Organization for Flora Neotropica

Chrysobalanaceae
Author(s): Ghillean T. Prance
Source: Flora Neotropica, Vol. 9, Chrysobalanaceae (Supplement) (Mar. 8, 1989), pp. 1-267
Published by: New York Botanical Garden Press on behalf of Organization for Flora
Neotropica
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FLORA NEOTROPIC

MONOGRAPH 9S
CHRYSOBALANACEAE

by
Ghillean T. Prance

l\-I- TROPICOF CANCER

FLORA
N E OTROPICA~

TROPIC OF CAPRICORN

Publishedfor

Organizationfor Flora Neotropica

by
The New York BotanicalGarden
New York
Issued 8 March 1989
 Copyright ? 1989
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 = Floraneotropica.
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-dc19 85-647083
AACR 2 MARC-S
Libraryof Congress [8508]
ISBN 0-89327-338-4
 CHRYSOBALANACEAE
GHILLEANT. PRANCE1

TABLE OF CONTENTS
Abstract . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Resumo ..................................................................... ......... 1
Introduction .................................................................................. 2
Embryology .............................. .................................................... 3
Rare Species ........................................................................ .......... 3
Systematic Treatment .......................................................................... 4
Chrysobalanus............................................................................ 4
Licania . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6
Parinari ....... .... ......... ...................... ............. .. .................... ... .. 56
Exellodendron .................................................................. .......... 59
Maranthes ................... ................................................ .......... 59
Couepia .................................................................................. 59
Hirtella.................................................................................. 79
Acioa ................................................................................. .. 96
Neocarya ................................................................................. 97
Acknowledgments ............................................................................. 97
LiteratureCited ............................................................................... 97
NumericalList of Taxa, IncludingNotes on Rarityand Endangerment,
with Index to DistributionMaps .............................................................. 99
SupplementalList of ExsiccataeStudiedSince 1972 Monograph.................................... 107
Second SupplementalList of Exsiccatae.......................................................... 124
DistributionMaps ................................................................ .... 127
Index of Local Names .......................................................................... 263
Index of ScientificNames ...................................................................... 263

ABSTRACT
Sincethe FloraNeotropicaaccountof the Chrysobalanaceae was publishedin 1972, much
new materialhas been collected. This supplementto the earlier monographhas studied
6170 new collectionsand includesthe descriptionsof 67 new taxa that have been described
sincethe monograph,64 of whicharepublishedas new speciesin this work.Fulldescriptions
are given of all new species and they are numberednext to their closest relatives in the
originalmonograph.New generickeys are providedto incorporateall the new species and
otherminorchangesin taxonomythat weremade in the light of the bettermaterialof many
species.

RESUMO
Depois a publicacaoda familia Chrysobalanaceae(1972) em Flora Neotropica muito
materialbotanica foi coletada. Foram estudadas6170 colecoes novas e descritas67 taxa
novas, incluido 64 apresentadasnesta obra. Describcesde tudos esp6cies descritasdesde
1972 e chaves novas paracada genero sao foricidas.

1New York BotanicalGarden, Bronx, New York 10458, U.S.A. Presentaddress:Director, Royal Botanic
Gardens,Kew, Richmond, SurreyTW9 3AB, United Kingdom.
1
2 Flora Neotropica
INTRODUCTION
This supplement to my "Flora Neotropica"
monograph of the Chrysobalanaceae(Prance,
1972) has become necessary because of the
amountof collectingactivityover the last decade
and a half. The 6170 additionalspecimensstud-
ied have included 67 new taxa, numerousrange
extensions and other missing data for species
which were describedfrom incomplete material
in 1972. The addition of the new taxa, as well
as the increasingly recognized morphological
variation of some previously poorly known
species,means that the 1972 keys are out of date
and do not functionfor some of the new material.
I have, therefore,broughttogether all this new
informationinto a supplement,which should be
used in conjunctionwith the 1972 monograph.
In 1972, 7762 specimens were studied, so a
79.48%increaseof the collections has occurred
in a 16 year period. This has produced 64 new
species, or one for every 96.4 collections, indi-
catingthe still poorlycollected statusof the neo-
tropics.
Most of the new taxa have alreadybeen pub-
lished in a series of mainly regionalpapers(Ber-
lin & Prance,1978;Prance, 1973, 1974a, 1974b,
1974c, 1976,1977a, 1979a, 1979b, 1979c, 1979d,
1981, 1984, 1986a).Additionalinformationhas
also been given in Prance (1975), and the Suri-
nam species were treated in Prance and G6rts
van Rijn (1976). The other purposeof this work
is to publish the hitherto unpublisheddistribu-
tion maps of Chrysobalanaceae,most of which
werenot includedin the originalmonograph,and
from which extensive data for several biogeo-
graphicalpapers(for example, Mori et al., 1981;
Prance, 1974d, 1977b, 1979e, 1982a; Prance &
Mori, 1983) have been taken. Since the distri-
butionmapsare,togetherwith distributionaldata
from other families such as the Caryocaraceae
and Dichapetalaceae,the basis for much of my
work on centersof endemism, it is importantto
presentthe maps here. The dots representpres-
ence on a degree squareof latitude, ratherthan
the exact locality of a collection.
Althoughthereare a largenumberof neotrop-
ical species of Chrysobalanaceaethat are still in-
adequatelycollected, the family is much better
known today than it was in 1970 when work on
the first monographwas completed. This sup-
plement discusses briefly new generalinforma-
tion about the family, and then presents data
about each of the species for which significant
new informationhas been gathered.These data
include significantrangeextensions, description
of organsnot previouslydescribed,especiallythe
fruit of several species, and any changes in cir-
cumscription based on the new material. The
new speciesareplacedand numberednext to the
species to which they are most closely related
usinga decimalpoint afterthe number.It is grat-
ifying to note that while many new species have
been found, they all fit extremely well into the
generaas previouslycircumscribed.The only ge-
neric change is in the circumscriptionof Acioa.
This genusoriginallyincludedthe Africanspecies
with ligular stamens, which we have now con-
cluded (Prance& White, 1979) should be sepa-
rated into the genus Dactyladenia.At the same
time, in the Americasthe circumscriptionof the
genus cannot be confined to the three species
with ligularstamens. We also concludedthat in
spite of its free stamens, Couepiaedulis(Prance)
Pranceshouldbe placedwhereI originallyplaced
it before the flowers were known, in the genus
Acioa.
Theamountof newinformationcollectedabout
Chrysobalanaceaein the last 15 yearsis a tribute
to the collectingeffortsof many people through-
out the neotropics. It has enabled us to bring
"FloraNeotropica"from a series based on very
preliminaryinformation to much more defini-
tive treatments,which will certainlybe reflected
in the futuremonographsof the series.
Keys are provided in the systematic section
only for those generain which there have been
changessince 1972.
Apart from an importantcontributionon the
embryologyof Chrysobalanaceae(Tobe & Ra-
ven, 1984, see separatesection), studies of the
pollen (Demchenko,1973;Patelet al., 1983) and
various papers by this author, there have been
few importantpapers on the family since those
reviewed in Prance (1972). However, some re-
gional treatments of the family, based on the
original monograph [Ecuador(Prance, 1979d),
Venezuela(Prance, 1982b), Surinam(Prance&
Gortsvan Rijn, 1976), and the Guianas(Prance,
1986b)], also included considerablymore ma-
terial than the 1972 monograph.Spichigerand
Masson's(1984) review of most of the Peruvian
species of Chrysobalanaceae,in an account of
the species in the ArboretumJenaroHerrerain
RareSpecies
Loreto Department, was also largely based on
the "FloraNeotropica"account.
The monographsof the 61 African species of
Chrysobalanaceaehave also allowed an inter-
esting comparison between the two continents
(Letouzey& White, 1976, 1978a, 1978b;White,
1976).This also led to the removalof the African
species with a staminalligule from the neotrop-
icalAcioato a distinctgenusDactyladeniaWelw.
(Prance& White, 1979). Likewisethe species of
Malesian Chrysobalanaceaehave recentlybeen
revisedand theirrelationshipsto neotropicaltaxa
arenow betterunderstood(Prance,1979b, 1987,
in press).Finally,a detailedworldwidetreatment
of the genera of Chrysobalanaceae(Prance &
White, 1988) has been completed.
EMBRYOLOGY
At the time of the 1972 monographlittle de-
tailed workhad been done on the embryologyof
the Chrysobalanaceae. RecentlyTobe and Raven
(1984) studiedthe embryologyof threeneotrop-
ical species-Chrysobalanus icaco L., Licania
apetala (E. Mey.) Fritsch, and L. michauxii
Prance. This has at least provided some basic
data for the tribe Chrysobalaneae.The most im-
portantembryologicalfeaturesare given below.
Antherstetrasporangiate,the walls five layers
thick, formation of the Basic type; epidermis
persistentand often tanniniferous,endothecium
fibrous, the two middle layers ephemeral, the
tapetum glandularwith two-nucleatecells. Cy-
tokinesis in the microsporemother cell simul-
taneous; microspore tetrad usually tetrahedral
and occasionally decussate; pollen grains two-
celled when shed. Ovule anatropousand tenui-
nucellate,the nucellus very small; all archespo-
rial cells developing directly into megaspore
mother cells, undergoing meiosis; megaspore
tetradslinear;chalazalmegasporefunctional,de-
velopinginto a modifiedtype of the Polygonum-
type embryo sac; mature embryo sac four-nu-
cleate, comprisingan egg, two synergidsand a
central nucleus which is probably triploid; an-
tipodal cells absent from the beginning,hypos-
tase differentiatingin Chrysobalanusbut not Li-
cania; the nucellar tissue soon disintegrating
except the megaspore and embryo sac. Ovule
bitegmic,the inner integument(ii) and outer in- Flora Neotropica it was recommended that
tegument(oi) initiated desmally,the ii up to five monographers comment on and point out rare
or eightcells thick and the oi more than five cells andendangeredspeciesin theirgroups.Although
3
thick; inner epidermis of ii developing into the
endotheliumwhich directlybordersthe embryo
sac and accumulates starch grains; micropyle
formed by both integuments. Fertilizationpo-
rogamous;endospermformationof the Nuclear
type;seed exalbuminous.Young seed coat com-
posed of both testa and tegmen; exotegmen fi-
brous;exotestaone-layered,composedof cuboid
tanniniferouscells; endotesta a few layers thick
composed of unspecialized small cells; micro-
testa 30-50 cells thick in C. icaco, composed
mostly of vessels, 3-4 layered and lacking vas-
cular tissue in L. michauxii. Mature seed me-
sotestal.
The embryologicaldatagiven above show that
thereis little differencebetweenthe species stud-
ied. The Chrysobalanaceaediffer from the Ro-
saceae, to which it has frequentlybeen related,
in the tenuinucellateovule, the very small nu-
cellus that disintegratesearly, and the presence
of an endothelium.Tobe and Raven conducted
their investigation to consider the possible re-
lationship of the Chrysobalanaceaeto the Myr-
tales, but concluded that it is embryologically
quitedifferentfromthatwell-definedorder.They
proposed a relationshipto the Theales, which
share with Chrysobalanaceaea tenuinucellate
ovule, a small disintegratingnucellus, an endo-
thelium,Nuclear-typeendospermformation,and
an exalbuminousseed. However,the Thealesdif-
fer from Chrysobalanaceaein their one-celled
ovule, archesporium,an Allium-typeembryosac,
and persistentantipodalcells. In their argument
Tobeand Ravenmisrepresentthe Chrysobalana-
ceae as lacking stipules. Since there are major
embryologicaldifferencesbetween the Chryso-
balanaceaeand both the Rosales and Theales,it
seemsunlikelythatthe familyis anybetterplaced
in the Theales than in the Rosales and a more
definite placementcan only be made after con-
siderabledata are obtainedfrom other fields, as
well as embryologicalinformationfrom a much
widerrangeof Chrysobalanaceae,insteadofjust
threespeciesin the two most closely relatedgen-
era.
RARE SPECIES
At the 1986 meeting of the Organizationfor
4 Flora Neotropica
ourknowledgeof tropicalgroupssuchas Chryso-
balanaceaeis far from adequate, the collection
data arenow certainlysufficientto pick out a few
of the rarestspecies. All rarespecies are marked
in the specieslist precedingthe list of exsiccatae,
and it gives the rarestand possibly most endan-
gered.The species listed as rareinclude all those
known from a single collection and not re-col-
lected since 1972 as well as others that appear
to be particularlythreatenedby virtue of their
habitat.Speciesdescribedsince 1972 and known
from a singlecollectionare only listed if they are
from particularlythreatenedareas such as At-
lanticcoastalforestsof Brazilor fromRond6nia,
as it is too early to judge their frequencyfrom
SYSTEMATICTREATMENT
1. Chrysobalanus
Linnaeus
Revised Key to Species of Chrysobalanus 1-3. Chrysobalanus venezuelanus Prance, sp.
1. Leavesorbicularto ovate-elliptic,the apexretuse
to roundedor minutely and bluntly acuminate;
midribglabrousbeneath. 1. C. icaco.
1. Leavesellipticto oblong,with a distinctacumen
5-15 mm long; midribsparselyhirsutebeneath.
2. Leaves chartaceous, acuminate; branches
conspicuouslylenticellate;inflorescencegray-
puberulous. 2. C. cuspidatus. cellatis, inflorescentiisferrugineo-pubescentibus
2. Leavesthicklycoriaceous,caudate;branches
not conspicuouslylenticellate;inflorescence
3. C. venezuelanus.
ferrugineous.
1-1. Chrysobalanus icaco Linnaeus, Sp. pl. 1:513.
1753.
ChrysobalanusinteriorSmall, Man. s. e. fl. 645. 1933.
Type. U.S.A. Florida:Hammocks, Long Key, Ev-
erglades,Small & Carter3166 (lectotype,here des-
ignated,NY).
The synonym C. interior was overlooked in
Prance (1972) and in the treatment for the ge-
neric flora of the southeastern U.S. (Prance, 1970).
This name, although validly published, was not
picked up in either the "Index Kewensis" or the
"Gray Herbarium Card Index." There are no
significant differences and C. interior falls well
within the range of variation of C. icaco as cir-
cumscribed in Prance (1972).
The distribution of C. icaco will be found in
Figure 20.2
2
Figures 20 to 155 are grouped at the end of the
monograph.
collectingactivity.The endangermentstatusused
in the list is based on the collection record and
on the currentcondition of the habitatin which
they grow. Poorly known species from relatively
undisturbedareas such as the Guianas are not
generallylisted unless they are known only from
a single collection.
I consider as endangered31 of the 100 rarest
species.These aremainlyfrom areasthat arefoci
of forest destruction such as Central America,
Choc6, Rond6nia, and eastern Brazil. The fact
that there are at least 100 rare species, 31 of
which are endangered,shows the desperateneed
for a greaterconservationeffortof the neotrop-
ical forests.
nov. Type. Venezuela.Bolivar:IcabaruiRiver
region, headwaterof Rio Hacha, 450-850 m,
3 Jan 1956 (fl),A. L. Bernardi2777 (holotype,
NY; isotype, NY). Fig. 1.
Species a C. cuspidatofoliis coriaceis, latiori-
bus, ramulis juvenilibus haud conspicue lenti-
differt.
Treeto 10 m tall, the young branchessparsely
puberulousto glabrescent.Leaf lamina oblong-
elliptic, coriaceous, 5.5-11 x 2.2-4.2 cm, cu-
neate at base, caudate at apex, the acumen 10-
17 mm long, glabrousand shiny above, glabrous
beneathexceptfor appressedhairson midriband
primary veins of lower surface of very young
leaves; midrib prominulous above, prominent
beneath;primaryveins 6-8 pairs,widely spaced,
prominulouson both surfaces;petioles 2-4 mm
long, terete, rugulose, glabrous. Stipules, ca-
ducous, membranous,axillary.Inflorescencesof
few-floweredcymules insertedon rachisto 5 cm
long, the rachis and branches ferrugineous-pu-
bescent; bracts and bracteoles ovate, membra-
nous, puberulous on exterior, ca. 1 mm long,
caducous.Flowers3 mm long, insertedin small
cymules; receptaclecampanulate,tomentellous
on exterior,tomentosewithin;calyxlobes round-
ed, tomentellous;petalsfive, ovate, glabrous;sta-
mens 14-15, insertedaroundcompletecircle,the
filamentsshortly exserted,pubescent;ovary in-
Systematic Treatment 5

Gfwysobarlana

a( cmj
,C~ 0
.

r ?I i
~~~~~~~~~~~~~~~~~v10

I~~~~~~~~~~~~~~~~~
-V~~~~

/Th II h4.?-"

'A ~ ~ ~)vc A: .

FIG. 1. Chrysobalanusvenezuelanus(Bernardi2777). A, habit; B, leaf undersurface;C, flower buds; D,

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

sertedat base of receptacle,lanate;style equalling
filamentsin length. Fruit (only seen young) ob-
long, deeply costate; exocarp glabrous except
when very young.
Distribution(Fig. 21). Forest on terrafirmein
easternVenezuela.
Additionalspecimensexamined.VENEZUELA.
BOLiVAR:W base of Amaruay-tepui,5?56'N,62?17'W,
5 May 1986 (fl), Hoist & Liesner 2788 (MO, NY); 4
km W of El Pauji, Rio Chaberu,750-900 m, 12 Nov
1985 (y fr), Liesner 19919 (MO, NY); Distr. Piar, Rio
Aparaman, SW corer of Amaruay-tepui, 5?54'N,
62?15'W,22 Apr 1986 (fl),Liesner&Hoist 20192 (MO,
NY); QuebradaLos Brasileros,4.5 km SW of Icabaru,
4?20'N,61?48'W,480 m, 16 Dec 1978 (fl),Steyermark
et al. 117696 (NY, VEN).
This species differs from the closely related
6 FloraNeotropica

Chrysobalanuscuspidatus(Fig. 21) of the Lesser South America. The genus is predominantly

Antillesin the more robustinflorescencebranch-
es with a ferrugineous,not graypubescence,the
broader, more coriaceous leaves, and the less
conspicuouslylenticellateyoung branches.This
is a most interesting occurrence of the genus
Chrysobalanussince it is frominlandcontinental
coastal (C. icaco) and from cloud forest in the
LesserAntilles (C. cuspidatus).The new species,
C. venezuelanus,is found at elevations between
450 and 850 m and is thus in a habitat very
similar to that of C. cuspidatusin the montane
forests of Guadeloupeand Martinique.

2. Licania Aublet
Revised Key to Species of Licania3
Since the genus is large,now containing 190 species, the key is divided for convenienceinto three
parts. Note: Species 1-152 are fully describedin Prance(1972) and the 38 species describedsince
1972 or in this work have complete descriptionsfollowing these keys.
1. Stamenslongerthan calyx lobes. Key A: SubgenusMoquileasections Moquileaand Leptobalanus.
1. Stamensequal to or shorterthan calyx lobes.
2. Stamens 10-25. Key B: SubgenusMoquileasections Microdesmiaand
Leptobalanuspro parte;subgenusParinariopsis.
2. Stamens3-9. Key C: SubgenusLicania.

Key A
Specieswith 8-60 stamens,exsertedbeyond calyx lobes.
1. Petals present;stamens 11-60. (For contrastingstatementsee p. 9.) Subg.Moquileasect. Moquilea.
2. Leaf undersidewith an appressedlanate-arachnoidpubescence.
3. Inflorescenceof fasciculate,short,dense-floweredracemes;primaryveins impressedabove (Pan-
ama). 5.3. L. fasciculata.
3. Inflorescenceof lax, branched,racemose panicles or cymose panicles; primaryveins usually
prominulousor plane above.
4. Primaryleaf veins 8-15 pairs, the lamina oblong-lanceolate.
5. Petioles 1-3 mm long; fruit 2.5-3 mm long, with smooth exterior,pericarpthin; inflores-
cence of small cymose panicles(SE U.S.A.). 1. L. michauxii.
5. Petioles 5-7 mm long;fruit4-7 mm long, with a crustaceous,verrucoseexterior,pericarp
thick;inflorescenceof racemosepanicles (Bolivia, La Paz). 2. L. boliviensis.
4. Primaryleaf veins 16-29 pairs, the lamina oblong to elliptic.
6. Inflorescenceramiflorous,borne on young woody twigs well below shoot apex.
7. Flowers(calyxand receptacle)6-7 mm long,youngleafundersurfacewith shaggydense
ferrugineouspubescence;leaves 27-47 cm long (Colombia,Valle, 0-50 m).
5.1. L. gentryi.
7. Flowers2-5 mm long, leaf undersurfacewith compact appressedpubescenceor with
caducouspubescence,becomingglabrouswhen mature;leaves 16-33 cm long.
8. Leaf undersurfaceglabrouswhen mature, with a lanate caducous pubescenceon
youngestleaves; flowersca. 2 mm long (Colombia,Valle, 0-50 m). 46. L. velata.
8. Leafundersurfacewith a compactpersistentgray-lanateor brownpubescence;flow-
ers 3-5 mm long.
9. Leaf undersurfaceferrugineousto brown-tomentose,secondaryvenation not
conspicuouslyreticulate(Colombia;Ecuador;Peru). 5. L. macrocarpa.
9. Leaf undersurfacegray-tomentellous,conspicuouslyreticulatewith parallelsec-
ondaryvenation;leaves thickly coriaceous.
10. Inflorescence15-30 cm long, much-branched,flowersdenselypacked,brac-
teoles caducous(Colombia;Ecuador). 4. L. durifolia.
10. Inflorescence8-15 cm long, unbranchedor with a few branches;bracteoles
persistent,membranous(Ecuador). 4.2. L. grandibracteata.

3 Note that Licania

cecidiophora,L. mexicana, and L. tepuiensisare not included in these keys since they
were describedfrom incompletematerial.
SystematicTreatment 7
6. Inflorescenceterminalor axillary,but on leafy shoot apex.
11. Flowers 1.5-2 mm long;leaves 12-19 x 3-6 cm, brown-pubescentbeneath,without
conspicuouslyreticulatevenation (Colombia). 148. L. veneralensis.
11. Flowers3-5 mm long; leaves 15-35 x 6-13 cm, gray-pubescentbeneath,often with
conspicuouslyreticulateparalleltertiaryvenation.
12. Inflorescenceand flowerswith light-brown-tomentellous pubescence,leavesgrad-
ually taperingfrom mid-pointto acuminateapex;centerprimaryveins far apart
(18-22 mm), the secondaryvenation obscure(Colombia,Choc6). 3. L. maritima.
12. Inflorescenceand flowerswith ferrugineouspubescence;leaves taperingfromwell
above midpoint,centerprimaryveins 5-13 mm apart,with conspicuousparallel
secondaryvenation.
13. Leaves 6-9.5 x 2-3 cm; petioles 4-5 mm long (Venezuela,Lara).
5.4. L. montana.
13. Leaves 12-46 x 3-26 cm; petioles 6-12 mm long.
14. Inflorescencesof racemosepanicles,to 18 cm long; petioles 10-12 mm
long;primaryveins 16-19 pairs,pedicels5 mm long (Colombia,Antio-
quia). 5.2. L. cabrerae.
14. Inflorescencesunbranchedracemes,or little-branchedpanicles,5-13 cm
long. Petioles 6-8 mm long, primaryveins 21-24 pairs;flowerssessile
(Peru,San Martin). 4.1. L. filomenoi.
2. Leaf undersideglabrouswhen mature(sometimeslanate when young).
15. Inflorescenceand exteriorof flowersglabrousor sparselypuberulous.
16. Leaf broadlyacuminate,the acumen 4-8 mm long; flowers2.5-3 mm long; pedicels 2-5
mm long (Brazil,Bahia). 6. L. salzmannii.
16. Leaf with well-developedfinely pointed acumen 8-12 mm long; flowers4-4.5 mm long;
pedicels 1-3 mm long.
17. Inflorescenceandexteriorof flowerspuberulous;petioles 10-12 mm long;leaveselliptic,
5.5-9 cm broad(Peru,Loreto). 7. L. klugii.
17. Inflorescenceand exterior of flowers glabrescent;petioles 5-7(-9) mm long; leaves
oblong to oblong-elliptic,2-5.5 cm broad (Guianas;PeruvianAmazonia).
8. L. guianensis.
15. Exteriorof flowersand usuallyinflorescencedensely tomentose.
18. Inflorescenceeither of dense pyramidalor cymose panicleswith a glabrousto puberulous
rachis,or a muchbranchedpaniclewith the primarybranchesbranchedinto smallbranches
bearing2 or more flowers.
19. Inflorescencea much branchedpanicle 10-25 cm long, with the primarybranches
branchedinto small branches;leaves 5.5-10.5 cm broad (Ecuador). 13.1. L. dodsonii.
19. Inflorescenceof dense pyramidalor ratherlax cymose panicles 1-5 cm long;leaves 1-
4.5 cm broad.
20. Inflorescenceshortand dense, the rachisglabrous;petioles4-6 mm long (Mexico).
9. L. retifolia.
20. Inflorescencelax, the rachispuberulous;petioles 1-3 mm long (SE U.S.A.).
1. L. michauxii.
18. Inflorescenceof lax racemesor racemosepanicles,the rachispubescent.
21. Pedicels 3-7 mm long; leaves with finely pointed acumen 1.5-2.5 cm long (Brazil,
Amazonas). 10. L. longipedicellata.
21. Pedicels0.5-2.5 mm long; leaf acumenratherblunt, less than 10 mm long.
22. Inflorescenceof predominantlyaxillary racemes or few-branchedpanicles (NE
Brazil). 11. L. tomentosa.
22. Inflorescenceof predominantlyterminalmuch-branchedpaniclesor racemosepan-
icles, rarelyaxillary,then much branched.
23. Leaveswith a lanatepubescencewhenyoung;petioles8-16 mm longand very
slender,not more than 1.2 mm thick (W Indies;Venezuela). 12. L. pyrifolia.
23. Leavesglabrouseven when young;petioles 3-12 mm long, stout, more than
1.5 mm thick.
24. Leafbases cuneate;stamensca. 30; petiolescanaliculate(W Indies;Ven-
ezuela). 13. L. leucosepala.
24. Leafbases cordate,roundedor subcuneate;stamens 12-23; petioles usu-
ally terete,rarelycanaliculate.
25. Leaves oblong-lanceolate,primaryveins slightlyimpressedabove.
26. Petioles 10-13 mm long;leaves 10-14.5 x 2.5-5.2 cm; primary
veins 11-17 pairs;stamens 17 (Peru;Brazil,W Amazonia).
14. L. angusataa.
8 Flora Neotropica
26. Petioles 5-7 mm long; leaves 14-25 cm long x 4.5-8 cm; pri-
mary veins 18-22 pairs;stamens 12-14 (Brazil,Para).
14.1. L. anneae.
25. Leavesovate to oblong(to oblong-lanceolatein L. platypus);primary
veins plane or prominulousabove.
27. Leaves 13-30 cm long; primaryveins 13-22 pairs, leaf bases
sometimes subcordate.
28. Flowersborne in small cymuleson short tertiarybranches;
petioles 12-16 mm long, canaliculate(Venezuela).
23.1. L. tachirensis.
28. Flowersborne on primaryinflorescencebranches;petioles
8-14 mm long, terete.
29. Leaves oblong to oblong-lanceolate,3.5-8 cm broad;
primaryveins 15-22 pairs;petioles 9-14 mm long;leaf
base roundedto subcuneate(CentralAmerica).
15. L. platypus.
29. Leaves elliptic, 7.5-9.5 cm broad;primaryveins 14-
15 pairs;petioles ca. 8 mm long; leaf base subcordate
(Brazil,Maranhao). 19. L. maranhensis.
27. Leaves 5-15.5 cm long (to 21 cm in L. guatemalensis);primary
veins 8-13 pairs (to 15 in L. britteniana);leaf bases subcuneate
to rounded(to subcordatein L. britteniana).
30. Petals not unguiculate,equallingor only shortlyexceeding
calyx lobes, filamentsfar exceedingpetals.
31. Flowers4-5 mm long; stipuleslinear-filamentous,5-7
mm long (Mexico). 16. L. gonzalezii.
31. Flowers 2-3.5 mm long; stipules short-subulate,to 2
mm long.
32. Petioles 2-5 mm long; leaves obovate or elliptic,
with cuspidate,mucronate,or abruptapex.
33. Inflorescenceonce-branched;leaves obovate
to elliptic,4.5-8.5 cm long (Brazil;Colombia;
Peru, Amazonia). 17. L. egleri.
33. Inflorescence twice-branched;the flowersborne
in small groupson short secondarybranches;
leaves elliptic, 10-21 cm long (Guatemala).
18.3. L. guatemalensis.
32. Petioles 6-14 mm long; leaves oblong to oblong-
elliptic, apex graduallyacuminate(cuspidatein L.
kallunkiae).
34. Inflorescencerachis andbranchessparselygray-
pubescent.
35. Leafapexcuspidate,the base subcuneate;
stamens 11-12 (Panama).
18.2. L. kallunkiae.
35. Leaf apex acuminate,the base rounded;
stamens 16-25 (Brazil,Amazonia;Peru).
20. L. fritschii.
34. Inflorescencerachisand branchestomentose.
36. Inflorescencesmall and little-branched;
flowers3-4 mm long;ovaryglabrous;pet-
ioles glabrous(Panama).
18.1. L. chiriquiensis.
36. Inflorescencelarge and much-branched;
flowers 2-3 mm long; ovary pubescent;
petioles tomentose when young.
37. Flower buds globose; flowers 2 mm
long; leaves 5.5-11 x 2-6 cm
(Guianas;Brazil;Venezuela).
18. L. minutiflora.
37. Flowerbudsovoid;flowers2.5-3 mm
long; leaves 8-16 x 3-7.5 cm (Co-
lombia;Peru;Brazil,Amazonia).
21. L. britteniana.
SystematicTreatment 9
30. Petals unguiculate,much exceedingcalyx lobes; filaments
scarcelyexceedingpetals.
38. Flowers 5-6 mm long; pedicels 2-2.5 mm long; ovary
pilose (Brazil,Amazonia). 22. L. unguiculata.
38. Flowers 3 mm long; pedicels 0-0.25 mm long; ovary
glabrescent(Peru;Brazil,Amazonia). 23. L. longipetala.
1. Petals absent;stamens 8-14. Subg.Moquileasect. Leptobalanus.
39. Leaf undersideglabrousor lanate, the venation not prominent,the stomatalcavities absent.
40. Most flowersborne in small groupsor cymules on distinct secondarybranches(peduncles)of
inflorescence;pedunclesover 2 mm long.
41. Rachis of inflorescencedensely yellow-villous-tomentose(Venezuela;Brazil,Amazonia).
24. L. wurdackii.
41. Rachis of inflorescencegray-puberulous.
42. Inflorescenceshort and compact, 3-5 cm long; flowers 4-5 mm long; leaves ovate
(NE Brazil). 25. L. turbinata.
42. Inflorescencelonger and spreading,6-20 cm long; flowers 2-3.5 mm long; leaves
oblong to elliptic.
43. Flowers3.5-5 mm long;petioles deeplycanaliculate;with two prominentglands
at lamina base;rachisarachnoid-pubescent when young (Amazonia). 26. L. lata.
43. Flowers2-3 mm long;petiolestereteto shallowlycanaliculate;laminabase eglan-
dular;rachispuberulous(Guianas;Brazilto S Amazonia). 27. L. apetala.
40. Flowerssessile or nearlyso on primarybranchesof inflorescence.
44. Leaf apex predominantlyobtuse to rounded.
45. Rachisof inflorescencedenselyvillous-tomentose;flowersca. 4 mm long (Venezuela;
Brazil,NW Amazonia). 24. L. wurdackii.
45. Rachis of inflorescencegray-puberulous; flowers3 mm long (Guianas;Brazil,Ama-
zonia). 28. L. parvifolia.
44. Leaf apex distinctlyacuminate(rarelyonly bluntlyacuminate,L. jefensis).
46. Flowers4-5 mm long;receptaclecupuliform.
47. Leaves glabrous beneath, inflorescencegray-villous-tomentose(Brazil, Mato
Grosso). 29. L. maguirei.
47. Leaveswith short persistentwaxy lanate-pulverulentpubescencebeneath;inflo-
rescencebrown-puberulous to tomentellouswhenmature(AmazonianColombia;
Peru;Brazil). 26. L. lata.
46. Flowers2-3.5 mm long; receptacleurceolateor campanulate.
48. Stipulespersistenton young branches;petioles densely tomentose when young,
2.5-3.5 mm thick;upperportion of ovary glabrous(centralBrazil). 30. L. gardneri.
48. Stipulesusuallycaducous;petioles sparselylanate-pubescent,puberulous,or gla-
brous, to 2 mm thick;upperportion of ovary pubescent.
49. Inflorescencesparselyhirsutulousor yellow-arachnoid-tomentose; flowerex-
terioryellow-brown-tomentose; petioles 2-3 mm long, canaliculate.
50. Inflorescencesparselyhirsutulous;leaf apex finely acuminate,the acu-
men 5-10 mm long (N Colombia). 31. L. cuspidata.
50. Inflorescencearachnoid-tomentose; leafapexacuteorbluntlyacuminate,
the acumen 3-5 mm long (Panama). 31.1. L. jefensis.
49. Inflorescencepuberulousto tomentose;flowerexteriorgray-tomentose;pet-
ioles 3-11 mm long, tereteor shallowlycanaliculatetowardsbase.
51. Leaf bases subcordate(Panama). 31.2. L. morii.
51. Leaf bases cuneateor subcuneateto rounded.
52. Leaf undersurfacerufous-brown-pubescent, with deeply reticulate
venation;petioles 8-11 mm long;leaf apex cuspidate;inflorescence
brown-tomentellous(Colombia,Valle). 31.3. L. cuatrecasasii.
52. Leafundersurface glabrousorrarelywhite-arachnoid-pubescent, ve-
nation smooth, not deeply reticulate;petioles 3-8 mm long; leaf
apex acuminate;inflorescencebrownor gray-puberulous to tomen-
tose.
53. Rachis and branchesof inflorescencetomentose;petioles usu-
ally bearingtwo distinct glands(CentralAmerica).
32. L. sparsipilis.
53. Rachis and branches of inflorescence puberulous;petioles
eglandular.
54. Inflorescence andflowerexteriorsparselypuberulous;leaves
thicklycoriaceous,the apex finelyattenuatewith acumen
10-25 mm long (Guianas;N AmazonianBrazil).
27.1. L. granvillei.
10 Flora Neotropica

54. Inflorescenceusuallydenselygray-puberulous; leaveschar-

taceous, the apex acuminatewith acumen 3-15 mm long
(Colombia;Peru;Venezuelato S Brazil). 27. L. apetala.
39. Leafundersurface with stomatalcavitiesor verydeeplycut reticulationresemblingstomatalcavities,
lanate-pubescentin mouth of cavities.
55. Inflorescencewith secondarybranches;flowerson distinct secondarybranchesof inflorescence
that are at least 2 mm long.
56. Flowersin small distinct groupsor in cymules on short secondarybranches(peduncles);
leaf uppersurfacesmooth, the base roundedto cuneate.
57. Inflorescence15-22 cm long, ferrugineous-arachnoid; leaf undersidebrownwhen dry
(Guyana). 35. L. persaudii.
57. Inflorescence6-8 cm long, gray-arachnoidor puberulous;leaf undersidegray when
dry (Guianas;Brazil,N Amazonia). 36. L. sprucei.
56. Flowers more or less clusteredin large groups on primaryand secondarybranches of
inflorescence,but withoutdistinctpeduncles;leaf uppersurfaceusuallypapillose,the base
often cordate(Brazil,Amazonia and south). 37. L. sclerophylla.
55. Inflorescencewith primarybranchesonly;flowerson primarybranchesof inflorescence,sessile
or rarelywith short pedunclesnot exceeding 1 mm in length.
58. Pedicels 2 mm long (in bud), to 4 mm in flower(Guianas). 38. L. albiflora.
58. Pedicels not exceeding2 mm, often shorter.
59. Leaf reticulationextremely prominent on upper surface;fruit exterior appressed-
sordid-yellow-velutinous.
60. Petioles 5-13 mm long; flowersca. 3 mm long, borneon slenderbranchlets0.5-
1 mm thick; receptaclecampanulate;flowersand inflorescencegray-tomentose
(Panama;Venezuela;Guianas;Brazil,Amazonia). 39. L. longistyla.
60. Petioles 15-20 mm long; flowersca. 4 mm long, borne on branchletsca. 2 mm
thick;receptaclecupuliform;inflorescenceand flowersbrown-tomentose(Colom-
bia, Pacific). 40. L. fuchsii.
59. Leaf reticulationprominulouson upper surface;fruit exterior glabrousor brown-
tomentose.
61. Young branchesdensely tomentellous,with a thick corky bark(centralBrazil).
41. L. humilis.
61. Young branchesglabrousor nearlyso, the barkthin.
62. Flowers 3.5-5 mm long; leaf apex mucronate(Guyana). 42. L. foveolata.
62. Flowers2-3 mm long;leaf apex roundedto acuminate(Venezuela;Guianas;
centraland AmazonianBrazil). 43. L. octandra.

Key B
Specieswith 10-25 stamens,equallingor shorterthan calyx lobes.
1. Petals present.
2. Ovaryinsertedat base of receptacle;bractsand bracteolessmall and not enclosinggroupsof flower
buds. Subg.Moquileasect. Microdesmia.
3. Leaf undersidewith stomatal cavities or deeply cut reticulation;filamentsconnate over half of
lengthin groups,densely pubescentthroughout.
4. Pubescenceof inflorescencelight gray;primaryleaf veins prominulousabove (NE Brazil).
44. L. rigida.
4. Pubescenceof inflorescenceferrugineous;primaryleaf veins slightlyimpressedabove (Central
America;W South Americato Peru). 45. L. arborea.
3. Leaf undersidewith inconspicuousvenation, lackingstomatalcavities; filamentsfree almost to
base, glabrouson upperportion.
5. Leaves 15-36 cm long; stipules 12-22 mm long.
6. Leafundersurfacewith caducous,lanatepubescence;anthersreniform;inflorescenceborne
on woody defoliatedstalks (Colombia,Valle). 46. L. velata.
6. Leafundersurfacehirsute;anthersdeltoid;inflorescenceterminal(FrenchGuiana;Brazil,
Amapa). 149. L. amapaensis.
5. Leaves less than 14 cm long; stipules 2-10 mm long; flowersin terminalor axillaryinflores-
cences on young branchlets.
7. Inflorescenceof unbranchedsubspikesor racemes.
8. Flowers ca. 1.5 mm long, borne in clusters along rachis, the rachis, 4-15 mm long
(Colombia,Boyaca). 47. L. subarachnophylla.
8. Flowers ca. 2.5 mm long, not clustered,the rachis 2-4 mm long (Peru, Madre dos
Dios). 47.1. L. tambopatensis.
Systematic Treatment 11

7. Inflorescenceof panicles,or a panicle of cymules.

9. Leaf undersidedensely lanate-pubescent.
10. Leaf apex with a well-developedacumen 3-5 mm long; stamensca. 25; exterior
of receptacleferrugineous(Colombia,Antioquia). 48. L. salicifolia.
10. Leafapex obtuseto broadlyacuminate,the acumennot exceeding1 mm; stamens
ca. 15; exteriorof receptaclebrown(Brazil,Goifs). 49. L. araneosa.
9. Leaf undersideglabrous.
11. Inflorescencea panicle of small cymules along the rachis and short secondary
branches;petioles 2-5 mm long; stamens 8-10 (Old World).
Sect. Hymenopus.147. L. splendens,L. palawanensis.
11. Inflorescenceof panicles,flowersnot groupedin cymules;petioles 6-11 mm long;
stamens 14-25 (New World).
12. Leaf acumen 3-7 mm long; leaves 5-10 cm long; stamens ca. 25; flowers
sessile on primarybranchesof inflorescence,denselybrown-tomentose(east-
ern Brazil). 50. L. silvatica.
12. Leafacumen 10-15 mm long;leaves 7.5-14 cm long;stamens 12-14; flowers
predominantlyon short secondarybranchesof inflorescence,short gray-to-
mentose (Colombia,Valle). 51. L. chocoensis.
2. Ovary inserted laterally;bracts and bracteoleslarge and enclosing small groups of flower buds
(Venezuela;Guianas;Brazil,Amazonia). Subg.Parinariopsis.52. L. licaniiflora.
1. Petalsabsent.
13. Leaf undersideglabrous,or with an easily removed lanate pubescence,the reticulationsshallow,
stomatalcavities absent. Subg.Moquileasect. Leptobalanus.
14. Leaf undersurfaceglabrous;flowersin groupson distinct secondarybranchesof inflorescence;
stamens 10-19 (Amazonia).
15. Stamensca. 19; leaves 15-20 cm long, with long acumen 11-19 mm long; flowersca. 5
mm long (Brazil,N Amazonia). 33.1. L. joseramosii.
15. Stamens 10-11; leaves 3-8.5 cm long, with acumen 0-6 mm long; flowers2-3 mm long
(Venezuela;Brazil,Amazonia). 33. L. emarginata.
14. Leaf undersidelanate; flowers sessile on primarybranchesof inflorescence;stamens ca. 22
(Colombia,Valle). 34. L. calvescens.
13. Leaf undersidewith stomatalcavities or with deep reticulations,pubescencedense and fillingthe
hollows betweenvenation. Subg.Licania sect. Licania (pro parte).
16. Receptacleconical;petioles usuallycanaliculate,8-13 mm long; midrib impressedon upper
surfacetowardslamina base; leaves dryinglight brown(Guianas;Brazil,N Amazonia).
108. L. majuscula.
16. Receptaclebarrel-shaped;petioles terete, 3-6 mm long; midrib prominulousat base above,
but becomingimpressedapically;leavesdryingdarkbrown(Colombia;Peru;Venezuela;Brazil,
Amazonia). 140. L. mollis.

Key C
Specieswith 3-9 stamens,equallingor shorterthan calyx lobes.
1. Leaf undersideglabrousor sparselyhirsute or hispid along venation; petals present or absent. (For
contrastingstatementsee p. 13.)
2. Petals present.(For contrastingstatementsee p. 13.)
3. Leaves sparselyhirsutealong venation of lower surfaceor hispid at least on midrib. Sect. Hirsuta.
4. Inflorescenceand flowers sparselyhirsutulous;leaf lamina appearingdeeply rugose above;
primaryveins and venation deeply impressedon uppersurface.
5. Leaf oblong, apex acuminate(Brazil,Amazonia). 53. L. hirsuta.
5. Leaf ovate to ovate-elliptic,apex roundedto obtuse (CentralAmerica). 54. L. costaricensis.
4. Inflorescenceand flowersdensely pubescent;leaf lamina not rugose,primaryveins plane or
impressedand venation more or less plane on uppersurface.
6. Young stems, leaf midrib, and lower inflorescencebrancheshispid (Venezuela).
57.1. L. hispida.
6. Young stems puberulous;leaf midrib hirsute;inflorescenceand exteriorof flowerswith
shortgraypubescenceor densely ferrugineous-tomentose.
7. Inflorescenceand exteriorof flowerswith a shortgraypubescence(Trinidad;Venezuela;
Guianas;Brazil,Amazonia). 69. L. heteromorpha.
7. Inflorescenceand exteriorof flowersdensely ferrugineous-tomentose.
8. Leaf chartaceous,apex acuminate(Bolivia, Amazonia). 55. L. krukovii.
8. Leaf thick-coriaceous,apex roundedto apiculate.
12 Flora Neotropica
9. Primaryleaf veins 6-9, prominulousabove; petioles 3-4 mm long (Guyana;
Venezuela). 56. L. lasseri.
9. Primary leaf veins 12-20, slightly impressed above; petioles 7-15 mm long
(Guyana;Brazil,Amazonia). 57. L. latifolia.
3. Leaf undersideglabrous. Sect. Hymenopus.
10. Flowersin small cymuleson shortdistinct secondarybranchesof inflorescenceat least 1 mm long.
11. Leaveswith a finely pointed acumen 15-30 mm long; flowersca. 1 mm long; ovary and style
pubescent(Colombia,Valle). 58. L. minuscula.
11. Leaves roundedor acute to broadlyacuminate,the acumen 2-8.8 mm long; flowers2-5 mm
long; ovary and style glabrousor pubescent.
12. Inflorescencecompact and triangular,to 8 cm long; rachis and branchesglabrous,the
rachisnot lenticellate;peduncles2-4 mm long;exteriorof flowersglabrous(CentralAmer-
ica). 59. L. operculipetala.
12. Inflorescencelax and spreading,over 8 cm long, or short, lax and of small cymules on
centralrachis;rachisand branchespuberulous,the rachisoften lenticellate;peduncles0.5-
2 mm long; exteriorof flowerspuberulous.
13. Inflorescenceof small cymules on a central rachis 1.5-10 cm long; stamens 8-10;
receptacleoblique in bud; maturefruit to 1.5 cm long, usually smaller,not costate
(Old World,Asia and Pacific). 147. L. splendens,L. palawanensis.4
13. Inflorescencelax and spreading,over 8 cm long;stamens5-7; receptaclesymmetrical
in bud; maturefruit 1.5-4 cm long, costate (New World).
14. Flowers 4-5 mm long; inflorescencerachis 2-3 mm thick; leaves thickly coria-
ceous (Venezuela). 60.1. L. pakaraimensis.
14. Flowers 1.5-3 mm long; inflorescencerachis 1 mm thick;leaves chartaceousor
thinly coriaceous.
15. Leafapexrounded;reticulationintricate(Trinidad;Venezuela;Guianas;Bra-
zil, Amazonia). 69. L. heteromorpha.
15. Leaf apex acuminate;reticulationlax (Amazonia). 60. L. reticulata.
10. Flowersborne mainly on primarybranchesof inflorescenceor on secondarybranches,but not in
pedunculatecymules.
16. Leaves narrowlyoblong with nearly parallelsides; stamens usually slightly exceeding calyx
lobes; mouth of receptaclefilled by a dense lanate mass; petals slightlyunguiculate.
17. Receptaclenarrowlyurceolate,densely arachnoid-pubescent; flowersin dense glomerules
on primarybranches(Guyana;Brazil,Amazonia;Peru). 61. L. arachnoidea.
17. Receptaclecampanulate,puberulous;flowersnot densely glomerulate.
18. Flowers 1.5-2 mm long; leaves to 17 cm long, usually smaller; stipules to 5 mm,
caducous(Brazil,Amazonia). 62. L. oblongifolia.
18. Flowers2.5-3 mm long;leaves usuallyexceeding16 cm in length(to 40 cm); stipules
to 15 mm, subpersistent(Guianas;Brazil,E Amazonia). 63. L. macrophylla.
16. Leaves ovate to oblong-lanceolate,but sides converging;stamens shorterthan calyx lobes;
mouth of receptaclewith short deflexedhairs only; petals with broad simple bases.
19. Exteriorof flowersand rachis and branchesof inflorescenceglabrousor sparselyhirsu-
tulous.
20. Leaf apex caudateto cuspidate.
21. Leaveselliptic, coriaceous,the apex caudate;petioles 5-7 mm long (Fr. Guiana;
Brazil,Amazonia;Colombia;Peru). 64. L. caudata.
21. Leaves narrowlyoblong, chartaceous,the apex cuspidate;petioles 1-3 mm long
(Brazil,Mato Grosso). 64.1. L. miltonii.
20. Leaf apex acute to acuminate.
22. Stipuleslargeand foliaceous,caducous;exteriorof flowersglabrous(Venezuela).
65. L. latistipula.
22. Stipulessmall, lanceolate,persistent;exteriorof flowershirsutulous.
23. Leaves 9-27 cm long, thick-coriaceous,the apex abruptlyshort-acuminate;
primaryveins plane above (Guianas;Brazil,Parf). 66. L. divaricata.
23. Leaves 7-11 cm long, membranous,the apex with a well-developedacumen
5-9 mm long;primaryveins slightlyimpressedabove (CostaRica;Venezuela;
Brazil,Para). 67. L. glabriflora.
19. Exteriorof flowersand usually rachis and branchesof inflorescencedensely puberulous
to tomentose;pubescencecompletelycoveringexteriorof calyx.

4 A third Malesian
species of Licania, L. fusicarpa (Kosterm.)Prance,probablybelongs here, but is known
only from fruitingmaterial.
SystematicTreatment 13

24. Midribbroadtowardsbase, 2-3.5 mm thick;leaves very thick-coriaceous;stipulesto

15 mm long, subpersistent;anthersdeltoid or nearlyso (Venezuela;Guyana;Brazil,
Amazon). 68. L. intrapetiolaris.
24. Midrib narrowertowards base, 1-2 mm thick; leaves membranousto coriaceous;
stipulesto 8 mm long, persistentto caducous;anthersreniform.
25. Leaf apex long acuminate,base cuneate,petioles eglandular,stipules small, ax-
illary,leaf base confluentinto petiole.
26. Leaves9-18 cm long x 4.2-7 cm broad;flowersbrownpubescent(Surinam;
Brazil,Amazonia). 69.1. L. laevigata.
26. Leaves 4-5 cm long x 2.5-3.8 cm broad; flowers gray pubescent(Brazil,
Amazonia). 69.2. L. occultans.
25. Leafapexbluntlyacuminate,base subcordate,rounded,subcuneate;petiolesusu-
ally glandular,leaf basenot confluent(Trinidad;Venezuela;Guianas;Amazonia).
69. L. heteromorpha.
2. Petals absent. Sect. Hymenopus.
27. Leaves thin and membranous,the base cuneate, the venation equally prominent on both
surfaces;receptacleurceolate(E-centralBrazil). 70. L. glazioviana.
27. Leavesthick and coriaceous,the base usuallyroundedto cordate,rarelyrounded-subcuneate
(to cuneate in L. marleneae),the venation obscure on upper surface;receptacleconical to
globose-cupuliform.
28. Leaves roundedto obtuse at apex, rarelyexceeding9 cm in length.
29. Flowersca. 3 mm long; petioles terete (E Brazil). 71. L. littoralis.
29. Flowersca. 2 mm long; petioles usuallycanaliculate.
30. Inflorescenceand flowerswith brownpubescence;stamens 3; venation of lower
surfaceof leaf often papillose;stipulesintrapetiolar(Venezuela;Guyana).
72. L. fanshawei.
30. Inflorescenceand flowersgray-puberulousor glabrescent;stamens5; venation of
leaf lower surfacesmooth-papillose;stipulesadnateto extremebase of petiole.
31. Leaf 2.5-6.5 cm long, the apex and base rounded(Guianas). 73. L. irwinii.
31. Leaf 5.5-9.5 cm long, the apex acute, the base cuneate(Brazil,centralAma-
zonia). 73.1. L. marleneae.
28. Leaves distinctlyacuminateat apex, usuallyexceeding8 cm in length.
32. Midribslightlyimpressedabove; petioles canaliculate(FrenchGuiana).74. L. cyathodes.
32. Midribprominulousabove; petioles terete.
33. Flowers2.5-3.5 mm long, ferrugineous-brown-pubescent; stipules4-7 mm long,
persistent(Venezuela;Guianas;Brazil,Amazonia). 75. L. polita.
33. Flowers1.5 mm long,gray-brown-pubescent; stipulessmall,caducous(Colombia;
Venezuela;Brazil,Para). 76. L. silvae.
1. Leafundersidepubescent,denselyarachnoid-lanateor pulverulent-farinaceous-pubescent (not hirsute);
petals absent.
34. Flowersin small cymules, on long slendersecondarybranches(peduncles)less than 0.5 mm thick
and attachedto primaryinflorescencebranches;pedicels usually 0.25-3 mm long, rarelyabsent;
fruit often very small, usuallynot exceeding2 cm in length. Sect. Cymosa.
35. Leaf undersidewith stomatalcavities or thick, coarse,deeply cut venation, lanate-arachnoid.
36. Bracteoleslarge and enclosing groups of flower buds; flowers subsessile;peduncles of
cymules short and ratherthick, 1-3 mm long (Venezuela;Guianas). 7. L. densiflora.
36. Bracteolessmall and not enclosinggroupsof flowerbuds; flowerspedicellate;peduncles
of cymuleslong and slender.
37. Inflorescenceand exterior of flowers ferrugineous-tomentose; young stems ferrugi-
neous-tomentellous(fruit3-5 mm long, stipitate,densely tomentose, in L. cuprea).
38. Flowersca. 2 mm long; stipulesca. 5 mm long; stamens 3 (Guyana). 78. L. cuprea.
38. Flowers ca. 6 mm long; stipules 7-10 mm long; stamens 6-7 (Brazil,Espirito
Santo). 78.1. L. arianeae.
37. Inflorescenceand exteriorof flowersgray-puberulousor gray- or brown-tomentose;
young stems glabrousor puberulous(fruit 1-2 mm long, not stipitate).
39. Midrib impressedfor entire length;petioles 7-14 mm long; leaves oblong-lan-
ceolate (Brazil,Amazonia). 79. L. impressa.
39. Midribplanethroughoutor impressedat baseonly;petioles2-10 mm long;leaves
ovate to oblong.
40. Leafapex rounded;leaves orbicular-ovate,petioles 8-10 mm long; stamens
7 (E Brazil). 80.1. L. santosii.
40. Leafapex acute to acuminate;leaves elliptic;petioles 2-7 mm long;stamens
3-7.
14 Flora Neotropica

41. Leaf undersidewith lanate pubescenceobscuringvenation;stamens 6-

7 (centralBrazil). 80. L. dealbata.
41. Leaf undersidepubescentonly in the mouth of stomatal cavities, the
reticulationsglabrous;stamens 3-5.
42. Leaves ovate, broadestnear base, 3-8.5 cm long; stipules axillary
(widespread). 88. L. hypoleuca.
42. Leaves oblong-elliptic,broadestabout middle, 5-13 cm long; sti-
pules adnate to petiole base (Venezuela;Guianas; Brazil, Ama-
zonia). 81. L. pallida.
35. Leaf undersidewith a very fine, plane or prominulousvenation, usuallypulverulent-farinaceous.
43. Exteriorof flowersand branchesof inflorescenceentirelyglabrousor glabrescent;fruitexterior
often dryingpurple.
44. Leaves oblong-lanceolateto oblong, the lower surfacewhite-lanate(Brazil,Amazonia).
82. L. gracilipes.
44. Leaves ovate to oblong, the lower surfacesparselygray-pulverulent-farinaceous (Vene-
zuela;Guianas;Brazil,Amazonia). 83. L. parvifructa.
43. Exteriorof flowersand usuallybranchesof inflorescencepuberulousto tomentose;fruitexterior
grayto brown.
45. Leaf apex rounded,the marginsrevolute;inflorescenceglabrous(Brazil,Para, Bahia).
84. L. cymosa.
45. Leaf apex acuminate,the marginsnot revolute;inflorescenceusually puberulousor to-
mentose.
46. Fruit elongate-pyriform,to 2.5 cm long, the exterior with short rufous-velutinous
pubescence;eitherinflorescenceandexteriorof flowersbrown-tomentellousor petioles
7-12 mm long.
47. Petioles3-6 mm long,terete;inflorescenceand flowersbrown-tomentellous; leaves
triangularor nearlyso (W Indies). 85. L. ternatensis.
47. Petioles 7-12 mm long, canaliculate;inflorescenceand flowersgray-puberulous;
leaves oblong (Trinidad;Guianas;Amazonia). 86. L. membranacea.
46. Fruit ovoid to pyriform,rarelyexceeding 1.2 cm long; exteriorof flowersgray-pub-
erulousor glabrescent;petioles 3-6 mm long.
48. Leaves 6.5-16 cm long; midrib and primaryveins prominenton upper surface;
rachisand branchesof inflorescenceglabrescent(Brazil,Amapa). 87. L. piresii.
48. Leaves 2.5-10 cm long; midrib plane or prominulouson upper surface;rachis
and branchesof inflorescencepuberulous.
49. Inflorescencespreading,the rachis 1-15 mm thick; lower leaf surfacepul-
verulent(Venezuela). 87.1. L. furfuracea.
49. Inflorescencecompact,the rachis0.5 mm thick;lower surfacetomentellous
(widespread). 88. L. hypoleuca.
34. Flowerssessile or subsessileon primarybranchesof inflorescenceor on short (less than 0.5 mm long)
thick pedunclesonly; fruit rarelyless than 2 cm long.
50. Leaf undersidepulverulent-farinaceous. Sect. Pulverulenta.
51. Leaf with roundedto acute apex, the marginsusuallyrevolute.
52. Flowers 1.5-2 mm long; receptaclecampanulate-cupuliform.
53. Young branchesand inflorescencedensely tomentose; inflorescenceto 4 cm long,
recurved(Venezuela;Guyana). 89. L. boyanii.
53. Youngbranchesand inflorescencebranchesglabrous;inflorescenceusuallyexceeding
4 cm long, erect (Guyana). 90. L. buxifolia.
52. Flowersca. 3 mm long; receptacleurceolate.
54. Leaves predominantlyorbicular,occasionallyoblong-elliptic,the apex roundedto
retuse(NW AmazonianVenezuela;Brazil). 91. L. orbicularis.
54. Leaves elliptic, the apex acute.
55. Receptaclenarrowlyurceolate-cylindrical; calyx lobes lanceolate(Brazil,Ron-
d6nia). 92. L. niloi.
55. Receptaclebroadlyurceolate;calyx lobes deltoid (Venezuela;Guianas;Brazil,
Amazonia). 93. L. coriacea.
51. Leaf with well-developedacumen,the marginsnot revolute.
56. Flowers 3-4 mm long; receptacleurceolate.
57. Leaf apex caudate;leaves 4-5.5 cm long, chartaceous;exterior of receptaclered-
brown-pubescentcontrastingwith whitepubescenceon interiorof calyxlobes (Brazil,
Rondonia). 95.1. L. teixeirae.
57. Leaf apex acuminateor acute;leaves 4-15 cm long, usually coriaceous;exteriorof
receptacleand calyx lobes gray-pubescent.
SystematicTreatment 15

58. Venationof leaf undersideminutelyreticulate,forminga networkwith less than

0.25 mm between reticulations;reticulationapparentbecause of absence of
pubescenceon veins; leaves thin-chartaceous;stipules usuallycaducous(Peru;
Colombia;Brazil,Amazonia). 94. L. urceolaris.
58. Venation and reticulationcoarse, with 1-2 mm between reticulations;pubes-
cence obscuringmuch of veins; leaves thick-coriaceous;stipulespersistent.
59. Leaf apex with finely pointed, well-developedacumen;rachis of inflores-
cence glabrous(CostaRica; Panama;Guianas;Brazil,Para). 95. L. affinis.
59. Leafapexacuteorwith shortbluntacumen;rachisofinflorescencepubescent
(Venezuela;Guyana;Brazil,Amazonia). 93. L. coriacea.
56. Flowers 1.5-2 mm long;receptaclecampanulate.
60. Primaryveins slightly impressed on upper surface;fruit exterior sordid-rufous-pubescent;
branchesof inflorescencedensely tomentose to puberulous.
61. Leaves thick-coriaceous;stipulescaducous;flowersin clusterson short thick peduncles;
stamens 3 (Colombia,PacificCoast). 96. L. glauca.
61. Leaves thin, chartaceous-membranous; stipulespersistent;flowerson primaryand sec-
ondarybranchesof inflorescence;stamens 6-7 (Guianas;Brazil,Amapa, Parf).
97. L. davillifolia.
60. Primaryveins plane or prominenton upper surface;fruit exteriorglabrous,dryingyellow;
branchesof inflorescenceglabrousto puberulous.
62. Leaves (11-)13-18 cm long, 4-8 cm broad(Guianas;Brazil,Amazonia). 98. L. elliptica.
62. Leaves 4-10(-12) cm long, 2-5.5 cm broad (Venezuela;Guianas; Brazil, Amazonia;
Bolivia). 99. L. canescens.
50. Leaf undersidedensely lanate-arachnoidor with stomatalcavities, never pulverulent.
Subg.Licania sect. Licania.
63. Flowers6-7.5 mm long, stamensoften connate for half their lengthor free.
64. Stamens connate for half their length;leaves prominentlyreticulatebut without stomatal
cavities;petioles eglandular(Guyana;Surinam). 100. L. couepiifolia.
64. Stamensfree to base; leaves with conspicuousstomatalcavities;petioles with 2 glandsnear
base of lamina (Brazil,Amapf). 101.1. L. naviculistipula.
63. Flowersnot exceeding5.5 mm in length;stamensfree almost to base.
65. Leaf base distinctlycordateor subcordate;leaves usuallyovate-orbicular.
66. Leavestriangular-ovate,10-16 cm long, membranous(Peru,Loreto). 101. L. trigonioides.
66. Leaves orbicularto ovate, 3-9 cm long, usuallycoriaceous.
67. Young stems hispid; lower surfaceof leaves with hirsutulous-hispidvenation, the
apex with well-developedacumen (Venezuela,Amazonas). 102. L. cordata.
67. Young stems puberulousto tomentose;lower surfaceof leaves glabrousor lanateon
venation, the apex acute or bluntlyacuminate.
68. Flowers4.5-5.5 mm long;receptacleurceolate;stipules5-6 mm long(Venezuela,
Amazonas). 103. L. foldatsii.
68. Flowers 1.5-3.5 mm long; receptaclecampanulate;stipules 1-3 mm long.
69. Leaves submembranous,the lower surfacewith deeply cut venation, the
pubescenceoccurringin cavities and hardto remove;fertilestamens5-11.
70. Stamens 8-11; petioles ca. 5 mm long; stipules 3-6 mm long (Ama-
zonian Colombia;Venezuela;Brazil). 140. L. mollis.
70. Stamens5-6; petioles 1.5-3 mm long; stipules 2-2.5 mm long (Ama-
zonian Venezuela;Colombia). 104. L. hebantha.
69. Leavesthick-coriaceous,the lowersurfacewith shallowvenation,not form-
ing cavities, the lanate pubescenceeasily rubbedoff; fertilestamens 3.
71. Pubescenceobscuringvenation;stipulesadnateto extremebase of pet-
iole (Venezuela,Bolivar). 105. L. steyermarkii.
71. Pubescencenot obscuringvenation;stipulesaxillary(Venezuela).
106. L. subrotundata.
65. Leaf base roundedto cuneate;leaves only rarelyovate-orbicular.
72. Midriband primaryveins distinctlyimpressedon leaf uppersurface.
73. Leaf undersidewith well-developedstomatalcavities filled with lanatepubescence;
petioles 8-17 mm long.
74. Stomatalcavities conspicuousbecauseof glabrousnerves and veins; leaves 4-
8.5 cm long (Venezuela;Brazil,Amazonas). 107. L. crassivenia.
74. Stomatalcavities less conspicuousbecauseof puberulousnerves and venation;
leaves (8-)10-25 cm long.
75. Flowersca. 1.5-2.5 mm long;inflorescencemuchbranched,predominantly
axillary;fertilestamens 3 with 4 sterilestaminodes;petioles conspicuously
2-4-glandular(Surinam). 108.1. L. jimenezii.
16 Flora Neotropica

75. Flowers3-5 mm long;inflorescencelittlebranched,terminal;fertilestamens

6-11, staminodesabsent;petiole glandsinconspicuous.
76. Receptacleconical,4-5 mm long;fruittomentellous,the stipe 2-6 mm
long; leaf undersidebrown-lanate(Guianas;Brazil,Amazonas).
108. L. majuscula.
76. Receptaclecampanulate,2.5-3 mm long; fruit pulverulent,the stipe
8-15 mm long;leaf undersidewhite-lanate(Venezuela;Guianas;Brazil,
Amazonas). 109. L. alba.
73. Leaf undersidewithout stomatal cavities but often prominentlyreticulate;petioles 2-6(-8) mm
long.
77. Exteriorof receptaclesparselypuberulous,the pubescencenot completelycoveringthe surface;
leaves oblong-lanceolate(widespread). 114. L. kunthiana.
77. Exteriorof receptacledenselytomentoseto tomentellous,the pubescencecompletelycovering
the surface;leaves oblong to elliptic.
78. Leaves 2.5-5.5 cm long, not prominentlyreticulatebeneath;petioles 1.5-3 mm long
(Venezuela,Bolivar). 110. L. hitchcockii.
78. Leaves 5-22 cm long, prominentlyreticulatebeneath;petioles 4-8 mm long.
79. Flowers ca. 3.5-5 mm long; receptaclebroadly cupuliform;leaves orbicularwith
roundedapex, the undersidehirsutealong venation (Guyana). 111. L. sandwithii.
79. Flowers 2-3 mm long; receptaclecampanulate;leaves oblong to elliptic, acute to
acuminateat apex, the undersideusuallytomentellous,rarelyhirsuteon venation.
80. Leafundersurfacewith parallelsecondaryveins, givinga prominentlyreticulate
appearance;pubescencebrown;primaryveins 7-10 (Venezuela;Guianas;Brazil,
Amazonia). 112. L. laxiflora.
80. Leaf undersurfacewith diffusesecondaryveins, less prominent;pubescenceru-
fous; primaryveins 5-6 (Venezuela;Guianas). 113. L. rufescens.
72. Primaryveins and usuallymidrib plane or prominenton uppersurface.
81. Exteriorof flowersand inflorescencebranchesgray-puberulous,the pubescencenot completely
coveringsurfaces.
82. Receptaclecampanulate;leaves without stomatalcavities (widespread). 114. L. kunthiana.
82. Receptacleurceolate;leaves with shallow stomatalcavities (Brazil,Rond6nia).
115. L. bellingtonii.
81. Exteriorof flowersdensely tomentellousor tomentose, the pubescencecompletelycoveringsur-
faces.
83. Leaf apex rounded,obtuse or bluntly acute, or rarelyshort-apiculate(never acuminate).
84. Lanatepubescenceof leaf undersidehard to remove, covering deep stomatal cavities;
receptacleurceolate(Guyana). 116. L. compacta.
84. Lanatepubescenceof leaf undersideeasily removed,revealinglittle protrudingvenation,
stomatal cavities absent;receptacleusually campanulate,urceolatein L. ovalifoliaand
L. savannarumonly.
85. Receptacleurceolate;stipules 3-5 mm long, adnateto petiole well above base.
86. Stamens 3; leaves thickly coriaceous;petioles 4-7 mm long (Guianas;Brazil,
Amapa). 117. L. ovalifolia.
86. Stamens9-11; leaves chartaceous;petioles 1.5-2.5 mm long(Brazil,Amazonas).
119.1. L. stewardii.
85. Receptacleusuallycampanulate,rarelyurceolate;stipules 1-3 mm long, axillaryor
adnateto extremebase of petiole.
87. Leaves ovate-orbicular,the apex often shortlyapiculate.
88. Lanatepubescenceobscuringvenation;stipules adnate to extremebase of
petiole (Venezuela,Bolivar). 105. L. steyermarkii.
88. Lanatepubescencenot obscuringvenation;stipulesaxillary(Venezuela).
106. L. subrotundata.
87. Leaves oblong-lanceolateto elliptic, the apex rounded,acute or acuminate.
89. Stipulesadnateto petiolebase;petioles8-12 mm long;inflorescencebranch-
es very sparselypuberulous;stamens 5 (Brazil,Bahia). 138. L. bahiensis.
89. Stipules axillary,caducous;petioles 2-7 mm long; inflorescencebranches
densely puberulousor tomentellous;stamens 3.
90. Leaves oblong-elliptic,sparselylanate, the undersideminutely reticu-
late;primaryveins 9-11; petioles4-7 mm long;receptaclecampanulate
(Colombia). 118. L. caldasiana.
90. Leavesoblongto oblong-lanceolate,the undersidedenselyfarinaceous-
lanate with obscuredvenation; primaryveins 5-8; petioles 2-3 mm
long;receptaclecampanulate-urceolate (Brazil;Venezuela,Amazonas).
119. L. savannarum.
Systematic Treatment 17

83. Leafapex distinctlyacuminateor sharplyacute.

91. Leaves 3-4 cm long, with two conspicuousglands at junction of petiole with upper surfaceof
lamina (Guyana). 120. L. microphylla.
91. Leavespredominantlylarge,exceeding5 cm, lackingconspicuousglands.
92. Stamens3.
93. Leaf undersurfacewith stomatalcavities.
94. Leaf venation glabrousor glabrescent,hence conspicuous,with glands present at
base of lower surface;leaves coriaceous;stipules less than 1.5 mm broad at base
(Peru;Brazil;ColombianAmazonia). 121. L. triandra.
94. Leaf venation pubescent,lanate pubescencecoveringentire leaf and obscuringsto-
matalcavitiesof lowersurface,lackingglandsat leafbase;leaveschartaceous;stipules
2.5 mm broadat base (Brazil,Para). 121.1. L. tocantina.
93. Leaf undersurfacewith prominentvenation but no stomatalcavities, the venation pu-
bescentand hence less conspicuous.
95. Stipulescaducous,petioles glabrousor tomentose;receptaclecampanulate.
96. Petioles tomentose when young;leaves elliptic to oblong, with well-developed
acumen;primaryveins 7-9 pairs (Venezuela;Guianas;Brazil). 122. L. discolor.
96. Petioles glabrous;leaves oblong-lanceolate,finely apiculate;primaryveins 10-
12 pairs (Brazil,Amazonia). 123. L. apiculata.
95. Stipulespersistent,petioles glabrousor puberulouswhen young;receptaclecampan-
ulate or urceolate.
97. Inflorescence8-15 cm long, spreading,with numerousprimarybranches;lower
leafsurfacebrown-lanate;receptaclecampanulate(Colombia;Venezuela;Guian-
as; Brazil,Amazonia, Bahia). 124. L. micrantha.
97. Inflorescence5-6 cm long, with a few primarybranches;lower leaf surface
whitish-gray-pubescent; receptacleurceolate.
98. Leaf apex with a finely pointed acumen;leaves thin-membranous(French
Guiana;Brazil,Amapa). 125. L. pruinosa.
98. Leaf apex acute to bluntlyacuminate;leaves coriaceous.
99. Leaves thinly coriaceous,4.5-10.5 x 2-5.5 cm; flowers4-5 mm long;
interiorof receptaclepubescenton upperportion (E-centralBrazil).
126. L. nitida.
99. Leavesthicklycoriaceous,3-5.5 x 2-3 cm; flowers2 mm long;interior
of receptacleglabrouson upperportion (Brazil,NW Amazonia).
126.1 L. aracaensis.
92. Stamens4-8(-10).
100. Leaf underside with deep, extremely conspicuous stomatal cavities, the pubescence
confinedto cavities made obvious by almost glabrousvenation.
101. Inflorescenceand flowersferrugineous-pubescent; stipules 1-2 mm long x 1-1.5
mm broad at base, persistent,ferrugineous;leaf apex acute to bluntlyacuminate
(E-centralBrazil). 127. L. riedelii.
101. Inflorescenceand flowers gray-puberulous;stipules 2-4 mm long, 0.2-0.5 mm
broad at base, persistentor caducous,pubescentbut not ferrugineous;leaf with
well-developedacumen.
102. Bracteolespersistent,lanceolate;uppersurfaceof midribimpressed;stipules
caducous(Brazil,Amazonia). 128. L. bracteata.
102. Bracteolescaducous,triangular;uppersurfaceof midribplane;stipulessmall
but persistent.
103. Inflorescencebranchesthick; flowers subsessile;fruit exterior ferru-
gineous-velutinous(Venezuela;Colombia;Guianas;Brazil,Amazon-
ia). 129. L. parviflora.
103. Inflorescencebranchesslender;flowersdistinctlypedicellate;fruitex-
teriorreddish-brown,short-pulverulent(Venezuela;Brazil,Amazon-
as). 81. L. pallida.
100. Leaf undersidewith poorly developed stomatalcavities or none, venation pubescent.
104. Petioles 1.5-2 cm long; inflorescencerachisand branchesglabrousor glabrescent
(Guianas;Brazil,Para). 130. L. robusta.
104. Petiolesto 1 cm long (to 1.5 cm in L. bahiensisand L. lamentanda);inflorescence
rachisand branchesusuallydensely tomentose or tomentellous.
105. Inflorescencepredominantlyof axillaryand terminalspikes,terminalinflo-
rescencesrarelylittle branchedor with minute spikes along rachis.
106. Leaves lanceolate, the lower surface deeply reticulate,with poorly
developedstomatalcavities (Venezuela,Amazonas;Brazil,Roraima).
131. L. lanceolata.
18 Flora Neotropica

106. Leavesoblongto ovate-elliptic,the lowersurfacenot deeplyreticulate,

lackingstomatalcavities.
107. Flowers 2.5 mm long; receptaclebroadlycupuliform;inflores-
cence puberulous;reticulatevenation of leaf undersideconspic-
uous (E-centralBrazil). 132. L. spicata.
107. Flowers 1.5-2 mm long; receptaclecampanulate;inflorescence
tomentose;venation of leaf undersideinconspicuousexcept in
L. nelsonii.
108. Flowers in short, dense, minute spikes attached to long
rachis(Surinam). 133. L. stricta.
108. Flowerssolitaryalong rachisor in dense glomerules,but
not in minute spikes.
109. Leafundersurfaceconspicuouslyreticulate;stipules
adnate to base of petiole; receptaclerufous-pubes-
cent; flowersdensely and evenly clusteredalong in-
florescencerachis;largetree. 135.1. L. nelsonii.
109. Leaf undersurfacelanate, not conspicuouslyreticu-
late; stipulesaxillary;receptaclegray-brownpubes-
cent; flowersin dense glomerulesor small groups;
small shrubto medium sized tree (15 m).
110. Leaves thin and membranous, the acumen
finelypointed;flowersin denseglomerules;in-
florescences largely axillary (Venezuela;
Guianas;Brazil,Bahia, Parf).
134. L. leptostachya.
110. Leavesthick and coriaceous,the acumenusu-
ally blunt;flowersin small groupsor solitary
but not glomerulate;inflorescenceslargelyter-
minal (Venezuela;Guianas;Brazil,Amazon-
ia). 135. L. incana.
105. Inflorescenceof terminaland sub-terminalracemosepanicles.
111. Leaves lanceolate;low shrubor subshrub(Venezuela,Amazonas;Brazil,Roraima).
131. L. lanceolata.
111. Leaves ovate to oblong-lanceolate;trees or tall shrubs.
112. Stipules 3-10 mm long and at least 1 mm broad at base, distinctlyadnate to petiole or
intrapetiolar,persistentand obvious.
113. Midribdistinctlyimpressedabove; bracteoles0.2-1.5 mm long.
114. Inflorescenceferrugineous-pubescent; petioles canaliculate;stipulesadnateto
petiole well away from axil (Bolivia;Brazil,Para). 136. L. paraensis.
114. Inflorescencegray-puberulous; petiolesterete;stipulesintrapetiolaror adnate
to inside of extremebase of petiole.
115. Leaves with a finely pointed acumen;primaryveins 11-12 pairs, the
lower surfaceglandularat base (Colombia,Amazonas;Brazil,W Ama-
zonia). 137. L. vaupesiana.
115. Leaves with blunt acumen;primaryveins 6-8 pairs, the lower surface
eglandularat base (E-centralBrazil). 138. L. bahiensis.
113. Midribplane or prominulousabove; bracteoles1.5-3 mm long.
116. Leavesmembranous;flowersca. 2 mm long, gray-tomentellous;inflorescence
gray-puberulous;stamens 5 (Brazil,Amapa). 139. L. maxima.
116. Leaves coriaceous;flowers 3-3.5 mm long, brown-tomentose;inflorescence
brown-tomentose;stamens 8-11 (AmazonianColombia;Venezuela;Brazil).
140. L. mollis.
112. Stipules usually less than (rarelyexceeding)2.5 mm long and very narrowto base, on
outside of axils or adnateto extremebase of petiole, caducousor persistent,often incon-
spicuous.
117. Stipulesadnate to extremebase of petiole, persistentor subpersistent.
118. Petioles 7-12 mm long, remainingtomentellouseven with age; laminae 7-
16 x 3-7.5 cm; stipules sub-persistent(Brazil,Para and central).
141. L. blackii.
118. Petioles 2-6 mm long, becomingglabrouswith age; lamina 3-12 x 2.2-5.5
cm; stipules persistent.
SystematicTreatment 19

119. Inflorescencelax and spreading;leaf with a finelypointed acumen 8-20 mm

long, the lower surface farinaceous-lanate,shallowly reticulate(Colombia;
Venezuela;Guianas;Brazil,Amazonia). 129. L. parviflora.
119. Inflorescenceand flowersdensely crowded;leaf with blunt acumen 2-
13 mm long, the lower surface lanate, deeply reticulate(Venezuela;
Guianasto E-centralBrazil). 114. L. kunthiana.
117. Stipulesaxillary,caducousor persistent.
120. Petioles 11-15 mm long; flowers3.5-4 mm long (Brazil,Bahia). 130.1. L. lamentanda.
120. Petioles 2-10 mm long; flowers2-3 mm long.
121. Leaf undersurfacewith smooth inconspicuousreticulation,the pubescence easily re-
moved; primaryveins widely spaced, 1.2-2.5 cm apart;exteriorof receptaclevelutinous-
pubescent(Brazil,Amazonas). 142. L. rodriguesii.
121. Leaf undersidewith deeply cut reticulation,and hence pubescencehard to remove;pri-
mary veins not more than 1 cm apart;exteriorof receptacletomentellous.
122. Receptacleglobose;upper surfaceof youngestleaves appressed-strigose,soon be-
coming glabrous;inflorescenceof axillaryspikes and terminalpanicles (E-central
Brazil). 132. L. spicata.
122. Receptacleusuallycampanulate;upper surfaceof youngestleaves glabrous;inflo-
rescenceof terminaland subterminalracemosepanicles.
123. Leavesoblong-lanceolate;midribslightlyimpressedon uppersurface(E-cen-
tral Brazil). 143. L. indurata.
123. Leaves ovate-ellipticto oblong;midrib usuallyplane on uppersurface.
124. Inflorescencemuch branched,spreading,lax; lowerleaf surfacerufous-
pubescent(Venezuela;Guianas;Brazil). 113. L. rufescens.
124. Inflorescencedenselycrowded,compact,little-branched;lowerleaf sur-
face gray-brown-pubescent.
125. Petioles4-7 mm long,terete;receptacleandcalyxlobestomentose
on exterior.
126. Leavesfinelyacuminate,the acumen8-12 mm long;stipules
caducous,ca. 1 mm long (Ecuador). 144.1. L. harlingii.
126. Leaves bluntlyacuminateto obtuse;stipulespersistent,1-
3 mm long.
127. Petioles soon becoming glabrous;leaf apex bluntly
acuminate;young fruitpyriform(E-centralBrazil).
144. L. hoehnei.
127. Petioles tomentose, becoming less so with age; leaf
apex obtuse to acute; young fruit cylindricalwhen
young, becomingpyriform(Panama;Trinidad;Ven-
ezuela). 145. L. cruegeriana.
125. Petioles 10-12 mm long, canaliculate;receptacleand calyx lobes
velutinouson exterior(Brazil,Bahia). 146. L. belemii.

Additional Notes and Descriptions Fruit globose with warty surface,9-10 cm in

of Species of Licania diam.; exocarpglabrous,crustaceous;mesocarp
1.5-1.8 cm thick, fibrous;endocarp ca. 3 mm
The notes on the species that follow referonly thick, hard and woody, glabrouswithin.
to those species for which interestingand useful Distribution(Fig. 59). Endemicto coastal for-
new data have been collected since the mono- ests of Choc6, Colombia.
graph(Prance, 1972).
Additionalspecimenexamined.COLOMBIA.CHOCO:
Mun.El Valle,Jan1985(fr),P. Perezs.n. (MEDEL).
This species was describedfrom a single flow-
Subgenus Moquilea Section Moquilea
ering specimen. The fruits are large and rather
2-3. Licania maritima Prance, Fl. Neotrop. similar to those of L. macrocarpa,and are said
Monogr.9: 44. 1972. to be edible.
20 FloraNeotropica
2-4. Licania durifolia Cuatrecasas, Fieldiana,
Bot. 27: 61. 1950.
a & so~ , This species,which has extremelylargeleaves
E
IX D IX el for the genus, was known only from two widely
0 0
separatedcollectionsin 1972. It is a high altitude
dI t0
forestspeciesthat has quite a wide rangebut still
remains poorly known.
CO i O O Distribution(Fig. 36). Montane forests 500-
e l ^ e: ^ e X
r < ,,. vE ,
2000 m in Colombia, Ecuador,and Peru.
<i _ss^'ss.g.s.ili E Additional examined.
specimens ECUADOR. 1?20'S,
~
(fr),Oldeman&Arevalo11 (QCA).
77?28'W
o PERU. HUANUCO: Prov. Huanuco W of division,
1400m, 27 Jul 1940(flbud),Asplund12538(S).
Local name. Auka Indian:maeneyowae.
? ~ CO
Licaniamacrocarpa,L. durifolia,and theirrel-
1 o 0 0 0k
&C
' , ' 0 0) 0
C atives form a complex of closely relatedspecies,
0) E 0) all of an Andeanand westerndistribution.They
are characterizedby having very long leaves (for
Licania),which aregenerallycoveredby a dense
lanatepubescence,unusualin section Moquilea.
The previously described species in this group
c X X X X X X X X
"
are L. durifolia,L. macrocarpa,L. maritima,L.
COCN ri rI - I
(- -
-C 00 Cl4 f)
velata, and L. veneralensis.Closely related but
o) (N cl Cl1 - - with smaller leaves are the two high montane
species L. cabreraeand L. montana. The prob-
lem with this group is the paucity of material
C4 If
available.There are only a few collections of all
I i I I
these species, thus still making their evaluation
0I) 0) N - n C e) n 0 difficult.However,a carefulstudyof the material
rd) - and a new comparisonwith all of the types in-
?6 oO 6 d 60
dicates that there are three new species to be
addedto this complex. These are describedas L.
filomenoi,L. gentryi,and L. grandibracteata. The
o 0) S~
C) C~
differencesbetween these species are summa-
rized in Table I.
0)
*)0 ) 0 cl Cl C CD
24.1. Licania filomenoiPrance, sp. nov. Type.
0 C C CO Peru. San Martin:Prov. MariscalCaceres, 12
km W of Tocache Nuevo, Rio Tocache, 500
m, 13 Mar 1979 (fl), Gentryet al. 25665 (ho-
lotype, NY; isotypes, F, MO). Fig. 2.
A L. durifolioinflorescentiis5-13 cm (haud
15-30 cm) longis, floribushaud agglomeratisex-
tus tomentosis, bracteolispersistentibusdiffert.
0~~~~V IAI
Treeto 8 m, the youngbranchesglabrous,len-
Tb,
d
~~oo d
In~~~ ticellate. Leaf lamina oblong-lanceolate,coria-
ceous, 20-32 x 7-10 cm, roundedat base (apices
St all damaged), glabrous above, with short gray
appressed pubescence beneath; midrib promi-
nent on both surfaces,glabrousabove, sparsely
 i.

C'

A.

r, ;'"~

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

FIG. 2. Licaniafilomenoi (Gentryet al 25665). A, habit;B, bract;C, flowerbud; D, section of flowerbud;

E, ovary and style; F, stamen;G, H, petals;I, young fruit.
21
22 FloraNeotropica

puberulousbeneath;primaryveins 21-24 pairs, conspicuous parallelreticulatepattern;petioles

slightly impressed above, prominent beneath; 8-12 mm long, puberulous,terete, eglandular.
secondaryveins more or less parallel on lower Stipulesintrapetiolar,ca. 2 mm long, persistent.
surface, forming a conspicuous reticulate pat- Inflorescencesramiflorous,little-branchedrace-
tern; petioles 6-8 mm long, terete, eglandular, mose panicles, to 15 cm long, the rachis and
sparselypuberulous.Stipulesnot seen. Inflores- branchesbrown-shaggy-tomentose. Bracts4-4.5
cencesof unbranchedracemesor little-branched cm long, membranous,glabrousexcept for mar-
panicles, 5-13 cm long, axillaryor subterminal, gins. Flowers 5-6 mm long, solitary and not
the rachisand branchesferrugineoustomentose. densely crowdedon inflorescencebranches.Re-
Bractsat base of inflorescencelarge and persis- ceptaclecupuliform,sessile, sparselypuberulous
tent, 25-35 cm long, chartaceous;bracteoles 5- on upperpartof exterior,tomentoseonly around
15 mm long, shortly tomentellous on exterior, base, denselytomentosewithin. Calyxlobes five,
glabrescentwithin. Flowersca. 5 mm long, sol- acute, puberulous on exterior, tomentellous
itaryalongrachisand primarybranchesof inflo- within. Petals five, white. Stamens ca. 40, in-
rescence. Receptacle cupuliform, sessile, ferru- sertedaroundcomplete circle, exserted,connate
gineous-browntomentose on exterior, densely at extremebase, with dense villous mass at base
tomentose within. Calyx lobes five, acute, to- of filaments.Ovaryinsertedat base of receptacle,
mentose on exterior. Petals five, white, tomen- lanate. Style lanate for two-thirdsof length, ex-
tellous on exterior,glabrouswithin. Stamensca. ceeding filaments.Fruit not seen.
35, insertedaroundcompletecircle,the filaments Distribution(Fig. 42). Known only from An-
exserted (seen only in buds). Ovary inserted at dean foothills of Ecuador.
base of receptacle,villous-pubescent.Style hir- Habitat. Primaryforest.
sute on lower portion. Youngfruit globose, exo-
Additional specimens examined. ECUADOR.
carp lenticellate. AZUAY:Between Rio Blanco and Rio Norcay on rd.
Distribution(Fig. 38). Known only from the Chancanceo
to Molleturo,1520m, 4 Jun 1943 (st),
type collection from San Martin, Peru. Steyermark52829 (F, NY). PICHINCHA: Reserva Fo-
Habitat.Matureflatlandforeston lateriticsoil. restalEndesa,Rio Silanche,0?5'N,79002'W,
650-700
This species is closely relatedto Licania duri- m, 10 Jun 1984(st),Jaramillo6672 (NY, QCA),14
Aug 1984 (st),Jaramillo6960 (NY, QCA),18 Mar
folia, with which it shares a similar parallelre- 1985 (st), Jaramillo 7527 (NY, QCA).
ticulatevenationpatternof the leafundersurface,
but from which it differs in the much smaller Closely relatedto Licania durifoliaand L. fi-
inflorescence,less densely crowded flower, and lomenoi,this speciesshareswiththem leaveswith
persistentbractsand bracteoles.It is named for a graypubescenceand conspicuousparallelsec-
FilomenoEncarnaci6n,botanistof Iquitos,Peru. ondary venation. It differs from L. durifoliain
the smaller unbranchedinflorescencewith less
24.2. Licania grandibracteataPrance, sp. nov. dense flowers,the persistentbracteolesand the
Type. Ecuador.Pichincha:Reserva Florestal sparsepubescence of the receptacle.It differsfrom
Endesa, Rio Silanche, 0?5'N, 79?02'W,650- both species in the ramiflorousinflorescenceand
700 m, 7 Dec 1984 (fl),J. Jaramillo 7413 (ho- from L. filomenoi in the much sparser pubes-
lotype, NY; isotype, QCA). Fig. 3. cence of the inflorescenceand flowers, and the
largermembranousbracteoles.
A L. durifolioinflorescentiisminoribus,rami-
floris,floribushaudagglomeratis,receptaculoex-
tus sparsepuberulo,bracteolispersistentibusdif- 2-5. LicaniamacrocarpaCuatrecasas,Fieldiana,
fert. Bot. 27: 107. 1951.
Treeto 30 m tall, the youngbranchesglabrous,
Distribution(Fig. 59). Lowlandforeststo 600
conspicuouslylenticellate. Leaf lamina oblong, m in
Colombia, Ecuador,and Peru.
coriaceous,31-33 x 7-13.5 cm, roundedat base,
bluntlyacute at apex, with short appressedgray- Additionalspecimensexamined. COLOMBIA. CHocO:
brownpubescencebeneath;midribprominulous 3 km W of Tutunend6,7 Jan 1981 (st), Gentryet al.
30324(MO,NY).
above, prominentbeneath;primaryveins 21-24 ECUADOR. LOS RIOS-PICHINCHA: El Centinela,
pairs,impressedabove, prominentbeneath;sec- Montaiiasde Lla, 6 Feb 1979 (fl), Dodson et al. 7516
ondary veins more or less parallel, forming a (MO), 2 Oct 1979 (fr),Dodson et al. 8672 (MO, NY);
Systematic Treatment 23

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

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

~~~r- I
r~~yr I ~ ~ ~ ";.i~~Tki~~e~~!'-?~~;~;.
? ~ ~ no

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

flwr etin F vayan tye G etl
24 FloraNeotropica
CerroAntisana,NE of Borja(fl), Grubbet al. 1045 a very small ovary at the base, and the greater
(NY). numberof stamens.It is the largest-leavedspecies
PERU:PASCO: Prov.Oxapampa, PalcazuValley,Is-
cozacin,22 Jan1984(fl),Foster7961(MO,NY). of Licania so far described.It is related to the
other large-leaved,lanate pubescent species of
2-5.1. LicaniagentryiPrance,sp. nov. Type. Co- Licania subgenusMoquilea,but is quite distinct
lombia. Valle: Bajo Calima, 15 km N of Bue- from them by the leaf size and shape and the
naventura, 3?56'N, 77?08'W, 50 m, 16 Feb ramiflorousnatureof the inflorescence.
1983 (fl), A. Gentry et al. 40355 (holotype,
2-5.2. Licania cabrerae Prance, Brittonia 28:
MO; isotype, NY). Fig. 4.
210-212, fig. 1. 1976. Type. Colombia. An-
Species a L. cabreraeaffinis, foliis 27-47 cm tioquia: Estaci6n ExperimentalForestal Pie-
longis x 22-26 cm latis, inflorescentiisrami- dras Blancas, 13 Jul 1957 (fl), I. CabreraR.
floris,receptaculocrasso,staminibuscirca60 dif- 94 (holotype, COL).
fert.
Tree 10 m tall. Leaf lamina elliptic, charta- Small tree, the young branchesferrugineous-
ceous, 27-47 cm x 22-26 cm, roundedat base, lanatepubescent,soon glabrate.Leaflamina ob-
abruptlymucronateat apex, the mucro 5-7 mm long to oblong-lanceolate,coriaceous, 12-17 x
long, densely ferrugineous-lanate-pubescent be- 3-4.5 cm, subcuneateat base, acuminateat apex,
neath, lanate above when young, becoming gla- the acumen8-15 mm long, lanate-tomentoseon
brous with age, the pubescence persisting only upper surfacewhen young, soon becoming gla-
along veins; midrib prominent beneath, plane brous, ferrugineous-lanate-tomentosebeneath
above, with a ferrugineous-lanate-pubescence when young, becoming gray-tomentellouswith
contrastingwith the caducouslighterpubescence age; midrib prominulous above, lanate when
of uppersurfaceand lateralveins; primaryveins young, prominent beneath, gray-tomentellous;
25-29 pairs, prominent beneath, slightly im- primaryveins 16-19 pairs, plane above, prom-
pressed above; petioles 20-25 mm long, ferru- inent beneath; petioles 10-12 mm long, terete,
gineous-lanate-pubescent, terete, eglandular. lanate-tomentose when young. Stipules mem-
Stipulesaxillary,ca. 12 mm long, membranous, branous,triangular,caducous. Inflorescencesof
sparselylanateon exterior,glabrouson innersur- racemosepaniclesto 18 cm long, the rachisand
face. Inflorescencesof panicles borne on the branchesdenselyferrugineous-tomentose; bracts
woody branches, the rachis and branches to- and bracteoles triangular,membranous, ferru-
mentellous;bractslargeand membranousto 20 gineous-pubescent,glabrouson innersurface,re-
mm long, persisting at base of inflorescence; flexed, caducous.Flowerbuds enclosed by large
bracteolesmembranous,to 20 mm long, tomen- navicular,amplexicaulbracts.Flowers 6-7 mm
tellous on exterior,glabrouswithin. Flowers 6- long. Receptacleglobose, ferrugineous-lanateon
7 mm long, sessile. Receptaclecupuliform,fer- exterior,witha thickrowofdeflexedhairsaround
rugineous-lanate-pubescent on exterior,the walls faucalannuluswithin;pedicels ca. 0.5 mm long.
thick, restricting the inner cavity, tomentose Calyx lobes five, triangular,acute, ferrugineous-
within. Calyxlobes five, triangular,lanateon ex- lanateon exterior,brown-tomentosewithin.Pet-
terior, tomentellous within. Petals five, white. als five, white,tomentellouson exterior,glabrous
Stamensca. 60, insertedin severalrows around within. Stamens 35-40, inserted in a complete
a complete circle, the filaments glabrous, ex- circle,the filamentsshortlyexserted,freeto base,
ceeding calyx lobes, free to base, with a dense with a few scatteredhairs on lower portion, pli-
ring of reflexedhairs aroundbase on interiorof cate in bud. Ovarypilose. Style pilose almost to
receptacle.Ovary insertedat base of receptacle, apex. Fruit unknown.
small, tomentose. Style longer than filaments, Distribution(Fig. 30). Known only from the
villous for three-fourthsof length.Fruitnot seen. type gatheringcollectedin flowerin July. A high
This species (distribution in Fig. 40), most altitude species, the type from 2550 m.
closely relatedto the highlandLicania cabrerae,
differsin the verymuchlargerandbroaderleaves, 2-5.3. Licania fasciculata Prance, Acta Ama-
in the inflorescenceborne on woody branches, z6nica8: 579-581, fig.2. 1978. Type. Panama.
the muchthickerwallsof the receptaclewith only Colon: Zona de Santa Rita, 31 Aug 1972 (fl),
SystematicTreatment 25

"-~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~
'
.. ......
'i~~~~~~~~~~~~~~~~~~~~~~~~~~~'
i
IVI]'

D.~~~~~~~~~~~~~~?:
o~.:: . .
rr
'~~~~~~~~~~~~~~~~~,?'
E.?

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

X C.

3
.i
enmI '
.ri}~!
'~.i[11.'I
i i'i :
:

FIG. 4. Licania gentryi (Gentry40355). A, leaf and leaf undersurface,B, inflorescence;C, flower bud; D),
flower;E, petals, left inner surface,right outer surface;F, flowersection.
26 FloraNeotropica

Correa A. & Dressier 1815 (holotype, MO; 1975 (fl),Steyermark,R. Smith & C. Espinoza
isotype, NY). 111541 (holotype, NY; isotype, VEN).
Tree 12 m tall, the young branchestomentel- Treeto 25 m tall, the young branchessparsely
lous, becomingglabrousand prominentlylenti- tomentose soon becoming glabrous and lenti-
cellate with age. Leaf lamina oblong to oblong cellate. Leaf lamina, narrowly elliptic, charta-
lanceolate, thickly coriaceous, 9-13.5 x 2.8-4 ceous, 6-9.5 x 2-3 cm, subcuneateat base, acu-
cm, cuneateat base, acuminateat apex, the acu- minate at apex, the acumen 3-5 mm long,
men 4-7 mm long, abrupt,glabrousabove, with glabrousabove, shortly appressed-tomentellous
a compactappressedgray-lanatepubescencebe- beneath;midrib prominulousabove, pubescent
neath; midrib prominulous and tomentellous towardsbase only, prominentbeneath, sparsely
above, prominentbeneath;primaryveins 15-19 lanate-arachnoid-pubescent; primaryveins 15-
pairs, prominent beneath, slightly impressed 17 pairs,prominulouson both surfaces;petioles
above; petioles 5-10 mm long, tomentellous,te- 4-5 mm long, terete, tomentellous, eglandular.
rete. Stipules axillary, 3 mm long, persistent, Stipules caducous (not seen). Inflorescencesof
membranous,puberulous.Inflorescencesof short racemoseterminaland subterminalpanicles, 8-
fasciculateracemes 1-2.5 cm long with flowers 12 cm long;the rachisand branchesferrugineous
densely clustered,completely obscuringthe ra- tomentose. Bractsand bracteolesmembranous,
chis. Bractsand bracteoles4-7 mm long, ovate, caducous, tomentose on exterior, sparsely pu-
tomentellous, subpersistent. Flowers 6-7 mm bescentwithin.Flowers4-5 mm long. Receptacle
long. Receptacle cupuliform,tomentose on ex- globose, sessile, ferrugineous-tomentoseon ex-
terior,tomentosewithin. Calyxlobes five, acute, terior,tomentose within, with a row of deflexed
tomentellouson both surfaces.Petalsfive, white, hairsaroundthe faucalannulus.Calyxlobes five,
puberulouson exterior.Stamensca. 60, inserted acute, triangular,ferrugineous-tomentoseon ex-
in complete circle; filaments far exceeding the terior, brown-tomentellouswithin. Petals five,
calyx lobes, glabrous,united at base, the basal tomentellous on exterior,tomentellous towards
fused portion bent inwards, villous. Ovary in- apex and glabroustowardsbase on innersurface,
serted at base of receptacle,pilose. Style villous the margins ciliate. Stamens 30-35, inserted
for two-thirdsof length. Fruit not seen. around a complete circle, the filaments shortly
Distribution. Figure 38. exserted,freeto base. Ovarylanate.Style lanate-
Additionalspecimenexamined.PANAMA.PANAMA: pilose equalling filaments in length. Fruit un-
El Llano-CartiRd., 7.5 km N of PanamericanHwy., known.
23 Jan1977(fl),Folsom1435(NY). Distribution(Fig. 65). A cloud forest species
collectedat 1300-1500 m; still knownonly from
Licaniafasciculatadiffersfromall otherspecies the type gathering,collectedin flowerin March.
in the section by the fasciculate,densely flow-
ered, racemose inflorescence.It is most closely
related to the species group of L. maritima, L. 2-7. Licania klugii Prance,Fl. Neotrop. Mono-
cabrerae,L. durifolia, L. montana, L. macro- gr. 9: 47. 1972.
carpa, and L. veneralensis,and has the same pu- Additional materialof this species, originally
bescenceand venation patternof the leaf under- based on a single collection, shows it to be quite
surface.It differsin the inflorescenceand in the distinct.
smallerleaves with fewer primaryveins, which
are impressedon the uppersurface.Licaniafas- Additionalspecimensexamined.PERU. LORETO:
ciculata also has a greaternumber of stamens Maynas, RioMom6n,24 Nov 1977(y fr),RimachiY.
3274 (NY). SAN MARTIN: QuebradaHicte, Rio Hua-
than the other species listed above. It is a most llaga,26 May1964(fl),SchunkeV.6502(F).
distinctspecies,not easilyconfusedwith any oth-
er in the genus.
2-9. Licania retifolia Blake, Contr. Gray Herb.
2-5.4. LicaniamontanaPrance,Brittonia28:212, 52: 66. 1917.
fig. 2. 1976. Type. Venezuela. Lara:Vic. La- This poorlyknown specieswas representedby
guna Negra, Loma de Los Naranjos, 24 Mar the type from Guerrero,Mexico and one other
SystematicTreatment27

collection in Prance (1972). Recent collections panulate,tomentellous on exterior, densely to-

have extended the rangeto El Salvador. mentose within; pedicels 1-2 mm long, articu-
Distribution.Figure 73. late.Calyxlobes five, acute,tomentellouson both
surfaces. Petals five, white, glabrous. Stamens
Additional specimens examined. MEXICO.
GUERRERO:Rinc6n Viejo, 15 Apr 1960 (fl), Kruse261 30-35, insertedaround complete circle, the fil-
12
(ENCB), Apr 1964 (fl),Kruse1256 (ENCB). aments exserted. Ovary inserted at base of re-
ELSALVADOR.AHUACHAPAN: CerroLa Piedradel ceptacle,tomentose. Style hirsuteon lower half.
Filo, 13?51'N,89?56'W,30 Jan 1980(fl), Witsberger Fruit not seen.
813 (NY).
Habitat. Tropicalrain forest.
This can be a largetree. Additionalspecimenexamined.ECUDAOR.ESME-
Local name. El Salvador:mulo. RALDAS: Lita,600 m, 20 May1987(fl),VanderWerff,
Dodson&Palacios9540(NY,MO).
2-10. LicanialongipedicellataDucke,Bull.Mus.
Hist. Nat. (Paris),s6r. 2, 4: 725. 1932. This species belongs to section Moquileaand
is most closely related to Licania leucosepala,
Distribution.Figure 57. butdiffersin the muchbroaderleaveswithwidely
Additional specimensexamined. PERU.AMAZONAS:spaced secondary veins and in the loosely
RioSantiago, 17Sep1979(y fr),Huashikat branchedinflorescencewith articulationsat each
Caterpiza,
662 (MO,NY). branch quite unlike most species of Licania,
BRAZIL.AMAZONAS: Tabatinga,Sao Leopoldo, 13 whichhave racemosepanicles.L. dodsoniiis also
Oct 1976(fl),Bragaet al. 3127(INPA). close to L. minutiflorabut differs in the inflo-
Local name. Peru:yukuku. rescence,the broaderleaves that are subcuneate
ratherthan roundedat the base, and acuminate
2-13.1. Licania dodsoniiPrance,sp. nov. Type. rather than cuspidate at the apex, and in the
Ecuador.Esmeraldas:Mun. de Lita, 19 km N greaternumberof stamens.
of Lita, road to San Lorenzo, ca. 650 m, 10
May 1987 (fl),Acevedo&Daly 1657 (holotype, 2-14.1. LicaniaanneaePrance,Brittonia31: 250,
NY). Fig. 155. fig. 2. 1979. Type. Brazil. Para: Cuiaba-San-
taremHwy., km 1305, vie. Igarap6Jose Preto,
Species a L. leucosepalaaffinis, foliis latiori- 22 Nov 1977 (fl), Prance et al. 25652 (holo-
bus, nervis secundariis8-19 mm distantibus,in-
florescentiislaxis multiramosis,ramisarticulatis type, MG; isotypes, AAU, NY).
differt. Tree7 m tall, the youngbranchessparselypu-
Tree 10-15 m tall, the young branches gla- berulous,soon becomingglabrous.Leaf lamina
brous, conspicuously lenticellate. Leaf lamina oblong-lanceolate, coriaceous,14-25 x 4.5-8 cm,
elliptic,chartaceousto thinly coriaceous,9.5-17 roundedat the base, shortlyacuminateor acute
x 5.5-10.5 cm, rounded to subcuneate at base, at the apex (acumen 2-6 mm long), glabrous
abruptlyacuminateat apex,the acumen3-5 mm above, pilose with a few sparseappressedhairs
long, glabrouson both surfaces;midrib plane to beneath,especiallyon the primaryveins; midrib
prominulous above, prominent and appressed glabrousandprominulousabove, prominentand
villous pubescentbeneath;primaryveins 13-15 puberulousbeneath;primaryveins 18-22 pairs,
pairs, plane to slightly impressedabove, prom- slightly impressed above, prominent and pu-
inent beneath; petioles 5-7 mm long, terete, berulous beneath, arcuate and conspicuously
eglandular, sparsely puberulous, glabrescent. anastomosingca. 2-3 mm from the margin;pet-
Stipules not seen. Inflorescences of much iole 5-7 mm long, rugulose,slightlycanaliculate,
branched panicles, the primary branches with puberulouswhen young. Stipulescaducous(not
short thick secondarybranches bearing groups seen). Inflorescencesof terminaland axillaryra-
of 2-3 flowers,the brancheswith articulationsat cemose panicles, the rachis and brancheslight
junctions, the rachisand branchesappressedto- brown-tomentellous.Bracts 5-18 mm long, ob-
mentellous. Bracts and bracteoles minute, tri- long to lanceolate,membranous,persistent,to-
angular,puberulous,caducouswhile in earlybud. mentellouson the exterior,sparselypuberulous
Flowers5-6 mm long. Receptaclebroadlycam- within; bracteoles ovate, persistent, membra-
28 FloraNeotropica

nous, glabrous or with a few sparse hairs and spicuously lenticellate. Leaf lamina oblong-el-
ciliate margins.Flowers3-3.5 mm long, inserted liptic, subcoriaceous,5-8.5 x 2.8-3.8 cm, sub-
on primary branches of the inflorescence.Re- cuneateat base, acuminateat apex, the acumen
ceptaclecupuliform,extremelyregularand round 5-6 mm long, glabrous, midrib prominulous
at the apex, short brown-tomentellouson exte- above, prominentbeneath;primaryveins 8-11
rior,tomentellouswithin. Calyxlobes five, acute, pairs, prominulous above, prominent beneath;
triangular,small, tomentellouson both surfaces, petioles 5-7 mm long, glabrous, terete, eglan-
but more sparsely on the inner surface. Petals dular,with very distinct articulationat junction
five, oblong,white,glabrouswith ciliatemargins, with branch.Stipulescaducous(not seen). Inflo-
caducous. Stamens 12-14, inserted around a rescences(not fully developed in specimen ex-
complete circle, filamentsslightly exceedingthe amined) of terminal and axillary panicles, the
calyx lobes, glabrous,connate at base forminga rachisand branchesyellow-browntomentellous.
ring 1 mm tall that is hirsute on both surfaces. Bractsand bracteolestriangular,caducous,pub-
Ovary insertedat the base of the receptacle,to- erulouson exterior.Flowers3-3.5 mm long. Re-
mentose-pilose.Style lanate almost to the apex. ceptacle broadly campanulate,tomentellous on
Fruit not seen. exterior,hirsutewithin;pedicelsca. 0.5 mm long.
Distribution.Figure 23. Calyxlobes five, acute,tomentellouson exterior,
Habitat.Disturbedforestbeside road on mar- puberulouswithin. Petals five, glabrouswith cil-
ginally flooded and non-floodedareas. iate margin. Stamens 13-15, inserted around a
This speciesbelongsto subgenusMoquileasec- complete circle;filamentsslightly exserted, free
tion Moquilea, and is quite distinct within the to base. Ovaryinsertedat base of receptacle,gla-
section.It is closestto Licaniaangustata,a species brous or sparselyhispid. Style sparselyhirsute.
of western Amazonia, but differs in the much Fruit not seen.
larger leaves, the shorter petioles, the higher Distribution.Figure 33.
number of primaryveins, the largerbracts, the Habitat. Cloud forest over 1200 m.
fewerstamens,and the distinctive appressedpu-
Additionalspecimenexamined.PANAMA.PANAMA:
bescence of the lower leaf surface. CerroJefe,30 Sep1978(fl),Hammel4823(MO,NY).
2-16. LicaniagonzaleziiMiranda,Bol. Soc. Bot. Licania chiriquiensisbelongsto subgenusMo-
Mexico 29: 36. 1965. quilea section Moquileaand is most closely re-
lated to L. minutiflora,a species of the Guianas
This poorly known but distinct species was and Amazonia. It differs in the
smaller, less
described from two collections from the States branchedinflorescencewith much
largerflowers,
of Nayarit and Jalisco. A collection from Guer- the smaller leaves with
longer petioles, the al-
rero has now been added. most glabrous ovary and in the more tapered,
Distribution.Figure42. less caudateleaf apex.
Additionalspecimenexamined.MEXICO.GUERRERO:
Rincon Viejo, 700 m, 2 Apr 1961 (fl), Kruse 625
(ENCB).
2-18.2. Licania kallunkiae Prance, Acta Ama-
z6nica 8: 583, fig. 4. 1978. Type. Panama.Co-
The field notes state that the flowersare white 16n:Santa Rita Rd., 17 km from Boyd-Roo-
and very aromaticand are "pollinated"by Lep- sevelt Hwy., 450 m, 14 Mar 1975 (fl), Mori &
idoptera,Hymenoptera,and Diptera. Kallunki5052 (holotype, NY; isotype, MO).

2-18.1. Licania chiriquiensis Prance, Brittonia
29: 154, fig. 1. 1977. Type. Panama.Chiriqui:
CerroColorado,50 km N of San Felix, cloud
forest, 1200-1500 m, 17 Aug 1975 (fl), Mori
& Dressier7778 (holotype,NY; isotype, MO).
Treeto 12 m tall, the young branchessparsely
puberulous,soon becoming glabrous and con-
Tree14 m tall, the youngbranchesvery sparse-
ly puberulous,soon becomingglabrous,not con-
spicuously lenticellate.Leaf lamina oblong, co-
riaceous, 5.5-10.5 x 2.2-4.0 cm, subcuneateat
base, cuspidateat apex, acumen 6-10 mm long,
glabrouson both surfaces;midrib plane above,
prominentbeneath, glabrous;primaryveins 9-
12 pairs,almostplaneand inconspicuouson both
SystematicTreatment29

surfaces, glabrous;petioles 6-7 mm long, gla- 1971 (fl), Contreras10742 (holotype,LL;iso-

brous, canaliculate,eglandular.Stipules small, type, US).
lanceolate,puberulous,caducous.Inflorescences Tree ca. 30 m, the young branchesglabrous.
terminal and axillary panicles 5-11 cm long,
Leaf lamina elliptic, 10-21 x 5.5-9.5 cm, the
3-branched,therachisandbranchessparselygray- baseroundedto subcuneate,the apexwith a short
puberulous.Bractsand bracteolescaducous(not acumen4-7 mm long, glabrouson both surfaces;
seen).Flowers2.5-3 mm long,bornein 2-3-flow-
ered cymules attached to primarybranches by primaryveins 8-10 pairs, prominulouson both
short secondarybranchesor peduncles. Recep- surfaces, secondaryvenation prominulous and
tacle campanulate,gray-puberulouson exterior, conspicuouslyreticulateon both surfaces;midrib
tomentose within;pedicels ca. 1 mm long, gray- prominulousand flattenedon both surfaces,gla-
brous; petioles 3-5 mm long, slightly canalicu-
puberulous. Calyx lobes five, acute, gray-pu-
berulous on both surfaces. Petals five, white. late, glabrous.Stipulessmall, triangular,axillary.
Stamens 11-12, inserted in a complete circle; Inflorescencesterminal panicles to 13 cm long,
the flowersborne in small groups on short sec-
filaments exceeding calyx lobes, free to base.
ondary branches, the rachis puberulous, soon
Ovaryinsertedat base of receptacle,almost gla-
brouswith only a few hairs.Stylelanateon lower glabrescent, the branches gray-tomentellous.
Bractsand bracteolessmall, ovate, tomentellous,
portion, equallingfilamentsin length. Fruit not
seen. persistent.Flowersca. 2.5 mm long. Receptacle
campanulate, tomentellous on exterior. Calyx
Distribution(Fig. 51). Known from only one lobes small, to 1 mm long, triangular,tomentel-
collection from Costa Rica and one from Pan- lous on exterior,puberulouswithin.Petalswhite,
ama. tomentellous on exterior, sticking together and
Habitat. Wet forest.
dehiscingin a calyptra-likemass, 1.2 mm long.
Additionalspecimenexamined.COSTARICA. Stamens 13-15, inserted in a complete circle;
LIMON:HoneCreek,8 kmS of Cahuita,30 m, 17Jul filamentsfar exceedingcalyx lobes, free to base,
1976(fl),J. &K. Utley5488(MO). glabrous.Ovary inserted at base of receptacle,
Licania kallunkiaebelongs to subgenusMo- tomentose on exterior.Style glabrous.Fruit not
quileasectionMoquilea.It is most closely related seen.
to L. minutiflorafromthe Guianasand Northern Distribution(Fig. 43). Known only from the
Amazonia,but it differsin the inflorescencewith typegatheringfromhighforest.Floweringin May.
the flowersborne in small groups on short ter- Licania guatemalensis belongs to subgenus
tiaryinflorescencebranches,in the smallerleaves MoquileasectionMoquilea,but is not easilycon-
with a more cuspidateacumen,and in the fewer fused with any other species in the section. The
stamens.Licania kallunkiaehas only 11-12 sta- leaves appearquite differentfrom other species
mens, fewerthanany otherspeciesof the section, in the section in their venation and flattened
but its exsertedstamens and presenceof petals, midrib with short, thick petiole. It is probably
closest to L. kallunkiaefrom Panama,but differs
as well as its similarityto L. minutiflora,all place
it in section Moquilea. It differs from another in the much longer leaves with abruptacumen,
relatedand recentlydescribedspecies from Pan- the shorter,thicker petioles, the smaller petals,
ama in the same section, L. chiriquiensis,in the and the glabrousstyle.
smaller flowerswith a gray-puberulousnot yel-
low-tomentellous indumentum, the spreading, 2-21.1. LicaniacecidiophoraPrance,Biotropica
much longer inflorescences, and the thinner, 10: 85, fig. 2A-E. 1978. Type. Peru. Amazo-
chartaceousleaves with a more cuspidate atten- nas: East of militarypost Chavez Valdivia, 2-
uate apex. 12 Feb 1974 (fl bud),Ancuash 752 (holotype,
NY; isotypes, F, MO).
2-18.3. LicaniaguatemalensisLundell,Wrightia Large tree to 30 m tall, the young branches
5: 39. 1974; Prance,Acta Amaz6nica 8: 583. puberulous,becoming glabrous with age. Leaf
1978. Type. Guatemala.Izabal:Between Seja lamina oblong, coriaceous, 9-12 x 3-5 cm,
and Fronterason Peten-GuatemalaRd., 6 May roundedat base and slightly decurrentinto the
30 Flora Neotropica

petiole, acuminateat apex, the acumen 5-8 mm tradaBelmonte,12Sep1975(fr),Cordeiro

755(INPA),
long, glabrous and shining above, with a ca- 12 Sep 1975(fl),Mota& Coelho86 (INPA).
ducous lanate pubescence beneath, becoming
glabrous with age; midrib prominent on both 2-23. Licania longipetala Prance, Fl. Neotrop.
surfaces;primary veins 14-19 pairs, prominu- Monogr.9: 62. 1972.
lous above, prominent beneath;petioles 16-22 Fruitnarrowlyellipsoid, 6-7 x 1-1.5 cm; exo-
mm long, canaliculate,slightlywinged on upper
carp smooth and glabrous;mesocarp thin and
portion, tomentellous when young, eglandular. fleshy; endocarp thin, hard and bony, sparsely
Stipulesinterpetiolar,small, triangular.Inflores- puberulouswithin.
cences of racemose panicles (only young ones Distribution.Figure 57.
seen), the rachis and branchesferrugineous-to-
mentose. Bracts and bracteoles triangular,pu- Additionalspecimensexamined.PERU.LORETO: Rio
berulouson exterior,with ciliate margins.Flow- Nanay,Quebrada de MapaCocha,23 Feb 1977(fr),
RimachiY.2838 (NY);Rio Amazonas,Quebrada de
ers buds only seen. Receptacle campanulate, Yanayacu, 3 Mar1977(fr),RimachiY.2866 (NY).
brown-tomentose on exterior, lanate-hirsute BRAZIL.AMAZONAS:
Tefe, Vila Nogueira, 17 Oct
within. Calyxlobes five, acute,tomentose on ex- 1975 (fl),D. Coelho& Damiaos.n. (INPA53295).
terior. Petals five, puberulouson exterior, with PARA:RioItacaiunas, 7 Oct1970(fl),
SerraBuritirana,
Pires & Belem 12676 (IAN, NY). RONDONIA:
Rio Ma-
ciliate margin. Stamens ca. 14, inserted around 29 (INPA).
chado,20 Oct 1978(fr),Goulding
a complete circle; filaments free (short and in-
cluded in buds). Ovaryinsertedat base of recep- The fruits of this species are eaten by fish
tacle, sparselylanate-pubescent.Style glabrous. (Goulding, 1980).
Fruit not seen.
Distribution.Figure 30. 2-23.1. LicaniatachirensisPrance,sp. nov. Type.
Venezuela.Tachira:Rio San Buena, 10 km W
Additionalspecimensexamined. PERU.AMAZONAS: of La
N of Cenepa,aboveChinkan EntseCreek,26 Feb1973 Fundacion,7?47'N,71?46'W,13-15 Mar
(st),Berlin902 (MO,NY), Mar1973(st),Berlin976 1980 (fl bud),R. Liesneret al. 9606 (holotype,
(MO,NY);N of CenepaalongChinkanEntseCreek, NY; isotypes, MO, VEN). Fig. 5.
25 Jul 1974(st),Berlin1799(MO,NY).
L. sectio Moquilea ab omnibus speciebus in-
Local name. Aguaruna,Jivaro:duship. florescentiisterminalibus,paniculatise cymulis
Uses. The small roundgallswhich form on the pluribus 2-4-floris breviter pedunculatis con-
leaf undersurfaceare used by the AguaruanaIn- structis differt. Petiolus 12-16 mm longus, ca-
dians as beads for their ceremonial capes (see naliculatus.
Berlin & Prance, 1978). Tree 8 m tall, the young branches glabrous.
This species belongs either to Licania subge- Leaf lamina oblong, coriaceous, 18-21 x 6.5-8
nus Moquilea section Moquilea or section Mi- cm, roundedat base, abruptlyacuminateat apex,
crodesmia.Openflowerswherethe stamenlength the acumen ca. 5 mm long, glabrous on both
is apparentare needed to determineto which of surfaces;midrib prominulousabove, prominent
these two sections it belongs. Licania cecidi- and glabrousbeneath;primaryveins 13-16 pairs,
ophorais most distinct, and easily recognizedby prominulousabove, prominentbeneath;petioles
the long petioles with slightly swollen base and 12-16 mm long, glabrous,rugulose,weaklycan-
by the lamina decurrentinto its upper portion. aliculate.Stipulesnot seen. Inflorescencesof ter-
minal and subterminalmuch branchedpanicles,
2-22. Licania unguiculata Prance, Fl. Neotrop. the flowers in small cymules on short tertiary
Monogr.9: 60. 1972. branches,the rachis and branches puberulous.
Distribution. Figure 83. Bracts and bracteoles triangular,membranous,
caducous. Receptacle campanulate, tomentel-
Additional specimens examined. BRAZIL. lous on exterior,tomentosewithin, sessile. Calyx
AMAZONAS: Manaus-Caracarai Rd.,km 57, 13 Sep 1976 lobes five, acute,tomentellouson exterior.Petals
(fl),Mota 594 (INPA),km 159,20 Sep 1974 (fl),Prance
et al. 22721 (INPA, NY), km 27, 19 Mar 1970 (fr), five, glabrous.Stamens 12-15, insertedin a com-
Rodrigues8770 (INPA);Rio Curicuriari,3 Nov 1971 plete circle (seen only in bud). Ovaryinsertedat
(fl), Prance et al. 16055 (INPA, NY). RONDONIA:
Es- base of receptacle, pilose. Fruit not seen.
SystematicTreatment 31

B.

youngbuds);D, petal.
32
Distribution.Figure 81.
Habitat. Primaryforested area on sandy soil,
700-1000 m.
This species is known also from specimensin
bud. It is quite distinctfrom any othercongener.
It belongs to section Moquilea,but differsfrom
all other species in the large inflorescencewith
the flowers borne in small cymules. The large
leaves with long canaliculatepetioles distinguish
it from most species.
Subgenus Moquilea Section Leptobalanus
2-27.1. Licania granvillei Prance, Proc. Kon.
Ned. Akad. Wetensch. Ser. C. 89: 114-116.
1986. Type. French Guiana. Saul, Monts La
Fumee, 3?37'N,53012'W,200-400 m, 21 Aug
1982 (fl),Mori & Boom 14764 (holotype,NY;
isotype, CAY).
Treeto 30 m tall, the young branchessparsely
puberulous,soon becomingglabrous.Leaf lam-
ina oblong,thicklycoriaceous,6-13 x 2.3-6 cm,
cuneate to subcuneateat base, finely acuminate
at apex, the acumen 10-25 mm long, glabrous
on both surfaces,withoutstomatalcavities;mid-
rib prominent on both surfaces, glabrous;pri-
mary veins 12-16 pairs, prominulous on both
surfaces,glabrous;petioles 5-8 mm long, can-
aliculate above, glabrous, eglandular,rugulose.
Stipules caducous (not seen). Inflorescencesof
racemosepanicles 6-12 cm long, the rachis and
branchessparselyappressedpubescent, appear-
ing dark because pubescence does not form a
complete covering. Bracts and bracteoles mi-
nute, 0.25 mm long, membranous.Flowers2-3
mm long, sessile along primarybranchesof in-
florescence. Receptacle campanulate, sparsely
puberulousto glabrouson exterior, densely to-
mentose within, sessile. Calyx lobes five, acute,
puberulouson both surfaces,with tomentellous
margins. Petals absent. Stamens 12, inserted
around complete circle, the filaments exserted
beyond calyx lobes, glabrous, but densely to-
mentose around base, free to base. Ovary in-
sertedat base of receptacle,sparselylanate.Style
exserted, sparselytomentose for half of length.
Fruit globose to ellipsoid, 5.5 cm long x 3.5-4
cm broad, glabrousand lenticellateon exterior;
mesocarpfleshy, 3-4 mm thick;endocarphard,
bony, 1 mm thick, glabrouswithin.
Flora Neotropica
Distribution(Fig. 42). Upland forest on terra
firmein hills from Colombia to FrenchGuiana.
Specimensexamined.COLOMBIA.AMAZONAS:Rio
Apaporis,RaudalYayacopi,0?5'S,70?30'W,800', 18
Feb 1982 (fr), Schultes& Cabrera15456 (NY); Jino-
goje, 700', 27 Feb 1982 (fr),Schultes& Cabrera15660
(NY).
VENEZUELA.AMAZONAS:Trail south from Cerro
Neblinabasecampon Rio Mawarinuma fromclear
waterstreamto topof firstsmallhills,0?50'N,66?1 'W,
3 May 1984 (fl, fr), Gentry& Stein 47121 (NY).
FRENCHGUIANA.Saul,MontsLaFum6e,200-
400 m, 26 Aug 1982(fl),Mori&Boom14772(CAY,
NY), 28 Aug 1982(fl),Mori& Boom14790(CAY,
NY), 1 Sep 1982 (fl),Mori &Boom 14828 (CAY,NY),
28 Mar1983(fr),Mori&Pipoly15428(CAY,NY), 1
Apr 1983 (fr), Mori & Pipoly 15476 (CAY, NY), 16
Apr 1983 (fr),Mori & Pipoly 15523 (CAY, NY).
BRAZIL.AMAZONAS:Serrada Neblina,betweenPal-
mitoandTatuCamp,400-600 m, 21 Dec 1965(fr),
Silva & Brazao 60697 (NY).
This species is close to Licania apetala, es-
peciallyto its var. aperta,but differsin the larger
flowers, the larger,thicker leaves with a much
more attenuateapex, and the minute bracteoles.
The Brazilianand Colombianmaterialin fruit
was kept aside as a possible new species at the
time of my monograph(Prance, 1972), and it is
now possibleto describeit with the new material
from French Guiana. This species is known in
both localitiesfrom the hill slopes of mountains.
It is named for J-J. de Granvillewho has done
much to interpretthe flora of French Guiana.
2-31.1. Licania jefensis Prance, Brittonia 28:
215-216, fig. 5.1976. Type. Panama.Panama:
Summit of CerroJefe, 2 Apr 1969 (fl), Dwyer
et al. 5047 (holotype, NY; isotype, MO).
Shrubto 2 m tall, the young branchestomen-
tose, soon becoming glabrous.Leaf lamina el-
liptic, coriaceous,3-6 x 1.8-3.5 cm, roundedat
base, acute to acuminateat apex, the acumen 3-
5 mm long, glabrouson both surfaceswhen ma-
ture, when young with a lanate, caducousindu-
mentum which is more persistentbeneath,with
scatteredglands on lower surface,without sto-
matal cavities, midrib prominent on both sur-
faces; primary veins 8-11 pairs, prominulous
above, prominentbeneath; secondaryvenation
conspicuouslyprominulousandreticulateon both
surfaces; petioles 2.5-4 mm long, tomentose,
eglandular,rugulose,slightly canaliculate.Stip-
SystematicTreatment
ules small, axillary, caducous. Inflorescencesof
terminaland subterminalracemosepanicles, 6-
11 cm long, the rachis and branches yellow-
brown-tomentose.Bractsand bracteolesca. 0.5
mm long, tomentose on exterior, persistent.
Flowersca. 2 mm long. Receptaclecampanulate,
sessile, tomentoseon exterior,pilose within. Ca-
lyx lobes five, tomentoseon exterior,puberulous
within. Petals absent. Stamens ca. 13, inserted
around complete circle; filaments far exserted,
free to base. Ovarypilose. Style glabrous,equal-
ling filamentsin length.Fruitglobose, ca. 25 cm
in diam. epicarpsmooth and glabrous;mesocarp
thin and fleshy;endocarphard, bony, thin.
Distribution(Fig. 50). Known only from the
cloud forestsof Panama,floweringin Marchand
April.
Additionalspecimensexamined.PANAMA.PANAMA:
9.4 km N of GoofyLake,900 m, 11 Mar 1977(fl),
Folsomet al. 1996(MO,NY);CerroJefe,1000m, 11
Jun 1975 (y fr), Mori 6532 (NY), 29 Aug 1975 (fr),
Mori7990(MO,NY).
2-31.2. Licaniamorii Prance,Brittonia28: 215,
fig.4.1976. Type.Panama.Panama:El Llano-
CartiRd., 12 km from Inter AmericanHwy.,
15 Feb 1975 (fl), Mori & Kallunki 4665 (ho-
lotype, NY; isotype, MO).
Treeto 15 m tall, the young branchestomen-
tellous, soon becoming glabrous. Leaf lamina
elliptic, coriaceous,7-11 x 2.7-5.5 cm, subcor-
date at base, acuminateat apex, the acumen 8-
12 mm long, glabrousand shiny above, with a
shortglaucousappressedpubescencebeneathand
with conspicuous palisade glands, without sto-
matal cavities; midrib prominulous above,
prominent beneath; primary veins 7-9 pairs,
prominulousabove, prominentbeneath;petioles
3-6 mm long, sparselytomentellous,weaklycan-
aliculate. Stipules caducous, not seen. Inflores-
cencesof terminalor subterminalracemosepan-
icles, 7-12 cm long, the rachis and branches
sericeous-tomentellous.Bracts and bracteoles
triangular,eglandular,ca. 1 mm long, caducous.
Flowersca. 2 mm long. Receptaclecampanulate,
sessile or with short pedicel to 1.5 mm long,
tomentoseon exterior,pilose within.Calyxlobes
five, acute,tomentellouson both surfaces.Petals
absent. Stamens ca. 13, inserted around com-
pletecircle,the filamentsfar-exserted,freeto base.
33
Ovarypilose. Styleglabrousexceptaroundbase,
equallingfilamentsin length. Fruit unknown.
Distribution(Fig. 65). Known only from the
type gatheringfrom moist forest at 350 m, col-
lected in flowerin February.
2-32. Licania sparsipilis Blake, Contr. Gray
Herb. 52: 67. 1917.
Distribution.Figure79.
Additionalspecimensexamined.MEXICO.OAXACA:
30kmS ofrd.PalomarestoUxpanapa,
betweenarroyo
HumacaandRio Verde,300 m, 20 Apr1985(fl), W.
Thomaset al. 3575(NY).
ELSALVADOR.AHUACHAPAN:
Rio ElVenado,Fin-
ca SanBenito,6 Mar1979(y fr),Castros.n. (NY).
PANAMA. COLON:Santa Rita Ridge, 2 Mar 1975
(fl),Mori&Kallunki4914(MO,NY).
2-32.1. LicaniacuatrecasasiiPrance,Acta Ama-
zonica 8: 577. 1978. Type. Colombia. Valle:
Alto Yunda,Rio Anchicaya,1000 m, Oct 1972
(fl), Hilty 0-1 (holotype, US; isotype, NY).
Treeto 30 m tall, the young branchespuber-
ulous, soon glabrate.Leaf lamina elliptic, cori-
aceous, 8-12.5 x 3.5-8 cm, subcuneateat base,
cuspidateat apex, the acumen 10-15 mm long,
slightlycurved,glabrousabove, with a compact
brown-lanatepubescencebeneath, without sto-
matalcavities;primaryveins 10-12 pairs,prom-
inent beneath, prominulous above; midrib
prominent on both surfaces;petioles 8-11 mm
long, tomentellouswhen young,tereteor slightly
canaliculate,eglandular,transverselyrugulose.
Stipules caducous (not seen). Inflorescencesof
racemose panicles usually once-branched,oc-
casionallywith secondarybranchesup to 10 cm
long, the rachis and branchesbrown-tomentel-
lous. Bractsand bracteolesovate, ca. 1 mm long,
persistent, tomentellous on exterior, entire,
eglandular.Flowersca. 2.5 mm long, sessile on
primary and secondary branches of inflores-
cence. Receptaclebroadlycampanulate,tomen-
tose on exterior,pilose within. Calyx lobes five,
acute, tomentellous on exterior, puberulous
within. Petals absent. Stamens 10-12, inserted
in a completecircle;filamentsfarexceedingcalyx
lobes, free to base, glabrous.Ovary inserted at
base of receptacle,villous aroundbase, but gla-
brous above. Style glabrous,equallingfilaments
in length. Fruit not seen (25-30 mm long ac-
cordingto field notes).
34 FloraNeotropica
Distribution(Fig. 34). Known only from up- Licania mexicana was described from poor
land forest of El Valle, Colombia. materialwith only old flowers present, making
it difficultto relateto other species. It belongsto
Additional specimenexamined.COLOMBIA. VALLE:
BajoCalima,15kmN of Buenaventura, 18Feb 1983 subgenusMoquilea, either section Moquilea or
(fl),Gentry&Juncosa40467(MO,NY). sectionLeptobalanus,dependingon the presence
or absence of petals, which cannot be observed
This speciesfromthe highlandsof Vallecomes in the old flowers.It
from an area in need of furtherexploration.It is probablybelongsto section
Leptobalanus and seems to be most closely re-
most closely related to Licania apetala and L. lated to the CentralAmerican
species L. spar-
sparsipilis,but differs from both in the dense sipilis.It differsin the shortinflorescencebranch-
brown-lanatepubescence of the leaf undersur-
es, the less acuminateleaveswith shorterpetioles,
face, which is set in the deeply reticulate sec- and the
greaternumber of stamens.
ondary and tertiaryvenation, as well as in the
leaf's long cuspidateacumen, the largerpetioles
withoutthe twoglandsofL. sparsipilis,thebrown- 2-33.1. Licania joseramosiiPrance, Acta Ama-
tomentellous pubescence of the inflorescences, z6nica 8: 581, fig. 3. 1978. Type. Brazil.Ama-
and the very small, broadlycampanulaterecep- zonas: Manaus-CaracaraiRd., km 130, 6 Jan
tacle. 1976 (fl), Monteiro & Ramos 29 (holotype,
INPA 54340; isotype, NY).
2-32.2. Licania mexicana Lundell, Wrightia 5:
40. 1974; Prance, Acta Amazonica 8: 585. Small tree 5 m tall, the young branchesgla-
1978. Type. Mexico. Sinaloa:BetweenRancho brous. Leaf lamina oblong to oblong-lanceolate,
Del Pinioand Chele, 22 May 1943 (fl),Lundell coriaceous, 13-20 x 4-6.5 cm, cuneate at base,
13023 (holotype, LL; isotype, MICH). finely acuminate at apex, the acumen 15-20 mm
long,glabrouson both surfaces,withoutstomatal
Tree 10 m tall, the young brancheslanate to cavities; primaryveins 9-14 pairs, prominulous
puberulousand soon glabrous.Leaf lamina nar- on both surfaces;midribprominenton both sur-
rowlyoblongto lanceolate,coriaceous,6.5-13 x faces;petioles 4-5 mm long, rugulosetomentel-
2.5-4 cm, cuneateat base, graduallyattenuateto lous when young, terete, with two glands near
acuteapex,glabrousabove, with a caducouslan- junction with lamina. Stipules linear, ca. 6 mm
ate pubescence beneath when young, stomatal long, hispidulous, caducous. Inflorescencesof
cavities absent; midrib prominulous above, panicleswith long thick centralrachisand short,
prominent beneath; primary veins 6-8 pairs, thin, lateral branches bearing 1-3 flowers, the
prominulouson both surfaces,secondaryvena- rachis and branches tomentellous. Bracts and
tion prominulous and conspicuously reticulate bracteoles lanceolate, subpersistent, tomentel-
on both surfaces;petioles 1.5-3.5 mm long, ru- lous on exterior, glabrous within, entire, with
gose, terete, lanate when very young, soon be- long, thin acumen,eglandular.Flowersca. 5 mm
coming glabrous. Stipules axillary, triangular, long. Receptacle campanulate,gray-tomentose
persistent,2 mm long, lanate when young. In- on exterior,tomentose within. Calyx lobes five,
florescenceof terminal panicles 2-3.5 cm long, acute, tomentose on exterior.Petals absent. Sta-
with short primarybranches (only 2 old inflo- mens ca. 19, inserted around complete circle;
rescences seen) gray-brown, the rachis and filamentsslightlyexceedingcalyx lobes, free al-
branches tomentellous. Bracts and bracteoles most to base, glabrousexcept for pilose annular
small, ca. 1 mm long, persistent,tomentellous, ring.Ovaryinsertedat base of receptacle,lanate.
ovate-triangular.Receptacle campanulate, to- Styleglabrous,equallingfilamentsin length.Fruit
mentellouson exterior,pilosewithin.Calyxlobes not seen.
triangular,1 mm long, acute, reflexed, tomen- Licania joseramosii (distributionin Fig. 51),
tellous. Petals? (old flowers only present). Sta- is a most distinct species that cannot be easily
mens 14-15, inserted around complete circle, confusedwith any otherin the genus.It is related
exsertedbeyond calyx lobes; filamentsglabrous, to L. emarginataand L. calvescensbut differsin
insertedat baseof receptacle.Styleglabrous.Fruit a large number of characterssuch as the much
not seen. longerleaves, the largerflowers,and the distinc-
Distribution(Fig. 60). This species is known tive inflorescence.It differsfrom L. emarginata
only from the type. in the greaternumber of stamens, and from L.
SystematicTreatment 35

calvescensin the flowers borne in small groups 2. Leaves broadly ovate to oblong, obtuse to
on secondaryinflorescencebranchesand in the bluntlyacuminate,the acumen 1-5 mm long;
uppersurfaceof leaf dryingbrown;youngin-
glabrousleaves. SuperficiallyL. joseramosii also florescencewith sparse gray-browntomen-
resembles L. longipedicellatain subgenus Mo- tum. a. subsp. octandra.
quileasectionMoquileabut differsin the smaller, 2. Leaves oblong-lanceolatewith a well-devel-
thickerleaves, the much less branchedinflores- oped finely pointed acumen 5-13 mm long;
cence, the smaller flowers, and the absence of upper surface of leaf drying gray or green;
young inflorescenceusually with a rufous-
petals. brownarachnoidindumentum.
b. subsp.pallida.
2-34. LicaniacalvescensCuatrecasas,Fieldiana, 1. Leaves 14-29 x 4.5-7 cm, long-acuminateat
Bot. 27: 64. 1950. apex, the acumen 12-28 mm long.
c. subsp.grandifolia.
This distinct specieswas known only from the
type from El Valle in Prance(1972). Since then 2-43c. L. octandra subsp. grandifolia Prance,
I have seen two furthersterile inventory collec- Acta Amazonica 4(1): 18. 1974. Type. Brazil.
tions that may be referredto this Pacificcoastal Amazonas: Rio Javari, behind Estirao de
forest species (Fig. 30). Equador, 9 Aug 1973 (fl), Lleras et al. P17270
(holotype, INPA; isotype, NY).
Additionalspecimensexamined.COLOMBIA.
CHoc6:TrailTubad6to Quibd6-Tutunendo Rd., 17 Specimens examined. COLOMBIA. AMAZONAS:Rio
Jan1979(st),Gentry&RenteriaA. 24331(MO,NY). Loreto-Yacu(st), Glenboski206 (NY). PERU.LORETO:
VALLE:BajoCalima,N of Buenaventura, lowerRio Rio Tacha, Curaray,18 Sep 1972 (fl), Croat 20372
SanJuan,8 Dec 1981(st),Gentry35478(MO,NY). (AAU, MO, NY); Maynas,Mishana,Rio Nanay half-
way between Iquitos and Santa Maria de Nanay, 31
2-38. Licania albiflora Fanshawe & Maguire, May 1978 (st),Gentryet al. 22387 (NY);Maynas,Dtto.
FernandoLores,QuebradaTamshiyacu,10 Jan 1977
Bull. TorreyBot. Club 75: 318. 1948. McDaniel& Rimachi21133
(fr), (NY); Maynas,Iqui-
This species was known from two collections tos, Carreterade PefiaNegra,10 Feb 1977(fr),Rimachi
Y 2797 (NY); Prov. Loreto:Nauta, Rio Marafion,9
from Guyanaand Surinamin Prance(1972). A Nov 1982 (fl buds), Vdsquez& Jaramillo3444 (NY);
thirdcollectionfromFrenchGuianahas recently Maynas,Pto. Almendras(Rio Nanay), 19 Feb 1985 (fl
been added,showingit to be rarebut widespread buds), Vasquez& Jaramillo6243 (NY).
in the Guianas. See Figure23.
Section Microdesmia
Additional
specimen FRENCH
examined. GUIANA. Subgenus Moquilea
La
Saiil, Fumee 27
Trail, Mar1983(fr), &
Mori Pipoly 2-45. Licania arborea Seemann, Bot. voy. Her-
15411(NY).
ald 3: 118,t. 25. 1853.
2-39. Licanialongistyla(Hookerf.) Fritsch,Ann. This species, common from Mexico through
K. K. Naturhist.Hofmus. 4: 56. 1889. Central America, was known by only three col-
lections from South America at the time of Prance
This species, common in Venezuela, the
(1972). It is apparently more widespread in South
Guianas,and westernand centralAmazonia,has
America, although it may well have been intro-
recentlybeen collected in Panama;yet another duced there indigenous traders because of its
Panama-Amazondisjunct.See Figure 58. by
use as an oilseed.
Additional
specimen examined.PANAMA. SANBLAS: Distribution. Figure 26.
El Llano-CartiRd., 9?19'N,78?55'W,300 m, 8 Jan
1985(fl),de Neverset al. 4446(MO,NY). Additional specimens examined. COLOMBIA.
CHOCO:Mun. Riosucio, ParqueNacional Los Katios,
2-43. Licania octandra(Hoffmannseggex Roe- 1 Dec 1976 (fr),Le6n 632 (MO).
VENEZUELA.ZULIA:Aricuiza, 19 Dec 1972 (fl),
mer & Schultes)Kuntze, Revis. gen. pl. 217. Veillon131 (US, VEN) [possiblycultivated].
1891. Fig. 68. PERU. HUANUCO: Prov. Pachitea,Rio Pachiteanr.
Mielde Abeja,9 Mar 1967(fr),SchunkeV 1737 (COL,
A new subspecies was described in Prance US); Dist. Honoria, Caserio Leoncio Prado, 29 Nov
(1974a). The three subspecies may be distin- 1963 (fl), Lao Magin 103 (F). LORETO:Maynas, trail
guishedby the following key. from Indianaon Rio Amazonasto Rio Napo, 24 May
22199
1978 (fl), Gentryet al. (MO, NY).
1. Leaves 3-12 x 2-4 cm, the apex obtuse to acu- BRAZIL.ACRE:Rio Macaua,Mun. Sena Madureira
minate, the acumen 1-13 mm long. (fl), Lima & Souza 231 (INPA, NY).
36 Flora Neotropica

2-47. Licania subarachnophylla Cuatrecasas, subarachnophyllain the subgenusMoquileasec-

Fieldiana,Bot. 27: 110. 1951. tion Microdesmia.These four are a closely re-
lated group of species each well separatedfrom
Distribution (Fig. 80). This little-collected one another geographicallyor ecologically. Li-
species,known only from the type and one other cania salicifolia is a montane species of Antio-
collection, from Boyaca and Meta in Colombia, quia, Colombia,which differsin its much longer
has now been found in Venezuela. It seems to inflorescenceswith flowersin clustersand in the
be endemic to the galleryforestsalong the rivers longerlanceolateleaves; L. subarachnophyllais
of the savannas. a speciesof the galleryforestsof the Llanoswhich
Additionalspecimenexamined.VENEZUELA. also differsin the much longerinflorescencewith
APURE:Distr.PedroCamejo, alongQuebrada ElPorve- the flowersin clustersand in its smallerflowers;
nir, 6?39'N,67017'W,21 Feb 1979 (fl),Davidse& Gon- L. araneosa is a species of the galleryforests of
zalez 15541 (MO, NY). a restrictedareaof the planaltoof CentralBrazil,
which differs in its smaller flowers, larger
2-47.1. Licania tambopatensisPrance, sp. nov. branched inflorescences,and more coriaceous
Type. Peru. Madre de Dios: TambopataNa- leaves.
tureReserve,LagunaCoco Cocha, 5.2 km east This species is named for the TambopataNa-
of lodge, 3 Jun 1986 (fl), V.A. Funk, B. Kahn ture Reserve, a remarkablesanctuary for the
& S. Wiser 8415 (holotype, NY; isotype, species diversity of the upper Amazon region
US). Fig. 6. where the highest known level of species diver-
Ab L. araneosa floribus 2 mm longis, inflo- sity has been recordedfor many organisms.
rescentiisracemosis, foliis chartaceisdiffert.
Shrub,the young branchestomentose becom- 2-48. Licania salicifolia Cuatrecasas,Fieldiana,
Bot. 27: 111. 1951.
ing glabrouswith age. Leaf lamina oblong, char-
taceous, 3.8-6 x 1.5-2.4 cm, cuneate at base, This species was known from a single rather
mucronateat apex, the mucro 1-2 mm long, gla- poor type specimen from Antioquia, Colombia.
brousabove except on midrib,appressedlanate- It is a high altitude endemic still only known
pubescentbeneath; midrib prominent beneath, from the type localityin the vicinity of Rionegro
prominulousand tomentose above when young; (not the Rio Negro as erroneously stated in
primary veins 11-14 pairs, prominulous be- Prance, 1972!). Espinal T. (1981) wrote a com-
neath, plane above; petioles 2-4 mm long, to- mentaryon the rarityof this species and pointed
mentose, terete, eglandular.Stipules linear, ax- out that residentsof Comfamaknow this as "the
illary, caducous, membranous. Inflorescences raretree." See Figure 76.
of terminal and axillary racemes, 2-4 cm, the
rachis tomentose. Bracts and bracteoles mem- Additionalspecimensexamined.COLOMBIA.
ANTIOQUIA:Rionegro, HaciendaComfama, Jul 1981
branous, caducous, triangular, tomentellous on (st), Espinal T. et al. 4537 (MEDEL),3 Dec 1981 (fl),
exterior,glabrouswithin.Flowers2-2.5 mm long, EspinalT.&A.LennaT.4608(MEDEL), 3 July1981
sessile, denselypackedalong rachisbut not clus- (fl),R. Jaramillos.n. (COL,NY).
tered. Receptaclecupuliform,sessile, brown-to-
mentose and lanate on exterior,densely tomen- Subgenus Licania Section Hirsuta
tose within. Calyx lobes acute, triangular,lanate
on both surfaces. Petals small, caducous, the 2-53. Licania hirsuta Prance, Fl. Neotrop.
marginsciliate. Stamensca. 15, insertedaround Monogr.9: 93. 1972.
complete circle, equalling or only slightly ex- This species was described from two collec-
ceeding the calyx lobes, glabrous, free to base. tions in CentralAmazonas, Brazil,and has now
Ovary inserted at base of receptacle.Style gla- also been found in Amapa. See Figure46.
brous except at base. Fruit not seen.
Distribution(Fig. 62). Known only from the Additionalspecimensexamined.BRAZIL.AMAPA:
collected Road Tinguilimto BraSo,km 21, 2 Jun 1969 (fl), N.
type along the edge of the lake. T. Silva 2096 (IAN, NY). AMAZONAS: Mun. Humaita,
This new species belongs close to the species rd. Humaita-Porto Velho, km 70, 5 May 1982 (fl),
groupof Licania araneosa,L. salicifolia,and L. Teixeiraet al. 253 (INPA,NY).
 SystematicTreatment 37

A.

iN.N K

ii~i""~~~~mm

FIG. 6. Licania tambopatensis(Funk et al. 8415). A, habit;B, leaf undersurface;C, inflorescence;D, bract
and bracteole;E, flower;F, flowersection;G, ovary.

2-57.1. Licania hispida Prance, sp. nov. Type. rachidibusinflorescentiarumbasim versus his-
Venezuela.Amazonas: Dept. Rio Negro, Ce- pidis unica est.
rro Aratitiyope, 2?10'N, 65?34'W, SSW of Tree6 m tall, the young branchesdensely his-
Ocamo, 990-1670 m, 24-28 Feb 1984 (fl, fr), pid. Leaf lamina oblong to oblong-elliptic, co-
J. A. Steyermark et al. 130185 (holotype, riaceous, 7-11 x 2.5-5.2 cm, rounded and un-
NY). Fig. 7. equal at base, apiculateor bluntly acuminate at
apex, sparsely hispid on primary venation be-
Speciessectione Hirsutaepertinens,in qua ra- neath, otherwiseglabrouson both surfaces;mid-
mulis juvenilibus, costis petiolis que hispidis, rib prominulousabove, prominentbeneath,his-
38 Flora Neotropica

,.'~~ 1cm.Y~gBB~~ 1
12cm.

cmEE;cm.

G~~. D. "" B.,,~~0

FIG. 7. Licania hispida (Steyermark 130185). A, habit; B, stem and petiole; C, leaf undersurface; D, flower;
E, flower section; F, young infructescence; G, young fruit.

pid beneathespeciallyon lower portion;primary tion of rachis hispid, sparsely tomentellous on

veins 9-12 pairs, plane above, prominent be-
neath,the lower ones sparselyhispid;petioles 4-
7 mm long, terete, eglandular,hispid. Stipules
linear, 5-6 mm long, subpersistent. Inflores-
cencesof little-branchedpanicles,the lower por-
upperportionand on branches.Bractsand brac-
teoles 1 mm long, lanceolate,persistent,tomen-
tellous. Flowersin bud ca. 1.5 mm long, sessile
on primarybranchesof inflorescence.Receptacle
campanulate,sessile, gray-tomentoseon exteri-
SystematicTreatment 39

or, tomentose within. Calyx lobes five, acute, posite to three calyx lobes, shorter than calyx
tomentose on both surfaces.Petals five, minute. lobes,thefilamentsconnateforlowerthird.Ovary
Stamens five, inserted around complete circle; pilose. Style pilose. Fruit unknown.
filamentsshorterthancalyxlobes, free,glabrous. Distribution (Fig. 70). Known only from the
Ovaryinsertedat base of receptacle,lanate.Style type collection from 1400 m, collected in flower
glabrousexcept at extreme base. Fruit (imma- in May.
ture) globose, exocarp ferrugineous-pubescent;
endocarpthin, hard. 2-61. Licania arachnoidea Fanshawe & Ma-
Distribution(Fig. 46). Known only from the guire, Bull. TorreyBot. Club 75: 318. 1948.
type from forest at base of high bluffs of an ig- Distribution(Fig. 25). A recent collection of
neous rock mountain.
this specieswas made in LoretoDept., Peru.Pre-
This new species belongs to section Hirsuta
it was knownonly from the Guianasand
and is closely relatedto the other species in that viously
from a single collection from western Brazil. It
section and in section Hymenopus. It is quite
adds to the growing list of species disjunct in
distinct nonetheless,and easily distinguishedby
easternandwesternAmazoniathatarenot found
the hispidpubescenceof the stems,petioles,mid-
in CentralAmazonia,suchas Licaniaguianensis
rib and lower part of the inflorescencerachis.
and Couepia parillo.
This type of hispid pubescence,common in the
genus Hirtella, does not occur in other species Additional specimen examined. PERU. LORETO:
of Licania. Licania hispida, with acute or only Maynas, Hondocochanr. Santa Cecilia, Rio Maniti,
11 Nov 1976 (fl), Encarnaci6n970 (NY, US).
bluntly acuminate leaves is also distinguished
from relatedspecies by the leaf shape.
2-64. Licania caudata Prance, Fl. Neotrop.
Monogr.9: 103. 1972.
Subgenus Licania Section Hymenopus Fruitoblong,25 x 8 mm; exocarpsmooth and
2-60.1. LicaniapakaraimensisPrance,Brittonia glabrous,often slightlyrugulose,wrinkledwhen
28:218, fig.6. 1976. Type. Venezuela.Bolivar: dry; mesocarpthin, 0.25 mm thick, fleshy;en-
SierraPakaraima,headwatersof Rio Paragua, docarp thin 0.5 mm thick, bony, slightly pu-
3?40'N, 63?0'W, May 1973 (fl), Steyermark berulouswithin.
107357 (holotype, NY; isotype, VEN). Additional specimens examined. COLOMBIA.
Treeto 10 m tall, the young branchessparsely VAUPES: Mitfi,25 Apr 1975 (fr),Zarucchi1278 (COL).
FRENCHGUIANA. Saul,MontsLa Fumee, 12 Oct
tomentellous,soon glabrateand lenticellate.Leaf 1982 (st), Boom & Mori 1997 (NY).
lamina narrowlyoblong, coriaceous,7.5-11.5 x PERU. LORETO: Rio Nanay above Bellavista, 1 Jun
2.5-4.5 cm, cuneateat base, acuminateat apex, 1976 (fl), Rimachi Y. 2326 (NY); Dept. Sapuena,Ar-
the acumen 3-12 mm long, glabrous on both boretumJenaro Herrera,4 Jun 1974 (fl), Diaz 51-A
surfaces;primaryveins 11-17 pairs,planeabove, (IAN); Pucallpa, 1 Aug 1980 (fr), Salazar 662 (MO,
NY), 6 Jun 1960(fl),Woytkowski5773(S,US). UCAYALI:
prominulousand glabrousbeneath;petioles 3-5 Km 86 Pucallpa-TingoMariaRd., Jul 1978(fr),Froeh-
mm long, terete,rugulose,glabrousor with a few ner 362 (MO, NY).
appressedhairs,eglandular.Stipulesaxillary,to- BRAZIL.ACRE: Cruzeirodo Sul, 12 Feb 1976 (fr),
Marinho 134 (IAN). AMAZONAS:Rio Cuieiras, nr. Lago
mentellous,caducous.Inflorescencesof terminal de Peixe-Boi, Sep 1972 (fr), O. Pires & Honda 185
panicles with flowersgroupedin small cymules (INPA, NY); Manaus-PortoVelho Rd., between Rio
on peduncles1 mm long, the rachisand branches Castanhoand Araca, 10 Oct 1974 (st), Prance et al.
tomentellous,the rachisca. 2 mm thick at base. 22775 (INPA, NY), 12 Jul 1972 (fl), M. F. da Silva
Bractsandbracteolesca. 0.5 mm long,triangular, 486 (INPA, NY). PARA: Rd. Oriximinato Obidos, km
70 at Rio Cumina-Mirim,14 Sep 1980 (fr), Cid et al.
persistent.Flowersca. 2.5 mm long. Receptacle 2545 (INPA, NY).
campanulate,gray-tomentellouson exterior,to-
mentellouswithin;pedicels0.25 mm long. Calyx This species was described from three collec-
lobes five, acute, gray-tomentellouson exterior, tions from the vicinity of Manaus. It has turned
glabrouswithin. Petalsfive, caducous,remaining out to be a widespread and common Amazonian
attachedto one anotherand caducousin a single species, occurring in both inundated and terra
calyptra-likeunit. Stamens five, unilateral,op- firme forest. See Figure 31.
40 FloraNeotropica
2-64.1. Licania miltonii Prance, Acta Amazo- noco Deltaof Venezuelabut also occursin French
nica 13: 24. 1983. Type. Brazil.Mato Grosso: Guiana. See Figure 54.
Aripuana,km 238 ofrd. BR 174,NuicleoJuina, Additional specimenexamined.FRENCH GUIANA.
area urbana, 17 Jan 1979 (fl), M. G. Silva & Pistede St. Elie,19 Oct 1977(st),Lescure758 (NY).
A. Pinheiro4296 (holotype,MG;isotype,NY).
Tree5 m tall, the young branchessparselypu- 2-67. Licania glabriflora Prance, Fl. Neotrop.
berulous,soon becoming glabrous.Leaf lamina Monogr.9: 104. 1972.
narrowlyoblong, chartaceous,4-10.2 x 1.3-3.8 Distribution(Fig. 41). This species, described
cm, cuneateat base,cuspidate-acuminate at apex, from Venezuela and French Guiana in Prance
the acumen 6-12 mm long, glabrous on both (1972), has now been collectedin adjacentBrazil
surfaces; midrib slightly prominulous above, and Surinam,but also in Costa Rica, giving it a
prominent beneath, with a few stiff appressed disjunctdistributionsimilarto that of L. affinis.
hairs on both surfaces;primaryveins 6-8 pairs, The CentralAmericanmaterialdiffersonly in
plane above, slightly prominulousbeneath;pet- having three ratherthan five stamens and often
ioles 1-3 mm long, terete, eglandular,with few four ratherthan five petals. In all other respects
sparseappressedhairs.Stipulesintrapetiolar,lin- it resemblesthe SouthAmericanspecimensstud-
ear, persistent,ca. 2 mm long. Inflorescencester- ied.
minal and axillary little-branchedpanicles 1-3
cm, the rachis and branches very sparsely hir- Additionalspecimensexamined.COSTARICA.
HEREDIA:FincaLa Selva,Rio PuertoViejo, 14 Feb
sutulous. Bracts and bracteoles minute, mem- 1982 (fl), Hammel 11149 (NY).
branous, sparsely hirsutulous on exterior, per- SURINAM.LelyMountains,SW plateau,29 Sep
sistent, entire. Flowers minute, 1-1.5 mm long, 1975 (fl), Lindeman& Stoffers525 (S).
borne solitary on short primarybranchesof in- FRENCHGUIANA.Riv. GrandInini,SaintBa-
florescence.Receptaclecampanulate,glabrousor tardeau,11 Sep 1970 (fl), GranvilleB3787 (CAY).
BRAZIL.AMAPA: Between Porto Platon and Serra
sparsely hirsutulous on exterior, tomentose on do Navio, Oct-Dec 1976 (st), Rosa 1165 (MG).
exterior;pedicels 0.2 mm long. Calyxlobes five,
acute, glabrous except for a few stiff appressed Note that in Prance (1972: 105) this species
hairson exterior,the marginsciliate. Petals five, was erroneously cited as Licania glabrifolia.
tomentellouswithin towards apex, glabrousbe-
2-69a. Licania heteromorphaBentham, J. Bot.
neath,puberulouson outersurface.Stamensfive,
fertile,insertedon shortthick filaments,connate (Hooker)2: 221. 1840. var. heteromorpha.
at base. Ovary inserted at base of receptacle, This is the commonest and most collected
sparselyhirsute.Fruitellipsoid, ca. 2 cm long x Amazonianspecies of Licania. In Prance(1972)
1 cm broad;exocarpsmooth, glabrous;mesocarp I also cited material from Rio de Janeiro for
thin; endocarp0.5 mm thick, glabrouswithin. which I was uncertainif it was of cultivated or-
Distribution(Fig. 64). Forest on terra firme, igin or for the Glaziou specimenspirated.A new
clay soil in Mato Grosso. collection shows that this species is a native of
Additionalspecimenexamined.BRAZIL.MATO the easternBrazilianforestsand is anotherAm-
GROSSO: Aripuana,NuicleoJuina, 28 May 1978 (fr), azon-easternBrazil disjunct(Fig. 44).
M. G. Silva & Rosario 4684 (MG).
Additionalspecimenexamined.BRAZIL.ESPIRITO
This species, most closely related to Licania SANTO: Mun. Linhares,Res. Florestalda Cia Vale do
caudata, differsin the much smaller, narrower, Rio Doce, 14 Dec 1981 (fl),H. C. de Lima 1660 (NY,
chartaceousleaves with cuspidate not caudate RB).
apices and shorter petioles, the smaller flowers 2-69e. Licania heteromorphaBentham var. re-
with fewer stamens with free filaments, and volutaPrance,Acta Amazonica 13: 24. 1983.
smaller inflorescences.
Type. Brazil. Amazonas: 20 km NW of Ma-
naus,Tarumadevelopmentarea,21 Mar 1981
2-65. Licania latistipula Prance, Fl. Neotrop. (fl), B. W. & S. P. Nelson 1058 (holotype, INPA;
Monogr.9: 103. 1972. isotype, NY).
This species, with distinctively large stipules, Leaves2-4.5 x 1.1-2.3 cm, the marginsrevo-
was described from two collections in the Ori- lute; anthers deltoid.
Systematic Treatment 41

, _A. . . _.
~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~l-i
=:z- ..

0-1~~~~~~~
E. C~~~~~~~~i
.~

FIG. 8. Licanialaevigata(Irwinet al. 54990;M. Silva80). A, habit;B, flower;C, flowersection;D, petal;

E, ovary;F, floraldiagram;G, young fruit.

Habitat. Forest on terra firme, clay soil.
This variety is quite distinct from var. heter-
omorphaand the other varieties of the species
by its much smaller, thickly coriaceous leaves
with revolute margins and a rounded to retuse
apex. It is possibly a distinct species, but I hes-
itate to describeit as such on the basis of a single
collection and in this complex, because there is
a great deal of morphologicalvariation in var.
heteromorpha.It also differsfrom othervarieties
ofL. heteromorphain the deltoid anthers,which
also occurin the closely relatedL. intrapetiolaris.
The anthersare distinctive because in the dried
materialthe pointedapex is much darkerin color
than the broad basal portion.
2-69.1. LicanialaevigataPrance,sp. nov. Type.
Brazil.Amazonas:Manaus-CaracaraiRd., km
45, Reserva Biologica do INPA, 4 Apr 1972
(fl),M. F. Silva&L. Coelho80 (holotype,INPA
35442; isotype, NY). Fig. 8.
42 FloraNeotropica

Species sectione Hymenopus pertinens, a L. servesof the INPA/WWFMinimumCriticalRe-

reticulata fructibus globosis exocarpio tomen- serve Size Projectnear to Manaus.It is now ap-
tellis, foliis laevigatis haud reticulatis, floribus parentthattheyaredifferentspeciesthatareeasily
extus brunneotomentellis sessilibus differt. distinguishedin the field.
Treeto 20 m tall, the youngbranchesglabrous. Licania laevigatais closest to L. reticulatabut
Leaf lamina oblong, coriaceous, 9-18 x 4.2-7 differsin the smooth, not reticulateleaves that
cm, cuneateat base, acuminateat apex, the acu- are more cuneate at the base, the round tomen-
men 4-10 mm long, glabrouson both surfaces, tose fruit ratherthan an ellipsoid, glabrousand
shiny above; midrib prominulousabove, prom- costate one, the flowersborne sessile on the pri-
inent beneath; primary veins 7-10 pairs, pro- mary inflorescencebranchesand the brown to-
minulous on both surfaces,widely spaced with mentellous pubescence of the exterior of the
1.2-1.8 mm betweenveins;petioles5-8 mm long, flowers.
with confluent leaf base, glabrous, eglandular.
Stipulesaxillary,caducous.Inflorescencesof ter- 2-69.2. LicaniaoccultansPrance,sp. nov. Type.
minal and subterminalracemose panicles, the Brazil. Amazonas: Manaus-ItacoatiaraRd.,
rachispuberulous.Bractsand bracteolesminute, km 31, CEPLACResearchStation,6 Nov 1973
triangular,tomentose, subpersistent.Flowersca. (fl), Steward & Ramos P17669 (holotype,
2 mm long, inserted on primary inflorescence INPA; isotype, NY). Fig. 9.
branches. Receptacle urceolate, brown-tomen- A L. heteromorphafoliis longe acuminatis,pe-
tellous on exterior, tomentose within; pedicels tiolis eglandulosis,laminis confluentibusdiffert.
0.5-1 mm long. Calyx lobes five, triangular,to- Treeto 30 m tall, the youngbranchesglabrous,
mentellous on both surfaces.Petals five, pubes- with largeconspicuouslenticels.Leaf laminaob-
cent on exterior, with ciliate margins. Stamens
long-elliptic,chartaceous,4.5-8 x 2.5-4 cm, cu-
6-7, insertedaroundthree-fourthsof circle with neate at base, acuminateat apex, the acumen 4-
tooth-like staminodesopposite;filamentsshort- 9 mm long, cuspidate,glabrouson both surfaces;
er than calyx lobes, free, glabrous;anthers del- midrib prominulousabove, prominentbeneath;
toid. Ovary inserted at base of receptacle, to-
primaryveins 5-7 pairs, prominulous on both
mentellouson exterior;style pubescenton lower
surfaces,widely spaced;petioles 2-5 mm long,
portion. Young fruit globose, exocarp densely with confluent leaf base, glabrous, eglandular.
short-ferrugineous-tomentellous. Stipulesaxillary,small, caducous.Inflorescences
Distribution (Fig. 62). Terra firme forest of of terminaland subterminalracemosepanicles,
CentralAmazonia and Surinam. the rachis sparselygray-puberulous.Bracts and
Additionalmaterial. SURINAM. FrederikTop, 3 bracteolestriangular,minute, 0.5 mm long, per-
km SSEof JulianaTop, 23 Aug 1963 (youngfr), Irwin sistent, puberulous. Flowers ca. 1.5 mm long,
et al. 54990(NY). insertedon primaryinflorescencebranches.Re-
BRAZIL.AMAZONAS:Sao Gabriel,27 May 1948 (y
fr),Black48-2902 (IAN,NY);ReservaFlorestalDucke, ceptacle campanulate, sparsely puberulous on
Manaus,6 Apr 1967 (fl), Byron& Elias 67-20 (INPA, exterior,tomentellouswithin;pedicelsca. 0.5 mm
NY), 27 Mar 1952 (fl), L. Coelho s.n. (INPA 5209, long. Calyx lobes five, triangular,puberulouson
NY), 29 Sep 1976 (st), Mello s.n. (INPA 60170); 24 both surfaces.Petals five, sparselypubescenton
Sep 1976 (st),Adairde Oliveiras.n. (INPA 74791), 19 exterior, the
Aug 1976 (st),Adairde Oliveiras.n. (INPA 60162), 16
margins ciliate. Stamens 5-6, in-
Jan 1976 (st), Adairde Oliveiras.n. (INPA 59936), 14 serted around three-fourthsof circle; filaments
Sep 1971 (y fr), Pranceet al. 14735 (INPA, NY). shorter than calyx lobes, free, glabrous. Ovary
insertedat base of receptacle,puberulouson ex-
This species and the next (Licania occultans)
terior;stylevillous on lowerhalf.Fruitunknown.
are both close to Licania heteromorpha and I
Distribution(Fig. 62). Forest on terrafirmeof
have previously placed some of the material
CentralAmazonia.
within that species. The description of these
species begins the dismemberment of that large Additionalspecimenexamined.BRAZIL.AMAZONAS:
and morphologically diverse species. The sepa- Maues,57?43'W,3?23'S,23 Jul 1983(fl),S. R. Hill
ration of L. laevigata and L. occultans is made
13163(INPA,NY).
possible by the collection of large population This species differs from Licania heteromor-
samples from the Reserva Ducke and the Re- pha in the long acuminateleaves which are char-
Systematic Treatment 43

3cm.[

.. ' .-.. . . .. .- .

~,a',,,,,?W'
.,',,'4g,,i..~';/~ / .
I ,4< (
44 Flora Neotropica

taceous and conduplicate,and in the eglandular actuallymuch more widespread,but only infre-
petioles with the leaf base confluentinto them. quentlycollected.It has been collectedin several
places in Colombia and Venezuela. See Figure
2-72. Licania fanshawei Prance, Fl. Neotrop. 78.
Monogr.9: 112. 1972. Additionalspecimensexamined.COLOMBIA.META:
Sierradela Macarena, CentralMountains, northridge,
Distribution (Fig. 38). This species was de- 30 Dec 1949
(fr),Philipson& Idrobo2005 (BM, NY).
scribedfrom materialcollectedin Venezuelaand VENEZUELA.TACHIRA: Rio San Buena, 10 km W
Guyana. Recent collections show it to be dis- of La Fundaci6n,15 Mar 1980 (fr),Liesneret al. 9584
tributedthroughoutthe Guianas. (MO, NY, VEN). ZULIA:Km 45 of Lara-ZuliaRd., 31
Mar 1979 (fl, fr), Bunting& Fucci 7124 (NY).
Additional specimens examined.SURINAM. No lo-
cality, 13Nov 1971(fr),Jimenez-Sda1608 (LBB,NY);
Fallawatra,22Nov 1971(st),Jimenez-Sda 1642(LBB, Subgenus Licania Section Cymosa
NY).
FRENCH GUIANA.Saiil,MontsLaFum6e,27Aug 2-78.1. Licania arianeaePrance,sp. nov. Type.
1982 (fl), Mori & Boom 14780 (NY). Brazil. Espirito Santo: Reserva Florestal
CVRD, Linhares,19 May 1980 (fl), D. A. Foli
2-73.1. Licania marleneaePrance,Brittonia28: 228 (holotype, RB; isotype, NY).
218, fig. 7. 1976. Type. Brazil. Amazonas: Figs. 10, 27.
Manaus-PortoVelho Hwy. between Rio Cas- Species sectione Cymosae pertinens;a L. cu-
tanho and Rio Tupana, 18 Jul 1972 (fl),M. F. prea staminibus 6-7, stipulis 6-10 mm longis,
da Silva 873 (holotype, INPA). floribus 5-6 mm longis, a L. riedeliifloribusin
Treeto 15 m tall, the youngbranchesglabrous, cymulis 2-3-floris dispositis, stipulisfloribusque
lenticellate. Leaf lamina, oblong-elliptic,coria- maioribus,foliis oblongo-ellipticisdiffert.
Treeto 20 m tall, the young branchesdensely
ceous, 5.5-9.5 x 3-4.7 cm, cuneateat base,acute
at apex, glabrouson both surfaces;midrib plane ferrugineous-tomentose. Leaf lamina oblong to
oblong-elliptic, coriaceous, 7-11 x 3.5-5 cm,
above, prominulousbeneath; primaryveins 7-
11 pairs, plane or prominulousabove, promin- rounded to subcuneateat base, bluntly acumi-
ulous beneath;petioles 2-3 mm long, glabrous, nate or acute at apex, the acumen 0-5 mm long,
with conspicuous deep-set sto-
rugulose,eglandular.Stipulesacute,ca. 0.75 mm glabrousabove,
matal cavities beneath, filledwith a lanatewhite
long, persistent, sparselypuberulous,adnate to
base of petiole. Inflorescencesof terminal and pubescence; midrib impressed for entire length
subterminalbranched panicles; rachis sparsely above, prominent and ferrugineous-tomentose
puberulous, becoming glabrous with age, the beneath;primaryveins 10-15 pairs,planeabove,
branchespuberulous.Bracts and bracteoles ca. prominentand with a few sparsehairs beneath;
0.5 mm long, persistent,triangular,puberulous petioles 6-8 mm long, densely ferrugineous-pu-
on exterior.Flowers2-2.5 mm long. Receptacle bescent,canaliculate.Stipules6-10 mm long, 1.5-
2 mm broad at base, lanceolate,persistent,ad-
campanulate,sessile, sordid-tomentellouson ex-
tomentose within. lobes nate to extremebase of petiole. Inflorescencesof
terior, Calyx five, acute,
tomentellous on exterior, puberulous within. terminaland subterminalpanicles,the rachisand
Petals absent. Stamens five, unilateral,inserted branchesdenselyferrugineous-tomentose. Bracts
and bracteoles minute, ca. 1 mm long, persistent.
opposite to three sepals; filaments glabrous,
shorterthan calyx lobes. Ovarypilose. Style gla- Flowers 5-6 mm long, inserted in small groups
brous. Fruit unknown. ofcymules attachedto primarybranchesby short
Distribution(Fig. 60). Known only from the thick peduncles; receptacle urceolate, slightly
type, from rain foreston terrafirme,collectedin gibbous, tomentose on exterior, lanate within.
flowerin July. Calyx lobes acute, tomentose on both surfaces.
Petalsabsent. Stamens6-7, unilateralwith large
2-76. Licaniasilvae Prance,Fl. Neotrop. Mono- lanate ridge opposite bearingshort staminodes,
filaments shorter than calyx lobes, unequal in
gr. 9: 115. 1972.
length. Ovaryinsertedat base of receptacle,vil-
This species, described from material from lous; style exceeding filamentsin length, lanate
BrazilianAmazonia in Para and Amazonas, is for entire length. Fruit not seen.
Systematic Treatment 45

11

3cm.

FIG. 10. Licania arianeae (Foli 228). A, habit; B, leaf undersurface; C, flower; D, flower section; E, stamen;
F, ovary.
F, ovary.

Additional specimen examined. BRAZIL. ESPIRITO is closest to the Guiananspecies Licania cuprea,
SANTO: Reserva Florestal de Linhares, 22 May 1972 but differs in the largerflowers, larger stipules,
(fl), Lino 45 (NY, RB).
gibbous receptacle,leaf shape, and greaternum-
This most distinct species belongs to section ber of stamens. It is one of the few species of
Cymosa, but has thicker peduncles bearing the Licania with extremely conspicuous, exposed
cymules than in other species of that section. It stomatalcavities on the leafundersurface;in this
46 Flora Neotropica

it superficiallyresembles L. riedelii, but differs base of filaments,tomentose on lower half. Fruit

in the inflorescenceof cymules and the much not seen.
largerstipules. Distribution(Fig. 77). Known only from the
type collection, from the seaside.
2-79. Licania impressa Prance, Fl. Neotrop. Licania santosiibelongsto section Cymosaon
Monogr.9: 118. 1972. accountof the cymose inflorescencebranches.It
is closely related to L. dealbata, from which it
Distribution.Figure49. differsin the broaderleaves with blunterapices,
Additional specimens examined. BRAZIL. the longer petioles, the pubescenceof the lower
AMAZONAS: Manaus-Itacoatiara Rd.,km69-70, 5 Sep surfacenot obscuringthe venation, and the gray-
1973(fl),Pranceet al. 17520(INPA,NY);Rio Cuiei- ish-brownpubescenceof the inflorescence.It dif-
ras,2 kmbelowRioBrancinho, 12Sep1973(fl),Prance
et al. 17820 (INPA, NY). PARA: Serrado Santar6m,18 fers from L. hypoleuca,another species of the
Aug1916(fl),DuckeMG16355(MG);TomeAqiu,29 same section occurringin the same region,in the
Dec 1977(fl),Nascimento 340(MG,NY),30Dec 1977 stomatalcryptsand conspicuouslyreticulateve-
(fr),Nascimento 363 (MG,NY). nation, the rounderleaves with blunt apices, and
the less branchedinflorescencewith largerflow-
2-80.1. Licania santosii Prance, Revista Brasil. ers and more stamens.
Bot. 2: 28, fig. 1. 1979. Type. Brazil. Bahia:
Itacare, 15 Oct 1968 (fl), J. Almeida 150 & T. 2-81. Licaniapallida Spruceex Sagot,Ann. Sci.
S. dos Santos 150 (holotype,CEPEC;isotypes, Nat. Bot., ser. 6, 15: 306. 1883.
AAU, FHO, MO, NY, U, US).
The first Peruviancollection of this Guianan
Small tree 4 m tall, the young branchesgla- and eastern Amazonian species has been col-
brous, conspicuously lenticellate. Leaf lamina lected (Fig. 69); see also Licania arachnoidea.
ovate-orbicular, coriaceous, becoming thickly Additional
coriaceouswith age, 6-10 x 4-6.5 cm, rounded Rio specimenexamined.PERU.AMAZONAS:
Santiago,65 km N of Pinglo,4 Dec 1979 (fl),
at base, rounded to slightly acute at apex, gla- Huashikat1425(MO,NY).
brous on upper surface,the lower surfacewith
stomatal crypts filled by dense lanate pubes- 2-84. Licania cymosa Fritsch, Ann. K. K. Na-
cence, the venation reticulate and conspicuous turhist.Hofmus. 4: 47. 1889.
since it is not coveredby the pubescence;midrib
plane above, prominentbeneath;primaryveins Thislittleknownspecieswasstudiedfromthree
7-8 pairs, plane above, prominentbeneath;pet- highly disjunctcollectionsin 1972. Severalmore
ioles 8-10 mm long, terete, sparselypuberulous from Bahia now have been collected (Fig. 35).
when young, eglandular,slightly rugulose.Stip- Additionalspecimensexamined.BRAZIL.BAHIA:
ules adnate to extreme base of petioles, persis- Mun.Ilheus,FazendaBarrado Manguinho,29 Sep
tent, lanceolate,ca. 2 mm long. Inflorescencesof 1980 (fl),MattosSilva et al. 1136 (CEPEC,NY), 5 Feb
little-branchedpanicles 3-4 cm, the rachis and 1982 (fr),MattosSilva et al. 1401 (CEPEC,NY); Una,
2 1971(fr),Pinheiro1558(NY).
branchesbrown-tomentellous.Bractsand brac- Oliven9a, Sep
teoles 1-2 mm long, ovate, persistent, tomen- Local name. oiti.
tellous on exterior.Flowersca. 2.5 mm long, in
few-floweredcymulesattachedto rachisand pri- 2-87. LicaniapiresiiPrance,Fl.
Neotrop.Mono-
mary branches by long slender peduncles. Re- gr. 9: 124. 1972.
ceptacle campanulate, brown-tomentellous on
exterior,tomentosewithin;pedicels0.2 mm long. Distribution(Fig. 73). This species was de-
Calyxlobes five, acute,tomentellous-puberulous scribedfrom threecollectionsfrom Amapa,Bra-
on both surfaces.Petals absent. Stamens seven, zil. It has recentlybeen collectedin Guyanasince
insertedaroundcomplete circle, anthersand fil- my accountfor the floraof the Guianaswas pub-
aments of variable size; filaments shorter than lished (Prance, 1986b).
calyxlobes, glabrous,freeto base. Ovaryinserted Additionalspecimensexamined.GUYANA.WEST
at base of receptacle,villous. Style extendingto DEMERARAREGION:Mabura Hillarea,5?20'N,58?40'W,
Systematic Treatment 47

3 Jun 1986, Pipoly 7513 (NY, US), 4 Jun 1986, Pipoly 2-94. LicaniaurceolarisHookerf., Fl. bras.14(2):
7524 (NY, US), Pipoly 7537 (NY, US).
15. 1867.

2-87.1. Licania furfuraceaPrance,Brittonia28: This species has now been collected over a
221, fig. 8. 1976. Type. Venezuela.Bolivar:El much wider area includingPeru (Fig. 84).
Dorado-SantaElena Rd., km 251-253, 4 Jan Additionalspecimensexamined.COLOMBIA.
1975 (fl), Steyermark113900 (holotype, NY; Florencia,Rio Orteguaza,21 Mar 1965 (fr),
CAQUETA:
isotype, VEN). Garcia-Barriga18202 (COL), Garcia-Barriga18209
(COL,US).
Treeto 20 m tall, the young branchessparsely PERU.LORETO: Requena,Rio Ucayali,Arboretum
Jenaro Herrera,Jul-Sep 1976(fl),Bernardi5-36 (G,
puberulous,becoming glabrousand lenticellate NY);Yanomono,Explorama TouristCamp,trailto
with age. Leaf lamina oblong, chartaceous,3.5- RioNapo,19 Feb1981(fr),Gentryet al. 31502(MO,
5.5 x 1.5-2.5 cm, subcuneateat base, acuminate NY);Maynas,PuertoAlmendras,14 Feb 1977 (fr),
at apex,theacumen3-7 mm long,glabrousabove, Revilla2345 (MO, NY).
with a waxy pulverulent-furfuraceouspubes-
cence beneath; midrib plane or prominulous 2-95. Licania affinis Fritsch, Ann. K. K. Na-
above, prominentbeneath;primaryveins 9-10 turhist.Hofmus. 4: 50. 1889.
pairs, plane above, prominulousbeneath; peti- Distribution(Fig. 22). This species was de-
oles 2.5-5 mm long, sparselypuberulous,can- scribed from
Guyana. Quite common in the
aliculate. Stipules axillary, 1-2 mm long,
Guianas, it was reportedonly from that region
lanceolate, sparsely puberulous,persistent. In- and adjacent Brazil in Prance (1972). Later it
florescencesof terminal and axillarycompound was collected in Panama and added to the in-
panicles,the flowersbornein small 2-3-flowered creasing list of Panama-Guiana disjuncts, as
cymuleson shortpeduncles;rachisand branches pointedout in Prance(1974a). Its CentralAmer-
puberulous.Bractsand bracteolesca. 5 mm long, ican rangehas recentlybeen expandedinto Costa
oblong, persistent, tomentellous on exterior. Rica, showingthat it exists in suitableforestsites
Flowersca. 2 mm long. Receptacleurceolate,to- well into CentralAmerica.
mentellouson exterior,appressed-puberulous on
interior;pedicels 0.25 mm long. Calyx lobes five, Additionalspecimensexamined.COSTARICA.
HEREDIA: Rio Peje,SW sectorof La Selva,24 May
acute, tomentellous on exterior, puberulous 1982
(fl), Hammel 12476 (NY).
within. Petals absent. Stamens five, unilateral, PANAMA.PANAMA:El Llano-Carti Rd., 10 km N
inserted opposite three sepals; filaments gla- of PanAmerican Hwy.,12Dec 1973(fl),Gentry8884
brous,freeto base,shorterthancalyxlobes.Ovary (MO, NY). COLON:SantaRita, Feb 1968 (fl), G6mez-
tomentose. Style puberulous.Fruit unknown. Pompaet al. 3347 (MEXU),Gomez-Pompaet al. 3354
10 1971 (fr),Lao et al. 10 (MO, NY).
Distribution(Fig. 40). This species is known (MEXU), Aug
from the type gathering,collected in flower in
2-95.1. Licaniateixeirae Prance,sp. nov. Type.
Januaryin tall forest, beside a small stream. Brazil. Rondonia: Minera~aoSanta Barbara,
25 May 1982 (fl), L. O. A. Teixeira 728 (ho-
lotype, INPA 104465; isotypes, FHO, NY).
Subgenus Licania Section Pulverulenta
Fig. 11.
2-90. LicaniabuxifoliaSandwith,Bull.Misc. In-
form. 1931: 369. 1931. Species sectione Pulverulentaepertinens, L.
urceolariaaffinis,foliis chartaceis4-5.5 mm lon-
Distribution(Fig. 29). This species,only known gis, apice caudato, receptaculourceolato, extus
from three collections in Prance (1972), is still brunneo-tomentosodiffert.
known only from a restrictedarea of Guyana. Tree,the youngbranchesglabrous,lenticellate.
Leaf lamina narrowlyovate, chartaceous,4-5.5
Additionalspecimensexamined.GUYANA.Ma- x 1.7-2.3 cm, subcuneateand slightly unequal
zaruni-PotaroDistrict,Bartica-PotaroRd., 24 mi S of
at base, caudateat apex, the acumen 1.2-1.8 mm
Bartica, 15 Aug 1976 (fl), Mori et al. 8088 (NY); 19
mi SW of Bartica, 17 Aug 1976 (fl), Mori et al. 8131 long, glabrousabove, with a waxy pulverulent-
(NY). farinaceous pubescence beneath; midrib gla-
48
?..-n
FIG. 11. Licania teixeirae (Teixeira 728). A, habit; B, flower;C, flower section; D, apical view of flower;
E, leaf undersurfaceshowing pulverulentpubescence.
brous and plane above, prominentbeneath;pri-
mary veins 7-9 pairs, inconspicuous on both
surfaces;petioles 4-6 mm long, glabrous, can-
aliculate. Stipules linear, 1 mm long, adnate to
base of petiole. Inflorescencesof terminal and
axillarylittle-branchedracemose panicles or ra-
cemes, the rachisand branchestomentellous,the
apicalpanicles3-4 cm long, the axillaryracemes
1-2 cm long. Bracts and bracteoles lanceolate,
persistent,ca. 0.5 mm long, tomentellous.Flow-
ers ca. 3 mm long, sessile on primarybranches
of inflorescence.Receptacleurceolate,sessile, ru-
fous-brown-tomentellouson exterior, densely
tomentellouswithin. Calyx lobes five, acute, ru-
fous-brown-tomentellouson exteriorcontrasting
sharply with gray-white-tomentellous pubes-
cence of inner surface. Petals absent. Stamens
five, inserted around 2/3 of circle, the filaments
shorter than calyx lobes, tomentellous. Ovary
Flora Neotropica
D.
. .. 2mm t r
inserted at base of receptacle, sparsely tomen-
tellous on exterior.Style equallingfilaments,la-
nate-pubescent.Fruit unknown.
Distribution(Fig. 81). Known only from the
type collection.
This speciesis quite distinctfrom otherspecies
of Licania and is easily recognized in the dry
condition by the contrastingcolors of the recep-
tacle and interiorof the calyx lobes. It is a mem-
ber of section Pulverulentaand differs from all
other species in the leaf shape and the thin char-
taceousleaves. This speciescomes fromthe same
locality as another recentlydescribedspecies of
Licania, L. bellingtonii.
Subgenus Licania Section Licania
2-100. LicaniacouepiifoliaPrance,Fl. Neotrop.
Monogr.9: 134. 1972.
Systematic Treatment 49

Distribution(Fig. 32). This species was de- Distr., Falawatra,Nov 1971 (fl),Jimenez-Sda
scribedfrom a single collection from Guyana.A 1549 (holotype,NY; isotype, LBB 14282).
secondcollectionfromSurinamextendsits range. Treeto 26 m tall, the young branchestomen-
Additionalspecimenexamined.SURINAM. Sipali- telloussoon becomingglabrous.Leaf laminaob-
wini area (fr), Oldenburgeret al. 1256 (LBB). long-elliptic,chartaceous,14-19 x 6-7.5 cm;base
subcuneate,apex acuminate,the acumen 10-18
2-104. Licania hebantha Martius ex Hooker f., mm long, glabrous above, densely lanate be-
Fl. bras. 14(2): 17. 1867. neath, deeply reticulatevenation beneath;mid-
rib impressedabove, tomentelloustowardsbase,
Distribution. Figure 46.
prominentbeneath;secondaryveins 9-11 pairs,
Additional specimens examined. COLOMBIA. plane above, prominentbeneath;petioles 15-20
Monochoa,Sabanade Mosco, 30Dec 1976
AMAZONAS: mm long, weakly canaliculate, shortly tomen-
(fl),Sastre&ReaichelD. 5060 (P);Rio Caqueta,Arara- tellous, with two to many glands towardsbase.
cuara,7 Jan 1977 (fl), Sastre&ReaichelD. 5178 (NY,
P). Stipulesnot seen. Inflorescencesof terminaland
axillary panicles, ca. 2.5 cm long; rachis and
2-105. Licania steyermarkii Maguire, Fieldiana, branches tomentellous. Bracts and bracteoles
Bot. 28: 254. 1952. ovate-lanceolate,0.6 mm long, persistent, pu-
bescent. Flowers 1.5 mm long. Receptaclecam-
Distribution. Figure 80. panulate, tomentellous on exterior, tomentose
Additional specimens examined. VENEZUELA. within; pedicels 0.4 mm long. Calyx lobes five,
AMAZONAS:Dept.Rio Negro,lowerRio Pacimoni,8 tomentose. Petals absent. Stamens seven, uni-
Feb 1981 (fl), Huber & Medina 5847 (NY, VEN). lateral, included, three large and fertile, four
BOLiVAR:Betweenkm 251.5 and 253 road El Dorado, smaller and sterile; filaments glabrous, free to
SantaElena,4 Jan 1975 (st),Steyermark111391 (NY); base.Ovarylanate-pubescent.Stylelanate.Fruits
Los Brasileros,16
4.5 km SWof Icabaru,Quebrada
Dec 1978 (fl), Steyermarket al. 117776 (NY). pyriform,to 5 cm long including stipe of 1.5-2
cm; exocarpvelutinous,ferrugineous-pubescent;
2-107. Licania crassiveniaSpruceex Hooker f., mesocarpthin; endocarp 1 mm thick, hard and
Fl. bras. 14(2): 14. 1867; Prance,Fl. Neotrop. fibrous,sparselyhirsutewithin.
Monogr.9: 137. 1972. Fig. 32. Distribution(Fig. 50). Known only from the
rain forests of Guyanaand Surinam.
Until recentlythis specieswasknownonlyfrom
the Sprucetype. The materialcited below keys Additionalspecimensexamined.GUYANA.Mu-
rudiMts., MazoaHill, 2?15'N,59?10'W,12 Nov 1982
to L. crassiveniaand is included there for the (fl), Stofferset al. 30143 (NY, U).
present,but has much largerleaves than the type SURINAM.SipaliwiniSavannaarea,forestW of
(10-13 x 4-5.5 cm as comparedto 5-8 x 1.5- Meyers farm, Jan 1970 (fr), Oldenburgeret al. 1213
3.8) and a yellow-brown-tomentellousinflores- (NY, U), Oldenburgeret al. 1225 (NY, U).
cence ratherthan gray. The leaves of the Her- Local name. rode kwepi.
ndndez& Dezzeo materialare exactly the same This species is most distinct and not easily
shape as those of the type, graduallytapering confusedwith any other species. The numerous
towardsthe apex from nearto the base;they also axillaryinflorescences,and the two types of sta-
have the extremely conspicuous stomatal cavi- mens distinguishthis species.The largerstamens
ties that are made obvious by the surrounding are fertile,the smallerones have well-developed
glabrousvenation. This could be two different anthersbut abortivepollen. Licaniajimenezii is
species but I hesitate to describe another one probably closest to L. alba and L. robusta. It
until further material is collected to show the differsfrom L. alba in the much smaller brac-
morphologicalvariation. teoles and flowers,the less conspicuousleaf re-
Additionalspecimenexamined.VENEZUELA. ticulation, the longer petioles, and the stamens,
BOLiVAR: Sorochoroyen, Sifontes Dept., 5?13'N, and from L. robustain the smaller flowers, the
611 1'W, 20 Apr 1985 (fl), Herndndez& Dezzeo 128 pubescent inflorescence,the glandularpetioles,
(MYF,NY). the caducousstipules,and the impressedmidrib.

2-108.1. Licania jimenezii Prance, Acta Ama- 2-117. Licania ovalifolia Kleinhoonte, Recueil
zonica 2(1): 7. 1972. Type. Surinam.Nickerie Trav. Bot. Neerl. 30: 180. 1933. Fig. 69.
50
Additional specimens examined. SURINAM. Fal-
lawatra, 7 Jan 1972 (st), Jimenez-Sda 1688 (LBB, NY).
BRAZIL. AMAPA:Contagem, Oct-Dec 1976 (st), Rosa
1061 (MG).
2-119.1. Licania stewardiiPrance,Brittonia28:
223, fig. 9. 1976. Type. Brazil. Amazonas:
Manaus-CaracaraiRd., km 130, 13 Feb 1974
(fl), Steward et al. P20251 (holotype, INPA;
isotypes, FHO, NY, US).
Small tree 3-5 m tall, the young branchespu-
berulous, soon glabrate. Leaf lamina oblong,
chartaceous,4-8 x 2.2-4 cm, rounded or sub-
cordateat base, rounded,acute or retuseat apex,
glabrousabove with a waxy lanateappressedpu-
bescence beneath; midrib slightly impressed
above, prominent beneath; primary veins 7-9
pairs, plane above, prominentbeneath, the sec-
ondary venation conspicuouslyreticulate;peti-
oles 1.5-2.5 mm long, terete,eglandular,tomen-
tellouswhenyoung.Stipuleslinear,2-3 mm long,
persistent,tomentellous, adnate to petioles. In-
florescences of terminal or subterminal little
branchedracemose panicles or simple racemes
2-7 cm long; rachis and branchesgray-tomen-
tellous. Bracts and bracteoles narrowlylanceo-
late, 1-2 mm long, tomentellouson exterior,per-
sistent. Flowers 3-3.5 mm long. Receptacle
campanulate,gray-tomentellouson exterior,la-
nate within; pedicels 0.25-0.5 mm long. Calyx
lobes five, acute, gray-tomentellouson exterior,
puberulouswithin. Petals absent. Stamens9-11,
insertedaroundcomplete circle,shorterthan ca-
lyx lobes; filamentsconnate at base, the united
partlanate.Ovarydenselylanate.Styleequalling
filaments.Fruit unknown.
Distribution (Fig. 80). This species is known
only from the vicinity of the type locality where Additionalspecimensexamined.BRAZIL.PARA:Rio
it wasverycommonin low campinaforestaround
Tocantins, Tucurui, Igarap6
a sandstone rock outcrop in an area where the Lisboa et al. 1382 (MG, NY), 21 Aug 1980 (fr), Ro-
originalforest has now been destroyed. drigueset al. 10259 (INPA).
Additional specimens examined. BRAZIL.
AMAZONAS: Manaus-Caracarai Rd., km 130, 16 Dec
1975 (fl), D. Coelho et al. 708 (INPA 54246, NY), 25
Aug 1976 (fl), Davis & D. Coelho 60303 (UEC), 13 Feb
1974 (fl), A. Loureiro et al. s.n. (INPA 47911, NY), 15
Feb 1974 (fl), INPA 47955 (INPA), 6 Jan 1976 (fl),
Monteiro & Ramos 33 (INPA 54344, NY), 25 May
1974 Rodrigues et al. 9284 (INPA), 22 Mar 1978 (y
fr), N. T. Silva 4578 (MG, NY); km 350, 18 Nov 1977
(fl), Steward 88 (INPA, NY), 14 Feb 1974 (fl), Steward
P20326 (FHO, INPA, NY), 16 Feb 1974 (fl), Steward
FloraNeotropica
P20401(FHO,INPA,NY),4 Oct 1977(fl),Ramos&
D. Coelho755 (INPA).
2-121.1. Licania tocantina Prance, Acta Ama-
zonica 13: 28. 1983. Type. Brazil. Para: Rio
Tocantins,Tucurui,BreuBranco,11 May 1978
(fl), M. G. Silva & R. Bahia 3508 (holotype,
MG; isotypes, INPA, NY). Fig. 82.
Tree 20 m tall, the young branches sparsely
puberulous,soon glabrate.Leaf lamina elliptic,
chartaceous,8-15 x 3.5-7.2 cm, subcuneateat
base, acuminateat apex, the acumen 7-12 mm
long,glabrousabove, with stomatalcavitiesfilled
by denselanatepubescencebeneath;midribplane
or slightly impressed above; petioles 4-6 mm
long, with two large conspicuous glands when
young, canaliculate,tomentellous when young,
becoming glabrousand rugulosewith age. Stip-
ules triangular,ca. 5 mm long, ca. 2.5 mm broad
at base, inserted on base of petioles, persistent.
Inflorescencesof terminal and subterminalra-
cemose panicles, the rachis and branches pu-
berulous. Bracts and bracteoles 1-2 mm long,
triangular,persistent, puberulous on both sur-
faces. Flowers ca. 1.5 mm long, sessile on pri-
marybranchesof inflorescence.Receptaclecam-
panulate, sessile, tomentose on exterior and
within. Calyx lobes five, acute, puberulouson
both surfaces.Petals absent. Stamensthree,uni-
lateral;filaments shorter than calyx lobes, gla-
brous. Ovary inserted at base of receptacle,pi-
lose. Style equalling filaments, hirsute three-
fourths of length. Fruit pyriformca. 2 cm long
(5 cm stipe); exocarp ferrugineous-pubescent;
mesocarpthin; endocarphard, granular,1 mm
thick.
2 Feb1980(fl),
Cagancho,
This species is most closely relatedto Licania
triandra, from which it differs in the thinner,
chartaceousleaves with deep stomatal cavities
covered by a lanate pubescence,which also ex-
tendsoverthe venation,makingthe cavitiesmuch
less conspicuousthan in L. triandra.It also dif-
fers in the broaderstipules adnateto the petiole
and in the petiole rather than leaf bases glan-
dular.
SystematicTreatment51
2-124. Licania micrantha Miquel, Stirp. suri- iform;exocarpglabrous,wrinkledwhen dry, the
nam. select. 20. 1850. stipe 5-10 mm in dry fruit.
Distribution(Fig. 25). This species is appar-
Distribution(Fig. 63). This species is wide- ently endemic to the summit of the sandstone
spreadin Venezuela, Amazonia, and the Guia- mountain Serra Araca at about 1000 m, flow-
nas. Its knownrangehas now been extendedwest ering in February.
of the Andes from recentcollectionsfrom Choc6
and Valle in Colombia, and also to Atlantic Additional specimens examined. BRAZIL.
AMAZONAS: SerraAraca, 1 Feb 1978 (fr),Rosa & Lira
coastal Brazilin the forests of Bahia. 2337 (IAN, NY), 22 Feb 1984 (fr), Tavareset al. 114
(INPA,NY).
Additionalspecimensexamined.COLOMBIA.
CHOCO: RioFujiad6,afluente
delRioSanJuan,4?36'N, 2-128. Licania bracteata
76?54'W,7 Apr 1979 (fl), Forero et al. 4788 (COL,
Prance, Fl. Neotrop.
MO). VALLE:Mun. Buenaventura,Concesi6n Cart6n Monogr. 9:155. 1972.
de Colombia, 15 Nov 1979 (fl), vanRoodenet al. 358
Distribution(Fig. 29). This species was de-
(NY, U), 6 Dec 1979 (fl), van Rooden et al. 553 (NY,
U). scribed from considerablematerialfrom the vi-
BRAZIL.BAHIA: Mun. Una, Maruim,28 Apr 1981 cinity of Manaus, Brazil,where it is quite com-
(st),Moriet al. 13774(CEPEC, NY), 1 May1981(st), mon in the forestson terrafirme.Recentlyit has
Mori et al. 13884 (NY), 13 May 1981 (st), Mori et al. also been collected in Peru.
14000(NY);Esta;aoExperimental LemosMaia,24
Oct 1980 (fr),Rylands42/80 (NY), 11 Nov 1980 (st), Additionalspecimensexamined.PERU.LORETO:
Rio
Rylands56/80 (NY). Nanay,SantaMaria,30 May1963(fl,fr),Ar6steguiV.
105(US);RioNanay,28 Oct 1965(fl),A. G.Ruiz224
(NY,US).
2-126.1. LicaniaaracaensisPrance,Brittonia28:
223, fig. 10. 1976. Type. Brazil. Amazonas: Local name. Peru:parinari.
SerraAraca, 1?N,63?W,10 Feb 1975 (fl),Pires
15027 (holotype, IAN; isotypes, INPA, NY). 2-130.1. Licania lamentanda Prance, sp. nov.
Tree2 m tall, the young branchespuberulous, Type. Brazil.Bahia:Municipalityof Ilheus, 4
km N of Oliven9a on road to Ilheus, 19 Apr
soon becoming glabrous and lenticellate. Leaf
1981 (fl), S. A. Mori et al. 13673 (holotype,
lamina oblong, conduplicate,coriaceous, 3-5.5
x 2-2.8 cm, subcuneateat base, acuteto bluntly CEPEC; isotype, NY). Fig. 12.
acuminate at apex, the acumen 2-6 mm long, Species sectione Licania pertinens.Folia 12-
glabrous above, densely lanate-tomentose be- 16.5 cm longa, petiolo 1.1-1.5 cm longo, velu-
neath; midrib plane above, prominulous be- tino-tomentello. Flores 4.5-5.5 mm longi, sta-
neath; primary veins 6-8 pairs, plane above, minibus 6-7, ovario densissime tomentoso.
prominulousbeneath;petioles 3-4 mm long, te- Tree,the youngbranchesshortlytomentellous,
rete, sparsely puberulous, eglandular.Stipules becoming glabrous with age. Leaf lamina ob-
persistent,lanceolate, 1-1.5 mm long, adnateto long-elliptic, coriaceous, 12-16.5 x 6-8.5 cm,
base of petiole. Inflorescencesterminaland sub- subcuneateto roundedat base, roundedto apic-
terminal racemose panicles, 4-7.5 cm long; ra- ulate at apex, glabrousabove, with a short ap-
chis and branchespuberulous.Bractsand brac- pressed-lanatebrownpubescencebeneath;mid-
teoles triangular, persistent, tomentose on rib prominulous above, prominent beneath;
exterior, ca. 0.75 mm long. Flowers ca. 2 mm primaryveins 10-12 pairs, prominulousabove,
long. Receptacle urceolate, sessile, yellow-to- prominentbeneath, secondaryvenation promi-
mentellouson exterior,tomentose towardsbase nulous,ca. more or less parallelat 90?to primary
and glabrousabove on interior.Calyxlobes five, veins; petioles 1.1-1.5 cm long, velutinous-to-
acute, tomentellous on exterior, puberulous mentellous,terete or weaklycanaliculateabove,
within. Petals absent. Stamens three, unilateral, eglandular.Stipules caducous (not seen). Inflo-
opposite three calyx lobes; filaments glabrous, rescencesof racemose panicles, 10-14 cm long,
shorterthan calyx lobes. Ovarytomentose. Style the rachis and branches shortly tomentellous.
puberulous,shorterthan calyx lobes. Fruit pyr- Bracts and bracteoles membranous,persistent,
52 Flora Neotropica

F.in(t3A,
;Clwr,foeeo.ai;B efudr

3cnr.l \I ~~~s~~llj
I 111
Illit'CV

I , . . . . . . . .

FIG. 12. Licania lamentanda (Mori et al. 13673). A, habit; B, leaf undersurface; C, flower; D, flower section.
SystematicTreatment 53

ovate, 1.5-3 mm long, tomentellouson exterior, Distribution(Fig. 56). This species, common
puberulous within. Flowers 4.5-5.5 mm long. along rivermarginsin the Guianasand northern
Receptaclecampanulate,3.5 mm long, tomen- Para, has recentlybeen found to be disjunct in
tellous on exterior,tomentosewithin, subsessile. Atlantic Coastal Brazil, where it also grows on
Calyx lobes five, acute, triangular,ca. 1.5 mm river margins. This adds yet another Amazo-
long, tomentellous on both surfaces.Petals ab- nian-Bahiandisjunct, similar in distributionto
sent. Stamens 6-7, shorterthan calyx lobes, in- Licaniacymosawhich,however,is rarerin Ama-
serted around three-fourthsof a circle. Ovary zonia.
extremelydenselytomentose,thick-walled.Style Additionalspecimenexamined.BRAZIL.BAHIA:Rio
pubescenton lowerportion,glabrousabove.Fruit Sto.Antonio,Mun.Andarai,19Jun1984(fl),Hatsch-
ellipsoid, 5-6.5 mm long x 4-4.5 cm broad; bach & Kummrow48068 (MBM, NY).
exocarp densely velutinous tomentellous; peri-
carphard,woody, ca. 6 mm thick, lanatewithin.
Distribution(Fig. 53). Restingaon sandy soil, 2-135.1. Licanianelsonii Prance,sp. nov. Type.
25 m altitude,growingwith many individualsof Brazil.Amazonas:3 km south of SerraAraca,
piaCabapalm in Bahia. 0?49'N,63?19'W,29 Feb 1984 (fl), W.A. Ro-
drigueset al. 10501 (holotype,INPA; isotype,
Additional specimens examined. BRAZIL. BAHIA: NY). Figs. 13, 62.
Mun.Ilheus,FazendaBarrado Manquinho,km 10
roadPontal-Olivenca, 5 Feb 1982(fl,fr),L.A.Mattos Speciesa sectio Licania pertinens,a L. incana
Silvaet al. 1440(CEPEC, NY);Mun.Uruguca, 28 km et L. leptostachyafoliis subtus conspicae reti-
NE of Urucucaon roadto SerraGrande,2 Dec 1979
culatis, stipulis a base peciolarum adnatis, re-
(fr),Mori et al. 13063 (CEPEC,NY).
ceptaculo rufo-pubescente,floris dense fascicu-
This most distinct new species is not easily latis, arboribusgrandisdiffert.
confused with others. It differs from related Treeto 30 m tall, the young branchesbrown-
species in section Licania in the large flowers and tomentose. Leaf lamina oblong-elliptic,charta-
leaves. The thick-walledovary with a dense al- ceous to coriaceous,3-6.5 x 1.8-3.2 cm, round-
most wall-like covering of hair and the seven ed to slightly subcordateat base, acuminate at
stamensalso distinguishit from most species of apex, the acumen 1-3 mm long, densely white-
the section.The long petiolesdifferentiateit from lanate pubescent beneath; midrib slightly im-
all except L. robusta,which has glabrousinflo- pressedabove,prominentbeneath;primaryveins
rescencebranchesand many other differences. 5-7 pairs, plane to lightly impressed above,
The specific epithet, meaning "fit to be prominentbeneath;venation conspicuouslyre-
mourned,"is derivedfrom the pitiful stateof the ticulate beneath in mature leaves; petioles 1-2
forests of eastern Brazil where most of the ex- mm long, terete, eglandular,tomentose when
citing novelties like this species are lamentably young.Stipulesadnateto base of petioles, linear,
on the verge of extinction. persistent,to 3 mm long. Inflorescencesof ter-
minal and axillary densely crowded spikes, 1-
2-131. Licania lanceolata Prance, Fl. Neotrop. 4.5 cm long,the rachisdenselytomentose.Bracts
Monogr.9:158. 1972. and bracteoles ovate, persistent, rufous-pubes-
1 mm long. Flowers ca. 2 mm long,
Distribution(Fig. 53). This species, common cent, ca.
in the VenezuelanAmazonian savannas, is ac- sessile and densely clusteredalong inflorescence
rachis. Receptacle campanulate,sessile, rufous
tually more widespread. It has now been col- tomentose on tomentose within. Calyx
lected in the savannas of Roraima, Brazil, an exterior,
lobes five, acute, cream when fresh, rufous-to-
interestingdistributionthat is seen for a number mentose on exterior when
of differentspecies, for example Barcella odora dry, pale brown-to-
mentose on interior.Petalsabsent.Stamensfive,
(Trail)Drude (Arecaceae). inserted opposite three calyx lobes; filaments
Additional specimen examined. BRAZIL. RORAIMA: shorter than calyx lobes, free, glabrous. Ovary
Manaus-Caracarai Rd., km 530, in basin of Rio Ana- insertedat baseof receptacle,denselypilose.Fruit
ua, 12 Feb 1979 (fl), Rodrigueset al. 10129 (INPA). unknown.
Habitat. Igap6 forest in sandy soil.
2-134. Licania leptostachya Bentham, J. Bot.
(Hooker)2: 220. 1840. Additionalspecimenexamined.BRAZIL.AMAZONAS:
54 Flora Neotropica

~i c, B. n

FIG. 13. Licania nelsonii (Rodrigues10501). A, habit;B, leaf undersurface;C, inflorescence;D, flowerand
bracteole;E,
bracteole; flowersection;
E, flower section;F,
F, ovary;
ovary;G,
G, floral
floraldiagram.
diagram.

3 kmsouthof SerraAraca,0?49'N,63?19'W,16 Mar This species is named for Bruce WalkerNel-

1984 (fl), Miralha67 (INPA, NY).
son, whose work in Brazilhas made possible the
This species belongs to section Licania and is Projeto Flora Amazonica series of expeditions,
most closely relatedto L. incana and L. leptos- one of which collected the material of this new
tachya. It is a much larger tree than either of species.
those species and differsin the densely crowded
inflorescenceswitha rufouspubescencewhendry,
in the stipules which are adnate to the petioles 2-136. Licania paraensis Prance, Fl. Neotrop.
rather than axillary, in the leaves which are Monogr.9: 163. 1972.
broadestquite nearto their base, and in the con- This poorlyknownspecieswas describedfrom
spicuouslyreticulateleaf undersurface. materialcollected near Santarem,Brazil(Ducke
SystematicTreatment55
RB 18818) and in Pando, Bolivia (Pranceet al. ploramaCamp,RioAmazonas,14Jul 1983(st),Gen-
6532) (Fig. 70). One collection from Manaus, try et al. 43029 (MO, NY). SANMARTIN: Distrito To-
cacheNuevo,Quebrada
de Cachiyacu,
500-600m, 9
originallyassignedto this species, belongsto Li- May 1975 (fl), Schunke V. 8439 (MO, NY).
cania impressa Prance. Rodrigues & Loureiro
7236 (INPA 15798) is actuallythe latterspecies. This species is close to Licania blackiiof east-
ern Amazonia, it differs in the small axillary,
Additionalspecimensexamined.BRAZIL.PARA: Rio caducous
Jari,MonteDourado,30 Nov 1978(fl),Santos444 stipules,the greaternumberof primary
(MG,NY);Estrada Caracaru 17Oct1969 veins, and the shorter,darkerpubescenceof the
to Munguba,
(fl), N. T. Silva 2826 (NY). flowers.It is also close to the VenezuelanL. crue-
geriana, but differs in the largerleaves with a
2-137. LicaniavaupesianaKillip & Cuatrecasas, more acuminateapex, the smaller stipules, and
Fieldiana,Bot. 27: 105. 1951. the laxer, less branchedinflorescence.
Additionalspecimenexamined.BRAZIL.AMAZONAS:
igarap6de Serrinha,12Nov 1977(fl), 2-145. Licania cruegerianaUrban, Symb. An-
Mun.Serrinha,
Damido 2637 (INPA). till. 5: 352. 1908.

2-144.1. Licania harlingii Prance, Fl. Ecuador Distribution(Fig. 34). Common in Trinidad
10: 9-10, fig. 2. 1979. Type. Ecuador.Napo: and northernVenezuela,this species is also dis-
Ca. 6 km S of Puerto Napo, 8 Apr 1969 (fl), junct in Panama.
H. Lugo S. 1054 (holotype,GB; isotype, NY). Additional specimens examined. PANAMA.COLON:
Feb 1968 (fr),Duke 15256 (MO, NY). PANAMA: 5-
Tree to 26 m tall, the young branchespuber- 623mi N of El Llano, 8 Sep 1972 (fl), Gentry5800 (MO,
ulous, soon becoming glabrousand lenticellate NY).
with age. Leaf lamina ovate-elliptic, charta-
ceous, 7-10 x 3-5 cm, subcuneateat base, acu- 2-146. Licania belemii Prance, Fl. Neotrop.
minate at apex, the acumen 8-12 mm long, gla- Monogr.9: 172. 1972.
brousabove,denselylanate-pubescent belowwith
puberulousconspicuousvenation;midribslight- Distribution(Fig. 27). This species was de-
scribed from a single specimen from Belmonte
ly impressedabove, prominentbeneath;second-
ary veins 8-11 pairs,plane above, impressedbe- in Bahia. It appearsstill to be quite common in
neath;petioles 5-6 mm long,tomentellous, terete. the forests of eastern Brazil since several new
collections have been made there.
Stipulessmall, ca. 1 mm, axillary,caducous.In-
florescencesof terminal racemose panicles, the Additional specimens examined. BRAZIL. BAHIA:
rachis and branchesyellow-brown,puberulous. BetweenUrugucaand SerraGrande,16 Jul 1978(y
Bracts and bracteoles small, triangular-hastate, fr), Mori et al. 10252 (CEPEC,NY); Niicleo Colonial
deUna,betweenBR 101SaoJoseandBA265,29 Oct
persistent,tomentellouson exterior.Flowersca. 1978
(fr), Mori & Thompson 11033 (CEPEC,NY);
1.5 mm long, borne solitary along primary Una,Fazenda SaoRafael,16Dec 1968(fl),Santos320
branches of inflorescence.Receptacle globose, (NY);Camaca,estradaRio Branco,28 Jan 1971(fl),
sessile, brown-tomentellouson exterior, lanate Santos 1444 (NY). ESPIRITOSANTO: Linares, Reserva
within. Calyx lobes acute, tomentellouson both CVRD,11Jan1979(fl),Foli61/79(INPA);RioDoce,
do 14 1934(fr),Kuhlmann208
surfaces.Petalsabsent.Stamens5-6, slightlyuni- Lag6a Durao, May
(RB 35360).
lateral;filaments free, shorterthan calyx lobes,
glabrous. Ovary inserted at base of receptacle, Local names. oiti, milho-torrado-amarelo.
tomentellous.Style equallingfilaments,glabrous
or with a few hairs only. Fruit not seen. 2-148. Licania veneralensisCuatrecasas,Field-
Distribution(Fig. 43). WesternAmazonia, in iana, Bot. 27: 109. 1951; Prance,Brittonia28:
Ecuadorand Peru. 212-215. 1976.
Additionalspecimensexamined.ECUADOR.NAPO: This species, poorly known at the time of the
Apuya,ca. 6 km S of PuertoNapo,6 Apr 1969(fl), monograph,has been describedin detail(Prance,
Lugo S. 1039 (GB, NY). PERU. MADREDE DIOS: Prov. 1976), and several additional collections have
Manu,Cerrode Pantiacolla, RioPalatoa,13Dec 1985 now been studied. It has now been shown to
(fl), Foster 10996 (F, NY); Tambopata,Rio Tambo- L. durifoliacomplex, see p. 20.
pata, 12?49'S,18 Feb 1984 (fl), Gentryet al. 45582 belong to the
(MO, NY). LORETO: Maynas Prov. Yanamono Ex- Distribution.Figure85.
56 FloraNeotropica

Additional specimens examined. COLOMBIA.
CHOCO: Cabo Corrientes,Rio Parguera,27 May 1974
(st), Warner299 (COL).NARI/O: Tumaco, vic. of Sa-
lahonda,5 Jul 1955(fr),RomeroC. 5270 (COL).vALLE:
Bajo Calima, 9 May 1961 (fl bud), CabreraR. 553
(COL), 18 Jul 1961 (fl), CabreraR. 600 (COL).
2-149. Licania amapaensis Prance, Fl. Neotrop.
Monogr. 9: 174. 1972. Fig. 23.
Inflorescence paniculate, the flowers borne in
small cymules on short secondary branches.
Young fruit ovoid, densely ferrugineous-veluti-
nous-tomentose, longitudinally striate when dry;
mesocarp thin and fleshy; endocarp hard, 1.5
mm thick in young fruit, glabrouswithin.
This very distinct species with large stipules
was describedfroma singlespecimenwith young
flowerbuds only. A recentcollection, while also
only in bud, does have fruit and also adds to
inflorescencecharacters.
Additional specimens examined. FRENCH
GUIANA.Sail, MontsLa Fum6e,4 Nov 1982(st),
Mori & Boom 15161 (NY).
BRAZIL. TERR. AMAPA: Agua Fria,23 Oct 1979(fl,
fr),Austinet al. 7180 (MG, NY). PARA: Belem, 2 Mar
1955 (st), T. N. Guedes311 (IAN).

3. Parinari Aublet

Key to the American Species of Parinari

1. Stipules(5-)10-40 mm long, persistent,semiamplexicaul.
2. Leaf base subcordateto cordate,the apex blunt;inflorescencedensely crowded. 3.1. P. alvimii.
2. Leaf base roundedto acute, the apex acuminate,inflorescencelaxer.
3. Young brancheswith short soft pubescence;leaf base roundedto cordate. 1. P. campestris.
3. Young brancheswith dense stiff ferrugineous-brown hairs;leaf base cuneateto subcuneate.
3. P. rodolphii.
1. Stipulessmall, 1-4 mm long, caducous,axillaryand not claspingstem.
4. Leaf undersurfacewhite-lanate,the pubescenceobscuringthe stomatalcavities; subshrubor small
shrub. 16. P. obtusifolia.
4. Leaf undersurfacewith gray-brownpubescencenot entirely obscuringthe stomatal cavities; large
shrubsor trees.
5. Midribdistinctlyimpressedon leaf upper surfacefor entirelength.
6. Leaf apex roundedto apiculateor bluntlyacuminateon some leaves, with acumento 3 mm
long; primaryleaf veins 9-20 pairs.
7. Leaves orbicularto broadlyelliptic, 2.5-6 cm long, 1.5-4.5 cm broad,the primaryveins
9-14 pairs. 10. P. maguirei.
7. Leaves elliptic, 6.5-7.5 cm long, 3-4 cm broad,the primaryveins 15-20 pairs.
11. P. littoralis.
6. Leaf apex with a well-developedacumen 7-15 mm long; primaryleaf veins 21-30 pairs.
8. Leaves 9-17 cm long; petioles usually with two pairs of glands;flowers 6-9 mm long;
primaryleaf veins more than 3 mm apart. 2. P. montana.
8. Leaves 2-7.5 cm long;petioleswithoutdistinctglands;flowers5-6 mm long;primaryleaf
veins 1-2 mm apart. 12. P. parvifolia.
5. Midribimpressedonly on lower portion of uppersurface,or not impressed.
9. Leaf base distinctlycordateor subcordate.
10. Leaves ovate, the acumen 10-12 mm long; petioles 6-12 mm long, glandular;inflores-
cences much-branched,5-6 cm long. 13. P. cardiophylla.
10. Leavesoblong,the acumen1-3 mm long;petioles3-4 mm long,eglandular;inflorescences
little-branched,2-4 cm long. 17. P. romeroi.
9. Leaf base roundedto cuneate.
11. Primaryleaf veins 28-35. 14. P. parilis.
11. Primaryleaf veins 12-26.
12. Inflorescencea dense many-floweredcorymbosepanicle.
13. Inflorescenceand flowers with yellow-brownpubescence;leaf apex acute or
short-acuminate,the acumen 0-4 mm long; young brancheswith short pubes-
cence only. 5. P. occidentalis.
13. Inflorescenceand flowerswith silver-graypubescence;leaf apex with acumen
8-13 mm long;youngbrancheswith both short pubescenceand long stiffhairs.
9. P. klugii.
12. Inflorescencea lax panicle, not corymbose.
14. Leaf ovate, with a well-developedslenderacumen 9-16 mm long. 6. P. sprucei.
14. Leaf elliptic to oblong-lanate,the acumen0-12 mm long.
Systematic Treatment 57

15. Primaryveins 22-26 pairs;leaves 7.5-16 cm long.

16. Upper surfaceof midrib slightlyprominent;petioles terete.
8. P. brasiliensis.
16. Upper surfaceof midrib slightlyimpressedon lower portion;petioles
canaliculate. 15. P. chocoensis.
15. Primaryveins 16-21 pairs;leaves 3-9.5 cm long.
17. Young petioles canaliculate;leaf reticulationprominenton uppersur-
face. 7. P. pachyphylla.
17. Youngpetiolesterete;leaf reticulationinconspicuouson uppersurface.
4. P. excelsa.

Additional Notes and Descriptions Additionalspecimensexamined.BRAZIL:BAHIA:8

of Species of Parinari kmS of Itacare,16Oct 1968(fl),Almeida& T. S. dos
Santos164 (CEPEC, NY);2 km S of Itacare,14 Jun
Thedistributionof Parinarias a wholeis shown from1972(fl),T. S. dosSantos2305(CEPEC, NY);28 km
Urucucaon rd.to SerraGrande,15Jul 1978(fl),
in Figure86. Moriet al. 10243(CEPEC, NY);BomGosto,Ilheus,
Mar1943(st),Fr6es12711/80(A).
3-3.1. Parinari alvimii Prance, Revista Brasil.
Bot. 2: 37, fig. 6. 1979. Type. Brazil. Bahia: Local name. oiti-mirim.
Una, Fazenda Sao 29 Oct 1969 T. Parinarialvimiidiffersfrom all speciesof Par-
Rafael, (fl),
S. dos Santos 457 (holotype,CEPEC;isotypes, inari, except P. rodolphiiand P. campestris,in
its large,amplexicaulstipules.It is quitedifferent
FHO, NY).
fromthesetwo speciesin the inflorescence,which
Treeto 20 m tall, the young branchesdensely is compact and smaller with densely crowded
ferrugineous-tomentose,becomingglabrousand flowers,in the largerflowers,and the leaf shape,
conspicuouslylenticellatewith age. Leaf lamina cordate at base and blunt at apex. Parinari al-
oblong to ovate, coriaceous, 5-12 x 4-10 cm, vimiidiffersfrom P. littoralisin the stipules,the
subcordateto cordate at base, rounded to very larger leaves, the thicker, rufous petioles, the
bluntly acute at apex glabrousabove except on largerbracts and flowers, and the denser inflo-
midrib, tomentose and with stomatal cavities rescences. It differs from two eastern Brazilian
obscured by dense pubescence beneath, fre- species,P. excelsaand P. brasiliensis,in the stip-
quentlywith two glandsat junction with petiole, ules and bracts,the leaf shape, and the compact
midribprominentbeneath,slightlyimpressedand inflorescence.
tomentoseabove;primaryveins 17-25 pairs,ex-
tremely prominent beneath, impressed above; 3-9. Parinari klugii Prance, Fl. Neotrop. Mono-
petioles 3-8 mm long,tomentose,terete.Stipules gr. 9: 190. 1972.
20-35 mm long, semiamplexicaul, persistent,
membranous,exteriorpilose-tomentoseat cen- Fruitnarrowlyoblong,4-6 x 2.2-2.5 cm; exo-
ter of base, gray-lanateover rest of surface,but carpminutelylenticellate;mesocarpthin, fleshy;
with a few pilose appressedhairs,glabrouswith- endocarphard and fibrous,pubescentwithin.
in. Inflorescencesof terminal and subterminal Distribution (Fig. 88). This species was de-
dense-floweredpanicles, 3-5 cm long, the rachis scribed from a single collection from the De-
and brancheslight brown tomentose. Bracts to partmentof San Martin,Peru.Recentcollections
1 cm long, persistent, ferrugineous,tomentose indicate that it is quite common in Loreto De-
on exterior,enclosingflowerbudsin smallgroups. partment.
Receptacle subcampanulate-turbinate,brown- Habitat. It occurs on riverside temporarily
tomentose on exterior,tomentose within. Petals flooded forests and upland terra firme.
five, white, slightlyshorterthan calyx lobes. Sta-
mens 5-7, unilateral,with shorttooth-likestam- Representative additional specimens examined.
inodes opposite them. Ovary and lower two- PERU. AMAZONAS:Huambisha, Rio Santiago, below
thirds of style densely pilose. Fruit unknown. Caterpiza,20 Nov 1979 (fr), Tunqui105 (MO, NY).
LORETO: Prov. Requena,Rio Ucayali, ArboretumJe-
Distribution(Fig. 87). Knownonly fromcoast- naro Herrera,Jul-Sep 1976 (fl), Bernardi16334 (G,
al Bahia. NY); Rio Tacsha, Curaray,19 Sep 1972 (fl), Croat
Habitat. Littoralforest. 20443 (MO, NY); Rio Itaya nr. Palo Seca, 20 Mar
58 Flora Neotropica

1977 (fr), Gentryet al. 18466 (MO, NY). MADREDE (Steyermarket al. 117792). However,thereis no
DIOS:Rio Manfi,PakitzaStation,19 Nov 1980(fr), reason to believe that this representsany diver-
Foster5774(F). sificationsince neithercollectionhas a complete,
Local name. yakuku. intact inflorescence.The flowers of this species
aretypicalfor any Parinari,the only notablefea-
3-10. Parinari maguirei Prance, Fl. Neotrop. ture being the extremelylong pubescenceof the
Monogr. 9: 190. 1972, descr. emend. Type. style.
Guyana.KaieteurSavannas,11 May 1944 (fr),
Maguire & Fanshawe 23390 (holotype, NY; 3-11. Parinari littoralis Prance, Fl. Neotrop.
isotypes, F, M, US). Mongr.9: 191. 1972.
Treeto 15 m tall, the young branchestomen- Distribution(Fig. 88). This distinctspecieswas
tellous,becomingglabrouswith age.Leaf lamina describedfrom a single collection from Marau,
orbicularto elliptic,coriaceous,2.5-6 cm x 1.5- Bahia. It appearsto be confined to a small area
4.5 cm, roundedto subcordateat base, rounded of the coastal restingaforests of Bahia.
to apiculateat apex, glabrousabove, tomentose
Additionalspecimensexamined.BRAZIL.BAHIA:
and with stomatal cavities beneath; midrib Belmonteto ItapebiRd., km 21, 18 May 1979 (fl),
slightly impressed on upper surface,prominent Mattos Silva et al. 386 (CEPEC,NY); Marau, 3 Aug
beneath;primaryveins 9-14 pairs, plane above, 1967(fl),da Vinha12(NY);5 kmS of Marai(fl),Mori
prominentbeneath;petioles2-6 mm long,terete, & Carvalho 12001(CEPEC, NY);Ilh6us,21 kmfrom
tomentose.Stipulesaxillary,lanceolate, 1-3 mm Olivenqa,25 Oct 1972 (fl), Pinheiro1948 (NY).
long, caducous. Inflorescencesof terminal and Local name. oiti.
subterminalpaniclesto 8 cm long, the rachisand The fieldnotes of Mori & Carvalho12001 note
branches gray-brown-tomentellous.Receptacle that "the fruits are covered by billions of black
subcampanulate-turbinate,tomentose on exte- bees (Melipona)which remove the pericarp."
rior,denselypilose within and aroundthroatand
sparsely puberulous at base; pedicels 0.5 mm 3-14. Parinari parilis Macbride, Candollea 5:
long. Petals five, equallingcalyx lobes. Stamens 367. 1934.
6-7, unilateral,with shorttooth-like staminodes This species, known in 1972 only by two col-
opposite them. Ovary inserted at mouth of re-
lections, the holotype and a paratype,has now
ceptacle, pilose; style long-hirsute. Fruit ellip- been re-collectedseveral times (Fig. 91).
soid, 4-5 cm x 3-3.5 cm; exocarp verrucose;
Habitat. It appearsto be confinedto the sea-
mesocarpthin, fleshy;endocarphard, thick, ex-
ternally fibrous and granular, densely lanate sonally flooded tahuampaforest.
within. Representativeadditionalspecimensexamined.
PERU. LORETO:Rio Mazanabove La Libertad,10 Jul
Distribution(Fig. 90). Venezuelaand Guyana 1976(fl),Gentry&Revilla16638(MO,NY);Rio Na-
on the plateau sharedby these two countries. nayoppositeSantaClara,7 Apr1977(fr),Gentry 19094
Habitat. Collectedin savannamarginand gal- (MO,NY);Rio Napo,Quebrada de Zucusari,5 Apr
1979(fr),RimachiY.4378(NY);RioLoreto-Yacu, 10
lery forest. km fromColombianfrontier,6 Feb 1969(fl),Sastre
Additionalspecimensexamined.VENEZUELA. &Echeverry 641(P);RioTacsha,Curaray, 12Sep1972
BOLiVAR:E of Icabaru, 18 Dec 1978 (fl), Steyermark (fl),Croat20367(AAU,MO);Rio Yaguasyaca, trib-
et al. 117792(NY,VEN),18 Dec 1978(fl, fr),Stey- utary of Rio Ampiyacu, 7 Nov 1977 (fr), Gentry&
ermarket al. 117824(NY,VEN). Revilla20371 (MO, NY).
Local names. parinari,parinariblanco.
This species, describedfrom two fruitingcol-
lectionsfrom Guyana,has now been collectedin
3-15. Parinari chocoensis Prance, Fl. Neotrop.
flowerin nearbyVenezuela.The new collections
have characteristicsmall orbicularleaves with a Monogr.9: 194. 1972.
blunt apex which distinguish it from all other Distribution(Fig. 88). This species, described
neotropicalspeciesof Parinari.The inflorescence from a single collection from the Rio Baudo in
is ratherlax in one collection (Steyermarket al. Choc6, Colombia,has now been collectedin the
117824) and compact and clusteredin the other Rio San Juan region of the same department.
SystematicTreatment 59

Additional specimens examined. COLOMBIA. from two collections from the Department of
CHOCO:Hoya del Rio San Juan, Quebrada Cunperro,
below Noanama, 8 Apr 1979 (fr), Forero et al. 4862
Narifto,Colombia, is now also known from the
(COL,MO, NY); basin of Rio San Juan,Rio Taparal, nearbyforests of Esmeraldas,Ecuador.
Nov-Dec 1979 (fr),vanRoodenet al. 586 (COL,MO, Additionalspecimenexamined.ECUADOR.ESME-
NY). RALDAS:Borb6n,8 Sep 1965 (fr),Little&Dixon 21014
(NY,US).
3-17. Parinari romeroi Prance, Fl. Neotrop.
Local name. cuero de sapo.
Monogr. 9: 400. 1972.
Distribution (Fig. 92). This species, described

4. Exellodendron Prance
The distribution of the genus is shown in Figure 93, that of Exellodendronbarbatumand E.
cordatumin Figure94, and E. coriaceum,E. gardneri,and E. gracile in Figure95.

4-5. Exellodendrongracile (Kuhlmann)Prance, Additionalspecimenexamined.BRAZIL.ESPiRITO

Fl. Neotrop. Monogr.9: 200. 1972. SANTO: Reserva Florestal Linhares, Estrada 161, 22
Jan 1973 (fl), Spada 151 (INPA, RB 162365).
This most distinctivespeciesof Exellodendron
was known only by the fruitingtype in Prance Local name. guaiti-mirim.
It is now since it This is noted as a tree 34 m tall of the "mata
(1972). obviously very rare,
occurs only in the much destroyedforest of Es- de taboleiro."
pirito Santo, Brazil.A new collection shows that
E. gracile has flowerstypical of the genus.

5. Maranthes Blume
5-1. Maranthespanamensis(Standley)Prance&
White, Brittonia37: 76. 1985. Fig. 153.
Distribution.This species was placed in syn-
onymy under the Malesian Maranthes corym-
bosa by Prance (1972) and appearedto be re-
stricted to Cerro Jefe in Panama in this
hemisphere. It has now been collected in the
province of Heredia in Costa Rica and in Nic-
aragua,and so, like Licaniaaffinis,occursin these
moist forested areas of Central America. This
confirmsthat M. panamensisis native to Central
America and not introduced,as originallysug-
gestedby Prance(1968). The good materialnow
available confirms that the single neotropical
species of this otherwise African and Malesian
genus is quite distinct.
Additional specimens examined. NICARAGUA.
ZELAYA: Rio Barbereba,5 km from Nueva Guinea,
16?47'N,84?29'W,27 Aug 1982 (fr),Araquistain3149
(MO).
COSTA RICA. HEREDIA:Finca la Selva, Rio Puerto
Viejo, 25 Feb 1982 (fl), Hammel 11260 (NY), 5 Apr
1982 (fr),Hammel & Schatz 11577 (NY).
PANAMA. PANAMA: Cerro Jefe, 8 Aug 1968 (fr),
Correa& Dressier958 (MO, NY), 29 Jul 1967 (fr),
Dwyer & Gauger7324 (COL);Cerro Azure, 11 Mar
1977 (fl), Folsom et al. 1949 (MO, NY). CANALZONE:
PipelineRd., 9 km NW of Gamboa,29 Oct 1973 (fr),
Nee 7675 (MO, NY).

6. Couepia Aublet
Revised Key to Species of Couepia
1. Inflorescencea racemeor spike.
2. Leaves prominentlyreticulatebeneath,with conspicuousparallelprimaryveins; exocarpoften to-
mentellous.
3. Inflorescencewith short silver-graypubescence;receptacleturbinate;bracteolescaducous.
28. C. elata.
3. Inflorescencewith dense ferrugineousor gray-puberulouspubescence;receptaclecylindricalto
obconical;bracteolesusuallypersistent.
60 FloraNeotropica
4. Inflorescencegray puberulous;leaf venation only prominulous;fruit exteriorglabrousand
smooth. 6.1. C. bernardii.
4. Inflorescencedensely ferrugineous;leaf venation prominent;fruit exteriorverrucoseor to-
mentose.
5. Leaf undersurfacewith distinct stomatalcavities; fruit exteriorverrucose. 10. C. foveolata.
5. Leafundersurfacereticulate,but withoutstomatalcavities;fruitexteriorusuallypubescent,
rarelyverrucose.
6. Leaf acumen 5.5-18 mm long; receptaclelong and slender, 11-22 mm long. 7. C. parillo.
6. Leaf acumen 1-12 mm long;receptacleshort and thick, 3-10 mm long.
7. Inflorescencemany-flowered;leaves ovate to oblong-elliptic8.5-18 cm long.
9. C. canomensis.
7. Inflorescencefew-flowered;leaves elliptic, 2-6.5 cm long.
8. Leaves elliptic, 2.7-4 cm broad, lower surfacewith pubescencenot covering
entire surface,the venation conspicuous. 8. C. steyermarkii.
8. Leavesoblongto oblong-lanceolate,1.5-3.2 cm broad,lowersurfacewith lanate
pubescencecoveringentire surfaceand obscuringreticulatevenation.
8.1. C. canescens.
2. Leaves not prominentlyreticulatebeneath;fruit exocarpalwaysglabrous,smooth or verrucose.
9. Bracteolespersistent,and at least 3/4the length of receptacleor more than 10 mm long.
10. Flowersnot more than 4 mm long. 14. C. spicata.
10. Flowers6 mm long or more.
11. Flowers 16-75 mm long; stamensmore than 50; leaves 14-26 cm long.
12. Primaryleaf veins distinctly anastomosingat marginsto form a marginalvein;
leaves with roundedto subcordatebases;receptaclesubcylindricalor tubular,30-
50 mm long.
13. Flowers16-18 mm long;receptaclesubcylindrical,glabrousexceptfor reflexed
hairs at apex; stipulesto 15 mm long, caducous. 32. C. insignis.
13. Flowers60-75 mm long; receptacletubular,tomentosewithin almost to base;
stipules 20-35 mm long, persistent. 32.1. C cidiana.
12. Primaryleaf veins not anastomosing;leaf base cuneateto rounded;receptacletu-
bular, 12-15 mm long.
14. Leaveselliptic, 6-10.5 cm broad;primaryveins 10-12 pairs;receptacle20-25
mm long. 30. C. martinii.
14. Leaves oblong-lanceolate,3.5-7 cm broad;primaryveins 17-25 pairs;recep-
tacle 12-15 mm long. 31. C. bondarii.
11. Flowers 6-12 mm long; stamens 15-28; leaves 6-18 cm long.
15. Leaves 10-18 cm long, caudate-acuminate;exteriorof receptaclewith red-brown
pubescence;stamens 15-19; style pubescentat base only. 12. C. exflexa.
15. Leaves 6-13 cm long, apex acuminate;exteriorof receptaclewith light-brownse-
riceouspubescence;stamensca. 25; style pubescentfor /4 of length. 13. C. habrantha.
9. Bracteolescaducousor less than half the lengthof receptacle,under 10 mm long.
16. Receptaclepubescentinside to the base; filamentshirsute;leaves hirsutebeneath.
33. C. recurva.
16. Receptacleglabrousinside except at throat;filamentsglabrous;leaves with arachnoidin-
dumentumor glabrousbeneath.
17. Stamensless than 40.
18. Receptaclesubcylindrical,densely ferrugineous-sericeous.
19. Leaves 2.5-5 cm long; primaryveins 8-14 pairs, slightly impressed;petioles
2-5 mm long. 13.1. C. scottmorii.
19. Leaves 7-24 cm long; primaryveins 15-20 pairs,plane or slightlyimpressed;
petioles 6-10 mm long.
20. Leavesnarrowlyoblong,7-10 cm long, 2-3.3 cm broad,glabrousbeneath;
receptaclelight-brownon exterior;stamensca. 35. 13.2. C. carautae.
20. Leaves broadlyoblong, 12-24 cm long, 4.5-8.5 cm broad, ferrugineous-
lanate-pubescentbeneath;receptacleferrugineouson exterior;stamensca.
25. 11. C. magnoliifolia.
18. Receptaclecylindrical,with sparseappressedpubescenceonly.
21. Leaves obovate, 2.5-10 cm long, bluntly acuminate;bracteolespersistentor
subpersistent;stamens in complete circle. 34. C. obovata.
21. Leaves oblong-lanceolate,9-15 cm long, with long acumen; bracteoles ca-
ducous;stamensunilateral. 1. C. guianensis.
17. Stamensnumerous(more than 60).
22. Receptacleglabrousor with sparseappressedgraytomentum on exterior.
SystematicTreatment 61
23. Leaves 9-17 cm long, densely appressed-lanatepubescentbeneath;primary
veins 12-15 pairs. 35. C. williamsii.
23. Leaves 5-8 cm long, sparselypubescentbeneath;primaryveins 7-10 pairs.
35.2. C. marleneae.
22. Receptaclewith dense brownpubescenceon exterior.
24. Leaves oblong-lanceolate. 23. C. krukovii.
24. Leaves oblong to oblong-elliptic.
25. Primaryveins 24-30 pairs. 22. C. macrophylla.
25. Primaryveins 16-20 pairs.
26. Leaf apex with prominentwell-developedacumen. 36. C. chrysocalyx.
26. Leaf apex roundedto bluntlyacuminate. 24. C. latifolia.
1. Inflorescencea panicle.
27. Bractsand bracteolespersistentat flowering,at least half as long as receptacle.
28. Flowers20-25 mm long;exteriorof receptacleandcalyxlobeswitha longferrugineoussericeous
pubescence;leaves glabrous(rarelyglabrescent)beneath. 37. C. eriantha.
28. Flowers7-20 mm long;exteriorof receptacleshortlybrown-to gray-tomentose;leaves arach-
noid-pubescentbeneath.
29. Leaves thick and coriaceous;bracteolesalways persistingthrough flowering;rachis of
inflorescenceand receptaclelongitudinallystriate,or if not, petioles 13-18 mm long.
30. Flowers 7-15 mm long; leaf bases subcordate,rarelyrounded;rachisand receptacle
with longitudinalstriations. 15. C. bracteosa.
30. Flowers 18-22 mm long;leaf bases roundedto subcuneate;rachisand receptaclenot
longitudinallystriate. 17. C. belemii.
29. Leavesthin and membranous;bracteolespersistentonly in bud;rachisand receptaclenot
longitudinallystriate;petioles 4-8 mm long.
31. Stamensconnate at base for at least 1 mm; receptacletaperingto base, subturbinate.
38. C. trapezioana.
31. Stamensfree almost to base;receptaclesubcylindrical. 16. C. subcordata.
27. Bractsand bracteolesnot persistentat flowering,or small and inconspicuous.
32. Interiorof receptaclefilled with hairs to base.
33. Primaryveins 10-15; exteriorof receptacleand calyx lobes with sparseappressedpubes-
cence not completelycoveringsurface. 3. C. paraensis.
33. Primaryveins 17-28; exteriorof receptacleand calyx lobes denselypubescent,completely
coveringsurface.
34. Petioles canaliculateabove; flowers 12-17 mm long; exteriorof receptacleand calyx
lobes with short brownpubescence;stamens 38-40. 19. C. excelsa.
34. Petiolesnot canaliculate;flowers8-12 mm long;exteriorof receptacleand calyxlobes
with sparseshort graypubescence;stamens 20-35. 18. C. caryophylloides.
32. Interiorof receptacleglabrousexcept for deflexedhairs at throat.
35. Leaf undersidecompletelyglabrous;exteriorof receptacleusuallyalmost glabrous.
36. Peduncleselongated(30-80 cm long);exteriorof calyx lobes with 2 sessile glands.
37. Exteriorof receptaclepubescent;calyxtube 14-22 mm long,cylindrical-turbinate;
stamensca. 32; petioles 4-8 mm long. 41. C. longipendula.
37. Exteriorof receptacleglabrousor almost so; calyx tube campanulate,4-6 mm
long; stamens 16-21; petioles 10-12 mm long. 41.1. C. dolichopoda.
36. Pedunclesshort (less than 10 cm long);exteriorof calyx lobes eglandular.
38. Inflorescencea much-branchedcorymbosepanicle,rachisand branchesglabrous;
petioles 1.5-2.5 cm long; stamens 17-20; disc very thick with only small central
cavity. 8-4. Acioa edulis.
38. Inflorescencelittleor much-branched,not corymbose;rachisandbranchessparse-
ly puberulous;petioles4-12 mm long;stamens25-120; disc thinwithlargecentral
cavity.
39. Leaves oblong to oblong-lanceolate,prominentlyacuminate;inflorescence
little-branched,almost racemose;petioles 8-12 mm; stamensca. 110.
35.1. C. glabra.
39. Leaves ovate to elliptic, bluntly acuminate;inflorescencemuch-branched;
petioles 4-7 mm; stamens 25-45. 3. C. paraensis.
35. Leaf undersurfacelanate;exteriorof receptaclepubescent.
40. Exteriorof receptacleand calyx lobes sparselyappressed-puberulous, the pubescence
not forminga complete covering.
41. Stipulespersistent,adnateto the base of petiole;rachisof inflorescence3-5 mm
thick;petal marginsglabrous. 39. C. stipularis.
62 Flora Neotropica

41. Stipulescaducous,not adnateto base of petiole;rachisof inflorescence1-2.5 mm

thick;petal marginsciliate.
42. Leaf undersurfacewith distinct, prominulous,parallelsecondaryvenation,
90?to primaryveins. 6.1. C. bernardii.
42. Leaf undersurfacedensely lanate pubescentand smooth, venation obscured
beneathpubescence.
43. Leaves bluntly or shortly acuminateto obtuse;receptaclesubcampan-
ulate, 3-5 mm thick at top; inflorescencea much-branchedpanicle.
44. Leaves ovate to oblong, 2.5-8.5 cm broad. 3. C. paraensis.
44. Leaves oblong-lanceolate,1.5-4 cm broad. 5. C. maguirei.
43. Leaves prominentlyacuminate;receptaclecylindrical,ca. 2 mm thick
at top; inflorescenceoften with only short 2-3 floweredbranches.
45. Inflorescenceserect,much-branched,predominantlyterminalpan-
icles. 1. C. guianensis.
45. Inflorescenceslittle-branchedracemose panicles, axillary or pre-
dominantlyaxillary,but with a small terminalbranch.
46. Inflorescencessingle racemose, reflexed panicles; bracteoles
persistent. 40. C. reflexa.
46. Inflorescenceserect,terminaland in the upper2-6 axils;brac-
teoles caducous. 1. C. guianensis.
40. Exteriorof receptacleand calyx lobes with dense pubescenceforming a complete
covering.
47. Leaf undersurfaceprominentlyreticulate.
48. Leaveswith deep,well-definedstomatalcavities,2.5-6 cm long,bluntlyacute
at apex. 29.1. C. amaralae.
48. Leaveswithout stomatalcavities, acuminateat apex, or with poorlydefined
shallowcavities, then exceeding6 cm in length.
49. Receptaclegraduallytaperingto a long slenderpedicel;exteriorof re-
ceptacle rufous-pubescent;primaryveins not impressedon upper leaf
surface. 42. C. cognata.
49. Receptaclechangingabruptlyto a short thick pedicel;exteriorof recep-
tacle brown-pubescent;primaryveins impressedon upperleaf surface.
29. C. racemosa.
47. Leaf undersurfacewith plane to prominentvenation but not prominentlyretic-
ulate.
50. Receptacleslightlycurved anteriorlyin bud.
51. Leaf with midrib much impressedabove. 42. C. cognata.
51. Leaf with midrib plane above.
52. Leaves 4-7 cm long; primaryveins 7-10. 53. C. pernambucensis.
52. Leaves 9-18.5 cm long; primaryveins 12-23.
53. Exteriorof receptacleand calyxlobes gray-puberulous; primary
leaf veins 18-23, slightlyimpressedabove. 51. C. impressa.
53. Exteriorof receptacleand calyx lobes brown-tomentose;pri-
mary leaf veins 12-14, plane above. 43. C. multiflora.
50. Receptacleerect, not curved in bud.
54. Outersurfaceof petals distinctlypubescent.
55. Leaves 2.5-5.5 cm long; primaryveins 5-9.
56. Leavesoblong-lanceolateto oblong,acumen5-7 mm long,base
cuneate;petioles 5-7 mm long. 55. C. parvifolia.
56. Leavesorbicularto elliptic,acumen0-3 mm long,baserounded
to cordate;petioles 2 mm long. 20. C. uiti.
55. Leaves (4-)6-16 cm long; primaryveins 9-17.
57. Flowers6-12 mm long; receptaclecylindrical,ca. 2 mm thick
at top below calyx lobes.
58. Inflorescenceand flowers densely ferrugineous-sericeous;
flowers8-12 mm long;petioles 5-10 mm long;receptacle
not striateon exterior. 52. C. meridionalis.
58. Inflorescenceand flowerswith shortgraypubescence;flow-
ers 6-8 mm long;petioles 3-7 mm long;exteriorof recep-
tacle longitudinallystriate. 44. C. ulei.
57. Flowers 12-20 mm long;receptaclesubcampanulate,4-6 mm
thick.
SystematicTreatment 63

59. Pedicels 6-15 mm long; exteriorof receptacleand calyx

lobes short gray-sordid-I erulent. 21. C. cataractae.
59. Pedicels not exceeding6 mm in length;exteriorof recep-
tacle and calyx lobes spreadingbrown-tomentose.
60. Leaves 4-8.5 cm long, 2-3.8 cm broad, rufous-pu-
bescentbeneath,grayingwith age. 45. C. comosa.
60. Leaves 7.5-18 cm long, 3.5-9 cm broad,gray-pubes-
cent beneath.
61. Leavescoriaceous,roundedto bluntlyacuminate
at apex. 27. C. grandiflora.
61. Leaveschartaceous,prominentlyacuminate.
46. C. venosa.
54. Petals glabrousexcept for ciliate margins.
62. Receptaclecylindricalor subcylindrical.
63. Inflorescenceof denselyclusteredglomerules;primaryleaf veins
impressedabove. 52.1. C. coarctata.
63. Inflorescencea loosely branchedpanicle; primary leaf veins
plane to prominentabove.
64. Stamensca. 35; leaf base subcordateto rounded.
16. C. subcordata.
64. Stamens 15-28; leaf base roundedto cuneate.
65. Flowersborne on distinctlyarticulatedpedicels with
lowerpartof pedicelpersistentafterfall of some flow-
ers. 6.2. C. monteclarensis.
65. Pedicelsnot conspicuouslyarticulatedand no partre-
mainingafterflowerfall.
66. Leaves 10-16 cm long, roundedat base;primary
veins 14-19 pairs.
67. Receptaclecylindrical,usuallyglabrousatbase
within;stamensca. 24. 38. C. trapezioana.
67. Receptaclesubcylindrical,pubescentwithin
to base; stamensca. 16. 47.1. C. nutans.
66. Leaves 4.5-13 cm long, cuneate to rounded at
base; primaryveins 8-15 pairs.
68. Receptaclecylindrical;stamens 20-28.
6. C. sandwithii.
68. Receptaclesubcylindrical;stamens 11-21.
47. C. polyandra.
62. Receptaclebroadlycampanulateor turbinate.
69. Stamens 14-20.
70. Leaves ovate, 4-9 cm long; primaryveins 11-15; flowers
8-12 mm long; petioles 4-8 mm long. 25. C. ovalifolia.
70. Leavesoblong, 10-18 cm long;primaryveins 16-20; flow-
ers 11-15 mm long; petioles 10-15 mm long. 26. C. schottii.
69. Stamensmore than 28.
71. Receptaclebroadlyturbinate,flattened,3 mm long,almost
solid. 48. C. platycalyx.
71. Receptaclesubcampanulate,not flattened,4-12 mm long,
hollow.
72. Petioles 14-18 mm long;inflorescencedenselycrowd-
ed. 52.2. C. longipetiolata.
72. Petioles 2-10 mm long;inflorescencelax.
73. Leaves 2.5-5.5 cm long, 1.5-3.5 cm broad.
20. C. uiti.
73. Leaves 9-27 cm long, 4-12 cm broad.
74. Leaf marginsundulateand revolute.
49. C. rufa.
74. Leaf marginsplane.
75. Leaf with short and blunt acumen, the
lower surface rufous-pubescent;calyx
lobes rufous-pubescenton exteriorand
64
Additional Notes and Descriptions In Prance (1981) the circumscriptionof this
of Species of Couepia species was changed considerablyfrom that in
Prance(1972). Three species recognizedin 1972
Distribution of Couepiaas a whole is shown were
mergedand aretreatedas subspeciesof this
in Figure96. now ratherwell definedbut variable species.
6-1. CouepiaguianensisAublet,Hist. pl. Guiane
1: 519, t. 207. 1775. Type. French Guiana. Key ta Subspecies of Couepia guianensis
Aublets.n. (lectotype,BM).
AciaamaraWilldenow, Linn,Sp.pi. ed. 4. 3(1):7117.
1800.
AcioaamaraSteudel,NomencLf bot.ed. 1. 9. 1821..
Moquilea Nomeid.
couepiaSteudel, bot. ed. 2.2: 15.
1849.
Tree to 25 m tall, the young branchespuber-
ulous, soon becomingglabrous.Leaf lamina ob-
long to oblong-lanceolate,membranous to co-
riaceous, 4.5-16.5 x 2.5-5.5 cm, rounded to
subcuneateat base, acuminateat apex, the acu-
men 5-18 mm long,glabrousabove,denselygray-
to brown-lanate,or glabrous, or with a sparse
caducous pubescence beneath, frequently with
two glandsatjunctionwith petioae;primaryveiins
10-15 pairs, plane above, prominent beneath;
midrib prominulousabove, prominentbeneath;
petioles 3-9 mm long, canaliculateabove, pu-
bescentwhen young,becomingglabrousand ru-
gose with age. Stipules1-3 mm long, linear,early
caducous. Inflorescencesterminal and axillary
little-branchedpanicles or racemes, the rachis
and branches sparsely puberulousto glabrous.
Bracts and bracteoles minute, membranous,
ovate, caducous. Receptacle cylindricalto sub-
campanulate,4.5-10(12) mm long, 1-2.5 mm
broad below calyx, sparsely puberulousto gla-
brous externally,glabrouswithin except for de-
flexed hairs around throat; pedicels 0.5-4 mm
long. Calyxlobes five, rounded,2-2.5 mm long,
puberulous or glabrous externally. Petals five,
white, ciliate. Stamens 14-30, unilateral,insert-
ed aroundhalf of a circle with short staminodes
opposite them. Ovary villous. Style pubescent
for at least halfits length.Fruitroundedto ovoid,
3-4 cm long, 2.5-3 cm broad;exocarp smooth,
glabrous;mesocarp thin, fleshy; endocarpthin,
fragile,granularin texture,glabrouswithin.
FloraNeotropica
yellow-brown-pubescent within.
50. C. robusta.
75. Leaf with finely pointed acumen, 8-14
mm long, the lower surfacegray-pubes-
cent;calyxlobes gray-pubescenton both
surfaces. 54. C.froesii.
1. Infloresnces of racemes(rarelywith a few short
branctsBbearing 2 flowers),usuallyin at least 3
axils bulw the apex of branch;leaf lamina 9-
chartaceous;petioles6-9 mm long,
16.3 crmlksng,
receptadleatways narrowlycylindrical.Plants of
terrafirnrm. a. subsp. guianensis.
1. Infloresemmesof panicles, usually only 1 or 2
axils beliw apex; leaf lamina 5-10.5 cm long,
coriaceoussachartaceous;petioles3-6 mm long;
or subcampanulate.Plants
receptaclec%lindrical
of terrafirme or flooded river banks.
2. Leaves chartaceous,the underside glabrous
or sparselypubescent,plantsof floodedriver
banks. b. subsp. glandulosm.
2. Leaves thickLycoriaceous, the undersidte
plant ofterrafirme:.
denselylanate-lgpuescent,
c.. subsp. divaricatm.
6-la. Couepia g_i ensis AutIlf subsp. _pu-
nensis. Fig.. isM.
CouepialeptostachyiBenthamex Hooker f., FR.lmas.
14(2): 44.. 1867. Type. Brazil. Amazonas:Maumus
(fl),Spruce15(hiolotype, K; isotypes,BM, Q,GGE,
GOET, LE, M1NY, OXF, P))
CouepiaIvtolorBenoist, Bull Mus. Hist. NatL(aris)
29: 596. 1923. Type. French Guiana (fl), M&linon
s.n. (holotype,P).
CouepiasurinamensisKleinhoonte,RecueilTrav.Bot.
Needr 22: 390. 1925. Type. Surinam.Sectie O (fl),
B. W. 3080 (lectotype,U; isolectotypes,K, NY).
Leaf lamina chartaceous,9-16.3 cm long, the
lower surfacedensely lanate-pubescent;petioles
6-9 mm long. Inflorescencemainly of terminal
axillary racemes, usually in several axils below
the apex, a few with short branches bearing two
or three flowers. Receptacle 7-12 mm long, al-
ways narrowlycylindrical.Plants of terrafirme.
6-lb. Couepia guianensis Aublet subsp. glan-
dulosa (Miquel) Prance, Brittonia 33: 350.
1981. Fig. 105.
SystematicTreatment 65

CouepiaglandulosaMiquel,Stirp.surinam.select. 28. neate at base, apiculateat apex, the acumen 2-

1851. Type.Suriname(fl),Hostmann&Kappler859 8 mm long, glabrous above, with a short ap-
(holotype,U; isotypes, BM, GH, GOET,K, LE, M,
P, S). pressed-lanate caducous pubescence beneath,
Moquileaglandulosa(Miquel)Walpers,Ann. 2: 463. midribprominenton both surfaces,primaryveins
1852. 12-14 pairs, plane above, prominent beneath,
Couepia myrtifoliaBentham ex Hooker f., Fl. bras. secondaryvenation prominulous, more or less
14(2): 44. 1867. Type. Venezuela.Amazonas:San paralleland 90? to primaryveins; petioles 4-7
Carlos(fl), Spruce3681 (holotype,K; isotypes,BM,
BR, F, GH, GOET, LD, LE, NY, P).
mm long, tomentellous,becomingglabrouswith
age, rugulose, canaliculate above, eglandular.
Leaf lamina chartaceous to thinly coriaceous, Stipuleslinear-lanceolate,tomentellous,to 6 mm
5-10.5 cm long, the lower surface glabrous or long, caducous. Inflorescencesof terminal and
sparsely lanate-pubescent; petioles 3-6 mm long. axillary little-branched panicles with central
Inflorescences of terminal and axillary panicles rachis and short 2-3 flowered branches, or of
with small few-flowered branches, in only one or racemes, the rachis and branches puberulous.
two axils below the apex of branch. Receptacle Receptacle cylindrical, 7-8 mm long, sparsely
5-10 mm, subcampanulate to narrowly cylin- gray-puberulousexternally,glabrouswithin ex-
drical. Plants of flooded river banks. cept for deflexed hairs at throat;pedicels ca. 1
mm long. Calyx lobes five, rounded, 2.5 mm
6-1c. Couepia guianensis Aublet subsp. divari- long, puberulousexternally. Petals five, white,
cata (Huber) Prance, Brittonia 33: 351. the margins ciliate. Stamens 20-23, unilateral,
1981. Fig. 104. the arc of the circle opposite to them toothed.
tomentose. Style pubescenton lower por-
CouepiadivaricataHuber, Bol. Mus. ParaenseHist. Ovary
Nat. 6: 75. 1910. Type. Brazil.Para:Belem, 14 May tion only. Fruitovoid, ca. 5 cm long, 4 cm broad;
1901 (fl), HuberMG 2030 (holotype,MG; isotype, exocarp hard and smooth or crustaceouswhen
BM). dry, glabrous;mesocarpwoody, 6-10 mm thick
CouepiadivaricataHubervar.strictiusculaHuber,Bol. when
Mus. ParaenseHist. Nat. 6: 76. 1910. Type. Brazil. dry; endocarp thin and bony, glabrous
Para:Peixe-Boi, Sep 1908 (fl), R. S. RodriguesMG within.
8274 (lectotype,MG;isolectotypes,BM,P, RB 15104, Distribution(Fig. 97). WesternAmazonia.
U).
ParinarikrukoviiGleason, Bull. TorreyBot. Club 55: specimens
Representative examined.PERU.LORETO:
353. 1933. Type. Brazil. Terr. Rondania:Rio Ma- Prov.Requena,Rio Ucayali,Arboretum JenaroHer-
chado, Tabajara,Nov-Dec 1931 (fl), Krukoff1362 7-84(G,NY),27 Aug
rera,Jul-Sep1974(fr),Bernardi
(holotype,NY; isotypes, K, MICH). 1976 (fl bud), Bernardi16219 (G, NY).
BRAZIL.AMAZONAS: Mun.Humaita, Livramento,
Leaf lamina coriaceous, 6-10.5 cm long, the (NY,US);TresCasas,
Oct-Nov1934(fr),Krukoff6635
lower surface densely lanate-pubescent; petioles Sep-Oct 1934 (fr),Krukoff6381(NY, US).
3-6 mm long. Inflorescences of terminal and ax-
Local names. Brazil:uchirana.Peru:parinari
illary panicles with small few-flowered branches blanco.
in only one or two axils below the apex of branch.
The closely relatedspecies Couepiabernardii,
Receptacle 5-7 mm long, cylindrical. Plants of C. obovata,C. reflexa,and C. sandwithiiform a
terra firme.
superspecies(see Prance, 1972, for other similar
Specimen citations for these three subspecies
were given in Prance (1981). groupsin Couepia).The greateramount of ma-
terial available now enables me to group these
6-6.1. Couepia bernardii Prance, Brittonia 33: species togethermore clearly than was possible
in 1972. They are characterizedby the almost
347. 1981. Type. Peru. Loreto: Prov. Requena,
racemose inflorescence with small groups of
Rio Ucayali, Arboretum Jenaro Herrera,
flowerson short branches,thus technicallypan-
4?55'S, 73?45'W, Tree Number 6-6, Jul-Sep
1974 (fl), Bernardi 6-6 (holotype, NY; isotype, icles, the long tubularreceptaclethat is curved
in bud at the apex, and the reticulateleaf un-
G). dersurfacewith parallel secondary venation in
Tree, the young branches puberulous, soon be- all species except C. reflexa. I am maintaining
coming glabrous. Leaf lamina elliptic, charta- all four species at present since they do appear
ceous, 9-16 x 3.5-6.5 cm, rounded to subcu- to be distinct. Since C. reflexais only known by
66 FloraNeotropica
Table II
Differences between species of the Couepia obovata species group

C. bernardii C. obovata C. reflexa C. sandwithii

Leaf laminae(cm) 9-16 4-10 11-14 4.5-9
Apex acuminate apiculateor acumi- acuminate acuminate,roundedor
nate apiculate
Acumen length 2-8 2-5 8-10 5-6
(mm)
Flower sparselypu- glabrousor sparsely sparselypu- denselytomentellous
exterior berulous puberulous berulous
Leaves coriaceous chartaceousto coria- chartaceous coriaceous
ceous

a singletype collectionit is still hardto evaluate.
The other species appear to be separate.They
have distinct geographical distributions, al-
thoughC. obovataand C. bernardiiaresympatric
in Amazonian Peru. C. obovata is much more
widespread,in CentralAmazonia and the South
of the Guianas, however. C. sandwithiiis allo-
patric, found in the Orinoco Delta region and
Guyana. These species can be differentiatedby
the charactersgiven in Table II and in the key
which follows. In the original descriptionof C.
bernardii(Prance, 1981) I referredtwo collec-
tions from Venezuelato that species. These two
collections (Steyermark87610, 88108) actually
belong to C. sandwithii.
The species Couepia monteclarensisis also
closely related to this group of species, but is
more distinct. For differences see under that
species.
Key to Species of Couepia obovata
Species Group
1. Flowerexteriordenselytomentellous,forminga
completecovering;leaf undersideusuallyferru-
gineous-brown-tomentose C. sandwithii.
1. Flowerexteriorglabrousor sparselypuberulous,
not forminga completecovering;leaf underside
gray-puberulous.
2. Leaves membranous,venation smooth and
not reticulatebeneath,the apexwithlongacu-
men 8-10 mm long C. reflexa.
2. Leaves chartaceousto thinly coriaceous,re-
ticulate beneath with parallelsecondaryve-
nation, the apex bluntlyacute to acuminate.
3. Leaf lamina 4-10 cm long, oblong to el-
liptic, the apex acuminate C. obovata.
3. Leaf lamina 9-16 cm long, obovate, the
apex usuallyapiculate C. bernardii.
6-6.2. Couepia monteclarensisPrance, sp. nov.
Type. Brazil.Minas Gerais:EstagaoBiologica
Caratinga,19 Feb 1984 (fl), M. A. Lopes 113
(holotype,BHCB;isotype, NY). Fig. 14.
Species C. sandwithii et C. bernardiiaffinis,
inflorescentiislaxioribus,floribus,cum pedicellis
articulatismunitis (1-5 mm longis), petiolis 8-
10 mm longis diversa.
Tree,the young branchesglabrous,conspicu-
ously lenticellate. Leaf lamina oblong, charta-
ceous, 7.5-11 x 3.3-4.6 cm, subcuneateat base,
apiculate at apex, the apex 4-6 mm long, gla-
brous above, with a short-brown-lanatepubes-
cence beneath; midrib prominulous above,
prominent beneath; primary veins 8-10 pairs,
plane above, prominent beneath;petioles 8-10
mm long, glabrescent,canaliculate,with a single
gland on one side or eglandular.Stipules ca-
ducous (not seen). Inflorescencesof terminal
panicles, 6-8 cm long, with a centralrachis and
shortdichotomousbranchesbearing2 to 4 flow-
ers,the rachisandbranchesgray-puberulous.Re-
ceptacle cylindrical, 5-7 mm long, gray puber-
ulous externally, glabrous within except for
deflexed hairs at throat;pedicels 3-5 mm long,
articulatedat middle and with lower part per-
sistent after flowerdehiscence.Calyx lobes five,
rounded to slightly acute, 2-3 mm long, gray
puberulouson exterior. Petals five, white, gla-
brousexcept for ciliate margins.Stamens 18-20,
inserted almost around complete circle. Ovary
insertednearmouth of tube, lanate.Stylehirsute
for three-fourthsof length. Young fruit ovoid;
exocarp smooth, glabrous;mesocarp granular;
endocarpbony, glabrouswithin.
Distribution. Atlantic coast forest of Minas
Gerais.
Systematic Treatment 67

\I I 3cm. '1'1~~~~~~~~~~~~~~

PA .-t
]-~~~~~~~~~~~~~~~~~~.
E.
I

nwated;eclarem"' I

t GNllp^ 15 a mm!l t

FIG. 14. Couepia monteclarensis (Lopes 113). A, habit; B, flower; C, flower section; D, petal; E, floral
diagram; F, young fruit; G, fruit cross-section.
68 FloraNeotropica
Additionalspecimenstudied.BRAZIL.MINASGERAIS: the rachisdensely brown tomentose. Bractsand
22 May1984(y fr),P.
EstacaoBiol6gicade Caratinga, bracteoleselliptic, ca. 1 mm long, tomentose on
M. Andrade252 (BHCB,NY).
exterior, caducous. Receptacle campanulate,
This species is named for the MonteclaroRe- straight, not curved, 3-4 mm long, brown-to-
serve where it occurs in a vestige of Atlantic mentose on exterior,glabrouswithin except for
coastal forest that harborsthis and many other deflexedhairsat throat;pedicels0.5-1 mm long.
severely endangeredspecies. Calyx lobes five, acute, tomentose on exterior.
This species is most closely relatedto Couepia Petals five, white, glabrous.Stamens 12-16, uni-
sandwithii from the Guianas and C. bernardii lateralwitha fewshortstaminodesoppositethem.
from western Amazonia. It differs primarilyin Ovary villous. Style glabrousexcept at extreme
the largerflowersand in the inflorescencewhich base. Fruit globose; exocarp glabrous and ver-
is much more open and with the flowersborne rucose when mature; pericarp hard and thick,
in pairson long articulatedpedicels.In addition, undifferentiated,densely lanate within.
the petiolesof C. monteclarensisaremuch longer Distribution(Fig. 98). GuayanaHighland re-
and the secondaryveins wider apart. This is a gion of Venezuela.
most distinct and easily recognizedspecies. The
two collections come from a small remnant of Additionalspecimensexamined.VENEZUELA.
AMAZONAS:CerroDuida,SerraParima,54 kmNWof
the BrazilianAtlantic coastal forest on the Fa- Rio Orinoco,2?27'N, 63?56'W,18-23 May 1972 (fr),
zenda Montes Claros in eastern Minas Gerais. Steyermark106117 (NY), (fl), Steyermark105936
The fact that new species are still being collected (NY).BOLIVAR:Km 119S of El Dorado,12 Jan 1964
in what is left of those forests emphasizes the (fr),Steyermark 93035(NY,VEN);SE
&Dunsterville
21 May 1953 (fr),Steyer-
bluffsof Chimanta-tepui,
importanceof their conservation.It is fortunate mark75526(NY);Abucapa-tepui, NW of Chimanta,
that World Wildlife Fund and the Fundaqao 18 Apr 1955 (st), Steyermark75127 (NY).
Brasileiraparaa Conservaqaoda Naturezahave
a projectat Montes Claros.It is to be hoped that Couepiacanescensis closest to C. cognata,but
this importanttract of land can be permanently differsin the shorterracemoseinflorescence,the
shorter thicker pedicels, the calyx tube that is
preserved.
campanulateand erect, not curved, the shorter
6-8.1. Couepiacanescens(Gleason)Prance,Acta and caducousbracteoles,and the shorterpubes-
Bot. Venez. 9: 119-120. 1974. Type. Vene- cence of the inflorescenceand flowers.In Coue-
zuela. Amazonas: Cerro Duida (fl), Tate 870 pia cognata the inflorescenceis nearly always a
(holotype, NY; isotype, K). slightly branched panicle; only rarely is it re-
duced to a racemeand then it is longerthan that
Gleason,Bull.TorreyBot.Club59: of C. canescens.These two species are ecologi-
Parinaricanescens
370. 1931.
Couepiacognata(Steudel)Fritschvar.cognataproparte cally
distinct. Couepiacognatais a small tree of
sensuPrance,Fl. Neotrop.Monogr.9: 149. 1972. the forest on highermountain slopes.
Treeto 10 m tall, the young branchestomen-
6-13.1. CouepiascottmoriiPrance,sp. nov. Type.
tellous. Leaf lamina oblong to oblong-lanceo-
Panama. Prov. Panama:CerroJefe, 1000 m,
late, coriaceous,2-6.5 x 1.5-3.2 cm, cuneateto 14 Jul 1975 (fl), Mori 7116 (holotype, NY;
roundedat base, acuminateat apex with acumen
2-5 mm long, glabrousabove except on midrib, isotype, MO). Fig. 15.
denselybrown-lanatepubescentbeneath,the pu- Species C. magnoliifoliaeaffinis,sed foliis mi-
bescence obscuring rather prominent venation noribus, 2.5-5 cm longis, nervis primariis8-14
with parallel primary veins; midrib impressed jugis, petiolis 2-5 mm diversa; a C. habrantha
and tomentellous towards base above, promi- foliis minoribus, bracteolis minoribus caducis,
nentand lanatepubescentbeneath;primaryveins nervis primariisimpressisdiffert.
10-12 pairs, plane or slightly impressed above, Treeto 15 m tall, the young branchessparsely
prominent beneath; petioles 2-3 mm long, puberulous.Leaf lamina elliptic, thickly coria-
denselytomentose,terete.Stipulesca. 1 mm long, ceous, 2.5-5 x 1.6-3 cm, rounded to slightly
linear, tomentose, caducous. Inflorescencester- subcordateat base, apiculateto bluntlyacumin-
minal and subterminalracemes 1.5-3 cm long, ate at apex, the acumen 0-5 mm long, glabrous
SystematicTreatment 69

Fi
71 ?':~~~~~~~~~~~~~~~ m

scodmor
FI.1.Cupasctmri(oi716.A ai;B rut ,la nesufc;D lwr E lwrscin

F, petals.~~~~~~~~~~~~~~~~~~~~~~~~~~'~3

FIG. 15. Couepiascottmorii(Mori7116). A, habit;B, fruit;C, leafundersurface;D, flower;E, flowersection;

F, petals.
70 FloraNeotropica

above, densely brown-arachnoidpubescentbe- midrib prominulousabove, prominent and al-

neath; midrib prominulous above, prominent most glabrousbeneath,with only a few shortstiff
beneath;primaryveins 8-14 pairs, slightly im- hairs; primaryveins 16-20 pairs, plane above,
pressedabove, prominulousbeneath;petioles 2- prominentbeneath;petioles 6-10 mm long, gla-
5 mm long, canaliculate,rugulose, puberulous. brous, rugulose,terete. Stipules linear, to 1 mm
Stipules minute, lanceolate, caducous. Inflores- long, subpersistent.Inflorescencesterminal and
cences terminal and subterminalracemes 3.5- subterminalracemes,the rachisdensely yellow-
5.5 cm long, the rachis densely yellow-brown- brown-sericeous-tomentose. Bracteolesovate, 6-
tomentellous. Bracts and bracteoles caducous. 7 mm long, tomentose on exterior, caducous.
Receptaclesubcylindrical,4-6 mm long, densely Receptaclesubcylindrical,7-8 mm long, yellow-
tomentellouson exterior,glabrouswithin except brown-sericeouson exterior, glabrous within;
fordeflexedhairsat throat.Calyxlobesfive,acute. pedicels 0.5 mm long. Calyx lobes five, acute.
Petals five, sparselypubescenton exterior,with Petals five, glabrouson exterior,the marginscil-
ciliate margins.Stamensca. 26, insertedaround iate. Stamens ca. 35, inserted around complete
complete circle,glabrousexcept for mass of hair circle, filaments exserted. Ovary pilose. Style
aroundinteriorof unitedbasesof filaments.Style densely hirsuteto apex. Fruit not seen.
densely pubescentfor three-fourthsof its length. Habitat. Rain forest on clay soil.
Ovarydensely lanate. Fruit ellipsoid, 5 x 3 cm; Local name. milho torrado.
exocarpsmooth, glabrous;mesocarpthin, fleshy; Uses. The wood is used for railroadties.
pericarpca. 3 mm thick. This species belongs to the Couepia magno-
Distributionand habitat(Fig. 115). Cloud for- liifolia superspeciesand is the firstof that group
est dominatedby Clusiaand Colpothrinaxcookii to be found in eastern Brazil. The four related
at 1000 m, on CerroJefe, Panama. species occur in Amazonia and the Guianas. It
differsfrom C. spicata and C. habranthain the
Additionalspecimensexamined.PANAMA.PANAMA:
CerroJefe,1000m, Sep 1972(fr),Gentry6172(MO, smaller and caducous bracteoles, the narrow
NY);CerroJefetrailon ridgeNE fromsummit,1000 leaves and the greaternumberof stamens.It dif-
m, 18 Dec 1974(fr),Mori&Nee3741(MO,NY). fers from C. magnoliifoliaand C. reflexain the
This species also belongsto the Couepiamag- smaller, narrowlyoblong leaves, the glabrous,
not ferrugineous-pubescent, lower surfaceof the
noliifolia superspecies and extends that group and in the number of stamens. It
from the Guianasto Panama.It is relatedto, but leaves, greater
distinct habrantha and is most closely relatedto the CentralAmazonian
quite from, C. C. mag-
C. habrantha. Couepia carautae is completely
noliifolia, differingin the much smaller leaves differentfrom the
with fewerprimaryveins and shorterpetioles. It only two other species of east-
differsfrom C. carautaein the small and elliptic ern Brazil with racemose inflorescences,C. in-
leaves with a densely pubescentundersurface. signis and C. bondarii,both of which have much
largerleavesand flowers.Couepiacarautaeis one
6-13.2. CouepiacarautaePrance,sp. nov. Type. of the few species of Couepia without a lanate
on
Brazil.EspiritoSanto:Linhares,Reserva Flo- pubescence the lower leaf surfacesand in this
restal de CVRD, Estrada 143A6, km 1.430, character it differsfrom all other easternBrazil-
Talhao 601, 24 Jan 1978 (fl), J. Spada 31/78 ian species.
It is with that I name this species for
(holotype, INPA; isotype, NY). Figs. 16, 99. Dr. Pedro pleasure
Carautaof Rio de Janeiro, who has
SpeciesC. magnoliifoliaeaffinis,sed foliis sub- done much to provide interestingmaterial for
tus glabrisminoribusoblongo-lanceolatis,stami- my studies of Chrysobalanaceae.
nibus 35, receptaculoextus brunneo-tomentoso
differt;a C. habranthaefoliis subtusglabris,brac- 6-14.
teolis minoribuscaducis, staminibus 35 differt. CouepiaspicataDucke,Arq.Inst.Biol.Veg.
2: 36. 1935.
Tree 22 m tall, the young branchesglabrous.
Leaf lamina narrowlyoblong, coriaceous, 7-10 Distribution(Fig. 115). This species,knownin
x 2-3.3 cm, subcuneateat base, acuminate at 1972 only from the type found near Manaus,
apex, the acumen 4-7 mm long, finely pointed, appearsto be ratherwidely distributed.
glabrous above, glabrous and waxy beneath; Habitat. It is a species of foreston terrafirme.
Systematic Treatment 71

3cm.

. A

'" 5% mm.
GouAO,P
caraute

FIG. 16. CouepiacarautaePrance(Spada31/78). A, habit;B, leaf undersurface;C, flower;D, flowersection;

E, petal.

Additional specimens examined. BRAZIL. (INPA 60898), 10 Aug 1976 (fl),Mota &Monteiros.n.
AMAZONAS:Manaus-Itacoatiara Rd., km 135, 11 Jul (INPA 61341).
1975(fl),Monteiros.n. (INPA 50017);Novo Aripauna,
Nova Prainha,23 Jul 1976 (fr),Mota & Monteiros.n. 6-17. Couepia belemii Prance, Fl. Neotrop.
(INPA 60724), 31 Jul 1976 (fl), Mota & Monteiros.n. Monogr. 9: 228. 1972.
72 FloraNeotropica

Distribution(Fig. 97). This species, described 6-29.1. CouepiaamaralaePrance,sp. nov. Type.

from two collectionsfrom coastalBahia,appears Brazil.Amazonas:Base of SerraAraca, 3 km
to be quitecommon in the remnantsof the south- E of Rio Jauari,0?49'N,63?19'W,27 Feb 1984
ern Bahia wet forest. (fl), Pranceet al. 29261 (holotype,INPA; iso-
type, NY). Figs. 17, 97.
Additional specimens examined. BRAZIL. BAHIA:
Marau,18Jan1967(fl),Belem3169(IAN,NY, UB); Species C. racemosaeaffinis,foliis minoribus
20 km NW of Una, 27 Feb 1978 (fl), Mori et al. 9336 2.5-6 cm
longis, subtuscavis stomatalisprofun-
(CEPEC, NY). ESPIRITO SANTO:Reserva Florestal 4-5 mm longis differt.
CVRD,Linhares, 8 Nov 1977 (fl), Spada 007/77 dis, petiolis
(INPA). Shrub 1-4 m tall, the young branches ap-
pressedlanatepubescent,becomingglabrouswith
6-18b. Couepia caryophylloidesBenoist subsp. age. Leaf lamina oblong to oblong-elliptic,co-
glabraPrance,subsp. nov. Type. Brazil.Para: riaceous, 2.5-6 x 1.5-3.2 cm, roundedat base,
Santarem-Cuiabfa Hwy., km 82, 30 Mar 1984 acute or bluntly acuminate at apex, glabrous
(fl),Medeiros& Marinho58 (holotype,IAN). above, prominentlyreticulatebeneathwith con-
Fig. 99. spicuousdeep stomatalcavities filled with short
graypubescence;midribprominentand sparsely
A subsp. caryophylloidereceptaculointus gla- lanate
beneath, slightly impressed above; pri-
bro differt.
maryveins 9-10 pairs,prominentand lanatebe-
This materialis identicalto subsp. caryophyl-
neath,planeto slightlyimpressedabove;petioles
loides except for the glabrousinterior of the re- 4-5 mm
long,lanatewhenyoung,rugulose,terete,
ceptacle,a consistentcharacterin Couepia;I have eglandular.Stipules caducous. Inflorescencesof
thus recognized the new taxon, which is geo- little branched
panicles or racemes,axillaryand
graphicallydistant from subsp. caryophylloides, terminal,the rachisand branchesbrown-tomen-
at the subspecificlevel. tose. Bractsand bracteolesovate, caducous, to-
mentellous on both surfaces. Receptacle cam-
6-22. CouepiamacrophyllaSpruceex Hookerf.,
panulate-turbinate,3-4 mm long, tomentellous
Fl. bras. 14(2): 43. 1867. on exterior,glabrouswithin except for deflexed
Distribution(Fig. 108).The rangeof this species hairs at throat. Calyx lobes five, acute, 2 mm
has been considerablyextended by recent col- long, tomentellouson both surfaces.Petals five,
lections. white, glabrous.Stamens35-40, insertedaround
half of circle. Ovary densely villous. Style gla-
Additionalspecimensexamined.PANAMA.DARIEN: brous
RioUcurganti, 7 Jul1967(fl),Bristan1121(MO,NY). exceptfor deflexedhairsat base. Fruitglo-
BRAZIL.ACRE:Rio Caete,tributaryof Rio laco, 25 bose, 5 cm diam.; exocarpglabrous,lenticellate;
Sep 1978 (fl), J. Ramos et al. 651 (INPA). pericarp2-2.5 mm thick, hard and woody, to-
mentose within.
6-23. Couepia krukovii Standley, Publ. Field
Additional specimens examined. BRAZIL.
Mus. Nat. Hist., Bot. Ser. 17: 250. 1937. AMAZONAS:Margin of Rio Jauari,Araca,1 Mar1977
Distributionand habitat (Fig. 107). In 1972 (fr), Rosa & Cordeiro 1713(MG,NY);Rio Araca,4
Nov 1952(fr),Fr6es29277;3 km S of SerraAraca,
this species was still known only from the two 0?49'N,63?19'W et al. 10509(INPA,
(fl),Rodrigues
type collections, the type and a paratype,from NY),Rodrigues et al. 10528(INPA,NY).
the Rio Madeiraregion.It appearsto be confined
to varzeaforestsof the centralpart of the basins This distinctive new species belongs to the
of the Rios Madeiraand Purus, extending into Couepia species group with deeply reticulate
Bolivia. leaves. The lower leaf surfacehas deep stomatal
cavities more similarto those that occur in Par-
Additional specimens examined. BRAZIL. inari and a few species of Licania than to those
AMAZONAS: Rio PurusbetwenRedencaoandItaboca,
of other species of Couepia. It differs from C.
22 Nov 1971 (fl), Prance et al. 16315 (FHO, INPA,
MG,NY, US). foveolata,which also has stomatalcavities, in the
BOLIVIA.BENI: Rio Beni, CachuelaEsperanza,28 branched inflorescenceand in the short blunt
Feb 1924(fl),Meyer46 (Z). leaves, and from C. steyermarkiiand C. canes-
SystematicTreatment 73

W,.
74 Flora Neotropica

cens in the branchedinflorescence,the stomatal triangular,acute, tomentose on exterior,tomen-

cavities and the leaf shape. It differs from C. tose within except for glabrous lower portion,
racemosain the much smaller,blunt leaves, the 10-13 mm long. petals five, white, glabrouson
deep well-definedstomatal cavities, the shorter exterior, the margins ciliate. Stamens ca. 165,
petioles and the lanate pubescence of the leaf insertedarounda complete circle, fused at base
venation below. to form a ringca. 5 mm tall, the stamensinserted
It is with pleasurethat this speciesis dedicated in several rows on exterior of ring, the interior
to Ieda L. do Amaral,the Brazilianbotanistwho of ring densely tomentose. Ovary inserted near
was leaderof the Aracaexpedition on which the mouth of tube, densely villous. Style sparsely
type was collected. hirsutepubescenton lower third. Fruitnot seen.
Distributionand habitat (Fig. 99). Forest on
6-32. Couepiainsignis Fritsch, Ann. K. K. Na- terrafirme of westernPara, Brazil.
turhist.Hofmus. 5: 11. 1890. Fig. 106. Additionalspecimenexamined.BRAZIL.PARA: Rio
This distinctive species was described from Cachorro, 16kmNWof Cachoeira
Serrado Cachorro,
scantmaterial,all from Bahia,Brazilbut without Porteira,22 Jun 1980 (fl),Martinelli7052 (INPA,NY).
preciselocality,in Prance(1972). The nineteenth This species is closest to Couepiainsignis, C.
century material is now complemented by two martinii,and C. bondarii.It differsfrom all three
recentcollectionsindicatingthat this species stillin the tomentose interior of the receptacle,an
survives in the coastal forest. uncommonfeaturein the genus,occurringin only
Additional specimens examined. BRAZIL. BAHIA:
one other species with a racemoseinflorescence,
Mun.de Una, 27 km S of Olivenqa,2 Dec 1981(fl), the Andean C. recurva,which is otherwisevery
Carvalho &Lewis858 (CEPEC, NY);Restingade Oli- different. Couepia cidiana also differs from its
venca,1 Dec 1979(fl),MelloFilho3061(CEPEC). threeclosestrelativesin the much longerflowers,
the persistentand longer stipules and the large
6-32.1. Couepiacidiana Prance,Acta Amaz6n- number (26-35) of primaryleaf veins.
ica 13: 21. 1983. Type. Brazil. Para:Mun. de
Oriximina,Rio Paru do Oeste, left marginof 6-33. Couepia recurva Spruce ex Prance, Fl.
CachoeiraChuvisco, 7 Sep 1980 (fl), C. A. Cid Neotrop. Monogr.9: 242. 1972.
et al. 2261 (holotype, INPA 96602; isotypes,
Distribution(Fig. 114). This species was de-
FHO, NY).
scribedfrom a single Sprucecollection from Ec-
Treelet 8-10 m tall, the young branches to- uadorwithoutlocality.A second collectionfrom
mentose. Leaf lamina oblong-lanceolate,coria- the EcuadoreanAndes has now been examined.
ceous, 15-26 x 7-9.5 cm broad, subcordateat
Additional specimen examined. ECUADOR.
base, acuminateat apex, the acumen 7-10 mm TUNGURAHUA: La
short-lanate Victoria,valleyof RioPastaza,1300
long, glabrousabove, densely gray m, 1 Dec 1939,Asplund10055(S).
beneath with contrastingbrown-tomentoseve-
nation;midribprominulousabove, tomentellous 6-35.1.
when young; primaryveins 26-35 pairs, lightly CouepiaglabraPrance,Acta Amazonica
2(1): 10. 1973. Type. Brazil. Amazonas: Rio
impressedabove, prominentand tomentose be-
Cuieiras,just below mouth of Rio Branquin-
neath, anastomosingca. 3 mm from margin to
form a conspicuous marginal vein; secondary ho, Sep 1971 (fl), Prance, D. Coelho & Mon-
teiro 14942 (holotype, NY; isotypes, FHO,
veins more or less parallel,prominulous;petioles
5-7 mm long, terete,eglandular,denselytomen- INPA).
tose. Stipuleslinear,membranous,2-2.5 cm long, Tree 10 m tall, the young branchesglabrous.
persistent.Inflorescencesfew-floweredracemes, Leaf lamina oblong to oblong-lanceolate,coria-
the rachistomentose. Bractsand bracteolesper- ceous, 11-21 x 4-7 cm, base subcuneate,apex
sistent, 2-3.6 cm long, tomentose on exterior, acuminate, the acumen 5-9 mm long, usually
glabrouswithin, ovate, acuminate.Flowers 60- curved,glabrouson both surfaces;midribprom-
75 mm long. Receptaclecylindrical,3-4 cm long, inenton both surfaces;primaryveins 16-17 pairs,
densely velutinous-tomentellouson exterior,to- prominulous or plane above, prominent be-
mentose within to base, sessile. Calyxlobes five, neath;petioles 8-12 mm long, glabrous,canalic-
SystematicTreatment75

ulate, rugose, eglandular. Stipules lanceolate, for dense pilosity aroundbase of stamens.Calyx
membranous,ca. 2 mm long, caducous. Inflo- lobes five, rounded,sparselypuberulouson ex-
rescencesof terminaland axillarypanicles, 4.5- terior,the marginsciliate. Petals five, white, the
9 cm long, or rarelyof unbranchedracemes;ra- margins ciliate. Stamens numerous, inserted
chis and branches with minute sparse pubes- arounda complete circle. Ovary densely pilose.
cence. Bractsand bracteolesreceptaclecylindri- Stylepilose for half of length.Youngfruitovoid,
cal-turbinate, 15-20 mm long, sparsely exocarpsmooth, glabrous;mesocarpfleshy;en-
puberulousin bud, soon becomingglabrous,gla- docarphard, fragile.
brous within except for dense deflexed hairs Distribution(Fig. 108).Knownonlyfromsouth
around throat. Calyx lobes five, sparsely pub- of Manaus.
erulous when young, soon becoming glabrous, Habitat. Forest on terrafirme.
the margins ciliate. Petals five, glabrous, ca-
Additional specimens examined. BRAZIL.
ducous.Stamensca. 110, insertedarounda com- Manaus-PortoVelho Hwy., betweenRios
AMAZONAS:
pletecircle,the filamentstomentosearoundbase, CastanhoandAraca,12 Jul 1972(st),M. F. da Silva
glabrousfor most of length. Ovary pilose. Style et al. 490(INPA,NY), 11Jul 1972(fl),M. F. da Silva
pilose at base, glabrousabove. Fruit unknown. 467 (INPA,NY);km 113, 12 Jul 1972(fl),M. F. da
Distribution(Fig. 103). Known only from the Silvaet al. 267 (INPA);Furode Castanho,estrada
Araca,22Jul1972(fl),M.F. daSilva1027(NY);Nova
lower Rio Negro region of Brazil. Olinda,Rio Paca,tributaryof Rio Marimari,2 Jul
Habitat. Forest on terrafirme. 1983(fl),Todzia2295 (INPA,NY).
Additional specimen examined.BRAZIL. AMAZONAS: Couepiamarleneaeis most closely related to
Manaus-Caracarai Rd.,km 115,Rio Urubu,13 Aug C. williamsii,differingin the muchsmallerleaves
1974(fl),Pranceet al. 21636(INPA,NY). and few primarynerves, the sparse pubescence
Couepiaglabrais quiteunlikeanyof the species and more reticulatevenation of the leaf under-
alreadydescribed,but it is probablyclosest to C. surface, and the more prominent conspicuous
williamsii from which it differs in the usually glandsat the junction of the lamina and the pet-
paniculate inflorescence,the larger leaves, the iole of the leaves. In C. williamsiithe leaves are
almost glabrousstyle, the glabrousundersideof 9-18 cm long, and 2.5-5.5 cm broad, and have
the leaves, as well as in a number of additional 12-15 pairsof primaryveins. Couepiamarlene-
small characters. ae is also rathersimilarto C. glabra,but it is not
so closely related, differingin the unbranched
6-35.2. Couepia marleneaePrance, Acta Ama- racemoseinflorescence,the much smallerleaves,
z6nica4(1): 17. 1974. Type. Brazil.Amazonas: the very sparse but definite pubescence of the
Manaus-PortoVelho Rd., between Rios Cas- exteriorof the flowers,and the glandsat the lam-
tanho and Tupana, 17 Jul 1972 (fl), M. F. da ina base.
Silva et al. 822 (holotype,INPA; isotype,NY).
6-38. Couepia trapezioanaCuatrecasas,Britto-
Small tree to 8 m tall, the young branches
nia8: 197. 1956. Fig. 118.
sparsely puberulous soon becoming glabrous.
Leaf lamina oblong-lanceolate,coriaceous, 5-8 Additionalspecimensexamined.PERU.LORETO: Rio
x 1.5-2.4 cm, base subcuneate,apex acuminate, Ampiacu,PucaUrquillo,24 Sep 1972 (fl),Croat20663
(MO, NY); Rio Nanay, Mishana, 16 Aug 1978 (fl),
the acumen 5-15 mm long, with two glandsto- Ramirez C. 87 (MO,NY).
wards base at junction with petiole, glabrous
above, with sparselylanate pubescencebeneath; 6-41.1.
midrib glabrousand prominulousabove, prom- Couepia dolichopodaPrance, Brittonia
26: 302. 1974. Type. Peru. Loreto:Varadura
inent beneath;primaryveins 7-10 pairs, prom-
de Mazan, from Rio Amazonas to Rio Napo
inent beneath; petioles 4-7 mm long, sparsely
(fl),Croat19382 (holotype,NY; isotype,MO).
puberulousor glabrous,canaliculateabove. Stip-
ules triangular,1 mm long. Inflorescencesof ter- Tree to 30 m tall, the young branchescom-
minalracemes,the rachisgray-puberulous. Bracts pletely glabrous. Leaf lamina elliptic, coria-
small and caducous. Receptacle cylindrical, ceous, 9-11.5 x 3.8-5.3 cm, cuneate at base,
swollen towardsbase, 15-18 mm long, sparsely acuminateat apex, the acumen 10-15 mm long,
puberulouson exterior, glabrouswithin except glabrouson both surfaces;midrib prominulous
76 Flora Neotropica

on both surfaces, glabrous;primary veins 6-8 Additionalspecimensexamined.VENEZUELA.

BOLIVAR:ReservaForestalImataca,Rio CuyuniSector
pairs,almost planeon both surfaces;petioles 10- CaiioNegro,15Jan1983(fl),Stergiosetal. 4991(MO,
12 mm long, glabrous, shallowly canaliculate.
NY);El Dorado,3 May1957(fl),Couret243 (US).
Stipules not seen. Inflorescencesof pendulous
panicles on long pedunclesup to 75 cm, the ra- 647.1. CouepianutansPrance,Brittonia31: 248,
chis and branchesglabrous,branchessmall and
fig. 1. 1979. Type. Colombia. El Valle: Alto
numerous. Bracts and bracteoles 0.5-1.5 mm Yunda, Rio Anchicaya, 1000 m, May 1973
long, acute, glabrousexcept for the sparselycil- (fl),HiltyA-1(holotype,US;isotypes,NY, UC).
iate margins,subpersistent.Receptaclecampan-
ulate, the tube 4-6 mm long when mature,con- Treeto 25 m, the youngbranchessparselypu-
tracted into pedicels 10-15 mm long, glabrous berulous,soon becoming glabrous.Leaf lamina
within and without. Calyx lobes five, rounded, elliptic, coriaceous,13-16 x 5-7.5 cm, rounded
unequal,two considerablylongerthan the other at the base, acuminateat the apex, the acumen
three,sparselypuberulouswhen young,the mar- 4-5 mm long, glabrous above, densely arach-
gins ciliate, the exterior with two sessile con- noid-pubescent beneath; midrib prominulous
spicuousglands.Petals five, glabrous,caducous, above, prominentbeneath;primaryveins 17-19
red. Stamens 16-21, unilateral, with ca. eight pairs, plane above, prominentbeneath;petioles
tooth-like staminodes opposite to them. Ovary 5-8 mm long, weakly canaliculate above, pu-
sparsely villous. Style glabrous. Fruit obpyri- berulous,with two glands.Stipulescaducous,not
form, 4.5 cm long, 3.5 cm wide at broadestpoint seen. Inflorescencesof terminaland axillarypan-
which is well below mid-point, exocarpsmooth, icles to 7 cm long, the rachis and brancheswith
glabrous;mesocarp thick, hard, fibrous;endo- shortlightbrowntomentellouspubescence.Bracts
carp thin and bony. ovate, membranous,ca. 5 mm long, caducous;
Distribution(Fig. 101). WesternAmazonia in bracteolesoblong, 2-3 mm long, puberulouson
the Peru-Brazilfrontierregion. exterior,membranous,subpersistent.Receptacle
Habitat. Forest on terrafirme. subcylindrical,ca. 5 mm long, shortly tomen-
tellous on the exterior,glabrouswithin exceptfor
Additional specimens examined. PERU. LORETO: deflexed hairs at the throat.
nr.BrillaNueva,RioYaguasyacu, 8 Nov 1977 Calyx lobes five,
Maynas, tomentellous. Petals
(fr),Gentry& Revilla20385 (MO, NY); QuebradaSu- rounded, five, white, gla-
cusari,N sideof RioNapobelowMazan,6 Nov 1979 brousexcept for the ciliate margins.Stamensca.
(fr), Gentryet al. 27591 (MO, NY). 16, insertedin a complete circle. Ovary villous.
BRAZIL.AMAZONAS: Rio Javari, 10 Aug 1973 (fl),
Lleraset al. P17294 (INPA, NY), 17 Dec 1975 (fr),N.
Style tomentose on lower third only. Fruit not
T. Silva4028(IAN). seen.
Distribution(Fig. 109). Known only from the
Uses. The fruit (cotyledons)is edible. type gathering.
Couepiadolichopodais closest to C. longipen- Habitat. Collected at an altitude of 1000 m,
dulabut differsin the largerpetioles, the reduced where it is an uncommon upland species.
numberof stamens, the glabrouspeduncles,the Couepianutansis closely relatedto C. polyan-
shorter campanulatecalyx tube and the mark- dra, a Mexican and Central American species,
edly unequalcalyx lobes. the rangeof whichextendssouthinto CostaRica.
Couepianutansdiffersfrom C. polyandrain the
6-45. Couepiacomosa Bentham,J. Bot. (Hook- larger leaves with a great number of primary
er) 2: 215. 1840. veins,the slightlycanaliculatepetioles,the slightly
flowerswith a darkerbrown pubescence,
Distributionandhabitat(Fig. 100).This species larger
and the pendulousleaves.
was known until recentlyonly from Guyana. It
has now been collected in Venezuelaand is not
6-48. Couepia platycalyx Cuatrecasas, Field-
cited for that countryin my accountfor the Flora
de Venezuela(Prance, 1982b). The Guyanama- iana, Bot. 27: 66. 1950.
terial is from rocky places beside rivers and one This is one of the most distinctive species of
of the Venezuelancollections(Stergioset al. 4991) Couepia because of the flattened, almost solid
is from rain forest. There is no doubt that the receptacle.A third and fourth collection add to
materialof this distinct species is conspecific. the two studied in 1972.
SystematicTreatment 77

Distributionand habitat (Fig. 111). It is con- lenticellate.Leaflamina oblong to oblong-ellip-

fined to the high altitude Andean forests of Co- tic, thickly coriaceous,6-13 x 3.5-5.5 cm, sub-
lombia and Venezuela. cordateto roundedat base, apex rounded,acute
or bluntly acuminate, glabrous and lustrous
Additional specimens examined. COLOMBIA.
ANTIOQIA. Mun El Retiro,nr.quebradaLaAguadelo, above, densely velutinous-arachnoid-ferrugi-
2200-2300 m, 12 Apr 1980 (fr),Bernal& Galeano142 neous beneath;midrib slightlyimpressedabove,
(HUA). prominentand lanatebeneath;primaryveins 9-
VENEZUELA.LAA 8 km fromSanare,1500-1800 13 pairs, impressedabove, prominentbeneath;
m, 29 Jul 1979 (f), Meijeret al. VEN92(NY).
petioles 6-10 mm long, lanate, with two glands
at base (visible in youngpetioles only) becoming
6-51. Cnmepiaimipessa Prance, Fl. Neotrop.
rugulosewith age. Stipules 1-2 mm long, ovate,
Mongr. 9: 255. 1972. Fig. 106.
membranous, caducous. Inflorescencesof ter-
This species is now divided into two subspe- minal denselycrowdedpanicleswith many short
cies, diiinguished in the following key. branchesonly, the rachis and branchespuberu-
lous. Bractsand bracteolessmall, membranous,
1. Leavesoblong-ovate, 19-22cm long,4-8.5 cm
broad-primaryveins 19-22pairs. very early caducous.Receptaclecylindrical,7-9
a. subsp. impressa. mm long, with short dense puberulouspubes-
1. Leavesovate,9-13 cm long,4-7.5 cm broad; cence on exterior,glabrouswithin except for de-
primaryveins 14-19pairs. b. subsp.cabraliae. flexed hairs at throat and dense lanate pubes-
cence around stamen bases; pedicels 2-6 mm
6-51b. C. impressa Prance subsp. cabraliae
long, thin. Calyx lobes five, acute, with sessile
Prance,Revista Brasil.Bot. 2: 31. 1979. Type. glandsaroundmargins.Petals five, glabrousex-
Brazil. Bahia:Santa Cruz de Cabralia,5 Nov
cept for ciliate margins.Stamensca. 17, inserted
1966 (fl), Belkn & Pinheiro 2837 (holotype, arounda
complete circle. Ovary densely lanate.
CEPEC;isotypes, MG, NY). Style pubescent for three-fourthsof its length.
Leaves ovate, 9-13 cm long, 4-7.5 cm broad; Fruit elipsoid-pyriform,taperingtowardsapex,
primaryveins 14-19 pairs;petioles7-10 cm long 4.5 cm long, 3 cm broad;exocarp smooth, gla-
brous; pericarphard, fibrous 3 mm thick, gla-
Representative specimensexamined.BRAZIL.BA- brous within.
HIA: SantaCrz de Cabralia, 4 Nov 1966(fl), BeIMm
& Pinheiro2846 (CEPEC,NY), 8 Feb 1967 (y fr), Habitat. Restinga.
Belem & PineMro3315 (CEPEC,NY), 21 Oct 1978 Additional specimens examined. BRAZIL. BAHIA:
(fl),MFriet al. 10934(CEPEC,
NY);9 km E of Una, Mun.Marau,rd.to Portode Campinhos, 17kmfrom
3 Dec 1981 (t Carwvao& Lewis877 (CEPEC,NY); 7 Jan 1982(fl),Carvalho& Lewis1106(CE-
Portode Campinhos-MaraiRd., km 11, 26 Feb 1980 Marau,
PEC,NY); km 11, Portode Campinhos-Marai, 26
(fr),Carmlko et al. 216 km
(NY); rd.
12, Porto Seguro Feb 1980 (fl), Carvalhoet al. 194 (CEPEC,NY).
to EunlpoEis,9 Nov 1972 fr),Euponino309 (CEPEC,
NY), 7-9 km W of PortoSeguro,1 Dec 1978 (fl), This speciesis best distinguishedby the dense-
Eupoino 405 (CEPEC,NY); 4 km N of Uruguca,4
Nov 1978 (fl), Mori 11037 (CEPEC,NY); Ilheus, 12 ly clustered inflorescencewith numerous short
Nov 1979 (fl), T. S. dos Santos 1279 (CEPEC,NY). branches,also by the small sessile glandsaround
the marginof the calyx lobes, and the extremely
This is a common subspeciesof restinga. lustrousleaf upper surfaces.It is probablymost
Local name:oiti mirim. closely related to the more southerly C. merid-
ionalis, from which it also differs in the im-
6-52.1. CouepiacoarctataPrance,sp. nov. Type.
pressed venation of the leaf upper surface and
Brazil. Bahia: Mun. de Marafu, 10 km S of the glabrouspetals. Couepiacoarctatais named
Portode Campinhoson rd. to Ubaitaba,7 Jan for the crowded,clusteredinflorescence.It is not
1982 (fl, fr), Carvalho & Lewis 1116 (holotype,
easily confusedwith any other species of the ge-
CEPEC;isotype, NY). Figs. 18, 100. nus. It was collected in two places on coastal
Species a C. meridionaliaffinis, petalis extus restingaon the small peninsulathat runs north
glabris,nervis primariissupraimpressis,calycis from Marau.
lobis glandulissessilibus munitis differt.
Small tree to 6 m tall, the young branches 6-52.2. Couepia longipetiolataPrance, Revista
sparselypuberulous,soon becomingglabrousand Brasil. Bot. 2: 31, fig. 3. 1979. Type. Brazil.
 r.~?
'~~~~~~~~~~~~~~~~~~~~.'; ..

'" Y~~~~~~~~~~~~~X
l

?? I /? I ?1 ----I , ,?=, E

?.

"r;~~~~~~~~~~~~~
.
?:~
~~~~Cup..
~ ~ .-
.

coarctata
FIG.18. Coucpia ] d). A habit;B, fruit;C, leaf udersurface; ]D, flower
Carvalho& Zwwi$1] G, bud
of calyxlobe; F, flower section;
E, glandular margin petal.~~~~~?
eoa 78~~~~~~~~:
CO4C&t,,1

FIG.18. Couepia coarctata (Carvaho & Lewis 16). A, habit; B, fruit; C, leaf undersurface; D, flower bud;

marginof calyx lobe; F, flowersection;G, petal.

E, glandular

78
SystematicTreatment 79

Bahia:Itacare,Estradado Aeroporto,28 Feb tose almost to apex, slightly exceeding the fila-
1975 (fl), T. S. dos Santos 2935 (holotype,CE- ments. Fruit not seen.
PEC;isotypes, FHO, NY). Distribution(Fig. 107). Known only from two
collections from the forests of Bahia.
Tree 8 m tall, the young branches glabrous,
Habitat. Littoralforest.
conspicuouslylenticellate.Leaf lamina oblong,
coriaceous, 12-15 x 4-4.5 cm, cuneate at base, Additionalspecimenexamined.BRAZIL.BAHIA: 22
shortlyand abruptlyacuminateat apex, the acu- kmEof Ubaitaba,25 Aug1979(st),Mori12749(NY).
men 2-3 mm long, glabrous above except on Local name. oiti.
midrib, lanate brown-pubescentbeneath, ap- Couepialongipetiolatais related to the more
pearingsmooth but pubescencecoveringa deep- southernspecies C. meridionalis,differingin the
ly reticulatevenation with stomatalcrypts,mid- narrowerleaves, the much denserinflorescence,
rib prominent beneath, slightly impressed and the longer petioles and the glabrousexterior of
lanate on basal half above; primaryveins 11-13 the petals.It differsfrom C. schottiiin the longer
pairs, slightly prominent beneath, slightly im- petioles, the fewer primaryleaf veins, the dense
pressedto planeabove;petioles 14-18 mm long, inflorescence,and the greaternumberof stamens
canaliculateon uppersurface,lanate pubescent, insertedarounda complete circle. Couepialon-
transverselyrugose,2-3 mm thick.Stipulessmall, gipetiolatais not easily confusedwith any other
ovate, membranous,caducous.Inflorescencesof easternBrazilianspecies becauseof its long pet-
terminal and subterminal panicles, densely ioles and distinctive, densely crowded, short-
crowded,shortbranched,the rachisand branch- branchedinflorescence.
es rufous-tomentellous.Bractscaducous,to 5 mm
long, membranous, glabrous; bracteoles ca- 6-53. CouepiapernambucensisPrance,Fl. Neo-
ducous,ca. 1 mm long tomentellouson e : rior. trop. Monogr.9: 256. 1972.
Receptaclecampanulate-turbinate, 4-5 mm long,
rufoustomentellouson exterior,glabrouswithin Distribution(Fig. 111). This species,described
except for deflexed hairs at throat. Calyx lobes from a single collection from Pernambuco,has
acute,tomentellouson both surfaces.Petalsfive, now also been collected in Bahia.
cream,glabrouswith ciliatemargins.Stamens ca. Additionalspecimenexamined.BRAZIL.BAHIA: Una,
30, insertedarounda complete circle, far exsert- Fazenda SaoRafael,16Dec 1968(fl),T.S. dosSantos
ed. Ovaryvillous-pubescent.Stylepilose-tomen- 322 (CEPEC, NY).

7. Hirtella L.
Distributionof Hirtella as a whole is shown in Figure 119.

Key to Groups of Hirtella

1. Leaf base with myrmecophilousswellings. GroupA: SectionMyrmecophila.
1. Leafbase without myrmecophilousswellings. Section Hirtella.
2. Inflorescencea panicle (branched). GroupB.
2. Inflorescencea raceme(unbranched). Group C.

Key to Group A: Hirtella Section Myrmecophila

1. Inflorescencefasciculate(bunchedracemes)sometimes cauliflorous.
2. Stamens 4-5; ovary inserted at middle of receptacle;leaves 9-19 cm long, coriaceous,the lower
surfacebullate. 1. H. myrmecophila.
2. Stamens6-7; ovary insertedat mouth of receptacle;leaves 17-30 cm long, membranous,the lower
surfacenot bullate. 2. H. physophora.
1. Inflorescencean elongateracemeor panicle,never cauline.
3. Inflorescencea panicle,sometimes only slightlybranchedat base.
4. Inflorescenceonly slightlybranchedat base; leaves oblong-lanceolate;bractseglandular.
3. H. vesiculosa.
4. Inflorescencewith many small lateralbranches;leaves ovate; bractsglandular. 4. H. dorvalii.
80 Flora Neotropica

3. Inflorescencean elongateraceme,never branched.

5. Exteriorof receptacleand calyx lobes hirsutulous;inflorescence15-30 cm long. 5. H. guainiae.
5. Exteriorof receptacleand calyx lobes hispid only; inflorescence2-12 cm long.
6. Pedicels 4-6 mm, with sparsehirsutepubescence. 6. H. duckei.
6. Pedicels 8-15 mm with dense hirsutepubescence. 6.1. H. revillae.

Key to Hirtella Group B: Species without myrmecophilousswellings at

leaf base and with a paniculateinflorescence.
1. Bractsand/or pedicelsglandular(with stalkedor sessile glands).
2. Leavesglabrousbeneathwhen mature,or rarelywith a few short stiff appressedhairs.
3. Stamens6-7.
4. Bractswith a few sessile glands only; inflorescencerachisand branches,and flowerexterior
glabrousor only sparselyhirsutulous.
5. Inflorescencea spreadingbranchedpanicle,the rachisand branchesglabrous.
7. H. macrosepala.
5. Flowersbornein clustersof 2-3 alonga centralrachisratherthana branchedinflorescence,
giving a racemoseappearance,the rachisand brancheshirsutulous.
6. Leaflamina 6-9.5 cm long, 2.5-3.8 cm broad,the acumen 9-14 mm long;petioles 2-
2.5 mm long. 11.1. H. barnebyi.
6. Leaflamina 21-27 cm long, 6-9 cm broad,the acumen 1-5.5 mm long; petioles 8-10
mm long. 11.2. H. margae.
4. Bractswith numerousstalkedglands;inflorescencerachis and branches,and flowerexterior
pubescent.
7. Leaves 3.5-8.5 cm long, 1.4-4.5 cm broad;stipuleslinear,to 6 mm long (Africa).
90. H. zanzibarica.
7. Leaves (6.4-)8-13 cm long, 4-7 cm broad;stipulesdeltoid, to 2 mm long (neotropics).
8. Calyxlobes glandular;bractglandssmall, ca. 0.1 mm in diameter;rachisand branches
of inflorescencesparselypuberulous;leaves glabrousbeneath. 8. H. ulei.
8. Calyx lobes eglandular;bractglands large,ca. 0.5 mm in diam.; rachis and branches
of inflorescencetomentellous;leaves sparselyappressedpubescentbeneath.
9. H. glabrata.
3. Stamens3-5.
9. Glands large and solitary on a long stalk, arisingfrom pedicel or junction of inflorescence;
inflorescencedistinctlythyrsoid. 10. H. carbonaria.
9. Glands usually numerous, long or short stalked, arising from the margins of bracts and
bracteoles;inflorescencenot thyrsoid.
10. Inflorescencea long rachisbearingsmall clustersof flowerson very shortbranches;leaves
oblong-lanceolate. 11. H. araguariensis.
10. Inflorescencespreading,with many distinct lateralbranches,flowersnot in distinctclus-
ters; leaves oblong to elliptic.
11. Leaves 3-5.5 cm long, cordateat base;inflorescencescarcelybranched,hirsutulous.
12. H. cordifolia.
11. Leaves usuallycuneateor subcuneateat base (if subcordatethen 7-17 cm long and
inflorescenceglabrescent);inflorescencesusuallymuch branched.
12. Bracteolarglands few, usually either sessile on marginsor a single gland ter-
minatingapex;inflorescenceusuallypilose or if glabrescentthen corymbose;leaf
apex acuminate.
13. Leaves 12-17 cm long, roundedto subcordateat base; fertile stamens 5.
13. H. insignis.
13. Leaves 3.5-15 cm long, cuneateat base; fertile stamens 3.
14. Leaves 10-15 cm long; bractsmembranous,the glands several,borne
on margins;inflorescence7-18 cm long, terminal. 14. H. tocantina.
14. Leaves4-9 cm long;bractscoriaceous,with singlereflexedapicalgland
only; inflorescence3-11 cm long, axillaryor terminal. 15. H. piresii.
12. Bracteolarglandsnumerous,borne on the margins,usuallyshort-stalked;inflo-
rescencespreadingbut not corymbose,usuallyglabrescent,if tomentellousthen
leaf apex roundedto acute.
15. Leaves 1.8-4.5 cm long, roundedto acuminateat apex, thicklycoriaceous;
primaryveins 4-7 pairs. 37. H. bahiensis.
15. Leaves6-13.5 cm long, acuminateat apex,thinlycoriaceous;primaryveins
9-10 pairs.
SystematicTreatment 81

16. Stamens3; bracteolescoriaceous,ovate; petioles 3-4 mm long.

16. H. davisii.
16. Stamens4; bracteolesmembranous,lanceolate;petioles 1-2 mm long.
17. H. subglanduligera.
2. Leaves hirsutebeneathat least on principalvenation, usuallymore generallyso.
17. Calyxlobes with small stipitateglands.
18. Leaves orbicularto ovate-elliptic,3-6.5 cm long, retuseto mucronateat apex (rarelyacu-
minate);trunkwith corky bark;receptaclepubescentwithin to base. 18. H. ciliata.
18. Leavesoblong to ovate, 4.5-23 cm long, acute to acuminateat apex;trunkwith thin bark;
receptacleglabrouswithin except at throat.
19. Venationof leavesprominulousabove;inflorescencesubcorymbose,bearingnumerous
tightlyclusteredflowers;stamens 7. 19. H. hoehnei.
19. Venationof leaves impressedor prominulousabove; inflorescencelax, spreading,not
subcorymbose,the flowersloosely arranged;stamens 3-5.
20. Leaves subcordateat base, subconduplicate,the lower surface merely sparsely
appressed-pubescent. 29. H. adderleyi.
20. Leaves roundedto cuneate at base, rarelyweakly subcordatebut never subcon-
duplicate,the lower surfacehirsute. 20. H. glandulosa.
17. Calyx lobes eglandular.
21. Venationof leaves impressedabove, or the leaf surfacebullate.
22. Inflorescenceandflowersrufous-tomentose;bractsbearingveryfew sessileglands,these
often obscuredby pubescence. 42. H. obidensis.
22. Inflorescenceand flowerstomentellousto puberulous,not rufous;bractsbearingmany
obvious stalkedor sessile glands.
23. Leaf surfaceusuallydistinctlybullate,the base usuallysubcordate. 21. H. bullata.
23. Leaf uppersurfaceplane, but with impressedvenation, the base roundedto sub-
cuneate.
24. Bracteolarglandsca. 0.5 mm in diam., sessile or slightlystipitate;bracteoles
distinct, ovate. 22. H. americana.
24. Bracteolarglands ca. 0.1 mm in diam., on slenderstalks;bracteolesreduced
to a mass of glandswith a slightlyconnatebase. 27. H. tentaculata.
21. Venationof leaves prominulousor plane above, the leaf surfacenever bullate.
25. Stamens3.
26. Bracteolarglands large, numerous,apparent;inflorescencetomentellous,many-
flowered;exocarpglabrescent;endocarpthin and bony. 22. H. americana.
26. Bracteolarglands small, few; inflorescencehispid, few-flowered;exocarptomen-
tose; endocarpthick and fibrous. 23. H. guatemalensis.
25. Stamens4-7.
27. Exocarptomentose; endocarpthick and fibrous;bracts with few inconspicuous
stipitateor sessile glands.
28. Inflorescencehispid, the flowerslax; bractswith small short-stipitateglands;
stamens4. 23. H. guatemalensis.
28. Inflorescencetomentellous,bearingsmall clustersof flowerson a long thick
rachis;bractswith a few largesessileglandsor a singleterminalgland;stamens
6. 24. H. eriandra.
27. Exocarpglabrescent;endocarpthin and bony;bractswith many conspicuousstip-
itate glands,except in H. liesneri.
29. Inflorescencehispid-hirsute;bractswith short-stalkedglandsarisingfromtheir
margins,the secretorytip small, ellipsoid.
30. Inflorescencespredominantlyterminal;leaf basesroundedto weaklysub-
cordate. 25. H. paniculata.
30. Inflorescencespredominantlyaxillary;leaf bases deeply cordate.
26. H. deflexa.
29. Inflorescencepuberulousto tomentellous;bractseitherreducedto numerous,
long-stalkedclavateglandsarisingfrom the junction of the pedicelswith the
stem, or ovate with glands borne on their margins,the secretorytip large,
flattened,formingan expandedhead.
31. Glands arising from junction of pedicel and stem, with a long slender
stalkand a small secretorytip, the whole appearingclavate;youngstems
puberulousor sparselytomentellous. 28. H. macrophylla.
31. Glands arisingfrom the bracts,each gland with a thick stalk and a large
secretorytip, flattenedto forman expandedhead;youngstems with short
dense compact puberulence.
82 FloraNeotropica
32. Leaves4-7 cm long, conduplicate;inflorescence3.5-8 cm long;stip-
ules lanceolate. 29. H. adderleyi.
32. Leaves 6.5-15 cm long, not folded; inflorescence12-19 cm long;
stipulesdeltoid or linear-lanceolate.
33. Inflorescencesca. 19 cm long, with very short lateralbranches
giving a racemoseappearance;glands scarce,many bractswith
single stalkedglandat apex;inflorescencerachissparselyhispid
at base. 11.3. H. liesneri.
33. Inflorescences12-15 cm long, not appearingracemose;glands
numerouson bracteoles;inflorescencerachistomentellous,never
hispid.
34. Leavescordateat base;petioles 1-3 mm long;stipuleslinear,
5 mm long; flowersdensely crowded. 9.1. H. confertiflora.
34. Leavesroundedto subcordateat base;petioles5-7 mm long;
stipulesdeltoid, to 2 mm long; flowerslaxly arranged.
9. H. glabrata.
1. Bractsand pedicels eglandular.
35. Leaf apex retuse,rounded,or acute, never acuminate.
36. Stamens 5-6; leaf apex mostly roundedto retuse(or mucronulate). 30. H. punctillata.
36. Stamens3-4; leaf apex acute.
37. Leaves elliptic, thick-coriaceous;inflorescencecompact,to 3 cm long. 31. H. corymbosa.
37. Leaves oblong, chartaceous;inflorescencelax and spreading,3.5-17 cm long.
36. H. triandra.
35. Leaf apex distinctlyacuminate,sometimes abruptlyso and mucronateor cuspidate.
38. Stamens3.
39. Leaf base subcordate;inflorescence25-55 cm long, pendulous. 33. H. pendula.
39. Leaf base roundedto cuneate;inflorescence1-15 cm long, erect.
40. Leaves lanceolate,at least three times longerthan broad;inflorescenceand exterior
of flowersglabrescent. 32. H. barrosoi.
40. Leaves oblong to elliptic, not exceeding 2.5 times longer than broad; inflorescence
and exteriorof flowersusuallypubescent,rarelyglabrous.
41. Inflorescenceandexteriorof flowersglabrousor glabrescent,or leaf apexdistinctly
cuspidatewith a long fine acumen;inflorescences1-4 cm long, predominantly
axillary. 50. H. bicornis.
41. Inflorescenceand exteriorof flowerspubescentor tomentose;leaf apexacuminate
but never cuspidate;inflorescences2-17 cm long, predominantlyterminal.
42. Inflorescenceand youngstem hispid;calyx tube ca. 6 mm long, the lobes ca.
5 mm long; corollalobes ca. 8 mm long; stipules 6-10 mm long.
34. H. leonotis.
42. Inflorescenceusuallypuberulous,rarelyweaklyhirsute;youngstem glabrous
to tomentellous;calyx tube to 4 mm long, the lobes to 3 mm long; corolla
lobes to 5 mm long; stipules 2-6 mm long.
43. Inflorescencelittle-branched,almost racemose,but with a few branches
bearing two flowers, others one flower and a single pair of bracts at
junction with pedicel. 35. H. mutisii.
43. Inflorescencemuch-branched,the branchesusuallybearingmanybracts.
44. Leaves 4-14.5 cm long, 2-5.5 cm broad;pedicels 1-3 mm long.
36. H. triandra.
44. Leaves 15-17 cm long, 8-10 cm broad;pedicels 0-0.5 mm long.
38. H. latifolia.
38. Stamens4-9.
45. Inflorescenceappearingracemose,but in fact either a long centralrachisbearinggroups
of flowers on short branches,or with a few brancheswith only three flowerseach; leaf
base sometimes cordate.
46. Inflorescencealmost a raceme,but with a few branchesbearingtwo or three flowers;
flowerslaxly arranged,not borne in distinct groups. 85. H. hebeclada.
46. Inflorescencea long centralrachiswith groupsof flowerson short branches.
47. Bracteolesexceeding receptaclein length, persistent,enclosing buds; leaf base
subcuneate;exocarpglabrous. 39. H. suffulta.
47. Bracteolesmuch shorterthan receptacle,not persistent,not enclosingbuds, leaf
bases subcordateto subcuneate;exocarptomentose or glabrous.
48. Leaves 18-40 cm long, 8-16.5 cm broad,with 14-18 pairsof primaryveins,
and 2 glandsat junction of uppersurfaceof laminaand petiole;inflorescence
rufous-tomentellous. 40.1. H. magnifolia.
SystematicTreatment 83
48. Leaves 8-23 cm long, 3.3-8 cm broad, with 10-13 pairs of primaryveins,
the lamina base eglandular;inflorescencegray-puberulousor brown-tomen-
tellous.
49. Leaves subcordateat base, the lower surfaceglabrescent;exocarpgla-
brous. 40. H. elongata.
49. Leavesroundedto subcuneateat base, the lower surfacesparselypilose-
hirsute;exocarptomentose. 24. H. eriandra.
45. Inflorescencea much-branchedpaniclewithouta long centralrachis;leaf base roundedto
cuneate.
50. Inflorescenceand exteriorof flowerstomentose, hirsute,or hispid, usuallyrufousto
golden-brown.
51. Inflorescenceand young branchessparselyhispid;leaves membranous.
41. H. rodriguesii.
51. Inflorescenceand young branchestomentose or hirsute;leaves chartaceousto
coriaceous.
52. Venationof leaves slightlyimpressedabove;inflorescencerufous-tomentose.
42. H. obidensis.
52. Venation of leaves plane or prominulousabove; inflorescencebrown-to-
mentose or hirsute.
53. Leaves 2.5-6 cm long; inflorescence0.8-4 cm long.
54. Inflorescence0.8-1.5 cm long, clustered;leaves ovate, roundedat
base; stamens 7. 43. H. cowanii.
54. Inflorescence2-4 cm long, lax;leaves orbicularto oblong-orbicular,
usuallysubcordateat base; stamens 5. 44. H. orbicularis.
53. Leaves 5.5-17 cm long; inflorescence4-11 cm long.
55. Bracts4-6 mm long, ovate, up to halfthe lengthof receptacle;leaves
ovate, thick-coriaceous,shortly mucronateat apex; inflorescence
tomentose. 45. H. guyanensis.
55. Bracts 1-3 mm long, oblong, to base of receptacleonly; leaves el-
liptic, subcoriaceous,acuminate(never mucronate)at apex; inflo-
rescencehirsutulous-tomentose. 46. H. lightioides.
50. Inflorescenceand exteriorof flowersgray-puberulousto glabrescent.
56. Inflorescencea subcorymbosepanicle 4-6.5 cm long;petioles 7-8 mm long.
47. H. aramangensis.
56. Inflorescencea spreadingpanicle (not subcorymbose);petioles 0.5-5 mm long.
57. Lamina9-15 cm long,withdistinctbasalglands;inflorescencelax andspread-
ing, 7-19 cm long. 48. H. rasa.
57. Lamina 2.5-8(-9.5) cm long, the base eglandularor with indistinctglands;
inflorescenceshort and compact, 1-8 cm long.
58. Inflorescenceand exteriorof flowersdensely puberulous;leaves thick
and coriaceous,with prominentvenation,the apex acuminatebut never
cuspidate. 49. H. scabra.
58. Inflorescenceand exteriorof flowersglabrescentto puberulous;leaves
chartaceous,with plane to prominulousvenation, the apex acuminate
to cuspidate. 50. H. bicornis.

Key to HirtellaGroupC: Specieswithoutmyrmecophilous swellingsat

leafbaseandwith a racemoseinflorescence.
1. Glandspresenton bractsand bracteolesor pedicels (eitherstipitateor sessile).
2. Leavesnarrowlylinear-lanceolate,5.5-18 cm long, 0.5-2.2 cm broad;rheophytic.
51. H. angustissima.
2. Leaves ovate to oblong-lanceolate,3-30 cm long, 1.5-10 cm broad;not rheophytic.
3. Bractsand bracteolesbearingsessile glands only or with translucentglandularexcretions,the
glandsoftencoveringentiresurface,most frequentlypairedtowardbasesof bractsand bracteoles;
stipitateglandsabsent from bractsand pedicels,but bractapex occasionallyglandular.
4. Leaves oblong-lanceolate.
5. Leaves9-14 cm long;youngstemspuberulous;pedicels5-10 mm long;stipuleseglandular.
52. H. tenuifolia.
5. Leaves5-8 cm long;youngstems hispid;pedicels 12-16 mm long;stipuleswith numerous
stipitateglands. 52.1. H. radamii.
4. Leaves oblong to ovate.
6. Young stem hispid;inflorescencerachissparselypuberulous. 56. H. racemosa.
84 Flora Neotropica

6. Young stem and inflorescencerachispilose or puberulous,never hispid.

7. Youngstem andinflorescencethick-piloseor pilose-tomentose,the stemwith persistent
hairs 2-3 mm long; stamens 3-6.
8. Leaf marginshispid-ciliate;stamens 5-6; pedicels 1.5-3 mm long; bractsaxillary.
53. H. pilosissima.
8. Leafmarginseciliate;stamens3; pedicels3-6 mm long;bractsinsertedon pedicels.
34. H. leonotis.
7. Young stem and inflorescenceusually puberulous,sometimes tomentellous;stamens
4-6.
9. Leaves thin, membranous;stamens 4-5; ovary insertedmidwayup receptacle.
41. H. rodriguesii.
9. Leaves thick, coriaceous;stamens 5-6; ovary insertedat mouth of receptacle.
10. Bractsandbracteolescaducous,with minutetranslucentsecretions,sessileglands
presentor absent;pedicels 6-25 mm long.
11. Bractswith sessile glandsin addition to translucentsecretion;pedicels 6-
16 mm long,thick;inflorescence4.5-14 cm long;leaves sparselyappressed-
pubescenton venation or glabrescentbeneath. 54. H. gracilipes.
11. Bractswith translucentglandularsecretionsonly, otherglandsabsent;ped-
icels 12-25 mm long, very slender;inflorescence1.5-5 cm long; leaves
glabrousbeneath. 55. H. brachystachya.
10. Bractsand bracteolespersistent,with sessileglandsonly;pedicels 1.5-10.5 mm
long.
12. Primaryveins 6-10 pairs; leaves 3.5-18.5(-19.5) cm long, glabrous or
sparselyappressed-pubescent beneath;matureleaves plane, not bullate.
56. H. racemosa.
12. Primaryveins 11-15 pairs;leaves 19.5-25 cm long or if smaller(7-13 cm
long) then hirsutulousbeneath;matureleaves slightlybullate.
13. Leaves 19.5-25 cm long; primaryveins 13-15 pairs; lower surface
with few sparseappressedhairs. 57. H. juruensis.
13. Leaves 7-13 cm long; primaryveins 11-13 pairs;lower surfacehir-
sutulous. 58. H. kuhlmannii.
3. Eitherbractsand bracteolesbearingstipitateglands,or with one or more solitarystipitateglands
arisingfrom some pedicels, and bractsoften glandularat apex only.
14. Bracts and bracteoleseglandular,or with sessile glands and a glandularapex only; some
pedicels with 1-3 stipitateglands.
15. Leaves ovate, cordateat base; bractsand bracteoleswith a few sessile glands.
59. H. standleyi.
15. Leaves oblong to oblong-lanceolate,usuallyroundedto subcuneateat base, rarelysub-
cordate;bractsand bracteoleseglandularor with sessile glands.
16. Primaryveins 17-19 pairs;leaves 19-26 cm long; pedicels 8-12 mm long.
60. H. longifolia.
16. Primaryveins 8-15 pairs;leaves 8-18 cm long (to 24 cm in one specieswith short
pedicels 0.5-2 mm long);pedicels 0.5-9 mm long.
17. Pedicels slender,4.5-9 mm long (or if 3.5-5 mm then leaf constrictedabove
base);leaves glabrousbeneathexcept for pubescenceon midriband marginof
extremebase of lamina;bracteoleseglandularor glandular.
18. Fertilestamens 3; leaves oblong-elliptic;bractswith apical glands.
61. H. lemsii.
18. Fertile stamens 5-7; leaves elliptic to oblong-lanceolate;bractsglandular
or eglandular.
19. Leaves elliptic to oblong-elliptic,3.5-8 cm long; bractsusually with
sessile and apical glands. 56. H. racemosa.
19. Leaves oblong-lanceolate,8-18 cm long; bractseglandular.
20. Leaves subauriculatebecause of slight constrictionabove base;
primaryveins 11-13 pairs;ovary insertednear mouth of recep-
tacle;fertilestamens 5. 62. H. schultesii.
20. Leaves not constricted above base; primary veins 8-15 pairs;
ovary insertedat mouth or middle of receptacle;fertile stamens
4 or 6.
21. Primaryveins 12-15 pairs;secondaryvenation slightlyim-
pressed;ovaryinsertedmidwayup receptacle;fertilestamens
4. 63. H. paraensis.
SystematicTreatment 85
21. Primary veins 8-10 pairs; tertiary venation prominulous;
ovary insertedat mouth of receptacle;fertilestamens 6.
64. H. sprucei.
17. Pedicels 1-3 mm long, thick; leaves hirsutebeneathor glabrouswith hirsute
marginand midrib;bracteoleswith glandularapex and often sessile glands.
22. Leaves bullate above, with distinctlyimpressedvenation, secondaryve-
nation and reticulationsextremelyprominentbeneath. 65. H. lancifolia.
22. Leaves not bullate above, the venation prominent or plane, secondary
venation and reticulationsprominulousonly beneath.
23. Leavesacuminateat apex, the lower surfaceglabrousto sparselyhir-
sute. 66. H. burchellii.
23. Leaves usually mucronateat apex, the lower surfacedensely hirsute
or glabrous.
24. Inflorescencedensely tomentose;leaves mucronateat apex, hir-
sute beneath. 67. H. mucronata.
24. Inflorescencesparselypuberulous;leaves acuminateat apex, gla-
brescentbeneath. 56. H. racemosa.
14. Bractsand bracteoleswith numerousstipitateglands;pedicelseglandular.
25. Receptaclecylindrical,elongate. 79. H. couepiiflora.
25. Receptaclecampanulate.
26. Leaves bullateabove, with impressedvenation. 21. H. bullata.
26. Leaves plane, not bullateabove, with plane or prominulousvenation.
27. Leaves orbicular,roundedto retuseat apex; pedicels 7-20 mm long.
68. H. longipedicellata.
27. Leaves oblong to elliptic, acuminateat apex;pedicels 4-11(-15) mm long.
28. Stipulesbearingnumerousstipitateglands. 69. H. glandistipula.
28. Stipuleseglandular.
29. Inflorescenceshort, compact, densely flowered, 3-7 cm long; calyx
bearingnumerousglands;leaves dryinggray. 70. H. martiana.
29. Inflorescencelax, 3-28 cm long;calyx eglandularor the glandsscarce;
leaves dryinggreento brown.
30. Calyx lobes 5-6 mm long; stamens 7-8; flowers8-9 mm long.
82. H. angustifolia.
30. Calyxlobes 1.5-4 mm long;stamens3-6; flowersusually4-6 mm
long (7-9 mm in H. santosii).
31. Youngbranchesandlowerpartofinflorescencerachishispid-
setose.
32. Leaves 2.5-6.5 cm long; inflorescence3-5 cm long.
71. H. pimichina.
32. Leaves 6-15 cm long; inflorescence6-28 cm long.
33. Leavesentirelyglabrousbeneath,driedmaterialwith
silver metallic sheen. 72. H. subscandens.
33. Leaves hirsutebeneath,especiallyon principalve-
nation,driedmaterialwithoutsilvermetallicsheen.
34. Bracteolesoblong, broad towardbase, charta-
ceous, with short-stalkedglands,the apicester-
minatingin a short-stalkedgland.
25. H. paniculata.
34. Bracteoles linear-lanceolate,coriaceous, with
long-stalkedglands, the apices always termi-
natingin a long-stalkedgland. 73. H. hispidula.
31. Young branchestomentellousto puberulous;lower part of
inflorescenceglabrousto puberulousor tomentellous.
35. Inflorescencetomentoseor tomentellous.
36. Inflorescencetomentellous,4.5-27 cm long, leaves
broadestabove middle, primaryveins 8-12 pairs;
bract glands predominantlystipitate;stipules 2-3
mm long. 74. H. silicea.
36. Inflorescence tomentose, 6-10 cm long; leaves
broadestat middle,primaryveins 15-19 pairs;ped-
icels 5-6 mm long;bractglandspredominantlyses-
sile; stipules 4-5 mm long. 22.1. H. santosii.
86 FloraNeotropica
35. Inflorescenceglabrescentor very sparselysetose.
37. Stamens3; calyx glandular. 75. H. excelsa.
37. Stamens5-6; calyx eglandular.
38. Leafgraduallytaperedto apex from nearbase;
pedicels 3-5 mm long. 76. H. adenophora.
38. Leaf apex cuspidate;pedicels 8-15 mm long.
77. H. caduca.
1. Bractsand pedicels eglandular.
39. Inflorescenceclustered,fasciculate;leaves slightlybullate,8-12.5 cm long;ovaryinsertednearbase
of receptacle. 78. H. fasciculata.
39. Inflorescenceof elongateracemes;leaves usuallyplane, if bullatethen small; ovary insertedat or
near mouth of receptacle.
40. Receptacleelongate-cylindrical; calyx lobes glandular.
41. Inflorescencesdensely tomentose;leaves 10-15 cm long; flowers 17-20 mm long.
79. H. couepiiflora.
41. Inflorescencessparselypuberulous;leaves to 8.5 cm long; flowers7-9 mm long.
80. H. tubiflora.
40. Receptaclecampanulate;calyx lobes eglandular.
42. Leaves orbicular,predominantlyretuseto roundedat apex;pedicels 7-20 mm long.
68. H. longipedicellata.
42. Leaves oblong-lanceolateto ovate, acuminateat apex; pedicels to 15 mm long, usually
shorter.
43. Lower surface of leaf hirsute, pilose or hispid at least on primaryand secondary
venation.
44. Leaves oblong, oblong-lanceolate,or lanceolate,the venation sometimes deeply
impressedabove; fertile stamens 7-9.
45. Leaves oblong, 3.5-8 cm long, venation deeply impressedabove; flowers4
mm long. 81. H. floribunda.
45. Leaveslanceolateto oblong-lanceolate,5-15 cm long;venationplaneabove;
flowersca. 6 mm long. 82. H. angustifolia.
44. Leaves ovate to elliptic, the venation usually prominentabove, rarely slightly
impressedbut then leaves broadlyovate; fertile stamens 3-9.
46. Leafvenation slightlyimpressedabove;leaves broadestnearbase, 2.5-8 cm
long, subcordateat base; inflorescencedensely crowded,with short rachis.
83. H. rugosa.
46. Leaf venationlevel to slightlyprominentabove;leaves broadestat or above
middle, subcuneateto roundedat base;inflorescencelax with longerrachis.
47. Stamens 3. 35. H. mutisii.
47. Stamens 5-7.
48. Leafapex mucronateto abruptlyacuminate;pedicels2-3 mm long.
84. H. scaberula.
48. Leaf apex acute to acuminate but never mucronate or abruptly
acuminate;pedicels 5-8 mm long.
49. Inflorescenceandexteriorof flowersdenselytomentose;flowers
5-8 mm long. 85. H. hebeclada.
49. Inflorescenceand exteriorof flowerspuberulous,flowers 4-5
mm long. 56. H. racemosa.
43. Lower surface of leaf glabrous or with few sparse appressedhairs on midrib and
primaryveins.
50. Leaves narrowlylanceolate,5.5-18 cm long, 0.5-2.2 cm broad.
51. H. angustissima.
50. Leaves ovate to oblong.
51. Fertilestamens 7-9.
52. Petalsslightlyclawed,leaf base subcuneate;youngbranchesglabrescent;
inflorescenceof densely crowdedracemes. 70.1. H. parviunguis.
52. Petals not clawed;leaf base subcordateto subcuneate;young branches
puberulousto hispid;inflorescenceof single racemes.
53. Leaf base subcordate;calyx lobes 4-6 mm long; young branches
sparselyhispid. 82. H. angustifolia.
53. Leaf base subcuneateto rounded;calyx lobes 1.5-2.5 mm long;
young branchessparselypuberulous. 86. H. enneandra.
51. Fertilestamens 3-6.
54. Fertile stamens 3; inflorescenceeither 2-3-flowered or densely many-
floweredand tomentose;filamentsonly slightlyexceedingcalyx lobes.
SystematicTreatment 87
55. Leaves 7-13 cm long, 3-5 cm broad,chartaceous;inflorescence2-
3-flowered. 87. H. pauciflora.
55. Leaves 3.5-5.5 cm long, 1.5-2.7 cm broad,thick-coriaceous;inflo-
rescencemany-flowered. 88. H. glaziovii.
54. Fertile stamens 4-6; inflorescencepuberulousto tomentellous,many-
flowered;filamentsfar exceedingcalyx lobes.
56. Leaves thick coriaceous;inflorescence4-15 cm long. 56. H. racemosa.
56. Leaves chartaceous;inflorescence2-4 cm long.
57. Leafapexbluntlyacuminateor acute;petioles0.5-1 mm; flow-
ers ca. 3 mm long. 55.1. H. arenosa.
57. Leafapex caudate;petioles 1.5-2.5 mm; flowersca. 6 mm long.
55.2. H. conduplicata.

Additional Notes and Descriptions in even at throat except around base of ovary;
of Species of Hirtella pedicels8-15 mm long, hirsute.Calyxlobes five,
lanceolate,hispid on exterior.Petals five, white,
7-4. Hirtella dorvalii Prance, Fl. Neotrop. glabrous.Stamens six, the filamentsfar exceed-
Monogr.9: 273. 1972. ing calyx lobes. Style hirsute on lower portion
only. Ovaryinsertedat mouth of receptacle,gla-
Distributionand habitat(Fig. 128). This little brous aroundbase. Fruit not seen.
known myrmecophilousspecies was described Distribution(Fig. 128) and phenology.Known
froma singlespecimenby Prance(1972). Further only from the Rio Nanay in Peru, collected in
collections, all from the same area, near Cara- flowerfrom November throughMarch.
carai, indicate that this is a locally abundant Habitat. Upland forest on white sand, poorly
species occurringonly in low caatingaforest on drained,swampy.
sandy soils.
Additional specimens examined. PERU. LORETO:
Additional specimens examined. BRAZIL. TERR. Maynas,Mishana,Rio NanaybetweenIquitosand
RORAIMA: RioAjarani, 28 May1974(fl),Pires& Caval- SantaMariadeNanay,13Nov 1977(fl),Gentry 20678
(NY),25 Feb1981(fl),Gentryet al 31725(MO,NY),
cante 14375 (IAN, INPA), 1 Jul 1974 (fl),Pires&Leite
14849(IAN,INPA);Caracarai, 27 Apr1974(fl),Pires 19 Mar1982(fl),Gentryet al. 36438(MO,NY).
& Cavalcante14360 (IAN, NY).
Hirtella revillae belongs to the section Myr-
mecophila,in which the swollen ant cavities at
7-6.1. Hirtella revillae Prance,Acta Amazonica the leaf bases, and hispid pubescenceare char-
8: 587, 89, fig. 6. 1979. Type. Peru. Loreto: acteristicof all members.It differsfromthe other
Maynas,Rio Nanay, 4 km from Mishana, 150 species in the section in the very long pedicels
m, 19 Jan 1975 (fl), Gentry, Ayala & Revilla and in the distinctive dense hirsute pubescence
15807 (holotype,NY; isotype, MO). of the pedicelsandflowers,andthe glabrousovary
and mouth of the receptacle.It is most closely
Trees8 m tall, the youngbrancheshispid.Leaf
relatedto H. physophora.
laminaoblong,chartaceous-membranous, 19-22
x 8-10 cm, rounded at base and bearing two The inflorescence is of rather intermediate
length in comparisonto other species of section
swollenant cavities, abruptlyacuminateat apex,
Myrmecophila, which have either much more
the acumen 7-10 mm long, hirsute on venation
compact fasciculateinflorescencesor elongatera-
beneath, with sparse appressedhairs on upper cemes or
panicles. Only Hirtella duckei has ra-
surface; midrib prominent beneath, prominu- cemose inflorescencesas short as H. revillaebut
lous above, hirsute on both surfaces;primary
H. duckeidiffersin many other ways.
veins 13-17 pairs, prominentbeneath, promin-
ulous above. Stipules linear, persistent, hispid.
7-9.1. HirtellaconfertifloraPrance,Brittonia33:
Inflorescencesof axillaryracemes 5-6 cm long, 354. 1981. Type. Venezuela.Amazonas:Rio
the rachislight brown hispid, the lower pedicels
Coro-Coro, 5?35'N,50?10'W,22 Feb 1979 (fl),
longerthan the upperones giving a slightlycor- et al. 117921 (holotype,NY; iso-
Steyermark
ymboseappearance.Bractsandbracteoleslinear, MO, VEN).
types,
persistent, hispid. Flowers 8-10 mm long (ex-
cludingpedicels).Receptaclecampanulate,light Tree4 m tall, the young branchesdensely fer-
brownhispid-hirsuteon exterior,glabrouswith- rugineoustomentose. Leaf lamina oblong-ellip-
88 Flora Neotropica

tic, coriaceous, 10.5-12 x 4.8-5.7 cm, cordate 7-11.1. Hirtella barnebyi Prance, Brittonia 33:
at base, shortlyand abruptlyacuminateat apex, 352. 1981. Type. Brazil. Rond6nia: Point 22
the acumen 1-3 mm long, glabrous and shiny of RADAM SC-20-XD, margin of Rio Preto
above, with sparseappressedhirsutepubescence nr.rapids,30 Aug 1975 (fl),M. dosR. Cordeiro
on lower surface; midrib prominulous above, 703 (holotype, INPA 57032; isotypes, IAN,
prominentandappressedbeneath;primaryveins INPA 54595). Fig. 122.
10-13 pairs,prominulousabove, prominentbe-
Shrub 2 m tall, the young branches sparsely
neath, secondaryvenation prominulouson both
hirsutulous,soon becomingglabrous.Leaf lam-
surfaces;petioles 1-3 mm long, terete, tomen- ina oblong, chartaceous,6-9.5 x 2.5-3.8 cm,
tellous. Stipules lanceolate, to 5 cm long, ca-
roundedat base, acuminateat apex, the acumen
ducous,eglandular.Inflorescencesof terminaland 9-14 mm long, glabrousbeneathexceptfor a few
axillary panicles with central rachis and short stiff appressedhairs on venation;primaryveins
densely clustered lateral branches bearing 2-4 8-11 pairs, prominulouson both surfaces;mid-
flowers, 12-15 cm long, the rachisand branches rib prominulousabove, prominentbeneath;pet-
tomentellous.Bractsandbracteoles1-3 mm long,
ioles 2-2.5 mm long, sparselytomentellous, te-
triangularto lanceolate,persistent,with numer-
ous stipitate tack-like glands. Flowers 5-7 mm rete, eglandular.Stipules minute, persistent,ca.
1 mm long, tomentellous. Inflorescences8-12
long. Receptaclecampanulate,tomentellousex- cm long,of little-branchedpanicleswith a central
ternally,glabrouswithin exceptfor reflexedhairs rachis and small lateral branches bearing 2-3
at throat;pedicels3-7 mm long. Calyxlobes five,
flowersonly, the rachisand branchesvery sparse-
acute, eglandular,tomentellous externally, pu-
berulous within. Petals five. Stamens six, uni- ly hirsutulous.Bracts and bracteoles 1-1.5 mm
long, lanceolate, persistent, with sessile glands
lateral, the filaments far exceeding calyx lobes. on some, and caducousbracteoleswith largeter-
Style sparselyhirsuteon lower fourth.Ovaryin- minal gland present only in bud. Flowers 6-7
serted at mouth of receptacle,pilose. Fruit not
mm long. Receptacle campanulate,with a few
seen.
hairson exterior,glabrouswith-
Distribution (Fig. 126). Known only from the sparseappressed
in exceptat throat;pedicels5-7 mm long, sparse-
type collection,fromgalleryforestborderingtree
savanna. ly appressedhirsute.Calyxlobes five, reflexedin
hirsutulous externally,
This species is closest to Hirtellaglabrata,but open flowers, acute,
the margins
differs in the shorter, more compact inflores- densely gray-tomentellouswithin,
eglandular. Petals five, glabrous. Stamens six,
cence, the oblong leaves, the distinctly cordate the filamentsglabrous,or with a few
leaf base, the shorterpetioles and the long linear unilateral,
hairson lower 10 mm, freeto base, farexceeding
stipules. H. confertiflorais also close to H. ad- lobes.
Style hirsute up to one third of length.
derleyi, but differsin the longer leaves that are insertednearmouth of receptacle,villous.
not conduplicate,in the compact inflorescence Ovary
Fruit not seen.
with shorter,weakerbranches,and in the longer
This is most closely related to Hirtella ara-
secretorytip of the bracts.
guariensiswhich has a similar subracemosein-
florescence with a central rachis and short
7-11. Hirtella araguariensis Prance, Fl. Neo- branches bearing2-3 flowers.This type of inflo-
trop. Monogr.9: 278. 1972. rescence occurs in only a few species such as H.
eriandraand H. elongata, which are otherwise
Distribution(Fig. 121). This species,described quite distinctfromH. barnebyi.This speciesdif-
from a type collection from Amapa, is much fers from H. araguariensisin the smaller, less
more widely distributed, ranging from north of lanceolateleaves with rounded,not subcordate,
Manaus, east to Amapa. bases, in the sparsely hirsutulousinflorescence
and flowers, and in the much smaller stipules
Additionalspecimensexamined. BRAZIL.AMAPA: and bracteoles.
PortoGrande,29 Nov 1976 (fl),Rosa 1052 (MG,NY).
AMAZONAS: Rio Uatuma,Mun. Itapiranga, Igarap6Ca-
tita, 21 Aug 1979 (fl), Cid et al. 640 (INPA, NY); Rio 7-11.2. HirtellamargaePrance,Proc.Kon. Ned.
Pitinga,27 Aug 1979 (fl), Cid et al. 860 (INPA, NY). Akad. Wetensch. Ser. C. 89: 111-113. 1986.
SystematicTreatment
Type. Suriname.BrokopondoDistr.: Browns-
bergNature Park,21 Jan 1978 (fl, fr), Roberts
LBB16303 (holotype, NY). Fig. 138.
Small trees to 3 m tall, the young branches
glabrous, becoming lenticellate with age. Leaf
lamina oblong to oblong-lanceolate,coriaceous,
21-27 x 6-9 cm, roundedto subcordateat base,
acuminateat apex, the acumen 1-5.5 cm long,
glabrousbeneathexcept for a few stiffappressed
hairs on venation;midrib prominulousand gla-
brous above; prominent and with sparse ap-
pressedpubescencebeneath;primaryveins 15-
18 pairs, prominulous above, prominent be-
neath;petioles 8-10 mm long, rugulose,with ap-
pressed pubescence of stiff long hairs, terete,
eglandular.Stipuleslinear-lanceolate,persistent,
7-9 mm long, eglandular,appressedpubescent.
Inflorescencesof little-branchedpanicles with a
central rachis and small groups of flowers on
short branches and solitary flowers;the rachis
darkbrown-tomentellous.Bractsand bracteoles
3-6 mm long, lanceolate, persistent, appressed
tomentellous,with few sessile glandson margins
of most bracteoles.Flowers 5-7 mm long, soli-
tary or in small clusters on central rachis. Re-
ceptaclecampanulate,appressedhirsutulouson
exterior, glabrous within except for pilose de-
flexedhairsaroundthroat.Calyxlobesfive,acute,
appressedhirsutulouson exterior, gray-tomen-
tellous within, eglandular.Petals five, pale pur-
ple, glabrous.Stamens six, unilateral;filaments
far exceedingcalyx lobes. Style pilose on lower
portion, glabrousabove. Ovary insertednear to
mouth of receptacle,pilose. Fruit ovoid, ca. 3
mm long, 1.8-2.5 cm broad, appressed-hirsu-
tulous on exterior;mesocarpthin, fleshy;endo-
carp thin, hard-bony,densely hirsute within.
Additional
specimenexamined.GUYANA.Puruni,
2 Apr 1953 (fr),J. Boyan 75 (FD 7749) (NY).
Local name. Guyana:bokobokotokon.
This species, closest to Hirtella araguariensis
is distinguishedby the larger,more coriaceous
leaves, the hirsutulousdark pubescence of the
inflorescence,and the less groupedflowersof the
inflorescence.
7-11.3. Hirtella liesneri Prance, sp. nov. Type.
Venezuela.Tachira:Rio San Buena, 10 km W Distribution(Fig. 135). This species,described
of La Fundaci6n,7?47'N,71046'W,13-15 Mar from just two collections, appears to be quite
89
1980 (fl),R. Liesneret al. 9633 (holotype,NY;
isotypes, MO, VEN). Figs. 19, 136.
SpeciesH. araguariensiaffinis,inflorescentiis
puberulisversus basim hispidis, bracteolisglan-
duligeris,foliisque chartaceisdiversa.
Tree7 m tall, the youngbranchessparselyhis-
pid, becoming glabrous with age. Leaf lamina
oblong, chartaceous, 10-15 x 3.5-5 cm, sub-
cordateat base, acuminateat apex, the acumen
10-15 mm long, the lower surface with sparse
appressedhairs, especially on venation; midrib
prominulous above, prominent and appressed
pubescentbeneath;primary veins 12-16 pairs,
prominulouson both surfaces;petioles ca. 2 mm
long,terete,sparselyhispidwhen young.Stipules
linear,persistent,ca. 3 mm long, eglandular.In-
florescencesofterminalandaxillarypanicles,with
a long centralrachisand very short 3-5 flowered
lateralbranches,giving a racemoseappearance,
20-22 cm long, the rachispuberulous,the lower
partsparselyhispid. Bracteolestriangular,mem-
branous,the apex terminatingin a stalkedgland.
Flowers6-7 mm long. Receptaclecampanulate,
sparselyhispid on exterior,glabrouswithin ex-
cept for ring of deflexedhairs at throat;pedicels
2-4 mm long. Calyxlobes five, acute,eglandular,
sparselyhirsute on exterior,puberulouswithin.
Petalsfive, glabrous.Stamenssix, unilateral,the
filamentsfar exceedingcalyx lobes. Style hirsute
on lower third. Ovary inserted at mouth of re-
ceptacle, pilose. Young fruit ellipsoid; exocarp
sparsely appressed pubescent, becoming gla-
brous with age.
Habitat. Primaryforestedarea on sandy soil,
700-1000 m.
This species is quite distinct in its inflores-
cence, and is one of the few hirtellaswith a short-
branchedelongatedpaniculateinflorescencethat
resemblesa raceme.It is probablyclosest to Hir-
tella araguariensisfrom which it differs in the
sparselyhispid to puberulous(not tomentellous)
inflorescence,the largerbranchesand pedicels,
the fewerglandson the bracteoles,and the larger
chartaceousleaves.
7-13. Hirtella insignis Briquet ex Prance, Fl.
Neotrop. Monogr.9: 279. 1972.
90 Flora Neotropica

G. 19. Hirtellaliesneri
(Liesneret a 9633). A, habit;B, bracteole;C, flowerand petal; , flowersection.

FIG. 19. Hirtella liesneri(Liesneret al. 9633). A, habit;B, bracteole;C, flowerand petal;D, flower section.
SystematicTreatment 91

common in Atlanticcoastal forestsof Bahia and teoles ovate, persistent,tomentose with numer-
EspiritoSanto. ous sessileor shortlystipitateglandsaroundmar-
Km
Additionalspecimensexamined.BRAZIL.BAHIA:
gins. Flowers 7-9 mm long. Receptacle
9, rd.PortoSeguroto Eunopolis,8 Feb 1972(fl),Eu- campanulate,tomentellouson exterior,glabrous
ponino212(NY);5 kmN ofComandatuba, SEof Una, within except for deflexed hairs at throat;pedi-
25 Jan 1977 (fl), Harley 18256 (K); rd. Porto Seguro cels 5-6 mm long, tomentose, eglandularexcept
20 May1971(fl),T.S. dos
to SantaCruzde Cabralia, on pairedbracteoles.Calyxlobes five, acute, to-
Santos 1677 (NY); Camaca,24 May 1971 (fl), T. S. mentose on exterior, densely tomentose on in-
dosSantos1695(NY);Km 10,rd.Valenqa
to Guabim,
22 Feb 1975 (fl), T. S. dos Santos 2898 (CEPEC,NY). terior.Petals five, glabrous.Stamenssix, unilat-
ESPiRITO SANTO:Reserva Florestal da CVRD, Lin- eral. Ovary inserted in middle of receptacle,
hares, 8 May 1979 (fl), Foli 76/79 (INPA). pilose. Style pilose at base glabrousabove. Fruit
not seen.
7-16. Hirtella davisii Sandwith,Bull. Misc. In- Distribution(Fig. 147). Known only from the
form. 1935: 125. 1935. Fig. 128. type collection.
This species, previously known from Vene- Habitat. Forest.
This species is close to the CaribbeanHirtella
zuela, Guyana,and Brazil,has recentlybeen col-
lected in Nicaraguaand is anotherexample of a americana but differs in the unbranchedinflo-
Guiana-CentralAmericadisjunction.Perhapsit rescence, the less impressed leaf venation, and
in many other minor characters.It is not easily
has been overlooked, because it is a large tree,
confusedwith other species of Hirtella.
30 m tall, unlike most other species of Hirtella
whichareshrubsand smallunderstorytrees.This
7-29. Hirtella adderleyi Prance, Fl. Neotrop.
species is noted as 30 m tall in both the Nicara-
guan and Guyanancollections. Monogr.9: 296. 1972. Fig. 120.
Additionalspecimens examined. NICARAGUA. RIO Representativeadditionalspecimensexamined.
SAN JUAN: Rio SantaCruz, 1101 'N, 84?24'W,22 Mar VENEZUELA. AMAZONAS: Depto. Atures, E of Sa-
1985(fl),Moreno25541(MO);Rio Sabalo,11?02'N, nariapo,Apr 1979 (fl),Davidseet al. 6753 (NY); Cas-
erio de 22
Canaripo, Aug1978 Huber2439
(fl), (NY,
84?28'W,23 Mar 1985 (fl), Moreno25614 (MO).
VEN); SE foot of Cerro Moriche, 19 Feb 1979 (fl),
Huber 3205 (NY, VEN); Rio Ventuari, 10 km E of
7-22.1. Hirtella santosii Prance,Revista Brasil. Carmelitas,20 Feb 1979 (fl),Huber3253 (NY); 22 km
Bot. 2: 34, fig. 5. 1979. Type. Brazil. Bahia: S of confluenceof Rios Manapiareand Ventuari,27
Una, FazendaSao Rafael, 10 Dec 1968 (fl), T. Feb 1979 (fl), Huber3469 (NY); Serraniadel Parfi,N
S. dos Santos 300 (holotype, CEPEC; isotype, of upperRio Pari, 2 Mar 1979 (fl),Huber3589 (NY);
4 km N of Rio Sipapo, 28 Jul 1980 (fl), Huber 5577
NY). (NY, VEN); E slopes of Cerro Calentura, 5?56'N,
65?40'W, 11 Apr 1974 (fl), Jangoux 10111 (NY).
Tree 8 m tall, the young branches tomentel-
lous, becoming glabrouswith age, not conspic-
uously lenticellate. Leaf lamina elliptic, coria- 7-30. HirtellapunctillataDucke,Arch.Jard.Bot.
ceous, 7-12.5 x 2-5.5 cm, roundedto subcuneate Rio de Janeiro3: 268. 1922; Prance,Fl. Neo-
at base,acuminateat apex, the acumen4-10 mm trop. Monogr.9: 296. 1972. Fig. 143.
long, hirsute beneath on venation; midrib pro- This rathercommon species of the Guayana
minulousabove but in impressedchannel,prom- Highlandand the sandy caatingasof the upper
inent and denselyhirsutebeneath;primaryveins Rio Negro is interestingbecause in several col-
15-19 pairs, prominulous above but in im- lections (for example Huber 10313 and 10875)
pressedchannel, prominentand densely hirsute most of the leaves are opposite. It is the only
beneath;secondaryvenation slightly impressed species of Chrysobalanaceaeknown to have op-
above givinglightlybullateappearance,promin- posite leaves.
ulous beneath;petioles 3-4 mm long, terete, to-
mentellous,eglandular.Stipuleslinear, 4-5 mm 7-31. Hirtella corymbosaChamisso & Schlech-
long, subpersistent,tomentellous.Inflorescences tendal, Linnaea2: 545. 1827. Fig. 127.
of terminalracemesto 6-10 cm long, the rachis
and branchesdensely rufous-tomentose.Bracts Habitat. Growing on open restingawith sa-
small, persistent, lanceolate, tomentose; brac- vannas.
92 FloraNeotropica
Additional specimen examined. BRAZIL. BAHIA: pairs, prominent beneath, prominulous above;
Mun.de Mucuri,7 km NW of Mucuri,14 Sep 1978 midrib prominentbeneath,prominulousabove,
(fl), Mori et al. 10470 (CEPEC,NY). tomentellouson both surfaces;petioles 5-9 mm
long, 3.5-6 mm thick, tomentellous,eglandular,
7-34. Hirtella leonotis Pittier, Arb. Arbust.Ve- terete.
Stipules early caducous(not seen). Inflo-
nez. 2: 23. 1923. rescencesterminal panicles with a long central
Fruitovoid, ca. 2 cm in diam.;exocarpdensely rachis 12-18 cm long and many short few-flow-
tomentellous. ered lateral primary branches, the rachis and
Distribution (Fig. 136). Forests of northern branches rufous-tomentellous.Bracts and brac-
Colombia and Venezuela;in Venezuela only in teoles ovate, persistent, gray-brown-tomentel-
the FederalDistrict and State of Miranda. lous on both surfaces,eglandular.Flowers 5-6
mm long. Receptaclecampanulate,tomentellous
Additionalspecimensexamined. COLOMBIA. on
exterior,glabrouswithin except for sparsely
ANTIOQUIA: BuenosAires forestabove rd. to Anori, 26
Apr 1973 (fl), Soejarto et al. 4025 (MO); Autopista pilose
areaaroundthroat;pedicels 1-2 mm long,
Medellin-Bogoti, Setor Rio Samara-Rio Claro,400- tomentellous. Calyx lobes five, acute, gray-to-
1000 m, 19 Mar 1982 (fl), Herndndez& Albertde E. mentellous on both surfaces.Petals five, white,
244 (HUA). glabrous.Stamens 5-7, unilateralwith toothed
VENEZUELA.MIRANDA:ParqueNacional de Gua-
of to them, filaments far
topo (fr), Aristeguieta7096 (VEN); Cerros del Ba- portion ring opposite
chiller,25 Mar 1978(st),Steyermark&Davidse116868 exceedingcalyx lobes. Ovary inserted at mouth
(MO, NY, VEN), 25 Mar 1978 (fl, fr),Steyermark& of receptacle, pilose. Style glabrous. Fruit not
Davidse116933 (MO, NY, VEN). seen.
This species was known only from the type in Distribution(Fig. 138). Known only from Lo-
1972. Recent collections from Cerros de Ba- reto, Peru, and adjacentBrazil.
chiller and Guatopo show that it is a most dis- Habitat. Upland forest on terrafirme, under-
tinct species. The fruit can now be described.It story in open clearings.
is one of the few species of Hirtella that has a Additional specimens examined. PERU. LORETO:
densely tomentose exocarp. Prov.Requena,Arboretum JenaroHerrera,Jul-Sep
1976(fl),Bernardi1-56 (G, NY), 15 Nov 1974(fl),
7-37. Hirtella bahiensis Prance, Fl. Neotrop. Diaz s.n. (G), 7 Dec 1977 (st), Gentryet al. 21213
(MO, NY).
Monogr. 9: 307. 1972. Fig. 122.
Hirtella magnifolia has the largest leaves of
Additionalspecimensexamined.BRAZIL.BAHIA:Nr.
SantaCruzde Cabralia,21 Oct 1978(fl),Mori10936 any describedspecies of the genus, often attain-
(CEPEC,NY);Ilh6usRd. to Olivenga,12 Nov 1970 ing 40 cm in length on the fertilebranches.It is
(fl), T. S. dos Santos 1284 (NY). ESPIRITOSANTO: Re- most closely related to H. elongata and H. er-
serva FlorestalCVRD, Linhares,24 Nov 1978 (fl), I. iandra,but differsfrom both in the largerleaves
A. Silva 30 (INPA). with a greaternumberof primaryveins. It differs
furtherfrom H. elongata in the rufous-tomen-
7-40.1 Hirtella magnifolia Prance, Acta Ama- tellous pubescenceof the inflorescence,the two
zonica 8: 585, 587, fig. 5. 1979. Type. Brazil.
glandsat the junction of the uppersurfaceof the
Amazonas: Rio Javari, Estirao do Equador, leaf lamina and the petioles, the rounded not
21 Oct 1976 (fl), Prance et al. 23974 (holotype, subcordateleaf bases, and the longer inflores-
INPA; isotypes, FHO, MG, MO, NY, US). cence branches;and furtherfrom H. eriandrain
Tree to 10 m tall, the young branchesshortly the inflorescence branching and the laminar
tomentellous becoming glabrousand conspicu- glands.
ously lenticellatewith age. Leaf lamina oblong-
elliptic, chartaceous,18-40 x 8-16 cm, the base 7-48. Hirtella rasa Standley, Publ. Field Mus.
rounded, abruptlyacuminate at apex, the acu- Nat. Hist., Bot. Ser. 17: 252. 1937. Fig. 146.
men 7-13 mm long,curved,glabrousabove, with
a few stiff appressedhairs beneath on venation, Additionalspecimenexamined.PERU. LORETO:Pu-
callpa,km 75, 9 Sep 1980 (fl), Angelo 11 (MO, NY).
with two glands at junction of upper surfaceof
lamina and the petioles; primary veins 14-18 Local name. lobo apacharama.
SystematicTreatment
7-52.1. Hirtella radamii Prance, Acta Amaz6-
nica 12: 22. 1983. Type. Brazil.Rond6nia:Iga-
rape Preto, RADAM SC-20-XA-Ponto 27,
62?14'W,8?58'S,30 Jun 1975 (fl),J. C. daSilva
100 (holotype, MG). Fig. 143.
Shrub3 m tall, the youngbrancheshispid.Leaf
lamina oblong-lanceolate,chartaceous, 5-8 x
1.5-2.3 cm, subcordate at base, acuminate at
apex, the acumen3-5 mm long,glabrouson both
surfaces;midrib prominulousabove, prominent
beneath; primary veins 10-12 pairs, prominu-
lous and glabrouson both surfaces;petioles 1-
1.5 mm long terete, sparselyhispid, eglandular.
Stipuleslanceolate,to 5 mm long, membranous,
caducous, with numerous long-stipitateglands.
Inflorescencesof terminalracemesto 12 cm long,
the rachis glabrous.Bractsand bracteoles 1.5-3
mm long, oblong,with two sessileglandstoward
apex, a few of the youngestbracteoleswith one
or two stipitateglands in addition to the sessile
pairedglands.Flowersca. 5 mm long. Receptacle
campanulate, glabrous on exterior, glabrous
within except for deflexed hairs at throat;pedi-
cels 12-16 mm long, extremely slender, eglan-
dular. Calyx lobes five, acute, glabrouson exte-
rior, puberulouswithin, eglandular.Petals five,
glabrous. Stamens six, unilateral,the filaments
farexceedingthe calyx lobes, glabrous.Style hir-
sute for half of length. Ovary insertedat mouth
of receptacle,pilose. Fruit not seen.
Local name. caripe torrado.
This species is closest to Hirtella tenuifolia,
from which it differs in the smaller leaves, the
hispid pubescence of the young stem, and the
muchlongerpedicels.It differsfromH. racemosa
in the narrow,oblong-lanceolateleaves, the long
pedicels and the hispid young stems. It differs
from H. sprucei in the smaller leaves, the long
pedicels, the bracteolarglands and the absence
of pedicel glands.
7-55.1. Hirtella arenosa Prance, Brittonia 28:
227, fig. 11. 1976. Type. Brazil. Amazonas:
Manaus-ItacoatiaraRd., km 180, 7 Jul 1980
(fl), W. A. Rodrigues et al. 8257 (holotype,
INPA). Fig. 122.
Shrubto 3 m tall, the young branchespuber-
ulous, soon becoming glabrous. Leaf lamina
oblong-elliptic,chartaceous,3-5.5 x 1.2-2 cm,
cuneate at base, acute, rounded, or shortly
93
acuminate at apex, the acumen 0-4 mm long,
glabrous above, glabrous beneath except for
appressedpubescenceon midrib;midrib prom-
inulousabove,prominentbeneath;primaryveins
8-10 pairs prominulouson both surfaces;peti-
oles 0.5-1 mm long, terete,eglandular,glabrous
or with a few adpressed hairs. Stipules 2 mm
long, linear, subpersistent. Inflorescencester-
minal and subterminalracemes, 3-4 cm long,
the rachis sparselyhispid. Bractsand bracteoles
narrowlytriangular,persistent,eglandular,ca. 1
mm long, sparselyhispid on exterior.Flowers 3
mm long. Receptaclecampanulate,sparselyhis-
pid on exterior, almost glabrous within except
for a few appressed hairs with a ring of long
deflexedhairsaroundbase of staminalring;ped-
icels 5-7 mm long, eglandular,sparselyhispid.
Calyx lobes five, acute, reflexed,sparselyhispid
on exterior,denselygray-puberulouson interior.
Petals five, purple,glabrous.Stamens 5-6, uni-
lateral, far exserted. Ovary tomentose, inserted
at mouth of receptacle.Style pilose. Fruit un-
known.
Habitat and phenology.This speciesis known
from two white sand campinasof CentralAma-
zonia. Collectedin flowerin July.
Additional specimens examined. BRAZIL.
AMAZONAS:Manacapuru,Igarap6Branco,6 Jul 1958
(fl),Ferreira58-320(INPA,NY);Serrados 6 Lagos,
18Jul 1979(fl),Maiaet al. 663 (INPA).
7-55.2. Hirtella conduplicataPrance, Brittonia
28: 227, fig. 12. 1976. Type. Brazil.Amazonas:
Lagode Castanho-Mirim,25 Jun 1973 (fl), B.
Albuquerqueet al. 887 (holotype, INPA).
Fig. 125.
Treeto 20 m tall, the youngbranchessparsely
hispid. Leaf lamina conduplicate,oblong-ellip-
tic, chartaceous,3.5-5.5 x 2-2.5 cm, subcordate
at base, caudateat apex, the acumen 8-15 mm
long, glabrousabove, glabrousor with sparseap-
pressed pubescence on veins beneath, midrib
prominulouson both surfaces;primaryveins 5-
7 pairs, prominulouson both surfaces;petioles
1.5-2.5 mm long,terete,eglandular,sparselyhis-
pid. Stipules ca. 1 mm long, caducous, hispid.
Inflorescencesof terminal and axillaryracemes
2-3.5 cm long, the rachissparselyhispid. Bracts
and bracteoles ca. 1 mm long, oblong, acute,
eglandular,sparsely hirsute. Flowers 5-6 mm
long. Receptacle campanulate,sparsely hirsute
94 FloraNeotropica
on exterior,glabrouswithin except for circle of Amazonia. Although it is quite distinct by its
deflexedpilose hairs aroundstaminalring;ped- long, bracteateinflorescence,it is little collected
icels 6-10 mm long, eglandular,sparselyhirsute, and apparentlyrare. One recent collection has
thick.Calyxlobes five, rounded,reflexed,sparse- been made.
ly hirsute on exterior, densely gray-puberulous Additionalspecimenstudied:BRAZIL.PARA:Mun.
within. Petals five, glabrous,white. Stamenssix, de Itaituba,Santar6m-CuiabaHwy., km 780, Serrado
unilateral,far exserted,purple,connate and hir- Cachimbo,9?22'S,54?54'W,29 Apr 1983 (fl),Amaral
sute towards base. Ovary pilose, inserted near et al. 1041 (INPA, NY).
mouth of receptacle. Style hirsute. Fruit un-
known. 7-61. Hirtella lemsii L. O. Williams & Prance,
Habitat and phenology.This species was col- Fl. Neotrop. Monogr. 9: 334. 1972. Fig. 136.
lected only at the type locality in rain forest on
terrafirme, floweringin June. This species, described from a single collection
from Costa Rica, appears to be quite common
Additionalspecimenexamined.BRAZIL.AMAZONAS: in that
25 Jun 1973(fl),Albuquer- country, and the fruit is described for the
Lagode Castanho-Mirim, first time.
queet al. 886 (INPA,NY).
Fruit oblong, ridged, and with two small pro-
7-56d. Hirtella racemosa Lam. var. hispida trusions when dry, slightly tapered towards base,
Proc. Kon. Ned. Akad. 2.5-3 cm long; exocarp sparsely tomentose when
Prance, Wetenschapp.
Ser. C. 89: 113. 1986. Type. Brazil. Amapa: young; mesocarp thin and fleshy; endocarp hard,
Rio Falsino, 10 km above Rio Araguari, 26 thin.
Aug 1983 (fl), B. V. Rabelo et al. 2381 (ho- Additional specimens examined. COSTA RICA.
lotype, MG; isotype, NY). Fig. 146. HEREDIA:Rio PuertoViejo, 2 km above Rio Sarapiqui,
14 Jun 1968 (fl), Burger& Stolze 5784 (F); Finca La
The young brancheshispid. Leaf lamina 15- Selva, Puerto Viejo, 17 Sep 1969 (fl), Frankie 398C
22 x 6-7.5 cm; pedicels thin, 8-12 mm long. (F), 7 Mar 1970 (fl), Hartshorn809 (NY), Sep 1974
Habitat. Variety hispida is a small shrub of (fl), Hartshorn1523 (NY), 13 Aug 1961 (fl), Rosbach
3719 (GH). PUNTARENAS: Ridge betweenBanegasand
1.5 m of the understoryof forest on terrafirme. Rio Riyito, 7 km W of Rinc6n de Osa, 8 Oct 1984 (fr),
Additional specimens examined. FRENCH Grayum4090 (MO,NY); Lloronatrailto SanPedrillo,
GUIANA.HautCriqueBaboune,2 Aug 1981(fl),de CorcovadoNational Park,21 Jul 1977 (fr),Hartshorn
Granville
4723 (NY). 1878 (NY).
BRAZIL.AMAZONAS: Mun. de Axinim, lower Rio
Paca,4?07'S,58?58'W,1 Jul 1983(fl),Zarucchiet al. 7-67. Hirtella mucronata Prance, Fl.
2920 (INPA,NY). AMAPA: Rio Oiapoque,Mt. Alikene, Neotrop.
nr.Riv.Camopi,30 Sep1960(fl),Irwin48589a(NY). Monogr. 9: 339. 1972.
This variety was described to accommodate Distribution (Fig. 139). This species was based
specimensof Hirtellaracemosathat aredistinct- on a type from southern Guyana and two col-
ly hispid on the young stems. It has the large lections from the Rio Negro region of Brazil.
leaves characteristicof var. racemosa but the New material adds to this range.
longthin pedicelscharacteristicof var. hexandra. Additionalspecimensexamined.SURINAME.Lely
The latter characteris almost constant to sepa-
Mountains, 175 km SSE of Paramaribo,12 Oct 1976
rate the two varieties,althougha few collections (fl), Mori & Bolten 8459 (NY).
placed in var. racemosahave long thin pedicels BRAZIL. AMAZONAS:Manaus-Itacoatiara Rd., km
(e.g., Chagas 1252, INPA, NY). The above ma- 185, 15 Dec 1966 (fl), Pranceet al. 3653 (MG); Ma-
terial is obviously closest to H. racemosaand is naus-CaracaraiRd., km 220, Santo Ant6nio de Abo-
nari, 24 Nov 1976 (fl), Prance et al. 24257 (INPA,
best regardedas a varietyuntil furthercollections NY).
and field studies are made.
7-70.1. Hirtella parviunguis Prance, Revista
7-58. Hirtella kuhlmanniiPilger, Notizbl. Bot.
Brasil. Bot. 2: 34, fig. 4. 1979. Type. Brazil.
Gart. Berlin-Dahlem8: 538. 1923. Fig. 135.
Espirito Santo: Linhares, estrada da Povoa-ao
This specieswasknownin the monographfrom ao Linhares, 30 Mar 1971 (fl), T. S. dos Santos
the type and one other collection from southern 1512 (holotype, CEPEC; isotypes, FHO, NY).
SystematicTreatment 95

Treeto 12 m tall, the youngbranchessparsely 7-75. Hirtella excelsa Standley ex Prance, Fl.
appressed puberulous,glabrescent,lenticellate. Neotrop. Monogr.9: 345. 1972. Fig. 130.
Leaf lamina narrowlyoblong,coriaceous,4-7 x Additionalspecimensexamined.PERU. HUANUCO:
1.8-2.5 cm, cuneate to subcuneateat base, acu- Prov. Pachitea,BosqueNacionalde Iparia,Rio Pachi-
minate at apex, with acumen 2-5 mm long, gla- tea, 8 Aug 1967 (fr), Schunke V. 2137 (NY); Dist.
brousbeneath;primaryveins 10-14 pairs,prom- Honoria,CarreteraMielde Abejas,km 1.5, 8 Aug 1967
inulous on both surfaces;midrib prominulous (fr), Schunke V. 10 (F). SANMARTIN: Santa Margarita,
W of Nueva Aspusana, 6 Aug 1962 (fr), Mathias &
above, prominentbeneath,venationpapilloseon Taylor6108 (F).
upper surface; petioles 3-4 mm long, terete, BRAZIL.ACRE: Between Porangabaand Papagaio,
eglandular,sparselypuberulouswhen young,ru- Rio Jurua-Mirim,18 May 1981(fl),Maasetal. P13134
gulose. Stipules lanceolate 2-4 mm long, pub- (INPA, NY). MATOGROSSO:
Rio Tucunazinho, BR 174,
km 330, 8 Jun 1979 (fr),M. G. Silva & Rosdrio4826
erulouswhenyoung, subpersistent,adnateto ex-
(MG), 10 Jun 1979 (fr), M. G. Silva & Rosdrio4845
treme base of petiole, eglandular.Inflorescences (MG, NY).
of terminal and axillary densely crowded ra-
cemes 3-5.5 cm long, the rachissparselytomen- Local name. Peru: apacharama amarillo.
tellous. Bractsand bracteolesovate, 1.5-2.5 mm
7-79. Hirtella couepiiflora Prance, Fl. Neotrop.
long, persistentmembranous,eglandular.Flow-
ers ca. 7 mm long. Receptacle campanulate, Monogr. 9: 350. 1972. Fig. 127.
sparselypuberulouson exterior,glabrouswithin Additional specimens examined. FRENCH
except for deflexedhairs at throat;pedicels 3-6 GUIANA.Riv. Oyapock,PetitToucouchi,LagonTrois
mm long. Calyx lobes five, acute, sparselypu- Pitons, 6 Aug 1969 (fl), OldemanT447 (CAY).
BRAZIL. AMAPA:Clevelandia, Aug 1960 (st), Pires
berulous on exterior, tomentellous within, the 7700 (IAN).
margins eglandular.Petals five, glabrous, with
minute claw at base. Stamens seven, unilateral, Local name. Fr. Guiana: gaulette.
the filaments glabrous, far exceeding the calyx
lobes. Stylepilose, with a few hairson lowerhalf. 7-80. Hirtella tubiflora Cuatrecasas, Fieldiana,
Ovary inserted at mouth of receptacle,pilose. Bot. 27: 59. 1950.
Fruit not seen. Distribution (Fig. 150). This species was known
Distribution(Fig. 141). Atlantic coastal forest
only from the type collection from the coastal
and cacao plantations. lowlands of Valle in Colombia at 30-50 m, in
four collections
Additionalspecimenexamined. BRAZIL.ESPiRITO the Rio Calima region. Recently,
SANTO: CVRD Linhares,10 Jul 1979 (fl), Foli 80 have been made from the wet forests of Panama
(INPA). at about 800 m altitude, showing an interesting
distribution pattern.
Local name. macucurana.
Hirtellaparviunguisis distinct from all related Additionalspecimensexamined.PANAMA.PANAMA:
species by the small slightly clawed petals. It is CerroJefe, 26 Sep 1975 (fl), J. T. & F. Witherspoon
most closely relatedto H. martiana, a Planalto 8543 (MO, NY), 15 Feb 1982 (fl), Knapp3525 (MO,
NY). SANBLAS:El Llano-Carti Rd., km 19.1, 11 Mar
species, but, it differsnot only in the petals but 1985 (fl), Nevers & Herrera 5106 (MO, NY). VERAGUAS:
also in the eglandularbracteolesand calyx lobes, 3-4 km W of Santa F6, 2500' (fl), Nee 11288 (MO,
and the less pubescentinflorescenceand style. It NY).
COLOMBIA. CHOCO:Colombia-Panama frontier,
differsfromH. angustifolia,anothereasternBra- 24 Sep 1979 (fl), Barbosa1213 (COL).
zilian species, in the inflorescencesof densely
crowdedshortracemes,the subcuneateleaf bas- 7-87. Hirtella pauciflora Little, J. Wash. Acad.
es, and the puberulous (not hispid), young Sci. 38: 88. 1948. Fig. 141.
branches and inflorescences.This species also
Additionalspecimensexamined. ECUADOR. LOS
resemblesH. glaziovii from the same region,but RIOS: Jaunecheforest,CantonVinces, 1 Oct 1979 (fl),
differsin the petals, the greaternumber of sta- Dodsonet al. 8655 (NY);Rio PalenqueBiologicalSta-
mens, the much less tomentoseinflorescenceand tion, km 56, Quevedoto SantoDomingo, 6 Mar 1974
(fl), Dodson 5464 (US).
flowers,the largerflowers,and the more crowded
inflorescences. Local name. coquito.
96 FloraNeotropica

8. AcioaAublet

Additional Notes and Descriptions Tree25 m tall, the trunkslightlybuttressedto

of Species of Acioa 0.5 m, the youngbranchesglabrous.Leaf lamina
oblong, thickly coriaceous, 7-17 x 4.5-12 cm,
Distribution of Acioa as a whole is seen in roundedat base, acuminateat apex, the acumen
Figure 153. 2-6 mm long, glabrous on both surfaces, with
two glandsnear to base of lower surface;midrib
8-1. Acioa guianensisAublet, Hist. pl. Guiane prominulous above, prominent beneath; pri-
2: 698, t. 280. 1775.
mary veins 9-11 pairs, prominenton both sur-
Acioa remains the most poorly known genus faces; secondary venation prominent on both
of neotropicalChrysobalanaceae.Very few col- surfaces.Stipules linear, membranous,5-8 mm
lections have been made of this apparentlyvery long, glabrous,caducous. Inflorescencesmuch-
rare species. branched,slightlycorymbosepanicles, 5-10 cm
A recent collection of A. guianensis from the long, the rachis and branchesglabrous.Recep-
Municipalityof Manicoreon the MadeiraRiver tacle conical and slightly curved near base, 6-7
is the firstreliablecollection from CentralAma- mm long, glabrouson exterior,the interiorlined
zonia and confirmsthe likelihoodthat seeds col- by an extremely thick disc with only a small
lectedby Ducke and grownin the BotanicalGar- hollow, the interior with pilose hairs below in-
den of Rio de Janeiro are really from the Rio sertion of style, glabrous on other side. Calyx
Purus region. All previous collections of this lobes five, rounded,unequal,3-5 mm long, gla-
species had been made in French Guiana and brous with two glands on exterior, the margins
adjacentAmapa. See Figure 154. ciliate, appressed-puberulouson interior.Petals
five, white, caducous,glabrous,the marginscil-
Additionalspecimenexamined.BRAZIL.AMAZONAS: iate. Stamens
Mun. of Manicore:RADAMSB-20-XA,Point 9, 17-20, insertedin two rowsaround
60?53'W,6?50'S(fl), C. D. A. Mota s.n. (INPA 61659).
half of the thick staminal ring, the other half of
the ringwith staminodes,with a circleofdeflexed
8-3. Acioa schultesii Maguire,Brittonia7: 272. hairsinsertedon interiorat base of staminalring,
1951. the filaments glabrous, the anthers dorsifixed.
Ovary inserted at mouth of receptacle on the
This specieswasknownuntilrecentlyonlyfrom same side as stamens,glabrouson exterior,gla-
the type collectionfromthe Rio Dimite in Brazil. brous within, unilocular;ovules two. Style in-
It has now been collected from the regionof San sertedat base of ovarytowardsinterior,the swol-
Carlos de Rio Negro in Venezuela, where it is len base pilose, the filamentousportionglabrous,
apparentlyquite common, occurringfrequently equalling stamens in length. Fruit ellipsoid, 6-
in the IVIC ecological study sites in the region 7.5 cm long, 4-5 cm broad, exocarp glabrous,
(see Fig. 154). It is a tree 20-40 m tall, flowering lenticellate; mesocarp 12-14 mm thick; endo-
in June-Augustand known locally as pasista. carp thin and bony, fragile,glabrouswithin, the
cotyledons completely filling the central cavity.
Additionalspecimensexamined.VENEZUELA. Germinationepigeal, firstleaves opposite.
AMAZONAS: 4.3 km NE of SanCarlosde Rio Negro,
19 Jun 1978 Clark& Distribution.Mainly in the basin of the Rio
1?56'N,67?3'W, (fl), Maquirino
6661 (FHO, NY), 20 Jul 1978 (fl), Clark& Maquirino Purus and the central Solimoes.
6737 (FHO, NY), 3 Aug 1978 (fl), Clark& Maquirino Habitat. Forest on terra firme.
6740 (FHO, NY), 25 Jul 1978 (fl), Clark& Maquirino
6753 (fl) (NY), 25 Aug 1981 (fl), Clark8211(NY), 5 Additional specimens examined. BRAZIL.
km N of Boca de Casiquiare,1?57'N,67008'W,5 Feb AMAZONAS:Coari, 11 May 1974 (fr),CampbellP21130
1980 (fr), Liesner& Clark 9107 (MO, NY). (INPA, NY); 10 Feb 1975 (fl), Ramos P23251 (FHO,
INPA, MG, NY); Lago de Tefe, 9 May 1974 (st),
P21129 (INPA, NY), 20 Nov 1959 (fr),Ro-
8-4. Acioa edulis Prance, Acta Amaz6nica 2(1): Campbell
drigues& L. Coelho 1408 (INPA);Rio Ituxi, Boca do
12-16. 1971. Couepia edulis (Prance) Prance, Remanzinho,1933 (fl bud), Krukoff5822A(NY); Ma-
Acta Amazonica 5: 143. 1975. nua,Lagode Tefe, 13 Jul 1973 (fr),Lleraset al. P16655
Systematic Treatment
(INPA, NY); Santo Ant6nio de Iga, 19 Aug 1973 (fr),
Lleras et al. P17397 (INPA, NY).
Local name. castanhade cotia.
Uses. The fleshycotyledonsare edible and the
fruitsaregatheredin largenumbersby Brazilians
in the Rio Ituxi region. The kernel is eaten raw
or is crushed and added to their tapioca cakes
(beiju).Its oil is also extractedand used for cook-
ing and soap making.
The fact that I have alreadyplaced this most
distinctive new species in two genera indicates
9. Neocarya Prance
9-1. Neocarya macrophylla(Sabine) Prance in
F. White, Bull. Jard.Bot. Etat. 46: 308. 1976.
The eight genera treated above are those in-
digenousto the neotropics.The monotypicWest
Africangenus Neocaryais grownat the Summit Specimen examined. PANAMA. CANALAREA:Sum-
BotanicalGarden in Panama and so it is men- mit Garden, 6 May 1986 (fl), de Nevers et al. 7726
tioned brieflyhere. Neocaryais describedin full (MO, NY).
in Prance and White (1988), and is a segregate
ACKNOWLEDGMENTS
I am especiallygratefulto Mrs. Edith Topfler
for the preparationof most of the distribution
maps and for many hours of painstakingvol-
unteer work for this project. I also particularly
thank Carol Gracie for the preparationof the
final version of the maps, and Bobbi Angell for
drawings of the new species. I am grateful to
RosemaryLawlorand MickeyMaroncellifor the
typing and word processingof various drafts of
this manuscript and to H. David Hammond,
William R. Anderson, and two anonymous re-
viewers for reviewing an earlier draft. Most of
my fieldwork,which has been essential for an
understandingfor the family,has been supported
by a series of grants from the National Science
Foundation,latelyby grantBSR-8409536 which
is gratefullyacknowledged.I am gratefulto col-
laboratorsin many neotropicalherbariaand bo-
tanicalinstitutions,especiallyto the directorsand
staffs of the Instituto Nacional de Pesquisas da
Amaz6nia(INPA) in Manaus,Brazil,and of the
Museu ParaenseEmilio Goeldi (MG) in Belem,
Brazil.I am especiallygratefulto David Johnson
for much editorialassistance.
97
its problematicnature.It is quite unlikeany oth-
ers. It resemblesAcioa vegetativelyand in many
floralfeatures,but does not have what was pre-
viouslyconsideredto be the central,unitingchar-
acterof the genus:the stamensfusedinto a ligule.
A recentreview of the genera(Prance& White,
1988)showedthatit is best placedinAcioa rather
than Couepiaeven though it does not have the
fused stamens. This species forms a link with
Couepia,a genus in which fused stamens never
occur.
from the closely relatedgenusParinari.It differs
in the largegibbousreceptacle,the greaternum-
ber of stamensand the very differentfruit struc-
ture.
LITERATURECITED
Berlin, B. & G. T. Prance. 1978. Insect galls and
human ornamentation.The ethnobotanicalsig-
nificanceof a new species of Licania from Ama-
zonas, Peru. Biotropica10: 81-86.
Demchenko,N. I. 1973. The pollen morphologyof
the familyChrysobalanaceae. Pages69-73 in Pol-
len and sporemorphologyof recentplants(in Rus-
sian). Proc. 3rd Int. Palynol. Conf., Acad. Sci.
USSR.
Espinal T., S. 1981. El frbol raro de Comfama en
Rionegro.FloraAntioquenia1: 1-3.
Goulding,M. 1980. The fishes and the forest: Ex-
plorationsin Amazoniannaturalhistory.Univer-
sity of CaliforniaPress, Berkeley.
Letouzey,R. & F. White. 1976. Chrysobalanacees
nouvellesdu Camerounet du Gabon.Adansonia,
Ser. 2, 16: 229-243.
-- & . 1978a. Chrysobalanacees.In Flore
du Cameroun20: 1-128, 237-247. MuseumNa-
tional d'HistoireNaturelle,Paris.
& . 1978b. Chrysobalanacees. In Flore
du Gabon 24: 138, 194-201. Museum National
d'HistoireNaturelle,Paris.
Mori, S. A., B. M. Boom& G. T. Prance. 1981. Dis-
tributionpatternsandconservationof easternBra-
zilian coast foresttree species. Brittonia33: 233-
245.
Patel, V., J. J. Skvarla& P. H. Raven. 1983. Pollen
98 FloraNeotropica
ultrastructureof Chrysobalanaceae.Vidya 26: 1- tropicalspecies with relationto history,dispersal
10. and ecology,with specialreferenceto Chrysobala-
Prance,G. T. 1968. Maranthes(Chrysobalanaceae), naceae, Caryocaraceaeand Lecythidaceae.Pages
a new generic record for America. Brittonia 20: 59-87 in K. Larsen& L. B. Holm-Nielsen (eds.),
203-204. Tropicalbotany. AcademicPress, New York.
. 1970. The generaof Chrysobalanaceae in the . 1981. Notes on Couepiaand Hirtella(Chrys-
southeasternUnited States. J. Arold Arbor. 51: obalanaceae).Brittonia33: 347-356.
521-528. . 1982a. Forestrefuges:Evidencesfromwoody
. 1972. Monographof Chrysobalanaceae.Fl. angiosperms.Pages 137-151 in G. T. Prance(ed.),
Neotrop. Monogr.9: 1-406. Biologicaldiversificationin the tropics.Columbia
. 1973. New and interestingChrysobalanaceae University Press,New York.
from Amazonia. Acta Amaz6nica2(1): 7-16. . 1982b. Chrysobalanaceae.Flora de Vene-
. 1974a. Supplementarystudies of American zuela 14(2): 325-487.
Chrysobalanaceae.Acta Amaz6nica4(1): 17-23. 1984. New taxa of AmazonianChrysobala-
1974b. A new Peruvian species of chirop- naceae.Acta Amaz6nica13: 21-30.
terophilous Couepia (Chrysobalanaceae).Britto- 1986a. Studies on the flora of the Guianas.
nia 26: 302-304. 19:New taxa of Chrysobalanaceae for the floraof
1974c. A note on Couepiacognata (Steud.) the Guianas. Proc. Kon. Ned. Akad. Wetensch.,
Fritsch and related species (Chrysobalanaceae). Ser. C. 89: 111-116.
Acta Bot. Venez. 9: 119-122. . 986b. Flora of the GuianasSeries A: Pha-
1974d. Phytogeographicsupportfor the the- nerogams.Family 85. Chrysobalanaceae.Koeltz
ory of Pleistocene forest refugesin the Amazon ScientificBooks,Koenigstein,FederalRepublicof
Basin based on evidence for distributionpatterns Germany.
in Caryocaraceae,Chrysobalanaceae,Dichapeta- . 1987. Notulae de ChrysobalanaceisMale-
laceae and Lecythidaceae.Acta Amaz6nica 3(3): sianis Praecursoriae.Brittonia39: 364-370.
5-28. . In press. Chrysobalanaceae.In C. Kalkman
1975. The correctname for castanhade cutia (ed.), Flora Malesiana.
(Couepiaedulis (Prance)Prance-Chrysobalana- - & A. R. A. Gortsvan Rijn. 1976. Chrysobal-
ceae). Acta Amaz6nica5(2): 39-41. anaceae. Pages 524-555 in J. Lanjouw & A. L.
. 1976. Additions to neotropicalChrysobal- Stoffers.Additionsand correctionsto the Floraof
anaceae.Brittonia28: 209-230. Suriname2(2).
.1977a. Two new speciesfor the floraof Pan- -& S. A. Mori. 1983. Dispersaland distribu-
ama. Brittonia29: 154-158. tion of Lecythidaceaeand Chrysobalanaceae. Son-
. 977b. The phytogeographicsubdivisionsof derb. Naturwiss.Ver. Hamburg7: 163-186.
Amazoniaand their influenceon the selection of & F. White. 1979. Resurrectionof the genus
biologicalreserves.Pages 195-213 in G. T. Prance Dactyladenia (Chrysobalanaceae).Brittonia 31:
& T. S. Elias(eds.),Extinctionis forever.The New 483-487.
York BotanicalGarden,New York. & . 1988. A genericmonographof the
.1979a. New generaand species of Chryso- Chrysobalanaceaeand its relevance to practical
balanaceaefrom Malesia and Oceania. Brittonia and theoreticaltaxonomy and evolutionarybiol-
31: 79-95. ogy. Phil. Trans.Roy. Soc. B. 320: 1-184.
. 1979b. Two new species of neotropical Spichiger,R. &D. Masson. 1984. The Chrysobalana-
Chrysobalanaceae.Brittonia31: 248-252. ceae of the ArboretumJenaroHerrera.5th Con-
.1979c. New and interestingspeciesof Chrys- tributionto the study of the flora and vegetation
obalanaceae.Acta Amaz6nica8: 577-589. of the PeruvianAmazon. Candollea39: 13-44.
. 1979d. Chrysobalanaceae.In G. Harling& Tobe, H. & P. H. Raven. 1984. An embryological
B. Sparre(eds.),Floraof Ecuador10: 1-24. Stock- contributionto systematicsof the Chrysobalana-
holm, Sweden. ceae I. Tribe Chrysobalaneae.Bot. Mag. (Tokyo)
1979e. The taxonomy and phytogeography 97:397-411.
of the Chrysobalanaceaeof the Atlantic coastal White, F. 1976. The taxonomy,ecology and chorol-
forestsof Brazil.Revis. Brasil.Bot. 2: 19-39. ogy of the African Chrysobalanaceae(excluding
1979f. Distributionpatternsof lowlandneo- Acioa). Bull. Jard.Bot. Etat. 46: 265-350.
NumericalList of Taxa 99

Numerical List of Taxa, CollectingHistory, Status of Endangermentand Figure

Numbers. Taxa in boldfacedescribedsince 1972
Map Numberof Exsiccatae
Fig.
No. 1972 May 1987 Status'
1. Chrysobalanus
1. C. icaco L. ............. ................................................... 20 >10 217
2. C. cuspidatusGriseb. ex. Duss............................................ . 21 >10 3
3. C. venezuelanusPrance .....................................................21 0 5 R
2. Licania
1. L. michauxiiPrance.......-................. ..... 61 >10 46
2. L. boliviensisPrance ....... ...
................................. 28 3 0 R
59
3. L. maritima Prance.................................................................... 3 1
4. L. durifoliaCuatrec.............-.......................
....... 36 4 6
.
4.1. L. filomenoi Prance ............................................ 38 - 1 R
4.2. L. grandibracteataPrance .................................................38 - 4
5. L. macrocarpaCuatrec ......................... .........42
.......... 2 10
....................40
5.1. L. gentryiPrance ........................................ - 1 R
5.2. L. cabreraePrance.............-...................... 30 - 1 R
5.3. L. fasciculata Prance..........................-.......... 38 - 3
5.4. L. montanaPrance.................................................... 65 -1 R
6. L. salzmannii (Hook. f.) Fritsch.- ............................. 77 5 3
-51
7. L. klugii Prance.............................................. 1 2 R
8. L. guianensis(Aubl.)Griseb .....................-............. .. 43 >10 14
9. L. retifoliaBlake .... ..........................73
...
................. 2 1 R, E
10. L. longipedicellataDucke ..........................-........... 57 2 2 R
11. L. tomentosa(Benth.)Fritsch....................-................ 82 >10 11
12. L. pyrifoliaGriseb .............-73 .
............................ >10 24
13. L. leucosepalaGriseb. .......................................................55 10 14
..................155
13.1. L. dodsoniiPrance ........................................ - 2 R, E
14. L. angustataPrance .............................................. . 23 2 0 R
14.1. L. anneaePrance ............................................ 23 - 1 R
15. L. platypus(Hemsl.)Fritsch ............-....................... 72 >10 31
16. L. gonzalezii Miranda.................-......................... 42 2 2
17. L. egleriPrance ...................-.................. 37 >10 45
18. L. minutiflora(Sagot)Fritsch ...................-................... 64 >10 16
18.1. L. chiriquiensisPrance........................................... 33 - 2 R
......................51
18.2. L. kallunkiaePrance.....................-.........-.. - 2 R
18.3. L. guatemalensisLundell.........................-................... 43 - 2 R
19. L. maranhensisPrance...................................................... 60 1 0 R, E
20. L. fritschiiPrance ............................................... . 39 4 0 R
...
21. L. brittenianaFritsch....-...................... .. 29 >10 15
21.1. L. cecidiophoraPrance..................................................30 - 4
22. L. unguiculataPrance.-......................... ....... . 83 8 15
23. L. longipetalaPrance....................-.......-.....-..--. 57 3 7
.
23.1. L. tachirensisPrance.....................-........................ 81 - 1 R
24. L. wurdackiiPrance.........................-............. 85 - 12
...................84
25. L. turbinataBenth. ....................................... 9 2
26. L. lata Macbr.................... ............................. 53 >10 21
27. L. apetala (E. Mey.) Fritsch
a. var. apetala ..--.........................--....... 24 >10 132
b. var. aperta(Benth.)Prance.......................-............ 24 >10 65
. 42
27.1. L. granvilleiPrance ....................................................... -13
28. L. parvifoliaHuber............-.............................. 71 -45
29. L. maguireiPrance .......................................... 59 3 0 R
30. L. gardneri(Hook. f.) Fritsch ....................................... 40 >10 22
31. L. cuspidata(Rusby)Prance....................................... 34 1 0 R, E
31.1. L. jefensis Prance .......................................-....... 50 - 6
31.2. L. morii Prance..............-........-............... 65 - 1 R
100 FloraNeotropica
Continued
Map Number of Exsiccatae
Fig.
No. 1972 May 1987 Status'
32. L. sparsipilisBlake .................................................................................
79 7 6
32.1. L. cuatrecasasiiPrance .......................................................................
34- 2
32.2. L. mexicanaLundell .............................................................................
60- 1 R
33. L. emarginataHook. f. ......................................................................
389 2
33.1. L. joseramosiiPrance..............................
................
.. ..............
... 51- 1 R
34. L. calvescensCuatrec. ..-.--.............-....-......
......... 30 2 2
35. L. persaudiiFanshawe& Maguire. ............................................ 71 9 1
36. L. sprucei(Hook. f.) Fritsch . ............. ..... 79 >10 21
37. L. sclerophylla(Mart.ex Hook. f.) Fritsch......................... 78 >10 30
38. L. albifloraFanshawe& Maguire....................................23 2 0 R
39. L. longistyla(Hook. f.) Fritsch . ..................................
58 >10 47
40. L. fuchsii Prance....................................... 39
....................................... 1 0 R
41. L. humilis Cham. & Schlecht............................. ... 47 >10 34
42. L. foveolata Prance............................................................................
39 2 0 R
43. L. octandra(Hoffmgg.ex Roem. & Schult.)Kuntze
67 >10
a. subsp. octandra................................................................................. 63
b. subsp.pallida (Hook. f.) Prance........................................... 68 >10 76
c. subsp.grandifoliaPrance .........................................................
68 0 8
44. 74 >10
L. rigidaBenth. ....................................................................................... 9
45. 26 >10
L. arboreaSeem. ..................................................................................... 60
46. L. velataCuatrec . 1
....................................................................................
85 3
47. L. subarachnophyllaCuatrec.........................................................
80 2 2 R
47.1. L. tambopatensisPrance ....................................................................
62- 1 R
48. L. salicifoliaCuatrec ...........................................................................
76 1 4 R
49. L. araneosaTaub. .......................................................................
25 4 0 R
50. L. silvaticaGlaziou ex Prance 1
. .....................................................
79 0 R, E
51. L. chocoensisCuatrec..........................................................................
33 3 4
52. L. licaniiflora(Sagot)Blake . ........................................
56 >10 27
53. L. hirsutaPrance . ...................................
.. 46 2 7
54. L. costaricensisStandl.& Steyerm . .............................. 32 1 0 R, E
55. 4
L. krukoviiStandl. .................................................................................
51 4
56. L. lasseriMaguire.......................
.............................
......... 54 8 14
57. L. latifoliaBenth. ex Hook. f. .......................................................
54 >10 31
57.1. L. hispidaPrance ....................................................................................
46 -1 R
58. L. minusculaCuatrec .......................................................... 63 1 1 R
59. L. operculipetalaStandl.& L. 0. Wms. 68 2 0 R, E
60. 75 >10
L. reticulataPrance ............................................................................... 21
60.1. L. pakaraimensisPrance...............................................................
70- 5
61. L. arachnoideaFanshawe& Maguire. ...................-- ...... 25 2 1 R
62. L. oblongifoliaStandl....i.............................................. 66 >10 20
63. L. macrophyllaBenth. ..................................................... 58 >10 27
64. L. caudataPrance ..................................................................................
31 3 24
64.1. L. miltonii Prance ................................................................................
64- 2
65. 54 4
L. latistipulaPrance.............................................................................. 2
66. L. divaricataBenth. ..............................................................................
36 >10 5
67. L. glabrifora Prance.............................................................................
41 5 4
68. L. intrapetiolarisSpruceex Hook. f. ........................................ 50 8 31
69. L. heteromorphaBenth.
44
a. var. heteromorpha........................................................................
>10 213
b. var. glabra(Mart.ex Hook. f.) Prance ........................... 45 >10 93
c. var. subcordataFritsch................................................................
45 4 1
d. var. perplexansSandw. ...............................................................
45 >10 0
e. var. revolutaPrance.......................................................................- 1
69.1. L. laevigataPrance -..............................-.........
- 62-9 R
69.2. L. occultansPrance ..........................................................................
62- 2
70. L. glaziovianaWarm ..........................................................................
41 2 1 R
NumericalList of Taxa 101

Continued
Map Number of Exsiccatae
Fig.
No. 1972 May 1987 Status'
71. L. littoralisWarm.
8
a. var. littoralis....................................................................................
57 6
b. var. cuneataKuhlm ..................................................................... 0 R, E
72. L. fanshawei Prance.............................................................................
38 5 3
73. L. irwiniiPrance......................................................................................
503 2
73.1. L. marleneaePrance..............................................................................
60- 1 R
74. L. cyathodesR. Ben .................................................... 35 4 0
75. L. polita Spruceex Hook. f. ..................-............................ 72 >10 13
76. 3
L. silvae Prance ........................................................................................
78 10
77. >10
L. densifloraKleinh..............................................................................
36 35
78. L. cupreaSandw ....................................................................................
34 8 1
78.1. 27 -2
L. arianeae ................................................................................................... R
79. L. impressaPrance...............................................................................
49 4 6
80. L. dealbataHook. f ...........................................................................
35 >10 22
80.1. 77-
L. santosii Prance ........................................................................... 4
81. L. pallida Spruceex Sagot ...............................................................
69 >10 46
82. 42 >10
L. gracilipesTaub ................................................................................. 8
83. L. parvifructaFanshawe& Maguire...................................... 71 7 14
84. L. cymosa Fritsch-.............
- ..............
.......... 35 3 5
85. L. ternatensisHook. f. ex Duss ..-............... ............ 82 >10 3
86. L. membranaceaSagotex Lanes .............................................. 61 >10 20
87. 3
L. piresii Prance.......................................................................................
73 3 R
87.1. 40-
L.furfuraceaPrance............................................................................. 1 R
88. L. hypoleucaBenth.
a. var. hypoleuca.........-.............................. 48 >10 103
b. var.foveolata Prance................................................................2 0 R
89. L. boyaniiTutin ..-........................... ......... 28 5 4
90. L. buxifoliaSandw ...............................................................................
29 3 4
91. L. orbicularisSpruceex Hook. f. ................................................ 69 7 3
92. 66
L. niloi Prance ......................................................................................1 1 R, E
93. 32
L. coriaceaBenth ................................................................................>10 8
94. 84
L. urceolarisHook. f. .......................................................................... 8 17
95. 22 >10
L. affinisFritsch....................................................................................... 5
95.1. 81
L. teixeiraePrance .................................................................................-1 R, E
96. 41
L. glauca Cuatrec .................................................................................. 3 2
97. L. davillifoliaBenoist ....................................... 35 >10 5
98. 37
L. ellipticaStandl. ..................................................................................7 9
99. 31
L. canescensBenoist .............................................................................>10 89
100. L. couepiifoliaPrance ..........................................................................
32 1 1 R
100.1. L. naviculistipulaPrance...................................... 65 2 0 R
101. L. trigonioidesMacbr. ....................................... 82 1 0 R, E
102. L. cordataPrance ...............................................................................
33 5 3
103. 8
L. foldatsii Prance ..................................................................................
39 3
104. L. hebanthaMart. ex Hook. f. ........................................46 3 2
105. L. steyermarkiiMaguire..............-........................ 80 5 4
106. 81
L. subrotundataMaguire................................................................ 7 2
107. L. crassiveniaSpruceex Hook. f. ............................................... 32 1 8 R, E
108. >10
L. majusculaSagot ................................................................................
59 15
108.1. 50
L. jimenezii Prance ................................................................................- 4
109. 22
L. alba (Bern.)Cuatrec ..................................................................... >10 27
110. 46
L. hitchcockiiMaguire ....................................................................... 2 0 R
111. 77
L. sandwithiiPrance......................................................................... 1 0 R
112. 55 >10
L. laxifloraFritsch .............................................................................. 11
113. L. rufescensKlotzschex Fritsch................................... 76 >10 5
114. 52
L. kunthianaHook. f. ......................................................................... >10 60
115. 27
L. bellingtoniiPrance........................................................................... 1 0 R, E
102 FloraNeotropica
Continued
Map Number of Exsiccatae
Fig.
No. 1972 May 1987 Status'
116. L. compactaFritsch ....-..........................
..... 33 1 0 R
117. L. ovalifoliaKleinh .......................................................... 69 9 3
118. L. caldasianaCuatrec.......................................................30 3 0 R
119. L. savannarumPrance.......................................................77 >10 21
119.1. L. stewardiiPrance ...........................................................80 - 16
120. L. microphyllaFanshawe& Maguire..................................... 61 1 0 R
121. L. triandraMart.ex Hook. f. ................................ 83 7 7
82 -
121.1. L. tocantinaPrance ............................................................................... 3
122. L. discolorPilg............................................................... 36 >10 12
123. L. apiculataPrance ............................................. . 25 3 0 R
124. L. micranthaMiq .................................................................................
63 0 64
125. L. pruinosaBenoist ...............................................................................
73 5 0
126. L. nitida Hook. f. .................. .................
........... 66 9 5
126.1. L. aracaensisPrance .................................................. 25 - 4
127. L. riedeliiPrance.....................................................................................
75 4 2 R, E
128. L. bracteataPrance .........................................
.......................... 29 8 8
129. L. parvifloraBenth .........................................................70 >10 33
130. L. robustaSagot ....................................................
... 75 8 3
130.1. L. lamentandaPrance ................................................ 53 - 3 E
131. L. lanceolataPrance ............................................... 53 2 23
132. L. spicataHook. f. ...........................................................79 >10 1
133. L. strictaKleinh ........................................................... 80 1 0 R
134. L. leptostachyaBenth ...................................................... . 56 >10 46
135. L. incana Aubl. .....................................................................................
49 >10 21
135.1. L. nelsonii Prance ...........................................................62 - 2 R
136. L. paraensisPrance...........................................................70 4 6
137. L. vaupesianaKillip & Cuatrec........................................84 3 1
138. L. bahiensisPrance ....................... ...... ...
............ 27 1 1 R, E
139. L. maxima Prance .. .................. 60 1 1 R
140. L. mollis Benth. .......................................................................................
65 >10 43
141. L. blackiiPrance .................................................. 28 >10 19
142. L. rodriguesiiPrance......................................... 76 8 6
143. L. indurataPilg.............................................. 49 2 1 R
144. L. hoehneiPilg. ...............................................................47 >10 15
144.1. L. harlingiiPrance ...........................................................43 - 8
145. L. cruegerianaUrb .........................................................34 >10 10
146. L. belemii Prance ..................................................................................
27 1 6
147. L. splendens(Korth.)Prance........................-............
.......-- Asiatic
147.1. L. palawanensisPrance.................................................................
- Asiatic
147.2. L. fusicarpa(Kosterm.)Prance ....................................... - Asiatic
148. L. veneralensisCuatrec............................................................85 1 11
149. L. amapaensisPrance ...................................................... . 23 1 4
150. L. tepuiensisPrance..........................................................81 1 0 R
151. L. obtusifoliaFritsch -1.....................................- 0 R
152. L. roraimensisStandl..................................................... . 76 1 0 R
3. Parinari
1. P. campestrisAubl. .................................. .
.................. 87 >10 29
2. P. montanaAubl ....................................... 90 >10 14
3. P. rodolphiiHuber .................................................. 92 >10 15
3.1. P. alvimii Prance .....................................................
. 87- 2
4. P. excelsa Sabine..................................................... 89 >10 126
5. P. occidentalisPrance ........................................................91 4 8
6. P. spruceiHook. f. ................................................................................
92 7 21
7. P. pachyphyllaRusby ..........................................................................
91 >10 35
8. P. brasiliensis(Schott)Hook. f. .......................................... 87 2 3 R
9. P. klugii Prance ...............................................................88 1 11
Numerical List of Taxa 103

Continued
Map
Number of Exsiccatae
Fig.
No. 1972 May 1987 Status'
10. P. maguirei Prance ........................................................... 90 2 3
11. P. littoralis Prance ........................................................ 88 1 6
12. P. parvifolia Sandw. ..................
.................................... 91 4 1
13. P. cardiophylla Ducke ...................................................... 88 3 0 R
14. P. parilis Macbr ......................................................................................
91 2 12
15. P. chocoensis Prance ...................................................
................
......... 88 1 4
16. P. obtusifolia Hook. f. ....................-................ 90 >10 28
17. P. romeroi Prance ....................................................... 92 2 3
4. Exellodendron
1. E. coriaceum (Benth.) Prance ............................................... 95 >10 24
2. E. cordatum (Hook. f.) Prance ..............................................94 5 18
3. E. barbatum (Ducke) Prance ........................................................
94 >10 30
4. E. gardneri (Hook. f.) Prance .................................................. 95 6 10
5. E. gracile (Kuhlm.) Prance ..............................................................
95 1 2 R, E
5. Maranthes
1. M. panamensis (Standl.) Prance & White 153 4 12
6. Couepia
1. C. guianensis Aubl.
a. var. guianensis 1>.......10................................................ 105 >10 66
b. var. glandulosa (Miq.) Prance ............................................... 105 >10 18
c. var. divaricata (Hub.) Prance ............................................ 104 >10 14
(2.) C. glandulosa Miq. = C. guianensis var. glandulosa .. -
3. C. paraensis (Mart. & Zucc.) Benth.
112
a. subsp. paraensis .............................................................................. >10 36
b. subsp. glaucescens (Spruce ex Hook. f.) Prance ....... 112 4 114
c. subsp. cerradoana Prance ................................................112 4 4
(4.) C. leptostachya Benth. ex Hook. f. =
C. guianensis var. guianensis ............................................. -
5. C. maguirei Prance ....................................
. ....... 108 4 1
6. C. sandwithii Prance 1.. 115 4 4
6.1. C. bernardii Prance ........................................................ 97 - 14
6.2. C. monteclarensis Prance .......2................................... - 2
7. C. parillo DC ................................................................. 115 >10 32
8. C. steyermarkii Maguire ................................... ......... 116 1 R
8.1. C. canescens (Gleason) Prance . ................. .................. 98 1 3
9. C. canomensis (Mart.) Benth. ex Hook. f. ..........................98 >10 21
10. C.foveolata Prance.........................................................
. 103 7 3
11. C. magnoliifolia Benth. ex Hook. f. ........................................ 108 >10 21
12. C. exflexa Fanshawe & Maguire ........................................... 103 3 0
13. C. habrantha Standl ........................................................ 106 >10 7
13.1. C. scottmorii Prance .......................................................... 115 - I R, E
13.2. C. carautae Prance ................................ ...... . 99 - 1 R, E
14. C. spicata Ducke ................................................... 115 1 7
15. C. bracteosaBenth. .............................................. . 98 >10 40
16. C. subcordataBenth. ex Hook. f. .................................... . 116 >10 11
17. C. belemiiPrance....................................................
. 97 3 6
18. C. . 99
caryophylloides Benoist .................................................
a. subsp. caryophylloides ............................... >10 >10
b. ............
subsp. glabra Prance ....................................... .- 1
19. C. excelsa Ducke ...............................................102 4 2
20. C. uiti (Mart. & Zucc.) Benth. ex Hook. f. .........................117 >10 10
21. C. cataractae Ducke ................................................ 99 8 16
22. C. macrophyllaSpruceex Hook. f. ........................................
108 9 3
23. ........ ............
C. krukovii Standl ................................. .... 107 2 2
104 FloraNeotropica

Continued
Map Numberof Exsiccatae
Fig.
No. 1972 May 1987 Status'
24. C. latifolia Standl. ...............................................................
107 2 1
25. C. ovalifolia(Schott)Benth .......................................1.....111 >10 23
26. C. schottiiFritsch.................................................................
. 115 >10 8
27. C. grandiflora(Mart.& Zucc.)Benth. ex Hook. f........ 104 >10 27
28. C. elata Ducke ..................................................... 102 >10 7
29. C. racemosaBenth. ex Hook. f. ............................. ............ 113 34
29.1.C. amaralaePrance ...............................................
. . 97 - 12
30. C. martiniiPrance .......................................................... . 109 1 0 R, E
31. C. bondariiPrance ................................... 97 2 0 R, E
32. C. insignisFritsch ..........................................................106 5 2
32.1.C. cidiana Prance .................................................. 99 - 2
33. C. recurvaSpruceex Prance................................................. 114 1 1 R, E
34. C. obovataDucke .............................................
.. 110 >10 15
35. C. williamsiiMacbr ......................................................... . 118 >10 22
35.1.C. glabra Prance.......................................................103 -2 R
35.2.C. marleneaePrance .......................................... 108 - 8
36. C. chrysocalyx(Poepp. & Endl.)Benth. ex Hook. f. ...100 >10 42
37. C. erianthaSpruceex Hook. f............................................ 102 >10 5
38. C. trapezioanaCuatrec ...................................... 118 4 4
39. C. stipularisDucke ................................... 116 2 0 R
40. C. reflexaDucke ...................................................114 1 0 R
41. C. longipendulaPilg ................................................. 107 >10 12
41.1.C. dolichopodaPrance . .
....................... 101 - 11
42. C. cognata(Steud.)Fritsch
a. var. cognata ............................................................101 >10 3
b. var. majorPrance ................................... 101 3 0
c. var. membranaceaPrance .................................. .. . 101 2 0
43. C. multifloraBenth. ......... . 109
...................................... >10 5
44. C. uleiPilg ..................................................... 117 >10 52
45. C. comosa Benth .................. ...........
. .. . 100 >10 3
46. C. venosaPrance ...................................................... 118 9 1
47. C. polyandra(Kunth)Rose ................................... 113 >10 21
47.1. C. nutansPrance ...........................................
. 109 - 1 R, E
48. C. platycalyxCuatrec ............................... ...... 111 2 4 E
49. C. rufaDucke ......................................................... 114 >10 1
50. C. robustaHuber .................................................114 >10 22
51. C. impressaPrance
a. subsp. impressa........................................
................ . 106 4 3
b. subsp. cabraliaePrance . ....................................... 100 - 10
52. C. meridionalisPrance ..............................................109 1 0 R, E
52.1. C. coarctataPrance ......................................... 100 - 3
52.2. C. longipetiolataPrance .................................... 107 - 2 R, E
53. C. pernambucensisPrance ................ .................... 111 1 1
54. C. froesii Prance........................................................... 103 3 0 R
55. C. parvifoliaPrance.........................................................111 2 0 R, E
7. Hirtella
1. H. myrmecophilaPilg .......................... ....... 139 >10 17
2. H. physophoraMart. & Zucc ............................................. . 142 >10 51
3. H. vesiculosaSuesseng.................................................... . 152 2 0 R
4. H. dorvaliiPrance ...............................
..... 128 13
5. H. guainiae Spruceex Hook. f. ............................................. 132 >10 35
6. H. duckeiHuber . .........................................
.................. . 128 >10 28
6.1. H. revillaePrance ............................................146 - 2
7. H. macrosepalaSandw ................................... 137 6 3
8. H. ulei Pilg. ....................................................
. 152 10 20
9. H. glabrataPilg ............................................................. 132 >10 11
NumericalList of Taxa 105

Continued
Map Number of Exsiccatae
Fig.
No. 1972 May 1987 Status'
9.1. H. confertifloraPrance........................................ 126 - 1 R
10. H. carbonaria Little . ................
...................... 125 >10 3
11. H. araguariensis Prance ...................................................... 121 1 4
11.1. H. barnebyi Prance ..........................................................
. 122 - 2
11.2. H. margae Prance ..................................................................................
138 - 3
11.3. H. liesneri Prance ............................................. 136 - 1
12. H. cordifolia Prance . ................ .............................................................
126 2 0 R
13. H. insignis Briq. ex Prance 15-----.......................................... 135 2 7
14. H. tocantina Ducke 103-----------.. ...................................150 3 3
15. H. piresii Prance ----.......................................
130-- 143 0 8
16. H. davisii Sandw ................................8...
........................1280 9
17. H. subglanduligera Pilg ..................410................... 149 1 0 R, E
18. H. ciliata Mart. & Zucc. ..................................................... 126 >10 45
19. H. hoehneiPilg. .............................................................134 8 6
20. H. glandulosa Spreng. ..........................11>0......... 131 >10 99
21. H. bullata Benth ............................................................124 >10 54
22. H. americanaL ............................................... . 120 >10 65
22.1. H. santosii Prance ........ ...............................................147 - 1 R, E
23. H. guatemalensis Standl ................................. 133 >10 17
24. H. eriandraBenth .................. ...........
............................ 129 >10 49
25. H. paniculata Sw ... ........................... 140 >10 78
26. H. deflexaMaguire......................................
.127 1 4
27. H. tentaculataPoepp. & Endl ....................................................
150 >10 1
28. H. macrophylla Benth. ex Hook. f. .....................................138 >10 8
29. H. adderleyiPrance.... ..........................................
. 120 2 15
30. H. punctillata Ducke ............................
............. ... 143 >10 18
31. 127
H. corymbosaCham. & Schlecht ........................................ 4 0 R
32. H. barrosoi Prance .. ........... . 122 >10 0
33. H. pendulaSoland. ex Lam .................... ................... 141 2 1
34. H. leonotisPittier . ..... ......... 136 1 7
35. H. mutisiiCuatrec....................................... 139 8 5
36. H. triandraSw.
a. subsp. triandra .................................
.... 151 >10 187
b. subsp.punctulata(Miq.) Prance ........................... 152 >10 8
c. subsp. media (Standl.)Prance...........................................
152 >10 15
37. H. bahiensisPrance............................................... 122 3 3
38. H. latifolia Prance .. .1.......................................136
. 2 14
39. H. suffulta Prance . 149 6
.................................................................................. 15
40. H. elongataMart.& Zucc ................
.................... 129 >10 64
40.1 H. magnifolia Prance . .......................................138
. - 6
41. H. rodriguesiiPrance . ......................................147 5 12
42. H. obidensis Ducke ............................................... 140 >10 14
43. H. cowaniiPrance& Maguire ..-...-...........................
. 127 5 2
44. H. orbicularis Prance . ......................................140 3 0 R
45. .... 133
H. guyanensis(Fritsch)Sandw............................................ 8 0
46. H. lightioidesRusby . ............................................................................
137 3 8
47. H. aramangensis Prance .-. ...................... .. 121 1 0 R, E
48. H. rasa Standl ........ ...................... 146 2 1
49. H. scabra Benth. .................................................... 148 >10 26
50. H. bicornisMart.& Zucc.
a. var. bicornis......................................................123 >10 29
b. var. pubescens Ducke ............................................ . 123 >10 45
51. H. angustissima Sandw ............................
....... 121 8 1
52. H. tenuifolia Prance .... . .. 150 3 17
52.1. H. radamii Prance . 143 -
............................................................................... 1
53. H. pilosissima Mart. & Zucc ............... ..................... 142 >10 22
106 Flora Neotropica

Continued
Map Number of Exsiccatae
Fig.
No. 1972 May 1987 Status'
54. . 143
H. gracilipes(Hook. f.) Prance ............................................. >10 67
55. H. brachystachyaSpruceex Benth........................................ 125 >10 10
55.1.H. arenosaPrance ...................... ...... .. 122 - 4
55.2.H. conduplicataPrance .............................................................
125- 2
56. H. racemosaLam.
a. var. racemosa........................................................... 145 >10 282
b. var. hexandra(Willd. ex Roem. & Schult.)Prance 144 >10 340
146
c. var. glandipedicellataPrance.................................................. 2 0
d. var. hispidaPrance.......................................................................
146 - 3 R, E
57. H. juruensisPilg ................................................................................
135 3 1 R
58. H. kuhlmanniiPilg ............................................................. 135 2 1 R
59. H. standleyiBaehni & Macbr ..................... .........................149 2 7
60. H. longifoliaBenth. ex Hook. f. .-......................................... 137 1 0 R, E
61. H. lemsii L. O. Wms. & Prance ........................................... 136 1 10
62. H. schultesiiPrance .........................-...-.........................148 7 31
63. H. paraensisPrance..-.............-..-..................... . 141 >10 10
64. H. spruceiBenth ....................................................................................
148 >10 6
65. H. lancifoliaDucke ...........................-.................
.. 135 5 2
66. H. burchelliiBritton.....-....................-......................... 124 >10 24
67. H. mucronataPrance..................-.............................
..... 139 3 3
68. H. longipedicellataPrance.-..................................
............... 137 6 5
69. H. glandistipulaDucke ................
................. 132 8 2
70. H. martianaHook. f. .........................................................................
139 >10 11
70.1. H. parviunguisPrance ................................. ...........141 - 2
71. H. pimichina Lasser& Maguire.................................................. 143 4 1
72. H. subscandensSpruceex Hook. f .............................-.......... 149 4 1
73. H. hispidulaMiq ............................................................. 134 >10 78
74. H. silicea Griseb .....................-...............
. ....... ................148 >10 19
75. H. excelsa Standl.ex Prance ............................ ....... 130 4 10
76. H. adenophoraCuatrec.................. ............... ............ 120 3 0 R
77. H. caducaFanshawe& Maguire................................................. 125 6 1
78. H. fasciculata Prance .-................................................
.. 130 2 1
79. H. couepiifloraPrance.........-............................................. 127 2 2
80. H. tubifloraCuatrec ..............................
........................... 150 1 6
81. H. floribundaCham. & Schlecht ......................................... 130 >10 0
82. H. angustifoliaSchott ....................................................... 121 >10 8
83. H. rugosaThuill. ex Pers ...........................-...................... 147 >10 10
84. H. scaberulaSpruceex Hook. f. .................................-.......... 147 1 0 R
85. H. hebecladaMoric. ex DC. .......................-..-............... . 134 >10 11
86. H. enneandraCuatrec ......................... ......
.......-....... ...... 130 1 0 R
87. H. paucifloraLittle ..............................
.............. ........... 141 1 2
88. H. glazioviiTaub ...................................-.......................132 7 0
89. H. megacarpaR. A. Grah. ............................. ........ .. - African
90. H. zanzibaricaOliv ............................ ...........................African
91. H. cliffortianaVeil ..........................................................
. - I 0 R
92. H. pohlii Hook. f. ......................... .....-..... 1 0 R
8. Acioa
1. A. guianensisAubl ........................................................... 154 7 1
2. A. somnolensMaguire...................................................... 154 1 0 R
3. A. schultesiiMaguire......................................................... 154 2 10
4. A. edulis Prance .....................................................
........ 102 - 8
9. Neocarya
1. N. macrophylla(Sabine)Prance..............................-......... . - 1
(African
sp.)
'R = rare; E = endangered.
 List of Exsiccatae
Supplemental
SUPPLEMENTALLIST OF EXSICCATAE
This list includes materialnot cited in Prance(1972), largelyrecent collections made since that
date and a few older ones not seen previously.
Abadie, B., 50T (6-44)
Acero, E. et al., 57 (7-50b); 195 (2-27b); 729 (2-121)
Acevedo R., P. et al., 1657 (20-13.1)
Ackerly,D. 0., 07 (7-2); 152 (7-50a); 161 (7-56b)
Acosta M., A., 1 (2-17)
Adair de Oliveira,INPA59936, INPA60162 (2-69.1);
INPA73467(6-41);INPA73471,INPA73472(2-62);
INPA73476, INPA73477 (2-69a); INPA73483 (2-
64);INPA73484 (6-la); INPA74791 (2-69.1);INPA
s.n. (2-69a)
Adams, C. D. A., 14572 (1-1); YSB 86 (7-74)
Agostini, G., 305 (2-77); 1104 (7-36a)
Albertde Escobar,L. et al., 1414 (7-36a);3282 (2-51);
3525 (1-1)
Albuquerque,B. W. P. de et al., 646 (7-56a); 671 (7-
6); 672 (2-69b); 798 (2-81); 841 (2-36); 876 (7-2);
886 (7-55.2); 67/20 (2-69.1); 887 (7-55.2); 919 (2-
17); 950 (2-69b); 954 (2-81); 956 (2-69b); 998 (2-
81); 1010 (2-39); 1048 (2-81)
Alencar,L., 26 (7-56a);89 (7-24); 112 (2-69b); 161 (7-
56b); 238 (2-129); 249 (2-88); 251 (6-lb); 260 (2-
88); 390 (7-5); 486, 499 (3-6); 570 (7-55); 616 (3-
41); 725 (2-129)
Allem, A. et al., 7, 2324, 2391 (6-20); 2489 (2-28)
Allen, P. H., 3019 (2-68)
Almeida, E. de F. et al., 275 (2-80.1); 283 (4-2)
Almeida, J. et al., 150 (2-80.1); 164 (3-1.3); 185 (7-
36a); 1308, 1791 (6-25)
Aluisio de Sousa, J., INPA59853 (2-99); s.n.
INPA61445 (7-1); INPA59853 (2-99); INPA61868
(6-4);INPA61951 (7-6);70666 (2-62);70678 (6-11);
70679 (7-1); 70682 (6-34); INPA-s.n. (2-43b)
Amaral,D. L., 728 (7-20)
Amaral,I. L. et al., 11 (6-3b); 39 (7-40); 40 (2-75); 43
(2-129); 102 (2-27a);56 (6-29); 131 (2-69a); 176 (7-
73); 186 (2-124); 216 (2-75); 333 (2-114); 353 (3-4;
390 (2-69b);466 (2-26); 535 (2-129);588 (2-26) 596
(2-129); 602 (2-114); 703 (7-56a); 729 (7-40); 864
(7-20); 909 (7-56a); 1006 (7-19); 1022 (7-50a); 1041
(7-58); 1050 (7-36a); 1105(2-17); 1119(7-20); 1193
(7-56b);1271(7-73);1368(2-43b);1396(2-57);1435
(7-25); 1448 (6-43); 1498 (6-15); 1695, 1697, 1698
(6-7); 1708 (2-119.1); 1757 (7-11)
Ancuash,E., 4, 354 (6-36); 752 (2-21.1); 1282 (7-21)
Anderson,A. B., 207 (7-9); 354 (6-29)
Anderson,C. W., 14 (2-109); 57 (2-77); 325 (3-1)
Anderson,W. R., 6248, 6259 (2-80); 6873 (4-4); 7281
(7-54); 7420 (2-30); 7543 (2-54); 7898, 8085 (2-80);
8702 (7-20); 10071 (3-16); 10653 (7-24); 10771 (6-
9); 11037 (2-27a);11065(6-9); 11037 (2-27a);11065
(6-9); 11921 (7-24); 12064 (2-27b); 12079 (2-69b);
12208 (2-43b); 12303 (7-54)
Andrade,P. M. et al., 252 (6-6.2); 420 (6260) (2-114);
621 (6541) (7-64)
Andrews,L. M., 911 (2-1); 3-103 (2-40)
Angelo, L., 11 (7-48); 11 (2-7)
Antonio,T., 2384 (7-56b);3763 (7-56a);4324 (7-56b);
4599, 4665 (7-36a)
107
Aranda,A., 60 (2-15); 73 (7-36a)
Araquistain,M., 34 (1-1); 2181 (7-22); 3128 (7-36a);
3149 (5-1)
Arafijo,D., 1425(7-85);2151 (1-1);4041 (7-36b);4146
(1-1); 4536 (7-36b);4719 (6-26); 4727 (6-25); 4736
(1-1); 4932, 5261 (6-26); 5834, 6309, 6337 (1-1);
6400 (6-25); 6408, 7517 (1-1)
Araijo et al., s.n. INPA94326 (7-73)
Araijo, J. da S., 72 (2-6); 104 (7-56b)
Argent,G. C., 6479 (2-41)
Aristeguieta,L. et al., 4493 (7-56b);4990 (2-27); 5131
(2-27b);6433 (2-12);6477 (7-56b);6493 (6-3);6867
(1-1); 6908 (7-36a);6977 (3-7); 7096 (7-34); 12529
(7-22)
Armond, 175 (3-4)
Ar6steguiV., A., 105 (2-128)
Asplund,E., 10055 (6-33); 12538 (2-4)
Assis, J. S., 52 (4-2); 188 (2-80); s.n. (4-2)
Atwood, J. T., 4526, 4723, 5261 (1-1)
Austin, D. F. et al., 6994 (7-24); 6995 (2-43a); 7058
(7-50a); 7180 (2-149); 7185 (2-109); 7251 (2-134);
7315 (2-69a); 7368 (2-109); 7394 (2-43a); 7396
(3-4)
Avalone, C. L. et al., 26 (2-41)
Ayala, F., 615 (2-39); 2888 (6-16)
Aymard,G. et al., 371 (2-77);2536, 2637 (7-36a);3230
(6-3b);3343 (7-40);3483 (7-56b);3900 (2-109);3911
(7-74); 4074 (2-145); 4076 (2-109); 4142 (7-56)
Badillo, V. M., 1559 (7-56b)
BAFOG, 119 (3-1)
Bagazo,N., 166 (7-56a)
Bahia, R. P., 54 (7-56b)
Bahia,T. R., 129 (2-124); 130 (7-53); 145 (6-3b); 191
(2-62); 198 (2-124); 215 (6-41)
Baker,M. A., 6056, 6112 (6-36)
Baldwin,A. A., s.n. (2-1)
Balee,W. L. et al., 17 (6-la); 44 (2-99); 46 (2-86); 125
(2-99); 174 (2-114); 175, 182 (2-69a); 218 (2-99);
228 (2-86); 243, 256, 286 (2-69a); 308 (6-la); 325,
326 (4-3); 336 (2-69a);339, 354 (2-86); 421 (2-57);
426 (2-99);501 (2-69a);508 (2-99);581 (2-114);599
(2-69a); 664 (6-la); 948 (2-27a); 975 (2-99); 1053
(2-69a);1058 (7-56a);1087 (6-lc); 1089 (6-la); 1107
(2-99); 1116 (2-63); 1124 (6-la); 1157 (3-3); 1170,
1179 (2-99); 1186 (6-la); 1191 (2-63); 1273 (3-3);
1276 (2-99); 1296, 1316 (6-lc); 1359 (2-114); 1375,
1378, 1429 (6-la); 1439 (2-99); 1451, 1455 (6-la);
1484 (2-99); 1501, 1509, 1516, 1518, 1520, 1521,
1522, 1524 (6-la); 1550 (7-56b); 1656, 1685, 1747
(7-63); 1786 (7-14); 1804 (2-69a); 1806 (2-8); 1861
(2-27a); 1934 (7-66); 3808 (2-17); 4063 (2-124)
Bamps, P., 5378 (7-6)
Barbosa,C., 1213 (7-80)
Barbour,P. J., 5478 (7-56b); 5505 (7-36a); 5764 (7-
56a)
Barclay,A. S. et al., 3701 (7-22)
Barlow,F. D., 30/62B (7-56a)
108
Barrier,S. et al., 2618 (7-56a);3863 (7-39);3989, 4259
(2-86)
Barros,S., 79 (6-3c)
Barroso,G. M. et al., 181, 290 (7-18); 355 (7-56b)
Bastos,N. C. et al., 79 (7-20); 87, 117 (2-43a); 127 (7-
56b)
Bautista,H. P., 34 (3-2); 48 (2-135); 52, 55 (7-56b)
Bawa,K. S., 383 (6-15)
Beaman,J. H., 6092 (6-47); 6263, 6333 (7-36c); 6423
(6-47)
Beck, C. H., s.n. (2-1)
Beck, S. G., 1480 (7-56a);4049, 6915, 7158, 8319 (7-
36a); 10158, 10172 (2-28); 12247 (7-26a)
Becker,R., 54 (7-54)
Begazo, N., 38 (2-64); 88 (2-27a); 144 (7-50b); 159,
166, 201 (7-56a)
Belem, R. P. et al., 2837, 2846 (6-5lb); 3169 (6-17);
3315 (6-5 lb)
Bellido, L. C., 2, 35 (2-27a)
Bena, P., 1317 (2-27a); 1328 (2-52); 1757 (7-36a)
Benitez de Rojas, C. E., 2578 (2-89); 2579 (2-81)
Benson, W. W., 5671 (7-50); INPA92145 (7-50a)
Berg,C. C., 221 (6-25);617 (2-17); 684 (6-21); 686 (4-
3); 697 (2-81); 712 (2-69a); 719 (2-81); P18449 (7-
54);P18452, P18565 (2-28);P18654 (2-75);P18678
(3-4);P18693 (2-28);P19462 (2-60);P19751 (2-28);
P19793 (6-3b); P19796 (7-73); P19922 (2-99);
P19928 (2-sp.);P19940 (2-28)
Berlin, B., 354 (6-36); 902, 976, 1799 (2-21.1); 3529
(7-56b)
Bermudez,B., 6 (1-1)
Bernal,C. et al., 142 (6-48)
Bernardi,A. L., 1-56 (7-40.1); 1-95 (2-64); 1-163 (6-
15); 2-109 (6-44); 2-162, 2-163 (2-17); 3-51 (2-55);
3-99 (6-44); 3-124 (6-34); 4-142 (6-15); 5-36 (2-94);
5-120 (2-124);6-6 (6-6.1);6-74 (6-44);6-123 (7-41);
6-148 (2-141); 6-189 (2-27a); 6-192 (2-27); 7-2 (7-
6); 7-84 (6-6.1); 7-89 (7-41); 8-30 (2-94); 8-128 (6-
15); 8-139 (2-60); 8-151 (2-55); 9-78 (6-15); 9-157
(2-141); 1624 (3-4); 2030 (2-13); 2127 (3-3); 2777
(1-3); 3069 (7-21); 3081 (7-56d); 6628, 6682 (3-4);
6760 (7-25); 7111 (7-16); 7356 (2-122); 7394 (7-
56b); 7416, 7672 (7-36a); 7824 (6-3b); 7825, 7828
(2-77); 8011 (2-122); 16219 (6-6.1); 16233 (7-6);
16334 (3-9)
Berry,P. E., 600 (2-69b); 633 (7-62); 662 (7-56); 737
(7-62); 1352 (7-50b); 1519 (2-69a); 2066 (2-27a);
2073 (2-69a); 2149 (7-49); 2231 (3-4); 3383 (3-7)
Bethancourt,A. et al., 91 (7-56b)
Bhorai,M., 8781 (2-44)
Billiet, F. et al., 1023 (7-25); 1180 (2-69a); 1201 (2-
108); 1234 (2-124); 1265 (1-1); 1298 (7-25)
Bisby,F. etal., P18071 (2-69a);P18077 (2-82);P18085,
P18102 (4-3); P18107 (7-56a);P18123 (2-56a)
Bitaillon,C., 64 (3 sp.)
Black, G. A., 48-2902 (2-69.1); 48-3610 (2-63); 49-
8473 (3-3)
Blanco C., C. A., 55 (2-113); 67 (7-20); 194, 254 (2-
77);421 (2-109);533 (7-20);673 (7-16);764 (7-56b);
766 (2-27b); 885 (7-36a); 1105 (2-17); 1112 (7-56);
1141 (2-17); 1157 (6-4); 1166a(2-77); 1166b(2-93);
1204 (3-6); 1221 (6-4); 1255 (2-140)
Bokermann,W., 5732 (7-82)
Boom, B. et al., 1997 (2-64);4087 (7-53);4167 (7-46);
FloraNeotropica
4178 (7-56a);4245 (7-46); 4335, 4338, 4343, 4346,
4386,4387,4395 (2-43b);4424 (7-46);4426 (2-43b);
4428 (2-21); 4461 (2-43b); 4464 (7-46); 4473 (2-
43b); 4497 (7-46); 4630 (7-53); 4700, 4729 (7-56a);
4940 (6-14); 5084 (2-43b); 5367 (7-62); 5868 (2-
27.1); 5919, 5952 (2-26);6032 (7-56b);6089 (2-88);
6263 (3-4); 6359 (2-145); 6490 (2-13); 6492 (2-99);
6514 (3-4); 6534 (2-88); 6536 (2-43a);6542 (7-20);
6551 (3-4); 6574 (7-56b); 6587 (3-4); 6592 (2-13);
6607 (2-99);6623 (2-13);6633 (2-145);6636 (2-99);
6641 (2-43a); 6660 (2-145); 6676 (2-99); 6677 (2-
88); 6685 (2-99); 6838 (7-83)
BorjasM., G., 175 (7-27)
Bossio, H., 34 (2-69b)
Boucas,P. R. P. et al., 13 (2-52); 17 (2-114); 148 (7-
56b)
Boyan, J., 75 FD7749 (7-11.2)
Braga,M. M. N. et al., 145 (6-3a)
Braga,P. I. S. et al., 3127 (2-10); 3347 (7-56a)
Brand,J., 971 (3-7); 992 (2-45)
Brandbyge,J. et al., 30575 (7-40);31228 (7-56b);32659
(6-36); 35032 (2-8); 36119 (7-53); 36177 (7-22)
Breedlove,D. E. et al., 20898 (1-1);34086 (7-22);34130
(7-56b); 58466 (7-22)
Breteler,F. J., 4606 (3-7); 4799 (7-50b); 4881 (7-40);
4883 (7-56b);4959, 4961 (2-77);4994 (2-109);5043
(2-27a); 5078 (2-77); 5123 (2-13); 5133 (3-4); 5134
(7-16)
Bristan,N., 1121 (6-22); 1131 (7-36a)
Brown,S. et al., 1639 (1-1)
Bruijn,J. de, 1252 (3-7); 1626 (3-3); 1635 (2-77); 1638
(2-88); 1664 (7-36a);1688 (3-4); 1694(2-109); 1696,
1698 (3-4); 1716 (2-77); 1729 (2-114); 1735, 1736
(2-88)
Brumbach,W. C., 8332 (2-1);9263, 9327, 9655, 9739
(1-1)
Briinig,E. et al., 160 (2-68)
Bunting,G. S. et al., 3505 (7-25); 3744 (2-69a);3746,
3825 (7-28); 4002 (1-68); 4032 (3-1); 4084 (2-88a);
4091 (2-119); 4231 (2-83); 5521 (3-7); 5655 (7-22);
5889 (3-7); 5947 (2-12); 6224 (3-7); 6225 (7-22);
6543 (3-7); 6682 (1-1); 6854 (2-43a); 7055 (3-7);
7124 (2-76); 7152 (7-22);7219 (2-43a);7321 (7-20);
8457 (3-7); 8725 (7-36a); 8750 (2-76); 8758 (7-22);
8763 (2-76);8764 (3-7);8797 (7-36a);8836 (7-56b);
8949 (7-36a);8954, 9192 (3-7);9832 (2-114); 10151,
10216 (7-36a);10587 (7-56b);10759 (2-43a); 10800
(3-7); 10929 (7-56a); 11036 (2-43a); 11274 (3-7);
11507 (2-114); 11529 (2-43a); 11992 (7-36a)
Burch,D., 3409 (1-1)
Burger,W. C. et al., 5784 (7-61)
Buschbacher,R. J., 39 (21)
Busey, P., 354 (7-56a)
B. W., 3371 (6-1a); 3404 (7-50b); 3827 (2-130); 4818
(7-25)
Byron et al. (=Albuquerque,B. W. P. de), 67-20 (2-
69.1);67-34 (6-15);67-59 (2-39);67-70 (2-69a);264
(6-3); 370 (3-6); 372 (7-50a); 377 (2-27a); 509 (6-
3b); 577 (2-1); 592 (2-28); 594 (6-3b); 633 (7-40);
886 (7-55b);924 (2-57); 927 (2-17); 946 (2-28)
Cabrera,E., 3138 (1-1)
CabreraR., I., 94 (2-5.2); 553, 600 (2-148); 2590 (3-
4); 2597 (7-56b); 3217 (7-56a); 3359 (2-69a)
Cain, S. A., 12 (7-36a)
 List of Exsiccatae
Supplemental
Caldas,T. S. P., 21 (6-27)
Calder6n,C., 79-105 (2-88a); 2521 (7-53); 2534 (7-
56a); 2542, 2591 (6-29); 2634 (7-9); 2806 (7-56a);
2820 (2-37); 2936 (2-17); 2938 (2-88); 2944 (2-60)
Calder6n,M., 34 (1-1)
Calzada,J. I. et al., 868 (6-47); 7166 (1-1); 8542 (2-
45
Cambar, I., 111 (1-1)
Campbell,D. G. et al., 6254 (2-99);6341 (2-43b);6540
(7-21);6476 (2-114);6503 (2-64);6543 (2-99);6573
(2-43b); 7020 (7-50b); P20826 (7-55); P20859 (6-
11);P20868 (2-69b);P20872 (2-140);P20887 (7-6);
P20896 (2-99); P20897 (7-2); P21129, P21130 (8-
4); P21210 (7-36a);P21834 (2-140);P21918 (2-14);
P21963 (2-57); P21976 (2-27a); P22052 (6-3);
P22126 (6-9);P22271 (2-99);P22302 (2-36);P22305
(7-56a);P22313 (2-134);P22314 (2-13);P22315 (6-
3); P22336 (7-56a);P22370 (7-52);P22389 (2-134);
P22404 (2-36); P22457 (7-56b); P22470 (2-36);
P22563 (6-29)
Capus,F., 147 (2-114)
Carauta,J. P. P., 973 (7-56b); 2319 (7-85); 3391 (2-
11)
Cardona,F., 2489, 2555 (6-10)
Carnevali,G. et al., 1480 (7-9)
Carrasquilla,L., 363 (1-1)
Carreira,L. et al., 724 (7-66); 850 (7-73)
Carvalho,A.M. de et al., 193 (6-25); 194(6-52.1); 216
(6-51b); 353 (2-80a);469 (7-20); 609 (1-1); 647 (7-
82); 858 (6-32); 877 (6-51b); 1046 (7-20); 1077 (4-
4); 1106, 1116 (6-52.1); 1366 (2-11)
Casari,M. B. et al., 277 (6-25); 1073 (2-18)
Castellanos,A., s.n. INPA27546 (6-16); 24054 (7-54);
24110, 24150, 24180 (6-27); 24185, 24280 (2-41);
25263 (2-43a)
Castilla,A., 152 (2-27a)
Castillo,A., 31 (6-6a); 422 (2-27b); 692, 705 (7-36a);
736, 742 (7-56a);1311(3-4); 1407, 1417(2-77); 1526
(7-25); 1535 (6-3b); 1592 (2-83); 1798, 1848 (7-25)
Castro,J. H., s.n. (2-32)
Castro,M., 10 (2-45)
Cavalcante,P. B., 441 (7-56a); 1789 (6-3b); 2309 (2-
81); 2441 (7-56b); 2065, 2276 (6-15); 2477 (7-25);
2764 (6-15); 2768 (6-41);2809, 2889, 2951 (7-56a);
3041 (6-36); 3088 (2-69b); 3202 (6 sp.); 3276 (2-
27b); 3308 (3-6); 3371 (7-56b); 3373 (7-20); 3378
(2-43a)
Cedillo T., R., 577 (2-45)
Cerrato,B. C. A., 141 (1-1)
Cesar,A., 1 (7-73)
Cezario& M. G. Vieira, s.n. INPA 20606 (6-15)
Champagne,H., 49-1 (2-130); 49-2 (2-86), 49-3 (2-
109);49-4 (2-86)
Chan, C. et al., 813, 1483, 1615 (1-1)
ChavelasP., J. et al., ES2059, ES2124, ES2144, 2600
(7-56b);ES2818 (7-36); ES2960, 3343 (2-15)
Chiang,F. et al., 510 (2-45)
Christenson,G. M. et al., 1155 (2-114); 1161 (2-27);
1399 (2-69b); 1430 (2-140)
Chrostowski,M. S., 70-398 (7-73)
Churchill,H. W. et al., 4345 (7-56b)
Cid, C. A. et al., 59 (7-52); 228 (7-56a); 278 (2-88);
317 (7-56a); 365 (3-4); 428 (2-75); 431 (7-52); 487
(7-8);489 (7-56a);516 (3-4);640 (7-11);704 (7-56a);
109
860 (7-11); 1059 (6-41); 1131 (7-2); 1139 (2-93);
1153 (2-134); 1201 (6-29); 1230 (2-134); 1281 (6-
15); 1433, 1549 (7-56b); 1579 (6-3a); 1627 (4-3);
1644 (7-50b); 1645 (7-56a); 1671 (7-56b); 1769 (6-
3a); 1797 (7-56a);1899(7-1); 1912 (7-56a);1916(2-
43b); 1971 (7-15); 2089 (2-134); 2207 (6-3a); 2217
(7-56a); 2261 (6-32.1); 2333 (6-3a); 2437 (7-50a);
2534 (2-64);2542 (7-56a);2545 (7-66);2665, 2792,
2798 (7-56a); 2946 (7-66); 3048 (2-69a); 3184 (6-
29); 3211 (6-36); 3226 (2-69a); 3296 (6-29); 3349
(2-75);3427 (7-62);3509 (2-52);3514 (2-69b);3555
(6-3b);3556 (2-52);3575 (2-27a);3633 (2-129);3650
(2-27a); 3688 (2-129); 3696 (6-3b); 3728 (2-124);
3747 (2-129); 3778 (6-lb); 3810 (6-3b); 3856 (6-9);
3869 (7-9); 3948 (2-69b); 3895 (7-6); 3949 (7-56b);
3953 (2-69a);3993 (7-2);4003 (7-56b);4088 (6-36);
4162 (7-52); 4181 (7-56b); 4252 (2-69b); 4258 (6-
3a);4340, 4434, 4474 (2-41);4500 (7-56b);4530 (7-
73); 4536, 4574 (7-36a); 4600, 4656 (7-56a); 4661
(7-36a); 4682 (7-73); 4685 (7-56a); 4724, 4729 (7-
56b);4768, 4831, 4956 (7-36a);4995 (7-56a);5065
(6-la); 5259 (7-5); 5336 (6-7); 5343 (2-121); 5589
(2-83); 5662, 5860 (2-43b); 5956 (2-16); 5982 (7-
50b); 6052 (3-16); 6093 (6-27); 6103 (3-16); 6135
(7-20);6137 (6-27);6170 (2-124);6218 (7-56a);6228
(2-30);6232 (7-54);6253 (2-37);6318 (2-124);6341
(7-15);6368 (6-3c);6380 (2-37);6434, 6477 (2-114);
6501 (2-141); 6503 (2-27a);6505 (2-141); 6521 (2-
27a); 6571 (7-66)
Clark,H. L., 6491 (3-4); 6661, 6737, 6740, 6753 (8-
3); 6963 (2-69a); 7033 (2-82); 7113 (2-27a); 7165
(2-39);7250 (2-69a);7290 (7-8); 7337 (2-69a);7342
(2-82);7362 (2-69a);7406 (2-62);7473 (7-21); 7508
(2-69a);7527, 7629 (2-36);7669 (3-6);7778 (2-69a);
7779 (7-8); 7783 (2-57); 7789 (7-8); 7811 (2-39);
7817 (2-69b);7846 (2-57);7924 (2-39);7926 (2-76);
7926 (2-94); 7928 (2-24); 7981 (2-27a); 8019 (2-
129); 8032 (2-27a); 8057, 8063 (8-3); 8082 (3-la);
8157 (6-la); 8211 (8-3); 8232 (2-24); 8327 (7-8)
Clausen,R. T. et al., 6238 (2-1)
Coelho, D. et al., 356, 380 (2-52); 395 (6-3a);406 (2-
27a); 411 (6-3a); 708 (2-119.1); 817 (7-8); 843 (2-
36);861 (2-64);INPA5332 (2-124);s.n. INPA51497
(7-73); INPA53263 (7-40); INPA53289 (7-56);
INPA53292(6-3b);INPA53295(2-23);INPA53339
(2-43b);INPA92490 (7-6)
Coelho, L. et al., 17 (7-6); 92 (2-43b); 134 (7-73); 206
(2-109); 308 (2-94); 361 (7-6); 382 (6-41); 432 (7-
6); 479 (2-69b);492 (2-57); 496 (3-6); 522 (2-60a);
533 (2-69b); 561, 572, 650 (2-81); 651 (6-29); 708
(2-119a); 835 (2-69a); 1837 (7-54); 1955 (2-43b);
1975 (7-56b); 1979 (7-56a); 1986 (2-79); 5209 (2-
69.1); INPA25903 (2-112); s.n. INPA42005 (6-21);
s.n. INPA42095 (7-25); s.n. INPA42141 (7-20); s.n.
INPA42161, s.n. INPA42171 (6-3a); INPA53292
(6-3b)
Coker,W. C., s.n. (2-1)
Collares,J. E. R. et al., 110 (7-18); 121 (2-80)
Cominote,J., 84 (3-4)
Conant,D. S., 993 (7-56a); 1100 (7-1)
Conejos,J., 24 (2-77); 28 (3-3); 43 (3-4)
Contreras,E., 10715, 10716 (7-23); 10719 (7-56b);
10729 (7-23); 10742 (2-18.3)
Coradin,L. et al., 75 (7-20); 689 (7-25); 781, 784, 785
110
(7-56b);1051 (7-25); 1080 (7-73);4876 (7-25);4880,
5022,5023, 5068 (7-56b);6434 (2-80);6471 (7-56b);
7425 (2-80)
Cordeiro,I. et al., 89 (4-1); 98 (2-91); 119 (2-69a); 147
(2-99); 175 (6-la); 285 (7-49); 289 (2-165); 300 (2-
99); 306 (3-6); CFSC6544 (2-144); CFSC6782,
CFSC7551(7-54)
Cordeiro,M. dos R., 14 (2-99); 82 (2-43b); 100 (6-20);
105 (2-43b); 153 (3-4); 159 (2-43b); 169 (7-73); 263
(2-98);468 (6-21); 497 (7-56b);535 (2-43b);626 (2-
62); 695 (6-3b);703 (7-11.1);755 (2-22);759 (2-62);
814(2-37); 1027(2-69a); 1156(2-144); 1162(2-17);
1270 (2-27b);4679 (2-15)
CorreaA., M.D. et al., 689 (7-38); 958 (5-1); 1815 (2-
5.3); 4679 (2-15)
Correll,D. S. & H. B., 35873, 42500 (2-1); 45500,
49992 (1-1); 51946, 52557 (2-1)
Costich, D. et al., 809 (7-2); 847 (7-1); 927 (2-114);
961 (6-9)
Couret, 18 (7-73); 243 (6-45); 263 (2-18); 264 (2-77)
Cowan,C. et al., 2253 (1-1)
Cox, D. K., 3304 (7-56b)
Cremers,G., 4628 (7-73); 5091 (7-69a); 5338 (7-25);
5367 (2-134);6498, 6538 (7-56a);6546 (7-74);7011
(2-114); 7711 (1-1)
Croat,T. B., 8129 (7-36a);10018 (1-1); 13246 (7-36a);
14648 (2-88); 16751 (7-36a); 17723 (6-44); 17750
(2-69b);17867(6-44);18574(7-56a);19382 (6-41.1);
19883 (2-39); 20014 (2-17); 20218 (2-69b); 20332
(6-22); 20367 (3-14); 20372 (2-43c); 20410 (7-56a);
20443 (3-9); 20663 (6-38); 20673 (2-69b); 23256,
24187, 24211 (1-1); 24289 (7-36a); 24492, 24977
(1-1); 33627 (7-36a); 39057 (1-1); 45746 (2-45);
54099 (7-21); 54147 (7-49); 51405 (7-36a); 57982
(6-36); 59199 (7-56b); 61008 (1-1)
CuadrosV., H., 2133, 2181 (7-22)
Cruz,A. et al., 21 (7-36a); 151 (7-21); 170 (7-36a)
Cumana,L. J. et al., 779 (1-1)
Curran,H. M., 2M57, 2M74 (1-1)
Daly, D. C. et al., 141 (7-56a); 149 (4-3); 158 (7-56a);
181 (2-114); 259 (7-16); 369 (2-56b); 509 (2-141);
639 (2-56b);679 (2-69a);681 (7-52);703 (7-20);709
(7-18); 814 (2-108); 820 (2-69a);831 (6-50); 833 (2-
81); 839 (2-56b); 862 (2-63); 875 (2-56b); 882 (2-
63); 898 (2-27a);992 (7-56b); 1028 (7-24); 1056 (7-
56b); 1115 (6-1); 1117 (2-114); 1342 (2-69a); 1373
(2-27a); 1374 (7-56a); 1413 (7-24); 1511 (2-69a);
1516, 1552 (2-24); 2035 (7-56a);2069 (7-36a);2928
(2-99); 3825, 3850, 3927 (2-69a);3931 (7-24); 3943
(6-la); 3968 (2-39); 4037 (2-108)
Dambros,L. A., 82, 86 (6-27); 87, 119 (2-41); 138 (7-
54)
Damiao, C., 593 (6-34); 594 (2-22); 600 (6-15); 616
(2-99); 619 (6-44); 621 (6-3b); 633 (7-50b); 656 (2-
128); 674 (2-114); 695 (2-124); 702 (6-44); 718 (7-
50b)2491 (2-69b);2513 (2-124);2527 (2-69b);2531,
2579 (6-3b);2587, 2599 (2-27a);2637 (2-137);2712
(3-4); 2784 (2-69b); 2884, 2898 (2-27a); 2904 (6-
3b); 2908 (7-56a); 2924 (6-15); 3034 (6-36); 3067
(2-124); 3071 (6-lb); 3079 (2-27a); 3081 (2-69b)
Danin, A., 76-35-5 (2-45); 77-3-6 (7-56b)
Dantas, M., 12374 (6-3b); 12382 (7-40); 12391 (2-
124); 12392 (7-29)
D'Arcy, W. G. et al., 13662 (7-56a); 14273, 14297,
FloraNeotropica
14329, 14548 (7-56b); 14559 (1-1); 14596, 14653,
15015 (7-56b)
Daubenmire,R., 95 (2-15); 392 (7-56b); 489 (2-45);
716 (7-56b)
Davidse, G. et al., 2327 (1-1); 4306 (7-40); 4358 (2-
12); 4446 (7-56a); 4466 (3-7); 5415 (2-69b); 6753,
6953 (7-29); 12132A, 12208A (2-27b); 12260A (2-
69b); 12354 (7-56b); 12361 (2-18); 12385A (3-7);
12402 (2-69b); 12423 (7-56a); 12429, 12435 (6-3b);
12517 (7-56b); 12531, 12535 (2-27b); 12538 (2-69);
12540 (2-27b); 12553 (6-3b); 12625 (2-69b); 12637,
12714 (2-27b); 12737 (6-3b); 12807, 12855 (2-27b);
13015 (6-3b); 13055 (7-56b); 13109 (2-69); 13148
(2-27); 13243,13414 (2-27b);13838 (7-56b);13852,
13944 (2-88a);13953 (2-27b); 13979 (2-134); 13981
(7-56b);14090 (2-69b);14180(7-40); 14305 (2-88a);
14327(2-60b);14366, 14393(2-27b);14536 (2-88a);
14546 (2-69b);14654 (2-88a);14708 (2-27b); 14711
(6-3b);14765, 14842(2-27b); 15107 (2-56b); 15118,
15212 (6-3b); 15292 (7-56b); 15451 (6-3b); 15490
(2-88a); 15491 (7-56b); 15541 (2-47); 15550 (7-40);
15619 (2-140); 15662 (7-56); 15665 (2-69b); 15756
(2-140); 15761(7-56a);15768(2-140); 15769 (6-3b);
15788, 15894 (2-69b); 15907, 16005 (6-3b); 16022
(3-7); 16025 (7-56b); 16027 (2-88a); 16030 (2-134);
16093 (7-40); 16150 (2-69b); 16176 (3-7); 16184 (2-
27b); 16265, 16308 (2-122); 16276 (7-56b); 16291
(3-1); 16363 (2-43a); 16420 (7-20); 16426 (2-13);
16432, 17476 (2-77); 16753 (7-29); 16889 (2-121);
16979 (7-62); 17010 (2-27b); 17032 (2-119); 17088
(2-28); 17098 (2-131); 17144 (2-119); 17219 (6-3b);
17329 (2-69b); 17387 (2-28); 17389 (2-119); 17409
(2-131); 17666 (2-141); 17721 (2-39); 17837 (1-1);
18056 (2-52); 18304 (7-50a); 18372 (7-22); 18575
(2-114); 19155, 19188, 19482 (7-36a);20227 (1-1);
20999 (7-36a);22578a, 22747 (7-21); 22903 (7-25);
22962 (7-56a); 23267 (2-45); 23705 (2-88); 27217
(6-7); 27627 (2-27a);27671 (2-140); 27730 (2-27a);
27735 (2-102);27740 (2-88);27852 (7-8);27883 (2-
27b); 27886 (2-88); 27888 (2-69b); 27951 (7-40);
30157 (2-45)
Davidson, C. et al., 5206 (7-5); 5357 (7-53); 5437 (2-
140);8516 (7-36a);9774 (7-40); 9848 (6-44); 10581
(7-56a); 10604 (6-29)
Davila, H. R. et al., 2 (7-56b)
Davis, D. H., 180 (7-25); 204 (2-39); 814 (6-43); 890
(7-56b); 1657 (6-43)
Davis, E. W. et al., 1080 (7-56b)
Davis, H. A., 15609 (1-1)
Davis, P. H. et al., 2259 (7-20);60303 (2-119.1);60326
(7-56a);60337 (6-28)
De La Cruz,J. S., 3406 (2-66)
Delascio, F. C. et al., 5534 (2-27b); 7361 (4-3); 9662
(2-140); 11135 (6-3b); 11187 (7-56b); 11190 (6-3b);
11349 (7-56b); 11350, 11409 (6-3b)
Denslow, J. et al., 76 (6-57); 79-25 (7-61)
Dept. de BotanicaUSP, 3 (7-56a)
Deward,G. 155 (2-134)
Dezzeo, N., 293 (7-56a)
Dias, A. A., 46 (2-30)
Diaz, C. et al., 68 (7-5); 288 (6-3b); 397 (6-35); 438
(2-140);638 (7-40); 916 (2-68); 944 (7-40); 1527 (6-
44); 1751-18 (7-75); 1739-118 (2-43c); 1758-32,
1758-22 (2-27a)
 List of Exsiccatae
Supplemental
Diaz, M., 2-A (6-44); 51-A (2-64); 87A (2-141); 127A
(6-44); 162A (7-41); 6-52 (7-40.1)
Dieckman,L., 318 (2-88a)
Dionisio et al., 120 (6-46)
Dios Holmquist,J. de, 60 (6-3b)
Dodson, C. H. et al., 1170 (7-35); 2945 (2-69a); 5464
(7-87); 7516 (2-5); 8655 (7-87); 8672, 12407 (2-5)
Dressier, R. L., 3322 (7-38); 3416 (7-36a); 3736 (7-
38); 4622 (2-88)
Duarte,A. P., 5420 (2-4); 5607 (2-71a)
Ducke, A., HPB1550 (7-18); MG7367 (2-129);
MG7876 (7-63); MG10563 (2-57); MG10966 (2-
43a); MG12221 (2-129); MG15289 (2-76);
MG16153, MG16260 (2-43a); MG16355 (2-79);
MG16430 (2-60); MG16525 (2-43a); RB15137 (6-
18)
Duke,J. A., 5682 (7-56); 13513 (7-36a);15256 (2-145)
Dunbar,P. L., s.n. (1-1)
Duran, A. R. Camacho, 147 (7-56b)
Dwyer,J. D. et al., 511, 592 (1-1); 1047 (2-31.1);4298
(7-56b); 5047 (2-31.1); 7324 (5-1); 10986 (1-1);
12252, 15138 (7-56b)
EcheverryE., R., 2078 (2-56)
Egler,W. A., 577 (2-36)
Ehrendorfer,F., 74108-25 (2-122)
Eiten, G. & L., 3253 (2-41); 4665 (4-2); 8389 (6-27);
8461 (7-19); 8503, 8734 (7-20); 8904 (2-27a);9764
(6-27)
Elburg,J., LBB9373(6-18); LBB9387(3-1); LBB9440
(2-134); LBB9815 (7-56a)
Elcoro, S. et al., 210 (7-56a); 223 (2-56); 233 bis (2-
69a)
Emmerich,M., 4579 (2-41); 4601 (7-54); 4638 (7-20)
Emygdio,L. (see Mello Filho, L. E.)
Encarnaci6n,F., 970 (2-61); E1060 (2-69a); 1243 (2-
23); 1254 (6-44);25033 (2-27a);25037 (6-3b);25041
(2-27a);26124(7-41); 26183 (7-36a);26201 (2-121);
26303 (6-38); 26369 (2-27a); 26392 (2-69a); 26409
(2-98)
Erlanson,C. O., 78 (1-1)
Espejel,I., 5-3, 404 (1-1)
Espina,J. et al., 205 (6-36); 207 (3-6)
Espinal,M., 163 (7-56b)
EspinalT., S. et al., 4110 (2-4); 4537, 4608 (2-48)
Estrada,U. et al., 11 (2-45)
Euponino,A., 195 (7-64);212 (7-13);309 (6-5 lb); 361
(7-50); 405 (6-51b)
Faircloth,W. R., 1077 (2-1)
Falcao, M. et al., 143 (2-43b); 192 (6-15)
Fanshawe,D. B., 621, 638 (6-6)
Farifias,M. et al., 299 (2-88); 330 (7-21); 471 (7-8);
658 (2-69b);661 (2-27a)
Farney,C., 1101 (6-26)
Fennell,J. L. et al., 955 (1-1)
Fernandez,A. et al., 1021 (7-74); 1618 (7-25); 1706
(2-27a); 1738 (7-56b); 1779 (2-27a); 1783 (6-3b);
1784 (2-27a);2855, 2872 (7-56b);2873 (7-21); 3022
(7-40); 3531 (2-69a); 3561 (3-1); HPB2517 (7-18);
HPB3075(1-1);HPB3453(2-44);HPB3782(2-43a);
HPB6908 (6-51a); HPB8066 (7-56b); HPB7221,
HPB8811, HPB9032 (7-18); HPB9373 (2-37);
HPB10015 (4-2); HPB10016 (7-18); HPB10051 (2-
30);HPB10723(6-27);HPB10724(7-20);HPB10595
(2-30); HPB10928 (2-27a);s.n. (1985) (2-28)
111
Fernmndez C., J., 3844 (6-27)
Fernandez,M. M. et al., 5 (2-80)
Fernmndez P., A. et al., 6124 (2-24);6975, 7005 (7-21)
Ferrari,G., 1231 (2-27a); 1272 (2-12)
Ferreira,E., 58-320 (7-55.1)
Ferreyra,R., 4603, 4636 (7-56b)
Feucht,S. P., 743 (1-1)
Feuillet,C., 487 (7-74);985 (7-52); 1089 (2-124);2303
(2-86); 2314 (2-52)
Filho, L. C. de O., 27 (7-18); 28 (7-20)
Figueroa,V., 122 (7-56b)
Flores,J. S., 8256, 8322, 8800, 9323, 9867 (1-1)
Florschiitz,P. A. et al., 2810 (7-1 lb)
Foli, D. A., 49 (3-4); 61/79 (2-146); 76/79 (7-13); 80
(7-70.1); 89 (7-70a); 120 (2-6); 228 (2-78.1)
Folsom, J. P. et al., 1435 (2-5.3); 1932 (7-36a); 1949
(5-1); 1996 (2-31.1); 2774 (1-1); 3508 (2-88); 3539
(7-36a);5672 (6-47); 6781 (7-36a)
Fonseca,S., 350 (2-30)
Fonseca,W. N., 129 (4-2)
Fontella,J., 2275 (1-1)
Forero,E. et al., 4005 (2-58); 4410, 4424 (7-10); 4684
(7-35); 4788 (2-124); 4862 (3-15)
ForeroP., L., s.n. (1-1)
ForestService,FrenchGuiana,3511 (2-134); 3513 (6-
18); 3758 (7-25); 4237 (7-56a); 4503 (7-25); 6097
(2-52); 6231, 6440 (2-69a); 7259 (7-7); 7786, 7793
(6-2)
Foresta,H. de, 190 (2-109); 202 (2-69a);277 (2-109);
555 (7-2); 630 (7-74); 642 (7-25); 692 (2-149); 746
(7-56a);750 (2-134); 751 (2-63)
Fortinet al., 8565 (6-47)
Fosberg, F. R. et al., 29203 (6-44); 42591 (7-56b);
53830, 54087 (1-1)
Foster, R. B. et al., 2252 (2-15); 3091 (7-36a); 3725
(6-36);4043 (7-5);4074 (6-35);4299 (7-2);4385 (7-
53); 4568 (6-36); 4570 (3-14); 4683 (6-35); 4709,
4788 (7-56a); 5623 (2-21); 5774 (3-9); 5909 (2-21);
7961 (2-5); 8021 (6-36); 8733, 8926 (7-73); 10942
(2-88); 10996 (2-144.1)
Fournet,A., 445 (2-56b)
Frame,D., 201 (7-56a)
Frankie,G. W., 259C, 292C (7-56b); 398C (7-61)
Freeland,J. et al., 112 (6-47)
Freeman,O. M., s.n. (1-1); s.n. (2-1)
Froehner,C., 361 (2-39); 362 (2-64)
Fr6es, R. L., 41 (2-88); 12711/80 (3-3.1); 21321 (6-
35);22431 (6-lb); 22645 (6-la); 24434 (7-73);25483
(6-9);25505 (4-1);25928 (2-63);27517 (3-4);29044
(6-3b);29277 (6-29.1);29562 (7-24); 30435, 30654,
30688 (6-lb); 31898 (2-66);32159 (2-97);32311 (2-
63); 33653 (6-la); 34168 (2-97); 34888 (6-lb)
Funk, V. A. et al., 8415 (2-47.1)
Furlan,A. et al., CFSC7513(6-27);CFCR7138(7-20)
Garcia,C., 30 (2-45)
Garcia-Barriga, H. et al., 3431 (2-27b); 18202, 18207,
18209 (2-94); 18457 (2-129); 18528 (2-12); 18924
(7-36a);20323 (7-56b)
Gamier, 76 (7-56b);79 (6-7)
Gasche,J. et al., 28 (7-24); 103 (2-69b); 154 (7-5);211
(2-37); 216 (6-29)
Genelle, P. et al., 261 (2-1)
Gentry,A. H. et al., 385 (2-45); 388 (3-45); 5800 (2-
145);6172 (6-13.1);7605 (2-88);7723, 8364 (7-22);
112
8884 (2-95); 10418 (2-27a); 10780 (7-35); 10831 (6-
3b);10835(7-21); 10863 (2-68); 10936 (7-40); 10966
(2-24); 12905 (2-27a); 13306 (7-56b); 13439 (7-38);
14554 (7-56b); 14570 (7-25); 14624 (2-140);14641
(7-56b); 14943 (7-74); 15536 (6-3b); 15767 (2-27);
15807 (7-6.1); 16638 (3-14); 16701 (7-40); 17224
(1-1); 17482 (7-10); 18140, 18148 (7-22); 18318,
18455 (6-44); 18466 (3-9); 18527 (2-52); 18543 (7-
40); 19094(3-14); 19098 (2-21);20371 (3-14);20385
(6-41.1); 20678 (7-6.1); 20679 (7-5); 20834 (2-43b);
20842 (6-3b); 21045 (7-5); 21130 (6-15); 21209 (2-
43b); 21213 (7-40.1); 21452 (3-16); 22199 (2-45);
23387 (2-43c); 24116 (3-14); 24331 (2-34); 24962
(6-7); 25012 (2 sp.); 25057 (6-35); 25064 (3-14);
25069 (6-35); 25073 (2-124); 25193 (7-53); 25203
(7-5); 25310 (6-35); 25613 (7-21); 25665 (2-4.1);
25845 (7-56b);25866A (2-140);25913 (3-14);26030
(6-44); 26035 (2-140); 26072, 26173 (6-7); 26202
(2-69b); 26866 (7-36a); 27591 (6-41.1); 27642 (7-
62); 29704 (3-9); 30147, 30201 (2-51); 30324 (2-5);
31284 (7-56a); 31502 (2-94); 31657 (2-69a); 31701
(6-35); 31725 (7-6.1); 31733 (3 sp.); 31824 (7-50b);
31936 (7-73); 32085 (6-44); 32617 (6-47); 35478 (2-
34); 36188 (7-56a); 36227 (2-43b); 36238 (7-62);
36269 (2-43b); 36275 (7-62); 36291, 36304 (2-99);
36311 (7-50b); 36438 (7-6.1); 36440 (6-35); 36626
(7-73); 37115 (6-36); 37252 (7-56a); 38047, 38059
(7-36a); 38077 (3-4); 39243, 39318, 39329 (2-26);
39350 (2-69a); 39408 (2-128); 39412 (2-99); 39471
(2-124);40274 (2-46);40355 (2-5.1);40416 (2-124);
40467 (2-32.1); 40474 (7-56a); 41138 (3-7); 41332
(7-56b);41569 (7-53);41600 (2-69a);41608 (2-124;
41682 (2-123); 41712 (3 sp.); 41731 (2 sp.); 41895
(7-53); 41939 (3 sp.); 42074 (7-56a); 42805 (2-5);
43029 (2-144.1);43178 (6-44); 43195 (2-21);43351
(7-36a); 43403 (3-5); 43576 (6-24); 43653 (3-5);
43687 (7-46);43754 (2 sp.);43777 (2-21);43811 (6-
36); 44539 (3-5); 45582 (2-144.1); 45669 (7-75);
46170 (2-99); 46213 (2-43b); 46275 (6-36); 46277,
46278 (2-27b);46290 (2-83); 46377 (2-69a);46457
(6-lb); 46471 (7-21); 47062 (2-63); 47121 (2-27.1);
47379 (2-69b);47385 (3-1);47551 (7-22);47821 (3-
17);47961 (2-148);48461 (2-15);48513 (7-61);48953
(2-124); 48978 (6-la); 48988 (2-114); 49015 (6-la);
49045 (7-63); 49103 (2-63); 50319 (7-56a); 51140
(2-69a); 51214 (7-56a); 51396 (2-94); 51556 (2-21);
52175 (2-158)
Gibbs, P. et al., 2708 (7-54); 2738 (3-16); 2839 (7-20);
3363 (6-27); 3522 (7-82); 3524 (2-126); 3837 (2-
114); 5087 (2-80); 5165 (7-20)
Giovanni, F. di, 2 (2-69a); 77 (7-40); 80 (2-140)
Giulietti,A. M. et al., CFCR2254(2-144);CFCR3552,
CFCR6760(7-20)
Glaziou, A. F. M. et al., 5689, 7602, 9391, 13796
(3-4)
Glenboski,L., 206 (2-43c)
Godfrey,R. K. et al., 52402 (2-1); 53779, 58121 (1-1)
G6es, O. C. et al., 152 (2-143); 943, 1046 (3-4)
Goesnner,F., s.n. (6-3b)
Gomes, M. et al., 8, 92 (7-56a); 106 (7-56b); 145 (2-
37); 156 (3-4); 195 (2-37); 226 (7-56a); 381 (2-37);
432 (3-4); 447 (6-3b); 448, 476 (2-28); 477 (2-27a);
FloraNeotropica
554, 867, 1015 (7-50b); 1114 (6-la); 1252, 1479,
1484 (7-50b)
G6mez,L. D. etal., 18927(7-56b);20461 (2-95);21178
(7-56b);23057 (6-47); 23356 (2-15); 24100 (7-36a)
G6mez-Pompa,A. et al., 1283 (2-15); 3347, 3354 (2-
95); s.n. (2-88)
Gonggrijp,J. W., 284 (2-69a)
Gongora,E., 438, 518 (1-1)
Gonzales,A., 1159(2-21)
GonzalesL., L. A., 4752, 4757 (7-56b)
Gonziles-Quintero,L., 3435 (7-56b); 3505 (7-36a)
Goodland, R., 289 (7-56b); 712 (2-41); 755 (7-56b);
894 (6-27); 927 (3-16); 949 (7-18); 963 (7-54); 1074
(2-135)
Goodrum,P., 9 (2-1)
Gordon,B. L., 9 (2-15)
Gottsberger,G., 16-28782 (7-18); 16-29782, 18-29782
(4-2); 22-81182 (7-18); 15-15183 (6-21)
Gottsberger,I. S. et al., 225 (2-41); 368, 152-25771
(6-27)
Goulding,M., 29, 54 (2-23); 153 (6-44); 183 (6-23)
G6mez, L. D. et al., 23356 (2-15)
Grant,G. B., s.n. (2-1)
Granville,J. J. de, C-2 (7-74);C-17 (7-56a);21 (2-60);
C-51 (7-56a); 115 (7-2); C-143 (7-74); 488 (2-27b);
509 (7-2); 582 (2-134); 599 (2-50b); 951, 1090 (7-
56a); T1203 (7-52); 1250 (7-2); 1436, 1739 (7-73);
1740, 1791 (7-56a); 1802 (7-25); 1883 (2-88); 1946
(7-56a);2682 (6-7); 3134 (7-52); 3140 (7-73); 3249,
3673 (7-2); 3726 (2-88a);4028, 4136 (7-2);4321 (7-
25); 4606 (7-52); 4684 (2-134); 4687 (2-69a); 4700
(7-56a);4723 (7-56d);4731 (2-88);4838 (7-73);4867
(2-69a);4875 (2-99); 4979 (2-93); 5051 (7-2); 5234
(2-88);5242, 5429 (7-2);5537 (7-56a);5652 (7-50b);
5719, 5722 (2-124); 6121 (4-3); 6124, 6171 (7-74);
6181 (7-2);6183 (2-114);6259 (7-2);6271, 6571 (2-
64); 6727, 6954 (7-56a); 6993 (2-88); 7177 (7-52)
B3637 (2-77);B3704 (7-2);B3732 (6-lc); B3785 (7-
56a);B3787(2-67);B4499(7-73);B4507(7-2);B4611
(7-56a); B4673 (2-8); 4684 (2-134); B4797 (7-2);
B4849 (2-88); B4856 (7-56a); B4891 (6-2); B4906
(7-56a); B5061 (7-2); B5263 (6-18); B5376 (7-73);
B6326 (2-135); BC101 (2-109)
Grayum,M. et al., 4090 (7-61); 4971 (7-36a)
Greenman,J. M. et al., 5739, 5833 (1-1)
Grenand,P., 38 (2-99); 139, 227 (2-69a); 547 (7-20);
548, 614 (2-124); 642 (2-69b); 967 (3-2); 1506 (3-
3); 1582 (7-52); 1718 (2-134); 1774 (6-7); 1818 (2-
25); 1905 (2-69a); 2037 (7-50b); 2083 (7-73); 2138
(7-51); 2459 (2-140)
Grijalva,A., s.n. (1-1);2308 (2-15);3443, 3546 (2-45);
3760 (7-36a); 3890 (2-15)
Grubb,P. J. et al., 1045 (2-5)
Guanchez,F. et al., 108 (7-25); 251 (2-88); 252 (2-12);
396 (6-29); 409 (7-40); 423 (7-25); 510 (7-62); 720
(2-81);734, 793 (2-69a);804 (7-30);943 (2-83);965
(2-103);989 (2-81); 1098 (7-49); 1228 (2-69b); 1249
(3-6); 3258 (7-49); 3375 (3-1); 3454 (7-9); 3480 (2-
121); 3501 (2-140); 3536 (7-49); 3647 (2-99)
GuarimNeto, G. et al., 38 (7-6); 312 (2-43a)
Guedes,T. N., 311 (2-149)
Guevara,L. C. de, 1400, 2125 (2-12)
 List of Exsiccatae
Supplemental
Guimarbes,J. G., 17 (7-66); 1150, 1462 (2-41)
Gunn, C. R., 3357 (2-1)
GutierrezV., G., 638 (7-40); 2784 (2-17)
Guzman & Castro,60 (1-1); 156 (2-45); 559 (1-1)
GuzmanB., R., s.n. (2-15)
Hage,J. L. et al., 285 (7-20); 564 (7-36a); 1896 (2-11)
Hahn, W. et al., 444 (1-1)
Halle, F., 494 (7-52); 2807 (7-73)
Hammel, B. et al., 975, B1768 (7-56b);4823 (2-18.1);
11149 (2-67); 11260, 11577 (5-1); 12476 (2-95);
14137 (5-1)
Hamilton, C. et al., 2965 (7-56b); 3281 (7-36a)
Hansen, B. et al., 9268 (7-83)
Hanke, W., 73 (6-4)
Harley,R. M. etal., 15752(6-25);16097(2-114);17120
(1-1);17370(7-20);17401(2-88):18067(1-1);18256
(7-13); 18581 (7-20); 18585 (7-56b); 19058 (7-18);
19083 (6-20);20074 (7-20);21645 (2-30);21649 (7-
54); 21735, 21751 (2-80); 21779 (7-18); 22240 (7-
56b); CFCR7352(4-4); CFCR7515(7-18)
Harling,G. et al., 19409 (7-35)
Harriman,N. A. 16126 (7-56b)
Harschberger,J. N., s.n. (2-1)
Hartmann,R. L., 12005 (7-36a); 12178 (7-56a)
Hartshorn,G., 809 (7-61); 886 (2-15); 1287 (6-47);
1523 (7-61); 1625 (2-88a); 1803 (7-56b); 1850 (6-
47); 1878 (7-61); 2414 (7-56a); 2659 (7-41); 2662,
2670 (2-27a); 2742 (7-50b)
de Hass Sr.et al., 136 (7-54); 152 (2-126); 154 (2-43a);
229 (2-126); 232 (7-70)
Hatschbach,G., 17886, 20216 (7-85); 21790 (2-41);
24241 (7-54);24595 (2-30);24632 (6-27);25298 (7-
66); 29890 (7-54);30202 (7-20);31917 (3-16);32418
(2-30); 32424 (7-20); 33215 (2-30); 34633 (2-41);
34674 (2-30);34686 (7-19);35405 (7-20);38466 (7-
56b); 38936 (6-20); 40137 (2-37); 41243 (7-20);
41689 (7-54); 43158 (2-114); 43760, 43796 (2-41);
44093 (7-20); 44112 (7-18); 46750 (1-1); 47027 (7-
31);47033 (1-1);47303 (3-16);47791 (7-31);48068
(2-134); s.n. (2-144)
Hazlett, D., 3270 (6-47)
Heithaus,E. R., 48 (7-56b)
Henderson,A. et al., 267 (7-53); 278 (7-14); 343 (7-
52); 359 (7-78); 412 (3-4); 448 (7-56a);477 (6-3b).
Hendrix,L., 196 (7-5)
Hensold, N. et al., SPF20850 (6-27)
Herb. Forestaldel Peru, 125 (2-17)
Heringer,E. P. et al., 1247, 1430, 1492, 1498 (7-54);
1626 (7-20); 1631, 1687 (7-70); 1761 (2-43a); 1770
(7-54); 1782 (7-70); 1815 (2-41); 1894 (7-20); 1902,
1918 (2-43a); 1923, 1934 (7-54); 1957 (7-70); 2067
(7-54); 2243 (7-20); 2279 (7-66); 2419 (7-70); 2439
(2-126);2760 (3-4);2813 (2-132);3200 (2-27a);4527,
4584, 4805 (7-54); 5021, 5114 (7-20); 5197 (7-70);
5324 (2-43a); 5399 (7-70); 5409 (2-114); 5445 (6-
27); 5598, 5604 (2-27a);5625 (2-41); 5628 (2-27a);
5696 (2-41); 6258, 6367 (3-16); 6980 (2-43); 7079
(3-16); 7121 (7-20); 14074 (2-114); 14098, 15109
(7-54); 15181 (2-43a); 16852, 17520 (7-20); 17533
(6-27); 17658, 17792 (7-54); 17837 (2-27a); 17856
(7-54); 18075 (3-16); 18514 (7-54); 18549 (6-49)
HeringerSalles, A. E., 170 (7-56b)
113
Hermann,F. J., 11001 (7-40)
Hemrndez, H. S., s.n. (1-1)
Hernmndez,J. J. et al., 244 (7-34)
Hernndez, L. et al., 112 (6-8); 128 (2-107); 160 (3-1);
187 (2-69a); 188 (2-43a); 304 (2-69a); 337 (7-25);
405 (2-113); 442 (3-4)
Hemnandez, R., 83176 (2-45)
Heyde, N. M. et al., 179 (7-20)
Hilario, L., 32 (7-56b)
Hill, S. R. et al., 2421 (1-1); 12784 (7-9); 12945 (7-
56a); 13163 (2-69.2)
Hilty, S., A-1 (6-47.1); 0-1 (2-32.1)
Hoehne, W., 6171 (2-144)
Holdridge,L. R. etal., 2517 (7-36a);6457 (2-15);6526
(7-38)
Holm-Nielsen, L. et al., 19678 (7-56b); 20051 (6-44)
Hoist, B. et al., 2220 (7-49); 2235 (2-56); 2403 (2-68);
2592 (7-56b);2665 (2-91); 2669 (6-10); 2788 (1-3);
2813 (2-124); 2817 (2-91); 2839 (2-135)
Honda, M. et al., s.n. (2-69a);s.n. (6-37)
Hoock, D. K., s.n. (7-18); s.n. (7-25)
Hopkins, M. J. G. et al., 508 (7-56a);733 (3-4)
Howard,R. A. et al., 18133(7-36a);18538(1-1); 18675
(1-2); 18881, 19004, 19389 (7-36a); 19406, 20001
(1-2)
Huashikat,V., 622 (2-10);663 (7 sp.);783, 946 (7-24);
1425 (2-81)
Huber,O. et al., 442, 443 (2-24); 450 (7-68); 524 (2-
69b); 578 (6-56b); 598 (7-56b); 692 (3-4); 1021 (1-
1); 1217, 1243 (7-68); 1345, 1425 (7-56b); 1470 (7-
9); 1553 (2-119); 1561 (6-3b); 1574 (2-119); 1580
(2-27a); 1628 (2-131); 1669 (2-69b); 1788, 1805 (2-
119); 1868 (7-71); 2035 (2-131); 2355 (7-49); 2439
(7-29);2481 (2-68);2744 (2-88a);2747 (6-3b);2755
(2-119);2757 (6-3b);2763 (7-49);2821 (2-119);2870
(2-39); 3037 (7-49); 3038 (6-3b); 3048 (7-49); 3078
(7-56b); 3083 (2-140); 3165 (2-131); 3205 (7-29);
3229 (7-59);3253 (7-29);3263 (7-59);3274 (2-131);
3276 (2-119); 3280 (6-3b);3288 (7-59); 3298, 3333
(2-131); 3337 (7-25); 3344 (2-27b); 3366 (2-27a);
3397 (2-119); 3441, 3449 (7-29); 3466 (7-21); 3469
(7-29);3476 (7-59); 3589 (7-29);3663 (2-131);3678
(7-29); 3685 (2-28); 3690, 3740 (2-119); 3853 (7-
59); 3886 (2-68); 3914 (2-131); 3932 (2-68); 3960
(2-131);4093 (2-68);4112 (6-3b);4114 (7-211);4735
(2-88);4744 (7-56b);4760 (6-3b);4792 (7-56b);4795
(2-83);4871 (7-59);4899 (2-88);4921 (2-131); 5011
(7-8); 5079 (7-59); 5086, 5151 (2-68); 5162 (2-131);
5163 (7-29); 5179 (7-56b); 5185 (2-83); 5191 (2-
27b); 5314, 5343 (2-68); 5360 (7-8); 5412 (2-68);
5523 (2-131); 5577 (7-29); 5578 (2-68); 5820 (2-
27a);5821 (2-119);5834 (2-88a);5847 (2-105);5870
(2-119);5944,6076,6088 (2-131);6155 (7-29);6210
(7-21);6287 (7-56b);6288 (3-7);6326 (6-36a);6670,
7277 (7-49); 8231 (2-56); 8288 (7-20); 8399 (7-2);
8433 (2-105); 8448 (7-56b); 8554 (2-56); 9138 (2-
69a); 9152 (6-8); 9238 (2-69a); 9396 (7-21); 9655
(3-10);9696 (7-49);9899 (7-21);10253(7-49);10286
(7-21); 10313 (7-30); 10680 (7-8); 10682 (2-119a);
10852 (2-56); 10875 (7-30)
Huft, M. et al., 1954, 1982 (7-36a)
Hume, H. H., s.n. (2-1)
114
Hunt, D. R., 5506 (3-16)
IbarraM., G., 660, 857 (6-47)
Irwin, H. S. et al., 6070 (2-80); 18231 (7-20); 21229
(7-56b); 25543, 25651 (3-16); 26068 (7-54); 26586
(2-27); 27405 (2-126); 28029 (2-144); 30786 (4-4);
30895 (2-80); 30916 (4-4); 30917 (2-80); 31037 (7-
56b);31065 (4-4);31363, 31376, 31563 (4-2);47423
(2-60); 48589a (7-56d); 54990 (2-69.1)
Isern,J., 1179 (6-44)
Jacquemin,H., 1403 (7-56a); 1408 (2-109)
Jangoux,J. et al., 6 (2-134); 30 (7-24); 40 (4-3); 52 (2-
81); 80 (2-27a); 105 (6-8a); 152 (6-lc); 273 (7-56a);
296 (7-24); 372 (7-56b); 375, 479, 495 (2-27a);644
(7-24); 656 (7-56a); 687 (2-134); 900 (7-56b); 973,
977 (7-56a);1008 (7-56b);1011 (6-lc); 1159 (7-56a);
1519(2-134);1604,1629 (7-56a);1691(6-3a);10111
(7-29)
Janssen,A. et al., 464 (2-37); 469 (7-73); 478 (7-53);
523, 545 (7-66); 546 (7-24); s.n. on 17 VI 1980 (2-
43b)
Jaramillo,J., 6672 (2-4.2); 6861 (2 sp.); 6960, 7413,
7527 (2-4.2); s.n. (2-48)
JaramilloM., R. et al., 8257 (2-48)
Jenssen,J., E., 41fr, 54fr (2-69a)
Jesus, J. A. de, 505 (6-51b); 594 (7-20)
Jim6nez-Saa,H., 1207 (7-50b); 1246 (7-36a); 1260 (2-
27a); 1293 (7-22); 1294 (2-27a); 1518 (3-4); 1526
(2-134); 1543 (3-1); 1540(2-112); 1547 (2-88); 1549
(2-108.1); 1556 (6-1); 1572 (7-42); 1577, 1586 (2-
77); 1592 (2-88); 1599 (4-3); 1608 (2-72); 1612 (2-
88); 1642 (2-72); 1643 (2-43a); 1652 (2-69a); 1656
(3-11);1673(2-108);1683 (2-98);1688(2-117);1691
(2-112); 1692 (2-134); 1693 (2-108); LBB14332 (4-
3)
Judziewicz,E. J., 4481 (7-56b)
Juncosa,A., 1394 (2-4)
Jones, G. C. et al., 3418 (2-32)
de Jong et al., 15784 (2-77)
Judd, W. S., 1330 (1-1)
Kalloo, M. B., B.240 (2-86)
Kayap,R., 49 (7-56a); 674 (6-36)
Keel, S., 218 (2-27a);222 (6-3b); 304 (7-24)
King, S. R., 426 (7-40)
Kirkbride,J. H. et al., 1000 (2-45); 1133 (7-36a); 1365
(7-36); 1447, 1448, 2527 (7-36a); 2915 (7-50a)
Klug, G., 2503 (6-3b)
Knapp,S. et al., 1255, 1896, 2271, 2785 (7-56b);2786
(7-18);2876 (7-25);2974, 3017, 3270, 3368 (7-56b);
3525 (7-80); 3935 (7-36a)
Kosei, I., 10 (7-56a)
Koyama,T. et al., 7292 (7-56b)
Kral,R., 31346, 35228, 35683, 35691 (2-1)
Krieger,P. L. et al., 12609(2-27a);12766 (6-3b);12795
(6-29); 12228 (6-3b); 12256 (7-73); 12816 (2-129)
Krukoff,B. A., 5822A (8-4); 6227, 6361 (2-81); 6381,
6635 (6-6.1); 6709 (6-3b); 7088 (2-17)
Kruse,H., 104 (2-45); 261 (2-9); 625 (2-16); 1256 (2-
9); s.n. (6-47)
Kubitzki, K., 75-53 (2-96b); 75-54 (6-44); 75-85 (6-
3b); 79-241 (7-25); 21726 (2-140); 85-30 (3-3)
Kuhlmann,J. G., 154 (3-4); 208 (2-146).
Kuhlmann,M. et al., 58 (6-4); 92 (6-la); 161 (6-lc)
FloraNeotropica
Labroy,s.n. (6-2); s.n. (6-9)
Lacerda,P. et al., 95 (7-56b); 137 (2-52)
Laguna,A., 16, 41 (7-36a)
LamonicaFreire,E. M. de, 15 (7-66)
Landrum,L. R., 4114 (7-85)
Lane, F., s.n. (7-54)
LannaSobrinho,J. P., 280 (6-25); 346 (6-16); 385 (7-
54); 642 (2-70); 686, 890 (3-4)
Lao, E. et al., 10 (2-95)
Lao Magin, R., 61, 62 (2-39); 72 (7-75); 103 (2-45);
5059 (2-43c)
Laughlin,R. M., 292 (2-45)
Lawesson,J. E., 43446 (6-36)
L.B.B. (LandsBosbeheerSuriname),11049 (7-15)
Leeds, A. N., 432 (1-1)
Leeuwenberg,A. J. M., 11837 (2-69a)
Leitao-Filho,H. F., 2074, 2091, 2161, 5742 (2-41);
7980 (7-20); 8281 (2-41); 12476 (7-54); 13118 (7-
85)
Leite, I. et al., 14 (6-44)
Lems, K., 5250 (2-130); 5272 (2-69a); 5373 (3-1);
64021702 (2-135); 650127 (7-61)
Lent, R. W., 2258 (2-88); 2319 (7-36c)
Le6n, H., 632 (2-45)
Lescure,J. P., 307 (7-56a); 390 (7-50b); 708 (7-18);
758 (2-65)
Lewis, G. P. et al., 1142 (6-20); CFCR6988(4-4)
Lewton,F. L., s.n. (2-1)
Liesner,R. L. et al., 370 (7-36a);3016 (2-15); 3374 (7-
8); 3580 (6-lc); 3591 (7-21); 3654 (2-69c); 3678 (7-
62); 3897 (6-36); 3940 (7-6); 3948 (7-40); 3970 (7-
5);4052 (6-lc); 4104 (2-69a);4191 (2-140); 5223 (7-
36a); 5572 (2-27b); 5780 (7-25); 5876 (2-77); 5885
(7-56a);5921 (2-77); 5957 (7-25);6087 (7-21); 6130
(7-62); 6378 (7-5); 6492 (6-36); 6530 (2-140); 6716
(7-62); 6903 (2-68); 6917, 7014 (2-69b); 7069 (6-
lb); 7136 (2-69b); 7209 (6-lb); 7236 (7-49); 7350
(2-39); 7451 (7-5); 7581 (7-21); 8466 (7-62); 8530
(7-5); 8550 (7-21); 8615 (2-69a);8621, 8638 (2-24);
8715 (7-40); 8742 (7-21); 8851 (2-69b); 8875 (2-
112); 8907 (2-27b); 8913 (6-13); 8917 (7-40); 9050
(7-55); 9107 (8-3); 9114 (2-68); 9143 (2-24); 9177
(7-36a); 9361 (2-43b); 9392 (7-22); 9434 (7-36a);
9507 (2-68);9538 (7-56a);9563 (6-3b);9580 (7-56);
9584 (2-76); 9606 (2-23.1); 9633 (7-102); 9887 (2-
114); 10404 (7-56a); 10424 (7-56b); 11009 (2-43a);
11027 (7-56b); 11085 (2-122); 11095 (7-36a);11170
(7-56a); 11195 (2-13); 11219 (7-25); 11293 (7-25);
11343 (2-122); 11345 (7-20); 11442, 11455 (7-56a);
12437 (3-7); 12471 (7-36a); 13063, 13354 (7-22);
13339 (2-43a); 13660 (7-36a); 13990 (2-68); 14051
(2-129); 14060 (2-77); 14070 (7-73); 14445 (7-36a);
16615 (7-30); 16624 (2-68); 17107 (3-1); 17231 (7-
56b); 17576 (7-25); 17847 (7-73); 17927 (2-83);
18344 (7-21); 18626 (7-43); 18666 (7-25); 19484 (2-
69a); 19507 (2-13); 19612 (7-56a); 19724 (7-49);
19796 (7-56a); 19852 (7-49); 19919 (1-3); 20100 (7-
56a);20133 (7-21);20192 (1-3);20517 (7-56a);20527
(2-56); 20562, 20627 (2-81); 20648 (2-56); 20675
(7-30); 20744 (2-135); 20811 (7-25); 20888 (2-56);
20943 (2-124); 20952 (2-81)
Lima, A., 390-68 (2-141)
SupplementalList of Exsiccatae
Lima, D. A., 65-4367 (7-50); 67-5059 (7-54); 68-5458
(7-18); 69-5595 (7-50)
Lima, D. P., 13083 (3-11)
Lima, E. et al., 45 (2-81)
Lima, H. C. de et al., 415 (2-144); 1019 (7-20); 1039
(7-54);1040 (7-20);1600 (2-69a);2178 (2-127);2275
(2-114); 2683 (3-4)
Lima, J. et al., 147, 160, 208 (7-56a);231 (2-45); 269
(7-56a);493 (2-18); 550 (7-56a)
Lima, J. P. de S., 89 (2-41); 101 (2-28); 104 (7-54)
Lima, R., 105 (2-114)
Lindeman,J. C., 64 (7-7); 162 (3-1);223 (6-7); 235 (7-
73); 257 (2-88); 329 (7-73); 384 (7-56a); 479 (6-4);
503 (7-56a); 525 (2-67); 531 (7-73); 677 (7-56a);
7947 (2-18); LBB15315 (2-63)
Lino, A. M., 9 (6-26); 45 (2-78.1); 71 (2-71)
Liogier,A. H. et al., 19025, 28174 (7-36a);28322 (7-
83); 29196, 29296 (1-1); 31856 (7-36a); 32784 (7-
83); 34476 (7-36a); 35577 (2-44); s.n. (1-1);
Lisboa, P. et al., 235 (7-73); 481 (2-75); 522 (7-73);
595 (2-114); 621 (2-141); 630 (6-3b); 712 (7-54);
1132(3-4);1205(7-56b);1377(2-24);1382(2-121.1);
1485 (2-27a); INPA52980, INPA53023 (7-56a);
INPA53045 (2-43b); INPA53135 (7-56a);
INPA53161 (2-99); INPA53215 (3-4); INPA53233
(2-37); INPA53235 (2-30); INPA53238 (2-28)
Little, E. L. et al., 443 (7-36); 13775, 20668 (1-1);
21014 (3-17); 21659, 21660, 23787, 25013 (1-1);
25020 (7-22); 25052 (2-88); 25053 (7-56b); 25058
(7-22); 25070 (7-36a);25101 (7-22); 25194 (7-36c);
25307 (7-36a); 25313 (7-88); 25314 (3-36c); 25326
(7-56b);25386 (7-23)
Lizot, J., s.n. (2-81); 66 (2-68); 75/46 (2-43b)
Lleras,E. etal., P16597 (7-56b);P16639 (6-34);P16655
(8-4);P16665 (6-3b);P16893 (6-36);P16962 (6-35);
P16977 (2-62); P17025 (7-6); P17058, P17142 (7-
62); P17196 (3-5); P17216 (7-95); P17270 (2-43c);
P17294 (6-41.1); P17392 (6-16); P17397 (8-4);
P17483 (6-29); P17488 (7-40); P17501 (2-124);
P17508 (7-73); P17561 (7-1); P17564 (7-56a);
P17569 (7-8); P19571 (2-57); P19665 (2-69b);
P19676 (7-6)
Lobo, M. G. A. et al., 3, 38, 70, 120, 163 (7-56b); 174
(6-15); 184 (2-114); 270 (4-3); 314 (7-24); 1102 (2-
43a)
L6pes-Forment,W., 1356 (6-47)
Lopes, M. A. et al., 113 (6-6.2); 176 (6261) (2-114)
Lopez-Palacios,S. et al., 4467 (6-3b); 4479 (2-12)
Loureiro,A. et al., s.n. (2-28); s.n. (2-29); s.n. (2-43);
s.n. (2-69a);s.n. (2-81);s.n. (3-4);s.n. (6-3a);s.n. (6-
4); s.n. (6-9); s.n. (6-11); s.n., s.n. (6-15); s.n. (6-28);
s.n. (6-29); s.n., s.n. (7-73); INPA37544 (7-56b);
INPA37629(7-2);INPA37826 (2-27a);INPA37848
(7-56a); INPA38099 (3-2); INPA38929 (2-28);
INPA38985 (7-56b); INPA39510 (7-56a); INPA-
47911, INPA47955 (2-119.1); INPA47967 (6-4);
INPA47973(2-27a);INPA48112(2-57);INPA48136
(3-4); INPA48139 (2-43b); INPA48149 (2-69b);
INPA48160, INPA48165 (2-43b); INPA48206 (2-
57); INPA48600 (2-140)
Loureiro,R. L., 11 (2-18)
Lourteig,A., 2340 (7-85)
115
Lowrie,S. R. et al., 115 (7-24); 284, 302 (7-56a);483
(2-141); 545, 567 (7-66); 568, 706 (7-56a)
Luetzelburg,P. von, 28349 (2-44)
Lugo, H. S., 1039, 1054 (2-144.1); 2144, 2160, 2202,
3230 (7-56a); 3491, 3576 (6-36); 3681, 3715, 3775
(7-56b);4286 (7-28)
Lundell,C. L., 13023 (2-32.2)
Luteyn,J. L., 8622, 8979, 8997 (6-36); 9234 (7-36a);
11506 (7-83)
Maas, P. J. M. et al., 386 (2-98); 391, 1757 (7-56a);
3517, 3576 (6-15); 3656 (7-56b);3799 (7-73); 3822
(2-77); 3817, 3836, 3872 (7-25); 4527 (7-21); 5187
(2-106);5171 (7-68); 5490, 5544 (7-25) 5838 (2-56);
P12658 (2-114); P12696 (7-73); P12762 (3-2);
P12846 (2-22); P13041 (7-56a); P13076 (7-40);
P13107 (7-5); P13134 (7-75); LBB10811 (2-134)
Maasola,J., 9 (7-22)
MacBryde,B. et al., 1376 (6-36)
MacDougall,T., s.n., s.n., s.n. (7-56b)
Macedo,M. et al., 150 (7-54); 235 (2-28); 282 (2-41);
480 (2-56a);495 (6-21); 553 (7-20); 609 (2-37); 846
(3-15); 1169 (2-11); 1297, 1395 (7-54); 1629, 1630
(2-28)
Macedo,R., s.n. INPA55750 (6-50)
Maciel, A., 124 (2-28); 128 (7-54); 326 (7-19)
Maciel, G., 5 (1-1)
Maciel, U. N. et al., 72 (7-56b); 83 (6-3a); 88 (7-24);
127 (7-50a); 133 (2-27a); 166 (6-3a); 260, 303 (7-
56b); 321 (4-1); 477 (2-69a); 717 (2-8)
Madison, M. T. et al., 135 (7-56a); 175 (2-69b); 195
(2-88); PFE211 (6-3b); PFE303 (7-2); 358, 650 (7-
62); 6175 (2-69b); 6195 (2-88); 6211 (3-3b); 6303
(7-5)
Magnago,H., 107 (2-124); 112,230 (3-16);272 (2-41);
s.n. INPA58151 (2-129)
Maguire,B. et al., 30975 (6-3b);37707 (7-56b);47080
(2-36)
Maia,L. A. et al., 26 (7-56a);112 (2-69b); 161 (7-56b);
189 (7-24); 238 (2-129); 458 (2-35); 486 (3-6); 570
(7-55); 663 (7-55.1); 725 (2-129)
Makrinius,E., 691 (7-56b)
Malpica,A. A., 1 (2-17)
Mantovani,W., 993 (6-27)
MarcanoBerti,L. A. et al., 119 (3-4); 211 (2-99); 284
(2-77); 575 (2-12); 652 (2-88); 682, 687, 792 (2-77);
861 (7-36a); 896 (7-22); 918 (7-36a); 1523 (3-7);
2603 (3-4); 2521 (6-3b);2617 (2-73); 2620 (7-56b);
2858 (2-27); 2951 (7-22); 3029 (7-36a);21-1-77 (3-
1);67-1-77, 52-2-77 (2-39); 56-2-77 (7-25);67-2-77
(1-1);77-2-77 (3-4); 13-3-77(7-25);6-10-78 (2-114);
64-979 (2-69a); 86-979 (8-3); 164-979 (3-7); 468-
979 (7-36a); 19-980 (6-la); 41-980 (2-27b); 54-980
(7-56b);58-980 (3-7);60-980 (7-56b);43-981 (2-68);
47-981 (3-4); 89-981, 99-981 (2-69a); 135-981 (7-
49); 178-981 (3-7); 200-981, 216-981 (2-43a);290-
981 (2-12); 982-157 (7-36a);983-018 (2-114); 983-
022 (3-7); 983-075 (2-43a)
Marinho,L. R., 125 (2-27); 134 (2-64); 158 (2-26);248
(6-35);267 (2-99); 382 (2-129);395 (2-69b);405 (2-
57); 447 (3-6);472 (2-69b);473 (2-57); 502 (2-69a);
561 (7-5)
Marshall,S. A. et al., 6562 (1-1)
116
Martin,R. T. et al., 1329 (7-56a); 1380, 1672 (2-1);
1676 (6-9); 5387 (7-56b)
Martinelli,G. et al., 80 (6-25); 372 (7-29); 2189 (4-5);
4114 (6-25); 6121 (2-71a); 6789 (7-42); 6859 (2-
135);6904 (7-25); 7052 (6-32.1); 7095 (6-3b);7156,
7166 (2-134);7177 (3-4);7226 (2-99);7231 (2-134);
7234 (2-99); 7244 (2-129); 7374 (6-3a); 7375 (7-
56b);7470 (7-54); 8466 (3-4); 8739 (7-54); 9649 (7-
20); 9671 (6-51b)
MartinezS., E., 1539 (1-1); 5798 (7-56b)
Martins,H. F., 336 (2-11); 354, 414 (6-25)
Martins,O., MG8155 (2-99)
Martins,T., 18 (2-41)
Martius,K. P. F. von, s.n. (3-4); 1840 (3-16)
Mathias,M. E. et al., 6108 (7-75)
Mattos Filho, A., 345 (7-54); 348 (7-20)
MattosSilva, L. A. et al., 386 (3-11); 390 (7-20); 1072
(2-?); 1073 (6-?); 1136 (2-84); 1176 (1-1); 1196 (2-
71); 1211(2-80a);1401,1402(2-84); 1440(2-130.1);
1586 (7-20); 1700 (6-51b); 1839 (2-84); 1870 (2-
71a); 1920 (2-88); 1984 (2-114)
Maury,C., 277 (3-16)
Mautone,L. et al., 118 (6-25)
Mazzeo, P. M., 199 (2-1)
McDaniel,S., 2660, 2496 (6-3b); 16280 (7-40); 17187
(2-8); 20530 (6-44); 21133 (2-43c); 22346 (6-44)
McPherson,G., 6979, 8017 (7-36a);8213 (7-56a);8231
(7-36a);8457 (7-56a);8479 (2-15);8577 (7-38);8771
(7-36a);9015 (7-38); 9498 (2-31.1); 9612 (7-80)
Medeiroset al., 58 (6-18b)
Medina,E., 291 (7-40); 322 (2-27a); 533 (2-119); 542
(2-103)
Meehan,s.n. (1-1)
Meijer,W. et al., VEN92 (6-48)
Meijeraan,J. W., 19 (7-7)
Mejia, M. et al., 6254, 6581, 10197 (1-1); 11174 (7-
36a); 23689 (1-1)
Melin, D., 41 (6-3a); 173 (2-27)
Mello, F. et al., 2 (6-11); 16 (2-69a);24 (7-50b);25 (3-
4); 43, 45 (2-64); 48 (2-69a); 55 (7-41); 71 (7-24);
114(7-56b);s.n. INPA51803 (6-3b);INPA55359 (2-
69a);INPA53361, INPA57834 (6-50); INPA57842
(2-39); INPA57898 (6-34); INPA57962 (6-35);
INPA57963(7-24);INPA57971(2-129);INPA60170
(2-69.1)
Mello Filho, L. E., 3061 (6-32); 3618 (7-54)
Mello Silva, R. et al., CFCR7548 (7-18); CFCR8313
(7-54); CFCR8397(7-70)
Mendonsa,S., 2 (7-73)
Meneces,E. et al., 340 (3-5); 669 (7-50b); 706 (2-99);
2061 (2 sp.)
Meneces,S., 360 (2-21); 669 (7-50a)
Mennega,A. M. W. et al., 882 (2-135)
Mercado,R., 9 (6-47)
Meredith,H. B., s.n. (2-1); s.n. (1-1)
Meyer, G., 46 (6-23); 78 (2-43b); 105 (7-66); 202 (7-
56a); 294 (2-43b)
Miers, H., RB179792 (7-85)
Mileski, E., 66 (7-66); 155 (6-3c); 168 (7-54); 171 (2-
27a); 187 (6-3c); 350 (6-3a)
Miralha,J. M. S., 49 (7-2); 63 (7-8); 67 (2-135.1); 74,
84, 87, 92 (2-43a)
FloraNeotropica
Miranda,C. A., 168 (2-80); 199 (4-2)
Miranda,F. E. et al., 299 (2-134); 309 (7-56a);378 (2-
136); 407 (7-30); 491 (2-69a); 539 (7-56b); 595 (6-
lb); 614 (7-24); 700 (7-56b)
Mocquerys,1021 (1-1)
Molina, D., 139 (7-22)
Molina, R., B-3 (2-18); 31 (3-6)
Molino, I., G416 (1-1)
Monsalve B., M., 152 (2-124); 481 (2-148); 445, 467
(2-46); 523 (3-15); 651 (2-148); 758 (2 sp.); 805 (2-
148)
Montalvo, E A., 4450 (7-56b)
Monteiro,O. P. et al., 19 (2-124); 29 (2-33.1); 32, 33
(2-119.1); 45 (1-1); 127 (6-3b); 133 (2-39); 162 (2-
88); 187 (7-56a); 375 (2-26); 412 (7-56b); 454 (6-
36); 516 (6-35); 532 (7-2); 779 (2-43b);781 (7-56a);
803 (7-24);804 (2-75);877 (2-43b);895 (7-56a);922
(6-34); 925 (6-9); 931 (2-43b); 936 (2-62); 945 (2-
142); 1032 (7-50b); 1059, 1067 (2-43b); 1071 (3-4);
1245 (7-1); 1270 (6-44); 1291 (2-27); 1363 (2-62);
1419 (2-69b); 1458 (6-3b); s.n. INPA50017 (6-14);
s.n. INPA50852 (2-43b); INPA50875 (2-69a); s.n.
INPA53419 (7-56a);INPA53534 (7-1); s.n. INPA-
50017 (6-14); s.n. (6-34)
Montouchet,P., 2202 (7-56b)
Mora, L. E., 2515 (1-1); 4424 (6-36)
Moreno,P. P. etal., 7263, 12059b, 12072 (1-1); 12173,
14570, 14612 (7-56b); 15205 (1-1); 16078B (7-22);
22530, 22830, 22876, 22957 (2-45); 23014, 23072
(7-23); 23130 (7-36c); 23166 (2-88); 23233 (7-36c);
23258 (7-23);23449 (2-45);23567 (7-22);23748 (7-
36a);23994 (7-22);24061 (7-36a);24627, 24980 (7-
56b); 25541 (7-16); 25576 (7-56a); 25614 (7-16)
Moretti,C., 1264 (3-4); 4510 (6-7)
Mori, S. A. et al., 2075, 2370 (7-38); 2876 (7-56a);
3653 (7-38); 3741 (6-13.1); 4091 (7-56a); 4188 (2-
95);4665 (2-31.2);4914 (2-31.2);5052 (2-18.2);5148
(7-38);5171 (2-88);5306 (7-36a);5543 (7-56a);6532
(2-31.1);7116 (6-13.1);7778 (2-18.1);7990 (2-31.1);
8015 (2-66);8048 (2-135);8064 (6-15); 8081 (2-89);
8088 (2-90) 8107 (2-109); 8131 (2-90); 8140 (7-25);
8158 (3-1);8181 (2-39); 8217 (2-109);8218 (2-113);
8224 (7-56a);8231 (2-77); 8306, 8330 (2-135);8401
(2-108);8459 (7-67); 8532 (6-7); 8536 (7-56a);8573
(6-2);8590 (7-2); 8672 (7-56a);8678 (4-3); 8862 (2-
112);8880 (2-69a);9002 (7-24);9003 (2-129); 9008
(6-29);9009 (2-129);9010 (7-40);9015 (6-3b);9065
(6-44); 9115 (7-40); 9234 (6-44); 9336 (6-17); 9598
(6-25); 9757 (1-1); 10241 (2-146); 10243 (3-3.1);
10252 (2-146); 10470 (7-31); 10871, 10934 (6-51b);
10936 (7-37); 11033 (2-146); 11037 (6-51b); 12001
(3-11); 12246 (7-20); 12354 (7-70); 12749 (6-52.2);
13063, 13673 (2-130.1); 13774 (2-124); 13819 (2-
144);13884(2-124);13989(2-88);13995(2-6);14000
(2-124); 14017 (6-17); 14575 (1-1); 14718 (2-109);
14763 (2-69a);14764, 14772(2-27.1); 14780(2-72);
14782 (2-43a); 14783 (7-39); 14790 (2-27.1); 14791
(2-43a); 14814 (2-99); 14821 (6-7); 14828 (2-27.1);
14830 (2-99); 14835 (7-73); 14870 (7-56a); 14897
(7-74); 14963 (7-39); 14978 (2-69a); 15160 (3-2);
15161 (2-149); 15201 (2-69a); 15230 (7-39); 15232
(2-112); 15244 (2-86); 15254(2-112); 15285 (7-20);
SupplementalList of Exsiccatae
15299 (2-69a); 15403 (1-112); 15411 (2-38); 15428
(2-27.1); 15435 (2-27a); 15461 (2-108); 15476 (2-
27.1); 15501 (2-99); 15523 (2-27.1); 15650 (2-8);
15796 (7-42); 15802 (2-99); 15822 (2-112); 15840
(2-76); 15855 (7-50b); 15859 (6-19); 15879 (7-24);
15902 (2-124); 15944 (2-69a); 16006 (2-17); 15975,
16007 (6-50); 16038, 16089 (7-42); 16120 (2-88a);
16143 (7-42); 16172 (2-39); 16177, 16180 (7-42);
16181 (2-108); 16214 (7-42); 16228 (2-99); 16234
(2-88a); 16290 (6-50); 16313 (2-99); 16312 (2-124);
16316 (7-24); 16323 (6-50); 16349 (2-69a); 16398
(6-50); 16402 (7-24); 16414 (2-43a); 16424 (7-42);
16444 (7-24); 16468 (7-42); 16476 (2-99); 16516 (2-
27a);16528(2-63);16762(7-20);16792(3-16);16842
(7-20); 16843 (7-51); 17186 (6-7); 17193 (2-141);
17201 (7-2); 17216 (2-64); 17219 (6-15); 17224 (2-
69a); 17238 (7-2); 17244 (3-12); 17245 (7-50b);
17290, 17332, 17380, 17422 (2-69a); 17476 (2-39);
17500 (7-24); 17547 (2-109); 17553 (2-99); 17578
(2-69a); 17579 (7-20); 17606 (2-39); 17658 (7-20);
17707 (2-109); 17713 (7-20); 17731 (2-86); 17961
(7-39);17977(2-27c);17980(2-109);18012(2-27.1);
18049 (2-99); 18080 (1-13); 18116 (7-50a); 18186
(2-69a)
Morillo, G. et al., 3460 (7-55); 3650 (2-99); 5121 (2-
69a); 5135 (7-21); 5314 (7-5); 5436 (7-21); 6948 (6-
3b)
Mosier, C. A. et al. 49 (1-1)
Mota, C. D. et al., 86 (2-22); 89 (2-62); 138 (6-3b);
191 (2-37); 246 (7-5); 739 (2-124); s.n. INPA60383
(2-141);INPAs.n.,INPA60396(2-69a);INPA60444,
INPA60480 (7-56b); INPA60611 (7-56a); INPA-
60614, INPA60635 (6-29); INPA60656 (6-13);
INPA60676(7-64);INPA60721(2-69a);INPA60724
(6-14); INPA60732 (7-50a); INPA60869 (7-56a);
INPA60898(6-14);INPA60927(7-56a);INPA60933
(6-29); INPA61336 (7-30); INPA61341 (6-14);
INPA61558(7-25);INPA61609(2-62);INPA61613
(2-36); INPA61659 (8-1); INPA61672 (6-3a); s.n.
(6-29); s.n. (7-56a)
Museu Goeldi, Belem, 9628 (2-57)
Narvaez, E., 633 (1-1)
Nascimento,O. C. et al., 20 (6-44); 29 (6-28); 191 (2-
69b);250 (6-2);259 (2-69b);279 (7-21); 340 (2-79);
352 (7-24); 363 (2-79);453 (7-56a);560 (6-3b); 571
(7-56b); 604 (2-69b); 611 (2-88a); 811 (6-lb); 867
(7-20); 875 (2-43a)
Nee, M. et al., 7675 (5-1); 11288 (7-80); 17773 (2-81);
24631 (2-15);27960 (2-45); 28191 (6-47);28329 (2-
15);28850 (1-1); 30848, 30999 (2-26); 31863 (6-3b)
Nehlin, S. 0., s.n. (7-33)
Neill, D., 1794 (7-36a); 1896 (7-2); 2829, 3103, 3136
(2-45); 3755 (7-23); 3795 (7-22); 4044 (7-23); 4104
(7-22); 4124 (7-56b); 4174 (7-36a); 4175, 4318 (7-
22); 4338 (7-23); 4508 (7-56b);4566 (1-1)
Nelson, B. W. et al., 344 (6-27);630 (3-5);804 (2-27b);
807 (2-69a);862 (7-73);871 (6-la); 913 (2-140);931
(2-27a);933, 939 (2-140);956 (2-27a); 1058 (2-69e);
1075 (7-56b); 1185 (6-3a); 1401 (7-56b); 2617 (2-
1);P21071 (2-57)
Nelson, C. et al., 4253 (7-56b)
Nelson, E. B., 4481, 4533 (1-1)
117
Nepomuceno,V., HPB2489 (2-43a)
Nevers, G. de et al., 3816 (7-38); 4446 (2-39); 4618
(1-1); 4773 (7-56b); 5106 (7-80); 5440 (2-5); 5767,
5840 (2-95);6446, 6558 (7-56b);7112 (2-145);7124
(2-88a);7553, 7581, 7603 (5-1); 7726 (9-1)
Nevling, L. R. et al., 2604 (7-56b)
Noblick, L. R., 1622, 2164 (7-18); 2864 (7-20); 3088
(7-18)
Nunes, E., HPB5947 (4-2)
Obando,R., 17 (1-1)
Oldeman, R. A. A. et al., 11 (2-4); 198 (2-27a); 291
(7-2); 1650, 1672 (7-56a); 1839 (6-18); 1871 (2-97);
1954, 2166 (2-99); 2237 (2-69a); 2773 (7-2); 2868
(7-56a);3097 (2-134);3223 (7-2);B492 (7-73);B551
(2-52); B898 (7-2); B1298 (2-69a); B1712 (2-97);
B1734A(7-50b);B1771(7-56a);B1875(7-74);B2031
(2-69a);B2514 (6-18); B2524 (7-50b);B2676 (6-7);
B2668 (2-129); B2688 (2-26); B3389 (6-2); B3458
(7-56a); B3464 (2-134); B3507 (7-56b); B3538 (2-
98);B3993 (7-2);B3999 (7-73);B4065 (2-86);B4353
(2-8);T226 (2-97);T280 (2-69a);T351 (2-97);T370
(7-50b); T447 (7-79); T486 (2-27a); T593 (2-27b);
T597, T648 (2-27c);T726 (2-124);T741 (6-2);T761
(2-69a);T919 (7-2); T933 (6-7)
Oldenburger,F. H. F. et al., 99 (7-56);232 (7-20);479
(2-122);483 (2-108); 620 (7-56b);961 (2-39); 1179
(2-112); 1213, 1225 (2-108.1); 1244 (7-56a); 1256
(2-100); 1416 (7-50a)
Oliveira,A. R. de, INPA58658 (6-4); INPA58700 (2-
99); INPA59593 (6-9); INPA59622 (2-27);
INPA59668 (2-43b); INPA59761, INPA59774 (2-
62); INPA59825 (6-15); INPA59877 (2-124);
INPA59908(6-9);INPA59936(2-69.1);INPA59967
(2-27); INPA59901 (2-69b); INPA60162, INPA-
73491 (2-69.1)
Oliveira,E. de, 302 (6-lc); 455 (4-3);2130 (6-1b);2655
(6-la); 3586, 4054 (2-69a); 4343 (2-63); 4471 (2-
69a); 5111 (7-56b); 5666 (3-1); 5617 (6-la); 5639
(6-1c);5709 (2-124); 5721 (6-lb); 5840 (6-50); 5962
(2-52); 6217, 6230 (7-56b); 6283 (2-69a); 6379 (7-
56b); 6494 (2-52)
Oliveira,H. B., 9(2-82)
Oliveira, M., 3017 (2-69); 3792 (2-43); 3922 (2-99);
3973 (2-136); 4909 (7-20); 5325 (7-56b); 5378 (2-
52); 6210 (2-43)
Oliveira,M. C. C., 2 (6-25)
Oliveira,P. E., 3 (7-54)
Oliveira,P. I., 508 (4-2)
Oliveira,R. F. de, 485 (1-1)
Ongley,J. C., P21739 (2-60)
O'Neill, H., 8625, 8626, 8627 (7-56b)
Oren,D. C., 3 (7-30)
Orlandi,R. P., 66 (2-80); 599 (4-2)
OrtegaU., A., 33 (2 sp.);73, 74 (2-88); 76 (2-27a); 110
(2-144.1); 146 (6-36); 154 (2-27a)
Ortega,F., 565 (1-1); 768 (2-12); 1691, 2517 (7-36a)
Ortiz, F., 1383 (7-23); 1974 (7-22)
Ortiz, R. T., 592, 1024 (7-56b); 2638 (7-22)
Osmarino(P. Monteiro)et al., s.n. (6-4); s.n. (6-44)
Pab6n E., M., s.n. (7-30)
Pabst, G., 5219 (3-4); 9559 (2-44)
Palacios,W. et al., 827 (6-36)
118
Paray,L., 523, 1817 (7-56b); 1944 (7-36a); 2017 (6-
47)
Passos, B. C. dos, 1073 (7-20); 1082 (6-3b)
Paula,J. Elias de, 684 (7-20); 1180 (1-1); 1497 (2-11);
1725 (6-20)
Pedrosa,J. S. et al., 1107 (1-1)
Pena, B. S., 73 (7-56b);82 (3-4); 99 (2-52); 184 (7-24);
242 (7-50b); 262 (7-50a); 324 (2-69a); 338 (7-25);
409 (6-3a);464 (2-27a);484 (2-37); 486 (7-54); 542
(6-la); 688 (2-62); 2002 (7-20)
Pennington,T. D. et al., 10152 (2-45); 10687 (6-16);
21675 (7-8)
Pereira,B. A. S., 59, 64 (6-27); 318 (2-43a)
Perez,J. L. A. et al., 4837 (7-56a)
Perez,P., s.n. (2-3)
Peters, C., 39 (6-44); 188 (6-16); 60-84 (6-41.1); 84-
11 (6-44)
Phelps, K. D., 511 (7-43)
Philcox, D., 3573 (2-17); 7774 (1-1); 7966 (7-25)
Philipson,W. R. et al., 2005 (2-76)
Pic6n, G. et al., 914 (7-25); 1045 (7-49)
Pinheiro,R. S., 448 (2-11); 1016 (2-36a); 1177 (3-36a);
1492 (2-11); 1558 (2-84); 1948 (3-11); 2078 (2-138);
2120 (7-36b)
Pinto, G. C. P., 176/81 (4-2); 54/82 (7-54)
Pinto, P. et al., 828 (7-40); 1172 (2-69b); 1244 (7-56b);
1256 (2-69b); 1269, 1317, 1344 (7-56b); 1345 (2-
69b); 1472 (7-56b); 1574 (2-69b); 1674 (7-22); 1902
(1-1)
Piniate,P. et al., s.n. (6-36); 1017 (3-6)
Pio, L. C., 13 (6-27)
Pipoly, J. J., 4535, 4692, 4795 (7-22); 4992 (7-56b);
6724 (2-39); 6730 (6-15); 6743 (4-3); 6751 (2-122);
6758, 6778 (2-43a); 6780 (2-102); 6788 (6-29.1);
6798 (6-7); 6823 (2-57); 6828 (2-17); 6892 (2-122);
7372 (2-135); 7356 (6-15); 7360 (6-42a); 7374 (6-
15); 7426 (6-42a); 7440 (2-90); 7446 (7-56a); 7497
(2-93);7507 (2-109);7513, 7524, 7537 (2-87);7701,
7760 (2-135);7820 (7-21);7828, 7927 (2-135);7941
(7-21)
Pirani,J. R. et al., 1216 (7-20); 1217 (2-37); 1250 (2-
114); 1292 (2-37); 1341 (3-16); 1352 (2-27a);
CFCR461,CFCR864(7-20);CFCR877,CFSC7971
(7-54); CFCR8277(6-27); CFCR8282(7-20)
Pires,J. M. et al., 74 (3-4); 108 (7-1); 184 (2-69a); 185
(2-64); 197 (6-3a); 210 (6-lc); 290 (6-34); 1622 (6-
3c);2914 (2-144);3195, 3544 (7-40);4146, 4923 (6-
la); 5293 (2-98); 5409 (2-63); 5864 (2-36); 5978 (2-
98); 6326 (2-17); 7124 (3-3); 7184 (2-69a);7700 (7-
79); 7925 (6-9); 10302 (6-la); 10543 (2-99); 10695
(4-3); 10755 (2-69a); 10781 (4-3); 10844 (2-69a);
10938 (2-63); 11005 (2-69a); 11012 (2-63); 11014
(6-la); 11029 (2-69a); 11055, 11065, 11066, 11137
(2-63); 11265 (2-69a); 11296 (2-52); 11359 (2-8);
11484(4-3); 11786(2-8); 11903 (3-2); 11931 (6-la);
12189 (2-69a); 12196 (2-18); 12214, 12245 (7-36a);
12394 (7-56a); 12556 (2-81); 12573 (7-36a); 12581
(7-56a); 12676 (2-23); 13077 (6-36); 13120 (6-50);
13149 (3-1); 13291 (2-86); 13295 (6-4); 13342 (7-
18); 13649 (6-4); 13737 (2-8); 13739 (7-36a); 13868
(2-99); 13915 (4-1); 13921 (2-27a); 13952 (6-7);
13957 (2-140); 13970A (4-1); 13972 (2-69b); 13977
FloraNeotropica
(6-3);13991(2-140);14008(3-6); 14022(4-1); 14044
(2-27a); 14059 (6-7); 14064 (2-141); 14072 (2-57);
14088 (2-75); 14157 (2-88); 14172 (2-93); 14192 (7-
49); 14206, 14217 (2-88); 14239 (6-7); 14249 (3-4);
14272 (2-93); 14360, 14375 (7-4); 14440 (6-3b);
14508 (3-1); 14542 (2-69a); 14548 (2-88); 14597 (3-
1); 14658 (6-3); 14755 (2-27a); 14760 (4-1); 14761
(7-25); 14738 (2-69b); 14783 (2-69a); 14849 (7-4);
14910(2-114); 15027(2-126.1);15939(7-20); 16135
(7-18); 16215 (6-3c); 16414 (7-66); 16560 (3-4);
16641 (6-27); 16713 (2-27b); 16811 (2-18); 16832
(3-4); 16930 (2-18); 16939 (7-21); 16967 (7-54);
17066 (7-66); 17135 (2-30); 50346 (6-7); 50930 (6-
10)
Plowman, T. et al., 7003 (2-69b); 7016 (7-40); 7699
(3-7);7736 (2-27b);7739 (7-56a);7784 (7-56b);9591
(7-63); 9674 (7-56b); 9837 (7-63); 10001 (3-16);
11371 (7-54); 11452, 11466 (7-56a); 12224 (7-5);
12390 (2-69a);12397 (2-129); 12414(2-124); 12448
(7-73); 12457 (2-129); 12458 (6-3b); 12466 (3-4);
12489 (7-40); 12525 (2-99); 12533 (7-56); 12725 (7-
20)
Poole, J. M., 1679 (2-69b); 1797 (2-88); 1798 (6-lb);
1804 (2-88); 1855 (7-56a); 1995 (2-27a);2092 (3-6);
2094 (2-129); 2118 (2-124)
Popenoe,J., 103 (6-47); 161, 242 (1-1); 718 (2-1)
Porter,D. M., 4078 (7-36a)
Poterima,L. B., 6385 (3-4)
Prance,G. T. et al., 3653 (7-67); 8821 (3-4); 10492 (2-
77); 10614(7-73); 10675, 10694(3-4); 10883 (2-18);
11144, 11323 (7-56a); 11460 (2-18); 11471 (4-1);
11480 (2-27a); 11492 (7-56a); 11506 (6-3); 11513
(2-27a); 11585 (6-3a); 11631 (6-29); 11793 (3-2);
11842 (7-5); 11883 (2-43); 11939 (2-94); 11943 (6-
28); 12058 (6-35); 12079 (6-36); 12151 (7-56b);
12227 (2-99); 12228, 12344 (7-73); 12345 (7-56b);
12560 (6-36); 13336 (7-56a); 13348 (2-88); 13349
(6-44); 13353 (2-36); 13404 (3-4); 13419 (7-56a);
13462 (7-56b); 13507 (2-27a); 13638 (2-77); 13693
(3-4); 13732 (7-56a); 13849 (7-6); 13868 (7-40);
13882 (7-6); 13883 (7-56a); 13925 (6-29); 13963,
13975 (2-69b); 13990 (7-56a); 14015 (8-4); 14024
(2-43b); 14070 (2-81); 14077 (7-6); 14079 (7-40);
14092 (2-17); 14095 (6-34); 14155 (7-40); 14158 (2-
94); 14159(2-62);14169 (2-17); 14172(2-99); 14178
(2-43b); 14194 (2-88); 14252 (2-17); 14276 (2-43b);
14320 (2-99); 14324 (7-56b); 14374 (6-4); 14388 (2-
56b); 14644 (2-69a); 14719, 14720 (1-1); 14735 (2-
69.1); 14755 (7-36a); 14787 (6-3a); 14839 (6-29);
14924 (6-3b); 14931 (2-27a); 14942 (6-35.1); 14981
(7-2); 14987 (2-27a); 14995 (2-36); 15001 (2-88);
15012 (6-15); 15013 (2-69a); 15018 (2-88); 15026
(2-98); 15030 (2-69a); 15043 (2-99); 15053 (6-3a);
15150 (2-69a); 15116 (6-3b); 15134 (7-24); 15166
(7-56a); 15265 (2-77); 15271 (7-56b); 15274 (6-2);
15336 (2-69a); 15388 (7-62); 15397 (2-77); 15426
(2-24); 15433 (6-3b); 15463 (2-88); 15503 (2-93);
15505 (4-1); 15604 (7-72); 15745 (2-52); 15820 (7-
62); 15935 (6-34); 16014 (6-2); 16035 (6-5); 16040
(2-99); 16048 (7-62); 16055 (2-22); 16141 (7-41);
16143 (6-41); 16167 (2-88); 16196 (4-1) 16215 (2-
27a); 16229 (2-69b); 16315 (6-23); 16368 (6-37);
 Listof Exsiccatae
Supplemental
16401 (7-56); 16408 (2-39); 16452 (7-62); 16518 (6-
44); 16521(6-41); 16524(2-43b);16526(7-2); 16529
(6-29); 16540, 16542 (1-1); 16545 (2-1); 16548 (7-
21); 16562(6-42a);16563(6-15);16564(2-78);16772
(7-36a); 17519 (6-4); 17520 (2-79); 17541 (2-69a);
17575 (7-2); 17750 (7-1); 17751, 17800 (2-69a);
17816 (2-119); 17820 (2-79) 17828 (7-73); 17897
(2-124); 17948 (2-82); 17950(2-60); 17985 (2-27a);
18000, 18001 (2-69a); 18051 (6-29); 18237 (7-56a);
18301 (2-69a); 18304 (6-3b); 18310 (2-17); 18727
(7-1); 18739(6-4); 18747 (6-9); 18759 (6-15); 18840
(6-27); 18920 (3-16); 19040 (2-114); 19061 (7-54);
19078 (7-20); 19161 (2-144); 19329 (2-37); 19361
(7-66); 20012 (2-68); 20199 (7-56a); 20555 (7-6);
20591 (2-69b);20592 (2-28); 20611 (2-69a);20626
(6-3b); 20633 (2-88); 20729 (6-11); 20768 (6-15);
21006 (6-16); 21007 (2-85); 21032 (7-8); 21629 (7-
6); 21636 (6-35.1);21639 (7-8);21652 (6-29);21653
(7-56a); 21661 (7-2); 21670 (2-114); 22611 (2-64);
22622 (6-41); 22658 (7-56a); 22682 (7-73); 22693
(2-39); 22721 (2-22); 22775 (2-64); 22807 (2-121);
22822 (2-17);22832 (2-99);22843 (2-21);22850 (2-
124); 22858 (2-43b); 22939 (2-28); 22972 (2-33);
22977 (7-56b);22986 (2-43b);23028 (2-64); 29997
(2-17);23131 (4-1);23253 (2-26);23366 (7-2);23379
(6-11); 23395 (6-9); 23481 (6-29); 23505 (2-26);
23508 (7-9); 23548 (7-56b);23595 (6-9); 23601 (7-
1);23762 (6-35);23789 (7-62);23816 (6-35); 23826
(7-41); 23860 (7-56a); 23938 (6-7); 23971 (6-29);
23974 (7-40.1);24123 (7-56a);24163 (7-62); 24166
(7-40); 24177 (7-62); 24215 (2-27a); 24216 (4-1);
24238 (7-56b);24251 (2-124); 24257 (7-67); 24268
(2-36);24269 (6-41);24287 (6-4);24297 (4-3);24539
(2-129); 24574 (7-2); 24731 (7-63); 24759 (2-28);
24800 (2-141); 24875 (2-75); 24895 (7-20); 25001
(7-50a); 25065 (2-57); 25087 (2-81); 25140 (2-75);
25190 (7-20);25264 (2-75);25190 (7-20);25264 (2-
17);25531 (2-69a);25574 (4-3);25575 (2-27b);25642
(2-39); 25652 (2-14.1); 25677 (2-99); 25789 (7-2);
25843 (2-69a); 25885 (4-3); 26127 (2-43b); 26130,
26190 (2-28); 26219 (2-43b); 26302 (6-20); 26318
(6-3b);26319, 26323 (4-1); 26324 (6-3a);26327 (4-
1);26343 (6-21);26352,26359 (2-27a);26372,26509
(7-36a); 26564 (2-8); 26630 (6-16); 27992 (7-56b);
28053 (2-51); 28076 (1-1); 28134 (2-11); 28154 (1-
1); 28078 (7-2); 28100 (7-74); 28154 (1-1); 28510
(7-56b);28171 (2-145); 28373 (7-21); 28376 (7-25);
28399 (7-21); 28448 (2-56); 28453 (2-69b); 28472
(7-56b);28693 (2-23);28762 (2-114);28868 (6-29);
28869 (7-49);28875 (2-93);28903 (2-57);28908 (4-
3);28919 (2-43a);28941,29000 (7-30);29037,29091
(2-135);29098 (2-69a);29137 (2-135);29163 (7-49);
29181 (2-69a);29261 (6-29.1);29331 (6-57);29357,
29389 (7-83); 29420 (6-29.1); 29443 (6-la); 29469
(7-62); 29513 (6-9); 29663 (2-69a); 29675 (6-29a);
29715 (6-9); 29721 (7-49); 29738 (2-37); 29741 (2-
36); 29749 (6-29.1); 29766 (7-49); 29768 (6-29a);
29809, 29815 (2-119a); 29834 (2-131); 29843 (6-
29.1);29856 (4-1);29877 (2-39);29913 (2-18);29931
(2-140);29947 (7-56a);29941 (2-88);30011 (2-140);
30053 (7-25); 30110 (7-36a)
Prevost, M. F. et al., 311 (2-56a); 343 (7-25); 355 (2-
119
109); 358 (2-86); 378 (7-56a); 393 (2-114); 437 (2-
36); 448 (2-56a);932 (3 sp.); 1059 (2-134); 1168 (7-
52); 1201 (2-114); 1495 (6-7); 1518 (7-52); 1574 (2-
69a)
Procter,J., 4737 (7-56a);4738 (2-135)
Pruski,J., 1511 (1-1)
Pulle, A. A., 182 (3-1)
Pulle-Lutz,1130 (7-36b)
Pyron,J. H. et al., 3145 (2-1)
Quintero,A., 2204 (7-36a)
Rabelo,B., 152 (2-43a); 164 (3-4); 230 (2-36); 605 (7-
18); 1528 (7-25); 1827 (3-4); 1839 (2-39); 2181 (7-
56a); 2193 (2-39); 2249 (3-4); 2275 (2-43a); 2285
(7-52);2380 (7-50b);2381 (7-56d);2475 (2-18);2477
(7-24);2478,2480,2482 (2-43a);2486 (7-50b);2491
(2-114); 2660 (2-39); 2695 (3-4); 2712 (2-8); 2774
(3-4); 2792 (6 sp.); 2885 (2-114); 2922 (7-18); 2925
(2-69a);2928 (7-56a);2931 (2-69);2938 (2-69a)2976
(2-43a);2990 (2-69a);3018 (3-2); 3020 (6-50); 3029
(2-69a);3038 (3-2); 3058 (2-39);3072 (2-69a);3128
(2-109);3135 (6-15);3147 (2-69a);3197 (6-la); 3223
(3-2)
Ramcharan,E. K., 438 (1-1)
Ramia, M., 7176 (2-27b)
RamirezC., R., 4 (6-34); 18 (7-40); 87 (6-38); 104 (2-
5); 1083 (7-40); 1094 (2-140)
Ramos, J. F. et al., 90 (6-41); 138 (7-56a); 157 (2-26);
378 (2-69a);382 (6-11);429, 433 (7-5);637 (7-56a);
646 (7-66); 651 (6-22); 755 (2-119.1); 792 (7-65);
851 (2-17); 865 (7-56a);877 (7-30); 914 (2-63); 940
(3-3); 942 (2-136); 945 (2-114); 1048 (2-63); 1087
(2-27a); 1090 (2-17); 1154 (7-56b); 1165, 1196 (2-
69a); 1507 (7-63); 1589 (2-81); 1668 (7-63);P21800
(6-11);P23251(8-4);INPA62116(7-50a);INPA2157
(6-9); INPA62177 (7-56b); INPA62187 (7-56a);
INPA62246 (7-56b); INPA62279, INPA62293 (7-
24); INPA62319 (6-34)
Ramos, R., 174 (7-22)
Ramsey, G. W. et al., 137 (2-1)
Rankin,J., 27 (2-64); 29 (2-69a); 137 (2-108)
Ratter,J. A., 2074, 2163 (2-41); 3232 (2-30); 3557 (7-
66); 3619 (6-27); 3739 (3-16); 3845 (2-4); 3941 (2-
30); 3956 (2-28); 4439 (2-27a); 4441 (2-30); 5076
(2-18)
Rau, E. A., s.n. (2-1)
Ravelo, O., 33 (2-113)
Reederet al., LBB12309(2-69a)
Reis, G., 149 (2-43a)
Reis, L. Q., s.n. INPA57832, s.n. INPA57833 (6-50)
Reitz, R. et al., 18086 (3-4)
RenteriaA., H., 1460 (1-1)
Renteria,E. et al., 2186 (2-56a)
Revilla, J., 171 (7-40); 359 (2-27a); 383 (2-27); 402
(6-3b);406 (2-37);479, 580, 676 (6-44);690 (2-69b);
698 (6-3b); 816 (6-14); 1120 (7-56b); 1167 (6-6a);
1483 (7-5); 1666 (2-17); 1814 (7-40); 1843 (7-56b);
1903 (2-140);2138 (6-la); 2218, 2247 (2-140);2345
(2-94); 2421 (6-44); 2465, 4071, 4075 (6-3b); 4156
(4-1);4537 (7-56a);7062 (2-22); 7086 (2-128);7106
(2-57);8372 (7-24);8380 (2-69a);8414 (2-17); 8416
(2-67);8421 (2-136);8433 (2-63);8434 (2-17);8529,
8531 (7-56b);8666 (6-21)
120 Flora Neotropica

Reyna R., N., 31, 132-I (2-128) (3-6);436 (6-21);513(2-94); 540 (6-41);672 (2-141);
ReynelR., C., 366 (2-21);422 (7-73);444 (7-36a);498 732 (2-69a); 847 (7-56a); 882 (7-50a); 888 (2-37);
(3-9); 558 (2-27a); 566 (7-41); 643 (2-43b); 653 (7- 914 (7-20); 916 (2-43b); 961, 988 (7-18); 1006 (2-
41); 695 (2-45); 850a (7-56a); 967 (7-36a) 63); 1052 (7-11); 1061 (2-117); 1133 (7-50b); 1134
Riba, R. et al., 292 (2-15) (2-109); 1163 (7-20); 1165 (2-67); 1353 (7-69); 1423
Ribanov, J. et al., 151 (2-43b); 169, 205 (2-108); 218, (6-43);1452(7-56b);1467(2-27b);1498(7-25);1564
245 (2-88); 268 (7-56a); 277 (7-50b); 281 (2-108); (2-119.1);1622(2-28); 1689 (2-36); 1697 (4-3); 1707
295, 303 (3-4); 319 (2-81); 325 (2-124); 345 (2-81); (7-8);1713(6-29.1);1795(2-112);1799(2-109);1926
347 (7-50b); 353 (7-1); 390 (2-36); 391 (2-69a) (3-9); 1931 (3-4); 1967 (2-75); 1988 (2-28); 2025 (7-
Ribeiro, B. G. S., 142 (6-13); 151 (6-la); 183 (2-52); 54); 2046 (2-43b);2098 (7-73); 2210 (3-4); 2308 (2-
273 (7-56b); 277, 299 (6-21); 307 (7-56a); 447 (6- 114); 2310 (2-69a); 2331 (7-56a); 2337 (2-126.1);
3a); 484 (6-37); 752 (2-119); 844, 1013 (7-8); 1090 2405 (6-50); 2568 (2-56b); 2708 (2-86); 2724 (7-
(7-50a); 1105, 1107 (2-52); 1307 (3-1); 1511 (7-18); 56a); 2727 (2-79a); 2753 (2-99); 2848 (2-86); 2891
1548 (7-24); 1584 (7-56a); 1608 (7-24); s.n. (7-30) (2-37); 2923 (7-24); 2924 (2-99); 2930 (6-12); 2942
Richardson,W. D., 744 (7-25) (2-27a); 2954 (2-134); 3097 (2-27b); 3120 (2-122);
Rico Gray, V., 124, 444 (1-1) 3124 (2-18); 3148 (3-3); 3155 (6-50); 3157 (2-81);
Riedel, L., s.n. (3-16); 405 (2-25); 1111 (2-30); 1521 3310, 3586 (7-36a); 3651 (7-53); 3655 (7-14); 3607
(2-27b); 1869 (7-85); 2319 (3-16); 2841 (6-27) (7-28)
Riedel, W. D., 1371 (6-16); 1667 (3-4) Rosario, C. da S., 9 (7-73)
Rimachi Y., M., 1803 (2-52); 1807 (7-40); 1854 (2- RosarioT., A. J., 82 (2-45)
69b); 1861 (2-60); 1881 (2-140); 1884 (7-40); 2264 Rosbach,G. B., 3719 (7-61)
(6-35); 2302 (7-40); 2324 (7-53); 2326 (2-64); 2424 Ruiz, A. G. et al., 224 (2-128)
(6-3b);2740 (7-53); 2789 (6-35);2797 (2-43c);2809 Ruiz C., J., 30 (2-43c); 500 (2-22)
(6-44); 2838, 2866 (2-23); 3136 (2-69b); 3165 (2- Ruiz, D., 182 (7-22)
52); 3250 (6-36); 3260 (6-26); 3274 (2-7); 3277 (2- Ruiz, T. et al., 3949 (2-69b)
69b); 3357 (6-44); 3374 (7-56b); 3391 (7-5); 3465 Rutkis, E., 55 (2-26); 285 (6-3b); 712 (2-27b)
(6-lb); 3472 (2-26); 3493 (3-9); 3643 (7-56a); 3681 Rylands,A., 42/80 (2-124); 53 (2-99); 56/80 (2-124)
(2-69a); 3685 (3-4); 4208 (7-24); 4257 (2 sp.); 4267 Rzedowski,J., 22516 (2-15)
(6-3b);4378 (3-14); 4622 (2-26); 4681 (7-40) Sabatier,D., 50 (2-63); 75 (2-69a); 71 (2-65); 110 (2-
Rizzini, G. M., 158 (7-36a) 99); 119 (6-7); 130 (2-35); 205 (6-la)
Rizzo, J. A. et al., 2990 (2-114); 4312 (7-54) Safford,W. E., s.n. (2-1)
Roa, A., 365 (6-41.1); 435 (2-82) SagasteguiA., A., 550, 6866 (7-56b)
Robbins, S. B., 5793 (1-1) Salazar,A., 2, 662 (2-64)
Robert,A., 567b (6-27) Saldana,655 (3-4)
Roberts,L., LBB14763 (2-117); LBB16303 (7-11.2) Sanchez,E., 8 (1-1)
Robertson,K. R. et al., 243 (2-77) SanchezV., P. et al., 424 (7-36a)
Robleto, W., 8 (1-1); 194 (2-15); 372, 408 (2-45); 641 Sandino,J. C. et al., 797 (1-1); 1822 (2-45); 2239 (1-
(7-22); 739 (6-47) 1);2678 (7-22); 3012 (1-1); 3630 (2-15); 3949, 4046
Rodrigues,I. A., 234 (7-54) (1-1); 4312 (6-47); 4395, 4407 (2-15); 4447 (2-45);
Rodrigues,R. S., MG8813 (2-86) 4746 (7-22); 4772 (7-36c); 4963 (2-45)
Rodrigues,W. A. et al., 887f(2-124); 1408 (8-4); 5402 Santino, 284 (7-54)
(6-la); 7263 (2-136); 8257 (7-55.1); 8353 (2-27a); Santos,A., 84 (7-36a)
8770 (2-22); 8815 (2-27a); 8832 (4-1); 9064 (6-28); Santos,F. S., 193 (6-49); 409 (2-lOOa)
9097 (2-53); 9278 (2-88); 9284 (2-119.1); 9317 (7- Santos,J. L. dos et al., 681 (2-119.1); 717 (2-131);758
8); 9405 (7-56a); 9408 (7-54); 9411 (7-50a); 9621 (2-165)
(2-69a); 10064 (2-43); 10129 (2-131); 10224 (2-57); Santos,J. U., 255 (7-73)
10230(7-56b);10259 (2-121.1);10497 (2-57); 10501 Santos,M. R., 11 (2-69b);57 (2-22); 120 (2-69b); 162
(2-135.1); 10505 (6-15); 10509, 10528 (6-29.1); (2-129); 166 (3-4); 281 (2-69a); 286 (7-24); 309 (7-
10542 (2-37); 10556 (2-69a); 10559 (6-7); 10609, 56a); 377 (2-39); 396 (2-27a); 444 (2-136); 466 (7-
10616 (2-57); 10630 (6-15); 10635, 10642 (2-57) 50b); 472 (2-69a);485 (2-27a); 507 (2-99); 532 (7-
RodriguezA., A. A. et al., 57 (7-56b) 63); 569 (7-56a);643 (2-69a); 652 (3-4); 686 (2-39)
Rodriguez,B., 1419 (7-15) Santos,R. R., 49 (4-3)
Rogers,G., 27 (7-5) Santos, T. S. dos et al., 164 (3-18); 300 (7-22.1); 309
Rojac, C., 4117 (3-16) (6-26); 311 (3-18); 320 (2-146); 322 (6-53); 326 (2-
Rojas, M., 50 (2-15) 114); 457 (3-3.1); 526 (7-36a); 570 (7-64); 945 (2-
RomeroCastafieda,R., 1212(7-21);5270 (2-148);5570 43a); 1279 (6-51b); 1284 (7-37); 1402 (7-20); 1441
(3-17); 8445 (7-56a) (7-36a); 1444 (2-146); 1512 (7-70.1); 1677, 1695 (7-
Rombouts, H. E., 321 (7-20) 13); 1837 (2-43a); 1995 (7-36a); 2287 (7-20); 2305
Rooden, J. van et al., 358 (2-124); 536 (2-32); 553 (2- (3-3.1); 2349, 2468 (2-43a); 2693 (7-36a); 2898 (7-
124); 586 (3-15) 13); 2935 (6-52.2); 2984 (7-20)
Rosa, M., 74 (7-36b) Sarmiento,A. C., 604 (4-2); 614/80, 651 (7-18)
Rosa, N. A. et al., 33 (2-134); 76 (7-56a); 170 (2-28); Sastre,C. et al., 514 (6-44); 641 (3-14); 671 (7-56a);
242 (2-37);373 (2-140);378 (2-24);381 (2-27a);384 839, 854 (7-40); 872 (7-56b); 1027 (7-40); 1311 (2-
 List of Exsiccatae
Supplemental
69a); 1363 (7-56a); 1397, 1415 (7-73); 1617, 1816
(7-56a);2357 (7-62);2427 (2-69b);2436 (7-62);2437
(2-68); 3127 (7-5); 3268 (7-56b); 3273 (7-35); 3350
(2-69b);3358 (7-55);3370 (7-56b);3443 (3-1);3486
(2-68);4107 (7-73);4156(7-18); 4348 (2-134);4426,
4536 (2-112); 5060 (2-104); 5070 (7-5); 5106 (7-2);
5178 (2-104); 5225 (7-40); 5587, 5674 (7-73); 5688
(7-56a);5990 (7-18); 5991 (7-56a); 6110 (3-1)
Saunders,J., 153, 223 (1-1); J 233 (7-22); 417 (1-1);
530, 574 (7-56b); 743 (1-1); 1197 (7-23); 1304 (7-
56b)
Sauvain,M., 156 (2-88)
Schatz, G. E. et al., 779 (7-56b)
Scheiner,P., 50 (2-114); 58 (7-54)
Schmidt,E., 26 (3-5)
Schnell,R., 11129 (7-25); 11134 (1-1); 11425 (7-56);
11728 bis (7-56a); 11751 (7-2); 11849 (7-56); 11899
12040 12069 12089
(7-73); (3-1); (6-lb); (6-1c);
12114(3-1)
Schomburgk,R. H., 318 (2-134)
Schubert,B. G., 42 (1-1)
R. E. et al., 6520 (6-3b); 12979 (3-6); 13416
Schultes,
(2-69b);14913(2-39); 15456, 15660(2-27.1); 16856
(2-69b); 19161 (7-21); 19398 (7-56b); 19421 (7-5);
19602 (2-69b);26038 (7-56b);26106 (6-9); 26139a
(6-13); 26184a (7-28)
SchunkeV., J., 10 (7-75); 16 (2-68); 1737 (2-45); 2008
(7-36a); 2050 (2-43b); 2063 (7-56a); 2137 (7-75);
2187 (2-69a);2194 (7-56a); 2587 (2-43b); 3344 (6-
36); 3525 (7-56a); 3794 (7-28); 4103 (2-141); 4128
(7-56a); 4389, 4610 (7-36a); 4767 (6-36); 4900 (2-
129);5622, 6210 (7-21); 6247 (6-16); 6314 (2-129);
6447 (7-73); 6502 (2-7); 6518 (7-28); 6637 (7-56a); Silva, M. N., 271 (7-20); 390 (3-4); 403 (7-56b)
6671, 7222 (7-21); 8439 (2-144.1); 11926 (6-34)
Schwacke,C. A. W., 256, 4256 (3-4)
SEF(StudiesofEcuadoreanForests),9269 (2-60);10321
(6-36)
Seymour,F. C., 3787 (7-56b);4593, 5957 (1-1)
Shepherd,G. J., 7443 (7-19)
Shepherd,J. D., 197 (7-34)
Shepherd,W. O., 62, 145 (2-1)
Sidney, (Fonseca)et al., 199 (7-54); 1302, 424 (2-30)
Silva, A. F., s.n. INPA68839 (6-29)
Silva, A. S. L. da et al., 8 (2-99); 65 (2-141); 132 (2-
129); 134 (2-81); 144 (2-56b); 220 (6-4); 426 (3-2);
452 (6-57); 511 (6-14); 534 (6-13); 540 (7-5); 569
(6-9); 1980 (2-134)
Silva, F. C. F. da, 35 (7-54); 82 (2-144); 237 (2-44)
Silva, I. A., 8 (2-69a); 30 (7-37); 94 (2-114); 131 (7-
85); 136 (7-64); 267, 268 (3-4); 283 (6-17)
Silva, J. A., 105 (7-56b); 178 (2-124); 193 (2-99); 202
(2-27b); 242 (2-81); 278 (3-4); 290 (2-17); 301 (6-
3b); 338 (2-27a); 356 (2-129)
Silva, J. C. da, 40 (7-19); 100 (7-52.1)
Silva, M. F. F. etal., 1102 (2-43a);1382 (2-134); 1404,
1446 (7-56a)
Silva, Manoel, s.n. INPA27688 (2-112)
Silva, MarleneF. da et al., 45 (2-57); 46 (7-56a); 50 Simpson, D. R. et al., 51 (2-43b); 76 (7-40); 732 (7-
(2-62); 80 (2-69.1); 127 (4-1); 135 (6-3b); 146 (4-1);
177 (7-73); 193 (7-2); 223 (6-15); 224 (2-124); 228, Skog, L. et al., 5643 (7-56b)
243 (2-69a); 248 (6-29); 262 (2-83); 267 (6-35.2); Slane,V., 12 (1-1)
337 (2-43b); 348 (2-17); 360 (2-142); 391 (2-69b); Smith, C. E., Jr., 6034 (6-3b); 6054 (2-12)
121
465 (6-15); 467 (6-35.2); 486 (2-64); 490 (6-69.1);
516 (2-69a);527 (7-1); 535 (7-56b);562 (2-75); 569
(7-6); 705 (2-99); 732 (7-1); 782 (7-20); 788 (7-1);
791 (7-6);822 (6-35.2);824 (2-83);828 (2-142);873
(2-73.1);874 (2-21); 901 (2-83); 914, 922 (3-2); 930
(2-99);942 (3-2); 967 (6-34) 981 (2-88); 993 (6-29);
1027 (6-35.2); 1151 (7-56a); 1186 (2-28); 1207 (2-
69b); 1228 (3-6); 1339 (2-27a); 1365 (2-140); 1466
(3-6); 1474 (7-62); 1510 (7-21); 1532 (2-140); 1538
(2-27a);1618(7-55);1620 (7-56a);1630(2-28); 1671
(7-55); 1701(2-69a);1815, 1866(2-69b);1918, 1932
(3-4); 2043 (2-27b);2230 (6-15)
Silva, Milton G. da et al., 716 (2-86); 892 (6-3a);936,
945 (2-27a); 1147 (2-60); 1181 (6-37); 1220 (2-81);
1304 (6-3a); 1408 (6-37); 1914 (2-69a);2190 (6-21);
2345, 2471,2686 (7-56a);2716 (7-24);2718 (7-56a);
2759 (7-18); 2818 (1-1); 2837 (7-56b); 3122 (3-4);
3128 (7-15); 3217 (2-17); 3230 (2-37); 3269 (2-30);
3327 (2-37); 3408 (2-63); 3496 (7-56b); 3508 (2-
121.1); 3527 (2-37); 3554 (2-69a); 3570, 3599 (7-
53); 3620 (6-3a); 3630 (2-129); 3705 (7-56a); 3754
3792 (2-27a);3793 (6-21);3805 (2-22); 3825
(7-24);
3873 (6-21); 3962 (2-52); 4281 (2-27); 4296
(6-3a);
(2-64.1); 4586 (2-41); 4662 (2-57); 4684 (2-64.1);
4744 (7-56a); 4746 (3-4); 4753 (7-20); 4826, 4845
(7-75); 4892 (7-20); 4906 (2-41); 4945 (7-54); 5008
(2-41); 5020 (6-27); 5026 (2-41); 5207 (7-30); 5237
(6-1c); 5279 (7-30); 5324 (2-81); 5348 (6-lc); 5402
(2-43b); 5442 (2-81); 5455 (7-56a); 5489 (2-69a);
5498 (6-50); 5500 (2-57); 5513 (7-56a);5522 (2-81);
5524 (7-69); 5528 (7-42); 5534, 5535 (6-50); 5537
6574 (2-92);6589 (2-53);7121 (6-29.1);7122
(6-la);
(2-119.1)
Silva,N. T. da, 593 (6-lc); 838 (2-8); 1045 (6-50); 1756
(3-4); 1796 (4-3); 1825 (2-114); 1831 (7-24); 1841
(6-la); 1857 (3-4); 1881 (6-50); 1901 (7-36a); 1969
(7-50b);2093 (2-39);2096 (2-53);2170 (7-27);2203
(2-53); 2215 (7-42); 2294 (6-la); 2329 (2-83); 2380
(3-4); 2385 (7-15); 2386 (7-42); 2408 (6-la); 2420
(7-56a);2512 (7-42);2517 (6-1a);2547 (7-56a);2621
(6-7);2683 (6-la); 2768, 2782 (2-69a);2826 (2-136);
2833 (6-la); 2879, 2911, 2925 (2-124); 2950 (7-24);
3170 (6-lc); 3203 (3-4); 3223 (6-la); 3241 (3-4);
3292 (7-50b); 3296 (6-la); 3310 (3-2); 3311 (3-4);
3316 (6-la); 3342 (6-50); 3391 (7-36a);3424 (6-la);
3556 (2-17);3615 (2-141);3631 (2-17);3727 (2-75);
3742 (2-129); 3874 (2-140); 3894 (7-21); 3897 (7-
8); 3926 (2-43a);3950 (6-la); 3978 (2-99); 4028 (6-
41.1); 4174 (7-62);4456 (7-56b);4459 (2-17); 4490
(2-27a);4492 (2-129); 4521 (6-lb); 4578 (2-119.1);
4609 (2-43b); 4612 (2-36); 4753 (2-69a); 4755 (2-
142); 4764 (6-21); 4796, 4805 (6-3a); 4810 (2-30);
4817 (7-56b); 4821 (2-27a); 4827 (6-20); 4837 (7-
54); 5064 (2-37); 5086 (7-24); 57170 (3-16); 60697
(2-27.1)
Silva, S. B. da et al., 383 (7-18)
Silva Costa, J. da, 1221 (2-28)
53); 761 (7-40)
122
Smith, D. et al., 1173 (2-98); 8409 (7-56b)
Smith, E., 8 (2-21)
Smith, F. D., Jr., s.n. (2-12)
Smith, R. F., V1592 (2-76); 4301, 4302 (2-12)
Smith, S. F. et al., 121, 190, 234, 297, 340, 381 (7-
56a); 590 (7-53); 7856 (2-69b)
Snow, D. W., 8 (7-25); 21 (7-26); 37 (7-25)
Soares,S. et al., s.n. on 18 II 1985 (2-122)
Sobrinho,J. F. et al., 322 (7-18); 890 (3-4)
Soejarto,D. D. et al., 360 (1-1); 449 (2-12); 694 (2-
69a); 2396 (6-la); 4025 (7-34)
Solomon,J. C., 6111 (7-56a);6205 (7-66); 6509, 7605
(7-36a); 7659 (7-56b); 7788 (7-53); 7856 (2-69b);
7944 (7-66); 8502 (7-46); 8784 (7-36a);9393 (7-46)
Sonkin, L., 340 (3-4)
Soria S., M. A., 20 (2-22)
Soto Nuiiez, J. C. et al., 32 (2-45)
Souza, A. B. de, 78 (6-25); 83 (4-2)
Souza, D. S., 242 (1-1)
Souza,J. L., 69 (7-56a)
Souza, H. M. de, IAC21452 (7-54)
Spada,J., 007/77 (6-17); 148 (6-26); 151 (4-5); 193 (2-
43a); 210 (7-13); 329 (2-114); 31/78 (6-13.2); 67/78
(2-43a);79/78 (2-71)
Sperling,C. et al., 5792 (7-56a); 5794 (7-36a); 5940
(2-43a);5942 (3-4);5945, 5977 (2-99);6029 (7-36a);
6028 (7-53); 6048 (7-36a);6072 (7-73);6102 (7-15);
6124 (2-134);6176 (7-73);6178 (7-15);6251 (2-17);
6327 (7-65);6334 (2-43b);6370 (7-36a);6437,6443,
6640 (7-56a)
Spetzman,L. A., 539 (1-1)
Spichigeret al., 1004 (7-41); 1170 (7-40)
Spongberg,S. A. et al., 17197 (2-1)
Stahel,G., 322 (2-88); 353 (2-18)
Stannard,B. et al., CFCR5960(2-144)
Stein, B A. et al., 1329 (7-56b); 1369 (7-36a); 1471 (7-
55); 1486 (7-62)
Stergios,B. et al., 2683 (3-1); 2787 (7-56a); 3179 (2-
69b); 3464 (1-1); 4112 (2-88); 4208 (7-5); 4420 (7-
21); 4685, 4881 (7-36a);4991 (6-45); 5535 (2-27b);
5718 (7-22); 5793 (7-36a);6175 (6-45); 7944 (3 sp.);
8455, 8460 (7-40); 8470 (2-27); 8619, 8623 (7-56b)
Stevens, W. D. et al., 5453 (2-45); 7204, 7478 (7-22);
7555, 7634 (7-23); 7642 (7-56b); 7714 (1-1); 7802
(7-22); 7818 (1-1); 7853 (7-56b); 7866 (2-88); 7902
(1-1);8811 (2-22); 8152 (7-56b);8283 (7-36c);8469
(7-56b); 9669 (2-45); 10473 (1-1); 10475 (7-56b);
10478, 10686 (1-1); 12104, 12769 (7-56b); 17147
(2-45); 17560 (2-15); 17746 (1-1); 18644 (7-56b);
19456, 19563, 19584, 19606 (1-1); 19826 (2-88);
19828 (7-56b); 19993 (1-1);20058 (2-88);20075 (7-
56b); 20635 (2-88); 20636 (7-56b); 20875 (1-1);
21460 (7-22); 23357 (2-15)
Stevenson,N. S., YALE10697 (7-23)
Steward,W. C. et al., 36 (2-69a); 38 (2-36); 54 (6-19);
61 (7-73); 88 (2-119.1); 110(2-33); 130(2-134); 134
(6-3b); 174 (7-21); 289 (2-27); 341 (2-27a); 390 (2-
69b); 511 (7-56a); P17669 (2-69.2); P17680 (3-4);
P17697 (7-8); P20128 (6-29); P20241 (2-69a);
P20251 (2-119.1); P20304 (6-4); P20310 (2-27);
P20326 (2-119.1); P20328 (2-88); P20369 (2-36);
P20401 (2-119.1)
Steyermark,J. A. et al., 52829 (2-4.2); 75127, 75526
FloraNeotropica
(6-8.1);86613 (3-3); 87137 (7-74); 87610, 88108 (6-
6);93035 (6-8.1);101826(3-7);102626(3-1);102951
(2-27a);102953 (2-119); 102974 (2-27a);103013 (7-
21); 103225(6-3b); 103247 (2-131); 103260(2-119);
104154 (2-56); 104223 (7-26); 104352 (2-124);
104492 (7-26); 104547 (2-69a); 104777 (7-22);
105503 (2-56); 105936 (6-8.1); 106087 (2-69a);
106117 (6-8.1); 106359 (7-49); 106411 (2-26);
106412(2-28);106645(7-49);106847(7-22);107132
(7-50b); 107357 (2-60.1); 107456 (2-68); 108611 (7-
56b);108939,109173(2-135);109900(2-12);111391
(2-105); 111507 (2-69a);111512(2-145); 111541(2-
5.4); 111609(7-22); 112386 (7-21); 113206, 113240
(2-89); 113875 (6-3b); 113900 (2-87.1); 114424 (3-
1); 114811 (2-39); 115119 (7-73); 115131 (2-8);
115141(2-39);115545(3-7);116339(7-36a);116506
(3-7); 116808 (2-86); 116841 (2-114); 116868,
116931, 116933 (7-34); 117028 (2-39); 117588 (2-
56); 117617 (2-68); 117651 (7-50b); 117696 (1-3);
117776 (2-105); 117792, 117824 (3-10); 117839 (7-
25); 117921 (7-9.1); 118135 (2 sp.); 119155 (6-3b);
119388 (2-43a); 119397 (2-57); 119454, 119576 (6-
la); 119749 (7-36a);120004 (7-22); 120609 (2-69a);
120732 (7-74); 121338 (7-36a); 121636 (2-69a);
121703 (7-74); 121791 (7-36a); 121800 (7-22);
121858 (7-36a); 122046 (2 sp.); 122072 (2-106);
122405 (6 sp.); 122715, 122853, 122913, 123127,
123184, 123300 (7-36a); 123392 (3-7); 123659 (7-
68); 124598, 124638(7-36a);124748 (6 sp.); 124924
(7-36a); 125670 (2-88); 125859, 126219 (2-68);
129177 (3-4); 129367 (2-124); 130185 (2-57.1);
130873 (2-124); 130906A (2-99); 131173 (2-27b);
131203(3-4); 131406(6-3b); 131527 (7-25); 131653
(3-4); 131676 (7-56b); 131888 (2-68); 131889 (7-
56a); 131957 (2-81); 132163 (2-135)
Stoffers,A. L. et al., 114 (2-108); 140 (2-90); 178 (7-
56b); 222 (7-36a); 254 (7-73); 300 (2-108a); 404,
492, 513 (7-25); 517 (6-43); 3659 (7-56a);30143 (2-
108.1)
Strang,H. E., 208 (6-25)
Strudwick,J. J. et al., 3042 (7-56b);3347 (2-134);3587,
3589, 3614, 3689 (7-56b); 3766 (2-134); 3789 (7-
20); 4047 (7-56b);4245 (7-50a);4352 (7-20); 4395,
4450 (7-56b)
Sucre,D., 1053 (6-25); 3525 (7-36b);3949 (1-1);4275
(7-85);5730 (7-36b);6384 (7-82);7939 (6-25);9350,
9384 (7-56b); 9579 (6-25); 10248 (1-1); 10299 (2-
44); 11352 (7-36b)
Svensson,B. et al., 685 (1-1)
Sytsma, K. J. et al., 1551 (7-38); 3183, 3258, 3325,
3335, 3557, (7-56b)
Takeuchi,s.n. INPA7809 (2-22)
Tamashiro,J. Y., 6554 (2-30); 8765 (7-82)
Tamayo, F., 3541 (2-27a)
Tate, G., 870 (6-8.1)
Tavares,A. S., 114 (2-126.1)
Tawjoeran,J. A., LBB14443 (4-3)
Taylor,E. L., 1298 (7-18); 1301 (7-56a)
Teixeira,L. O. A., 43 (7-56b); 122 (2-36); 157 (6-34);
253 (2-53); 563 (7-56a); 728 (2-95.1); 798 (7-55a);
804 (7-56b); 927 (6-la); 975 (7-6); 1201 (7-50b);
1203 (6-9); 1222 (2 sp.); 1238 (2-69a); 1246 (3-4);
1365 (7-66); 1511 (7-56a); 1581 (2-60)
 List of Exsiccatae
Supplemental
Tellez, O., 2002, 2098 (7-22)
Terceros,W. et al., 21 (7-36a)
Terezo, E. F., s.n. INPA139843 (7-25)
Thomas,W. W., 3195 (7-30); 3575 (2-32); 3598, 3672
(7-56b);3747 (6-47);3808 (2-17);3810 (2-124);3836
(2-37); 3876 (2-144); 4091 (2-56a); 4095 (2-27a);
4154 (7-20); 4309 (2-30); 4324 (2-37); 4329 (7-20);
4404 (2-114);4457 (7-20);4506, 4523, 4525 (2-27a);
4572 (2-28); 4656 (7-66)
Thore, R. F. et al., 57938, 57955 (2-1)
Tidestrom,I., 4182 (1-1)
Tjon Lim San, R., LBB14813(6-1)
Todzia, C. et al., 2212 (6-9); 2226 (7-9); 2283 (7-6);
2295 (6-35.2); 2328 (2-129)
Torres,A.M., 1804 (1-1)
TorresC., R., 639 (6-47)
Torres, J., 85 (6-16); 93 (6-35); 290 (6-3b); 830 (2-
140); 919 (6-36)
Trigos,R. C. (see Cedillo T., R.)
Trinidade,L., 30070 (2-127)
Troth, R. G., 1117 (2-12)
Trujillo,B. et al., 3663 (7-74); 3765 (7-56a); 3789 (2-
12); 4497 (7-56a); 4534 (7-25); 4655, 5323 (2-12);
5518 (7-20); 5774 (6-3b); 5816, 5847 (7-56b);5940
(7-25); 6058 (2-26); 14980 (7-40); 15043, 15168,
15243 (7-25); 15262 (7-56b); 15316a (6-3b); 15338
(2-114); 15361a (2-88a); 15381 (7-56b); 15458 (3-
4); 15473 (2-69a); 16224 (2-39); 17354 (7-25)
Tunqui, S., 105 (3-9)
Tyson, E. L., 7367 (7-56b)
Ucan ek, E et al., 627, 1084 (1-1)
Uhl, C. F., 458 (7-56a)
Ule, E., 4214 (6-50)
Univ. Brasilia,TaxonomyClass, 189 (2-114); 519 (3-
16)
Uribe U., L., 127 (7-25)
Utley, J. et al., 5488 (2-18.2)
Utrera,A., 81 (7-49)
Valle, M. A., 46 (2-69b); 149 (7-56b)
Van der Werff,H. et al., 6998 (2-88); 9540 (2-13.1)
Van Hall, C. J. B., 47 (2-66); 49a (6-1); 49b (2-135);
51 (2-27a)
Varela,J. R. C., s.n. (1-1)
Vasconcelos,H. L., C2-7, H13 (2-69a); H16 (2-81);
H19 (6-28); J7 (2-76); J38 (7-50b); J53 (2-76); J59
(2-142); S20 (2-69a)
Vasquez,R. et al., 110 (6-44); 182 (2-27); 184 (6-3b);
1008 (6-6.1); 1102 (6-44); 1220 (7-56b);1547 (2-99);
1255 (6-44); 1309 (6-6a); 1392 (2-99); 2085 (6-6a);
3387 (7-56a); 3463 (6-7); 3444 (2-43c); 3472 (7-5);
3479 (7-40);3488 (2-69a);3572 (7-56b);3599, 3640
(2-27b);3696 (6-44);3759 (2-52);3762 (6-29);3860
(2-36a);3976 (2-26);3995 (2-69a);3997 (6-34);4000,
4011, 4023, 4043 (2-69a);4263 (6-3a);4312 (2-21);
4460 (2-39); 4492 (7-5); 4681 (6-36); 4788 (2-43b);
4973 (7-36a);5204 (6-36); 5255 (6-3b);5263 (2-52);
5267 (2-60); 5271, 5497 (7-53); 5637 (7-24); 5650
(7-56b); 5655, 5656 (6-35); 5698 (2-62); 5782 (6-
35); 5812 (2-39); 6124 (2-128); 6127 (7-5); 6135 (2-
88a); 6147 (2-69a); 6152 (2-94); 6166 (7-53); 6197
(6-62);6220 (2-88a);6243 (2-43c);6411 (3-4); 6435
(2-23); 6467 (6-44); 6622 (2-22)
Vaz, A. M. S. F., 158 (7-18)
123
Veillon, J. P., 2/v (2-27); 57 (7-36a); 131 (2-45)
Velasquez,N., 28 (2-45)
Vellow, 299 (3-4)
VenturaA., F., 976 (7-36); 19786 (7-36c); 20011 (7-
36a); 20203 (7-36c)
Viera,M. G. et al., 8, 13 (7-56a); 72 (2-124); 123 (6-
3a); 134 (7-50a); 151 (7-20); 281 (7-56a); 633 (2-
41); 750 (2-57); 877 (2-114); 929 (2-82); 949 (6-27);
977 (2-17); 995 (4-4)
Vilhena,R., 46 (7-56a); 143 (7-50a); 144 (6-3a); 147
(4-1); 211, 266 (7-50a);289 (4-1); 304 (2-27b);305
(6-3a)
Vincelli, P. C., 532 (2-88); 586 (1-1) 966 (7-25); 1012
(2-124); 1055 (2-69a); 1066 (2-39)
Vinha, S. G. da, 12 (3-11)
Vital, D. M. et al., s.n. (1981) (7-54)
Voeks, R., 28 (2-130.1); 53 (6-51b);72 (7-37)
Vogl, C., s.n. (3-7)
Wachenheim,115 (2-69a)
Wagner,R. J., 1747 (7-83)
Warer, R. H., 299 (2-148)
Webster,G. L. et al., 9806 (2-69a)
Wendt, T. et al., 3302 (7-36c); 3325 (2-15); 3724 (7-
36c); 3899 (7-56b)
WesselsBoer,J. G., 2073 (2-109); 2074 (7-56a);2317
(6-13); 2334 (7-21); 2405 (7-62)
West, E., s.n. (2-44)
Wherren,L., 16 (7-73); 105 (2-98)
Whetstone,R. D., 13350, 14376 (2-1); 14501 (1-1);
14514 (2-1)
White, S. et al., 460 (7-22)
Whitefoord,C., 2761 (2-88a); 2828 (7-56b); 3084 (1-
1); 3151 (7-56b);3290 (7-36a)
Widgren,J. F., s.n. (3-4)
Wilbur,R. L. et al., 10847 (2-14)
Williams,LI., 3173 (6-16); 12510 (2-27); 14914 (4-1);
16049 (2-69b)
Williams,L. O. et al., 15962 (6-3b); 26513 (2-45)
Williamson,C. S., s.n. (2-1)
Witherspoon,J. T. & F., 8543 (7-80)
Witsberger,D., 813 (2-9)
Wolfe, F., 12181 (7-36a)
Wood, C. W., 292, 410 (2-145)
Woodbury,R. O., s.n., 30464 (7-83)
Worthington,R. D., 13561 (1-1)
Woytkowski,F., 5773 (2-64); 6318 (6-44)
Young, K. et al., 146 (7-56a); 1037 (7-75); 1047 (7-
56a)
Zaandam,C., 6764 (6-la)
Zabala,A., 80, 102, 162 (7-16)
Zanoni,T. et al., 12668, 15904, 17737, 21206, 25782
(1-1)
Zappi,D. C. et al., CFCR8476(2-43a)
Zarucchi,J. L. et al., 1221 (6-9); 1278 (2-64); 1944 (3-
6); 1983 (7-25); 2038 (7-25); 2548 (7-8); 2574 (2-
57); 2580 (2-124);2585 (7-20); 2858 (7-1); 2862 (7-
9); 2920 (7-56d); 3079, 3100 (35.2); 3162 (2-129);
3174, 3186 (2-140); 3396A (2-69b); 3401 (2-27b);
3452 (2-83);3491 (2-69b);3533 (2-39); 3529 (7-40);
3583 (3-4); 3626 (6-36); 3630 (2-24); 3634 (7-56b);
3666 (6-3b); 3667 (2-69b); 3689 (2-140); 3698 (2-
69a); 3773 (3-4)
124
SECOND SUPPLEMENTALLIST OF EXSICCATAE
Acevedo, P. et al., 1607 (7-2); 1657 (2-13.1)
Acosta P., R. et al., 1653 (7-56b)
Alvarez,H. J. et al., 34 (1-1)
Amaral,I. L. et al., IG2-6-173, TF-2-15 (2-57)
Aumeeruddy,73 (2-114); 92 (2-109)
Axelrod, F., 610 (7-83)
Ayala, F. et al., 2888 (6-16); 3023 (2-43b); 3572 (7-
56b)
AymardC., G. et al., 4825 (2-68); 4872 (7-49); 5355,
5425 (7-25); 5552 (7-36a)
Bal6e,W. et al., 2658 (7-24); 2737, 2829, 2836, 2836,
2882 (6-la); 2894 (2-69a); 2902, 2925, 2928, 2929
(6-la); 2943 (2-67); 2982, 3004 (6-la); 3018 (6-lc);
3031 (7-56b); 3056 (6-lc)
Bamps, P., 5478 (6-27)
Barrier,S., 3863 (7-16); 3989,4259, 4814 (2-77);4976
(7-20);5003 (7-50b);5022 (2-114);5183 (7-16);5191,
5058 (2-77)
Beck, S. G., 10013 (2-69b); 10160 (2-69b)
Billiet, F. et al., 1457 (2-135); 1458 (7-18); 1809 (2-
109)
Boom, B. M., 6838, 6881 (7-83); 7119 (2-135); 7127
(6-7);7136 (2-78);7142,7145 (6-42a);7363 (2-108);
7529 (7-74)
Brant,A. E. et al., 1055 (7-56b)
Calzada,J. I., 5997 (7-56b)
Castillo,A., 1340, 1518 (7-25); 1653 (2-81); 1859 (2-
137); 2094, 2173 (7-25); 2208 (6-34); 2305 (2-83);
2306 (2-121); 2317 (2-83); 2414 (7-56b);2438 (3-4)
Cid Ferreira,C. A., 5463 (7-50a); 5490 (2-43b); 5668
(7-28);5706, 5762 (2-43b);5847, 5848 (7-56b);5981
(7-1);6660 (2-62);6674 (2-124);6689 (2-129);6696
(2-52); 6737 (2-97); 6833 (6-7); 6844 (2-75); 6850
(2-81); 6894 (2-39); 6897 (3-4); 6903 (2-27); 6911,
6916 (3-4); 6917 (2-8); 6946 (7-11); 7033 (6-la);
7064 (6-lb); 7070 (2-27b);7078 (3-4);7105 (2-140);
7143 (2-119); 7145 (7-55); 7160 (2-27b); 7185 (6-
44); 7209 (3-9); 7248 (6-44); 7254 (2-69b);7262 (2-
124); 7291 (3-9); 7326 (6-3b); 7650 (2-134); 7663
(2-93); 7681 (3-4); 7703 (2-93); 7709 (7-56a); 7719
(7-39); 7750, 7766 (2-36); 7770 (2-83); 7783 (3-4);
7795 (6-3b);7812 (2-36);7820 (2-88); 7856 (7-50b);
7857 (6-la); 7863 (7-56b); 7917 (4-3); 7949 (6-la);
7957 (7-56a); 7968 (2-134); 7978 (6-3a); 8039 (2-
134); 8041 (2-52); 8054 (6-19); 8073 (2-97); 8143
(3-4); 8167 (2-75); 8180 (7-52); 8183 (3-4); 8190 (2-
69a); 8194 (7-9); 8214 (7-1)
Coradin,L. et al., 5809 (7-18); 6132 (7-56b)
Cremers,G., 4664 (2-63); 9945 (7-2)
Croat,T. B. et al., 64503 (7-56b)
Daly, D. C. et al., 4348 (8-4); 4387 (6-44.1); 4390 (2-
27b); 4408 (6-9); 4461 (2-69a); 5210 (2-4.2); 5426
(2-69a); 5447 (2-22); 5565 (7-55); 5568 (4-1); 5635
(6-34); 5670 (2-46b)
Davidse, G. et al., 30899, 30941 (1-1); 31993 (7-56b);
32033, 32393 (7-36a); 32745, 32934, 33165 (1-1)
Descoings,B. et al., 20068 (6-7)
Diaz S., C. et al., 2483 (6-3a)
Dodson, C. et al., 14621 (6-la)
Dubs, B., 44 (6-27); 237 (2-28); 351 (6-20); 358, 398
(2-18); 458 (2-28)
FloraNeotropica
Faber-Langendoen, D. et al., 287 (2-124); 352 (2-96);
439 (7-56a);572 (2-148); 660 (2-144.1); 663 (3-17);
689 (2-46);693 (3-15);867 (2-148);883 (2-124);979
(2-148); 1084 (2-51)
Farney,C. et al., 1218 (6-46); 1451 (6-13.2)
Fernandez,A., 2942 (7-25); 3311 (2-68); 3339 (7-28)
Fernmndez Casaset al., 4065 (2-41)
Feuillet, C., 1089 (2-77); 1374 (2-135); 1388 (3-1);
1419 (2-124); 1445 (2-69a); 2262 (7-56b); 2303 (2-
124);3579 (7-25); 3809 (7-2); 3866 (7-52); 4034 (7-
25); 4297 (7-2)
Filho, A. C. et al., 4672 (3-8)
Fleury,M., 327 (2-124)
Foresta,H. de, 228 (2-149); 618 (7-52); 733 (2-69a);
750 (2-134)
Forget,279 (7-25); 323, 324, 327 (2-77); 377 (7-74)
Fournet,A., 211 (7-52)
Foster, R. B. et al., 11562 (7-21)
Garcia,R. et al., 335 (1-1)
Garcia-Barriga H., 20911 (7-8)
Garwood,N. et al., 231 (7-36a)
Gentry,A. etal., 25069 (6-35); 53381 (2-32.1); 53676
(3-15); 53738 (2-5.1); 53780 (2-148); 54188 (2-64);
54361 (2-5); 54458 (6-22); 54965 (2-43b);56813 (2-
144.1); 56842 (7-35); 56884 (2-148); 56892 (2-40);
56919 (2-46); 56947 (7-35); 56973 (2-124); 57015
(7-10); 57018 (2-46); 57062 (2-148); 57063 (2-124);
57516 (7-56a); 57675 (7-75); 57677 (7-56a); 57951
(7-75)
G6mez, L. D. et al., 23350 (7-22)
G6mez, S., 178 (7-22)
Granville, J. J. de et al., 604 (7-73); B.3787 (2-67);
5484 (2-56a); 5719, 5722 (2-77); 6121 (4-3); 6183
(2-77);6423 (2-67);6641 (2-27a);7366 (7-73); 7407
(7-2); 7467 (2-88); 7651 (7-52); 7786, 8004 (7-74);
8019 (2-88); 8035 (7-52); 8087 (2-77); 8144 (2-88);
8201 (2-77); 8678 (7-74); 8820 (7-2); 9072 (6-7);
9184 (7-52); 9254 (7-56b); 9638 (6-lb); 10243 (7-
25)
Grayum,M. H. et al., 5731 (7-36a);6252 (7-56b)
Guanchez,F. et al., 4575 (2-83); 4606 (2-43a); 4796
(2-69a);4799 (6-3b)
Haase, R., 634 (2-69b)
Hammel, B. et al., 7972 (7-61); 11048 (2-88); 14427
(2-13.1); 14490 (5-1); 14586 (7-36c)
Hartshorn,G. et al., 2919 (7-50a)
Henderson,A. et al., 461 (2-115)
Heringer,E. P., 17167 (3-16)
Hernmndez,L. et al., 128 (2-107); 175 (2-68); 479 (2-
107); 550 (7-20)
Hoff, M., 5002 (1-1)
Hoist, B. et al., 2640 (7-74); 3046 (7-68);3052 (7-56b);
3199 (2-83); 3260 (7-25); 3436 (2-69a)
Huber, O. et al., 11774 (7-26); 11778 (2-57); 11798
(3-10); 11973 (6-7); 12062 (2-69b)
IbarraM., G. et al., 2284 (7-36c)
Jacquemin,H., 2016 (7-50b)
Jansen-Jacobs,M. J. et al., 75 (7-56b); 136 (6-43); 210
(2-134); 231 (7-56a);238 (2-135); 347 (7-36a);495
(2-27b);496 (6-15)
Jaramillo,J., 7457 (2-4.2); 9036 (2-27a); 9066 (7-22)
Second SupplementalList of Exsiccatae
Keel, S. et al., 264 (6-3b)
Kral, R., 72077 (7-49)
Kubitzki, K. et al., 79-105 (2-88); 79-43 (7-20); 84-
293 (6-3b)
Kvist, L. P. et al., 187A (7-51)
Lane, C. & R. Gieschen, 50 (1-1)
Le6n, H. et al., 1347 (7-35); 1561 (7-10)
Liesner,R. et al., 21204 (2-83); 21216 (7-21); 21384
(7-68); 21511 (7-21); 21836 (2-83); 21839 (7-25);
21873 (2-105)
Lima, H. C. de, 2689 (6-41); 3155 (4-5)
Lindeman,J. C. et al., 784 (2-108)
Maguire,B. et al., 37515 (2-119); 56834 (6-27)
Maciel, U. N. et al., 720 (7-36a); 810 (7-75); 823 (7-
20); 829 (7-18)
Martinelli,G. et al., 11935 (6-46)
McPherson,G., 7449 (2-31.1); 10024 (2-18.2); 10222
(2-88); 10272 (7-56b); 10278 (2-88); 10588 (6-13.1);
10610 (2-5.3); 10647 (2-31.1); 10804 (2-88)
Miralha,J. M. S. et al., BO-1-81, BO-1-114, BO-2-
190 (2-43a)
MonsalveB.,M., 1128(2-5.1);1278(2-40);1297, 1308
(2-124); 1503 (2-148)
Moraes,M., 513 (7-66); 524 (7-73)
Moretti,C., 872 (6-7); 925 (2-86); 985 (2-69a)
Mori, S. A. et al., 15152 (2-64); 18484 (7-74); 18534
(7-50b); 18522 (6-34)
Nee, M., 31393 (7-56a)
Neill, D. et al., 7269 (7-53);7217 (2-45);7462 (7-50b);
7471 (7-56a);7667 (2-5)
Nelson, B. W. et al., 301 (7-9, 7-56a); 445, 454, 467
(6-3b)
Nevers, G. de et al., 7519 (2-31.2); 7589 (2-88)
Nunez, P. et al., 5370 (7-36a); 5828 (7-66); 6181 (7-
75)
Oldeman,R. A. A., 1535 (2-134); 1825 (2-63); 2433
(2-69a); B.2819 (2-27a); B.2278 (2-86); B3119 (2-
87); B3127 (2-27.1); B3559, T655 (6-lb)
Padilla,F., 191 (7-22)
Palacios,W. et al., 1122 (2-64); 1159 (7-53); 1269 (6-
36); 1484 (7-50b)
Pennington,T. D. et al., 12230 (6-36)
Pimentel,J. & R. Garcia,97-A (1-1)
Pipoly,J. J. et al., 8024 (2-77); 8146 (7-56a);8179 (2-
69b);8181, 8241 (2-108);8337 (2-86);8428 (6-42a);
8429 (6-15); 8432 (2-135); 8438 (6-42a); 8444 (2-
73); 8445 (2-135); 8462 (6-42a); 8583 (2-90); 8589
(3-12);8597 (2-99);8815 (2-69.1);8704 (2-90);8849
(2-68); 8850 (2-69a); 8853 (2-109); 8929 (2-69a);
8930 (2-68); 8934 (2-112); 8940 (2-124); 8942 (2-
109);8945 (2-124);8955 (2-108);8959 (6-15); 8962
(2-108); 9140 (2-135); 9159 (6-15); 9334, 9394 (7-
25);9432 (3-1);9536 (2-93);9562,9581 (2-78);9596
(6-15); 10741 (2-69a); 10842 (7-21)
Prevost, M. F., 1201 (2-114); 1301 (2-134); 1574 (2-
69a); 1698 (3-1); 1806 (7-2)
Proctor,G. R., 41577 (7-83)
Pruski,J. et al., 3205 (7-1); 3352 (2-63); 3381 (2-18);
3354 (7-56a)
Puig, H., 10256 (2-35)
Queiroz,L. P. de, 1825 (7-56b); 1827 (7-20)
Ramirez,J. O. et al., 28 (2-114)
Riera, 49 (2-77); 450 (6-15); 514 (6-la); 586 (7-50b);
125
587 (6-7); 589 (2-35); 624 (2-69a); 671, 731, 735,
766, 767, 768, 769, 770, 804 (2-77); 830 (1-1); 841
(3-3); 872 (6-la); 890 (2-77); 1090 (2-109)
Rivas, R. M., 122 (1-1)
Rivero, R., 883 (7-36a)
Rodrigues,W. A., IG1-8A-589 (2-57);IG1-10-464 (6-
15);IL8-22 (2-119.1)
Roosmalen,M. van, 12 (7-73)
Rosa, N. A. et al., 4287 (7-24); 4392 (3-4)
Rosales, J. et al., 15, 17 (6-3b); 19 (2-27a); 33 (7-25);
52 (7-56b); 84, 109 (6-3b)
Rosario,C. S., 96 (7-24); 101 (7-56a); 137 (7-18)
Rutkis, E. & K. Udris, 634 (3-7)
Sabatier,D., 88 (2-27.1); 165 (2-86); 484 (2-77); 509,
572 (7-15); 644, 676 (7-15); 725 (2-77); 830 (7-74);
849 (2-134); 866 (2-95); 893 (2-86); 925 (2-77); 939
(7-16);1003(2-63);1022(2-124);1028(2-27.1);1066
(2-97); 1072 (7-73); 1104 (2-99)
Santos,J. L. dos et al., 681 (2-119.1); 717 (2-131);732
(4-1); 758 (2-119.1)
Sastre,C. et al., 8016 (7-25); 8129 (7-56b)
Sauvein,M., 218 (2-8); 401 (2-77); 589 (7-56a)
Serv. Forestier(Fr. Guiana),85M (2-124)
Sherman,C., 151 (7-22)
Silva, M. G. et al., 3122 (3-4); 4684 (2-64); 5827 (2-
88); 6216 (7-56a);6301 (2-30)
Silva, N. T. et al., 4594 (2-128)
Skog, L. et al., 7476 (7-18)
Smith, Damon, 193 (7-61)
Smith, S. F. et al., 1074 (7-66)
Sobel,G. L. et al., 4568 (2-69a);4569 (2-39);4643 (2-
141); 4671A (2-69a); 4673A (2-124); 4704 (2-52);
4759 (2-39);4764 (6-36);4818 (2-39);4836 (2-27a);
4860 (3-4)
Solabarrieta,S., 155 (7-22)
Stein, B. A. et al., 3936 (6-44); 3945A (7-5)
Stergios,B. et al., 5073 (2-69a);7319 (7-40); 7419 (2-
102); 7472 (2-69b); 7473 (3-6); 7549 (2-140); 7550
(7-56a);7580 (2-24);7609 (6-36);8066 (2-24);8084
(7-40); 8157 (2-27a); 8175 (2-69b);9863 (2-68)
Stevens, W. D., 24575 (2-88); 24627 (7-36c); 24628
(2-95)
Steyermark,J. A. et al., 125689(2-77); 125863(2-27a);
126228 (2-77)
Taylor,C. M., 7633 (1-1)
Thomas,W. W. et al., 4926 (2-43b);4951 (7-56a);4969
(7-53);5005 (2-43b);5050 (6-15);5070 (7-50b);5163
(2-17); 5192 (2-99); 5194 (7-56a); 5275, 5258 (2-
99); 5340 (7-9); 5343 (2-69a); 5414 (7-56b); 5439
(7-42);5442 (2-99);5456 (7-56a);5458 (2-88);5612
(6-27)
Thorne,R. F. et al., 48111 (1-1); 48566 (2-1)
Torres,G. A., 117(1-1)
TorresC., R. et al., 86 (7-56b)
Valdespino,I. A. et al., 254, 377 (2-88a)
Vasquez,R. et al., 2411 (7-53); 2412 (2-10); 2729 (2-
5); 2788 (6-41a);2808 (6-7); 2821 (6-35); 2898 (2-
26); 2949 (2-52); 3168 (7-40); 3213, 6050 (7-36a);
6648, 6694 (2-8); 6769 (2-124); 6780 (6-6.1); 6832,
6856 (7-5); 7150 (2-144.1); 7192 (2-62); 7446 (2-
69a);7457 (2-39); 7517 (2-69a);7610 (2-140);7615
(2-88);7622 (7-41);7628 (7-40);7660 (2-140);7679
(7-5); 7868 (7-56); 7909 (2-27a);7916 (7-40); 7926
126
(2-69b); 7963, 8031 (2-69a); 8098 (6-36); 8112 (6-
41.1); 8177 (2-5); 8188, 8203 (2-43b)
Ventura,E. & E. Lopez, 781, 811, 838 (7-56b)
VenturaA., F., 20795, 21298, 21541 (7-56b)
Vera, S. E. N., 215 (2-68)
Viellescazes,A., 447,471, 502 (2-77);513 (7-50b);534
(7-56b)
Villiers,J. J., 2246 (2-114); 2729 (7-15); 3719, 3780,
3781, 3786, 3787, 3788, 3797, 3830, 3843, 3859 (2-
77); 3884 (7-16); 3947, 6064 (2-77); 6603 (7-16)
FloraNeotropica
Wendt,T. et al., 3373 (2-32)
Werff,H. van der et al., 540 (2-13.1)
Woodbury,R. O. et al., 1-71 (1-1)
Zanoni, T. et al., 11795 (1-1); 29406 (6-36a); 32064,
32446 (1-1); 32568 (7-36a)
Zarucchi,J. L. et al., 3593 (2-69b);4889 (2-88)
Zaruma,J., 558 (7-53); 753 (6-36); 809 (7-50b)
Zona, S., 136 (2-1)
90 80 70 60 50

0.b o o s
50m --------
128 Flora Neotropica

_? ^^*60 . ._____
__60_

cs'
V-' C. cuspidatus C. venezuelanus
IS ?C4
FIG.
2.DtitooCrsa ""lau u C e0

_ _ _ _ _ _ _ _ _ _ _0 0

0 HABITAT 0 HABITAT
Slope forest _ ______Terra firme forest _ |
AO fNo
JFIM1AI- stibuJti C a ui C j nAISIOnD
JF MAIMnd
FRUIT
FLOWER_I
I
I I
I I III
I 1 II I - I
I/
1
IT
IIIIFRU
IFLOW W ER
I
r

60
_60
FIG. 21. Distribution of Chrysobalanus cuspidatus and C. venezuelanus.
Distribution Maps 129

80 70 60 50
0
/^\ ^ L. affinis
0

1 o

10

FRUIT I

80 70 60
80 70 60 50

I\ ~L. alba
0
0

'" -^
HABI.itAT.o~ -y
Terra irme forest

FRUITM*ii I /..I . . ..
80 70 60 b

FIG. 22. Distribution of Licania affinis and L. alba.

130 Flora Neotropica

60 50 60 50
0o L. albiflora 0 1 L. amapaensis
0

00 ~ ~ ~ ~ ~ 0
0
1S- --------------- ~------

RUIT aifra L.

10

70 60 50

FIG.23 DstbutionofLicaniaalbilora,L.amapaensis, andL. anneae

L. angustata,
~
HABITAT Fi-- .\ ....
HABITAT
.

Terra firme forest - Seasonally floode orest

J FIM AMIIJIJIAIS INDJ IFIMIAMIJIJ IAIS OINID
1~~FRUIT~
'1J
CFLOWER:::^:
~ ~ l ~~ ~
~FRUIT
JY~ 10
I I I I I I I I
ilIig
/ i
FLOWERi iI II i L_
70 60 60 __ 50

FIG. 23. Distribution of Licania albiflora, L. amapaensis, L. angustata, and L. anneae.

Distribution Maps 131

70 60 50 40
Ic o- L. apetala var aperta
L ^ CC^""'^

ol

Savanna margins _____ + vqf \ / \ /} ( ; /- \^ 10

Seasonally flooded forest \' v^- |/ /'1 5 \\\/) \ /^
JF\FM/A/ J/J A IOII I I
FRUIT d : \
FLOWERI************
X| s\< | / \ N^ v

70 60 50 40

Savanna
0 apetala var. apetala
~--L.

10 TT^ i
( 'u0

FFABITAT 24. Distribution of.Licania.apet

Beaches
Gallery forest" }\
Savanna margins
Seasonally flooded forest
J
"JFMIAIMJJ IAISO0 N Q .......
-
FLOWER;:
70 60 50 40

FIG. 24. Distribution of Licania apetala.

132 Flora Neotropica

60O 50 60 50
__

' L. apiculata L. aracaensis

.----- 1 0 lC?--- , 10
_ _ _ _ _ _ Io . _ _

I' i
"I iI

ABITAT I BITAT I

Beaches ( , Montane & Cloud forest (.

7JF_MAjJ J A S CN1Q
_ ( J FIMAIMIJIJIASIOINIDI
FRUIT I FRUII II I I ) 1

FIG.
2 D , a.. . ar

50w )50

Terra firme forest __ /\ \^ & S (

L ~ JIFIMIAiM.JoJIA
" _ S_ND. \ .../ 2. JF.MAIMJTJ
IJAIS.N
T
' 0FRUIT 1111~1 1 11 1 1 1 / I LIFRUIT
:l i
11 1I " 1
^\ FLOWERR 10
1 L I I/____ 1
70 60 50

FIG. 25. Distribution of Licania apiculata, L. aracaensis, L. arachnoidea, and L. araneosa.

Distribution Maps 133

100 90 80

-.^r^^'f "VL. *arborea

0 / o

0 0aOo:

?~~~~~~~~R)rj~~~~~~~~~~~~

Dry forest, semideciduous

FIG. 26. Distribution
\ of Licania arborea. ? ?
SIOND
IJIFIMINJIJIAIS h A

100 90 80 70
134 Flora Neotropica

50 40 50 40
arianeae L. bahiensis

.
-------- - -- : I
./-~. ..- -^-.."'"^^' ' l

0r

T r
JIF_IMIAIMJIJIAISIOINIDI
forest Terra firme
FRUIT | | | | i | I I I i i i | 'O HABITAT \ :|/
F IND I
IVJ IJJASS
2 .FLOWER I I I I II! I I 2:1 E

? ...MIA~JIJAI SIOINlol
,.f......I\.SIN I I I JFABWJJAT

1Te firm forest___...... ~~';Terra

- firme forest -

2CFLOWER * ~"~"~ZIIg'0 n ^
--H--444--^---J>:FW
I 11 1

\ V - I0_- //IFRUIT I

40 50

FIG. 27.
FIG. of Licania
Distributionof
27. Distribution Licania arianeae, L. bahiensis,
arianeae,L. L. belemii,
bahiensis,L. belemii, and L. bellingtonii.
and L. bellingtonii.
Distribution Maps 135

70 60 50
Vo o70 ITX ?_60' / blackii
5 h_L.

Gallery forest

Seasonally forest
flooded r~~~~~~~~~~~~~c~',-
\ ) / \ (
'"
JFMAMJJ
M I ..
ASOND
J......?/.. JK s -? / .
FRUIT 0--^
| g l | | | | | 0 1@1@11 ) /\ s X -. |.. .
m., J (
FLOWER :::ggi^ ,. ylUUl | , J r i >
70
- ___60 50
70 60 60 50

boliviensiso
",^L. L. boyanii

28 sr
FIG. Di o f btoLi c ab , n.byni
icii L.blvess

Terra firme forest White sand forest or campina > (

FRUIT.AT 2 D it i a FR
IT,. b nJ L. b n
FLOWER MN III :II 1g _ >s FLOWER I _:.................
*
~ bU
770 -50
FIG. 28. Distribution of Licania blackii, L. boliviensis, and L. boyanii.
136 Flora Neotropica

80 70 60 50

T ?" - --
^ L. bracteata
IIw ' |

"JIF AIMIJIJIAISIOIN
_ - D Y 0\ ( S ?
FRUIT I I /I I I , I ,\
FLOWER, * *- 1T1z1I _ y

80 70 60 50

L. britteniana

Terra firme forest \ / ///

Seasonally flooded forest \

80 70 60 50
80 70 60 50

cli S^X -' _. -L. buxifolia 0

White sand forest or campina _ -' / /- .'- \

-
FLOWER I" _ __ \;\( > ---n
_- 8U - 70 i60 U 0

FIG. 29. Distributionof Licania bracteata,L. britteniana,and L. buxifolia.

Distribution Maps 137

L. cabrerae L. caldasiana

o0o

H 0

Montane & Cloud forest T o te (

JF MAM,J A S C5ND J
FRUIT | _ | X X IR T \

70 80 70

0
''''^^ 71
FL OWE-R I I I FLOWER~~~
8070807

. 80 80 70
__ __0 70__

FIG. 30. Distribution of Licaniacabrerae,L. caldasiana(exact locality in Colombia unknown), L. calvescens,

and L. cecidiophora.
138 Flora Neotropica

70 60 50

L..-. aL.canescens

/
Sersonally f forest
looded X) 1 / x \ f \
I-
, S^Mi^AsonD/
~\\ZZZL / '
.....................
^.^ , p'

Licaiaan1e FI.3.Dstiuino

FRUIT _*_* ____** .. l \ . /

FLOWER___W * __ '* / \

FI.3.Dsrbto
FIG. 31. DistributionoffLcni aecn
Licania canescens n L..cuaa0
and caudata.
Distribution Maps 139

60____80
70

^L.
^10 coriacea 0
;yL. costaricensis

_ 60 0_ 800

o _ __ - _ __ _ __ _ 6 _
_0__ 6
3
oTerra rine forest ain A c i

6L0ueioi0 L

FIG I. IIoI o e I. I I I I a L a i

FIG. 32. Distributionof Licania co uepiifolia, and L.

crassivenia.
140 Flora Neotropica

80 70 80 70

L. chiriquiensis L. chocoensis

6^ v
0^', ) " 0

HABITAT
Montane & Cloud forest __ Terra firme forest'-

U /."1"1 FRUIT
III I IF
I U I-

60II 6070

0 i- L. compacta I o

. -.10..

0
0 6 :

FIG. 33 Distbutionof Liania hiriquiensis,L. hooensis, L. ompata, and L. ordata.

F~~IG~~',.
3.D b f a u L.chocesavannas,Loata.

FIG. 33. Distribution of Licania chiriquiensis, L. chocoensis,

compacta, L. L.andcordata.
Distribution Maps 141

_70 60 70 60

L. cruegeriana L. cuatrecasasii

o o~~~~~~
o
Terra trine forest M & Clu fores
o~~~~~~~
|FLOWEFI| I II I 101E
_1 1 I A
_^g_
60 5 70 6

I B I ^ ^ "
00 ? 7"I ^0 6 0

FI.3. itibto f iaiacugein,L.careaaii .cpra ndL usiaa

FIG. 34. Distributionof Licania cruegeriana,

cuatrecasas,i, L. L. cuprea,and L. cuspidata.
142 Flora Neotropica

L. cyathodes L. cymosa

HABITAT /~J/ ~~I~-~

'THABITAT" f
Seasonally flooded forest Terra firme forest
) I |
IJIFIMIAIMIJIJIAISIOINIDI DI
IJIFIMAIMIJIJIAISOINI
FRUIT I I I i1 I 1 I- 1- / - IFRUIT II I I II
[FLOWERI
1 Il I I I I I I

L. davillifolia L. dealbata

Terra firme forest HABIT

Seasonally flooded forest Cerrado
FI I3 IAISION Di
I AIMIJJ
J- thJoFdL.ALMsJaJ
IAaSIOA L
NID
FRUIT I A 11 I / i FRUIT I I
FLIWERFLOWER
:: :g lI !_1FLOWER : g I I

FIG. 35. Distribution of Licania cyathodes, L. cymosa, L. davillifolia, and L. dealbata.

Distribution Maps 143

_60 _60 50

Terra firme forest __ ( \ on

ntTerra
M fnrme forestudforest
JIFIMIATJIJIAIS / /)
IONID \0 IJIFIMIAI^^J
IIAISIOINDI /-^1
I E * I FRUIT II I I I I1 1
10 FLOWER I1Ig g11 1, 1 X,^ i . FLOWER _
60 60 '*60 50

FIG.^^ L. divarica ra . ds durifolia

1 I1ABIT
AT

F 60 50 8070or Ld_ c L d l

Savanna_______________ It) / I ABITAT

Terra firme forest- e
rrMontane &
Cloudf forest _
-
JIFIMIA JASND--
0JJ JAFIM- NJIMAAs,/y -\ / , AM ,J - D /
FRUIT j IFRUIT ----I I --I-I/I
FLOWERE
*gg g l WWI ....... I FLOWER I
gI mm IgI
60 50 80 70

FIG.36. Distribution
Lica0ia of andL. durifolia.
densiflora,L.discolor,L. divaricata,
144 Flora Neotropica

80
- 70 ' 60_50
--0
's"
i
f o; .
' L egleri

HLABITAT \ (''"*/ ^ ......

i^r-^Q......... . . . .
Secondary forest
U Terra firme forest
t, :
g ?, - /
-)
,
.W /
- )
1
_ < - ,_ .\- -
Seasonally flooded forest / ^ > I \ I t /
_/A _J J AS
JFMAM *N D
1D v \ . f|

_
o
^
Ter im oet __^ ^- U
j \ \ \
- -JFAM
-- -
jA'O D*'I
^ / /\ N i^K/ ,f) ^ V-
FR T -g /--< \\ '\ I L^?/

-
FRU I oI.A
I II Ig |^, ( I \ ,) )_

80 70 60 50

FIG. 37. Distribution of Licania egleri and L. elliptica.

Distribution Maps 145

0 60 5 p
7______0 70

?
/^P^; 10^^^-__ 1

Terra|irmeforetTera firme forest

_LEFIMAIJ I L.8070IJ
I J
80 7IF0MI
fasciculata
0
lNIDI I
r H~ fir \
I.ABIT
L.- lomenoi|
HABIT 11\MAM^i^SQNDHABITAT
HABITAT
AT JATf rT^AJ )\\ (

FIG.38. Distribution of Liania L..

emargnata, fanshaW::i ::L.fa:ilata,andL. ..........
70 60 60 --0
80 70 80 70

L. fasciculata L. filomenoi.
.1

UV'^ / ~ ":;t'RUI
~ ~' ~ ~ ~~~T
~X

FRUIT |\ | |firme
|Terra | forest
/
Terra firme forest-
SIOINID
HABIT AT =2JIF5yAJAZ1A
Terra firme forestFRUIT I
FLlOW iER

FLOWER
I - \ ' " ^0
7
80
____7 0 80 70

FIG. 38. Distribution of Licania emarginata, L. fanshawei, L. fasciculata, and L. filomenoi.

146 Flora Neotropica

70 60 70 60

L. foldatsii L. foveolata
10 o
2

,~
'-^-^-^, .....-'-^-f i,'^' (e^^^--^J--------- -

Savanna ____ _ .. HABITAT

.
Savanna margins _/ I Slope forest
JIFIMAIMIJIJIAISONIDI J IFIMAIMJIJIASONrISI N
/ .
FRUIT |Aile | | [ 1I \ FRUIT
.

70 -0 70 60
60 50 80 7

oo L. fritschii L. fuchsii

10 0o

!U8_______7
?

0IG 3 Dtuof fot,fvaLftiaLf

HABIT
Seasonally flooded forest Terra firme forest

FRUITI9 ID I I I oI I I f dI f aI .fr I I
FLOWER101 I II I1 I i i!1I W
LO ER II I
80 70

FIG. 39. Distribution of Licania foldatsii, L. foveolata, L. fritschii, and L. fuchsii.

Distribution Maps 147

10

7!F-

70 ~.0. 60 50 0 07

Seasonally flooded forest

flooded forest '( I , Terra firme forest / / /
Seasonally y

FRUIT II
FLO
WERI Jll .I..!.i. !
6 0 50 5 80 70
IlFRIT f |I | | | | ||II T _ ___

FIG. 40. Distbuton of Lcanfurfuracea L. gentryiand L. gardneri.

148 Flora Neotropica

. 70
...7 Q

I eL. 4 glabriflora

FRUIT ^UIT .---__*___ _,^

* -<^ ^ | | r T L
FLO W ER I I I 4 i I I _ _ _
_ __
080 70 60 50

L. glauca L. glaziovianal
10

* 0 54040
8^70;~

FIG. 41. Distribution of Licania glabriflora, L. glauca, and L. glazioviana.

Distribution Maps 149

^ B
i_m
11Q___ ______ M60
L. gonzalezii .. L. gracilipes

FRUIT_ _RUIT T__i__ / I

FLOWER
_i llI _ 1 1i| II !
|..OWER..... - _
110 1(0 _ _60
80__________70 70 60

L. grandibracteata d-- L. granvillei

011
FLOWERI
I 14 I I! I 1 FOWER::

JIFMI 1MJ JIISO II D h(--l----- f ol77ArS~ i

FRUIT I I I1 I II I I I I~j I-IIRU IT I j~ I I IJr-' ~cr 0

FLWE J II 1 I1 I1~ "~~..:: ??Lnl

IIL0 E

0 ~ ~ 0 1 7

0I.4.Dsrbto fL ozlzi .gaiie,L rnivcet,adL rnili

80 ' 70 670
7' _

FIG. 42. Distribution ofL. gonzalezii, L. gracilipes, L. grandibracteata, and L. granvillei.

150 Flora Neotropica

80 70 60 50

'^ % ,? ? L. guianensis
,
0 .

90 0

0 z /t X
=60(09(0000 6

L. guatemalensis, L. harlingii
and

Terra firmmeforest / \

HABITAT HABITAT
Terra firme forest Terra firme forest

- n o Ies
J aJAISOINDI
JIFiMAuIMI L a i an
LA.MJA lng
I FRUIT
FRUIT I I _ T/
/
V / Il I
FRUIT
IFRUIT Iell I I II i 1 I le!l Io I I( /
I FLOWER|
FLOWER l ? 1 _ _ _- ?/0 J.Sl...'...
FOWEIIl?,
IFLOWERJOIJ,I -?" J} l O
mole _ _^

FIG. 43. Distribution of Licania guianensis, L. guatemalensis, and L. harlingii.

Distribution Maps 151

700 50
0
- ^ o?L. heteromorpha
-v ? N ^ var.yheteromorpha

.,-'Y '. I ^
.... 0'-^S?,<4^^B^ l. '^.

.....

FRUIT,;,,** ******* ^J0. . \ \

FLOWER .. . ..

FIG. 44. Distribution of Licania heteromorpha var. heteromorpha.

T
FRUITA - 1 1 1 O y I
152 Flora Neotropica

80 70 60 50

,
o;-N =_ heteromorpha
U^-"L. O
:'^ '
y//oI var. glabra

60 50 0

Seasona Il flooded forest . Sao ly \\

A forest ( \ \

10 FIMIAIMJ JJALS OINID ^\ ---

^ --aij^
II
1|FRUIT
I I I
-1 F7 p'--
I (0
IFLOWERIaS I \ I
_60_8 50

G var. perplexans var. subcordata 0

HABITAT AI
Terraf foirm o e forest Sd

*
60FRUTT *"; . ..... 1RUT 4 k
FLOWER u OW E....

FIG. 45. Distribution of Licania heteromorpha vars. glabra, perplexans, and subcordata.
Distribution Maps 153

70 60 650

L. hebantha L. hirsuta

HA ITAT
I ^/y-Terra Iirme
0 forest /

Amazonian caatinga Seasonally flooded forest /i

FLOWERI I e I I I I EI I E 1 - I

_ 70 60: 70 60

FL. L. hispida ,L. hitchcockii

?1 0
0 0 0

Montane & Cloud forest Slope forest

FRUIT 1 IS I IFRUIT 1* A

760 7- g..70 60

FIG. 46. Distribution of Licania hebantha, L. hirsuta, L. hispida, and L. hitchcockii.

154 Flora Neotropica

70 60 50 40

****-***
...../
^ ^ ^ ^FMAM ^ ASOND !!ss^^
j^/...

< *i (\^ L. humilis

Terra firme forest I ) t 1 '-

'FLOWERI _ 1_ 111
g _ _111,_ I ^a I __/ _
2CiS~~~~~~~~~~~~
70 60 50 40

ITAT i h

FIG. 47. Distrbution ofLicania hoehneiand L. humilis.

 90 80 70 60

0
00 0 .

_jS^--- T^ t^l00 *

HABITAT~~~~~~~~~
var.. hypoleuca (^(/
/{ ^ ^10
margins
~JSavanna var foveolatata
HABITAToveola

1FRUIT IOI*IIOI@I@I@I*I'
FIG. 48. Distributionof Licaniahypoleu.a.

HABITGAT var. hypoleuca o h

Savanna I
Savanna margins FIABITAT, v ar. foveolai
Terra forest
firme forest
Terra firme ____ Terra firme
Terra forest
firme forest ____ V^l / /
JF AMJJA1AIs NID \ I FRUIT F MI(J J A
FRUIT
JIFIMAMIJIJ
101 *****
AS
10 FRUIT IJIFI--I
A AS IONI
..,\I.IAIsMNIDI
A0
FLIII-1010 ^gI 0
IFLOWERgS0 ggg*0Z~LLI
Jg* \ FOWER
_FLOWER:^: ""J':^-~\ "
50
9080 ~~~80 70
70 60
60

FIG. 48. Distribution of Licania hypoleuca.

156 Flora Neotropica

60 50

0
0 L. impressa 0 0 L. incana

Iv ,'

0 40

FRUIT
II~I I I I I^ itI0I In R p e
FLOWER
IG. i 1D:: Ln....I I FLOWERi and

50 40 __ 60

4 D L. indurata( a L. inpae
..A. ........
.

^
~~oy~rr~
l~ ~ y ~-~. Gallery forest
Montane & Cloud forest
: I
-iJIFIMIAIMIJ
-IJIAIS
- H-11- - FRUIT FLIOR
NID
-
RI T OWER <i
50*~ 40 60

FIG. 49. Distributionof Licania impressa,L. incana, L. idurata, and L. inpae.

Distribution Maps 157

i L. intrapetiolaris o ^
O
L. irwinii

\<~~~~~~~~~~~~~~~~~~~~~~~?

HABITAT > J A y r / r
^.\
'
1uTerra firme forest ____ ( T 0 Soefrest ______
\ \
J FMALMJJASOIN D ( \ JF_IA IAJjAISiO5ND A)
FRUIT * * \ ;^ 1 FRUIT _*
R _i8"""_
FLOWER _"_ I I I I _ I LFLOWER __I
_ fI 1__

70 70 1J1 60

FIG.
5 Dsi0~~~~~~~
t iaL
n. fjei fen ns
it is
p ande ne z ii
L . jim

FiABITAT

forest
Montane& Clou Terra fire forest
lo(IN
JIFIMIAI14JIJIAIS I i I IFIJEI J IIJAISIOIN
D
FRUIT FRUIT
IrJIRU nI J I I I
FLOWER ?T II I~l 14 OWERI 1.EI I II
bu 1_ 0 bu

FI.5.Ditiuin fLcna rii,L.

nrptilrs .
jefensisanL.jm ez.
0 b
FI.5.Dsrbto f iai nrptoais .iwni .jfnss n .jmnzi
158 Flora Neotropica

7-O-?

L. joseramAosii r I L. kallunkia
_ I .
60 _ _

" 70
^ 700

FIA I Ftbi o iABliTATe L kru

0 10 10
D

51
FIG. 51. L. L. kiug
Distribution of Licania joseramosii, L. kallunkiae,
kallunkiae L. krukovii.
klugii, and L
Distribution Maps 159

70 60 50 40

C|: --- -----__ L. kunthiana

J7

F. . sbo fc a na 0

i \ ~~ L1
IY?\
`?-~

.i. ??,I~~~~~~~~~~~~~~~~
.......A'ON /
........ ........ f
,M..,J
?-~~~~~~~~~
~~~~~~~~~~~~~~~~~~~~~~~~~.....
o?/

FLAITA

FL 1 O00111
7060504
c.c? ~ ~ FG.5.Dsrbuino ianaknhaa

Seasonally flooded forest O

70 60 50 40

FIG. 52. Distribution of Licania kunthiana.

160 Flora Neotropica

50 40_ 70

0 L. lamentanda L. lanceolata
0

805 70 0- 0 0

HABITAT \ \ >-y (
.
FG 53 Dsriu io of0 acoaa...
icai ........
. .. a etna,L

trme forest
Terrda . 4~~ <, / } )
sand
Ji Jorest AlSI1 _N( ' JIFIMIAMIJ J A N D

80 _5706050

HG. 53. Distributionof Licania lamentanda,L. lanceolata,and L. lata.

Distribution Maps 161

70 60 50

L. latifolia

op-AI

Terra firme forest ....... . 7. .

J\
J_FIMIAMjJSO N DD J A / 10
FRUIT I1
FLOWER,"
_;g I 1 Wv
1 1 1W |F V m II
60 5O
60 50 60 50

0 o

--lo --10 ^^
/3^^-

Slope forest HABITAT ,

Gallery forest___________ / Terra firme forest.
54. FasAIMIJ,adL
IJFIG. D stbuin o Lcni a . J J AiSOINpDl
FRUIT ji ~iI I I I I OI FRUIT 111 III I ! ! !!/!
FLOWERI__. J. J J
___ , FLOWER _ J
'
'-6-0.............. 50 ' 60 50

FIG. 54. Distribution of Licania latifolia, L. lasseri, and L. latistipula.

162 Flora Neotropica

80 70 60 50

.Tea firme forest :i . / i' /

J
F
FRUIT
M..........

80 70 60 50

FIG.L laoL. I leucosepala

iI I
FLOWER II- L**'Jol .,jyl I f

Terra firme forest --- ~ ~. :

..80 . ...70 60 50

FIG. 55. Distribution of Licania laxifora and L. leucosepala.

Distribution Maps 163

70 60 5 0 40

tTAT
HABrI
0aN1
1FLOER10 { )\ 7
Gallery forest

70 60 50 _____)40 _
.0r70 60 50

Seasonally flooded forest i1 I \ .

0

FIG. 56. Distributionof Licania leptostachyaand L. icanifor a

Gallety forest

Seasonally flooded forest / p/

FRUIT

80 70 60 50

FIG. 56. Distribution of Licania leptostachya and L. licaniiflora.

164 Flora Neotropica

50 40

| L. littoralis L. longipedicellata
4

010. / .6.0 L 5 I.

HABIT AT : o '-,J
Restinga HABITA

Sesn---al flo de I
foSeasonally flooded forest
JJJ IASOINID
JIF1MAIM IJ IAISION D
FLOWER _ I I
1!l I WER
iJ -
50 40 70

"Y \))6 ?^ ^ ^ r'~l L.Jongipetala

FRUIT II!! !el F I |

Seasonally flooded forest

80 70 60 50

FIG. 57. Distribution of Licania littoralis, L. longipedicellata, and L. longipetala.

Distribution Maps 165

80 70 60 50

L'. L. longistyla
10~~,,,? ~0 ,k ^^-~

Gallery forest /* ^
Seasonally flooded forest
\\

FRUI 1 M I-I I I I I

a u 70 60 50
80 70 60 50

FIG.
/... ofLicania longistyla and
58.Distribution
"'. O o L.macrophylla 0

Seasonally flooded forest

"
uu.. 73 60 50

FIG. 58. Distribution of Licania longistyla and L. macrophylla.

166 Flora Neotropica

60 70 560 50

?L.macrocarpa 0- L maguirei
^

...................1
1 o -~- ;,.-. v-o0
<,,.,

80 70 60 50
JIFIM60
IM50
I80 NJ7

ZZ2I

0.

F JUFIAT MTJRUAS1ONTD J FIMAMJIJJIAISOND

-I-

Seasonally flooded forest ( j ) Beaches /,

"
FRUIT _ * _I f******I*-.
) FRUIT -) _
10 FLOWER _a__ _ J/ / / FLOWER I I I I 2
60 50 80 70

FIG. 59.
FIG. Distributionof
59. Distribution Licania macrocarpa,
of Licania L. maguirei,
macrocarpa,L. maguirei,L. majuscula,
L. majuscula,
L.and maritima.
and L. maritima.
Distribution Maps 167

50 40 60 50

o L. maranhensis -JL. marleneae

0

...
..........It I I i' 7~7"t~-' ..

0 0

Terra firme forest

JIA IS /
20 HABITAT TR\fJFI TAFIMIAMIJ( |
Gallery forest
50 40 _ ~60 50
60 5 110

FIG.6 D L. maxiamaar\ L.m- L aamexicana.

FIG. 6~~~~0

HABITAT
Terra firme forest H10 ABITAT
IIIJIFIM J ~Z ZZ lJ ISIO NID
MIAIMIJIJIA
iFRUIT i
"IAS^OND
FRUIT J 1
JFLOWERi.! ,I!!I_I I IIFLOWER
i/
60 _ 50 110 100

FIG. 60. Distribution of Licania maranhensis, L. marleneae, L. maxima, and L. mexicana.

168 Flora Neotropica

70 60 50

i fo oI
L. membranacea

Terra firme forest \ )

10

FLOWER ___ _ *_ I \ **
I A /I /* * : ...........10 I 1
60 50

90_______8 _____s60 50

r - L. michauxii L. microphyllal10

3;0 3 00I
oABITA HABITI

FRUIT T-II I I 1 FRUIT / /

.WE 066 g IfFLOWAERg
Ig I :
YU 80 60 bU

FIG. 61. Distributionof Licania membranacea,L. michauxii,and L. microphylla.

Distribution Maps 169

60 60

0. L. laevigata LL. nelsonii

0 "" :
1

HABITAT HABITA
Terra firme forest /\ Swamps j \
NDA-|
I/ M -
IFRUIT \ FRUIT I I I
I |I I
FLOWER I: \ I L
FOWER'
I\ I I I

60 50 70
G. . Ditibton o c l a . o L c n .
0 L. occultanso L. tambopatensis
?
--------- '

0 FRUIT
1 :::
I::I:

60 50 70

FIG. 62. Distribution of Licania laevigata, L. nelsonii, L. occultans, and L. tambopatensis.

170 Flora Neotropica

70 508 O

L. micrantha

0 !
0

Terra firme forest

99 9 ^
FRUIT i? -^ ?/ /'^ /"**..-...-.o... :' j
FLowERgs I j4N-10 '0
70 60 50 40
I 90 r _ LI

^
o 0 -)A
HABITAT0~~ ^ -

FLOWER
0 80 70 60

FIG. 63. Distribution of Licania micrantha and L. minuscula.

Distribution Maps 171

70 6050 40
-
~'-ye O-L.
" ei::,
.... minutiflora

.. .... .

10

"JIFIMAMJIJ'IISIOI'I 1
TERr
I r f t1 1. 1 t.---E-l
- - I
_ 60 50 40

-.....
yj~ ....1/l'"
':....L ' miltonii

FIG 64 Dstrbuionof icniamiutilor ad L mltoii

0 _
0 Terra firme forest
F I G. 64. Distribution
of-Licania
minutifora and L. miltonii.
172 Flora Neotropica

70__ 70o_

L. montana , f o) oxwL. mollis

FHABITAT
HABITAT Beaches
Montane & Cloud forest / Seasonally flooded forest .

1 FA J IAIS
JRFIMIAIMIJIJ
I I oAISINIDI1
II I Il f 10
FLOWERUI i g I I
FLOWER FLOWER
"I
-
70 D
U W

80 _ 50 40

L. morii y L. naviculistipula

FIG.
65. Distribution
ofLicania
montana,
L.mollis, andL~~~~~~~~~~~~~~~.........
L.morEi, ....n u l
HABITAT BITATI
Terra firme forest, Terra firme forest
J. FMA
F JJ ,A S ON,
MJJA,M J IFIMIAI J IJ A- IS FND
FRUIT FRUIT11- '
FLOWER FLOWER I I I
?
70 o 4U

FIG. 65. Distribution of Licania montana, L. mollis, L. morii, and L. naviculistipula.

Distribution Maps 173

70 50 40

L._niloi 10 L. nitida
_' 0

HABITAT HABITAT
fr
TCerrafIrme Or
forest rrado
IJ FMAMIJIJASI IAISIOINIDI
JIFIMIAIMIJI-J
FL I OWERI +...gg.. L______
10 70 40
80 70 60 50

?/,){j^l
?~"-'v-~} L. oblongifolia
o--~ ., . ,,.: e ,...........
.......

Terra firme forest

J FMAIMIJJIASOI
NIDU ,, ',:, . ...o ......./'
I
FRUIT iee illlle ;.0-" o)-.... .........
('

80 70 60 50

FIG. 66. Distribution of Licania nilo, L. nitida, and L. oblongifolia.

174 Flora Neotropica

70 60 50 40

,C L. octandra subsp. octandra

, I.

.. ..

/'
.._.J~-':MAMJ D ::< / . ........
70 60 50 40

..........
HAB T ... ...... ?
Gallery-|@/,O
IGallery forest
f, . /1O fI "
Distribution Maps 175

80 70 60 50

r L . o ctandra subsp. pallidal

D,.I > \</ ~~~~~ / I

Terra firme forest

Jz - 1AMwIII
F J AS ND
NS S J AMJ / D

80 70 80

F .isr L.
o octandra L.
L operculipetala

petaI
la

~
FRUIT I FRUIT /
FLOWER I__1 11 / FLOWER_"_ * I I _
80 70 70
")

FIG. 68. Distribution of Licania octandra subsp. pallida, L. octandra subsp. grandifolia, and L. operculi-
petala.
176 Flora Neotropica

70 60 _ 60 50

670 60 50

)Jo^ %L. pallida

o ,.^~~~
. .

0~~

Terra firme forest

lFI AMJ IJIAISI D \i\ a /
FRUIT M \ i.) ~-/( \ V < ;7I
FLOWER F) wi --? .
_-770 60 50

FIG. 69. Distribution of Licania orbicularis, L.

L. ovalifolia, and L.
L. pal/ida.
pallida.
Distribution Maps 177

70 606 _ 0

, L. pakaraimensis L. paraensis
12C

?TFRUIT

HABI'TAT ,/<' . '.

HABITAT:":(
/J^-r ^ .......

FLOER
IiI ! il , FLOWRO I I
.00

80
bu 7070 6060 5050

[^esl^~r >4 ^,._- 7

1 ally flood d f r s _ * - .^^^\k)/ X . . 1
^ ^ ^^ _
:3!MAMliASOND^-^-'.^i/ Jff/\77z^^:-7t^^^~~~,?
?ro l *** ( A /
[ijgsissftg-^ / ^iO^w
' ^ -^
/A/ 'M /^
8U 73 ^'
M
S FIG70floodeditiuino
onalne forest iaapkriess .preni,adL avfoa

\FRUIT/ of |L1n a L aap/ /

FIG. 70. L. andparviflora.

Distribution of Licania pakaraimensis, L. paraensis,
178 Flora Neotropica

70 60 50

x, L. parvifolia
o
~

'r1
Iz 2" Fv A J Av [ N r.' '""'.... ) / . .

' L
I~~~~
~~FRUIT
J|F|M|A*MJ|J|A|SI y........ }\ ~...
X

Seasonally flooded forest

FRUIT

0 ~~~ - 0 1 '5
Gallery
7O, forest
70 60~ ~ ~ 600 500 60

,IG7 L.
parvifructa a persaudi
L.

FIG. 71. Distributionof Licania parvfolia, L. parvifructa,and L. persaudi.

e1 /
HABITAT~ / /~
A
~
firme
r:erra forest
i.Terra ___ > firme forest _

FIG.__ 6 60
7iboiaiapf,L________6
Distribution Maps 179

100 90 80

_ - ----- .. ....L. platypus

..

A0ITT

Gallery forest
IFIMJA|MIJIJIASlONID I (

_8_?__10 0 9 0 70
70

__80_ _____70 60 50

FIG. 72. Distribution of Licania platypusand L. polita

Seasonally flooded forest 0/ / ? >\ 1 j *\ 7 ] / / 1

JFMA JJASO ND // _ Y /
t00
. ) (
180 Flora Neotropica

50 50

Terra firme
foreso t:; ___ *'"' " ^
........5"y " . flooded
S...easonally. : forest
iMM
FRUI____T_I!i ASN N
_FMAIMJlAIS I _ 50 lOiND
FRUIT ) I FRUIT / i
_
FLOWER i _lll I _! / /i _
IFLOWER
..5050
70 60 100 90

L. pyrifolia ..--.-. L. retifolia

0
7_Bo~~
0
- ..---, .'
,ro^

HABITAT 'ABITAT
Gallery forest Gallery forest
Terra firme forest Terra firme forest
J IFA IMJ JASOND
jNID J AIS NID
FRUIT I i FRUIT
FLOWER FLOWE

'_o
m7 60-uo u90

FIG. 73. Distribution of Licania piresii, L. pruinosa, L. pyrifolia, and L. retifolia.

Distribution Maps 181

70 60 50 40
^
^ .or.r ?..L. rigida2o
"? 0

o 50 4

Arid formations, caatinQa ai ~'" "

.... /

FRUIT

I / ' ~ o L. rigida (cultivated)

0 80 70 60---

FIG. 74. Distribution of Licania rigida.

182 Flora Neotropica

80 70 60

0
.0*^ ^^ o Y L. reticulata

Terra firme forest - --

Seasonally flooded forest - - 10

J A S D I
;FRUIT * 7 ) \I

8^
,,__70
? -60 50
50 40 50

L. riedelii L. robusta

10
0

'HAAHATT 0 10

^
""**....^ ~"~\~"~~'~---^Terra firme forest

FIG. 75. Distribution of Licania reticulata, L. riedelii, and L. robusta.

Distribution Maps 183
I
60 50 60 50

.: L. rodriguesii L. roraimensis

10'~~~~~~~~~~~~~~\ I~?c-1

HA^^BITAT1 V A ( F
ILHABITAT 10
Terra firm st ontane C d forest
Cloud f
J F1AMJJASIOND )/.ONDMJ F JJ S ND
IFRUIT i
/e / I FRUIT
_../ _
FLOWER I *le*
* 1"i : II I * . .J IFLOWER ! J
'"
60 560 b
^_ _ _ _ _ _ _ __60
_ 80 70

FIG -. o
. L. rufescens L. salicifolia

Slope forest ' Gallery forest /

) \ /, / '
JFMA'ND -"-- /[\ f ' --- ! IAIS
IJIFMIAIMIJIJ NID./
10 FRUIT_ -------^
I1- I i I-/,;)
-1FRUIT I ~ -
IFLOWERI
PLOWgR~":::^^ij|_j~~7r----lo II I i i II
bo o L 7o80

FIG. 76. Distribution of Licania rodriguesii,L. roraimensis, L. rufescens, and L. salicifolia.

184 Flora Neotropica

50 40 __60

L. salzmannii L. sandwithii

FIG. 77. Distribution

of Licania sazmannii, L. sandwithii,L. santsii, and L. savannarum.

Restinga Terra firme forest

FRUIT IFRUIT II !I Ii L
FLOWERII i I i?iI IW .. I I .... I
50 40 60
Rest5nga 4020 Terra s|
50 40 7mr0
70 60 A \

L. santosii L.-?santos..
7.DsrbtooLiaiLsadih-
FIGL . L. . savannarum
L. savannarum
and

b) 4070 (0

FIG.77. Distribution
ofLicania L.sandwithii,
salzmannii, L.santosii,
andL.savannarum.
Distribution Maps 185

Cerrado : ' ,'~ ~l:~)~, L."sclerophylla

f.,/~ ,t

forest
70'Gallery 50
/

Terra firme forest ./ . \ I // //// /

Savanna margins 1\) , _//j ) . /
Seasonally flooded forest . \) * / / ;: / /
/I J VJjA:
JFiMAJJJASOND N ; / /', I
'' :
FLOWER F --.(
.
|
________170 _____ 60 40-- 50
70 60 50
?
.T)I~~L.
\^ / silvae

70 60 b

FIG. 78. Distribution of Licania sclerophylla and L. silvae.

186 Flora Neotropica

5Q 40 . ... _90_ _ 80

L. silvatica ""
I L. sparsipilis
10,; , 0; V6610<21

......9 ~
•.....:'............. .......

HAPBITAT '
/ -Gallery Galle ryforest
forest
Cerrado -Terra firme forest,
Terraed o Terra firme forest

FRUITE I I I I I I FRUIT frsI*___ n

50
5O 4 40 4O 60
60

,IG ....DitL. sp atita, L.s L.. L. sprucei

FIG.
79. Distribution
of Licania L.sparsipilis,
silvatica, L.spicata,
andL.sprucei.

FG79Dsbtno/slacLsaspl, a/dL.sei
Distribution Maps 187

60 70 60

L. stewardii.. L. steyermarkii
0 0

' % . .... 1.o ... ...

I
AIjFM
wi MA/ 1 101NDJ
IStjrND
]MIJIJAIAISO \ -I iII iJ I
FLOWER S ND ..
FRUIT_ _!1
iFLOWER
I *(I ITI I I _ *

60 60

, L.stricta L. subarachnophylla

....... HABITA

60bU

10 HABIT Gallery forest

Seasonally flooded
floodd forest Terra firme forest
IJIFIMIAMJIJIAISONID Ja IFIMAMJIJIAISO
FRUIT FRUIT
I
I 1 1 l 1H
FLWR ":
IFLOWER

FIG. 80. Distribution of Licania stewardii, L. steyermarkii, L. stricta, and L. subarachnophylla.

188 Flora Neotropica

70 60 __70

70 7

-,... .....

^ HABITAT
HABITAT^ \ \^ J) ( '^^
Montane & Cloud forest N / Slope forest|M|AIM|J I
rS
- ~ -
- Ig-- 1-----l-1
--FLOWER I FLOWERI .
FRUIT IIIII I I I I I I I 1 1FRUIT
\ II--
I : II II I I \/
:FL:OWER I:1 1 ,I I ,FLOWER _ I I I,

70_0 0
_/v^ L. teixeirae L. tepuiensis
o o

Terra Cfirmest Soe forest \

ofLicania
0 81. Distribution
FIG. subrotundata,
L.tachirensis,
teixeirae, L. L.
and
tepuiensis

Terra firme forest ?, Montane-& Cloud forest

JIFMIAIMIJ J F MAMJJAS nD
III ! AM AISIOND . f/
[JF IJIFIMIAIMIJJAIJSI^ISOIFNID
S!qlNID ! I
FRUIT I III I
FLOWER
WER I I I MMII

FIG. 81. Distribution of Licania subrotundata, L. tachirensis, L. teixeirae, and L. tepuiensis.

Distribution Maps 189

60 so

L. ternatensis L. tocantina

1 00

Slope
I?,
forest
JIFIMIAM J J ~
AIS ONID
\ ^^'Terra
5J
firme forest-
FlMAWMAJSJNA1r
2 1
0

FRUIT i~*~ _ /t
S_FRUIT
FLOWER _I :ggg_ ff:I FLOWE _ _ _ _----H/

50 40 I 80 70

I
,,
tomentosa, L. L trigonioides
.a

BI
HAFITAT / \ \ \
Restinga

FRUIT- 10 HABITAT
LOWER! I IIi$11Ig$1 . I Terra firme forest /
JIFIMIAIMJIJIAISlOtNID
FRUIT
L igid
FIG8.i s otiu Linatnts,LtcnnFLOWER
4U
4U80 70I 11FLOWERII I I 1N1 1 1 1 1:1 1 1</70 I I

FIG. 82. Distribution of Licania ternatensis, L. tocantina, L. tomentosa, and L. trigonioides.

190 Flora Neotropica

80 70 60 50

* L. triandra
'^ .... . ... .

..,. ~~ ~~~~~~ .. ....

TerraBITAT~~~~~~~~~~~~~~~~~~~~~
firme forest _ \ \ I i \ ~ A
J FMAltJ J AISIOND - ^ >
I ) rl \ \ (

FLOWER_III_* ! , ^ /^ S^\ V / L / //I I

?U 70 60 50
80 70 ____60
___ _______50
~~I~~~~~~~~~~~~~~~~~
"~~~~'~/~" " "ooj
0!?-.:~- L.unguiculata
FR i-g
FLW o _ (.; /^ v In \? /y//

HABITAT"'""T
~
. .
.. ... y
FITerra firme foresta .
(
Seasonally flooded forest "\ r ^ > /
JFIMAIMJ J A SND^- )^ / V M t (

8U 60 50_

FIG. 83. Distribution of Licania triandra and L. unguiculata.

Distribution Maps 191

/L. turbinata] L. urceolaris

1 O

,. 1".
u<^^^^^t~~~~~~~~~~~~~~~`O

Restinga 10HABITAT/1
Terra firme forest Terra firme forest
J|FIM|AIMIJIJIAISIOINID MJ IJIAIS CINID
IJIF|MIA(
FRUIT I 1 I I I I I I I _ _ FRUIT I *I I 1
FLOWER:_ II I I I I a
40 70
70 60 50 40

L. vaupesiana

Seasonally flooded forest

FRUIT I III I I I I
I111 I
^\( / I/ yr 0u
FLOWER; F I IIII v, / /yf 1 . _
70 6 0 4

FIG. 84. Distributionof Licania turbinata,L. urceolaris,and L. vaupesiana.

192 Flora Neotropica

80 70 80 70

L. velata L. veneralensis
10..-/ /'
~",~c~~,o11~....-10 ol10

? o ::
~ ? ~, ~
Io ____ J o . -- '-.J.=,--- 0, "-I
...%~. ~

-
HABITA^ - ^r^^V /^ ^ HABITAT-
Terra firme forest '- Terra firme forest -- I
l J
JIFIMIAIMIJ O .'I 'JFI I I
F!1.FRUITII
I !FLOWER I I I I '~I I I FRUIT
Ill i I!1lel 1 :LFLOWER
a I ...... I I I I PI!!"..

_1 U 80 70 60
0', 0
10~1
- 0
L. wurdackii

10 0

'0 o"'I -
0-
i--L---4T.--0

o o

FL {,OA (X1
JIFMAIMIJIJASND
A\ .H , II
L ggE 11 11 1II!I I . ) t. I I f . (\ /I // )f v1
8U ~ ~~70 60

FIG. 85. Distributionof Licania velata,L. veneralensis,and L. wurdackii.

Distribution Maps 193

8 0 60 50

CParinari Composite Map

0O

.0 0

0
F W-EROO."*@ @-@'@ f 0

u.. I 8 un...
D .fh

................. e */-^^ 7^^ r*^>^ . ..

^^Fs
h~~~~~~~~~~~~~~~~~~~~~~~..1
I\~
IV \ ^^

" ;?~~~

30 FRUIT ****0 10 ** ** , /
rY- /
194 Flora Neotropica

50 40 50 40

P. alvimii (7|.) P. brasiliensis

l J~Fs sJA
JS~.. i .......

e
Terra firm forest
_ _ __
FRUIT __--- 201 HABITATI
FLOWER
_r _ _ _ _ _;____ I I Terra firme forest j
50 40 4
70 60 50

8 Dr onofPrri al P. campestris

Galleryforest'- 1-

10 FLOWER /JTTJ7\)-"-7'-----^^ .....-^-)

gigBigSgtigg^
~70 60~ 0

FIG. 87. Distribution of Parinari alvimii, P. brasiliensis, and P. campestris.

Distribution Maps 195

60 50 80 70

P. cardiophylla P. chocoensis
10 10

0;~~~i
o

"''HEITADFHA 7
M AISIOINIDI
;T
Terra firme forest
10*
y Terra firme forest 2__.
JI J IAISIAOINIQI \ I IJIFIMIA8J IJ|AISIO NI
FRUITII I I I II I I I IFRU IT
[FLOWER I:II I I ! ! III i
~...
60 50 8"0 70
80 8 b 70 50 40
70
FIG.
/.Dsrbtonof p. klugiio a P. i P. littoralis

\' 0 10

* HABITAT
V^TAT \0 ;\
HABITAT, Restinga
Terra firme s firme Terra forest
.t 2
^ AFMI - J-J AISFOND IS 1 JJ'FM A J J A S OINID
FRUIT i v\o ' \ FRUIT
ov
FLOWER FLOWER
Eir J
, 70 50 40

FIG. 88. Distribution of Parinari cardiophylla, P. chocoensis, P. klugii, and P. littoralis.

196 Flora Neotropica

70 605

1~0
o
o P. excelsa
10
10

HABITAT ....... . _ _
0 ___1_

b u i o
FG89Distri of Pa rn a e xcelsa..??
r -

3C Terra firme forest

JAFMAMJJASOND \

______ 70 60 50 40

FIG. 89. Distribution of Parinari excelsa.

Distribution Maps 197

P. maguirei P. obtusifolia
............................

.......

HABITAT HABITAT

JIFIMIA JJJIA SCN D IXA JIFIMIA_ ,71J,A S C N DI /

FRUIT _-- -- I I I - I FRUIT * _"i * g _ /
FLOWER= I II FLOWERgg ogiggg1gS /
60 40
7 ,o0 50

90 Dsrbto P iP.
FIG.'^ montana

! / ~ c . .....
~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~c~~

. .-' /
oJ F A J'" N D ...........-'t
FLOWER" : . ... . .. . . . ... .

Terra firme forest I

FRUIT 9 " 0
* * ( ,71 f \ 7 A /

70 60 50

FIG. 90. Distribution of

Distribution of Parinari maguirei, P.
Parinari P. obtusifolia, and P. montana.
and P. montana
198 Flora Neotropica

0P'~~~60

Semideciduous forest ___ . Seasonally flooded forest J< <

JF MAMJJASOIND/ 50 .- 7F0EAMJJAs
. oND O)/
IFRU
TA_re____/'_ FRUI
HA_ _i~_ _ _~//_ \11
70 60 _

OWI. R ^P p/ ,~ l rv |If^li 0
P. par la P. P occidentalis

^^ ~
~~HABITAT-- ~7Z-:--74-JL_': /{ -^ HABITATY' ^ ^ '^^^ J ~^\

FIG.
91. Distribution of Parinari pachyphya,
P. P. pariis,
parvifoia, P. and
occiden talis
Terra firme forest Tera a e for

9
FRUIT i o P1
o a ay
---c FRUIT A a P id ai
Distribution Maps 199

P. rodolphii

- ' (
FRU / / )/

Terra firme forest

80 70 70 60

FIG 92 P. romeroi,andP. sprucei.ucei

Distributionof Parinarirodolphii,

HABITAT HABITT
Terra firme forest Seasonally flooded forest
J,FMAM,1 JI1AISOD JF A J AS D
FRUIT
FLOWER
'
8U 70 70 6'

FIG. 92. Distribution of Parinari rodolphii, P. romeroi, and P. sprucei.

200 Flora Neotropica

70 650 40'
0.
Exellodendron Composite Map

Distbution of the genus

FIG 93 Exellodendron.-.

; J/ ]AND/
JFIM:iA i

_ 70 0 60460 50

FIG. 93. Distribution of the genus Exellodendron.

Distribution Maps 201

70 60 .0

"Y~
...:: _... E. barbatum

1E I
60 50 40

770 5 40

'
'7 0 60 5 0 40

FIG. 94.
FIG. of Exellodendronbarbatumand
Distributionof
94. Distribution and E. cordatum.
cordatum.
202 Flora Neotropica

70 60 50

.ogr E. grE. coriaceum

Gallery forest... . .' -

.. .
' ......
Savanna mar ins
Seasonally flooded forest

FRUIT _i FRUIT -

60 50
50 40 50 40

E. gardneri E. gracile

FLOWER_I __ __
- -_ LOWER I I i I I I

HABITAT^ ? HABITAT 2
) ^--j~l
Cerrado_______________ Gallery forest

JFIMAMJ IJA SIOINIDI cr JFM AMIJ J A S gOiND

FRUIT gm I 11 I I [- RUI
IT
--IFRU ? * I I -
FLOWER I8I FLOWER II:II
40 4'

FIG. 95. Distribution of Exellodendron coriaceum, E. gardneri, and E. gracile.

 ___ 100_____ 90 80 ___ 70 __ _ __ 60

aCouepia
\ -onianca^itin
/

r o~~~~~~~~~~~~~~~~~~~eF
O0 I0 v -?E~~~~~~~2 1

0~~~~~~~~~~~~~~~~~~~~~~

Terr fimsfres

Montane & Cloud forest~~~~~~~~~~~~~~~~~~~~~~I ?10,O~~~~~~

i

Whitesand fo /rest or campina

Slop ire forest

|Seondary&Clu forest \ - - - --V iX
___ ______ - 3 .s^9.
X / T X7. "
ap |\^
Seaonally f loodd forest ?

^
forest \ ^^- ^
|Becodr ____ \ . 'f a ; I/ W

^'
SeaIsonally tlooded forest j-
,/<^.
4 l &4.
y j?|
forIG
Secondar 9. isriutonofth gnu Cueia

_ ~~100 90 80 o
7, ,^6 ~5

FIG. 96. Distributionof the genus Couepia.

204 Flora Neotropica

60 504n

10o --' C. amaralae 1 C. belemii

10HABITAT
"" Ar (\ \
A i ) f I/ TATA
Z"^?~HAI j
J| Jii J S-

Savanna margins ) OI FRUIT _I ___

I
Amazonian caatinga__ __
60 4F5'

70 50 40
10 10 .
C. bernardii C. bondarii

' '7 0 '

'~~~~~~~~~~
FIG. 9 .Dsrbto
fCupaaaaa,C
eei,C
enri,adC
odri

T1erra firme forest .. .........

IG. 97 J IIs o C m C b . / . .
FRUIT .':" " '""""'4
I
FLOWER. I III II 'M I
Distribution Maps 205

80 70 60 50

^.~3 -
.: C. bracteosa
,,t_J___f^.f'y

[FLOWER X
.,,
*.jo -
iA)l _ -. I

Secondary forest
Terra firme forest

80 770 60 50
60 70 60
"
C. canescens o? C. canomensis
^
p' "o o 0 /IiS ^ ./C ']

'
. Slope orest / f
Terra firme forest .\

HABITA r i o o C b , c e adCcnoes.

60 70 60

FIG. 98. Distributionof Couepiabracteosa,C. canescens,and C. canomensis.

206 Flora Neotropica

50 40 60 50

1 / C. carautae C. caryophylloides

HABIT HABITAT--7
Terra firme forest Terra frme forest 0
(
FRUIT JIF
IMIAMI
J IJ N I I J JIA S OIND
-FRUITI II FRUITI Im 1 /A
MI I IIk
IFLOWER I I1Im1[1@1
1 I.L|| I I I I I i111
IFLOWERI I'
40 60 50

60 50 60

F9. is C. cataractae andC.

cidiana

Beaches
Rheophyte H/A "0 A
Seasonally flooded forest Terra firme forest
J F M AMJAJIJoINAI /I yJ FIMIAIMJIJIAISIOMNIDT
FRUIT Li /I ....' FRUIT
FLOWER_ IM01g I 1_J FLOWER
660 50

FIG. 99. Distribution of Couepia carautae, C. caryophylloides, C. cataractae, and C. cidiana.

Distribution Maps 207

1
FRUBITAT';., r / C. chrysocalyx

'
0 0" ' ' ' '- 0
-'--.z
FLoWERI" 1E1zzE1 .....
J ':..
? -e.v

Terra ~ -
firme forest ? , . /~~ /,.
Secondary forest
IJ, F'A'J
I II AI . J - ' / )
FRUIT l ( 3 o- . :
FLOWER '""
iil l ,- l '".'''** ''''_ '1
80 _ 70 W0 50
540 4060 50

FI.. C.
Ducoarctatoua c , C. C. comosa

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

.^ ^ B T T
..0~~ \ - - ^ *K .....
..':'
HA )
_

HABITATi Beaches
7^?P^~^--^
Restinga heophyte
J
JIF^M2 JA S I--- IF AIMI J IJA SO ND
FRUIT I
AFR|)|JJST--
---
40 60 50

FIG. 100. Distribution of Couepia chrysocalyx, C. coarctata, and C. comosa.

208 Flora Neotropica

60 60
loi, C. cognata C. cognata
-"-... ~-var. cognata10 7' (, var. major
?"-

/: . '.i> M.

Savanna margins ( 10 HABITAT

Secondary forest/ \ Terra firme forest 0
J|F|M|A|M|JJ|A|S|O|NP \ JJASO D
JFMAMJ D\

60 80 70

0 C. cognata / C. dolichopoda
var. membranacea

(
Savanna ___ \) ) / \ Terra firme forest , o

JIFIM|AiJ|JA|S|O|ND /iD ) IJIFIMAIMIJIJIAISIONID

0'FRUIT
-I 1 I '
i.........F I I \I I
|IFLOWER_IIII
l I I I FLOWER
O6080 70

FIG. 101. Distribution of Couepia cognata and C. dolichopoda.

Distribution Maps 209

60 60

C. eriantha Acioa edulis

XHABIToAT.......0 HABITAT
Terra firme 0
Seasonally flooded forest forest
JFIMAMJIJLAISoD \ JFMMJ J A S 0 N ^1
FRUIT I I \ FRUIT * **
FLOWER gg___ I // //1 FLOWER _g__ ___ 2 \)/J
Oi_0
70 60 60_50

10^ ^0 ........

HABITAT, HABITAT 0
Terra firme forest Terra firme forest
JFIMA^ JIJIAISONID
ASN0JFMA'J|| 1 IAISIONID / (
FRUIT 1 1hdIII
1 1 I T
FLOWERI 1 1 1 1 1E\\) I |IFLOWER
70
60___ 60 50

FIG. 102. Distributionof Couepiaeriantha,Acioa edulis, C. elata, and C. excelsa.

210 Flora Neotropica

60 60

00 0 . exflexa C. foveolata

0 1o0o

HABITAT 3 ui e ABITAT .
D o i a C
Terrafirme forest TSeasonally flooded forest
\

FRUIT
IIIIIIcIIII_I
FLOWER
JIF D
MAMAIS.ND i C. FfeaAIMI
FLOWWERI(^
JJIAISd Cl

I 6N -^ . _
60

I0sioaa.oC. froesii C. glabra

FHABITAT/........... HABITAT
Terra firme forest Terra firme forest

FRUIT _ n [f FRUIT
FLOWER FLOW II I:I is/
60
^^^^^^^^^__^^___60
FIG. 103. Distribution of Couepia exflexa, C. foveolata, C. froesii, and C glabra.
Distribution Maps 211

70 60 50 40

..:erra...~... ..;.~_..
...fi...foe..:::70 ......f .^- ..... . ..

,?-. . .

HABIT I I I I

7 50t?4
?60

FRU ISO ...............

JIFMAM
IJIJI A IN
FIG.104. Distribution of Couepia grandiflora an

F W
ER ............ :
I

7060 60 5 0

FIG. 104. Distributionof Couepia and C

grandifora guianensis subsp. divaricata.
212 Flora Neotropica

70 60 50

I o. . -
--. C. guianensis subsp. glandulosa

Seasonally floded forest \

.
70 60 50 0
yo 10
o\3 .,,.,- -

FG1C. guianensis subsp. guianensis

0 0

FG105 DisrbtoofCupaginnisus.gadlsansus.uaess

70 60 5O

FIG. 105. Distributionof Couepiaguianensis subsp. glandulosaand

subsp. guianensis.
Distribution Maps 213

60 50 40

C. habrantha \ C. impressa :
f '
?subsp.0 cabraliae

01 '
I:::' . . . . ..I.C 1

HABITAT S V/HATT
0 Terra firme forest TerrHABITAT for-e-
Seasonally flooded forest / Restinga -
J FIMAIMJIJ AISIOINID JIF1\M AMJIJ IAIS ND
~ -
FLOWERgggJ_I _I IIi /j / TFLOWERUI
R

50 40 50 40

C. impressa | .'.i , C. insignis

subsp. impressa
/ :..............

HiABITAt, Restinga
Terra firme forest ITerra firme forest
2f-g 20
IE IJFIMIAIJ IJIAISIOINIDI IJFIMIAMJIJIAISIOINID
-- --
FRUIT I I I I I I IFRUIT I I
FLOWERI |I I I I I I IMll-E25 ---
4U 4U

FIG. 106. Distribution of Couepia habrantha, C. impressa, and C. insignis.

214 Flora Neotropica

~..
~""~'~
............ ' '
, I///'
~0'.. I-.~'~
~. ~ C . latifolia

HABLTAT
Seasonall flooded
C, HABITAT 10
forest R
Terra firme forest
FT-MTAJO.1 I J
JIFIMIAIM J J A SNIIND
FRUIT ,
*__****,* I ! FRUIT I
:::WER :__=:; _S_ _ E- -B
\YVFLOWER

60 50 40

IG C.
l.ongipendulao IColongipetiolat
C.

Terra firme T esringa

f
forestJ s
1 JFMAMJJ AI .....i.......... 2J MA J J AIS0 ND-
10FRUIT IongIpenIa 0/ 10 FRUIT I -I
FLOWER
60
F
IM
I: H-H
40

FIG. 107. Distribution of Couepia krukovii, C. latifolia, C. longipendula, and C. longipetiolata.

Distribution Maps 215

70 60 60 50
0 1
1^^^C. macrophylla C. magnolifolia 0
.......... ... .........

i 0
Terra firme forest..^ Terra firme forest )
20 . JFMA__ JASJOJ ND , _-2C 10 IJ FIfAIMIJIJIAISOND ,/ |
FRUIT I__FRUIT I 110 _ _

I.____________________60 6_ _ 60
0
d'
^C. marl.
maguirei C. 'eneae

0HABI 0I C magnoliifolia,C. magiei, and C maleneae.

Rheophyte ________ _ ) ) / Terra firme forest //

I JIFIA>JIJIAIO
" // I IJIFIMIAIJJASNDI f .1) \ /
1C IFRUIT 1 1 I L X ./. z IFRUIT I I IIII1 11111 // /A
FLOWER 10 I I I
FLOWER I II
- 60O
I60 ________060
FIG. 108. Distribution of Couepiamacrophylla, C. magnoliifolia, C. maguirei,and C. marleneae.
216 Flora Neotropica

60 50 50

C. martinii / LC. meridionalis

. Terra firme forest

JF|MAIMIJIJIAISE NID1 JFIAIM JJAS ND
XI
FRUIT ~ - FRUIT_ /
FLOWER FLOWER
60-50 50 40
60 80 70

F-IG.
1 C. multiflora C. nutans
0
0^ ^^^
........-------- --\~~~~~~~~~;
.0

'0 5 O 40^ -
6O\ 5^-^/''-

HABITAT
savanna
IFCABITAT.
Savanna margins Montane & Cloud forest
F
JG1 M0 D sibIAISoOIN uiD JIFa ji MAli J iSDloND
FLOWERJ
I_,1_60
1.J1? -WE-7R "i IE
800
FIG. 109. DistributionofCouepia martinii,C. meridionalis,C multiflora,and C. nutans.
Distribution Maps 217

80 70 60 50

^ .0 S.^ o,o&tA c. obovata

0 0

-
.0 0A-- )r

'
N ,,,/ \ /
....; ,,/ 1.

Terra firme forest

-
FRUIT
CR- g - *: vf A. /1 \ / / /
FLOWi ./1__/K
//' j .J \
______
80___ 70 60 50
8F 1tf701O i oCu60aObat ad50

10. DistributionofCouepiaobovata
FIG.1 npa rillo,l
1: L

2
Savanna__________-'-"I I

l
IFNIAIMIJIJIAISIOINID ...........

Terra firme forest .-/

FRUITA
Secondary forest ________ -- x-^ -j .. . ..,..

80 70 50
60

FIG. 110. Distribution of Couepia obovata and C. parillo.

218 Flora Neotropica

C. ovalifolias C. parvifolia

HABITATk P I IHABITAT

~~FLOWERI ~
g"g"~SMg" ......E| ,'
*FLOWERg"""" .__...........

T) C.pernambucensis
.. C. platycalyx

FIG 11 .. b fo ..... f C o . b s .

firme Terra forest Montane&

_ JIFMIAIMIJ
I IJIAISIONIDI -II NI
IJIFIMIAIMIJIAISIOI -/
FRUIT I -p-
I 1I 1111|1 FIRUIT
IFLOWER
FIG. I
11. Distribution of Couepia I I.1-.I C.
:I ovalifolia, FLOWERC. pernambucensis, and
II parvifolia, C.
plat70calyx.
40

FIG. 111. Distribution of Couepia ovalifolia, C. parvifolia, C. pernambucensis, and C. platycalyx.

Distribution Maps 219

_ 0 70 60

?k?^Vy C
.t^C. paraensi1
0 Bec .,/. <..._subsp.glaucescens
/.es~

Gallery forest ......

0 Seasonally flooded forest , - . _. A
./'---) . --~- ...
S0N\IIT
|ZZ lD iFMAM IA , W L (1 | \ / 1l

0
0
-:::--C. paraensis i_ C paraensis?
!'~"'-? ,subsp. paraensis subsp. cerradoanda

0 10

HA
BITA H

60 50 50 40

FIG 12.. Distribution of Couepia paraensis.

220 Flora Neotropica

100 90 80

300 ( C. polyandra
^]

HABITAT
Gallery forest _ 'o 51
Terra firme forest \ /
Ii I ND f
V FRUIT
IJFMAMJJAS
gi _0I , /

110 100 90 80
80 70 6030

C. racemosa

HABITAT \
10 Terra firme forest -

Seasonally flooded forest

Secondary forest '/ \ L
White sand forest or campina /
\ / ///

| tJ]FMMIAIMIJM|J|ASS )IQI
A )\ L1I \
L
FG80 1.Dsrb60 70 50

FIG. 113. Distribution of Couepia polyandra and C. racemosa.

Distribution Maps 221

80 70 5

C. recurva C. reflexa

'1 O0
0
HABIT^__AT RinHABITAT
Montane & Cloud forest Terra firme forest
JIFIM_AMJJ JASOND
JA / IJ
JFIMIAIMIJIJIAISO ND /
FRUIT ____I _i_ _ / IFRUIT II /
FLOWER _ * _/=_ IFLOWER __ I IlG!
Iu 70u
60 50 50 40

C. robusta,
FIG. 14. Distributionof Couepiarecurva,
C. refexa, and C. rufa.

^V^ V HABITAT
HABITAT Restinga
Terra firme forest ^ ~ Terra firme forest
__
JFIMIAIMIJIJ IAISIOINIDI '''t
JJ'' A - 1 -
JFMAMJJASJND -
FRUIT I I I I I /I IFRUIT I
4 : 1i"i:iii
LFLOWE^ llFLowER-g
I --: g
bUG14Dsiu 50 ru 40
FIG. 114. Distribution of Couepia recurva, C. reflexa, C. robusta, and C. rufa.
222 Flora Neotropica

~_______0__________ 50 40

C. sandwithii C. schottii
10
0

c. scottmorii 00 ..... s
c.?i~~~~~~~

HABITAT J , ./. . .HABITAT

Terra firme forest \

I Terra firme forest

F AtJIAISION D IJIFIMAIMIJ
JIG
FRUITI iIM II I1 II \s IhIFI IT
FRUI JIAISIdNID
I
FLOWERI
i I ll I i ll I FLOWERS
1_
60 40
.__ 80 70 60

FI 1 C. scottmorii andC.
spicatai

HABITAT HABITAT ..
Montane & Cloud forest 10 Terra firme forest. . .........

FRUIT I II II I IFRUIT
HIT1/
FLOWER ::::::g:FSII:: ler -LOWER
80 70 go

FIG. 115. Distribution of Couepia sandwithii, C. schottii, C. scottmorii, and C spicata.

Distribution Maps 223

60 , 60

C. steyermarkii C. stipularis
10 I -
1o (

Montane oud forest\ Terra firme forest

IFIMAIM
--- JAso ND JIFIMAMJIJ IAIS N \
/) I / FRUIT
FRUIT I I I _ Il /I) I
_- _ _ - - ___ I
60 60
80 70 60 50
'
/y'. \-'. sub/rdC.
{err/
,;M:jAO:

MA!l
JFR iIAI -: I

.
80 .o o7 50
..

FIG. 116. Distribution of Couepia steyermarkii, C. stipularis, and C subcordata.

224 Flora Neotropica

70 60 504

C. ulti

50

HABITAT

^
| \TELw R: ilFMIMIJIJ (A)S\ N --- -; A 2
BiiSjai5i^.)-
FRUIT |N|D| <> . 'I ^
____ ->
-.-._
_FLO\VER_
: g |."._
B -__ t '--
/ -~^
80 70 60 50

FIG. 117. Distbution of Couepiauti and C. ule.

FIG.117. Distribution
of Couepiauiti andC.ulei.
Distribution Maps 225

60 50 40

C. trapezioana f /jC. venosa

'
1O 0

- HABITAT
10.HABITAT --$--hOl I

t030 ^
0^nffi^flS^
FLOWERI I I I I I I I * I I I L\ FLOWER I 111
g _

FI.180 Do 70no 6CtCvna adCwlimi

i 0

60 4U 50

80 70 60 50
 100 90 80 70 60
------------- H
-----------

A 0;S 0

HABITAT
10Rheophyte
Montane & Cloud forest

Atazonian caatinga
avanna margins
Savanna .
Cerrado.......
Secondary forest
Coastal thickets ____
Arid formations. caatinaa
2 White
10 sand forest
White sand or campina
orest or campina \ - -
Terra firma forest
Distributionoe
Gallery forest margenus FIG. 119.
Slope forest
Beaches
Restinga
Seasonally flooded forest

100 90 80 70 050

FIG. 119. Distributionof the genus

Distribution Maps 227

70 60 80

H.adderleyi H. adenophora
-
in - ----- T.
r,,
7.\^-^^-^g^ ______^ 10-- 10
- - - --O

0.

FL l
IcT
T
OWE0 I 0

JIFIMAIJM,A SOND . ? ^ VIFIMIA JJ ASN ID

| IVE
FRUIT |II
(I|FRUIT ; II FRUITI I
- ! 1.|,/I!
1, ^^ - ,
FLOWER ggggssass8-- US^^ \

90 800 60

10

o r o~~~~~~~~~~~~~~~~~~~~~
lkz v

I-

90 80 70

G. 120. of Hirtellaadderley, H. adenophora,and H.americana.

Distribution
FRUIT.i
FLOWER 0i
O ,6 o ~ ~ 80706

FIG.120 Ditriutin o Hitela aderlyi,H. denphoa, nd . aeriana

FIG. 120.
Distribution of Hirtella adderleyi, H. adenophora, and ame
H. ricana.
228 Flora Neotropica
I
. -- -.? , 50
50 -- ___4__50 40 1 60 50
/ H. angustifolia H. angustissima

~(-'-I)~I

HABITAT / 1 HABITAT -
Terra firme forest
TRhophyte
FRUI fLI I I IFIAIMIJ
I I JAIS ND
FLOWERI I I 11 1~FRUIT ER
I I I

60
5^^0 ________4_____508,50
0___________0_0______ 80 70

: H. araguariensis H. aramangensis

FiBA F7-- "'^:' T^~ 10 HABITAT

Terra firme forest / yTerra firme forest
JIFIMAIMJJI|J
IAS X II J
IJIFMIAM
JAISJOINIDI ( oo
FRUIT I _FRUI I
1--1/ I1I IFRU I Ji
FLOWER. LOWER iI I I i I
60 50 80 70

FIG. 121. Distribution of Hirtella angustifolia, H. angustissima, H. araguariensis, and H. aramangensis.

Distribution Maps 229

60 50 40

.-.-'H.arenosa H. bahiensis

10
HABITAT F(AB /( ( ( J1 HABITAT
White sand forest or campina Terra firme forest
FRI JIFIMMAI|JIJIAISoIOIN j 1170fiAIMIJJ
AISIOINIDI
FRUIT I I I I I I-I IF I I 'f---- I II RII
WE'_-
IFLOWER
- I I T-l-I-I ell
14/ ,I I I ~
604U

70 _ 50 40

H. barnebyi, and
H.H.
barrosoiID

HABITAT _ ----- >- ---I- F ABITAT

Terra firme forest Terra firme forest

FLOWERIG
12 isrbt o Hrel a , HFL.OahEnRs

FIG. 122. Distribution of Hirtella arenosa, H. bahiensis, H. barnebyi, and H. barrosoi.

230 Flora Neotropica

70 60 50 40

ELOWER
IH.
I/^^ \ bicornis
O var. bicornis

Terra firme forest

Restinga
Beaches
Savanna margins

*
FRUIT H * f )
FLOWER
________0 40
80 70 60 50

oO^^V^,^?^J S"^H. bicornis

^l^ ?
ST(/ ^var. pubescens

1 Terra firme forest

FIG. 123. Distributionof Hirtella bicornis.

Distribution Maps 231

80
__?^^"*^^70 60 70 60 50

......;.'.....~~ .
... .
. ~
. .

JF"IG1Dsb o o( H70 l 60

1 .1
Gallery forest _

Gallery -^~~`
forest
'erra~~~~~
fim oet
M.^^____ /
-
____________ T^^S.""^
//^Y(
:
'^?2
^
--T^ ^ - -/-
/
^~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~
__ "
_j V "v *^- -^^ ^
^
"^^- 0

Terra firme forest ..

HG24 itrbton of HIrelblatan fbucl

Savanna G. 124. Distribution of Hirtella bullata and H. burcheli.

232 Flora Neotropica

70 70 00
Gallery forestGaller forest \ (
Seasonally flooded forest / -Savanna margins / ) f \

70 . 60
_I6 I
,,60 , m

0 / H. carbonaria io _H. conduplicata10

0
\I 0

JI JFMA 1
FRUIT JIFIMIAMJIJjAISIOINIDI-_ FRUIT
J /F/.' SN
"
isl
FLOWER15 tggri FbOWER
H c

70_60 80 50

FIG.
125. Distributionof Hirtellabrachystachya,H. caduca, H. carbonaria,and H.
conduplicata.

FCoastal thicketsI ,_\ HABITAT

Mangallove margins I TGallerra
frme forest

FIG. 125. Distribution of Hirtella brachystachya, H. caduca, H. carbonaria, and H. conduplicata.

Distribution Maps 233

70 60 50 4

^'^X^ -H. ciliata

0S r 'i 0

HABIT ATi |giii * ^ V

706060 60

H. confertiflora OC H. cordifolia

FHABITATG 126. Dtbi HiABITAT fr a H orflia.

Gallery forest &\\''- Savanna W
0 JIFIMIAMJ IJiAIS OFN D J F MA JJAS ND
FRUIT | I I > -
I II A 0 FRUIT
FLOFLWERI I I I I
ELWE II :II /

FIG. 126. Distribution of Hirtella ciliata, H. confertiflora, and H. cordifolia.

234 Flora Neotropica

s40 50

H. corymbosa H. couepiiflora

'?.i i

40 50 iS-0

60
60_____________________ _50
o, o H. cowanii o H. deflexa

12. D
FHABITATG. b /fi . "ym AT
HABIT
Montane & Clou forestGaller
foreste :
JFMAMJJASNP J FIMIAIJ IJASON
\[
FRUIT I FRUIT __I___ I \
1)
FLOWERI li I lI_
/(1 ( FLOWER _~_~__ _ _ ~_I ( \
60 _ 60

FIG. 127. Distribution of

Distribution of Hirtella
Hirtella corymbosa, H.
H. couepiifiora,
couepiiflora, H. and H.
cowanii, and
H. cowantii, H. defiexa.
deflexa.
Distribution Maps 235

i0m~~~~70
00, ..o H. davisii

Terra firme forest / / / /

80 7 0 60 50
0
"^^H^^'^MB^^"B^^B^^50 70 60

? H. dorvali H. duckei
rf
101

IW: g
:F/LOWEg II I \ F

FIG. 128. Distribution of Hirtella davisii, H. dorvalii, and H. duckei.

G. 128. Distributionof Hirtella.ds, H. dra, and H. ducke.

236 Flora Neotropica

80 70

'- H. elongata

70

Gallery forest Y* 3*3i 7(,

R
,2i0 |*

FLO
| ER 80
80 7.0 60
60 50 0
50

80 70 60 50

01 ~ ~ FG 129 Ditiuino itlaeogtn. ...........

Secondary '
forestI D st riuto1 o l e t an H
Seasonally flooded forest \ r \
O ND \
JIFIMAIM Jl IAISOINIDI \ I
^ V
FRUIT
*Ig~ H 114 ^\
I "~i"' Y- 1 w ^
8U 70 60 50

FIG. 129. Distribution of Hirtella elongata and H. eriandra.

Distribution Maps 237

80 70 70

?t enneandra
AH. ;.. " H. excels

Y~3
FRI I v I'-v/ _ I I II
*y_ yI R

Montane & Cloud forest Terra firme forest-

FRUIT !I1
I 11 ! FRUIT ? --:7-"'- --
IFLOWER
8 0_/__ 1 1 I1 . i7-J i FLOWER I
0.70 !
_ 60 , 50 50 40

o H. fasiculata H floribunda

1
FIG.O~~~~~~~~~~~~!

Terra firme forest \

./ '
FJI3FIMIAAMIjIJ ^IS ONIDI30IN
- J
FRUIT I I -'1 FRUIT I I' I I I
FLOWER_ I I i I)'
_III:I _ ::1 FgLOWER ! I
U 5 74U

FIG. 130. Distribution of Hirtella enneandra, H. excelsa, H. fasciculata, and H. floribunda.

238 Flora Neotropica

70 60 50
^^^ l-^o^^ ~~~~~~~~~~
?e H. glandulosa
0
O 0
1~~~~~~ /
0~~~~~~~~
o 0
0~~~~~~~~~~~

io
'^^^^

..... .... .. ........::

7-
0 *
1 -- f "^ -- 'p ^ 7 - ~ Y ~ - V
-T^
1y Y 1
-

BITAT~~~~~~~~~~~~~~~ ....
..........

J F A J JA

3C Savan,a .... ..
:;_____________\ ^--(

----
FRUIT ~ ~~ ~ ------^ '^Z // ~-- -

I I
!Savanna margins _____ - ~- - ~
iip~ Y7 ^ ^
i
iele!ll elel30
,FLOWERlell!iillell!l~ele!tel llllel ...._F l RUIT
70 60 50 40

FIG. 131. Distribution of Hirtella glandulosa.

Distribution Maps 239

60 50

10. btH. glabrata,1 H. glandistipula

HABITAT
avanna
econdary forest HABITA
hite sand forest or campina Terra firme forest 10
1 JFMAF
MA Ml J AIS 0 NIDI J-I )
MAIMIJIJIA SkND I
FRUIT 0 FRUIT
FLOWER B 1/I E 1 I FLOWER
6050

H. glaziovii H. guainiae

0
5I.-^^ ^^ 8^T^0 ^^0
^ 0

HABITAT BITAT
Terra firme forest Terra firme forest
JF AMJJ AS ONDJ fM J SN
FRUIT I I I I I I I I I ITI
|pFRU III /I |
FLOWER ________lgft FLOWER M"[gbbI 1-SSl 0
____I-I
L_^^^^^^^ ^^40^ _ _80 -U

FIG. 132. Distributionof Hirtellaglabrata,H. glandistipula,H. glaziovii, and H. guainiae.

240 Flora Neotropica

70 60 50

A/ / -/ . gracilipes
',,\H.

1~_ ____ 90_________ 80 _ _ __ 60__

0 1

F guatemalensis
iraH. H.a guyanensis
. ::
L
WERf:g .. :' ...........

FIG0 I

133. D
FIG.90 n of0 l 60
FIG. 133. Distributionof Hirtella
H. guatemalensis,
gracilipes, and
H. guyanensis.
Distribution Maps 241

80 70 60 50

^^^^:^? H. hispidula 10

4 0

FRUIT
aiFLOWER iFLOWER FLOWER
80
50__.o 70 60
50 4 40 60 50
_

FIG. 134. Distributionof Hirtella hispidula,H. hebeclada,and H hoeh nei

0 H.hoHAeIThAT-

50 40 1 60 50
242 Flora Neotropica

50 40 60 %/ ir50u
H. insignis H. juruensis

10~ ~ ~ ~ ~ ~~ ~10

AT.T.-..
.. . . ....

1 0

BITAT
Restinga - J/ I

FRUIT FRUIT I
FLOWERI Ii- 1I1111 ____ 'FLOWER
40__ 60 50
60 50 50 40

H. kuhimannii H. lancifolia

0 ABITAT 10
HABSecondary forest
Terra firme forest ( T erra firme forest
^
IJFMIAIMIJJIAISOINID
/n\ I_ JFMA^JJA As
-/,IS o /
FRUIT !FRUIT
I II I I!
FLOWER _I _ / .
)FLOWER
60 50 _ 50 40

FIG. 135. Distribution of Hirtella insignis, H. juruensis, H. kuhlmannii, and H. lancifolia.

Distribution Maps 243

80 70 80

{FLOWER---H. = latifolia H. lemsii

10 -M 010 0

0/D~~
0

0 0
70 V

Terra firme forest Terra firme forest

H. leonotis H. liesneri

Terra firme forest Slope forest /

FRI J3FM.AJDJAoND I IJFM JAISH N sn - .

II
OI I I I FRUIT
I II I I I LOWE
I-I
II
II [:L
|FLOWER_I- 1Fl 7 - I
70^

FIG. 136. Distribution of Hirtella latifolia, H. lemsii, H. leonotis, and H. liesneri.

 244 Flora Neotropica

70 70

- H. lightioides H. longifolia

10, 7 6-- 10

3
~

Terra firme forestfHtligod,H Amazonian caatinga t,a .

FRUIT I FRUIT

70 70__________
____7_______60 _60_______________ 50

H. longipedicellata 0 H. macrosepala 0

'
L
HABITAT
Savanna margins F _ITAT
Slope forest Terra firme forest

J(FIMMAIMIJJAS OND J JASON D .....

FRUIT L:::I: --------R
RUT-
10I I
TR-U
FLOWER I ^ .w "
I_^ \ FLOWER
_60 70 60 50

FIG. 137. Distribution of Hirtella lightioides, H. longifolia, H. longipedicellata, and H. macrosepala.

Distribution Maps 245

80 70 60 50

H. magnifoliaS:7- H. macrophylla
H.

,
forest : .
_0_Secondary ........,, .

880 70 60 50

.
1. H. magnifolia a nd H. margae

Seasonafirme foodrest Terra fe fos ' 0

I JIAISO\
IJIFFIMIAIMIJI NII /
~
rFLOWERFI I I = = =II
_\ = il !.'?" F
b o
G80 70

HABITAT \ F0A
Terra firme forest Terra firme forest

FRUIT ____________ ^\ ) FRUIT *_ I

FLOWER _ _ -- FLOWER_
80 70 6U 56

FIG. 138. Distribution of Hirtella macroPhylla, H. magnifolia, and H. margae.

246 Flora Neotropica

50 40 60 50

- H-.martiana > H. mucronata

?10F .. ........

HABITAT- - .....
Swamps I HABITAT 10
Gallery forest . Terra firme forest
IJIAISIA
|J|FIM|A|MIJJ NID
|A|S|O|N|D|I J F_AIQJ
FRUIT 1 1 1 I I I I FRUIT ----- _ _
FLOWERI 0g;gggI lO 11 F
LFLOWER g J
40 50
80 70 60 50

H. mutisii H. myrmecophila
10 10
.0

'erra firme forest __[erra firme forest 1 j

0 "~/U b'HABIT_T U ....o
139.Distribution
FIG. ofHirtella martiana, H. mucronata, H. mutis, andH. myrmecophla.

FIG. 139. Distribution of Hirtella martiana, H. mucronata, H. mutisii, and H. myrmecophila.

Distribution Maps 247

60 50 60

H. obidensis %
H. orbicularis
(,. o ,
1 03

Terra firme forest Montane & Cloud forest

HG. 10 itbtnbor s H. paniculata

FRU II F L O,...

80 7060 50

FIG. 140. Distribution of Hirtella obidensis, H. orbicularis, and H. paniculata.

FIG. 140. Distributionof Hirtella obidensis, H. orbicularis, and H. paniculata.

248 Flora Neotropica

50 50 40
,)H. H. parviunguis
: paraensis

0. 0

FHABITAT
/ /I HABTij

--
50 50 402
90 80 70 60

G1 H. pauciflora ' CH. pendula

0 0 0
(.

-^'0 0
Id.HABII-AT
1j^^V^-\J^ t-II0HABITAT ^ 0
Terra firme forest . Terra firme forest
ZZZZJ AIJ IJ IA NID00 "^ / <Z AM
J IFM J IA SON D0
FRUIT B I .' /y FRUIT
FLOWERI I: I I I I I I I: I`~IFLOWER ^
/^
90 80 70 60

FIG. 141. Distribution of Hirtella paraensis, H. parviunguis, H. pauciflora, and H. pendula.

Distribution Maps 249

80 70 60 50

fort,,.
Terrafirme
0

F 2 i7 6. . 5
..
I IJIFIG. JIJ IoI S li

FIG ofHirtella
142.Distribution ilosissima
andH. physophora.

n of
FG14.Dsrbto Hit
piloism
el
nHhspoa
250 Flora Neotropica

70 60 60 50

H. pimichina H. piresi

l ..

HABITAT? // /J -HABITA..
/r^ / _ _,__.~_T 0
.......f^
Savanna margins Terra firme forest
JFIF AM
AJ TJ AS N
ND) 1FMAJ F= AI
1FRUI FRUI T I I III I

---
1 ~i10
^s:^. 'C

0~~~O7

FIGOW. Distributn of H
143. .a p pii H. p til , ad H. r mii
60 50 70

FIG. 143. Distribution of Hirtella pimichina, H. piresii, H. punctillata, and H. radamii.

 100 90 70 60
80,
o, H. r
l2(
C^^^^^--^^ .~ C:

10

10:BI
TATI

....' .......

White sand forest or campina( /

Terra firme forest'-'

JSav'7nna " =l ) -J
/---i- \
Savanna margins~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~^
,\ ;
FRUIT-I i ***__* **, .
FLOWER
100 90 80 70 60

FIG. 144. Distribution of Hirtella racemosa var. hexandra.

252 Flora Neotropica

80 70 60 50

H. racemosa
10
10 *<-
Ooe
-
^^^P^^^
?^ ---- var. racemosa lo

? ?i 0

10
'[

HABITAT
firme forest -- '
Terra
"
Gallery forest . O. . -
Seasonally flooded forest / I/ _
0
White sand forest or campina '
AIFIMIA 4JIIJIAISI 0N IDI
FRUIT
FLOWER
80 70 60 50

FIG. 145. Distribution of Hirtella racemosa var. racemosa.

Distribution Maps 253

80 70 50

/ H. racemosa H. racemosa
var. glandipedicellata var. hispida
0

l , (e
80 - - 10 .. ....
20 HABITAT- .0

FRUITII11I ~ F
II I I I
FLOWER

80
80 70
70 ...... 50
80 70 80 70

?
.^ rasa| | | | /: H. revillae

' /U

' 0
20

Ho e H -H.revillae
rasa

20HABITAT .. BITAT
Terra fiorest forest
SlopTerra
firme ( ,\' White
White sand
sand forest
forest or
or campina
campi'"'
JIFIMAMJ IJIAS - 7h'te ; IJ;ANi.,'
JFMAJJASND" o0
sIa'!j;l~
FRUIT UIT I
FLOWER! i I IFLOWRI
I I I I,
70 80

FIG. 146. Distributionof Hirtella racemosavar. glandipedicellata,H. racemosavar. hispida,H. rasa, and
H. revillae.
254 Flora Neotropica

70 60 70 60

H. rodriguesii H. rugosa

''I Wl" I I0I 0

'TTHABITAT
/Bi\
HAB17AT-^4' ~ >
f .
Terra firme forest \Montane & Cloud forest -
I"
JIFIMIAIMIJIJIAISND\ J IIAIMJ IAISoNI

50 40 I 80 70

G 1... H. santosii r a H.ascaberula

HABITAT^k \ ^0
--:????./ 0.ABITAT 0

Terra firme forest Amazonian caatinga

JIFIMIAIJI As AoNNJIFIMIA1J
I I I IJIAISND
FRUIT IFRUIT I I I I I_ I I I
FLOWER__I
II I I I I I ^- FLOWER I_______II I
4U80 Tu70

FIG. 147. Distribution of Hirtella rodriguesii, H. rugosa, H. santosii, and H. scaberula.

Distribution Maps 255

70

g H. scabra H. schultesii
10

10I 120^ .. I I I .y .

/ v
HABITAT
HABITHAT ^/ ^ M^
Savanna IJ F M J JWSTerra \ t
firme foresi .e_
IJ i AS A
FRUIT
FLOWER FOlO_WE_ll iTJ( I
Is I

0
FIG.148. Distribution
of Hirtella H. silicea,
scabra,H.schultesi, H.'/2sprucei
and

10C0

vTerra
time forest .Terra firme forest

_~60 _ - bU _ _ 4u

G.
148.Distribution scabra,
oHirtlla H.schultesii,
silicea,H. and
H.sprucei.

FIG. 148. Distribution of Hirtella scabra, H. schultesii, H. silicea, and H. sprucei.

256 Flora Neotropica

80 70 70

r H. standleyi H. subglanduligera

~~>FRUIT . I I-
LFRUIT- I I_
__ 01r 8_
7 _ 1750_= -?c'HxIAS0
71 0 0 -606

0 50
60 60

1
HABIT11\IAT /I II\T-A IL I I I I

I
NIFLOlWER___g__ __I |I 1FLWE T IE I SI

FIG. 149. Distributionof Hirtella standleyi,H. subglanduligera,H. subscandens,and H. suffulta.

Distribution Maps 257

60 50 60 50

H. tentaculata H. tocantina

Seasonally flooded forest tubifBITAT H.

Terra firme forest Terra firme forest X
JIFIMIAIMI
JIJIAISIOIN eJ FMAIMIJ
IJIAIS
I
FRUIT _ _ _ _ 6___ _ FRUIT //
FLOWER
_ _:_I
'*ggS_rl I @1 1I I =IFLI J
60 50 50

60 50 80

r'oLH. tenuifolia H. tubiflora

s
Secondary forest .----^ 1 HABITAT^-_ /==
Terra firme forest / . . Terra firme forest >1

IFRUITI|I I|I IIJ||IN11 I l Y

JMX 2/ 1 ^^ 1

FIG. 150. Distributionof Hirtella tentaculata,H. tocantina,H. tenuifolia,and H. tubiflora.

 100 90_ 80 70
^
/ "I ^~ <r;^^^ ^3^^
^
50

Savanna margins s

Slope forest

--3------
---FRUIT------2

.'I
FIG. 151. Distributionof Hirtellatriandrasubsp
Distribution Maps 259

90 50 40

-H. triandra H triandra

subsp. media20 1 subsp. punctulata

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

HABITAT
HABITAT Terra firme forest
Terra firme forestSlope lope forest
forest -
-- JJ IAIS
JIFIMIAIMI
_ - OIND IJIFIMAIMIJIJ ID
IAIOI N
FRUIT - L-FRUIT I I I I I I I I I
FL~OWERI T11l
__1 II@
FLOWER I1___
II 1 ___I@ FLOWERi:::II I r
rILOWERI Bs*g 1 L __
90 40
U'-60 70 60

'..- JH.ulei H. vesiculosa

- 0

-o- -- 0
,

HABITAT .. ..
Savanna .....
White sand forest or campina /0
Beaches

FRUIT= '! FR! 1 II I I II I

FLOWER : :FLOWER . III I I /
70

FIG. 152. Distributionof Hirtella triandrasubsp. media, H. triandrasubsp. punctulata,H. ulei, and H.
vesiculosa.
260 Flora Neotropica

90 8070

1^\-_ ^^ panamensis 0
o
-JMaranthes

olF RUI

0 ~~ ~ I-rr
FLOWER I _ _ _ __ _. _

Slope forest
Amazonian caatinga
Terra firme forest 1
__I-80____-70_ _ 60 50

FIG. 153. Distributionof Maranthespanamensisand the genus Acioa.

Distribution Maps 261

60
0 o , I50 _7 60_I

- <:A. guianensis A. schultesii

Esl /M I\I A N|D

I0 (J)\

l ). ... ....... HABIT

........
9Terra
2orest
firme l Amazonian
_ caating_______
iJA .:' N
FI1 Dio Aaa RUITs
IA. !1 A s I

_ 80
80 70
70 60 5050

FIG.,,15.rio?. A
.sA.suii-a somnolens

Terra firmGforest rI I g an ni ch ti a s .

80 70 60 50

FIG. 154. Distribution of Acioa guianensis, A. schultesii, and

A. somnolens.
262 Flora Neotropica

HABITA

'Ir\ 1 )'II iJI)I MJIJ

1' II() EL1
\~~~~~~~~~~~~~~~~7
FI.15 Lcnadosni(Aeeo&
sectin; E petl; F,ovar andstyl. ay159.A
Inst: dstriutionof L ait
dodonii,oenn lwe u;C,foe; ,foe

FIG. 155. Licania dodsonii (Acevedo & Daly 1659). A, habit; B, opening flower bud; C, flower; D, flower
section; E, petal; F, ovary and style. Inset: distribution of L. dodsonii.
Indexof ScientificNames 263
INDEX OF LOCALNAMES
apacharamaamarillo(Peru)95 lobo apacharama92 parinari58
bokobokotokon89 macucurana95 parinari(Peru)51
carip6torrado93 maeneyowaie(AukaIndian)20 parinariblanco 58, 65
castanhade cotia 97 milho torrado70 pasista 96
coquito 95 milho-torrado-amarelo55 rode kwepi 49
cuero de sapo 59 mulo (El Salvador)27 uchirana(Brazil)65
duship 30 oiti 46, 55, 58 yakuku58
gaulette(Fr. Guiana)95 oiti-mirim 57, 77 yukuku(Peru)27
guaiti-mirim59

INDEX OF SCIENTIFICNAMES
New names and combinationsare in bold face and synonymsare in italic. Page numbersin bold
face indicate primarypage references.Page numberswith an asterisk(*) indicate pages with illus-
trations or maps.

Acia amara 64 divaricata65

Acioa 2, 3, 96, 97, 260* divaricatavar. strictiuscula65
amara64 dolichopoda75, 76, 104, 208*
edulis 96, 106, 209* edulis 2, 96
guianensis96, 106, 261* elata 104, 209*
schultesii96, 106, 261* eriantha104, 209*
somnolens 106, 261* excelsa 103, 209*
Arecaceae53 exflexa 103, 210*
Barcellaodora 53 foveolata 72, 103, 210*
Caryocaraceae2 froesii 104, 210*
Chrysobalanaceae2, 3, 4, 91 glabra74, 75, 104, 210*
Chrysobalaneae3 glandulosa65, 103
Chrysobalanus3, 4 grandiflora104, 211*
cuspidatus6, 99, 128* guianensis64
icaco 3, 4, 6, 99, 127* subsp. divaricata65, 103, 211*
interior4 subsp.glandulosa64, 103, 212*
venezuelanus4, 5*, 6, 99, 128* subsp.guianensis64, 103, 212*
Clusia 70 habrantha70, 103, 213*
Colpothrinaxcookii 70 impressa77, 104, 213*
Couepia59, 64, 65, 72, 97, 203* subsp.cabraliae77, 104, 213*
amaralae72, 73*, 104, 204* subsp.impressa 104, 213*
belemii 71, 103, 204* insignis 70, 74, 104, 213*
bernardii65, 66, 68, 103, 204* krukovii72, 103, 214*
bondarii70, 74, 104, 204* latifolia 104, 214*
bracteosa103, 205* leptostachya64, 103
canescens68, 72, 103, 205* longipendula76, 104, 214*
canomensis 103, 205* longipetiolata77, 79, 104, 214*
carautae70, 71*, 103, 206* macrophylla72, 103, 215*
caryophylloides72, 103, 206* magnoliifolia70, 103, 215*
subsp. caryophylloides72, 103 maguirei103, 215*
subsp.glabra72, 103 marleneae75, 104, 215*
cataractae103, 206* martinii74, 104, 216*
chrysocalyx104, 207* meridionalis77, 79, 104, 216*
cidiana74, 104, 206* monteclarensis66, 67*, 68, 103
coarctata77, 78*, 104, 207* multiflora104, 216*
cognata68, 104, 208* myrtifolia65
var. cognata68, 104, 208* nutans76, 104, 216*
var. major 104, 208* obovata 65, 66, 104, 217*
var. membranacea104, 208* ovalifolia 104, 218*
comosa 76, 104, 207* paraensis219*
264
subsp.cerradoanda103, 219*
subsp. glaucescens103, 219
subsp. paraensis103, 219*
parillo 39, 103, 217*
parvifolia 104, 218*
perambucensis 79, 104, 218*
platycalyx76, 104, 218*
polyandra76, 104, 220*
racemosa 104, 220*
recurva74, 104, 221*
reflexa65, 70, 104, 221*
robusta 104, 221*
rufa 104, 221*
sandwithii65, 66, 68, 103, 222*
schottii 79, 104, 222*
scottmorii68, 69*, 103, 222*
spicata70, 103, 222*
steyermarkii72, 103, 223*
stipularis104, 223*
subcordata103, 223*
surinamensis64
trapezioana75, 104, 225*
uiti 103, 224*
ulei 104, 224*
venosa 104, 225*
versicolor64
williamsii 75, 104, 225*
Dactyladenia2, 3
Dichapetalaceae2
Exellodendron59, 200*
barbatum59, 103, 201*
cordatum59, 103, 201*
coriaceum59, 103, 202*
gardneri59, 103, 202*
gracile59, 103, 202*
Hirtella79, 91, 92, 226*
adderleyi88, 91, 105, 227*
adenophora106, 227*
americana91, 105, 227*
angustifolia95, 106, 228*
angustissima105, 228*
araguariensis88, 89, 105, 228*
aramangensis105, 228*
arenosa93, 106, 229*
bahiensis92, 105, 229*
bamebyi 88, 105, 229*
barrosoi 105, 229*
bicoris 105, 230*
var. bicomis 105, 230*
var. pubescens 105, 230*
brachystachya106, 232*
bullata 105, 231*
burchellii 106, 231*
caduca 106, 232*
carbonaria105, 232*
ciliata 105, 223*
cliffortiana106
condifolia 105, 233*
conduplicata93, 106, 232*
confertiflora87, 88, 105, 233*
cordifolia 105, 233*
corymbosa91, 105, 234*
couepiiflora95, 106, 234*
cowanii 105, 234*
davisii 91, 105, 235*
FloraNeotropica
deflexa 105, 234*
dorvalii87, 104, 235*
duckei 87, 104, 235*
elongata88, 92, 105, 236*
enneandra106, 237*
eriandra88, 92, 105, 236*
excelsa95, 106, 237*
fasciculata106, 237*
floribunda106, 237*
glabrata88, 104, 239*
glandistipula106, 239*
glandulosa105, 238*
glaziovii 95, 106, 239*
gracilipes106, 240*
guainiae 104, 239*
guatemalensis105, 240*
guyanensis105, 240*
hebeclada106, 241*
hispidula 106, 241*
hoehnei 105, 241*
insignis89, 105, 242*
juruensis 106, 242*
kuhlmannii94, 106, 242*
lancifolia 106, 242*
latifolia 105, 243*
lemsii 94, 106, 243*
leonotis 92, 105, 243*
liesneri 89, 90*, 105, 243*
lightioides 105, 244*
longifolia 106, 244*
longipedicellata106, 244*
macrophylla105, 245*
macrosepala104, 244*
magnifolia92, 105, 245*
margae88, 105, 245*
martiana95, 106, 246*
megacarpa106
mucronata94, 106, 246*
mutisii 105, 246*
myrmecophila104, 246*
obidensis 105, 247*
orbicularis105, 247*
paniculata105, 247*
paraensis106, 248*
parviunguis94, 95, 106, 248*
pauciflora95, 106, 248*
pendula 105, 248*
physophora87, 104, 249*
pilosissima 105, 249*
pimichina 106, 250*
piresii 105, 250*
pohlii 106
punctillata91, 105, 250*
racemosa93, 106
var. glandipedicellata106, 253*
var. hexandra94, 106, 251*
var. hispida94, 106, 253*
var. racemosa94, 106, 252*
radamii93, 105, 250*
rasa92, 105, 253*
revillae87, 104, 253*
rodriguesii105, 254*
rugosa 106, 254*
santosii 91, 105, 254*
scaberula106, 254*
Index of ScientificNames 265

scabra 105, 255* cordata 101, 140*

schultesii 106, 255* coriacea 101, 139*
sect. Myrmecophila79, 87 costaricensis100, 139*
silicea 106, 255* couepiifolia48, 101, 139*
sprucei 106, 255* crassivenia49, 101, 139*
standleyi 106, 256* cruegeriana55, 102, 141*
subglanduligera105, 256* cuatrecasasii33, 100, 141*
subscandens106, 256* cuprea45, 101, 141*
suffulta105, 256* cuspidata99, 141*
tentaculata105, 257* cyathodes 101, 142*
tenuifolia93, 105, 257* cymosa 46, 53, 101, 142*
tocantina 105, 257* davillifolia 101, 142*
triandra105 dealbata46, 101, 142*
subsp. media 105, 259* densiflora101, 143*
subsp. punctulata105, 259* discolor 102, 143*
subsp. triandra105, 258* divaricata100, 143*
tubiflora95, 106, 257* dodsonii27, 99, 262*
ulei 104, 259* durifolia20, 22, 26, 55, 99, 143*
vesiculosa 104, 259* egleri99, 144*
zanzibarica106 elliptica 101, 144*
Licania3, 6, 39, 72 emarginata100, 145*
subg. Licania36, 39, 44, 47, 48 fanshawei44, 101, 145*
sect. Cymosa44, 45, 46 fasciculata24, 26, 99, 145*
sect. Hirsuta36, 39 filomenoi20, 21*, 22, 99, 145*
sect. Hymenopus39 foldatsii 101, 146*
sect. Licania48, 53, 54 foveolata 100, 146*
sect. Pulverulenta47, 48 fritschii99, 146*
subg. Moquilea19, 24, 28, 29, 30, 32, 34, 35, 36 fuchsii 100, 146*
sect. Leptobalanus32, 34 furfuracea47, 101, 147*
sect. Microdesmia30, 35, 36 fusicarpa102
sect. Moquilea 19, 20, 27, 28, 29, 30, 32, 34, 35 gardneri99, 147*
subg. Parinariopsis6, 11 gentryi20, 24, 99, 147*
affinis47, 59, 101, 129* glabriflora40, 100, 148*
alba 101, 129* glabrifolia(sphalm.)40
albiflora35, 100, 130* glauca 101, 148*
amapaensis56, 102, 130* glazioviana 100, 148*
angustata28, 99, 130* gonzalezii28, 99, 149*
anneae 27, 99, 130* gracilipes101, 149*
apetala3, 32, 34, 99 grandibracteata20, 22, 23*, 99, 149*
var. aperta32, 99, 131* granvillei32, 99, 149*
var. apetala99, 131* guatemalensis29, 99, 150*
apiculata 102, 132* guianensis39, 99, 150*
aracaensis51, 102, 132* harlingii55, 102, 150*
arachnoidea39, 46, 100, 132* hebantha49, 101, 153*
araneosa36, 100, 132* heteromorpha40, 41, 42
arborea35, 100, 133* var. glabra 100, 152*
arianeae44, 45*, 101, 134* var. heteromorpha100, 151*
bahiensis 102, 134* var. perplexans100, 152*
belemii 55, 102, 134* var. revoluta40, 100
bellingtonii48, 101, 134* var. subcordata100, 152*
blackii 55, 102, 135* hirsuta36, 100, 153*
boliviensis 99, 135* hispida37, 38*, 100, 153*
boyanii 101, 135* hitchcockii 101, 153*
bracteata51, 102, 136* hoehnei 102, 154*
britteniana99, 136* humilis 100, 154*
buxifolia47, 101, 136* hypoleuca46, 101, 155*
cabrerae20, 24, 26, 99, 137* var. foveolata 101, 155*
caldasiana102, 137* var. hypoleuca 101, 155*
calvescens35, 100, 137* impressa46, 55, 101, 156*
canescens 101, 138* incana 54, 102, 156*
caudata39, 40, 100, 138* indurata102, 156*
cecidiophora29, 30, 99, 137* inpae 156*
chiriquiensis28, 29, 99, 140* intrapetiolaris41, 100, 157*
chocoensis 100, 140* irwinii 101, 157*
compacta 102, 140* jefensis 32, 99, 157*
266
jimenezii 49, 101, 157*
joseramosii34, 100, 158*
kallunkiae28, 29, 99, 158*
klugii 26, 99, 158*
krukovii 100, 158*
kunthiana101, 159*
laevigata41*, 42, 100, 169*
lamentanda51, 52*, 102, 160*
lanceolata53, 102, 160*
lasseri 100, 161*
lata 99, 160*
latifolia 100, 161*
latistipula40, 100, 161*
laxiflora101, 162*
leptostachya53, 54, 102, 163*
leucosepala27, 99, 162*
licaniiflora100, 163*
littoralis 101, 164*
var. cuneata 101
var. littoralis 101
longipedicellata27, 35, 99, 164*
longipetala30, 99, 164*
longistyla35, 100, 165*
macrocarpa19, 20, 22, 26, 99, 166*
macrophylla100, 165*
maguirei99, 166*
majuscula101, 166*
maranhensis99*, 167*
maritima19, 20, 26, 99, 166*
marleneae44, 101, 167*
maxima 102, 167*
membranacea101, 168*
mexicana34, 100, 167*
michauxii 3, 99, 168*
micrantha51, 102, 170*
microphylla102, 168*
miltonii 40, 100, 171*
minuscula 100, 170*
minutiflora27, 28, 29, 99, 171*
mollis 102, 172*
montana20, 26, 99, 172*
morii 33, 99, 172*
naviculistipula101, 172*
nelsonii 53, 54*, 102, 169*
niloi 101, 173*
nitida 102, 173*
oblongifolia100, 173*
obtusifolia 102
occultans42, 100, 169*
octandra35
subsp. grandifolia35, 100, 175*
subsp. octandra35, 100, 174*
subsp. pallida 35, 100, 175*
operculipetala100, 175
orbicularis101, 176*
ovalifolia 49, 102, 176*
pakaraimensis39, 100, 177*
palawanensis102
pallida46, 101, 176*
paraensis54, 102, 177*
parviflora102, 177*
parvifolia99, 178*
parvifructa101, 178*
FloraNeotropica
persaudii100, 178*
piresii 46, 101, 180*
platypus99, 179*
polita 101, 179*
pruinosa102, 180*
pyrifolia99, 180*
reticulata26, 42, 100, 182*
retifolia99, 180*
riedelii 46, 102, 182*
rigida 100, 181*
robusta53, 102, 182*
rodriguesii102, 183*
roraimensis102, 183*
rufescens101, 183*
salicifolia36, 100, 183*
salzmannii99, 184*
sandwithii 101, 184*
santosii 46, 101, 184*
savannarum102, 184*
sclerophylla100, 185*
silvae 44, 101, 185*
silvatica 100, 186*
sparsipilis33, 34, 100, 186*
spicata 102, 186*
splendens 102
sprucei 100, 186*
stewardii50, 102, 187*
steyermarkii49, 101, 187*
stricta 102, 187*
subarachnophylla36, 100, 187*
subrotundata101, 188*
tachirensis30, 31*, 99, 188*
tambopatensis36, 37*, 100, 169*
teixeirae47, 48*, 101, 188*
tepuiensis 102, 188*
teratensis 101, 189*
tocantina50, 102, 189*
tomentosa99, 189*
triandra50, 102, 190*
trigonioides101, 189*
turbinata99, 191*
unguiculata30, 99, 190*
urceolaris47, 101, 191*
vaupesiana55, 102, 191*
velata 20, 100, 192*
veneralensis20, 26, 55, 102, 192*
wurdackii99, 192*
Maranthes59
corymbosa59
panamensis59, 103, 260*
Moquilea64
couepia64
glandulosa65
Myrtales3
Neocarya97
macrophylla97, 106
Parinari56, 57, 58, 72, 97, 193*
alvimii 57, 102, 194*
brasiliensis57, 102, 194*
campestris57, 102, 194*
canescens68
cardiophylla103, 195*
chocoensis 58, 103, 195*
Indexof ScientificNames
excelsa 57, 102, 196*
klugii 57, 102, 195*
krukovii65
littoralis57, 58, 103, 195*
maguirei58, 103, 197*
montana 102, 197*
obtusifolia 103, 197*
occidentalis 102, 198*
267
pachyphylla102, 198*
parilis58, 103, 198*
parvifolia 103, 198*
rodolphii 57, 102, 199*
romeroi59, 103, 199*
sprucei 102, 199*
Rosaceae3
Theales 3