Organization for Flora Neotropica

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

Author(s): C. C. Berg, R. W. A. P. Akkermans, E. C. H. van Heusden
Reviewed work(s):
Source: Flora Neotropica, Vol. 51, Cecropiaceae: Coussapoa and Pourouma, with an Introduction
to the Family (Apr. 11, 1990), pp. 1-208
Published by: New York Botanical Garden Press on behalf of Organization for Flora Neotropica
Stable URL: http://www.jstor.org/stable/4393813 .
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FLORA NEOTROPIC

MONOGRAPH 51

CECROPIACEAE: COUSSAPOA AND

POUROUMA, WITH AN INTRODUCTION
TO THE FAMILY

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

. i ., """ - -- - -------------------
///A "^C^ -
TIOPIC

OF CANCER

NEOTROPICsAh

Publishedfor

Organizationfor Flora Neotropica

by
The New York BotanicalGarden
New York

Issued 11 April 1990

 Copyright ? 1990
The New York Botanical Garden

Published by
The New York Botanical Garden
Bronx, New York 10458
International Standard Serial Number 0071-5794

Library of Congress Cataloging in Publication Data

Flora neotropica. - Monograph no. 1 - New York: Published
for Organization for Flora Neotropica by the New York
Botanical Garden, 1968-
v.: ill.; 26 cm.
Irregular.
Each issue has distinctive title.
Separatelycatalogedand classifiedin LC before monographno. 40.
ISSN 0071-5794 = Flora neotropica.
1. Botany-Latin America-Classification-Collected works. 2. Botany-
Tropics-Classification-Collected works. 3. Botany-Classification-Col-
lected works. I. Organizationfor Flora Neotropica. II. New York Botanical
Garden.
QK205.F58 581.98'012-dcl9 85-647083
AACR 2 MARC-S
Libraryof Congress [8508]
ISBN 0-89327-352-x
 CECROPIACEAE:COUSSAPOAAND POUROUMA,
WITH AN INTRODUCTION TO THE FAMILY

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

CONTENTS
Abstract .................................................................. 2
Introductionto the Family (C. C. BERG) ....... ..................................... ............ 2
History and Circumscriptionof the Family ..... ................. ............................ 2
Descriptionof the Family ......... ........................................ .. .............. 3
Key to the NeotropicalGenera................................................ ............. 3
Genus Descriptionof Cecropia.... ......................................................... 3
Coussapoa(C. C. BERGANDR. W. A. P. AKKER NS)............................................ 4
Introduction ............................................................................. 4
Taxonomic History ....................................................................... 4
Morphology........... ........................ ........................................... 4
Anatomy (K. BoNSEN ) .................................................................... 8
Distributionand Ecology......................... ......................................... 10
SystematicPosition of the Genus........................................................... 15
SystematicArrangementof the Species...................................................... 15
Taxonomy ........ ........................ .............................................. 16
GenericDescription .................................................................. 16
Keys to the Species .................. ........ ......................................... 17
Treatmentsof the Speciesand Subspecies .... .................................... 28
Unnamed Collections .... ........................ .......... .......................... 108
Names Not Included.................................................................. 109
Pourouma (C. C. BERGANDE. C. H. VANHEUSDEN)
.............................................. 110
Introduction............................................................................. 110
Taxonomic History ....................................................................... 111
Morphology .............................................................................. 111
Phenology ............................................................................... 113
Pollination.................. ....
......................................................... 113
Dispersal ............................................................................... 114
Relations with Ants ....................................................................... 114
Distributionand Ecology ......... ...... ................................ ................... 114
SystematicPosition of the Genus.... ..................... .................................. 115
SystematicArrangementof the Species...................................................... 115
Uses .................................................................................... 116
Taxonomy ............................................................................... 116
GenericDescription .... .. ........................................................ , ... 116
Key to the Speciesand Subspecies...... ................................................ 117
Treatmentsof the Speciesand Subspecies ............................................. 122
Unnamed Collections ............. . ............................................ 193
Names Excluded ................................................................. .... 194
Acknowledgments ............................................................................ 194
LiteratureCited .............. .. ...... ................................... ..................... 194
Numerical List of Taxa ......... .............................................................. 196
List of Exsiccatae............................................................................. 197
Index of VernacularNames . ................................................................... 205
Index of ScientificNames ................................... .............................. .. .. 206

Arboretumand BotanicalGarden(ARBOHA),University of Bergen,N-5067 Store Milde, Norway.

2
Institute for Systematic Botany, State University of Utrecht, Heidelberglaan2, 3508 TC Utrecht, The
Netherlands.
3 Institute for Systematic Botany, State University of Utrecht, Heidelberglaan2, 3508 TC
Utrecht, The
Netherlands.
4Institute for Systematic Botany, State
University of Utrecht, Heidelberglaan2, 3508 TC Utrecht, The
Netherlands.
1
2 FloraNeotropica

ABSTRACT
Berg,C. C. (Arboretumand BotanicalGarden[ARBOHA],Universityof Bergen,N-5067
Store Milde, Norway), R. W. A. P. Akkermansand E. C. H. van Heusden (Institutefor
SystematicBotany, State University of Utrecht, Heidelberglaan2, 3508 TC Utrecht, The
Netherlands).Cecropiaceae:Coussapoaand Pourouma.FloraNeotropica51: 1-000. 1990.
The presentmonographcomprisesthe revisions of two of the three Neotropicalgeneraof
the Cecropiaceae:Coussapoaand Pourouma.For Coussapoa46 species are recognizedat
present;15 of them have been describedas new duringthe presentstudy.All or most species
are hemi-epiphytes.They have entire leaves and are dioecious. The small simple flowers
are bornein uni- to pluricapitateinflorescences.The staminateflowershave one stamen or
two or threeconnatestamens.The fruitsare small. The genusrangesfrom southernMexico
to south-easternBrazil and is absent in the West Indies. Most species are componentsof
lowlandrainforesthabitats.Few speciesarewide-spread,most specieshave relativelysmall
or even very small (known)rangesof distribution.The species are morphologicallyclear-
cut and most of them ratheruniform.For severalreasons,e.g., lack of staminatematerial,
the speciesare (provisionally)arrangedin alphabeticalsequences.Littleis knownabout the
biology of the genus. For Pourouma25 species are presentlyrecognized.Four taxa have
been describedas new during the preparationof this revision. All species are terrestrial,
usuallysmall to medium-sizedtrees. They have entire to palmatelylobed to partedleaves
and are dioecious. The pistillate flowers are pedicellate and are arrangedin repeatedly
branchedto subumbellateinflorescences.The small staminate flowers are pedicellateor
sessileand occur + loosely glomerateor in globoseheads.The stamensarefreein all species
but one. The fruits are relativelylarge.The genus rangesfrom CentralAmericato south-
easternBraziland is absent in the West Indies. Most of them occurin + disturbedand/or
+ open places in lowland rain forest. Besides a number of clear-cutand slightly variable
species the genus comprisesa numberof (very)variableand + complex species:P. bicolor,
P. cucura,P. guianensis,P. melinonii,P. mollis, and P. tomentosa.The formerthree and
the latterthreespeciesareoftendifficultto separatefromeachother(andsome otherspecies).
These six taxa and some othersshow a considerableplasticity,especiallywith regardto the
leaf shape, leaf dimension, and composition of the indument,and most prominentlyso in
the Upper Amazon Basin. Little is known about the biology of the genus. P. cecropiifolia
is in cultivationas a fruit tree.

INTRODUCTION TO genera: three (Cecropia, Coussapoa and Pou-

THE FAMILY rouma)that are Neotropical,two (Musangaand
Myrianthus)African,and an Asian-Australasian
The present monograph is comprised of re- genus (Poikilospermum).The majority of the
visions of two Neotropical genera of the Ce- members of the family constituted subfamily
cropiaceae,Coussapoaand Pourouma.To place Conocephaloideaein Engler'sMoraceae(Engler,
these revisions in a propersetting, there is pro- 1889), but Poikilospermumand some of the
vided first a history and circumscriptionof the species presentlyincludedin it were includedin
family, along with a brief discussionof the char- the Urticaceae.Chew Wee-lek (1963) proposed
acters that separatethe family from the related rankingthe four microspermousgenera(Cecro-
Moraceae and Urticaceae, a description of the pia, Coussapoa,Poikilospermumand Musanga)
family and the type genus, Cecropia,and a key under the Urticaceaeand leave the two macro-
to the Neotropical genera. spermousgenera(Myrianthusand Pourouma)in
the Moraceae.Corer (1962), however,regarded
all six generaas membersof the Urticaceae.The
History and Circumscription
of the Family creation of a new family appearedto be an ap-
propriateresolution to an unsatisfactorysitua-
The family was proposedby Berg(1978a) and tion.
accepted by Cronquist(1981). It comprises six Severalfeaturessuggestthat the Cecropiaceae
Introduction 3

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

to the Moraceae. Poikilospermumshows more radially-incised;venationpinnate,(sub)palmate,
urticaceousfeatures, including the typical urti- trinervate,or radial; stipules fully amplexicaul
caceouscystoliths,than the other five genera(cf. and connate.Inflorescencesin the leaf axils, usu-
Berg, 1989c;Bonsen & ter Welle, 1983). All gen- ally in pairs,unisexual,branched,with the flow-
era match the Urticaceae in the (sub)basalat- ers ? solitaryor clusteredin heads or spikes, or
tachmentof the ovule and the absence of milky unbranchedwith a singlehead or spike,bracteate
sap. They are, however, clearly different from or ebracteate.Staminateflowerswith 2-4 free or
the Urticaceaein the absenceof urticaceous(i.e., connate tepals; stamens 1-4; pistillode absent.
explosive)stamens.Such stamensare inflexedin Pistillate flowerswith 2-4 connate tepals; pistil
the bud and, at anthesis, bend suddenly and one, ovary free from the perianth, unilocular;
elasticallyoutward,throwingout the pollen. The ovule one, (sub)basally attached; stigma one,
African and Neotropical genera have a truly penicillateto peltate. Fruit an achene or some-
woody habit (cf. Bonsen& ter Welle, 1983). Oth- times (in Coussapoa) ? drupaceous,enveloped
er featureswhichcharacterizethe familyarestrict by a ? fleshy perianth;seed small, with endo-
dioecy,waterysap thatturnsblackafterexposure sperm,or largeand without endosperm;embryo
to the air, the common occurrenceof palmately- straight,cotyledons equal and flat or thickened.
incised leaves and, in certain species, radially- A pantropicalfamily with six generaand ap-
incised, peltate leaves, large stipules (connected proximately 180 species.
with the protectionof young inflorescences),and
adventitious aerial roots. While these roots are
manifest as stilt-roots in most genera, in Cous- Key to the Neotropical Genera
sapoa they are associated with the hemi-epi- 1. Laminapeltate,radially-incised. ...... Cecropia.
phytic habit and in Poikilospermumwith the 1. Laminabasallyattached,entireorpalmately-in-
climbing habit. cised.
The inflorescencesof the Cecropiaceaevary 2. Staminate flowerswith 1 or with2 or 3 con-
natestamens;pistillateflowerssessile,in glo-
from being branched, with a + loose arrange- bose(to ellipsoid)heads;fruitssmall;stipule
ment of the flowers(as in some staminateinflo- scarsusuallyascending........... Coussapoa.
rescencesof Pourouma),to unbranched,with a 2. Staminate flowerswith2-4 freestamens(or,
single globose flower head (as in some pistillate in P. napoensis,
3-4 connatestamens); pistil-
inflorescencesof Coussapoa),to a subumbellate lateflowerspedicellate(tosubsessile), solitary
orin non-capitateclusters; fruitslarge;stipule
inflorescence(as in Pouroumaminor),or clusters scarshorizontal................. Pourouma.
of spikes enveloped by a spathe (as in most Ce-
cropiaspecies).
In all Cecropiaceaethe fruitsare envelopedby CecropiaLoefling,Iter Hispan. 272. 1758, nom.
conserv. Type. C. peltata Linnaeus.
a ? fleshy perianth, and with regardto polli-
nation and dispersal,the Cecropiaceaeare more Trees, terrestrial,usually with stilt-roots; in-
similar to the Moraceaethan to the Urticaceae. ternodes usually hollow. Lamina peltate, radi-
The genus Cecropiashows the most derived ally incised, venation radial,petiole mostly with
charactersin the family, not only with regardto 1-2 trichilia(patchesof dense indument,usually
leaves and inflorescences,but also in the features
with trichomesformingMiillerianbodies)at the
connected with the adaptationto a mutualistic base. Inflorescencesdigitateclustersof spikes(or
relationwith ants, as well as in the peculiarde-a single spike), usually enveloped by a spathe
tachment (abscission) of the anthers (cf. Berg, until anthesis,interfloralbractsabsent. Perianth
1977b). tubular;stamens two; stigma penicillate to pel-
tate. Fruit small, dry, enveloped by a greenish
Description of the Family fruitingperianth.
Withapproximately80 speciesthroughoutthe
Cecropiaceae
Neotropics, with a distinct concentration of
Trees or shrubs,terrestrialor hemi-epiphytic, species in the Andean region.
with aerial roots, and watery sap turningblack Revisions of Coussapoa and Pourouma are
upon exposure.Leaves in spirals;lamina basally presentedsequentiallybelow.
4 Flora Neotropica

COUSSAPOA Engler'ssystem of the Moraceae(Engler,1889).

This subfamilywas transferredto the Urticaceae
Introduction by Corner(1962), but has recently been recog-
nized as a separatefamily Cecropiaceae(Berg,
Coussapoa,one of the three Neotropical gen- 1978a; Cronquist, 1981).
era of the Cecropiaceae,is a well-definedgenus.
The majorityof its species are arboreoushemi-
epiphytes,a peculiarlife-formsharedwith a few Morphology
othergeneralike Ficus(subg. Urostigma),Clusia, -
Habit. Coussapoaspeciesare shrubsor more
and Schefflera.The species are rathersimilar in small to medium-sized trees (up to
commonly
their ecology and show little variation in mor- ca. 20 m but they can sometimesform huge
tall),
phological characters.Knowledge of their biol- wide-crownedtrees up to 40 m tall, e.g., C. cur-
ogy is very limited. ranii and C. trinervia.On labels the height is
The poorstateof botanicalexplorationof many not alwayscorrectlystated,becausefor
probably
regionsof the Neotropics has adverselyaffected epiphytesthe host often appearsto be included.
the presentrevision, as is evident from the fact Most
that it has not proved possible to make a sys- Coussapoa species are hemi-epiphyticor
tematic arrangementof the species and from the issecondarilyterrestrial.Probablyonly C. trinervia
basically terrestrial.As in Ficus, Coussapoa
way in which the keys have had to be construct- species can be epilithic. The (hemi-)epiphytic
ed. In about one-thirdof the species only one of habit is evident in the presenceof aerial
the sexesis known.Manyspeciesareknownfrom growth roots.Theseaerialrootscan formbasketsof roots
only one or at most just a few specimens. More aroundthe stem of the host tree. Sometimes,the
than one-quarterof the species now known have aerial root consists of a main (vertical)
been recognizedas new during the preparation axis with at system
more or less regularlydistancedpairs
of this revision; many of them are based on re- of
horizontalbranchesclaspingthe trunkof the
cent collections. The increase in the number of host tree. In
general,hemi-epiphyticCoussapoa
speciessuggeststhat furtherexplorationwill lead species are not true (or
to the discovery of additional species. vigorous) stranglerslike
hemi-epiphyticFicus species.
Most Coussapoa species have straight, stiff
Taxonomic History branches.The smaller branches can be hollow
(widelyso in C. asperifolia,but narrowlyin sev-
Coussapoawas establishedby Aublet (1775), eralotherspecies,e.g., C. villosa).Whenbranch-
who describedtwo species from FrenchGuiana. es are hollow they are often inhabitedby biting
Names were added by Trecul (1847), Klotzsch ants (Azteca?)or stingingants (often in the case
(1847), Miquel (1853), and later on by Standley of C. villosa).These insects bore openingsto the
(1919, 1924, 1930, 1937a, 1940), Mildbraed interiorof the branches.Like (most?)other gen-
(1928, 1942), and Cuatrecasas(1951, 1956b), era of the Cecropiaceae,when wounded Cous-
among others,so that ca. ninety names were cre- sapoa species exude a (more or less mucilagi-
ated for thirty-fourof the taxa recognizedin the nous)waterysap whichturnsblackafterexposure
present paper. In the course of the preparation to the air.
of present treatment 15 new species were de- Indumentum.-Four types of trichomes can
scribed(Akkermans& Berg, 1982; Berg, 1983). be recognized:
Taxonomic, mostly floristic, treatments of the
genusareto be foundin the workof Trecul(1847), 1. Unicellulararachnoid(cobwebby)hairs.These
Flora brasiliensis(Miquel, 1853), Flora of Peru very thin, crinkled, interwoven hairs of dif-
(Macbride,1937), Floraof Panamai(Woodson& ferent lengths are mostly white, sometimes
Schery, 1960), Flora of Suririame(Berg, 1975), brownish.
and Flora Costaricensis (Burger, 1977). Thus, 2. Unicellularnon-arachnoid,thickerand most-
since the treatmentsby Trecul and Miquel, the ly moreor less straighthairsof differentlengths
genus has never been studied as a whole. and white, yellow, or brown.
The genus belongs to a group of genera that 3. Pluricellularhairs-dark or pale brown,short
constitute the subfamily Conocephaloideaein and subgloboseto longand moniliform.These
Coussapoa
hairs are very common on young leaves, sti-
pules, inflorescences,etc. In dry materialthey
are more or less granular(powdery)in ap-
pearance.
4. Pearl bodies (or glands)-hyaline, succulent,
more or less club-shaped, and confined al-
most entirely to the leaves. These trichomes
are found in specimens of Coussapoamicro-
carpaand C. villosacultivatedin greenhouses.
Pearlbodies are probablycommon in the ge-
nus, but are often not found in the field be-
causethey areeasilydetached;moreover,they
are probablyharvestedby ants (cf. O'Dowd,
1982).
As with many other genera of the Cecropi-
aceae, these four types of hairs are also often
encounteredin Coussapoaspecies. Many species
have a dense or rather dense indumentum. In
several species arachnoidhairs are lacking and
most of these species have sparse indumentum
as well.
Leaves.-The petiole in most species is rela-
tively long-up to half the length of the lamina.
There is a tendency towards shorteningof the
petiole (to one-tenthof the length of the lamina)
in species with derived (i.e., obovate, trinerved)
types of lamina.The relativelengthof the petiole
is not subjectto greatvariation as is the case in
related genera, such as Myrianthus and Pou-
rouma.
The length of the lamina varies from more
than 50 cm to less than 10 cm and more or less
parallelsthe change in the shape of the lamina
from cordiform,subreniform,or broadly ovate
to elliptic or oblong to obovate or subobovate;
at the same time, this correlateswith a gradation
in the leaf base from cordate to truncate to
roundedto obtuse to subacute.
The venationis pinnateand brochidodromous
with a more or less regularparalleltertiaryve-
nation in the intercostalarea.The venation may
tend towardsubpalmate(C. vannifolia),or may
be more or less clearly trinervate(C. trinervia,
C. cinnamomifolia),with this developmentbeing
combined with a change towards a (sub)obvate
lamina. In ovate leaves the basal pair of lateral
vein branchesis mostly branched.Especiallyin
large leaves, the other lateral veins show little
branching(being mostly only furcate);however,
in C. asperifoliathe otherlateralveins aremostly
branched.Branchingof the lateral veins is less
5
common in ellipticleaves, and in obovate leaves
normally there is none. Venation types in the
genus can be seen in Figure 1.
The ovate laminawith branched,basal,lateral
veins and a relativelylong petiole can be regard-
ed as the least derived type of leaf in the genus,
as it shows the closest resemblanceto leaves in
othergeneraof the family, especiallyPourouma.
The leaves of P. minorresemblethose of C. con-
tortaand the leaves of P. tomentosaand P. me-
liononiiarereminiscentof those of C. villosaand
C. nitida, respectively.The obovate, trinervate
lamina with a short petiole can be regardedas a
derived type of leaf. Anothertype of leaf, which
can also be regardedas derived, is that found in,
e.g., C. parvifoliaand C. contorta.This type of
leaf has an ellipticto subovatelaminawith many
unbranchedlateral veins, a very regularvena-
tion, and a relativelyshortpetiole.These features
are characteristicmore of the Moraceaethan of
the Cecropiaceae.
The indumentumof the leavesalso showssome
variationin denseness and occurrenceof differ-
ent types of hairs.The occurrenceof a dense and
varyingindumentumappearsto be weakly cor-
relatedwith primitivenessin other leaf charac-
ters.
C. contorta is distinct in having domatium-
like cavities in the axils of the lateralveins.
The stipulesareconnate,fullyamplexicauland
caducous, usually leaving oblique scars, char-
acteristicfor the genus. The length varies con-
siderablyin the genus.One can roughlyrecognize
three categories:long stipules (5-15 cm long or
longer),medium-longstipules(1-5 cm long)and
shortstipules(less than 1 cm long).Longstipules
give protectionfor a longertime to youngleaves
and developing inflorescences,but a correlation
between the type of inflorescenceor habit pref-
erence and the length of the stipules is not ap-
parent.
Inflorescences.- Coussapoa,like all othergen-
era of the Cecropiaceae,is dioecious. Monoe-
cious specimens are rare.
The inflorescencesoccur in pairs in the leaf
axils, a featurecommon in the Cecropiaceaeand
related families. The ramificationof the inflo-
rescenceis basicallysimilarto that usuallyfound
in Myrianthus(see: De Ruiter, 1976) and Pou-
rouma. The common pedunclebears at its apex
(at least) three branches, a pattern repeated in
secondaryand furtherramifications.However,
6 Flora Neotropica

Ni

3 4 5
FIG. 1. Schematicdrawingsof types of venation in Coussapoa:1, most lateralveins branched;2, 3, only
the basal pair of lateralveins branched;4, lateralveins unbranched,the basal pair reachingthe marginabove
the middle; 5, three-nerved.
Morphology
FIG. 2. Schematic drawings of types of pistillate inflorescences in Coussapoa.
FIG. 2. Schematicdrawingsof types of pistillateinflorescencesin Coussapoa.
reductionof the numberof branchesto two, re-
sultingin a (sub)dichotomousramification,is also
found, especially in the ultimate ramification,
and may occurdown to the primaryramification
as well. Pistillateinflorescencesare schematized
in Figure2.
The sessile flowers are arrangedin terminal
globoseheads.Solitary(often more or less abort-
ed) flowersmay occur elsewherein the inflores-
cence,but theirpresenceis mostlyconnectedwith
points of branching.Solitary flowersare rare in
pistillate inflorescences.
The staminateinflorescencesare normallyre-
peatedlybranchedand bearsmallterminalheads.
In pistillateinflorescences,however,the number
of flower heads is often reduced to one, due to
reductionof the ramificationand/or to fusion of
the flower heads. In some species, e.g., C. bre-
vipes, C. duquei, the flower heads are ellipsoid
to clavate and combined with a very short pe-
duncle.
The inflorescencesare bracteatein the major-
ity of the Coussapoaspecies.The bractsaremostly
confinedto the flowerheads, but sometimes oc-
cur elsewherein the inflorescence,as in the sta-
minate inflorescencesof C. tessmannii. In some
species,e.g., C. microcarpaand C. viridifolia,the
interfloralbracts are very small; they may also
be few in numberor sometimes even absent. In
C. parviceps,the only species in which the pis-
tillate flowers are connate (or cohering), inter-
floralbractsare absent in the pistillate inflores-
cence but presentin the staminate. This lack of
interfloralbractsis apparentlya derived state.
The interfloralbractsare more or less distinct-
ly spathulate,often with the lower partvery nar-
row (and stipe-like). Relatively large interfloral
7
bractscan be subpeltateor more or less cucullate.
Sometimes, these bracts are truly peltate--as in
C. contorta.Bracts outside the flower heads are
mostly scale-likeand ovate to oblong.
Staminate Flowers.-The small staminate
flowershave a tubularperianthwhich is 3-lobed
at the apex. The number of stamens is 3, 2, or
1. In 3- and 2-staminateflowersthe stamensare
fused, forming a stalk with, respectively, 6 (or
occasionally5) or 4 thecaeat the apex. The num-
ber of stamens is very constant and may prove
to be the (only?)basisfor subdivisionof the genus
and arrangementof the species. The length of
the filaments varies from about as long as the
perianthto more than twice as long as the peri-
anth, the differences being distinctive at the
species level.
Pistillate Flowers.-The small pistillate flow-
ers have a tubular perianth with a narrow (al-
most)entireopeningthatallowsthe stylethrough.
The flowersare free,but in C. parvicepsthe mar-
gins of the flattened apices of the perianths of
adjacent flowers cohere. The ovary is free and
the stigma is penicillateto subpeltate.
Staminateand pistillate flowersand their as-
sociated bracts are shown schematicallyin Fig-
ure 3.
Fruits and Seeds.-The small fruits are free
from the perianth. The pericarp consists of a
ratherthickand crustaceousendocarpand a very
thin to ratherthick, slightly fleshy to mucilagi-
nous layer. Thus, depending on the featuresof
the exocarp,the ellipsoid to ovoid, 1-3 mm long
fruitlets are achene-like or more or less drupa-
ceous. Fruits are depicted schematicallyin Fig-
ure 4.
In C. asperifolia,while the exocarpis present
8 FloraNeotropica

1 2 3 4 5 6 7 a ' 9.
FIG. 3. Schematicdrawingsof flowersand interfloralbractsin Coussapoa:1-3, staminateflowers, 1, with
three connate stamens, 2, with two connate stamens, 3, with one stamen; 4, pistillate flower; 5-9, various
interfloralbracts.

as a very thin (membranaceous)outer layer, the ochorous,often by frugiferousbirds, but also by

mesocarpis relativelythick and expandsin such
a way as to push the endocarpbody out of the
perianth,the apex of which becomes torn in the
process. Whetherthis phenomenon is confined
to C. asperifoliaor also occurs in other species
is not clear.At any rate,it is not generalin Cous-
sapoa,as previouslyassumed(Berg,1975, 1977a).
In many (or all?) species the fruiting perianth
becomes more or less juicy, and it is also often
colored-orange, yellow, or brown. Sometimes,
as in C. villosa, the perianth remains greenish,
at least the apical part.
Coussapoais one of the four microspermous
genera of the Cecropiaceae.In the genus mi-
crospermyis relatedto the (hemi-)epiphytichab-
it, as well as to the occurrenceof a mucilaginous
layer (cf. Berg, 1983b).
Phenology, Pollination, and Dispersal. -
Availableherbariumannotationssuggestthat in
general Coussapoa species flower (and fruit)
throughoutthe year.
Nothing is known about pollination in Cous-
sapoa. Neither the structureof the inflorescences
nor the features of the stamens suggest the oc-
currenceof wind pollination, however.
In most cases dispersal is probably endozo-
monkeys(Dr. M. van Roosmalen, pers.comm.).
Exozoochorous dispersal may occur when the
exocarpis more or less mucilaginous(andsticky),
as in C. asperifolia.
Anatomy
By K. BONSEN
SecondaryXylem.-General Properties.Col-
or: light yellowish brown.Grain:straight,some-
times slightly interlocked. Texture: medium.
Odor and taste:not characteristic.Specificgrav-
ity: 0.50-0.75.
Structure.Based on eight species, nine speci-
mens, the growthrings are faint or absent. Ves-
sels diffuse,roundto oval, solitary(25-83%),and
in radial multiples and irregularclusters of 2-8
(-21), (0-1)1-6(-11) per sq mm, diameter(200-)
220-300(-340) gm. Vessel memberlength(400)
475-600(-725) Mm.Perforationssimple. Inter-
vascularpits alternate,round or polygonal, 10-
15 Lm.Thin-walledtyloses are usually present.
Fibersnon-septate,diameter 18-25,m, walls 2-
3.5 tm, L/W ratio 2-5, gelatinousfibersusually
present.Pits simple, mainly on the radialwalls.

a ad tfg
1 2 34 5 6
FIG. 4. Schematicdrawingsof fruits in Coussapoa:1-4, C. asperifoliatype, 1, 2, fruit in perianth,3, fruit
in opened perianth,4, endocarpbody; 5, 6, C. villosatype, fruit in perianth.a, embryo;b, endosperm;c, testa;
d, endocarp;e, mucilaginousmesocarp;f, exocarp;g, perianth.
Anatomy
, ,. : ?
. . . .

Ii
;~ ~
~~~~:':-
~ ~~.: II~C
...? :
%,-:

? ~ ~' ~ i 2.''? ..','

'" ? "'."' ..

:i... t
' . ...-I. : .,
:. '
%;.: ,, '
I,~~~~~.
F? '
I - i. "' ~,c .?.h

? ~
t*l..:.
1;.::.:.;. ... ~ ~,,~x
.....~ .
"~~~-i
FiG. 5. Woo ofCo p~ltf'l9
, ,cos ci~ ~ lsci~(.22

Length(875-)1100-1800(-2175) .m, F/V ratio Leaf Anatomy. -Structure. Based on 11

2.5-3.7. species, 18 specimens.
Rays heterogeneous,uniseriate(21-35%) and In surface view. Indumentum of thin, inter-
multiseriate(3-)4-7(-9) per mm, sheath cells woven, unicellulararachnoidhairs,abaxialpres-
presentor absent. Uniseriate rays mainly com- ent or absent; unicellular needle-shaped, often
posed of square to upright cells, ray height wavy hairs (mostly on abaxial surface,rarelyon
(200-)300-500(-980) gtm. Multiseriate rays adaxial surface);adaxial, glandular hairs with
composedof uprightand procumbentcells, ver- multicellular,globularheads on 3-5-celled, uni-
tically compounded 1-10%, (450-)700-1100 seriate stalks, mostly in groups of 2-4, present
(-1600) ,m high, 3-6 cells in width, uniseriate or absent;abaxial,uniseriate,6-10-celled,curved,
parts(0-)1-2(-8) cells. glandularhairswith globularto elongatedheads,
Parenchymaparatracheal,banded, irregular, mostly present;and conical papillae sometimes
wavy, (0-)1-2(-3) per mm, (3-)5-9(-15) cells in present.Epidermalcells polygonal;adaxial cells
width. Strands5-8(-14) cells, length (530-)600- overlying large crystalliferousmesophyll cells
710(-870) Mm,containingsome to many rhom- forming a rosette. Stomata confined to abaxial
bic crystals.Radial latex tubules have been ob- surface,anomocytic,averagelengthof guardcell
served in Coussapoalatifolia (see Fig. 5). pairs 15-20 gm, averagewidth 12-18 um. Hy-
The wood of Coussapoaresemblesthat of My- dathodes formed by 10-15 water pores each,
rianthusand some species of Pourouma. The presentor absenton adaxialsurface.Minorveins
presenceof confluent-bandedparenchymain the usually very prominentin abaxial epidermis.
wood of Coussapoadistinguishesthis genusfrom In transverse section. Lamina bifacial. Epi-
the wood of Cecropiaand Musanga (Bonsen & dermal cells small, especially abaxiallybetween
ter Welle, 1983). the veins. Adaxialepidermalcellssometimeswith
10 Flora Neotropica

silicified outer walls. Stomata sometimes raised
above level of unspecializedcells. Adaxial hy-
podermis of two or three layers of parenchyma
cells, includingmucilage cells except for C. vil-
losa. Mesophyll consisting of one layer of pali-
sade cells, sometimes subdivided loose spongy
tissue, with or without an intermediate layer.
Veins with sclerenchymatousvertical bundle
sheathextensions (touchingadaxialhypodermis
and abaxial epidermis). Midrib with a flat or
raised adaxial surfaceand a prominentlyraised
abaxial surface;peripheralgroundtissue paren-
chymatousto collenchymatous,interspersedwith
mucilage cells; vascular system composed of a
closed or variously interruptedcylinder, partly
or whollysurroundedby sclerenchymafibers,and
enclosing one or two rows of bundles which are
situatedin the same directionas the most abaxial
bundleof the cylinder.Vascularsystemof petiole
similar.Crystalspresentas drusesthroughoutthe
mesophyll, in the petiole and midrib; rhombic
crystalssometimes present. Distinguishmentof
the inner structureof the leaves of Coussapoa
from those of the other Neotropicalgenera, Ce-
cropia and Pourouma,can be accomplishedon
the basis of the petiole vascular system type
(Bonsen& ter Welle, 1983). Some differencesin
the inner structureof some species within the
genus Coussapoa are mentioned by Renner
(1907).
Distribution and Ecology
The genus,beingdistinctlyassociatedwith the
rain forest habitat, is widespread in tropical
America, but is absent in the West Indies; see
Figures6-9.
Most species are components of lowland rain
forests.A few species,e.g., Coussapoavillosaand
C. asperifolia,extend into submontaneor mon-
tane forest. Some species are associated with
higheraltitudes(from ca. 800 m) in the Guiana
(=Guayana) Highland region, e.g., C. argentea
and C. viridifolia,while othersarefoundin higher
altitudes (from ca. 900 m or from ca. 1400 m)
in the Andean region, e.g., C. cinnamomifolia
and C. duquei;C. crassivenosais found in both
regions.Some speciesalso occurin drierhabitats,
such as savanna (C. viridifolia)or semi-decidu-
ous forest (C. purpusii). Most lowland species
appearto occurboth in uplandand riverinefor-
est. A few species, like C. nitida,appearto show
a preferencefor riverside(varzea)forest. C. tri-
nervia, the only truly terrestrialspecies of the
genus, is confined to riverbanks, apparently
(only?)of black-waterrivers.
Only two species, C. villosaand C. asperifolia,
have an extendeddistribution.Most of the other
species are confinedto, or at least distinctly as-
sociated with, one of the phytogeographicre-
gions, such as eastern Brazil, Amazon Basin,
eastern Guiana (=Guayana Lowland) region,
western Guiana (=Guayana Highland) region,
Andean region, Pacificcoastal region, and Cen-
tralAmerica.Many speciesare only knownfrom
a very small area.
Severalspecieswith medium-sizedareasshow
a discontinuousdistribution.It is, however, not
certainwhetherthese discontinuitiesare real or
due to insufficientexploration.The Amazonian
C. trinerviahas a disjunct distribution, which
appearsto be relatedto its ecology (see above).
C. crassivenosais known from four remote re-
gions. C. latifolia and C. asperifoliassp. asperi-
folia, common taxa of the easternGuianaregion
to easternParf, show similargapsin distribution
and are found segregatedin more central parts
of the Amazon Basin as well. If these gaps are
real,the same factorsmay have causedthe west-
ern Parf gap in the distributionof C. nitida, as
well as the disjunct distribution of the species
pairs C. arachnoidea-C. orthoneuraand C. le-

FIG. 6. Distributionof Coussapoaangustifolia,C. arachnoidea,C. argentea,C. asperifoliassp. asperifolia,

C. asperifoliassp. magnifolia, C. asperifoliassp. rhamnoides,C. batavorum,C. brevipes,C. chocoensis, C.
cinnamonea,C. cinnamomifolia,C. contorta,C. crassivenosa,and C. cupularis.
FIG. 7. Distributionof Coussapoacurranii,C. duquei,C. echinata,C.ferruginea,C.floccosa, C. glaberrima,
C. herthae, C. latifolia, C. leprieurii,C. macerrima,C. microcarpa,C. microcephala,C. napoensis, and C.
parvifolia.
FIG. 8. Distributionof Coussapoalongepedunculata,C. manuensis,C. nitida, C. nymphaeifolia,C. oligo-
cephala, C. orthoneura,C. ovalifolia, C. pachyphylla,C. parviceps,C. purpusii,C. scabra, and C. sprucei.?
unnamedcollection aff. C. ovalifolia.
Distribution and Ecology 11

0 C.angustifolia Aublet

OC.arachnoidea Akkermans 6 Berg

r
^^-^^'.L_ n //i C ^ 4-Lc<'' ^/ ^ A C.argentea Akksrmans & Berg

~
!
- -' --/\i . _ C.asper.folia Tr.cul
\ \.f C( J / 0 a. ssp. asperifolla
A b. spp. magnifolla (Trdcul) Akkermans 6 Berg

- ' i \ Sc. spp. rhamnoides (Standley) Akkermans & Berg

.
, ,X ? _ _ j**C.cinnamoa ifolla MiIdbraed
|__ /-J?Z- _,'nX Sjo . C.contorta Cuatrecasas
...C
A C.crassivenosa Mlldbraed
C r vlp P C.cupularis Akkermsns & Berg
S ?

0 C.brevipes Pittlere a
A C.chocoensls Cuetrecasas
12 Flora Neotropica

v* r

r'^S'9n 7 J (^^'^^J^'^ - ^^Q^0 C.duquei Standley

A C.echinata Akkermans Berg
* C.curranli Blake i .e -
T C.ferruginea Trecul

| ? (% 'J/' / { , t
__l|: C.p_rvi fo ^g d C-latifolia Aublet

A C.glaberrima
Burger
* C.herthee

C.floccosa y
i
Akkermans Berg . B

S
Mildbred

C.cerricroc Akkerpa Berg

(SchQtt) izzini ~ -

0 C'microcephalaTrdcul '
V' C.parvifolia Standley
A C.napoensis Akkermmns & Berg

,.
Distribution and Ecology 13

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

.AC.oligo ala Donnael Smith

C.orthoneura Stand ey

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

HC.pcyhl tcuatl
Tra
Aknn ~t C.oval 1 i fol ia %

C.parviceps Standey\

C.scabra Akkermans 6 Berg

*C.sprucel Mildbraed
\
 I C. tessmannll MIldbrwd

A C. fulvescens C.C. Berg

VC. tolimensis C.C. Berg

Ce. voiaria C.C. Berg

C Mil dbraed
1.trinervia
A C.vannifolia Cuatrecasas

*C.villosa Poeppig 6 Endlicher

AC.viridifolia Cuatrecasas

FIG. 9. Distribution of Coussapoafulvescens, C. tessmannii, C. tolimensis, C. trinervia,C. vannifolia, C.

villosa,C. viridifolia,and C. valaria (name is misspelledin legend).
Systematic Position
prieurii-C. sprucei. This type of distribution
may be connectedwith the distributionof Pleis-
tocene rain forest refugia (cf. Prance, 1977).
Another remarkablediscontinuity in distribu-
tion occurswith C. cinnamomifolia,a species of
medium altitudes in the Andean region. This
discontinuityshows resemblancesto that of two
partial areas of C. crassivenosa.The Ecuador-
Bolivia disjunctionis probablynot genuine and
more likely to be due to inadequateexploration
in Peru.
The discontinuities in the distribution of C.
microcarpa,especially the isolated localities in
Perambuco and Paraiba,could be relatedto the
more or less isolated occurrenceof forest with
rain forest elements on low mountain tops.
In the presenttreatment,severalpairsof closely
related species are recognized,e.g., C. crassive-
nosa-C. ferruginea, C. latifolia-C. viridifolia.
These pairs of species have allopatricdistribu-
tions.
Systematic Position of the Genus
Coussapoa,togetherwith the Neotropicalgen-
era Cecropiaand Pourouma,the Africangenera
Musangaand Myrianthus,and the Asiatic genus
Poikilospermum,constitute the family Cecropi-
aceae (Berg, 1978).
Poikilospermum,geographicallyisolated from
the othergenera,has more characters,especially
anatomicalones, in common with the Urticaceae
than the other genera (cf. Bonsen & ter Welle,
1983). The latter authors,indeed, suggesttrans-
ferringthe genus back to the Urticaceae.Never-
theless,consideringthe distributionof characters
throughoutthe (complex of) families Cecropi-
aceae, Urticaceae, and Moraceae, placing Poi-
kilospermumin the Cecropiaceaeappearsto be
justified, althoughnot beyond all doubt.
Both Coussapoaand Poikilospermumare pre-
dominantlyhemi-epiphytic(cf. Chew Wee-Lek,
1963). However, the two generadiffer in habit!
Coussapoaspeciesareshrubsor treeswith straight
and stiffbranches,while Poikilospermumspecies
are more or less liana-like with long and weak
(scrambling)branches(cf. ChewWee-Lek,1963).
The two genera resemble each other in the
branched inflorescenceswith terminal globose
heads, but the basic ramificationof the inflores-
cence appearsto be different.Dichotomousram-
ificationappearsto be basic in Poikilospermum,
but is derived in Coussapoa(see above). Expul-
15
sion of the diasporesis anotherfeaturewhich the
two generahave in common; in Coussapoait is
(probably)confinedto C. asperifoliaand in Poi-
kilospermumit is only found in subg. Liguli-
stigma(cf. ChewWee-Lek,1963). This phenom-
enon, in combination with microspermy, is
probably associated with the hemi-epiphytic
habit. In Coussapoathe mucilaginouslayer that
swells and pushes the endocarp out of the
perianth is formed by the middle layer of the
pericarp.In Poikilospermum,however, the ex-
pulsion of the fruits is achieved by the swelling
of the inner surfaceof the perianth, which be-
comes mucilaginousduring the ripening of the
fruit (cf. Chew Wee-Lek, 1963).
Judgingfrom (the nature of) the similarities
and differences,the two genera are ecological
equivalentsratherthan systematicallyclosely re-
lated. Morphologicalas well as anatomicalchar-
acters (cf. Bonsen & ter Welle, 1983) suggesta
closerrelationshipof Coussapoato the otherfive
generaof the family, especiallyto the Neotrop-
ical ones. Coussapoa matches Cecropiain mi-
crospermy.The two generashow similaritiesin
the flowers, but otherwise are quite different.
More similaritiesare found between Coussapoa
and Pourouma,althoughthe lattergenus is mac-
rospermous.The inflorescencesof Coussapoare-
semble the staminate ones of some Pourouma
species, e.g., P. tomentosa and P. mollis; some
Pouroumaspecies have leaves stronglyreminis-
cent of those common in Coussapoa(see above).
Systematic Arrangement
of the Species
In the presentrevision the species are treated
in alphabeticalorder,as it is not possible to pro-
pose a satisfactorysystematicarrangement.
For most charactersclear discontinuities in
theirvariationarelacking.Primitiveand derived
state (as the extremesof a morphologicalseries)
can be inferred,like lamina broadlyovate-nar-
rowly obovate, basal lateralveins branched-un-
branched, lateral veins numerous-few, petiole
long-short,stipuleslong-short,indumentumwith
all types of hairs and ? dense-less diverse and
? sparse, inflorescencesrepeatedly branched-
unbranched,interfloralbractspresent-absent,and
stamens three-two. See Table I.
In some species (e.g., C. nymphaeifolia,C. vil-
losa, and C. tessmannii) a predominance of
"primitive" characterscan be found or a pre-
16 Flora Neotropica

Table I numberof stamensin arrangingthe species,since

Extremesin a morphologicalcharacterseries in for aboutone-thirdof the speciesrecognizedma-
Coussapoa,inferredto be primitive and derived terial belongingto only one sex is known.
respectively. Acceptanceof the number of stamens as the
basis for subdividingthe genus would imply the
Primitivestates Derivedstates occurrenceof parallel (or homologous) differ-
Laminabroadlyovate Laminanarrowly ob- entiation of many characters,e.g., in the differ-
ovate entiationof the leaf shape,dimensions,venation,
Basallateralveins Basallateralveinsun- indumentum,and length of the petiole, in three
branched branched
Lateralveinsnumerous Lateralveinsfew groupsof species (judgingfrom the comparison
Petiolelong Petioleshort of charactersin the taxa of which the numberof
Stipuleslong Stipulesshort stamens is known).
Indumentwithall types Indumentlessdiverse
of hairsand? dense and+ sparse
Inflorescencesrepeatedly Inflorescences un- Taxonomy
branched branched
Interfloralbractspresent Interfloral bractsabsent 1. CoussapoaAublet, Hist. pl. Guiane 2: 955.
Stamens3 Stamen1 1775;Trecul,Ann. Sci. Nat. Bot. Ser. 3, 8: 92.
1847; Miquel, Fl. bras. 4(1): 131. 1853; Mac-
bride,Publ.FieldMus. Bot. 13(2.2):295. 1937;
Woodson& Schery,Ann. MissouriBot. Gard.
47: 168. 1960; Berg, Fl. Suriname5(1): 279.
dominanceof"derived"charactersin others(e.g.,
C. cinnamomifolia, C. trinervia,and C. viridi- 1975; Burger,Fieldiana Bot. 40: 131. 1977;
Akkermans& Berg, Proc. Kon. Ned. Akad.
folia), with the majority of the species some- Wetensch.Ser. C, 85(4): 441. 1982. Lectotype
wherein between.A few specieshave exceptional
characters:C. trinerviahas a trulyterrestrialhab- species (Berg, Fl. Suriname5(1): 279. 1975):
it, C. parvicepshas fused pistillate flowers; C. Coussapoalatifolia Aublet.
contortahas domatium-likecavities in the axils Hemi-epiphytic (or epiphytic) or terrestrial
of the lateral veins; C. asperifoliahas abruptly trees or shrubs with aerial roots or stilt-roots.
thickened twigs with wide cavities and, more- Leavesin spirals,entire,venation pinnateto tri-
over, a thickmucilaginousmesocarp,and C.pur- nervate, margin entire to subcrenate, stipules
pusii is (often?)deciduous. fused, fully amplexicaul,usuallyleaving oblique
The differencein the number of stamens ap- scars. Inflorescencesmostly in pairs in the leaf
pearsto be a promisingbasis for the arrangement axils, branched or in pistillate ones often un-
of the species or the subdivision of the genus. branched,bracteate(mostly only with interfloral
The significanceof these differenceslies not only bracts)or ebracteate.Flowersin globose (to el-
in the absence of transitionsbut also in the dis- lipsoid or clavate)terminalheads, free or pistil-
tributionranges:the predominanceof specieswith late ones sometimes connate, perianthtubular,
3-staminateflowersin CentralAmerica and the in staminateflowers(2-)3(-4)-lobed, in pistillate
Pacificcoastal regionof Colombiaand Ecuador; ones entire; stamens two or three and connate,
the predominanceof species with 2-staminate or one; ovary free, stigma penicillateto subpel-
flowersin easternSouthAmerica,especiallyeast- tate. Fruit a drupelet or almost an achene, en-
ern Brazil;and the concentrationof species with closed in the enlarged + fleshy perianth;seed
uni-staminateflowersin the Upper Amazon Ba- with endosperm,embryo straightwith flat and
sin. It is not yet possible, however, to use the equal cotyledons and relatively short radicle.
Coussapoa 17

Keys to the Species of Coussapoa

Generalkey
Keys to the species in:
1. CentralAmerica and Mexico
2. Colombia
3. Venezuela
4. the Guianas2 2
5. Ecuador
6. Peru and Bolivia " \ .7 /
7. Amazonian Brazil
8. EasternBrazil

General Key to the Species of Coussapoa

1. Interfloralbractsabsent.
2. Basal lateralveins branched.
3. Lamina beneath with dense indumentum of minute hairs, at least in the areoles and on the
reticulum.
4. Indumentum hairsonly!)predominantly
(unicellular white;pistillateinflorescences
unicapi-
tate; stamens 3.
5. Intercostalvenation prominentbeneath;Costa Rica. .............. 29. C. nymphaeifolia.
5. Intercostalvenation plane beneath;Panama. ............................. 6. C. brevipes.
4. Indumentum(unicellularhairsonly!)at leastpartlybrownishor yellow;pistillateinflorescences
usuallypluricapitate;stamen 1 or unknown;Amazon Basin or Colombiaat ca. 1900 m.
6. Laminausuallyelliptic; lateralveins 5-9 pairs. ..................... See 12. C. cupularis.
6. Laminausuallyovate; lateralveins 8-15 or 14-22 pairs.
7. Lateralveins 8-15 pairs;peduncleof the pistillateinflorescence1-2.5 mm thick.
8. Intercostalvenation (almost) plane; indumentum of petiole consisting of dense
minute hairs and distinctlylonger,yellowish hairs;Amazon Basin ..... 28. C. nitida.
8. Intercostalvenation prominent;indumentumof the petiole consistingof hairs not
distinctlydifferentin length;Colombia (ca. 1900 m). ............. 45. C. tolimensis.
7. Lateralveins 14-22 pairs;peduncleof the pistillateinflorescence6-10(-15) mm thick;
Colombia (Amazonas). ......................................... 8. C. cinnamomea.
3. Laminabeneathwith sparseindumentumor glabrous,at leastin the areolesand on the reticulum.
9. Stipules0.5-1 cm long; Venezuela(Bolivarand Amazonas),Guyana,and Brazil(Amazonas,
Roraima). ........................................ .................. 43. C. viridifolia.
9. Stipulesat least 1 cm long.
10. Lateralveins 3-6 pairs, the basal pair reachingthe marginat or above the middle of the
lamina;pistillate flowerscohere;stamens 3; CentralAmericaand Pacificcoastal region
of Colombia and Ecuador. ...........................................34. C. parviceps.
10. Lateralveins 7 or more pairs,the basal pair reachingthe marginbelow the middle of the
lamina;if less than 7 pairs of lateralveins then the pistillateflowersfree and stamen 1.
11. Lamina and leafy twigs glabrous;base of the lamina cordate;Colombia (Choco).
.............................................................. 5. C. batavorum .
18 Flora Neotropica

11. Lamina and leafy twigs at least sparselypuberulous;base of the lamina acute to
truncate.
12. Stipules(normally)1-4 cm long.
13. Marginof the laminarevoluteat the base;Pacificcoastalregion(Colombia).
......................................................... 46. C. valaria.
13. Marginof the lamina not revoluteat the base; Amazon Basin and Central
America.
14. Laminacoriaceous,mostly ovate; stamen 1; Amazon Basin (and Pan-
ama?). ............................................. 32. C. ovalifolia.
14. Laminasubcoriaceous,elliptic to subobovate;stamens 2; Costa Rica.
........................................ 23. C. macerrima.
12. Stipules4-13 cm long; AmazonianEcuadorand Colombia. .... 27. C. napoensis.
2. Basal lateralveins unbranched(or occasionallywith a single branch).
15. Stipules0.5-1 cm long;stamens2; common peduncleor branchesof the pistillateinflorescence
not distinctly broadenedtowards the flower heads; eastern Venezuela, Guyana, and Brazil
(Amazonas, Roraima). ................................................... 43. C. viridifolia.
15. Stipules(normally)at least 1 cm long, if shorter,then stamen 1 and branchesof the pistillate
inflorescencemore or less distinctlybroadenedtowardsthe flowerheads.
16. Marginof the lamina distinctlyrevolute towardsthe base; pistillateinflorescencesmostly
unicapitate;stamens 2; easternBrazil. ................................ 25. C. microcarpa.
16. Marginof the lamina plane, or if more of less revolute, then the pistillate inflorescences
pluricapitateand stamen 1; Amazon Basin or CentralBrazil.
17. Lateralveins 2-8; basal pair of lateralveins reachingthe marginabove to just below
the middle of the lamina and/or (often) the lamina beneathwith dense, minute hairs
in the areoles.
18. Intercostalvenation prominentbeneath;lamina beneathand inflorescenceswith
dense, white, arachnoidindumentum;Brazil(Amapa) .......... 2. C. arachnoidea.
18. Intercostalvenation plane beneath;lamina and inflorescenceswithout arachnoid
indumentum, or if present, then mostly sparse and soon disappearing;Upper
Amazon Basin.
19. Basallateralveins reachingthe marginabove to just below the middle of the
lamina;stipules mostly without or with sparse,long, yellow hairs;branches
of the pistillate inflorescenceslightly broadenedtowards the flower heads;
Upper Amazon Basin and Colombia (Santander). .......... 31. C. orthoneura.
19. Basallateralveins reachingthe marginbelow to just above the middle of the
lamina;stipules with dense, long, yellow hairs;branchesof the pistillatein-
florescencestronglybroadenedtowardsthe flowerheads;Brazil(R6ndonia).
......................................................... 12. C. cupularis.
17. Lateralveins (normally)7-13 or 7-21 pairs, basal pair reachingthe marginfar below
the middle of the lamina;lamina beneathglabrousor with sparsehairs in the areoles.
20. Stipules(normally)1-4 cm long.
21. Laminacoriaceous,mostly ovate; stamen 1; Amazon Basin (and Panama?).
......................................................... 32. C. ovalifolia.
21. Laminasubcoriaceous,elliptic to subobovate;stamens 2; Costa Rica. .....
........................................ 23. C. macerrima.
20. Stipules4-13 cm long; AmazonianEcuadorand Colombia. ...... 27. C. napoensis.
1. Interfloralbractspresent(sometimes small and/or few!).
22. Laminatrinervateor with only the basal pairof lateralveins prominentand the otherlateralveins
weak and inconspicuous.
23. Laminatrinervate;intercostalvenation prominentbeneath;Amazon Basin .... 40. C. trinervia.
23. Laminawith only the basal pair of lateralveins prominentand the other lateralveins weak
and inconspicuous;intercostalvenation beneath; Bolivia (La Paz), Ecuador(Pastaza),and
Colombia (Antioquia). ..............................................9. C. cinnamomifolia.
22. Laminapinnatelyveined.
24. Laminaabove with white arachnoidhairs,at least (sub)persistenton the midrib;easternBrazil.
25. Stipulesca. 0.5 cm long ............................................... 13. C. curranii.
25. Stipules 1-2.5 cm long ................................................ 17. C. floccosa.
24. Laminaabove glabrous(sometimesexcept for the base of the midrib)or scabrousto scabrid-
ulous with short, rigidhairs;not in easternBrazil.
26. Laminascabrousto scabridulousabove.
27. Stipulesca. 6 cm long;laminabeneathwitharachnoidhairs;basallateralveinsbranched;
Peru (Loreto)and Ecuador(Napo). ........................ 22. C. longepedunculata.
27. Stipules up to 3.5 cm long, mostly less than 1 cm long; lamina beneath lacking
arachnoidhairs, or if presentthen the basal lateralveins unbranched.
Coussapoa 19

28. Leafytwigs (rather)abruptlythickenedand distinctlyhollow; widespread ....

............................................................ 4. C asperifolia.
28. Leafy twigs graduallythickenedand solid.
29. Lateralveins 6-13 pairs;laminasubcoriaceous;southernAmazon Basin. ...
............................... ................ ........... 37. C. scabra.
29. Lateralveins 6-7 pairs;laminacoriaceous;FrenchGuianaand Brazil(Para).
........................................................ 21. C. leprieurii.
26. Laminasmooth and glabrousabove.
30. Basal lateralveins unbranched(or sometimes with 1 or 2 branches).
31. Lamina beneath with domatium-likecavities in the axils of the lateral veins;
interfloralbractslargeand peltate;westernEcuadorand Colombia. ..........
....................................... ...................... 10. C. contorta.
31. Laminabeneathwithoutdomatium-likecavities;interfloralbractsrelativelysmall
and spathulateto subpeltate.
32. Stipules0.5-1 cm long.
33. Lateralveins 2-3(-4) pairs.
34. Apex of the lamina acute;CentralAmerica. ...... 18. C. glaberrima.
34. Apex of the lamina roundedto obtuse; Surinam,French Guiana,
and Brazil(Amapaand Para). .................... 1. C. angustifolia.
33. Lateral veins at least 4 pairs.
35. Youngleaves with arachnoidindumentum;laminaobovate to sub-
obovate; lateral veins 7-10 pairs; terminal buds slender;eastern
Brazil. .................... ..................... 13. C. curranii.
35. Young leaves withoutarachnoidindumentum;laminaelliptic, or if
(sub)obovate,then lateral veins 4-7 pairs and/or terminal buds
swollen;not in easternBrazil.
36. Terminalbuds swollen;Upper Amazon Basin and Colombia.
........................................ 35. C. parvifolia.
36. Terminalbuds slender.
37. Lateralveins 4-7 pairs.
38. Hairson the main veins beneathof equallength;Low-
er Amazon Basin, Surinam,and FrenchGuiana. ...
.............. ...................... 20. C. latifolia.
38. Hairs on the main veins beneathof differentlengths;
eastern Venezuela, Guyana, and Brazil (Amazonas,
Roraima). ......................... 43. C. viridifolia.
37. Lateralveins 7-11 pairs; Upper Amazon Basin and Co-
lombia. ................................ 35. C. parvifolia.
32. Stipulesat least 1 cm long.
39. Lateralveins 2-3(-4) pairs.
40. Laminaglabrous;CentralAmerica. .............. 18. C. glaberrima.
40. Laminabeneath with (partlyarachnoid)indumentum;Venezuela.
........................................ 3. C. argentea.
39. Lateralveins at least 4 pairs.
41. Laminabeneathwith dense indumentum.
42. Arachnoidhairspresenton the laminabeneath,at leastin young
leaves.
43. Arachnoidhairs brownish.
44. Lateralveins 10-14 pairs;pistillateinflorescencesuni-
capitate, head ellipsoid to clavate; staminate flower
heads 4-8 mm in diameter;Andean Colombia and
Ecuador(1400-2000 m). ............... 14. C. duquei.
44. Lateralveins at most 10(-11);pistillateinflorescences
pluricapitate,or if unicapitatethen the headsglobose;
staminateflowerheads 1-3 mm in diameter.
45. Lateralveins 4-5 pairs; lamina oblong; French
Guiana and Brazil(Amapa). ..... 16. C.ferruginea.
45. Lateralveins (4-)5-8(-11) pairs; lamina mostly
elliptic to ovate.
46. Hairson the smallerveins beneathappressed;
Guianas ................ 26. C. microcephala.
46. Hairs on the smallerveins beneath (at least
partly)patent.
47. Stamens slightly longer than the peri-
20 Flora Neotropica

anth;pistillateflowerheads 1-3; eastern

Brazil. ............... 33. C. pachyphylla.
47. Stamensdistinctlylongerthan the peri-
anth; pistillate flowerheads 3-7; Vene-
zuela (Bolivar),Brazil (Roraima),Am-
azonian Bolivia and Ecuador,and
Panama. ............ 11. C. crassivenosa.
43. Arachnoidhairs whitish.
48. Laminamostly ovate;lateralveins 9-16 pairs;stamen
1;peduncleof the pistillateinflorescence2-4 mm thick;
Amazon Basin. .................... 39. C. tessmannii.
48. Lamina mostly elliptic to oblong;lateralveins 5-12
pairs; stamens 3; peduncle of the pistillate inflores-
cence ca. 1 mm thick.
49. Stipules0.5-1.5(-2) cm long; heads of staminate
inflorescencemany, 3-4 mm in diameter;heads
of pistillateinflorescence2-5; perianthglabrous;
Amazon Basin. .................... 38. C. sprucei.
49. Stipules 1-5 cm long; heads of staminateinflo-
rescence3-10, 4-6 mm in diameter;headsof pis-
tillate inflorescence1(-3); perianthminutely pu-
berulous;CentralAmericato Mexico..........
............................. 30. C. oligocephala.
42. Arachnoidhairs lackingon the lamina beneath, or if present
then sparseand (soon) disappearing.
50. Lateralveins 5-8 pairs;easternBrazil. .. 33. C. pachyphylla.
50. Lateralveins 8-12 pairs;not in easternBrazil.
51. Marginof the laminarevoluteat the base;UpperAm-
azon Basin and Colombia. ........... 35. C. parvifolia.
51. Marginof the lamina not revoluteat the base.
52. Petiole 1-3 cm long; stipules 1.5-2.5 cm long;
pistillateinflorescencesunicapitate,the peduncle
0.5-1 cm long; Colombia(Choc6). 7. C. chocoensis.
52. Petiole 3-9 cm long;stipules2-6 or 3-20 cm long;
pistillate inflorescencespluricapitate,or if uni-
capitatethen the peduncleat least 1 cm long.
53. Heads of pistillate inflorescence6-10; sta-
mens 3; Pacificcoastalregionof Ecuadorand
Colombia. ................... 19. C. herthae.
53. Heads of pistillate inflorescence1(-2); sta-
mens 2 or unknown.
54. Laminabroadlyellipticto obovate,apex
shortlyacuminate;pistillateflowerheads
ca. 4 mm, in fruit up to 12 mm in di-
ameter;AmazonianPeru ...........
...................... 24. C. manuensis.
54. Laminamostly ovate to subovate,apex
acute to obtuse; pistillate flower heads
ca. 5-10 mm, in fruit up to 40 mm in
diameter;widespread....... 42. C. villosa.
41. Laminabeneathwith sparseindumentum.
53. Marginof the lamina distinctlyrevolutetowardsthe base.
56. Stipules 1-7 cm long;terminalbuds slender;easternBrazil
...................................... 25. C. microcarpa.
56. Stipulesup to 1.3 cm long;terminalbuds usuallyswollen;
Upper Amazon Basin and Colombia. ..... 35. C. parvifolia.
55. Marginof the lamina plane or entirelyrevolute.
57. Leafytwigs(rather)abruptlythickenedand with wide cav-
ities; stamen 1; widespread ............... 4. C. asperifolia.
57. Leafy twigs graduallythickenedand solid or with narrow
cavities.
58. Stipulesglabrousor puberulous.
59. Basallateralveins reachingthe marginabove the
Coussapoa 21

middle of the lamina; base of the lamina acute;

pistillateinflorescencespluricapitate; Panama. ..
............................... 15. C. echinata.
59. Basal lateral veins mostly reachingthe margin
belowthe middleof the lamina;baseof the lamina
mostly obtuse to (sub)cordate.
60. Stamens 3; pistillate inflorescencesunicapi-
tate; northernCentralAmerica to Mexico.
............................. 36. C. purpusii.
60. Stamens2; pistillateinflorescencespluricap-
itate; South America.
61. Hairs on the main veins beneath equal
in length and arachnoidhairs lacking;
Surinam, French Guiana, and Brazil
(Amapa,Para, and Amazonas). ......
......................... 20. C. latifolia.
61. Hairs on the main veins beneathof dif-
ferentlengthsand/or on the main veins
and margin subpersistent arachnoid
hairs;Guianas....... 26. C. microcephala.
58. Stipules(sub)sericeous,(sub)hirsute,(sub)villous,etc.
62. Stamens3; pistillateflowerheads 6-10, 5-9 mm
in diameter;Pacificcoastalregionof Ecuadorand
Colombia.
63. Petiole 5-10 cm long, or if shorter,then sta-
mens 3, pistillateflowerheads6-10, 5-9 mm
in diameter;Pacificcoastalregionof Colom-
bia and Ecuador.
64. Stipules2-6 cm long. ..... 19. C. herthae.
64. Stipules 1.5-2 cm long. .. 44. C.fulvescens.
63. Petiole up to 5 cm long; stamens 2; flower
heads 1-5; not in Pacificcoastalregion.
62. Stamens2; pistillateflowerheads 1-3, or if more
than 3 then 2-5 mm in diameter.
64. Stamensslightlylongerthanthe perianth;pis-
tillateflowerheads 1-3; easternBrazil......
........................ 33. C. pachyphylla.
64. Stamensdistinctlylongerthan the perianth;
pistillateflowerheads (1-)3-15; Guianas. ..
........................ 26. C. microcephala.
30. Basal lateralveins branched.
65. Lamina beneath with (rather)dense, (rather)conspicuous and (sub)persistent
arachnoidindumentum.
66. Arachnoidhairs on the lamina brownish.
67. Lateralveins 4-8(-11) pairs;pistillate inflorescencesmostly pluricapi-
tate; staminateflowerheads ca. 2-3 mm in diameter.
68. Hairs on the smallerveins beneath(at least partly)patent.
69. Stamensslightlylongerthanthe perianth;pistillateflowerheads
1-3; easternBrazil. ....................... 33. C. pachyphylla.
69. Stamens distinctly longer than the perianth;pistillate flower
heads 3-7; Venezuela(Bolivar),Brazil(Roraima),Amazonian
Bolivia and Ecuador,and Panamf. ......... 11. C. crassivenosa.
68. Hairs on the smallerveins beneathappressed;Guianas..........
............................................ 26. C. microcephala.
67. Lateralveins (7-)10-24 pairs;pistillateinflorescencesusually unicapi-
tate; staminateflowerheads 4-10 mm in diameter.
70. Commonpeduncleof the staminateinflorescence1-2 cm long, that
of the pistillateinflorescence0.5-0.8 cm long;pistillateheadsellip-
soid to clavate;AndeanEcuadorand Colombia(1400-2000 m). ..
.................................................. 14. C. duquei.
70. Commonpeduncleof the staminateinflorescence2-6 cm long, that
of the pistillate inflorescence(1-)2-13 cm long; pistillate flower
heads (sub)globose;widespread. ..................... 42. C. villosa.
22 FloraNeotropica
66. Arachnoidhairs on the lamina whitish.
71. Lateralveins ca. 10 or 9-24 pairs;lamina mostly longer than 15 cm;
stipules mostly longerthan 2 cm.
72. Laminabeneathwith densebrownhairson the mainveins;common
peduncleof the staminateinflorescenceca. 9 cm long; Amazonian
Peru and Ecuador. ....................... 22. C. longepedunculata.
72. Laminabeneathwith whitish hairs on the main veins; peduncleof
the staminateinflorescenceup to 6 cm long.
73. Stamen 1; pistillate inflorescencespluricapitateand common
peduncle2-3 cm long; Amazon Basin. ....... 39. C. tessmannii.
73. Stamens2 or 3; pistillateinflorescencesunicapitate,or if 2-3-
capitatethen the common pedunclelongerthan 3 cm.
74. Stamens3; pistillateflowerheads ellipsoid to clavate;pe-
duncle of the pistillateinflorescence1-2 cm long; Central
America. .......................... 29. C. nymphaeifolia.
74. Stamens 2; peduncleof the pistillateinflorescence(1-)2-
13 cm long; pistillate flower heads (sub)globose;wide-
spread. ................................... 42. C. villosa.
71. Lateralveins 4-10(-11) pairs;lamina mostly shorterthan 15 cm.
75. Hairs on the smallerveins beneathappressed;Guianas..........
............................................ 26. C. microcephala.
75. Hairs on the smallerveins beneath(at least partly)patent.
76. Stipulesup to 2 cm long.
77. Hairs on the stipules predominantlyweak and crinkled;
stamens 2; pistillateflowerheads 1-3; easternBrazil. ...
..................................... 33. C. pachyphylla.
77. Hairson the stipulespredominantlystiffand straight;sta-
mens 3; pistillateflowerheads 2-6; AmazonianBrazil. ..
.......................................... 34. C. sprucei.
76. Stipules3-20 cm long;widespread. .............. 42. C. villosa.
65. Laminabeneath without arachnoidindumentum,or if presentthen sparseand
soon disappearing.
78. Leafy twigs (rather)abruptlythickened and with wide cavities; stamen 1;
widespread. ............................................. 4. C. asperifolia.
78. Leafy twigs graduallythickened, solid or with narrowcavities; stamens 2
or 3.
79. Lateralveins 4-10(-11) pairs.
80. Laminaalmost as long as broad;pair of main basal veins reaching
the marginfar above the middle of the lamina;most lateralveins
branched;Pacificcoastalregionof Ecuadorand Colombia. ......
............................................... 41. C. vannifolia.
80. Laminamostly distinctlylongerthan broad,or if almost as long as
broadthen only the basal pair of lateralveins branched.
81. Hairs on the smallerveins beneathappressed;Guianas. .....
........................................ 26. C. microcephala.
81. Hairs on the smallerveins beneathat least partlypatent.
82. Stamen 1; pistillate inflorescencespluricapitateand the
commonpeduncle2-3 cm long and 2-4 cm thick,perianth
of the pistillate flower(almost)glabrous;Amazon Basin.
...................................... 39. C. tessmannii.
82. Stamens 2 or 3; pistillateinflorescencesunicapitate,or if
pluricapitatethen the common pedunclelongerthan 3 cm,
less than 2 mm thick;and/or the perianthof the pistillate
flowerdensely puberulous.
83. Stamens3; pistillateinflorescencewith 6-10 heads, 5-
9 (in fruitup to 15) mm in diameterand the common
peduncle 3-6 mm long; Pacific coastal region of Ec-
uadorand Colombia. ................. 19. C. herthae.
83. Stamens2; pistillateinflorescenceswith 1-3 heads,or
if more then the heads 3-5 mm in diameterand the
common peduncle 1-3 cm long.
84. Stamensslightlylongerthan the perianth;pistil-
Coussapoa 23
late flower heads 1-3; perianth of the pistillate
flowerglabrous;easternBrazil. .. 33. C. pachyphylla.
84. Stamensdistinctlylonger than the perianth;pis-
tillateflowerheads3-7, or if less then the perianth
of the pistillateflowerdensely puberulous.
85. Staminateflowerheads2-3 mm in diameter;
pistillate flower heads 3-7, and 2-5 mm in
diameter; Venezuela (Bolivar), Brazil (Ro-
raima),AmazonianBoliviaandEcuador,and
Panama. ................. 1. C. crassivenosa.
85. Staminateflowerheads 5-10 mm in diame-
ter;pistillateflowerheads 1(-3)and 5-10 mm
in diameter;widespread. ....... 42. C. villosa.
79. Lateralveins at least 10 pairs.
86. Stamen 1; pistillate inflorescencespluricapitateand the common
peduncle2-3 cm long and 2-4 cm thick; perianthof the pistillate
flower(almost)glabrous;Amazon Basin. ......... 39. C. tessmannii.
86. Stamens2 or 3; pistillateinflorescencesunicapitate,or if pluricap-
itate then the common pedunclelongerthan 3 cm, less than 2 mm
thick,and/orthe perianthof the pistillateflowerdenselypuberulous.
87. Stamens 3; pistillateflowerheads 6-10; Pacificcoastal region
of Ecuadorand Colombia. ..................... 19. C. herthae.
87. Stamens2; pistillateflowerheads 1(-3); widespread.........
........................................ 42. C. villosa.

1. Key to the CoussapoaSpeciesin CentralAmericaandMexico

1. Interfloralbractsabsent.
2. Laminabeneathglabrousor with sparseindumentum.
3. Lateralveins 3-6 pairs,the basal pair reachingthe marginabove or at the middle of the lamina.
.. ............... ....................................................... 34. C. parviceps.
3. Lateralveins 7-13 pairs, the basal pair reachingthe marginbelow the middle of the lamina.
4. Stipules 1-2 cm long; lamina 5-13 x 4-7 cm. ........................... 23. C. macerrima.
4. Stipules2-3 cm long; lamina 13-20 x 9-13 cm. ................. See under 32. C. ovalifolia.
2. Laminabeneathwith dense indumentum.
5. Intercostalvenation prominentbeneath;stipuleswith brownishindumentum(unicellularhairs
only). ........................................................ .. .... 39. C. nymphaeifolia.
5. Intercostalvenation plane beneath;stipuleswith whitish indumentum(unicellularhairs only!).
........................................................................... 6. C brevipes.
1. Interfloralbractspresent.
6. Laminabeneathwith dense indumentum,partlyarachnoidindumentum.
7. Arachnoidindumentumon the lamina brownish. .......................... 11. C. crassivenosa.
7. Arachnoidindumentumon the lamina whitish.
8. Basal pair of lateralveins unbranched. ................................. 30. C. oligocephala.
8. Basal pair of lateralveins branched.
9. Stamens3; (common)peduncleof the pistillateinflorescence1-2 cm long, pistillateflower
heads ellipsoid to obovoid. ...................................... 39. C. nymphaeifolia.
9. Stamens2; (common) peduncleof the pistillateinflorescence(1-)2-13 cm long, pistillate
flowerheads (sub)globose. ............................................. 42. C. villosa.
6. Laminabeneathglabrousor with sparseindumentum,arachnoidindumentumlacking.
10. All or most lateralveins branched;leafytwigsmoreor less abruptlythickened,with wide cavities. C
......... . ..................................................... asperifolia.
10. All or most lateral veins unbranched;leafy twigs graduallythickened, solid or with narrow
cavities.
11. Lateralveins 2-3 pairs. ............................................ 18. C. glaberrima.
11. Lateralveins 4-6 pairs.
12. Basal pair of lateralveins unbranched;pistillateinflorescencesunicapitate..........
...................................... ............................ 36. C. purpusii.
12. Basal pair of lateralveins (faintly)branched;pistillateinflorescencespluricapitate...
................................................................. 15. C. echinata.
24 Flora Neotropica

2. Key to the Coussapoa Species in Colombia

1. Interfloralbractsabsent.
2. Lateralveins 2-8 pairs, basal pair reachingthe marginabove (or at) the middle of the lamina.
3. Basal pair of lateralveins unbranched;stamen 1; pistillateflowersfree. ........ 31. C. orthoneura.
3. Basal pair of lateralveins branched;stamens 3; pistillateflowerscohere. .........34. C. parviceps.
2. Lateralveins at least 8 pairs, or if less then basal pair reachingthe marginbelow the middle of the
lamina.
4. Laminain the areolesbeneathglabrousor with sparsehairs.
5. Stipules2-3 cm long. .................................................... 46. C. valaria.
5. Stipules4-13 cm long.
6. Base of the lamina cordate;basal pair of lateralveins reachingthe marginabove or at the
middle of the lamina. .............................................. 5. C. batavorum.
6. Base of the lamina (sub)obtuseor roundedto truncate;basal pair of lateralveins reaching
the marginbelow the middle of the lamina ........................... 27. C. napoensis.
4. Laminain the areolesbeneathwith dense, minute hairs.
7. Indumentumof the petiole consistingof dense, minute hairs and distinctlylonger,yellowish
hairs. .................................................................... C. 28. nitida.
7. Indumentumof the petiole consistingof short hairs, not distinctlydifferentin length.
8. Indumentumof the leafy twigs and stipules(unicellularhairs only) brown(ish);petiole 5-
7 mm thick. ..................................................... 8. C. cinnamomea.
8. Indumentumof the leafy twigs and stipules (unicellularhairs only) white or yellowish;
petiole 2-4 mm thick.
9. Intercostalvenation almost plane;pistillateinflorescencesunbranched ... 6. C. brevipes.
9. Intercostalvenation prominent;pistillateinflorescencesbranched .... 45. C. tolimensis.
1. Interfloralbractspresent.
10. Laminatrinervate........................................................... 40. C. trinervia.
10. Laminapinnatelyveined.
11. Laminabeneathwith arachnoidindumentum.
12. Arachnoidindumentumon the lamina beneathbrown(ish).
13. Lateralveins 4-8(-11) pairs;pistillate inflorescencespluricapitate;staminateflower
heads ca. 2-3 mm in diameter ................... .............. 11. C. crassivenosa.
13. Lateralveins (7-)10-24 pairs;pistillateinflorescences(usually)unicapitate;staminate
flowerheads 4-10 mm in diameter.
14. Commonpeduncleof the staminateinflorescence1-2 cm long,thatof the pistillate
inflorescence0.5-0.8 cm long; pistillateflowerheads ellipsoid to clavate. .....
........................... ................................... 14. C. duquei.
14. Commonpeduncleof the staminateinflorescence2-6 cm long,thatof the pistillate
inflorescence(1-)2-13 cm long; pistillateflowerheads (sub)globose. .. 42. C. villosa.
12. Arachnoidindumentumon the lamina beneathwhitish.
15. Stamen 1;pistillateinflorescencepluricapitateandthe commonpeduncle2-3 cm long,
2-4 mm thick;perianthof the pistillatefloweralmost glabrous...... 39. C. tessmannii.
15. Stamens2 or 3; pistillateinflorescenceunicapitate,or if pluricapitatethenthe common
pedunclelongerthan 3 cm, less than 2 mm thick, and/or the perianthof the pistillate
flowerdensely puberulous.
16. Stamens 3; pistillateflowerheads 6-10. ......................... 19. C. herthae.
16. Stamens 2; pistillateflowerheads 1(-3). ........................ 42. C. villosa.
11. Laminabeneathwithout arachnoidindumentum.
17. Lamina beneath with domatium-likecavities in the axils of the lateralveins; interfloral
bractslargeand peltate.................................. ............. 10. C. contorta.
17. Lamina beneath without domatium-likecavities; interfloralbracts relatively small and
spathulateto subpeltate.
18. Stipulesglabrousor sparselypuberulous ............................ 34. C. parviceps.
18. Stipuleswith + dense indumentum.
19. Basal lateralveins distinctlybranched.
20. Laminaabout as long as broad;lateralveins 5-9 pairs. ..... 41. C. vannifolia.
20. Laminadistinctlylongerthan broad;lateralveins 8-12 pairs.... 19. C. herthae.
19. Basal lateralveins unbranched(or sometimes with 1-2 branches).
21. Lateralveins 5-6 pairs;intercostalvenation inconspicuous .............
................................................... 9. C. cinnamomifolia.
21. Lateralveins 7-12 pairs;intercostalvenation conspicuous.
22. Stipules0.3-1.3 cm long. ............................ 35. C. parvifolia.
22. Stipules 1.5-2.5 cm long.
Coussapoa 25

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

cences unbranched. .............................. 7. C. chocoensis.
23. Lateralveins 5-10 pairs;petiole 5-10 cm long; pistillate inflores-
cences branched. ............................... 44. C.fulvescens.

3. Key to the Coussapoa Species in Venezuela

1. Interfloralbractsabsent.
2. Stipules0.5-1 cm long; stamens 2; pistillateflowerheads 1-3. .................... 43. C. viridifolia.
2. Stipules(0.5-)1-7 cm long; stamen 1; pistillateflowerheads 2-10. ............... 31. C. orthoneura.
1. Interfloralbractspresent.
3. Laminatrinervate ............................................................. 40. C. trinervia.
3. Laminapinnatelyveined.
4. Laminabeneathwith arachnoidindumentum.
5. Arachnoidindumentumon the lamina brownish . ...................... 11. C. crassivenosa.
5. Arachnoidindumentumon the lamina whitish.
6. Lateralveins 2-3(-4) pairs ........ ..................................... 3. C. argentea.
6. Lateralveins 4-24 pairs . .................. ........................... 42. C. villosa.
4. Laminabeneathwithout arachnoidindumentum.
7. Leafytwigs (rather)abruptlythickenedwith wide cavities;stamen 1 ......... 4. C. asperifolia.
7. Leafytwigs graduallythickenedsolid or with narrowcavities.
8. Laminawith patenthairs on the venation beneath;stipules 1-5 cm long. .............
....................................................11. C. crassivenosa.
8. Laminawith appressedhairs on the venation beneath;stipules 0.3-1(-1.3) cm long.
9. Lateralveins 7-11 pairs;terminalbudsusuallyswollen;marginof the lamina ? revolute
at the base. ....................................... .... 35. C. parvifolia.
9. Lateralveins 4-7 pairs;terminalbuds slender;marginof the lamina not revolute at
the base ........................................................ 43. C. viridifolia.

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

1. Laminaabove scabrous.
2. Basal lateralveins reachingthe marginabove the middle of the lamina. ............4. C. asperifolia.
2. Basal lateralveins reachingthe marginbelow the middle of the lamina ............. 21. C. leprieurii.
1. Laminaabove smooth.
3. Laminabeneathwith arachnoidindumentum.
4. Arachnoidindumentumon the lamina brownish.
5. Laminabeneathwith patenthairs on the venation.
6. Lateralveins 4-5 pairs;lamina oblong. ............................... 16. C.ferruginea.
6. Lateralveins (4-)5-8 pairs;lamina mostly elliptic to ovate to obovate or to suborbicular.
..........................1.............. ...... . . .. 1 . C. crassivenosa.
5. Laminabeneathwith appressedhairs on the venation .................. 26. C. microcephala.
4. Arachnoidindumentumon the lamina whitish.
7. Lateralveins 2-3(-4)pairs. ......................... ... .... ................. 3.C. argentea.
7. Lateralveins 4-11 pairs. .......................... .................. 26. C. microcephala.
3. Laminabeneathwithout arachnoidindumentum.
8. Lateralveins 2-4 pairs. .................. ..... ........ .................... 1. C. angustifolia.
8. Lateralveins 4-11 pairs.
9. Leafytwigs (rather)abruptlythickenedwith wide cavities; stamen 1. .........4. C. asperifolia.
9. Leafytwigs graduallythickenedand solid or with narrowcavities.
10. Lateralveins 7-11 pairs;terminalbuds usuallyswollen. .............. 35. C. parvifolia.
10. Lateralveins 4-7 pairs;terminalbuds slender.
11. Hairs on the lamina beneathof about equal length .................. 20. C. latifolia.
11. Hairs on the lamina beneathof differentlengths ... ................ 43. C. viridifolia.

5. Key to the Coussapoa Species in Ecuador

1. Interfloralbractsabsent.
2. Basal pair of lateralveins unbranched(or occasionallya single branch).
3. Lateralveins 2-7 pairs,the basal pair reachingthe marginabove to just below the middle of the
lamina. ................................................................. 3 1. C. orthoneura.
26 Flora Neotropica

3. Lateralveins (5-)7-21 pairs, the basal pair reachingthe margin (far) below the middle of the
lamina.
4. Stipules 1-4 cm long; apex of the lamina mostly acute to shortlyacuminate. . 32. C. ovalifolia.
4. Stipules4-13 cm long; apex of the lamina mostly obtuse. .................. 27. C. napoensis.
2. Basalpair of lateralveins branched.
5. Stipulesglabrousor sparselypuberulous .....................................34. C. parviceps.
5. Stipulessubvelutinous,subhirsute,or subsericeous.
6. Stipules 1-4 cm long; apex of the lamina mostly acute to shortlyacuminate. .. 32. C. ovalifolia.
6. Stipules4-13 cm long; apex of the lamina mostly obtuse. .................. 27. C. napoensis.
1. Interfloralbractspresent.
7. Laminatrinervateor only the basal pair of lateralveins strongand the other lateralveins weak and
inconspicuous.
8. Laminatrinervate;intercostalvenation prominentbeneath. .... ............ 40. C. trinervia.
8. Lamina with only the basal pair of lateral veins strong and the other lateral veins weak and
inconspicuous;intercostalvenation plane beneath. ............... ........9. C. cinnamomifolia.
7. Laminapinnatelyveined.
9. Laminabeneathwith arachnoidindument.
10. Arachnoidindumentumon the lamina beneathbrown(ish).
11. Lateralveins 4-8(1 1) pairs;pistillateinflorescencespluricapitate;staminateflowerheads
ca. 2-3 mm in diameter. ........................................ 11. C. crassivenosa.
11. Lateralveins (7-)10-24 pairs; pistillate inflorescences(usually unicapitate;staminate
flowerheads 4-10 mm in diameter.
12. Commonpeduncleof the staminateinflorescence1-2 cm long, that of the pistillate
inflorescence0.5-0.8 cm long; pistillateflowerheads ellipsoid to clavate. .......
...................................... . ........................ . C. duquei.
14.
12. Commonpeduncleof the staminateinflorescence2-6 cm long, that of the pistillate
inflorescence(1)2-13 cm long; pistillateflowerheads (sub)globose.... 42. C. villosa.
10. Arachnoidindumentumon the lamina beneathwhitish.
13. Stamens2; pistillateflowerheads 1(-3). ................................. 42. C. villosa
13. Stamens 3; pistillateflowerheads 3-10.
14. Common peduncle 8-20 cm long; base of the lamina cordateto rounded........
...................................................... 22. C longepedunculata.
14. Common peduncleup to 7 cm long;base of the lamina usuallyobtuse to rounded.
............................................................... 19. C. herthae.
9. Laminabeneathwithout arachnoidindumentum.
15. Laminabeneathwith domatium-likecavitiesin the axils of the lateralveins;interfloralbracts
largeand peltate. ...................................................... 10. C. contorta.
15. Laminabeneathwithoutdomatium-likecavities;interfloralbractsrelativelysmalland spath-
ulate to subpeltate.
16. All lateralveins usually branched;pistillateinflorescencesusually unicapitate;stamen
1.......... ..................................... . ..1. ..C. asperifolia.
16. Only the basal pair of lateralveins usuallybranched;pistillateinflorescencespluricap-
itate; stamens 3.
17. Stipulesglabrousor sparselypuberulous ............................... 34. C. parviceps.
17. Stipuleswith dense indumentum.
18. Laminaabout as long as broad;lateralveins 5-9 pairs. ....... 41. C. vannifolia.
18. Laminadistinctlylongerthan broad;lateralveins 8-12 pairs.
19. Common peduncle8-20 cm long;base of the lamina cordateto rounded.
..................................... 22. C. longepedunculata.
19. Common peduncleup to 7 cm long;base of the lamina usuallyobtuse to
rounded. .............................................. 19. C. herthae.

6. Key to the CoussapoaSpecies in Peru and Bolivia

1. Interfloralbractsabsent.
2. Basal lateralveins unbranched(or occasionallywith a single branch).
3. Lateralveins 7-21 pairsand the laminabeneath(almost)glabrousin the areoles. .. 32. C. ovalifolia.
3. Lateralveins 2-9 pairs and/or the lamina beneathwith dense, minute hairs in the areoles.
4. Basallateralveins reachingthe marginabove or just below the middle of the lamina;stipules
mostlywithoutor with sparselong yellowhairs;branchesof the pistillateinflorescenceslightly
broadenedtowardsthe flowerheads ..................................... 31. C. orthoneura.
4. Basallateralveins reachingthe marginbelow to just above the middle of the lamina;stipules
Coussapoa 27

with dense, long, yellow hairs; branches of the pistillate inflorescencestronglybroadened

towardsthe flowerheads ................................................ 12. C. cupularis.
2. Basal lateralveins branched.
5. Lateralveins 5-9 pairs;lamina usuallyelliptic. ........................... See 12. C. cupularis.
5. Lateralveins 8-15 or 14-22; lamina usuallyovate.
6. Indumentumof the petiole consistingof dense minute hairs and distinctlylonger,yellowish
hairs;lateralveins 8-15 pairs;peduncleof the pistillateinflorescence1-2 mm thick;Amazon
Basin. ........................... .................................... 28. C. nitida.
6. Indumentumof the petiole consistingof short hairs, not differingdistinctlyin length;lateral
veins 14-22 pairs; peduncle of the pistillate inflorescence6-10(-15) mm thick; Colombia
(Amazonas). ........................................................ 8. C. cinnamomea.
1. Interfloralbractspresent.
7. Laminatrinervateor only the basal pair of lateralveins strongand the other lateralveins weak and
inconspicuous.
8. Laminatrinervate;intercostalvenation prominentbeneath. .................... 40. C. trinervia.
8. Lamina with only the basal pair of lateral veins strong and the other lateral veins weak and
inconspicuous;intercostalvenation plane beneath. ....................... 9. C. cinnamomifolia.
7. Laminapinnatelyveined.
9. Laminabeneathglabrousor with sparseindumentum.
10. Basal lateral veins unbranched;stipules up to 1(-1.3) cm long; margin of the lamina +
revoluteat the base. .................................................. 35. C. parvifolia.
10. Basaland/or other lateralveins usuallybranched;stipulesup to 3.5 cm long;marginof the
lamina not revolute at the base. ......................................... 4. C. asperifolia.
9. Laminabeneathwith dense indumentum.
11. Laminabeneathwithout arachnoidindumentumand basal pair of veins unbranched.....
.................................................................... 24. C. manuensis.
11. Laminabeneathwith arachnoidindumentumand basal pair of lateralveins branched.
12. Stamens3; common peduncle8-20 cm long; pistillateflowerheads 3-10 ...........
............... ........................................... 22. C. longepedunculata.
12. Stamens2 or 1;common peduncleup to 6 cm long, or if longerin pistillateinflorescences
then a single flowerhead.
13. Stamens2; pistillateinflorescencesunicapitate,or if pluricapitatethen the perianth
of the pistillateflowerdensely puberulous. ......................... 42. C. villosa.
13. Stamen 1; pistillateinflorescencespluricapitateand perianthof the pistillateflower
almost glabrous. ........................................ ... 39. C. tessmannii.

7. Key to the Coussapoa Species in Amazonian Brazil

1. Interfloralbractsabsent.
2. Lateralveins 2-8 pairs.
3. Stipules0.5-1 cm long;stamens2; pistillateflowerheads 1-3 and common peduncleor branches
not thickenedbelow these heads. ........................................... 43. C. viridifolia.
3. Stipulesusuallylongerthan 2 cm, if shorterthan 1 cm then stamen 1 or pistillateflowerheads
2-10 and the branchesof the inflorescencemore or less thickenedbelow these heads.
4. Intercostalvenation prominentbeneath;laminabeneathand inflorescenceswith dense, white
arachnoidindumentum;LowerAmazon Basin. .......................... 2. C. arachnoidea.
4. Intercostalvenationplanebeneath;laminaandinflorescenceswithoutarachnoidindumentum,
or if presentthen mostly soon disappearing;Upper Amazon Basin.
5. Basal lateral veins reachingthe margin above to just below the middle of the lamina;
stipulesmostly without or with sparse,long, yellow hairs;branchesof the pistillateinflo-
rescenceslightlythickenedbelow the flowerheads. .................... 31. C. orthoneura.
5. Basal lateral veins reachingthe margin below to just above the middle of the lamina;
stipules with dense, long, yellow hairs; branchesof the pistillate inflorescencestrongly
thickenedbelow the flowerheads. .................................... 12. C. cupularis.
2. Lateralveins (7-)9-22 pairs.
6. Basal lateralveins (mostly unbranched;lamina beneath in the areolesglabrousor sparselypu-
berulous. ................................................................. 32. C. ovalifolia.
6. Basal lateralveins branched;lamina beneathin the areoleswith dense, minute hairs.
7. Petiole 2-4 mm thick, with dense, minute hairs and sparse,much longer,yellowishhairs.
......................................................................... 28. C. mitida.
7. Petiole 5-7 mm thick, with short hairs about the same in length. ......... 8. C. cinnamomea.
28 Flora Neotropica

1. Interfloralbractspresent.
8. Laminatrinervate ............................................................. 40. C. trinervia.
8. Laminapinnatelyveined.
9. Laminascabrousabove.
10. Lateralveins unbranched. ................................................ 37. C. scabra.
10. All or most lateralveins branched .......................................4. C. asperifolia.
9. Laminasmooth above.
11. Lamina trinervate. ..................................................... 40. C. trinervia.
11. Laminapinnatelyveined.
12. Laminabeneathwith dense, partlyarachnoidindumentum.
13. Arachnoidindumentumon the lamina brownish.
14. Lateralveins 4-5 pairs;lamina oblong. ..................... 16. C.ferruginea.
14. Lateralveins (4-)5-8 pairs;lamina mostly elliptic to ovate to obovate or to
suborbicular............................................ 11. C. crassivenosa.
13. Arachnoidindumentumon the lamina whitish.
15. Stamens 2; pistillate inflorescences(usually)unicapitate;perianthof the pis-
tillate flowerdensely puberulous. .............................. 42. C. villosa.
15. Stamen 1;pistillateinflorescencespluricapitate;perianthof the pistillateflower
almost glabrous. ........................................ 39. C. tessmannii.
12. Laminabeneathglabrousor with sparseindumentumand arachnoidindumentumlack-
ing.
16. Lateralveins 2-4 pairs. .......................................1. C. angustifolia.
16. Lateralveins 4-11 pairs.
17. Lateralveins 7-11 pairs;terminalbuds usuallyswollen. ...... 35. C. parvifolia.
17. Lateralveins 4-7 pairs;terminalbuds slender.
18. Leafytwigs (rather)abruptlythickenedand with wide cavities;stamen 1.
...................................................... . C. asperifolia.
18. Leafytwigs graduallythickenedand solid or with narrowcavities.
19. Hairs on the lamina beneathdistinctlydifferentin lengths. ........
................................................. 43. C. viridifolia.
19. Hairs on the lamina beneathabout equal in length ..... 20. C. latifolia.

8. Key to the CoussapoaSpecies in EasternBrazil

(Rio Grande do Sul to Pernambuco)
1. Stipulesup to 1 cm long. ........................................... 13. C. curranii.
1. Stipulesat least 1 cm long.
2. Laminaabove with subpersistent,white, arachnoidindumentum. ................... 17. C. floccosa.
2. Laminaabove glabrous.
3. Stamensmuch longer than the perianth;pistillateinflorescencesmostly unicapitate,(common)
peduncle 1-2.5 cm long; stipulesup to 7 cm long;leaf marginusuallyrevolutetowardsthe base.
........................................................................ 25. C. microcarpa.
3. Stamensas long as the perianth;pistillateinflorescencesmostly pluricapitate,common peduncle
2.5-4 cm long; stipules up to 2 cm long; leaf marginentirelyrevolute or plane...............
....................................................................... 33. C. pachyphylla.

1. CoussapoaangustifoliaAublet, Hist. pl. Gui-

to subobtuse),marginentire;upper surfacegla-
ane 2: 956, t. 363. 1775;Miquel,Fl. bras.4(1):
138. 1853;Berg.Fl. Suriname5(1): 283. 1975. brous, lower surface glabrous or sparsely ap-
Type. French Guiana. Without locality, -, pressed-puberulous; lateral veins 2-4 pairs,
Aublets.n. (BM, leaf fragment). straightor curved,basalpairunbranched,reach-
Fig. 10.
ing the marginfar above the middle of the lam-
Shrubor tree,mostly hemi-epiphytic,up to 30 ina;intercostalvenationmostly plane;petiole 1-
m tall. Leafy twigs 3-8 mm thick, glabrousor 5 cm long, glabrous or sparsely appressed-pu-
sparsely,minutely puberulous,lenticels numer- berulous; stipules 0.5-1 cm long, glabrous or
ous and conspicuous. Lamina coriaceous ob- sparselyappressed-puberulous. Staminate inflo-
ovate or subobovate (to oblanceolate or ellip- rescences poorly branched, heads 3-5, some-
tic to oblong), 2-11 x 1-6 cm, apex obtuse to times partlyfused,globose, 5-8 mm diam.;com-
roundedor emarginate,base acuteto cuneate(or mon peduncle 0.5-1.5 cm long, minutely
Coussapoa 29

FIG. 10. Coussapoaangustifolia:1, leafy twig with pistillate inflorescences(Schulz 10344a); 2, staminate
inflorescences(Pireset al. 50631).

bracts spathulate to subpeltate, apex

puberulous;perianth 1 mm high, glabrous;sta- floral
men one, as long as the perianth.Pistillateinflo- ciliolate.
rescencesunbranchedor rarelybranched;heads Distribution(Fig. 6). Surinam,FrenchGuiana,
1(-2), globose, ca. 5 mm, in fruit up to 20 mm and Brazil(Amapaand Para);in uplandand riv-
diam.; (common) peduncle 0.3-3 cm mi- erine forest.
long,
nutely puberulous; perianth 1(-2) mm high, gla- Specimens studied.SURINAM.Brownsberg, 6 Sep
brous;fruitingperianthyellow to orange.Inter- 1915 (e), BW 772 (A, K, NY, U); BrokopondoLake,
30
10 Sep 1965 (st), VanDonselaar2569 (U); Wilhelmina
Mts., Zuid:River, 45 km above confluencewith Lucie
River, 25 Sep 1963 (9), Irwinet al. 57574 (B, G, MO,
NY, OXF, R, S, U); CoppenameRiver, Bitagron,28
Jul 1954 (9),Lindeman6362 (F, U); ZanderijSavanna,
28 Aug 1954 (st), Lindeman 6545 (U); Zanderij, 15
Apr 1954 (st), Lindeman 6600 (MO, U); Saramacca
River, Jacobkondre,19 Jun 1944 (6), Maguire23867
(A, F, K, NY, U, US); SaramaccaRiver, nr. Kwat-
tahede,22 Jun 1944 (2), Maguire23942 (A, F, K, MO,
NY, P, RB, U); WilhelminaMts., S of Julianatop,7
Aug 1963 (8),Schulz10344a (NY, U); SurinameRiver,
Gansee, 8 Jul 1908 (9), Tresling55 (U).
FRENCH GUIANA. Without locality, - (st), Au-
blet s.n. (BM, type collection);Itany River, island in
rapids, 30 Sep 1961 (9), BAFOG(=Graimleux) 7938
(CAY, U); MaroniRiver, 15 Sep 1903 (9), Geay 3285
(P), OyapockRiver,nr. GrandRoche Fall, 17 Jul 1960
(9), Maguireet al. 47049 (G, GH, LE, NY, OXF, U,
US); nr. Cayenne, - (9), Martin s.n. (BM, BR, K);
MaroniRiver, 1854 (9),Melinon 172 (LE,P); Camopi
River, below Grand Tamouri Creek, 2 Feb 1968 (2),
Oldemanet al. 207 (CAY,P, U, US);SinnamaryRiver,
1.5 km above CourciboRiver, 7 Aug 1967 (9), Olde-
man B1167 (CAY);KourouRiver, 2 km above Cr6ole,
18 Sep 1967 (9), Oldeman B1324 (CAY, U); Oropu
River, 1.5 km above DegradLalande,16 Oct 1965 (9),
Oldeman1619 (CAY);ApprouagueRiver, 5 km above
MaripaCreek, 3 Feb 1967 (9), Oldeman2476 (CAY,
P, U); OyapockRiver, above MoutouciFalls, 15 May
1970 (6), OldemanB3243 (CAY, U); Sinnamary,road
to St. Elie, 18 Jul 1977 (6), Sastre 5500 (P, U).
BRAZIL. AMAPA: Rio Araguari,between 1055'N, between1?55'N,51059'Wand2?5'N,51056'W,1 Sep
51?51'W and 2?5'N, 51056'W, 1 Sep 1961 (9), Pires et 1961(v),Pireset al. 50656(IAN,M, MG,NY, OXF,
al. 50631 (NY, U, US). PARA:Mun. Oriximina,road US, VEN,typecollection).
to CachoeiraPorteira,18 Jun 1980 (9), Cid et al. 1053
(U); Rio Paru do Oeste, CachoeiraChuvisco, 17 Sep
1980 (9), Cid et al. 2223 (U), Ilhas de Breves, Furo
Macujubim,16 Nov 1922 (9), Ducke (HJBR) 18461
(RB);Rio Trombetas,CachoeiraPorteira,8 Jun 1978
(9), N. T. Silva et al. 4757 (U).
This species, with few lateral veins in a mostly
subobovate lamina, appears to be related to C.
trinervia. In its leaf characters C. angustifolia is
transitional to the trinervate state of the latter
species. It also shows many similarities to the
Central American C. glaberrima.
2. Coussapoa arachnoidea Akkermans & C. C.
Berg, Proc. Kon. Ned. Akad. Wetensch. Ser.
C, 85(4): 442. 1982. Type. Brazil. Amapa: Rio
Araguari, between 1?55'N, 51?59'W and 2?5'N,
51?56'W, 1 Sep 1961 (2), Pires et al. 50656
(holotype, IAN; isotypes, M, MG, NY, OXF,
US, VEN). Fig. 11.
Tree. hemi-epiphytic, up to 35 m tall. Leafy
twigs 3-8 mm thick, with sparse, arachnoid hairs
and yellowish, straight hairs on the scars of the
FloraNeotropica
stipules. Lamina coriaceous,elliptic to ovate or
to lanceolate,(6-)9-16 x (2-)3-7 cm, apexacute,
base obtuseto acute,marginentire;uppersurface
glabrous, lower surface in the areoles rather
densely,minutelypuberulous,on the main veins
sparse,ratherlong, yellowish hairs, initially the
whole surfacedenselycoveredwith white arach-
noid hairs;lateralveins 5-8 pairs,almoststraight,
basalpairunbranched,reachingthe marginat or
just belowthe middle;intercostalvenationrather
prominent;petiole3-6 cm long,villouswithwhite
arachnoidhairs,glabrescent;stipules5-8 cm long,
villous with white arachnoidhairs, mixed with
ratherlongand straightappressed,yellowishhairs
and with reddishbrownpluricellularhairs. Sta-
minate inflorescencesunknown. Pistillate inflo-
rescencesbranched;heads 2-4, globose ca. 4-5
mm diam.; common peduncle 1.5-2 cm long,
denselyvillous with yellowishto white arachnoid
hairs, mixed with straight appressedyellowish
hairsand with reddish-brownpluricellularhairs,
apices of the branches broadened towards the
heads; perianth ca. 1 mm high, minutely pu-
berulous.Interfloralbractsabsent.
Distribution (Fig. 6). Brazil (Amapa).
Specimensstudied.BRAZIL.AMAPA:Rio Araguari,
Coussapoaarachnoideaappearsto be closely
relatedto C. orthoneura,from which it differsin
the prominentintercostalvenationand the dense
covering of white arachnoidhairs on the leaves
and inflorescences.Staminateinflorescencesare
needed to determine the validity or not of the
presumedrelationship.It may prove to be dis-
tinct from C. orthoneuraonly at the subspecific
level.
3. CoussapoaargenteaAkkermans& C. C. Berg,
Proc.Kon. Ned. Akad.Wetensch.Ser.C, 85(4):
443. 1982. Type. Venezuela.Bolivar: 119 km
S of El Dorado, 12 Jan 1964 (6), Steyermark
et al. 92991 (holotype,VEN; isotypes,NY, U,
US). Fig. 12.
Tree,hemi-epiphyticor terrestrial,up to 30 m
tall. Leafy twigs 3-5 mm thick, brownish pu-
berulousto strigoseor partlyto hirsuteand with
brown to white arachnoidhairs. Lamina coria-
ceous, obovate to elliptic, occasionallyovate or
Coussapoa 31

FG. 11. Coussapoa1 ly tg wh p e is (s et cm.

FIG. 11. Coussapoaarachnoidea:1, leafy twig with pistillateinflorescences(Pireset al. 50656).

oblong,2-12 x 1-6 cm, apex obtuse to acute or ering of silvery arachnoidhairs, deciduous with
acuminate, base obtuse, margin entire; upper age; lateral veins 2-3(-4) pairs, slightly curved,
surfaceglabrous,lowersurfacesparselyto dense- basalpairunbranched,reachingthe marginabove
ly appressed-puberulousand with a dense cov- the middle of the lamina; intercostal venation
32 Flora Neotropica

c.

FIG. 12. Coussapoaargentea:1, leafy twig with pistillateinflorescences(Steyermark&Nilsson 487); 2, leafy

twig with staminateinflorescences(Steyermark& Nilsson 388).
Coussapoa
slightly prominent to almost plane; petiole 1-2
cm long,brownishpuberulousto strigoseor part-
ly to hirsuteand with brown to white arachnoid
hairs;stipules 1-5 cm long, denselycoveredwith
brownishcurled to arachnoidhairs and whitish
straighthairs. Staminate inflorescencesrepeat-
edly branched;headsmany,globose,ca. 2-3 mm
diam.; common peduncle 2-3 mm long, puber-
ulous to hirtellous; perianth ca. 1 mm high,
3-lobed, glabrous;stamens two, exceeding the
perianth. Pistillate inflorescencesunbranched;
head globose, ca. 4 mm diam.; peduncle 2-3.5
cm long, puberulous,and with silvery arachnoid
hairs; perianthca. 1 mm high, glabrous.Inter-
floral bractsspathulateto subpeltate,sometimes
few.
Distribution (Fig. 6). Venezuela (Bolivar);
mostly in moist forest on table mountains, be-
tween (450-)1000 and 1300 m.
Specimensstudied.VENEZUELA. BOLIVAR: Road
El Dorado-SantaElena,km 109, 23 Feb 1959 (6), Ber-
nardi7238 (G); upperRio Cuyuni,Cumbrede La Es-
calera, 1000 m, 20-21 Aug 1962 (6), Maguire et al.
46879 (U, US, VEN); nr. CerroUei, between Luepa
and CerroVenamo, 1100 m, 20-22 Apr 1960 (6),Stey-
ermark& Nilsson 388 (NY, US, VEN);NE of Luepa,
1300 m, 23 Apr 1960 (v), Steyermark& Nilsson 487
(NY, VEN); 119 km S of El Dorado, 12 Jan 1964 (6),
Steyermarket al. 92991 (NY, U, US, VEN, type col-
lection).
Coussapoa argentea is similar to C. micro-
cephalain manyfeatures.The speciesdiffersfrom
the latterin the smallernumberof lateralveins,
the basalpairof which reachesthe marginbelow
the middle of the lamina.Moreover,the pistillate
inflorescenceis unicapitate,being normallyplur-
icapitate in C. microcephala.The two species
appear to differ ecologically (altitudinalcondi-
tions) and may prove to be distinct only at the
subspecies level.
4. CoussapoaasperifoliaTrecul, Ann. Sci. Nat.
Bot. Ser. 3, 8: 96. 1847; Miquel, Fl. bras.4(1):
135. 1853; Macbride, Publ. Field Mus. Bot.
13(2.2): 296. 1937; Berg, Fl. Suriname 5(1):
284. 1975. Lectotype.Suriname.Without lo-
cality, 1843 ($), Hostmann 1189 (holotype,P;
isotypes, B, G, K, LE, U).
Tree or shrub, hemi-epiphytic or terrestrial, up
to 25 m tall. Leafy twigs 3-20 mm thick, (usually)
33
abruptlythickenedjust belowthe apexandwidely
hollow, (rather)sparselypuberulousto hispidu-
lous, often also with brown pluricellularhairs,
sometimes partlyhirtellous.Lamina coriaceous
to subcoriaceous,obovate to subobovate to el-
liptic to oblong to suborbicularto ovate or to
cordiform, 7-32 x 4-24 cm, apex rounded to
obtuse to emarginate,base obtuse to roundedto
truncateto cordate,marginentire to subcrenate;
upper surface scabrouswith minute rigid hairs
to smooth and glabrous, lower surface densely
to sparselypuberulouswith curved (to straight)
hairs or only minutely puberulouson the main
veins, sometimes partly hirtellous, in juvenile
leaves sometimes the whole surface his-
pid(ulous);lateralveins 4-9 pairs, usually all or
most of them branched,sometimesthe basalpair
unbranchedor faintlybranchedbut some of the
otherlateralveins poorlybranched,(main)basal
pair reachingthe marginat, below, or above the
middleof the lamina;intercostalvenationprom-
inent to plane; petiole 1.5-15 cm long, sparsely
to denselyminutelypuberulous,sometimespart-
ly hirtellous;stipules 0.5-3.5 cm long, puberu-
lous to hispidulous,often also with brownpluri-
cellular hairs, sometimes partly hirtellous.
Staminate inflorescencesbranched;heads many,
freeorpartlyfused,(sub)globose,3-10 mm diam.;
common peduncle 2-8 cm long, puberulous;
perianthca. 1 mm high, minutelypuberulousor
glabrous;stamenone, as long as or slightlylonger
than the perianth. Pistillate inflorescenceun-
branchedor branched;heads 1-6, free or partly
fused, 3-12 mm, in fruit up to 30 mm diam.;
(common) peduncle 1-9.5 cm long, puberulous
to almost glabrous;perianth 1.2 mm high, mi-
nutely puberulousor glabrous.Interfloralbracts
(sub)spathulateto (sub)peltate,minutely puber-
ulous at the apex.
C. asperifoliadiffersfrom the other Coussapoa
species:(1) The endocarpbody is pushed out of
the perianththroughthe torn apex of the peri-
anth, due to the expansion of the mucilaginous
mesocarp (see p. 8). (2) Furthermore,the leafy
twigs widen ratherabruptlyjust below the apex
and have wide cavities, which areoften occupied
by ants.
Within this widespread and polymorphic
speciesthreesubspeciescan be recognized.These
three subspeciesare not well separatedmorpho-
logicallyor geographically.
34 Flora Neotropica

Key to the Subspecies of Coussapoa asperifolia

1. Lamina scabrousto scabridulousabove, densely (to rathersparsely)puberulouswith curved hairs on
the smallerveins beneath;intercostalvenation prominent;lamina mostly obovate. ....a. ssp. asperifolia.
1. Laminasmooth (to scabridulous)above, glabrousor sparselypuberulouson the smallerveins, sparsely
to ratherdenselyminutelypuberulouswith straighthairson the main veins beneath;intercostalvenation
plane to prominent.
2. All or most lateral veins branched,sometimes poorly so and the basal pair unbranched;lamina
mostly ovate to cordiformor elliptic;pistillateheads mostly 1, more than 10 mm, in fruitup to 35
mm in diam. ........................................................... ..b. ssp. magnifolia.
2. Few or none of the lateralveins (poorly)branched,the basal pair unbranchedor faintly branched;
lamina mostly obovate to subobovate;pistillate heads 3-5, ca. 3-6 mm, in fruit up to 15 mm in
diam ........ ........................................................... c. ssp. rhamnoides.

4a. Coussapoaasperifoliassp. asperifolia
Fig. 13.
CoussapoaficinaStandley,Publ.FieldMus.Bot. 17:
161. 1937.Type.Brazil.Amazonas:Nr. Manaus,
roadto Aleixo,12Aug-1Sep1936(9),Krukoff7966
(holotype,NY;isotypes,A, BM,F, K, LE,MO,S,
U).
CoussapoacayennensisHawkes,Phytologia3: 30. 1948.
Type.FrenchGuiana.Nr.Cayenne,17Jul1921(9),
Broadway880 (holotype,NY; isotypes,GH, NY,
US).
Lamina mostly obovate to subobovate or el-
liptic to oblong, mostly 10-17 cm long, base
mostly obtuse to rounded; upper surface sca-
brouswith a densecoveringof minuterigidhairs,
sometimes scabridulous,lower surface on the
smaller veins mostly densely puberulous with
curved(to straight)hairs;lateralveins 4-7 pairs,
all or most of them branched,the (main) basal
pairreachingthe marginabove the middle of the
lamina; intercostalvenation prominent;petiole
1.5-9 cm long; stipules 0.5-1.5 cm long. Sta-
minate inflorescenceswith heads up to 10 mm
diam.;common peduncleup to 4.5 cm long. Pis-
tillateinflorescenceswith mostly, sometimestwo,
(partlyfused) heads, mostly more than 1 cm, in
fruitup to 2.5 cm diam.; (common) peduncle 1-
3 cm long.
Distribution(Fig. 6). Eastern Guiana region
(Surinamand FrenchGuiana),in Brazilin Ama-
pa and eastern Para and also, apparentlysegre-
gated from the main area, in the centralpart of
the Amazon Basin;in uplandand riverineforest.
This distribution shows striking similarities to
that of C. latifolia.
Specimensstudied.SURINAM.Section0, 11 Nov
1915(st),BW1322(U);Corantijn
River,Kabouri,26
Jun1916(st),BW2070(U);MaratakkaRiver,25 Nov
1917(9),BW3430(F,NY, U, US);Section0, 10Aug
1921 (6), BW 5262 (A, U); without locality, 1843 (8),
Hostmann 1189 (B, G, K, LE, P, U, lectotypecollec-
tion); Lely Mts., 19 Sep 1975 (st), Lindemanet al. 34
(U); ConcessionHaenen, W of mouth of Coppename
River, 25 Mar 1954 (st), Lindeman 5709 (U); upper
Litani River, 9 Aug 1937 (a), Rombouts803 (A, K,
NY, U); PlantationLustrijk,Dec 1837 (8), Splitgerber
448 (=122) (L, P); MaratakkaRiver, 20 Aug 1976 (2),
Teunissen(LBB) 16031 (U).
FRENCH GUIANA. Itany River, island in rapids,
30 Sep 1962 (9), BAFOG(=Graimleux) 7942 (CAY,
U); nr. Cayenne,17 Jul 1921 (9), Broadway880 (GH,
NY, US, type collection of C. cayennensis);Oyapock
River, Les Trois Sauts,27 Mar 1975 (9), Grenand966
(CAY);Anse Bruyere,15 Aug 1978 (a), Grenand1596
(U); nr. Cayenne,- (9), Leman s.n. (P); nr. Cayenne,
Mar 1900 (st), Lemee s.n. (P);without locality, - (9),
Leprieurs.n. (P);Kuataka,AntecumePata, confluence
of Itany River and MarouiniRiver, 14 Nov 1977 (9),
Moretti837 (CAY); ApprouagueRiver, between Sa-
pokaye Creek and Grand Canori Falls, 23 Oct 1968
(9), Oldeman T243 (CAY, GH, NY, P, U); Oyapock
River, between Notaye Creekand Caiman Creek, 10
Jul 1969 (st), OldemanT366 (CAY,U); OyapockRiv-
er, Savane-Roche,Baton Pilon, 12 Jul 1969 (6), Olde-
man B2538 (CAY, U); OyapockRiver, above Oscar,
18 Jun 1970 (9), OldemanB3420 (CAY,P, U); without
locality, 1820 (9),Perrotets.n. (P);Sinnamary,road to
St. Elie, km 8, 28 Aug 1979 (st), Provost 738 (U);
Acarouany,1856 (st), Sagot s.n. (P); nr. Kourou, 14
Dec 1978 (9), Sastre 6417 (CAY, P, U); without lo-
cality, 23 Jun 1921 (9), Wachenheim426 (P).
BRAZIL. AMAPA:Rio Araguari,between 1?2'N,
51?15'Wand 0?57'N,51?29'W,13 Sep 1961 (6), Pires
et al. 50913 (NY, OXF, R, U, US). AMAZONAS:
Nr.
Manaus,BR. 17, ca. km 9, 5 Aug 1955 (9), D. Coelho
(INPA)1577 (U), 8 Jul 1955 (8), Chagas(INPA)1364
(U); nr. Manaus,Colonia Joao Alfredo, 21 Aug 1947
(9), Ducke 2103 (IAN, R, U); nr. Manaus,Pensador,
Aug 1910($), Ule8836 (G, K, MG);Manaus-Caracarai
road,ca. km 130, 13Feb 1974(9),Stewardetal. P.20255
(MO, U). PARA:Res. Florest,Rio Aura, 18 Oct 1978
(9), Carautaet al. 3011 (RB);Belem, 21 Oct 1915 (st),
Ducke (HAMP) 15804 (BM); Mun. Oriximinf, Rio
Mapuera,19 Aug 1986 (a), Ferreiraet al. 7831 (BG);
Arama,2 Nov 1900 (9),Huber(HAMP)1873 (G);Rio
Jan, rd. to Munguba,km 15, 13 Jul 1964 (9), N. T.
Silva 2166 (U).
.".
~:.,, . .
36 Flora Neotropica

4b. Coussapoaasperifoliassp. magnifolia(Tre- Specimens studied. PANAMA. CANALZONE:Fort

cul) Akkermans& C. C. Berg,Proc. Kon. Ned. San Lorenzo, 13 Sep 1947 (a), Alien 5120 (BM, G,
Akad. Wetensch.Ser. C, 85(4): 445. 1982. MO); BarroColoradoIsland, 27 Apr 1968 (a), Croat
5124 (DUKE, F, MO), (9), Croat5125 (F), 1 Oct 1968
Fig. 14. (9), Croat 6588 (MO), 10 Feb 1969 (8), Croat 7859
(MO),13 Jul 1969(6),Foster1101 (DUKE,F), Chagres,
CoussapoamagnifoliaTrecul,Ann. Sci. Nat. Bot. Ser. 22 Jun 1860 (9 + d), Hayes 354 (NY, syntypesof C.
3, 8: 98. 1847; Miquel, Fl. bras. 4(1): 139. 1853; chagresiana);nr.FortSherman,16Nov 1955(2), John-
Macbride, Publ.FieldMus.Bot.13(2.2):297. 1937; ston 1709 (A);E of Indio Tower,2 Apr 1956 (9),John-
Woodson& Schery,Ann.MissouriBot. Gard.47: ston 1768 (A, MO); Barro Colorado Island, 16 Feb
169. 1960. Type. Peru.Withoutlocality, - (2), Ruiz 1932 (a), Woodworth& Vestal606 (A, F, MO).
& Pavon s.n. (or 4) (holotype, FI; isotypes, B, BM, COLOMBIA. ANTIOQUIA: Rio Porce, 1000 m, Dec
F, US). 1962 (2), Espinal 882 (COL). BOYACA:El Humbo re-
CoussapoaruiziiKlotzsch,Linnaea20:529. 1847.Type. gion, 130 km N ofBogoti, 1200-1300m, 19 May 1933
Peru.Withoutlocality, - (9),Ruiz & Pavons.n. (or (8),Lawrance810 (A, F, G, K, MO, US, typecollection
4) (holotype,B; isotypes, BM, F, FI, US). of C. hypochlora).CAQUETA: La Tagua, 5 May 1953
CoussapoahypochloraStandley,Publ. Field Mus. Bot. (9),Romero-Castaneda4211 (COL,MO). CHOCO: Nr.
17: 162. 1937. Type. Colombia.Boyaca:El Humbo Helipad,Rio Truando,atjunctionwithQuebradaBuche,
region, 130 km N of Bogota, ca. 1150 m, 19 May 1 Apr 1968 (a), Duke 15742 (U); El Valle, beach, 5
1933 (6), Lawrance810 (holotype,F; isotypes,A, G, Aug 1976 (9), Gentryet al. 17217 (MO, NY, U); Rio
K, MO, US). El Valle,betweenEl Valle and villagenr. mouthof Rio
Coussapoacardonae Pittier, Bol. Soc. Venez. Cienc. Mutatt, 10 Aug 1976 (8), Gentryet al. 17493 (U); 5
Nat. 8: 257. 1943, as cardonaei.Type. Venezuela. km E of Istmo de San Pablo, ca. 12 km W of Las
Bolivar:Rio Caura,sabanasde Canaracuni,Jan 1942 Animas, 12 Jan 1979 (2), Gentryet al. 24074 (MO,U).
(9), Cardona374 (holotype, VEN; isotypes, GH, S, CUNDINAMARCA:La Palma, road to Pacho, Rio Murca,
US). 1150-1400 m, 27 Jul 1947 (2), Garcia-Barriga12408
CoussapoachagresianaHawkes,Phytologia3: 30. 1948. (US). SANTANDER: Rio Opon, Atlantico railway, 5 Jul
Syntypes.Panama.CanalZone,Chagres,22 Jun 1860 1958 (6), Garcia-Barriga16239 (NY). VALLE:Nr.
(6 and 9), Sutton Hayes 354 (NY). Buenaventura,15 Nov 1979 (8), VanRoodenet al. 258
(U). VAUPES: Rio Vaup6s,YurupariFalls, 12 Apr 1953
Lamina mostly ovate to cordiform or (8), Schulteset al. 18996 (F), Nov 1951 (2),Schulteset
elliptic al. 19758
to sometimes 17- (GH, MO, US).
suborbicular, obovate, mostly VENEZUELA. AMAZONAS:Between Yavita and
25 cm long, base often subcordateto cordate; Maroa, 6-19 Jul 1969 (st), Bunting et al. 3959 (MY,
upper surfacesmooth and glabrous,sometimes U); upperRio Orinoco,Raudalde los Guaharibos,24
scabridulous,lower surfaceon the smallerveins Jul 1951 (2), Croizat343 (F, US); Yavita, 10 Jul 1972
(9), Lizot (1972-) 1 (US, VEN); San Carlos de Rio
glabrous or sparsely puberulous, on the main Negro, 7 Mar 1942 (9), LI. Williams14675 (A, F, G,
veins sparsely to densely minutely puberulous K, NY, RB, US, VEN). BOLIVAR: Rio Caura,sabanas
with straighthairs, sometimes on the main veins de Canaracuni,Jan 1942 (9), Cardona374 (A, S, US,
partly hirtellous;lateral veins 4-9 pairs, all or VEN, type collection of C. cardonae);Rio Cuyuni,El
most of them branched,sometimesthe basalpair Chibau,2 km S of Rio Chibau,1 Sep 1962 (9),Maguire
et al. 53517 (F, NY, OXF, US, VEN);Rio Cuyuni,NE
not or faintly branched but some of the other of Luepa, 1300 m, 23 Apr 1960 (9), Steyermark&
lateralveins poorlybranched,basalpairreaching Nilsson 507 (NY, US, VEN);Ptari-tepui,1600 m, 11
the marginat, below, or above the middle of the Nov 1964 (st), Steyermark60002 (F, VEN);between
lamina;intercostalvenation plane to prominent; Santa Teresita de Kavanayen and Rio Pacairao, Que-
bradaOparu-ma,1065-1220 m, 20 Nov 1944(9),Stey-
petiole 4-17 cm long; stipules 0.5-3.5 cm long. ermark60432 (F, VEN);between Rio Paramichiand
Staminate inflorescenceswith the heads up to 6 Salto de Chalimano,NW of SerraniaPia-soi, 5 Jan
mm diam.; common peduncle up to 8 cm long. 1962 (9), Steyermark90583 (US, VEN); Rio Parami-
Pistillate inflorescencesmostly with one head, chi, between mouth of Rio Paramichiand Salto de
more than 1 cm, in fruitup to 3 cm diam.;(com- Chalimano,8-9 Jan 1962 (9), Steyermark90736 (K,
NY, US); Rio Cuyuni, 140-142 km S of El Dorado,
mon) peduncle 1.5-9.5 cm long. 1300-1380 m, 22-28 Dec 1970 (9), Steyermarket al.
Distribution(Fig. 6). Amazon Basin (Bolivia, 104348 (MY, NY). DELTAAMACURO:
Cariapo, 13 Dec
Brazil, Peru, Ecuador,Colombia, and Venezue- 1953 (9),Littleet al. 15990 (VEN);Dto. Antonio Diaz,
la), westernGuianaregion(Guyana,Venezuela), CanfoGuiniquina,5 Feb 1977 (6),Marcano-Bertiet al.
84-22-77 (U); Rio Cuyubini,above CasaCuyubini,12
and in the centraland Pacificcoastal part of Co- Nov 1960 (9),Steyermark87480 (G, GH, NY, U, US,
lombia, as far north as Panama;in upland and VEN); Dto. Antonio Diaz, Caio Htioba, tributaryof
riverineforests up to 2000 m. Rio Araquoa,16Oct 1977(st),Steyermarketal. 114759
 Coussapoa 37

FIG. 14. Coussapoaasperifoliassp. magnifolia: 1, leafy twig with pistillate inflorescences(Harling et al.
14733); 2, staminateinflorescence(Garcia-Barriga16239).
38 Flora Neotropica

(MO);Dto. Antonio Diaz, upperpartof CafnoJobure, 4c. Coussapoa asperifolia ssp. rhamnoides
7 Apr 1955 (2), Wurdack286 (F, NY). (Standley)Akkermans& C. C. Berg,Proc.Kon.
GUYANA. Upper RupununiRiver, nr. Dadanawa,
24-29 Jul 1922 (9),De la Cruz1729 (F, GH, MO, NY); Ned. Akad. Wetensch. Ser. C, 85(4): 445.
Mazarunistation,7 May 1943 (2),Fanshawe1278 (FD 1982. Fig. 15.
4014) (K, NY); EssequiboRiver, Tiger Creek, 3 Jun
1943 (6), Fanshawe 1337 (FD 4073) (K, NY); Esse- CoussapoarhamnoidesStandley,Publ.Field Mus.Bot.
17: 166. 1937.Type.Brazil.Amazonas:Mun.Sao
quibo River, Bartica,8 Jul 1952 (6), Fanshawe 3409 Paulode Olivenca, 11 Sep-26 Oct 1936 (9),Krukoff
(FD 6973) (K, NY); Pomeroon River, Kabakaburi,19
8406(holotype,NY;isotypes,A, BM,F, G, K, LE,
Aug 1918 (2),Hohenkerks.n. (FD 738) (K, OXF);Ber-
bice River, Apr 1889 (2), Jenman 4947A (K); Deme- MO,S, U).
rara River, - (8), Parkers.n. (K); Ekaria,Rosedale,
Aug 1924 (2), Persaud116 (B, F, K, NY). Lamina obovate to subobovate or elliptic to
ECUADOR.NAPO:3 mi SW of Tena, 28 Aug 1960 oblong, 7-16 cm long, base obtuse to rounded
(2), Grubb et al. 1498 (K, NY). PASTAZA:Nr. Mera, (to subcordate);upper surface smooth and gla-
ca. 1000 m, 10 Dec 1955 (9), Asplund18780 (S); be- brous;lower surfaceon the main veins sparsely
tween Paloraand Shell, 4 Mar 1980 (st), Berg & Ak-
kermans 1119 (U), (9), Berg & Akkermans1133 (U); minutely puberulouswith straighthairs; lateral
nr. Mera, 17 Jan 1977 (9), Harling et al. 14733 (GB), veins 4-6 pairs, unbranchedor some of the lat-
18 Mar 1940 (8), Lugo 83 (S). eral veins (poorly)branched,the basal pair un-
PERU. Without locality (2), Ruiz & Pavon s.n. (or branchedor faintlybranched,reachingthe mar-
4) (B, BM, F, FI, US, type collectionof C. magnifolia gin above the middle of the lamina;petiole 2-7
and C. ruizii). JUNiN: La Merced, 10-24 Aug 1923 (2),
Macbride5447 (F, C). LORETO: Prov. Maynas, Pefia cm long; stipules 0.5-2 cm long. Pistillate inflo-
Negra, 25 km SW of Iquitos, 1 Aug 1972 (st), Croat rescenceswith (1-)3-5 (sometimes partlyfused)
18643 (MO,NA); Prov. Requena,Rio Ucayali,Jenaro heads, 3-6 mm, in fruit up to 15 mm diam.;
Herrera,below Requena,9 Dec 1977 (2), Gentryet al. (common) peduncle 1-4 cm long.
21319 (MO, U); Prov.
Maynas, nr.
Quistococha, 26
Distribution (Fig. 6). Amazon Basin (Brazil,
May 1978 (6), Gentryet al. 22280 (MO, U); Prov.
Maynas, Rio Nanay, below Bellavista, 31 May 1972 Peru);in upland and riverside(varzea)forest.
(6), McDaniel 16092 (NA); Prov. Maynas,Dist. Iqui-
tos, Puerto Almendra, Rio Nanay, 1 Nov 1976 (6), Specimens studied. PERU. LORETO:Rio Tacsha
Revilla 1260 (MO, U). SAN MARTIN:Prov. Marichal Curaray,18 Sep 1972 (2), Croat 20370 (F, GH, NA,
Caceres,Dist. TocacheNuevo, FundoGranChaparral, NY, P, RB).
11 Nov. 1975 (6), Schunke 8666 (U); Prov. Mo- BRAZIL.AMAZONAS: Rio Solimoes, Rio Bia, afflu-
yobamba, 1906 (st), Weberbauer4472 (G). ent of Rio Jutai, 5 Nov 1975 (Q),L. Coelhoet al. 332
BRAZIL. ACRE:Cruzeirodo Sul, 2 Mar 1976 (2), (U); Barcelos,7 Sep 1962 (Q),Duarteet al. 6970 (RB);
Marinho 348 (IAN). AMAZONAS:
Rio Negro, Sao Ga- Mun.SaoPaulode Olivenca,nr.Palmares,11Sep-26
brielde Cachoeira,28 Feb 1975(9),Cordeiro367 (IAN); Oct 1936 (2), Krukoff8406(A, BM, F, G, K, LE, MO,
mouth of Rio Caiari, affluentof Rio Negro, 15 Sep NY, S, U, type collectionof C. rhamnoides);Manaus-
1952 (?), Fro6s et al. 28585 (IAN); mun. Humaiti, Itacoatiararoad,Res. Flor.Ducke,1 Sep 1966 (2),
nr. Livramento on Rio Livramento, 12 Oct-6 Nov Pranceet al. 2153 (INPA, NY, U). PARA:Rio Tapajos,
1934 (2), Krukoff6662 (A, B, BM, F, G, K, LE, MO, Periquito,27 Jul 1923 (2), Ducke(HJBR) 18460 (RB).
NY, RB, S, U, US); nr. Manaus,1906 (2) Lambroys.n.
(P); Rio Negro, Uanari, nr. Sao Gabriel,31 Oct 1947
(6), Pires 805 (IAN, NY), 2 Nov 1947 (9), Pires 830 5. Coussapoa batavorumAkkermans & C. C.
(NY);Rio Negro,betweenSao Gabrieland Sao Felipe, Berg, Proc. Kon. Ned. Akad. Wetensch. Ser.
base of SerraUanari, Oct 1947 (6), Schultes & Pires C, 85(4): 447. 1982. Type. Colombia. Valle:
8976 (GH, U); Rio Negro, Sao Gabrielde Cachoeira,
Nr. Buenaventura, 5 Dec 1979 (9), Van Rood-
Jan-Aug 1852 (6), Spruce 2260 (BR, K, P). MATO
GRosso:Rio Aripuafia,nr. HumboldtCenter,59?21'N, en et al. 700 (holotype, COL;isotypes, AAU,
10?12'S,19 Oct 1973 (st), Berg s.n. (U); Rio Juruena, K, NY, U, US). Fig. 16.
airport, 17 Jul 1977 (9), M. G. Silva et al. 3368 (U).
PARA:Almeirim,Barreirada Velha Pobre, 6 Jul 1919 Tree,terrestrial,ca. 15 m tall. Leafytwigs 10-
(2), Ducke(HBJR)13605 (RB);Rio Brancode Obidos, 19 mm thick, glabrous. Lamina coriaceous,
21 Jul 1918 (6), Ducke (HAMP) 17132 (MG).
RONDONIA: Nr. Porto Velho, 28 Jun 1952 (9), J. F. broadlyovate to cordiform,36-40 x 34-37 cm,
Silva 227 (IAN);Rio Madeira,CachoeiraMisericorde, apex obtuse to emarginate,base cordate,margin
Riberao, 2 Aug 1968 (9), Pranceet al. 6718 (NY, U). subcrenate;both surfacesglabrous;lateralveins
RORAIMA: West facingslopesof SerraTepequem,1200 10-13
pairs, 3-5 of which arise at the base of the
m, 20 Feb 1967 (2), Pranceet al. 4588 (NY, U).
BOLIVIA. LA PAZ:Prov. S. Yungas, basin of Rio midrib, straight, main basal pair branched,
Bopi, San Bartolome,nr. Calisaya, 1-22 Jul 1939 (2), reachingthe margin at or above the middle of
Krukoff10123 (A, F, G, K, MO, NY, S, U). the lamina;intercostalvenation almost plane to
Coussapoa
FIG. 15. Coussapoaasperifoliassp. rhamnoides:1, leafy twig with pistillateinflorescences(Krukoff8406).
slightly prominent;petiole 10-12 cm long, dis-
tinctly ribbedand densely minutely puberulous;
stipules 8-10 cm long, densely to sparselyred-
dish-brownpuberulousand with longer straight
white hairs. Staminate inflorescencesunknown.
Pistillate inflorescencesunbranched or poorly
branched;heads 1-2(-3), free or partly fused,
globose to ovoid, ca. 8-10 mm, in fruit up to 27
mm diam.; common peduncle 0.5-2 cm long,
ratherdenselypuberulousto hirtellous;perianth
39
1 cm
1(-3) mm high, white puberulous;fruitingperi-
anth brown. Interfloralbractsabsent.
Distribution (Fig. 6). Colombia (Choco); in
riversideforest. Known only from the type col-
lection.
Knowledge of the staminate inflorescenceis
requiredin orderto determinethe relationships
of this species. In the shape, size, and sparse
indumentumof the lamina, it resembles C. as-
perifolia ssp. magnifolia as well as forms of C.
40 Flora Neotropica

FIG. 16. Coussapoabatavorum:1, leafy twig with pistillateinflorescences(Van Roodenet al. 700).
 Coussapoa 41

vannifoliahaving leaves with a sparseindumen- NY); Cocle del Norte, 23 Aug 1978 (2),Hammel 4475
tum. (U); ca. 11 km SW of CerroBraja,9?25'N, 79?35'W,
2 May 1981(8),Sytsmaet al. 4225 (U). PANAMA: Cerro
Jefe, 3 Mar 1979 (8), d'Arcy12329 (U); road to Cerro
6. CoussapoabrevipesPittier, Contr.U.S. Natl. Campana,3 km from InteramericanHighway, 18 Sep
Herb.18:225. 1917;Woodson & Schery,Ann. 1968 (6), Correaet al. 1031A (DUKE, F, MO); Cerro
MissouriBot. Gard.47:171. 1960. Type. Pan- Campana,9 Apr 1970 (9), Croat 14212 (GH, MO);
amf. San Bias: Nr. Puerto Obaldia, hills of CerroJeferegion,La Eneida,25 Mar 1968 (2),Dressier
3454 (DUKE, MO); 10 km N of Margarita,on road
Sperdi, Sep 1911 (v), Pittier 4386 (holotype, to Madronio,31 Jan 1979 (8),Hammel 6006 (U); 3 mi
US; isotype, B). Fig. 17. NE of Altos de Pacora,CampoTres, 10 Mar 1973 (6),
Liesner515 (K, MO, NY, US); Altos de Campana,28
Tree,terrestrialor hemi-epiphytic,up to 15 m May 1977 (8), Mendez 15 (MO); 6.5 km N of Lago
tall. Leafy twigs 4-11 mm thick, more or less Azul, 13 Jan 1974 (9), Nee 9281 (MO, NY, U); El
densely white (appressed-)puberulousand with Llano-Cartiroad, 8 mi from Pan AmericanHwy., 17
reddish-brownpluricellularhairs, or also with Apr 1981 (2), Sytsma 4024 (U); CerroCampana, 18
Sep 1968 (a), Weaveret al. 1677 (DUKE). SANBLAS:
sparsearachnoidhairs.Laminacoriaceous,ovate Road El Llano-Carti-Tupile,1 mile from continental
to elliptic or occasionallyobovate, 5-30 x 2.5- divide, 30 Mar 1973 (9), Liesner 1309 (DUKE, GH,
18 cm, apex obtuse to acute or to subcordate, MO, NY, US); nr. PuertoObaldia,hills of Sperdi,Sep
base obtuseto roundedto subcordate,sometimes 1911(9),Pittier4386 (B, US, typecollection);Rio Acla,
7 Feb 1979 (2), Sugden 415
acute, margin entire or subcrenatetowards the 8048'38"N,77040'30"W,
(K).
apex; upper surfaceglabrous,lower surfaceap-
pressed-puberulous,densely so in the areoles, Coussapoa brevipes shows morphological af-
and on the main veins often with white arach- finities especially with C. chocoensis but also with
noid hairswhicharesoon deciduous;lateralveins C. contorta.
6-18 pairs, straightor slightlycurved,basal pair
branched,reachingthe marginbelow the middle 7. Coussapoachocoensis Cuatrecasas,Caldasia
of the lamina;intercostalvenation almost plane; 7:287.1956. Type. Colombia.Choc6:Rio San
petiole 1-7 cm long, 2-4 mm thick, appressed- Juan,Quebradadel Taparal,30 May 1946 (2),
puberulousand often also with white arachnoid Cuatrecasas 21446 (holotype, F). Fig. 18.
hairs which are infrequentlydeciduous;stipules
3-10 cm long, densely covered with appressed Tree, hemi-epiphytic. Leafy twigs 3-6 mm
white puberulous hairs, mixed with reddish- thick,brownishsubhirsuteto hirtellous.Lamina
brown pluricellularhairs and often also with coriaceous,ovate to elliptic, 8-16 x 4-12, apex
sparseto densewhitearachnoidhairsand/orlong shortly acuminate,base roundedto subcordate,
margin entire; upper surface glabrous, lower sur-
yellowishstraighthairs.Staminateinflorescences face densely appressed-puberulous in the areoles,
branched;heads4-20, globose,ca. 4-5 mm diam.;
common peduncle 0.5-3 cm long, densely cov- sparsely puberulous to hirtellous on the smaller
ered with short white puberulousand reddish- veins, densely subhirsute, subvelutinous, hirtel-
brownpluricellularhairs;perianthca. 1 mm high, lous, strigose or strigillose on the main veins;
lateral veins 10-12 pairs, curved and distinctly
densely minutelypuberulous;stamens three, far
looping, basal pair unbranched or sparsely
exceeding the perianth. Pistillate inflorescences
branched,reachingthe marginbelow the middle
unbranched;heads globose to ellipsoid, ca. 5-7 of the lamina;intercostalvenation ratherprom-
mm, in fruit up to 14 mm diam.; peduncle0.4-
inent; petiole 1-3 cm long, brown subhirsuteto
1.5(-2.5) cm long, denselywhite puberulousand
also with reddish-brownpluricellularhairs;peri- subvelutinous;stipules 1.5-2.5 cm long, brown
anth ca. 1-2 mm high, densely minutely pub- (sub)hirsute. Staminate inflorescences unknown.
Pistillate inflorescences unbranched; head (glo-
erulous;fruitingperianthorange.Interfloralbracts bose or ellipsoid?),ca. 15 mm diam.; peduncle
absent.
0.5-1 cm long, hirtellousto subhirsute;perianth
Distribution(Fig. 6). Panama,in forestsup to
ca. 2 mm high, minutely brown puberulous.In-
800 m.
terfloral bracts subspathulate, hirtellous at the
Specimens studied.PANAMA: COCLE:Nr.LaMesa, apex.
4 kmfromElValle,15Sep1968(Y),Correaet al. 1002
(DUKE,F);foothillsof CerroPil6n,nr.ElValle,5 Oct Distribution(Fig. 6). Colombia (Choc6);low-
1967 (Q),Dukeet al. 14712 (MO,US). COLON: Nr. land, riverside. Known only from the type col-
Guasimo, 22 Apr 1970 (Q),Croat9973A (F, GH, MO, lection.
 A'
11 I

c1m

.. .
FIG. 17. Coussapoabrevipes:1, leafy twig with staminateinflorescences(Weaver& Foster 1677); 2, pistillatein
'?

'Cous':
'.' ... . . c e :1..

~~~~~~~~~~~~~~~~~~~~~~~~~~~?~
\~~~~~~~~~~~~~~~~~~~~~? '' ;:
?~. .: ..8~~~~~~~~~~~~~~~
:7
?:::
.:; 0 .-; - :. .

..-.
i'1'::"
,~~ ~ j. 'i ' :
." :,,; ' .?

?'"'",...?:

. ;... ?.
iI.' ?

?.:.;?...;
':I~~~ ~ cm

~~~~'"' ?~~~~~
"~~~~~~~~~~~~~~~~~~~' i,:~..~~~~~~~~~~~~~~~~~~
....:?.;t~' . .
(.:??

~~~~~. ~~~~~~~I
' ~~~~~~~~~~~~~~~~~
'-~~~~ . '? "'

::i.: ' ??
~~~~~~~~~~~~~
?'"....:',,: .

FIG. 18. Coussapoa ':."

~~?
''(...'".1
chocoensis: ~~~~~~~
'"!;:"i,~'i .,. 1,
~ ::'"..?..
leafy twig with pistillate inflorescences (Cuatrecasas 214

~
.. \ ..?. . .

FIG. 18.Coussapoa chocoensis: , leafy twig withpistillate

iniorescences
(Cuatrecasas
44 Flora Neotropica

This speciesresemblesC. brevipes,fromwhich x 1-5 cm, apex (sub)acuteto obtuse,base obtuse

it clearly differsin the brown indumentum and to acute, margin entire;upper surfaceglabrous,
the presenceof interfloralbracts. lower surfaceglabrousor appressed-puberulous
in the lowerpart;lateralveins 3-6 pairs,the basal
8. CoussapoacinnamomeaCuatrecasas,Calda- pairs strong, unbranched,reaching the margin
sia 7: 288. 1956. Type. Colombia. Amazonas: far below the middle of the lamina, the other
TrapecioAmaz6nico, Rio Loretoyacu,28 Sep lateralveins weakand inconspicuous;intercostal
1946 (9), Schultes & Black 8269 (=Black & venation plane; petiole 1-2 (or -3) cm long,
Schultes 46-137) (holotype, F; isotypes, A, denselybrownishappressed-puberulous; stipules
IAN, K, US). Fig. 19. 0.5-2.5 cm long, brownishsubsericeousto sub-
velutinous. Staminate inflorescences poorly
Tree,hemi-epiphyticor terrestrial,up to 8 m heads 2-7, partly fused, globose, ca.
tall. Leafy twigs 9-22 mm thick, brownish branched;
2-3 mm diam.;commonpeduncle1.5-3 cm long,
(sub)velutinous, glabrescent. Lamina coria- brownish
puberulous,perianthca. 1 mm high,
ceous, (broadly)ovate, 16-38 x 12-28 cm, apex
obtuse to (acute),base roundedto truncateor to sparsely minutely puberulous;stamensthree,just
exceeding the perianth. Pistillate inflorescences
cordate, margin subcrenate;upper surface gla- unbranched or
poorly branched; heads 1-2,
brous, lower surface densely puberulousin the sometimes
partlyfused, globose, ca. 4-7 mm, in
areoles, (rather)sparsely appressed puberulous fruit to 9 mm
to substrigoseon the veins; lateral veins 14-22 up diam.; (common) peduncle2-
5 cm long, brownishpuberulous;perianthca. 1
pairs, (almost) straight, basal pair branched, mm
high, glabrous;fruitingperianthyellow. In-
reachingthe marginbelow the middle of the lam- bracts spathulateto subpeltate or pel-
terfloral
ina; intercostalvenation (almost) plane; petiole tate.
5-15 cm long, 5-7 mm thick; shortly subvelu-
Distribution (Fig. 6). Known in Colombia
tinous, also brownpluricellularhairs.Staminate Ecuador(Pastaza),and Bolivia (La
inflorescencesunknown.Pistillate inflorescences (Antioquia),
Paz); in forest at altitudes between (400-?) 800
branched;heads 2-4, often partly fused, sub- and 1200 m.
globose, in fruitup to 40 mm in diam.;common
peduncle 2-4 cm long, peduncle and branches Specimens studied. COLOMBIA. ANTIOQUIA: Mun.
6-10(-15) mm thick, shortly velutinous; peri- San Luis, rd. Medellin-Bogota,quebradaLa Tebaida,
anth glabrous.Interfloralbractsabsent. 1000-1100 m, 22 Jun 1987 (2), Callejas et al. 3997
Distribution (Fig. 6). Colombia (Amazonas, (BG);nr. SanLuis, 900-1000 m, 21 Jun 1982 (6),Her-
nandezet al. 369 (HUA).
TrapecioAmaz6nico);riversideforest. ECUADOR. PASTAZA: Nr. Mera, ca. 1000 m, 14
Specimensstudied.COLOMBIA.AMAZONAS: Tra- Feb 1955 (9), Asplund 18840 (S), 28 Oct 1938 (6),
pecioAmaz6nico,Rio Loretoyacu, 22 Nov 1945(2), Schultze-Rhonhof2943(B, type collection).
Duque-Jaramillo2179 (COL),Oct 1945 (2), Schultes BOLIVIA.LAPAZ:Prov. Larejaca,Copacabana,10
6693 (F, GH, U, US), 28 Sep 1946 (2),Schultes&Black km S of Mapiri,850-950m, 8 Oct-15Nov 1939(6),
8269(=Black&Schultes46-137)(F,GH,IAN,K,US, Krukoff11275 (A, F, G, K, MO, NY, S, U, US), (9)
type collection). Krukoff11276(A, F, G, MO,S, U, US).
C. cinnamomeahas many featuresin common In many characters,and especiallyin the sub-
with C. tessmannii, but in most parts is larger trinervatelamina, C. cinnamomifoliais similar
and more robustthan the latterspecies. The two to C. trinervia.The 3-staminate flowers, how-
species appearto be closely related. ever, suggestrelationshipswith other species, in
particularwith C. parviceps.
9. Coussapoa cinnamomifoliaMildbraed, No- The Colombian collection differs somewhat
tizbl. Bot. Gart. Berlin 15: 783. 1942. Type. from the other collections in the longer petiole
Ecuador.Pastaza:Nr. Mera, ca. 1000 m, 28 (up to 3 cm) and peltate and largerbracts.
Oct 1938 (i), Schultze-Rhonhof2943 (holo-
type, B). Fig. 20. 10.
CoussapoacontortaCuatrecasas,Caldasia7:
Tree,mostly terrestrial,up to 35 m tall. Leafy 289. 1956. Type. Colombia. Valle: Pacific
twigs 2-5 mm thick, densely brownish ap- Coast, Rio Naja, braza Aji, Calle Larga, 28
pressed-puberulous,glabrescent.Lamina coria- Feb 1943 (6), Cuatrecasas 14284 (holotype,
ceous, oblong to subovate or subobovate, 3-13 F). Fig. 21.
Coussapoa 45

1
cm
"~ [

FIG. 19. Coussapoa cinnamomea: 1, leafy twig with pistillate inflorescences (Schultes 6693 (inflorescences)
and Schultes& Black 8269 (leaf and stipules)).
46 Flora Neotropica

1 cm

FIG. 20. Coussapoacinnamomifolia:1, leafytwig with pistillateinflorescences(Krukoff11276);2, staminate

inflorescences(Krukoff11275).

Shrubor tree,hemi-epiphyticor terrestrial,up tic or oblong to ovate or obovate, 5-26 x 1.5-

to 20 m tall. Leafytwigs 3-6 mm thick, yellowish 13 cm, apex acute to acuminate, base acute to
appressed-puberulousor sometimes partly hir- obtuse and occasionally to cordate, margin en-
tellous. Lamina chartaceousto coriaceous,ellip- tire;uppersurfaceglabrous,lower surfaceon the
Coussapoa 47

X { cm

FIG. 21. Coussapoa contorta: 1, leafy twig with pistillate inflorescences (Javita & Epling 1097); 2, leafy twig
with staminate inflorescences (Cuatrecasas 14284).

mainveins rathersparselyappressed-puberulous the smaller veins rathersparselypuberulousto

and sometimes, only in the axils of the lateral glabrous;lateralveins 6-15 pairs,slightlycurved,
veins on more or less distinct domatium-like basal pair unbranched,reachingthe marginbe-
cavities,distinctlylongerstiffyellowishhairs,on low the middle of the lamina; intercostalvena-
48 Flora Neotropica

tion plane; petiole 1-8 cm long, appressed-pu- small, ca. I cm long stipules, the basal pair of
berulous; stipules 2-4 cm long, yellowish lateralveins reachingthe margin of the lamina
appressed-puberulousto (sub)sericeous.Stami- at aboutthe middle, and the ratherdensebrown-
nate inflorescence poorly branched; heads 6-10, ish hairs on the perianthof the pistillate flower.
globose, ca. 2-3 mm diam.; common peduncle It couldrepresenta distinctspeciesor subspecies.
1-3 cm long, densely puberulousto hirtellous;
perianth ca. 1 mm high, minutely puberulous;
stamens three, far exceeding the perianth. Pis- 11. CoussapoacrassivenosaMildbraed,Notizbl.
tillate inflorescences mostly unbranched, occa- Bot. Gart. Berlin 15: 784. 1942. Type. Ecua-
sionally branched;heads 1-2, globose, 5-7 mm, dor. Pastaza:Nr. Mera,ca. 1000 m, 7 Sep 1938
in fruitup to 10 mm diam.; (common)peduncle (a), Schultze-Rhonhof2783 (holotype, B).
1-2 cm long, densely puberulousto hirtellous; Fig. 22.
perianth1-2 mm high, minutelypuberulous.In-
terfloralbracts peltate, relatively large, densely CoussapoasteyermarkiiCuatrecasas,Fieldiana
Bot. 28(1): 212. 1951. Type. Venezuela. Bo-
puberulous.
Distribution(Fig. 6). Colombia and Ecuador, livar: Ptari-tepui, 1600 m, 10 Nov 1944 (Y),
Pacific coastal region, also in Costa Rica?; in Steyermark 60002a (holotype, F; isotype,
forest up to 1500 m altitude. VEN).

Specimensstudied. COLOMBIA. CAUCA?: Juntas,

Tree, often hemi-epiphytic, up to 20 m tall.
- (2), LehmannBT694 (A, GH, K, L, NY). CHocO: Leafytwigs 6-9 mm thick, densely (pale)brown
Rio San Juan,Quebradade Taparal,30 May 1946 (6), puberulousto hirtellousor to subhirsuteto stri-
Cuatrecasas21496 (F); Rio San Juan, between Que- gose. Lamina coriaceous, oblong to elliptic to
brada La Sierpe and QuebradaEl Quicharo,27 Mar suborbicularto subobovateor to
1979 (6), Foreroet al. 4099 (COL, MO, U); Rio San (sub)ovate,4-
nr.
Juan, Palestina, 26 Mar 1979 (6),Foreroet al. 4110 24(-43) x 2-14(-20) cm, apex acuminate to ob-
(COL,MO, U); Rio San Juan, Quebradade Taparal, tuse and to rounded,base obtuse to rounded(or
5 Dec 1979 (9), Van Rooden et al. 699 (U). NARIRO: to acute), marginentire;uppersurfaceglabrous,
Mun. Iscuand6,QuebradaLa Berrera,28 Nov 1955 lowersurface
strigoseto subsericeouson the main
(6), Romero-Castaieda 5512 (COL). VALLE:Pacific
Coast, Rio Naja, Calle Larga,28 Feb 1943 (6), Cua- veins, sparsely(along marginof the lamina) pu-
trecasas14284 (F, type collection);PacificCoast, Rio berulous to hirtellous, in addition covered
Cajambre,5-15 May 1944 (9), Cuatrecasas17647 (F). with (rather) dense, sometimes very sparse,
ECUADOR. BOLIVAR: Valley of Rio Tablas, 1300 (sub)persistent(pale)brownarachnoidhairs;lat-
m, 1 Oct 1943 (st),AcostaSolis 6008 (F). CARCHI: Rio eral veins
Blanco,tributaryof Rio SanJuan,above Chical, 1300- (4-)5-8 pairs, slightly curved, basal
1500 m, 25 Sep 1979 (Q),Gentryet al. 26525 (SEL, pair branched (or sometimes unbranched),
U). ESMERALDAS: Rio Onzole, EsteroChontaduro,14 reachingthe margin at or above the middle of
Jul 1966 (Q),Javita et al. 1097 (MO, NY, S, US). IM- the lamina; intercostal venation (very) promi-
BABURA: Collapi,4 Jul 1949 (2), Acosta Solis 12797
nent; petiole 1-4 cm long, (puberulousto) sub-
(F). ELORO:Trail to Sambotambo,Rio Moro Moro,
S of Buenaventura,highwayto Portovele, 1035-1800 strigoseto hirsute;stipules 1-5 cm long, brown
m, 29 Aug 1943 (2), Steyermark54214 (F). PICHINCHA:subsericeousto subhirsute.Staminate inflores-
Nr. SantoDomingode los Colorados,30 Aug 1930 (6), cences branched;heads 4-20, globose, ca. 2-3
Benoist 3010 (P); road Santo Domingo de los Colo- mm diam.; common peduncle 1-4.5 cm
long,
rados-Quininde(km 170-175), 2 Sep 1949 (st),Acosta
Solis 13687, 4 Sep 1949 (8), AcostaSolis 13713 (F). (rather)denselypuberulousto hirtellousto strig-
illose; perianthca. 1 mm high, minutely brown
The taxonomicrelationshipsof C. contortaare puberulous;stamenstwo, farexceedingthe peri-
not clear.Overallsimilaritiessuggestaffinitywith anth.Pistillateinflorescencesbranched;heads 3-
C. brevipes, but a connection with C. oligoceph- 7, globose, 2-5 mm diam.;common peduncle 1-
ala is also possible. The species is distinguished 3 cm long, brownishpuberulousto hirtellousto
by the domatium-likecavities in the axils of the strigillose;perianthca. 1-1.5 mm high,glabrous.
lateralveins and by the largeinterfloralbracts. Interfloralbractsspathulate,often the lowerpart
Collection Sanchez & Zamora 512, from Cos- very narrow,minutely puberulousat the apex.
ta Rica, San Jose, below La Hondrua, Parque Distribution(Fig. 6). Presently known from
Nacional BraulioCarrillo,2 Jan 1984 (2),(AAU) the followingdisjunctareas:the easternGuiana
differs from the collections cited above in the region (Venezuela and northern Brazil), Ama-
Coussapoa
I.
FIG. 22. Coussapoacrassivenosa:1, leafy twig with pistillate inflorescences(Steyermarket al. 104296); 2,
leafy twig with staminateinflorescences(Krukoff11085).
zonian Bolivia, Amazonian Ecuador, and Pan-
ama; in forests, mostly between 700 and
1700 m.
Specimens studied. PANAMA. PANAMA: Road El
Llano-Carti,9'20'N, 78?50'W,7 Jan 1981 (d), Hahn
347 (U).
VENEZUELA. AMAZONAS:Brazilian border, ca.
2?27'24'N, 63056'W, 22 May 1972 (a), Steyermark
106118 (U, VEN). BoLnvAR: Meseta del Jaua, Cerro
Sarisarinama,700 m, 10 Feb 1976 (a), Brewer-Carias
s.n. (COL,MY);Ptari-tepui,1600 m, 10 Nov 1944 (2),
Steyermark60002a (F, VEN, typecollectionofC. stey-
ermarkil);QuebradaO-paru-ma,between Santa Ter-
esita de Kavanayenand Rio Pacairao, 1065-1220 m,
20-21 Nov 1944 (st), Steyermark60364 (F, VEN);
Ptari-tepui, 1220 m, 28 Nov 1944 (st), Steyermark
60673 (F); Chimanta-tepui(Torono-tepui), 1700 m,
21 May 1953 (9), Steyermark75522 (F, NY, VEN);
Rio Cuyuni, ca. 142 km S of El Dorado, 1300-1380
m, 22-28 Dec 1970 (2), Steyermarket al 104296 (MY,
49
1 cm
NY); Mesetadel Jaua,CerroSarisarinama,700 m, 12-
15 Feb 1974 (6), Steyermarket al. 108988 (F, K, NY);
6 km N of Mision de Santa Teresita de Kavanayen,
1100 m, 21 Feb 1978 (st), Steyermarket al. 115549
(U, VEN).
ECUADOR. MORONA-SANTIAGO:
Cord. de Cutucui,
5-10 km from Logrono, 1200-1500 m, 7-9 Oct 1975
(2), Little et al. 635 (COL,Q). PASTAZA:Road Puyo-
Tena, ca. 8 km, Tnte. Hugo Ortiz, ca. 950 m, 4 Mar
1980 (8),Berg&Akkermans1120 (U); betweenPalora
and Shell, ca. 900 m, 4 Mar 1980 (6), Berg & Akker-
mans 1129 (st),Berg&Akkermans1131 (U); nr. Mera,
ca. 1000 m, 7 Sep 1938 (8),Schultze-Rhonhof2783(B,
type collection), 20 Nov 1938 (9), Schultze-Rhonhof
3006 (B).
BRAZIL. RoRAIMA: Nr. Auaris, 4?3'N, 64?22'W,
760-800 m, 7 Feb 1969 (6),Pranceet al 9679 (F, GH,
K, NY, P, S, U, US); SerraSurucucu,26 Jan 1975 (e),
Ribeiro 612 (IAN, MG, RB), 23 Jan 1975 (9), Rosa
296 (IAN).
BOLIVIA.LAPAZ:Prov. Larecaja,Copacabana,ca.
10 km S of Mapiri, 850-950 m, 8 Sep-15 Nov 1939
(8), Krukoff11085 (A, F, G, K, MO, NY, S, US).
50
This species has a remarkable,apparentlydis-
continuous, distribution. It appears to be very
closely related to C. ferruginea. The two taxa
could prove to be conspecific,althoughprobably
distinct at the subspecificlevel. Several speci-
mens fromVenezuelaand the RoraimaTerritory
(Brazil)aredistinctin havingrelativelybroad(to
suborbicular)leaves.
12. Coussapoa cupularis Akkermans & C. C.
Berg, Proc. Kon. Ned. Akad. Wetensch. Ser.
C, 85(4): 448. 1982. Type. Brazil. Rond6nia:
P6rto Velho, 5 Jun 1952 (9), J. F. Silva 69
(holotype, IAN). Fig. 23.
Trees,terrestrial.Leafy twigs 3-7 mm thick,
brownish puberulousto hirtellous. Lamina co-
riaceous, elliptic to obovate, 5-10 x 3-7 cm,
apex obtuseto shortlyacuminate,base obtuse to
(sub)cordate,marginentire to subcrenate;upper
surfaceglabrous,lowersurfaceon the midriband
lateralveins sparselyto ratherdenselyappressed-
puberulousto strigillose,for the rest densely(ap-
pressed-)puberulous;lateral veins 5-8 pairs,
slightly curved, basal pair unbranchedor occa-
sionallywith a singlebranch,reachingthe margin
below to just above the middle of the lamina;
intercostalvenation plane; petiole 2-5 cm long,
denselypuberulousto strigillose;stipules2-5 cm
long, yellow to brownishsubsericeousto subhir- 13. CoussapoacurraniiBlake,Contr.U.S. Natl.
sute. Staminate inflorescencesunknown. Pistil-
late inflorescencesbranched;heads 3-4, globose,
ca. 4-6 mm, in fruit up to ca. 10-15 mm diam.;
common peduncle 2-4 cm long, puberulousto Coussapoa
hirtellous,branchesbelow the headsmore or less
strongly broadened (more or less cupuliform); Serrada Estrella,21 May 1877(3), Glaziou8934
perianthca. 1-2 mm high, subpapillatewith mi-
nute thick brown hairs. bractsabsent. US).
Interfloral
Distribution(Fig. 6). Brazil(Rond6nia);in for- 22: 72. 1940. Type. Brazil. Minas Gerais: Mun.
est of terrafirme.
Specimensstudied.BRAZIL.RONDONIA:
Road Por-
km 117,Rio Jamari,15 Aug 1968
to Velho-Cuiaba,
(2), Pranceet al. 6969(U);8 kmfromPortoVelho,5
Jun1952(2), J. F. Silva69 (IAN,typecollection),26
Jun 1952 (2), J. F. Silva 211 (IAN).
Coussapoacupularis,representedby threecol-
lectionsfromthe same area,appearsto be closely
related to C. orthoneura,and may even prove
not to be distinctat the species level. At present,
the speciesdiffersfrom C. orthoneurain the basal
lateralveins which mostly reach the marginbe-
low the middle of the lamina, the broadestpart
Flora Neotropica
of which tends to be below the middle; in the
stronglybroadened,almost cupuliformapices of
the head-bearingbranchesof the pistillateinflo-
rescences;and in the relativelydense brown pa-
pillate apex of the perianthof the pistillateflow-
er. Moreover, the stipules are densely covered
with long yellowish hairs, which is uncommon
in C. orthoneura.Coussapoacupularisalso shows
resemblancesto C. nitida.
Becauseof the occurrenceof more or less con-
spicuouslybroadenedbranchesbelow the pistil-
late flowerheads and other similarities,as in the
indumentum, C. cupularis, C. orthoneura,C.
arachnoidea,and C. nitida appearto constitute
a rather distinct group among the taxa with
ebracteateinflorescencesand unistaminateflow-
ers.
CollectionHuashikat 1585, Peru, Amazonas,
basin of Rio Santiago,65 km N of Pinglo, near
Caterpiza,19 Dec 1979 (9) (MO, U), resembles
C. cupularisin many characters.The main dif-
ferenceis the branchingof the basallateralveins.
Less importantare the greaternumberof lateral
veins (8-9 pairs)and some differencesin the in-
dumentum. In the leaf venation the specimen
approachesC. nitida.The identity of this collec-
tion is uncertain.
Herb. 20: 237. 1919. Type. Brazil.Bahia:Rio
Gongoji, 1 Oct-30 Nov 1915 (9), Curran 8
(holotype, US; isotype, RB). Fig. 24.
warburgianaMildbraed, Notizbl.Bot.Gart.
Berlin10:416. 1928.Type.Brazil.Rio de Janeiro:
(holotype,B;isotypes,BR,G, K, LE,MO,P, S, U,
CoussapoaincomitataStandley,Publ. Field Mus. Bot.
Tombos, Fazenda da Cachoeira, 29 Jul 1935 (9),
Mello Barreto1795 (holotype,F).
Tree, (mostly?) terrestrial,up to 40 m tall.
Leafy twigs 4-6 mm thick, glabrousor sparsely
puberulous, sometimes initially with sparse
arachnoid hairs. Lamina (sub)coriaceous,ob-
ovate or subobovate,4-19 x 1-9 cm, apex ob-
tuse to rounded to emarginateor shortly acu-
minate,baseobtuseto acute,marginentire;upper
surfaceglabrousor with sparse(to ratherdense)
white arachnoid hairs which soon disappear,
lowersurfacesparselypuberulous,oftenalso with
(mostly sparse)deciduous arachnoidhairs; lat-
Coussapoa 51

1 cm

FIG. 23. Coussapoacupularis:1, leafy twig with pistillateinflorescences(J. F. Silva 69).

eral veins 7-10 pairs, basal pair unbranchedor petioles 1-5 cm long, glabrousto sparselypub-
sometimesfaintlybranched,reachingthe margin erulous or also with sparse to dense arachnoid
below the middle of the lamina; intercostalve- hairs; stipules 0.4-0.6 cm long, appressed-pub-
nationslightlyprominent,the smallerveins plane; erulous to strigilloseto subsericeous(to subhir-
52 Flora Neotropica

I
1 cm

FIG. 24. Coussapoacurranii:1, leafy twig with staminateinflorescences(Glaziou8934).

sute). Staminate inflorescencesbranched;heads US, type collection). EsPiarro SANTO:Res. Flor. de

severalto many, globose, ca. 2 mm diam.; com- Linhares,24 May 1978 (d), DAF 004 (GUA). MINAS
GERAIS:Mun. Tombos, Fazendada Cachoeira,29 Jul
mon peduncle 1-5 cm long, sparselyto densely 1935 (9), Mello Barreto 1795 (F). Rio DEJANEIRO:
puberulousor glabrous;perianthca. 1 mm high, Corcovado, 1857 (2), Casaretto617 (G); Givea-Sao
sparselyminutely puberulous;stamens two, ex- Conrado,Jun 1960 (2), Duarte5239 (M, NY, RB, US);
ceeding the perianth. Pistillate inflorescences Serrada Estrella,Petr6polis,21 May 1877 (a), Glaziou
8934 (B, BR, G, K, LE, MO, P, U, US, type collection
branched;heads 3-6, globose,ca. 4-6 mm diam.; of C. warburgiana);
common peduncle 1-4 cm long, patent to ap- Corcovado,29 Jun 1876 (st), Gla-
ziou 8934a (P); Rio de Janeiro,Tijuca, Jan 1837 (2),
pressed puberulous;perianthca. 1-2 mm high, Guillemin1339 (P); Rio de Janeiro,JardimBotanico,
minutely puberulous. Interfloral bracts sub- 3 Sep 1926 (2), Kuhlmanns.n. (U); Cordeira,Fazenda
spathulateto subpeltate,puberulousat the apex. SantaClara,Feb 1970 (6), Lisboa s.n. (R); Rio de Ja-
Distribution(Fig. 7). EasternBrazil,from Rio neiro, Jardim BotAnico,29 Jul 1922 (8), Occhioniet
al. (HJBR) 5864 (GUA, RB, U), 25 Jul 1922 (9), Oc-
de Janeiro to Bahia; in moist forests up to chioni et al. (HJBR) 5917 (RB).
500 m.
studied.BRAZIL.Withoutlocality,2 Dec
Specimens Coussapoa curranii is very closely related to
1968 (8), Gomeset al. 2818 (RB).BAHIA: C. floccosa. Affinities of these two species with
Rio Gongoji,
1 Oct-20Nov 1915(9),Curran8 (RB(HJBR13070), other Coussapoa are not clear.
 Coussapoa 53
14. CoussapoaduqueiStandley,Publ. Field Mus. PASTAZA:Rd. Hollin-Loreto-Coca,between Rio Pu-
Bot. 22: 71. 1940. Type. Colombia. Caldas: cunoandCaceriode Guamani,1200m, 12 Dec 1987
(6), Cer6nM. 2955 (BG);nr. Mera,ca. 1400m, 22
Palestina, 1500 m, 15 Jun 1938 (d), Duque- Nov 1938(9),Schultze-Rhonhof3020 (B,typecollec-
Jaramillo 1767 (holotype, F; isotypes, A, K, tion of C. apoda).PICHINCHA:
Rd. Nanegalito-Pacto,
NY, US). Fig. 25. 5 km W of Tulipe,1300-1500m, 22 Jul 1980 (6),
Holm-Nielsen
et al. 24519(BG).
Coussapoa apodaMildbraed,Notizbl.Bot.Gart.Ber-
lin 15:784. 1942.Type.Ecuador.
Pastaza:Nr.Mera, C. duquei is closely related to C. villosa, es-
ca. 1400 m, 22 Nov 1938 (9),Schultze-Rhonhof3020 pecially to the form with brown arachnoid in-
(holotype,B). dumentum(see p. 104). The speciesis distinctin
the shortly pedunculatepistillate inflorescence
Tree,(secondarily?)terrestrial,up to 30 m tall.
withan ellipsoidto clavatehead.This taxoncould
Leafytwigs 5-10 mm thick, hirtellousto hirsute. be
Lamina coriaceous,elliptic to ovate, 8-18 x 5- regardedas distinct only at the subspecies
11 cm, apexobtuseto acuteto shortlyacuminate, level.
base subacuteto obtuse (or to subcordate),mar- 15.
CoussapoaechinataAkkermans& C. C. Berg,
gin entire to subcrenate;uppersurfaceglabrous, Proc.Kon. Ned. Akad.Wetensch.Ser.C, 85(4):
lower surface puberulous to hirtellous on the
449. 1982. Type. Panama.Panama:CerroJefe,
veins, the whole surface densely covered with Altos de Pacora, Rio Diablo, 15 km NE of
(sub)persistent pale yellow-brown arachnoid Cerro Azul village, 5 Jan 1975 (9), Gentry&
hairs; lateral veins 10-14 pairs, straight, main Mori 13408 (holotype, U; isotype, MO).
basalpair(sometimesfaintly)branched,reaching
the marginbelow the middle of the lamina; in- Fig. 26.
tercostal venation prominent;petiole 2-10 cm Tree, hemi-epiphytic. Leafy twigs 4-6 mm
long, (rather)sparselypuberulous,with whitish thick, glabrous or sparsely appressed-puberu-
to yellowish-brownarachnoid hairs which are lous. Lamina coriaceous, oblong to elliptic to
also sometimes deciduous;stipules2-7 cm long, subobovate,9-20 x 4-10 cm, apex acuteto acu-
white to brown strigoseto hirsuteand with red- minate, base acute, marginentire;uppersurface
dish-brownarachnoidhairs. Staminate inflores- glabrous,lowersurface(purplishor reddish),gla-
cencesbranched;heads many, globose to oblate, brousor on the veins sparselypuberulous;lateral
ca. 4-8 mm diam.; common peduncle 1-2 cm veins 4-5 pairs,slightlycurved,basalpair(faint-
long, puberulousto hirtellous, sometimes also ly) branched,reachingthe marginabove the mid-
with arachnoidhairs; perianth ca. 1 mm high, dle of the lamina; intercostal venation slightly
yellowish-brownpuberulous;stamens two, far prominent to plane; petiole 2-6 cm long, gla-
exceeding the perianth. Pistillate inflorescences brous; stipules 1.5-2.5 cm long, sparsely ap-
unbranched;heads ellipsoid to clavate, ca. 1.5 pressed-puberulous. Staminate inflorescences
x 1 cm; peduncle 0.5-0.8 cm long, puberulous branched;heads numerous, ca. 2 mm diam.;
to hirtellous hairs, sometimes also with arach- common peduncle ca. 1.5-2 cm long, puberu-
noid hairs; perianthca. 1-2 mm high, densely, lous; perianthca. 1 mm high, glabrous;stamens
minutely puberulous. Interfloral bracts (sub) three.Pistillateinflorescencesbranched;heads3-
spathulate,puberulousat the apex. 8, globose, ca. 4-6 mm, in fruit up to 8 mm
Distribution(Fig. 7). Andean region (Colom- diam.; common peduncle 1-3.5 cm long, with
bia and Ecuador);in (sub)montaneforest, be- sparsepatentshorthairs,occasionallymixedwith
tween 1200 and 2000 m. some longerhairs;flowersfree, perianthca. 1-2
mm high, (when dry) with an acute apex, gla-
Specimensstudied.COLOMBIA.ANTIOQUIA: Me- brous;
dellin, ca. 1500 m, - (st), Toro 120c (US). CALDAS: fruitingperianthorange.Interfloralbracts
Cord.Central,Palestina,1500 m, 15 Jun 1938 (8), narrowly spathulate, short, sparsely, minutely
Duque-Jaramillo 1767(A, F, K, NY, US, typecollec- puberulous.
tion). NORTEDESANTANDER: Ocafa, Jun 1845 (9),Pur- Distribution (Fig. 7). Panama (Panama and
die s.n. (K). San Bias);in wet forest, up to 800 m.
ECUADOR.NAPO:SE of Borja,ca. 1900 3
m, Aug
1960 (d), Grubbet al. 1205 (K, NY); NE of Borja,ca. Specimens studied. PANAMA. PANAMA:Road El
1800 m, 15 Aug 1960 (9), Grubbet al. 1283 (K, NY); Llano-Carti,km 17-20, 8 Mar 1974 (9),Dressier4636
nr. Borja, 1650 m, 16 Aug 1975 (9), Little et al. 212 (F); CerroJefe, Altos de Pacora region, Rio Diablo,
(Q), 1850 m, 16 Aug 1975 (8), Little et al. 213 (Q). 15 km E of CerroAzul village, 5 Jan 1975 (2), Gentry
54 Flora Neotropica

2 ~ ? I

wo t :1cm

FIG. 25. Coussapoaduquei: 1, leafy twig with pistillate inflorescences(Grubbet al. 1283); 2, staminate
inflorescences(Grubbet al. 1205).
Coussapoa SS

'1 R~~~~~~~~~~~~~~~~~~~~~~~~?

: e' r ]; e

FIG. 26. Coussapoa

echinata: 1, leafy twig with pistillateinflorescences
4636).i?
?. (Dressler
56 Flora Neotropica

G.27. oussapoaferruginea:1, leafy twigwith staminateinflorescences(Irwin

amet

FIG. 27. Coussapoaferruginea:1, leafy twig with staminateinflorescences(Irwinet al. 48631).

& Mori 13408 (MO, U, type collection). SAN BLAS: dense persistent brown arachnoid hairs; lateral
Nusagandi, rd. to Carti, 18 Jul 1984 (a), McDonagh et
veins 4-5(-6) pairs, straight, basal pair un-
al. 114(MO).
branched,reachingthe marginbelow the middle
The species shows similaritiesto C. parviceps, of the lamina; intercostal venation prominent;
fromwhichit is distinctin the shapeof the leaves, petiole 0.5-1.5 cm long, denselybrownhirtellous
the free pistillate flowers, and the presence of to subhirsute; stipules 1.5-3 cm long, brown
interfloralbracts in the pistillate heads. On the (sub)hirsuteto strigoseor partlywhitish subseri-
other hand, C. echinata shows similaritiesto C. ceous. Staminate inflorescencesbranched;heads
herthae,especiallyin the acute apex of the fruit- several to many, ca. 2-3 mm diam.; common
ing perianth(in dry material). peduncleca. 1 cm long, densely hirtellous;peri-
anthca. 1 mm high,denselypuberulous;stamens
16. CoussapoaferrugineaTrecul,Ann. Sci. Nat. two, far exceeding the perianth.Pistillate inflo-
Bot. Ser. 3, 8: 93. 1847; Miquel, Fl. bras.4(1): rescencesunknown.Interfloralbractsspathulate,
137. 1853;Berg,Fl. Suriname5(1): 286. 1975. puberulous.
Distribution (Fig. 6). French Guiana and the
Type. FrenchGuiana. Without locality, 1838
(6),Leprieurs.n.(holotype,P; isotypes,G, GH, adjacent part of Amapa (Brazil); in riverine for-
est.
L, U). Fig. 27.
Tree, hemi-epiphytic. Leafy twigs 3-7 mm Specimens studied. FRENCH GUIANA. Without
locality, 1838 (d),Leprieurs.n. (F, G, GH, L, P, R, U,
thick, brown(ish) (sub)hirsute. Lamina coria- type collection), 1840 (8), Leprieurs.n. (FI, G, GH, K,
ceous, lanceolateto oblong, 2-8 x 1-4 cm, apex P).
acute to subobtuse, base acute, margin entire; BRAZIL.AMAPA: Rio Oiapoque,betweenmouth of
surface lower surface Camopi River and CachoeiraCamaraua,3 Oct 1960
upper glabrous, appressed-
(d),Irwinet al. 48631 (F, K, MO, NY, U, US).
puberulousto strigose on the main veins, pu-
berulous to hirtellous (to subhirsute) on the The species is similar in many characters to
smaller veins, the whole surface covered with C. crassivenosa. However, it is distinct from the
Coussapoa
FIG. 28. Coussapoafloccosa:1, leafy twig with pistillateinflorescences(Irwin2058).
latter in having a narrowerlamina, a smaller
number of lateral veins, and a relatively short
petiole.Pistillateinflorescencesareneededto de-
cide whetherthis taxon is distinct from C. cras-
sivenosaat the specificor only intraspecificlevel.
17. CoussapoafloccosaAkkermans& C. C. Berg,
Proc.Kon. Ned. Akad.Wetensch.Ser.C, 85(4):
451. 1982. Type. Brazil.Minas Gerais:Vi9osa,
16 Nov 1935 (2), Kuhlmann2074 (holotype,
RB; isotypes, RB, U, US). Fig. 28.
Tree or shrub, (mostly?) hemi-epiphytic.
Leafy twigs 10-15 mm thick, (rather)densely
57
puberulousto (sub)hirsute.Lamina coriaceous,
elliptic to ovate or obovate, 10-25 x 6-17 cm,
apex rounded to obtuse or to emarginate,base
obtuse to roundedto cordate,marginentire;up-
per surface often densely covered with white
arachnoidhairswhich laterdisappear,lower sur-
face hirtellousto subtomentellouson the veins,
the whole surface sparsely to densely covered
with white arachnoidhairs, later deciduous;lat-
eralveins 7-10 pairs,(main)basal pairdistinctly
branched,reachingthe margin at or above the
middle of the lamina; intercostalvenation very
prominent;petiole 1.5-5 cm long, tomentose to
densely hirtellous;stipules 1-2.5 cm long, white
to brownishappressed-puberulous to strig(ill)ose
58 FloraNeotropica
to subsericeous(to subhirsute),sometimes also mm high,glabrous.Interfloralbracts(sub)peltate,
with sparsearachnoidhairs. Staminate inflores- nearlyglabrous.
cencesbranched;heads numerous,globose, ca. 4 Distribution(Fig. 7). CentralAmerica:Costa
mm diam.; common peduncle 1.5-4 cm long, Rica (Puntarenas),Nicaragua(Bluefieldsand Rio
puberulous;perianthca. 1 mm high, puberulous; San Juan), Panama(San Blas);in wet (riverside
stamens two, exceeding the perianth. Pistillate or swamp)forest.
inflorescencesbranched;heads 2-3, partlyfused,
(sub)globose,5-7 mm diam.; common peduncle Specimensstudied.NICARAGUA.BLUEFIELDS:
Rio
1-3 cm long, densely puberulous;perianth 1-2 Kama,betweenKamaand CanioValentin,10 Mar
1966(2), Proctoret al. 27080(F, US, typecollection).
mm high, glabrous. Interfloral bracts sub- Rio SANJUAN:1 km NW of Rio Santa Cruz, 22 Feb
spathulate,puberulousat the apex. 1984 (2), Moreno23629 (BG).
Distribution(Fig. 7). Brazil(MinasGerais,Vi- COSTA RICA. PUNTARENAS: Peninsulade Osa, 8
km S of Rinc6n, 28 Feb 1965 (9), Jimenez 3016 (F).
qosa)in forest. PANAMA. SANBLAS:El Llano-Cartird., km 19.1,
Specimens studied. BRAZIL. MINASGERAIS: Est. 1 Feb 1985(2),Neverset al. 4804(WIS),3 Nov 1985
(st), Neverset al. 6186 (BG). SANBLAS-PANAMA:
Exp.de AguaLimpa,Feb 1968(2), Gomeset al. 2819 Llano-Carti El
(RB);Vi9osa,10 Nov 1958(2), Irwin2058 (F, NY, rd.,28 Aug1982(d),Hamiltonetal. 1062
R, US),31 Aug1934(st),Kuhlmann 1832(US),1935 (BG).
(st),Kuhlmann 2073(RB,US),16Nov 1935(2),Kuhl- This speciesappearsto be one of a groupwhich
mann2074(RB,U, US, typecollection),31 Aug1934
(st), Kuhlmann 2117 (RB), 1935 (9), Kuhlmann2207 also includesC. angustifoliaand C. trinerviaand
(GUA),8 Oct 1976(9),G.Rodrigues et al. 877 (RB); is characterizedby shortlypetiolate,more or less
Capolivinba, nr. Rio Novo, Sep 1895(6),Schwacke distinctlyobovate leaves with few lateralveins,
11895 (B, P). and sparseindumentum.
This species, known only from the surround-
ings of Vicosa, is very closely relatedto C. cur- 19. CoussapoaherthaeMildbraed,Notizbl. Bot.
ranii; but the two taxa are sufficientlydifferent Gart. Berlin 14: 29. 1938. Type. Ecuador.Pi-
for them each to be regardedas species in their chincha: San Carlos de los Colorados, 5 Oct
own right. 1935 (d),Schultze-Rhonhof953 (holotype,B).
Fig. 30.
18. CoussapoaglaberrimaW. Burger,Phytolo-
Tree, mostly hemi-epiphytic,up to 25 m tall.
gia 26: 422. 1973; Burger,Fieldiana Bot. 40:
133, t. 22. 1977. Type. Nicaragua.Bluefields: Leafy twigs 5-13 mm thick densely, yellow to
Rio Kama,betweenKamaand CanioValentin, brown, strigoseto hirsuteto subvillous.Lamina
10 Mar 1966 (2), Proctoret al. 27080 (holo- coriaceous, elliptic to ovate to obovate or to ob-
long, 16-32 x 10-20 cm, apex acuminate to
type, F; isotype, US). Fig. 29.
acute, base obtuse to rounded or occasionally
Tree, hemi-epiphytic or terrestrial, small. subcordate,margin entire or subcrenate;upper
Leafy twigs 2-5 mm thick, glabrous. Lamina surfaceglabrous,sparselyto ratherdensely pub-
subcoriaceous,obovate to narrowelliptic, 2-14 erulous to hirtellous or on the main veins to
x 1-6 cm, apex acute,base acuteto obtuse, mar- subsericeous,sometimes with white arachnoid
gin entire; both surfacesglabrous;lateral veins hairs which soon disappear;lateral veins (4-)
2-3 pairs, slightlycurved, basal pair indistinctly 8-11 pairs,curved,basalpairdistinctlyto faintly
branched, reaching the margin far above the branched, reaching the margin usually below,
middleof the lamina;intercostalvenation slight- sometimes at or just above the middle of the
ly prominent;petiole 0.5-2 cm long, glabrous; lamina, sometimes also other lateral veins
stipules 1.2 cm long, glabrous.Staminate inflo- branched,a distinctsubmarginalvein in the low-
rescencesbranched;heads ca. 10-12, ca. 2 mm er partof the lamina;intercostalvenation (often
diam.; common peduncle 1.5-3 cm long, gla- very) prominent;petiole (1-)3-9 cm long, pu-
brous;perianthca. 1 mm high,glabrous;stamens berulousto hirtellous(to subhirsute);stipules2-
two, just exceedingthe perianth.Pistillateinflo- 6 cm long, densely reddish- to yellowish-brown
rescencespoorly branched;heads 2-3, globose, subsericeous to subhirsute to subvillous. Sta-
ca. 6 mm, in fruitup to 15 mm diam.; common minate inflorescencesbranched; heads many,
peduncle 2-4 cm long, glabrous;perianthca. 1 globose, 2-3 mm in diam.; common peduncle
Coussapoa
1 cm
FIG. 29. Coussapoaglaberrima:1, leafy twig with pistillateinflorescences(Proctoret al. 27080).
(1-?)3-7 cm long, puberulousto hirtellous;peri-
anth ca. 1 mm high, glabrous;stamensthree, far
exceeding the perianth. Pistillate inflorescences
branched;heads 6-10, sometimes partly fused,
(sub)globose,5-9 mm, in fruitup to 13 mm diam.;
common peduncle 3-6 cm long, 2-3 mm thick,
puberulousto hirtellous;perianthca. 2 mm high,
minutely puberulous;fruiting perianth orange.
Interfloralbracts spathulateto subpeltate,pub-
erulousat the apex.
Distribution(Fig. 7). Pacificcoastalregion(Ec-
59
uador and southern Colombia); in wet forest, up
to 1200 m.
Specimens studied. COLOMBIA. NARuio:Bay of
Tumaco, Rio Yanaje, 6 Jul 1955 (9), Romero-Casta-
ineda5283 (COL).
ECUADOR. CARCHI: Rio San Juan, nr. Chical, 12
km below Maldonado,1200 m, May 1978 (2), Madison
et al. 4764 (AAU, F, QCA, SEL,U); El Pail6n, 45 km
below Maldonado, 28 Nov 1979 (d), Madison et al.
7101 (SEL,U). ESMERALDAS: Nr. Lita, W of Rio Lita,
9 Feb 1981 (2), Berg 1259 (QCA, U). GUAYAS: Nr.
Bucay(=Grl.Elizalda),8-15 Jun 1945(9),CampE3674
60 Flora Neotropica

2 I!
cm1 1

FIG. 30. Coussapoa herthae: 1, leafy twig with pistillate inflorescences (Schulze-Rhonhof 1953); 2, staminate
inflorescences (Dodson 5776).
Coussapoa 61

(NY, U). IMBABURA: Parambas,30 May 1949 (9),Acos- diam.;common peduncle0.5-3 cm long, sparse-
ta Solis 12661 (F). Los Rios: Rio PalenqueBiol. St.,
km 56 on road Quevedo-Santo Domingo de los Co- ly to densely appressed-puberulous;perianthca.
0.5-1 mm high, glabrous;stamens two, just ex-
lorados, 27 Feb 1975 (6), Dodson 5776 (MO, QCA),
29 May 1976 (2), Dodson 6082 (MO), 20 Mar 1980 ceeding the perianth. Pistillate inflorescences
(a), Dodson et al. 9539 (SEL, U), 13 Feb 1974 (6), branched;heads 2-15, globose, ca. 3-4 mm, in
Gentry9892 (MO, U). PICHINCHA: 35 km N of Santo fruit up to 8 mm diam.; common peduncle 1-3
Domingode los Colorados,nr.bridgeover Rio Blanco, cm long, sparselyto densely appressed-puberu-
3 Feb 1974 (2), Gentry9606 (MO, U); San Carlosde
los Colorados,5 Oct 1935 (6),Schultze-Rhonhof1953 lous; perianth ca. 1-1.5 mm high, glabrous.
(B, type collection). Interfloral bracts small, subpeltate to (sub)
spathulate,minutely puberulousat the apex.
Coussapoa herthae shows general similarity to Distribution(Fig. 7). Western Guiana region
C. nymphaeifolia and in the echinate (dry) fruit-
(Surinam and French Guiana), in Brazil from
ing head to C. echinata.
The species is very variable in the shape and
Amapa througheasternPara to Maranhao;fur-
venation of the leaves.
ther, apparentlysegregatedfrom the main area
in the central part of the Amazon Basin; forest
of terrafirme.
20. Coussapoalatifolia Aublet, Hist. pl. Guiane
2: 955, t. 362. 1775; Trecul, Ann. Sci. Nat.
studied.SURINAM.Brownsberg,
Specimens 3 Sep
Bot. Ser. 3, 8: 94. 1847; Miquel, Fl. bras.4(1): 1915 (9), BW(Stahel & Gonggrijp)732 (K, MO, NY,
135. 1853;Berg,Fl. Suriname5(1): 280. 1975. P), 19 Sep 1931 (9), Van Emden s.n. (F, U, US), 23
Type. FrenchGuiana.Withoutlocality, - (9), Sep 1931 (st), VanEmden LVI (U); WilhelminaMts.,
Aublets. n. (holotype, BM). Fig. 31. upperGran Rio, Maup6dam, 20 Feb 1926 (st), Exp.
WilhelminaGebergte196 (U); without locality, - (2),
Coussapoa obovata Miquel, Het Instituut(1842):200, Focke s.n. (GH); plantationBergendaal,- (9), Focke
t. 3. 1843.Type.Surinam. PlantationBergendaal,- 9 (L, type collection of C. obovata= C. latifolia var.
(2), Focke 9 (holotype,L). obovata);plantationBlauberg,- (9), Focke 418 (U);
Coussapoalatifolia Aublet var. obovata (Miquel) Mi- Distr. Nickerie,ca. 20 km SW of Avanaverodam site,
quel,Fl. bras.4(1):135. 1853. 14 Nov 1976 (9),Heydeet al. 78 (U); WilhelminaMts.,
Coussapoa froesiiStandley,Publ.FieldMus.Bot.17: LucieRiver,2-5 km belowconfluencewithOost River,
162. 1937. Type. Brazil. Maranhao:Rio Maracas- 8 Sep 1963 (9), Irwinet al. 55487 (MO, NY, OXF, R,
sume region, 14 Sep 1932 (2), Fr6es 1907 (holotype, S, U, US); Lely Mts., 19 Sep 1975 (st), Lindemanet
NY; isotypes, A, G, K, U). al. 33 (U), 19 Sep 1975 (8), Lindemanet al. 73 (U), 4
Oct 1975 (9), Lindemanet al. 749 (K, U), 5 Oct 1975
Shrub or tree, mostly hemi-epiphytic, up to 35 (st), Lindemanet al. 799 (U); Jodensavanne-Mapane
m tall. Leafy twigs 4-9 mm thick, glabrous. Lam- Creekarea,camp 8, 21 Dec 1954 (st),Lindeman6938
ina coriaceous or subcoriaceous, ovate to elliptic, (U); CoppenameRiver, Doksi Creek,11 Oct 1954 (6),
271 (C, NY, U); TanjimamaRiver, Kodji
sometimes oblong to obovate, occasionally to Mennega Creek,25 Nov 1954 (6),Mennega521 (A, C, NY, U);
suborbicular,4-18(-25) x 3-15 cm, apex acute Voltzberg,22 Sep 1956 (st), Schulz (LBB) 7787 (U);
to shortlyacuminateto obtuse to rounded,base Jodensavanne-MapaneCreekarea, 5 May 1967 (st),
acute to obtuse or to subcordate,marginentire, Vreden(LBB) 11747 (U). -
FRENCH GUIANA. Without locality, (2), Aublet
plane or more or less revolute towardsthe base; s.n. (BM, type collection), Feb 1868 (2), Aubry-Le-
uppersurfaceglabrous,lower surfaceglabrousor comptes.n. (FI); Sinnamary,road to St. Elie, 22 Oct
sparselyappressed-puberulous withhairsof about 1981 (9),Billiet et al. 1108 (U); Montagnede Kaw, 13
equal length; lateral veins 4-7 pairs, (slightly) Dec 1954 (9), Cowan38774 (F, K, NY, P, US); Mont
Belvedere, 18 Nov 1974 (2), De Granvilleet al. B5191
curved,basal pairsmostly branched,sometimes (CAY, U); Oyapock River, Myuli Creek, Les Trois
(especially)in narrowleaves unbranched,reach- Sauts, 15 Sep 1977 (6), Grenand1478 (CAY, U); with-
ing the marginat to below or (especiallyin ob- out locality, - (9), Leman s.n. (P); Cayenne, - (9),
ovate leaves) above the middle of the lamina, Martins.n. (or 112) (K);without locality, - (9), Mel-
the otherlateralveins sometimespoorlybranched inon s.n. (A, P); Maroni River, 1864 (9), Melinon 18
(GH, K, LE,P);AntecumePata,at confluenceof Itany
(furcate); intercostal venation (almost) plane; Riverand MaroniRiver, 14 Nov 1977 (2), Moretti836
petiole 1-7 cm long, sparselyto ratherdensely (CAY); Kourou River, 3 km above Couy Creek, 19
minutely appressed-puberulous;stipules 0.5- Sep 1967 (6), OldemanB1338 (CAY, U); Cayenne,-
1.5(-2.5) cm long, sparselyto densely whitish to (st),Poiteaus.n. (FI);Acarouany,1857 (6),Sagot 1136
brownish appressed-puberulous, sometimes (B, BM, G, GH, K, S, U); Sinnamary,roadto St. Elie,
15 Sep 1978 (6), Sastre 6130 (P, U).
brownish subvelutinous. Staminate inflores- BRAZIL. AMAPA:"Gafcho area," 21 Oct 1979 (9),
cencesbranched;heads many, globose, ca. 1 mm Austinet al. 7149 (U); Rio Araguari,between 1?26'N,
FIG. 31. Coussapoa latifolia: 1, leafy twig with pistillate inflorescences (Ducke (HJBR) 18456); 2, staminate inflor
Coussapoa
51?58'Wand 109'N,51?52'W,11 Sep 1961 (2), Pires
et al. 50864 (NY, OXF, P, R, U, US); Rio Araguari,
nr. 1?11'N,52?8'W,1 Oct 1964 (6), Pires et al. 51409
(F, GH, NY, U); Col6niaTorrao,2?25'N,51015'W,29
Aug 1962 (2), Pires et al. 52656 (NY, OXF, U, US).
AMAZONAS:
Road Manaus-Caracarai, km 97, 30 Aug
1979 (e), Cidet al. 942 (U); roadManaus-PortoVelho,
km 235, IgarapeAcu, 24 Nov 1973 (d), Lleras et al.
P19661 (F, M, MO, NY, P, S, U); road Manaus-Ca-
racarai,km 159, 20 Sep 1974 (6), Pranceet al. 22720
(K, MO, NY, S, U, US). MARANHAO: Rio Maracas-
sume region, 14 Sep 1932 (2), Frogs 1907 (A, G, K,
NY, U, type collection of C. froesii). PARA:Col6nia
Benjamin Constant, Braganoa, 21 Nov 1908 (2),
Anonymous(HAMP)9783 (G);Sta.Izabel,10 Sep 1922
(2), Ducke(HJBR)18456 (RB);Col6niaAugustoMon-
tenegro,IgarapePitor6, 18 Sep 1958 (2), Frogs34651
(IAN); road Belem-Brasilia,km 93, 19 Sep 1959 (2),
Kuhlmannet al. 263 (NY, US); road Belem-Brasilia,
km 301-306, 7 Aug 1960 (2), Oliveira985 (U); Rio
Jari,Monte Dourado, 16 Nov 1967 (2), Oliveira3550
(NY), 9 Nov 1967 (2), Oliveira3599(NY), 8 Nov 1967
(9),Oliveira3682 (NY), 23 Jan 1968 (st), Oliveira3954
(NY), roadCapanema-Maranhao, km 96,27 Oct 1965
(2), Prance et al. 1720 (MO, NY, S, U, US), km 98,
21 Aug 1964 (2), Prance et al. 58783 (U); Peixe Boi,
26 Oct 1907 (2), Seguiera(HAMP)8808 (BM, G, U);
Rio Jari,Monte Dourado, 15 Oct 1968 (d),N. T. Silva
1207 (NY), 28 Oct 1968 (2), N. T. Silva 1321 (NY, U,
US); Rio Jari, between Pilao and Repartimento, 12
Nov 1968 (2), N. T. Silva 1380 (IAN, NY, U, US).
Coussapoa latifolia is very closely related to
C. viridifolia. The species shows affinities with
C. microcephala, C. microcarpa, C. pachyphylla,
and also with the Central American C. macer-
rima. Because of similarities in the vegetative
parts, C. latifolia can be confused with C. ortho-
neura, especially specimens with elliptic leaves.
21. Coussapoa leprieurii Benoist, Bull. Mus. Hist.
Nat. Paris 30:103. 1924. Type. French Guiana.
Without locality, - (6), Leprieur s.n. (holo-
type, P). Fig. 32.
Tree, hemi-epiphytic, up to 10 m tall. Leafy
twigs 5-8 mm thick, brown(ish) puberulous to
hirtellous to hirsute, solid. Lamina coriaceous,
(broadly) ovate to (broadly) elliptic to (broadly)
obovate to oblong, 5-18 x 1.5-12 cm, apex ob-
tuse to subacuminate to acute, base subcordate
to obtuse or acute, margin entire, often more or
less revolute, especially towards the base; upper
surface scabrous, lower surface puberulous, on
the main veins to hirtellous, sometimes also white
arachnoid hairs present; lateral veins 6-7 pairs,
straight or curved, basal pair unbranched, reach-
63
ing the margin at or below the middle of the
lamina, the other lateral veins sometimes
branched;intercostalvenation prominent;peti-
ole 1.5-3 cm long, puberulousto hirtellous;stip-
ules 0.5-1 cm long, yellowish to brown hirsute
to subsericeous;terminalbuds often more or less
swollen. Staminate inflorescences branched;
heads many, globose, ca. 6 mm diam.; common
peduncle4-7 cm long, puberulousto hirtellous;
perianthca. 1 mm high,glabrous;stamensthree,
far exceeding the perianth. Pistillate inflores-
cences unbranched;heads globose, ca. 5-8 mm
diam.; peduncle0.5-2.5 cm long, puberulousto
hirtellous;perianthca. 1-2 mm high, glabrous.
Interfloralbracts spathulate to subpeltate, pu-
berulous.
Distribution (Fig. 7). In (eastern) French
Guiana and in Brazilin easternPara;in upland
and riverineforest.
Specimens studied. FRENCH GUIANA. Without
locality(3), Leprieurs.n. (P, type collection);Ap-
prouague River,betweenSapokayeCreekandGrand
CanoriFalls,23 Oct1968(6),OldemanT243a(CAY);
Sapkaye Creek,22 Oct1968(3),Oldeman B1946(CAY,
K, P, U); Approuague River,MapaouFalls,26 Jan
1970(6),OldemanB2861(CAY,U);OyapockRiver,
3 km aboveCamopi,8 Apr 1970(6), Oldeman3062
(CAY,P, U).
BRAZIL.PARA:Ilhasde Breves,FuroMacujubim,
16 Nov 1912 (6), Ducke(HJBR)18462 (RB);Alto
Quatipuir,Dec 1899 (6),Huber(HAMP)1775 (F, MG).
Coussapoaleprieurii,one of the few species in
which the lamina is scabrousabove, appearsto
be ratherclosely relatedto C. sprucei.Specimens
with an elliptic to oblong lamina having arach-
noid hairsbeneathare very similarto C. sprucei.
Specimens with obovate leaves are reminiscent
of (the unrelated)C. asperifoliassp. asperifolia.
22. CoussapoalongepedunculataAkkermans&
C. C. Berg,Proc. Kon. Ned. Akad. Wetensch.
Ser. C. 85(4): 453. 1982. Type. Peru. Loreto:
Prov. Maynas, QuebradaSucusari,Llachapa
camp of Explorama,N side of Rio Napo, be-
low Mazan,6 Nov 1979 (6),Gentryet al. 27566
(holotype, U; isotype, MO). Fig. 33.
Tree, up to 25 m tall, hemi-epiphyticor ter-
restrial.Leafy twigs 1-1.5 cm thick, brown hir-
sute. Lamina coriaceous,oblong to elliptic, 15-
50 x 7-30 cm, apex obtuse, base cordate to
roundedmarginrepandto subcrenate;uppersur-
64
FG2 3,si1e tg ts
FIG. 32. Coussapoa leprieurii: 1, leafy twig with staminate inflorescences (Oldeman B1946); 2, pistillate
inflorescences (Oldeman 3062).
face scabridulousto smooth, sparsely hispidu-
lous to hirtellous,often ? bullate, lower surface
sparselyhirtellous on the veinlets, between the
lateralveins white arachnoidhairs, ratherdense
or sparse and then concentratedat the margin;
lateralveins 9-11 pairs, more or less curved, at
leastthe lowerones branched,basalpairreaching
the marginfar below the middle of the lamina;
intercostalvenation prominent;petiole 3-10 cm
long, brownhirsute;stipules 2-6 cm long, some-
times subpersistent,hirsuteto subsericeous,with
brown hairs of differentlengths. Staminate in-
florescences rather poorly branched; common
Flora Neotropica
iore s l a 1 cm
peduncle9-16 cm long, the branchesin two clus-
tersand all or some of them very short,peduncle
and branches brown hirtellous to pubescent;
heads many, (sub)globose,3-5 mm diam.; peri-
anth ca. 1 mm high, sparselyminutely puberu-
lous; stamensthree, exceedingthe perianth.Pis-
tillate inflorescences branched; heads 3-10,
(sub)globose,5-8 mm, in fruitup to 15 mm diam.;
common peduncle 8-20 cm long, peduncle and
branchesbrownhirtellousto subtomentose;peri-
anth ca. 1 mm long, glabrous;fruitingperianth
red. Interfloralbractsca. 1 mm long, spathulate,
subpeltate,at the apex puberulous.
Coussapoa 65

?~~~~~~~~~~~~~~~~~~~~~~~~~~~L
~ ' ~/ ~
1o g . '',
..
_ i , ., '? ... ~ ~ ~.'. ~ ~ ~~~~~~~~~~.
,...

?'
..? . . ... .

: ~. -
or ?- -.' .
''.

"'

???~ ~~~~~~~~~~'
?
-r
.'
. . ' - ..;
.'
..

*~~
-, ~ ~~~~ -..s

??.?

,, -
... ,,

~~~~~~~~~~~~~~~~~~...
-~~~~~~~~~~~~
~ ~
: ~ ~~ ,. ....

''' =, .
I' ~~~~~~ .~~~~~~~~~~~.
'? IY~~~
.'? ..- *-. .I

r~~~~~~~~~~~~~~~~. o o-

..,.

I
'~'~~~~~~~~~~~~~~~~~~~~~'
~'C ?1 .'E":? .

..? -?~~~~1
?~ ",

'~~~~~~~~'

?~~~~~~~~~?
c. ?
FIG 33 Cossaoa onepeunclat: 1 lafytwi wih samiat
iniorsceces(Gntr etal.2756)
66 FloraNeotropica

Distribution (Fig. 8). Peru (Loreto) and Ec- and allied species. In the shape and venation of
uador (Napo); in non-inundatedor periodically the leaves it resembles C. contorta,from which
inundatedforest. it clearlydiffersin the 2-staminateflower.
Specimensstudied.ECUADOR.NAPO:ParqueNa-
cional Yasuni, 16-19 Jan 1988 (d), Ceron M. 3495 24. CoussapoamanuensisC. C. Berg,Proc.Kon.
(BG), 9-19 Jan 1988 (9), Neill et al. 8289 (BG). Ned. Akad.Wetensch.Ser.C, 86(3):305. 1983.
PERU. LORETO: Prov. Maynas,QuebradaSucusari, Type. Peru. Madrede Dios: ParqueNacional
Llachapacamp of Explorama,N side of Rio Napo, del Manu, Rio Manu, Cocha Cashu Station,
belowMazan,6 Nov 1979(a),Gentry etal.27588(MO,
Prov.Maynas,PuertoAlmendras, 26 Oct 1980 (v), Foster 5641 (holotype, F).
U, typecollection);
12Jan1982(9),Vdsquez 2852(BG);Iquitos,RioItaya, Fig. 35.
BuenaSuerte,16 Nov 1986(9), Vdsquez et al. 8380
(BG);Nauta,12Dec 1986(9),Vdsquez etal.8600(BG). Tree, hemi-epiphytic,25 m tall. Lamina co-
riaceous,broadlyelliptic to obovate, 10.5-19 x
C. longepedunculataresembles C. nymphaei- 8-13 cm, apex shortlyacuminate,base rounded
folia in many features. It is distinct from the to subcordate,marginentireto subcrenate;upper
latter especially in the indumentum, the wider surface glabrous or sparsely puberulous at the
(looser)venation, and in the scabridulousupper base of the midrib, lower surface densely pu-
surface of the lamina. The species also shows berulousto hirtellous(to subtomentose);lateral
affinitieswith C. spruceiand C. leprieurii. veins 8-12 pairs, almost straight,the basal pair
unbranched,reachingthe marginbelow the mid-
23. CoussapoamacerrimaAkkermans& C. C. dle of the lamina; intercostalvenation more or
Berg, Proc. Kon. Ned. Akad. Wetensch. Ser. less prominent;petiole 3-5 cm long, puberulous
C, 85(4): 455. 1982; Burger,FieldianaBot. 40: to hirtellous;stipulesca. 6 cm long, chartaceous,
133, t. 22. 1977 (sub C. contorta).Type. Costa subsericeous (to subhirsute), with rather con-
Rica.Puntarenas:Above PalmarNorte de Osa, spicuous (parallel)venation. Staminate inflores-
22 Feb 1951 (d),Allen 5949 (holotype, F; iso- cences unknown. Pistillate inflorescencesun-
types, A, F). Fig. 34. branched;heads globose, ca. 4 mm, in fruit up
to 12 mm diam.; peduncle 2.2-2.7 cm long, pu-
Tree, up to 25 m tall. Leafy twigs 2-4 mm
thick,sparselyappressed-puberulous, on the scars berulous;perianthca. 1 mm high, sparselymi-
of the stipules distinctly longer hairs. Lamina nutely puberulous;fruitingperianthyellow. In-
bractsspathulate,puberulousat the apex.
subcoriaceous,elliptic to oblong to subobovate, terfloral
5-13 x 4-7 cm, apex shortly acuminate, base Distribution (Fig. 8). Peru (Madre de Dios).
acuteto obtuse, marginentire;uppersurfacegla- Known only from the type collection.
C. manuensis shows no distinct affinitieswith
brous, lower surface on the main veins rather
with hairs of dif- any of the other Coussapoaspecies.
sparselyappressed-puberulous
ferent lengths or partly strigillose;lateral veins
25. Coussapoamicrocarpa(Schott)Rizzini, Du-
7-13 pairs, straight to curved, basal pair un-
senia 1(5): 295. 1950. Fig. 36.
branchedreachingthe marginbelow the middle
of the lamina;intercostalvenation plane;petiole BrosimummicrocarponSchott in Sprengel,Syst. Veg.
2-5 cm long, sparselyappressed-puberulous with 4 (Curr.Post.,App.):403. 1827.Type.Brazil.With-
hairs of different lengths; stipules 1-2 cm out locality, - (Y),Schott s.n. (holotype, W, de-
stroyed).
long, (sub)sericeous. Staminate inflorescences Coussapoaschottii Miquel, Fl. bras. 4(1): 137. 1853,
branched; heads many, globose, ca. 1-2 mm as a synonym of Brosimummicrocarpon.
diam.; common peduncle 2-3 cm long, ap- CoussapoaschottiiMiquel var. lanceolataMiquel, Fl.
bras.4(1): 137. 1853. Type. Brazil.Withoutlocality,
pressed-puberulous; perianthca. 1 mm high,gla- - (Y),Pohl s.n. (holotype,U; isotypes, B, M (herb.
brous;stamens two, far exceedingthe perianth. Zuccarini105 sub forma angustifolia)).
Pistillate inflorescences unknown. Interfloral CoussapoaschottiiMiquel var. longifoliaMiquel, Fl.
bractsabsent. bras.4(1): 137. 1853. Type. Cult.bot. gardenMin-
Distribution(Fig. 7). Costa Rica (Puntarenas); chen, 1835 (2), herb. Zuccarini103 (holotype,M).
in forest at 450 m. Known only from the type Coussapoafontanesiana Trecul, Ann. Sci. Nat. Bot.
Ser. 3, 8: 94. 1847. Syntypes. Brazil. Sao Paulo:
collection. Withoutlocality, - (8 and 2), Gaudichaud922 (lec-
This speciesmay be relatedto C. microcephala totype (Q),P).
Coussapoa
FIG. 34. Coussapoa macerrima: 1, leafy twig with staminate inflorescences (Allen 5949)
Shrubor tree,terrestrialor occasionallyhemi-
epiphytic,up to 20 m tall. Leafy twigs 2-5 mm
thick, white to yellowish appressed-puberulous
to hirtellousor to hirsute, sometimes also with
white to brownisharachnoidhairs. Lamina co-
riaceous,subovate to ovate oblong to elliptic to
67
I
1 cm
(sub)obovateor to lanceolate, 3-21 x 2-7 cm,
apex shortly acuminate to acute or to obtuse,
base acute to obtuse to rounded, marginentire,
usuallyrevolute towardsthe base; upper surface
glabrous,lower surfaceglabrousor sparselyap-
pressed-puberulous(to strigose) to (sub)hirsute
 FIG.c3.m cousapoa anuenss 1 leves sipue n itlaeiforsecs(otr54)

FIG. 35. Coussapoamanuensis:1, leaves, stipulesand pistillateinflorescences(Foster5641

Coussapoa 69

"-; ~~~~~~~~~~~~~~~~~~~~~~'?
~C ?: ' .. ?

~~~~~~~~~~~~~~.. U ? :

:7:I
.i

??
1~~~~~~~~~~~~~ ..

~ ~ ~ ~
~~~~~~~~~~~~~~~.
?: : .??,.

~~~~~~~~~~~~~~~~.2
?~ ~~~~~~~~~~?

~~?. ?'''" .:? .. ".'

'
".i ii{. ..
I-

FIG. 36. Coussapoa microcarpa: l,leafy twig with pistillate inflorescences (Mexia 5115); 2, leafy twig with
staminate inflorescence (Hage 31).

on the main veins; lateral veins 6-11 pairs,
straight, basal pair unbranched, reaching the
margin below the middle of the lamina; inter-
costal venation plane to slightlyprominent;pet-
iole 1-4 cm long, appressed-puberulousto hir-
tellous to (sub)hirsute;stipules 1-7 cm long,
yellowish puberulousto subsericeousto hirsute
or also with white to brownish arachnoidhairs.
Staminate inflorescencesbranched;heads 5-9,
globose, ca. 2-3 mm diam.; common peduncle
1-2 cm long, puberulousto hirtellous;perianth
ca. 1 mm high, minutely puberulous;stamens
two, far exceeding the perianth.Pistillate inflo-
rescencesunbranchedor rarelybranched;heads
1(-5), globose, ca. 3-5 mm, in fruitup to 10 mm
diam.; (common) peduncle 2-5 cm long, pu-
70
berulous to hirtellous; perianth ca. 1 mm high,
glabrous; fruiting perianth yellow and orange. In-
terfloral bracts small, narrowly spathulate, often
only a few are sometimes absent.
Distribution (Fig. 7). Brazil, from Rio Grande
do Sul to Espirito Santo, common in forests up
to 1800 m and in restinga vegetation; moreover,
apparently in more or less isolated populations
further north, in Bahia (near Ilheus) and in Pa-
raiba and Pernambuco (on the top of low moun-
tains?).
Specimensstudied. BRAZIL. Without (certain)lo-
cality, 1814-1817 (9), Bowie& Cunningham57 (BM),
- (d and 2), Burchell3148 (GH, K, P), - (6), Glaziou
1942 (P), - (6), Glaziou 2016 (P), - (6), Glaziou 6009
(C), - (9), Grahams.n. (K), - (2), Pohl s.n. (B, M
(herb.Zuccarini105), U, type collectionof C. schottii
var. lanceolata), 11 Nov 1822 (d), Riedel s.n. (LE), 5
Oct 1863 (st), Warmings.n. (C), 1835 (d), (cult.in hort.
bot. Munchen),herb.Zuccarini103 (M, typecollection
of C. schottiivar. longifolia).BAHIA:Nr. Ilheus, 1 Apr
1965 (2), Belem et al. 644 (U); without locality, - (9),
Blanchets.n. (BM,M, U); nr. Ilheus, 1836 (2), Blanchet
2327 (G, M, P), 30 Nov 1970 (6),Emygdio3018 (=Em-
merich3556) (R); Itabuna, 16 Dec 1966 (9),Emygdio
et al. 2447 (=Emmerich et al. 3002 = Andradeet al.
2340) (R);nr. Ilheus,23 Nov 1970 (6),Hage 31 (GUA,
U), 24 Nov 1971 (2),Pinheiro1716 (U); CasteloNovo,
Oct 1821 (2), Riedels.n. (LE),Mar 1822 (2), Riedel664
(LE). ESPiRITOSANTO:Mun. Linhares, Res. Biol. Soor-
etama, 14 Mar 1972 (2), Sucre 8690 (U). MINASGERMS:
LagoaSanta, - (6),Lund 139 (C);Viqosa,Fazendade
Aguada,29 Sep 1930 (2), Mexia 5115 (A, BM, F, G,
GB, GH, K, MO, NY, S, U); Lagoa Santa, - (?),
Warming1917 (C), 13 Oct 1863 (?), Warming1918
(C), 5 Oct 1863 (6), Warming1919 (C), 29 Dec 1864
(8), Warming 1942 (C). PARAiBA:Serro Araripe, Santa
Ana, Aug 1921 (2), Luetzelburg12471 (M). PARANA:
Mun. Guaratuba,Garuva, 28 Jul 1960 (9), Duarte et
al. 5328 (F, RB); Jacarehy,27 Sep 1908 (st), Dusen
6635 (GH, NY); Caita de Agua, 28 Aug 1910 (st),
Dusen 10134 (NY); Morretes,7 Sep 1910 (2), Dusen
10186 (F, GH, K, L, MO, NY, P, S);Jacarehy,25 Mar
1911 (6),Dusen 11419 (F, GH, NY); Rio Cubatao,28
Dec 1911 (8), Dusen 13653 (F, GH, NY, S); Jacarehy,
26 Mar 1914 (6), Dusen 14718 (F, GH, K, MO, NY,
P), 11 May 1915 (st), Dusen 17030 (NY); CerroAzul,
2 Oct 1949 (6),Hatschbach1475 (S);Mun. Paranagua,
Rio Cachoeirinha,25 Aug 1951 (6), Hatschbach2458
(RB);Mun. Paranagua,PortoD. Pedro2?,31 Jan 1961
(8), Hatschbach7807 (RB);Mun. CerroAzul, Turvo,
6 Feb 1961 (6), Hatschbach7833 (RB);between Cu-
ritiba and Paranagua,10 Nov 1948 (6), Hatschbach
16304 (F, K); Mun. Guaraquecaba,Rio do Cedro, 8
Nov 1968 (9), Hatschbach18680 (C, MO, NY); Mun.
Bocaiuvado Sul, Sesmaria,Rio Capivari,11 Nov 1968
(6),Hatschbach20253 (C);Mun. Guaratuba,Rio Tup-
itinga,29 Apr 1972 (6),Hatschbach29629 (NA);Para-
nagua,7 Mar 1914 (2), Jonnson5a (F, GH, K, NY, S);
Flora Neotropica
Portode Cima, 26 Jun 1914 (6),Jonsson610a (F, GH,
K, NY, S); Rio Sao Joao, W of road to Joinville, N of
Garuva, 16 Jul 1966 (st), Lindemanet al. 1857 (U);
Serrado Mar, nr. old road Curitiba-Morretes,Bella
Vista, 25 Jul 1967 (st), Lindemanet al. 5682 (U); nr.
Pontaldo Sul, 20 Jun 1967 (st), Lindemanet al. 5748
(U); Caioba, 35 km S of Paranagua,10 Nov 1947 (9),
Tessmanns.n. (MO); Mun. Guaraqueqaba,Serrinha,
13 Apr 1967 (6), Tessmann s.n. (MO). PERNAMBUCO:
"Gurgaf"(=Gurjao?),14 May 1952 (8),Ducke&Lima
104 (R). Rio DEJANEIRO:Rio de Janeiro, - (2), Blan-
chets.n. (NY), 1835 (8),Blanchets.n. (FI);Itatiaia, 10
Jan 1910 (st), CamposPorto863 (RB);Rio de Janeiro,
CollegioAnglo-Brasileiro,22 Sep 1916 (2), Constan-
tino (HJBR) 19689 (RB);Restingade Itapeba,nr. Ca-
nal das Taxas, 22 May 1963 (2), Carauta178 (GUA,
K); Rio de Janeiro,Gavea, 11 Aug 1974 ($), Carauta
1716 (F, GUA, K, U); Rio de Janeiro,Morroda Urca,
17 Apr 1975 (st), Carauta1780 (F), (2), Carauta1781
(GUA);Rio de Janeiro,BotanicalGarden,9 Nov 1978
(2), Carauta3050 (RB); Rio de Janeiro,Alto da Boa
Vista, 7 Aug 1974 (2), Carautaet al. 1714 (F, GUA,
RB); between Serrade Macah6 and Novo Friburgo,
Sao Pedro, 18 Oct 1977 (2), Carautaet al. 2722 (GUA,
U); Rio de Janeiro,Pao de Acucar,26 Aug 1979 (2),
Carautaet al. 3166 (GUA, U); Turin, 1857 (2), Cas-
aretto643 (G); JoatingaJoa, 24 Mar 1959 ($), Duarte
et al. 4650 (RB);Dois Irmaos, 19 Feb 1921 (2),Ducke
et al. (HJBR) 16361 (RB); Meio Serra, old road to
Petr6polis,3 Nov 1928 (9),Ducke(HJBR)21200 (RB);
Serrade Itatiaia,Mont Serrat,21 Oct 1903 (2), Dusen
2160 (S); Rio de Janeiro, Prainha, 4 Dec 1978 (2),
Ferreira493 (RB); Serra dos Orgaos, Apr 1837 (6),
Gardner732 (BM, K); Rio de Janeiro, Gavea, Aug
1837 (9), Gardner5632 (BM, K);Teres6polis,1100 m,
31 Jan 1978 (9), Gentryet al. 918 (MO, U); Rio de
Janeiro,Gavea, 19 Jan 1861 (st), Glazious.n. (P), 28
Aug 1861 (st), Glaziou s.n. (P); Rio de Janeiro, Co-
pacabana,4 May 1866 (9), Glaziou 1013 (C, NY, P);
Rio de Janeiro,Gavea, 28 Apr 1867 (6), Glaziou1138
(C, P); Rio de Janeiro, - (st), Glaziou 1881 (G); Rio
de Janeiro,Gavea, 28 Nov 1869 (6), Glaziou4937 (P);
Rio de Janeiro,Copacabana,1871 (6), Glaziou 4938
(C:K, P); Rio de Janeiro,Tijuca,6 Feb 1871 (2), Gla-
ziou 6009 (K, P); Rio de Janeiro, 12 Aug 1876 (2),
Glaziou8936 (A, K, LE, P); Serrados Orgaos,23 Mar
1880 (2), Glaziou12166 (C, F, G, K, LE,P);Petr6polis,
BairroAmoeda, Dec 1943 (8), Goes et al. 815 (GUA,
RB); Recreio dos Bandeirantes,30 km W of Rio de
Janeiro, 5 Mar 1964 (6), Lems s.n. (NY); Rio de Ja-
neiro, 1839 (6), Luschnaths.n. (LE);betweenSerrade
Macah6and Novo Friburgo,1000 m, 18 Oct 1977 (2),
Maas et al. 3325 (K, U); Petr6polis,Vale das Videiras,
1800 m, 6 Jan 1973 (2), Martinelli 150 (U); Rio de
Janeiro, 1867 (8), Miers s.n. (BM); Rio de Janeiro,
Tijuca, 1879 (2), Miers s.n. (BM);Rio de Janeiro,Jul
1878 (6), Miers 2714 (K, P); Rio de Janeiro, Tijuca,
Jul 1878 (9),Miers3792 (K);Rio de Janeiro,Pedreiro,
Jul 1878 (6), Miers 3858 (K, P); Restingade Jacare-
pagua,Res. Biol., 12 Apr 1967 (6),Moreira46 (F, GH,
GUA, US); Rio de Janeiro,Corcovado,20 Sep 1974
(8),Mosen 2614 (S);Rio de Janeiro,Sep 1862 (9), Na-
deauds.n. (P);Rio de Janeiro,Copacabana,Oct 1862
Coussapoa
(a),Nadeauds.n. (P);Rio de Janeiro,BotanicalGarden,
8 Mar 1936 (9), Occhioni(HJBR) 35333 (RB); Serra
dos Orgaos,Rio Paquequer,4 Mar 1949 (9), Pereira
39 (RB); Rio de Janeiro, Botanical Garden, 23 May
1961 (9),Pereira5670 (B, M); Rio de Janeiro,Gavea,
3 Oct 1927 (9), Pessoal do Horto Florestal 654 (RB);
Rio de Janeiro,Tijuca,14Jun 1969 (st),Plowman2919
(=Sucre 5209) (GH, K, US); Rio de Janeiro, 1829 (a),
Riedel 7 (LE), Sep-Dec 1831 (9), Riedel 75 (LE, NY,
RB, US), 1832 (st), Riedel et al. s.n. (LE), 1832 (st),
Riedelet al. I (LE);Teres6polis,Rio Imbui,2 Oct 1952
(9), Rizzini 144 (RB); Belem, Oct 1867 (9), Schwacke
1067 (RB); Rio de Janeiro, Corcovado, 2 Nov 1883
(9), Schwacke6783 (RB), 11 Mar 1883 (2), Schwacke
8418 (R); Rio de Janeiro, Copacabana,25 Oct 1967
(9),Sucre 1783 (F, GUA, K, RB);Cabo Frio, Restinga
do Per6, 14 Sep 1968 (9), Sucre3633 (GUA, U); Res-
tingade Jacarapagua,8 May 1969 (9),Sucre4989 (U);
Parades da Subida,Pedrada Panela,3 Nov 1971 (9),
Sucre 7878 (U); Rio de Janeiro,Gavea, Nov 1899 (8),
Ule s.n. (R); Rio de Janeiro,Copacabana,- (9), Ule
1693 (P);Itatiaia,1935 (a),Zikan (HJBR)29279 (RB).
Rio GRANDEDO SUL: Fazenda das Almas, nr. Pal-
mares,Jan 1945 (8),Buck(in Rambo)26425 (B);P6rto
Alegre,1 Oct 1957 (9), Camargo1849 (B);MorroTer-
es6polis, Dec 1940 (8), Leite 413 (A), Dec 1942 (8),
Leite 872 (A); P6rto Alegre, 2 Jun 1893 (9), Malme
826 (S), 26 Feb 1902 (6), Malme 1428 (R, S), 16 Oct
1932 (d and 9), Rambo 426 (F, MO, S), 10 Nov 1946
(9),Rambo27092 (FI, S), 2 Oct 1948 (9),Rambo37782
(C), 1 Dec 1948 (9), Rambo 38427 (B);Lami, nr. Via-
mao, 3 Jan 1949 (6),Rambo39413 (F);EstacaoPereci,
14 Jan 1949 (9),Rambo 39780 (P);Ilha das Flores,22
Apr 1949 (9), Rambo 41173 (C, K, US); Lagoa dos
Barros,nr. Osoria, 14 Feb 1949 (9),Rambo 44743 (P);
Fazendado Arroio,nr. Osoria,4 Jan 1950 (a),Rambo
45136 (K);Lagoados Quadros,nr.Torres,21 Feb 1950
(a),Rambo 45904 (K);nr. Tramandai,5 Mar 1950 (a),
Rambo 46145 (P); without locality, - (9), Sello 241
(B, P, US); Rio Tapuchy, 1837 (6), Tweedie29 (K);
Col6nia Sao Pedro Torres, 12 Nov 1968 ($), Vianna
et al. 5465 (U). SANTACATARINA:
Rio Claro, Linha de
Joinville, 30 Oct 1882 (6), Anonymuss.n. (RB); Ilha
da SantaCatarina,8 Dec 1950 (9), Duarteet al. 3410
(F, K, NY, RB, U); Ibirama,13 Dec 1953 (8), Gevieski
85 (B, NY, S, US); Rio Pirai,6 Jan 1950 (6),Hans 335
(R, RB); Brusque,Mata Sao Pedro, 30 Dec 1949 (a),
Klein 79B (= Veloso97) (RB, US); Morrode Fazenda,
Itajai,25 Mar 1954 (6), Klein 772 (S, US); Morrodos
Conventos, E of Ararangua,5 Nov 1972 (6), Lima
20795 (U), 15 Nov 1971 (9),Lindemanet al. 9105 and
9110 (U); Sombrio, 3 Sep 1945 (8), Reitz 1912 (F, R,
NY); Sao Franciscodo Sul, 9 Jan 1951 (9),Reitz 3809
(US); Campo do Massiambfi,Palhoca, 24 Sep 1953
(a),Reitz et al. 975 (NY, S); Morrode Fazenda,Itajai,
3 Mar 1954 ($),Reitz et al. 1688 (NY, S, US); Sombrio,
8 Feb 1946 (8), Reitz et al. 2008 (US); Cunhas,Itajai,
5 Aug 1954 (8), Reitz et al. 2027 (US); Morrode Res-
sacada,Itajai,29 Mar 1957 (9),Reitz et al. 2913 (US);
Morrodo Bau, Itajai, 1 Nov 1951 (6),Reitz 4167 (NY,
S);TrdsBarras,Garuva,Sao Franciscodo Sul, 24 Aug
1957 (9),Reitz et al. 4692 (NY); Campodo Massiam-
bi, Palhoca, 19 Dec 1952 (9), Reitz et al. 4965 (F, G,
71
NY, S, U), 20 Dec 1952 (6), Reitz et al. 5030 (F, G,
NY, S, U); Mina Velha,Garuva,Sao Franciscodo Sul,
8 Nov 1957 (9), Reitz et al. 5611 (B, US); Serrado
Matador,Rio do Sul, 26 Jan 1959 (2),Reitz et al. 8294
(G, M);Ilhada SantaCatarina,6 Mar 1962 (8),Sehnem
7993 (B);P6rtoBelo, IlhaJoaoda Cunha,31 Mar 1957
(Q),L. B. Smith et al. 12307 (NY, R, US). SAO PAULO:
Mun. Picinguaba,Picinguababeach, 2 Oct 1975 (9),
Araujoet al. 848 (RB);Mun.Iguape,Morrodas Pedras,
Aug 1915 (8), Brade 7887 (R), 1917 (9), Brade 7949
(R, RB);Mun. Iguape,IguapeIsland, 19 Feb 1965 (2),
Eiten et al. 6214 (MO, US); withoutlocality, 1917 (6),
Frazas(HJBR)13059 (RB);1833 (2), Gaudichaud992
(P);SaoPaulo,Parquedo Estado,4 Sep 1933(6),Hoehne
30923 (F, GUA, NY, P); road Caraguatatuba-Uba-
tuba, 10 Oct 1968(9),LeitaoFilho 685 and686 (GUA);
Santos, 1 Dec 1874 (2), Mosen 2941 (C, S).
This species appears to be related to C. latifolia
and C. microcephala. It is quite variable in the
shape and dimensions of the lamina. A form with
relatively long and narrow leaves is cultivated in
several European greenhouses and (in The Neth-
erlands) as an indoor ornamental.
26. Coussapoa microcephala Tr6cul, Ann. Sci.
Nat. Bot., Ser. 3, 8: 96. 1847; Miquel, Fl. bras.
4(1): 136. 1853; Berg, Fl. Suriname 5(1): 282.
1975. Type. Guyana. Pomeroon River, - (6),
Schomburgk 876 (holotype, P; isotypes, BM,
F, FI, NY). Fig. 37.
Coussapoafagifolia Klotzsch,Linnaea20: 528. 1847;
Miquel, Fl. bras. 4(1): 136. 1853. Type. Guyana.
Pomeroon River, - (8), Schomburgk 1366 (=876?)
(holotype,B; isotype, K).
CoussapoacuneataMiquel, Fl. bras. 4(1): 138. 1853.
Type. Guyana. Berbice River, - (6), Schomburgk
287 (holotype, U; isotypes, BM, G, GH, K, L, P,
OXF, U).
Tree, often hemi-epiphytic, up to 20 m tall.
Leafy twigs 2-9 mm thick, sparsely to densely
yellowish appressed-puberulous to hirtellous to
subhirsute or also with white to brown arachnoid
hairs. Lamina coriaceous (to subcoriaceous),
ovate to subovate or elliptic to oblong, 3-27 x
1.5-16 cm, apex acuminate to acute, base round-
ed to obtuse, margin entire, + revolute towards
the base; upper surface glabrous, lower surface
densely to very sparsely appressed-puberulous
(on the main veins with hairs of different lengths)
or also densely to sparsely covered with white to
pale brown arachnoid hairs, disappearing or more
or less persistent on the main veins and the mar-
gin; lateral veins 4-11 pairs, straight or slightly
curved, basal pair branched or unbranched, in
 I
1cm

FIG. 37. Coussapoamicrocephala:ssp. microcephala:1, leafytwig with pistillateinflorescences(Maguire24588); 2, lea

(Gleason405).
Coussapoa
ovate to ellipticleaves reachingthe marginbelow
or sometimes at the middle of the lamina; in-
tercostal venation more or less prominent to
plane; petiole 0.5-5 cm long, appressed-puber-
ulous to hirtellousor also with arachnoidhairs;
stipules 1.2-5 cm long, yellowish puberulousto
subsericeous,often also with brownisharachnoid
hairs;terminalbuds slender.Staminate inflores-
cencesbranched;heads six to many, globose, ca.
2 mm diam.; common peduncle 1-4 cm long,
puberulous;perianth ca. 1 mm high, glabrous;
stamens two, far exceeding the perianth.Pistil-
late inflorescencesbranched, occasionally un-
branched;heads (1-)3-15, sometimes fused,glo-
bose, 2-5 mm, in fruit up to 10 mm diam.;
(common) peduncle 1-10 cm long, puberulous;
perianth ca. 1 mm high, glabrous. Interfloral
bracts (sub)spathulate,minutely puberulous at
the apex.
Distribution(Fig. 7). Common in Guyana,ap-
parentlyrarein Surinamand FrenchGuiana;in
forest, often along streams, at altitudes up to
900 m.
Specimensstudied.GUYANA. NorthwestDistrict,
Kaituma River, 27 Oct 1908 (a), Anderson 188 (K,
NY); CuyouniaCreek, - (8), Appun296 (K); Potaro
River, nr. KaieteurFalls, 18 Feb 1962 ($), Cowan et
al. 1868 (F, K, US), 18 Feb 1962 (9),Cowanet al. 1880
(F, K, US); NorthwestDistrict,Anabisi River, 15 Feb
1922 (9),De la Cruz1367 (GH, NY); upperRupununi
River,nr. Dadanawa,14 Jun 1922 (a),De la Cruz1501
(F, GH, MO, NY); betweenDemeraraRiver and Ber-
bice River, 5?50'N, 15-19 Jul 1922 (6), De la Cruz
1577 (F, GH, MO, NY); upper MazaruniRiver, ca.
60?10'W,22 Sep-6 Oct 1922 (8), De la Cruz2241 (F,
GH, MO, NY, US), De la Cruz 2387 (F, GH, MO,
NY, US);DemeraraRiver,Malali,30 Oct-5 Nov 1922
(6),De la Cruz2636 (F, GH, K, NY); upperMazaruni
River, Kamakusa,23-29 Nov 1922 (9), De la Cruz
2813 (F, GH, MO, NY); DemeraraRiver, Malali, 30
Oct-5 Nov 1922, De la Cruz 3070 (NY); Essequibo
River,WiniperuCreek,2 Nov 1939 (9),Fanshawe295
(FD 3031) (K, NY, U); MazaruniRiver, Kurupung
River, Sep 1925 (a), Gleason405 (K, NY); PotaroRiv-
er, Tumatumari,4-6 Jul 1921 (9), Gleason415 (GH,
K, NY); PotaroRiver, nr. KaieteurFalls, 17 Jan 1954
(a), Irwin 183 (US); MazaruniRiver, Sep 1880 (9),
Jenman 652 (K); upper Demerara River, nr. Great
Falls, Sep 1887 (2), Jenman 3995 (K, NY); Berbice
River, May 1889 (2), Jenman 4947 (K), without lo-
cality,Jun 1889 (e), Jenman 5315 (K);DemeraraRiv-
er, Mar 1898 (9), Jenman 7310 (K, S, U); Pomeroon
River, nr. Macasuma,Feb 1904 (e), Jenman 7933 (K);
KupurungRiver, 22 Nov 1922 (9), Leng 177 (NY);
WaineRiver, 10-15 km S of Kwabanna,10 Aug 1977
(9), Maas et al. 2478 (F, K, U); S of Timehri, Mr.
Thompson'sfarm, 17 Oct 1979 (a), Maas et al. 3622
73
(U); PakaraimaMts., Kamarang,8 Sep 1979 (6),Maas
et al. 4126 (U); Kaieteur Gorge, 14 May 1944 (6),
Maguireet al. 23449 (F, G, K, MO, NY, P, U, US);
DemeraraRiver, Comaka,Apr 1933 (6), Persaud222
(F);EssequiboRiver, MoraballiCreek,nr. Bartica,18
Sep 1929 (8),Sandwith290 (K, NY, RB), 19 Sep 1929
(2),Sandwith302 (K, U), 7 Oct 1929 (6),Sandwith394
(K, P, US); without locality, - (?), Schomburgks.n.
(F, G), - (6), Schomburgk 118 (B, K), - (9), Schom-
burgk167 (G, K, P); BerbiceRiver, 1837 (6), Schom-
burgk287 (BM, G, GH, K, L, OXF, P, U, type col-
lection of C. cuneata); Pomeroon River (?), - (6),
Schomburgk876 (by error276) (BM, F, FI, G, NY, P,
type collectionof C. microcephala),Schomburgk1366
(=?876) (B, K, type collection of C. fagifolia); Potaro
River, nr. Amatuk,26 Aug 1959 (8), Whitton179 (K).
SURINAM. Without locality, - (9), Andersons.n.
(BM);Tafelberg,1 Sep 1944 (9),Maguire25488 (A, F,
G, K, MO, NY, U, US).
FRENCHGUIANA.Roadto Brazil,km 6, at bridge
over Compt6 River, ca. 51 km S of Cayenne, 1 Jan
1977 (2), Mori 8864 (CAY, MO, NY, U).
Coussapoa microcephala is closely related to
C. argentea.Becauseof more or less distinctsim-
ilarities in the dimensions, shape, margin, and
venation of the leaves these two species together
with C. latifolia, C. viridifolia, C. parvifolia, C.
microcarpa, C. pachyphylla (and C. macerrima)
constitute a more or less distinct group within
the taxa having 2-staminate flowers.
27. Coussapoa napoensis Akkermans & C. C.
Berg,Proc. Kon. Ned. Akad. Wetensch., Ser.
C. 85(4): 457. 1982. Type. Ecuador. Napo:
Coca, 17 Jan 1973 (6), Lugo 2820 (holotype,
GB). Fig. 38.
Tree,up to 25 m tall, terrestrialor hemi-epi-
phytic. Leafytwigs 5-12 mm thick, sparselypu-
berulous,and mostly also with sparse,distinctly
longer, straight to curved, stiff hairs. Lamina co-
riaceous,broadlyovate to broadlyelliptic, 7-30
x 6-22 cm, apex obtuse, occasionally acumi-
nate, base subobtuseto rounded or to truncate,
margin entire to subcrenate; upper surface gla-
brous, lower surface on the main veins with ap-
pressedstraighthairsof differentlengthsand with
sparsearachnoidhairswhichlaterdisappear;lat-
eral veins (5-)7-12(-15) pairs, almost straight,
basal pair (sometimes faintly) branched, reaching
the margin below the middle of the lamina; in-
tercostal venation slightly prominent to almost
plane; petiole 3-10 cm long, with appressed
straight hairs of different lengths, glabrescent;
stipules 4-13 cm long, subvelutinous, also with
distinctly longer straight stiff hairs. Staminate
74 Flora Neotropica

~.I.

I cm.

FIG. 38. Coussapoanapoensis:1, leafy twig with staminateinflorescences(Lugo 2857).

Coussapoa 75

inflorescencesrepeatedlybranched;heads many, sparsely(to ratherdensely) covered with arach-

globose, 2-3 mm diam.; common peduncle2.5- noid hairs which are deciduous;lateralveins 8-
6 cm long, puberulousto hirtellous;perianthca. 15 pairs, mostly straight,basal pair (normally)
0.5-1 mm high, glabrous;stamen one, far ex- branched,reachingthe marginbelow the middle
ceeding the perianth. Pistillate inflorescences of the lamina;intercostalvenation(almost)plane;
branched; heads 3-9, subglobose, 6-10 mm petiole 3-13 cm long, 2-4 mm thick, densely
diam.; common peduncle 3-4 cm long, puber- brownishto white puberulousandwith distinctly
ulous to subhirsute;perianthca. 1 mm high, gla- longeryellowish appressedto patent hairs;stip-
brous. Interfloral bracts absent. ules 2.5-10 cm long, yellowishto brownishsub-
Distribution(Fig. 7). Ecuador(Napo) and Co- sericeous to hirsute or predominantlydensely
lombia (Putumayo);Amazonian forest. puberulous. Staminate inflorescence branched;
heads numerous,free,globose, ca. 2 mm diam.,
Specimensstudied.COLOMBIA. PUTUMAYO: Rio
often
Putumayo, PifiufiaNegro, 20 Nov 1940 (6), Cuatre- several solitaryflowerson the branchesof
casas10714(F, US). the inflorescence;commonpeduncle2-5 cm long,
ECUADOR.NAPO:Nr. Coca, 17 Jan 1973 (6),Lugo ratherdensely puberulousto hirtellous to sub-
2820(GB,typecollection), 22 Jan1973(6),Lugo2909 hirsute;perianthca. 1 mm
(GB);RioPayamino, nr.PayaminoCapihuara, 19Jan high, glabrousor mi-
1973(6),Lugo2857 (GB);Shushufindi, 4 Aug 1975 nutely puberulous; stamen one, farexceedingthe
(e), Little et al. 43 (Q);20 km W of Coca, 22-23 April perianth. Pistillate inflorescence branched; heads
1985 (v), Palacios et al. 6387 (BG). 2-7, free, globose, ca. 4-6, in fruit up to 20 mm
This species has many features in common diam.; common peduncle2-7 cm long, peduncle
with C. ovalifolia. It is distinct from the latter and branches 1-2 mm thick, ratherdensely pu-
berulous to hirtellous,apicesof the branchesoften
mainly in the longer stipules (of fertile twigs!), more or less
the smallernumberof lateralveins, and the larger broadened towardsthe heads;peri-
flowerheads of the staminateinflorescence.The anth ca. 1 mm high, sparselyminutelywhite pu-
two species could prove to be distinct only at the berulous to glabrous. Interfloralbractsabsent.
Distribution (Fig. 8). Amazon Basin (Brazil
subspecificlevel. and Peru), especially along the Amazon River;
mostly in riverside (varzea) forest.
28. CoussapoanitidaMiquel,Fl. bras.4(1): 113,
t. 44. 1853. Type. Brazil. Para: "Jaguary" Specimensstudied. COLOMBIA.AMAZONAS: Rio
(=Jaguarari?),- (e), Martius s.n. (lectotype, Caqueta, Puerto Cordoba, 13 Nov 1912 (8), Ducke
(HAMP)12238 (MG);TrapecioAmazonico,nr.mouth
M; isolectotypes,B?, BR, U). Fig. 39. of Rio Loretoyacu,26 Nov 1945 (2), Duque-Jaramillo
2207 (COL);Rio Loretoyacu,Oct 1945 (9), Schultes
CoussapoaintermediaMiquel, Fl. bras. 4(1): 133, t. 6726 (F, GH), Sep 1946 (2), Schulteset al. 8380 (F).
43. 1853. Type. Brazil.Withoutlocality("prov.Pa-
raensiet Rio Negro"),- (8),Martiuss.n. (holotype, PERU. LORETO:Rio Itaya, ca. 5 km above Iquitos,
6 Aug 1972 (a), Croat18850 (MO, U); Prov. Maynas,
M;isotype,B). Rio Ampiyacu,betweenPebas and mouth of Rio Ya-
CoussapoaschunkeiStandley,Publ. Field Mus. Bot.
22: 72. 1940. Type. Peru.Loreto:Rio Mazan,Gam- guasyacu,7 Nov 1977 (2), Gentryet al. 20381 (MO,
itanacocha,27 Jan 1935 (6), Schunke130 (holotype, U); Rio Ampiyacu,abovejunctionof Rio Yaguasyacu,
9 Nov 1977 (2), Gentryet al. 20495 (MO, U); Rio
F; isotypes, A, NY, US).
Yavari,opposite Paumari,23 Nov 1977 (a), Gentryet
Tree,hemi-epiphyticor terrestrial,up to 30 m al. 20802 (U); Rio Nanay, Mishana,halfwaybetween
tall. Leafytwigs 5-12 mm thick, densely brown- Iquitos and SantaMariade Nanay, 26 Feb 1979 (st),
ish puberulousto shortlyvelutinous,often partly Gentryet al. 25124 (MO, U); lower Rio Mom6n, nr.
Iquitos,8 Dec 1979 (9),J. Jones et al. 9712 (LAM,U),
(sub)hirsute.Lamina (sub)coriaceous,ovate (to 9 Dec 1979 (8), J. Jones et al. 9769 (LAM, U); Rio
elliptic), 8-30 x 4-20 cm, apex acute to obtuse, Ampiyacu, Puca Urquillo, 2 Apr 1977 (9), Plowman
base obtuse to rounded to truncateto cordate, et al. 6555 (F, GH, K, U); Rio Ampiyacu,Pebas, 16
Jul 1976 (a), Revilla 805 (MO); Rio Mom6n, Mo-
margin entire to subcrenate;upper surfacegla- noncillo, 17 Aug 1976 (9), Revilla 1099 (MO, U); Rio
brous, lower surface densely minutely puberu- Mom6n, above Bellavista, 19 Sep 1975 (9), Rimachi
lous in the areolesand on the reticulum,usually 1943 (NA); Rio Mazan,Gamitanacocha,27 Jan 1935
sparselypuberulousto hirtellouson the (parallel) (8), Schunke130 (A, F, NY, US, type collection of C.
tertiaryvenation, (appressed-)puberulous to stri- schunkei).
BRAZIL. Without locality (Prov. Paraensiet Rio
gose or to sparsely hirsute on the midrib and Negro), - (a),Martiuss.n. (B, M, type collectionof C.
lateral veins, sometimes the whole surface intermedia);- (a + 9, monoecious!),Poeppigs.n.(BR).
76 Flora Neotropica

FIG. 39. Coussapoanitida: 1, leafy twig with pistillateinflorescences(Schultes6726).

Coussapoa 77

ACRE: Rio Purus,nr. mouth of Rio Macaua,tributary 21 cm, apexacuteto obtuseto shortlyacuminate,
of Rio Iaco, 21 Aug 1933 (6), Krukoff5596 (A, BM, base cordate,marginentire or subcrenate;upper
F, G, K, LE, M, MO, NY, Q, S, U, US). AMAPA: Rio
surfaceglabrous,lower surfacedensely minutely
Jari, 2 km E of Arumanduba,27 Jun 1961 (2), W. A.
Egler et al. 46027A (IAN, MG, MO, NY, S, U). puberulousin the areoles and on the reticulum,
AMAZONAS: Rio Solim6es, B6ca de Manaquiri, 3 Oct (rather)sparselyhirtellousto puberulouson the
1973 (6), Berg et al. P17591 (F, M, MO, P, S, U), 3 other veins, on the main veins and marginalso
Oct 1973 (2), Berg et al. P17592 (F, MO, K, P, S, U);
Rio Japura,Mun. Maraa, Ilha dos Macacos, 30 Oct sparsewhite arachnoidhairs;lateralveins 9-11
1977 (8), Damiao 2503 (INPA);Rio Japura,Costa do pairs, slightly curved, basal pairs branched,
Jacitaria,8 Dec 1977 (8), Damido 2817 (INPA); Rio reachingthe marginbelow the middle of the lam-
Solimoes, Fonte Boa, 9 Oct 1945 (6), Ducke 1795 (A, ina, other lateral veins often poorly branched
F, K, MG, NY, R, RB, US); Rio Japura,19 Sep 1904 (furcate);intercostal venation very prominent;
(9), Ducke (HAMP)6782 (S, U); Rio Ia, 9 Sep 1906
(9), Ducke (HAMP)7715 (BM, G); Rio Tonantins,26 petiole 8-13 cm long, sparsely to densely mi-
Oct 1949 (6), Frogs25531 (IAN); Mun. Sao Paulo de nutelypuberulousand sparselycoveredwith dis-
Olivenca,nr. Palmares,11 Sep-26 Oct 1936 (6), Kru- tinctly longerstraightstiff hairs;stipules0.5-3.5
koff8401 (A, B, BM, F, G, K, LE, NY, P, S, U, US); cm long,brownishsubsericeousand with a dense
Rio Solimoes,Mamia,20 Jan 1924(8),Kuhlmann1178
(RB);Rio Javari,behindPalmeiras(armypost), 6 Aug covering of reddish-brownpluricellularhairs.
1973 (8), Lleras et al. P17197 (C, K, M, MO, NY, U, Staminateinflorescencesbranched;heads many,
US). PARA: Rio Tajapurir, Ant6niaLemos,IgarapePix- globose, ca. 4-6 mm diam.; common peduncle
una, 19 Jul 1948 (2), Black 48-2949 (IAN); Rio Ta- 1.5-3 cm long, puberulousto hirtellous, on the
japuri, Ilha Sao Sebastiao,Nazar6, 31 Jul 1948 (6), branchesto tomentellous, mixed with sparseto
Black48-3017 (IAN);Gurupa,27 Jan 1916 (6),Ducke
(HJBR) 13066 (=HAMP 16003) (BM, RB); Rio Ta-
densereddish-brownpluricellularhairs;perianth
japurfi,- (), Martiuss.n. (FI);"Jaguary"(=?Jaguar- ca. 1 mm high, densely minutely puberulous;
ari),Aug (9),Martiuss.n. (B, BR, M, U, lectotypecol- stamens three, exceedingthe perianth.Pistillate
lection), "InsulaeArchipelagiParaensis,"Aug (d and inflorescencesunbranched; heads ellipsoid to
2), Martiuss.n. (or 2673) (M, L, LE);Rio Jari,between
Monte Douradoand Caracura,18 Nov 1967 (9), Oliv- obovoid, ca. 20 x 10, in fruitup to 15 mm diam.;
eira 3627 (IAN, NY); Rio Jari,betweenMonte Dour- peduncle 1-2 cm long, reddish-brownpuberu-
ado and Patricia, 27 Mar 1970 (6), N. T. Silva 303 lous to hirtellous;perianthca. 2 mm high,dense-
(IAN). ly reddish-brownpuberulous.Interfloralbracts
Coussapoa nitida can easily be confused with subpeltate,often only a few or sometimesabsent.
C. tessmannii, due to the strong overall similar- Distribution(Fig. 8). Costa Rica (Alajuela);in
ities. However, C. nitida is clearly distinct from rain forest up to 800 m.
the latterin the absenceofinterfloralbracts.Oth- Specimensstudied.COSTARICA.ALAJUELA: Road
er differencesoccur in the indumentum, espe- to San CarlosValley, Buena Vista, 16 Apr 1903, (2),
cially in that of the petiole, and in the diameter Cook & Doyle 157 (US, type collection);canton San
of the peduncleof the pistillateinflorescence.C. Carlos,Villa Quesada,21 Feb 1939 (8),A. Smith 1632
HEREDIA: Nr. Puerto Viejo de Sarapiqui,3
nitida appearsto be related to C. cupularisand (F, MO).
Apr 1974 (2), Hartshorn1436 (U).
more remotely to C. orthoneuraand C. arach-
noidea. Although C. nymphaeifoliais reminiscentof
C. villosa in general appearanceand also to C.
in the nature of the pistillate inflores-
29. Coussapoa nymphaeifolia Standley, Proc. duquei
cence, these species are probablyunrelated.The
Biol. Soc. Wash. 37: 50. 1924; Burger, Field-
iana Bot. 40: 134, t. 22. 1977. Type. Costa occurrenceof 3-staminateflowersand othersim-
ilarities in leaf charactersand the staminatein-
Rica. Alajuela: Road to San Carlos valley,
Buena Vista, 16 Apr 1903 (2), Cook & Doyle
florescencesuggest a relationship with C. lon-
157 (holotype, US). gepedunculata.
Fig. 40.
Tree, hemi-epiphytic or terrestrial,ca. 20 m 30. CoussapoaoligocephalaDonnell Smith,Bot.
tall. Leafy twigs 5-17 mm thick, densely mi- Gaz. 40: 11. 1905. Type. Guatemala.Alta Ve-
nutely puberulousor also with dense reddish- rapaz: Cubilguitz, Apr 1904 (6), Von Tuerk-
brown pluricellularhairs. Lamina coriaceous, heim 8659 (=11.942) (holotype, US; isotype,
(broadly)elliptic to ovate, (9-)24-29 x (6-)18- B). Fig. 41.
 78 FloraNeotropica

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.?' ?

:~~~~~~~~~~~~'-
- ,??? I I 'I'~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~
.I . ,. "

?I r.~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~
?::~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~
'~~~~~~~~?: . .
.

P~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~

' ~ ~ ~ ~ ~ ~ ~ ~~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~~~~~~~ "

.? ~~~~~~
?~~~~~~~~~~~~
''~
? ~' 'y~ ~~~~~~~~~~~~~
~. . c , ,,.

~~~~~~~~~~~. .

F~~~~~~~~~~~~~~~~~~~~...:
?~~~~~~~~~~~~~~~~~~~~~~~~~~~ ?':. : ,::
..T!

?:".
?. jI .?~

-L
~~~~~~~~~~ I~~~~~~~~~~~~~~~~~~~~~ .....?:;:"li?b i"

~~~~~~~~~~~ ~ ??.. .*
~~~~~~~~~~:
......
';:-:F:..~,?; "'
~~;.? .~~~~~~~~~~~~~~' ?
'~~~~~~~~~~~~~~ ;?r
'..:i.- ; .

~~~~~~~~~~~~~~~~~~~~~~~~~~~'-?.-
I.':! ?-:'
, i'

i;, : ,...
?~~~~~~~~~~~~~~~~~~~~~~1
"
"'

FIG.40. Coussapoa
nymphaeifolia:
1, leafytwigwithstaminate
inflorescences,
2,leaf(A.Smith1632).
Coussapoa 79

I
1 cm

FIG. 41. Coussapoa oligocephala: 1, leafy twig with staminate inflorescences(Schipp 999); 2, pistillate
inflorescences(Contreras5787); 3, leaf (Contreras5787).
80
Shrubor tree,hemi-epiphyticor terrestrial,up
to 20 m tall. Leafy twigs 4-7 mm thick, ap-
pressed-puberulousto hirtellousto strigilloseor
to hirsute. Lamina (sub)coriaceous,elliptic to
oblong to (sub)ovateto (sub)obovate,4-21 x 1-
7 cm, apex shortlyacuminateto acute to obtuse,
base obtuse to subcordate,marginentire;upper
surfaceglabrous,lower surfaceminutely puber-
ulous to glabrousin the areoles sparselyhirtel-
lous on the smaller veins, the main veins gla-
brous, the whole surfacewith (persistent)white
arachnoidhairs;lateralveins (5-)8-12, straight,
basal pair unbranched,reachingthe marginbe-
low the middle of the lamina; intercostalvena-
tion prominent;petiole (1-)2-6 cm long, sparsely
to densely minute puberulousto hirtellous, or
hairsmixed with longerpatentto appressed,and
sometimes also arachnoidhairs;stipules 1-5 cm
long, (rather)sparselyappressed-puberulous(to
brownish subsericeous). Staminate inflores-
cences branched;heads 3-10, globose, ca. 4-6
diam.; common peduncle 1-5 cm long, with pu-
berulous to hirtellous to subtomentellous;peri-
anth ca. 1 mm high, minutely puberulous;sta-
mens three, far exceedingthe perianth.Pistillate
inflorescencesunbranchedor occasionallypoorly
branched;heads 1(-3), globose, ca. 5-8 mm, in
fruitup to 10 mm diam.; (common)peduncle 1-
5 cm long, ca. 1 mm thick, puberulousto hir-
tellous to subtomentellous;perianth ca. 1 mm
high, minutely puberulous;perianth orange or
yellow. Interfloralbracts(sub)spathulateto sub-
peltate, minutely puberulousat the apex.
Distribution(Fig. 8). NorthernCentralAmer-
ica (Guatemalaand Belize) to southernMexico
(Tabasco);in lowland (often riverine)forest.
Specimensstudied.MEXICO.TABASCO: La Palma,
Balancan,6 Jun 1939 (6), Matuda3300 (A, F, K, NA,
NY).
BELIZE. EL CAYODISTRICT:
Nr. El Cayo, 1933 (d),
Chanek220 (F, K);betweenMillionaroandCueras, zonas: Mun.
30 May 1973 (6), Dwyer 10799 (U); Vaca, 12 May
1938 (6), Gentle2602 (A, F, K, MO); nr. Millionaro,
30 May 1973 (6), Gentry7727 (MO, NY); Valentin,
Jun-Jul 1936 (6), Lundell6224 (F, GH, NY, S, US),
Jun-Jul 1936 (2), Lundell 6350 (C, F, GH, MO, NY,
S, US); Macaw Bank, 3 Apr 1969 (9), Proctor30273 CoussapoawilliamsiiCuatrecasas,FieldianaBot. 28(1):
(BM, MO). TOLEDO DISTRICT: Bolo Camp, 13 Apr
1944(6),Gentle4528(DUKE);HoneyCamp-Orange Isla Solitaria,betweenTamatamaand Esmeralda,7
Walk,Sep1928(st),Lundell2 (BM,F, K),24 Oct1929
(6),Lundell648 (F, K, MO, NY, S, US);Toledo, 1906-
1907 (6), Peck 497 (GH, K, NY, U); Punta Gorda, 7 CoussapoapranceiC. C. Berg,Acta Bot. Neerl. 27(1):
Aug 1932 (6), Schipp999 (A, BM, F, G, GH, K, MO,
NY, S).
Flora Neotropica
GUATEMALA. ALTAVERAPAZ:Sebol, 21 Apr 1964
(8), Contreras4449 (NY, S); El Cacao, Trece Aguas,
- (9), Cooks.n. (F, US); CerroChinaja,betweenFinca
Yalpemechand Chinnaja,1-2 Apr 1942 (st), Steyer-
mark45663 (F);Cubilgultz,Apr 1904 (6), VonTuerck-
heim 8659 (=11.942) (B, US, type collection).IZABAL:
Rio Dulce, road Seja-Cienaga,km 5, 11 Jul 1970 (2),
Contreras10195 (BM, DUKE, MO, U); Rio Dulce, 21
Jul 1936 (2), Hatch et al. s.n. (F); lower Rio Oscuro,
SW of LakeIzabal,27 Apr 1966 (9), G. C. Jones et al.
3152 (NY, US);LakeIzabal,JaguaCreek,31 May 1965
(9),SnedakerC39 (F), 28 May 1966 (9),SnedakerD88
(A, F); Rio Dulce, between Livingstoneand 6 mi up
river, 14 Apr 1940 (8),Steyermark39460 (F);Rio Frio,
17 Dec 1941(st),Steyermark39934 (F, US);Rio Dulce,
21 May 1939 (8), Wilson381 (F). PETEN:Vaxactun,
30 May 1931 (2), Bartlett 12356 (GH, NY, S, US);
Tikal, 31 Jul 1959 (2), Contreras61 (NY, S);LakeItza,
between Remate and San Andr6s, 18 Apr 1960 (9),
Contreras845 (F, NY, S, US); old roadto Machaquila,
Dolores, 27 Apr 1961 (8), Contreras2220 (NY, S); nr.
San Andr6s,Apr 1962 (2), Contreras3559 (F, NY, S);
Macanche,18 May 1966 (9),Contreras5787 (GH, US);
Cadenasroad, km 138, La Cumbre,25 Sep 1966 (9),
Contreras6231 (BM, MO, U); Lake Yaxha, archeo-
logical camp on north shore, 18 Jun 1973 (st), Croat
24663 (MO, NY); nr. Carmelita,25 Jun 1942 (st), F.
E. Egler 42-229 (F); 5 mi S of Tikal, 19 Jun 1973 (9),
Gentry8348 (U); nr. San Andr6s, 2 May 1933 (9),
Lundell3170 (F); La Libertad,31 May 1933 (9),Lun-
dell 3535 (F, S);Tikal, 30 Apr 1959 (6),Lundell15930
(S), 2 Jul 1959 (8), Lundell 16122 (F, S), 4 Jul 1959
(9), Lundell 16162 (G, NY, S, US), 27 Jul 1969 (6),
Ortiz 200 (BM, F); San Benito, nr. Playa Planca, 26
Apr 1970 (8), Ortiz1030 (BM, F, MO, NY); Tikal, 29
May 1971 (9), Ortiz 1810 (BM, F, US); CerroCeibal,
betweenRio SantaM6nicaand mouth of Rio Martin,
30 Apr 1942 (st), Steyermark46103 (F); Rio Macha-
quila, N of El Cambio, 25 Apr 1942 (6), Steyermark
45981 (F).
Coussapoa oligocephala is strongly reminis-
centof the AmazonianC. sprucei,especiallywhen
the specimens have an elliptic to subovate lam-
ina with subcordate base.
31. Coussapoa orthoneura Standley, Publ. Field
Mus. Bot. 17: 165. 1937. Type. Brazil. Ama-
Sao Paulo de OlivenSa, nr. Pal-
mares, 26 Oct-11 Dec 1936 (2), Krukoff8518
(holotype, NY; isotypes, A, B, BM, F, G, K,
LE, MO, SU, U, US). Fig. 42.
212. 1951. Type. Venezuela.Terr.Fed. Amazonas:
May 1942 (6),LI. Williams15240 (holotype,F; iso-
types, A, G, NY, RB, US, VEN).
11. 1978. Type. Brazil.Amazonas:Rio Cuieras,nr.
Jarada,17 Sep 1973 (9), Pranceet al. 18032 (holo-
Coussapoa
FIG. 42. Coussapoaorthoneura:1, leafy twig with pistillateinflorescences(Prance14092); 2, leafy twig with
staminateinflorescences(Schulteset al. 13774).
type,INPA;isotypes,C, F, K, M, MICH,MO,NY,
P, SU, U, US).
Tree,hemi-epiphyticor terrestrial,up to 35 m
tall. Leafy twigs 2-6 mm thick, glabrous or
sparsely, puberulous, sometimes hirtellous.
Lamina (sub)coriaceous,obovate to (broadly)el-
liptic to oblong, 2-15(-24) x 1-8(-11) cm, apex
81
1cm
acuteto shortlyacuminateor obtuse,base obtuse
to acute or truncate,marginentireor subcrenate;
uppersurfaceglabrous,lower surfaceglabrousor
sparselyto denselyappressed-puberulous,some-
times substrigoseor with white arachnoidhairs
later deciduous;lateralveins 2-7 pairs, straight
or slightlycurved,basal pairsunbranchedreach-
82 Flora Neotropica

ing the margin above to just below the middle PERU. AMAZONAS: Prov. Bagua, Dist. Cenepa,
of the lamina;intercostalvenation(almost)plane; Nueva Nazareth,nr. mouth of Rio Imaza,on Rio Ma-
raiion,26 Jan-18 Feb 1967 (2), Tillett671-33 (YEN).
petiole 1-7 cm long, glabrousor appressed-pu- LORETO: Prov. Maynas,Rio Nanay,nr. MoronCocha,
berulous to strigillose or to hirtellous; stipules N of Iquitos, 21 Mar 1977 (6), Gentryet al. 18528
(0.5-)1-7 cm long, sparselyto denselyappressed- (MO,U), Rio Nanay,nr. CaserioSantaClara,nr. Iqui-
puberulousor to mostly sparselyyellowish sub- tos, 19 Nov 1976 (2), Revilla 1847 (U).
sericeous (to subhirsuteor to subvillous), occa- BRAZIL.AMAZONAS: Manaus, Igarap6do Leao, 3
Sep 1957 (6), L. Coelho (INPA) 5728 (U); Nogueira-
sionally also with sparse to rather dense white Tef'e,26 Oct 1972 (2), Danta 12383 (INPA);Tonan-
arachnoid hairs. Staminate inflorescences tins, 7 Feb 1944 (9),Ducke 1528 (NY, US); Rio Purus,
branched;heads many, sometimes partlyfused, Quinta de Sao Christovao, 18 Jan 1873 (6), Glaziou
globose, ca. 1-6 mm diam.; common peduncle 10070a (P);Mun.SaoPaulode Olivenca,nr.Palmares,
1-3 cm long, with (dense)appressed-puberulous, 11 Sep-26 Oct 1936 (2), Krukoff8518 (A, BM, F, G,
K, LE,NY, S, U, US, typecollection),(6),Krukoff8587
sometimes also with reddish-brownpluricellular (F); Mun. Sao Paulo de Olivenca, Belem Creek, 26
hairs;perianthca. 1 mm high, glabrousor min- Oct-11 Dec 1936 (6), Krukoff8967 (A, B, BM, F, G,
utely puberulous; stamen one, just exceeding K, LE, MO, NY, S, U); Lago do Tefe, 10 km above
the perianth. Pistillate inflorescences branched; Tefe, 26 Jul 1971 (8), McDanielet al. 2666 (US); Rio
heads 2-10, sometimes partlyfused, 3-6 mm, in Ituxi, nr. B6ca do Curuquete,9 Jul 1971((), Pranceet
al. 14029 (F, GH, K, M, S, U); Rio Cuieras,just below
fruit up to 15 mm diam.; common peduncle 1- mouth of Rio Branchino,29 Sep 1971 (6), Prance et
3 cm long, (densely) appressed-puberulous, al. 15039 (F, GH, K, M, P, S, U); Rio Cuieras,nr.
sometimes partly hirtellous,apices of the head- Jarada,17 Sep 1973 (2), Pranceet al. 18032 (C, F, K,
bearingbranchesoften (especiallyin fruit)more M, MICH, MO, NY, P, S, U, US, type collection of
C. prancei);road Manaus-Itacoatiara,km 29, 14 Sep
or less swollen; perianthca. 1-2 mm high, gla- 1974 (2), Pranceet al. 22729 (INPA, U); Mun. Pura-
brous or sparselypapillate at the apex; fruiting quequara,road to Manaus, Panama, 6 Jun 1963 (2),
perianthyellow to red. Interfloralbractsabsent. W. A. Rodriqueset al. 5266 (U); Rio Castanho,road
Distribution (Fig. 8). Upper Amazon Basin to Careiro,11 Jul 1972 (8),M. Silva et al. 466 (U); Rio
Uaup6s, Taracua,Rio Tikie, 28 Jan-9 Feb 1948 (6),
(Brazil, Peru, Ecuador,and Venezuela);in up- Schulteset al. 9685 (GH, K);Rio Uaup6s,Panur6,Oct
landand riverine(varzea)forest;also knownfrom 1852-Jan 1853 (8), Spruce2498 (BM, G, GH, K, LE,
CentralColumbia (Santander,at ca. 1000 m). NY, OXF, P).
Specimens studied. COLOMBIA. AMAZONAS:Rio Coussapoa orthoneura is a variable species,
Igara-Parana,affluentof Rio Putumayo,La Chorrera, especially in the shape, dimensions, and second-
19 Dec 1973 (9), Gascheet al. 29 (U). GUAINiA: Rio venation of the lamina and also in the in-
Guainia,Manacal,21 Oct 1977 (2), Espina et al. 219 ary
(COL). SANTANDER: Rio Negro, El Play6n, 1000 m, - dumentum of the stipules. More or less distinct
(st), Franco& Garces419 (COL).VAUPES:Upper Rio regional forms, like the small-leaved form found
Vaup6s,La Jirisa,9 Jan 1944 (a), Guttierrezet al. 562 near Manaus, at the time described as C. prancei,
(COL,GH); upperRio Vaup6s,betweenLa Jarisaand have proved not to be
LasBocas, 12 Jan 1944 (Q),Guttierrezet al. 595 (COL); sufficiently different to be
Rio Apaporis,nr. Cachiverade Jirijirimo,8 Jul 1951 recognized as distinct (infraspecific) taxa. Cous-
(Y),Schultes et al. 12985 (F); Rio Apaporis,between sapoa arachnoidea appears to be closely related
Rio PacoaandRio Kananari,27 Aug 1951 (6),Schultes to C. orthoneura, which shows affinities espe-
et al. 13774 (COL,GH, U); Rio Apaporis,Raudalde
cially to C. cupularis and C. nitida. In the veg-
Jirijirimo,below mouth of Rio Kananari,14 Feb 1952 etative
(9), Schulteset al. 15324 (GH, US). parts the species can easily be confused
VENEZUELA. AMAZONAS: Rio Casiquiare, be- with C. latifolia, C. viridifolia and C. microceph-
tween Lago de Vasiva and Rio Pacimoni, 1853-1854 ala, but it is quite distinct from these three species
(a),Spruce3231 (B, C, G, K, P, S); Rio Ocama,above in the ebracteate inflorescence and the staminate
confluencewith Rio Orinoco,nr. OcamaMission, 23- flower.
24 May 1972 (8), Steyermark106167 (U, VEN); Isla
Solitaria,between Tamatamaand Esmeralda,7 May
1942 (6), Ll. Williams15240 (A, F, G, type collection 32. Coussapoa ovalifolia Trecul, Ann. Sci. Nat.
of C. williamsii);Esmeralda,14 May 1942 (a),LI. Wil- Bot. S6r. 3, 8: 95. 1847; Miquel, Fl. bras. 4(1):
liams 15363 (G, NY, US, VEN). 139. 1853; Macbride, Publ. Field Mus. Bot.
ECUADOR.NAPO:Misahualli,mouth of Rio Misa-
hualli, 3 Mar 1980 (st), Berg & Akkermans1111 (U); 13(2.2): 298. 1937. Type. Peru. Huanuco: Ma-
roadCoca-LagoAgrio, 9 km NE of Rio Coca, 20 Mar cora ?, - (2), Ruiz & Pavon s.n. (or 2) (holo-
1980 (st), Brandbygeet al. 30258 (AAU). type, FI; isotypes, F, US). Fig. 43.
Coussapoa 83

FIG. 43. Coussapoaovalifolia:1, leafy twig with pistillateinflorescences(Krukoff5657);2, staminateinflo-

rescences(Ule 9316).
84
CoussapoahirsutaTr6cul,Ann. Sci. Nat. Bot. Ser. 3,
8: 97. 1847;Macbride,Publ.FieldMus. Bot. 13(2.2):
297. 1937. Type. Peru. Huanuco: Macora, - (9),
Ruiz & Pavons.n. (or 5) (holotype,FI; isotype, B).
CoussapoaacutifoliaKlotzsch,Linnaea20: 529. 1847;
Macbride,Publ. Field Mus. Bot. 13(2.2):296. 1937.
Type.Peru.Huanuco:Cochero(fideMacbride,1937),
- (9),Ruiz & Pavons.n. (or 7) (holotype,B; isotype,
FI).
CoussapoapuberulaKlotzsch,Linnaea20: 529. 1847.
Type. Peru. Huinuco: "Chinchao-Pillao-Pozuzo-
Mucona,"- (9),Ruiz &Pavons.n. (or 6) (holotype,
B; isotype, F).
Coussapoasetosa Klotzsch, Linnaea 20: 528. 1847.
Type. Peru.Huinuco: Macora,- (9),Ruiz & Pavon
s.n. (or 5) (holotype,B; isotype, FI).
Tree, hemi-epiphytic, up to 20 m tall. Leafy
twigs 3-9 mm thick, glabrous or puberulous, often
also with distinctly longer straight stiff hairs.
Lamina coriaceous, ovate to subovate (or elliptic
to obovate), 7-32 x 2-16 cm, apex shortly acu-
minate to acute (to obtuse), base obtuse to round-
ed or acute to truncate, margin entire or sub-
crenate; upper surface glabrous, lower surface
glabrous or with the main veins having appressed
straight hairs of different lengths, often initially
also with sparse or sometimes rather dense white
arachnoid hairs on the main veins and along the
margin; lateral veins 7-21 pairs, almost straight
to curved, basal pair unbranched or in large leaves
poorly branched, reaching the margin far below
the middle of the lamina; intercostal venation
almost plane to slightly prominent; petiole 2-9
cm long, glabrous or with appressed hairs of dif-
ferent lengths; stipules 1-4 (in rapidly growing
shoots up to 13) cm long, subsericeous (to sub-
hirsute). Staminate inflorescences repeatedly
branched; heads numerous, globose, ca. 1(-2) mm
diam.; common peduncle 1.5-3 cm long puberu-
lous to hirtellous, occasionally also with sparse
arachnoid hairs; perianth 0.5-1 mm high, gla-
brous; stamen one, just exceeding the perianth.
Pistillate inflorescences branched or sometimes
unbranched; heads 1-5, subglobose, 5-8 mm, in
fruit up to 15 mm diam.; (common) peduncle 2-
6 cm long, puberulous to hirtellous; perianth ca.
1-2 mm high, glabrous; fruiting perianth brown-
ish. Interfloral bracts absent.
Distribution (Fig. 8). Upper Amazon Basin
(Bolivia, Peru, Brazil and Ecuador); in upland
and riverine forest, up to 900 m.
Specimensstudied.ECUADOR.NAPO:LagoAgrio,
3 Apr 1980 (9),Brandbygeet al. 30421 (AAU). PASTA-
ZA: Between Palora and Shell, 4 Mar 1980 (st), Berg
Flora Neotropica
& Akkermans1132 (U). ZAMORA-CHINCHIPE: Road
to Guayzimi, 27 km NNE of Zamora, 17 Sep 1975
(st), Little et al. 424 (COL,Q).
PERU. AMAZONAS: E ofHuampmai, 4 Jul 1974 (2),
Berlin1531 (MO,U). HuANuco:Prov. LeoncioPrado,
TingoMaria,7 Dec 1981 (9),Plowmanet al. 11179 (U);
Chinchoa-Pillao-Pozuzo-Mucuna,- (2), Ruiz & Pa-
von s.n. (or 6), (B, F, lectotypecollection of C. puber-
ula); Cochera, - (2), Ruiz & Pavon s.n. (or 7) (B, F,
FI, US, type collectionof C. acutifolia);Macora,- (2),
Ruiz & Pavons.n. (or 2) (F, FI, US, type collectionof
C. ovalifolia),- (2), Ruiz & Pavon s.n. (or 7) (B, FI,
type collection of C. hirsutaand C. setosa);Prov. Pa-
chitea,Dtto. Honoria,BosqueNac. Iparia,SchunkeV.
1162 (F, G, NY, US). JUNIN:Valley of Rio Paucar-
tambo, nr. Perenebridge, 19 Jun 1929 (a), Killipet al.
25341 (F, NY, S, US);Panchistrail,betweenSanNico-
las and Azupizfi, 6 Jul 1929 (2), Killip et al. 26095
(NY);La Merced,10-24 Aug 1928 (a),Macbride5594
(F).LORETO: Prov.Maynas,Rio Nanay,Mishana,half-
way between Iquitos and Santa Maria de Nanay, 26
Jul 1979 (st), Gentry25151 (MO, U); Rio Marafion,
Yanache,2 Mar 1924 (9), Kuhlmann1540 (U); Que-
bradaAucaya, 12 May 1973 (9), Rimachi 359 (NA);
Rio Maranion,betweenIquitos and Pongo de Manse-
riche, 1924 (9), Tessmann3922 (G); MatariCocha, 3
Oct 1968 (d), TorresM. 482 (K). MADREDE DioS:
Parque Nacional del Manu, Cocha Cashu, between
Punaguaand Tayakome, 17-24 Aug 1974 (2), Foster
et al. 3412 (F). SAN MARTIN: Eslab6n, between Sa-
posoa and Tingede Saposoa,Rio Saposoa,8 Sep 1948
(9), Ferreyra4813 (US); between Gramaloteand Sa-
posoa, 29 Apr 1962 (a), Woytkowski7306 (GH, US).
BRAZIL.ACRE: Mouth of Rio Macaua,tributaryof
Rio laco, 26 Aug 1933 (9), Krukoff5657 (A, BM, F,
G, K, LE, M, S, U); nr. Tarauaca,25 Sep 1970 (2),
Pranceet al. 7371 (U); Rio Acre, SeringalSao Fran-
cisco, Aug 1911 (9), Ule 9315 (G, K, L, MG), May
1911 (a), Ule 9316 (G, K, L, MG, US).
BOLIVIA.SANTA CRUZ:BuenaVista, Jun 1915 (8),
Steinbach1484 (GH, U).
Coussapoa ovalifolia is closely related to C.
napoensis. The collection Gentry & Mori 14011,
Panama, Cerro Tacaruna, premontane wet for-
est, 1300 m, 30 Jan 1975 (MO, U), probably
represents a separate taxon, possibly distinct at
the subspecific level from the Amazonian rep-
resentatives of C. ovalifolia. The collection is
similar in most characters to the other collections
of C. ovalifolia,but is distinctmainly in the short
(1-1.5 cm long) peduncle of the pistillate inflo-
rescence andthe branching of the basal lateral
veins-features which hardly justify recognition
as a distinct species. Staminate material is need-
ed to confirm the present supposition.
33. Coussapoa pachyphylla Akkermans & C. C.
Berg, Proc. Kon. Ned. Akad. Wetensch. Ser.
C, 85(4): 459. 1982. Type. Brazil. Bahia: Mun.
Coussapoa
lcm.
FIG. 44. Coussapoapachyphylla:1, leafy twig with pistillateinflorescences(Mori & Santos 11655).
SantaCruzde Cabralia,old roadto SantaCruz,
ca. 15 km NW of P6rto Seguro, 3 Apr 1979
(9), Mori & dos Santos 11655 (holotype, CE-
PEC;isotype, U). Fig. 44.
Shrub, often (hemi-)epiphytic,up to 5 m tall
(or taller?). Leafy twigs 4-8 mm thick, rather
densely to sparsely appressed-puberulous,also
withdistinctlylongerstraightstiffhairsandbrown
pluricellularhairs, glabrescent. Lamina coria-
ceous, ovate to elliptic (or obovate), 6-16(-24)
x 3.5-10(-19), apex acuteto subacuminate,base
acute, obtuse or sometimes subcordate,margin
entireto subcrenate;uppersurfaceglabrous,low-
er surfacedensely to sparselypuberulousto sub-
tomentose, initially often with sparse brownish
arachnoidhairs,especiallyalong the margin;lat-
eralveins 5-8 pairs,basalpairbranchedor some-
times (in small leaves) unbranched,reachingthe
85
margin below the middle of the lamina; inter-
costal venation more or less prominent; petiole
0.5-3(-5) cm long, densely to sparsely puberu-
lous, often also with distinctly longer straight stiff
hairs; stipules 1-2 cm long, densely covered with
short reddish-brown arachnoid hairs. Staminate
inflorescences branched; heads ca. 6-12, some-
times fused, globose, ca. 1-2 mm diam.; com-
mon peduncle 1-2 cm long, densely puberulous;
perianth ca. 0.5 mm high, nearly glabrous; sta-
mens two, as long as the perianth. Pistillate in-
florescences unbranched or few-branched; heads
1-3, sometimes partly fused, ca. 5-8 mm, in fruit
up to 13 mm diam.; (common) peduncle 2.5-4
cm long, 1-1.5 mm thick, densely puberulous;
perianth 1(-2) mm high, glabrous. Interfloral
bracts, few, subpeltate to spathulate.
Distribution (Fig. 8). Brazil (Bahia); in humid
forests in the coastal region.
86 Flora Neotropica

Specimensstudied.BRAZIL.BAHIA: Road Una-Il- Distribution (Fig. 8). Costa Rica, Panama, and
heus, ca. km 21, 2 Apr 1980 (st), Berg et al. 1153 in the Pacific coastal region of Colombia and
(CEPEC,K, NY, RB, U); Mun.SantaCruzde Cabralia, northern
old roadto SantaCruz,ca. 15 km NW of PortoSeguro, Ecuador; in forests up to 1250 m.
3 Apr 1979 (Q),Mori et al. 11655 (CEPEC,U, type
collection); Ilh6us-Repartimento(Fortuna), 21 Nov Specimens studied. COSTA RICA. ALAJUELA: Nr.
1944 (8), Vellozo 737 (R). San Ram6n, Cataratas(Los Angeles)de San Ram6n,
17 Apr 1935 (9),Brenes20542 (F, NY, type collection
This species appears to be related to C. latifolia of C. brenesii);SanCarlos,BuenaVista, FincaLa Con-
and allied taxa. stancia,3 Mar 1963 (2), Jimenez433 (F, MO);Rio San
Rafael,2 km W of La Marina,Llanurade San Carlos,
21 Feb 1966 (6),MolinaR. et al. 17694 (F);SanCarlos,
34. Coussapoa parviceps Standley, Proc. Biol. La Sucre, 1025 m, 2 Mar 1939 (6),A. Smith 1650 (A,
Soc. Wash. 37: 51. 1924; Burger, Fieldiana F, MO), 1 Mar 1939 (2), A. Smith 1939 (6),A. Smith
Bot. 40: 136, t. 22. 1977. Type. Costa Rica. 1650 (A, F, MO), 1 Mar 1939 (2), A. Smith 1663 (F).
CARTAGO:Rio Pejibaye,between Rio Taus and Que-
Puntarenas: Agua Buena valley, nr. Cafnas
brada Azul, 28 May 1972 (st), Lent 2538 (COL, F);
Gordas, 1100 m, Feb 1897 (9), Pittier 11166 Morariade Turrialba,21 Apr 1975 (st), Poveda 951
(holotype, US; isotype, MO). Fig. 45. (F);nr. Orosi, 30 Mar 1924 (6), Standley39792 (US).
PUNTARENAS:
CaniasGordas, Finca Loma Linda, 1140
CoussapoabrenesiiStandley,Publ.Field Mus. Bot. 18: m, 23 Feb 1973 (9),Busey628 (F, GH, MO);Rio Java,
383. 1937. Type. Costa Rica. Alajuela:nr. San Ra- Las CrucesBot. Garden, 16 Mar 1978 (9), Hartshorn
m6n, Cataratas(LosAngeles)de SanRam6n, 17 Apr 2157 (U); AguaBuenavalley, nr. CaniasGordas, 1100
1935 (Q),Brenes20542 (holotype,F; isotype, NY). m, Feb 1897 (2), Pittier 11166 (MO, US, type collec-
CoussapoaoligoneuraMildbraed,Notizbl. Bot. Gart. tion).
Berlin10: 415. 1928. Type. Colombia.Without lo- PANAMA. CHIRIQUi: BuricaPeninsula,Mellize, 6
cality, 19 Feb 1892 (9), Triana 866 (holotype, B; mi S of Puerto Armuelles, 5 Mar 1973 (st), Liesner
isotype, P). 417A (MO).COCLE: Road La Pineda-ElCope,nr. saw-
mill above El Cop6, 1000 m, 20 May 1978(6),Hammel
Tree, hemi-epiphytic or terrestrial, up to 30 m 2576 (U). PANAMA:Road El Llano-Carti, km 8-12,
tall. Leafy twigs 4-8 mm thick, glabrous or pu- 13 May 1973 (st),Nee et al. 8822 (MO,NY, U). VERA-
berulous. Lamina coriaceous, broadly ovate to GUAS: NW of SantaF6, 2.7 km from EscuelaAgricola
elliptic or suborbicular, sometimes tending to Alto de Piedra,30 Mar 1975 (2), Moriet al. 5360 (MO,
obovate, 7-24 x 4-18 cm, apex shortly acumi- NY).
COLOMBIA.Withoutlocality,19 Mar- (2), Triana
nate, sometimes almost rounded or obtuse, base 866 (B, P, type collection of C. oligoneura).NARIuO:
rounded to truncate or subcordate, margin en- Barbacoas,May 1853 (9), Triana 1866 (NY, photo-
tire; upper surface glabrous, lower surface gla- graph).VALLE: Rio Cajambre,Barco,21-30 Apr 1944
brous, occasionally sparsely puberulous to to- (2), Cuatrecasas17351 (F); Rio Anchicaya,Alto Yun-
da, 1000 m, May 1972 (2), Hilty M87 (MO).
mentellous; lateral veins 3-6 pairs, slightly ECUADOR. ESMERALDAS:Alto Tambo, 23 Sep 1965
curved, basal pair branched, reaching the margin (8), Little et al. 21136 (NY, Q, US).
at or above the middle of the lamina; intercostal
venation plane or slightly prominent; petiole Coussapoa parviceps differs from other Cous-
(0.5-)2-7 cm long, occasionally sparsely puber- sapoa species in the connate pistillate flowers,
ulous; stipules 1.5-5 cm long, glabrous or sparse- which is probably the reason why interfloral
ly puberulous. Staminate inflorescences branched; bracts, present in staminate inflorescences, are
heads many, globose, ca. 1 mm diam.; common lacking in the pistillate ones.
peduncle 1-5 cm long, glabrous or sparsely pu- Several characters, particularly as exhibited in
berulous; perianth ca. 0.5 mm high, glabrous; the collection Little & Dixon 21136, suggest
stamens three, just exceeding the perianth. Pis- taxonomic relationship to C. cinnamomifolia.
tillate inflorescences branched; heads 3-12, glo- The collections Liesner 417A and Nee et al.
bose, ca. 2-3 mm, in fruit up to 6 mm diam.; 8822, both from Panama, presumably represent
common peduncle 1-4 cm long, glabrous or juvenile specimens of the species. They are dis-
sparsely puberulous; flowers connate, perianth tinct in having narrower (oblong to subovate)
ca. 1 mm high, glabrous. Interfloral bracts lack- and larger (up to 50 cm long) laminae with up
ing or in the staminate heads a few, more or less to 18 pairs of lateral veins, the basal pair of which
distinctly spathulate (more or less cucullate), gla- reaches the margin below the middle of the lam-
brous or ciliolate bracts. ina; petiole up to 20 cm long; the densely puberu-
Coussapoa 87

Fa1to4 I"
FIG. 4.osppc 11 w .cm

FIG. 45. Coussapoaparviceps:1, leafy twig with pistillateinflorescences(Jimenez433).

lous twigs, petioles, and stipules (both sides!); 35. Coussapoa parvifolia Standley, Publ. Field
and in the presenceof white arachnoidhairs on Mus. Bot. 17: 165. 1937. Type. Brazil. Ama-
the margin and midrib on the lower surface of zonas: Mun. Sao Paulo de Olivenga, nr. Pal-
the lamina. The charactersof these specimens mares, 11 Sep-26 Oct 1936 (Q),Krukoff8273
arenot includedin the speciesdescription.Cous- (holotype, NY; isotypes, A, BM, F, G, K, LE,
sapoaparvicepsappearsto be the only Coussapoa MO, S, U, US). Figs. 46, 47.
species with pronounceddimorphism of leaves CousspoacornifoliaStandley,Publ.FieldMus.Bot.
dependingupon age. 17: 160. 1937.Type.Brazil.Amazonas:Mun.Sao
 FIG. 46. Coussapoaparvifolia:1, leafy twig with pistillateinflorescences(Krukoff8539);2, leafy twig with
staminateinflorescences(Pranceet al. 9054); 3, leaf (Pranceet al. 22999); 4, stipules (Krukoff8539).
Coussapoa 89

F: 1,ly tg wh 1 cm

FIG. 47. Coussapoaparvifolia:1, leafy twig with pistillateinflorescences(Krukof 8273).

Paulode Oliven:a,Bel6mCreek,26 Oct-11 Dec ger brownish patent hairs, soon disappearing;
1936(d),Krukoff8897(holotype,NY; isotypes,A, stipules 0.4-1(-1.3) cm long, appressed-puber-
F, G, K, LE,MO,P, S, U, US). ulous to subsericeous, terminal buds mostly
Coussapoamicrocephalasubsp. cornifoliaAkkermans
&C.C.Berg,Proc.Kon.Ned.Akad.Wetensch., Ser. swollen (containing young inflorescences).Sta-
C, 85(4): 457. 1982. minateinflorescencesrepeatedlybranched;heads
ca. 15-30, globose, ca. 2-3 mm diam.; common
Tree,up to to 20 m tall, hemi-epiphytic.Leafy peduncle 1.5-2 cm long, rathersparsely puber-
twigs 2-5 mm thick, rathersparselyappressed- ulous or partlyhirtellous;perianthca. 1 mm high,
puberulous,sometimes on the youngerpartsalso minutelyciliolate;stamenstwo, filamentsjust to
longer,brownish,patent(to appressed)hairs,soon far exceeding the perianth. Pistillate inflores-
disappearing.Lamina coriaceous to subcoria- cencesunbranchedor branchedheads (1-5, glo-
ceous, lanceolate to narrowlyovate, sometimes bose, 4-8 mm diam.; (common) peduncle 1-2.5
subobovate,4-14(-21) x 1.5-5.5(-11) cm, apex cm long, puberulous perianth ca. 1 mm high,
acuteto acuminate,base acute to obtuse, margin glabrous.Interfloralbracts(sub)spathulate,mi-
entire, towards the base + revolute; upper sur- nutely puberulous.
face glabrous, lower surface ? densely ap- Distribution(Fig. 8). Upper Amazon Basin, in
pressed-puberulousto glabrous; lateral veins Brazil (Amazonas), Peru (Loreto), Venezuela
(4-)7-11 pairs, basal pair unbranched,reaching (Amazonas),also in Extra-AmazonianColombia
the marginbelow or in subobovateleaves above (Antioquiaand Vichada);in non-inundatedfor-
the middle of the lamina; intercostal venation est.
slightly prominent; petiole 0.5-3(-4) cm long, Specimensstudied. COLOMBIA. ANTIOQuIA: Rd.
appressed-puberulousto glabrousalso with lon- Mutat&-Pavarand6, betweenhaciendasLa Esperanza
90
and Mocari, 150 m, 6 Mar 1987 (6), Fonnegraet al.
1798 (BG, HUA). VICHADA: Gaviotas,CainoAriba, 19
Feb 1973 (9), Cabrera2668 (COL).
VENEZUELA. AMAZONAS: Rio Cucucucuma,be-
low Raudal Pacure, 9 Nov 1950 (6), Maguire et al.
29408 (NY); Dpto. Atabapo, 15 km SE of San Fer-
nando de Atabapo, 10-16 Feb 1988 (d),Stergioset al.
11530 (BG).
BRAZIL.AMAZONAS: Mun. Sao Paulo de Olivenoa,
nr. Palmares, 11 Sep-26 Oct 1936 (2), Krukoff8273
(A, F, G, K, LE, MO, NY, P, S, U, US, type collection
of C. parvifolia),(2), Krukoff8539 (A, BM, F, G, LE,
MO, NY, P, S, U, US); Mun. Sao Paulo de Olivenca,
Belem Creek, 26 Oct-11 Dec 1936 (6), Krukoff8658
(A, BM, F, G, K, LE,MO,NY, P, S, U, US), (6),Krukoff
8897 (A, BM, F, G, K, LE,MO, NY, P, S, U, US, type
collectionof C. cornifolia);rd.Manaus-Itacoatiara,km
65-70, 23 Oct 1963 (8), Oliveira2764 (IAN); rd. Ma-
naus-Itacoatiara,km 61, Res. Flor. Walter Egler, 17
Dec 1968 (6), Prance et al. 9054 (M, MO, NY, P, S,
U); rd. Manaus-PortoVelho, km 510, 5 km N of Rio
Purusinho,17 Oct 1974 (6),Pranceet al. 22999 (INPA,
NY, U, US). PARA:Rd. Cuiaba-Santar6m,km 1417,
Rio Itapacura,25 Nov 1977 (6), Prance et al. 25748
(MO, RB, U).
This species belongs to a group with two sta-
mens and leaf margins which mostly are revolute
at the base, comprising e.g., C. microcarpa and
C. microcephala.
Prance et al. 9054 is aberrant in having
(sub)obovate leaves.
36. Coussapoa purpusii Standley, Publ. Field
Mus. Bot. 8: 6. 1930. Type. Mexico. Veracruz:
Zacualpan, Apr 1928 (6), Purpus 11162 (ho-
lotype, F). Fig. 48.
Tree, hemi-epiphytic or terrestrial, often (?)
deciduous, up to 20 m tall. Leafy twigs 3-6 mm
thick, glabrous or puberulous on the scars of the
stipules. Lamina chartaceous, ovate to elliptic or
to obtuse, occasionally obovate, 4-16 x 1-7 cm,
apex acute to acuminate to obtuse, base obtuse
sometimes subacute or subcordate, margin en-
tire; both surfaces glabrous; lateral veins 4-6 pairs,
curved, basal pair unbranched reaching the mar-
gin below, at, or slightly above the middle of the
lamina; intercostal venation plane; petiole 1-5
cm long, glabrous; stipules 1-3 cm long, glabrous
or sparsely to densely appressed-puberulous.
Staminate inflorescences branched; heads 2-6,
globose, ca. 5-8 mm diam.; common peduncle
2-4 cm long, glabrous; perianth ca. 1 mm high,
glabrous or sparsely minutely puberulous; sta-
mens three, far exceeding the perianth. Pistillate
inflorescences unbranched; heads globose, 4-6
mm, in fruit up to 10 mm diam.; peduncle 3-4
FloraNeotropica
cm long, glabrous; perianth 1-2 mm high, gla-
brous; fruiting perianth yellow or orange. Inter-
floral bracts (sub)spathulate, sparsely puberulous
at the apex.
Distribution (Fig. 8). Southern Mexico and
north-western Guatemala; in evergreen forest in
moist places (often along streams) or in semi-
deciduous forest; up to 1700 m.
Specimensstudied.MEXICO.CHIAPAS: OvandoMt.,
17 Dec 1936 (6), Matuda 446 (MO, NY, US); Santa
Rita, Mapastepec,Jan 1938 (6), Matuda 2020 (A, F,
K, NA, NY); between Guatimoc and Cacahuatan,3
Dec 1941 (9), Miranda 1737 (US); NW of San Fer-
nando,NW ofTuxtla, 1000m, 2 Apr 1950(9),Miranda
6164 (US); nr. El Ocote, 30 km NW of Ocozocuantla,
24 Mar 1951 (9),Miranda6273 (US), GUERERO: Distr.
Montesde Oca, San Antonio, 18 Apr 1938 (9),Hinton
14018 (K, NY, P, US). JALISCO: SierraMadre Occi-
dental, NW of San Sebastian,Las Mesitas,Arroyode
las Caaitas,1700 m, 15 Mar 1927 (6),Mexia 1872 (A,
BM, F, GH, MO, NY, US). VERACRUZ: Paso del In-
genio, Jun 1971 (9), Calzada325 (U); withoutlocality,
1930 (st), Purpuss.n. (B, F, K); Zacualpan,Mar 1908
(6),Purpus5996 (BM,F, GH, MO,NY, US), Mar 1930
(8), Purpus11161 (K), Apr 1928 (6),Purpus11162 (F,
type collection), Jun 1930 (9), Purpus 11162 (A, K,
MO), Sep 1930 (9), Purpus 11162 (C), Apr 1931 (9),
Purpus11162 (C), May 1933 (6 and 9), Purpus11162
(A), May 1934 (6 and 9), Purpus 11162 (K, US), Sep
1929 (8), Purpus11182 (A).
GUATEMALA. QUEZALTENANGO: Colomba, ca.
1000 m, 28 Dec 1934 (6), Skutch 2023 (BM, F, GH,
NY). SANMARCOS: FincaEl Porvenir,on PotreroMa-
tasan, Rio Cabuis,Volcan Tajumulco, 1000-1300 m,
12 Mar 1940 (9), Steyermark37602 (F), 25 Mar 1943
(st), Steyermark52334 (F).
The species appears to be related to C. parvi-
ceps.
37. CoussapoascabraAkkermans& Berg,Proc.
Kon. Ned. Akad.Wetensch.Ser. C 85(4): 460.
1982. Type. Brazil.Rondonia:Rio Ouro Pre-
to, affluentof Rio PacaasNovos, 18 Sep 1923
(9), Kuhlmann 470 (HJBR 19836) (holotype,
RB; isotypes, RB, U). Fig. 49.
Tree. Leafy twigs 2-5 mm thick, puberulous
to subhirsute; internodes solid. Lamina charta-
ceous, elliptic to ovate or obovate, 4-10 x 2.5-
7 cm, apex acute to shortly acuminate, base ob-
tuse to acute, margin entire; upper surface, sca-
brous, lower surface, scabridulous, sparsely to
rather densely puberulous to hirtellous; lateral
veins 6-13 pairs, straightto curved, basal pair
unbranched,reachingthe margin far below the
middle of the lamina; intercostal venation slight-
ly prominent; petiole 1-3 cm long, densely pu-
 FIG. 48. Coussapoapurpusii:1, leafy twig with pistillate inflorescences(Hinton 14018); 2, leafy twig with
staminateinflorescences(Skutch2023).
92 Flora Neotropica

1cm

FIG. 49. Coussapoascabra: 1, leafy twig with pistillateinflorescences(Kuhlmann470).

berulousto hirtellous;stipules 0.5-0.7 cm long, lamina show similarities to those found in C.

subhirsute. Staminate inflorescencesunknown. parvifoliaand C. macerrima.
Pistillate inflorescencesunbranched or poorly
branched;heads 1-5, globose, 2-4 mm diam., in 38.
fruit up to 6 mm; (common) peduncle 1-3 cm CoussapoaspruceiMildbraed,Notizbl. Bot.
Gart.Berlin10:415.1928. Type. Brazil.Ama-
long, puberulous;perianth ca. 1 mm high, gla- zonas: "Rio Negra," probably Rio Taruma,
brous.Interfloralbracts(sub)spathulate,minute-
Dec 1854 (9), Spruce 3782 (holotype, B; iso-
ly puberulousat the apex. BM, BR, C, G, GH, K, LE, NY, OXF,
Distribution(Fig. 8). Brazil,R6ndonia.Known types,
P). Fig. 50.
only from the type collection.
The position of this species,whose leaves have Tree or shrub, mostly hemi-epiphytic, up to
a characteristicscabrousand chartaceouslami- 30 m tall. Leafytwigs 3-7 mm thick, puberulous
na, is unclear. The shape and venation of the to hirtellousto brownishhirsute,sometimes also
Coussapoa
2 1cm
FIG. 50. Coussapoasprucei: 1, leafy twig with pistillate inflorescences(Spruce3782); 2, leafy twig with
staminateinflorescences(Pranceet al. 14976).
with white arachnoidhairs. Lamina coriaceous,
oblong to elliptic, subovate or to subovate, 5-18
x 3-8 cm, apex shortlyacuminateto acute, base
subcordateto rounded,marginentire;uppersur-
face glabrous, lower surface puberulous in the
areoles,hirtellous(to tomentellous)on the small-
er veins, appressed-puberulous on the mainveins,
sometimes also with a few distinctly longer ap-
pressed hairs, the whole surface covered with
rather dense to sparse (sub)persistent white
arachnoidhairs;lateralveins 5-10 pairs, slightly
curved, basal pair unbranched or inconspic-
uously branched,reachingthe marginbelow the
middle of the lamina;intercostalvenation (very)
prominent;petiole 1-4 cm long,puberulous,often
also having distinctly longer straight hairs and
white arachnoid hairs; stipules 0.5-1.5(-2) cm
long, brownishsericeousto subhirsute,terminal
buds often more or less swollen. Staminate in-
florescencesbranched;heads many, globose, ca.
3-4 mm diam.; common peduncle (2-)4-8 cm
long,puberulousto hirtellous;perianthca. 1 mm
high, glabrous;stamens three, far exceeding the
perianth.Pistillateinflorescencesbranched;heads
2-5, globose, ca. 3-5 mm in fruit up to 8 mm
diam.;common peduncle2-8 cm long, ca. 1 mm
thick, puberulous to hirtellous; perianth ca. 1
mm high, glabrous.Interfloralbractsspathulate
to subpeltate,puberulous,mostly conspicuous.
Distribution(Fig. 8). Brazil, Amazon basin,
nearManaus,and in the basin of the Rio Trom-
betas;in forest of terrafirme.
93
Specimensstudied. BRAZIL. AMAZONAS: Nr. Ma-
naus, Rio Taruma, 27 Oct 1943 (6), Ducke 1354 (A,
F, NY, R, RB, US); nr. Manaus,C6loniaJoao Alfredo,
27 Sep 1946 (8), Ducke 1999 (A, NY, R, RB, U); nr.
Manaus,Rio Taruma,CachoeiraGrande,17 Oct 1929
(9), Ducke(HJBR)23609 (RB);Manaus, 17 Oct 1929
(9), Killip & Smith 30179 (NY); Rio Cuieras, 2 km
belowmouthofRio Brancinho,27 Sep 1971 (6),Prance
et al. 14976 (K, M, MO, NY, S, U), 12 Sep 1973 (8),
Pranceet al. 17847 (F, K, M, MO, NY, P, S, U); "Rio
Negro," possibly Rio Taruma, Dec 1854 (9), Spruce
3782 (B, BM, BR, C, G, GH, K, LE, NY, OXF, P,
type collection).PAIA:Rio Mapuera,11 Dec 1907 (2),
Ducke (HAMP)9096 (MG, US).
Coussapoa sprucei appears to be closely related
to C. leprieurii. It differs distinctly from the latter
in the smooth and glabrous upper surface of the
lamina. Other differences are of little signifi-
cance. C. sprucei also resembles the Central
American C. oligocephala, from which it differs
mainly in the length of the stipules and to a lesser
extent in the nature of the indumentum. This
group of three species is probably related to the
large-leaved taxa C. nymphaeifolia and C. lon-
gepedunculata, judging from the similarities in
indumentum, leaf venation, and number of sta-
mens.
39. Coussapoa tessmannii Mildbraed, Notizbl.
Bot. Gart. Berlin 10: 413. 1928; Macbride,
Publ. Field Mus. Bot. 13(2.2): 298. 1937. Type.
Peru. Loreto: Mouth of Rio Santiago, 3 Dec
1924 (p), Tessmann 4673 (holotype, B; iso-
types, G, S). Fig. 51.
94 Flora Neotropica

FIG. 51. Coussapoa tessmannii: 1, leafy twig with pistillate inflorescences (Krukoff 8994); 2, staminate
inflorescences (Krukoff 1187).
Coussapoa
Tree or shrub,(mostly) hemi-epiphytic,up to
20 m tall. Leafy twigs 5-13 mm, brown puber-
ulous to strigilloseto hirtellousto strigose or to
(sub)velutinous,often also with a dense covering
of (long) dark brown pluricellular hairs and
sometimeswhitisharachnoidhairsas well. Lam-
ina coriaceous,ovate to elliptic, 13-32 x 11-20
cm, apex subacute to obtuse to rounded, base
obtuse to subcordate,marginentire to subcren-
ate;uppersurfaceglabrous,lowersurfacedensely
minutely puberulousin the areoles and on the
reticulum,to (sparsely)hirtellouson the (paral-
lel) tertiary venation, appressed-puberulousto
strig(ill)oseon the midrib and lateral veins, the
whole surface with rather sparse to dense sub-
persistentwhite to pale brown arachnoidhairs;
lateralveins 9-16 pairs,straight,main basal pair
branched,sometimes faintly so (in small leaves
sometimes unbranched),reachingthe marginfar
below or sometimesjust below the middle of the
lamina; intercostal venation very to slightly
prominent;petiole 3.5-9 cm long, 2-5 mm thick,
brown(ish)puberulousto hirtellousto subvelu-
tinous, often also with dark brown pluricellular
hairsand sometimes arachnoidhairs;stipules2-
3(-9) cm long, yellowish to brown strigose to
subhirsuteor also with darkbrown pluricellular
hairs. Staminate inflorescencesbranched;heads
many, crowded at the end of the branches,
(sub)globose,2-4 mm diam., often fused; com-
mon peduncle 1-3 cm long, brown puberulous
to hirtellousto short-velutinousor also (densely)
coveredwith darkbrownpluricellularhairs;peri-
anth ca. 1 mm high, puberulous;stamen one,
just exceeding the perianth. Pistillate inflores-
cences branched;heads 2-7, (sub)globose,5-10
mm, in fruit up to 15 mm diam.; common pe-
duncle 2-3 cm long, peduncle and branches2-
4 mm thick, appressedpuberulousto hirtellous
to short-velutinousor also with darkbrownplur-
icellularhairs;perianthca. 1 mm high, (almost)
glabrous,sometimesmuriculate.Interfloralbracts
(sub)spathulateto subpeltate,puberulousat the
apex; extrafloral bracts often present on the
branchesof staminate inflorescences.
Distribution(Fig. 9). Amazon Basin (Brazil,
Peru, and Colombia); in upland and riverside
forest.
Specimens examined. COLOMBIA. VAUPES:
Rio
Papuri,nr. MontfortMission,3 Sep 1943(s), P. H.
Allen3103(COL).
95
Prov. Maynas,Rio Yaguasyuca,af-
PERU. LORETO:
nr.BrilloNuevo,2 Mar1978
fluentof RioAmpiyacu,
(2), Balick et al. 1021 (MO, U); road Zungarocha-
PuertoAlmendra,3 Dec 1964 (2), Dodson et al. 3017
(F);RioYavari,behindAngamoGarrison, 5 Oct1973
(a),Lleraset al. P17178(F, K, NY, P, S, U); Puerto
Almendra,nr. Rio Nanay,23 Feb 1977 (a),Revilla
2378 (U); mouthof Rio Santiago,3 Dec 1924 (2),
Tessmann4673(B, G, S, typecollection).
BRAZIL.AMAZONAS: Rio Negro, Uaup6s,Jauarite,
23 Aug 1945 (9), Froes 21264 (NY); Mun. Sao Paulo
de Olivenca,BelemCreek,26 Oct-11Dec 1936(2),
Krukoff8994(A, G, K, LE,NY, P, S, U); roadMa-
naus-PortoVelho,km 379, Rio Jutai,12 Oct 1974
(9), Prance et al. 22836 (MG, NY, U); Rio Javari, 7
hours above Paumari, 16 Oct 1976 (2), Prance et al.
23846 (NY, U). PARA: Cupary,Sep 1931 (6), da Costa
90 (IAN);upperRio Cupary,Sep 1931 (a),Krukoff
1147(A, BM,G, K, NY, P, S, U), Krukoff1187(A,
BM,G, K, MO,NY, S, U); roadSantarem-Palhao,
km 70, 15 Sep 1969(8),M. Silvaet al. 2591 (F, K,
MG, NY, S, U, US). RONDONIA: Porto Velho, 15 Sep
1975 (a), Mota et al. 154 (U).
In the vegetative parts C. tessmannii shows
similaritiesto C. villosa, but it is distinct from
the latter in having a ramified,and thus pluri-
capitate,pistillate inflorescenceand a unistami-
nate flower. C. tessmannii shows distinct affin-
ities with C. cinnamomea,e.g., in the relatively
thick peduncleand branchesof the pistillatein-
florescence,and it can be easily confused with
C. nitida. Several collections made in the Rio
Tapajos area (da Costa 90, Krukoff1147 and
1187, and M. Silva et al. 2591) differ from the
other collections in the presenceof a dense cov-
ering of darkbrown moniliformpluricellular(=
granular)hairs on the leafy twig, petiole, and
inflorescence.Moreover,the lamina tends to be
elliptic rather than ovate (as is normal in the
species).In the vegetativeparts,these collections
show strikingsimilaritiesto the aberrantpistil-
late collection (Pranceet al. 26402) assigned to
C. villosa (see p. 104).
40. Coussapoa trinervia Spruce ex Mildbraed,
Notizbl. Bot. Gart.Berlin10:416. 1928. Type.
Venezuela. Terr. Fed. Amazonas: Rio Casi-
quiare, between Lago de Vasiva and Rio Pa-
cimoni, 1853-1854 (2),Spruce3464 (holotype,
B;isotypes,BM, BR, G, GH, K, LE,NY, OXF,
P). Fig. 52.
Tree,terrestrial,up to 20 m tall, mostly with
stilt roots. Leafy twigs 3-8 mm thick, glabrous.
Lamina coriaceous, subobovate to obovate to
 1cm

2
FIG. 52. Coussapoatrinervia:1, leafy twig with pistillateinflorescences(Berg et al. P18453); 2, leafy twig
with staminateinfiorescences(Schunke371).
 Coussapoa
oblong to elliptic, 4-14 x 1.5-7 cm, apex obtuse
to shortly acuminate, base (sub)acute to cuneate,
margin entire; both surfaces glabrous; trinervate,
the single pair of lateral veins, unbranched,
reaching the margin near or at the apex of the
lamina; intercostal venation prominant; petiole
1-4 cm long, glabrous; stipules 0.5-1.5 cm long,
glabrous or occasionally appressed-puberulous.
Staminate inflorescences branched; heads 3-9,
often partly fused, globose, 2-5 mm diam.; com-
mon peduncle 1-3 cm long, glabrous or sparsely
puberulous; perianth ca. 1 mm high, glabrous;
stamen one, just exceeding the perianth. Pistil-
late inflorescences unbranched; heads globose, ca.
5-7 mm, in fruit up to 10 mm diam.; peduncle
1-3 cm long, glabrous or sparsely puberulous;
perianth ca. 1 mm high, glabrous. Interfloral
bracts subspathulate to peltate, sparsely puber-
ulous at the apex.
Distribution (Fig. 9). Amazon Basin (Brazil,
Peru, Ecuador, Colombia, and Venezuela); the
main area apparently in the north-western part
of Amazonia-elsewhere in smaller, more re-
stricted, areas; on periodically inundated banks
(always?) of black-water rivers.
Specimens studied. COLOMBIA. CAQUETA:
Rio Ca-
queta,Solano, 8 km SE ofTres Esquinas,below mouth
of Rio Orteguaza,7 Mar 1945 (st), Little et al. 9629
(COL); Rio Apaporis, between Rio Pacoa and Rio
Kananari,Soratama,16 Aug 1961 (d), Schulteset al.
13589 (GH, U).
VENEZUELA. AMAZONAS: Rio Manariche,Kalo-
hi-teri, - (2), Lizot 83 (VEN);Rio Yatua,just above
PiedraArauicaua,28 Sep 1957(2), Maguireet al. 41620
(K), Esmeralda-Ocana,Mavaca,23 Apr 1968 (2), Me-
dina 397 (VEN);Rio Casiquiare,betweenLagode Va-
siva and Rio Pacimoni, 1853-1854 (9), Spruce 3464
(B, BM, BR, G, GH, K, LE, NY, OXF, P, type collec-
tion); Rio Yatua,just above PiedraArauicaua,16 Jul
1959 (9), Wurdacket al. 43485 (K).
ECUADOR. NAPO:Rio Cuyabeno, ca. 0?10'S,
76?55'W, 16 Feb 1980 (d), Berg & Akkermans1038
(U), 17 Feb 1980 (2), Berg & Akkermans1048 (U), 18
Feb 1980 (2), Berg & Akkermans1058 (U); Laguna
Cuyabeno,7 Aug 1980(9), Jaramilloet al. 2888 (AAU),
PASTAZA:Rio Capihuari,23 Jul 1980 (st), Ollgaardet
al. 35068 (AAU).
PERU. LORETO: QuebradaCuninico,7 Jul 1972 (2),
Croat17784 (F, NY); Rio TacshaCurary,18 Sep 1972
(a), Croat 20411 (C, F, GH, L, MO, NA, NY, P, S);
Rio Yavari, Emilia, above Atalaiadel Norte, 22 Nov
1977 (2), Gentryet al. 20766 (MO, U); Prov. Maynas,
Rio Mom6n, tributaryof Rio Nanay, nr. Iquitos, 9
Dec 1979 (2), J. Jones et al. 9788 (LAM, U); Rio Na-
nay, QuebradaMorropon,26 Jun 1972 (2), McDaniel
16344 (NA); Rio Nanay, Mapa Cocha, 17 Mar 1976
(a),McDanielet al. 20555 (NA);Rio Nanay, Quebrada
Morropon,31 May 1976 (2), Rimachi 2297 (F, MO),
97
Rio Mazan,Gamitanococha,14 Mar 1935(6),Schunke
371 (A, F, NA, NY).
BRAZIL. AMAZONAS:Rio Negro Basin, Tapuru-
quara, 21 Oct 1971 (9), Prance et al. 15764 (F, GH,
K, NY, S, U); Rio Uaup6s,Panur6,Oct 1852-Jan 1853
(Y),Spruce2616 (B, BM, BR, C, G, GH, K, LE, NY,
OXF, P). MARANHXO: Turiaga,Astonas, 6 Dec 1978
(2),Rosa et al. 2874 (U). MATOGROSSO: Rio Aripuafia,
nr. HumboldtCenter, 10?12'S,59?21'W,10 Oct 1973
(a),Berg et al. P18409 (F, K, MO, NY, P, U, US), 12
Oct 1973 (2), Berg et al. P18453 (K, MO, NY, S, U,
US); Rio Juruena,Cachoeirade Sao Joao da Barra,3
Jun 1977 (a), Rosa et al. 2053 (NY); Rio Juruena,
IgarapeChuini, 15 Jul 1977 (2),M. G. Silva et al. 3345
(U). PARA:Mun. Itapiranga,Rio Uamata, nr. Igarap6
Santa Luzia, 16 Aug 1974 (2), Cid et al. 435 (U), nr.
Cachoeirado Tucumari, 19 Aug 1974 (Y),Cid et al.
511(U);Belem,betweenValde CaesandSaoJoaquin,
7 Nov 1922 (2), Ducke (HJBR) 18455 (RB); Belem,
Sep-Oct1961(st),Pires51736(NY).
This remarkablespecies, probably the only
truly terrestrialspecies, resembles terrestrialfig
trees (like F. trigonaL. f.) in habit and has me-
lastomataceousleaves. It appearsto be confined
to periodicallyinundated banks of black-water
rivers;this may explainthe disjunctdistribution.
Coussapoa trinervia appears to be closely related
to C. angustifolia, as the two species have many
characters in common. The difference in the
numberof stamenssuggeststhat C. trinerviaand
C. cinnamomifolia are not related, in spite of
similarities in the leaf venation.
41. Coussapoa vannifolia Cuatrecasas, Caldasia
7: 296. 1956. Type. Colombia. Valle: Rio An-
chicayf, between Sabuletas and Quebrada Ta-
tabro, 28-29 Sep 1946 (9), Cuatrecasas 22048
(holotype, F). Fig. 53.
CoussapoarotundaLittle, Phytologia 18: 196. 1969.
Type. Ecuador.Esmeraldas:Junction of Rio Hoja
Blancaand Rio Hualpi, 50 km S of Borb6n, 14 Sep
1965 (6), Little & Dixon 21056 (holotype,US; iso-
types, NY, Q).
Tree, terrestrial or hemi-epiphytic, up to 20 m
tall. Leafy twigs 5-9 mm thick, white to brown
hirtellous to hirsute. Lamina coriaceous, broadly
ovate to cordiform to subreniform or to subor-
bicular, 4-22 x 4-25 cm, apex rounded to ob-
tuse, base cordate, occasionally obtuse, margin
entireor subcrenate;uppersurfaceglabrous,low-
er surfaceratherdenselywhite puberulousto hir-
tellous on the main veins, otherwiseglabrousor
sparsely puberulous to hirtellous on the tertiary
(parallel) venation; lateral veins 5-9 pairs, in-
cluding 2-4 pairs of basal veins, basal veins
 FG53 Cosaovanfla1leftwgwtstmntifoecne Apud141'
....
I
cm
.......

FIG. 53. Coussapoa vannifolia: 1, leafy twig with staminate infiorescences (Asplund 1640
Coussapoa 99

branched,the upperpair of basal veins reaching CoussapoavellereaKlotzsch, Linnaea20: 527. 1847;

the marginat or above the middle of the lamina, Miquel, Fl. bras. 4(1): 139. 1853; Macbride,Publ.
Field Mus. Bot. 13(2.2):299. 1937. Syntypes.Peru,
the otherlateralveins oftenpoorlybranched(fur- Huanuco:Macora, - (6 and 2), Ruiz & Pavon s.n.
cate);intercostalvenation almost plane;petioles (or 3) (holotype (2), B; isotype (2), FI; isotypes (8),
1-6 cm long, (rather)densely white to brownish BR, G, OXF, US).
hirtellousto hirsute;stipules2-8 cm long, dense- Coussapoamartiana(Miquel,Fl. bras.4(1): 132, t. 42.
1853. Type. Brazil.Withoutlocality("prov.Paraen-
ly puberulousto hirtellousto strigillose,mostly si et Rio Negro"),- (2), Martiuss.n. (holotype,M).
also with brown pluricellularhairs. Staminate CoussapoasubincanaMiquel,Fl. bras.4(1): 134, t. 45.
inflorescencesbranched;headsmany,globose,ca. 1853. Type. Brazil.Withoutlocality, - (6),Martius
1.2 mm diam.; common peduncle 1-2 cm long, s.n. (holotype,M; isotypes, BR, M, U).
hirtellous to subtomentose;perianth ca. 1 mm CoussapoapanamensisPittier,Contr.U.S. Natl. Herb.
18: 226. 1917; Woodson & Schery,Ann. Missouri
high, apex minutely puberulous;stamens three, Bot. Gard. 47: 169, t. 59. 1960; Burger,Fieldiana
just exceeding the perianth. Pistillate inflores- Bot. 40:135, t. 22. 1977. Type. Panama.Col6n:Rio
cences branched;heads 3-5, free or sometimes Fat6, Jul 1911 (9), Pittier 3892 (holotype,US; iso-
partlyfused, (sub)globose,6-9 mm diam.; com- types, B, GH, NY).
mon peduncle1.5-2.5 cm long, hirtellousto sub- Coussapoadonnell-smithiiMildbraed,Notizbl. Bot.
Gart. Berlin 10: 414. 1928. Type. Costa Rica. Car-
tomentose; perianth ca. 1 mm high, papillate. tago: Rio Turrialba,Mar 1894 (2), Donnell Smith
Interfloralbractspresent,subpeltate,at the apex 4826 (holotype,US; isotypes, GH, K, US).
minutely puberulous,in pistillate inflorescences CoussapoagrandicepsKillip in Macbride,Publ. Field
often only a few. Mus. Bot. 13(2.2): 297. 1937. Type. Peru. Loreto:
Distribution(Fig. 9). Ecuadorand Colombia, LowerRio Huallaga,Yan6n, 5-11 Sep 1929 (2), Kil-
lip & Smith 29246 (holotype,US).
Pacific coastal region;in lowland rain forest. CoussapoastandleyiMacbride,Publ. Field Mus. Bot.
13(2.2): 298. 1937. Type. Peru. Huanuco:Huama-
Specimens studied. COLOMBIA. CAUCA:Rio Tim- lies, between Monzon and Rio Huallaga, - (9), We-
biqui, - (8), Lehmann BT948 (B, GH, K, NY). VALLE: berbauer3702 (holotype,B, destroyed;isotype, G).
Rio Callima, between Pailon and El Coco, 23 May Coussapoaaraneosa Standley,Publ. Field Mus. Bot.
1946 (Q),Cuatrecasas21256 (F); Rio Anchicaya,be- 17: 158. 1937. Type. Brazil. Amazonas:Mun. Hu-
tween Sabuletasand Quebradade Tatabro,28-29 Sep mayta,Tres Casas, 14 Sep-11 Oct 1934 (2), Krukoff
1946 (Q),Cuatrecasas22048 (F, type collection). 6524 (holotype,F;isotypes,A, BM, K, LE,MO, NY,
ECUADOR. ESMERALDAS: Junction of Rio Hoja S, U, US).
Blancaand Rio Hualpi, 14 Sep 1965 (6), Little et al. CoussapoabolivianaStandley,Publ. Field Mus. Bot.
21056 (NY, Q, US, type collectionof C. rotunda).Los 17: 159. 1937. Type.Bolivia. SantaCruz:Prov. Ichi-
Rios: Rio Palenque,alongroadaftercrossingRio Bimbe lo, Rio Vibora, 5 Oct 1926 (2), Steinbach7567 (ho-
and Rio Waija, 7 Oct 1976 (Q),Dodson et al. 6513 lotype, F; isotypes, A, GH, K, MO, S, U).
(MO, QCA). PICHINCHA: Nr. Santo Domingo de los CoussapoaeggersiiStandley,Publ.Field Mus. Bot. 17:
Colorados, 18 May 1955 (a), Asplund 16401 (S), 20 160. 1937. Type. Ecuador.Manabi:El Recreo, Dec
Aug 1920 (st), Benoist 3042 (P, U), 5 Feb 1979 (Q), 1891 (6),Eggers14165 (holotype,F; isotypes,prob-
Dodson 7416 (SEL,U), 5 Feb 1979 (6), Dodson et al. ably in A, BR, L, M, US).
7616 (MO), 9 Apr 1892 (9), Sodiro 193/12 (B). CoussapoaembiranaStandley,Publ. Field Mus. Bot.
17:161. 1937. Type.Brazil.Amazonas:Basinof Rio
In the shape and venation of the leaves, C.
Jurua,nr. mouth of Rio Embira (tributaryof Rio
vannifoliais reminiscentof broad-leavedforms Tarauaca),26 Jun 1933 (6),Krukoff4994 (holotype,
of C. asperifoliassp. magnifolia but also of C. NY; isotypes, A, G, K, U, US).
batavorum.Becauseof the differencein the num- CoussapoalawranceiStandley,Publ. Field Mus. Bot.
ber of stamens,a taxonomicrelationshipwith C. 17: 164. 1937. Type. Colombia.Boyaca:El Humbo,
130 km N of Bogota,ca. 1000 m, 26 Apr 1933 (6),
asperifoliais unlikely. Lawrance769 (holotype,F; isotypes,A, B, F, FI, G,
K, MO, S, US).
42. CoussapoavillosaPoeppig& Endlicher,Nov. CoussapoalehmanniiStandley,Publ. Field Mus. Bot.
17: 164. 1937. Type. Ecuador.Azuay:WesternAn-
2:
gen. sp. pl. 33, t. 147. 1838; Trecul, Ann.des of Cuenca, Chacayacu, - (2), Lehmann 5606
Sci. Nat. Bot. S6r. 3, 8: 99. 1847; Miquel, Fl. (holotype,F; isotype, K).
bras. 4(1): 138. 1853; Macbride, Publ. Field CoussapoasubcrenataStandley,Publ. Field Mus. Bot.
Mus. Bot. 13(2.2): 299. 1937. Type. Peru. 17: 166. 1937. Type. Brazil.Amazonas:Mun. Tefe,
Huanuco:Casapi(fideMacbride,1937) ($ and Rio Solim6es,Paleta,21 May 1933 (6),Krukoff4518
2 ?) Poeppigs.n. (holotype, W, destroyed;iso- (holotype,F; isotypes, A, BM, G, K, M, MO, NY,
S, U, US).
types (6), B, OXF). Fig. 54. CoussapoadanielisCuatrecasas,Caldasia7: 290. 1956.
100 Flora Neotropica

1 cm

FIG. 54. Coussapoa villosa: 1, leafy twig with pistillate inflorescences (Steyermark et al. 95170); 2, twig with
stipules and staminate inflorescences (Elias 631); 3, pistillate inflorescences (Donnell Smith 4826).
Coussapoa 101

Type.Colombia.Antioquia:Jardin,1800 m, Dec lutinous or to (sub)hirsute,often also with red-

1941 (2), Bro. Daniel 2652 (holotype,US; isotypes,
dish-brown pluricellularhairs and sometimes
COL,F). arachnoidhairs;perianthca. 1 mm high,densely,
CoussapoamacarenensisCuatrecasas,Caldasia7: 292.
1956.Type.Colombia.Meta:Sierradela Macarena, minutely puberulous;stamens two, (rather)far
CainoEntrada,24 Dec 1949 (5),Phillipson& Idrobo exceeding the perianth. Pistillate inflorescence
1914(holotype,BM;isotypes,COL,US). mostlyunbranched,sometimespoorlybranched;
CoussapoamacarenensisCuatrecasasvar. antioquien- heads 1(-4), free or occasionally partly fused,
sis Cuatrecasas,Caldasia7: 293. 1956.Type.Co-
lombia,Antioquia: RioCauca,PuentoValdiva,17- (sub)globose,ca. 5-10 mm, in fruitup to 40 mm
20 Feb 1942 (2), Metcalf& Cuatrecasas30093 (ho- diam.; (common)peduncle(1-)2-13 cm long, 1-
lotype,F; isotypes,A, MO,US). 4 mm thick, densely to sparselypuberulousto
CoussapoamutisiiKillip& Cuatrecasas, Caldasia7: hirtellousto shortlyvelutinousor to (sub)hirsute,
293. 1956. Type. Colombia.Withoutlocality, - (2),
Mutis646 (holotype,US). often also with reddish-brownpluricellularhairs
Coussapoaplanitiensis Cuatrecasas,Caldasia7: 295. and sometimes arachnoidhairs;perianthca. 1-
1956.Type.Colombia.Vaupes:Rio Guayabero, 8 2 mm high, densely minutelywhite to brownish
Nov 1939 (6), Cuatrecasas7510 (holotype,US). puberulous;fruitingperianthat the apex green,
Treeor shrub,hemi-epiphyticor terrestrial,up below the apex orange. Interfloral bracts
to 35 m tall. Leafytwigs 5-15 mm thick, sparsely (sub)spathulate to subpeltate,white to brownish
to densely white (to brown) puberulousto hir- puberulousat the apex.
tellous to (sub)hirsuteto (sub)villous. Lamina Distribution(Fig. 9). CentralAmerica (from
coriaceous, broadly ovate to subovate, some- Guatemala to Panama),the Andean region (Ec-
times to elliptic, occasionally to obovate, 6-60 uador to Venezuela), extending to the coastal
x 5-40 cm, apex obtuse to acute, sometimes mountain range of Venezuela, and wide-spread
rounded to emarginate,base obtuse to cordate (especially)in the (Upper)Amazon Basin;in up-
(sometimes deeply cordate with overlapping land, riverine, and mountain forest, up to 2400
lobes),marginentireto subcrenate;uppersurface m; often in secondaryvegetation.
glabrous,lower surface sparselyto densely mi-
Specimensstudied. GUATEMALA.IZABAL: Peten
nutely puberulousin the areolesand mostly also road, 2 km from Fronteras,Rio JuanVicente, 11 May
on the reticulum,hirtellousto hispidulouson the 1971 (8), Contreras10763 (DUKE, MO, P, US); Qui-
(parallel)tertiary venation, sometimes puberu- rigua,15-31 Mar 1922 (6),Standley23766 (GH, NY),
lous to (sub)hirsute,on the midriband the lateral 26-27 Apr 1939 (9), Standley 72288 (F, NY); Lago
veins sparsely,sometimes ratherdensely,puber- Izabal, between Punta dos Reales and Punta de Le-
17 Apr 1940 (2), Steyermark39618 (F).
ulous to hirtellous or to (sub)hirsute,the whole chuga,
HONDURAS. ATLANTIDA: La Ceiba, 22 Sep 1916
surfacecovered with sparseto dense whitish to (9), Dyer A85 (F, US); LancetillaValley, nr. Tela, 6
brownish subpersistentarachnoid hairs; lateral Dec 1927-20 Mar 1928 (st), Standley 53230 (A, F,
veins (7-)9-24, straightor slightlycurved, main US), (9),Standley53991 (A, F, US), (st),Standley54325
basal pair branched,reachingthe marginbelow (A, F), (2), Standley 55155 (A, F, US), (st), Standley
55445 (A, F, US), (6),Standley56623 (F).COMAYAGUA:
the middle of the lamina, in large leaves some- LakeYojoa, Pito Solo, nr. Pito Solo, 16 Apr 1951 (6),
times also with other lateral veins poorly P. H. Alien 6186 (F, GH), 8 Aug 1932 (st), Edwards
branched(furcate);intercostalvenation more or 400 (A, F, US); 15 km E of LakeYojoa, 20 May 1974
less prominent;petiole (2-)3-20 cm long, white (6),Hazeletts.n. (MO);nr. Pito Solo, 14 Aug 1973 (2),
Hazelett 664 (MO);Lake 29 Jul-10 Aug 1951
(to yellowish) puberulous to hirtellous to (6), Howardet al. 605 (A);Yojoa, nr. Pito Solo, 18 Apr 1945
(sub)hirsuteor to (sub)villous;stipules 3-20 cm (2), J. V. Rodriguez2929 (F); Rio Tempemechin,ca.
long, puberulousto hirtellousto strigoseto hir- 6 km N of Pito Solo, 15 Apr 1951 (d),L. O. Williams
sute, mostly also with dense (moniliform) red- et al. 18014 (F, GH). COPAN: Rio Copan, 2 mi E of
dish-brown pluricellularhairs and sometimes Copanruins, 27 Aug 1975 (8),Molina et al. 30747 (F,
MO). CORTES: Naciementodel Rio Lindo, nr. Jaral,9
arachnoid hairs. Staminate inflorescence Apr 1947 (6), Standleyet al. 7081 (F); Rio Londo, 8
branched,occasionallyunbranched;heads (1-)2- Apr 1945 (6),L. O. Williamset al. 12394 (F, GH);Rio
30(-ca. 50), often partly fused, (sub)globoseto Lindo, nr. Carrizal,12 Apr 1951 (st), L. O. Williams
ovoid to oblate, often more or less irregularin et al. 17817 (F), GRACIAS A Dios: Rio Platano, 0-4
hours upstreamRas, 23 May 1973 (9), Gentryet al.
shapeand dimensions,ca. 5-10(-20) mm diam.; 7530 (MO, U).
common peduncle 2-6 cm long, densely to NICARAGUA. BLUEFIELDS: Finca Santa Rosa, ca.
sparselypuberulousto hirtellous to shortly ve- 2.5 km ENE of Rama, 5 Apr 1966 (6), Proctoret al.
102 Flora Neotropica

Nr. La Libertad, 29 May-

27336 (F, US). CHONTALES: CricamolaValley, 17 Feb-l Mar 1928 (8), Cooper538
1 Jun 1947 (st), Standley 8861 (F). JINOTEGA:
Cord. (F, K); ChanguinolaValley, 10 Jan 1924 (2), Dunlap
Isabelia,Macizos de PeniasBlancas, 7 May 1976 (9), 299 (F, US);nr. ChiriquiLagoon,9 Nov 1940 (st), Von
Neill 7179 (MO). MATAGALPA: Rio Yasica, 20 km E Wedel 1548 (S), 22 Nov 1940 (6), Von Wedel 1733
of Matagalpa,25 May 1977 (2), Neill 1984 (U); 10-15 (GH, US), 26 Oct 1941 (9), Von Wedel2879 (GH, MO,
km NE of Matagalpa,16 Jan 1963 (2), L. O. Williams US);Rio Cricamola,betweenFincaSt. Louisand Kon-
et al. 24005 (F). NUEVASEGOVIA: Rio Solonli, 5 km kintoe, 12-16 Aug 1938 (6), Woodsonet al. 1893 (F,
N of Jalapa,5 Apr 1977 (8), Neill 1653 (U). ZELAYA:MO, NA, NY). CANALZONE:Barro Colorado Island,
ComarcaEl Hormiguero,W of Rio Uli, 6 May 1977 7 Feb 1969 (2), Croat7768 (F, MO), 10 Feb 1969 (6),
(6), Neill 1919 (U). Croat7839 (MO),21 Jun 1971(2),Croat15064(DUKE,
COSTA RICA. ALAJUELA: Between La Palma and F, GH, MO, NY), 25 Jun 1968 (2), Foster631 (DUKE);
Rio Platanillo, 19 Feb 1964 (6), Jimenez 1750 (US); Rio Providencia,5 Dec 1973 (st), Gentry8707 (MO,
NE-base of Volcan Arenal, 5 km W of FortunaSan U); Gatun, 4 Mar 1860 (9), Hayes 1008 (NY); Mindi
Carlos,29 Apr 1972 (6), Lent 2520 (F, MO, U, US); Hills, 28 Mar 1956 (st),Johnston1725 (A);BarroCol-
N side of Volcan Arenal, QuebradaGuillermina,21 oradoIsland, 19 Mar 1932 (9), Shattuck832 (F, MO);
Apr 1973 (st), Lent et al. 3388 (F). CARTAGO: Road nr. Fort Sherman, 15 Jan 1924 (st), Standley 30924
Turrialba-Moravia de Chirrip6,betweenTuis and Bajo (US); Barro Colorado Island, 18-24 Nov 1925 (2),
Pacuare, 15 Nov 1975 (8), Burger et al. 10040 (F); Standley41170 (US), 12 Mar 1931 (6), Wilson127 (F,
Turrialba,10 Dec 1952 (2), Cordoba295 (NA); Rio MO),6 Feb 1932 (6), Woodworthet al. 445 (A, F, MO),
Turrialba,Mar 1894 (2), DonnellSmith 4826 (GH, K, 22 Feb 1932 (9), Woodworthet al. 660 (A, F, MO).
US, type collectionof C. donnell-smithii);Atirro,Apr CHIRIQUI: La Fortunahydroelectricproject, 1200 m,
1896 (2), Donnell Smith 6770 (BM, F, GH, K, US); 22 Mar 1978 (2), Hammel 2176 (U). COLON: Rio
Rio Raventaz6n,nr. Turrialba,13 Feb 1971 (6), Gilles Guache, 1 Oct 1972 (st), Gentry6314 (MO);road to
et al. 10191 (F, GH); Rio Pejibaye,betweenRio Taus Portobelo,Villa Alondra, 13 Apr 1971 (6), Lao et al.
and QuebradaAzul, 28 May 1972 (2), Lent 2535 (F, 21 (DUKE, MO);Rio Fat6, Jul 1911, (2), Pittier3892
MO, U); Turrialba,May 1847 (st), Oersted14317 (C, (B, GH, NY, US, type collection of C. panamensis).
part of the type collection of Urostigmaintermargin- PERLASARCHIPELAGO: San Jose Island, 28 May 1945
ale), May 1898 (2), Pittier12288 (B, BM, GH, P, US), (2), Erlanson 240 (GH, NA, NY), 29 Mar 1945 (9),
1 Sep 1920(st),Rowleet al. 856 (NY, US);nr.Pejivalle, Johnston580 (GH), 2 Apr 1945 (6), Johnston606 (FI,
Jan 1940 (6), Skutch4612 (A, MO, NA, US), 7-8 Feb GH, MO), (st), Johnston 608 (GH), 5 Apr 1945 (2),
1926 (6), Standley et al. 47791 (US). GUANACASTE: El Johnston652 (BM,DUKE, FI, GH, MO, P, S), 10 Apr
Arenal, 18-19 Jan 1926 (2), Standleyet al. 45182 (F, 1945 (2), Johnston678 (GH).
US); nr. Tilerfn, 10-31 Jan 1926 (6), Standley et al. COLOMBIA. Without locality, - (2), Mutis 646
44550 (US), Standleyet al. 44968 (F, US), (2), Standley (US, typecollectionofC. mutisii),(a),Mutis4558 (US),
et al. 45661 (US), 3 Jan 1964 (2),L. O. Williamset al. - (2), Triana 867 (BM, K, P), - (6), Triana s.n. (COL).
26529 (F, US). LIMON: Road Bribri-Bratsi,Rio Six- AMAZONAS:Nr. PuertoNarifio,24 Jul 1965(6),Lozano
aola, 12 Feb 1977 (8), Burgeret al. 10474 (DUKE, F, 640 (COL), 28 Jan-7 Feb 1969 (6), Plowman et al.
MO); Las Diamantas, Rubber Expt. Station, 5 May 2362 (F, GH, K, NY, S), 13 Sep 1963 (2), Soejarto856
1943 (st),Dayton3019 (F);betweenPorteteand Moin, (A). ANTIOQUIA: Cocorna, Jun 1937 (st), Bro. Daniel
13 Feb 1965 (2), Jimenez 2887 (F); nr. mouth of Rio 1793 (US);Jardin, 1800 m, Dec 1941 (2), Bro. Daniel
Blanco,N of Moin, 13 Feb 1965 (9), Lent 349 (COL); 2652 (COL,F, US, type collectionof C. danielis);Rio
ca. 4.5 km N of El Carmen, 17 Mar 1973 (st), Lent Porce, 1100 m, 19 Sep 1963 (2), Espinal 1314 (COL);
3279 (F); Zent, Oct 1904 (6 and 2), Pittier s.n. (GH, Rio Cauca,PuertoValdiva, 17-20 Feb 1942 (2), Met-
US); affluenceof Rio Parisminaand Rio Reventaz6n, calf& Cuatrecasas30093 (A, MO, US, type collection
Suerre,3 Oct 1951 (st), Shank et al. 4294 (F);nr. Gua- of C. macarenensisvar. antioquensis).BOLiVAR:150
piles, 12-13 Mar 1924 (6), Standley 37259 (US); Rio km N of Barrancabermeja, Mico-Ahumadocamp, ca.
Reventaz6n, Finca Montecristo,below Cairo, 18-19 8?15N, 74?4'W,25 Aug 1966 (6), De Bruijn 1123 (K,
Feb 1926 (6), Standley et al. 48594 (F, US); Puerto M, MO, NY, S, U, VEN). BOYACA: El Humbo, 130
Viejo, Apr 1895 (2), Tonduz 9520 (F, US); PUNTARE- km N of Bogotf,ca. 1000 m, 26 Apr 1933 (6),Lawrance
NAS:Golfito, 27 Jul 1951 (8),P. H. Allen 6280 (F, GH, 769 (A, B, F, FI, G, K, MO, S, US, type collectionof
US); BuricaPeninsula,QuebradaPalito, 6 Mar 1973 C. lawrancei).CALDAS:Salamina,Aug 1943 (?), Bro.
(st), Croat 22635 (MO). SAN JOSt: Guayabo, 3 Feb Tomas 1923 (US). CAQUETA: Rio Caquetf, Rio Orte-
1908 (st), Ferrys.n. (F); Rio Vargas,Tabarcia,Mora, guaza, nr. Potosi, 7 Mar 1945 (2), Little et al. 9641
22 Apr 1963 (2), Jimenez 657 (COL,F); San Isidro El (NY, US), 15 Mar 1945 (8),Littleet al. 9789 (NY, US).
General,3 Mar 1966 (6), Molina et al. 18289 (F, US); CHOC6:Between Alto Curicheand Camp Curiche,E
El General,Mar 1939 (6), Skutch4256 (A, K, MO, S, od Boca Curiche,20 May 1967 (2), Duke et al. 11273
US), (2), Skutch 4262 (A, K, MO, S, US), (6), Skutch (US);road Medellin-Quibd6,6 km W of Siete, 38 km
4267 (A, K, MO, S, US); Las Vueltas,Tucirrique,Feb W of Bolivar, 1400 m, 6 Jan 1979 (2), Gentryet al.
1899 (8), Tonduz 13280 (BM, F, G, K, LE, MO, P, 23713 (MO, U). CUNDINAMARCA: 5 km W of El Salto
US). de Tequendama,on road to El Colegio, 1800 m, 14
PANAMA. Without locality (Isthmusof Panama), Jul 1972 (8),Barclayet al. 3593 (NA);betweenEl Salto
1859-1860 (st) Hayes 414 (NY), Hayes 867 (NY), (2), and El Colegio, 1480 m, 10 Mar 1940 (st), Cuatrecasas
Hayes 986 (NY), (2), Hayes s.n. (BM). BocAs DELTORO: 8291 (F, US); nr. Albfn, Aug 1962 (2), Ferndndez-
Coussapoa 103
Perez F3 (COL);3 km N of Tena, nr. La Lagunade et al. 95170 (NY, U, US, VEN); nr. Canoabo, 5 Mar
PedroPalo, 2080 m, 9 May 1952 (8),Ferndndez-Perez 1964 (8), Trujillo6141 (MY). DepartamentoFederal:
et al. 1458 (US); Mun. La Mesa, road to Mosquera, Nr. Caracas,1940 (6),Bro. Elias 130 (F), Jan 1931 (6),
1800 m, 5 Aug 1963 (8), Soejarto 327 (A, K); Rio Bro. Elias 631, 1 Oct 1972 (6),Hoyos 2043 (VEN), 17
Bogota, 1600 m, 1909 (a), Uribe-Uribe364 (COL). Nov 1975 (2), Olivas.n. (MY, VEN), 24 Nov 1975 (9),
META: Rio Negro, 20 km W of Villaviicencio,24 Mar Oliva s.n. (VEN). FALCON: Distr. Acosta, Mun. Jacura,
1972 (Q),Barclayet al. 3294 (NA, US); nr. Villavicen- Cerrade la Mina, 13 Feb 1961 (st), Ruiz-Terdn423
cio, 1-15 Aug 1946 (8),Duque-Jaramillo3957 (COL), Res. For. Rio Tocuyu,
(MER). FALCON-YACACUY:
- (a), Karstens.n. (LE);Sierrade la Macarena,Calno QuebradaEl Charalito,Aug 1970 (9),Blanco 958 (NY,
Entrada,24 Dec 1949 (6), Phillipsonet al. 1914 (BM, VEN). MERIDA: Merida(?), 26 Jun 1956 (9), Bernardi
COL,US, type collection of C. macarenensis),13 Jan s.n. (MY). MIRANDA: Cerros del Bachiller, between
1950 (2), Phillipsonet al. 2095 (BM);nr. Villavicencio, QuebradaCorozaland QuebradaSantaCruz,S of San-
Llanode SanMartin,Feb 1856 (9), Triana1866 (COL, ta Cruz,18-19 Mar 1978 (st),Steyermarketal. 116463
F, K, NY). VALLE:Rio Sanquinini, La Laguna, 1250- Bosques de Yumare, 10 Feb
(MO, U, VEN). YARACUY:
1400 m, 10-20 Dec 1943 (st), Cuatrecasas15589 (F); 1959 (8),Bernardi7017 (FI, G, MO); 35 km N of San
Gibraltar,N of Las Brisas,2100-2200 m, 25 Oct 1946 Felipe, 19 Feb 1954 (6),Little 16245 (VEN);El Guare-
(2), Cuatrecasas 22526 (F). VAUPES:Rio Guayabero, mal, between Valencia and San Felipe, Feb 1919 (8),
8 Nov 1939 (a), Cuatrecasas7510 (US, type of C. Pittier8404 (GH, US, VEN); Rio Taria, La Llanada
planitensis). de Taria, 5 Mar 1973 (st), Romero 548 (MY). ZULIA:
VENEZUELA. Without locality, 26 Jun 1956 (2), QuebradaPerayra, affluent of Rio Tokuku, SW of
Bernardis.n. (8), 4 Mar 1957 (6), Bernardis.n. (NY), Machiques,29 Aug 1967(St),Steyermark99829 (VEN).
1865 (a), Ernst 324 (BM), 1930-1934 (a), Vogl 1379 ECUADOR. AZUAY:Chacayuca, western Andes of
(BR). AMAZONAS: Raudal de los Guaharibos, 24 Jul Cuenca, - (9), Lehmann 5606 (F, K, type collection
1951 (st), Croizat312 (US); Salto Salas, 20 Aug 1951 of C. lehmannii). ESMERALDAS:Borb6n, 11 Sep 1965
(2), Croizat 541 (US). ARAGUA:Cerro de Avila, 1 May (8), Little et al. 21047 (NY, Q, US); 10 km SE of Es-
1859 (8), Cruegers.n. (K); Colonia Tovar, May 1854 meraldas,29 Sep 1965 (9), Little et al. 21172 (NY, Q,
(2), Fendler1237 (GH, K), Jul 1854 (a),Fendler1237a US). GUAYAS:Nr. Naranjito, 6-7 Jul 1945 (st), Camp
(GH, MO);nr. Maracay,30 Jan 1857 (6),Fendler1237 E3626 (NY, U); Baloa, - (9),Eggers14165 (A, L, LE,
(G), 1860 (9), Fendler 1237a (G), Rancho Grande, 6 M, US). IMBABURA:
Nr. Lita, 7 Feb 1981 (8), Berg
Jun 1963 (st), Montaldo 3435 (MY); Rio Chuao, El 1241 (U). Los Rios: Pichilingue, 18 May 1943 (2),
Medio, 15 Mar 1926 (8), Pittier12124 (US, VEN);nr. Little6461 (F, US); Rio PalenqueBiol. Station,km 56
Maracay,2 Apr 1926 (9), Pittier 12142 (US, VEN), 2 on road Quevedo-Santo Domingo de los Colorados,
Apr 1926 (2), Pittier12144 (A, G, M, MO, NY, VEN); 11 Feb 1973 (6), Dodson 5241 (MO, QCA, SEL, U),
road Maracay-Choroni,nr. El Castafio, 18 Feb 1937 27 Feb 1975 (6),Dodson5778 (QCA,US), 29 Jul 1972
(a), Pittier 13985 (F, K, NY, US, VEN); Cerros de (6),Dwyer10305 (MO), 15 Feb 1974 (9), Gentry9934
Guamitas,1100 m, 27 Feb 1948 (a),Pittieret al. 15733 (MO, QCA, U), 22 Feb 1974 (6), Gentry10127 (MO,
(US, VEN);ParqueNacionalHenriPittier,31 Oct 1975 QCA, U); canton Vinces, hacienda Santa Lucia, 18-
(2), H. Rodriguez399 (MY); nr. El Castaino,23 Mar 28 Oct (8), Mexta 6573 (F, NA). MANABI: El Recreo
1953 (2), Schnee 1337 (MY);road Maracay-Choroni, Dec 1891 (8), Eggers 14165 (A, BR, L, M, US, type
20 Dec 1948 (6), Standen20 (MY);nr. El Castaino,21 collection of C. eggersii), 1897 (9), Eggers 15615 (F,
Feb 1968 (2), Trujillo8765 (MY);Guamitas, 12 May LE, P). MORONA-SANTIAGO: Macas, 18 Mar 1956 (2),
1938 (a), Ll. Williams 10071 (B, F, VEN), between Asplund19835 (S); 5 km SW of Macas, 26 Jan 1981
Guamitasand El Lim6n, 28 Jan 1940 (a),Ll. Williams (9), Berg 1222 (U); Cord. de Cucutfi, 5-10 km E of
12348 (F, S, US, VEN); nr. El Castafio, 18 Feb 1937 Logronio, 1200-1500 m, 7-9 Oct 1975 (6), Little et al.
(st), Ll. Williams 13895 (US). BARINAS:Road Barini- 602 (COL, Q). NAPO: Rio Cuyabeno, ca. 0?10'S,
tas-Barinas,Mar 1953 (9),Aristeguieta1699 (MY,NY, 76?55'W, 16 Feb 1980 (2), Berg & Akkermans1033
US, VEN); nr. Barinitas, QuebradaParangula,Mar and 1040 (U), 19 Feb 1980 (6), Berg & Akkermans
1953 (9), Aristeguieta1704 (MY, NY, VEN); conflu- 1062 (U); Misahualli,mouth of Rio Misahualli,3 Mar
ence of Rio Curbaciticoand Rio Madredel Monte, 10 1980 (9),Berg&Akkermans1113 (U); 1 km S of Lago
Dec 1954 (a),Bernardi1724 (F, FI, K, VEN);Alto del Agrio, 5 Nov 1974 (6), Gentry12474 (MO, U); Rio
Aguada,Pedreza, 19 Feb 1955 (9), Bernardi1980 (F, Putumayo,just above mouth of Rio Gueppi, 19 May
NY); Barinitas,26 Jun 1956 (st), Bernardi3354 (NY); 1978 (9), Gentry 22095 (MO, U). PICHINCHA: Road
betweenAltamiraand Filo de Mertiuez,nr. Calderas, SantoDomingode los Colorados- Quinind6(km 170-
4 Mar 1957 (6),Bernardi6348 (G);Res. For.Ticoporo, 175), 1 Sep 1949 (st), Acosta Solis 13629 (F); Santo
Rio Bumbum,3 May 1964 (8),Breteler3889 (G, MO, Domingo de los Colorados,20 Aug 1930 (st), Benoist
NY, P, S, U, US, VEN); Rio Caparo,2-5 km above 3044 (P, S, U, US); nr. Alluriquin,bridge over Rio
dam, 12 Mar 1980 (9), Liesner et al. 9443 (MO, U); Toachi, 23 Feb 1981 (6), Berg 1301 (U); road Santo
Cuidad Bolivia, 12 Jan 1968 (a), R. F. Smith 3313 Domingo de los Colorados-Chona,km 5, 3 Apr 1943
(VEN);Res. For. Caparo,E of El Canton, 10 Apr 1968 (8),Little6155 (K, Q);SantoDomingo dos Colorados,
(2), Steyermarket al. 102083 (U, VEN), 12 Apr 1968 21 Apr 1943 (9), Little 6328 (F, K, NY, Q, US); nr.
(2), Steyermark et al. 102211(F, MY, VEN).CARABOBO:Alluriquin,at confluenceof Rio Alluriquinand Rio
Bejuma,18 Feb 1954 (8),Little 16230 (VEN);Rio San Toachi, 14 Mar 1967 (8), Sparre 14822 (S). ZAMORA-
Gian, S of Borburata,27-28 Mar 1966 (9),Steyermark CHINCHIPE: Quebrada Las Pavas Pavas, 14 km from
104 Flora Neotropica

Loja, 1800 m, 21 Aug 1975 (st), Samaniego et al. 92 K,NY,P,R,S,U, US);RioPurus,betweenLagoQuati

(COL,Q). andLagoArima,20 Jun1971(6),Pranceet al. 13438
PERU. Without locality, - (9), Dombey s.n. (F, P, Jun 1851(8),
(K,M, MO,NY, S, U, US);Manaquiri,
US), 1835 (6), Matthews2068 (K, OXF, U), 1829 (6), Spruce1608(BM,BR,C,G, GH,K,LE,NY,P, OXF).
Poeppigs.n. (B, OXF, type collection).AMAZONAS: Rio MATOGROSSO: Rio Aripuana,nr. Humboldt Center,
Cenepa,Aintami Creek,24 Nov 1972 (6), Berlin 372 ca.10?12'S, 59021'W, 12Oct1973(9),Bergetal.P18455
(MO, U); Pongo de Manseriche,1924 (9), Tessmann (F, K, MO,NY, S, U, US), 22 Oct 1973(6),Berget
4696 (B, NY); Rio Santiago,Caterpiza,20 Nov 1979 al. P19841(F, K, MO,NY, P, S, U, US);Sarar6,14?
(8), Tunqui103 (U). Huanuco:Pampayacu,nr. Huana- 50'S,59'45'10"W, 4 Aug 1972(6),Pireset al. 16461
co, 20 Jan 1927 (6), Kanehira99 (GH);Prov. Leoncio (U). PARA: AltoTapaj6s,Rio Curucii1-5 km above
Prado, Tingo Maria, 7 Dec 1981 (6), Plowman et al. MissaoCururui,7?35'S,57?31'W,17 Feb 1974 (9),An-
11180 (U); Macora,- (9),Ruiz & Pavons.n. (or 3) (B, derson11033(MO,U);GlebaBacaja,belowmouthof
FI, type collection of C. vellera),- (5), Ruiz & Pavon Rio Bacaja,22 Nov 1980(2), Pranceet al. 26402(U).
s.n. (BR, G, OXF, US); Huamalies,betweenMonzon RORAIMA: Rio Mucajai,Posto 17 Mar 1971 (6),Prance
and Rio Huallaga, - (9), Weberbauer3702 (G, type et al. 11066(K,M, NY, P, S, U).
collection of C. standleyi).JUNIN:La Merced,Puente BOLIVIA.SANTACRUZ:Prov. Ichilo, Rio Vibora,
Quimiri,24 Oct 1950 (6),Nunez 2928 (US);Sanabeni, 5 Oct 1926(2), Steinbach 7567(A, F, GH,K, MO,S,
4 Sep 1960 (8), Woytkowski5956 (MO, U). LORETO: U, typecollectionof C. boliviana).
Prov. Maynas,Rio Itaya,above PuertoBelen, 16 Nov
1978 (9),Diaz et al. 618 (U); Prov. Maynas,Rio Itaya,
nr. Palo Seco, ca. 40 km above Iquitos, 20 Mar 1977
(6), Gentryet al. 18458 (MO, U); Prov. Requena,Rio Coussapoavillosais a widespreadspecies(like
Tapiche, tributaryof Rio Ucayali, 1 hour above Re- C. asperifolia)and is extremelyvariablein many
quena, 8 Dec 1972 (,), Gentryet al. 21291 (MO, U); characters.Although the occurrenceof some of
lower Rio Huallaga,Yan6n, 5-11 Sep 1929 (9), Killip the featuresis associatedwith certainpartsof the
et al. 29249 (NY, US, type collectionof C. grandiceps); branched pistillate inflores-
Rio Marafion,lower Rio Ampiyacu, 6 Mar 1977 (6), species area, e.g.,
Prance et al. 24687 (U); Prov. Maynas, Rio Nanay, cences in Central America and brownarachnoid
Mishana, 1978 (6), Ramirez 1084 (U); Rio Ucuyali, hairs in Colombia and adjacent partsof Ecuador,
Yarinacocha,between 10?Sand mouth, 1923 (6), Tess- infraspecifictaxa cannot readily be recognized.
mann 3461 (B, G, NY, S); lower Rio Huallaga,Yuri- Some specimensfromthe
peripheryof the species
maguas,26 Oct 1929 (6),LI. Williams4179 (F);Puerto areaand/or with featuresnearthe extremeof the
Arturo, Yurimaguas, 22 Nov 1929 (2), LI. Williams
5349 (B, F). (LORETO?): Rio Zungarococha,28 Jan variationmust be mentioned.Barclayet al. 3593
1978 (6),Ayala et al. 1436 (MO, U). MADRE DE Dios: (Colombia)has staminateinflorescenceswith ex-
Prov.Tambopata,affluenceof Rio TambopataandRio tremely numerous (ca. 50) heads; Gentryet al.
Le Torre, TambopataNat. Res., ca. 30 km SSW of 23713
Puerto Maldonado, 16 May 1980 (9), Balbour 5372 (Colombia) has pistillate inflorescences
(U); ParqueNacional del Manu, Rio Manu, Pakitza
with very short (less than 2 cm long) peduncles
station, 9 Nov 1980 (6), Foster 5721 (F). (in this respectit formsa transitionto C. duquei);
BRAZIL.Without locality, - (9), Martiuss.n. (M, Prance et al. 26402 (Brazil,Pari) is peculiarin
typecollectionof C. martiana),- (6),Martiuss.n. (BR, thatthe headof the pistillateinflorescenceis lobed
M, U, typecollectionof C.subincana).ACRE: Rio Acre,
and obliquelyattachedto the peduncle.This last
Itui,4 Nov 1923 (9),Kuhlmann759 (RB, U); Tarauaca,
1-2 km E of Rio Tarauaca,24 Sep 1968 (2), Pranceet collection matches some more or less aberrant
al. 7530 (F, GH, K, NY, P, R, S, U, US); Rio Jurui- collectionof C. tessmannii(see p. 95) in the dense
Mirim, Aug 1901 (6), Ule 5717 (B, G, K, L, MG). coveringof darkbrownmoniliformpluricellular
AMAZONAS: Rio Solimoes, Mun. Tefe, Paleta, 21 May
1933 (6), Krukoff4518(A, BM, F, G, K, M, MO, NY, (=granular)hairson the peduncle,leafytwig,and
S, U, US, type collection of C. subcrenata);nr. mouth petiole.
of Rio Embira,tributaryof Rio Tarauaca,26 Jun 1933 Typification of C. villosa: It is unknown
(6),Krukoff4994(A, G, K, NY, U, US, type collection whetherthe destroyedtype materialin W con-
of C. embirana);Mun. Humaita,Tres Casas, 14 Sep- sisted of a pistillateand a staminatespecimenas
11 Oct 1934 (9), Krukoff6165 (A, F, FI, K, MO, NY,
the drawingin Poeppig& Endlicher(1838) sug-
S, U, US), Krukoff6524 (A, BM, F, G, K, LE, MO,
NY, S, U, US, type collection of C. araneosa);Rio gests. The specimensin B and OXF which could
Madeira,Boa Hora, 4 Sep 1923 (2), Kuhlmann368 replacethe holotype materialare staminate.
(RB);Rio Japura,- (6), Martiuss.n. (G);Lagode Ma- Typificationof C. eggersii:Eggers made sev-
nacapuru,10 Oct 1972 (2), 0. Pires 248 (U); Tefe, 23 eralcollectionsundernumber14165- staminate
Jul 1972 (9),PLK& Urbano12296 (INPA);road Boca
do Acre-Rio Branco,km 1-4, 19 Sep 1966 (6),Prance and pistillateones from differentlocalitiesin Ec-
et al. 2306 (F, GH, K, P, R, S, U); Rio Demini, To- uador (El Recreo and Baloa)-at several dates.
totobi, 28 Feb 1969 (2), Prance et al. 10348 (F, GH, These collectionshave been mixed up. The spec-
Coussapoa 105
imen in F on which C. eggersii is based is sta- zonas), Guyana (Upper Mazaruni),and Brazil
minate and possibly from El Recreo.In the orig- (Roraima);in rain forest and savannas, up to
inaldescriptionthe specimenis wronglydescribed 1250 m.
as pistillate.
Specimens studied. VENEZUELA. AMAZONAS:
"Alto Orinoco," 1951 (2), Croizat959 (F, US); Cerro
43. CoussapoaviridifoliaCuatrecasas,Fieldiana Duida, CanioNegro, 25-26 Aug 1944 (2), Steyermark
Bot. 28(1): 213. 1956. Type. Venezuela. Bo- 57955(F, VEN,typecollectionof C. viridifolia var.
livar:0-8 km N of SantaTeresitade Kavana- tenuifolia).BOLIVAR:Region of Rio Icabaruand Rio
Hacha,2 Jan 1956(2), Bernardi2744 (F, FI, NY, VEN);
yen, 23 Nov 1944 (6), Steyermark60456 (ho- betweenSantaTeresitade Kavanayen andravineca.
lotype, F; isotype, VEN). Fig. 55. 8 km NW of it, 23 Nov 1944 (8), Steyermark60451
(F,VEN,typecollection).
Coussapoa Cuatrecasas
viridifolia var.tenuifoliaCua- GUYANA. Kamarang River,belowmouthofUtschi
Fieldiana
trecasas, Bot.28(1):214. 1956.Type.Ven- Creek,30 Oct1960(8),Tillettet al. 45869(F,NY, U).
ezuela.Bolivar:CerroDuida,CanioNegro,25-26 BRAZIL. AMAZONAS: Serrade Neblina, Rio Cau-
Aug 1944(9),Steyermark 57955(holotype,F; iso- aburi-RioMaturaca, 600-1300m, 16 Dec 1965(2),
type,VEN). N. T. Silva et al. 60670 (U). RORAIMA:
Serra Surucucu,
2 Feb 1975 (9),Ribeiro655 (IAN), 1 Feb 1975 (2), Rosa
Treeor shrub,often hemi-epiphytic,up to 20 339 (IAN).
m tall. Leafy twigs 2-5 mm thick, appressed-
This speciesappearsto be closely relatedto C.
puberulousto substrigoseto hirtellous,with hairs
differingdistinctlyin length.Lamina coriaceous latifolia. The two are allopatricand apparently
to subcoriaceous,obovate to elliptic to oblong, ecologically distinct, but morphologicallyvery
2.5-10 x 1-6 cm, apex obtuse to shortly acu- close. The two taxa show (minor) differencesin
minate or roundedto emarginate,base obtuse to the leaf venation, the length of the stipules, and
acute to cuneate to rounded, margin entire to the indumentumof the leaves. The morpholog-
ical differencessuggestthe possibility of distinc-
subcrenate;uppersurfaceglabrous,lowersurface
tion at the subspecificlevel.
sparselypuberulousto substrigoseon the main
veins, the hairs differingdistinctlyin length;lat- The followingthree species have been discov-
eral veins 4-7 pairs, basal pair unbranchedor
ered and describedafter submissionof the post-
faintlybranchedreachingthe marginat or above reviewmanuscriptof the revision and are,there-
the middle of the lamina, sometimes some of the
other lateralveins poorly branched(furcate);in- fore, not placed in the alphabeticalsequence.
tercostalvenation plane to more or less promi-
44. Coussapoafulvescens C. C. Berg, Brittonia
nent; petiole 1-3 cm long, sparsely (to rather 42: 59-65. 1990. Type. Colombia.Choc6:Rd.
densely) appressed-puberulousto substrigose,
with hairs differingdistinctly in length;stipules Quibdo-Bolivar,ca. km 50, 9 Jul 1986 (2),
0.5-1 cm long, densely appressed-puberulous to Berg 1546 (holotype, COL; isotypes, AAU,
subsericeous; terminal buds slender. Staminate BG, F, HUA, MO, NY). Fig. 56.
inflorescencesbranched;heads 3-14, globose,ca. Tall, hemi-epiphytic tree. Leafy twigs ca. 10
2-4 mm diam.; common peduncle 1-3 cm long, mm thick, white-appressed-puberulous,partly
sparsely puberulous;perianth ca. 1 mm high, also brownish-hirsute, lenticels conspicuous.
glabrous;stamens two, just exceeding the peri- Lamina coriaceous,oblong to subobovate, 18-
anth. Pistillate inflorescences unbranched or 26 x 9-15 cm, apex short-acuminateto obtuse,
poorlybranched;heads 1-3, globose,ca. 3-6 mm, base obtuse, margin entire; upper surface gla-
in fruit up to 8 mm diam.; (common) peduncle brous, lower surface sparsely appressed-puber-
0.5-2.5 cm long, sparsely(to ratherdensely)pu- ulous; lateral veins 7-9 pairs, basal pair un-
berulous;perianthca. 1 mm high, glabrous.In- branched(or very faintlybranched)reachingthe
terfloralbractsoften only a few (sometimeslack- marginat orjust below the middle of the lamina,
ing?),small,subspathulateto subpeltate,minutely usuallythe secondand/orthirdlateralveins (from
puberulousat the apex. the base) with one or two strong branches,ter-
Distribution(Fig. 9). EasternGuiana (=High- tiaryvenationdistinct,slightlyprominent;petiole
land) region, in Venezuela (Bolivar and Ama- 5-10 cm long,appressed-puberulous; stipules1.5-
cm
Coussapoa 107

FXG. 1,leafy with

twig
56. Coussapoafillvescens: (Berg
in~lorescences
Distillate x0.75.
1546),

r;:~~~~~~~~~~~~~~~~~~'
?~~~~~~~~~~~~~~~~~~~~~~'
"'
":~~~~~~~~~~~~~~~.:?
1~~?. ~ .

?i I?:i,
FIG.56. Coussaoafulvescens:
1leafy twig wihpistillate (Beg 1546),
inflorescence x0.75
108 Flora Neotropica

2 cm long, yellowish-to brown-(sub)hirsute.Pis- Specimensstudied.COLOMBIA.TOLIMA: 3.4 km

tillate inflorescences branched; heads 10-20, from Las Juntas, 1940 m, 9 Apr 1984 (v), Escobaret
sometimes partlyfused, (sub)globose,3-5 cm in al. 4219 (HUA, type collection).
diam.;common peduncle3-6 cm long, ca. 2 mm This species resemblesC. nitida, known from
thick, appressed-puberulous and partly also the Amazon Basin and mostly occurringin riv-
brownish-hirtellous;perianthca. 2 mm high,gla- ersidevegetation.It differsclearlyfrom the latter
brous, + distinctlyribbed.Interfloralbractssub- in the prominent tertiary and quartemaryve-
ulate to narrowlyspathulate,with a few hairs at nation at the lower surfaceof the lamina and in
the apex. the red-coloredfruitingperianth.
Distribution(Fig. 9). Colombia (Choc6), only
known from the type locality, in lowland sec- 46. Coussapoavalaria C. C. Berg, Brittonia42:
ondarygrowth. 59-65.1990. Type.Colombia.Valle:BajoCali-
Specimensstudied. COLOMBIA.CHOC6:Rd. ma, rd. to Dindo, 25 Sep 1986 (2), Monsalve
Quibdo-Bolivar, ca.km 50, 9 Jul 1986(9),Berg1546 B. 1176 (holotype, CUVC; isotypes, BG,
(AAU,BG,COL,F, HUA,MO,NY, typecollection). MO). Fig. 58.
This species is closely related to C. herthae, Tree, hemi-epiphytic. Leafy twigs 5-10 mm
known from western Ecuadorand Narifio (Co- thick, brown strigose,at least at the scars of the
lombia). It differs from the latter, e.g., in the stipules, glabrescent.Lamina coriaceous, ellip-
indumentof the leaves, the venation, the shorter tic, 15-17 x 11-14 cm, apex roundedto shortly
stipules, the more numerousand smallerflower acuminate,base roundedto subcordate,margin
headsof the pistillateinflorescence.The two taxa subentire,at the base revolute;uppersurfacegla-
might prove to be distinct only at the subspecific brous, lower surfacestrigilloseat the base of the
level. C. fulvescens may also be related to C. midrib, on the midrib and lateral veins with a
echinata. soon disappearingwhite arachnoid indument;
lateralveins 8-10 pairs, basal pair + branched,
45. Coussapoatolimensis C. C. Berg, Brittonia reachingthe marginat or just below the middle
42: 59-65. 1990. Type. Colombia.Tolima:3.4 of the lamina, tertiary venation almost plane;
km from Las Juntas, 1940 m, 9 Apr 1984 (9), petiole 5-6.5 cm long, glabrousor brownishstri-
Escobaret al. 4219 (holotype, HUA). gose, stipules 2-3 cm long, ratherdensely whit-
Fig. 57. ish appressed-puberulousto -strigillose,or also
with white arachnoid hairs. Pistillate inflores-
Tree, 18 m tall. Leafy twigs 5-12 mm thick, cences branched;heads 3-9, 3-4 (in fruit 7-9)
yellowish- to whitish-hirtellous.Lamina coria- mm diam.; common peduncle 2-4 cm
long,
ceous, ovate to elliptic, 10-21 x 8-16 cm, apex densely minutely puberulous;perianth ca. 1.5
shortly acuminate (to acute), base rounded to mm high, minutely puberulousaround the ap-
subcordate,margincoarselycrenateto subentire; erture.Interfloralbractsabsent.
upper surface with sparse hairs on the midrib, Distribution 9). Colombia(Valle),in low-
lower surfacehirtellous on the veins, in the re- land forest. (Fig.
ticulum minutely puberulous;lateral veins 12-
14 pairs, basal pair branched,reachingthe mar- Specimensstudied.COLOMBIA.VALLE: BajoCali-
below the middle of the ma, rd.to JuanchacoPalmeras,10Jul1984(9), Gentry
gin lamina, tertiaryand et al. 47838
(BG);BajoCalima,rd.to Dindo,25 Sep
quarterary venation and reticulumprominent; 1986 (9),MonsalveB. 1176 (BG, MO, type collection).
petiole 4-6 cm long, 2-4 mm thick, hirtellous,
hairs not distinctly differentin length; stipules This species resemblesboth C. ovalifoliaand
4-12 cm long, denselyyellowish-puberulousand C. parviceps.It differsfrom these species in, e.g.,
partly also hirsute. Pistillate inflorescences the venation and the numberand dimensionsof
branched;heads 4-5, globose (to subdiscoid),in the flowerheads of the pistillate inflorescences.
fruit 0.7-1.3(-1.5) mm in diam.; common pe-
duncle 2.5-3 cm long, ca 2-2.5 mm thick, hir- Unnamed Collections
tellous; perianth ca. 1.5 mm high, at the apex Gentry& Mori 14011 from Panama:Discussed
puberulous,in fruitred. Interfloralbractsabsent. under C. ovalifolia.
Distribution(Fig. 9). Colombia(Tolima),only Huashikat 1585 from Peru:Discussed under C.
known from the type locality. cupularis.
 Coussapoa 109

FIG. 57. Coussapoatolimensis: 1, leafy twig with pistillateinflorescences(Albertde Escobaret al. 4219), x0.6.

Names Not Included of discordant elements: staminate inflores-

cences of a Coussapoaspecies and leafy twigs
Coussapoa dealbata Andre, Ill. Hort. 17: 17. of some species not belongingto the Urticales,
1870--nomen nudum. but possibly to the Lauraceae.The latterparts
CoussapoadolichandraCuatrecasasCaldasia7: constitute the lectotype, chosen here.
287. 1956-based on collection Cuatrecasas CoussapoaemarginataKillip in Macbride,Publ.
17458 (Colombia,Valle: Rio Cajambre,5-15 Field Mus. Bot. 13(2.2): 296. 1937 = Pourou-
May 1944 (holotypeand isotype, F)-consists ma minor Benoist.
110
FIG. 58. Coussapoavalaria:1, leafy twig with pistillateinflorescences(Gentryet al 47838), x0.55.
CoussapoakrukoviiStandley, Publ. Field Mus.
Bot. 17:1937 = PouroumatomentosaMiquel.
CoussapoalaevigataPoeppig& Endlicher,Nov.
gen. sp. pl. 2: 33. 1838-based on material
from Brazil (Amazonas:Tefe). The holotype
in W has been destroyed. Isotypes have not
been traced. The identity is uncertain. The
name possibly applies to C. villosa.
CoussapoaleopoldiiMicheli, Rev. Hort. (1894):
251. 1894-nomen nudum.
Coussapoa obovata Warburgex Glaziou, Bull.
Soc. Bot. France 59 (Mem. 3g): 645. 1913-
nomen nudum.
Coussapoapittieri Standley in Hoyos, Los Ar-
bolesde Caracas,ed. 2 (Monogr.24. Soc. Cienc.
Nat. La Salle, Caracas,Venezuela)p. 242, tab.
(of C. villosa)-nomen nudum.
Coussapoaplicata Warburgex Ule, Bot. Jahrb.
40: 143. 1907- nomen nudum.
CoussapoarekoiStandley,Contr.U.S. Natl. Herb
20: 211. 1919 = Poulsenia armata (Miquel)
Standley:see Mildbraed,Notizbl. Bot. Gart.
Berlin 10: 413. 1928; Standley,Trop. Woods
33: 4. 1933.
Urostigma intermarginale Liebmann, Kongel.
DanskeVidensk.-Selsk.Skr.Ser.5,2:328.1851
Flora Neotropica
= Ficus intermarginalis(Liebmann) Miquel,
Ann Bot. Lugd.Bat.3:297. 1867;see Standley,
Publ. Field Mus. Bot. 18(1): 383. 1937.
POUROUMA
Introduction
Pourouma is the smallest of the three Neo-
tropicalgeneraof the Cecropiaceae.It is a well-
definedgenus of small to medium-sized trees of
the rain forestareasof South and CentralAmer-
ica, and has not drawn much attention, except
that the fruits of P. cecropiifoliaare suitable for
human consumption. The genus is rather uni-
form in its ecology. Besides a number of clear-
cut species, mostly of limited distribution, it
comprises several more wide-spreadand rather
complex groups of taxa with a confusing varia-
tion in leaf shapes and indument, and to which
the followingquotation applies:"The species of
Pouroumaare perhapsthe most ambiguousand
vexatious of the AmericanMoraceae,a dubious
distinction"(Woodson& Schery,1960:166). The
lack of sufficientfield work must be deemed a
greatdisadvantagein the preparationof the pres-
Morphology 111

ent revision, especially with regardto the vari- Theyareoftenverydenseand forma powdery
ation in the leaves. layerin driedmaterial.The densenessof these
hairs is often variable.
Taxonomic History 2. Unicellulararachnoid(cobwebby)hairs.These
very thin, crinkled, interwoven hairs of dif-
The genus Pouroumawas establishedby Au- ferent length are mostly white, sometimes
blet (1775) who described one species, P. gui- brownish(in P. ferruginea).Short arachnoid
anensis. Names were added by Martius(in Spix hairs are always found in the areoles at the
andMartius,1831, 1843),Tr6cul(1847), Klotzsch lowerleaf surface.Longerarachnoidhairscan
(1847), and Miquel (1853), and in this century be found on the smaller veins, and in some
by Rusby (1910), Benoist (1922, 1924), Mild- species (e.g., P. ferrugineaand P. tomentosa)
braed (1927, 1928), Macbride (1930), Ducke also on the main veins, fromwhichthey often
(1932a, 1932b, 1947), Standley (1937a, 1937b, disappearwith age. Longarachnoidhairsalso
1939),Cuatrecasasand Standley(in Cuatrecasas, occuron the petioles,stipules,leafytwigsand/
1951, 1956b),Cuatrecasas(1954, 1956a, 1956b, or the inflorescencesin some species.
1956c, 1967), Woodson and Schery(1960), and 3. Unicellularnon-arachnoid,thickerand most-
finally by Berg and Kooy (1982), Berg and van ly more or less straighthairs. Short hairs are
Heusden (1988), and Berg (1989, 1990). usually whitish, the longer ones mostly yel-
Since the comprehensivetreatments of Pou- low(ish).They can be found on most partsof
rouma by Trecul (1847) and Miquel (1853) the the plant. In some species, forms with only
genus has only been treated for regional floras: short, whitish hairs occur beside forms pos-
Peru (Macbride, 1937), Panama (Woodson & sessing longer yellowish hairs. In several
Schery, 1960), Costa Rica (Burger, 1977), and species, all or some specimens have short,
Surinam(Berg, 1975). rigid hairs on the upperor sometimes on the
lower leaf surface.
Morphology
Leaves.-The leavesaremedium-sizedto large,
Habit.-Pourouma species are usually small
entire or palmatelyincised. In taxa with incised
to medium-sizedtrees,oftenwith stilt-rootsfrom
leaves the first formed leaves are entire; subse-
the lower part of the trunk, as in most of the
leaves sooner or later, frequentlyor al-
other terrestrial,non-climbingspecies of Cecro- quently
ways, become incised. In adult specimens, es-
piaceae.In some species (e.g.,P. hirsutipetiolata) on floweringbranches,entireleaves can
the trunkmay become buttressed.The architec- pecially
be differentin shape and
ture of the trees of P. minoris that of the model again found, although
texturefrom the leaves in the juvenile stage.En-
of Aubreville (F. Halle, pers. comm.). The
tire and incised leaves may occur on the same
branchesmay be thick (as in specieswith incised
branchlet.Leaves similarto those formedin the
leaves)or ratherthin (as in specieswith medium-
subjuvenile stage often appear again on sucker
sized, entire leaves). shoots of adult specimens with entire leaves.
The wood (andothervegetativeparts)of some
Leaves are always incised in adult specimens of
species (e.g., P. myrmecophila)give out a pecu- Pourouma
liarodor,indicatedas 'spearmint','winter-green', cecropiifolia,and probablyalso in P.
myrmecophila, P. oraria, P. stipulacea, and P.
'balsam',or 'bengie'. For other species (e.g., P. villosa.Incisedleaves
appearto occurconstantly
quianensis)a 'balsam' odor is reported for the in some of species in which entire
fruits. subspecies
leaves are common. In species in which only
When cut the plant parts exude a watery sap
entireleaves are found, the leaves in thejuvenile
which is clear or sometimes reddish (e.g., in P.
and subjuvenile stages are basically similar to
bicolor);after exposureto the air it turns black. those of adult
Indumentum.-Three types of trichomes can specimens. A number of species
appear to be constantlyentire-leaved(e.g., P. bo-
be recognized:
livarensis,P. minor, P. ovata, and P. phaeotri-
1. Pluricellularhairs,often more or less globose, cha). In this groupof speciesthe leaves may vary
pale brown to dark brown or purple. These from predominantlyovate to elliptic to oblong,
hairs occur on most young parts of the plant. or even to obovate.
112
Leaf texture varies from chartaceousor sub-
coriaceousto coriaceous,andleafindumentfrom
dense (with the three types of trichomes repre-
sented)to sparse.
The venation varies from pinnate in entire
leaves to subpalmatein incised leaves. In species
in which palmatelyincised leaves occur,the bas-
al lateralveins even of entireleaves arenormally
branched,while in speciesin whichincisedleaves
are not formed, the basal lateralveins are often
unbranchedor only faintlyor sparselybranched.
The tertiaryvenation is scalariform.
The leaf marginis entire or often more or less
faintly crenate. In most species the petioles are
relativelylong, with a considerablevariation in
length on the same twig. Relatively short peti-
oles, only slightlyvaryingin length,are found in
a few species (e.g., P. formicarum). In P. for-
micarum and P. myrmecophilathe base of the
petiole is swollen and has an abaxial slit. These
two species are myrmecophilous.
The mostly largestipulesareconnateand fully
amplexicaul,and appearto be of importancefor
the protection of young inflorescences-espe-
cially staminateinflorescenceswith flowerswith
a connate (more or less urceolate)perianthand
the anthers alreadyexserted before anthesis, as
in P. tomentosa.In some species the stipulesare
short, e.g., P. ovata. While in most species the
stipulesare caducous,they are (sub)persistentin
P. myrmecophila,P. oraria, and P. stipulacea.
The stipulesleave a horizontalannularscar,not
oblique as in Coussapoa.This featuremay help
to distinguishsterilespecimensof the two genera,
of which the leaves can be very similar.
The leaf charactersmentionedare more or less
correlated.In taxa with incised leaves the leaves
tend to be large,often subcoriaceousand densely
hairy, with relatively long petioles, often quite
variablein length, and often with long stipules.
Contrariwise,in taxawith entireleaves,the leaves
tend to be medium-sized,usuallycoriaceousand
often rathersparselyhairy. They have relatively
short petioles, more or less constant in length,
and the stipules tend to be small.
Inflorescences.-The inflorescencesare nearly
always ramified, basically with three branches
departingfrom the top of the peduncle.Further
ramificationis usually (sub)dichotomous.The
primaryramificationcan be dichotomousas well.
In pistillateinflorescencesthe ramificationis often
more reduced than in the staminate ones, with
a tendencyto a subumbellatearrangementof the
FloraNeotropica
flowers. A subumbellatepistillate inflorescence
is characteristicfor Pouroumaminor.Staminate
inflorescencesare often more or less dorsiven-
trally flattened.
The flowersof the staminateinflorescencesare
more or less closely clusteredat the end of the
branches. In several species the flowers form
(sub)globoseheads.
The inflorescencesare mostly ebracteate,al-
thoughminutebractsare found in staminateand
pistillate inflorescencesof P. cucura,and some-
what larger(caducous)bracts at the base of the
peduncleare sometimes found in other species.
Staminate Flowers.-Staminate flowers are
pedicellateor sessile and tetramerous,or some-
times trimerous.The tepalsarefree,basallycon-
nate, or almost entirely connate, and then they
form an urceolateor infundibuliformperianth.
The stamens are short in species with free or
basallyconnatetepals,but in specieswith almost
entirely connate tepals the filaments are longer
than the tepals. Probablyin connectionwith the
rathernarrowopeningof the urceolateperianth,
the anthersareoutsidethe perianth(long?)before
anthesis. The filamentsare free, sometimes ba-
sally connate,but in Pouroumamelinoniisubsp.
glabratathey canbe entirelyconnate,thuslargely
similar to the androecium characteristic for
Coussapoa.In the center of the flower one can
usuallyfinda tuft of hairs,which mightrepresent
a pistillode.
Pistillate Flowers.-Pistillate flowers are al-
ways pedicellate.The perianthis tubularwith an
entireor slightlylobed margin.The ovary is free
from the perianth.The shortstyle bearsa stigma
that is subpeltateor peltateto knob-like,in Pou-
rouma minor.
Fruitsand Seeds.- The fruithas a drypericarp
and is completelyenclosed by the enlargedperi-
anth. The inner layerof the fruitingperianthbe-
comes soft and whitish and the otherlayer(skin)
? leatheryand at full maturityoften purple to
black or red-brown.The whole (fruit and peri-
anth)can be indicatedas a pseudodrupe(or func-
tional drupe).
The pedicel under maturefruits may become
reddish (a contrasting color against black?).
Whetherthis color is causedby a changeof color
of tissues of the pedicelor by the presenceof red-
brown pluricellularhairs is not clear.
The fruiting perianth is reported to be aro-
matic for some species (e.g., Pouroumabicolor).
Pourouma and the African Myrianthus are
Morphology;Phenology;Pollination
macrospermous,in contrast to the other four
generaof the Cecropiaceae.
GeneralRemarks.-The greatestvariation of
characters(mainly,but not only of the leaves) in
the genus is found among taxa in which incised
leaves do not occur constantly.Only part of this
variation is connected with distribution. Taxa
with constantlyincised leaves in adultspecimens
and those with entire leaves in all stages of de-
velopmenttend to be ratheruniform,even if they
have a relativelywide distribution.
It is somewhatdifficultto indicatewhich char-
acters can be regardedas derived or as special-
ized. In severalof its characters(especiallyof the
vegetative parts) Pouroumais (like the African
genus Myrianthus)intermediatebetween Cecro-
pia (and the Africangenus Musanga)and Cous-
sapoa(andthe MalesiangenusPoikilospermum).
From about the middle of the spectrumof vari-
ation in Pourouma one could indicate a pro-
gressiontowardsCecropia,endingin a form with
thickbranchesand large,constantlyincisedleaves
with many incisions (but never peltate as in Ce-
cropia),represented,e.g., by P. cecropiifoliaand
P. cuspidata;and another progressiontowards
Coussapoa,ending in a form with ratherslender
branches and medium-sized, constantly entire
leaves. As indicated (p. 5), there is an overlap
of the leaf charactersof Pourouma and Cous-
sapoa, and sterilematerialof both generacan be
easily confused.
In addition to these charactersone can also
regardas derived:sparseindument(e.g.,Pourou-
ma ovata), elliptic to obovate leaves (e.g., P.
phaeotrichaand P. bolivarensis),unbranchedlat-
eralveins (e.g., P. acuminataand P. ovata),peti-
olulateleaf segments(in taxawith incisedleaves),
as in P. tomentosasubsp.persecta,shortpetioles
of constant length (P. formicarum),swollen and
saccate base of the petiole (P. formicarum, P.
myrmecophila),shortstipules(e.g.,P. ovata),sta-
minate flowers in globose heads (P. melinonii
and P. myrmecophila),subumbellate pistillate
inflorescences(P. minor), elongate peduncle of
the pistillate inflorescence(P. ferrugineaand P.
ovata), staminate flowers with fused tepals (P.
mollis), stamens with partlyor fully connate fil-
aments (P. melinonii and P. napoensis respec-
tively), and knob-like stigma (P. minor).
The above summation of derived characters
and specializationsshows that they tend to be
'at the Coussapoaside' of the variationspectrum
in Pourouma.
113
Phenology
From label data alone it is difficultif not im-
possible to draw conclusions about the phenol-
ogy of a particularspecies.
In an area explored (Montagne de Trinite,
FrenchGuiana,shortrainyseason,Jan-Feb 1984)
none of the numerous Pourouma specimens of
different species was bearing flowers or fruits.
This suggestedseasonal flowering.The combi-
nation of label data of all Pouroumacollections
made in the Guianas gives a patternindicating
a main floweringperiod in August-October(in
the long dry season) and a fruitingperiod which
may extend to February,thus from the end of
the long dry season into a short rainy season.
According to Croat (1978) P. bicolor fruits in
Panamain the end of the dry season. According
to Martius(in Spix & Martius, 1831) P. cecro-
piifolia can bear (ripe?)fruits in May and No-
vember. The occurrenceof these two periods of
fruitinggets some supportfrom the label data of
the examinedmaterialof this species.Falcaoand
Lleras (1980) found that P. cecropiifoliain the
Manausregionwas floweringat the height of the
rainyseason (April-June),and producinga crop
of ripe fruit at the end of the dry season to the
beginningof the next rainy season.
The periodbetweenfloweringand bearingripe
fruitappearsto be longerin P. cecropiifoliathan
in (most?)other species of Pourouma.The dis-
crepancybetween the data supplied by Falcao
andLlerasand the observationsof Martiusmight
be caused by the presenceof several strains. In
the publication"UnderexploitedTropicalPlants
with Promising Economic Value" (National
Academy of Sciences, Washington,D.C., 1975)
a fruiting period of three months in the rainy
season is mentioned for P. cecropiifolia.
Pollination
Little is known about pollination in Pourou-
ma. Neither morphologicalfeatures(of inflores-
cences and flowers)nor habit and habitat(small
to medium-sizedtrees in forests)suggestthe oc-
currenceof wind pollination.The herbariumma-
terial indicates that most pistils develop into
fruits,which suggestsan effectivepollinationsys-
tem. Falcao and Lleras (1980) found a number
of bee species as pollinators of P. cecropiifolia.
The bees collect pollen in the staminate inflo-
rescencesand fly to pistillateinflorescences.The
114
authors do not give indications about the at-
tractant(s)of the pistillate inflorescences.One
may wonder whether the pluricellular('glandu-
lar') hairs, which occur so abundantlyin inflo-
rescencesof most Pouroumaspecies (as a pow-
dery layer, as in dry material?),may play a role
(as attractantor a substitutefor pollen?).
Dispersal
Again, little is known about dispersalof Pou-
rouma seeds. One may assume that the 'fruits'
with a fleshy mesocarp-likeinner layer of the
enlargedperianth, and a purple to black (or a
red-brown)exocarp-like outer layer (skin) are
eaten by several frugiverousarboreal animals;
monkeys are mentioned in label data.
The presenceof very long pedunclesof pistil-
late inflorescencesin P. ovata and P. ferruginea
could be an adaptationto dispersalby bats.
Relations with Ants
Pouroumaformicarum and P. myrmecophila
aremyrmecophilous.These specieshave swollen
and saccatebasesof the petiole ("domatia"),usu-
ally occupiedby small ants. Accordingto Benson
(1985) these ants belong to genusAllomerusand
presumably eat food bodies, produced on the
inner surfaceof the domatia. Sometimes, as in
P. mollis subsp. triloba(collection Prance et al.
7907) and most collections of P. napoensis,ants
(of the genus Crematogaster)build tubularnests
on the leafy twigs and on the lower leaf surface.
The nest-buildingon the lowerleaf surface(only
if the main venation is subpalmate?)startsin the
axils of the main veins; then the nests look like
domatia. Lateron these nests are often extended
as tubes. A peculiaraspect is the use of the long,
ratherstiff hairs (occurringon various parts of
the plant)in the nest building,perhapsas a pro-
tective cover of the nests.
Distribution and Ecology
The distributionrangeof Pouroumais almost
the same as that of Coussapoa.Pouroumais con-
fined to rain forest areas, but is notably absent
in the West Indies and southern Mexico. Most
arelowlandspecies,foundat altitudesup to 1000
m, occasionally up to ca. 1500 m, and excep-
tionally (P. tomentosa in Bolivia and Peru) up
FloraNeotropica
to ca. 1800 or 1900 m. Some taxa(P. bolivarensis
and P. guianensis subsp. venezuelensis)appear
to be confinedto altitudesof about 1000 m (600-
1400 m). The unnamedcollections Gentryet al.
22866 and D. N. Smith et al. 4743 (see unnamed
collections 1, p. 193) may representa montane
taxon. Most species are components of non-in-
undatedforest, but P. acuminatais often found
in (occasionally?)inundated forest. Pourouma
(probably)needs light for germination and is
commonly found in placeswith more or less dis-
tinct tracesof previous disturbanceof the forest
(e.g., chablis) or in rather open forest (e.g., on
ratherpoor ? sandy soils) and secondaryforest.
A numberof species have a wide distribution.
All these species are representedin the Amazon
Basin and the Guianas. Two, P. bicolorand P.
minor, extend to CentralAmerica, and P. mel-
inonii to Panama.Pouroumamollis,P. velutina,
and P. guianensis also occur in eastern Brazil,
the latter in the coastal mountain rangeof Ven-
ezuela as well. Pourouma tomentosa, however,
is confinedto the Amazon Basin and the Guian-
as. Most of these wide-spreadspecies are very
variable in a confusing way. In five of these
species, two or more infraspecifictaxa are rec-
ognized.
The otherspeciesare much more uniformand
have much smaller or even very small distri-
bution ranges. Only one of the this group of
species (P. hirsutipetiolata,from northern and
westernColombia) is subdivided into two sub-
species(butin a more clearway than in the wide-
spreadspecies).This species and P. oraria(from
westernColombia)are the only ones not occur-
ringin the Amazon Basin or the Guianas.Of the
remainingspecies, three (P. bolivarensis,P. sau-
lensis, and P. stipulacea), are confined to the
Guianaregion,all probablywith very small dis-
tributionranges.The others are confinedto the
Amazon Basin, or almost so as P. acuminata,P.
cucura,and P. ovata (found in the Guineanpart
of Venezuela).
The present range of distributionof the very
uniformP. cecropiifoliamight be largelyanthro-
pogenous.
The Amazon Basin and the Guianasin all re-
spects representthe center of the genus. Three
centersof distributioncan be distinguished:
(a) The Upper Amazon Basin (down to ca.
60?W) with Pourouma acuminata, P. cecropi-
ifolia, P. cucura,P. cuspidata,P. elliptica,P. fer-
Distribution and Ecology
ruginea,P. formicarum, P. herrerensis,P. myr-
mecophila,P. napoensis,P. ovata,P. phaeotricha,
as well as P. bicolorsubsp. tessmannii,P. mollis
subsp. triloba,and P. tomentosa subspp. apicu-
lata, persecta, and tomentosa. A concentration
of taxa is found in the north-westernpartof this
region.
(b) The Guiana(or Guayana)region,the Low-
er Amazon Basin (Brazil:easternParaand Ama-
pa),includingan extensionto easternBrazil,with
the following taxa confined or distinctly con-
nected to this area: Pourouma bolivarensis,P.
saulensis,as well as P. bicolorsubsp. digitata,P.
melinoniisubsp.melinonii,P. mollis subsp.mol-
lis, P. tomentosasubsp.essequiboensisand subsp.
maroniensis,and P. velutina.
(c) The PacificCoastalregion (Colombiaand
Ecuador)to Panama, with extensions to Gua-
temalaand throughnorthernColombiato north-
westernVenezuela,with Pouroumahirsutipetio-
lata, P. oraria, P. bicolor subsp. chocoana and
subsp.scobina,and P. melinoniisubsp.glabrata.
The featuresof the East-Brazilianpopulations
of P. guianensis, P. mollis, and P. velutina,re-
semble those of the materialof the same species
in the Lower Amazon Basin and adjacentparts
of the Guianas.
Pouroumabicolor,P. melinonii,P. mollis and
P. tomentosahave one (or two) morphologically
distinctentities (subspecies)distinctlyconnected
with the Guiana (Lowland)region and the ad-
jacent part of the Amazon Basin. In P. bicolor
and P. melinonii one or two disjunctsubspecies
occur in the third center of the genus and in P.
guianensisa disjunctsubspeciesoccursin north-
ern Venezuela.
In the wide-spreadspecies the materialin the
Upper Amazon Basin is morphologicallymore
variable than in the other parts of their distri-
butionranges.Pouroumacucurasharesthis trait.
In most partsof the distributionrangeof Pou-
rouma the taxa (even those at the subspecific
level) can be recognized,and well-collectedma-
terial of adult specimens can readily be identi-
fied. In contrast,the situationin the Upper Am-
azon Basin, especially in its northwesternpart,
is ratherunclearwith regardto the delimitation
of subspecies (even species in some cases). In
several species (e.g., P. bicolor,P. mollis, P. to-
mentosa) morphologicallyaberrantcollections
have been made, e.g., in the valley of the Rio
Santiago(Peru,Amazonas).
115
In some cases (P. cucura and P. tomentosa
subsp.persecta)the presentdata suggestthe oc-
currenceof a disjunctionbetweenpopulationsin
the southwesternpart and those in the north-
westernpart of the Amazon Basin.
The genus appearsto be underrepresentedin
a zone from southeasternto northwesternPara
largelyconsistingof'mata de cip6' and north of
the Rio Solim6es of patchesof'campos de terra
firme'(Pires, 1973). A similar disjunctivegap is
found in Coussapoa(see p. 10).
Systematic Position of the Genus
Pourouma shows many morphologicalsimi-
laritiesto the Africangenus Myrianthus(with 7
species), revised by de Ruiter (1976). The sim-
ilaritiesare foundin the habit, the leaf shapeand
dimension and their infraspecificvariation,and
the (variable)lengthof the petiole. Furthermore,
the staminate inflorescencesshow similarities,
but the pistillate ones are different,as they are
capitatein Myrianthus,wherethe pistillateflow-
ers are sessileand clusteredin a singlehead. Both
generaaremacrospermousin contrastto the oth-
er genera of the Cecropiaceae.The two genera
also show strong ecological similarities and are
found in similartypes of habitat.These two gen-
era may also be regardednot only as genetically
relatedbut also as the most primitiveamongthe
Cecropiaceae.
The leaves of some taxa, Pourouma cecropi-
ifoliaandP. asperasubsp.digitata,resemblethose
of Cecropiabecause of the great number of in-
cisions, but the leaves are never peltate as they
always are in Cecropia. Leaves of some other
taxa resembleleaves of some Coussapoaspecies
(see p. 5).
Systematic Arrangement
of the Species
On the basis of overall similarities, several
groups of species can be recognized. Not all
species can be placed with certaintyinto one of
them, however, as the staminate inflorescences
arenot known(as in P. bolivarensis,P. saulensis,
and P. stipulacea).
One group of species is characterizedby hav-
ing the tepals of the staminatefloweralmost en-
tirely connate, forming an urceolateinfundibu-
liformperianthwith the filamentsof the stamens
116
longerthan the tepals. The staminateflowersare
usually clustered in distinct, terminal, globose
heads. The arachnoidindument often occurson
the lateral veins of the lamina beneath or also
on other parts, such as the stipules. The lamina
is usually smooth above. The leaves are entire
or palmatelyincised. The basal part of the leaf
marginis formed by the basal part of the basal
lateralvein in all (or most?) species. Pourouma
mollis, P. melinonii, P. hirsutipetiolata,P. to-
mentosa, P. herrerensis,and P. napoensiscom-
prises this group, as may P. acuminata and P.
stipulacea, of which the staminate flowers are
still unknown.
About equally well-definedis a groupformed
by Pouroumamyrmecophila,P.formicarum,and
P. phaeotricha,in which the laminae are mostly
scabrousabove. The staminateflowers,with (al-
most) free tepals and short filaments, are clus-
tered in distinct, terminal, globose heads. The
hairsof the tepals and the stigma are long (com-
pared with other species). The lamina is entire
or 3-lobed and the tendencytowardentireleaves
is strong.The base of the petiole of P. myrme-
cophila and P. formicarum is swollen and sac-
cate. These two species are myrmecophilous.
Lesswell-definedis a groupof speciesin which
the lamina is usually scabrousand/or has more
than 7 incisions. The staminateflowersare more
or less clustered, but not in globose heads. In
most of the species the lamina is incised. This
groupis formedby Pouroumaguianensis,P. as-
pera, P. velutina,P. cucura,P. cuspidata,and P.
cecropiifolia.
The rest of the species cannot be satisfactorily
grouped,exceptfor pairsof speciesshowingmore
or less distinct similarities, such as Pourouma
minor and P. bolivarensisor P. ovata and P.
saulensis.
Pouroumaovata and P. ferrugineaboth have
pistillateinflorescenceswith verylong peduncles.
But they are so differentin other features that
this similarityappearsnot to indicate common
origin, but rather a similar adaptation (to dis-
persalby bats?).
Uses
Most, if not all, Pouroumaspecies have edible
fruits. As far as known, only P. cecropiifoliais
cultivated as a fruit tree, however. The taste of
Flora Neotropica
the fleshy(mesocarp-like)innerlayerof the peri-
anthstronglyresemblesthat ofgrapes.P. cecropi-
ifolia (Amazon grape, uva, uvilla, imbauibado
vinho) is one of the taxa mentioned in "Under-
exploited Tropical Plants with Promising Eco-
nomic Value" (National Academy of Sciences,
Washington,D.C., 1975).P. cecropiifoliacan start
reproducing(thusformingfruits)a few yearsafter
planting,as small trees.OtherPouroumaspecies
may have fruits as tasty as those of P. cecropi-
ifolia, but the fleshy mesocarp-likelayer is usu-
ally thinnerand many of them become medium-
sized trees before fruiting. Pourouma cecropi-
ifolia fruits are used to make a sweet wine. For
a detailed study on the reproductionand prop-
erties of the fruits of P. cecropiifoliasee Falcao
and Lleras(1980).
Taxonomy
PouroumaAublet, Hist. pl. Guiane 2: 891. 1775;
Tr6cul, Ann. Sci. Nat. Bot., Ser. 3, 8: 100.
1847; Miquel in Martius,Fl. bras. 4(1): 122.
1853; Bentham in Bentham & Hooker, Gen.
pi. 3(1): 357. 1880; Baillon, Hist. pl. 6: 141.
1875-1876; Engler in Engler & Prantl, Nat.
Pflanzenfam.3(1): 95. 1889; Benoist, Arch.
Bot. Mem. 5: 29. 1931; Woodson & Schery,
Ann. MissouriBot. Gard.47: 165. 1960. Berg
in Lanjouw& Stoffers,Fl. Suriname5(1): 265.
1975. Burger,Fieldiana Bot. 40: 200. 1977;
Berg& van Heusden, Proc. Kon. Ned. Akad.
Wetensch., Ser. C., 91(2): 105. (1988). Type
species. PouroumaguianensisAublet.
Trees,terrestrial,often with stilt-roots.Leaves
in spirals, entire or palmately incised, margin
entire to subcrenate;stipules fused, fully am-
plexicaul.Inflorescencesin the leaf axils, mostly
in pairs, branched(or to subumbellatein pistil-
late inflorescences), mostly ebracteate, some-
times bracteate;staminateflowerssessile or ped-
icellate, tepals (3-)4, free or connate; stamens
(3-)4, mostly free, sometimes connate;pistillate
flowerspedicellate,perianthtubular;ovary free,
stigma(sub)peltateor sometimes-knob-like.Fruit
an achene, large, surroundedby the enlarged,
more or less fleshy perianthwith a purplish to
blackish or red-brownouter layer at full matu-
rity; seed without endosperm, embryo straight,
cotyledons thick, radicle short.
Pourouma 117

Key to the Species and Subspecies of Pourouma

1. Base of the petiole swollen.
2. Petiole 1-1.5 cm long ..................................................... 8. P. formicarum.
2. Petiole 8-27 cm long ..................................................... 7. P. myremcophila.
1. Base of the petiole not swollen.
3. Stipules(sub)persistent.
4. Laminasmooth above.
5. Stipuleswith dense, white, arachnoidhairs;Guyana. ...................... 14. P. stipulacea.
5. Stipuleswithout white, arachnoidhairs;Colombia .............................. 18. P. oraria.
4. Laminascabrousabove ....................................... 3e. P. bicolorsubsp. chocoana.
3. Stipulescaducous.
6. Laminamore or less scabrousabove.
7. Laminaentire.
8. Lateralveins 2 x 7-10 (in large leaves up to 13), the basal pair unbranched(or faintly
branched);lamina mostly elliptic to oblong or to subobovate.
9. Pluricellularhairson the leafytwigsdense;staminateflowersin globoseheads;pistillate
inflorescencewith 11-15 flowers. .......... ......................... 9. P. phaeotricha.
9. Pluricellularhairson the leafy twigs sparseor lacking;staminateflowersnot in globose
heads;pistillateinflorescencewith 2-8 flowers. ........................ 2. P. velutina.
8. Lateralveins 2 x 10-25, the basal pair branched;lamina mostly ovate.
10. Laminaabove yellowish-hirsuteto -hirtellousover the whole surface.
11. Staminate flower with connate tepals and filaments longer than the perianth;
(fruiting)perianthof pistillateflowervelutinous. . ............. 25. P. herrerensis.
11. Staminateflowerwith (almost)freetepalsand filamentsshorterthanthe perianth;
(fruiting)perianthof pistillateflowerhispidulousto hirtellous........ 5. P. cucura.
10. Laminaabove yellowish- to whitish-hirtellousonly on the main veins.
12. White, arachnoidhairs on the main veins of the lamina beneathand/or also on
the stipules, petioles, leafy twigs, and peduncles ...........................
.......................................... 13b. P. tomentosasubsp. apiculata.
12. White, arachnoidhairs only in the areoles or also on the smaller veins of the
lamina beneath.
13. Hairs on the lamina beneath(more or less) appressed. .......... 3. P. bicolor.
14. Stipuleshairy inside. ...................... 3a. P. bicolorsubsp. bicolor.
14. Stipulesglabrousinside. ............... 3b. P. bicolorsubsp. tessmannii.
13. Hairs on the lamina beneathat least partlypatent.
15. Stipulesinside denselyhairy;pluricellularhairson the leafytwigs sparse
or lacking;pistillateinflorescenceswith 2-8 flowers ........ 2. P. velutina.
15. Stipulesinside glabrous,or if with sparsehairs, then pluricellularhairs
on the leafy twigs usually(rather)dense;pistillateinflorescencesusually
with at least 10 flowers.
16. Pluricellular(brown)hairs sparseor lackingon the leaftytwigs and
petioles .. ............................... .......... 5. P. cucura.
16. Pluricellular(brown)hairs(rather)dense on the leafy twigs and the
petioles.
17. Arachnoidhairson the laminabeneathconfinedto the areoles;
tepalsof the staminateflowers(almost)free;apexof the pedicels
at fruitnot widened.. ....... la. P. guianensissubsp. guianensis.
17. Arachnoid hairs on the lamina beneath also on the smaller
veins; tepals of the staminateflowersconnate;apex of the ped-
icels at fruit widened (aberrantform from Peru)............
.................................. 10. P. mollis subsp. triloba.
7. Laminalobed to parted.
18. Stipuleshairy inside.
19. Laminawith the hairs beneathon the veins more or less appressedand the smaller
veins plane or only slightlyprominent. ............................... 3. P. bicolor.
20. Lamina(usually)5-7(-9)-fid to -parted.
21. Laminawith arachnoidindumentbeneath,coveringalso the almost plane,
smallerveins.
22. Stipules subpersistent;midsegment of the lamina oblong to elliptic;
leafy twigs often hirsute;Panamaand westernColombia ..........
......... ..................... ...... 3e. P. bicolorsubsp. chocoana.
118 Flora Neotropica

22. Stipules caducous;midsegment of the lamina lanceolate;leafy twigs

never hirsute;Guianasand LowerAmazon Basin (to southernVene-
zuela?). ................................. 3c. P. bicolorsubsp. digitata.
21. Laminawith arachnoidindumentbeneath(almost)confinedto the areoles,
the smallerveins more or less prominent;CentralAmericaand the Pacific
coastal region of Colombia, Ecuador,and northwestVenezuela(also Am-
azonian Peru?). ............................. 3d. P. bicolorsubsp. scobina.
20. Lamina(usually)3-lobed to -fid.
23. Laminawith arachnoidindumentbeneath,coveringalso the almost plane
smallerveins or with arachnoidindumentconfinedto the areoles,but then
the spots of white indumentdistinctlyseparatedby the thick smallerveins;
pistillateinflorescencesmostlywithup to 25(-35) flowers;Guianas,Amazon
Basin, to northernColombia. ................. 3a. P. bicolorsubsp. bicolor.
23. Laminawith arachnoidindumentbeneath(almost)confinedto the areoles,
the smallerveins more or less prominentand thin; pistillateinflorescences
with up to 50 flowers;CentralAmerica, the Pacific coastal region of Co-
lombia, Ecuador,and to northwesternVenezuela(also AmazonianPeru?).
........................................... 3d. P. bicolorsubsp. scobina.
19. Laminawith hairsbeneath(partly)patenton the veins, the smallerveins prominent.
24. Leafytwigs yellow-hirsuteand/or the whole uppersurfaceof the lamina hirtel-
lous; fruitingperianthrarelyscabrous;Upper Amazon Basinand northernVen-
ezuela. ..................................... .............. 1. P. guianensis.
25. Lamina 3-5-fid to -parted;leafy twigs and petioles with long yellow hairs;
stipulesalwayshairyinside;fruitingperianth2-2.5 cm long;northernVen-
ezuela. ............................ lb. P. guianensissubsp. venezuelensis.
25. Lamina entire, 5-7-fid to -parted,or if 3-5-fid to -parted,then the leafy
twigsand the petioleswithoutlong yellow hairs,the stipulesglabrousinside
and/or the fruitingperianth 1.2-2 cm long; Guiana region,easternBrazil,
Amazon Basin to easternColombia. ..... la. P. guianensissubsp. guianensis.
24. Leafytwigspaleyellow-hirtellous(to subtomentose);uppersurfaceof the lamina
only hirtellouson the main veins; fruitingperianthusually scabrous;Central
America,the Pacificcoastalregionof Colombiaand Ecuador(also Amazonian
Peru?). ................................ ...... 3d. P. bicolorsubsp. scobina.
18. Stipulesglabrousinside.
26. Laminayellow-hirsuteto -hirtellousabove on the whole surface.
27. Brown, pluricellularhairs on the leafy twigs and petioles sparse or lacking;
arachnoidhairs on the lamina beneathalso on the smallerveins.
28. Lamina 7-fid to -parted,usually longer than 30 cm; perianthof pistillate
flowerhirtellousto hirsute;filamentsconnate. ............ 24. P. napoensis.
28. Lamina at most 5-parted,usually up to 20 cm long; perianthof pistillate
flowerhispidulous;filamentsfree. ........................... 5. P. cucura.
27. Brown,pluricellularhairs on the leafy twigs and petioles (rather)dense and/or
the arachnoidhairs on the lamina beneath (almost) confined to the areoles.
.......................................... la. P. guianensissubsp. guianensis.
26. Lamina yellow-hirsuteto -hirtellous or whitish strigose above only on the main
veins.
29. Pluricellular(brown)hairs sparseor lackingon the leafy twigs and petioles.
30. Lamina 7-fid to -parted,usually longer than 30 cm; perianthof pistillate
flowerhirtellousto hirsute;filamentsconnate. ............ 24. P. napoensis.
30. Lamina at most 5-parted,usually up to 20 cm long; perianthof pistillate
flowerhispidulous;filamentsfree. ........................... 5. P. cucura.
29. Pluricellular(brown)hairs (rather)dense on the leafy twigs and petioles.
31. Laminawith more or less appressedhairs on the veins beneath. ........
........................................ 3b. P. bicolorsubsp. tessmannii.
31. Laminawith (partly)patent hairs on the veins .......................
...................................... la. P. guianensissubsp. guianensis.
6. Laminasmooth above.
32. Laminascabrousbeneath.
33. Stipulesglabrousinside. ..........................................6. P. cuspidata.
33. Stipuleshairy inside. ................................. 3a. P. bicolorsubsp. bicolor.
32. Laminasmooth beneath.
34. Lamina 7-11-parted(to -fid).
Pourouma 119

35. Leafy twigs hirsute, or if glabrousor puberulous,then the incisions reaching

down to the petiole and the segmentsoften petiolulate.
36. Laminahirsuteto hirtellousabove, at least on the main veins...........
...................................................... 24. P. napoensis.
36. Lamina(sub)glabrousabove. ............. 13c. P. tomentosasubsp.persecta.
35. Leafy twigs puberulous;incisions usually not entirelydown to the petiole and
leaf segmentswith a broaderbase.
37. Baseof the laminausuallydeeplycordate;fruitingperianth1.5-3.5 cm long;
Upper Amazon Basin to easternColombia. .............. 4. P. cecropiifolia.
37. Base of the lamina often truncateto shallowlycordate;fruitingperianthto
ca. 1.5cm long;GuianasandLowerAmazonBasin(to southernVenezuela?).
............................................ 3c. P. bicolorsubsp. digitata.
34. Laminaentire or at most 5-parted.
38. Petiole with a mixtureof minute whitish hairs,long yellowishhairs,and sparse
brown pluricellularhairs.
39. Staminateflowerswithconnatetepalsandfilamentslongerthanthe perianth;
(fruiting)perianthof pistillateflowervelutinous. ......... 25. P. herrerensis.
39. Staminateflowerswith (almost) free tepals and filamentsshorterthan the
perianth;(fruiting)perianthhispidulousto hirtellous. .......... 5. P. cucura.
38. Petiole with indumentotherwise.
40. Lamina 3-lobed to 5-parted.
41. Arachnoidhairs of indument confined to the lamina beneath in the
areolesor also the smallerveins.
42. Laminabeneathwith the hairs on the veins appressed.
43. Stipules hairy inside; tepals of the staminate flowersalmost
free.
44. Leafytwigs with appressedhairs. ............ 3. P. bicolor.
Repeat20-20 for subspecies;see also unnamedcollection
#5.
44. Leafytwigs with patenthairs. ............... 17. P. villosa.
43. Stipulesglabrousinside; tepals of the staminateflowerscon-
nate.
45. Leafy twigs sparsely appressed-puberulousto strigose.
...................................... 11. P. melinonii.
46. Base of the lamina truncateto subcordate(occasion-
ally deeplycordate);fruitingperianthusuallydensely
puberulousto subvelutinous;stamens with the fila-
ments (almost)free;Guianasand Amazon Basin. ..
.................. 1la. P. melinoniisubsp. melinonii.
46. Base of the lamina deeply cordate;fruitingperianth
almost glabrous;stamens with the filamentsmostly
connate, Panama,northernand westernColombia.
1lb. P. melinoniisubsp. glabrata.
45. Leafytwigs hirsute. . 12b. P. hirsutipetiolatasubsp. hispida.
42. Laminabeneathwith the hairs on the veins patent.
47. Leafytwigs with appressedhairs ........................
.......................... 1la. P. melinoniisubsp. melinonii.
47. Leafy twigs with patent hairs.
48. Basalpartof the marginof the laminaformedby the basal
lateralveins;tepalsof the staminateflowersforthe greater
partconnate;fruitingperianthvelutinous. .... 10. P. mollis.
49. Pistillateinflorescenceswith the flowersin two more
or less distinctclusters;Guianas,easternBrazil,Low-
er Amazon Basin. ........ 1Oa. P. mollis subsp. mollis.
49. Pistillateinflorescenceswith the flowersusuallymore
or less diffuselydistributed;UpperAmazonBasinand
easternColombia. ....... 1Ob.P. mollis subsp. triloba.
48. Basalpartof the marginof the laminaseparatedfrom the
basal lateralveins by mesophyll;tepals of the staminate
flowersfree or up to halfway connate; fruitingperianth
sparselypuberulous ....................... 17. P. villosa.
41. Arachnoidhairs also on the petiole, the main veins of the lamina be-
120 Flora Neotropica

neath, the leafy twigs, the stipules, the pedunclesand/or the perianth
of the pistillateflower.
50. Peduncleof the pistillateinflorescence32-48 cm long;tepalsof the
staminatefloweralmsot free ..................... 16. P. ferruginea.
50. Peduncleof the pistillateinflorescence2-13 cm long; tepals of the
staminateflowerconnate.
51. Lowerleaf surfacetomentoseto subvelutinous ...........
................................ lOa. P. mollis subsp. mollis.
51. Lower leaf surface appressed-puberulousto strigose on the
main veins. ............................... 13. P. tomentosa.
52. Stipuleshairy inside. ....13b. P. tomentosasubsp. apiculata.
52. Stipulesglabrousinside.
53. Leafytwigs yellow-hirsute(or glabrous). ..........
.................. 13c. P. tomentosasubsp.persecta.
53. Leafytwigs puberulousto hirtellous.
54. Base of the lamina deeply cordate. ...........
......... 13d. P. tomentosasubsp. essequiboensis.
54. Base of the lamina truncateto subcordate.
55. Apex of the laminaacuminate;headsof sta-
minateflowers2-3 mm in diameter;Guian-
as, Amapa, and near Manaus. ...........
....... 13e. P. tomentosasubsp. maroniensis.
55. Apex of the lamina usuallyrounded;heads
of staminateflowers 4-6 mm in diameter;
Upper Amazon Basin. ..................
......... 13a. P. tomentosasubsp. tomentosa.
40. Laminaentire.
56. Basal lateralveins branched.
57. Laminabroadestabove or in the middle.
58. Arachnoidhairs only on the lamina beneathin the areolesor
also on the smallerveins.
59. Leafy twigs and lamina beneath on the main veins with
appressedhairs. ...................... 22. P. bolivarensis.
59. Leafy twigs and lamina beneath on the main veins with
patenthairs.
60. Basal(most)partof the marginof the laminaformed
by the basallateralveins;tepalsof the staminateflow-
er for the greaterpart connate. .......... 10. P. mollis.
Repeat 48-48 for subspecies.
60. Basal (most) part of the marginof the lamina sepa-
ratedfromthe basallateralveins by mesophyll;tepals
of the staminateflowerfreeor up to halfwayconnate.
61. Stipuleshairy inside. .............. 17. P. villosa.
61. Stipulesglabrousinside. ......... 21. P. elliptica.
58. Arachnoid hairs also on the petiole, the main veins of the
laminabeneath,the leafytwigs,the stipules,the peduncleand/
or the perianthof the pistillateflower.....................
........................ 13a. P. tomentosasubsp. tomentosa.
57. Laminabroadestbelow the middle.
62. Laminawith ? patent hairs on the main veins beneath.
63. Basalpartof the marginof the laminaformedby the basal
lateralveins; tepalsof the staminateflowerfor the greater
part connate.
64. Leafytwigs with patenthairs. .......... 10. P. mollis.
Repeat 49-49 for subspecies;see also unnamedcol-
lection #2.
64. Leafytwigs with appressedhairs. ................
.................. 1 a. P. melinoniisubsp. melinonii.
63. Basalpartof the marginof the laminaseparatedfrom the
basal lateralveins by mesophyll;tepals of the staminate
flowerfree or up to halfwayconnate. ........ 17. P. villosa.
See also unnamedcollection #4.
62. Laminawith appressedhairs on the main veins beneath.
Pourouma 121
65. Arachnoid hairs confined to the areoles or also on the
smallerveins on the lamina beneath.
66. Stipulesdensely or sparselyhairyinside.
67. Staminateflowerswith connate tepals and fila-
ments longer than the perianth;(fruiting)peri-
anth of pistillate flower velutinous; stipules
sparselyhairyinside. .......... 25. P. herrerensis.
67. Staminateflowerswith (almost) free tepals, fil-
amentsshorterthan the perianth;(fruiting)peri-
anthhispidulous,scabrous;stipulesdenselyhairy
inside. .............. 3a. P. bicolorsubsp. bicolor.
See also unnamedcollections#1, #3, and #4.
66. Stipulesglabrousinside.
68. Leafy twigs sparsely appressed-puberulousto
strigose. ...................... 11. P. melinonii.
Repeat 46-46 for subspecies.
68. Leafytwigs hirsute ........................
..... 12a. P. hirsutipetiolatasubsp. hirsutipetiolata.
65. Arachnoidhairsalso on the petiole, the main veins of the
laminabeneath,the leafytwigs,the stipules,the peduncles
and/or the perianthof the pistillateflower.
69. Peduncleof the pistillateinflorescence32-48 cm long;
tepals of the staminatefloweralmost free. ........
.................................. 16. P. ferruginea.
69. Peduncleof the pistillateinflorescence2-13 cm long;
tepals of the staminateflowerconnate. ...........
.................................. 13. P. tomentosa.
70. Stipuleshairy inside........................
.............. 13b. P. tomentosasubsp. apiculata.
70. Stipulesglabrousinside.
71. Apexof the laminausuallyroundedto emar-
ginate, sometimes short-acuminate;heads
of the staminateinflorescences4-6 mm in
diameter;Upper Amazon Basin..........
......... 13a. P. tomentosasubsp. tomentosa.
71. Apex of the laminaacuminate;heads of the
staminateinflorescences2-3 mm in diam-
eter; Guianas, Amapa, and near Manaus.
....... 13e. P. tomentosasubsp. maroniensis.
56. Basallateralveins unbranched.
72. Stipules 1-3 cm long.
73. Leafytwigs white-puberulousto subglabrous.
74. Lateralveins 2 x 7-12; stipulesglabrousinside;peduncle
of the pistillateinflorescence3-5 cm long. . 19. P. saulensis.
74. Lateralveins 2 x 9-21; stipuleshairyinside;peduncleof
the pistillateinflorescenceup to 32 cm long. .. 20. P. ovata.
73. Leafytwigs yellow-sericeousto -hirtellous.... 9. P. phaeotricha.
72. Stipules3-13 cm long.
75. Lowersurfaceof the laminainitiallyentirelyor largelycovered
with white or brownish,arachnoidhairs;peduncleof the pis-
tillate inflorescenceup to ca. 50 cm long. ..... 16. P. ferruginea.
75. Lowersurfaceof the laminawith white arachnoidhairsin the
areolesonly (or sometimes extendingto the main veins); pe-
duncle of the pistillateinflorescenceup to ca. 10 cm long.
76. Lamina with dense yellow hairs on the main veins be-
neath.
77. Leafytwigs with very short and much longeryellow
hairs. ............................... 21. P. elliptica.
77. Leafytwigs with hairs of about equal length. ......
...................................... 23. P. minor.
76. Laminawith sparseand inconspicuoushairson the main
veins beneath.
78. Hairs on leafy twigs very short ..... 25. P. herrerensis.
78. Hairson leafytwigslong or absent. ... 15. P. acuminata.
122
1. PouroumaguianensisAublet,Hist. pl. Guiane
2: 892, t. 341. 1775; Miquel in Martius, Fl.
bras.4(1): 127. 1853; Berg,Fl. Suriname5(1):
267. 1975;Croat,FloraBarroColoradoIsland
360. 1978. Type. French Guiana. Sinnamary
R., withoutdate (Y),Aublets.n. (holotype,BM,
only photographsin NY and U seen;isotypes?,
LE, S).
Tree, up to 30 m tall. Leafy twigs 3-15 mm
thick, white-puberulous,white- to yellow-hirtel-
lous, yellow-(sub)velutinousor yellow-hirsute,
with dense to/or (very) sparse, brown, pluricel-
lularhairs;lenticelsratherconspicuous.Lamina
3-fid to 3-lobed, 3- or 5-fid (to -lobed or to -part-
ed), sometimes 5-7-fid to -partedca. 10-25(-45)
x 10-25(-45) cm, or entire(to 3-lobed),broadly
ovate to elliptic (to oblong), (4-)10-20 x (2-)7-
14 cm, subcoriaceousto chartaceous,apex acu-
minate, base cordate, the sinus rather shallow
and wide or deep and narrow,often with over-
lappinglobes,sometimestruncate,in entireleaves
to rounded(to subacute);uppersurfacescabrous,
on the main veins hirtellous or hirsute or the
whole surfacehirtellousto subvelutinous,lower
surface subtomentose to subvelutinous or hir-
tellous, or on the smaller veins to puberulous,
hairs on main veins often + appressed,white,
arachnoidhairs usually (almost)confinedto the
areoles;tertiaryvenationand reticulum+ prom-
inent (and conspicuous) beneath; lateral veins
(7-)12-26 pairs, basal pair branched;petiole 4-
25(-40) cm long, whitish-yellow-puberulousto
-hirtellous to -hirsute and with dense to (very)
sparse,brown,pluricellularhairs;stipules(2-)4-
15 cm long, caducous, outside whitish- (to pale
yellow-)subsericeousto -subhirsute to -subto-
mentose or to -subvelutinousor yellow-hirsute,
and with dense to (very) sparse, brown, pluri-
cellularhairs,insideglabrousor with sparse,white
hairs or with ratherdense, yellow hairs. Stami-
nate inflorescencesup to 20 cm long and 12 cm
wide; peduncle 4-6 cm long, peduncle and
branchespuberulousto hirtellousto (sub)hirsute
or to subvelutinous, often with dense, brown,
pluricellularhairs;flowers + clustered,but not
in (sub)globoseheads;tepalslanceolate,(almost)
free; filamentsshorterthan the tepals. Pistillate
inflorescencesin fruit up to 20 cm long and 12
cm wide; peduncle (3-)6-15(-20) cm long, in-
dument of the peduncleand branchessimilarto
that of the staminate inflorescence; flowers
FloraNeotropica
(8-)10-25; pedicel 0.2-0.5 cm, in fruit0.4-1 cm
long, perianth 2-4 mm long, subvelutinous,
sometimes sparsely puberulous,stigma subpel-
tate, 1.5-2 mm diam., shortly pilose. Fruiting
perianthpurple,(sub)ovoidto ellipsoid, 1.2-2 or
2-2.5 cm long, mostly (sub)velutinous, some-
times sparsely puberulous, scabrous or almost
glabrous.
This taxon can be difficultto distinguishfrom
P. bicolor.The two taxa have overlappingdis-
tributionsand areeach very variablewith, more-
over, quite an overlap of charactersand their
variation.Consideringthe variationin both taxa,
one would be inclinedto combine the two. How-
ever, as distinct morphologicalentities occur in
the same areas without having clear intermedi-
ates it is likelythat they aregeneticallyseparated;
only in a few cases do the collected specimens
show ? intermediatefeatures.
Pourouma guianensis can be distinguished
from P. bicolorin the followingways:
(a.) The indument of the inner surfaceof the
stipules. In subsp. guianensis the inner surface
of the stipulesis usuallyglabrousor with sparse,
white hairs. However, in some forms of subsp.
guianensisand in subsp. venezuelensisthe inner
surfaceof the stipules is generallycovered with
(ratherdense)yellow hairs,apparentlycorrelated
with the yellow-hirsuteindument on the leafy
twigs, the outer surfaceof the stipules,and other
parts.Pouroumabicolor(except for subsp. tess-
mannii.) is likewise characterizedby the occur-
rence of dense, whitish to yellow hairs on the
inner surfaceof the stipules, but in this species
(except for subsp. chocoana.)this indument co-
occurswith an indumenton the leafy twigs, pet-
ioles, and outer surfaceof the stipulesconsisting
of short, appressedhairs.
(b.) The indumenton the smallerveins of the
lamina beneath.In P. guianensisthis consists of
relatively long, patent hairs. In P. bicolor the
hairs on both the main veins and the smaller
veins is mostly appressed,only occasionallypa-
tent on the smallerveins and then mostly shorter
than in P. guianensis.Only in severalspecimens
of P. bicolorsubsp. scobina does the indument
resemblethat normallyoccurringin P. guianen-
sis.
(c.) The prominenceof the reticulumand oc-
currenceof the white-arachnoidindumenton the
lamina beneath. In P. guianensis the reticulum
is prominentand, moreover,conspicuousas the
Pourouma 123

white-arachnoidindumentis usuallyconfinedto PouroumascabraRusby, Bull. New York Bot. Gard.

the areoles.In P. bicolorthe reticulumis, in most 6: 498. 1910. Type. Bolivia. Pando:Santa Barbara,
30 Aug 1902 (9),R. S. Williams1560 (holotype,NY;
of the subspecies, merely slightly prominent to
isotype, US).
planeand usuallyinconspicuousas the arachnoid PouroumaradulaBenoist,Bull.Mus. Hist. Nat. (Paris)
indumentusuallyalso occurson the smallerveins. 28: 320. 1922; Woodson & Schery,Ann. Missouri
However, in a form of P. bicolorsubsp. bicolor, Bot. Gard. 47: 166, t. 58. 1960. Type. Colombia.
Without locality, without date (a), Triana860 (ho-
occurringin the westernpart of the rangeof the
lotype, P; isotypes, E, NY).
subspecies,the arachnoidindument is confined Pourouma subtrilobaRusby, Mem. New York Bot.
to the very smallareolessurroundedby relatively Gard.7: 232. 1927. Type. Bolivia. La Paz: Nr. Tu-
thick veinlets, and in P. bicolor subsp. scobina mapasa, 12 Oct 1921 (9), Cardenas1990 (holotype,
the arachnoidindumentis also more or less con- NY; isotypes, GH, US).
fined to the areoles, but in this subspecies this PouroumasubstrigosaMildbraed,Notizbl. Bot. Gart.
Berlin-Dahlem10:192. 1927;Macbride,Publ.Field
featureis combined with ? prominent smaller Mus. Nat. Hist., Bot. Ser., 13(2.2):293. 1937. Type.
veins. Peru. Loreto:Pongo de Manseriche,26 Nov 1924
In P. guianensistwo allopatricsubspeciescan (8), Tessmann 4642 (holotype, B, isotypes, B, G,
be recognized. NY).
PouroumamildbraedianaStandley,Publ. Field Mus.
Nat. Hist., Bot. Ser., 17: 183. 1937. Type. Brazil.
Amazonas:Mun. Sao Paulo de Olivenoa, nr. Pal-
Key to the Subspecies mares, 11 Sep-26 Oct 1936 (a), Krukoff8386 (ho-
of Pourouma guianensis lotype,NY; isotypes,A, BR, F, G, LE,MO, P, S, U,
US).
1. Lamina3-5-fidto -parted;leafytwigsandpeti-
oles with long yellow hairs;stipuleshairyinside; Leafy twigs pale to bright yellow-hirtellous to
fruitingperianth2-2.5 cm long; northernVen- -subvelutinous to -hirsute, brown, pluricellular
ezuela ................ b. subsp. venezuelensis. hairs dense to sparse. Lamina entire, 3-5-lobed
1. Lamina entire, 5-7-fid to -parted,or if 3-5-fid
to -parted,then the leafy twigs and the petioles to -parted, or sometimes 5-7-parted, base shal-
without long yellow hairs, the stipules glabrous lowly to deeply cordate or sometimes rounded;
inside,and/orthe fruitingperianth1.2-2 cm long; lateral veins mostly 10-16, sometimes up to 20,
Guianaregion,easternBrazil,Amazon Basin to or occasionally up to 25 pairs; stipules outside
easternColombia. ........ a. subsp. guianensis.
pale to bright yellow-hirtellous, -subvelutinous,
-subtomentose or -hirsute, inside glabrous or
la. Pourouma guianensis Aublet subsp. sometimes
hairy. Fruitingperianth 1.2-2 cm long,
guianensis. Fig. 59. mostly densely hairy, sometimes sparsely
hairy
Pouroumapalmata Poeppig & Endlicher,Nov. gen. or scabrous.
sp. pi. 2: 29, t. 141. 1838; Tr6cul, Ann. Sci. Nat. Distribution (Fig. 60). Throughout the Ama-
Bot., Ser. 3, 8: 104. 1847; Miquel in Martius, Fl. zon Basin, extending to eastern Colombia and
bras. 4(1): 126. 1853; Macbride,Publ. Field Mus.
Nat. Hist., Bot. Ser., 13(2.2):293. 1937. Type. Peru. the Guiana region; occurring disjunctly in east-
San Martin:Tocache, Aug 1830 (6 and 9), Poeppig ern Brazil (from Pernambuco to Santa Catarina);
s.n. or 1881 (holotype or syntypes, W, destroyed; mostly in non-inundated forest, sometimes in
lectotype, G (Qspecimen!),fragmentof 6 specimen periodically inundated (varzea) forest; mostly at
in F).
low altitudes, in Venezuela (Bolivar) up to ca.
PouroumaacutifloraTrecul, Ann. Sci. Nat. Bot., S&r.
3, 8: 105. in
1847; Miquel Martius, Fl. bras. 4(1): 1300 m, in Bolivia up to 1500 m.
126. 1853. Type. Brazil. Rio de Janeiro:Without
locality, 1839 (6), Guillemin1024 (holotype,P). Specimens examined. Colombia.Without locality,
PouroumacinerascensMartiusex Miquel in Martius, withoutdate (d),Triana860 (E, NY, P, type collection
Fl. bras. 4(1): 125, t. 37. 1853. Type. Brazil.Ama- of P. radula),withoutdate (6 and 9), Trianas.n. (E, P).
zonas:RioJapura,Maripi,withoutdate(6),Martius META: PuertoLopez,3 Aug 1944 (st),Liitle et al. 8423
s.n. (holotype,M;isotypes,M, U). (F);Villavicencio,Llano de San Martin,without date
PouroumaheterophyllaMartiusex Miquel in Martius, (9), Karstens.n. (GOET,LE).
Fl. bras. 4(1): 125. 1853. Type. Brazil. Amazonas: VENEZUELA. AMAZONAS: Rio Manavicke, Indian
Rio Japura,Dec 1819 (st, juv), Martiuss.n. (holo- village "Kalchi-Teri,"without date (juv.), Lizot s.n.
type, M; isotypes, M, U). (VEN);SierraParima, 1300 m, 18-23 May 1972 (st),
Pouroumafulginea Miquel in Martius,Fl. bras. 4(1): Steyermark106086 (NY); Parima Mts., nr. Simara-
129. 1853.Type.Brazil.AmazonBasin,withoutdate wochi, Rio Matacuni,18 Apr-23 May 1973 (d),Stey-
(Q),Martiuss.n. (holotype,M; isotype, U). ermark 107154 (F, U, VEN). BOLIVAR:Reserva Fo-
124 Flora Neotropica

JII'd~~~~~~~i

_ 4~, ,. ,e \

as t af +

POUROUMA einerascenx
Pourouma125
restal"LaParagua,"Rio Asa, Jun 1970(st,juv),Blanco
810 (VEN); slopes of QuebradaO-paru-ma,between
Santa Teresitade Kavanayenand Rio Pacairao(trib-
utary of Rio Mouak), alt. 1065-1220 m, 20-21 Nov
1944 (2), Steyermark60407 (F, VEN); 45 km N of
Tumeremo,Sierrade Nuria, 5-8 Feb 1961 (st), Stey-
ermark89134 (VEN); Rio Nichare (tributaryof Rio
Caura),nr. the confluence with Rio Cicuta, 25 Apr
1966 (2), Steyermark et al. 95711 (VEN). DELTA
AMACURO:E of Rio Grande, ENE of El Palmar, 29
Nov-18 Dec 1964 (2),MarcanoBerti471 (NY, VEN),
19 Jan 1965 (2),MarcanoBerti611 (F, NY, VEN);Rio
Amacuro,Venezuela-Guyanafrontier,Imataca Mts.,
downstreamfrom San Victor, 4 Nov 1960 (st), Stey-
ermark87307 (NY, VEN).
GUYANA. Kartabo region, Kartabo, 22 Jul 1920
(st, juv) Bailey 18 (GH), Bailey 19 (GH), Bailey 20
(GH), 20 Aug 1920 (st), Bailey 146 (GH), Bailey 147
(GH);Pomeroondistrict,Kamwatta,21 Sep 1921 (st),
Cruz 1175 (NY); between DemararaR. and Berbice
R., 19 Jul 1922 (9),de la Cruz1657 (F, MO, NY, US);
KurumaikabraCreek,EssequiboR., 22 Nov 1930 (2),
FD 1011 (FHO);TakutuCreekto PuruniR., Mazaruni
R., 25 Oct 1944 (6), Fanshawe2050 (=FD 4786) (A,
F, NY, U, US); Mazarunistation,forestnursery,1945
(st, juv), FD 4970 (U); FD 5011 (U, S); KanukaMts.,
Iramaipang,Nov 1948 (6), FD 5936 (NY); 11/2mile
along Bartica-Potarord., withoutdate (juv),FD 6914
(U); Tumatumari,18 Jun-8 Jul 1921 (st), Gleason174
(GH, NY); PakaraimaMts., just N of ParuimaMis-
sion, 19 Oct 1981 (9), Maas et al. 5858 (U); basin of
ShodikarCreek,tributaryof EssequiboR., 8-22 Jan
1938 (2), A. C. Smith 2845 (A, F, G, MO, NY, P, S,
U, US); MarudiMts., 02?15'N,59?10'W,12 Nov 1982
(2), Stofferset al. 311, 312 (U).
SURINAM. Raleighfalls,base of Voltzberg,Nature
Reserveon the CoppenameR., 18 Nov 1979 (9),Mori
et al. 8662 (NY, U).
FRENCHGUIANA. Withoutlocality,withoutdate
(2), Aublets.n. (BM, LE, ?S, type collection of P. gui-
anensis);nr. Saul3 Oct 1973 (9),Granvilleet al. B.5071
(U); Paul Isnardregion, 5 km of Citron, 12 Nov 1982
(2), Granville5284 (U); Oyapock R., Trois Sauts, 13
Jul 1774 (st), Grenand445 (CAY); Zidock Ville, 26
Aug 1976 (st), Grenand1352 (CAY);ComteR., Cacao,
25 Feb 1965 (2), Halle 1132 (U, P); nr. Sail, Jan 1975
(2), Moretti 102 (CAY); Comte R., ca. 60 km S of
Cayenne,26 Feb 1965 (9),Oldeman1179 (CAY);Saiil,
19 Oct 1971 (2), OldemanB.4108 (CAY, P, U); Kour-
ou, without date (9), (herb.)Richards.n. (P).
ECUADOR.NAPO:Rio Pucino,firstmajortributary
of Rio Aguarico, above bridge at Aguarico, 10 Feb
1974 (2), Gentry 9824 (U); Rio Napo, 8 km below
PuertoMishualli,25-27 Mar 1986 (2) Neill et al. 7082
(BG).
PERU. AMAZONAS:
6 Apr 1973 (2), Ancuash 178 (F,
GH, MO); Rio Santiago,nr. Caterpiza,9 Nov 1979
FIG. 59. Pouroumaguianensissubsp. guianensis.From Martius,Flora Brasiliensis4(1). 1853: tab. 37 (as
P. cinerascens).Leafytwig with staminateinflorescence,leaf apex, partof staminateinflorescence(28), diagram
of staminateflower(D), staminateflower(1).
(9), Huashikat 1216 (BG); Rio Cenepa, 30 Jan 1973
(6), Kayap268 (F, GH, MO), 21 Feb 1973 (9), Kayap
393 (F, GH); QuebradaHuampami, 5 Jul 1974 (9),
Kayap 1057 (F, GH, MO). HuANuco:Tingo Maria,
13 Aug 1940 (9),Asplund13020 (G, A, S, US); Pachita,
Codo de Pozuzo, 24 Oct 1982 (6), Foster 9373 (BG);
Distr. Churubamba,Rio Cayumba, 11 Sep 1936 (9),
Mexia 8172 (F, G, GB, GH, MO, NA, NY, S, U, UC,
US);betweenMonz6nand Rio Huallaga,withoutdate
(9), Weberbauer 3639 (G). LORETO: Prov.CoronelPor-
tillo, Dist. Calleria,rd. Pucallpa-Huanuco,km 34, 23
May 1968 (6), CastilloS. 11 (DUKE, F, P, US); Que-
brada Shanuce, above Yurimaguas,11 Jul 1972 (2),
Croat 18016 (F, MO, NY); Rio Napo, Caserio Bella
Vista,22 Sep 1972 (6),Croat20161 (C, DUKE, F, GH,
NA, NY); Rio Ucayali, Jenaro Herrera,7 Dec 1977
(st), Gentryet al. 21205 (MO,U); Prov.Maynas,Yana-
mono ExploramaTouristCamp, 15 Jul 1983 (st), Gen-
tryet al. 43066 (BG);Pongode Manseriche,nr. mouth
of Rio Santiago,2 Dec 1931 (d), Mexia 6201 (F, G,
GB, GH, MO, NY, S, U, UC, US), 26 Nov 1924 (8),
Tessmann4642 (B, G, NY, type collection of P. sub-
strigosa). MADREDEDIos: Parque Nacional del Manu,
Rio Manu,CochaCashuStation,Jul 1978 (st), Foster
et al. 6568 (F);Rio Tambopata,at mouthof Rio D'Or-
bigny, 1 and 2 Mar 1981 (st), Gentryet al. 31816 and
31933 (U). PASCO: Prov. Oxapampa,Vivero, Puerto
Bermudez,16 Jan 1983 (st), Gentryet al. 42029 (BG).
SANMARTIN: Tocache, Aug 1830 (6 and 9), Poeppig
s.n. (or 1881) (F, G, type collection of P. palmata);
MaynasAlto, withoutdate (9), Poeppigs.n. (B, OXF);
withoutlocality,withoutdate (6),Poeppigs.n. (LE,P);
Distr. Tocache, rd. to Almendras,7 Aug 1969 (2), J.
SchunkeV.3309 (F, G, US);rd.to Shunthe,E ofPuente
de Palo Blanco, 14 Jul 1974 (9), J. Schunke V. 7415
(MO, NY, U).
BRAZIL.Withoutlocality, 1839 (3),Guillemin1024
(P, type collection of P. acutiflora);"AmazonBasin,"
without locality,without date (9), Martiuss.n. (M, U,
type collection of P. fulginea). ACRE: Cruzeirodo Sul,
21 Feb 1976 (9),Marinho273 (IAN), 18 Feb 1976 (2),
Monteiroet al. 477 (INPA,MG);rd.Rio Branco-Porto
Acre, km 83, 10 Oct 1980 (9),Nelson 681 (BG) = Cid
et al. 2870 (U); rd. Sena Madureira-RioBranco,nr.
km 7, 30 Sep 1968 (6),Pranceet al. 7670 (GH, INPA,
M, NY, R, S, U, US). AMAZONAS:
Rio Antimari 30
Feb 1904 (st), Huber(MG) 4244 (MG);Rio Madeira,
nr. Calama,7 Nov 1931 (9), Krukoff1297 (G, NY, P,
S, U); nr. mouth of Rio Embira,14 Jan 1933 (9),Kru-
koff4817(A, F, G, MO,NY, S, U, US);Mun.Humaita,
Tres Casas, 14 Sep-11 Oct 1934 (o), Krukoff6238(B,
BR, F, LE,MO, NY, S, U, US);Sao Paulode Olivenca,
nr. Palmares11 Sep-26 Oct 1936 (9),Krukoff8369(A,
B, BR, F, G, LE, NY, P, U), (6),Krukoff8386(A, BR,
F, G, LE, MO, NY, P, S, U, US, type collection of P.
mildbraediana);Sao Paulo de Olivenca,CreekBelem,
26 Nov-11 Dec 1936 (2), Krukoff8652(A, BR, F, G,
 r~~~
II ~~~
I P. guiamnonsi

^%^_ 1 ^vf'^^/9^^ ^^\?^^^/ v *\ subsp. quirnnisL

i~subap.
vwwwcLensnir,

---------- - -- - '--- - - p l
---- ^l
-

Pourou-
FIG. 60. Distributionof Pouroumaguianensissubsp.guianensis,P. guianensissubsp.venezuelenyis',
ma velutina.
Pourouma127

LE, MO, NY, P, S, U, US); Mun. Coari, Rio Coari, da Divisa, 19 Dec 1964 (6), Hatschbach 12049 (F);
20 Apr 1976 (st),Magnagoet al. (INPA)58046 (INPA); aboveAntonia,18Jan 1966(9),Hatschbachet al. 13537
Rio Japura,Dec 1819 (st, juv), Martius s.n. (M, U, (F, NY, P, RB, U, US); Mun. Guaraquecaba,Rio Ban-
type collectionofP. heterophylla);Rio Japura,Mapiri, anal, 8 Dec 1970 (9), Hatschbach25765 (C, MO, NA,
withoutdate (8),Martiuss.n. (M, U, type collectionof S, UC); Rio do Costa, 9 Feb 1972 (9), Hatschbach
P. cinerascens);Manaus-Itacoatiarard., km 27, Re- 29129 (NA). PERNAMBUCO:Quipapa, Eng. Brejino 22
serva FlorestalDucke, 30 Jun 1976 (st), Mello (INPA) Mar 1967 (9), Andrade-Lima67-4983 (F). Rio DE
57509 (INPA);km 26,6 Jan 1977 (st),Nascimento303 JANEIRO: Tingua, 1 Oct 1946 (6), Brade et al. 18619
(INPA); Rio Canuma,Mun. Nova Olinda do Norte, (GUA, RB, U); Palmeiras, 13 Jan 1877 (st), Glaziou
Nov 1976 (st), Monteiro1295 (INPA);Rod. BR-174, 8935 (E, P);Serrados Orgaos,Tabuinha,21 Mar 1980
Reserva Biologica do INPA, 28 Sep 1976 (st), Mota (st), Glaziou12173 (P, mixed with Cecropiaglaziovii);
740 (INPA); Reserva Florestal Ducke, 11 Jan 1976 Serrada Estrella,nr. Mandioca,15 May 1880 (6), Gla-
(st),Nascimento387 (INPA),Nov 1972 (st),Rodrigues ziou 12174 (C, E, P), without date (6), Glaziou20408
9219 (INPA);rd. Humaitato Labrea,km 80, between (C, P); Matta do Teizeira Borges,nr. Gavea, 12 Dec
Rios Ipixuna and Itaparana,29 Jan 1970 (9), Prance 1928 (9), Pessoal do Horto Florestal650 (RB); Matta
et al. 3260 (F, INPA, NY, R, S, U, US); basin of Rio dos Tres Barros,nr. Gavea, 14 Mar 1927 (9), Pessoal
Demeni, nr. Totobi, 25 Feb 1969 (9), Prance et al. do Horto Florestal 651 (RB). RONDONIA:Mineraao
10233 (C, G, INPA, MO, NY, R, S, U, US); Manaus- CampoNovo, 100 km SW of Ariquemes,10 Oct 1979
P6rtoVelho rd. BR-319, km 175, betweenRio Tupana (9), Zarucchi 2744 (INPA, NY). RORAMA:Rio Catri-
andRio Igapo-aqu,11 Oct 1974 (9),Pranceet al. 22803 mani, 13 Feb 1975 (9), Pires 15088 (IAN); SerraTe-
(INPA, MO, U). BAHIA: Agua Preta, 13 Oct 1937 (a), pequem,west facingslopes, 1200 m, 17 Feb 1967 (9),
Bondars.n. (GUA); Rio das Contas, 3 Oct 1919 (8), Pranceet al. 4438 (G, INPA, M, NY, P, R, S, U, US);
Curran19 (C, F, MO, NY, US); rd. from Camaca- Indian trail Surucucu-Uaica,nr. Uaica airstrip,Rio
Canavieiro,Apr 1965 (9),Magalhaes733 (M);nr. Uru- Uraricoeira,2 Mar 1971 (9), Prance et al. 10836 (F,
cuca,4 Nov 1978 (9),Moriet al. 11046 (RB,U); Ilh6us, GH, INPA, M, NY, P, S, U); between Botamatatedi
Sep 1821 (9 and d), Riedel 2 or s.n. (LE, mixed with and Maita, 9 Feb 1971 (9), Prance et al. 13579 (F,
sterilespecimenof P. mollis), 10 Nov 1944 (9), Velloso INPA, NY, U, US). SANTACATARINA:Brusque, 15 Feb
723 (R). CEARA: Without locality, without date (st), 1951 (9), Klein 89 (US); nr. Blumeneau,18 Jan 1955
Allemaoet al. 446 (R);Serrade Baturit6,Sep 1897 (st), (9),Klein 110 (NY, S, UC, US); Brusque,15 Feb 1952
Huber 238 (MG). EspiRITOSANTO:9 Aug 1965 (9), (9), Veloso89 (RB);nr. Blumeneau,Nov 1888 (6), Ule
Belem 1471 (MG, NY, U); rd. BR-5 (BR-101),Morro 991 (F, HGB, P, US). SAOPAULO: Ubatuba, 8 Jan
Dantos, 8 Aug 1965 (9), Lanna Sobrinho1011 (FEE- 1985 (9), Gentryet al. 49346 (BG).
MA, U); nr. Linhares, 1 Oct 1971 (6), T. S. Santos BOLIVIA. LA PAZ:Nr. Tumapasa, 12 Oct 1921
2035 (MG, NY, U). MATOGROSSO:Rd. to Fontanilha, (9), Cardenas1990 (GH, NY, US, type collectionof P.
15 Sep 1976 (9), Gomeset al. 326 (INPA);Sarar6,10 subtriloba);16 km N ofCarrasco,1500 m, 31 Oct 1984
Aug 1978 (9),Pireset al. 16547 (MG);Nuicleode Hum- (9), Solomon et al. 12655 (BG). PANDO:Mukden, Jun-
boldt, (9), Roth 36 (INPA);Rio Juruena,road to Ari- Dec 1979 (9), Izawa 9 (U); Prov. N. Suarez,20 Aug
puana, km 5, 9 Jul 1977 (a), M. G. Silva et al. 3305 1978 (9), Mences 759 (MG);W bank of Rio Madeira,
(MG);Aripuana,rd. to airport,2 Jun 1979 ($), M. G. nr.Abufia,22 Jul 1970 (9),Pranceet al. 6253 (B, INPA,
Silva et al. 4750 (MG). MINASGERAIS:Viosa, 16 May NY, S, U, UC, US), 19 Nov 1968 (9), Prance et al.
1935 (9), Kuhlmann 2075 (US), 1936 (9), Kuhlmann 8670 (F, INPA, MG, NY, S, U); Prov. N. Suarez,San
(RB) 2200 (RB);Mun. Caratinga,Faz. Montes Claros, Pedro 13 Oct 1977 (9), Terceros1400 (INPA); Santa
23 Jan 1980 (9), Nsihimura 39 (FEEMA,GUA, U); Barbara,30 Aug 1902 (9), R. S. Williams1560 (NY,
Rio Novo, - (9), Schwacke 10393 (P). PARA:Serra US, type collection of P. scabra).
dos Carajas,SerraNorte, ca. 20 km N of Amza Ex-
plorationcamp, 17 Oct 1977 (9),Berg et al. 598 (MG, Vernacular names. Colombia. Meta: papa-
U); Cuiaba-Santaremrd., km 1183, 2 Feb 1977 (9), quillo. Venezuela. Bolivar: amia-yek; Delta
Berg et al. 772 (F, MG, MO, RB, S, U); Breves, Oct- Amacuro:chaparrode agua. Guyana. buruma,
Nov 1957 (a), Pires et al. 6663 (U); Rio Itacaiunas, Surinam. Manbospapaja. French
affluentof Rio Tocantins, SerraBuritirama,Jul 1970 sandpaper.
(9),Pireset al. 12570 (IAN);Capanema-Maranhaord. Guiana. Bois canon, kuluma, kalate, (Wayapi).
BR-22, km 96, 29 Oct 1965 (9),Pranceet al. 1777 (F, Peru.Amazonas:shuiya(Huambisa),sugkama(t)
GH, NY, P, S, U, US);GlebaBacaja,just below mouth shuiya, tsakap suiya (Huambisa);Huanuco:pa-
of Rio Bacaja21 Nov 1980 (st), Prance et al. 26375
payadel monte;Loreto:uvilla;San Martin:uvil-
(U); Santar6m,km 70 Estradado Palhao, Ramal do uvilla blanca. Brazil. Bahia:itararanga,tara-
Caetetu,16 Sep 1969 (a),M. Silva et al. 2623 (G, NY, la,
S, U, US). PARANA:P6rto de la 17
Cima, Jul 1911 (st), ranga blanca; Maranhao:kaymbe'y (Ka'apor);
Dusen 11942 (S); Jacarehy(=Sao Joao), 8 Oct 1915 Mato Grosso; imbaubarana, imbauba-torem;
(st), Duskn 17239 (F, NY, S), 22 Nov 1915 (9 and 6), Para:amapatior mapati;Parana:imbaubarana,
Dusen' 17345 (F, GH, NY, S); Morretes, 1904 (st),
de
Dusen s.n. (S); 14 Aug 1911 (st), Dusen s.n. (F, GH, pau jacu; Pernambuco:embaubada mata.
NY, S);Mun.Guaratuba,PedraBrancado Araraquara, This subspeciesis very variablein the density
3 Nov 1960 (9), Hatschbach7411 (R, U, US), 3 Nov and length of the indument. The materialfrom
1960 (6),Hatschbach7466 (US);Mun. Guarataba,Rio eastern Brazil often has 5-fid leaves, the indu-
128
ment is rathershort, and the leaf base is deeply
to shallowlycordate.Some collections from the
southernpart of the Amazon Basin (Bolivia and
adjacentparts of Brazil)usually have 5-7-fid to
-parted leaves, while elsewhere in the Amazon
Basin the leaves are 3-lobed to -fid or entire.
Most collectionsfrom the Guianaregionand the
Lower Amazon Basin have 3-fid leaves with a
shallowlycordate to truncateleaf base or entire
leaves; the indument is short, and the brown,
pluricellularhairson the twigs are usuallydense.
Material from the Upper Amazon Basin is
muchmorevariable.Some of the collectionshave
almost elliptic (to oblong)entireleaves. Another
groupof collectionshave entireleaves (oftenwith
relatively short petioles) or 3-fid to -parted,
sometimes 5-partedleaves with the lobes rather
close together, due to a smaller angle in which
the main basallateralveins departfrom the mid-
rib. These collections often have sparsepluricel-
lular hairs on the leafy twigs, and those with
entire leaves resemble P. velutina,from which
they differ in a greaternumber of lateral veins.
A thirdgroupof specimenshave 3-fid to -parted
leaves with a deeply cordateleaf base, often with
overlapping lobes. Several of these collections
have relatively long yellow hairs on the leafy
twigs and stipules, or have a ratherdensely hir-
tellous upper leaf surface. The collections with
long, yellow hairsusuallyhave stipulesin which
the inner surfaceis ratherdensely hairy, unlike
the other collections of subsp. guianensiswhich
have the inner surfaceof the stipulesglabrousor
sometimes with very sparse, white hairs. Three
collections(Croat20616, Krukoff4817and 8369)
have 3-fid,deeplycordateleaves andlong,yellow
hairson variousparts,includingthe innersurface
of the stipules and more numerouslateralveins
(up to 25 pairs),in medium-sizedleaves less than
1 cm fromeach other.Othercollectionsofsubsp.
guianensis(andcollectionsof subsp.venezuelen-
sis) have the lateralveins more than 1 cm from
each other.
The materialexamineddoes not provide clear
indicationsaboutthe natureof the variousforms
within the subspecies no more than about the
distribution of these forms. Material received
from an inventory of a forest in the Upper Rio
Moa region (Brazil,Acre) by Campbelland col-
laboratorsindicates the presenceof several dis-
Flora Neotropica
tinct forms of the subspecies in the same area
and the absence of intermediates between these
forms.
In several specimens from the Upper Amazon
Basin the leafy twigs, petiole, stipules, and lower
surface of the lamina are covered by a white
layer, actually the mycelium of some fungus which
apparently uses the pluricellular hairs as a sub-
strate.
lb. Pourouma guianensis Aublet subsp. vene-
zuelensis (Cuatrecasas) C. C. Berg & van Heus-
den, Proc. Kon. Ned. Akad. Wetensch., Ser.
C. 91(2): 106. 1988. Fig. 61.
PouroumavenezuelensisCuatrecasas,Bol. Soc. Venez.
Ci. Nat. 15: 107. 1954. Type. Venezuela.Aragua:
Parque Nacional Henri Pittier, 20 Nov 1953 (6),
Aristeguieta2005 (holotype, F, not seen; isotype,
VEN).
Leafy twigs yellow-hirsute, brown, pluricellu-
lar hairs dense. Lamina 3-5-fid to -parted, base
? deeply cordate; lateral veins 16-26 pairs; stip-
ules outside yellow-hirsute, inside with yellow
hairs. Fruiting perianth 2-2.5 cm long.
Distribution (Fig. 60). Northern Venezuela
(Aragua, Carabobo, Miranda, and Yaracuy),
coastal mountains; in forest at altitudes between
ca. 1000 and 1200 m.
Specimens examined. VENEZUELA. ARAGUA:
Par-
que Nacional "H. Pittier," 20 Nov 1953 (6), Ariste-
guieta2005 (VEN,type collectionof P. venezuelensis),
12 Mar 1964 (6),Ijjdsz-Madriz28 (VEN),24 Dec 1946
(9),Lasser2202 (VEN);RanchoGrande,30 Dec 1946
(6), Lasser 2267 (VEN); Guamitas, 5 Nov 1947 (6),
Pittieret al. 15641 (US, VEN); ParqueNacional "H.
Pittier,"6 Jan 1968 (st), R. F. Smith V-3321 (VEN);
Chorni, 39 Apr-1 May 1972 (2), Steyermarket al.
105863 (NY, VEN);AguaAmarilla, 12 Nov 1947 (6),
Tamayo 3370 (VEN), Rancho Grande,26 Mar 1938
(9), LI. Williams 9983 (A, F, GH, NY, US, VEN).
CARABOBO: Between Valencia and Campanero, 7 Mar
1860 (2), Fendler2420 (G); Cumbrede Valencia,Pt.
Cabello,without date (6), Karstens.n. (LE);E of Los
Tangues,S of Borburata,31 Mar 1966 (9).Steyermark
et al. 95378 (NY, VEN, U). MIRANDA: Guatopo, 19
Nov 1956(9),Bernardi5696 (VEN),Bernardis.n. (NY);
San Juan, 18 km SW of Cupira, 2-7 Sep 1977 (6),
Gonzalezet al. 1319 (BG);S of SantaCruz,Mar 1978
(2), Steyermarket al. 116884 (BG); 11 km SSE of El
Guapo,27-28 Mar 1978 (2), Steyermarket al. 116966
(U). YARACUY: N of Salom, 4 Mar 1982 (st), Liesner
et al. 12394 (U).
Pourouma 129

~ I I

V CRIA
DE AGRICULTUPA
MINISTERIO
IERDBAJiONACIONAL DF VENEZUFLA

FIG. 61. Pourouma guanenss Abl

subsp. Leafy t wigwith pistillate inflorescences (Steyermark et
ial. 105863).sden tr: a
RI%,?%t a.
Ce.: C,C, Blrg
D Utrecht r'. ...X'lr , A.
in-He
E,C.I. van usden ,,,. J.&M Cm"t

FIG. 61. Pouroumaguianensissubsp. venezuelensis.Leafytwig with pistillateinflorescences(Steyermarket

al. 105863).

Vernacularnames.Venezuela.Carabobo:tam- guished by smaller leaves and smaller fruiting

bor; Miranda:yagrumonegro. perianths.
This subspeciesis very uniformcomparedwith
subsp.guianensis.Some collections of the latter 2. PouroumavelutinaMartiusex Miquelin Mar-
subspecies from the Upper Amazon Basin re- tius, Fl. bras. 4(1): 130, t. 41. 1853; Berg, Fl.
semble subsp. venezuelensis, but are distin- Suriname5(1): 269. 1975. Type. Brazil.Para:
130 Flora Neotropica

Mouth of Rio Tocantins, Aug 1819 (6), Mar- widenedapex;perianth4-6 mm long, yellowish-
velutinous,at the apex with brown, pluricellular
tius s.n. (lectotype, M, chosen here; isolecto-
type, M). hairs; stigma peltate, 1.5-3 mm diam., with
Fig. 62.
sparse,yellow hairs and dense, brown, pluricel-
Pouroumasteyermarkii Standley& Cuatrecasas in lular hairs. Fruitingperianthblack, ovoid to el-
Cuatrecasas,FieldianaBot.28(1):210. 1951.Type. lipsoid or sometimes to oblongoid, 1.2-1.8 x
Venezuela. betweenRioKaruai
Bolivar:Ptari-tepui,
andfirstridgeaboveRio Karuai,28 Nov 1944(v), 0.6-1 cm, densely to sparselyhairy.
Steyermark 60665(holotype,F; isotype,VEN). Distribution (Fig. 60). Venezuela (Amazonas
and Bolivar), Surinam, French Guiana, Ama-
Tree, up to 15 m tall. Leafy twigs 2-6 mm zonianBrazil(Amapa,Amazonas,Mato Grosso,
thick, brownish- to yellowish- to whitish-seri- and Para)and Peru (Loreto),and easternBrazil
ceous to -puberulousand sometimeswith brown, (Bahia and Espirito Santo); in non-inundated
arachnoidhairs.Lamina entire,ellipticto ovate, forest, mostly at low altitudes, in Venezuelaup
rarelyto obovate, 6-30 x 2-20 cm, coriaceous to ca. 1200 m.
to subcoriaceousor sometimeschartaceous,apex
acuminate, base rounded to acute, sometimes Specimens examined. VENEZUELA. AMAZONAS:
truncateor subcordate;upper surfacescabrous, Atures, 20 Apr 1978 (9), Davidse et al. 15365 (MO,
U); Camani, 24 Apr 1971 (9), Foldats 114-1A (NY,
the main veins brown-tomentoseto -sericeous, VEN). BOLIVAR: Lower S-facingslopes of Ptari-tepui
sometimes with orange or orange-brown,pluri- betweenRio Karuaiand firstridgeabove Rio Karuai,
cellularhairs;lower surfacewhitish-sericeousto ca. 1200 m, 28 Nov 1944 (9) Steyermark60665 (F,
VEN, type collection of P. steyermarkii).
-velutinous,sometimeswith brown,pluricellular SURINAM. Sectie O, 11 Nov 1915 (st), BW 1368
hairs, white arachnoidhairs in the areoles and (U); Tibiti savanna, 20 Jan 1949 (st), Lanjouwet al.
on the smaller veins; lateral veins 7-13 pairs, 1761 (NY, U); Jodensavanne,MapaneCreek, 13 Jun
basal pair branchedor unbranched,tertiaryve- 1953 (st), Lindeman4057 (U); 1 Oct 1953 (st), Linde-
nation prominent beneath; petiole 1.5-20 cm man 4794 (U); UpperCoppenameR., 6 Aug 1954 (st),
Lindeman6418 (U).
long, yellow- to brown-sericeousto -tomentose, FRENCH GUIANA. St. Laurent,10 Oct 1956 (9),
and sometimes with brown, pluricellularhairs; BAFOG7562 (CAY,NY, P, U); Gregoire, 26 Jan 1972
stipules2-12 cm long, caducous,outside yellow- (9), Deward 138 (CAY, P, U); Maroni R., 1891 (6),
sericeous to -velutinous, and often with brown Gandoger19 (P); Orapu, 26 Oct 1970 (9), Oldeman
B.3786A (A, CAY, P, U); Acarouany,1858 ($), Sagot
to yellow, pluricellularhairs, inside yellow-se- 1163 (BR, F, G, GH, GOET,NY, P, S, U); Godebert,
riceous to -velutinous. Staminate inflorescences 20 Dec 1920 (6), Wachenheim271 (P).
up to 12 cm long and 8 cm wide; peduncle 2-7 PERU. LORETo: Rio Nanay, trail from Astoria to
cm long, peduncle and branchesyellowish-seri- Rio Mazan, 9 Jan 1976 (9), McDaniel et al. 20403
ceous to -velutinousor to -tomentose, and often (NA).
BRAZIL.AMAPA: Mun. Mazagao,75-80 km WSW
with dense, brown, pluricellularhairs;flowers of Macapa, 5-10 km SW of Rio Prato, 20 Dec 1984
sessile, diffusely distributedalong the ultimate (9),Daly et al. 3936 (BG);Clevelandia,2 Aug 1960 (6),
branches;tepals 1.3-1.8 mm long, freeor basally Westra 47303 (FHO, GH, MG, NY, U, UC, US).
AMAZONAS: Manaus, 9 Aug 1962 (6), Chagas (INPA)
connate,white-puberulous,filamentsshorterthan
the perianth,free.Pistillateinflorescencesin fruit 1655 (INPA,U); Rio Uatuma,Itapiranga,27 Aug 1979
(9), Cid 849 (U); Manaus,8 Jul 1933 (9 and 6), Ducke
up to 12 cm long and 7 cm wide; peduncle 2.5- (RB) 25247 (INPA, RB); Manaus-Itacoatiarard., km
7 cm long, peduncleand branchesyellowish- to 26, 4 Jan 1977 (st),Nascimento262 (INPA);trackfrom
brownish-sericeousto -velutinous,sometimesto Boca de Acre airstripto Monte Verde, 21 Sep 1966
-tomentose or to -puberulous,and with dense, (9),Pranceet al. 2469 (INPA, MO, NY, R, S, U, UC,
US);Manaus,Aleixord.,km 11, 2 May 1974(6),Prance
brown, pluricellularhairs;flowers 2-8, + dis- et al. 21011 (INPA,MG, MO, NY, S, U, US);Manaus,
tinctly in two clusters;pedicel up to 0.4 cm long, Binda Creekrd., 26 Jun 1961 (9 and 6), Rodrigueset
in fruit up to 1 cm, already at anthesis with a al. 2892 (INPA as 2092, MG, RB, U); Manaus Pas-

--9

FIG. 62. Pouroumavelutina.From Martius,FloraBrasiliensis4(1). 1853:tab. 41. Leafytwigwith staminate

inflorescences,diagramof staminateflower(D), partof staminateinflorescence(28), staminateflower(1), tepals
(4), stamens (7).
Pourouma 131

'
'~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~r
'~i "
,?il:~~~~~~~~~~

,^ > 3 \

POUROUMA vebliainas.
132
sarinhord., 14 Sep 1961 (9),Rodrigueset al. 3267 (U),
9 Aug 1962 (2),Rodrigueset al. 4585 (U). BAHIA:Road
fromRod. SaoJose-Buerarena to Una, 7 Jul 1980 (juv),
Berget al. 1143 (U); Una, 29 Aug 1978 (2), Mori et al.
11026 (MG, RB, U); between Vit6ria and Salvador,
without date (2), Sello s.n. (B). ESPIRITOSANTO:Rio
Doce, Velho-Pancasrd., 20 Sep 1930 (6),M. Kuhlmann
383 (RB);Linhares,11 Mar 1972 (st), Sucre8684 (U).
MATO GROSSO:Aripuana,28 Mar 1977 (st), Gomes et
al. 1021 (INPA), 12 Apr 1977 (st), Gomeset al. 1196
(INPA), without date (9), Rylands 15 (INPA); Mun.
Sinop, 6-9 km E of BR-163, on rd. to FazendaLon-
drina,24 Sep 1985 (9), Thomaset al. 4018 (BG);Mun.
Sinop-Colider,BR-080, ca. 91 km E of junction with
BR-163, SerraFormosa,3 Oct 1985 (9), Thomaset al.
4179 (BG); Mun. Vila Bela SantissimaTrinidade, 4
km S of borderofRond6nia, 3 Nov 1985 (9), Thomas
et al. 4810 (BG). PARA:Belem, 2 Jul 1914 (8), Ducke
(MG) 15350 (G, MG, P, US); Belem, 18 Jul 1899 (2),
Huber(MG) 1640 (G, MG, P, S, U); Gurupa(2), Ducke
(MG) 15935 (MG); Rio Xingfi, Vit6ria-Ponte Nova
rd., 7 Aug 1918 (9), Ducke (MG) 17167 (MG); B6a
Vista, 15 Oct 1897 (2), Guedes(MG) 1235 (G, MG, P,
S, U); Bel6m-Brasiliard., km 93, 19 Aug 1959 (6),M.
Kuhlmannet al. 52 (GUA, MG, R, S, US); mouth of
Rio Tocantins,Aug 1819 (8),Martiuss.n. (M, lectotype
collection of P. velutina);Jacuarary(=Jaguarari),21
Aug 1819 (8), Martiuss.n. (M); Bel6m, 12 Aug 1945
(9), Pires et al. 161 (RB); Pirelli, Jul 1958 (a), Pires
7015 (U); Belem, Sep-Oct 1961 (9), Pires 51882 (NY,
U, US);Ilhade Breu,5 Oct 1965 (9),Pranceet al. 1548
(F, GH, NY, P, S, U, US); Cuiaba-Santaremrd., km
163, 10 Nov 1977 (2), Prance et al. 25167 (F, INPA,
MO, RB, U), km 941, 13 Nov 1977 (9), Pranceet al.
25363 (MO, RB, S, U, US); Santar6m-Palhaord., km
35, 28 Aug 1969 (9),M. Silva et al. 2423 (F, GH, MG,
NY, S, U, US).
Vernacular names. Venezuela. Amazonas: cu-
cura. Surinam. Boroma or boroma ibeberobana
(Arawak), bospapaja, yarayara (Carib). French
Guiana. Bois canon, papaye apici (Paramaka).
Brazil. Amazonas and Mato Grosso: imbafibara-
na; Para: mapatirana.
This species can sometimes be very difficult
to distinguish from forms ofP. guianensis subsp.
guianensis with entire leaves and sparse pluri-
cellular hairs on the leafy twigs.
The lectotype is chosen from the two (syntype)
collections of Martius cited above.
3. Pourouma bicolor Martius, Syst. mat. med.
bras. 34. 1843; Miquel in Martius, Fl. bras.
Key to the Subspecies of Pourouma bicolor
1. Stipulesglabrousinside. ........................................................
1. Stipuleshairy inside.
2. Arachnoidhairs on the lower leaf surfaceconfinedto the areolesor almost so.
Flora Neotropica
4(1): 129, t. 39. 1853. Type. Colombia. Ama-
zonas:Rio Caqueta,PuertoMiraia, Jan 1820
(Q),Martiuss.n. (holotype, M; isotype, U).
Tree, up to 15 m tall. Leafy twigs 3-15 mm
thick, yellowish- to whitish-puberulous to
-(sub)sericeous,or yellow-hirsute,andwith sparse
to dense, brown, pluricellularhairs. Lamina en-
tire, ovate to subovate or elliptic to oblong, 5-
25 x 3-18 cm, 3-5-lobed to -parted, or 5-9-
parted,13-30(-42) x 13-30(-40) cm, coriaceous
to chartaceous,apex acuminate(to obtuse),base
(obtuseto) truncateto deeplycordate;uppersur-
face scabrous to scabridulous or sometimes
smooth, hairyon the main veins, puberulouson
the smaller veins, lower surfacesmooth, some-
times scabridulous,yellowish-appressed-puber-
ulous on the main veins, arachnoidhairs in the
areoles and on the smallerveins, or confinedto
the areoles; lateral veins (6-)12-25(-40) pairs,
basal pair branched, tertiary venation almost
plane beneath; petiole (2-)4-30(-46) cm long,
appressed-puberulousto -strigose, sometimes
yellowish-hirtellousto -(sub)tomentoseor yel-
low-hirsute;stipules 2-13.5(-25) cm long, out-
side yellowish-to whitish-appressed-puberulous
to -(sub)sericeousor sometimes yellowish-hir-
tellous to -(sub)tomentoseor yellow-hirsute,in-
side ? densely hairy, sometimes glabrous, ca-
ducous. Staminate inflorescencesup to 17 cm
long and 14 cm wide; peduncle 2.5-7 cm long,
yellow-puberulousand with dense (most dense
on the branches), dark-brown or red-brown,
pluricellularhairs;flowersusuallysessile, ? clus-
tered, but not in (sub)globoseheads; tepals 1-2
mm long, free or basally connate, (sparsely)pu-
berulous;filamentsshorterthan the tepals. Pis-
tillateinflorescencesin fruitup to 26 cm long and
14 cm wide; peduncle 1.5-14 cm long peduncle
and brancheswith indument similar to that of
the staminateinflorescences;flowers 6-40; ped-
icel 0.2-0.5 cm long, in fruit0.4-1 cm long;peri-
anth 2-4 mm long, scabrous;stigmapeltate, 1-2
mm in diam. Fruitingperianthpurpleto black,
ovoid to ellipsoid, ca. 1.5 cm long, scabrous(or
smooth), sparselyhairy.
In P. bicolorfive subspeciescan be recognized.
. subsp. tessmannii.
Pourouma 133
3. Smallerveins (reticulum)of the lamina beneathplane or almost so, the veinlets ratherthick and
the spots of arachnoidindument distinctly separated;lamina mostly entire or 3-lobed; Upper
Amazon Basin to northernColombia. ...................................... a. subsp. bicolor.
3. Smallerveins (reticulum)of the lamina beneathmore or less prominent,the spots of arachnoid
indument mostly not distinctly separated;lamina mostly 5-7-parted;CentralAmerica to the
Pacific coastal region of Colombia and Ecuadorand to northwesternVenezuela(also in Ama-
zonian Peru?). .................................... ....................... d. subsp.scobina.
2. Arachnoidhairs on the lower leaf surfacealso coveringthe smallerveins, usuallycoveringthe area
betweenthe paralleltertiaryveins.
4. Laminausually 5-7-parted;pistillateinflorescencesoften with up to 50 flowers.
5. Stipulessubpersistent;segmentsofthe laminarelativelybroad;leafytwigsoftenyellow-hirsute;
Panamaand westernColombia. ....................................... e. subsp. chocoana.
5. Stipulescaducous;segmentsof the laminarelativelynarrow;leafytwigsneverhirsute;Guianas
and the LowerAmazon Basin (to southernVenezuela?). ................... c. subsp. digitata.
4. Lamina usually entire or 3-lobed; pistillate inflorescencesmostly with up to 25(-35) flowers;
Guianaregion,Amazon Basin to northernColombia. ........................ a. subsp. bicolor.

3a. Pourouma bicolor Martius subsp. bicolor. Specimensexamined.COLOMBIA.Withoutlocal-

Fig. 63.
PouroumaasperaTr6cul,Ann. Sci. Nat. Bot., Ser. 3,
8: 102. 1847; Standley,Publ. Field Mus. Nat. Hist.,
Bot. Ser., 18(1): 391. 1937; Standley& Steyermark,
FieldianaBot. 24(4):52.1946. Type.FrenchGuiana.
Without locality, without date (9), Poiteau s.n. (ho-
lotype, P; isotypes, LE, P).
PouroumacrassivenosaMildbraed,Notizbl. Bot. Gart.
Berlin-Dahlem10:419.1928. Type. Bolivia.La Paz:
Nr. Mapiri,Sep 1907 (2), Buchtien2050 (holotype,
US; isotypes, B, NY).
PouroumalawranceiStandley,Publ. Field. Mus. Nat.
Hist., Bot. Ser., 17: 183. 1937. Type. Colombia.Bo-
yaca: El Humbo, 31 Mar 1933 (2), Lawrance727
(holotype,F; isotypes, A, E, G, MO, S, UC).
PouroumaschultesiiCuatrecasas,Caldasia7: 303. 1956.
Type. Colombia.Amazonas-Vaup6s:Rio Apaporis,
Jinogoje,25 Sep 1952 (2), Schultes& Cabrera17615
(holotype,F).
PouroumacamaratanaCuatrecasas,Acta Bot. Venez.
2(5-8): 202.1967. Type.Venezuela.Bolivar:Auyan-
tepui,above valley of Camaratana,18 May 1964 (2),
Steyermark 94810 (holotype, US; isotypes, NY,
VEN).
Leafy twigs appressed-puberulous to strigose.
Lamina entire to 3-lobed, sometimes 3-fid, oc-
casionally (in subjuvenile material?) 5-parted;
upper surface scabrous, sometimes smooth, low-
er surface smooth, sometimes scabridulous, the
arachnoid hairs in the areoles and on the smaller
veins or confined to the areoles, but then the
smaller veins relatively thick and the spots of
arachnoid hairs clearly separated; stipules inside
glabrous, caducous. Pistillate inflorescences
mostly with 3-25 flowers.
Distribution (Fig. 68). Throughout the Ama-
zon Basin and the Guiana region, extending to
northern Colombia; in non-inundated forest at
low altitudes, in Bolivia up to ca. 900 m.
ity, without date (2), Triana 861 (E, P). AMAZONAS:
Rio Caqueta,PuertoMirafia,1820 (2), Martiuss.n. (M,
U type collections of P. bicolor) (M, U).
AMAZONAS-VAUPES: Rio Apaporis, Jinogoj6, 25 Sep
1952 (2), Schultes& Cabrera17615 (F, type collection
*of P. schultesii). ANTIOQUIA: Confluence of Rios Ite
and Tamarinto Rio Cimatarra,ca. 38 km W of Bar-
rancabermeja,24 Nov 1967 (6), Bruijn 1497 (F, M,
MO, NY, S, U, US, VEN), 26 Nov 1967 (6), Bruijn
1514 (F, M, MO, NY, S, U, US, VEN), (st), Bruijn
1516 (F, M, MO, NY, S, U, US, VEN), (8),Bruijnl517
(F, M, MO, NY, S, U, US, VEN), 27 Nov 1967 (6),
Bruijn 1526 (F, MO, NY, S, U, US, VEN). BOYACA:
El Humbo, 31 Mar 1933 (6), Lawrance727 (A, E, F,
G, MO, S, UC, type collectionof P. lawrancei).META:
Footpath between Rio Giiejar and Cana Guapayita,
CafnoYerli, 20-28 Dec 1950 (2), Idroboet al. 786 (F,
MO,NY, US);Villavicencio(2), Karstens.n. (LE);Sier-
ra de la Macarena,CafnoCiervo, 12 Jan 1950 (2), Phil-
ipson et al. 2086 (S, US). PUTUMAYO: Santa Rosa on
Rio Guamfies, 26 Nov 1966 (2), Pinkley 556 (S).
SANTANDER: Region Carare-Op6n (Capote), nr. Rio
Magdalena, 6 Jul 1968 (9), Garcia et al. 9 (US). Vi-
CHADA:3 km S ofLinea Roja,ParqueNacionalNaturel
"El Tuparro,"14 Mar 1985 (2), Zarucchiet al. 3720
(BG).
VENEZUELA. AMAZONAS:Reserva Forestal "El
Sipapo,"Rio Sipapo,May 1971 (2), Blanco 1149 (NY,
US, VEN);Rio Marawinuma,nr. Cerrode La Neblina
Base Camp, 6 Feb 1984 (st), Liesner 15681 (BG), 28
Feb 1984 (9), Liesner 16299 (BG); Depto. Atures, N
side of Rio Cataniapo,45 km SE of PuertoAyacucho,
13 May 1980 (2), Steyermarket al. 122390 (U); Depto.
Atabapo,Rio Cunucunuma,betweenCerroDuida and
Cerro Huachamacahi,Jan-Feb 1982 (9), Steyermark
et al. 12585 (BG);Rio Marawinuma,nr. Cerrode La
Neblina Base Camp, 5 May 1984 (6), Thomas 3370
(BG). BOLIVAR: Rio Chizca, Uriman, 5-8 Sep 1953
(st),Bernardi910 (NY, VEN);Rio Hacha,28 Dec 1955
(2), Bernardi2657 (G, NY,); slopes ofQuebradaO-po-
ru-ma,betweenSantaTeresitade Kavanayenand Rio
Pacairao(tributaryof Rio Mouak), 20-21 Nov 1944
(st), Steyermark60401 (F); lower SW slopes of Chi-
manta-tepui(Torono-tepui),nr. Rio Tirica, 24 May
134 Flora Neotropica

!ie.
z
'.
~.~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~:'
?I
' ' ~ ~ ~
r-? ..
*~~~.

~'~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~.

.,~~~~~~~~ :

$ Y~~j-'"'~2-'"-...

.. . ~~~~~~~~~~~~~~~~~~~~~~~~~~~~~
. : ,,-
i
~. ~~~~~~~~

~
ii!:..-~~

-P~~~~~~~~~~~~~~~~~~,~
,..~

?f
:,, ,

POURi~A biol.
Pourouma 135

1953 (st), Steyermark75540 (F, NY, VEN); Sierrade ReservaFlorestalDucke,Nov 1972(st),Rodrigues9206

Lema, lowland between Rio Chicanan and Rio Ay- (INPA).MATOGROSSO: Aripuana,21 Feb-2 Aug 1977
aiche, betweenPuertaLemaand base of the Sierra,24 (2 and st), Gomeset al 676, 786, 843, 966, 979, 1052,
Aug 1961 (9), Steyermark89457 (NY, VEN);frontier 1072, 1079, 1088, 1123, 1176, 2399 (INPA); Ari-
Venezuela-Brazil,NE ofSerraniaPia-soi, 5-6 Jan 1962 puana,nr. HumboldtCentre,rd. to Rio Juruena,8 Oct
(st), Steyermark90644 (VEN); Auyan-tepui, above 1973 (6),Pranceet al. 18250 (INPA, NY, U), 30 Nov
valley of Camarata,alt. 1500-1530 m, 18 May 1964 1978 (2), Rylands 52 (INPA). PARA:Rios Mojfi and
(9), Steyermark94180 (NY, US, VEN, type collection Acara, S of Belem, ca. 7 km N of Mojfi, 3 Jun 1969
of P. camaratana);Rio Caura,nr. Salto Para, 15-17 (6),Austinet al. 4140 (MO);Faro,6 Jan 1920 (2), Ducke
Jan 1977 (9), Steyermarket al. 113096 (U). (RB) 13057 (RB);betweenRio Pacajaand Rio Muira-
SURINAM. ForestReserveZanderij1, 24 Oct 1917 piranga,SW of Ilha de Breu, 20 Sep 1965 (2), Prance
(9),BW3376 (MO,U); Watramiri,8 Oct 1918 (9),BW et al. 1397 (NY, US); BR-22, km 64, 25 Aug 1964 (6),
4033 (NY, U), 13 Dec 1920 (9), BW 5036 (U); Sectie Prance et al. 58862 (F, GH, NY, S, U, US); Belem
0, Feb 1943 (9), Stahel (Woodherb.Sur.) 166 (A, F, (IAN) 17 Oct 1963 (2), N. T. Silva 57832 (NY, U, US).
NY, U, UC). RONDONIA: FortePrincipeda Beira,Estradado Igarape
FRENCH GUIANA. S of Saul, 12 Feb (9), Leeu- da Viiva, 5 Jan 1962 (2), Rodrigueset a. 4237 (U).
wenberg11784 (CAY); without locality, (9), Poiteau RORAIMA: Indiantrail from SurucucuW to Uaica be-
s.n. (LE, P, type collection of P. aspera). tween Botamatediand Maitf, 10 Feb 1971 (2), Prance
ECUADOR.NAPO:6 km SE of Los Sachas, 14 Apr et al. 13582A (INPA, F, GH, M, NY, P, S, U).
1985 (9),Baker6017 (BG);5 km N of Coca, on Coca- BOLIVIA.LAPAZ:Nr. Mapiri,Sep 1907 (2), Buch-
Lago Agrio rd., 16 Mar 1980 (st), Brandbygeet al. tien 2050 (B, NY, US, type collectionof P. crassiveno-
30194 (AAU); El Chuncho, Estaci6n INIAP, 2 Oct sa), Sep 1907 (6),Buchtien2122 (NY, US), 12-30 Sep
1987 (st), Palacios 2059 (BG). 1939 (9), Krukoff10867(A, F, G, MO, NY, S, U, UC,
PERU. AMAZONAS: Rio Santiago, nr. Caterpiza, 14 US), Sep 1939 (8), Krukoff10874 (F, NY), 8 Oct-15
Nov 1979 (9), Huashikat1241 (U), 28 Nov 1979 (2), Nov 1939 (2), Krukoff11254 (F, NY).
Rio Mom6n, 4 Sep 1972 (9),
Junqui 149 (U). LORETO:
Croat 19997 (MO);Maynas,Yaguasyacu,tributaryof Vernacular names. Colombia. Antioquia-Bo-
Rio Ampiyacu, below Borro Indian village of Brilla livar: cirpe (or sirpe) hembra, cirpe (or sirpe)
Nueva, 7 Nov 1977 (9), Gentryet al. 20356 (MO);Rio
macho; Boyaca: cormi; Meta: caimar6n de mico.
Ucayali, JenaroHerrera,7 Dec 1977 (9), Gentryet al.
21193 (MO, U); Rio Javari, between Rio CuraqaR. Putumayo: otsepacho tsaha (Kofan). Venezuela.
and Tambaqui,28 Oct 1976 (9), Prance et al. 24188 Amazonas: cucura or cucure, sadha'fhi; Bolivar:
(INPA, MG, U). cay-bari-cay, cay-i-wari-cay-yek, sarasara. Suri-
BRAZIL. Without locality (9), Burchel9792 (GH, nam. Boroma (Arawak), pourouma or puruma
LE,P, underthe same numberP. mollissubsp.mollis).
ACRE:Upper Rio Moa, FazendaArizona, 10-16 Oct (Carib). Peru. Amazonas: pauishuina, tsakap sui-
1985 (st), Campbellet al. 6650 (BG); Rio Jurua,nr. ya (Huambisa). Brazil. Acre: imbafbarana.
Cruzeirodo Sul, 22 Oct 1966 (9), Prance et al. 2754 Amazonas: imbafiba; Mato Grosso: imbafibar-
(INPA, MG, U). AMAPA:Rio Oiapoque, 1-3 km N of ana, tamaoquare; Para: garguaba, mapatirana.
CachoeiraTres Saltos, 12 Sep 1960 (6), Irwin et al.
48192 (GH, M, NY, P, U, US). AMAZONAS: 27 Sep Bolivia. La Paz: uva menueda.
1973 (8), Berg et al. P. 18152 (INPA, MO, NY, P, U); In and towards the eastern part of the species
Manaus-Caracarai rd., km 148, 26 Sep 1973 (6),Bisby range (from Colombia to Bolivia) the arachnoid
et al. P.18118 (F, INPA, MO, NY, S, U, US); Rio hairs on the lower leaf surface are confined to the
Negro,Uaupes, 22 Jan 1960 (9),Cavalcante777 (MG); areoles in combination with an increase of the
Serrade Neblina, Rio Cauaburi,CampTucano, 5 Dec
1965 (9),Maguireet al. 60333 (F, MG, NY, P, U, US); thickness of the smaller veins, causing distinct
BenjaminConstant, 16 Oct 1956 (9), Meyer Drees 9 separation of the "islets" of arachnoid indument.
(INPA, U); Manaus,Reserva FlorestalDucke, 6 Dec The transition of the normal form of subsp. bi-
1976 (2), Nascimentoet al. (INPA)66338) (INPA);Rio color, with the arachnoid indument extending
Negro, Uaup6s, 1 May 1947 (9), Pires 502 (NY); Rio over the relatively thin smaller veins to the form
Negro, Sao Gabriel, 1 May 1947 (6), Pires 587 (NY);
Tapuruquara,rd. to airport, 17 Oct 1917 (9), Prance de scribed above, is rather gradual. Both forms
et al. 15366 (INPA, M, MO, NY, S, U, US); Manaus, may occur in the same region. In the eastern part

FIG. 63. Pouroumabicolorsubsp. bicolor.From Martius,Flora Brasiliensis,4(1). 1853: tab. 39, Leafytwig
with fruitingpistillateinflorescence,fruitingperianth(17), fruitingperianthand fruit(1711),pericarp(43), fruit
(19), pericarpand seed (21), seed (19).
136 FloraNeotropica

?3X3l cm t. '- ?

A 3

FG.64. 94).'ur inflorescence (Tunqui

Pourouma bicolor subsp. tessmanni. Leafy twigs with pistillate
(! L ..' i. _ ....... '

i-" .':..
Pourouma 137

of the speciesrangethe upperleaf surfaceis often

to obovate, base truncateto shallowlycordateor
smooth. See unnamed collection #1 (p. 193). sometimes deeply cordate; upper surface sca-
brousto scabridulous,sometimes smooth, lower
3b. PouroumabicolorMartiussubsp.tessmannii surfacewith appressedhairs on the main veins,
arachnoidhairsin the areolesand on the smaller
(Mildbraed)C. C. Berg& van Heusden, Proc.
Kon. Ned. Akad.Wetensch.,Ser.C 91(2): 106. veins; smallerveins (almost)plane:stipulesout-
1988. side with appressedhairs,inside hairy,caducous.
Fig. 64
Pistillate inflorescences mostly with 10-50 flow-
PouroumatessmanniiMildbraed,Notzibl. Bot. Gart. ers.
Berlin-Dahlem10: 192. 1927. Type. Peru. Loreto: Distribution (Fig. 68). Guyana, Surinam,
Pongo de Manseriche,8 Oct 1924 (6), Tessmann French Guiana, and Brazil (Amapa and Parf),
4236 (holotype,B; isotypes, NY, S, US).
and in southeastern
probably in non-
Venezuela;
Leafy twigs appressed-puberulousto strigose. inundatedforest at low altitudes.
Lamina entire or occasionally 3-lobed to -fid; Specimens exmained. VENEZUELA. AMAZONAS:
upper surface scabrous, lower surface smooth, Nr. Cerrode La NeblinaBase Camp, 19 Apr 1984 (st),
appressed hairs on the main veins, arachnoid Gentryet al 46752 (BG).
hairs in the areoles and on the smaller veins; GUYANA.NW slopes of KanukuMts., drainageof
Moku-MokuCreek,Takututributary,31 Mar-16 Apr
smallerveins (almost)plane;stipulesoutsidewith 1938 (2), A. C. Smith 3564 (B, F, LE, MO, NY, P, S,
appressedhairs, inside glabrous,caducous.Pis- U, US).
tillate inflorescences mostly with 3-16 flowers. SURINAM. Watramiri,9 Jun 1916 (st), BW 2030
Distribution (Fig. 68). Peru (Amazonas and (U); Kaboeri,30 Oct 1920,B W 4866 (U); Brownsberg,
4 Oct 1923 (S), BW 6292 (U); Distr. Brokopondo,2
Loreto);in non-inundatedforest. km S of Gansee,27 Apr 1964 (st),Donselaar1262 (U);
Nassau Mts., at km 0.2 on terraceof MarowijneR.,
Specimensexamined.PERU. AMAZONAS: Rio San- 16 Feb 1949 (st), Lanjouwet al. 2213 (F, NY, RB, U);
tiago, 65 km N of Teniente Pinglo, QuebradaCater- Distr. Saramacca,nr. Groningen, 19 Feb 1954 (st),
piza, 13 Oct 1979 (2), Huashikat912 (U), 30 Jan 1980 Lindeman5468 (U); Rikanau,nr. Moengo,6 Jun 1954
(9), Huashikat1872 (U); Rio Santiago,nr. La Paz, 12
Nov 1979 (2), Tunqui7 (BG);Rio Santiago,Quebrada (st), Lindeman 6123 (U); S of Moengotapoe, 14 Jun
1954 (st), Lindeman6166 (U); Jodensavanne-Mapane
Caterpiza, 17 Nov 1979 (6 and 2), Tunqui 94 (U). Creek area, 10 Dec 1954 (st), Lindeman 6765 (U);
HuANuco:Prov. Leoncio Prado, 80 km NE of Tingo Distr. Nickerie, area of KabaleboDam project,4 km
Maria,nr. La Divisoria, 18 Jan 1976 (2), Gentryet al. NW of rd., km 39.5, 4 Nov 1981 (st), Lindeman& de
1604 (DUKE, MO, NY). LORETO: 22 km S of km 86 Roon 750 (U); Lely Mts., E rd. on plateau 1, 29 Sep
on Pucallpa-TingoMariahwy., 11 Feb 1981(2),Gentry 1976 (juv),Lindeman& Stofferset al. 539 (U), 22 Jan
et al. 31197 (U); Pongo de Manseriche,nr. mouth of 1970 (2), Oldenburgeret al. 1120 (U); Sipaliwiniarea,
Rio Santiago,8 Oct 1924 (d), Tessmann4236 (B, NY, Brazilianfrontier,2.5 km S of SipaliwiniR., Feb 1970
S, US, type collection of P. tessmannii). et al. 1406 (U);WinnanaCreek,Upper
(st), Oldeburger
KabouirCreek,CorantyneR., 20 May 1960(st),Schulz
Vernacularnames. Peru. Amazonas: shuiya (LBB)8316 (U); Section 0, ca. 5?N, 55?W,Nov 1942
(Huambisa),tanta shuiya, tentarshuina(Huam- (a),Stahel 123A (U), Dec 1942 (S),Stahel (Woodherb.
bisa). Sur.) 166B (A, NY, U, UC).
This subspeciesis similar to subsp. bicolorin FRENCHGUIANA:Cayenne,1 Dec 1955 (2),BAF-
OG (=Bena) 1050 (CAY, F, U); upper Oyapock R.,
most featuresbut the stipulesareglabrousinside. Zidockville, 6 Sep 1977 (st), Grenand1442 (CAY,U);
Saul, 8 Dec. 1982 (2), Mori et al. 15350 (BG);deserted
3c. Pourouma bicolor Martius subsp. digitata villageof EauClaire,about 7 km N of Belizon, 12 Feb
1966 (9), Oldeman 2030 (CAY); Yaroupi R., 8 km
(Trecul)C. C. Berg& van Heusden,Proc.Kon. downstreamof Saut Tainoua, 18 Apr 1970 (st), Olde-
Ned. Akad. Wetensch., Ser. C 91(2): 106. man 3115 (CAY, P, U); withoutlocality,withoutdate
1988. Fig. 65. (9), Poiteau s.n. (B, P, type collection of P. digitata),
without date (8), Poiteau s.n. (P); OyapoqueR., Ca-
PouroumadigitataTr6cul,Ann. Sci. Nat. Bot., S6r. 3, mopi, 8 Mar 1976 (9), Sastre 4350 (P, U); Mt. St.
8: 106. 1847; Miquel in Martius,Fl. bras.4(1): 124. Marcel,28 Mar 1976 (9), Sastre 4579 (P).
1853. Type. FrenchGuiana.Withoutlocality,with- BRAZIL.AMAPA:Serrado Navio, 1961 (2),Aubre-
out date (9), Poiteau s.n. (lectotypeP, chosen here; ville 144 (P, U, US); rd. to Clevelandia,14 Oct 1950
isolectotypeB). (9),Fr6es 26631 (IAN), 13 Oct 1960 (a), Irwin48682
(GH, IAN, NY, U, US); Rio Oiapoque,Mt. Tipac, 16
Leafy twigs appressed-puberulousto strigose. Oct 1960 (8), Irwin48755 (IAN, NY); Rio Oiapoque,
Lamina 5-7(-9)-parted, midsegmentlanceolate Mt. Tipac, 16 Oct 1961 (6), Pires et al. 51624 (B, F,
138 Flora Neotropica

_l

? ..J

- i; . . . . .
'"

* . . . ..
.ru ,| x. | |.. |

FIG. 65. Pouroumabicolor subsp. digitata. Leaf and pistillate inflorescences BAFOG 1050).

FIG. 65. Pourouma bicolor subsp. digitata. Leaf and pistillate inflorescences (BAFOG 1050).

NY, R, UC, US). PARA:Rio Jari, Monte Dourado, 4
Feb 1969 (9), Cavalcante2297 (MG); 25-35 and 65
km SW of Tucurui,4 and 18-20 Nov 1981 (9), Daly
et al. 1208 and 1474 (BG);Rio Jari, Monte Dourado,
15 Nov 1968 (9), N. T. Silva 1411 (NY, U, US), 15
Nov 1968 (8), N. T. Silva 1412 (NY, U).
Vernacular names. Surinam. boroma (Ara-
wak), bospapaja,pouroumaor puruma(Carib),
yarayara(Carib).French Guiana. a'ilea (Waya-
pi), male bois canon or bois canon, kulumatelelel
(Wayapi).Brazil. Para:mapatirana.
Pourouma 139

This subspeciesresemblesP. cecropiifoliavery 30 Apr 1939 (st),Standley72667 (F);betweenVirginia

much. It is thereforesomewhatdoubtfulwhether and Lago Izabal, 5 Apr 1940 (st), Steyermark38916
Gentryet al. 46752 (fromVenezuela,Amazonas) (F).
BELIZE. Belmopan,21 Jun 1973 (6), Croat24831
belongsto subsp.digitataor representsajuvenile (MO); Stann Creek district, Middlesex, 5 May 1939
form of P. cecropiifolia.Subspeciesdigitata dif- (9), Gentle 2787 (A, F, NA, NY, US); Stann Creek
fers from P. cecropiifoliain the smaller number valley, 18 Mar 1940 (9),Gentle3265 (A, NY); Monkey
of incisions of the lamina and usuallya scabrous R., 18 Oct 1942 (6), Gentle 4211 (DUKE); El Cayo
to scabridulousupper surfaceof the lamina. district,HummingbirdHwy., 16 Apr 1955 (9), Gentle
8675 (F, S, US); Maya Mts., 18 Jun 1930 (9), Schipp
Nests of small ants are sometimes found in 127 (A, F, G, GH, MO, NY, S).
between the bases of the main veins. HONDURAS. Rio Esperanza,28 Feb 1903 (6), Wil-
Fromthe two syntypecollections,Leprieurs.n. son 606 (NY, US). ATLANTIDA: San Alejo, 22-27 Apr
and Poiteaus.n. (staminatespecimens),the latter 1947 (st),Standley7878 (F);Tela, 4 Dec 1927-20 Mar
1928 (st), Standley 52900 (A, F, US), (9), Standley
is chosen as lectoptypecollection. 53951 (F, US). CORTES: Agua Azul, 9 Feb 1952 (9),
Allen6401 (F, GH, US). GRACIAS ADios: Rio Platano,
17-23 May 1973 (9), Clewellet al. 4142 (MO).
3d. Pourouma bicolor Martius subsp. scobina NICARAGUA.ZELAYA: PuertoCabezas,2 Dec 1927
(Benoist) C. C. Berg & van Heusden, Proc. (9),Englesing 50 (F), (st), Englesing 52A (F);between
Kon. Ned. Akad.Wetensch.,Ser.C 91(2): 106. Bluefieldsand GinneyPoint, 24 Mar 1949 (st),Molina
1988. Fig. 66. R. 1963 (GH);El Recreo-ElPijibayerd., 11 May 1949
(st), Standley19904 (F);rd. to ColoniaManantiales,S
PouroumascobinaBenoist,Bull.Mus.Hist.Nat.(Paris) of ridgeof Serraniasde Yolania, 29-31 Oct 1977 (st),
28: 320. 1922; Woodson & Schery,Ann. Missouri Stevens4839 (BG);ca. 14.3 km N of El Enpalme,along
Bot. Gard.47: 167. 1960.Type:CostaRica. San new rd. to Mina Nueva America, 27 Apr 1978 (9),
Jose: Tucurrique,Dec 1898 (i), Tonduz12930 (ho- Stevens8323 (BG).
lotype, P; isotypes, F, GH, M, NY, S, US). COSTA RICA. ALAJUELA: Cariblanco,Quebrada
PouroumacrassiveniaCuatrecasas,Revista Acad. Co- Arrayanes,1 Jul 1973 (6),Lent 3547 (DUKE, F, MO,
lomb. Ci. Exact.9(36/37): 340. 1956. Type. Colom- U). CARTAGO:
Turrialba, 20 May 1951 (9), Le6n 3501
bia.Valle:RioDigua,Piedrade Moler,24 Aug1943 (NA). HEREDIA:
Puerto Viejo, Rio Sarapiqui,19 Jan
(9), Cuatrecasas15071 (holotype, F; isotypes, GH, 1974 (st), Hartshorn1348 (F), 1 Oct 1974 (9), Hart-
NY). shorn 1539 (F, U); Rio Sarapiqui, Tirimbina, 16 Jul
1971 (6), Lent 2004 (F, G, MO, NY, U, US); Rio
Leafy twigs appressed-puberulousto strigose Sarapiqui,La Virgen, 23 Jul 1979 (9), Stevens 13341
or hirtellous. Lamina usually 5-7(-9)-parted, (U). LIMON:Madrede Dios, 26 Feb 1949 (9),Holdridge
sometimes 3-lobed to -parted, midsegment 2514 (US); Rio Hondo, Aug 1901 (9), Pittier 16163
(G, US). PUNTARENAS:
Valley of Rio Diquis, between
mostly lanceolate to oblong, sometimes elliptic Union and San Pedro, 29 Mar 1898 (9), Pittier 12105
to obovate, base + deeply cordate;uppersurface (BR, P, US). SANJOSE:San Pablo, 2 Mar 1966 (9),
scabrous(to scabridulous),lowersurfacesmooth, Jimenez 3816 (F);San Isidro,El General,2 Mar 1966
hairs on the main vein appressedto patent, ar- (9), Molina R. 18227 (F, NY, US); El General, Jan
achnoid hairs (almost) confined to the areoles 1939 (6), Skutch4083 (A, MO, NY, S, US), Feb 1939
but the spots of arachnoidhairsmostly not clear- (9), Skutch4225 (A, MO, NY, S); Rio de las Vueltas,
Tucurrique,Dec 1898 (6), Tonduz12930 (F, GH, M,
ly separatedby the smaller veins; smaller veins NY, P, S, US, type collection of P. scobina).
more or less prominent;stipulesoutside with ap- PANAMA. COCLE: Valley of Rio Anton, 1000 m,
pressed or patent hairs, inside hairy, caducous. 27 Sep 1946 (9), Allen 3742 (BR, E, F, G, MO, P).
Pistillateinflorescenceswith (10-)20-50 (ormore COLON: SantaRita, Feb 1968 (9), Gomez-Pompa3385
(MO). DARIEN: Cerro Pirre, 14 Dec 1962 (9), Duke
?) flowers. 6588 (MO); Pidiaque, 28 Mar 1966 (9), Duke 8035
Distribution(Fig. 68). CentralAmerica (from (MO); between Cerro Pierre and Piji Vasal, 15 Nov
Guatemala to Panama) and the Pacific coastal 1977 (st), Folsom 6377 (MO);Cocalito,8-9 Apr 1978
region of Colombia and Ecuador,and to north- (st), Garwood759 (F); CerroPirre, 1000-1400 m, 29
Dec 1972(9),Gentryetal. 6997 (F, MO);Rio Setigandi,
westernVenezuela,possibly also in Peru(Loreto 19 Apr 1980 (9), Gentryet al. 28609 (U); Manene,23
and San Martin);in non-inundatedforest at al- Dec 1980 (9), Hartman 12176 (U); betweenPaya and
titudes up to ca. 1600 m. Pucro, 12 Jun 1959 (9), Steam et al. 423 (G, GH, LE,
US); PediaqueHill, nr. Rio Sabanaand Rio Lara, 16
Specimensexamined.GUATEMALA.ALTAVERA- Jul 1966 (2), Tyson et al. 4734 (DUKE, GH, MO);
PAZ:FincaCubilguitz,Mar 1913 (Q),Tuerckheim4082 Cana, 17 Apr-8 Jun 1908 (9),R. S. Williams983 (NY,
(US). IZABAL:Puerto Mendez, 12 Jun 1970 (2), Con- US). PANAMA: El Llano-Cartird., 20 Feb 1973 (9),
treras10026 (DUKE, F, U, US); El Estor,7 Mar 1972 Correaet al. 1854 (F, MO); CerroAzul, 26 Jul 1970
(2), Contreras11195 (DUKE, P, S, US); Entre Rios, (st), Croat 11591 (F, MO); between Cerro Azul and
140 Flora Neotropica

' I?

inch ? . J e . td. .

UNITDO STATES NATIONALli55UII5553 -N-

FIG. 66. Pouroumabicolorsubsp. scobina. Leafy twig with pistillateinflorescences(Allen6401).

CerroJefe, 6 Aug 1968 (Q),Dressier3579 (DUKE, GH, COLOMBIA.CHOC6:Rio Curiche,camp Curiche,

MO);El Llano-Cartird., 17 Apr 1981 (Q),Sytsma4018 25 May 1967 (9), Duke 11599 (US). VALLE: Rio Digua,
(U). SANBLAS:Cangandi,10 Feb 1986 (st), Neverset Piedrade Moler, 24 Aug 1943 (2), Cuatrecasas15071
al. 7178 (BG), 27 Mar 1986 (st), Nevers et al. 7542 (F, GH, NY, type collection of P. crassivenia);Rio
(BG). Sanquinini,La Laguna,14 Dec 1943 (d), Cuatrecasas
Pourouma
15520 (F, GH, NY), (9), Cuatrecasas15534 (F, GH,
NY).
VENEZUELA. BARINAS: Pedreza, Feb 1955 (2),
Bernardi1986 (NY, VEN);rd. Barinitas-ElCacao,21
Sep 1982 (9),Marcano-Bertet al. 982-154 (U). MERI-
DA:Caiio Amarillo, 5 Feb 1955 (2), Bernardi 1919 (F,
NY, VEN); El Vigia, 14 Jun 1979 (6), Marcano-Bert
et al. 238-979 (U). TACHIRA:
La Fria, 22 Dec 1965 (2),
Breteler4920 (MO, U); 35 km SSE of San Cristobal,
20-21 Mar 1981 (2), Liesner et al. 10944 (BG); San
Joaquinode Navay, 6 Nov 1979 (2), Stevermarket al.
119396 (U). ZULIA: Dtto. Col6n,nr. CasuiguaEl Cubo,
21 Jan 1979 (9), Bunting6836 (BG);Sierrade Perija,
SW ofMachiques, 31 Aug 1967 (2), Steyermark99945
(U, VEN).
ECUADOR.CARCHI: Maldonado,25 Sep 1979 (6),
Gentryet al. 26530 (U). ELORO:Piedras,20 Jun 1943
(st),Little 6640 (F, US). ESMERALDAS: SantoDomingo
de los Colorados-Quininderd., 14 Apr 1943, Acosta
S. 13605 (F);Quininde,12-14 Apr 1943(2), Little6241
(F, US), (st), Little 6258 (F, UC, US); San Lorenzo,20
Apr 1943 (2), Little 6301 (F, U, UC, US). IMBABURA:
Collapi,4 Jun 1949 (st),AcostaS. 12884 (F). Los Rios:
Rio PalenqueBiol. Station, 16 Aug 1976 (st), Dodson
6156 (MO), 2 Oct 1976 (st), Dodsonet al. 6312 (MO), 3e. Pouroumabicolor Martius subsp. chocoana
(6), Dodson et al. 6327 (GB, MO, U), 2 Feb 1974 (9), (Standley)C. C. Berg & van Heusden, Proc.
Gentry9552 (MO, U). PICHINCHA: Santo Domingo de
los Colorados,30 Aug 1930 (6), Benoist (P, S, U), 17
Oct 1961 (2), Cazaletet al. 5037 (FHO, NY, UC, US),
19 Oct 1961 (6), Cazalet et al. 5072 (FHO, NY, UC, Pourouma chocoanaStandley,Publ.FieldMus.Nat.
US).
PERU.LORETO: Prov.Maynas,YanamonoExplora-
ma TouristCamp,26 Jun 1983 (st), Gentryet al. 42168
SAN MARTIN:Rioja-Pomacochas rd., below
(BG).
Venceremos, 1600 m, 8 Feb 1984 (9), Gentryet al. Pourouma
45160 (BG).
Vernacular names. Guatemala. Guarumo de
montafia. Belize. Trumpet tree. Honduras.
Guarumo de montafia, mano de le6n. Nicaragua.
Guarumo macho, pacica, yahal (Moskito Indi-
ans). Costa Rica. Chumico, guarumo de mon-
tafia, lija. Panama. Darien: guarumo macho, pi-
raejo. Colombia. Choc6: uva. Venezuela. Barinas:
tinajero. Merida: orumo de monte. Ecuador. Es-
meraldas: guagay, guarumo de montafia, uva, uva
del monte.
The lamina is mostly 5-7-fid to -parted, but
in Panama 3-lobed laminas occur. The number
of flowers in the pistillate inflorescence is often
50 or more. In Guatemala and Panama collec-
tions with (10-20) flowers in the pistillate inflo-
rescence have been made. In several collections
in the southern part of the subspecies range (Ec-
uador to Panama) the indument of the lamina
beneath, the petioles, stipules, peduncles and leafy
twigs is patent (hirtellous to (sub)tomentose) in-
stead of appressed as in most collections of this
141
subspecies and in that of the other subspecies,
except for subsp. chocoana. Such specimensare
hardlydistinguishablefrom P. guianensissubsp.
guianensiswith regardto the leaf indument (see
p. 122). Fortunatelythese taxa can mostly be
distinguishedon the presenceof hairs on the in-
ner surface of the stipules and are, moreover,
geographicallyseparated.However, two recent
collections from Peru (San Martin;Gentryet al.
45160) and (Loreto, Gentryet al. 42168) may
also belong to subsp. scobina.The indumenton
the petiole, the leafy twigs, and the main veins
of the laminabeneathis denserthan usual in the
collections from Central America and western
Ecuador. The latter collection has, moreover,
bright-yellowhairs on the stipules and 9-parted
leaveswith subpetiolulatesegments.The identity
of these two collections is ratheruncertain.
Kon. Ned. Akad.Wetensch.,Ser.C 91(2): 106.
1988. Fig. 67
Bot.Ser.,22:73. 1940.Type:Colombia.Cho-
Hist.,
c6:Junctionof Rio CondotoandRio SanJuan,20
Apr1939(2),Killip&A. C.Smith35095(holotype,
F;isotype,US).
johnstoniiWoodson,Ann. MissouriBot.
Gard.47: 166.1960;Burger,FieldianaBot.40:201.
1977.Type.Panama.CanalZone,W of Lim6nBay,
GatunLocks,MaruTowers,27 Mar1956(Q),John-
ston1714(holotype,MO;isotype,A).
Leafy twigs patent- to appressed-puberulous
and at least on the scarsof the stipules(sparsely)
yellow-hirsute.Lamina mostly 5-fid to -parted,
sometimes 3-lobed to -fid or entire,occasionally
(in subjuvenilematerial?)7-parted,midsegment
broadlyelliptic to oblong, base deeply or some-
times shallowlycordate;uppersurfacescabrous,
lowersurfacesmooth,appressedhairson the main
veins, arachnoidhairs in the areoles and on the
smallerveins, mostly coveringthe areasbetween
the paralleltertiaryveins; smallerveins (almost)
plane;stipulesoutsideyellow-hirsuteto -subseri-
ceous, inside hairy, subpersistent.Pistillate in-
florescencesmostlywith up to 50 (ormore?)flow-
ers.
Distribution(Fig. 68). Panama (Canal Zone
and Col6n) to western Colombia (Choc6 and
Valle);in non-inundatedforestat altitudesup to
ca. 1000 m.
142 Flora Neotropica

-..

movririg-. o

Ievsed for YXora eotrei''

Pouroum bicolar orths* subsp, chocoasa

?Staodisy) C.C. krg & van Seusden

Dec~L~E~I!t.: C.c. Barg Utrechtr 19L6

OPPANABMt

der*-o lno Tru4. . 5

un~err rortbO~h
ofr trwil, tr--, : rtire tr,-
~rp~ an~~~~~~~~~~~~~~~~~~~rt-in~f4etedm
tkr~- ~-~:ht brow:1., ,('!'t wi';
~ag
t.einch99 Jaa i irr-f~'Lr hc~riz,.ntml rineg; 1
c~r~.
en
~ ~ ~ r C.RAVfJR~rok;%s~l
Il
6,FR
. Feb. r
pCv.bl
; z, 1c60
%N0 p
Ftrol..'
"'Cf.7

FIG. 67. Pouroumabicolorsubsp. chocoana.Leafy twig with staminateinflorescence(Croat8097).

Specimensexamined.PANAMA. Without locality, ColoradoIsland, 16 Aug 1927 (st), Kenoyer312 (US),

1861 (st), Hayes 32 (E), without date (2), Hayes 348
(NY). CANAL ZONE: Nr. Gamboa,8 Feb 1966 (2), Blum
2039 (MO), 19 Jun 1968 (st); BarroColoradoIsland,
22 Feb 1969 (d), Croat8097 (F, MO, NY), (9), Croat
8100 (F, MO), 14 Mar 1969 (st), Croat8648 (MO),23
May 1970 (st), Croat 10343 (MO), 9 Jun 1970 (st),
Croat 10830 (MO); nr. Gamboa, Foster et al. 584
(DUKE, NY), 18 Dec 1972 (d), Gentry6679 (MO);
GatunLake, Gatun Locks,27 Mar 1956 (2), Johnston
1714 (A, MO, type collectionof P. johnstonii);Barro
27 Oct 1931 (st),Shattuck260 (A, F, MO),4 Dec 1931
(d),Shattuck522 (F, MO, US) and 525 (F, MO);N of
Frijoles, 19 Dec 1923 (st), Standley27479 and 27502
(US). CoIkN:Donese district,CampBatija,9-14 May
1968 (9), Holdridge6254A (MO).
COLOMBIA.CHOC6:Quibdo-Istminard., 48 km
S of Quibdo,nr. Certegui,8 Jan 1979 (9), Gentryet al
23835 (BG);Quibdo-Tutunendord., 3 km W of Tu-
tunendo,7 Jan 1981 (st), Gentryet al. 30278 (U);junc-
tion of Rio Condoto and Rio San Juan, 20 Apr 1939
 O^^Jgy -' i biccoolr P. bicolor subsp.

* ^ ^ ^ ~
_' "( *?'7 * nde p. biColor subsp. digitate
*' -
-r~ E^^ "*^ s.cbina Aand P. bicolor subsp.

P. bicolor *P. ccrepifo.

- '- s
_SAs. L ? [

FIG. 68. Distribution of Pourouma bicolor subsp. bicolor, P. bicolor subsp. chocoana, P. bicolor subsp.
digitata, P. bicolor subsp. scobina, P. bicolor subsp. tessmannii, and P. cecropiifolia.
144
(9),Killip35095 (F, US, typecollectionofP. chocoana).
NARIMO: Mun. Barbacoas,Barbacoas-Juninrd., 1050
m, 7 Aug 1982 (9), Mora 2295 (US); Mun. Iscuande,
Rio Sequi6n,22 Nov 1955(d),Romero-Castaneda 5484
(AAU). VALLE: Rio Yurumangui,Veneral,28 Jan-10
Feb 1944 (a), Cuatrecasas15876 (F); Rio Calima, La
Trojita, 19 Feb-10 Mar 1944 (2), Cuatrecasas16315
(F);LaEsperanza,6-7 Mar 1944(d),Cuatrecasas16755
(F); San Isidro, 2-5 May 1944 (8), Cuatrecasas17357
(F);Rio Cajambre,Silva, 5-15 May 1944 (st), Cuatre-
casas 17451 and 17479 (F);Bajo Calima,ca. 15 km N
of Buenaventura,18 Feb 1983 (st), Gentryet al. 40479
(BG);Bajo Calima,rd. to JuanchacoPalmares,18 Jul
1984 (2), Gentryet al. 48303 (BG).
Vernacularnames:Panama.Mangabe.Colom-
bia. Antioquia:sirpo;Nariiio: guagay.
This subspecies can be easily distinguished
from subsp. scobinaby the arachnoidindument
coveringthe smallerveins (betweenthe parallel
tertiaryveins), and often also by the relatively
broadsegmentsof the lamina. Some of the spec-
imens are yellow-hirsuteon the leafy twigs, or
also on the stipules, petioles, main veins of the
lamina above and/or the peduncles.Such spec-
imens resemble P. oraria very much. The two
taxa differ in the smooth upper surface of the
lamina, the smooth fruiting perianth, and the
concentrationof hirsuteindumentat the adaxial
side of the petiole. The resemblancesbetween
these two taxa are so strikingthat we sometimes
wondered whether they could be two forms of
the same species (distinct from P. bicolor).
4. Pourouma cecropiifolia Martius in Spix &
Martius,Reise Bras.3:1130. 1831, as Puruma
cecropiaefolia;Martius,Syst. mat. med. bras.
34. 1843; Miquel in Martius, Fl. bras. 4(1):
123, t. 36. 1843; Macbride,Publ. Field Mus.
Nat. Hist., Bot. Ser., 13(2.2):291. 1937. Type.
Brazil. "In silvis prov. Paraensiset Rio Ne-
gro,"Dec 1819 (st), Martiuss. n. (lectotypeM,
chosen here). Fig. 69.
PouroumamultifidaTrecul, Ann. Sci. Nat. Bot., Ser.
3, 8: 107. 1847; Miquel in Martius, Fl. bras. 4(1):
123, 1853. Type. Brazil?Without locality, without
date (9), Anonymus(=Ferreira?)s.n. (holotype, P;
isotype, P).
Pouroumasapida Karsten,Fl. Columb. 2: 19, t. 110.
1862.Type.Colombia.Meta:Villavicencio,nr.Jira-
mene, Oct 1872 (a), Karstens.n. (holotype,GOET;
isotype?,LE-with fragmentsof staminateinflores-
cences!).
PouroumaedulisDufresne,Ill. Hort.20:70. 1873.Based
on (living?)material from Colombia, collected by
Linden).
PouroumauviferaRusby,Mem. New YorkBot. Gard.
FloraNeotropica
7: 231. 1927.Type.Bolivia.Beni:Rurrenabaque,
Oct 1921(9),Rusby1599(holotype,NY; isotypes,
F, GH,US).
Tree, up to 20 m tall. Leafy twigs 5-30 mm
thick,(sparsely)white-pubescentand with sparse
to dense, brown, pluricellularhairs, often gla-
brescent.Lamina 7-1 1-parted,rarely3-5-lobed,
ca. 10-40 x 13-60 cm, coriaceous, apex acu-
minate to acute, base deeply cordate, with or
withoutoverlappinglobes;uppersurfacesmooth,
white puberulouson the main veins and some-
times with sparse,red-brown,pluricellularhairs
usuallyon the smallerveins, lowersurfacewhite-
or sometimes yellow-, usually appressed-pubes-
cent to -puberulouson the veins, occasionally
brown, pluricellularhairs on the main veins,
arachnoidhairs in the areoles, often also on the
smaller veins; lateralveins ca. 15-30 pairs, ter-
tiary venation almost plane beneath;petiole 8-
50 cm long,glabrousor occasionallywith brown,
pluricellularhairs;stipules2-20 cm long, outside
white-appressed-puberulous andwith(dense)red-
brown,pluricellularhairs,inside denselyyellow-
hirsuteand sometimes with brown,pluricellular
hairs, caducous. Staminate inflorescencesup to
27 cm long and 17 cm wide; peduncle 2-11 cm
long, peduncle and branchesyellow- or white-,
usually appressed-puberulousand usually with
dense, (dark-or red-)brown,pluricellularhairs;
flowers sessile, ? diffuselydistributedalong the
ultimate branches;tepals ca. 2.5 mm long, free
or occasionally basally connate, ciliolate; fila-
ments shorterthan the tepals, free. Pistillate in-
florescencesin fruit up to 24 cm long and 22 cm
wide; peduncle 2-13.5 cm long, peduncle and
brancheswith indument similar to that of the
staminateinflorescence;flowers 25-170; pedicel
up to 0.6 cm, in fruit up to 2 cm long; perianth
4-9 mm long; white- to yellow-puberulousand
usually with (dark or red-) brown, pluricellular
hairs;stigmapeltate,ca. 1.5-2 mm diam.,sparsely
white-puberulous.Fruitingperianthdarkpurple
to black, ovoid to subglobose, 1.5-3.5 x 0.7-2
cm, densely to sparsely puberulousand some-
times with brown, pluricellularhairs.
Distribution(Fig. 68). ExtendingfromColom-
bia (Meta) throughAmazonian Ecuador,Peru,
and Brazil(Acreand westernAmazonas)to Am-
azonian Bolivia, also in Amazonian Venezuela;
in non-inundatedforest, at altitudes up to ca.
1000 m. Often cultivated and their occurrence
often indicating previous human inhabitation.
Pourouma 145

? ^T
\Sr~~u..I .

,J,

0f W/+ O.+ 14+ I+

P OUR 0 ITMA cecropiaefolia.

FIG. 69. Pourouma cecropiifolia. From Martius, Flora Brasiliensis 4(1). 1853: tab. 36. Leaf, stipule (32),
staminateinflorescence,diagramof staminate flower (D), part of staminate flower (28), staminateflower (1),
tepals (4), stamens (7).
146 Flora Neotropica

Also cultivated in some places outside the species al. 43137 (BG); 2-8 Aug 1929 (9), Killipet al. 27381
range, e.g., Bahia (Brazil). (NY, US); Yurimaguas,24-25 Aug 1929 (2), Killipet
al. 27932 (F, NY, US); Iquitos,26 Sept 1929 (2),Killip
Specimensexamined.COLOMBIA.Without local- et al. 29839 (F, NY, US), Apr 1930 (6), Klug 1185 (F,
ity, without date (9), Triana 862 (E, NY, P), without NY, US), May 1930 (2), Klug 1326 (F, NY, US); Rio
date (9), Triana 1892 (NY). AMAZONAS: Rio Boiauasu, Marafion,Perseveranca,5 Feb 1924(2), Kuhlman1337
28 Nov 1945 (st), Duque-Jaramillo2261 (NY); Rio (F, RB);Pongode Manseriche,11 Dec 1931 (2), Mexia
Loretoyacu,Barracon,25 Dec 1945 (2), Duque-Jara- 6257 (F, G, GB, NY, S, U, UC, US); Prov. Requena,
millo 2446 (NY); between Rio Kananariand Rio Pa- nr. JenaroHerrera,4 Nov 1986 (2), Peters 187 (BG);
coa, 1-5 Dec 1951 (6),Garcia-Barriga13871 (US);Rio Rio Yaguasyacu,BrilloNuevo, 16 Apr 1977 (2), Plow-
Igara-Parana, La Chorrera,20 Sep 1973(2), Sastre2274 man et al. 6903 (F, GH);Rio Yucayali, 1923 (a), Tess-
(G, P); Los Monos, 24 Sep 1978 (st), Pabon E. 573 mann 3054 (G, S); Rio Itaya,Paraiso,9 Oct 1929 (2),
(herb.Araracuara);Rio Caraparana,El Encanto, 22- Ll. Williams3347 (F, US); Yurimaguas,4 Nov 1929
28 May 1942 (st), Schultes 3826 (NA), (2), Schultes (9), Ll. Williams3984 (F), 10 Nov 1929 (2), LI. Wil-
3862 (F); Rio Igara-Parana,La Chorrera,4-10 1942 liams 4627 (F). MADREDEDios: Rio Manu, Cocha
(2), Schultes 3901 (F); Rio Boiauasu, Nov 1945 (2), Cashu Station, 25 Nov 1980 (2), Foster 5905 (F), 15
Schultes6805 (F, US). CAQUETA: Las Guacamayas,28 Sep 1979 (st), Foster 7001 (F); Tambopata, 19 Feb
Apr 1944 (9), Little 7756 (US); Morelia, 7 Oct 1941 1984 (st), Gentryet al 45660 (BG). PASCO: Prov. Ox-
(6),vonSneiderns.n. (S).META:Villavicencio,nr.Jira- apampa,Iscozacin, 24 Jun 1982 (2), D. Smith et al.
mene, Oct 1872 (9),Karstens.n. (GOET,LE),(6),Kar- 2089 (BG). SAN MARTIN:Puerto Pizana, 27 Jul 1973
sten s.n. (GOET, LE, type collection of P. sapida). (9), J. Schunke V. 6485 (F, GH, MO, NY). UCAYALI:
PUTUMAYO: Rio Putumayo, 28 Aug 1966 (9), Pinkley Prov. CoronelPortillo, LagoYarinacocha,Povenir, 5
405 (S); PuertoLim6n, 27-28 Feb 1942 (st), Schultes Nov 1984 (2), Maas et al. 6201 (U); Prov. Coronel
3343 (F). VAUPES:Mitui,10 Sep 1939 (9), Cuatrecasas Portillo, km 13 of rd. from Campo Verde, km 34 of
et al. 6737 (F); Upper Rio Guaviare,9 Nov 1939 (6), rd. Pucallpa-TingoMaria,7 Nov 1984 (2), Maas et al.
Cuatrecasas7602 (F, US); Mitui,7-8 Nov 1952 (2), 6225 (U).
Humbertet al. 27221 (P, U, US); nr. Yavarate,20 Nov BRAZIL.Withoutlocality, without date (2), Anon-
1952 (6),Romero-Castaneda3647 (NY);Mitu, 26 Sep ymus (=Ferreira?)s.n. (P, type collection of P. multi-
1976 (6), Zarucchi2145 (INPA). fida). ACRE: Rd. Abuna-Rio Branco,km 242-246, 20
VENEZUELA. AMAZONAS:Cerro Duida, base of Jul 1968 (2), Forero et al. P.6419 (INPA, NY, R, S,
Duida, Jan-Feb 1939 (st), Farinas et al. 347 (NY, U, U); mouth of Rio Macauhan,8 Aug 1933 (2), Krukoff
VEN);Depto. Atures, 35 km SE of PuertoAyacucho, 5327 (A, F, G, LE, M, MO, NY, S, U, UC, US), (a),
25 Sep 1980 (9), Guanchez239 (VEN);Rio Orinoco, Krukof 5332 (A, F, G, M, MO, NY, S, U, UC); Rio
betweenCafnoGrullaand Grulla,8 Apr 1978 (9),Mo- Branco, ParqueZoobotanico da Univ. Fed., 15 Oct
rillo et al. 7384 (VEN);Depto. Atapabo,nr. Culebra, 1980 (2), Nelson 705 (BG);Tarauaca,24 Sep 1968 (2),
Rio Cunucunuma,22 Mar-4 Apr 1983 (st), Steyer- Pranceet al. 7503 (F, INPA, M, NY, P, R, S, U, US);
market al. 129163 (BG, VEN). Sao Francisco, Jun 1911 (2), Ule 9314 (G, MG).
ECUADOR. MORONA-SANTIAGO: 5 km SW of Ma- AMAZONAS: Rio Javari,Atalaiado Norte, withoutdate
cas, 26 Jan 1981 (6), Berg 1223 (BG, U). NAPO:Tena, (2), Braga et al. 3178 (INPA);Manaus(cult.!),15 Apr
20 Oct 1939 (9), Asplund9462 (R); San Pablo de los 1955 (2), Chagas(INPA)962 (INPA, U), 29 Jul 1978
Secoyas, 19 Aug 1980 (9), Asanza C. 32929 (AAU); (6),Falcao 215 (INPA);Marco,29 Jan 1969 (2), Croat
Aiiangu,30 May-21 Jun 1982 (st),SEF8764 and 8930 7633 (MO);Rio Japura,mouthof Rio Apaporis,4 Dec
(U); 3 km E of Atahualpa, 10 km E of Puerto Napo, 1912(a),Ducke(MG)12366 (MG);Rio Solimoes,Fon-
13 Dec 1985 (9), Neill et al. 7044 (BG). PASTAZA:Rio teboa, 8 Jun 1945 (9),Fr6es21048 (NY, US); Tefe, 19
Curaray,Loricachi,30 May 1980 (9), Jaramilloet al. Jul 1972 (9),Kriegeret al. (INPA)2268 (INPA);mouth
31516 (AAU). of Rio Embira, 1 Jul 1933 (2), Krukoff5109 (A, F, G,
PERU. AMAZONAS: Rio Cenepa, 13 Dec 1972 (2), M, MO, NY, S, UC, US); Sao Paulo de Olivenca, 11
Berlin 523 (F, GH, NY), 4 Aug 1974 (9), Berlin 1999 Sep-26 Oct 1936 (2), Krukoff8332(BR, F, G, LE,MO,
(NY). HUANUCO: Tingo Maria, 10 Aug 1940 (9), As- P, S, U, US), (2), Krukoff8469 (A, B, BR, F, G, LE,
plund 12933 (S, US), 10 Jul 1957 (st), Ellenberg2288 MO,NY, P, S, U, US);Manaus,EstradaRoas de Maio,
(U); Aucayacu,27 Sep 1967 (9), Lao et al. s.n. (F, G, Armando(cult.!),25 Sep 1973 (8), Lisboa 27 (INPA);
US); PuertoInca, 7 Dec 1968 (2), J. Schunke V. 2824 Rio Papori, Trindade, 13 Dec 1928 (st), Luetzelburg
(F, G, GH, NY, US). JUNiN:La Merced, 10-24 Aug 23913 (R), 27 Dec 1928 (2), Luetzelburg23887(M, R);
1923 (9),Macbride5446 (F, GH, US). LORETO: Iquitos, Rio Curicuriari,8 Jul 1979 (2), Maia et al. 497 (INPA);
29 Oct 1970 (2), Asplund14124 (G, S);Zungarococha, various collections without precise indications of lo-
28 Jan 1978 (st), Ayala et al. 1446 (U); Rio Ampiaco, calities (probablymost of them along Rio Amazonas
Puca Urquilla,24 Feb 1972 (9), Croat20662 (DUKE, and Rio Japurf,(Dec) 1819-(Feb) 1820 (6 or st), Mar-
MO, NY, U); Iquitos, 8 Aug 1906 (9), Ducke (MG) tius s.n. (G, M, includinglectotypeof P. cecropiifolia);
7581 (G, MG); El Sacramento,89 km S of Pucallpa, Taparuquara,22 Oct 1971 (2), Prance et al. 15791
22 Jul 1957 (st), Ellenberg2474 (U); halfwaybetween (INPA, NY, U), Jul-Aug 1967 (9), Schultes 24547
Iquitos and SantaMariade Nanay, 30 May 1978 (st), (INPA);betweenBarcellosand Sao Gabriel,Dec 1851
Gentryet al. 22460 (U); Prov. Maynas, Yanamono (2), Spruce2023 (B, BR, CGE,P);Rio Jurua,Bom Fin,
ExploramaTourist Camp, 15 Jul 1983 (st), Gentryet Oct 1900 (2), Ule 5265 (B, F, G, HBFG, MG, NY);
Pourouma 147

Rio Jurua,Marary,8 Oct 1900 (c), Ule 5266 (HBG, 1951(e),Garcia-Barriga14002(holotype,US;iso-

MG). type,NY).
BOLIVIA.BENI:Prov. Moxos, S of Moxos, 18 Sep
1976(9),Menecesetal.338(MG,MO);Rurrenabaque, Tree, up to 30 m tall. Leafy twigs 4-8 mm
Oct 1921 (9), Rusby 1599 (F, GH, NY, US). PANDO: thick, densely to sparsely yellow-hirsute,these
Prov.Nicolas Suarez,Mukden,Jun-Dec 1979 (2), Iza-
wa21 (U). long hairs often intermixedwith much shorter,
whitishhairs,sometimesalso with sparse,brown,
Vernacularnames. Colombia. bochoa tsaha pluricellularhairs, sometimes (sub)glabrousor
(Kofan),caimaron,caimaronsilvestre, uva, uva subvelutinous.Lamina entire,ovate to subovate
del monte, uvilla, uvo; Amazonas: gurucana. or elliptic to oblong and 6-20 x 2.5-12 cm or
Venezuela: cocura, sadajii. Peru. sacha uvilla, 3-lobedto -parted(or in subjuvenilematerialup
uvilla, suia; Pasco: shewantaqui.Brazil:cucura, to 5-parted,sometimeswith the segmentsalmost
cucuva,imbauba(or ambauva)mansa,imbafba petiolulate)and up to ca. 25 x 25 cm, coriaceous
(or ambauva)do vinho, mapati, puruma,puru- to subcoriaceous,apex acuminate, acute, apic-
ma-y, sucuuiba,uva. Bolivia. uva silvestre;Beni: ulate,or sometimesemarginate,base subcordate
tanaribe,uva del monte. to truncateto roundedto subacute;uppersurface
This species shows strongsimilaritiesto P. bi- ? scabrous,occasionallysmooth, yellow-hirsute
color, especially to the subspecies digitata and to -hirtellouson the whole surfaceor only on the
scobina. It mainly differs in the greater number main veins, lowersurfaceyellow-hirsute(to -hir-
of segmentsof the leaves of adultspecimens(nor- tellous) to -substrigoseon the main veins, white
mally 7-11 versus normally 5-7 in subsp. digi- arachnoidhairsin the areolesand on the smaller
tata and subsp. scobina),the smooth uppersur- veins; lateral veins 10-20 pairs, the basal pair
face,and the sparserindumenton the leafytwigs. branched,tertiaryvenation prominentbeneath;
The lamina of subsp.digitataand subsp.scobina petiole 3-17 cm long, yellow-hirsute,intermixed
is sometimes smooth and subjuvenilespecimens with short,whitishhairs;stipules2-12.5 cm long,
may have up to 9 leaf segments, which makes outside yellow-hirsute to -strigose, inside gla-
distinction of the taxa on the basis of morpho- brous, caducous.Staminate inflorescencesup to
logical characterssometimes impossible. For- 12 cm long and 4.5 cm wide; peduncle3-6.5 cm
tunately,the taxa areallopatric(or almost so, see long, peduncle and branches yellow-hirsute;
p. 139). flowers sessile, rarelypedicellate, clustered,but
The lectotypeofP. cecropiifoliais selectedfrom not in globose heads;tepals almost free, (sparse-
nine Martius syntype collections listed above, ly) puberulent;filaments free, shorter than the
that of P. sapidafrom two Karsten(syntype)col- tepals. Pistillate inflorescencesin fruit up to 19
lections. cm longand 10.5 cm wide;peduncle3-9 cm long,
The fully mature fruitingperianthof most (if peduncleandbranchesyellow-hirsute;flowers 20-
not all) Pourouma species is suitable for human 25; pedicel up to 0.6 cm long, in fruit up to 1
consumption, but P. cecropiifoliais the only cm; perianth0.3-0.9 mm long, hispidulous (or
species in cultivation (see p. 116). hirtellous);stigma peltate, ca. 2 mm in diam.,
The fact that the species is so often found in whitishpuberulous.Fruitingperianthblue-black,
(formerly) inhabited places suggests that the ellipsoid to oblongoid to subglobose, ca. 1-1.5
present distribution is (at least) partly anthro- cm long, hispidulous(or hirtellous).
pogenous. Distribution (Fig. 74). Colombia (Amazonas
and Vaupes), Venezuela (Amazonas and Boli-
5. PouroumacucuraStandley & Cuatrecasasin var),Ecuador(Napoand Morona-Santiago), Peru
Cuatrecasas, Fieldiana Bot. 28(1): 211. 1951. (Loreto and Madre de Dios) and Brazil (Amapa,
Type. Venezuela. Amazonas: Capihuara, Alto Amazonas,and Mato Grosso),extendingto east-
Casiquiare,5 Jun 1942 (Q),LI. Williams15812 ern Venezuela(Bolivar);in non-inundatedor in
(holotype, F; isotypes, G, NY, RB, US). periodicallyinundated (varzea or igap6) forest;
Fig. 70. mostly at low altitudes,sometimes (in Ecuador)
up to ca. 1500 m.
PouroumagarcianaCuatrecasas,Caldasia7:298. 1956.
Type. Colombia.Amazonas-Vaupes:Rio Apaporis, Specimensexamined.COLOMBIA.AMAZONAS:
Rio
betweenRio Kananariand Rio Pacoa,1-15 Dec Apaporis,abovemouthof Rio Kananari,Soratama,
148 Flora Neotropica

~ ~ ~ ~ ~
dl;i?~~~~~~~~~~~~~~~~~~~~~~~~~,T!
::. ~ ~ ~ ~ ?~

IWITi NAIONAL to PiQtJLAS IA AMAI7AA4A

5352ew J /rurourm eurura

Starl,l.

.F2;?Z,3!: -~ '; T.'l ^near

-
rlpu
Rio Jurw-.
, Hufb,b't C t.r.
10 ':"
- t
59"'?1'N:
disturbed f"rest r"nterra firrie. ;ie, .
Nx hnc. lt . W*itbhtilt.
7.t. i.- ; -
oiua to whitte benestb, 3 partim-l r *r
^* 'v-~~~ 1
70.
"
cucra
-'
FIG.~~~~~~~~~~~~~~~~~~1
gla
fol~avt., pwerianthb,
p.-dofv
inftovescence
papper &
(rnce
nireenesl
Puroua~ et "
Lefwgwihpitllat"r
: l. 84)
brown. Exuding br iw -]
t 'nning to blacsk. "UmbinublI 'A
,' L

O.T.Pr nce, F.A.ilsby, C...Berg.

W.C.St_fw d, E. Lleras, TT . s..,

-- ctber A
ii____ _ _ .
FIG. 70 Pourouma cu. atIg~~~~~with ~
pist~ilat a"n: i.8
flrM..
PA.
s ( r'ane, et. 20.

FIG. 70. Pourouma cucura. Leafy twig with pistillate inflorescences (Prance et al. 18240).

15-19 Dec 1951 (9), Garcia-Barriga14106 (NY, US), VENEZUELA. AMAZONAS:

18 Mar 1952 (9), Schulteset al. 15973 (F), Jan 1952
(6), Schultes et al. 19835 (US). AMAZONAS-VAUPES: Rio
Apaporis, between Rio Kananariand Rio Pacoa, 1-
15 Dec 1951 (9), Garcia-Barriga14002 (GH, NY, US,
type collection of P. garciana).
Capihuara, Alto Casi-
quiare, 5 Jun 1942 (9), Ll. Williams 15812 (G, NY,
RB, VEN, US, type collection of P. cucura).BOUVAR:
Rio Tirica, Oct 1947 (9), Cardona2170 (MO, VEN).
ECUADOR.NAPO:Rio Cuyabeno,16 Feb 1980 (8),
Berg&Akkermans1041 (U), Berg&Akkermans1042
Pourouma
(U), 17 Feb 1980 (6), Berg&Akkermans1045 (U), (2),
Berg & Akkermans1053 (U), 20 Feb 1980 (st), Berg
& Akkermans1070 (U); Rio Aguarico, Shushufindi,
26 Dec 1974 (2), Vickers86 (F).
PERU. LORETO: Rio Ucayali, JenaroHerrera,Jul-
Sep 1976 (6), Bernardis.n. (U); Rio Nanay, Puerto
Almendras,22 Feb 1979 (st), Gentryet al. 24900 (U);
Rio Yaguasyacu,tributaryof Rio Ampiyacu, 9 Nov
1977 (6), Gentryet al. 20515 (MO, U); Rio Nanay,
halfwaybetweenIquitosand SantaMariade Nanay,26
Feb 1979 (st), Gentryet al. 25100, 25050 (U); E of
Tamshiyacu,below SerafinFilomeno, 19 Mar 1979
(2), Gentryet al. 25854 (U); Rio Nanay, betweenIqui-
tos andSantaMaria,de Nanay,23 Feb 1979 (2), Gentry
et al. 31655 (U);below SerafinFilomeno, 19 Mar 1979
(2), Gentryet al. 25854 (U); Rio Yaguasyacu,nr. Brillo
Nuevo, 16 Apr 1977 (st),Plowmanet al. 6906 (F, GH);
Rio Nanay, nr. Iquitos, 14 Nov 1976 (st),Revilla 1818
(U); rd. to Pena Negra, 25 km from Iquitos, 30 Nov
1976 (9),Rimachi2724 (DUKE, F, MO, NY) Rio Na-
nay, above BellaVista, 18 Jan 1977 (2), Ramachi2765 6. PouroumacuspidataMildbraed,Notizbl. Bot.
(F, MO); rd. from Orellanato Sta. Catalena,km 5, 9
Dec 1982 (2), C. R. Rodrigues 944 (BG). MADREDE
Dios: Rd. Maldonado-Quincemil, 10 Jul 1968 (6),
AcostaM. 12 (BG, DUKE, F, NY, P); Tambopata,22
Feb 1984 (st), Gentryet al. 45922 (BG).
BRAZIL.ACRE:Reserva INCRA Santa Luzia, km
40, BR-364, 5-19 Oct 1984 (st), Campbellet al. 6927
(BG). AMAPA: Serrodo Navio, P6rto Platon, 10 Oct-
15 Dec 1976 (st), Rosa 1091 (MG), 10 Oct-15 Dec Pourouma
1976 (st), Rosa 1159 (MG). AMAZONAS:Rio Japura,
Vila Bittancourt,20 Nov 1982 (8), Amaral et al. 604
(BG); Manaus-Itacoatiara rd., km 26, 12 Aug 1976
Oliveira 60534 Rio De-
(st), (INPA) (INPA);Tototobi,
mini, 25 Feb 1969 (9),Pranceet al. 10231 (INPA, MG,
MO, NY, U, US); Manaus-Itacoatiarard., km 26, 2
Oct 1965 (st),Rodrigues8066 (INPA).MATOGROSSO:
Aripuana,28 Jan 1977 (st), Gomeset al. 614 (INPA),
21 Feb 1977 (st), Gomes et al. 844 (INPA), 29 Mar
1977 (st), Gomes et al. 1040 (INPA); Aripuana,nr.
HumboldtCentre,rd. to Rio Juruena,8 Oct 1973 (2),
Pranceet al. 18240 (F, INPA, MG, MO, NY, P, S, U,
US).
Vernacular names. Venezuela. Amazonas: cu-
cura; Bolivar: kaibarikei (Arekuna), sarasara
(Camaracoto). Ecuador. Santiago-Zamora: guar-
umo colorado. Peru. Loreto: dacha uvillos, uvil-
la; Brazil. Amazonas: imbaubarana branca;
Amapa: imbaubarana branca, inbaubarana folha
peluda; Mato Grosso: imbaiuba or umbauiba, im-
baubarana.
The species is very variable. The long, yellow
hairs on the leafy twigs, petioles and stipules vary
from very dense to sparse. The leaves of (adult)
specimens collected in Colombia, and Venezuela
and in Amapa and Amazonas (Brazil) normally
have entire and elliptic leaves. The collections
made in Mato Grosso (Brazil) have on the same
twig entire and 3-parted leaves. The material from
Ecuador and Peru mostly have ovate and often
149
3-lobedleaves (similarto the common leaf shape
in P. bicolorsubsp.bicolor).Subjuvenilematerial
fromthesetwo countriesoftenhaveup to 7-parted
leaves, with the segments often petiolulate.
The species may prove to consist of two or
three morphologically readily distinguishable
entities, mainlybasedon differencesin the shape
and incisions of the lamina. The relativelysmall
number of collections prevent establishingnew
infra-specifictaxa.
The identity of the two Rosa collections from
Amapa(Serrodo Navio), consistingonly of (fal-
len?)leaves, is not certain.The petioles and the
main veins of the lamina beneath are densely
yellow-hirsute.
Gart. Berlin-Dahlem10: 417. 1926; Warburg
ex Ule, Bot. Jahrb.Syst.40:150. 1907, nomen.
Type. Brazil. Acre: Confluenceof Rio Jurua-
Mirim and Rio Juruf, Aug 1901 (6), Ule 5719
(lectotypeB, chosen here;isolectotypes,F, G,
GH, MG).
Cuatrecasas,Caldasia7: 304.
tergoscabra
1956.Type.Colombia. Caqueta: RioCaqueta, below
mouthof Rio Orteguaza, Solano,8 km SEof Tres
4 Mar1945(v),Littleetal. 9527(holotype,
Esquinas,
F; isotypes, NY, US).
Tree, up to 30 m tall. Leafy twigs 8-12 mm
thick, white-hirtellousto -puberulous,and with
usually ratherdense, brown, pluricellularhairs.
Lamina 3-lobed or 3-5 parted, 10-35 x 18-45
cm, coriaceous,apex long acuminate,base cor-
date, with or without overlappinglobes; upper
surface smooth, hirtellous or occasionally also
withbrown,pluricellularhairson the main veins,
lower surface strigose, main veins hispidulous
and sometimes with brown, pluricellularhairs,
arachnoidhairs confined to the areoles; lateral
veins 13-30 pairs,tertiaryvenation + prominent
beneath;petiole 11-40 cm long, white-hispidu-
lous, and with brown,pluricellularhairs;stipules
7-13 cm long, outside white-appressed-puberu-
lous and with brown, pluricellularhairs, inside
sparsely white-puberulousto subglabrous,ca-
ducous. Staminate inflorescencesup to 11 cm
long and 4.5 cm wide; peduncle 4-6 cm long,
(densely) white-puberulous and with (dense),
brown, pluricellularhairs;flowers sessile, + dif-
fusely distributedalong the ultimate branches;
tepals ca. 1 mm long, free or basally connate,
sparselyhairy;filamentsshorterthan the tepals.
150
Pistillate inflorescencesin fruit up to 12.5 cm
long and 8 cm wide; peduncle 4-6 cm long, pe-
duncle and branches with indument similar to
that of the staminateinflorescences;flowers 10-
35; pedicel0.2-0.4 cm, in fruitup to 0.8 cm long;
perianth 2-6 mm long, hispidulous and with
brown,pluricellularhairs;stigmapeltate,ca. 1.5
mm diam., white puberulous.Fruitingperianth
ovoid to ellipsoid, 12-15 x 5-11 mm, scabrous.
Distribution (Fig. 74). Colombia (Caqueta) Ec-
uador(Napo),Peru(Pasco),and Brazil(Acreand
Amazonas);in non-inundatedforest.
Specimens examined. COLOMBIA. CAQUETA:So-
lano, 8 km SE of Tres Esquinas,Rio Caqueta,below
mouth of Rio Orteguaza,4 Mar 1945 (2), Little et al.
9527 (F, GH, NY, US, type collection of P. tergosca-
bra).
ECUADOR. NAPO:Afiangu,30 May-21 Jun 1982
(st), SEF 8532 (U).
PERU. PASCO: Prov Oxapampa,Rio Iscozacin,Ca-
beza de Mono, 12 Jun 1983 (st), Gentryet al. 41890
(BG).
BRAZIL. ACRE:Reserva INCRA Santa Luzia, km
40, BR-364, 5-19 Oct 1984 (st), Campbellet al. 6834
(BG);confluenceof Rio Jurua-Mirimand Rio Jurua,
Aug 1901 (6), Ule 5719 (B, F, G, GH, MG, lectotype
collectionofP. cuspidata),Aug 1901 (2), Ule 5719 (B,
G). AMAZONAS: Sao Paulode Olivenca, 11 Sep-26 Oct
1936 (Q),Krukoff8373 (A, F, NY, S, U).
This species, characterizedby incised leaves
and the smooth uppersurface,but scabrouslow-
er surface of the lamina, may be related to P.
bicolor.
Ule collected staminate and pistillate speci-
mens under the same number 5719. The sta-
minate collectionis selectedas the lectotypecol-
lection.
7. Pourouma myrmecophila Ducke, Bull. Mus.
Hist. Nat. (Paris),Ser. 2, 4: 723. 1932. Type.
Brazil. Amazonas: Manaus, Cachoeira do
Mindui,8 Jul 1929 (d),Ducke(RB)23607 (lec-
totype, RB, chosen here; isolectotypes, G, P,
S, US). Fig. 71.
Tree, up to 8 m tall, or shrub, myrmecophi-
lous. Leafy twigs 0.6-1 cm thick, densely yel-
low(ish)-hirsute,glabrescent,and with reddish-
brown,pluricellularhairs.Lamina 3-5-lobed, to
-fid, 13-38 x 15-42 cm, subcoriaceousto char-
taceous,rarelycoriaceous,midsegmentrelative-
ly long,apexacuminate,base subcordateto deep-
ly cordate;uppersurfacescabrous,yellow-hirsute
to -hirtellous on the main veins, elsewhere
Flora Neotropica
sparsely so or only hispidulous, lower surface
pale yellow-hirtellouson the veins, white, arach-
noid hairs in the areoles or sometimes also on
the smallerveins; lateralveins 14-17 pairs, the
basal pair usually branched, tertiary venation
prominentbeneath;petiole 8-27 cm long, whit-
ish-hirsuteto -hirtellousto -subhispid,at the base
swollenand saccate;stipules2-7.5 cm long, out-
side yellowish-hirsute,inside sparselyhairy,ca-
ducous. Staminate inflorescences up to 12 cm
long and 6 cm wide; peduncle 1-3.5 cm long,
peduncleand branchespuberulousor partlyhir-
tellous;flowerssessile,clusteredin globoseheads,
these 2-3 mm diam.; tepals ca. 0.5-1 mm long,
basallyconnate,hirtellous;filamentsshorterthan
the tepals. Pistillate inflorescencesbranched,in
fruit up to 8 cm long and 7 cm wide; peduncle
3-4 cm long, indumentor peduncleand branches
similar to that of the staminate inflorescences;
flowers 8-ca. 30; pedicel up to 0.3 cm long, in
fruit up to 0.5 cm; perianth 0.3-0.4 cm long,
(densely) hirsute-hirtellous,usually arachnoid
hairs near the apex; stigma peltate, ca. 1.5 mm
diam., hirtellous.Fruitingperianthpurple,ovoid
to ellipsoid, 1-1.5 cm long, puberulous, also
sparselyyellow-hirsute,sometimes with brown,
pluricellularhairs.
Distribution(Fig. 74). Colombia (Amazonas),
Peru (Loreto), and Brazil (Amazonas);in non-
inundatedforest.
Specimens examined. COLOMBIA. AMAZONAS:
CantoAduche,12 Nov 1981(2),La Rotta101 (herb.
Araracuara),4 Mar 1982 (2),La Rotta 139 (herb.Ara-
racuara).
PERU. LORETO: 13 Jan 1976 (2),Gentryet al. 15876
(U); PuertoAlmendras,11 Nov 1984 (2), Maas et al.
6249 (U), 13 Jan 1976 (2),McDanielet al. 20436 (U);
Rio Nanay, PuertoAlmendras,29 Nov 1976 (2), Re-
villa 1966 (U).
BRAZIL.AMAZONAS: Mun. Tefe, Lago de Tefe, 14
Dec 1982 (8),Amaralet al. 76113 (BG);Rio Cuieras,
mouth of Rio Branchinho,Sep 1973 (st), Berg 276
(MO, U); upper Rio Alalau, 24 Feb 1968 (2), Bovan
267 (INPA);BR-17, km 11, 4 Aug 1955 (6), Chagas
(INPA) 1564 (INPA, U); Manaus-Mindfird., 3 Nov
1955 (9), L. Coelho(INPA)2265 (INPA);Mun. Jutai,
Copessfi,nr. P6rto Alfonso, 24 Oct 1986 (2), Daly et
al. 4129 (BG), 24 Oct 1986 (6),Daly et al. 4135 (BG);
Manaus,Aleixo rd., 29 Dec 1942 (9),Ducke1149 (MG,
MO, NY, R, US), 23 Sep 1945 (6), Ducke 1764 (A, F,
MG, NY, R, US), 19 Sep 1929 (2), Ducke(RB) 23606
(RB, US); Manaus,Cachoeirado Mindu, 8 Jul 1929
(8),Ducke(RB)23607 (G, P, RB, S, lectotypecollection
of P. myrmecophila);Manaus,BR-17, km 3, without
date (a),Francisco(INPA)2186 (INPA);nr. P6rtoVel-
ho, 8 Sep 1923 (a), Kuhlmann (RB) 19843 (B, U);
Pourouma 151

~~~~~~~~~~~~~~~~~~~~~~~~..

44deS A._ S 4V

?Cm Ira= BDT~OANWAL GARDENN

AMAZ6.%-I
VMVMW~TNACIONAL DE PESQUMA%A DA
gftt. at A? .m

-Gd , - v~~~~~~~~~~~~~T?e
m e h. y tg
' S Anto NAntni
,24313 x 22 depAbonIariA ,M-.anaus-I
d-.meterl
Pouroum&220 aForest
Crcrintuits on terra

? brown.
mateat Abonari. I:anaur,-
S:nto Antonio de
Forest an terra
C rac&raj, Km 220.

br&ct brown.
Prne,RJ
G.T. vember 2619?
Negat've imeiach Coelho- J.F. Ramosj_
L,F,~L.F ol.
... _ .ln:
.% '1??~1 11 55551 31| 51wf*-ffi0stI....
nr. C=ll 21cr rwr ar
JtPAN n^
: 71i~ 21RK 3F1N41
1w

myrmecophila. LeafS twig with pistillate inflorescences (Prance et al. 24313).

FIG. 71. Pourouma

Manaus, Reserva Florestal Ducke, 6 Dec 1976 (st),
Nascimentoet al. (INPA)66396 (INPA);Manaus-Ita-
coatiarard., km 192, 13 Sep 1974 (6), Penningtonet
al. P.22647 (INPA, NY); Humaita-Labreard., km 80,
27 Jan 1970 (9), Pranceet al. 3253 (F, INPA, NY, R,
S, U, US); Rio Cuieras,mouth of Rio Brancinho,24
Sep 1971 (d),Prance et al. 14795 (F, INPA, NY, U);
Rio Javari, Attalaia, 14 Oct 1976 (9), Prance et al.
23776 (G, INPA, MG, NY, U); Manaus-Caracarai rd.,
km 220, SantaAntonio de Abonari,26 Nov 1976 (2),
152 FloraNeotropica
Pranceet al. 24313(INPA,MO,NY, S, U, US);Ma- perianth(sparsely)hirsute;stigmapeltate,ca. 1.5
naus-Itacoatiara rd.,km79, 1 Sep1965(8),Rodrigues cm diam., subhirsute.Fruitingperianthovoid to
et al. 7069 (U), km 74, 3 Sep 1965(d), Rodrigues et
al. 7076(INPA). ellipsoid, ca. 1.5 cm long, blue-black, yellow-
hirsute.
Vernacularnames.Brazil.Amazonas:embau- Distribution(Fig. 74). Brazil (Amazonas);in
baranaor imbafbarana. non-inundatedforest.
From the syntype collections of P. myrmeco-
Specimensexamined. BRAZIL. AMAZONAS: Mun.
phila, Ducke (RB) 23606 and 23607 and Kuhl- Sao Paulode Olivenca,RioJandiatuba,27 Nov 1986
mann (RB) 19843, Ducke(RB) 23607, is chosen (9),Dalyet al. 4456(BG);nr.SaoPaulode Olivenca,
as the lectotype collection. 5 Dec 1986(8),Dalyet al. 4485(BG);Tonantins, Rio
The collection Kuhlmann (RB) 150082, ac- Solim6es,7 Feb 1944(9),Ducke1917(F,NY, R, type
collectionof P. formicarum);Sao Antoniade Ia, 2
cordingto the label from Rio de Janeiro,is prob-
either from Amazonas or was cultivated in May 1945(8).Fr6es20870 (NY, US), 19 Aug 1973
ably (8).Lleraset al. P.17411(F, INPA,MO,NY, S, U,
Rio de Janeiro. US);Carauari, PocoJurua,7 Jul 1980(8),A. S. Silva
Vegetativeparts(e.g., the wood) are (strongly) et al. 476 (BG,INPA).
aromatic, smelling like spearmintor, according Vernacularnames. Brazil.Amazonas:imbai-
to other labels, like "bengue."
binha, mapati.
This species matches P. myrmecophilain the
8. PouroumaformicarumDucke, Trop. Woods swollen and saccatebase of the petiole, a feature
90: 9. 1947. Type. Brazil.Amazonas:Rio So- connectedwith myrmecophily.The two species
limoes, Tonantins,7 Feb 1944 (9),Ducke 1917 also show strongresemblancesin the indument,
(not 1916!) (holotype, R, not indicated by staminateinflorescencesand flowersin the pis-
Ducke, designatedhere; isotypes, F, NY). tillateflowers.Theyarequitedistinctin the shape
Fig. 72. of the lamina and in the length of the petiole.
"The branchletsare inhabitedby stinging'ji-
Tree, up to ca. 10 m tall, or shrub, myrme-
quitaia' ants which nest among the hairs"(Daly
cophilous. Leafy twigs ca. 6 mm thick, yellow- et al.
to brownish-hirsuteand with sparse,brownplu- 4456).
ricellularhairs.Laminaentire,subovate,(14-)18-
32 x 6-12 cm, chartaceous to subcoria- 9. Pouroumaphaeotricha Mildbraed, Notizbl.
Bot. Gart.Berlin-Dahlem10:193.1927. Type.
ceous, apex abruptlyshort-acuminate,base ob- Peru. Loreto:Iquitos, 20 Aug 1925 (2), Tess-
tuse to subacute;upper surfacescabrousto sca-
mann 5364 (holotype, B; isotypes, S, US).
bridulous,yellow-hirsute,lowersurface+ densely
yellow-hirsuteon the main veins to (very)sparse- Fig. 73.
ly hirsute on the smaller veins, arachnoid hairs Tree, up to 10 m tall. Leafy twigs 4-6 mm
in the areoles and on the smaller veins; lateral thick, yellow-sericeousto -hirtellous, and with
veins 10-20 pairs, the basal pair unbranched, dense long, brown pluricellularhairs. Lamina
tertiary venation slightly prominent beneath; entire, oblong to obovate, 8-23 x 3-9 cm, co-
petiole 1-1.5 cm long, yellow-hirsute,at the base riaceousto subcoriaceous,apex acuminate,base
swollenand saccate;stipules3-5 cm long,outside acute to rounded;upper surfacesmooth or sca-
yellow-hirsute,inside glabrous, caducous. Sta- brous,yellow-hirtellouson the main veins, lower
minate inflorescencesup to 12 cm long and to 6 surfaceyellow-hirsuteon the veins and sparseor
cm wide; peduncle 2-4 cm long, peduncle and sometimes dense, orange, pluricellular hairs,
branches yellow-hirtellous to -hirsute;flowers arachnoidhairsconfinedto the areolesor almost
sessile, clusteredin globose heads, these 2-3 mm absent; lateral veins 8-10 pairs, basal pair un-
in diam.; tepals ca. 1 mm long, almost free, yel- branched,tertiaryvenation prominentbeneath;
lowish-hirtellous;filaments shorterthan the te- petiole 2-6 cm long, indumentsimilarto that of
pals. Pistillate inflorescencesbranched, in fruit the twigs;stipules1-4.5 cm long, outside yellow-
up to 7.5 cm long and 3.5 cm wide; peduncleca. sericeousto -velutinous, inside sparselyyellow-
4 cm long, peduncleandbrancheswith indument hairy to glabrous,caducous.Staminate inflores-
similar to that of the staminate inflorescences; cencesup to 10 cm long and 6 cm wide;peduncle
flowers 5-6; pedicel in fruit up to 0.5 cm long; 1.5-3 cm long, white- to yellow-sericeousto -ve-
Pourouma 153

R.:

17. 411...
. ur Uri? fr)i' -r : *
If*,,r,
, '*^>r i /'.:
X>.t'i. L:
E.< . ?J.1 W.C... . c O,, k..y
.
penin* .l betLw.ei
P Cul *.
;i5 Behi m Ant r .i e J .

FG Poru a'o1r1ma1m.11 1ea i 11gw h s i InfloFre"nce tiete.s e

72_..

'__A 35 likl , i* -.
1 . ......
212*.

FIG. 72. Pouroumaformicarum. Leafy twig with staminate inflorescences (Lleras et al. P.17411).

lutinous, and with dense (but on the ultimate petals.Pistillateinflorescencesin fruitup to 9 cm

branchessparse)brown,pluricellularhairs;flow- long and 8 cm wide; peduncle 2-5 cm long, pe-
ers sessile, clusteredin globose heads, these ca. duncle and branchesyellow- to white-hirtellous
2 mm diam.; tepals ca. 0.7 mm long, free or to sericeous;flowers 11-15; pedicel 0.1-0.2 cm
basallyconnate,hairy;filamentsshorterthan the long, in fruit up to 0.9 cm; perianthca. 2 mm
154 Flora Neotropica

HAtM OF PLRU

Pourouma phaeotricha mildbr.

Maynas: Dtto. Iquitos. lio :lar.na,

Caserio de Iishana trail to :
IItjt I( :,
|tg
N ~ative i?nch 1 2 de Yarana, uplands, "uvilla",.
green.
FIG.73. Porio p ec fL.
:- R~oX// r-5355S 1 I21 31 41 5 Dececber 4, 1976
II all C Manuel Rimachi Y. 2725
/ . -,,,,,,,I,,,h,,,l.,,I,,,,

FIG. 73. Pouroumaphaeotricha.Leafy twigs with pistillateinflorescences(Rimachi Y. 2725).

Pourouma
long, yellow-velutinous,with long, brown,pluri-
cellularhairs;stigma peltate, ca. 1.5 mm diam.,
yellow-puberulous.Fruitingperianthovoid to al-
most ellipsoid, 1.1-1.5 x 0.7-0.9 cm long, pu-
berulous.
Distribution(Fig. 85). Peru(Loreto)and Brazil
(Amazonas,along Rio Javari);in non-inundated
forest.
Specimensexamined.PERU. LORETO:
Rio Nanay,
Casariode Mishana3 Dec 1976(9),Davidson5203
(MO,U, US),4 Dec 1976(9),Rimachi2725(DUKE,
F, NY);Iquitos,3 Dec 1976(9),Tessmann5364 (B,
S, US, typecollectionof P. phaeotricha).
Rio Javari, Estirao do Ec-
BRAZIL. AMAZONAS:
uador,8 Aug1973(8),Lleraset al. P.17242(F,INPA,
MO,NY, U, US),21 Oct1976(v),Pranceet al. 23966
(G, INPA,NY, U).
Vernacularnames.Peru. Loreto:Sachauvilla,
uvilla.
The features of the staminate inflorescences
and flowersand the pistillateflowerssuggestclose
relationshipsto P. heterophyllaand P. formica-
rum. However, P. phaeotrichalacks the swollen
petiole base.
The two collections from the Brazilianbank
of Rio Javarihave leaves with a scabrousupper
surfacein contrastto the other collections.
10. PouroumamollisTrecul,Ann. Sci. Nat. Bot.,
Ser. 3, 8: 102, 1847; Miquel in Martius, Fl.
bras.4(1): 127. 1853; Berg,Fl. Suriname5(1):
272, t. 15b. 1975. Type. FrenchGuiana.With-
out locality,withoutdate (2),Leprieurs.n. (lec- anth black or purplish, ovoid to ellipsoid or to
totype, P. chosen by Berg, Fl. Suriname5(1):
272. 1975; isolectotype, G).
Tree, up to 30 m tall. Leafy twigs 4-10 mm
thick, rather sparsely to densely white-puberu-
lous (to -subvelutinous)to yellow-hirsute, and
with sparse to dense, purple to reddish-brown,
pluricellularhairs, occasionally sparse, white,
arachnoidhairs. Lamina entire, ovate to elliptic
(or obovate), 6-25(-35) x 4-20(-30) cm or 3-5-
lobed to -parted, ca. 10-25(-40) x 10-25(-40)
cm, coriaceous(to subcoriaceous),apex acumi-
nate, base acute, rounded to truncate, or sub-
cordate;upper surface smooth, sometimes sca-
brous, hairy on the main veins or subglabrous, 1. Laminausuallyentire;pistillateinflorescences
occasionally yellow-hirsute, lower surface,
to to hirtellous to clusters;Guianas,LowerAmazonBasin,and
sparsely densely puberulous
subtomentose,often partof the hairson the main 1. Laminausually3-5 parted;pistillateinflores-
veins appressed,arachnoidhairs in the areoles,
155
mostly also on the smaller veins, occasionally
also on the leaf margin;lateralveins (6-)10-25
(-30) pairs,basalpairbranchedor sometimes(in
obovate to elliptic leaves) unbranched,the basal
part of these veins formingthe basal part of the
leaf margin(thus not separatedfrom the margin
by mesophyll), reticulumand tertiaryvenation
(rather)planebeneath;petiole 3-22(-35) cm long,
sparselyto denselywhite-puberulous(to -subve-
lutinous)to yellow-hirsute,occasionallyalso with
longwhite,arachnoidhairs;stipules3-16 cm long,
outsideyellow(ish)-(sub)velutinous to -hirsuteto
-(sub)sericeous,inside glabrous or sometimes
sparsely hairy, caducous. Staminate inflores-
cencesup to 10 cm long and 5 cm wide;peduncle
2-6 cm long, peduncleand brancheswhitish- to
yellow-puberulous(to -subvelutionous)to -hir-
tellous;flowerssessileor pedicellate,most of them
in globose heads, these 3-4 mm diam.; tepals
0.5-1 mm long, connate, forminga ? urceolate
perianth,hairy; filaments longer than the peri-
anth, free. Pistillate inflorescencesin fruit up to
18 cm long and 10 cm wide; peduncle 3-12 cm
long,peduncleand brancheswith indumentsim-
ilarto that of the staminateinflorescence;flowers
10-30, in two clustersor more diffuselydistrib-
uted in the inflorescence;pedicel up to 1 cm, in
fruitup to 1.5 cm long, often alreadyat anthesis
with a strongly (+ capula-like)widened apex;
perianth 0.5-1 cm long, yellow(ish)-(sub)ve-
lutinous, apex ? tuberculate;stigma peltate, ca.
2 mm diam., white-puberulous.Fruiting peri-
oblongoid, 1-2 x 0.7-1.5 cm, usually densely
hairy.
This species is very closely related to P. me-
linonii. We had consideredrecognizingthe two
taxa only at the subspecificlevel, but the lack of
intermediatesin areas where the two taxa grow
together indicates that they are probablygood
species.
Two allopatricsubspeciescan be recognized.
Key to the Subspecies of
Pourouma mollis
with the flowersin two, moreor less distinct
easternBrazil................. a. subsp.mollis.
cencesusuallywiththe flowersdiffuselydistrib-
156
uted;UpperAmazonBasinto easternColombia.
............................ b. subsp. triloba.
lOa. Pourouma mollis Trecul subsp. mollis.
Fig. 75b-f.
Lamina usually entire, sometimes 3-lobed or
to 5-parted, subjuvenile material usually 3-5-
parted; upper surface smooth, with hairs on the
main veins or subglabrous. Pistillateflowers ar-
ranged in two + distinct clusters, due to the
shortening of one (or two) of the primary branch-
es and the short pedicels, being at fruit up to 0.7
cm long.
Distribution (Fig. 74). Eastern Guyana, Suri-
nam, French Guiana, Amazonian Brazil
(Amampa and Para), and eastern Brazil (Bahia
and Espirito Santo); in non-inundated forest at
low altitudes.
Specimensexamined.GUYANA. EssequiboR., nr.
Rockstone, 1 Aug 1921 (juv), Gleason670 (GH, NY,
US), 4 Nov 1979 (2), Maas et al. 3938 (U).
SURINAM. PeninicaCreek,20 Apr 1950 (st), BW
1128 (U); Sectie 0, 15 Sep 1916 (st), BW2410 (NY),
U), 19 Dec 1919 (2), BW 4494 (NY, U), 25 Feb 1921
(2), BW5048 (U, UC), 15 Nov 1921 (2), BW 5440 (A,
RB, U, US), 28 Jan 1922 (9), BW 5581 (MO, U), 24
Nov 1922 (2), BW 5992 (F, U); nr. Gansee, 26 May
1964 (juv), Donselaar 1373 (U); nr. Brownsberg,14
Oct 1964 (st), Donselaar 1681 (U), 4 Jan 1966 (juv),
Donselaar2924 (U);GranDam, 13 Oct 1966 (st),Don-
selaar 3688 (U); MapaneCreek,Feb 1964 (2), Elburg
(LBB) 9843 (MO, NY, U), 7 Mar 1963 (2), LBB 9424
(U); SurinameR., without date (8 and 9), Hostmann
(& Kappler)1272 (B, CGE, G, GH, LE, NY, OXF, P,
S, U, mixed with P. melinoniisubsp. melinonii);Nas-
sau Mts., 16 Feb 1949 (9), Lanjouwet al. 2106 (NY,
U); Jodensavanne,27 Mar 1953 (st), Lindeman3631
(U), 16 May 1953 (st), Lindeman 3989 (U), 20 Dec
1954 (st), Lindeman6923 (U).
FRENCHGUIANA. Massifdes Emerillons,source
of ApprouageR., 19 Sep 1980 (8), Cremers6726 (U);
upperCriqueNouciri, 13 Dec 1983 (st), Cremers8271
(U);withoutlocality,withoutdate (2), Leprieurs.n. (G,
P, lectotype collection of P. mollis); Orapu, 13 Oct
1966 (9),OldemanB.644 (CAY,P, U); pistede St. Elie,
km 16, 19 Apr 1982 (st), Riera 573 (U).
BRAZIL.Withoutlocality,withoutdate(a),Burchell
9792 (GH, P, the same collectionnumberis also used
for a collection of P. guianensis subsp. guianensis).
AMAPA: Serra do Navio, 1961 (st), Rodrigues2984
(INPA); between P6rto Platon and Serrado Navio,
1976 (st), Rosa 1158 (MG). BAHIA:Without locality,
without date (6), Blanchet2361 (or 94) (FHO, G, LE,
P); P6rto Seguro,20 Sep 1968 (6), Almeida et al. 71
(NY, U); Agua Preta, 23 Feb 1937 (9), Bondar 2169
(GUA); Rio das Contas,3 Oct 1919 (8), Curran21 (C,
F, NY, US); Ilheus, 1848 (8),Luschnaths.n. (BR),Sep
1821 (a),Riedel2 (LE);CasteloNovo, 15 Sep 1944 (2),
Velloso 1083 (R). ESPiRITOSANTO:Linhares, 18 Aug
Flora Neotropica
1978 (6), DAF 027 (GUA); Reserva de Linhares,30
Oct 1930 (6), Kuhlmann446 (U); Linhares,Corriogo
de Durao, 2 Jan 1972 (9), Sucre 8303 (U). PARA:Faro,
6 Jan 1920 (6), Ducke (RB) 13058 (RB); Obidos, 27
Dec 1913 (9), Ducke(MG) 15261 (MG);Belem, 1 Feb
1928 (6), Ducke (RB) 19455 (RB); Rio Guama; Sao
Miguel, Oct 1906 (6), Goeldi(MG) 7731 (G, MG, S,
U); AcaraiMts., Rio Taruini,20 Nov 1952 (6), Guppy
656 (=FD 7671) (NY, US); Belem, 19 Sep 1963 (6),
Oliveira2597 (IAN), 26 Sep 1963 (9), Oliveira3071
(IAN), 3 Sep 1966 (3),Pires 10123 (IAN), 24 Aug 1967
(), Pires et al. 10578 (IAN), 2 Sep 1967 (6), Pires et
al. 10809 (IAN), Sep-Oct 1961 (9), Pires 51676 (F,
NY, US); Cuiaba-Santar6mrd., km 919,13 Nov 1977
(9), Prance et al. 25337 (F, MG, RB, U); Mun. Al-
meirim, Monte Dourado, 2 Dec 1978 (6), Santos 455
(U); Belem, 12 Jan 1966 (6),M. Silva 354 (MG),4 Oct
1942 (6), M. B. Silva 130 (NY, US).
Vernacular names. Surinam. Boroma (Aro-
wak), bospapaja, granboesipapaja, yarayara
(Carib).French Guiana. Bois canon male, bois
canon sauvage. Brazil. Amapa: uva de macaco.
Bahia: itararanga,tararanga,tararangavermel-
ha; Maranhao: ama'yrary (Ka'apor); Para: im-
bauiba,imbauibarana vermelha, mapatirana,ka-
moyuwa (Wai-wai),sa-ouro (Mawayan).
In Amapaand Parathis subspeciescan be con-
fused with a form of P. melinonii subsp. meli-
nonii with patenthairs on the lower leaf surface.
However, the latter has sparse, appressed hairs
on the leafy twigs and stipules and the flowers in
the pistillate inflorescence are diffusely distrib-
uted, thus not in two clusters as in subsp. mollis.
Hostmann & Kappler 1272 (6) belongs to P.
melinonii subsp. melinonii while Hostmann (&
Kappler) 1272 (9 and 6) belongs to P. mollis subsp.
mollis.
The description of P. mollis was based on the
collections Blanchet 2361 (from Brazil, Bahia);
Hostmann (from Suriname), and Leprieur 141.
The latter was selected as the lectotype collection
(Berg, 1975).
1Ob. Pourouma mollis Trecul subsp. triloba
(Trecul) C. C. Berg & van Heusden, Proc. Kon.
Ned. Akad.Wetensch.,Ser.C, 91(2): 107. 1988.
PouroumatrilobaTrecul, Ann. Sci. Nat. Bot., Ser. 3,
8:104. 1847 (Aug);Miquelin Martius,Fl. bras.4(1):
126. 1853; Macbride,Publ. Field Mus. Nat. Hist.,
Bot. Ser., 13(2.2):294. 1937. Type. Peru.Huanuco:
Macora,without date (6!), Ruiz & Pavon s.n. (lec-
totype P, chosen here;isolectotypes,G, OXF, US).
PouroumajussiaeanaTrecul,Ann. Sci. Nat. Bot., Ser.
3, 8: 106. 1847; Miquel in Martius,Fl. bras. 4(1):
124. 1853. Type. Peru. Buena Vista, without date
(2), Anonymuss.n. (holotype,P).
 P. cucura P. cuspidats
.?

GP. 7rorPcabrL P

leo myrsoMaphile P.

subsp. tizab

r~~~~~~~~~~~~~~~~~~~
'' ? ~~~~ulSlsp.
grlabrat

~~~I 'I ~~~~Ubp. mebli inmii

FIG. 74. Distribution of Pourouma cucura, P. cuspidata, P. formicarum, P. melinonii subsp. glabrata, P.
melinonii subsp. melinonii, P. mollis subsp. mollis, P. mollis subsp. triloba, and P. myrmecophila.
158 Flora Neotropica

~, a-

e t~~~e7 f~~
Pourouma159

Pourouma triloba Klotzsch, Linnaea 20: 526. 1847 V.10650(BG).UYACALLI: Prov.CoronelPortillo,Pu-

(Sep). Type. Peru. Huanuco:Macora,without date callpa-TingoMariard., 1980(9),Trucios9 (BG).
(9!),Ruiz & Pavon s.n. (holotype, B; isotypes, BR, BRAZIL.ACRE: UpperRio Moa,nr.FazendaAri-
G, OXF). zona,24-30 Sep1984(st),Campbell et al. 8383(BG);
Pouroumajaramilloi Cuatrecasas,Caldasia 7: 300. mouthof RioMacauhan, 5 Aug1933(d),Krukoff5309
1956. Type. Colombia.Meta:Sierrade la Macarena, (F,G, GB,M,MO,NY,S,U, UC,US);SenaMadurei-
CafnoCiervo, 19 Jan 1950 (2), Philipson&Jaramillo ra, Rio Caete,7 Oct 1968(d), Pranceet al. 7907 (F,
2133 (holotype,BM, not seen; isotypes, F, US). GH,NY, P, R, S, U, US).
Lamina usually 3-5-parted, sometimes entire; Vernacularnames.Peru.Amazonas:suia, suir
upper surface sometimes scabrous; petiole oc- shuina (Huambisa),uhukamsuiya;San Martin:
casionally with white-arachnoid hairs. Pistillate obija, uvilla.
flowers usually more or less diffusely distributed This subspeciesis more variable than subsp.
in the inflorescence, sometimes in two ? distinct mollis and may prove to consist of severalinfra-
clusters; pedicels in fruit up to 1 cm long. specific taxa. In particularHuashikat 944, Hu-
Distribution (Fig. 74). Colombia (Meta), Peru ashikat 1027, Tunqui296 (all from Peru, Ama-
(Amazonas, Loreto, Madre de Dios, Pasco and zonas, valley of Rio Santiago)and Gentryet al.
San Martin), and Brazil (Acre); in non-inundated 15621 (from Loreto, Rio Nanay) are a distinct
forest at low altitudes. set of collections. They have entire leaves with
a ? scabrousupperleaf surface,the basal lateral
Specimensexamined.COLOMBIA.META:SanJuan veins are often unbranched,and the stipules are
de Arama, Rio Guejar,26 Jan 1951 (9), Idroboet al.
1316 (US);Sierrade la Macarena,CainoCiervo, 19 Jan ratherdensely hairy inside.
1950 (9),Philipson&Jaramillo2133 (BM, P, US, type Prance et al. 7907 has relatively many lateral
collection of P. jaramilloi). veins, up to ca. 30 pairs, whereasin other col-
ECUDADOR.NAPO:Lumbaqui,25 Nov 1987 (8), lections there are at most 25 pairs.
Penningtonet al. 12300 (BG). Schunke V. 7729 and 10650, Foster 5791, D.
PERU.Withoutlocality,withoutdate(9),Anonymus
s.n. (P, type collection of P. jussiaeana). AMAZONAS:N. Smith 5134, and Trucios9 have entireleaves
Rio Cenepa,30 Dec 1972 (9),Berlin746 (F, GH, MO); and the pistillate inflorescencestend to the type
Rio Santiago,nr.Caterpiza,15 Oct 1979 (9),Huashikat of inflorescencenormal in subsp. mollis.
944 (U), 24 Oct 1979 (2), Huashikat 1027 (BG); Rio The descriptionof P. trilobaTr6culwas based
Huampi, 26 Jul 1974 (8), Kayap 1306 (MO, US); Rio on
Santiago,nr. Caterpiza,12 Dec 1979 (9), Tunqui296
both staminateand pistillatematerialcollect-
(BG).HUANUco:Tingo Maria, 13 Jul 1957 (st), Ellen- ed by Ruiz & Pavon, the staminate material is
berg2297 (U); Macora,withoutdate (6),Ruiz & Pavon chosen as the lectotype collection, the pistillate
s.n. (G, OXF, P, US, lectotypecollection of P. triloba materialhas been used for the descriptionof P.
Tr6cul),withoutdate (2), Ruiz & Pavons.n. (B, BR, G, trilobaKlotzsch.
OXF, type collectionof P. triloba
Klotzsch);Pampaya-
cu, 5 Jun 1927 (d),Sawada21 (F). JUNiN:Prov. Satipo,
GranPajonal,S of Chequitaro,22 Sep 1983 (8),D. N. 11. PouroumamelinoniiBenoist,Bull.Mus.Hist.
Smith 5134 (BG).LORETO: Prov. Maynas,Rio Nanay, Nat. (Paris)28: 318. 1922; Berg,Fl. Suriname
PuertoAlmendras,5 Jan 1976 (9), Gentryet al. 15621
5(1): 274, t. 15a. 1975. Type. FrenchGuiana.
(BG);22 km S of km 86 on Pucallpa-TingoMariard., La Mana, Sep 1846 (9), Sagot 990 (lectotype,
11 Feb 1981 (2), Gentryet al. 31179 (BG);Prov. May-
nas, YanamonoExploramaTouristCamp,27 Jun 1983 P. chosenby Berg,Fl. Suriname5(1):274.1975;
(st), Gentryet al. 42232 (BG), 8 Jul 1983 (st), Gentry isolectotype,B).
et al. 42793 (BG);Yurimaguas,Oct-Nov 1929 (6),LI.
Williams4688 (F). MADRE DEDios: Rio Manu, Pak- Tree, up to 30 m tall. Leafy twigs 4-12 mm
itza Station, 20 Nov 1980 (9), Foster 5781 (BG, F); thick, sparselywhitish, mostly appressed-puber-
Rio Manu, CochaCashuStation,Jul 1978 (st), Foster ulous to strigose, and with purple to reddish-
et al. 6569 (F). SANMARTIN: Prov. MariscalCaceres,
Rio Uchiza, 24 Jul 1974 (a),J. Schunke V. 7729 (MO, brown,pluricellularhairsand occasionallysparse,
NY, U); Prov.MariscalCaceres,Dist. TocacheNuevo, white, arachnoidhairs. Lamina entire, ovate (to
Quebradade Ischichini, 12 Oct 1978 (2), J. Schunke elliptic) or 3-lobed, 6-25(-35) x 4-16(-27) cm

FIG.75. a. Pouroumamelinoniisubsp.melinonii.Leafytwigwithpistillateinflorescences(Irwinetal. 47832).

b-f. P. mollis subsp. mollis. b. Leafytwig with pistillateinflorescences(Elburg(LBB)9843). c. Staminateflower
(Pireset al. 51625). d. Pistillateflower.e. Ovule f. Seed and embryo (BW 5581).
160
or 3-5-parted and ca. 10-27 x 10-27 cm, co-
riaceous, apex acuminate, base acute, rounded
to truncateto subcordateor cordate;upper sur-
face smooth, hairy on the main veins or subgla-
brous, lower surface(rather)sparselyappressed-
puberulousto strigose or patent-puberulousto
hirtellousor subglabrouson the veins, arachnoid
hairs in the areoles or also on the smallerveins;
lateral veins 12-20 pairs, basal pair branched,
the basal part of these veins forming the basal
part of the leaf margin(thus not separatedfrom
the marginby mesophyll), tertiaryvenation al-
most plane beneath;petiole 3-22 cm long, stri-
gillose to strigose or puberulous to hirtellous;
stipules(2-)4-17 cm long, outsidesparselywhite-
strigose, inside glabrous, caducous. Staminate
inflorescencesup to 12 cm long and 12 cm wide;
peduncle 2-6 cm long, peduncle and branches
sparsely to rather densely puberulous;flowers
sessile or pedicellate, most of them in globose
heads, these 3-4 mm diam.; tepals 0.5-1 mm
long, connate, forming a ? urceolate perianth,
sparselyto densely hairy; filamentslonger than
the tepals, free or + connate. Pistillate inflores-
cencesin fruitup to 17 cm long and 11 cm wide;
peduncle 3-12 cm long, peduncle and branches
with indument similar to that of the staminate
inflorescence;flowers 10-30; pedicel up to 1 cm,
in fruit up to 1.5 cm long; perianth 0.5-1 cm
long, (rather)sparsely puberulous, apex tuber-
culate;stigma peltate, ca. 2 mm diam. Fruiting
perianth dark violet or dark maroon, ovoid to
ellipsoidto oblongoid, 1-2 x 0.7-1.5 cm, sparse-
ly to densely puberulous to subvelutinous or
subglabrous.
This species is very closely relatedto P. mollis
from which it is not alwayseasy to separate.One
of the main reasons for treatingP. melinonii as
distinct at the species level is the co-occurrence
of P. mollissubsp. mollisand P. melinoniisubsp.
melinoniiin the Guianaswithout indications of
interbreeding.P. melinoniican be distinguished
from P. mollis by the sparse indument in most
parts of the plant, but especiallyon the stipules;
only the fruitingperianthis sometimes densely
puberulousto subvelutinous. The sparse indu-
ment on the stipules and leafy twigs of P. meli-
nonii is whitish and usually appressed,whereas
the dense indument on the stipules and leafy
twigsof P. mollis is yellow and usually + patent.
Within P. melinoniitwo allopatricsubspecies
can be recognized.
FloraNeotropica
Key to the Subspecies of
Pourouma melinonii
1. Lamina usually entire, if 3-5-parted, then the
baseusuallytruncateto subcordate;fruitingperi-
anth sparselyor densely hairy;stamens usually
with freefilaments;AmazonBasinand Guianas.
.......................... a. subsp. melinonii.
1. Lamina3-5-parted,basemoreor less deeplycor-
date; fruiting perianths subglabrous;stamens
(often) with connate filaments; Panama and
northernand westernColombia..............
........................... b. subsp. glabrata.
lla. Pourouma melinonii Benoist subsp.
melinonii. Fig. 75a.
PouroumaapaporiensisCuatrecasas,Caldasia7: 297.
1956. Type. Colombia. Amazonas-Vaupes:Rio
Apaporis,Lagunasdel Chucuro,22 Nov 1951 (9),
Garcia-Barriga13643 (holotype,US; isotype, NY).
Pourouma apaporiensis Cuatrecasas forma macro-
phyllaCuatrecasas,Caldasia7:298. 1956. Type.Co-
lombia. Vaup6s:Rio Cubiyi, 8 Dec 1943 (9), Allen
3251 (holotype,US, isotype, MO).
Leafy twigs hairy or subglabrous. Lamina
mostly entire with a rounded to truncate base,
sometimes 3-lobed with a truncate to subcordate
base, occasionally 3-parted with a deeply cordate
base; lower surface on the veins ? sparsely hairy,
mostly with appressed hairs, sometimes with
patent hairs. Stamens with (almost) free fila-
ments. Fruiting perianth sparsely or + densely
puberulous to subvelutinous.
Distribution(Fig. 74). Colombia (Amazonas),
Venezuela (Amazonas and Bolivar), Guyana,
Surinam, French Guiana, Peru (Amazonas and
Loreto), and Brazil (Amapa, Amazonas, Mato
Grosso, and Para); in non-inundated forest at
low altitudes.
Specimensexamined.COLOMBIA.AMAZONAS: Rio
Apaporis,betweenRio Pacoa and Rio Kananari,15-
19 Dec 1951 (9), Garcia-Barriga14116 (NY, US).
AMAZONAS-VAUPES: Rio Apaporis,Lagunasdel Chu-
ruco,22 Nov 1951(9),Garcia-Barriga13643 (NY, US,
type collectionof P. apaporiensis).VAUPES:Cabeceras
Rio Cubyyfi,8 Dec 1943 (9), Allen 3251 (MO, type
collectionof P. apaporiensisForma macrophylla).
VENEZUELA. AMAZONAS: Upper Rio Orinoco,
1951 (9), Croizat962 (US);base of CerroDuida, Jan-
Feb 1969 (st), Farinas et al. 490 (U, VEN); Rio Cai-
quiare,Rio Vacivaand Rio Yatua,Apr 1853(6),Spruce
3176 (C, CGE, E, GH, GOET, MG, NY, OXF, P).
BOLiVAR: Rio Chirea Uriman, 5 Aug 1953 (st), Ber-
nardi902 (NY, VEN); Rio Icabaruand Rio Hacha, 7
Jan 1956 (9),Bernardi2808 (G, NY); Mun. Urdaneta,
Rio Chinarok,10 Jan 1986 (9), Hernandez317 (BG);
Pourouma 161

betweenRio Chicananand Rio Ayaiche,24 Aug 1961 (Arawak),bospapaja,granboesipapaja,poeroe-

(6), Steyermark89452 (NY, VEN);SierraIchun, Salto ma or puruma(Carib),yarayara(Carib).French
Maria Espuma, 29 Dec 1961 (6), Steyermark90408
Guiana. Bois canon, papaye apici (Paramaka).
(NY); El Tigre, 2 May 1939 (9), LI. Williams 12008
(F, S, US, VEN). Peru. Amazonas:dakamtazshuiya,mimpashui-
GUYANA. PakaraimaMts., Kamanafalls, 25 Aug ya washi shuina;Loreto:uvilla. Brazil.Amazo-
1961 (6), Maguire et al. 45945 A (NY, US); upper nas: imbaibarana;Para:mapatirana.
MazaruniR., Kako R., 23 Sep 1960 (6), Tillettet al. The collectionfromAmazonianColombia,in-
45495 (U).
SURINAM. MarowijneR., withoutdate (6), (Host- cluding the type collection of P. apaporiensis,
mann &) Kappler1272 (LE, M, P); Grote Zwiebel- have deeply cordate leaves, like the materialof
zwamp, 22 Sep 1948 (2), Lanjouwet al. 399 (NY, U); subsp.glabrata,but in contrastto the other col-
Jodensavanne,4 Jul 1961 (st), Schulz (LBB) 8976 (F, lections of subsp. melinonii.Pistillatespecimens
U). from this area can be distinguishedfrom those
FRENCHGUIANA. Withoutlocality,withoutdata
(2), Aublets.n. (BM, S), 1986 (6), Melinon 457 (A, G, of subsp. glabratain the dense indument of the
P, U), 1863 (2),Melinons.n. (P), 1863 (6),Melinons.n. perianthof the pistillate flower.
(E, LE, NY, P, US), (8), Poiteau s.n. (LE), 1865 (st) The descriptionof P. melinoniiwas based on
Sagot s.n. (P);BolateeCreek,2 Nov 1948 (st),BAFOG the collections Kappler 1272 (from Suriname)
4272 (CAY, U);'St. Pierre Creek, 28 Feb 1959 (st),
Cantonnementde Cayennes.n. (CAY);Trois Sauts,28 and three collections made in French Guiana:
Oct 1974 (2), Lescure343 (CAY); Cayenne,without Sagot 517 (p.p.), Sagot 990, and Melinon 457.
date (2),Martins.n. (BR, F, MO, U); Cacao,S of Cay- Sagot 990 has been selectedas the lectotypecol-
enne, 21 Apr 1965 (6), Oldeman1243 (CAY);piste de lection (Berg,1975). The (Hostmann&)Kappler
St. Elie, 1Jun 1981(st),Sabatier51 (CAY);Acarouany,
May 1858 (6),Sagot 517 (F, P; underthe same number
collection has the same number as a collection
the lectotypecollectionof P. laevis);LaMana,Sep 1856 of Hostmann(&Kappler)of P. mollissubsp.mol-
(2), Sagot 990 (B, P, lectotype collection of P. meli- lis.
nonii), 1856 (2), Sagot 990bis (G); Godebert, 13 Jul
1921 (2), Wachenheim232 and s.n. (P).
PERU. AMAZONAS:
Monte Chichijam, 24 Apr 1973 lib. PouroumamelinoniiBenoistsubsp.glabra-
(2),Ancuash290 (F, GH, MO); Rio Santiago,nr..Ca- ta C. C. Berg& van Heusden,Proc. Kon. Ned.
terpiza,7 Sep 1977 (2),Huashikat431 (BG);Rio Cusu, Akad. Wetensch., Ser. C, 91(2): 108. 1988.
Yucui, 12 Mar 1973 (2), Kayap 571 (GH). LORETO: Type. Panama. Canal Zone: Pipeline Rd., 10
Rio Tacsha,Curaray,19 Sep 1972 (6), Croat20447 (C,
km NW of Gamboa, 14 Dec 1973 (9), Berg &
DUKE, F, GH, MO, NY).
BRAZIL. AMAPA: Rio Jaue, 3 km E of confluence Nee 355 (holotype,U; isotypes, BG, COL, K,
with Rio Oiapoque,26 Aug 1960 (2),Irwinet al. 47832 MO, NY, PMA). Fig. 76.
(GH, NY, U, US); Rio Mutura,22 Sep 1960 (2),Irwin
et al. 48439 (NY, U, US); Mun. Mazagao,81 km NW Leafytwigssubglabrous.Lamina usually5- or
of Macapa,20 Dec 1984 (2), Mori et al. 17478 (BG). 3-parted with a distinctly cordate base, some-
AMAZONAS: Mun. Tefe, Rio Solimoes, Lago de Tefe, times entireand then with a cordateto subacute
15 Oct 1982 (2),Amaral et al. 101 (BG);Rio Japura,
Vila Bittancourt,19 Nov 1982 (2), Amaral et al. 595 base; lower surface on the main veins subgla-
(BG);Manaus-Caracarai rd., km 148, 27 Sep 1973 (2), brous. Stamens often with + connate filaments.
Berget al. P. 18136 (F, INPA, MO, NY, P, S, U, US); Fruitingperianth(sub)glabrous.
Mun. Anori, rd. Amori-Anama, 27 Nov 1975 (8), D. Distribution(Fig. 74). Panamaand Colombia
Coelho et al. 672 (INPA); Rio Solimoes, Belem, 16
Dec 1948 (6), Fr6es 23725 (IAN), 25 Mar 1976 (st), (Antioquia);in non-inundatedforest at low al-
Mello s.n. (INPA); Manaus-Itacoatiarard., km 26, 2 titudes.
Sep 1966 (6), Prance et al. 2180 (F, GH, INPA, NY,
P, R, S, U, US). MATOGROSSO: Brasilia-Acrehwy., Specimens examined. PANAMA. CANALZONE:
215 km, west of Rio Juruena,2 Sep 1963 (6),Maguire PipelineRd.,ca. 10kmNWof Gamboa,14Dec 1973
et al. 56511 (F, NY, RB, U, US). PARA:Rio Guama, (9), Berg& Nee 355 (BG, COL,K, MO, NY, PMA, U,
Sao Miguel,Oct 1906 (2), Goeldi(MG) 7732 (G, MG); typecollectionof P. melinoniisubsp.glabrata),12Oct
Rio Jari, Monte Dourado, 16 Nov 1967 (2), Oliveira 1971 (9), Gentry2046 (MO);Gamboa, 21 Nov 1971
3736 (IAN), 22 Jan 1968 (2), Oliveira3936 (NY), 30 S of El Real, nr.
(9), Gentry2657 (MO, NA). DARIEN:
Canamine, 21 Aug 1987 (9), McPherson11510 (BG).
Jan 1961 (9), Oliveira4047 (NY), 9 Sep 1968 (2), N. SAN BLAS:Rio Cangandi,nr. Cangandi,18 Feb 1984
T. Silva 921 (NY, U), 11 Sep 1968 (6);N. T. Silva 949
(9), Nevers4929 (BG).
(NY, U), 16 Sep 1968 (2) N. T. Silva 995 (IAN, NY). COLOMBIA.ANTIOQUIA:38 km W of Barranca-
24 Feb1967(8),Bruijn1496(F,M,MO,NY,
bermeja,
Vernacular names. Venezuela. Bolivar: kai- S, U, US, VEN), 25 Feb 1967 (9), Bruijn 1502 (F, M,
warikai, yagrumo-sunsun. Surinam. Boroma MO,NY, S, U, US, VEN),1 Mar 1967 (st),Bruijn
162 Flora Neotropica

1546(F, M, MO,NY, S, U, US, VEN),2 Feb 1967 leafytwigs,andthe moreprominentsmallerveins

(st),Bruijn1555 (F, M, MO,NY, S, U, US, VEN); at the laminabeneath,and the broadersegments
Anori,6-12 Sep 1973(9), Soejarto4283 (MO). of the incised leaves. It differsfrom P. mollis in
Vernacularnames.Colombia.Antioquia:cirpe the cordate leaf base of the incised leaves and
macho. the more prominentsmallerveins on the lamina
beneath.
12. PouroumahirsutipetiolataMildbraed, No- Two (probably)allopatric subspecies can be
tizbl. Bot. Gart.Berlin-Dahlem10: 420. 1928. recognized.
Type.Colombia.Bolivar:Norosi-Tiquisotrail,
lands of Loba, Apr-May 1916 (Q),Curran116
Key to the Subspecies of
(holotype, US, fragmentin B). Pourouma hirsutipetiolata
Tree,up to ca. 30 m tall, with butresses.Leafy
1. Laminaentire;northernColombia...........
twigs 6-20 mm thick, yellow-hirsuteand with .................... a. subsp. hirsutipetiolata.
minute, whitish, appressedor patent hairs and 1. Lamina3-lobedto -parted;westernColombia
usually also with brown, pluricellular hairs. andEcuador................. b. subsp.hispida.
Laminaentire,ovate (to ellipticor suborbicular),
10-25 x 7-18 cm, or 3-lobed to -partedand ca.
10-35 x 10-35 cm, coriaceous,apex acuminate, 12a. PouroumahirsutipetiolataMildbraedsubsp.
base cordateor subcordateor sometimes round- hirsutipetiolata. Fig. 77.
ed; uppersurfacesmooth,hairyon the mainveins Lamina entire.
or subglabrous,lower surface (rather)sparsely Distribution(Fig. 81). Colombia (Antioquia,
strigoseto strigilloseon the veins or puberulous Bolivar,and Santander);in non-inundated,low-
to hirtellouson the smallerveins, arachnoidhairs land forest.
in the areoles, usually extending to the lateral
examined.COLOMBIA.ANTIOQUIA: 38
veins; lateral veins (5-)12-15 pairs, basal pair kmSpecimens
W ofBarrancabermeja, 1 Mar 1967(9),Bruijn1547
branched, tertiary and quarterary venation (F, M, MO, NY, S, U, US, VEN). BOLIVAR: Norosi-
ratherprominentbeneath;petiole 3-20 cm long, Tiquisio trail,Landsof Loba, Apr-May 1916 (2), Cur-
yellow-hirsute,usuallyalso with minute, whitish ran 116 (B, US, type collection of P. hirsutipetiolata).
hairs;stipules4-24 cm long, outside yellow-hir- SANTANDER: Bucaramanga,11 Dec 1977 (2), Renteria
sute (to -subsericeous), and also with much et al. 45 (MO);rd. to Citimarra,9 Dec 1979 (2), Ren-
teriaetal.2093(HUA);16kmSEof Barrancabermeja,
shorter,whitish hairs,inside glabrous,caducous. 26 Aug 1954 (9),Romero-Castaneda4715 (MO, US).
Staminate inflorescencesup to 10 cm long and
8 cm wide; peduncle 3-4 cm long, densely pub- Vernacularnames.Colombia.Santander:sirpe
erulousor also hirtellous;flowerssessile, most of or sirpo, urumo blanco.
them in subgloboseheads,these ca. 3 mm diam.;
tepals 1-1.5 mm long, connate, forming a 12b. PouroumahirsutipetiolataMildbraedsubsp.
(sub)infundibuliformperianth, often with an hispida(Standley& Cuatrecasas)C. C. Berg&
obliqueapex,sparselyhairy;filamentslongerthan van Heusden, Proc. Kon. Ned. Akad. We-
the tepals, free. Pistillate inflorescencesin fruit tensch., Ser. C, 91(2): 108. 1988. Fig. 78.
up to 22 cm long and 10 cm wide; peduncle 5- PouroumahispidaCuatrecasas,Standley& Cuatreca-
13 cm long; peduncle and branches minutely sasin Cuatrecasas, Caldasia7: 299. 1956.Type.Co-
puberulous,sparsely so on the peduncle, more lombia.Valle:Rio Anchicaya,LaPlanta,5 Aug 1943
densely on the branchesand very densely on the (9), Cuatrecasas14881 (holotype,F).
pedicels;flowers 7-17; pedicel 0.5-1 cm long, in Lamina 3-lobed to -parted.
fruit up to 1.5 cm long; perianth5-10 cm long, Distribution(Fig. 81). Colombia(Choc6,Valle)
densely short-velutinous;stigma peltate, ca. 1.5 and Ecuador (Esmeraldas);in non-inundated,
mm diam., white puberulous.Fruitingperianth lowland forest.
dark violet or black, ellipsoid to ovoid, ca. 1.5-
2 x 1-1.5 cm, densely minutely puberulous. Specimensexamined.COLOMBIA.CHocO:LaMo-
jarra, 5 Nov 1983 (2), Juncosa 1253 (BG).VALLE: Rio
This species is closely relatedto P. melinonii,
Anchicaya, La Planta, 5 Aug 1943 (2), Cuatrecasas
fromwhichit differsin the shapeof the staminate 14881 (F, type collection of P. hispida);Rio Calima,
flowers,the long yellow hairs on the petioles and LaTrojita,19 Feb-10 Mar1944 (9),Cuatrecasas16826
Pourouma 163

FIG.
76 Prom mn b gbith pitt inre
FLORA PANAMENSIS
Br
b*. . .. , .,
A _l I - p _
p ,
f-nTt.
rrrO 10. I wW

- . r - ^ , O^ t a r rol e
. .,, ^ V4
]..p
2m 20 - x? 30 *Iwith
L n_5 5tul
;il1t Sotwo MP bl z
^ ^I
'
I.-it^^^;.;'' *-i FrtouM mal.nrt^.?,Bf??.ft [,..; CC Berg i * ,
*'*^ 'AM9 i^ fe tr ^"b-t* cc*B"^*t'a"*l*)^ - i^. ?: .-
'Fu.: C.C. arit o llreah twig t ;
i.

FIG. 76. Pourouma melinonii subsp. glabrata. Leafy twig with pistillate inflorescences (Berg et al. 355).

(F); Rio Dagua, between Las Cascadas and Alto de (9), Jdtiva et al. 2037 (NY, U, UC, US); Rio Hoja
Yundo, 7 Jul 1984 (9), Gentryet al. 48341 (BG). Blancaand Rio Hualpi, 15 Sep 1965 (d), Little et al.
ECUADOR. ESMERALDAS:
Rio Pambil, Estero Bra- 21070 (F, MO, NY, US); Rio Guayllabamba,Quinin-
vo, 6 Jul 1966 (d),Jdtiva303 (NY), Jdtivaet al. 1079 d6, 4 Oct 1965 (d),Little et al. 21225 (NY, US).
(NY, UC, US); Panadero,5 km from the river, 26 Jul
1967 (d),Jdtivaet al. 2033 (MO, NY, US), 26 Jul 1967 Vernacular names. Ecuador. Esmeraldas: uva.
164 Flora Neotropica

FORESAL
INSTIIWJO AMERICANO
ULATIMO
1I|e1 D . att 1eSs U| I A

h
1g|nc i
:.. .. 2 ..:.
ta -..
. rir, .oula
11, 2 _34 5 3s t
. . . .
.... -,
*.., . ?

;' - ;
UNITED STATES

Pouriou hirauriprt iolata Mtldbread subap.

*!37'0h3 hr*,rup,,-tolat ... . -.

C . .Berg m1a ilo

NToAL Hn
NATiONAL
UI
HERSA1IU
De.
E.C.K. van Keusden
Utrech
IatmD1l 1986 bm

FIG. 77. Pouroumahirsutipetiolatasubsp. hirsutipetiolata.Leafy twig with pistillateinflorescences(Bruijn

1547).

13. PouroumatomentosaMartiusex Miquel in hirsute, often also with white, arachnoid hairs,
Martius,Fl. bras.4(1): 128, t. 38. 1853. Type. pluricellularhairslackingor very sparse.Lamina
Colombia.Amazonasor Caqueta:Araracuara, entire, ovate to elliptic (to oblong), (4-)10-20
Dec 1819 (Q),Martiuss.n. (lectotype,M, cho- (-40) x (3-)5-15(-27) cm or 3-5-lobed to -part-
sen here;isolectotype,U). ed or 5-9-partedand ca. 10-35 x 10-35 cm, co-
riaceous,apexacuminate,roundedor sometimes
emarginate,base acute to obtuse, rounded,trun-
Tree,up to ca. 30 m tall. Leafytwigs 4-15 mm cate, or cordate; upper surface smooth, some-
thick, white-puberulousor partly also yellow- times scabrous,hairy on the main veins or only
Pourouma
m.m,---
* _ . 1
FIG. 78. Pouroumahirsutipetiolatasubsp. hispida. Leafy twig with staminate inflorescences(Little e a.
21225).
21225).
on the midrib, in scabrous leaves hispidulous
over the whole surface, lower surface rather
sparselyappressed-puberulous to strigoseon the
(main) veins, arachnoidhairs in the areoles,usu-
ally extending to the midrib, but on the main
veins ? disappearingwith age; lateral veins 7-
25 pairs, the basal pair branchedor sometimes
(in elliptic leaves) unbranched,the basal part of
165
2'. A . .. ..;
these veins forming the basal part of the leaf
margin(thus not separatedfrom the marginby
mesophyll),tertiary(andquarternary) veins often
ratherprominentbeneath;petiole 2-26 cm long,
white-puberulous or yellow-(sub)hirsute, and
usuallyalso white,arachnoidhairs;stipules(2-)6-
18 cm long, outside whitish-strigoseto yellow-
hirsute or -hirtellous,and with dense to sparse,
166 Flora Neotropica

white, arachnoidhairs, usuallyalso pluricellular
hairs, inside glabrous or with dense or sparse,
yellow(ish) hairs, caducous. Staminate inflores-
cences up to 12 cm long and 12 cm wide; pe-
duncle 2-5 cm long, peduncle and branches +
denselypuberulous,often also with white, arach-
noid hairs;flowers sessile or pedicellate,most of
them in (sub)globoseheads,these 3-5 mm diam.;
tepals 0.5-1 mm long, connate, forminga more
or less distinctly urceolateperianth,sparselyto
densely hairy; filaments longer than the tepals,
free. Pistillate inflorescencesin fruit up to 20 cm
long and 13 cm wide; peduncle 2-13 cm long,
peduncleand branchedsparselyto densely pub-
erulous or also yellow-(sub)hirsute,often with
white, arachnoid hairs; flowers pedicellate (or
subsessile),pedicel up to 0.5 cm long, in fruitup
to 1.5 cm long; perianth5-10 mm long, mostly
densely, yellowish, short-velutinous,sometimes
sparselypuberulous,or also with white, arach-
noid hairs;stigma(sub)peltate,ca. 1.5 mm diam.
Fruitingperianthblack or purple, oblongoid to
ellipsoid to ovoid, 1.5-2 x 0.8-1.5 cm, mostly
densely, yellowish-puberulousto -velutinous or
-subhirtellous.
This species is closely relatedto P. mollis and
P. melinonii. It is distinguishableby the occur-
rence of ? dense, arachnoidhairs on the leafy
twigs, stipules,petioles, main veins at the lower
leaf surfaceand/orthe inflorescences,and, more-
over, by the lack or scarcityof pluricellularhairs
on the leafy twigs.
Five subspecies,partlyallopatric,can be rec-
ognized.A few specimensfromPeru(Amazonas)
cannot be satisfactoralyinsertedin one of these
subspecies.They may representanothersubspe-
cies. These collections (Kayap201, Tunqui217,
Huashikat 966, 1432 and 1482) largely match
the collections under subsp. tomentosa,but the
arachnoidindument is almost lacking on most
parts, except for the main veins of the lamina
beneathand the petioles. Moreover,the hairson
the fruitingperianthare much shorter,and the
apex of the leaves is short-acuminate.They are
provisionallyplaced under subsp. tomentosa.
From the two syntype collections made by
Martius,one is chosenas the lectotypecollection.

Key to the Subspeciesof Pourouma tomentosa

1. Stipuleshairy inside. .................................. ... ................. b. subsp. apiculata.
1. Stipulesglabrousinside.
2. Leafytwigs yellow-hirsute(or glabrous);lamina 5-9-parted. ..................... c. subsp.persecta.
2. Leafytwigs puberulousor hirtellous;lamina entire or 3-5-parted.
3. Base of the lamina more or less deeply cordate;lamina 3-5-parted. ...... d. subsp. essequiboensis.
3. Base of the lamina roundedto truncateor subcordate;lamina usuallyentire.
4. Lamina 3-5-fid to -parted ............................................ a. subsp. tomentosa.
4. Laminaentire.
5. Apex of the lamina rounded to emarginate,(or short-acuminatein aberrantspecimens
fromPeru,Amazonas);headsof the staminateinflorescence4-5 mm diam.;upperAmazon
Basin. .......... ................................. a. subsp. tomentosa.
5. Apex of the lamina acuminate;heads of the staminate inflorescence2-3 mm diam.;
Guianas,Amapa, and near Manaus. ............................. e. subsp. maroniensis.

13a. Pourouma tomentosa Miquel subsp. roundedor emarginate(or sometimes short-acu-

tomentosa. Fig. 79. minate),base roundedto truncate,or (in incised
PouroumaalbistipulataCuatrecasas,Bol. Soc. Venez. leaves)to subcordate;uppersurfacesmooth;stip-
Ci. Nat. 17: 92. 1956. Type. Venezuela.Amazonas: ules outside with dense, white-arachnoidhairs,
Rio Guainia, Victorino, 22 Mar 1942 (9), LI. Wil- inside glabrous.Heads of the staminateinflores-
liams14833(holotype,F; isotypes,F, G, NY, RB, cences ca. 4-6 mm in diam. Pistillate
S, US, VEN). flowers +
diffuselydistributedin the inflorescence,some-
Leafy twigs and petioles white-puberulous. times clusteredin two groups;peduncle mostly
Lamina entire, elliptic to ovate, sometimes (in up to 6(-7) cm long; perianth without dense,
Colombiaand Ecuador)3- or 5-parted,if entire, white, arachnoid indument, in fruit yellowish
then oftenthicklycoriaceous,+ plicate,and apex short-velutinous.
 /+ AJ ++

PO R O ULMA tomentosa.

FIG. 79. Pouroumatomentosasubsp. tomentosa.From Martius,Flora Brasiliensis4(1). 1853: tab. 38. Leafytwig with
section of pistillateflower(17), stigma (13), pericarp(19), seed (21), cotyledons(24).
168 Flora Neotropica

Distribution (Fig. 81). The north-western part See also unnamedcollectionsundernumber2

of the Upper Amazon Basin; in non-inundated (p. 193).
forest, at low altitudes but in Peru (Amazonas)
up to 2000 m. 13b. Pouroumatomentosa Miquel subsp. api-
culata (Benoist) C. C. Berg & van Heusden,
Specimens examined. COLOMBIA. AMAZO-
NAS-CAQUETA:Araracuara, Dec 1819 (9), Martiuss.n. Proc. Kon. Ned. Akad. Wetensch., Ser. C,
(M, U, lectotypecollectionof P. tomentosa),Feb 1820 91(2): 108. 1988.
(st), Martiuss.n. (M, U). VAUPES?:Lagosdel Paso, 25
Dec 1975 (6), Roa 252 (INPA). PouroumaapiculataSpruceex Benoist,Bull.Mus.Hist.
VENEZUELA. AMAZONAS:Conuco, 2 Dec 1977 Nat. (Paris)28: 321. 1922. Type. Brazil.Amazonas:
(st), Liesner4105 (U); CerroDuida, 23 Aug 1944 (st), Rio Uaupes, nr. Panur6(=Ipanor6),Oct 1852-Jan
Steyermark57872 (F); CerroYapacana,3 May 1970 1853 (Q),Spruce2865 (holotype,P; isotypes,B, BR,
(st), Steyermarket al. 103026 (U, Ven);Victorina,Rio C, CGE, E, F, G, GH, GOET,LE, MG, NY, OXF,
Guainia,22 Mar 1942 (9),LI. Williams14833 (G, NY, P, US).
RB, US, VEN, type collection of P. albistipulata). PouroumaapiculataSpruceex Mildbraed,Notzbl. Bot.
ECUADOR.NAPO:8 km SE of Tena, 30 Sep 1960 Gart. Berlin-Dahlem10: 419. 1928. Based on the
(9), Grubbet al. 1681 (NY); ParqueNacional Yasuni, specimenof Spruce2865 in B.
9-19 Jan 1988 (9), Neill et al. 8158 (BG). CoussapoakrukoviiStandley, Publ. Field Mus. Nat.
12-18 km on trail E of La Peca
PERU. AMAZONAS: Hist., Bot. Ser., 17: 163. 1937. Type. Brazil.Ama-
in Serraniade Bagua, 1800-1950 m, 14 Jun 1978 (8), zonas:Mun. Sao Paulo de Olivenca, 11 Sep-26 Oct
Gentryet al. 22865 (BG);Rio Santiago,Caterpiza,17 1936 (a), Krukoff8223 (holotype,NY; isotypes, A,
Oct 1979 (9), Huashikat 966 (BG), 4 Dec 1979 (9), F, G, GH, LE, MO, P, S, U, US).
Huashikat1432 (U), 10 Dec 1979 (9), Huashikat1482
(BG);Rio Cenepa,QuebrackYutai-entsa,22 Jan 1973 Leafytwigsyellow-hirsute(sometimesonly on
(9), Kayap201 (F, GH, NO); Rio Santiago,Quebrada the scars of the stipules), this indument often
Caterpiza,3 Dec 1979 (9), Tunqui217 (BG). LORETO: extended to the petioles. Lamina usually entire
Prov. Requena,ArboretumJenaroHerrera,Aug-Sep and ovate, sometimes 3-lobed or (in
1976 (6),Bernardis.n. (arbre6/27) (U); Prov. Maynas, subjuvenile
Dist. Iquitos,ZungaroCocha, 13 Dec 1967 (9), Reyna material?)3-5-parted, apex usually acuminate,
R. 37 (BG, F). base broadlyacute to truncateor (in subjuvenile
BRAZIL.AMAZONAS: Tefe, 25 Sep 1940 (st), Black material?)occasionallyshallowlycordate;upper
47-1520 (IAN); Manaus-Itacoatiararoad, km 26 (st), surfacesmooth or scabrous;stipulesoutside with
L. Coelho(INPA)5223 (INPA);5 Jun 1976 (st), Har- white arachnoidhairs,inside with
oldo (INPA) 57512 (INPA), 26 Oct 1965 (6), Loureiro (rather)dense,
hairs. Heads of the staminate inflores-
(INPA)16461 (INPA), 19 Mar 1976 (st),Mello(INPA) yellow
55388 (INPA);Manaus,Dec 1850-Mar1851(9), Spruce cences 3-4 mm diam. Perianthof the staminate
951 (M), Spruce 1219 (P), Spruces.n. (B, CGE, E, F, flower densely puberulous.Peduncle of the pis-
G, GOET, LE, NY, OXF, P). tillate inflorescences mostly up to 8 cm long. Pis-
Vernacular names. Colombia. Chiricaba (Kar- tillateflowers usually in two ? distinct clusters;
ijona), guaumoutognac (Tukano); Amazonas- perianth yellow-velutinous,withoutdense,white,
Vaupes: ambauiba do vinho. Venezuela. Ama- arachnoid hairs.
zonas: cocora, cocora montanero, cucura. Peru. Distribution(Fig.81).Venezuela(Bolivar),Peru
Amazonas: paiu shuiya (Huambisa), paui shuina; (Loreto),and Brazil (Amazonas, Mato Grosso,
Loreto: sacha uvilla. Brazil. Amazonas: imbaiba and Roraima);in non-inundatedforest.
(or ambauva) do vinho.
Neill et al. 8158 (from Ecuador) and Rao 252 Specimens examined. VENEZUELA, AMAZONAS:
Depto. Rio Negro,nr.Cerrode LaNeblinaBaseCamp,
(from Colombia) differ from the other collections 22 Feb 1984 (9), Liesner16175 (BG), 4 May 1984 (2),
in the 3- or 5-parted lamina, but match in other Thomas3349 (BG).BOLiVAR:Rio Hacha,Rio Icabaru,
features subsp. tomentosa. Grubbet al. 1681 (from 2 Jan 1956 (8), Bernardi2731 (F, NY), Rio Carfin,Apr
1945 (a), Cardona1222 (US, VEN).
Ecuador) has 5-parted leaves and the postillate PERU. LORETO: Prov. Maynas, Rio Nanay, Mis-
flowers clustered as in subsp. apiculata, but the hana, 6 Jan 1983 (st), Gentryet al. 39264 (BG).
yellow hairs normally occurring on the leaves of BRAZIL.AMAZONAS: Sao Paulo de Olivenca,7 Oct
that subspecies are wanting. Gentry et al. 22865 1931 (8), Ducke s.n. (RB); Humaita, 12 Oct-6 Nov
1934 (9), Krukoff6887 (A, BR, F, G, LE, MO, NY,
(from Peru, Amazonas, 1800-1950 m!) is another
collection which could not be placed satisfacto- RB, S, U, US); Sao Paulo de Olivenca, 11 Sep-26 Oct
1936 (8), Krukoff8223 (A, F, G, GH, LE, MO, NY,
rily. It matches subsp. tomentosa, except for the P, S, U, US, type collection of Coussapoakrukovii);
yellow-hirsute leafy twigs and stipules. Democracia, 31 Aug 1923 (2), Kuhlmann255 (BG);
Pourouma 169

Varadouro do Morcego,31 Aug 1923(8),Kuhlmann PERU. LORETO: Prov. Alto Amazonas,Rio Pastaza,
291(RB);Manaus-Itacoatiarard.,km57, 15Sep1976 Andoar, 21 Nov 1980 (2), Vdsquezet al. 809 (BG).
(a),Mota646(INPA);km 159,20 Sep1974(8),Prance PAsco:Prov.Oxapampa,Distr.Iscozacin,22 Sep 1986
et al. 22708(INPA,MG,U), 291 (RB);Tef6,28 Nov (8), Pariona et al. 954 (BG).
1959 (st),Rodrigueset al. 1434 (INPA);Rio Solim6es, BRAZIL. AMAZONAS:
Sao Paulo de Olivenca, 11
FonteBoa,26 Oct 1968(6),M. Silva2199 (G, MG); Sep-26 Oct 1936 (9), Krukoff8325 (A, BR, F, G, LE,
Ipanor6 ("Panur6"),Oct 1852-Jan 1853 (9), Spruce NY, PS,U).
2865(B,BR,C, CGE,E, F, G, GH,GOET,LE,MG, BOLIVIA. COCHABAMBA: Torora, Jungas of Jana
NY, OXF, P, US, typecollectionofP. apiculata).MATO Maya, 1800 m, 24 Jun 1947 (2), Cardenas3973 (US)
GRosso:Rio Aripuana,NucleoPioneiroHumboldt, LAPAZ:Larecaja,Mapiri,12-30 Sep 1939 (8),Krukoff
Rio Juruenard.,25 Oct 1973(8),Berget al. P.19878 10861 (A, F, G, MO, NY, S, U), 8 Oct-15 Nov 1939
(F, INPA,MO,NY, P, S, U, US);Aripuana,18 Feb- (9), Krukoff11062 (A, F, G, MO, NY, S, U, UC, US,
9 May 1977 (st), Gomeset al. 826, 1517, 1540, 1588, type collection of P. tomentosasubsp.persecta).
1598, 1604, 1610, 1679 (INPA), 8 Nov 1978 (2), Ry-
lands30 (INPA);Mun.VilaBelada Santissima Trin- Vernacular names. Brazil. Amazonas: mapati
idad,5 km S of borderof Rond6nia,2 Nov 1985(2), or mapaty; Mato Grosso: imbafibarana. Bolivia.
Thomaset al. 4765 (BG). RONDONIA: Rd. to Abunato
Guajara-Mirim, 1 km N of Riberao,25 Jul 1968(a), Cochabamba: uva de monte.
Pranceet al. 6292 (G, INPA, NY, S, U, US). RORAIMA: Cardenas 3973 (from Bolivia) differs from the
Anaris, 12 Feb 1969 (2), Prance et al. 9857 (F, GH, others in having the pistillate flowers in two clus-
INPA,M, NY, P, K, S, U, US),Serrados Surucucus, ters in the inflorescences.
15 Feb 1969 (2), Prance9952 (C, INPA, MO, MY, R,
S, U). Vdsquez et al. 809 (from Peru) differs from the
other collections in the absence of long yellow
Vernacularnames. Venezuela. Bolivar: sara- hairs on the leafy twigs. The same applies to
sara(Arekuna).Brazil.Amazonas:imbauibarana, collection Korning et al. 47618 which has, more-
purumai. over, distinctly petiolulate leaf segments.

13c. Pourouma tomentosa Miquel subsp. per-

secta C. C. Berg & van Heusden, Proc. Kon. 13d. Pourouma tomentosa Miquel subsp. esse-
Ned. Akad.Wetensch.,Ser.C, 91(2): 108.1988. quiboensis (Standley) C. C. Berg & van Heus-
Type. Bolivia. La Paz: Prov. Larecaja,Copa- den, Proc. Kon. Ned. Akad. Wetensch., Ser.
cabana,10 km S ofMapiri, 8 Oct-15 Nov 1939 C 91(2): 109. 1988.
(2), Krukoff11062 (holotype, U; isotypes, A, Pourouma essequiboensisStandley, Lloydia 2: 175.
F, G, MO, NY, S, UC, US). Fig. 80. 1939; Berg, Fl. Suriname 5(1): 276. 1975. Type.
Guyana.EssequiboR., nr. mouth of Onoro Creek,
Leafytwigsand petioles yellow-hirsute,some- 15-24 Dec 1937 (d),A. C. Smith 2731 (holotype,F;
times glabrous. Lamina 5-9-parted, with inci- isotypes, A, G, Mo, NY, P, S, U, US).
sions down to (near)the petiole, sometimes with
the segmentspetiolulate,occasionallyentire,apex
Leafy twigs and petioles without long yellow
acuminate, base cordate to subcordate;upper hairs. Lamina 3-5-parted, apex acuminate, base
surface smooth; stipules outside with sparse, ? deeply cordate; upper surface smooth; stipules
white, arachnoidhairs,inside with sparseyellow with rather dense, white, arachnoid hairs, inside
hairs. Heads of the staminate inflorescences ca.
glabrous. Heads of the staminate
5 mm in diam. Peduncle of the pistillate inflo- ca. 3 mm diam. Peduncle of the inflorescences
pistillate inflo-
rescencesoften longerthan 8 cm. Pistillateflow- rescences
mostly up to 6(-7) cm long. Pistillate
ers + diffuselydistributedin the inflorescenceor
flowers ? diffusely distributed in the inflores-
sometimes clusteredin two groups;perianth of cence. Perianth of the
pistillate flower yellow-
the pistillate flower yellow-velutinous, without
velutinous, without dense, white, arachnoid hairs.
dense, white, arachnoidhairs. Distribution (Fig. 81). Guyana and Brazil
Distribution(Fig. 81). Brazil (Amazonas and
(Amazonas and Para); in non-inundated forest.
MatoGrosso),Peru(Loreto),and Bolivia (LaPaz
and Cochabamba);in non-inundatedforest, at Specimensexamined.GUYANA. Mouth of Onoro
altitudesup to 1800 m. Creek, 15-24 Dec 1937 (6), A. C. Smith 2731 (A, F,
G, GH, MO, NY, P, S, U, US, type collection of P.
Specimensexamined.ECUADOR. NAPO:Aiiangu, essequiboensis).
30 May-21 Jun 1982 (st), SEF 8823 (U), 1-15 Feb BRAZIL.AMAZONAS: Itapiranga,Rio Uatuma, 27
1986 (2), Korninget al. 47618 (BG). Aug 1979 (C),Cid et al. 847 (BG, INPA); Rio Jurua,
170 Flora Neotropica

_I

SAt L arsed
forkFloa ed.rrovc e.e

FIG. 80. Pourouma tomentosa subsp. persecta. Leafy twig with pistilate inflorescences (Krukoff 8325).

L U- a SL I-. ITtA

Revised w ora fmbro

picn aT.

Saracura,22 Aug 1975 (9), D. Coelho et al. (INPA)
53383 (INPA, U); Codajas,Rio Capitari,3 Sep 1950
Cuiaba-Santaremrd., km
(d),Fr6es26520 (US).PARA:
1180, 17 Nov 1977(2), A. S. Silva et al. 215 (BG, COL,
K, MO, U).
13e. Pourouma tomentosa Miquel subsp. ma-
roniensis(Benoist)C. C. Berg& van Heusden,
Proc.Kon. Ned. Akad.Wetensch.,Ser.C 91(2):
109. 1988. Fig. 82a.

FIG. 81. Distributionof Pouroumaherrerensis,P. hirsutipetiolatasubsp. hirsutipetiolata,P. hirsutipetiolata

subsp. hispida, P. napoensis,P. tomentosasubsp. apiculata,P. tomentosasubsp. essequiboensis,P. tomentosa
subsp. maroniensis,P. tomentosasubsp.persecta,and P. tomentosasubsp. tomentosa.
Pourouma 171

_ j qlq, PIF P. t osantosastubsp. t lontosa

__ / *~ _~ C~Osa iF~
* Pwith entire lamina

I-~~S Z 1^ ^ ^ 0with incised

ab^P~ 0?^P "L^ ^fr/I ^-^ Smrine

P..hir?utiptiolat. ,3
,wrr@e s

1A . sqa ounu
^Sfti^"1~^PA ?- 'AP. naponsis

i-r g^ -r ----- ---p.-' b-------AS"s"eioovsZ

'l^^ ^P0 ? C sulbOp. mflroni.nsis

.h a,~_ is
,ns
172
FIG. 82. a. Pouroumatomentosasubsp. maroniensis.Leafand pistillateinflorescences(Lanjouwet al. 432).
b. Pouroumaminor.Leafy twig with pistillateinflorescences(Pranceet al. 10089).
Pourouma maroniensis
Benoist.Bull.Mus.Hist.Nat.
(Paris)28:319. 1922;Berg,Fl. Suriname5(1):276,
t. 14a. 1975.Type.FrenchGuiana.Godebert,23
Jun1921(d),Wachenheim ser.2: 392 (holotype,P).
Leafy twigs and stipules without long yellow
hairs. Lamina entire and ovate, only in subju-
venile materialto 3-parted,but then the base to
truncateor at most subcordate,apex acuminate;
uppersurfacesmooth;stipulesoutsidewith dense,
white, arachnoidhairs,inside glabrous.Heads of
the staminate inflorescencesnormally 2-3 mm
diam. Perianthof the staminateflower sparsely
puberulous.Peduncle of the pistillate inflores-
cencesmostly up to 7 cm long. Pistillateflowers
Flora Neotropica
+ diffusely distributed in the inflorescence. Peri-
anth of the pistillate flower sparsely white-pub-
erulous or yellow-velutinous, often with dense,
white, arachnoid hairs.
Distribution (Fig. 81). Eastern Surinam, French
Guiana, and Brazil (Amapa and Para); in non-
inundated forest.
Specimensexamined.SURINAM. Moengo, 25 Aug
1951 (st), BBS 298 (U); Gansee, 21 Apr 1964 (st),
Donselaar1226 (U);GroteZwiebelzwamp,23 Sep 1948
(9), Lanjouwet al. 432 (F, NY, U); Lely Mts., 2 Oct
1975 (st), Lindemanet al. 701 (U).
FRENCH GUIANA. St. Laurent, Feb 1956 (9),
BAFOG7228 (NY, P, U); Mana,3 Mar 1956 (9), BAF-
OG 7330 (NY, P); Charvein,29 Jun 1914 (d), Benoist
Pourouma 173

465(P);TroisSauts,Zidockville, 28Jan1975(2),Gren- type collectionof P. stipulacea);1-6 km W of Ka-

and 688 (CAY);ManaR., SautSabbat,16 Jul 1981 marang,MazaruniR., 21 Aug 1977(st),Maaset al.
(2),Granville 4531 (BG);Godebert,23 Jun 1921(6), 2592 (U).
Wachenheim 392 (P, typecollectionof P. maronien-
sis). This speciesshares(sub)persistentstipuleswith
BRAZIL. AMAPA: Serrado Navio, 4 Jan 1985 (6), P. oraria and P. bicolorsubsp. chocoana. Most
Mori et al. 17672 (BG), 25 Sep 1961 (6), Pires et al. the white, arachnoidindument on
51193(B,FHO,INPA,NY, P, U, US);Rio Araguari, features,e.g.,
9 Oct 1961(6),Pireset al. 51625(FHO,NY, U, US); most parts,suggestrelationshipto P. tomentosa.
Serrado Navio, 1961(st),Rodrigues 2986 and2987 Accordingto Boom (et al. 8102) the exudate
(INPA).PARA: RioJari,MonteDourado,22 Jan1968 is red and viscous.
(2), Oliveira3936 (NY).
Vernacular names. Surinam. Boroma (Ara- 15. PouroumaacuminataMartiusex Miquel in
wak), boesipapja, granboesipapaja. French Martius,Fl. bras 4(1): 130. t. 40. 1853; Mar-
Guiana.Bois canon, bouchi papaye(Paramaka), tius, Syst. mat. med. bras. 34. 1843, nomen.
kukuma,wilaupiyua(Wayapi). Type. Brazil.Amazonas:Rio Japura,between
The morphologicaldifferencesbetweensubsp. Maripiand Parana-Mirim,Dec 1819 (9), Mar-
maroniensis,subsp.apiculata and subsp. tomen- tius s.n. (holotype, M; isotype, U). Fig. 84.
tosa are not strong.
Pourouma populifolia Standley,Publ.FieldMus.Nat.
Hist.,Bot.Ser.,17: 184. 1937.Type.Brazil.Ama-
14. Pouroumastipulacea C. C. Berg, Brittonia zonas:Mun.SaoPaulode Olivenca,nr. Palmares,
1 Sep-26Oct1936(9),Krukoff8427 (holotype,NY;
34(1): 37, t. 2. 1982. Type. Guyana. Bartica- isotypes,A, BR,F, G, LE,P, S, U, US).
Potarord., 25 Jan 1943 (2), Fanshawe 1105 =
FD 3851 (holotype, K). Fig. 83. Tree, up to 26 m tall. Leafy twigs 4-7 mm
thick, (sparsely)yellow-hirsute.Lamina entire,
Tree,up to 6 m tall. Leafytwigs 1-2.5 cm thick, (broadly)ovate to deltoid or to elliptic, 5-19 x
yellow-hirsute.Lamina 5-parted, 25-50 x 35- 3-10 cm, coriaceous,apex(long)acuminate,base
75 cm, coriaceous,apex acuminate,base deeply rounded to acute;
upper surface smooth, main
cordate,often with lobes overlapping;uppersur- veins sparsely hairy, lower surface white-, ap-
face smooth, hirtellouson the main veins, lower pressed-puberulouson the main
veins, minutely
surface white-puberulousand with sparse, or- puberulouson the smaller
veins, arachnoidhairs
ange, pluricellularhairs on the veins, arachnoid in the areoles,sometimes extendingto the main
hairs ? confinedto the areoles;lateralveins 20- veins; lateral veins 7-14
pairs, basal pair un-
30 pairs, tertiary(and quartemary)veins prom- branched,
tertiary venation slightly prominent
inent beneath;petiole 23-65 cm long, sparsely beneath; petiole
sparsely appressed-puberulous
yellow-hirsute;stipules 15-20 cm long, outside to glabrescent,and usuallysparselyhirsute;stip-
yellow-hirsuteand with sparse to dense, white, ules 5-10 cm long, outside yellow-hirsuteand
arachnoidhairs,and with sparse,minute, orange with
dense, white, arachnoidhairs, occasionally
pluricellularhairs,inside sparselyyellow-hirsute, with minute, appressed,white hairs, inside gla-
subpersistent. Staminate inflorescences un- brous, caducous. Staminate inflorescencesun-
known. Pistillate inflorescencesin fruit up to 15 known. Pistillate
inflorescencesin fruit up to 10
cm long and 8 cm wide; peduncles 10 cm long, cm
long and 4 cm wide;peduncle4.5-8 cm long,
peduncle and branches sparsely yellow-hirsute peduncle and branches puberulous(velutinous
and with (dense),white, arachnoidhairs;flowers on the ultimate
branches);flowers4-17; pedicels
ca. 15; pedicel in fruit 0.5-1 cm long; stigma 0.3-0.6 cm
long, in fruitup to 1.5 cm long;peri-
peltate,2 mm diam. Fruitingperianthovoid, ca. anth 2-5 mm long, densely yellow-velutinous,
2 x 1 cm, yellow-hirsuteand with dense, white-
apex papillose;stigmapeltate, 1.5-2 mm diam.,
arachnoidhairs. sometimes puberulous.Fruitingperianthovoid
Distribution(Fig. 85). Guyana;in non-inun- to elliptic, ca. 1-1.6 x 1
cm, densely yellow-
dated forest at low altitudes. velutinous.
Specimens examined. GUYANA.Nr.Kamerang, 20 Distribution(Fig. 85). Brazil(Amazonas)and
Jun1987(st),Boomet al. 8102(BG);Bartica-Potaro Peru(Loreto);in periodicallyinundated(varzea)
rd., 25 Jan 1943 (2), Fanshawe 1105 (=FD 3851) (K, forest or also in non-inundatedforest.
174 Flora Neotropica

- g

FIG. 83. Pourouma stipulacea. Leaf and pistillate inflorescence (Fanshawe 1105); twig with stipules (Maas
et al 2592).

FIG. 84. Pourouma acuminata. From Martius, Flora Brasiliensis 4(1). 1853: tab. 40. Leafy twig with pistillate
inflorescences, fruiting perianth and fruit (17), fruit (19), fruiting perianth (4311), pericarp and seed (19, 21),
seed (21), embryo (23), radicle (25).
Pourouma 175

..4

I0711~~~~~~~~~~~~~~~~~ 41 al

9- . . .. ,~ act ninat

I'OU'R M
AL aeminnata.
 +~~~5 P. acuminata fei\`f `~
.--ID P.
p ~rugines

4~074' -~-3~

P. oraria

-P. <ovata

I P. saulansis

0 P. stipLaacea

?I ?~~~~~~~~~~~~~~~. ils

I? ? 1L ' / ~~~~~~~~~~~~~~~~
Pnv~vthshrther

amrersiP.

FIG.85. istrbutin ofouroma auminta, . boivarnsis P. llipica,P.ferugiea,P. bmivrnorsPioas a

FIG 85 Ditriutin o Poroua aumiata P.bolvarnsi, P eliptca,P. errgina, . mnor P.oraia
ovata, P P. haotica,P
P.P.ovta phaetricha, P. sulnss,P
saulesis, P. siplaea
stipulace, ndP.viloa
aP. ph wtricha
Pourouma
Specimensexamined.VENEZUELA.BOLvAR: Mun.
ForaneoAripao, CarnoFatua, 2-5 May 1988 (st), Ay-
mardet al. 6778 (BG).
PERU. LORETO: Rio Itaya, 5 km above Iquitos, 8
Aug 1972 (9), Croat 18888 (C, DUKE, F, GH, NA,
NY, P); Rio Itaya,Nuevo Esperanza,10 Oct 1975 (9),
McDanielet al. 20362 (NA);Rio Momon, 12 Oct 1977
(2), Rimachi 3256 (MO).
BRAZIL.AMAZONAS: Rio Japura,Vila Bittancourt,
11Nov 1982(9),Amaraletal. 473(BG);Rio ICa,1877-
1878(2),Jobert682 (P);Mun.SaoPaulode Olivena,
nr. Palmares,11 Sep-26Oct 1936(9),Krukoff8427
(A,BR,F,G,LE,MO,NY,P, S,U, US,typecollection
of P. populifolia);
RioJapura,Parana-Mirim, Dec 1819
(2), Martiuss.n. (M, U, typecollectionof P. acumi-
nata);Rio Ia, Oct 1877 (9), (herb.)Schwacke560 (R,
RB).
Vernacularnames. Peru. Loreto:sacha uvilla,
uvilla. Brazil.Amazonas:mapati or mapaty.
This speciesis ratheruniform.It resemblesthe
formofP. herrerensis with oblongto ellipticleaves
and unbranchedbasal lateral veins, but can be
distinguishedfrom the latterby the indumentof
the leafy twigs, consisting of only long hairs or
wanting.
Glaziou 10070 (accordingto the label from
Brazil, Espirito Santo, Itapemerim)is probably
the same collection as Schwacke560.
16. PouroumaferrugineaStandley, Publ. Field
Mus. Nat. Hist., Bot. Sr., 17:181. 1937. Type.
Brazil.Amazonas:Mun. Sao Paulo de Oliven-
ca,Belem Creek,26 Oct- 1 Dec 1936 (6),Kru-
koff 8807 (holotype, NY; isotypes, A, F).
Fig. 86.
Tree, up to 20 m tall. Leafy twigs ca. 10-15
mm thick, white-puberulousand with dense,
purplish, pluricellularhairs; lenticels conspicu-
ous. Lamina entire,ovate and 11-32 x 8-23 cm
or 3-5-fid and up to ca. 45 x 45 cm, coriaceous,
? plicate,apex roundedto acuminate,base trun-
cate to cordate;upper surfacesmooth, hairy on
the main veins, lower surface sparsely subseri-
ceous to strigose on the main veins, the whole
surfacecovered with white or brownish,arach-
noid hairs,which disappearfrom the main veins;
lateralveins 8-16 pairs,the basal pairbranched,
occasionallyunbranched,tertiaryvenationplane
to slightly prominentbeneath;petiole 4-30 cm
long, with dense whitish, arachnoidhairs;stip-
ules 3-13 cm long, outside hispid to substrigose,
inside glabrous, caducous. Staminate inflores-
cences up to 30 cm long and up to 20 cm wide;
peduncle 2-9 cm long, peduncle and branches
177
white-puberulous,densely so on the ultimate
branches;flowers sessile, not densely clustered;
tepals ca. 1.5 mm long, almost free, densely pu-
berulous;filamentsshorterthan the tepals. Sta-
minate inflorescencesunknown. Pistillate inflo-
rescencesup to 70 cm long and 16 cm wide;
peduncle32-48 cm long, peduncleand branches
with indument similar to that of the staminate
inflorescences;flowers7-13; pedicel 0.7-1.3 cm
long, in fruit up to 5 cm; perianthca. 1 cm long,
densely white-puberulous;stigma peltate, ca. 3
mm diam.; bilobate, densely puberulous.Fruit-
ing perianthglobose, ca. 2 cm diam., apiculate,
yellow-short-velutinous.
Distribution(Fig. 85). Brazil (Amazonas),in
non-inundatedforest.
Specimensexamined.BRAZIL.AMAZONAS: Rio So-
lim6es, mouth of Rio Jutai, 5 Sep 1975 (2), L. Coelho
339 (INPA,U); SaoPaulode Olivenca,10 Oct 1931
($), Ducke (RB) 25244 (RB); Sao Paulo de Olivenca,
CreekBelem,26 Oct-11Dec 1936(6),Krukoff8807
(A, F, NY, type collectionofP.ferruginea);Tonantins,
19Feb1944(2),Ducke1527(A,F, IAN,MG,NY, R,
UC, US);Rio Solim6es,Sao Antoniode Ila, 2 Dec
1948 (2), Fr6es23693 (IAN);Manaus-Porto Velho
hwy.,BR-319,km 380, 2 km S of Rio Jutai,13 Oct
1974 (6), Pranceet al. 22859 (INPA, MG, U).
Vernacularnames.Brazil.Amazonas:mapati-
rana.
This species matches P. ovata in the longly
pedunculate,pendulous pistillate inflorescences
withgloboseand apiculatefruitingperianths.The
two species are not closely related.
See unnamedcollection number 3 (p. 193).
17. PouroumavillosaTrecul,Ann. Sci. Nat. Bot.,
Ser. 3, 8: 103. 1847; Miquel in Martius, Fl.
bras. 4(1): 127. 1853. Type. French Guiana.
Without locality, 1840 (6), Leprieurs.n. (ho-
lotype, P). Fig. 87.
Pourouma laevisBenoist,Bull.Mus.Hist.Nat.(Paris)
28: 319. 1922;Berg,Fl. Suriname5(1):270. 1975.
Type.FrenchGuiana.Acarouany,1858(6), Sagot
517(lectotype, P, chosenhere;isolectotypes,
B, BR,
F, G, GH,GOET,P, S, U, US).
Tree, up to 30 m tall. Leafy twigs 5-15 mm
thick, yellow(ish)-puberulousto -subvelutinous
to -subhirsute,or (at least on the scars of the
stipules) pale yellow- to whitish-villous, with
dense, brown, pluricellularhairs. Lamina 3-5-
lobed to -fid, sometimes entire and ovate, rarely
obovate to oblong, 5-40 x 4-42 cm, coriaceous,
rarelychartaceous,apex acuminate,base deeply
178 Flora Neotropica

?~~~~~~~~~~~~~?
•. .. '". '

la"?fr~~~~~~~
~~~L~~r~~AI
~~~~R~~~?? . . ' !':,"..:
Ir?i . :.:.::..
r~~~~~~~~':. . . ...............

' ~ ~ ~ TO
;: -
~ ~
..194~ ~ ~ .
2 Neg|tive
nch
..

?
,~'~ .... ,i
F; ~~~ ~~ ~
rr
4: ~ ~~ ~ ~~ ~~ ~,'~- ,~ ...............~,':..:,,.
,.~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~
? ' ':.-L,.
. . ~. ,> ~ ~

r~ -..-.:. 4~
~ ~~~~ .
~~~~~~~~~~~~~~~~~~?'',,
. ' .
.... _

-?.
._~ ... ':'.-,.C.~ . . . . .

IG86Poruafruine.
Laytwi ihpsilt nirsecDce12)

Jo - .
......":.... .....: .

FIG. 86. Pourouma ferruginea. Leafytwig with pistillate inflorescence(Ducke 1527).

cordateto subcordate,sometimes with overlap- pluricellularhairs on the main veins, and white,
ping lobes; upper surface smooth, hairy on the arachnoidhairsin the areolesand on the smaller
main veins, lower surface white-puberulousto veins; lateral veins 6-20 pairs, the basal pair
yellow-hirsuteon the veins, usuallywith brown, branched(and their basal parts separatedfrom
Pourouma 179

2 3 4 5
11

THNENEW YORK BOTANICAL GARDEN

'
No". '.3 : j ",-'.

kNut.v Wnch 1of2fpmribo.

Old G.D: car.p I!.?"..

Faysetr AIrstri'.)

stilt roots. Fril r'-

d
odor:?'"
'.ciLnnr'.mon-lik

BOTANICAf!' ? O iO

FIG. 87. Pourouma villosa. Leafy twig with pistillate inflorescences (Mori et al. 8583).

the margin by mesophyll), tertiary venation noid hairs;stipules3-25 cm long, outsideyellow-

slightlyprominentbeneath;petiole6-35 cm long, hirsuteto -velutinousor pale yellow- to whitish-
yellow-hirsuteand with long and short, appres- villous,alsowithdense,brown,pluricellularhairs,
sesd, whitehairs,andwith (dense),brown,arach- inside yellow-hirsute to glabrous; caducous.
180 Flora Neotropica

Staminate inflorescencesup to 21 cm long and and Serro do Navio, 10 Oct-15 Dec 1976 (st), Rosa
12 cm wide; peduncle2-10 cm long, yellow-pu- 1126 (MG). AMAZONAS: Manaus,23 Oct 1969 (8), L.
Coelho 15 (INPA, U), 17 Jan 1931 (2), Ducke (RB)
berulous and with dense brown, pluricellular 25243 (RB), 2 Apr 1976 (st), Macedos.n. (INPA), 24
hairs;flowers sessile, mostly clusteredat the ul- Oct 1978 (8), Rodrigueset al. 10082 (INPA), 1850-
timate branches;tepals 1-2 mm long, ovate to 1851 (8), Spruces.n. (G), Mar 1912 (2), Ule 8839 (G,
lanceolate, (almost) free or up to halfway con- MG). MARANHAO: Rod. Belem-Brasilia,km 380-375,
28 Aug 1960 (2), Oliveira 1074 (IAN). PARA:Faro, 3
nate, sparselyhairy; filaments shorter than the Jan 1920 (9), Ducke (MG) 10736 = (RB) 13056 (MG,
tepals. Pistillate inflorescences in fruit up to 19 RB);Rio Branco,Obidos, Serrada Boa Vista, 23 Dec
cm long and 10 cm wide;peduncle2-12 cm long, 1913 (9),Ducke(MG)25225 (MG);upperRio Cupary,
peduncle and branches yellow-puberulous to between Rio Xingu and Rio Tapaj6s, Sep 1931 (2),
-hirtellousand with dense, brown to red-brown Krukoff1197 (A, G, NY, P, S, U); Rio Tapaj6s,Boa
or darkbrown,pluricellularhairs;flowersca. 10- Vista, 1931 (2), Monteiroda Costa 92 (F); Rio Jari,
Monte Dourado, 14 Jan 1968 (a), Oliveira3915 (NY).
40; pedicels up to 0.6 cm long, in fruit up to 1.2 RONDONIA: P6rto Velho-Cuiabahwy., SantaBarbara,
cm; perianth 4-8 mm long, yellow-puberulous 16 Aug 1968 (8), Prance et al. 7160 (F, GH, INPA,
and with dense, dark brown, pluricellularhairs; NY, P, S, U, US).
stigma peltate, 1.5-2.5 mm diam. Fruitingperi-
Vernacularnames. Surinam. Boroma (Aro-
anth purple,(sub)ovoidto ellipsoid, ca. 1.5-2 x
wak), granboesipapaja,kobe (Djoeka),manbos-
0.7-1.5 cm, yellow-puberulousand with sparse
papaja,pourouma or puruma (Carib),yarayara
to dense, dark brown, pluricellularhairs.
(Carib).FrenchGuiana.Bois canon,(bouchi)pa-
Distribution (Fig. 85). Surinam, French pate or papaye (Paramaka).Brazil. Amapa:im-
Guiana, and Brazil (Amapa, Amazonas, Para, bafibabengue;Para:imbafibabranca,imbafiba
and Rond6nia);mostly in non-inundatedforest,de cheiro uvilha.
sometimes in riverine forest, at low alti- The relationshipswith otherPouroumaspecies
tudes. are not clear. The collections from Amazonas
Specimensexamined.SURINAM. Moengo, 16 Aug and Rond6nia, three collections from Para (the
1951 (st), BBS 297 (U); Brownsberg,15 Jul 1916 (st), two Ducke collections and Oliveira 3915) and
BW2095 (U), 13 Sep 1917 (6), BW 3255 (NY, U), 7 the type collection of P. villosa, have long pale
Sep 1917 (6),BW3302 (B, F, MO, P, U), 20 Sep 1918 yellow hairs on the petioles and leafy twigs, but
(), BW 4010 (U, UC), 17 Sep 1921 (9), BW 5390 (F,
RB, U), 28 Jun 1924 (6), BW 6509 (U); Brownsweg, they are similar to the other collections in all
16 Nov 1964 (st), Donselaar 1888 (U); Wilhelmina other features.Togetherwith the overlapin dis-
Mts., 10 Sep 1963 (8),Irwinet al. 55576 (B, FHO, GH, tribution this is a reason not to recognize the
M, MO, NY, U, US), 20 Sep 1963 (6), Irwin et al. form with long hairs as a distinct infraspecific
55884 (FHO, GH, NY, S, U, US); Fallawatra,6 Nov
1971 (2), Jimenez-Saa 1563 = LBB 14296 (NY, U); taxon.
Lely Mts., 29 Sep 1975 (st), Lindemanet al. 540 (U), Cut fruitgive out a cinnamon-likeodor (Mori
5 Oct 1975,Lindemanet al. 790 (U);ParisJacobCreek, et al. 8583).
28 Jun 1965 (st), Maas et al. (LBB) 11017 (U); Kayser The description of P. laevis was based on Ben-
Mts., 29 Oct 1976 (9), Moriet al. 8535 (F, NY, U, US); oist 341, Sagot 517, and Sagot 972. Sagot 517
SectieO, Nov 1942 (9), Stahel (Woodherb.Sur.) 123B
(U), Dec 1942 (9), Stahel (Woodherb.Sur.) 166A (U).
is chosen as the lectotype collection.
FRENCHGUIANA.Withoutlocality(9),Aublets.n.
(G), 1840 (6), Leprieurs.n. (P, type collection of P. 18. PouroumaorariaStandley& Cuatrecasasin
villosa);Charvein-Acarouanyrd., km 1, 27 Oct 1953 Cuatrecasas,Caldasia7: 301. 1956;Woodson,
(9),BAFOG93-M(CAY, P, U); St. Laurent,Dec 1955 Ann. MissouriBot. Gard.47:167.1960. Type.
(9), BAFOG7087 (CAY,NY, P, U); Routede Cayenne,
km 14, 28 Oct 1957 (8), BAFOG7745 (CAY, NY, P); Colombia.Valle:Rio Yurumagui,Veneral,28
Charvein,(9), Benoist 341 (P); Acarouany, 1858 (6), Jan 1944 (2), Cuatrecasas15720 (holotype,F).
Sagot 517 (F, G, GH, GOET, P, S, U, U S, lectotype
collection of P. laevis;in F and P, materialof P. mel- Tree, up to 20 m tall. Leafy twigs up to 15 mm
inonii subsp. melinoniiunderthe same number),1856 thick,yellow-puberulousand with dense, brown,
(9), Sagot 970 (P), (9), Sagot 972 (B, BR, F, G, P, S, pluricellularhairs. Lamina 7-fid, 28-32 x 40
U), (9), Sagot s.n. (U), (st), Sagot s.n. (G).
BRAZIL. AMAPA: 35 km ESE of Oiapoque,2 Dec cm, coriaceous,apexacuminate,basedeeplycor-
1984 (9), Daly et al. 3790 (BG);Rio Amapari,Rancho date with overlapping lobes; upper surface
SantaAna, 30 Sep 1961 (6), Pires et al. 51390 (FHO, smooth, main veins hairy, lower surfacepuber-
IAN, INPA, MG, MO,NY, US);betweenP6rtoPlaton ulous on the veins, arachnoidhairs confined to
Pourouma 181

the areoles;lateralveins up to ca. 30 pairs, ter- pressed-puberulouson the main veins, arach-
tiary and quarternaryvenation prominent be- noid hairsin the areolesand on the smallerveins;
neath; petiole 30 cm long, yellow-hirsuteat the lateralveins 7-12 pairs, basal pair unbranched,
adaxial side, for the rest sparselyhairy;stipules tertiaryvenation plane beneath;petiole 2-9 cm
10-12 cm long, outside yellow-hirsute,and with long, with some (very) sparse,white hairs;stip-
white appressedhairs, occasionallywith brown, ules 1-5 cm long, outside yellow-appressed-pu-
pluricellularhairs,inside yellow-hirsute,subper- berulous,inside glabrous,caducous. Staminate
sistent.Staminateinflorescences about25 cm long inflorescencesunknown.Pistillate inflorescences
and 15 cm wide;peduncleca. 12 cm long,yellow- in fruitup to 8 cm long and 5 cm wide;peduncle
puberulous;flowers in clustersof up to 10 flow- 2-3.5 cm long, peduncleand branchesminutely
ers; tepals free, ca. 2 mm long; filamentsshorter yellow-puberulous,the ultimate branchesmore
than the tepals. Pistillate inflorescencesin fruit denselyhairy,and with brown,pluricellularhairs;
up to 22 cm long and 18 cm wide; peduncle 14- flowers7-10; pedicel0.1-0.5 cm long, in fruitup
15 cm long, peduncle and branchesyellow-pu- to 1 cm; perianth ca. 1.5-2 mm long, yellow-
berulous and with dense, dark-brownor red- puberulous;stigma bilobed, ca. 2.5 mm diam.
brown or brown, pluricellularhairs;flowers ca. Fruitingperianthovoid, ca. 2 x 0.8 cm, sparsely,
100; pedicels ca. 5 mm long, in fruit up to 10 minutely appressed-puberulous.
mm long; perianth3-5 mm long, yellow-puber- Distribution(Fig. 85). FrenchGuianaand Bra-
ulous and with dense, dark-brown, pluricel- zil (Amapa);in non-inundatedforest.
lular hairs; stigma peltate, 1.5-2 mm diam.
Specimens examined.
FRENCHGUIANA.Saiil,20
Fruitingperianthovoid to ellipsoid, ca. 15 x 10 Dec 1976(Q),Granville
2744 (CAY),Saul,km 2.7 on
mm, + sparselyhairy. trailon MontagneBelvedereSud, 21 Oct 1971 (v),
Distribution(Fig. 85). Colombia (Choc6 and OldemanB.4132(CAY,P, U, VENtypecollectionof
Valle);in lowland forest. P. saulensis).
BRAZIL.AMAPA:
109 km SSE of Oiapoque,5 Dec
Specimens examined. COLOMBIA. CHOC6:Be- 1984(v),Moriet al. 17185(BG).
tweenCerteguiandLasAnimas,17Aug1976(9),Gen-
tryet al. 17804(BG,U);N of Certegui,13 Jun 1982 Vernacularname.FrenchGuiana.Bois canon.
(8), Gentryet al. 36797 (JUAM). VALLE:Costa del This species shows similaritiesto P. ovata in
Pacifico,Rio Yurumangui, Veneral,28 Jan 1944(9), the indument, the short stipules, and the un-
Cuatrecasas 15720(F, typecollectionof P. oraria). branched,
basal, lateral veins. The species are
This species resemblesP. bicolorsubsp. cho- distinct in the numberof lateralveins, the length
coana in many features (see p. 144). It differs of the peduncleof the pistillateinflorescence,and
from the latter in the smooth upper surface of the shape of the fruitingperianth.
the lamina,the smooth fruitingperianth,and the
concentrationof the long hairsat the adaxialside 20. Pouroumaovata
of the petiole. P. oraria may be much closer to Trecul,Ann. Sci. Nat. Bot.,
Ser. 3, 8: 101. 1847, Miquel in Martius, Fl.
P. bicolor (subsp. chocoana) than suggestedby bras.4(1): 132.1853. Type.Brazil.Para:With-
the arrangementoftaxa in the presenttreatment. out locality,without date (d),Ferreiras.n. (ho-
lotype, P). Fig. 89.
19. Pouroumasaulensis C. C. Berg, Brittonia
34(1): 36, t. 1. 1982. Type. French Guiana. Pouroumalongipendula Ducke,Notizbl.Bot. Gart.
km 2.7 on trail of Berlin-Dahlem 11: 581. 1932.Type.Brazil.Ama-
Sail, MontagneBelvedere zonas:Manaus,fallsof Rio Taruma;17 Oct 1929
Sud, 21 Oct 1971 (9), Oldeman B.4132 (ho- (v),Ducke(RB)23608(holotype,RB,not seen;iso-
lotype, U; isotypes, CAY, P, VEN). Fig. 88. types,B, G, S, US).
Tree, up to 11 m tall. Leafy twigs 3-5 mm Tree, up to 25 m tall. Leafy twigs 4-9 mm
thick, minutelypuberulousto glabrous.Lamina thick,sparselywhite-appressed-puberulous, more
entire, (broadly)ovate to elliptic, 6-14 x 3-9.5 densely haired at the nodes. Lamina entire, el-
cm, coriaceous, apex (sub)acuminate, base lipticto ovate, 7-26 x 3-13 cm, coriaceous,apex
roundedto obtuse;upper surfacesmooth, some shortacuminate,base acuteto obtuseto rounded
sparse,white hairsand brown,pluricellularhairs or sometimes truncate;upper surface smooth,
on the main veins, lower surface white-, ap- sparselyhairyand occasionallywith some brown,
182 Flora Neotropica

[--

-
'
.' ? "

i. "

Q~~~~~~~~~~~," ..

FIG. 88. Pourouma saulensis. Leafy twig, (schematic) pistillate inflorescence, and fruiting perianth (Granville~~~~~~
2744); pstillat flower OldemanB.413')

FIG.88. Pouroumasaulensis.Leafytwig,(schematic)pistillateinflorescence,and fruitingperianth(Granville

2744); pistillateflower(OldemanB.4132).

pluricellularhairs on the midrib, lower surface glabrescent, ultimate branches usually more
white-puberulouson the veins, arachnoidhairs densely yellow- to orange-puberulous;flowers
confinedto the areoles;lateralveins 9-21 pairs, sessile or occasionally pedicellate, mostly in
basal pair unbranchedor occasionallybranched, subglobose heads, these 3-7 mm diam.; tepals
tertiary venation slightly prominent beneath; ca. 1 mm long, connate, rather densely hairy;
petiole 2-7.5 cm long, subglabrousto sparsely filamentsshorterthan the tepals. Pistillate inflo-
white hairy;stipules 1.5-3 cm long, outside yel- rescencesin fruit up to 38 cm long and 11 cm
low-velutinous,usuallywith brown,pluricellular wide; peduncle up to 32 cm long, peduncle and
hairs, inside yellow-(or orange-brown- or branchespuberulousto almost glabous;flowers
white-)sericeous,caducous. Staminate inflores- 7-20; pedicel 0.2-2.4 cm long, in fruit 1-3 cm,
cences up to 10 cm long and 9.5 cm wide; pe- obconical,conspicuouslylenticellate;perianth3-
duncle 1-5.5 cm long, sparsely puberulous to 5 mm long, denselyshort-velutinous;stigmapel-
Pourouma 183

| I 3 -2^
2 A?:.

Ak~~~~~~~~~~~~?IIk

TrMw Tons BOTANICAL

GAIM'W
n
liBlTIm lA L AiASSPA MZI.
i. ii~L m.---.-V---lmDI,

_ r kd
L oree
,.l._ l,-~v ?
of tioe rmchliho.
Frest om terra firme on cly.
Tm 1in x 20cm; frvit green, maturing red,
w clew
. light bow. ri. oraa
wdll g- -
' '*--- J 'l .* ovd'sember
1-8....

FIG. 89. Pouroumaovata. Leafy twig with pistillateinflorescences(Stewardet al. 107).

184
tate, bilobed, ca. 1.5-2 mm diam. Fruiting peri-
anth red (mature?), (depressed-)globose, ca. 1.5
cm diam., puberulous.
Distribution (Fig. 85). Colombia (Vaup6s),
Venezuela (Bolivar), Peru (Loreto), and Brazil
(Acre, Amazonas, and western Para); in non-
inundated forest.
Specimensexamined.COLOMBIA.VAUPES:Mon-
fort, 30 Nov 1952 (9), Romero-Castaneda3848 (F,
MBG,NY); Rio Apaporis,Soratama,26 Mar 1952 (2),
Schulteset al. 16009 (C, U, US).
VENEZUELA.BOLIVAR: SierraIchfin,Salto Maria
Espuma,28 Dec 1961 (6), Steyermark90368 (NY, U,
VEN).
PERU. LORETO: Prov. Requena, Rio Ucayali, Ar-
boretumJenaroHerrera,Aug-Sep 1976 (st), Bernardi
5.16.5 (U), Aug-Sep 1976 (9), Bernardis.n. (tree 4/8)
(U), 3 Jan 1974 (9),Diaz 22-A (G);Prov. Maynas,Rio
Nanay,Mishana,23 Mar 1979 (st), Gentryet al. 26049
(U), 24 Mar 1979 (st), Gentryet al. 26177 (U), 12 Jan
1983 (st), Gentryet al. 39379 (BG); Prov. Requena,
ArboretumJenaroHerrera,9 Dec 1980 (9), Vdsquez
et al. 1004 (BG).
BRAZIL.ACRE:Cruzeirodo Sul, 11 Feb 1976 (9),
Monteiroet al. 309 (INPA), 1 Mar 1976 (2), Ramos et
al. 204 (INPA).AMAZONAS: Manaus,ReservaFlorestal
Ducke, 27 Nov 1957 (st), D. Coelho (INPA) 5979
(INPA), 13 Sep 1957 (6), L. Coelho(INPA) 5808 (U);
Manaus,waterfallsof Rio Taruma, 17 Oct 1929 (9),
Ducke (RB) 23608 (B, G, RB, S, US, type collection
of P. longipendula);Manaus,ReservaFlorestalDucke,
17 Oct 1929 (9), Killip et al. 30139 (A, F, NY, US);
Humaita,7-18 Nov 1934 (2), Krukoff7073 (A, B, BR,
F, G, GB, LE, MO, NY, RB, S, U, US); Sao Paulo de
Olivenca, 26 Nov 1936 (9), Krukoff8695 (A, B, BR,
F, G, MO, NY, P, S, U, US); Manaus-P6rtoVelho rd.,
km 240, 21 Nov 1973 (9), Lleras et al. P.19586 (F,
INPA, MO, NY, P, S, U, US); Manaus,ReservaFlo-
restalDucke, 10 Nov 1965 (9),Loureiro(INPA)16968
(U); Torquato-Tapajosrd., km 118, 28 Aug 1975 (9),
Loureiroet al. (INPA) 50623 (INPA); Manaus-Cara-
caraird., km 57, 16 Sep 1976 (9), Mota 680 (INPA);
Tapuruquara,4 Jul 1972 (9), Prance et al. 15393 (F,
GH, INPA, M, NY, P, S, U); km 245, 13 Mar 1974
(9), Prance et al. 20454 (INPA, MO, NY, S, U, US),
km 380, 13 Oct 1974 (9), Prance et al. 22864 (INPA,
MO, NY, P, S, U, US); Manaus, Reserva Florestal
Ducke, 21 Aug 1965 (9), Rodrigues 5449 (NY, U);
Manaus-Itacoatiarard., 3 Sep 1965 (9), Rodrigueset
al. 7081 (U), 26 Nov 1965 (9), Rodrigueset al. 7292
(U); Manaus-Caracaraird., km 60, 20 Aug 1976 (6),
Shima et al. 14 (INPA); between Rio Castanhoand
Rio Tupanan, 14 Jul 1972 (8), M. F. Silva et al. 742
(U); 18 Jul 1972 (9), M. F. Silva et al. 869 (INPA);
Manaus-P6rtoVelho rd.,km 250, 5 Jan 1974 (9),Stew-
ard et al. P.20156 (INPA, U). PARA:Withoutlocality,
(6),Ferreiras.n. (P, typecollectionof P. ovata);Cuiaba-
Santaremrd., km 1230, (9), Prance et al. 25530 (F,
MO, RB, S, U); Rio Trombetas(6), N. T. Silva et al.
4740 (MG, U). RORAIMA: Manaus-Caracaraird., km
343, 18 Nov 1977 (9), Stewardet al. 107 (U).
Flora Neotropica
Vernacularnames.Peru.Loreto:chullachaqui,
chullachaquiblanco, chullachaquicaspi, sacha
uvillos. Brazil.Amazonas:imbafbarana.
This speciesresemblesP.ferrugineain the long-
pedunculate and pendulous pistillate inflores-
cences and the globose and apiculate fruiting
perianth,but is in other featuresquite distinct
from the latter. In the vegetative parts it shows
resemblancesto P. saulensisin severalcharacters
(seep. 181).
21. Pourouma elliptica Standley, Publ. Field
Mus. Nat. Hist, Bot. Ser.,17:181. 1937. Type.
Brazil.Amazonas:Mun. Sao Paulode Oliven-
9a, nr. Palmares,11 Sep-26 Oct 1936 (6),Kru-
koff8388 (holotype, NY; isotypes, A, BR, F,
G, LE, MO, P, S, U). Fig. 90.
Tree, up to ca. 15 m tall. Leafy twigs ca. 10
mm thick, densely yellow-hirsute and yellow-
subhispidulous,thus a mixtureof hairsdistinctly
differentin length, moreover, with sparse red-
dish-brown,pluricellularhairsand sparse,white,
arachnoidhairs. Lamina entire, elliptic to ob-
long, 13-30 x 6.5-20 cm, coriaceous, apex
abruptly,short-acuminate,base acute to round-
ed to truncate;upper surface smooth, hairy on
the main veins, to puberulous on the smaller
veins, lower surface yellow-(sub)hirsuteon the
main veins, to puberulouson the smallerveins,
white, arachnoidhairs in the areoles and on the
smallerveins, lateralveins 10-16 pairs,the basal
pair unbranched or sparsely and/or faintly
branched, tertiary venation almost plane be-
neath;petiole 5-13.5 cm long,yellow-hirsuteand
with sparseto dense,white, arachnoidhairs;stip-
ules 3-9 cm long, caducous, outside yellow-hir-
sute, inside glabrous. Staminate inflorescences
up to 19 cm long and up to 14 cm wide;peduncle
4-7 cm long, yellow-hirsuteon the peduncle to
hispidulouson the ultimatebranches,sometimes
also white, arachnoidhairs;flowers sessile, not
denselyclustered;tepalsca. 1.5 mm long, almost
free, densely hispidulous;filamentsshorterthan
the tepals. Pistillate inflorescencesunknown.
Distribution(Fig. 85). Brazil (Amazonas);in
non-inundatedforest.
Specimensexamined. BRAZIL. AMAZONAS: Espe-
ranga,mouth of Rio Janvari,21 Sep 1931 (6), Ducke
(RB)25246 (RB);Sao Paulode Olivenca,nr. Palmares,
11 Nov-26 Oct 1936 (6), Krukoff8388 (A, BR, F, G,
LE, MO, NY, P, S, U, type collection of P. elliptica).
Pourouma 185

I I

,E Fn CL1 STPE
|SOP Y r.

Tr ern rt. 'mh; trunk 41 G

Safeo AmSaonm: r u.odeO .

near Plmamres. Sept. Il-OCt. 26.1M91
.di-tb.td thlu
.: th.New YO3k hlod= 6 tc .'bre 4937

FIG. 90. Pouroumaelliptica.Leafy twig with staminateinflorescence(Krukoff8388).

186 FloraNeotropica

Vernacularnames. Brazil. Amazonas:mapati 5(1): 278, t. 14b. 1975; Burger, Fieldiana Bot.
or mapaty. 40: 202, t. 22. 1977. Type. French Guiana.
Saint Jean de Maroni, 17 Mar 1914 (9), Benoist
22. PouroumabolivarensisC. C. Berg,Brittonia 960 (lectotype, P, chosen here, cf. Berg, Fl.
34(1): 39, t. 3. 1982. Type. Venezuela.Bolivar: Suriname 5(1): 278. 1975). Fig. 82b.
CerroVenamo, SW part, upper Rio Venamo,
PouroumaaureaUle ex Mildbraed,Notizbl. Bot. Gart.
10 Jan 1964 (9), Steyermarket al. 92936 (ho- Berlin-Dahlem10: 418. 1928;Ule, Bot Jahrb.Syst.
lotype, VEN; isotypes, NY, U). Fig. 91.
40:149. 1907, nomen.Type.Brazil.Acre:Rio Jurua-
Mirim, Aug 1901 (s), Ule 5718 (holotype, B; iso-
Tree, up to 15 m tall. Leafy twigs 5-8 mm types, G, MG).
thick, puberulous.Lamina entire, broadlyellip-Pouroumafolleata Macbride,Publ. Field Mus. Nat.
tic to obovate, sometimes broadlyovate or sub- Hist., Bot. Ser., 8(2): 114. 1930, l.c. 13(2.2): 292.
orbicular, 5-19 x 3-14 cm, coriaceous, apex 1937.Type.Peru.Junin;Chanchamayovalley, 1924-
1927 (8), C. Schunke416 (holotype,F; isotype, B).
shortlyacuminateto roundedto emarginate,base PouroumaisophlebiaStandley,Publ. Field Mus. Nat.
acute to obtuse; upper surfacesmooth, sparsely Hist., Bot. Ser., 17: 182. 1937. Type. Brazil.Ama-
white-hirtellousor glabrouson the midrib,lower zonas: Mun. Sao Paulo de Oliven9a,nr. Palmares,
surfaceappressedwhitish-pubescenton the main 11 Sep-26 Oct 1936(9),Krukoff8037(holotype,NY;
veins, arachnoidhairs in the areolesand ? cov- isotypes,A, BR, G, LE, P, S, U, US).
PouroumasubplicataStandley,Publ. Field Mus. Nat.
eringthe smallerveins; lateralveins 6-10 pairs, Hist., Bot. Ser., 17: 184. 1937. Type. Brazil. Acre:
basal pair and sometimes the second pair Nr. mouthof Rio Macauhan,4 Aug 1933(9), Krukoff
branched, tertiary venation plane or slightly 5282 (holotype,F; isotypes,A, G, LE, M, MO, NY,
prominent beneath;petiole 1-6.5 cm long, pu- S, U, UC, US).
PouroumaumbellataStandley,Publ. Field Mus. Nat.
berulous;stipules 2-6 cm long, outside yellow- Hist., Bot. Ser., 17: 185. 1937. Type. Brazil.Ama-
puberulous, inside glabrous, caducous. Stami- zonas:Mun.Humaita,betweenRio Livramentoand
nate inflorescencesunknown. Pistillate inflores-Rio Ipixuna,7-18 Nov 1934 (9), Krukoff7071 (ho-
cencesin fruitup to 10.5 cm long and 2 cm wide; lotype, F; isotypes, A, BR, G, GB, LE, MO, NY,
peduncle 3-7 cm long, peduncle and branches RB, S, U, US).
CoussapoaemarginataKillip ex Macbride,Publ.Field
yellow-puberulous;flowers 7-12; pedicel 0.2-1 Mus. Nat. Hist., Bot. Ser., 13(2.2):296. 1937. Type.
cm long, in fruit up to 2 cm; perianthca. 3 mm Peru.Loreto:Mishuyacu,nr.Iquitos,24-28 Sep 1929
long, puberulous;stigma peltate, more or less (9), Killip & A. C. Smith 29955 (holotype, US; iso-
bilobed, 2-4 mm diam. Fruiting perianth ma- types, B, F).
PouroumacuatrecacasiiStandleyin Cuatrecasas,Re-
roon-red,ca. 1.8 x 1 cm, puberulous. vista Acad. Colomb.Ci. Exact.9(36/37): 339. 1956.
Distribution(Fig. 85). Venezuela(Bolivar);in Type. Colombia.Vaupes:Mandi, nr. Mitfi, 24 Oct
humid, (sub)-montane forest, at altitudes be- 1939 (9), Cuatrecasas7299 (holotype,US).
tween 900 and 1350 m. Pouroumaumbellifera Burger,Phytologia26:430. 1973.
Type. Costa Rica. Heredia:Nr. Rio Sarapiqui,Ti-
Specimensexamined.VENEZUELA.BOLIVAR: Cer- rimbina, 21 Jan 1972 (9), Lent 2327 (holotype, F;
ro Venamo,Rio Venamo,8 Jan 1964(9),Steyermark isotypes,B, US).
et al. 92850(U, VEN);CerroVenamo,SWpart,upper
Rio Venamo,alt.950-1000m, 10 Jan1964(9),Stey- Tree, up to ca. 35 m tall. Leafy twigs 2-13 mm
ermarket al. 92936(NY,U, VEN,typecollectionof thick, yellow-sericeous to -hirsute, or appressed-
P.bolivarensis);
RioCuyuni,RioAnawaray-parui,1300- puberulous, almost glabrous, some-
1350m, 25 Dec 1970(st),Steyermark et al. 104466 times with occasionally
(NY,VEN). sparse brown, pluricellular hairs.
Lamina entire, narrowly to broadly obovate to
Vernacularnames. Venezuela. Bolivar: cay- (broadly) elliptic, rarely to oblong, 4-45 x 1-15
wari-cay-yek(Arekuna). cm, coriaceous to chartaceous, apex (long) acu-
This species resemblesP. minor, from which minate to mucronate or rounded to emarginate,
it differsin the sparserindument,branchedpis- base acute to rounded, rarely to retuse; upper
tillate inflorescences,and branchedbasal lateral surface smooth, yellow-sericeous, hirtellous, hir-
veins. sute, or puberulous on the midrib, occasionally
brown, pluricellular hairs on the main veins, low-
23. Pouroumaminor Benoist, Bull. Mus. Nat. er surface yellow-sericeous to -velutinous or to
Hist. (Paris)30: 103. 1924;Berg,Fl. Suriname -hirsute on the main veins (minutely) puberulous
Pourouma 187

. . .

FIG. 91. Pouroumabolivarensis.Leafy twig with pistillate inflorescences(Steyermarket al. 92936); sterile
leafy twig (Steyermarket al. 104466).

to hirtellous on the smaller veins, orange, pluri- branched, tertiary venation (almost) plane be-
cellular hairs present or not, arachnoid hairs in neath;petiole 0.7-15 cm long, yellow-velutinous
the areoles, usually ? extending to the smaller to -sericeous or hirsute;stipules 1-15 cm long,
veins; lateral veins 6-26 pairs, basal pair un- outside yellow-sericeousto -velutinous, some-
 188 Flora Neotropica

times -hirtellous, sometimes with orange-brown, (F, VEN), (2), Steyermark60414 (F, VEN); Rio Chi-
pluricellular hairs, inside glabrous, caducous. canan, PuertaLema, SE of El Dorado, 24 Aug 1961
Staminate inflorescences up to 12 cm long and (2), Steyermark89506 (NY, U, VEN); Venezuelan-
Brazilianfrontier,SerraniaPia-soi, 5-6 Jan 1962 (st),
7.5 cm wide; peduncle 1.5-8.5 cm long, yellow-
Steyermark90620 (VEN).
sericeous to -velutinous; flowers sessile or pedi- GUYANA.KanukuMts., Iramaipang,Nov 1948(6),
cellate, diffusely distributed along the ultimate Wilson-Browne549 (=FD 5948) (NY).
branches or occasionally + clustered; tepals 1.5- SURINAM. Kabalebo, 12 Nov 1976 (st), Heyde et
al. 32 (U), 13 Nov 1976 (9),Heyde et al. 39 (U); River
3 mm long, connate, puberulous to almost gla-
Nassau, Mts, 16 Feb 1949 (2), Lanjouwet al. 2214
brous; filaments shorter than the tepals. Pistillate (NY, U); ParisJacobCreek,29 Jun 1965 (st), Maas et
inflorescences in fruit up to 12 cm long and 7 cm al. (LBB) 11027 (U); TapanahonyR., at mouth of
wide, subumbellate; peduncle 1-9 cm long, yel- PaloemeuR., Aug 1959 (st), Schulz8161 (U); Goddo,
low-sericeous; flowers 2-11; pedicel 0.2-1.2 cm 26 Jan 1926 (9), Stahel (Exp. Wilhelminageb.)82 (F,
U); Voltzberg,Troon(LBB) 16310 (U).
long, in fruit up to 3.5 cm; perianth 2-6 mm FRENCH GUIANA. St. Laurent,13 Jan 1950 (9),
long, usually densely hairy; stigma knob-shaped, BAFOG5042 (P), 2 Jan 1956 (2), BAFOG7152 (CAY,
variously lobed, 2.5-6 mm diam. Fruiting peri- NY, P, U), 16 Jan 1956 (2), BAFOG7173 (CAY, NY,
anth ovoid with an apiculate apex, ca. 1-2.5 x P, U), 8 Feb 1956 (2), BAFOG7261 (NY, P), 10 Feb
1-1.5 cm, reddish, hairy. 1956 (2), BAFOG7268 (CAY, NY, P, U); Saint Jean
de Maroni, 17 Mar 1914 (2), Benoist 960 (P, lectotype
Distribution (Fig. 85). From Nicaragua to the collection of P. minor); 16 Mar 1914 (2), Benoist 978
Amazon Basin and the Guiana regions; in non- (P); Aug 1973 (st), Garnier112 (CAY);betweenSaut
inundated forest at altitudes up to ca. 1500 m. Emerillonand Etats-Unis,21 Aug 1970 (st), Granville
581 (CAY,P, U); Trois Sauts,3 Nov 1974 (9),Lescure
Specimensexamined.NICARAGUA.Rio SanJuan, 422 (CAY); upper ApprouageR., Creek Par6pou,7
nr. CainoChotaleno, 20 km NW of El Costillo, 7-9 Feb 1967 (st), Oldeman2504 (CAY, NY, P, U); Ap-
Mar 1978 (st), Neill 3389 (BG). prouageR., Pierrette,14 Sep 1968 (st), Oldeman2821
COSTA RICA. ALAJUELA:15 Sep 1973 (6), Hart- (CAY, P); Godebert,without date (st), Wachenheim
shorn 1295 (F); Corazonde Jesus, 29 Mar (6), Hart- 18 (P), 23 Jun 1921 (6), Wachenheim467 (P), 13 Jul
shorn 1431 (F, U). HEREDIA: Tirimbina, 12-15 Aug 1921 (st), Wachenheims.n. (A, E, U, US).
1971 (st), Burgeret al. 8137 (F); Las Horquetas,25 ECUADOR. NAPO:Rd. Coca-Lago Agrio, 9 km NE
May 1973 (6), Hartshorn1224 (F, MO, U); nr. Rio of Rio Coca, 20 Mar 1980 (st),Brandbygeet al. 30252
Sarapiquiat Tirimbina,21 Jan 1972 (Q),Lent 2327 (F, (AAU);Guayusa,2 hr upstreamRio Coca from Coca,
GH, MO, NY, U, US, type collection of P. umbelli- 24 Mar 1980 (st), Brandbygeet al. 30327 (AAU);Afn-
fera); La Virgen,23 Jul 1979 (6), Stevens 13340 (U). angu,30 May-21 Jun 1982 (st), SEF8617, 9033, 9159
LIMON: BetweenSiquirresand Turrialba,10 Oct 1972 (U); MORONA-SANTIAGO: nr. Bomboiza, 27 Sep 1985
(st), Holdridge6818 (F, MO, U); Guapiles, 12-13 Mar (st), ShakaimRBAE-26 (BG).
1924 (st),Standley37121 (US);Cairo, 18-19 Feb 1926 PERU. AMAZONAS: Rio Santiago, nr. Caterpiza, 2
(juv), Standleyet al. 48600 (US). Nov 1979 (6),Huashikat1150 (U); mouth of Rio San-
PANAMA. COCLE: El Cope, 1 Sep 1977 (6), Berg tiago, 8-13 Oct 1962 (6), Wurdack2169 (F, G, GH,
400 (U); 25 Nov 1977 (st),Folsomet al. 6469 (U). SAN NY, P, S, UC). HUANUCO: Rio Pachitea,Tournavista,
BLAS:CerroSanJose,YarBirea,5 Feb 1986 (st),Nevers 8 Aug 1967 (6), J. Schunke V. 2139
(F, G, GH, NY,
et al. 6997 (MO). US); Prov. LeonciaPrado,Tingo Maria, 17 Feb 1964
COLOMBIA. AMAZONAS-VAUPES: Rio Apaporis, (st), Vasquez 17 (F). JUNiN: San Ram6n, 1925
Aug (6),
betweenRio Kananariand Rio Pacoa, 1-15 Dec 1951 C. SchunkeA-95 (F, NY, US);
ChanchamayoValley,
(2), Garcia-Barriga13836 (NY), 28 Sep 1951, Schultes 1924-1927 (8), C. Schunke416 (B, F, type collection
et al. 14148 (U). ANTIOQUIA: Between Dos Bocas and ofP.folleata). LORETO:
Puerto Almendras, 7 Aug 1973
Anori, 24-31 May 1973 (6), Soejartoet al. 4063 (A, (st),Ayala 341 (U); Rio Nanay, halfwaybetweenIqui-
MO). CHOCO: Novita, CerroTorra, 22 Feb 1977 (2), tos and SantaMariade Nanay, 31 May 1975 (st), Gen-
Forero et al. 3145 (RB, U). META:Sierrade la Ma- try et al. 22381 (U), 23 Mar 1979 (st), Gentryet al.
carena, 19 Jan 1950 (6), Philipson et al. 2134 (US). 26049 (U), 24 Mar 1979 (st), Gentryet al. 26177 (U);
VALLE:Rio Anchicaya,Alto Yunda,1000 m, May 1972 Prov. Maynas,Yanamono ExploramaTourist Camp,
(Q),Hilty M-86 (MO). VAUPES: Mandi, nr. Mitfi, 24 1 Jul 1983 (st), Gentryet al. 42568 (BG);Mishuyacu,
Oct 1939 (2), Cuatrecasas7299 (US, type collectionof nr. Iquitos, 24-28 Sep 1929 (2), Killip& Smith 29955
P. cuatrecacasii);Rio Inirida,SanJoaquin,27 Jan 1955 (US, B, F, type collection of Coussapoaemarginata);
(6), Fernandez2019 (US). Yurimaguas-Tarapotord., km 14, 8 Nov 1967 (2),
VENEZUELA. AMAZONAS:El Dorado-Sta. Elena Soria S. 21 (F, G, US). MADREDE DIos: Rio Manu,
rd., S of El Dorado, km 79, 25 Feb 1959 (2), Bernardi CochaCashustation,2 Sep 1976 (9),Fosteret al. 5019
7250 (F, G, NY, VEN);SierraParima,Simarawochi, (F), 23 Oct 1979 (6), Foster et al. 7199
18 Apr-23 May 1973 (Q),Steyermark107064 (F, U). bopata, mouth of Rio (F); Rio Tam-
D'Orbigny, 4 Mar 1981 (st),
BOLiVAR:Between Santa Teresita de Kavanayen and Gentryet al. 31938, 31976
Rio Pacairao,20-21 Nov 1944 (6), Steyermark60399 shu Station,8 (U); Rio Manu,CochaCa-
Aug 1983 (st), Gentryet al. 43454 (BG);
Pourouma
upperRio Madrede Dios, 17 Jun 1954 (6),Rauh 1609
(F). UCAYALI: Pucallpa-TingoMaria rd., San Alejan-
dro, Gentryet al. 16181 (F, MO, NY); Pucallpa-Tingo
Mariard., km 88, 13 Mar 1982 (st), Gentryet al. 36309
(BG).
BRAZIL.ACRE: Upper Rio Moa, FazendaArizona,
10-15 Oct 1985(st),Campbellet al. 6250 (BG);Abuna-
Rio Brancohwy., km 242-246, 18 Jul 1968 (6),Forero
et al. 6367 (C, INPA, MO, NY, S, UC, US); nr. mouth
of Rio Macauhan,tributaryof Rio Yaco, 4 Aug 1933
(9), Krukoff5282 (A, G, LE, M, MO, NY, S, U, UC,
US, type collectionof P. subplicata);rd. Brasileia-As-
sis, km 16, 3 Nov 1980 (9),Lowrieet al. 710 (U); Sena
Madureira,3 Oct 1968 (9), Prance et al. 7792 (GH,
INPA, M, NY, P, R, S, U); Rio Jurua-Mirin,Aug 1901
(6), Ule 5718 (B, G, MG, type collection of P. aurea).
AMAPA: Confluenceof Rio Oiapoqueand Rio Ian6,27
Aug 1960 (6), Irwin et al. 47874 (F, FHO, GH, MG,
NY, RB, US); Rio Oiapoque, nr. mouth of Rio Ya-
roupi, 24 Sep 1960 (6), Irwin et al. 48468 (M, MG,
NY, P, RB, UC); Rio Oiapoque,nr. CachoeriaSanta
Maria, 21 Aug 1961 (6), Pires et al. 50419 (FHO, G,
MG, NY, S, US);40 min up Rio Falcino, 14 Nov 1961
(8), Pires et al. 50941 (F, MO, NY); Serrado Navio,
26 Sep 1961 (Q),Pireset al. 51247 (GH, MG, NY, MO,
U, US); AMAZONAS: Manaus-Itacoatiara rd., Reserva
FlorestalDucke,6 Aug 1976 (st),Aluisio(INPA)70809
(INPA);Uaup6s, SerraSao Gabriel, 14 Feb 1959 (9),
Cavalcante606 (MG);Rio Uatumi, Itapiranga,24 Aug
1979 (6), Cid et al. 721 (INPA, U); Rio Jurua, Sta.
Rosa, 15 Aug 1975 (st), D. Coelhoet al. (INPA)51350
(INPA), 22 Aug 1975 (st), D. Coelho et al. (INPA)
52381 (INPA); Manaus-Itacoatiarard., Reserva Flo-
restalDucke, 28 Sep 1962 (8),Duarte7194 (INPA, M);
Manicor6, nr. Santa F6, 8-11 Sep 1934 (9), Krukoff
6041 (A, B, BR, F, LE, MO, NY, S, U, US); Humaita,
plateaubetweenRio Livramentoand Rio Ipixuna,7-
18 Nov 1934 (9), Krukoff7071 (A, BR, G, GB, LE,
MO, NY, RB, S, S, U, US, type collection of P. um-
bellata);Sao Paulo de Olivenca,nr. Palmares,11 Sep-
26 Oct 1936 (9), Krukoff8073 (A, BR, G, LE, MO,
NY, P, S, U, US, typecollectionofP. isophlebia);upper
Rio Negro Uaupes, 12 Feb 1959 (9), Rodrigues884
(U); Tefe, 24 Nov 1959 (st), Rodrigueset al. 1424
(INPA); Manaus-Itacoatiorard., Reserva Florestal
Ducke, 19 Oct 1965 (6),Rodrigues7532 (INPA);Tefe,
23 Aug 1979 (6), Rodrigueset al. 10182 (INPA);Ma-
naus-P6rtoVelho rd., betweenRio Castanhoand Rio
Tapuna,18 Jul 1972 (8),M. F. Silva et al. 903 (INPA).
MATOGROSSO:Aripuana, 18 Jun 1975 (8), Cordeiro
140 (US), 25 Jan-14 May 1977 (st), Gomeset al. 547,
593, 632, 635, 646, 655, 673, 690, 759, 789, 810, 813,
838, 872, 891, 924, 926, 931, 933, 955, 1033, 1118,
1120, 1129, 1156, 1175, 1255, 1319, 1336, 1338, 1356,
1378, 1419, 1423, 1425, 1444, 1446, 1534, 1568, 1570,
1577, 1584, 1593, 1594, 1595, 1636, 1640, 1641, 1692,
1731, 1745, 1769, 1822, 1823, 1867, 1893, 1917, 1983
(INPA), 14 Jul 1977 (6), Gomeset al. 2170 (INPA), 19
Jan 1979 (9), 29 May 1976 (6), Monteiro et al. 1119
(INPA), M. G. Silva et al. 4326 (MG), 1 Jan 1979 (6),
M. G. Silva et al. 4738 (MG), 8 Oct 1979 (9),Rylands
25 (INPA);Mun. Vila Bela da SantissimaTrinidade,
4 km S of border with Rond6nia, 3 Nov 1985 (2),
189
Thomaset al. 4794 (BG).PARA:Cuiaba-Santaremrd.,
km 919, 13 Nov 1977 (9),Pranceet al. 25336 (F, RB,
U). RONDONIA: Mun. PresidenteMedici, rd. Cuiaba-
P6rto Velho, km 300, 25 Jun 1984 (d),Cid et al. 4798
(BG);Mun. Ouro Preto, rd. Cuiaba-P6rtoVelho, km
353, 39 Jun 1984 (6), Cid et al. 4919 (BG);Mun. Pi-
menta Bueno, Guapor6,7 Nov 1964 (st), Vieiroet al.
985 (U). RORAIMA: Serrados Surucucus,19 Feb 1969
(2), Pranceet al. 10089 (G, INPA, NY, R, S, U, US),
20 Feb 1969 (2), Prance et al. 10124 (INPA, NY, S,
U, US); Rio Negro, left bank of Paranado Marara,7
Feb 1977 (9), M. R. Santos 172 (MG, U).
BOLIVIA.BENI:Rio Beni, CachuelaEsperanza,29
Nov 1922 (6),Meyer403 (U). PANDO: Barrola,17 km
from Cobija, 18 May 1977 (st), Meneces 613 (MG,
MO); San Francisco, 50 km from Cobija to Puerto
Rico, 1 Jun 1977 (6), Meneces 647 (MO);rd. Cobija
to Extrema,Villa Marieta,24 Jun 1978 (2), Meneces
725 (INPA);3 km aboveAbuna,13km 1968(a),Prance
et al. 8378 (B, INPA, NY, S, U, UC, US); Rio Na-
reuda, Nicolas Suarrez,SW of Cobija, 31 Jul 1982
(5), Sperling et al. 6430 (BG).
Vernacular names. Colombia. Vaupes: uva sil-
vestre. Venezuela. Bolivar: majagua, cay-wari-
cay-yek, coiwaricoi-yek, kai-wa-rei-kei-yek. Su-
rinam. Bospapaja, granboesipapaja. French
Guiana. Bois canon, bois canon male or male
bois canon, bouchi papaie and papaye (Para-
maka), kulumasi (Wayapi). Peru. Amazonas:
sacha uvila, shuvija (Aguaruna); Huanuco and
San Martin: uvilla or uvilla blanca; Loreto: uvilla
lanuda. Brazil. Amapa: mapatirana; Amzonas:
imbaibarana, purumai, tourem; Mato Grosso:
caramuri, imbaubarana, torena. Bolivia. Pando:
ambaibochi, ambaibillo.
The species is rather variable in the dimen-
sions and shape of the leaves. The indument
mostly consists of appressed hairs but patent hairs
occur in the Upper Amazon Basin, e.g., in the
type collection ofP. aurea. The number of lateral
veins varies from 6 to ca. 25 pairs, but in some
collections (of juvenile material?) probably be-
longing to P. minor, e.g., Ayala 341 and Gentry
et al. 22381, the number of lateral veins may rise
to 40 pairs.
The species is a clear-cut one, characterized
by the subumbellate pistillate inflorescence and
the knob-like stigma. It shows resemblances of
P. bolivarensis (see p. 186).
The fruits are eaten by the monkey Cebus apel-
la (Foster et al. 5019).
From the three syntype collections from French
Guiana (Benoist 960, Benoist 978, and Wach-
enheim 18) Benoist 960 has been selected as the
lectotype collections (Berg, 1975).
190 Flora Neotropica

Pourouma napoensisand P. herrerensishave fromthe latterin the staminateflowerswith their

been included in this revision after submission connatetepalsand fullyconnatefilaments.These
of the (pre-review)manuscriptand are,therefore, featuressuggesta close relationshipto P. hirsu-
placed at the end of the systematic treatment, tipetiolata.
and not after P. hirsutipetiolataand P. mollis,
respectively. 25. PouroumaherrerensisC. C. Berg,Candollea
44(2): 513. 1989. Type, Peru, Loreto, Prov.
24. Pouroumanapoensis C. C. Berg, Brittonia Requena, Jenaro Herrera, Reserva Forestal,
42: 59-65. 1990. Type. Ecuador. Napo: Re- tree4/122, 2 Oct 1985 (d),Spichigeret al. 1995
serva Biologia Jatun Sacha, 14 Aug 1987 (6), (holotype, G; isotype, BG). Fig. 93.
Palacios 1874 (holotype, QCNE; isotypes, Tree, up to 25 m tall. Leafy twigs 3-6 mm
AAU, BG, MO, NY, QAME). Fig. 92. thick, minutely puberulousto hispidulous (and
+ scabrous)or also with much longer, yellow
Tree, up to 25 m tall. Leafy twigs 1-2.5 cm
hairs, pluricellularhairs dark brown and rather
thick, yellow-hirsute,pluricellularhairs absent. Laminaentire,oblongto ellipticto ovate,
Lamina 7-fid to -parted,ca. 25-45 x 25-45 cm, sparse.
8-20 x 3-11.5 cm, (sub)coriaceous,apex short-
coriaceous,apex acuminate,base deeply cordate base acute to rounded;uppersurface
with overlappinglobes; uppersurfacesmooth to acuminate,
sparsely appressed-puberulous on the midrib or
slightlyscabrous,yellow-hirsuteto -hirtellouson the whole surfacecoveredwith
the whole surface or only on the main veins, long, yellowhairs,
lower surfaceappressed-puberulous on the main
lower surfaceyellow-hirsuteto -hirtellouson the
veins or also with long yellow hairs, arachnoid
veins, arachnoidhairs in the areoles and on the hairsin the areolesand on the
reticulum;lateral
reticulum;lateral veins 16-25 pairs; petiole ca. veins 9-14 basal pair unbranched or
25-40 cm long,yellow-hirsute;stipules10-16 cm pairs,
branched,the basal part of these veins forming
long, outside yellow-hirsute,inside glabrous,ca- the basal
ducous. Staminate inflorescencesca. 12-22 cm part of the leaf margin (thus not sep-
arated from the margin by mesophyll), tertiary
long, ca. 8-14 cm wide; peduncle 4-8 cm long, venation
slightly prominent; petiole 3-10 cm
peduncleand branchesyellow-hirtellousto -pu-
long, minutely puberulousor also with long, yel-
berulous;flowers (sub)sessile,in globose heads, low 3.5-8 cm long, caducous,out-
these 7-12 mm diam.;perianth,tubular,1-2 mm hairs;stipules
side densely puberulousand with brown, pluri-
high, puberulousat the apex; stamens 3-4, fila- cellular
ments completleyconnate, 4-6 mm long. Pistil- hairs, or also with long, yellow hairs,
sometimes only a few appressed ones, inside
late inflorescencesup to 10 cm long and up to
12 cm wide, branched;peduncle 3-5 cm long, sparsely short-pubescent. Staminate inflores-
cences up to 10 cm long and up to 9 cm wide;
peduncleand branchesyellow-hirsuteto -hirtel-
peduncle 2-6 cm long, peduncle and branches
lous;flowers ca. 40-70; pedicel 0.3-0.5 cm, in
fruit up to 1.2 cm long; perianthca. 5 mm long, densely puberulousto short-velutinousor also
with long, yellow hairs;flowers sessile or pedi-
yellow-hirsuteto -hirtellous, sparsely so in the
cellate,most of them in globoseheads,numerous
uppermostpart;stigma peltate. and 3-4 mm diam. or less (5-12) and 5-8 mm
Distribution(Fig. 81). Ecuador(Napo),in non-
inundatedforest, at low altitudes. diam.; tepals ca. 1 mm long, connate, forminga
+ urceolate perianth, densely hairy; filaments
Specimensexamined. ECUADOR. NAPO:Reserva exceedingthe tepals,free.Pistillateinflorescences
BiologicaJatunSacha,Rio Napo,8 kmbelowPuerto up to 14 cm long and 9 cm wide; peduncle and
Misahualli, 4 Sep1987(2), Cer6nM. 2019(BG);8 km branchesshort-velutinous;flowers 7-25, in 1-4
SE of Tena,30 Aug 1960(2), Grubbet al. 1545 (K, + distinct clusters,
pedicels up to 0.7 cm long;
NY);ReservaBiologicaJatunSacha,Rio Napo,8 km
belowPuertoMisahualli, 2 Oct1986(2),Palacios1337 perianth yellow-velutinous, except for the apex;
(BG),14 Aug 1987(a),Palacios1874(BG,typecol- stigma subpeltate.
lectionof P. napoensis);Lumbaqui,12 May1987(a), Distribution(Fig. 81). Peru (Loreto),in non-
Pennington et al. 12244(BG,K). inundatedforest.
In the shape,dimensions,and indumentof the Specimens examined. PERU. LORETO: Prov. Re-
lamina this species resemblesP. oraria.It differs quena,JenaroHerrera, Aug-Sep1976(6),Bernardi s.n.
Pourouma 191

f
In

i
bi

FIG.92. Pouroumanapoensis.a. Leaf(Grubbet a. 1545). b. Stipulesand staminateinflorescence(Pennington

et al. 12244). c. Pistillate inflorescence(Grubbet al. 1545). d. Staminateflower (Penningtonet al 12244). e.
Pistillateflower(CeronM. et al 2019).
 .At

FIG.93. Pouroumaherrerensis.a. Leafytwigwith staminateinflorescences(Spichigeret al. 1995). b. Pistillate

inflorescence(Spichigeret al. 1999). c. Leaf (Vasquezet al. 2623).
Pourouma 193

(BG),9 Dec 1977(d),Gentryet al. 21346(U);Prov. tepalsof the staminteflower,and the sparseplur-
Maynas,PuertoAlmendras, 11Nov 1984(9),Maaset icellularhairs.
al. 6267(BG,U);Prov.Requena,JenaroHerrera, Re- Thesespecimensmay representa distinctcloud
servaForestal,2 Oct 1985(d),Spichigeret al. 1995
(BG,G, typecollection),17 Sep 1982(6), Spichiger et forest species (see p. 114).
al. 1996(BG,G),9 Oct1982(st),Spichiger et al. 1997
(BG,G), 14 Sep 1982(d),Spichigeret al. 1998(BG, Rio Corrientes,betweenTiente
2. PERU. LORETO:
G), 10 Nov 1982(9),Spichigeret al. 1999 (BG,G); LopezandPuestoAvanzado,4 Apr1977(st),Gentry
Prov.Maynas,PuertoAlmendras,17 Apr 1980 (6), etal. 19048(BG);SantaRosa,1-5 Sep1929(st),Killip
Vdsquez et al. 167 (BG),21 Oct 1981(9), Vdsquez et &A. C. Smith28800(F, NY).
al. 2623(BG);Prov.Maynas,RioNanay,Mishana,24 BRAZIL.ACRE: ReservaINCRASantaLuzia,BR-
Apr1986(9), Vdsquez et al. 7535(BG). 364,km40, 5-19 Oct 1984(st),Campbell et al. 6820
(NY)and7689(BG).
This species has a strangemixtureof features.
The staminateinflorescencesand flowersresem- These have ovate to elliptic leaves (16-32 x
ble those of P. mollis,the pistillateinflorescences 12-22 cm) with a deeply cordatebase. The leafy
and flowersthose of P. mollis subsp. trilobaand twigs, petioles, stipules, and main veins at the
P. acuminata. In the indument it differs from lower surface of the lamina have dense, long,
both. One of the forms of this species has very patent, yellow hairs. The stipules are glabrous
short and stiff hairs, resultingin a slightly sca- inside. The presence of arachnoidhairs on the
broussurfaceof the leafytwigs,petioles, stipules, petiole, main veins of the lamina beneath, and
and main veins of the laminabeneath.The other stipulessuggestaffinitiesto P. tomentosa.These
form has this short indument mixed with much collections may representa new subspecies of
this species.
longer yellow hairs, as in P. cucura,and bears,
moreover, long yellow hairs on the upper leaf 3. VENEZUELA.AMAZONAS: Nr. Yavita, 21 Apr
surface,like some forms of P. cucura.Some of 1970 (st), Steyermark et al. 102874(BG,MO, NY,
the specimenshave oblongto ellipticleaves with VEN);Dept.RioNegro,Cerrode la Neblina,RioMa-
the basal pair of lateralveins unbranched;such warinuma, 8-10Jan1984(st),Steyermarket al. 129773
(MO).
specimensbear some resemblanceto P. acumi-
nata. Other specimens (with long hairs) have These collections have thickly coriaceous
ovate leaves and the basal pairs of lateral veins leaves, thick leafy twigs with conspicuous lenti-
branched. However, both types of leaves may cels, and dense, almost black, pluricellularhairs
occur on the same plant, or even on the same on the leafy twigs and stipules. In these features
branch. they match P. ferruginea,but the characteristic,
The materialwith a scabrousupper leaf sur- dense (white to brownish)villous-arachnoidin-
face, provisionally kept in P. mollis subsp. tri- dument on the lower surface of the lamina is
loba, resemblesP. herrerensisin the variationof wantingand the stipulesare denselyhairyinside.
the leaves. These collections might representan abberant
form of P. ferrugineaor perhapsof P. melinonii
subsp. melinonii.
Unnamed Collections
4. COLOMBIA.VAUPES:Alto Vaupes,25 Nov 1975
1. PERU. AMAZONAS:
Serraniade Bagua,ca. 12- (9),Roa251 (INPA).
18 km on trailE of La Peca, 1800-1950m, lower
montanecloudforest,14 Jun 1978(6), Gentryet al. This collection has leafy twigs with yellow-
22866 (U). SANMARTIN:Prov. Rioja, rd. PedroRuiz- hirsuteindumentand conspicuouslenticels.The
Moyobamba, km390,Venceremos, 2100m, 7-9 Aug indument of the
1983(st),D. N. Smithet al. 4742(MO). leafy twigs and the leaf surface
resemblesthat of P. cucura,but the thickly co-
The indumentof the leafy twigs,stipules,pet- riaceous,somewhatplicate lamina has the hairs
ioles, and lower surface of the lamina largely on its uppersurfaceconfinedto the main veins.
matches that of P. bicolor subsp. bicolor. The Moreover, the stipules are hairy inside and the
specimens differfrom that taxon in the smooth reticulumon the lamina beneath is very prom-
and puberulousuppersurfaceof the lamina, the inent. The fruiting perianth is scabrous. This
broadly ovate lamina, the slender (ca. 15 x 10 specimenshas a mixture of charactersof P. cu-
cm) staminateinflorescence,the oblongto elliptic cura and P. bicolorsubsp. bicolor.
194 FloraNeotropica

5. ECUADOR.NAPO:Aiiangu, 1-15 Feb 1986 (st), E. Kieft (Utrecht), and some elements of that
Korning& Thomson47801and47807(BG). unfinishedstudy have been used by the present
The latterhas most charactersin common with authors(C.C.B.and E.C.H.vH.). The senior au-
the form of P. bicolorsubsp. bicolorwith an en- thoris indebtedto the Universityof Bergen,Fac-
tire lamina, being smooth above. However, the ulty of Mathematicsand Natural Sciences, for
indument of the leafy twigs, petiole, and lower coveringtravelingexpenses,and to the Botanical
Institute at Bergen for their assistance. The il-
leaf surfaceresemblesthat of P. cucura.The for-
mercollectionwith 5-partedleaves and incisions lustrations were preparedby Ms. Siri Herland
down to the petiole may representthe juvenile (Bergen), Mr. H. Rypkema(Utrecht),and Mr. T.
formof the lattercollection.The mixtureof char- Schipper(Utrecht).The authorsare indebted to
acters suggestspossible hybridization. Ms. KathleenWellingfor processingpart of the
text, to Drs. W. H. A. Hekking (Utrecht), Drs.
Names Excluded E. Simonis (Utrecht), Ms. Edit Fjaere (Bergen)
and Ms. FrancinaBerg (Bergen)for their assis-
Pouroumaparaensis-nomen (in schedule)giv- tance in preparingthe list of exsiccatae, and to
en by Huber;refersto Goeldi(MG) 7732 = P. Drs. E. Simonis for his contributionsto the prep-
melinonii subsp. melinonii. aration of the lists of specimens examined and
PouroumaretusaBentham-nomen, see Miquel the distributionmaps.
in Martius,Fl. bras. 4(1): 128. 1853; refersto
Spruces. n. from Brazil,Amazonas,Manaus- LITERATURECITED
syntype collection of P. tomentosa.
PouroumatrianaeA. Richter-nomen (in sche- Akkermans,R. W. A. P. & C. C. Berg. 1982. New
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PouroumauleiWarburgex Ule-nomen; see Ule, piaceae),and keys to its species. Proc. Kon. Ned.
Bot. Jahrb. Syst. 40: 132. 1907; refersto Ule Akad. Wetensch.,Ser. C, 85(4): 441-471.
Aublet, J. B. C. F. 1775. Histoire des plantes de
9314 = P. cecropiifolia. Guiane francaise2. Paris.
Benoist, R. 1922. Descriptionsd'especes nouvelles
ACKNOWLEDGMENTS du genre Pourouma(Moracees).Bull. Mus. Nat.
Hist. Nat. (Paris)28: 318-321.
The authorsof the Coussapoarevision (C.C.B. . 1924. Descriptions d'especes nouvelles de
and R.W.A.P.A.) are much indebted to Dr. N. Phan6rogamesde la Guyanefrancaise.Bull. Mus.
Nat. Hist. Nat. (Paris)30: 103-105.
G. Bisset (ChelseaCollege,London) for correct- . 1933. Lesbois de la Guyanefranqaise.Mora-
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T. Prance & T. E. Lovejoy, Key environments:
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NUMERICAL LIST OF TAXA

1. Coussapoa 1-28. C. nitidaMiquel
1-1. C. angustifoliaAublet 1-29. C. nymphaeifoliaStandley
1-2. C arachnoideaAkkermans& C. C. Berg 1-30. C. oligocephalaDonnell Smith
1-3. C. argenteaAkkermans& C. C. Berg 1-31. C. orthoneuraStandley
1-4. C. asperifoliaTr6cul 1-32. C. ovalifoliaTr6cul
a. subsp. asperifolia 1-33. C. pachyphyllaAkkermans& C. C. Berg
b. subsp. magnifolia(Trecul)Akkermans& C. 1-34. C. parvicepsStandley
C. Berg 1-35. C. parvifoliaStandley
c. subsp. rhamnoides(Standley)Akkermans& 1-36. C. purpusiiStandley
C. C. Berg 1-37. C. scabraAkkermans& C. C. Berg
1-5. C. batavorumAkkermans& C. C. Berg 1-38. C. spruceiMildbraed
1-6. C. brevipesPittier 1-39. C. tessmanniiMildbraed
1-7. C. chocoensisCuatrecasas 1-40. C. trinerviaMildbraed
1-8. C. cinnamomeaCuatrecasas 1-41. C. vannifoliaCuatrecasas
1-9. C. cinnamomifoliaMildbraed 1-42. C. villosaPoeppig& Endlicher
1-10. C. contortaCuatrecasas 1-43. C. viridifoliaCuatrecasas
1-11. C. crassivenosaMildbraed 1-44. C. fulvescensC. C. Berg
1-12. C. cupularisAkkermans& C. C. Berg 1-45. C. tolimensisC. C. Berg
1-13. C. curraniiBlake 1-46. C. valariaC. C. Berg
1-14. C. duqueiStandley 2. Pourouma
1-15. C. echinataAkkermans& C. C. Berg 2-1. P. guianensisAublet
1-16. C.ferrugineaTrecul a. subsp. guianensis
1-17. C. floccosa Akkermans& C. C. Berg b. subsp.venezuelensis(Cuatrecasas)C. C. Berg
1-18. C. glaberrimaBurger & van Heusden
1-19. C. herthaeMildbraed 2-2. P. velutinaMiquel
1-20. C. latifoliaAublet 2-3. P. bicolorMartius
1-21. C. leprieuriiBenoist a. subsp. bicolor
1-22. C. longepedunculataAkkermans& C. C. Berg b. subsp. tessmannii(Mildbraed)C. C. Berg&
1-23. C. macerrimaAkkermans& C. C. Berg van Heusden
1-24. C. manuensisC. C. Berg c. subsp. digitata (Tr6cul) C. C. Berg & van
1-25. C. microcarpa(Schott)Rizzini Heusden
1-26. C. microcephalaTrecul d. subsp. scobina (Benoist) C. C. Berg & van
1-27. C. napoensisAkkermans& C. C. Berg Heusden
Exsiccatae 197

e. subsp.chocoana(Standley)C. C. Berg& van a. subsp. tomentosa

Heusden b. subsp. apiculata(Benoist)C. C. Berg& van
2-4. P. cecropiifoliaMartius Heusden
2-5. P. cucuraStandley& Cuatrecasas c. subsp.persectaC. C. Berg& van Heusden
2-6. P. cuspidataMildbraed d. subsp.essequiboensis(Standley)C. C. Berg&
2-7. P. myrmecophilaDucke van Heusden
2-8. P. formicarumDucke 3. subsp. maroniensis(Benoist) C. C. Berg &
2-9. P. phaeotrichaMildbraed van Heusden
2-10. P. mollis Tr6cul 2-14. P. stipulaceaC. C. Berg
a. subsp. mollis 2-15. P. acuminataMiquel
b. subsp.triloba(Tr6cul)C. C. Berg&van Heus- 2-16. P. ferrugineaStandley
den 2-17. P. villosaTrecul
2-11. P. melinoniiBenoist 2-18. P. orariaStandley& Cuatrecasas
a. subsp. melinonii 2-19. P. saulensisC. C. Berg& Kooy
b. subsp. glabrataC. C. Berg& van Heusden 2-20. P. ovataTr6cul
2-12. P. hirsutipetiolataMildbraed 2-21. P. ellipticaStandley
a. subsp. hirsutipetiolata 2-22. P. bolivarensisC. C. Berg
b. subsp.hispida(Standley& Cuatrecasas)C. C. 2-23. P. minorBenoist
Berg& van Heusden 2-24. P. napoensisC. C. Berg
2-13. P. tomentosaMiquel 2-25. P. herrerensisC. C. Berg

LIST OF EXSICCATAE
Some recentcollections have been added to this list without being cited in the lists of specimens.
Acosta M., A., 12 (2-5).
AcostaSolis, M., 6008 (1-10); 12661 (1-19); 12797 (1-
10);12884, 13605(2-3d);13629(1-42);13687, 13713
(1-10).
Alencar,L., 497 (2-4).
Allemao, F. et al., 446 (2-la).
Allen, P. H., 3103 (39); 3251 (2-1 a); 3742 (2-3d);
5120 (1-4b); 5949 (1-23); 6186, 6280 (1-42); 6401
(2-3d).
Aluisio, J., 70809 (2-23).
Amaral, I. L. et al., 101 (2-11a); 473 (2-15); 478 (1-
40); 595 (2-1 la); 604 (2-5); 666 (2-23); 761 (2-7);
IG2-10-328, IG2-16-509 (2-5).
Ameida, J. et al., 71 (2-10a).
Ancuash,E., 178 (2-la); 290 (2-1 la).
Anderson,C. W., 188, s.n. (1-26).
Andrade,A. et al., 2340 (1-25).
AndradeLima, D. de, see Lima. D.
Anonymus, 9783 (1-20); s.n. (2-4, 1-25).
Appun, C. F., 296 (1-26).
Araujo,D. S. Dunn de et al., 848 (1-25).
Aristeguieta,L., 1699, 1704 (1-42); 2005 (2-16).
Asanza C., E., 32929 (2-4).
Asplund, E., 9462, 12933 (2-4); 13020 (2-la); 14124
(2-4); 16401(1-41);18780(1-4b); 18840(1-9); 19835
(1-42).
Aublet, J. B. C. F., s.n. (1-20, 2-la, 2-1 la, 2-17).
Aubreville,A., 144 (2-3c).
Aubry-Lecompte,C. E., s.n. (1-20).
Austin, D. F. et al., 4140 (2-3a); 7149 (1-20).
Ayala, F. (et al.), 341 (2-spec.); 1436 (1-42); 1446 (2-
4).
Aymard,G. et al., 3990 (2-23); 6778 (2-15).
BAFOG,93-M (2-17); 1050 (2-3c);4272 (2-1 la); 5042
(2-23);7087 (2-17), 7152, 7173 (2-23);7228 (2-13e);
7261, 7268 (2-23); 7330 (2-13e); 7562 (2-2); 7745
(2-17); 7938 (1-1); 7942, s.n. (1-4a).
Bailey, I. W., 18, 19, 20, 146, 147 (2-la).
Baker,M. A., 6017 (2-3a).
Balick,M. et al., 1021 (1-39).
Barbour,P. J., 5372 (1-42).
Barclay,A. S. et al., 3294, 3593 (1-42).
Barrier,S., 5114 (2-13e).
Bartlett,H. H., 12356 (1-30).
Bastos, M. A., 2051 (2-la); 2220 (2-23).
BBS, 3003 (1-20).
BBS (=BosbeheerSuriname),297 (2-17); 298 (2-13a);
1128 (2-la).
Belem, R. P. (et al.), 644 (1-25); 1471 (2-la).
Benoist, R., 341 (2-17); 465 (2-13e); 960, 978 (2-23);
3003 (2-3d); 3010 (1-10); 3042 (1-41); 3044 (1-42).
Berg,C. C. (et al.), 276 (2-7); 355 (2-1 lb); 400 (2-23);
BG.598, BG.772 (2-la); 1033 (1-42); 1038 (1-40);
1040 (1-42); 1041, 1042, 1045 (2-5); 1048 (1-40);
1053 (2-5); 1058 (1-40); 1062 (1-42); 1070 (2-5);
1111 (1-31); 1113 (1-42); 1119 (1-4b); 1120, 1129,
1131 (1-11); 1132 (1-32); 1133 (1-4b); 1143 (2-2);
1153 (1-33); 1222 (1-42); 1223 (2-4); 1241 (1-42);
1259 (1-19); 1301, (1-42); 1223 (2-4); 1546 (1-44);
P.17591, P.17592 (1-28), P.18136 (2-1 la); P.18152
(2-3a);P.18409, P.18453 (1-40); P.18455, P.19841
(1-42); P.19878 (2-13b), s.n. (1-4b, 1-42, 2-13a).
Berlin,B., 372 (1-42); 523 (2-4); 746 (2-lOb);1531 (1-
32); 1999 (2-4).
Bernardi,L., 902 (2-1 la); 910 (2-3a);1724(1-42); 1919
(2-3d); 1980 (1-42); 1986 (2-3d); 2657 (2-3a); 2731
(2-13b);2744 (1-43);2808 (2-1la); 3354 (1-42);5696
(2-lb); 6348, 7017 (1-42); 7238 (1-3); 7250 (2-23);
5.16.5. (2-20);s.n. (tree3/100) (2-25); s.n. (tree4/8)
(2-23); s.n. (tree 6/27) (2-13a);s.n. (1-42, 2-lb, 2-5,
2-13a).
Bisby, F. et al., P.18118 (2-3a).
Billiet, F. et al., 1108 (1-20).
Black,G. A. (et al.), 46-137 (1-8);47-1520 (2-13a);48-
2949, 48-3017 (1-28).
Blanchet,J. S., 160, 2327 (1-25); 2361 (2-lOa);s.n. (1-
25).
Blanco,C., 810 (2-la); 958 (1-42); 1149 (2-3a).
198
Blum, J. L., 2039 (2-3e).
Boerboom,J., 9199 (2-10a).
Bondar,G., 2169 (2-10a);s.n. (2-la).
Boom, B. et al., 7812 (1-4b); 7870 (1-9); 8102 (2-14).
Bowie, J. et al., 57 (1-25).
Boyan, R., 267 (2-7).
Brade,A. C. (et al.), 7887, 7949 (1-25); 18619 (2-la).
Braga,P. L. S., 3178 (2-4).
Brandbyge,J. et al., 30194 (2-3a);30252 (2-23);30258
(1-31); 30327 (2-23); 30421 (1-32).
Brenes,A. M., 20542 (1-34).
Breteler,F. J., 3889 (1-42); 4920 (2-3d).
Brewer-Carias,C., s.n. (1-11).
Broadway,W. E., 880 (1-4a).
Bruijn,J. de, 1123 (1-42); 1496 (2-1 lb); 1497 (2-3a);
1502 (2-llb); 1514, 1516, 1517, 1526 (2-3a); 1546
(2-1 lb); 1547 (2-12a); 1555 (2-1 lb).
Buchtien,0., 2050, 2122 (2-3a).
Buck, P., 26425 (1-25).
Bunting,G. S. (et al.), 3959 (1-4b); 6836 (2-3b).
Burchell,W. J., 3148 (1-25); 9792 (2-3a and 2-lOa).
Burger,W. C. et al., 8137 (2-23); 10040, 10474 (1-42).
Busey, P., 628 (1-34).
BW (=Boschwezen)732 (1-20);772 (1-1); 1322 (1-4a);
1368 (2-2); 2030 (2-3c); 2070 (1-4a); 2095 (2-17);
2410 (2-lOa);3255, 3302 (2-17); 3376 (2-3a); 3430
(1-4a);4010 (2-17);4033 (2-3a);4494 (2-1Oa);4866
(2-3c); 5036 (2-3a);5048 (2-10a);5262 (1-4a);5390
(2-17); 5440, 5581, 5992 (2-10a);6292 (2-3c); 6509
(2-17).
Cabrera,J., 2668 (1-35).
Callejas,R. et al., 3997 (1-9).
Calzada,J. I., 325 (1-36).
Camargo, -, 1849 (1-25).
Camp, W. H., E3626 (1-42); E3674 (1-19).
Campbell,D. G. et al., 6820 (2-unnamed);6834 (2-6),
6927 (2-5); 7001 (2-la); 7157 (2-3a); 7264 (2-23);
7600 (2-3a); 7689 (2-unnamed);8065, 8127, 8267
(2-la); 8344 (2-23); 8383 (2-lOb);8463 (2-23); 8537
(2-la); 8844 (2-4); 8997 (2-3a); 9120 (2-lOb);9521
(2-3a); 9610 (2-la).
CamposPorto, P., 863 (1-25).
Cantonnementde Cayenne,s.n. (2-1 la).
Carauta,J. P. P. (et al.), 178, 1714, 1716, 1780, 1781,
2722; 3011 (1-4a); 3050, 3166 (1-25).
Cardenas,M., 1990 (2-1a); 3973 (2-13c).
Cardona,F., 374 (1-4b); 1222 (2-13b); 2170 (2-5).
Casaretto,J., 617 (1-13); 643 (1-25).
CastilloS., M., 11 (2-la).
Cavalcante,P., 606 (2-23); 777 (2-3a); 2297 (2-3c).
Cazolet,P. C. D. et al., 5037, 5072 (2-3d).
Cer6nM., C. E. (et al.), 252 (2-4); 261 (2-la); 2019 (2-
24); 2955 (1-14); 2993 (1-9); 3389 (1-32); 3495 (1-
22).
Chagas,J., 962 (2-4); 1364 (1-4a); 1564 (2-7); 1655 (2-
2).
Cid (Ferreira),C. A. et al., 435, 511 (1-40); 721 (2-23);
847 (2-13d); 849 (2-2); 942 (1-20); 1053, 2223 (1-
1); 2870 (2-la); 3297 (2-20); 6200 (2-2); 7031 (2-
30); 7831 (1-4a).
Claes, F., 26 (2-4).
Clewell,A. et al., 4142 (2-3d).
Flora Neotropica
Coelho, D. (et al.), 672 (2-1 la); 1577 (1-4a); 5979 (2-
20); 51350, 52381 (2-23); 52383 (2-13d).
Coelho, L. (et al.), 15 (2-17); 332 (1-4c); 339 (2-16);
2265 (2-7); 5223 (2-13a); 5728 (1-31); 5808 (2-20).
Constanino,D., 19689 (1-25).
Contreras,E., 61, 845, 2220, 3559, 4449, 5787, 6231
(1-30); 10026 (2-3d); 10195 (1-30); 10763 (1-42);
11195 (2-3d).
Cook, O. F. et al., 157 (1-29); s.n. (1-30).
Cooper,G. P., 538 (1-42).
Cordeiro,M. R., 140 (2-23); 367 (1-4b).
Cordoba,J. J., 295 (1-42).
Correa,M. D. et al., 1002, 1031A (1-6); 1854 (2-3d).
Costa, M. da, 90 (1-39).
Cowan,R. S. (et al.), 1868, 1880 (1-26); 38774 (1-20).
Cremers,G., 6726, 8271 (2-lOa).
Croat,T. B., 5124,5125,6588 (1-4b);7633 (2-4);7768,
7839 (1-42); 7859 (1-4b); 8097, 8100, 8648 (2-3e);
9973A, 14212(1-6); 10343, 10830 (2-3e); 11591 (2-
3d); 15064 (1-6); 17784 (1-40); 18016 (2-la); 18643
(1-4b); 18850 (1-28); 18888 (2-15); 19997 (2-3a);
20370 (1-4c); 20411 (1-40); 20447 (2-1 a); 20616
(2-la); 20662 (2-4); 22635 (1-42); 24663 (1-30);
24831 (2-30); 59353 (2-3a).
Croizat,L., 312 (1-42); 343 (1-4b);541 (1-42);959 (1-
43); 962 (2-1la).
Crueger,H., s.n. (1-42).
Cruz, J. S. de la, 1175 (2-la); 1367 (1-26); 1501 (1-
26a); 1577 (1-26); 1657 (2-la); 1729 (1-4b); 2241,
2387, 2636, 2813, 3017 (1-26).
Cuatrecasas,J. (et al.), 6737 (2-4); 7299 (2-23); 7510
(1-42);7602 (2-4);8291 (1-42); 10714 (1-27); 14284
(1-10); 14881 (2-12b); 15071, 15520, 15534 (2-3d);
15589 (1-42); 15720 (2-18); 15876, 16315, 16755
(2-3e); 16826 (2-12b); 17251 (1-34); 17357 (2-3c);
17451, 17479 (2-3e); 17647 (1-10); 21256 (1-41);
21446 (1-7); 21496 (1-10); 22048 (1-41); 22526 (1-
42).
Curran,H. M., 8 (1-13); 19 (2-la); 21 (2-1Oa);116 (2-
12a).
DAF, 004 (1-13); 027 (2-lOa).
Daly, D. C. et al., 1208, 1474 (2-3c);3790 (2-17);3936
(2-2);4129, 4135 (2-7); 4457 (2-16); 4456, 4485 (2-
8), 5356 (1-11).
Damazio, L. B., s.n. (1-25).
Damiao, C., 2503, 2817 (1-28); 680 (2-20), see also
Mota, C. Damiao A. de.
Damiao, J. P. L. et al., 1805 (2-3a); 1822 (2-13b).
Daniel, H., 1793, 2652 (1-42).
Danta, M., 12383 (1-31).
D'Arcy,W. G., 12329 (1-6).
Davidse, G. et al., 15365 (2-2).
Davidson, C., 5203 (2-9).
Dayton, W. A., 3019 (1-42).
Denslow, J., 2698 (2-3e).
Deward,G., 27 (2-17).
Diaz, C. et al., 618 (1-32).
Diaz, M., 22A (2-20).
Dodson, C. H. (et al.), 3017 (1-39); 5241 (1-42); 5776
(1-19); 5778 (1-42); 6082 (1-19); 6156, 6312, 6327
(2-3d);6513, 7416, 7616 (1-41); 9539 (1-19); 15201
(1-34).
Doeve, -, 3003 (1-20).
Exsiccatae 199

Dombey, J., s.n. (1-42). Focke, H. C., 9, 418, s.n. (1-20).

Donnell Smith, J., 4826, 6770 (1-42). Foldats, E., 114-1A (2-2).
Donselaar, J. van, 1226 (2-13e); 1262 (2-3c); 1373, Folsom, J. P. et al., 6377 (2-3d); 6469 (2-23).
1681 (2-lOa);1888 (2-17); 2569 (1-1); 2924 (2-lOa); Fonnegra,R. et al., 1798 (1-35).
3333 (2-3c); 3788 (2-lOa). Forero,E. et al., 3145 (2-23); 4099, 4110(1-10); 6367
Dressier,R. L., 3454 (1-6); 3579 (2-3d); 4636 (1-15). (2-23), P.6419 (2-4).
Duarte, A. P. (et al.), 3410, 4650, 5328 (1-25); 6970 Forest DepartmentBritishGuyana,see FD.
(1-4a); 7194 (2-23); 9785 (2-1 la); s.n. (1-13). Foresta,H. de., 526 (2-13e); 676, 677, 4970, 5011 (2-
Ducke, A. (et al.), 104 (1-25); 1149 (2-7); 1354 (1-38); la).
1527 (2-16); 1528 (1-20); 1764 (2-7); 1795 (1-28); Foster, R. B. (et al.), 584 (2-3e); 631 (1-42); 1101 (1-
1917 (2-8); 1999 (1-38); 2103 (1-4a);6782, 7715 (1- 4b); 3412 (1-32); 5019 (2-23); 5641 (1-24); 5719 (2-
28); 9096 (1-38); 10736 (2-17); 12238 (1-28); 12366 10b);5721 (1-42); 5903 (2-4); 6568 (2-la); 6569 (2-
(2-4); 13056 (2-17); 13057 (2-3a); 13058 (2-10a); 10b); 7001 (2-4); 7194 (2-10b); 7199 (2-23); 7200,
13065 (1-4b); 13066 (1-28); 14499 (2-7); 15255 (2- 7201 (2-la); 7995 (2-lOb);8023, 9373 (2-la).
17);15261 (2-lOa);15350(2-2); 15804(1-4a); 15935 Francisco,2186 (2-7).
(2-2); 16003 (1-28); 16321 (1-25); 17132 (1-4b); Franco,J. et al., 419 (1-31).
17164 (2-7); 17167 (2-2); 18455 (1-40); 18456 (1- Frazao,A., s.n. (1-25).
20); 18460 (1-4c); 18461 (1-1); 18462 (1-21); 19455 Froes, R. de Lemos (et al.), 1907 (1-20); 20870 (2-8);
(2-1Oa);21200 (1-25);23606, 23607 (2-7);23608 (2- 21048 (2-4); 21264 (1-39); 23693 (2-16); 23725 (2-
20);23609 (1-38);25243 (2-17);25244 (2-16);25245 lla); 25531 (1-28); 26520 (2-13d); 26631 (2-3c);
(2-13b); 25246 (2-21); 25247 (2-2); 25248 (1-38); 28585 (1-4b); 34651 (1-20).
s.n. (2-13b).
Gandoger,M., 19 (2-2).
Duke, J. A. (et al.), 6588, 8035 (2-3d); 11273 (1-42); H., 12408(1-4b);13643(2-1la); 13836
11599 (2-3d); 14712 (1-6); 15742 (1-4b). Garcia-Barriga,
(2-23); 13871 (2-4); 14002, 14106 (2-5); 14116 (2-
Dunlap, V. C., 299 (1-42). 1la); 16239 (1-4b).
Duque-Jaramillo,J. M., 1767 (1-4); 2179 (1-8); 2207 J. H. et al., 9 (1-3a).
2446 Garcia,
(1-28); 2261, (2-4); 3957 (1-42).
Dus6n, P., 2160, 6635, 10134, 10186, 11419 (1-25); Gardner, G., 732, 5632, 5858 (1-25).
11942 (2-la); 13653, 14718, 17030 (1-25); 17239, Gamier, Bro. A., 112 (2-23).
17345 (2-la); s.n. (1-25, 2-la). Garwood,N. C., 759 (2-3d).
Dwyer,J. D., 4734 (2-3d); 10305 (1-42); 10799 (1-30). Gasche,J. et al., 29 (1-31).
Dyer, F. J., A-85 (1-42). Gaudichaud,C., 992 (1-25).
Edwards,J. B., 400 (1-42). Geay, M. F., 3285 (1-1).
Eggers,H. F. A. von, 14165, 15615 (1-42). Gentle,P. H., 2602 (1-30);2787, 2917, 3265, 4211 (2-
Egler,F. E., 42-229 (1-30). 3d); 4528 (1-30); 6312, 6997, 8675 (2-3d).
Egler,W. A. et al., 46027 (1-28). Gentry,A. H. (et al.), 918 (1-25); 2046, 2657 (2-1 lb);
Eiten, G. et al., 6214 (1-25). 6314 (1-42); 6679 (2-3e); 7530 (1-42); 7727, 8348
Ellenberg,H., 2288 (2-4); 2297 (2-lOb);2474 (2-4). (1-30); 8707 (1-42); 9552 (2-3d); 9606 (1-19); 9824
Elias, Bro., 130, 631 (1-42). (2-la); 9892 (1-19)9934, 10127, 12474(1-42);13408
Emden, W. C. van, L-VI, s.n. (1-20). (1-15); 14011 (see 1-32); 15621 (2-lOb); 15876 (2-
Emmerich,M. (et al.), 3003, 3556 (1-25). 7); 16041 (2-3b);16181 (2-23); 17217, 17493 (1-4b);
Emygdiode Mello Filho, L. (et al.), 412, 2340, 2447, 17804 (2-18); 18458 (1-42); 18528 (1-31); 19048 (2-
3018 (1-25). unnamed);20381 (1-28); 20356 (2-3a); 20495 (1-
Englesing,F. C., 50, 52a (2-3d). 28); 20515 (2-5); 20766 (1-40);20802 (1-28);21167
Erlanson,C. 0., 240 (1-42). (1-35); 21193 (2-3a); 21205 (2-la); 21291 (1-42);
Ernst,W. R., 324 (1-42). 21319 (1-4b);21346 (2-25);22095 (1-42);22280 (1-
Escobar,L. Albertde et al., 4219 (1-45). 4b); 22381 (2-spec.); 22460 (2-4); 22865 (2-13a);
Espina,J. et al., 219 (1-31). 22866 (2-unnamed);23713 (1-42); 23835 (2-3e);
Espinal,T. S., 822 (1-4b); 1314 (1-42). 24074 (1-4b); 24900, 25050, 25078, 25100 (2-5);
Falcao, M., 215 (2-4); 1105 (2-14). 25124 (1-28);25151 (1-32);25527 (2-23);25854 (2-
Fanshawe,D. B., 1105 (2-14); 2050 (2-la); 3031 (1- 5);26049, 26117 (2-23);26525 (1-10);26530 (2-3d);
26); 4970, 5011 (2-1a). 27194 (2-23); 27566 (1-22); 27799 (2-spec.);28609
Farifias,J. et al., 347 (2-4); 490 (2-1 la). (2-3d); 30278 (2-3e); 31179 (2-10b); 31197 (2-3b);
FD (=Forest DepartmentBritish Guyana), 1011 (2- 31271 (2-23); 31655 (2-5); 31816, 31933 (2-la);
la); 3031 (1-26); 3851 (2-14); 4014, 4073 (1-4b); 31938, 31976, 36231 (2-23); 36241 (2-4); 36309 (2-
4090, 4970, 5011, 5936, 6914 (2-la); 6973 (1-4b). 23); 36797 (2-18); 39264 (2-13b); 39379 (2-20);
Fendler,A., 1237, 1237a (1-42); 2420 (2-lb). 40479 (2-3e); 41890 (2-6); 41894 (2-13); 42029 (2-
FernandezP., A. (et al.), F3, 1458 (1-42); 2019 (2-23). la);42168 (2-3d);42232 (2-1Ob);42568 (2-23);42793
Ferreira,C. A. Cid, see Cid (Ferreira),C. A. (2-lOb); 43137 (2-4); 43454 (2-23); 45160 (2-3d);
Ferreira,V. F., 493 (1-25); s.n. (2-20). 45660 (2-4); 45915 (2-la); 45922 (2-5); 46154 (1-
Ferreyra,R., 4813 (1-32). 40);46752 (2-3c);47838 (1-46);48303 (2-3e);49026
Ferry,J. F., s.n. (1-42). (2-lOa);49127 (2-1 la); 49346 (2-13); 51089, 51111
Feuillet,C. P., 1070 (2-23). (2-23); 51154 (2-la); 51177 (2-5); 51207 (2-23);
200
51355 (2-19); 51515 (2-23); 53622 (2-3e); 53829 (1-
10); s.n. (2-5).
Gevieski, A., 85 (1-25).
Gillis, W. T. et al., 10191 (1-42).
Glaziou, A. F. M., 81, 1013, 1138, 1942, 2016, 4937,
6009 (1-25); 8934, 8934a (1-13); 8935 (2-la); 8936
(1-25); 10070a (1-31); 10070 (2-15); 12166 (1-25);
12173 (2-la); 16350, s.n. (1-25); 20408 (2-la).
Gleason, H. A., 174 (2-la); 405, 415 (1-26); 670 (2-
10a).
Goeldi, A., 7731 (2-10a); 7732 (2-1 la); 7773 (2-lOa).
Goes, O. C. et al., 815 (1-25).
Gomes, M. (et al.), 326 (2-la); 547, 593 (2-23); 614
(2-5); 632, 635 (2-23); 636 (2-13); 646, 655, 673 (2-
23); 676 (2-3a); 690, 759 (2-23); 786 (2-3a); 789,
810, 813 (2-23); 826 (2-13b);838 (2-23); 843 (2-3a);
844 (2-5); 872, 891, 924, 926, 931, 933, 955; 966,
979 (2-3a);1021 (2-2); 1033 (2-23); 1040 (2-5); 1048
(2-23); 1052, 1072, 1079, 1088 (2-3a); 1118, 1120
(2-23); 1123 (2-3a); 1129, 1156, 1175; 1176 (2-3a);
1196 (2-2); 1255, 1319, 1336, 1338, 1356 (2-23);
1357 (2-2); 1378, 1419, 1423, 1425, 1444, 1446 (2-
23); 1517 (2-13b); 5134 (2-23); 1540 (2-13b); 1534,
1568, 1570, 1577, 1584 (2-23); 1588 (2-13b); 1593
(2-25); 1594, 1595(2-23); 1598, 1604, 1610(2-13b);
1636, 1640, 1641 (2-23); 1679 (2-13b); 1692, 1731,
1745, 1769, 1822, 1823, 1867, 1893, 1917, 1983,
2170 (2-23); 2399 (2-3a).
Gomes, V. et al., 2818 (1-13); 2819 (1-17).
Gomez Pompa, A. et al., 3385 (2-3d).
Gonzalez,A. et al., 1319 (2-1b).
Graham,Mrs., s.n. (1-25).
Graimleux,-, 7938 (1-1); 7942 (1-4a).
Granville,J.-J.de (et al.), 581 (2-23);2744 (2-19);4531
(2-13e); B.5071 (2-la); B.5191 (1-20); 5284 (2-la);
5642 (2-23).
Grenand,P., 445 (2-la); 688 (2-13e);966 (1-4a); 1352
(2-la); 1442 (2-3c); 1478 (1-20); 1596 (1-4a).
Grubb,P. J. et al., 1545 (2-24); 1681 (2-13a).
Grubbe,J. P. et al., 1205, 1283 (1-14); 1498 (1-4b).
Guanchez,F., 239 (2-4).
Guedes, M. or T., 1235 (2-2).
Guillemin,J.-B. A., 1024 (2-la); 1339 (1-13).
Guppy, N., 656 (2-lOa).
Gutierrez,G. et al., 562 (1-31).
Haber,W. A., 1545 (1-34).
Hage, J. L., 31 (1-25).
Hahn, W., 347 (1-11).
Halle, F., 1132 (2-la).
Hamilton, B. et al., 1062 (1-18).
Hammel, B., 2176, 2576 (1-34); 4475, 6006 (1-6).
Hans, D., 335 (1-25).
Harling,G et al., 14733 (1-4b).
Haroldo, 57512 (2-13a).
Hartman,R. L., 12176 (2-3d).
Hartshorn,G. S., 1224, 1295(2-23); 1348 (2-3d); 1431
(2-23); 1436 (1-29); 1539 (2-3d); 2157 (1-34).
Hatch, W. R. et al., s.n. (1-30).
Hatschbach,G., 1475, 2458 (1-25); 7411, 7466 (2-la);
7807, 7833, 16304, 18680, 20253 (1-25); 25765,
29129 (2-la); 29629, 41730 (1-25).
Hayes, S., 32, 348 (2-3e); 354 (1-4b); 414, 867, 986,
1008, s.n. (1-42).
Hazlett, D., 664, s.n. (1-42).
Flora Neotropica
Hernandez,J. J. et al., 369 (1-9).
HerreraCh., G., 501, 503 (1-34); 505 (1-29).
Heyde, N. M. et al., 32, 39 (2-23); 78 (1-20).
Hilty, S., M-86 (2-23); M-87 (1-34).
Hine, R., P-1609 (2-23).
Hinton, G. B., 14018 (1-36).
Hohenkerk,L. S., 738 (1-4b).
Hoehne, F. C., 30923 (1-25).
Holdridge,L. R., 2514 (2-3d); 6254a (2-3e); 6818 (2-
23).
Holm-Nielsen, L. et al., 24519 (1-14); 24856 (1-10);
26331, 26610 (1-42).
Hoist, B. K. et al., 2421 (2-1 la); 2719 (2-la); 3195 (2-
3a).
Hostmann,F. W. R., 1189 (1-4a).
Hostmann, F. W. (& A. Kappler), 1272 (2-lOa and
2-1 la).
Howard,R. A. et al., 605 (1-42).
Hoyos, J., 2043 (1-42).
Huashikat,V., 431 (2-1 la); 912 (2-3b); 944 (2-lOb);
966 (2-13a); 1027 (2-10b); 1150(2-23); 1216 (2-la);
1241 (2-3a); 1432, 1482 (2-13a); 1585 (see 1-12);
1872 (2-3b).
Huber,J. E., 238 (2-la); 1640 (2-2), 1775 (1-21), 1873
(1-4a), 4244 (2-la).
Humbert,H. et al., 27221 (2-4).
Idrobo,J. M. et al., 786 (2-3a); 1316 (2-lOb).
Ijjasz-Madriz, -, 28 (2-1b).
Irwin,H. S. (et al.), 183 (1-26); 2058 (1-17); 4783 (2-
lla); 47874 (2-23); 48192 (2-3a); 48439 (2-1la);
48468 (2-23); 48631 (1-16); 48682, 48755 (2-3c);
55487 (1-20); 55576, 55884 (2-17); 57574 (1-1).
Izawa,K., 9 (2-la); 21 (2-4).
Janse, C. 0., 276 (2-3d).
Jaramillo,J. et al., 2088 (1-40); 4586, 31516 (2-4).
Jativa,C. (etal.), 303, 1079 (2-12b);1097 (1-10);2033,
2037 (2-12b).
Jenman,G. S., 652, 3995, 4947 (1-26); 4947A (1-4b);
5315 (1-26); 6310 (2-la); 7310, 7933 (1-26).
Jimenez, M. A., 433 (1-35); 657, 1750, 2887 (1-42);
3016 (1-18); 3816 (2-3d).
Jim6nezSaa, H., 15631 (=LBB 14296) (2-17).
Jobert, -, 682 (2-15).
Johnston,I. M., 580, 606, 608, 652, 678 (1-42); 1709
(1-4b); 1714 (2-3e), 1725, 1768 (1-42).
Jones, G. C. et al., 3152 (1-30).
Jones, J. et al., 9712, 9769 (1-28); 9788 (1-40).
Jonsson,G., 5A, 610a (1-25).
Juncosa,A., 1253 (2-12b).
Kanehira,R., 99 (1-42).
Kappler,A., 1272 (2-11a).
Karsten,H., s.n. (1-42, 2-la, 2-lb, 2-3a, 2-4).
Kayap,R., 201 (2-13a); 268, 393 (2-la); 571 (2-1 la);
1057 (2-la); 1306 (2-10b).
Kenoyer,L. A., 312 (2-3e).
Killip,E. P. (etal.), 25341, 26095 (1-32);27381,27932
(2-4), 28800 (2-unnamed);29246 (1-42); 29839 (2-
4); 29955 (2-23); 30139 (2-20); 30179 (1-38); 35095
(2-3e).
Kinloch,J. B., s.n. (1-30).
Klein, R. M., 89 (2-la); 79B, 772 (1-25); 1110 (2-la).
Klug,G., 1185, 1326 (2-4).
Exsiccatae 201

Korning,J. et al., 47618 (2-13c), 47641, 47645 (2-3a); Lindeman,J. C. (et al.), 1857 (1-25); 3631, 3689 (2-
47801, 47807 (2-unnamed). 10a);4057 (2-2); 5468 (2-3c); 5682 (1-25); 5709 (1-
Kosei Izawa (see also Izawa, K), 9 (2-la); 21 (2-4). 4a);5748 (1-25);6123,6166 (2-3c);6362 (1-1);6418
Krieger,P(adre)L. (as PLK)(et al.), 12268 (2-4); 12296 (2-2); 6445, 6600 (1-1); 6765 (2-3c); 6923 (2-lOa);
(1-42). 6938 (1-20); 9105, 9110 (1-25).
Krukoff,B. A., 1147, 1187 (1-39); 1197 (2-17); 1297 Lindeman,J. C. & A. C. de Roon, 747a (2-3c); 828a,
(2-la); 4518 (1-42); 4817 (2-la); 4994 (1-42); 5109 842b (2-3a).
(2-4); 5282 (2-23); 5309 (2-lOb); 5327, 5332 (2-4); Lindeman,J. C. & A. L. Stofferset al., 33 (1-29); 34
5596 (1-28); 5657 (1-32); 6041 (2-23); 6165 (1-42); (1-4a); 73 (1-20); 540 (2-17); 701 (2-13e); 749 (1-
6238 (2-la); 6524 (1-42);6662 (l-4b); 6887 (2-13b); 20); 790 (2-17); 799 (1-20).
7071 (2-23); 7073 (2-20); 7966 (1-4a); 8073 (2-23); Lisb6a, B., s.n. (1-13, 1-25).
8223 (2-13b); 8273 (1-35); 8325 (2-13c); 8332 (2- Lisb6a,P. L., 27 (2-4).
4); 8369 (2-5); 8223 (2-13b); 8273 (1-35); 8325 (2- Little,E. L. (et al.), 43 (1-27); 212, 213 (1-14); 424 (1-
13c); 8332 (2-4); 8369 (2-la); 8373 (2-6); 8386 (2- 32);602 (1-42);635 (1-11);6155 (1-42);6241, 6258,
la); 8388 (2-21); 8401 (1-28); 8406 (1-4c); 8427 6301 (2-3d); 6328, 6461 (1-42); 6640 (2-3d); 7756
(2-15); 8469 (2-4); 8518 (1-31); 8539 (1-35); 8587A (2-4); 8423 (2-la); 9527 (2-6); 9629 (1-40); 9641,
(1-31); 8652 (2-la); 8658 (1-35); 8695 (2-20); 8755 9789 (1-42); 15990 (1-4b); 16230, 16245, 21047 (1-
(1-35); 8807 (2-16); 8897 (1-35); 8967 (1-31); 8994 42); 21056 (1-41); 21070 (2-12b); 21136 (1-34);
(1-39); 10123 (1-4b); 10861 (2-13c); 10867, 10874 21172 (1-42); 21225 (2-12b).
(2-3a); 11062 (2-13c); 11085 (1-11); 11254 (2-3a); Lizot, J., 83 (1-40); 1972-1 (1-4b);s.n. ("Cl") (2-la).
11275 (1-9). Lleras,E. et al., P.17178, P.17197 (1-39); P.17242 (2-
Kuhlmann,J. G. (et al.), 255 (2-13b);263 (1-20); 291 9); P.17411 (2-8); P.17484 (1-4a); P.19586 (2-20);
(2-13b); 368 (1-42); 382 (2-2); 446 (2-10a);470 (1- P.19661 (1-20).
37); 759 (1-42); 1178 (1-28); 1337 (2-4); 1540 (1- Loureiro, A. (et al.), 16461 (2-13a); 16568, 16968,
32); 1832, 2073, 2074 (1-17); 2075 (2-la); 2177 (1- 50623 (2-20).
17); 2200 (2-la); 2207 (1-17); 5004 (1-25); 19843, Lowrie,S. R. et al., 710 (2-23).
150082 (2-7); s.n. (1-13). Lozano,C. G., 640 (1-42).
Kuhlmann,M. et al., 52 (2-2). Luetzelburg,Ph. von, 12471 (1-25); 23887, 23913 (2-
Lambroy, -, s.n. (1-4b). 4).
Lanjouw,J. et et al., 399 (2-1 la); 432 (2-13e); 1761 Lugo, M., 83 (1-4b); 2820, 2857, 2909, 3563 (1-27).
(2-2); 2106 (2-10a); 2113 (2-3c); 2114 (2-23). Lund, P. W., 139 (1-25).
LannaSobrinhoJ. de Paula et al., 376, 1011 (2-la). Lundell,C. L., 2, 648, 3170, 3535, 6224, 6350, 15930,
Lao, E. A. et al., 21 (1-42). 16122, 16162 (1-30).
Lao, M. R. et al., s.n. (2-4). Luschnath,B., s.n. (1-25); 40, s.n. (2-lOa).
Lasser,T., 2202, 2267 (2-lb). Maas, P. J. M. et al., 2478 (1-26); 2592 (2-14); 3325
Lawrance,A. E., 727 (2-3a); 769 (1-42); 810 (1-4b). (1-25);3622 (1-26), 3938 (2-1Oa);4126 (1-26);4679,
LBB (='s Lands Bosbeheer Suriname), 8000 (2-17); 4740, 4746, (1-19); 5858 (2-la); 6201, 6225 (2-4);
8161 (2-23); 8316 (2-3c); 8976 (2-1 la); 9199, 9424, 6249 (2-7);6267 (2-25); 11017 (2-17); 11027 (2-23).
9843 (2-10a), 11017(2-17); 11027 (2-23); 11747 (1- Macbride,J. F., 5446 (2-4); 5447 (1-4b); 5594 (1-32).
20); 14296 (2-17); 16031, 16310 (1-4a). Macedo, R., 55753 (2-17).
Leeuwenberg,A. J. M., 11784 (2-3a). Madison,M. T. et al., 4764, 7101 (1-19).
Lehmann,F. C., BT-694 (1-10); 948 (1-41); 5606 (1- Magalhaes,G. Mendes, 733 (2-la).
42). Magnago, - et al., 58046 (2-la).
Leitao Filho, H. P., 685, 686 (1-25). Maguire,B. (etal.),4945A (2-1la); 23449 (1-26);23867,
Leite, J. E., 413, 892 (1-25). 23942 (1-1); 24588 (1-26); 29408 (1-35); 41620 (1-
Leman,D. S. (herb.),s.n. (1-4a, 1-20). 40); 46879 (1-3); 47049 (1-1); 53517 (1-4a); 56511
Lemee, A. M. V., s.n. (1-4a); s.n. (2-3a, 2-3c). (2-1 la); 60333A (2-3a).
Lems, K., s.n. (1-25). Maia, L. A. et al., 497 (2-4).
Leng, H., 177 (1-26). Malme, G. O. A., 826, 1428 (1-25).
Lent, R. W. (et al.), 349 (1-42); 2004 (2-3d); 2327 (2- Marcano-Berti,L. (et al.), 471, 611 (2-la); 238-979 (2-
23); 2520, 2535 (1-42); 2538 (1-34); 3279, 3388 (1- 3d); 84-2-77 (1-4b); 982-154 (2-3d).
42); 3547 (2-3d); 5320 (1-42). Marinho,L. R., 273 (2-la); 348 (1-4b).
Le6n, J., 3501 (2-3d). Martin,J., s.n. (1-20, 2-la, 2-3c, 2-1 la).
Leprieur,F. R., 141 (2-lOa); s.n. (1-4a, 1-16, 1-21, Martinelli,G., 150 (1-25).
2-17). Martius,K. F. P. von, 2620 (2-2); 2673 (1-28); s.n. (1-
Lescure,J. P., 343 (2-1 la); 422 (2-23); 2075 (2-5). 28, 1-42, 2-la, 2-2, 2-3a, 2-4, 2-7, 2-13a, 2-15).
Liesner, R. L. (et al.), 417A (1-34); 515, 1309 (1-6), Mathews,S., 2060 (1-42).
4105 (2-13a); 8750 (1-31); 9443 (1-42); 10944 (2- Matuda,E., 446, 2020 (1-36); 3300 (1-30).
3d); 12394 (2-lb); 16175 (2-13b); 16299 (2-3a); McDaniel,S. (et al.), 2666 (1-31); 16092 (1-4b); 16344
22180 (1-11); 24462 (2-la). (1-40);20362 (2-15);20403 (2-2);20438 (2-7);20555
Lima, A., see Lima, D. de Andrade (1-40).
Lima, D. de Andrade(et al.), 20795 (1-25); 58-3185 McDonagh,J. F. et al., 114 (1-15).
(2-2); 67-4983 (2-la). McPherson,G., 7594 (1-6); 11510 (2-1 lb).
Lima, R., 21 (2-lOa). Medina, E., 379 (1-40).
 202
Melinon, E., 18 (1-20); 172 (1-11); 457 (2-11a); s.n.
(1-20, 2-1 la).
Mello Barreto,H., 1795 (1-13).
Mello, F., 55.388 (2-13a); 55.444 (2-1 la); 57509 (2-
la).
Mendez,R., 15 (1-6).
Meneces, E. (et al.), 338 (2-4); 613, 647, 725 (2-23);
759 (2-la).
Mennega,A. M. W., 271 (1-6); 521 (1-20).
Metcalf,R. D. et al., 30093 (1-42).
Mexia, Y., 1872 (1-36); 5115 (1-25); 6573 (1-42).
MeyerDrees, E., 5595 (2-3a).
Meyer,D. G., 403 (2-23).
Miers,J., 2714, 3792, 3858 (1-25); 6201 (2-la); 6257
(2-4); 8172 (2-la); s.n. (1-25).
MirandaBastos, A., 2051 (2-la); 2220 (2-23).
Miranda,F., 1737, 6164, 6273, 7180 (1-36).
MolinaR., A. et al., 1963 (2-3d); 17694 (1-34); 18227
(2-3d); 18289, 30747 (1-42).
Monsalve B., M., 1176 (1-46); 1259 (1-4b).
Montaldo,P., 3435 (1-42).
Monteiroda Costa, R. C., 90 (1-42); 92 (2-17).
Monteiro,O. P. (et al.), 309 (2-20); 477 (2-la); 1119
(2-23); 1295 (2-la).
Mora, L. E., 2295 (2-3e).
Morales, -, 38 (1-41).
Moreira,A. S., 46 (1-25).
Moreno,P. P., 23629 (1-18).
Moretti,C., 102 (2-la); 836 (1-20); 837 (1-4a).
Mori, S. A. (et al.), 5360 (1-34); 8583 (2-17); 8662 (2-
la); 8864 (1-26); 10408 (2-4); 11025 (2-lOa); 11026
(2-2); 11046 (2-la); 11655 (1-33); 14940 (2-19);
15127, 15342 (2-13e); 15350 (2-3c); 15776 (1-40);
16046 (2-la); 17185 (2-19); 17478 (2-1la); 17672
(2-13e).
Morillo, G. et al., 7384 (2-4).
Mosen, H., 2614, 2941 (1-25).
Mota, C. Domiao A. de (et al.), 154 (1-39); 680 (2-20);
740 (2-la); see also Domino, C.
Mota, M. G. et al., 1040 (2-5).
Mutis, J. C., 646, 649, 4558 (1-42).
Nadeaud,J., s.n. (1-25).
Nascimento, R. J. (et al.), 262 (2-2); 303, 387 (2-la);
66338 (2-3a); 66396 (2-7).
Nee, M. (et al.), 8822 (1-34); 9281 (1-6); 30877 (2-
13a).
Neill, D. (et al.), 1653, 1919, 1984, (1-42); 3389 (2-
23); 7044 (2-4); 7082 (2-la); 7179 (1-42); 8158 (2-
13a);8289 (1-22).
Nelson, B., 680, 681 (2-la); 705 (2-4).
Nevers, G. de (et al.), 4320 (1-6); 4697, 4789 (1-11);
4929 (2-1 lb); 4982 (1-15); 5062, 5502, 6152 (1-6);
6186 (1-18); 6194 (1-6); 6997 (2-23); 7170, 7542 (2-
3d).
Nishimura,A., 39 (2-1a).
Nufez, O. V., 2928 (1-42).
Nuiiez, P., 1823, 5761, 5946 (2-23); 5949 (2-la).
Occhioni, P. (et al.), 5864, 5917 (1-13); 8228, 35333
(1-25).
Oersted,A. S., 14317 (1-42).
Oldeman, R. A. A. (et al.), 207 (1-1); T.243 (1-4a);
T.243a (1-21); 366 (1-4a);B.644 (2-10a);B.1176 (1-
1); 1179 (2-la); 1243 (2-1 la); B.1324 (1-1); B.1338
(1-20); 1619 (1-1);B.1946 (1-21); 2030 (2-3c);2476
Flora Neotropica
(1-1);2504(2-23);B.2538(1-4a);2821 (2-23);B.2861,
3062 (1-21); 3115 (2-3c); B.3243 (1-1); B.3420 (1-
4a); B.3786A (2-2); B.4108, B.4132 (2-la).
Oldenburger,F. H. F. et al., 1120, 1406 (2-3c).
Oliva, F., s.n. (1-42).
Oliveira,E. de, 985 (1-20); 1974 (2-17); 2597 (2-10a);
2764 (1-35); 3071 (2-1Oa);3550, 3599 (1-20); 3627
(1-28);3682 (1-20); 3736 (2-1 la); 3915 (2-17); 3936
(2-13e); 3954 (1-20); 4047 (2-1 la); 60534 (2-5).
011gaard,B. et al., 35068 (1-49); 57061 (2-3a).
Ortega, -, s.n. (1-34).
Ortiz, R. T., 200, 1030, 1810 (1-30).
Palacios, W. (et al.), 1874 (2-24), 2059 (2-3a); 6387
(1-27).
Pariona,W. et al., 954 (2-13c).
Parker,C. S., s.n. (1-4b).
Peck, M. E., 497 (1-30).
Pennington,T. D. (et al.), 12244 (2-24), P.22647 (2-
7).
Pereira,A. B., 39 (1-25).
Pereira, L., 5670 (1-25).
Perrottet,G. S., s.n. (1-4a).
Persaud,A. C., 116 (1-4b); 222 (1-26).
Pessoal do Horto Florestal,650, 651 (2-la); s.n. (1-
25).
Peters,C., 187 (2-4).
Petilon, -, 154 (1-4a).
Philipson,W. R. et al., 1914 (1-42); 2086 (2-3a);2095
(1-42); 2133 (2-lOb);2134 (2-23).
Pinheiro,G. S., 1716 (1-25).
Pinsley, H. V., 405 (2-4); 556 (2-3A).
Pires, J. Murqa(et al.), 502, (2-39); 507, 542, 587 (2-
3a);805, 830 (1-4b);6663 (2-la); 7015 (2-2); 10123,
10758, 10794, 10809 (2-lOa); 12570, 15088 (2-la);
16461 (1-42); 16547 (2-1a);50419 (2-23); 50631 (1-
1); 50656 (1-2); 50864 (1-20); 50913 (1-4a); 51193
(2-13e); 51247 (2-23); 51390 (2-17); 51409 (1-20);
51624 (2-3c); 51625 (2-13e); 51676 (2-10a); 51736
(1-40); 51882 (2-2); 52656 (1-20).
Pires, O. et al., 248 (1-42).
Pittier, H. (et al.), 2286 (2-3e); 3892 (1-42); 4386 (1-
6); 8404 (1-42); 11166 (1-34); 12105 (2-3d); 12124,
12142, 12144, 12288, 13985 (1-42); 15641 (2-1b);
15733 (1-42); 16163 (2-3d); s.n. (1-42).
Plowman,T. C. (etal.), 2362 (1-42); 2919 (1-25);6555
(1-28); 6903 (2-4); 6906 (2-5); 11179 (1-32); 11180
(1-42).
Poeppig,E., 1881 (2-la); s.n. (1-28, 1-42, 2-la, 2-3c).
Pohl, J. E., s.n. (1-25, 1-28).
Poiteau,P. A., s.n. (1-20, 2-3c, 2-3a, 2-1 la, 2-3a, 2-3c,
2-1 la).
Poveda, L. J., 951 (1-34).
Prance,G. T. (et al.), 1379 (2-3a); 1548 (2-2); 1720 (1-
20); 1777 (2-la); 2153 (1-4c);2180 (2-1 la); 2306 (1-
42); 2469 (2-2); 2754 (2-3a);3253 (2-7); 3260, 4438
(2-la); 4588 (1-4b);6253 (2-la); 6292 (2-13b);6419
(2-4); 6718 (1-4b); 6969 (1-31); 7160 (2-17); 7371
(1-32); 7503 (2-4); 7530 (1-42); 7670 (2-la); 7792
(2-23);7907 (2-lOb),8378 (2-23);8670 (2-la); 9054
(1-35);9679(1-11);9857, 9952 (2-13b);10089,10124
(2-23); 10231 (2-5), 10233 (2-la); 10348 (1-42);
10836 (2-la); 11066, 13438 (1-42); 13579 (2-la);
13582-A (2-3a); 14029, 14146 (1-31); 14795 (2-7);
14976 (1-38); 15039 (1-34); 15366 (2-3a); 15393 (2-
Exsiccatae
20); 15764 (1-40); 15791 (2-4); 17242 (2-9); 17847
(1-38); 18032 (1-31); 18204 (2-23), 18240 (2-5);
18250 (2-3a); 19878 (2-13b); 20454 (2-20), 21011
(2-2); 22708 (2-13b); 22720 (1-20); 22729 (1-31);
22740 (1-20); 22803 (2-la); 22836 (1-20); 22859 (2-
16);22864 (2-20); 22999 (1-35); 23776 (2-7);23846
(1-39);23966 (2-9);24188 (2-3a);24313 (2-7);24687
(1-42);25167, 25336 (2-2);25337 (2-10a);25363 (2-
2); 25530 (2-20); 25748 (1-35);26375 (2-la); 26402
(1-42); 58783 (1-20); s.n. (2-5).
Pr6vost,M. F. et al., 738 (1-4a); 982, 983 (2-3c).
Proctor,G. R., 27080 (1-18); 27336 (1-42); 30273 (1-
30).
Pruski,J. et al., 3235 (2-13b); 3239 (2-20).
Purdie,W., s.n. (1-14).
Purpus,C. A., 5996, 11161, 11162, 11182, s.n. (1-36).
Rambo, B., 426, 27092, 37782, 38427, 39413, 39780,
41173, 44743, 45136, 45904, 46145 (1-25).
Ramirez,C. R., 1084 (1-31).
Ramos, J. et al., 204 (2-20).
Rauh, W., P-1609 (2-23).
Reineck,E. M. et al., 446 (1-25).
Reitz, R. (et al.), 975, 1688, 1912, 2008, 2027, 2913,
3809, 4176, 4692, 4965, 5030, 5611, 8294 (1-25).
Renteria,E. et al., 45, 2093 (2-12a).
Revilla, J., 805, 1099 (1-28); 1260 (1-4b); 1818 (2-5);
1847 (1-31); 1966 (2-7); 2387 (1-39); 3387 (1-28).
Reyno, R. N., 37 (2-13a).
Ribeiro, B. G. S., 612 (1-11); 665 (1-43).
RichardL. C. (herb.).,s.n. (1-20, 2-la).
Riedel, L. (et al.), 1 (1-25); 2 (2-la and 2-lOa); 7 (1-
25); 37, 46 (2-la); 75 (1-25); 160 (2-la); 664; s.n.
(1-25, 2-la).
Riera, B. J. J.-L., 206 (2-3c); 301, 571 (2-17); 573 (2-
10a);714 (2-la); 723 (2-13e); 923 (2-la).
Rimachi,Y. M., 359 (1-32); 1943 (1-28); 2297 (1-40);
2724 (2-5); 2725 (2-9); 2765 (2-5); 3256 (2-15).
Rizzini, C. T., 415, 1114 (1-25).
Roa T., A., 251 (2-unnamed);252 (2-13a).
Rodrigues,C. R., 944 (2-5).
Rodrigues,G. et al., 877 (1-17).
Rodrigues,W. A. (et al.), 884 (2-23); 1415 (2-7); 1424
(2-23); 1434 (2-13b);2092 (2-2) = 2892 (2-2); 2984
(2-1Oa);2986, 2987 (2-13e);3267 (2-2);4237 (2-3a);
4585 (2-2); 5266 (1-31); 5449 (2-20); 7069, 7076 (2-
7);7081,7292 (2-20);7532 (2-23);8065 (2-13);8066
(2-5); 9204 (2-la); 9206 (2-3a); 9219 (2-la); 10082
(2-17); 10182 (2-23); IG1-2A-70 (2-5).
Rodriguez,H., 399 (1-42).
Rodriguez,J. V., 2929 (1-42).
Rohr, J. B. von, 151 (1-4a).
Rombouts, H. E., 803 (1-4a).
Romero, F., 548 (1-42).
Romero-Castaieda,R., 3647 (2-4); 3840 (2-5); 3848
(2-20);4211 (1-4b);4715 (2-12a);5283 (1-19); 5484
(2-3e); 5512 (1-10).
Rooden, J. van et al., 258 (1-4b); 699 (1-10); 700 (1-
5).
Rosa, N. A. (et al.), 296 (1-11); 339 (1-43); 1091 (2-
5); 1126 (2-17); 1158 (2-lOa); 1159 (2-5); 1265 (2-
23); 2053, 2874 (1-40).
Rowlee, W. W. et al., 856 (1-42).
Ruiz, H. & J. Pavon, 2 (1-32); 3 (1-42); 4 (1-4b); 5, 6,
7 (1-32); s.n. (1-4b, 1-32, 1-42, 2-lOb).
203
Ruiz-Teran,L., 423 (1-42).
Rusby, H. H., 1599 (2-4).
Rylands,A. B., 15 (2-2); 25 (2-23); 30 (2-13b); 52 (2-
3a); 59 (2-2).
Sabatier,D. R. L., 501 (2-1 la).
Sagot,P. A., 517, 970, 972 (2-17);990, 990bis (2-1 la);
1136 (1-20); 1163 (2-2); s.n. (1-4a, 1-20, 2-17).
Samaniego,A. et al., 92 (1-42).
Sandwith,N. Y., 290, 302, 394 (1-26).
Santos,M. R., 172 (2-23); 455 (2-lOa).
Santos,T. S. dos, 2035 (2-la).
Sastre,C., 2274 (2-4); 4350, 4579 (2-3c); 5500 (1-1);
6130 (1-20); 6417 (1-4a).
Sawada,M., 21 (2-1Ob).
Schipp,W. A., 127 (2-3d); 999 (1-30).
Schnee,L., 1337 (1-42).
Schomburgk,R., 118, 167, 287, 876, 1366, s.n. (1-26).
Schubert,B. G. et al., 748 (2-3d).
Schultes,R. E. (et al.), 3343, 3826, 3862, 3901 (2-4);
6693 (1-8); 6726 (1-28); 6805, 8175 (2-4); 8269 (1-
8); 8380 (1-28); 8976 (1-4b); 9685, 12985 (1-31);
13589(1-40); 13774(1-31); 14148(2-23); 15324(1-
31); 15973(2-5); 16009(2-20); 16112(1-31); 16334,
16367 (1-42); 17615 (2-3a); 18969, 19758 (1-4b);
19835 (2-5); 24547 (2-4).
Schultze-Rhonhof,H., 1953 (1-19); 2783 (1-11); 2943
(1-9); 3006 (1-11); 3020 (1-14).
Schulz,J. P. (etal.), 7787 (1-20); 8000 (2-17); 8161 (2-
23); 8316 (2-3c); 8976 (2-1 la); 10344a (1-1).
Schunke,C., 416, A-95 (2-23).
Schunke,J. M., 130 (1-28); 371 (1-40).
SchunkeV., J., 1162 (1-32); 2139 (2-23); 2824 (2-4);
3309, 3552 (2-la); 6485 (2-4); 7415 (2-la); 7729 (2-
10b);8487 (2-23); 8666 (1-4b); 10650 (2-lOb).
Schwacke,W., 560 (2-15); 1072, 6783, 8418 (1-25);
10393 (2-la); 11895 (1-17).
SEF(=Studiesof EcuadorianForest),8532 (2-6);8617
(2-23); 8764 (2-4); 8823 (2-13c); 8930 (2-4); 9033,
9159 (2-23); 10366, 10378 (2-spec.).
Sehnem,A., 7993 (1-25).
Sello(w),F., 241 (1-25); s.n. (2-2).
Shakaim,S., RBAE-26(2-23).
Shank,P. J. et al., 4294 (1-42).
Shattuck,O. E., 260, 522, 525 (2-3e); 832 (1-42).
Shepherd,D., 364 (2-23); 376, 379 (2-3a).
Shima, D., 14 (2-20).
Sigueira,R., 8808 (1-20).
Silva, A. S. Lima de et al., 215 (2-13d);476 (2-8).
Silva, J. F., 69, 211 (1-20); 227 (1-4b).
Silva, M., 354 (2-lOa).
Silva, M. Barbosada, 130 (2-lOa).
Silva, M. F. et al., 466 (1-31); 742 (2-20); 869 (2-lOa);
903 (2-23); 2166 (1-4a); 2199 (2-13b); 2423 (2-2);
2591 (1-39); 2623 (2-la); 2712 (1-4a).
Silva, M. G. et al., 3345 (1-40); 3368 (1-4b); 4326,
4738 (2-23); 4750 (2-la).
Silva, N. T. (et al.), 921, 949, 995 (2-1 la); 1207, 1321,
1380 (1-20); 1411, 1412 (2-3c); 3013 (1-28); 4757
(1-10); 60670 (1-43).
Skutch,A. F., 2023 (1-36); 4083 (2-3a); 4225 (2-3d);
4256, 4262, 4267, 4612 (1-42); 4740 (2-20); 57832,
58862 (2-3a).
Smith, A., 1632 (1-29); 1663, 1650 (1-34).
204
Smith, A. C., 2731 (2-13d); 2753, 2799 (1-35); 2845
(2-1a); 3564 (2-3c).
Smith, D. N. (et al.), 1090 (1-32); 2089 (2-4); 4743 (2-
unnamed);5134 (2-10b); 5300 (2-4).
Smith, L. B. et al., 12307 (1-25).
Smith, R. F., 3221 (2-lb); 3313 (1-42).
Smith, S. F. et al., 695 (2-3a).
Snedaker,S. C., C-39, D-88 (1-30).
Sneidern,K. von., s.n. (2-4).
Sodiro, A., 193/12 (1-41).
Soejarto,D. D., 327, 856 (1-42); 2917 (2-23); 3385 (2-
3e); 3489, 4063 (2-23); 4283 (2-1 lb).
Solomon, J. C. (et al.), 9254 (2-23); 12655 (2-la).
Soria S., M. A., 21 (2-23).
Sparre,B., 14822 (1-42).
Sperling,C. R. et al., 6430 (2-23).
Spichiger,G. et al., 1995, 1996, 1997, 1998, 1999 (2-
25).
Splitgerber,F. L., 448 (1-4a).
Spruce,R., 951, 1219 (2-13a); 1509 (1-4a); 1608 (1-
42); 2023 (2-4); 2260 (1-4b); 2498 (1-31); 2616 (1-
40); 2865 (2-13b);3176 (2-1 la); 3231, 3464 (1-40);
3782 (1-38); s.n. (2-13a, 2-17).
Stahel,G.et al., 732 (1-20).
Stahel,G., (Exp.WilhelminaGebergte)196 (1-20).
Stahel,G., (Woodherb.Suriname)82 (2-23); 123A (2-
3c); 123B (2-17); 166 (2-3a); 166A (2-17); 166B (2-
3c).
Standen, -, 20 (1-42).
Standley,P. C. (et al.), 7081 (1-42); 7878 (2-3d);8861,
12288 (1-42); 19904 (2-3d); 23766 (1-42); 27479,
27502 (2-3e);30924 (1-42); 37121 (2-23); 37259 (1-
42); 39792 (1-34); 41170, 44550, 44968, 45182,
45661, 47791, 48594 (1-42); 48600 (2-23); 52900
(2-3d); 53230 (1-42); 53951 (2-3d); 53991, 53425,
55155, 55445, 56623, 72288 (1-42); 72667 (2-3d).
Steinbach,J., 1484 (1-32); 7567 (1-42).
Stergios,B. et al., 11530 (1-35).
Ster, W. L. et al., 423 (2-3d).
Stevens,W. D., 4839, 8323 (2-3d);13340(2-23); 13341
(2-3d).
Steward,W. C. et al., 107, P.20156 (2-20); P.20255
(1-4a).
Steyermark,J. A. (et al.), 388, 487 (1-3); 507 (1-4b);
37602 (1-36); 38916 (2-3d);39460 (1-30); 39618 (1-
42); 39934, 45663, 45981, 46103 (1-30); 52334 (1-
36);54214(1-10);54533 (1-42);57872(2-13a);57955
(1-43); 60002 (1-4b); 60002a, 60364 (1-11); 60399
(2-23); 60401 (2-3a); 60407 (2-1a); 60414 (2-23);
60432 (1-4b); 60456 (1-43); 60665 (2-2); 60673 (1-
11);60800 (2-23);75522 (1-11);75540 (2-3a);87307
(2-la); 87480 (1-4b); 89134 (2-la); 89452 (2-1 la);
89457 (2-3a);89506 (2-23); 90368 (2-20);90408 (2-
1la); 90583 (1-4b);90620 (2-23);90644 (2-3a);90736
(l-4b); 92850, 92936 (2-22); 92991 (1-3); 94180 (2-
3a);95170(1-42); 95378 (2-lb); 95711 (2-la); 99829
(1-42);99945 (2-3d);102082, 102211(1-42); 102874
(2-unnamed);103026(2-13a);104296(1-11);104348
(1-4b); 104466 (2-23), 105863 (2-lb); 106086 (2-
la); 106118 (1-11); 106167 (1-31); 107064 (2-23);
113096(2-3a);114759(1-4b);115549(1-11);116463
(1-42); 116884 (1-4b); 116966 (2-1b); 119396 (2-
Flora Neotropica
3d); 122390, 125857 (2-3a); 129163 (2-4); 129733
(2-unnamed);131962 (2-1 la).
Stoffers,A. L. et al., 244 (2-17); 265, 311, 312 (2-la);
319 (2-17).
Sucre,D., 1783, 3633, 4989, 5209, 5330, 7878 (1-25);
8303 (2-lOa);8684 (2-2); 8690 (1-25).
Sugden,A., 415 (1-42).
Sytsma,K. J., 4018 (2-3d); 4024, 4225 (1-6).
Tamayo, F., 3370 (2-lb).
Teixeira,L. O. A. (et al.), 61 (2-3a); 849 (1-4B).
Terceros,W., 1400 (2-la).
Tessmann,G., 3054 (2-4); 3416 (1-42); 3922 (1-32);
4236 (2-3b);4642 (2-la); 4673 (1-39); 4696 (1-42);
s.n. (1-25); 5364 (2-9).
Teunissen,P., 16031 (1-4a).
Thomas, W. W., 3349 (2-13b); 3370 (2-3a).
Tillett,S. S. (et al.), 45869 (1-43);671-33 (1-3 1);49495
(2-1la).
Tomas, Bro., 1923 (1-42).
Tonduz, A., 9520 (1-42); 12930 (2-3d); 13280 (1-42).
Toro, R. A., 120C (1-14).
Torres,M. J., 482 (1-32).
Toth, M. J., 482 (1-32).
Tresling,J., 55 (1-1).
Triana,J. J., 860 (2-la); 861 (2-3a);862 (2-4); 866 (1-
34); 867 (1-42); 1846 (2-4); 1866 (1-34, 1-42); s.n.
(1-42).
Troon, F. van et al., 16310 (2-23); s.n. (2-la).
Trucios,T., 9 (2-1Ob).
Trujillo,B., 6141, 8765 (1-42).
Tuerkheim,H. von, 4082 (2-3d); 8659 (1-30).
Tunqui, S., 7, 94 (2-3b); 103 (1-42); 149 (2-3a); 217
(2-13a); 296 (2-10b).
Tweedie,J., 29 (1-25).
Tyson, E. L. et al., 4734 (2-3d).
Ule, E. H. G., 991 (2-la); 1693 (1-25); 5265, 5266 (2-
4); 5717 (1-42); 5718 (2-23); 5719 (2-6); 8836 (1-
4a); 8839 (2-17); 9314 (2-4); 9315, 9316 (1-32); s.n.
(1-25).
Uribe-Uribe,L., 364 (1-42).
VasquezA., R. et al., 17 (2-23); 167 (2-25); 809 (2-
13c); 2396 (2-la); 2491 (2-23); 2553, 2559 (1-42);
2623 (2-25); 2630 (2-7); 2635 (2-5); 2539 (1-39);
2719 (2-1 lb); 2852 (1-22); 2861 (2-23); 3965, 4201
(2-7); 5111 (2-9); 5433 (1-31); 5745 (2-la); 5853 (2-
9);6110 (2-23);6156 (2-20);6942 (1-42);7104, 7522
(1-31); 7535 (2-25); 7669 (1-4b);8380, 8600 (1-22).
Veloso, H., 97 (1-25).
Vellozo, -, 723 (2-la).
Vellozo, H. P., 89, 723 (2-la); 1083 (2-lOa); 737 (1-
33).
Vianna,E. C. et al., 5465 (1-25).
Vickers,W. T., 86 (2-5).
Vieira,M. G. de et al., 985 (2-23); 1004 (2-20).
Villiers,J. F., 2088, 2111 (2-23).
Vogl, C., 1379, s.n. (1-42).
Vreden,J., 11717(1-20).
Wachenheim,H., 18 (2-23); 232 (2-1 la); 271 (2-2);
392 (2-13e);426 (1-4a);467 (2-23); s.n. (1-20, 2-la,
2-23).
Warming,J. E. B., 1942, s.n. (1-25).
Weaver,R. et al., 1677 (1-6).
Exsiccatae 205

Weberbauer,A., 3639 (2-la); 3702 (1-42); 4472 (1-
44b).
Wedel, H. von, 1548, 1733, 2879 (1-42).
Westra,L. Y. T., 47303 (2-2).
Whitton, B. A., 179 (1-25).
Williams, LI. (et al.), 3347, 3984 (2-4); 4179 (1-42);
4627 (2-4); 4688 (2-lOb);5349 (1-42); 9983 (2-lb);
10071 (1-42); 12008 (2-lla); 12348, 12394, 13895
(1-42); 14675 (1-4b); 14833 (2-13a); 15240, 15363
(1-31); 15812 (2-5).
Williams,L. O. et al., 17817, 18014, 24005, 26529 (1-
42).
Williams,R. S., 501 (2-lb); 983 (1-3d); 1560 (2-la).
Wilson-Browne,G., 549 (2-23).
Wilson, C. L., 127 (1-42); 381 (1-30).
Wilson, P., 606 (2-3d).
Woodson, R. E. et al., 1893 (1-42).
Woodworth,R. H. et al., 445 (1-42); 606 (1-4b); 660
(1-42).
Woytkowski,F., 5926 (1-42); 7306 (1-32).
Wurdack,J. J. (et al.), 286 (1-4b); 2169 (2-23); 43485
(1-40).
Yele, -, DR-23 (1-26b).
Zarucchi,J. L., 2145 (2-4); 2744 (2-la); 3720 (2-3a);
3272 (2-3e).
Zikan,J. F., s.n. (1-25).
Zuccarini,J. G. (herb.), 103 (1-25).

INDEX OF VERNACULAR NAMES OF POUROUMA

amapati(la) 127 cucuva (4) 147
ama'yrary(1Oa)156 dacha uvillos (5) 149
ambafba do vinho (13a) 168 dakamtazshiuya(1 la) 161
ambauvamansa (4) 147 embaubada mata (la) 127
ambauvado vinho (4, 13a) 147, 168 embaubarana(7) 152
a'ilea (3c) 138 garguaba(3a) 135
ambaibochi(23) 189 granboesipapaja (10a, la, 13e, 17, 23) 156, 161, 173,
ambaibillo(23) 189 180, 189
amia-yek(la) 127 guagay(3d) 141
bochoa tsaha (4) 147 guarumocolorado(5) 149
boesipapaja(13e) 173 guarumode montafia(3d) 141
bois canon (la, 2, 3c, la, 13e, 17, 19, 23) 127, 132, guarumomacho (3d) 141
138, 161, 173, 180, 181, 189 guaumoutognac(13a) 168
bois canon male (10a, 23) 156, 189 gurucana(4) 147
bois canon sauvage(10a) 156 hembra(3a) 135
boroma (2, 3a, 3c, 10a, Ila, 13e, 17) 132, 135, 138, imbafba (4, 5, 10a) 147, 149, 156
156, 161, 173, 180 imbaubabengue(17) 180
boroma ibeberobana(2) 132 imbaubabranca(17) 180
bospapaja(2,3c, 10a, 1la, 23) 132, 138, 156, 161,189 imbaubade cheirouvilha(17) 180
bouchi papaie (23) 189 imbaubado vinho (4, 13a) 147, 168
bouchi papaye(13e, 17, 23) 173, 180, 189 imbaiba mansa (4) 147
buruma(la, 3c) 127, 138 imbaibarana (la, 2, 3a, 5, 7, lla, 13b, 13c, 20, 23)
caimar6n(4) 147 127, 132, 135, 149, 152, 161, 169, 180, 189
caimar6nde mico (3a) 135 imbafbaranabranca(5) 149
caimar6nsilvestre(4) 147 imbaubaranafolha peluda(5) 149
caramuri(23) 189 imbaubaranavermelha(10a) 156
cay-bari-cay(3a) 135 imbauba-tor6m(la) 127
cay-wari-cay-yek(3a, 22, 23) 135, 186, 189 imbaubinha(8) 152
chaparrode agua (la) 127 itararanga(la, 10a) 127, 156
chiricaba(13a) 168 kaibarikei(5) 149
chullachaqui(20) 184 kai-wa-rei-kei-yek(23) 189
chullachaquiblanco (20) 184 kaiwarikai(1 a) 161
chullachaquicaspi (20) 184 kalate(la) 127
chumico (3d) 141 kamoyuwa(10a) 156
cirpe (3a) 135 kaymbe'y(la) 127
cirpe macho (3a, 1 b) 135, 162 kukuma(13e) 173
cocora(13a) 168 kuluma(la) 127
cocura(4) 147 kulumasi(23) 189
coiwaricoi-yek(23) 189 kulumatelelel(3c) 138
cormi (3a) 135 lija (3d) 141
cucura(2, 3a, 4, 5) 132, 135, 147, 149 majagua(23) 189
cucure(3a) 135 male bois canon (3c, 23) 138, 189
206 Flora Neotropica

manbospapaja(la, 17) 127, 180 sirpe macho (3a) 135

mangab6(3e) 144 sirpo (3e, 12a) 144, 162
mano de leon (3d) 141 sucufba (4) 147
mapati(la, 4, 8, 13c, 15, 21) 127, 147, 152, 169, 177, sugkama(t)shuiya (la) 127
186 suia(4, lOb) 147, 159
mapatirana(2, 3a, 3c, 10a, la, 16, 23) 132, 135, 138, suir shuina(lOb) 159
156, 161, 177, 189 tamaoquare(3a) 135
mapaty(13c, 15, 21) 169, 177, 186 tambor(lb) 129
mimpa shuiya washi shuina ( la) 161 tanaribe(4) 147
obija (1Ob)159 tanta shuiya (3b) 137
orumo de monte (3d) 141 tararanga(lOa) 156
otsepacho(3a) 135 tararangablanca(la) 127
pacica (3d) 141 tararangavermelha(lOa) 156
papaie (17) 180 tentarshuina (3b) 137
papaquillo(la) 127 tinajero(3d) 141
papayadel monte (la) 127 torena(23) 189
papaye(17) 180 tourem (23) 189
papayeapici (2, 1la) 132, 161 trumpettree (3d) 141
pau de jacu (la) 127 tsaha (3a) 135
pai shuina (3a, 13a) 135, 168 tsakapsuiya (la, 3a) 127, 135
pai shuiya (13a) 168 uhukamsuiya(10b) 159
piraejo(3d) 141 umbaouba(5) 149
pourouma(3a, 3c, lla, 17) 135, 138, 161, 180 urumoblanco (12a) 162
puruma(3a, 3c, lla, 17) 135, 138, 161, 180 uva (3d, 4, 12b) 141, 147, 163
puruma(4) 147 uva de macaco (lOa) 156
purumai(13b, 23) 169, 189 uva de(l) monte (3d, 4, 13c) 141, 147, 169
puruma-y(4) 147 uva silvestre(4, 23) 147, 189
sachauvilla (4, 9, 13a, 15, 23) 147, 155, 168, 177, 189 uva medueda(3a) 135
sacha uvillos (20) 184 uvilla(la,4, 5, 9, 10b, 1a, 15,23) 127, 147, 149, 155,
sadajii(4) 147 159, 161, 177, 189
sadha'fhi(3a) 135 uvilla blanca(la, 23) 127, 189
sandpaper(la) 127 uvilla lanuda(23) 189
sa-ouro(10a) 156 uvo (4) 147
sarasara(3a, 5, 13b) 135, 149, 169 wilaupiyua(13e) 173
shewantoqui(4) 147 yagrumonegro (lb) 129
shuiya (la, 3b) 127, 137 yagrumo-sunsun(10a) 156
shuvija(23) 189 yahal (3d) 141
sirpe (3a, 12a) 135, 162 yaryara(2, 3c, lOa, lla, 17) 132, 138, 156, 161, 180

INDEX OF SCIENTIFIC NAMES

Synonymsarein italics.Pagenumbersin boldfaceindicateprimarypagereferences.Pagenumbers
with an asterisk(*) indicate pages with illustrationsor maps.
Allomeris 114 argentea10, 11*, 30, 32*, 73
Azteca 4 asperifolia4, 5, 7, 8*, 10, 11*, 15, 16, 33, 34 (key),
Brosimum66 99, 104
microcarpon66 subsp. asperifolia10, 11*, 34, 35*, 63
Cebusapella 189 subsp. magnifolia 11*, 36, 37*, 39, 99
Cecropia2, 3, 9, 10, 15, 113, 115 subsp. rhamnoides11*, 38, 39*
peltata3 batavorum11*, 38, 40*, 99
Cecropiaceae2, 3, 5, 15, 110, 113 boliviana99
Clusia 4 brenesii86
Coussapoa2, 3, 4, 5, 6*, 7*, 8*, 9, 10, 15, 16, 17-28 brevipes7, 11*, 41, 42*
(keys), 112, 113, 114, 115, 194 cardonaei36
acutifolia84 cayennensis34
angustifolia11*, 28, 29*, 58, 97 chagresiana34
apoda 53 chocoensis 11*, 41, 43*
arachnoidea10, 11*, 30, 31*, 50, 77, 82 cinnamomea 11*, 44, 45*, 95
araneosa99 cinnamomifolia5, 10, 11*, 15, 16, 44, 46*, 86, 97
Index of ScientificNames 207
contorta5, 7, 11*, 41, 44, 47*, 48, 66 planitiensis 101
cornifolia87 plicata 110
crassivenosa10, 11*, 15, 48, 49*, 56 prancei80, 82
cuneata 71 puberula84
cupularis11*, 50, 51*, 77, 82 purpusii10, 13*, 16, 90, 91*
curranii4, 12*, 50, 52*, 58 rekoi 110
danielis 99 rhamnoides38
dealbata 109 rotunda97
dolichandra109 ruizii 36
donnell-smithii99 scabra 13*, 90, 92*
duquei 7, 10, 12*, 53, 54*, 77, 104 schottii66
echinata 12*, 53, 55*, 61, 108 var. lanceolata66
eggersii99, 104, 105 var. longifolia66
emarginata109, 186 schunkei75
embirana99 setosa 84
fagifolia 71 sprucei 13*, 15, 63, 66, 80, 92, 93*
ferruginea12*, 15, 56* standleyi99
ficina 34 steyermarkii48
floccosa 12*, 52, 57* subcrenata99
fontanesiana 66 subincana99
froesii 61 tessmannii 14*, 15, 44, 77, 93, 94*, 104
fulvescens 14*, 105, 107*, 108 tolimensis 14*, 108, 109*
glaberrima12*, 58, 59* trinervia4, 5, 10, 14*, 16, 44, 59, 95, 96*, 97
grandiceps99 valaria108, 110*
herthae12*, 56, 58, 60*, 61, 108 vanniifolia5, 14*, 39, 97, 98*
hirsuta84 vellerea99
hypochlora36 villosa 4, 5, 8*, 10, 14*, 15, 53, 77, 95, 99, 100*,
incomitata50 110
intermedia75 viridifolia 7, 10, 14*, 15, 16, 63, 82, 105, 106*
krukovii110, 168 var. tenuifolia105
laevigata 110 volaria 14*
latifolia9*, 10, 12*, 15, 16, 61, 62*, 63, 71, 73, 82, warburgiana50
86, 105 williamsii80
var. obovata61 Crematogaster114
lawrancei99 Ficus 4, 110
lehmannii99 intermarginalis110
leopoldii 110 Moraceae2, 3, 5, 15, 110
leprieurii12*, 15, 63, 64*, 66, 93 Musanga2, 9, 15, 113
longepedunculata13*, 63, 65*, 77, 93 Myrianthus2, 5, 9, 15, 112, 113, 115
macarenensis101 Poikilospermum2, 3, 15, 113
var. antioquiensis101 subgen.Ligulistigma15
macerrima12*, 63, 66, 67*, 73, 92 Poulsenia 110
magnifolia36 armata 110
manuensis 13*, 66, 68* Pourouma2, 3, 5, 9, 10, 15, 110, 111, 113, 114, 116,
martiana99 117-121 (key), 194
microcarpa5, 7, 12*, 15, 63, 66, 69*, 73, 90 acuminata113, 114, 116, 173, 175*, 176*, 193
microcephala12*, 33, 63, 66, 71, 82, 90 acutiflora123
subsp. corifolia 89 albistipulata166
subsp. microcephala72* apaporiensis160, 161
mutisii 101 forma macrophylla160
napoensis 12*, 73, 74*, 84 apiculataBenoist 168
nitida 5, 10, 13*, 50, 75, 76*, 77, 82, 108 apiculataMildbraed168
nymphaeifolia13*, 15, 61, 66, 77, 78*, 93 aspera 116, 133
obovata61, 110 subsp.digitata 115
oligoneura86 aurea 186, 189
oligocephala13*, 48, 77, 79*, 80, 93 bicolor 111, 112, 114, 115, 122, 123, 132, 132-133
orthoneura10, 13*, 30, 50, 63, 77, 80, 81*, 82 (key)
ovalifolia 13*, 82, 83*, 84, 108 subsp.bicolor 123,133, 134*, 143*, 149, 193, 194
pachyphylla13*, 63, 73, 84, 85* subsp.chocoana 115, 141, 142*, 143*, 181
panamensis99 subsp. digitata 115, 137, 138*, 139, 143*
parviceps7, 13*, 16, 44, 56, 86, 87*, 90, 108 subsp.scobina115, 122, 123,139, 140*, 141, 143*,
parvifolia5, 12*, 73, 87, 88*, 89*, 92 144
pittieri 110 subsp. tessmannii 115, 136*, 137, 143*
208 Flora Neotropica

bolivarensis 111, 113, 114, 115, 116, 176*, 186, subsp. mollis 115, 156, 157*, 158*, 159, 160
187*, 189 subsp. triloba 114, 115, 156, 157*, 193
camaratana133 multifida144
cecropiifolia110, 111, 113, 114, 116,139, 143*,144, myrmecophila111, 112, 113, 114, 115, 116, 150,
145*, 147, 194 151*, 152, 157*
chocoana 141 napoensis 113, 114, 115, 116, 171*, 190, 191*
cinerascens123 oraria 111, 112, 114, 115, 176*, 180, 190
crassivenia139 ovata 111, 112, 113, 114, 115, 116, 176*, 177,181,
crassivenosa133 183*
cuatrecasasii186 palmata 123
cucura 112, 114, 115, 116, 147, 148*, 157*, 193, paraensis194
194 phaeotricha1 111,113, 115,116,152,154*, 155,176*
cuspidata 113, 114, 116, 149, 157* populifolia173
digitata 137 radula 123
edulis 144 retusa 194
elliptica 114, 176*, 184, 185* sapida 144, 147
essequiboensis169 saulensis 114, 115, 116, 176, 181, 182*, 184
ferruginea111, 113, 114, 116, 176*,177, 178*, 184, scabra 123
193 scobina 139
folleata 186 schultesii133
formicarum112, 113, 114,115,116,152,153*, 155, steyermarkii130
157* stipulacea111, 112, 114, 115, 116, 173, 174*, 176*
fulginea 123 subplicata186
garciana 147 substrigosa123
guianensis 111, 114, 115, 116, 122, 123 subtriloba123
subsp.guianensis122, 123, 124*, 126*, 128, 129, tergoscabra149
132 tessmannii 137
subsp. venezuelensis 114, 122, 126*, 128, 129* tomentosa5, 15, 110, 111, 112, 114, 115, 116,164,
herrerensis115, 116, 171*, 177, 190, 192*, 193 166 (key), 193, 194
heterophylla123, 155 subsp. apiculata115, 168, 171*, 173
hirsutipetiolata111, 114, 115, 116, 162 (key), 190 subsp. essequiboensis115, 169, 171*
subsp. hirsutipetiolata162, 164* subsp. maroniensis115, 170, 171*, 172*
subsp. hispida 162, 165* subsp. persecta113, 115, 169, 170*, 171*
hispida 162 subsp. tomentosa 115, 166, 167*, 171*, 173
isophlebia186 trianae 194
jaramilloi 159 trilobaKlotzsch 159
johnstonii 141 trilobaTrecul 156, 159
jussiaeana 156 ulei 194
laevis 180 umbellata186
lawrancei133 umbellifera186
longipendula181 uvifera144
maroniensis172 velutina 114, 115, 116, 126*, 128, 129, 131*
melinonii 5, 113, 114, 115, 116, 155, 159, 160 (key) venezuelensis128
subsp.glabrata112, 115, 157*, 161, 163* villosa 111, 176*, 177, 179*
subsp. melinonii 115, 156, 157*, 158*, 160, 193, Schefflera4
194 Urostigma 10
mildbraediana123 intermarginala110
minor 3, 5, 109, 111, 112, 113, 114, 116, 176*,186, Urticaceae2, 3, 15
189
mollis 15, 114, 115, 116, 155, 155-156 (key), 160,
193