
64. Leafy twigs <strong>with</strong> patent hairs. .......... 10. P. mollis.<br />
Repeat 49-49 for subspecies; see also unnamed collection<br />
#2.<br />
64. Leafy twigs <strong>with</strong> appressed hairs. ................<br />
.................. 1 a. P. melinonii subsp. melinonii.<br />
63. Basal part of the margin of the lamina separated from the<br />
basal lateral veins by mesophyll; tepals of the staminate<br />
flower free or up to halfway connate. ........ 17. P. villosa.<br />
See also unnamed collection #4.<br />
62. Lamina <strong>with</strong> appressed hairs on the main veins beneath.
<strong>Pourouma</strong> 121<br />
65. Arachnoid hairs confined to the areoles or also on the<br />
smaller veins on the lamina beneath.<br />
66. Stipules densely or sparsely hairy inside.<br />
67. Staminate flowers <strong>with</strong> connate tepals <strong><strong>an</strong>d</strong> filaments<br />
longer th<strong>an</strong> the peri<strong>an</strong>th; (fruiting) peri<strong>an</strong>th<br />
of pistillate flower velutinous; stipules<br />
sparsely hairy inside. .......... 25. P. herrerensis.<br />
67. Staminate flowers <strong>with</strong> (almost) free tepals, filaments<br />
shorter th<strong>an</strong> the peri<strong>an</strong>th; (fruiting) peri<strong>an</strong>th<br />
hispidulous, scabrous; stipules densely hairy<br />
inside. .............. 3a. P. bicolor subsp. bicolor.<br />
See also unnamed collections #1, #3, <strong><strong>an</strong>d</strong> #4.<br />
66. Stipules glabrous inside.<br />
68. Leafy twigs sparsely appressed-puberulous to<br />
strigose. ...................... 11. P. melinonii.<br />
Repeat 46-46 for subspecies.<br />
68. Leafy twigs hirsute ........................<br />
..... 12a. P. hirsutipetiolata subsp. hirsutipetiolata.<br />
65. Arachnoid hairs also on the petiole, the main veins of the<br />
lamina beneath, the leafy twigs, the stipules, the peduncles<br />
<strong><strong>an</strong>d</strong>/or the peri<strong>an</strong>th of the pistillate flower.<br />
69. Peduncle of the pistillate inflorescence 32-48 cm long;<br />
tepals of the staminate flower almost free. ........<br />
.................................. 16. P. ferruginea.<br />
69. Peduncle of the pistillate inflorescence 2-13 cm long;<br />
tepals of the staminate flower connate. ...........<br />
.................................. 13. P. tom entosa.<br />
70. Stipules hairy inside........................<br />
.............. 13b. P. tomentosa subsp. apiculata.<br />
70. Stipules glabrous inside.<br />
71. Apex of the lamina usually rounded to emarginate,<br />
sometimes short-acuminate; heads<br />
of the staminate inflorescences 4-6 mm in<br />
diameter; Upper Amazon Basin ..........<br />
......... 13a. P. tomentosa subsp. tomentosa.<br />
71. Apex of the lamina acuminate; heads of the<br />
staminate inflorescences 2-3 mm in diameter;<br />
Gui<strong>an</strong>as, Amapa, <strong><strong>an</strong>d</strong> near M<strong>an</strong>aus.<br />
....... 13e. P. tomentosa subsp. maroniensis.<br />
56. Basal lateral veins unbr<strong>an</strong>ched.<br />
72. Stipules 1-3 cm long.<br />
73. Leafy twigs white-puberulous to subglabrous.<br />
74. Lateral veins 2 x 7-12; stipules glabrous inside; peduncle<br />
of the pistillate inflorescence 3-5 cm long. . 19. P. saulensis.<br />
74. Lateral veins 2 x 9-21; stipules hairy inside; peduncle of<br />
the pistillate inflorescence up to 32 cm long. .. 20. P. ovata.<br />
73. Leafy twigs yellow-sericeous to -hirtellous .... 9. P. phaeotricha.<br />
72. Stipules 3-13 cm long.<br />
75. Lower surface of the lamina initially entirely or largely covered<br />
<strong>with</strong> white or brownish, arachnoid hairs; peduncle of the pistillate<br />
inflorescence up to ca. 50 cm long. ..... 16. P. ferruginea.<br />
75. Lower surface of the lamina <strong>with</strong> white arachnoid hairs in the<br />
areoles only (or sometimes extending to the main veins); peduncle<br />
of the pistillate inflorescence up to ca. 10 cm long.<br />
76. Lamina <strong>with</strong> dense yellow hairs on the main veins beneath.<br />
77. Leafy twigs <strong>with</strong> very short <strong><strong>an</strong>d</strong> much longer yellow<br />
hairs. ............................... 21. P. elliptica.<br />
77. Leafy twigs <strong>with</strong> hairs of about equal length. ......<br />
...................................... 23. P. m inor.<br />
76. Lamina <strong>with</strong> sparse <strong><strong>an</strong>d</strong> inconspicuous hairs on the main<br />
veins beneath.<br />
78. Hairs on leafy twigs very short ..... 25. P. herrerensis.<br />
78. Hairs on leafy twigs long or absent. ... 15. P. acuminata.
122 Flora Neotropica<br />
1. <strong>Pourouma</strong> gui<strong>an</strong>ensis Aublet, Hist. pl. Gui<strong>an</strong>e<br />
2: 892, t. 341. 1775; Miquel in Martius, Fl.<br />
bras. 4(1): 127. 1853; Berg, Fl. Suriname 5(1):<br />
267. 1975; Croat, Flora Barro Colorado Isl<strong><strong>an</strong>d</strong><br />
360. 1978. Type. French Gui<strong>an</strong>a. Sinnamary<br />
R., <strong>with</strong>out date (Y), Aublet s.n. (holotype, BM,<br />
only photographs in NY <strong><strong>an</strong>d</strong> U seen; isotypes?,<br />
LE, S).<br />
(8-)10-25; pedicel 0.2-0.5 cm, in fruit 0.4-1 cm<br />
long, peri<strong>an</strong>th 2-4 mm long, subvelutinous,<br />
sometimes sparsely puberulous, stigma subpel-<br />
tate, 1.5-2 mm diam., shortly pilose. Fruiting<br />
peri<strong>an</strong>th purple, (sub)ovoid to ellipsoid, 1.2-2 or<br />
2-2.5 cm long, mostly (sub)velutinous, some-<br />
times sparsely puberulous, scabrous or almost<br />
glabrous.<br />
This taxon c<strong>an</strong> be difficult to distinguish from<br />
P. bicolor. The two taxa have overlapping dis-<br />
tributions <strong><strong>an</strong>d</strong> are each very variable <strong>with</strong>, more-<br />
Tree, up to 30 m tall. Leafy twigs 3-15 mm<br />
thick, white-puberulous, white- to yellow-hirtellous,<br />
yellow-(sub)velutinous or yellow-hirsute, over, quite <strong>an</strong> overlap of characters <strong><strong>an</strong>d</strong> their<br />
<strong>with</strong> dense to/or (very) sparse, brown, pluricel- variation. Considering the variation in both taxa,<br />
lular hairs; lenticels rather conspicuous. Lamina one would be inclined to combine the two. How-<br />
3-fid to 3-lobed, 3- or 5-fid (to -lobed or to -part- ever, as distinct morphological entities occur in<br />
ed), sometimes 5-7-fid to -parted ca. 10-25(-45) the same areas <strong>with</strong>out having clear intermedix<br />
10-25(-45) cm, or entire (to 3-lobed), broadly ates it is likely that they are genetically separated;<br />
ovate to elliptic (to oblong), (4-)10-20 x (2-)7- only in a few cases do the collected specimens<br />
14 cm, subcoriaceous to chartaceous, apex acu- show ? intermediate features.<br />
minate, base cordate, the sinus rather shallow <strong>Pourouma</strong> gui<strong>an</strong>ensis c<strong>an</strong> be distinguished<br />
<strong><strong>an</strong>d</strong> wide or deep <strong><strong>an</strong>d</strong> narrow, often <strong>with</strong> over- from P. bicolor in the following ways:<br />
lapping lobes, sometimes truncate, in entire leaves (a.) The indument of the inner surface of the<br />
to rounded (to subacute); upper surface scabrous, stipules. In subsp. gui<strong>an</strong>ensis the inner surface<br />
on the main veins hirtellous or hirsute or the of the stipules is usually glabrous or <strong>with</strong> sparse,<br />
whole surface hirtellous to subvelutinous, lower white hairs. However, in some forms of subsp.<br />
surface subtomentose to subvelutinous or hir- gui<strong>an</strong>ensis <strong><strong>an</strong>d</strong> in subsp. venezuelensis the inner<br />
tellous, or on the smaller veins to puberulous, surface of the stipules is generally covered <strong>with</strong><br />
hairs on main veins often + appressed, white, (rather dense) yellow hairs, apparently correlated<br />
arachnoid hairs usually (almost) confined to the <strong>with</strong> the yellow-hirsute indument on the leafy<br />
areoles; tertiary venation <strong><strong>an</strong>d</strong> reticulum + prom- twigs, the outer surface of the stipules, <strong><strong>an</strong>d</strong> other<br />
inent (<strong><strong>an</strong>d</strong> conspicuous) beneath; lateral veins parts. <strong>Pourouma</strong> bicolor (except for subsp. tess-<br />
(7-)12-26 pairs, basal pair br<strong>an</strong>ched; petiole 4- m<strong>an</strong>nii.) is likewise characterized by the occur-<br />
25(-40) cm long, whitish-yellow-puberulous to rence of dense, whitish to yellow hairs on the<br />
-hirtellous to -hirsute <strong><strong>an</strong>d</strong> <strong>with</strong> dense to (very) inner surface of the stipules, but in this species<br />
sparse, brown, pluricellular hairs; stipules (2-)4- (except for subsp. choco<strong>an</strong>a.) this indument co-<br />
15 cm long, caducous, outside whitish- (to pale occurs <strong>with</strong> <strong>an</strong> indument on the leafy twigs, petyellow-)subsericeous<br />
to -subhirsute to -subto- ioles, <strong><strong>an</strong>d</strong> outer surface of the stipules consisting<br />
mentose or to -subvelutinous or yellow-hirsute, of short, appressed hairs.<br />
<strong><strong>an</strong>d</strong> <strong>with</strong> dense to (very) sparse, brown, pluri- (b.) The indument on the smaller veins of the<br />
cellular hairs, inside glabrous or <strong>with</strong> sparse, white lamina beneath. In P. gui<strong>an</strong>ensis this consists of<br />
hairs or <strong>with</strong> rather dense, yellow hairs. Stami- relatively long, patent hairs. In P. bicolor the<br />
nate inflorescences up to 20 cm long <strong><strong>an</strong>d</strong> 12 cm hairs on both the main veins <strong><strong>an</strong>d</strong> the smaller<br />
wide; peduncle 4-6 cm long, peduncle <strong><strong>an</strong>d</strong> veins is mostly appressed, only occasionally pabr<strong>an</strong>ches<br />
puberulous to hirtellous to (sub)hirsute tent on the smaller veins <strong><strong>an</strong>d</strong> then mostly shorter<br />
or to subvelutinous, often <strong>with</strong> dense, brown, th<strong>an</strong> in P. gui<strong>an</strong>ensis. Only in several specimens<br />
pluricellular hairs; flowers + clustered, but not of P. bicolor subsp. scobina does the indument<br />
in (sub)globose heads; tepals l<strong>an</strong>ceolate, (almost) resemble that normally occurring in P. gui<strong>an</strong>enfree;<br />
filaments shorter th<strong>an</strong> the tepals. Pistillate sis.<br />
inflorescences in fruit up to 20 cm long <strong><strong>an</strong>d</strong> 12 (c.) The prominence of the reticulum <strong><strong>an</strong>d</strong> occm<br />
wide; peduncle (3-)6-15(-20) cm long, in- currence of the white-arachnoid indument on the<br />
dument of the peduncle <strong><strong>an</strong>d</strong> br<strong>an</strong>ches similar to lamina beneath. In P. gui<strong>an</strong>ensis the reticulum<br />
that of the staminate inflorescence; flowers is prominent <strong><strong>an</strong>d</strong>, moreover, conspicuous as the
<strong>Pourouma</strong> 123<br />
white-arachnoid indument is usually confined to<br />
the areoles. In P. bicolor the reticulum is, in most<br />
of the subspecies, merely slightly prominent to<br />
pl<strong>an</strong>e <strong><strong>an</strong>d</strong> usually inconspicuous as the arachnoid<br />
indument usually also occurs on the smaller veins.<br />
However, in a form of P. bicolor subsp. bicolor,<br />
occurring in the western part of the r<strong>an</strong>ge of the<br />
subspecies, the arachnoid indument is confined<br />
to the very small areoles surrounded by relatively<br />
thick veinlets, <strong><strong>an</strong>d</strong> in P. bicolor subsp. scobina<br />
the arachnoid indument is also more or less confined<br />
to the areoles, but in this subspecies this<br />
feature is combined <strong>with</strong> ? prominent smaller<br />
veins.<br />
In P. gui<strong>an</strong>ensis two allopatric subspecies c<strong>an</strong><br />
be recognized.<br />
Key to the Subspecies<br />
of <strong>Pourouma</strong> gui<strong>an</strong>ensis<br />
1. Lamina 3-5-fid to -parted; leafy twigs <strong><strong>an</strong>d</strong> petioles<br />
<strong>with</strong> long yellow hairs; stipules hairy inside;<br />
fruiting peri<strong>an</strong>th 2-2.5 cm long; northern Venezuela<br />
................ b. subsp. venezuelensis.<br />
1. Lamina entire, 5-7-fid to -parted, or if 3-5-fid<br />
to -parted, then the leafy twigs <strong><strong>an</strong>d</strong> the petioles<br />
<strong>with</strong>out long yellow hairs, the stipules glabrous<br />
inside, <strong><strong>an</strong>d</strong>/or the fruiting peri<strong>an</strong>th 1.2-2 cm long;<br />
Gui<strong>an</strong>a region, eastern Brazil, Amazon Basin to<br />
eastern Colombia. ........ a. subsp. gui<strong>an</strong>ensis.<br />
la. <strong>Pourouma</strong> gui<strong>an</strong>ensis Aublet subsp.<br />
gui<strong>an</strong>ensis. Fig. 59.<br />
<strong>Pourouma</strong> palmata Poeppig & Endlicher, Nov. gen.<br />
sp. pi. 2: 29, t. 141. 1838; Tr6cul, Ann. Sci. Nat.<br />
Bot., Ser. 3, 8: 104. 1847; Miquel in Martius, Fl.<br />
bras. 4(1): 126. 1853; Macbride, Publ. Field Mus.<br />
Nat. Hist., Bot. Ser., 13(2.2): 293. 1937. Type. Peru.<br />
S<strong>an</strong> Martin: Tocache, Aug 1830 (6 <strong><strong>an</strong>d</strong> 9), Poeppig<br />
s.n. or 1881 (holotype or syntypes, W, destroyed;<br />
lectotype, G (Q specimen!), fragment of 6 specimen<br />
in F).<br />
<strong>Pourouma</strong> acutiflora Trecul, Ann. Sci. Nat. Bot., S&r.<br />
3, 8: 105. 1847; Miquel in Martius, Fl. bras. 4(1):<br />
126. 1853. Type. Brazil. Rio de J<strong>an</strong>eiro: Without<br />
locality, 1839 (6), Guillemin 1024 (holotype, P).<br />
<strong>Pourouma</strong> cinerascens Martius ex Miquel in Martius,<br />
<strong>Pourouma</strong> scabra Rusby, Bull. New York Bot. Gard.<br />
6: 498. 1910. Type. Bolivia. P<strong><strong>an</strong>d</strong>o: S<strong>an</strong>ta Barbara,<br />
30 Aug 1902 (9), R. S. Williams 1560 (holotype, NY;<br />
isotype, US).<br />
<strong>Pourouma</strong> radula Benoist, Bull. Mus. Hist. Nat. (Paris)<br />
28: 320. 1922; Woodson & Schery, Ann. Missouri<br />
Bot. Gard. 47: 166, t. 58. 1960. Type. Colombia.<br />
Without locality, <strong>with</strong>out date (a), Tri<strong>an</strong>a 860 (ho-<br />
lotype, P; isotypes, E, NY).<br />
<strong>Pourouma</strong> subtriloba Rusby, Mem. New York Bot.<br />
Gard. 7: 232. 1927. Type. Bolivia. La Paz: Nr. Tu-<br />
mapasa, 12 Oct 1921 (9), Cardenas 1990 (holotype,<br />
NY; isotypes, GH, US).<br />
<strong>Pourouma</strong> substrigosa Mildbraed, Notizbl. Bot. Gart.<br />
Berlin-Dahlem 10:192. 1927; Macbride, Publ. Field<br />
Mus. Nat. Hist., Bot. Ser., 13(2.2): 293. 1937. Type.<br />
Peru. Loreto: Pongo de M<strong>an</strong>seriche, 26 Nov 1924<br />
(8), Tessm<strong>an</strong>n 4642 (holotype, B, isotypes, B, G,<br />
NY).<br />
<strong>Pourouma</strong> mildbraedi<strong>an</strong>a St<strong><strong>an</strong>d</strong>ley, Publ. Field Mus.<br />
Nat. Hist., Bot. Ser., 17: 183. 1937. Type. Brazil.<br />
Amazonas: Mun. Sao Paulo de Olivenoa, nr. Pal-<br />
mares, 11 Sep-26 Oct 1936 (a), Krukoff 8386 (ho-<br />
lotype, NY; isotypes, A, BR, F, G, LE, MO, P, S, U,<br />
US).<br />
Leafy twigs pale to bright yellow-hirtellous to<br />
-subvelutinous to -hirsute, brown, pluricellular<br />
hairs dense to sparse. Lamina entire, 3-5-lobed<br />
to -parted, or sometimes 5-7-parted, base shal-<br />
lowly to deeply cordate or sometimes rounded;<br />
lateral veins mostly 10-16, sometimes up to 20,<br />
or occasionally up to 25 pairs; stipules outside<br />
pale to bright yellow-hirtellous, -subvelutinous,<br />
-subtomentose or -hirsute, inside glabrous or<br />
sometimes hairy. Fruitingperi<strong>an</strong>th 1.2-2 cm long,<br />
mostly densely hairy, sometimes sparsely hairy<br />
or scabrous.<br />
Distribution (Fig. 60). Throughout the Ama-<br />
zon Basin, extending to eastern Colombia <strong><strong>an</strong>d</strong><br />
the Gui<strong>an</strong>a region; occurring disjunctly in east-<br />
ern Brazil (from Pernambuco to S<strong>an</strong>ta Catarina);<br />
mostly in non-inundated forest, sometimes in<br />
periodically inundated (varzea) forest; mostly at<br />
low altitudes, in Venezuela (Bolivar) up to ca.<br />
1300 m, in Bolivia up to 1500 m.<br />
Specimens examined. Colombia. Without locality,<br />
<strong>with</strong>out date (d), Tri<strong>an</strong>a 860 (E, NY, P, type collection<br />
Fl. bras. 4(1): 125, t. 37. 1853. Type. Brazil. Ama- of P. radula), <strong>with</strong>out date (6 <strong><strong>an</strong>d</strong> 9), Tri<strong>an</strong>a s.n. (E, P).<br />
zonas: Rio Japura, Maripi, <strong>with</strong>out date (6), Martius META: Puerto Lopez, 3 Aug 1944 (st), Liitle et al. 8423<br />
s.n. (holotype, M; isotypes, M, U).<br />
(F); Villavicencio, Ll<strong>an</strong>o de S<strong>an</strong> Martin, <strong>with</strong>out date<br />
<strong>Pourouma</strong> heterophylla Martius ex Miquel in Martius, (9), Karsten s.n. (GOET, LE).<br />
Fl. bras. 4(1): 125. 1853. Type. Brazil. Amazonas: VENEZUELA. AMAZONAS: Rio M<strong>an</strong>avicke, Indi<strong>an</strong><br />
Rio Japura, Dec 1819 (st, juv), Martius s.n. (holo- village "Kalchi-Teri," <strong>with</strong>out date (juv.), Lizot s.n.<br />
type, M; isotypes, M, U).<br />
(VEN); Sierra Parima, 1300 m, 18-23 May 1972 (st),<br />
<strong>Pourouma</strong> fulginea Miquel in Martius, Fl. bras. 4(1): Steyermark 106086 (NY); Parima Mts., nr. Simara-<br />
129. 1853. Type. Brazil. Amazon Basin, <strong>with</strong>out date wochi, Rio Matacuni, 18 Apr-23 May 1973 (d), Stey-<br />
(Q), Martius s.n. (holotype, M; isotype, U). ermark 107154 (F, U, VEN). BOLIVAR: Reserva Fo-
124 Flora Neotropica<br />
JII'd~~~~~~~i<br />
_ ,e<br />
4~, ,. \<br />
as t af +<br />
POUROUMA einerascenx
<strong>Pourouma</strong>125<br />
restal "La Paragua," Rio Asa, Jun 1970 (st,juv), Bl<strong>an</strong>co<br />
810 (VEN); slopes of Quebrada O-paru-ma, between<br />
S<strong>an</strong>ta Teresita de Kav<strong>an</strong>ayen <strong><strong>an</strong>d</strong> Rio Pacairao (tributary<br />
of Rio Mouak), alt. 1065-1220 m, 20-21 Nov<br />
1944 (2), Steyermark 60407 (F, VEN); 45 km N of<br />
Tumeremo, Sierra de Nuria, 5-8 Feb 1961 (st), Steyermark<br />
89134 (VEN); Rio Nichare (tributary of Rio<br />
Caura), nr. the confluence <strong>with</strong> Rio Cicuta, 25 Apr<br />
1966 (2), Steyermark et al. 95711 (VEN). DELTA<br />
AMACURO: E of Rio Gr<strong><strong>an</strong>d</strong>e, ENE of El Palmar, 29<br />
Nov-18 Dec 1964 (2), Marc<strong>an</strong>o Berti 471 (NY, VEN),<br />
19 J<strong>an</strong> 1965 (2), Marc<strong>an</strong>o Berti 611 (F, NY, VEN); Rio<br />
(9), Huashikat 1216 (BG); Rio Cenepa, 30 J<strong>an</strong> 1973<br />
(6), Kayap 268 (F, GH, MO), 21 Feb 1973 (9), Kayap<br />
393 (F, GH); Quebrada Huampami, 5 Jul 1974 (9),<br />
Kayap 1057 (F, GH, MO). HuANuco: Tingo Maria,<br />
13 Aug 1940 (9), Asplund 13020 (G, A, S, US); Pachita,<br />
Codo de Pozuzo, 24 Oct 1982 (6), Foster 9373 (BG);<br />
Distr. Churubamba, Rio Cayumba, 11 Sep 1936 (9),<br />
Mexia 8172 (F, G, GB, GH, MO, NA, NY, S, U, UC,<br />
US); between Monz6n <strong><strong>an</strong>d</strong> Rio Huallaga, <strong>with</strong>out date<br />
(9), Weberbauer 3639 (G). LORETO: Prov. Coronel Portillo,<br />
Dist. Calleria, rd. Pucallpa-Hu<strong>an</strong>uco, km 34, 23<br />
May 1968 (6), Castillo S. 11 (DUKE, F, P, US); Que-<br />
Amacuro, Venezuela-Guy<strong>an</strong>a frontier, Imataca Mts., brada Sh<strong>an</strong>uce, above Yurimaguas, 11 Jul 1972 (2),<br />
downstream from S<strong>an</strong> Victor, 4 Nov 1960 (st), Stey- Croat 18016 (F, MO, NY); Rio Napo, Caserio Bella<br />
ermark 87307 (NY, VEN).<br />
Vista, 22 Sep 1972 (6), Croat 20161 (C, DUKE, F, GH,<br />
GUYANA. Kartabo region, Kartabo, 22 Jul 1920 NA, NY); Rio Ucayali, Jenaro Herrera, 7 Dec 1977<br />
(st, juv) Bailey 18 (GH), Bailey 19 (GH), Bailey 20 (st), Gentry et al. 21205 (MO, U); Prov. Maynas, Y<strong>an</strong>a-<br />
(GH), 20 Aug 1920 (st), Bailey 146 (GH), Bailey 147 mono Explorama Tourist Camp, 15 Jul 1983 (st), Gen-<br />
(GH); Pomeroon district, Kamwatta, 21 Sep 1921 (st), try et al. 43066 (BG); Pongo de M<strong>an</strong>seriche, nr. mouth<br />
Cruz 1175 (NY); between Demarara R. <strong><strong>an</strong>d</strong> Berbice of Rio S<strong>an</strong>tiago, 2 Dec 1931 (d), Mexia 6201 (F, G,<br />
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),<br />
Kurumaikabra Creek, Essequibo R., 22 Nov 1930 (2), Tessm<strong>an</strong>n 4642 (B, G, NY, type collection of P. sub-<br />
FD 1011 (FHO); Takutu Creek to Puruni R., Mazaruni strigosa). MADRE DE DIos: Parque Nacional del M<strong>an</strong>u,<br />
R., 25 Oct 1944 (6), F<strong>an</strong>shawe 2050 (=FD 4786) (A, Rio M<strong>an</strong>u, Cocha Cashu Station, Jul 1978 (st), Foster<br />
F, NY, U, US); Mazaruni station, forest nursery, 1945 et al. 6568 (F); Rio Tambopata, at mouth of Rio D'Or-<br />
(st, juv), FD 4970 (U); FD 5011 (U, S); K<strong>an</strong>uka Mts., bigny, 1 <strong><strong>an</strong>d</strong> 2 Mar 1981 (st), Gentry et al. 31816 <strong><strong>an</strong>d</strong><br />
Iramaip<strong>an</strong>g, Nov 1948 (6), FD 5936 (NY); 11/2 mile 31933 (U). PASCO: Prov. Oxapampa, Vivero, Puerto<br />
along Bartica-Potaro rd., <strong>with</strong>out date (juv), FD 6914 Bermudez, 16 J<strong>an</strong> 1983 (st), Gentry et al. 42029 (BG).<br />
(U); Tumatumari, 18 Jun-8 Jul 1921 (st), Gleason 174 SAN MARTIN: Tocache, Aug 1830 (6 <strong><strong>an</strong>d</strong> 9), Poeppig<br />
(GH, NY); Pakaraima Mts., just N of Paruima Mis- s.n. (or 1881) (F, G, type collection of P. palmata);<br />
sion, 19 Oct 1981 (9), Maas et al. 5858 (U); basin of Maynas Alto, <strong>with</strong>out date (9), Poeppig s.n. (B, OXF);<br />
Shodikar Creek, tributary of Essequibo R., 8-22 J<strong>an</strong> <strong>with</strong>out locality, <strong>with</strong>out date (6), Poeppig s.n. (LE, P);<br />
1938 (2), A. C. Smith 2845 (A, F, G, MO, NY, P, S, Distr. Tocache, rd. to Almendras, 7 Aug 1969 (2), J.<br />
U, US); Marudi Mts., 02?15'N, 59?10'W, 12 Nov 1982 Schunke V. 3309 (F, G, US); rd. to Shunthe, E ofPuente<br />
(2), Stoffers et al. 311, 312 (U).<br />
de Palo Bl<strong>an</strong>co, 14 Jul 1974 (9), J. Schunke V. 7415<br />
SURINAM. Raleigh falls, base of Voltzberg, Nature (MO, NY, U).<br />
Reserve on the Coppename R., 18 Nov 1979 (9), Mori BRAZIL. Without locality, 1839 (3), Guillemin 1024<br />
et al. 8662 (NY, U).<br />
(P, type collection of P. acutiflora); "Amazon Basin,"<br />
FRENCH GUIANA. Without locality, <strong>with</strong>out date <strong>with</strong>out locality, <strong>with</strong>out date (9), Martius s.n. (M, U,<br />
(2), Aublet s.n. (BM, LE, ?S, type collection of P. gui- type collection of P. fulginea). ACRE: Cruzeiro do Sul,<br />
<strong>an</strong>ensis); nr. Saul 3 Oct 1973 (9), Gr<strong>an</strong>ville et al. B.5071 21 Feb 1976 (9), Marinho 273 (IAN), 18 Feb 1976 (2),<br />
(U); Paul Isnard region, 5 km of Citron, 12 Nov 1982 Monteiro et al. 477 (INPA, MG); rd. Rio Br<strong>an</strong>co-Porto<br />
(2), Gr<strong>an</strong>ville 5284 (U); Oyapock R., Trois Sauts, 13 Acre, km 83, 10 Oct 1980 (9), Nelson 681 (BG)<br />
Jul 1774 (st), Gren<strong><strong>an</strong>d</strong> 445 (CAY); Zidock Ville, 26<br />
Aug 1976 (st), Gren<strong><strong>an</strong>d</strong> 1352 (CAY); Comte R., Cacao,<br />
25 Feb 1965 (2), Halle 1132 (U, P); nr. Sail, J<strong>an</strong> 1975<br />
(2), Moretti 102 (CAY); Comte R., ca. 60 km S of<br />
Cayenne, 26 Feb 1965 (9), Oldem<strong>an</strong> 1179 (CAY); Saiil,<br />
19 Oct 1971 (2), Oldem<strong>an</strong> B.4108 (CAY, P, U); Kourou,<br />
<strong>with</strong>out date (9), (herb.) Richard s.n. (P).<br />
ECUADOR. NAPO: Rio Pucino, first major tributary<br />
of Rio Aguarico, above bridge at Aguarico, 10 Feb<br />
1974 (2), Gentry 9824 (U); Rio Napo, 8 km below<br />
Puerto Mishualli, 25-27 Mar 1986 (2) Neill et al. 7082<br />
(BG).<br />
PERU. AMAZONAS: 6 Apr 1973 (2), Ancuash 178 (F,<br />
GH, MO); Rio S<strong>an</strong>tiago, nr. Caterpiza, 9 Nov 1979<br />
= Cid<br />
et al. 2870 (U); rd. Sena Madureira-Rio Br<strong>an</strong>co, nr.<br />
km 7, 30 Sep 1968 (6), Pr<strong>an</strong>ce et al. 7670 (GH, INPA,<br />
M, NY, R, S, U, US). AMAZONAS: Rio Antimari 30<br />
Feb 1904 (st), Huber (MG) 4244 (MG); Rio Madeira,<br />
nr. Calama, 7 Nov 1931 (9), Krukoff 1297 (G, NY, P,<br />
S, U); nr. mouth of Rio Embira, 14 J<strong>an</strong> 1933 (9), Krukoff4817<br />
(A, F, G, MO, NY, S, U, US); Mun. Humaita,<br />
Tres Casas, 14 Sep-11 Oct 1934 (o), Krukoff6238 (B,<br />
BR, F, LE, MO, NY, S, U, US); Sao Paulo de Olivenca,<br />
nr. Palmares 11 Sep-26 Oct 1936 (9), Krukoff8369 (A,<br />
B, BR, F, G, LE, NY, P, U), (6), Krukoff8386 (A, BR,<br />
F, G, LE, MO, NY, P, S, U, US, type collection of P.<br />
mildbraedi<strong>an</strong>a); Sao Paulo de Olivenca, Creek Belem,<br />
26 Nov-11 Dec 1936 (2), Krukoff8652 (A, BR, F, G,<br />
FIG. 59. <strong>Pourouma</strong> gui<strong>an</strong>ensis subsp. gui<strong>an</strong>ensis. From Martius, Flora Brasiliensis 4(1). 1853: tab. 37 (as<br />
P. cinerascens). Leafy twig <strong>with</strong> staminate inflorescence, leaf apex, part of staminate inflorescence (28), diagram<br />
of staminate flower (D), staminate flower (1).
~~~ II ~~~ I<br />
P. guiamnonsi<br />
^%^_ 1 ^vf'^^/9^^ ^^\?^^^/ v * \ subsp. quirnnisL<br />
---- ---------- - --<br />
^l -<br />
i~subap. vwwwcLensnir,<br />
- '--- - - p l<br />
FIG. 60. Distribution of <strong>Pourouma</strong> gui<strong>an</strong>ensis subsp. gui<strong>an</strong>ensis, P. gui<strong>an</strong>ensis subsp. venezuelenyis',<br />
Pourou-<br />
ma velutina.
<strong>Pourouma</strong>127<br />
LE, MO, NY, P, S, U, US); Mun. Coari, Rio Coari,<br />
20 Apr 1976 (st), Magnago et al. (INPA) 58046 (INPA);<br />
Rio Japura, Dec 1819 (st, juv), Martius s.n. (M, U,<br />
type collection ofP. heterophylla); Rio Japura, Mapiri,<br />
<strong>with</strong>out date (8), Martius s.n. (M, U, type collection of<br />
P. cinerascens); M<strong>an</strong>aus-Itacoatiara rd., km 27, Reserva<br />
Florestal Ducke, 30 Jun 1976 (st), Mello (INPA)<br />
57509 (INPA); km 26,6 J<strong>an</strong> 1977 (st), Nascimento 303<br />
(INPA); Rio C<strong>an</strong>uma, Mun. Nova Olinda do Norte,<br />
Nov 1976 (st), Monteiro 1295 (INPA); Rod. BR-174,<br />
Reserva Biologica do INPA, 28 Sep 1976 (st), Mota<br />
740 (INPA); Reserva Florestal Ducke, 11 J<strong>an</strong> 1976<br />
(st), Nascimento 387 (INPA), Nov 1972 (st), Rodrigues<br />
9219 (INPA); rd. Humaita to Labrea, km 80, between<br />
Rios Ipixuna <strong><strong>an</strong>d</strong> Itapar<strong>an</strong>a, 29 J<strong>an</strong> 1970 (9), Pr<strong>an</strong>ce<br />
et al. 3260 (F, INPA, NY, R, S, U, US); basin of Rio<br />
Demeni, nr. Totobi, 25 Feb 1969 (9), Pr<strong>an</strong>ce et al.<br />
10233 (C, G, INPA, MO, NY, R, S, U, US); M<strong>an</strong>aus-<br />
P6rto Velho rd. BR-319, km 175, between Rio Tup<strong>an</strong>a<br />
<strong><strong>an</strong>d</strong> Rio Igapo-aqu, 11 Oct 1974 (9), Pr<strong>an</strong>ce et al. 22803<br />
(INPA, MO, U). BAHIA: Agua Preta, 13 Oct 1937 (a),<br />
Bondar s.n. (GUA); Rio das Contas, 3 Oct 1919 (8),<br />
Curr<strong>an</strong> 19 (C, F, MO, NY, US); rd. from Camaca-<br />
C<strong>an</strong>avieiro, Apr 1965 (9), Magalhaes 733 (M); nr. Urucuca,<br />
4 Nov 1978 (9), Mori et al. 11046 (RB, U); Ilh6us,<br />
Sep 1821 (9 <strong><strong>an</strong>d</strong> d), Riedel 2 or s.n. (LE, mixed <strong>with</strong><br />
sterile specimen of P. mollis), 10 Nov 1944 (9), Velloso<br />
723 (R). CEARA: Without locality, <strong>with</strong>out date (st),<br />
Allemao et al. 446 (R); Serra de Baturit6, Sep 1897 (st),<br />
Huber 238 (MG). EspiRITO SANTO: 9 Aug 1965 (9),<br />
Belem 1471 (MG, NY, U); rd. BR-5 (BR-101), Morro<br />
D<strong>an</strong>tos, 8 Aug 1965 (9), L<strong>an</strong>na Sobrinho 1011 (FEE-<br />
MA, U); nr. Linhares, 1 Oct 1971 (6), T. S. S<strong>an</strong>tos<br />
2035 (MG, NY, U). MATO GROSSO: Rd. to Font<strong>an</strong>ilha,<br />
15 Sep 1976 (9), Gomes et al. 326 (INPA); Sarar6, 10<br />
Aug 1978 (9), Pires et al. 16547 (MG); Nuicleo de Humboldt,<br />
(9), Roth 36 (INPA); Rio Juruena, road to Aripu<strong>an</strong>a,<br />
km 5, 9 Jul 1977 (a), M. G. Silva et al. 3305<br />
(MG); Aripu<strong>an</strong>a, rd. to airport, 2 Jun 1979 ($), M. G.<br />
Silva et al. 4750 (MG). MINAS GERAIS: Viosa, 16 May<br />
1935 (9), Kuhlm<strong>an</strong>n 2075 (US), 1936 (9), Kuhlm<strong>an</strong>n<br />
(RB) 2200 (RB); Mun. Caratinga, Faz. Montes Claros,<br />
23 J<strong>an</strong> 1980 (9), Nsihimura 39 (FEEMA, GUA, U);<br />
Rio Novo, - da Divisa, 19 Dec 1964 (6), Hatschbach 12049 (F);<br />
above Antonia, 18 J<strong>an</strong> 1966 (9), Hatschbach et al. 13537<br />
(F, NY, P, RB, U, US); Mun. Guaraquecaba, Rio B<strong>an</strong><strong>an</strong>al,<br />
8 Dec 1970 (9), Hatschbach 25765 (C, MO, NA,<br />
S, UC); Rio do Costa, 9 Feb 1972 (9), Hatschbach<br />
29129 (NA). PERNAMBUCO: Quipapa, Eng. Brejino 22<br />
Mar 1967 (9), Andrade-Lima 67-4983 (F). Rio DE<br />
JANEIRO: Tingua, 1 Oct 1946 (6), Brade et al. 18619<br />
(GUA, RB, U); Palmeiras, 13 J<strong>an</strong> 1877 (st), Glaziou<br />
8935 (E, P); Serra dos Orgaos, Tabuinha, 21 Mar 1980<br />
(st), Glaziou 12173 (P, mixed <strong>with</strong> Cecropia glaziovii);<br />
Serra da Estrella, nr. M<strong><strong>an</strong>d</strong>ioca, 15 May 1880 (6), Glaziou<br />
12174 (C, E, P), <strong>with</strong>out date (6), Glaziou 20408<br />
(C, P); Matta do Teizeira Borges, nr. Gavea, 12 Dec<br />
1928 (9), Pessoal do Horto Florestal 650 (RB); Matta<br />
dos Tres Barros, nr. Gavea, 14 Mar 1927 (9), Pessoal<br />
do Horto Florestal 651 (RB). RONDONIA: Mineraao<br />
Campo Novo, 100 km SW of Ariquemes, 10 Oct 1979<br />
(9), Zarucchi 2744 (INPA, NY). RORAMA: Rio Catrim<strong>an</strong>i,<br />
13 Feb 1975 (9), Pires 15088 (IAN); Serra Tepequem,<br />
west facing slopes, 1200 m, 17 Feb 1967 (9),<br />
Pr<strong>an</strong>ce et al. 4438 (G, INPA, M, NY, P, R, S, U, US);<br />
Indi<strong>an</strong> trail Surucucu-Uaica, nr. Uaica airstrip, Rio<br />
Uraricoeira, 2 Mar 1971 (9), Pr<strong>an</strong>ce et al. 10836 (F,<br />
GH, INPA, M, NY, P, S, U); between Botamatatedi<br />
<strong><strong>an</strong>d</strong> Maita, 9 Feb 1971 (9), Pr<strong>an</strong>ce et al. 13579 (F,<br />
INPA, NY, U, US). SANTA CATARINA: Brusque, 15 Feb<br />
1951 (9), Klein 89 (US); nr. Blumeneau, 18 J<strong>an</strong> 1955<br />
(9), Klein 110 (NY, S, UC, US); Brusque, 15 Feb 1952<br />
(9), Veloso 89 (RB); nr. Blumeneau, Nov 1888 (6), Ule<br />
991 (F, HGB, P, US). SAO PAULO: Ubatuba, 8 J<strong>an</strong><br />
1985 (9), Gentry et al. 49346 (BG).<br />
BOLIVIA. LA PAZ: Nr. Tumapasa, 12 Oct 1921<br />
(9), Cardenas 1990 (GH, NY, US, type collection of P.<br />
subtriloba); 16 km N ofCarrasco, 1500 m, 31 Oct 1984<br />
(9), Solomon et al. 12655 (BG). PANDO: Mukden, Jun-<br />
Dec 1979 (9), Izawa 9 (U); Prov. N. Suarez, 20 Aug<br />
1978 (9), Mences 759 (MG); W b<strong>an</strong>k of Rio Madeira,<br />
nr. Abufia, 22 Jul 1970 (9), Pr<strong>an</strong>ce et al. 6253 (B, INPA,<br />
NY, S, U, UC, US), 19 Nov 1968 (9), Pr<strong>an</strong>ce et al.<br />
8670 (F, INPA, MG, NY, S, U); Prov. N. Suarez, S<strong>an</strong><br />
Pedro 13 Oct 1977 (9), Terceros 1400 (INPA); S<strong>an</strong>ta<br />
Barbara, 30 Aug 1902 (9), R. S. Williams 1560 (NY,<br />
(9), Schwacke 10393 (P). PARA: Serra US, type collection of P. scabra).<br />
dos Carajas, Serra Norte, ca. 20 km N of Amza Exploration<br />
camp, 17 Oct 1977 (9), Berg et al. 598 (MG, Vernacular names. Colombia. Meta: papa-<br />
U); Cuiaba-S<strong>an</strong>tarem rd., km 1183, 2 Feb 1977 (9), quillo. Venezuela. Bolivar: amia-yek; Delta<br />
Berg et al. 772 (F, MG, MO, RB, S, U); Breves, Oct- Amacuro: chaparro de agua. Guy<strong>an</strong>a. buruma,<br />
Nov 1957 (a), Pires et al. 6663 (U); Rio Itacaiunas,<br />
affluent of Rio Toc<strong>an</strong>tins, Serra Buritirama, Jul 1970 s<strong><strong>an</strong>d</strong>paper. Surinam. M<strong>an</strong>bospapaja. French<br />
(9), Pires et al. 12570 (IAN); Cap<strong>an</strong>ema-Mar<strong>an</strong>hao rd. Gui<strong>an</strong>a. Bois c<strong>an</strong>on, kuluma, kalate, (Wayapi).<br />
BR-22, km 96, 29 Oct 1965 (9), Pr<strong>an</strong>ce et al. 1777 (F, Peru. Amazonas: shuiya (Huambisa), sugkama(t)<br />
GH, NY, P, S, U, US); Gleba Bacaja, just below mouth shuiya, tsakap suiya (Huambisa); Hu<strong>an</strong>uco: paof<br />
Rio Bacaja 21 Nov 1980 (st), Pr<strong>an</strong>ce et al. 26375<br />
(U); S<strong>an</strong>tar6m, km 70 Estrada do<br />
paya del monte; Loreto: uvilla; S<strong>an</strong> Martin: uvil-<br />
Palhao, Ramal do<br />
Caetetu, 16 Sep 1969 (a), M. Silva et al. 2623 (G, NY, la, uvilla bl<strong>an</strong>ca. Brazil. Bahia: itarar<strong>an</strong>ga, tara-<br />
S, U, US). PARANA: P6rto de la Cima, 17 Jul 1911 (st), r<strong>an</strong>ga bl<strong>an</strong>ca; Mar<strong>an</strong>hao: kaymbe'y (Ka'apor);<br />
Dusen 11942 (S); Jacarehy (=Sao Joao), 8 Oct 1915 Mato Grosso; imbaubar<strong>an</strong>a, imbauba-torem;<br />
(st), Duskn 17239 (F, NY, S), 22 Nov 1915 (9 <strong><strong>an</strong>d</strong> 6), Para: amapati or mapati; Par<strong>an</strong>a: imbaubar<strong>an</strong>a,<br />
Dusen' 17345 (F, GH, NY, S); Morretes, 1904 (st),<br />
Dusen s.n. (S); 14 Aug 1911 (st), Dusen s.n. pau de jacu; Pernambuco: embauba da mata.<br />
(F, GH,<br />
NY, S); Mun. Guaratuba, Pedra Br<strong>an</strong>ca do Araraquara,<br />
This subspecies is very variable in the density<br />
3 Nov 1960 (9), Hatschbach 7411 (R, U, US), 3 Nov <strong><strong>an</strong>d</strong> length of the indument. The material from<br />
1960 (6), Hatschbach 7466 (US); Mun. Guarataba, Rio eastern Brazil often has 5-fid leaves, the indu-
128 Flora Neotropica<br />
ment is rather short, <strong><strong>an</strong>d</strong> the leaf base is deeply tinct forms of the subspecies in the same area<br />
to shallowly cordate. Some collections from the <strong><strong>an</strong>d</strong> the absence of intermediates between these<br />
southern part of the Amazon Basin (Bolivia <strong><strong>an</strong>d</strong> forms.<br />
adjacent parts of Brazil) usually have 5-7-fid to<br />
-parted leaves, while elsewhere in the Amazon<br />
Basin the leaves are 3-lobed to -fid or entire.<br />
Most collections from the Gui<strong>an</strong>a region <strong><strong>an</strong>d</strong> the<br />
Lower Amazon Basin have 3-fid leaves <strong>with</strong> a<br />
shallowly cordate to truncate leaf base or entire<br />
leaves; the indument is short, <strong><strong>an</strong>d</strong> the brown,<br />
pluricellular hairs on the twigs are usually dense.<br />
Material from the Upper Amazon Basin is<br />
much more variable. Some of the collections have<br />
almost elliptic (to oblong) entire leaves. Another<br />
group of collections have entire leaves (often <strong>with</strong><br />
relatively short petioles) or 3-fid to -parted,<br />
sometimes 5-parted leaves <strong>with</strong> the lobes rather<br />
close together, due to a smaller <strong>an</strong>gle in which<br />
the main basal lateral veins depart from the midrib.<br />
These collections often have sparse pluricellular<br />
hairs on the leafy twigs, <strong><strong>an</strong>d</strong> those <strong>with</strong><br />
entire leaves resemble P. velutina, from which<br />
they differ in a greater number of lateral veins.<br />
A third group of specimens have 3-fid to -parted<br />
leaves <strong>with</strong> a deeply cordate leaf base, often <strong>with</strong><br />
overlapping lobes. Several of these collections<br />
have relatively long yellow hairs on the leafy<br />
twigs <strong><strong>an</strong>d</strong> stipules, or have a rather densely hirtellous<br />
upper leaf surface. The collections <strong>with</strong><br />
long, yellow hairs usually have stipules in which<br />
the inner surface is rather densely hairy, unlike<br />
the other collections of subsp. gui<strong>an</strong>ensis which<br />
have the inner surface of the stipules glabrous or<br />
sometimes <strong>with</strong> very sparse, white hairs. Three<br />
collections (Croat 20616, Krukoff4817 <strong><strong>an</strong>d</strong> 8369)<br />
have 3-fid, deeply cordate leaves <strong><strong>an</strong>d</strong> long, yellow<br />
hairs on various parts, including the inner surface<br />
of the stipules <strong><strong>an</strong>d</strong> more numerous lateral veins<br />
(up to 25 pairs), in medium-sized leaves less th<strong>an</strong><br />
1 cm from each other. Other collections ofsubsp.<br />
gui<strong>an</strong>ensis (<strong><strong>an</strong>d</strong> collections of subsp. venezuelensis)<br />
have the lateral veins more th<strong>an</strong> 1 cm from<br />
each other.<br />
The material examined does not provide clear<br />
indications about the nature of the various forms<br />
<strong>with</strong>in the subspecies no more th<strong>an</strong> about the<br />
distribution of these forms. Material received<br />
from <strong>an</strong> inventory of a forest in the Upper Rio<br />
Moa region (Brazil, Acre) by Campbell <strong><strong>an</strong>d</strong> collaborators<br />
indicates the presence of several dis-<br />
In several specimens from the Upper Amazon<br />
Basin the leafy twigs, petiole, stipules, <strong><strong>an</strong>d</strong> lower<br />
surface of the lamina are covered by a white<br />
layer, actually the mycelium of some fungus which<br />
apparently uses the pluricellular hairs as a substrate.<br />
lb. <strong>Pourouma</strong> gui<strong>an</strong>ensis Aublet subsp. venezuelensis<br />
(Cuatrecasas) C. C. Berg & v<strong>an</strong> Heusden,<br />
Proc. Kon. Ned. Akad. Wetensch., Ser.<br />
C. 91(2): 106. 1988. Fig. 61.<br />
<strong>Pourouma</strong> venezuelensis Cuatrecasas, Bol. Soc. Venez.<br />
Ci. Nat. 15: 107. 1954. Type. Venezuela. Aragua:<br />
Parque Nacional Henri Pittier, 20 Nov 1953 (6),<br />
Aristeguieta 2005 (holotype, F, not seen; isotype,<br />
VEN).<br />
Leafy twigs yellow-hirsute, brown, pluricellular<br />
hairs dense. Lamina 3-5-fid to -parted, base<br />
? deeply cordate; lateral veins 16-26 pairs; stipules<br />
outside yellow-hirsute, inside <strong>with</strong> yellow<br />
hairs. Fruiting peri<strong>an</strong>th 2-2.5 cm long.<br />
Distribution (Fig. 60). Northern Venezuela<br />
(Aragua, Carabobo, Mir<strong><strong>an</strong>d</strong>a, <strong><strong>an</strong>d</strong> Yaracuy),<br />
coastal mountains; in forest at altitudes between<br />
ca. 1000 <strong><strong>an</strong>d</strong> 1200 m.<br />
Specimens examined. VENEZUELA. ARAGUA: Parque<br />
Nacional "H. Pittier," 20 Nov 1953 (6), Aristeguieta<br />
2005 (VEN, type collection of P. venezuelensis),<br />
12 Mar 1964 (6), Ijjdsz-Madriz 28 (VEN), 24 Dec 1946<br />
(9), Lasser 2202 (VEN); R<strong>an</strong>cho Gr<strong><strong>an</strong>d</strong>e, 30 Dec 1946<br />
(6), Lasser 2267 (VEN); Guamitas, 5 Nov 1947 (6),<br />
Pittier et al. 15641 (US, VEN); Parque Nacional "H.<br />
Pittier," 6 J<strong>an</strong> 1968 (st), R. F. Smith V-3321 (VEN);<br />
Chorni, 39 Apr-1 May 1972 (2), Steyermark et al.<br />
105863 (NY, VEN); Agua Amarilla, 12 Nov 1947 (6),<br />
Tamayo 3370 (VEN), R<strong>an</strong>cho Gr<strong><strong>an</strong>d</strong>e, 26 Mar 1938<br />
(9), LI. Williams 9983 (A, F, GH, NY, US, VEN).<br />
CARABOBO: Between Valencia <strong><strong>an</strong>d</strong> Camp<strong>an</strong>ero, 7 Mar<br />
1860 (2), Fendler 2420 (G); Cumbre de Valencia, Pt.<br />
Cabello, <strong>with</strong>out date (6), Karsten s.n. (LE); E of Los<br />
T<strong>an</strong>gues, S of Borburata, 31 Mar 1966 (9). Steyermark<br />
et al. 95378 (NY, VEN, U). MIRANDA: Guatopo, 19<br />
Nov 1956 (9), Bernardi 5696 (VEN), Bernardi s.n. (NY);<br />
S<strong>an</strong> Ju<strong>an</strong>, 18 km SW of Cupira, 2-7 Sep 1977 (6),<br />
Gonzalez et al. 1319 (BG); S of S<strong>an</strong>ta Cruz, Mar 1978<br />
(2), Steyermark et al. 116884 (BG); 11 km SSE of El<br />
Guapo, 27-28 Mar 1978 (2), Steyermark et al. 116966<br />
(U). YARACUY: N of Salom, 4 Mar 1982 (st), Liesner<br />
et al. 12394 (U).
<strong>Pourouma</strong><br />
~ I I<br />
MINISTERIO DE AGRICULTUPA V CRIA<br />
IERDBAJiO NACIONAL DF VENEZUFLA<br />
FIG. 61. <strong>Pourouma</strong> gu<strong>an</strong>enss Abl subsp. Leafy t wig <strong>with</strong> pistillate inflorescences (Steyermark et<br />
ial. 105863).sden tr: a<br />
D Ce.: C,C, Blrg<br />
E,C.I. v<strong>an</strong> in- He usden<br />
Utrecht r'.<br />
...X'lr , A. R I%, ? %t<br />
,,,. J.&M<br />
a.<br />
Cm"t<br />
FIG. 61. <strong>Pourouma</strong> gui<strong>an</strong>ensis subsp. venezuelensis. Leafy twig <strong>with</strong> pistillate inflorescences (Steyermark et<br />
al. 105863).<br />
Vernacular names. Venezuela. Carabobo: tambor;<br />
Mir<strong><strong>an</strong>d</strong>a: yagrumo negro.<br />
This subspecies is very uniform compared <strong>with</strong><br />
subsp. gui<strong>an</strong>ensis. Some collections of the latter<br />
subspecies from the Upper Amazon Basin resemble<br />
subsp. venezuelensis, but are distin-<br />
129<br />
guished by smaller leaves <strong><strong>an</strong>d</strong> smaller fruiting<br />
peri<strong>an</strong>ths.<br />
2. <strong>Pourouma</strong><br />
velutina Martius ex Miquel in Mar-<br />
tius, Fl. bras. 4(1): 130, t. 41. 1853; Berg, Fl.<br />
Suriname 5(1): 269. 1975. Type. Brazil. Para:
130<br />
Flora Neotropica<br />
Mouth of Rio Toc<strong>an</strong>tins, Aug 1819 (6), Mar- widened apex; peri<strong>an</strong>th 4-6 mm long, yellowishtius<br />
s.n. (lectotype, M, chosen here; isolecto- velutinous, at the apex <strong>with</strong> brown, pluricellular<br />
type, M).<br />
Fig. 62. hairs; stigma peltate, 1.5-3 mm diam., <strong>with</strong><br />
sparse, yellow hairs <strong><strong>an</strong>d</strong> dense, brown, pluricel-<br />
<strong>Pourouma</strong> steyermarkii St<strong><strong>an</strong>d</strong>ley & Cuatrecasas in lular hairs. Fruiting peri<strong>an</strong>th black, ovoid to el-<br />
Cuatrecasas, Fieldi<strong>an</strong>a Bot. 28(1): 210. 1951. Type.<br />
Venezuela. Bolivar: Ptari-tepui, between Rio<br />
lipsoid or sometimes to<br />
Karuai<br />
oblongoid, 1.2-1.8 x<br />
<strong><strong>an</strong>d</strong> first ridge above Rio Karuai, 28 Nov 1944<br />
0.6-1<br />
(v),<br />
cm, densely to sparsely hairy.<br />
Steyermark 60665 (holotype, F; isotype, VEN). Distribution (Fig. 60). Venezuela (Amazonas<br />
<strong><strong>an</strong>d</strong> Bolivar), Surinam, French Gui<strong>an</strong>a, Ama-<br />
Tree, up to 15 m tall. Leafy twigs 2-6 mm zoni<strong>an</strong> Brazil (Amapa, Amazonas, Mato Grosso,<br />
thick, brownish- to yellowish- to whitish-seri- <strong><strong>an</strong>d</strong> Para) <strong><strong>an</strong>d</strong> Peru (Loreto), <strong><strong>an</strong>d</strong> eastern Brazil<br />
ceous to -puberulous <strong><strong>an</strong>d</strong> sometimes <strong>with</strong> brown, (Bahia <strong><strong>an</strong>d</strong> Espirito S<strong>an</strong>to); in non-inundated<br />
arachnoid hairs. Lamina entire, elliptic to ovate, forest, mostly at low altitudes, in Venezuela up<br />
rarely to obovate, 6-30 x 2-20 cm, coriaceous to ca. 1200 m.<br />
to subcoriaceous or sometimes chartaceous, apex<br />
acuminate, base rounded to acute, sometimes Specimens examined. VENEZUELA. AMAZONAS:<br />
truncate or<br />
Atures, 20<br />
subcordate; upper surface<br />
Apr 1978 (9), Davidse et al. 15365 (MO,<br />
scabrous, U); Cam<strong>an</strong>i, 24 Apr 1971 (9), Foldats 114-1A (NY,<br />
the main veins brown-tomentose to -sericeous, VEN). BOLIVAR: Lower S-facing slopes of Ptari-tepui<br />
sometimes <strong>with</strong> or<strong>an</strong>ge or or<strong>an</strong>ge-brown, pluri- between Rio Karuai <strong><strong>an</strong>d</strong> first ridge above Rio Karuai,<br />
cellular hairs; lower surface whitish-sericeous to ca. 1200 m, 28 Nov 1944 (9) Steyermark 60665 (F,<br />
-velutinous, sometimes <strong>with</strong> VEN, type collection of P. steyermarkii).<br />
brown, pluricellular SURINAM. Sectie O, 11 Nov 1915 (st), BW 1368<br />
hairs, white arachnoid hairs in the areoles <strong><strong>an</strong>d</strong> (U); Tibiti sav<strong>an</strong>na, 20 J<strong>an</strong> 1949 (st), L<strong>an</strong>jouw et al.<br />
on the smaller veins; lateral veins 7-13 pairs, 1761 (NY, U); Jodensav<strong>an</strong>ne, Map<strong>an</strong>e Creek, 13 Jun<br />
basal pair br<strong>an</strong>ched or unbr<strong>an</strong>ched, tertiary ve- 1953 (st), Lindem<strong>an</strong> 4057 (U); 1 Oct 1953 (st), Lindenation<br />
prominent beneath; petiole 1.5-20 cm m<strong>an</strong> 4794 (U); Upper Coppename R., 6 Aug 1954 (st),<br />
Lindem<strong>an</strong> 6418<br />
long, yellow- to brown-sericeous to<br />
(U).<br />
-tomentose, FRENCH GUIANA. St. Laurent, 10 Oct 1956 (9),<br />
<strong><strong>an</strong>d</strong> sometimes <strong>with</strong> brown, pluricellular hairs; BAFOG 7562 (CAY, NY, P, U); Gregoire, 26 J<strong>an</strong> 1972<br />
stipules 2-12 cm long, caducous, outside yellow- (9), Deward 138 (CAY, P, U); Maroni R., 1891 (6),<br />
sericeous to -velutinous, <strong><strong>an</strong>d</strong> often <strong>with</strong> brown G<strong><strong>an</strong>d</strong>oger 19 (P); Orapu, 26 Oct 1970 (9), Oldem<strong>an</strong><br />
B.3786A<br />
to yellow, pluricellular hairs, inside (A, CAY, P, U); Acarou<strong>an</strong>y, 1858 ($),<br />
yellow-se-<br />
Sagot<br />
1163 (BR, F, G, GH, GOET, NY, P, S, U); Godebert,<br />
riceous to -velutinous. Staminate inflorescences 20 Dec 1920 (6), Wachenheim 271 (P).<br />
up to 12 cm long <strong><strong>an</strong>d</strong> 8 cm wide; peduncle 2-7 PERU. LORETo: Rio N<strong>an</strong>ay, trail from Astoria to<br />
cm long, peduncle <strong><strong>an</strong>d</strong> br<strong>an</strong>ches yellowish-seri- Rio Maz<strong>an</strong>, 9 J<strong>an</strong> 1976 (9), McD<strong>an</strong>iel et al. 20403<br />
ceous to -velutinous or to -tomentose, <strong><strong>an</strong>d</strong> often (NA).<br />
BRAZIL. AMAPA: Mun.<br />
<strong>with</strong> Mazagao, 75-80 km WSW<br />
dense, brown, pluricellular hairs; flowers of Macapa, 5-10 km SW of Rio Prato, 20 Dec 1984<br />
sessile, diffusely distributed along the ultimate (9), Daly et al. 3936 (BG); Clevel<strong><strong>an</strong>d</strong>ia, 2 Aug 1960 (6),<br />
br<strong>an</strong>ches; tepals 1.3-1.8 mm long, free or basally Westra 47303 (FHO, GH, MG, NY, U, UC, US).<br />
connate, white-puberulous, filaments shorter th<strong>an</strong> AMAZONAS: M<strong>an</strong>aus, 9 Aug 1962 (6), Chagas (INPA)<br />
the peri<strong>an</strong>th, free. Pistillate inflorescences in fruit<br />
1655 (INPA, U); Rio Uatuma, Itapir<strong>an</strong>ga, 27 Aug 1979<br />
(9), Cid 849<br />
up to 12 cm long <strong><strong>an</strong>d</strong> 7<br />
(U); M<strong>an</strong>aus, 8 Jul 1933 (9 <strong><strong>an</strong>d</strong> 6), Ducke<br />
cm wide; peduncle 2.5-<br />
(RB) 25247 (INPA, RB); M<strong>an</strong>aus-Itacoatiara rd., km<br />
7 cm long, peduncle <strong><strong>an</strong>d</strong> br<strong>an</strong>ches yellowish- to 26, 4 J<strong>an</strong> 1977 (st), Nascimento 262 (INPA); track from<br />
brownish-sericeous to -velutinous, sometimes to Boca de Acre airstrip to Monte Verde, 21 Sep 1966<br />
-tomentose or to -puberulous, <strong><strong>an</strong>d</strong> <strong>with</strong> dense, (9), Pr<strong>an</strong>ce et al. 2469 (INPA, MO, NY, R, S, U, UC,<br />
brown, pluricellular hairs; flowers 2-8, + dis- US); M<strong>an</strong>aus, Aleixo rd., km 11, 2 May 1974 (6), Pr<strong>an</strong>ce<br />
et al. 21011 (INPA, MG, MO, NY, S, U, US); M<strong>an</strong>aus,<br />
tinctly in two clusters; pedicel up to 0.4 cm long, Binda Creek rd., 26 Jun 1961 (9 <strong><strong>an</strong>d</strong> 6), Rodrigues et<br />
in fruit up to 1 cm, already at <strong>an</strong>thesis <strong>with</strong> a al. 2892 (INPA as 2092, MG, RB, U); M<strong>an</strong>aus Pas-<br />
FIG. 62. <strong>Pourouma</strong> velutina. From Martius, Flora Brasiliensis 4(1). 1853: tab. 41. Leafy twig <strong>with</strong> staminate<br />
inflorescences, diagram of staminate flower (D), part of staminate inflorescence (28), staminate flower (1), tepals<br />
(4), stamens (7).<br />
--9
<strong>Pourouma</strong> 131<br />
'~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~r<br />
'~i '<br />
"<br />
,?il:~~~~~~~~~~<br />
, ^ > 3<br />
\<br />
POUROUMA vebl iainas.
132 Flora Neotropica<br />
sarinho rd., 14 Sep 1961 (9), Rodrigues et al. 3267 (U), 4(1): 129, t. 39. 1853. Type. Colombia. Ama-<br />
9 Aug 1962 (2), Rodrigues et al. 4585 (U). BAHIA: Road zonas: Rio<br />
from Rod. Sao Jose-Buerarena to Una, 7 Jul 1980 Caqueta, Puerto Miraia, J<strong>an</strong> 1820<br />
(juv),<br />
Berg et al. 1143 (U); Una, 29 Aug 1978 (2), Mori et al. (Q), Martius s.n. (holotype, M; isotype, U).<br />
11026 (MG, RB, U); between Vit6ria <strong><strong>an</strong>d</strong> Salvador,<br />
<strong>with</strong>out date (2), Sello s.n. (B). ESPIRITO SANTO: Rio Tree, up to 15 m tall. Leafy twigs 3-15 mm<br />
Doce, Velho-P<strong>an</strong>cas rd., 20 Sep 1930 (6), M. Kuhlm<strong>an</strong>n thick, yellowish- to whitish-puberulous to<br />
383 (RB); Linhares, 11 Mar 1972 (st), Sucre 8684 (U). -(sub)sericeous, or yellow-hirsute, <strong><strong>an</strong>d</strong> <strong>with</strong> sparse<br />
MATO GROSSO: Aripu<strong>an</strong>a, 28 Mar 1977 (st), Gomes et to dense, brown, pluricellular hairs. Lamina enal.<br />
1021 (INPA), 12 Apr 1977 (st), Gomes et al. 1196<br />
(INPA), <strong>with</strong>out date (9), Ryl<strong><strong>an</strong>d</strong>s 15 (INPA); Mun.<br />
tire, ovate to subovate or elliptic to oblong, 5-<br />
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,<br />
24 Sep 1985 (9), Thomas et al. 4018 (BG); Mun. parted, 13-30(-42) x 13-30(-40) cm, coriaceous<br />
Sinop-Colider, BR-080, ca. 91 km E of junction <strong>with</strong> to chartaceous, apex acuminate (to obtuse), base<br />
BR-163, Serra Formosa, 3 Oct 1985 (9), Thomas et al. (obtuse to) truncate to deeply cordate; upper sur-<br />
4179 (BG); Mun. Vila Bela S<strong>an</strong>tissima Trinidade, 4<br />
km<br />
face<br />
S of border ofRond6nia, 3<br />
scabrous to<br />
Nov 1985 (9), Thomas<br />
scabridulous or sometimes<br />
et al. 4810 (BG). PARA: Belem, 2 Jul 1914 (8), Ducke smooth, hairy on the main veins, puberulous on<br />
(MG) 15350 (G, MG, P, US); Belem, 18 Jul 1899 (2), the smaller veins, lower surface smooth, some-<br />
Huber (MG) 1640 (G, MG, P, S, U); Gurupa (2), Ducke times scabridulous, yellowish-appressed-puber-<br />
(MG) 15935 (MG); Rio Xingfi, Vit6ria-Ponte Nova ulous on the main<br />
rd., 7 Aug 1918 (9), Ducke (MG) 17167 (MG); B6a<br />
veins, arachnoid hairs in the<br />
Vista, 15 Oct 1897 (2), Guedes (MG) 1235 areoles <strong><strong>an</strong>d</strong> on the<br />
(G, MG,<br />
smaller<br />
P,<br />
veins, or confined to<br />
S, U); Bel6m-Brasilia rd., km 93, 19 Aug 1959 (6), M. the areoles; lateral veins (6-)12-25(-40) pairs,<br />
Kuhlm<strong>an</strong>n et al. 52 (GUA, MG, R, S, US); mouth of basal pair br<strong>an</strong>ched, tertiary venation almost<br />
Rio Toc<strong>an</strong>tins, Aug 1819 (8), Martius s.n. (M, lectotype pl<strong>an</strong>e beneath; petiole<br />
collection of P.<br />
(2-)4-30(-46) cm<br />
velutina); Jacuarary (=Jaguarari), 21<br />
long,<br />
Aug 1819 (8), Martius s.n. (M); Bel6m, 12 Aug 1945 appressed-puberulous to -strigose, sometimes<br />
(9), Pires et al. 161 (RB); Pirelli, Jul 1958 (a), Pires yellowish-hirtellous to -(sub)tomentose or yel-<br />
7015 (U); Belem, Sep-Oct 1961 (9), Pires 51882 (NY, low-hirsute; stipules 2-13.5(-25) cm long, out-<br />
U, US); Ilha de Breu, 5 Oct 1965 (9), Pr<strong>an</strong>ce et al. 1548 side yellowish- to whitish-appressed-puberulous<br />
(F, GH, NY, P, S, U, US); Cuiaba-S<strong>an</strong>tarem rd., km to<br />
163, 10 Nov 1977 (2), Pr<strong>an</strong>ce et al. 25167<br />
-(sub)sericeous or sometimes yellowish-hir-<br />
(F, INPA,<br />
MO, RB, U), km 941, 13 Nov 1977 (9), Pr<strong>an</strong>ce et al. tellous to -(sub)tomentose or yellow-hirsute, in-<br />
25363 (MO, RB, S, U, US); S<strong>an</strong>tar6m-Palhao rd., km side ? densely hairy, sometimes glabrous, ca-<br />
35, 28 Aug 1969 (9), M. Silva et al. 2423 (F, GH, MG, ducous. Staminate inflorescences up to 17 cm<br />
NY, S, U, US).<br />
long <strong><strong>an</strong>d</strong> 14 cm wide; peduncle 2.5-7 cm long,<br />
yellow-puberulous <strong><strong>an</strong>d</strong> <strong>with</strong> dense (most dense<br />
Vernacular names. Venezuela. Amazonas: cu- on the br<strong>an</strong>ches), dark-brown or red-brown,<br />
cura. Surinam. Boroma or boroma ibeberob<strong>an</strong>a<br />
pluricellular hairs;flowers usually sessile, ? clus-<br />
(Arawak), bospapaja, yarayara (Carib). French<br />
tered, but not in (sub)globose heads; tepals 1-2<br />
Gui<strong>an</strong>a. Bois c<strong>an</strong>on, papaye apici (Paramaka). mm long, free or basally connate, (sparsely) pu-<br />
Brazil. Amazonas <strong><strong>an</strong>d</strong> Mato Grosso: imbafibaraberulous;<br />
filaments shorter th<strong>an</strong> the tepals. Pisna;<br />
Para: mapatir<strong>an</strong>a.<br />
tillate inflorescences in fruit up to 26 cm long <strong><strong>an</strong>d</strong><br />
This species c<strong>an</strong> sometimes be very difficult 14 cm wide; peduncle 1.5-14 cm long peduncle<br />
to distinguish from forms ofP. gui<strong>an</strong>ensis subsp. <strong><strong>an</strong>d</strong> br<strong>an</strong>ches <strong>with</strong> indument similar to that of<br />
gui<strong>an</strong>ensis <strong>with</strong> entire leaves <strong><strong>an</strong>d</strong> sparse pluri- the staminate inflorescences; flowers 6-40; pedcellular<br />
hairs on the leafy twigs.<br />
icel 0.2-0.5 cm long, in fruit 0.4-1 cm long; peri-<br />
The lectotype is chosen from the two (syntype) <strong>an</strong>th 2-4 mm long, scabrous; stigma peltate, 1-2<br />
collections of Martius cited above.<br />
mm in diam. Fruiting peri<strong>an</strong>th purple to black,<br />
ovoid to ellipsoid, ca. 1.5 cm long, scabrous (or<br />
3. <strong>Pourouma</strong> bicolor Martius, Syst. mat. med. smooth), sparsely hairy.<br />
bras. 34. 1843; Miquel in Martius, Fl. bras. In P. bicolor five subspecies c<strong>an</strong> be recognized.<br />
Key to the Subspecies of <strong>Pourouma</strong> bicolor<br />
1. Stipules glabrous inside. ........................................................ . subsp. tessm <strong>an</strong>nii.<br />
1. Stipules hairy inside.<br />
2. Arachnoid hairs on the lower leaf surface confined to the areoles or almost so.
<strong>Pourouma</strong> 133<br />
3. Smaller veins (reticulum) of the lamina beneath pl<strong>an</strong>e or almost so, the veinlets rather thick <strong><strong>an</strong>d</strong><br />
the spots of arachnoid indument distinctly separated; lamina mostly entire or 3-lobed; Upper<br />
Amazon Basin to northern Colombia. ...................................... a. subsp. bicolor.<br />
3. Smaller veins (reticulum) of the lamina beneath more or less prominent, the spots of arachnoid<br />
indument mostly not distinctly separated; lamina mostly 5-7-parted; Central America to the<br />
Pacific coastal region of Colombia <strong><strong>an</strong>d</strong> Ecuador <strong><strong>an</strong>d</strong> to northwestern Venezuela (also in Amazoni<strong>an</strong><br />
Peru?). .................................... ....................... d. subsp. scobina.<br />
2. Arachnoid hairs on the lower leaf surface also covering the smaller veins, usually covering the area<br />
between the parallel tertiary veins.<br />
4. Lamina usually 5-7-parted; pistillate inflorescences often <strong>with</strong> up to 50 flowers.<br />
5. Stipules subpersistent; segments ofthe lamina relatively broad; leafy twigs often yellow-hirsute;<br />
P<strong>an</strong>ama <strong><strong>an</strong>d</strong> western Colombia. ....................................... e. subsp. choco<strong>an</strong>a.<br />
5. Stipules caducous; segments of the lamina relatively narrow; leafy twigs never hirsute; Gui<strong>an</strong>as<br />
<strong><strong>an</strong>d</strong> the Lower Amazon Basin (to southern Venezuela?). ................... c. subsp. digitata.<br />
4. Lamina usually entire or 3-lobed; pistillate inflorescences mostly <strong>with</strong> up to 25(-35) flowers;<br />
Gui<strong>an</strong>a region, Amazon Basin to northern Colombia. ........................ a. subsp. bicolor.<br />
3a. <strong>Pourouma</strong> bicolor Martius subsp. bicolor.<br />
Fig. 63.<br />
<strong>Pourouma</strong> aspera Tr6cul, Ann. Sci. Nat. Bot., Ser. 3,<br />
8: 102. 1847; St<strong><strong>an</strong>d</strong>ley, Publ. Field Mus. Nat. Hist.,<br />
Bot. Ser., 18(1): 391. 1937; St<strong><strong>an</strong>d</strong>ley & Steyermark,<br />
Fieldi<strong>an</strong>a Bot. 24(4): 52.1946. Type. French Gui<strong>an</strong>a.<br />
Without locality, <strong>with</strong>out date (9), Poiteau s.n. (ho-<br />
lotype, P; isotypes, LE, P).<br />
<strong>Pourouma</strong> crassivenosa Mildbraed, Notizbl. Bot. Gart.<br />
Berlin-Dahlem 10:419.1928. Type. Bolivia. La Paz:<br />
Nr. Mapiri, Sep 1907 (2), Buchtien 2050 (holotype,<br />
US; isotypes, B, NY).<br />
<strong>Pourouma</strong> lawr<strong>an</strong>cei St<strong><strong>an</strong>d</strong>ley, Publ. Field. Mus. Nat.<br />
Hist., Bot. Ser., 17: 183. 1937. Type. Colombia. Bo-<br />
yaca: El Humbo, 31 Mar 1933 (2), Lawr<strong>an</strong>ce 727<br />
Specimens examined. COLOMBIA. Without locality,<br />
<strong>with</strong>out date (2), Tri<strong>an</strong>a 861 (E, P). AMAZONAS:<br />
Rio Caqueta, Puerto Mirafia, 1820 (2), Martius s.n. (M,<br />
U type collections of P. bicolor) (M, U).<br />
AMAZONAS-VAUPES: Rio Apaporis, Jinogoj6, 25 Sep<br />
1952 (2), Schultes & Cabrera 17615 (F, type collection<br />
*of P. schultesii). ANTIOQUIA: Confluence of Rios Ite<br />
<strong><strong>an</strong>d</strong> Tamar into Rio Cimatarra, ca. 38 km W of Bar-<br />
r<strong>an</strong>cabermeja, 24 Nov 1967 (6), Bruijn 1497 (F, M,<br />
MO, NY, S, U, US, VEN), 26 Nov 1967 (6), Bruijn<br />
1514 (F, M, MO, NY, S, U, US, VEN), (st), Bruijn<br />
1516 (F, M, MO, NY, S, U, US, VEN), (8), Bruijnl517<br />
(F, M, MO, NY, S, U, US, VEN), 27 Nov 1967 (6),<br />
Bruijn 1526 (F, MO, NY, S, U, US, VEN). BOYACA:<br />
El Humbo, 31 Mar 1933 (6), Lawr<strong>an</strong>ce 727 (A, E, F,<br />
G, MO, S, UC, type collection of P. lawr<strong>an</strong>cei). META:<br />
(holotype, F; isotypes, A, E, G, MO, S, UC). Footpath between Rio Giiejar <strong><strong>an</strong>d</strong> C<strong>an</strong>a Guapayita,<br />
<strong>Pourouma</strong> schultesii Cuatrecasas, Caldasia 7: 303. 1956. Cafno Yerli, 20-28 Dec 1950 (2), Idrobo et al. 786 (F,<br />
Type. Colombia. Amazonas-Vaup6s: Rio Apaporis, MO, NY, US); Villavicencio (2), Karsten s.n. (LE); Sier-<br />
Jinogoje, 25 Sep 1952 (2), Schultes & Cabrera 17615 ra de la Macarena, Cafno Ciervo, 12 J<strong>an</strong> 1950 (2), Phil-<br />
(holotype, F).<br />
ipson et al. 2086 (S, US). PUTUMAYO: S<strong>an</strong>ta Rosa on<br />
<strong>Pourouma</strong> camarat<strong>an</strong>a Cuatrecasas, Acta Bot. Venez. Rio Guamfies, 26 Nov 1966 (2), Pinkley 556 (S).<br />
2(5-8): 202.1967. Type. Venezuela. Bolivar: Auy<strong>an</strong>- SANTANDER: Region Carare-Op6n (Capote), nr. Rio<br />
tepui, above valley of Camarat<strong>an</strong>a, 18 May 1964 (2), Magdalena, 6 Jul 1968 (9), Garcia et al. 9 (US). Vi-<br />
Steyermark 94810 (holotype, US; isotypes, NY, CHADA: 3 km S ofLinea Roja, Parque Nacional Naturel<br />
VEN).<br />
"El Tuparro," 14 Mar 1985 (2), Zarucchi et al. 3720<br />
(BG).<br />
Leafy twigs appressed-puberulous to strigose. VENEZUELA. AMAZONAS: Reserva Forestal "El<br />
Lamina entire to 3-lobed, sometimes 3-fid, oc- Sipapo," Rio Sipapo, May 1971 (2), Bl<strong>an</strong>co 1149 (NY,<br />
casionally (in subjuvenile material?) 5-parted; US, VEN); Rio Marawinuma, nr. Cerro de La Neblina<br />
Base<br />
upper surface scabrous, sometimes smooth, low- Camp, 6 Feb 1984 (st), Liesner 15681 (BG), 28<br />
Feb 1984 (9), Liesner 16299<br />
er surface<br />
(BG); Depto.<br />
smooth, sometimes<br />
Atures, N<br />
scabridulous, the side of Rio Cat<strong>an</strong>iapo, 45 km SE of Puerto Ayacucho,<br />
arachnoid hairs in the areoles <strong><strong>an</strong>d</strong> on the smaller 13 May 1980 (2), Steyermark et al. 122390 (U); Depto.<br />
veins or confined to the areoles, but then the Atabapo, Rio Cunucunuma, between Cerro Duida <strong><strong>an</strong>d</strong><br />
smaller veins relatively thick <strong><strong>an</strong>d</strong> the spots of<br />
Cerro Huachamacahi, J<strong>an</strong>-Feb 1982 (9), Steyermark<br />
et al. 12585<br />
arachnoid hairs<br />
(BG); Rio<br />
clearly separated; stipules inside<br />
Marawinuma, nr. Cerro de La<br />
Neblina Base Camp, 5 May 1984 (6), Thomas 3370<br />
glabrous, caducous. Pistillate inflorescences (BG). BOLIVAR: Rio Chizca, Urim<strong>an</strong>, 5-8 Sep 1953<br />
mostly <strong>with</strong> 3-25 flowers.<br />
(st), Bernardi 910 (NY, VEN); Rio Hacha, 28 Dec 1955<br />
Distribution (Fig. 68). Throughout the Ama- (2), Bernardi 2657 (G, NY,); slopes ofQuebrada O-pozon<br />
Basin <strong><strong>an</strong>d</strong> the Gui<strong>an</strong>a region, extending to<br />
ru-ma, between S<strong>an</strong>ta Teresita de Kav<strong>an</strong>ayen <strong><strong>an</strong>d</strong> Rio<br />
Pacairao (tributary of Rio<br />
northern Colombia; in<br />
Mouak), 20-21 Nov 1944<br />
non-inundated forest at<br />
(st), Steyermark 60401 (F); lower SW slopes of Chilow<br />
altitudes, in Bolivia up to ca. 900 m. m<strong>an</strong>ta-tepui (Torono-tepui), nr. Rio Tirica, 24 May
134 Flora Neotropica<br />
!ie.<br />
~.~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~:'<br />
' '<br />
?I<br />
r-? *~~~. ..<br />
z<br />
' ~ ~ ~<br />
~'~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~.<br />
.,~~~~~~~~ :<br />
$ Y~~j-'"'~2-'"-...<br />
i ~<br />
~~~~~~~~~~~~~~~~~~~~~~~~~~~~~<br />
.. . . : ,,-<br />
ii!:..-~~<br />
~<br />
-P~~~~~~~~~~~~~~~~~~,~<br />
,..~<br />
:,, ,<br />
POURi~A biol.<br />
?f<br />
'.<br />
. ~~~~~~~~
<strong>Pourouma</strong> 135<br />
1953 (st), Steyermark 75540 (F, NY, VEN); Sierra de Reserva Florestal Ducke, Nov 1972 (st), Rodrigues 9206<br />
Lema, lowl<strong><strong>an</strong>d</strong> between Rio Chic<strong>an</strong><strong>an</strong> <strong><strong>an</strong>d</strong> Rio Ay- (INPA). MATO GROSSO: Aripu<strong>an</strong>a, 21 Feb-2 Aug 1977<br />
aiche, between Puerta Lema <strong><strong>an</strong>d</strong> base of the Sierra, 24 (2 <strong><strong>an</strong>d</strong> st), Gomes et al 676, 786, 843, 966, 979, 1052,<br />
Aug 1961 (9), Steyermark 89457 (NY, VEN); frontier 1072, 1079, 1088, 1123, 1176, 2399 (INPA); Ari-<br />
Venezuela-Brazil, NE ofSerr<strong>an</strong>ia Pia-soi, 5-6 J<strong>an</strong> 1962 pu<strong>an</strong>a, nr. Humboldt Centre, rd. to Rio Juruena, 8 Oct<br />
(st), Steyermark 90644 (VEN); Auy<strong>an</strong>-tepui, above 1973 (6), Pr<strong>an</strong>ce et al. 18250 (INPA, NY, U), 30 Nov<br />
valley of Camarata, alt. 1500-1530 m, 18 May 1964 1978 (2), Ryl<strong><strong>an</strong>d</strong>s 52 (INPA). PARA: Rios Mojfi <strong><strong>an</strong>d</strong><br />
(9), Steyermark 94180 (NY, US, VEN, type collection Acara, S of Belem, ca. 7 km N of Mojfi, 3 Jun 1969<br />
of P. camarat<strong>an</strong>a); Rio Caura, nr. Salto Para, 15-17 (6), Austin et al. 4140 (MO); Faro, 6 J<strong>an</strong> 1920 (2), Ducke<br />
J<strong>an</strong> 1977 (9), Steyermark et al. 113096 (U).<br />
(RB) 13057 (RB); between Rio Pacaja <strong><strong>an</strong>d</strong> Rio Muira-<br />
SURINAM. Forest Reserve Z<strong><strong>an</strong>d</strong>erij 1, 24 Oct 1917 pir<strong>an</strong>ga, SW of Ilha de Breu, 20 Sep 1965 (2), Pr<strong>an</strong>ce<br />
(9), BW3376 (MO, U); Watramiri, 8 Oct 1918 (9), BW et al. 1397 (NY, US); BR-22, km 64, 25 Aug 1964 (6),<br />
4033 (NY, U), 13 Dec 1920 (9), BW 5036 (U); Sectie Pr<strong>an</strong>ce et al. 58862 (F, GH, NY, S, U, US); Belem<br />
0, Feb 1943 (9), Stahel (Woodherb. Sur.) 166 (A, F, (IAN) 17 Oct 1963 (2), N. T. Silva 57832 (NY, U, US).<br />
NY, U, UC).<br />
RONDONIA: Forte Principe da Beira, Estrada do Igarape<br />
FRENCH GUIANA. S of Saul, 12 Feb (9), Leeu- da Viiva, 5 J<strong>an</strong> 1962 (2), Rodrigues et a. 4237 (U).<br />
wenberg 11784 (CAY); <strong>with</strong>out locality, (9), Poiteau RORAIMA: Indi<strong>an</strong> trail from Surucucu W to Uaica bes.n.<br />
(LE, P, type collection of P. aspera).<br />
tween Botamatedi <strong><strong>an</strong>d</strong> Maitf, 10 Feb 1971 (2), Pr<strong>an</strong>ce<br />
ECUADOR. NAPO: 6 km SE of Los Sachas, 14 Apr et al. 13582A (INPA, F, GH, M, NY, P, S, U).<br />
1985 (9), Baker 6017 (BG); 5 km N of Coca, on Coca- BOLIVIA. LA PAZ: Nr. Mapiri, Sep 1907 (2), Buch-<br />
Lago Agrio rd., 16 Mar 1980 (st), Br<strong><strong>an</strong>d</strong>byge et al. tien 2050 (B, NY, US, type collection of P. crassiveno-<br />
30194 (AAU); El Chuncho, Estaci6n INIAP, 2 Oct sa), Sep 1907 (6), Buchtien 2122 (NY, US), 12-30 Sep<br />
1987 (st), Palacios 2059 (BG).<br />
1939 (9), Krukoff10867 (A, F, G, MO, NY, S, U, UC,<br />
PERU. AMAZONAS: Rio S<strong>an</strong>tiago, nr. Caterpiza, 14 US), Sep 1939 (8), Krukoff 10874 (F, NY), 8 Oct-15<br />
Nov 1979 (9), Huashikat 1241 (U), 28 Nov 1979 (2), Nov 1939 (2), Krukoff 11254 (F, NY).<br />
Junqui 149 (U). LORETO: Rio Mom6n, 4 Sep 1972 (9),<br />
Croat 19997 (MO); Maynas, Yaguasyacu, tributary of Vernacular names. Colombia. Antioquia-Bo-<br />
Rio Ampiyacu, below Borro Indi<strong>an</strong> village of Brilla livar: cirpe (or sirpe) hembra,<br />
Nueva, 7<br />
cirpe (or<br />
Nov 1977 (9), Gentry et al. 20356<br />
sirpe)<br />
(MO); Rio<br />
Ucayali, Jenaro Herrera, 7 Dec 1977 (9), Gentry et al. macho; Boyaca: cormi; Meta: caimar6n de mico.<br />
21193 (MO, U); Rio Javari, between Rio Curaqa R. Putumayo: otsepacho tsaha (Kof<strong>an</strong>). Venezuela.<br />
<strong><strong>an</strong>d</strong> Tambaqui, 28 Oct 1976 (9), Pr<strong>an</strong>ce et al. 24188 Amazonas: cucura or cucure, sadha'fhi; Bolivar:<br />
(INPA, MG, U).<br />
cay-bari-cay, cay-i-wari-cay-yek, sarasara. Suri-<br />
BRAZIL. Without locality (9), Burchel 9792 (GH, nam. Boroma<br />
LE, P, under the same number P. mollis (Arawak), pourouma or<br />
subsp.<br />
puruma<br />
mollis).<br />
ACRE: Upper Rio Moa, Fazenda Arizona, 10-16 Oct (Carib). Peru. Amazonas: paui shuina, tsakap sui-<br />
1985 (st), Campbell et al. 6650 (BG); Rio Jurua, nr. ya (Huambisa). Brazil. Acre: imbafbar<strong>an</strong>a.<br />
Cruzeiro do Sul, 22 Oct 1966 (9), Pr<strong>an</strong>ce et al. 2754 Amazonas: imbafiba; Mato Grosso: imbafibar-<br />
(INPA, MG, U). AMAPA: Rio Oiapoque, 1-3 km N of<br />
<strong>an</strong>a,<br />
Cachoeira Tres Saltos, 12 Sep 1960 (6), Irwin<br />
tamaoquare; Para:<br />
et al.<br />
garguaba, mapatir<strong>an</strong>a.<br />
48192 (GH, M, NY, P, U, US). AMAZONAS: 27<br />
Bolivia. La Paz: uva menueda.<br />
Sep<br />
1973 (8), Berg et al. P. 18152 (INPA, MO, NY, P, U); In <strong><strong>an</strong>d</strong> towards the eastern part of the species<br />
M<strong>an</strong>aus-Caracarai rd., km 148, 26 Sep 1973 (6), Bisby r<strong>an</strong>ge (from Colombia to Bolivia) the arachnoid<br />
et al. P.18118 (F, INPA, MO, NY, S, U, US); Rio hairs on the lower leaf surface are confined to the<br />
Negro, Uaupes, 22 J<strong>an</strong> 1960 (9), Cavalc<strong>an</strong>te 777 (MG); areoles in<br />
Serra de Neblina, Rio Cauaburi, Camp Tuc<strong>an</strong>o, 5<br />
combination<br />
Dec<br />
<strong>with</strong> <strong>an</strong> increase of the<br />
1965 (9), Maguire et al. 60333 (F, MG, NY, P, U, US);<br />
thickness of the smaller veins, causing distinct<br />
Benjamin Const<strong>an</strong>t, 16 Oct 1956 (9), Meyer Drees 9 separation of the "islets" of arachnoid indument.<br />
(INPA, U); M<strong>an</strong>aus, Reserva Florestal Ducke, 6 Dec The tr<strong>an</strong>sition of the normal form of subsp. bi-<br />
1976 (2), Nascimento et al. (INPA) 66338) (INPA); Rio<br />
color, <strong>with</strong> the arachnoid indument<br />
Negro, Uaup6s, 1 May 1947 (9), Pires 502 (NY); Rio<br />
extending<br />
Negro, Sao Gabriel, 1 May 1947 (6), Pires 587<br />
over the<br />
(NY); relatively thin smaller veins to the form<br />
Tapuruquara, rd. to airport, 17 Oct 1917 (9), Pr<strong>an</strong>ce de scribed above, is rather gradual. Both forms<br />
et al. 15366 (INPA, M, MO, NY, S, U, US); M<strong>an</strong>aus, may occur in the same region. In the eastern part<br />
FIG. 63. <strong>Pourouma</strong> bicolor subsp. bicolor. From Martius, Flora Brasiliensis, 4(1). 1853: tab. 39, Leafy twig<br />
<strong>with</strong> fruiting pistillate inflorescence, fruiting peri<strong>an</strong>th (17), fruiting peri<strong>an</strong>th <strong><strong>an</strong>d</strong> fruit (1711), pericarp (43), fruit<br />
(19), pericarp <strong><strong>an</strong>d</strong> seed (21), seed (19).
136 Flora Neotropica<br />
A 3<br />
?3X3l cm<br />
t. '- ?<br />
FG. 64. <strong>Pourouma</strong> bicolor subsp. tessm<strong>an</strong>ni. Leafy twigs <strong>with</strong> pistillate 94).'ur inflorescence (Tunqui<br />
(!<br />
L<br />
..'<br />
i-"<br />
i. _ .......<br />
'<br />
.':..
<strong>Pourouma</strong> 137<br />
of the species r<strong>an</strong>ge the upper leaf surface is often<br />
smooth. See unnamed collection #1 (p. 193).<br />
3b. <strong>Pourouma</strong> bicolor Martius subsp. tessm<strong>an</strong>nii<br />
(Mildbraed) C. C. Berg & v<strong>an</strong> Heusden, Proc.<br />
Kon. Ned. Akad. Wetensch., Ser. C 91(2): 106.<br />
1988. Fig. 64<br />
<strong>Pourouma</strong> tessm<strong>an</strong>nii Mildbraed, Notzibl. Bot. Gart.<br />
Berlin-Dahlem 10: 192. 1927. Type. Peru. Loreto:<br />
Pongo de M<strong>an</strong>seriche, 8 Oct 1924 (6), Tessm<strong>an</strong>n<br />
4236 (holotype, B; isotypes, NY, S, US).<br />
Leafy twigs appressed-puberulous to strigose.<br />
Lamina entire or occasionally 3-lobed to -fid;<br />
upper surface scabrous, lower surface smooth,<br />
appressed hairs on the main veins, arachnoid<br />
hairs in the areoles <strong><strong>an</strong>d</strong> on the smaller veins;<br />
smaller veins (almost) pl<strong>an</strong>e; stipules outside <strong>with</strong><br />
appressed hairs, inside glabrous, caducous. Pistillate<br />
inflorescences mostly <strong>with</strong> 3-16 flowers.<br />
Distribution (Fig. 68). Peru (Amazonas <strong><strong>an</strong>d</strong><br />
Loreto); in non-inundated forest.<br />
Specimens examined. PERU. AMAZONAS: Rio S<strong>an</strong>tiago,<br />
65 km N of Teniente Pinglo, Quebrada Caterpiza,<br />
13 Oct 1979 (2), Huashikat 912 (U), 30 J<strong>an</strong> 1980<br />
(9), Huashikat 1872 (U); Rio S<strong>an</strong>tiago, nr. La Paz, 12<br />
Nov 1979 (2), Tunqui 7 (BG); Rio S<strong>an</strong>tiago, Quebrada<br />
Caterpiza, 17 Nov 1979 (6 <strong><strong>an</strong>d</strong> 2), Tunqui 94 (U).<br />
HuANuco: Prov. Leoncio Prado, 80 km NE of Tingo<br />
Maria, nr. La Divisoria, 18 J<strong>an</strong> 1976 (2), Gentry et al.<br />
1604 (DUKE, MO, NY). LORETO: 22 km S of km 86<br />
on Pucallpa-Tingo Maria hwy., 11 Feb 1981 (2), Gentry<br />
et al. 31197 (U); Pongo de M<strong>an</strong>seriche, nr. mouth of<br />
Rio S<strong>an</strong>tiago, 8 Oct 1924 (d), Tessm<strong>an</strong>n 4236 (B, NY,<br />
S, US, type collection of P. tessm<strong>an</strong>nii).<br />
Vernacular names. Peru. Amazonas: shuiya<br />
(Huambisa), t<strong>an</strong>ta shuiya, tentar shuina (Huambisa).<br />
This subspecies is similar to subsp. bicolor in<br />
most features but the stipules are glabrous inside.<br />
3c. <strong>Pourouma</strong> bicolor Martius subsp. digitata<br />
(Trecul) C. C. Berg & v<strong>an</strong> Heusden, Proc. Kon.<br />
Ned. Akad. Wetensch., Ser. C 91(2): 106.<br />
1988. Fig. 65.<br />
<strong>Pourouma</strong> digitata Tr6cul, Ann. Sci. Nat. Bot., S6r. 3,<br />
8: 106. 1847; Miquel in Martius, Fl. bras. 4(1): 124.<br />
1853. Type. French Gui<strong>an</strong>a. Without locality, <strong>with</strong>-<br />
out date (9), Poiteau s.n. (lectotype P, chosen here;<br />
isolectotype B).<br />
Leafy twigs appressed-puberulous to strigose.<br />
Lamina 5-7(-9)-parted, midsegment l<strong>an</strong>ceolate<br />
to obovate, base truncate to shallowly cordate or<br />
sometimes deeply cordate; upper surface scabrous<br />
to scabridulous, sometimes smooth, lower<br />
surface <strong>with</strong> appressed hairs on the main veins,<br />
arachnoid hairs in the areoles <strong><strong>an</strong>d</strong> on the smaller<br />
veins; smaller veins (almost) pl<strong>an</strong>e: stipules outside<br />
<strong>with</strong> appressed hairs, inside hairy, caducous.<br />
Pistillate inflorescences mostly <strong>with</strong> 10-50 flowers.<br />
Distribution (Fig. 68). Guy<strong>an</strong>a, Surinam,<br />
French Gui<strong>an</strong>a, <strong><strong>an</strong>d</strong> Brazil (Amapa <strong><strong>an</strong>d</strong> Parf),<br />
<strong><strong>an</strong>d</strong> probably in southeastern Venezuela; in noninundated<br />
forest at low altitudes.<br />
Specimens exmained. VENEZUELA. AMAZONAS:<br />
Nr. Cerro de La Neblina Base Camp, 19 Apr 1984 (st),<br />
Gentry et al 46752 (BG).<br />
GUYANA. NW slopes of K<strong>an</strong>uku Mts., drainage of<br />
Moku-Moku Creek, Takutu tributary, 31 Mar-16 Apr<br />
1938 (2), A. C. Smith 3564 (B, F, LE, MO, NY, P, S,<br />
U, US).<br />
SURINAM. Watramiri, 9 Jun 1916 (st), BW 2030<br />
(U); Kaboeri, 30 Oct 1920, B W 4866 (U); Brownsberg,<br />
4 Oct 1923 (S), BW 6292 (U); Distr. Brokopondo, 2<br />
km S of G<strong>an</strong>see, 27 Apr 1964 (st), Donselaar 1262 (U);<br />
Nassau Mts., at km 0.2 on terrace of Marowijne R.,<br />
16 Feb 1949 (st), L<strong>an</strong>jouw et al. 2213 (F, NY, RB, U);<br />
Distr. Saramacca, nr. Groningen, 19 Feb 1954 (st),<br />
Lindem<strong>an</strong> 5468 (U); Rik<strong>an</strong>au, nr. Moengo, 6 Jun 1954<br />
(st), Lindem<strong>an</strong> 6123 (U); S of Moengotapoe, 14 Jun<br />
1954 (st), Lindem<strong>an</strong> 6166 (U); Jodensav<strong>an</strong>ne-Map<strong>an</strong>e<br />
Creek area, 10 Dec 1954 (st), Lindem<strong>an</strong> 6765 (U);<br />
Distr. Nickerie, area of Kabalebo Dam project, 4 km<br />
NW of rd., km 39.5, 4 Nov 1981 (st), Lindem<strong>an</strong> & de<br />
Roon 750 (U); Lely Mts., E rd. on plateau 1, 29 Sep<br />
1976 (juv), Lindem<strong>an</strong> & Stoffers et al. 539 (U), 22 J<strong>an</strong><br />
1970 (2), Oldenburger et al. 1120 (U); Sipaliwini area,<br />
Brazili<strong>an</strong> frontier, 2.5 km S of Sipaliwini R., Feb 1970<br />
(st), Oldeburger et al. 1406 (U); Winn<strong>an</strong>a Creek, Upper<br />
Kabouir Creek, Cor<strong>an</strong>tyne R., 20 May 1960 (st), Schulz<br />
(LBB) 8316 (U); Section 0, ca. 5?N, 55?W, Nov 1942<br />
(a), Stahel 123A (U), Dec 1942 (S), Stahel (Woodherb.<br />
Sur.) 166B (A, NY, U, UC).<br />
FRENCH GUIANA: Cayenne, 1 Dec 1955 (2), BAF-<br />
OG (=Bena) 1050 (CAY, F, U); upper Oyapock R.,<br />
Zidock ville, 6 Sep 1977 (st), Gren<strong><strong>an</strong>d</strong> 1442 (CAY, U);<br />
Saul, 8 Dec. 1982 (2), Mori et al. 15350 (BG); deserted<br />
village of Eau Claire, about 7 km N of Belizon, 12 Feb<br />
1966 (9), Oldem<strong>an</strong> 2030 (CAY); Yaroupi R., 8 km<br />
downstream of Saut Tainoua, 18 Apr 1970 (st), Oldem<strong>an</strong><br />
3115 (CAY, P, U); <strong>with</strong>out locality, <strong>with</strong>out date<br />
(9), Poiteau s.n. (B, P, type collection of P. digitata),<br />
<strong>with</strong>out date (8), Poiteau s.n. (P); Oyapoque R., Camopi,<br />
8 Mar 1976 (9), Sastre 4350 (P, U); Mt. St.<br />
Marcel, 28 Mar 1976 (9), Sastre 4579 (P).<br />
BRAZIL. AMAPA: Serra do Navio, 1961 (2), Aubreville<br />
144 (P, U, US); rd. to Clevel<strong><strong>an</strong>d</strong>ia, 14 Oct 1950<br />
(9), Fr6es 26631 (IAN), 13 Oct 1960 (a), Irwin 48682<br />
(GH, IAN, NY, U, US); Rio Oiapoque, Mt. Tipac, 16<br />
Oct 1960 (8), Irwin 48755 (IAN, NY); Rio Oiapoque,<br />
Mt. Tipac, 16 Oct 1961 (6), Pires et al. 51624 (B, F,
138 Flora Neotropica<br />
? ..J<br />
'" - i; . . . . .<br />
.ru<br />
_l<br />
*<br />
,| x. . . . | |.. | . .<br />
FIG. 65. <strong>Pourouma</strong> bicolor subsp. digitata. Leaf <strong><strong>an</strong>d</strong> pistillate inflorescences BAFOG 1050).<br />
FIG. 65. <strong>Pourouma</strong> bicolor subsp. digitata. Leaf <strong><strong>an</strong>d</strong> pistillate inflorescences (BAFOG 1050).<br />
NY, R, UC, US). PARA: Rio Jari, Monte Dourado, 4<br />
Feb 1969 (9), Cavalc<strong>an</strong>te 2297 (MG); 25-35 <strong><strong>an</strong>d</strong> 65<br />
km SW of Tucurui, 4 <strong><strong>an</strong>d</strong> 18-20 Nov 1981 (9), Daly<br />
et al. 1208 <strong><strong>an</strong>d</strong> 1474 (BG); Rio Jari, Monte Dourado,<br />
15 Nov 1968 (9), N. T. Silva 1411 (NY, U, US), 15<br />
Nov 1968 (8), N. T. Silva 1412 (NY, U).<br />
Vernacular names. Surinam. boroma (Ara-<br />
wak), bospapaja, pourouma or puruma (Carib),<br />
yarayara (Carib). French Gui<strong>an</strong>a. a'ilea (Waya-<br />
pi), male bois c<strong>an</strong>on or bois c<strong>an</strong>on, kulumatelelel<br />
(Wayapi). Brazil. Para: mapatir<strong>an</strong>a.
<strong>Pourouma</strong> 139<br />
This subspecies resembles P. cecropiifolia very<br />
much. It is therefore somewhat doubtful whether<br />
Gentry et al. 46752 (from Venezuela, Amazonas)<br />
belongs to subsp. digitata or represents a juvenile<br />
form of P. cecropiifolia. Subspecies digitata dif-<br />
30 Apr 1939 (st), St<strong><strong>an</strong>d</strong>ley 72667 (F); between Virginia<br />
<strong><strong>an</strong>d</strong> Lago Izabal, 5 Apr 1940 (st), Steyermark 38916<br />
(F).<br />
BELIZE. Belmop<strong>an</strong>, 21 Jun 1973 (6), Croat 24831<br />
(MO); St<strong>an</strong>n Creek district, Middlesex, 5 May 1939<br />
(9), Gentle 2787 (A, F, NA, NY, US); St<strong>an</strong>n Creek<br />
valley, 18 Mar 1940 (9), Gentle 3265 (A, NY); Monkey<br />
R., 18 Oct 1942 (6), Gentle 4211 (DUKE); El Cayo<br />
district, Hummingbird Hwy., 16 Apr 1955 (9), Gentle<br />
8675 (F, S, US); Maya Mts., 18 Jun 1930 (9), Schipp<br />
fers from P. cecropiifolia in the smaller number<br />
of incisions of the lamina <strong><strong>an</strong>d</strong> usually a scabrous<br />
to scabridulous upper surface of the lamina.<br />
Nests of small <strong>an</strong>ts are sometimes found in 127 (A, F, G, GH, MO, NY, S).<br />
between the bases of the main veins.<br />
HONDURAS. Rio Esper<strong>an</strong>za, 28 Feb 1903 (6), Wil-<br />
From the two syntype collections, Leprieurs.n. son 606 (NY, US). ATLANTIDA: S<strong>an</strong> Alejo, 22-27 Apr<br />
<strong><strong>an</strong>d</strong> Poiteau s. n. (staminate specimens), the latter 1947 (st), St<strong><strong>an</strong>d</strong>ley 7878 (F); Tela, 4 Dec 1927-20 Mar<br />
1928<br />
is chosen as lectoptype collection.<br />
(st), St<strong><strong>an</strong>d</strong>ley 52900 (A, F, US), (9), St<strong><strong>an</strong>d</strong>ley<br />
53951 (F, US). CORTES: Agua Azul, 9 Feb 1952 (9),<br />
Allen 6401 (F, GH, US). GRACIAS A Dios: Rio Plat<strong>an</strong>o,<br />
17-23<br />
3d. <strong>Pourouma</strong> bicolor Martius subsp. scobina May 1973 (9), Clewell et al. 4142 (MO).<br />
NICARAGUA. ZELAYA: Puerto Cabezas, 2 Dec 1927<br />
(Benoist) C. C. Berg & v<strong>an</strong> Heusden, Proc. (9), Englesing 50 (F), (st), Englesing 52A (F); between<br />
Kon. Ned. Akad. Wetensch., Ser. C 91(2): 106. Bluefields <strong><strong>an</strong>d</strong> Ginney Point, 24 Mar 1949 (st), Molina<br />
1988. Fig. 66. R. 1963 (GH); El Recreo-El Pijibaye rd., 11 May 1949<br />
(st), St<strong><strong>an</strong>d</strong>ley 19904 (F); rd. to Colonia M<strong>an</strong><strong>an</strong>tiales, S<br />
<strong>Pourouma</strong> scobina Benoist, Bull. Mus. Hist. Nat. (Paris) of ridge of Serr<strong>an</strong>ias de Yol<strong>an</strong>ia, 29-31 Oct 1977 (st),<br />
28: 320. 1922; Woodson & Schery, Ann. Missouri Stevens 4839 (BG); ca. 14.3 km N of El Enpalme, along<br />
Bot. Gard. 47: 167. 1960. Type: Costa Rica. S<strong>an</strong> new rd. to Mina Nueva America, 27 Apr 1978 (9),<br />
Jose: Tucurrique, Dec 1898 (i), Tonduz 12930 (ho- Stevens 8323 (BG).<br />
lotype, P; isotypes, F, GH, M, NY, S, US).<br />
COSTA RICA. ALAJUELA: Caribl<strong>an</strong>co, Quebrada<br />
<strong>Pourouma</strong> crassivenia Cuatrecasas, Revista Acad. Co- Array<strong>an</strong>es, 1 Jul 1973 (6), Lent 3547 (DUKE, F, MO,<br />
lomb. Ci. Exact. 9(36/37): 340. 1956. Type. Colom- U). CARTAGO: Turrialba, 20 May 1951 (9), Le6n 3501<br />
bia. Valle: Rio Digua, Piedra de Moler, 24 Aug 1943 (NA). HEREDIA: Puerto Viejo, Rio Sarapiqui, 19 J<strong>an</strong><br />
(9), Cuatrecasas 15071 (holotype, F; isotypes, GH, 1974 (st), Hartshorn 1348 (F), 1 Oct 1974 (9), Hart-<br />
NY).<br />
shorn 1539 (F, U); Rio Sarapiqui, Tirimbina, 16 Jul<br />
1971 (6), Lent 2004 (F, G, MO, NY, U, US); Rio<br />
Leafy twigs appressed-puberulous to strigose Sarapiqui, La Virgen, 23 Jul 1979 (9), Stevens 13341<br />
or hirtellous. Lamina usually 5-7(-9)-parted, (U). LIMON: Madre de Dios, 26 Feb 1949 (9), Holdridge<br />
sometimes 3-lobed to -parted, midsegment<br />
2514 (US); Rio Hondo, Aug 1901 (9), Pittier 16163<br />
mostly l<strong>an</strong>ceolate to oblong, sometimes (G, US). PUNTARENAS: Valley of Rio Diquis, between<br />
elliptic Union <strong><strong>an</strong>d</strong> S<strong>an</strong> Pedro, 29 Mar 1898 (9), Pittier 12105<br />
to obovate, base + deeply cordate; upper surface (BR, P, US). SAN JOSE: S<strong>an</strong> Pablo, 2 Mar 1966 (9),<br />
scabrous (to scabridulous), lower surface smooth, Jimenez 3816 (F); S<strong>an</strong> Isidro, El General, 2 Mar 1966<br />
hairs on the main vein appressed to patent, ar- (9), Molina R. 18227 (F, NY, US); El General, J<strong>an</strong><br />
achnoid hairs (almost) confined to the areoles<br />
1939 (6), Skutch 4083 (A, MO, NY, S, US), Feb 1939<br />
but the spots of arachnoid<br />
(9), Skutch 4225<br />
hairs mostly not clear-<br />
(A, MO, NY, S); Rio de las Vueltas,<br />
Tucurrique, Dec 1898 (6), Tonduz 12930 (F, GH, M,<br />
ly separated by the smaller veins; smaller veins NY, P, S, US, type collection of P. scobina).<br />
more or less prominent; stipules outside <strong>with</strong> ap- PANAMA. COCLE: Valley of Rio Anton, 1000 m,<br />
pressed or patent hairs, inside hairy, caducous. 27 Sep 1946 (9), Allen 3742 (BR, E, F, G, MO, P).<br />
Pistillate<br />
COLON: S<strong>an</strong>ta<br />
inflorescences <strong>with</strong> (10-)20-50 (or more<br />
Rita, Feb 1968 (9), Gomez-Pompa 3385<br />
(MO). DARIEN: Cerro Pirre, 14 Dec 1962 (9), Duke<br />
?) flowers.<br />
6588 (MO); Pidiaque, 28 Mar 1966 (9), Duke 8035<br />
Distribution (Fig. 68). Central America (from (MO); between Cerro Pierre <strong><strong>an</strong>d</strong> Piji Vasal, 15 Nov<br />
Guatemala to P<strong>an</strong>ama) <strong><strong>an</strong>d</strong> the Pacific coastal 1977 (st), Folsom 6377 (MO); Cocalito, 8-9 Apr 1978<br />
region of Colombia <strong><strong>an</strong>d</strong> Ecuador, <strong><strong>an</strong>d</strong> to north- (st), Garwood 759 (F); Cerro Pirre, 1000-1400 m, 29<br />
Dec 1972<br />
western Venezuela, possibly also in<br />
(9),<br />
Peru<br />
Gentryet al. 6997 (F, MO); Rio Setig<strong><strong>an</strong>d</strong>i,<br />
(Loreto 19 Apr 1980 (9), Gentry et al. 28609 (U); M<strong>an</strong>ene, 23<br />
<strong><strong>an</strong>d</strong> S<strong>an</strong> Martin); in non-inundated forest at al- Dec 1980 (9), Hartm<strong>an</strong> 12176 (U); between Paya <strong><strong>an</strong>d</strong><br />
titudes up to ca. 1600 m.<br />
Pucro, 12 Jun 1959 (9), Steam et al. 423 (G, GH, LE,<br />
US); Pediaque Hill, nr. Rio Sab<strong>an</strong>a <strong><strong>an</strong>d</strong> Rio Lara, 16<br />
Specimens examined. GUATEMALA. ALTA VERA- Jul 1966 (2), Tyson et al. 4734 (DUKE, GH, MO);<br />
PAZ: Finca Cubilguitz, Mar 1913 (Q), Tuerckheim 4082 C<strong>an</strong>a, 17 Apr-8 Jun 1908 (9), R. S. Williams 983 (NY,<br />
(US). IZABAL: Puerto Mendez, 12 Jun 1970 (2), Con- US). PANAMA: El Ll<strong>an</strong>o-Carti rd., 20 Feb 1973 (9),<br />
treras 10026 (DUKE, F, U, US); El Estor, 7 Mar 1972 Correa et al. 1854 (F, MO); Cerro Azul, 26 Jul 1970<br />
(2), Contreras 11195 (DUKE, P, S, US); Entre Rios, (st), Croat 11591 (F, MO); between Cerro Azul <strong><strong>an</strong>d</strong>
140 Flora Neotropica<br />
' I?<br />
UNITDO STATES NATIONAL li55UII5553<br />
inch ?<br />
. J e . td. .<br />
-N-<br />
FIG. 66. <strong>Pourouma</strong> bicolor subsp. scobina. Leafy twig <strong>with</strong> pistillate inflorescences<br />
(Allen 6401).<br />
Cerro Jefe, 6 Aug 1968 (Q), Dressier 3579 (DUKE, GH,<br />
MO); El Ll<strong>an</strong>o-Carti rd., 17 Apr 1981 (Q), Sytsma 4018<br />
(U). SAN BLAS: C<strong>an</strong>g<strong><strong>an</strong>d</strong>i, 10 Feb 1986 (st), Nevers et<br />
al. 7178 (BG), 27 Mar 1986 (st), Nevers et al. 7542<br />
(BG).<br />
COLOMBIA. CHOC6:<br />
Rio Curiche, camp Curiche,<br />
25 May 1967 (9), Duke 11599 (US). VALLE: Rio Digua,<br />
Piedra de Moler, 24 Aug 1943 (2), Cuatrecasas 15071<br />
(F, GH, NY, type collection of P. crassivenia); Rio<br />
S<strong>an</strong>quinini, La Laguna, 14 Dec 1943 (d), Cuatrecasas
<strong>Pourouma</strong><br />
15520 (F, GH, NY), (9), Cuatrecasas 15534 (F, GH, subspecies <strong><strong>an</strong>d</strong> in that of the other subspecies,<br />
NY).<br />
except for subsp. choco<strong>an</strong>a. Such specimens are<br />
VENEZUELA. BARINAS: Pedreza, Feb 1955 (2),<br />
Bernardi 1986 (NY, VEN); rd. Barinitas-El Cacao, 21 hardly distinguishable from P. gui<strong>an</strong>ensis subsp.<br />
Sep 1982 (9), Marc<strong>an</strong>o-Bert et al. 982-154 (U). MERI- gui<strong>an</strong>ensis <strong>with</strong> regard to the leaf indument (see<br />
DA: Caiio Amarillo, 5 Feb 1955 (2), Bernardi 1919 (F, p. 122). Fortunately these taxa c<strong>an</strong> mostly be<br />
NY, VEN); El Vigia, 14 Jun 1979 (6), Marc<strong>an</strong>o-Bert distinguished on the presence of hairs on the inet<br />
al. 238-979 (U). TACHIRA: La Fria, 22 Dec 1965 (2), ner surface of the<br />
Breteler 4920 (MO, U); 35 km SSE of S<strong>an</strong> stipules <strong><strong>an</strong>d</strong> are, moreover,<br />
Cristobal,<br />
20-21 Mar 1981 (2), Liesner et al. 10944 (BG); S<strong>an</strong> geographically separated. However, two recent<br />
Joaquino de Navay, 6 Nov 1979 (2), Stevermark et al. collections from Peru (S<strong>an</strong> Martin; Gentry et al.<br />
119396 (U). ZULIA: Dtto. Col6n, nr. Casuigua El Cubo, 45160) <strong><strong>an</strong>d</strong> (Loreto, Gentry et al. 42168) may<br />
21 J<strong>an</strong> 1979 (9), Bunting 6836 (BG); Sierra de Perija, also<br />
SW ofMachiques, 31 Aug 1967 (2), Steyermark 99945<br />
belong to subsp. scobina. The indument on<br />
the<br />
(U, VEN).<br />
petiole, the leafy twigs, <strong><strong>an</strong>d</strong> the main veins<br />
ECUADOR. CARCHI: Maldonado, 25 Sep 1979 (6), of the lamina beneath is denser th<strong>an</strong> usual in the<br />
Gentry et al. 26530 (U). EL ORO: Piedras, 20 Jun 1943 collections from Central America <strong><strong>an</strong>d</strong> western<br />
(st), Little 6640 (F, US). ESMERALDAS: S<strong>an</strong>to Domingo Ecuador. The latter collection has, moreover,<br />
de los Colorados-Quininde rd., 14 Apr 1943, Acosta<br />
S. 13605 (F); Quininde, 12-14 Apr 1943 (2), Little 6241 bright-yellow hairs on the stipules <strong><strong>an</strong>d</strong> 9-parted<br />
(F, US), (st), Little 6258 (F, UC, US); S<strong>an</strong> Lorenzo, 20 leaves <strong>with</strong> subpetiolulate segments. The identity<br />
Apr 1943 (2), Little 6301 (F, U, UC, US). IMBABURA: of these two collections is rather uncertain.<br />
Collapi, 4 Jun 1949 (st), Acosta S. 12884 (F). Los Rios:<br />
Rio Palenque Biol. Station, 16 Aug 1976 (st), Dodson<br />
6156 (MO), 2 Oct 1976 (st), Dodson et al. 6312 (MO), 3e. <strong>Pourouma</strong> bicolor Martius subsp. choco<strong>an</strong>a<br />
(6), Dodson et al. 6327 (GB, MO, U), 2 Feb 1974 (9), (St<strong><strong>an</strong>d</strong>ley) C. C. Berg & v<strong>an</strong> Heusden, Proc.<br />
Gentry 9552 (MO, U). PICHINCHA: S<strong>an</strong>to Domingo de Kon. Ned. Akad. Wetensch., Ser. C 91(2): 106.<br />
los Colorados, 30 Aug 1930 (6), Benoist (P, S, U), 17<br />
Oct 1961 (2), Cazalet et<br />
1988.<br />
al. 5037 (FHO, NY, UC, US),<br />
Fig. 67<br />
19 Oct 1961 (6), Cazalet et al. 5072 (FHO, NY, UC, <strong>Pourouma</strong> choco<strong>an</strong>a St<strong><strong>an</strong>d</strong>ley, Publ. Field Mus. Nat.<br />
US).<br />
Hist., Bot. Ser., 22: 73. 1940.<br />
PERU. LORETO: Prov. Maynas, Y<strong>an</strong>amono<br />
Type: Colombia. Cho-<br />
Explora- c6: Junction of Rio Condoto <strong><strong>an</strong>d</strong> Rio S<strong>an</strong><br />
ma Tourist Camp, 26 Jun 1983 (st), Gentry et al.<br />
Ju<strong>an</strong>, 20<br />
42168<br />
(BG). SAN<br />
Apr 1939<br />
MARTIN:<br />
(2), Killip & A. C. Smith 35095<br />
Rioja-Pomacochas rd., below<br />
(holotype,<br />
F;<br />
Venceremos, 1600 m, 8 Feb<br />
isotype, US).<br />
1984 (9), Gentry et al. <strong>Pourouma</strong><br />
45160<br />
johnstonii Woodson, Ann. Missouri Bot.<br />
(BG).<br />
Gard. 47: 166. 1960; Burger, Fieldi<strong>an</strong>a Bot. 40: 201.<br />
1977. Type. P<strong>an</strong>ama. C<strong>an</strong>al Zone, W of Lim6n Bay,<br />
Vernacular names. Guatemala. Guarumo de Gatun Locks, Maru Towers, 27 Mar 1956 (Q), Johnston<br />
1714<br />
montafia. Belize. Trumpet tree. Honduras.<br />
(holotype, MO; isotype, A).<br />
Guarumo de montafia, m<strong>an</strong>o de le6n. Nicaragua.<br />
Guarumo macho, pacica, yahal (Moskito Indi<strong>an</strong>s).<br />
Costa Rica. Chumico, guarumo de montafia,<br />
lija. P<strong>an</strong>ama. Darien: guarumo macho, piraejo.<br />
Colombia. Choc6: uva. Venezuela. Barinas:<br />
tinajero. Merida: orumo de monte. Ecuador. Esmeraldas:<br />
guagay, guarumo de montafia, uva, uva<br />
del monte.<br />
The lamina is mostly 5-7-fid to -parted, but<br />
in P<strong>an</strong>ama 3-lobed laminas occur. The number<br />
of flowers in the pistillate inflorescence is often<br />
50 or more. In Guatemala <strong><strong>an</strong>d</strong> P<strong>an</strong>ama collections<br />
<strong>with</strong> (10-20) flowers in the pistillate inflorescence<br />
have been made. In several collections<br />
in the southern part of the subspecies r<strong>an</strong>ge (Ec-<br />
Leafy twigs patent- to appressed-puberulous<br />
<strong><strong>an</strong>d</strong> at least on the scars of the stipules (sparsely)<br />
yellow-hirsute. Lamina mostly 5-fid to -parted,<br />
sometimes 3-lobed to -fid or entire, occasionally<br />
(in subjuvenile material?) 7-parted, midsegment<br />
broadly elliptic to oblong, base deeply or sometimes<br />
shallowly cordate; upper surface scabrous,<br />
lower surface smooth, appressed hairs on the main<br />
veins, arachnoid hairs in the areoles <strong><strong>an</strong>d</strong> on the<br />
smaller veins, mostly covering the areas between<br />
the parallel tertiary veins; smaller veins (almost)<br />
pl<strong>an</strong>e; stipules outside yellow-hirsute to -subsericeous,<br />
inside hairy, subpersistent. Pistillate inflorescences<br />
mostly <strong>with</strong> up to 50 (or more?) flowers.<br />
uador to P<strong>an</strong>ama) the indument of the lamina<br />
beneath, the petioles, stipules, peduncles <strong><strong>an</strong>d</strong> leafy<br />
twigs is patent (hirtellous to (sub)tomentose) instead<br />
of appressed as in most collections of this<br />
Distribution (Fig. 68). P<strong>an</strong>ama (C<strong>an</strong>al Zone<br />
<strong><strong>an</strong>d</strong> Col6n) to western Colombia (Choc6 <strong><strong>an</strong>d</strong><br />
Valle); in non-inundated forest at altitudes up to<br />
ca. 1000 m.<br />
141
142 Flora Neotropica<br />
movririg-. o<br />
-..<br />
Ievsed for YXora eotr ei''<br />
Pouroum bicolar orths* subsp, chocoasa<br />
?Staodisy) C.C. krg & v<strong>an</strong> Seusden<br />
Dec~L~E~I!t.: C.c. Barg Utrechtr 19L6<br />
OPPANABMt<br />
der*-o lno Tru4. . 5<br />
un~err<br />
t.e inch99 Jaa<br />
en<br />
~rp~<br />
i<br />
~ ~ ~ r<br />
rortbO~h ofr trwil, tr--, : rtire tr,<strong>an</strong>~~~~~~~~~~~~~~~~~~~rt-in~f4etedm<br />
tkr~- ~-~:ht brow:1., ,('!'t wi';<br />
~ag<br />
irr-f~'Lr hc~riz,.ntml rineg; c~r~. 1<br />
%N0 p<br />
C.RAVfJR~rok;%s~l<br />
6,FR pCv.bl Ftrol..'<br />
Il . Feb. r ; z, 1c60<br />
"'Cf.7<br />
FIG. 67. <strong>Pourouma</strong> bicolor subsp. choco<strong>an</strong>a. Leafy twig <strong>with</strong> staminate inflorescence<br />
(Croat 8097).<br />
Specimens examined. PANAMA. Without locality,<br />
1861 (st), Hayes 32 (E), <strong>with</strong>out date (2), Hayes 348<br />
(NY). CANAL ZONE: Nr. Gamboa, 8 Feb 1966 (2), Blum<br />
2039 (MO), 19 Jun 1968 (st); Barro Colorado Isl<strong><strong>an</strong>d</strong>,<br />
22 Feb 1969 (d), Croat 8097 (F, MO, NY), (9), Croat<br />
8100 (F, MO), 14 Mar 1969 (st), Croat 8648 (MO), 23<br />
May 1970 (st), Croat 10343 (MO), 9 Jun 1970 (st),<br />
Croat 10830 (MO); nr. Gamboa, Foster et al. 584<br />
(DUKE, NY), 18 Dec 1972 (d), Gentry 6679 (MO);<br />
Gatun Lake, Gatun Locks, 27 Mar 1956 (2), Johnston<br />
1714 (A, MO, type collection of P. johnstonii); Barro<br />
Colorado Isl<strong><strong>an</strong>d</strong>, 16 Aug 1927 (st), Kenoyer 312 (US),<br />
27 Oct 1931 (st), Shattuck 260 (A, F, MO), 4 Dec 1931<br />
(d), Shattuck 522 (F, MO, US) <strong><strong>an</strong>d</strong> 525 (F, MO); N of<br />
Frijoles, 19 Dec 1923 (st), St<strong><strong>an</strong>d</strong>ley 27479 <strong><strong>an</strong>d</strong> 27502<br />
(US). CoIkN: Donese district, Camp Batija, 9-14 May<br />
1968 (9), Holdridge 6254A (MO).<br />
COLOMBIA. CHOC6: Quibdo-Istmina rd., 48 km<br />
S of Quibdo, nr. Certegui, 8 J<strong>an</strong> 1979 (9), Gentry et al<br />
23835 (BG); Quibdo-Tutunendo rd., 3 km W of Tutunendo,<br />
7 J<strong>an</strong> 1981 (st), Gentry et al. 30278 (U); junction<br />
of Rio Condoto <strong><strong>an</strong>d</strong> Rio S<strong>an</strong> Ju<strong>an</strong>, 20 Apr 1939
_' "(<br />
O^^Jgy -' i biccoolr<br />
P. bicolor subsp.<br />
-r~ E^^<br />
-<br />
_SAs.<br />
*<br />
L ?<br />
*?'7 ^ ^<br />
[<br />
*'<br />
"*^<br />
^ ~ * nde<br />
s.cbina<br />
A<strong><strong>an</strong>d</strong><br />
p. biColor subsp. digitate<br />
-<br />
P. bicolor subsp.<br />
P. bicolor *P. ccrepifo.<br />
'- s<br />
FIG. 68. Distribution of <strong>Pourouma</strong> bicolor subsp. bicolor, P. bicolor subsp. choco<strong>an</strong>a, P. bicolor subsp.<br />
digitata, P. bicolor subsp. scobina, P. bicolor subsp. tessm<strong>an</strong>nii, <strong><strong>an</strong>d</strong> P. cecropiifolia.
144 Flora Neotropica<br />
(9), Killip 35095 (F, US, type collection ofP. choco<strong>an</strong>a). 7: 231. 1927. Type. Bolivia. Beni: Rurrenabaque,<br />
NARIMO: Mun. Barbacoas, Barbacoas-Junin rd., 1050 Oct 1921 (9), Rusby 1599 (holotype, NY; isotypes,<br />
m, 7 Aug 1982 (9), Mora 2295 (US); Mun. Iscu<strong><strong>an</strong>d</strong>e, F, GH, US).<br />
Rio Sequi6n, 22 Nov 1955 (d), Romero-Cast<strong>an</strong>eda 5484<br />
(AAU). VALLE: Rio Yurum<strong>an</strong>gui, Veneral, 28 J<strong>an</strong>-10 Tree, up to 20 m tall. Leafy twigs 5-30 mm<br />
Feb 1944 (a), Cuatrecasas 15876 (F); Rio Calima, La<br />
Trojita, 19 Feb-10 Mar 1944 (2), Cuatrecasas 16315<br />
thick, (sparsely) white-pubescent <strong><strong>an</strong>d</strong> <strong>with</strong> sparse<br />
(F); La Esper<strong>an</strong>za, 6-7 Mar 1944 (d), Cuatrecasas 16755<br />
to dense, brown, pluricellular hairs, often gla-<br />
(F); S<strong>an</strong> Isidro, 2-5 May 1944 (8), Cuatrecasas 17357 brescent. Lamina 7-1 1-parted, rarely 3-5-lobed,<br />
(F); Rio Cajambre, Silva, 5-15 May 1944 (st), Cuatre- ca. 10-40 x 13-60 cm, coriaceous, apex acucasas<br />
17451 <strong><strong>an</strong>d</strong> 17479 (F); Bajo Calima, ca. 15 km N minate to acute, base deeply cordate, <strong>with</strong> or<br />
of Buenaventura, 18 Feb 1983 (st), Gentry et al. 40479 <strong>with</strong>out<br />
(BG); Bajo Calima, rd. to Ju<strong>an</strong>chaco Palmares, 18 Jul overlapping lobes; upper surface smooth,<br />
1984 (2), Gentry et al. 48303 (BG).<br />
white puberulous on the main veins <strong><strong>an</strong>d</strong> sometimes<br />
<strong>with</strong> sparse, red-brown, pluricellular hairs<br />
Vernacular names: P<strong>an</strong>ama. M<strong>an</strong>gabe. Colom- usually on the smaller veins, lower surface whitebia.<br />
Antioquia: sirpo; Nariiio: guagay. or sometimes yellow-, usually appressed-pubes-<br />
This subspecies c<strong>an</strong> be easily distinguished cent to -puberulous on the veins, occasionally<br />
from subsp. scobina by the arachnoid indument brown, pluricellular hairs on the main veins,<br />
covering the smaller veins (between the parallel arachnoid hairs in the areoles, often also on the<br />
tertiary veins), <strong><strong>an</strong>d</strong> often also by the relatively smaller veins; lateral veins ca. 15-30 pairs, terbroad<br />
segments of the lamina. Some of the spec- tiary venation almost pl<strong>an</strong>e beneath; petiole 8-<br />
imens are yellow-hirsute on the leafy twigs, or 50 cm long, glabrous or occasionally <strong>with</strong> brown,<br />
also on the stipules, petioles, main veins of the pluricellular hairs; stipules 2-20 cm long, outside<br />
lamina above <strong><strong>an</strong>d</strong>/or the peduncles. Such spec- white-appressed-puberulous <strong><strong>an</strong>d</strong> <strong>with</strong> (dense) redimens<br />
resemble P. oraria very much. The two brown, pluricellular hairs, inside densely yellowtaxa<br />
differ in the smooth upper surface of the hirsute <strong><strong>an</strong>d</strong> sometimes <strong>with</strong> brown, pluricellular<br />
lamina, the smooth fruiting peri<strong>an</strong>th, <strong><strong>an</strong>d</strong> the hairs, caducous. Staminate inflorescences up to<br />
concentration of hirsute indument at the adaxial 27 cm long <strong><strong>an</strong>d</strong> 17 cm wide; peduncle 2-11 cm<br />
side of the petiole. The resembl<strong>an</strong>ces between long, peduncle <strong><strong>an</strong>d</strong> br<strong>an</strong>ches yellow- or white-,<br />
these two taxa are so striking that we sometimes usually appressed-puberulous <strong><strong>an</strong>d</strong> usually <strong>with</strong><br />
wondered whether they could be two forms of dense, (dark- or red-)brown, pluricellular hairs;<br />
the same species (distinct from P. bicolor). flowers sessile, ? diffusely distributed along the<br />
ultimate br<strong>an</strong>ches; tepals ca. 2.5 mm long, free<br />
4. <strong>Pourouma</strong> cecropiifolia Martius in Spix & or occasionally basally connate, ciliolate; fila-<br />
Martius, Reise Bras. 3:1130. 1831, as Puruma ments shorter th<strong>an</strong> the tepals, free. Pistillate incecropiaefolia;<br />
Martius, Syst. mat. med. bras. florescences in fruit up to 24 cm long <strong><strong>an</strong>d</strong> 22 cm<br />
34. 1843; Miquel in Martius, Fl. bras. 4(1): wide; peduncle 2-13.5 cm long, peduncle <strong><strong>an</strong>d</strong><br />
123, t. 36. 1843; Macbride, Publ. Field Mus. br<strong>an</strong>ches <strong>with</strong> indument similar to that of the<br />
Nat. Hist., Bot. Ser., 13(2.2): 291. 1937. Type.<br />
staminate inflorescence; flowers 25-170; pedicel<br />
Brazil. "In silvis prov. Paraensis et Rio Ne- up to 0.6 cm, in fruit up to 2 cm long; peri<strong>an</strong>th<br />
gro," Dec 1819 (st), Martius s. n. (lectotype M,<br />
4-9 mm long; white- to yellow-puberulous <strong><strong>an</strong>d</strong><br />
chosen here). Fig. 69. usually <strong>with</strong> (dark or red-) brown, pluricellular<br />
hairs; stigma peltate, ca. 1.5-2 mm diam., sparsely<br />
white-puberulous. Fruiting peri<strong>an</strong>th dark purple<br />
to black, ovoid to subglobose, 1.5-3.5 x 0.7-2<br />
cm, densely to sparsely puberulous <strong><strong>an</strong>d</strong> sometimes<br />
<strong>with</strong> brown, pluricellular hairs.<br />
<strong>Pourouma</strong> multifida Trecul, Ann. Sci. Nat. Bot., Ser.<br />
3, 8: 107. 1847; Miquel in Martius, Fl. bras. 4(1):<br />
123, 1853. Type. Brazil? Without locality, <strong>with</strong>out<br />
date (9), Anonymus (=Ferreira?) s.n. (holotype, P;<br />
isotype, P).<br />
<strong>Pourouma</strong> sapida Karsten, Fl. Columb. 2: 19, t. 110.<br />
1862. Type. Colombia. Meta: Villavicencio, nr. Jiramene,<br />
Oct 1872 (a), Karsten s.n. (holotype, GOET;<br />
isotype?, LE-<strong>with</strong> fragments of staminate inflorescences!).<br />
<strong>Pourouma</strong> edulis Dufresne, Ill. Hort. 20: 70. 1873. Based<br />
on (living?) material from Colombia, collected by<br />
Linden).<br />
<strong>Pourouma</strong> uvifera Rusby, Mem. New York Bot. Gard.<br />
Distribution (Fig. 68). Extending from Colombia<br />
(Meta) through Amazoni<strong>an</strong> Ecuador, Peru,<br />
<strong><strong>an</strong>d</strong> Brazil (Acre <strong><strong>an</strong>d</strong> western Amazonas) to Amazoni<strong>an</strong><br />
Bolivia, also in Amazoni<strong>an</strong> Venezuela;<br />
in non-inundated forest, at altitudes up to ca.<br />
1000 m. Often cultivated <strong><strong>an</strong>d</strong> their occurrence<br />
often indicating previous hum<strong>an</strong> inhabitation.
<strong>Pourouma</strong> 145<br />
,J,<br />
? ^T \Sr~~u. I<br />
. .<br />
0f W/+ O.+ 14+ I+<br />
P OUR 0 ITMA cecropiaefolia.<br />
FIG. 69. <strong>Pourouma</strong> cecropiifolia. From Martius, Flora Brasiliensis 4(1). 1853: tab. 36. Leaf, stipule (32),<br />
staminate inflorescence, diagram of staminate flower (D), part of staminate flower (28), staminate flower (1),<br />
tepals (4), stamens (7).
146 Flora Neotropica<br />
Also cultivated in some places outside the species<br />
r<strong>an</strong>ge, e.g., Bahia (Brazil).<br />
al. 43137 (BG); 2-8 Aug 1929 (9), Killip et al. 27381<br />
(NY, US); Yurimaguas, 24-25 Aug 1929 (2), Killip et<br />
al. 27932 (F, NY, US); Iquitos, 26 Sept 1929 (2), Killip<br />
et al. 29839 (F, NY, US), Apr 1930 (6), Klug 1185 (F,<br />
Specimens examined. COLOMBIA. Without locality,<br />
<strong>with</strong>out date (9), Tri<strong>an</strong>a 862 (E, NY, P), <strong>with</strong>out NY, US), May 1930 (2), Klug 1326 (F, NY, US); Rio<br />
date (9), Tri<strong>an</strong>a 1892 (NY). AMAZONAS: Rio Boiauasu, Marafion, Persever<strong>an</strong>ca, 5 Feb 1924 (2), Kuhlm<strong>an</strong> 1337<br />
28 Nov 1945 (st), Duque-Jaramillo 2261 (NY); Rio (F, RB); Pongo de M<strong>an</strong>seriche, 11 Dec 1931 (2), Mexia<br />
Loretoyacu, Barracon, 25 Dec 1945 (2), Duque-Jara- 6257 (F, G, GB, NY, S, U, UC, US); Prov. Requena,<br />
millo 2446 (NY); between Rio K<strong>an</strong><strong>an</strong>ari <strong><strong>an</strong>d</strong> Rio Pa- nr. Jenaro Herrera, 4 Nov 1986 (2), Peters 187 (BG);<br />
coa, 1-5 Dec 1951 (6), Garcia-Barriga 13871 (US); Rio Rio Yaguasyacu, Brillo Nuevo, 16 Apr 1977 (2), Plow-<br />
Igara-Par<strong>an</strong>a, La Chorrera, 20 Sep 1973 (2), Sastre 2274 m<strong>an</strong> et al. 6903 (F, GH); Rio Yucayali, 1923 (a), Tess-<br />
(G, P); Los Monos, 24 Sep 1978 (st), Pabon E. 573 m<strong>an</strong>n 3054 (G, S); Rio Itaya, Paraiso, 9 Oct 1929 (2),<br />
(herb. Araracuara); Rio Carapar<strong>an</strong>a, El Enc<strong>an</strong>to, 22- Ll. Williams 3347 (F, US); Yurimaguas, 4 Nov 1929<br />
28 May 1942 (st), Schultes 3826 (NA), (2), Schultes (9), Ll. Williams 3984 (F), 10 Nov 1929 (2), LI. Wil-<br />
3862 (F); Rio Igara-Par<strong>an</strong>a, La Chorrera, 4-10 1942 liams 4627 (F). MADRE DE Dios: Rio M<strong>an</strong>u, Cocha<br />
(2), Schultes 3901 (F); Rio Boiauasu, Nov 1945 (2), Cashu Station, 25 Nov 1980 (2), Foster 5905 (F), 15<br />
Schultes 6805 (F, US). CAQUETA: Las Guacamayas, 28 Sep 1979 (st), Foster 7001 (F); Tambopata, 19 Feb<br />
Apr 1944 (9), Little 7756 (US); Morelia, 7 Oct 1941 1984 (st), Gentry et al 45660 (BG). PASCO: Prov. Ox-<br />
(6), von Sneidern s.n. (S). META: Villavicencio, nr. Jira- apampa, Iscozacin, 24 Jun 1982 (2), D. Smith et al.<br />
mene, Oct 1872 (9), Karsten s.n. (GOET, LE), (6), Kar- 2089 (BG). SAN MARTIN: Puerto Piz<strong>an</strong>a, 27 Jul 1973<br />
sten s.n. (GOET, LE, type collection of P. sapida). (9), J. Schunke V. 6485 (F, GH, MO, NY). UCAYALI:<br />
PUTUMAYO: Rio Putumayo, 28 Aug 1966 (9), Pinkley Prov. Coronel Portillo, Lago Yarinacocha, Povenir, 5<br />
405 (S); Puerto Lim6n, 27-28 Feb 1942 (st), Schultes Nov 1984 (2), Maas et al. 6201 (U); Prov. Coronel<br />
3343 (F). VAUPES: Mitui, 10 Sep 1939 (9), Cuatrecasas Portillo, km 13 of rd. from Campo Verde, km 34 of<br />
et al. 6737 (F); Upper Rio Guaviare, 9 Nov 1939 (6), rd. Pucallpa-Tingo Maria, 7 Nov 1984 (2), Maas et al.<br />
Cuatrecasas 7602 (F, US); Mitui, 7-8 Nov 1952 (2), 6225 (U).<br />
Humbert et al. 27221 (P, U, US); nr. Yavarate, 20 Nov BRAZIL. Without locality, <strong>with</strong>out date (2), Anon-<br />
1952 (6), Romero-Cast<strong>an</strong>eda 3647 (NY); Mitu, 26 Sep ymus (=Ferreira?) s.n. (P, type collection of P. multi-<br />
1976 (6), Zarucchi 2145 (INPA).<br />
fida). ACRE: Rd. Abuna-Rio Br<strong>an</strong>co, km 242-246, 20<br />
VENEZUELA. AMAZONAS: Cerro Duida, base of Jul 1968 (2), Forero et al. P.6419 (INPA, NY, R, S,<br />
Duida, J<strong>an</strong>-Feb 1939 (st), Farinas et al. 347 (NY, U, U); mouth of Rio Macauh<strong>an</strong>, 8 Aug 1933 (2), Krukoff<br />
VEN); Depto. Atures, 35 km SE of Puerto Ayacucho, 5327 (A, F, G, LE, M, MO, NY, S, U, UC, US), (a),<br />
25 Sep 1980 (9), Gu<strong>an</strong>chez 239 (VEN); Rio Orinoco, Krukof 5332 (A, F, G, M, MO, NY, S, U, UC); Rio<br />
between Cafno Grulla <strong><strong>an</strong>d</strong> Grulla, 8 Apr 1978 (9), Mo- Br<strong>an</strong>co, Parque Zoobot<strong>an</strong>ico da Univ. Fed., 15 Oct<br />
rillo et al. 7384 (VEN); Depto. Atapabo, nr. Culebra, 1980 (2), Nelson 705 (BG); Tarauaca, 24 Sep 1968 (2),<br />
Rio Cunucunuma, 22 Mar-4 Apr 1983 (st), Steyer- Pr<strong>an</strong>ce et al. 7503 (F, INPA, M, NY, P, R, S, U, US);<br />
mark et al. 129163 (BG, VEN).<br />
Sao Fr<strong>an</strong>cisco, Jun 1911 (2), Ule 9314 (G, MG).<br />
ECUADOR. MORONA-SANTIAGO: 5 km SW of Ma- AMAZONAS: Rio Javari, Atalaia do Norte, <strong>with</strong>out date<br />
cas, 26 J<strong>an</strong> 1981 (6), Berg 1223 (BG, U). NAPO: Tena, (2), Braga et al. 3178 (INPA); M<strong>an</strong>aus (cult.!), 15 Apr<br />
20 Oct 1939 (9), Asplund 9462 (R); S<strong>an</strong> Pablo de los 1955 (2), Chagas (INPA) 962 (INPA, U), 29 Jul 1978<br />
Secoyas, 19 Aug 1980 (9), As<strong>an</strong>za C. 32929 (AAU); (6), Falcao 215 (INPA); Marco, 29 J<strong>an</strong> 1969 (2), Croat<br />
Aii<strong>an</strong>gu, 30 May-21 Jun 1982 (st), SEF8764 <strong><strong>an</strong>d</strong> 8930 7633 (MO); Rio Japura, mouth of Rio Apaporis, 4 Dec<br />
(U); 3 km E of Atahualpa, 10 km E of Puerto Napo, 1912 (a), Ducke (MG) 12366 (MG); Rio Solimoes, Fon-<br />
13 Dec 1985 (9), Neill et al. 7044 (BG). PASTAZA: Rio teboa, 8 Jun 1945 (9), Fr6es 21048 (NY, US); Tefe, 19<br />
Curaray, Loricachi, 30 May 1980 (9), Jaramillo et al. Jul 1972 (9), Krieger et al. (INPA) 2268 (INPA); mouth<br />
31516 (AAU).<br />
of Rio Embira, 1 Jul 1933 (2), Krukoff 5109 (A, F, G,<br />
PERU. AMAZONAS: Rio Cenepa, 13 Dec 1972 (2), M, MO, NY, S, UC, US); Sao Paulo de Olivenca, 11<br />
Berlin 523 (F, GH, NY), 4 Aug 1974 (9), Berlin 1999 Sep-26 Oct 1936 (2), Krukoff8332 (BR, F, G, LE, MO,<br />
(NY). HUANUCO: Tingo Maria, 10 Aug 1940 (9), As- P, S, U, US), (2), Krukoff 8469 (A, B, BR, F, G, LE,<br />
plund 12933 (S, US), 10 Jul 1957 (st), Ellenberg 2288 MO, NY, P, S, U, US); M<strong>an</strong>aus, Estrada Roas de Maio,<br />
(U); Aucayacu, 27 Sep 1967 (9), Lao et al. s.n. (F, G, Arm<strong><strong>an</strong>d</strong>o (cult.!), 25 Sep 1973 (8), Lisboa 27 (INPA);<br />
US); Puerto Inca, 7 Dec 1968 (2), J. Schunke V. 2824 Rio Papori, Trindade, 13 Dec 1928 (st), Luetzelburg<br />
(F, G, GH, NY, US). JUNiN: La Merced, 10-24 Aug 23913 (R), 27 Dec 1928 (2), Luetzelburg23887 (M, R);<br />
1923 (9), Macbride 5446 (F, GH, US). LORETO: Iquitos, Rio Curicuriari, 8 Jul 1979 (2), Maia et al. 497 (INPA);<br />
29 Oct 1970 (2), Asplund 14124 (G, S); Zungarococha, various collections <strong>with</strong>out precise indications of lo-<br />
28 J<strong>an</strong> 1978 (st), Ayala et al. 1446 (U); Rio Ampiaco, calities (probably most of them along Rio Amazonas<br />
Puca Urquilla, 24 Feb 1972 (9), Croat 20662 (DUKE, <strong><strong>an</strong>d</strong> Rio Japurf, (Dec) 1819-(Feb) 1820 (6 or st), Mar-<br />
MO, NY, U); Iquitos, 8 Aug 1906 (9), Ducke (MG) tius s.n. (G, M, including lectotype of P. cecropiifolia);<br />
7581 (G, MG); El Sacramento, 89 km S of Pucallpa, Taparuquara, 22 Oct 1971 (2), Pr<strong>an</strong>ce et al. 15791<br />
22 Jul 1957 (st), Ellenberg 2474 (U); halfway between (INPA, NY, U), Jul-Aug 1967 (9), Schultes 24547<br />
Iquitos <strong><strong>an</strong>d</strong> S<strong>an</strong>ta Maria de N<strong>an</strong>ay, 30 May 1978 (st), (INPA); between Barcellos <strong><strong>an</strong>d</strong> Sao Gabriel, Dec 1851<br />
Gentry et al. 22460 (U); Prov. Maynas, Y<strong>an</strong>amono (2), Spruce 2023 (B, BR, CGE, P); Rio Jurua, Bom Fin,<br />
Explorama Tourist Camp, 15 Jul 1983 (st), Gentry et Oct 1900 (2), Ule 5265 (B, F, G, HBFG, MG, NY);
<strong>Pourouma</strong> 147<br />
Rio Jurua, Marary, 8 Oct 1900 (c), Ule 5266 (HBG, 1951 (e), Garcia-Barriga 14002 (holotype, US; iso-<br />
MG).<br />
type, NY).<br />
BOLIVIA. BENI: Prov. Moxos, S of Moxos, 18 Sep<br />
1976 (9), Meneces et al. 338 (MG, MO); Rurrenabaque, Tree, up to 30 m tall. Leafy twigs 4-8 mm<br />
Oct 1921 (9), Rusby 1599 (F, GH, NY, US). PANDO: thick, densely to sparsely yellow-hirsute, these<br />
Prov. Nicolas Suarez, Mukden, Jun-Dec 1979 (2), Izawa<br />
21 long hairs often intermixed <strong>with</strong> much shorter,<br />
(U).<br />
whitish hairs, sometimes also <strong>with</strong> sparse, brown,<br />
Vernacular names. Colombia. bochoa tsaha pluricellular hairs, sometimes (sub)glabrous or<br />
(Kof<strong>an</strong>), caimaron, caimaron silvestre, uva, uva subvelutinous. Lamina entire, ovate to subovate<br />
del monte, uvilla, uvo; Amazonas: guruc<strong>an</strong>a. or elliptic to oblong <strong><strong>an</strong>d</strong> 6-20 x 2.5-12 cm or<br />
Venezuela: cocura, sadajii. Peru. sacha uvilla, 3-lobed to -parted (or in subjuvenile material up<br />
uvilla, suia; Pasco: shew<strong>an</strong>taqui. Brazil: cucura, to 5-parted, sometimes <strong>with</strong> the segments almost<br />
cucuva, imbauba (or ambauva) m<strong>an</strong>sa, imbafba petiolulate) <strong><strong>an</strong>d</strong> up to ca. 25 x 25 cm, coriaceous<br />
(or ambauva) do vinho, mapati, puruma, puru- to subcoriaceous, apex acuminate, acute, apicma-y,<br />
sucuuiba, uva. Bolivia. uva silvestre; Beni: ulate, or sometimes emarginate, base subcordate<br />
t<strong>an</strong>aribe, uva del monte.<br />
to truncate to rounded to subacute; upper surface<br />
This species shows strong similarities to P. bi- ? scabrous, occasionally smooth, yellow-hirsute<br />
color, especially to the subspecies digitata <strong><strong>an</strong>d</strong> to -hirtellous on the whole surface or only on the<br />
scobina. It mainly differs in the greater number main veins, lower surface yellow-hirsute (to -hirof<br />
segments of the leaves of adult specimens (nor- tellous) to -substrigose on the main veins, white<br />
mally 7-11 versus normally 5-7 in subsp. digi- arachnoid hairs in the areoles <strong><strong>an</strong>d</strong> on the smaller<br />
tata <strong><strong>an</strong>d</strong> subsp. scobina), the smooth upper sur- veins; lateral veins 10-20 pairs, the basal pair<br />
face, <strong><strong>an</strong>d</strong> the sparser indument on the leafy twigs. br<strong>an</strong>ched, tertiary venation prominent beneath;<br />
The lamina of subsp. digitata <strong><strong>an</strong>d</strong> subsp. scobina petiole 3-17 cm long, yellow-hirsute, intermixed<br />
is sometimes smooth <strong><strong>an</strong>d</strong> subjuvenile specimens <strong>with</strong> short, whitish hairs; stipules 2-12.5 cm long,<br />
may have up to 9 leaf segments, which makes outside yellow-hirsute to -strigose, inside gladistinction<br />
of the taxa on the basis of morpho- brous, caducous. Staminate inflorescences up to<br />
logical characters sometimes impossible. For- 12 cm long <strong><strong>an</strong>d</strong> 4.5 cm wide; peduncle 3-6.5 cm<br />
tunately, the taxa are allopatric (or almost so, see long, peduncle <strong><strong>an</strong>d</strong> br<strong>an</strong>ches yellow-hirsute;<br />
p. 139).<br />
flowers sessile, rarely pedicellate, clustered, but<br />
The lectotype ofP. cecropiifolia is selected from not in globose heads; tepals almost free, (sparsenine<br />
Martius syntype collections listed above, ly) puberulent; filaments free, shorter th<strong>an</strong> the<br />
that of P. sapida from two Karsten (syntype) col- tepals. Pistillate inflorescences in fruit up to 19<br />
lections.<br />
cm long <strong><strong>an</strong>d</strong> 10.5 cm wide; peduncle 3-9 cm long,<br />
The fully mature fruiting peri<strong>an</strong>th of most (if peduncle <strong><strong>an</strong>d</strong> br<strong>an</strong>ches yellow-hirsute;flowers 20not<br />
all) <strong>Pourouma</strong> species is suitable for hum<strong>an</strong> 25; pedicel up to 0.6 cm long, in fruit up to 1<br />
consumption, but P. cecropiifolia is the only cm; peri<strong>an</strong>th 0.3-0.9 mm long, hispidulous (or<br />
species in cultivation (see p. 116).<br />
hirtellous); stigma peltate, ca. 2 mm in diam.,<br />
The fact that the species is so often found in whitish puberulous. Fruiting peri<strong>an</strong>th blue-black,<br />
(formerly) inhabited places suggests that the ellipsoid to oblongoid to subglobose, ca. 1-1.5<br />
present distribution is (at least) partly <strong>an</strong>thro- cm long, hispidulous (or hirtellous).<br />
pogenous.<br />
Distribution (Fig. 74). Colombia (Amazonas<br />
<strong><strong>an</strong>d</strong> Vaupes), Venezuela (Amazonas <strong><strong>an</strong>d</strong> Boli-<br />
5. <strong>Pourouma</strong> cucura St<strong><strong>an</strong>d</strong>ley & Cuatrecasas in var), Ecuador (Napo <strong><strong>an</strong>d</strong> Morona-S<strong>an</strong>tiago), Peru<br />
Cuatrecasas, Fieldi<strong>an</strong>a Bot. 28(1): 211. 1951. (Loreto <strong><strong>an</strong>d</strong> Madre de Dios) <strong><strong>an</strong>d</strong> Brazil (Amapa,<br />
Type. Venezuela. Amazonas: Capihuara, Alto Amazonas, <strong><strong>an</strong>d</strong> Mato Grosso), extending to east-<br />
Casiquiare, 5 Jun 1942 (Q), LI. Williams 15812 ern Venezuela (Bolivar); in non-inundated or in<br />
(holotype, F; isotypes, G, NY, RB, US). periodically inundated (varzea or igap6) forest;<br />
Fig. 70. mostly at low altitudes, sometimes (in Ecuador)<br />
up to ca. 1500 m.<br />
<strong>Pourouma</strong> garci<strong>an</strong>a Cuatrecasas, Caldasia 7:298. 1956.<br />
Type. Colombia. Amazonas-Vaupes: Rio Apaporis, Specimens examined. COLOMBIA. AMAZONAS: Rio<br />
between Rio K<strong>an</strong><strong>an</strong>ari <strong><strong>an</strong>d</strong> Rio Pacoa, 1-15 Dec Apaporis, above mouth of Rio K<strong>an</strong><strong>an</strong>ari, Soratama,
148 Flora Neotropica<br />
dl;i?~~~~~~~~~~~~~~~~~~~~~~~~~,T!<br />
~ ~ ~ ~ ~ ~ ~ ~ ~ ?~<br />
::.<br />
IWITi NAIONAL to PiQtJLAS IA AMAI7AA4A<br />
5352 ew J Starl,l. /rurourm eurura<br />
.F2;?Z,3!: -~ '; T.'l<br />
^* 1 " -'<br />
FIG.~~~~~~~~~~~~~~~~~~1<br />
'v-~~~ 70. cucra<br />
^near<br />
-<br />
rlpu , Hufb,b't C t.r. - t<br />
Rio Jurw-. 59"'?1'N: 10 ':"<br />
disturbed f"rest r"n terra firrie. ;ie, .<br />
Nx hnc. lt . W*itbh tilt. 7.t. i.- ; -<br />
gla oiua to whitte benestb, 3 partim-l r *r<br />
fol~avt., p.-dofv pweri<strong>an</strong>thb, papper & "r "<br />
inftovescence brown. Puroua~ Exuding Lefwgwihpitllat<br />
nireenesl br iw (rnce -] et :<br />
t 'nning to blacsk. "UmbinublI 'A ,' L<br />
O.T.Pr nce, F.A.ilsby, C...Berg.<br />
W.C.St_fw d, E . Lleras, T . s..,<br />
--<br />
ii____ _ _ . ctber A<br />
FIG. 70 <strong>Pourouma</strong> cu. atIg~~~~~ <strong>with</strong> pist~ilat ~<br />
flrM.. a"n: s (<br />
PA. r' <strong>an</strong>e, et.<br />
i.8<br />
l. 84)<br />
FIG. 70. <strong>Pourouma</strong> cucura. Leafy twig <strong>with</strong> pistillate inflorescences (Pr<strong>an</strong>ce et al. 18240).<br />
15-19 Dec 1951 (9), Garcia-Barriga 14106 (NY, US),<br />
18 Mar 1952 (9), Schultes et al. 15973 (F), J<strong>an</strong> 1952<br />
(6), Schultes et al. 19835 (US). AMAZONAS-VAUPES: Rio<br />
Apaporis, between Rio K<strong>an</strong><strong>an</strong>ari <strong><strong>an</strong>d</strong> Rio Pacoa, 1-<br />
15 Dec 1951 (9), Garcia-Barriga 14002 (GH, NY, US,<br />
type collection of P. garci<strong>an</strong>a).<br />
VENEZUELA. AMAZONAS: Capihuara, Alto Casiquiare,<br />
5 Jun 1942 (9), Ll. Williams 15812 (G, NY,<br />
RB, VEN, US, type collection of P. cucura). BOUVAR:<br />
Rio Tirica, Oct 1947 (9), Cardona 2170 (MO, VEN).<br />
ECUADOR. NAPO: Rio Cuyabeno, 16 Feb 1980 (8),<br />
Berg & Akkerm<strong>an</strong>s 1041 (U), Berg & Akkerm<strong>an</strong>s 1042<br />
20.
<strong>Pourouma</strong><br />
(U), 17 Feb 1980 (6), Berg & Akkerm<strong>an</strong>s 1045 (U), (2), 3-lobed leaves (similar to the common leaf shape<br />
Berg & Akkerm<strong>an</strong>s 1053 (U), 20 Feb 1980 (st), Berg in P. bicolor<br />
& Akkerm<strong>an</strong>s 1070 (U); Rio<br />
subsp. bicolor). Subjuvenile material<br />
Aguarico, Shushufindi,<br />
26 Dec 1974 (2), Vickers 86 from these two countries often have<br />
(F).<br />
up to 7-parted<br />
PERU. LORETO: Rio Ucayali, Jenaro Herrera, Jul- leaves, <strong>with</strong> the segments often petiolulate.<br />
Sep 1976 (6), Bernardi s.n. (U); Rio N<strong>an</strong>ay, Puerto The species may prove to consist of two or<br />
Almendras, 22 Feb 1979 (st), Gentry et al. 24900 (U); three morphologically readily distinguishable<br />
Rio Yaguasyacu, tributary of Rio Ampiyacu, 9 Nov<br />
1977 (6), Gentry et al. 20515 (MO, U); Rio entities,<br />
N<strong>an</strong>ay,<br />
mainly based on differences in the shape<br />
halfway between Iquitos <strong><strong>an</strong>d</strong> S<strong>an</strong>ta Maria de N<strong>an</strong>ay, 26 <strong><strong>an</strong>d</strong> incisions of the lamina. The relatively small<br />
Feb 1979 (st), Gentry et al. 25100, 25050 (U); E of number of collections prevent establishing new<br />
Tamshiyacu, below Serafin Filomeno, 19 Mar 1979 infra-specific taxa.<br />
(2), Gentry et al. 25854 (U); Rio N<strong>an</strong>ay, between Iqui- The<br />
tos <strong><strong>an</strong>d</strong> S<strong>an</strong>ta Maria, de N<strong>an</strong>ay, 23 Feb 1979 identity of the two Rosa collections from<br />
(2), Gentry<br />
et al. 31655 (U); below Serafin Filomeno, 19 Mar 1979 Amapa (Serro do Navio), consisting only of (fal-<br />
(2), Gentry et al. 25854 (U); Rio Yaguasyacu, nr. Brillo len?) leaves, is not certain. The petioles <strong><strong>an</strong>d</strong> the<br />
Nuevo, 16 Apr 1977 (st), Plowm<strong>an</strong> et al. 6906 (F, GH); main veins of the lamina beneath are densely<br />
Rio N<strong>an</strong>ay, nr. Iquitos, 14 Nov 1976 (st), Revilla 1818<br />
yellow-hirsute.<br />
(U); rd. to Pena Negra, 25 km from Iquitos, 30 Nov<br />
1976 (9), Rimachi 2724 (DUKE, F, MO, NY) Rio N<strong>an</strong>ay,<br />
above Bella Vista, 18 J<strong>an</strong> 1977 (2), Ramachi 2765 6. <strong>Pourouma</strong> cuspidata Mildbraed, Notizbl. Bot.<br />
(F, MO); rd. from Orell<strong>an</strong>a to Sta. Catalena, km 5, 9<br />
Gart. Berlin-Dahlem 10: 417.<br />
Dec 1982 (2), C. R. Rodrigues 944 (BG). MADRE 1926;<br />
DE<br />
Warburg<br />
Dios: Rd. Maldonado-Quincemil, 10 Jul 1968 (6),<br />
ex Ule, Bot. Jahrb. Syst. 40:150. 1907, nomen.<br />
Acosta M. 12 (BG, DUKE, F, NY, P); Tambopata, 22 Type. Brazil. Acre: Confluence of Rio Jurua-<br />
Feb 1984 (st), Gentry et al. 45922 (BG).<br />
Mirim <strong><strong>an</strong>d</strong> Rio Juruf, Aug 1901 (6), Ule 5719<br />
BRAZIL. ACRE: Reserva INCRA S<strong>an</strong>ta Luzia, km<br />
40, BR-364, 5-19 Oct 1984 (st), Campbell et al. 6927<br />
(lectotype B, chosen here; isolectotypes, F, G,<br />
(BG). AMAPA: Serro do Navio, P6rto Platon, 10 Oct- GH, MG).<br />
15 Dec 1976 (st), Rosa 1091 (MG), 10 Oct-15 Dec <strong>Pourouma</strong><br />
1976 (st), Rosa 1159 (MG). AMAZONAS: Rio tergoscabra Cuatrecasas, Caldasia 7: 304.<br />
Japura, 1956.<br />
Vila Bitt<strong>an</strong>court, 20 Nov 1982 (8), Amaral et al. 604<br />
Type. Colombia. Caqueta: Rio Caqueta, below<br />
mouth of Rio<br />
(BG); M<strong>an</strong>aus-Itacoatiara rd., km 26, 12 Aug 1976<br />
Orteguaza, Sol<strong>an</strong>o, 8 km SE of Tres<br />
(st), Oliveira (INPA) 60534 (INPA); Tototobi, Rio De- Esquinas, 4 Mar 1945 (v), Little et al. 9527 (holotype,<br />
mini, 25 Feb 1969 (9), Pr<strong>an</strong>ce et al. 10231<br />
F; isotypes, NY, US).<br />
(INPA, MG,<br />
MO, NY, U, US); M<strong>an</strong>aus-Itacoatiara rd., km 26, 2 Tree, up to 30 m tall. Leafy twigs 8-12 mm<br />
Oct 1965 (st), Rodrigues 8066 (INPA). MATO GROSSO:<br />
Aripu<strong>an</strong>a, 28 J<strong>an</strong> 1977 (st), Gomes et al. 614 thick, white-hirtellous to -puberulous, <strong><strong>an</strong>d</strong> <strong>with</strong><br />
(INPA),<br />
21 Feb 1977 (st), Gomes et al. 844 (INPA), 29 Mar usually rather dense, brown, pluricellular hairs.<br />
1977 (st), Gomes et al. 1040 (INPA); Aripu<strong>an</strong>a, nr. Lamina 3-lobed or 3-5 parted, 10-35 x 18-45<br />
Humboldt Centre, rd. to Rio Juruena, 8 Oct 1973 (2), cm, coriaceous, apex long acuminate, base cor-<br />
Pr<strong>an</strong>ce et al. 18240 (F, INPA, MG, MO, NY, P, S, U, date, <strong>with</strong> or <strong>with</strong>out overlapping lobes; upper<br />
US).<br />
surface smooth, hirtellous or occasionally also<br />
Vernacular names. Venezuela. Amazonas: cu- <strong>with</strong> brown, pluricellular hairs on the main veins,<br />
cura; Bolivar: kaibarikei (Arekuna), sarasara lower surface strigose, main veins hispidulous<br />
(Camaracoto). Ecuador. S<strong>an</strong>tiago-Zamora: guar- <strong><strong>an</strong>d</strong> sometimes <strong>with</strong> brown, pluricellular hairs,<br />
umo colorado. Peru. Loreto: dacha uvillos, uvil- arachnoid hairs confined to the areoles; lateral<br />
la; Brazil. Amazonas: imbaubar<strong>an</strong>a br<strong>an</strong>ca; veins 13-30 pairs, tertiary venation + prominent<br />
Amapa: imbaubar<strong>an</strong>a br<strong>an</strong>ca, inbaubar<strong>an</strong>a folha beneath; petiole 11-40 cm long, white-hispidupeluda;<br />
Mato Grosso: imbaiuba or umbauiba, im- lous, <strong><strong>an</strong>d</strong> <strong>with</strong> brown, pluricellular hairs; stipules<br />
baubar<strong>an</strong>a.<br />
7-13 cm long, outside white-appressed-puberu-<br />
The species is very variable. The long, yellow lous <strong><strong>an</strong>d</strong> <strong>with</strong> brown, pluricellular hairs, inside<br />
hairs on the leafy twigs, petioles <strong><strong>an</strong>d</strong> stipules vary sparsely white-puberulous to subglabrous, cafrom<br />
very dense to sparse. The leaves of (adult) ducous. Staminate inflorescences up to 11 cm<br />
specimens collected in Colombia, <strong><strong>an</strong>d</strong> Venezuela long <strong><strong>an</strong>d</strong> 4.5 cm wide; peduncle 4-6 cm long,<br />
<strong><strong>an</strong>d</strong> in Amapa <strong><strong>an</strong>d</strong> Amazonas (Brazil) normally (densely) white-puberulous <strong><strong>an</strong>d</strong> <strong>with</strong> (dense),<br />
have entire <strong><strong>an</strong>d</strong> elliptic leaves. The collections brown, pluricellular hairs; flowers sessile, + difmade<br />
in Mato Grosso (Brazil) have on the same fusely distributed along the ultimate br<strong>an</strong>ches;<br />
twig entire <strong><strong>an</strong>d</strong> 3-parted leaves. The material from tepals ca. 1 mm long, free or basally connate,<br />
Ecuador <strong><strong>an</strong>d</strong> Peru mostly have ovate <strong><strong>an</strong>d</strong> often sparsely hairy; filaments shorter th<strong>an</strong> the tepals.<br />
149
150 Flora Neotropica<br />
Pistillate inflorescences in fruit up to 12.5 cm<br />
long <strong><strong>an</strong>d</strong> 8 cm wide; peduncle 4-6 cm long, peduncle<br />
<strong><strong>an</strong>d</strong> br<strong>an</strong>ches <strong>with</strong> indument similar to<br />
that of the staminate inflorescences; flowers 10-<br />
35; pedicel 0.2-0.4 cm, in fruit up to 0.8 cm long;<br />
peri<strong>an</strong>th 2-6 mm long, hispidulous <strong><strong>an</strong>d</strong> <strong>with</strong><br />
brown, pluricellular hairs; stigma peltate, ca. 1.5<br />
mm diam., white puberulous. Fruiting peri<strong>an</strong>th<br />
ovoid to ellipsoid, 12-15 x 5-11 mm, scabrous.<br />
prominent beneath; petiole 8-27 cm long, whitish-hirsute<br />
to -hirtellous to -subhispid, at the base<br />
swollen <strong><strong>an</strong>d</strong> saccate; stipules 2-7.5 cm long, outside<br />
yellowish-hirsute, inside sparsely hairy, ca-<br />
Distribution (Fig. 74). Colombia (Caqueta) Ecuador<br />
(Napo), Peru (Pasco), <strong><strong>an</strong>d</strong> Brazil (Acre <strong><strong>an</strong>d</strong><br />
Amazonas); in non-inundated forest.<br />
Specimens examined. COLOMBIA. CAQUETA: Sol<strong>an</strong>o,<br />
8 km SE of Tres Esquinas, Rio Caqueta, below<br />
mouth of Rio Orteguaza, 4 Mar 1945 (2), Little et al.<br />
9527 (F, GH, NY, US, type collection of P. tergoscabra).<br />
ECUADOR. NAPO: Afi<strong>an</strong>gu, 30 May-21 Jun 1982<br />
(st), SEF 8532 (U).<br />
PERU. PASCO: Prov Oxapampa, Rio Iscozacin, Cabeza<br />
de Mono, 12 Jun 1983 (st), Gentry et al. 41890<br />
(BG).<br />
BRAZIL. ACRE: Reserva INCRA S<strong>an</strong>ta Luzia, km<br />
40, BR-364, 5-19 Oct 1984 (st), Campbell et al. 6834<br />
(BG); confluence of Rio Jurua-Mirim <strong><strong>an</strong>d</strong> Rio Jurua,<br />
Aug 1901 (6), Ule 5719 (B, F, G, GH, MG, lectotype<br />
collection ofP. cuspidata), Aug 1901 (2), Ule 5719 (B,<br />
G). AMAZONAS: Sao Paulo de Olivenca, 11 Sep-26 Oct<br />
1936 (Q), Krukoff 8373 (A, F, NY, S, U).<br />
ducous. Staminate inflorescences up to 12 cm<br />
long <strong><strong>an</strong>d</strong> 6 cm wide; peduncle 1-3.5 cm long,<br />
peduncle <strong><strong>an</strong>d</strong> br<strong>an</strong>ches puberulous or partly hirtellous;flowers<br />
sessile, clustered in globose heads,<br />
these 2-3 mm diam.; tepals ca. 0.5-1 mm long,<br />
basally connate, hirtellous; filaments shorter th<strong>an</strong><br />
the tepals. Pistillate inflorescences br<strong>an</strong>ched, in<br />
fruit up to 8 cm long <strong><strong>an</strong>d</strong> 7 cm wide; peduncle<br />
3-4 cm long, indument or peduncle <strong><strong>an</strong>d</strong> br<strong>an</strong>ches<br />
similar to that of the staminate inflorescences;<br />
flowers 8-ca. 30; pedicel up to 0.3 cm long, in<br />
fruit up to 0.5 cm; peri<strong>an</strong>th 0.3-0.4 cm long,<br />
(densely) hirsute-hirtellous, usually arachnoid<br />
hairs near the apex; stigma peltate, ca. 1.5 mm<br />
diam., hirtellous. Fruitingperi<strong>an</strong>th purple, ovoid<br />
to ellipsoid, 1-1.5 cm long, puberulous, also<br />
sparsely yellow-hirsute, sometimes <strong>with</strong> brown,<br />
pluricellular hairs.<br />
Distribution (Fig. 74). Colombia (Amazonas),<br />
Peru (Loreto), <strong><strong>an</strong>d</strong> Brazil (Amazonas); in noninundated<br />
forest.<br />
This species, characterized by incised leaves<br />
<strong><strong>an</strong>d</strong> the smooth upper surface, but scabrous low-<br />
er surface of the lamina, may be related to P.<br />
bicolor.<br />
Ule collected staminate <strong><strong>an</strong>d</strong> pistillate speci-<br />
mens under the same number 5719. The sta-<br />
minate collection is selected as the lectotype col-<br />
lection.<br />
7. <strong>Pourouma</strong> myrmecophila Ducke, Bull. Mus.<br />
Hist. Nat. (Paris), Ser. 2, 4: 723. 1932. Type.<br />
Brazil. Amazonas: M<strong>an</strong>aus, Cachoeira do<br />
Mindui, 8 Jul 1929 (d), Ducke (RB) 23607 (lec-<br />
totype, RB, chosen here; isolectotypes, G, P,<br />
S, US). Fig. 71.<br />
Tree, up to 8 m tall, or shrub, myrmecophilous.<br />
Leafy twigs 0.6-1 cm thick, densely yellow(ish)-hirsute,<br />
glabrescent, <strong><strong>an</strong>d</strong> <strong>with</strong> reddishbrown,<br />
pluricellular hairs. Lamina 3-5-lobed, to<br />
-fid, 13-38 x 15-42 cm, subcoriaceous to chartaceous,<br />
rarely coriaceous, midsegment relatively<br />
long, apex acuminate, base subcordate to deeply<br />
cordate; upper surface scabrous, yellow-hirsute<br />
to -hirtellous on the main veins, elsewhere<br />
sparsely so or only hispidulous, lower surface<br />
pale yellow-hirtellous on the veins, white, arach-<br />
noid hairs in the areoles or sometimes also on<br />
the smaller veins; lateral veins 14-17 pairs, the<br />
basal pair usually br<strong>an</strong>ched, tertiary venation<br />
Specimens examined. COLOMBIA. AMAZONAS:<br />
C<strong>an</strong>to Aduche, 12 Nov 1981 (2), La Rotta 101 (herb.<br />
Araracuara), 4 Mar 1982 (2), La Rotta 139 (herb. Araracuara).<br />
PERU. LORETO: 13 J<strong>an</strong> 1976 (2), Gentry et al. 15876<br />
(U); Puerto Almendras, 11 Nov 1984 (2), Maas et al.<br />
6249 (U), 13 J<strong>an</strong> 1976 (2), McD<strong>an</strong>iel et al. 20436 (U);<br />
Rio N<strong>an</strong>ay, Puerto Almendras, 29 Nov 1976 (2), Revilla<br />
1966 (U).<br />
BRAZIL. AMAZONAS: Mun. Tefe, Lago de Tefe, 14<br />
Dec 1982 (8), Amaral et al. 76113 (BG); Rio Cuieras,<br />
mouth of Rio Br<strong>an</strong>chinho, Sep 1973 (st), Berg 276<br />
(MO, U); upper Rio Alalau, 24 Feb 1968 (2), Bov<strong>an</strong><br />
267 (INPA); BR-17, km 11, 4 Aug 1955 (6), Chagas<br />
(INPA) 1564 (INPA, U); M<strong>an</strong>aus-Mindfi rd., 3 Nov<br />
1955 (9), L. Coelho (INPA) 2265 (INPA); Mun. Jutai,<br />
Copessfi, nr. P6rto Alfonso, 24 Oct 1986 (2), Daly et<br />
al. 4129 (BG), 24 Oct 1986 (6), Daly et al. 4135 (BG);<br />
M<strong>an</strong>aus, Aleixo rd., 29 Dec 1942 (9), Ducke 1149 (MG,<br />
MO, NY, R, US), 23 Sep 1945 (6), Ducke 1764 (A, F,<br />
MG, NY, R, US), 19 Sep 1929 (2), Ducke (RB) 23606<br />
(RB, US); M<strong>an</strong>aus, Cachoeira do Mindu, 8 Jul 1929<br />
(8), Ducke (RB) 23607 (G, P, RB, S, lectotype collection<br />
of P. myrmecophila); M<strong>an</strong>aus, BR-17, km 3, <strong>with</strong>out<br />
date (a), Fr<strong>an</strong>cisco (INPA) 2186 (INPA); nr. P6rto Velho,<br />
8 Sep 1923 (a), Kuhlm<strong>an</strong>n (RB) 19843 (B, U);
<strong>Pourouma</strong><br />
~~~~~~~~~~~~~~~~~~~~~~~~..<br />
44deS A._ S 4 V<br />
?Cm Ira= BDT~OANWAL GARDENN<br />
VMVMW~T NACIONAL DE PESQUMA%A DA AMAZ6.%-I<br />
gftt. at A? .m<br />
-Gd , - v~~~~~~~~~~~~~T?e m e h. y tg<br />
' S Anto NAntni<br />
,24313<br />
x 22 d-.meterl de pAbonIariA ,M-.<strong>an</strong>aus-I<br />
Crcrintuits 220 aForest on terra<br />
Pouroum&<br />
? mateat<br />
S:nto Antonio de Abonari.<br />
brown.<br />
C rac&raj, Km 220. Forest<br />
I:<strong>an</strong>aur,<strong>an</strong><br />
terra<br />
br&ct brown.<br />
Prne,RJ<br />
Negat've<br />
G.T.<br />
ime<br />
ol. vember 2619?<br />
iach<br />
L,F,~L.F Coelho- J.F. Ramosj_ ... _ .ln:<br />
.% '1??~ 1 1 1 55551 nr. C=ll 21 cr 31 | 51 wf *-ffi0st I....<br />
rwr ar<br />
1w JtPAN n^<br />
:<br />
21RK 3F1N41<br />
71i~<br />
FIG. 71. <strong>Pourouma</strong> myrmecophila. LeafS twig <strong>with</strong> pistillate inflorescences (Pr<strong>an</strong>ce et al. 24313).<br />
M<strong>an</strong>aus, Reserva Florestal Ducke, 6 Dec 1976 (st),<br />
Nascimento et al. (INPA) 66396 (INPA); M<strong>an</strong>aus-Itacoatiara<br />
rd., km 192, 13 Sep 1974 (6), Pennington et<br />
al. P.22647 (INPA, NY); Humaita-Labrea rd., km 80,<br />
27 J<strong>an</strong> 1970 (9), Pr<strong>an</strong>ce et al. 3253 (F, INPA, NY, R,<br />
151<br />
S, U, US); Rio Cuieras, mouth of Rio Br<strong>an</strong>cinho, 24<br />
Sep 1971 (d), Pr<strong>an</strong>ce et al. 14795 (F, INPA, NY, U);<br />
Rio Javari, Attalaia, 14 Oct 1976 (9), Pr<strong>an</strong>ce et al.<br />
23776 (G, INPA, MG, NY, U); M<strong>an</strong>aus-Caracarai rd.,<br />
km 220, S<strong>an</strong>ta Antonio de Abonari, 26 Nov 1976 (2),
152 Flora Neotropica<br />
Pr<strong>an</strong>ce et al. 24313 (INPA, MO, NY, S, U, US); Ma- peri<strong>an</strong>th (sparsely) hirsute; stigma peltate, ca. 1.5<br />
naus-Itacoatiara rd., km 79, 1 Sep 1965 (8), Rodrigues cm diam., subhirsute. Fruiting peri<strong>an</strong>th ovoid to<br />
et al. 7069 (U), km 74, 3 Sep 1965 (d), Rodrigues et<br />
al. 7076 (INPA).<br />
ellipsoid, ca. 1.5 cm long, blue-black, yellowhirsute.<br />
Vernacular names. Brazil. Amazonas: embau- Distribution (Fig. 74). Brazil (Amazonas); in<br />
bar<strong>an</strong>a or imbafbar<strong>an</strong>a.<br />
non-inundated forest.<br />
From the syntype collections of P. myrmecophila,<br />
Ducke (RB) 23606 <strong><strong>an</strong>d</strong> 23607 <strong><strong>an</strong>d</strong> Kuhl- Specimens examined. BRAZIL. AMAZONAS: Mun.<br />
Sao Paulo de Olivenca, Rio J<strong><strong>an</strong>d</strong>iatuba, 27 Nov 1986<br />
m<strong>an</strong>n (RB) 19843, Ducke (RB) 23607, is chosen (9), Daly et al. 4456 (BG); nr. Sao Paulo de Olivenca,<br />
as the lectotype collection.<br />
5 Dec 1986 (8), Daly et al. 4485 (BG); Ton<strong>an</strong>tins, Rio<br />
The collection Kuhlm<strong>an</strong>n (RB) 150082, ac- Solim6es, 7 Feb 1944 (9), Ducke 1917 (F, NY, R, type<br />
cording to the label from Rio de J<strong>an</strong>eiro, is collection of P.<br />
prob-<br />
formicarum); Sao Antonia de Ia, 2<br />
ably either from Amazonas or was cultivated in<br />
May 1945 (8). Fr6es 20870 (NY, US), 19 Aug 1973<br />
(8). Lleras et al. P.17411 (F, INPA, MO, NY, S, U,<br />
Rio de J<strong>an</strong>eiro.<br />
US); Carauari, Poco Jurua, 7 Jul 1980 (8), A. S. Silva<br />
Vegetative parts (e.g., the wood) are (strongly) et al. 476 (BG, INPA).<br />
aromatic, smelling like spearmint or, according<br />
to other labels, like<br />
Vernacular names. Brazil. Amazonas: imbai-<br />
"bengue."<br />
binha, mapati.<br />
This species matches P. myrmecophila in the<br />
8. <strong>Pourouma</strong> formicarum Ducke, Trop. Woods swollen <strong><strong>an</strong>d</strong> saccate base of the petiole, a feature<br />
90: 9. 1947. Type. Brazil. Amazonas: Rio So- connected <strong>with</strong> myrmecophily. The two species<br />
limoes, Ton<strong>an</strong>tins, 7 Feb 1944 (9), Ducke 1917 also show strong resembl<strong>an</strong>ces in the indument,<br />
(not 1916!) (holotype, R, not indicated by staminate inflorescences <strong><strong>an</strong>d</strong> flowers in the pis-<br />
Ducke, designated here; isotypes, F, NY). tillate flowers. They are quite distinct in the shape<br />
Fig. 72. of the lamina <strong><strong>an</strong>d</strong> in the length of the petiole.<br />
Tree, up to ca. 10 m<br />
"The<br />
tall, or<br />
br<strong>an</strong>chlets are inhabited<br />
shrub, myrmeby<br />
stinging 'jicophilous.<br />
Leafy twigs ca. 6 mm quitaia' <strong>an</strong>ts which nest<br />
thick, yellowamong<br />
the hairs" (Daly<br />
to<br />
et<br />
brownish-hirsute <strong><strong>an</strong>d</strong><br />
al.<br />
<strong>with</strong> sparse, brown 4456).<br />
pluricellular<br />
hairs. Lamina entire, subovate, (14-) 18-<br />
32 x 6-12<br />
9.<br />
cm, chartaceous<br />
<strong>Pourouma</strong><br />
to subcoria-<br />
phaeotricha Mildbraed, Notizbl.<br />
Bot. Gart. Berlin-Dahlem<br />
ceous, apex abruptly short-acuminate, base<br />
10:193.1927.<br />
ob-<br />
Type.<br />
tuse to<br />
Peru. Loreto:<br />
subacute; upper surface scabrous to sca-<br />
Iquitos, 20 Aug 1925 (2), Tessm<strong>an</strong>n<br />
5364<br />
bridulous, yellow-hirsute, lower surface +<br />
(holotype, B; isotypes, S,<br />
densely<br />
US).<br />
yellow-hirsute on the main veins to (very) sparse-<br />
Fig. 73.<br />
ly hirsute on the smaller veins, arachnoid hairs Tree, up to 10 m tall. Leafy twigs 4-6 mm<br />
in the areoles <strong><strong>an</strong>d</strong> on the smaller veins; lateral thick, yellow-sericeous to -hirtellous, <strong><strong>an</strong>d</strong> <strong>with</strong><br />
veins 10-20 pairs, the basal pair unbr<strong>an</strong>ched, dense long, brown pluricellular hairs. Lamina<br />
tertiary venation slightly prominent beneath; entire, oblong to obovate, 8-23 x 3-9 cm, copetiole<br />
1-1.5 cm long, yellow-hirsute, at the base riaceous to subcoriaceous, apex acuminate, base<br />
swollen <strong><strong>an</strong>d</strong> saccate; stipules 3-5 cm long, outside acute to rounded; upper surface smooth or scayellow-hirsute,<br />
inside glabrous, caducous. Sta- brous, yellow-hirtellous on the main veins, lower<br />
minate inflorescences up to 12 cm long <strong><strong>an</strong>d</strong> to 6 surface yellow-hirsute on the veins <strong><strong>an</strong>d</strong> sparse or<br />
cm wide; peduncle 2-4 cm long, peduncle <strong><strong>an</strong>d</strong> sometimes dense, or<strong>an</strong>ge, pluricellular hairs,<br />
br<strong>an</strong>ches yellow-hirtellous to -hirsute; flowers arachnoid hairs confined to the areoles or almost<br />
sessile, clustered in globose heads, these 2-3 mm absent; lateral veins 8-10 pairs, basal pair unin<br />
diam.; tepals ca. 1 mm long, almost free, yel- br<strong>an</strong>ched, tertiary venation prominent beneath;<br />
lowish-hirtellous; filaments shorter th<strong>an</strong> the te- petiole 2-6 cm long, indument similar to that of<br />
pals. Pistillate inflorescences br<strong>an</strong>ched, in fruit the twigs; stipules 1-4.5 cm long, outside yellowup<br />
to 7.5 cm long <strong><strong>an</strong>d</strong> 3.5 cm wide; peduncle ca. sericeous to -velutinous, inside sparsely yellow-<br />
4 cm long, peduncle <strong><strong>an</strong>d</strong> br<strong>an</strong>ches <strong>with</strong> indument hairy to glabrous, caducous. Staminate infloressimilar<br />
to that of the staminate inflorescences; cences up to 10 cm long <strong><strong>an</strong>d</strong> 6 cm wide; peduncle<br />
flowers 5-6; pedicel in fruit up to 0.5 cm long; 1.5-3 cm long, white- to yellow-sericeous to -ve-
<strong>Pourouma</strong> 153<br />
FG<br />
R.:<br />
17. 411...<br />
. ur Uri? fr)i' -r : *<br />
If*,, r, , '*^>r ,, i /'.:<br />
X>.t 'i. E.< L: . ?J.1 W.C...<br />
. c O k ..y .<br />
penin * .l ei<br />
P Cul betLw. *.<br />
;i5 Behi m Ant r .i e J .<br />
72_.. Poru a'o1r1ma1m.11 1ea i 11g w h s i InfloFre"nce tiete. s e<br />
'__A 35<br />
212*.<br />
likl , 1 . -.<br />
i* ......<br />
FIG. 72. <strong>Pourouma</strong>formicarum. Leafy twig <strong>with</strong> staminate inflorescences (Lleras et al. P.17411).<br />
lutinous, <strong><strong>an</strong>d</strong> <strong>with</strong> dense (but on the ultimate<br />
br<strong>an</strong>ches sparse) brown, pluricellular hairs;flowers<br />
sessile, clustered in globose heads, these ca.<br />
2 mm diam.; tepals ca. 0.7 mm long, free or<br />
basally connate, hairy; filaments shorter th<strong>an</strong> the<br />
petals. Pistillate inflorescences in fruit up to 9 cm<br />
long <strong><strong>an</strong>d</strong> 8 cm wide; peduncle 2-5 cm long, peduncle<br />
<strong><strong>an</strong>d</strong> br<strong>an</strong>ches yellow- to white-hirtellous<br />
to sericeous; flowers 11-15; pedicel 0.1-0.2 cm<br />
long, in fruit up to 0.9 cm; peri<strong>an</strong>th ca. 2 mm
154 Flora Neotropica<br />
<strong>Pourouma</strong> phaeotricha<br />
HAtM OF PLRU<br />
mildbr.<br />
Maynas: Dtto. Iquitos. lio :lar.na,<br />
Caserio de<br />
IItjt<br />
Iish<strong>an</strong>a trail to :<br />
|tg N ~ative i ?nch 1 2 de Yar<strong>an</strong>a, upl<strong><strong>an</strong>d</strong>s, I( :,<br />
"uvilla",.<br />
green.<br />
FIG. 73. Porio p ec fL.<br />
:- R~oX// r-5355S 1 I 21 31 41 5 Dececber 4, 1976<br />
/ . -,,,,,,,I,,,h,,,l.,,I,,,, II all C M<strong>an</strong>uel Rimachi Y. 2725<br />
FIG. 73. <strong>Pourouma</strong> phaeotricha. Leafy twigs <strong>with</strong> pistillate inflorescences<br />
(Rimachi Y. 2725).
<strong>Pourouma</strong> 155<br />
long, yellow-velutinous, <strong>with</strong> long, brown, pluri- mostly also on the smaller veins, occasionally<br />
cellular hairs; stigma peltate, ca. 1.5 mm diam., also on the leaf margin; lateral veins (6-)10-25<br />
yellow-puberulous. Fruiting peri<strong>an</strong>th ovoid to al- (-30) pairs, basal pair br<strong>an</strong>ched or sometimes (in<br />
most ellipsoid, 1.1-1.5 x 0.7-0.9 cm long, pu- obovate to elliptic leaves) unbr<strong>an</strong>ched, the basal<br />
berulous.<br />
part of these veins forming the basal part of the<br />
Distribution (Fig. 85). Peru (Loreto) <strong><strong>an</strong>d</strong> Brazil leaf margin (thus not separated from the margin<br />
(Amazonas, along Rio Javari); in non-inundated by mesophyll), reticulum <strong><strong>an</strong>d</strong> tertiary venation<br />
forest.<br />
(rather) pl<strong>an</strong>e beneath; petiole 3-22(-35) cm long,<br />
sparsely to densely white-puberulous (to -subve-<br />
Specimens examined. PERU. LORETO: Rio N<strong>an</strong>ay,<br />
Casario de Mish<strong>an</strong>a 3 Dec 1976 (9), Davidson 5203 lutinous) to yellow-hirsute, occasionally also <strong>with</strong><br />
(MO, U, US), 4 Dec 1976 (9), Rimachi 2725 (DUKE, long white, arachnoid hairs; stipules 3-16 cm long,<br />
F, NY); Iquitos, 3 Dec 1976 (9), Tessm<strong>an</strong>n 5364 (B, outside yellow(ish)-(sub)velutinous to -hirsute to<br />
S, US, type collection of P. phaeotricha). -(sub)sericeous, inside glabrous or sometimes<br />
BRAZIL. AMAZONAS: Rio Javari, Estirao do Ecuador,<br />
8 Aug 1973 (8), Lleras et al. P.17242 (F, INPA,<br />
sparsely hairy, caducous. Staminate inflores-<br />
MO, NY, U, US), 21 Oct 1976 (v), Pr<strong>an</strong>ce et al. 23966 cences up to 10 cm long <strong><strong>an</strong>d</strong> 5 cm wide; peduncle<br />
(G, INPA, NY, U).<br />
2-6 cm long, peduncle <strong><strong>an</strong>d</strong> br<strong>an</strong>ches whitish- to<br />
yellow-puberulous<br />
Vernacular names. Peru. Loreto: Sacha<br />
(to -subvelutionous) to -hiruvilla,<br />
tellous;flowers sessile or pedicellate, most of them<br />
uvilla.<br />
in<br />
The features of the staminate inflorescences<br />
globose heads, these 3-4 mm diam.; tepals<br />
0.5-1 mm<br />
<strong><strong>an</strong>d</strong> flowers <strong><strong>an</strong>d</strong> the pistillate flowers suggest close<br />
long, connate, forming a ? urceolate<br />
peri<strong>an</strong>th,<br />
relationships to P. heterophylla <strong><strong>an</strong>d</strong> P.<br />
hairy; filaments longer th<strong>an</strong> the periformica<strong>an</strong>th,<br />
free. Pistillate<br />
rum. However, P. phaeotricha lacks the swollen<br />
inflorescences in fruit up to<br />
18 cm<br />
petiole base.<br />
long <strong><strong>an</strong>d</strong> 10 cm wide; peduncle 3-12 cm<br />
The two collections from the<br />
long, peduncle <strong><strong>an</strong>d</strong> br<strong>an</strong>ches <strong>with</strong> indument sim-<br />
Brazili<strong>an</strong> b<strong>an</strong>k<br />
ilar to that of the staminate<br />
of Rio Javari have leaves <strong>with</strong> a scabrous<br />
inflorescence;flowers<br />
upper<br />
surface in<br />
10-30, in two clusters or more<br />
contrast to the other collections.<br />
diffusely distributed<br />
in the inflorescence; pedicel up to 1 cm, in<br />
fruit<br />
10. <strong>Pourouma</strong> mollis Trecul, Ann. Sci. Nat.<br />
up to 1.5 cm long, often already at <strong>an</strong>thesis<br />
Bot., <strong>with</strong> a<br />
Ser. 3, 8: 102, 1847; Miquel in Martius, Fl.<br />
strongly (+ capula-like) widened apex;<br />
bras. 4(1): 127. 1853; Berg, Fl. Suriname<br />
peri<strong>an</strong>th 0.5-1 cm long, yellow(ish)-(sub)ve-<br />
5(1):<br />
272, t. 15b. 1975. Type. French Gui<strong>an</strong>a.<br />
lutinous,<br />
Withapex<br />
? tuberculate; stigma peltate, ca.<br />
2 mm<br />
out locality, <strong>with</strong>out date (2), Leprieur s.n.<br />
diam., white-puberulous. Fruiting peri-<br />
(lec- <strong>an</strong>th black or<br />
totype, P. chosen by Berg, Fl. Suriname<br />
purplish, ovoid to ellipsoid or to<br />
5(1):<br />
272.<br />
oblongoid, 1-2 x 0.7-1.5 cm, usually<br />
1975;<br />
densely<br />
isolectotype, G).<br />
hairy.<br />
This species is very closely related to P. melinonii.<br />
We had considered recognizing the two<br />
taxa only at the subspecific level, but the lack of<br />
intermediates in areas where the two taxa grow<br />
together indicates that they are probably good<br />
species.<br />
Two allopatric subspecies c<strong>an</strong> be recognized.<br />
Tree, up to 30 m tall. Leafy twigs 4-10 mm<br />
thick, rather sparsely to densely white-puberu-<br />
lous (to -subvelutinous) to yellow-hirsute, <strong><strong>an</strong>d</strong><br />
<strong>with</strong> sparse to dense, purple to reddish-brown,<br />
pluricellular hairs, occasionally sparse, white,<br />
arachnoid hairs. Lamina entire, ovate to elliptic<br />
(or obovate), 6-25(-35) x 4-20(-30) cm or 3-5-<br />
lobed to -parted, ca. 10-25(-40) x 10-25(-40)<br />
cm, coriaceous (to subcoriaceous), apex acumi-<br />
nate, base acute, rounded to truncate, or sub-<br />
cordate; upper surface smooth, sometimes sca-<br />
brous, hairy on the main veins or subglabrous,<br />
occasionally yellow-hirsute, lower surface,<br />
sparsely to densely puberulous to hirtellous to<br />
subtomentose, often part of the hairs on the main<br />
veins appressed, arachnoid hairs in the areoles,<br />
Key to the Subspecies of<br />
<strong>Pourouma</strong> mollis<br />
1. Lamina usually entire; pistillate inflorescences<br />
<strong>with</strong> the flowers in two, more or less distinct<br />
clusters; Gui<strong>an</strong>as, Lower Amazon Basin, <strong><strong>an</strong>d</strong><br />
eastern Brazil. ................ a. subsp. mollis.<br />
1. Lamina usually 3-5 parted; pistillate inflorescences<br />
usually <strong>with</strong> the flowers diffusely distrib-
156 Flora Neotropica<br />
uted; Upper Amazon Basin to eastern Colombia.<br />
............................ b. subsp. triloba.<br />
lOa. <strong>Pourouma</strong> mollis Trecul subsp. mollis.<br />
Fig. 75b-f.<br />
Lamina usually entire, sometimes 3-lobed or<br />
to 5-parted, subjuvenile material usually 3-5-<br />
parted; upper surface smooth, <strong>with</strong> hairs on the<br />
main veins or subglabrous. Pistillateflowers ar-<br />
r<strong>an</strong>ged in two + distinct clusters, due to the<br />
shortening of one (or two) of the primary br<strong>an</strong>ch-<br />
es <strong><strong>an</strong>d</strong> the short pedicels, being at fruit up to 0.7<br />
cm long.<br />
Distribution (Fig. 74). Eastern Guy<strong>an</strong>a, Suri-<br />
nam, French Gui<strong>an</strong>a, Amazoni<strong>an</strong> Brazil<br />
(Amampa <strong><strong>an</strong>d</strong> Para), <strong><strong>an</strong>d</strong> eastern Brazil (Bahia<br />
<strong><strong>an</strong>d</strong> Espirito S<strong>an</strong>to); in non-inundated forest at<br />
low altitudes.<br />
1978 (6), DAF 027 (GUA); Reserva de Linhares, 30<br />
Oct 1930 (6), Kuhlm<strong>an</strong>n 446 (U); Linhares, Corriogo<br />
de Durao, 2 J<strong>an</strong> 1972 (9), Sucre 8303 (U). PARA: Faro,<br />
6 J<strong>an</strong> 1920 (6), Ducke (RB) 13058 (RB); Obidos, 27<br />
Dec 1913 (9), Ducke (MG) 15261 (MG); Belem, 1 Feb<br />
1928 (6), Ducke (RB) 19455 (RB); Rio Guama; Sao<br />
Miguel, Oct 1906 (6), Goeldi (MG) 7731 (G, MG, S,<br />
U); Acarai Mts., Rio Taruini, 20 Nov 1952 (6), Guppy<br />
656 (=FD 7671) (NY, US); Belem, 19 Sep 1963 (6),<br />
Oliveira 2597 (IAN), 26 Sep 1963 (9), Oliveira 3071<br />
(IAN), 3 Sep 1966 (3), Pires 10123 (IAN), 24 Aug 1967<br />
(), Pires et al. 10578 (IAN), 2 Sep 1967 (6), Pires et<br />
al. 10809 (IAN), Sep-Oct 1961 (9), Pires 51676 (F,<br />
NY, US); Cuiaba-S<strong>an</strong>tar6m rd., km 919,13 Nov 1977<br />
(9), Pr<strong>an</strong>ce et al. 25337 (F, MG, RB, U); Mun. Almeirim,<br />
Monte Dourado, 2 Dec 1978 (6), S<strong>an</strong>tos 455<br />
(U); Belem, 12 J<strong>an</strong> 1966 (6), M. Silva 354 (MG), 4 Oct<br />
1942 (6), M. B. Silva 130 (NY, US).<br />
Vernacular names. Surinam. Boroma (Arowak),<br />
bospapaja, gr<strong>an</strong>boesipapaja, yarayara<br />
(Carib). French Gui<strong>an</strong>a. Bois c<strong>an</strong>on male, bois<br />
Specimens examined. GUYANA. Essequibo R., nr.<br />
c<strong>an</strong>on sauvage. Brazil. Amapa: uva de macaco.<br />
Rockstone, 1 Aug 1921 (juv), Gleason 670 (GH, NY, Bahia: itarar<strong>an</strong>ga, tarar<strong>an</strong>ga, tarar<strong>an</strong>ga vermel-<br />
US), 4 Nov 1979 (2), Maas et al. 3938 (U).<br />
ha; Mar<strong>an</strong>hao: ama'yrary (Ka'apor); Para: im-<br />
SURINAM. Peninica Creek, 20 Apr 1950 (st), BW bauiba, imbauibar<strong>an</strong>a vermelha, mapatir<strong>an</strong>a, ka-<br />
1128 (U); Sectie 0, 15 Sep 1916 (st), BW2410 (NY),<br />
U), 19 Dec 1919 (2), BW 4494 (NY, U), 25 Feb 1921 moyuwa (Wai-wai), sa-ouro (Maway<strong>an</strong>).<br />
(2), BW5048 (U, UC), 15 Nov 1921 (2), BW 5440 In<br />
(A, Amapa <strong><strong>an</strong>d</strong> Para this subspecies c<strong>an</strong> be con-<br />
RB, U, US), 28 J<strong>an</strong> 1922 (9), BW 5581 (MO, U), 24 fused <strong>with</strong> a form of P. melinonii subsp. meli-<br />
Nov 1922 (2), BW 5992 (F, U); nr. G<strong>an</strong>see, 26 May nonii <strong>with</strong> patent hairs on the lower leaf surface.<br />
1964 (juv), Donselaar 1373 (U); nr. Brownsberg, 14<br />
Oct 1964 (st), Donselaar 1681 (U), 4<br />
However, the latter has<br />
J<strong>an</strong> 1966<br />
sparse, appressed hairs<br />
(juv),<br />
Donselaar 2924 (U); Gr<strong>an</strong> Dam, 13 Oct 1966 (st), Donon<br />
the leafy twigs <strong><strong>an</strong>d</strong> stipules <strong><strong>an</strong>d</strong> the flowers in<br />
selaar 3688 (U); Map<strong>an</strong>e Creek, Feb 1964 (2), Elburg the pistillate inflorescence are diffusely distrib-<br />
(LBB) 9843 (MO, NY, U), 7 Mar 1963 (2), LBB 9424 uted, thus not in two clusters as in subsp. mollis.<br />
(U); Suriname R., <strong>with</strong>out date (8 <strong><strong>an</strong>d</strong> 9), Hostm<strong>an</strong>n Hostm<strong>an</strong>n & Kappler 1272<br />
(& Kappler) 1272<br />
(6) belongs to P.<br />
(B, CGE, G, GH, LE, NY, OXF, P,<br />
S, U, mixed <strong>with</strong> P.<br />
melinonii<br />
melinonii subsp. melinonii); Nas-<br />
subsp. melinonii while Hostm<strong>an</strong>n (&<br />
sau Mts., 16 Feb 1949 (9), L<strong>an</strong>jouw et al. 2106 (NY, Kappler) 1272 (9 <strong><strong>an</strong>d</strong> 6) belongs to P. mollis subsp.<br />
U); Jodensav<strong>an</strong>ne, 27 Mar 1953 (st), Lindem<strong>an</strong> 3631 mollis.<br />
(U), 16 May 1953 (st), Lindem<strong>an</strong> 3989 (U), 20 Dec The description of P. mollis was based on the<br />
1954 (st), Lindem<strong>an</strong> 6923 (U).<br />
collections Bl<strong>an</strong>chet 2361<br />
FRENCH GUIANA. Massif des Emerillons, source<br />
(from Brazil, Bahia);<br />
of Approuage R., 19 Sep 1980 (8), Cremers 6726<br />
Hostm<strong>an</strong>n<br />
(U);<br />
(from Suriname), <strong><strong>an</strong>d</strong> Leprieur 141.<br />
upper Crique Nouciri, 13 Dec 1983 (st), Cremers 8271 The latter was selected as the lectotype collection<br />
(U); <strong>with</strong>out locality, <strong>with</strong>out date (2), Leprieur s.n. (G, (Berg, 1975).<br />
P, lectotype collection of P. mollis); Orapu, 13 Oct<br />
1966 (9), Oldem<strong>an</strong> B.644 (CAY, P, U); piste de St. Elie,<br />
km 16, 19 Apr 1982 (st), Riera 573 (U).<br />
1Ob. <strong>Pourouma</strong> mollis Trecul subsp. triloba<br />
BRAZIL. Without locality, <strong>with</strong>out date (a), Burchell (Trecul) C. C. Berg & v<strong>an</strong> Heusden, Proc. Kon.<br />
9792 (GH, P, the same collection number is also used Ned. Akad. Wetensch., Ser. C, 91(2): 107. 1988.<br />
for a collection of P. gui<strong>an</strong>ensis subsp. gui<strong>an</strong>ensis).<br />
AMAPA: Serra do Navio, 1961 (st), Rodrigues 2984 <strong>Pourouma</strong> triloba Trecul, Ann. Sci. Nat. Bot., Ser. 3,<br />
(INPA); between P6rto Platon <strong><strong>an</strong>d</strong> Serra do Navio, 8:104. 1847 (Aug); Miquel in Martius, Fl. bras. 4(1):<br />
1976 (st), Rosa 1158 (MG). BAHIA: Without locality, 126. 1853; Macbride, Publ. Field Mus. Nat. Hist.,<br />
<strong>with</strong>out date (6), Bl<strong>an</strong>chet 2361 (or 94) (FHO, G, LE, Bot. Ser., 13(2.2): 294. 1937. Type. Peru. Hu<strong>an</strong>uco:<br />
P); P6rto Seguro, 20 Sep 1968 (6), Almeida et al. 71 Macora, <strong>with</strong>out date (6!), Ruiz & Pavon s.n. (lec-<br />
(NY, U); Agua Preta, 23 Feb 1937 (9), Bondar 2169 totype P, chosen here; isolectotypes, G, OXF, US).<br />
(GUA); Rio das Contas, 3 Oct 1919 (8), Curr<strong>an</strong> 21 (C, <strong>Pourouma</strong> jussiae<strong>an</strong>a Trecul, Ann. Sci. Nat. Bot., Ser.<br />
F, NY, US); Ilheus, 1848 (8), Luschnath s.n. (BR), Sep 3, 8: 106. 1847; Miquel in Martius, Fl. bras. 4(1):<br />
1821 (a), Riedel 2 (LE); Castelo Novo, 15 Sep 1944 (2), 124. 1853. Type. Peru. Buena Vista, <strong>with</strong>out date<br />
Velloso 1083 (R). ESPiRITO SANTO: Linhares, 18 Aug (2), Anonymus s.n. (holotype, P).
.?<br />
P. cucura P. cuspidats<br />
GP. 7rorPcabrL P<br />
leo myrsoMaphile<br />
P.<br />
r~~~~~~~~~~~~~~~~~~~<br />
subsp.<br />
tizab<br />
'' ? ~~~~ulSlsp. grlabrat<br />
~~~I 'I ~~~~Ubp. mebli inmii<br />
FIG. 74. Distribution of <strong>Pourouma</strong> cucura, P. cuspidata, P. formicarum, P. melinonii subsp. glabrata, P.<br />
melinonii subsp. melinonii, P. mollis subsp. mollis, P. mollis subsp. triloba, <strong><strong>an</strong>d</strong> P. myrmecophila.
158 Flora Neotropica<br />
~,<br />
a-<br />
e t~~~e7 f~~
<strong>Pourouma</strong>159<br />
<strong>Pourouma</strong> triloba Klotzsch, Linnaea 20: 526. 1847<br />
(Sep). Type. Peru. Hu<strong>an</strong>uco: Macora, <strong>with</strong>out date<br />
(9!), Ruiz & Pavon s.n. (holotype, B; isotypes, BR,<br />
G, OXF).<br />
<strong>Pourouma</strong> jaramilloi Cuatrecasas, Caldasia 7: 300.<br />
1956. Type. Colombia. Meta: Sierra de la Macarena,<br />
Cafno Ciervo, 19 J<strong>an</strong> 1950 (2), Philipson & Jaramillo<br />
2133 (holotype, BM, not seen; isotypes, F, US).<br />
V. 10650 (BG). UYACALLI: Prov. Coronel Portillo, Pucallpa-Tingo<br />
Maria rd., 1980 (9), Trucios 9 (BG).<br />
BRAZIL. ACRE: Upper Rio Moa, nr. Fazenda Arizona,<br />
24-30 Sep 1984 (st), Campbell et al. 8383 (BG);<br />
mouth of Rio Macauh<strong>an</strong>, 5 Aug 1933 (d), Krukoff5309<br />
(F, G, GB, M, MO, NY, S, U, UC, US); Sena Madureira,<br />
Rio Caete, 7 Oct 1968 (d), Pr<strong>an</strong>ce et al. 7907 (F,<br />
GH, NY, P, R, S, U, US).<br />
Lamina usually 3-5-parted, sometimes entire; Vernacular names. Peru. Amazonas: suia, suir<br />
upper surface sometimes scabrous; petiole oc- shuina (Huambisa), uhukamsuiya; S<strong>an</strong> Martin:<br />
casionally <strong>with</strong> white-arachnoid hairs. Pistillate obija, uvilla.<br />
flowers usually more or less diffusely distributed This subspecies is more variable th<strong>an</strong> subsp.<br />
in the inflorescence, sometimes in two ? distinct mollis <strong><strong>an</strong>d</strong> may prove to consist of several infraclusters;<br />
pedicels in fruit up to 1 cm long. specific taxa. In particular Huashikat 944, Hu-<br />
Distribution (Fig. 74). Colombia (Meta), Peru ashikat 1027, Tunqui 296 (all from Peru, Ama-<br />
(Amazonas, Loreto, Madre de Dios, Pasco <strong><strong>an</strong>d</strong> zonas, valley of Rio S<strong>an</strong>tiago) <strong><strong>an</strong>d</strong> Gentry et al.<br />
S<strong>an</strong> Martin), <strong><strong>an</strong>d</strong> Brazil (Acre); in non-inundated 15621 (from Loreto, Rio N<strong>an</strong>ay) are a distinct<br />
forest at low altitudes.<br />
set of collections. They have entire leaves <strong>with</strong><br />
a ? scabrous upper leaf surface, the basal lateral<br />
Specimens examined. COLOMBIA. META: S<strong>an</strong> Ju<strong>an</strong> veins are often<br />
de Arama, Rio Guejar, 26 J<strong>an</strong> 1951 (9), Idrobo et al.<br />
unbr<strong>an</strong>ched, <strong><strong>an</strong>d</strong> the stipules are<br />
1316 (US); Sierra de la Macarena, Caino Ciervo, 19 J<strong>an</strong><br />
rather densely hairy inside.<br />
1950 (9), Philipson & Jaramillo 2133 (BM, P, US, type Pr<strong>an</strong>ce et al. 7907 has relatively m<strong>an</strong>y lateral<br />
collection of P. jaramilloi).<br />
veins, up to ca. 30 pairs, whereas in other col-<br />
ECUDADOR. NAPO: Lumbaqui, 25 Nov 1987 (8), lections there are at most 25 pairs.<br />
Pennington et al. 12300 (BG).<br />
PERU. Without locality, <strong>with</strong>out date<br />
Schunke V. 7729 <strong><strong>an</strong>d</strong> 10650, Foster 5791, D.<br />
(9), Anonymus<br />
s.n. (P, type collection of P. jussiae<strong>an</strong>a). AMAZONAS: N. Smith 5134, <strong><strong>an</strong>d</strong> Trucios 9 have entire leaves<br />
Rio Cenepa, 30 Dec 1972 (9), Berlin 746 (F, GH, MO); <strong><strong>an</strong>d</strong> the pistillate inflorescences tend to the type<br />
Rio S<strong>an</strong>tiago, nr. Caterpiza, 15 Oct 1979 (9), Huashikat of inflorescence normal in subsp. mollis.<br />
944 (U), 24 Oct 1979 (2), Huashikat 1027 (BG); Rio The<br />
Huampi, 26 Jul 1974 (8), Kayap 1306 (MO, US); Rio<br />
description of P. triloba Tr6cul was based<br />
S<strong>an</strong>tiago, nr. Caterpiza, 12 Dec 1979 (9), Tunqui 296 on both staminate <strong><strong>an</strong>d</strong> pistillate material collect-<br />
(BG). HUANUco: Tingo Maria, 13 Jul 1957 (st), Ellen- ed by Ruiz & Pavon, the staminate material is<br />
berg 2297 (U); Macora, <strong>with</strong>out date (6), Ruiz & Pavon chosen as the lectotype collection, the pistillate<br />
s.n. (G, OXF, P, US, lectotype collection of P. triloba material has been used for the description of P.<br />
Tr6cul), <strong>with</strong>out date (2), Ruiz & Pavon s.n. (B, BR, G,<br />
OXF, type collection of P. triloba<br />
triloba Klotzsch.<br />
Klotzsch); Pampayacu,<br />
5 Jun 1927 (d), Sawada 21 (F). JUNiN: Prov. Satipo,<br />
Gr<strong>an</strong> Pajonal, S of Chequitaro, 22 Sep 1983 (8), D. N. 11. <strong>Pourouma</strong> melinonii Benoist, Bull. Mus. Hist.<br />
Smith 5134 (BG). LORETO: Prov. Maynas, Rio N<strong>an</strong>ay, Nat. (Paris) 28: 318. 1922; Berg, Fl. Suriname<br />
Puerto Almendras, 5 J<strong>an</strong> 1976 (9), Gentry et al. 15621<br />
(BG); 22 km S of km 86 on Pucallpa-Tingo Maria 5(1): 274, t. 15a. 1975. Type. French Gui<strong>an</strong>a.<br />
rd.,<br />
11 Feb 1981 (2), Gentry et al. 31179 (BG); Prov. La<br />
May- M<strong>an</strong>a, Sep 1846 (9), Sagot 990 (lectotype,<br />
nas, Y<strong>an</strong>amono Explorama Tourist Camp, 27 Jun 1983 P. chosen by Berg, Fl. Suriname 5(1): 274.1975;<br />
(st), Gentry et al. 42232 (BG), 8 Jul 1983 (st), Gentry isolectotype, B).<br />
et al. 42793 (BG); Yurimaguas, Oct-Nov 1929 (6), LI.<br />
Williams 4688 (F). MADRE DE Dios: Rio M<strong>an</strong>u, Pak- Tree, up to 30 m tall. Leafy twigs 4-12 mm<br />
itza Station, 20 Nov 1980 (9), Foster 5781 (BG, F); thick, sparsely whitish, mostly appressed-puber-<br />
Rio M<strong>an</strong>u, Cocha Cashu Station, Jul 1978 (st), Foster ulous to<br />
et al. 6569 (F). SAN MARTIN: Prov. Mariscal<br />
strigose, <strong><strong>an</strong>d</strong> <strong>with</strong> purple to reddish-<br />
Caceres,<br />
Rio Uchiza, 24 Jul 1974 (a), J. Schunke V. 7729 (MO,<br />
brown, pluricellular hairs <strong><strong>an</strong>d</strong> occasionally sparse,<br />
NY, U); Prov. Mariscal Caceres, Dist. Tocache Nuevo, white, arachnoid hairs. Lamina entire, ovate (to<br />
Quebrada de Ischichini, 12 Oct 1978 (2), J. Schunke elliptic) or 3-lobed, 6-25(-35) x 4-16(-27) cm<br />
FIG. 75. a. <strong>Pourouma</strong> melinonii subsp. melinonii. Leafy twig <strong>with</strong> pistillate inflorescences (Irwin et al. 47832).<br />
b-f. P. mollis subsp. mollis. b. Leafy twig <strong>with</strong> pistillate inflorescences (Elburg (LBB) 9843). c. Staminate flower<br />
(Pires et al. 51625). d. Pistillate flower. e. Ovule f. Seed <strong><strong>an</strong>d</strong> embryo (BW 5581).
160 Flora Neotropica<br />
or 3-5-parted <strong><strong>an</strong>d</strong> ca. 10-27 x 10-27 cm, co-<br />
Key to the Subspecies of<br />
riaceous, apex acuminate, base acute, rounded<br />
<strong>Pourouma</strong> melinonii<br />
to truncate to subcordate or cordate; upper surface<br />
smooth, hairy on the main veins or subgla-<br />
1. Lamina usually entire, if 3-5-parted, then the<br />
base<br />
brous, lower surface usually truncate to subcordate; fruiting peri-<br />
(rather) sparsely appressed- <strong>an</strong>th sparsely or densely hairy; stamens usually<br />
puberulous to strigose or patent-puberulous to <strong>with</strong> free filaments; Amazon Basin <strong><strong>an</strong>d</strong> Gui<strong>an</strong>as.<br />
hirtellous or subglabrous on the veins, arachnoid .......................... a. subsp. m elinonii.<br />
hairs in the areoles or also on the smaller veins; 1. Lamina 3-5-parted, base more or less deeply corlateral<br />
veins 12-20 pairs, basal pair br<strong>an</strong>ched,<br />
date; fruiting peri<strong>an</strong>ths subglabrous; stamens<br />
(often) <strong>with</strong> connate filaments; P<strong>an</strong>ama <strong><strong>an</strong>d</strong><br />
the basal part of these veins forming the basal northern <strong><strong>an</strong>d</strong> western Colombia ..............<br />
part of the leaf margin (thus not separated from ........................... b. subsp. glabrata.<br />
the margin by mesophyll), tertiary venation almost<br />
pl<strong>an</strong>e beneath; petiole 3-22 cm long, strilla.<br />
<strong>Pourouma</strong> melinonii Benoist<br />
gillose to strigose or puberulous to<br />
subsp.<br />
hirtellous; melinonii.<br />
stipules (2-)4-17 cm long, outside sparsely white-<br />
Fig. 75a.<br />
strigose, inside glabrous, caducous. Staminate <strong>Pourouma</strong> apaporiensis Cuatrecasas, Caldasia 7: 297.<br />
inflorescences up to 12 cm long <strong><strong>an</strong>d</strong> 12 cm wide; 1956. Type. Colombia. Amazonas-Vaupes: Rio<br />
peduncle 2-6 cm long, peduncle <strong><strong>an</strong>d</strong> br<strong>an</strong>ches Apaporis, Lagunas del Chucuro, 22 Nov 1951 (9),<br />
Garcia-Barriga 13643 (holotype, US; isotype,<br />
sparsely to rather<br />
NY).<br />
densely puberulous; flowers <strong>Pourouma</strong> apaporiensis Cuatrecasas forma macrosessile<br />
or pedicellate, most of them in globose phylla Cuatrecasas, Caldasia 7:298. 1956. Type. Coheads,<br />
these 3-4 mm diam.; tepals 0.5-1 mm lombia. Vaup6s: Rio Cubiyi, 8 Dec 1943 (9), Allen<br />
long, connate, forming a ? urceolate 3251<br />
peri<strong>an</strong>th, (holotype, US, isotype, MO).<br />
sparsely to densely hairy; filaments longer th<strong>an</strong> Leafy twigs hairy or subglabrous. Lamina<br />
the tepals, free or + connate. Pistillate inflores- mostly entire <strong>with</strong> a rounded to truncate base,<br />
cences in fruit up to 17 cm long <strong><strong>an</strong>d</strong> 11 cm wide; sometimes 3-lobed <strong>with</strong> a truncate to subcordate<br />
peduncle 3-12 cm long, peduncle <strong><strong>an</strong>d</strong> br<strong>an</strong>ches base, occasionally 3-parted <strong>with</strong> a deeply cordate<br />
<strong>with</strong> indument similar to that of the staminate base; lower surface on the veins ? sparsely hairy,<br />
inflorescence; flowers 10-30; pedicel up to 1 cm, mostly <strong>with</strong> appressed hairs, sometimes <strong>with</strong><br />
in fruit up to 1.5 cm long; peri<strong>an</strong>th 0.5-1 cm patent hairs. Stamens <strong>with</strong> (almost) free filalong,<br />
(rather) sparsely puberulous, apex tuber- ments. Fruiting peri<strong>an</strong>th sparsely or + densely<br />
culate; stigma peltate, ca. 2 mm diam. Fruiting puberulous to subvelutinous.<br />
peri<strong>an</strong>th dark violet or dark maroon, ovoid to Distribution (Fig. 74). Colombia (Amazonas),<br />
ellipsoid to oblongoid, 1-2 x 0.7-1.5 cm, sparse- Venezuela (Amazonas <strong><strong>an</strong>d</strong> Bolivar), Guy<strong>an</strong>a,<br />
ly to densely puberulous to subvelutinous or Surinam, French Gui<strong>an</strong>a, Peru (Amazonas <strong><strong>an</strong>d</strong><br />
subglabrous.<br />
Loreto), <strong><strong>an</strong>d</strong> Brazil (Amapa, Amazonas, Mato<br />
This species is very closely related to P. mollis Grosso, <strong><strong>an</strong>d</strong> Para); in non-inundated forest at<br />
from which it is not always easy to separate. One low altitudes.<br />
of the main reasons for treating P. melinonii as<br />
distinct at the species level is the co-occurrence Specimens examined. COLOMBIA. AMAZONAS: Rio<br />
of P. mollis subsp. mollis <strong><strong>an</strong>d</strong> P. melinonii subsp. Apaporis, between Rio Pacoa <strong><strong>an</strong>d</strong> Rio K<strong>an</strong><strong>an</strong>ari, 15-<br />
19 Dec 1951<br />
melinonii in the Gui<strong>an</strong>as <strong>with</strong>out indications of<br />
(9), Garcia-Barriga 14116 (NY, US).<br />
AMAZONAS-VAUPES: Rio Apaporis, Lagunas del Chuinterbreeding.<br />
P. melinonii c<strong>an</strong> be distinguished ruco, 22 Nov 1951 (9), Garcia-Barriga 13643 (NY, US,<br />
from P. mollis by the sparse indument in most type collection of P. apaporiensis). VAUPES: Cabeceras<br />
parts of the pl<strong>an</strong>t, but especially on the stipules; Rio Cubyyfi, 8 Dec 1943 (9), Allen 3251 (MO, type<br />
only the fruiting peri<strong>an</strong>th is sometimes<br />
collection of P.<br />
densely<br />
apaporiensis Forma macrophylla).<br />
VENEZUELA. AMAZONAS:<br />
puberulous to subvelutinous. The sparse indu-<br />
Upper Rio Orinoco,<br />
1951 (9), Croizat 962 (US); base of Cerro Duida, J<strong>an</strong>ment<br />
on the stipules <strong><strong>an</strong>d</strong> leafy twigs of P. meli- Feb 1969 (st), Farinas et al. 490 (U, VEN); Rio Cainonii<br />
is whitish <strong><strong>an</strong>d</strong> usually appressed, whereas quiare, Rio Vaciva <strong><strong>an</strong>d</strong> Rio Yatua, Apr 1853 (6), Spruce<br />
the dense indument on the stipules <strong><strong>an</strong>d</strong> 3176<br />
leafy (C, CGE, E, GH, GOET, MG, NY, OXF, P).<br />
twigs of P. mollis is yellow <strong><strong>an</strong>d</strong><br />
BOLiVAR: Rio Chirea Urim<strong>an</strong>, 5<br />
usually +<br />
Aug 1953<br />
patent.<br />
(st), Bernardi<br />
902 (NY, VEN); Rio Icabaru <strong><strong>an</strong>d</strong> Rio Hacha, 7<br />
Within P. melinonii two allopatric subspecies J<strong>an</strong> 1956 (9), Bernardi 2808 (G, NY); Mun. Urd<strong>an</strong>eta,<br />
c<strong>an</strong> be recognized.<br />
Rio Chinarok, 10 J<strong>an</strong> 1986 (9), Hern<strong><strong>an</strong>d</strong>ez 317 (BG);
<strong>Pourouma</strong> 161<br />
between Rio Chic<strong>an</strong><strong>an</strong> <strong><strong>an</strong>d</strong> Rio Ayaiche, 24 Aug 1961 (Arawak), bospapaja, gr<strong>an</strong>boesipapaja, poeroe-<br />
(6), Steyermark 89452 (NY, VEN); Sierra Ichun, Salto ma or<br />
Maria Espuma, 29 Dec 1961<br />
puruma (Carib),<br />
(6), Steyermark 90408<br />
yarayara (Carib). French<br />
(NY); El Tigre, 2 May 1939 (9), LI. Williams 12008<br />
Gui<strong>an</strong>a. Bois c<strong>an</strong>on, papaye apici (Paramaka).<br />
(F, S, US, VEN).<br />
Peru. Amazonas: dakamtazshuiya, mimpashui-<br />
GUYANA. Pakaraima Mts., Kam<strong>an</strong>a falls, 25 Aug ya washi shuina; Loreto: uvilla. Brazil. Amazo-<br />
1961 (6), Maguire et al. 45945 A (NY, US); upper nas: imbaibar<strong>an</strong>a; Para: mapatir<strong>an</strong>a.<br />
Mazaruni R., Kako R., 23 Sep 1960 (6), Tillett et al. The collection from Amazoni<strong>an</strong><br />
45495 Colombia, in-<br />
(U).<br />
SURINAM. Marowijne R., <strong>with</strong>out date (6), (Host- cluding the type collection of P. apaporiensis,<br />
m<strong>an</strong>n &) Kappler 1272 (LE, M, P); Grote Zwiebel- have deeply cordate leaves, like the material of<br />
zwamp, 22 Sep 1948 (2), L<strong>an</strong>jouw et al. 399 (NY, U); subsp. glabrata, but in contrast to the other col-<br />
Jodensav<strong>an</strong>ne, 4 Jul 1961 (st), Schulz (LBB) 8976 (F, lections of subsp. melinonii. Pistillate specimens<br />
U).<br />
FRENCH GUIANA. Without locality, <strong>with</strong>out data<br />
from this area c<strong>an</strong> be distinguished from those<br />
(2), Aublet s.n. (BM, S), 1986 (6), Melinon 457 (A, G, of subsp. glabrata in the dense indument of the<br />
P, U), 1863 (2), Melinon s.n. (P), 1863 (6), Melinon s.n. peri<strong>an</strong>th of the pistillate flower.<br />
(E, LE, NY, P, US), (8), Poiteau s.n. (LE), 1865 (st) The description of P. melinonii was based on<br />
Sagot s.n. (P); Bolatee Creek, 2 Nov 1948 (st), BAFOG<br />
the collections<br />
4272 (CAY, U);'St. Pierre Creek, 28 Feb 1959 Kappler 1272 (from<br />
(st),<br />
Suriname)<br />
C<strong>an</strong>tonnement de Cayenne s.n. (CAY); Trois Sauts, 28 <strong><strong>an</strong>d</strong> three collections made in French Gui<strong>an</strong>a:<br />
Oct 1974 (2), Lescure 343 (CAY); Cayenne, <strong>with</strong>out Sagot 517 (p.p.), Sagot 990, <strong><strong>an</strong>d</strong> Melinon 457.<br />
date (2), Martin s.n. (BR, F, MO, U); Cacao, S of Cay- Sagot 990 has been selected as the lectotype colenne,<br />
21 Apr 1965 (6), Oldem<strong>an</strong> 1243 (CAY); piste de lection<br />
St. Elie, 1 Jun 1981 (st), Sabatier 51 (Berg, 1975). The (Hostm<strong>an</strong>n &) Kappler<br />
(CAY); Acarou<strong>an</strong>y,<br />
May 1858 (6), Sagot 517 (F, P; under the same number collection has the same number as a collection<br />
the lectotype collection of P. laevis); La M<strong>an</strong>a, Sep 1856 of Hostm<strong>an</strong>n (& Kappler) of P. mollis subsp. mol-<br />
(2), Sagot 990 (B, P, lectotype collection of P. meli- lis.<br />
nonii), 1856 (2), Sagot 990bis (G); Godebert, 13 Jul<br />
1921 (2), Wachenheim 232 <strong><strong>an</strong>d</strong> s.n. (P).<br />
PERU. AMAZONAS: Monte Chichijam, 24 Apr 1973 lib. <strong>Pourouma</strong> melinonii Benoist subsp. glabra-<br />
(2), Ancuash 290 (F, GH, MO); Rio S<strong>an</strong>tiago, nr.. Ca- ta C. C. Berg & v<strong>an</strong> Heusden, Proc. Kon. Ned.<br />
terpiza, 7 Sep 1977 (2), Huashikat 431 (BG); Rio Cusu, Akad. Wetensch., Ser. C, 91(2): 108. 1988.<br />
Yucui, 12 Mar 1973 (2), Kayap 571 (GH). LORETO:<br />
Rio Tacsha, Curaray, 19 Sep 1972 (6), Croat 20447<br />
Type. P<strong>an</strong>ama. C<strong>an</strong>al Zone: Pipeline Rd., 10<br />
(C, km NW of<br />
DUKE, F, GH, MO, NY).<br />
Gamboa, 14 Dec 1973 (9), Berg &<br />
BRAZIL. AMAPA: Rio Jaue, 3 km E of confluence Nee 355 (holotype, U; isotypes, BG, COL, K,<br />
<strong>with</strong> Rio Oiapoque, 26 Aug 1960 (2), Irwin et al. 47832 MO, NY, PMA). Fig. 76.<br />
(GH, NY, U, US); Rio Mutura, 22 Sep 1960 (2), Irwin<br />
et al. 48439 (NY, U, US); Mun. Mazagao, 81 km NW Leafy twigs subglabrous. Lamina usually 5- or<br />
of Macapa, 20 Dec 1984 (2), Mori et al. 17478 (BG). 3-parted <strong>with</strong> a distinctly cordate base, some-<br />
AMAZONAS: Mun. Tefe, Rio Solimoes, Lago de Tefe, times entire <strong><strong>an</strong>d</strong> then <strong>with</strong> a cordate to subacute<br />
15 Oct 1982 (2), Amaral et al. 101 (BG); Rio Japura,<br />
Vila Bitt<strong>an</strong>court, 19 Nov 1982 (2), Amaral et al. 595 base; lower surface on the main veins subgla-<br />
(BG); M<strong>an</strong>aus-Caracarai rd., km 148, 27 Sep 1973 (2), brous. Stamens often <strong>with</strong> + connate filaments.<br />
Berg et al. P. 18136 (F, INPA, MO, NY, P, S, U, US); Fruiting peri<strong>an</strong>th (sub)glabrous.<br />
Mun. Anori, rd. Amori-Anama, 27 Nov 1975 (8), D. Distribution (Fig. 74). P<strong>an</strong>ama <strong><strong>an</strong>d</strong> Colombia<br />
Coelho et al. 672 (INPA); Rio Solimoes, Belem, 16<br />
Dec 1948 (6), Fr6es 23725 (IAN), 25 Mar 1976 (Antioquia); in non-inundated forest at low al-<br />
(st),<br />
Mello s.n. (INPA); M<strong>an</strong>aus-Itacoatiara rd., km 26, 2 titudes.<br />
Sep 1966 (6), Pr<strong>an</strong>ce et al. 2180 (F, GH, INPA, NY,<br />
P, R, S, U, US). MATO GROSSO: Brasilia-Acre Specimens examined. PANAMA. CANAL ZONE:<br />
hwy.,<br />
215 km, west of Rio Juruena, 2 Sep 1963 Pipeline Rd., ca. 10 km NW of Gamboa, 14 Dec 1973<br />
(6), Maguire<br />
et al. 56511 (F, NY, RB, U, US). PARA: Rio (9), Berg & Nee 355 (BG, COL, K, MO, NY, PMA,<br />
Guama,<br />
U,<br />
Sao Miguel, Oct 1906 (2), Goeldi (MG) 7732 type collection of P. melinonii subsp.<br />
(G,<br />
glabrata), 12 Oct<br />
MG);<br />
Rio<br />
1971<br />
Jari, Monte Dourado, 16 Nov 1967 (2), Oliveira (9), Gentry 2046 (MO); Gamboa, 21 Nov 1971<br />
3736 (IAN), 22 J<strong>an</strong> 1968 (2), Oliveira 3936 (NY), 30<br />
(9), Gentry 2657 (MO, NA). DARIEN: S of El Real, nr.<br />
C<strong>an</strong>a<br />
J<strong>an</strong> 1961 (9), Oliveira 4047 (NY), 9 Sep 1968 (2), N.<br />
mine, 21 Aug 1987 (9), McPherson 11510 (BG).<br />
SAN BLAS:<br />
T. Silva 921<br />
Rio<br />
(NY, U), 11 Sep 1968 (6); N. T. Silva 949<br />
C<strong>an</strong>g<strong><strong>an</strong>d</strong>i, nr. C<strong>an</strong>g<strong><strong>an</strong>d</strong>i, 18 Feb 1984<br />
(NY, U), 16 Sep 1968 (2) N. T. Silva 995 (9), Nevers 4929 (BG).<br />
(IAN, NY). COLOMBIA. ANTIOQUIA: 38 km W of Barr<strong>an</strong>cabermeja,<br />
24 Feb 1967 (8), Bruijn 1496 (F, M, MO, NY,<br />
Vernacular names. Venezuela. Bolivar: kai- S, U, US, VEN), 25 Feb 1967 (9), Bruijn 1502 (F, M,<br />
warikai, yagrumo-sunsun. Surinam. Boroma MO, NY, S, U, US, VEN), 1 Mar 1967 (st), Bruijn
162<br />
1546 (F, M, MO, NY, S, U, US, VEN), 2 Feb 1967<br />
(st), Bruijn 1555 (F, M, MO, NY, S, U, US, VEN);<br />
Anori, 6-12 Sep 1973 (9), Soejarto 4283 (MO).<br />
Vernacular names. Colombia. Antioquia: cirpe<br />
macho.<br />
12. <strong>Pourouma</strong> hirsutipetiolata Mildbraed, No-<br />
tizbl. Bot. Gart. Berlin-Dahlem 10: 420. 1928.<br />
Type. Colombia. Bolivar: Norosi-Tiquiso trail,<br />
l<strong><strong>an</strong>d</strong>s of Loba, Apr-May 1916 (Q), Curr<strong>an</strong> 116<br />
(holotype, US, fragment in B).<br />
Tree, up to ca. 30 m tall, <strong>with</strong> butresses. Leafy<br />
twigs 6-20 mm thick, yellow-hirsute <strong><strong>an</strong>d</strong> <strong>with</strong><br />
minute, whitish, appressed or patent hairs <strong><strong>an</strong>d</strong><br />
usually also <strong>with</strong> brown, pluricellular hairs.<br />
Lamina entire, ovate (to elliptic or suborbicular),<br />
10-25 x 7-18 cm, or 3-lobed to -parted <strong><strong>an</strong>d</strong> ca.<br />
10-35 x 10-35 cm, coriaceous, apex acuminate,<br />
base cordate or subcordate or sometimes rounded;<br />
upper surface smooth, hairy on the main veins<br />
or subglabrous, lower surface (rather) sparsely<br />
strigose to strigillose on the veins or puberulous<br />
to hirtellous on the smaller veins, arachnoid hairs<br />
in the areoles, usually extending to the lateral<br />
veins; lateral veins (5-)12-15 pairs, basal pair<br />
br<strong>an</strong>ched, tertiary <strong><strong>an</strong>d</strong> quarterary venation<br />
rather prominent beneath; petiole 3-20 cm long,<br />
yellow-hirsute, usually also <strong>with</strong> minute, whitish<br />
hairs; stipules 4-24 cm long, outside yellow-hirsute<br />
(to -subsericeous), <strong><strong>an</strong>d</strong> also <strong>with</strong> much<br />
shorter, whitish hairs, inside glabrous, caducous.<br />
Staminate inflorescences up to 10 cm long <strong><strong>an</strong>d</strong><br />
8 cm wide; peduncle 3-4 cm long, densely pub-<br />
Flora Neotropica<br />
leafy twigs, <strong><strong>an</strong>d</strong> the more prominent smaller veins<br />
at the lamina beneath, <strong><strong>an</strong>d</strong> the broader segments<br />
of the incised leaves. It differs from P. mollis in<br />
the cordate leaf base of the incised leaves <strong><strong>an</strong>d</strong><br />
the more prominent smaller veins on the lamina<br />
beneath.<br />
Two (probably) allopatric subspecies c<strong>an</strong> be<br />
recognized.<br />
Key to the Subspecies of<br />
<strong>Pourouma</strong> hirsutipetiolata<br />
1. Lamina entire; northern Colombia. ..........<br />
.................... a. subsp. hirsutipetiolata.<br />
1. Lamina 3-lobed to -parted; western Colombia<br />
<strong><strong>an</strong>d</strong> Ecuador. ................ b. subsp. hispida.<br />
12a. <strong>Pourouma</strong> hirsutipetiolata Mildbraed subsp.<br />
hirsutipetiolata. Fig. 77.<br />
Lamina entire.<br />
Distribution (Fig. 81). Colombia (Antioquia,<br />
Bolivar, <strong><strong>an</strong>d</strong> S<strong>an</strong>t<strong><strong>an</strong>d</strong>er); in non-inundated, lowl<strong><strong>an</strong>d</strong><br />
forest.<br />
Specimens examined. COLOMBIA. ANTIOQUIA: 38<br />
km W ofBarr<strong>an</strong>cabermeja, 1 Mar 1967 (9), Bruijn 1547<br />
(F, M, MO, NY, S, U, US, VEN). BOLIVAR: Norosi-<br />
Tiquisio trail, L<strong><strong>an</strong>d</strong>s of Loba, Apr-May 1916 (2), Curr<strong>an</strong><br />
116 (B, US, type collection of P. hirsutipetiolata).<br />
SANTANDER: Bucaram<strong>an</strong>ga, 11 Dec 1977 (2), Renteria<br />
et al. 45 (MO); rd. to Citimarra, 9 Dec 1979 (2), Renteria<br />
et al. 2093 (HUA); 16 km SE of Barr<strong>an</strong>cabermeja,<br />
26 Aug 1954 (9), Romero-Cast<strong>an</strong>eda 4715 (MO, US).<br />
Vernacular names. Colombia. S<strong>an</strong>t<strong><strong>an</strong>d</strong>er: sirpe<br />
erulous or also hirtellous; flowers sessile, most of or sirpo, urumo bl<strong>an</strong>co.<br />
them in subglobose heads, these ca. 3 mm diam.;<br />
tepals 1-1.5 mm long, connate, forming a 12b. <strong>Pourouma</strong> hirsutipetiolata Mildbraed subsp.<br />
(sub)infundibuliform peri<strong>an</strong>th, often <strong>with</strong> <strong>an</strong> hispida (St<strong><strong>an</strong>d</strong>ley & Cuatrecasas) C. C. Berg &<br />
oblique apex, sparsely hairy; filaments longer th<strong>an</strong> v<strong>an</strong> Heusden, Proc. Kon. Ned. Akad. Wethe<br />
tepals, free. Pistillate inflorescences in fruit tensch., Ser. C, 91(2): 108. 1988. Fig. 78.<br />
up to 22 cm long <strong><strong>an</strong>d</strong> 10 cm wide; peduncle 5- <strong>Pourouma</strong> hispida Cuatrecasas, St<strong><strong>an</strong>d</strong>ley & Cuatreca-<br />
13 cm long; peduncle <strong><strong>an</strong>d</strong> br<strong>an</strong>ches minutely sas in Cuatrecasas, Caldasia 7: 299. 1956. Type. Copuberulous,<br />
sparsely so on the peduncle, more lombia. Valle: Rio Anchicaya, La Pl<strong>an</strong>ta, 5 Aug 1943<br />
densely on the br<strong>an</strong>ches <strong><strong>an</strong>d</strong> very densely on the (9), Cuatrecasas 14881 (holotype, F).<br />
pedicels; flowers 7-17; pedicel 0.5-1 cm long, in Lamina 3-lobed to -parted.<br />
fruit up to 1.5 cm long; peri<strong>an</strong>th 5-10 cm long, Distribution (Fig. 81). Colombia (Choc6, Valle)<br />
densely short-velutinous; stigma peltate, ca. 1.5 <strong><strong>an</strong>d</strong> Ecuador (Esmeraldas); in non-inundated,<br />
mm diam., white puberulous. Fruiting peri<strong>an</strong>th lowl<strong><strong>an</strong>d</strong> forest.<br />
dark violet or black, ellipsoid to ovoid, ca. 1.5-<br />
2 x 1-1.5 cm, densely minutely puberulous. Specimens examined. COLOMBIA. CHocO: La Mo-<br />
This species is closely related to P. jarra, 5 Nov 1983 (2), Juncosa 1253 (BG). VALLE: Rio<br />
melinonii,<br />
Anchicaya, La Pl<strong>an</strong>ta, 5 Aug 1943 (2), Cuatrecasas<br />
from which it differs in the shape of the staminate 14881 (F, type collection of P. hispida); Rio Calima,<br />
flowers, the long yellow hairs on the petioles <strong><strong>an</strong>d</strong> La Trojita, 19 Feb-10 Mar 1944 (9), Cuatrecasas 16826
<strong>Pourouma</strong> 163<br />
FIG.<br />
FLORA 76 Prom mn b gbith pitt inre PANAMENSIS Br<br />
b*. . . , .,<br />
A _l I - p , 10 . I w W<br />
rrrO p _ f-nTt.<br />
;<br />
-<br />
^<br />
. r<br />
^I<br />
- ^ , O^ t a r r o l e . .,, ^ V4<br />
]..p<br />
2m 20 - x ? 30 * I <strong>with</strong> L n_5 ;il1t Sotwo MP 5 tul bl z<br />
'<br />
I.-it^^^;.;'' *-i FrtouM mal.nrt^.?, Bf??.ft [,..; C Berg i * ,<br />
i^. ?: .-<br />
'Fu.: C.C. arit o llreah twig t<br />
i.<br />
*'*^ 'AM9 i^ fe tr ^"b-t* cc*B"^*t'a"*l*)^ -<br />
FIG. 76. <strong>Pourouma</strong> melinonii subsp. glabrata. Leafy twig <strong>with</strong> pistillate inflorescences (Berg et al. 355).<br />
(F); Rio Dagua, between Las Cascadas <strong><strong>an</strong>d</strong> Alto de<br />
Yundo, 7 Jul 1984 (9), Gentry et al. 48341 (BG).<br />
ECUADOR. ESMERALDAS: Rio Pambil, Estero Bravo,<br />
6 Jul 1966 (d), Jdtiva 303 (NY), Jdtiva et al. 1079<br />
(NY, UC, US); P<strong>an</strong>adero, 5 km from the river, 26 Jul<br />
1967 (d), Jdtiva et al. 2033 (MO, NY, US), 26 Jul 1967<br />
(9), Jdtiva et al. 2037 (NY, U, UC, US); Rio Hoja<br />
Bl<strong>an</strong>ca <strong><strong>an</strong>d</strong> Rio Hualpi, 15 Sep 1965 (d), Little et al.<br />
21070 (F, MO, NY, US); Rio Guayllabamba, Quinin-<br />
d6, 4 Oct 1965 (d), Little et al. 21225 (NY, US).<br />
Vernacular names. Ecuador. Esmeraldas: uva.
164 Flora Neotropica<br />
INSTIIWJO FORESAL ULATIMO AMERICANO<br />
1I|e1 D . att 1eS s U | I A<br />
h<br />
1g|nc i :..<br />
.. 2 ..:.<br />
-. .<br />
11, 2 _34 5 ta<br />
.<br />
rir, .oula 3s t<br />
. . . .<br />
.... *.., -, . ?<br />
UNITED STATES ;' - ;<br />
Pouriou hirauriprt iolata Mtldbread subap.<br />
*!37'0h3 hr*,rup,,-tolat ... . -.<br />
NToAL Hn UI De. C . . Berg Utrech IatmD1l 1986 m1a<br />
ilo bm<br />
NATiONAL HERSA1IU E.C.K. v<strong>an</strong> Keusden<br />
FIG. 77. <strong>Pourouma</strong> hirsutipetiolata subsp. hirsutipetiolata. Leafy twig <strong>with</strong> pistillate inflorescences (Bruijn<br />
1547).<br />
13. <strong>Pourouma</strong> tomentosa Martius ex Miquel in<br />
Martius, Fl. bras. 4(1): 128, t. 38. 1853. Type.<br />
Colombia. Amazonas or Caqueta: Araracuara,<br />
Dec 1819 (Q), Martius s.n. (lectotype, M, cho-<br />
sen here; isolectotype, U).<br />
Tree, up to ca. 30 m tall. Leafy twigs 4-15 mm<br />
thick, white-puberulous or partly also yellow-<br />
hirsute, often also <strong>with</strong> white, arachnoid hairs,<br />
pluricellular hairs lacking or very sparse. Lamina<br />
entire, ovate to elliptic (to oblong), (4-)10-20<br />
(-40) x (3-)5-15(-27) cm or 3-5-lobed to -parted<br />
or 5-9-parted <strong><strong>an</strong>d</strong> ca. 10-35 x 10-35 cm, coriaceous,<br />
apex acuminate, rounded or sometimes<br />
emarginate, base acute to obtuse, rounded, truncate,<br />
or cordate; upper surface smooth, sometimes<br />
scabrous, hairy on the main veins or only
<strong>Pourouma</strong> 165<br />
m.m,---<br />
* _ . 1 2'. A . .. . . ;<br />
FIG. 78. <strong>Pourouma</strong> hirsutipetiolata subsp. hispida. Leafy twig <strong>with</strong> staminate inflorescences (Little e a.<br />
21225).<br />
on the midrib, in scabrous leaves hispidulous<br />
over the whole surface, lower surface rather<br />
sparsely appressed-puberulous to strigose on the<br />
(main) veins, arachnoid hairs in the areoles, usually<br />
extending to the midrib, but on the main<br />
veins ? disappearing <strong>with</strong> age; lateral veins 7-<br />
25 pairs, the basal pair br<strong>an</strong>ched or sometimes<br />
(in elliptic leaves) unbr<strong>an</strong>ched, the basal part of<br />
these veins forming the basal part of the leaf<br />
margin (thus not separated from the margin by<br />
mesophyll), tertiary (<strong><strong>an</strong>d</strong> quarternary) veins often<br />
rather prominent beneath; petiole 2-26 cm long,<br />
white-puberulous or yellow-(sub)hirsute, <strong><strong>an</strong>d</strong><br />
usually also white, arachnoid hairs; stipules (2-)6-<br />
18 cm long, outside whitish-strigose to yellowhirsute<br />
or -hirtellous, <strong><strong>an</strong>d</strong> <strong>with</strong> dense to sparse,
166 Flora Neotropica<br />
white, arachnoid hairs, usually also pluricellular This species is closely related to P. mollis <strong><strong>an</strong>d</strong><br />
hairs, inside glabrous or <strong>with</strong> dense or sparse, P. melinonii. It is distinguishable by the occuryellow(ish)<br />
hairs, caducous. Staminate inflores- rence of ? dense, arachnoid hairs on the leafy<br />
cences up to 12 cm long <strong><strong>an</strong>d</strong> 12 cm wide; pe- twigs, stipules, petioles, main veins at the lower<br />
duncle 2-5 cm long, peduncle <strong><strong>an</strong>d</strong> br<strong>an</strong>ches + leaf surface <strong><strong>an</strong>d</strong>/or the inflorescences, <strong><strong>an</strong>d</strong>, moredensely<br />
puberulous, often also <strong>with</strong> white, arach- over, by the lack or scarcity of pluricellular hairs<br />
noid hairs; flowers sessile or pedicellate, most of on the leafy twigs.<br />
them in (sub)globose heads, these 3-5 mm diam.; Five subspecies, partly allopatric, c<strong>an</strong> be rectepals<br />
0.5-1 mm long, connate, forming a more ognized. A few specimens from Peru (Amazonas)<br />
or less distinctly urceolate peri<strong>an</strong>th, sparsely to c<strong>an</strong>not be satisfactoraly inserted in one of these<br />
densely hairy; filaments longer th<strong>an</strong> the tepals, subspecies. They may represent <strong>an</strong>other subspefree.<br />
Pistillate inflorescences in fruit up to 20 cm cies. These collections (Kayap 201, Tunqui 217,<br />
long <strong><strong>an</strong>d</strong> 13 cm wide; peduncle 2-13 cm long, Huashikat 966, 1432 <strong><strong>an</strong>d</strong> 1482) largely match<br />
peduncle <strong><strong>an</strong>d</strong> br<strong>an</strong>ched sparsely to densely pub- the collections under subsp. tomentosa, but the<br />
erulous or also yellow-(sub)hirsute, often <strong>with</strong> arachnoid indument is almost lacking on most<br />
white, arachnoid hairs; flowers pedicellate (or parts, except for the main veins of the lamina<br />
subsessile), pedicel up to 0.5 cm long, in fruit up beneath <strong><strong>an</strong>d</strong> the petioles. Moreover, the hairs on<br />
to 1.5 cm long; peri<strong>an</strong>th 5-10 mm long, mostly the fruiting peri<strong>an</strong>th are much shorter, <strong><strong>an</strong>d</strong> the<br />
densely, yellowish, short-velutinous, sometimes apex of the leaves is short-acuminate. They are<br />
sparsely puberulous, or also <strong>with</strong> white, arach- provisionally placed under subsp. tomentosa.<br />
noid hairs; stigma (sub)peltate, ca. 1.5 mm diam. From the two syntype collections made by<br />
Fruiting peri<strong>an</strong>th black or purple, oblongoid to Martius, one is chosen as the lectotype collection.<br />
ellipsoid to ovoid, 1.5-2 x 0.8-1.5 cm, mostly<br />
densely, yellowish-puberulous to -velutinous or<br />
-subhirtellous.<br />
Key to the Subspecies of <strong>Pourouma</strong> tomentosa<br />
1. Stipules hairy inside. .................................. ... ................. b. subsp. apiculata.<br />
1. Stipules glabrous inside.<br />
2. Leafy twigs yellow-hirsute (or glabrous); lamina 5-9-parted. ..................... c. subsp. persecta.<br />
2. Leafy twigs puberulous or hirtellous; lamina entire or 3-5-parted.<br />
3. Base of the lamina more or less deeply cordate; lamina 3-5-parted. ...... d. subsp. essequiboensis.<br />
3. Base of the lamina rounded to truncate or subcordate; lamina usually entire.<br />
4. Lamina 3-5-fid to -parted ............................................ a. subsp. tomentosa.<br />
4. Lamina entire.<br />
5. Apex of the lamina rounded to emarginate, (or short-acuminate in aberr<strong>an</strong>t specimens<br />
from Peru, Amazonas); heads of the staminate inflorescence 4-5 mm diam.; upper Amazon<br />
Basin. .......... ................................. a. subsp. tomentosa.<br />
5. Apex of the lamina acuminate; heads of the staminate inflorescence 2-3 mm diam.;<br />
Gui<strong>an</strong>as, Amapa, <strong><strong>an</strong>d</strong> near M<strong>an</strong>aus. ............................. e. subsp. maroniensis.<br />
13a. <strong>Pourouma</strong> tomentosa Miquel subsp.<br />
tomentosa. Fig. 79.<br />
<strong>Pourouma</strong> albistipulata Cuatrecasas, Bol. Soc. Venez.<br />
Ci. Nat. 17: 92. 1956. Type. Venezuela. Amazonas:<br />
Rio Guainia, Victorino, 22 Mar 1942 (9), LI. Williams<br />
14833 (holotype, F; isotypes, F, G, NY, RB,<br />
S, US, VEN).<br />
Leafy twigs <strong><strong>an</strong>d</strong> petioles white-puberulous.<br />
Lamina entire, elliptic to ovate, sometimes (in<br />
Colombia <strong><strong>an</strong>d</strong> Ecuador) 3- or 5-parted, if entire,<br />
then often thickly coriaceous, + plicate, <strong><strong>an</strong>d</strong> apex<br />
rounded or emarginate (or sometimes short-acuminate),<br />
base rounded to truncate, or (in incised<br />
leaves) to subcordate; upper surface smooth; stipules<br />
outside <strong>with</strong> dense, white-arachnoid hairs,<br />
inside glabrous. Heads of the staminate inflorescences<br />
ca. 4-6 mm in diam. Pistillate flowers +<br />
diffusely distributed in the inflorescence, sometimes<br />
clustered in two groups; peduncle mostly<br />
up to 6(-7) cm long; peri<strong>an</strong>th <strong>with</strong>out dense,<br />
white, arachnoid indument, in fruit yellowish<br />
short-velutinous.
+ AJ ++<br />
P O R O ULMA tomentosa.<br />
FIG. 79. <strong>Pourouma</strong> tomentosa subsp. tomentosa. From Martius, Flora Brasiliensis 4(1). 1853: tab. 38. Leafy twig <strong>with</strong><br />
section of pistillate flower (17), stigma (13), pericarp (19), seed (21), cotyledons (24).
168<br />
Distribution (Fig. 81). The north-western part<br />
of the Upper Amazon Basin; in non-inundated<br />
forest, at low altitudes but in Peru (Amazonas)<br />
up to 2000 m.<br />
Specimens examined. COLOMBIA. AMAZO-<br />
NAS-CAQUETA: Araracuara, Dec 1819 (9), Martius s.n.<br />
(M, U, lectotype collection of P. tomentosa), Feb 1820<br />
(st), Martius s.n. (M, U). VAUPES?: Lagos del Paso, 25<br />
Dec 1975 (6), Roa 252 (INPA).<br />
VENEZUELA. AMAZONAS: Conuco, 2 Dec 1977<br />
(st), Liesner 4105 (U); Cerro Duida, 23 Aug 1944 (st),<br />
Steyermark 57872 (F); Cerro Yapac<strong>an</strong>a, 3 May 1970<br />
(st), Steyermark et al. 103026 (U, Ven); Victorina, Rio<br />
Guainia, 22 Mar 1942 (9), LI. Williams 14833 (G, NY,<br />
RB, US, VEN, type collection of P. albistipulata).<br />
ECUADOR. NAPO: 8 km SE of Tena, 30 Sep 1960<br />
(9), Grubb et al. 1681 (NY); Parque Nacional Yasuni,<br />
9-19 J<strong>an</strong> 1988 (9), Neill et al. 8158 (BG).<br />
Vernacular names. Colombia. Chiricaba (Kar-<br />
ijona), guaumoutognac (Tuk<strong>an</strong>o); Amazonas-<br />
Vaupes: ambauiba do vinho. Venezuela. Ama-<br />
zonas: cocora, cocora mont<strong>an</strong>ero, cucura. Peru.<br />
Amazonas: paiu shuiya (Huambisa), paui shuina;<br />
Loreto: sacha uvilla. Brazil. Amazonas: imbaiba<br />
(or ambauva) do vinho.<br />
Neill et al. 8158 (from Ecuador) <strong><strong>an</strong>d</strong> Rao 252<br />
(from Colombia) differ from the other collections<br />
in the 3- or 5-parted lamina, but match in other<br />
features subsp. tomentosa. Grubb et al. 1681 (from<br />
Ecuador) has 5-parted leaves <strong><strong>an</strong>d</strong> the postillate<br />
flowers clustered as in subsp. apiculata, but the<br />
yellow hairs normally occurring on the leaves of<br />
that subspecies are w<strong>an</strong>ting. Gentry et al. 22865<br />
(from Peru, Amazonas, 1800-1950 m!) is <strong>an</strong>other<br />
collection which could not be placed satisfacto-<br />
rily. It matches subsp. tomentosa, except for the<br />
yellow-hirsute leafy twigs <strong><strong>an</strong>d</strong> stipules.<br />
Flora Neotropica<br />
See also unnamed collections under number 2<br />
(p. 193).<br />
13b. <strong>Pourouma</strong> tomentosa Miquel subsp. api-<br />
culata (Benoist) C. C. Berg & v<strong>an</strong> Heusden,<br />
Proc. Kon. Ned. Akad. Wetensch., Ser. C,<br />
91(2): 108. 1988.<br />
<strong>Pourouma</strong> apiculata Spruce ex Benoist, Bull. Mus. Hist.<br />
Nat. (Paris) 28: 321. 1922. Type. Brazil. Amazonas:<br />
Rio Uaupes, nr. P<strong>an</strong>ur6 (=Ip<strong>an</strong>or6), Oct 1852-J<strong>an</strong><br />
1853 (Q), Spruce 2865 (holotype, P; isotypes, B, BR,<br />
C, CGE, E, F, G, GH, GOET, LE, MG, NY, OXF,<br />
P, US).<br />
<strong>Pourouma</strong> apiculata Spruce ex Mildbraed, Notzbl. Bot.<br />
Gart. Berlin-Dahlem 10: 419. 1928. Based on the<br />
specimen of Spruce 2865 in B.<br />
<strong>Coussapoa</strong> krukovii St<strong><strong>an</strong>d</strong>ley, Publ. Field Mus. Nat.<br />
PERU. AMAZONAS: 12-18 km on trail E of La Peca Hist., Bot. Ser., 17: 163. 1937. Type. Brazil. Amain<br />
Serr<strong>an</strong>ia de Bagua, 1800-1950 m, 14 Jun 1978 (8), zonas: Mun. Sao Paulo de Olivenca, 11 Sep-26 Oct<br />
Gentry et al. 22865 (BG); Rio S<strong>an</strong>tiago, Caterpiza, 17 1936 (a), Krukoff 8223 (holotype, NY; isotypes, A,<br />
Oct 1979 (9), Huashikat 966 (BG), 4 Dec 1979 (9), F, G, GH, LE, MO, P, S, U, US).<br />
Huashikat 1432 (U), 10 Dec 1979 (9), Huashikat 1482<br />
(BG); Rio Cenepa, Quebrack Yutai-entsa, 22 J<strong>an</strong> 1973 Leafy twigs yellow-hirsute (sometimes only on<br />
(9), Kayap 201 (F, GH, NO); Rio S<strong>an</strong>tiago, Quebrada the scars of the stipules), this indument often<br />
Caterpiza, 3 Dec 1979 (9), Tunqui 217 (BG). LORETO: extended to the petioles. Lamina usually entire<br />
Prov. Requena, Arboretum Jenaro Herrera, Aug-Sep <strong><strong>an</strong>d</strong> ovate, sometimes 3-lobed or (in<br />
1976<br />
subjuvenile<br />
(6), Bernardi s.n. (arbre 6/27) (U); Prov. Maynas,<br />
Dist. Iquitos, Zungaro Cocha, 13 Dec 1967 (9), Reyna material?) 3-5-parted, apex usually acuminate,<br />
R. 37 (BG, F).<br />
base broadly acute to truncate or (in subjuvenile<br />
BRAZIL. AMAZONAS: Tefe, 25 Sep 1940 (st), Black material?) occasionally shallowly cordate; upper<br />
47-1520 (IAN); M<strong>an</strong>aus-Itacoatiara road, km 26 (st), surface smooth or scabrous; stipules outside <strong>with</strong><br />
L. Coelho (INPA) 5223 (INPA); 5 Jun 1976 (st), Haroldo<br />
(INPA) 57512 (INPA), 26 Oct 1965 (6), Loureiro (rather) dense, white arachnoid hairs, inside <strong>with</strong><br />
(INPA) 16461 (INPA), 19 Mar 1976 (st), Mello (INPA) yellow hairs. Heads of the staminate inflores-<br />
55388 (INPA); M<strong>an</strong>aus, Dec 1850-Mar 1851 (9), Spruce cences 3-4 mm diam. Peri<strong>an</strong>th of the staminate<br />
951 (M), Spruce 1219 (P), Spruce s.n. (B, CGE, E, F, flower densely puberulous. Peduncle of the pis-<br />
G, GOET, LE, NY, OXF, P).<br />
tillate inflorescences mostly up to 8 cm long. Pistillate<br />
flowers usually in two ? distinct clusters;<br />
peri<strong>an</strong>th yellow-velutinous, <strong>with</strong>out dense, white,<br />
arachnoid hairs.<br />
Distribution (Fig. 81).Venezuela (Bolivar), Peru<br />
(Loreto), <strong><strong>an</strong>d</strong> Brazil (Amazonas, Mato Grosso,<br />
<strong><strong>an</strong>d</strong> Roraima); in non-inundated forest.<br />
Specimens examined. VENEZUELA, AMAZONAS:<br />
Depto. Rio Negro, nr. Cerro de La Neblina Base Camp,<br />
22 Feb 1984 (9), Liesner 16175 (BG), 4 May 1984 (2),<br />
Thomas 3349 (BG). BOLiVAR: Rio Hacha, Rio Icabaru,<br />
2 J<strong>an</strong> 1956 (8), Bernardi 2731 (F, NY), Rio Carfin, Apr<br />
1945 (a), Cardona 1222 (US, VEN).<br />
PERU. LORETO: Prov. Maynas, Rio N<strong>an</strong>ay, Mish<strong>an</strong>a,<br />
6 J<strong>an</strong> 1983 (st), Gentry et al. 39264 (BG).<br />
BRAZIL. AMAZONAS: Sao Paulo de Olivenca, 7 Oct<br />
1931 (8), Ducke s.n. (RB); Humaita, 12 Oct-6 Nov<br />
1934 (9), Krukoff 6887 (A, BR, F, G, LE, MO, NY,<br />
RB, S, U, US); Sao Paulo de Olivenca, 11 Sep-26 Oct<br />
1936 (8), Krukoff8223 (A, F, G, GH, LE, MO, NY,<br />
P, S, U, US, type collection of <strong>Coussapoa</strong> krukovii);<br />
Democracia, 31 Aug 1923 (2), Kuhlm<strong>an</strong>n 255 (BG);
<strong>Pourouma</strong> 169<br />
Varadouro do Morcego, 31 Aug 1923 (8), Kuhlm<strong>an</strong>n PERU. LORETO: Prov. Alto Amazonas, Rio Pastaza,<br />
291 (RB); M<strong>an</strong>aus-Itacoatiara rd., km 57, 15 Sep 1976 Andoar, 21 Nov 1980 (2), Vdsquez et al. 809 (BG).<br />
(a), Mota 646 (INPA); km 159, 20 Sep 1974 (8), Pr<strong>an</strong>ce PAsco: Prov. Oxapampa, Distr. Iscozacin, 22 Sep 1986<br />
et al. 22708 (INPA, MG, U), 291 (RB); Tef6, 28 Nov (8), Pariona et al. 954 (BG).<br />
1959 (st), Rodrigues et al. 1434 (INPA); Rio Solim6es, BRAZIL. AMAZONAS: Sao Paulo de Olivenca, 11<br />
Fonte Boa, 26 Oct 1968 (6), M. Silva 2199 (G, MG); Sep-26 Oct 1936 (9), Krukoff 8325 (A, BR, F, G, LE,<br />
Ip<strong>an</strong>or6 ("P<strong>an</strong>ur6"), Oct 1852-J<strong>an</strong> 1853 (9), Spruce NY, PS, U).<br />
2865 (B, BR, C, CGE, E, F, G, GH, GOET, LE, MG, BOLIVIA. COCHABAMBA: Torora, Jungas of J<strong>an</strong>a<br />
NY, OXF, P, US, type collection ofP. apiculata). MATO Maya, 1800 m, 24 Jun 1947 (2), Cardenas 3973 (US)<br />
GRosso: Rio Aripu<strong>an</strong>a, Nucleo Pioneiro Humboldt, LA PAZ: Larecaja, Mapiri, 12-30 Sep 1939 (8), Krukoff<br />
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<br />
(F, INPA, MO, NY, P, S, U, US); Aripu<strong>an</strong>a, 18 Feb- (9), Krukoff 11062 (A, F, G, MO, NY, S, U, UC, US,<br />
9 May 1977 (st), Gomes et al. 826, 1517, 1540, 1588, type collection of P. tomentosa subsp. persecta).<br />
1598, 1604, 1610, 1679 (INPA), 8 Nov 1978 (2), Ryl<strong><strong>an</strong>d</strong>s<br />
30 (INPA); Mun. Vila Bela da S<strong>an</strong>tissima Trin- Vernacular names. Brazil. Amazonas: mapati<br />
idad, 5 km S of border of Rond6nia, 2 Nov 1985 (2), or<br />
Thomas et al. 4765 (BG). RONDONIA: Rd. to Abuna to<br />
mapaty; Mato Grosso: imbafibar<strong>an</strong>a. Bolivia.<br />
Guajara-Mirim, 1 km N of Riberao, 25 Jul 1968 Cochabamba: uva de monte.<br />
(a),<br />
Pr<strong>an</strong>ce et al. 6292 (G, INPA, NY, S, U, US). RORAIMA: Cardenas 3973 (from Bolivia) differs from the<br />
Anaris, 12 Feb 1969 (2), Pr<strong>an</strong>ce et al. 9857 (F, GH, others in having the pistillate flowers in two clus-<br />
INPA, M, NY, P, K, S, U, US), Serra dos Surucucus, ters in the inflorescences.<br />
15 Feb 1969 (2), Pr<strong>an</strong>ce 9952 (C, INPA, MO, MY, R,<br />
S, U).<br />
Vdsquez et al. 809 (from Peru) differs from the<br />
other collections in the absence of long yellow<br />
Vernacular names. Venezuela. Bolivar: sara- hairs on the leafy twigs. The same applies to<br />
sara (Arekuna). Brazil. Amazonas: imbauibar<strong>an</strong>a, collection Korning et al. 47618 which has, morepurumai.<br />
over, distinctly petiolulate leaf segments.<br />
13c. <strong>Pourouma</strong> tomentosa Miquel subsp. per-<br />
secta C. C. Berg & v<strong>an</strong> Heusden, Proc. Kon.<br />
Ned. Akad. Wetensch., Ser. C, 91(2): 108.1988.<br />
Type. Bolivia. La Paz: Prov. Larecaja, Copa-<br />
cab<strong>an</strong>a, 10 km S ofMapiri, 8 Oct-15 Nov 1939<br />
(2), Krukoff 11062 (holotype, U; isotypes, A,<br />
F, G, MO, NY, S, UC, US). Fig. 80.<br />
13d. <strong>Pourouma</strong> tomentosa Miquel subsp. esse-<br />
quiboensis (St<strong><strong>an</strong>d</strong>ley) C. C. Berg & v<strong>an</strong> Heus-<br />
den, Proc. Kon. Ned. Akad. Wetensch., Ser.<br />
C 91(2): 109. 1988.<br />
<strong>Pourouma</strong> essequiboensis St<strong><strong>an</strong>d</strong>ley, Lloydia 2: 175.<br />
1939; Berg, Fl. Suriname 5(1): 276. 1975. Type.<br />
Guy<strong>an</strong>a. Essequibo R., nr. mouth of Onoro Creek,<br />
Leafy twigs <strong><strong>an</strong>d</strong> petioles yellow-hirsute, some- 15-24 Dec 1937 (d), A. C. Smith 2731 (holotype, F;<br />
times glabrous. Lamina 5-9-parted, <strong>with</strong> inci- isotypes, A, G, Mo, NY, P, S, U, US).<br />
sions down to (near) the petiole, sometimes <strong>with</strong><br />
the segments petiolulate, occasionally entire, apex Leafy twigs <strong><strong>an</strong>d</strong> petioles <strong>with</strong>out long yellow<br />
acuminate, base cordate to subcordate; upper hairs. Lamina 3-5-parted, apex acuminate, base<br />
surface smooth; stipules outside <strong>with</strong> sparse, ? deeply cordate; upper surface smooth; stipules<br />
white, arachnoid hairs, inside <strong>with</strong> sparse yellow <strong>with</strong> rather dense, white, arachnoid hairs, inside<br />
hairs. Heads of the staminate inflorescences ca.<br />
glabrous. Heads of the staminate inflorescences<br />
5 mm in diam. Peduncle of the pistillate inflo- ca. 3 mm diam. Peduncle of the pistillate inflorescences<br />
often longer th<strong>an</strong> 8 cm. Pistillateflow- rescences mostly up to 6(-7) cm long. Pistillate<br />
ers + diffusely distributed in the inflorescence or<br />
flowers ? diffusely distributed in the infloressometimes<br />
clustered in two groups; peri<strong>an</strong>th of cence. Peri<strong>an</strong>th of the pistillate flower yellowthe<br />
pistillate flower yellow-velutinous, <strong>with</strong>out<br />
velutinous, <strong>with</strong>out dense, white, arachnoid hairs.<br />
dense, white, arachnoid hairs.<br />
Distribution (Fig. 81). Guy<strong>an</strong>a <strong><strong>an</strong>d</strong> Brazil<br />
Distribution (Fig. 81). Brazil (Amazonas <strong><strong>an</strong>d</strong><br />
(Amazonas <strong><strong>an</strong>d</strong> Para); in non-inundated forest.<br />
Mato Grosso), Peru (Loreto), <strong><strong>an</strong>d</strong> Bolivia (La Paz<br />
<strong><strong>an</strong>d</strong> Cochabamba); in non-inundated forest, at Specimens examined. GUYANA. Mouth of Onoro<br />
altitudes up to 1800 m.<br />
Creek, 15-24 Dec 1937 (6), A. C. Smith 2731 (A, F,<br />
G, GH, MO, NY, P, S, U, US, type collection of P.<br />
Specimens examined. ECUADOR. NAPO: Aii<strong>an</strong>gu, essequiboensis).<br />
30 May-21 Jun 1982 (st), SEF 8823 (U), 1-15 Feb BRAZIL. AMAZONAS: Itapir<strong>an</strong>ga, Rio Uatuma, 27<br />
1986 (2), Korning et al. 47618 (BG).<br />
Aug 1979 (C), Cid et al. 847 (BG, INPA); Rio Jurua,
170 Flora Neotropica<br />
_ I<br />
Revised<br />
SAt L arsed forkFloa e d.rrovc e. e<br />
FIG. 80. <strong>Pourouma</strong> tomentosa subsp. persecta. Leafy twig <strong>with</strong> pistilate inflorescences (Krukoff 8325).<br />
Saracura, 22 Aug 1975 (9), D. Coelho et al. (INPA)<br />
53383 (INPA, U); Codajas, Rio Capitari, 3 Sep 1950<br />
(d), Fr6es 26520 (US). PARA: Cuiaba-S<strong>an</strong>tarem rd., km<br />
1180, 17 Nov 1977 (2), A. S. Silva et al. 215 (BG, COL,<br />
K, MO, U).<br />
w<br />
L U - a SL I-. ITtA<br />
ora fm bro picn a T.<br />
13e. <strong>Pourouma</strong> tomentosa Miquel subsp. ma-<br />
roniensis (Benoist) C. C. Berg & v<strong>an</strong> Heusden,<br />
Proc. Kon. Ned. Akad. Wetensch., Ser. C 91(2):<br />
109. 1988. Fig. 82a.<br />
FIG. 81. Distribution of <strong>Pourouma</strong> herrerensis, P. hirsutipetiolata subsp. hirsutipetiolata, P. hirsutipetiolata<br />
subsp. hispida, P. napoensis, P. tomentosa subsp. apiculata, P. tomentosa subsp. essequiboensis, P. tomentosa<br />
subsp. maroniensis, P. tomentosa subsp. persecta, <strong><strong>an</strong>d</strong> P. tomentosa subsp. tomentosa.
<strong>Pourouma</strong><br />
qlq,<br />
_ j PIF P. t os<strong>an</strong>tosa stubsp. t lontosa<br />
__ / *~ _~ *<br />
C~ Osa iF~ P <strong>with</strong> entire lamina<br />
ab^P~ 0?^P "L^ I-~~S<br />
^fr/I Z 1^ ^<br />
P..hir?utiptiolat.<br />
1A<br />
^-^<br />
^ 0<strong>with</strong> incised<br />
Smrine<br />
,wrr@e ,3<br />
s<br />
^Sfti^"1~^PA ?- 'AP. naponsis<br />
i-r g^ -r ----- ---p.-' b-------AS"s"eioovsZ<br />
'l^^ ? C<br />
^P0<br />
. sqa ounu<br />
sulbOp. mflroni.nsis<br />
.h a,~_<br />
,ns is<br />
171
172 Flora Neotropica<br />
FIG. 82. a. <strong>Pourouma</strong> tomentosa subsp. maroniensis. Leaf <strong><strong>an</strong>d</strong> pistillate inflorescences (L<strong>an</strong>jouw et al. 432).<br />
b. <strong>Pourouma</strong> minor. Leafy twig <strong>with</strong> pistillate inflorescences (Pr<strong>an</strong>ce et al. 10089).<br />
<strong>Pourouma</strong> maroniensis Benoist. Bull. Mus. Hist. Nat.<br />
(Paris) 28: 319. 1922; Berg, Fl. Suriname 5(1): 276,<br />
t. 14a. 1975. Type. French Gui<strong>an</strong>a. Godebert, 23<br />
Jun 1921 (d), Wachenheim ser. 2: 392 (holotype, P).<br />
Leafy twigs <strong><strong>an</strong>d</strong> stipules <strong>with</strong>out long yellow<br />
hairs. Lamina entire <strong><strong>an</strong>d</strong> ovate, only in subju-<br />
venile material to 3-parted, but then the base to<br />
truncate or at most subcordate, apex acuminate;<br />
upper surface smooth; stipules outside <strong>with</strong> dense,<br />
white, arachnoid hairs, inside glabrous. Heads of<br />
the staminate inflorescences normally 2-3 mm<br />
diam. Peri<strong>an</strong>th of the staminate flower sparsely<br />
puberulous. Peduncle of the pistillate inflores-<br />
cences mostly up to 7 cm long. Pistillate flowers<br />
+ diffusely distributed in the inflorescence. Peri-<br />
<strong>an</strong>th of the pistillate flower sparsely white-pub-<br />
erulous or yellow-velutinous, often <strong>with</strong> dense,<br />
white, arachnoid hairs.<br />
Distribution (Fig. 81). Eastern Surinam, French<br />
Gui<strong>an</strong>a, <strong><strong>an</strong>d</strong> Brazil (Amapa <strong><strong>an</strong>d</strong> Para); in non-<br />
inundated forest.<br />
Specimens examined. SURINAM. Moengo, 25 Aug<br />
1951 (st), BBS 298 (U); G<strong>an</strong>see, 21 Apr 1964 (st),<br />
Donselaar 1226 (U); Grote Zwiebelzwamp, 23 Sep 1948<br />
(9), L<strong>an</strong>jouw et al. 432 (F, NY, U); Lely Mts., 2 Oct<br />
1975 (st), Lindem<strong>an</strong> et al. 701 (U).<br />
FRENCH GUIANA. St. Laurent, Feb 1956 (9),<br />
BAFOG 7228 (NY, P, U); M<strong>an</strong>a, 3 Mar 1956 (9), BAF-<br />
OG 7330 (NY, P); Charvein, 29 Jun 1914 (d), Benoist
<strong>Pourouma</strong> 173<br />
465 (P); Trois Sauts, Zidockville, 28 J<strong>an</strong> 1975 (2), Gren- type collection of P. stipulacea); 1-6 km W of Ka<strong><strong>an</strong>d</strong><br />
688 (CAY); M<strong>an</strong>a R., Saut Sabbat, 16 Jul 1981 mar<strong>an</strong>g, Mazaruni R., 21 Aug 1977 (st), Maas et al.<br />
(2), Gr<strong>an</strong>ville 4531 (BG); Godebert, 23 Jun 1921 (6), 2592 (U).<br />
Wachenheim 392 (P, type collection of P. maroniensis).<br />
This species shares (sub)persistent stipules <strong>with</strong><br />
BRAZIL. AMAPA: Serra do Navio, 4 J<strong>an</strong> 1985 (6), P. oraria <strong><strong>an</strong>d</strong> P. bicolor subsp. choco<strong>an</strong>a. Most<br />
Mori et al. 17672 (BG), 25 Sep 1961 (6), Pires et al.<br />
51193 (B, FHO, INPA, NY, P, U, US); Rio<br />
features, e.g., the white, arachnoid indument on<br />
Araguari,<br />
9 Oct 1961 (6), Pires et al. 51625 (FHO, NY, U, US);<br />
most parts, suggest relationship to P. tomentosa.<br />
Serra do Navio, 1961 (st), Rodrigues 2986 <strong><strong>an</strong>d</strong> 2987 According to Boom (et al. 8102) the exudate<br />
(INPA). PARA: Rio Jari, Monte Dourado, 22 J<strong>an</strong> 1968 is red <strong><strong>an</strong>d</strong> viscous.<br />
(2), Oliveira 3936 (NY).<br />
Vernacular names. Surinam. Boroma (Ara-<br />
wak), boesipapja, gr<strong>an</strong>boesipapaja. French<br />
Gui<strong>an</strong>a. Bois c<strong>an</strong>on, bouchi papaye (Paramaka),<br />
kukuma, wilaupiyua (Wayapi).<br />
The morphological differences between subsp.<br />
maroniensis, subsp. apiculata <strong><strong>an</strong>d</strong> subsp. tomen-<br />
tosa are not strong.<br />
14. <strong>Pourouma</strong> stipulacea C. C. Berg, Brittonia<br />
34(1): 37, t. 2. 1982. Type. Guy<strong>an</strong>a. Bartica-<br />
Potaro rd., 25 J<strong>an</strong> 1943 (2), F<strong>an</strong>shawe 1105 =<br />
FD 3851 (holotype, K). Fig. 83.<br />
Tree, up to 6 m tall. Leafy twigs 1-2.5 cm thick,<br />
yellow-hirsute. Lamina 5-parted, 25-50 x 35-<br />
75 cm, coriaceous, apex acuminate, base deeply<br />
cordate, often <strong>with</strong> lobes overlapping; upper surface<br />
smooth, hirtellous on the main veins, lower<br />
surface white-puberulous <strong><strong>an</strong>d</strong> <strong>with</strong> sparse, or<strong>an</strong>ge,<br />
pluricellular hairs on the veins, arachnoid<br />
hairs ? confined to the areoles; lateral veins 20-<br />
30 pairs, tertiary (<strong><strong>an</strong>d</strong> quartemary) veins prominent<br />
beneath; petiole 23-65 cm long, sparsely<br />
yellow-hirsute; stipules 15-20 cm long, outside<br />
yellow-hirsute <strong><strong>an</strong>d</strong> <strong>with</strong> sparse to dense, white,<br />
arachnoid hairs, <strong><strong>an</strong>d</strong> <strong>with</strong> sparse, minute, or<strong>an</strong>ge<br />
pluricellular hairs, inside sparsely yellow-hirsute,<br />
subpersistent. Staminate inflorescences unknown.<br />
Pistillate inflorescences in fruit up to 15<br />
cm long <strong><strong>an</strong>d</strong> 8 cm wide; peduncles 10 cm long,<br />
peduncle <strong><strong>an</strong>d</strong> br<strong>an</strong>ches sparsely yellow-hirsute<br />
15. <strong>Pourouma</strong> acuminata Martius ex Miquel in<br />
Martius, Fl. bras 4(1): 130. t. 40. 1853; Mar-<br />
tius, Syst. mat. med. bras. 34. 1843, nomen.<br />
Type. Brazil. Amazonas: Rio Japura, between<br />
Maripi <strong><strong>an</strong>d</strong> Par<strong>an</strong>a-Mirim, Dec 1819 (9), Mar-<br />
tius s.n. (holotype, M; isotype, U). Fig. 84.<br />
<strong>Pourouma</strong> populifolia St<strong><strong>an</strong>d</strong>ley, Publ. Field Mus. Nat.<br />
Hist., Bot. Ser., 17: 184. 1937. Type. Brazil. Amazonas:<br />
Mun. Sao Paulo de Olivenca, nr. Palmares,<br />
1 Sep-26 Oct 1936 (9), Krukoff8427 (holotype, NY;<br />
isotypes, A, BR, F, G, LE, P, S, U, US).<br />
Tree, up to 26 m tall. Leafy twigs 4-7 mm<br />
thick, (sparsely) yellow-hirsute. Lamina entire,<br />
(broadly) ovate to deltoid or to elliptic, 5-19 x<br />
3-10 cm, coriaceous, apex (long) acuminate, base<br />
rounded to acute; upper surface smooth, main<br />
veins sparsely hairy, lower surface white-, ap-<br />
pressed -puberulous on the main veins, minutely<br />
puberulous on the smaller veins, arachnoid hairs<br />
in the areoles, sometimes extending to the main<br />
veins; lateral veins 7-14 pairs, basal pair un-<br />
br<strong>an</strong>ched, tertiary venation slightly prominent<br />
beneath; petiole sparsely appressed-puberulous<br />
to glabrescent, <strong><strong>an</strong>d</strong> usually sparsely hirsute; stip-<br />
ules 5-10 cm long, outside yellow-hirsute <strong><strong>an</strong>d</strong><br />
<strong>with</strong> dense, white, arachnoid hairs, occasionally<br />
<strong>with</strong> minute, appressed, white hairs, inside gla-<br />
brous, caducous. Staminate inflorescences unknown.<br />
Pistillate inflorescences in fruit up to 10<br />
cm long <strong><strong>an</strong>d</strong> 4 cm wide; peduncle 4.5-8 cm long,<br />
peduncle <strong><strong>an</strong>d</strong> br<strong>an</strong>ches puberulous (velutinous<br />
<strong><strong>an</strong>d</strong> <strong>with</strong> (dense), white, arachnoid hairs; flowers on the ultimate br<strong>an</strong>ches);flowers 4-17; pedicels<br />
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,<br />
2 mm diam. Fruitingperi<strong>an</strong>th ovoid, ca. <strong>an</strong>th 2-5 mm<br />
2 x<br />
long, densely yellow-velutinous,<br />
1 cm, yellow-hirsute <strong><strong>an</strong>d</strong> <strong>with</strong> dense, white- apex papillose; stigma peltate, 1.5-2 mm diam.,<br />
arachnoid hairs.<br />
sometimes puberulous. Fruiting peri<strong>an</strong>th ovoid<br />
Distribution (Fig. 85). Guy<strong>an</strong>a; in non-inun- to elliptic, ca. 1-1.6 x 1 cm, densely yellowdated<br />
forest at low altitudes.<br />
velutinous.<br />
Specimens examined. GUYANA. Nr. Kamer<strong>an</strong>g, 20 Distribution (Fig. 85). Brazil (Amazonas) <strong><strong>an</strong>d</strong><br />
Jun 1987 (st), Boom et al. 8102 (BG); Bartica-Potaro Peru (Loreto); in periodically inundated (varzea)<br />
rd., 25 J<strong>an</strong> 1943 (2), F<strong>an</strong>shawe 1105 (=FD 3851) (K, forest or also in non-inundated forest.
174 Flora Neotropica<br />
- -<br />
g<br />
FIG. 83. <strong>Pourouma</strong> stipulacea. Leaf <strong><strong>an</strong>d</strong> pistillate inflorescence (F<strong>an</strong>shawe 1105); twig <strong>with</strong> stipules (Maas<br />
et al 2592).<br />
FIG. 84. <strong>Pourouma</strong> acuminata. From Martius, Flora Brasiliensis 4(1). 1853: tab. 40. Leafy twig <strong>with</strong> pistillate<br />
inflorescences, fruiting peri<strong>an</strong>th <strong><strong>an</strong>d</strong> fruit (17), fruit (19), fruiting peri<strong>an</strong>th (4311), pericarp <strong><strong>an</strong>d</strong> seed (19, 21),<br />
seed (21), embryo (23), radicle (25).
<strong>Pourouma</strong> 175<br />
I0711~~~~~~~~~~~~~~~~~ 41 al<br />
9- . . .. ,~ act ninat<br />
..4<br />
I'OU'R M A L aeminnata.
+~~~5 P. acuminata .--ID P.<br />
fei\`f `~ p ~rugines<br />
4~074' -~-3~<br />
-P.
<strong>Pourouma</strong> 177<br />
Specimens examined. VENEZUELA. BOLvAR: Mun. white-puberulous, densely so on the ultimate<br />
For<strong>an</strong>eo Aripao, Carno Fatua, 2-5 May 1988 (st), Ay- br<strong>an</strong>ches; flowers sessile, not densely clustered;<br />
mard et al. 6778 (BG).<br />
PERU. LORETO: Rio Itaya, 5 km above Iquitos, 8 tepals ca. 1.5 mm long, almost free, densely pu-<br />
Aug 1972 (9), Croat 18888 (C, DUKE, F, GH, NA, berulous; filaments shorter th<strong>an</strong> the tepals. Sta-<br />
NY, P); Rio Itaya, Nuevo Esper<strong>an</strong>za, 10 Oct 1975 (9), minate inflorescences unknown. Pistillate inflo-<br />
McD<strong>an</strong>iel et al. 20362 (NA); Rio Momon, 12 Oct 1977 rescences up to 70 cm long <strong><strong>an</strong>d</strong> 16 cm wide;<br />
(2), Rimachi 3256 (MO).<br />
peduncle 32-48 cm<br />
BRAZIL. AMAZONAS: Rio Japura, Vila<br />
long, peduncle <strong><strong>an</strong>d</strong> br<strong>an</strong>ches<br />
Bitt<strong>an</strong>court,<br />
11 Nov 1982 (9),Amaral etal. 473(BG); Rio ICa, 1877<strong>with</strong><br />
indument similar to that of the staminate<br />
1878 (2), Jobert 682 (P); Mun. Sao Paulo de Olivena, inflorescences;flowers 7-13; pedicel 0.7-1.3 cm<br />
nr. Palmares, 11 Sep-26 Oct 1936 (9), Krukoff 8427 long, in fruit up to 5 cm; peri<strong>an</strong>th ca. 1 cm long,<br />
(A, BR, F, G, LE, MO, NY, P, S, U, US, type collection densely white-puberulous;<br />
of P.<br />
stigma peltate, ca. 3<br />
populifolia); Rio Japura, Par<strong>an</strong>a-Mirim, Dec 1819 mm<br />
(2), Martius s.n. (M, U, type collection of P. acumi- diam.; bilobate, densely puberulous. Fruitnata);<br />
Rio Ia, Oct 1877 (9), (herb.) Schwacke 560 (R, ing peri<strong>an</strong>th globose, ca. 2 cm diam., apiculate,<br />
RB).<br />
yellow-short-velutinous.<br />
Distribution<br />
Vernacular names. Peru. Loreto: sacha<br />
(Fig. 85). Brazil (Amazonas), in<br />
uvilla, non-inundated forest.<br />
uvilla. Brazil. Amazonas: mapati or mapaty.<br />
This species is rather uniform. It resembles the Specimens examined. BRAZIL. AMAZONAS: Rio Soform<br />
ofP. herrerensis <strong>with</strong> oblong to elliptic leaves lim6es, mouth of Rio Jutai, 5 Sep 1975 (2), L. Coelho<br />
339<br />
<strong><strong>an</strong>d</strong> unbr<strong>an</strong>ched basal lateral veins, but c<strong>an</strong> be<br />
(INPA, U); Sao Paulo de Olivenca, 10 Oct 1931<br />
($), Ducke (RB) 25244 (RB); Sao Paulo de Olivenca,<br />
distinguished from the latter by the indument of Creek Belem, 26 Oct-11 Dec 1936 (6), Krukoff8807<br />
the leafy twigs, consisting of only long hairs or (A, F, NY, type collection ofP.ferruginea); Ton<strong>an</strong>tins,<br />
w<strong>an</strong>ting.<br />
19 Feb 1944 (2), Ducke 1527 (A, F, IAN, MG, NY, R,<br />
Glaziou 10070 (according to the label from<br />
UC, US); Rio Solim6es, Sao Antonio de Ila, 2 Dec<br />
1948 (2), Fr6es 23693 (IAN); M<strong>an</strong>aus-Porto Velho<br />
Brazil, Espirito S<strong>an</strong>to, Itapemerim) is probably hwy., BR-319, km 380, 2 km S of Rio Jutai, 13 Oct<br />
the same collection as Schwacke 560.<br />
1974 (6), Pr<strong>an</strong>ce et al. 22859 (INPA, MG, U).<br />
16. <strong>Pourouma</strong> ferruginea St<strong><strong>an</strong>d</strong>ley, Publ. Field<br />
Mus. Nat. Hist., Bot. Sr., 17:181. 1937. Type.<br />
Brazil. Amazonas: Mun. Sao Paulo de Oliven-<br />
ca, Belem Creek, 26 Oct- 1 Dec 1936 (6), Kru-<br />
koff 8807 (holotype, NY; isotypes, A, F).<br />
Fig. 86.<br />
Tree, up to 20 m tall. Leafy twigs ca. 10-15<br />
mm thick, white-puberulous <strong><strong>an</strong>d</strong> <strong>with</strong> dense,<br />
purplish, pluricellular hairs; lenticels conspicuous.<br />
Lamina entire, ovate <strong><strong>an</strong>d</strong> 11-32 x 8-23 cm<br />
or 3-5-fid <strong><strong>an</strong>d</strong> up to ca. 45 x 45 cm, coriaceous,<br />
? plicate, apex rounded to acuminate, base truncate<br />
to cordate; upper surface smooth, hairy on<br />
the main veins, lower surface sparsely subsericeous<br />
to strigose on the main veins, the whole<br />
surface covered <strong>with</strong> white or brownish, arachnoid<br />
hairs, which disappear from the main veins;<br />
lateral veins 8-16 pairs, the basal pair br<strong>an</strong>ched,<br />
occasionally unbr<strong>an</strong>ched, tertiary venation pl<strong>an</strong>e<br />
to slightly prominent beneath; petiole 4-30 cm<br />
long, <strong>with</strong> dense whitish, arachnoid hairs; stipules<br />
3-13 cm long, outside hispid to substrigose,<br />
inside glabrous, caducous. Staminate inflorescences<br />
up to 30 cm long <strong><strong>an</strong>d</strong> up to 20 cm wide;<br />
peduncle 2-9 cm long, peduncle <strong><strong>an</strong>d</strong> br<strong>an</strong>ches<br />
Vernacular names. Brazil. Amazonas: mapatir<strong>an</strong>a.<br />
This species matches P. ovata in the longly<br />
pedunculate, pendulous pistillate inflorescences<br />
<strong>with</strong> globose <strong><strong>an</strong>d</strong> apiculate fruiting peri<strong>an</strong>ths. The<br />
two species are not closely related.<br />
See unnamed collection number 3 (p. 193).<br />
17. <strong>Pourouma</strong> villosa Trecul, Ann. Sci. Nat. Bot.,<br />
Ser. 3, 8: 103. 1847; Miquel in Martius, Fl.<br />
bras. 4(1): 127. 1853. Type. French Gui<strong>an</strong>a.<br />
Without locality, 1840 (6), Leprieur s.n. (holotype,<br />
P). Fig. 87.<br />
<strong>Pourouma</strong> laevis Benoist, Bull. Mus. Hist. Nat. (Paris)<br />
28: 319. 1922; Berg, Fl. Suriname 5(1): 270. 1975.<br />
Type. French Gui<strong>an</strong>a. Acarou<strong>an</strong>y, 1858 (6), Sagot<br />
517 (lectotype, P, chosen here; isolectotypes, B, BR,<br />
F, G, GH, GOET, P, S, U, US).<br />
Tree, up to 30 m tall. Leafy twigs 5-15 mm<br />
thick, yellow(ish)-puberulous to -subvelutinous<br />
to -subhirsute, or (at least on the scars of the<br />
stipules) pale yellow- to whitish-villous, <strong>with</strong><br />
dense, brown, pluricellular hairs. Lamina 3-5lobed<br />
to -fid, sometimes entire <strong><strong>an</strong>d</strong> ovate, rarely<br />
obovate to oblong, 5-40 x 4-42 cm, coriaceous,<br />
rarely chartaceous, apex acuminate, base deeply
178 Flora Neotropica<br />
?~~~~~~~~~~~~~?<br />
;: - ~<br />
?<br />
.<br />
'<br />
.. '".<br />
~<br />
~~~L~~r~~AI ~~~~R~~~??<br />
Ir ?i<br />
~ ..194 ~ . ~ ~<br />
. . ' !':,"..:<br />
r~~~~~~~~':.<br />
. :.:.::.. . . ........... ....<br />
Neg|tive nch ~ ~ ..<br />
TO 2<br />
rr<br />
F; ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ,' ~- ,~ ...............~,':..:,,.<br />
? ' ':.-L,. . . ~ . ,> ~ ~<br />
la"?fr ~~~~~~~<br />
,~'~ .... ,i 4: ,.~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~<br />
r~ ~~~~~~~~~~~~~~~~~~?'',,<br />
-..-.:. ~ 4~ ~~~~<br />
-?. ._~ ...<br />
Jo - ......":.... .<br />
.....: .<br />
FIG. 86. <strong>Pourouma</strong> ferruginea. Leafy twig <strong>with</strong> pistillate inflorescence (Ducke 1527).<br />
....<br />
. . ' . ':'.-,.C.~ . . . . . _<br />
IG86Poruafruine. Laytwi ihpsilt nirsecDce12)<br />
cordate to subcordate, sometimes <strong>with</strong> overlap-<br />
ping lobes; upper surface smooth, hairy on the<br />
main veins, lower surface white-puberulous to<br />
yellow-hirsute on the veins, usually <strong>with</strong> brown,<br />
pluricellular hairs on the main veins, <strong><strong>an</strong>d</strong> white,<br />
arachnoid hairs in the areoles <strong><strong>an</strong>d</strong> on the smaller<br />
veins; lateral veins 6-20 pairs, the basal pair<br />
br<strong>an</strong>ched (<strong><strong>an</strong>d</strong> their basal parts separated from<br />
'
<strong>Pourouma</strong><br />
11<br />
'.ciLnnr'.mon-lik<br />
2 3 4 5<br />
THNE NEW YORK BOTANICAL GARDEN<br />
No". '.3 : j ", -'.<br />
Old G.D: car.p I!. ? "..<br />
Faysetr AIrstri'.)<br />
kNut.v Wnch 1of 2fpmribo.<br />
stilt roots. Fril r'd<br />
odor:?'"<br />
BOTANICAf!' ? O iO<br />
FIG. 87. <strong>Pourouma</strong> villosa. Leafy twig <strong>with</strong> pistillate inflorescences (Mori et al. 8583).<br />
the margin by mesophyll), tertiary venation<br />
slightly prominent beneath; petiole 6-35 cm long,<br />
yellow-hirsute <strong><strong>an</strong>d</strong> <strong>with</strong> long <strong><strong>an</strong>d</strong> short, appres-<br />
sesd, white hairs, <strong><strong>an</strong>d</strong> <strong>with</strong> (dense), brown, arach-<br />
'<br />
179<br />
noid hairs; stipules 3-25 cm long, outside yellowhirsute<br />
to -velutinous or pale yellow- to whitishvillous,<br />
also <strong>with</strong> dense, brown, pluricellular hairs,<br />
inside yellow-hirsute to glabrous; caducous.
180 Flora Neotropica<br />
Staminate inflorescences up to 21 cm long <strong><strong>an</strong>d</strong> <strong><strong>an</strong>d</strong> Serro do Navio, 10 Oct-15 Dec 1976 (st), Rosa<br />
12 cm wide; peduncle 2-10 cm 1126<br />
long, yellow-pu-<br />
(MG). AMAZONAS: M<strong>an</strong>aus, 23 Oct 1969 (8), L.<br />
Coelho 15 (INPA, U), 17 J<strong>an</strong> 1931 (2), Ducke<br />
berulous <strong><strong>an</strong>d</strong> <strong>with</strong> dense<br />
(RB)<br />
brown, pluricellular 25243 (RB), 2 Apr 1976 (st), Macedo s.n. (INPA), 24<br />
hairs; flowers sessile, mostly clustered at the ul- Oct 1978 (8), Rodrigues et al. 10082 (INPA), 1850timate<br />
br<strong>an</strong>ches; tepals 1-2 mm long, ovate to 1851 (8), Spruce s.n. (G), Mar 1912 (2), Ule 8839 (G,<br />
l<strong>an</strong>ceolate, (almost) free or up to halfway con- MG). MARANHAO: Rod. Belem-Brasilia, km 380-375,<br />
28<br />
nate, sparsely hairy; filaments shorter th<strong>an</strong> the Aug 1960 (2), Oliveira 1074 (IAN). PARA: Faro, 3<br />
J<strong>an</strong> 1920 (9), Ducke (MG) 10736 = (RB) 13056 (MG,<br />
tepals. Pistillate inflorescences in fruit up to 19 RB); Rio Br<strong>an</strong>co, Obidos, Serra da Boa Vista, 23 Dec<br />
cm long <strong><strong>an</strong>d</strong> 10 cm wide; peduncle 2-12 cm long, 1913 (9), Ducke (MG) 25225 (MG); upper Rio Cupary,<br />
peduncle <strong><strong>an</strong>d</strong> br<strong>an</strong>ches yellow-puberulous to between Rio Xingu <strong><strong>an</strong>d</strong> Rio Tapaj6s, Sep 1931 (2),<br />
-hirtellous <strong><strong>an</strong>d</strong> <strong>with</strong> dense, brown to red-brown Krukoff 1197 (A, G, NY, P, S, U); Rio Tapaj6s, Boa<br />
or dark<br />
Vista, 1931 (2), Monteiro da Costa 92<br />
brown, pluricellular hairs;flowers ca. 10-<br />
(F); Rio Jari,<br />
Monte Dourado, 14 J<strong>an</strong> 1968 (a), Oliveira 3915 (NY).<br />
40; pedicels up to 0.6 cm long, in fruit up to 1.2 RONDONIA: P6rto Velho-Cuiaba hwy., S<strong>an</strong>ta Barbara,<br />
cm; peri<strong>an</strong>th 4-8 mm long, yellow-puberulous 16 Aug 1968 (8), Pr<strong>an</strong>ce et al. 7160 (F, GH, INPA,<br />
<strong><strong>an</strong>d</strong> <strong>with</strong> dense, dark brown, pluricellular hairs; NY, P, S, U, US).<br />
stigma peltate, 1.5-2.5 mm diam. Fruiting peri- Vernacular names. Surinam. Boroma (Aro<strong>an</strong>th<br />
purple, (sub)ovoid to ellipsoid, ca. 1.5-2 x<br />
wak), gr<strong>an</strong>boesipapaja, kobe (Djoeka), m<strong>an</strong>bos-<br />
0.7-1.5 cm, yellow-puberulous <strong><strong>an</strong>d</strong> <strong>with</strong> sparse papaja, pourouma or puruma (Carib), yarayara<br />
to dense, dark brown, pluricellular hairs.<br />
(Carib). French Gui<strong>an</strong>a. Bois c<strong>an</strong>on,<br />
Distribution<br />
(bouchi) pa-<br />
(Fig. 85). Surinam, French pate or papaye (Paramaka). Brazil. Amapa: im-<br />
Gui<strong>an</strong>a, <strong><strong>an</strong>d</strong> Brazil (Amapa, Amazonas, Para, bafiba bengue; Para: imbafiba br<strong>an</strong>ca, imbafiba<br />
<strong><strong>an</strong>d</strong> Rond6nia); mostly in non-inundated forest, de cheiro uvilha.<br />
sometimes in riverine forest, at low alti- The relationships <strong>with</strong> other <strong>Pourouma</strong> species<br />
tudes.<br />
are not clear. The collections from Amazonas<br />
Specimens examined. SURINAM. Moengo, 16 <strong><strong>an</strong>d</strong><br />
Aug Rond6nia, three collections from Para (the<br />
1951 (st), BBS 297 (U); Brownsberg, 15 Jul 1916 (st), two Ducke collections <strong><strong>an</strong>d</strong> Oliveira 3915) <strong><strong>an</strong>d</strong><br />
BW2095 (U), 13 Sep 1917 (6), BW 3255 (NY, U), 7 the type collection of P. villosa, have long pale<br />
Sep 1917 (6), BW3302 (B, F, MO, P, U), 20 Sep 1918<br />
(), BW 4010 (U, UC), 17 Sep 1921 (9), BW<br />
yellow hairs on the<br />
5390<br />
petioles <strong><strong>an</strong>d</strong> leafy twigs, but<br />
(F,<br />
RB, U), 28 Jun 1924 (6), BW 6509 (U); Brownsweg,<br />
they are similar to the other collections in all<br />
16 Nov 1964 (st), Donselaar 1888 (U); Wilhelmina other features. Together <strong>with</strong> the overlap in dis-<br />
Mts., 10 Sep 1963 (8), Irwin et al. 55576 (B, FHO, GH, tribution this is a reason not to recognize the<br />
M, MO, NY, U, US), 20 Sep 1963 (6), Irwin et al. form <strong>with</strong> long hairs as a distinct<br />
55884 infraspecific<br />
(FHO, GH, NY, S, U, US); Fallawatra, 6 Nov<br />
1971 (2), Jimenez-Saa 1563 = taxon.<br />
LBB 14296 (NY, U);<br />
Lely Mts., 29 Sep 1975 (st), Lindem<strong>an</strong> et al. 540 Cut fruit<br />
(U),<br />
give out a cinnamon-like odor (Mori<br />
5 Oct 1975, Lindem<strong>an</strong> et al. 790 (U); Paris Jacob Creek, et al. 8583).<br />
28 Jun 1965 (st), Maas et al. (LBB) 11017 (U); Kayser The description of P. laevis was based on Ben-<br />
Mts., 29 Oct 1976 (9), Mori et al. 8535 (F, NY, U, US); oist<br />
Sectie O, Nov 1942<br />
341,<br />
(9), Stahel (Woodherb. Sur.) 123B<br />
Sagot 517, <strong><strong>an</strong>d</strong> Sagot 972. Sagot 517<br />
(U), Dec 1942 (9), Stahel (Woodherb. Sur.) 166A<br />
is chosen as the<br />
(U).<br />
lectotype collection.<br />
FRENCH GUIANA. Without locality (9), Aublet s. n.<br />
(G), 1840 (6), Leprieur s.n. (P, type collection of P. 18. <strong>Pourouma</strong> oraria St<strong><strong>an</strong>d</strong>ley & Cuatrecasas in<br />
villosa); Charvein-Acarou<strong>an</strong>y rd., km 1, 27 Oct 1953 Cuatrecasas, Caldasia 7: 301.<br />
(9), BAFOG 93-M(CAY, P, U); St.<br />
1956;<br />
Laurent, Dec<br />
Woodson,<br />
1955<br />
(9), BAFOG 7087<br />
Ann. Missouri<br />
(CAY, NY, P, U); Route<br />
Bot.<br />
de<br />
Gard. 47:167.1960.<br />
Cayenne,<br />
Type.<br />
km 14, 28 Oct 1957 (8), BAFOG 7745 (CAY, NY, P);<br />
Colombia. Valle: Rio Yurumagui, Veneral, 28<br />
Charvein, (9), Benoist 341 (P); Acarou<strong>an</strong>y, 1858 (6), J<strong>an</strong> 1944 (2), Cuatrecasas 15720 (holotype, F).<br />
Sagot 517 (F, G, GH, GOET, P, S, U, U S, lectotype<br />
collection of P. laevis; in F <strong><strong>an</strong>d</strong> P, material of P. mel- Tree, up to 20 m tall. Leafy twigs up to 15 mm<br />
inonii subsp. melinonii under the same number), 1856 thick, yellow-puberulous <strong><strong>an</strong>d</strong> <strong>with</strong> dense, brown,<br />
(9), Sagot 970 (P), (9), Sagot 972 (B, BR, F, G, P, S, pluricellular hairs. Lamina 7-fid, 28-32 x 40<br />
U), (9), Sagot s.n. (U), (st), Sagot s.n. (G).<br />
BRAZIL. AMAPA: 35 km ESE of Oiapoque, 2 Dec<br />
cm, coriaceous, apex acuminate, base deeply cor-<br />
1984 (9), Daly et al. 3790 (BG); Rio Amapari, R<strong>an</strong>cho date <strong>with</strong> overlapping lobes; upper surface<br />
S<strong>an</strong>ta Ana, 30 Sep 1961 (6), Pires et al. 51390 (FHO, smooth, main veins hairy, lower surface puber-<br />
IAN, INPA, MG, MO, NY, US); between P6rto Platon ulous on the veins, arachnoid hairs confined to
<strong>Pourouma</strong> 181<br />
the areoles; lateral veins up to ca. 30 pairs, ter- pressed-puberulous on the main veins, arachtiary<br />
<strong><strong>an</strong>d</strong> quarternary venation prominent be- noid hairs in the areoles <strong><strong>an</strong>d</strong> on the smaller veins;<br />
neath; petiole 30 cm long, yellow-hirsute at the lateral veins 7-12 pairs, basal pair unbr<strong>an</strong>ched,<br />
adaxial side, for the rest sparsely hairy; stipules tertiary venation pl<strong>an</strong>e beneath; petiole 2-9 cm<br />
10-12 cm long, outside yellow-hirsute, <strong><strong>an</strong>d</strong> <strong>with</strong> long, <strong>with</strong> some (very) sparse, white hairs; stipwhite<br />
appressed hairs, occasionally <strong>with</strong> brown, ules 1-5 cm long, outside yellow-appressed-pupluricellular<br />
hairs, inside yellow-hirsute, subper- berulous, inside glabrous, caducous. Staminate<br />
sistent. Staminate inflorescences about 25 cm long inflorescences unknown. Pistillate inflorescences<br />
<strong><strong>an</strong>d</strong> 15 cm wide; peduncle ca. 12 cm long, yellow- in fruit up to 8 cm long <strong><strong>an</strong>d</strong> 5 cm wide; peduncle<br />
puberulous; flowers in clusters of up to 10 flow- 2-3.5 cm long, peduncle <strong><strong>an</strong>d</strong> br<strong>an</strong>ches minutely<br />
ers; tepals free, ca. 2 mm long; filaments shorter yellow-puberulous, the ultimate br<strong>an</strong>ches more<br />
th<strong>an</strong> the tepals. Pistillate inflorescences in fruit densely hairy, <strong><strong>an</strong>d</strong> <strong>with</strong> brown, pluricellular hairs;<br />
up to 22 cm long <strong><strong>an</strong>d</strong> 18 cm wide; peduncle 14- flowers 7-10; pedicel 0.1-0.5 cm long, in fruit up<br />
15 cm long, peduncle <strong><strong>an</strong>d</strong> br<strong>an</strong>ches yellow-pu- to 1 cm; peri<strong>an</strong>th ca. 1.5-2 mm long, yellowberulous<br />
<strong><strong>an</strong>d</strong> <strong>with</strong> dense, dark-brown or red- puberulous; stigma bilobed, ca. 2.5 mm diam.<br />
brown or brown, pluricellular hairs; flowers ca. Fruitingperi<strong>an</strong>th ovoid, ca. 2 x 0.8 cm, sparsely,<br />
100; pedicels ca. 5 mm long, in fruit up to 10 minutely appressed-puberulous.<br />
mm long; peri<strong>an</strong>th 3-5 mm long, yellow-puber- Distribution (Fig. 85). French Gui<strong>an</strong>a <strong><strong>an</strong>d</strong> Braulous<br />
<strong><strong>an</strong>d</strong> <strong>with</strong> dense, dark-brown, pluricel- zil (Amapa); in non-inundated forest.<br />
lular hairs; stigma peltate, 1.5-2 mm diam.<br />
Fruiting peri<strong>an</strong>th ovoid to ellipsoid, ca. 15 x 10<br />
Specimens examined. FRENCH GUIANA. Saiil, 20<br />
Dec 1976 (Q), Gr<strong>an</strong>ville 2744 (CAY), Saul, km 2.7 on<br />
mm, + sparsely hairy.<br />
trail on Montagne Belvedere Sud, 21 Oct 1971 (v),<br />
Distribution (Fig. 85). Colombia (Choc6 <strong><strong>an</strong>d</strong> Oldem<strong>an</strong> B.4132 (CAY, P, U, VEN type collection of<br />
Valle); in lowl<strong><strong>an</strong>d</strong> forest.<br />
P. saulensis).<br />
BRAZIL. AMAPA: 109 km SSE of Oiapoque, 5 Dec<br />
Specimens examined. COLOMBIA. CHOC6: Be- 1984 (v), Mori et al. 17185 (BG).<br />
tween Certegui <strong><strong>an</strong>d</strong> Las Animas, 17 Aug 1976 (9), Gentry<br />
et al. 17804 (BG, U); N of Certegui, 13 Jun 1982 Vernacular name. French Gui<strong>an</strong>a. Bois c<strong>an</strong>on.<br />
(8), Gentry et al. 36797 (JUAM). VALLE: Costa del This species shows similarities to P. ovata in<br />
Pacifico, Rio Yurum<strong>an</strong>gui, Veneral, 28 J<strong>an</strong> 1944 (9), the indument, the short stipules, <strong><strong>an</strong>d</strong> the un-<br />
Cuatrecasas 15720 (F, type collection of P. oraria). br<strong>an</strong>ched, basal, lateral veins. The species are<br />
distinct in the number of lateral veins, the length<br />
of the peduncle of the pistillate inflorescence, <strong><strong>an</strong>d</strong><br />
the shape of the fruiting peri<strong>an</strong>th.<br />
This species resembles P. bicolor subsp. choco<strong>an</strong>a<br />
in m<strong>an</strong>y features (see p. 144). It differs<br />
from the latter in the smooth upper surface of<br />
the lamina, the smooth fruiting peri<strong>an</strong>th, <strong><strong>an</strong>d</strong> the<br />
concentration of the long hairs at the adaxial side<br />
of the petiole. P. oraria may be much closer to<br />
P. bicolor (subsp. choco<strong>an</strong>a) th<strong>an</strong> suggested by<br />
the arr<strong>an</strong>gement oftaxa in the present treatment.<br />
19. <strong>Pourouma</strong> saulensis C. C. Berg, Brittonia<br />
34(1): 36, t. 1. 1982. Type. French Gui<strong>an</strong>a.<br />
Sail, km 2.7 on trail of Montagne Belvedere<br />
Sud, 21 Oct 1971 (9), Oldem<strong>an</strong> B.4132 (ho-<br />
lotype, U; isotypes, CAY, P, VEN). Fig. 88.<br />
Tree, up to 11 m tall. Leafy twigs 3-5 mm<br />
thick, minutely puberulous to glabrous. Lamina<br />
entire, (broadly) ovate to elliptic, 6-14 x 3-9.5<br />
cm, coriaceous, apex (sub)acuminate, base<br />
rounded to obtuse; upper surface smooth, some<br />
sparse, white hairs <strong><strong>an</strong>d</strong> brown, pluricellular hairs<br />
on the main veins, lower surface white-, ap-<br />
20. <strong>Pourouma</strong> ovata Trecul, Ann. Sci. Nat. Bot.,<br />
Ser. 3, 8: 101. 1847, Miquel in Martius, Fl.<br />
bras. 4(1): 132.1853. Type. Brazil. Para: Without<br />
locality, <strong>with</strong>out date (d), Ferreira s.n. (holotype,<br />
P). Fig. 89.<br />
<strong>Pourouma</strong> longipendula Ducke, Notizbl. Bot. Gart.<br />
Berlin-Dahlem 11: 581. 1932. Type. Brazil. Amazonas:<br />
M<strong>an</strong>aus, falls of Rio Taruma; 17 Oct 1929<br />
(v), Ducke (RB) 23608 (holotype, RB, not seen; isotypes,<br />
B, G, S, US).<br />
Tree, up to 25 m tall. Leafy twigs 4-9 mm<br />
thick, sparsely white-appressed-puberulous, more<br />
densely haired at the nodes. Lamina entire, elliptic<br />
to ovate, 7-26 x 3-13 cm, coriaceous, apex<br />
short acuminate, base acute to obtuse to rounded<br />
or sometimes truncate; upper surface smooth,<br />
sparsely hairy <strong><strong>an</strong>d</strong> occasionally <strong>with</strong> some brown,
182 Flora Neotropica<br />
[--<br />
i. "<br />
Q~~~~~~~~~~~,"<br />
.'<br />
? "<br />
FIG. 88. <strong>Pourouma</strong> saulensis. Leafy twig, (schematic) pistillate inflorescence, <strong><strong>an</strong>d</strong> fruiting peri<strong>an</strong>th (Gr<strong>an</strong>ville~~~~~~<br />
2744); pstillat flower Oldem<strong>an</strong>B.413')<br />
FIG. 88. <strong>Pourouma</strong> saulensis. Leafy twig, (schematic) pistillate inflorescence, <strong><strong>an</strong>d</strong> fruiting peri<strong>an</strong>th (Gr<strong>an</strong>ville<br />
2744); pistillate flower (Oldem<strong>an</strong> B.4132).<br />
pluricellular hairs on the midrib, lower surface<br />
white-puberulous on the veins, arachnoid hairs<br />
confined to the areoles; lateral veins 9-21 pairs,<br />
basal pair unbr<strong>an</strong>ched or occasionally br<strong>an</strong>ched,<br />
tertiary venation slightly prominent beneath;<br />
petiole 2-7.5 cm long, subglabrous to sparsely<br />
white hairy; stipules 1.5-3 cm long, outside yel-<br />
low-velutinous, usually <strong>with</strong> brown, pluricellular<br />
hairs, inside yellow-(or or<strong>an</strong>ge-brown- or<br />
white-)sericeous, caducous. Staminate inflores-<br />
cences up to 10 cm long <strong><strong>an</strong>d</strong> 9.5 cm wide; pe-<br />
duncle 1-5.5 cm long, sparsely puberulous to<br />
glabrescent, ultimate br<strong>an</strong>ches usually more<br />
densely yellow- to or<strong>an</strong>ge-puberulous; flowers<br />
sessile or occasionally pedicellate, mostly in<br />
subglobose heads, these 3-7 mm diam.; tepals<br />
ca. 1 mm long, connate, rather densely hairy;<br />
filaments shorter th<strong>an</strong> the tepals. Pistillate inflorescences<br />
in fruit up to 38 cm long <strong><strong>an</strong>d</strong> 11 cm<br />
wide; peduncle up to 32 cm long, peduncle <strong><strong>an</strong>d</strong><br />
br<strong>an</strong>ches puberulous to almost glabous; flowers<br />
7-20; pedicel 0.2-2.4 cm long, in fruit 1-3 cm,<br />
obconical, conspicuously lenticellate; peri<strong>an</strong>th 3-<br />
5 mm long, densely short-velutinous; stigma pel-<br />
..<br />
-<br />
'
<strong>Pourouma</strong> 183<br />
Ak~~~~~~~~~~~~?IIk<br />
i.<br />
| I 3 -2^ 2 A? :.<br />
T rM w Tons BOTANICAL GAIM'W<br />
liBlTIm n lA L AiASS PA MZI.<br />
ii~L m.---.-V---lmDI,<br />
_ r ,.l. _ l,-~v kd L<br />
of tioe rm ch liho.<br />
oree<br />
?<br />
wdll<br />
Frest om terra firme on cly.<br />
Tm 1in x 20cm; frvit green, maturing red,<br />
clew light bow . ri. oraa<br />
w<br />
' g- '*--- J<br />
.<br />
'l -<br />
.*<br />
ovd'sember 1-8....<br />
FIG. 89. <strong>Pourouma</strong> ovata. Leafy twig <strong>with</strong> pistillate inflorescences<br />
(Steward et al. 107).
184 Flora Neotropica<br />
tate, bilobed, ca. 1.5-2 mm diam. Fruiting peri-<br />
<strong>an</strong>th red (mature?), (depressed-)globose, ca. 1.5<br />
cm diam., puberulous.<br />
Distribution (Fig. 85). Colombia (Vaup6s),<br />
Venezuela (Bolivar), Peru (Loreto), <strong><strong>an</strong>d</strong> Brazil<br />
(Acre, Amazonas, <strong><strong>an</strong>d</strong> western Para); in non-<br />
inundated forest.<br />
Vernacular names. Peru. Loreto: chullachaqui,<br />
chullachaqui bl<strong>an</strong>co, chullachaqui caspi, sacha<br />
uvillos. Brazil. Amazonas: imbafbar<strong>an</strong>a.<br />
This species resembles P. ferruginea in the longpedunculate<br />
<strong><strong>an</strong>d</strong> pendulous pistillate inflorescences<br />
<strong><strong>an</strong>d</strong> the globose <strong><strong>an</strong>d</strong> apiculate fruiting<br />
peri<strong>an</strong>th, but is in other features quite distinct<br />
from the latter. In the<br />
Specimens examined. COLOMBIA. VAUPES: Mon-<br />
vegetative parts it shows<br />
fort, 30 Nov 1952 (9), Romero-Cast<strong>an</strong>eda 3848 resembl<strong>an</strong>ces to P. saulensis in several characters<br />
(F,<br />
MBG, NY); Rio Apaporis, Soratama, 26 Mar 1952 (2), (seep. 181).<br />
Schultes et al. 16009 (C, U, US).<br />
VENEZUELA. BOLIVAR: Sierra Ichfin, Salto Maria<br />
Espuma, 28 Dec 1961 (6), Steyermark 90368 (NY, U, 21. <strong>Pourouma</strong> elliptica St<strong><strong>an</strong>d</strong>ley, Publ. Field<br />
VEN).<br />
Mus. Nat. Hist, Bot. Ser., 17:181. 1937. Type.<br />
PERU. LORETO: Prov. Requena, Rio Ucayali, Ar-<br />
Brazil. Amazonas: Mun. Sao Paulo de Olivenboretum<br />
Jenaro Herrera, Aug-Sep 1976 (st), Bernardi<br />
5.16.5 (U), Aug-Sep 1976 (9), Bernardi s.n. (tree 4/8) 9a, nr. Palmares, 11 Sep-26 Oct 1936 (6), Kru-<br />
(U), 3 J<strong>an</strong> 1974 (9), Diaz 22-A (G); Prov. Maynas, Rio koff8388 (holotype, NY; isotypes, A, BR, F,<br />
N<strong>an</strong>ay, Mish<strong>an</strong>a, 23 Mar 1979 (st), Gentry et al. 26049 G, LE, MO, P, S, U). Fig. 90.<br />
(U), 24 Mar 1979 (st), Gentry et al. 26177 (U), 12 J<strong>an</strong><br />
1983 (st), Gentry et al. 39379 (BG); Prov. Requena, Tree, up to ca. 15 m tall. Leafy twigs ca. 10<br />
Arboretum Jenaro Herrera, 9 Dec 1980 (9), Vdsquez mm thick, densely yellow-hirsute <strong><strong>an</strong>d</strong> yellowet<br />
al. 1004 (BG).<br />
BRAZIL. ACRE: Cruzeiro do Sul, 11 Feb 1976 subhispidulous, thus a mixture of hairs<br />
(9),<br />
distinctly<br />
Monteiro et al. 309 (INPA), 1 Mar 1976 (2), Ramos et different in length, moreover, <strong>with</strong> sparse redal.<br />
204 (INPA). AMAZONAS: M<strong>an</strong>aus, Reserva Florestal dish-brown, pluricellular hairs <strong><strong>an</strong>d</strong> sparse, white,<br />
Ducke, 27 Nov 1957 (st), D. Coelho (INPA) 5979 arachnoid hairs. Lamina entire, elliptic to ob-<br />
(INPA), 13 Sep 1957 (6), L. Coelho (INPA) 5808 (U); long, 13-30 x 6.5-20 cm, coriaceous, apex<br />
M<strong>an</strong>aus, waterfalls of Rio Taruma, 17 Oct 1929 (9),<br />
Ducke (RB) 23608 (B, G, RB, S, US, type collection abruptly, short-acuminate, base acute to roundof<br />
P. longipendula); M<strong>an</strong>aus, Reserva Florestal Ducke, ed to truncate; upper surface smooth, hairy on<br />
17 Oct 1929 (9), Killip et al. 30139 (A, F, NY, US); the main veins, to puberulous on the smaller<br />
Humaita, 7-18 Nov 1934 (2), Krukoff 7073 (A, B, BR, veins, lower surface yellow-(sub)hirsute on the<br />
F, G, GB, LE, MO, NY, RB, S, U, US); Sao Paulo de main<br />
Olivenca, 26 Nov 1936 (9), Krukoff 8695<br />
veins, to puberulous on the smaller veins,<br />
(A, B, BR,<br />
F, G, MO, NY, P, S, U, US); M<strong>an</strong>aus-P6rto Velho rd., white, arachnoid hairs in the areoles <strong><strong>an</strong>d</strong> on the<br />
km 240, 21 Nov 1973 (9), Lleras et al. P.19586 (F, smaller veins, lateral veins 10-16 pairs, the basal<br />
INPA, MO, NY, P, S, U, US); M<strong>an</strong>aus, Reserva Flo- pair unbr<strong>an</strong>ched or sparsely <strong><strong>an</strong>d</strong>/or faintly<br />
restal Ducke, 10 Nov 1965 (9), Loureiro (INPA) 16968 br<strong>an</strong>ched, tertiary venation almost pl<strong>an</strong>e be-<br />
(U); Torquato-Tapajos rd., km 118, 28 Aug 1975 (9),<br />
Loureiro et al. (INPA) 50623 (INPA); M<strong>an</strong>aus-Cara- neath; petiole 5-13.5 cm long, yellow-hirsute <strong><strong>an</strong>d</strong><br />
carai rd., km 57, 16 Sep 1976 (9), Mota 680 (INPA); <strong>with</strong> sparse to dense, white, arachnoid hairs; stip-<br />
Tapuruquara, 4 Jul 1972 (9), Pr<strong>an</strong>ce et al. 15393 (F, ules 3-9 cm long, caducous, outside yellow-hir-<br />
GH, INPA, M, NY, P, S, U); km 245, 13 Mar 1974 sute, inside glabrous. Staminate inflorescences<br />
(9), Pr<strong>an</strong>ce et al. 20454 (INPA, MO, NY, S, U, US),<br />
km 380, 13 Oct 1974 (9), Pr<strong>an</strong>ce et al. 22864 up to 19 cm<br />
(INPA,<br />
long <strong><strong>an</strong>d</strong> up to 14 cm wide; peduncle<br />
MO, NY, P, S, U, US); M<strong>an</strong>aus, Reserva Florestal 4-7 cm long, yellow-hirsute on the peduncle to<br />
Ducke, 21 Aug 1965 (9), Rodrigues 5449 (NY, U); hispidulous on the ultimate br<strong>an</strong>ches, sometimes<br />
M<strong>an</strong>aus-Itacoatiara rd., 3 Sep 1965 (9), Rodrigues et also white, arachnoid hairs; flowers sessile, not<br />
al. 7081 (U), 26 Nov 1965 (9), Rodrigues et al. 7292<br />
(U); M<strong>an</strong>aus-Caracarai rd., km 60, 20 Aug 1976<br />
densely clustered; tepals ca. 1.5 mm long, almost<br />
(6),<br />
Shima et al. 14 (INPA); between Rio Cast<strong>an</strong>ho <strong><strong>an</strong>d</strong> free, densely hispidulous; filaments shorter th<strong>an</strong><br />
Rio Tup<strong>an</strong><strong>an</strong>, 14 Jul 1972 (8), M. F. Silva et al. 742 the tepals. Pistillate inflorescences unknown.<br />
(U); 18 Jul 1972 (9), M. F. Silva et al. 869 (INPA); Distribution (Fig. 85). Brazil (Amazonas); in<br />
M<strong>an</strong>aus-P6rto Velho rd., km 250, 5 J<strong>an</strong> 1974 (9), Stew- non-inundated forest.<br />
ard et al. P.20156 (INPA, U). PARA: Without locality,<br />
(6), Ferreira s.n. (P, type collection of P. ovata); Cuiaba- Specimens examined. BRAZIL. AMAZONAS: Espe-<br />
S<strong>an</strong>tarem rd., km 1230, (9), Pr<strong>an</strong>ce et al. 25530 (F, r<strong>an</strong>ga, mouth of Rio J<strong>an</strong>vari, 21 Sep 1931 (6), Ducke<br />
MO, RB, S, U); Rio Trombetas (6), N. T. Silva et al. (RB) 25246 (RB); Sao Paulo de Olivenca, nr. Palmares,<br />
4740 (MG, U). RORAIMA: M<strong>an</strong>aus-Caracarai rd., km 11 Nov-26 Oct 1936 (6), Krukoff 8388 (A, BR, F, G,<br />
343, 18 Nov 1977 (9), Steward et al. 107 (U). LE, MO, NY, P, S, U, type collection of P. elliptica).
<strong>Pourouma</strong> 185<br />
I I<br />
STPE<br />
,E Fn CL1 |SOP Y<br />
r.<br />
Tr e rn rt. 'mh; trunk 41 G<br />
Safe o AmSaonm: r u.odeO<br />
near Plmamres. Sept. Il-OCt. 26. 1M91<br />
.<br />
.di- tb.td .: thlu th. New YO3 k<br />
hlod= 6 tc .'bre 4937<br />
FIG. 90. <strong>Pourouma</strong> elliptica. Leafy twig <strong>with</strong> staminate inflorescence<br />
(Krukoff<br />
8388).
186 Flora Neotropica<br />
Vernacular names. Brazil. Amazonas: mapati<br />
or mapaty.<br />
22. <strong>Pourouma</strong> bolivarensis C. C. Berg, Brittonia<br />
34(1): 39, t. 3. 1982. Type. Venezuela. Bolivar:<br />
Cerro Venamo, SW part, upper Rio Venamo,<br />
10 J<strong>an</strong> 1964 (9), Steyermark et al. 92936 (ho-<br />
lotype, VEN; isotypes, NY, U). Fig. 91.<br />
5(1): 278, t. 14b. 1975; Burger, Fieldi<strong>an</strong>a Bot.<br />
40: 202, t. 22. 1977. Type. French Gui<strong>an</strong>a.<br />
Saint Je<strong>an</strong> de Maroni, 17 Mar 1914 (9), Benoist<br />
960 (lectotype, P, chosen here, cf. Berg, Fl.<br />
Suriname 5(1): 278. 1975). Fig. 82b.<br />
<strong>Pourouma</strong> aurea Ule ex Mildbraed, Notizbl. Bot. Gart.<br />
Berlin-Dahlem 10: 418. 1928; Ule, Bot Jahrb. Syst.<br />
40:149. 1907, nomen. Type. Brazil. Acre: Rio Jurua-<br />
Mirim, Aug 1901 (s), Ule 5718<br />
Tree, up to 15 m tall.<br />
(holotype, B; iso-<br />
Leafy twigs 5-8 mm types, G, MG).<br />
thick, puberulous. Lamina entire, broadly ellip- <strong>Pourouma</strong> folleata Macbride, Publ. Field Mus. Nat.<br />
tic to obovate, sometimes broadly ovate or sub- Hist., Bot. Ser., 8(2): 114. 1930, l.c. 13(2.2): 292.<br />
orbicular, 5-19 x 3-14 1937.<br />
cm, coriaceous, apex Type. Peru. Junin; Ch<strong>an</strong>chamayo valley, 1924-<br />
1927<br />
shortly acuminate to rounded to emarginate, base<br />
(8), C. Schunke 416 (holotype, F; isotype, B).<br />
<strong>Pourouma</strong> isophlebia St<strong><strong>an</strong>d</strong>ley, Publ. Field Mus. Nat.<br />
acute to obtuse; upper surface smooth, sparsely Hist., Bot. Ser., 17: 182. 1937. Type. Brazil. Amawhite-hirtellous<br />
or glabrous on the midrib, lower zonas: Mun. Sao Paulo de Oliven9a, nr. Palmares,<br />
surface appressed whitish-pubescent on the main 11 Sep-26 Oct 1936 (9), Krukoff8037 (holotype, NY;<br />
veins, arachnoid hairs in the areoles <strong><strong>an</strong>d</strong> ? cov- isotypes, A, BR, G, LE, P, S, U, US).<br />
<strong>Pourouma</strong><br />
ering the smaller veins; lateral<br />
subplicata<br />
veins 6-10<br />
St<strong><strong>an</strong>d</strong>ley, Publ. Field Mus. Nat.<br />
pairs, Hist., Bot. Ser., 17: 184. 1937. Type. Brazil. Acre:<br />
basal pair <strong><strong>an</strong>d</strong> sometimes the second pair Nr. mouth of Rio Macauh<strong>an</strong>, 4 Aug 1933 (9), Krukoff<br />
br<strong>an</strong>ched, tertiary venation pl<strong>an</strong>e or slightly 5282 (holotype, F; isotypes, A, G, LE, M, MO, NY,<br />
prominent beneath; petiole 1-6.5 cm long, pu- S, U, UC, US).<br />
berulous; stipules 2-6<br />
<strong>Pourouma</strong><br />
cm<br />
umbellata<br />
long, outside St<strong><strong>an</strong>d</strong>ley, Publ. Field Mus. Nat.<br />
yellow- Hist., Bot. Ser., 17: 185. 1937. Type. Brazil. Amapuberulous,<br />
inside glabrous, caducous. Stami- zonas: Mun. Humaita, between Rio Livramento <strong><strong>an</strong>d</strong><br />
nate inflorescences unknown. Pistillate inflores- Rio Ipixuna, 7-18 Nov 1934 (9), Krukoff 7071 (hocences<br />
in fruit up to 10.5 cm long <strong><strong>an</strong>d</strong> 2 cm wide; lotype, F; isotypes, A, BR, G, GB, LE, MO, NY,<br />
peduncle 3-7 cm long, peduncle <strong><strong>an</strong>d</strong> br<strong>an</strong>ches RB, S, U, US).<br />
<strong>Coussapoa</strong> emarginata Killip ex<br />
yellow-puberulous; flowers 7-12; pedicel 0.2-1<br />
Macbride, Publ. Field<br />
Mus. Nat. Hist., Bot. Ser., 13(2.2): 296. 1937. Type.<br />
cm long, in fruit up to 2 cm; peri<strong>an</strong>th ca. 3 mm Peru. Loreto: Mishuyacu, nr. Iquitos, 24-28 Sep 1929<br />
long, puberulous; stigma peltate, more or less (9), Killip & A. C. Smith 29955 (holotype, US; isobilobed,<br />
2-4 mm diam. Fruiting peri<strong>an</strong>th ma- types, B, F).<br />
roon-red, ca. 1.8 x 1<br />
<strong>Pourouma</strong> cuatrecacasii<br />
cm, puberulous.<br />
St<strong><strong>an</strong>d</strong>ley in Cuatrecasas, Revista<br />
Acad. Colomb. Ci. Exact.<br />
Distribution<br />
9(36/37): 339. 1956.<br />
(Fig. 85). Venezuela (Bolivar); in Type. Colombia. Vaupes: M<strong><strong>an</strong>d</strong>i, nr. Mitfi, 24 Oct<br />
humid, (sub)-mont<strong>an</strong>e forest, at altitudes be- 1939 (9), Cuatrecasas 7299 (holotype, US).<br />
tween 900 <strong><strong>an</strong>d</strong> 1350 m.<br />
<strong>Pourouma</strong> umbellifera Burger, Phytologia 26: 430. 1973.<br />
Type. Costa Rica. Heredia: Nr. Rio Sarapiqui, Ti-<br />
Specimens examined. VENEZUELA. BOLIVAR: Cer- rimbina, 21 J<strong>an</strong> 1972 (9), Lent 2327 (holotype, F;<br />
ro Venamo, Rio Venamo, 8 J<strong>an</strong> 1964 (9), Steyermark isotypes, B, US).<br />
et al. 92850 (U, VEN); Cerro Venamo, SW part, upper<br />
Rio Venamo, alt. 950-1000 m, 10 J<strong>an</strong> 1964 (9), Stey- Tree, up to ca. 35 m tall. Leafy twigs 2-13 mm<br />
ermark et al. 92936 (NY, U, VEN, type collection of thick, yellow-sericeous to -hirsute, or appressed-<br />
P. bolivarensis); Rio Cuyuni, Rio Anawaray-parui, 1300-<br />
1350<br />
puberulous,<br />
m, 25 Dec 1970 occasionally almost<br />
(st), Steyermark et al.<br />
glabrous, some-<br />
104466<br />
times <strong>with</strong><br />
(NY, VEN).<br />
sparse brown, pluricellular hairs.<br />
Lamina entire, narrowly to broadly obovate to<br />
Vernacular names. Venezuela. Bolivar: cay- (broadly) elliptic, rarely to oblong, 4-45 x 1-15<br />
wari-cay-yek (Arekuna).<br />
cm, coriaceous to chartaceous, apex (long) acu-<br />
This species resembles P. minor, from which minate to mucronate or rounded to emarginate,<br />
it differs in the sparser indument, br<strong>an</strong>ched pis- base acute to rounded, rarely to retuse; upper<br />
tillate inflorescences, <strong><strong>an</strong>d</strong> br<strong>an</strong>ched basal lateral surface smooth, yellow-sericeous, hirtellous, hirveins.<br />
sute, or puberulous on the midrib, occasionally<br />
brown, pluricellular hairs on the main veins, low-<br />
23. <strong>Pourouma</strong> minor Benoist, Bull. Mus. Nat. er surface yellow-sericeous to -velutinous or to<br />
Hist. (Paris) 30: 103. 1924; Berg, Fl. Suriname -hirsute on the main veins (minutely) puberulous
<strong>Pourouma</strong> 187<br />
. . .<br />
FIG. 91. <strong>Pourouma</strong> bolivarensis. Leafy twig <strong>with</strong> pistillate inflorescences (Steyermark et al. 92936); sterile<br />
leafy twig (Steyermark et al. 104466).<br />
to hirtellous on the smaller veins, or<strong>an</strong>ge, pluri-<br />
cellular hairs present or not, arachnoid hairs in<br />
the areoles, usually ? extending to the smaller<br />
veins; lateral veins 6-26 pairs, basal pair un-<br />
br<strong>an</strong>ched, tertiary venation (almost) pl<strong>an</strong>e be-<br />
neath; petiole 0.7-15 cm long, yellow-velutinous<br />
to -sericeous or hirsute; stipules 1-15 cm long,<br />
outside yellow-sericeous to -velutinous, some-
188 Flora Neotropica<br />
times -hirtellous, sometimes <strong>with</strong> or<strong>an</strong>ge-brown, (F, VEN), (2), Steyermark 60414 (F, VEN); Rio Chipluricellular<br />
hairs, inside glabrous, caducous. c<strong>an</strong><strong>an</strong>, Puerta Lema, SE of El Dorado, 24 Aug 1961<br />
Staminate inflorescences up to 12 cm long <strong><strong>an</strong>d</strong><br />
(2), Steyermark 89506 (NY, U, VEN); Venezuel<strong>an</strong>-<br />
Brazili<strong>an</strong> frontier, Serr<strong>an</strong>ia<br />
7.5 cm wide; peduncle 1.5-8.5 cm<br />
Pia-soi, 5-6 J<strong>an</strong> 1962 (st),<br />
long, yellow- Steyermark 90620 (VEN).<br />
sericeous to -velutinous; flowers sessile or pedi- GUYANA. K<strong>an</strong>uku Mts., Iramaip<strong>an</strong>g, Nov 1948 (6),<br />
cellate, diffusely distributed along the ultimate Wilson-Browne 549 (=FD 5948) (NY).<br />
br<strong>an</strong>ches or occasionally + clustered; tepals 1.5- SURINAM. Kabalebo, 12 Nov 1976 (st), Heyde et<br />
al. 32<br />
3 mm long, connate, puberulous to almost (U), 13 Nov 1976 (9), Heyde et al. 39<br />
gla-<br />
(U); River<br />
Nassau, Mts, 16 Feb 1949 (2), L<strong>an</strong>jouw et al. 2214<br />
brous; filaments shorter th<strong>an</strong> the tepals. Pistillate (NY, U); Paris Jacob Creek, 29 Jun 1965 (st), Maas et<br />
inflorescences in fruit up to 12 cm long <strong><strong>an</strong>d</strong> 7 cm al. (LBB) 11027 (U); Tap<strong>an</strong>ahony R., at mouth of<br />
wide, subumbellate; peduncle 1-9 cm long, yel- Paloemeu R., Aug 1959 (st), Schulz 8161 (U); Goddo,<br />
low-sericeous; flowers 2-11; pedicel 0.2-1.2 cm<br />
26 J<strong>an</strong> 1926 (9), Stahel (Exp. Wilhelminageb.) 82 (F,<br />
long, in fruit up to 3.5 cm; peri<strong>an</strong>th 2-6 mm<br />
U); Voltzberg, Troon (LBB) 16310 (U).<br />
FRENCH GUIANA. St. Laurent, 13 J<strong>an</strong> 1950 (9),<br />
long, usually densely hairy; stigma knob-shaped, BAFOG 5042 (P), 2 J<strong>an</strong> 1956 (2), BAFOG 7152 (CAY,<br />
variously lobed, 2.5-6 mm diam. Fruiting peri- NY, P, U), 16 J<strong>an</strong> 1956 (2), BAFOG 7173 (CAY, NY,<br />
<strong>an</strong>th ovoid <strong>with</strong> <strong>an</strong> apiculate apex, ca. 1-2.5 x P, U), 8 Feb 1956 (2), BAFOG 7261 (NY, P), 10 Feb<br />
1-1.5<br />
1956<br />
cm, reddish,<br />
(2), BAFOG 7268<br />
hairy.<br />
(CAY, NY, P, U); Saint Je<strong>an</strong><br />
de Maroni, 17 Mar 1914<br />
Distribution<br />
(2), Benoist 960<br />
(Fig. 85). From Nicaragua to the<br />
(P, lectotype<br />
collection of P. minor); 16 Mar 1914 (2), Benoist 978<br />
Amazon Basin <strong><strong>an</strong>d</strong> the Gui<strong>an</strong>a regions; in non- (P); Aug 1973 (st), Garnier 112 (CAY); between Saut<br />
inundated forest at altitudes up to ca. 1500 m. Emerillon <strong><strong>an</strong>d</strong> Etats-Unis, 21 Aug 1970 (st), Gr<strong>an</strong>ville<br />
581 (CAY, P, U); Trois Sauts, 3 Nov 1974 (9), Lescure<br />
Specimens examined. NICARAGUA. Rio S<strong>an</strong> Ju<strong>an</strong>, 422 (CAY); upper Approuage R., Creek Par6pou, 7<br />
nr. Caino Chotaleno, 20 km NW of El Costillo, 7-9 Feb 1967 (st), Oldem<strong>an</strong> 2504 (CAY, NY, P, U); Ap-<br />
Mar 1978 (st), Neill 3389 (BG).<br />
prouage R., Pierrette, 14 Sep 1968 (st), Oldem<strong>an</strong> 2821<br />
COSTA RICA. ALAJUELA: 15 Sep 1973 (6), Hart- (CAY, P); Godebert, <strong>with</strong>out date (st), Wachenheim<br />
shorn 1295 (F); Corazon de Jesus, 29 Mar (6), Hart- 18 (P), 23 Jun 1921 (6), Wachenheim 467 (P), 13 Jul<br />
shorn 1431 (F, U). HEREDIA: Tirimbina, 12-15 Aug 1921 (st), Wachenheim s.n. (A, E, U, US).<br />
1971 (st), Burger et al. 8137 (F); Las Horquetas, 25 ECUADOR. NAPO: Rd. Coca-Lago Agrio, 9 km NE<br />
May 1973 (6), Hartshorn 1224 (F, MO, U); nr. Rio of Rio Coca, 20 Mar 1980 (st), Br<strong><strong>an</strong>d</strong>byge et al. 30252<br />
Sarapiqui at Tirimbina, 21 J<strong>an</strong> 1972 (Q), Lent 2327 (F, (AAU); Guayusa, 2 hr upstream Rio Coca from Coca,<br />
GH, MO, NY, U, US, type collection of P. umbelli- 24 Mar 1980 (st), Br<strong><strong>an</strong>d</strong>byge et al. 30327 (AAU); Afnfera);<br />
La Virgen, 23 Jul 1979 (6), Stevens 13340 (U). <strong>an</strong>gu, 30 May-21 Jun 1982 (st), SEF8617, 9033, 9159<br />
LIMON: Between Siquirres <strong><strong>an</strong>d</strong> Turrialba, 10 Oct 1972 (U); MORONA-SANTIAGO: nr. Bomboiza, 27 Sep 1985<br />
(st), Holdridge 6818 (F, MO, U); Guapiles, 12-13 Mar (st), Shakaim RBAE-26 (BG).<br />
1924 (st), St<strong><strong>an</strong>d</strong>ley 37121 (US); Cairo, 18-19 Feb 1926 PERU. AMAZONAS: Rio S<strong>an</strong>tiago, nr. Caterpiza, 2<br />
(juv), St<strong><strong>an</strong>d</strong>ley et al. 48600 (US).<br />
Nov 1979 (6), Huashikat 1150 (U); mouth of Rio S<strong>an</strong>-<br />
PANAMA. COCLE: El Cope, 1 Sep 1977 (6), Berg tiago, 8-13 Oct 1962 (6), Wurdack 2169 (F, G, GH,<br />
400 (U); 25 Nov 1977 (st), Folsom et al. 6469 (U). SAN NY, P, S, UC). HUANUCO: Rio Pachitea, Tournavista,<br />
BLAS: Cerro S<strong>an</strong> Jose, Yar Birea, 5 Feb 1986 (st), Nevers 8 Aug 1967 (6), J. Schunke V. 2139 (F, G, GH,<br />
et al. 6997<br />
NY,<br />
(MO).<br />
US); Prov. Leoncia Prado, Tingo Maria, 17 Feb 1964<br />
COLOMBIA. AMAZONAS-VAUPES: Rio Apaporis, (st), Vasquez 17 (F). JUNiN: S<strong>an</strong> Ram6n, Aug 1925 (6),<br />
between Rio K<strong>an</strong><strong>an</strong>ari <strong><strong>an</strong>d</strong> Rio Pacoa, 1-15 Dec 1951 C. Schunke A-95 (F, NY, US); Ch<strong>an</strong>chamayo Valley,<br />
(2), Garcia-Barriga 13836 (NY), 28 Sep 1951, Schultes 1924-1927 (8), C. Schunke 416 (B, F, type collection<br />
et al. 14148 (U). ANTIOQUIA: Between Dos Bocas <strong><strong>an</strong>d</strong> ofP.folleata). LORETO: Puerto Almendras, 7 Aug 1973<br />
Anori, 24-31 May 1973 (6), Soejarto et al. 4063 (A, (st), Ayala 341 (U); Rio N<strong>an</strong>ay, halfway between Iqui-<br />
MO). CHOCO: Novita, Cerro Torra, 22 Feb 1977 (2), tos <strong><strong>an</strong>d</strong> S<strong>an</strong>ta Maria de N<strong>an</strong>ay, 31<br />
Forero<br />
May 1975<br />
et al. 3145<br />
(st), Gen-<br />
(RB, U). META: Sierra de la Ma- try et al. 22381 (U), 23 Mar 1979<br />
carena, 19<br />
(st),<br />
J<strong>an</strong> 1950<br />
Gentry et al.<br />
(6), Philipson et al. 2134 (US). 26049 (U), 24 Mar 1979 (st), Gentry et al. 26177<br />
VALLE: Rio<br />
(U);<br />
Anchicaya, Alto Yunda, 1000 m, May 1972 Prov. Maynas, Y<strong>an</strong>amono Explorama Tourist<br />
(Q), Hilty M-86<br />
Camp,<br />
(MO). VAUPES: M<strong><strong>an</strong>d</strong>i, nr. Mitfi, 24 1 Jul 1983 (st), Gentry et al. 42568<br />
Oct<br />
(BG);<br />
1939<br />
Mishuyacu,<br />
(2), Cuatrecasas 7299 (US, type collection of nr. Iquitos, 24-28 Sep 1929 (2),<br />
P.<br />
Killip & Smith 29955<br />
cuatrecacasii); Rio Inirida, S<strong>an</strong> Joaquin, 27 J<strong>an</strong> 1955 (US, B, F, type collection of <strong>Coussapoa</strong><br />
(6), Fern<strong><strong>an</strong>d</strong>ez 2019<br />
emarginata);<br />
(US).<br />
Yurimaguas-Tarapoto rd., km 14, 8 Nov 1967 (2),<br />
VENEZUELA. AMAZONAS: El Dorado-Sta. Elena Soria S. 21 (F, G, US). MADRE DE DIos: Rio<br />
rd., S of El<br />
M<strong>an</strong>u,<br />
Dorado, km 79, 25 Feb 1959 (2), Bernardi Cocha Cashu station, 2 Sep 1976<br />
7250<br />
(9), Foster et al. 5019<br />
(F, G, NY, VEN); Sierra Parima, Simarawochi, (F), 23 Oct 1979 (6), Foster et al. 7199<br />
18 Apr-23 May 1973<br />
(F); Rio Tam-<br />
(Q), Steyermark 107064 (F, U). bopata, mouth of Rio<br />
BOLiVAR:<br />
D'Orbigny, 4 Mar 1981<br />
Between S<strong>an</strong>ta<br />
(st),<br />
Teresita de Kav<strong>an</strong>ayen <strong><strong>an</strong>d</strong> Gentry et al. 31938, 31976<br />
Rio Pacairao, 20-21<br />
(U); Rio<br />
Nov 1944<br />
M<strong>an</strong>u, Cocha Ca-<br />
(6), Steyermark 60399 shu Station, 8 Aug 1983 (st), Gentry et al. 43454 (BG);
<strong>Pourouma</strong> 189<br />
upper Rio Madre de Dios, 17 Jun 1954 (6), Rauh 1609 Thomas et al. 4794 (BG). PARA: Cuiaba-S<strong>an</strong>tarem rd.,<br />
(F). UCAYALI: Pucallpa-Tingo Maria rd., S<strong>an</strong> Alej<strong>an</strong>- km 919, 13 Nov 1977 (9), Pr<strong>an</strong>ce et al. 25336 (F, RB,<br />
dro, Gentry et al. 16181 (F, MO, NY); Pucallpa-Tingo U). RONDONIA: Mun. Presidente Medici, rd. Cuiaba-<br />
Maria rd., km 88, 13 Mar 1982 (st), Gentry et al. 36309 P6rto Velho, km 300, 25 Jun 1984 (d), Cid et al. 4798<br />
(BG).<br />
(BG); Mun. Ouro Preto, rd. Cuiaba-P6rto Velho, km<br />
BRAZIL. ACRE: Upper Rio Moa, Fazenda Arizona, 353, 39 Jun 1984 (6), Cid et al. 4919 (BG); Mun. Pi-<br />
10-15 Oct 1985 (st), Campbell et al. 6250 (BG); Abuna- menta Bueno, Guapor6, 7 Nov 1964 (st), Vieiro et al.<br />
Rio Br<strong>an</strong>co hwy., km 242-246, 18 Jul 1968 (6), Forero 985 (U). RORAIMA: Serra dos Surucucus, 19 Feb 1969<br />
et al. 6367 (C, INPA, MO, NY, S, UC, US); nr. mouth (2), Pr<strong>an</strong>ce et al. 10089 (G, INPA, NY, R, S, U, US),<br />
of Rio Macauh<strong>an</strong>, tributary of Rio Yaco, 4 Aug 1933 20 Feb 1969 (2), Pr<strong>an</strong>ce et al. 10124 (INPA, NY, S,<br />
(9), Krukoff 5282 (A, G, LE, M, MO, NY, S, U, UC, U, US); Rio Negro, left b<strong>an</strong>k of Par<strong>an</strong>a do Marara, 7<br />
US, type collection of P. subplicata); rd. Brasileia-As- Feb 1977 (9), M. R. S<strong>an</strong>tos 172 (MG, U).<br />
sis, km 16, 3 Nov 1980 (9), Lowrie et al. 710 (U); Sena BOLIVIA. BENI: Rio Beni, Cachuela Esper<strong>an</strong>za, 29<br />
Madureira, 3 Oct 1968 (9), Pr<strong>an</strong>ce et al. 7792 (GH, Nov 1922 (6), Meyer 403 (U). PANDO: Barrola, 17 km<br />
INPA, M, NY, P, R, S, U); Rio Jurua-Mirin, Aug 1901 from Cobija, 18 May 1977 (st), Meneces 613 (MG,<br />
(6), Ule 5718 (B, G, MG, type collection of P. aurea). MO); S<strong>an</strong> Fr<strong>an</strong>cisco, 50 km from Cobija to Puerto<br />
AMAPA: Confluence of Rio Oiapoque <strong><strong>an</strong>d</strong> Rio I<strong>an</strong>6, 27 Rico, 1 Jun 1977 (6), Meneces 647 (MO); rd. Cobija<br />
Aug 1960 (6), Irwin et al. 47874 (F, FHO, GH, MG, to Extrema, Villa Marieta, 24 Jun 1978 (2), Meneces<br />
NY, RB, US); Rio Oiapoque, nr. mouth of Rio Ya- 725 (INPA); 3 km above Abuna, 13 km 1968 (a), Pr<strong>an</strong>ce<br />
roupi, 24 Sep 1960 (6), Irwin et al. 48468 (M, MG, et al. 8378 (B, INPA, NY, S, U, UC, US); Rio Na-<br />
NY, P, RB, UC); Rio Oiapoque, nr. Cachoeria S<strong>an</strong>ta reuda, Nicolas Suarrez, SW of Cobija, 31 Jul 1982<br />
Maria, 21 Aug 1961 (6), Pires et al. 50419 (FHO, G, (5), Sperling et al. 6430 (BG).<br />
MG, NY, S, US); 40 min up Rio Falcino, 14 Nov 1961<br />
(8), Pires et al. 50941 (F, MO, NY); Serra do Navio, Vernacular names. Colombia. Vaupes: uva sil-<br />
26 Sep 1961 (Q), Pires et al. 51247 (GH, MG, NY, MO, vestre. Venezuela. Bolivar: majagua, cay-wari-<br />
U, US); AMAZONAS: M<strong>an</strong>aus-Itacoatiara rd., Reserva<br />
Florestal Ducke, 6 Aug 1976 (st), Aluisio (INPA) 70809<br />
cay-yek, coiwaricoi-yek, kai-wa-rei-kei-yek. Su-<br />
(INPA); Uaup6s, Serra Sao Gabriel, 14 Feb 1959<br />
rinam.<br />
(9), Bospapaja, gr<strong>an</strong>boesipapaja. French<br />
Cavalc<strong>an</strong>te 606 (MG); Rio Uatumi, Itapir<strong>an</strong>ga, 24 Aug Gui<strong>an</strong>a. Bois c<strong>an</strong>on, bois c<strong>an</strong>on male or male<br />
1979 (6), Cid et al. 721 (INPA, U); Rio Jurua, Sta. bois c<strong>an</strong>on, bouchi papaie <strong><strong>an</strong>d</strong> papaye (Para-<br />
Rosa, 15 Aug 1975 (st), D. Coelho et al. (INPA) 51350 maka), kulumasi (Wayapi). Peru. Amazonas:<br />
(INPA), 22 Aug 1975 (st), D. Coelho et al. (INPA) sacha<br />
52381 (INPA); M<strong>an</strong>aus-Itacoatiara rd., Reserva Flo- uvila, shuvija (Aguaruna); Hu<strong>an</strong>uco <strong><strong>an</strong>d</strong><br />
restal Ducke, 28 Sep 1962 (8), Duarte 7194 (INPA, M);<br />
S<strong>an</strong> Martin: uvilla or uvilla bl<strong>an</strong>ca; Loreto: uvilla<br />
M<strong>an</strong>icor6, nr. S<strong>an</strong>ta F6, 8-11 Sep 1934 (9), Krukoff l<strong>an</strong>uda. Brazil. Amapa: mapatir<strong>an</strong>a; Amzonas:<br />
6041 (A, B, BR, F, LE, MO, NY, S, U, US); Humaita, imbaibar<strong>an</strong>a, purumai, tourem; Mato Grosso:<br />
plateau between Rio Livramento <strong><strong>an</strong>d</strong> Rio Ipixuna, 7-<br />
18 Nov 1934 (9), Krukoff 7071<br />
caramuri, imbaubar<strong>an</strong>a, torena. Bolivia. P<strong><strong>an</strong>d</strong>o:<br />
(A, BR, G, GB, LE,<br />
MO, NY, RB, S, S, U, US, type collection of P. um- ambaibochi, ambaibillo.<br />
bellata); Sao Paulo de Olivenca, nr. Palmares, 11 Sep- The species is rather variable in the dimen-<br />
26 Oct 1936 (9), Krukoff 8073 (A, BR, G, LE, MO, sions <strong><strong>an</strong>d</strong> shape of the leaves. The indument<br />
NY, P, S, U, US, type collection ofP. isophlebia); upper mostly consists of appressed hairs but patent hairs<br />
Rio Negro Uaupes, 12 Feb 1959 (9), Rodrigues 884<br />
(U); Tefe, 24 Nov 1959<br />
occur in the<br />
(st), Rodrigues et al. 1424<br />
Upper Amazon Basin, e.g., in the<br />
(INPA); M<strong>an</strong>aus-Itacoatiora rd., Reserva Florestal type collection ofP. aurea. The number of lateral<br />
Ducke, 19 Oct 1965 (6), Rodrigues 7532 (INPA); Tefe, veins varies from 6 to ca. 25 pairs, but in some<br />
23 Aug 1979 (6), Rodrigues et al. 10182 (INPA); Ma- collections (of juvenile material?) probably benaus-P6rto<br />
Velho rd., between Rio Cast<strong>an</strong>ho <strong><strong>an</strong>d</strong> Rio<br />
Tapuna, 18 Jul 1972 (8), M. F. Silva et al. 903<br />
longing to P. minor, e.g., Ayala 341 <strong><strong>an</strong>d</strong> Gentry<br />
(INPA). et<br />
MATO<br />
al.<br />
GROSSO: Aripu<strong>an</strong>a, 18 Jun 1975 (8), Cordeiro 22381, the number of lateral veins may rise<br />
140 (US), 25 J<strong>an</strong>-14 May 1977 (st), Gomes et al. 547, to 40 pairs.<br />
593, 632, 635, 646, 655, 673, 690, 759, 789, 810, 813, The species is a clear-cut one, characterized<br />
838, 872, 891, 924, 926, 931, 933, 955, 1033, 1118, by the subumbellate pistillate inflorescence <strong><strong>an</strong>d</strong><br />
1120, 1129, 1156, 1175, 1255, 1319, 1336, 1338, 1356, the knob-like<br />
1378, 1419, 1423, 1425,<br />
stigma. It shows resembl<strong>an</strong>ces of<br />
1444, 1446, 1534, 1568, 1570,<br />
1577, 1584, 1593, 1594, 1595, 1636, 1640, 1641, 1692,<br />
P. bolivarensis (see p. 186).<br />
1731, 1745, 1769, 1822, 1823, 1867, 1893, 1917, 1983 The fruits are eaten by the monkey Cebus apel-<br />
(INPA), 14 Jul 1977 (6), Gomes et al. 2170 (INPA), 19 la (Foster et al. 5019).<br />
J<strong>an</strong> 1979 (9), 29 May 1976 (6), Monteiro et al. 1119 From the three syntype collections from French<br />
(INPA), M. G. Silva et al. 4326 (MG), 1 J<strong>an</strong> 1979 (6),<br />
M. G. Silva et al. 4738 (MG), 8 Oct 1979<br />
Gui<strong>an</strong>a<br />
(9), Ryl<strong><strong>an</strong>d</strong>s<br />
(Benoist 960, Benoist 978, <strong><strong>an</strong>d</strong> Wach-<br />
25 (INPA); Mun. Vila Bela da S<strong>an</strong>tissima Trinidade, enheim 18) Benoist 960 has been selected as the<br />
4 km S of border <strong>with</strong> Rond6nia, 3 Nov 1985 (2), lectotype collections (Berg, 1975).
190 Flora Neotropica<br />
<strong>Pourouma</strong> napoensis <strong><strong>an</strong>d</strong> P. herrerensis have<br />
been included in this revision after submission<br />
of the (pre-review) m<strong>an</strong>uscript <strong><strong>an</strong>d</strong> are, therefore,<br />
placed at the end of the systematic treatment,<br />
<strong><strong>an</strong>d</strong> not after P. hirsutipetiolata <strong><strong>an</strong>d</strong> P. mollis,<br />
respectively.<br />
24. <strong>Pourouma</strong> napoensis C. C. Berg, Brittonia<br />
42: 59-65. 1990. Type. Ecuador. Napo: Re-<br />
serva Biologia Jatun Sacha, 14 Aug 1987 (6),<br />
Palacios 1874 (holotype, QCNE; isotypes,<br />
AAU, BG, MO, NY, QAME). Fig. 92.<br />
Tree, up to 25 m tall. Leafy twigs 1-2.5 cm<br />
thick, yellow-hirsute, pluricellular hairs absent.<br />
Lamina 7-fid to -parted, ca. 25-45 x 25-45 cm,<br />
coriaceous, apex acuminate, base deeply cordate<br />
<strong>with</strong> overlapping lobes; upper surface smooth to<br />
slightly scabrous, yellow-hirsute to -hirtellous on<br />
the whole surface or only on the main veins,<br />
lower surface yellow-hirsute to -hirtellous on the<br />
veins, arachnoid hairs in the areoles <strong><strong>an</strong>d</strong> on the<br />
reticulum; lateral veins 16-25 pairs; petiole ca.<br />
25-40 cm long, yellow-hirsute; stipules 10-16 cm<br />
long, outside yellow-hirsute, inside glabrous, caducous.<br />
Staminate inflorescences ca. 12-22 cm<br />
long, ca. 8-14 cm wide; peduncle 4-8 cm long,<br />
peduncle <strong><strong>an</strong>d</strong> br<strong>an</strong>ches yellow-hirtellous to -pu-<br />
In the shape, dimensions, <strong><strong>an</strong>d</strong> indument of the<br />
lamina this species resembles P. oraria. It differs<br />
from the latter in the staminate flowers <strong>with</strong> their<br />
connate tepals <strong><strong>an</strong>d</strong> fully connate filaments. These<br />
features suggest a close relationship to P. hirsu-<br />
tipetiolata.<br />
25. <strong>Pourouma</strong> herrerensis C. C. Berg, C<strong><strong>an</strong>d</strong>ollea<br />
44(2): 513. 1989. Type, Peru, Loreto, Prov.<br />
Requena, Jenaro Herrera, Reserva Forestal,<br />
tree 4/122, 2 Oct 1985 (d), Spichiger et al. 1995<br />
(holotype, G; isotype, BG). Fig. 93.<br />
Tree, up to 25 m tall. Leafy twigs 3-6 mm<br />
thick, minutely puberulous to hispidulous (<strong><strong>an</strong>d</strong><br />
+ scabrous) or also <strong>with</strong> much longer, yellow<br />
hairs, pluricellular hairs dark brown <strong><strong>an</strong>d</strong> rather<br />
sparse. Lamina entire, oblong to elliptic to ovate,<br />
8-20 x 3-11.5 cm, (sub)coriaceous, apex shortacuminate,<br />
base acute to rounded; upper surface<br />
sparsely appressed-puberulous on the midrib or<br />
the whole surface covered <strong>with</strong> long, yellow hairs,<br />
lower surface appressed-puberulous on the main<br />
veins or also <strong>with</strong> long yellow hairs, arachnoid<br />
hairs in the areoles <strong><strong>an</strong>d</strong> on the reticulum; lateral<br />
veins 9-14 pairs, basal pair unbr<strong>an</strong>ched or<br />
br<strong>an</strong>ched, the basal part of these veins forming<br />
the basal part of the leaf margin (thus not separated<br />
from the margin by mesophyll), tertiary<br />
venation slightly prominent; petiole 3-10 cm<br />
berulous; flowers (sub)sessile, in<br />
long, minutely puberulous or also <strong>with</strong> long, yelglobose<br />
heads, low<br />
these 7-12 mm diam.; peri<strong>an</strong>th, tubular, 1-2 mm<br />
hairs; stipules 3.5-8 cm long, caducous, outside<br />
high, puberulous at the apex; stamens 3-4, filadensely<br />
puberulous <strong><strong>an</strong>d</strong> <strong>with</strong> brown, pluricellular<br />
ments completley connate, 4-6 mm long. Pistilhairs,<br />
or also <strong>with</strong> long, yellow hairs,<br />
sometimes<br />
late inflorescences up to 10 cm long <strong><strong>an</strong>d</strong> up to<br />
only a few appressed ones, inside<br />
12 cm wide, br<strong>an</strong>ched; peduncle 3-5 cm sparsely short-pubescent. Staminate infloreslong,<br />
cences<br />
peduncle <strong><strong>an</strong>d</strong> br<strong>an</strong>ches yellow-hirsute to -hirtel- up to 10 cm long <strong><strong>an</strong>d</strong> up to 9 cm wide;<br />
lous; flowers ca. 40-70; pedicel 0.3-0.5 cm, in<br />
peduncle 2-6 cm long, peduncle <strong><strong>an</strong>d</strong> br<strong>an</strong>ches<br />
fruit up to 1.2 cm long; peri<strong>an</strong>th ca. 5 mm densely puberulous to short-velutinous or also<br />
long, <strong>with</strong><br />
yellow-hirsute to -hirtellous, sparsely so in the long, yellow hairs; flowers sessile or pedicellate,<br />
most of them in globose heads, numerous<br />
uppermost part; stigma peltate.<br />
<strong><strong>an</strong>d</strong> 3-4 mm diam. or less<br />
Distribution (Fig. 81). Ecuador (Napo), in non-<br />
(5-12) <strong><strong>an</strong>d</strong> 5-8 mm<br />
inundated forest, at low altitudes.<br />
diam.; tepals ca. 1 mm long, connate, forming a<br />
+ urceolate peri<strong>an</strong>th, densely hairy; filaments<br />
Specimens examined. ECUADOR. NAPO: Reserva exceeding the tepals, free. Pistillate inflorescences<br />
Biologica Jatun Sacha, Rio Napo, 8 km below Puerto up to 14 cm long <strong><strong>an</strong>d</strong> 9 cm wide; peduncle <strong><strong>an</strong>d</strong><br />
Misahualli, 4 Sep 1987 (2), Cer6n M. 2019 (BG); 8 km br<strong>an</strong>ches short-velutinous; flowers 7-25, in 1-4<br />
SE of Tena, 30 Aug 1960 (2), Grubb et al. 1545 (K,<br />
NY); Reserva Biologica Jatun Sacha, Rio Napo, 8 km<br />
+ distinct clusters, pedicels up to 0.7 cm long;<br />
below Puerto Misahualli, 2 Oct 1986 (2), Palacios 1337 peri<strong>an</strong>th yellow-velutinous, except for the apex;<br />
(BG), 14 Aug 1987 (a), Palacios 1874 (BG, type col- stigma subpeltate.<br />
lection of P. napoensis); Lumbaqui, 12 May 1987 (a), Distribution (Fig. 81). Peru (Loreto), in non-<br />
Pennington et al. 12244 (BG, K).<br />
inundated forest.<br />
Specimens examined. PERU. LORETO: Prov. Requena,<br />
Jenaro Herrera, Aug-Sep 1976 (6), Bernardi s. n.
<strong>Pourouma</strong> 191<br />
bi<br />
f<br />
In<br />
i<br />
FIG. 92. <strong>Pourouma</strong> napoensis. a. Leaf(Grubb et a. 1545). b. Stipules <strong><strong>an</strong>d</strong> staminate inflorescence (Pennington<br />
et al. 12244). c. Pistillate inflorescence (Grubb et al. 1545). d. Staminate flower (Pennington et al 12244). e.<br />
Pistillate flower (Ceron M. et al 2019).
.At<br />
FIG. 93. <strong>Pourouma</strong> herrerensis. a. Leafy twig <strong>with</strong> staminate inflorescences (Spichiger et al. 1995). b. Pistillate<br />
inflorescence (Spichiger et al. 1999). c. Leaf (Vasquez et al. 2623).
<strong>Pourouma</strong><br />
(BG), 9 Dec 1977 (d), Gentry et al. 21346 (U); Prov.<br />
Maynas, Puerto Almendras, 11 Nov 1984 (9), Maas et<br />
al. 6267 (BG, U); Prov. Requena, Jenaro Herrera, Reserva<br />
Forestal, 2 Oct 1985 (d), Spichiger et al. 1995<br />
(BG, G, type collection), 17 Sep 1982 (6), Spichiger et<br />
al. 1996 (BG, G), 9 Oct 1982 (st), Spichiger et al. 1997<br />
(BG, G), 14 Sep 1982 (d), Spichiger et al. 1998 (BG,<br />
G), 10 Nov 1982 (9), Spichiger et al. 1999 (BG, G);<br />
Prov. Maynas, Puerto Almendras, 17 Apr 1980 (6),<br />
Vdsquez et al. 167 (BG), 21 Oct 1981 (9), Vdsquez et<br />
al. 2623 (BG); Prov. Maynas, Rio N<strong>an</strong>ay, Mish<strong>an</strong>a, 24<br />
Apr 1986 (9), Vdsquez et al. 7535 (BG).<br />
This species has a str<strong>an</strong>ge mixture of features.<br />
The staminate inflorescences <strong><strong>an</strong>d</strong> flowers resemble<br />
those of P. mollis, the pistillate inflorescences<br />
<strong><strong>an</strong>d</strong> flowers those of P. mollis subsp. triloba <strong><strong>an</strong>d</strong><br />
P. acuminata. In the indument it differs from<br />
both. One of the forms of this species has very<br />
short <strong><strong>an</strong>d</strong> stiff hairs, resulting in a slightly scabrous<br />
surface of the leafy twigs, petioles, stipules,<br />
<strong><strong>an</strong>d</strong> main veins of the lamina beneath. The other<br />
form has this short indument mixed <strong>with</strong> much<br />
longer yellow hairs, as in P. cucura, <strong><strong>an</strong>d</strong> bears,<br />
moreover, long yellow hairs on the upper leaf<br />
surface, like some forms of P. cucura. Some of<br />
the specimens have oblong to elliptic leaves <strong>with</strong><br />
Unnamed Collections<br />
1. PERU. AMAZONAS: Serr<strong>an</strong>ia de Bagua, ca. 12-<br />
18 km on trail E of La Peca, 1800-1950 m, lower<br />
mont<strong>an</strong>e cloud forest, 14 Jun 1978 (6), Gentry et al.<br />
22866 (U). SAN MARTIN: Prov. Rioja, rd. Pedro Ruiz-<br />
Moyobamba, km 390, Venceremos, 2100 m, 7-9 Aug<br />
1983 (st), D. N. Smith et al. 4742 (MO).<br />
The indument of the leafy twigs, stipules, petioles,<br />
<strong><strong>an</strong>d</strong> lower surface of the lamina largely<br />
matches that of P. bicolor subsp. bicolor. The<br />
specimens differ from that taxon in the smooth<br />
<strong><strong>an</strong>d</strong> puberulous upper surface of the lamina, the<br />
broadly ovate lamina, the slender (ca. 15 x 10<br />
cm) staminate inflorescence, the oblong to elliptic<br />
193<br />
tepals of the staminteflower, <strong><strong>an</strong>d</strong> the sparse pluricellular<br />
hairs.<br />
These specimens may represent a distinct cloud<br />
forest species (see p. 114).<br />
2. PERU. LORETO: Rio Corrientes, between Tiente<br />
Lopez <strong><strong>an</strong>d</strong> Puesto Av<strong>an</strong>zado, 4 Apr 1977 (st), Gentry<br />
et al. 19048 (BG); S<strong>an</strong>ta Rosa, 1-5 Sep 1929 (st), Killip<br />
& A. C. Smith 28800 (F, NY).<br />
BRAZIL. ACRE: Reserva INCRA S<strong>an</strong>ta Luzia, BR-<br />
364, km 40, 5-19 Oct 1984 (st), Campbell et al. 6820<br />
(NY) <strong><strong>an</strong>d</strong> 7689 (BG).<br />
These have ovate to elliptic leaves (16-32 x<br />
12-22 cm) <strong>with</strong> a deeply cordate base. The leafy<br />
twigs, petioles, stipules, <strong><strong>an</strong>d</strong> main veins at the<br />
lower surface of the lamina have dense, long,<br />
patent, yellow hairs. The stipules are glabrous<br />
inside. The presence of arachnoid hairs on the<br />
petiole, main veins of the lamina beneath, <strong><strong>an</strong>d</strong><br />
stipules suggest affinities to P. tomentosa. These<br />
collections may represent a new subspecies of<br />
this species.<br />
3. VENEZUELA. AMAZONAS: Nr. Yavita, 21 Apr<br />
1970 (st), Steyermark et al. 102874 (BG, MO, NY,<br />
VEN); Dept. Rio Negro, Cerro de la Neblina, Rio Mathe<br />
basal pair of lateral veins unbr<strong>an</strong>ched; such warinuma, 8-10 J<strong>an</strong> 1984 (st), Steyermarket al. 129773<br />
specimens bear some resembl<strong>an</strong>ce to P. acumi- (MO).<br />
nata. Other specimens (<strong>with</strong> long hairs) have These collections have thickly coriaceous<br />
ovate leaves <strong><strong>an</strong>d</strong> the basal pairs of lateral veins leaves, thick leafy twigs <strong>with</strong> conspicuous lentibr<strong>an</strong>ched.<br />
However, both types of leaves may cels, <strong><strong>an</strong>d</strong> dense, almost black, pluricellular hairs<br />
occur on the same pl<strong>an</strong>t, or even on the same on the leafy twigs <strong><strong>an</strong>d</strong> stipules. In these features<br />
br<strong>an</strong>ch.<br />
they match P. ferruginea, but the characteristic,<br />
The material <strong>with</strong> a scabrous upper leaf sur- dense (white to brownish) villous-arachnoid inface,<br />
provisionally kept in P. mollis subsp. tri- dument on the lower surface of the lamina is<br />
loba, resembles P. herrerensis in the variation of w<strong>an</strong>ting <strong><strong>an</strong>d</strong> the stipules are densely hairy inside.<br />
the leaves.<br />
These collections might represent <strong>an</strong> abber<strong>an</strong>t<br />
form of P. ferruginea or perhaps of P. melinonii<br />
subsp. melinonii.<br />
4. COLOMBIA. VAUPES: Alto Vaupes, 25 Nov 1975<br />
(9), Roa 251 (INPA).<br />
This collection has leafy twigs <strong>with</strong> yellow-<br />
hirsute indument <strong><strong>an</strong>d</strong> conspicuous lenticels. The<br />
indument of the leafy twigs <strong><strong>an</strong>d</strong> the leaf surface<br />
resembles that of P. cucura, but the thickly co-<br />
riaceous, somewhat plicate lamina has the hairs<br />
on its upper surface confined to the main veins.<br />
Moreover, the stipules are hairy inside <strong><strong>an</strong>d</strong> the<br />
reticulum on the lamina beneath is very prom-<br />
inent. The fruiting peri<strong>an</strong>th is scabrous. This<br />
specimens has a mixture of characters of P. cu-<br />
cura <strong><strong>an</strong>d</strong> P. bicolor subsp. bicolor.
194<br />
5. ECUADOR. NAPO: Aii<strong>an</strong>gu, 1-15 Feb 1986 (st),<br />
Korning & Thomson 47801 <strong><strong>an</strong>d</strong> 47807 (BG).<br />
The latter has most characters in common <strong>with</strong><br />
the form of P. bicolor subsp. bicolor <strong>with</strong> <strong>an</strong> entire<br />
lamina, being smooth above. However, the<br />
indument of the leafy twigs, petiole, <strong><strong>an</strong>d</strong> lower<br />
leaf surface resembles that of P. cucura. The former<br />
collection <strong>with</strong> 5-parted leaves <strong><strong>an</strong>d</strong> incisions<br />
down to the petiole may represent the juvenile<br />
form of the latter collection. The mixture of characters<br />
suggests possible hybridization.<br />
Names Excluded<br />
<strong>Pourouma</strong> paraensis-nomen (in schedule) given<br />
by Huber; refers to Goeldi (MG) 7732 = P.<br />
melinonii subsp. melinonii.<br />
<strong>Pourouma</strong> retusa Bentham-nomen, see Miquel<br />
in Martius, Fl. bras. 4(1): 128. 1853; refers to<br />
Spruce s. n. from Brazil, Amazonas, M<strong>an</strong>aussyntype<br />
collection of P. tomentosa.<br />
<strong>Pourouma</strong> tri<strong>an</strong>ae A. Richter-nomen (in schedula);<br />
refers to Tri<strong>an</strong>a 862 = P. cecropiifolia.<br />
<strong>Pourouma</strong> ulei Warburg ex Ule-nomen; see Ule,<br />
Bot. Jahrb. Syst. 40: 132. 1907; refers to Ule<br />
9314 = P. cecropiifolia.<br />
ACKNOWLEDGMENTS<br />
The authors of the <strong>Coussapoa</strong> revision (C.C.B.<br />
<strong><strong>an</strong>d</strong> R.W.A.P.A.) are much indebted to Dr. N.<br />
G. Bisset (Chelsea College, London) for correct-<br />
ing the English text. They th<strong>an</strong>k Mr. Bonsen (In-<br />
stitute for Systematic Bot<strong>an</strong>y, Utrecht) for con-<br />
tributing the chapter on the wood <strong><strong>an</strong>d</strong> leaf<br />
<strong>an</strong>atomy, <strong><strong>an</strong>d</strong> Drs. E. C. H. v<strong>an</strong> Heusden (In-<br />
stitute for Systematic Bot<strong>an</strong>y, Utrecht) for pre-<br />
paring the distribution maps. They are grateful<br />
to the curators of the cited herbaria for putting<br />
their collections at the disposal of the authors.<br />
Gr<strong>an</strong>ts from the Netherl<strong><strong>an</strong>d</strong>s Foundation for Ad-<br />
v<strong>an</strong>cement of Tropical Research (WOTRO) to<br />
C.C.B. enabled part of the collections <strong><strong>an</strong>d</strong> data<br />
on which the treatment is based to the gathered<br />
during several South Americ<strong>an</strong> trips. They also<br />
th<strong>an</strong>k Dr. L. B. Holm-Nielsen (Bot<strong>an</strong>ical Insti-<br />
tute, Aarhus) for his help during fieldwork in<br />
Ecuador. The drawings were prepared by Ms. Siri<br />
Herl<strong><strong>an</strong>d</strong> (Bergen), Miss E. M. Hupkens v<strong>an</strong> der<br />
Elst (Utrecht), <strong><strong>an</strong>d</strong> Mr. T. Schipper (Utrecht).<br />
The revision of <strong>Pourouma</strong> was started by Drs.<br />
Flora Neotropica<br />
E. Kieft (Utrecht), <strong><strong>an</strong>d</strong> some elements of that<br />
unfinished study have been used by the present<br />
authors (C.C.B. <strong><strong>an</strong>d</strong> E.C.H.vH.). The senior au-<br />
thor is indebted to the University of Bergen, Fac-<br />
ulty of Mathematics <strong><strong>an</strong>d</strong> Natural Sciences, for<br />
covering traveling expenses, <strong><strong>an</strong>d</strong> to the Bot<strong>an</strong>ical<br />
Institute at Bergen for their assist<strong>an</strong>ce. The il-<br />
lustrations were prepared by Ms. Siri Herl<strong><strong>an</strong>d</strong><br />
(Bergen), Mr. H. Rypkema (Utrecht), <strong><strong>an</strong>d</strong> Mr. T.<br />
Schipper (Utrecht). The authors are indebted to<br />
Ms. Kathleen Welling for processing part of the<br />
text, to Drs. W. H. A. Hekking (Utrecht), Drs.<br />
E. Simonis (Utrecht), Ms. Edit Fjaere (Bergen)<br />
<strong><strong>an</strong>d</strong> Ms. Fr<strong>an</strong>cina Berg (Bergen) for their assis-<br />
t<strong>an</strong>ce in preparing the list of exsiccatae, <strong><strong>an</strong>d</strong> to<br />
Drs. E. Simonis for his contributions to the preparation<br />
of the lists of specimens examined <strong><strong>an</strong>d</strong><br />
the distribution maps.<br />
LITERATURE CITED<br />
Akkerm<strong>an</strong>s, R. W. A. P. & C. C. Berg. 1982. New<br />
species <strong><strong>an</strong>d</strong> combinations in <strong>Coussapoa</strong> (Cecro-<br />
piaceae), <strong><strong>an</strong>d</strong> keys to its species. Proc. Kon. Ned.<br />
Akad. Wetensch., Ser. C, 85(4): 441-471.<br />
Aublet, J. B. C. F. 1775. Histoire des pl<strong>an</strong>tes de<br />
Gui<strong>an</strong>e fr<strong>an</strong>caise 2. Paris.<br />
Benoist, R. 1922. Descriptions d'especes nouvelles<br />
du genre <strong>Pourouma</strong> (Moracees). Bull. Mus. Nat.<br />
Hist. Nat. (Paris) 28: 318-321.<br />
. 1924. Descriptions d'especes nouvelles de<br />
Ph<strong>an</strong>6rogames de la Guy<strong>an</strong>e fr<strong>an</strong>caise. Bull. Mus.<br />
Nat. Hist. Nat. (Paris) 30: 103-105.<br />
. 1933. Les bois de la Guy<strong>an</strong>e fr<strong>an</strong>qaise. Moracees.<br />
Arch. Bot. Mem. 5(1): 29-49.<br />
Benson, W. W. 1985. Amazoni<strong>an</strong> <strong>an</strong>t-pl<strong>an</strong>ts. In G.<br />
T. Pr<strong>an</strong>ce & T. E. Lovejoy, Key environments:<br />
Amazonia. Chapter 13. Pergamon Press. Oxford-<br />
New York-Toronto-Sydney-Fr<strong>an</strong>kfurt.<br />
Berg, C. C. 1975. <strong>Coussapoa</strong>. In J. L<strong>an</strong>jouw & A. L.<br />
Stoffers, Flora of Suriname 5(1): 279-286. Leiden.<br />
1977a. Abscission of <strong>an</strong>thers in Cecropia<br />
Loefl. Acta Bot. Neerl. 26: 417-419.<br />
. 1977b. Urticales, their differentiation <strong><strong>an</strong>d</strong><br />
systematic position. In K. Kubitzki (ed.), Flowering<br />
pl<strong>an</strong>ts, evolution <strong><strong>an</strong>d</strong> classification of higher<br />
categories. P1. Syst. Evol. Suppl. 1: 349-374.<br />
. 1978a. <strong>Cecropiaceae</strong>, a new family of the<br />
Urticales. Taxon 27(1): 39-44.<br />
. 1978b. New Amazoni<strong>an</strong> species. <strong>Coussapoa</strong><br />
pr<strong>an</strong>cei <strong><strong>an</strong>d</strong> Perebea mennegae (Moraceae). Acta<br />
Bot. Neerl. 27: 11-15.<br />
. 1983a. A new species of <strong>Coussapoa</strong> (<strong>Cecropiaceae</strong>)<br />
from Peru. Proc. Kon. Ned. Akad. Wetensch.,<br />
Ser. C, 86(3): 305-307.<br />
. 1983b. Dispersal <strong><strong>an</strong>d</strong> distribution in Urticales-An<br />
outline. In K. Kubitzki (ed.), Dispersal
Literature Cited 195<br />
<strong><strong>an</strong>d</strong> distribution. Sonderb. Naturwiss. Vereins liens. Notizbl. Bot. Gart. Berlin-Dahlem 11: 579-<br />
Hamburg 7: 219-229.<br />
591.<br />
. 1988. The genera Trophis <strong><strong>an</strong>d</strong> Streblus (Mo- . 1932b. Especes nouvelles de pl<strong>an</strong>tes de<br />
raceae) remodelled. Proc. Kon. Ned. Akad. We- l'Amazonie Bresilienne. Bull. Mus. Hist. Nat.<br />
tensch., Ser. C, 91(4): 345-362.<br />
(Paris), Ser. 2, 4: 720-749.<br />
. 1989. <strong>Pourouma</strong> herrerensis C. C. Berg, a . 1947. New forest trees <strong><strong>an</strong>d</strong> climbers of the<br />
new species of<strong>Cecropiaceae</strong> from Amazoni<strong>an</strong> Peru. Amazon. Trop. Woods 90: 7-30.<br />
Contribution to the study of the flora <strong><strong>an</strong>d</strong> vege- Engler,G.H.A. 1889. InG.H. A.Engler&K. Pr<strong>an</strong>tl,<br />
tation of Peruvi<strong>an</strong> Amazonia. XV. C<strong><strong>an</strong>d</strong>ollea 44(2): Nattirliche Pfl<strong>an</strong>zenfamilien 3(1): Moraceae: 66-<br />
513-516.<br />
98. Leipzig.<br />
1990. New species of <strong>Coussapoa</strong> <strong><strong>an</strong>d</strong> Pourou- FalcAo, M. de Aguiar & E. Lleras. 1980. Aspectos<br />
ma (<strong>Cecropiaceae</strong>). Brittonia 42: 59-65.<br />
fenologicos, ecol6gicos e de produtividade do<br />
1989c. Systematics <strong><strong>an</strong>d</strong> phylogeny of Urti- Mapati (<strong>Pourouma</strong> cecropiifolia Mart.). Acta<br />
cales. Pages 193-220 in S. Blackmore & P. R. Amazonica 10(4): 711-724.<br />
Cr<strong>an</strong>e (eds.), Evolution, systematics <strong><strong>an</strong>d</strong> fossil his- Hawkes, A. H. 1948. Notes on the Moraceae-I.<br />
tory of Hamamelidae. Vol. 2. 'Higher' Hama- Phytologia 3: 29-32.<br />
melidae. Oxford University Press, Oxford. Killip, E. P. 1937. <strong>Coussapoa</strong> gr<strong><strong>an</strong>d</strong>iceps. In J. F.<br />
& E. C. H. v<strong>an</strong> Heusden. 1988. Studies on Macbride, Flora of Peru: Moraceae. Publ. Field.<br />
the flora of the Gui<strong>an</strong>as 36. New taxa <strong><strong>an</strong>d</strong> com- Mus. Nat. Hist., Bot. Ser., 13(2.2): 296.<br />
binations in <strong>Pourouma</strong> (<strong>Cecropiaceae</strong>). Proc. Kon. Klotzsch, J. F. 1847. Beitrage zu einer Flora de Ae-<br />
Ned. Akad. Wetensch., Ser. C, 91(2): 105-116. quinoctial-Gegenden: Artocarpeae. Linnaea 20:<br />
& F. Kooy. 1982. Three new species of Pou- 523-525.<br />
rouma (<strong>Cecropiaceae</strong>) of the Gui<strong>an</strong>a Region. Brit- Liebm<strong>an</strong>n, F.M. 1851. MexicosogCentral-Americas<br />
tonia 34(1): 36-41.<br />
neldeagtige pl<strong>an</strong>ter (Ordo: Urticaceae) indbeffat-<br />
Blake, S. F. 1919. New South Americ<strong>an</strong> spermato- tende familierne: Urticeae, Moreae, Artocarpeae<br />
phytes collected by B. M. Curr<strong>an</strong>. Contr. U.S. Natl. og Ulmaceae. Kongel. D<strong>an</strong>ske Vidensk. Selsk. Skr.<br />
Herb. 20: 237-245.<br />
Ser. 5 (Naturvidensk. Math. Afd.), 2: 285-343.<br />
Bonsen, K. & B. J. H. ter Welle. 1983. Comparative Little, E. L. 1969. New tree species from Esmeraldas,<br />
wood <strong><strong>an</strong>d</strong> leaf <strong>an</strong>atomy of the <strong>Cecropiaceae</strong> (Urti- Ecuador. Phytologia 18: 195-208.<br />
cales). Bull. Mus. Nat. Hist. Nat. (Paris) Ser. 4, Macbride, J. F. 1930. Peruvi<strong>an</strong> Spermatophytes. 4.<br />
5(sect. B, Ad<strong>an</strong>sonia 2): 151-177.<br />
Other Peruvi<strong>an</strong> pl<strong>an</strong>ts, chiefly new species. Publ.<br />
Burger, W. C. 1973. Notes on the flora of Costa Rica, Field Mus. Nat. Hist., Bot. Ser., 8(2): 113-130.<br />
3: New species in the Moraceae. Phytologia 26:<br />
1937. Flora of Peru: Moraceae. Publ. Field<br />
421-434.<br />
Mus. Nat. Hist., Bot. Ser., 13(2.2): 274-331.<br />
1977. Flora Costaricensis: <strong>Coussapoa</strong>. Field- Martius, C. F. P. 1843. Systema materiae medicae<br />
i<strong>an</strong>a Bot. 40: 131-137; <strong>Pourouma</strong> 40: 200-203. vegetabilis Brasiliensis. Leipzig.<br />
Chew Wee-Lek. 1963. A revision of the genus Poi- Mildbraed, J. 1927. Pl<strong>an</strong>tae Tessm<strong>an</strong>ni<strong>an</strong>ae Perukilospermum.<br />
Gard. Bull. Singapore 20: 1-104. vi<strong>an</strong>ae VI. Moraceae. Nitzbl. Bot. Gard. Berlin-<br />
Corner, E. J. H. 1962. The classification of Mora- Dahlem 10: 183-194.<br />
ceae. Gard. Bull. Singapore 19: 187-252.<br />
. 1928. Neue Arten von <strong>Coussapoa</strong> und Pou-<br />
Croat, T. B. 1978. Flora of Barro Colorado Isl<strong><strong>an</strong>d</strong>. rouma. Aubl. Notizbl. Bot. Gart. Berlin-Dahlem<br />
St<strong>an</strong>ford, California.<br />
10:413-420.<br />
Cronquist, A. 1981. An integrated system of classi- .1938. Moraceae. In L. Diels, Neue Arten aus<br />
fication of flowering pl<strong>an</strong>ts. Columbia University Ecuador. Notizbl. Bot. Gart. Berlin-Dahlem 14:<br />
Press, New York.<br />
29.<br />
Cuatrecasas, J. 1951. In J. A. Steyermark, Bot<strong>an</strong>ical .1942. Moraceae II. In L. Diels, Neue Arten<br />
exploration of Venezuela--I: Moraceae. Fieldi<strong>an</strong>a aus Ecuador. Notizbl. Bot. Gart. Berlin-Dahlem<br />
Bot. 28(1): 210-216.<br />
15: 783-785.<br />
.1954. Dos Moraceas y dos Compuestas nue- Miquel, F. A. W. 1843. Observations bot<strong>an</strong>icae de<br />
vas de Venezuela. Bol. Soc. Venez. Ci. Nat. 15: quibusdam pl<strong>an</strong>tis, quas in colonia Surinamensi<br />
107-111.<br />
legit vir. graviss. H. C. Focke. Het Instituut 2(1842):<br />
. 956a. Notas a la Flora de Colombia, XIV. 185-205, t. 3. 1843.<br />
Revista Acad. Colomb. Ci. Exact. 9(36/37): 325- . 1853. Urticales. In C. F. P. von Martius,<br />
341, <strong>with</strong> photographs.<br />
Flora Brasiliensis 4(1): <strong>Coussapoa</strong>: 131-139, t. 42-<br />
.1956b. Morfceas nuevas de Colombia. Cal- 45. <strong>Pourouma</strong>: 122-131, t. 36-41.<br />
dasia 7: 287-304.<br />
O'Dowd, D. J. 1982. Pearl bodies as <strong>an</strong>t food: An<br />
1956c. Notas a la Flora de Venezuela. Bol. ecological role of some leaf emergences of tropical<br />
Soc. Venez. Ci. Nat. 17: 80-97.<br />
pl<strong>an</strong>ts. Biotropica 14(1): 40-49.<br />
. 1967. Moraceae. In J. A. Steyermark, Flora Pires, J. Murqa. 1973. Tipos de vegetaco da Amadel<br />
Auy<strong>an</strong>-tepui. Acta Bot. Venez. 2(5-8): 202- z6nia. O Museu Goeldi no Ano do Sesquicentena-<br />
205.<br />
rio. Belem.<br />
Ducke, A. 1932a. Neue Arten aus der Hylea Brasi- Pittier, H. 1917. New <strong><strong>an</strong>d</strong> noteworthy pl<strong>an</strong>ts from
196 Flora Neotropica<br />
Colombia <strong><strong>an</strong>d</strong> Central America-6. Contr. U.S.<br />
Natl. Herb. 18: 225-259.<br />
.1943. Notasdendrologicasde Venezuela. VIII.<br />
Bol. Soc. Venez. Ci. Nat. 8(56): 257-264.<br />
Poeppig, E. F. & S. Endlicher. 1838. Nova genera ac<br />
species pl<strong>an</strong>tarum 2. Leipzig.<br />
Pr<strong>an</strong>ce, G. T. 1977. The phytogeographic subdivisions<br />
of Amazonia <strong><strong>an</strong>d</strong> their influence on the selection<br />
of biological reserves. In G. T. Pr<strong>an</strong>ce &<br />
T. S. Ellias (eds.), Extinction is forever. The New<br />
York Bot<strong>an</strong>ical Garden, New York.<br />
Renner, 0. 1907. Beitrige zur <strong>an</strong>atomie und systematiek<br />
der Artocarpeen und Conocephaleen, inbesondere<br />
der Gattung Ficus. Bot. Jahrb. Syst. 39:<br />
319-448.<br />
Ruiter, G. de. 1976. Revision of the genera Myri<strong>an</strong>thus<br />
<strong><strong>an</strong>d</strong> Mus<strong>an</strong>ga (Moraceae). Bull. Jard. Bot.<br />
Etat. 46: 471-510.<br />
Rusby, H. H. 1910. New species from Bolivia, collected<br />
by R. S. Williams- 1. Bull. New York Bot.<br />
Gard. 6: 487-517.<br />
1927. Descriptions of new genera <strong><strong>an</strong>d</strong> species<br />
of pl<strong>an</strong>ts collected on the Mulford biological exploration<br />
of the Amazon valley. Mem. New York<br />
Bot. Gard. 7: 205-384.<br />
Smith, J. Donnell. 1905. Undescribed pl<strong>an</strong>ts of Guatemala<br />
<strong><strong>an</strong>d</strong> other Central Americ<strong>an</strong> republics. Bot.<br />
Gaz. 40: 1-11.<br />
NUMERICAL LIST OF TAXA<br />
1. <strong>Coussapoa</strong><br />
1-1. C. <strong>an</strong>gustifolia Aublet<br />
1-2. C arachnoidea Akkerm<strong>an</strong>s & C. C. Berg<br />
1-3. C. argentea Akkerm<strong>an</strong>s & C. C. Berg<br />
1-4. C. asperifolia Tr6cul<br />
a. subsp. asperifolia<br />
b. subsp. magnifolia (Trecul) Akkerm<strong>an</strong>s & C.<br />
C. Berg<br />
c. subsp. rhamnoides (St<strong><strong>an</strong>d</strong>ley) Akkerm<strong>an</strong>s &<br />
C. C. Berg<br />
1-5. C. batavorum Akkerm<strong>an</strong>s & C. C. Berg<br />
1-6. C. brevipes Pittier<br />
1-7. C. chocoensis Cuatrecasas<br />
1-8. C. cinnamomea Cuatrecasas<br />
1-9. C. cinnamomifolia Mildbraed<br />
1-10. C. contorta Cuatrecasas<br />
1-11. C. crassivenosa Mildbraed<br />
1-12. C. cupularis Akkerm<strong>an</strong>s & C. C. Berg<br />
1-13. C. curr<strong>an</strong>ii Blake<br />
1-14. C. duquei St<strong><strong>an</strong>d</strong>ley<br />
1-15. C. echinata Akkerm<strong>an</strong>s & C. C. Berg<br />
1-16. C. ferruginea Trecul<br />
1-17. C. floccosa Akkerm<strong>an</strong>s & C. C. Berg<br />
1-18. C. glaberrima Burger<br />
1-19. C. herthae Mildbraed<br />
1-20. C. latifolia Aublet<br />
1-21. C. leprieurii Benoist<br />
1-22. C. longepedunculata Akkerm<strong>an</strong>s & C. C. Berg<br />
1-23. C. macerrima Akkerm<strong>an</strong>s & C. C. Berg<br />
1-24. C. m<strong>an</strong>uensis C. C. Berg<br />
1-25. C. microcarpa (Schott) Rizzini<br />
1-26. C. microcephala Trecul<br />
1-27. C. napoensis Akkerm<strong>an</strong>s & C. C. Berg<br />
Spix, J. B. & C. F. P. von Martins. 1831. Reise in<br />
Brasiliens 3. Miinchen.<br />
St<strong><strong>an</strong>d</strong>ley, P. C. 1919. Studies of tropical Americ<strong>an</strong><br />
ph<strong>an</strong>erogams-No. 3. Contr. U.S. Natl. Herb 20(6):<br />
173-220.<br />
1924. Nine new species of pl<strong>an</strong>ts from Central<br />
America. Proc. Biol. Soc. Wash. 37: 49-54.<br />
. 1930. Studies of Americ<strong>an</strong> pl<strong>an</strong>ts-III. Publ.<br />
Field Mus. Nat. Hist., Bot. Ser., 8(1): 3-73.<br />
. 1933. Poulsenia, a genus oftrees ofthe family<br />
Moraceae. Trop. Woods 33: 4-5.<br />
- . 1937a. StudiesofAmeric<strong>an</strong>pl<strong>an</strong>ts-VII. Publ.<br />
Field Mus. Nat. Hist., Bot. Ser., 17(2): 155-284.<br />
1937b. Flora of Costa Rica: Moraceae. Publ.<br />
Field Mus. Nat. Hist., Bot. Ser., 18(1): 378-392.<br />
1939. In A. C. Smith, Notes on a collection<br />
of pl<strong>an</strong>ts from British Gui<strong>an</strong>a. Lloydia 2:161-218.<br />
1940. Studies of Americ<strong>an</strong> pl<strong>an</strong>ts-X. Publ.<br />
Field Mus. Nat. Hist., Bot. Ser., 22(2): 65-129.<br />
Trecul, A. 1847. Sur la famille des Artocarp6es. Ann.<br />
Sci. Nat. Bot., Ser. 3, 8: 38-157.<br />
Ule, E. 1907. Die Pfl<strong>an</strong>zenformationen des Amazonas-Gebietes.<br />
Bot. Jahrb. Syst. 40: 114-172.<br />
Woodson, R. E. & R. W. Schery. 1960. Flora of P<strong>an</strong>ama:<br />
Moraceae. Ann. Missouri Bot. Gard. 47(2):<br />
114-178.<br />
1-28. C. nitida Miquel<br />
1-29. C. nymphaeifolia St<strong><strong>an</strong>d</strong>ley<br />
1-30. C. oligocephala Donnell Smith<br />
1-31. C. orthoneura St<strong><strong>an</strong>d</strong>ley<br />
1-32. C. ovalifolia Tr6cul<br />
1-33. C. pachyphylla Akkerm<strong>an</strong>s & C. C. Berg<br />
1-34. C. parviceps St<strong><strong>an</strong>d</strong>ley<br />
1-35. C. parvifolia St<strong><strong>an</strong>d</strong>ley<br />
1-36. C. purpusii St<strong><strong>an</strong>d</strong>ley<br />
1-37. C. scabra Akkerm<strong>an</strong>s & C. C. Berg<br />
1-38. C. sprucei Mildbraed<br />
1-39. C. tessm<strong>an</strong>nii Mildbraed<br />
1-40. C. trinervia Mildbraed<br />
1-41. C. v<strong>an</strong>nifolia Cuatrecasas<br />
1-42. C. villosa Poeppig & Endlicher<br />
1-43. C. viridifolia Cuatrecasas<br />
1-44. C. fulvescens C. C. Berg<br />
1-45. C. tolimensis C. C. Berg<br />
1-46. C. valaria C. C. Berg<br />
2. <strong>Pourouma</strong><br />
2-1. P. gui<strong>an</strong>ensis Aublet<br />
a. subsp. gui<strong>an</strong>ensis<br />
b. subsp. venezuelensis (Cuatrecasas) C. C. Berg<br />
& v<strong>an</strong> Heusden<br />
2-2. P. velutina Miquel<br />
2-3. P. bicolor Martius<br />
a. subsp. bicolor<br />
b. subsp. tessm<strong>an</strong>nii (Mildbraed) C. C. Berg &<br />
v<strong>an</strong> Heusden<br />
c. subsp. digitata (Tr6cul) C. C. Berg & v<strong>an</strong><br />
Heusden<br />
d. subsp. scobina (Benoist) C. C. Berg & v<strong>an</strong><br />
Heusden
Exsiccatae 197<br />
e. subsp. choco<strong>an</strong>a (St<strong><strong>an</strong>d</strong>ley) C. C. Berg & v<strong>an</strong><br />
Heusden<br />
2-4. P. cecropiifolia Martius<br />
2-5. P. cucura St<strong><strong>an</strong>d</strong>ley & Cuatrecasas<br />
2-6. P. cuspidata Mildbraed<br />
2-7. P. myrmecophila Ducke<br />
2-8. P. formicarum Ducke<br />
2-9. P. phaeotricha Mildbraed<br />
2-10. P. mollis Tr6cul<br />
a. subsp. mollis<br />
b. subsp. triloba (Tr6cul) C. C. Berg & v<strong>an</strong> Heusden<br />
2-11. P. melinonii Benoist<br />
a. subsp. melinonii<br />
b. subsp. glabrata C. C. Berg & v<strong>an</strong> Heusden<br />
2-12. P. hirsutipetiolata Mildbraed<br />
a. subsp. hirsutipetiolata<br />
b. subsp. hispida (St<strong><strong>an</strong>d</strong>ley & Cuatrecasas) C. C.<br />
Berg & v<strong>an</strong> Heusden<br />
2-13. P. tomentosa Miquel<br />
LIST OF EXSICCATAE<br />
a. subsp. tomentosa<br />
b. subsp. apiculata (Benoist) C. C. Berg & v<strong>an</strong><br />
Heusden<br />
c. subsp. persecta C. C. Berg & v<strong>an</strong> Heusden<br />
d. subsp. essequiboensis (St<strong><strong>an</strong>d</strong>ley) C. C. Berg &<br />
v<strong>an</strong> Heusden<br />
3. subsp. maroniensis (Benoist) C. C. Berg &<br />
v<strong>an</strong> Heusden<br />
2-14. P. stipulacea C. C. Berg<br />
2-15. P. acuminata Miquel<br />
2-16. P. ferruginea St<strong><strong>an</strong>d</strong>ley<br />
2-17. P. villosa Trecul<br />
2-18. P. oraria St<strong><strong>an</strong>d</strong>ley & Cuatrecasas<br />
2-19. P. saulensis C. C. Berg & Kooy<br />
2-20. P. ovata Tr6cul<br />
2-21. P. elliptica St<strong><strong>an</strong>d</strong>ley<br />
2-22. P. bolivarensis C. C. Berg<br />
2-23. P. minor Benoist<br />
2-24. P. napoensis C. C. Berg<br />
2-25. P. herrerensis C. C. Berg<br />
Some recent collections have been added to this list <strong>with</strong>out being cited in the lists of specimens.<br />
Acosta M., A., 12 (2-5).<br />
Acosta Solis, M., 6008 (1-10); 12661 (1-19); 12797 (1-<br />
10); 12884, 13605 (2-3d); 13629(1-42); 13687, 13713<br />
(1-10).<br />
Alencar, L., 497 (2-4).<br />
Allemao, F. et al., 446 (2-la).<br />
Allen, P. H., 3103 (39); 3251 (2-1 a); 3742 (2-3d);<br />
5120 (1-4b); 5949 (1-23); 6186, 6280 (1-42); 6401<br />
(2-3d).<br />
Aluisio, J., 70809 (2-23).<br />
Amaral, I. L. et al., 101 (2-11a); 473 (2-15); 478 (1-<br />
40); 595 (2-1 la); 604 (2-5); 666 (2-23); 761 (2-7);<br />
IG2-10-328, IG2-16-509 (2-5).<br />
Ameida, J. et al., 71 (2-10a).<br />
Ancuash, E., 178 (2-la); 290 (2-1 la).<br />
Anderson, C. W., 188, s.n. (1-26).<br />
Andrade, A. et al., 2340 (1-25).<br />
Andrade Lima, D. de, see Lima. D.<br />
Anonymus, 9783 (1-20); s.n. (2-4, 1-25).<br />
Appun, C. F., 296 (1-26).<br />
Araujo, D. S. Dunn de et al., 848 (1-25).<br />
Aristeguieta, L., 1699, 1704 (1-42); 2005 (2-16).<br />
As<strong>an</strong>za C., E., 32929 (2-4).<br />
Asplund, E., 9462, 12933 (2-4); 13020 (2-la); 14124<br />
(2-4); 16401 (1-41); 18780 (1-4b); 18840(1-9); 19835<br />
(1-42).<br />
Aublet, J. B. C. F., s.n. (1-20, 2-la, 2-1 la, 2-17).<br />
Aubreville, A., 144 (2-3c).<br />
Aubry-Lecompte, C. E., s.n. (1-20).<br />
Austin, D. F. et al., 4140 (2-3a); 7149 (1-20).<br />
Ayala, F. (et al.), 341 (2-spec.); 1436 (1-42); 1446 (2-<br />
4).<br />
Aymard, G. et al., 3990 (2-23); 6778 (2-15).<br />
BAFOG, 93-M (2-17); 1050 (2-3c); 4272 (2-1 la); 5042<br />
(2-23); 7087 (2-17), 7152, 7173 (2-23); 7228 (2-13e);<br />
7261, 7268 (2-23); 7330 (2-13e); 7562 (2-2); 7745<br />
(2-17); 7938 (1-1); 7942, s.n. (1-4a).<br />
Bailey, I. W., 18, 19, 20, 146, 147 (2-la).<br />
Baker, M. A., 6017 (2-3a).<br />
Balick, M. et al., 1021 (1-39).<br />
Barbour, P. J., 5372 (1-42).<br />
Barclay, A. S. et al., 3294, 3593 (1-42).<br />
Barrier, S., 5114 (2-13e).<br />
Bartlett, H. H., 12356 (1-30).<br />
Bastos, M. A., 2051 (2-la); 2220 (2-23).<br />
BBS, 3003 (1-20).<br />
BBS (=Bosbeheer Suriname), 297 (2-17); 298 (2-13a);<br />
1128 (2-la).<br />
Belem, R. P. (et al.), 644 (1-25); 1471 (2-la).<br />
Benoist, R., 341 (2-17); 465 (2-13e); 960, 978 (2-23);<br />
3003 (2-3d); 3010 (1-10); 3042 (1-41); 3044 (1-42).<br />
Berg, C. C. (et al.), 276 (2-7); 355 (2-1 lb); 400 (2-23);<br />
BG.598, BG.772 (2-la); 1033 (1-42); 1038 (1-40);<br />
1040 (1-42); 1041, 1042, 1045 (2-5); 1048 (1-40);<br />
1053 (2-5); 1058 (1-40); 1062 (1-42); 1070 (2-5);<br />
1111 (1-31); 1113 (1-42); 1119 (1-4b); 1120, 1129,<br />
1131 (1-11); 1132 (1-32); 1133 (1-4b); 1143 (2-2);<br />
1153 (1-33); 1222 (1-42); 1223 (2-4); 1241 (1-42);<br />
1259 (1-19); 1301, (1-42); 1223 (2-4); 1546 (1-44);<br />
P.17591, P.17592 (1-28), P.18136 (2-1 la); P.18152<br />
(2-3a); P.18409, P.18453 (1-40); P.18455, P.19841<br />
(1-42); P.19878 (2-13b), s.n. (1-4b, 1-42, 2-13a).<br />
Berlin, B., 372 (1-42); 523 (2-4); 746 (2-lOb); 1531 (1-<br />
32); 1999 (2-4).<br />
Bernardi, L., 902 (2-1 la); 910 (2-3a); 1724(1-42); 1919<br />
(2-3d); 1980 (1-42); 1986 (2-3d); 2657 (2-3a); 2731<br />
(2-13b); 2744 (1-43); 2808 (2-1 la); 3354 (1-42); 5696<br />
(2-lb); 6348, 7017 (1-42); 7238 (1-3); 7250 (2-23);<br />
5.16.5. (2-20); s.n. (tree 3/100) (2-25); s.n. (tree 4/8)<br />
(2-23); s.n. (tree 6/27) (2-13a); s.n. (1-42, 2-lb, 2-5,<br />
2-13a).<br />
Bisby, F. et al., P. 18118 (2-3a).<br />
Billiet, F. et al., 1108 (1-20).<br />
Black, G. A. (et al.), 46-137 (1-8); 47-1520 (2-13a); 48-<br />
2949, 48-3017 (1-28).<br />
Bl<strong>an</strong>chet, J. S., 160, 2327 (1-25); 2361 (2-lOa); s.n. (1-<br />
25).<br />
Bl<strong>an</strong>co, C., 810 (2-la); 958 (1-42); 1149 (2-3a).
198 Flora Neotropica<br />
Blum, J. L., 2039 (2-3e).<br />
Coelho, D. (et al.), 672 (2-1 la); 1577 (1-4a); 5979 (2-<br />
Boerboom, J., 9199 (2-10a).<br />
20); 51350, 52381 (2-23); 52383 (2-13d).<br />
Bondar, G., 2169 (2-10a); s.n. (2-la).<br />
Coelho, L. (et al.), 15 (2-17); 332 (1-4c); 339 (2-16);<br />
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).<br />
Bowie, J. et al., 57 (1-25).<br />
Const<strong>an</strong>ino, D., 19689 (1-25).<br />
Boy<strong>an</strong>, R., 267 (2-7).<br />
Contreras, E., 61, 845, 2220, 3559, 4449, 5787, 6231<br />
Brade, A. C. (et al.), 7887, 7949 (1-25); 18619 (2-la). (1-30); 10026 (2-3d); 10195 (1-30); 10763 (1-42);<br />
Braga, P. L. S., 3178 (2-4).<br />
11195 (2-3d).<br />
Br<strong><strong>an</strong>d</strong>byge, J. et al., 30194 (2-3a); 30252 (2-23); 30258 Cook, O. F. et al., 157 (1-29); s.n. (1-30).<br />
(1-31); 30327 (2-23); 30421 (1-32).<br />
Cooper, G. P., 538 (1-42).<br />
Brenes, A. M., 20542 (1-34).<br />
Cordeiro, M. R., 140 (2-23); 367 (1-4b).<br />
Breteler, F. J., 3889 (1-42); 4920 (2-3d).<br />
Cordoba, J. J., 295 (1-42).<br />
Brewer-Carias, C., s.n. (1-11).<br />
Correa, M. D. et al., 1002, 1031A (1-6); 1854 (2-3d).<br />
Broadway, W. E., 880 (1-4a).<br />
Costa, M. da, 90 (1-39).<br />
Bruijn, J. de, 1123 (1-42); 1496 (2-1 lb); 1497 (2-3a); Cow<strong>an</strong>, R. S. (et al.), 1868, 1880 (1-26); 38774 (1-20).<br />
1502 (2-llb); 1514, 1516, 1517, 1526 (2-3a); 1546 Cremers, G., 6726, 8271 (2-lOa).<br />
(2-1 lb); 1547 (2-12a); 1555 (2-1 lb).<br />
Croat, T. B., 5124,5125,6588 (1-4b); 7633 (2-4); 7768,<br />
Buchtien, 0., 2050, 2122 (2-3a).<br />
7839 (1-42); 7859 (1-4b); 8097, 8100, 8648 (2-3e);<br />
Buck, P., 26425 (1-25).<br />
9973A, 14212(1-6); 10343, 10830 (2-3e); 11591 (2-<br />
Bunting, G. S. (et al.), 3959 (1-4b); 6836 (2-3b). 3d); 15064 (1-6); 17784 (1-40); 18016 (2-la); 18643<br />
Burchell, W. J., 3148 (1-25); 9792 (2-3a <strong><strong>an</strong>d</strong> 2-lOa). (1-4b); 18850 (1-28); 18888 (2-15); 19997 (2-3a);<br />
Burger, W. C. et al., 8137 (2-23); 10040, 10474 (1-42). 20370 (1-4c); 20411 (1-40); 20447 (2-1 a); 20616<br />
Busey, P., 628 (1-34).<br />
(2-la); 20662 (2-4); 22635 (1-42); 24663 (1-30);<br />
BW (=Boschwezen) 732 (1-20); 772 (1-1); 1322 (1-4a); 24831 (2-30); 59353 (2-3a).<br />
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-<br />
2410 (2-lOa); 3255, 3302 (2-17); 3376 (2-3a); 3430 43); 962 (2-1la).<br />
(1-4a); 4010 (2-17); 4033 (2-3a); 4494 (2-1Oa); 4866 Crueger, H., s.n. (1-42).<br />
(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-<br />
(2-17); 5440, 5581, 5992 (2-10a); 6292 (2-3c); 6509 26a); 1577 (1-26); 1657 (2-la); 1729 (1-4b); 2241,<br />
(2-17).<br />
2387, 2636, 2813, 3017 (1-26).<br />
Cabrera, J., 2668 (1-35).<br />
Cuatrecasas, J. (et al.), 6737 (2-4); 7299 (2-23); 7510<br />
Callejas, R. et al., 3997 (1-9).<br />
(1-42); 7602 (2-4); 8291 (1-42); 10714 (1-27); 14284<br />
Calzada, J. I., 325 (1-36).<br />
(1-10); 14881 (2-12b); 15071, 15520, 15534 (2-3d);<br />
Camargo, -, 1849 (1-25).<br />
15589 (1-42); 15720 (2-18); 15876, 16315, 16755<br />
Camp, W. H., E3626 (1-42); E3674 (1-19).<br />
(2-3e); 16826 (2-12b); 17251 (1-34); 17357 (2-3c);<br />
Campbell, D. G. et al., 6820 (2-unnamed); 6834 (2-6), 17451, 17479 (2-3e); 17647 (1-10); 21256 (1-41);<br />
6927 (2-5); 7001 (2-la); 7157 (2-3a); 7264 (2-23); 21446 (1-7); 21496 (1-10); 22048 (1-41); 22526 (1-<br />
7600 (2-3a); 7689 (2-unnamed); 8065, 8127, 8267 42).<br />
(2-la); 8344 (2-23); 8383 (2-lOb); 8463 (2-23); 8537 Curr<strong>an</strong>, H. M., 8 (1-13); 19 (2-la); 21 (2-1Oa); 116 (2-<br />
(2-la); 8844 (2-4); 8997 (2-3a); 9120 (2-lOb); 9521 12a).<br />
(2-3a); 9610 (2-la).<br />
DAF, 004 (1-13); 027 (2-lOa).<br />
Campos Porto, P., 863 (1-25).<br />
Daly, D. C. et al., 1208, 1474 (2-3c); 3790 (2-17); 3936<br />
C<strong>an</strong>tonnement de Cayenne, s.n. (2-1 la).<br />
(2-2); 4129, 4135 (2-7); 4457 (2-16); 4456, 4485 (2-<br />
Carauta, J. P. P. (et al.), 178, 1714, 1716, 1780, 1781, 8), 5356 (1-11).<br />
2722; 3011 (1-4a); 3050, 3166 (1-25).<br />
Damazio, L. B., s.n. (1-25).<br />
Cardenas, M., 1990 (2-1a); 3973 (2-13c).<br />
Damiao, C., 2503, 2817 (1-28); 680 (2-20), see also<br />
Cardona, F., 374 (1-4b); 1222 (2-13b); 2170 (2-5). Mota, C. Damiao A. de.<br />
Casaretto, J., 617 (1-13); 643 (1-25).<br />
Damiao, J. P. L. et al., 1805 (2-3a); 1822 (2-13b).<br />
Castillo S., M., 11 (2-la).<br />
D<strong>an</strong>iel, H., 1793, 2652 (1-42).<br />
Cavalc<strong>an</strong>te, P., 606 (2-23); 777 (2-3a); 2297<br />
D<strong>an</strong>ta, M., 12383 (1-31).<br />
(2-3c).<br />
Cazolet, P. C. D. et al., 5037, 5072 D'Arcy, W. G., 12329 (1-6).<br />
(2-3d).<br />
Cer6n M., C. E.<br />
Davidse, G. et<br />
(et al.), 252<br />
al., 15365<br />
(2-4); 261<br />
(2-2).<br />
(2-la); 2019 (2-<br />
24); 2955<br />
Davidson, C., 5203<br />
(1-14); 2993 (1-9); 3389<br />
(2-9).<br />
(1-32); 3495 (1- Dayton, W. A., 3019 (1-42).<br />
22).<br />
Denslow, J., 2698 (2-3e).<br />
Chagas, J., 962 (2-4); 1364 (1-4a); 1564 (2-7); 1655 (2- Deward, G., 27 (2-17).<br />
2).<br />
Diaz, C. et al., 618 (1-32).<br />
Cid (Ferreira), C. A. et al., 435, 511 (1-40); 721 (2-23); Diaz, M., 22A (2-20).<br />
847 (2-13d); 849 (2-2); 942 (1-20); 1053, 2223 (1- Dodson, C. H. (et al.), 3017 (1-39); 5241 (1-42); 5776<br />
1); 2870 (2-la); 3297 (2-20); 6200 (2-2); 7031 (2- (1-19); 5778 (1-42); 6082 (1-19); 6156, 6312, 6327<br />
30); 7831 (1-4a).<br />
(2-3d); 6513, 7416, 7616 (1-41); 9539 (1-19); 15201<br />
Claes, F., 26 (2-4).<br />
(1-34).<br />
Clewell, A. et al., 4142 (2-3d).<br />
Doeve, -, 3003 (1-20).
Exsiccatae<br />
Dombey, J., s.n. (1-42).<br />
Focke, H. C., 9, 418, s.n. (1-20).<br />
Donnell Smith, J., 4826, 6770 (1-42).<br />
Foldats, E., 114-1A (2-2).<br />
Donselaar, J. v<strong>an</strong>, 1226 (2-13e); 1262 (2-3c); 1373, Folsom, J. P. et al., 6377 (2-3d); 6469 (2-23).<br />
1681 (2-lOa); 1888 (2-17); 2569 (1-1); 2924 (2-lOa); Fonnegra, R. et al., 1798 (1-35).<br />
3333 (2-3c); 3788 (2-lOa).<br />
Forero, E. et al., 3145 (2-23); 4099, 4110(1-10); 6367<br />
Dressier, R. L., 3454 (1-6); 3579 (2-3d); 4636 (1-15). (2-23), P.6419 (2-4).<br />
Duarte, A. P. (et al.), 3410, 4650, 5328 (1-25); 6970 Forest Department British Guy<strong>an</strong>a, see FD.<br />
(1-4a); 7194 (2-23); 9785 (2-1 la); s.n. (1-13). Foresta, H. de., 526 (2-13e); 676, 677, 4970, 5011 (2-<br />
Ducke, A. (et al.), 104 (1-25); 1149 (2-7); 1354 (1-38); la).<br />
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-<br />
1917 (2-8); 1999 (1-38); 2103 (1-4a); 6782, 7715 (1- 4b); 3412 (1-32); 5019 (2-23); 5641 (1-24); 5719 (2-<br />
28); 9096 (1-38); 10736 (2-17); 12238 (1-28); 12366 10b); 5721 (1-42); 5903 (2-4); 6568 (2-la); 6569 (2-<br />
(2-4); 13056 (2-17); 13057 (2-3a); 13058 (2-10a); 10b); 7001 (2-4); 7194 (2-10b); 7199 (2-23); 7200,<br />
13065 (1-4b); 13066 (1-28); 14499 (2-7); 15255 (2- 7201 (2-la); 7995 (2-lOb); 8023, 9373 (2-la).<br />
17); 15261 (2-lOa); 15350(2-2); 15804(1-4a); 15935 Fr<strong>an</strong>cisco, 2186 (2-7).<br />
(2-2); 16003 (1-28); 16321 (1-25); 17132 (1-4b); Fr<strong>an</strong>co, J. et al., 419 (1-31).<br />
17164 (2-7); 17167 (2-2); 18455 (1-40); 18456 (1- Frazao, A., s.n. (1-25).<br />
20); 18460 (1-4c); 18461 (1-1); 18462 (1-21); 19455<br />
Froes, R. de Lemos (et al.), 1907<br />
(2-1Oa); 21200 (1-25); 23606, 23607 (2-7); 23608<br />
(1-20); 20870 (2-8);<br />
(2- 21048 (2-4); 21264 (1-39); 23693<br />
20); 23609 (1-38); 25243 (2-17); 25244 (2-16); 25245<br />
(2-16); 23725 (2lla);<br />
25531 (1-28); 26520<br />
(2-13b); 25246 (2-21); 25247 (2-2); 25248<br />
(2-13d); 26631 (2-3c);<br />
(1-38); 28585<br />
s.n.<br />
(1-4b); 34651 (1-20).<br />
(2-13b).<br />
Duke, J. A. (et al.), 6588, 8035 (2-3d); 11273<br />
G<strong><strong>an</strong>d</strong>oger, M., 19 (2-2).<br />
(1-42);<br />
11599 (2-3d); 14712 (1-6); 15742 Garcia-Barriga, H., 12408 (1-4b); 13643<br />
(1-4b).<br />
(2-1 la); 13836<br />
Dunlap, V. C., 299 (2-23); 13871 (2-4); 14002, 14106<br />
(1-42).<br />
(2-5); 14116 (2-<br />
1<br />
Duque-Jaramillo, J. M., 1767 (1-4); 2179 (1-8); 2207<br />
la); 16239 (1-4b).<br />
(1-28); 2261, 2446 (2-4); 3957 Garcia, J. H. et<br />
(1-42).<br />
al., 9 (1-3a).<br />
Dus6n, P., 2160, 6635, 10134, 10186, 11419 (1-25);<br />
Gardner, G., 732, 5632, 5858 (1-25).<br />
11942 (2-la); 13653, 14718, 17030 (1-25); 17239, Gamier, Bro. A., 112 (2-23).<br />
17345 (2-la); s.n. (1-25, 2-la).<br />
Garwood, N. C., 759 (2-3d).<br />
Dwyer, J. D., 4734 (2-3d); 10305 (1-42); 10799 (1-30). Gasche, J. et al., 29 (1-31).<br />
Dyer, F. J., A-85 (1-42).<br />
Gaudichaud, C., 992 (1-25).<br />
Edwards, J. B., 400 (1-42).<br />
Geay, M. F., 3285 (1-1).<br />
Eggers, H. F. A. von, 14165, 15615 (1-42).<br />
Gentle, P. H., 2602 (1-30); 2787, 2917, 3265, 4211 (2-<br />
Egler, F. E., 42-229 (1-30).<br />
3d); 4528 (1-30); 6312, 6997, 8675 (2-3d).<br />
Egler, W. A. et al., 46027 (1-28).<br />
Gentry, A. H. (et al.), 918 (1-25); 2046, 2657 (2-1 lb);<br />
Eiten, G. et al., 6214 (1-25).<br />
6314 (1-42); 6679 (2-3e); 7530 (1-42); 7727, 8348<br />
Ellenberg, H., 2288 (2-4); 2297 (2-lOb); 2474 (2-4). (1-30); 8707 (1-42); 9552 (2-3d); 9606 (1-19); 9824<br />
Elias, Bro., 130, 631 (1-42).<br />
(2-la); 9892 (1-19) 9934, 10127, 12474 (1-42); 13408<br />
Emden, W. C. v<strong>an</strong>, L-VI, s.n. (1-20).<br />
(1-15); 14011 (see 1-32); 15621 (2-lOb); 15876 (2-<br />
Emmerich, M. (et al.), 3003, 3556 (1-25).<br />
7); 16041 (2-3b); 16181 (2-23); 17217, 17493 (1-4b);<br />
Emygdio de Mello Filho, L. (et al.), 412, 2340, 2447, 17804 (2-18); 18458 (1-42); 18528 (1-31); 19048 (2-<br />
3018 (1-25).<br />
unnamed); 20381 (1-28); 20356 (2-3a); 20495 (1-<br />
Englesing, F. C., 50, 52a (2-3d).<br />
28); 20515 (2-5); 20766 (1-40); 20802 (1-28); 21167<br />
Erl<strong>an</strong>son, C. 0., 240 (1-42).<br />
(1-35); 21193 (2-3a); 21205 (2-la); 21291 (1-42);<br />
Ernst, W. R., 324 (1-42).<br />
21319 (1-4b); 21346 (2-25); 22095 (1-42); 22280 (1-<br />
Escobar, L. Albert de et al., 4219 (1-45).<br />
4b); 22381 (2-spec.); 22460 (2-4); 22865 (2-13a);<br />
Espina, J. et al., 219 (1-31).<br />
22866 (2-unnamed); 23713 (1-42); 23835 (2-3e);<br />
Espinal, T. S., 822 (1-4b); 1314 (1-42).<br />
24074 (1-4b); 24900, 25050, 25078, 25100 (2-5);<br />
Falcao, M., 215 (2-4); 1105 (2-14).<br />
25124 (1-28); 25151 (1-32); 25527 (2-23); 25854 (2-<br />
F<strong>an</strong>shawe, D. B., 1105 (2-14); 2050 (2-la); 3031 (1- 5); 26049, 26117 (2-23); 26525 (1-10); 26530 (2-3d);<br />
26); 4970, 5011 (2-1a).<br />
27194 (2-23); 27566 (1-22); 27799 (2-spec.); 28609<br />
Farifias, J. et al., 347 (2-4); 490 (2-1 la).<br />
(2-3d); 30278 (2-3e); 31179 (2-10b); 31197 (2-3b);<br />
FD (=Forest Department British Guy<strong>an</strong>a), 1011 (2- 31271 (2-23); 31655 (2-5); 31816, 31933 (2-la);<br />
la); 3031 (1-26); 3851 (2-14); 4014, 4073 (1-4b); 31938, 31976, 36231 (2-23); 36241 (2-4); 36309 (2-<br />
4090, 4970, 5011, 5936, 6914 (2-la); 6973 (1-4b). 23); 36797 (2-18); 39264 (2-13b); 39379 (2-20);<br />
Fendler, A., 1237, 1237a (1-42); 2420 (2-lb).<br />
40479 (2-3e); 41890 (2-6); 41894 (2-13); 42029 (2-<br />
Fern<strong><strong>an</strong>d</strong>ez P., A. (et al.), F3, 1458 (1-42); 2019 (2-23). la); 42168 (2-3d); 42232 (2-1 Ob); 42568 (2-23); 42793<br />
Ferreira, C. A. Cid, see Cid (Ferreira), C. A.<br />
(2-lOb); 43137 (2-4); 43454 (2-23); 45160 (2-3d);<br />
Ferreira, V. F., 493 (1-25); s.n. (2-20).<br />
45660 (2-4); 45915 (2-la); 45922 (2-5); 46154 (1-<br />
Ferreyra, R., 4813 (1-32).<br />
40); 46752 (2-3c); 47838 (1-46); 48303 (2-3e); 49026<br />
Ferry, J. F., s.n. (1-42).<br />
(2-lOa); 49127 (2-1 la); 49346 (2-13); 51089, 51111<br />
Feuillet, C. P., 1070 (2-23).<br />
(2-23); 51154 (2-la); 51177 (2-5); 51207 (2-23);<br />
199
200 Flora Neotropica<br />
51355 (2-19); 51515 (2-23); 53622 (2-3e); 53829 (1-<br />
10); s.n. (2-5).<br />
Gevieski, A., 85 (1-25).<br />
Gillis, W. T. et al., 10191 (1-42).<br />
Glaziou, A. F. M., 81, 1013, 1138, 1942, 2016, 4937,<br />
6009 (1-25); 8934, 8934a (1-13); 8935 (2-la); 8936<br />
(1-25); 10070a (1-31); 10070 (2-15); 12166 (1-25);<br />
12173 (2-la); 16350, s.n. (1-25); 20408 (2-la).<br />
Gleason, H. A., 174 (2-la); 405, 415 (1-26); 670 (2-<br />
10a).<br />
Goeldi, A., 7731 (2-10a); 7732 (2-1 la); 7773 (2-lOa).<br />
Goes, O. C. et al., 815 (1-25).<br />
Gomes, M. (et al.), 326 (2-la); 547, 593 (2-23); 614<br />
(2-5); 632, 635 (2-23); 636 (2-13); 646, 655, 673 (2-<br />
23); 676 (2-3a); 690, 759 (2-23); 786 (2-3a); 789,<br />
810, 813 (2-23); 826 (2-13b); 838 (2-23); 843 (2-3a);<br />
844 (2-5); 872, 891, 924, 926, 931, 933, 955; 966,<br />
979 (2-3a); 1021 (2-2); 1033 (2-23); 1040 (2-5); 1048<br />
(2-23); 1052, 1072, 1079, 1088 (2-3a); 1118, 1120<br />
(2-23); 1123 (2-3a); 1129, 1156, 1175; 1176 (2-3a);<br />
1196 (2-2); 1255, 1319, 1336, 1338, 1356 (2-23);<br />
1357 (2-2); 1378, 1419, 1423, 1425, 1444, 1446 (2-<br />
23); 1517 (2-13b); 5134 (2-23); 1540 (2-13b); 1534,<br />
1568, 1570, 1577, 1584 (2-23); 1588 (2-13b); 1593<br />
(2-25); 1594, 1595(2-23); 1598, 1604, 1610(2-13b);<br />
1636, 1640, 1641 (2-23); 1679 (2-13b); 1692, 1731,<br />
1745, 1769, 1822, 1823, 1867, 1893, 1917, 1983,<br />
2170 (2-23); 2399 (2-3a).<br />
Gomes, V. et al., 2818 (1-13); 2819 (1-17).<br />
Gomez Pompa, A. et al., 3385 (2-3d).<br />
Gonzalez, A. et al., 1319 (2-1b).<br />
Graham, Mrs., s.n. (1-25).<br />
Graimleux, -, 7938 (1-1); 7942 (1-4a).<br />
Gr<strong>an</strong>ville, J.-J. de (et al.), 581 (2-23); 2744 (2-19); 4531<br />
(2-13e); B.5071 (2-la); B.5191 (1-20); 5284 (2-la);<br />
5642 (2-23).<br />
Gren<strong><strong>an</strong>d</strong>, P., 445 (2-la); 688 (2-13e); 966 (1-4a); 1352<br />
(2-la); 1442 (2-3c); 1478 (1-20); 1596 (1-4a).<br />
Grubb, P. J. et al., 1545 (2-24); 1681 (2-13a).<br />
Grubbe, J. P. et al., 1205, 1283 (1-14); 1498 (1-4b).<br />
Gu<strong>an</strong>chez, F., 239 (2-4).<br />
Guedes, M. or T., 1235 (2-2).<br />
Guillemin, J.-B. A., 1024 (2-la); 1339 (1-13).<br />
Guppy, N., 656 (2-lOa).<br />
Gutierrez, G. et al., 562 (1-31).<br />
Haber, W. A., 1545 (1-34).<br />
Hage, J. L., 31 (1-25).<br />
Hahn, W., 347 (1-11).<br />
Halle, F., 1132 (2-la).<br />
Hamilton, B. et al., 1062 (1-18).<br />
Hammel, B., 2176, 2576 (1-34); 4475, 6006 (1-6).<br />
H<strong>an</strong>s, D., 335 (1-25).<br />
Harling, G et al., 14733 (1-4b).<br />
Haroldo, 57512 (2-13a).<br />
Hartm<strong>an</strong>, R. L., 12176 (2-3d).<br />
Hartshorn, G. S., 1224, 1295 (2-23); 1348 (2-3d); 1431<br />
(2-23); 1436 (1-29); 1539 (2-3d); 2157 (1-34).<br />
Hatch, W. R. et al., s.n. (1-30).<br />
Hatschbach, G., 1475, 2458 (1-25); 7411, 7466 (2-la);<br />
7807, 7833, 16304, 18680, 20253 (1-25); 25765,<br />
29129 (2-la); 29629, 41730 (1-25).<br />
Hayes, S., 32, 348 (2-3e); 354 (1-4b); 414, 867, 986,<br />
1008, s.n. (1-42).<br />
Hazlett, D., 664, s.n. (1-42).<br />
Hern<strong><strong>an</strong>d</strong>ez, J. J. et al., 369 (1-9).<br />
Herrera Ch., G., 501, 503 (1-34); 505 (1-29).<br />
Heyde, N. M. et al., 32, 39 (2-23); 78 (1-20).<br />
Hilty, S., M-86 (2-23); M-87 (1-34).<br />
Hine, R., P-1609 (2-23).<br />
Hinton, G. B., 14018 (1-36).<br />
Hohenkerk, L. S., 738 (1-4b).<br />
Hoehne, F. C., 30923 (1-25).<br />
Holdridge, L. R., 2514 (2-3d); 6254a (2-3e); 6818 (2-<br />
23).<br />
Holm-Nielsen, L. et al., 24519 (1-14); 24856 (1-10);<br />
26331, 26610 (1-42).<br />
Hoist, B. K. et al., 2421 (2-1 la); 2719 (2-la); 3195 (2-<br />
3a).<br />
Hostm<strong>an</strong>n, F. W. R., 1189 (1-4a).<br />
Hostm<strong>an</strong>n, F. W. (& A. Kappler), 1272 (2-lOa <strong><strong>an</strong>d</strong><br />
2-1 la).<br />
Howard, R. A. et al., 605 (1-42).<br />
Hoyos, J., 2043 (1-42).<br />
Huashikat, V., 431 (2-1 la); 912 (2-3b); 944 (2-lOb);<br />
966 (2-13a); 1027 (2-10b); 1150(2-23); 1216 (2-la);<br />
1241 (2-3a); 1432, 1482 (2-13a); 1585 (see 1-12);<br />
1872 (2-3b).<br />
Huber, J. E., 238 (2-la); 1640 (2-2), 1775 (1-21), 1873<br />
(1-4a), 4244 (2-la).<br />
Humbert, H. et al., 27221 (2-4).<br />
Idrobo, J. M. et al., 786 (2-3a); 1316 (2-lOb).<br />
Ijjasz-Madriz, -, 28 (2-1b).<br />
Irwin, H. S. (et al.), 183 (1-26); 2058 (1-17); 4783 (2-<br />
lla); 47874 (2-23); 48192 (2-3a); 48439 (2-1 la);<br />
48468 (2-23); 48631 (1-16); 48682, 48755 (2-3c);<br />
55487 (1-20); 55576, 55884 (2-17); 57574 (1-1).<br />
Izawa, K., 9 (2-la); 21 (2-4).<br />
J<strong>an</strong>se, C. 0., 276 (2-3d).<br />
Jaramillo, J. et al., 2088 (1-40); 4586, 31516 (2-4).<br />
Jativa, C. (etal.), 303, 1079 (2-12b); 1097 (1-10); 2033,<br />
2037 (2-12b).<br />
Jenm<strong>an</strong>, G. S., 652, 3995, 4947 (1-26); 4947A (1-4b);<br />
5315 (1-26); 6310 (2-la); 7310, 7933 (1-26).<br />
Jimenez, M. A., 433 (1-35); 657, 1750, 2887 (1-42);<br />
3016 (1-18); 3816 (2-3d).<br />
Jim6nez Saa, H., 15631 (=LBB 14296) (2-17).<br />
Jobert, -, 682 (2-15).<br />
Johnston, I. M., 580, 606, 608, 652, 678 (1-42); 1709<br />
(1-4b); 1714 (2-3e), 1725, 1768 (1-42).<br />
Jones, G. C. et al., 3152 (1-30).<br />
Jones, J. et al., 9712, 9769 (1-28); 9788 (1-40).<br />
Jonsson, G., 5A, 610a (1-25).<br />
Juncosa, A., 1253 (2-12b).<br />
K<strong>an</strong>ehira, R., 99 (1-42).<br />
Kappler, A., 1272 (2-11 a).<br />
Karsten, H., s.n. (1-42, 2-la, 2-lb, 2-3a, 2-4).<br />
Kayap, R., 201 (2-13a); 268, 393 (2-la); 571 (2-1 la);<br />
1057 (2-la); 1306 (2-10b).<br />
Kenoyer, L. A., 312 (2-3e).<br />
Killip, E. P. (etal.), 25341, 26095 (1-32); 27381,27932<br />
(2-4), 28800 (2-unnamed); 29246 (1-42); 29839 (2-<br />
4); 29955 (2-23); 30139 (2-20); 30179 (1-38); 35095<br />
(2-3e).<br />
Kinloch, J. B., s.n. (1-30).<br />
Klein, R. M., 89 (2-la); 79B, 772 (1-25); 1110 (2-la).<br />
Klug,G., 1185, 1326 (2-4).
Exsiccatae 201<br />
Korning, J. et al., 47618 (2-13c), 47641, 47645 (2-3a); Lindem<strong>an</strong>, J. C. (et al.), 1857 (1-25); 3631, 3689 (2-<br />
47801, 47807 (2-unnamed).<br />
10a); 4057 (2-2); 5468 (2-3c); 5682 (1-25); 5709 (1-<br />
Kosei Izawa (see also Izawa, K), 9 (2-la); 21 (2-4). 4a); 5748 (1-25); 6123,6166 (2-3c); 6362 (1-1); 6418<br />
Krieger, P(adre) L. (as PLK) (et al.), 12268 (2-4); 12296 (2-2); 6445, 6600 (1-1); 6765 (2-3c); 6923 (2-lOa);<br />
(1-42).<br />
6938 (1-20); 9105, 9110 (1-25).<br />
Krukoff, B. A., 1147, 1187 (1-39); 1197 (2-17); 1297 Lindem<strong>an</strong>, J. C. & A. C. de Roon, 747a (2-3c); 828a,<br />
(2-la); 4518 (1-42); 4817 (2-la); 4994 (1-42); 5109 842b (2-3a).<br />
(2-4); 5282 (2-23); 5309 (2-lOb); 5327, 5332 (2-4); Lindem<strong>an</strong>, J. C. & A. L. Stoffers et al., 33 (1-29); 34<br />
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-<br />
6238 (2-la); 6524 (1-42); 6662 (l-4b); 6887 (2-13b); 20); 790 (2-17); 799 (1-20).<br />
7071 (2-23); 7073 (2-20); 7966 (1-4a); 8073 (2-23); Lisb6a, B., s.n. (1-13, 1-25).<br />
8223 (2-13b); 8273 (1-35); 8325 (2-13c); 8332 (2- Lisb6a, P. L., 27 (2-4).<br />
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-<br />
13c); 8332 (2-4); 8369 (2-la); 8373 (2-6); 8386 (2- 32); 602 (1-42); 635 (1-11); 6155 (1-42); 6241, 6258,<br />
la); 8388 (2-21); 8401 (1-28); 8406 (1-4c); 8427 6301 (2-3d); 6328, 6461 (1-42); 6640 (2-3d); 7756<br />
(2-15); 8469 (2-4); 8518 (1-31); 8539 (1-35); 8587A (2-4); 8423 (2-la); 9527 (2-6); 9629 (1-40); 9641,<br />
(1-31); 8652 (2-la); 8658 (1-35); 8695 (2-20); 8755 9789 (1-42); 15990 (1-4b); 16230, 16245, 21047 (1-<br />
(1-35); 8807 (2-16); 8897 (1-35); 8967 (1-31); 8994 42); 21056 (1-41); 21070 (2-12b); 21136 (1-34);<br />
(1-39); 10123 (1-4b); 10861 (2-13c); 10867, 10874 21172 (1-42); 21225 (2-12b).<br />
(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).<br />
11275 (1-9).<br />
Lleras, E. et al., P.17178, P.17197 (1-39); P.17242 (2-<br />
Kuhlm<strong>an</strong>n, J. G. (et al.), 255 (2-13b); 263 (1-20); 291 9); P.17411 (2-8); P.17484 (1-4a); P.19586 (2-20);<br />
(2-13b); 368 (1-42); 382 (2-2); 446 (2-10a); 470 (1- P. 19661 (1-20).<br />
37); 759 (1-42); 1178 (1-28); 1337 (2-4); 1540 (1- Loureiro, A. (et al.), 16461 (2-13a); 16568, 16968,<br />
32); 1832, 2073, 2074 (1-17); 2075 (2-la); 2177 (1- 50623 (2-20).<br />
17); 2200 (2-la); 2207 (1-17); 5004 (1-25); 19843, Lowrie, S. R. et al., 710 (2-23).<br />
150082 (2-7); s.n. (1-13).<br />
Loz<strong>an</strong>o, C. G., 640 (1-42).<br />
Kuhlm<strong>an</strong>n, M. et al., 52 (2-2).<br />
Luetzelburg, Ph. von, 12471 (1-25); 23887, 23913 (2-<br />
Lambroy, -, s.n. (1-4b).<br />
4).<br />
L<strong>an</strong>jouw, J. et et al., 399 (2-1 la); 432 (2-13e); 1761 Lugo, M., 83 (1-4b); 2820, 2857, 2909, 3563 (1-27).<br />
(2-2); 2106 (2-10a); 2113 (2-3c); 2114 (2-23). Lund, P. W., 139 (1-25).<br />
L<strong>an</strong>na Sobrinho J. de Paula et al., 376, 1011 (2-la). Lundell, C. L., 2, 648, 3170, 3535, 6224, 6350, 15930,<br />
Lao, E. A. et al., 21 (1-42).<br />
16122, 16162 (1-30).<br />
Lao, M. R. et al., s.n. (2-4).<br />
Luschnath, B., s.n. (1-25); 40, s.n. (2-lOa).<br />
Lasser, T., 2202, 2267 (2-lb).<br />
Maas, P. J. M. et al., 2478 (1-26); 2592 (2-14); 3325<br />
Lawr<strong>an</strong>ce, A. E., 727 (2-3a); 769 (1-42); 810 (1-4b). (1-25); 3622 (1-26), 3938 (2-1Oa); 4126 (1-26); 4679,<br />
LBB (='s L<strong><strong>an</strong>d</strong>s Bosbeheer Suriname), 8000 (2-17); 4740, 4746, (1-19); 5858 (2-la); 6201, 6225 (2-4);<br />
8161 (2-23); 8316 (2-3c); 8976 (2-1 la); 9199, 9424, 6249 (2-7); 6267 (2-25); 11017 (2-17); 11027 (2-23).<br />
9843 (2-10a), 11017(2-17); 11027 (2-23); 11747 (1- Macbride, J. F., 5446 (2-4); 5447 (1-4b); 5594 (1-32).<br />
20); 14296 (2-17); 16031, 16310 (1-4a).<br />
Macedo, R., 55753 (2-17).<br />
Leeuwenberg, A. J. M., 11784 (2-3a).<br />
Madison, M. T. et al., 4764, 7101 (1-19).<br />
Lehm<strong>an</strong>n, F. C., BT-694 (1-10); 948 (1-41); 5606 (1- Magalhaes, G. Mendes, 733 (2-la).<br />
42).<br />
Magnago,<br />
Leitao Filho, H. P., 685, 686 (1-25).<br />
Leite, J. E., 413, 892 (1-25).<br />
Lem<strong>an</strong>, D. S. (herb.), s.n. (1-4a, 1-20).<br />
Lemee, A. M. V., s.n. (1-4a); s.n. (2-3a, 2-3c).<br />
Lems, K., s.n. (1-25).<br />
Leng, H., 177 (1-26).<br />
Lent, R. W. (et al.), 349 (1-42); 2004 (2-3d); 2327 (2-<br />
23); 2520, 2535 (1-42); 2538 (1-34); 3279, 3388 (1-<br />
42); 3547 (2-3d); 5320 (1-42).<br />
Le6n, J., 3501 (2-3d).<br />
Leprieur, F. R., 141 (2-lOa); s.n. (1-4a, 1-16, 1-21,<br />
2-17).<br />
Lescure, J. P., 343 (2-1 la); 422 (2-23); 2075 (2-5).<br />
Liesner, R. L. (et al.), 417A (1-34); 515, 1309 (1-6),<br />
4105 (2-13a); 8750 (1-31); 9443 (1-42); 10944 (2-<br />
3d); 12394 (2-lb); 16175 (2-13b); 16299 (2-3a);<br />
22180 (1-11); 24462 (2-la).<br />
Lima, A., see Lima, D. de Andrade<br />
Lima, D. de Andrade (et al.), 20795 (1-25); 58-3185<br />
(2-2); 67-4983 (2-la).<br />
Lima, R., 21 (2-lOa).<br />
- et al., 58046 (2-la).<br />
Maguire, B. (et al.), 4945A (2-1 la); 23449 (1-26); 23867,<br />
23942 (1-1); 24588 (1-26); 29408 (1-35); 41620 (1-<br />
40); 46879 (1-3); 47049 (1-1); 53517 (1-4a); 56511<br />
(2-1 la); 60333A (2-3a).<br />
Maia, L. A. et al., 497 (2-4).<br />
Malme, G. O. A., 826, 1428 (1-25).<br />
Marc<strong>an</strong>o-Berti, L. (et al.), 471, 611 (2-la); 238-979 (2-<br />
3d); 84-2-77 (1-4b); 982-154 (2-3d).<br />
Marinho, L. R., 273 (2-la); 348 (1-4b).<br />
Martin, J., s.n. (1-20, 2-la, 2-3c, 2-1 la).<br />
Martinelli, G., 150 (1-25).<br />
Martius, K. F. P. von, 2620 (2-2); 2673 (1-28); s.n. (1-<br />
28, 1-42, 2-la, 2-2, 2-3a, 2-4, 2-7, 2-13a, 2-15).<br />
Mathews, S., 2060 (1-42).<br />
Matuda, E., 446, 2020 (1-36); 3300 (1-30).<br />
McD<strong>an</strong>iel, S. (et al.), 2666 (1-31); 16092 (1-4b); 16344<br />
(1-40); 20362 (2-15); 20403 (2-2); 20438 (2-7); 20555<br />
(1-40).<br />
McDonagh, J. F. et al., 114 (1-15).<br />
McPherson, G., 7594 (1-6); 11510 (2-1 lb).<br />
Medina, E., 379 (1-40).
202 Flora Neotropica<br />
Melinon, E., 18 (1-20); 172 (1-11); 457 (2-11a); s.n.<br />
(1-20, 2-1 la).<br />
Mello Barreto, H., 1795 (1-13).<br />
Mello, F., 55.388 (2-13a); 55.444 (2-1 la); 57509 (2-<br />
la).<br />
Mendez, R., 15 (1-6).<br />
Meneces, E. (et al.), 338 (2-4); 613, 647, 725 (2-23);<br />
759 (2-la).<br />
Mennega, A. M. W., 271 (1-6); 521 (1-20).<br />
Metcalf, R. D. et al., 30093 (1-42).<br />
Mexia, Y., 1872 (1-36); 5115 (1-25); 6573 (1-42).<br />
Meyer Drees, E., 5595 (2-3a).<br />
Meyer, D. G., 403 (2-23).<br />
Miers, J., 2714, 3792, 3858 (1-25); 6201 (2-la); 6257<br />
(2-4); 8172 (2-la); s.n. (1-25).<br />
Mir<strong><strong>an</strong>d</strong>a Bastos, A., 2051 (2-la); 2220 (2-23).<br />
Mir<strong><strong>an</strong>d</strong>a, F., 1737, 6164, 6273, 7180 (1-36).<br />
Molina R., A. et al., 1963 (2-3d); 17694 (1-34); 18227<br />
(2-3d); 18289, 30747 (1-42).<br />
Monsalve B., M., 1176 (1-46); 1259 (1-4b).<br />
Montaldo, P., 3435 (1-42).<br />
Monteiro da Costa, R. C., 90 (1-42); 92 (2-17).<br />
Monteiro, O. P. (et al.), 309 (2-20); 477 (2-la); 1119<br />
(2-23); 1295 (2-la).<br />
Mora, L. E., 2295 (2-3e).<br />
Morales, -, 38 (1-41).<br />
Moreira, A. S., 46 (1-25).<br />
Moreno, P. P., 23629 (1-18).<br />
Moretti, C., 102 (2-la); 836 (1-20); 837 (1-4a).<br />
Mori, S. A. (et al.), 5360 (1-34); 8583 (2-17); 8662 (2-<br />
la); 8864 (1-26); 10408 (2-4); 11025 (2-lOa); 11026<br />
(2-2); 11046 (2-la); 11655 (1-33); 14940 (2-19);<br />
15127, 15342 (2-13e); 15350 (2-3c); 15776 (1-40);<br />
16046 (2-la); 17185 (2-19); 17478 (2-1la); 17672<br />
(2-13e).<br />
Morillo, G. et al., 7384 (2-4).<br />
Mosen, H., 2614, 2941 (1-25).<br />
Mota, C. Domiao A. de (et al.), 154 (1-39); 680 (2-20);<br />
740 (2-la); see also Domino, C.<br />
Mota, M. G. et al., 1040 (2-5).<br />
Mutis, J. C., 646, 649, 4558 (1-42).<br />
Nadeaud, J., s.n. (1-25).<br />
Nascimento, R. J. (et al.), 262 (2-2); 303, 387 (2-la);<br />
66338 (2-3a); 66396 (2-7).<br />
Nee, M. (et al.), 8822 (1-34); 9281 (1-6); 30877 (2-<br />
13a).<br />
Neill, D. (et al.), 1653, 1919, 1984, (1-42); 3389 (2-<br />
23); 7044 (2-4); 7082 (2-la); 7179 (1-42); 8158 (2-<br />
13a); 8289 (1-22).<br />
Nelson, B., 680, 681 (2-la); 705 (2-4).<br />
Nevers, G. de (et al.), 4320 (1-6); 4697, 4789 (1-11);<br />
4929 (2-1 lb); 4982 (1-15); 5062, 5502, 6152 (1-6);<br />
6186 (1-18); 6194 (1-6); 6997 (2-23); 7170, 7542 (2-<br />
3d).<br />
Nishimura, A., 39 (2-1a).<br />
Nufez, O. V., 2928 (1-42).<br />
Nuiiez, P., 1823, 5761, 5946 (2-23); 5949 (2-la).<br />
Occhioni, P. (et al.), 5864, 5917 (1-13); 8228, 35333<br />
(1-25).<br />
Oersted, A. S., 14317 (1-42).<br />
Oldem<strong>an</strong>, R. A. A. (et al.), 207 (1-1); T.243 (1-4a);<br />
T.243a (1-21); 366 (1-4a); B.644 (2-10a); B.1 176 (1-<br />
1); 1179 (2-la); 1243 (2-1 la); B.1324 (1-1); B.1338<br />
(1-20); 1619 (1-1); B. 1946 (1-21); 2030 (2-3c); 2476<br />
(1-1); 2504(2-23); B.2538(1-4a); 2821 (2-23); B.2861,<br />
3062 (1-21); 3115 (2-3c); B.3243 (1-1); B.3420 (1-<br />
4a); B.3786A (2-2); B.4108, B.4132 (2-la).<br />
Oldenburger, F. H. F. et al., 1120, 1406 (2-3c).<br />
Oliva, F., s.n. (1-42).<br />
Oliveira, E. de, 985 (1-20); 1974 (2-17); 2597 (2-10a);<br />
2764 (1-35); 3071 (2-1Oa); 3550, 3599 (1-20); 3627<br />
(1-28); 3682 (1-20); 3736 (2-1 la); 3915 (2-17); 3936<br />
(2-13e); 3954 (1-20); 4047 (2-1 la); 60534 (2-5).<br />
011gaard, B. et al., 35068 (1-49); 57061 (2-3a).<br />
Ortega, -, s.n. (1-34).<br />
Ortiz, R. T., 200, 1030, 1810 (1-30).<br />
Palacios, W. (et al.), 1874 (2-24), 2059 (2-3a); 6387<br />
(1-27).<br />
Pariona, W. et al., 954 (2-13c).<br />
Parker, C. S., s.n. (1-4b).<br />
Peck, M. E., 497 (1-30).<br />
Pennington, T. D. (et al.), 12244 (2-24), P.22647 (2-<br />
7).<br />
Pereira, A. B., 39 (1-25).<br />
Pereira, L., 5670 (1-25).<br />
Perrottet, G. S., s.n. (1-4a).<br />
Persaud, A. C., 116 (1-4b); 222 (1-26).<br />
Pessoal do Horto Florestal, 650, 651 (2-la); s.n. (1-<br />
25).<br />
Peters, C., 187 (2-4).<br />
Petilon, -, 154 (1-4a).<br />
Philipson, W. R. et al., 1914 (1-42); 2086 (2-3a); 2095<br />
(1-42); 2133 (2-lOb); 2134 (2-23).<br />
Pinheiro, G. S., 1716 (1-25).<br />
Pinsley, H. V., 405 (2-4); 556 (2-3A).<br />
Pires, J. Murqa (et al.), 502, (2-39); 507, 542, 587 (2-<br />
3a); 805, 830 (1-4b); 6663 (2-la); 7015 (2-2); 10123,<br />
10758, 10794, 10809 (2-lOa); 12570, 15088 (2-la);<br />
16461 (1-42); 16547 (2-1a); 50419 (2-23); 50631 (1-<br />
1); 50656 (1-2); 50864 (1-20); 50913 (1-4a); 51193<br />
(2-13e); 51247 (2-23); 51390 (2-17); 51409 (1-20);<br />
51624 (2-3c); 51625 (2-13e); 51676 (2-10a); 51736<br />
(1-40); 51882 (2-2); 52656 (1-20).<br />
Pires, O. et al., 248 (1-42).<br />
Pittier, H. (et al.), 2286 (2-3e); 3892 (1-42); 4386 (1-<br />
6); 8404 (1-42); 11166 (1-34); 12105 (2-3d); 12124,<br />
12142, 12144, 12288, 13985 (1-42); 15641 (2-1b);<br />
15733 (1-42); 16163 (2-3d); s.n. (1-42).<br />
Plowm<strong>an</strong>, T. C. (etal.), 2362 (1-42); 2919 (1-25); 6555<br />
(1-28); 6903 (2-4); 6906 (2-5); 11179 (1-32); 11180<br />
(1-42).<br />
Poeppig, E., 1881 (2-la); s.n. (1-28, 1-42, 2-la, 2-3c).<br />
Pohl, J. E., s.n. (1-25, 1-28).<br />
Poiteau, P. A., s.n. (1-20, 2-3c, 2-3a, 2-1 la, 2-3a, 2-3c,<br />
2-1 la).<br />
Poveda, L. J., 951 (1-34).<br />
Pr<strong>an</strong>ce, G. T. (et al.), 1379 (2-3a); 1548 (2-2); 1720 (1-<br />
20); 1777 (2-la); 2153 (1-4c); 2180 (2-1 la); 2306 (1-<br />
42); 2469 (2-2); 2754 (2-3a); 3253 (2-7); 3260, 4438<br />
(2-la); 4588 (1-4b); 6253 (2-la); 6292 (2-13b); 6419<br />
(2-4); 6718 (1-4b); 6969 (1-31); 7160 (2-17); 7371<br />
(1-32); 7503 (2-4); 7530 (1-42); 7670 (2-la); 7792<br />
(2-23); 7907 (2-lOb), 8378 (2-23); 8670 (2-la); 9054<br />
(1-35); 9679 (1-11); 9857, 9952 (2-13b); 10089,10124<br />
(2-23); 10231 (2-5), 10233 (2-la); 10348 (1-42);<br />
10836 (2-la); 11066, 13438 (1-42); 13579 (2-la);<br />
13582-A (2-3a); 14029, 14146 (1-31); 14795 (2-7);<br />
14976 (1-38); 15039 (1-34); 15366 (2-3a); 15393 (2-
Exsiccatae<br />
20); 15764 (1-40); 15791 (2-4); 17242 (2-9); 17847 Ruiz-Ter<strong>an</strong>, L., 423 (1-42).<br />
(1-38); 18032 (1-31); 18204 (2-23), 18240 (2-5); Rusby, H. H., 1599 (2-4).<br />
18250 (2-3a); 19878 (2-13b); 20454 (2-20), 21011 Ryl<strong><strong>an</strong>d</strong>s, A. B., 15 (2-2); 25 (2-23); 30 (2-13b); 52 (2-<br />
(2-2); 22708 (2-13b); 22720 (1-20); 22729 (1-31); 3a); 59 (2-2).<br />
22740 (1-20); 22803 (2-la); 22836 (1-20); 22859 (2- Sabatier, D. R. L., 501 (2-1 la).<br />
16); 22864 (2-20); 22999 (1-35); 23776 (2-7); 23846 Sagot, P. A., 517, 970, 972 (2-17); 990, 990bis (2-1 la);<br />
(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).<br />
(1-42); 25167, 25336 (2-2); 25337 (2-10a); 25363 (2- Sam<strong>an</strong>iego, A. et al., 92 (1-42).<br />
2); 25530 (2-20); 25748 (1-35); 26375 (2-la); 26402 S<strong><strong>an</strong>d</strong><strong>with</strong>, N. Y., 290, 302, 394 (1-26).<br />
(1-42); 58783 (1-20); s.n. (2-5).<br />
S<strong>an</strong>tos, M. R., 172 (2-23); 455 (2-lOa).<br />
Pr6vost, M. F. et al., 738 (1-4a); 982, 983 (2-3c). S<strong>an</strong>tos, T. S. dos, 2035 (2-la).<br />
Proctor, G. R., 27080 (1-18); 27336 (1-42); 30273 (1- Sastre, C., 2274 (2-4); 4350, 4579 (2-3c); 5500 (1-1);<br />
30).<br />
6130 (1-20); 6417 (1-4a).<br />
Pruski, J. et al., 3235 (2-13b); 3239 (2-20).<br />
Sawada, M., 21 (2-1 Ob).<br />
Purdie, W., s.n. (1-14).<br />
Schipp, W. A., 127 (2-3d); 999 (1-30).<br />
Purpus, C. A., 5996, 11161, 11162, 11182, s.n. (1-36). Schnee, L., 1337 (1-42).<br />
Rambo, B., 426, 27092, 37782, 38427, 39413, 39780, Schomburgk, R., 118, 167, 287, 876, 1366, s.n. (1-26).<br />
41173, 44743, 45136, 45904, 46145 (1-25). Schubert, B. G. et al., 748 (2-3d).<br />
Ramirez, C. R., 1084 (1-31).<br />
Schultes, R. E. (et al.), 3343, 3826, 3862, 3901 (2-4);<br />
Ramos, J. et al., 204 (2-20).<br />
6693 (1-8); 6726 (1-28); 6805, 8175 (2-4); 8269 (1-<br />
Rauh, W., P-1609 (2-23).<br />
8); 8380 (1-28); 8976 (1-4b); 9685, 12985 (1-31);<br />
Reineck, E. M. et al., 446 (1-25).<br />
13589(1-40); 13774(1-31); 14148(2-23); 15324(1-<br />
Reitz, R. (et al.), 975, 1688, 1912, 2008, 2027, 2913, 31); 15973(2-5); 16009(2-20); 16112(1-31); 16334,<br />
3809, 4176, 4692, 4965, 5030, 5611, 8294 (1-25). 16367 (1-42); 17615 (2-3a); 18969, 19758 (1-4b);<br />
Renteria, E. et al., 45, 2093 (2-12a).<br />
19835 (2-5); 24547 (2-4).<br />
Revilla, J., 805, 1099 (1-28); 1260 (1-4b); 1818 (2-5); Schultze-Rhonhof, H., 1953 (1-19); 2783 (1-11); 2943<br />
1847 (1-31); 1966 (2-7); 2387 (1-39); 3387 (1-28). (1-9); 3006 (1-11); 3020 (1-14).<br />
Reyno, R. N., 37 (2-13a).<br />
Schulz, J. P. (etal.), 7787 (1-20); 8000 (2-17); 8161 (2-<br />
Ribeiro, B. G. S., 612 (1-11); 665 (1-43).<br />
23); 8316 (2-3c); 8976 (2-1 la); 10344a (1-1).<br />
Richard L. C. (herb.)., s.n. (1-20, 2-la).<br />
Schunke, C., 416, A-95 (2-23).<br />
Riedel, L. (et al.), 1 (1-25); 2 (2-la <strong><strong>an</strong>d</strong> 2-lOa); 7 (1- Schunke, J. M., 130 (1-28); 371 (1-40).<br />
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);<br />
(1-25, 2-la).<br />
3309, 3552 (2-la); 6485 (2-4); 7415 (2-la); 7729 (2-<br />
Riera, B. J. J.-L., 206 (2-3c); 301, 571 (2-17); 573 (2- 10b); 8487 (2-23); 8666 (1-4b); 10650 (2-lOb).<br />
10a); 714 (2-la); 723 (2-13e); 923 (2-la).<br />
Schwacke, W., 560 (2-15); 1072, 6783, 8418 (1-25);<br />
Rimachi, Y. M., 359 (1-32); 1943 (1-28); 2297 (1-40); 10393 (2-la); 11895 (1-17).<br />
2724 (2-5); 2725 (2-9); 2765 (2-5); 3256 (2-15). SEF (=Studies of Ecuadori<strong>an</strong> Forest), 8532 (2-6); 8617<br />
Rizzini, C. T., 415, 1114 (1-25).<br />
(2-23); 8764 (2-4); 8823 (2-13c); 8930 (2-4); 9033,<br />
Roa T., A., 251 (2-unnamed); 252 (2-13a).<br />
9159 (2-23); 10366, 10378 (2-spec.).<br />
Rodrigues, C. R., 944 (2-5).<br />
Sehnem, A., 7993 (1-25).<br />
Rodrigues, G. et al., 877 (1-17).<br />
Sello(w), F., 241 (1-25); s.n. (2-2).<br />
Rodrigues, W. A. (et al.), 884 (2-23); 1415 (2-7); 1424 Shakaim, S., RBAE-26 (2-23).<br />
(2-23); 1434 (2-13b); 2092 (2-2) = 2892 (2-2); 2984 Sh<strong>an</strong>k, P. J. et al., 4294 (1-42).<br />
(2-1Oa); 2986, 2987 (2-13e); 3267 (2-2); 4237 (2-3a); Shattuck, O. E., 260, 522, 525 (2-3e); 832 (1-42).<br />
4585 (2-2); 5266 (1-31); 5449 (2-20); 7069, 7076 (2- Shepherd, D., 364 (2-23); 376, 379 (2-3a).<br />
7); 7081,7292 (2-20); 7532 (2-23); 8065 (2-13); 8066 Shima, D., 14 (2-20).<br />
(2-5); 9204 (2-la); 9206 (2-3a); 9219 (2-la); 10082 Sigueira, R., 8808 (1-20).<br />
(2-17); 10182 (2-23); IG1-2A-70 (2-5).<br />
Silva, A. S. Lima de et al., 215 (2-13d); 476 (2-8).<br />
Rodriguez, H., 399 (1-42).<br />
Silva, J. F., 69, 211 (1-20); 227 (1-4b).<br />
Rodriguez, J. V., 2929 (1-42).<br />
Rohr, J. B. von, 151<br />
Silva, M., 354 (2-lOa).<br />
(1-4a).<br />
Rombouts, H. E., 803<br />
Silva, M. Barbosa da, 130 (2-lOa).<br />
(1-4a).<br />
Romero, F., 548 Silva, M. F. et al., 466<br />
(1-42).<br />
(1-31); 742 (2-20); 869 (2-lOa);<br />
Romero-Castaieda, R., 3647 (2-4); 3840 (2-5); 3848<br />
903 (2-23); 2166 (1-4a); 2199 (2-13b); 2423 (2-2);<br />
(2-20); 4211 (1-4b); 4715 (2-12a); 5283 (1-19); 5484<br />
2591 (1-39); 2623 (2-la); 2712 (1-4a).<br />
(2-3e); 5512 (1-10).<br />
Silva, M. G. et al., 3345 (1-40); 3368 (1-4b); 4326,<br />
Rooden, J. v<strong>an</strong> et al., 258 (1-4b); 699 (1-10); 700 4738<br />
(1- (2-23); 4750 (2-la).<br />
5).<br />
Silva, N. T. (et al.), 921, 949, 995 (2-1 la); 1207, 1321,<br />
Rosa, N. A. (et al.), 296 (1-11); 339 (1-43); 1091 (2-<br />
1380 (1-20); 1411, 1412 (2-3c); 3013 (1-28); 4757<br />
5); 1126 (2-17); 1158 (2-lOa); 1159 (2-5); 1265 (2- (1-10); 60670 (1-43).<br />
23); 2053, 2874 (1-40).<br />
Skutch, A. F., 2023 (1-36); 4083 (2-3a); 4225 (2-3d);<br />
Rowlee, W. W. et al., 856 (1-42).<br />
4256, 4262, 4267, 4612 (1-42); 4740 (2-20); 57832,<br />
Ruiz, H. & J. Pavon, 2 (1-32); 3 (1-42); 4 (1-4b); 5, 6, 58862 (2-3a).<br />
7 (1-32); s.n. (1-4b, 1-32, 1-42, 2-lOb).<br />
Smith, A., 1632 (1-29); 1663, 1650 (1-34).<br />
203
204 Flora Neotropica<br />
Smith, A. C., 2731 (2-13d); 2753, 2799 (1-35); 2845 3d); 122390, 125857 (2-3a); 129163 (2-4); 129733<br />
(2-1a); 3564 (2-3c).<br />
(2-unnamed); 131962 (2-1 la).<br />
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);<br />
unnamed); 5134 (2-10b); 5300 (2-4).<br />
319 (2-17).<br />
Smith, L. B. et al., 12307 (1-25).<br />
Sucre, D., 1783, 3633, 4989, 5209, 5330, 7878 (1-25);<br />
Smith, R. F., 3221 (2-lb); 3313 (1-42).<br />
8303 (2-lOa); 8684 (2-2); 8690 (1-25).<br />
Smith, S. F. et al., 695 (2-3a).<br />
Sugden, A., 415 (1-42).<br />
Snedaker, S. C., C-39, D-88 (1-30).<br />
Sytsma, K. J., 4018 (2-3d); 4024, 4225 (1-6).<br />
Sneidern, K. von., s.n. (2-4).<br />
Tamayo, F., 3370 (2-lb).<br />
Sodiro, A., 193/12 (1-41).<br />
Teixeira, L. O. A. (et al.), 61 (2-3a); 849 (1-4B).<br />
Soejarto, D. D., 327, 856 (1-42); 2917 (2-23); 3385 (2- Terceros, W., 1400 (2-la).<br />
3e); 3489, 4063 (2-23); 4283 (2-1 lb).<br />
Tessm<strong>an</strong>n, G., 3054 (2-4); 3416 (1-42); 3922 (1-32);<br />
Solomon, J. C. (et al.), 9254 (2-23); 12655 (2-la). 4236 (2-3b); 4642 (2-la); 4673 (1-39); 4696 (1-42);<br />
Soria S., M. A., 21 (2-23).<br />
s.n. (1-25); 5364 (2-9).<br />
Sparre, B., 14822 (1-42).<br />
Teunissen, P., 16031 (1-4a).<br />
Sperling, C. R. et al., 6430 (2-23).<br />
Thomas, W. W., 3349 (2-13b); 3370 (2-3a).<br />
Spichiger, G. et al., 1995, 1996, 1997, 1998, 1999 (2- Tillett, S. S. (et al.), 45869 (1-43); 671-33 (1-3 1); 49495<br />
25).<br />
(2-1 la).<br />
Splitgerber, F. L., 448 (1-4a).<br />
Tomas, Bro., 1923 (1-42).<br />
Spruce, R., 951, 1219 (2-13a); 1509 (1-4a); 1608 (1- Tonduz, A., 9520 (1-42); 12930 (2-3d); 13280 (1-42).<br />
42); 2023 (2-4); 2260 (1-4b); 2498 (1-31); 2616 (1- Toro, R. A., 120C (1-14).<br />
40); 2865 (2-13b); 3176 (2-1 la); 3231, 3464 (1-40); Torres, M. J., 482 (1-32).<br />
3782 (1-38); s.n. (2-13a, 2-17).<br />
Toth, M. J., 482 (1-32).<br />
Stahel, G.et al., 732 (1-20).<br />
Tresling, J., 55 (1-1).<br />
Stahel, G., (Exp. Wilhelmina Gebergte) 196 (1-20). Tri<strong>an</strong>a, J. J., 860 (2-la); 861 (2-3a); 862 (2-4); 866 (1-<br />
Stahel, G., (Woodherb. Suriname) 82 (2-23); 123A (2- 34); 867 (1-42); 1846 (2-4); 1866 (1-34, 1-42); s.n.<br />
3c); 123B (2-17); 166 (2-3a); 166A (2-17); 166B (2- (1-42).<br />
3c).<br />
Troon, F. v<strong>an</strong> et al., 16310 (2-23); s.n. (2-la).<br />
St<strong><strong>an</strong>d</strong>en, -, 20 (1-42).<br />
Trucios, T., 9 (2-1Ob).<br />
St<strong><strong>an</strong>d</strong>ley, P. C. (et al.), 7081 (1-42); 7878 (2-3d); 8861, Trujillo, B., 6141, 8765 (1-42).<br />
12288 (1-42); 19904 (2-3d); 23766 (1-42); 27479, Tuerkheim, H. von, 4082 (2-3d); 8659 (1-30).<br />
27502 (2-3e); 30924 (1-42); 37121 (2-23); 37259 (1- Tunqui, S., 7, 94 (2-3b); 103 (1-42); 149 (2-3a); 217<br />
42); 39792 (1-34); 41170, 44550, 44968, 45182, (2-13a); 296 (2-10b).<br />
45661, 47791, 48594 (1-42); 48600 (2-23); 52900 Tweedie, J., 29 (1-25).<br />
(2-3d); 53230 (1-42); 53951 (2-3d); 53991, 53425, Tyson, E. L. et al., 4734 (2-3d).<br />
55155, 55445, 56623, 72288 (1-42); 72667 (2-3d). Ule, E. H. G., 991 (2-la); 1693 (1-25); 5265, 5266 (2-<br />
Steinbach, J., 1484 (1-32); 7567 (1-42).<br />
4); 5717 (1-42); 5718 (2-23); 5719 (2-6); 8836 (1-<br />
Stergios, B. et al., 11530 (1-35).<br />
4a); 8839 (2-17); 9314 (2-4); 9315, 9316 (1-32); s.n.<br />
Ster, W. L. et al., 423 (2-3d).<br />
(1-25).<br />
Stevens, W. D., 4839, 8323 (2-3d); 13340 (2-23); 13341 Uribe-Uribe, L., 364 (1-42).<br />
(2-3d).<br />
Vasquez A., R. et al., 17 (2-23); 167 (2-25); 809 (2-<br />
Steward, W. C. et al., 107, P.20156 (2-20); P.20255 13c); 2396 (2-la); 2491 (2-23); 2553, 2559 (1-42);<br />
(1-4a).<br />
2623 (2-25); 2630 (2-7); 2635 (2-5); 2539 (1-39);<br />
Steyermark, J. A. (et al.), 388, 487 (1-3); 507 (1-4b); 2719 (2-1 lb); 2852 (1-22); 2861 (2-23); 3965, 4201<br />
37602 (1-36); 38916 (2-3d); 39460 (1-30); 39618 (1- (2-7); 5111 (2-9); 5433 (1-31); 5745 (2-la); 5853 (2-<br />
42); 39934, 45663, 45981, 46103 (1-30); 52334 (1- 9); 6110 (2-23); 6156 (2-20); 6942 (1-42); 7104, 7522<br />
36); 54214(1-10); 54533 (1-42); 57872(2-13a); 57955 (1-31); 7535 (2-25); 7669 (1-4b); 8380, 8600 (1-22).<br />
(1-43); 60002 (1-4b); 60002a, 60364 (1-11); 60399 Veloso, H., 97 (1-25).<br />
(2-23); 60401 (2-3a); 60407 (2-1a); 60414 (2-23); Vellozo, -, 723 (2-la).<br />
60432 (1-4b); 60456 (1-43); 60665 (2-2); 60673 (1- Vellozo, H. P., 89, 723 (2-la); 1083 (2-lOa); 737 (1-<br />
11); 60800 (2-23); 75522 (1-11); 75540 (2-3a); 87307 33).<br />
(2-la); 87480 (1-4b); 89134 (2-la); 89452 (2-1 la); Vi<strong>an</strong>na, E. C. et al., 5465 (1-25).<br />
89457 (2-3a); 89506 (2-23); 90368 (2-20); 90408 (2- Vickers, W. T., 86 (2-5).<br />
1 la); 90583 (1-4b); 90620 (2-23); 90644 (2-3a); 90736 Vieira, M. G. de et al., 985 (2-23); 1004 (2-20).<br />
(l-4b); 92850, 92936 (2-22); 92991 (1-3); 94180 (2- Villiers, J. F., 2088, 2111 (2-23).<br />
3a); 95170(1-42); 95378 (2-lb); 95711 (2-la); 99829 Vogl, C., 1379, s.n. (1-42).<br />
(1-42); 99945 (2-3d); 102082, 102211 (1-42); 102874 Vreden, J., 11717(1-20).<br />
(2-unnamed); 103026 (2-13a); 104296 (1-11); 104348 Wachenheim, H., 18 (2-23); 232 (2-1 la); 271 (2-2);<br />
(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,<br />
la); 106118 (1-11); 106167 (1-31); 107064 (2-23); 2-23).<br />
113096 (2-3a); 114759 (1-4b); 115549(1-11); 116463 Warming, J. E. B., 1942, s.n. (1-25).<br />
(1-42); 116884 (1-4b); 116966 (2-1b); 119396 (2- Weaver, R. et al., 1677 (1-6).
Exsiccatae 205<br />
Weberbauer, A., 3639 (2-la); 3702 (1-42); 4472 (1-<br />
44b).<br />
Wedel, H. von, 1548, 1733, 2879 (1-42).<br />
Westra, L. Y. T., 47303 (2-2).<br />
Whitton, B. A., 179 (1-25).<br />
Williams, LI. (et al.), 3347, 3984 (2-4); 4179 (1-42);<br />
4627 (2-4); 4688 (2-lOb); 5349 (1-42); 9983 (2-lb);<br />
10071 (1-42); 12008 (2-lla); 12348, 12394, 13895<br />
(1-42); 14675 (1-4b); 14833 (2-13a); 15240, 15363<br />
(1-31); 15812 (2-5).<br />
Williams, L. O. et al., 17817, 18014, 24005, 26529 (1-<br />
42).<br />
Williams, R. S., 501 (2-lb); 983 (1-3d); 1560 (2-la).<br />
Wilson-Browne, G., 549 (2-23).<br />
Wilson, C. L., 127 (1-42); 381 (1-30).<br />
Wilson, P., 606 (2-3d).<br />
Woodson, R. E. et al., 1893 (1-42).<br />
Woodworth, R. H. et al., 445 (1-42); 606 (1-4b); 660<br />
(1-42).<br />
Woytkowski, F., 5926 (1-42); 7306 (1-32).<br />
Wurdack, J. J. (et al.), 286 (1-4b); 2169 (2-23); 43485<br />
(1-40).<br />
Yele, -, DR-23 (1-26b).<br />
Zarucchi, J. L., 2145 (2-4); 2744 (2-la); 3720 (2-3a);<br />
3272 (2-3e).<br />
Zik<strong>an</strong>, J. F., s.n. (1-25).<br />
Zuccarini, J. G. (herb.), 103 (1-25).<br />
INDEX OF VERNACULAR NAMES OF POUROUMA<br />
amapati (la) 127<br />
cucuva (4) 147<br />
ama'yrary (1Oa) 156<br />
dacha uvillos (5) 149<br />
ambafba do vinho (13a) 168<br />
dakamtazshiuya (1 la) 161<br />
ambauva m<strong>an</strong>sa (4) 147<br />
embauba da mata (la) 127<br />
ambauva do vinho (4, 13a) 147, 168<br />
embaubar<strong>an</strong>a (7) 152<br />
a'ilea (3c) 138<br />
garguaba (3a) 135<br />
ambaibochi (23) 189<br />
gr<strong>an</strong>boesipapaja (10a, la, 13e, 17, 23) 156, 161, 173,<br />
ambaibillo (23) 189<br />
180, 189<br />
amia-yek (la) 127<br />
guagay (3d) 141<br />
bochoa tsaha (4) 147<br />
guarumo colorado (5) 149<br />
boesipapaja (13e) 173<br />
guarumo de montafia (3d) 141<br />
bois c<strong>an</strong>on (la, 2, 3c, la, 13e, 17, 19, 23) 127, 132, guarumo macho (3d) 141<br />
138, 161, 173, 180, 181, 189<br />
guaumoutognac (13a) 168<br />
bois c<strong>an</strong>on male (10a, 23) 156, 189<br />
guruc<strong>an</strong>a (4) 147<br />
bois c<strong>an</strong>on sauvage (10a) 156<br />
hembra (3a) 135<br />
boroma (2, 3a, 3c, 10a, Ila, 13e, 17) 132, 135, 138, imbafba (4, 5, 10a) 147, 149, 156<br />
156, 161, 173, 180<br />
imbauba bengue (17) 180<br />
boroma ibeberob<strong>an</strong>a (2) 132<br />
imbauba br<strong>an</strong>ca (17) 180<br />
bospapaja(2, 3c, 10a, 1 la, 23) 132, 138, 156, 161,189 imbauba de cheirouvilha (17) 180<br />
bouchi papaie (23) 189<br />
imbauba do vinho (4, 13a) 147, 168<br />
bouchi papaye (13e, 17, 23) 173, 180, 189<br />
imbaiba m<strong>an</strong>sa (4) 147<br />
buruma (la, 3c) 127, 138<br />
imbaibar<strong>an</strong>a (la, 2, 3a, 5, 7, lla, 13b, 13c, 20, 23)<br />
caimar6n (4) 147<br />
127, 132, 135, 149, 152, 161, 169, 180, 189<br />
caimar6n de mico (3a) 135<br />
imbafbar<strong>an</strong>a br<strong>an</strong>ca (5) 149<br />
caimar6n silvestre (4) 147<br />
imbaubar<strong>an</strong>a folha peluda (5) 149<br />
caramuri (23) 189<br />
imbaubar<strong>an</strong>a vermelha (10a) 156<br />
cay-bari-cay (3a) 135<br />
imbauba-tor6m (la) 127<br />
cay-wari-cay-yek (3a, 22, 23) 135, 186, 189<br />
imbaubinha (8) 152<br />
chaparro de agua (la) 127<br />
itarar<strong>an</strong>ga (la, 10a) 127, 156<br />
chiricaba (13a) 168<br />
kaibarikei (5) 149<br />
chullachaqui (20) 184<br />
kai-wa-rei-kei-yek (23) 189<br />
chullachaqui bl<strong>an</strong>co (20) 184<br />
kaiwarikai (1 a) 161<br />
chullachaqui caspi (20) 184<br />
kalate (la) 127<br />
chumico (3d) 141<br />
kamoyuwa (10a) 156<br />
cirpe (3a) 135<br />
kaymbe'y (la) 127<br />
cirpe macho (3a, 1 b) 135, 162<br />
kukuma (13e) 173<br />
cocora(13a) 168<br />
kuluma (la) 127<br />
cocura (4) 147<br />
kulumasi (23) 189<br />
coiwaricoi-yek (23) 189<br />
kulumatelelel (3c) 138<br />
cormi (3a) 135<br />
lija (3d) 141<br />
cucura (2, 3a, 4, 5) 132, 135, 147, 149<br />
majagua (23) 189<br />
cucure (3a) 135<br />
male bois c<strong>an</strong>on (3c, 23) 138, 189
206 Flora Neotropica<br />
m<strong>an</strong>bospapaja (la, 17) 127, 180<br />
sirpe macho (3a) 135<br />
m<strong>an</strong>gab6 (3e) 144<br />
sirpo (3e, 12a) 144, 162<br />
m<strong>an</strong>o de leon (3d) 141<br />
sucufba (4) 147<br />
mapati (la, 4, 8, 13c, 15, 21) 127, 147, 152, 169, 177, sugkama(t) shuiya (la) 127<br />
186<br />
suia(4, lOb) 147, 159<br />
mapatir<strong>an</strong>a (2, 3a, 3c, 10a, la, 16, 23) 132, 135, 138, suir shuina (lOb) 159<br />
156, 161, 177, 189<br />
tamaoquare (3a) 135<br />
mapaty (13c, 15, 21) 169, 177, 186<br />
tambor (lb) 129<br />
mimpa shuiya washi shuina ( la) 161<br />
t<strong>an</strong>aribe (4) 147<br />
obija (1Ob) 159<br />
t<strong>an</strong>ta shuiya (3b) 137<br />
orumo de monte (3d) 141<br />
tarar<strong>an</strong>ga (lOa) 156<br />
otsepacho (3a) 135<br />
tarar<strong>an</strong>ga bl<strong>an</strong>ca (la) 127<br />
pacica (3d) 141<br />
tarar<strong>an</strong>ga vermelha (lOa) 156<br />
papaie (17) 180<br />
tentar shuina (3b) 137<br />
papaquillo (la) 127<br />
tinajero (3d) 141<br />
papaya del monte (la) 127<br />
torena (23) 189<br />
papaye (17) 180<br />
tourem (23) 189<br />
papaye apici (2, 1 la) 132, 161<br />
trumpet tree (3d) 141<br />
pau de jacu (la) 127<br />
tsaha (3a) 135<br />
pai shuina (3a, 13a) 135, 168<br />
tsakap suiya (la, 3a) 127, 135<br />
pai shuiya (13a) 168<br />
uhukamsuiya (10b) 159<br />
piraejo (3d) 141<br />
umbaouba (5) 149<br />
pourouma(3a, 3c, lla, 17) 135, 138, 161, 180 urumo bl<strong>an</strong>co (12a) 162<br />
puruma(3a, 3c, lla, 17) 135, 138, 161, 180 uva (3d, 4, 12b) 141, 147, 163<br />
puruma (4) 147<br />
uva de macaco (lOa) 156<br />
purumai (13b, 23) 169, 189<br />
uva de(l) monte (3d, 4, 13c) 141, 147, 169<br />
puruma-y (4) 147<br />
uva silvestre (4, 23) 147, 189<br />
sacha uvilla (4, 9, 13a, 15, 23) 147, 155, 168, 177, 189 uva medueda (3a) 135<br />
sacha uvillos (20) 184<br />
uvilla(la,4, 5, 9, 10b, 1a, 15,23) 127, 147, 149, 155,<br />
sadajii (4) 147<br />
159, 161, 177, 189<br />
sadha'fhi (3a) 135<br />
uvilla bl<strong>an</strong>ca (la, 23) 127, 189<br />
s<strong><strong>an</strong>d</strong>paper (la) 127<br />
uvilla l<strong>an</strong>uda (23) 189<br />
sa-ouro (10a) 156<br />
uvo (4) 147<br />
sarasara (3a, 5, 13b) 135, 149, 169<br />
wilaupiyua (13e) 173<br />
shew<strong>an</strong>toqui (4) 147<br />
yagrumo negro (lb) 129<br />
shuiya (la, 3b) 127, 137<br />
yagrumo-sunsun (10a) 156<br />
shuvija (23) 189<br />
yahal (3d) 141<br />
sirpe (3a, 12a) 135, 162<br />
yaryara (2, 3c, lOa, lla, 17) 132, 138, 156, 161, 180<br />
INDEX OF SCIENTIFIC NAMES<br />
Synonyms are in italics. Page numbers in boldface indicate primary page references. Page numbers<br />
<strong>with</strong> <strong>an</strong> asterisk (*) indicate pages <strong>with</strong> illustrations or maps.<br />
Allomeris 114<br />
Azteca 4<br />
Brosimum 66<br />
microcarpon 66<br />
Cebus apella 189<br />
Cecropia 2, 3, 9, 10, 15, 113, 115<br />
peltata 3<br />
<strong>Cecropiaceae</strong> 2, 3, 5, 15, 110, 113<br />
Clusia 4<br />
<strong>Coussapoa</strong> 2, 3, 4, 5, 6*, 7*, 8*, 9, 10, 15, 16, 17-28<br />
(keys), 112, 113, 114, 115, 194<br />
acutifolia 84<br />
<strong>an</strong>gustifolia 11*, 28, 29*, 58, 97<br />
apoda 53<br />
arachnoidea 10, 11*, 30, 31*, 50, 77, 82<br />
ar<strong>an</strong>eosa 99<br />
argentea 10, 11*, 30, 32*, 73<br />
asperifolia 4, 5, 7, 8*, 10, 11*, 15, 16, 33, 34 (key),<br />
99, 104<br />
subsp. asperifolia 10, 11*, 34, 35*, 63<br />
subsp. magnifolia 11*, 36, 37*, 39, 99<br />
subsp. rhamnoides 11*, 38, 39*<br />
batavorum 11*, 38, 40*, 99<br />
bolivi<strong>an</strong>a 99<br />
brenesii 86<br />
brevipes 7, 11*, 41, 42*<br />
cardonaei 36<br />
cayennensis 34<br />
chagresi<strong>an</strong>a 34<br />
chocoensis 11*, 41, 43*<br />
cinnamomea 11*, 44, 45*, 95<br />
cinnamomifolia 5, 10, 11*, 15, 16, 44, 46*, 86, 97
Index of Scientific Names 207<br />
contorta 5, 7, 11*, 41, 44, 47*, 48, 66<br />
cornifolia 87<br />
crassivenosa 10, 11*, 15, 48, 49*, 56<br />
cuneata 71<br />
cupularis 11*, 50, 51*, 77, 82<br />
curr<strong>an</strong>ii 4, 12*, 50, 52*, 58<br />
d<strong>an</strong>ielis 99<br />
dealbata 109<br />
dolich<strong><strong>an</strong>d</strong>ra 109<br />
donnell-smithii 99<br />
duquei 7, 10, 12*, 53, 54*, 77, 104<br />
echinata 12*, 53, 55*, 61, 108<br />
eggersii 99, 104, 105<br />
emarginata 109, 186<br />
embir<strong>an</strong>a 99<br />
fagifolia 71<br />
ferruginea 12*, 15, 56*<br />
ficina 34<br />
floccosa 12*, 52, 57*<br />
font<strong>an</strong>esi<strong>an</strong>a 66<br />
froesii 61<br />
fulvescens 14*, 105, 107*, 108<br />
glaberrima 12*, 58, 59*<br />
gr<strong><strong>an</strong>d</strong>iceps 99<br />
herthae 12*, 56, 58, 60*, 61, 108<br />
hirsuta 84<br />
hypochlora 36<br />
incomitata 50<br />
intermedia 75<br />
krukovii 110, 168<br />
laevigata 110<br />
latifolia 9*, 10, 12*, 15, 16, 61, 62*, 63, 71, 73, 82,<br />
86, 105<br />
var. obovata 61<br />
lawr<strong>an</strong>cei 99<br />
lehm<strong>an</strong>nii 99<br />
leopoldii 110<br />
leprieurii 12*, 15, 63, 64*, 66, 93<br />
longepedunculata 13*, 63, 65*, 77, 93<br />
macarenensis 101<br />
var. <strong>an</strong>tioquiensis 101<br />
macerrima 12*, 63, 66, 67*, 73, 92<br />
magnifolia 36<br />
m<strong>an</strong>uensis 13*, 66, 68*<br />
marti<strong>an</strong>a 99<br />
microcarpa 5, 7, 12*, 15, 63, 66, 69*, 73, 90<br />
microcephala 12*, 33, 63, 66, 71, 82, 90<br />
subsp. corifolia 89<br />
subsp. microcephala 72*<br />
mutisii 101<br />
napoensis 12*, 73, 74*, 84<br />
nitida 5, 10, 13*, 50, 75, 76*, 77, 82, 108<br />
nymphaeifolia 13*, 15, 61, 66, 77, 78*, 93<br />
obovata 61, 110<br />
oligoneura 86<br />
oligocephala 13*, 48, 77, 79*, 80, 93<br />
orthoneura 10, 13*, 30, 50, 63, 77, 80, 81*, 82<br />
ovalifolia 13*, 82, 83*, 84, 108<br />
pachyphylla 13*, 63, 73, 84, 85*<br />
p<strong>an</strong>amensis 99<br />
parviceps 7, 13*, 16, 44, 56, 86, 87*, 90, 108<br />
parvifolia 5, 12*, 73, 87, 88*, 89*, 92<br />
pittieri 110<br />
pl<strong>an</strong>itiensis 101<br />
plicata 110<br />
pr<strong>an</strong>cei 80, 82<br />
puberula 84<br />
purpusii 10, 13*, 16, 90, 91*<br />
rekoi 110<br />
rhamnoides 38<br />
rotunda 97<br />
ruizii 36<br />
scabra 13*, 90, 92*<br />
schottii 66<br />
var. l<strong>an</strong>ceolata 66<br />
var. longifolia 66<br />
schunkei 75<br />
setosa 84<br />
sprucei 13*, 15, 63, 66, 80, 92, 93*<br />
st<strong><strong>an</strong>d</strong>leyi 99<br />
steyermarkii 48<br />
subcrenata 99<br />
subinc<strong>an</strong>a 99<br />
tessm<strong>an</strong>nii 14*, 15, 44, 77, 93, 94*, 104<br />
tolimensis 14*, 108, 109*<br />
trinervia 4, 5, 10, 14*, 16, 44, 59, 95, 96*, 97<br />
valaria 108, 110*<br />
v<strong>an</strong>niifolia 5, 14*, 39, 97, 98*<br />
vellerea 99<br />
villosa 4, 5, 8*, 10, 14*, 15, 53, 77, 95, 99, 100*,<br />
110<br />
viridifolia 7, 10, 14*, 15, 16, 63, 82, 105, 106*<br />
var. tenuifolia 105<br />
volaria 14*<br />
warburgi<strong>an</strong>a 50<br />
williamsii 80<br />
Crematogaster 114<br />
Ficus 4, 110<br />
intermarginalis 110<br />
Moraceae 2, 3, 5, 15, 110<br />
Mus<strong>an</strong>ga 2, 9, 15, 113<br />
Myri<strong>an</strong>thus 2, 5, 9, 15, 112, 113, 115<br />
Poikilospermum 2, 3, 15, 113<br />
subgen. Ligulistigma 15<br />
Poulsenia 110<br />
armata 110<br />
<strong>Pourouma</strong> 2, 3, 5, 9, 10, 15, 110, 111, 113, 114, 116,<br />
117-121 (key), 194<br />
acuminata 113, 114, 116, 173, 175*, 176*, 193<br />
acutiflora 123<br />
albistipulata 166<br />
apaporiensis 160, 161<br />
forma macrophylla 160<br />
apiculata Benoist 168<br />
apiculata Mildbraed 168<br />
aspera 116, 133<br />
subsp. digitata 115<br />
aurea 186, 189<br />
bicolor 111, 112, 114, 115, 122, 123, 132, 132-133<br />
(key)<br />
subsp. bicolor 123,133, 134*, 143*, 149, 193, 194<br />
subsp. choco<strong>an</strong>a 115, 141, 142*, 143*, 181<br />
subsp. digitata 115, 137, 138*, 139, 143*<br />
subsp. scobina 115, 122, 123,139, 140*, 141, 143*,<br />
144<br />
subsp. tessm<strong>an</strong>nii 115, 136*, 137, 143*
208 Flora Neotropica<br />
bolivarensis 111, 113, 114, 115, 116, 176*, 186,<br />
187*, 189<br />
camarat<strong>an</strong>a 133<br />
cecropiifolia 110, 111, 113, 114, 116,139, 143*, 144,<br />
145*, 147, 194<br />
choco<strong>an</strong>a 141<br />
cinerascens 123<br />
crassivenia 139<br />
crassivenosa 133<br />
cuatrecasasii 186<br />
cucura 112, 114, 115, 116, 147, 148*, 157*, 193,<br />
194<br />
cuspidata 113, 114, 116, 149, 157*<br />
digitata 137<br />
edulis 144<br />
elliptica 114, 176*, 184, 185*<br />
essequiboensis 169<br />
ferruginea 111, 113, 114, 116, 176*, 177, 178*, 184,<br />
193<br />
folleata 186<br />
formicarum 112, 113, 114,115,116,152,153*, 155,<br />
157*<br />
fulginea 123<br />
garci<strong>an</strong>a 147<br />
gui<strong>an</strong>ensis 111, 114, 115, 116, 122, 123<br />
subsp. gui<strong>an</strong>ensis 122, 123, 124*, 126*, 128, 129,<br />
132<br />
subsp. venezuelensis 114, 122, 126*, 128, 129*<br />
herrerensis 115, 116, 171*, 177, 190, 192*, 193<br />
heterophylla 123, 155<br />
hirsutipetiolata 111, 114, 115, 116, 162 (key), 190<br />
subsp. hirsutipetiolata 162, 164*<br />
subsp. hispida 162, 165*<br />
hispida 162<br />
isophlebia 186<br />
jaramilloi 159<br />
johnstonii 141<br />
jussiae<strong>an</strong>a 156<br />
laevis 180<br />
lawr<strong>an</strong>cei 133<br />
longipendula 181<br />
maroniensis 172<br />
melinonii 5, 113, 114, 115, 116, 155, 159, 160 (key)<br />
subsp. glabrata 112, 115, 157*, 161, 163*<br />
subsp. melinonii 115, 156, 157*, 158*, 160, 193,<br />
194<br />
mildbraedi<strong>an</strong>a 123<br />
minor 3, 5, 109, 111, 112, 113, 114, 116, 176*, 186,<br />
189<br />
mollis 15, 114, 115, 116, 155, 155-156 (key), 160,<br />
193<br />
subsp. mollis 115, 156, 157*, 158*, 159, 160<br />
subsp. triloba 114, 115, 156, 157*, 193<br />
multifida 144<br />
myrmecophila 111, 112, 113, 114, 115, 116, 150,<br />
151*, 152, 157*<br />
napoensis 113, 114, 115, 116, 171*, 190, 191*<br />
oraria 111, 112, 114, 115, 176*, 180, 190<br />
ovata 111, 112, 113, 114, 115, 116, 176*, 177,181,<br />
183*<br />
palmata 123<br />
paraensis 194<br />
phaeotricha 1 111,113, 115,116,152,154*, 155,176*<br />
populifolia 173<br />
radula 123<br />
retusa 194<br />
sapida 144, 147<br />
saulensis 114, 115, 116, 176, 181, 182*, 184<br />
scabra 123<br />
scobina 139<br />
schultesii 133<br />
steyermarkii 130<br />
stipulacea 111, 112, 114, 115, 116, 173, 174*, 176*<br />
subplicata 186<br />
substrigosa 123<br />
subtriloba 123<br />
tergoscabra 149<br />
tessm<strong>an</strong>nii 137<br />
tomentosa5, 15, 110, 111, 112, 114, 115, 116,164,<br />
166 (key), 193, 194<br />
subsp. apiculata 115, 168, 171*, 173<br />
subsp. essequiboensis 115, 169, 171*<br />
subsp. maroniensis 115, 170, 171*, 172*<br />
subsp. persecta 113, 115, 169, 170*, 171*<br />
subsp. tomentosa 115, 166, 167*, 171*, 173<br />
tri<strong>an</strong>ae 194<br />
triloba Klotzsch 159<br />
triloba Trecul 156, 159<br />
ulei 194<br />
umbellata 186<br />
umbellifera 186<br />
uvifera 144<br />
velutina 114, 115, 116, 126*, 128, 129, 131*<br />
venezuelensis 128<br />
villosa 111, 176*, 177, 179*<br />
Schefflera 4<br />
Urostigma 10<br />
intermarginala 110<br />
Urticaceae 2, 3, 15