Organization for Flora Neotropica

Chrysobalanaceae
Author(s): Ghillean T. Prance
Source: Flora Neotropica, Vol. 9, Chrysobalanaceae (Jun. 8, 1972), pp. 1-409
Published by: New York Botanical Garden Press on behalf of Organization for Flora
Neotropica
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FLORA NEOTROPICA
Monograph No. 9
CHRYSOBALANACEAE

by
Ghillean T. Prance

C.^,\ (~ 0 ~,"\ ~ TROPIC OP CANCER

_ _ ,------- ----- -------- I....

FLORA
NEOTROPIC

TROPIC OF CAPRICORN

Publishedfor
Organizationfor Flora Neotropica
by
Hafner PublishingCompany
NewYork
June 8, 1972.
 Copyright? 1972

HAFNER PUBLISHING COMPANY, INC.

Publishedby
Hafner Publishing Company, Inc.
866 Third Avenue
New York, New York 10022

Libraryof CongressCatalogCardNumber 70 -180014

Printed in the U.S.A.

 A MONOGRAPH OF NEOTROPICAL CHRYSOBALANACEAE

GHILLEANT. PRANCE*

INTRODUCTION

This monograph was intended to be the second major publication on the

Chrysobalanaceae, and was preceded by work on a world-wide survey of the family
at the generic level (Prance, in press) which, however, has had little effect on the
circumscription of the genera presented here. The earlier study has met with a
series of unforeseen delays in publication, and will in fact be preceded by this mono-
graph of the neotropical species. The full details of the genera on a world-wide basis
will be published shortly, and only relevant aspects of the earlier study are presented
here.
The richness of Chrysobalanaceae in the tropics of the New World is reflected
by the fact that the present treatment accounts for nearly two-thirds of the species of
the family. During the past several years I have made repeated visits to tropical
South America and have studied in the field 92 of the neotropical species. Some of
the commonest ones have been encountered many times in widely separated stations,
affording me opportunities to assessand measure variation in living plants. However,
as is the case with so many tropical species, a large number of Chrysobalanaceae
are known only by the type collections. At this time our knowledge of such species is
obviously only provisional.
Throughout this study of the family, attempts have been made to uncover
additional taxonomic characters in the wood, pollen, leaf anatomy, and vascular
anatomy, but these categories have offered little below the generic level. No reports
of chromosome number have yet been published and my own attempts to obtain
counts thus far have been unsuccessful. Thus it is clear that at the specific level, the
present treatment is based almost entirely on morphological characters. These seem
generally satisfactoryto distinguish the species except in the several species complexes
where experimental studies would no doubt contribute to better understanding.

TAXONOMIC HISTORY

In the first edition of SpeciesPlantarumLinnaeus (i 753) treated just two species:

Chrysobalanus icaco,which was placed in Polyandria Monogynia, and Hirtellaamericana,
which was listed under Triandria Monogynia. Their generic names for purposes of
valid publication (Article I3, note 2 of the I96I edition of the International Codeof
BotanicalNomenclature) are associated with the subsequent descriptions given in the
fifth edition of Linnaeus' GeneraPlantarum(I754), and Hirtellahad been detailed still
earlier in HortusCliffortianus by Linnaeus (1738).
* B. A. Krukoff Curator of Amazonian Botany, The New York Botanical Garden.

1
2 Flora Neotropica

Knowledge of the group was greatly extended by Aublet (I 775) who described
six South American genera, which were later assigned to it by the various authors
mentioned below. Four of Aublet's genera, Acioa, Couepia,Licaniaand Parinariare
still recognized by most botanists, though their contents have varied widely in
different treatments in the past. Moquilea,here regarded as a synonym of Licania,was
separated from the latter genus by Aublet because of his use of the Linnaean sexual
system of classification, and as the result of his apparently having confused the calyx
with the corolla, and the bracteoles with the calyx. Aublet placed Licaniain Pentan-
dria Monogynia; Parinari,Couepiaand Moquileain Icosandria Polygynia; and Acioa
in Monadelphia Dodecandra. The sixth genus, Ferolia,based on a sterile specimen
and Barrere's pre-Linnean description (I74I), was included in a group of plants
imperfectly known. Many subsequent authors have merged it with Parinari,but more
recently it has been shown that Ferolia is synonymous with the Moraceous genus
Brosimum,Pfeiffer (1926), Standley (I929). The nomenclatural difficulties involved
in this synonymy are discussedby Berg (i 968) in a proposal for the conservation of the
name Brosimumover Ferolia.
In 1789, two important works dealing with Chrysobalanaceae were published:
one (Schreber, I789) based on the artificial system of Linnaeus; the other (Jussieu,
1789) laying the foundation for all subsequent natural systems.
In the eighth edition of the GeneraPlantarum,Schreber (1789, I791) corrected
some mistakes of earlier workers, including Aublet's description of Licania,but his
alteration of the names of several genera and, in some cases, their circumscription,led
to much subsequent confusion, both taxonomic and nomenclatural. Schreberchanged
the name of Licaniato Hedycreaand placed it with Hirtellain Pentandria Monogynia.
Parinariwas re-named Petrocarya and placed in Heptandria Monogynia. Chrysobalanus
was transferred to Icosandria Monogynia. Acioa became Acia and was placed in
Monadelphia Dodecandra. Of the remaining genera of Aublet, Couepiawas regarded
as doubtfully distinct from Acia. Moquileaand Feroliawere not even mentioned.
Jussieu (1789) brought all these genera together for the first time as part of the
family Rosaceae, which he divided into seven tribes, two of them including but not
restricted to genera of Chrysobalanaceae as defined herein. One of these tribes,
named Prockiae, was defined, "Germen unicum supernum monostylum. Fructus
unilocularis mono aut polyspermus. Arbores aut frutices, interdum petalis carentes."
It contained Tigarea,Delima,Prockiaand Hirtella,only the last of which is still retained
in Chrysobalanaceae. The other tribe, the Amygdaleae, bore the synopsis, "Germen
unicum supernum monostylum. Nux mono aut disperma, nuda aut saepis drupacea.
Arbores aut frutices." It contained eleven genera: Licania,Grangeria,Chrysobalanus,
Cerasus,Prunus, Armeniaea,Amygdalus,Moquilea,Couepia,Acioa and Parinarium1of
which the first three and the last four are true Chrysobalanaceae. Grangeria, based on
a Commerson specimen from Reunion, was described here for the first time.
Robert Brown (I818) was the first author to give family status to this group
which he named Chrysobalaneae. It should be noted that Brown's circumscription
of the family, especially as to the genera he assigned to it, stands alone among all
subsequent interpretations as the forerunner of the circumscription adopted here
and in Prance (in press). Brown characterized the family by its basal style, erect em-
bryo and ovule, and a tendency to zygomorphy. He listed as component genera:
Chyrsobalanus, Moquilea,Grangeria,Couepia,Acioa,Licania,Hirtella, Thelira,and Parinar-
ium. Thelira,described by Du Petit Thouars (I806) without indications of species, is in
fact a Mascarene Hirtella.
1 A latinized form of the Aublet name, which I regard as an orthographic variant. In this history
Parinari and similar variants of other generic names are spelled in each case in the way used by the
author being discussed.
Introduction 3

De Candolle in his Prodromus(I825) treated the Chrysobalaneae of Robert

Brown as the first tribe of his Rosaceae. He included Chrysobalanus, Moquilea,Couepia,
Acioa, Parinarium,Grangeria,Licania, Thelyraand Hirtella. He also described a new
genus, Lecostemon, from a drawing by Mocifio and Sesse and placed it in Genera
Rosaceis affinia together with the genus Trilepisium.The identity of Lecostemon has
been the subject of much uncertainty, but was finally shown to be congeneric with
SloaneaL. (Tiliaceae) by Standley (I944). It was also fully discussed in Prance
(1968).
During the nineteenth century many South American Chrysobalanaceae were
incorrectly interpreted or erroneously assigned to genera, eg, among many similar
instances, Hoffmannsegg proposed Hirtellaoctandra(in fact a species of Licania)and
his manuscript name was later published by Roemer & Schultes; similarly E. Meyer
(1825) misplaced Licania apetalain Hirtella. Martius (1827) redefined Moquileato
include Couepiaand Acioaalthough he had not seen the type specimens of any of these
genera. Subsequently both he and Zuccarini (Martius & Zuccarini, I832; Zuccarini,
i832) excluded Acioa but continued to regard Moquileaand Couepiaas congeneric,
although in fact they only included species of Couepiain the genus they called Moquilea.
The confusion in their work is largely nomenclatural. Martius and Zuccarini clarified
the distinction between Hirtellaand Couepia(which they mistakenly called Moquilea)
by transferring H. polyandraand H. dodecandra to Couepia.Hirtellawas restricted to
species with less than o stamens and a thin endocarp, an interpretation supported
in my work.
Lindley (I836) accepted the Chrysobalanaceae as a family and placed it be-
tween the Rosaceae and Leguminosae. The genera he included are: Chrysobalanus,
Moquilea,Couepia,Acioa, Parinarium,Grangeria,Licania, Thelyra,Hirtella, Stylobasium
and Prinsepia.He also listed the synonyms known to him. Prinsepiawas laterremovedto
the Prunoideaeby Hooker (1865). Stylobasium, an Australiangenus describedby Desfon-
taines (I819), was long an anomalous member of the Chrysobalanceae, and has
been removed to the Sapindales (Prance. 1965).
Meisner ( 1837) was the first to make a key to the genera of the Chrysobalanaceae
(Chrysobalaneae) which he treated as the first tribe of his Rosaceae. As it included
several misplaced genera, little light was shed on true relationships. His genera are
RoydsiaRoxb. (now placed in the Capparidaceae), TrilepisiumThou. (a genus still of
uncertain position), Lecostemon, Stylobasium,Licania,Parinarium, Chrysobalanus, Grangeria,
BartlingiaBrongn. (= PultenaeaSm., Leguminosae, Papilionatae), Hirtella, Moquilea
sensu Mart., BalantiumDesv. ex Ham. and Thelyra.I have examined the type collec-
tion of Balantiumand find that it is a Parinari.The first four of Meisner's genera were
only doubtfully included. He followed Martius in including Acioaand Couepiaunder
Moquilea.
Endlicher (I840, I842) treated the Chrysobalanaceae (Chrysobalaneae) as a
family in the order Rosiflorae and listed Chrysobalanus, Hirtella, Licania, Moquilea,
Parinarium,Thelyra,Grangeriaand PrinsepiaRoyle as well as three genera dubiae,
Lecostemon, Trilepisiumand Stylobasium. Again Acioaand Couepiawere given as synonyms
of Moquilea.
In an account of the South American genera, Bentham (I840) realized that
generic limits in the Chrysobalanaceae had been previously confused and gave a
summary of the genera known to him: Parinarium,Chrysobalanus, Aloquilea,Couepia,
Grangeria,Hirtellaand Licania.He did not see a good enough specimen of Acioa to
decide its status. He restored Couepiato generic rank, transferredall the post-Aublet
species of Moquileato it. He also discussed the affinities of the Chrysobalanaceaewith
the Leguminosae and the Rosaceae.
The account of the group (as Rosaceae: Chrysobalaneae) by J. D. Hooker
 4 Flora Neotropica

(i865) in the GeneraPlantarumof Bentham and Hooker was much more compre-
hensive than all previous accounts. He included: Chrysobalanus, Licania, Moquilea,
Grangeria, Parinarium,Trichocarya,Parastemon, Hirtella,Couepia,Griffonia,Lecostemon
and
Stylobasium. Prinsepiawas correctly placed in the Prunoideae (Pruneae) in spite of its
basal style. Acioawas still treated as part of Couepia,but a new genus, Griffonia,was
described for an African species of Acioa. Thelyra(Hirtella)was mistakenly reduced to
synonymy under Parinari,by Hooker.
Parastemon had been described by A. de Candolle (i 842) to accommodate a new
Asiatic species (Embeliaurophylla)he had mistakenly placed in Myrsinaceae a few
years previously (1834). De Candolle still harbored doubt about the relationships of
his new genus and tentatively placed it in Olacineae. Trichocarya was published by
Miquel (I855) and based on two Asiatic species which Korthals had recently pub-
lished in other genera as Angelesiasplendens and Diemeniaracemosa.Miquel apparently
had only seen these names on herbarium specimens. In Prance (in press) reasons are
given for placing Angelesiain Licaniaand Diemeniain Parastemon.
Two years later, in his account for Martius's Flora Brasiliensis,J. D. Hooker
(1867) made an outstanding contribution to our knowledge of this family by describ-
ing all the Brazilian species of Chrysobalanus, Licania, Moquilea, Hirtella, Couepia,
Parinariumand Lecostemon known at that time. He again treated Acioa as part of
Couepia.Recent studies summarized in Prance (I968) have shown that the species
referred to Lecostemon by Hooker in fact belong to Rhabdodendron in the unigeneric
family, Rhabdodendraceae.
Baillon (I868) gave reasons for regarding Thelyraas a Hirtella. He also com-
mented on the difficulty of subdividing the Chrysobalanaceae. In his HistoireBaillon
(i 869), included the Chrysobalanaceaein the Rosaceae. He is the first author to have
arranged the genera into groups based on the symmetry of the flower. His "Group a"
contained Chrysobalanus, Licania (including Moquilea), Lecostemonand Stylobasium.
"Group b" comprised Grangeria,Hirtella,Couepia,"Parinari,"Acioa and Parastemon.
He did not see material of Trichocarya. Baillon, following Grisebach (I857), united
Moquilea with Licania,and reduced Griffoniato synonymy under Acioa.Certain errors
notwithstanding, particularly concerning the illustrationsBaillon's views, have been
largely accepted by later workers.
Fritsch (i888), on morphological grounds, considered the Chrysobalanaceae to
be a well defined family, clearly separable from its nearest allies, the Rosaceae and
Leguminosae. He delineated three subfamilies and, by means of a sketchy key,
designated the following generic assignments: (i) Chrysobalaneae including Chryso-
balanus,Licania,Grangeria,Hirtella,Couepia,Acioa,Parinarium,Angelesia,Diemeniaand
Parastemon;(2) Lecostemoneae containing Lecostemon; and (3) Stylobasieae contain-
ing Stylobasium. The following year (1889) he published a valuable conspectus of all
known species of Licania, together with detailed reasons for uniting Licania and
Moquilea.
Shortly afterwards, in I891, Focke provided a review of the entire assemblage
in his account of the Rosaceae for Engler & Prantl's Die NatiirlichenPflanzenfamilien.
Under the subfamily Chrysobalanoideae he included only one tribe which was in
turn divided into two subtribes, Chrysobalaninae and Hirtellinae. The genera
placed in the Chrysobalaninae are Chrysobalanus, Grangeria,Moquilea(kept separate
from Licania),Licania,Lecostemon and Stylobasium.The Hirtellinae included Hirtella,
Couepia,Acioa, Parinarium,Angelesiaand Parastemon.Focke's generic diagnoses were
very brief and his keys to the genera did not employ the most diagnostic characters.
Subsequent work has not supported his subtribal groupings. Lecostemon and Stylo-
basiumwere included with some reservationsand he suggested a relationship for them
with the Phytolaccaceae. At about this time an extremely detailed account of the
Introduction 5

anatomy of the stem and leaf of eleven genera of the Chrysobalanaceae (including not
only Stylobasium and Lecostemon but also Prinsepia)was published by Kuster (I897).
Although Kuster himself drew no conclusions from the very well characterized series
of anatomical features, his work has some interesting taxonomic implications as has
been indicated by Hallier (1903). Hallier suggested the transfer of Stylobasiumto
Prunoideae(Amygdaloideae) somewhere near Prinsepia and Dichotomanthes;that
Moquilea,Licania,and Angelesiashould be united as a single genus and that Lecostemon,
while truly of the Chrysobalanaceae,is isolated within the family. Hallier's groupings
of the genera (excepting the inclusion of Lecostemon) are largely supported by my work.
All widely used generic synonyms are mentioned in the foregoing historical
summary but during the Nineteenth Century many others were published, mostly
by authors with little apparent knowledge of the group; there are at least eleven for
the genus Parinari(sens. lat.) alone. Further details of the American synonymy are
given under appropriate taxa in the systematic treatment.
Since the publication of Die NatiirlichenPflanzenfamilien only three new genera
have been described by authors other than myself. One of these, Geobalanus Small I
consider synonymouswith Licania.In 1905 Engler described Magnistipulafrom Africa.
Most authors since then have united it, at least in part, with Hirtella,but recent work
has shown that it is, in fact, distinct. Another African genus, Afrolicania,was described
by Mildbraed in 1921. It differs from Licaniain certain floral and fruit charactersand
also is distinctive in its secondary xylem.
In I905 Gilg & Pilger described a South American genus, Rhabdodendron which
they placed in the Rutaceae. Subsequently, the species of Lecostemon
as interpreted by
Hooker (1865), by Focke (1891) and by other workerswere transferredto Rhabdoden-
dronby Huber (g9o9b) who believed the original Lecostemon of de Candolle to be a
different plant. The taxonomic history of Lecostemon and of Rhabdodendron and their
relationship to the Tiliaceae and Phytolaccaceae respectively was fully discussed
by Prance (1968).
During the present century several important papers on the floral anatomy of
the Chrysobalanaceae have been published (Bonne, 926a, b, 1928; Juel, 1915;
Morvillez, 1918, etc.).
Recent authors have been more or less equally divided about whether the group
should be treated as a separate family or as a subfamily of Rosaceae. Widely divergent
views on its phylogenetic relationshipshave been expressed, Hailler (i923) suggested
relationship to the Linaceae, Bonne (1926a) to the Tropaeolaceae, Hauman (I95I)
to the Geraniaceae, and Gutzwiller (I96I) to the Sapindaceae. On the basis of floral
structure,anatomy and pollen, I believe that the family belongs between the Rosaceae
and the Leguminosae. Further details are presented in Prance (in press).
In a forthcoming publication entitled A Synopsisof Chrysobalanaceae, Prance (in
press), I present a worldwide revision of the generic limits of the family based on ana-
tomy, pollen structure, blastogeny and morphology. This work considerably changed
the circumscription of the genus, Parinari.Parinarihad been a heterogeneous assort-
ment of species held together by a single unnatural character, the false septum of the
ovary. The difference between species of Parinariwere far greater than those between
long-established genera such as Couepiaand Hirtella.To restore consistency in generic
concept I concluded that Parinarihad to be divided into seven genera. This made
little change in the American genera except for the segregation of five anomalous
species of Parinariinto the new genus Exellodendron. The generic synopsis arising from
this work recognizes the following 17 genera: i. Tribe Chrysobalaneae: Chrysobalanus,
Licania (incl. Geobalanus and Angelesia);Afrolicania;Parastemon;2. Tribe Hirtelleae:
Hirtella, Couepia,Acioa, Magnistipula,Grangeria,Hunga, Parinari and the following
segregates of Parinari: Atuna (Asia), Bafodeya (Africa), Exellodendron(America),
6 Flora Neotropica

Kostermanthus(Asia), Maranthes(Africa & Asia), and Neocarya(Africa). (Hungais a

new genus from New Caledonia.)

ANATOMY

Primary stem and leaf.

In the detailed study of primary stem and leaf anatomy by Kiister (I897), a
description of the family as a whole was presented, as well as of each genus indivi-
dually. This study was based on 76 species of Chrysobalanaceae, and has formed the
basis for all later descriptions of the primary anatomy of the family.
The whole mesophyll consistsof palisade tissue. Hairs are unicellular and simple.
Glands are frequently present at the base and on the undersurface of the leaf. The
epidermis is composed of cells with mucilaginous walls in Chrysobalanus, Hirtellaand
Parinari,but is papillose on the lower surface of some species of Couepiaand on the
upper surface of Licania.The epidermal cells are frequently palisade-like. Hypoderm
is present in most genera. Stomata are confined to the lower surfaceand are rubiaceous.
Fiber-like spicular cells occur in the mesophyll of some species of Couepiaand Licania.
The vascular bundles of the veins are surrounded by sclerenchymatous fibers which
have characteristic U-shaped thickenings in transversesections; the bundles are verti-
cally transcurrent in some species. The petiole in transverse section has an abaxial
closed ring of xylem and phloem which shows various modifications toward the
adaxial side in different genera and species, but the modifications are not well
correlated with taxonomic groupings. Silicified membranes are universal, and silica
bodies are present in the epidermal cells and in the neighborhood of the leaf veins in
all genera except Parastemon;they also occur in the mesophyll of some species of
Couepiaand Licania.
The young stem has cork of superficial origin. The pericycle includes strands of
primary fibers which subsequently become united by stone cells, the latter showing
horseshoe-shaped thickenings in transverse section. The xylem forms a continuous
cylinder transversedby narrow rays. The vessels have simple perforations. Idioblasts
containing tanin are present in phloem of some species of various genera. Silica bodies
occur in the pericycle, medullary rays and pith of all genera except Parastemon.

Secondary xylem (Fig I A-E).

Growth rings are often present, marked by periodic wider spacings of paren-
chyma bands; the rings may be present or absent in different samples of the same
species. Vessels are typically exclusively solitary, but occasionally in radial multiples
ot 2-5 cells, especially in Acioa,and are usually in pairs in Afrolicania;typically they
are obliquely arranged and in most genera are moderately large; the mean tangential
diameter is 200-300 ,p; they are always fewer than 20 per sq mm, and in some species
fewer than 5 per sq mm; they are without spiral thickenings; perforations are ex-
clusively simple; intervascularpitting is alternate, never minute; the ray-vesselpits are
large, of various shapes, and alternate; tyloses are frequent, especially in Magnistipula,
Maranthesand Licania and are occasionally sclerosed in Licania; the mean vessel
length is o.6-o.8 mrm. Parenchyma is exclusively apotracheal and abundant and
usually occurs in numerous fine continuous bands usually i (locally 2) cells wide;
there are 6-i bands per mm; less often the bands are 2-3 cells wide and are less than
6 per mm in Parinarisens. strict.; the bands are obscure in Neocaryaand Afrolicania;in
Afrolicaniathere is a reticulate pattern of uniseriate bands; the parenchyma strands
are up to i6 cells high; gum deposits are frequent. Rays are predominantly uniseriate,
Introduction 7

A B

E H
FIG i. Secondary xylem and pollen of Chrysobalanaceae. A, Licania heteromorpha, X.S. x 8
(PRFw 7I78); B, Hirtella triandra,T.S. x8 (Cooper& Slater 279); C, Couepia glandulosa, T.S.X 32
(Stahel 353); D, Licania tomentosa,T.S. x 8 (FHOw 20926); E, Hirtella triandra, R.S. x 8 (Cooper&
Slater 279). F, Chrysobalanus
icaco, x 380 (Broadway8626); G-H, Exellodendronbarbatum,x 380 (Krukoff
I"37).

exclusively so in Chrysobalanus, Exellodendron,Grangeria,Hirtella and Licaniaand with

some biseriate rays in all other genera except Afrolicaniain which they are predomi-
nantly biseriate; rays are never more than 2-seriate and are commonily more than
i mm high in Couepia,Parastemon,Parinari, Maranthes,Atuna and Afrolicania; in
Afrolicaniathey are higher than in any others and are commonly over 2 mm high;
they are usually heterogeneous (Kribs Type III) but almost homogeneous in Para-
stemon;gum deposits are abundant; the pits to axial parenchyma are large. Silica
inclusions are present in the ray cells of many species. Fibers are usually thick-walled,
but somewhat thinner in Parinariand Neocaryaand in some species of Licania; they
8 Flora Neotropica

have numerous distinctly bordered pits on the tangential walls, which are more
numerous and often biseriate in wider cells bordering vessels; pits on the radial walls
are almost entirely limited to areas in contact with the rays; the mean length is
1.3-I.4 mm.
The wood is moderately to very hard and heavy. Specific gravity is 0.75-I.10.
It is hard to saw owing to abundant silica inclusions in the ray cells in all species, and
in the parenchyma of a few species. The silica content has been summarized in detail
by Amos (1952).
The uniformity of the xylem yields few generic or specific charcters. In the
taxonomy of the group, the anatomy of the secondary xylem is most useful to define
the limits of the family, and as supportingevidence for separatingit fromthe Rosaceae.

POLLEN

The pollen grains have 3 furrows without any special features except occasional
equatorial constrictions; some species have grains with 3 and other grains with 4
furrows; there are no distinct pores; the grains are usually distinctly triangular in
shape in polar view, except when 4-furrowed; they are elliptical to circular in equator-
ial view and are oblate-spheroidal, prolate-spheroidal or subprolate in shape as in-
dicated by the ratio- i00 polar length : the equatorial length =85-150. The size is
very variable from one genus to another but is rather less variable among species of
each genus; the polar area is usually small, sometimes medium, but never large. The
exine is medium to rather thick with very little patterning on the walls; it is usually
scabrous to verrucose, but never striate.
Chrysobalanaceae pollen is very uniform, and, as with the anatomical features,
is most useful to define the family limits.
Pollen of Chrysobalanaceae is easily distinguishable from that of the Rosaceae
but is also similar to it. It is different in the markedly triangular shape in polar view
in the expanded grain. The grains in the Rosaceae are rounder to weakly triangular,
usually with more distinctive pores, and often with more patterning on the walls;
they frequently have a distinct wedge-shaped protusion from the middle of the
furrow which is obvious in polar view. The pollen of Chrysobalanaceaedoes not have
such a protusion from the furrows, Erdtman (I952) states, "pollen morphological
objections cannot be raised against regarding the Chrysobalanaceae as a separate
family." The pollen of Chrysobalanaceae is illustrated in Fig i, F-H.

THE RANK AND POSITION OF THE CHRYSOBALANACEAE

Robert Brown (i818), who first described the group as a family, has been
followed in this view by: Bartling (1830), Zuccarini (I832), Lindley (i836), Endlicher
(1840), Bentham (1840), Walpers (1843), Miquel (i 855), Agardh (I858), Warming
and Mobius (i 9 I), Wettstein (I 935), Exell (I 944), Gundersen (I950), Dandy
(1960), Melchior (1964), Takhtajan (1966), Cronquist (1968). All of these authors
have placed the family next to the Rosaceae. However, the authors of the most widely
used systems of classificationshave usually treated the group as a tribe or subfamily of
Rosaceae, eg, de Candolle (I825), Hooker (i865) in Bentham and Hooker, Focke
(1 89 I) in Engler and Prantl and Hutchinson (1926, 1959).
Introduction 9

Fritsch (i888), in a careful study of the group, regarded it as a family between

Rosaceae and Leguminosae, and his views have been reiterated by many subsequent
workers. Wettstein (i933) placed the Chrysobalanaceae in this position, and com-
mented that one of his reasonsfor regarding it as a family was the distinctive anatomy.
Hallier (1903) concluded from Kuister's(I897) detailed account of the anatomy
of the leaf and primary stem that the group should be separated from the Rosaceae
as a family. Juel (i 915), who correctly interpreted the ovary of the Chrysobalanaceae
for the first time, considered it to be so different from that of the Rosaceae that the
group should be elevated to family rank, but remain near Rosaceae. On the basis of
differences he discovered in leaf-trace anatomy, Morvillez (I918) reached a similar
conclusion. Bonne (i928) made a detailed study of the pedicel and floral vascular
anatomy, concluding that the Chrysobalanaceaeare quite distinct from true Rosaceae.
Both Metcalfe and Chalk (I950) and Erdtman (I952) imply that no objections could
be raised against treating the group as a family on the basis of wood anatomy and
pollen-grain structure, respectively. Details of the most important anatomical
differences between Chrysobalanaceae and Rosaceae are summarized in Table I.
There are also several characterscommon to all, or most, Chrysobalanaceae but onlv
of sporadic occurence in Rosaceae.
The characters tabulated (Table I) clearly show that the Chrysobalanaceae,
although sufficiently distinct from Rosaceae to be treated as a separate family, is
nevertheless still closely related to it.
Various phylogenists during the present century have suggested diverse relation-
ships for the Chrysobalanaceaewith other families. Hallier (I 923) united the Chryso-
balanaceae, Dichapetalaceae and Trigoniaceae to form a single family, which he
suggested is derived from the Linaceae and Polygalaceae. In 1915, Juel, after point-
ing out the trimerous nature of the ovary of the Chrysobalanaceae, compared its
essentially gynobasic2 structure with Limnanthes but he did not suggest a relationship
between the two families. Bonne (I 926a) appears to have misread Juel's paper, for
she suggested a phylogenetic connection between Chrysobalanaceae and Limnan-
thaceae or Tropaeolaceae. Hauman (I 951) proposeda relationshipwith Tropaeolaceae
and Geraniaceae. Gutzwiller (I961), whose knowledge was based chiefly on Stylo-
basium(now excluded from the family), Parinariand Chrysobalanus, suggested that the
Chrysobalanaceae, along with the Connaraceae, should be transferredto the Sapin-
dales, and placed next to the Sapindaceae.
Wood structure, anatomy, pollen and morphology do not confirm a close rela-
tionship with any of the above families. There are no grounds for removing the
Chrysobalanaceaefrom the order Rosales. It stands as a family between the Rosaceae
and the Leguminosae. This is further discussed by Prance (in press).

Table I
Summary of differences between Chrysobalanaceae and Rosaceae

Chrysobalanaceae Rosaceae

Primarystemand leaf anatomy

(I) silica deposits abundant and occurring (i) silica absent
in five different forms
(2) stone cells in the pericycle thickened on (2) stone cells in the pericycle not so
only one side, thus appearing U-shaped thickened
in transverse section
(3) stomata rubiaceous (3) stomata ranunculaceous

Gynobasic is used here and in other places in this work to refer to the style which is attached by
2
its base to receptacular tissue below the ovary.
10 Flora Neotropica

(4) petiole with an abaxial closed ring of (4) petiole various but not as in
xylem and phloem which is variously Chrysobalanaceae
modified on the abaxial side

PedicelAnatomy
(i) pedicel articulate at an active (i) pedicel inarticulate or if articulate
meristematic zone without an active meristem

WoodAnatomy
(I) vessels oblique, infrequent, usually (i) vessels not oblique, abundant,
200-300 /e diam usually less than Io00 diam

(2) parenchyma banded, composed of (2) parenchyma sparse, not banded,

long strands composed of short strands

(3) rays predominantly uniseriate (3) rays multiseriate

(4) intervascular pitting large (4) intervascular pitting never large,

usually minute

(5) silica abundant in the rays (5) silica absent

Pollen
(1) pollen markedly triangular in (i) pollen usually not triangular or only
expanded grain weakly triangular

(2) always without a protrusion in the (2) often with a wedge-shaped protrusion
mouth of the furrow in the mouth of the furrow

(3) grain scarcely patterned (3) grain often distinctly patterned

Floral anatomy
(I) ovary basically of 3 carpels with a (I) style never gynobasic
gynobasic style

(2) ovule erect (2) ovule pendulous

(3) flowers at least slightly zygomorphic in (3) flowers actinomorphic

all genera except Chrysobalanus

ACKNOWLEDGMENTS

First I would especially like to thank Dr. Bassett Maguire, who encouraged me
to undertake this monograph of the neotropical Chrysobalanaceae. Dr. Maguire, for
many years interested in the family, had accumulated much material and informa-
tion. When I came to New York in I963, he generously handed over to me all his
material and information and has since been a continual source of encouragement.
I am also grateful to Dr. William C. Steere and the staff of The New York
Botanical Garden for their support of this work. I am grateful to the National
Science Foundation for the grants GB-464I, and GB-7356, to support field work
which has been a vital part of this study. I should like to thank the directors of the
numerous herbaria consulted, for loans and facilities when visiting them. These
herbaria are too many to list but they are all cited in the systematic part of this work.
I am particularly grateful to the herbaria that provided me facilities while I was in the
field. In this regard warm thanks are due to the following individuals; Dr. Djalma
Batista, until recently Director of the Instituto Nacional de Pesquisas da Amazonia,
Introduction 11

Manaus, Brazil; Dr. William Rodrigues, Chief of the Botanical Section of the same
Institution; Dr. J. Mur9a Pires of the Instituto Agronomico do Norte, Belem, Para,
Brazil; Dr. J. P. Schulz of the Forest Service, Paramaribo, Suriname; Prof. J.
Purseglove, formerly Professor of Botany at the University of the West Indies,
Trinidad; and Dr. T. Lasser, Director of the Instituto Botanico, Caracas, Venezuela.
I am especially grateful to those people who have done much to help me during
the final stages of the preparation of this manuscript, to Dr. Rogers McVaugh who
made many helpful suggestions, to Dr. Tetsuo Koyama who checked the Latin
descriptions, to Mr. Enrique Forero who helped in many ways, particularly by
checking the Spanish names, and most of all to Dr. Howard S. Irwin Jr. for the many
hours which he devoted to editing this manuscript, and who gave me much help and
encouragement throughout this study.
I would like to thank Mr. Charles C. ClareJr., Mrs. Julia Loken and Mr. David
Woolcott for preparation of all the line drawings and maps in this work. I am grateful
to Mr. Frank White, Curator of the Forest Herbarium, Oxford, for much advice and
encouragement. I should like to thank my wife for her valuable help with many
aspects of this work.
 SYSTEMATIC TREATMENT

In addition to the general introduction, a few comments about the systematic

section are necessary to clarify the descriptions.
Leaf size measurementsare taken from fertile branches, and are often much larger
on sterile, sucker, and shoot material, but since such material has been collected in
only a few species it is better to present only measurements from fertile branches.
Flower size has been measured from the tip of the largest calyx lobe to the articula-
tion of the pedicel unless otherwise stated. In the species description the surfaceof the
midrib is treated separately from the description of the leaf surface. In most descrip-
tions of the fruit the epicarp, mesocarp, and endocarp are described separately, but
in some cases there is no clear-cut differentiation into these layers, and the term
pericarp is used to include the entire casing of the fruit.
In the two largest genera, Hirtellaand Licania,there is naturally greater mor-
phological variation and a greater number of characters are involved, consequently
a discussion and definitions of some terms used are given before the keys to these
genera.
All herbarium material examined of most species has been cited below the species
descriptions. In the case of extremely common species, only selected specimens are
cited in the text, and the rest are all included in the list of exsiccatae at the end. In
these cases the list of specimen citations in the text is preceded by "Representative
collections."
Local names have been taken both from the literature and from herbarium
specimens.
Abbreviations used:
cult =-cultivated
fl = flowering
fr = fruiting
frag =-fragment
st = sterile
Herbaria abbreviations are the standard ones from Index Herbariorum.One
additional herbarium, not cited in the index, has been consulted; ie HST = Herbario
Sergio Tavares, of Sudene, Recife, Brazil.
The specimens cited in quotes are dubious records, and probably do not re-
present the true distribution of the species. This mainly refers to specimens pirated
by Glaziou.

CHRYSOBALANACEAE R. BROWN. Chrysobalaneae (family) R. Br. in Tuckey,

Narrat. Exp. Zaire Congo. 433. I818; Bartling, Ord. Nat. P1. 405-406. I830;
Endl., Gen. P1. I251. I840; Suppl. 2: 95. I942; Martius & Zuccarini, Abh.
Akad. Miinchen I: 372-391. I832; Zuccarini, Flora 15(Beibl.2): 78-93. I832;
C. Muell. in Walp. Ann. Bot. Syst. 4: 640-49. I858; Griseb., Fl. Brit. W. Ind.
229-231. I864.
ChrysobalanaceaeLindley, Nat. Syst. Bot. ed. 2. 158. i836; Benth. in Jour. Bot. Hooker a:
210-23. I840; Fritschin Verh. Zool.-Bot. Ges. Wien 38: 93. i888; Warming& Mobius,

12
Chrysobalanus 13

Handb. Syst. Bot. ed. 3. 393. I9II; Wettstein, Handb. Syst. Bot. ed. 4. i: 771. 1933;
Gundersen, Fam. Dicot. 109. 1950; Melchior in Engl., Syllab. Pflanzenfam. ed. 12. 2: 219.
I964; Takhtajan, Systema Phyl. Magnoliophytorum 267-268. 1966; Cronquist, Evol. &
Classif. Fl. PI. 235-236. I968; Prance, Synopsis of Chrysobalanaceae (in press). Rosaceae
Chrysobalaneae de Candolle, Prodr. 2: 525-29. I825; Meisner, P1. Vasc. Gen. i: IOI-IO3.
1837; Hook. f. in Benth. & Hook., Gen. P1. 1(2): 607-609. I865; Hook. f., Mart. Fl. Bras.
14(2): 5-56. I867; Hallier in Abh. Naturh. Ver. Hamburg I8: 1-98. 1903. Rosaceae
Chrysobalanoideae Focke in Engl. & Prantl, Nat. Pflanzenfam. 3(3): 53. 1894; Fawcett &
Rendle, Fl. Jamaica 3: 253-257. I914; MacBride, Fl. Peru 1070-83. 1938; Kleinhoonte in
Pulle, Fl. Suriname 2(I): 426-56. 1939; Hauman in Bull. Jard. Bot. Brux. 21: 167-98.
I95I; Lem6e, Fl. Guyane Fran;. 2: I6-30. 1952. Amygdalaceae sensu Standl., Trees &
Shrubs of Mexico 343-45. I 920. Rosaceae tribe Prockieae pro parte and tribe Amygdaleae
pro parte-Jussieu, Gen. P1. 340-342. I789. Pre-Jussieureferencesto thefamily: Aublet, PI.
Guiane I: 119, 514-22; 2: 698. 1775. Schreb., Linn., Gen. P1. ed. 8. 153, I6o, 245, 336,
458, I 789.

Flowers actinomorphic to zygomorphic, hermaphrodite or rarely polygamous,

markedly perigynous. Receptacle of varied shape and size, short to elongate, often
gibbous at the base; lobes 5, imbricate, often unequal, erect or reflexed. Disc always
present, forming a lining to the receptacle. Petals (4-)5, sometimes absent, inserted
on the margin of the disc, commonly unequal, imbricate, deciduous, rarely dis-
tinctly unguiculate. Stamens indefinite, from 2-IOO (or to 300 in one species of
Couepia),inserted on the margin of the disc, in a complete circle or unilateral, all
fertile or some without anthers and often reduced to small staminodes; filaments
filiform, free, connate or ligulately connate, very short to elongate, included to far
exserted; anthers small, 2-locular, longitudinally dehiscent, glabrous or rarely
slightly hairy. Ovary basically of three carpels but usually with only one developed,
the other 2 aborted or vestigial, variously attached to (the base, middle or mouth of)
receptacle, sessile or with a short gynophore, pubescent or villous, each carpel uni-
locular with two ovules, or bilocular (owing to a false partition) with one ovule in
each locule. Ovules erect, with micropyle at the base. Style filiform, basally attached,
stigma 3 lobed or truncate. Fruit a dry or fleshy drupe of varied size, interior often
densely hairy; endocarp much varied, thick or thin, fibrous or bony, often with a
special mechanism for seedling escape. Seed erect, exalbuminous, the testa mem-
braneous; cotyledons amygdaloid, plano-convex, fleshy, sometimes ruminate. Ger-
mination hypogeal with the first leaves opposite or alternate, in all American genera
except Maranthes(epigeal with opposite first leaves).
Trees, shrubs or suffrutices. Leaves simple, entire, alternate, often coriaceous,
glabrous or with an indumentum on undersurface;petioles often with 2 lateral glands.
Stipules 2, minute to large (to 5.0 cm long), caducous or persistent, of varied shape.
Inflorescence racemose, paniculate or cymose; flowers bracteate and usually bibrac-
teolate.
About 420 species in I7 genera distributed throughout the lowland tropical
regions of both hemispheres.
TYPE GENUS.ChrysobalanusLinnaeus [derived from the Greek words Chrysos-
(golden) and balanos- (fruit or acorn)].

Key to American Genera of Chrysobalanaceae

i. Ovary inserted at or near base of receptacle, flowers actinomorphic.
Tribe i. Chrysobalaneae.
2. Stamens exserted and connate in groups, with hairy filaments; endocarp longitudinally
ridged; inflorescence cymose or a raceme of cymules. i. Chrysobalanus.
2. Stamens included or exserted and with glabrous free filaments; endocarp seldom ridged;
inflorescence usually a racemose panicle rarely a raceme or raceme of cymules or cymose.
2. Licania.
14 Flora Neotropica

i. Ovary inserted at or near mouth of receptacle; flowers zygomorphic. Tribe 2. Hirtelleae.

3. Carpels bilocular, fruit bilocular often with one loculus only developing.
4. Stamens included, 6-8 fertile; receptacle hollow, interior pubescent.
5. Leaf underside with stomatal cavities which are filled by lanate pubescence;
endocarp thick with a rough surface and with two basal stoppers, fruit exterior
verrucose. 3. Parinari.
5. Leaf underside glabrous or lanate-pubescent but no cavities; endocarp thin, with a
smooth surface without basal stoppers, fruit exterior smooth. 4. Exellodendron.
4. Stamens exserted, I8-6o fertile; receptacle solid, interior glabrous. 5. Maranthes.
3. Carpels and fruit unilocular.
6. Staminal filaments free almost to base, not united into a ligule.
7. Stamens (I o-) I4-300; fruit with no lines of dehiscence; endocarp thick and often
fragile. 6. Couepia.
7. Stamens 3-10; fruit opening by longitudinal lines of weakness; endocarp hard
and thin. 7. Hirtella.
6. Staminal filaments united into a ligule for the greater part of their length. 8. Acioa.

Tribe i. Chrysobalaneae, see Prance, Synopsis, in press (Genera I & 2).

Ovary inserted at or near the base of the receptacle.

i. Chrysobalanus Linnaeus, (Gen. P1. ed. I. 365. 1737) Sp. P1. 514. 1753;
Gen. P1. ed. 5. 229. I754; Browne, Nat. Hist. Jamaica. 250, t. I58. I756;
Plum., P1. Am. 151, t. 158. 1755; Jacq., Sel. Stirp. Am. Hist. 155, t. 94-.
1763; DC., Prodr. 2: 525. I825; Tussac, Fl. Antill. 91, t. 31. Meissn.,
Gen. P1. IO02.I837; Endl., Gen. P1. 6407. I840; Griseb., Fl. Brit. W. Ind.
229. I86o; Hook. f. in Benth. & Hook., Gen. P1. i: 606. I865; Hook. f.,
Mart. Fl. Bras. 14(2): 7. I867; Focke in Engl. & Prantl, Nat. Pflanzenfam.
3(3): 57. I894; Fawcett & Rendle, Fl. Jamaica 3: 253. I914; Kleinh. in Pulle,
Fl. Suriname 2(I): 429. 1939; Howard, Jour. Arnold Arb. 45: 279. 1964;
Prance, Synopsis (in press), Jour. Arnold Arb. 51: 523-526. I970.
IcacoPlumier, Nov. P1. Am. Gen. 43. 1703 nom illegit.

Small trees and shrubs with hermaphrodite flowers. Leaf underside glabrous or
with a few stiff appressed hairs, the stomata visible at x 25, with two glands (some-
times obscure) at base of lamina. Bracts and bracteoles small, eglandular. Inflores-
cences terminal or axillary cymules. Receptacle cupuliform, the interior and exterior
puberulous. Calyx lobes 5, acute. Petals 5, longer than the calyx lobes. Stamens 12-26,
sometimes shorter than the others, arranged in a complete or nearly complete circle;
filaments hairy, united at the base for a short part of their length, about twice as
long as calyx lobes. Ovary inserted at base of the receptacle, densely pilose; carpel
unilocular; ovules 2. Style pubescent. Fruit a small fleshy drupe; epicarp smooth,
ridged; endocarp hard, thin, glabrous on interior, smooth on exterior with 4-8
prominent longitudinal ridges which correspond to the lines of fracture that allow
the seedling to escape.
TYPE SPECIES.Chrysobalanusicaco Linnaeus.
DISTRIBUTION. Tropical Africa and America, mainly in coastal regions. Natural-
ized in Asia and Pacific Islands. In America from Florida through Central America,
to southern Brazil and the West Indies.

Taxonomic History of Chrysobalanus

Chrysobalanuswas validly published by Linnaeus in the I753 edition of Species

Plantarum, although it had appeared in the literature prior to that date (Linnaeus,
Chrysobalanus 15

1737). The specific name C. icacowas taken from Plumier (I 703). Linnaeus also cited
several other earlier names, some of which are certainly not referable to Chrysobalanus,
such as Myrobalanus of Sloane and Condondum of Rumphius. If it were not for the type
specimens in the Linnaean herbarium, Chrysobalanus would be difficult to typify
because of the different elements cited by Linnaeus. However, his conception of the
species was obviously based on the herbarium material which is now in the Linnaean
Herbarium, and it correspondswith Plumier's (I 703) description and (I755) illustra-
tion. The Linnaean herbarium contains three sheets of C. icaco. One sheet was
collected by Patrick Browne from Jamaica, and bears Linnaeus' script, therefore,
must be regarded as the holotype. The other two sheets are from Suriname, but the
collector is not identified. Linnaeus gave the distribution of ChrysobalanusasJamaica,
Brazil and Suriname.
Since the time of Linnaeus Chrysobalanus has been cited and dealt with in numer-
ous publications, without any alteration to the generic circumscription. It has also
been beautifully illustrated several times, eg by Tussac (I827). Some of these refer-
ences are given here above the generic and specific descriptions.
Chrysobalanus was apparently firstrecognized in Africa by Solander when annotat-
ing material in the Banks Herbarium as Chrysobalanus ellipticus.Solander did not
publish this, but his name was published by Sabine (I824). In the same publication
Sabine published C. luteus,another new African species. Since the latter species was
based only on a description given to Sabine by Don, and since there is no herbarium
material, it will always remain a dubious species. C. ellipticus,however, is a Chryso-
balanus.Shortly after the publication of Sabine, Schumacher (I827) published C.
orbicularisfrom Liberia. This species is conspecific with the Linnaean species C. icaco
from the New World
More recently several other species of Chrysobalanus have been described both
from America and Africa. All the subsequent species from America, except C.
cuspidatus,belong within the polymorphic species C. icaco.This genus has been little
confused with other genera. However, its closeness to Licaniawas suggested by the
opinions of Gomez de la Maza (I887) and Morales (I887), who initiated the merging
of Licaniawith Chrysobalanus by making some of the new combinations. The position of
Chrysobalanus, and its relationship with Licaniawill be more fully discussed (Prance,
in press).

Key to American Species of Chrysobalanus

i. Leaves orbicular to ovate-elliptic, apex retuse to rounded or minutely and bluntly acuminate;
midrib glabrous beneath. I. C. icaco.
i. Leaves elliptic to oblong, apex with a distinct acumen 5.0-I2.0 mm long; midrib sparsely
hirsute beneath. 2. C. cuspidatus.

i. Chrysobalanus icaco Linnaeus, Sp. P1. I: 513. 1753; Plum., P1. Am. 15I,
t. I58. I755; Browne, Nat. Hist. Jamaica. 250, t. I5(5). I756; Jacq. Sel.
Stirp. Am. Hist. I55, t. 94. I763; DC., Prodr. 2: 525. I825; Tussac, Fl.
Antill. 91, t. 31. 1827; Griseb., Fl. Brit. W. Ind. 229. I864; Hook. f., Mart.
Fl. Bras. 14(2): 7. I867; Fawcett & Rendle, Fl. Jamaica 3: 253. I914;
Kleinh. in Pulle, Fl. Suriname 2(I): 429. I939. Fig 2.
Prunusicaco Labat, Itin. Am. 3: 30. I722 nom illegit.
C. pellocarpusG. F. W. Meyer, Prim. Fl. Esseq. I93. I818. Type. Herb. Meyer5r, Guyana, fl
(holotype, GOET).
C. purpureusMiller, Gard. Dict. ed. 8 (2). I768.
16 Flora Neotropica

C. orbicularisSchumacher in Schum., Beskr. Guin. P1. 232. I827.

C. icacovar pellocarpus(G. F. W. Meyer) Hooker f., Mart. Fl. Bras. 14(2): 7. i867.
C. icacovar ellipticus(Solander ex Sabine) Hooker. f., Mart. Fl. Bras. 14(2): 7. I867.
C. savannarum Britton, Bull. Torrey Club 48: 33I. 1922. Type. Britton, Hazen & Freeman 2005,
Trinidad, fl fr (holotype, NY).
C. ellipticusSolander ex Sabine, Trans. Hort. Soc. London 5: 453. I824.
C. icacovar genuinusStehle & Quentin, Fl. Guad. & Dep. & Mart. 2(3): 48. I949.

Shrub or small tree to 5 m tall, the branches glabrous and lenticellate. Leaves
orbicular to ovate-elliptic, 2.0-8.0 cm long, 1.2-6.0 cm broad, retuse or rounded or
with a short blunt acumen not exceeding 2.0 mm at apex, subcuneate at base,
glabrous on both surfaces; primary veins inconspicuous; petioles 2.0-4.0 mm long.
Stipules deciduous, I.O0-3.0mm long. Inflorescences small terminal and axillary,
cymules with gray-brown tomentum. Receptacle cupuliform, tomentose on interior
and exterior. Calyx lobes 5, rounded to acute. Petals exceeding calyx lobes, white,
glabrous. Stamens I2-26; filaments joined for part of their length in small groups,
densely hairy. Ovary pilose. Fruit ovate to obovate, 1.8-5.0 cm long; epicarp smooth
with longitudinal ridges; mesocarp thin and fleshy; endocarp thin and hard with
smooth, ridged exterior.
TYPE. PatrickBrownesn, Jamaica, In Linnaeus Herb. No. 64I, fl (holotype,
LINN)
DISTRIBUTION.Mainly in coastal areas on sandy dunes, beaches, scrub or thickets,
or less frequently inland in sandy savannas or open river banks, ranging from Mexico
through Central America on both coasts, Florida, throughout the West Indies and
Bahamas, around the northern and eastern coast of South America from Colombia
to southern Brazil. Also distributed in the coastal regions of West Africa from Guinea
to Angola. Cultivated and naturalized in the Seychelles and Fiji. In America flower-
ing and fruiting continuously.
Representative collections. U.S.A. Florida: Black 49-4371 fl (NY); Blodgettsn, fl (BM, GH);
Britton53 fl (F, NY); I23 fr (F, NY); I24 fl (NY); Burgess718 fl (NY); Curtiss728 fl fr (A, F, GH, K,
NY, P, US); 5845 fl (A, GH, K, NY, P, US); Eaton 1103 fl fr (ECON, F, K); Garbersn fl fr (F, NY,
P, US); Hasslersn fl (NY); Hitchcock87 fl (F, GH, NY, US); Le Roy sn fl (NY); Moldenke751 fl (K,
NY, US); 87I fl (K, MO, NY); 5442 st (NY); Pollard & Collins 247 fl (BM, F, NY); Rehder822 fl
(A, K); Rhodes 8427 fl (FHO, US); J7.K. Small 2106 fr (NY); 2110 fl (NY); 4130 fl fr (NY); 7244 fr
(GH, NY, US); J. K. Small et al. 1315 fl (F, NY); I448 fr (NY); 1449 fl (NY); 3166 fr (NY); 3312
fl (NY); Small and Nash 13 fl (NY); 129 st (NY); Tracy 7540 fl (BM, F, GH, NY, US); 924I fl fr (F,
GH, NY, US). MEXICO. Tamaulipas: Palmer i73 fr (A, F, GH, MEXU, MICH, MO, NY, US);
485 fr (BM, F, GH, K, NY, US); Pringle 6629 fl (A, BM, BR, F, GH, K, MEXU, MICH, MO, NY,
P, US); Le Sueur553 fl (F, GH, US). Oaxaca: Conzatti3262 fl (US); 4402 st (US); 4473 fr (US);
ElmoreD I5 fl (F, MICH, US); Nelson 2604 fr (GH, US); Seler 1782 fr (GH). Tabasco: Barlow 7/3C
fl (MEXU); 30o/56Afl (MEXU); Matuda3132 fl (A, F, MEXU, MICH, NY). Chiapas: Purpus342 fl
(US); I0512 fl fr (NY, US). Campeche: SteereI900 fl (F, MICH); 3398 fl (MEXU). Yucatan: Goldman
615 fl (US); 6i6 fl (US); Lindensn fl (P); Miranda 8126 fl (MEXU). Veracruz: Palmer 469 fr (US);
Rotvirosa729 fl (K, US). Without precise locality: J7urgensensn fr (GH); Sesse & Mociio 2129 fl (F).
GUATEMALA. Bernouilli854 fl (NY); Friedrichsthal sn fl (K, NY); Hatch & Wilsonsn fl (F); Heyde &
Lux 3959 fl (MO, NY, US); Johnson1167 fr (LA, US); 7. D. Smith 1481 fr (GH); 1804 fl (GH, K, NY,
US); Standley 73038 st (F); Steyermark 34529 fl (F); 37783 fl (ECON, F); 38717 fl (F); 39432 fl (F);
39606 fl (F); Wilson39I fl (F). BRITISH HONDURAS. Bartlett 112z8 st (MICH, NY, US); Gentle
1248 fl fr (A, F, K, MO, NY); Hunt354 fl (BM, US); Karling I9 fl (F, K); Kluge 13 st (FHO); Lundell
620 fr (A, F, GH, K, MO); 38I7 st (MICH, NY); 4352 fl (F, MICH); Peck21 fl (GH, K); I74 fl (K);
367 fl (GH, K); 368 fl (GH, K); Schipp65 fl (A, BM, F, GH, K, MICH, MO, US); 932 fr (A, BM, F,
GH, K, MICH, MO, NY). HONDURAS. Carleton629 fl (A, US); Mitchell3 fl (GH); Peterssn fl (F);
Standley53007 fl (A); 53997 fl (F); L. 0. Williams & Molina R. 14465 fr (F); Wilson 14 fl (US); runcker
4673 fl (A, F, MICH). EL SALVADOR. Calderon 623 fr (GH, US); Carlson 1053 fl fr (F); 1122 fl (F);
Standley19492 fr (GH, US); 22023 fl (FHO, GH, MO, US). NICARAGUA. C. F. Baker2147 fr (A.
ECON, GH, K, US); Garnier41z6 fl (A, F); Greenman 5739 fl (MO); Lety 62 fl (P); Molina R. 1819 fl
(F); 14790 fl (F); Shimek& C. L. Smith95 fl (F); C. L. Smith i fl (F, GH, MEXU, MICH, US); Standley
11284 fl (F); COSTA RICA. Birkley 2030 fl fr (US); Brenes I2250 st (A, F, NY); J7imenez 2848 fl (F);
 Chrysobalanus 17

Chrysobalanus in America

T '~. .I..~ ... .

U' _. ....'~~~~~~~~~~~~~~~~?:--..'.

.... ..'

?=~~~~~~~~~~~~~~~~~.
....oa/nu .....
~~=Chrysoba/anus cuspidatus~~~~~~~~~~~~~.

'~au
FI
cc,x025
.Gorpi ,Crsblns
iTrbto n
upEts
aito .5
la hp Crsbln. - HV ry-

* = Chrysoba/lanus icaco i-
= Chrysoba/anus cuspidafus

FIG 2. Geographic distribution and variation in leaf shape of Chrysobalanus.

A-F, H-V, Chryso-
balanusicaco, x o.25. G, Chrysobalanus
cuspidatus,x 0.25.
18 Flora Neotropica

Rowles I20 fr (US); Shank & Molina R. 4344 fr (GH); Tonduz 7022 fl fr (BM, GH, K, US); 7023 fr
(BM, F, GH, K, MO, NY, US); 10004 fr (BM, BR, P, US); 10023 fr (BM, P); ro403 fl (K, NY);
Weber6218 fl (GH). PANAMA. Elmore645 fl fr (F, MICH); Erlanson78 fl (GH, US); Fendler 0o7
fl (BM, K, OXF, P, US); Hinds sn fl (K); Johnston292 fr (GH); 802 fr (GH); 938 fr (GH, MO);
Seemannsn fl (BM, GOET); von Wedel2936 fl (F); R. S. Williams 400 fl (US). BERMUDA. Brown,
Britton & Worthy1639 fl fr (NY, Cult.). CUBA. Britton2135 fl (NY, US); Britton,Earle et al 6504 fl
fr (NY); Io309 fl (K, NY, US); Combs436 fl (F, GH, K, P, NY); Curtiss280 fl fr (A, BM, F, GH,
K, NY, P, US); Earle 702 fl (F, NY, US); Ekman 13330 fl (NY); Howard5597 fr (GH, NY, US);
6017 fr (GH, NY); Jack 8776 fl (P); Jennings9 fl fr (GH, NY, US); Killip 42611 fl (BR, K, US); 42884
fl (GH, NY, US); Leon &Cazanas5908 fr (NY, P); Pollard & Palmer I96fl fr (F, GH, NY, US); Sagra
sn fl (P); Shafer378 fl (NY, US); 710 fl (F, NY, US); 715 fl fr (GH, NY, US); 1078 fl (GH, NY, US);
o079ga ffr (F, NY, US); 3995 fl (F, NY, US); 7720 fr (GH, NY); 8302 fr (GH, K, NY, US); 10772 fl
(F, NY, US); iroi8 fr (NY, US); 11759 fr (F, GH, NY, US); VanHermannsn fr (NY); Wright 1607
fl (BM, GH, K, NY, P, US). BAHAMAS. Baker9 fl (K); Brace 4569 st (F, NY); 6980 fr (F, NY);
X. L. Britton89 st (NY); E. F. Britton6587 fl fr (NY); N. L. Britton& Brace528 fl fr (K, NY); Britton &
Millspaugh2516 fl fr (F, NY, US); 2709 st (F, NY); Coopersn fl (NY); Degener18770 fl fr (NY); 18839
fl fr (NY); Eggers 4060 fl (BM, K); Howard 10087 fl (NY, US); Millspaugh 2175 fl (F); 2280 fl (F);
9202 fl (NY); Nash & Taylor 1302 fr (F, NY); NorthropIr5 fl (A, F, GH, NY); Richardsn fl (P); Small
& Carter8760 fl (F, K, NY, US); Wight 6i fl (F, GH, K, NY). CAYMAN ISLANDS. Kings 355 st
(BM); Millspaugh 1265 fl (F); Proctor15223 fl (GH); Rothrock228 fl fr (F); JAMAICA. Harris 6721 fl
fr (F, NY); 8515 fl fr (A, BM, NY); 11166 fl fr (BM, F, K, NY); Marsh 299 fl (K); McGrath8521 fr
(A, NY); Metcalfesn fl (BM); Prior sn fl (K); Wight 174 fl (F, NY). HISPANIOLA. Dominican Re-
public: Abbott2213 fl (F, GH, US); 2863 fr (GH, US); 2882 fl (US); Augusto595 fl (NY); 634 fl fr
(NY); 638 fl (NY); Eggers 1640 fl (GOET, NY, US); 2456 fl (A, BR, P, US); Ekman5808 fl (K, S);
12205 fl (A, US); 12477 fl fr (K, S, US); 15775 fl (A); Fuertes191 fl (BM, BR, F, GH, GOET, K, LD,
NY, US). Haiti: Ekman9903 fl (US); E. C. & G. M. Leonard13950 st (A, NY); Nash 62 fr (F, K, NY);
394 fl fr (F, NY); Poiteausn fr (P). PUERTO RICO. Britton7818 fl fr (NY); Britton& Hess 2692 fl
fr (NY, US); Britton& Shafer1573 fl (NY, US); Cowell559 fl fr (NY, US); Goll, Cook& Collins864 fr
(NY, US); Heller 7 fl (F, K, LD, NY, US); Millspaugh284 fl (F, NY); Mohn 138 fl fr (F, GH, NY);
Plie sn fr (P); Shafer2934 fl fr (F, NY, US); 3165 fr (NY, US); Sintenis587 fl (BM, K, GOET, LD, NY,
P, US); 5735 fl (LD, NY); 5793 fl (BR, F, GH, K, LD, NY, P, US); 6645 fl (BM, GH, K, NY, US);
Underwood& Griggs 251 st (NY, US); 947 fl fr (NY, US); 948 fl (NY, US). VIRGIN ISLANDS.
Tortola: Fishlock371 fl fr (A, GH, NY). St. Thomas: Eggers 134 fl (BR, K, P); Richardsn fl (P). St.
John: Morrow 148 fl (US). St. Croix: Borgesensn fl (BR, P); Ricksecker353 fl (F, P). LEEWARD
ISLANDS. Anguila: ProctorI8778 fr (A). St. Martin: Stoffers2410 fl (A, U). St. Bartholomew: Forsstrom
sn fl fr (NY); Questel124 fl (NY). St. Kitts: Britton & Cowell211 fl fr (NY); Fairchildsn fl (A); Walshsn
fl (NY). St. Jan: Britton & Shafer286 st (NY, US); 564 fr (NY, US). St. Eustatius: Boldingh88oB fl fr
(K). Barbuda: Box 658 fl (BM). Saba: Boldingh22goB fr (NY). Guadeloupe: Duss 2212 fl (NY, US);
2730 fr (NY); 2731 fl fr (NY); Proctor19855 fr (A, BM); Quentin& Bena 5541 fl (US); Rodriguez3218
fl (A). La Desirade: Proctor21316 fl (A). WINDWARD ISLANDS. Dominica: Cowan1597 fl fr (NY
US); Hodge582 fl (BM, GH; 583fr (BM, GH, US); 584 fl fr (GH, US); 585 fl (GH); Lloyd613 fl fr
(NY); Ramagesn (BM, K); Walsh sn fr (NY). Martinique: Drake i fl (P); Duss 1906 fl (NY, US);
Egler 39-200 fr (NY); Hahn g91fr (BM, K, P); 528 fr (K, NY, US); 951 fl (BR, K, P); Stehle4380 fr
(US). St. Lucia: Andersonsn fl (K); P. Beard 1097 fl (GH, US); Howard 11347 fl fr (A); PlIe sn fl fr
(P); Sauer & Sauer2509 fl (F); St. Vincent: Cooley8365 fl (GH); Eggers 6722 fl (A, P); Guildingsn fl
(K); Morton5658 fl (GH); H. H. & G. W. Smith 198 fl (NY). Barbados: Gooding336 fl (BM); Eggers
7145 fl (A); Robertson5343 fl (K). Grenada: Broadwaysn fr (FHO, NY). COLOMBIA. Isla de San
Andres: Proctor3322 fl (US). Magdalena: Dawe 429 fl (US); H. H. Smith 811 fl fr (A, BM, BR, COL,
GH, K, MICH, NY, P, US); RomeroC. 789 fl fr (COL). Atlantico: Dugandet al 3485 fl (COL); Paul 22
fl (US). Bolivar: Curran256 fl (US); Garcia-Barriga13441 fl (COL, US); Heriberto85 fl (US); Killip &
Smith 14093 fl (A, GH, NY, US). Norte de Santander: Garganta1133 fl (F). Antioquia: Archer421 fl
(US); Barkleyet al 17 C 3.56 fl (COL. F); Feddema1832 fl fr (MICH, US); Haught4540 fl (CDL, US);
Huertes& Herndndez4072 fr (COL); Scolniket al 19 An 549 fl (US). Choc6: Ferndndez244 fl (COL, NY,
US); Killip & Cuatrecasas 309054 fl (A, COL, F, US). Cundinamarca: Perez-Arbeldez592 fl (COL, US).
Valle: Cuatrecasas 16177 fl fr (COL, F); Jaramillo 1250 fl (F). Tolima: Josephsn fr (US). Huila: Lehmann
7556 fl fr (F, K). Narinio: RomeroC. 5368 fr (COL); 5551 fr (COL). Without precise locality: Cuming
1289 fr (K); Dawe 429 fl (K); Holton r fl (K); Lehmann98 fl (K); Moritz 1227 fl (BR, P); Seemann
1095 fl (K); Trianasn fl (P). VENEZUELA. Aragua: Pittier 12185 fl (NY, US, VEN). Dist. Federal:
Curran& Haman847 fl (GH); Pittier 12433 fl (A, NY, US, VEN). Sucre: Potter5189 fl (GH); Lasser&
Vareschi3892 fr (VEN). Monagas: Wurdack& Monachino39535 fl (NY, P). Terr. Delta Amacuro:
Bond, Gillin & Brown58 fl fr (NY, US); Gines4849 fl (US); 5171 fl (US); 5178 fl (US); 5232 fl (US);
Wurdack353 fl fr (NY). Tachira: Garcia-Barriga13312 fl (COL, US, VEN). Bolivar: Holt & Gehringer 71
fl (NY, US); Steyermark 86272 fr (NY); 86273 fr (NY); Ll. Williams 11909 fr (F). TRINIDAD. Britton
Chrysobalanus 19

et al 1632 fl (NY, US); 2005 fl fr (GH, NY, US); Broadwayo124 fl (P); 2169 fl (BR, F, MICH, US);
6680 fl (F, FHO, K, US); 7659 fl (BM, K, US); 8o14 fl (BM, LA); Eggers Io86 fl (BR, P); Fendlersn
fl (BM, NY); Graf 4 fl (NY); Howard o0492 fl fr (A, NY); Kuntze980 fl (NY); 981 fl (NY); Lockhartsn
fl fr (K); Prance2102 fr (NY); 2103 fr (NY); Purseglove6387 fl (TRIN); Schacksn fl (K); Warming188 fl
(C, F); R. 0. Williams 12493 fr (K, TRIN); 12495 fl fr (K, TRIN). GUYANA. Anderson232 fl (K);
Appun298 fl (K); Bartlettsn fr (K); De La Cruz 990 fl (GH, NY, US); 1536 fl fr (F, GH, NY, US);
2015 fl (F, GH, NY, US); 3656 fl (F, GH, NY, US); 3932 fl (F, GH, NY, US); 4226 f (F, GH, K, NY,
US); 4443 fl (F, GH, NY, US); Fanshawe15 (F.D. 2568) fl fr (FHO, K); 2702 (F.D. 5500) fr (K, NY,
US); Harrison741 fl (K, NY, TRIN); Hitchcock16883 fl (GH, NY, US); Im Thurnsn fl (P); Jenman298
fl (P); 540 fl (P); 2258 fl (BM, K); 4666 fl (F, P); 4976 fl (K); Parkersn fl (K); Tutin 94 fl (BM, K,
US); Schomburgk 39 fl (BR, K); 92 fl (BM, K, OXF, P); 220 fl (BM, K, OXF, P, US); Wilson-Browne 595
(FD. 5967) fl (K, NY). SURINAME. Hostmannsn fl (BM, NY); 476 fl (K); Irwin,Prance,Soderstrom, &
Holmgren55224 fl (F, NY, US); 55915 fl (F, GH, MICH, NY, US); Kramer& Hekking2988 fl (NY);
Lanjouw& Lindeman574 fl (K, NY); 1202 fl (K, NY); Schweinfurth sn fl fr (NY); Splitgerber210 fl (P);
Stahel 43 fl (NY). FRENCH GUIANA. Aubrgville24 fl (P); Benoist779 fl (P); i658 fl (P); 1709 fl (P);
Blacket al sn fr (NY); Cowan38890 fl fr (NY, P, US); Cowan& Maguire38045 fl fr (F, NY, US); Gando-
ger 116 fl (P); Lemeesn fl (P); Leprieursn fl (P); Martin sn fl (BR, P); Melinon 130 fl (P); 197 fl fr (P);
Poiteausn fl (K); Sagot 202 fl (BM, BR, K, P). ECUADOR. Johansen 14 fl (US); Mille Io59 fl (F).
BRAZIL. Amazonas: (Cult.), FrSes21590 fl (K); Rodrigues& Chagas1889 fl fr (INPA 8261); 4918 fr
(INPA I3451). Para: Black 48-2934 fl (IAN, NY); 48-3614 fl fr (IAN, NY); DuckeMG 2508 fl (BM,
RBI; Egler I54 fl (INPA, MG 21994, US); Fr6es22840 st (IAN, NY); Killip & Smith30382 fl fr (NY.
RB, US); Pires51898 fr (MICH, NY, RB, US); Prance& Silva 58762 fl fr (NY, US); Prance& Penning-
ton 1271 f fr (NY); 1304 fl (GH, NY). Maranhao: Frdes 8 fl (US); 1992 fl (A, BM, F, K, MICH,
NY, P, US); 21590 fl fr (NY, SP, US); Ducke MG 434 fl (BM); Snethlager56a fl (F). Ceara: Drouet
2455 fl (GH, MICH, NY, SP, US); 2676fl (GH, NY, SP, US). Rio Grande do Norte: WurdackBi89
fl fr (NY); Tavares52-64 fl (HST, IPA, US); 53-335 fl (HST, IPA). Pernambuco: Gardner1150 fl (BM,
GH, OXF); Pickel 201 fl (IPA); Sobrinhosn fl (IPA, RB). Bahia: Belem & Pinheiro 2160 fl (UB);
2165 fr (UB); 2512 fl (UB); Blanchet1822 fl (NY, P); 2299 fl (P); Duarte6065 fl (NY, RB); Gardner902
fl (GH, K, P); 1150 fl (K, P); Martius 511 fl (BR); Pires 3433 fl (NY, P); Riedelsn fl (US). Espirito
Santo: Mattos 10788 fl (SP). Guanabara: Glaziou 10698 fl (K, P); Emygdio 1197 fl (NY); Miers 3889
fl (P); Riedelsn fl (US); RestingaI (Segadas-Vianna et al) 757 fl (NY); 803 fl fr (GH, NY, RB); 878 fl fr
(GH, NY, SP). Sao Paulo: Hoehne29359 fl fr (F, NY, SP); Kuhlmann2983 fl (SP). Without precise
locality: Blanchet1254 fr (BM); 1415 fl (BM); 1822 fl (BM).

LOCAL NAMESAND USES. Most widely known by the names Cocoa Plum, Fat Pork
and Icacoor modifications of the latter name: eg Cacco(Mexico), Zicaque(Antilles),
Icacillo (Venezuela). Other names are: Guyana: Caramio;Suriname: Koenatepie,
Kodibiassiballi, Koelimiro (Arawak), Pruim (Sur.), Koenoto-Epoe, Erejoeroe (Kaar.).
Brazil: Ajuru-branco, Cajuru,Goajuru', Oajuru.The fruit is edible and in Colombia and
Venezuela it is cooked, bottled in syrup, and sold commercially.
The extreme variation in leaf shape and size, and in fruit size exhibited by this
species has led to the various names cited in synonymy. In analyzing this variation
carefully, I could find no basis for the recognition of more than one species. There is
little correlation among the variable characters, and no obvious grounds exist for the
creation of subspecific taxa. Therefore I do not follow Hooker (I867), who divided
the New World material of this species into three varieties. In this treatment var
pellocarpa,and var ellipticusare not recognized as separate taxa. I have observed C.
icacoin the field in Trinidad, Suriname and Brazil. Frequently the different forms
grow side by side without any ecological separation, varying principally in the leaf
and the fruit. The leaf varies, even on one individual, from an extremely small to a
large size, and in shape from orbicular to elliptic. The fruit may be up to 4.5 cm long,
or it may be as small as o.8 mm long. The larger fruits are more conspicuously costate
than the smaller ones, and tend to have a thicker more fleshy mesocarp. There is also
variation in the pigment of the fruit, the mature fruit may be deep purple or black or
it may be yellow. The prefix Chryso (golden) suggests that the material described by
Linnaeus was yellow-skinned. The fruit ofC. icacois eaten by man, although only
the larger more fleshy fruit make good eating. The differences in fruit type are com-
parable to those between good and bad varieties of plums (PrunusdomesticaL.),
 20 Flora Neotropica

and where C. icaco is used commercially obviously there is some selection of the plants
in cultivation. At present it seems best to maintain C. icaco as a single polymorphic
species. As it is impossible to subdivide this species using information based on her-
barium material and ecology alone, experimental cultivation of plants of the different
morphological types in controlled environments might suggest a more satisfactory
basis for taxonomic status. The two attempts that I have made to examine the
chromosomes of C. icaco have also proved unsuccessful, but I hope to make further
attempts at a later date.
African material of this species seems as variable as the American. Hauman
(I951) gave reasons for regarding C. orbicularis as a species separate from C. icaco,
but as none of his differences holds true when a large amount of material is studied, I
have concluded that C. orbicularisis identical with C. icaco. The African material has
been placed into three species C. orbicularis,C. ellipticus and C. atacorensis.The last is
probably a distinct species, but C. ellipticus, with elliptic acuminate leaves, merges,
both morphologically and geographically, gradually into C. icacowith orbicular retuse
leaves. C. ellipticus must be regarded as part of C. icaco, but I have not studied the
Africa material in enough detail to determine whether it merits subspecific recogni-
tion.
Studies on the origin of cultivated plants raise the question of the original
province of C. icaco whether its occurrence on both continents is the result of its
carriage from one continent to the other during the time of the slave trade. Present
evidence strongly suggests that C. icaco is a native of both Africa and America. In
America C. icaco has a very wide range for a species which was introduced compara-
tively recently and is not primarily a cultivated plant. Most of the fruits used are
harvested from wild plants in sandy, littoral areas. C. icaco occurs well inland in some
of the remotest savannas in the Guianas where it would be hard to argue for its intro-
duction. For example, I have collected it in such a savanna near the Wilhelmina
Gebergte of Suriname. Chrysobalanusis long-established in Africa by the distribution
of the distinct species, C. atacorensis,through the Congo Basin to Zambia, as well as by
the distribution of the more acute-leaved forms of C. icacowhich in the past have been
named C. ellipticus. Although C. icaco is variable in America, it is just as variable in
Africa and grows in natural rather than man-made habitats. The bicontinental
distribution of C. icaco and of Parinari excelsa will be discussed in further detail in a
separate paper.

2. Chrysobalanus cuspidatus Grisebach ex Duss, (Grisebach, Fl. Brit. W.

Ind. 711 . 864, nom. nud.) ex Duss, Fl. Ant. Fran9aise 25g. 1867; Urb.,
Symb. Antill. 5' 351. I907; Howard, Jour. Arnold Arb. 45: 279. 1964.
Fig 2 G.
Licania oliganthaA. C. Smith, Jour. Arnold Arb. 28: 333. 1947. Type. Beard 492, St. Lucia
(holotype, A; isotypes, K, LE, MO, NY).

Small tree, the short-pubescent young branches becoming lenticellate with age.
Leaves oblong to elliptic, 4.0-II.0 cm long, 2.0-4.5 cm broad, apex with well
developed acumen 5.0-I2.0 mm long, base rounded to subcuneate, both surfaces
glabrous, sparsely hirsute on midrib beneath, the primary veins inconspicuous;
petioles 3.0-6.0 mm long, sparsely pubescent when young. Stipules subpersistent,
lanceolate, 5.o-8.o mm long, intrapetiolar. Inflorescences few-flowered terminal and
axillary cymules with brown pubescence. Receptacle cupuliform, pubescent inside
and outside. Calyx lobes broadly ovate. Petals 5, exceeding calyx lobes. Stamens ca
Chrysobalanus 21

12, the filaments joined in groups at base, pubescent. Ovary and style pilose. Fruit
obovate, ca 2.7 cm long; epicarp smooth and ridged; mesocarp thin and fleshy;
endocarp hard and thin 5-7 longitudinal ridges.
TYPE. Duss 3633, Guadeloupe, (holotype, B, lost; lectotype, NY); 3476 fr
(paratype, NY); 154, Martinique, fr (paratypes, F, GH, NY, US).
DISTRIBUTION. Forest, low woodland and wooded slopes, confined to the Lesser
Antilles. Flowering Aug-Dec.
LEEWARD ISLANDS. St. Kitts: Wadsworthet al 523 st (A); 575 fl (A); 588 fr (A). WINDWARD
ISLANDS. Dominica: Eggers6i8 fl (BR, GH, GOET, P); 986 fl (K); Imraysn fl (GOET, K). Martini-
que: Stehle6519 fl (US); 6805 fl (US). St. Lucia: Beard488 fl (A, K, NY); Howard1638 fl (A); Proctor
21621 fl (A).

LOCALNAMES.Guadeloupe: Icaquemontagne.Martinique: Tifeuilles; Zicac des bois.

Excluded American Names and Species

(L.) Morales, An. Acad. Ci. Mod. Fisic. & Nat. Habana 47: 39I.
I. C. americanus
1887. =Hirtella americanaL.
2. C. humilis (Cham. & Schlecht.) Kuntze, Rev. Gen. 3(2): 76. I891. =Licania
humilisCham. & Schlecht.
3. C. incanus Rafin., New Fl. Am. 3: 26. 1836. =Licania michauxii Prance.
4. C. incanus(Aubl.) G6mez de la Maza, Fl. Cuba 39. 1887. =Licania incanaAubl.
5. C. macrophyllusSchott in Spreng., Linn. Syst. Veg. ed. I6. 4, App. 407. 1828.
=Couepia schottii Fritsch.
6. C. montanus(Aubl.) G6mez de la Maza, Fl. Cuba 39. 1887. =Parinari montanaAubl.
7. C. oblongifolius Michx., Fl. Bor. Am. I: 283. I803 =Licania michauxii Prance.
8. C. ovalifolius Schott. in Spreng., Linn. Syst. Veg. ed. 16. 4, App. 406. 1828.
=Couepia ovalifolius (Schott) Benth.
9. C. pallidus (Small) L. B. Smith, Rhodora 48: 136. I946. = Licania michauxii Prance.
Io. C. prunifolius Rafin., New Fl. Am. 3: 26. 1836 in obs. (see p 43, sub Licania
michauxii).
1. C. retusus Rafin., New Fl. Am. 3: 26. 1836. =Licania michauxii Prance.
I2. C. rugosus(Thuill. ex Pers.) G6mez de la Maza, Dicc. Bot. Nom. Vulg. Cub. &
Puerto Riquenos. o10. 1889. =Hirtella rugosaThuill. ex Pers.
I3. C. sublanatus Kuntze, Rev. Gen. 3(2): 761. I89g.=Licania humilis Cham. &
Schlecht.
14. C. subundulatus Brign. in Mem. Soc. Ital. Mod. II. I: 71. 1862. =probably
Couepiasp (type not seen).
I5. C. triandrus(Sw.) Morales, Ann. Acad. Ci. Med. Fisic. & Nat. Habana 43: 390.
1887. = HirtellatriandraSw.

2. Licania Aublet, P1. Guiane I: I19, t. 45. 1775; Don, Gen. Syst. 2: 479. 1832,
"Lincania" sphalm.; DC., Prodr. 2: 527. I825; Benth., Jour. Bot. Hooker
2: 212-215, 218-222. 1840; Hook. f., Mart. Fl. Bras. 14(2): 8-26. 1867;
Fritsch, Ann. Naturh. Mus. Wien 4: 33-60. 1889; Benoist, Bull. Mus. Hist.
Nat. Paris 25: 512. 1919; Sandwith, Kew Bull. 1931: 369-74. I93I;
Kleinhoonte in Pulle, Fl. Surin. 2(I): 430-444. I939; Lemee, Fl. Guyane
franc. 2: I6-24. I952; Prance, Synopsis (in press),Jour. Arnold Arb. 51:
521-528. 1970.
MoquileaAublet, P1. Guiane i: 521, t.208. 1775.
DahuroniaScopoli, Introd. 2I7. I777.
HedycreaSchreber, Linn. Gen. P1. ed. 8. I: I60. 1789.
 22 Flora Neotropica

Hirtella sensu E. Meyer, Nov. Acta Acad. Leop.-Carol. 21: 803. 1825 et auct. div. non L.
AngelesiaKorthals, Nederl. Kruidk. Arch. 3: 384. 1854.
TrichocaryaMiquel, Fl. Ind. Bat. I: 258. I855. quoad T. splendenstantum.
Chrysobalanus auct. div. non L. sensu G6mez de la Maza, Fl. Cuba 39. 1887, pro parte quoad
C. incanatantum; sensu Kuntze, Rev. Gen. 3(2): 76. I891.
GeobalanusSmall, Fl. Miami. 80. 1913.
CoccomeliaRidley, Jour. As. Soc. Str. Branch 72: I83. I920.

Small to large trees or shrubs, rarely suffrutices. Flowers hermaphrodite. Leaves

entire, glabrous above at maturity, lanate or pulverulent or strigose or glabrous or
with stomatal cavities on lower surface. Petioles with two or more sessile glands, or
eglandular. Bracts and bracteoles usually eglandular, usually small, very rarely large
and enclosing small groups of flower buds. Inflorescences most frequently sparsely
branched racemose panicles, less frequently a panicle of cymules or a spike. Receptacle
I.o-8.o mm long, varying from subglobose or campanulate to urceolate, the interior
always pubescent. Calyx lobes 5, acute. Petals (4-)5 or absent. Stamens 3-40, uni-
lateral or inserted in a complete circle; filaments usually free to base, rarely connate,
usually glabrous, long-exserted beyond the calyx lobes to short and included. Ovary
inserted at or near the base of the receptacle, rarely slightly laterally inserted, usually
pilose; carpel unilocular, ovules 2. Style filiform, of varied length. Fruit a small to
large, dry or flesy drupe, exterior varying from densely tomentose to pulverulent or
glabrous, the surface smooth or verrucose; pericarp sometimes in one layer, more
often divided into a fleshy mesocarp and a hard woody or bony endocarp which has
no special mechanism for seedling escape. Seed large, erect, filling the loculus.
Germination hypogeal, first leaves alternate.
TYPE SPECIES. Licania incana Aublet. The name Licania is derived from the
vernacular name "Calignia", intended to be an anagram.
DISTRIBUTION. Largely New World from Mexico, Florida, West Indies to
Southern Brazil, with one widespread species in the Asiatic tropics. Most abundant in
the lowland forests and savannas of the Guianas and the Amazon Basin.

Taxonomic History of Licania

The history of this genus up to 1889 was well summarized by Fritsch (1889),
and to present all information in this one summary much of Fritsch's account is re-
peated here. Licaniawas first described by Aublet ('775) who based it on the single
species L. incana.However, Aublet misunderstood the floral structure, interpreting
the calyx as a corolla and the bracteoles as a calyx. In the same work Aublet des-
cribed Moquilea,which he did not relate to Licaniabecause his work was based on the
Linnaean sexual system of classification. The very different number of stamens be-
tween Aublet's species of Licaniaand Moquilearesulted in the placement of these two
genera in different groups. Scopoli (1777) used the new but illegitimate name
Dahuroniafor Moquileaof Aublet. Schreber in his edition of GeneraPlantarum(1789)
corrected Aublet's description of Licaniaand pointed out the absence of a corolla.
Although his generic diagnosis is free of fault, he gave the new illegitimate name
Hedycrea to Licania. Jussieu (1789), the first to relate Moquilea to Licania, considered
these two genera as members of his subfamily Amygdaloideae. De Candolle (I825)
in the Prodromusreferred to Licania two species previously described in Hirtella, H.
octandraHoffmannsegg. ex Roem. & Schult. and H. apetala E. Mey. Martius (1827)
united Couepiaand Actoa with Moquilea, gave a detailed description of his conception of
the expanded genus Moquilea, and described the new species Moquilea canomensis which
was later transferred to Couepia by Bentham (i 840). With this act the confusion be-
tween Moquilea and Couepia began. In 1832 Martius & Zuccarini discussed Moquilea
Licania 23

further. With the exception of Aublet's species of Moquilea all the rest of the species,
which Martius placed in Moquileahave at a later date (Bentham, I840; Hooker,
1867; Prance, in press) been transferredto the genus Couepia.Meisner (1837) in his
GeneraPlantarum,retained Couepiaand Acioa as sections of Moquilea,although he
suggested that restoration of their generic status was justified. Endlicher ( 1840) held a
similar interpretation of Moquilea.None of these authors appears to have seen the
Aublet material.
One of the first significant contributions to our knowledge of Licaniawas made
by Bentham (1840). He clarified the definition of the genera of the Chrysobalanaceae,
and recognized Moquilea,Couepia(incl. Acioa) and Licania as separate genera. He
correctly placed all previously described species of Moquilea,except M. guianensis
Aubl. in Couepia,and he described the new species M. tomentosa. He divided Licania
into five sections with the comment that these were perhaps worthy of generic rank.
His sections were:
I. Batheogyne.Calyx broadly campanulate; petals absent; fertile stamens Io-I2,
unilateral, shortly exserted: L. turbinataBenth.
2. Leptobalanus. Calyx broadly campanulate; petals absent; stamens io, inserted
in a complete circle, all fertile, exserted: L. pendulaBenth.; L. floribundaBenth.; L.
pubiforaBenth.; L. humilisCham. & Schlecht.; L. turiuvaCham. & Schlecht.
3. Microdesmia. Calyx ovate; petals 5, small, or absent; fertile stamens about o0,
unilateral, included: L. mollisBenth.
4. Eulicania. Calyx ovate or subglobose; petals absent; fertile stamens 3-5,
unilateral, included: L. leptostachya Benth.; L. incanaAubl.; L. crassifoliaBenth.; L
coriaceaBenth.; L. parvifloraBenth.
5. Hymenopus.Calyx ovate or subglobose; petals 4-5; fertile stamens 5-8,
unilateral, included: L. divaricataBenth.; L. heteromorpha Benth.
Thus Bentham recognized 14 species of Licania, a few of which have been
merged, as well as two species of Moquilea.
During the subsequent years new species were added to most sections. Blume
(i856) included Couepiaand Acioain Moquilea.Up to I857 Licaniaand Moquileawere
always regarded as separate genera. Even though relatively few species were known
at that time, the only character separating them was the greater number of stamens
in Moquilea.Grisebach (I857) found plants with intermediate numbers of stamens,
which led to his merging Moquilea(sensu Bentham) with Licania.
J. D. Hooker, in Bentham and Hooker's (I865) GeneraPlantarum,retained the
two genera Licania and Moquilea and expanded Moquilea to include Bentham's
sections of Licania,Leptobalanusand Batheogyne.In his treatment of the genus in
Martius' Flora Brasiliensis (1867), Hooker again retained Licania and Moquilea
separately. The only character distinguishing these genera was filament length, the
exserted stamens of A/Ioquileacontrasting with the included stamens of Licania.
Hooker maintained two sections in Licania,Hymenopus (including sect Microdesmia of
Bentham) and Eulicania. He recognized the sections Eumoquileaand Leptobalanus
(including sect. Batheogyne of Bentham) in Moquilea.
Baillon (1869) and later Eichler (1878) followed Grisebach in merging Licania
and Moquilea.
Fritsch (I889) published a conspectus of the genus Licania.This useful work is
the only summary of an entire genus of the Chrysobalanaceae published since De
Candolle's Prodromus. While Fritsch'swork brought together much information, it was
only a conspectus and not a critical revision of the genus. He reduced several names
to synonymy, most of which I have followed here, but he did not see any material of
many species, and understandably misinterpreted some. He gave convincing evi-
dence for uniting Moquileawith Licaniaand recognized 60 species. It is not clear why
24 Flora Neotropica

later authors did not accept this position. For example, Focke (1891), in Engler and
Prantl's Pflanzenfamilien,re-est; blished Licania and Moquilea as separate genera.
During the present century, authors have been divided in opinion as to the status of
Licaniaand Moquilea.Many new species of Licaniahave been described since the time
of Fritsch'sConspectus;but no critical revision of the entire genus has been undertaken.
Huber (9goga) established a new subgenus, Parinariopsis,to accommodate his new
The most noteworthy accounts during the present century are
species, L. parinarioides.
three regional revisions of a few species of the genus: Benoist (I919) published notes
on the species of Licaniafrom French Guiana; Macbride (I 938) revised the Peruvian
species; and Kleinhoonte (1939) in revising the Suriname representation included 18
species.
The preceding account shows that this large genus has never been compre-
hensively revised, and was thus badly in need of attention at the outset of the present
work.

Specific and Sectional Characters in Licania

Since this is the largest and most complex genus in the family, a large number of
specific characters are used herein. To facilitate use of the keys and the descriptions
some of the characters are discussed here:

THE LEAF Fig 3.

One of the most important taxonomic characters is the nature of the lower
surface of the leaf. There are five main types, any one of which may be characteristic
of an entire section or of a group of closely related species.
i. In the simplest type the leaf underside is entirely glabrous,and the venation is
usually inconspicuous (Fig 3 E).
2. There are a few related species with a strigose-hirsutulouspubescence along the
veins and reticulations, but the intervening spaces are glabrous (Fig 3 A).
3. Some of the species of subgenus Licaniahave a distinct pulverulent-furfuraceous
indumentum. In the Chrysobalanaceae this feature is known only in these closely
related species (Fig 3 B).
4. The commonest type of indumentum is dense lanate-arachnoid pubescence
which covers the entire undersurface. The venation may be extremely prominent to
almost plane (Fig 3 C).
5. Stomatalcavities,as discussed by Homes et al (I951) and Prance (in press), is a
term used frequently in the following descriptions of species of Licania.A leaf is said
to have stomatal cavities when the venation is deeply reticulate, and the veins are
flattened on the surface to make the leaf riddled with small cavities which contain the
stomata. The cavities are always filled at the mouth with a lanate pubescence. In
species with well developed cavities the cavities are a most distinctive feature. The
hair-filled cavities suggest that the phenomenon is perhaps a xerophytic adaptation
(Fig 3 D). Stomatal cavities occur in Licaniasubgenera Leptobalanus and Licania,as
well as in two species of Couepiaand nearly all species of Parinari.
Several types of leaf glands are found in Licania,especially in subgenus Moquilea,
and elsewhere in species with glabrous leaves. The glands are, however, poor taxono-
mic characters. They show a marked similarity to those of Prunus,as discussed and
illustrated by Kalkmann (i 965). Glands are found on the lower surface of the lamina,
at the termination of the primary veins, and in many species on the petioles. I am
inclined to agree with Kalkmann and others about the homology of all leaf glands of
this type, marginal, laminar, and petiolar, especially in view of the range of variation.
Licania 25

A B

,~
:I~~~.~

~,~
~:,~:',.???' ~.~~~~~~~~~,.~.~ 1 ..

._..

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

~.
~

it
I~'
?..:
"'.?
:~ I.i
'<~ (>
.. .
?):'? ' .
~~

' ~ ~ ~ ~ ~ Loke~
?:?~~~~~~~~~.:.

."'F c:ansc.
B L. ( re

30397); C, L. incana (Maguire 24234); D, L. albiftora(Fanshawe,678). E, L. heteromorpha

(De La Cruz
44"), all x6.
26 Flora Neotropica

In Licaniabasal glands may be on the lamina surface, at the junction of the lamina
and petiole or well down on the petiole. The position of the glands seems to be most
frequently constant within a species.

THE STIPULES

The most important feature of the stipules is their position. Many species of
Licaniahave stipules which are distinctly adnate to the petiole, a feature that does not
occur elsewhere in the family. This feature is most frequent in subgenus Licania,and
does not occur in subgenera Moquileaor Leptobalanus. In other species the stipules are
axillary and may be either intrapetiolar or on the outside of the petiole.

INFLORESCENCEFig 4.
i. Many species of Licania are characterized by a few-branched compound
raceme termed here a racemose panicle(Fig 4 A). It is often easy to recognize Licania
from other genera by this character alone.
2. A much-branched panicle so common in most other genera of the family is
rare in Licania.(Fig 4 B).
3. An inflorescence type that occurs in most of the subgenera in groups of re-
lated species is a panicleof cymules,where the flowers are borne in small groups of 2-6
in cymules on slender distinct peduncles. (Fig 4 C). This feature, which is useful for
the delimitation of some species, appears to have arisen independently several times
in the genus.
4. A spike with the flowers remote or densely crowded on the rachis is character-
istic of a few species (Fig 4 D).

THE FLOWER

Most of the floral characters used are self explanatory. Flower length has been
measured from the tip of the longest calyx-lobe to the articulation of the pedicel.
The stamens are exserted when they obviously exceed the calyxlobes.In most species
this is easy to observe, but in a few the stamens are intermediate between the exserted
and included condition. The shape of the receptacle is constant within a species, and
three main types occur: cupuliform, campanulate, and urceolate (Fig 4 E-G).

Subgenera and Sections of Licania

Unlike most other genera in the family, Licania can be validly divided into
subgenera and sections. The differencesbetween the subgenera and sections are small,
thus minimizing grounds for elevating such taxa to generic rank. Reasons for uniting
Moquileawith Licaniahave been discussedelsewhere in greater detail (Prance, in press).
The sections adopted here agree closely with those originally proposed by Bentham
(1840).

Synopsis of Subgenera and Sections of Licania

I. Subgenus Moquilea (Aublet) Prance, Atas do Simp6sio s6bre a Biota Amazonica
4: 224. I967.

MoquileaAublet, P1. Guiane I: 521. 1775; Hook, f., Mart. Fl. Bras. 14(2): 19. 1867.

Stamens 8-50, inserted in a complete circle, the filaments usually far exceeding
Licania 27

A B

C D

FIG 4. Inflorescence and Receptacle Characters in Licania A-F, Inflorescences. A, racemose

panicle; B, panicle; C, panicle of cymules; D, spike. E-G, Receptacle: E, cupuliform; F, campanulate;
G, urceolate.
28 Flora Neotropica

or rarely equalling the calyx lobes; petals present or absent; ovary inserted at base of
receptacle; bracteoles small; leaf undersurface glabrous, lanate, or with stomatal
cavities.
TYPE SPECIES.
Licania guianensis (Aublet) Grisebach
i. Section Moquilea (Aublet) Prance, Jour. Arnold Arb. 51: 527. I970.
Stamens I4-50, far exceeding calyx lobes; petals present; leaf undersurface
glabrous or lanate.
ii. Section Leptobalanus Bentham, Jour. Bot. Hooker 2: 212. I840; Hook. f.,
Mart. Fl. Bras. I4(2) :2I I867. Incl. section Batheogyne Bentham, Jour. Bot.
Hooker 2: 212. 1840.
Stamens 8-I4 (22 in L. calvescens);usually far exceeding calyx lobes (equalling
calyx lobes in L. emarginata);petals absent; leaf undersurface glabrous, lanate, or
with stomatal cavities.
TYPE SPECIES. Licania apetala (E. Meyer) Fritsch
iii. Section Microdesmia Bentham emend, Jour. Bot. Hooker 2: 2 2. I840.
Stamens 8-25, equalling calyx lobes in length; petals present; leaf undersurface
glabrous, lanate, or with stomatal cavities.
TYPE SPECIES.
Licania rigida Bentham
2. Subgenus Parinariopsis Huber, Bol. Mus. Emilio Goeldi 5: 368. I909.
Stamens 16-25, inserted in a complete circle; filaments equalling or slightly
exceeding the calyx lobes; petals present; ovary inserted laterally at side of receptacle;
bracteoles large and enclosing groups of buds; leaf undersurfacedeeply reticulate and
lanate.
TYPE SPECIES.
Licania parinarioides Huber
3. Subgenus Licania
Stamens 3-8(-I I), unilateral or inserted in a complete circle; filaments shorter
than or equalling the calyx lobes; petals present or absent; ovary inserted at base of
receptacle; bracteoles usually much shorter than or rarely equalling calyx; leaf
undersurface variously glabrous, hirsute, pulverulent, lanate, or with stomatal
cavities.
Licania incana Aublet
TYPE SPECIES.
i. Section Hirsuta Prance, Atas do Simp6sio s6bre a Biota Amazonica 4: 226.
i967.
Leaf undersurface hirsute at least on venation, not tomentose; petals present;
flowers borne on primary branches of inflorescence, never in cymules.
TYPE SPECIES.Licania latifolia Bentham
ii. Section Hymenopus Bentham, Jour. Bot. Hooker 2: 212. I840; Section
Hymenopussensu Hook. f., Mart. Fl. Bras. 14(2): 9. I867 pro parte.
Leaf undersurface entirely glabrous; petals present or absent; flowers borne on
primary branches of inflorescence or in small cymules on slender peduncles.
TYPE SPECIES.
Licania divaricataBentham
iii. Section Cymosa Prance, Atas do Simp6sio s6bre a Biota Amazonica 4: 226.
I967.
Leaf undersurface pulverulent-farinaceousor lanate, or with stomatal cavities;
petals absent; flowers borne in small cymules on short secondary branches of inflores-
cence.
TYPE SPECIES.
Licania hypoleucaBentham
iv. Section Pulverulenta Prance, Atas do Simp6sio s6bre a Biota Amazonica
4: 226. I967.
Leaf undersurface pulverulent-furfuraceous; petals absent; flowers borne on
primary branches of inflorescence.
Licania coriaceaBentham
TYPE SPECIES.
Licania 29

v. Section Licania
Leaf undersurfacelanate or with stomatal cavities; petals absent; flowers borne
on primary branches of inflorescence.
TYPE SPECIES.
Licania incana Aublet

Key to Subgenera and Sections of Licania

i. Stamens far exceeding calyx lobes. Subgenus Moquilea.
2. Petals present; stamens 14-50. Section Moquilea.
2. Petals absent; stamens 8-14. Section Leptobalanus.
I. Stamens equalling or shorter than calyx lobes.
3. Stamens 10-25.
4. Petals present.
5. Ovary inserted at base of receptacle; bracteoles small.
Subgenus MoquileaSection Microdesmia.
5. Ovary inserted laterally at side of receptacle; bracteoles large and enclosing groups of
flower buds. Subgenus Parinariopsis
4. Petals absent.
6. Leaf underside glabrous or with easily removed lanate pubescence.
Subgenus MoquileaSection Leptobalanus.
6. Leaf undersurface with deep reticulation and persistent pubescence.
Subgenus Licania Section Licania.
3. Stamens 3-8. Subgenus Licania.
7. Leaf undersurface completely glabrous; petals present or absent.
Section Hymenopus.
7. Leaf undersurface pubescent; petals present or absent.
8. Leaf underside sparsely hirsute on venation; petals present. Section Hirsuta.
8. Leaf underside densely lanate or pulverulent or with stomatal cavities; petals absent.
9. Inflorescence a panicle of cymules borne on long slender pedicels.
Section Cymosa.
9. Inflorescence a racemose panicle or a spike, without cymules.
10. Leaf underside pulverulent-furfuraceous. Section Pulverulenta.
Io. Leaf underside never pulverulent, but lanate-pubescent or with stomatal cavities.
Section Licania.

Key to Species of Licania

i. Stamens far exserted (filaments twice as long as calyx lobes). (For contrasting statement see p 32).
Subgenus Moquilea.
2. Petals present; stamens I4-50. (For contrasting statement see p 31). Section Moquilea.
3. Leaf underside with an appressed lanate-arachnoid pubescence.
4. Primary leaf veins 8-15, the lamina oblong-lanceolate.
5. Petioles 1.0-3.0 mm long; fruit 2.5-3.0 mm long, with smooth exterior, pericarp
thin; inflorescence of small cymose panicles. (Southeastern U.S.A.) i. L. michauxii.
5. Petioles 5.0-7.0 mm long; fruit 4.0-7.0 mm long, with a crustaceous, verrucose
exterior, pericarp thick; inflorescence of racemose panicles. (Bolivia-La Paz.)
2. L. boliviensis.
4. Primary leaf veins 16-25 pairs; leaves oblong-elliptic.
6. Leaves gradually tapering from about mid point to acuminate apex; center primary
leaf veins far apart (average 18.22 mm apart); bracteoles coriaceous, persistent. (Colombia)
3. L. maritima.
6. Leaves tapering from well above mid point, apex abruptly caudate; center primary
leaf veins close together (average 11.22 mm apart); bracteoles caducous or thin and
membraneous.
7. Mesocarp ca 2.0 mm thick, without parallel fibers, the thin endocarp adnate to
it; leaf underside gray-pubescent to ferrugineous-pubescent; flowers densely
clustered; exterior of flowers yellow-brown-pubescent. (Colombia, Ecuador.)
4. L. durifolia.
7. Mesocarp ca o0.0 mm thick, with parallel fibers; the endocarp detached from it;
leaf underside ferrugineous pubescent; flowers solitary; exterior of flower ferrugi-
neous. (Colombia.) 5. L. macrocarpa.
30 Flora Neotropica

3. Leaf underside glabrous when mature (sometimes caducous-lanate when young).

8. Inflorescence and exterior of flowers glabrous or sparsely puberulous.
9. Leaves broadly acuminate, the acumen 4.0-8.0 mm long; flowers 2.5-3.0 mm long;
pedicels 2.0-5.0 mm long. (Brazil-Bahia.) 6. L. salzmannii.
9. Leaves with well developed finely pointed acumen 8.o-I2.0 mm long; flowers
4.0-4.5 mm long; pedicels 1.0-3.0 mm long.
o0. Inflorescence and exterior of flowers puberulous; petioles Io.o-I2.0 mm long;
leaves elliptic 5.5-9.0 cm broad. (Peru-Loreto.) 7. L. klugii.
o0. Inflorescence and exterior of flowers glabrescent; petioles 5.0-7.0 (-9) mm
long; leaves oblong to oblong-elliptic; 2.0-5.5 cm broad. (Guianas & Amazon.)
8. L. guianensis.
8. Exterior of flowers and usually inflorescence densely tomentose.
I . Inflorescence of dense pyramidal or rather lax cymose panicles, the rachis glabrous
to puberulous; leaf apex rounded-apiculate or mucronate.
I2. Inflorescence short and dense, the rachis glabrous; petioles 4.0-6.0 mm long.
(Mexico.) 9. L. retifolia.
I2. Inflorescence lax, the rachis puberulous; petioles 1.0-3.0 mm long. (South-
eastern U.S.A.) i. L. michauxii.
1. Inflorescence of lax racemes or racemose panicles, the rachis pubescent; leaf apex
usually acuminate.
I3. Pedicels 3.0-7.0 mm long; leaves with finely pointed acumen 1.5-2.5 cm long.
(Brazil-Amazonas.) io. L. longipedicellata.
13. Pedicels o.5-2.5 mm long; leaf acumen rather blunt, less than I0.0 mm long.
14. Inflorescence of predominantly axillary racemes or few-branched panicles.
(Brazil-Amazon & the Northeast.) I . L. tomentosa.
I4. Inflorescence of predominantly terminal much-branched panicles or racemose
panicles.
I5. Leaves with a caducous lanate pubescence when young; petioles 8.o-I6.o
mm long and very slender not more than 1.2 mm thick. (W. Indies,
Venezuela.) 12 L. pyrifolia.
15. Leaves glabrous even when young; petioles 4.0-I2.0 mm long, thick, more
than I.5 mm thick.
16. Leaf bases cuneate; stamens ca 30; petioles canaliculate. (W. Indies,
Venezuela.) 13. L. leucosepala.
I6. Leaf bases cordate, rounded or subcuneate; stamens 14-23; petioles
usually terete, rarely canaliculate.
17. Leaves oblong-lanceolate. (Amazon.) 14. L. angustata.
17. Leaves ovate to oblong.
i8. Leaves 13.0-30.0 cm long; primary veins 14-22 pairs.
I9. Leaves oblong to oblong-lanceolate, 3.5-8.0 cm broad;
primary veins 15-22 pairs; petioles 9.0-i4.0 mm long; leaf
base rounded to subcuneate.(Central America.) 15. L. platypus.
i9. Leaves elliptic, 7.5-9.5 cm broad; primary veins I4-15 pairs;
petioles ca 8.o mm long; leaf base subcordate. (Brazil-Maranhao.)
I9. L. maranhensis.
i8. Leaves 5.0-I5.5 cm long; primary veins 8-I3(-I5).
20. Petals not clawed, equalling or only shortly exceeding calyx
lobes; filaments far exceeding petals.
21. Flowers 4.o-5.0 mm long; stipules linear, filamentous,
3.0-7.0 mm long; plants of Mexico. (Mexico.)
i6. L. gonzalezii.
21. Flowers 2.0-3.5 mm long; stipules short-subulate to 2.0 mm
long; plants of South America.
22. Petioles 2.0-5.0 mm long; leaves usually obovate, apex
cuspidate to mucronate. (Amazon.) 17. L. egleri.
22. Petioles 6.0-I2.0 mm long; leaves oblong to oblong-
elliptic; apex acuminate, never mucronate.
23. Leaves 4.0-I2.0 cm long; flower buds globose;
flowers ca 2.0 mm long; petioles usually pubescent.
(Guianas, Amazon.) 18. L. minutiflora.
23. Leaves 8.0-I5.5 cm long; flower buds ovoid; flowers
ca 3.0 mm long; petioles glabrous when mature.
24. Some primary veins I2.0-I6.0 mm apart; young
Licania 31

stems not conspicuously lenticellate; inflorescence

and flowers brown-pubescent; midrib extremely
prominent above. (Amazon.) 20. L. fritschii.
24. All primary veins 8.0-I 2.0 mm apart; young stems
conspicuously lenticellate; inflorescence and
flowers gray-tomentellous; midrib plane above.
(Amazon.) 21. L. britteniana.
20. Petals unguiculate, much exceeding calyx lobes; filaments
scarcely exceeding petals.
25. Flowers 5.o-6.o mm long; pedicels 2.0-2.5 mm long;
ovary pilose. (Amazon.) 22. L. unguiculata.
25. Flowers 3.0 mm long; pedicels 0-0.25 mm long; ovary
glabrescent. (Amazon.) 23. L. longipetala.
2. Petals absent; stamens 8-I4. Section Leptobalanus.
26. Leaf underside glabrous or lanate, the venation not prominent, the stomatal cavities
absent.
27. Most flowers borne in small groups or cymules on distinct secondary branches (pedun-
cles) of inflorescence; peduncles over 2.0 mm long.
28. Rachis of inflorescence densely yellow-villous-tomentose. (Venezuela, Amazon.)
24. L. wurdackii.
28. Rachis of inflorescence gray-puberulous.
29. Inflorescence short and compact, 3.0-5.0 cm long; flowers 4.0-5.0 mm long;
leaves ovate. (northeastern Brazil.) 25. L. turbinata.
29. Inflorescence longer and spreading; flowers 2.0-3.5 mm long; leaves oblong
to elliptic.
30. Flowers 3.0-3.5 mm long; petioles deeply canaliculate; 2 prominent
glands at lamina base; rachis arachnoid-pubescent when young. (Amazon.)
26. L. lata.
30. Flowers 2.0-2.8 mm long; petioles terete to shallowly canaliculate;
lamina base eglandular; rachis puberulous. (Guiana, Amazon,
southern Brazil.) 27. L. apetala.
27. Flowers sessile or nearly so on primary branches of inflorescence.
3I. Leaf apex predominantly obtuse to rounded.
32. Rachis of inflorescence densely villous-tomentose; flowers ca 4.0 mm long.
(Amazon.) 24. L. wurdackii.
32. Rachis of inflorescence gray-puberulous; flowers 3.0 mm long. (Guiana,
Amazon.) 28. L. parvifolia.
3 . Leaf apex distinctly acuminate.
33. Flowers 4.0-5.0 mm long; receptacle cupuliform.
34. Leaves glabrous beneath, inflorescence gray-villous-tomentose. (Brazil-
Mato Grosso.) 29. L. maguirei.
34. Leaves with short persistent waxy lanate-pulverulent pubescence beneath;
inflorescence brown-puberulous to tomentellous when mature. (Amazon.)
26. L. lata.
33. Flowers 2.0-3.5 mm long; receptacle urceolate or campanulate.
35. Stipules persistent on young branches; petioles densely tomentose when
young, 2.5-3.5 mm thick; upper portion of ovary glabrous. (Southern
Brazil.) 30. L. gardneri.
35. Stipules usually caducous; petioles sparsely lanate-pubescent, puberulous,
or glabrous, to 2.0 mm thick; upper portion of ovary pubescent.
36. Inflorescence sparsely hirsutulous; exterior of flowers yellow-brown-
tomentose; petioles 2.0-3.0 mm long, deeply canaliculate. (Colombia-
Magdalena.) 31. L. cuspidata.
36. Inflorescence puberulous to tomentose; flower exterior gray-tomentose;
petioles 3.0-7.0 mm long; terete or shallowly canaliculate towards base.
37. Rachis and branches of inflorescence tomentose; petioles usually bear-
ing two distinct glands. (Central America.) 32. L. sparsipilis.
37. Rachis and branches of inflorescence puberulous; petioles eglandular.
(Guiana, Amazon.) 27. L. apetala.
26. Leaf undersurface with stomatal cavities or very deeply cut reticulation resembling
stomatal cavities, lanate-pubescent in mouth of cavities.
38. Inflorescence with secondary branches, flowers on distinct secondary branches of
inflorescence which are at least 2.0 mm long.
32 Flora Neotropica

39. Flowers in small distinct groups or in cymules on short secondary branches

(peduncles); leaf upper surface smooth, the base rounded to cuneate.
40. Inflorescence 15.0-22.0 cm long, ferrugineous-arachnoid; leaf underside brown
when dry. (Guiana.) 35. L. persaudii.
40. Inflorescence 6.o-8.0 cm long, gray-arachnoid or puberulous; leaf underside
gray when dry. (Guiana, Amazon.) 36. L. sprucei.
39. Flowers more or less clustered in large groups on primary and secondary branches
of inflorescence, but without distinct peduncles; leaf upper surface usually papillose
base often cordate. (Brazil-Amazon and south.) 37. L. sclerophylla.
38. Inflorescence with primary branches only; flowers on primary branches of inflorescence,
sessile or rarely with short peduncles not exceeding I.0 mm long.
41. Pedicels 2.0 mm long (in bud), to 4.0 mm in flower. (Guiana.) 38. L. albiflora.
41. Pedicels not exceeding 2.0 mm, often shorter.
42. Leaf reticulations extremely prominent on upper surface; fruit exterior appressed
sordid-yellow-velutinous.
43. Petioles 5.0-13.0 mm long; flowers ca 3.0 mm long; borne on slender branch-
lets 0.5-1.0 mm thick; receptacle campanulate; flowers and inflorescence
gray-tomentose. (Guiana, Amazon.) 39. L. longistyla.
43. Petioles 15.0-20.0 mm long; flowers ca 4.0 mm long, borne on branchlets
ca 2.0 mm thick; receptacle cupuliform; inflorescence and flowers brown-
tomentose. (Colombia, Pacific.) 40. L. fuchsii.
42. Leaf reticulations prominulous on upper surface; fruit exterior glabrous or
brown-tomentose.
44. Young branches densely tomentellous, with a thick corky bark. (Central
Brazil.) 41. L. humilis.
44. Young branches glabrous or nearly so, the bark thin.
45. Flowers 3.5-5.0 mm long; leaf apex mucronate. (Guiana.)
42. L.foveolata.
45. Flowers 2.0-3.0 mm long; leaf apex rounded to acuminate. (Venezuela,
Guiana, Amazon, Central Brazil.) 43. L. octandra.
i. Stamens included (filaments not exceeding calyx lobes or only very slightly exceeding them by
ca 0.2 mm).
46. Stamens 10-25. (For contrasting statement see p 33).
47. Petals present.
48. Ovary inserted at base of receptacle; bracts and bracteoles small and not enclosing
groups of flower buds. Subgenus MoquileaSection Microdesmia.
49. Leaf underside with stomatal cavities or deeply cut reticulation; filaments connate
for half of length in groups, densely pubescent.
50. Pubescence of inflorescence light gray; primary leaf veins prominulous above.
(northeastern Brazil.) 44. L. rigida.
50. Pubescence of inflorescence ferrugineous; primary leaf veins slightly impressed
above. (Central America, western South America.) 45. L. arborea.
49. Leaf underside with inconspicuous venation, stomatal cavities absent; filaments
free almost to base, glabrous above.
5I. Leaves 15.0-36.0 cm long; stipules 12.0-22.0 mm long; inflorescences on woody
defoliated stems 5.0- 1.0 mm thick. (Colombia-Valle.) 46. L. velata.
5I. Leaves under 14.0 cm long; stipules 2.0-Io.o mm long; flowers in terminal
inflorescences on young branchlets.
52. Inflorescence of unbranched subspikes, the flowers in small clusters along
rachis. (Colombia-Boyaca.) 47. L. subarachnophylla.
52. Inflorescence of panicles, or panicle of cymules.
53. Leaf underside densely lanate-pubescent.
54. Leaf apex with a well developed acumen 3.0-5.0 mm long; stamens
ca 25; exterior of receptacle ferrugineous. (Colombia-Antioquia.)
48. L. salicifolia.
54. Leaf apex obtuse to broadly acuminate, acumen not exceeding .0 mm;
stamens ca 15; exterior of receptacle brown. (Brazil-Goiis.)
49. L. araneosa.
53. Leaf underside glabrous.
55. Inflorescence a panicle of small cymules along the rachis and short
secondary branches; petioles 2.0-5.0 mm long; stamens 8-io. Species
of Old World. (Sect. Hymenopus.) 147. L. splendens.
55. Inflorescence of panicles, flowers not grouped in cymules; petioles
6.0-I I.0 mm long; stamens I4-25. Plants of New World.
Licania 33

56. Leaf acumen 3.0-7.0 mm long; leaves 5.0-10.0 cm long; stamens

ca 25; flowers sessile on primary branches of inflorescence, densely
brown-tomentose. (Southern Brazil.) 50. L. sylvatica.
56. Leaf acumen 10.0-15.0 mm long; leaves 7.5-14.0 cm long; stamens
ca I4; flowers predominantly on short secondary branches of in-
florescence; short gray-tomentose. (Colombia-Valle.)
5I. L. chocoensis.
48. Ovary inserted laterally; bracts and bracteoles large and enclosing small groups of
flower buds. (Guiana, Amazon.) 52. L. licaniaeflora.
Subgenus MoquileaSection Parinariopsis.
47. Petals absent.
57. Leaf underside glabrous, or with an easily removed lanate pubescence, the reticulations
shallow, the stomatal cavities absent. Subgenus MoquileaSection Leptobalanus.
58. Leaf underside glabrous; flowers in groups on distinct secondary branches of
inflorescence; stamens Io- I. (Amazon.) 33. L. emarginata.
58. Leaf underside lanate; flowers sessile on primary branches of inflorescence; stamens
ca 22. (Colombia-Valle.) 34. L. calvescens.
57. Leaf underside with stomatal cavities or with deep reticulations, pubescence dense and
filling the hollows between venation. Subgenus Licania Section Licania.
59. Receptacle conical; petioles usually canaliculate, 8.o-I3.0 mm long; midrib im-
pressed on upper surface towards lamina base; leaves drying light brown. (Guiana.)
108. L. majuscula.
59. Receptacle barrel-shaped; petioles terete, 3.0-6.0 mm long; midrib prominulous
at base but becoming impressed on upper portion; leaves drying dark brown.
(Amazon, Venezuela.) I40. L. mollis.
46. Stamens 3-8(-9). Subgenus Licania.
60. Leaf underside glabrous or sparsely hirsute along venation; petals present or absent. (For
contrasting statement see p 35).
61. Petals present (For contrasting statement see p 34).
62. Leaves sparsely hirsute along venation of lower surface. Section Hirsuta.
63. Inflorescence and flowers sparsely hirsutulous; primary veins and venation
deeply impressed on upper surface of lamina, appearing deeply rugose.
64. Leaf apex acuminate, leaves oblong. (Amazon.) 53. L. hirsuta.
64. Leaf apex rounded to obtuse; leaves ovate to ovate-elliptic. (Central America.)
54. L. costaricensis.
63. Inflorescence and flowers densely pubescent; primary veins plane or impressed,
and venation more or less plane on upper surface of lamina, never rugose.
65. Inflorescence and exterior of flowers with a short gray pubescence. (Guiana,
Amazon.) 69. L. heteromorpha.
65. Inflorescence and exterior of flowers densely ferrugineous-tomentose.
66. Leaf apex acuminate, leaves membraneous. (Bolivia, Amazon.)
55. L. krukovii.
66. Leaf apex rounded to apiculate, leaves thick-coriaceous.
67. Primary leaf veins 6-9, prominulous above; petioles 3.0-4.0 mm long.
(Guiana.) 56. L. lasseri.
67. Primary leaf veins 12-20, slightly impressed above; petioles 7.0-15.0
mm long. (Amazon.) 57. L. latifolia.
62. Leaf underside glabrous. Section Hymenopus.
68. Flowers in small cymules on short distinct secondary branches of inflorescence
at least I.0 mm long.
69. Leaves with a finely pointed acumen 15.0-30.0 mm long; flowers ca I.0 mm
long; ovary and style pubescent. (Colombia-Valle.) 58. L. minuscula.
69. Leaves rounded or acute to broadly acuminate (the acumen 2.0-8.8 mm
long); flowers ca 2.0 mm long; ovary and style glabrous or pubescent.
70. Inflorescence compact and triangular, to 8.o mm long; rachis not lenti-
cellate; peduncles 2.0-4.0 mm long; exterior of flowers glabrous. (Central
America) 59. L. operculipetala.
70. Inflorescence lax and spreading, over 8.o cm long, or short lax and of small
cymules on central rachis; rachis and branches puberulous, the rachis
often lenticellate; peduncles 0.5-2.0 mm long; exterior of flowers puber-
ulous.
71. Inflorescence of small cymules on central rachis 1.5-10.0 cm long;
stamens 8-io; receptacle oblique in bud; mature fruit to 1.5 cm long
34 Flora Neotropica

usually smaller, not costate. Species of Old World. (Asia and Pacific.)
146. L. splendens.
71. Inflorescence lax and spreading, over 8.o cm long; stamens 5-7;
receptacle symetrical in bud; mature fruit 1.5-4.0 cm long, costate.
Species of New World.
72. Leaf apex rounded; reticulation intricate. (Guiana, Amazon.)
69. L. heteromorpha.
72. Leaf apex acuminate; reticulation lax. (Amazon.)
60. L. reticulata.
68. Flowers borne mainly on primary branches of inflorescence or on secondary
branches, but not in pedunculate cymules.
73. Leaves narrowly oblong with nearly parallel sides; stamens usually slightly
exceeding calyx lobes; mouth of receptacle filled by a dense lanate mass; petals
slightly clawed.
74. Receptacle narrowly urceolate, densely arachnoid-pubescent; flowers in
dense glomerules on primary branches. (Guiana.) 61. L. arachnoidea.
74. Receptacle campanulate, puberulous; flowers not densely glomerulate.
75. Flowers ca 1.5 mm long; leaves to 15.0 cm long, usually smaller;
stipules to 5.0 mm, caducous. (Amazon.) 62. L. oblongifolia.
75. Flowers ca 3.0 mm long; leaves usually exceeding i6.0 cm long (to 38.0
cm); stipules to I5.0 mm, subpersistent. (Guiana, Brazil-Pari.)
63. L. macrophylla.
73. Leaves ovate to oblong-lanceolate, but sides converging; stamens shorter than
calyx lobes; mouth of receptacle with short deflexed hairs only; petals with
broad simple bases.
76. Rachis and branches of inflorescence and exterior of flowers glabrous or
sparsely hirsutulous.
77. Leaf apex caudate. (Brazil-Amazon.) 64. L. caudata.
77. Leaf apex acute to acuminate.
78. Stipules large and foliaceous, caducous; exterior of flowers glabrous.
(Venezuela.) 65. L. latistipula.
78. Stipules small, lanceolate, persistent; exterior of flowers hirsutulous.
79. Leaves 9.0-27.0 cm long, thick-coriaceous, apex abruptly short-
acuminate; primary veins plane above. (Guianas.)
66. L. divaricata.
79. Leaves 7.0-I I.0 cm long, membraneous, apex with a well de-
veloped acumen 5.0-9.0 mm long; primary veins slightly im-
pressed above. (Venezuela, Brazil-Para.) 67. L. glabrifora.
76. Exterior of flowers and usually rachis and branches of inflorescence
densely puberulous to tomentose, pubescence completely covering exterior
of calyx.
80. Midrib broad towards base, 2.0-3.5 mm thick; leaves very thick-
coriaceous; stipules to 15.0 mm long, subpersistent; anthers deltoid or
nearly so. (Venezuela.) 68. L. intrapetiolaris.
80. Midrib narrower towards base, I.0-2.0 mm thick; leaves membraneous
to coriaceous; stipules to 8.0 mm long, persistent to caducous; anthers
reniform. (Guiana, Amazon.) 69. L. heteromorpha.
61. Petals absent. Section Hymenopus.
81. Leaves thin and membraneous, bases cuneate, the venation equally prominent on
both surfaces; receptacle urceolate. (E.-central Brazil.) 70. L. glazioviana.
81. Leaves thick and coriaceous, base usually rounded to cordate, rarely rounded-
subcuneate, the venation obscure on upper surface; receptacle conical to globose-
cupuliform.
82. Leaves rounded to obtuse at apex, rarely exceeding 9.0 cm long.
83. Flowers ca 3.0 mm long; petioles terete. (Eastern Brazil.) 71. L. littoralis.
83. Flowers ca 2.0 mm long; petioles usually canaliculate.
84. Inflorescence and flowers with brown pubescence; stamens 3; venation of
lower surface of leaf often papillose; stipules intrapetiolar. (Guiana.)
72. L. fanshawei.
84. Inflorescence and flowers gray-puberulous; stamens 5; venation of leaf
lower surface smooth-papillose; stipules adnate to extreme base of
petiole. (Guianas.) 73. L. irwinii.
82. Leaves distinctly acuminate at apex, usually exceeding 8.o cm long.
Licania 35

85. Midrib slightly impressed above; petioles canaliculate. (Guiana.)

74. L. cyathodes.
85. Midrib prominulous above, petioles terete.
86. Flowers 2.5-3.5 mm long; ferrugineous-brown-pubescent; stipules 4.0-7.0
mm long, persistent. (Amazon.) 75. L. polita.
86. Flowers 1.5 mm long, gray-brown-pubescent; stipules small, caducous.
(Brazil-Para.) 76. L. silvae.
60. Leaf underside pubescent, densely arachnoid-lanate or pulverulent (not hirsute); petals
absent.
87. Flowers in small cymules, on long slender secondary branches (peduncles) less than
0.5 mm thick and attached to primary inflorescence branches; pedicels usually 0.25-
3.0 mm long; rarely absent; fruit often very small usually not exceeding 2.0 cm long.
(For contrasting statement see p 36). Section Cymosa.
88. Leaf underside with stomatal cavities or thick, coarse, deeply cut venation below,
lanate-arachnoid.
89. Bracteoles large and enclosing groups of flower buds; flowers subsessile; peduncle
short and rather thick. (Guiana.) 77. L. densiflora.
89. Bracteoles small and not enclosing groups of flower buds; flowers pedicellate;
peduncles long and slender.
90. Inflorescence and exterior of flowers ferrugineous-tomentose; young stems
ferrugineous-tomentellous; fruit densely tomentose, 3.0-5.0 mm long, stipi-
tate. (Guiana.) 78. L. cuprea.
90. Inflorescence and exterior of flowers gray-puberulous or gray-tomentose;
young stems glabrous or puberulous; fruit I.0-2.0 mm long, not stipitate.
9I. Midrib impressed for entire length; petioles 7.0-14.0 mm long; leaves
oblong-lanceolate. (Amazon.) 79. L. impressa.
9I. Midrib plane throughout or impressed at base only; petioles 2.0-7.0 mm
long; leaves ovate to oblong.
92. Leaf underside with lanate pubescence obscuring venation; stamens
6-7. (Southern Brazil.) 80. L. dealbata.
92. Leaf underside pubescent only in the mouth of stomatal cavities;
reticulations glabrous; stamens 3-5.
93. Leaves ovate, broadest near base, 3.o-8.5 cm long; stipules axillary.
(Amazon.) 88. L. hypoleuca.
93. Leaves oblong-elliptic, broadest about middle, 5.o-I3.0 cm long;
stipules adnate to petiole base. (Amazon, Guiana.) 8I. L. pallida.
88. Leaf underside with a very fine, plane or prominulous venation, usually pulverulent-
farinaceous.
94. Exterior of flowers and branches of inflorescence entirely glabrous or glabres-
cent; fruit exterior often drying purple.
95. Leaves oblong-lanceolate to oblong, the lower surface white-lanate. (Amazon.)
82. L. gracilipes.
95. Leaves ovate to oblong, lower surface sparsely gray-pulverulent-farinaceous.
(Guiana, Amazon.) 83. L. parvifructa.
94. Exterior of flowers and usually branches of inflorescence puberulous to tomen-
tose; fruit exterior gray to brown.
96. Leaf apex rounded; margins revolute; inflorescence glabrous. (Brazil-
Bahia.) 84. L. cymosa.
96. Leaf apex acuminate; margins not revolute; inflorescence usually puberulous
or tomentose.
97. Fruit elongate-pyriform, to 2.5 cm long, the exterior with short rufous-
velutinous pubescence; either inflorescence and exterior of flowers brown-
tomentellous or petioles 7.0-12.0 mm long.
98. Petioles 3.0-6.0 mm long, terete; inflorescence and flowers brown-
tomentellous; leaves triangular or nearly so. (W. T.ldies.)
85. L. ternatensis.
98. Petioles 7.0-I2.0 mm long, canaliculate; inflorescence and flowers
gray-puberulous; leaves oblong. (Trinidad, Guiana, Amazon.)
86. L. membranacea.
97. Fruit ovoid to pyriform, rarely exceeding 1.2 cm long, the exterior with
short sordid-pulverulent indumentum; inflorescence and exterior of flowers
gray-puberulous or glabrescent; petioles 3.0-6.0 mm long.
99. Leaves 6.5-I6.o cm long; upper surface of midrib and primary veins
36 Flora Neotropica

prominent; rachis and branches of inflorescence glabrescent. (Brazil-

Amapa.) 87. L. piresii.
99. Leaves 2.5-I0.0 cm long, upper surface of midrib plane or prominu-
lous; rachis and branches of inflorescence puberulous. (Central
America, Colombia, Venuzeula, Guiana, Amazon.) 88. L. hypoleuca.
87. Flowers sessile or subsessile on primary branches of inflorescence or on short (less than
0.5 mm) thick peduncles only. Flowers usually sessile; fruit rarely less than 2.0 cm long.
1oo. Leaf underside pulverulent-farinaceous. Section Pulverulenta.
IoI. Leaves with rounded to acute apex, the margins usually revolute.
o02. Flowers 1.5-2.0 mm long; receptacle campanulate-cupuliform.
103. Young branches and inflorescence densely tomentose; inflorescence to
4.0 cm long, recurved. (Guiana.) 89. L. boyanii.
103. Young branches and inflorescence branches glabrous; inflorescence
usually exceeding 4.0 cm long, erect. (Guiana.) 90. L. buxifolia.
102. Flowers ca 3.0 mm long; receptacle urceolate.
I04. Leaves predominantly orbicular, occasionally oblong-elliptic, the apex
rounded to retuse. (Amazon.) 91. L. orbicularis.
104. Leaves elliptic, the apex acute.
105. Receptacle narrowly urceolate-cylindrical; calyx lobes lanceolate.
(Brazil-Rond6nia.) 92. L. niloi.
o05. Receptacle broadly urceolate; calyx lobes deltoid. (Guiana, Amazon.)
93. L. coriacea.
o10. Leaves with well developed acumen, the margins not revolute.
Io6. Flowers 3.0 mm long; receptacle urceolate.
107. Venation of leaf underside minutely reticulate, forming a network less
than 0.25 mm between spaces; reticulation apparent because of absence
of pubescence from nerves; leaves thin-chartaceous; stipules usually
caducous. (Amazon.) 94. L. urceolaris.
107. Venation and reticulation coarse, with I.0-2.0 mm between reticula-
tions; pubescence obscuring much of veins; leaves thick-coriaceous;
stipules persistent.
o08. Leaves 7.0-I5.0 cm long, the apex with finely pointed, well developed
acumen; rachis of inflorescence glabrous. (Guiana, Brazil-Para.)
95. L. affinis.
Io8. Leaves 4.0-9.0 cm long, the apex acute or with short blunt acumen;
rachis of inflorescence pubescent. (Guiana, Amazon.) 93. L. coriacea
io6. Flowers 1.5-2.0 mm long; receptacle campanulate.
o19. Primary veins slightly impressed on upper surface; fruit exterior sordid-
rufous-pubescent; branches of inflorescence densely tomentose to puberu-
lous.
I I. Leaves thick-coriaceous; stipules caducous; flowers in clusters on
short thick peduncles; stamens 3. (Colombia-West coast.)
96. L. glauca.
IIo. Leaves thin, chartaceous-membraneous; stipules persistent; flowers
on primary and secondary branches of inflorescence; stamens 6-7.
(Guiana, Brazil-Par,.) 97. L. davillaefolia.
109. Primary veins plane or prominent on upper surface; fruit exterior glab-
rous, drying yellow; branches of inflorescence glabrous to puberulous.
I I I. Leaves (I I-) I3.0-18.0 cm long; 4.5-7.5 cm broad. (Guiana, Amazon.)
98. L. elliptica.
I I . Leaves 4.0-10.0 (-I2.0) cm long. (Guiana, Amazon.) 99. L. canescens.
Ioo. Leaf underside densely lanate-arachnoid or with stomatal cavities, never pulveru-
lent. Section Licania.
II2. Flowers 6.0-7.0 mm long; stamens connate for half their length. (Guiana.)
Also L. naviculistipulaPrance, see supplement. IOO.L. couepifolia.
I2. Flowers not exceeding 5.5 mm long; stamens free almost to base.
I 13. Leaf base distinctly cordate or subcordate; leaves usually ovate-orbicular.
114. Leaves triangular-ovate, io.o-I6.ocm long, membraneous. (Peru-
Loreto.) IoI. L. trigonioides.
114. Leaves orbicular to ovate, 3.0-9.0 cm long, usually coriaceous.
115. Young stems hispid; lower surface of leaves with hirsutulous-hispid
venation, the apex with well developed acumen. (Venezuela-
Amazonas.) 102. L. cordata.
Licania 37

I 5. Young stems puberulous to tomentose; lower surface of leaves glab-

rous or lanate on venation, the apex acute or bluntly acuminate.
I16. Flowers 4.5-5.5 mm long; receptacle urceolate; stipules 5.0-6.0
mm long. (Venezuela-Amazonas.) 103. L. foldatsii.
I 6. Flowers 1.5-3.5 mm long; receptacle campanulate; stipules I.o-
3.0 mm long.
117. Leaves submembraneous, the lower surface with deeply cut vena-
tion; pubescence occuring in cavities and hard to remove;
fertile stamens 5-9.
1 8. Stamens 8-I I; petioles ca 5.0 mm long; stipules 3.0-6.0 mm
long. (Amazon, Venezuela.) I40. L. mollis.
118. Stamens 5-6; petioles 1.5-3.0 mm long; stipules 2.0-2.5 mm
long. (Colombia-Amazonas.) Io4. L. hebantha.
II7. Leaves thick-coriaceous, the lower surface with shallow vena-
tion, not forming cavities, lanate pubescence easily rubbed off;
fertile stamens 3.
119. Pubescence obscuring venation; stipules adnate to extreme
base of petiole. (Venezuela-Bolivar.) Io5. L. steyermarkii.
19. Pubescence not obscuring venation; stipules axillary.
(Venezuela.) o6. L. subrotundata.
13. Leaf base rounded to cuneate, leaves only rarely ovate-orbicular.
120. Midrib and primary veins distinctly impressed on leaf upper surface.
21. Leaf underside with well developed stomatal cavities filled with lanate
pubescence; petioles 8.0- 7.0 mm long.
122. Stomatal cavities conspicuous because of glabrous nerves and
veins: leaves 4.0-8.5 cm long. (Venezuela-Amazonas.)
107. L. crassivenia.
122. Stomatal cavities less conspicuous because of puberulous nerves
and venation; leaves (8.o-)10.0-25.0 cm long.
I23. Receptacle conical, 4.0-5.0 mm long; fruit tomentellous, the
stipe 2.0-6.0 mm long; leaf underside brown-lanate. (Guiana.)
108. L. majuscula.
123. Receptacle campanulate, 2.5-3.0 mm long; fruit pulverulent,
the stipe 8.0-x5.0 mm long; leaf underside white-lanate.
(Guiana.) 09. L. alba.
121. Leaf underside without stomatal cavities but often prominently reti-
culate; petioles 2.0-6.0(-8.0) mm long.
I24. Exterior of receptacle sparsely puberulous, not completely cover-
ing the surface; leaves oblong-lanceolate. (Guiana.)
114. L. kunthiana.
124. Exterior of receptacle densely tomentose to tomentellous, com-
pletely covering the surface; leaves oblong to elliptic.
125. Leaves 3.0-5.0 cm long, not prominently reticulate beneath;
petioles 1.0-3.0 mm long. (Venezuela-Bolivar.)
I o. L. hitchcockii.
125. Leaves 5.-I5.0 cm long, prominently reticulate beneath;
petioles 4.0-8.0 mm long.
i26. Flowers ca 4.0 mm long; receptacle broadly cupuliform;
leaves ovate-orbicular with rounded apex, underside hirsute
along venation. (Guiana.) II. L. sandwithii.
126. Flowers 2.0-3.0 mm long; receptacle campanulate; leaves
oblong to elliptic, acute to acuminate at apex; underside
usually tomentellous rarely hirsute on venation.
127. Leaf undersurface with parallel secondary veins, giving
prominently reticulate appearance; pubescence brown;
primary veins 7-I0. (Guiana, Amazon.) 112. L. laxiflora.
127. Leaf undersurface with diffuse secondary veins; pubes-
cence rufous; primary veins 5-6. (Guiana.)
I 13. L. rufescens.
120. Primary veins and usually midrib plane or prominent on upper surface.
128. Exterior of flowers and inflorescence branches gray-puberulous, not
completely covering surface.
129. Receptacle campanulate; leaves without stomatal cavities.
(Guiana, Brazil.) 14. L. kunthiana.
38 Flora Neotropica

129. Receptacle urceolate; leaves with shallow stomatal cavities.

(Brazil-Rondonia.)
115. L. bellingtonii.
128. Exterior of flowers densely tomentellous or tomentose, completely
covering surface.
I30. Leaf apex rounded, obtuse or bluntly acute, or rarely short-
apiculate (never acuminate.)
131. Lanate pubescence of leaf underside hard to remove, covering
deep stomatal cavities; flowers 3.o-5.o mm long; receptacle
urceolate. (Guiana.) i6. L. compacta.
131. Lanate pubescence of leaf underside easily removed, revealing
little protuding venation, stomatal cavities absent; flowers less
than 3.0 mm long; receptacle usually campanulate.
132. Receptacle urceolate; stipules 4.0-5.0 mm long, adnate to
petiole well above base. (Guiana.) ii7. L. ovalifolia.
132. Receptacle usually campanulate, rarely urceolate; stipules
1.0-3.0 mm long, axillary or adnate to extreme base of
petiole.
133. Leaves ovate-orbicular, the apex often shortly apiculate.
134. Late pubescence obscuring venation; stipules adnate
to extreme base of petiole. (Venezuela-Bolivar.)
105. L. steyermarkii.
134. Lanate pubescence not obscuring venation; stipules
axillary. (Venezuela.) io6. L. subrotundata.
I33. Leaves oblong-lanceolate to elliptic.
135. Leaves oblong-elliptic, the underside minutely reti-
culate; primary veins Io-II; petioles 4.0-7.o mm
long; receptacle campanulate. (Colombia.)
i 8. L. caldasiana.
135. Leaves oblong to oblong-lanceolate, the underside
densely lanate with obscured venation; primary veins
7-9; petioles 2.0-3.0 mm long; receptacle campanulate-
urceolate. (Brazil, Venezuela-Amazonas.)
19. L. savannarum.
130. Leaf apex distinctly acuminate or acute.
136. Leaves 3.0-4.0 mm long; with two conspicuous glands at
junction of petiole with upper surface of lamina. (Guiana.)
120. L. microphylla.
I36. Leaves predominantly larger, lacking such conspicuous glands.
I37. Stamens 3.
138. Leaf undersurface with poorly developed stomatal cavities
filled by lanate pubescence, the venation glabrous or
glabrescent and hence conspicuous; glands present at
base of lower surface. (Brazil, Colombia-Amazon.)
121. L. triandra.
138. Leaf undersurface with prominent venation but no stoma-
tal cavities, the venation pubescent and hence less conspicu-
ous.
139. Stipules caducous, 0.5-I.5(-2.0) mm long; petioles often
tomentose; receptacle campanulate.
140. Petioles tomentose when young; leaves elliptic to
oblong, with well developed acumen; primary veins
7-9 pairs. (Guiana.) 122. L. discolor.
140. Petioles glabrous; leaves oblong-lanceolate, finely
apiculate; primary veins 10-12 pairs. (Amazon.)
123. L. apiculata.
I39. Stipules persistent, 2.0-3.0 mm long; petioles glabrous
or puberulous when young; receptacle campanulate or
urceolate.
I41. Inflorescence 8.o-I5.0 cm long, spreading, with
numerous primary branches; lower leaf surface
brown-lanate; receptacle campanulate. (Guiana,
Amazon.) 124. L. micrantha.
141. Inflorescence 5.o-6.o cm long, with a few primary
Licania 39

branches; lower leaf surface whitish-gray-pubescent;

receptacle urceolate.
142. Leaf apex with a finely pointed acumen; leaves
thin-membraneous. (Guiana.) 125. L. pruinosa.
142. Leaf apex acute to bluntly acuminate; leaves
thick-coriaceous. (E.-central Brazil.)
126. L. nitida.
I37. Stamens 4-8(-I0).
143. Leaf underside with deep, extremely conspicuous stomatal
cavities, the pubescence confined to cavities made obvious
by almost glabrous venation.
144. Inflorescence and flowers ferrugineous-pubescent;
stipules 3.0-4.0 mm long x I.o-I.5 mm at base,
persistent, ferrugineous; leaves acute to bluntly acumi-
nate. (E.-central Brazil.) I27. L. riedelii.
I44. Inflorescence and flowers gray-puberulous; stipules
i.0-3.0 mm long, 0.2-0.5 mm at base, persistent or
caducous, pubescent but not ferrugineous; leaves with
well developed acumen.
145. Bracteoles persistent, lanceolate; upper surface of
midrib impressed; stipules caducous. (Amazon.)
128. L. bracteata.
I45. Bracteoles caducous, triangular; upper surface of
midrib level; stipules small but persistent.
146. Inflorescence branches thick; flowers subsessile;
fruit exterior ferrugineous-velutinous. (Vene-
zuela, Amazon.) 129. L. parviflora.
146. Inflorescence branches slender; flowers distinctly
pedicellate; fruit exterior reddish brown short-
pulverulent. (Venezuela, Brazil-Amazonas.)
8i. L. pallida.
143. Leaf underside with poorly developed stomatal cavities or
none; venation pubescent.
I47. Petioles 1.5-2.0 cm long; inflorescence rachis and
branches glabrous or glabrescent. (Guiana, Brazil-
Para.) 130. L. robusta.
147. Petioles to I.ocm long; inflorescence rachis and
branches usually densely tomentose or tomentellous.
148. Inflorescence predominantly of axillary and terminal
spikes, terminal inflorescence rarely little branched
or with minute spikes along rachis.
149. Leaves lanceolate, lower surfacedeeply reticulate,
with poorly developed stomatal cavities. (Vene-
zuela-Amazonas.) 13 . L. lanceolata.
I49. Leaves oblong to ovate-elliptic, lower surface
not deeply reticulate, lacking stomatal cavities.
I50. Flowers 2.5 mm long; receptacle broadly
cupuliform; inflorescence puberulous; reti-
culate venation of leaf underside conspicuous.
(E.-central Brazil.) 132. L. spicata.
I50. Flowers 1.5-2.0 mm long; receptacle campan-
ulate; inflorescence tomentose; venation of
leaf underside inconspicuous.
151. Flowers in short dense minute spikes at-
tached to long rachis. (Guiana.)
I33. L. stricta.
151. Flowers solitary along rachis or in dense
glomerules, but not in minute spikes.
152. Leaves thin and membraneous, the
acumen finely pointed; flowers in dense
glomerules; inflorescences largely axil-
lary. (Guiana, Brazil-Para)
I34. L. leptostachya.
40 Flora Neotropica

152. Leaves thick and coriaceous, the acumen

usually blunt; flowers in small groups
or solitary but not glomerulate; in-
florescence largely terminal. (Guiana,
Amazon.) 135. L. incana.
148. Inflorescence of terminal and sub-terminal race-
mose panicles.
I53. Leaves lanceolate; low shrub or suffrutex.
(Venezuela-Amazonas.) 13 . L. lanceolata.
153. Leaves ovate to oblong-lanceolate; trees or high
shrubs.
154. Stipules 3.0-7.0 mm long and at least I.0 mm
broad at base; distinctly adnate to petiole or
intrapetiolar, persistent and obvious.
155. Midrib distinctly impressed above; brac-
teoles o.2-I.o mm long.
I56. Inflorescence ferrugineous-pubescent;
petioles canaliculate; stipules adnate to
petiole well away from axil. (Brazil-
Para.) 136. L. paraensis.
I56. Inflorescence gray-puberulous; petioles
terete; stipules intrapetiolar or adnate
to inside of extreme base of petiole.
157. Leaves with a finely pointed acumen;
primary veins 11-12 pairs, lower
surface glandular at base. (Colom-
bia-Amazon.) I37. L. vaupesiana.
157. Leaves with blunt acumen; primary
veins 6-8 pairs; lower surface eglan-
dular at base. (E.-central Brazil.)
I38. L. bahiaensis.
155. Midrib not distinctly impressed above;
bracteoles 1.5-2.0 mm long,
158. Leaves membraneous; flowers ca 2.0
mm long, gray-tomentellous; inflores-
cence gray-puberulous; stamens 5.
(Brazil-Amap,.) I39. L. maxima.
158. Leaves coriaceous; flowers 3.0-3.5 mm
long, brown-tomentose; inflorescence
brown-tomentose; stamens 8-I I. (Brazil,
Venezuela-Amazon.) I40. L. mollis.
I54. Stipules usually less than (rarely exceeding)
2.5 mm long, and very thin to base, on out-
side of axils or adnate to extreme base of
petiole, caducous or persistent, often incon-
spicuous.
159. Stipules adnate to extreme base of petiole,
persistent or subpersistent.
I60. Petioles 7.0-12.0 mm long, remaining
tomentellous even on older leaves;
laminas 7.0-I6.0 cm long, 3.0-7.5 cm
broad; stipules subpersistent. (Brazil-
Pari and central.) 141. L. blackii.
I60. Petioles 2.0-5.0 mm long, becoming
glabrous with age; laminas 3.0-12.0
cm long, 2.2-5.5 cm broad; stipules per-
sistent.
I6I. Inflorescence lax and spreading;
leaves with a finely pointed acumen
8.0-20.0 mm long; lower surface
farinaceous-lanate, shallowly reti-
culate. (Guiana, Amazon.)
129. L. parviflora.
Licania 41

I6I. Inflorescence and flowers densely

crowded; leaves with blunt acumen
2.0-I3.0 mm long, lower surface lan-
ate, deeply reticulate. (Guiana to E.-
central Brazil.) I 14. L. kunthiana.
i59. Stipules axillary, caducous or persistent.
162. Leaf undersurface with smooth incon-
spicuous reticulation, the pubescence
easily removed; primary veins widely
spaced 1.2-2.5 cm apart; exterior of
receptacle velutinous-pubescent. (Brazil-
Amazon.) 142. L. rodriguesii.
162. Leaf underside with deeply cut reticu-
lation, and hence pubescence hard to
remove; primary veins not more than
I.0 cm apart; exterior of receptacle
tomentellous.
163. Receptacle globose; upper surface
of youngest leaves appressed-strigose,
soon becoming glabrous;inflorescence
of axillary spikes and terminal
panicles. (E.-central Brazil.)
132. L. spicata.
163. Receptacle usually campanulate;
upper surface of youngest leaves
glabrous; inflorescence of terminal
and subterminal racemose panicles.
I64. Leaves oblong-lanceolate; mid-
rib slightly impressed on upper
surface. (E.-central Brazil.)
143. L. indurata.
I64. Leaves ovate-elliptic to oblong;
midrib usually plane on upper
surface.
I65. Inflorescence much branched,
spreading, lax; lower leaf sur-
face with rufous pubescence.
(Guiana.) I 3. L. rufescens.
I65. Inflorescence densely crowded;
compact, little-branched; lower
leaf surface gray-brown-pubes-
cent.
i66. Petioles 4.o-7.o mm long,
terete; receptacle and calyx
lobes tomentose on exterior.
167. Petioles soon becoming
glabrous; leaf apex
bluntly acuminate;
young fruit pyriform.
(E.-central Brazil.)
I44. L. hoehnei.
167. Petioles tomentose, be-
coming less so with age;
leaf apex obtuse to acute;
young fruit cylindrical
when young, becoming
pyriform. (Trinidad, Ve-
nezuela.) I45. L. cruegeriana.
i66. Petioles Io.o-I2.0 mmlong,
canaliculate; receptacle and
calyx lobes velutinous on
exterior. (Brazil-Bahia.)
I46. L. belemii.
42 Flora Neotropica

i. Licania subgen Moquilea sect Moquilea

i. Licania michauxii Prance, Jour. Arnold Arb. 5I: 526. I970.

oblongifoliusMichaux, Fl. Bor.-Am. I: 283. 1803; Pursh, Fl. Am. Sept. I: 329. 1814;
Chrysobalanus
Nuttall, Gen. I: 301. I818; Elliot, Bot. S. Car. & Georgia I: 539. 1821; DC., Prodr. 2:
526. 1825; Torrey & Gray, Fl. N. Am. I: 406. I840; Chapman, Fl. So. U.S. II9. i86o;
Sargent, Silva 4: I. I892; C. Mohr, P1. Life Alab. 553. I901; Small, Fl. Southeastern U.S.
569. 1903; Ahles et al, Rhodora 60: I6. I958. Type. Michauxsn fl (P).
ChrysobalanusretususRafinesque, New Fl. Am. 3: 26. 1836. Type. Nuttall (in Herb. Collins) sn,
U.S.A., fl (BM).
Geobalanusoblongifolius(Michaux) Small, Fl. Miami 81, 200. I913, Manual of Southeastern Fl.
646, 1933.
GeobalanuspallidusSmall, Fl. Miami 81, 200. 1913; Small, Manual of Southeastern Fl. 646. 1913.
Type. Small & Carter71I, U.S.A., Florida, fl (lectotype, NY).
pallidus (Small) L. B. Smith, Rhodora 48: I36. 1946.
Chrysobalanus
ChrysobalanusincanusRafinesque, New Fl. Am. 3: 26. 1836. Type. Michauxsn fl (P).

Low suffrutex to ca 40.0 cm tall, the young stems glabrous, not lenticellate.
Leaves lanceolate to oblong-lanceolate to rarely oblong-elliptic, serrulate, coriaceous,
4.0-I 1.0 cm long, . I-3.5 cm broad, acute to rounded or, rarely retuse at apex, often
mucronate, cuneate to subcuneate at base, glabrous or the lower surface gray-lanate-
arachnoid; midrib prominent and glabrous above; primary veins 8-13 pairs, promi-
nulous on both surfaces; reticulations conspicuous on both surfaces; petioles I.o-3.0
mm long, glabrous, terete, eglandular. Stipules 0.5-1.0 mm long, lanceolate, coriace-
ous, persistent or caducous, extrapetiolar. Inflorescences rather lax terminal and
subterminal cymose panicles, the rachis and branches puberulous. Flowers ca 2.5 mm
long, in small groups on long slender primary and secondary inflorescence branches.
Bracts and bracteoles minute, ovate, persistent, puberulous on exterior. Receptacle
campanulate, tomentose on exterior, tomentellous within; pedicels I.0-3.5 mm long.
Calyx lobes acute, tomentellous on both sufaces. Petals 5, densely pubescent. Stamens
I4-I7, inserted in a complete circle; filaments exceeding calyx lobes, connate at base,
glabrous. Ovary inserted at base of receptacle, glabrous or sparsely villous. Style
equalling filaments, glabrous except for pubescent base. Fruit ovoid, 2.5-3.0 cm
long; epicarp smooth, glabrous; pericarp thin, hard, fibrous, with no lines of de-
hiscence, sparsely pilose within.
TYPE. Michauxsn, U.S.A. Georgia, fl (holotype, P).
DISTRIBUTION.Pine barrens, sand dunes and oak scrubland of southeastern
United States. Flowering between December and August, especially from April to
June, fruiting June to September.
Representative collections: NORTH AMERICA. U.S.A., Louisiana: Cocks3132 fl (A). Missis-
sippi: Demaree33429 fl (GH); Earle sn fr (NY); Gerhartio fl (NY); Pollard Ir20 fl fr (F, GH, MO, NY,
US); Skehansn fl fr (A, MO, US); Underwoodsn fl (NY); Woodson& Schery80 fl (MO); Webster&
Wilbur3441 fl (GH, US). Alabama: C. F. Baker sn fr (MO, NY); Beaumontsn fl (GH); Bush 367 fr
(A, NY); Demaree35889 fl fr (GH); Drushel4649 fl (P); Gatessn fl (NY); Jewett sn fl (P); Mackenzie4038
fl (NY); Mohrsn fl (F, MICH, NY, US); Webster& Wilbur3547 fl fr (GH, US). Georgia: Beyrich132 fl
(BM); Boole 1141 fl (GH); Correll5494 fl (GH); Cronquist5332 fl (GH, NY, US); De Wolf 1779 fl (GH);
Graysn fl (K); R. M. Harper698 fl (BM, K, NY, US); 983 fl (F, GH, MO, NY, P, US); 1433 fr (F,
GH, MO, NY, US); Le Contesn fl (NY); McVaugh8634 fl (GH, MICH) ;J. K. Smallsn fl (A, F, K, NY);
Torreysn fl (K). South Carolina: Ahles & Bell 18217 fr (GH, NY); Le Contesn fl (P). Florida: Black 49-
4347 fl (NY); Britton274 fl fr (F, NY); Chapmansn fl (BM, GH, NY, US); Curtiss727 fl fr (A, BM, FI,
GH, K, MICH, NY); 4212 fl (NY, US); 5779 fr (A, GH, K, MO, NY, P); Ford 4416 fl (BM); Foster,
L. B., & D. L. Smith 1355 fl (BM, BR, F, GH, K, LA, LD, MICH, MO, NY, US); 1356 fl (BM, BR,
GH, K, LA, LD, MICH, NY, US); Godfrey53336 fl (GH, NY); 55531 fl (GH, NY); Lakela 24962 fl
(GH); Moldenke333 fl (K, MO, NY); 961 fl (K, NY, US); 5463 fl (NY); Rugelsn fl (BM, BR, K, NY);
Licania 43

217 fl (BM, K, NY, P); J. K. Smallet al 2479 st (NY); 3377 fl (NY); 3383 fl (NY); 6505 fl (NY); 6559
fr (GH, NY, US); 9988 fl (K); 11360 fl (NY); 11508 fr (NY);J. P. Standley 176fl (A, BM, F, GH, US);
Sunkill527 fl (K); 568 fl (K); Tracy7451 fl (A, BM, F, GH, MO, NY, US).

LOCALNAMES.Gopherapple, Groundoak.
Details of the nomenclature of this species have been elucidated elsewhere
(Prance, in press). Rafinesque (New Fl. Am. 3: 26. 1836) described another
species, Chrysobalanus prunifoliusRaf., which has commonly been referred to this
species. I have been unable to find type material of C.prunifolius,but judging from the
description it is not L. michauxii.The small leaves, the few-flowered racemes, and the
pod-like fruit of C.prunifoliusare all unlike L. michauxii.The specific epithets of all the
synonyms cited are already occupied in Licania. For these reasons the Rafinesque
name is not used, and a new name was proposed for the species commonly known as
Chrysobalanus oblongifolius.
Geobalanus(Chrysobalanus) pallidus is placed here in synonymy. The original
difference given by Small is not sound. Pubescence of both the ovary and the leaf
undersurfacevary from glabrous to lanate. In many cases the pubescence of the leaf
is caducous and the older leaves are glabrous. This was the only difference given by
Small to separate C. pallidusand C. oblongifolius. Although Small did not cite a type in
his description of C. pallidus,the specimen, Small & Carter7rI, at NY was annotated
by Small as the type and hence is chosen here as a lectotype.
Chrysobalanus incanusRafinesque, which also corresponds to the material des-
cribed as C. pallidus,was based on Michaux's original description of C. oblongifolius.
Rafinesque elevated a variety to specific rank without seeing the material at all. I
do not consider the material with pubescent leaf undersurfacesworthy of taxonomic
rank, because of the gradation in this character.
This species has often been said to have a stellate pubescence. Kiister (Bot.
Centralb. 69: I35-I37. I897) worked on the anatomy of the Chrysobalanaceae and
originally reported the stellate pubescence, which has led to the mention of stellate
pubescence in connection with the family in several other publications. Attribution
of a stellate pubescence to L. michauxiihas been recorded erroneouslyas the result of a
mixed gathering of herbarium material. The widely distributed herbarium material
of Curtiss727 contains, in addition to L. michauxii,sterile branches of Quercus pumila
Walt. This species of Quercusdoes have stellate pubescence on the lower leaf surface.
Q. pumilahas the same suffruticosehabit as L. michauxii,and could easily be mistaken
for it in the sterile condition, except for the pubescence.

2. Licania boliviensis Prance, sp nov Fig 6 E.

Arbor vel frutex, ramulis juvenilibus lanato-arachnoideis pubescentibus mox

glabris. Folia alternata petiolata, petiolo 5.0-7.0 mm longo, juvenile arachnoideo
mox tomentello, eglanduloso, leviter canaliculato; laminae oblongo-lanceolatae vel
oblongae, coriaceae, 7.0-I .0 cm longae, 2.0-4.0 cm latae, basi cuneatae, apice in
acuminem 2.0-5.0 mm longum contractae, supra glabrae subtus breviter lanato-
tomentosae; costa media supra prominente, glabra; costis secundariis I1-15 jugis,
subtus prominentibus, supra planis. Stipulae filamentosae ad 20.0 mm longae,
membranaceae, caducae. Flores circa 2.0 mm longi in paniculis racemosis terminali-
bus vel subterminalibus dispositi, haud pedunculati, rachi ramisque lanato-arach-
noideis. Bracteae bracteolaeque 1.0-2.5 mm longae, lanceolatae, flores excedentes,
membranaceae, caducae. Receptaculum globosum, sessile,extus lanato-arachnoideum
44 Flora Neotropica

pubescens, intus tomentosum. Calycis-lobi acuti. Petala 5, glabra. Stamina circa 25

in orbem completum disposita; filamenta glabra, exserta. Ovarium ad basim
receptaculi insertum, lanato-pubescens. Stylus e basi ovarii ortus, calycis-lobos
excedens, praeter basim glaber. Drupa globosa, extus crustacea, verrucosa, glabra;
pericarpio ad i.o cm lato, lignoso, fibroso, intus glabro.
TYPE. Krukoff11233, Boliva, La Paz Dept., Larecaja Province, io km. south of
Mapiri, fl bud, (holotype, NY; isotypes, A, F, LA, MICH, MO, S, US).
DISTRIBUTION. Known only from the Mapiri region of Bolivia, fruiting in
November.
BOLIVIA. La Paz Dept.: Buchtein1721 st (NY, US); Krukoffr1104 fr (A, F, LA, MAD, MICH,
MO, NY, S, US).

This species is most distinct from others in the section because of the oblong-
lanceolate leaves with short sordid-lanate pubescence on their lower surface. The
large fruit with a crustaceous exterior also distinguishes it from most species of the
genus. In addition, L. boliviensisoccurs outside the geographical range of most other
species of Licania.

3. Licania maritima Prance, sp nov Fig 5.

Arbor media; ramulis juvenilibus puberulis mox glabris. Folia afternata petio-
lata; petiolo I3.0-20.0 mm longo, tereti, eglanduloso, juvenilo tomentello; laminae
coriaceae, oblongo-ellipticae, 23.0-30.0 cm longae, 6.5-Io.o cm latae, apice in
acuminem 7.0-I5.0 mm longum contractae, basi rotundatae vel subcordatae; supra
glabra, subtus brunneo-arachnoideo-pubescentes, eglandulosae; costa media supra
prominente, glabra; costis secundariis i6-i8 jugis, supra leviter prominentibus,
subtus prominentibus. Stipulae lanceolatae, subpersistentes, puberulae, apicibus
filamentosis. Flores 3.5-5.0 mm longi in paniculis terminalibus subterminalibusque
dispositi, rachi ramisque breviter tomentellis. Bracteae bracteolaeque 2.0-3.5 mm
longae, ovatae, persistentes. Receptaculum cupuliforme, sessile, extus tomentellum,
intus tomentosum. Calycis-lobi acuti, utrinque tomentosi. Petala 5, glabra. Stamina
circa 23, in orbem completum disposita; filamenta glabra, longe exserta. Ovarium
ad basim receptaculi insertum, lanato-pubescens. Stylus e basi ovarii ortus, ad apicem
hirsutus. Fructus mihi ignoti.
TYPE.Duke9659, Colombia, Choco, south of Curiche, fl (holotype, NY).
DISTRIBUTION. Known only from the vicinity of Curiche, Department of Choc6,
Colombia, where it grows on the beach. FloweringJanuary to May.
COLOMBIA. Choc6: near Curiche, Duke irzog fl (NY); 11679 fl (BTT3).

This species is most closely related to L. macrocarpa and L. durifolia,but differs

from both species in the leaf shape and the well spaced primary veins, the thick
persistent bracteoles, the stipules with a long filamentous apex, and the short distinct
secondary branches of the inflorescence.

4. Licania durifolia Cuatrecasas, Fieldiana Bot. 27: 6i. I950.

Large tree to 25.0 m tall, the young branches tomentose, soon becoming glab-
8 Battelle Memorial Institute, Columbus, Ohio.
Licania 45

48A
,,B n.

\ -^ : ?.....

'
: / * Y

FIG 5. Licania maritima(Duke 11679). A, flower bud, x 5; B, flower bud section, x ; C, ovary
and style, x I5; D, petal, x Io; E, inflorescence, Xo.5; F, habit, x o.5; G, leaf lower surface, X5.
46 Flora Neotropica

rous and conspicuously lenticellate, Leaves oblong to oblong-elliptic, coriaceous,

io.o-24.0 cm long, 3.5-I 2.0 cm broad, caudate to mucronate-acuminate at apex,
with acumen 2.0-7.0 mm long, rounded to subcordate at base, glabrous above,
brown-appressed-arachnoidbeneath; palisade glands present at base of lamina only;
midrib prominent above, glabrous; primary veins I8-23 pairs, impressed above,
prominent beneath; secondary veins parallel or nearly so; petioles 5.0-Io.o mm long,
short-tomentose, eglandular, terete or shallowly canaliculate. Stipules 5.0-8.0 mm
long, caducous, subulate, intrapetiolar. Inflorescences terminal and axillary race-
mose panicles to 25.0 cm long, the rachis and branches densely brown-tomentose.
Bracts at base of inflorescence branches broadly ovate, to 22.0 mm long, other bracts
and bracteoles ovate, 2.0-20.0 mm long, caducous, tomentose on exterior. Flowers
3.5-4.0 mm long, solitary but densely crowded and sessile on primary branches of
inflorescence. Receptacle campanulate-cupuliform, sessile, brown-tomentose on
exterior, villous within. Calyx lobes acute, tomentose on both surfaces.Petals 5, white,
glabrous with ciliate margins. Stamens 25-30, inserted in a complete circle; filaments
far exceeding calyx lobes, connate at base 0.75 mm. Ovary inserted at base of
receptacle, lanate. Style hirsute almost to apex, equalling filaments. Fruit globose
6.o-8.o cm diameter; epicarp glabrous, crustaceous; mesocarp ca 2.0 mm thick,
without parallel fibers; endocarp adnate to mesocarp, thin, hard, glabrous within.
TYPE. Cuatrecasas22251, Colombia, Valle, fl (holotype, F; isotypes, COL, MAD,
P); Cuatrecasas
I533I, Colombia, Valle, fl (paratypes, COL, F, MAD, US).
DISTRIBUTION.Upland forest in western Colombia and Ecuador at Iooo000-
2000 m altitude. Flowering in October and November.
ECUADOR. Lugo44 fl (S); Steyermark
52829 st (F, NY).

This species is distinct from all other species of this subgenus except L. macro-
carpaand L. maritima,in the pubescence of the leaf undersurface, the number of
primary veins, and in the flower. It differsfrom L. macrocarpa in the fruit having a thin
mesocarp which lacks the characteristic parallel fibers of L. macrocarpa, in the endo-
carp which is adnate to the mesocarp, in the yellow-brown pubescence of the in-
florescence, the gray pubescence of the leaf underside, the thicker inflorescence
branches, and in the more densely clustered flowers. The differences between L.
durifoliaand L. maritimaare given under the latter species.

5. Licania macrocarpa Cuatrecasas, Fieldiana Bot. 27: Io7. 1951.

Tree to 20 m tall, the young branches lanate, soon becoming glabrous and
conspicuously lenticellate. Leaves oblong-elliptic, coriaceous, I4.0-28.0 cm long,
3.8-IO.5 cm broad, cuspidate at apex, with acumen 5.0-9.0 mm long, rounded at
base, glabrous above, with dense rufous lanate-arachnoid pubescence beneath;
midrib prominent above, lanate, becoming glabrous with age; primary veins I7-19
pairs, prominent beneath, slightly impressed above; 8.0-20.0 mm long, lanate when
young, becoming glabrous with age, terete, eglandular. Stipules triangular-lanceolate,
3.0-7.0 mm long, intrapetiolar, persistent, pubescent when young. Inflorescences
axillary racemose panicles, the rachis and branches rufous-pubescent. Bracts and
bracteoles triangular, membraneous, glabrescent, subpersistent. Flower buds globose,
ca 3.5 mm diameter, sessile on primary and secondary inflorescence branches.
Receptacle cupuliform, sessile, rufous-tomentellous on exterior, tomentose within.
Calyx lobes acute, tomentose. Petals 5. Stamens ca 25-30, inserted in a complete
circle; filaments exceeding calyx lobes. Ovary inserted at base of receptacle, lanate.
Licania 47

Style hirsute on lower portion, equalling filaments. Fruit globose, 8.0-9.0 cm dia-
meter; epicarp glabrous, crustaceous; endocarp ca Io.o mm thick, fibrous with
parallel fibers, hard; endocarp detached from mesocarp, glabrous within. Seed
globose, 5.5-6.5 cm in diameter.
TYPE. Cuatrecasas I7740, Colombia, Valle, fr (holotype, F; isotype, COL).
DISTRIBUTION. Known only from the lowland Pacific coastal forest of Colombia.
Collected in flower in May.
COLOMBIA. Choc6: Cuatrecasas& Llano24219 fl (NY, US).

Most closely related to L. durifolia;see under that species.

6. Licania salzmanii (Hooker f.) Fritsch, Ann. Naturh. Mus. Wien 4: 49.
1889.

MoquileasalzmanniiHook. f., Mart. Fl. Bras. 14(2): 21. 1867.

Tree, the young branches glabrous, conspicuously lenticellate. Leaves elliptic,

coriaceous, 5.0-Io.o cm long, 2.0-4.5 cm broad, acuminate at apex, subcuneate at
base, shining and glabrous above; palisade glands absent; midrib prominulous and
glabrous above; primary veins 9-11 pairs, inconspicuous on both surfaces; petioles
5.o-6.o mm long, glabrous, canaliculate, eglandular. Stipules caducous (not seen).
Inflorescences terminal and axillary racemose panicles, the rachis and branches
sparselyappressed-puberulous.Bracts and bracteolesminute, linear, caducous. Flowers
2.5-3.0 mm long, solitary on primary branches of inflorescence. Receptacle cupuli-
form, exterior and interior sparsely puberulous; pedicels 2.0-5.0 mm long. Calyx
lobes acute, narrow, puberulous on both surfaces. Petals 5, sparsely puberulous.
Stamens 20-25, inserted in a complete circle; filaments exserted beyond calyx lobes
free almost to base. Ovary inserted at base of receptacle, lanate. Style lanate at base
only. Fruit unknown.
TYPE. Salzmannsn, Brazil, Bahia, fl (holotype, K; isotypes, G, LE, MO, P).
DISTRIBUTION. Known only from Bahia in Brazil. Flowering August to October.
BRAZIL. Bahia: Dahlgrensn fl (F); P. da Silva 27743 fl (A, F, NY, S, SP). Espirito Santo:
Kuhlmann162 st (NY, RB); RB 4811o fl (NY, RB).

LOCAL NAMES. Oiti, Oitizeiro, Oiti cor6.

7. Licania klugii Prance, sp nov Fig 6 A-D.

Arbor parva, ramulis juvenilibus glabris lenticellatis. Folia alternata petiolata;

petiolo Io.o- 2.0 mm longo, eglanduloso, glabro, canaliculato; laminae ellipticae, cori-
aceae, I2.0-I8.o cm longae, 5.5-9.0 cm latae, basi rotundatae vel subcuneatae, apice
in acuminem I0.0-15.0 mm longum contractae, utrinque glabrae, subtus glandulosae;
costa media supra prominente, glabra; costis secundariis 10-I2 jugis, subtus promi-
nentibus, supra planis vel leviter impressis. Stipulae caducae (non visae). Flores
3.5-4.0 cm longi, breviter pedunculati, in paniculis racemosis terminalibus dispositi,
rachi ramisque sparse puberulis. Bracteae bracteolaeque I.0-2.5 mm longae, ovatae,
caducae. Receptaculum campanulato-cupuliforme, extus sparse puberulum, intus
pilosum; pedicelli ad 1.2 mm longi. Petala 5, margine ciliata. Stamina circa 36, in
48 Flora Neotropica

orbem completum disposita; filamenta glabra, ad basim libera, exserta. Ovarium ad

basim receptaculi insertum, pilosum. Stylus e basi ovarii ortus, cum filamentis
aequans, pilosus. Fructus mihi ignoti.
TYPE. Klug 2221, Peru, Loreto, Rio Putumayo, fl (holotype, NY; isotypes,
F, GH, K, US).
DISTRIBUTION. Known only from the type gathering made in riverine forest of
Amazonian Peru.
This species is most closely related to L. guianensis,but differs in the broader more
elliptic leaves with longer petioles, and in the puberulous exterior of the flowers.

8. Licania guianensis (Aublet) Grisebach, Abh. Ges. Wiss. Gottingen 7: I98.

I857; Fritsch, Ann. Naturh. Mus. Wien 4: 53. 1889.

MoquileaguianensisAublet, P1. Guiane I: 521, t. 208. 1775; DC., Prodr. 2: 526. I825.

Small to medium sized tree to 15 m tall, the young branches glabrous, in-
conspicuously lenticellate. Leaves oblong-elliptic to oblong, coriaceous, 7.0-I8.0 cm
long, 2.0-5.5 cm broad, the apex with finely pointed acumen 8.0-22.0 mm long,
cuneate to subcuneate at base, glabrous on both surfaces; palisade glands sparsely
distributed on undersurface, especially towards lamina base; midrib prominulous
above, glabrous; primary veins 8-I2 pairs, prominulous beneath, plane above;
petioles 5.0-7.0 mm long, canaliculate, glabrous, eglandular. Stipules caducous (not
seen). Inflorescences terminal and axillary racemose panicles, the rachis and branches
with only a few short hairs otherwise glabrous. Flowers 4.0-6.0 mm long, on primary
and secondary branches of inflorescence. Bracts and bracteoles i.0-4.0 mm long,
caducous. Receptacle cupuliform, almost glabrous on exterior, densely lanate within;
pedicels I.0-3.0 mm long. Calyx lobes glabrous or sparsely puberulent, acute, the
margins ciliolate. Petals 5, the margins ciliolate. Stamens ca 35, inserted in a complete
circle; filaments far exceeding calyx lobes, glabrous, free almost to base. Ovary
inserted at base of receptacle, villous. Style equalling filaments, the lower portion
densely lanate. Fruit oblong to globose, to 5.0 cm diameter; epicarp smooth and
glabrous, drying black; mesocarp thin, fleshy, fibrous; endocarp thin, fragile,
glabrous within.
TYPE.Aubletsn, French Guiana, st (BM).
Riverine forest of the Guianas, the Amazon delta, and Peru.
DISTRIBUTION.
Flowering November to April.
VENEZUELA. Terr. Delta Amacuro: Steyermark 87301 fr (NY, US, VEN); 87429 fl (F, GH, NY
US); 87735 fr (NY, US). GUYANA. Appun707 fl (K); Boyan39 (F. D. 7723) fl (K, NY, US); De La
Cruz 1283 fl (GH, NY, US); 3031 fl (F, GH, NY); 3484 fl (GH, NY, US); 4591 fl (F, GH, LA, MO,
NY, US); Hohenkerk39A fl (K); Jenman3615 fl (K); 3842 fl (K); 4282 fl (K); 6967 fl (F, K, NY);
6998 fl (F, K); 7125 fl (GH, K, MO, NY, US); Maguire & Fanshawe22949 fl (NY); Schomburgk 905 fl
(BM, K, NY, P); ioo6 fl (BM, K, P); 1382 fl (BR); 1719 fl (K). SURINAME. Lanjouw& Lindeman
3461 fl (K, NY). FRENCH GUIANA. Halle 570 fl (P); Melinon219 fl (P); 263 fl (P); 401 fl (BR, GH,
NY, P); 413 fl (P); 433 fl (P); 495 fl (K, P); Poiteausn fl (P); Sagot 203 fl (BM, BR, GOET, K, P, S,
U); Wachenheim sn fl (BM, F, K, NY, US); 198 fl (P). PERU. Loreto: Schunke344 fl (A, NY, US).
BRAZIL. Par,: Black & Foster48-3386 fl (IAN); DuckeMG 15527 fl (BM, INPA, MG, P, RB, US);
RB 18787 fl (RB, S); RB 18788 fl (RB, S); RB 18789 fl (RB); Frdes20427 fl (IAN, K, US); Pires 1428
fl (IAN, INPA, NY); 2672 fl (IAN, NY); Pires & Black 738 fl (P, RB); A. Silva sn fr (IAN); N. T. Silva
337 fl (NY, P). Terr. Amapa: Frdes25870 fl (NY, RB).

LOCALNAME.Guyana: Swamp Kauta.

Licania 49

B C

1/

..".
Y Y
' o ':~~~~~~~~~~~~.
.............

FIG 6. Speciesof Licania.A-D, L. klugii(Klug 222r); A, habit, x 0o5;B, flower, x 5; C, flower

section, x 5; D, petal, x io. E, L. boliviensis
(Krukoff
r1233); habit, x o'5.
50 Flora Neotropica

9. Licania retifolia Blake, Contr. Gray Herb. 52: 66. 1917.

Small tree 5.0-8.0 m tall, the young branches glabrous, not lenticellate. Leaves
oblong to oblong-lanceolate, coriaceous, 4.0-I2.0 cm long, I.6-4.6 cm broad, obtuse
to acute at apex, cuneate at base, shining and glabrous above, glabrous beneath,
with two glands at lamina base, other palisade glands rare; midrib plane and
glabrous above; primary veins 7-10 pairs, prominulous on both surfaces, slender;
petioles 2.0-6.0 mm long, glabrous, terete, eglandular. Stipules minute, intrapetiolar,
caducous. Inflorescences small dense pyramidal panicles, the rachis and branches
glabrous or sparsely puberulous near flowers. Flowers 3.0-3.5 mm long, solitary or in
small groups on short secondary branches of inflorescence. Bracts and bracteoles
minute, ovate, persistent. Receptacle campanulate, gray-tomentose on exterior,
villous-tomentose within; pedicels 1.5 mm long. Calyx lobes acute, tomentellous on
both surfaces. Stamens I5, inserted in a complete circle; filaments far exceeding
calyx lobes, connate at base for I.0 mm. Ovary inserted at base of receptacle, glabrous.
Style glabrous, equalling or exceeding filaments. Fruit unknown.
TYPE. Langlasse992, Mexico, Guerrero, fl (holotype, GH; isotypes, K, US).
DISTRIBUTION. Known only from the type and one other gathering from west-
central Mexico.
MEXICO. Sinaloa: Lundell 13023 fl (MICH).

Licaniaretifoliais most closely related to L. michauxii,but differs in the inflores-

cence, the longer petioles and in the growth form.

IO. Licania longipedicellata Ducke, Bull. Mus. Hist. Nat. Paris II. 4: 725.
i932.

Small tree, the young branches glabrous. Leaves oblong-linear, coriaceous,

i6.0-33.0 cm long, 5.0-12.0 cm broad, cuspidate-acuminate at apex, with acumen
I5.0-25.0 mm long, rounded at base, glabrous on both surfaces; palisade glands
absent; midrib prominulous above, glabrous; primary veins I3-I7 pairs, prominent
beneath, plane or nearly so above; petioles 7.0-10.0 mm long, glabrous, eglandular,
terete. Stipules linear, to 20.0 mm long, membraneous, glabrous, subpersistent,
intrapetiolar. Inflorescences terminal panicles, the rachis and branches short-
puberulous. Flowers 6.0-7.0 mm long, in clusters on primary and secondary branches
of inflorescence. Bracts at base of branches ovate, to 6.o mm long; other bracts and
bracteoles ovate, to 3.5 mm long, persistent, puberulous on exterior. Receptacle
cupuliform, sericeous-velutinous on exterior, tomentellous within, with a mass of
lanate hairs around the throat; pedicels 3.0-7.0 mm long, with 2.0-3.0 mm below the
articulation. Calyx lobes acute, sericeous-tomentose on both surfaces. Petals 5,
puberulous, white. Stamens 35-45, inserted in a complete circle; filamentsfree to base,
glabrous, far exceeding calyx lobes. Ovary inserted at base of receptacle, lanate.
Style lanate for half of length, equalling the filaments. Fruit unknown.
TYPE. DuckeRB 23603, Brazil, Amazonas, fl (holotype, RB; isotypes, K, P, S,
US).
DISTRIBUTION. Riverine forest in the upper part of BrazilianAmazonia. Flower-
ing September to October.
BRAZIL.Amazonas:Duckeo041fl (GH, IAN, K, MG, MO, NY, R, US).

This distinctive species is easily recognized by the long pedicels, and the finely-
pointed cuspidate leaf apex.
Licania 51

ii. Licania tomentosa (Bentham) Fritsch, Ann. Naturh. Mus. Wien 4: 52.
I889.
MoquileatomentosaBentham, Jour. Bot. Hooker 2: 215. 1840; Hook f., Mart. Fl. Bras. 14(2): 2I.
1867.
PleraginaodorataArruda da Camara ex Koster, Trav. Bras. 499. 1816; Prance, Mem. N. Y. Bot.
Gard. 15(1): I34. 1966, nom illegit.

Tree to 20.0 m tall, the young branches lanate-tomentose, soon becoming

glabrous. Leaves oblong-elliptic to oblong-lanceolate, chartaceous, 4.5-I2.0 cm long,
1.5-4.5 cm broad, apiculate-acuminate at apex, with acumen 2.0-4.0 mm long,
cuneate to subcuneate at base, farinaceous-lanate on both surfaces when young, be-
coming glabrous with age, the lamina base with two glands, but other palisade glands
absent; midrib prominulous above, lanate-pubescent when young; primary veins
7-IO pairs, thin, prominulous beneath, level or nearly so and inconspicuous above;
petioles 4.0-6.0 mm long, lanate when young, becoming glabrous with age, with 2
glands, shallowly canaliculate. Stipules caducous, linear, membraneous, intrapetiolar.
Inflorescences little-branched racemose panicles or racemes, predominantly axillary,
the rachis sparsely gray-tomentose. Flowers ca 3.0 mm long, solitary but densely
crowded along rachis. Receptacle campanulate, sparsely gray-tomentose on exterior,
tomentose within; pedicels I.0-2.0 mm long. Calyx lobes acute, tomentellous on
exterior, puberulous within. Petals 5, almost glabrous, with ciliate margins. Stamens
ca 30, inserted in a complete circle; filaments far exceeding calyx lobes, slighty
connate at base. Ovary inserted at base of receptacle, lanate. Style hirsute most of its
length, equalling or exceeding filaments. Fruit oblong; epicarp smooth, drying black;
pericarp thick, fragile, fibrous, glabrous within.
TYPE. Gardner992, Brazil, Pernambuco, fl (holotype, K; isotypes, BM, CGE, GH,
NY, OXF, P, S, US).
Native of the coastal restinga forest of northeastern Brazil, but
DISTRIBUTION.
widely cultivated outside the region. Flowering June to November in northeastern
Brazil.
BRAZIL. Amazonas: (cult), CoelhoINPA 1286 fl (IAN, INPA, MG, NY); INPA 2966 fr (INPA,
NY). Par,: (cult.), Baker 64 fl fr (GH, K, LE, NY, P, U, US); Cavalcante234 fl (IAN, MG); Huber
3015 fl (BM, INPA, MG, P, RB, US); MG 9360 fl (BM, F, MG, U); MG 15224 fl (U); Kuhlmann&
Jimbo 9 fl (FHO, IAN, MG, SP). Maranhio: (cult), Frdes 11963 fl (MICH, NY, US). Piaui: Gardner
2559 fl (BM, GH, K, NY, OXF, P). CearA: Dahlgrensn fl (F); Ducke 15i8 fl (MG, RB); MG i616 fl
(MG); Eugenio711 fl (R, RB, S); Allemao& Cysneiros562 fl (P, R). Paraiba: Pickel 1175 fl (IPA); 2841
fl (IPA). Pernambuco: Pickel 1878 fl fr (IPA); Prance4811 st (NY). Bahia: Velloso1065 fl (R). Guana-
bara: Aguiar & Souza 241 fl (RB 138188); Glaziou237 fl (BR, P); I514 fl (BR, F, FI, K, P, R); 7879 fl
(P); 10703 fl (K, P); Kuhlmann1001 fl (RB 136937); Miers3890 st (K); Saldanha5430 fl (R); Whitford3
fl (GH, K, S); Widgrensn fl (S). Santa Catarina: Klein 2731 fl (BM).

LOCAL NAMESAND USES. Oiti, Oiti da Praia. Cultivated widely for its edible fruit
and as a shade tree because of its low and very broad crown. Also cultivated in South
Africa, as in the Botanic Gardens Durban (Thorns& Strey5323 fl K) and Mauritius
(Bot. Gdn. 98, 23I fl K).
Hooker described two varieties in Flora Brasiliensis,both based on Gardner
collections from northeastern Brazil. The two varieties were distinguished from the
typical variety by their leaf shape. However, with the more abundant material now
available, there is found a complete range in leaf shape from oblong-ovate to oblong-
linear. As there are no breaks in this range of variation no basis exists for maintaining
these two varieties (var latifolia Hook. f., and var angustifolia Hook. f., both in Mart.
Fl. Bras. 14(2): 2I. 1867.)
The status of Pleragina odorata Arruda has been discussed elsewhere (Prance
and
I966a), where it was shown that this species must be included with L. tomentosa,
that the name P. odoratawas not validly published.
52 Flora Neotropica

I2. Licania pyrifolia Grisebach, Fl. Brit. W. Ind. 230. i860; Fritsch, Ann.
Naturh. Mus. Wien 4: 53. I889.
Moquileapyrifolia (Grisebach) R. O. Williams, Fl. Trinidad & Tobago I: 314. 1932.
Moquileamacrocarpa Pittier, Bol. Soc. Venez. Ci. Nat. 4: 352. I938, synon nov. Types. Pittier 8202,
Venezuela, Carabobo, fl (holotype, VEN; isotypes, F, GH, MICH, MO, US); 12230,
Venezuela, Guarico, fl (paratypes, A, NY, US, VEN); 1140 fr (paratypes, GH, NY, P,
VEN).

Small to medium-sized tree, often with spreading branches, the young branches
lanate, soon becoming glabrous, inconspicuouslylenticellate. Leaves broadly elliptical
to oblong, chartaceous, 4.5-11.0 cm long, 2.0-5.5 cm broad, acute to abruptly
apiculate at apex, with acumen to 3.0 mm long, rounded to cuneate at base, shining
and glabrous above, with caducous lanate pubescence beneath, becoming glabrous
with age; palisade glands present beneath; midrib plane above, lanate; primary
veins 6-8 pairs, thin, prominulous beneath, inconspicuous above; petioles 8.0-I6.o
mm long, sparsely pubescent when young, terete, eglandular, not more than 1.2 mm
thick. Stipules caducous, linear, membraneous. Inflorescencesspreading terminal and
subterminal panicles, the rachis and branches brownish-gray-tomentose. Flowers
4.0-4.5 mm long, solitary or in small groups on primary and short secondary branches
of inflorescence. Bracts and bracteoles 0.5-3.0 mm long, caducous. Receptacle
cupuliform, brownish-gray-tomentoseon exterior, tomentose within; pedicels 1.0-2.0
mm long with ca 2.0 mm of stalk below the articulation. Calyx lobes acute, tomentose
on exterior, puberulous within. Petals 5, glabrous with ciliate margins. Stamens 25-
35, inserted in a complete circle; filaments far exceeding calyx lobes, free to base.
Ovary inserted at base of receptacle, lanate. Style hirsute almost to apex, equalling
filaments. Fruit oblong, to I I.0 cm long and 5.0 cm broad; epicarp densely verrucose,
glabrous; pericarp thick, fragile, fibrous, undifferentiated, glabrous within.
TYPE. Purdie sn, Trinidad, fl (K).
DISTRIBUTION. Lowlandforest and riverine forest in open places, Martinique to
Trinidad and adjacent Venezuela.
WINDWARD ISLANDS. Martinique: Belangersn fl (P); 622 fl (P); 747 fl (P); Hahn 1124 fl
(BM, BR, FI, K, LE, P, R). COLOMBIA. Boyaca: Garcia-Barriga18528 fl (COL, NY). VENEZUELA.
Aragua: TamayoI42I fl (F, US, VEN). GuArico: Aristeguieta4607 fl (VEN); 4988 st (VEN); Castellanos
25 fl (VEN); Pittier 12402 fr (VEN). Apure: Ramia586 fr (VEN). Bolivar: Holt & Gehriger88 a fr (US,
VEN); Humbolt & Bonpland1059 fr (F, VEN); Little & Petit 15912 fr (VEN); LI. Williams 12694 fr
(F, VEN); 12917 fr (F, VEN); Wulffsn fl (VEN). Terr. Amazonas: LI. Williams 15772 fr (F). TRINI-
DAD. Broadway2884 fl (GH, K, NY, US); Cranssn fl (K, TRIN); Cruegersn fl (K, NY); Evans sn fl
(TRIN); Prance2112 fr (NY); Rogerssn fl (TRIN); R. 0. Williams 12052 fl (K, NY); 12450 fr (TRIN).

LOCAL NAME AND USES. Venezuela: Merecure.Cultivated for the edible fruit.
The distinctions indicated by Pittier between this species and Moquileamacrocarpa
cannot stand, partly because the Trinidad material does not have unguiculate petals.
Licaniapyrifoliais remarkable for its petioles, which are very slender for their length.
to
The leaf shape, the petioles, and the inflorescence distinguish it from L. leucosepala,
which it is most closely related.

13. Licania leucosepala Grisebach, Abh. Ges. Wiss. Gottingen 7: I98. 1857;
Fritsch, Ann. Naturh. Mus. Wien 4: 53. 1889.
Moquilealeucosepala(Grisebach) R. O. Williams, Fl. Trinidad & Tobago I: 3I5. 1932.

Small tree, the young branches glabrous and lenticellate. Leaves oblong to
oblong-lanceolate, 6.0-14.0 cm long, 2.0-5.0 cm broad, coriaceous, acuminate at
Licania 53

apex, with acumen 4.0-9.0 mm long, cuneate to subcuneate at base, glabrous on both
surfaces, the lower surface with palisade glands especially at base; midrib almost
plane above, lanate when young; primary veins 8- I pairs, thin, prominulous be-
neath, inconspicuous above; petioles 4.0-6.0 mm long, lanate when young, becoming
glabrous with age, canaliculate, eglandular. Stipules linear, ca 4.0 mm long, mem-
braneous, caducous. Inflorescences spreading terminal and axillary panicles, the
rachis and branches puberulous to densely brown-tomentose. Bracts and bracteoles
I.0-3.5 mm long, caducous. Flowers ca 4.0 mm long, solitary or in small groups on
primary and short secondary branches of inflorescence. Receptacle cupuliform-
campanulate, densely brown-tomentose on exterior, villous-tomentose within;
pedicels ca 1.5 mm long. Calyx lobes acute, tomentellous on both surfaces. Petals 5,
sparsely pubescent, the margins ciliate. Stamens ca 30, inserted in a complete circle;
filaments far exceeding calyx lobes, slightly connate. Ovary inserted at base of
receptacle, tomentose. Style hirsute almost to apex, exceeding filaments. Fruit
globose or ellipsoid, 3.5-5.0 cm long; epicarp verrucose; pericarp thick, undifferen-
tiated, fragile, fibrous, glabrous within.
TYPE. Duchassaingsn, Guadeloupe, fl (holotype, GOET; isotypes, K, P).
DISTRIBUTION. Forest in the Antilles, Trinidad and adjacent Venezuela. Flower-
ing December to May.
LEEWARD ISLANDS, Guadeloupe: Questel 754 fl (P, US); 876 fl (US). WINDWARD
ISLANDS. Dominica: Duss I53 fr (NY); Eggers 753 fl (BR, GH, LE, P, R); io6i fl (K); Jones sn fl
(K). St. Lucia: Ramagesn fl (BM, K). St. Vincent: H. H. & G. W. Smith 478 fl (BM, GH, K, NY, S);
1566 fl (BM, K); 1838 fr (GH, K, NY). VENEZUELA. Sucre: Steyermark6284I fr (NY, VEN).
Bolivar: Bernardi7125 fl (K, US). TOBAGO. Broadway4483 fl (BM, F, GH, K, MO, NY, S,U, US).
TRINIDAD. Dannouse6653 fr (NY); sn fl (NY, TRIN).

14. Licania angustata Prance, sp nov Fig 7 A-C.

Arbor magna, ramulis juvenilibus tomentellis mox glabris. Folia alternata

petiolata; petiolo IO.O-I3.0mm longo, juvenili tomentoso, eglanduloso, leviter
canaliculato; laminae oblongo-lanceolatae, coriaceae, I0.0-I4.5 cm longae, 2.5-5.2
cm latae, basi rotundatae vel subcuneatae, apice in acuminem 7.0-12.0 mm longum
contractae, utrinque glabrae, subtus basi cum glandulis duobus munitae; costa media
supra prominente, sparse pubescente, mox glabra; costis secundariis I 1-17 jugis,
supra leviter impressis, subtus prominentibus. Stipulae ovatae vel lineares, .o0-2.0
mm longae, intrapetiolares, caducae. Flores circa 3.5 mm longi, in paniculis race-
mosis terminalibus vel subterminalibus dispositi, haud pedunculati, rachi ramisque
dense villoso-tomentosis. Bracteae bracteolaeque lanceolatae, ad 2.2 mm longae,
persistentes, extus tomentosae. Receptaculum campanulato-cupuliforme, extus
tomentosum, intus villoso-tomentosum; pedicelli circa 0.5 mm longi. Calycis-lobi
acuti, tomentosi, intus puberuli. Petala 5, sparse pubescentia. Stamina circa 17, in
orbem completum disposita; filamenta glabra, longe exserta. Ovarium ad basim
receptaculi insertum, lanato-pubescente. Stylus e basi ovarii ortus, calycis-lobos
excedens. Fructus mihi ignoti.
TYPE.Krukoff8445, Brazil, Amazonas, municipality of Sao Paulo de Olivenca,
near Palmares, fl (holotype, NY; isotypes, A, BM, BR, F, K, LA, LE, MO, P, S, US).
DISTRIBUTION. Lowland non-flooded forest in western Amazonia. Flowering
September and October.
PERU. Amazonas: Mouth of Rio Santiago, Tessmann4502 fl (NY).
54 Flora Neotropica

.A.

A D

-
.. ..........."

\'.. .....................

?x7 ?,5 :

1 ~ :;
~~~
'
.. ~ ~ ~ ~~~~~~~~~~~~~~~~~~~
LIRI

IG 7.Seiso
G, flwer ectin iai.AG (Eler 87);D,
x.D-, .agsaa(rkjfhbit,x 0.;E,flowr,
45;A 6
ai,x.;B fowersecton,lwr 7.5
L.eglei x7.5;F,
pea,xI.HK
G, .rtci Prs~Sla65) , ai,x05 J lwr .;K lwrscin
x7 .5.~~~~~~~~~"

FIG 7. Species of Licania. A-C, L. angustata (Krukoff8445); A, habit, xo.5; B, flower, x6;
C, flower section x 6. D-G, L. egleri(Egler887); D, habit, x o.s; E, flower, x 7.5; F, flower section, x 7.5;
G, petal, x i o. H-K, L.fritschii (Pires & Silva 6731); H, habit, x o.s; J, flower, x 7.5; K, flower section,
Licania 55

Licaniaangustatais most closely related to L. britteniana,

but differs in the oblong-
lanceolate leaves and the larger flowers.

I5. Licania platypus (Hemsley) Fritsch, Ann. Naturh. Mus. Wien 4: 53.
1889; Standl., Contr. U. S. Natl. Herb. 23: 344. 1922.
MoquileaplatypusHemsley, Diag. P1. Nov. Mex. 9. I880; Biol. Centr. Am. Bot. I: 366. I88o;
Hook. Ic. P1. 27, t. 2618-9. I899.

Medium to large-sized tree to 30 m tall, sometimes buttressed, the young

branches glabrous and lenticellate. Leaves oblong to oblong-lanceolate, coriaceous,
I3.o-20.0 cm long (to 30.0 cm long on sterile branches), 3.5-8.o cm broad, short-
acuminate at apex, with acumen 2.0-I0.0 mm long, rounded to subcuneate at base,
glabrous on both surfaces, with two glands at base of lamina on underside; palisade
glands rare or absent; midrib glabrous above, plane or slightly impressed; primary
veins I5-22 pairs, prominulous beneath, level above or nearly so; petioles 9.0-I4.0
mm long, glabrous, terete, eglandular. Stipules 2.0-4.0 mm long, ovate, persistent,
intrapetiolar. Inflorescences terminal racemose panicles, the rachis and branches
with a short sparse gray pubescence. Flowers 3.0-4.0 mm long, solitary or in small
groups on primary and secondary branches of inflorescence. Bracts and bracteoles
I.o-6.o mm long, caducous. Receptacle cupuliform, with short gray pubescence on
exterior, densely hirsute within; pedicels o.5-2.0 mm long. Calyx lobes acute,
pubescent on both surfaces. Petals 5, glabrous with ciliate margins, caducous. Stamens
I5-20, inserted in a complete circle; filaments slightly exceeding calyx lobes, glabrous,
free almost to base. Ovary inserted at base of receptacle, villous. Style villous on lower
portion. Fruit ellipsoid to ovoid, up to 15.0 cm long at maturity; epicarp glabrous,
verrucose; mesocarp thick, fibrous, and fleshy; endocarp hard and thin, glabrous
within.
TYPES.Cuming 1272, Panama, fl (lectotype, K; isotype, MO), Levy 222, Nicara-
gua (paratypes, F, K, P).
DISTRIBUTION. Forest, commonest near rivers, but also in forest away from rivers,
from southern Mexico through Central America to Northern Colombia.
MEXICO. Veracruz: Gomez-Pompa887 st (A); 922 fr (MEXU); LI. Williams 86o5 st (A, F);
8722 st (MICH, S). Puebla: Bravo385 st (MEXU); Pennington& Sarukhdn9277 fr (FHO, NY). Guer-
rero: Palmer428 fr (F, K). Oaxaca: Pennington& Sarukhdn7326fl (FHO, MEXU, NY); 9326fl (MEXU
NY). Chiapas: Matuda 16380 fl (F, MEXU, MICH); 16450 fl (F, MEXU, MICH); Miranda7256 fl
(MEXU); Without precise locality: Bourgeau3124 st (GH, LE); LiebmannI747 fl (BM, NY); 1753 st
(F); I754 st (GH). GUATEMALA. Aguilar1661 fr (F); FAO-FYDEP 21 fr (F, NY); Heyde & Lux 6421
fl (BM, GH, K, NY); Johnson1070 st (LA); Kellerman5670 fr (MEXU, MICH); Standley72644 st (F);
Steyermark 44656 st (A, F); 45350 st (A, F); 46132 st (F); 46465 st (A, F); 48073 fl (A, NY, S); 49?3L3'
st (F). BRITISH HONDURAS. Cambellsn fl (K); Schipp308 st (A, BM, F, GH, K, NY, S). HON-
DURAS. Molina R. 3637 fl (F, GH); Standleyi88I5 st (F); 53845 st (A, F); 55732 st (A, F). EL SAL-
VADOR. Allen 6885 fl (F, GH, NY); Calderon902 fl (GH, MO, NY); Standley20532 St (GH, NY);
22228 St (GH, NY); L. 0. Williams & Calderdn15224 St (GH); L. O. Williams & Molina R. 15224
(F, GH). NICARAGUA. Standley11517 st (F). COSTA RICA. Brenes 12170 st (F); 12170a fr (F);
Pittier 12102 fl (P). PANAMA. Allen 930 fl (BR, F, GH, K, NY, P, S, U); 941 fl (F, FI, K, MICH,
MO, NY, P, S, U); Cooper& Slater256 st (F, NY); Johnston659 fr (GH, P, U); Kluge31 fl (F); Pittier
3347 fl (GH); 6971 fr (GH); 6998 st (NY); Shattuck575 st (F); Zetek4351 st (F). COLOMBIA. Bolivar:
Curran146 fl (US); RomeroC. 1179 st (COL, US). TRINIDAD. Cult, Prance2113 fr (NY).

LOCAL NAMES AND USES. Mexico: Zin zapote, Zapote cabello, Zapote amarillo,
Zapote borracho,Mesonsapote. Guatemala: Moshpin, Urraco. Honduras: Urraco. British
Honduras: Monkey apple. Salvador: Sunzapote, Sunza. Costa Rica: Zapote. Panama:
Sangre. Colombia: Chupa. This species has an edible fruit which is much sought after
by birds and animals, but little eaten by man.
56 Flora Neotropica

x6. Licania gonzalezii Miranda, Bol. Soc. Bot. Mex. 29: 36, fig 2. I965.

Tree, the young branches puberulous, becoming glabrous and lenticellate with
age. Leaves elliptic, chartaceous, 10.0-I3.0 cm long, 4.5-6.2 cm broad, abruptly
short-acuminate-mucronate at apex, with acumen 2.0-5.0 mm long, rounded to
subcuneate at base, glabrous on both surfaces; palisade glands absent; midrib
glabrous above, prominulous; primary veins 9-I pairs, prominent beneath, pro-
minulous above; petioles 5.0-8.0 mm long, pubescent, shallowly canaliculate,
eglandular. Stipules filamentous, membraneous, to 7.0 mm long, subpersistent,
axillary. Inflorescences terminal and subterminal racemose panicles, the rachis and
branches puberulous. Flowers 4.0-5.0 mm long, densely crowded along primary
branches of inflorescence. Bracts and bracteoles ovate, I.0-4.0 mm long, persistent.
Receptacle campanulate-cupuliform, tomentellous-puberulouson exterior, tomentose
within; pedicels 0.25-I.0 mm long, Calyx lobes acute, puberulous on both surfaces.
Petals 5, ciliate on margins, not clawed. Stamens I4-45, inserted in a complete circle;
filaments far exceeding calyx lobes, glabrous, free almost to base. Ovary inserted at
base of receptacle, pilose. Style equalling filaments, pilose almost to apex. Fruit un-
known.
TYPES. Ortega 41, Mexico, Nayarit, fl (holotype, MEXU; isotype, US); Miranda
9062, Mexico, Jalisco, st (paratype, MEXU).
Known only from the type gatherings from western Mexico. This
DISTRIBUTION.
species is closest to L. platypusbut differs in a large number of minor features. It is
distinguished mainly by the smaller elliptic leaves with fewer primary veins, and the
shorter petioles. This is the most northerly species in Mexico.

17. Licania egleri Prance, sp nov Fig 7 D-G, Fig 8.

Arbor parva vel media, ramulis juvenilibus glabris lenticellatis. Folia alternata
petiolata; petiolo 2.0-5.0 mm longo, glabro, rugoso, tereti, eglanduloso; laminae
obovatae vel ellipticae, coriaceae, 4.5-8.5 cm longae, 2.0-4.0 cm latae, basi rotun-
datae vel subcuneatae, apice retusae vel breviter apiculatae, utrinque glabrae, subtus
glandulosae et basi cum glandulis duobus munitae; costa media supra prominente,
glabra; costis secundariis 8-I3 jugis, utrinque leviter prominentibus, inconspicuis.
Stipulae parvae, I.0-2.0 mm longae, ovatae, membranaceae, intrapetiolares, sub-
persistentes. Flores 2.0-2.5 mm longi, in paniculis racemosis terminalibus subter-
minalibusque dispositi, haud pedunculate, rachi ramisque puberulis, Bracteae
bracteolaeque 0.5-I.5 mm longae, ovatae, persistentes, membranaceae, sparse
pubescentes. Receptaculum campanulatum, extus griseo-tomentellum, intus tomen-
tosum; pedicelli 0.5-i.0 mm longi. Calycis-lobi acuti, extus tomentelli, intus puberuli.
Petala 5, pubescentia. Stamina circa 15, in orbem completum disposita; filamenta
exserta, ad basim libera, glabra. Ovarium ad basim receptaculi insertum, villoso-
tomentosum. Stylus e basi ovarii ortus, calycis-lobos excedens, villosus. Drupa ovoidea
4.0-5.0 cm longa, extus laevis glabra.
TYPE. Egler 887, Brazil, Para, Tapaj6s, Rio Cururu, fl (holotype, NY; isotypes,
IAN, MG 23789).
DISTRIBUTION. Open non-flooded forest, beside rivers and in tall cerrado(cerradao)
in Amazonian Brazil and the northern limits of the Planalto of Central Brazil.
Flowering May to October.
COLOMBIA.Amazonas:Schultes& Cabrera15599 fr (GH, NY, US). BRAZIL. Amazonas:
Frdes 23272 fl (IAN); 23891 fl (IAN, US); 25040 fl (F, IAN, NY); 25173 fl (IAN, NY); 26459 fr
& Loureiro
(IAN, NY); Krukof885i fl (A, BR, LE, MO, NY, S, U); Rodrigues 5828 fl (INPA 14780,
Licania 57

* a ^A A

FIG8. o grp . g, L L ni
mi

FIG 8. Geographic distribution of species of Licania. * L. egleri,A L. minutiflora.

NY). Para: Egler 828 fl (IAN, MG 23791). Terr. Rond6nia: Maguire, Pires, Maguire & Silva 56538 fl
(NY); Cordeiro& Silva 87 fr (IAN, NY); 225 fl (IAN, NY); Pranceet al 6692 fl (INPA, NY). Mato
Grosso: Irwin & Soderstrom6651 fl (NY); Lima 58-3139 fl (IPA, NY, RB); 58-3175 fl (IPA, RB);
Pires 56823 fl (UB). Goias: Prance & Silva 58991 fl (NY, US).

This species, although widespread, has been overlooked until now because of the
confusion in the circumscription of L. minutiflora. Licaniaegleriis related to L. minuti-
flora but differs in the shorter petioles and in the smaller obovate leaves with a retuse
to mucronate apex. This species is named for the late Dr. W. A. Egler of the Museu
Goeldi, Belem, who, until his untimely death, contributed much to our understand-
ing of the Amazonian flora.

I8. Licania minutiflora (Sagot) Fritsch, Ann. Naturh. Mus. Wien 4: 54.
I889. Fig 8.
MoquileaminutifloraSagot, Ann. Sci. Nat. Bot. VI. I5: 308. 1883.
Licania rondoniiPilger, Notizbl. Bot. Gart. Berlin 7: 540. I923, synon nov. Type. Hoehne4602,
Brazil, Mato Grosso, fl (holotype, B, lost; isotype, SP).
MoquilearipariaGleason, Bull. Torrey Club 51: 192. 1934, synon nov Types. Frdes 1961, Brazil,
Maranhao, fl (holotype, NY; isotypes, BM, F, K, MICH, MO, P, S); 1935 fl (paratypes,
A, BM, F, K, MICH, MO, NY, P, S, US).
Licaniariparia(Gleason) Standley, Publ. Field Mus. Bot. 17: 258. 1937.
58 Flora Neotropica

Large tree to 45.0 m tall, sometimes with low-buttressed trunk, the young
branches glabrous. Leaves ovate to oblong-elliptic, coriaceous, 5.5-11.0 cm long,
2.0-6.o cm broad, apiculate-acuminate at apex, with acumen I.o-6.o cm long,
rounded to subcuneate at base, glabrous on both surfaces, usually without palisade
glands; midrib prominulous above, sparsely pubescent when young; primary veins
8- 3 pairs, prominulous beneath; petioles 6.0-i I.o mm long, tomentose when young,
glandular or eglandular, terete. Stipules ovate, i.5 mm long, persistent, intrapetiolar.
Inflorescences spreading terminal and subterminal panicles, the rachis and branches
brown-tomentose. Bracts and bracteoles 0.2-1.0 mm long, caducous, tomentose on
exterior. Flowers ca 2.0 mm long, solitary or in small groups on primary branches of
inflorescence. Receptacle campanulate-cupuliform, densely brown-tomentose on
exterior, tomentose within; pedicels ca I.o mm long. Calyx lobes acute, tomentose on
both surfaces. Petals 4-5, cream-white, sparsely pubescent. Stamens I4-21, inserted
in a complete circle; filaments far exceeding calyx lobes, slightly connate at base.
Ovary inserted at base of receptacle, lanate. Style hirsute almost to apex, equalling
or exceeding filaments. Fruit ellipsoid, 3.5-4.5 cm long; epicarp minutely lenticellate,
glabrous; mesocarp thick, fibrous; endocarp thin, hard, glabrous within.
TYPE. Melinon sn, French Guiana, fl (holotype, P; isotypes, A, F, G, K, NY, R,
US).
DISTRIBUTION. Non-flooded forest in the Guianas and northern Brazilian
Amazonia. Flowering April to November.
VENEZUELA. Terr. Amazonas: Maguire,Cowan& Wurdack29448 fl (F, NY, US). GUYANA.
Davis 259 (F.D. 2250) fl (K); Fanshawe598 (F.D. 3334) fl (K, NY, U); 2004 (F.D. 4740) fl (K, NY, P, S,
U, US). SURINAME. Schulz 7947 fl (NY, U). BRAZIL. Amazonas: DuckeRB 35565 fl (K, RB). Terr.
Roraima: Pranceet al 4590 fr (INPA, NY). Para: Black 50-9963 fl (NY). Maranhao: Froes3404o fl
(IAN); Moses 38 fl (BM).

The only difference between the type specimens of L. minutiflora and L. ripariais
in the leaf size. Since there is a complete gradation of leaf size between the two type
specimensand in the material subsequentlycollected, L. ripariais reduced to synonymy.

I9. Licania maranhensis Prance, sp nov Fig 21.

Arbor?, ramulis juvenilibus tomentellis mox glabris. Folia alternata petiolata;

petiolo circa 8.0 mm longo, juvenilo tomentello, eglanduloso, tereti; laminae sub-
coriaceae, ellipticae, I5.0-I8.5 cm longae, 7.5-9.5 cm latae, apice in acuminem circa
5.0 mm longum contractae, basi subcordatae, utrinque glabrae, cum glandulis
duobus basim versus munitae; costa media supra leviter prominente, sparse adpresso-
pubescens; costis secundariis I4-I5 jugis, subtus prominentibus. Stipulae caducae
(haud visae). Alabastra circa 2.0 mm longa. Flores in paniculis terminalibus axillari-
busque dispositi; rachi ramisque sparse tomentellis. Bracteae bracteolaeque 1.0-3.5
mm longae, membranaceae, persistentes. Receptaculum subsessile, campanulato-
cupuliforme, extus tomentosum, intus tomentosum. Calycis-lobi acuti, extus tomen-
tosi. Petala 5. Stamina circa 20, in orbem completum disposita; filamenta glabra
(probabiliter longae exserta, in alabastro tantum visa). Ovarium ad basim recepta-
culi insertum, pilosum. Stylus e basi ovarii ortus. Fructus mihi ignoti.
TYPE. Lisb6a MG 2462, Brazil, Maranhao, Barra da Corda, fl bud (holotype,
NY; isotype, RB).
DISTRIBUTION. Known only from the type collected from a river margin in central
Maranhao, Brazil.
Licania 59

This species is closest to L. minutiflora,but differs in the much longer broader

leaves with more primary veins and more tapered apices, and in the subcordate leaf
bases.

20. Licania fritschii Prance, sp nov Fig 7 H-K.

Arbor, ramulis juvenilibus glabris, haud lenticellatis. Folia alternata petiolata;

petiolo 6.o-Io.omm longo tereti vel leviter canaliculato, glabro, eglanduloso;
laminae ellipticae, coriaceae, 9.0-15.0 cm longae, 4.0-7.0 cm latae, basi rotundatae,
apice in acuminem, 4.0-Io.0 mm longum contractae, utrinque glabrae, subtus
glabrae, subtus sparse glandulosae et basi cum duobus glandulis munitae; costa
media supra prominente, glabra; costis secundariis 9- I jugis, utrinque leviter
prominentibus. Stipulae minutae, subulatae, glabrae, subpersistentes,intrapetiolares.
Flores 3.0-3.5 mm longi, haud pedunculati, in paniculis terminalibus subterminali-
busque dispositi, rachi ramisque sparse puberulis. Bracteae bracteolaeque 0.3-0.8
mm longae, puberulae, caducae. Receptaculum campanulato-cupuliforme, extus
brunneo-puberulum, intus tomentosum; pedicelli 0.5-1.0 mm longi. Calycis-lobi
acuti, utrinque puberuli. Petala 5, sparse pubescentia, margine ciliata. Stamina
16-25, in orbem completum disposita; filamenta longe exserta, ad basim libera,
glabra. Ovarium ad basim receptaculi insertum, piloso-tomentosum. Stylus e basi
ovarii ortus, calycis-lobos excedens, pilosus. Fructus mihi ignoti.
TYPE. Pires & Silva 673I, Brazil, Para, Belem, fl (holotype, NY; isotypes,
IAN, UB).
DISTRIBUTION. Periodically inundated forest in Amazonian Brazil. Flowering
October to February.
BRAZIL. Amazonas:Borba, Rio Madeira, DuckeRB 35564 fl (K); Sao Paulo de Olivenga,
885ofl (A, F, K, NY). Para:Belem,DuckeRB 19784fl (K, RB).
Krukoff

This species is closest to L. britteniana but differs in the larger,

and to L. minutiflora
thickly coriaceous leaves with glabrous petioles, and in the pubescence of the in-
florescence.I have named this species for Dr. Carl Fritsch in recognition of his contri-
bution to our knowledge of Licania.

21. Licania britteniana Fritsch, Osterr. Bot. Zeitschr. 42: 6. I892.

Licaniapallida Britton, Bull. Torrey Club 17: 9. 1890. Type. Rusby2442.
MoquileaelataPilger, Notizbl. Bot. Gart. Berlin6: I36. 1914. Type. Ule 9446, Brazil,Acre, fl
(K, MG 14292).
Licaniadlata(Pilger)Pilgerex L1.Williams,Publ. Field Mus. Bot. 15: 174. I936; Macbride,Fl.
Peru 1070. 1938.

Large tree to 50.0 m tall, the young branches glabrous and lenticellate. Leaves
oblong-elliptic to oblong, coriaceous, 8.o-I5.5 cm long, 3.0-7.0 cm broad, acuminate
at apex, with acumen 3.0-7.0 mm long, rounded to subcordate at base, glabrous on
both surfaces; palisade glands present beneath, with two glands at junction of
lamina and petiole on lower surface; midrib glabrous above, plane or nearly so;
primary veins 9-I5 pairs, prominulous above, prominent beneath; petioles 8.o-
I2.0 mm long, with sparse arachnoid pubescence when young, soon glabrous,
shallowly canaliculate, eglandular. Stipules 2.0 mm long, subulate, glabrous, per-
sistent, intrapetiolar. Inflorescences terminal racemose panicles, the rachis and
60 Flora Neotropica

branches gray-tomentose. Flowers 2.0-3.0 mm long, solitary but densely crowded on

primary and secondary branches of inflorescence. Bracts and bracteoles ca I.5 mm
long, caducous, tomentose on exterior. Receptacle campanulate, gray-tomentose on
exterior, pilose-tomentosewithin; pedicels 0.5 mm long. Calyx lobes acute, tomentose
on exterior, puberulous within. Petals 5, sparsely pubescent, the margins ciliate.
Stamens I6-23, inserted in a complete circle; filaments far exceeding calyx lobes,
their bases slightly connate. Carpels I(-2), the second carpel occasionally develop-
ing. Ovary inserted at base of receptacle, lanate. Style pilose almost to apex, equalling
filaments. Fruit ovoid to 8.o cm long; epicarp crustaceous; mesocarp thick and hard;
endocarp thin, hard, sparsely pubescent within.
TYPE.Rusby2442, Bolivia, Rio Beni, fl (holotype, NY; isotypes, BM, F, GH, K,
MICH, P, US).
DISTRIBUTION. Non-flooded forest and periodically flooded forest in western
Amazonia. Flowering July to December.
COLOMBIA.Amazonas:Schultes& Black8588 fl (COL, F, GH, IAN, K NY, US). PERU.
Loreto:Ellenberg2412st (U); Tessmann 3215 fl (S); 3250 fl (NY, S); 5468 fl (NY, S); LI. Williams5064
fl (BM, F, S, US). BRAZIL. Amazonas:Kuhlmann RB 17985 fl (K, P, RB, S, US); Krukoff 5165 fl
(A, BM, F, K, LE, MICH, MO, NY, S, US); 5635 fl (A, BM, F, K, MICH, MO, NY, S, US); 6139 fl
(A, BM, BR, F, K, LE, MICH, MO, NY, RB, S, US); 6306fr (A, BR, F, IAN, K, LE, MICH, MO,
NY, RB, US); Pranceet al 2524 fl (INPA, NY). Pari: DuckeMG 15I75 fl (BM, MG, P, RB, US); Silva,
651 fl (IAN).

The nomenclature of this species has been somewhat confused in the past, be-
cause of both Moquileapallida Hook f. and Licaniapallida Spruce ex Sagot. The latter
name, from Spruce's manuscript, was only cited in synonymy by Hooker in Flora
Brasiliensis,but it was used later as a specific epithet in 1883 by Sagot. This legitimate
usage of L. pallida was not cited in Index Kewensis, resulting in the general and
illegitimate application of L. pallidato L. britteniana, even though Fritsch had clarified
the position in 1892 when he proposed the latter name. In dedicating this species for
N. L. Britton, Fritsch mis-spelled Britton's name, but his original spelling must stand.

22. Licania unguiculata Prance, sp nov Fig 9 F-K.

Arbor magna, ramulis juvenilibus puberulis mox glabris, lenticellatis. Folia

alternata petiolata; petiolo 2.0-5.0 mm longo, tereti, glabro, rugoso, subtus cum
glandulis duobus munito; laminae ovatae vel ellipticae, coriaceae, 5.5-I6.0 cm
longae, 3.5-6.8 cm latae, basi rotundatae vel subcuneatae, apice in acuminem 3.0-
15.0 mm longum contractae, utrinque glabrae, subtus glandulosae; costa media
supra prominente, glabra; costis secundariis 7-Io jugis, subtus prominentibus, supra
leviter prominentibus. Stipulae lanceolatae, 2.0-4.0 mm longae, intrapetiolares,
membranaceae, persistentes. Flores 5.0-6.0 mm longi, pedunculati, in paniculis
racemosis terminalibus subterminalibusque dispositi, rachi ramisque puberulis.
Bracteae bracteolaeque 0.3-1.0 mm longae, ellipticae, persistentes, extus puberulae.
Receptaculum campanulatum, extus tomentellum, intus dense villoso tomentosum;
pedicelli 2.0-2.5 mm longi. Calycis-lobi acuti, utrinque tomentelli. Petala 5, pubes-
centia, unguiculata, persistentia, calycis-lobos excedentia. Stamina circa I4, in
orbem completum disposita, filamenta glabra, calycis-lobos duplo excedentia,
pubescentia, basi connata. Ovarium ad basim receptaculi insertum, villoso-tomen-
tosum. Stylus e basi ovarii ortus, calycis-lobos excedens, ad apicem hirsutus, Fructus
immaturus elongato-cylindricus; epicarpio laevi; pericarpio fibroso, intus glabro.
TYPE. DuckeRB 23599, Brazil, Rio Negro, above mouth of Rio Curicuriari, fl
(holotype, K; isotype, RB).
 Licania 61

.I .(V/;.\

B C

FIG 9. Spec~~~ieso iai.AE.wraki(aur ta328;A ai,x05 ,foe,x5

C , flowrscin ;D lwrpbsec,x5 , ri,xx(re 73) -,L nucl

(F6s
,3);F abtxo.;Gflwrx5H,foeseto,x;,flwrpecee,5;,

li''IY ~
peta~(~?J~ G FV, x7.5

FIG 9. Species of Licania.A-E, L. wurdackii(Maguireet al 36238); A, habit, x o5; B, flower, X 5;

C, flower section, x ; D, flower pubescence, x 5; E, fruit, X i (Frdes 27934). F-K, L. unguiculata
(Froes 21133); F, habit, x o.; G, flower, x s; H, flower section, x s; J, flower pubescence, x s; K,
petal, x 7.5/
62 Flora Neotropica

DISTRIBUTION. Non-flooded forest in western Brazilian Amazonia. Flowering

October to December.
BRAZIL. Amazonas: DuckeRB 25025 fl (K, RB); Krukojf8640 fl (A, BM, BR, F, K, MO, NY,
P, S, US); FerreiraINPA 5880 fl (INPA); Frdes21133 fl (F, IAN, K, NY, US); Oliveira2285 fl (IAN).
Terr. Roraima: KuhlmannRB 3061 fr (K, RB). Para: (cult), Silva 59729 fl fr (IAN, NY).

Although this distinct species is placed in section Moquilea,the stamens are only
shortly exserted. They appear less exserted because of the unusually long, persistent
petals, which hide the filaments. Licaniaunguiculata is closely related to L. longipetala,
both species possessing unguiculate exserted petals which are unique in the genus.

23. Licania longipetala Prance, sp nov

Arbor magna, ramulis juvenilibus glabrescentibus mox glabris, lenticellatis.

Folia alternata petiolata; petiolo 3.0-5.0 mm longo, tereti, glabro, rugoso, subtus
cum glandulis duobus munito; laminae oblongo-ellipticae vel ellipticae, coriaceae,
8.o-i6.o cm longae, 3.0-6.5 cm latae, basi rotundatae vel subcuneatae, apice in
acuminem 5.0-I5.0 mm longum contractae, utrinque glabrae, sutus glandulosae;
costa media supra leviter prominente, glabra; costis secundariis 9-12 jugis, subtus
prominentibus, supra leviter prominentibus. Stipulae lanceolatae, circa 3.0 mm
longae, intrapetiolares, membranaceae, persistentes. Flores 3.0 mm longi, haud
pedunculati, in paniculis racemosis terminalibus subterminalibusque dispositi,
rachi ramisque puberulis. Bracteae bracteolaeque 0.3-I.0 mm longae, ovatae,
persistentes, extus puberulae. Receptaculum campanulatum, subsessile, extus
griseo-tomentellum, intus dense villoso-tomentosum; pedicelli 0-0.25 mm longi.
Calycis-lobis acuti, utrinque tomentelli. Petala 5, pubescentia, unguiculata, persis-
tentia, calycis-lobos excedentia. Stamina circa 14, in orbem completum disposita,
filamenta glabra, calycis-lobos dupla excedentia, pubescentia, basi connata. Ovarium
ad basim receptaculi insertum, glabrescentum vel glabrum. Stylus e basi ovarii ortus,
calycis-lobos excedens, sparse hirsutus. Fructus mihi ignoti.
TYPE. Schunke288, Peru, Loreto, Rio Mazan, Gamitanacocha, fl (holotype, NY;
isotypes, A, F, S, US).
DISTRIBUTION. Periodically flooded forest in Amazonian Peru and Brazil.
Flowering January to April.
BRAZIL. Amazonas: Borba, Rio Madeira, Ducke RB 23600 fl (K, RB); ParanA do Autaz
Mirim; Rodrigues& Mello 7771 fl (INPA 17077, NY).

LOCALNAMEANDUSES.Peru: Apacharana.The Chamas Indians of Peru burn the

bark, and use the ashes in ceramics to harden the clay used forjars, pitchers, etc.
This species is close to L. unguiculata,but distinct by the much smaller, nearly
sessile flowers and the glabrous ovary.

2. Licania subgen Moquilea sect Leptobalanus

The species of this section are very closely related and constitute about the most
complex species group in the genus. The floral structure is uniform, the only differ-
ence being small variations in size and shape and in stamen number. Although there
is some variation in the fruits, it is hard to use these organs in the taxonomy because
of the inadequate fruiting material presently available. The most reliable differences
Licania 63

are in the inflorescenceand in vegetative characters. Because of the complexity of the

group, and because many small and insignificant characters have been used hitherto
as a basis for a new species, it is necessary to reduce many of these into synonymy.
The species of section Leptobalanus fall into two groups. I. Those with glabrous or
lanate-pubescent leaf undersides, with no trace of stomatal cavities, and 2. those
with stomatal cavities on the leaf underside. Although the presence or absence of
stomatal cavities divides the group nicely into two, in the past there has been con-
fusion between these groups because this very character has been largely overlooked.
L. apetala(without cavities) and L. octandra(with cavities) were both described in the
early nineteenth century, and many later authors have confused these two species,
citing synonyms under the wrong species.
In the group without stomatal cavities, ten species are recognized. One species,
L. apetala,has a wide geographical range and represents the center of the variation.
The other nine are more local segregates. The position is similar in the group
with stomatal cavities, where seven species are recognized. One, L. octandra,extends
over a wide geographical range, and is very variable. The two species L. apetalaand
L. octandrarequire more experimental ecological study in order to elucidate their
biology.

24. Licania wurdackii Prance, sp nov Fig 9 A-E, Fig Io.

Arbor parva, ramulis juvenilibus lanato-arachnoideis pubescentibus mox

glabris, haud lenticellatis. Folia alternata petiolata; petiolo 5.0-9.0 mm longo, dense
tomentoso, glanduloso, leviter canaliculato vel tereti; laminae oblongae vel oblongo-
ovatae, coriaceae, 5.5-10.0 cm longae, 2.2-5.3 cm latae, basi rotundatae vel sub-
cuneatae, apice obtusae, juveniles cum pilis deciduis utrinque lanato-pubescentes
mox glabrae, subtus glandulosae; costa media leviter prominente; costis secundariis
7-10 jugis, subtus prominentibus, supra planis. Stipulae lineares, membranaceae,
3.0-4.0 mm longae, subpersistentes. Flores circa 4.0 mm longi, sessiles vel pedun-
culati, elongati cymosi, in paniculis terminalibus subterminalibusque dispositi, rachi
ramisque dense flavo-brunneis piloso-tomentosis. Bracteae primariae lanceolatae, ad
I2.0 mm longum, caducae, aliae bracteae bracteolaeque lanceolatae vel ovatae,
eglandulosae, ad 3.0 mm longum, persistentes, extus tomentosae, intus glabrae.
Receptaculum campanulatum, sessile, extus tomentosum, intus villoso-tomentosum.
Calycis-lobi acuti, utrinque pubescentes. Petala nulla. Stamina 8-o0, in orbem
completum disposita, filamenta glabra, longe exserta, ad basim libera. Ovarium ad
basim receptaculi insertum, basim versus dense villoso-pubescens, apice sparse
villosum. Stylus e basi ovarii ortus, calycis-lobos longe excedens, subtus villosus,
supra glaber. Drupa lanceolato-fusiformis, extus laevis, glabra sparse verrucosa;
mesocarpio tenue; endocarpio tenue, fragile, intus sparse hirsuto.
TYPE. Maguire, Wurdack& Bunting36238. Venezuela, Terr. Amazonas, Rio
Atabapo, fl (holotype, NY).
Riverine forest and river banks of Amazonian
DISTRIBUTION. Venezuela and
adjacent regions. Flowering August to January.
VENEZUELA. Terr. Amazonas: Rio Atabapo, Maguire, Wurdack& Bunting 36215 fl (NY);
36255 fl (NY); Pto. Ayacucho, LI. Williams 14961 fr (A, F, US, VEN); 15951 fr (F, US, VEN). Apure:
Rio Cinaruco, Wurdack& Monachino41350 fl (NY); Without precise locality: Ernstsn fl (US). BRAZIL.
Amazonas: Rio Negro, DuckeRB 25026 fl (K); Fr6es 27934 fr (NY, R, US).

This species is closest to L. apetala,from which it differs in the pubescence of the

inflorescence and young leaves, the blunter leaves, and the larger flowers.;
64 Flora Neotropica

+DatBA
cpid

G 14,

'
L

FIG io. Variation and geographical distribution of Licania apetalaand related species. ? =-L.
cuspidata; + =L. wurdackii;A =L. apetalavar aperta; ? =L. apetala var apetala; A-L, Licaniaapetala,
leaves, X 0.3; flowers and fruit, x o.6; A, Maguire et al 35673; B, Maguire et al 30769; C, Berti I30; D,
Pranceet al 4271; E, B. W. 4811; F, Pranceet al 4768; G, Rodrigues446; H, Pranceet at 3361; J, Black et a
50-10747; K, Pranceet al 59622; L, Pires et al 1 140.
Licania 65

25 Licania turbinata Bentham, Jour. Bot. Hooker 2: 218. 1840; Fritsch, Ann.
Naturh. Mus. Wien 4: 55. 1889.
(Bentham)Hookerf., Mart. Fl. Bras. 14(2): 23. 1867.
Moquileaturbinata

Tree to I5.0 m or shrub, the young branches glabrous, not lenticellate. Leaves
ovate, 3.0-I I.0 cm long, 2.0-6.0 cm broad, coriaceous, obtuse to bluntly acuminate
at apex, with acumen 0-4.0 mm long, rounded at base, glabrous on both surfaces;
palisade glands absent; midrib prominulous above, glabrous; primary veins 8-II
pairs, prominulous on both surfaces; petioles 3.0-6.0 mm long, sparsely pubescent
when young, soon becoming glabrous, eglandular, shallowly canaliculate. Stipules
caducous. Inflorescencesshort terminal panicles to I .0ocm long, the primary branches
short, 0.5-1.8 cm long, the rachis and branches sparsely gray-puberulous. Flowers
4.0 mm long, in small cymules on short secondary branches of inflorescence. Bracts
and bracteoles 0.5-I.0 mm long, ovate, eglandular, sub-persistent, the margins
entire or serrulate. Receptacle broadly campanulate, gray-puberulous on exterior,
villous within; pedicels ca 0.5 mm long. Calyx lobes acute, puberulous on both
surfaces. Petals absent. Stamens II-12, inserted in a complete circle; filaments ex-
ceeding calyx lobes, free almost to base, glabrous. Ovary inserted at base of receptacle,
villous at base, glabrous above. Style hirsute on lower portion only, equalling fila-
ments. Young fruit globose; epicarp smooth, glabrous, drying black; mesocarp thin;
endocarp hard, thin, the interior hirsute.
TYPE.Gardner1149, Brazil, Pernambuco, fl (holotype, K; isotypes, BM, CGE,
GH, NY, OXF, P, S, US, W).
DISTRIBUTION. Coastal forest and restinga forest in eastern Brazil.
BRAZIL. Bahia: Belem & Pinheiro3130 fl (NY, UB); 3136 fl (NY, UB); 3144 fl (NY, UB);
3165 fl (NY, UB); 3191 fl (NY, UB); 3341 fr (NY, UB); Duarte 8057 fl (NY, RB); Riedelsn (I821) fl
(NY, US).

When Bentham first described this species, he created a new section for it.
However, there was no apparent ground for his section Batheogyne,and it was not
used by Hooker (1867) in his account of the genus in FloraBrasiliensis.

26. Licania lata Macbride, Candollea 5: 369. I934; Fl. Peru I07. I938.

Tree to 20.0 m tall, the young branches villous, becoming glabrous with age,
not lenticellate. Leaves oblong-elliptic to elliptic, coriaceous, 9.o-2I.o cm long,
3.0-8.5 cm broad, acuminate at apex, with acumen 7.0-20.0 mm long, rounded to
subcuneate at base, glabrous above, with persistent lanate-pulverulent pubescence
beneath, becoming sparse with age; palisade glands rare; with 2 large sessile glands
at junction of lamina and petiole on lower surface; midrib prominent above, glabrous;
primary veins IO-I2 pairs, prominent on both surfaces; petioles 6.o-Io.o mm long,
puberulous, becoming glabrous, terete or shallowly canaliculate, eglandular. Stipules
caducous. Inflorescences terminal and axillary racemose panicles, the rachis and
branches brown-puberulous, sometimes arachnoid when young. Flowers 4.0-5.0 mm
long, sessile on primary branches of inflorescence or on short peduncles in small
groups. Bracts and bracteoles I.0-4.0 mm long, ovate, persistent, puberulous on
exterior, the margins entire or serrulate. Receptacle campanulate-cupuliform, sessile,
tomentose on exterior, densely villous within. Calyx lobes acute, tomentellous on ex-
terior, puberulous within. Petals absent. Stamens ca I4, inserted in a complete circle;
filaments far exceeding calyx lobes, free to base, glabrous. Ovary inserted at base of
66 Flora Neotropica

receptacle, densely pilose. Style villous almost to apex, equalling filaments. Fruit
globose, 2.0-3.0 cm in diameter; epicarp smooth, glabrous, drying black; mesocarp
very thin, fleshy; endocarp very thin, 0.5 mm thick, hard, sparsely hirsute within.
TYPES.Klug 560, Peru, Loreto, fl (holotype, F; isotype, NY); 587 (paratypes,
NY, US).
DISTRIBUTION. Non-flooded forest and secondary scrub on sandy soil in western
Amazonia. Flowering October to February.
COLOMBIA. Amazonas: Garcia-Barriga13863 fr (COL, US). VENEZUELA. Terr. Amazonas:
Maguire, Wurdack& Bunting36758 fl (GH, NY, US); 36767 fl (F, NY, US). PERU. Loreto: Schunke350
fl (A, F, IAN, NY, P, S. US). BRAZIL. Amazonas: ChagasINPA 355 fl (IAN, INPA, NY); Chagas&
CoelhoINPA 3438a fr (IAN, INPA, NY); CoelhoINPA 2911 fl (IAN, INPA, NY); INPA 2998 fl (IAN,
INPA); INPA 3057a fr (INPA, NY); Ducke I90o fl fr (F, IAN, NY, R, US); RB 23596 fl (K, RB); Pires
& Black o102 fl (GH, IAN); INPA 603 fr (INPA, NY); 6062 fr (IAN, INPA, NY); Rodrigueset al 2772 fl
(INPA 9315); 2805 fl (INPA 10155); 3304 fr (INPA Io67I); 3863 fl (INPA 10428); 4086 fr (INPA
10651); 4105 fr (INPA Io670, NY); 4196 fr (INPA 10762, NY); 4919 fl (INPA 13452, NY); 5018 fr
(INPA 13679, NY).

LOCALNAMES.Brazil: Caripe, Macucu bobo.

27. Licania apetala (E. Meyer) Fritsch, Ann. Naturh. Mus. Wien 4: 54. I889;
Kleinh. in Pulle, Fl. Suriname 2(I): 443. 1939, Fig. Io.
HirtellaapetalaE. Meyer, Nova Acta Acad. Leop.-Carol. 21: 803. 1825; DC., Prodr. 2: 529. I825.

Tree to 40.0 m tall, the young branches glabrous or sparsely puberulous, be-
coming glabrous, not lenticellate. Leaves oblong-ovate, elliptic, or oblong-lanceolate,
coriaceous, 3.7-I4.0 cm long, 1.3-6.0 cm broad, the apex usually with well developed
acumen 3.0-I8.0 mm long, rarely obtuse to rounded, rounded to subcuneate at base,
glabrous on upper surface, glabrous or less often with persistent or caducous lanate
pubescence on lower surface; midrib prominulous above, glabrous or sparsely
pubescent when young; primary veins 7-12 pairs, prominent on lower surface,
prominulous above; petioles 3.0-6.0 mm long, glabrous or sparsely pubescent when
young, eglandular, terete. Stipules linear to 4.0 mm long, membraneous, rarely
persistent. Inflorescencesracemose panicles, the rachis and branches gray-puberulous.
Flowers 2.0-3.0 mm long, in small groups of cymules on short secondary branches
(peduncles) of inflorescence or sessile on primary branches of inflorescence. Bracts
and bracteoles 0.2-I.0 mm long, ovate to lanceolate, persistent. Receptacle cam-
panulate, short-puberulousto tomentose on exterior, tomentose within, sessile or with
short pedicels to 5.0 mm long. Calyx lobes acute, puberulous to tomentose on exter-
ior, puberulous to glabrous within. Petals absent. Stamens ca Io, inserted in a com-
plete circle; filaments far exceeding calyx lobes, free almost to base. Ovary inserted
at base of receptacle, villous. Style villous at base only, glabrous above, equalling
filaments. Fruit globose to narrowly lanceolate-fusiform; epicarp smooth, glabrous;
mesocarp very thin, fleshy; endocarp thin, fragile, fibrous, sparselypubescent within.

Key to the Varieties of Licaniaapetala

I. Flowers borne in small groups of cymules on distinct secondary branches of the inflorescence,
more than 2.0 mm long. Fruit usually elongate-lanceolate. a. var apetala.
I. Flowers predominantly sessile on primary branches of inflorescence; fruit globose.
b. var aperta.
Licania 67

27a. Licania apetala var apetala

Licania pendula Bentham, Jour. Bot. Hooker 2: 218. I840. Type. Schomburgk906, Guyana, fl
(holotype, K; isotypes, BM, CGE, F, GH, LE, NY, OXF, P, US, W).
Moquileapendula(Bentham) Hooker f., Mart. Fl. Bras. I4(2): 22, t. 5. 1867.
Licania apetalavar pendula(Bentham) Fritsch, Ann. Naturh. Mus. Wien 4: 55. 1889.
LicaniafloribundaBentham, Jour. Bot. Hooker 2: 219. 1840. Type. Schomburgk897, Guyana, fl
(holotype, K; isotypes, BM, CGE, F, GH, L, OXF, P, US).
Moquileafloribunda(Bentham) Hooker f., Mart. Fl. Bras. 14(2): 2I. 1867.
Moquilea turiuvasensu Hooker f., Mart. Fl. Bras. 14(2): 25. I867 pro parte, non Licania turiuva
Chamisso & Schlechtendal.
LicaniadahlgreniiStandley, Publ. Field Mus. Bot. I7: 255. I937, synon nov. Type. Dahlgren876,
Brazil, Piaui, fl (holotype, F; isotypes, P, S, US).
Licania hylaea Cuatrecasas, Brittonia 8: 198. 1956, synon nov. Type. Garcia-Barrigar14oo,
Colombia, Amazonas, fl (holotype, US; isotypes, COL, NY).
MoquileaorinocensisRusby, Descr. New Sp. S. Am. P1. 27. 1920, synon nov. Type. Rusby& Squires
426 fr (holotype, NY; isotypes, F, K, M, MO).

Flowers borne in small groups of cymules on distinct secondary branches of

inflorescence which are more than 2.0 mm long; fruit usually elongate-lanceolate.
TYPE. Meyersn, Suriname, fl (GOET, M).
DISTRIBUTION. Periodically flooded forest, river margins, savanna margins, and
open river beaches on sandy soil; abundant in the Guianas and Amazonia and
frequent in gallery forest further south.
Representative collections: COLOMBIA. Vaup6s: Gutierrez& Schultes567 fl (NY); 883 fl (GH);
Maguire& Wurdack35673 fr (F, NY, US); Schultes& Cabrera13085 fl (COL, US); 16662 fl (BM, BR, F,
GH, NY, US). Amazonas: Schultes& Black 8288 fl (F, GH, K, NY, US). VENEZUELA. Anzoategui:
Steyermark 61435 fl (NY). Terr. Delta Amacuro: Steyermark 93070 fl (NY). Bolivar: Ll. Williams 12603 fr
(F, US, VEN); 15077 fr (VEN); Wurdack& Monachino41138 fl (NY); 40971 fl (F, GH, MO, NY, RB,
US); 41o39 fl (F, GH. MICH, NY, US). Terr. Amazonas: Maguire, Cowan& Wurdack30769 fl (NY,
US); Maguire& Wurdack34594 fl (NY, US); 34781 fl (GH, NY, US); Maguire,Wurdack& Bunting37526
fl (F, GH, NY, US); 37578 fl (NY, US); Maguire, Wurdack& Maguire42635 fl (F, GH, MO, NY, US);
Wurdack& Adderley42735 fl (F, MO, NY); Ll. Williams 13849 fl (F, VEN). GUYANA. De la Cruz3963
fl (F, GH, MO, NY, US); 4539 fl (F, GH, MO, NY, US); Jenman 6214 fl (K); 7316 fl (K). SURI-
NAME. BBS 0o32 st (BBS, K, U); 1124 st (BBS, U); B.W. 51 fl (U); 488 fl (U); 1446 st (U); 3352 fl
(A, U); 4io6fl (IAN, K, NY); 4811 fl (K, U); 608 fl (A, U); 6io6fr (U);Focke 1368fl (U); Hostmann
363 fl (BM, F, K, LE, NY, OXF, P, S, U); Kappler2117 fl (GOET, NY, P); Mennega316fl (A, BR, U);
357 fl (A, U); Stahel 6i fl (A, NY, U); Versteeg389 fl (U); 422 fl (U). FRENCH GUIANA. For. Serv.
116I fr (U); 1175 st (U); 1317 fl (U); Lemeesn fl (P); Leprieursn fl (A, BM, BR, F, GH, K, LA, NY, P,
US); Martin 157 fl (BM); Melinonr18 fl (P); 199 fl (P); Patrissn fl (F); Perrottetsn fl (P). PERU. Loreto:
Schunke53 fl (F, NY, US); Klug 729 fl (F, NY); Wurdack2524 fl (F, LE, MICH, NY, US). BRAZIL.
Amazonas: CoelhoINPA 7264 fr (INPA); Ducke MG 11580 fr (BM, MG, RB); RB 25027 fl (K, RB);
Frommet al 1401 fl (R); Froes 22061 fr (IAN, NY, U); 22449 fl (IAN); 22863 fl (IAN, NY); 25198 fl
(IAN, NY, RB); 28133 fl (IAN, UB); 33572 fr (IAN); 33736fl (IAN); KuhlmannRB 17979 fl (K, RB);
17980 fr (K, RB); Mello & CoelhoINPA 3335 fl (INPA, NY); Oliveira2842 fl (IAN); Pires 8i fl (NY);
Pires & CoelhoINPA 13241 fl (INPA, NY); Pires & Silva 7972 fl (IAN); Pranceet al 3675 fl (INPA, NY,
SP); 3692 fr (INPA, NY); 4660 fl (INPA, NY); 4768 fr (INPA, NY); Rodrigueset al 207 fl (IAN, INPA
4317, MG); 289 fl (IAN, INPA 4399, MG); 2040 fl (INPA 8417); 4786 fl (INPA 13313, NY); 5004 r-
(INPA 13665, NY); 7295 fl (INPA 16633, NY); Spruce534 fl (K); 1179 fr (NY); 1230 fl (BM); 1437 fl
(K, P); 1830 fl (GH, K, LE, NY, OXF, P); 1831 fl (BM, K, LE, NY, P); 2707 fl (BM, BR, C, LE, P,
OXF). Terr. Roraima: Kuhlmann121 fl (INPA, RB 2809); 927 fl (RB 4538); Pranceet al 3974 fl (INPA,
NY); 4I33 fl (INPA, NY). Para: DuckeMG 6904 fl (MG, RB); MG 6905 fl (BM, RB); I5566 fl (MG,
RB); Ducke RB 19797 fl (K); Black & Ledoux50-o1634 fl (IAN, NY); 50-10690 fl (IAN, NY); 50-
o0747 fl (IAN, NY, UB); Oliveira6 fr (IAN); Pires 51856 fl (F, MO, NY, UB); Pires & Silva 4355 fl
(IAN, NY); Prance& Silva 58948 fl (F, NY, UB, US); Prance& Pennington1690 fl (F, GH, IAN, MO,
NY); Silva 637 fl (IAN, UB). Maranhao: Froes33991 fl (IAN); Viana2 fl (RB); Piaui: LisboaMG 2377
fl (RB 15197). Ceara: Lisb?aMG 2428 fl (RB 15221). Mato Grosso: KuhlmannRB 17891 fl (K); 17978
fl (K, RB). Goias: Prance& Silva 59622 fl (F, NY, UB, US); 59629 fl (NY, UB). Dist. Federal: Irwinet
al 10176 fl (NY). Guanabara: Glaziou 10701 fl (K, P).

LOCAL NAMES.Colombia: Ta-fee-a (Tarimuka), Ran-hoo (Makuna), Amaree

68 Flora Neotropica

0 - 70 60 50 40

, ., ~ . _ _
/'_3
10,0~~4

*.^
I Iv\ 'c \ - .....

FIG i . Geographic distribution of species of Licania. * L. longistyla,A L. sclerophylla,4 L.

fuchsii, ? L. albiflora.

(Yakuna). Venezuela: Mamoncillo. Suriname: Kauta, Kwepi. Brazil: Cariperana,

Caripe.

27b. Licania apetala var aperta (Bentham) Prance, stat nov

LicaniaapertaBentham, Jour. Bot. Hooker 2: 218. I840.
Licania pubifloraBentham, Jour. Bot. Hooker 2: 219. I840. Type. Schomburgk136, Guyana, fl
(holotype, K; isotypes, BM, CGE, NY, OXF, P).
Licania affinis Kuntze, Rev. Gen. I: 217. I891, nom. illegit., non Fritsch. Type. Kuntze o014,
Trinidad, fl fr (K, US).
LicaniakuntzeanaUrban, Symb. Antill. 5: 353. I908, synon nov. Type. Kuntze 1014, Trinidad fl
fr (K, US).
Moquileakuntzeana(Urban) R. 0. Williams, Fl. Trinidad & Tobago I: 314. 1932.
LicaniacaracasanaKlotzsch ex Hooker f., Mart. Fl. Bras. 14(2): 25. I867 nom nud in syn.

Flowers predominantly sessile on primary branches of inflorescence; fruit

globose.
TYPE.Schomburgk 593, Guyana, fl (holotype, K; isotypes, BM, BR, CGE, GH, L,
NY, OXF, P).
DISTRIBUTION. Riverine forest, river banks and fresh water beaches, and savanna
margins of the Guianas and Amazonia.
Representative collections: VENEZUELA. Zulia: Tejera79 fl (US). Portuguesa: Pittier I2073 fl
(A, NY, US, VEN). Dist. Federal: Otto 969 fl (BR, GH, K, NY, P). Miranda: Aristeguieta513r fl
Licania 69

(VEN). Guarico: Aristeguieta4505 fl (VEN); 5954 fr (VEN). Anzoategui: Aristeguieta3004 fl (NY,

VEN). Terr. Delta Amacuro:Berti120 fl (BR, F, NY, P, U); 130 fr (BR, NY, RB). Barinas:Bernardi
o08i fl (K, VEN). Bolivar: Aristeguieta5889 fl (VEN); Bailey & Bailey 1422 fl (A, US); 1438 fl (US);
I658 fl (US); Maguire, Steyermark& Maguire 53500 fl (F, NY, US); LI. Williams 12678 fr (VEN);
12679 fr (F, US); Wurdack215 fr (NY, US); Wurdack& Monachino41254 fl (A, F, MICH, NY, RB,
UB, US). TRINIDAD. Bot. Gdn. 5700 fl (TRIN); Britton & Hazen 691 fr (GH, NY, US); Broadway
6761 fl (BM, K, NY). GUYANA. Qyelch& McConnell308 fl (BM, K); Schomburgk 230 fl (BM, NY, P);
252 fl (P); 349 fl (K); 402 fl (K); A. C. Smith3275 fl (F, IAN, K, LE, MO, NY, P, U, US); 3519 fr
(A, F, IAN, K, LE, MO, NY, P, U, US). BRAZIL.Amazonas:Cardona
1254fl (F, US, VEN); Ducke
2212 fl (COL, IAN, NY, P, R); RB 23595 fl (K, RB); Poeppig2877 fl (BR, GOET, P); 2880 fl (OXF);
Pranceet al 3361 fl (INPA, NY); Rodrigues& Osmarino5689 fl (INPA I4401, NY); 5699 fl (INPA
I4411, NY). Terr. Roraima: Kuhlmann927 fl (F, RB 4538); Pranceet al 4049 fl (INPA, NY); 4271 fr
(INPA, NY); M. Silva 107 st (MG); 191 fr (MG); Ule 7886 fl (K, MG, US). Para: DuckeMG 10492
fl (BM, RB); 10248 fl (BM, RB); MG 16590 fl (R); 16592 fl (R); Monteiroda Costa317 fl (IAN, R, US);
Pires et al 6320 fl (IAN, NY); SchwackeIII-163 fl (GOET, R, RB); Sprucesn fl (NY). Acre: Prance
et al 2692 fl (INPA, NY). Terr. Rondonia: Silva 405 fl )NY). Mato Grosso: Prance & Silva 59129 fl
(NY, UB); 59394 fl (F, MO, NY, RB, UB, US); 594r1 fl (NY, RB, UB, US). Goias: Burchell7185 fl
(GH); Prance& Silva 59616 fl (F, NY, RB, UB, US); 59629 fl (F, NY, RB, UB, US); 59637 fl (F, NY,
RB, UB, US).

LOCALNAMES.Venezuela: Mamoncillorebalsero,Tacamahaco.
Brazil: Cariperana.

This species exhibits much more variation than most others of Licania.However,
the different variables are not correlated, and there is a complete range with no well
defined gaps. If this species were to be treated as more than one, it would be necessary
to describe a great many, based on minutiae, with no geographical correlations. The
principal and most striking variant within this species is regarded here as a variety.
The species as a whole obviously requires experimental study.
Several of the synonyms cited above reflect the confusion that has attended
earlier interpretations. One important difference in the synonomy presented here is
that several of the names cited were previously associated with L. octandra(L. turiuva
Cham. & Schl.). In FloraBrasiliensisHooker cited L. apertaand L. pubifloraas synonyms
of Moquileaturiuva.This was adopted by Fritsch (1889) in his conspectus of the genus.
Pilger (I923) correctly observed that L. apertadiffers from L. octandra in the nervation
of the leaf undersurface. L. octandraresembles L. apetalain most features, but differs
significantly in that the leaf underside has stomatal cavities. This earlier citation of
these two species as synonyms of L. octandrahas led to much material of L. apetala
being erroneously identified as L. octandra or as L. turiuva.

28. Licania parvifolia Huber, Bol. Mus. Emilio Goeldi 5: 367. 1909.
Licania arachnitesStandley, Lloydia 2: 181. 1939, synon nov. Type. A. C. Smith 2696, Guyana, fl
(holotype,F; isotypes,A, IAN, K, MAD, MO, NY, P, U, US).

Small to medium-sized tree, the young branches puberulous, soon becoming

glabrous, not lenticellate. Leaves ovate-elliptic to oblong, coriaceous, 3.3-8.5 cm
long, 1.3-3.5 cm broad, obtuse to acute at apex, cuneate to cordate at base, glabrous
above, caducous-lanate beneath; midrib prominulous above, glabrous; primary
veins 8- I pairs, prominulous on both surfaces; petioles 3.0-4.o mm long, hirsute
when young, becoming glabrous, shallowly canaliculate to terete, eglandular.
Stipules linear, to 4.0 mm long, membraneous, persistent. Inflorescences spreading
racemose panicles, the rachis and branches gray-puberulous. Flowers 3.0 mm long,
70 Flora Neotropica

in small cymules on short secondary branches of inflorescence or sessile on primary

branches. Bracts and bracteoles 0.5-I.0 mm long, ovate, pubescent on exterior,
entire, persistent, eglandular. Receptacle campanulate, sessile, tomentose to villous-
tomentose on exterior, tomentose within. Calyx lobes acute, tomentose on both
surfaces. Petals absent. Stamens 9-Io, inserted in a complete circle; filaments far
exceeding calyx lobes, glabrous, free almost to base. Ovary inserted at base of recept-
acle, pilose. Style pilose on lower portion only, equalling or exceeding filaments.
Fruit unknown.
TYPE.DuckeMG 8979, Brazil, ParA,fl (holotype, MG; isotype, G).
DISTRIBUTION. Open riverine localities, especially, river banks and islands in
Guyana and the southern part of Amazonian Brazil.
GUYANA. Boyan 9r (F. D. 7053) fl (K, MICH, NY, U); Drakesn fl (NY); Fanshawe1564 (F. D.
4300) fl (K, NY, U); Im Thurnsn fl (K); Persaud8 fl (F); 97 fl (F, K, NY, S, US). BRAZIL. Para
Froes34175 fl (IAN). Maranhao: GomesMourasn fl (INPA 20849). Mato Grosso: Kuhlmann312 fl (R);
Malme 2703 fl (R, S); Moore564 fl (BM, K, NY, P, R); Prance& Silva 59194 fl (F, NY, RB, UB).

This species is close to L. apetala,and is perhaps part of it, however, it differs in

the blunt, usually obtuse, leaves and in the habitat, apparently growing only beside
rivers in rocky places.

29. Licania maguirei Prance, sp nov Fig 12 A-D.

Frutex, ramulis juvenilibus puberulis mox glabris, haud lenticellatis. Folia

alternata petiolata, petiolo 3.0-7.0 mm longo, juvenilo hirsutulo, tereti, subtus sub
lamina cum glandulis duobus sessilibus munito; laminae oblongo-ovatae, coriaceae,
6.5-I7.0 cm longae, 2.4-6.5 cm latae, basi rotundatae vel subcordatae, apice in
acuminem 7.o-I7.0 mm longum contractae, utrinque glabrae, haud glandulosae;
costa media supra glabra, prominente; costis secundariis Io-I2 jugis, subtus promi-
nentibus, supra planis. Stipulae lineares, caducae. Flores circa 4.0 mm longi, haud
pedunculati, in paniculis dense multi-ramosis terminalibus subterminalibusque
dispositi, rachi ramisque dense griseo-tomentosis. Bracteae primariae ovatae ad 7.0
mm longum, caducae, bracteae aliae bracteolaeque, 1.5-3.0 cm longae, ovatae
persistentes extus tomentosae, margine serratae. Receptaculum cupuliforme, sessile,
extus tomentosum, intus villoso-tomentosum. Calycis-lobi acuti, utrinque tomentosi.
Petala nulla. Stamina circa I4, in orbem completum disposita; filamenta glabra, ad
basim libera, longe exserta. Ovarium ad basim receptaculi insertum, dense villosum.
Stylus e basi ovarii ortus, subtus villosus, calycis-lobos excedens. Fructus mihi ignoti.
TYPE. Maguire,Pires,Maguire& Silva56470, Brazil, Mato Grosso, Rio Juruena,
Brasilia to Acre highway, fl (holotype, NY).
DISTRIBUTION. Riverine forest, known only from the State of Mato Grosso,
Brazil. Collected in flower September to November.
BRAZIL. Mato Grosso: Serra do Roncador near Rio Turvo, Sidney1203 fl (UB); Chavantina,
Sidney350 fl (NY, UB).

Licania maguirei is most closely related to L. parvifolia from which it differs in the
larger flowers, the narrow acuminate leaves, and the larger number of stamens. It is
also close to L. gardneribut differs in the large flowers, the cupuliform receptacle, and
the more nearly plane venation of the leaf upper surface.
Licania 71

-- .

4ff

.... J

FIG 12. Species of Licania.A-D, L. maguirei(Maguireet al 5647o); A, habit, x o.5; B, flower, -x 5;

C, flower section, x 5; D, flower pubescence, x 5. E, L. silvatica(Glaziouo10702), habit, x o.5. F-K,
L. foveolata (Boyan28); F, habit, x o.5; G, flower, x 5; H, flower section, x 5; J, flower pubescence,
x 5; K, young fruit, x 5.
72 Flora Neotropica

30. Licania gardneri (Hooker f.) Fritsch, Ann. Naturh. Mus. Wien 4: 56.
I889.
MoquileagardneriHooker f., Mart. Fl. Bras. 14(2): 21. I867.
Pilger, Notizbl. Bot. Gart. Berlin 8: 540. 1923, synon nov. Type. Hoehne
Licania mattogrossensis
4262, Brazil, Mato Grosso, fl (B, lost; isotype, R; photo, NY).
Moquileamattogrossensis (Pilger) Malme, Ark. Bot. 23A(4): II. 1930.

Small tree or shrub, the young branches puberulous, soon becoming glabrous,
not lenticellate. Leaves elliptic to oblong, coriaceous, 9.0-I6.5 cm long, 5.o-8.5 cm
broad, acuminate at apex, with acumen 5.o-I2.o mm long, rounded to subcordate
at base, glabrous on both surfaces; palisade glands occasional on lower surface;
midrib prominulous above, glabrous when mature, sparsely pubescent towards base
when young; primary veins 1-14 pairs, prominent on both surfaces; petioles 4.0-
7.0 mm long, densely tomentose when young, terete, eglandular. Stipules linear,
membraneous, 4.0-5.0 mm long, subpersistent. Inflorescences terminal and subter-
minal racemose panicles, the rachis and branches sparsely gray-puberulous. Flowers
2.5-3.5 mm long, sessile or on short pedicels not exceeding I.5 mm long. Bracts and
bracteoles I.0-2.5 mm long, ovate, puberulous on exterior, entire, eglandular.
Receptacle campanulate, sessile, gray-puberulous on exterior, tomentose within.
Calyx lobes acute, tomentose on both surfaces. Petals absent. Stamens ca i2, inserted
in a complete circle; filaments far exceeding calyx lobes, slightly connate at base,
glabrous. Ovary inserted at base of receptacle, villous at base, glabrous above. Style
sparsely hirsute on lower half, equalling filaments. Fruit (immature) globose, ex-
terior smooth, glabrous.
TYPES.Gardner4539, Brazil, Minas Gerais, fl (lectotype, K; isolectotype, BM);
Gardner 2564, Brazil, Piaui, fl fr (paratypes,BM, CGE, F, G, K, M, NY, OXF, P, US);
Pohl i929, Brazil, Goias, fl (paratype, W); Pohl 2283, Brazil, GoiAs,fl (paratypes, NY,
W).
DISTRIBUTION. Riverine forest bordering cerrado,and in cerradoitself in the
Planalto of Central Brazil. Flowering May to January.
BRAZIL. Pard: Silva 709 fl (IAN). Maranhao: Lisb6a MG 2482 fl (RB I5195). Piaui: Snethlage
623 fl (F). Mato Grosso: Hoehne4549 fl (R); Irwin & Soderstrom6744 fl fr (NY); Irwin et al 16718 fl
(NY); s6888 fl (NY); I7034 fl (NY); Lima58-3083 fl (IPA); Malme 1757 fl (S); 1757a fl (S); Prance&
Silva 59117 fr (F, NY, RB, US); 59441 fr (F, NY, UB, US). Goiss: Hunt 6127 fl (K, NY); 6283 fl
(K, NY); Irwin et al 17707 fl (NY); 17923 fl (NY); Maguire,Pires, Maguire & Silva 56160 fl (F, NY,
UB, US); Pires & Black 1658a fl (IAN, NY); Pohl sn fl (probably paratype, BR); Prance& Silva 58569
fl (NY, RB, UB, US); 59514 fr (NY, UB, US); Sidney221 fl (UB); 279 fl (UB). Minas Gerais: Macedo
1128 fl (BM, MO, SP); 2550 fl (NY, S). Without precise locality: Martius991 fl (BM, K, MO, NY).

Malme (Ark. Bot. Stockh. 23A:(4): II. 1931) questioned whether L. matto-
grossensiswas distinct from L. gardneri,a doubt founded in fact as the names apply to
a single species.

31. Licania cuspidata (Rusby) Prance, comb nov Fig Io.

MoquileacuspidataRusby, Descr. New Sp. S. Am. P1. 27. 1920.

Tree to 20.0 m tall, the young branches glabrous, not lenticellate. Leaves oblong
to oblong-ovate, coriaceous, 4.5-8.o cm long, 1.5-3.5 cm broad, acuminate at apex,
with acumen 5.0-I0.0 mm long, rounded to subcuneate at base, glabrous on both
surfaces; palisade glands occasional; midrib prominent above, glabrous; primary
veins 0-I2 pairs, prominulous on both surfaces; petioles 2.0-3.0 mm long, glabrous,
eglandular, canaliculate. Stipules caducous. Inflorescences terminal and axillary
Licania 73

racemose panicles, the rachis and branches sparselyhirsutulous.Bracts and bracteoles

ca I mm long, ovate, hirsute on exterior, persistent, the margins slightly serrate.
Flowers 2.5-3.0 mm long, sessile on primary branches of inflorescence. Receptacle
campanulate, sessile, ferrugineous hirsutulous-tomentoseon exterior, villous-tomen-
tose within. Calyx lobes acute, tomentose on exterior, puberulous within. Petals
absent. Stamens Io, inserted in a complete circle; filaments far exceeding calyx lobes,
free to base. Ovary inserted at base of receptacle, densely villous. Style glabrous
except at very base, equalling filaments. Fruit unknown.
TYPE. H. H. Smith I773, Colombia, Magdalena, fl (holotype, NY; isotypes,
A, BM, F, GH, K, MICH, P, US).
DISTRIBUTION. Known only from the type collection from montane forest in the
Santa Marta mountains of Colombia.
This species is close to L. apetala,differing mainly in the hirsutulouspubescence
of the inflorescence and the shorter petioles.

32. Licania sparsipilis Blake, Contr. Gray Herb. 52: 67. I9I7.

Medium-sized tree, the young branches glabrous, or puberulous and soon be-
coming glabrous, not lenticellate. Leaves elliptic to oblong, coriaceous, 5.0-I3.0 cm
long, 2.2-5.5 cm broad, acuminate at apex, with acumen 6.o-I5.0 mm long, cuneate
to subcuneate at base, glabrous on both surfaces; palisade glands present beneath;
midrib prominulous above, glabrous; primary veins 8-o1 pairs, prominulous on both
surfaces; petioles 4.0-7.0 mm long, sparsely lanate becoming glabrous, shallowly
canaliculate, with two sessile glands. Stipules caducous (not seen). Inflorescences
terminal and axillary racemose panicles, the rachis and branches brown-tomentose.
Flowers 2.0 mm long, sessile on primary branches of inflorescence. Bracts and brac-
teoles ca i mm long, ovate, tomentose on exterior, entire to serrate, eglandular.
Receptacle campanulate, sessile, tomentose on exterior, lanate-tomentose within.
Calyx lobes acute, puberulous on both surfaces. Petals absent. Stamens 8-Io, in-
serted in a complete circle; filaments far exceeding calyx lobes, free to base, glabrous.
Ovary inserted at base of receptacle, villous at base, glabrescent above. Style
hirsute at base, glabrous above, equalling filaments. Fruit globose, 1.5-2.0 cm dia-
meter; epicarp smooth, glabrous; mesocarp thin, fleshy; endocarp thin, hard, glab-
rous within.
TYPE. Peck 858, British Honduras, fl (holotype, GH; isotypes, K, NY).
DISTRIBUTION. Forest on low wet ground in Guatemala and British Honduras.
GUATEMALA.Steyermark 38537 fl bud (F). BRITISH HONDURAS. Bartlett13095 fl (A,
MICH, NY, S); Gentle3535 fl (A, F, MEXU, MICH, MO, NY, U); 8584 fl (BM, F, MICH, S);
SchippI02 fl (A, BM, F, GH, K, MICH, MO, NY, S); 598 fr (A, BM, F, GH, K, MICH, MO, NY, S).

LOCALNAME. British Honduras: Bastard Pigeon Plum.

33. Licania emarginata Spruce ex Hooker f., Mart. Fl. Bras. 14(2): I5. 1867;
Fritsch, Ann. Naturh. Mus. Wien 4: 47. i889.

Medium to large-sized tree, the young branches glabrous, not conspicuously

lenticellate. Leaves ovate-elliptic to elliptic, coriaceous, 3.0-8.5 cm long, 1.5-5.0 cm
broad, rounded or obtuse to bluntly acuminate at apex, with acumen o-6.o mm long,
rounded at base, glabrous and shining on both surfaces; palisade glands absent;
74 Flora Neotropica

midrib glabrous and prominent above; primary veins 7-9 pairs, prominent beneath,
prominulous above; petioles 2.0-4.0 mm long, sparsely lanate when young, soon
becoming glabrous, deeply canaliculate, with two sessile glands at junction with
lower surface of lamina. Stipules minute, ovate, caducous, intrapetiolar. Inflores-
cences panicles, the rachis and branches puberulous. Flowers 2.0-3.0 mm long,
in small cymules on distinct secondary branches of inflorescence which are 3.0-10.0
mm long. Bracts and bracteoles 0.5-1.5 mm long, ovate, caducous, entire. Receptacle
campanulate, puberulous to gray-tomentose on exterior, rather sparsely tomentose
within; pedicels ca 0.5 mm long. Calyx lobes acute, puberulous on both surfaces.
Petals absent. Stamens Io-II, inserted in a complete circle; filaments shorter than
calyx lobes, glabrous, free to base. Ovary inserted at base of receptacle, pilose. Style
pilose at base, glabrous above, equalling filaments. Fruit unknown.
TYPE.Spruce2699, Brazil, Amazonas, fl (holotype, K; isotypes, BM, BR, CGE,
F, GH, GOET, LD, LE, NY, OXF, P).
DISTRIBUTION. Non-flooded forest in Venezuelan Guyana and Amazonia, and
Amazonian Brazil. Flowering November to January.
VENEZUELA. Bolivar: Wurdack& Monachino41099 fl (NY). BRAZIL. Amazonas: Frdes12435
fl (A, F, NY); Krukoff6860 fl (A, BM, BR, F, IAN, K, LE, MICH, MO, NY, RB, S, US); 7036 st
(A, BM, BR, F, IAN, K, LE, MICH, MO, NY, RB, S, US); 7058 fl (A, BM, BR, F, IAN, K, LE,
MICH, MO, NY, RB, S, US); 8761 fl (A, BM, BR, F, K, LE, MO, NY, P, S, US); 8869 fl (A, BM,
BR, F. K, LE, MO, NY, P, S); 8907 fl (A, BM, BR, F, K, LE, MO, NY, P, US).

This species differs from the other species of section Leptobalanus,except L.

in having included stamens. Since it fits in this section in all other respects I
calvescens,
have placed it here. This species clearly demonstrates that Licania and Moquilea
cannot be separated on the basis of the included or exserted stamens. Licaniaemarginata
resembles species formerly placed in Moquilea (subgenus Moquileain the present
account) in all respects except the included stamens which are characteristic of
Licania(or subgenus Licania).It is because of species such as this that I have followed
the workers who have considered Moquileaas part of Licania.Licaniacalvescensand
L. emarginata demonstrate that the staminal character is not reliable, and on the basis
of the other characters examined, I find no valid reasons for regarding Moquileaas a
distinct genus.

34. Licania calvescens Cuatrecasas, Fieldiana Bot. 27: 64. I950.

Large tree, the young branches puberulous, soon becoming glabrous, not
conspicuously lenticellate. Leaves oblong to oblong-lanceolate, coriaceous, 6.o-
10.5 cm long, 2.3-3.8 cm broad, cuspidate-acuminate at apex, with acumen 7.0-
i7.o mm long, subcuneate to rounded at base, glabrous above, the lower surfacewith
a ferrugineous-lanatepubescence which is easily removed; palisade glands frequent;
midrib prominent above, puberulous toward base; primary veins Io-II pairs,
prominent on lower surface, prominulous above; petioles 3.0-5.0 mm long, puberu-
lous, terete, eglandular. Stipules caducous (not seen). Inflorescencesterminal panicles,
the rachis and branches puberulous. Flowers ca 2.0 mm long, sessile on primary and
secondary branches of inflorescence. Bracts and bracteoles 0.5-1.5 mm long, broadly
ovate, persistent, minutely serrate to entire. Receptacle campanulate, sessile, puberu-
lous on exterior, tomentellous within. Calyx lobes acute, puberulous on both surfaces.
Petals absent. Stamens ca 22, inserted in a complete circle; filaments apparently
equalling calyx lobes (seen only in unopened buds), slightly connate at base. Ovary
inserted at base of receptacle, the lower portion villous-tomentose. Style villous at
base, glabrous above, equalling filaments. Fruit unknown.
Licania 75

TYPES.Cuatrecasas15835, Colombia, Valle, fl (holotype, F; isotype, COL);

17431, Colombia, Valle, st (paratypes, F, MAD).
Cuatrecasas
DISTRIBUTION. Known only from the type gatherings from the Pacific coastal
forest of Colombia.
The ovary is not altogether glabrous as stated in the original description. The
stamens are included in the type specimen because it is in bud only. However, the
filaments are not folded over in the buds and are probably not exserted in the open
flower. This can only be established with certainty when open flowers are collected.
Although I place this species in section Leptobalanus because of the apetalous flowers,
its position here is tenuous owing to the large number of stamens. Further material
of this species is required for certainity of its position within the genus.

35. Licania persaudii Fanshawe & Maguire, Bull. Torrey Club 75: 375. I948.

Tree to 30.0 m tall, the young branches glabrous, not conspicuously lenticellate.
Leaves elliptic to oblong, coriaceous, 9.0-20.0 cm long, 3.5-8.0 cm broad, abruptly
apiculate-acuminate at apex, with acumen 8.0-12 o mm long, subcuneate to rounded-
subcuneate at base, glabrous on upper surface, the lower surface with rather open
stomatal cavities obscured by a dense lanate pubescence at the mouth; midrib
prominent above, gabrous; primary veins 8-o0 pairs, prominent below; petioles
7.0-10.0 mm long, puberulous to glabrescent, transverselyrugose, terete to shallowly
canaliculate, with two glands at junction with lamina. Stipules caducous (not seen).
Inflorescences terminal and axillary racemose panicles, the rachis and branches
arachnoid-pubescent, becoming puberulous with age. Flowers 3.0-4.0 mm long,
mostly in small cymules on short distinct secondary inflorescencebranches 2.0-5.0 mm
long; some flowers sessile on primary branches. Bracts and bracteoles ovate, 2.0-
3.5 mm long, persistent, often slightly serrate, eglandular, sparsely pubescent on
exterior. Receptacle campanulate, the exterior short-gray-pubescent, villous-
tomentose within; pedicels 0-0.2 mm long. Calyx lobes acute, pubescent on both
surfaces. Petals absent. Stamens 9-10, inserted in a complete circle; filaments far
exceeding calyx lobes, glabrous, free to base. Ovary inserted at base of receptacle,
villous. Style villous at base, the upper portion glabrous, equalling or exceeding
filaments. Fruit globose, 1.8-2.0 cm diameter; epicarp smooth, glabrous, drying
black; mesocarp thin, fleshy; endocarp thin, fibrous, puberulous within.
TYPES.Persaud95, Guyana, fl (lectotype, NY; isolectotypes, F, K, US); De La
Cruz 1670, Guyana, fl (paratypes, F, GH, MO, NY, US); FanshaweIo86 (F.D. 3822),
Guyana, fl bud (paratypes, K, NY); I359 (F.D. 4095), fr (paratypes, K, NY, P, U);
1700 (F.D. 4436), fl (paratypes, K, NY).
DISTRIBUTION. Confined to primary forests of Guyana. Flowering January to
July.
GUYANA. Andersonsn fl (K); Davis 350 (F.D. 2343) fl (FHO, K); Fanshawe3417 (F.D. 698z) fl
(K, NY, U); Martyn273 fl (K); Persaud 6z1fr (F, K, NY, S).

LOCAL NAME.Kauta (Arawak).

36. Licania sprucei (Hooker f.) Fritsch, Ann. Naturh. Mus. Wien 4: 55. 1889.
MoquileaspruceiHooker f, Mart. Fl. Bras. 14(2): 22, t. 6. I867.

Small to medium-sized tree to 20.0 m tall, the young branches glabrous or

76 Flora Neotropica

puberulous. Leaves oblong to oblong-elliptic, thick-coriaceous, 6.0-20.0 cm long,

2.5-6.5 cm broad, acuminate at apex, with acumen 4.0-I5.0 mm long, rounded to
subcuneate at base, the upper surface glabrous, well developed stomatal cavities on
lower surface, venation plane, with slit-like apertures to cavities filled with lanate
pubescence; midrib prominent above, glabrous; primary veins 9- I pairs, prominent
below; petioles 6.o-io.o mm long, sparsely puberulous, becoming glabrous, terete to
shallowly canaliculate, rugose, with two glands on upper surface at base of lamina.
Stipules caducous (not seen). Inflorescencesterminal and axillary panicles, the rachis
and branches sparselypuberulous. Flowers ca 3.0 mm long, in small cymules on short
secondary inflorescence branches 2.0-5.0 mm long. Bracts and bracteoles ca i mm
long, ovate, persistent, serrate, often with stipitate glands. Receptacle campanulate,
puberulous on exterior, villous-tomentose within; pedicels ca 0.5 mm long. Calyx
lobes acute, puberulous on both surfaces. Petals absent. Stamens I0, inserted in a
complete circle; filaments exceeding calyx lobes, free almost to base, glabrous. Ovary
inserted at base of receptacle, villous. Style villous for most of length, equalling
filaments. Fruit globose; exterior smooth, glabrous, drying black; mesocarp thin,
fleshy; endocarp thin, fibrous, puberulous within.
TYPE.Sprucei80o, Brazil, Amazonas, fl (holotype, K; isotypes, BM, CGE, GH,
GOET, LD, LE, M, NY, P).
DISTRIBUTION. Primary forest on non-flooded ground and secondary forest in
southern Guyana and in the Manaus region of Brazil. Flowering June to October.
GUYANA. A. C. Smith 2611 fr (A, F, K, M, NY, P, S, US). BRAZIL. Amazonas: Ducke307 fl
(A, F, K, MO, NY, R, RB, S, US); RB 23597 fl (K, RB); RB 23598 fl (K, RB); Mello INPA 2086 fl
(IAN, INPA, MG, NY); Jobert434 fl (P); Labroysn fl (P); Pires & Black 1034 fl (IAN); Riedel i43I
fl (US). 'Guanabara: Glaziou9788 fl (K, P).'4

LOCAL NAME. Caripe.

37. Licania sclerophylla (Martius ex Hooker f.) Fritsch, Ann. Naturh. Mus.
Wien 4: 56. 1889. Fig. II.
MoquileasclerophyllaMartius ex Hooker f.. Mart. Fl. Bras. 14(2): 23, t. 7. 1867.
LicaniaasperaStandley, Publ. Field Mus. Bot. 17: 254. 1937, synon. nov. Type. Monteiroda Costa
279, Brazil, Para, fl (holotype, F; isotypes, IAN, US).

Small to medium-sized tree to 15.0 m tall, the young branches pubescent, be-
coming glabrous with age. Leaves elliptic to ovate-elliptic, coriaceous, 9.0-23.0 cm
long, 4.0-I0.5 cm broad, acute to acuminate at apex, with acumen o-8.o mm long,
rounded to cordate at base, the upper surfaceglabrous and papillose, the lower surface
bearing well developed stomatal cavities with flattened venation leaving small slit-
like lanate-pubescent apertures; midrib prominent above, lower portion densely
tomentose; primary veins 8-I o pairs, prominent beneath; petioles 6.o- 0.0 mm long,
terete, eglandular, densely tomentose. Stipules linear 3.o-7.0 mm long, mem-
braneous, intrapetiolar, subpersistent. Inflorescences paniculate, the rachis and
branches densely brown tomentose. Flowers 2.0-3.0 mm long, borne in clusters on
short secondary branches of inflorescence, and on primary branches. Bracts and
bracteoles I.0-3.0 mm long, lanceolate to ovate, usually entire, rarely slightly serrate.
Receptacle campanulate, tomentose on exterior, villous-tomentose within; pedicels
4 There are
many Glaziou collections labelled Rio de Janeiro which are in fact Amazonian plants.
Most of these were pirated from Schwacke and Jobert collections, and the Glaziou collection cited
here is undoubtedly an example of this. Throughout this monograph the pirated Glaziou collections are
indicated by single quotation marks.
 Licania 77

0.5-i.5 mm long. Calyx lobes acute, tomentose on exterior, puberulous within.

Petals absent. Stamens ca o1, inserted in a complete circle; filaments far exceeding
calyx lobes, free to base, glabrous. Ovary inserted at base of receptacle, densely
villous. Style villous on lower portion, equalling or exceeding filaments. Fruit
globose, to 3.0 cm diameter; epicarp densely appressed-pubescent; pericarp un-
differentiated, thin, fragile, sparselyvillous-pubescent within.
TYPES.Martius1467, Brazil, Minas Gerais, fl (lectotype, M); Pohl 2226, Brazil,
Minas Gerais, fl (paratypes, BR, M, NY, W); Spruce840, Brazil, Para, fl (paratypes,
K, NY, P); 987 fr (paratypes, K, NY, P).
DISTRIBUTION. Non-flooded forest, riverine forest or cerradoin eastern and
southern Amazonia and the Planalto of central Brazil.
BRAZIL. Par,: Black 48-3313 fl (IAN); Ducke MG 11397 fl (BM, INPA, MG, RB); 11969 fr
(MG); I2026 fl (MG, RB); Fires & Silva 4627 fl (NY); Pires, Black, Wardack& Silva 6325 fr (IAN, NY).
Terr. Amapa: Pires & Cavalcante52276 fl (F, GH, IAN, MG, NY, US). Maranhao: Frdes 11702 fr
(A, F, K, MO, NY, S, U, US); 11879 fl (A, F, MICH, NY, U); Lisboa MG 2458 fl (BM, MG, NY,
RB 5234, US); Pires & Black I745a fl (IAN, NY). Piaui: Snethlage669 fl (F). Terr. Rond6nia: Black &
Cordeiro52-14472 fl (NY); Pranceet al 6000 fr (INPA, NY). Mato Grosso: Kuhlmann296 fl (R); Prance&
Silva 59212 fl(NY, RB, UB, US); 59404 fr (NY, P, RB, UB, US); Sidney221 fl (UB). Goias: Burchell
7638 fl (GH); Macedo3624 fl (IAN, NY, RB, S); Prance& Silva 58479 fl (F, MO, NY, UB, US).

In the original description Hooker described three varieties, Moquileasclerophylla

var pohlianaHook. f., var scabraHook. f., and var myristicoides Hook. f., based on the
four type gatherings. These taxa were founded on small differences of leaf shape. In
light of the material at hand, it is not possible to consider these as varieties, because
there is a complete gradation in the leaf characters used by Hooker. Two of these
varieties, var scabraand var myristicoides, were maintained by Fritsch (i889), when
he transferredthis species to Licania.
Licania sclerophyllawas originally described in Moquilea under which Licania
sclerophyllaMart. mss was cited in synonymy. Although IndexKewensislists the name
as Licaniasclerophylla Mart. ex Hook. f., Fritsch (1889) was the first to use L. sclero-
phyllalegitimately, and he must be regarded as the author of the new combination.
is close to L. longistyla,but differs in a number of small, well corre-
L. sclerophylla
lated features, which are tabulated below.

L. sclerophylla
I. At least some flowers borne on
secondary branches of inflorescence, in
dense clusters
2. Inflorescence yellow-brown tomentose
3. Leaf base usually cordate less frequently
rounded
4. Upper surface of leaf usually papillose
5. Midrib usually pubescent above
6. Stamens twice as long as calyx lobes
L. longistyla
Flowers borne on primary branches of
inflorescence, solitary
Inflorescence gray tomentose
Leaf base rounded to subcuneate
Upper surface of leaf rarely papillose
Midrib usually glabrous above
Stamens 3-4 times as long as calyx lobes

38. Licania albiflora Fanshawe & Maguire, Bull. Torrey Club 75: 3I8. I948.
Fig. Io.
Tree to 25.0 m tall, the young branches puberulous, becoming glabrous with age.
Leaves obovate to oblong-elliptic, coriaceous, 11.0-21.0 cm long, 5.0-I I.0 cm broad,
acuminate at apex, acumen 3.0-6.0 mm long, rounded to subcuneate at base,
glabrous and papillose on upper surface, the lower surface with well developed
stomatal cavities with prominent unflattened venation describing rather open
78 Flora Neotropica

cavities, the mouth of each lanate-pubescent; midrib prominulous above, densely

pubescent when young, becoming glabrous with age; primary veins 7-IO pairs,
prominent beneath; petioles 7.0-14.0 mm long, tomentose, becoming less so with age,
eglandular, terete. Stipules caducous (not seen). Inflorescencesterminal and axillary
racemose panicles, the rachis and branches gray-brown villous-tomentose. Flowers
ca 2.5 mm long, solitary on primary branches of inflorescence. Bracts and bracteoles
1.5-4.0 mm long, lanceolate, entire, eglandular, the exterior tomentose. Receptacle
campanulate, tomentose on exterior, villous within; pedicels slender, 3.5-5.0 mm
long in flower, ca 2.0 mm in bud. Calyx lobes acute, tomentose on exterior, sparsely
puberulous within. Petals absent. Stamens I2-13, inserted in a complete circle;
filaments far exceeding calyx lobes, free to base, glabrous. Ovary inserted at base of
receptacle, densely villous. Style glabrous except at base, equalling or exceeding
filaments. Fruit unknown.
TYPE. Fanshawei678 (F.D. 4414), Guyana, fl (holotype, NY; isotype, K).
Primary forest of the Guianas, collected only twice.
DISTRIBUTION.
SURINAME. Maguire,Schulz, Soderstrom& Holmgren54100 fl (F, MO, NY, US).

This little known species is readily distinguished from the other related species
by the length of the pedicels.

LOCAL NAME. Suriname: Kwepi.

39. Licania longistyla (Hooker f.) Fritsch, Ann. Naturh. Mus. Wien 4: 56.
1889. Fig. I .
MoquilealongistylaHooker f., Mart. Fl. Bras. 14(2): 24, t. 8. 1867.

Small to medium-sized tree to 20.0 m tall, the young branches puberulous, soon
becoming glabrous. Leaves elliptic, coriaceous, 6.0-23.0 cm long, 3.5-I0.0 cm broad,
acuminate at apex, with acumen 3.0-I2.0 mm long, rounded to subcuneate at base,
glabrous on upper surface, rarely papillose, the lower surface bearing well developed
stomatal cavities with flattened venation leaving small slit-like lanate-pubescent
apertures; midrib prominent above, pubescent towards base, becoming glabrous
with age; primary veins 9-I2 pairs, prominent below; petioles 5.0-I3.0 mm long,
terete, eglandular, densely pubescent, becoming glabrescent with age. Stipules linear,
3.0-8.0 mm long, membraneous, intrapetiolar, subpersistent. Inflorescences terminal
and axillary racemose panicles, the rachis and branches gray-tomentellous. Flowers
ca 3.0 mm long, solitary on primary branches of inflorescence. Bracts and bracteoles
2.0-5.0 mm long, ovate to lanceolate, serrulate, usually with stipitate glands. Re-
ceptacle campanulate, gray-tomentose on exterior, densely villous-tomentose within;
pedicels 0.5-I.5 mm long. Calyx lobes acute, tomentose on exterior, puberulous
within. Petals absent. Stamens ca Io, inserted in a complete circle; filaments far ex-
ceeding calyx lobes, free to base, glabrous. Ovary inserted at base of receptacle,
villous. Style villous on lower portion, equalling or exceeding filaments. Fruit globose,
2.5-3.5 mm in diameter; epicarp with appressed sordid yellow-velutinous pubes-
cence; pericarp undifferentiated, thin, fragile, fibrous, sparsely villous within.
TYPE. Spruce3232, Venezuela, Terr. Amazonas, fl (holotype, K; isotypes, BM,
BR, GH, LE, NY, OXF, P).
DISTRIBUTION.
Periodically flooded forest of the Guianas and Amazonia.
VENEZUELA. Terr. Delta Amacuro: Steyermark87506 fr (F, GH, NY, US); Wurdack306 fr
(NY). Terr. Amazonas: Frdes 21495 fl (F, K, NY, S, US). GUYANA. Jenman 430r fl (K); 4650 fl
Licania 79

(BM, K, NY, U); Rudgesn fr (BM). PERU. Loreto: LI. Williams 937 st (F). BRAZIL. Amazonas:
Byron & Elias 67-59 fl (INPA, NY); DuckeRB 35566 fl (RB); Loureiro& CoelhoINPA 15512 fl (INPA,
NY); Pranceet al 3154 fl (INPA, NY); 3282 fl (INPA, NY); 5013 fl (INPA, NY); Rodrigueset al 5557
fr (INPA 14230, NY, RB); 5941 fl (INPA I4885, NY, RB); 6946 fl (INPA 15499, NY). ParA: Black &
Foster 48-3360 fl (IAN, P); Calvalcante470 fl (INPA, MG 22827); DuckeMG 15967 fl (BM, MG, RB,
US); 16418 fl (3M, MG, RB); RB 18791 fr (RB); Frdes20438 fr (IAN, NY); 33059 fr (IAN); Guedes
238 fl (COL, NY, P); HuberMG 26I7 fl (BM, MG, P, RB, US); Museu Goeldi 10123 fl (BM, INPA,
MG, RB, US); Ledoux287 fl fr (IAN); Oliveira3636 fl (IAN, NY); Pires 1473 fl (IAN, NY); 1486 fl
(IAN, NY, R); 4333 fl (IAN, NY, US); 51867 fl (K, NY, RB, US); Prance,Pennington& Silva 1653 fl
(GH, NY).

LOCAL NAMES. Venezuela: MerecurilloBlanco.Brazil: Caripe,Uchi de Cotia.

40. Licania fuchsii Prance, sp nov Fig 27F-H

Arbor, ramulis juvenilibus puberulis mox glabris, lenticellatis. Folia alternata

petiolata, petiolo I5.0-20.0 mm longo, tereti, pubescente mox glabro, eglanduloso;
laminae oblongae, coriaceae, 15.0-24.0 cm longae, 6.0-io.o cm latae, apice in
acuminem 6.0-I2.0 mm longum contractae, basi rotundatae vel subcuneatae, supra
glabrae, subtus cum cavis stomatalis foveolatae; costa media supra prominente,
glabra; costis secundariis 10-I2 jugis, subtus prominentibus, supra leviter prominenti-
bus. Stipulae caducae (haud visae). Flores circa 4.0 mm longi, haud pedunculati, in
paniculis racemosis terminalibus dispositi, rachi ramisque brunneo-tomentosis.
Bracteae primariae ovatae, ad 15.0 mm longae, subpersistentes,extus puberulae, aliae
bracteae bracteolaeque lanceolatae vel oblongae, parvae, eglandulosae, persistentes.
Receptaculum complanato-cupuliforme, subsessile,extus brunneo-tomentosum, intus
puberulum. Calycis-lobi acuti, extus tomentosi, intus tomentelli. Petala nulla.
Stamina 8, in orbem completum disposita, filamenta glabra, ad basim libera, exserta.
Ovarium ad basim receptaculi insertum, dense lanato-pubescentum. Stylus e basi
ovarii ortus, calycis-lobos excedens, puberulus. Drupa globosa; epicarpio sordido-
pubescentio; pericarpio tenue, fibroso.
TYPE. Fuchs 22036, Colombia, Choc6, Rio Baud6, 13.5 km above mouth, fl fr
(holotype, NY; isotype, US).
DISTRIBUTION. Known only from the type gathering from the Pacific coastal
forest of Colombia.
This species is most closely related to L. longistyla,but differs in the larger in-
florescence with much thicker branches, the larger flowers, the flattened receptacle
with a large prominent disc, the lanate mass surrounding the ovary, the thicker
leaves, and the longer petioles.

41. Licania humilis Chamisso & Schlechtendal, Linnaea 2: 549. I826;

Fritsch, Ann. Naturh. Mus. Wien 4: 59. I889.

Moquileahumilis (Chamisso & Schlechtendal) Hooker f., Mart. Fl. Bras. 14(2): 26. 1867.
Chrysobalanus humilis (Chamisso & Schlechtendal) Kuntze, Rev. Gen. 3(2): 76. I891.
Chrysobalanus sublanatusKuntze, Rev. Gen. 3(2): 76. 1891, synon nov. Type. Kuntze sn, Brazil,
Mato Grosso, fl (NY).
Licaniaulei Taubert, Bot. Jahrb. 21: 428. I896, synon nov. Type. Ule 3187, Brazil, Minas Gerais,
fl (holotype, B, lost; isotypes, HBG, R).

Low tree or shrub to 6.0 m tall, the young stems densely tomentellous, with
thick corky bark. Leaves ovate-elliptic to oblong, thick-coriaceous,4.0-10.5 cm long,
80 Flora Neotropica

2.5-6.5 cm wide, rounded, obtuse, or bluntly acuminate at apex, with acumen

0-8.0 mm long, rounded to subcuneate at base, glabrous and shining on upper sur-
face, stomatal cavities on lower surface, the rounded veins describing open cavities
filled with lanate pubescence; midrib prominent above, lanate when young; primary
veins 7-10 pairs, prominent below; petioles 2.0-6.0 mm long, densely tomentose,
terete, eglandular. Stipules linear, to 4.0 mm long, tomentose, caducous. Inflores-
cences paniculate, the rachis and branches brown-tomentose. Flowers ca 3.0 mm
long, in sessile clusters on primary branches of inflorescence. Bracts and bracteoles
ovate to lanceolate, I.0-2.0 mm long, persistent, entire or with one or two small
teeth on each side. Receptacle campanulate, sessile, tomentose on exterior, villous-
tomentose within. Calyx lobes acute, tomentose on exterior, tomentose to puberulous
within. Petals absent. Stamens 9-I2, inserted in a complete circle; filaments far ex-
ceeding calyx lobes, free to base, glabrous. Ovary inserted at base of receptacle,
villous. Style pilose for most of length, equalling filaments. Fruit ovoid, 2.0-2.5 cm
long, I 3-2.0 cm broad; epicarp smooth, glabrous, drying black; mesocarp thick,
fleshy; endocarp thin, fragile, sparsely hirsute within.
TYPE. Sellow sn, Brazil, Minas Gerais, fl (holotype, LE; isotype, BR; isotype
fragments F, M, P).
DISTRIBUTION. Cerrados of the Planalto of Central Brazil. Flowering March to
September.
BRAZIL. Mato Grosso: Hoehne2186 fl (R); Irwin et al 17127 fl (NY); 17352 fl (NY); Irwin &
Soderstrom6430 fr (NY); Maguire,Pires, Maguire & Silva 56234 fl (NY, UB, US); 56400 fr (NY, US);
Robert497 fr (BM); Schwacke4476 fl (RB); Veloso1201 fl (RB 87400). GoiSs: Duarte 8244 (M 507) fl
(RB, UB); Glaziou21116 fl (K, P, R); Heringer1o466 fl (UB); Irwinet al I3435 fl (NY); 17613 fl (NY);
17864 fl (NY); Lane 8 fl (SP). Dist. Federal: Heringer9272 fr (RB, UB). Minas Gerais: Damazio RB
47985 fl (RB); Lundsn fl (C); Macedo702 fl (MO, S, SP); Magalhaes 17996 fl (NY); 19240 fl (NY);
RegnellIII-152 fl fr (BR, S, US); Riedel482 fl (US); Robert497b fr (K); St. Hilaire 499 fl (P); Santos &
Castellanos24230 fl (RB); Warmingsn fl (NY). Sao Paulo: BarrosRB 47979 fl (RB); Burchell5075 fl (K);
5130 fl (K); Campos30 fl (US); Emmerich2906 fl (R); Handro692 fl (SP); 890 fl (SP); M. Kuhlmann
3455 fl (SP); Lofgren930 fl (SP); Mattos 136 fl (F, RB 10346, NY, UB); RegnellIII-152a fl (BR, S);
Riedel2218 fl (NY, US); Toledo& GehrtSP 4319o fl (SP).

LOCALNAME.Marmelito do campo.

Pilger, in his description of L. ulei, mentioned that it differs from L. humilisin the
verruculose branches, the venation of the upper surface of the lamina, and the
tomentose calyx lobes. In light of the further material studied there is no correlation
or consistency in the above characters, and I have therefore reduced L. ulei to
synonymy. L. humilisis the most distinctive species of this section of Licania.It is a
gnarled shrub or small tree of the cerradowith distinct thick corky bark and cinereous
pubescence on the young branches. It has much more open stomatal cavities than
other related species.

42. Licania foveolata Prance, sp nov Fig 12 F-K.

Arbor ad I2. m alta, ramulis juvenilibus glabris haud lenticellatis. Folia

alternata petiolata; petiolo 4.o-8.o mm longo, sparse pubescente mox glabro,
canaliculato, rugoso, sub laminis cum glandulis duobus sessilibus munito; laminae
obovatae vel oblongo-obovatae, coriaceae, 3.0-11.0 cm longae, 1.7-6.o cm latae,
basi rotundatae vel subcuneatae, apice mucronatae in acuminem 4.0-8.0 mm longum
contractae, supra glabrae, subtus cum cavis stomatalis foveolatae, lanato-pubescentes;
costa media supra leviter prominente; costis secundariis 9- 2 jugis, subtus prominen-
Licania 81

tibus, supra leviter prominentibus. Stipulae lineares, 3.0-4.0 mm longae, mem-

branaceae, subpersistentes. Flores 3.5-4.0 mm longi, haud pedunculati, in paniculis
racemosis terminalibus axillaribusque dispositi, rachi ramisque brunneo-tomentosis.
Bracteae bracteolaeque ovatae, I.0-4.0 mm longae, eglandulosae, extus pubescentes.
Receptaculum campanulatum, sessile, extus brunneo-tomentosum, intus villoso-
tomentosum. Calycis-lobi acuti, utrinque tomentosi. Petala nulla. Stamina Io, in
orbem completum disposita; filamenta glabra, ad basim libera, longe exserta.
Ovarium ad basim receptaculi insertum, villoso-pubescens. Stylus e basi ovarii ortus,
calycis-lobos excedens, subtus villosus, supra glaber. Fructus immaturus globosus,
extus laevis, glaber.
TYPE. Boyan 28 (F.D. 7852), Guyana, Upper Mazaruni River, Aga Creek, fl
(holotype, K; isotypes, NY, U).
Riverine forest, collected only in Guyana.
DISTRIBUTION.
GUYANA. Arubaru River, near Haiamatipu Mt., Pinkus i80 fl (BR, F, GH, LE, MO, NY, S).

This species is closest to L. octandra,but differs in the much larger flowers and the
mucronate leaf apex. The name, L. foveolata,was suggested by the stomatal cavities
of the leaf underside.

43. Licania octandra (Hoffmansegg ex Roemer & Schultes) Kuntze, Rev.

Gen. 217. I89I; Pilger, Notizbl. Bot. Gart. Berlin 8: 539. 1923; Macbride,
Fl. Peru 1072. 1938. Fig I3.
HirtellaoctandraHoffmansegg ex Roemer & Schultes, Linn. Syst. Veg. ed. 9. 5: 274. I819; DC.,
Prodr. 2: 529. I825.
Licania turiuvaChamisso & Schlechtendal, Linnaea 2: 550. 1827, nom illegit.
Moquileaturiuva(Chamisso & Schlechtendal) Hooker f., Mart. Fl. Bras. 14(2): 25. I867; Fritsch,
Ann. Naturh. Mus. Wien 4: 57. 1889, pro parte, syn L. aperta,L. pubifloraexclusum.

Small to medium-sized tree, the young branches glabrous. Leaves broadly ovate
to oblong-lanceolate, 3.0-12.0 cm long, 2.0-4.0 cm broad, obtuse to acuminate at
apex, with acumen 1.0-13.0 mm long, rounded to subcuneate at base, glabrous and
shining above, with well developed stomatal cavities beneath, with venation flattened
around small slit-like apertures to the cavities, the mouth of the cavities glabrous to
lanate; midrib prominulous above, glabrous; primary veins 8-I3 pairs, prominent
on lower surface, plane or nearly so above; petioles tomentose, becoming less so with
age, terete or rarely shallowly canaliculate, with two sessile glands at or near junction
with lower surface of lamina, the glands often obscured by pubescence. Stipules
linear, to 5.0 mm long, membraneous, hirsutulous, subpersistent,on young branches
only. Inflorescencesracemose panicles, the rachis and branches sparsely gray-brown-
tomentose or arachnoid-pubescent. Flowers 2.0-3.0 mm long, sessile or nearly so on
primary branches of inflorescence, solitary or in small groups. Bracts and bracteoles
1.0-4.0 mm long, persistent, tomentose on exterior, often serrate,with stipitate glands.
Calyx lobes acute, tomentose on both surfaces. Receptacle campanulate, sessile or
with short pedicels to 0.2 mm long, tomentose on exterior, densely villous within.
Petals absent. Stamens 8-I2, inserted in a complete circle; filaments far exceeding
calyx lobes, free to base, glabrous. Ovary inserted at base of receptacle, villous. Style
villous at base only, upper portion glabrous, equalling filaments. Fruit globose to
elongate-lanceolate, to 2.5 cm long; epicarp glabrous or with a light brown appressed
pubescence; mesocarp thin and fleshy; endocarp hard, thin, fibrous, glabrous or
sparsely hirsute when young within.
82 Flora Neotropica

''''~ ~~~~~~~~~~~~~~~~~~~~'
AA

?~ ~~~~~~~~~~~~~~~~~~~??

20 200 s n

1 ~~~~~~~~~f'
.. I . _/~
C Da
Licania 83

Key to Subspecies of Licaniaoctandra

i. Leaves broadly ovate to oblong, obtuse to bluntly acuminate at apex, with acumen i.0-5.0
mm long; upper surface of leaf drying brown; young inflorescence with sparse gray-brown-
tomentum. a. subsp octandra.
I. Leaves oblong-lanceolate, with a well developed finely pointed acumen 5.0-I3.0 mm long;
upper surface of leaf drying gray; young inflorescence usually with rufous-brown arachnoid
pubescence. b. subsp pallida.

43a. Licania octandra subsp octandra

Licania octandra(Hoffmansegg ex Roemer & Schultes) Kuntze, Rev. Gen 217. 1891.
Licania bothynophylla Martius, Flora 24(Beibl. 2): 15. I841. Type. Martius 449, Brazil, Rio de
Janeiro, fl (holotype, M; isotypes, BR, F, GH, K, LE, M, NY, P).
Moquileabothynophylla (Martius) Hooker f., Mart. Fl. Bras. 14(2): 26. 1867.
Moquileautilis Hooker f., Mart. Fl. Bras. 14(2): 24, t. 7. I867. Types. Spruce133, Brazil, Para, st
(paratype, K); 877 fl (lectotype, K; isolectotypes, LD, NY, P); Sagot969, French Guiana,
fl (paratypes, BM, GH, K, NY, P, S, US).
Licaniautilis (Hooker f.) Fritsch, Ann. Naturh. Mus. Wien 4: 56. 1889.
LicaniasellowianaKlotzsch ex Hooker f., Mart. Fl. Bras. I4(2): 26. 1867. nom nud, in syn.
Licania takutuensisStandley, Lloydia 2: 182. I939, synon nov. Type. A. C. Smith3302, Guyana,
fl (holotype, F; isotypes, A, IAN, K, LE, NY, P, S, US).
Licania hookerivar obtusaHuber, Bol. Mus. Emilio Goeldi 5: 368. 1909, synon nov. Type. Ducke
MG 8371, Brazil, Para, fl (holotype, MG; isotypes, BM, G).

Leaves broadly ovate to oblong, obtuse to bluntly acuminate at apex, with

acumen I.0-5.0 mm long; upper surface of leaf drying brown; young inflorescence
with sparse gray brown tomentum.
TYPE. Siebersn, Brazil, Para, fl (holotype, B, Herb. Willdenow 4851; photo, NY).
Forest, both on flooded and non-flooded ground but especially in
DISTRIBUTION.
open riverine habitats and at savanna margins, widely distributed from northern
Venezuela through the Guianas and easern Amazonia to northeast and eastern-
central Brazil, and in the gallery forest of the Planalto of central Brazil.
Representative collections: VENEZUELA. Monagas: Steyermark61759 fr (F, NY, US, VEN).
Barinas: Aristeguieta1653 fl (NY, VEN); Bernardi1119 fl (NY, VEN); Breteler3927 fr (VEN). M6rida:
Bernardi2346 fl (NY, VEN); Grunwaldsn fl (VEN); Lasser2368 fl (NY). Bolivar: Steyermark 86865 st
(VEN). GUYANA. De La Cruz3499 fl (NY, US); Fanshawe2587 (F.D. 5375) fl (A, K, NY, S, U, US);
Im Thurnsn fl (K); Irwin 238 fl (US); Jenman42 fl (P); 4865 fl (K); 6630 fl (K, NY); A. C. Smith3107 fl
(A, F, IAN, K, LE, MO, NY, P, S, U, US). SURINAME. Lindeman6230 st (NY, U); 6371 st (U);
Schulz 7630 st (U); Stahel 6ia st (IAN, NY); VanDonselaar2946 st (U). FRENCH GUIANA. Melinon
sn (I863) fl (A, F, K). PERU. Loreto: Tessmann3446 fl (NY, S). BRAZIL. Terr. Roraima: Pranceet al
3967 fl (INPA, NY); 4196 fl (INPA, NY); 4235 fl (INPA, NY); 4520 fl (INPA, NY); 4576 fl (INPA,
NY). Para: Burchell9316 fl (K, NY, P, US); 9629 fr (P); Black 47-996 fl (IAN, INPA); DuckeMG 8371
fl (MG); RB 15192 fl (K); RB I8795 fl (K, RB); RB i8796fl (K, RB); RB 18824 fl (K, RB); RB 19798
fl (K, MG 16153, RB); RB 19800 fl (K, MG I6260, RB); Egler 842 fl (IAN, NY); Froes30379 fr (IAN,
UB); Silva 554 fl (IAN); Pires, Black & Dobzhansky4132 fl (IAN). Terr. Amapa: Pires & Silva 4812 fr
(IAN, NY, P). Maranhao: Duckesn (Registro16) fl (IAN, NY); Frdes28524 fl (IAN). Ceara: Castellanos
& Duarte523 fl (HB); Guedes628 fl (IAN, MG, NY, R). Paraiba: Lima 65-4340 fr (IPA); Moraes894 fl
&
(NY, RB 86440). Pernambuco: Leal & Silva 154 fl (IPA, RB); Ducke& Lima33 fl (R). Goias: Prance
Silva 58524 fl (NY, RB, UB, US); Glaziou 21115 fl (BR, K, P). Dist. Federal: Irwin & Soderstrom 5257
fl (NY, UB); Pires, Silva & Souza 9546 fl (NY, UB); 9640 fl (NY, UB). Bahia: Blanchet216 fl (P);
Lima 61-3919 fl (IPA). Minas Gerais: Duarte2967 fl (NY, RB 71913); Glaziou 1478 fl (BR, K, NY, P,

FIG I3. Leaf shape variation and geographical distribution of Licania octandra.A, A, B, E,
L. octandrasubsp octandra;A, Fanshawe2587; B, Pranceet al 3967; E, Mikan sn. 0, C, D, L. octandrasubsp
pallida; C, Krukoff4688; D, Cordeiroet al 145; leaves, x 0.25.
84 Flora Neotropica

US); Irwin et al 7821 fr (NY); 9618 fr (NY); St. Hilaire 2062 fl (P). Rio de Janeiro: Bello 6178 fl (R).
Guanabara: Duarte5831 fl (NY, RB 124045); Glaziou 1385 fl (BR, P); Kuhlmann156I fl (RB); Langs-
dorffsnfl (US); Pessoalde HortoFlorestalRB 136939 fl (RB); Riedel sn fl (NY, US); Schwackesn fl (R);
VitorioRB 136938 fr (RB). Without locality, Rio de Janeiro or Guanabara: Gaudichaud113 fr (P);
Luschnathsn fl (OXF); Mikan sn fl (NY, W).

LOCAL NAMESAND USES. Venezuela: Clavellino, sorore,Palo de hierro, Guaray blanco.

Suriname: Kauta(Arawak), Kwepi.Brazil: Caripe,Cocao.An extract of the bark is used
by several tribes of Indians for hardening pottery.

43b. Licania octandra subsp pallida (Hooker f.) Prance, stat nov
Moquileapallida Hooker f., Mart. Fl. Bras. 14(2): 25. 1867.
Licaniapallida (Hooker f.) Kuntze, Rev. Gen. 217. 891 nom illegit.
LicaniahookeriFritsch, Ann. Naturh. Mus. Wien 4: 56. 1889, synon nov. Type. Spruce3302.
Licania egensisFritsch, Ann. Naturh. Mus. Wien 4: 58. 1889, synon nov. Types. Poeppig2531,
Brazil, Amazonas, fl fr (lectotype, W; isotypes, G, LE); 2770 fl (paratype, W).
LicaniastenocarpaStandley, Publ. Field Mus. Bot. 17: 258. I937, synon nov. Type Krukoff8553,
Brazil, Amazonas, fr (holotype, NY; isotypes, A, BM, BR, F, MO, P, S, US).

Leaves oblong-lanceolate, with a well developed finely pointed acumen 5.0-

13.0 mm long; upper surface of leaf drying gray; young inflorescence usually with
rufous-brown arachnoid pubescence.
TYPE.Spruce3302, Venezuela, Terr. Amazonas, fl (holotype, K; isotypes, BM,
BR, CGE, GH, GOET, LD, LE, P, OXF, RB).
Non-flooded forest in western and central Amazonia.
DISTRIBUTION.
COLOMBIA. Caqueta: RomeroC. 3964 fl (COL). PERU. Loreto: LI. Williams 133r fl (F).
BRAZIL. Amazonas: Aluisio i85 fr (INPA); Black 48-2471 st (IAN, NY); CoelhoINPA 5178 st (INPA);
Ducke MG 7623 fl (BM); RB 25028 fl (RB); Froes 23252 fl (IAN, NY); GouveaINPA o066fl (IAN,
INPA, NY); Krukoff4688 fl (A, F, K, LE, MICH, MO, NY, S, US, U); 4915 fl (A, F, LE, MICH,
MO, NY, RB, S, U); 8560 fr (A, BR, F, K, LA, LE, MO, NY, S, U); Loureiro& CoelhoINPA 15514 fl
(INPA); I5518 fl (INPA); OsmarinoINPA 20738 fr (INPA, NY); Pranceet al 2504 fr (INPA, NY);
Rodrigueset al 1423 fl (INPA 7754); i808 fr (INPA 8179); I884 fr (INPA 8256); 2952 fl (INPA 14896,
NY); 5819 fl (INPA I4771, NY); 5952 fl (INPA 14896); 5974 fl (INPA 14918, NY, RB); 6886 fl
(INPA 15436, NY); 6951 fl (INPA 15525, NY); 6969 fl (INPA 15542, NY); Schultes24559 fl (INPA).
Terr. Roraima: Pranceet al 9537 fl (INPA, NY). Para: Black, Egleret al 19980 fl (IAN, NY); MG 16223
fl (MG); RB 18755 fl (K). Acre: Krukoff5629 fl (A, F, LE, MO, NY, US); Pranceet al 7785 fr (INPA,
NY). Terr. Rond6nia: Cordeiro& Silva 145 fl (IAN, NY); Pranceet al 5242 fl (INPA, NY); 5579 fl
(INPA, NY); 5678 fl (INPA, NY); 6512 fl (INPA, NY). BOLIVIA. Pando: Pranceet al 6130 fl (INPA,
NY).

LOCAL NAME. Caripe.

The two subspecies of L. octandraare markedly distinct from one another at the
extremes of their variation. However, as certain intermediates occur it is not always
easy to distinguish between them. Since the variation is well correlated geographi-
cally and the two subspecies are allopatric (see Fig I3), it seems best to regard these
variants as two subspecies. Two collections, Maguireet al 55681 fr (NY), and Cordeiro
& Silva309 fl (IAN, NY), are intermediate between the subspecies.
Licaniaoctandrais most closely related to L. sprucei,but differs by its sessile flowers.
The taxonomic problems within this species are very similar to those already dis-
cussed under L. apetala.Licaniaoctandrais a species that needs further experimental
work, and falls into the category of an ochlospecies.
Confusion between L. octandraand L. apetalabegan with Hooker's account of the
genus in Flora Brasiliensis.He treated L. octandraunder the illegitimate name of
Licania 85

L. turiuva,the latter having been created by Chamisso and Schlechtendal to accommo-

date Hirtellaoctandra.Hooker cited L. apertaand L. pubifioraas synonyms of L. turiuva.
However, these two names are in fact synonyms of L. apetala.Fritsch (i 889) followed
Hooker's circumscription. The type of L. octandra(the same as that of L. turiuva)has
stomatal cavities on the leaf undersurface, but those of L. apertaand L. pubiflorado
not. Pilger (I923) the first to realize the confusion, studied the type of L. octandraand
matched it with two of the type specimens of L. utilis rather than with L. apertaand
L. pubiflora.Pilger also suggested that L. egensiswas the same as L. octandra.I have not
seen the Berlin type specimen (in the Willdenow herbarium) of L. octandra,but
photographs of it show the stomatal cavities as described by Pilger, who did see the
actual type. A study of the material now available shows that L. bothynophylla, L.
takutuensis,and L. stenocarpaare all synonymous with L. octandra.Apart from the
charactersused in support of the two subspeciesdifferentiatedhere, the other variable
features are not well correlated. Scatter diagrams did not indicate any other in-
fraspecific taxa. Subspecies octandraitself is very variable in its morphology.

3. Licania subgen Moquilea sect Microdesmia Bentham

This group contains species which are related to subgenus Moquilea,but differ
by having short stamens. It must be admitted that this section is perhaps the least
natural of those recognized here. The component species are for the most part clearly
distinct from one another. The geographical distribution of this section is unusual in
that the component species occur in coastal Colombia and in northeast and south
Brazil, peripheral to the center of distribution of the genus. This section, the species
of which have short but numerous stamens, shows that the staminal characters used
in the past to separate the general Licaniaand Moquileaare not reliable since all
species in this section are intermediate in these characters.

44. Licania rigida Bentham, Jour. Bot. Hooker a: 220. 1840; Hook. f., Mart.
Fl. Bras. 14(2): I3. I867; Fritsch, Ann. Naturh. Mus. Wien 4: 45. I889.
PleraginaumbrosissimaArruda da Camara ex Koster, Trav. Bras. 500. I816 nom. illegit; see
Prance. Mem. N.Y. Bot. Gard. 15(1): '34. 1966.

Small tree to 15.0 m tall, with spreading crown, the young branches lanate to
tomentellous, soon becoming glabrous and lenticellate. Leaves oblong to elliptic,
6.o-I3.0(-I6.0) cm long, 2.8-6.5 cm broad, coriaceous, rounded to retuse at apex,
rounded to cordate at base, glabrous and shining on upper surface, the lower surface
with deeply reticulate venation quite or nearly describing stomatal cavities, with
lanate pubescence among but not on venation; midrib prominulous above, puberu-
lous towards base when young; primary veins II-I6 pairs, prominent on lower
surface, prominulous above; petioles 5.o-8.o cm long, tomentose when young, be-
coming glabrescent with age, terete, with two sessile glands. Stipules linear, to o0.0
mm long, membraneous, caducous. Inflorescences racemose panicles, the rachis and
branches gray-tomentose. Flowers 2.5-3.5 mm long, in small groups, sessile on
primary branches of inflorescence. Bracts and bracteoles 1.5-2.5 mm long, ovate,
tomentose on exterior, persistent, entire to serrulate, eglandular. Receptacle cam-
panulate, gray-tomentose on exterior, tomentose within; pedicels to 0.5 mm long.
Calyx lobes acute, tomentose on exterior, tomentellous within. Petals 5, densely
86 Flora Neotropica

pubescent. Stamens ca 14; filaments equalling calyx lobes, connate to about halfway
from base, densely pubescent. Ovary attached to base of receptacle, villous. Style
equalling calyx lobes, villous nearly to apex. Fruit elliptic, 4.0-5.5 cm long; epicarp
smooth, drying green or black; mesocarp thin, fleshy; endocarp thin, fibrous, fragile,
fibers arranged longitudinally promoting longitudinal dehiscence, sparsely pubescent
within.
TYPE.Gardner1592, Brazil, Ceara, fl (holotype, K; isotypes, BM, F, G, GH, NY
OXF, P, US).
Dry forest and gallery forest of northeastern Brazil.
DISTRIBUTION.
BRAZIL. Piaui: Alencari6 fl (RB I36927); Frdes 11687 st (NY). Cear,: Allemao& Cysneiros560
fl (R); Agr. Chef.RB 78265 fl (NY); Cutler8264 fr (A, ECON, F, GH, NY, US); 8375 fl (A, ECON, F,
GH, US); Dahlgren932 fr (A, F, P); DuckeMG 1102 fr (MG, RB); MG 1603 fl (BM, MG, RB); Lisboa
MG 2438 fr (MG, RB); O. Martins MG 902 fr (BM, MG, RB); M. Silva 343 st (MG); Vice-Consul
sn st (K); Virgilissn fl (A, F, GH, MICH, MO, NY, P, S, US). Rio Grande do Norte: Castellanos22867 fl
(R); 23005 fr (R); Sobrinho80 fl (HST); Tavares683 fl (HST). Paraiba: Luetzelburg12531 fl (A, F, M,
US); 26616 fl (M); 26619 fl fr (M); 26642 fl (M, NY, US); 28105 fl (M); 28106 fl (M); 28107 fl (M);
28108 fl (M); 28109 fl (M); 28112 fl (M); 28113 fl (M); Moraes892 fl (NY, P); Pickel3947 st (IPA);
Sobrinhosn fl fr (IPA). Guanabara: Glaziou 10700 fl (K, P); KuhlmannRB 41453 cult, fl (NY, RB). Sao
Paulo: Santoro& Siviero10032 cult fl (SP); ToledoSP 8850 cult fl (SP).

LOCAL NAMEAND USES. Oiticica. Licania rigida is grown for its seeds from which oil
is extracted. The oil is used as a substitute for tung oil in paint (Bull. Imp. Inst. 44:
I5. I946). Brazil exported 16,606 metric tons in 1941. The kernels contain 61% oil,
which is extracted with petrol ether (see also Tropenflanzer 206-2O1. 1931). The
wood is also used for building purposes. This species is cultivated outside its natural
range, eg in Trinidad, (Prance2113A fl, NY), but is not used commercially outside
Brazil.

45. Licania arborea Seeman, Bot. Voy. Herald 118, t. 25. 1853; Standl.,
Contr. U.S. Natl. Herb. 23: 343. I922.
LicaniaselerianaLoesner, Verh. Bot. Ver. Brand. 53: 55. 1911 . Types. Tonduz 13808, Costa Rica,
fl (paratypes, GH, K, NY); 13890 fr (paratypes, BM, GH, K); Seler i660, Mexico,
Oaxaca, fl (isotypes, GH, US); 1992 (paratype, not seen).
Licania retusaPilger, Notizbl. Bot. Gart. Berlin 6: 137. 1914; Macbride, Fl. Peru 1072. I938.
Type. Ule 9598, Brazil, Acre, fl (holotype, B, lost; isotypes, K, L, U).
Licania bullatifoliaCuatrecasas, Fieldiana Bot. 27: 63. I950. Type. Cuatrecasaso1865, Colombia,
Putumayo, fl (holotype, F; isotypes, COL, NY, P, US).

Tree to 20.0 m tall, the young branches pubescent, soon becoming glabrous.
Leaves ovate-orbicular to oblong, 5.0-12.0 cm long, 2.5-8.0 cm broad on fertile
branches (usually much larger on sterile branches), glabrous and shining above,
lower surface with deep reticulation quite or nearly describing stomatal cavities,
lanate-pubescent among venation, sometimes with dense lanate pubescence obscur-
ing venation when young, obtuse, rounded, or retuse at apex, cordate to rounded at
base; midrib prominulous above, glabrous; primary veins II-I8 pairs, prominent
beneath, slightly impressed above; petioles 5.0-i2.0 mm long, terete, tomentose
when young, glabrescent, with two sessile glands near lower surface of lamina.
Stipules to 3.0 mm long, linear, intrapetiolar, membraneous, caducous. Inflores-
cences terminal and axillary racemose panicles, the rachis and branches densely
brown to ferrugineous-tomentose. Flowers 2.5-3.0 mm long, solitary and densely
clustered on primary and secondary branches of inflorescence. Bracts and bracteoles
deltoid to lanceolate, to 1.5 mm long, persistent, tomentose on exterior, puberulous
within, entire to serrulate, eglandular. Receptacle campanulate, sessile, tomentose
Licania 87

on exterior, villous within. Calyx lobes acute, tomentose on both surfaces. Petals 5,
small, oblong, pubescent. Stamens 8-I2, inserted in a complete circle; filaments
equalling calyx lobes, connate in groups for half their length, densely villous. Ovary
inserted at base of receptacle, densely pilose. Style equalling filaments, pilose through-
out. Fruit oblong, to 3.0 cm long; epicarp smooth and glabrous, drying black or
green; mesocarp thin and fleshy; endocarp thin, soft and fibrous, fibers arranged
longitudinally, glabrous within.
TYPE. Cuming I og, Panama, fl (BM, K, MO).
DISTRIBUTION.Dry plains, slopes, and scrub forest, through Central America
from Mexico to Panama, and infrequent in South America in Colombia and Peru.
MEXICO. Michoacan: Leavenworth& Hoogstraal452 st (F); 1448 st (GH); 1612 st (F, MO);
Maury 5313 fl (NY). Mexico: Hinton3021 fl (BM, K, S); 3832 fr (BM, F, K, NY); Matuda 30680 st
(MEXU); Maury 4698 st (NY); Paray3365 fl (MEXU). Morelos: Gdmez-Pompa1216 fl (GH, MEXU).
Guerrero: Hinton5357 fr (A); 5957 fr (K); IooI5 fr (GH, K, NY); Miranda 1915 fr (MEXU); Niiuez
sn fl (MEXU); Palmer342 fl (A, F, K, MEXU, MO, NY); Pennington& Sarukhdn 9477 fr (NY). Chiapas:
Becerra9 fl (MEXU); Miranda4955 fl (MEXU); Purpus9183 fr (F, GH, MEXU, MO, NY). Oaxaca:
Conzatti4455 fl (GH); Juzepczuk 1 66 fl (F, LE); Nelson 2363 fr (GH); Pennington& Sarukhdn9337 st
(FHO, MEXU); Ton3759 fr (NY). GUATEMALA.Aguilar1717 fl (F); Kellerman 7063 fl (F); Pittier
1899 fl (NY); Standley 60138st (A, NY); 75388 st (F); 76037 st (F); 89271fr (F); Steyermark 30236 st
(F); 3I399 st (F); 47761st (A). HONDURAS. MolinaR. 853 st (F, GH); 11899fr (F, NY); Standley 91
st (F); 24558 st. (F): Standley
et al 1276 st (F); 1290 st (F); 5847 st (F); 7338 st (F); L. 0. Williams
i8i60 fr (GH); L. O. Williams & Molina R. 12210 st (F, GH, MICH); 12563 st (F, GH); 12830 fl
(F, GH); i8005 fr (F, GH). EL SALVADOR.Calderon 2030 fl (NY); Standley20035st (NY); 20327fl
(GH, NY); 20429 fl (GH, NY); 21206 st (GH, NY); Standley& Padilla V. 3143 st (F); 3401 fl (F);
3564 st (F); Tucker9I5 fr (F, K, MICH, NY, P). NICARAGUA. Bunting& Licht717 fr (NY); Standley
9170 st (F). COSTA RICA. Burger& RamirezR. 4066 fl (F); Jimenez3101 fr (F, NY); Ledn988 st (F);
Standley & Valerio44901 st (A); Valerio492 fr (F). PANAMA. Allen 1741 fl (F, GH, MO, NY);
Piper5713 st (NY); Pittier 2841 fl (NY); Seeman508 fl (GH, K, S); sn fl (GOET); Stern& Chambers135
st (F, MO, NY); R. S. Williams338 fl (NY). COLOMBIA. Magdalena: Dugand & Peten369:816 st
(F); Dugand 6311 st (COL); 6314 st (US). Bolivar: Curran303 st (GH, US). PERU. Amazonas:
2504 st (U).
Ellenburg

LOCAL NAMESAND USES. Mexico: Cacahuananche, Cacahuate, Cacahoanantzin,

Quirindolcacahuananche, Cana dulce,Frailecillo,Palo de fraile, Totopostle.Guatemala:
Encina.El Salvador: Roble,Canillamula.Costa Rica: Alcornoque. Panama: Rasca.This
species has durable wood which is used in general construction. The fruit, rich in an
inflammable oil, is often strung on sticksfor illuminating purposes. The oil is extracted
in large quantities for use in candles, soap, axle grease etc (see Standl., Contr. U. S.
Natl. Herb, 23: 343. 1922.)
Licaniaselerianawas described by Loesner because it differed from L. arboreain
the less pubescent leaf underside and the caducous petals. L. selerianawas later reduced
to synonymy by Standley (1922), although he admitted that he did not see authentic
material of it. It was restored to specific rank by Cuatrecasas (1950), but no detailed
reasons were given for regarding L. selerianaas a distinct species. On the basis of the
generous material examined in this study, I find that there is a complete range in the
pubescence of the lamina lower surface and in the time of dehiscence of the petals
between L. arboreaand L. seleriana,and that these two characters are not correlated
with one another nor with the additional small differences between the type speci-
mens of the two species. Consequently I treat L. selerianaas a synonym of L. arborea.
Furthermore, I have examined carefully the types of two South American names,
L. bullatifoliaand L. retusa,and can find no consistent differences to separate them
from L. arborea.In the original description L. bullatifoliawas distinguished by the
bullate leaves with a much deeper basal sinus. Several of the Central American
collections of L. arboreashow a tendency to bullate leaves and an equally deep basal
sinus (eg Allen 174I from Panama). The flowers of this species are interesting because
88 Flora Neotropica

of the filaments which are connate in groups and densely pubescent. The species most
closely related to L. arboreais L. rigidafrom the northeast of Brazil, differing in the
pubescence of the inflorescence and leaves, the slightly impressed leaf venation, and
the shorter rounder fruit. The flowers and fruit of these two species are very similar.
The fruit is rich in oil, and is used commercially in both regions. Perhaps the rather
unusual distribution of L. arboreaderives from its cultivation for economic purposes.

46. Licania velata Cuatrecasas, Fieldiana Bot. 27: 60. 1950.

Large tree to 35.0 m tall, the young branches tomentose, soon becoming glabrous.
Leaves oblong to oblong-elliptic, I5.0-36.0 cm long, 6.o-i I.o cm broad, cuspidate-
acuminate at apex, with acumen 5.0-1 I.0 mm long, subcuneate at base, caducous-
lanate on both surfaceswhen young, becoming glabrous with age; midrib prominent
and glabrous above when mature; primary veins II-I2 pairs, prominent on lower
surface, prominulous above; petioles 8.o-i8.o mm long, arachnoid when young,
becoming puberulous, terete, eglandular. Stipules i5.o-20.0mm long, linear, mem-
braneous, intrapetiolar, caducous, adnate to base of petiole. Inflorescences axillary
panicles, on nodes of mature woody branches, the rachis and branches arachnoid.
Flowers ca 2.5 mm long, in small sessile clusters on primary and secondary branches
of inflorescence.Bracts and bracteoles ovate, equalling flowers, eglandular, pubescent
on exterior, entire. Receptacle campanulate-urceolate, sessile, arachnoid on exterior,
villous-tomentose within. Calyx lobes acute, lanate on exterior, puberulous within.
Petals 5, puberulous. Stamens I i-I5, inserted in a complete circle; filaments equall-
ing the calyx lobes, united at base only, densely villous. Ovary inserted at base of
receptacle, villous-lanate. Style densely villous throughout, equalling filaments.
Fruit unknown.
TYPES.Cuatrecasas21179, Colombia, Valle, fl (lectotype, F; isolectotypes, COL,
MAD, P); 21179A fl (paratypes, COL, F, P).
DISTRIBUTION.Forest in the Pacific coastal regions of Colombia. Known only
from the type gatherings.
This species has a ramiflorous inflorescence, a character that is shared with only
one other species of Licania,L. arachnoidea,
which is not closely related.

47. Licania subarachnophylla Cuatrecasas, Fieldiana Bot. 27(2): I I0. 1951.

Tree, the young branches tomentose, soon becoming glabrous. Leaves elliptic to
oblong-elliptic, coriaceous, 4.5-7.0 cm long, 2.0-3.6 cm broad, rounded or short-
mucronate at apex, subcuneate at base, glabrous above, caducous-lanate beneath;
palisade glands absent; midrib prominent above, tomentellous; primary veins I I-12
pairs, prominent on lower surface; petioles 2.0-4.0 mm long, tomentellous, terete,
eglandular, rugose. Stipules 3.0-6.0 mm long, linear, membraneous, intrapetiolar,
caducous. Inflorescences axillary and terminal unbranched spikes, the rachis arach-
noid. Flowers ca 1.5 mm long, in glomerules along rachis. Bracts and bracteoles
1.5-4.0 mm long, oblong-ovate to lanceolate, persistent, tomentose on exterior,
glabrous within, entire. Receptacle cupuliform, sessile, lanate on exterior, lanate
within. Calyx lobes acute, lanate on exterior, glabrous within. Petals 5, ovate, ciliate.
Stamens ca 15, inserted in a complete circle; filaments equalling or slightly exceeding
the calyx lobes, slightly connate at base. Ovary inserted at base of receptacle, densely
lanate. Style lanate. Fruit unknown.
Licania 89

TYPE. Haught 2629, Colombia, Boyaca fl (holotype, F; isotypes, A, COL, NY,

US).
Known only from the forest of Central Colombia.
DISTRIBUTION.
& Jaramillo17I45 fl (COL).
COLOMBIA.Meta: Garcia-Barriga

This species is distinguished by the spiciform inflorescence.

48. Licania salicifolia Cuatrecasas, Fieldiana Bot. 27(2): I I. 1951.

Small tree, the young branches puberulous, soon becoming glabrous, con-
spicuously lenticellate. Leaves oblong-lanceolate, coriaceous, 4.5-9.5 cm broad,
acuminate at apex, with acumen 3.0-6.0 mm long, cuneate at base, glabrous above,
densely gray-lanate-arachnoid beneath; midrib glabrous and prominulous above;
primary veins IO-I4 pairs, prominent beneath, slightly impressed above; petioles
4.0-5.0 mm long, lanate, becoming puberulous, terete, eglandular, transversely
rugose. Stipules linear, membraneous, pubescent, 4.0-8.0 mm long, caducous.
Inflorescences racemose panicles, gray-tomentose. Flowers 1.5-2.0 mm long, sessile
on primary branches of inflorescence.Bracts and bracteoles ovate to linear, to 2.5 mm
long, persistent, tomentose on exterior, glabrouswithin, entire, eglandular. Receptacle
cupuliform-globose, sessile, densely ferrugineous-tomentose on exterior, tomentose
within. Calyx lobes acute, tomentose on both surfaces. Petals 1.5 mm long, ovate,
ciliate. Stamens ca 25, inserted in a complete circle; filaments apparently shorter than
calyx lobes (seen only in unopened buds). Ovary inserted at base of receptacle,
villous-tomentose. Style lanate throughout. Fruit unknown.
TYPE. Daniel 477, Colombia, Antioquia, fl (holotype, US; isotypes, COL, F,
GH, NY).
DISTRIBUTION. Known only from the type, collected beside the Rio Negro of
Colombia.

49. Licania araneosa Taubert, Bot. Jahrb. 21: 428, 1896.

Tree, the young branches tomentellous, becoming glabrous with age, not lenticel-
late. Leaves oblong-lanceolate, coriaceous, 2.0-7.0 cm long, 1.3-3.0 cm broad, acute
or short-mucronate at apex, with acumen not exceeding I.o mm long, cuneate at
base, glabrous above, with easily rubbed off lanate pubescence beneath; midrib
prominulous above, lanate; primary veins 9-I pairs, prominulous beneath, plane
above; petioles 2.0-3.0 mm long, lanate, eglandular, terete. Stipules ca I.o mm long,
linear, coriaceous, persistent, extrapetiolar. Inflorescencesterminal and axillary little
branched panicles, the rachis and branches tomentellous. Flowers 1.5 mm long,
sessile on primary and secondary branches of inflorescence. Bracts and bracteoles
0.5-2.5 mm long, ovate to liner, pubescent on exterior. Receptacle cupuliform, sessile,
tomentose on exterior, densely tomentose within. Calyx lobes acute, tomentose on
exterior, glabrous within. Petals 5, sparsely pubescent, ciliate. Stamens I5-I7,
inserted in a complete circle; filaments equalling calyx lobes, free to base, glabrous.
Ovary inserted at base of receptacle, pilose. Fruit (immature) rotund; epicarp
glabrous, verrucose; mesocarp thin, fleshy; endocarp thin, granular, fragile.
TYPE. Ule 2885, Brazil, Goias (holotype, B, lost; lectotype, HBG; isotype, R;
fragment F).
90 Flora Neotropica

DISTRIBUTION.Cerradoand gallery forest of the Planalto of Central Brazil,

of
apparently very restricted range. Flowering in January.
BRAZIL. GoiAs: Ule 459 fl (P), (probably same as type gathering); Irwinet al 11833 fl (topotype,
NY). Dist. Federal: Irwin et al 18229 fr (NY).

50. Licania silvatica Glaziou ex Prance Fig 12 E.

L. silvaticaGlaziou, Bull. Soc. Bot. France 53(Mem. 3): 195. I906 nom.nud.

Frutex vel arbor?, ramulis juvenilibus tomentosis mox glabris. Folia alternata
petiolata; petiolo 6.0-9.o mm longo, tomentoso, eglanduloso, canaliculato, sub
lamina subtus cum glandulis duobus munito; laminae oblongae vel oblongo-lanceo-
latae, coriaceae, 6.o-Io.o cm longae, 2.2-3.5 cm latae, basi rotundatae, apice in
acuminem 5.0-8.0 mm longum contractae, utrinque glabrae, subtus sparse glandu-
losae; costa media supra glabra, plana; costis secundariis I I-I4 jugis, subtus leviter
prominentibus supra leviter impressis. Stipulae lineares, 8.0-I2.0 mm longae,
membranaceae, intrapetiolares, subpersistentes. Flores circa 2.0 mm longi, haud
pedunculati, in paniculis terminalibus dispositi, rachi ramisque griseo-brunneis
tomentosis. Bracteae bracteolaeque parvae, ovato-lanceolatae, persistentes, extus
tomentellae. Receptaculum cupuliforme, sessile, extus tomentosum, intus villoso-
tomentosum. Calycis-lobi acuti, utrinque tomentosi. Petala 5, puberula. Stamina
circa 25, in orbem completum disposita; filamenta glabra, versus basim breviter
connata, calycis-lobos aequantia (ut videtur in alabastro). Ovarium ad basim recep-
taculi insertum, villoso-pubescentum. Stylus e basi ovarii ortus, calycis-lobosaequans,
villosus. Fructus mihi ignoti.
TYPE. Glaziou10702, Brazil, Espirito Santo, fl (holotype, C; isotypes, BR, K, P).
DISTRIBUTION. Known only from the type gathering collected in cerrado of
Espirito Santo in Brazil.
LOCAL NAME. Oiti silvestre.
This species, never described by Glaziou, is in fact most distinct. It is closest to
L. araneosabut differs by its larger glabrous leaves with more numerous primary
veins, and in the larger number of stamens.

5I. Licania chocoensis Cuatrecasas, Fieldiana Bot. 27: 65. 1950.

Tree, the young branches glabrous. Leaves ovate-elliptic to oblong-elliptic,

coriaceous, 8.0-I4.0 cm long, 3.5-5.5 cm broad, cuspidate-acuminate at apex, with
acumen I2.0-I5.0 mm long, rounded to subcuneate at base, glabrous on both surfaces
palisade glands absent; midrib prominent above, puberulous towards base; primary
veins 9-12 pairs, prominent beneath, scarcely visible above; petioles 8.0-1 I.0 mm
long, puberulous, becoming glabrous with age, terete, eglandular. Stipules caducous
(not seen). Inflorescences terminal panicles, the rachis and branches short-tomen-
tellous. Flowers 2.0-2.5 mm long, in small groups or solitary on short secondary
branches (peduncles) of inflorescence. Bracts and bracteoles caducous. Receptacle
campanulate-cupuliform, short-tomentellous on exterior, densely tomentose within;
pedicels 0.5-I.0 mm long. Calyx lobes acute, tomentellous on both surfaces. Petals 5,
puberulous on exterior, white. Stamens ca 14, inserted in a complete circle, filaments
equalling calyx lobes, connate at base, villous at base only. Ovary inserted at base of
receptacle, lanate. Style villous on lower portion. Fruits ovate-elliptic, 4.0 cm long;
epicarp verrucose; mesocarp thin, fleshy; endocarp, thin, fibrous, glabrous within.
Licania 91

TYPES.Cuatrecaasa 21455, Colombia, Choc6, fl (holotype, F; isotype, P); 17752,

Colombia, Valle, fl (paratypes, COL, F).
Confined to the Pacific coastal forests of Colombia.
DISTRIBUTION.
15941fr (COL, F).
COLOMBIA.Valle: Cuatrecasas

4. Licania subgen Parinariopsis

52. Licania licaniaeflora (Sagot) Blake, Contr. Gray Herb. 42: 66. I9I7.
MoquilealicaniaefloraSagot. Ann. Sci. Nat. Bot. VI. 15: 308. 1883.
LicaniabracteosaFritsch, Ann. Naturh. Mus. Wien 4: 54. 1889, nom. illegit.
LicaniaparinarioidesHuber, Bol. Mus. Emilio Goeldi 5: 368. I909; Macbr. Fl. Peru 1073. I938,
synon nov. Type. DuckeMG 896r, Brazil, Para, fl (holotype, MG; isotypes F, RB I5180)
Licaniaparinarioidesvar latifolia Maguire, Brittonia 7: 397. 1952. Type. DuckeMG i6299, Brazil,
Para, fl (holotype, US; isotypes, BM, MG, RB).
LicaniacapinensisHuber, Bol. Mus. Emilio Goeldi 6:7I. I9I0, synon nov. Type. HuberMG 946,
Brazil, Para, fl (holotype, MG; isotype, BM).
LicaniahuberianaMaguire, Brittonia 7: 398. 1952, synon nov. Type. DuckeRB i8809, Brazil, Para,
fl (holotypes, US; isotypes, P, RB, S).
LicaniaobovatifoliaMaguire, Brittonia 7: 398. 1952, synon nov. Types. Frdes22566, Brazil, Ama-
zonas, fl (holotype, NY; isotypes, COL, IAN, SP); SchunkeI51, Peru, Loreto, fl (paratypes,
A, F, NY, U, US).
Licaniawilson-brownei Maguire, Brittonia 7: 399. 1952, synon nov. Type. Wilson-Browne482 (F.D.
5888), Guyana fl (holotypes, NY; isotypes, K, US).

Small to large sized tree to 30.0 m tall, the young branches hispid-puberulous,
soon becoming glabrous and conspicuously lenticellate. Leaves obovate, elliptic to
oblong, coriaceous, 5.0-23.0 cm long, 2.5-Io.0 cm broad, apiculate to mucronate at
apex, with acumen 0-5.o mm long, glabrous above, conspicuously reticulate on
lower surface with quite or nearly parallel secondary veins, the veins appressed-
strigose with intervening spaces gray-lanate-pubescent; midrib impressed its entire
length above, glabrous except when young; primary veins 8-17 pairs, prominent
beneath, impressed above; petioles 7.0-20.0 mm long, terete, strigose-puberulous,
eventually glabrous, usually with two prominent medial glands, sometimes eglandular
or glands near lamina base. Stipules to 13.o mm long, ovate, membraneous, caducous,
intrapetiolar. Inflorescences spreading terminal panicles, the rachis and branches
tomentellous. Flowers 5.o-6.o mm long, densely disposed along primary and secon-
dary branches of inflorescence. Bracts and bracteoles 3.0-6.0 mm long, triangular,
enclosing small groups of flowers, tomentellous on exterior, glabrescent within.
Receptacle campanulate, slightly tapered at base but sessile, tomentose on exterior,
tomentose within. Calyx lobes acute, tomentellous on both surfaces. Petals 5, oblong,
short-clawed, puberulous on exterior. Stamens 18-25, inserted in a complete circle;
filaments slightly exceeding calyx lobes, free nearly to base, hirsutulous. Ovary
inserted laterally at the side of receptacle, villous. Style equalling filaments, pubescent
throughout. Fruit oblong, 6.o-8.o cm long, 3.0-4.5 cm broad, stipitate when young,
becoming sessile by expansion of the stipe; epicarp covered with large conspicuous
lenticels; mesocarp thick; endocarp thin, hard, fibrous, glabrous within.
TYPES. Martin sn, French Guiana, fl (lectotype, P; isolectotype, K); Melinon sn,
French Guiana, fl (paratypes, F, K, NY, P); 422 fl (paratype, P); 423 fl (paratype,
P, US).
Open forest at river margins, savanna margins and in secondary
DISTRIBUTION.
forest of the Guianas and Amazonia.
92 Flora Neotropica

VENEZUELA. Terr. Amazonas: Breteler4862 fl (VEN). GUYANA. Fanshawe1785 (F.D. 4521)

fr (K, NY, U). SURINAME. Lanjouw & Lindeman2663 st (K, NY, U); Lindeman& Cowan70or st
(NY, U); 7024 St (NY, U); van Donselaar1134 st (U): 1172 St (U); 1493 St (U). FRENCH GUIANA.
Benoist950 st (U); For. Serv. i69M fr (P, U); 1328 fl (U); 6242 fl (P); 7730 fl (P, U). PERU. Loreto:
Klug 1526 fl (F, NY, US); Tessmann5219 fl (NY); Ellenberg2504 st (U). BRAZIL. Amazonas: Ducke
RB 23594 fl (RB); Krukoff4880fl (A, F, MICH, MO, NY). Pari: Black50-9895 fl (IAN, NY); Capucho347
fl (F, IAN); DuckeMG 109I2 fl (BM, MG, US); MG 11758 fl (RB I5181); MG I6299 fl (BM, MG,US);
MG 16415 fl (BM, MG, US); Froes33585 fl (IAN); Frdes& Pires 24I24 fr (IAN, NY); GuedesMG 2583
fr (BM, MG, RB 5177); Pena 20 fr (IAN); Pires3r42 fr (IAN, NY); Pires,Frdes& Silva 5624 fl (IAN);
Pires & Silva 10590 fl (IAN, NY); o0611fl (IAN, NY); Sampaio5460 fl (R). Acre: Pranceet al 2978 fl
(INPA, NY).

LOCAL NAMES. Guyana: Marishiballi.Suriname: Anaura,Zwaetefoengoe.French

Guiana: Gaulette,Gris-gris.Brazil: Copuda,Pajurarana,Pianchirana.
The specimens bearing the five names cited in synonymy were analysed carefully,
and after preparing scatter diagrams, it was concluded that there is insufficient
correlation in the variation to regard L. licaniaeflora
as more than one species. There is
some range in leaf shape and size, but it is not large. The only relatively constant
feature of those characters showing variation is the absence of prominent petiolar
glands from the Guiana material. However, this character varies in presence even
among different collections from the same tree.
Licanialicaniaefiorais a very distinct species, and I follow Huber in regarding it
as the sole component of a separate subgenus. This subgenus is characterized by the
lateral position of the ovary and the large bracteoles. It is more closely related to
subgenus Moquileathan to subgenus Licania.

Licania subgen Licania

It is shown in the synopsis of the sections of subgenus Licaniathat the nature of

the lower surface of the lamina is here considered as a primary character. It is felt
that the use of this character in the definition of the sections not only brings together
groups of related species, but also makes the sections comparatively easy to recognize
from poor material.

5. Licania subgen Licania sect Hirsuta

53. Licania hirsuta Prance, sp nov Fig 14 A-D.

Frutex ad 3.0 m altus, ramulis juvenilibus hirsutulis mox glabris, haud lenticel-
latis. Folia alternata petiolata; petiolo 7.0-8.0 mm longo, hirsutulo, tereti, eglandul-
oso; laminae oblongo-ellipticae, coriaceae, I2.o-i9.0 cm longae, 5.0-8.5 cm latae,
basi cuneatae, apice in acuminem 3.0-6.0 mm longum breviter contractae, supra
glabrae, subtus praeter costas venasque pilis hirsutas glabrae; costa media supra
pubescente basi impressa, summa plana vel impressa; costis secundariis 9-io jugis,
supra impressis,subtus prominentibus;venis supra leviter impressis,sulcatis. Stipulae
lineares, 4.0-5.0 mm longae, coriaceae, extrapetiolares, persistentes. Flores 1.5 mm
longi, haud pedunculati, in paniculis racemosis terminalibus vel axillaribus dense
dispositi, rachi ramisque sparse hirsutis. Bracteae ovatae, ad 5.0 mm longum,
Licania 93

K L

FIG I4. Species of Licania.A-D, L. hirsuta(Rodrigueset al 5844); A, habit, x o.5; B, flower, X 7.5;
C, flower section, x 7.5; C, flower pubescence; D, petal, x 5. E-L, L. reticulata(INPA 4V4z, Ducke
RB 19796); E,4habit, x 0.5; F, flower, x 7.5; G, petal, x 7.5; H, flowersection, x 7.5; J, flowerpubescence,
x 7.5; K, fruit, x o.5; L, fruit section, x o.5.
94 Flora Neotropica

persistentes; bracteolae minutae, persistentes, extus hirsutae. Receptaculum cam-

panulatum, sessile, extus hirsutulum, intus tomentosum. Calycis-lobi acuti, utrinque
puberuli. Petala 5, pubescentia, haud unguiculata. Stamina 7-9, in orbem completum
disposita; filamenta glabra, calycis-lobis breviora, ad medium connata. Ovarium ad
basim receptaculi insertum, villosum. Stylus e basi ovarii ortus, ad medium villosus.
Fructua mihi ignoti.
TYPE. Froes 26137. Brazil, Amazonas, Rio Tefe, fl (holotype, NY; isotypes,
IAN, NY).
DISTRIBUTION. Flooded forest or non-flooded forest in Brazilian Amazonia.
BRAZIL. Amazonas: Manaus, Rodrigues& Loureiro5844 fl (INPA 14796, NY).

This species is most closely related to the Central American L. costaricensis.

It is
not very close to any other Amazonian species, differing in the sulcate leaves, and in
the nearly glabrous inflorescence and flowers.

54. Licania costaricensis Standley & Steyermark, Publ. Field Mus. Bot.
22: 335. I940.

Medium sized tree, the young branches hirsute, soon becoming glabrous and
lenticellate. Leaves ovate to ovate-elliptic, coriaceous, IO.o-I5.0 cm long, 6.o-io.5
cm broad, rounded to obtuse at apex, subcuneate at base; upper surface glabrous,
rugose-sulcate to bullate; lower surface glabrous except for hirsute pubescence along
venation; palisade glands rare; midrib slightly impressed above, densely pubescent;
primary veins 7-9 pairs, impressed above, prominent beneath; petioles 6.o-Io.o mm
long, hirsutulous, eglandular, terete, transversely rugose. Stipules caducous (not
seen). Inflorescences axillary racemose panicles, the rachis and branches sparsely
hirsutulous. Flowers i.5 mm long, sessile on primary branches of inflorescence, in
densely crowded clusters. Bracts and bracteoles 0.2-I.5 mm long, ovate-elliptic,
persistent, hirsutulous on exterior. Receptacle campanulate, hirsutulous on exterior,
tomentose within; pedicels 0.2 mm long. Calyx lobes acute, puberulous on both
surfaces. Petals 5, pubescent, not clawed. Stamens 3, unilateral; filaments shorter
than calyx lobes, glabrous, free to base. Ovary inserted at base of receptacle, pilose-
tomentose. Style pubescent throughout, equalling filaments. Fruit unknown.
TYPE.A. Smith1779, Costa Rica, fl (holotype, F; isotypes, MO, NY, US).
DISTRIBUTION. Known only from the type gathering which was collected in hill
pastures in the Alajuela province of Costa Rica.

55. Licania krukovii Standley, Publ. Field Mus. Bot. 27: 256. 1937.

Tree to 25.0 m tall, the young branches densely ferrugineous-tomentose, be-

coming lenticellate with age. Leaves oblong to oblong-lanceolate, chartaceous,
I1.0-20.0 cm long, 4.0-10.0 cm broad, acuminate at apex, with acumen 3.0-Io.o mm
long, rounded to subcuneate at base, glabrous or sparsely hirsute when young on
upper surface, hirsute on veins and nerves beneath; palisade glands absent; midrib
ferrugineous-tomentoseabove, plane; primary veins I3-I5 pairs, plane and hirsute
above, prominent beneath, arcuate towards lamina margins; petioles 4.0-7.0 mm
long, densely pilose-tomentose, terete, with two glands towards base. Stipules
intrapetiolar, to I.o cm long, broadly ovate, acute, persistent, tomentose on exterior.
Licania 95

Inflorescences spreading racemose panicles, the rachis and branches ferrugineous-

pilose-tomentose. Flowers 2.0-2.5 mm long, sessile and in small clusters on primary
and secondary branches of inflorescence. Bracts and bracteoles oblong-lanceolate,
I.5 mm long, acute, persistent, ferrugineous-tomentose on exterior. Receptacle
campanulate, sessile, densely pilose-tomentose on exterior, pilose within. Calyx
lobes acute, pubescent on both surfaces. Petals 5, pubescent. Stamens 5, inserted in a
nearly complete circle; filaments shorter than calyx lobes, free to base, glabrous.
Ovary inserted at base of receptacle, tomentose. Style pubescent throughout. Fruit
unknown.
TYPE.Krukoff5070, Brazil, Amazonas, fl (holotype, F; isotypes, A, BM, K, LE,
M, MICH, MO, NY, S, US).
DISTRIBUTION.Forest on high ground in western Amazonia, and upland in
Bolivia.
BOLIVIA. La Paz: Buchtein74I fl (US); Krukoff10996 fl (A, F, K, MICH, MO, NY, S, US);
I 246 fl (A, F, K, MICH, MO, NY, S, US).

56. Licania lasseri Maguire, Fieldiana Bot. 28: 253. I952.

Tree to 20.0 m tall, the young branches densely rufous-villous-tomentose.Leaves

elliptic to oblong, coriaceous, 4.0-9.0 cm long, 2.2-6.5 cm broad, rounded to
shortly apiculate at apex, rounded to subcordate at base, sparsely hirsute on upper
surface when young, soon becoming glabrous, sparsely hirsute beneath on venation;
palisade glands rare; midrib prominulous and often pubescent; primary veins 6-9
pairs, prominent beneath, prominulous and often pubescent above; petioles 3.0-4.0
mm long, densely ferrugineous-tomentose, terete, with two glands at junction with
lower surface of lamina. Stipules intrapetiolar, 4.0-5.o mm long, ovate, acute, tomen-
tose on exterior, persistent. Inflorescences spreading racemose panicles, the rachis
and branches brown-tomentose. Flowers ca 2.0 mm long, sessile on primary branches
of inflorescence. Bracts and bracteoles oblong-lanceolate, acute, 2.0-2.5 mm long,
tomentose on exterior. Receptacle campanulate, sessile, brown-tomentose on exterior,
pilose within. Calyx lobes acute, tomentose on both surfaces. Petals 5, minute, pubes-
cent, white. Stamens 5, inserted in a complete circle; filaments equalling or shorter
than calyx lobes and united at base only, the anthers deltoid. Ovary inserted at base
of receptacle, pilose. Style pubescent on lower half, equalling filaments. Fruit
(immature) epicarp rufous-velutinous.
TYPES.Lasser I730, Venezuela, Bolivar, fl (holotype, NY; isotype, F); Steyermark
59360, Venezuela, Bolivar, st (paratypes, F, NY).
DISTRIBUTION. Open forest on slopes, and savanna and river margins in Guyana
and adjacent Venezuela. Collected in flower in May and September.
VENEZUELA. Bolivar: Steyermark& Dunsterville92373 fr (NY, VEN); 92952 fr (NY, VEN);
Steyermark& Nilsson 393 fr (VEN); Steyermark& Aristeguieta75 fr (NY, VEN). GUYANA. Tillett,
Tillett & Boyan 44912 fl (F, GH, NY); 45234 fl (GH, NY).

57. Licania latifolia Bentham ex Hooker f., Mart. Fl. Bras. I4(2): i . 1867;
Fritsch, Ann. Naturh. Mus. Wien 4: 43. 1889.
Licania obovataBentham ex Hooker f., Mart. Fl. Bras. I4(2): I. I867. Fritsch, Ann. Naturh.
Mus. Wien. 4:43. I889, synon nov. Type. Spruce1569, Brazil, Amazonas, fl (holotype, K;
isotypes, BM, CGE, GH, GOET, K, LD, LE, M, NY, OXF, P).
96 Flora Neotropica

Tree to 25.0 m tall or shrub, the young branches pubescent, soon becoming
glabrous and lenticellate. Leaves obovate to oblong-elliptic, thick-coriaceous, 6.0-
20.0 cm long, 4.5-10.5 cm broad, rounded to bluntly apiculate at apex, rounded to
subcuneate at base, glabrous on upper surface, prominently reticulate beneath, with
sparse hirsute pubescence on venation; palisade glands absent; midrib plane above,
tomentose when young; primary veins 12-20 pairs, slightly impressed above, promi-
nent beneath, arcuate near lamina margins; petioles 7.0-15.0 mm long, rufous-brown
tomentose, terete, usually with two or more medial sessile glands. Stipules intrapetio-
lar, 3.0-8.0 cm long, triangular, acuminate, membraneous, persistent. Inflorescences
spreading racemose panicles, the rachis and branches ferrugineous-tomentose.
Flowers ca 2.5 mm long, sessile, in dense glomerules on branches of inflorescence.
Bracts and bracteoles to 1.5 mm long, ovate, persistent, tomentose on exterior.
Receptacle urceolate, sessile, ferrugineous-tomentoseon exterior, tomentose within.
Calyx lobes acute, tomentose on both surfaces. Petals 5, ca I mm long, pubescent.
Stamens 6-Io, inserted in a complete circle; filaments connate for half of length,
shorter than calyx lobes. Ovary inserted at base of receptacle, velutinous-tomentose.
Style pilose throughout, equalling filaments. Fruit globose; epicarp rufous-velutinous;
pericarp thin, hard, fibrous; sparsely pubescent within when young.
TYPE. Spruce457, Brazil, Para, fr (holotype, K; isotypes, CGE, NY, OXF, P).
DISTRIBUTION. Non-flooded forest in Guyana and Amazonian Brazil. Flowering
July to December.
GUYANA. Atkinson91 fl (BM, MICH); Fanshawe 2837 (F.D. 5838) fl (K, NY). BRAZIL.
Amazonas: Froes 25353 fl (IAN, NY, RB); Krukof 66i8 fr (A, BM, BR, F, K, LE, MICH, MO, NY,
RB, S, US); 7084 fr (A, BM, BR, F, K, MO, NY, RB, S, US); Pires 21 fr (COL, INPA, NY, P);
Pranceet al 2489 fr (INPA, NY). Para: Black 47-1854 fr (IAN, NY); Burchell9625 fl (K, P, US);
9778 fr (K, NY, P); Cavalcante1746 fr (MG); DuckeMG 15475 fl (BM, MG, RB, US, P); MG 15799 fr
(BM, INPA, MG, US); MG 16341 fl (BM, MG, RB, US); RB 15187 fl (RB); HuberMG 6982 fr (MG);
Kuhlmann& Jimbo 117 fl (MG); Oliveira356 fr (IAN); 895 fr (IAN); 2619 fl (IAN); R. S. Rodrigues
MG 8761 fl (BM, MG, P, R, RB, US); A. C. Smith2992 fr (A, F, MO, NY).

All the species in section Hirsuta are closely related. L. latifolia is nearest L.
krukoviibut differs in the blunt apex and greater thickness of the leaves. It is also
similar to L. lasseribut differs in the larger number of primary leaf veins as well as in
the greater number of stamens which have fused filaments. Further material is
needed to determine whether L. krukoviiis indeed a distinct species. Its separate
recognition here is based on the small differences used by Standley to distinguish it
from L. latifolia, and which are valid in the material seen by me.

6. Licania subgen Licania sect Hymenopus

58. Licania minuscula Cuatrecasas, Fieldiana Bot. 27(2): 113. 1951.

Licania minutifloraCuatrecasas, Fieldiana Bot. 27: 64. I950, nom illegit, non minutiflora(Sagot)
Fritsch.

Large tree, the young branches puberulous, becoming glabrous but not lenticel-
late. Leaves elliptic-lanceolate, chartaceous, I I.0-22.0 cm long, 4.2-8.5 cm broad,
cuspidate at apex, with finely pointed acumen I4.0-30.0 mm long, cuneate at base,
glabrous above, glabrous or with very few short appressed hairs beneath; palisade
glands frequent; midrib prominulous above, glabrous; primary veins 7-9 pairs,
Licania 97

slightly impressed above, prominent beneath, arcuate and anastomosing near lamina
margin; petioles 4.0-5.0 mm long, terete, glabrous, eglandular. Stipules lanceolate,
to 5.0 mm long, hirsutulous, caducous. Inflorescences terminal and axillary panicles,
the rachis and branches sparselypuberulous. Flowers ca I.o mm long, in small groups
on short secondary branches of inflorescence. Bracts and bracteoles minute, ovate,
persistent, sparsely puberulous on exterior, entire. Receptacle turbinate, tomentose
within; pedicels ca 0.5 mm long. Calyx lobes acute, sparsely pilose on exterior,
glabrous within. Petals 5, pubescent, not clawed. Stamens 3, unilateral; filaments very
short, glabrous, free to base. Ovary inserted at base of receptacle, pilose. Style pilose
throughout, shorter than filaments. Fruit unknown.
TYPES.Cuatrecasas21546, Colombia, Choc6, fl (holotype, F; isotype, COL);
21234, Colombia, Valle, fl (paratypes, COL, F).
DISTRIBUTION. Known only from the type gatherings collected in the Pacific
coastal region of Colombia. Flowering in May.

59. Licania operculipetala Standley & L. O. Williams, Ceiba 3: 46. I952.

Large tree to 35.0 m tall, buttressed to about 2.5 m, the young branches glabrous
but not lenticellate. Leaves elliptic, coriaceous, 12.0-1 7.0 cm long, 4.0-6.0 cm broad,
obtuse or short-acuminate at apex, with acumen to 7.0 mm long, subcuneate at base,
glabrous on both surfaces; palisade glands frequent beneath; midrib prominulous
above, glabrous; primary veins 8-i2 pairs, prominulous on both surfaces; petioles
5.0-7.0 mm long, glabrous, terete, with two glands at junction with lower
surface of lamina. Stipules to 2.0 mm long, subulate, intrapetiolar, persistent.
Inflorescences terminal panicles, the rachis and branches almost glabrous. Flowers
1.5 mm long, in small groups on short secondary branches (peduncles) of inflores-
cence. Bracts and bracteoles minute, persistent, entire. Receptacle turbinate, almost
glabrous on exterior, tomentose within; pedicels 0.5-I.0 mm long. Calyx lobes acute,
glabrous, ciliate. Petals 5, pubescent. Stamens 5, inserted in a complete circle;
filaments shorter than calyx lobes, glabrous, free to base. Ovary inserted at base of
receptacle, glabrous. Style glabrous, scarcely exceeding ovary. Fruit unknown.
TYPE.Allen 6030, Costa Rica, fl (holotype, EAP, not seen; isotypes, GH, US).
DISTRIBUTION.Known only from two collections from the forested hills of
Puntarenas Province of Costa Rica. Flowering March.
COSTA RICA. Allen 6032 fl (F, MEXU).

can be distinguished from related species by the shape of the

Licaniaoperculipetala
inflorescence,the glabrous ovary, and the glabrous exterior of the receptacle and calyx
lobes.

60. Licania reticulata Prance, sp nov Fig 14 E-L.

Arbor, ramulisjuvenilibus glabris, lenticellatis. Folia alternata petiolata; petiolo

7.0-12.0 mm longo, glabro, tereti, subtus sub lamina cum glandulis duobus sessilibus
munito; laminae oblongo-ellipticae vel oblongae, coriaceae, 8.0-24.0 cm longae,
3.3-Io0. cm latae, basi rotundatae vel subcuneatae, apice in acuminem 2.0-5.0 mm
longum contractae, utrinque glabrae reticulatae, subtus glandulosae; costa media
supra glabra, plana; costis secundariis I -I3 jugis utrinque prominentibus. Stipulae
98 Flora Neotropica

ad 2 mm longae, membranaceae, lanceolatae, extrapetiolares, caducae. Flores

2.5-3.0 mm longi, breviter pedunculati, cymosi, in paniculis terminalibus dispositi,
rachi lenticellati, rachi ramisque sparse puberulis. Bracteae bracteolaeque ca I mm
longae, ovatae, caducae. Receptaculum campanulatum, extus puberulum, intus
tomentellum; pedicelli 0.25-1.5 mm longi. Calycis-lobi acuti, utrinque puberuli.
Petala 5, sparse pubescentia. Stamina 6-7, in orbem completum disposita; filamenta
pubescentia, ad medium connata, calycis-lobos aequantia. Ovarium ad basim
receptaculi insertum, sparsevillosum. Stylus e basi ovarii ortus, calycis-lobosaequans,
lanato-pubescens. Drupa oblonga, ad 7.0 cm longus, extus laevis, glabra, longitudi-
naliter costata; mesocarpio tenue, carnoso; endocarpio tenue, duro, intus glabro.
TYPE. I.N.P.A. 414I, Brazil, Amazonas, Manaus, fl (holotype, NY; isotypes,
IAN, INPA).
DISTRIBUTION. Non-flooded and periodically flooded forest in Amazonian Peru
and Brazil.
PERU. Loreto: Tessman5370 fr (NY). BRAZIL. Amazonas: Manaus, INPA 5730 fl (INPA, NY);
ChagasINPA 952 fl (INPA, NY); Ducke488 fl (A, F, IAN, K, MO, NY, R, RB 35569, S, US); 2209 fl
(IAN, NY, R, US); RB 19796 fr (K, RB); RB 23602 fl (K, RB); RB 25031 fl (K, RB); Pranceet al 5012
fl (INPA, NY).

Licaniareticulatais related to L. heteromorpha but differs in the glabrous costate

fruit, the pedunculate groups of flowers, and the connate filaments. The prominent
reticulation of the upper surface of the leaf makes this an easily recognized species.

61. Licania arachnoidea Fanshawe & Maguire, Bull. Torrey Club 75: 3I8.
1948.

Medium-sized tree, the young branches puberulous, soon becoming glabrous.

Leaves narrowly oblong-lanceolate with parallel sides, coriaceous, I7.o-25.0 cm
long, 4.5-7.0 cm broad, acuminate to acute at apex, with acumen to I I.o mm long,
rounded at base, glabrous on both surfaces; palisade glands absent; midrib promi-
nent and glabrous above; primary veins 12-14 pairs, prominulous above, prominent
beneath; petioles 7.0-9.0 mm long, glabrous, canaliculate, rugose, with two disc-
shaped, medial glands above. Stipules linear, 10.0-30.0 mm long, submembraneous,
intrapetiolar, persistent. Inflorescences axillary racemose panicles, ramiflorous, the
rachis and branchessparsely arachnoid. Flowers 3.5-4.0 mm long, in dense glomerules
on primary branches of inflorescence. Primary bracts ovate, to 8.o mm long, other
bracts and bracteoles 1.0-3.0 mm long, persistent, pubescent on exterior, glabrous
within, often slightly serrate.Receptacle narrowly urceolate, sessile, densely arachnoid
on exterior, lanate within. Calyx lobes acute, pubescent on exterior, glabrous within.
Petals 5, densely pubescent, some shortly unguiculate. Stamens 5 fertile, inserted
in a complete circle, with about 4 sterile staminodes; filaments slightly exceeding
calyx lobes, connate for half of length, with a dense lanate mass surrounding them
and filling the mouth of the receptacle. Ovary inserted at base of receptacle, lanate-
tomentose. Style pubescent throughout, equalling filaments. Fruit unknown.
TYPE.FanshaweI495 (F.D. 4231), Guyana, fl (holotype, NY; isotypes, IAN, K).
DISTRIBUTION.Non-flooded forest in Guyana and Amazonian Brazil.
BRAZIL. Amazonas: Frdes20910 fl (K, NY, US).

Licaniaarachnoidea but is distinct in a number of im-

is close to L. macrophylla,
portant featureswhich were listed by Fanshawe & Maguire in the original description.
Licania 99

The most significant differences are the arachnoid pubescence and disposition of the
flowers.

62. Licania oblongifolia Standley, Publ. Field Mus. Bot. 17: 257. I937.

Tree to 30.0 m tall, the young branches glabrous, not lenticellate. Leaves oblong,
with parallel sides, coriaceous, 6.o-I 7.0 cm long, 2.5-6.0 cm broad, rounded at apex,
acute to short-acuminate, with acumen to 5.0 mm long, rounded to subcordate at
base, glabrous on both surfaces; palisade glands occasional; midrib prominulous
above, glabrous; primary veins 9-15 pairs, prominent beneath, prominulous above;
petioles 5.0-I 1.0 mm long, glabrous, shallowly canaliculate, usually with two sessile
glands at base of lamina. Stipules linear, 3.0-5.0 mm long, submembraneous, persis-
tent. Inflorescences axillary panicles, the rachis and branches puberulous. Flowers
1.5-2.0 mm long, solitary on primary and secondary branches of inflorescence.
Bracts and bracteoles 0.2-1.0 mm long, persistent, puberulous on exterior, entire.
Receptacle campanulate, puberulous on exterior, puberulous within; pedicels
0.5-1.0 mm long. Calyx lobes acute, puberulous on both surfaces. Petals 5, densely
lanate, usually short-unguiculate. Stamens 5-6 fertile, inserted in a complete circle;
filaments equalling calyx lobes, slightly connate at base, with a dense lanate mass
surrounding the filaments and filling the mouth of receptacle. Ovary inserted at base
of receptacle, lanate-tomentose. Style pubescent throughout, equalling filaments.
Fruit globose to ovoid, to 6.o cm wide and 9.0 cm long; epicarp glabrous, smooth,
or crustaceous-pulverulent, the surface deeply and broadly rugose when mature;
mesocarp 4.0-7.0 mm thick, fibrous, hard when dry; endocarp thin, granular, fragile,
glabrous and shining within.
TYPES.Krukoff6812, Brazil, Amazonas, fr (holotype, F; isotypes, A, BR, K, LE,
MICH, MO, NY, RB, S, US); 7216 fr (paratypes, A, BR, F, K, LE, MICH, MO,
NY, RB, S, US); I452, Brazil, Mato Grosso, fr (paratypes, A, F, K, MICH, MO,
NY, P, S, US).
DISTRIBUTION. Non-flooded forest in Amazonian Brazil. Flowering August to
December.
BRAZIL. Amazonas: ChagasINPA 3438b fr (NY); CoelhoINPA 3057b fr (IAN, INPA); 5986 fr
(INPA); Ducke303 fl (A, F, K, MO, NY, R, S, US); RB 23601 fl (K); RB 35553 fr (RB); RB 35568 fl
(K, RB); RB 35570 fl (RB); FerreiraINPA 5912 fl (IAN, INPA, NY); Froes26300 fr (IAN); Guedes20
fl(IAN, NY); LoureiroINPA 16139 fl (INPA); I6566 fl (INPA, NY); Rodrigues5520 fl (INPA 14193,
NY); 7398fr (INPA 16799, NY); Rodrigues& Coelho7266fl (INPA 16586, NY). Par,: Black 47-2150
st (IAN, NY). Mato Grosso: Maguire,Pires, Maguire & Silva 56604 fl (NY).

LOCALNAMES.Castanha, Pajurd-rana, Macucu chiador.

Licaniaoblongifoliais close to L. macrophyllabut differs by its much smaller flowers
and leaves. This is a species of western Amazonia, while L. macrophylla is of eastern
Amazonia and the Guianas, their ranges scarcely overlapping. Field notes indicate
that L. oblongifoliais a species of non-flooded forest while L. macrophyllais a species of
flooded forest.

63. Licania macrophylla Bentham, Jour. Bot. Kew Misc. 2: 240. I850;
Hook. f., Mart. Fl. Bras. 14(2): 9. I867; Fritsch, Ann. Naturh. Mus.
Wien 4: 42. 1889; Benoist, Bull. Mus. Hist. Nat. Paris 25: 512. 1919;
Kleinh. in Pulle, Fl. Suriname 2(I): 43I. 1939.

Medium to large-sized tree to 30.0 m tall, the young branches almost glabrous.
100 Flora Neotropica

Leaves narrowly oblong with parallel margins, coriaceous, I3.o-40.0 cm long,

4.5-Io.5 cm broad, acuminate to obtuse at apex, with acumen 6.o-i .o mm long,
rounded to cordate at base, glabrous on both surfaces; palisade glands absent;
midrib prominulous above, glabrous; primary veins I2- 6 pairs, prominent beneath,
prominulous above; petioles 6.0-I4.0 mm long, glabrous, usually shallowly canali-
culate, with two sessile glands at base of lamina. Stipules linear, 6.0-i5.0 mm long,
submembraneous, intrapetiolar, persistent. Inflorescences axillary racemose panicles
on woody stems, the rachis and branches sparsely puberulous. flowers 2.5-3.0 mm
long, solitary or in small groups on primary and secondary branches of inflorescence.
Bracts and bracteoles ovate to lanceolate, 1.0-3.0 mm long, puberulous on both
surfaces, peristent, entire. Receptacle campanulate to shortly urceolate, puberulous
on exterior, arachnoid within; pedicels 0.25-o.5 mm long. Calyx lobes acute, puberu-
lous on both surfaces. Petals 5, pubescent, shortly unguiculate. Stamens 5 fertile,
with 5-6 staminodes, inserted in a complete circle; filaments slightly exceeding calyx
lobes. connate at base to form a short tube, with a dense lanate mass surrounding
filaments and filling mouth of receptacle. Ovary inserted at base of receptacle, lanate-
tomentose. Style pubescent throughout, equalling filaments. Fruit ovoid to pyriform,
to 8.o cm diameter; epicarp smooth, glabrous or crustaceous-verrucose,drying black,
mesocarp ca 8.o mm thick, hard when dry; endocarp thin, granular, glabrous within.
TYPE.SpruceI39, Brazil, Para, fl (holotype, K; isotypes, CGE, NY, OXF, P).
DISTRIBUTION. Periodically flooded forest in the Guianas and eastern Amazonian
Brazil.
SURINAME. B.B.S. I48 st (BBS, U); 1031 st (BBS, U); B.W. 32 fl (U); 3687 fr (U); 4307 st
(U); 6I84 st (A, MO, U); vanDonselaar1519 st (NY, U); Focke102I fl (U); Heinsdyk62 st (U); Lanjouw
& Lindeman326 st (NY, U); 420 st (U); 6i8 fl (NY, U); 3458 st (U); Mennega331 fl (U); Stahel I43 fl
(A, NY, U); Wullschlaegel1650 st (BR, GOET, NY, U); I651 fl (BR). FRENCH GUIANA. Bettenfeld
sn st (NY, P); For. Serv. i49M st (P, U); 6087 st (P); 7480 st (P); Melinonsn (1863) fl (A, BM, BR, F,
GH, LA, NY, P, US); 117 fl (P); 209 fl (P); 1ogo st (K); Sagot 462 st (P); 1o09 fl (K, P); Wachenheim
124 fl (BR, P); 185 fl (P); 349 fl (P). BRAZIL. Amazonas: INPA 6028 fr (INPA, NY); Silva & Brazao
60774 fl (NY). Para: Archer782I fr (K, S); 7824 fl (F, K, NY, US); 7903 fl (F, K, NY, S); Black &
Foster48-3414 fl (IAN, INPA, U); DuckeMG I5529 fl (BM, RB); RB 21325 fl (K, RB); Frdes32438 fl
(IAN, INPA, R); HuberMG 1147 fl (BM, MG, RB); Lima & Black 48-3o18 fl (IPA); Pires 1840 fl
(IAN, NY, RB); 51870 st (NY); Prance& Silva 58813 fl (F, NY, UB, US); Prance& Pennington1752 fl
(F, GH, NY); R. S. RodriguesMG 8772 fl (MG, RB). Terr. Amapa: Bastos 54 fl (F, INPA, NY, RB
96385); Froes25928 fr (IAN); 26632 fl (NY, SP); Pires, Rodrigues& Irvine51470 fl (F, INPA, MG, NY,
US); Westra48556 fl (F, MG, NY, US).

LOCAL NAMESAND USES. Suriname: Sponshout, Sponshoedoe,Anaura (Arawak),

French Guiana: Gris-griscoumate.Brazil: Anauera,Pintadinha.
Alauna, Sergeants-kloot.
The wood is used in boat building and general construction.

64. Licania caudata Prance, sp nov Fig I6 J-M.

Arbor parva ramulis juvenilibus glabris, lenticellatis. Folia alternata petiolata;

petiolo 5.0-7.0 mm longo, glabro, eglanduloso, leviter canaliculato; laminae ovato-
ellipticae, subcoriaceae, 8.0-i I.0 cm longae, 3.5-5.5 cm latae, basi cuneatae apice
in acuminem 7.0-9.0 mm longum contractae, utrinque glabrae, subtus glandulas
paucas ferentes; costa media supra glabra, leviter prominente; costis secundariis
8-9 jugis, subtus prominentibus supra planis. Stipulae caducae (non visae). Flores
circa 2.0 mm longi, breviter pedunculati, in paniculis racemosis terminalibus
axillaribusque dispositi, rachi ramisque glabris vel sparse hirsutulis.Bracteae bracteo-
laeque minutae, lanceolatae, caducae. Receptaculum campanulatum, extus sparse
 Licania 101

.~, I

. ..

'.
. . .. .
? ~ ~ ~ ~~ ~ ~ ~ ~ ~ ~ ~ ~ ~~~~~~?
..

~"?(~~
~t'l
Y~F?I ~ ~ . ... .
~"~ ,~i
.r :' ? ES
.,.h .' .i/'
?!:i'.~. .
?1~~~~~~.1 .~.
? ".i ./~".~? ~'~"? . . j
.:?'??~~ ? ..........
...
'~,:X:I;?L~:;~,~Ci?"~_,;~ .:, :..... : ?\ .
. c.....
?;r'~' ?.-... . ..,.?. ....
, ...~ . .:?-~~~~~~~~~~~~~~..
:.~.._..~. -. ?. ........
~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~...
.
.-':":..-. .,.~~~~~~~1
~.~-..,.,~.~.:. . ?~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~I;
......, ..~~~~~~~~~~~~~~~~~~*

r~~~~~~~~~~~ .
E, flower, x
X I; D, flower section, x 5; IO..; F, petal, x 8.

: :?t;~~~~~~~~~

a, ~ ~ .:.
x z; D, FIGz5.Liani x ltisi~~la(Brti346 ~bal 3). x ~8 , hbi, xo.; B fuit x;C
frit~'ton
zS; E, flower,x xo; F, petal,
flower section,

FIG I5. Licania latistipula(Berti 346, Zabala3z). A, habit, x o.s ; B, fruit, x I; C. fruit setion,
102 Flora Neotropica

Species Licania.A-D,L.anshawei(Steyermark87557);Aus,
DISTRIBUTION6. of habit, x5;B,x.o; flower,
C,, flower
flower section,
sectionL.x IO;
Amax D, flower
zonas:Manaus; pubescence, x IO.
flower pubescence, E-H, L.L. glabriflora
o. E-H, (Steyermark
glabriflora E, habit,
87907); E,
(Steyermark
87907); habit,
x .5; F, flower flower
12(INPA.5 X 12.5; G, section, X 12.5 H, flower pubescence, x J-M, L. caudata(D.
Coelho3946); J, habit, x o-s;
o.5; K, flower, x 5; L, flower section, x 5; M, flower pubescence, x 5.
5.

hirsutulum, intus tomentosum; pedicelli circa 0.5 mm longi. Calycis-lobi acuti,

utrinque sparse hirsutuli. Petala 5, sparse pubescentia. Stamina 7-8 in orbem com-
pletum disposita; filamenta glabra, ad medium connata, cum calycis-lobis aequantia.
Ovarium ad basim receptaculi insertum, tomentosum. Stylus e basi ovarii ortus,
calycis-lobos aequans, ad apicem pubescens. Fructus mihi ignoti.
TYPE. D. CoelhoINPA 3946, Brazil, Amazonas, Manaus, fl (holotype, NY;
isotypes, IAN,
isotypes, INPA, MG).
IAN, INPA, MG).
DISTRIBUTION. Non-flooded forest in the region of Manaus, Brazil.
BRAZIL. Amazonas:'Manaus; Rodrigues,Coelho& Lima 2772 fl (INPA); Rodrigues& Osmarino
6894 fl (INPA 15445).
Licania 103

Licaniacaudatais most closely related to L. heteromorpha

from which it differs in the
caudate leaf apex, the almost glabrous exterior of the flowers and inflorescence, and
the connate filaments.

65. Licania latistipula Prance sp nov Fig 15.

Arbor magna, ramulis juvenilibus glabris lenticellatis. Folia alternata petiolata;

petiolo 3.0-8.0 mm longo, eglanduloso, glabro, tereti; laminae oblongo-ellipticae,
6.0-I5.5 cm longae, 2.5-7.5 cm latae, basi subcuneatae, apice acutae vel breviter
acuminatae, utrinque glabrae; costa media supra leviter prominente, glabra; costis
secundariis 7-9 jugis, supra planis, subtus leviter prominentibus. Stipulae ovatae,
foliaceae, glabrae, 5.0-I7.0mm longae, 3.0-I.0 mm latae, coriaceae, subpersis-
tentes, axillares. Flores circa I.o mm longi, haud pedunculati, in paniculis terminali-
bus axillaribusque dispositi, rachi ramisque glabrae. Bracteae bracteolaeque minutae,
glabrae, persistentes. Receptaculum sessile, campanulatum, extus glabrum, intus
tomentosum. Calycis-lobi acuti, extus glabri. Petala 5, pubescentia, margine ciliata.
Stamina 3, unilateralia, connata, calycis-lobos aequantia. Ovarium ad basim recepta-
culi insertum, glabrescente vel sparse tomentosum. Stylus e basi ovarii ortus, calycis-
lobis brevior, tomentosum. Drupa oblonga, 2.0-3.0 cm longa, extus laevis, glabra;
mesocarpio tenue, carnoso; endocarpio tenue, duro, lignoso, fibroso, intus glabro.
TYPE. Berti 346, Venezuela, Terr. Delta Amacuro, east of Rio Grande, east-
northeast of El Palmar, near boundary of Bolivar State, fl (holotype, VEN; isotype,
VEN).
DISTRIBUTION. Forest on high ground, collected only in the Orinoco Delta in
Venezuela.
VENEZUELA. Terr. Delta Amacuro,eastnortheastof El Palmar, ZabalaI fr (VEN); 3I fr
(VEN); 142 st (VEN).

LOCAL NAME. Pildn Nazareno.

Licanialatistipulais easily distinguished from other species of Licaniaby the large
broad foliaceous stipules. It is most closely related to L. divaricatabut, in addition to
the stipules, it differs in part by the glabrous inflorescence,the less prominent primary
veins, the smaller flowers, and the fewer stamens with united filaments. It differs from
L. glabrifoliapartly in the glabrous inflorescence, the larger leaves with blunt apices,
the eglandular glabrous petioles, and the fewer stamens with united filaments. It
differs from L. caudatain the blunt leaf apices, the terete stipules, the smaller flowers,
the fewer stamens, and the almost glabrous ovary.

66. Licania divaricata Bentham, Jour. Bot. Hooker 2: 22 . 1840; Sandwith,

Kew Bull. 1931: 37I. 1931; Kleinh. in Pulle, Fl. Suriname a(I): 433. I937.
(Bentham)Fritsch,Ann. Naturh. Mus. Wien 4: 45
Benthamvar divaricata
Licaniaheteromorpha
1889.

Small to medium-sized tree to I8.o m tall, the trunk often with stilt roots; the
young branches puberulous, soon becoming glabrous and lenticellate. Leaves elliptic
to oblong-elliptic, thick-coriaceous, 9.0-27.0 cm long, 4.5-0o.0 cm broad, abruptly
short-acuminate at apex, with acumen 2.0-10.0 mm long, subcuneate at base,
glabrous on both surfaces;palisade glands present; midrib plane to slightly impressed
104 Flora Neotropica

above, glabrous; primary veins 6-7 pairs, extremely prominent beneath, inconspicu-
ous above; petioles 5.0-9.o mm long, glabrous, terete, with two sessile glands near
lamina base. Stipules lanceolate, coriaceous, to 4.0 mm long, adnate to petiole base,
subpersistent.Inflorescencespredominantly axillary panicles, the rachis and branches
sparsely appressed-hirsutulous.Flowers ca 1.5 mm long, solitary but densely crowded
on primary and secondary branches of inflorescence. Bracts and bracteoles 0.5-2.0
mm long, oblong to lanceolate, hirsutulous, persistent. Receptacle campanulate, al-
most sessile, sparsely hirsutulous on exterior, sparsely pubescent within. Petals 5,
pubescent, not clawed. Stamens 7, inserted in a complete circle; filaments slightly
shorter than calyx lobes, connate for one third of length, glabrous. Ovary inserted at
base of receptacle, villous. Style equalling filaments, hirsute throughout. Fruit ovoid;
epicarp smooth, glabrous, drying black; mesocarp thin, fleshy; endocarp thin,
granular, fragile, glabrous within.
TYPE.Schomburgk 463, Guyana, fl (holotype, K; isotypes, BM, CGE, G, L, NY,
OXF, P).
DISTRIBUTION. Forest and savanna margins of the Guianas and adjacent Brazil,
on sandy soil.
GUYANA. Davis 587 (F.D. 2628) fl (FHO, K); Fanshawe 1315 (F.D. 4051) fl (K, NY); Hohenkerk
zo4 fl (F); Tutin232 fl (BM, K, RB,
705 fl (K); Irwin13r fl (US); 184fr (US); 383 fl (MICH); Persaud
U, US); 294 fl (BM, K, RB, U, US). SURINAME.B.B.S. io fl (U); B.W. 1301St (U); I496St (U);
1590 St (K, NY, U); 2825 fl (U); 2909 fl (A, MO, U); Heyligers343 st (U); 524 st (U); Lanjouw&
Lindeman1833 st (U); Lindeman3896 st (U); 4137 fl (NY, U); 4205 fr (NY, U); 6375 st (U); Stahel
158 st (A, K, NY, U); 158a st (U). BRAZIL. Par,: Frdes31920 fl (IAN).

LOCALNAMES. Guyana: Buruburuli.Suriname: Boroborelli(Arawak), Itoeloetano

japopalli,Anaura, Sabaneanaura.
I agree with those authors who have maintained this as a species separate from
L. heteromorpha, rather than with Fritsch who regarded it as a variety, because L.
in the leaf venation and in the
divaricatadiffers from the variable L. heteromorpha,
pubescence of the inflorescence and flowers.

67. Licania glabriflora Prance, sp nov Fig I6 E-H.

Arbor parva vel media, ramulis juvenilibus hirsutulis mox glabris lenticellatis.
Folia alternata petiolata; petiolo 2.0-5.0 mm longo, tereti, eglanduloso, hirsuto vel
hirsutulo; laminae oblongae vel oblongo-ellipticae, submembranaceae, 7.0-I .0 cm
longae, 2.5-4.2 cm latae, basi cuneatae, apice in acuminem 5.0-9.0 mm longum
contractae, utrinque glabrae, subtus saepius prope petiolum glandulis duobus
munitae, costa media supra leviter prominenti, hirsutula, mox glabra; costis secun-
dariis 8-iojugis, subtus prominentibus, supra impressis. Stipulae ad 3.5 mm longae,
lanceolatae, persistentes, intrapetiolares, hirsutulae. Flores circa 2.0 mm longi, haud
vel brevissime pedunculati, in paniculis racemosis saepissime axillaribus, vel ter-
minalibus dispositi, rachi ramisque sparse hirsutis. Bracteae bracteolaeque minutae,
lanceolatae, caducae. Receptaculum campanulatum, breviter pedicellatum, extus
hirsutulum, intus tomentosum. Calycis-lobi acuti, sparse hirsutuli. Petala 5, dense
pubescentia. Stamina 5 in orbem 3/4 disposita; filamenta glabra, ad medium con-
nata, cum calycis-lobis aequantia. Ovarium ad basim receptaculi insertum, pilosum.
Stylus e basi ovarii ortus, calycis-lobos aequans, ad apicem hirsutus. Fructus mihi
ignoti.
TYPE.Steyermark 87907, Venezuela, Bolivar: Sierra Imataca, Rio Reforma, fl
(holotype, NY).
Licania 105

Non-flooded forest in Venezuela to Para, Brazil.

DISTRIBUTION.
FRENCH GUIANA. Oldeman 2463 (fr P). BRAZIL. Par,: Belterra,Black47-1738 fl (IAN,
555 fl (IAN, UB).
NY); 47-1912, fl (IAN, NY); Bel1m-Brasiliaroad, Km x67, Oliveira

Licaniaglabrifoliabelongs to the L. heteromorpha complex, but differs from L.

heteromorpha in the hirsutulous inflorescence and flowers, the connate stamens, and the
slightly impressed primary leaf veins. The present species is also closely related to
L. caudatabut differs in the leaf shape, and the terete petioles, the more hirsutulous
petioles, the inflorescencebranching pattern and the leafvenation.

68. Licania intrapetiolaris Spruce ex Hooker f., Mart. Fl. Bras. 14(2): II.
1867; Fritsch, Ann. Naturh. Mus. Wien 4: 4. 1889.

Tree to 20.0 m tall, the young branches puberulous or hispid, becoming glabrous
with age. Leaves broadly elliptic to oblong-elliptic, 9.0-23.0 cm long, 4.5-I4.0 cm
broad, rounded to acute at apex, rounded at base, glabrous on both surfaces; midrib
plane or nearly so above, glabrous; primary veins 8-12 pairs, extremely prominent
below, prominulous above; petioles 4.0-7.0 mm long, terete, glabrous or sparsely
puberulous, sometimes hispid when young, with two prominent protuding medial
glands. Stipules linear, to I5.0 mm long, persistent, intrapetiolar. Inflorescences
terminal and subterminal racemose panicles, the rachis and branches tomentellous.
Flowers 2.0-2.5 mm long, sessile along primary and secondary branches of in-
florescence. Bracts and bracteoles 1.0-2.0 mm long, lanceolate, persistent, tomentel-
lous on exterior, glabrous within, Receptacle campanulate, tomentellous on exterior,
tomentose within, sessile. Calyx lobes acute, puberulous. Petals 5, pubescent. Stamens
ca 5, inserted in a complete circle; filaments shorter than calyx lobes, glabrous, free
to base; anthers deltoid. Ovary inserted at base of receptacle, tomentose. Style rising
to base of filaments, hirsute throughout. Fruit globose, ca 1.5 cm diameter; epicarp
ferrugineous-velutinous; mesocarp thin; endocarp thin, fibrous, glabrous within.
TYPE.Spruce3539, Venezuela, Terr. Amazonas, fl (holotype, K; isotypes, BM,
BR, CGE, LE, NY, OXF, P).
DISTRIBUTION. Lowland forest in open places in Guyana and Amazonian
Venezuela.
VENEZUELA. Terr. Amazonas: Maguire,Cowan & Wurdack3oo008 fr (F, NY, US); Maguire &
Wurdack35556 fr (NY); Maguire, Wurdack& Bunting37652 fr (NY, US); LI. Williams I3987 fr (VEN).
GUYANA. Fanshawez680 (F.D. 44I6) fr (K); i68I (F.D. 4417) fl (K, NY). BRAZIL. Terr. Roraima:
Pranceet al 10121 fl (INPA, NY).

LOCALNAME.Guyana Buruburuli.
but differs in the large thick leaves with a
This species is close to L. heteromorpha,
broad midrib up to 3.5 mm thick at the base as well as in the deltoid anthers.

69. Licania heteromorpha Bentham, Jour. Bot. Hooker 2: 22 . I840; Hook.

f., Mart. Fl. Bras. 14(2): I2. I867; Sagot, Ann. Sci. Nat. Bot. VI. 15: 307.
1883; Fritsch, Ann. Naturh. Mus. Wien 4: 44. 1889; Benoist, Bull. Mus.
Hist. Nat. Paris 25: 512. 1919; Sandwith, Kew Bull. 1931: 370. I93I;
Kleinh. in Pulle, Fl. Suriname 2: 433. 1939. Fig. i8.

Tree to 30.0 m tall, occasionally slightly buttressed, the young branches hispid
106 Flora Neotropica

or puberulous, soon becoming glabrous. Leaves elliptic-orbicular to oblong-lanceolate

coriaceous, 4.5-22.0 cm long, 2.0-10.o cm broad, most frequently rounded at apex
but varying from retuse to acute or bluntly acuminate, rounded to cuneate or rarely
subcordate at base, glabrous on both surfaces; midrib prominulous above, glabrous;
primary veins 6-14 pairs, plane or nearly so above, prominent beneath; petioles
2.0-I2.0 mm long, glabrous when mature, usually shallowly canaliculate, less often
terete, most frequently bearing two protuding prominent medial glands, rarely
eglandular. Stipules 1.5-3.5 mm long, membraneous, subpersistent or caducous,
linear. Inflorescences terminal and subterminal racemose panicles, the rachis and
branches puberulous to tomentellous. Flowers 1.5-3.0 mm long, in small groups along
primary and secondary branches of inflorescence, sessile or subsessile. Bracts and
bracteoles 0.3-2.0 mm long, linear to ovate, persistent. Receptacle campanulate,
tomentellous to tomentose on exterior, tomentose within. Calyx lobes acute, puberu-
lous on both surfaces. Petals 5, pubescent. Stamens 5-7, inserted in a complete circle;
filaments shorter than calyx lobes, free to base. Ovary inserted at base of receptacle,
lanate to short-pubescent. Style rising to base of anthers, pubescent. Fruit round to
oblong; epicarp smooth or less frequently markedly costate, velutinous-pubescent or
glabrous and verrucose; mesocarp thin, fleshy; endocarp very thin, granular, hard,
glabrous within.

Key to the Varieties of Licaniaheteromorpha

i. Leaf base distinctly cordate. c. var subcordata.
I. Leaf base rounded to cuneate.
2. Fruit deeply costate, often oblong; leaves usually oblong-lanceolate, cuneate at base;
young stem never hispid. b. var glabra.
2. Fruit terete or very slightly costate; leaves broader, more oblong, usually subcuneate to
rounded at base; young stem usually hispid.
3. Leaf reticulation lax. d. var perplexans.
3. Leaf reticulation intricate. a. var heteromorpha.

69a. Licania heteromorpha var heteromorpha

Licania heteromorphaBentham, Jour. Bot. Hooker 2: 22. 1840.
Licania benthamiiIIooker f., Mart. Fl. Bras. 14(2): 12. 1867, synon nov; Fritsch, Ann. Naturh.
Mus. Wien 4: 45. I889. Type. Spruce3278, Venezuela, Terr. Amazonas, fl (holotype, K;
isotypes, BM, BR, C, CGE, GH, GOET, LD, LE, NY, OXF, P).
Licaniaprismatocarpa Spruce ex Hooker f., Mart. Fl. Bras. 14(2): I9. 1867; Fritsch, Ann. Naturh.
Mus. Wien 4: 59. I889, synon nov. Type. Spruce3490, Brazil, Amazonas, fr (hclotype, K;
isotypes, BR, CGE, GH, LD, LE, NY, OXF, P).
Licania biglandulosaGrisebach ex Urban, Symb. Antill. 5: 354. x908, synon nov. Types. Crueger
o05, Trinidad fl (syntype, GOET; isosyntype, K); Eggers 113 (nv); Trin. Bot. Gard. 1439 fl
(isosyntype, S); Eggers5842, Tobago, fl (isosyntypes, K, S).

Leaves ovate to oblong, usually elliptic, rounded to subcuneate at base, the

reticulation intricate; fruit smooth or rarely slightly costate, usually globose, less
frequently ellipsoid.
TYPE. Schomburgk 873, Guyana, fl (holotype, K; isotypes, BM, F, GH, L, NY,
OXF, P, US).
An abundant forest species of Trinidad, Venezuela, the Guianas
DISTRIBUTION.
and Amazonian Brazil, commonest in periodically flooded forest, but occuring else-
where.
Licania 107

Representative collections: VENEZUELA. Carabobo: Steyermark 95334 fr (NY, VEN). Sucre:

Steyermark& Agostini91377 fl fr (NY, VEN). Terr. Delta Amacuro: Steyermark 87282 fr (F, GH, NY,
VEN); 87409 fl (NY, US); 87451 fl (F, MO, NY, US, VEN); Wurdack& Monachino39657 fr (F, GH,
LE, MICH, MO, NY, SP). Apure: Velez2275 fr (VEN); 2543 fr (VEN); Wurdack& Monachino41363
fr (F, NY). Bolivar: Bernardi1509 fr (VEN); sn fl (VEN); Cardona389 fl (VEN); 2573 fl (NY, US,
VEN); Killip 37470 fr (NY, VEN); Maguire & Maguire 29000 fl (NY, US); 29005 fr (NY, US);
Maguire, Steyermark& Maguire 46981 fr (F, NY); 53747 fr (F, NY, US); Steyermark 75428 fr (F, NY,
VEN); 75562 fr (F, NY, VEN); 88ro8 fr (NY); Steyermark& Nilsson 260 fr (NY, US); 484 fr (VEN);
Steyermark & Wurdack8 fl (NY, VEN); 1315 fr (VEN); Wurdack& Monachino40933 fl (F, GH, NY, US).
Terr. Amazonas: Croizat ro4oA fl (NY, US); Level 1o2 fl (NY, US); Maguire & Politi 27953 fr (GH,
NY, US); 28208 fr (F, MO, NY); Maguire,Cowan& Wurdack29487 fl (NY); Pannier& Schwabe1098 fl
(VEN); LI. Williams I4707 fl (A, F, US, VEN); 15227 fr (F, US); 15939 fr (F); Wurdack& Adderley
42769 fl (GH, NY, US); 43573 fl (F, MO, NY, US). TRINIDAD & TOBAGO. Baker & Simmonds
14919 fl (K, TRIN); Broadway4070 fl (NY, P, S, U); 4466 fr (BM, F, GH, K, MO, NY, U, US);
6445 fl (BM, F, K); 6634 fr (BM, FHO, K, MO, S, US); Brooks12484 fr (FHO, K, TRIN); Britton,
Hazen & Mendelson1811 fl (GH, K, NY); Eggers 113 fl (P); 1439 fl (TRIN, S); 1458 fl (K); 5842 fl
(BR, LE, US); Lodgesn fl (K); Marshall 11930 fl (FHO, K, TRIN); 12688 fr (FHO, K, NY, TRIN);
Mckaysn (FHO); Prance2109 fl (NY); Swabey& Dean 13139 fr (TRIN); R. O. Williams 11138 fr (K).
GUYANA. Anderson39 fl (K, NY); Cowan39392 fl (F, K, NY, P, RB, US); De la Cruz 2542 fl (F, GH,
MO, NY, US); 3829 st (F, GH, MO, NY, US); 4411 fl fr (F, K, MICH, NY, US); 4419 fl (F, GH, K,
MICH, NY, US); 4524 fl (F, GH, MO, NY, US); Fanshawe607 (F.D. 3343) fl (K, NY, U); 1420
(F.D. 4156) fl (K, NY, U); 2469 (F.D. 5205) fr (A, K, NY, U, US); 2866 (F.D. 6013) fl (K, NY);
Jenman3917 fl (K, NY); Persaud94 fl (F, K, NY, S, US); Schomburgk 908 fl (K, NY, P); 986 fl (K, NY,
P, W); 1357 fl (K, P); Sandwith604 fl (K, LE, NY, P, RB, S, U, US); Tillett & Tillett 45578 fr (GH,
NY, US). SURINAME. B.W. 479 fr (U); 1538 fr (U); 1578 fr (A, U); 1970 fl (U); 2257 fl (A, U);
2312 fl (U); 2604 fr (U); 3396 fr (IAN, K, MO, U); 5659 fl (U); 6i80 fl (IAN, K, U); 6693 fl (A,
MO, U); Geyskesi8 fl (NY); Heyligers 164 fl (U); Hostmann1038 fl (BM, K, LE, P); Irwin, Prance,
Soderstrom& Holmgren55425 fl (F, MO, NY, UB, US); Lanjouw1238 fl (U); Lanjouw& Lindeman916 fr
(IAN, K, NY); Lindeman6870 st (NY, U); Stahel 41 fl (A, IAN, K, NY, U); Tullehen483 fl (U); van
Donselaar2668 fr (NY, U). FRENCH GUIANA. Benoist 1498 fl (P); For. Serv. 22M fl (P, U); 1123 fl
(U); 7526 fl (NY, P, U); 7987 fl (MG, U); Leprieursn fl (BM, K, P, US); Maguire,Pires & Maguire
47052 fl (F, NY); Martin sn fl (BM, K, US); Melinonsn fl fr (A, BR, NY, P); 339 fl (P); Sagot 843 fl
(BM, K, P, S); 974 fl (P); Wachenheim sn fl (BM, K, US); 208 fl (P); 235 fl (P); 253 fl (P). BRAZIL.
Amazonas: Black 48-2701 fr (NY); 48-2902 fl (IAN, NY); ChagasINPA 201 fr (INPA, NY); Ducke
580 fl (F, IAN, MG, MO, NY, R, US); 2149 fl (IAN, NY, P, R); MG 7111 fr (BM, MG, RB); Frdes
22799 st (IAN, NY); 23274 fl (NY); 23929 fr (IAN, UB); 25465 fl (IAN, NY, RB); 33223 fl (IAN);
Krukoff4796 fl (A, BM, F, K, MICH, MO, NY, S, US); 6489 fr (A, BM, BR, F, K, LE, MICH,
MO, NY, RB, S); 6898 fl (A, BM, BR, K, LE, MICH, NY, RB); 6913 fl (A, K, MO, NY, S); 7028
fr (A, BM, BR, F, K, LE, MICH, MO, NY, RB, S, US); 8646 fl (A, BM, BR, F, K, LE, NY, P, S);
8962 fl (A, NY, S); Pranceet al 2596 fl (INPA, NY); 2617 fl (INPA, NY); 3081 fl (INPA, NY); 3099
fl (INPA, NY); 3615 fl (INPA, NY); 365I fl (INPA, NY); 3728 fr (INPA, NY); 8215 fl (INPA, NY);
Riedel 1304 fl (US); Rodrigueset al 2227 fl (INPA 8637); 2536 fr (INPA 9070); 2948 st (INPA 10313);
4776 fl (INPA 13303, NY); 4827 fl (INPA 13360, NY); 5003 fl (INPA 13664; NY); 7123 fl (INPA
15785); 7357 fr (INPA I6749); 7700 fr (INPA 17005); Schultes& Ldpez9280 fl (IAN, NY, US); 9948
fl (US); Spruce1471 fr (BM, K, LE, NY, P); 1649 fl (BM, GH, GOET, K, LE, OXF, P); Traill 210
fl (K); Ule 8913 fl (K, MG), Terr. Roraima: Pranceet al 9762 fl (INPA, NY). ParA: Black & Ledoux
50-10723 fr (IAN, NY); Cavalcante1497 fr (MG); DuckeMG 8514 fl (BM, MG, RB); 86o0 fl (BM, MG,
RB); 8651 fl (BM, RB); 9087 fl (BM, RB); 11708 fr (BM, MG, RB); RB i88o0 fl (RB, S); 19770 fl
(K); 19786 fl (K, RB); 19789 fl (K, RB); Frdes20415 fr (F, IAN, K, NY, US); 34618 fl (IAN); Guedes
30 fl (IAN, NY); Mexia 5996 fl (BM, F, GH, K, LA, MICH, MO, NY, S, U, US); Oliveira II8 fr
(IAN); 272 fr (IAN); Pires 51821 fr (COL, K, LE, NY); Pires & Silva 4638 fl (IAN, NY); Prance&
Silva 58959 fl (NY, UB, US); Prance & Pennington1513 fl (F, GH, NY); 1882 fr (F, GH, NY); 1951 fr
(F, GH, IAN, NY); 2033 fr (F, GH, NY); 2077 fr (F, GH, NY); R. S. Rodrigues19776 fl (K). Terr.
AmapA: Black 49-8500 fl (IAN, NY); Froes25745 fl (IAN, NY, RB); 25958 fl (IAN, LE); Cowan38212
fl (NY, US); Ducke MG 4829 fl (BM, RB); RB 15167 fl (K, MG 4766); Irwin, Egler & Pires 47I47 fl
(MG, NY); 47I95 fl (F, MG, NY); Irwin & Westra47184 fr (F, MG, NY, UB, US); Pires, Rodrigues&
Irvine50690 fr (F, MG, NY, US); 51044 fl (F, GH, IAN, MG, NY, US); 51400 fl (MG, NY); 51407 fr
(F, INPA, MG, MO, NY, UB, US); 51418 fr (F, IAN, MG, NY, US); Pires & Cavalcante52459 fl
(IAN, MG, NY, US); 52565 fr (F, IAN, MG, NY, US). Maranhao: Froes 1915 fl (A, BI i-, F, K,
MICH, MO, NY, P, S, US); 1976 fr 'A, BM, F, K, MICH, MO, NY, P, S, US). Acre: Krukoff5252 fl
(A, BM, F, K, MICH, MO, NY, RB, S, US); 5379 fl (A, BM, F, LE, MICH, MO, NY, US); Prance
et al 7260 fr (INPA, NY). Terr. Rond6nia: Blacket al 52-14636 fl (NY); Pranceet al 6965 fr (INPA, NY);
8803 fr (INPA, NY); 8996 fr (INPA, NY). Mato Grosso: Lima 58-3157 fl (IPA, RB). Guanabara:
108 Flora Neotropica

Carcerelli62 fl (RB 47114); 76 fl (NY, RB 78244); Glaziou9789 fl (P); 13711 fl (R); 13671 fl (P); Goes&
Dionisio 790 fl (RB).

LOCALNAMES.Venezuela: Oroy-uara-yek (Arekuna), Guayabo.Trinidad: Cofee-

wood, Wild cocoa.Guyana: Cococow,Kairiballi,Buruburuli.Suriname: Anaura,Kriki-
Kairiballi,Japopare,Ingibarki.Brazil: Caripe-rana,
mauroeroe, Macucu,Macucu-fnfo.

69b. Licania heteromorpha var glabra (Martius ex Hooker f.) Prance, stat
nov
Licaniaglabra Martius ex Hooker f., Mart. Fl. Bras. I4(2): lo. 1867.
LicaniacostataSpruce ex Hooker f., Mart. Fl. Bras. 14(2): Io. 1867, nom nud in syn.

Leaves oblong to oblong-lanceolate, cuneate at base, reticulation lax; fruit

elongate, costate.
TYPES.Spruce3503, Venezuela, Terr. Amazonas, fr (lectotype, K; isotypes, NY,
P); Martiussn (Jan. 1820), Brazil, Amazonas, fl (paratype, M); sn (Feb. 1820) fl
(paratype, M); Spruce2197, Brazil, Amazonas, fr (paratypes, BM, CGE, GOET, K,
LD, LE, M, NY, OXF, P); 3023 fl (paratypes, BR, K, P).
DISTRIBUTION.Riverine forest mainly in western Amazonia, but scattered
throughout the basin.
COLOMBIA. Vaup6s: Garcia-Barriga14212 fr (COL, US); Schultes& Cabrera16615 fr (F, GH
NY, US); 17025 fr (COL, GH, NY, US); 19768 fl (GH, NY, US); Schultes& Ldpez9472 fr (F, GH
IAN, US). Amazonas: Schultes3977 fr (COL, F, US); Schultes& Lpez 9246fl (GH, US). VENEZUELA
Bolivar: Wurdack& Monachino39896 fr (F, FHO, GH, NY, UB, US). Terr. Amazonas: Farinas et al
658 fr (NY); Maguire, Cowan & Wurdack32052 fr (FHO, NY, US); Maguire & Wurdack34862 fr
(F, IAN, K, NY); 34986 fr (NY, RB); Wurdack& Adderley42731 fr (FHO, NY, US); Schwabesn fl
(VEN); Ll. Williams13024 st (K, VEN); 15602 fr (VEN). PERU. Loreto: Schunke346 fl (A, NY, US);
Tessmansn fl (NY). BRAZIL. Amazonas: Baldwin 3453 fl fr (F, NY, S, US); Black 48-2788 fr (IAN,
NY); DuckeRB 25033 fr (K, P, RB, US); Frdes22240 fr (IAN, NY, U); 22709 fl (NY); Krukoff4744
fr (BM, F, K, MICH, MO, NY, S, US); 4846 fr (A, BM, F, K, LE, MICH, MO, NY, S); 7271 fr
(A, BM, BR, LE, MO, NY, S, U, US); Pires 458 fr (IAN, NY); 652 fl (IAN); Schultes& Ldpez8895
fl (GH, IAN, K, US); Ule 6i 8 fl (K, MG 5996). Terr. AmapA: Irwin,Pires & Westra4836oa fr (F, NY);
47862 fr (F, MG, NY). Terr. Rond6nia: Pranceet al 6838 fr (INPA, NY).

LOCALNAME.Colombia: komkow.
In the original description Hooker cites Spruce's numbers I472 and I649 as
L. glabra.However, he also cites these same numbers as L. heteromorpha. This confusion
is indicative of the close relationship of these species. Both numbers are assigned to
var heteromorpha as defined here.

69c. Licania heteromorpha var subcordata Fritsch, Ann. Naturh. Mus.

Wien 4: 45. 1889.
Licaniasubcordata
Fritsch, Osterr. Bot. Zeitschr. 44: i8. 1894.

Leaves ovate, subcordate at base, the reticulation intricate; fruit globose, smooth.
TYPES. Pohl 4402, Brazil, Amazonas, fl (lectotype, W); Martius sn fr (paratype,
M).
Riverine forest in Amazonian Brazil.
DISTRIBUTION.
BRAZIL. Amazonas: Frdes25148 fr (IAN, NY, RB); 25455 fr (IAN, NY, RB).
Licania 109

69d. Licnaia heteromorpha var perplexans Sandwith, Kew Bull. 1931:

37I. I93'.

Leaves ovate-elliptic, rounded to subcuneate at base, the reticulation lax; fruit

globose, smooth.
TYPES.Schomburgk 824, Guyana, fl (holotype, K; isotype, NY); 1443 fl (para-
type, K); Jenman3902 fl (paratypes, K, U); 7336 fl (paratypes, F, K); Hohenkerk
F.D. 39B fl (paratype, K); SandwithI40 fr (paratypes, K, P, S, US); 433 fr (paratypes,
K, NY, RB, U).
DISTRIBUTION. Confined to the forests of Guyana.
GUYANA. Boyan75 (F.D. 7759) fl (K, NY, U); Fanshawe1047 (F.D. 3783 fl (K, NY); Davis 383
(F.D. 2379) fl (FHO, K, U); Hohenkerk139A fr (K).

LOCAL NAMES
Kairiballi, Kautaballiballi.
Several authors, notably Fritsch (1889) and Sandwith (193i), have commented
on the amazing variability within this species, especially with regard to leaf and fruit
morphology. This variation is far more extensive than in most other species of Licania.
However, there is remarkably little correlation among the different variables. Thus,
when several methods were used to examine the variation, none revealed any really
distinct segregates. This species is widely distributed geographically and ecologically,
and fortunately there is a large amount of herbarium material to work with. Analysis
of the material available revealed four distinct varieties, each of which has rather
restrictedgeographical distributionand may be definedby a numberof smalldifferential
characters. These varieties correspond to former species or varieties. Variety glabra
is most distinct at the extreme of its variation where the fruit is deeply costate and the
leaves are narrowly oblong-lanceolate. However, it shows complete gradation into
var. heteromorpha. Much flowering material lacking fruits of var glabrais almost in-
distinguishable from var. heteromorpha. The deeply costate fruit of var glabra is an
important character for specific delimitation. However, within L. heteromorpha there
is a complete range in this character from a terete unridged globose fruit to a deeply
costate oblong one. The costate var glabraoccurs in Amazonian Brazil, Venezuela
and Colombia. Outside this range there are occasional specimens with costate fruits
in var heteromorpha, as in Steyermark 87282 from Delta Amacuro, Venezuela, and
Wurdack& Monachino41363 from the State of Apure, Venezuela. These two collec-
tions have slightly costate fruit, but much less than in typical variety glabra. The
variation in the various leaf characters is similarly ill-defined. The fruit exterior may
be velutinous or verrucose and glabrous. In the other species fruit indumentum is
usually constant.
Often one can pick out two collections of L. heteromorpha and conclude that they
represent two separate species. However, study of all cited material has revealed
extraordinaryphenotypic plasticity of this appropriately named species, suggesting a
free exchange of genetic material across its entire range. Experimental study is called
for, hopefully to shed further light on the genetic mechanisms of variability. Licania
heteromorpha is a good example of an ochlospecies, as described by White (I962).
Licaniaprismatocarpa, described by Hooker (i867), has always been placed among
the dubious species because of inadequate type material. Recently, topotype material
of L. prismatocarpa has been collected (Schultes& Lopez 9280). This matches and
supplements the type well, and taken alone, appears to be rather distinct from L.
heteromorpha. However, as with the other synonyms given under L. heteromorpha, L.
prismatocarpa is linked to the total picture by a complete range of intermediates and
cannot be regarded as a taxon separate from L. heteromorpha var heteromorpha.
There is a type of gall which appears to be very frequent on specimens of L.
110 Flora Neotropica

16

i,,,.,,i, '"',,

."'.... .....

FIG 17. Geographic distribution and variation in Licania heteromorpha.? L. heteromorpha var
heteromorpha; var glabra; A L. heteromorpha
0 L. heteromorpha var subcordata.I, (Steyermark87282); 2,
(Steyermark 87409); 3, (Wurdacket al 41363); 4, (Wurdacket al 43573); 5, (Steyermark 90519); 6, (Broad-
way 4466); 7, (B.W. 6796); 8, (Pires et al 51418); Io, (Irwin et al 47147); ii, (Frdes20415); 12, (Spruce
3503); I3, (Krukoff7271); 14, (Schulteset al 17025); 15, (Irwin et al 48360a); I6, (Frdes25455), leaves
x o. 15; fruits x 0.45.

heteromorpha,occuring in most varieties. Galled material has been found throughout

the range of the species (eg Persaud 94, Guyana; Sagot 843, French Guiana; Krukoff
5279, Brazil-Acre; Steyermark87409, Venezuela-Delta Amacuro; Velez 2275, Vene-
zuela-Apure).
Benoist (Bull. Mus. Nat. Hist. Paris 25: 513. i919.) based his variety grandifolia
on the larger size of the leaves of a single specimen (Martin sn). Since there is so much
variation in leaf size throughout the range of L. heteromorpha,and since large leaved
specimens are frequent, var grandifolia is considered as part of var heteromorpha.

70. Licania glazioviana Warming, Kjoeb. Vidensk. Meddel. 1874: 68. 1874;
Fritsch, Ann. Naturh. Mus. Wien 4: 48. I889.

Tree? Young branches glabrous and conspicuously lenticellate. Leaves oblong-

lanceolate to narrowly elliptic, membraneous, 3.0-6.5 cm long, I.4-2.5 cm broad,
bluntly acuminate or obtuse at apex, with acumen to 5.0 mm long, cuneate at base,
glabrous on both surfaces; palisade glands present; midrib prominulous above,
glabrous; primary veins 1o-12 pairs, prominulous on both surfaces, but rather in-
distinct; venation papillose on lower surface; petioles 2.0-4.0 mm long, glabrous,
canaliculate, eglandular. Stipules lanceolate, I.5-2.5 mm long, glabrous, adnate to
petiole base, persistent. Inflorescences short axillary and terminal racemose panicles,
Licania 111

the rachis and branches almost glabrous. Flowers ca 3.5 mm long, solitary and sessile
on primary branches of inflorescence. Bracts and bracteoles 0.5-I.O mm long, ovate,
persistent, almost glabrous. Receptacle urceolate, sessile, tomentellous on exterior,
tomentose within. Calyx lobes acute, tomentellous on both surfaces. Petals absent.
Stamens 5, unilateral, with fused sterile portion of ring opposite to them; filaments
connate at base only, shorter than calyx lobes, pubescent. Ovary inserted at base of
receptacle, villous. Style pubescent throughout, equalling filaments. Young fruit
pyriform; epicarp with a waxy indumentum; pericarp thin, hard, tomentellous with-
in.
TYPE.Glaziou2561, Brazil, Guanabara, fl (holotype, C; isotypes, BR, K, P).
Known only from the coastal restingas of Guanabara, Brazil,
DISTRIBUTION.
BRAZIL. Guanabara: Duarte 4839 fr (NY, RB 11052).

Licaniaglaziovianais quite distinct from related species in the thin membraneous

leaves with cuneate bases, and in the urceolate receptacle.

71. Licania littoralis Warming, Kjoeb. Vidensk. Meddel. 1874: 67. I874;
Fritsch, Ann. Naturh. Mus. Wien 4: 47. I889.

Tree or shrub, the young branches puberulous, becoming glabrous and lenticel-
late with age. Leaves obovate to ovate-elliptic, thick-coriaceous, 3.0-I .o cm long,
1.2-6.5 cm broad, retuse to obtuse at apex, rounded to cordate or rarely cuneate at
base, glabrous on both surfaces; palisade glands occasional; midrib prominulous
above, glabrous; primary veins 7-9 pairs, prominent beneath, almost plane above;
petioles 2.0-3.0 mm long, glabrous, rugose, terete, eglandular. Stipules lanceolate,
I.o mm long, adnate to petiole base, subpersistent. Inflorescences terminal and sub-
terminal racemose panicles, the rachis and branches puberulous. Flowers ca 3.0 mm
long, solitary or in small groups on primary branches of inflorescence. Bracts and
bracteoles 0.5-I.0 mm long, persistent, ovate, entire, tomentellous on exterior.
Receptacle campanulate, sessile, tomentellous on exterior, tomentose within. Calyx
lobes acute, tomentellous on both surfaces. Petals absent. Stamens 4, unilateral;
filaments shorter than calyx lobes, free to base, glabrous. Ovary inserted at base of
receptacle, short-tomentose. Style pubescent throughout, equalling filaments. Fruit
pyriform, to 3.0 cm long; epicarp smooth with a waxy indumentum; pericarp thin,
hard, fibrous, tomentose within.

Key to the Varieties of Licanialittoralis

i. Leaf basesroundedto cordate. a. var littoralis.
I. Leaf bases cuneate. b. var cuneata.

71a. Licania littoralis var littoralis

LicanialittoralisWarming, Kjoeb. Vidensk. Meddel. I874: 67. I874.

Leaf bases rounded to cordate.

112 Flora Neotropica

TYPE.Glaziou6i68, Brazil, Guanabara, fl (holotype, C; isotypes, F, K, P).

DISTRIBUTION. Confined to the coastal restingas and coastal forest of eastern
Brazil from Paraiba to Guanabara.
BRAZIL. Paraiba: Lima 60-3434 fl (IPA); Mattos9709 fl (SP); Santos1092 fl (R). Pernambuco:
Lima 49-209 fl (IAN, IPA, NY); 49-290 fr (IAN, IPA, NY); 52-1200 fl (IAN, IPA); 55-2072 fr
(IPA). Guanabara: MachadoRB 75836 fr (RB), topotype.

LOCALNAMES.Oiti, Disbota.

7ib. Licania littoralis var cuneata Kuhlmann, Ann. Prim. Reun. Sul-Am.
Bot. 1(3): 77. I940.
Leaf bases cuneate.
TYPE.Kuhlmann 208, Brazil, Espirito Santo, fl (holotype, RB34I58; isotype, NY).
DISTRIBUTION. Known only from the type gathering from coastal forest of
Espirito Santo in Brazil.

72. Licania fanshawei Prance, sp nov Fig 16 A-D.

Arbor magna, remulis juvenilibus puberulis mox glabris lenticellatis. Folia

alternata petiolata; petiolo 3.0-5.0 mm longo, glabro, rugoso, leviter canaliculato,
eglanduloso; laminae ovato-ellipticae, coriaceae, 4.0-I I. cm longae, 2.5-7.0 cm
latae, basi rotundatae, apice obtusae, utrinque glabrae, subtus glandulosae, saepe
pustulatae, prope petiolum cum glandulis duobus munitae; costa media supra leviter
prominente, glabra; costis secundariis 7-8 jugis, utrinque leviter prominentibus
subtus saepe pustulatis. Stipulae ad 2.0 mm longae, caducae, intrapetiolares. Flores
circa 2.0 mm longi, haud pedunculati, in paniculis terminalibus subterminalibusque
dispositi, rachi ramisque fulvo-tomentellis vel puberulis. Bracteae bracteolaeque
0.3-I.O mm longae, triangulares, persistentes, extus puberulae. Receptaculum
campanulatum, sessile, extus fulvo-tomentellum, intus tomentosum. Petala nulla.
Stamina 3, unilateralia, filamenta glabra, ad basim libera, calycis-lobis breviora.
Ovarium ad basim receptaculi insertum, villosum. Stylus e basi ovarii ortus,
cum filamentis aequans, ad apicem pubescens. Fructus mihi ignoti.
TYPE.Steyermark 87557, Venezuela, Terr. Delta Amacuro, Rio Cuyubini, Cerro
La Palma, fl (holotype, NY).
DISTRIBUTION.
Primary forest of Guyana and the Orinoco delta.
GUYANA. Moraballi Creek: Fanshawe265 (F.D. 3001) fl (BM, K, NY, P, S, U); Barabara
Creek, Lower Essequibo River, Fanshawe944 (F.D. 3680) fl (K, NY); Mazaruni Station, Fanshawe
650 (F.D. 3386) fl (K, NY); Issano road, Fanshawe3516 (F.D. 7112) fl (K, NY, P, S, U).

LOCALNAME.Guyana: Kautaballiballi.
This species is nearest to L. irwinii,but differsin the frequently papillose venation,
the pubescence of the flowers and inflorescence, and the fewer stamens. It differs
in the rounded leaves with blunt apices.
from L. cyathodes
Licania 113

73. Licania irwinii Prance, sp nov Fig i8 A-E.

Arbor, ramulisjuvenilibus glabris, lenticellatis. Folia alternata, petiolata; petiolo

2.0-4.0 mm longo, glabro, eglanduloso, leviter canaliculato; laminae ovato-orbicu-
lares vel ellipticae, crasse coriaceae, margine recurvatae, 2.0-6.5 cm longae, 1.2-4.0
cm latae, basi rotundatae, apice rotundatae vel obtuso-apiculatae, utrinque glabrae,
subtus glandulosae haud pustulatae; costa media supra glabra, leviter prominente;
costis secundariis 6-I2 jugis, subtus leviter prominentibus, supra planis. Stipulae
parvae, crassae, ad 1.5 mm longum, subpersistentes,cum basi petioli adnatae. Flores
2.5-3.0 mm longi, haud pedunculati, in paniculis terminalibus subterminalibusque
dispositi, rachi glabrescente, ramis puberulis. Bracteae bracteolaeque ad i.o mm
longae, triangulares, persistentes, extus puberulae. Receptaculum campanulatum,
sessile, extus sordido-tomentellum, intus tomentosum. Calycis-lobi acuti, utrinque
tomentelli. Petala nulla. Stamina 5, unilateralia, filamenta glabra, calycis-lobis
breviora. Ovarium ad basim receptaculi insertum, villosum. Stylus e basi ovarii
ortus, calycis-lobis breviora, ad apicem pubescens. Drupa pyriformis cum stipite'
I2.0 mm longo, 3.5 cm longa, extus sordido-pulverulenta, leviter rugosa in sicco;
mesocarpio tenue, carnoso; endocarpio duro, 2.5 mm lato, lignoso, intus sparse
pubescente.
TYPE. Irwin, Prance, Soderstrom & Holmgren 55002, Suriname, Wilhelmina
Mountains, fl (holotype, NY).
Forested slopes of Suriname and French Guiana.
DISTRIBUTION.
SURINAME. Moengo, Lanjouw& Lindeman454 fl (K, NY, U). FRENCH GUIANA. Aubreville
278 fr (P).

This species is distinct from others with glabrous leaves by the rounded leaf apex
but
and the long stipe of the fruit. It is closely related to species of section Pulverulenta
differs in the glabrous leaves as well as the long stipe of the fruit.

74. Licania cyathodes R. Benoist, Bull. Mus. Hist. Nat. Paris 25: 513. 1919.

Small tree, the young branches glabrous and lenticellate. Leaves oblong to
oblong-lanceolate, coriaceous, 5.0-9.0 cm long, I.8-4.5 cm broad, acuminate at
apex, with acumen 6.0-I2.0 mm long, subcuneate at base, glabrous on both surfaces;
palisade glands occasional; midrib slightly impressed above, glabrous; primary veins
7-10 pairs, prominent beneath, more or less plane and inconspicuous above; petioles
3.0-5.0 mm long, glabrous, eglandular, canaliculate; stipules lanceolate, 3.0-4.0 mm
long, glabrous, coriaceous, persistent, adnate to base of petiole. Inflorescences short
terminal and subterminal racemose panicles, the rachis puberulous, with branches
puberulous to tomentellous. Flowers 2.0-2.5 mm long, in small groups on very short
secondary inflorescence branches. Bracts and bracteoles ovate, 0.5-2.0 mm long,
persistent, tomentose on exterior, glabrous within, entire. Receptacle campanulate,
tomentose on exterior, tomentose within; pedicels 0.25-0.50 mm long. Calyx lobes
acute, tomentellous on both surfaces.Petals absent. Stamens 5-7, unilateral; filaments
shorter than calyx lobes, free to base, glabrous. Ovary inserted at base of receptacle,
villous-tomentose. Style pubescent to apex, equalling filaments. Fruit unknown.
TYPES.Sagotsn, French Guiana, fl (lectotype, P); Benoist778, French Guiana,
fl (paratype, P); Martinsn, French Guiana, fl (paratypes, BM, K, NY, P).
Collected only in the riverine forests of French Guiana.
DISTRIBUTION.
Ir4, fl (BM, F, GH, K, NY, P, US).
FRENCH GUIANA. Wachenheim
114 Flora Neotropica

GII '; w \ 0 v

.....~-:?' v,: ?~, ,," _',,.,'"*',. ......'............

\\ ~ :-..:,..'",
~~~~ "-1, ',r ,
? ?
!:i' ~.iI ~ ~ ~

FI 8.Seieso 7) ,hbt .;B

l50,Abdil (Irwne iai.A-,Lrii
flwr ;C lwrscin
u,
x ;D ri,xz E ri eto,xz -,Lmrsa(uk
RB
25032, Nlaguire et al 56066); F, ~;
habit,
x G, flower, x
~ 5; H, x z 5; J, fruit,
flower, x 8. K-M,~~~~~~~~r
L. ta598;K
(Pac
si l ai,xo5
a ,foe,x~;M lwrscin o

FIG i8. Species of Licania.A-E, L. irwinii (Irwin et al 5002, Aubre'ville278); A, habit, x 0.5; B,
flower bud, x I5; C, flower section, x I5; D, fruit, x I; E, fruit section, x i. F-J, L. impressa(Ducke
RB 25032, Maguireet al 56066); F, habit, x I; G, flower, x I5; H, flower, x I5; J, fruit, x 8. K-M,
L. silvae (Pranceet al 58968); K, habit, x 0.5; L, flower, x io; M, flower section, x o.
Licania 115

75. Licania polita Spruce ex Hooker f., Mart. Fl. Bras. 14(2): I7, t.4. I867;
Fritsch, Ann. Naturh. Mus. Wien 4: 49. I889.
LicaniapoeppigiiFritsch, Ann. Naturh. Mus. Wien 4: 49. i889, synon nov. Type. Poeppig2785,
Brazil, Amazonas, fl (holotype, W; isotypes, BR, G, GOET).
LicanialaurifoliaHuber, Bol. Mus. Emilio Goeldi 5: 365. 1909, synon nov. Types. DuckeMG 7958,
Brazil, Pard, fl (holotype, MG; isotype, BM); 9052 fr (paratype, MG).
Licanialaxa Fanshawe & Maguire, Bull. Torrey Club 75: 321. I948, synon nov. Type. Fanshawe
1665 (F.D. 4401), Guyana, fl (holotype, NY; isotype, K).

Medium to large-sized tree, the young branches puberulous, becoming glabrous

with age. Leaves ovate to elliptic, thick-coriaceous, 7.o-I8.o cm long, 3.o-1.o0 cm
broad, acute to acuminate at apex, with acumen to Io.o mm long, subcordate or
rounded to subcuneate at base, glabrous on both surfaces; palisade glands present;
venation often minutely papillose on lower surface; midrib plane to prominulous
above, glabrous; primary veins 6-Io pairs, prominent beneath, prominulous above;
petioles 6.0-14.0 mm long, terete, rugose, eglandular, short-velutinouswhen young.
Stipules lanceolate, 4.0-7.0 mm long, coriaceous, glabrous, persistent to subpersistent,
adnate to base of petioles. Inflorescences terminal racemose panicles, the rachis and
branches puberulous to short-tomentellous. Flowers 2.5-3.5 mm long, densely
crowded in small groups on primary branches of inflorescence. Bracts and bracteoles
lanceolate to elliptic, often equalling flowers, puberulous on both surfaces,persistent,
entire. Receptacle campanulate, sessile, short-velutinous-tomentose on exterior,
tomentose within. Calyx lobes acute, tomentellous on both surfaces. Petals absent.
Stamens 6-7, unilateral; filaments shorter than calyx lobes, free to base, sparsely
pubescent. Ovary inserted at base of receptacle on short gynophore, densely villous.
Style equalling calyx lobes, pubescent throughout. Fruit pyriform; epicarp ferrugin-
eous-tomentose; pericarp thin, undifferentiated, fragile, granular, fibrous, sparsely
hirsute within.
TYPE. Spruce2676, Brazil, Amazonas, fl (holotype, K; isotypes, BM, BR, C,
CGE, F, GH, GOET, LD, LE, NY, OXF, P).
DISTRIBUTION. Non-flooded and periodically flooded forest in the Guianas and
Amazonian Brazil.
VENEZUELA. Bolivar: Cardona2203 fr (US, VEN); Ll. Williams iI306 fr (US, VEN). Terr.
Amazonas: LI. Williams 15680 fr (US, VEN). SURINAME. B.W. 3817 fr (NY, U); Stahel 257 st
(A, U). BRAZIL. Amazonas: Fr6es22510 fl (IAN); Krukoff6668fl (A, BM, BR, F, IAN, K, LE, MICH,
MO, NY, RB, S, US); 8138 fl (A, BM, F, K, LE, MO, NY, S); 8276 fl (A, BM, BR, F, K, LE, MO,
NY, P, RB, S, US); 8507 fl (A, BM, BR, F, K, LE, MO, NY, P, S, US); Kuhlmann1044 fl (K, RB
17970); Spruce2762 fl (BM, BR, F, GOET, LD, LE, OXF, P). Para: DuckeMG 15019 fl (BM, G, NY,
RB 15150); RB i88i6 fl (K, NY). Mato Grosso: Kuhlmann304 fl (R).

LOCALNAME.Venezuela: Peruz(Arekuna).
Licaniapolita differs from related species in the large flowers and stipules as well
as the large, thick-coriaceous leaves. Specimens documenting the three names re-
duced to synonymy here do not differ in any significant features. L. polita displays a
wide range of variation in leaf shape and size, which has led to the proposal of the
names cited as synonyms.

76. Licania silvae Prance, sp nov Fig 18 K-M.

Arbor, ramulis juvenilibus sparse puberulis mox glabris tomentellis. Folia

alternata petiolata; petiolo 2.o-4.0 mm longo, glabro, rugoso, tereti, supra sub lamina
glandulis duobus munito; laminae oblongae vel oblongo-ellipticae, coriaceae,
116 Flora Neotropica

6.0-I 1.5 cm longae, 2.5-5.5 cm latae, basi rotundatae vel subcuneatae, apice acutae
vel in acuminem 2.0-6.0 mm longum contractae, utrinque glabrae, subtus venulis
pustulatis; costa media supra glabra, leviter prominente; costis secundariis 7-8 jugis,
utrinque leviter prominentibus. Stipulae minutae, lanceolatae, coriaceae, extra-
petiolares, subpersistentes.Flores circa I.5 mm longi, haud pedunculati, in paniculis
terminalibus subterminalibusquedispositi, rachi ramisque dense puberulis. Bracteae
bracteolaeque 0.2-0.6 mm longae, ovatae, persistentes, extus puberulae. Recepta-
culum campanulatum, sessile, extus tomentellum, intus tomentosum. Calycis-lobi
acuti, utrinque tomentelli. Petala nulla. Stamina 3, unilateralia, filamenta sparse
pubescentia, calycis-lobis breviora, ad basim libera. Ovarium ad basim receptaculi
insertum, tomentosum. Stylus e basi ovarii ortus, ad apicem villosus, calycis-lobos
aequans. Fructus mihi ignoti.
TYPE. Prance & Silva 58968, Brazil, Para, Belem-Brasilia road; 30 km south
of Gurupi, fl (holotype, NY; isotype, UB).
DISTRIBUTION. Non-flooded forest in Amazonian Brazil. Flowering August to
September.
BRAZIL. Amazonas: Rio Purus, Boca do Acre, Pranceet al 2534 fl (INPA, NY). Par4: Bel6m-
Brasilia highway, km Ioo, N. T. Silva 597 fl (IAN); 611 fl (IAN, UB).

Licaniasilvaelies near L. cyathodes, but differs in the prominent midrib of the leaf
upper surface, the terete petioles, the fewer stamens, the sessileflowers disposed on the
primary inflorescence branches, and the small caducous stipules. L. silvae is also
closely related to L. fanshawei,but differs in the acute leaf apex, the terete petioles,
the stipules adnate to the outside of the petioles, and in the pubescence of the in-
florescence and flowers.

7. Licania subgen Licania sect Cymosa

77. Licania densiflora Kleinhoonte, Rec. Trav. Bot. Neerl. 22: 383. 1925;
Kleinh. in Pulle, Fl. Suriname 2(I): 439. I939; Sandwith, Kew Bull.
1931. 371. 1931.
Licania kanukuensisStandley, Lloydia 2: I82. 1939; Maguire, Bull. Torrey Club 75: 320. I948.
Type. A. C. Smith3420, Guyana, fl (holotype, F; isotypes, A, IAN, K, LE, MO, NY, U,
US).

Tree to 30.0 m tall, the young branches tomentose, soon becoming glabrous and
lenticellate. Leaves oblong to elliptic, coriaceous, 5.0-I5.0 cm long, 2.0-7.5 cm broad,
acute to shortly acuminate at apex, with acumen 3.0-7.0 mm long, rounded to sub-
cuneate at base; midrib slightly impressed or plane above, glabrous; primary veins
9-II pairs, prominent beneath, plane above; petioles 3.o-Io.o mm long, terete,
lanate-tomentose, eglandular. Stipules linear, 2.0-7.0 mm long, pubescent, subper-
sistent, adnate to extreme base of petiole. Inflorescences densely flowered terminal
and subterminal panicles, the rachis and branches densely tomentose. Flowers 4.0-
5.0 mm long, in small groups of cymules attached to rachis and primary branches by
short thick peduncles. Bracts and bracteoles ca 2.0 mm long, persistent, enclosing
buds in small groups, tomentose on exterior. Receptacle campanulate, subsessile,
tomentose on exterior, tomentose within. Calyx lobes acute, tomentellous on both
Licania 117

surfaces. Petals absent. Stamens 5-7, unilateral; filaments shorter than calyx lobes,
glabrous. Ovary inserted at base of receptacle, villous. Style equalling filaments,
lanate for most of length. Fruit pyriform, with short stipe ca 5.0 mm long; epicarp
rufous-tomentose; pericarp thin, very hard, fibrous, densely hirsutulous within.
TYPES. B.W.5346, Suriname, fl (lectotype, U); B.W.2039 fl (paratype, U);
Pulle 452 st (paratypes, NY, U).
DISTRIBUTION. Primary forest on high ground and slopes in the Guianas and
adjacent Venezuela and Brazil. Flowering March to November.
VENEZUELA. Terr. Delta Amacuro: Berti 137 st (NY); Blanco 197 fr (VEN); 254 fr (VEN);
Little 6000ofr (VEN); 17674 fr (VEN); Steyermark87592 fr (F, NY); Wurdack339 fr (F, NY, US);
Zabala 26 fl (VEN); 65 fl (VEN); 69 fl (VEN). Bolivar: Bernardi2951 fl (VEN); 3032 fl (VEN);
6491 fl (VEN); Berti 284 st (VEN); Blanco33 fl (VEN); 294 fr (VEN); Maguire, Steyermark& Maguire
46754 fr (NY); Steyermark 86370 fr (VEN); 86644 fr (VEN); 86758 st (VEN); 86976 fr (VEN); 88030
st (F, NY); 89076 fr (VEN). Guyana. Anderson57 fl (BM, K, NY); Fanshawe671(F.D. 3407) fl (K, NY);
J. Boyan88(F.D. 7772) fl (K, NY, U); Jenman3604 fl (BM); Sandwith336 fl (K, P, RB); 341 fl (K, NY,
RB, U, US); A. C. Smith3563 fl (A, NY). SURINAME. Irwin, Prance,Soderstrom& Holmgren55434 fl
(F, GH, NY, US); Maguire24024 fl (A, F, GH, MO, NY, US). BRAZIL. Terr. Amapa: Irwin 48741 fr
(GH, NY).

LOCAL NAMES. Venezuela: Guaray, Hierrito, Merecurillo. Guyana: Buhurada

(Arawak), Marishiballi.
Licaniadensiflorais distinct from related species by its large bracteoles and the
inflorescence with the cymules on short thick peduncles.

78. Licania cuprea Sandwith, Kew Bull. I937: io8. I937.

Tree to 20.0 m tall, the young branches densely fulvous-tomentose, inconspi-

cuously lenticellate. Leaves elliptic, oblong to oblong-lanceolate, coriaceous, 3.5-
I6.o cm long, 1.8-6.5 cm broad, acuminate or acute at apex, with acumen to I2.0
mm long, rounded at base, glabrous on upper surface, the lower surface with deep
reticulation obscured by dense rufous-brown arachnoid-tomentose pubescence;
midrib impressed above, tomentellous on lower portion; primary veins 7-10 pairs,
prominent beneath, slightly impressed toward base of upper surface; petioles 5.0-
8.o mm long, densely tomentose, shallowly canaliculate to terete, the glands obscure.
Stipules linear, persistent, ca 5.0 mm long, adnate to extreme base of petiole. In-
florescences terminal and axillary panicles, the rachis and branches densely rufous-
brown-tomentose.Flowers ca 2.0 mm long, solitary or in small groups on long slender
peduncles attached to primary branches of inflorescences. Bracts and bracteoles
I.0-2.5 mm long, lanceolate, tomentose, persistent. Receptacle campanulate,
tomentose on exterior, tomentose within; pedicels ca 0.25 mm long, Calyx lobes acute,
tomentose on both surfaces. Petals absent. Stamens 3, unilateral; filaments shorter
than calyx lobes, glabrous, free to base. Ovary inserted at base of receptacle, densely
villous. Style equalling filaments, pilose throughout. Fruit globose, with a long stipe
1.5-2.0 cm long; epicarp densely ferrugineous-velutinous-tomentose;pericarp thin,
fibrous, fragile, sparsely hirsutulous within.
TYPES.Jenman 6300, Guyana, fl (lectotype, K); Davis 426 (F.D.242I), Guyana,
fr (paratypes, FHO, K, U); AndersonF.D.456, Guyana, fr (paratype, K); Hohenkerk
fl
F.D.8oI, Guyana, (paratype, K).
DISTRIBUTION. Known only from Wallaba forest and savanna margins in Guyana.
Flowering October to February.
118 Flora Neotropica

GUYANA. Cowan39277 fr (GH, K, NY, P, US); De La Cruz 1776 fr (F, GH, MO, NY, US);
2648 fl (F, GH, MO, NY, US); 2865 fl (F, GH, NY, US); Fanshawei33(F.D. 2742) fr (FHO, S, U).

LOCALNAMES.Araudanni, Muri Kautaballi (Arawak), Unikiakia, Kunoko.

79. Licania impressa Prance, sp nov Fig 18 F-J.

Arbor magna vel media, ramulis juvenilibus puberulis haud lenticellatis. Folia
alternata petiolata; petiolo 7.0-I4.0 mm longo, primo tomentoso, canaliculato,
saepissime glanduloso; laminae oblongo-lanceolatae, coriaceae, 5.5-I3.0 cm longae,
1.8-4.0 cm latae, basi rotundatae vel subcuneatae, apice in acuminem 3.0-7.0 mm
longum contractae, supra glabrae, subtus dense lanato-arachnoideae pubescentes,
venulis profunde reticulatis vel cum cavis stomatalis; costa media supra glabra,
impressa; costis secundariis 9-I2 jugis, subtus prominentibus, supra planis vel leviter
impressis. Stipulae lanceolatae, coriaceae, subpersistentes,cum basi petioli adnatae.
Paniculacompositaterminalibus vel axillaribus, cymulis pluribus2 vel 3 floris breviter
pedunculatis constructa; rachi ramisque griseo-brunneis tomentosis; flores circa 1.5
mm longi. Bracteae bracteolaeque parvae, ovatae, persistentes, extus tomentosae.
Receptaculum campanulatum, extus tomentosum, intus tomentosum; pedicelli circa
0.25 mm longi. Calycis-lobi acuti, utrinque tomentosi. Petala nulla. Stamina 6-7,
in orbem completum disposita, filamenta glabra, ad basim libera, calycis-lobis
breviora. Ovarium ad basim receptaculi insertum, pilosum. Stylus e basi ovarii ortus,
calycis-lobis brevior, ad apicem pubescens. Fructus immaturus pyriformis, extus
sordido-tomentellus.
TYPE. DuckeRB25o32, Brazil, Amazonas, Manaus, fl (holotype, K; isotype, RB).
DISTRIBUTION. Non-flooded forest in eastern and central part of Brazilian
Amazonia.
BRAZIL. Amazonas: Manaus, Rodrigues7698 fl (INPA 17003). Para: Bel6m-Brasilia road, Km.
13, Maguire,Pires, Maguire & Silva 56066 fr (NY); Santar6m, M. Silva & Souza 2621 fr (NY, MG).

Licaniaimpressamay be distinguished from all other species of the genus with this
inflorescence type, by the oblong-lanceolate leaves with an impressed midrib, and
the long petioles.

80. Licania dealbata Hooker f., Mart. Fl. Bras. I4(2): 14. I867; Fritsch, Ann.
Naturh. Mus. Wien 4: 45. 1889.

Low suffrutex or shrub, the young branches puberulous, becoming glabrous

with age, obscurely lenticellate. Leaves oblong to oblong-ovate, coriaceous, 4.0-7.0
cm long, 2.0-3.5 cm broad, acuminate or acute at apex, with acumen to 5.0 mm
long, glabrous on upper surface, the lower surfacewith deep reticulation obscured by
dense lanate white pubescence; midrib plane or slightly impressed above, sparsely
pubescent towards base; primary veins 6-8 pairs, prominent beneath, more or less
plane above; petioles 2.0-5.0 mm long, pubescent, eglandular, terete. Stipules
lanceolate, 3.0-5.0 mm long, coriaceous, persistent, adnate to extreme base of petiole.
Inflorescences terminal and axillary panicles, the rachis and branches gray-puberu-
lous. Flowers ca 2.5 mm long, in few-flowered cymules attached to primary branches
by long slender peduncles. Bracts and bracteoles 0.2-I.0 mm long, ovate, persistent,
Licania 119

puberulous on exterior. Receptacle campanulate, gray-tomentellous on exterior,

tomentellous within; pedicels ca 0.25 mm long. Calyx lobes acute, tomentellous-
puberulous on both surfaces. Petals absent. Stamens 6-7, unilateral; filaments shorter
than calyx lobes, glabrous, free to base. Ovary inserted at base of receptacle, villous.
Style equalling filaments, sparsely pubescent throughout. Fruit pyriform, ca 2.0 cm
long; epicarp with short-appressed-velutinouspubescence; mesocarp thick, hard,
fibrous; endocarp thin, hard, fibrous, hirsutulouswithin.
TYPES.Gardner 2836, Brazil,Pernambuco,fl (lectotype, K; isolectotype,BM, CGE);
Martiussn, Brazil, Bahia, fl (paratype, M).
DISTRIBUTION.
Cerradosof the Planalto of Central Brazil. Flowering March to
June.
BRAZIL. Goias: Dawson 14812 fl (NY); Glaziou21113 fl (BR, K, LE, P, R, S); Irwin et al 14521
fl (NY); Rosa 57 fl (RB). Dist. Federal: Irwin & Soderstrom6070 fl (UB); Irwin et al 15332 fl (NY);
Pires, Silva & Souza 9199 fl (NY, UB); Prance & Silva 58995 fl (NY, US). Bahia: Irwin et al I4719 fl
(NY); 14720 fl (NY). Minas Gerais: Irwin et al 23881 fl (NY).

LOCALNAME.Manja-croite.
Licaniadealbatamay be distinguished from nearly related species by the deeply
reticulate leaf undersurface with white-lanate pubescence, the number of stamens,
and the low stature.

8i. Licania pallida Spruce ex Sagot emend, Ann. Sci. Nat. Bot. VI. 15: 306.
1883 pro parte quoad L. parvifloraBenth. var pallida tantum; Fritsch,
Osterr. Bot. Zeitschr. 42: 6. I892.
varpallida Hooker f., Mart. Fl. Bras. 14:(2): I8. I867; Fritsch, Ann. Naturh. Mus.
Licaniaparviflora
Wien 4: 51. I889.
Licaniapallida Spruce ex Hooker f., Mart. Fl. Bras. 14(2): I8. I867, nom nud, in syn.

Tree to 30.0 m tall, the young branchesglabrous and lenticellate. Leaves oblong-
elliptic, coriaceous, 5.0o-3.0 cm long, 2.0-6.5 cm broad, the apex with a finely
pointed acumen 6.0-i5.0 mm long, rounded-subcuneate at base, glabrous on upper
surface; lower surface with well developed stomatal cavities filled with lanate pubes-
cence, the reticulation glabrescent and cavities conspicuous; midrib plane above, or
slightly impressed toward base, glabrous; primary veins 7-0o pairs, prominent below,
more or less plane above; petioles 5.0-7.0 mm long, glabrescent, eglandular, rugose,
usually shallowly canaliculate. Stipules 2.0-3.0 mm long, linear, coriaceous, sub-
persistent, adnate to extreme base of petiole. Inflorescences terminal and axillary
branched panicles, the rachis and branches slender, puberulous. Flowers 1.5-2.0 mm
long, usually on long slender peduncles attached to primary branches of inflores-
cence, rarely almost sessile on branches of inflorescence. Bracts and bracteoles
persistent, 0.2-1.0 mm long, puberulous. Receptacle campanulate, tomentellous on
exterior, tomentellous to puberulous within; pedicels ca 0.25 mm long. Calyx lobes
acute, puberulous on both surfaces. Petals absent. Stamens 3-5, unilateral; filaments
shorter than calyx lobes, glabrous, free to base. Ovary inserted at base of receptacle,
tomentose. Style equalling filaments, sparsely pubescent throughout. Fruit oblong;
epicarp with short sordid reddish-brownpulverulent pubescence; pericarp thin, hard,
fibrous, sparsely hirsutulous within.
TYPES.Spruce1576,Brazil, Amazonas, fl (lectotype, P; isotypes,BR, CGE, F, K, LE,
M, NY, OXF); I635 fl (paratype, K).
DISTRIBUTION. Non-flooded forest in Venezuela and Brazilian Amazonia.
120 Flora Neotropica

VENEZUELA. Bolivar: Bernardi2658 fr (VEN); 2755 fl (FI, NY, VEN); Steyermark 90409 fl
(NY, VEN). Terr. Amazonas: Spruce3458 fr (K). BRAZIL. Amazonas: Aluisio r66 fl (INPA); Frdes
22671 fl (A, IAN, NY); 22549 fl (IAN, NY); 26390 fl (IAN, NY, UB); Krukoff6897 fr (F, K, MICH,
NY, RB, S, U, US); Pranceet al 3067 fl (INPA, NY); 10o46 fr (INPA, NY); Rodrigues5520 fl (INPA
14193, NY); 8319 fr (INPA, NY). Terr. Roraima: Pranceet al 4507 fr (INPA, NY); 4524 fr (INPA,
NY). Par,: Ducke RB 18825 fl (K, RB); Kuhlmann& Jimbo 95 fl (IAN); Pires 6981 fr (IAN); 7037
(IAN); Pires & Silva 10869 fl (IAN, NY); R. S. RodriguesMG 9650 fl (K, RB 19769). Terr. Rond6nia:
Pranceet al 6846 fl (INPA, NY).

LOCALNAME.Brazil: Macucu Sangue.

Licaniapallida is very closely related to L. parviflora,but differs in the more
slender inflorescence branches, the pedicellate flowers borne in small groups on
distinct peduncles, and the shorter sordid reddish-brown indumentum of the fruits.
The well developed stomatal cavities distinguish L. pallida from other species with
this type of inflorescence.
The nomenclature of this species has been much confused in the past, as the
specific epithet pallidahas been applied to three differentspecies of Licania.The earliest
mention of L. pallida Spruce was by Hooker who cited it in synonymy under L.
parvifloravar pallida, this leading to the later specific citation in IndexKewensis.Sagot
treated L. pallida as a species in 1883, taking it to include both L. parvifloravar pallida
and var subfalcata,but basing the description more on the characters of var pallida.
As this was the first legitimate use of pallida as a specific epithet in Licania,L. pallida
Sagot must take priority over L. pallida of Britton, I890 ( =L. brittenianaFritsch) or
L. pallida (Hook. f.) Kuntze, 1891 (=L. hookeriFritsch and L. octandra(Hoffmgg. ex
R. & S.) Kuntze). In spite of the fact that Fritsch (Osterr. Bot. Zeitschr. 42: 6. 1892)
showed that the correct name for this species is L. pallida Sagot, the epithet pallida
has frequently been applied illegitimately to other species by later authors.

82. Licania gracilipes Taubert, Bot. Jahrb. 15(Beibl. 34): 8. 1892.

LicaniaduckeiMaguire, Trop. Woods 93: 29. 1948, synon nov. Type. Ducke248, Brazil, Amazonas,
fl (holotype, NY; isotypes, K, R, RB).

Tree to 20.0 m tall or shrub, the young branches glabrous, obscurely lenticellate.
Leaves lanceolate to oblong-lanceolate, coriaceous, 3.5-8.0 cm long, I.3-3.5 cm
broad, the apex with a finely pointed acumen IO.O-I5.0 mm long, cuneate or sub-
cuneate at base, glabrous on upper surface, the lower surface with short dense white
persistent lanate pubescence; venation shallowly reticulate; midrib glabrous, more
or less plane above; primary veins 8-I5 pairs, prominulous beneath, plane above;
petioles 3.0-6.0 mm long, glabrous, eglandular, shallowly canaliculate. Stipules ca
I.5 mm long, linear, coriaceous, persistent, adnate to extreme base of petiole.
Inflorescences terminal and axillary branched panicles, the rachis and branches
glabrous. Flowers ca I.5 mm long, in cymules on long slender peduncles attached to
primary inflorescence branches. Bracts and bracteoles minute, persistent, glabrous.
Receptacle campanulate, glabrous on exterior, tomentellous within; pedicels o.5-
3.0 mm long. Calyx lobes acute, puberulous on both surfaces. Petals absent. Stamens
3, unilateral; filaments shorter than calyx lobes, glabrous, free to base. Ovary in-
serted at base of receptacle, tomentose. Style rising to base of filaments, pubescent.
Fruit to 1.8 mm long, pyriform; epicarp pulverulent, drying reddish; mesocarp thin;
endocarp hard, thin, sparsely hirsutulous within.
TYPE. Glaziou 13800, Brazil, without locality, fl (holotype, B, lost; isotypes, K, P).
DISTRIBUTION. Non-flooded forest in Amazonian Brazil.
Licania 121

BRAZIL. Amazonas: ChagasINPA 5549 fl (INPA, NY); DuckeRB 35573 fl (K, RB); Froes26247
fr (IAN); Schultes& Lopez 9748 fl (GH, K, NY, US); SchwackeIII-292 fl (GOET, R). Pari: Ducke
RB I5157 fl (K, MG 17219, RB). Terr. Rondbnia: Maguire,Pires, Maguire & Silva 56550 fr (F, GH,
LE, MICH, NY); Pires 56789 fr (UB).

The Index Kewensis mistakenly cited this species originally as L. gracilis. This
error was corrected in the addenda to the same volume. However, the original mis-
take has still led to the species being referred to as L. gracilisfrom time to time, and
hence caused confusion with L. gracilisKleinh., which is not closely related.
Licaniagracilipesis distinct from related species by the white lanate-arachnoid
pubescence of the leaf underside. Although the type of this Amazonian species is a
Glaziou number, the specimens were presumably collected by another. As is so often
the case in Amazonian species, allegedly collected by Glaziou, and thus purporting
to be from southern Brazil, the type is more probably a duplicate of a Schwacke
number (ie Schwacke 111-292, cited above). The association of L. gracilis with
southern Brazil led Maguire to overlook this name when he described L. duckei,
which is a synonym of L. gracilis.

83. Licania parvifructa Fanshawe & Maguire, Bull. Torrey Club 75: 374.
1948.

Large tree, the young branches glabrous and lenticels inconspicuous. Leaves
ovate to oblong, coriaceous, 4.0-9.0 cm long, 2.5-3.5 cm broad, apex with a fine
acumen 5.0-I8.o mm long, rounded, and subconduplicate at base, glabrous on upper
surface, with a short pulverulent-farinaceouspubescence on lower surface, the vena-
tion shallow; midrib glabrous above, more or less plane or slightly impressed at base;
primary veins 7-I I pairs, prominent beneath, plane above; petioles 3.0-6.0 mm long,
glabrous to hirsutulous, eglandular, canaliculate. Stipules 1.0-2.0 mm long, linear,
coriaceous, persistent, adnate to extreme base of petiole. I nflorescencesterminal and
axillary branched panicles, the rachis and branches glabrescent. Flowers ca I.5 mm
long, in cymules on long slenderpeduncles attached to primary inflorescencebranches.
Bracts and bracteoles 0.2-1.5 mm long, persistent. Receptacle campanulate, glabres-
cent on exterior, tomentellous within; pedicels I.0-3.0 mm long. Calyx lobes acute;
puberulous on both surfaces. Petals absent. Stamens 3, unilateral; filaments shorter
than calyx lobes, glabrous, free to base. Ovary inserted at base of receptacle, hirsutu-
lous. Style rising to base of calyx lobes, pubescent throughout. Fruit to I.5 mm long,
pyriform; epicarp pulverulent; mesocarp thin; endocarp hard, thin, sparsely hir-
sutulous within.
TYPES. Abraham 258, Guyana, fl fr (lectotype, NY; isolectotypes, K, U); Fan-
shawe 588(F.D.3324) fr (paratypes, NY, P).
DISTRIBUTION. Primary forests of the Guianas and Amazonian Brazil.
VENEZUELA. Bolivar: Wurdack& Monachino41040 fr (LE, NY); 41175 fr (F, GH, MICH,
NY, RB). GUYANA. Hohenkerk86 fr (K). BRAZIL. Amazonas: Coelho9 fl (INPA 6519, IAN). Terr.
Amapi: Pires, Rodrigues& Irvine505I5 fl (F, GH, LE, MG, NY, RB); Pires & Cavalcante52228 fl
(GH, IAN, MG, NY).

LOCALNAMES.Guyana: Marishiballi, Unikiakia.

This species is very close to L. gracilipesTaub., but is distinguishedby the broader
leaves which bear thinner, less tenacious farinaceouspubescence on the lower surface,
and by the more rounded leaf bases.
122 Flora Neotropica

84. Licania cymosa Fritsch, Ann. Naturh. Mus. Wien 4: 47. 1889.

Tree, the young branches glabrous, prominently lenticellate. Leaves oblong-

elliptic, thick-coriaceous, 2.0-5.0 cm long, 1.3-3.2 cm broad, revolute at margins,
rounded at apex, subcuneate at base, glabrous on upper surface, the lower surface
persistently gray-farinaceous;midrib plane above, glabrous; primary veins 5-6 pairs,
prominulous beneath, plane above; petioles 1.5-2.0 mm long, glabrous, eglandular,
terete. Stipules linear, persistent, I.0-2.0 mm long, adnate to extreme base of petiole.
Inflorescences terminal and axillary cymose panicles; the rachis and branches glab-
rous. Flowers ca 3.0 mm long, in small cymules on slender peduncles attached to
rachis and primary branches of inflorescence.Bracts and bracteoles 0.5-I .0 mm long,
lanceolate, puberulous, persistent. Receptacle urceolate, sessile, short-gray-tomentose
on exterior, tomentose within. Calyx lobes acute, tomentellous on both surfaces.
Petals absent, Stamens 5-6, unilateral; filaments shorter than calyx lobes, glabrous,
free to base. Ovary inserted at base of receptacle, villous. Style equalling filaments,
pilose. Fruit globose, 2.0-2.5 cm diam.; epicarp waxy; pericarp thick, fibrous,
sparsely hirsutulous within.
TYPE. Blanchet3200, Brazil, Bahia, fl (holotype, W; isotypes, F, G, GOET, OXF,
P).
Known only from three inadequate collections from central and
DISTRIBUTION.
eastern Brazil.
BRAZIL. Para: DuckeMG 2870 fl (BM, MG, RB). Bahia: Belem & Pinheiro3082 fr (NY, UB).

This species is close to L. buxifolia,but differs in the cymose inflorescence with

long, slender peduncles.

85. Licania ternatensis Hooker f. ex Duss, (Hooker f., Kew Bull. 1893: 251.
1893, nom nud,) ex Duss, Fl. Antill. Franc. 259. I897; Urban, Symb.
Antill. 5: 352. I907.

Licaniahypoleucasensu Grisebach, Fl. Brit. W. Ind. 230. I860, non Benth.

Medium-sized tree, the young branches puberulous, becoming glabrous and

lenticellate with age. Leaves ovate-elliptic, coriaceous, 4.5-I7.0 cm long, 2.0-8.0 cm
broad, acuminate at apex, with acumen 8.0-I8.o mm long, rounded and subcondu-
plicate at base, glabrous on upper surface, the lower surface with short gray lanate-
arachnoid pubescence; midrib glabrous and more or less plane above; petioles
3.0-6.0 mm long, glabrous, eglandular, terete or slightly canaliculate, rugose.
Stipules ca i.o mm long, lanceolate, coriaceous, persistent, adnate to extreme base of
petiole. Inflorescencesterminal and axillary panicles, the rachis and branches tomen-
tellous to puberulous. Flowers ca 1.5 mm long, in cymules on long slender peduncles
attached to primary inflorescence branches. Bracts and bracteoles 0.2-0.8 mm long,
persistent,puberulous. Receptacle campanulate, tomentellous on exterior and within;
pedicels ca 0.25 mm long. Calyx lobes acute, tomentellous on both surfaces. Petals
absent, Stamens 3-5, unilateral; filaments shorter than calyx lobes, glabrous, free to
base. Ovary inserted at base of receptacle, tomentose. Style rising to base of filaments,
sparsely pubescent throughout. Fruit pyriform, 2.0-2.5 cm long; epicarp rufous-
tomentose; mesocarp thin, fleshy; endocarp hard, thin, woody, hirsutulous within.
TYPES.Duss 2868, Guadeloupe, fl (isosyntypes, NY, US); Duss I903, Martinique,
fl (isosyntypes, NY, US).
Licania 123

Forests and forested slopes of the LesserAntilles from Guadeloupe

DISTRIBUTION.
to Grenada.
LEEWARD ISLANDS. Gaudeloupe: Questel4745 fl (US). WINDWARD ISLANDS. Dominica:
Eggers742 fl (BR, GH, GOET, LE, P, R); 1073 fl (K, US); Fishlock22 fl (BM, F, GH, K, MO, NY,
P, S, US); Hodge578 fr (BM, GH, MO, NY, US); 1237 fr (GH, US); 13I9 st (GH); 2067 st (GH);
2672 fr (US); 2840 st (GH); 3590 St (GH); Imray 129 fr (GH, K, LE); i66 fl (K); sn fl (GOET);
Lloyd 761 fr (NY); Ramagesn (i888) fr (BM, K, S); sn (1889) fr (GH, K). St. Lucia: Beard 192 fr
(A, K, NY, U); Webster,Ellis & Miller 9264 fl (A, MICH, US); 9334 fl (A, MICH, S, U, US). St.
Vincent: H. H. & G. W. Smith 1730 fl (K, NY). Grenada: Broadway57 fl (K); Webster,Ellis & Miller
9530 fl (US).

LOCAL NAMESAND USES. Bois diable, Bois de masse, Bois fer, Bois gris, Bois resolu.
As the vernacular names suggest the wood is very hard. It is used for subterraneanand
submerged structuresas well as to make charcoal.
The differences between this species and L. membranacea are very small, but as
they appear to be constant, the two are retained separately. The name L. ternatensis
is that of Duss (1897), who must be regarded as the author, rather than Urban who
did not describe L. ternatensis
until I907 but to whom the epithet has been attributed
in most earlier works. Urban also noted that L. ternatensisallegedly grew in Trinidad,
but that he had not seen the material. All material from Trinidad named L. ternatensis
that I have examined actually belongs to L. membranacea. Licaniaternatensis,as defined
here, appears to be confined to the Antilles. It differs from L. membranacea in the
shorter petioles which are not conspicuously canaliculate, the tomentose, more
densely pubescent branches of the inflorescence, and the more triangular leaves.

86. Licania membranacea Sagot ex Lanessan, P1. Util. Colon. Franc. I30.
I886; Sandwith, Kew Bull. I931: 373. I931.
LicaniagalibicaR. Benoist,Bull. Mus. Hist. Nat. Paris25: 515. I919.Type. Sagot o081.

Tree to 32.0 m tall, the young branches puberulous, becoming glabrous and
lenticellate with age. Leaves oblong, coriaceous, 8.0-19.0 cm long, 3.7-7.8 cm broad,
the apex with fine acumen I0.0-25.0 mm long, rounded to subcuneate at base,
glabrous on upper surface, the lower surface with short appressed lanate-arachnoid
pubescence, the venation shallow; midrib plane or prominulous above, glabrous;
primaryveins 7- o pairs, prominent beneath, plane above; petioles 8.o-i2.0 mm long,
glabrous or glabrescent, eglandular, canaliculate, transversely rugose. Stipules
3.0-7.0 mm long, linear, coriaceous, persistent, adnate to extreme base of petiole.
Inflorescences terminal and axillary panicles, the rachis and branches puberulous.
Flowers ca 1.5 mm long, in cymules on long slender peduncles attached to primary
branches of inflorescence. Bracts and bracteoles 0.5-I.0 mm long, persistent, puber-
ulous. Receptacle campanulate, tomentellous on exterior, tomentellous within;
pedicels 0.25-I.o mm long. Calyx lobes acute, tomentellous on both surfaces. Petals
absent. Stamens 3-5, unilateral; filaments shorter than calyx lobes, glabrous, free to
base. Ovary inserted at base of receptacle, tomentose. Style equalling filaments,
lanate. Fruit pyriform, ca 2.5 cm long; epicarp brown rufous tomentose; mesocarp
thin, fleshy; endocarp hard, thin, woody, hirsutulous within.
TYPE. Sagot o08i, French Guiana, fl (holotype, P; isotypes, BM, BR, GOET,
K, NY, S).
DISTRIBUTION. Forested slopes and non-flooded forest in Trinidad, the Guianas
and Venezuela, and in Brazil east of Belem.
124 Flora Neotropica

VENEZUELA. Carabobo: Steyermark 95233 fr (U, VEN). Miranda: Steyermark 90088 fr (F, NY,
VEN); Bernardi5719 st. (VEN). Guarico: AristeguietaI77I fl (NY). Sucre: Steyermark& Agostini91o27
fl (NY, VEN); 91361 fl (NY, VEN). TRINIDAD. Bot. Gdn. 6420 fl (K, TRIN); Cruegersn fl (K);
Marshall 11912 fl (K, NY, TRIN); 12678 fl (TRIN); Prance 2105 fr (NY); Purdie sn fl (GH, K);
Richardson12668 fl (K, NY). FRENCH GUIANA. Bettenfeldsn fl (P); Lemeesn fl (P); Melinon60 fl (P).
BRAZIL. Para: DuckeRB 18813 fl (RB); GoeldiMG 8233 fl (MG, RB); Oliveira379 fl (IAN); Pires &
Silva 10542 fl (IAN, NY); o0649 fl (IAN, NY); 11237 fr (NY); 11285 fl (IAN, NY); Prance & Silva
5886i fr (F, GH, NY, UB); 58877 fr (F, GH, NY, RB); Prance& Pennington1938 fl (F, GH, IAN, NY);
2005 fl (F, GH, IAN, NY); R. S. RodriguesMG 8275 fl (MG, RB).

LOCALNAMES.Venezuela: Belero, Soberbio.Brazil: Pintadinha.

87. Licania piresii Prance, sp nov Fig 19 J-M.

Arbor parva vel media, ramulisjuvenilibus lenticellatis. Folia alternatapetiolata;

petiolo 3.0-4.0 mm longo, glabro, eglanduloso, leviter canaliculato, rugoso; laminae
oblongae, coriaceae, 6.0-I5.0 cm longae, 3.5-6.0 cm latae, basi subcuneatae, apice
in acumen caudatum 5.0-I2.0 mm longum contractae, supra glabrae, subtus griseo-
pulverulento-farinaceae pubescentes, venulis planis; costa media supra glabra,
prominente; costis secundariis 7-9 jugis, subtus prominentibus, supra leviter promi-
nentibus. Stipulae 2.0-3.0 mm longae, lineares, parvae, coriaceae, persistentes, cum
basipetioli adnatae. Panicula composita axillaribusvel terminalibus,cymulispluribus2
vel 3 floris breviter pedunculatis constructa; rachi ramisque glabris vel glabrescentis;
flores circa 2.0 mm longi. Bracteae bracteolaeque 0.2-0.7 mm longae, persistentes,
puberulae. Receptaculum campanulatum, extus tomentellum, intus tomentellum;
pedicelli 0.25-0.5 mm longi. Calycis-lobi acuti, utrinque tomentelli. Petala nulla.
Stamina 4-5, unilateralia, filamenta glabra, ad basim libera, calycis-lobis breviora.
Ovarium ad basim receptaculi insertum, tomentellum. Stylus e basi ovarii ortus,
praeter basim glaber. Fructus mihi ignoti.
TYPE. Pires, Rodrigues & Irvine 50435, Brazil, Terr. Amapa, Rio Araguari, fl
(holotype, NY).
DISTRIBUTION. Collected only in non-flooded forest in Territorio do Amapa,
Brazil.
BRAZIL. Terr. Amapa: Rio Araguari, Pires, Rodrigues& Irvine50762 fl (MG, NY); Rio Oiapo-
que, Maguire,Pires & Maguire47108 fl (MG, NY).

This species is most closely related to L. hypoleuca,but is distinct in the larger

leaves, the glabrous inflorescence,and the pulverulent furfuraceouspubescence of the
lower leaf surface. It is with pleasure I name this species for Dr. J. M. Pires who has
contributed so much to our knowledge of the Brazilian flora, and who has extended
much help to me.

88. Licania hypoleuca Bentham, Bot. Voy. Sulph. 91, t. 32. I844; Fritsch,
Grisebach, Fl. Brit.
Ann. Naturh. Mus. Wien. 4: 47. 1889, non L. hypoleuca
West Ind. 230. i86o.

Tree to 35.0 m tall, the young branches glabrous, lenticellate. Leaves ovate to
oblong-lanceolate, membraneous to coriaceous, 2.5-IO.O(-I I.5) cm long, 0.8-4.5
(-5.5) cm broad, the apex tapering to a fine acumen, 5.o-I6.o mm long, usually
subcuneate at base but varying from rounded to cuneate, glabrous on upper surface,
 125
125
Licania

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^; ^^
\v - \

FIG i9. Species of Licania.A-D, L, couepifolia(A. C. Smith2839); A, habit, x 0.5; B, flower, x 5;

C, flower section, x 5; D, flower pubescence, x 5; E-H, L. niloi (Silva 392); E, habit, x 0.5; F, flower
pubescence, x i o; G, flower, X i 0; H, flower section, x 1o. J-M, L. piresii (Pires et al 50435); J, habit
XO.5; K, flower, X 12.5; L, flower section, X 12.5; M, flower pubescence, X 12.5.
126 Flora Neotropica

the lower surface short gray-pulverulent-farinaceous or sordid-lanate; venation

shallowly reticulate; midrib plane to prominulous above, glabrous; primary veins
5-9 pairs, prominulous beneath, plane above; petioles 2.0-6.0 mm long, glabrous or
pubescent and then becoming glabrous with age, transversely rugose, eglandular,
terete or canaliculate. Stipules I.0-2.0 mm long, linear, coriaceous, persistent, adnate
to extreme base of petiole. Inflorescences terminal and axillary branched panicles,
the rachis and branches sparsely puberulous. Flowers 1.5 mm long, on long slender
peduncles attached to primary inflorescence branches. Bracts and bracteoles 0.2-0.5
mm long, persistent. Receptacle campanulate, gray-puberulous to tomentellous on
exterior, tomentellous within; pedicels 0.25-1.0 mm long, Calyx lobes acute, puberu-
lous on both surfaces. Petals absent. Stamens 3-5, unilateral; filaments shorter than
calyx lobes, glabrous, free to base. Ovary inserted at base of receptacle, lanate. Style
sparsely hirsute throughout. Fruit pyriform to ovoid, usually less than I.5 cm long
rarely 2.0 cm long; epicarp with sordid pulverulent indumentum; mesocarp thin;
endocarp thin, hard, fibrous, hirsutulous within.

Key to the Varieties of Licania hypoleuca

i. Leaf undersurface lanate to pulverulent; stomatal cavities absent. a. var hypoleuca.

i. Leaf undersurface with stomatal cavities which are filled by a lanate pubescence.
b. varfoveolata.

88a. Licania hypoleuca var hypoleuca

LicaniahypoleucaBentham, Bot. Voy. Sulph. 91, t. 32. 1844.

Licania microcarpaHooker f., Mart. Fl. Bras. 14(2): 15. 1867, synon nov; Fritsch, Ann. Naturh.
Mus. Wien 4: 47. 1889; Macbride, Fl. Peru 1072. 1938. Type. Spruce3696, Brazil, Amazonas
fl (holotype, K; isotypes, BM, BR, C, CGE, F, GH, GOET, LD, LE, NY, OXF, P).
Licaniagrisea Kleinhoonte, Rec. Trav. Bot. N6erl. 22: 382. 1925, synon nov; in Pulle, Fl. Suri-
name 2(1): 436. I939. Type. B.W. 3392, Suriname, fl (lectotype, U; isotypes, A, MO).
Licaniaparvifolia Pittier, Bol. Soc. Venez. Ci. Nat. 4: 351. 1938, synon nov, nom illegit non
parvifolia Huber. Type. Holt & Gehringer69, Venezuela, Bolivar, fl (holotype, VEN;
isotypes, F, NY, US).
Licania parvifloraBentham var. conduplicataMaguire, Fieldiana Bot. 28: 253. 1952, synon nov.
Type. Steyermark 60277, Venezuela, Bolivar, fl (holotype, NY; isotypes, F, VEN).

Leaf undersurface lanate to pulverulent; stomatal cavities absent.

TYPE. Hinds sn, Panama, fl fr (holotype, K; isotype, LE).
DISTRIBUTION. A wide-ranging variety on sandy soils in forest and on savannas
from southern Mexico through central America, Colombia, Venezuela, the Guianas
and scattered localities in Amazonian Brazil.
Representative collections: MEXICO. Veracruz: Miranda 8506 fl (MEXU). Tabasco: Matuda
3054 fl (A, F, MEXU, MICH, NY); 3110 fl (A, F, K, MEXU, MICH, NY); 3142 fr (A, F, MEXU,
MICH, NY); 3565 fr (A, F, MEXU, MICH, NY). Chiapas; Enriquez7291 fr (MEXU); Miranda7291
fr (US). GUATEMALA. Jones & Facey3272 fl (NY); 3288 fl (NY); Standley25106 fl (GH); 72615 fl
(A, F, NY); 73052 st (F); 79082 st (F); Steyermark
38046 st (F); 38842 st (F); 39509 st (F); 44638 st (F);
45525 st (A, F, MICH); Whitford& Stadtmiller68 fl (GH). BRITISH HONDURAS. Bartlett 11290 fr
(A, MEXU, NY); I3034 fl (F, MICH, S); Gentle1249 (A, F, GH, K, MICH, MO, NY, U); 1182 fr
(A, F, GH, K, MICH, MO, NY); 1287 st (A, F, GH, K, MICH, MO, NY); 1585 fl (A, K, MICH,
MO, NY); 2744 fl (A, F, K, MEXU, MICH, NY); 3342 fl (A, MEXU, MICH, MO, NY); 5570 fr
(BM, MICH, S); 6193 fl (F, MICH); 7028 St (F, MICH); 7081 fr (F, MICH); 9i88 fr (BM, F, MICH);
9744 fl (BM, MICH); Kinloch234 st (K); I70 fr (K); Lundell66oi fr (F, GH, MICH, NY, S); 6963 fl
(F, GH, K, MEXU, MICH, NY, S); Peck471 fi (GH, K); Schipp103 fl (A, BM, F, GH, K, LE, MICH,
MO, NY, S); 292 fl (A, BM, F, GH, K, MICH, MO, NY, S). HONDURAS. Standley53774 st (A, F);
Theime5464 fl (BM, GH, K). NICARAGUA. Standley20152 st (F). PANAMA. Dwyer 1973 fr (MO);
Licania 127

7085 fr (MO); Duke 8053 fr (MO); 8093 fr (MO); 8131 fr (MO); Johnston1782 fr (A); Pittier 5491 fl
(F); Shattuck823 fr (F); Standley31323 st (A); 41067 st (A); Tysonet al 3859 fr (MO). COLOMBIA.
Meta: Pinto, Bischler& Jaramillo 255 fr (P). Vaupss: Cuatrecasas7245 fl (COL, F, NY, US); Schultes&
Cabrera12832 fl (BM, COL, F, GH, NY, US); 12887 fl (BM, COL, GH, NY, US); 13536 fl (BM, COL,
F, GH, LE, NY, US). VENEZUELA. Bolivar: Berti 290 st (VEN); Blanco30 fl (VEN); 41 fl (VEN);
75 fl (VEN); 290 fr (VEN); Steyermark 86459 fr (VEN). Terr. Amazonas: Breteler4720 fl (U); Foldats
3626 fl (VEN); Jordansn fl (NY, VEN); Maguire & Wurdack35723 fl (MICH, NY); Maguire, Wurdack
& Maguire4z663 fl (F, NY); Maguire, Wurdack& Keith 41833 fr (NY, RB); 41891 fr (MO, NY, RB);
Maguire, Wurdack& Bunting36378 fl (NY, SP); Wurdack& Adderley43242 fl (NY); 43612 fl (F, GH,
LE, NY). GUYANA. De La Cruz 2715 fr (F, GH, MO, NY, US); 2858 fr (F, GH, MO, NY, US);
Fanshawe66i(F.D. 3397) fl (K, NY, U); Davis 288(F.D. 2629) fr (FHO, K); 369(F.D. 2365) fl (FHO,
K, U); Gleason247 fr (GH, NY). SURINAME. B.B.S.I96 fr (K); B.W. 1365st (U); 17295st (U);
28136fl (K, NY); 24i86st (U); 30605fr (IAN, NY); 36325st (U); 3860 fl (IAN, K, NY); 58825 fr (U);
6113 fl (A, MO, U); 6864 fl (K, NY); van Donselaar 1200 st (U); 1897 st (U); 2915 st (U); 3036 fr
(NY, U); Lems 650224 st (NY); Stahel 15oa fr (IAN, K, NY, U). FRENCH GUIANA. Benoist625 fr
(P); Melinon76fl (P). PERU. Loreto: Klug 1525 fl (F, NY, US). San Martin: Klug 3740 fl (A, BM, F,
GH, K, MO, NY, S, US). BRAZIL. Amazonas: CoelhoINPA 6519 fl (INPA); Frdes 23329 fl (IAN,
UB), 25229 fr (IAN, NY); 25204 fl (IAN, NY); 28777 fl (IAN); Froes & Addison29037 fl (IAN, UB);
Pranceet al 4746 fl (INPA, NY); 4875 fl (INPA, NY); 4937 fl (INPA, NY); 508I fl (INPA, NY). Terr.
Roraima: Ducke2233 fl (IAN, NY, R). Terr. Rond6nia: J. F. Silva 173 (IAN). Mato Grosso: Kuhlmann
1922 fl (R, RB, SP). Bahia: Belim & Pinheiro3186 fl (NY, UB); 3203 fl (NY, UB); 3209 fr (NY, UB).

LOCAL NAMES. Guatemala: Chozo.Venezuela: Hierrillo,HierrilloBlanco.Guyana:

IronMary, Unikiakia.Suriname: OenikiakiaDjamaro.Brazil: Caripe.
The names here reduced to synonymy applied to specimens which were analysed
with the use of scatter diagrams, and no grounds for keeping them apart were found,
since the characters used by earlier authors showed gradation into one another.

88b. Licania hypoleuca var foveolata Prance, nov var

A var hypoleucodiffert foliorum laminis subtus cum cavis stomalibus munitis.

Differs from var hypoleuca in the presence of stomatal cavities on the leaf under-
side.
TYPE.Schultes& L6pez9719, Brazil, Amazonas, Rio Curicuriary, fl (holotype,
US; isotype, IAN).
Forest in the region of the Rio Curicuriary.
DISTRIBUTION.
BRAZIL. Amazonas, Rio Curicuriary. DuckeRB 35571 fl (K).

This material differs only in the presence of stomatal cavities on the leaf under-
side. Since this is an important character in the genus, this variety is recognized here.

8. Licania subgen Licania sect Pulverulenta

89. Licania boyanii Tutin, Jour. Bot. 78: 255. 1940.

Large tree to 25.0 m tall, the young branches sparsely tomentellous, becoming
glabrous with age, lenticellate. Leaves ovate-orbicular to elliptic, thick-coriaceous,
6 = Paratypes of L. grisea.
128 Flora Neotropica

2.5-5.5 cm long, .5-3.5 cm broad, rounded at apex, rounded to subcuneate at base,

glabrous above, with waxy-brown pulverulent-furfuraceouspubescence beneath, the
margins slightly recurved; midrib impressed above, glabrous; primary veins 5-7
pairs, prominulous beneath, plane or slightly impressed above; petioles 1.5-3.0 mm
long, terete, rugose, eglandular, sparsely pubescent. Stipules I.0-3.0 mm long,
lanceolate, persistent, tomentose when young, adnate to base of petiole. Inflores-
cences short terminal and axillary racemose panicles, 3.0-6.0 mm long, the rachis
and recurved branches with short dense sordid-furfuraceouspubescence. Flowers ca
I.5 mm long, sessile, solitary on primary branches of inflorescence. Bracts and
bracteoles minute, persistent, triangular, with three partially fused bracteoles sub-
tending each flower. Receptacle campanulate, sessile, tomentellous on exterior,
tomentose within. Calyx lobes acute, tomentellous on both surfaces. Petals absent.
Stamens 3, unilateral, opposite three calyx lobes; filaments very short, glabrous, free
to base. Ovary inserted at base of receptacle, lanate, globose. Style shorter than calyx
lobes, pubescent at base only. Fruit ca 2.8 cm long, globose, with base narrowed into
a stipe; epicarp pulverulent; pericarp undifferentiated, hard, granular, velutinous
within.
TYPE. Tutin293, Guyana, fl (holotype, BM; isotypes, K, NY, RB, U, US).
DISTRIBUTION. Wallaba forest and forest on slopes in Venezuela and Guyana.
Flowering April to June.
VENEZUELA. Bolivar: Cardona2303 st (US). GUYANA. De La Cruz 3408 fr (GH, NY, US);
Irwin 256 fl (US); Tutin 230 fl (BM, K, RB, U, US).

LOCAL NAME. Guyana: Thick-skinnedMarishiballi.

90. Licania buxifolia Sandwith, Kew Bull. 1931: 369. I93I.

Large tree, the young branches glabrous, lenticellate. Leaves ovate-elliptic to

elliptic, thick coriaceous, 3.0-I2.0 cm long, 1.5-6.2 cm broad, usually rounded to
obtuse at apex, rarely mucronate, subcuneate at base, the margins revolute, glabrous
above, with gray waxy pulverulent-furfuraceouspubescence beneath; midrib glab-
rous above, impressed; primary veins 5-9 pairs, prominent beneath, impressedabove;
petioles 2.0-6.0 mm long, glabrous, terete, with two rather obscure glands at junction
with upper surface of lamina. Stipules ca I.o mm long, lanceolate, coriaceous, sub-
persistent, adnate to base of petiole. Inflorescences terminal and axillary little-
branched racemose panicles, 5.0-I5.0 cm long, the rachis and branches glabrous.
Flowers 1.5-2.0 mm long, solitary or in small groups attached to primary branches
by very short peduncles. Bracts and bracteoles ca 0.2 mm long, persistent, triangular,
pubescent on exterior. Receptacle campanulate, sessile,gray-tomentellouson exterior,
tomentellous within. Calyx lobes acute, tomentellous on both surfaces. Petals absent.
Stamens 5, unilateral; filaments very short, glabrous. Ovary inserted at base of
receptacle, globose, tomentose. Style rising to base of filaments, pubescent at base
only. Fruit unknown.
TYPES.Sandwith327, Guyana, fl (holotype, K; isotypes, NY, RB, U); Jenman
4110 fl (paratypes, BM, K, NY, US).
DISTRIBUTION. Known only from Wallaba forest in Guyana.
GUYANA. Davis sn (F.D. 2309) fl (BM, FHO, K, U).

LOCALNAME.Marishiballi (Arawak).
Licania 129

9I. Licania orbicularis Spruce ex Hooker f., Mart. Fl. Bras. 14(2): 13. I867;
Fritsch, Ann. Naturh. Mus. Wien 4: 46. 1889.

Small to medium-sized tree, the young branches puberulous, becoming glabrous

and lenticellate with age. Leaves predominantly orbicular but occasionally elliptic,
thick-coriaceous, 3.0-10.0 cm long, 1.8-7.0 cm broad, retuse rounded or acute at
apex, rounded to subcuneate at base, the margins revolute, glabrous above, with
waxy gray pulverulent-furfuraceouspubescence beneath; midrib glabrous and plane
above; primary veins 7-II pairs, prominent beneath, prominulous above; petioles
2.0-5.0 mm long, puberulous, becoming glabrescent with age, terete, with two rather
obscure glands at lamina base. Stipules 1.0-4.0 mm long, deltoid, coriaceous, adnate
to base of petiole, subpersistent. Inflorescences terminal and axillary little-branched
racemose panicles, 6.0-1o.o cm long, the rachis and branches tomentellous. Bracts
and bracteoles lanceolate, ca 2.0 mm long, in small groups on primary branches of
inflorescence. Receptacle urceolate, sessile, light brown tomentose on exterior,
sparsely puberulous within. Calyx lobes acute, tomentellous on both surfaces. Petals
absent. Stamens 2-4, unilateral; filaments shorter than calyx lobes, glabrous. Ovary
inserted at base of receptacle, pubescent. Style rising to base of filaments, pubescent.
Fruit pyriform, ca 4.0 cm long, 2.0-2.5 cm broad; epicarp glabrous, drying yellow
and wrinkled; mesocarp very thin; endocarp hard to 3.0 mm thick, granular and
fibrous, velutinous within.
TYPE.Spruce3760, Venezuela, Terr. Amazonas, fl (holotype, K; isotypes, BM,
BR, CGE, NY, P).
DISTRIBUTION. Riverine forest and savanna margins in Amazonian Venezuela
and adjacent Brazil. Flowering October to January.
VENEZUELA. Terr. Amazonas: Maguire, Wurdack& Bunting 36641 fr (F, GH, NY, US);
37478 fr (F, NY, US); 37579 fr (NY, US); Maguire29260 fl (GH, NY, US); LI. Williams I4292 fr (F,
US, VEN). BRAZIL. Amazonas: Frdes27889 fr (IAN, UB).

92. Licania niloi Prance, sp nov Fig 19 E-H.

Arbor media, ramulis juvenilibus glabris haud lenticellatis. Folia alternata

petiolata; petiolo 3.0-5.0 mm longo, glabro, tereti, rugoso, cum glandulis duobus
sessilibus munito; laminae oblongo-ellipticae, crasse coriaceae, 4.5-9.0 cm longae,
2.0-4.2 cm latae, basi rotundatae vel subcuneatae, apice acutae, supra glabrae,
subtus indumento ceroso pulverulento-furfuraceomunitae; costa media supra glabra,
plana; costis secundariis 5-7 jugis, subtus prominentibus, supra planis. Stipulae 2.0
mm longae lanceolatae, coriaceae, persistentes, cum petiolo adnatae. Flores 3.0-4.0
mm longi, haud pedunculati, in paniculis brevis terminalibus vel axillaribus dis-
positi, rachi ramisque tomentellis. Bracteae bracteolaeque 0.5-3.0 mm longae, ovatae,
persistentes,extus puberulae. Receptaculum angustatum, cylindrico-urceolatum, basi
inflatum, sessile, extus tomentellum, intus puberulum. Calycis-lobi lanceolati,
utrinque puberuli. Petala nulla. Stamina 3, unilateralia; filamenta glabra, ad basim
libera, calycis-lobis breviora. Ovarium ad basim receptaculi insertum, breviter
pubescens. Stylus e basi ovarii ortus, calycis-lobis brevior, ad apicem puberulentum.
Fructus mihi ignoti.
TYPE. Nilo T. Silva392, Brazil, Terr. Rondonia, Porto Velho, fl (holotype, NY;
isotype, IAN).
DISTRIBUTION. Known only from the type gathering collected in primary forest.
Licania niloi is easily distinguished from any other Licaniaby the shape of the
130 Flora Neotropica

receptacle, which is extremely long and slender with a swollen base, and by the
narrow, lanceolate calyx lobes. It is most closely related to L. coriaceabut differs in
the characters given above.

93. Licania coriacea Bentham, Jour. Bot. Hooker 2: 221. 1840; Sagot, Ann.
Sci. Nat. Bot. VI. 15: 306. 1883; Fritsch, Ann. Naturh. Mus. Wien 4: 49.
I889.

LicaniapallidulaStandley, Bull. Torrey Club 67: 286. 1940, synon nov. Type. Pinkus89, Venezuela,
Bolivar, fr (holotype, F; isotypes, GH, K, NY, US).

Tree to 25.0 m tall, the young branches puberulous, soon becoming glabrous
and lenticellate. Leaves elliptic to oblong-ovate, thick-coriaceous, 3.0-12.0 cm long,
1.5-5.0 cm broad, acute or acuminate at apex, with acumen to 6.0 mm long, cuneate
to subcuneate at base, the margins revolute, glabrous above, with waxy gray pulveru-
lent-furfuraceous pubescence beneath; midrib glabrous above, plane or slightly
impressed; primary veins 6-8 pairs, prominulous beneath, plane above; petioles
3.0-6.0 mm long, eglandular, terete, pulverulent, becoming glabrescent with age.
Stipules 2.0-4.0 mm long, lanceolate, coriaceous, adnate to base of petiole, persistent.
Inflorescenceslittle branched terminal and axillary racemose panicles, the rachis and
branches tomentellous to puberulous. Flowers 3.0 mm long solitary or in small
groups along primary branches of inflorescence. Bracts and bracteoles 0.5-2.5 mm
long, deltoid, persistent, tomentellous on exterior. Receptacle urceolate, sessile,
tomentellous on exterior, and within. Calyx lobes acute, tomentellous on both sur-
faces. Petals absent. Stamens 4-5, unilateral; filaments shorter than calyx lobes,
glabrous, free to base. Ovary inserted at base of receptacle, tomentose. Style equalling
filaments, tomentose. Fruit pyriform, 2.5-3.0 cm long; epicarp glabrous, drying
yellow and wrinkled; mesocarp thin; endocarp ca 3.0 mm thick, hard, granular and
fibrous, velutinous within.
TYPE.Schomburgk 50, Guyana, fl (holotype, K; isotypes,BM, BR, CGE, G, L, OXF,
P, US).
DISTRIBUTION. Non-flooded forest in the Guianas and Central Amazonia.
VENEZUELA. Bolivar: Steyermark90840 fl (F, NY, P, US, VEN). GUYANA. Appun 1752 fl
(K); Beccarisn fl (FI, K); Davis 264 (F.D. 2255) fl (K); 452 (F.D. 2447) fl (FHO, K); Drake sn fr (K);
Guppy408 (F.D. 7424) fl (NY); Hohenkerk676 fr (K); Jenman 4041 fl (K, U); Pinkus 62 (F.D. 2834) fl
(FHO, K); A. C. Smith2138 fl (A, F, K, MO, NY, P, S, U, US); 2226 fl (A, F, K, MO, NY, P, S, U,
US). BRAZIL. Amazonas: Rodrigues& Osmarino6911 fr (INPA 15462, NY); 6997 fl (INPA 15659
NY). Para: Ducke MG 8997 fr (MG, RB); RB 19792 fr (K, RB). Terr. Rond6nia: Black, Cordeiro,&
Francisco52-14661 st (IAN, NY).

LOCAL NAMES. Guyana: Marishiballi, Unikiakia.

94. Licania urceolaris Hooker f., Mart. Fl. Bras. 14(2): 15. I867; Fritsch,
Ann. Naturh. Mus. Wien 4: 47. 1889.

Tree to 25.0 m tall, the young branches puberulous, becoming glabrous and
lenticellate with age. Leaves oblong-ovate, chartaceous to coriaceous, 5.5-12.5 cm
long, 2.5-6.2 cm broad, the apex with a well developed acumen 4.0-12.0 mm long,
rounded to subcuneate at base, the margins plane, glabrous above, with short waxy
gray-pulverulent-furfuraceous pubescence beneath; venation glabrous beneath;
Licania 131

midrib glabrous above, plane or slightly impressed; primary veins 6-Io pairs,
prominulous beneath, plane above; petioles 3.0-6.0 mm long, terete or shallowly
canaliculate, puberulous, becoming glabrous with age, with two medial sessile
glands. Stipules ca 2.0 mm long, lanceolate, coriaceous, adnate to base of petiole,
caducous or subpersistent. Inflorescences little-branched terminal and axillary race-
mose panicles, the rachis and branches tomentellous. Flowers 3.0-4.0 mm long,
solitary or in small groups along primary branches of inflorescence. Bracts and
bracteoles 0.5-1.5 mm long, deltoid, persistent, tomentellous on exterior. Receptacle
urceolate, sessile, tomentose on exterior, tomentellous within. Calyx lobes acute,
tomentellous on both surfaces. Petals absent. Stamens 5, inserted in an arc of ca
270?; filaments shorter than calyx lobes, glabrous, free to base. Ovary inserted at base
of receptacle, tomentose. Style equalling filaments, sparsely pubescent to glabrous.
Fruit (immature) pyriform; epicarp glabrous, drying yellow and wrinkled; mesocarp
thin; endocarp thick, hard, pubescent within.
TYPE. Spruce 2422, Brazil, Amazonas, fl (holotype, K; isotypes, BM, BR, C,
CGE, GH, GOET, LD, LE, NY, OXF, P).
DISTRIBUTION. Non-flooded forest in western Amazonia. Flowering August to
October.
COLOMBIA. Vaupds: Schultes& Cabrera13818 fl (COL, GH, US). BRAZIL. Amazonas: Elias
313 fl (MG); Krukoff6817 fr (A, BM, BR, F, K, LE, MICH, MO, NY, RB, S, US); 6926 st (A, BM,
BR, F, K, LE, MICH, MO, NY, RB, S, US); 8725 fl (A, BM, BR, F, K, LE, NY, P, S); 8751 fl
(A, BM, BR, F, K, LE, MO, NY, P, S, US); 8886 fl (A, BM, BR, F, K, LE, MO, NY, P, S).

95. Licania affinis Fritsch, Ann. Naturh. Mus. Wien 4: 50. 1889; Benoist,
Bull. Mus. Hist. Nat. Paris 25: 516. I919; Kleinh. in Pulle, Fl. Suriname
2(I): 440. I939.
L. coriaceaSagot (non Benth.), Ann. Sci. Nat. Bot. VI. 15: 306. I883.

Small tree, the young branches glabrous, lenticellate. Leaves elliptic to oblong,
thick-coriaceous, 5.0-I5.5 cm long, 2.5-6.7 cm broad, acuminate at apex, with
acumen 3.0-I2.o mm long, subcordate to subcuneate at base, the margins plane,
glabrous above, with waxy brown pulverulent-furfuraceouspubescence beneath;
midrib glabrous above, prominulous; primary veins 4-7 pairs, arcuate, prominent
beneath, prominulous above; petioles 3.0-6.0 mm long, glabrous, rugose, shallowly
canaliculate, with two obscure sessile glands near to lamina. Stipules linear, 3.0-5.0
mm long, coriaceous, persistent, adnate to base of petiole. Inflorescence of little-
branched terminal and axillary racemose panicles, the rachis and branches glabrous
to glabrescent. Flowers ca 3.0 mm long, on primary branches of inflorescence.Bracts
and bracteoles 0.3-1.0 mm long, deltoid, persistent, puberulous on exterior. Recep-
tacle urceolate, sessile, tomentellous on exterior, sparsely puberulous within. Calyx
lobes acute, tomentellous on both surfaces. Petals absent. Stamens 3, unilateral;
filaments shorter than calyx lobes, glabrous, free to base. Ovary inserted at base of
receptacle, short-tomentose. Style equalling filaments, pubescent throughout. Fruit
to 4.0 cm long with stipe ca i.o cm long, pyriform; epicarp glabrous, drying yellow
and wrinkled; mesocarp thin; endocarp thick, hard, granular and fibrous, sparsely
hirsutulous within.
TYPE. Schomburgk822, Guyana, fl (holotype, W; isotypes, CGE, G, K, P).
DISTRIBUTION. Riverine and periodically flooded forest in the Guianas and the
state of Para in Brazil.
GUYANA. Cowan & Soderstrom 1361 fl (BR,
2173 fr (NY, US); Jenman 6624 fl (K); Schomburgk
K). SURINAME. Kappler1968 fl (GOET, LE, P, S, U); Lanjouw & Lindeman327 fl (NY); 332 fl
132 Flora Neotropica

(K, NY, U); Wullschlaegel1659 fl (BR, GOET). FRENCH GUIANA. For. Serv. 7204 fr (NY, P, U);
7354 fr (NY, P); 7392 fr (NY, P); Melinonsn fl (BM, F, GH, K, NY, US); 12 fr (P); 287 fl (K, P);
Richardsn fr (P). BRAZIL. Par,: Black, Froes & Ledoux50-9808 fr (IAN, NY); Huber MG 1894 fr
(BM, MG).

LOCALNAMES.French Guiana: Lebi koko, Bois Gaulette.

96. Licania glauca Cuatrecasas, Fieldiana Bot. 27(2): 109. 951.

Tree to I5.0 m tall, the young branches glabrous, sparingly lenticellate. Leaves
ovate-elliptic, thick-coriaceous, 5.0-13.0 cm long, 3.5-8.0 cm broad, abruptly
acuminate or acute at apex, with acumen I.0-5.0 mm long, rounded at base, the
margins slightly revolute, glabrous above, with waxy gray pulverulent-furfuraceous
pubescence beneath; midrib glabrous above, usually slightly impressed; primary
veins 5-7 pairs, prominent beneath, slightly impressed above; petioles 3.0-5.0 mm
long, glabrous, eglandular, shallowly canaliculate. Stipules lanceolate, I.0-2.0 mm
long, coriaceous, persistent, adnate to base of petiole. Inflorescences terminal and
axillary panicles, the rachis and branches ferrugineous-tomentellous. Flowers ca
2.0 mm long, in small groups on short thick peduncles attached to primary branches
of inflorescence' Bracts and bracteoles deltoid, to 2.0 mm long, persistent, tomentose
on exterior. Receptacle campanulate, ferrugineous-tomentoseon exterior, tomentose
within. Calyx lobes acute, tomentellous on both surfaces. Petals absent. Stamens 3,
unilateral; filaments shorter than calyx lobes, glabrous, free to base. Ovary inserted
at base of receptacle, hirsute. Style equalling filaments, hirsute throughout. Fruit
globose, ca 3.7 cm in diameter; epicarp with short appressed rufous pubescence;
mesocarp thin; endocarp hard, woody, ca 4.0 mm thick, hirsutulous within.
TYPE. Killip & Cuatrecasas39000, Colombia, Valle, fl fr (holotype, F; isotypes,
COL, K, MO, NY, S, US).
DISTRIBUTION.
Pacific coastal forest of Colombia.
COLOMBIA. Narinio: Killip & Garcia3321I st (BM). Without locality: Triana 214 fl (P).

The robust leaves and stems of L. glaucadistinguish it from all other species with
pulverulent leaf indumentum.

97. Licania davillaefolia R. Benoist, Bull. Mus. Hist. Nat. Paris 25: 513. 1919.

Large tree, the young branches glabrous, lenticellate. Leaves oblong-elliptic,

chartaceous, 6.0-21.0 cm long, 3.5-8.5 cm broad, short-acuminate at apex, with
acumen 3.0-Io.o mm long, rounded at base, the margins plane, glabrous above, with
waxy gray pulverulent-furfuraceouspubescence beneath; midrib glabrous above
impressed; primary veins 7-10 pairs, prominent beneath, impressedabove; secondary
nerves more or less parallel, prominent beneath; venation of upper surface usually
papillose; petioles 6.o-io.o mm long, glabrescent to puberulous, shallowly canali-
culate, eglandular or with inconspicuous glands, rugose. Stipules 1.0-3.0 mm long,
linear, coriaceous, persistent, adnate to base of petiole. Inflorescences terminal and
axillary spreading branched panicles, the rachis and branches tomentellous to
puberulous. Flowers 1.5-2.0 mm long, solitary on primary and secondary branches of
inflorescence. Bracts and bracteoles 0.3-1.0 mm long, deltoid to lanceolate, persis-
tent, exterior tomentellous. Receptacle campanulate, sessile, tomentose on exterior,
tomentose within. Calyx lobes acute, tomentose on both surfaces. Petals absent.
Licania 133

Stamens 6-7, unilateral; filaments shorter than calyx lobes, glabrous, free to base.
Ovary inserted at base of receptacle, tomentose. Style equalling filaments, sparsely
pubescent throughout. Fruit ovoid to pyriform, ca 3.2 cm long; epicarp densely and
compactly rufous-pubescent; pericarp thick, granular, fibrous, densely hirsutulous
within.
TYPE. Melinonsn, French Guiana, Maroni, fl (lectotype, P; isolectotypes, A,
BM, F, GH, K, R, US); MelinonI44, French Guiana, Mana, fl (paratype, P).
DISTRIBUTION. Riverine forest and non-flooded forest in the Guianas and eastern
Amazonia.
SURINAME. van Donselaar1151 St (U); 1467 fl (NY, U). FRENCH GUIANA. Benoist 1085 st
(P); For. Serv.46M fr (U); 53M fl (P, U); 132 st (U); I73M st (U); Halle 726 fr (P); Wachenheim 13
st (P); 137 st (P). BRAZIL. Para: Black 47-1975 fr (IAN, NY); Ducke MG 16225 fr (MG, RB); RB
19774 fl fr (K, MG I6225); RB 19779 fl (K, RB); RB 19793 fr (RB); KuhlmannRB 17969 fr (K, RB);
Pires & Silva 4695 fl (IAN, INPA, US); Prance& Pennington2050 fr (F, GH, NY); LI. Williams & Silva
18232 fr (NY); no col. RB 19793 fr (K). Terr. Amapa: Irwin, Pires & Westra47781 fl (MG, NY); 48037
fl (IAN, NY); Luetzelburg20232 fl (R); Pires, Rodrigues& Irvine50652 st (IAN, NY); 50903 fl (IAN,
MG, NY).

LOCAL NAMES.French Guiana: Bois Gaulette,Kokolebi,Gris-gris.Brazil: Pajurarana.

Licania davillaefoliahas often been confused with other species, but is easily
distinguished from those bearing a pulverulent leaf indumentum by its membraneous
leaves, which also display impressed venation.

98. Licania elliptica Standley, Publ. Field Mus. Bot. 17: 255. I937.
LicaniapaniculataFanshawe & Maguire, Bull. Torrey Club 75: 323. I948, synon nov. Type.
Maguire 24068, Suriname, fl (holotype, NY; isotypes, A, BR, F, K, US).

Medium-sized tree, the young branches glabrous, lenticellate. Leaves elliptic,

coriaceous, (10.5-) I2.0-I9.0 cm long, 4.0-8.0 cm broad, the apex with well developed
acumen 5.0-15.0 mm long, rounded to cuneate at base, the margins plane; upper
surface glabrous, with two glands at margin towards base; lower surface with waxy
gray pulverulent-furfuraceouspubescence; midrib glabrous above, prominulous;
primary veins 8-9 pairs, prominent beneath, prominulous above; petioles 4.0-7.0 mm
long, glabrous, shallowly canaliculate, often with two rather obscure medial, sessile
glands. Stipules to 3.0 mm long, linear, coriaceous, persistent, adnate to extreme base
of petiole. Inflorescences terminal and axillary spreading branched panicles, the
rachis and branches glabrous or sparsely puberulous. Flowers ca 1.5 mm long, in
small groups on short thick peduncles attached to primary and secondary branches
of inflorescence. Bracts and bracteoles 0.3-1.0 mm long, deltoid, persistent, puberu-
lous on exterior. Receptacle campanulate, sessile, tomentellous on exterior, puberu-
lous within. Calyx lobes acute, puberulous on both surfaces. Petals absent. Stamens
3-5, unilateral; filaments shorter than calyx lobes, glabrous, free to base. Ovary
inserted at base of receptacle, tomentose. Style equalling filaments, sparselypubescent
throughout. Fruit pyriform, ca 3.6 cm long; epicarp glabrous, drying wrinkled and
yellow; mesocarp thin; endocarp thick, hard, fibrous, hirsutulouswithin.
TYPE. Krukoff5014, Brazil, Amazonas, fl (holotype, F; isotypes, A, BM, K, LE,
M, MICH, MO, NY, S, U, US).
DISTRIBUTION. Primary forest in Suriname and Amazonian Brazil.
SURINAME. Maguire24066 fl (A, BR, F, K, MO, NY, US); Stahel281 st (A, GH, K, NY, S, U);
vanDonselaar1040 st (U). BRAZIL. Amazonas: Frdes23885 fr (IAN); Pranceet al 3470 fr (INPA, NY).
Para: Froes31996 fl (IAN).
134 Flora Neotropica

LOCALNAME.Suriname: Marishiballihariraroe.
This species is very similar to L. canescens,but, to judge from the admittedly
limited material at hand, still distinct, especially by the much larger leaves.

99. Licania canescens R. Benoist, Bull. Mus. Hist. Nat. Paris 25: 514. I919.

Tree to 20.0 m tall, the young branches glabrous, lenticellate. Leaves elliptic to
oblong-elliptic, coriaceous, 4.0-I2.0 cm long, 2.0-5.5 cm broad, acuminate at apex,
with acumen 4.0-I5.0 mm long, rounded to cuneate at base, the margins plane,
glabrous on upper surface, the lower surfacewith waxy gray pulverulent-furfuraceous
pubescence; midrib glabrous above, plane; primary veins 6-9 pairs, prominent
beneath, plane above; petioles 3.0-5.0 mm long, glabrous, terete or shallowly canali-
culate, eglandular, transversely rugose. Stipules 2.0-4.0 mm long, linear, persistent,
adnate to base of petiole. Inflorescences terminal and axillary racemose panicles, the
rachis and branches glabrous or sparsely puberulous. Bracts and bracteoles 0.5-2.0
mm long, deltoid, persistent, exterior puberulous. Flowers ca 2.0 mm long, in small
groups on primary branches of inflorescence. Receptacle campanulate, sessile,
tomentellous on exterior, tomentose within, Calyx lobes acute, tomentellous on both
surfaces. Petals absent. Stamens 5, unilateral; filaments shorter than calyx lobes,
glabrous, free to base. Ovary inserted at base of receptacle, tomentose. Style equalling
filaments, sparsely pubescent throughout. Fruit pyriform, ca 3.0 cm long; epicarp
glabrous, drying yellow and wrinkled; mesocarp thin; endocarp hard, thick, fibrous,
hirsutulous within.
TYPE.Melinonsn, French Guiana, Maroni, fl (lectotype, P; isolectotypes, GH,
K); Wachenheim 43, Maroni, fl (paratype, P).
DISTRIBUTION. Riverine and periodically flooded forest in the Guianas, and
eastern, central and southern Amazonia.
VENEZUELA. Terr. Amazonas: Maguire, Cowan & Wurdack30397 fr (NY). GUYANA. Fan-
shawe 2836 (F.D. 5837) fl (K, NY); Guppy653(F.D. 7668) fl (NY). SURINAME. B.B.S. 1o04 st (NY,
U); B.W. 3662 st (K); Lanjouw& Lindeman357 st (IAN, K, NY); Lindeman5874 st (NY, U); 5947 st
(NY, U); vanDonselaar1814 st (U); 2300 st (U). FRENCH GUIANA. For. Dept. 56M fl (P); Melinon
sn (BM, F, GH, NY, R: all possible isotypes); Wachenheim401 fl (GH, P). BRAZIL. Amazonas:
CoelhoINPA 2974 fr (IAN, INPA, NY); Coelho& Mello INPA 426i fl (INPA, NY); Ducke 2211 fr
(IAN, NY, R); MG 2898 fl (MG, RB); Froes 23856 st (IAN, NY); 25226 fr (IAN, NY); Frdes &
Addison29177 (IAN); Krukoff8764 fl (BR, F, LE, NY, P, U, US); Fires 361 fr (IAN, NY); Pranceet al
3821 fr (INPA, NY); Rodrigues2946 st (INPA 10301). Para: Ducke MG 14858 fl (RB 5152); MG
14948 fr (RB I978I); Oliveira623 fl (IAN, UB); 793 fl (IAN); 823 fl (IAN); 3922 fr (IAN, NY); Pires
51830 fr (NY); Pires, Black & Dobzhansky4134 st (IAN, NY); Pires & Silva o1543 fl (IAN, NY);
R. S. RodriguesMG 8276 fl (K, RB 19783); Silva 590 fl (IAN). Terr. Rond6nia: Prance et al 5546 fl
(INPA, NY); 8257 fr (INPA, NY). BOLIVIA. Pando: Pranceet al 6475 fl (INPA, NY).

LOCALNAMES.Guyana: Kwefe (Mawayan). Suriname: Foengoe. Witte Foengoe.

Brazil: Caripe.

9. Licania subgen Licania sect Licania

Ioo. Licania couepifolia Prance, sp nov Fig 19 A-D.

Arbor parva, ramulis juvenilibus tomentellis mox glabris lenticellatis. Folia

Licania 135

alternata petiolata; petiolo circa 4.0 mm longo, dense villoso-pubescens, eglanduloso,

tereti; laminae oblongae vel oblongo-lanceolatae, coriaceae, 10.0-I7.0 cm longae,
2.8-6.5 cm latae, basi rotundatae, apice in acuminem 8.o-I5.0 mm longum con-
tractae, supra glabrae nitidae, subtus lanato-pubescens, venulis profunde reticulatis,
parallelis, hirsutis; costa media impressa, dense pubescente; costis secundariis subtus
prominentibus, hirsutis, supra impressis. Stipulae lanceolatae, tomentosae, 5.0-6.0
mm longae, coriaceae, persistentes,cum basi petioli adnatae. Flores 6.5-7.0 mm longi,
solitarii, breviter pedunculati, in paniculis racemosis axillaribus vel in ramis lignosis
dispositi, rachi ramisque rufo-tomentellis. Bracteae bracteolaeque I.0-2.5 mm longae
lanceolatae, extus tomentellae. Receptaculum campanulato-urceolatum, pro genere
magnum, extus brunneo-tomentosum. Calycis-lobi acuti, extus tomentosi, intus
puberuli. Petala nulla. Stamina 8, in orbem completum insertum; filamenta calycis-
lobos aequantia, ad medium connata, pubescentes. Ovarium ad basim receptaculi
insertum, villosum. Stylus e basi ovarii ortus, calycis-lobos aequans, arachnoideo-
pubescens. Fructus mihi ignoti.
TYPE. A.C. Smith2839, Guyana, Shodikar Creek, Essequibo River. fl (holotype,
NY; isotypes, A, F, K, MAD, S, U, US).
DISTRIBUTION. Known only from the type gathering, collected in lowland forest
in Guyana, in January.
This species is distinct from all other species of Licaniain the flower and in-
florescence.However, both the basal insertion of the ovary and the absence of the petals
indicate that it belongs to Licania.The leaves strongly suggest those species of Couepia
in the C. canomensis complex, but this type of leaf with parallel venation also occurs in
several other species of Licania.

IoI. Licania trigonioides Macbride, Candollea 5: 368. I934; Fl. Peru 1073.
I938.

Low shrub, the young branches densely ferrugineous-hirstulous, lenticellate.

Leaves oblong-ovate to elliptic, membraneous, broadest near base, io.o-i6.o cm
long, 4.0-5.6 cm broad, acuminate at apex, with acumen 12.0-20.0 mm long, cordate
at base, glabrous above, with short dense appressed lanate gray pubescence beneath;
midrib and veins and leaf margins hirsutulous beneath; midrib plane above, villous;
primary veins IO-I4 pairs, prominulous on both surfaces; petioles 2.0-2.5 mm long,
villous, terete, eglandular. Stipules linear, 4.0-7.0 mm long, villous, coriaceous,
persistent, intrapetiolar. Inflorescences terminal and axillary racemose panicles, the
rachis and branches pilose. Flowers ca 3.0 mm long, solitary and sessile on primary
branches of inflorescence. Bracts and bracteoles I.0-2.0 mm long, oblong-lanceolate,
persistent, to half length of flowers, pilose on exterior. Receptacle campanulate,
sessile, gray-tomentellous on exterior, tomentellous within. Calyx lobes acute,
puberulous on both surfaces.Petals absent. Stamens 5-7, inserted in a complete circle;
filaments shorter than calyx lobes, pilose below, glabrous above, free almost to base.
Ovary inserted at base of receptacle, pilose. Style equalling filaments, lanate. Fruit
unknown.
TYPE.Klug 449, Peru, Loreto, fl (holotype, F; isotypes, NY, US).
DISTRIBUTION. Known only from the type, collected in forest near Iquitos in
October.
This poorly known species is apparently quite distinct from any other Licania.
The leaf shape is uncommon and few other species have pilose-villous leaves and
inflorescences.
136 Flora Neotropica

102. Licania cordata Prance, sp nov Fig 20 D-G.

Frutex vel arbor parva, ramulis juvenilibus hispidis mox glabris lenticellatis.
Folia alternata, breviter petiolata; petiolo 0.5-1.5 mm longo, piloso-tomentoso, tereti,
eglanduloso; laminae ovatae, coriaceae, 2.8-6.5 cm longae, 1.4-4.0 cm latae, basi
cordatae, apice in acuminem 3.0-7.0 mm longum contractae, supra glabrae, subtus
lanato-pubescentes, reticulatae; costa media supra villosa, leviter impressa; costis
secundariis 7-10 jugis, subtus prominentibus, pilosis vel hispidis, supra planis.
Stipulae lineares, ad 2.0 mm longum, pilosae, extrapetiolares, persistentes. Flores
circa I.5 mm longi, haud pedunculati, in spicis terminalibus vel axillaribus dispositi,
rachi hispido. Bracteae bracteolaeque I.0-2.0 mm longae, ovatae, persistentes, extus
tomentosae. Receptaculum campanulatum, sessile, extus tomentosum. Calycis-lobi
acuti, utrinque tomentelli. Petala nulla. Stamina 5-6, in orbem completum vel 3/4
disposita; filamenta glabra, ad basim libera, calycis-lobis breviora. Ovarium ad
basim receptaculi insertum, villosum. Stylus e basi ovarii ortus, ad apicem pubescens,
calycis-lobis brevior. Drupa immatura ovoidea, extus rufo-tomentosa.
TYPE.Maguire& Wurdack 34505, Venezuela, Terr. Amazonas, Cerra Yapacana,
fl (holotype, NY).
Known only from the savannas around Cerro Yapacana, where
DISTRIBUTION.
it is reported to be abundant. Flowering November to March.
VENEZUELA. Terr. Amazonas: Cerro Yapacana, Maguire,Cowan & Wurdack30497 fr (NY);
30785 fl (NY); Maguire, Wurdack& Bunting36615 fl (NY); Maguire, Wurdack& Keith 41487 st (NY).

Licaniacordatais distinct most of all by its small ovate leaves with cordate bases
as well as by the hispid stems. These characters distinguish it from L. hebantha,to
which it is most closely related.

103. Licania foldatsii Prance, sp nov Fig 20 H-M.

Frutex vel arbor parva, ramulisjuvenilibus tomentosis. Folia alternata petiolata;

petiolo 8.0- 2.0 mm longo, tereti, eglanduloso, tomentoso; laminae ovato-orbiculares
vel ovatae, crasse coriaceae, 5.5-I4.0 cm longae, 3.5-10.0 cm latae, basi saepissime
subcordatae vel raro rotundatae, apice acutae, rotundatae vel retusae, supra glabrae,
subtus dense lanato-arachnoideo-pubescentes, profunde reticulatae; costa media
supra plana vel leviter impressa, praeter basim glabra; costis secundariis 8-13 jugis,
subtus prominentibus, supra planis. Stipulae 4.0-7.0 mm longae, lanceolatae, persis
tentes, coriaceae, cum petiolo adnatae. Flores circa 5.0 mm longi, haud pedunculati,
in paniculis racemosis sparse ramosis terminalibus subterminalibusque dispositi,
rachi ramisque brunneo-tomentosis. Bracteae bracteolaeque acutae, ad 4.5 mm
longae, persistentes, extus tomentosae. Receptaculum urceolatum, sessile, extus
tomentosum, intus tomentosum. Petala nulla. Stamina 3, unilateralia; filamenta
glabra, ad basim libera, calycis-lobis breviora. Ovarium ad basim receptaculi
insertum, pilosum. Stylus e basi ovarii ortus, calycis-lobis brevior, ad apicem pubes-
cens. Drupa globosa vel pyriformis, ad 4.0 cm in diametro, extus glabra, rugosa in
sicco; mesocarpio tenui; endocarpio 3.0-4.0 mm crasso, duro, fibroso, intus dense
hirsutulo.
TYPE. Foldats 3857, Venezuela, Terr. Amazonas, Rio Atabapo, fl (holotype,
VEN 49613; isotype, NY).
DISTRIBUTION. Savannas and river margins in the upper Orinoco and Rio Negro
region of Venezuela. Flowering September to November.
 Licania 137

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~~~,::,,, ~ ",.."' '

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....ml ,,

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FIGao ~ ~ odt~ -C,?

peis fL~a~a L.snw~h
' R.Bya ), .5;Bhabt,x
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flower -,L s
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.5 aitE,flwe,~ o;F
,yugfut
seto,~~o xa -,L l3~,355
odt~(odr
flower'' 33,35) ,hbt .;
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FIG 20. Species of Licania.A-C, L. sandwithii(R. Boyan92); A, habit, X 0 .5; B, flower, x i o; C,

flower section, xi o. D-G, L. cordata(Maguireet al 30497, 34505); D, habit, Xo.5; E, flower, xi 0; F,
flower section, x ixo; G, young fruit, X 2. H-M, L, foldatsii (Foldats3633, 3857) ; H, habit, x 0 .5; J,
flower, x 7.5; K, flower section, x 7.5; L, fruit, x o.5; M, fruit section, x 0.5.
138 Flora Neotropica

VENEZUELA. Terr. Amazonas: Rio Atabapo, Foldats 3709 fl (NY, VEN 49600); 3633 fl
(NY, VEN 49602); LI. Williams 13848 fr (US); 13849 fr (VEN 9763); Rio Guainia, LI. Williams
I4913 fr (F, US, VEN); Yavita, LI. Williams 14077 fr (F, US, VEN 2198); 4II19 fr (F, VEN).

LOCALNAME.Guayabito.
Licaniafoldatsiiis nearest L. subrotundata,
but is distinguished from this and related
species by the large flowers and stipules.

104. Licania hebantha Martius ex Hooker f., Mart. Fl. Bras. I4(2): 7. 1867;
Fritsch, Ann. Naturh, Mus. Wien 4: 49. I889.

Shrub or small tree, the young branches puberulous, soon becoming glabrous and
lenticellate. Leaves ovate-orbicular or elliptic, submembraneous, 2.5-9.0 cm long,
2.2-5.0 cm broad, rounded to acute at apex, subcordate at base, glabrous above,
with appressed lanate pubescence beneath obscuring the deeply reticulate venation;
midrib slightly impressed above, hirsutulous towards base; primary veins 7-I3 pairs,
prominent beneath, plane or slightly impressed above; petioles 1.5-3.0 mm long,
terete, eglandular, tomentose. Stipules linear, 2.0-2.5 mm long, coriaceous, persistent.
adnate to extreme base of petiole. Inflorescences terminal and subterminal racemose
panicles, the rachis and branches tomentose. Flowers ca 3.0 mm long, sessile on
primary branches of inflorescence. Bracts and bracteoles 0.5-I.5 mm long, elliptic,
persistent, tomentose on exterior. Receptacle campanulate, sessile, tomentose on
exterior, lanate-pilose within. Calyx lobes acute, tomentose on exterior, puberulous
within. Petals absent. Stamens 5-6, unilateral; filaments shorter than calyx lobes,
glabrous, free to base. Ovary inserted at base of receptacle, pilose. Style equalling
filaments, pubescent throughout. Fruit unknown.
TYPE.Martiussn, Colombia, Amazonas, fl (holotype, M; isotype, fragment K).
DISTRIBUTION. Savannas in Amazonian Colombia.
COLOMBIA. Vaup6s: Schultes5453 fl (COL, K, US). Amazonas: Maguire,Maguire & Ferndndez
44I56 fl (NY).

105. Licania steyermarkii Maguire, Fieldiana Bot. 28: 254. I952.

Tree to 25.0 m tall, the young branches puberulous, becoming glabrous and
lenticellate with age. Leaves broadly ovate to ovate-orbicular, thick-coriaceous,
4.5-8.5 cm long, 2.5-6.o cm broad, acute or short-caudate at apex, rounded or
slightly subcordate at base, glabrous above, appressed arachnoid-lanate below, and
with shallowly reticulate venation; midrib plane above, glabrous except sometimes
sparsely pubescent at extreme base; primary veins 5-6 pairs, prominent beneath,
plane or prominulous above; petioles 3.0-5.0 mm long, sparsely puberulous-hirsutu-
lous, eglandular, shallowly canaliculate or terete. Stipules linear, coriaceous, persis-
tent, adnate to extreme base of petiole. Inflorescences terminal and subterminal
racemose panicles, the rachis and branches tomentellous to tomentose. Flowers ca
2.0 mm long, sessile on primary branches of inflorescence. Bracts and bracteoles
0.4-1.5 mm long, ovate, persistent,tomentellous on exterior. Receptacle campanulate,
sessile, tomentose on exterior, tomentellous within. Calyx lobes acute, tomentellous
on both surfaces. Petals absent. Stamens 3, unilateral; filaments shorter than calyx
lobes, glabrous, free to base. Ovary inserted at base of receptacle, pilose. Style
equalling filaments, pilose almost to apex. Fruit unknown
Licania 139

TYPE.Steyermark 60768, Venezuela, Bolivar, fl (holotype, NY; isotypes, F, MO,

VEN); 60008 st (paratypes, F, NY); 60465 fl (paratypes, F, NY, US, VEN).
DISTRIBUTION. Forests in Venezuela and Colombia. Flowering September to
December.
COLOMBIA.Vaupes:Schultes& Cabrera
13846fl (BM, F, GH, LE, NY, US). VENEZUELA.
Bolivar: Bernardi2737 fl (K, NY, VEN).

LOCAL NAMES. Venezuela: Macara-yek, Hierrito.

io6. Licania subrotundata Maguire, Fieldiana Bot. 28: 255. I952.

Small tree, the young branches puberulous, becoming glabrous and lenticellate.
Leaves broadly ovate to ovate-orbicular, thick-coriaceous, 4.0-I2.0 cm long, 3.0-7.5
cm broad, rounded to acute at apex, cordate to rounded at base, glabrous on upper
surface, the lower surface with appressed lanate farinaceous pubescence; venation
conspicuous but not deeply reticulate; midrib glabrous above, slightly impressed;
primary veins 6-9 pairs, prominent beneath, prominulous above; petioles 4.0-6.0 mm
long, glabrescent or glabrous, shallowly canaliculate, with two sessile glands at junc-
tion with upper surface of lamina. Stipules 1.0-3.0 mm long, linear, coriaceous,
persistent, extrapetiolar. Inflorescencesterminal and subterminal panicles, the rachis
and branches puberulous. Flowers ca 3.0 mm long, sessile on primary and secondary
branches of inflorescence.Bracts and bracteoles lanceolate, persistent, to 4.0 mm long,
puberulous on exterior. Receptacle campanulate, sessile, tomentellous on exterior,
tomentellous within. Calyx lobes acute, tomentellous on both surfaces. Petals absent.
Stamens 3, unilateral; filaments shorter than calyx lobes, glabrous, free to base.
Ovary inserted at base of receptacle, pilose. Style equalling calyx lobes, sparsely
pubescent throughout. Fruit globose to pyriform, to 3.5 cm in diameter; epicarp
furfuraceous, drying yellow and wrinkled; mesocarp very thin; endocarp 2.0 mm
thick, hard, granular, hirsutulous within.
TYPE. Steyermark 56917, Venezuela, Dist. Federal, fr (holotype, NY; isotypes,
F, VEN).
DISTRIBUTION. Confined to the highland cloud forest of the Federal District of
Venezuela. Flowering November to March.
VENEZUELA. Dist. Federal: Delgado I20 fl fr (F, US, VEN); 434 fr (VEN); 445 fr (VEN);
et al 90865 fr (NY, US, VEN); 9587i fl (NY, VEN); Tamayo144 fl fr (US, VEN).
Steyermark

LOCALNAME. Icaquito.
Licania subrotundata appears to be endemic to the mountains in the Federal
District of Venezuela, and is there confined to the cloud forest. Growing at about
2,000 m altitude, it is one of the very few truly montane members of the Chryso-
balanaceae.

107. Licania crassivenia Spruce ex Hooker f., Mart. Fl. Bras. 14(2): 14.
1867; Fritsch, Ann. Naturh. Mus. Wien 4: 49. 1889.

Large shrub, the young branches glabrous, obscurely lenticellate. Leaves ovate-
lanceolate, coriaceous, 5.0-8.0 cm long, 1.5-3.8 cm broad, tapering to a fine point at
140 Flora Neotropica

the apex, with acumen 3.0-7.0 mm long, subcuneate at base, glabrous above; lower
surface with well developed stomatal cavities which are filled by lanate pubescence,
the venation glabrous and hence the cavities conspicuous; midrib slightly impressed
above, glabrous; primary veins 7-IO pairs, prominent beneath, impressed above;
petioles 8.o-i2.o mm long, glabrous, shallowly canaliculate, eglandular. Stipules to
2.0 mm long, lanceolate, puberulous, caducous. Inflorescences terminal racemose
panicles, the rachis and branches gray-puberulous. Flowers ca 3.0 mm long, sessile on
primary branches of inflorescence. Bracts and bracteoles to 2.0 mm long, lanceolate,
persistent, puberulous. Receptacle campanulate, sessile, gray-puberulouson exterior,
lanate within. Calyx lobes acute, puberulous on both surfaces. Petals absent. Stamens
5-7, unilateral; filaments shorter than calyx lobes, glabrous, free to base. Ovary
inserted at base of receptacle, pilose. Style rising to base of filaments, lanate. Fruit
unknown.
TYPE.Spruce2678, Brazil, Amazonas, fl (holotype, K; isotypes, BM, BR, CGE,
OXF, P).
DISTRIBUTION. Known only from the type, collected in the caatingas of the Rio
Vaupes.
Licaniacrassiveniais easily recognized by the leaves with impressed venation on
the upper surface, and with deep stomatal cavities beneath, as well as by the petioles
which are longer than in most related species.

Io8. Licania majuscula Sagot, Ann. Sci. Nat. Bot. VI. 15: 306. 1883; Fritsch,
Ann. Naturh. Mus. Wien 4: 43. 1889; Benoist, Bull. Mus. Hist. Nat. Paris
25: 516. I9I9.
LicaniahostmanniiFritsch, Ann. Naturh. Mus. Wien 4: 42. 1889; Kleinh. in Pulle, Fl. Suriname
2(I):432. 1939, synon nov. Type. Hostmann& KapplerI250, Suriname, fl (holotype, W; isotypes,
BM, C, CGE, F, K, LE, NY, P, S, U).

Tree to 22.0 m tall the young branches tomentose becoming glabrous with age.
Leaves elliptic to obong-ovate, coriaceous, 7.5-18.0 cm long, 2.8-9.0 cm broad,
cuspidate to acuminate at apex, with acumen 2.0-12.0 mm long, rounded at base,
glabrous above, the venation deeply reticulate on lower surface, revealing open
stomatal cavities or deep reticulations, lanate between the venation or in the mouth
of the cavities; midrib slightly impressed above for entire length; primary veins 8-Io
pairs, prominent beneath, slightly impressedabove; petioles 8.0-I 3.0 mm long, lanate,
usually deeply canaliculate, with two or more pairs of sessile, medial glands. Stipules
linear-lanceolate, 3.0-8.0 mm long, adnate to base of petioles, persistent. Inflores-
cences terminal and axillary racemose panicles, the rachis and branches tomentose.
Flowers 4.0-5.0 mm long solitary or in small groups, sessile on primary and secon-
dary branches of inflorescence. Bracts and bracteoles I.O-3.5 mm long, oblong to
lanceolate, persistent, entire, eglandular, tomentose on exterior. Receptacle conical,
sessile, tomentose on exterior, villous-tomentose within. Calyx lobes acute, tomentose
on both surfaces. Petals absent. Stamens 8-I i, inserted in a complete circle; filaments
equalling calyx lobes, slightly connate at base, glabrous except for deflexed hairs at
base. Ovary inserted at base of receptacle, pilose. Style equalling filaments, lanate
throughout. Fruit globose when mature, 5.0-6.0 cm diameter, with short stipe,
pyriform when young; epicarp velutinous-pubescent; pericarp thick, undifferen-
tiated, fibrous, densely pubescent within.
TYPE.Melinonsn (1862), French Guiana, fl (holotype, P; isotypes, BM, GH, K,
NY, P, US).
Licania 141

Upland forest and high riverine forest in the Guianas. Flowering

DISTRIBUTION.
August to November.
GUYANA. Davis I5o(F.D. 2141) fl (K); 3I5(F.D. 2308) fl (K, NY, U); 433(F.D. 2438) fl
(FHO, K); De La Cruz 2032 fl (NY); Fanshawe127(F.D. 2736) fl (FHO, K, U); 9o4(F.D. 3640) fr (U);
3515(F.D. 71II) fl (K, NY); Sandwith1236 fl (K, NY, S, U); A. C. Smith3095 fr (F, GH, IAN, LE,
MO, NY, P, S, U, US); Wilson-Browne 224(F.D. 5701) fl (K, NY); 3o6(F.D. 5800) fl (K, NY). SURI-
NAME. B. W. 628 fl (U); 1699 st (U): 1738 st (U); 1748 st (U); 3999 fl (MO, NY, U); 6263 fl (U);
6455 fl (IAN, K, NY, U); Irwin, Prance,Soderstrom& Holmgren54564 fl (F, MO, NY, US); 54785 fl
(F, GH, NY, US); 54937 fl (F, GH, NY, US); 54988 fl (F, GH, MO, NY, P, US); 55387 fl fr (F, GH,
NY, P, US); 55667 fl (F, GH, NY, US); 55769 fl (F, MO, NY, US); Lanjouw911 fr (NY); Schulz
1034a fl (BBS, U); van Emdensn fl (IAN, K, NY). FRENCH GUIANA. Benoist I80 fl (P); 1089 st
(P); For. Serv.62M fl (P, U); 77M fl (P, U); 7440 st (P, U); 7485 st (P, U); Gandoger
95 fl (P); Wachen-
heimsn fl (BM, F, K, US); 23 st (P); i80 st (P).

LOCALNAMES.Guyana: Kauta, Kautaballi. Suriname: Vonkhout,Foengoe,Bokohoko-

kandiamaroe, Marishiballitataro.French Guiana: Koko-Baaka.
When he described L. hostmannii, Fritsch proposed two forms based on the single
type gathering and distinguished by leaf shape and size. There is considerable range
in leaf form offering no grounds for subspecific taxa. Fritsch had not seen material
of L. majuscula,
whose type is matched by that of L. hostmannii.

o09. Licania alba (Bernouilli) Cuatrecasas, Contr. U. S. Natl. Herb. 35(6):

588. I964.

Theobroma alba Bernouilli, Denkschr. Schw. Naturf. Ges. 24(3): 14. 1871.
Licania venosaRusby, Descr. New S. Am. P1. 26. I920. Type. Rusby & Squires423, Venezuela,
Terr. Delta Amacuro, fl (holotype, NY; isotypes, A, BM, F, GH, MICH, MO, S, US,
VEN).
LicanialongifoliaR. Benoist, Bull. Mus. Hist. Nat. Paris 28: 252. I922. Type collections. Richard
sn, French Guiana, fl (syntype, P); Melinon202 st (syntype, P).

Tree to 35.0 m tall, the young branches pulverulent, becoming glabrous with age.
Leaves oblong-elliptic to elliptic, coriaceous, 9.0-27.0 cm long, 4.0-Io.o cm broad,
acuminate at apex, with acumen 3.0-Io.o mm long, rounded at base, glabrous above;
the lower surface with deep stomatal cavities rather obscured by dense lanate pubes-
cence; midrib impressed above for entire length, pulverulent near base but becoming
glabrous with age; primary veins 8-I2 pairs, prominent beneath, impressed and
glabrous above; petioles 9.0-I7.0 mm long, short-tomentellous-pubescent, canali-
culate, usually with medial glands. Stipules elliptic, to 7.0 mm long, subpersistent,
axillary. Inflorescences terminal and axillary racemose panicles, the rachis and
branches tomentellous-pubescent. Flowers 2.5-3.0 mm long, sessile on primary and
secondary branches of inflorescence. Bracts and bracteoles ovate, to 4.0 mm long,
persistent, often with paired glands at base, tomentellous on exterior. Receptacle
campanulate, sessile, tomentellous on exterior, tomentose within. Calyx lobes acute,
tomentellous on both surfaces. Petals absent. Stamens 6-8, inserted in a complete
circle; filaments shorter than calyx lobes, pubescent, free to base. Ovary inserted at
base of receptacle, villous. Style equalling filaments, lanate throughout. Fruit
pyriform, to 9.0 cm long, including stipe to I5.0 mm long; epicarp sordid ferrugine-
ous-brown pulverulent; mesocarp thin, very hard, sparsely hirsutulous within.
TYPE. Appun I, Guyana, st (K).
DISTRIBUTION. Primary forest in the Guianas and adjacent Venezuela and Brazil.
VENEZUELA. Terr. Delta Amacuro: Bernardi7505 fr (NY, VEN); Berti 8I fr (BR, NY); 172 st
(F, NY, US); i85 fr (F, US); Blanco36i fl fr (VEN); Breteler3850 st (NY, U, VEN); Steyermark
87r27
142 Flora Neotropica

fl (MICH, NY, US, VEN); Wurdack& Monachino39569 fr (F, GH, IAN, NY, P, RB, US). Bolivar:
Bernardi2140 fl (VEN); 3030 st (VEN); 7169 fr (VEN); Steyermark 86684 st (VEN). GUYANA. Aitkin
56 fl (K); Davis 280(F.D. 2271) fl (K); De La Cruz 2377 fl (F, GH, MO, NY, US); 4349 fl (F, GH, K,
MO, NY, US); 4384 fl (F, GH, K, MICH, MO, NY, US); For. Dept. 6268 st (BM, FHO, P); 6269 fl
(BM, FHO, P); Jenman2393 st (K); 3839 fl (K); 4012 fl (K, NY, US); 6627 fl (K); Maguire 40493 st
(F, K, NY, RB, US); Persaud13 fr (F, GH, K, NY); Sandwith261 fl (K, P, RB, U); 299 fl (K, NY, RB).
FRENCH GUIANA. Cowan38781 fl (NY, P). BRAZIL. Amazonas: Cavalcante739 fr (MG 23313);
Silva & Brazco 60705 fl (NY). Terr. Amapa: Frdes26706 fl (IAN, NY); Pires, Rodrigues& Irvine50557
fl (MG, NY).

LOCALNAMES. Venezuela: Hierrito,Hierro.Guyana: Kautaballi.

but is consistently different
This common species is closely related to L. majuscula,
in a number of important characters: the campanulate receptacle, the fewer stamens,
the white pubescence with deeper stomatal cavities, and the long stipe of the fruit all
separate L. alba from L. majuscula.

Ino. Licania hitchcockii Maguire, Mem. N. Y. Bot. Gard. 9: 477. 1957.

Small tree, the young branches puberulous, becoming glabrous with age. Leaves
ovate-elliptic to oblong-elliptic, thick-coriaceous, 2.5-5.5 cm long, 1.5-3.0 cm broad,
acute or bluntly acuminate at apex, with acumen 2.0-5.0 mm long, rounded at base,
glabrous above, the lower surface densely lanate-farinaceous;midrib impressed above
for entire length, glabrous except at base; primary veins 6-7 pairs, prominent
beneath, slightly impressed above; petioles 1.5-3.0 mm long, puberulous, axillary.
Stipules ca i.0 mm long, subpersistent, axillary. Inflorescences short terminal and
subterminal axillary panicles, the rachis and branches puberulous. Bracts and
bracteoles to I.o mm long, ovate, persistent, puberulous on exterior. Flowers ca
3.0 mm long, sessile on primary branches of inflorescence. Receptacle sessile, with
tomentellous exterior, tomentose within. Petals absent. Stamens included. Ovary
inserted at base of receptacle, villous. Fruit to 3.5 cm long, pyriform; epicarp glab-
rous, pulverulent; pericarp thin, hard, sparsely hirsutulous within.
TYPE. Phelps & Hitchcock 407, Venezuela, Bolivar, fr (holotype, NY; isotype,
VEN).
DISTRIBUTION. Known from only two inadequate collections from forested slopes
in Venezuela.
VENEZUELA. Bolivar: Steyermark
88582 st (F, NY).

III. Licania sandwithii Prance, sp nov Fig 20 A-C.

Arbor magna, ramulis juvenilibus rufo-hirsutulismox glabris lenticellatis. Folia

alternata petiolata; petiolo 4.0-5.0 mm longo, dense hirsutulo, tereti, eglanduloso;
laminae ovato-orbiculares, crasse coriaceae, 6.o-II.5 cm longae, 4.0-7.5 cm latae,
basi rotundatae, apice rotundatae vel acutae, supra glabrae nitidae, subtus profunde
reticulatae, lanato-farinaceo-pubescentes; costa media supra leviter impressa, basim
versus hirsutula, subtus hirsutula; costis secundariis 7-9 jugis, subtus prominentibus,
hirsutulis, supra glabris impressis. Stipulae ellipticae, 3.0-5.0 mm longae, hirsutulae,
intrapetiolares, persistentes. Flores 3.5-5.0 mm longi, haud pedunculati, in paniculis
racemosis terminalibus axillaribusque dispositi, rachi ramisque rufo-tomentosis.
Bracteae bracteolaeque ovatae, ad 3.0 mm longum, persistentes, extus tomentosae.
Licania 143

Receptaculum lato-cupuliforme, sessile, extus dense rufo-tomentosum. Calycis-lobi

acuti, utrinque tomentosi. Petala nulla. Stamina 6-8, unilateralia; filamenta glabra,
ad basim libera, calycis-lobis breviora. Ovarium ad basim receptaculi insertum, vil-
losum. Stylus e basi ovarii ortus, villosus, calycis-lobis brevior. Fructus mihi ignoti.
TYPE.R. Boyan92(F.D.79I6), Guyana, Upper Mazaruni District, fl (holotype,
NY; isotype, K).
DISTRIBUTION. Known only from the type gathering from primary forest in
Guyana.
Licaniasandwithiiis most closely related to L. laxifloraand L. couepiifolia.It differs
in the ovate-orbicular leaves with a blunt apex, and in the cupuliform receptacle;
the flowers are larger than those of L. laxiflora,and smaller than those of L. couepiifolia.
It is with pleasure that I name this species for the late Mr. Noel Sandwith, one of the
outstanding students of the South American flora, who was always so helpful to me.

112. Licania laxiflora Fritsch, Ann. Naturh. Mus. Wien 4: 46. I889.
Licaniagracilis Kleinhoonte, Rec. Trav. Bot. N6erl. 22: 382. I925. Type. B. W.379I, Suriname,
fl (holotype, U; isotypes, K, NY).

Tree to 30.0 m tall, the young branches puberulous or rufous-hirsutulous,be-

coming glabrous with age. Leaves ovate to elliptic, coriaceous, 5.0-22.0 cm long,
2.5-9.0 cm broad, acuminate or rarely acute at apex, with acumen 2.0-Io.o mm long,
rounded at base, glabrous above, the lower surface with deeply reticulate often
hirsutulous venation, with the area between veins lanate; midrib slightly impressed
above, tomentellous towards base; primary veins 7-10 pairs, prominent beneath,
impressed and glabrous; secondary veins more or less parallel; petioles 4.0-8.0 mm
long, densely rufous-tomentose, terete, eglandular or with medial glands. Stipules
elliptic to lanceolate, 2.0-5.0 mm long, pubescent, intrapetiolar, subpersistent.
Inflorescences terminal and subterminal racemose panicles, the rachis and branches
rufous-tomentose. Flowers 2.5-3.0 mm long, sessile on primary branches of in-
florescence. Bracts and bracteoles ovate, to 3.0 mm long, persistent, tomentose on
exterior. Receptacle campanulate, sessile, tomentose on exterior, tomentose within.
Calyx lobes acute, tomentose on both surfaces. Petals absent. Stamens 6-8, inserted
in a complete circle; filaments shorter than calyx lobes, glabrous, free to base. Ovary
inserted at base of receptacle, villous. Style equalling filaments, villous throughout.
Fruit round to pyriform, to IO.Ocm long including the short stipe to Io.o mm long;
epicarp rufous-velutinous; mesocarp thin, fleshy; endocarp thick, hard, sparsely
hirsutulous within.
TYPE.Schomburgk 976, Guyana, fl (holotype, W; isotypes, K, P).
DISTRIBUTION. Non-flooded forest and clump Wallaba forest in the Guianas and
Amazonian Brazil.
VENEZUELA. Bolivar: Bernardi7275 fl (NY). GUYANA. J. Boyan 48(F.D. 7732) fr (K, NY,
U); R. Boyan 44(F.D. 7868) fl (K, NY, U); Fanshawe ioig(F.D. 3755) fl (K, NY); Io2o(F.D. 3756) fl
(K, NY); Io32(F.D. 3768) fl (K, NY, U, US); ro59(F.D. 3795) fl (K, NY); Maguire 40533 fl (F, K,
NY, RB, US); Maguire & Fanshawe23511 fr (A, BR, F, K, MO, NY, US); Maguire,Maguire & Wilson-
Browne46247A fr (F, NY); Sandwith586 fl (K, NY, RB, U); 7z1 st (K); Schomburgk 54 fl (K). SURI-
NAME. Lanjouw& Lindeman2805 st (NY, U). BRAZIL. Amazonas: DuckeRB 35575 fr (RB); INPA
4883 st (NY); Mello & CoelhoINPA 4183 fl (INPA); Rodrigues& Coelho5572 fl (INPA 14244, NY).

LOCAL Venezuela: Hierrito.Guyana: Kauta.Suriname: Landhout,Kwepie-

NAMES.
rang.
144 Flora Neotropica

113. Licania rufescens Klotzsch ex Fritsch, Ann. Naturh. Mus. Wien 4: 52.
I889.
LicaniarufescensKlotzsch in Schomb., Faun. & Fl. Brit. Guian. 103. 1848 nom nud.

Tree to 35.0 m tall, the young branches puberulous. Leaves oblong-elliptic,

coriaceous, 5.0-Io.0 cm long, 1.8-5.5 cm broad, acuminate or acute at apex, with
acumen to 8.o mm long, subcuneate at base, glabrous above, with rufous-lanate
pubescence beneath obscuring deeply reticulate venation; midrib usually impressed
above, glabrous; primary veins 5-8 pairs, prominent beneath, usually slightly im-
pressed above; petioles 5.o-8.o mm long, tomentellous, shallowly canaliculate,
eglandular. Stipules lanceolate, to 3.0 mm long, intrapetiolar, pubescent, caducous.
Inflorescences terminal and subterminal racemose panicles, the rachis and branches
rufous-tomentellous. Flowers ca 2.0 mm long, sessile on primary branches of in-
florescence. Bracts and bracteoles 0.5-I.0 mm long, ovate, tomentellous on exterior,
persistent. Receptacle sessile, campanulate, tomentose on exterior, tomentose within.
Calyx lobes acute, tomentellous on both surfaces. Petals absent. Stamens 5-6, uni-
lateral; filaments shorter than calyx lobes, free to base, glabrous. Style rising to base
of filaments, pilose throughout. Fruit unknown.
TYPE. Schomburgk 6o0 (935B), Guyana, fl (holotype, W; isotypes, BR, CGE, K,
NY, P).
Primary forest on slopes and high ground in the Guianas and
DISTRIBUTION.
adjacent Venezuela and Brazil. Flowering September to December.
VENEZUELA. Terr. Delta Amacuro: Berti 244 st (NY); 426 st (VEN); Blanco 357 fl (VEN);
Zabala 33 st (VEN); 35 st (VEN). Bolivar: Pinkus 62 fl (F, GH, NY, S, US). GUYANA. Maguire,
Maguire & Wilson-Browne46I36A st (NY, US); P 5o(F.D. 2822) fl (FHO, K, U); Schomburgk 935 fl
(K); Wilson-Browne38o(F.D. 5794) fl (NY, US). SURINAME. Maguire 24795 fl (A, BR, F, K, LE,
MO, NY, US). BRAZIL. Para: A. silva 474 fl (IAN, NY).
LOCALNAMES.Venezuela: Hierillo, Hierrillo negro. Brazil: Cariperana.

114. Licania kunthiana Hooker f., Mart. Fl. Bras. 14(2): I6. 1867; Benoist,
Bull. Mus. Hist. Nat. Paris 25: 516. I919.
LicaniahypagyreaMalme, Ark. Bot. 23(4): I2. 1930, synon nov. Type. Malme 2378, Brazil, Mato
Grosso, fl (holotype, S; isotypes, LD, R).
Licania incanaauct., non. Aublet, Benth. Jour. Bot. Hooker 2: 220. 1840; Fritsch, Ann. Naturh.
Mus. Wien 4: 48. I889.
Licania parvifora Bentham var submembranacea Maguire, Fieldiana Bot. 28: 254. 1952. Types.
Steyermark62173, Venezuela, Monagas, fl (holotype, NY; isotypes, F, VEN); 6o093,
Venezuela, AnzoStegui, fl (paratypes, NY, US, VEN).

Tree to 25.0 m tall, the young branches puberulous, soon becoming glabrous and
lenticellate. Leaves oblong-ovate to oblong-lanceolate, membraneous to subcoriace-
ous, 3.0-8.5 cm long, .3-5.0 cm broad, acuminate at apex, with acumen 2.0-13.0 mm
long, rounded to cuneate at base, glabrous above, densely lanate-farinaceousbeneath;
upper surface of midrib plane or slightly impressed, glabrous except at extreme base;
primary veins 7-9 pairs, prominent beneath, plane above; secondary veins reticulate;
petioles 2.0-5.0 mm long, tomentellous to puberulous, becoming glabrous with age,
shallowly canaliculate to terete, eglandular. Stipules lanceolate, 2.0-3.0 mm long,
persistent, adnate to extreme base of petiole. Inflorescences terminal and axillary
racemose panicles, the rachis and branches sparsely puberulous. Flowers 1.5-2.0 mm
long, sessile and densely crowded on primary branches of inflorescence. Bracts and
bracteoles 0.2-0.8 mm long, triangular. Receptacle campanulate, sessile, sparsely
gray-puberulous or rarely tomentellous on exterior, tomentose within. Calyx lobes
acute, puberulous on both surfaces. Petals absent. Stamens 5-6, unilateral; filaments
Licania 145

shorter than calyx lobes, glabrous. Ovary inserted at base of receptacle, lanate. Style
lanate, equalling filaments. Fruit oblong-elliptical, up to 2.0 cm long, I.Ocm broad,
with a tapered base; epicarp yellow-brown-pulverulent, sometimes becoming glab-
rous with age; pericarp thin. hard, fibrous, hirsutulous within.
TYPE. Schomburgk 728, Guyana, fl (lectotype, K; isolectotypes, BM, BR, CGE,
GH, L, NY, OXF, P, US); Gardner5448, Brazil, Guanabara, fl (paratypes, BM,
CGE, K); Miers3814, Brazil, Guanabara, fl (paratypes, K, P).
DISTRIBUTION. A wide-ranging species from the Guianas to southern Brazil in
non-flooded forest, riverine forest, or secondary forest.
VENEZUELA. Bolivar: Bernardi1463 fl (NY). GUYANA. Fanshawe841(F.D. 3577) fl (K, NY,
U); 84iA(F.D. 3577A) fr (K, NY, P, U); 937(F.D. 3673) fr (K, NY, U); 2946 (F, D.6276) fl (A, K, NY,
S, U). FRENCH GUIANA. Bena 1299 fr (U); For. Serv.o109 fl (U); Leblond464 fl (P); Perrottetsn fl (P);
Poiteausn fr (K); Martin sn fl (K); Sagot 968 fr (BM, BR, GOET, K, NY, P, S, U); Talbot sn fr (K).
BRAZIL. Amazonas: Ducke 2137 fl (IAN, INPA, NY, R, RB); Frdes 26295 fl (IAN, NY); 33075 fl
(IAN); Poeppig2531 fl (LE); Pranceet al 4808 fr (INPA, NY); Rodrigues& Chagas1308 fl (IAN, INPA
7637). Terr. Roraima: Pranceet al 4624 fl (INPA, NY). Para: Archer8076 fi fr (F, IAN, K, NY, S);
Black 48-3475 fl (IAN); 54-17013 fl (NY, P, US); Black & Ledoux50-10233 fl (IAN); DuckeMG 4945
fl (BM, MG, P, RB, US); MG 15752 fl (BM, MG, P, R, US); RB 15173 fl (RB); RB 18821 fr (K);
KuhlmannRB 138217 fr (RB); Oliveira32 fl (IAN); 3136 fl (IAN); Pires 8121 fl (IAN, NY); 51884 st
(NY, UB); Prance& Silva 58750 fl (F, GH, NY, RB, UB); Prance& Pennington1564 fl (F, GH, IAN,
NY); 1932 fl (GH, NY); 2022 fr (GH, IAN, NY); Silva 1463 fl (IAN, NY). Terr. AmapA: Cowan38270
fr (NY, US). Maranhao: Froes 21 fl (US); 2004 fl (A, BM, F, K, MICH, MO, NY, P, S, US); 24250
fr (IAN); 25618 fl (IAN, NY); 25622 fl (IAN); TavaresrIIo fr (HST). Cear,: JNascimento 28 fl (RB
47227). Pernambuco: Lima 65-4311 fr (IPA); Prance & Lima 4808 fr (NY); Tavares 1208 fr (HST);
Teixeira2573 fl (HST). GoiSs: Burchell7369 fl (GH, K, P); Glaziou 21114 fl (BR, K, LE, R); Maguire,
Pires, Maguire & Silva 56139 fl (F, NY); Prance & Silva 58643 fl (F, GH, NY, RB, UB); 5947I fr (F,
GH, NY, UB); 59614 fr (F, GH, MO, NY, UB). Minas Gerais: Heringer2584 fl (NY, RB); Mexia
5409 fr (F, GH, IAN, LA, MICH, MO, NY, S, U, US); 5460 fr (F). Rio de Janeiro: Andrade&
Emmerich55I fl (R 11873); Glaziou 738 fl (BR); 7603 fl (BR, F, K, LE, NY, P, R, S). Guanabara:
Pessoaldo HortoFlorestalRB 136926 fr (RB); RB 136932 fl (RB); RB 136934 fl (RB). Without locality,
Rio deJaneiro or Guanabara: Gaudichaud1170 fr (P); Riedelsn fl (NY, US). Sao Paulo: HandroSP 28197
fl (NY, SP); SP 31063 fr (SP); HoehneSP 27385 fl (NY, SP); SP 27563 fl (NY, SP); Kuhlmann& Kiihn
SP 66047 fl (SP).

LOCAL NAMES. Venezuela: Carbonero. Guyana: Unikiakia. French Guiana:

Gaulettegrigri. Brazil: Ascindiuva,Cariperana,Uilho torrado,Bafo de boi.
Although much confused with L. incanaand other species, L. kunthianais quite
distinct. It differs from L. incanain the sparse pubescence of the inflorescence and
flowers, the membraneous leaves, and the fruit shape. As specimens named L.
hypagyreafall well within the variation of this species, this name must therefore be
considered a synonym.

I 5. Licania bellingtonii Prance, sp nov Fig 21.

Arbor magna, ramulis juvenilibus sparse puberulis, mox glabris lenticellatis.

Folia alternata petiolata, petiolo 3.o-5.0 mm longo, sparse pubescenti, tereti,
eglanduloso; laminae oblongo-ellipticae, coriaceae, 6.o-8.5 mm longae, 2.0-4.0 cm
latae, apice in acuminem 6.o-Io.o mm longum contractae, basi cuneatae, supra
glabrae subtus cum cavis stomatalis vadosis munitae; costa media supra plana vel
basim versus leviter impressa, primo tomentella mox glabra; costis secundariis 8-io
jugis, supra planis, subtus prominentibus. Stipulae lanceolatae, 3.0-5.0 mm longae,
persistentes, sparse puberulae, cum basi petioli adnatae. Flores circa 3.0 mm longi,
haud pedunculati, in paniculis terminalibus subterminalibusque dispositi, rachi
ramisque sparse puberulis. Bracteae bracteolaeque o.2-I.O mm longae, ovatae,
persistentes. Receptaculum urceolatum, sessile, extus sparse puberulum, intus
146 Flora Neotropica

tomentosum. Calycis-lobi acuti, utrinque sparse tomentelli. Petala nulla. Stamina 5,

unilateralia; filamenta glabra, basi libera, calycis-lobis breviora. Ovarium ad basim
receptaculi insertum, tomentosum. Stylus e basi ovarii ortus, calycis-lobis brevior,
ad apicem lanato-pubescens. Fructus mihi ignoti.
TYPE. Prance& Ramos7000, Brazil, Terr. Rondonia, Santa Barbara, 15 km. east
of Km. 117 of Porto Velho to Cuiaba highway, fl (holotype, NY; isotypes, INPA,
US).
DISTRIBUTION. Known only from the type collection from non-flooded forest,
flowering in August.
This species is closest to L. kunthiana,but differs in the urceolate receptacle and
the stomatal cavities of the leaf undersurfaces. It is also close to L. pallida but differs
in the sessile flowers, the urceolate receptacle, and in the persistent stipules which are
adnate to the petiole.
I have named this species for Paul Bellington, missionary in Porto Velho, and
amateur naturalist, who suggested that we visit the region which includes the type
locality of this species, and who extended help in many ways during my field work
in Rondonia Territory.

II6. Licania compacta Fritsch, Ann. Naturh. Mus. Wien 4: 52. 1889.

Tree (?), the young branches tomentellous, becoming glabrous with age. Leaves
oblong, thick-coriaceous, 3.0-5.5 cm long, 1.5-3.2 cm broad, rounded to acute at
apex, rounded at base, glabrous above, the lower surface with dense brown-lanate
pubescence obscuring the deep stomatal cavities; midrib plane or slightly impressed
above, glabrous; primary veins 9-I pairs, prominent below, plane above; petioles
3.0-4.0 mm long, terete, eglandular, tomentellous when young. Stipules 3.0-4.0 mm
long, lanceolate, tomentellous when young, persistent, adnate to base of petiole.
Inflorescences terminal racemose panicles, the rachis and branches tomentellous.
Flowers 3.5-4.5 mm long, sessile on primary branches of inflorescence. Bracts and
bracteoles ovate, persistent, to 3.0 mm long, tomentellous on exterior. Receptacle
ureolate, sessile, tomentellous on exterior, tomentose within. Calyx lobes acute,
tomentellous on exterior, puberulous within. Petals absent. Stamens 4, unilateral;
filaments shorter than calyx lobes, glabrous. Ovary inserted at base of receptacle,
villous. Style equalling filaments, pubescent. Fruit unknown.
TYPE. Schomburgk519(812), Guyana, fl (holotype, W; isotypes, BR, CGE, F,
G, K, L, NY, OXF, P).
DISTRIBUTION. Known only from the type from the vicinity of Mt. Roraima on
the western frontier of Guyana.
This poorly known species is quite distinct from related species. It has stipules
similar to those of L. mollis,but differs from that species in leaf shape and in smaller
number of stamens.

17. Licania ovalifolia Kleinhoonte, Rec. Trav. Bot. Neerl. 30: I8o. i933;
Kleinh. in Pulle, Fl. Suriname 2(I): 442. 1939.
Licaniastahelii Kleinhoonte, Rec. Trav. Bot. Neerl. 30: i8i. 1933, synon nov; Kleinh. in Pulle.
Fl. Suriname 2(I): 442. I939. Type. B.W. 68i8, Suriname, fl (holotype, U; isotypes, IAN,
K, NY).

Tree, the young branches puberulous, soon becoming glabrous and conspi-
Licania 147

h^ A ; , 0 'U
?

'.
X

/^\\ s \ \ "'*-~~~~~~~...
.... .

FI ~ peis f iana ADLblngoii(Pae tal7oo; ,hai, o.;B,fowr

x~~
8;Clwrsct,x8D,oayx 5.EHL.mrness(iba46)E,vr,5
bu~~~~~~~d,x8C,flwrbdscinx8;.hai,xo5
flower

FIG 21. Species of Licania. A-D, L. bellingtonii(Pranceet al 7000); A, habit, x 0.5; B, flower,
x8; C, flower section, x8; D, ovary, x 15. E-H, L. maranhensis(Lisboa 2462); E, ovary, x i5; F,
flower bud, x8; G, flower bud section, x 8; H. habit, x 0o.5.
148 Flora Neotropica

cuously lenticellate. Leaves oval to elliptic, thick-coriaceous, 3.5-Io.0 cm long,

2.0-6.0 cm broad, the margins often recurved, usually rounded at apex but fre-
quently acute, rounded at base, glabrous above, the lower surface densely rufous-
lanate; midrib slightly impressed above, glabrous; primary veins 5-7 pairs, prominu-
lous beneath, plane or slightly impressed above; petioles 4.0-7.0 mm long, tomentel-
lous when young, usually eglandular, terete. Stipules lanceolate, 4.0-6.0 mm long,
persistent, pubescent when young, adnate to lower portion of petiole. Inflorescences
terminal and axillary racemose panicles, the rachis and branches rufous-tomentellous.
Flowers 3,0-4.0 mm long, sessile on primary branches of inflorescence. Bracts and
bracteoles 1.5-2.5 mm long, ovate, persistent, exterior tomentellous. Receptacle
urceolate, sessile, tomentose on exterior, tomentose within. Calyx lobes acute,
tomentellous on exterior, puberulous within. Petals absent. Stamens 3, unilateral;
filaments shorter than calyx lobes, glabrous. Ovary inserted at base of receptacle,
pilose. Style equalling filaments, pubescent throughout. Fruit unknown.
TYPE. B.W.6457, Suriname, fl (holotype, U; isotypes, IAN, K, NY).
DISTRIBUTION. Primary forest in the Guianas. Flowering January to May.
SURINAME. B.B.S. I027 st (U); IIo6 st (U); Burger21 fl (U); L.B.B. 9185 st (U); Lindeman
5209 st (NY, U). FRENCH GUIANA. For. Serv. 64M st (U); 7697 fl (P, U).

LOCALNAMES.Suriname: Foengoe, Zandhout. French Guiana: Bois Gaulette, Gris-

gris, Wetikoko.
Licaniaovalifoliais most closely related to L. micrantha,but differs in the leaves
with a blunt apex. See also note under L. micrantha.Kleinhoonte distinguished be-
tween L. ovalifoliaand L. stahelii by leaf shape: broadly oval in L. ovalifoliaand
narrowly oval in L. stahelii.The material now available shows continuous variation
from broad to narrow leaves. No additional characters were discovered. Conse-
quently, L. staheliiis reduced to synonymy here.

Ii8. Licania caldasiana Cuatrecasas, Fieldiana Bot. 27(2): 64. I950.

Tree, the young branches puberulous, becoming glabrous and lenticellate with
age. Leaves elliptic to oblong-elliptic, coriaceous, 5.0-10.0 cm long, 2.5-5.0 cm broad,
rounded or rarely acute at apex, rounded at base, glabrous above, densely gray-lanate
beneath; midrib plane above, glabrous, primary veins 9-II pairs, prominent be-
neath, plane above; petioles 4.0-7.0 mm long, shallowly canaliculate, eglandular,
tomentellous when young. Stipules caducous (not seen). Inflorescences terminal and
subterminal racemose panicles, the rachis and branches tomentellous. Flowers
1.5-2.0 mm long, densely crowded and sessile along primary branches of inflores-
cence. Bractsand bracteoleso.8-2. mm long, ovate, persistent,tomentose on exterior.
Receptacle globose, sessile, tomentose on exterior, and within. Calyx lobes acute,
tomentose on both surfaces. Petals absent. Stamens 3 (rarely 4), unilateral; filaments
shorter than calyx lobes, glabrous, free to base. Ovary inserted at base of receptacle,
villous-tomentose. Style rising to base of filaments, glabrous except for the villous base.
Fruit unknown.
TYPE. Mutis 3868, Colombia, fl (holotype, US; isotypes, F, MA); II14 fl
(paratypes, MA, US); 1130 fl (paratypes, MA, US).
DISTRIBUTION. Known only from the type gatherings, which lack detailed field
notes.
see notes and differences
Licaniacaldasianais most closely related to L. savannarum;
under that species.
 Licania 149

I 9. Licania savannarum Prance, sp nov Fig 22 D-H.

Frutex vel arbor parva, ramulis juvenilibus puberulis mox glabris lenticellatis.
Folia alternata petiolata, petiolo 2.0-3.0 mm longo, glabro, eglanduloso, rugoso,
tereti; laminae ellipticae vel oblongae vel oblongo-lanceolatae, coriaceae, 3.0-8.0 cm
longae, 1.2-3.5 cm latae, basi cuneatae vel subcuneatae, apice rotundatae vel
acutae, supra glabrae, subtus dense lanato-farinaceo-pubescentes;costa media supra
glabra, plana; costis secundariis 5-8 jugis, subtus leviter prominentibus, supra planis.
Stipulae ad I.o mm longae, lineares, caducae, extrapetiolares. Flores 2.5-3.0 mm
longi, haud pedunculati, in paniculis axillaribus terminalibusque dispositi, rachi
ramisque dense puberulis. Bracteae bracteolaeque 0.5-I.5 mm longae, lineares,
persistentes, extus tomentellae. Receptaculum urceolatum, sessile, extus rufo-brun-
neo-tomentosum, intus tomentosum. Calycis-lobi acuti, utrinque tomentosi. Petala
nulla. Stamina 3, unilateralia; filamenta glabra, calycis-lobis breviora, ad basim
libera. Ovarium ad basim receptaculum insertum, tomentosum. Stylus e basi ovarii
ortus, calycis-lobis brevior, ad medium hirsutus. Drupa (immatura tantum visi)
pyriformis, extus glabra, rugoso in sicco; mesocarpio tenui; endocarpio crasso,
lignoso, duro, intus hirsutulo.
TYPE. Maguire, Wurdack& Bunting37584, Venezuela, Terr. Amazonas, Rio
Pacimoni, fl (holotype, NY).
DISTRIBUTION. Savannas or open places by river margins in the upper Rio
Orinoco region of Venezuela and the Rio Negro region of Brazil.
VENEZUELA. Terr. Amazonas: Rio Pacimoni, Maguire, Wurdack& Bunting37604 fl (NY);
Cerro Yapacana, Maguire& Wurdack
34500 fl (NY); Rio Atabapo, Wurdack
& Adderley
42967 fl
(NY); Vareschi7998 fr (VEN). BRAZIL. Amazonas: Rio Curicuriary, Ducke RB 25030 fl (K); Rio
Cauburi,Silva & Brazao60791 fl (NY); 60966 fl (NY); 60976 fl (NY); Rio Urubu, Froes25240 fl
(IAN, NY, RB); Pranceet al 5050 fr (INPA, NY); Rodrigues300 fl (IAN, INPA 4410, MG, NY); Rio
Vaupes, Igarape Jurupari, Froes28220 fl (IAN, UB).

This species is most closely related to L. caldasiana,but differs in the narrower

leaves with fewer primary veins, the shorterglabrous terete petioles, and the urceolate
receptacle.

120. Licania microphylla Fanshawe & Maguire, Bull. Torrey Club 75: 322.
I948.

Tree, the young branches puberulous soon becoming glabrous and lenticellate.
Leaves ovate-elliptic, submembraneous, 3.0-4.0 cm long, I.0-2.0 cm broad, acumi-
nate at apex, with acumen 3.0-7.0 mm long, rounded at base, glabrous above, lower
surface with dense lanate pubescence which is easily removed; midrib plane above,
glabrous; primary veins 5-6 pairs, prominulous beneath, plane above; petioles 2.0-
4.0 mm long, puberulous, terete, with two large sessile glands at junction with upper
surface of lamina. Stipules 2.0 mm long, linear, puberulous, persistent, axillary.
Inflorescences short terminal and subterminal racemose panicles, the rachis and
branches puberulous. Flowers not seen. Bracts and bracteoles 0.5-I.0 mm long,
triangular, puberulous, persistent. Fruit pyriform, I.o cm long, with stipe 3.0-4.0 mm
long; epicarp densely pulverulent; pericarp thin, hard, fibrous, villous pubescent
within.
TYPE.FanshaweI078(F.D.38i4), Guyana, fr (holotype, NY; isotype, K).
DISTRIBUTION. Known only from the type gathering, collected in forest in the
Potaro River region of Guyana.
150 Flora Neotropica

LOCAL NAME. Unikiakia.

This poorly known species is distinct by its very small leaves and large petiolar
glands.

12I. Licania triandra Martius ex Hooker f., Mart. Fl. Bras. I4(2): i8. I867.
LicaniamicranthaMiquel var triandraFritsch, Ann. Naturh. Mus. Wien 4: 50. 1889.
Licania pulchraveniaKillip & Cuatrecasas, Fieldiana Bot. 27(2): io6. 1951, synon nov. Type
Cuatrecasas7054, Colombia, Vaup6s, fl (holotype, COL; isotype, fragments F).

Tree to 26.0 m tall, the young branches puberulous, soon becoming glabrous
and lenticellate. Leaves ovate-elliptic to oblong, thick-coriaceous, 7.0-I 7.0 cm long,
4.0-I0.0 cm broad, the margins plane, caudate-acuminate or acuminate at apex, with
acumen 5.0-I5.0 mm long, rounded at base, glabrous above, the lower surface with
deeply reticulate venation and poorly developed stomatal cavities, the venation
glabrous or glabrescent, hence conspicuous, usually with several conspicuous glands
on lower surface of lamina towards base; midrib plane above, glabrous; primary
veins 7-9 pairs, prominent beneath, plane above; petioles 5.0-9.0 mm long, puberu-
lous when young, soon becoming glabrous, terete, eglandular. Stipules 3.0-4.0 mm
long, lanceolate, glabrous, persistent, axillary. Inflorescences terminal and subter-
minal racemose panicles, the rachis and branches tomentellous. Flowers ca 2.0 mm
long, sessile on primary branches on inflorescence. Bracts and bracteoles 0.5-I.0 mm
long, triangular, persistent, puberulous on exterior. Receptacle campanulate, sessile,
tomentose on exterior, tomentose within. Calyx lobes acute, tomentellous on exterior,
puberulous within. Petals absent. Stamens 3, unilateral; filaments shorter than calyx
lobes, glabrous. Ovary inserted at base of receptacle, tomentose. Style equalling
filaments, pilose throughout. Fruit pyriform, to 3.5 cm long; epicarp short-
velutinous; pericarp thin, hard, fibrous, sparsely hirsutulous to glabrous within.
TYPES. Spruce 2490 (2409 some sets), Brazil, Amazonas, fl (lectotype, K; iso-
lectotypes, BR, C, CGE, GH, GOET, LD, LE, NY); Martiussn, Brazil, Amazonas or
Para, fl (paratype, M).
DISTRIBUTION. Non-flooded forest in Colombian and Brazilian Amazonia.
Flowering July to October.
COLOMBIA. Amazonas: Schultes& Cabrera12977 fl (NY, US); 16004 fr (BM, F, LE, NY, US);
i6977 fl (GH, NY, US). BRAZIL. Amazonas: Duarte7129 fl (INPA 14653); Froes21093 fl (IAN, NY).
Para: DuckeRB 19777 fl (K, RB).

Licaniatriandrais closely related to L. micrantha,

but in light of the recent material
it remains distinct. L. pulchravenia matches the type of L. triandrawell, the two types
even having glands at the base of the lower surface of the lamina. See also note under
L. micrantha.

X22. Licania discolor Pilger, Notizbl. Bot. Gart. Berlin 6: 137. I914.

Tree to 35.0 m tall, the young branches tomentellous, soon becoming glabrous
and lenticellate. Leaves oval to oblong, thick-coriaceous, 3.0-9.0 cm long, 1.5-4.5
cm broad, the apex with finely pointed acumen 4.0-12.0 mm long, subcuneate at
 Licania 151

base, glabrous above, densely lanate below; midrib slightly impressedabove, glabrous;
primary veins 7-9 pairs, prominent beneath, plane above; petioles 3.0-7.0 mm long,
tomentellous when young, terete, eglandular. Stipules to 3.0 mm long, lanceolate,
deciduous. Inflorescences terminal racemose panicles, the rachis and branches
tomentose. Flowers I.5-2.0 mm long, sessile on primary branches of inflorescence.
Bracts and bracteoles to i.o mm long, ovate, subpersistent, tomentellous on exterior.
Receptacle campanulate, tomentose on exterior and within; pedicels o-0.25 mm
long. Calyx lobes acute, tomentose on exterior, puberulous within. Petals absent.
Stamens 3, unilateral; filaments shorter than calyx lobes, glabrous. Ovary inserted at
base of receptacle, villous. Style rising to base of filaments, sparselypubescent through-
out. Fruit (immature) pyriform, with a short stipe; epicarp rufous-velutinous.
TYPE. Ule 8393, Brazil, Terr. Roraima, fl (holotype, B, lost; lectotype, K;
isotypes, L, LA, MG).
DISTRIBUTION. Forest on high ground, especially on slopes, in the Guianas and
adjacent Venezuela and Brazil. FloweringJuly to November.
VENEZUELA. Bolivar: Maguire, Wurdack& Bunting35890 fl (F, GH, MO, NY, US, VEN);
Steyermark86592 st (VEN). GUYANA. A. C. Smith 2270 fl (A, F, K, MO, NY, P, S, U, US); 2923 fr
(A, F, K, MO, NY, P, S, U, US); Wilson-Browne48o(F.D. 5886) fl (K, NY); 484(F.D. 5890) fl (K,
NY). SURINAME. Irwin ,Prance,Soderstrom& Holmgren54926 fl (F, GH, MO, NY); 55681 fl (F, MO,
928 fl (INPA, RB 4539,
NY, US). BRAZIL.Terr. Roraima:Black51-13885 fl (IAN, NY); Kuhlmann
S); Pranceet al 4210 st (INPA, NY); 4224 fr (INPA, NY); 4352 fr (INPA, NY); 9359 st (INPA, NY).

123. Licania apiculata Prance, sp nov Fig 22 J-L.

Frutex, ramulis juvenilibus glabris haud lenticellatis. Folia alternata petiolata;

petiolo 3.0-4.0 mm longo, primo glabrescente mox glabro, tereti, eglanduloso;
laminae oblongae vel oblongo-lanceolatae, coriaceae, 5.0-Io.o cm longae, 1.7-3.5
cm latae, basi cuneatae, apice apiculatae, supra glabrae nitidae, subtus dense lanato-
pubescentes; costa media supra plana, glabra; costis secundariis I0-I2 jugis, utrinque
leviter prominentibus. Stipulae lineares, ad 2.0 mm longae, glabrae, caducae, cum
basi petiolo adnatae. Flores 2.0-3.0 mm longi, haud pedunculati, in paniculis race-
mosis terminalibus dispositi, rachi ramisque puberulis vel tomentellis. Bracteae
bracteolaeque I.0-3.0 mm longae, lineares, parvae, persistentes, extus puberulae.
Receptaculum campanulatum, sessile, extus tomentosum, intus tomentellum. Calycis-
lobi acuti, utrinque tomentelli. Petala nulla. Stamina 3, unilateralia; filamenta glabra
ad basim libera, calycis-lobis breviora. Ovarium ad basim receptaculi insertum,
tomentellum. Stylus e basi ovarii ortus, ad apicem pubescens. Fructus (immaturus
tantum visi) extus sordido pubescens, pyriformis.
TYPE. Rodrigues& Coelho4881, Brazil, Amazonas, Rio Cuieiras, fl (holotype,
NY; isotype, INPA I34I4).
DISTRIBUTION. On beaches and open habitats beside rivers, known only from the
Rio Cuieiras in Brazil.
BRAZIL. Amazonas: Rio Cuieiras, Rodrigues& Wilson3970 fl (INPA o0535); 3998 fr (INPA
o0563).

This species is nearest L. micranthabut differs in the narrower apiculate leaves

and the greater number of primary leaf veins. It is also related to L. savannarum,
from
which it differs in the apiculate leaves with more primary veins, the campanulate
receptacle, and the less branched inflorescence.
152 Flora Neotropica

124. Licania micrantha Miquel, Stirp. Surin. Sel. 20. 1850; Hook. f., Mart.
Fl. Bras. 14(2): i8. 1867; Fritsch, Ann. Naturh. Mus. Wien 4: 50. 1889;
Kleinh. in Pulle, Fl. Suriname 2(I): 441. I939.
Licania helvolaSpruce ex Hooker f., Mart. Fl. Bras. 14(2): I8. I867, nom nud.
LicaniaanisophyllaStandley, Publ. Field Mus. Bot. 17: 253. 1937, synon nov. Type. Krukoff8451,
Brazil, Amazonas, fr (holotype, NY; isotypes, A, BM, BR, F, K, MO, NY, S, US).

Tree to 30.0 m tall, the young branches puberulous, soon becoming glabrous and
lenticellate. Leaves ovate-elliptic to oblong, thick-coriaceous, 4.5-I5.0 cm long, 2.3-
7.0 cm broad, margins plane, usually acuminate or caudate at apex, rarely acute,
with acumen 2.0-14.0 mm long, rounded at base, glabrous above, the lower surface
with dense brown lanate pubescence which is easily rubbed off; midrib more or less
plane above, glabrous; primary veins 5-7 pairs, prominent beneath, plane above;
petioles 4.0-12.0 mm long, puberulous when young, soon becoming glabrous, terete,
eglandular. Stipules lanceolate to 7.0 mm long, glabrous, persistent, adnate to ex-
treme base of petiole or axillary. Inflorescencesterminal and axillary much-branched
racemose panicles, the rachis and branches tomentellous to puberulous. Flowers
ca 2.0 mm long, sessile on primary branches of inflorescence. Bracts and bracteoles
0.5-I.0 mm long, ovate, persistent, exterior puberulous. Receptacle campanulate,
sessile, tomentellous on exterior, tomentose within. Calyx lobes acute, puberulous on
both surfaces. Petals absent. Stamens 3, unilateral; filaments shorter than calyx lobes,
glabrous. Ovary inserted at base of receptacle, pilose. Style equalling filaments,
pubescent throughout. Fruit pyriform, to 5.0 cm long, including stipe 5.0-10.0 mm
long; epicarp with waxy pulverulent indumentum, wrinkled when dry; pericarp thin,
hard, fibrous, densely hirsutulous within.
TYPE. HostmannI257, Suriname, fl (holotype, U; isotypes, F, GH, K, LE, NY,
P).
Non-flooded forest in the Guianas, Venezuela and Amazonia.
DISTRIBUTION.
COLOMBIA. Amazonas: Garcia-Barriga14463 fr (COL). VENEZUELA. Bolivar: Bernardi
i538 st (NY); Steyermark& Dunsterville93019 fl (NY, VEN); Steyermark60758 st (F, NY, US). Terr.
Amazonas: CardonaI479 fr (VEN); Maguire, Wurdack& Bunting365r9 fr (GH, NY, US). GUYANA.
De La Cruz 2739 fl (F, GH, MO, NY, US); Fanshawe204(F.D. 2940) fl (K); 324(F.D. 3060) fl (F, K,
NY, S, U); Jenman 1314 fl (K, U); Persaud4I pro parte fr (NY); 79 fr (F, K, NY). SURINAME.
B.B.S. 197 fr (K, U); B. W. 56 fl (U); I507 fl (U); i9i6 st (U); 2302 st (U); 3410 fl (A, K, MO, NY);
4058 fl (U); 4479 st (U); 5039 fl (A, U); 5149 st (U); 5385 st (U); 6008 fl (K, U); 6034 fl (NY);
6io8 fr (U); 6i86 st (U); Lanjouw& Lindeman401 fl (K, NY); Lindeman4599 st (NY, U); 4971 st (NY,
U); 6781 st (NY, U); Stahel80 fl (A, K, NY, U); 86 fl (A, K, NY, U). FRENCH GUIANA. For. Serv.
1o6 fl (U); 1174 fr (U). BRAZIL. Amazonas: Byron67-83 st (INPA, NY); ChagasINPA 246 fl (IAN,
INPA, NY); CoelhoINPA 4062 fl (INPA); Fr6es & Addison29076 fl (IAN, UB); Kuhlmann1004 fl
(INPA, NY, RB 17971, SP); Rodrigueset al 5436fl (INPA I4105); 5563 fl (INPA I4236, NY); 6928 fl
(INPA 1548I, NY); 6944 fl (INPA I5497, NY); 7245 fl (INPA I6179, NY); 7702 fl (INPA 17007);
Spruce2690 fl (BM, BR, CGE, F, GH, GOET, K, LD, LE, NY, OXF, P, S). Pari: Froes 32374 fl
(IAN); 32508 fl (IAN); Pires & Silva 10678 fl (IAN, NY); Pranceet al 4992 fr (INPA, NY); 51o8 fr
(INPA, NY); Silva 479 fl fr (NY); 496 fl (IAN, NY). Terr. Amapa: Froes 25882 fr (IAN, NY, RB);
26609 fl (IAN); 26660 fl (IAN, LE). Guanabara: Glaziou II943a cult fl (P, R).

LOCALNAMES.Venezuela: Macara-yek(Arekuna). Guyana: Marishiballi (Arawak).

Suriname: Foengoe, Man Foengoe,Braka Foengoe, Marishiballi,Santihoedoe.French
Guiana: Bois Gaulette.Brazil: Pintadinha,Macucuifarinha seca.
Licaniamicrantha, L. ovalifolia,and L. triandraare very closely related, but as they
do have a few correlated distinguishing characters, I maintain them as separate
species. Licaniatriandradiffers from L. micranthain the deeply reticulate venation of
the leaf undersurfacewith shallow stomatal cavities, the velutinous pubescence of the
fruit exterior, and the greater number of primary leaf veins (7-9). Licania ovalifolia
differs from L. micrantha in the leaves with an obtuse apex, the slightly recurved
Licania 153

C B

JL oke.r K L

FIG. 22. Species of Licania.A-C, L. riedelii(Riedel5z8); A, habit, x 0.5; B, flower section, x 7.5;
C, flower, x 7.5. D-H, L. savannarum(Maguire et al 34500, 37604); D, habit, x o.5; E, fruit, x 1.5; F,
fruit, transverse section, x 1.5; G, flower section, x 7.5; H, flower, x 7.5. J-L, L. apiculata(Rodrigueset
al 4881); J, habit, x o.5; K, flower, x 7.5; L, flower section, x 7.5.
154 Flora Neotropica

margins, and the larger stipules. I find no differences between L. micranthaand L.

anisophylla.

I25. Licania pruinosa R. Benoist, Bull. Mus. Hist. Nat. Paris 25: 5I6. I919.

Tree to 20.0 m tall, the young branches glabrous, conspicuously lenticellate.

Leaves oblong to oblong-elliptic, membraneous, 5.0-Io.o cm long, 2.0-5.0 cm broad,
acuminate at apex, with acumen 4.0- I.0 mm long, cuneate at base, glabrous above,
with short gray-brown lanate farinaceous pubescence beneath; midrib plane above,
glabrous; primary veins 6-8 pairs, prominent beneath, plane above; petioles 5.0-
6.0 mm long, terete, glabrous, eglandular. Stipules linear, ca 3.0 mm long, glabrous,
persistent, axillary. Inflorescences terminal racemose panicles, the rachis glabrescent,
branches puberulous to tomentellous. Flowers 3.5-5.0 mm long, on primary branches
of inflorescence. Bracts and bracteoles 0.2-0.5 mm long, triangular, persistent,
puberulous on exterior. Receptacle urceolate, sessile, tomentellous on exterior,
hirsute within. Calyx lobes acute, puberulous on both surfaces. Petals absent.
Stamens 3, unilateral; filaments shorter than calyx lobes, glabrous. Ovary inserted at
base of receptacle, pilose. Style equalling filaments, hirsute. Fruit unknown.
TYPE.Richardsn, French Guiana, fl (holotype, P).
DISTRIBUTION. River and island margins in French Guiana and Terr. Amapa
of Brazil. Flowering August to October.
BRAZIL. Terr. Amapa: Irwin,Pires & Westra47836 fl (F, IAN, MG, NY, RB); 47914 fl (F,
GH, IAN, MG, NY); 48122 fl (IAN, MG, NY); 48344fl (F, GH, IAN, MG, NY).

126. Licania nitida Hooker f., Mart. Fl. Bras. 14(2): I 7. I867.
Benthamvar nitida(Hookerf.) Fritsch,Ann. Naturh.Mus. Wien 4: 49. I889.
Licaniacoriacea

Shrub or low tree, the young branches glabrous, conspicuously lenticellate.

Leaves oblong to oblong-elliptic, thick-coriaceous, 4.5-10.5 cm long, 2.0-5.5 cm
broad, acute to bluntly acuminate at apex, with acumen to Io.o mm long, rounded to
subcuneate at base, glabrous above, densely lanate beneath; midrib plane above,
glabrous; primary veins 6-8 pairs, prominent beneath, prominulous above; petioles
4.0-7.0 mm long, glabrescent or glabrous, terete to flattened, eglandular. Stipules
linear, ca 3.0 mm long, glabrous, persistent, adnate to extreme base of petiole.
Inflorescences terminal racemose panicles, the rachis and branches tomentellous.
Flowers 4.0-5.0 mm long, on primary branches of inflorescence.Bracts and bracteoles
0.8-1.5 mm long, linear, persistent, tomentellous on exterior. Receptacle urceolate,
sessile, tomentellous on exterior, tomentose within. Calyx lobes acute, tomentellous
on exterior, puberulous within. Petals absent. Stamens 3 (-4), unilateral; filaments
shorter than calyx lobes, glabrous. Ovary inserted at base of receptacle, pilose. Style
equalling filaments, pilose. Fruit pyriform, ca 3.0 cm long, on stipe ca 8.o mm long;
epicarp sordid-pulverulent; pericarp thin, hard, fibrous, tomentose within.
TYPE. Blanchet3290, Brazil, Bahia, fl (holotype, BM; isotypes, BR, F, G, K,
LE, NY, P, W).
Eastern-central Brazil.
DISTRIBUTION.
BRAZIL. Minas Gerais: Kuhlmann174 fr (RB 136933); Netto 242 fl (GH, R); St. Hilaire Bi967
fl (P); C499 fl (P). Guanabara: AlmeidaRB 136929 fl (RB); RB 136931 fl (RB). Sao Paulo: Burchell
3235 fl (GH, K, NY, P, US); Lifgren 4180 fl (SP); Mosen2837 fl fr (C, LD, LE, R, S).
Licania 155

Fritsch treated this species as a variety of L. coriacea,but there are many important
differences between L. nitidaand L. coriacea,especially in the indumentum of the leaf
undersurface and the flower size. Licanianitidais most closely related to L. pruinosa,
but differs in the blunter leaf apices, the more pubescent inflorescence, and the
coriaceous leaves. It is also related to L. spicatawhich differsin the spicate unbranched
inflorescence, the globose receptacle, and the large number of stamens.

127. Licania riedelii Prance, sp nov Fig 22 A-C.

Frutex, ramulis juvenilibus puberulis, mox glabris lenticellatis. Folia alternata

petiolata; petiolo 3.0-5.0 mm longo, tereti, eglanduloso, puberulo; laminae ellipticae,
crasse coriaceae, basi rotundatae, apice acutae, supra glabrae nitidae, subtus cum
cavis stomatalis faucis lanato-pubescentibus, venulis glabris conspicuis; costa media
supra impressa, primo tomentosa; costis secundariis 9-I o jugis, subtus prominentibus,
supra planis, vel leviter impressis. Stipulae lineares, 1.5-2.0 mm longae, persistentes,
pubescentes, cum basi petioli adnatae. Flores 2.5-3.0 mm longi, haud pedunculati,
in paniculis densis racemosis terminalibus axillaribusque dispositi, rachi ramisque
ferrugineo-pubescentibus. Bracteae bracteolaeque 0.5-2.0 mm longae, ovatae,
persistentes, extus ferrugineo-pubescentes.Receptaculum campanulatum, subsessile,
extus ferrugineo-tomentosumintus tomentosum. Calycis-lobi acuti, utrinque tomen-
telli. Petala nulla. Stamina 5, unilateralia; filamenta glabra, basi libera, calycis-lobis
breviora. Ovarium ad basim receptaculi insertum, tomentosum. Stylus e basi ovarii
ortus, calycis-lobis brevior, ad apicem pubescens. Drupa pyriformis, 1.5 cm longa,
extus tomentella; mesocarpio 2.0-3.0 mm crasso; endocarpio tenui, duro, intus
pubescente.
TYPE. Riedel 508, Brazil, Minas Gerais, Serra da Caraca, fl (holotype, NY;
isotype, US).
DISTRIBUTION. Forested region of East Central Brazil.
BRAZIL. Minas Gerais, Serra da CaraCa,Langsdorff sn, fl (LE); Ouro Preto, Gomes2409 fr
(RB 47988); RB 47989fl (RB). Rio de Janeiro: Serrade Petropolis,Vit6rio
RB 48468fr (NY, RB).

Licania riedeliiis easily distinguished from related species by the conspicuous

stomatal cavities of the leaf undersurface.

128. Licania bracteata Prance, sp nov Fig 23 K-M.

Arbor magna, ramulis juvenilibus puberulis, mox glabris lenticellatis. Folia

alternata petiolata; petiolo 7.0-10.0 mm longo, juvenili tomentello, tereti, eglandulo-
so; laminae ellipticae vel oblongo-ellipticae, coriaceae, 5.5-I6.5 cm longae, 2.5-8.0
cm latae, supra glabrae, subtus cum cavis stomatalis faucis lanato-pubescentibus,
venulis glabris conspicuis; costa media supra impressa, glabra; costis secundariis
8-I jugis, subtus prominentibus, supra planis conspicuis. Stipulae lanceolatae, circa
4.0 mm longae, puberulentes, subpersistentes,cum basi petioli adnatae. Flores circa
2.0 mm longi, haud pedunculati, in paniculis racemosis terminalibus axillaribusque
dispositi, rachi ramisque puberulis. Bracteae bracteolaeque lanceolatae ad 2.5 mm
longum, persistentes, extus puberulae. Receptaculum campanulatum, subsessile,
extus griseo-tomentellum, intus lanatum. Calycis-lobi acuti utrinque tomentelli.
Petala nulla. Stamina 5, unilateralia; filamenta pubescentia, calycis-lobis breviora.
156 Flora Neotropica

Ovarium ad basim receptaculi insertum, villosum. Stylus e basi ovarii ortus, calycis-
lobis brevior, lanato-pubescens. Drupa oblonga, ad 7.0 cm longa, 4.0 cm lata, extus
velutino-pubescens; mesocarpio tenui; endocarpio circa 6.0 mm crasso, lignoso duro,
intus glabro.
TYPE.Ducke293 (2nd collection), Brazil, Amazonas, Manaus, fl (holotype, K;
isotypes, IAN, US).
DISTRIBUTION. Non-flooded forest in the vicinity of Manaus and the lower Rio
Negro.
BRAZIL. Amazonas: Manaus, Ducke293 (Ist collection) fl (F, GH, K, NY, R, S, US); 2165 fr
(GH, IAN, NY, R); RB 35567 fl (RB); CoelhoINPA 6004 st (INPA); Pires & Black 936 fl (GH, IAN,
NY); Ferreira147 fl (INPA 5905); Rio Negro, Marabitanus, Pires 469 fl (IAN, NY).

LOCAL NAME. Cariperana.

Licaniabracteatais most closely related to L. robusta,from which it differs in the
more pubescent inflorescence, the broader stipules, the better developed stomatal
cavities with glabrous reticulation, the distinctive bracts, and the shorter petioles.
The name was suggested by the distinctive persistent lanceolate bracts.

129. Licania parviflora Bentham, Jour. Bot. Hooker 2: 221. 1840; Hook. f.,
Mart. Fl. Bras. I4(2): i8. 1867; Fritsch, Ann. Naturh. Mus. Wien 4: 5I.
1889. Fig 23 E-H.

Tree to 25.0 m tall, the young branches glabrous and conspicuously lenticellate.
Leaves oblong-ovate to oblong-lanceolate, coriaceous to submembraneous, 4.5-12.5
cm long, 2.0-5.5 cm broad, the apex with a finely pointed acumen 5.0-20.0 mm
long, often subfalcate, rounded to subcuneate at base, glabrous above, the lower
surface with well developed stomatal cavities filled with lanate pubescence, or smooth
and lanate-tomentose without stomatal cavities, the reticulation glabrous and
conspicuous; midrib plane above, glabrous; primary veins 7-II pairs, prominent
beneath, plane above; petioles 4.0-6.0 mm long, terete, eglandular, puberulous, soon
becoming glabrous. Stipules linear, to 4.0 mm long, glabrous, persistent, adnate to
extreme base of petiole. Inflorescencesracemose panicles with puberulous to glabres-
cent rachis and tomentellous to glabrescent branches. Flowers ca 2.0 mm long, on
primary branches of inflorescence.Bracts and bracteoles 0.2-0.5 mm long, triangular,
subpersistent, puberulous on exterior. Receptacle campanulate, sessile, gray-tomen-
tellous on exterior, tomentose within. Calyx lobes acute, tomentellous on exterior,
puberulous within. Petals absent. Stamens 5-6, inserted in a nearly complete circle;
filaments shorter than calyx lobes, glabrous. Ovary inserted at base of receptacle,
tomentose. Style equalling filaments, pubescent throughout. Fruit oblong-pyriform,
2.5-3.0 cm long; epicarp with sordid pulverulent-velutinousferrugineouspubescence;
pericarp thin, hard, fibrous, sparsely hirsutulous within.
TYPE. Schomburgk 977, Brazil, Amazonas, fl (holotype, K; isotypes, BM, CGE, G,
L, OXF, P).
DISTRIBUTION. Periodically flooded and riverine forest in Venezuela, and western
and central Amazonia.
COLOMBIA. Vaupes: Gutierrez & Schultes862 fr (GH); 875 fl (GH); 882 fl (GH); 907 fr
(COL, GH). VENEZUELA. Bolivar: Cardona II77 fl (NY, VEN); 2557 fr (US, VEN);
2569 fl (NY, US, VEN); 2821 fl (F, US, VEN); Steyermark& Wurdack4 fl fr (F, NY, VEN); LI.
Williams 12063 fr (F, K, US); Wurdack2i6 fl fr (NY, US, VEN). Terr. Amazonas: Ll. Williams I4755
fr (F, US, VEN); Maguire & Wurdack34801 fr (F, GH, NY); Maguire, Wurdack& Bunting37415 fl
(F, NY). GUYANA. Atkinson88 fl (BM, GH, MICH); Lang & Persaud242 fl (F); Pinkusr19 fl (BR, F,
Licania 157

GH, IAN, MO, NY, S, US); 260 fr (F, GH, NY, US); P69(F.D. 2841) fl (FHO, K, NY, S). BRAZIL.
Amazonas: DuckeRB 19780 fl (K, RB); Froes 25244 fl (IAN, NY, RB); 25556 fl (IAN, NY, RB, UB);
2618 fr (IAN, NY); INPA 6117 fr (IAN, INPA); Krukoff6670 fl (A, BR, F, IAN, K, LE, MICH, MO,
NY, RB, S, US); 7002 fl (A, BR, F, K, LE, MICH, MO, NY, RB, S, U, US); 7237 fr (A, BM, BR,
F, K, LE, MICH, MO, NY, RB, S, U, US); Martiussn fl (BR, K, M); Pires 176 fl (IAN, NY); i80 fl
(IAN, NY); Poeppig2770 fl (LE, P, W); Pranceet al 3356 fr (INPA, NY); Schwacke385 fr (R, RB);
Spruce1953 fl (F, GH, LE, M, NY); 2885 fl (F, GH, GOET, LD, LE, NY, OXF, P); 33I4 fl (BM, GOET
K, LD, LE, NY, OXF, P); Rodrigueset al 1404 fl (INPA 7735); 2206 fl (INPA 8596); 7138 fr (INPA
15800, NY). Para: DuckeMG 2983 fl (RB 15175); MG 1o672 fl (RB I5171); RB 18808 fl (RB); Mexia
6047 fl (F, GH, LA, MICH, MO, S, U, US); Pires, Black, Wurdack& Silva 6399 fr (IAN, NY, RB);
Pires & Silva 4402 fl (IAN, INPA, R). Terr. Roraima: Pranceet al 4039 fl (INPA, NY). Terr. Rond6nia:
Maguire,Pires, Maguire & Silva 56696 fl (F, GH, NY); Pranceet al 8769 fl (INPA, NY). Mato Grosso:
KuhlmannRB I7973 fl (K, RB).

LOCALNAMES.Venezuela: Hierito, Huaje (Guariquena), Huipanupare (Banita),

Kuepi (Arekuna), Pasa. Brazil: Caripe, Miwari-ek (Akawai).
Many varieties of this species have been described, but they are either not
distinct or show closer affinities with other species. The confused status of the alleged
varieties results from long-term misinterpretation of L. parvifloracompounded in
some instances by lack of reference to existing types. A summary of the varieties and
their presently interpreted status is given below.
i. var subfalcata Spruce ex Hooker f., Mart. Fl. Bras. 14(2): I8. I867. Types.
SpruceI429, 1953, 2885, 3314. In light of further material now available this variety is
not distinct from other material and is considered here as part of L. parviflora,because
there is an overlap in the leaf characters used to separate var subfalcata.
2. var pallida Spruce ex Hooker f., Mart. Fl. Bras. 14(2): i8. i867. =Licania
pallida Spruce ex Sagot.
3. var grandifolia Hooker f., Mart. Fl. Bras. I4(2): i8. 1867. =Licania vaupesiana
Cuatrecasas.
4. var conduplicata Maguire, Fieldiana Bot. 28: 253. I952. =Licania hypoleuca
Bentham.
5. var submembranaceaMaguire, Fieldiana Bot. 28: 254. I952. =Licania kunthiana
Hooker f.
Fritsch (1889, p. 5I) recognized two forms (f. brevifoliaand f. longifolia)of this
species. These were based on leaf size, but in light of the additional material they
cannot be maintained, since leaf size shows wide variation without correlation with
other characters.

130. Licania robusta Sagot, Ann. Sci. Nat. Bot. VI. 15: 306. 1883; Fritsch,
Ann. Naturh. Mus. Wien 4: 32. 1889; Benoist, Bull. Mus. Mus. Hist. Nat.
Paris 25: 515. 19I9; Sandwith, Kew Bull. 193I: 373, I93I; Kleinh. in
Pulle, Fl. Suriname 2(I): 440. 1939.
Kleinh., Rec. Trav. Bot. Nderl.22: 384. 1925.Type. B. W. 2729, Suriname,
Licaniapachystachya
fl (U).

Tree to 25.0 m tall, the young branches glabrous, conspicuously lenticellate.

Leaves elliptic, coriaceous, 8.0-25.0 cm long, 4.5-8.5 cm thick, acuminate at apex,
with acumen 5.o-I5.0 mm long, rounded at base, glabrous above, the lower surface
with dense brown-lanate pubescence obscuring the prominent deeply reticulate
venation; midrib prominulous above, glabrous; primary veins 6-8 pairs, prominent
beneath, prominulous above; petioles 1.2-2.0 cm long, glabrous, eglandular, canali-
culate. Stipules linear, ca 4.0 mm long, glabrous, persistent, adnate to base of petiole.
158 Flora Neotropica

Inflorescences terminal and subterminal racemose panicles, the rachis glabrous or

glabrescent, lenticellate, with puberulous to tomentellous branches. Flowers 2.5-3.0
mm long, solitary or in small groups on primary branches of inflorescence. Bracts and
bracteoles 0.3-1.5 mm long, triangular to linear, subpersistent, exterior puberulous.
Receptacle narrowly campanulate to urceolate, subsessile, exterior tomentellous,
interior tomentose. Calyx lobes acute, tomentellous on both surfaces. Petals absent.
Stamens 5-6, unilateral; filaments shorter than calyx lobes, glabrous. Ovary inserted
at base of receptacle, tomentose. Style equalling filaments, lanate. Fruit (immature)
oblong; epicarp with short yellow-brown pubescence.
TYPE.Melinonsn (1862), French Guiana, fl (holotype, P; isotypes, A, BM, F,
GH, K, NY, R, US).
DISTRIBUTION. Primary forest in the Guianas and Para in Brazil.
SURINAME. B.B.S. 202 fl (U); 1107 st (U); B.W. II fl (U); 1300 st (U); 1460 st (U); i6i6 fi
(U); 2658 fl (A, MO); 2686 fl (IAN, K, NY); 2729 fl (U); 2806 fl (U); 2818 fl (A, MO, U); 2882 fr
(MO, U); 2906 fr (IAN, K, NY); 3098 fr (U); 3768 fl (U); 5387 fr (U); 5863 fr (U); 6156 pro parte st
(U); 6816 fl (IAN, K, NY); Stahel8oA fl (A, IAN, K, NY, U). FRENCH GUIANA. For. Serv. 7676 fl
(P, U); Wachenheimii fl (NY, P). BRAZIL. Pari: DuckeRB i8815 fl (K, RB); Oliveira230 fl (IAN);
Pires 51818 st (NY).

LOCALNAMES.Suriname: Grootblad,Foengoe, Bongro, Goro-goro,Marishiballi.

French Guiana: Weti-koko.
Most of the Suriname collections cited above are from two trees in forest reserves
that have been re-collected many times.
Licaniarobustamay be distinguished from related species by the glabrous lenti-
cellate rachis of the inflorescence, and by the long petioles.

I31. Licania lanceolata Prance, sp nov Fig 23 A-E.

Frutex vel frutex suffruticosa, ramulis juvenilibus tomentellis, mox glabris.

Folia alternata petiolata; petiolo 2.0-Io.0 mm longo, tereti, eglanduloso, juvenili
tomentoso; laminae lanceolatae, basim versus latissimae, crasse coriaceae, 3.5-6.0 cm
longae, 1.0-2.5 cm latae, basi rotundatae, apice acutae, supra glabrae, subtus dense
lanato-pubescentes, profunde reticulatae vel cum cavis stomatalis munitae; costa
media supra leviter impressa, basim versus pubescente; costis secundariis 7-IO jugis,
subtus prominentibus, supra planis arcuatis ante marginem confluentibus. Stipulae
minutae, ad I.o mm longum, lineares, puberulae, subpersistentes, extrapetiolares.
Flores circa 2.0 mm longi, haud pedunculati, in paniculis racemosis pauci-ramosis
terminalibus axillaribusque et spicis axillaribus dispositi, rachi ramisque brunneo-
tomentellis. Bracteae bracteolaeque 0.3-I.o mm longae, ovatae, persistentes, extus
puberulae. Receptaculum campanulatum, sessile, extus tomentosum, intus lanatum.
Calycis-lobi acuti, utrinque tomentelli. Petala nulla. Stamina 6-8, in orbem comple-
tum disposita; filamenta glabra, ad basim libera, calycis-lobis breviora. Ovarium ad
basim receptaculi insertum, villosum. Stylus e basi ovarii ortus, calycis-lobis brevior,
ad apicem villosus. Drupa oblonga, parva, extus sordido-pubescens;pericarpio tenue,
fibroso, fragile, intus hirsutulo.
TYPE. Maguire & Wurdack35568, Venezuela, Terr. Amazonas, Rio Guainia,
Sabana El Venado, fl (holotype, NY).
Known only from the savannas of the upper Orinoco region of
DISTRIBUTION.
Venezuela.
VENEZUELA. Terr. Amazonas: Rio Ventuari, Sabana Moyo, Level30, fl (NY).
Licania 159

I= .
D E

I /:

......y.
'..

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

... : . ....,....!...

....~.....

A=4'

FIG 23. Speciesof Licania.A-E, L. lanceolata (Level30, Maguireet al 35568); A, habit, x o0.; B,
flower, x Io; C, flower section, x Io; D, fruit, x 2.5; E, fruit section, x 2.5. F-J, L. paraensis(DuckeRB
i88i8, 19772); F, habit, x o.-; G, flower, x Io; H, flower section, x Io; J, young fruit, x 4; K-M, L.
bracteata(Ducke293); K, habit, xo-5; L, flower, x Io; M, flower section x Io.
160 Flora Neotropica

This species is most closely related to L. incana,but it differs in the leaves by the
distinctive shape and the deeply reticulate lower surface.

132. Licania spicata Hooker f., Mart. Fl. Bras. 14(2): i6. 1867; Fritsch, Ann.
Naturh. Mus. Wien 4: 47. I889.
MoquileaorganensisMiers, Jour. Linn. Soc. Bot. 17: 374. 1879. Type. Miers 4095, Brazil, Rio de
Janeiro, fl (BM, K, P).
Licaniaorganensis(Miers) Fritsch, Ann. Naturh. Mus. Wien 4: 59. 1889.

Tree, the young branches puberulous, becoming glabrous and lenticellate with
age. Leaves oblong, subcoriaceous, 4.5-9.0 cm long, 1.5-3.7 cm broad, the apex with
finely pointed acumen 5.0-I5.0 mm long, subcuneate to cuneate at base; upper sur-
face glabrous or sparsely strigose when young, densely yellow-brown lanate below;
midrib slightly impressed above, glabrous; primary veins 8-I pairs, prominent be-
neath, plane above; petioles 6.0-0.9 mm long, tomentellous, canaliculate, eglandular.
Stipules deciduous (not seen). Inflorescences axillary and terminal, the axillary un-
branched spikes, the terminal few-branched racemose panicles, the rachis and
branches puberulous to tomentellous. Flowers ca 2.5 mm long, on rachis and primary
branches of inflorescence. Bracts and bracteoles 0.3-0.8 mm long, ovate, persistent,
tomentellous on exterior. Receptacle cupuliform-globose, sessile, tomentellous to
puberulous on exterior, tomentose within. Calyx lobes acute, tomentellous on exterior,
puberulous within. Stamens 5-6, slightly unilateral; filaments shorter than calyx
lobes, glabrous. Ovary inserted at base of receptacle, villous. Style equalling filaments,
villous throughout. Fruit (immature) with densely tomentose epicarp.
TYPE.Miers 4095, Brazil, Rio de Janeiro, fl (holotype, K; isotypes, BM, P).
DISTRIBUTION. Known only from forested regions around Rio de Janeiro.
Flowering January to April.
BRAZIL. Rio de Janeiro: Binot 40 fl (P); For. Serv. 40 fl (R 21337); Glaziou 8399 fl (BR, F, K,
LE, P, R, S, US); i943 fl (A, BR, F, K, LE, NY, P); 19137 fl (A, BR, GH, K, LE, NY, P); Miers sn
fl (A, NY probable isotypes); Mus. Nac. 6323I fl (R); Vellososn fl (R 38617); sn fl (R 38619).

LOCAL NAME. Milho cozido.

133. Licania stricta Kleinhoonte, Rec. Trav. Bot. Neerl. 22: 380. I925;
Kleinh. in Pulle, Fl. Suriname 2(I): 438. I939.

Tree (?), the young branches tomentellous, becoming glabrous with age. Leaves
oblong-ovate, coriaceous, 5.5-9.0 cm long, 2.5-4.0 cm broad, the apex with finely
pointed acumen 6.o-io.o mm long, subcuneate at base, glabrous above, densely
yellow-brown-lanate below, the pubescence easily removed; midrib plane above,
glabrous except at base; primary veins 7-Io pairs, prominulous beneath, plane above;
petioles 3.0-4.0 mm long, terete, eglandular, tomentellous when young. Stipules to
2.0 mm long, linear, pubescent, adnate to extreme base of petiole. Inflorescences ter-
minal and subterminal spikelets attached to rachis ca Io.o cm long, the rachis and
branches tomentose when young. Flowers ca 2.0 mm long, in dense spikelets along
rachis. Bracts and bracteoles ovate, 1.0-2.5 mm long, densely crowded along spike-
let, persistent, tomentose on exterior. Receptacle campanulate, sessile, tomentose on
exterior, tomentose within. Calyx lobes acute, tomentellous. Petals absent. Stamens
Licania 161

5-6, unilateral; filaments shorter than calyx lobes, glabrous. Ovary inserted at base
of receptacle, villous. Style equalling filaments, pubescent throughout. Fruit unknown.
TYPE.B.W.3463, Suriname, fl (holotype, U).
DISTRIBUTION. Known only from the type gathering, collected beside the Mara-
takka River, Suriname.
The status of this species is rather dubious. It is very close to L. incanaand L.
and may be an aberrant form of one of them. The only difference from
leptostachya,
L. incanais the inflorescence with short densely crowded spikelets.

134. Licania leptostachya Bentham, Jour. Bot. Hooker 2: 220. I840; Fritsch,
Ann. Naturh. Mus. Wien 4: 46. I889. Kleinh. in Pulle, Fl. Suriname
2(I): 434. I939.
Licaniaincanaauct non Aublet viz Hooker f., Mart. Fl. Bras. 14(2): I4. 1867.
Licania axilliflora Hochreutiner, Bull. N. Y. Bot. Gard. 6: 273. I9Io. Types. Schomburgk868,
Guyana, fl (holotype, NY; isotypes, K, IAN, P); Sagot 1281, French Guiana, fl (paratypes,
K, L, P, U).
Licania incanavar axillifloraSagot, Ann. Sci. Nat. Bot. VI.15: 305. 1883.
Licanialeptostachya var axilliflora(Sagot) Fritsch, Ann. Naturh. Mus. Wien 4: 46. 1889.

Small tree or shrub, the young branches puberulous to glabrescent, soon be-
coming glabrous and conspicuously lenticellate. Leaves ovate, membraneous, oblong-
ovate to oblong-lanceolate, 3.0-7.0 cm long, 1.5-3.5 cm broad, the apex usually with
a finely pointed acumen 2.0-15.0 mm long, cuneate to rounded at base, glabrous
above, densely brown-lanate below, the pubescence easily removed; midrib plane
above, pubescent towards base; primary veins 7-10 pairs, prominent beneath, plane
above; petioles 2.0-4.0 mm long, terete, eglandular, tomentose when young. Stipules
linear, to 3.0 mm long, pubescent, persistent, axillary. Inflorescences unbranched
spikes or glomerules, predominantly axillary, the rachis tomentose when young, be-
coming glabrescent in fruit. Flowers ca 2.0 mm long, in dense glomerules on rachis.
Bracts and bracteoles 0.5-1.5 mm long, triangular, persistent, puberulous on exterior.
Receptacle cupuliform- campanulate, sessile, tomentose on exterior, tomentose with-
in. Calyx lobes acute, tomentose on exterior, puberulous within. Petals absent.
Stamens 6-7, inserted in a complete circle; filaments shorter than calyx lobes, glab-
rous. Ovary inserted at base of receptacle, tomentose. Style equalling filaments,
tomentose at base, glabrous above. Fruit oblong, ca 2.5 cm long, including stipe ca
6.o mm long; epicarp short-tomentose; pericarp thin, hard, fibrous, densely hir-
sutulous-tomentose within.
TYPE. Schomburgkiii, Guyana, fl (holotype, K; isotypes, CGE, F, G, L, LE,
NY, US, W).
DISTRIBUTION.Riverine forest, river banks and river islands in the Guianas and
northeastern Amazonian Brazil.
VENEZUELA. Bolivar: Budowski40 fr (NY); 40-1 fr (NY). GUYANA. Graham354 fl (K);
Guppy595(F.D. 7610) fl (NY); Im Thurnsn fl (K); Myers5894 fl (K); Schomburgk 342 fl (BR); 1279 fl
(BR); A. C. Smith 2570 fl (A, F, K, MO, NY, P, S, US); 2653 fl (A, F, MO, NY, P, S, US); 3024 fl
(A, F, MO, NY, P, S, U, US); Wilson-Brownesn (F.D. 6491) fl (K, NY). SURINAME. B.W. 1ooI fl
(U); 2194 fl (NY, U); 3067 fl (U); 3624 fl (A); 3824 fl (NY); Dept. Lanbouw244 fl (U); Hulk 320 fl (U);
Irwin, Prance,Soderstrom& Holmgren55375 fl (F, GH, MO, NY, US); 55756 fl (F, GH, LE, NY, US);
Lanjouw727 fl (MO, U); 792 fl (U); Lanjouw& Lindeman895 st (U); Maguire24862 fl (A, BR, F, LE
MO, NY, US); Pulle 439 fl (IAN, K, NY); Tulleken519 fl (U); vanDonselaar1078st (NY, U). FRENCH
GUIANA. Benoist 726 fl (P); Irwin & Westra47470 fl (MO, NY, US); Jussieu sn fl (P); Poiteau sn fl
(FI, K, LE, P); Leprieursn fl (P); Melinon 122 fr (P); Wachenheim74 fl (P); 138 fl (P); 217 fl (P);
Westra48552fr (NY, US). BRAZIL. Para: DuckeMG 8929 fl (BM, MG, RB); MG i4996fl (RB 15155);
Fittkau & CoelhoINPA 12816 fl (INPA, NY); Froes 34133 fl (IAN); Froes & Black 24553 fl (IAN,
162 Flora Neotropica

NY); Oliveira 4095 fl (IAN, NY); 4651 fl (IAN, NY); Pires & Silva 4620 fl (IAN, NY); Prance &
Pennington1734 fr (GH, IAN, NY); 1762 fl fr (F, MO, NY); R. S. RodriguesMG 4043 fl (MG, RB
15156); Sampaio5547 fl (R); 5636 fl (R); 5712 fl (R). Terr. Amapi: Frdes& Black 27705 fr (IAN, UB);
Irwin & Westra47434 fl (F, IAN, MG, MO, NY, US); 47617 fl (F, IAN, MG, MO, NY, US); Pires,
Rodrigues& Irvine51055 fl (IAN, MG, MICH, NY, US). Maranhao: Froes I735 fl (A, F, K, MO, NY,
P, S, US); 3456 fl (IAN).

LOCAL NAMES. Venezuela: Hierrito. Suriname: Onitjatja, Sabana Foengoe. Brazil:

Caripe.
This species is very close to L. incana,but the considerable material available
reinforced with my own field observations make clear the distinctions between the
two. The principal differences are in the predominantly axillary inflorescence, and
the membraneous leaves with well developed acumen of L. leptostachya which is a
species of river margins and forest, while L. incanais confined to savannas throughout
its range. L. axilliflorawas described as a separate species because of its glomerulous
inflorescence. However, many herbarium sheets of L. leptostachya have both glomer-
ules and spikes on the same collection, negating any basis for the maintenance of
L. axillifloraas a species or variety.

135. Licania incana Aublet, P1. Guiane I: I19. 1775; DC., Prodr. 2: 527.
I825; Sandwith, Kew Bull. 1931: 372. 1931; Kleinh. in Pulle, Fl. Suriname
2(I): 437. I939.
Hedycreaincana(Aublet) G. F. Gmelin, Linn., Syst. Veg. ed. 13. 2: 428. 1796; Willd., Linn., Sp.
P1. ed. 4. I: 1210. 1797, nom illegit.
Chrysobalanus incanus(Aublet) G6mez de la Maza, Fl. Cuba 39. 1887.
LicaniacrassifoliaBentham, Jour. Bot. Hooker 2: 221. I840; Miquel, Stirp. Surin. Sel. 29. 1850;
Fritsch, Ann. Naturh. Mus. Wien 4: 48. I889. Type. Schomburgk 388 (381 some sets),
Guyana, fl (holotype, K; isotypes, CGE, L, NY, OXF, P, US, W).
Licania leptostachyaBentham var crassifolia (Bentham) R. Benoist, Bull. Mus. Hist. Nat. Paris
25: 512. 1919.

Shrub or rarely small tree, the young branches tomentellous to puberulous, soon
becoming glabrous and lenticellate. Leaves ovate to oblong, thick-coriaceous, 2.5-
8.5 cm long, 1.3-4.0 (-5.5) cm broad, .-cute to acuminate at apex, with acumen to
Io.o mm long, usually rounded at base, rarely subcuneate, glabrous above, densely
brown-lanate below, the pubescence easily removed; midrib plane or slightly im-
pressed above, glabrous; primary veins 7-9 pairs, prominulous beneath, plane above;
petioles 2.0-5.0 mm long, terete, eglandular, tomentellous when young. Stipules to
2.0 mm long, linear, persistent, pubescent, adnate to extreme base of petiole or
axillary. Inflorescences terminal and axillary spikes, occasionally slightly branched,
the rachis tomentose when young. Flowers ca 2.0 mm long, in small glomerules along
rachis. Bracts and bracteoles ovate to lanceolate, I.0-3.0 mm long, persistent,
tomentose on exterior. Receptacle campanulate, sessile, tomentose on exterior,
tomentose within. Calyx lobes acute, tomentose on exterior, puberulous within.
Petals absent. Stamens 5-7, slightly unilateral; filaments shorter than calyx lobes,
glabrous. Ovary inserted at base of receptacle, lanate. Style equalling filaments,
tomentose throughout. Fruit globose, ca 1.6 cm diameter, including stipe ca 2.0 mm
long; epicarp with short tomentellous pubescence; pericarp thin, hard, fibrous,
tomentose-hirsute within.
TYPE. Aublet sn, French Guiana, st (BM, P-Rousseau).
DISTRIBUTION. Savannas, savanna margins and savanna forests in Venezuela,
the Guianas and northern Amazonian Brazil.
Licania 163

Representative collections: VENEZUELA, Bolivar: Maguire, Steyermark& Maguire 53584 fl

(NY, US); Steyermark 89716 fl (NY). Terr. Amazonas: Maguire, Wurdack& Maguire 42473 fl (F, GH,
MO, NY, RB). GUYANA. Anderson586 fl (K); Bartlett8037 fl (K); Cowan39273fr (F, GH, NY, RB);
Cowan & Soderstrom1917 fl (NY, US); 1948 fl (NY, US); 2140 fl (NY, US); Davis 296(F.D. 2287) fl
(K); De La Cruz 1775 fl (F, GH, K, MO, NY, US); 1777 fl (F, GH, MO, NY, US); 2050 fl (F, GH,
MO, NY, US); 2661 fl (F, GH, MO, NY, US); 3828 fl (F, GH, K, MO, NY, US); Fanshawe633(F.D.
3369) fl (K, NY); Graham147 fr (K); Hitchcock16913 fl (GH, US); Hohenkerk676A fr (K); Irwin 6 fr
(US); Jenman3742 fl (K, U); Maguire,Bagshaw & Maguire40662 fl (NY); Maguire & Fanshawe23151
fl (A, NY); 23266 fl (A, F, K, MO, NY, US); 32243 fr (NY); 32603 fr (NY); Maguire & Maguire
45911 fl (NY); Sandwith1044 fl (K, NY, U); Tillett & Tillett 43846 fr (NY); Tutin 292 fl (K, RB, U,
US); 626 fl (K, RB); Waby8442 fl (K); Wilson-Browne467(F.D. 5877) fl (NY). SURINAME. B.B.S.
199 fr (K, NY, U); 200 fl (U); Boldingh3871 fl (U); 3883 fl (U); B. W. 132 fl (U); 413 fl (A, MO);
496 fr (U); 525 fl (U); 547 fl (U); 2382 fr (U); 2817 fl (U); 2881 fr (K, NY); 3920 fl (U); 4366 fr (U);
4806 fr (NY); 5384 fr (U); Florschiitz640 fl (NY, U); Focke 1281 fl (U); Heinsdijk3 st (U); 55 fl (U);
Heyligers117 fl (U); 746 fl (U); Jonker 149 fl (U); Hostmann& Kappler794 fl (F, GH, GOET, K, LE,
MO, NY, OXF, P, S); 1712 fl (U); Krukoff12316 fr (A, NY); Kuyper56i fr (U); Lanjouw152 fl fr (U)
438 fl (U); 1245 fr (K, NY); Lanjouw& Lindeman265 fl (K, NY); 563 fl (K, NY); 732 fl (NY); 1838
fl (K, NY); 3287 fl (NY); L.B.B. 8358 fl (U); 9131 fl (U); Lems5124 fl (NY); Lindeman3894 st (U);
4636 fl (U); Maguire24232 fl (NY, US); 24354 fl (A, BR, F, K, LE, MO, NY, US); 24438 fl (A, F, K,
NY, US); 24791 fl (A, F, K, NY, US); 40827 fl (COL, MICH, MO, NY); Maguire & Stahel23651 fl
fr (A, F, K, MO, NY, US); 24982 fl (A, F, K, NY, US); Pulle 56 fl (U); 143 st (U); 152 fl (U);
Samuels223 fr (A, GH, K, NY, P); Stahel 141 fl (A, K, NY, U); de Vriesesn fl (K); Wullschlaegel964 fl
(BR, GOET). FRENCH GUIANA. Benoist 146 fl (P); 278 fl (P); 555 fl (P); Cowan38031 fl (NY, P,
US); 38873 fr (F, GH, K, NY, P, RB, US); Melinonsn fl (P, R); 148 fl (P); Wachenheim sn fl (P, US);
317 fl (P). BRAZIL. Amazonas: Holt & Blake 517 fl (F, NY, P, RB, S, US). ParA: Black 48-3310 fl
(IAN, NY); 50-8675 fl (IAN, NY); 54-16850 fl (IAN, NY); Black et al 57-I9589 fl (IAN, NY);
DuckeMG 11288 fr (BM, MG, P, RB, US); MG 12563 fl (BM, MG, P, RB, US); RB 19787 fl (K, RB);
Egler 260 fl (UB); Pires 6039 fl (NY); Poeppig2985 fl (F, GOET, LE, NY, W); Prance,Pennington&
Pires 1287 fl (F, NY).

LOCALNAMES.Guyana: Marishiballi, Unikiakia. Suriname: Kwepilan, Marishiballi,

Onitjatja,SabanaFoengoe,Oenikiakia,Akoerieerepare.French Guiana: Caligni.
Material of L. leptostachyaand L. kunthianahas frequently been cited under L.
incana,and material of the latter species has frequently been cited under L. crassifolia.
This confusion arose because several authors did not refer to the Aublet type collec-
tion. Licaniacrassifoliadoes not differ from L. incanain any characters, and conse-
quently I follow those authors who regard it as a synonym of L. incana.Licania
and L. kunthianaare both distinct from L. incana;for the differences see
leptostachya
under the two former species.

136. Licania paraensis Prance, sp nov Fig 23 F-J.

Arbor magna, ramulis juvenilibus puberulis mox glabris. Folia alternata,

petiolata; petiolo 8.o-i5.0 mm longo, tomentello, canaliculato, glanduloso vel
eglanduloso; laminae ellipticae, submembranaceae, 9.0-22.0 cm longae, 5.5-I0.0 cm
latae, basi rotundatae vel subcuneatae, apice rotundatae vel mucronatae, supra
glabrae, subtus profunde reticulatae, griseo-lanato-pubescentes, venulis parallelis;
costa media supra glabra, impressa; costis secundariis I 0-I 2 jugis, subtus prominenti-
bus, supra planis. Stipulae lanceolatae, puberulae, 5.0-10.0 mm longae, cum petiolo
adnatae. Flores 2.5-3.0 mm longi, haud pedunculati, in paniculis racemosis termina-
libus subterminalibusque dispositi, rachi puberuli, ramis ferrugineo-tomentellis.
Bracteae bracteolaeque o.3-2.o mm longae, lanceolatae, subpersistentes, extus
puberulae. Receptaculum campanulato-cupuliforme, sessile, extus ferrugineo-
tomentellum, intus tomentosum. Calycis-lobi acuti, extus tomentelli, intus puberuli.
164 Flora Neotropica

Petala nulla. Stamina 6, in orbem completum inserta, filamenta glabra, ad basim

libera, calycis-lobis breviora. Ovarium ad basim receptaculum insertum, pilosum.
Stylus e basi ovarii ortus, ad apicem pilosus, calycis-lobis aequans. Drupa (immatura
tantum visa) extus velutino-rufo-pubescens.
TYPE. DuckeRB i88i8, Brazil, Para, Santarem, fl (holotype, K; isotypes, NY,
RB).
DISTRIBUTION. Non-flooded forest in Amazonian Brazil and Bolivia.
& Loureiro
BRAZIL. Amazonas:Manaus-Itacoatiararoad, Km 99, Rodrigues 7136 fl (INPA).
Para: Obidos, Ducke MG 15142 fl (K, MG, RB I9772). BOLIVIA. Pando: Rio Madeira opposite
Riberao, Pranceet al 6532 fl (INPA, NY).

This species is distinct by its densely crowded inflorescencewith short velutinous

pubescence on the flowers and branches. It is probably most closely related to L.
laxifloraand L. mollis,but is well differentiated from both species.

137. Licania vaupesiana Killip & Cuatrecasas, Fieldiana Bot. 27: I05. 1951.
Licania parvifora Bentham var grandifoliaHooker f., Mart. Fl. Bras. I4(2): 19. 1867; Fritsch
Ann. Naturh. Mus. Wien 4: 5I. 1889. Type. Martiussn, Colombia, Amazonas, fl (holotype
M).

Tree, the young branches puberulous, soon becoming glabrous and lenticellate.
Leaves elliptic, coriaceous, 7.0-I4.0 cm long, 3.7-5.5 cm broad, acuminate at apex,
with acumen 5.o-Io.o cm long, rounded at base, glabrous above, densely brown-
lanate beneath; midrib impressed above, pubescent towards base; primary veins
I-12 pairs, prominent beneath, plane above; petioles 4.0-7.0 mm long, puberulous,
terete, eglandular. Stipules linear, ca 5.0 mm long, intrapetiolar or adnate to inside
of extreme base of petiole, puberulous, persistent. Inflorescencesterminal and subter-
minal panicles, the rachis and branches puberulous. Flowers ca 2.0 mm long, in small
subsessile groups or solitary on primary branches of inflorescence. Bracts and bracte-
oles to i.o mm long, triangular, persistent, puberulous on exterior. Receptacle cam-
panulate, sessile, tomentose on exterior, tomentose within, Calyx lobes acute, tomen-
tellous on both surfaces. Petals absent. Stamens 5-7, unilateral; filaments shorter
than calyx lobes, glabrous. Ovary inserted at base of receptacle, tomentose. Style
equalling filaments, tomentose throughout. Fruit (immature) oblong; epicarp sordid-
ferrugineous pubescent, wrinkled when dry; pericarp thin, hard, woody, sparsely
hirsute within.
TYPE. Cuatrecasas6837, Colombia, Vaupes, fl (holotype, COL; isotype, frag-
ments F, US).
DISTRIBUTION. Riverine forest in Amazonian Colombia.
COLOMBIA. Amazonas: Schultes& Cabrera15563 fl (GH, NY, US).

LOCALNAME.Mooneeka.
When Hooker described L. parvifloravariety grandifolia,he commented that it
was perhaps a distinct species. Since the variety was described, it has not been re-
assessed. However, L. vaupesianawas recently described from the same region, and the
authors were apparently unaware of L. parvifloravar grandifolia.Licaniavaupesianais
certainly equal to variety grandifolia,and I agree with Cuatrecasas that it is a distinct
species. It is most notably different from L. parviflora,in the absence of stomatal
cavities from the lower surface of the leaf, the larger leaves and the much larger
number of primary leaf veins.
Licania 165

I38. Licania bahiensis Prance, sp nov Fig 24 G-J.

Arbor, ramulis juvenilibus glabris lenticellatis. Folia alternata petiolata; petiolo

5.0-6.0 mm longo, puberulo, tereti, eglanduloso, laminae obongo-ellipticae, coria-
ceae, 5.5-8.0 cm longae, 2.5-4.5 cm latae, basi rotundatae vel subcuneatae, apice
apiculatae vel in acuminem I.0-5.0 mm longum contractae, supra glabrae, subtus
dense lanato-pubescentes, venulis planis; costa media supra glabra, leviter impressa;
costis secundariis 6-8jugis, subtus prominentibus, supra planis. Stipulae circa 3.0 mm
longae, lanceolatae, puberulae, persistentes, cum basi petiolo adnatae. Flores circa
2.0 mm longi, haud pedunculati, in paniculis racemosis terminalibus axillaribusque
dispositi, rachi ramisque griseo-puberulis.Bracteae bracteolaeque 0.5-1.5 mm longae,
ovatae, persistentes, extus puberulae. Receptaculum campanulatum, sessile, extus
griseo-tomentellum, intus tomentosum. Calycis-lobi acuti, extus tomentelli, intus
puberuli. Petala nulla. Stamina 5, unilateralia; filamenta glabra, ad basim libera,
calycis-lobis breviora. Ovarium ad basim receptaculum insertum, villosum. Stylus e
basi ovarii ortus, pubescens, calycis-lobis brevior. Fructus mihi ignoti.
TYPE. H.P. Vellozo736, Brazil, Bahia, Ilheus, fl (holotype, R; isotype, NY).
DISTRIBUTION. Known only from the type gathering made in the coastal forest of
Bahia, Brazil.
Licaniabahiensisis closely related to L. micrantha,but differs in the larger number
of stamens and the whitish pubescence of the leaf undersurface. It is close to L.
vaupesiana,but differs in the leaf shape, the fewer primary veins, and the stipules.

139. Licania maxima Prance, sp nov Fig 24 A-C.

Arbor magna ad 32.0 m alta x 70.0 cm diametro; ramulis juvenilibus glabres-

centibus, lenticellatis. Folia alternata petiolata, petiolo 6.0-7.0 mm longo, tereti vel
leviter canaliculato, juvenili pubescente, eglanduloso; laminae oblongae, submem-
branaceae, 6.5-Io.5 cm longae, 2.5-4.7 cm latae, basi rotundatae vel subcuneatae,
apice in acuminem 4.0-8.0 mm longum contractae, supra glabrae, subtus lanato-
arachnoideae, profunde reticulatae; costa media supra glabra, plana vel basim
versus leviter impressa; costis secundariis 8-o1 jugis, subtus prominentibus, supra
planis. Stipulae 4.0-5.0 mm longae, lanceolatae, persistentes, cum petiolo adnatae.
Flores 2 mm longi, haud pedunculati, in paniculis racemosis terminalibus subter-
minalibusque dispositi, rachi ramisque griseo-puberulis. Bracteae bracteolaeque
ovatae vel lanceolatae, ad 2.0 mm longum, saepe cum floribus aequantes, persistentes,
extus puberulae. Receptaculum campanulatum, sessile, extus griseo-tomentellum,
intus tomentosum. Calycis-lobi acuti, utrinque tomentelli. Petala nulla. Stamina 5,
in orbem completum disposita; filamenta glabra, ad basim libera, calycis-lobis
breviora. Ovarium ad basim receptaculi insertum, villoso-tomentosum. Stylus e basi
ovarii ortus, calycis-lobis brevior, ad apicem pubescens. Fructus mihi ignoti.
TYPE. Maguire,Pires & Maguire47081, Brazil, Terr. Amapa, Rio Oiapoque, fl
(holotype, NY; isotypes, IAN, MG).
DISTRIBUTION. Known only from the type collection, made in riverine forest.
This distinct species is closest to L. mollis,but differs in the narrower membrane-
ous leaves, the smaller flowers, the thinly puberulous inflorescence, and the fewer
stamens.
166 Flora Neotropica

"e''.-
'Is' t n\
s /:r E \ ,
-' B c

E~~~~~~~~
'Y~~~~~~~~~'
L/ = A"'
-,~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~o,
... ......

~~~~~~~~~~~~~~G

FIG 24 Speciesof Licania.A-C, L. maxima(Maguire etal 4708z); A, habit, x o.5; B, flower, x i o;

C, flowersection, x Io. D-F, L. blackii(Pranceet al 58989); D, habit, x 0.5; E, flower, x io; F, flower
section, x i o. G-J, L. bahiensis( Vellozo736); G, habit, x o.5; H, flower, x Io; J, flower section, x Io.
Licania 167

I40. Licania mollis Bentham, Jour. Bot. Hooker 2: 219. I840; Hook. f., Mart.
Fl. Bras. 14(2): IO. I867; Fritsch, Ann. Naturh. Mus. Wien 4: 42. I889.
Licania lucida Macbride, Candollea 5: 369. 1934; Fl. Peru I072. 1938. Type. Klug 462, Peru,
Loreto, fl (holotype, F; isotypes, NY, US).

Tree to I5.0 m tall, the young branches pubescent, becoming glabrous and
lenticellate with age. Leaves ovate-elliptic to oblong, coriaceous, 5.0-I3.0 cm long,
2.3-7.0 cm broad, cuspidate to acuminate at apex, with acumen 3.0-18.0 mm long,
base rounded to subcordate, glabrous and shining above, the lower surface deeply
reticulate-venose and densely lanate; midrib impressed for most of length above,
becoming prominulous near base, tomentose; primary veins 9-I2 pairs, prominent
beneath, plane or nearly so above; petioles 3.0-6.0 mm long, tomentose, terete,
eglandular. Stipules linear-lanceolate, 3.0-6.0 mm long, erect, adnate to base of
petiole, persistent. Inflorescences terminal and axillary racemose panicles, densely
brown-tomentose. Flowers 3.0-3.5 mm long, sessileand crowded on primary branches
of inflorescence. Bracts and bracteoles I.o-3.5 mm long, oblong-lanceolate, persistent
entire, eglandular, tomentose on exterior. Receptacle broadly campanulate, sessile,
tomentose on exterior, tomentellous within. Calyx lobes acute, tomentose on exterior,
puberulous within. Petals absent. Stamens 8-II, inserted in an arc of ca 270?;
filaments shorter than calyx lobes, connate at base. Ovary inserted at base of recept-
acle, villous. Style equalling filaments, lanate throughout. Fruit globose but surface
more or less longitudinally much-ridged, to 2.5 cm in diameter; epicarp ferrugineous-
velutinous; pericarp thick, hard, glabrous within.
TYPE. Schomburgkgio, Brazil, Amazonas, fl (holotype, K; isotypes, BM, CGE,
F, G, GH, L, NY, OXF, US, W).
River banks and river islands and in riverine forest on sandy soil,
DISTRIBUTION.
in Amazonian Colombia, Venezuela and Brazil.
COLOMBIA. Vaup6s: Cuatrecasas7304 fl (COL, F, US); Garcia-Barriga13782 fl (COL, NY,
US); Gutierrez& Schultes66i fl (GH); Schultes& Ldpez9481 fl (F GH, IAN, NY, US). VENEZUELA.
Bolivar: Wurdack& Monachino40966 fl fr (F, GH, LE, MO, NY, UB, US). Terr. Amazonas: Maguire
Cowan& Wurdack30995 fl (F, GH, NY, US); Maguire & Wurdack34891 fr (F, NY, UB, US); 34892 fl
(F, GH, NY, US); 34894 fr (GH, NY, US); 35624 fl (F, GH, NY, US); 35729 fr (F, MO, NY, US);
Maguire, Wurdack& Bunting37537 fl (F, GH, NY, US); Spruce3314 fr (BR, K, RB); 3441 fl (BM, BR,
C, GOET, K, LD, LE, NY, OXF, P); LI. Williams 15254 fr (F, US, VEN). BRAZIL. Amazonas:
Baldwin3447 fl (IAN, US); Cavalcante623 fr (MG); Chagas& CoelhoINPA 3470 fl (IAN, INPA); Froes
22198 fr (IAN, U); Krukoff7267 fl (A, BM, BR, IAN, K, MICH, MO, NY, RB, S); Pranceet al 3339 fl
(INPA, NY); 3672 fl (INPA, NY); 3691 fl (INPA, NY); 4732 fr (INPA, NY); 4886 fr (INPA, NY);
Rodrigues902 fr (INPA 7138); J. S. Rodrigues,Pires & Silva 87 fr. (IAN, NY); Silva & Brazao 60974 fl
(NY). Terr. Roraima: Froes 23001 fl (IAN, NY, SP).

LOCAL NAME. Colombia: Juan-poop (Puinave).

I4I. Licania blackii Prance, sp nov Fig 24 D-F.

Frutex vel arbor, ramulis juvenilibus tomentellis mox glabris lenticellatis. Folia
alternata petiolata; petiolo 7.0-12.0 mm longo, velutino-tomentello-pubescente,
tereti, eglanduloso; laminae oblongo-ovatae vel oblongo-lanceolatae, coriaceae,
7.0-I6.0 cm longae, 3.0-7.5 cm latae, basi subcuneatae vel rotundatae, apice acutae
vel in acuminem, I.0-20.0 mm longum contractae, supra glabrae, nitidae, subtus
profunde reticulatae, lanato-arachnoideae; costa media supra plana vel leviter
impressa, glabra; costis secundariis 5-7 jugis, subtus prominentibus, supra planis.
Stipulae lineares, ad 3.0 mm longum, puberulae, subpersistentes, cum basi petioli
168 Flora Neotropica

adnatae. Flores circa 2.0 mm longi, haud pedunculati, in paniculis racemosis ter-
minalibus subterminalibusque dispositi, rachi ramisque tomentellis. Bracteae
bracteolaeque 0.2-1.3 mm longae, triangulares, persistentes, pubescentes. Recepta-
culum campanulatum, sessile, extus tomentosum, intus tomentosum. Calycis-lobi
acuti, utrinque tomentelli. Petala nulla. Stamina circa 6, unilateralia; filamenta
glabra, ad basim libera, calycis-lobis breviora. Ovarium ad basim receptaculi in-
sertum, villosum. Stylus e basi ovarii ortus, calycis-lobis brevior, ad apicem pubescens.
Drupa immatura pyriformis; epicarpio ferrugineo-velutino-tomentello; pericarpio
duro, tenue, fibroso, intus tomentoso.
TYPE.Prance& Silva58989, Brazil, Goias, Araguaina, fl (holotype, NY; isotype,
UB).
DISTRIBUTION. Gallery forest and secondary forest in open places in eastern
Amazonia and south to the borders of the Planalto of Central Brazil.
BRAZIL. Para: Ilha de Maraj6, Black 48-3482 fl (IAN, NY); Belem, Burchell9455 fl (K);
BraganCa, Museu Goeldi9781 fl (K, RB 19782); Santar6m, Ducke RB I9773 fl (K, MG i6355, RB);
Soure, Bastos RB 138187 fl (RB). Terr. Rond6nia: 8 km northeast of Porto Velho, Pranceet al 8237 fr
(INPA, NY). Mato Grosso: Serra do Roncador near Rio Turvo, Sidney 1281 fr (UB); Chavantina,
Sidney268 fl (NY, UB); Alto Araguaia, Maguire,Pires,Maguire& Silva56354 fl (NY). GoiSs: Araguaina,
Maguire, Pires, Maguire & Silva 56097 fl (NY); Filad6lfia, Prance& Silva 58512 fl (NY, US).

This species is most closely related to L. laxiflora,but differs in its foliar venation:
the primary veins plane (not impressed) on the upper surface, and the secondary
veins are reticulate rather than parallel. In addition, the flowers of L. blackiiare also
smaller than those of L. laxiflora.

142. Licania rodriguesii Prance, sp nov Fig 25 J-N.

Arbor magna, ramulis juvenilibus puberulis, mox glabris lenticellatis. Folia

alternata petiolata; petiolo 6.0-12.0 mm longo, glabrescente, eglanduloso, leviter
canaliculato vel tereti; laminae ellipticae vel oblongae, coriaceae, 9.0-I6.o cm
longae, 3.0-7.0 cm latae, basi subcuneatae, apice in acuminem 8.o-I5.0 mm longum
contractae, saepissime subfalcatae, supra glabrae, subtus dense lanato-pubescentes,
venulis laevis; costa media supra glabra, leviter prominente; costis secundariis 6-7
jugis, subtus prominentibus, supra planis, arcuatis. Stipulae lineares, glabrae, sub-
persistentes, extrapetiolares. Flores 2.0-2.5 mm longi, haud pedunculati, in paniculis
racemosis terminalibus axillaribusque dispositi, rachi ramisque tomentellis vel
puberulis. Bracteae bracteolaeque 0.5-I.5 mm longae, ovatae, persistentes, extus
puberulae. Receptaculum campanulatum, sessile, extus tomentellum, intus tomen-
tosum. Calycis-lobi acuti, utrinque tomentelli. Petala nulla. Stamina 5, unilateralia,
filamenta glabra, ad basim libera, calycis-lobis breviora. Ovarium ad basim recepta-
culi insertum, tomentosum. Stylus e basi ovarii ortus, calyc is-lobis brevior, ad apicem
pubescens. Drupa (immatura) extus rufo-velutino-tomentella; pericarpio tenue, duro,
fibroso, intus hirsutulo.
TYPE. DuckeRB 18812, Brazil, Para, Breves, fl (holotype, K; isotype, RB).
DISTRIBUTION. Non-flooded forest in Brazilian Amazonia. Flowering June to
September.
BRAZIL. Amazonas: Manaus, Byron & Elias 67-57 fl (INPA, NY); IJNPA4062 fl (INPA, NY);
Rodrigues& Osmarino7113 fl (INPA 15775, NY); Rodrigues& Coelho7806 fr (INPA 171I2, NY). Para:
Breves, Pires, Froes & Silva 5802 fl (IAN, NY, UB). Bel6m, Ducke RB 18814 fl (K, RB); RB 19788 fl
(K, RB).
Licania 169

frui .......

G H E F

'
":.; ""'., :' -' ,

?-

-I 5?Seie ofiai. AC . tpini Mgie339 ,hbt .5 fu ..; '"

... .??DH
'.ieto, . ...oa(iJcc g, 9 60;Dabt .5:E,foe,xIo
~foe
section, x ro; G, fruit, I.5;H,
x fruit section, x I.5.J-N, L. rodriguesii (Ducke RE r86rz, 1881g);j, habit,~~~..::11..

FIG 25. Species of Licania.A-C, L. tepuiensis(Maguire33389); A, habit, x 0.5; B, fruit, x 05.; C,

fruitsection, x 0.5. D-H, L. parviflora
(PinkusI9I, 198, 260); D, habit, x o.s; E, flower, x i o; F, flower
section, x i o; G, fruit, x I .5; H, fruit section, x I .5. J-N, L. rodriguesii(DuckeRB 18812, 18814); J, habit,
xo.5; K, flower, x io; L, flower section, x 10; M, fruit, x I .5; N, fruit section, x 1.5.
170 Flora Neotropica

LOCALNAME.Pajurazinho.
It is with pleasure that I name this very distinct species for Dr. William A.
Rodrigues, an active student of Amazonian botany, who has given me much help
with my field work in Amazonia.

143. Licania indurata Pilger, Notizbl. Bot. Gart. Berlin 8: 542. 1923.

Small tree, the young branches sparsely puberulous, soon becoming glabrous
and lenticellate. Leaves oval-lanceolate to oblong-lanceolate, thick-coriaceous,
3.0-7.0 cm long, I.0-2. cm broad, bluntly acuminate at apex, the acumen 8.0-
I .o mm long, cuneate at base, glabrous above, densely lanate beneath; midrib
slightly impressed above, glabrous; primary veins 5-7 pairs, prominent beneath,
plane above; petioles 5.0-8.0 mm long, pubescent, shallowly canaliculate, eglandular.
Stipules minute, caducous, axillary. Inflorescences terminal and subterminal race-
mose panicles, the rachis and branches ferrugineous-tomentellous.Flowers ca 2.5 mm
long, solitary or in pairs on primary branches of inflorescence. Bracts and bracteoles
to I.o mm long, triangular, persistent, puberulous on exterior. Receptacle campanu-
late-globose, sessile, tomentellous on exterior, tomentose within. Calyx lobes acute,
tomentellous on both surfaces. Petals absent. Stamens 5, unilateral; filaments shorter
than calyx lobes, glabrous. Ovary insertedat base of receptacle, villous. Style equalling
filaments, puberulous. Fruit unknown.
TYPE. Schwebel8i, Brazil, Sao Paulo, fl (holotype, B, lost; lectotype, R; isotype,
SP 567; photos, F, GH, US).
DISTRIBUTION. Known only from the state of Sao Paulo in Brazil.
BRAZIL. Sao Paulo: Damasio 47980 fl (NY).

LOCALNAME.Milho Cozido.
This species is as yet poorly known. In the original description Pilger cites the
type as herbarium F.C. Hoehne567. There has been much confusion with Hoehne
labels, and an examination of the type material at Sao Paulo and the Museu Nacional,
Rio de Janeiro, shows that the type is in fact Schwebel8i (labelled II-8i in some
collections), and that the Sao Paulo collection of Schwebel8i is number 56z of that
herbarium. Licaniainduratais a very distinct species and is most closely related to
L. hoehneiand L. cruegeriana.It differs in the oblong-lanceolate, thick-coriaceous
leaves, and the impressed midrib. It is also close to L. spicata,from which it differs in
the narrower leaves with fewer primary veins, the impressed midrib, and in the
terminal, much-branched inflorescence.

144. Licania hoehnei Pilger, Notizbl. Bot. Gart. Berlin 8: 54I. I923.

Tree or shrub, the young branches puberulous, soon becoming glabrous and
conspicuously lenticellate. Leaves oblong, coriaceous, 3.7-8.o cm long, 1.5-3.0 cm
broad, usually acute but varying from rounded to acuminate at apex, subcuneate at
base, glabrous above, persistently densely gray-lanate below, the reticulations
conspicuous but shallow; midrib plane or slightly impressed above, glabrous;
primary veins 7-9 pairs, prominent beneath, conspicuous above; petioles 4.0-7.0 mm
long, tomentose when young, soon becoming glabrous, terete, eglandular. Stipules
I.0-3.0 mm long, lanceolate, persistent, tomentellous. Inflorescences dense terminal
Licania 171

and subterminal racemose panicles, the rachis and branches tomentellous. Flowers
ca 2.0 mm long, solitary or in small groups densely crowded on primary branches of
inflorescence. Bracts and bracteoles 0.5-I.0 mm long, triangular, persistent, puberu-
lous on exterior. Receptacle campanulate, sessile, tomentellous on exterior, tomentose
within. Calyx lobes acute, puberulous on both surfaces. Petals absent. Stamens 5-7,
unilateral; filaments shorter than calyx lobes, glabrous. Ovary inserted at base of
receptacle, tomentose. Style equalling filaments, pubescent throughout. Fruit globose
to pyriform; epicarp densely rufous-pubescent; pericarp rather thick, hard, fibrous,
lanate-hirsutulous within.
TYPES.HoehneI49i, Brazil, Sao Paulo, fl fr (lectotype, SP; isotypes, A, US;
syntype, B, lost): Sellow57z3, Brazil, Sao Paulo (paratype, nv).
DISTRIBUTION. Forests of eastern-central Brazil.
BRAZIL. Minas Gerais: Duarte 225i fr (RB 68906); Kuhlmann35 fr (RB I36935); Paldcioset al
3543 fl (R); Magalhdes2298 fr (NY). Espirito Santo: Kuhlmann364 fl (RB 136930). Sao Paulo: Handro
SP 39715 fl (FHO, LE, SP); Hoehne22997 fl (NY, SP); 29539 fl (FHO, SP, US); Pickel5705 fr (IPA,
SP); Prance10457 fl (NY); Roth 894 fl (SP).

LOCALNAME.Milho torrado.
Licaniahoehneiis closest to L. indurata,but also near L. kunthiana,differing in the
more pubescent inflorescence, the tomentellous pubescence of the fruit, and the
blunter leaves.

145. Licania cruegeriana Urban, Symb. Antill. 5: 352. I908.

Licaniacrassifolia
auct. non Bentham,Griseb.,Fl. Brit.W. Ind. 71. I864.

Tree, the young branches puberulous, becoming glabrous and lenticellate with
age. Leaves oblong to elliptic, coriaceous, 4.0-9.0 cm long, 2.0-3.8 cm broad, bluntly
acuminate or acute at apex, with acumen 2.0-8.0 mm long, cuneate to rounded at
base, glabrous above, persistently densely brown-lanate below, the reticulations
shallow; midrib plane above, glabrous when mature; primary veins 7-9 pairs,
prominent beneath, plane above; petioles 4.0-6.0 mm long, tomentellous even when
mature, eglandular, terete. Stipules linear, to 3.0 mm long, subpersistent,puberulous,
axillary. Inflorescences dense terminal and subterminal racemose panicles, the rachis
and branches tomentose. Flowers ca 2.0 mm long, solitary or in small groups densely
clustered on primary branches of inflorescence.Bracts and bracteoles to I.5 mm long,
triangular, subpersistent, puberulous on exterior. Receptacle campanulate, sessile,
tomentose on exterior, tomentose within. Calyx lobes acute, tomentellous on both
surfaces. Petals absent. Stamens 5-7, unilateral; filaments shorter than calyx lobes,
glabrous. Ovary inserted at base of receptacle, tomentose. Style equalling filaments,
pubescent throughout. Fruit cylindrical when young, becoming pyriformat maturity;
epicarp brown-tomentellous; pericarp thin, fibrous, hard, lanate-hirsutulous within.
TYPE. Crueger205, (2065 some sets). Trinidad, fr (GOET, K, NY, TRIN).
DISTRIBUTION.Forests of Trinidad and Tobago and northern coastal Venezuela.
Flowering September to January.
4409 fr (BM, F, GH, NY); 4462 fr (C, GH, MO, US). VENEZUELA.
TOBAGO.Broadway
Carabobo: Stelling 840o fr (US). Aragua: Pittier 15640 fl (US); Delgado88 fl (F, US, VEN); Pittier &
Nakichenovich15640 fl (VEN); 15742 fr (US, VEN); Steyermark 89727 fl (F, NY, US, VEN); LI. Williams
10333 fl (A, US, VEN).

LOCALNAME.Venezuela: Palo de Hierro.

This species is closely related to L. hoehneifrom southern Brazil, but differsin the
narrowly cylindrical immature fruit, the more pointed leaves, and the pubescent
petioles.
172 Flora Neotropica

146. Licania belemii Prance, sp nov Fig 26 A-E.

Arbor 20.0 m alta, ramulis juvenilibus glabris. Folia alternata petiolata, petiolo
Io.o-I2.0 mm longo, primo puberulo mox glabro, rugoso, leviter canaliculato;
laminae oblongo-ovatae, coriaceae, 5.0-9.5 cm longae, 2.6-4.5 cm latae, basi sub-
subcuneatae, apice in acuminem 3.0-7.0 mm longum contractae, supra glabrae,
subtus lanato-tomentosae, reticulatae; costa media supra plana, glabra; costis
secundariis 6-8 jugis, subtus prominentibus, supra planis. Stipulae non visae. Flores
2.0-2.5 mm longi in paniculis racemosis terminalibus axillaribusque dispositi, rachi
ramisque sparse tomentellis. Bracteae bracteolaeque 0.2-0.5 mm longae, persistentes,
tomentellae. Receptaculum campanulatum, sessile, extus tomentello-velutinum,
intus tomentosum. Calycis-lobi acuti, utrinque tomentelli. Petala nulla. Stamina 5-6
unilateralia; filamenta glabra, basi libera, calycis-lobis breviora. Ovarium ad basim
receptaculi insertum, tomentosum. Stylus e basi ovarii ortus, calycis-lobis brevior, ad
apicem pubescens. Fructus mihi ignoti.
TYPE.Belem& Pinheiro3211, Brazil, Bahia, Belmonte, fl (holotype, UB, isotype,
NY).
Known only from the type collection from coastal forest in Bahia.
DISTRIBUTION.
Collected in flower in January.
Licaniabelemiiis closest to L. hoehnei,but differs in the longer, broader leaves, the
larger petioles, the lax inflorescence, and the velutinous exterior of the receptacle and
calyx lobes. It is related to L. bahiensisbut differs in the caducous stipules, the more
reticulate leaf undersurface, the midrib which is plane above, and the velutinous-
brown pubescence of the exterior of the receptacle and calyx lobes.

Asiatic Species
Since there is only a single Asiatic species in the genus, this was included in the
keys in order to complete this treatment of Licania. It is to be fully described else-
where in a paper that will deal with the extra-American taxa of the largely New
World genera.

147. Licania splendens (Korthals) Prance, comb nov (see Prance, in press).

Angelesia splendensKorthals, Nederl. Kruidk. Arch. 3: 384. 1854; Boerlage & Koorders, Ic.
Bogor. 4: 59, t. 96. 90oi.
LicaniaangelesiaBlume, M6langes Bot. (1855) from Hasskarl, Flora 41: 256. 1858, nom illegit.
Trichocarya (Korthals)Miquel,Fl. Ind. Bat. I: 358. I855.
splendens
Chrysobalanus (Korthals)Miquel,Fl. Ind. Bat. i: 358. I855. in syn.
splendens
Parinarium nitidumHookerf., Fl. Brit. Ind. 2: 3IO. I878. Types. Maingay619, Malay Peninsula,
fl (syntype,K); Griffith
sn, (nv).
Coccomelianitida(Hookerf.) Ridley,Jour. As. Soc. Str. Branch72: I83. 1920.
Parinari Elmer,Leafl. Philipp.Bot. Io: 3809. 1939,nom nud.
philippinensis
Licaniasplendens(Korthals)Prance& Kosterm.,Reinwardtia7: I84. 1965,nom illegit.

TYPE.Korthalssn, Sumatra, fr (holotype, L).

DISTRIBUTION. Thailand, Malay Peninsula, Borneo, Sumatra, and Philippines.

Imperfectly Known Species (not included in key)

The following species were either described from poor material, or have not yet
been verified by me through examination of the type specimens.
Licania 173

D E

-' : . '
'::.:,:..'",.:?
,'..:'!:'7';"-:
""..
:~/.'~'~ i~~~~~z
~. /
??;~'~;"---.. ~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~
L /G/ J ~

'$...??...~~~/. .......

,- , ' -.
~
--.~.,/.~,
i!
'LAR E '~~

FIG 26. Speciesof Licaniaand Parinari.A-E, Licaniabelemii(Bele'm& Pinheiro 321I1); A, habit,

Xo-.; B, flower, x Io; C, flowersection, x io; D, ovary, X 25; E, ovary section, X 25. F-N, Parinari
chocoensis(Fuchs & Zanella 2I793); F, habit, x o.; G, flower, x 6; H, flower section x 6; J, ovary,
x 20; K, ovary section, x 20; L, fruit, x o.s; M, fruit section, x 0.5; N, fruit, section of middle part,
X 1.5.
174 Flora Neotropica

148. Licania veneralensis Cuatrecasas (section Moquilea),Fieldiana Bot. 27:

I09. I95I.

Tree, the young branches glabrescent. Leaves oblong-elliptic, coriaceous,

I2.0-I8.0 cm long, 4.0-6.0 cm broad, short-acuminate at apex, the acumen 2.0-5.0
mm long, rounded at base, glabrous on both surfaces; midrib prominent above,
glabrous; primary veins 12-14 pairs, prominent beneath, plane above; petioles
7.0-8.0 mm long, glabrous, scaly, terete, eglandular; palisade glands frequent,
bearing two large glands at base of lower surface of lamina. Stipules caducous (not
seen). Inflorescences terminal panicles, the rachis and branches ferrugineous-arach-
noid. Flowers on primary branches and secondary branches of inflorescence. Bracts
o.8-I.5 mm long, navicular, membraneous, glabrous; bracteoles to 3.0 mm long,
ovate, puberulous on exterior. Flower buds globose, arachnoid-pubescent (mature
flowers not seen). Petals absent. Fruit unknown.
TYPE.Cuatrecasas 15786, Colombia, Valle, fl bud, (holotype, F; isotype, COL).
Although this species was described from poor material, it is certainly distinct.
It probably belongs to section Moquilea,but mature flowers are needed to be certain.
It does not appear to be very closely related to any other species.

149. Licania amapaensis Prance, sp nov Fig 27 A-E.

Arbor Io.o m alta, ramulis juvenilibus hispido-tomentosis. Folia alternata

petiolata; petiolo io.o-i6.o mm longo, tereti, hispido-tomentoso, cum glandulis
duobus sessilibus munito; laminae oblongae, chartaceae, 15.0-33.0 cm longae,
4.5-9.5 cm latae, apice in acuminem 3.0-I5.0 mm longum contractae, basi cuneatae,
subtus hirsutae, glandulosae; costa media supra plana, subtus prominente, sparse
hirsuta; costis secundariis I3-I7 jugis, supra impressis, subtus prominentibus sparse
hirsutis. Stipulae intrapetiolares, membranaceae, lanceolatae, persistentes, tomen-
tellae. Flores (ca 3.0 mm longi in alabastris) in paniculis terminalibus subterminali-
busque dispositi, rachi ramisque ferrugineo-tomentosis. Bracteae bracteolaeque
oblongae vel linearae, persistentes, tomentosae. Receptaculum campanulatum, extus
tomentosum, intus tomentosum; pedicelli circa o.5 mm longi, Calycis-lobi acuti,
extus tomentosi. Petala 5, glabra. Stamina o, in orbem 3/4 disposita, filamenta
glabra; anthera deltoidea. Ovarium ad basim receptaculi insertum, pilosum. Stylus
e basi ovarii ortus, puberulus. Fructus mihi ignoti.
TYPE. Pires, Rodrigues& Irvine51198, Brazil, Amapa; Rio Araguari, Serra do
Navio, fl (holotype, NY; isotypes, IAN, INPA, MG).
Known only from the type gathering, from non-flooded forest.
DISTRIBUTION.
This most distinct species is of uncertain position within the genus as all available
material is in early bud. It belongs to the subgenus Moquilea,but the section can be
determined only when mature flowers are collected. Notwithstanding the lack of
open flowers, description of this species seems in order here. Licaniaamapaensis is easily
recognized by the large leaves with impressed venation, and the large tomentose
stipules.

150. Licania tepuiensis Prance, sp nov Fig 25 A-C.

Arbor parva, ramulisjuvenilibus tomentellis, mox glabris. Folia alternata petio-

lata, petiolo 2.0-3.0 mm longo, tomentello, subt-tlscuir glandulis duobus obscuris
Licania 175

*x 0-5; section, xx4;

o.5; G, flower section, H flower, x 4-
4~;H,
176 Flora Neotropica

munito, tereti; laminae orbiculares, crasse coriaceae, 1.6-4.5 cm longae, 1.7-3.8 cm

latae, basi cordatae, apice rotundatae vel retusae, supra glabra, subtus cum cavis
stomatalis profundis, faucis cavorum lanato-pubescentibus; costa media supra glabra,
leviter impressa; costis secundariis 7-9 jugis, subtus leviter prominentibus, supra
planis. Stipulae caducae (haud visae). Flores mihi ignoti. Drupa ovoidea, circa 4.2 cm
longa x 2.2 cm lata, extus laevis, glabra; pericarpio crasso, duro circa 6.o mm lato,
fibroso, intus sparse pubescente.
TYPE.Maguire33389, Venezuela, Bolivar, Ilu-Tepui, Gran Sabana, fr (holotype,
NY).
DISTRIBUTION. Known only from the type gathering from low open woodland
at edge of mesa, 4500' elevation.
Even though the flowers are not yet known, I have described L. tepuiensis because
it is so distinct in other characters. It differs from all other species of Licaniapossessing
stomatal cavities by the small orbicular leaves with cordate bases. The unusually
thick pericarp of the fruit also distinguishes it from most species. Judging from the
fruit and leaves with such well developed stomatal cavities L. tepuiensisprobably
belongs to section Leptobalanus.

I5I. Licania obtusifolia Fritsch (subgenus Parinariopsis), Ann. Naturh. Mus.

Wien 4: 53. 1889.
MoquileaparvifloraBlume, Mus. Bot. Lugd. Bat. 2: 92. 1856.
Licaniaparviflora(Blume) Lemee, Fl. Guyane Frang. 2: 23. 1952, nom illegit non L. parviflora
Benth.

Tree, the young branches glabrous, lenticellate. Leaves oval to obovate, coriace-
ous, acute at base, the young leaves arachnoid-lanate, becoming prominently reti-
culate with age; upper surface of midrib impressed; petioles shallowly canaliculate.
Stipules intrapetiolar. Inflorescences paniculate, the rachis and branches velutinous.
Bracts and bracteoles lanceolate to ovate, persistent, pubescent on exterior. Recep-
tacle campanulate, tomentose on exterior, hirsutulous within. Calyx lobes acute.
Petals 5, ciliate. Stamens ca 20, inserted in a complete circle, slightly connate at base;
filaments equalling calyx lobes. Ovary inserted to the side of the receptacle, sparsely
pilose. Style villous at base. Fruit unknown.
TYPE. Hort. Bot. Paris sn, French Guiana (PCU missing, nv).
In his description of this species, Blume wrote "In Guyana Gallica circa ripas
fluminis Ojapok, unde specimina ex Horto botanico Parisiensi bene vole mecum sunt
communicata." Although the type specimen should be in the Hortus Botanicus of
Paris, no specimen of L. obtusifoliacould be found in that herbarium. Thus, the type
is apparently missing and without it this species will be hard to place with certainty.
Judging from the description of Blume, it is a member of subgenus Parinariopsis.The
stamen number and exsertion, the lateral ovary, and the impressed venation all
to which this species may possibly be assigned in due course.
suggest L. licaniaeflora,

152. Licania roraimensis Standley (section Licania), Bull. Torrey Club 47:
286. 1940.

Tree, the young branches puberulous, becoming glabrous and lenticellate with
age. Leaves elliptic, coriaceous, 9.0-I3.0 cm long, 4.5-7.0 cm broad, acuminate at
Licania 177

apex, with acumen 8.0-15.0 mm long, rounded at base, glabrous above, lower surface
with well developed stomatal cavities filled with lanate pubescence, the reticulations
glabrous; midrib glabrous above, plane; primary veins 5-6 pairs, prominent beneath
plane above; petioles 7.0-Io.o mm long, pubescent, becoming glabrous with age,
eglandular, canaliculate. Stipules caducous (not seen). Inflorescences terminal
panicles, the rachis and branches puberulous. Flowers not seen. Fruit (immature)
with a tomentose epicarp.
TYPE.Pinkus6i, Guyana, fr (holotype, F; isotypes, GH, K, NY, S, US).
This species was described from very poor material, and no new material
and L. parviflora.
is related to L. triandra,L. bracteata
matches it. Licaniaroraimensis
Names Without Descriptions
The following names were published by Klotzsch in Schomburgk, Faun. et Fl.
Guian. 1198-9. 1848. They had no description and are hence nominanuda.Since they
have occasionally appeared in the literature, they are listed here, together with
equivalents, where known.
L. conduplicataKlotzsch = ?L. apetala(E. Mey.) Fritsch
L. flavicansKlotzsch =L. compactaFritsch (Schomburgk 812, K).
L. guianensisKlotzsch =L. heteromorpha Benth. (Schonzburgk
I599, BR).
L. macrophylla Klotzsch =L. laxifloraFritsch (Schomburgk
54, K).
L. odorataKlotzsch
L. schomburgkiana Klotzsch =L. affinisFritsch (SchomburgkI36r, K).
L. surinamensisKlotzsch

Excluded Names and Species (New World only)

A. Licania
i. L. aubletiana (Blume) Lemee, Fl. Guyane Fran9. 2: 23. I952. =Acioa gtianensis
Aubl.
2. L. celativeniaStandl., Publ. Field Mus. Bot. 17: 254. I937. = Vantaneacelativenia
(Standl.) Cuatr. (Humiriaceae)
3. L. columbarum Stehle & Quentin, Fl. Guadeloupe & Martinique 2(3): 50. I949.
= Terminaliasp (Combretaceae).
4. L. exiguifoliaStandl., Bull. Torrey Club 67: 285. I940. =Couepiacognata(Steud.)
Fritsch.
5. L. oliganthaA.. C. Smith, Jour. Arnold Arb. 28: 33. I947. =Chrysobalanus
cuspidatus
Griseb.

B. Moquilea
6. M. aubletianaBlume, Mus. Bot. Lugd. Bat. 2: 92. I856. =Acioa guianensisAubl.
7. M. bracteosa(Benth.) Walp., Rep. 2: 7. I843. =Couepiabracteosa Benth.
8. M. canomensisMart., Nov. Gen. et Sp. P1. 2: 8o. I827. =Couepiacanomensis
(Mart.)
Benth. ex Hook. f.
9. M. chrysocalyxPoepp. & Endl., Nov. Gen. et Sp. 3: 75. I845. =Couepiachrysocalyx
(Poepp. & Endl.) Benth. ex Hook. f.
IO. M. comosa(Benth.) Walp., Rep. 2: 6. I843. =CouepiacomosaBenth.
I . M. couepiaSteud., Nom. ed 2. 2: 159. 1849. =CouepiaguianensisAubl.
12. M. eliti (Mart. & Zucc.) Walp., Rep. 2: 6. I843. =Couepiauiti (Mart. & Zucc.)
Benth. ex Hook. f.
13. M. glandulosa (Miq.) Walp., Ann. 2: 463. 1852. =Couepia glandulosa Miq.
14. M. grandifloraMart. & Zucc., Abh. Akad. Miinchen I: 388. I832. =Couepia
grandiflora(Mart. & Zucc.) Benth. ex Hook. f.
I5. M. hypoleuca Miq., Linnaea I9: 438. I847. =Couepiaovalifolia(Schott) Benth.
178 Flora Neotropica

I6. M. inaequalisPoepp. ex Fritsch, nom. nud. in syn., Ann. Naturh. Mus. Wien 5:
I2. I889. =Couepiasubcordata Benth. ex Hook. f.
17. M. kunthianaMart. & Zucc., Abh. Akad. Munchen i: 390. I832. =Couepia
polyandra(Kunth) Rose.
18. M. multiflora(Benth.) Walp., Rep. 2: 7. 1843. =CouepiamultifloraBenth.
19. M. paraensisMart. & Zucc., Abh. Akad. Munchen I: 390. I832. =Couepia
paraensis(Mart. & Zucc.) Benth. ex Hook. f.
20. M. parilla (DC.) Steud., Nom. ed. . 22: 159. I84I. =Couepiaparillo DC.
21. M. rufaBarb. Rodr., Hort. Flumin. 165. I895. =Couepiarufa Ducke and Couepia
bracteosaBenth.
22. M. steudeliana(Miq.) Walp., Ann. 2: 463. 1852. =Couepiacognata(Steud.) Fritsch
23. M. uiti Mart. & Zucc., Abh. Akad. Mtinchen I: 390. I832. =Couepiauiti (Mart.
& Zucc.) Benth. ex Hook. f.

Tribe 2. Hirtelleae Prance, in press (Genera 3-8).

Ovary inserted at least half-way up the receptacle, or at its mouth.

3. Parinari Aublet, P1. Guiane I: 514, t. 204-206. I775; Kleinhoonte in Pulle,

Fl. Suriname 2(I): 454. I939; Kosterm., Reinwardtia 7: I47. I965. Prance,
in press.
ParinariumJussieu,Gen. P1. 342. 1789; DC. Prodr. 2: 526. 1825 pro parte quoad sectio Petrocarya
tantum; Meisner, P1. Vasc. Gen. I: 102. 1836 pro parte; Endl., Gen. 1251. 1840 pro parte;
Hooker f. in Benth. & Hook., Gen. P1. i: 607. I865 pro parte; C. Muell. in Walp. Ann.
Bot. Syst. 4: 644. I857 pro parte quoad sectio Euparinariumtantum; Hooker f., Mart. Fl.
Bras. 14(2): 49. 1867 pro parte; Focke in Engl. & Prantl, Nat. Pflanzenfam. 3(3): 53. 1894.
Dugortia Scopoli, Introd. 217. 1777.
PetrocaryaSchreber, Linn., Gen. P1. ed. 8. I: 245. I789.
BalantiumDesvaux ex Hamilton, Prod. Fl. Ind. Occ. 34. I825.
LepidocarpaKorthals, Nederl. Kruidk. Arch. 3: 385, 1855.
Ferolia sensu Kuntze, Rev. Gen. 2I6. 1891 pro parte, quoad Parinari sensu mihi tantum non
FeroliaAublet.

Small to large trees or shrubs, occasionally suffrutices. Flowers hermaphrodite

4.o-7.o(-I.o ) mm long. Leaves entire, glabrous above with stomatal cavities filled
with lanate pubescence on lower surface (except in a few Asiatic species). Petioles
usually with two sessile glands. Bracts and bracteoles eglandular, usually enclosing
young flowers in small groups. Inflorescences much branched panicles. Receptacle
turbinate to campanulate, slightly swollen at one side, hollow, pubescent throughout
within, tomentose on exterior. Calyx lobes 5, acute. Petals 5. Stamens 6-8; filaments
not exceeding the calyx lobes, unilateral with staminodes inserted opposite them.
Ovary inserted laterally at the mouth of the receptacle, the carpels bilocular. Style
filiform, not exceeding the calyx lobes. Fruit a fleshy drupe with a verrucose epicarp;
mesocarp fleshy and often fibrous; endocarp hard, thick, and with a rough fibrous
surface which may be channelled, with two basal plugs or stoppers the detachment of
which allows the seedling to escape. Germination hypogeal, the first leaves alternate.
TYPE SPECIES.Parinari campestrisAublet. The name Parznari is derived from the
vernacular name in French Guiana.
DISTRIBUTION. Pan-tropical, in the American tropics from Colombia through the
Guianas, Amazon basin to southern Brazil and Bolivia; also in Trinidad.
In my synopsis (Prance in press) the circumscriptionof Parinariwas considerably
altered. However, this redefinition of the genus makes little change to the New World
Parinari 179

representatives, except for the segregation of P. barbata,P. cordata,P. coriacea,P.

gardneri,and P. gracileinto the new genus Exellodendron.
Most genera in the Chrysobalanaceae contain groups of closely related species,
but the greater number of species in each genus are well defined or taxonomically
isolated. This is not the case in Parinari,where it is almost true to say that the whole
genus is one large species complex distributed around the tropics. There are a few
very distinct species (eg P. congolanain Africa, P. canarioidesin Asia, and P. campestris
and P. montanain America). The majority of species are, however, so closely related
that they can be separated only by differential characters and a few weak diagnostic
ones. These species are usually geographically and ecologically distinct from one
another. This has been discussed elsewhere (Prance, in press).
Taxonomic History of Parinari
Parinariwas described by Aublet ( 775), who named the two species P. campestris
and P. montana.Parinaricampestriswas described with reasonable accuracy, but not
P. montana.The latter species was erroneously depicted in Aublet's plate with the
ovary inserted at the base of the receptacle tube and with a lateral style. In addition
to this it contained parts of two species, (P. montanaand P. rodolphiias defined here).
Scopoli (I777) described the genus Dugortiawhich is synonymous with Parinari.
This was realized by Necker (I 790), who cited Parinarias a synonym of Dugortia.
Jussieu (I789), in his GeneraPlantarum,Latinized the Aublet name to Parinarium.
Until recently, later authors describing new species used Parinariumrather than
Parinari.However, according to current rules of nomenclature, Parinariis correct and
not Jussieu's orthographic variant. Several authors appear to have even concluded
that Parinari Aublet and ParinariumJussieu were different genera. For example,
Walpers (I843) in his summary of the species of the genus Parinarium Juss. did not list
the two Aublet species. Miers (1879) stated explicitly that the two genera are different
and thought that Aublet species were Licanias. His conclusions were based largely on
Aublet's misinterpretationsof the flower of P. montana.
Jussieu was the first to connect the American Parinariwith its congeners of an-
other continent. He cited two Adansonian genera from Senegal, Mampataand JNeou,
as synonyms of Parinari.Both of these names are nomina nuda, but Mampatacorres-
ponds with Parinari,and Neou with the more recent segregate of Parinari,Neocarya
Prance (in press). Also in 1789, Schreber, in his edition of GeneraPlantarum,described
Parinariof Aublet under the new name Petrocarya.
In I824 Sabine described two African species of Parinari,P. macrophylla and P.
excelsa.The latter species also occurs in South America. De Candolle (I825), in the
Prodromus, divided Parinariinto two sections. Section Petrocarya included Aublet's two
New World species, and section Neocaryaincluded Sabine's two African species. With
the more adequate specimen material now available, it is found that these two sections
are biologically not valid. Desvaux (i825) described the genus Balantiumfrom Guiana,
which was reduced to synonymy under Parinariby Bentham (I840) in his useful
contribution to the knowledge of the genus. Bentham described two new American
species, P. brachystachyum and P. coriaceum, and placed both in his section Petrocarya.
The latter species has now been transferred to Exellodendron.
On publishing his new genus Cyclandrophora from Asia, Hasskarl (1842) quickly
reconsidered and in I843 transferredhis species C. glaberrima to Parinari.Hasskarlwas
is now clearly
thus the first to link the Aublet name to an Asiatic species. Cyclandrophora
recognized as separatefrom Parinari(Prance, in press).However, one Asiatic species of
Parinarisens. strict. was described as PetrocaryasumatranaJack (1822) prior to the
publication of Cyclandrophora.
From I843 onwards, when Parinariwas recognized on the continents of Africa,
180 Flora Neotropica

Asia and America, the number of species described began to grow rapidly. Parinari
soon grew by the acretion of all members of the Chrysobalanaceae with a bilocular
carpel. Details of the classification that resulted from the use of this single character
alone are given elsewhere (Prance, in press), and are not repeated here because
the New World representatives were largely unaffected. The only other notable
study of the New World Parinariwas Hooker's account (1867) for the FloraBrasiliensis,
in which seven new species were described.

Key to the American Species of Parinari

I. Stipules (5.0-)I0.0-40.0 mm long, persistent, semiamplexicaul.
2. Young branches with short soft pubescence; leaf base rounded to cordate.
I. P. campestris.
2. Young branches with dense stiff ferrugineous-brown hairs; leaf base cuneate to subcuneate.
3. P. rodolphii.
i. Stipules small 1.5-4.0 mm long, caducous, axillary and not clasping stem.
3. Leaf undersurface white-lanate, the pubescence obscuring the stomatal cavities; suffrutex
or small shrub. I6. P. obtusifolia.
3. Leafundersurface with gray-brown pubescence not entirely obscuring the stomatal cavities;
large shrubs or trees.
4. Midrib distinctly impressed on leaf upper surface for entire length.
5. Leaf apex rounded to apiculate or bluntly acuminate on some leaves, with acumen to
3.0 mm long; primary leaf veins 9-20 pairs.
6. Leaves orbicular to broadly elliptic, 2.5-5.0 cm long, 1.5-3.5 cm broad, the primary
veins 9-11 pairs. o1. P. maguirei.
6. Leaves elliptic, 6.5-7.5 cm long, 3.0-4.0 cm broad, the primary veins 15-20 pairs.
I. P. littoralis.
5. Leaf apex with a well developed acumen 7.0- 5.0 mm long; primary leaf veins 2 -27
pairs.
7. Leaves 9.0-I7.0 cm long; petioles usually with two pairs of glands; flowers 6.0-
9.0 mm long; primary leaf veins more than 3.0 mm apart. 2. P. montana.
7. Leaves 2.0-7.5 cm long; petioles without distinct glands; flowers 5.o-6.o mm
long; primary leaf veins 1.0-2.0 mm apart. I2. P. parvifolia.
4. Midrib impressed only on lower portion or not impressed.
8. Leaf base distinctly cordate. (Also P. romeroiPrance, see Supplement) 13. P. cardiophylla.
8. Leaf base rounded to cuneate.
9. Primary leaf veins 28-35. 14. P. parilis.
9. Primary leaf veins 12-26.
Io. Inflorescence a dense many-flowered corymbose panicle.
i. Inflorescence and flowers with yellow-brown pubescence; leaf apex acute
or short-acuminate, the acumen 0-4.0 mm long; young branches with short
pubescence only. 5. P. occidentalis.
II. Inflorescence and flowers with silver-gray pubescence; leaf apex with
acumen 8.0-12.0 mm long; young branches with both short pubescence
and long stiff hairs. 9. P. klugii.
o0. Inflorescence a lax panicle, not corymbose.
12. Leaves ovate, with a well developed slender acumen 9.0-I6.o mm long.
6. P. sprucei.
12. Leaves elliptic to oblong-lanceolate, the acumen 0-12.0 mm long.
I3. Primary veins 22-26 pairs; leaves 7.5-16.0 cm long.
I4. Upper surface of midrib slightly prominent; petioles terete.
8. P. brasiliensis.
14. Upper surface of midrib slightly impressed on lower portion; petioles
canaliculate 15. P. chocoensis.
13. Primary veins 16-21 pairs; leaves 3.0-9.5 cm long.
I5. Young petioles canaliculate; leaf reticulation prominent on upper
surface. 7. P. pachyphylla.
15. Young petioles terete; leaf reticulations inconspicuous on upper
surface. 4. P. excelsa.
Parinari 181

The American Species of Parinari

i. Parinari campestris Aublet, P1. Guiane I: 514, t. 206. 1775; DC., Prodr.
2: 527. I825; Sandw., Kew Bull. 1931: 375. 193I; Kleinh. in Pulle, Fl
Suriname 2(I): 455. I939.
Petrocaryacampestris(Aublet) Willdenow, Linn. Sp. PI., ed. 4. 2: 287. I8oo.
Balantium cordifoliumDesvaux ex Hamilton, Prod. Fl. Ind. Occ. 34. 1825. Type. Herb.
Desvauxsn fl (holotype, P).
Ferolia campestris(Aublet) Kuntze, Rev. Gen. 216. 1891.

Tree to 25.0 m tall, the young branches villous, becoming glabrous and grayish
with age. Leaves ovate, 6.0-I3.0 cm long, 3.0-6.5 cm broad, cordate at base, acumi-
nate at apex, with acumen 7.0-12.0 mm long, glabrous above, tomentose beneath;
primary veins I4-I6 pairs, prominent beneath; midrib slightly impressed above;
petioles 2.0-4.0 (-7.0) mm long, pubescent when young, terete, the glands incon-
spicuous. Two glands often present on young branches beneath point of petiole
abscission. Stipules broad, to 3.0 cm long, acute at apex, villous, semiamplexicaul,
persistent. Inflorescencesterminal and axillary panicles, the rachis and branches with
short brown pubescence. Receptacle subcampanulate-turbinate, short-brown-
tomentellous on exterior. Petals 5, white, shorter than calyx lobes. Stamens 7 fertile,
unilateral, with 7-8 filamentous staminodes opposite them. Ovary and lower portion
of style densely hirsute. Fruit oblong, 4.0-6.0 cm long, 2.0-3.0 cm broad; epicarp
lenticellate; mesocarp thin, fleshy; endocarp thick and hard, densely tomentose
within.
TYPE. Aublet sn, French Guiana, fr (BM).
DISTRIBUTION. River margins, savanna margins, and open forests in Trinidad,
the Guianas, and adjacent Venezuela and Brazil. Flowering continuously.
VENEZUELA. Monagas: Wurdack& Monachino39414 st (NY). Terr. Delta Amacuro: Budowski
99 fl (F); Little & Petit 15991 fl (VEN); Steyermark87493 st (F, NY, US); Wurdack352 fr (F, K, US).
Bolivar: Rusby& Squiressn fl (NY); 423a st (NY); Steyermark 86451 st (VEN). Terr. Amazonas: Foldats
3750 fl (NY, US, VEN); Level 134 fl (NY, US); Maguire, Wurdack& Bunting37420 fl (NY, US); 36545
fr (NY, US); Maguire, Wurdack& Maguire41936 fl fr (MICH, MO, NY, US); 42630 fr (F, NY, US);
LI. Williams 14048 fr (F, VEN); Wurdack& Adderley42885 fl (MICH, NY, US); 41352 fl (MICH, NY,
P, US). TRINIDAD. Broadway6i60 st (K); Britton, Hazen & Mendelson614 st (K, NY, TRIN);
Brooks12257 fl (K, TRIN); 12473 fl (NY, TRIN); Cruegersn fr (K); Dannousesn fl (TRIN); Eggers1051
fl (BR, GOET, LE, P); 1362 fl (K, US); Prance2111 st (NY); Richardson644 st (K, TRIN); R. O.
Williams 12432 st (K, TRIN). GUYANA. Davis i43(F.D. 2134) fl (K); De La Cruz 1195 fl (GH, NY,
US); 1713 fl (F, GH, MO, NY, US); 2370 fl (F, GH, MO, NY, US); 2720 fl (F, GH, MO, NY, US);
2780 fl (F, GH, MO, NY, US); 2857 fl (NY); 4595 fl (F, GH, K, MO, NY); Jenman3979 st (K); 4015
fl (BM, F, K, NY, US); 5073 fl (BM); 6625 fl (K, NY); 7731 fl (K); Persaud58 fr (F, K, NY); rIo fl
(BM, F, K, NY); 209 fr (F); Sandwith184 fl (K, NY, P, RB, US); Schomburgk 535 fl (BM, GH, K, NY);
564 fl (OXF, P); 848 fl (BR); 980 fl (BM, P); Spence52 fl (K). SURINAME. B. W. 3033 fl (A); 3174 fl
(IAN, K, MO, NY); 3527 fr (A); 3557 fr (IAN, K, NY); 4018 fl (A); 4351 fl (IAN, K, NY); 4422 fr
(MO); 4929 fl (A, MO); 5401 fl (IAN, K, MO, NY); Irwin, Prance,Soderstrom& Holmgren55226 fl
(F, NY, US); 55374 fl (F, GH, NY, RB, US); 55454 fl (F, MO, NY, US); 55995 fl fr (GH, MICH,
MO, NY, US); Kappler1824 fl (C, GOET, LE, P); Kegel 1500 fl (GOET); Lanjouw& Lindeman805 fl
(IAN, K, NY); 1307 fl (IAN, K, NY); Tresling396 fl (MG); Stahel 84 fl (A, IAN, K, NY). FRENCH
GUIANA. Benoist 270 fr (P); 317 st (P); 994 fl (P); 1519 fl (P); Black & Klein 54-17160 fr (NY);
For. Serv.34M st (P); 192M fl (P); 7633 st (P); 7700 st (P); Lemoine7824 fl (NY, P); Martin sn st (BM,
BR, F, K); Melinonsn fl (A, BM, F, K, NY, P, R, US); 4 fl (P); 46 fl (P); 187 fr (A); Poiteausn fl
(K, LE, P); Sagot 204 fl (BM, K, NY, P); Wachenheim12 fl (K, P); iIr st (P); 129 st (P). BRAZIL.
Amazonas: Frdes 23189 fr (IAN); Schultes9929 fl (IAN); 10029 fl (GH, IAN). Terr. Roraima: Prance
et al 4366 st (NY); 4601 fr (NY); 9503 st (INPA, NY). Para: Bastos i fl (RB I36942); Ducke 1265 fr
(IAN, K, MG 17779, NY, R, US); RB 18226 fl (K, NY, P, RB, US); Killip & Smith 30546 fr (A,
NY); KuhlmannRB 21027 fr (K); Oliveira4563 st (IAN, NY); 4706 st (IAN, NY); Poeppig3007 st
(F, GOET, LE, W). Terr. Amapa: Black 49-8311 fl (IAN, NY); Pires, Rodrigues& Irvine 50448 fr
(NY). Maranhao: Ducke781 fl (BM, MG); Froes31944 fr (IAN).
182 Flora Neotropica

LOCAL NAMES. Venezuela: Merecurillo, Guaray. Trinidad: Bois Baude. Guyana:

Buhurada,MahaicaBalli. Suriname: Boehoeroedoe, Boehoerada, Bohowoda,Foengoe,Tajama-
ma, Vonkhout, Koepesienoe.French Guiana: Foungouti.
This species is chosen as the generic type in preference to P. montanabecause of
the confusion over the circumscription of the latter species. This species, one of the
most distinct in the genus, is easily recognized by the large semiamplexicaul stipules
and the cordate leaf bases.

2. Parinari montana Aublet emend Ducke Fig 28 E-H.

ParinarimontanaAublet, P1. Guiane 1: 514, t. 205. 1775 pro parte quoad fructum tantum.
ParinarimontanaAublet emend Ducke, Revue Bot. Appl. Agr. Trop. 15: 8o. 935.
Petrocaryamontana(Aublet) Willdenow, Linn. Sp. P1. ed. 4. 287. I880 pro parte.
Chrysobalanus montanus(Aublet) G6mez de la Maza, Fl. Cuba 39. I887.
Feroliamontana(Aublet) Kuntze, Rev. Gen. 216. I891 pro parte.
Parinaripajura R. Benoist, Bull. Mus. Hist. Nat. Paris 28: 232. I922. Type. Huber MG 7045,
Brazil, Para, fl (holotype, P; isotypes, BM, G, K, MG, R).

Tree to 40.0 m tall, the young branches pilose, becoming glabrous and lenticel-
late with age. Leaves oblong to oblong-lanceolate, 9.0-I7.0 cm long, 3.o-6.5 cm
broad, cuneate to subcuneate at base, acuminate at apex, with acumen 5.0-9.0 mm
long, glabrous above, tomentose and with stomatal cavities beneath; upper surface of
midrib impressed, lower portion pubescent, with two pairs of medial glands,
terete. Stipules I.0-3.0 mm long, caducous. Inflorescences terminal and axillary
panicles 4.5-I .0 cm long, the rachis and branches short-tomentellous. Receptacle
subcampanulate-turbinate, yellow-brown-tomentellous on exterior. Petals 5, white,
shorter than calyx lobes. Stamens 7 fertile, with 7-8 staminodes opposite them. Ovary
and lower portion of style pilose. Fruits globose to oblong, up to Io.o cm in diameter;
epicarp densely verrucose; mesocarp thin and fleshy; endocarp hard and very thick,
the exterior deeply ridged and spikey, lanate within.
TYPE.Aubletsn, French Guiana, fr (BM fruit only, not vegetative material).
DISTRIBUTION. Non-flooded forest in French Guiana and northern Brazilian
Amazonia. Flowering June to November.
BRAZIL. Amazonas: Ducke 2279 fl (F, IAN, INPA, MG, NY, R, RB 71444, SP); RB 18225 fl
(K, RB, US); Froes 23319 fl (IAN); Rodrigues2699 fr (INPA 17004); 7699 fr (INPA 17004, NY);
Rodrigues& Loureiro7129 fr (INPA 15791, NY). Pari: Pires & Silva 1094I fr (IAN, NY); Io942 fr
(IAN, NY). Terr. Amap,: Cowan38269 fr (F, GH, NY, US).

LOCALNAMES.Brazil: Castanheirade pedra, Pajura de pedra.

The identity of Aublet's species P. montanahas been a matter of dispute for many
years, having been commented on by Huber (1904), Ducke (1935), Sandwith ( 93 ),
and Benoist (1922). Although none of the above workersappeared to have had all the
facts together, Ducke arrived at the correct solution by comparing the herbarium
material available to him in Brazil with Aublet's illustrations. In light of more recent
collections and my own consultation of the Aublet specimen at the British Museum,
it is now possible to clarify the position.
Ducke suggested that Aublet had confused two species in his original description
of P. montanaby using the fruit of one species and the sterile branches of another. Huber
and Ducke both adopted the fruit as the part to bear the name P. montana.The endo-
carp of the fruit figured by Aublet is distinct with deep furrows and large irregular
verrucose teeth (Fig 28 G-H), and is not easily confused with any other species of
Parinari. I have observed these characters in the specimen collected and used by
Ducke (Ducke 2279) and others in the Museu Goeldi, in the Aublet fruit in the
Parinari 183

carpological collection at the British Museum, and in a recent collection, Cowan

38269, from Serra do Navio, Terr. Amapa, Brazil. The leaves of all the post-Aublet
gatherings correspond exactly with the type specimen of P. pajura,which as Ducke
suggested, must be regarded as a synonym of P. montana.Benoist and Sandwith both
referredto P. montana in this sense under the name P. pajurasince they realized that it
is distinct from what had usually been termed P. montana,ie the species that corres-
ponds with the leaves described and figured by Aublet. This latter species was
described by Huber (I9Io) as P. rodolphii,which name must be retained, although
Huber was not fully aware of the problem when he described P. rodolphii.The dis-
tinctions between the two species P. montanaand P. rodolphiiwere elucidated in some
detail by Ducke. The two species differ not only in the type of endocarp but also in
the shape and size of the leaves and in the stipules (Fig 28).

3. Parinari rodolphii Huber, Bol. Mus. Emilio Goeldi 6: 77. I91o; Ducke
Rev. Bot. Appl. Agr. Trop. 15: i8i. 1935. Fig. 28 A-D.
ParinarimontanaAublet, P1. Guiane I: 514, t. 204. 1775 pro parte quoad ramulum et folia tantum.
Type: Aubletsn, French Guiana, st (BM, leaves only).
ParinarilucidissimaStandley, Lloydia 2: 183. 1939. Types. A. C. Smith3320, Guyana, fr (holotype,
F; isotypes, A, K, LE, MAD, MO, NY, P).
Petrocaryamontana(Aublet) Willdenow, Linn. Sp. PI., ed. 4. 2: 287. 800opro parte.
Feroliamontana(Aublet) Kuntze, Rev. Gen. 216. 1891 pro parte.

Tree to 40.0 m tall, the trunk often buttressedto about 2.0 m, the young branches
long and densely ferrugineous-pilose. Leaves elliptic to oblong-lanceolate, glabrous
above, tomentose and with stomatal cavities beneath, subcuneate to rounded at base,
acuminate at apex, with acumen 5.o-I5.0 mm long; primary veins 21-26 pairs,
prominent beneath, slightly impressed above; midrib impressed and usually pubes-
cent above; petioles 3.0-8.0 mm long, with two medial glands usually obscured by
dense pubescence, terete. Stipules to 4.0 cm long, semiamplexicaul, ferrugineous-
pubescent. Inflorescences terminal dense-flowered panicles to 6.0 cm long, the rachis
and branches ferrugineous-pilose. Receptacle subcampanulate-turbinate, brown-
tomentose on exterior. Petals 5, white, shorter than calyx lobes. Stamens 7 fertile,
with 7-8 short filamentous staminodes opposite them. Ovary and lower portion of
style densely pilose. Fruit ellipsoid, to 5.5 cm long; epicarp verrucose, glabrous;
mesocarp thin, fleshy; endocarp hard and thick, granular, fibrous, plane, not ridged
on exterior, lanate within.
TYPE. R. S. Rodrigues MG 9648, Brazil, Para, fl (holotype, MG; isotypes, K, P,
NY, R, RB).
DISTRIBUTION. Non-flooded forest or periodically flooded areas in the Guianas
and eastern Amazonian Brazil.
Representative collections: VENEZUELA. Terr. Delta Amacuro: Berti 84 fr (BR, NY, RB, US,
VEN); I74 fr (F, NY, VEN); 588 st (F, NY, US, VEN); Blanco 484 fl (NY); Steyermark87248 st (F,
NY, US, VEN); Wurdack& Monachino39675 fl (F, GH, K, NY, RB, US); Zabala 63 fr (VEN). Bolivar:
Bernardi 2138 fl (NY, US, VEN); Little 17615 st (VEN); Steyermark86972 st (VEN). GUYANA.
Fanshawe28o(F.D. 3016) fl (K, NY); Wilson-Browne525(F.D. 5926) fl (NY). FRENCH GUIANA.
Irwin, Pires & Westra47210 st (F, GH, IAN, MG, MO, NY, US); Martinsn fl (BM); Richardsn st (P);
Wachenheim sn st (P). BRAZIL. Para: Black 49-8547 fl (IAN, P); Ducke 1619 fl (A, F, IAN, MG, NY,
R, US); MG 15305 st (MG); MG I5644 fl (MG); MG 15806 fl (MG); RB 18337 fl (RB); Krukoff
5862a fl (F, K, MICH, MO, NY, US); Kuhlmann& Jimbo 229 fr (IAN, MG 24014, SP); 328 fl (FHO,
IAN, MG 24109, SP, US); Pires 3193 st (IAN); 3z94 fr (IAN); 6925 fr (IAN); 7424 fl (UB); 51851 st
(NY); Pires & Silva 4640 fl (IAN, INPA, NY); 10901 fl (IAN, NY); Prance& Silva 58753 fl (MO, NY,
US); 58865 fl fr (F, NY, US); Prance & Pennington1994 fr (GH, NY); R. S. RodriguesMG 9684 fl (BM,
MG, R, US). Terr. AmapA: Black 49-8473 fr (IAN, NY); Pires, Rodrigues& Irvine51474 fl (F, IAN,
INPA, MG, NY, US).
184 Flora Neotropica

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

X 0o.5; B, flower section, x 5; C, fruit section, x o.5; D, fruit, mesocarp removed from top half, x o.5.
E-H, P. montana(Ducke 2279); E, habit, x o.5; F, flower section, x 5; G, fruit section, x o.5; H, fruit,
mesocarp removed from top half, x o.5.

LOCALNAMES. Venezuela: Merecurillo, Merecure.French Guiana: Parinari. Brazil:

Parinari,Farinhaseca.
For details of the nomenclature, see under P. montana.I agree with Fanshawe &
Maguire (Bull. Torrey Club 75: 382. I948) that P. lucidissimais synonymous with P.
rodolphii.
Parinari 185

4. Parinari excelsa Sabine, Trans. Hort. Soc. London 5: 45I. 1824; Kleinh.
in Pulle, Fl. Suriname 2(I): 455. I939; Fanshawe & Maguire, Bull. Torrey
Club 75: 382. 1948.
Parinaribrachystachya Bentham, Jour. Bot. Hooker 2: 213. 1840; Hook. f., Mart. Fl. Bras. 14(2):
52, t. i8. 1867. Type. Schomburgk 785, Guyana, fl (holotype, K; isotypes, BM, BR, CGE,
GH, L, NY, OXF, P, US).
Petrocaryaexcelsa(Sabine) Steudel, Nom. ed. 2. 2: 309. 1849. non P. excelsaJack.
Parinari laxifora Ducke, Arch. Jard. Bot. Rio de Janeiro 3: 44. 1922, synon nov. Types. Ducke
RB 11051, Brazil, Amazonas fl (K, US); RB I5244 fr (P).
Parinarilaxifloravar lata Ducke, Arch. Inst. Biol. Veg. Rio de Janeiro 2: 33. 1935. Type. Ducke
RB 24181, Brazil, Amazonas, fl (holotype, RB; isotype, NY).
Parinari glazioviana Warming, Kjoeb. Vidensk. Meddel. 1874: 72. 1874, synon nov. Types.
Glaziou 752, Brazil, Guanabara, fl (syntype, C; isosyntypes, BR, P); 2128 fl (syntype, C;
isosyntypes, BR, K, P, R); 2560 fl (isosyntypes, BR, F, FI, K, P).
ParinarisylvestrisKuhlmann, Loefgrenia 13: I. I964. Types. Kuhlmann2736, Brazil, Sao Paulo, fl
(holotype, SP 76011 ); 2738 fl (paratypes, SP, UB); 2354 fl (paratype, SP).
Feroliaexcelsa(Sabine) Kuntze, Rev. Gen. 216. 1891.
FeroliaamazonicaKuntze, Rev. Gen. 216. I89I6.

Tree to 40.0 m tall, the trunk often buttressed, the young branches sparsely
pubescent, becoming glabrous with age. Leaves ovate to oblong-elliptic 3.0-9.0 cm
long, 1.5-5.0 cm broad, rounded to cuneate at base, acuminate at apex, with acumen
2.0-Io.o mm long, glabrous above, tomentose and with stomatal cavities beneath;
primary veins 13-20 pairs, prominent beneath; midrib plane to prominulous above,
glabrous; petioles 3.0-7.0 mm long, pubescent when young, with two sessile medial
glands (often becoming inconspicuous with age), terete. Stipules ca I.O mm long,
caducous. Inflorescences terminal rather lax panicles, the rachis and branches with
light brown to rufous pubescence. Receptacle subcampanulate-turbinate, tomentose
on exterior, pedicels 1.0-2.0 mm long. Petals 5, white, shorter than calyx lobes.
Stamens 7 fertile, unilateral with 7-8 short filamentous staminodes opposite them.
Ovary and base of style pilose. Fruit ellipsoid, 2.5-4.0 cm long, 1.8-2.5 cm broad;
epicarp densely verrucose; mesocarp thin and fleshy; endocarp hard and thick, the
exterior granular, fibrous, and slightly ridged, densely lanate within.
TYPE. G. Don sn, Sierra Leone, (lectotype, K).
DISTRIBUTION. (In neotropics). Non-flooded forest from Amazonian Colombia
and eastern Venezuela through the Guianas and Amazonian Brazil to eastern-central
Brazil.
COLOMBIA. Caqueta: RomeroC. 4109 fr (COL). VENEZUELA. Terr. Delta Amacuro: Berti
77 fl (BR, F, NY, RB, US, VEN); 124 fl (BR, F, NY, RB, VEN); 315 fr (F, VEN); 566 fr (F, NY
VEN); 576 fr (F). Bolivar: BernardiI540 st (NY); Cardona1174 fl (NY, US, VEN); 1176 fl fr (US,
VEN); Steyermark86374 fr (VEN); 86468 fr (VEN); 88489 fr (VEN). GUYANA. Fanshawe6iz(F.D.
3347) fr (K, NY, P); 67o(F.D. 3406) fl (F, IAN, K, NY, US); For. Dept. 6383 st (US); 6384 st (US).
SURINAME. B. W. 6462 fl (IAN, K, NY); 6599 fl (A); 6777 fr (NY). FRENCH GUIANA. For. Serv.
25M fr (NY, U); Martin sn fl (K). BRAZIL. Amazonas: CoelhoINPA 1578 fl (IAN, INPA, MG, NY);
INPA 4099 fl (INPA, NY); Ducke534 fl (A, F, K, MO, NY, R, US); 1347 fl (A, IAN, K, MG 17775,
NY, R, US); 1544 fr (A, IAN, MG 17776, NY, R); 1727 fl (A, F, IAN, K, MG, NY, R, US); 1824 fr
(A, F, IAN, K, NY); 2145 fl (COL, GH, IAN, MG 17774, NY, P, R); 2418 fl (K); MG 7372 fl (BM,
MG, RB); Ducke MG 15244 fr (MG, US); RB 24181 fl (NY); RB 35555 fl (K, P, RB, US); Ferreira
57-I2 st (INPA 5061); Froes23738fr (IAN); 26396fl (IAN, NY, SP, US); 26457 fr (IAN, NY, P, UB);
26532 fl (FHO, IAN, NY, SP); GoeldiMG 3882 fl (BM, MG, P, RB, US); Pranceet al 3157 fr (INPA,
NY); 10365 fr (INPA, NY); Rodrigues& Osmarino6007 fl (INPA 15057, NY, RB); Spruce 1577 fl
(BM, GOET, K, LE, MG, NY, OXF, P, RB). Terr. Roraima: KuhlmannRB 3060 fl (K, RB, US);
Pranceet al 4131 fr (NY); 4270 fr (NY). Pari: Frdes31971 fl (COL, IAN, NY, P, RB, US, SP); 31819
fl (FHO, IAN, P, SP, US); Ducke MG o1884 fl (MG, P, RB, US); MG 11051 fl (P); MG 16648 fr
(BM, MG); RB 15037 fl (RB); HuberMG 1876 fr (MG); Oliveira493 fr (IAN); 2092 fl (IAN, UB);
3580 fr (IAN, NY); 3768 fr (IAN, NY); 4179 fr (IAN, NY); 4606 fr (IAN, NY); Silva 926 fl (IAN,
6 The numerous African
synonyms of this species are listed elsewhere (Prance, in press).
186 Flora Neotropica

NY). Terr. Amapl: Frdes & Black 27278 st (IAN); 27517 fl (IAN); 32896 fl (IAN); Pires & Cavalcante
52476 fl (F, IAN, MG, NY, US); Pires, Rodrigues& Irvine50349 fl (F, GH, IAN, MG, NY, US);
51558 fr (IAN, MG, NY, US). Maranhao: Frdes 1909 fr (A, F, MICH, MO, P, US); 1982 fl (A, F, P,
US). Terr. Rond6nia: Pranceet al 8821 fr (INPA, NY). Mato Grosso: Krukoff1501 fr (A, BM, K,
MICH, NY, P); Rondon2450 fl (R). Espirito Santo: Kuhlmann154 fr (RB 34329). Rio de Janeiro:
Armond175 fr (R); Glaziou 7602 fl (BR, K, LE, US); 9391 fl (BM, BR, K, LE, NY); 13796 fl (BR, K,
LE); Glaziou & Schwacke5689 fl (R); Goes & Constantino 943 fr (RB 51671); 1o46 fl (RB 51670, NY);
Pabst 5219 fl (HB); Schwacke256 fl (GOET); 4256 fl (RB 47973); St. Hilaire 13 fl (P); Sellow 299 fl (R)
Without precise locality Rio de Janeiro or Guanabara: Nadeaudsn fr (F, P); Riedel 1667 fl (NY, US);
Saldanha655 fr (R); Saldanha& Glaziou 4993 fl (R); Schwackesn fl (R); Widgrensn fl (S).

LOCAL NAMES AND USES. Venezuela: Guari, Makara-yek, Merecurillo. Guyana:

Aiamoradan(Arawak). Suriname: Rosenkwarie,Bokohoton,Wokerebesoerie.French
Guiana: Foungouti.Brazil: Uchirana,Uchide Tambagui.Fruit edible. The oil from the
pulp is used for perfuming hair. The wood is used in construction, but is very hard.
The IntercontinentalDistribution.Oliver (1871) was the first to suggest that the
African species P. excelsaSabine is conspecific with the American P. brachystachya
Benth. This position was followed by Kleinhoonte (i939). The African species of
Parinarihave recently been treated in some of the regional floras, and P. excelsahas
been more clearly defined (see Graham I957, I960). Several other species were re-
duced to synonymy under P. excelsaby various authors (for a list of synonomy see
Prance, in press). In AfricaP. excelsais a widespread foresttree ranging from Guinea to
Tanzania, Zambia and Mozambique. Close examination of material of P. excelsa
from Africa and of P. brachystachya from America reveals no differences at all. It is
also noteworthy that P. brachystachya in the Guianas tends to have rather brittle
membraneous leaves, a feature paralleled in the West African coastal samples of
P. excelsa,which was formerly separated into the species P. tenuifoliaA. Chev.
Parinariexcelsaand relatedspecies.Redefinition of P. excelsain Africa requires re-
consideration of the American element of this species. P. laxiflorawas separated from
P. excelsaby a few minor characters, none of which is constant. The name P. laxiflora
in fact merely represents the Amazonian material of P. excelsa.Parinariglazioviana
from around Rio de Janeiro, Brazil represents the austral extreme of P. excelsa.The
minor differences employed to distinguish these two geographic extremes are not
constant, and there is no morphological discontinuity to separate P. laxifloraor P.
glaziovianafrom P. excelsa.In South America P. excelsais a forest species ranging from
northern Venezuela to southern Brazil. In both Africa and America it is the most
widely distributed species in the genus.
Parinariexcelsabelongs to a complex of very similar species, including P. brasilien-
sis (incl. P. pohlii), P. pachyphylla,P. sprucei,and P. occidentalis.These species are
closely related, but are separable from one another by a number of differential
characters as well as by a few weak diagnostic ones. A summary of their differences
is given in the comparative notes below the description of each of these five species.
Comparative note for P. excelsa.Leaves ovate to oblong-elliptic, 3.9-9.0 cm long,
1.5-5.0 cm broad, rounded to cuneate at base, acuminate at apex, with acumen
i.o-Io.o mm long. Primary veins 13-22. Petioles terete. Inflorescences rather lax
panicles, light-brown-rufous-pubescent.

5. Parinari occidentalis Prance, sp nov Fig 29 A-G.

Arbor alta, ramulis juvenilibus pubescentibus mox glabris lenticellatis. Folia

alternata petiolata, petiolo 5.0-I2.0 mm longo, glanduloso, tomentoso; laminae
ellipticae ad oblongo-lanceolatae, 7.0-I0.5 cm longae, 2.5-5.5 cm latae, basi rotun-
Parinari 187

datae vel subcuneatae, apice acutae vel in acuminem brevem ad 4.0 mm longum
contractae, superne glabrae, subtus cum cavis stomatalis griseo-tomentosae; costis
secundariis I7-20 jugis, subtus prominentibus. Stipulae circa I.o mm longae, caducae.
Flores in paniculis terminalibus corymbosis ubique argenteo-brunneo-tomentosis
dispositi. Bracteae bracteolaeque ovatae, cum calice aequantes. Receptaculum sub-
campanulatum, extus argenteo-brunneo-tomentosum,intus ubique pilis longis dense
tomentosum. Petala 5. Stamina 7, unilateralia, staminodiis opposita. Ovarium vil-
losum. Stylus e basi ovarii ortus, basi villosus. Fructus globosus, 3.5 cm diametro,
extus verrucosus, endocarpio extus fibroso, intus dense lanato tomentoso.
I
TYPE. Krukoff I r3, Bolivia, Dept. La Paz, Larecaja Province, Copacabana, fl
(holotype, NY; isotypes, A, F, K, MICH, MO).
Upland forest in Bolivia and western Brazil.
DISTRIBUTION.
BOLIVIA. La Paz: Larecaja Prov., Krukoff11ooo fr (A, F, K, MICH, NY, US); 11278 fl (A, F,
K, MICH, NY, SP). BRAZIL. Acre: Near mouth of Rio Macauhan, Krukoff5650 fr (A, F, K, LE,
MO, NY, US).

Comparative note. Leaves elliptic to oblong-lanceolate, 7.0-I0.5 cm long,

2.5-5.5 cm broad, rounded to subcuneate at base, acute or shortly acuminate at apex,
with acumen 0-0.4 mm long. Primary veins 17-20. Petioles terete. Inflorescences
dense-flowered corymbose panicles, yellow-brown-villous.
This species is distinguished mainly by its corymbose inflorescence.

6. Parinari sprucei Hooker f., Mart. Fl. Bras. 14(2): 53. 1867.
Feroliasprucei(Hookerf.) Kuntze, Rev. Gen. 216. I891.

Tree to 20.0 m tall, the young branches villous becoming glabrous and lenticel-
late with age. Leaves ovate to oblong-ovate, 9.0-I5.0 cm long, 4.5-6.0 cm broad,
rounded at base, acuminate at apex, with acumen 8.0-i6.0 mm long, glabrous above,
tomentose beneath; primary veins I5-I8 pairs, prominent beneath; midrib level
above; petioles 7.0-I3.0 mm long, terete, pubescent when young, with two glands at
base. Stipules caducous (not seen). Inflorescencesfew-flowered axillary and terminal
panicles not more than 2.0 cm long, the rachis and branches with short brown
pubescence. Receptacle subcampanulate-turbinate, pubescent on exterior. Petals 5,
white, shorter than calyx lobes. Stamens 7 fertile, unilateral with 7 short filamentous
staminodes opposite them. Ovary and lower portion of style densely pilose. Fruit
oblong, 5.0-6.0 cm long, 2.5 cm broad; epicarp densely verrucose; mesocarp fleshy;
endocarp hard, thick, the exterior granular and fibrous, slightly channelled, lanate
within.
TYPE. Spruce 2539, Brazil, Amazonas, fl (holotype, K; isotypes, BM, BR, CGE,
GH, GOET, LD, LE, NY, OXF, P).
Riverine forest in the upper Rio Negro region of Brazil and ad-
DISTRIBUTION.
jacent Colombia and Venezuela.
COLOMBIA. Vichada: Giovannisn (COL). Vaupes: Cuatrecasas69r7 fl fr (COL, F, P, US);
Ferndndez2221 fr (COL, F, IAN, US); Schultes& Cabrera14577 fr (F, NY, US). VENEZUELA. Terr.
Amazonas: Wurdack& Adderley43266 fr (NY, US). BRAZIL. Amazonas: Black 48-2663 fr (IAN, NY);
Froes 22168 fr (IAN, NY, U).

LOCALNAMEAND USES.Colombia: Varete. Fruit edible.

Comparative note. Leaves ovate to oblong-ovate 9.0-15.0 cm long, 4.5-6.0 cm
broad, rounded at base, acuminate at apex, with acumen 8.o-I6.o mm long. Primary
veins I5-I8. Petioles terete. Inflorescences small axillary and terminal few-flowered
panicles with short brown pubescence.
188 Flora Neotropica

~~~~~a ~ ,' '~i

F D~~~I
..... ....

ILli
A, L4
~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~?IL.:?
E "
'

A~~~~~~~~~~ C

cr:~~~~~~~~~~~~~~~~~~~~~~~~~~;
N

:~~~~~~~~~~~~
~: . i \ \ II&j
FIG ~~~~~_
29 pce fPrnr.AG .ocdnai Krlof100 17) ,hbtxo5 lwr
x ~; C,flower ection,x ~; D petal,x 10; E ovaryand stye X 10 , frui, X o ~G leaf urface
(upr,lft
fritsetin,x
owr rgt)
.;
~ -L . auie (Mgireeo.K a 330
, ea upe srac, i MP,P
Hhbi,x
klgi (Ku 380, M,hbt
Jfui
,N lwr
>< 5; 0 flowe sectin, x ~ P, lef surfae (uppr abov, lowe beneah), x

FIG 29. Species of Parinari.A-G,P. occidentalis(Krukoffrrooo,r 2x78); A, habit, xo.-5; B, flower,

x 5; C, flower section, x 5; D, petal, x i o; E, ovary and style, x i o; F, fruit, x 0.5; G, leaf surfaces
(upper, left; lower, right), x 5. H-L, P. maguirei(Maguireet a12339o); H, habit, x 0.5; J, fruit, X 0.5; K,
fruit section, x 0.5; L, leaf upper surface, x i. M-P, P. klugii (Kflug387o); M, habit, x 0.5; N, flower,
x 5; O~ flower section, x 5; P, leaf surface (upper above, lower beneath), x 5.
Parinari 189

7. Parinari pachyphylla Rusby, Descr. New Sp. S. Am. P1. 26. 1920.

Tree to 30.0 m tall, the young branches pubescent, soon becoming glabrous and
lenticellate. Leaves elliptic to oblong, 3.0-9.5 cm long, 1.7-4.5 cm broad, cuneate to
subcuneate at base, bluntly acuminate at apex, with acumen I.o-4.0 mm long,
glabrous above, tomentose and with stomatal cavities beneath; midrib prominulous
above, glabrous; primary veins 16-21 pairs, prominent beneath; petioles 4.0-9.0 mm
pubescent, with two medial glands, shallowly canaliculate. Stipules ca I.o mm long,
caducous. Inflorescences terminal and axillary panicles to 9.0 cm long, rather lax,
with short light brown pubescence. Receptacle subcampanulate-turbinate, short-
pubescent on exterior. Petals 5, white, equalling calyx lobes. Stamens 6-7 fertile,
with 7-8 short filamentous staminodes opposite them. Ovary and base of style pilose.
Fruit ellipsoid, to 3.0 cm long; epicarp verrucose; mesocarp thin, fleshy; endocarp
hard, thick, with a fibrous granular and plane exterior, tomentose within.
TYPE. H. H. Smith i775, Colombia, Magdalena, fl (holotype, NY; isotypes, A,
F, FI, GH, K, L, MICH, P, US).
DISTRIBUTION. Riverine forest and upland forest in Northern Colombia and
Venezuela.
COLOMBIA. Bolivar: Curran139 st (GH, US). Magdalena: Cuatrecasas& RomeroC. 24876 fr
(US); Espina & GiacomettoB2-A st (A, COL, F, NY); RomeroC. 223 fl (COL); 686 fr (COL, US).
VENEZUELA. Zulia: Aristeguieta,Blanco & Carrillo6771 fr (NY). Falc6n: Steyermark& Braun9449I
st (NY, VEN); Carabobo: Pittier7999 fr (GH, US, VEN). Dist. Federal: Delgado 192 fr (F, US, VEN);
399 fr (US, VEN). M6rida: Bernardi1215 fl (COL, FI, K, NY, VEN). Aragua: Curran913 fr (NY);
Lasser895 fr (US, VEN); Pittier oo103 fr (GH, NY, US, VEN); 13556 fr (F, MO, US, VEN); 63556
fr (US); LI. Williams 9972 fl fr (F, K, US, VEN). Bolivar: Steyermark88480 fr (F, GH, NY, US);
Wurdack& Monachino41z43 fr (MICH, NY).

LOCALNAMES.Colombia: Pereguetano.Venezuela: Merecurede montaia, Guari.

Comparative note. Leaves elliptic to oblong, 3.0-9.5 cm long, 1.7-4.5 cm broad,
cuneate to subcuneate at base, bluntly acuminate at apex, with acumen I.0-4.0 mm
long, the lamina conspicuously reticulate above. Primary veins I6-21. Petioles
canaliculate when young. Inflorescences terminal and axillary, rather lax panicles
with light brown pubescence.

8. Parinari brasiliensis (Schott) Hooker f., Mart. Fl. Bras. 14(2): 5I. 1867.
PetrocaryabrasiliensisSchott in Spreng., Linn. Syst. Veg., ed. I6. 4(2), (Cur. Post.): 405. I827.
Ferolia brasiliensis(Schott) Kuntze, Rev. Gen. 216. 1891.
Parinariumpohlii Hooker f., Mart. Fl. Bras. 14(2): 51, t. 17. 1867. Type. Pohl sn, Brasil, Minas
Gerais, fl (BR, M).
Feroliapohlii (Hooker f.) Kuntze, Rev. Gen. 2 6. I891.

Tree, the young branches villous. Leaves oblong, 7.5-I6.0 cm long, 2.5-6.5 cm
broad, rounded to subcuneate at base, acuminate at apex, with acumen 2.0- 1.0 mm
long, glabrous above, tomentose and with stomatal cavities beneath; midrib promi-
nulous above, pubescent towards base; primary veins 23-26 pairs, prominent be-
neath; petioles 5.0-7.0 mm long, pubescent, terete, with two medial glands. Stipules
to 2.0 mm long, caducous. Inflorescences short terminal densely flowered panicles
6.o cm long, the rachis and branches densely brown.tomentellous. Receptacle sub-
campanulate-turbinate, densely pubescent on exterior; pedicels I.0-4.0 mm long.
Petals 5, white, equalling the calyx lobes. Stamens 7 fertile, unilateral, with 7 short
filamentous staminodes opposite to them. Ovary and base of style pilose. Fruit un-
known.
190 Flora Neotropica

TPYE. Schott 4222, Brazil, Guanabara or Rio de Janeiro, fl (isotypes, K, NY).

DISTRIBUTION. Known only by the type collections from Minas Gerais and from
near Rio de Janeiro, Brazil.
Comparative note. Leaves oblong 7.5-I6.0 cm long, 2.5-6.5 cm broad, rounded
to subcuneate at base, acuminate at apex, with acumen 2.0- I.0 mm long. Primary
veins 23-26. Petioles terete. Inflorescences short densely flowered panicles ca 6.o cm
long, with dense brown pubescence.

9. Parinari klugii Prance, sp nov Fig 29 M-P.

Arbor ramulisjuvenilibus sparsepubescentibus. Folia alternata petiolata, petiolo

5.o-6.o mm longo, glanduloso, tomentoso; laminae oblongae, 6.5- 1.5 cm longae,
2.3-4.5 cm latae, basi rotundatae vel subcuneatae, apice in acuminem 8.0-12.0 mm
longum contractae, superne juveniliter sparse lanato-tomentosae maturitate glabres-
centes indumento mox deciduo, subtus cum cavis stomatalis, griseo-tomentosae;
nervis secundariis 19-22 jugis, subtus prominentibus. Stipulae ad 3.0 mm longae,
ovatae, caducae. Flores in paniculis terminalibus corymbosis ubique griseo-brunneo-
tomentosisdispositi,rachi ramisquesparsehirsutis.Bracteaebracteolaequeovatae, acu-
tae, calicem aequantes. Receptaculum subcampanulatum, extus griseo-tomentosum,
intus pilis longis ubique dense tomentosum. Calycis-lobi acuti. Petala 5, alba.
Stamina 7, unilateralia; staminodiis opposita. Ovarium villosum. Stylus e basi
ovarii ortus base villosus. Fructus mihi ignoti.
TYPE. Klug3870, Peru, San Martin, Juan Jui, Alto Rio Huallanga, fl (holotype,
NY; isotypes, BM, F, K, MO, US).
This specimen was included under the species P. parilis by Macbride (Fl. Peru
1074. 1938), but differs from P. parilis in many features, particularly in the shorter
leaves with fewer primary veins (28-32 in P. parilis), the shorter petioles, in the
corymbose inflorescence,and the pubescence of the inflorescenceand young branches.
Parinariklugiidiffersfrom P. excelsaand related species in the corymbose inflorescence,
the hirsute hairs on the petioles, young branches and inflorescence, and the striate
exterior of the receptacle tube.

Io. Parinari maguirei Prance, sp nov Fig 29 H-L.

Arbor parva ad I3.0 m alta. Folia alternata, pttiolata, petiolo 3.0-4.0 mm longo;
laminae orbiculares vel ellipticae, 2.5-5.0 cm longae, 1.5-3.5 cm latae, basi sub-
cuneatae vel rotundatae, apice rotundatae vel breviter apiculatae, superne glabrae,
subtus cum cavis stomatalis,breviter lanatis; costa supra impressa,subtusprominente;
nervis secundariis 9-11 jugis, subtus prominentibus. Stipulae ad i.o mm longae,
caducae. Flores mihi ignoti. Fructus drupaceus, ellipsoideus, 4.0-5.0 cm longus,
3.0-3.5 cm latus, extus dense verrucosus; mesocarpio carnoso; endocarpio duro,
crasso, obturamentibus duobus basalibus instructo, extus fibroso, intus dense lanato-
tomentoso.
TYPE. Maguire & Fanshawe 23390, Guyana, Kaieteur savannas, fr (holotype,
NY; isotypes, F, M, US).
Savannas of the Kaieteur Plateau.
DISTRIBUTION.
GUYANA. Kaieteur Plateau, Cowan& Soderstrom
2180 fr (NY, US).
Parinari 191

The leaf shape of this species is most distinctive. Since the majority of the species
of Parinari are distinguished principally by vegetative characters, it is possible to
describe this distinctive species from specimens lacking flowers. Parinarimaguireiis
easily recognized by the small broad orbicular leaves with a rounded or apiculate
apex. The outer surface of the endocarp is much smoother than in most species of
Parinari.It is granular and fibrous as is typical, but it is not channelled on the surface.

I. Parinari littoralis Prance, sp nov Fig 30.

Arbor media, ramulis juvenilibus glabris lenticellatis. Folia alternata breviter

petiolata, petiolo 3.o-4.0 mm longo, sparse puberulo mox glabro, eglanduloso;
laminae ellipticae, coriaceae, 6.4-7.5 cm longae, 3.0-4.0 cm latae, basi rotundatae,
apice acutae vel rotundatae vel breviter acuminatae, superne glabrae, subtus cum
cavis stomatalis faucis tomentosis munitae; costa media supra impressa; costis secun-
dariis 15-20 jugis, supra planis, subtus prominentibus. Stipulae parvae, ovatae,
persistentes. Flores in paniculis axillaribus compactis dense-florentibus dispositi,
rachi ramisque tomentosis. Bracteae bracteolaeque ovatae, calicem aequantes,
caducae. Receptaculum subcampanulatum, extus brunneo-tomentosum, intus ad
faucem pilosum, versus basim dense tomentosum. Petala 5. Stamina 7, unilateralia,
staminodiis opposita. Ovarium pilosum. Stylus e basi ovarii ortus, basi pilosus.
Fructus mihi ignoti.
TYPE. Beldm1833, Brazil, Bahia, Marau, fl (holotype, IAN; isotypes, NY, UB).
DISTRIBUTION. Known only from the type collection from coastal forest in Bahia,
flowering in October.
This species is quite distinct by the densely clustered axillary inflorescence. It
differs from the other species of eastern Brazil in the blunt leaf apex and the im-
pressed midrib.

12. Parinari parvifolia Sandwith, Kew Bull. 1931: 374. I931.

Large tree to 45.0 m high, the young branches villous, soon becoming glabrous
and lenticellate. Leaves elliptic to oblong-lanceolate, 2.0-7.5 cm long, I.0-3.0 cm
broad, stiffly coriaceous, cuneate (rarely slightly rounded) at base, acuminate at apex,
with acumen Io.o-I3.o mm long; primary veins 27-30 pairs, prominent beneath,
slightly impressed above; midrib slightly impressed above; petioles short, 2.0-5.0 mm
long, pubescent when young, with 2 inconspicuous glands near lamina, terete.
Stipules to I.0 mm long, caducous. Inflorescences terminal and axillary panicles, the
rachis and branches light-brown-tomentellous, 3.0-5.0 cm long, densely flowered.
Receptacle subcampanulate-turbinate, tomentose on exterior; pedicels I.0-2.0 mm
long. Petals 5, equalling calyx lobes. Stamens 7 fertile, with 7 short filamentous
staminodes opposite them. Ovary and base of style villous. Fruit unknown.
TYPES. Sandwith I39, Guyana, fl (holotype, K; isotype, NY); Jenman 3992,
Guyana, fl (paratypes, K, NY, US).
Primary forests of Guyana and eastern Amazonia.
DISTRIBUTION.
GUYANA. Jenman 20o fl (US). BRAZIL. Pari: Pires 6800 fl (IAN, SP).

LOCAL NAME. Hill Buhurada.

192 Flora Neotropica

"-'~
"(S,y'..
.,'i' ,

16

G H1

FIG 30. Species of Parinariand Couepia.A-E, Parinarilittoralis (Belem 1833); A, habit, x 0.5; B,
flower, x 5; C, flower section, x 6; D, ovary, x 5; E, petal, x i o. F-K, Couepiabelemii(Belim et al
3146); F, petal, x 2; G, ovary, X 4; H, flower, x i; J, flower section, x 2; K, habit, x 0.5.
Parinari 193

13. Parinari cardiophylla Ducke, Arch. Inst. Biol. Veg. Rio de Janeiro a:
33. I935.

Tree to 25.0 m high, the young branches puberulous, soon becoming glabrous
and lenticellate. Leaves ovate, 8.o-I2.0 cm long, 4.0-7.0 cm broad, cordate at base,
acuminate at apex, with acumen ca I2.0 mm long, glabrous above, tomentose and
with stomatal cavities beneath; primary veins 15-20 pairs, prominent beneath;
midrib prominulous above, petioles 6.0-i2.0 mm long, glabrous when mature,
terete, with two medial sessile glands. Stipules caducous. Inflorescences terminal and
axillary panicles 5.0-6.0 cm long, densely flowered, the rachis and branches with
short light-brown pubescence. Receptacle subcampanulate-turbinate, tomentellous
on exterior; pedicels I.o-3.0 mm long. Petals 5, white, just exceeding the calyx lobes.
Stamens 7 fertile, with 7 filamentous staminodes opposite them. Ovary and lower
portion of style pli ose. Fruit unknown.
TYPES.DuckeRB 24182, Brazil, Amazonas, fl (holotype, RB; isotypes, INPA, K,
NY, P); RB 24183 fl (paratypes, K, NY, US).
DISTRIBUTION. Primary forest in Brazilian Amazonia.
BRAZIL. Amazonas: Frdes & Addison29070 fl (IAN, UB). Par,: Oliveira4575 st (IAN, NY).

This species is as yet poorly known. It resembles P. campestrisin the leaf shape,
especially the cordate base, but differs by lacking the persistent large stipules of the
latter species. Further material is needed to clarify the status of this species, especially
its relationship to P. campestrisand P. sprucei.

I4. Parinari parilis Macbride, Candollea 5: 367. 1934; Fl. Peru 1074. I938.

Tree, the young branches glabrous, lenticellate. Leaves oblong-lanceolate,

8.0- 7.0 cm long, 2.5-5.5 cm broad, cuneate to subcuneate at base, acuminate at
apex, with acumen 6.0-I I.o mm long, glabrous above, with lanate pubescence and
stomatal cavities beneath; midrib prominulous above, glabrous; primary veins 28-32
pairs, prominent beneath; petioles 7.0-9.0 mm long, pubescent when young, be-
coming glabrous with age, without conspicuous glands, terete. Stipules to 2.0 mm
long, caducous. Inflorescences lax terminal panicles to 8.o cm long, the rachis and
branches with a silver gray pubescence. Receptacle subcampanulate-turbinate, gray-
tomentellous on exterior; pedicels I.0-2.0 mm long. Petals 5, white, equalling the
calyx lobes. Stamens 6-7 fertile, unilateral, with 7-8 filamentous staminodes opposite
them. Ovary and base of style pilose. Fruit unknown.
TYPES.LI. Williams1140, Peru, Loreto, fl (holotype, F); III3 fl (paratypes, F,
US).
DISTRIBUTION. Known only from the type gatherings, collected in lowland forest
of Amazonian Peru.
LOCALNAME.Upacha-umari.
This species may be distinguished by the large oblong-lanceolate leaves with
numerous primary veins.

15. Parinari chocoensis Prance, sp nov Fig 26 F-N.

Arbor ca 30.0 m alta, ramulis juvenilibus pubescentibus mox glabris lenticel-
latis. Folia alternata petiolata, petiolo 5.o-8.o mm longo, glanduloso, canaliculato;
194 Flora Neotropica
laminae oblongae, 8.0-15.5 cm longae, 2.5-4.7 cm latae, basi subcuneatae, apice in
acuminem 7.0-I3.0 mm longum contractae, superne glabrae, subtus cum cavis
stomatalis, tomentosae; costa media supra versus basim leviter canaliculata, versus
apicem piano vel leviter prominente; costis secundariis 22-26 jugis, subtus prominen-
tibus. Stipulae ca 2.0 mm longae, tomentosae, caducae. Flores in paniculis terminali-
bus subterminalibusquedispositi, rachi ramisque tomentellis. Bracteae bracteolaeque
2.5-4.0 mm longae, ovatae, caducae, calicem aequantae. Receptaculum subcam-
panulatum, extus griseo-tomentosum, intus pilis longis ubique dense viloso-tomento-
sum. Calycis-lobi acuti. Petala 5. Stamina 7, unilateralia; staminodiis opposita.
Ovarium villosum. Stylus e basi ovarii ortus, basi villosus. Drupa elongato-ellipsoidea,
ad 7.0 cm longa; epicarpio dense lenticellato, glabro; mesocarpio tenue, carnoso;
endocarpio duro, fibroso, 5.0-7.0 mm crasso, intus dense lanato-tomentoso.
TYPE. Fuchs & Zanella 21793, Colombia, Choc6, Rio Baud6, I8.5 km from
mouth, fl fr (holotype, NY; isotypes, COL, SRR7, US).
Known only from the type gathering from a high river bank.
DISTRIBUTION.
Collected in flower and fruit in February.
Parinarichocoensis is apparently different from any other species of Parinariby the
narrowly ellipsoid fruit (the fruit of several other species is still unknown). It is most
closely related to P. parilis but differs by the slightly impressed lower portion of the
midrib, the fewer primary veins, the dull upper surface of the lamina, and the shorter
inflorescences. It is also closely related to P. klugii,P. pachyphyllaand P. brasiliensisbut
differs from all three in the laxer inflorescence, the slightly impressed midrib, and
the narrowly ellipsoid fruit. It differs from each of these 3 species individually by
many additional characters.

i6. Parinari obtusifolia Hooker f., Mart. Fl. Bras. 14(2): 52, t.i8. I867;
Usteri, Fl. Sao Paulo 184. I91I.
Feroliaobtusifolia(Hooker f.) Kuntze, Rev. Gen. 216. I891.

Suffrutex or small shrub 0.2-3.0 m tall, the young branches tomentose, be-
coming glabrous and lenticellate with age. Leaves oblong-elliptic, 5.o-Io.o cm long,
2.5-4.0 cm broad, rounded to slightly cordate at base, obtuse to acute at apex,
glabrous or with short-ferrugineouspubescence, especially near nerves above; with
dense silvery white lanate appressed pubescence completely obscuring the stomatal
cavities beneath; midrib prominent above; primary veins 11-16 pairs, prominent
beneath, often ending in a small gland at margin; petioles 2.0-3.0 mm long, terete,
densely tomentose, with two medial sessile glands. Inflorescencesaxillary or terminal
panicles, with ferrugineousto brown pubescence. Stipules to 3.0 mm long, caducous.
Receptacle subcampanulate-turbinate,brown-tomentose on exterior. Petals 5, white,
equalling the calyx lobes. Stamens 7-8 fertile, with 7-8 staminodes opposite them,
some reduced to mere teeth. Ovary and base of style villous. Fruit oblong, 2.5-4.0 cm
long, 2.0 cm broad; epicarp verrucose; mesocarp fleshy; endocarp hard, thick, with
a rough channelled exterior, densely tomentose within.
TYPE.Gardner3I37, Brazil, Goias, fl (holotype, K; isotypes, BM, CGE, F, NY,
OXF, P).
Cerradoand open disturbed areas of the planalto of Central Brazil.
DISTRIBUTION.
BRAZIL. Mato Grosso: Malme I992a fl (R); sn fl (LD); Riedel459 fr (LE); Robert388 st (BM);
530 fl (BM, K). Goias: Duarte 8319 fr (RB 12358I); Glaziou 21112 fl (BR, K, LE, P, R); Irwin &
7340 fl (NY, UB); 7444 fl (NY, UB); Irwin et al 10813 fl (NY, UB); 13608 fr (NY); 14469 fr
Soderstrom
(NY); Macedo3466 fl (K, MO, NY, SP); Prance & Silva 59553 fl (MO, NY, US); Riedel2319 fl (NY,
7 Shell Research, Rijkswijk, Netherlands.
Parinari 195

US); St. Hilaire 869 fr (P). Dist. Federal: Belim 1926 fl (NY, UB); Gomes1107 fr (RB I07091); Heringer
8671/865 fl (UB); Irwin & Soderstrom 5058 fr (NY, UB); Irwin et al 7847 fl (NY); 8229 fl (NY); 9197 fr
(NY); Io580 fl (NY, UB); iiioi fr (NY); 11587 fr (NY); 11649 fr (NY); I3880 fr (NY); Magalhdes
9670 fl (RB 102880); Pabst 4958 fl (NY, RB); Pereira & Pabst 4652 fl (HB); 7376 fl (F, K); Prance &
Silva 58996 fl fr (F, NY, US); 59705 fl (MO, NY, UB, US); Pires, Silva & Souza 9058 fl (UB); 9059 fr
(UB); Santos & Sacco 27085 fl (NY); Sucre 852 fl (NY, RB, UB). Bahia: Irwin et al 14700 fr (NY).
Minas Gerais: Duarte 823 fl (HB); Heringer8709/903 fl (NY, RB, SP, UB); Macedo4261 fl (K, NY);
Magalhdes9854 fl (IAN); 19005 fl fr (NY). Sao Paulo: Brade 7462 fl (R, SP); HandroSP 43305 fi
(FHO, SP, US); Hoehne & GehrtSP 11978 fl (SP); Lifgren 1451 fl (SP); Toledo & GehrtSP 43I74 fl
(FHO, SP); Usteri3576 fi (K).

LOCALNAME.Fruta de ema.
This species is easily recognized by its suffruticose habit as well as by the lower
surface of the leaves which are densely white-lanate. Comparable to this species of
suffruticose habit in central Brazil is P. capensisHarv. of the sandy savanna regions of
southern and eastern Africa. P. capensis, however, differs from P. obtusifolia in many
characters such as the foliar shape and size and the pubescence of the leaf under-
surface and inflorescence.

Excluded Species of Parinari (New World only)

I. P. barbatumDucke, Arch. Jard. Bot. Rio de Janeiro 3: 45. 1922. =Exellodendron
barbatum(Ducke) Prance.
2. P. canescensGleason, Bull. Torrey Club 58: 370. I93I. =Couepia cognata (Steudel)
Fritsch.
3. P. cordatumHooker f., Mart. Fl. Bras. 14(2): 50. 1867. =Exellodendron cordatum
(Hooker f.) Prance.
4. P. coriaceumBentham, Jour. Bot. Hooker 2: 213. I840. =Exellodendron coriaceum
(Bentham) Prance.
5. P. gardneri Hooker f., Mart. Fl. Bras. 14(2): 50. 1867. =Exellodendron gardneri
(Hooker f.) Prance.
6. P. gracilis Kuhlmann, Ann. Prim. Reun. Sul-Am. Bot. 3: 78. I940. = Exellodendron
gracile (Kuhlmann) Prance.
7. P. guyanense Fritsch, Ann. Naturh. Mus. Wien 5: 14. I890. =Hirtella guyanensis
(Fritsch) Sandwith.
8. P. guyanensis (Aublet) Lemee, Fl. Guyane Franc. 2: 25. I952. =Brosimum (Mora-
ceae).
9. P. hostmannii Fritsch, Ann. Naturh. Mus. Wien 5: 13. 1890. =Couepia cognata
(Steudel) Fritsch.
io. P. krukovii Gleason, Bull. Torrey Club 60: 353. I933. =Couepia glandulosa
Miquel.
ii. P. pilosa Standley, Publ. Field Mus. Bot. 17: 259. I937. =Couepia canomensis
(Martius) Bentham ex Hooker f.

4. Exellodendron Prance, gen nov

Parinariumsensu Hooker f., Mart. Fl. Bras. 14(2): 49. I867 pro parte quoad P. cordatum,
P.
coriaceum,P. gardneritantum.
Parinariumsensu auct. viz P. barbataet P. gracile.
Parinari subgenus PellegriniellaHauman, Bull. Jard. Bot. Brux. 21: I89. 195I, pro parte, quoad
P. coriacea,et P. gardneritantum.

Ab Parinari sens. str. differt foliis subtus planis sine cavis stomatalis, petiolis
eglandulosis, bracteis bracteolisque parvioribus, drupa cum exocarpio laeve haud
196 Flora Neotropica

lenticellato, endocarpio laeve tenue breviter apiculato sine obturamentibus basalibus.

Trees or shrubs with hermaphrodite flowers 6.0-7.0 mm long. Leaves entire,
glabrous above, glabrous or densely lanate beneath, without stomatal cavities.
Petioles eglandular. Bracts and bracteoles eglandular, not enclosing the young
flowers in groups. Inflorescences terminal panicles. Receptacle subcampanulate-
turbinate, slightly swollen at one side, hollow, hairy throughout within, tomentose on
exterior. Calyx lobes acute. Stamens 7, not exceeding the calyx lobes, unilateral, with
small staminodes opposite them. Ovary inserted laterally at the mouth of the recept-
acle, the carpels bilocular. Style filiform, not exceeding the calyx lobes. Fruit a fleshy
drupe; epicarp smooth, not lenticellate; endocarp smooth, thin and hard, with an
appressed beak to one side of the base and a single line of dehiscence.
TYPE SPECIES. Exellodendron coriaceum (Bentham) Prance. The genus Exellodendron
is named for Dr. A. W. Exell, formerly Deputy keeper of the Herbarium of the
British Museum.
Venezuela, Guianas, Amazonian and Central Brazil; see Fig 32.
DISTRIBUTION.
In Exellodendronall species have very similar flower and fruit structureand cannot
be distinguished by characters of these structures. The species are, however, clearly
distinct by vegetative characters. The reasons for separation of Exellodendron from
Parinarihave been fully discussed elsewhere (Prance, in press) on a worldwide basis.

Key to the Species of Exellodendron

I. Leaf base distinctlycordateto subcordate;lamina3.0-5.5 cm long. 2. E. cordatum.
i. Leaf base rounded to cuneate; lamina 3.0-11.5 cm long.
2. Leafundersurface andreceptaclepersistentlydensewhite-arachnoid,
laminaovate-elliptic;
inflorescencesdenselyclustered. I. E. coriaceum.
2. Leaf undersurfaceglabrousor with a sparseeasily removedindumentum;lamina oblong;
inflorescencesratherlax.
3. Lamina 8.5-I 1.0 cm long, with a distinct well developed acumen 1.5-2.5 cm long.
3. E. barbatum.
3. Lamina to 7.0 cm long, the acumen less than 8.0 mm long.
4. Leaves coriaceous, with a distinct but shorter acumen 2.0-3.0 mm long, primary veins
prominent on lower surface. 4. E. gardneri.
4. Leaves chartaceous, long-apiculate (ca I.o cm long), primary veins inconspicuous.
5. E. gracile.

Taxonomic History of Exellodendron

Segregation of this group from Parinari has been fully explained elsewhere
(Prance et al 1969, Prance, in press), where it was shown that Parinarisensulatowas an
unnatural aggregation of all species of Chrysobalanaceae with a bilocular ovary. In
order to produce meaningful generic limits within the family, Parinariwas subdivided
into seven genera, one of which is Exellodendron.
When Ducke (1922) described Parinaribarbata,he observed that it differed in
many features from all other Parinariknown to him. Hauman (195 1) created the new
subgenus Pellegriniella,of Parinari,which appeared to have been based on the African
species P. tessmannii.At the end of the description he said, "Nous classons ici 2
especes americaines P. coriaceaBenth. et P. gardneriHook. f. et une espece africaine
P. tessmannii."The two American species are very different from P. tessmannii. Indeed,
they have since been transferredto Magnistipula(Prance I966b).
Exellodendron 197

I. Exellodendron coriaceum (Bentham) Prance, comb nov

ParinariumcoriaceumBentham, Jour. Bot. Hooker 2: 213. 1840; Hook. f., Mart. Fl. Bras. 14(2):
51. 1867.
Ferolia coriacea(Bentham) Kuntze, Rev. Gen. 216. I891.

Tree to Io.o m tall. Leaves oblong-elliptic, 9.0-12.0 cm long, 3.0-7.5 cm broad,

cuneate to subcuneate at base, acuminate at apex, with acumen I.0-1.5 cm long,
glabrous above, densely arachnoid beneath; midrib prominent on both surfaces;
primary veins 13-19 pairs, not prominent; petioles 5.o-6.o mm long, canaliculate
above. Stipules to 2.0 mm long, caducous. Inflorescences dense flowered axillary or
terminal panicles, much shorter than the leaves, with silver-gray pubescence. Bracts
and bracteoles I.0-3.0 mm long, caducous. Receptacle subcampanulate, swollen
slightly to one side, silver-gray-tomentoseon both interior and exterior. Calyx lobes
sharply pointed. Petals 5, white, caducous. Stamens 7, the staminodes represented
solely by swellings in the staminal circle or occasionally with I or 2 short filaments.
Ovary and style densely villous. Fruit round to oval, basally narrowed, 1.5-2.5 cm
long, 1.2-2.0 cm broad; epicarp smooth and glabrous; mesocarp thin and fleshy;
endocarp thin, hard, with a smooth surface, the interior sparsely lined with short
straight silvery hairs.
TYPE.Schomburgk 65, Guyana, fl (holotype, K; isotypes, BM, BR, CGE, GH, L,
NY, OXF)
DISTRIBUTION. Savannas, savanna margins and less frequently in periodically
flooded forest in Venezuela, Guyana and Amazonian Brazil.
VENEZUELA. Bolivar: Steyermark 86843 fl (VEN). Terr. Amazonas: Maguire,Cowan& Wurdack
31019 fl (F, NY). GUYANA. Schomburgk 93 fl (BM). BRAZIL. Amazonas: Baldwin 3463 fr (US);
Coelho14 (INPA 6736) fl (IAN, MG, NY); Ducke 1364 fl (A, F, IAN, MG 17777, R, US); 1700 fr
(A, F, IAN, MG 17778, NY, R, US); MG 10242 fr (BM, MG); RB 35r85 fl fr (NY, RB, US); Frdes
22630 fl (IAN, NY, SP, VEN); 25313 fl (IAN, NY, RB); 29594 fr (IAN); INPA 3247 fr (IAN, INPA,
MG, NY); 3710 fr (INPA, NY); Mello INPA 808 fl fr (IAN, INPA, NY); Riedelsn fl fr (OXF); 1406 fl
fr (A, US); Rodrigues& Coelho2049 fr (INPA 8427); 5780 fr (INPA I449, NY); Spruce1z8z fl (BM,
F, GOET, K, LD, LE, P). Terr. Roraima: Froes 22991 fl (IAN, NY); 23215 fl (NY). Para: Black
47-1758 fl (IAN); DuckeRB 10249 fr (MG); Frdes33563 fr (IAN); KuhlmannRB 21027 fr (US); Pires
& Silva 4401 fl (IAN, NY).

2. Exellodendron cordatum (Hooker f.) Prance, comb nov

Fig 31 A-C.
ParinariumcordatumHooker f., Mart. Fl. Bras. 14(2): 50. 1867.
Feroliacordata(Hooker f.) Kuntze, Rev. Gen. 2 6. 1891.

Small tree or shrub. Leaves ovate, 3.5-9.0 cm long, 2.25-3.50 cm broad, usually
distinctly cordate, rarely rounded at base, rounded at apex, or with a small acumen
to 3.0 cm long, glabrous above, glabrous or with densely lanate easily rubbed off
pubescence on lower surface; midrib prominent on both surfaces; primary veins
9-12 pairs, not prominent; petioles 4.0-8.0 mm long, canaliculate above, pubescent,
with 2 glands at lamina base. Stipules to 1.5 mm long, caducous. Inflorescences
terminal and axillary panicles. Bracts and bracteoles 1.0-2.0 mm long, caducous.
Receptacle subcampanulate, swollen slightly at one side, silvery gray-tomentose on
exterior, villous to base within. Calyx lobes acute. Petals 5, white. Stamens 7, with 7
filamentous staminodes opposite them. Ovary and base of style villous. Immature
fruit oblong; epicarp smooth, glabrous; mesocarp thin; endocarp thin, hard, with a
smooth surface, densely brown-pubescent within.
TYPE. Gardner2560, Brazil, Piaui, fr (holotype, K; isotypes, BM, CGE, US, W).
198 Flora Neotropica

^y E F|. cLokm4

3 1 Species of Exellodendron.
3I. A-C, E. cordatum(Irwinet al 14483); A, habit, x 0.5; B, flower,
x 3; C, flower section, x 3. D-G, E. barbatum(Irwin et al 55766, Pranceet al iz93); D, habit, x o.5; E,
young fruit, x o.5; F, flower, x 3; G, flower section, 3.
Exellodendron 199

DISTRIBUTION. Cerradosof the Planalto of central Brazil.

BRAZIL. Maranhao: Ducke MG 727 fl fr (MG, RB 15031). GoiSs: Irwin et al 14483 fl (NY).
Bahia: Irwin et al 14694 fl (UB); 148I3 fl (UB).

3. Exellodendron barbatum (Ducke) Prance, comb nov

Fig. 31 D-G.
ParinariumbarbatumDucke, Arch. Jard. Bot. Rio de Janeiro 3: 45. 1922.

Tree to 20.0 m tall. Leaves oblong, 8.o- I.o cm long, 2.5-3.5 cm broad,
cuneate at base, acuminate at apex, with acumen I.0-2.5 cm long, glabrous on both
surfaces; midrib prominent on both surfaces; primary veins 0o-I5 pairs, not promi-
nent; petioles 4.0-5.0 mm long, canaliculate above, usually covered by silvery gray
hairs. Stipules to 1.5 mm long, caducous. Inflorescences rather lax terminal or
axillary panicles, with short gray pubescehce. Bracts and bracteoles I.0-3.0 mm long,
membraneous. Receptacle subcampanulate, swollen slightly to one side, silvery gray-
tomentose on interior and exterior. Calyx lobes sharply pointed. Petals 5, white,
caducous. Stamens 7, with 7-8 filamentous staminodes opposite them. Ovary and
base of style villous. Fruit oblong, 2.0-2.5 cm long, I.O-I.5 cm broad; epicarp smooth
and glabrous; mesocarp thin and fleshy; endocarp thin, hard, with a smooth surface,
sparsely hirsutulous within.
TYPE.DuckeRB I6385, Brazil, Terr. Amapa, fl (holotype, RB; isotypes, F, NY,
P, R).
DISTRIBUTION. Non-flooded forest or periodically flooded forest in eastern
Venezuela, the Guianas and eastern Amazonian Brazil.
VENEZUELA. Bolivar: Steyermark86828 fr (VEN); 86864 fr (VEN). GUYANA. A. C. Smith
2609 fl (A, K, NY, P, US); 3559 fr (F, K, LE, NY, S). SURINAME. Lindeman6795 fr (NY, U);
Maguire, Schulz, Soderstrom& Holmgren54267 fr (F, NY); Irwin, Prance,Soderstrom& Holmgren55409 fl
(F, NY); 55766fl (F, NY); 55976fl (F, NY); Schulz & Elburgzoio6fr (BBS, U). FRENCH GUIANA.
Melinon 15 fr (P); i6 fr (P); 466 fr (P). BRAZIL. Amazonas: Pranceet al Io268 fr (INPA, NY). Terr.
Roraima: Pranceet al 4204 fl (NY); 4355 fr (NY). Pari: Capucho384 fl (F, IAN, RB); DuckeRB 19760 fl
(NY, P, RB); 18224 fl (K, RB); Krukoff1137 fl (A, BM, K, MICH, NY, P); Monteiroda Costa 1o9 fl
(IAN); Pires & Silva o0703 fl (IAN, NY); Prance & Pennington1931 fr (F, NY); Prance& Silva 58727 fl
(F, NY).

LOCAL NAMES.Venezuela: Guaray negro. Suriname: Ingibarki, Sirikoko. Brazil:

Cariperana.

4. Exellodendron gardneri (Hooker f.) Prance, comb nov

ParinariumgardneriHooker f., Mart. Fl. Bras. I4(2): 50, t. 17. 1867.
Feroliagardneri(Hooker f.) Kuntze, Rev. Gen. 216. I891.

Shrub to I.5 m tall. Leaves oblong-ovate, 3.5-6.5 cm long, 2.0-3.0 cm broad,

cuneate to rounded at base, short-acuminate at apex, with acumen 3.0-5.0 mm long,
glabrous above, glabrous or with sparse caducous pubescence beneath; midrib
prominent on both surfaces; primary veins 6-8 pairs, prominulous on both surfaces;
petioles 2.0-3.0 mm long, shallowly canaliculate, silvery gray-lanate. Stipules to
I.o mm long, caducous. Inflorescences terminal panicles, the rachis and branches
gray-puberulous.Bracts and bracteoles 0.5-2.0 mm long. Receptacle subcampanulate
swollen slightly at one side, silvery gray-puberulous to glabrescent on exterior
200 Flora Neotropica

J--

"" ':
; -., V- * , 1 ;
} , ~'? "?4A / -II'"

> *****'' I_}-''" *- s "^2

FIG 32. a E. coriaceum,0 E. barbatum, E. gardneri,

Geographic distribution of Exellodendron.
E. cordatum,
+ E. gracile.

Calyx lobes 5, pointed. Petals 5. Stamens 7, with short filamentous staminodes

opposite them. Ovary and base of style pilose. Fruit unknown.
TYPE.GardnerI266, Brazil, Goias, fl (holotype, K; isotype, OXF).
DISTRIBUTION. Cerradosof the Planalto of Central Brazil.
BRAZIL. GoiSs: Gardner3139 fl (BM, P); Irwin et al I4591 fr (NY). Minas Gerais: Glaziou
14677 fl (BR, F, IAN, K, LE, NY, P, US); 15943 fl (LE, P).

5. Exellodendron gracile (Kuhlmann) Prance, comb nov

Parinariumgracile Kuhlmann, Ann. Prim. Reun. Sul.-Am. Bot. 3:78. 1940.

Medium sized tree, branchlets gray-arachnoid. Leaves 3.o-6.o cm long, I.3-

2.2 cm broad, oblong-lanceolate, glabrous above, with a sparse easily rubbed off
lanate indumentum beneath, cuneate at base, narrowed to an acute apex, sometimes
acuminate; midrib slightly prominent on both surfaces; primary veins 6-8 pairs,
not prominent; petioles 3.o0-5.o0mm long, canaliculate above, tomentose. Inflores-
cences terminal panicles 2.0 cm long, the rachis and branches shortly pilose. Bracts
and bracteoles I.0-1.5 mm long, caducous. Receptacle gray-tomentellous on ex-
terior. Fruit ellipsoid, 3.o0cm long, 1.2 cm broad, basally narrowed to a stipe 5.0-
6.o mm long; epicarp smooth and glabrous; mesocarp thin and fibrous; endocarp
Parinari 201

hard and thin, with a smooth surface, the interior filled by a lanate mass of long
brown hairs.
TYPE. J. G. Kuhlmann279, Brazil, Espirito Santo, fr (holotype, RB 34160).
DISTRIBUTION. Known only from the type gathering from riverine forest of the
Rio Doce in Espirito Santo.
LOCALNAME.Oiti.

5. Maranthes Blume, Bijdr. Fl. Nederl. Ind. 89. 1825; Reichb., Consp. 204.
1829 "Maranthus";Kostermans, Candollea 20: io6. I965; Prance, Bol.
Soc. Brot. II. 40: I83. 1966; Brittonia 20: 203-204. 1968; in press.
Exitelia Blume, Fl. Jav., Praef. 7. 1828.
GrymaniaPresl, Epim. Bot. 193. 1851 pro parte quoad G. salicifolia tantum.
Parinarisubgen SarcostegiaBentham in Hooker, Niger Fl. 335. 1849.

Small to large trees with hermaphrodite flowers 9.0-40.0 mm long. Leaf under-
surface glabrous (or densely wooly-tomentose in some African species). Petioles
glandular or eglandular. Bracts and bracteoles eglandular, not enclosing the young
flowers in groups. Receptacle of varied shape but narrowed to the base, solid or nearly
solid (hollow and curved in one African species), glabrous inside at base; calyx lobes
rounded. Stamens I8-60, exserted far beyond calyx lobes, often forming a complete
circle, or slightly unilateral; filaments glabrous. Ovary of i (-2-3) carpels, inserted
laterally at the mouth of the receptacle, the carpels bilocular with one ovule in each
loculus. Fruit a fleshy drupe; epicarp smooth, glabrous, not verrucose; endocarp
very hard, fibrous with a roughish exterior, with two lateral plates which break away
during germination to allow seedling to escape. Germination epigeal, the first leaves
opposite.
TYPE SPECIES. Maranthescorymbosa Blume. The name Maranthesis probably
derived from the Greek word Maraino to wither or fade (opposite of Amaranthus
everlasting). This refers to the persistent mass of stamens and the calyx which remain
in a withered state, long after flowering.
DISTRIBUTION. Ten species in Africa, and one extremely widespread species in
Malesia, which also grows in Panama.
The Panamanian collections cited under M. corymbosa come from forest which
has not been recently disturbed, in three well separated localities in Panama. Since
the Isthmus of Panama has been subject to disturbance for several centuries as a land
route and then for a canal, the chances are good that areas where Marantheshas been
found were penetrated and disturbed in the past. As M. corymbosa grows naturally in
a wide range of habitats in Malesia, it is probable that it would readily naturalize in
Panama. On the other hand, if it were native to the New World, a species with such
wide ecological amplitude would probably be more widespread today than it is.
Therefore, it seems reasonable to conclude that Maranthesis only naturalized in the
New World.
Taxonomic History of Maranthes
This genus was first described by Blume (1825), and has usually been included
in Parinarias a separate subgenus, Sarcostegia.Blume himself later (i826) included it
in Parinari.Marantheswas merged with Parinarion the basis of the false septum of the
ovary. The species of Parinarisubgen Sarcostegiaare very different from Parinarisensu
stricto, and full reasons for reinstating Maranthesas a genus are given elsewhere
(Prance et al 1969, Prance; in press). Maranthesis in fact more closely related to
Couepiathan to Parinari.
202 Flora Neotropica

The American Species of Maranthes

i. Maranthes corymbosa Blume, Bijdr. Fl. Ned. Ind. 2:89. 1825; Prance,
Brittonia 20: 203-4. 1968; Prance, in press Fig. 33.
8 Exitelia Blume ex Spanoghe, Linnaea 15: I74. 1841.
corymbosa
Parinariumcorymbosum (Blume) Miquel, Fl. Ind. Bat. I(I): 356. 1855, Io84. 1858.
CouepiapanamensisStandley, Trop. Woods. 44: 22. 1935; Woodson & Schery, Ann. Missouri Bot.
Gard. 37: 178. I950. Type. G. P. Cooper279, Panama, fl (holotype, F; isotypes, GH, NY,
US).

Large tree to 25.0 m tall, the young branches puberulous or glabrous. Leaves
oblong to oblong-elliptic, 6.5-I3.0 cm long, 2.0-4.5 cm broad, subcuneate at base,
abruptly acuminate at apex, with acumen I .o-20.o mm long, glabrous on both
surfaces; primary veins 7-9 pairs, prominent beneath; petioles 4.0-6.0 mm long,
glabrous, terete, with two conspicuous glands near lamina base. Stipules to 3.0 mm
long, lanceolate, caducous. Inflorescences terminal corymbose panicles, the rachis
and branches almost glabrous, with a few appressed hairs only (often puberulous in
Asiatic material). Bracts and bracteoles minute, acute, caducous. Receptacle tur-
binate-campanulate, almost solid, 5.0-6.0 mm long, with a few appressed hairs on
exterior (to tomentellous in Malesia), glabrous within except for deflexed hairs at
throat; pedicels 4.0-7.0 mm long. Calyx lobes rounded. Petals 5, with ciliate margins.
Stamens ca 19, inserted in a semicircle. Ovary villous. Style pubescent at base only.
Fruit oblong-pyriform, to 6.o cm long, 2-celled; epicarp smooth, glabrous; mesocarp
thin, fleshy; endocarp ca 5.0 mm thick, hard, woody, densely lanate within.
TYPE.Blume 51r6,Java, fl (BO, L).
DISTRIBUTION. Forest on high ground in the Isthmus of Panama where it is
probably naturalized. Abundant in Malesia from the Malay Peninsula through the
Philippines, Indonesia, Tropical Australia, New Guinea and western Polynesia.
PANAMA: Dwyer I494 fr (MO, NY); Dwyer & Stimson8060 fr (MO, NY); SuttonHayes goo fl
(NY).

As the Panamanian material cited here is certainly conspecific with the Malesian
material, Blume's epithet must apply.

6. Couepia Aublet, P1. Guiane I: 519, t. 207. I775; DC., Prodr. 2: 526. 1825;
Benth., Jour. Bot. Hooker 2: 2I5. I840; Hooker f. in Benth. & Hook.,
Gen. P1. I: 608. I865; Hooker f., Mart. Fl. Bras. 14(2): 40-49. I867;
Focke in Engl. & Prantl, Nat. Pflanzenfam. 3(3): 59. I894; Macbride,
Fl. Peru 0I79. 1938; Kleinhoonte in Pulle, Fl. Suriname 2(I): 45I. I939;
Prance, in press.
Acia Schreber, Linn., Gen. P1. ed. 8. 2: 458. 1791, pro parte quoad syn. Couepiatantum.
PleraginaArruda da Camara ex Koster, Trav. 499. i816, pro parte quoad P. rufa tantum.
GrymaniaPresl, Epim. Bot. 193. I851, pro parte quoad G. polyandratantum.
Moquileaauct. non Aublet; Mart., Nov. Gen. et Sp. P1. 2: 80. 1827; Endl., Gen. I25I. I840;
Meisner, P1. Vasc. Gen. I: I02. 1836, pro parte excl. M. guianensiset Acioaguianensis.

Trees or shrubs with hermaphrodite flowers 7.0-40.0 mm long. Leaves entire,

usually glabrous above, glabrous or with dense lanate or arachnoid very rarely
hirsute indumentum beneath. Petioles with two sessile glands or eglandular, often
8 The numerous Malesian synonyms of this species are listed elsewhere, (Kostermans I965a,
Prance, in press). The description given here refers to the American element of this species.
Parinari 203

4 k

FIG 33. Maranthescorymbosa (Cooper279, Dwyer r494); A, habit, X 0.5; B, flower section, x 0.5; C,
ovary transverse section, x o.5; D, fruit, x o.5; E, fruit transverse section, x o.5.
204 Flora Neotropica

canaliculate. Bracts and bracteoles eglandular. Inflorescences axillary or terminal

panicles or racemes. Receptacle cylindrical to turbinate, but with much variation in
shape and size, rarely flattened, usually hollow, the base within commonly glabrous
(rarely hairy), with deflexed hairs at mouth, tomentose to glabrous on exterior.
Calyx lobes acute or rounded, often reflexed. Stamens (IO-)I4-IOO (to 300 in C.
macrophylla), most often in a complete circle, less frequently unilateral with staminodes
opposite them; filaments exceeding the calyx lobes, usually much entwined to form a
tangled mass. Ovary of I(-2-3) carpels, inserted laterally at mouth of receptacle;
carpels unilocular. Fruit a hard or fleshy drupe, with a verrucose, smooth or tomen-
tose epicarp; endocarp hard and roughish, with a characteristicallygranular exterior,
breaking up in an irregular manner on germination. Germination hypogeal, the
first leaves alternate.
TYPE SPECIES. Couepia guianensis Aublet. The name Couepia is derived from the
vernacular name in French Guiana.
DISTRIBUTION. Mexico and Central America, Colombia, Ecuador, through the
Guianas and Amazonia to southern Brazil.
Couepiacontains 8 groups of two or more closely related species which include
26 of the 55 species. The remaining 29 species are clearly distinct from any group of
species as well as from each other (ie, taxonomically isolated species in the sense of
White, 1962, p 75). Variation in the genus suggests no overall pattern on which to
base subgeneric taxa. The groups of related species do not represent any subgeneric
or supra-specifictaxa. This is confirmed by the fact that it is also impossible to group
the taxonomically isolated species into any meaningful subgeneric taxa, either to-
gether with or apart from the species groups.
The treatment of closely related species such as those in Couepiahas been sum-
marized by White (1962) in his study of speciation in Diospyros.White uses the
following categories: I. Superspecies with completely allopatric components.
2. Superspecies with imperfectly allopatric components, and 3. Sympatric species
groups. The genus Couepiacontains examples of all three categories of closely related
species. These terms will be used in reference to Couepiawithout further explanation
since they are clearly defined in White's paper, and have also been discussed by me
(Prance 1967). In this treatment of the genus the species groups are considered first,
and the species descriptions of each group are followed by a discussion of the taxo-
nomic problems within the group.
Taxonomic History of Couepia
The genus was described by Aublet (I775), and was maintained by Brown
(1818) and de Candolle (I825). Schreber (1789) reduced it to synonymy under
Acia which was his name for the genus of Acioaof Aublet. Martius (1827) described
Moquileachar.reformatus, which was a union of three of Aublet's genera, Acioa,Couepia,
and Moquilea.This circumscriptionof the genus was followed by Martius & Zuccarini
(1832), who described three new species of Moquileanow referred to Couepia(M.
grandiflora,M. paraensisand M. uiti) by Meisner (I837) and by Endlicher (I840).
Bentham (I840) realized that Moquileaand Couepiaof Aublet were separate
genera. He defined the differences between the two genera and was the first person
to divide Couepiaat the subgeneric level. He divided Couepiainto two sections.
I. Eucouepia:stamens more than 30, inserted in a complete circle.
2. Hemicouepia: stamens about 20, unilateral.
Bentham did not see material of Acioa,and consequently did not attempt to place it
in this work. However, Hooker (i865), in the GeneraPlantarum,cited Acioa as a
synonym of Couepia.In the same work the next genus was Griffonia,which represented
the African species of Acioa.
Couepia 205

Walpers (1843) did not follow Bentham, but included Couepiain his genus
Moquilea,which contained two sections.
I. Section Moquilea,containing M. grandiflora,M. uiti, M. kunthiana,M. paraensis
M. canomensis, and M. tomentosa.All except the last species are now considered as
Couepia.
2. Section Couepia,which contained the two subdivisions of Bentham.
I. Eucouepia,including M. comosaand M. bracteosa.
2. Hemicouepia,including M. multiflora.
Mueller (1857) also included Couepiaand Acioain his genus Moquileawhich he
divided into four subgenera, Eumoquilea,Microbalanus, Couepiaand Acioa. Each sub-
genus contained at least one species of Couepiaas defined in this work. In addition to
species of Couepiasubgen Eumoquileaalso contained species of Licania, and subgen
Acioaincluded Aublet's species of Acioa.
It was not until Fritsch (i 888) summarized the genera of the Chrysobalanaceae,
that Couepiawas purged of all extraneous entities. Fritsch recognized the three genera
of Aublet, Acioa,Couepia,and Moquilea.This circumscription was followed by Focke
(1891) in the Pflanzenfamilien, and has been adopted in all later treatments of the
family.
Sprengel (i827) transferredCouepiacanomensis to Hirtella.From time to time later
authors have misassignedvarious species of Couepiato Hirtella,but the two genera have
never been merged and they are clearly distinct. In recent work on the generic limits
of the Chrysobalanaceae (Prance et al 1969, Prance, in press), no change has been
made in the circumscription of Couepia.As I find no reliable characters to subdivide
Couepiainto subgenera or sections, these earlier infrageneric taxa are abandoned.

Key to Species of Couepia

i. Inflorescence a raceme or spike. (For contrasting statement see p. 206).
2. Leaves prominently reticulate beneath, with conspicuous parallel secondary veins.
3. Inflorescence with short sparse silver-gray pubescence; receptacle turbinate; bracteoles
caducous. 28. C. elata.
3. Inflorescence with dense ferrugineous pubescence; receptacle cylindrical to obconical;
bracteoles persistent.
4. Leaf undersurface with distinct stomatal cavities; fruit exterior verrucose.
o. C.foveolata.
4. Leaf undersurface reticulate, but without stomatal cavities; fruit exterior usually
pubescent.
5. Leaves with acumen 5.5-I8.0 mm long; receptacle long and slender, I 1.0-22.0
mm long. 7. C. parillo.
5. Leaves with acumen 2.5-12.0 mm long; receptacle short, 5.5-I0.0 mm long.
6. Inflorescence many-flowered; leaves ovate to oblong-elliptic, 8.5-18.0 cm long.
9. C. canomensis.
6. Inflorescence few-flowered; leaves elliptic, 4.0-6.o cm long. 8. C. steyermarkii.
2. Leaves not prominently reticulate beneath; fruit epicarp always glabrous, smooth or
verrucose.
7. Bracteoles persistent, and at least 3/4 the length of receptacle or more than I.0 mm
long.
8. Flowers not exceeding 4.0 mm long9. 14. C. spicata.
8. Flowers 6.o mm or more.
9. Flowers 16.0-25.0 cm long; stamens more than 50; leaves 14.0-25.0 cm long.
10. Primary leaf veins distinctly anastomosing at margins; flowers I6.o-i8.0 mm
long; leaves with rounded to subcordate bases; receptacle subcylindrical.
30. C. insignis.
8 All flower length measurements are from tip of calyx lobe to pedicel articulation.
206 Flora Neotropica

Io. Primary leaf veins not anastomosing; flowers 20.0-30.0 mm long; leaf base
cuneate to rounded; receptacle tubular.
I . Leaves elliptic, 6.0-I0.5 cm broad; primary veins I0-I2 pairs; receptacle
tube 20.0-25.0 mm long. 29. C. martinii.
I . Leaves oblong-lanceolate, 3.5-7.0 cm broad; primary veins 17-25 pairs;
receptacle tube I2.0-I5.0 mm long. 31. C. bondarii.
9. Flowers 6.0-I2.0 mm long; stamens 15-28; leaves 6.0-I8.o cm long.
12. Leaves Io.o-I8.o cm long, caudately acuminate; exterior of receptacle with
red-brown pubescence; stamens 15-19; style hairy at base only. 12. C. exfiexa.
I2. Leaves 6.0-i2.0 cm long, apex acuminate; exterior of receptacle with light
brown pubescence; stamens ca 25; style hairy for 3/4 of length. I3. C. habrantha.
7. Bracteoles caducous or less than half the length of receptacle, under I.0 cm long.
13. Receptacle hairy inside to the base; filaments hairy; leaves hirsute beneath.
33. C. recurva.
13. Receptacle glabrous inside except at throat; filaments glabrous; leaves with arach-
noid indumentum or glabrous beneath.
14. Stamens less than 26.
i5. Receptacle subcylindrical, with a dense ferrugineous-sericeous pubescence;
leaves I4.0-24.0 cm long x 5.5-8.5 cm broad. I . C. magnoliifolia.
I5. Receptacle cylindrical, with sparse appressed pubescence only; leaves
2.5-15.0 cm long x 1.5-5.0 cm broad.
i6. Leaves obovate, 2.5-I0.0 cm long, bluntly acuminate; bracteoles persis-
tent or subpersistent; stamens in complete circle. 34. C. obovata.
I6. Leaves oblong-lanceolate, 9.0-I5.0 cm long, with long acumen; bracteoles
caducous; stamens unilateral. 4. C. leptostachya.
14. Stamens numerous (more than 6o).
17. Receptacle glabrous or with sparse gray appressed tomentum on exterior.
35. C. williamsii.
17. Receptacle with dense brown pubescence on exterior.
i8. Leaves oblong-lanceolate. 23. C. krukovii.
18. Leaves oblong to oblong-elliptic.
19. Primary veins 24-30 pairs. 22. C. macrophylla.
19. Primary veins I6-20 pairs.
20. Leaf apex with prominent well developed acumen. 36. C. chrysocalyx.
20. Leaf apex rounded to bluntly acuminate. 24. C. latifolia.
I. Inflorescence a panicle.
21. Bracts and bracteoles persistent at flowering, at least half as long as receptacle.
22. Flowers 20.0-25.0 mm long; exterior of receptacle and calyx lobes with a long ferrugi-
neous sericeous pubescence; leaves glabrous (rarely glabrescent) beneath. 37. C. eriantha.
22. Flowers 7.0-20.0 mm long; exterior of receptacle shortly brown to gray tomentose;
leaves arachnoid pubescent beneath.
23. Leaves thick and coriaceous; bracteoles always persisting through flowering:
rachis of inflorescence and receptacle longitudinally striate, or if not petioles
13.0-18.0 mm long.
24. Flowers 7.0-I5.0 mm long; leaf bases subcordate rarely rounded; rachis and
receptacle with longitudinal striations. I5. C. bracteosa.
24. Flowers I8.0-22.0 mm long; leaf bases rounded to subcuneate; rachis and
receptacle not longitudinally striate. I7. C. belemii.
23. Leaves thin and membraneous; bracteoles persistent only in bud; rachis and
receptacle not longitudinally striate; petioles 6.0-i 1.0 mm long.
25. Stamens connate at base for at least I.0 mm; receptacle tapering to base, sub-
turbinate. 36. C. trapezioana.
25. Stamens free almost to base; receptacle subcylindrical. i6. C. subcordata.
21. Bracts and bracteoles not persistent at flowering; or small and inconspicuous.
26. Interior of receptacle filled with hairs to base.
27. Primary veins Io-I5; exterior of receptacle and calyx lobes with sparse appressed
pubescence not completely covering surface. 3. C. paraensis.
27. Primary veins 17-23; exterior of receptacle and calyx lobes densely pubescent,
completely covering surface.
28. Petioles canaliculate above; flowers I3.o-I7.0 mm long; exterior of receptacle
and calyx lobes with short brown pubescence; stamens more than 40. 19. C. excelsa.
28. Petioles not canaliculate; flowers 8.0-I2.0 mm long; exterior of receptacle and
calyx lobes with short sparse gray pubescence; stamens 20-35. i8. C. caryophylloides.
Couepia 207

26. Interior of receptacle glabrous except for deflexed hairs at throat.

29. Exterior of receptacle and calyx lobes sparsely appressed-puberulous, the pubes-
cence not forming a complete covering.
30. Stipules adnate to the base of petiole; rachis of inflorescence 3.0-5.0 mm thick;
petal margins glabrous. 37. C. stipularis.
30, Stipules caducous, not adhering to base of petiole; rachis of inflorescence
1.0-2.5 mm thick; petal margins ciliate.
31. Leaves bluntly or shortly acuminate to obtuse; receptacle subcampanulate,
3.0-5.0 mm thick at top.
32. Leaves ovate to oblong. 3. C. paraensis.
32. Leaves oblong-lanceolate. 5. C. maguirei.
31. Leaves prominently acuminate; receptacle cylindrical, ca 2.0 mm thick at
top.
33. Inflorescences erect, much-branched, predominantly terminal panicles.
34. Leaf undersurface densely short-arachnoid. I. C. guianensis.
34. Leaf undersurface glabrous or with sparse, easily rubbed-off pubes-
cence. 2. C. glandulosa.
33. Inflorescences little-branched racemose panicles which are axillary or
predominantly axillary but with a small terminal branch.
35. Inflorescences single racemose reflexed panicles; bracteoles persistent.
40. C. reflexa.
35. Inflorescences erect, terminal and in the upper 2-6 axils; bracteoles
caducous. 4. C. leptostachya.
29. Exterior of receptacle and calyx lobes with dense pubescence forming a complete
covering.
36. Leaf undersurface prominently reticulate.
37. Receptacle tapering gradually to a long slender pedicel; exterior of receptacle
rufous-pubescent; primary veins not impressed on upper surface. 42. C. cognata.
37. Receptacle changing abruptly to short thick pedicel; exterior of receptacle
brown-pubescent; primary veins impressed on upper surface. 29. C. racemosa.
36. Leaf undersurface with plane to prominent venation but not prominently reti-
culate.
38. Leaf undersurface glabrous; peduncles elongated (30.0-50.0 cm).
4I. C. longipendula.
38. Leaf undersurface usually pubescent, at least when young, rarely glabrous;
peduncle not elongated.
39. Receptacle slightly curved anteriorly in bud.
40. Leaves with midrib much impressed above. 42. C. cognata.
40. Leaves with midrib plane above.
41. Leaves 4.0-7.0 cm long; primary veins 7-I0. 53. C. pernambucensis.
4I. Leaves 9.0-1 I.0 cm long; primary veins 12-23.
42. Exterior of receptacle and calyx lobes gray-puberulous; primary
leaf veins 18-23, slightly impressed above. 51 C. impressa.
42. Exterior of receptacle and calyx lobes brown-tomentose; primary
leaf nerves 12-14, plane above. 43. C. multiflora.
39. Receptacle erect, not curved in bud.
43. Outer surface of petals distinctly pubescent.
44. Leaves 2.5-5.5 cm long; primary veins 5-9.
45. Leaves oblong-lanceolate to oblong, with acumen 5.0-7.0 mm
long, cuneate at base; petioles 5.0-7.0 mm long. 55. C. parvifolia.
45. Leaves orbicular to elliptic, with acumen 0-3.0 mm long, rounded
to cordate at base; petioles 2.0 mm long. 20. C. uiti.
44. Leaves (4.o-) 6.0-i6.0 cm long; primary veins 9-17.
46. Flowers 8.0-I2.0 mm long; receptacle cylindrical, ca 2.0 mm
thick at top below calyx lobes.
47. Inflorescence and flowers with dense ferrugineous sericeous
pubescence; flowers 8.0-I2.0 mm long; petioles 5.0-I0.0 mm
long; receptacle not striate on exterior. 52. C. meridionalis.
47. Inflorescence and flowers with short gray pubescence; flowers
6.o-8.o mm long; petioles 3.0-7.0 mm long; exterior of
receptacle longitudinally striate. 44. C. ulei.
46. Flowers I2.0-20.0 mm long; receptacle subcampanulate,
4.0-6.0 mm thick.
208 Flora Neotropica

48. Pedicels 6.0-15.0 mm long; exterior of receptacle and calyx

lobes short gray-sordid-puberulent. 2I. C. cataractae.
48. Pedicels not exceeding 6.0 mm long; exterior of receptacle
and calyx lobes spreading brown-tomentose.
49. Leaves 4.0-8.5 cm long x 2.0-3.8 cm broad, rufous-
pubescent beneath, graying with age. 45. C. comosa.
49. Leaves 7.5-I6.o cm x 3.5-8.0 cm broad, gray-pubescent
beneath.
50. Leaves thick, rounded to bluntly acuminate at apex.
27. C. grandifiora.
50. Leaves thin, prominently acuminate. 46. C. venosa.
43. Petals glabrous except for ciliate margins.
5I. Receptacle cylindrical or subcylindrical.
52. Leaves 4.5-12.0 cm long, cuneate to rounded at base; primary
veins 8-13.
53. Receptacle slender, cylindrical; stamens 20-28. 6. C. sandwithii.
53. Receptacle subcylindrical, broad; stamens 11-21. 47. C. polyandra.
52. Leaves I0o.o-7.0 cm long, subcordate to rounded at base;
primary veins 13-17.
54. Stamens ca 24, distinctly connate at base for I.0 mm; leaves
thick-coriaceous, rounded at base. 38. C. trapezioana.
54. Stamens ca 35, not distinctly connate at base; leaves thin-
membraneous, subcordate at base. I6. C. subcordata.
51. Receptacle broadly campanulate or turbinate.
55. Stamens I4-20.
56. Leaves ovate, 4.0-9.0 cm long; primary veins I I-I5; flowers
8.0-12.0 mm long; petioles 4.0-8.0 mm long. 25. C. ovalifolia.
56. Leaves oblong, io.o-I8.ocm long; primary veins I6-20;
flowers I I-.-I5.0 mm long; petioles 10.0-I5.0 mm long.
25. C. schottii.
55. Stamens more than 30.
57. Receptacle broadly turbinate, flattened, 3.0 mm long, almost
solid. 48. C. platycalyx.
57. Receptacle subcampanulate, not flattened, 5.0-12.0 mm long,
hollow.
58. Leaves 2.5-5.5 cm long x 1.5-3.5 cm broad. 20. C. uiti.
58. Leaves 9.0-27.0 cm long x 4.0-12.0 mm broad.
59. Leaf margins undulate and revolute. 49. C. rufa.
59. Leaf margins plane.
60. Leaves with short and blunt acumen, the lower sur-
face rufous-pubescent; calyx lobes rufous-pubescent
on exterior and yellow-brown-pubescent within.
50. C. robusta.
60. Leaves with finely pointed acumen, 8.o-I4.0 mm
long, the undersurface gray-pubescent; calyx lobes
gray-pubescent on both surfaces. 53. C.froesii.

A. The Couepia guianensis Superspecies (Species I-6).

I. Couepia guianensis Aublet, P1. Guiane I: 519, t. 207. I775; DC., Prodr.
2: 526. I825.
Acia amaraWilldenow, Linn., Sp. P1. ed. 4. 3(I): 717. i8oo.
Acioa amaraSteudel, Nom. ed. I. 9. I82I.
MoquileacouepiaSteudel, Nom. ed. 2. 2: 159. 1849.
CouepiaversicolorR. Benoist, Bull, Mus. Hist. Nat. Paris 29: 596. 1923; Kleinh., in Pulle, Fl.
Suriname 2(I): 453. 1939. Type. Melinonsn, French Guiana, fl (holotype, P).
CouepiasurinamensisKleinhoonte, Rec. Trav. Bot. N6erl. 22: 390. 1925. Type. B.W. 3080, Suri-
name, fl (lectotype, U).
Couepia 209

Medium to large-sized tree. Leaves oblong to oblong-lanceolate, 5.0-13.0 cm

long, 2.5-5.0 cm broad to 20.0 cm long on sterile branches, rounded at base, acumi-
nate at apex, with acumen 8.0-I8.0 mm long, glabrous above, densely gray lanate-
arachnoid beneath, often with two inconspicuous glands at base of lamina; midrib
glabrous above, prominent; primary veins o0-I5 pairs, prominent beneath; petioles
4.0-6.0 mm long, canaliculate above, villous when young, becoming glabrous when
mature. Stipules I.0-3.0 mm long, linear, caducous. Inflorescences terminal and
axillary panicles (few in upper leaf axils only), the rachis and branches sparsely gray-
brown pubescent. Bracts and bracteoles minute, ovate, caducous. Receptacle cylin-
drical, ca 6.o mm long, sparsely gray-brown puberulous on exterior, glabrous within
except for deflexed hairs at throat; pedicels 2.0 mm long. Calyx lobes rounded, 2.0
mm long. Petals 5, white, ciliate on margins, the surfaces glabrous. Stamens 14-20,
unilateral, with a toothed portion of ring opposite them. Ovary villous. Style pubes-
cent for half its length. Fruit round to ovate, 3.0-4.0 cm long, 2.5-3.0 cm broad;
epicarp smooth, glabrous; mesocarp thin, fragile, granular, glabrous within.
TYPE. Aublet sn, French Guiana, fr (BM).
DISTRIBUTION. Primary forest in the Guianas. Flowering July to October.
GUYANA. Fanshawe242(F.D. 2798) fl (BRG, K, NY, U). SURINAME. B.B.S. IIo3 st (BBS,
U); IIII St (BBS, U); B.W. 49a fl (U); I226st (U); 1333 fl (A, MO, U); 1339 st (U)1?; I755 st (U);
3027 fl (K, NY)10; 3371 fr (U)10; 3407 fr (U)10; 3584 fr (U)10; 5103 st (U); 6157 st (U); 6185 st (U);
6635 fl (NY, U); 6764 fr (NY); Van Emdensn fl (U); Heyligers466 st (U); Krukoff12293 fr (A, NY,
US); Lindeman4125 st (NY, U); 5794 st (U); 5923 st (NY, U); 6431 st (U). FRENCH GUIANA.
For. Serv.r12 fr (U); Melinonsn fl (P).

LOCAL NAMES. Suriname: Oenikiakia Djamaro, Kairiballi hohorodihoro,Apesia.

2. Couepia glandulosa Miquel, Stirp. Surin. Sel. 28. I850; Kleinh. in Pulle,
Fl. Suriname 2(I): 45I. I939.
Moquileaglandulosa(Miquel) Walp., Ann. 2: 463. 1852.
CouepiamyrtifoliaBentham ex Hooker f., Mart. Fl. Bras. 14(2): 44. I867, synon nov. Types.
Spruce3072, Brazil, Amazonas, fl (lectotype, K; isolectotypes, BM, BR, CGE, F, NY);
2262 fl (paratypes, BM, CGE, F, GH, GOET, K, LD, LE, M, NY, OXF, P); r74, Brazil,
Para, fl (paratypes, GH, K, NY, P).
CouepiathyrsifloraHooker f., Mart. Fl. Bras. 14(2): 45. I867, synon nov. Type. Spruce3681,
Venezuela, Terr. Amazonas, fl (holotype, K; isotypes, BM, BR, F, GH, GOET, LD, LE,
NY, P).
Parinari krukoviiGleason, Bull. Torrey Club 55: 353. 1933. Type. Krukoff1362, Brazil, Terr.
Rondonia, fl (holotype, NY; isotypes, K, MICH).

Tree to I2.o m high. Leaves oblong to oblong-lanceolate, 4.5-12.0 cm long,

2.2-4.5 cm broad, coriaceous to membraneous, rounded to subcuneate at base,
acuminate at apex, with acumen 5.0-I8.0 mm long, glabrous above, glabrous or with
sparse caducous pubescence beneath, with two glands at base of lower surface of
lamina (not always apparent); primary veins IO-I4 pairs, prominent beneath;
petioles 3.0-5.0 mm long, canaliculate above, pubescent when young. Stipules I.5-
3.0 mm long, caducous. Inflorescences terminal little-branched panicles, the rachis
and branches sparsely puberulous. Bracts and bracteoles minute, oblong, caducous.
Receptacle cylindrical, 4.5-7.0 mm long, often slightly curved anteriorly, sparsely
puberulous on exterior, glabrous within except for deflexed hairs at throat. Calyx
lobes rounded. Petals 5, white, ciliate on margins. Stamens I6-35, inserted in a
semicircle, with short staminodes opposite them. Ovary pilose. Style pubescent for
10= Paratypes of C. surinamensis.
210 Flora Neotropica

three fourths of length. Fruit elongate to ovoid, 2.5-3.0 cm long, 1.3-2.0 cm broad;
epicarp smooth, glabrous; mesocarp thin, fleshy; endocarp thin, fragile, granular in
texture, glabrous within.
TYPES. Hostmann &9 Kappler 859, Suriname, fl (holotype, U; isotypes, BM, GH,
GOET, K, LE, M, P).
DISTRIBUTION. Riverine forest, river banks and savanna margins in Venezuela,
the Guianas and the Amazon basin.
COLOMBIA. Meta: Jaramillo & Herndndez1027 fl (COL). Vaupes: Cuatrecasas6913 fl (COL,
F, NY, P, US); 7067 fl (COL, F, US); Ferndndez2338 fr (COL, US); Garcia-Barriga14282 fl (COL,
US); Schultes& Cabrera13145 fl (COL, GH, NY, US). Amazonas: Schultes& Cabrera16466 fl (GH,
LE, US); 17644 fl (GH, NY, US). VENEZUELA. Bolivar: Blanco51 fl (NY, VEN); 63 fl (NY, VEN);
265 fl (NY, VEN); Wurdack& Monachino40931 fl (F, GH, NY, US); 41028 fl (F, NY, US). Terr.
Amazonas: Level93 fl (F, GH, NY, US); LI. WilliamsI4640 fl (F, US); 15645 fr (F, VEN). GUYANA.
For. Dept. 334 fl fr (K); Jenman 6580 fl (BRG, K, NY). SURINAME. B. W. 2531 fr (U); 5427 fl (A,
MO, U); Dumortiersn fl (U); VanEmdenI333 fl (A); Hostmann1119 fl (BM, GH, K); Kappler2141 fl
(GOET, P); Lanjouw709 fl (U); 968 fl (IAN, K, NY, U); Versteeg172 fl (U); 715 fl (U). FRENCH
GUIANA. Halle 662 fl (P); Irwin, Pires & Westra48443 fl fr (F, GH, IAN, MG, NY, RB, US); Lemoine
(For. Serv.)7786 fl (MG, NY, U); 7793 fl (NY, U); 7850 fl (MG, NY, P, U); Melinonsn fl (BM, US); 18
fr (A, K, P); 54 fl (P); I49 fl (P); Sagot262 fl fr (K). BRAZIL. Amazonas: Black 48-2468 fr (IAN, LE,
NY); 48-2828 fr (IAN); 54-16957 fl (IAN, NY); Ducke RB 24993 fl (RB); RB 24994 fl (RB); Froes 2 1102
fl fr (F, IAN, K, NY, S, US); 21548 fr (IAN, K, NY); 22129 fl (COL, IAN, NY, U); 22431 fl (IAN);
22908 fl (NY); 33653 fr (IAN); 34888 fl (IAN); Oliveira2130 fl (IAN); Schultes8941 fl (IAN). Para:
Ducke 778 fl (F, IAN, MG, MO, NY, R, US); MG 9899 fl (MG); MG 12587 fl (BM, MG, RB); MG
12614 fl (BM, MG, RB); RB 15098 fl (U, US); MG 15632 fl (BM, MG, P, R, RB); RB 18227 fl (K,
U, US); RB 19755 fl (K, RB, U, US); Froes 30541 fl (IAN); 30588 fl (IAN); 30617 fl (IAN); 30654 fl
(IAN); Huber MG 854 fl (MG); Martius MG I6524 fl (BM, MG, RB); Pires & Silva 4614 fl (IAN,
INPA, NY); Prance& Pennington1265 fl (F, GH, NY); 1701 fr (F, MO, NY); Prance& Silva 58827 fl
fr (F, NY, US); SnethlageMG 9498 fl (MG). Terr. Amapi: Irwin, Pires & Westra47766 fl (F, GH,
IAN, MG, NY); 48390 fl (IAN, MG, NY). Maranhao: Frdes 1930 fl (A, BM, F, K, MO, NY, P, U,
US); 1945 fl (A, BM, F, K, MO, NY, P, U, US).

LOCALNAMES.Colombia: Wee-roo (Yakuna), Koo-yo (Puinave), Kan-he (Maka),

Cuyuy.French Guiana: Boliquin, Kobo, Kouebi.

3. Couepia paraensis (Martius & Zuccarini) Bentham, Jour. Bot. Hooker

2: 216. I840; Hook. f., Mart. Fl. Bras. I4(2): 48. 1867; Macbride, Fl.
Peru. io8i. 1938. Fig 34.
MoquileaparaensisMartius & Zuccarini, Abh. Akad. Munchen x: 390. I832.

Tree to 20.0 m tall, usually smaller. Leaves ovate to oblong, 5.5-I6.o cm long,
2.5-8.5 cm broad, rounded to cordate at base, blunt or short-acuminate at apex,
with acumen .0-5.0 mm long, glabrous above, usually densely gray to brown lanate-
arachnoid beneath, rarely glabrous or puberulous, two glands at base of lower surface
of lamina; primary veins IO-15 pairs, prominent beneath; midrib glabrous or
pubescent above; petioles 4.0-7.0 mm long, pubescent, canaliculate above. Stipules
I.0-2.0 mm long, caducous. Inflorescences terminal panicles, sparsely appressed-
pubescent. Bracts and bracteoles 0.5-2.5 mm long, ovate, caducous. Receptacle
subcylindrical to subcampanulate, 8.o-I2.0 mm long, sparsely appressed-pubescent
on exterior, completely hairy within or glabrous except for deflexed hairs at throat.
Calyx lobes rounded. Petals 5, white, ciliate on margins. Stamens 25-45, inserted in
half to two thirds of a circle, with short staminodes opposite them. Ovary pilose.
Style densely pubescent for three fourths of its length. Fruit ellipsoid, 3.5-4.0 cm long,
2.0-3.0 cm broad; epicarp smooth, glabrous; mesocarp thin, fleshy; endocarp thin,
fragile, granular in texture, the interior glabrous.
Couepia 211

V.Wiw +i /subsp. cerradoana

l 1'' | ?
o} C.0<paraensis subsp. paraensis
1 subsp. glaucescens
l IA gu.anens/s C.

/ B

FIG 34. Distribution of Couepiaparaensis and related species. A, C. paraensis subsp paraensis
(Fr6es25236), x 2.5; B, C. paraensissubsp glaucescens(Frdes26436) X 2.5.
212 Flora Neotropica

Key to the Subspecies of Couepiaparaensis

I. Interior of receptacle hairy throughout. a. subsp paraensis.
I. Interior of receptacle glabrous except for deflexed hairs at throat.
2. Bark of young stem thin; tree of forest and savanna margins. b. subsp glaucescens.
2. Young stem with a thick corky bark; tree of cerrado. c. subsp cerradoana.

3a. Couepia paraensis subsp paraensis Fig 34 A.

Couepiaparaensis(Martius & Zuccarini) Bentham, Jour. Bot. Hooker 2: 2I6. 1840.
Couepiarivalis Spruce, Not. Bot. Amaz. Andes I: I49. I908, nom nud.

Interior of receptacle hairy throughout; bark of young stem thin; growing in

sandy riverine localities.
TYPE.Martiussn, Brazil, Para, fl (lectotype, M).
Sandy river banks and beaches, and open riverine forest in eastern
DISTRIBUTION.
and central Amazonian Brazil.
BRAZIL. Amazonas: Ducke 1742 fl (F, GH, K, MG I7758, NY, US); MG 8409 fl (BM, MG);
Froes25086 fl (IAN, NY); 25236 fl (NY, RB); Pires 163 fl (COL, IAN, NY); Pranceet al 2649 fl (INPA,
NY); 3023 fr (INPA, NY); 3747 fl (INPA, NY); 4734 fr (INPA, NY); Rodrigues273 fl (INPA 4383,
MG, NY); 288 fl (INPA 4398, MG, NY); Rodrigueset al 2107 fl (INPA 9451); 2182 fr (INPA 8570);
2219 fr (INPA 8608, NY); 4816 fr (INPA 13349, NY); 4857 fl (INPA 13390, NY); 4998 fr (INPA
13659, NY); 6035 fl (INPA I5o86); 6775 fl (INPA 15239); Schwacke111-641 fl (GOET, R). Pari:
Black 47-1167A fl (IAN); 47-1765 fl (IAN, NY); Black & Ledoux50-10749 fl (NY); Capucho460 fl (F);
Cavalcante1669 fl (MG); 1726 fl (MG); Dahlgrensn fl (A, US); Derleysn fl (NY); Ducke 2948 fl (MG,
RB); 4924 fl (MG); MG 8409 fl (BM, RB); 8928 fl (BM, MG, RB); MG 10243 fr (MG); Monteiroda
Costa 132 fl (F); 277 fl (F, US); Pires & Silva 4350 fl (IAN, NY); Sampaio4978 fl (R); 5206 fl (R);
5251 fl (R); Spruce719 fr (K, P); 1014 fl (K, NY). Terr. Rond6nia: Pranceet al 6734 fl (INPA, NY).
Mato Grosso: Kuhlmann306 fl (R). "Guanabara: Glaziou 13798 fl (K, P)".11

3b. Couepia paraensis subsp glaucescens (Spruce ex Hook. f.) Prance, stat
nov. Fig. 34 B.
Spruce ex Hook. f., Mart. Fl. Bras. I4(2): 49. I867.
Couepiaglaucescens
CouepiaduckeiHuber, Bol. Mus. Emilio Goeldi 5: 371. I909, synon nov. Type. Ducke MG 8536,
Brazil, Amazonas, fl (holotype, MG; isotype, BM).

Interior of receptacle glabrous except for deflexed hairs at throat; bark of young
stem thin; growing in sandy riverine localities.
TYPE. Spruce 1752, Brazil, Amazonas, fl (holotype, K; isotypes, BM, CGE, F,
GH, GOET, LE, M, NY, OXF, P).
DISTRIBUTION.Sandy river banks and beaches and open riverine forest in
Venezuela and central and western Amazonia.
COLOMBIA. Arauca: Gomez 12 fl (US). VENEZUELA. Gusrico: Aristeguieta4589 fl (VEN);
5084 fr (VEN). Apure: Chardon247 fl (US, VEN); Codazzi & Pittier 9600 fl (NY, US); Codazzi sn fl
(VEN I900); Vlez 2274 fl (VEN); 2545 fr (VEN). Bolivar: Aristeguieta3005 fr (VEN); 5299 fr (VEN);
Avila i fl (VEN); Cardona2822 fl (VEN); Maguire & Maguire28998 fl (NY, US); 29021 fr (NY, US);
90816 fl (NY); Tamayo3439 fl (VEN); Velez 2545 fr (US); Ll. Williams
Ojasti sn fl (VEN); Steyermark
13305 fl (F, K, US); 11980 fr (VEN); 12037 st (S, VEN); Wurdack& Monachino39893 fl (NY, US);
40864 fl (GH, NY, US). Terr. Amazonas: Chaffanjon 332 fl (P); Curran1853 fl (NY); Maguire,Cowan&
Wurdack299o4A fl (NY, US); 30575 fl (NY, US); 30832 fl (NY, US); Maguire& Politi 27359 fl (F, NY,
US); 27854 fl (NY, US); 28143 fl (F, MO, NY, US); 28437 fl (F, GH, NY, US); Maguire, Wurdack
& Bunting36237 fl (NY, US); 36561 fl (NY, US); Molina & Barkleyi8V42 fl (US); LI. Williams 15937
fr (F, VEN); 15970 fr (F, US); Wurdack& Adderley42968 fl fr (NY, US). PERU. Loreto: Klug 1444 fl
11 See note on p 76
Couepia 213

(F, NY, US); 2520 fl (A, F, NY, U, US); Killip & Smith27184 fl (F, NY, US); 29963 fl (F, NY, US);
Tessmann5300 fl (NY); LI. Williams 1490 fl (A, F, US). BRAZIL. Amazonas: Ducke579 fl (F, IAN,
MG, MO, R, RB, US); 1329 fl (IAN, NY, R, RB, US); 2217 fr (NY, R); MG 7344 fl (BM, MG, RB);
RB 23591 fl (RB, US); RB 35562 fl (INPA, K, P, RB, U, US); Frdes20984 fl fr (F, IAN, K, NY, US);
22694 fl (IAN, IPA, NY); 26436fl (IAN, NY, U); Krukoff4576fl (A, BM, F, K, LE, MO, NY, U, US);
6709 fl (A, BM, BR, F, LE, MO, NY, U, US); Labroysn fl (P); Pires 169 fl (IAN, NY); Poeppig2503 fl
(F, GOET, LE, NY, OXF, P); Pranceet al 2648 fl (INPA, NY); 3350 fr (INPA, NY); 3369 fl (INPA,
NY); 4756 fr (INPA, NY); 6559 fr (INPA, NY); 6858 fl bud (INPA, NY); 8812 fr (INPA, NY);
10444 fr (INPA, NY); Rodrigues718 fl (INPA 6866, MG); 3921 fl (INPA I0486, NY); 5497 fl (INPA
14170, NY); 6720 fl (INPA 15I70); Rodrigues& Lima 2476 fl (INPA 9825); Rusby2678 fr (BM, F, GH,
K, LE, NY, P, US). Terr. Roraima: Pranceet al 4020 fl (INPA, NY). ParA: Ducke r249 fl (A, BM, F,
MO); 11665 fl (BM, MG, P, US); RB 19754 fl (K, RB, U, US); Martius2921 fl (M). Acre: Pranceet al
2964 fl (INPA, NY). Terr. Rond6nia: Cordeiro& Silva 161 st (NY). Mato Grosso: KuhlmannRB I7974
fl (RB, U, US).

LOCALNAMES. Venezuela: Querebere, Pilon. Brazil: Umarirana,Uchirana,Pajurd.

For a discussion of the subspecies paraensis and glaucescenssee note at the end of
this superspecies.

3c. Couepia paraensis subsp. cerradoana Prance, subsp nov. Ab. subsp
paraensis differt cortice suberoso crassissimo; receptaculo intus glabro.
TYPE.Prance& Silva58519, Brazil, Goias, Filadelfia, fl (holotype, NY; isotypes,
F, M, MO, UB).
DISTRIBUTION. Cerradoin the north of the Planalto of central Brazil.
BRAZIL.Maranhao:Carolina,Macedo4035fl (IAN, RB, SP, US); Pires& Black1718afl (IAN).
Para: AraguAia;Frdes30059 fl (IAN).

except for the thick corky bark

This subspeciesis identical to subspeciesglaucescens
which occurs from the young branches to the trunk. The interior of the receptacle is
glabrous, not filled with hairs as in subsp paraensiswhose geographical range comes
closer to that of subsp cerradoana Subspecies cerradoana
than to subsp glaucescens. grows
on the northermost limits of the cerrado whilst the other subspecies are confined to
the Amazonian forest regions. This subspecies and C. grandiflora are the only species of
Couepiawith a thick, corky bark. Subspecies cerradoana in the
differs from C. grandiflora
smaller flowers with glabrous petals (except for the ciliate margins), and in the
sparsely puberulous exterior of the receptacle.

4. Couepia leptostachya Bentham ex Hooker f., Mart. Fl. Bras. I4(2): 44.
1867; Ducke, Arq. Serv. Flor. Rio de Janeiro I(I): 3I. I939.
Huber,Bol. Mus. EmilioGoeldi6: 75. 1910,synonnov. Type. HuberMG2030fl
divaricata
Couepia
(holotype, MG; isotype, BM).
Couepiadivaricatavar strictiusculaHuber, Bol. Mus. Emilio Goeldi 6: 76. 1910. Types. R. S. Rodri-
gues MG 8274, Brazil, ParA, fl (syntype, MG; isosyntypes, BM, P, RB 15104, U); MG
9644 fl (syntype, MG; isosyntypes, BM, INPA, P).

Small tree. Leaves oblong-lanceolate, 6.o-I5.0 cm long, 2.0-5.o0 cm broad,

cuneate to subcuneate at base, acuminate at apex, with acumen I0.0-20.0 cm long,
glabrous above, with dense gray-brown-arachnoid pubescence beneath, the base of
lamina with two glands beneath; midrib prominent on both surfaces, glabrous above;
primary veins I2-I4 pairs, prominent beneath; petioles 5.0-8.0 mm long, pubescent
when young, conspicuously canaliculate above. Stipules ca I.O mm long, caducous.
214 Flora Neotropica

Inflorescences terminal and axillary racemes, or occasionally slightly branched

panicles, the rachis and branches with sparse appressed pubescence. Bracts and
bracteoles I.O-I.5 mm long, ovate, deciduous. Receptacle cylindrical ca 8.0 mm long,
sparsely gray-puberulous on exterior, glabrous within except for deflexed hairs at
throat; pedicels 2.0-4.0 mm long. Calyx lobes rounded, 2.0 mm long. Petals 5, white,
the margins ciliate. Stamens I5-22, unilateral, with a toothed portion of the circle
opposite them. Ovary tomentose. Style pubescent for three fourths of length. Fruit
oblong, 2.2-3.0 cm long; epicarp smooth, glabrous; mesocarp thin, fleshy; endocarp
fragile, of a granular texture, glabrous within.
TYPE. Spruce 1536, Brazil, Amazonas, fl (holotype, K; isotypes, BM, CGE, GH,
GOET, LE, M, NY, OXF, P).
DISTRIBUTION. Non-flooded forest in Amazonian Venezuela and Brazil. Flower-
ing May to September.
VENEZUELA. Terr. Amazonas: Croizat282 fl (NY). BRAZIL. Amazonas: ChagasINPA 199 fr
(INPA, MG, NY); Coelho& Mello INPA 3955 fl (IAN, INPA, MG, NY); DuckeRB 24992 fl (RB);
FerreiraINPA 5920 fr (INPA, MG, NY); Froes22675 fl (IAN); OsmarinoINPA 20739 fl (INPA, NY);
Pires 51816 st (IAN, NY); Pranceet al 3101 fr (INPA, NY); 9038 fr (INPA, NY); Rodrigues4891 fr
(INPA 13426); 5403 fl (INPA 14071, NY); 6012 fl (INPA I5062, NY); Rodrigues& Osmarino7883 fl
(INPA 17243, NY); 7893 fl (INPA 17253, NY). Pari: Black et al 57-19919 fl (IAN, NY, UB); Ducke
779 fl (F, IAN, MG 17747, MO, NY, R, US); MG 15336 fl (MG); MG 16319 fl (BM, MG, RB);
MG 17178 fl (MG); MG 17179 fl (MG); RB 15119 fl (RB); 18229 fl (RB); 18230 fl (RB); Goeldi
MG 8234 fl (BM, MG); Guedes256 fl (IAN, NY); MG 2141 fr (BM, MG, RB I5Ioo); INPA 9041 fl
(INPA); Kuhlmann& Jimbo 28 fl (FHO, IAN, MG 24017); 68 fl (FHO, INPA, MG 24021, SP, US);
92 fl (IAN, MG 24016, SP); I6I fr (IAN, MG 24013, SP); Moss sn st (US); Pires, Frdes& Silva 4860 fl
(IAN, NY); Pires, Black & Dobzhansky4146 fl (IAN, INPA, NY); Prance& Silva 58745 fr (F, NY, RB,
US); Registro44 fl (NY); Silva 62 fl (IAN, NY); 323 fl (IAN, U, US); 593 fl (IAN); Cavalcante222 fl
(MG). Terr. AmapA: Bastos 60 fl (F, INPA I6224, RB 96386, US); Oliveira3649 fr (IAN, NY); Pires,
Rodrigues& Irvine51045 fl (F, GH, INPA, NY, US); Silva 1133 fl (IAN, NY).

LOCAL NAMES. Brazil: Cumate, Oiti, Macuczi.

5. Couepia maguirei Prance, sp nov Fig 35 G-O.

Arbor vel frutex, ramulis juvenilibus sparse pubescentibus. Stipulae parvae,

caducae. Folia alternata, breviter petiolata, petiolo 4.0-5.5 mm longo, pubescenti,
supra canaliculato; laminae oblongo-lanceolatae, 5.o-II.0 cm longae, 1.5-4.0 cm
latae, apice in acuminem 3.0-6.0 mm longum contractae, basi rotundatae vel
cordatae, supra glabrae sat nitidae, subtus dense araneoso-tomentosae; costis secun-
dariis 1-i6 jugis, subtus prominentibus. Stipulae lineares, I.0-2.0 mm longae,
caducae. Flores in paniculis terminalibus dispositi; rachi ramisque sparsiuscule
griseo-pubescentibus; pediceli 2.0-3.0 mm longo. Bracteae bracteolaeque circa 0.5
mm longae, caducae. Receptaculum subcampanulatum, 8.o mm longum, extus
sparsiuscule griseo-tomentosum intus prope faucem pilis longis deflexis infra stamina
dense tomentosum, versus basim glabrum. Calycis-lobi 5, rotundati. Petala 5,
decidua, marginibus breviter ciliata. Stamina 55-60, in orbem completum disposita
Ovarium ad faucem receptaculi insertum, dense pilosum. Stylus e basi ovarii ortus,
pilosus. Drupa ovalis, extus glabra, laevis; endocarpio granulato, tenue fragilo, intus
glabro.
TYPE. Maguire 32714, Venezuela, Bolivar; Cerro Guaiquinima, Rio Paragua,
Isla Cassabe, fl (holotype, NY).
DISTRIBUTION. River islands and rocky places in Venezuela and the upper Rio
Negro, Brazil.
 Couepia 215

F(

M N 0 ~
':'.
G-, C.maguirei 3274) G, habit, xo.5; H, leafuppersurface,xo.;J, flower,x2.5; K,
(Maguire

FIG 35. Species of Couepia.A-F, C. sandwithii(Fanshawe7o8); A, habit, x 0.5; B, flower, x 2.5; C,

flower section, X 2.5; D, diagrammatic flower section, x 2.5; E, petal, X 2.5; F, ovary and style, X 2.
G-0, C. maguirei(Maguire32714); G, habit, x 0.5; H, leaf upper surface, x o.5; J, flower, x 2.5; K,
flower section, X2.5; L, diagrammatic flower section, x2.5; M, petal, x 2.5; N, ovary and style,
X2; O, fruit, xo.5.
216 Flora Neotropica

VENEZUELA.Terr.Amazonas:Rio GuainiabetweenComunidadand GuzmdnBlanco,

Maguire,Wurdack& Bunting35555 fr (NY); Alto Rio Negro. lI Williams14356 fr (F, US, VEN).
BRAZIL. Amazonas: Froes 21432 fl (K, NY, US).

This species is most closely related to C. paraensis,but differs in the smaller and
much narrower oblong-lanceolate leaves. It is with pleasure that I name it in honor
of Dr. Bassett Maguire whose studies of the genus Couepialaid the foundation for the
present work.

6. Couepia sandwithii Prance, s,) nov Fig 35 A-F.

Arbor magna, ramulis juvenilibus glabris. Stipulae parvae, deciduae. Folia

alternata, breviter petiolata, petiolo 4.0-6.0 mm longo, sparse pubescente, supra
canaliculato; laminae oblongae vel ellipticae, 4.5-9.0 cm longae, 2.0-3.5 cm latae, apice
in acuminem ad 6.0 mm longum contractae, basi cuneatae vel subcuneatae, superne
glabrae, subtus dense tomentosae; costis secundariis8-I4jugis, subtus prominentibus.
Stipulae circa I.o mm longae, caducae. Flores in paniculis terminalibus axillari-
busque dispositi; rachi ramisque breviter brunneo-tomentosis;pedicelli circa I.Omm
longi. Bracteae bracteolaeque ovatae, ad 3.0 mm longae. Receptaculum cylindricum,
3.0-4.5 mm longum, extus griseo-brunneo-tomentosum, intus prope faucem pilis
longis deflexis infra stamina dense tomentosum, versus basim glabrum. Calycis-lobi
5, rotundati vel acuti. Petala alba, decidua, margine breviter ciliata. Stamina 20-28,
in orbem completum vel 2/3 disposita, basi breviter connata. Ovarium ad faucem
receptaculi insertum, pilosum. Stylus e basi ovarii ortus, pilosus. Drupa ovalis, extus
laevis, glabra.
TYPE.Fanshawe708(F.D. 3444), Guyana, Mazaruni Station, fl (holotype, NY;
isotypes, BRG, K, U, US).
DISTRIBUTION. Primary forest in Guyana. FloweringJune to October.
GUYANA. Moraballi Creek, Fanshawe584(F.D. 3320) fl (BRG, F, K, NY); Apoteri, Rupununi
River, Davis 87(F.D. 2028) fl (BRG, K); Mazaruni Station: Fanshawe709 (F.D. 3445) fl (NY).

This species differs from C. guianensisand C. glandulosain the dense, appressed

pubescence of the receptacle, in the obtuse to bluntly acuminate leaves, and in the
persistent bracteoles. This is a large forest tree, while the other members of the C.
guianensisspecies complex are shrubs or at most small trees.
Taxonomic Discussion of the CouepiaguianensisSuperspecies
This, the largest and most complex group of species in Couepia,consists of im-
perfectly allopatric components. This group is characterized by the flowers with
subcylindricalreceptacles that are sparselypubescent on the exterior. The stamens are
usually unilateral and the style is densely pubescent for most of its length. The fruit
is round to ovoid with a smooth, glabrous epicarp and a granular rather thin and
fragile endocarp.
The most distinct species in this group is C. leptostachya. The inflorescence of this
species is characteristic. It is usually simply racemose or occasionally slightly branched
but always racemose in appearance. In addition to the terminal raceme the in-
florescence are borne in the last 2-6 leaf axils. The long tubular receptacle is also
distinctive. The remaining species of this group have a paniculate inflorescencewhich
is predominantly terminal. Couepialeptostachya is an Amazonian species described
originally from the Rio Negro, Amazonas, Brazil. With more recent material it has
become clear that it is a wide ranging species in the Amazon basin. (See Fig 34).
Couepia 217

Examination of all available material assignable to this complex has led to

acceptance of only three species. The most important differences among them are
summarized in the Table II.

Table II
Principal Differences among Species of CouepiaguianensisSuperspecies.

Character C. guianensis C. glandulosa C.paraensis

LEAF apex prominently acuminate prominently acuminate bluntly acuminate
or rounded
indumentum below dense, arachnoid glabrous or sparse dense, arachnoid
rarely glabrous

RECEPTACLE width + 2.0 mm + 2.0 mm 3.0-5.0 mm

shape subcylindrical subcylindrical subcampanulate
PEDICEL distinct, slender gradually tapering gradually tapering
from receptacle from receptacle

A comparison with the Aublet type specimen at the British Museum has shown
that C. guianensisis conspecific with C. versicolorand C. surinamensis.The fruit and the
indumentum of the leaf are exactly similar. Couepiaguianensisis most closely related
to the more inland C. glandulosa.
According to the type descriptions, C. myrtifoliadiffers from C. glandulosain the
shorter, broader, non-membraneous leaves lacking two glands at the lamina base, as
well as in the simpler, little-branched racemes and the shorterpedicels. In the original
description, C. thrysiflorawas said to differ from C. myrtifoliain the longer, broader
leaves, the densely flowered inflorescence, and the cylindrical receptacle. A cursory
examination of the type specimens of these three species showed that even the above
differences are insufficient. In a study of a considerable amount of additional material
of these species, it was found that there were no correlations among the above
characters, and that there is no additional information to justify maintaining these
three species. There is a complete range of leaf shape and size, and scatter diagrams
demonstrated no break in the foliar character. In almost every specimen, there are
two conspicuous and distinct glands at the leaf base. The densely flowered inflores-
cence typical of C. thrysiflora also occurs occasionally in specimens of C. myrtifoliaand
C. glandulosa.The only relatively constant character is the texture of the dried leaf.
The type gathering of C. glandulosaand other material from Suriname and French
Guiana has membraneous leaves, while material from further East has distinctly
coriaceous leaves. However, in several cases intermediates have been collected, in
which cases it is impossible to say whether the leaf is coriaceous or membraneous.
For these reasonsC. myrtifoliaand C. thyrsiflora are here regarded as part ofC. glandulosa.
Parinarikrukovii,of erroneousgeneric assignment, is also synonymouswith C.glandulosa.
Couepiaparaensis was first described by Martius. There are two Martius collections
at the Munich herbarium (Martius sn and 2921). One of these collections has a
feature rarely found in Couepia,the interior of the receptacle-tube densely pubescent
to the base. In the other Martius collection the receptacle is glabrous inside towards
the base. The large amount of recent material also shows this variation, as is shown in
Fig 34. This map indicates that the hairy receptacle occurs only in the western part
of the range of this species. Around Manaus both types occur together, but as yet no
intermediates have been discovered. Couepiaparaensisshows considerable variation in
stamen number, flower size and leaf shape, but none of these features is in any way
correlated with the receptacle type. As the original Martius specimens suggest, there
are no grounds for discerning two separate species. However, in all other species of
the Chrysobalanaceae having a pubescent interior of the receptacle, this character is
218 Flora Neotropica

diagnostic and correlated with others (eg in Parinarisp, Couepiaexcelsa,and C. caryo-

phylloides).Since this is an important feature in the family and since there is such
definite geographical distribution, with respect to it, there is a good basis for recog-
nizing subspecies. In the original description of C. paraensis,Martius mentioned that
the interior of the receptacle is hairy, so this element is taken to represent subspecies
paraensis.The glabrous element has been treated as subsp glaucescens.
Couepia paraensisis distinct from the other species already discussed by its blunter
and less acuminate leaves, the thicker subcampanulate receptacle, and the tendency
toward a larger number of stamens. In the original description, C. glaucescenswas
distinguished from C. paraensissolely by a difference in leaf and flower size. In C.
paraensisthe leaves are 3.0-5.0 inches long, the flowers 0.50-0.66 inches long; in C.
glaucescensthe leaves are 2.0-4.0 inches long, and the flowers 0.3 inches long. In fact,
flowers and leaves of both species vary considerably in length, scatter diagrams
showing no such distinction possible.
In addition to the four species discussed above, two new species belonging to this
complex have been described. One is from Venezuela and the other from Guyana.
The differences from other species in this complex are mentioned under the pertinent
species descriptions.

B. The Couepia parillo superspecies (Species 7-IO)

7. Couepia parillo A. P. de Candolle, Prodr. 2: 526. 1825; Prance, Atas do

Simp6sio sobre a Biota Amazonica 4: 220-222. 1967. Fig 36.
CouepiapaucifloraHuber, Bol. Mus. Emilio Goeldi 5: 372. I909, synon nov. Type. DuckeMG 8630
Brazil, Para, fl (holotype, MG; isotype, BM).
Couepiavillosa Maguire, Bull. Torrey Club 75: 378. I948, synon nov. Type. Maguire 24782,
Suriname, fl (holotype, NY; isotypes, BR, F, M, MO).
Moquileaparilla (A. P. de Candolle) Steudel, Nom. ed. 2. 2: 159. I841.
Tree to 20.0 m tall or shrub. Leaves oblong or oblong-elliptic, 5.0-15.5 cm long,
1.7-5.8 cm broad, acuminate at apex, with acumen 5.5-i8.0 mm long, rounded to
cuneate at base, glabrous above, prominently reticulate beneath, with silver-gray
pubescence between the reticulations; midrib impressed and pubescent above;
primary veins IO-I5 pairs, prominent beneath, slightly impressed above; petioles
2.0-9.0 mm long. Inflorescences racemes of 5-2I flowers, the rachis and branches
with ferrugineous-brownpubescence. Stipules 1.5-5.0 mm long, linear. Bracts and
bracteoles 2.0-6.0 mm long, linear, persistent. Receptacle slender, tubular, II.0-
22.0 mm long, with thick dense hirtellous pubescence, and some longer hairs on
exterior, glabrous within or with hairs nearly to base on proximal side beneath ovary
only; pedicels I .0-3.0 mm long. Calyx lobes acute. Petals 5, white, glabrous on margins.
Stamens 45-62, inserted almost in a complete circle, with a short portion of toothed
staminodes. Ovary densely villous. Style sparsely pubescent at base. Fruit globose,
ca 2.5 cm diameter; epicarp soft yellow-brown-velutinous; endocarp thin and fragile,
granular in texture, sparsely hirsute within.

FIG 36. Variation and geographical distribution of species of Couepia.A, A-C, C. parillo; A,
Boyan70; B, Gleason539; C, Tessmann3690. *, D, C. canomensis;D, Rodrigueset al 5937; leaves, x 0.25;
flowers, X I.25.
 Couepia 219

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k. ~~~~~~~~~~I
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-?

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;I<:^ ^'-^^tiiLiS 1*
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:C i.
" / *1c "**"''"'^^.V-
../-' %--,?..<
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220 Flora Neotropica

TYPE. Herb. Ventenatsn, French Guiana, fl (holotype, G).

DISTRIBUTION.Savannas or primary forest, and also common in secondary forest
in the Guianas and Amazonian Peru and Colombia.

COLOMBIA. Vaup6s: Ferndndez2305 fl (COL, US). Amazonas: Schultes & Black 46-342 fl
(IAN, US). PERU. Loreto: Klug z42 fl (F, NY, US); Tessmann3690 fl (NY, US); LI. Williams3780 fl
(BM, F). GUYANA. Boyan 7o(F.D. 7754) fl (K, NY, U); Davis 437(F.D. 2432) fl (BRG, K); De La
Cruz 1825 fl (F, GH, MO, NY, US); 3468 fl (F, GH, NY, US); Fanshawe377(F.D. 3133) fr (BRG);
902(F.D. 3638) fr (BRG, K, NY, U, US); I224(F.D. 3690) fl (BRG, K); i5o6(F.D. 4242) fl (K);
Gleason539 fl (GH, K, NY); Guppy266(F.D. 7202) fl (NY); 669(F.D. 7684) st (NY); Jenman 4184 fl
(K, NY); Persaud142 fl (F, K, NY); Sandwith315 fl (BRG, K, NY, RB, U); Whitton98 st (K); 288 fl
(K). FRENCH GUIANA. Cowan38854 fl fr (GH, NY, US); Richardsn fl (P). BRAZIL. Para: Oliveira
41 I fr (IAN, NY); Silva 1130 fl (IAN, NY). Terr. Amapa: Pires,Rodrigues& Irvine50843 fl (IAN, MG,
NY).

LOCALNAME.Guyana: Hatcheballi. Aimoradan.

8. Couepia steyermarkii Maguire, Fieldiana Bot. 28: 252. 1952.

Medium-sized tree. Leaves elliptic to oblong-ovate 4.0-6.0 cm long, 2.7-4.0 cm

broad, rounded to cuneate at base, shortly acuminate at apex, with acumen I.O-
3.0 mm long, glabrous above, prominently reticulate beneath, gray-lanate between
the reticulations; primary veins 9-I2 pairs, prominent beneath, impressed above;
midrib impressedabove, pubescent at base; petioles 5.0-7.0 mm long, short-hirtellous.
Stipules to 2.0 mm long, subulate, caducous. Inflorescences few-flowered axillary
racemes, the rachis and branches with dense ferrugineous pubescence. Bracts and
bracteoles I.0-3.0 mm long, ovate, persistent. Receptacle subsessile, cylindrical,
8.o-io.o mm long, short-hirtellous on exterior, glabrous within except for deflexed
hairs at throat. Calyx lobes acute, ovate. Petals 5, white, glabrous on margins.
Stamens inserted on two-thirds of the circle. Ovary densely villous. Style pubescent
at base. Fruit unknown.
TYPE.Steyermark 60025, Venezuela, Bolivar, fl bud (holotype, F; isotypes, NY,
VEN).
DISTRIBUTION. Known only from the type gathering collected in forests on the
southeastern slopes of Ptari-tepui.

9. Couepia canomensis (Martius) Bentham ex Hooker f., Mart. Fl. Bras.

14(2): 42. 1867; Ducke, Arq. Serv. Flor. Rio de Janeiro i(I): 29. 1939;
Prance, Atas do Simp6sio sobre a Biota Amazonica 4:220-222. 1967. Fig 36.
MoquileacanomensisMartius, Nov. Gen. et Sp. 2: 80. I827.
Hirtella canomensis(Martius) Sprengel, Linn. Syst. Veg. ed. I6. 4(Cur. Post.): 341. 1827.
ParinaripilosaStandley, Publ. Field Mus. Bot. 17: 257. 1937. Type. Krukoff70o fr (holotype, F;
isotypes, A, BM, BR, IAN, K, LE, MICH, MO, NY, U).

Shrub or small tree to Io.o m tall. Leaves ovate to oblong-elliptic, 8.5-I8.o cm

long, 3.0-8.0 cm broad, rounded to subcuneate at base, short-acuminate at apex,
with acumen 2.5-12.0 mm long, glabrous above, prominently reticulate beneath
with short gray-brown pubescence between reticulations; primary veins Io- 5 pairs,
prominent beneath, slightly impressed above; midrib impressed and usually pubes-
cent above; petioles 6.0-i I.o mm long, with dense ferrugineouspubescence. Stipules
3.0-8.0 mm long, linear, persistent. Inflorescences densely-flowered axillary racemes,
Couepia 221

the rachis and branches ferrugineous-hirtellous.Bracts and bracteoles 2.5-7.0 mm

long, persistent. Receptacle subcampanulate-cylindrical, 5.5-10.0 mm long, dense
ferrugineous-brownpubescent on exterior, glabrous within except for deflexed hairs
at throat; pedicels ca I.o mm long. Calyx lobes acute, hirsute on exterior. Petals 5,
white, glabrous on margins. Stamens 40-5I, inserted in a nearly complete circle,
with a short row of toothed staminodes opposite them. Ovary densely villous. Style
pubescent at base only. Fruit round to oblong, 3.5-5.0 cm long, 2.5-3.5 cm broad;
epicarp short-velutinous; endocarp thin, fragile, granular, sparsely-pubescent within.
TYPE. Martius sn, Brazil, Amazonas, fl fr (holotype, M), st (isotypes, BR, K).
DISTRIBUTION. Primary and secondary forest and clearings on non-flooded ground
in western and central Amazonia and southern Guyana.
COLOMBIA. Vaupes: Ferndndez2153 fr (COL, US); RomeroC. 3497 fl (COL); Schultes &
Cabrera13392 fl (GH, LE, US); 14962 fr (US); 17325 fl (F, GH, US). VENEZUELA. Terr. Amazonas:
Maguire,Cowan& Wurdack29312 fr (NY); LI. Williams14181 fr (F); I5055 fr (F, US, VEN). GUYANA.
Tillett & Tillett 45573 fl (NY, US). BRAZIL. Amazonas: CoelhoINPA 1259 fl (IAN, INPA, MG, NY);
3011 fl (INPA, NY); ChagasINPA 784 fr (INPA, NY); INPA 4206 fl (IAN, INPA, MG, NY); INPA
4299 fr (INPA, NY); Ducke464 fl (A, F, K, MO, NY, R, US); 1297 fl (A, F, IAN, MG 17752, NY, R,
US); RB 19750 fl (RB, U, US); RB 23590 fl (RB); RB 24989 fl (RB); RB 35560 fl (RB); Ferreira87/57
fr (INPA 5835, NY); Frdes21204 fl fr (F, IAN, K, NY, US); 24910 fl (IAN, UB); 25059 fl (IAN);
GoeldiMG 3993 fl (MG); Killip & Smith30031 fr (NY, RB, US); Krukoff6896 fr (A, BR, K, LE, MO,
NY, RB, U, US); 7983 fl (A, BM, BR, K, LE, MO, NY, P, U); Kuhlmann963 fr (RB I7972); Mello
INPA 2116 fr (INPA, NY); Pires 135 fr (IAN, K, NY); Pires & Silva 9725 fl (IAN); Pranceet al 3633 fr
(INPA, NY); 3850 fl fr (INPA, NY); Rodrigueset al 1360 fr (INPA 7690, NY); 2148 fr (INPA 8530);
2686 fr (INPA I0036); 2771 fr (INPA 10121); 2785 fr (INPA 10I35); 3486 fr (INPA 10036, NY);
3571 fr (INPA Io112, NY); 3585 fl (INPA 10135, NY); 4058 fr (INPA Io623, NY); 5367 fl (INPA
14036, NY, RB); 5937 fl (INPA 1488I, NY); 7135 fl (INPA 15797, NY); Spruce2318 fl (BM, BR, F,
K, LE, NY, OXF, P). Para: Egler 925 fl (IAN, MG 23678); Hoehne5152 fl (R).

LOCALNAMES.Brazil: Cuparana,Macuciu.

o1. Couepia foveolata Prance, sp nov Fig 37 E-J.

Arbor ad 30.0 m alta, ramulis juvenilibus hirtellis mox glabris. Folia alternata,
breviter petiolata; petiolo 5.0-9.0 mm longo, hirtelli pubescenti; laminae oblongae
3.5-9.5 cm longae, 1.5-3.7 cm latae, apice in acuminem 3.0-7.0 mm longum con-
tractae, basi subcuneatae vel cuneatae, supra glabrae sat nitidae, subtus reticulatae,
cavas stomatales ferentes lanato-tomentosae; costis secundariis 9-11 jugis, subtus
prominentibus; costa media supra impressa, versus basim pubescentia. Stipulae ca
I.o mm longae, subulatae, caducae. Flores in racemis axillaribus paucifloris dense
ferrugineo-brunneo-pubescentibus dispositi. Bracteae bracteolaeque oblongae, ad
7.0 mm longae, persistentes. Receptaculum subcylindricum, 8.o-Io.o mm longum,
extus hirtello pubescens, intus infra faucem pilis deflexis tomentosum, versus basim
glabrum. Calycis-lobi 5, acuti. Petala 5, decidua, margine glabra. Stamina circa 30,
unilateralia, staminodiis dentatis opposita. Ovarium villosum. Stylus e basi ovarii
ortus, versus basim pilosus. Drupa ovalis, 2.6 cm lata, 3.2 cm longa, extus verrucosa;
mesocarpio tenue, carnoso; endocarpio tenue, granulato, intus sparse hirsuto.
TYPE. Steyermark& Wurdack 62, Venezuela, Bolivar, Rio Caroni, fl (holotype,
NY; isotype, F).
DISTRIBUTION. Riverine forest in Venezuela.
VENEZUELA. Bolivar: Rio Caroni, Cardona2489 fr (US, VEN); 2555 fr (VEN); Chimanta
Massif, Steyermark74709 fl (F, NY, VEN); 75555 fr (F, NY, VEN); Uriman Region, Bernardi757
fl (NY), 904 fr (NY).

LOCALNAME.Makara (Arekuna).
222 Flora Neotropica

The relationship of this species with the C. parillosuperspeciesis discussed in the

general discussion of the superspecies.C.foveolatais also close to C. cognata,one of the
taxonomically isolated species. However, C. cognatais different in a number of
significant characters. In C. cognatathe inflorescence is paniculate, the inflorescence
and flowers have a characteristic hirsute pubescence, and the arachnoid pubescence
of the leaf undersurfacecompletely obscuresthe reticulate pattern of the leafvenation.
In addition to this C. cognatadoes not have stomatal cavities.
This species and Couepiaracemosaare the only species of Couepiaknown to have
distinct stomatal cavities on the undersurface of the leaf. This feature is almost uni-
versal in Parinariand is common in Licania.C.foveolatadiffers from C. racemosa in the
fewer-floweredunbranched inflorescence, the more prominent stomatal cavities and
the eglandular petioles.
Taxonomic Discussion of the Couepiaparillo Superspecies
This group is easily recognized by the conspicuous reticulation of the leaf under-
surfaces. The reticulation is conspicuous because of the prominent parallel secondary
leaf nerves connecting the primary veins, and the silver-gray pubescence in the
intervening spaces. Two of the taxonomically isolated species, C. elataand C. racemosa
also have this type of reticulation. In addition to the reticulate leaves, the C. parillo
superspecies is characterized by its racemose inflorescence with a dense ferrugineous-
brown hirtellous pubescence, its slender cylindrical receptacle tube usually with
villous hairs on the exterior, and the fruit with a softly tomentose exterior.
The Venezuelan material of this complex is markedly distinct from the rest. All
specimens collected so far from the Guayana region of Venezuela appear to belong to
C. steyermarkii and the new species C.foveolata.The latter is clearly distinct from others
of this group by the stomatal cavities of the leaf undersurfaces.Couepiafoveolata also
differsin having fruit with a verrucose exterior (the fruit ofC. steyermarkii is unknown).
Couepiafoveolata is closest to C. steyermarkii but, in addition to the stomatal cavities, it
differs in the longer, more oblong leaves which lack markedly impressed primary
veins, and in the longer leaf acumen. In C. steyermarkii the foliar nerves are much more
prominent. The floral and fruit charactersof these two species cannot be compared at
present because C. steyermarkii is as yet imperfectly known. Couepiafoveolataand C.
steyermarkii are two of the three species of Couepiaconfined to the Guayana region.
Couepiasteyermarkii is close to the other two species of the C. parillo superspecies, but
appears to be clearly distinct. It differs from C. parillo in the short receptacle and in
the elliptic leaves with a much shorter acumen, and from C. canomensis in the smaller
elliptic leaves, and in the small few-flowered inflorescences.
Although it may seem inconsistent to include C. canomensis and C. steyermarkii in
a species complex with C. parillo, it must be pointed out that some material (eg
Martinsn, BM, K, NY) which deviates markedly from the characteristicsof the type,
has been distributed under the name C. parillo. The type of C. parillo, apparently
overlooked hitherto, corresponds well with the type of C. pauciflora,necessitating
reduction of the latter name to synonymy. The material previously anotated as C.
parillo in some herbaria is described here as the new species C. martinii(p 238).
Couepiaparillo, described only synoptically by de Candolle in the "Prodromus", has
never been described more fully under that name. Couepia parillois most closely related
to C. canomensis and C. steyermarkii.
Couepiaparillo, as defined here, has a markedly disjunct geographical distri-
bution, (Fig 36). The Peruvian material, previously considered as C. canomensis,
unquestionably belongs to C. parillo. It would be premature to draw many conclu-
sions from this datum until further material has been collected, but such a pattern of
distribution is not unknown in other genera, (eg Connarusand Ficus) and several
Couepia 223

t, o
D 1IF

""" G

FIG 37. Species of Couepia.A-D, C. meridionalis(Mosin 3377); A, habit, x o.5; B, flower, x 2; C,

flower section, x 2; D, petal, x 3.5. E-J, C.foveolata(Steyermark et al 62, 74709); E, habit, x o.5; F, flower,
x i.5; G, flower section, x i .5; H, young fruit, x I; J, ovary section, x I5. K-O, C. pernambucensis (Lima
53-1206); K, habit, x o.5; L, flower, x 3; M, flower section, x 3; N, petal, x 5; O, fruit, x2.
224 Flora Neotropica

species of Couepia(e.g C. chrysocalyx),range from the Guianas through Brazil to Peru.

Couepiavillosa was separated from C. pauciflora(=C. parillo) in the original
description partially by its leaf shape, but because of great variation in leaf shape in
C. parillo, this character is not useful in defining species. Couepiavillosawas also dis-
tinguished by the fimbriate sterile portion of the staminal circle. However, a small
portion of the base of the staminal circle is often fimbriate in C.parilloand C. canomensis.
The only remaining significant character, the short villous tomentum of the in-
florescence and the flower, is shared with some specimens ofC. parillo (eg PersaudI42
NY), and many of C. canomensis(eg Tillett & Tillett 45573, NY). Thus there is no
evidence supporting the separate maintenance of C. villosa. Since C. parillo and C.
canomensisare very closely related and easily confused, a summary of the main
differences, representing the results of a scatter-diagram analysis of the main charac-
ters, is given below.
C. canomensis
Leaves ovate to oblong-elliptic, 8.5-i8.o cm long, 3.0-8.0 cm broad (mean
length : breadth ratio 2.II), with a short acumen 2.5-12.0 mm long (mean 5.9).
Inflorescences many-flowered racemes (mean 20 flowers). Receptacle short, obconi-
cal-tubular, 5.5-I0.0 mm long (mean 6.3). Fertile stamens 40-51.
C. parillo
Leaves oblong to oblong-elliptic, 5.0-I5.5 cm long, 1.7-5.8 m broad (mean
length: breadth ratio 2.63), with a well developed acumen 5.5-I8.o mm long (mean
10.3). Inflorescences sparingly-floweredracemes (5-21, mean 9 flowers). Receptacle
long, slender, tubular, I1.0-33.0 mm long (mean I5.8 mm). Fertile stamens 45-62.

C. The Couepia magnoliifolia Superspecies (Species 1-I 4).

ii. Couepia magnoliifolia Bentham ex Hooker f., Mart. Fl. Bras. 14(2): 43.
1867; Ducke, Arq. Serv. Flor. Rio deJaneiro I(I): 30. 1939.

Small to medium-sized tree. Leaves oblong, I4.0-24.0 cm long, 5.5-8.5 cm

broad, rounded to subcuneate at base, shortly acuminate at apex, with acumen
3.0-5.0 mm long, glabrous above, with dense rufous-brown arachnoid pubescence
beneath; primary veins I5-18 pairs, prominent beneath, slightly impressed above;
petioles 7.0-8.0 mm long, tomentose, terete. Stipules linear, to I.o mm long, mem-
braneous, caducous. Inflorescences dense axillary or terminal racemose spikes, the
rachis ferrugineous-brown-sericeous. Bracts ovate, 5.0-7.0 mm long, caducous.
Receptacle subcylindrical, 8.o-9.o mm long, ferrugineous-sericeous on exterior,
glabrous within except for deflexed hairs at throat; pedicels I.o mm long. Calyx
lobes rounded. Petals 5, white, with ciliate margins. Stamens ca 25, inserted in a
complete circle. Ovary pilose. Style densely pubescent for three fourths its length.
Fruit ellipsoid; epicarp smooth, glabrous; pericarp thick, fragile, glabrous within.
TYPE.SpruceI4r4, Brazil, Amazonas, fl (holotype, K; isotypes, CGE, F, GH,
GOET, LD, LE, NY, OXF, P, RB).
DISTRIBUTION. Known only from non-flooded forest in the vicinity of Manaus,
Brazil. Flowering February to June.
Couepia 225

BRAZIL. Amazonas: ChagasINPA 3576 fl (IAN, INPA, MG, NY); Ducke 156 fl (A, F, IAN, K,
MG 17755, MO, NY, R, US); RB 24987 fl (F, INPA, K, RB, U, US); RB 24988 fr (K, RB, UB);
RB 35559 fl (K, RB); INPA 9043 fl (INPA); Pranceet al 4696 fl (INPA, NY); Rodrigueset al 2165 fr
(INPA 9509); 2770 fr (INPA 9313, NY); 2965 fr (INPA 9509, NY); 2974 fr (INPA 95I8, NY).

LOCAL NAME. Pajurd.

12. Couepia exflexa Fanshawe & Maguire, Bull. Torrey Club 75: 376. I948.

Tree to 30.0 m tall, Leaves oblong, Io.o-i8.o cm long, 5.0-8.0 cm broad,

obtuse to cuneate at base, apiculate at apex, 7.0-Io.o mm long, glabrous above
(sparsely-ferrugineous-pubescenton young leaves), brown lanate-arachnoid beneath;
midrib prominent above, glabrous except at base; primary veins I4-I7 pairs, promi-
nent on lower surface; petioles IO.O-I2.0mm long, sparsely pubescent, eglandular.
Stipules minute. Inflorescences densely-flowered spikes 3.0-6.0 cm long with a
reddish-brown pubescence. Bracts 5.o-8.o mm long, ovate, equalling the receptacle
tube in length. Receptacle subsessile, 7.0-Io.o mm long, cylindrical but strongly re-
curved distally on anterior, dense red-brown pubescent on exterior, glabrous within
except for deflexed hairs at throat. Calyx lobes acute. Petals 5, white, deciduous,
ciliate on margins. Stamens 15-19, slightly connate at base, unilateral, with 5-6
short filamentous staminodes opposite them. Ovary pilose. Style sparsely pubescent
at base. Fruit oblong-oval; epicarp smooth, glabrous; pericarp thin and hard,
sparsely pubescent within.
TYPE. Fanshawe722 (F.D. 3458), Guyana, fl (lectotype, NY; isolectotypes,
BRG, K); 722a fr (paratypes, K, NY).
DISTRIBUTION.Known only from primary forest around Mazaruni Station,
Guyana.
GUYANA. Fanshawe723 (F.D. 3459) fl (NY).

This species is most closely related to C. magnoliifolia from which it differs in the
distinctive reddish-brown pubescence and the curvature of the receptacle as well as
in the persistence of the bracteoles. In C. magnoliifoliathe bracteoles are large but
tend to fall early.

13. Couepia habrantha Standley, Publ. Field Mus. Bot. 17: 249. I937;
Ducke, Arq. Serv. Flor. Rio de Janeiro I(i): 30. 1939.

Tree to I2.0 m tall. Leaves oblong-elliptic, 6.0-I3.0 cm long, 2.5-6.0 cm broad,

cuneate to rounded at base, acuminate at apex, with acumen 4.0-I4.0 mm long,
glabrous above, densely reddish-brownto gray-arachnoidbeneath; midrib prominent
and glabrous on upper surface; primary veins II-I5 pairs, prominent beneath;
petioles 5.o-8.o mm long, canaliculate above. Stipules to 2.0 mm long, caducous.
Inflorescences short, dense, axillary and terminal spikes, the rachis with ferrugineous-
brown pubescence. Bracts and bracteoles 7.0-9.0 mm long, ovate, acute, deciduous.
Receptacle subsessile, subcylindrical, 8.0-io.o mm long, densely ferrugineous-
sericeous on exterior, glabrous within except for deflexed hairs at throat. Calyx lobes
226 Flora Neotropica

rounded. Petals 5, white, ciliate on margins. Stamens ca 25, inserted in a semicircle,

with short filamentous staminodes opposite them. Style densely pubescent for three
fourths its length. Ovary pilose. Fruit ellipsoid; epicarp smooth, glabrous; mesocarp
thin, fleshy; pericarp thin, fibrous and fragile, glabrous within.
TYPE. Krukoff7252, Brazil, Amazonas, fl (holotype, F; isotypes, A, BM, BR,
IAN, K, MO, NY, U, US).
DISTRIBUTION. Riverine and non-flooded forest in Guyana and central Brazilian
Amazonia. Flowering December to May.
GUYANA. Maguire & Fanshawe23363 fl (A, BR, F, K, MO, NY, RB, US); 23569 fl (A, F, K,
NY, US). BRAZIL. Amazonas: Ducke 661 fl (F, IAN, K, MG 17762, MO, NY, R, US); 2127 fl fr
(COL, IAN, MG 17763, NY, P, R); 2232 fl (IAN, NY, R); Mello & CoelhoINPA 3490 fl (IAN, INPA,
MG, NY); Pranceet al 3718 fl (INPA, NY); 4636 fl (INPA, NY); 4934 fr (INPA, NY). Par,: Egler
1258 fl (MG 4305).

Closest to C. spicata,this species differs in the larger flowers 8.0-9.0 mm long,

the leaves, and in the style which is hairy for three-fourthsits length.

14. Couepia spicata Ducke, Archiv. Inst. Biol. Veg. Rio deJaneiro 2:36. I935.

Medium-sized tree. Leaves oblong-lanceolate, 4.6-9.0 cm long, I.4-2.8 cm

broad, cuneate at base, acuminate at apex, with acumen 6.o-I I.o mm long, glabrous
above, densely reddish-brown-arachnoid beneath; midrib prominent above, lower
portion pubescent; primary veins 9-I2 pairs, prominent on lower surface; petioles
3.0-6.0 mm long, pubescent, shallowly canaliculate. Stipules to 4.0 mm long,
lanceolate, caducous. Inflorescences dense-flowered terminal and axillary spikes, the
rachis with ferrugineous-brown sericeous pubescence. Bracts and bracteoles ovate,
3.0-4.0 mm long, equalling the flowers, persistent. Receptacle sessile, turbinate,
3.0-4.0 mm long, ferrugineous-brown-pubescenton exterior, glabrous within except
for deflexed hairs at throat. Calyx lobes rounded. Petals 5, white, ciliate on margins.
Stamens I4-I8, inserted in a semicircle, with a sterile portion opposite them. Ovary
pilose. Style glabrous except at base. Fruit unknown.
TYPE.DuckeRB 25002, Brazil, Amazonas, fl (holotype, RB; isotypes, IAN, K,
NY, P, U).
DISTRIBUTION. Known only from the type gathering collected in non-flooded
forest near Manaus, Brazil.
This species is distinct from others of this group by its much smaller flowers
(less than 4.0 mm long). It is close to and has been confused with C. habrantha, from
which it also differs in the glabrous style.

Taxonomic Discussion of CouepiamagnoliifoliaSuperspecies

There are nine species of Couepiawith the distinctive feature of large persistent
bracts and bracteoles. Four of these species belong to the superspecies under con-
sideration here, two form the C. bracteosaspecies group, and the other three, C.
eriantha,C. insignisand C. martinii,are taxonomically isolated species.
The C. magnoliifoliaspecies complex is a superspecieswith imperfectly allopatric
components. The difference beteen the species in this group are small but constant
and no change was necessary in the taxonomy of the four species. The differencesare
given briefly below each of the species descriptions.
Couepia 227

D. The Couepia bracteosa species Group (Species I5-I7).

15. Couepia bracteosa Bentham, Jour. Bot. Hooker 2: 215. I840; Hook. f.,
Mart. Fl. Bras. I4:(2): 45, t. 15. I867; Ducke, Arch. Mus. Nac. Rio de
Janeiro 22: 68. 9 0; Ducke, Arq. Serv. Flor. Rio de Janeiro I (): 31. 1939.
Tree to 20.0 m tall, usually smaller. Leaves ovate-elliptic, 8.0-22.0 cm long,
4.0-9.5 cm broad, subcordate at base, short-acuminate at apex, glabrous above, gray
to brown-lanate beneath; primary veins 12-25 pairs, prominent beneath; petioles
8.o-I8.o mm long, sparsely pubescent, shallowly canaliculate. Stipules 1.5-3.5 mm
long, subulate, deciduous. Inflorescences terminal panicles with short silver-gray
pubescence. Bracts ovate, often exceeding the receptacle tube, enclosing small groups
of buds, 7.0-I2.0 mm long, persistent. Receptacle subsessile, cylindrical, 7.0-I2.5
mm long, densely brown-pubescent on exterior, glabrous within except for deflexed
hairs at throat. Calyx lobes acute. Petals 5, white, sparsely ciliate on margins.
Stamens 25-40, connate at base of filaments, inserted in an arc of 270?, with a few
short filamentous staminodes opposite them. Ovary densely villous. Style pubescent.
Fruit globose, 8.0 cm long, 7.0 cm broad; epicarp verrucose or smooth; mesocarp
thick and fleshy; endocarp thin and rather fragile, granular, glabrous within.
TYPE. Schomburgk485, Guyana, fl (holotype, K; isotypes, BM, CGE, GH, L, OXF).
DISTRIBUTION. Primary and secondary forest on non-flooded grollnd in the
Guianas and Amazonian Brazil.
GUYANA. Anderson554 fl (BRG, K); De La Cruz 2114 fl (F, GH, NY, US); 2218 fl (F, GH, NY,
US); Davis 429 (F.D. 2424) fl (BRG, FHO, K, U); Fanshawe405 (F.D. 3141) fl (BRG); i333(F.D.
4069) fr (BRG, K); Irwin 8 fl (US); Jenman3952 fl (BRG, K); 4232 fl (K, NY); Persaud141 fr (F, K,
NY); Tutin 289 fr (K, U, US); 291 fl (BM, K, U, US). FRENCH GUIANA. For. Serv. i321 st (U);
Melinonsn fl (P). BRAZIL. Amazonas: Byron& Elias 67-34 fr (INPA, NY); ChagasINPA 865 fl (INPA,
NY); 1956fr (IAN, INPA, NY); 2137 fr (INPA); Ducke 184 fl fr (A F, IAN, K, MG, MO, NY, R, US);
662 fl (F, MG, MO, NY, R, US); 2213 fl (IAN, NY); RB 23589 fl (RB); RB 35563 fl (RB); Froes
26242 fr (IAN, LE); 26303 fl (IAN); Froes & Addison29048 fl (IAN); Guedes225 fl (IAN, NY); INPA
6065 fr (NY); 8078 fl (IAN); Pranceet al 2212 fr (INPA, NY); 3077 fl (INPA, NY); 4706 fr (INPA,
NY); Rodrigues957 fl (INPA 7193, NY); Rodrigues& Chagas1715 fl (IAN, INPA 8078, MG); 2390 fl
(INPA 8804, NY); Rodrigues& Loureiro7170 fl (INPA 15834, NY); Rodrigues,Pires & Silva 157 fl
(IAN); Silva 488 fr (IAN, NY); Spruce1496 fl (BM, F, GH, GOET, LD, LE, NY, OXF, P); 2003 fl
(BM, F, GOET, K, LD, LE, NY, OXF, P). Para: DuckeMG 15438 fl fr (BM, MG, P, RB, US); MG
16367 fl (BM, MG, P, R, RB, US); RB 19756 fl (K, US); RB 24990 fl (K, P, RB, US); KuhlmannRB
17967 fl (K, RB, U, US); Prance,Pennington& Silva 1579 fl (F, NY); Silva 59733 fl (F, GH, NY). Terr.
Rond6nia: Pranceet al 8796 fl (INPA, NY); 8793 fr (INPA, NY). 'Guanabara: Glaziou 13799 fl (K, P).'
BOLIVIA. Pando: Pranceet al 8758 fl (INPA, NY).

LOCALNAMESAND USES.Guyana: Aruadan (Arawak). Brazil: Marirana, Pajurd,

Pajurd de mata. Widely cultivated in the Amazon valley for its edible fruit.
Two varieties of this species have been described, based on variation in leaf size:
var grandifolia R. Ben., Bull. Mus. Hist. Nat. Paris 29: 597. 1923.
var minor Ducke, Arch. Jard. Bot. Rio de Janeiro 5: I 17. I930.
Since there is much variation and a complete gradation in leaf size, it is not
possible to maintain these varieties. At a later date Ducke himself (I939) stated that
var minor was based on a single specimen with smaller dimensions, and should not be
maintained as a separate taxon.

I6. Couepia subcordata Bentham ex Hooker f., Mart. Fl. Bras. 14(2): 46,
t. I6. I867; Macbride, Fl. Peru Io82. I938.
CouepiaamazonicaFritsch, Ann. Naturh. Mus. Wien 5: 2. I890. Type. Poeppig 2814, Brazil,
Amazonas, fl (holotype, W; isotypes, F, GOET, K, L, LE, NY, P).
MoquileainaequalisPoeppig ex Fritsch, Ann. Naturh. Mus. Wien 5: I2. 1890, nom nud, in syn.
228 Flora Neotropica

Medium-sized tree. Leaves oblong-elliptic, I2.0-I7.0 cm long, 5.5-7.5 cm broad,

membraneous, cordate to rounded at base, acuminate at apex, with acumen 3.0-
8.0 mm long, glabrous above, with short, thick, silver-gray to brown-arachnoid
pubescence beneath; primary veins I3-I7 pairs, prominent beneath; petioles 6.0-
7.0 mm long, pubescent, terete, eglandular. Stipules ca 2 mm long, deciduous.
Inflorescences terminal and axillary panicles, the rachis and branches with short
dense light brown pubescence. Bracts and bracteoles ovate, 6.o-I I.o mm long, often
as long as the receptacle tube, persistent up to early flowering. Receptacle subcylin-
drical, 9.0-I2.0 mm long, with short dense pubescence on exterior, glabrous within
except for deflexed hairs at throat; pedicels I.0-2.0 mm long. Calyx lobes acute.
Petals 5, white, ciliate on margins. Stamens ca 35, inserted in a complete circle. Ovary
villous. Style pubescent for two thirds its length. Fruit ellipsoid, 4.5-9.0 cm long,
3.0-5.0 cm broad; epicarp smooth, glabrous; mesocarp thick and fleshy; endocarp
thin, fragile, fibrous, glabrous within.
TYPE. Spruce1423, Brazil, Amazonas, fl (holotype, K; isotypes, CGE, LE, M,
NY, OXF, P).
DISTRIBUTION. Forest in central Amazonian Brazil.
BRAZIL. Amazonas: ChagasINPA 1496 fl (IAN, INPA, MG, NY); Ducke I98 fl fr (A, F, IAN,
K, MG, MO, NY, R, US); KuhlmannRB 17966 fl (K, P, RB, U, US); RB 17987 fr (RB, U, US);
Pranceet al 8z46 fr (INPA, NY); SchwackeIII-209 fl (R); 111-364 fl (GOET). Pari: DuckeRB 15120 fl
cult (RB).

LOCAL NAMES AND USES. Marirana, Umarirana. Cultivated in the vicinity of

Manaus as shade plant. The wood is used in carpentry and for carving.

17. Couepia belemii Prance, sp nov

Arbor media, ramulisjuvenilibus glabris rugosis. Stipulae caducae (haud visae).

Folia alternata petiolata; petiolo I3.o-20.0 mm longo, glabrescenti, rugoso, leviter
canaliculato; laminae oblongae vel oblongo-ellipticae, I i.o-i8.o mm longae, 4.0
7.0 cm latae, apice in acuminem 4.0-8.0 mm longum contractae, basi rotundatae
vel subcuneatae, superne glabrae, subtus dense lanato-arachnoideo-pubescentes;
costa media supra prominente, glabra, subtus prominente; costis secundariis 18-24
jugis, subtus prominentibus, supra planis. Flores in paniculis terminalibus dispositi,
rachi ramisque ferrugineo-tomentosis.Bracteae bracteolaeque magnae, 7.0-25.0 mm
longae, persistentes, coriaceae, extus tomentosae. Receptaculum subsessile,campanu-
latum, 8.0-I4.0 mm longum, extus dense tomentosum, intus prope faucem et infra
stylum pilis longis deflexis ornatum, versus basim glabrum. Calycis-lobi acuti,
7.0-Io.o mm longi, extus tomentosi. Petala 5, alba, margine ciliata. Stamina circa
30, in orbem completum insertum. Ovarium ad faucem receptaculi insertum,
lanato-pilosum. Stylus e basi ovarii ortus, tomentosus. Fructus mihi ignoti.
TYPE. Belem & Pinheiro 3146, Brazil, Bahia, Marau, fl (holotype, NY; isotype,
UB).
Known only from the coastal forest of Bahia, Brazil. Flowering
DISTRIBUTION.
in January.
BRAZIL. Bahia: Marau, Belem & Pinheiro3132 fl (NY, UB); 3139 fl (NY, UB).

This species is most closely related to C. bracteosa,from which it differs in the

larger leaves, the absence of striations from the inflorescence and receptacle, the
stamens inserted in a complete circle, and the more acute leaf bases. It is also close to
Couepia 229

C. erianthabut differs in the broader elliptic leaves with dense pubescence on the
lower surface, the smaller flowers with shorter pubescence, and the much longer
canaliculate petioles.

Taxonomic Discussion of the CouepiabracteosaSpecies Group

Two species of this group, C. bracteosaand C. subcordata, are sympatric. Couepia
bracteosais widely distributed, while C. subcordatais restricted to the region around
Manaus, Brazil. Both species are widely cultivated in Brazil for their fruit. C. subcor-
datais distinct from C. bracteosain the submembraneousleaves, in the greater number
of thinner and less prominent primary leaf veins, in the tomentose (not sericeous)
indumentum of the inflorescence, and in the absence of striations on the rachis of the
inflorescence and on the exterior of the receptacle. The third species, C. belemii,is
allopatric, and is from the east coast of Brazil. It is closely related to C. bracteosa
but
differs in the characters listed under the species description.

E. The Couepia caryophylloides Superspecies (Species I8-I9)

I8. Couepia caryophylloides R. Benoist, Bull. Mus. Hist. Nat. Paris 28: 253.
1922; Kleinh. in Pulle, Fl. Suriname 2(I): 454. I939, Prance, Atas do
Simp6sio sobre a Biota Amaz6nica 4: 220-221. I967.

Tree to 25.0 m tall. Leaves oblong-elliptic to oblong-lanceolate, IO.O-2I.ocm

long, 4.0-7.5 cm broad, rounded at base, acuminate at apex, with acumen 4.0-10.0
mm long, glabrous above, densely gray-brown-arachnoid beneath; primary veins
17-25 pairs, prominent beneath; midrib glabrous above; petioles 7.0-9.0 mm long,
canaliculate above, glabrous or sparsely pubescent. Stipules ca 2.0 mm long, cadu-
cous. Inflorescences dense terminal or axillary panicles 4.0-6.0 cm long, the rachis
and branches densely gray-pubescent. Bracts and bracteoles ovate, to 8.o mm long,
early caducous. Receptacle subcylindrical 5.0-7.0 mm long, longitudinally ridged
when dry, with short gray pubescence on exterior, interior lined by dense reflexed
hairs to base; pedicels I.o-3.o mm long. Calyx lobes rounded, ovate, 3.0-4.0 mm
long. Petals 5, white, ciliate on margins. Stamens 22-35, inserted in a semicircle,
with a toothed ridge of staminodes opposite them. Ovary villous. Style pubescent
almost to apex. Fruit round to oval; epicarp smooth, glabrous; mesocarp thin, fleshy;
endocarp granular, fragile, glabrous within.
TYPE. Wachenheim3-I , French Guiana, fl (lectotype, P); 2-274, French Guiana,
st (paratypes, A, BM, K, P, US).
DISTRIBUTION. Primary forest on high ground from Suriname to Terr. Amapa
in Brazil. Flowering June to December.
SURINAME. B. W. 6350 fl (IAN, K, NY); Lindeman3671 st (U); 4048 st (U); 4178 fl fr (NY, U);
6I09 st (NY, U); 6743 st (NY, U); Maguire24060 fl (A, F, K, MO, NY, US); Maguire,Schulz, Soderstrom
& Holmgren54037 fl (F, GH, MO, NY, US); 54368 fl (F, MO, NY, US); Irwin, Prance,Soderstrom&
Holmgren54903 fl (F, NY, US); 55717 fr (F, GH, MO, NY, US); Schulz 7212 fl (NY, U); roo5o fl
(BBS, U); 7799 st (U); Schulz & Elburg 10167 fl (BBS, U); van Donselaar1131 St (U); 1242 st (U);
1247 st (U). FRENCH GUIANA. For. Serv. 7110 st (P, U); 7138 fr (U); Irwin, Pires & Westra48622
fr (F, GH, NY). BRAZIL. Terr. Amapa: Irwin 48695 fl (F, GH, IAN, MO, NY, US).

LOCALNAMES.Suriname: Anauraballi,Anaura, Hoogland-anaura,BruinbladigeFoengoe.

French Guiana: Pirone,Boliquin.
230 Flora Neotropica

I9. Couepia excelsa Ducke, Arch. Jard. Bot. Rio de Janeiro 5: I I6. 1930;
Prance, Atas do Simp6sio sobre a Biota Amaz6nica 4: 220-221. I967.

Tree to 25.0 m tall. Leaves elliptic to oblong-lanceolate, I3.0-25.0 cm long,

4.0-7.0 cm broad, rounded to subcordate at base, acuminate at apex, with acumen
4.0-9.0 mm long, sparsely appressed-pubescentabove, becoming glabrous with age,
densely gray-arachnoid beneath; midrib sparsely pubescent above; primary veins
23-28 pairs, prominent on lower surface; petioles 8.0-o1.5 mm long, terete, pubes-
cent. Stipules to 3.0 mm long, caducous. Inflorescences short-branched panicles
or racemes, axillary or terminal, 3.0-6.0 cm long, the rachis and branches
brown tomentellous. Bracts and bracteoles, ovate, to 8.o mm long, caducous. Re-
ceptacle subcampanulate, 9.0 mm long, densely brown-pubescent on exterior, the
interior lined to base by dense reflexed hairs; pedicels 2.0-3.0 mm long. Calyx lobes
rounded, ovate, 4.5-6.o mm long. Petals 5, white, ciliate on margins. Stamens 38-40,
inserted in an arc 270?, sometimes with a short sterile toothed portion of circle
opposite them. Ovary villous. Style pubescent almost to apex. Fruit oval, 6.0-7.0 cm
long, 4.0 cm broad; epicarp smooth, glabrous; endocarp granular, fragile, glabrous
within.
TYPE.DuckeRB I9758, Brazil, Para, fl (holotype, RB; isotypes, IAN, K, P, U).
DISTRIBUTION. Non-flooded forest in eastern Amazonian Brazil. Flowering July
to October.
BRAZIL. Terr. Amapa: Pires, Rodrigues& Irvine50765 fl (F, GH, MG, NY, US); 51248 fl
(F, GH, NY, UB, US). 5I324 fl (F, NY, UB).
Taxonomic Discussion of the Couepiacaryophylloides Superspecies
The two foregoing previously little known species comprise a completely
allopatric superspecies. Recent collections have yielded much new information about
these two species. They are among the few species of Couepiain which the interior of
the receptacle bears hairs right to the base. The only other taxon with this feature and
likely to be confused with this group is C. paraensissubsp paraensis.That subspecies is
clearly distinct from the C. caryophylloides superspecies by its broader, shorter leaves
with fewer primary veins (less than I5). For these and other reasons C. paraensisis
considered part of the C. guianensisspecies complex. C. caryophylloides and C. excelsa
are two of the few species of Couepiathat rank among the large trees of the forest of
Amazonia and the Guianas. Couepiais predominantly a genus of small trees and
shrubs.
The type specimens of C. caryophylloides have oblong-elliptic leaves as have
supplemental specimens from nearby areas. However, recent material from the
Wilhelmina Gebergte, Suriname, has much more oblong-lanceolate leaves, suggesting

Table III
Differences between C. excelsaand C. caryophylloides
Character C. excelsa C. caryophylloides
Flower size 3.0-I 7.o mm 8.0-12.0 mm
Color of pubescence in brown silver-gray
inflorescence and fls.
Stamens more than 40 20-35
Leaf shape oblong-lanceolate oblong, elliptic or
oblong-lanceolate
Indumentum of midrib on sparsely pubescent glabrous
upper surface
Indumentum of petioles usually glabrous rarely pubescent
sparsely pubescent
Petioles terete canaliculate
Couepia 231

those of C. excelsa.In all other respects there is no doubt that this material belongs to
These two species are separated by a number of small characters,
C. caryophylloides.
which are summarized in Table III.

F. The Couepia uiti Superspecies (Species 20-21)

20. Couepia uiti (Martius & Zuccarini) Bentham ex Hooker f., Mart. Fl.
Bras. 14(2): 47. 1867.
Moquileauiti Martius & Zuccarini, Abh. Akad. Munchen i: 390. 1832.
CouepiamartianaHooker f., Mart. Fl. Bras. 14(2): 47. 1867, synon nov. Types. Martiussn, Brazil,
Piaui, fr (syntype M); Gardner1284, Brazil, Piaui, fr (syntype K).
CouepiaparaguariensisHassler, Fedde Repert. Nov. Sp. 7: 375. 1909, synon nov. Type. Fiebrig
i379, Paraguay, fl (G, K).
CouepiadahlgreniiStandley, Publ. Field Mus. Bot. I7: 249. 1937, synon nov. Type. Dahlgren986,
Brazil, Piauf, fl (holotype, F).

Tree to 4.0 m tall or shrub. Leaves orbicular to elliptic, 2.5-5.5 cm long, I.5-
3.5 cm broad, rounded to cordate at base, short-acuminate or apiculate at apex,
with acumen 1.0-3.0 mm long, sparsely tomentose above when young, becoming
glabrous with age, densely gray-arachnoid beneath; midrib pubescent above;
primary veins 5-9 pairs, prominent beneath; petioles 2.0 mm long, densely pubes-
cent, often with two conspicuous glands at lamina base, shallowly canaliculate.
Stipules to 6.o mm long, linear, membraneous. Inflorescences terminal panicles, the
rachis and branches with short gray-brown tomentum. Bracts and bracteoles ovate
to lanceolate, to 8.o mm long, deciduous. Receptacle subcampanulate, 5.o-8.o mm
long, with short light brown pubescence on exterior, glabrous within except for
deflexed hairs at throat; pedicels 3.0-5.0 mm long. Calyx lobes rounded. Petals 5,
white, sparsely pubescent, ciliate on margins. Stamens 30-60, inserted in a complete
circle. Ovary densely pilose. Style pubescent for two thirds its length. Fruit ovoid,
3.0-3.5 cm long, 1.5-2.0 cm broad; epicarp verrucose, glabrous; mesocarp thick,
fleshy; endocarp thin, fragile, of a granular texture, glabrous within.
TYPE.Martiussn, Brazil, Bahia, fl (holotype, M).
Savannas, cerrados, and rocky places beside streams and sandy
DISTRIBUTION.
riverbanks in central Brazil and Paraguay.
BRAZIL. Terr. Rond6nia: Pranceet al 5464 fl (INPA, NY); 6722 fl (INPA, NY); 8693 fr (INPA,
NY). Piaui: Ducke866 fr (MG, RB); Gardner o56 fl (K); 2561 fl (BM, GH, K, OXF); Lisb6a MG 2378
fl (BM, MG, P, RB, US). Pernambuco: Curran231 fr (GH, US). Mato Grosso: Diogo 301 fl (R);
Prance,Pires & Silva 59144 fl (F, NY, US); Sick B34 fl (RB 39293); VelosoI4Ii fl (RB 87401). Goias:
Burchell7665 fl (GH). Bahia: Blanchet2775 fl (BM, BR, F, K, LE, NY, OXF, P); Porto 2450 fl (RB
29752); Zehntner230 fl (NY, RB 6345); 1917 fl (M); 300I fl (M). Alagoas: Chase7808 fr (A, US).
Minas Gerais: Macedo1262 fl (MO, RB, SP, US); 2558 fl (NY, S, US); Glaziou 19139 fl (P). Sao Paulo:
GehrtSP 45851 fl (FHO, SP, US). PARAGUAY. Anisits 2182 fl (S); Palmersn fr (US).

LOCALNAMES.Oiti, Uiti.

21. Couepia cataractae Ducke, Arq. Serv. Flor. Rio de Janeiro (I): 3 I. 1939.

Shrub or small tree. Leaves oblong, 4.o-Io.o cm long, 1.5-5.o cm broad,

rounded to subcordate at base, rounded to obtuse at apex, glabrous above, densely
232 Flora Neotropica

gray-arachnoid beneath, usually becoming glabrous with age; primary veins IO-15
pairs, rather inconspicuous beneath; midrib prominent beneath, glabrous above
except at base; petioles 3.0-5.0 mm long, sparsely pubescent, canaliculate above.
Stipules linear, to 1.5 mm long, membraneous, deciduous. Inflorescences terminal
and axillary panicles, the rachis and branches with short dense gray pubescence.
Bracts and bracteoles 1.5-5.0 mm long, ovate, deciduous. Receptacle 5.0-7.0 mm
long, subcampanulate-turbinate, often slightly curved anteriorly, short gray-pubes-
cent on exterior, glabrous within except for deflexed hairs at throat; pedicels slender,
6.0-I3.0 mm long. Calyx lobes rounded. Petals 5, white, pubescent, ciliate on
margins. Stamens 60-90, inserted in a complete circle, the filaments very far exserted.
Ovary pilose. Style pubescent for three fourths its length. Fruit ovoid, with a verru-
cose epicarp; mesocarp thin; endocarp thin, fragile, granular, sparsely pubescent
within.
TYPE.Ducke RB 15136, Brazil, Para, fl (holotype, RB; isotypes, IAN, K, U,
US).
DISTRIBUTION. Rocky places beside rivers south of the Amazon in Para.
BRAZIL. Pari: Black 49-8029 fl (IAN, NY); Frdes20283 fl (IAN, NY, US); 23419 fr (IAN);
30162 fl (IAN, UB); DuckeMG 16388 fl (MG); OliveiraI787 fl (IAN, UB); Pires3565 fr (IAN); Silva
676 fl (IAN).

Taxonomic Discussion of the Couepiauiti Superspecies

Couepiauiti and C. cataractaeform an allopatric superspecies. According to the
original descriptions the difference between C. uiti, C. martiana,C. paraguariensis,and
C. dahlgreniiare in small variations in leaf, flower and inflorescence sizes, and in the
leaf indumentum and number of stamens. With the material of this complex now
available it was found that there is some uncorrelated variation in the characters
mentioned above, but that it is nevertheless not possible to maintain more than one
species. By virtue of its small leaves and the verrucose epicarp of the fruit C. uiti is
rather distinct from all other species of Couepiaexcept C. cataractae.Couepiauiti has a
more southern distribution than C. cataractae,but both grow in the same type of
habitat. C. cataractaeis distinct from C. uiti in its much larger pedicels and in the
longer, larger leaves with a rounded to obtuse apex. In C. uiti the pedicels do not
exceed 3.0 mm, and the shorter leaves have an acumen I.0-3.0 mm long.

G. The Couepia macrophylla Superspecies (Species 22-24).

22. Couepia macrophylla Spruce ex Hooker f., Mart. Fl. Bras. 14(2): 43,
t. I3. 1867; Macbride, Fl. Peru io8i. 1939.
CouepiaspeciosaPilger, Bot. Jahrb. 37: 539. I906. Type. Weberbauer
4649, Peru, (holotype, B,
lost; photo, F).

Small tree. Leaves oblong to oblong-elliptic, I9.0-22.0 cm long, 6.0-12.0 cm

broad on flowering branches (much larger on sterile branches, up to 40.0 cm long
and 14.0 cm broad), glabrous above, densely rufous-to-gray arachnoid beneath,
rounded to cordate at base, abruptly obtuse to abruptly acuminate at apex, with
acumen 4.0-8.0 mm long; primary veins 26-30 pairs, prominent beneath, slightly
impressed above; petioles thick, 8.o-I5.0 mm long, terete, sparsely pubescent, with
two medial sessile glands. Stipules I5.o-20.o mm long, linear, caducous. Inflores-
Couepia 233

cences dense-flowered racemes, the rachis with short reddish-brown pubescence.

Bracts and bracteoles ovate, to 15.0 mm long, persistent. Receptacle 20.0-25.0 mm
long, cylindrical, with short reddish brown pubescence on exterior, glabrous within
except for deflexed hairs at throat; pedicels 2.0-3.2 mm long, Calyx lobes rounded.
Petals 5, persistent, sparsely ciliate at margins. Stamens numerous (ca 280), inserted
in a complete circle, in many rows. Ovary villous. Style densely pubescent through-
out. Fruit unknown.
TYPE. Spruce 4049, Peru, Loreto, fl (holotype, K; isotype, OXF).
DISTRIBUTION. Non-flooded forest in western and central Amazonia.
ECUADOR. Grubbet al io45 fr (FHO, K, NY). Peru. Loreto: Klug 2089 fl (A, F, GH, K, MO,
NY, US); LI. Williams888 st (F, GH, US). BRAZIL. Amazonas: Krukoff5726fl (A, NY); Rodrigues&
Loureiro5744 st (INPA 14456, NY); Wickhamsn st (K).

23. Couepia krukovii Standley, Publ. Field Mus. Bot. 17: 250. I937.

Small to medium-sized tree. Leaves oblong-oblanceolate, 22.0-33.0 cm long,

6.0-7.5 cm broad, rounded to cordate at base, acuminate at apex, with acumen
7.0-10.0 mm long, glabrous on both surfacesexcept when very young; midrib promi-
nent above, glabrous when mature; primary veins I7-23 pairs, prominent beneath,
slightly impressed above; petioles thick, 3.0-6.0 mm long, glabrous, eglandular,
terete. Stipules linear, to T.5 cm long, caducous. Inflorescences densely-flowered
terminal and axillary spicate racemes, the rachis with short tomentellous brown
pubescence. Bracts and bracteoles 6.o-Io.o mm long, ovate, acute, persistent.
Receptacle subsessile, cylindrical, 2.5 cm long, slightly curved, with short dense
brown pubescence on exterior, glabrous within except for deflexed hairs at throat.
Calyx lobes rounded, i.o cm long. Petals 5, ciliate on margins. Stamens numerous
(ca 255), inserted in a complete circle. Ovary villous. Style pubescent for two thirds
its length. Fruit unknown.
TYPE. Krukoff 6208, Brazil, Amazonas, fl (holotype, F; isotypes, A, BM, BR, K,
MO, NY, U); 6007 fl bud (paratypes, A, BM, BR, F, K, LE, MO, NY, U, US).
DISTRIBUTION. Known only from the type gatherings collected in periodically
flooded riverine forest on the Rio Madeira between Manicore and Humaita.

24. Couepia latifolia Standley, Publ. Field Mus. Bot. 17: 251. I937; Ducke,
Arq. Serv. Flor. Rio deJaneiro I (): 30. 1939.

Tree. Leaves oblong-ovate, I7.o-24.o cm long, 5.0-I2.0 cm broad, glabrous

above, sparsely gray-arachnoid or glabrous beneath, rounded at base, rounded or
with short and blunt acumen at apex; primary veins I6-I9 pairs, prominent beneath,
slightly impressed above; petioles 4.0-9.0 mm long, pubescent, with two medial
glands, terete. Stipules lanceolate, ca 3.0 mm long, subpersistent. Inflorescences
axillary racemes, the rachis tomentellous. Bracts and bracteoles 4.0-8.0 mm long,
ovate, acute, persistent, tomentellous on exterior. Receptacle cylindrical, sessile,
I.5-2.5 cm long, with dense light-brown pubescence on exterior, glabrous within
except for deflexed hairs at throat. Calyx lobes rounded. Petals 5, persistent, ciliate
on margins. Stamens numerous, inserted in a complete circle. Ovary villous. Style
densely pubescent for three fourths its length. Fruit oblong, curved, 4.5 cm long,
234 Flora Neotropica

1.5 cm broad (in single fruit examined); epicarp smooth, glabrous; mesocarp thick,
fleshy; endocarp thin, of a granular texture, fragile, glabrous within.
TYPE.Krukoff6005, Brazil, Amazonas, fl fr (holotype, F; isotypes, A, BM, BR,
K, MO, NY, US).
DISTRIBUTION. Forest in periodically flooded areas. Collected from the basins of
the Rios Purus and Madeira.
BRAZIL, Amazonas: Pranceet al 2239 fl (NY); 7934 fl (INPA, NY).

This species is close to C. macrophylla

but differs in the smaller leaves with fewer
primary veins, and the smaller flowers.

Taxonomic Discussion of the Couepiamacrophylla

Superspecies
The three component species Couepiamacrophylla, C. latifoliaand C. krukoviiform
an imperfectly allopatric superspecies. All three are, at this time, so poorly known
that it is possible only to make preliminary comments. Although at present no
alteration in circumscription is necessary, the three species are very closely related.
As a group they have larger leaves and flowers than any other species of Couepia.

H. The Couepia ovalifolia Species Group (Species 25-26).

25. Couepia ovalifolia (Schott) Bentham, Jour. Bot. Hooker 2: 216. I840;
Hook. f., Mart. Fl. Bras. 14(2): 48. I867.
ovalifoliusSchott in Spreng,. Linn., Syst. Veg. ed. I6. 4(App.): 406. 1827.
Chrysobalanus
CouepiahypoleucaMiquel, Linnaea 19: 438. I847. Type. Claussen2023, Brazil, Guanabara, fl
(holotype, L; isotvpes, C, G, MO, P, U).

Dwarf shrub to medium-sized tree. Leaves ovate, 4.0-I0.5 cm long, 2.5-7.0

cm broad, rounded to cordate at base, short-acuminate at apex, with acumen I.o-
7.0 mm long, glabrous above, densely silver-gray-arachnoidbeneath, sometimes be-
coming glabrous with age; midrib prominent beneath, pubescent above when young;
primary veins 11-15 pairs, prominent beneath, impressed above; petioles 4.0-7.0
(-io.o) mm long, pubescent when young, shallowly canaliculate. Stipules to 2.0 mm
long, caducous. Inflorescences terminal panicles, the rachis and branches with short
light brown pubescence. Bracts and bracteoles oblong-ovate, 3.0-6.5 mm long,
caducous. Receptacle subcampanulate, 4.0-5.0 mm long, densely light-brown-
pubescent on exterior, glabrous within except for deflexed hairs at throat; pedicels
1.5-4.0 mm long. Calyx lobes rounded, 3.0-4.0 mm long. Petals 5, white, ciliate on
margins. Stamens 16-19, inserted in an arc of 240?, with a sterile portion opposite
them. Ovary densely pilose. Style pubescent half its length. Fruit elliptical; epicarp
smooth, glabrous or softly pubescent when young; mesocarp thin, fleshy; endocarp
thin and fragile, granular, glabrous within.
TYPE.Schottsn, Brazil, Guanabara, fl (holotype, W; isotypes, K, NY).
Restingas, littoral forest, sand plains, and sand bars on the coast
DISTRIBUTION.
of Brazil from Pernambuco to Rio de Janeiro. Flowering August to November.
BRAZIL. Pernambuco: A. Lima 55-2256 fl (IPA); Prance48o1 fr (NY). Bahia: Belem & Pinheiro
2837 fl (NY, UB); 2846 fl (NY, UB); 2978 fl (NY, UB); 3084 fl (NY, UB); 3227 fr (NY, UB); 3281 fr
(NY, UB); 3315 fr (NY, UB); Blanchet3387 fl (BM, BR, F, GH, K, LE, NY, P, RB). Minas Gerais:
St. Hilaire 293 fl (P). Espirito Santo: Brade 18344 st (RB 56596). Rio de Janeiro: Alston & Lutz 66 fl
Couepia 235

(R); Brade 12758 fl (RB 29402); MachadoRB 75835 fr (RB); Markgraf3038 fl (RB); RestingaI 772 fl
(NY, R); o040fl (NY, R); Riedel sn fl (NY, US); Segadas-Vianna527 fl (NY, R); Ule 7095 fl (R).
Guanabara: Alston & Lutz 136 fr (BM, R, U); Blitz 671 fl (US); Brade 1107 fl (GH); CapenemaRB
19200 fl (RB); Glaziou 78 fl (P, R); 896 fl (BR, P); 1384 fl (BR, P); 6167 fl (BR,K, P); 13433 fl (K,
LE); 16761 fl (P); Gomessn fl (K); Hans 132 fl (R); 162 fl (R); KuhlmannRB 2025 fl (RB); RB 139760
fl (RB); Lutz 67i fl (R, US); 931 fl (R); Maguire,Maguire& Pires 44577 fl fr (NY, US); MuseuNacional
36006 fl (R); Occhioni1157 fl (NY, RB); Pereira3530 st (RB); 4033 fl (RB); 4281 fl (RB); Santos5206
fl (R); 5231 fl (R); Segadas-Vianna2731 fl (NY, R); 3796 fr (R); 3813 fr (R).

Later authors have cited this species as C. ovatifolia,but as ovalifoliawas used in

the original description (in Chrysobalanus),this name must be regarded as valid.

26. Couepia schottii Fritsch, Ann. Naturh. Mus. Wien 5: I3. I89o.
Schott in Spreng., Linn., Syst. Veg. ed. I6. 4(App.): 407. 1827.
macrophyllus
Chrysobalanus

Tree. Leaves oblong, Io.o-I8.o cm long, 4.0-7.0 cm broad, subcordate at base,

rounded to short-acuminate at apex, with acumen I.0-3.0 mm long, glabrous above,
densely gray-brown-arachnoid beneath; primary veins I6-20 pairs, prominent
beneath, impressed above; petioles I2.0-I5.0 mm long, sparsely pubescent, shallowly
canaliculate. Stipules ca 2.0 mm long, caducous. Inflorescences terminal panicles,
the rachis and branches with short light brown pubescence. Bracts and bracteoles
ovate, acute, to 4.o mm long, caducous. Receptacle subcampanulate 5.o-6.o mm
long, with short light brown pubescence on exterior, glabrous within except for de-
flexed hairs at throat; pedicels 2.0 mm long. Calyx lobes rounded. Petals 5, ciliate
on margins. Stamens I5-I6, inserted in a semicircle, with a few small filamentous
staminodes opposite them. Ovary densely villous. Style pubescent for half its length.
Fruit spherical, ca 3.5 cm in diam.; epicarp smooth, glabrous; mesocarp thin, fleshy;
endocarp thin, granular, fragile, glabrous within.
TYPE. Schottsn, Brazil, Rio de Janeiro or Guanabara, fl (holotype, W; isotype,
NY).
DISTRIBUTION.Coastal restingas from Bahia to Rio de Janeiro. Flowering
August to November.
BRAZIL. Bahia: Bondar133 fl (RB); 2355 fl (SP); 2511 st (F, SP); 3012 fl (F); Duarte5980 fl
(RB); 8005 fl (RB). Rio de Janeiro: Glaziou & Schwacke5696 fl (R); Schwacke4242 fl (GOET). Guana-
bara: Boechat2I fl (R); Glaziou 2126 fl (GH, K, NY, P); Soti 12 fl (R); Ule 7096 fl (R). Rio de Janeiro
or Guanabara: Glaziou 7876 fl (F, K, P, US).

LOCAL NAME. Oiti boi.

Hooker cited this species in synonymy under Couepiaovalifoliain his account for
Flora Brasiliensis.Fritsch examined the two Schott specimens and realized that the
two species originally described as Chrysobalanusovalifoliusand Chrysobalanus macro-
phylluswere separate species. He created the new name Couepiaschottiifor the latter
because of the priority of C. macrophylla
Spruce ex Hooker f.

Taxonomic Discussion of the CouepiaovalifoliaSpecies Group

These two species form a sympatric species group. They grow together in the
restricted habitat of the coastal savannas and sand bars (restingas) of southern Brazil.
Couepiaovalifoliaand C. schottiiare closely related, but are separable by a number of
small features which are summarized in Table IV.
236 Flora Neotropica

Table IV

Difference between C. ovalifoliaand C. schottii

Character C. schottii C. ovalifolia

Leaf shape oblong ovate to oblong-ovate

length io.o-i8.o cm 4.0-10.0 cm
width 4.0-7.0 cm 3.0-3.5 cm
No. of primary veins 16-20 11-15
Petiole length 10.0-I5.0 mm 4.0-7.0 mm

I. Taxonomically Isolated Species (Species 27-55).

27. Couepia grandiflora (Martius & Zuccarini) Bentham ex Hooker f.,

Mart. Fl. Bras. I4(2): 46. 1867; Warming, Dansk. Vid. Selsk. Skrift. VI.
6: I90. 1892.
MoquileagrandifloraMartius & Zuccarini, Abh. Akad. Munchen I: 388. 1832.
Couepiaformosana Taubert, Bot. Jahrb. 21: 429. 1896. Type. Ule 2881, Brazil, fl (holotype, B, lost;
isotype, HBG).
CouepiasuberosaPilger, Notizbl. Bot. Gart. Berlin 8: 538. 1923. Type. Riedel504, Brazil, Minas
Gerais, fl (holotype, B, lost; isotype, OXF).

Tree to 8.o m tall, the branches pubescent when young, the older branches often
with a conspicuous thick cork covering ca 4.0 mm thick. Leaves oblong to ovate,
7.5-I8.o cm long, 3.5-9.0 cm broad, cordate to cuneate at base, rounded to bluntly
acuminate at apex, glabrous above, or with sparse ferrugineous pubescence when
young, shortly dense-arachnoid beneath; primary veins io0-6 pairs, prominent
beneath, prominulous above; midrib prominent on both surfaces; petioles 3.0-I3.0
mm long, pubescent, terete. Stipules to 2.0 mm long, subulate, caducous. Inflores-
cences terminal and axillary panicles, the rachis and branches with dense yellow-
brown pubescence. Bracts and bracteoles 1.5-5.0 mm long, ovate, membraneous,
caducous. Receptacle campanulate, 5.0-11.5 cm long, densely cinereous-pubescent,
glabrous within except for deflexed hairs at throat; pedicels 2.0-5.0 mm long.
Calyx lobes rounded. Petals 5, white, pubescent, ciliate on margins, caducous.
Stamens 50-I25, inserted in a complete circle. Ovary densely villous. Style pubescent
for most of its length. Fruit oblong; epicarp smooth, glabrous, drying black; pericarp
thick, fleshy, glabrous within.
TYPES.Martius sn, Brazil, Goias, fl (lectotype, M); Martiussn, Brazil, Minas
Gerais, fl (paratype, M).
DISTRIBUTION. Cerrados and savannas of the Planalto of central Brazil. Flowering
July to December.
BRAZIL. Maranhao: LisboaMG 2494 fl (BM, MG, RB). Piaui: Gardner2563 fl (BM, F, GH, K,
OXF, P, US). Mato Grosso: Hoehne27716 fl (R); Irwin & Soderstrom 6423 fl (NY); 6495 fl (NY); 66r5
fl (NY); 6711 fl (NY); Maguire, Pires, Maguire & Silva 56300 fl (NY); Malme 2356 fl (R); Oliveira
RB 67975 fl (RB); Pires 56825 fl (UB); 56948 fl (UB); Sidney 1261 fl (UB). Goias: Glaziou 20117 fl
(K, R); Macedo3905 fl (RB); Riedel 702 fl (K, NY); Ule 965 fl (P). Dist. Federal: Andrade351 fl (R);
424 fl (R); Cobra & Oliveira 174 fl (P, UB); Duarte & Santos99 fl (RB 26920, NY); Heringer8799 fl
(NY); 9288 fl (HB, NY, UB); 10529 fl (UB); Irwin et al 7968 fl (NY); Lane SP 69973 fl (SP); Macedo
3905 fl (RB); Prance & Silva 59043 fl (F, NY, RB). Minas Gerais: Barreto6783 fr (K); 6784 fl (F, K,
RB); 7289 fl (R, RB); 8732 fl (F); 9227 fl (R); Claussensn fl (BM, K); 256 fl (P); Duarte3269 fl (NY,
RB 71914); 9255 fl (RB 130313); Ferreira& Marquesi9 fl (SP); Gardner4538 fl (BM, K, NY); Heringer
4087 fl (NY, UB); 5368 fl (NY, UB); Irwin & Soderstrom 5548 fl (NY); Macedo715 fl (MO, SP, US);
.Netto243 fl (R); Warmingsn fl (C, F); L. O. Williams & Assis 7597 fl (BR, F, GH, K, MO, NY, R, RB,
SP, US); Magalhaes 6o05 fl (P); RegnellIII:152 fl (BR, US); St. Hilaire 982 fl (P); 1714 fl (P). Sao
Couepia 237

Paulo: Campos62 fl (SP); Felippe 52 fl (SP, US); Hoehne& GehrtSP 36764 fl (FHO, LE, SP, US);
Mattos & Mattos 8362 fl (NY); Pinho 26 fl (SP); Schwebeliri9 fl (R). Parana: Hatschbach2819 fl (US).
Without locality: Riedel zo28 fl (US).

LOCALNAME.Pitomba de leite.
The only difference allegedly separating C. suberosafrom C. grandiflora,is the
thick corky bark of the mature stems of the former. This feature is not at all constant
as the thickness of the bark is variable, and since this character is not correlated with
any other characters,C. suberosais reduced to synonymy. The thick bark is apparently
a common adaption for the annual fires of the dry season in the Brazilian cerrados.

28. Couepia elata Ducke, Arch. Inst. Biol. Veg. Rio de Janeiro 2: 35. 1935.

Large tree. Leaves ovate 4.0-7.0 cm long, 2.5-3.0 cm broad, obtuse to subcuneate
at base, caudately acuminate at apex, with acumen 6.0-9.0 mm long, glabrous above,
prominently reticulate beneath with silver-gray pubescence between reticulations;
midrib impressed above, pubescent on lower portion; primary veins 6-8 pairs,
prominent beneath; petioles 4.0-6.0 mm long, shortly pubescent. Stipules minute.
Inflorescences terminal and axillary few-flowered lax racemes, with short gray-
pubescence. Bracts and bracteoles minute, caducous. Receptacle obconical-turbinate,
4.0-5.0 mm long, sparsely pubescent on exterior, glabrous within except for de-
flexed hairs at throat; pedicels 2.5-3.5 mm long. Calyx lobes acute. Petals 5, white,
glabrous on margins, deciduous. Stamens 35-40, inserted in half a circle, with a row
of short-toothed staminodes opposite them. Ovary densely villous. Style glabrous
except at base. Fruit globose, 5.0-6.0 cm diameter; epicarp densely verrucose;
mesocarp thick; endocarp fragile, thick, granular.
TYPE.DuckeRB 2500o, Brazil, Amazonas, fl fr (holotype, RB; isotypes, K, P, U).
DISTRIBUTION. Non-flooded forest in the lower Rio Negro region of Brazil.
Flowering March to June.
BRAZIL. Amazonas: Ducke921 fl (F, GH, IAN, K, MG, MO, NY, R, US, all from the type tree);
INPA 9040 fl (INPA); Loureiro& CoelhoINPA 15532 fl (INPA, NY); Oliveira2979 st (IAN, UB);
Rodrigues& Loureiro5829 fl (INPA 1478I, NY); Rodrigues& Osmarino6962 fl (INPA 15535); 7638 fr
(INPA I6942, NY).

In the original description Ducke mentioned that this species has a very charac-
teristic bark, which in fact arises from the fact that the old bark exfoliates in scales
which at first remain attached, giving the trunk a shaggy appearance. Couepiaelata
is clearly distinct from the other species of Couepiawith prominently reticulate leaves
by the short sparse silver-gray pubescence of the inflorescences and flowers, the
minute caducous bracts and stipules, and the turbinate receptacle.

29. Couepia racemosa Bentham ex Hooker f., Mart. Fl. Bras. 14(2): 43, t. 14.
1867; Macbride, Fl. Peru. io8i. 1938; Ducke, Arq. Serv. Flor. Rio de
Janeiro i(I): 29. I939.
CouepiacalophlebiaStandley, Publ. Field Mus. Bot. 17: 248. I937. Type. Krukoff4913, Brazil,
Amazonas, fl (holotype, F; isotypes, A, BM, F, K, M, MO, NY, RB, U, US).

Tree to 30.0 m tall. Leaves elliptic to oblong, 5.0-I8.5 cm long, 2.0-7.5 cm

broad, rounded to cuneate at base, acuminate at apex, with acumen 4.5-20.0 mm
238 Flora Neotropica

long, glabrous above, prominently reticulate beneath, or with stomatal cavities, with
short appressed gray pubescence; midrib prominent beneath, slightly impressed
above; primary veins 8-I2 pairs, prominent beneath, slightly impressed above;
petioles 8.0-I2.0 mm long, with two sessile glands near to lamina base. Stipules ca
1.5 mm long, caducous. Inflorescenceslittle branched densely-floweredpanicles, the
rachis and branches with short brown tomentellous pubescence. Bracts and bracteoles
ovate to oblong, 2.0-6.0 mm long, caducous. Receptacle cylindrical-turbinate, 5-0-
7.5 mm long, with a short brown pubescence on exterior, glabrous within except for
deflexed hairs at throat. Calyx lobes rounded to acute. Petals 5, white, exceeding
calyx lobes, glabrous on margins. Stamens 24-30, unilateral, with a smooth glabrous
untoothed portion of the circle opposite them. Ovary densely villous. Style pubescent
at base only. Fruit elliptic, pointed at apex, 3.0-4.5 cm long, 2.0-3.5 cm broad;
epicarp verrucose;pericarp undifferentiated,rather fragile, sparselypubescent within.
TYPE.SpruceI776, Brazil, Amazonas, fl (lectotype, K; isolectotypes, OXF, P);
2322, Brazil, Amazonas, fl fr (paratypes, BM, CGE, F, GH, GOET, K, LD, M,
NY, OXF).
Non-flooded forest or on periodically flooded ground, riverine forest
DISTRIBUTION.
and river banks, and secondary forest in central and western Amazonia. Flowering
February to August.
COLOMBIA. Vaupes: Schultes& Cabrera15291 fr (F, NY, US). BRAZIL. Amazonas: Chagas
INPA 983 fl (IAN, INPA, MG, NY); CoelhoINPA 2907 fr (INPA, NY); INPA 3953 fl (INPA, MG, NY);
INPA 20880 fr (INPA); Ducke407 fl (A, F, K, MO, NY, R, US); 924 fl (F, IAN, MG, MO, NY, R, US);
2142 fl (IAN, MG, NY, R); 2208 fr (IAN, NY, R); MG 10461 fr (MG); MG 11658 fl (MG, P); MG
15684 fr (BM, MG); RB 15133 fl (K, RB, U); RB 24991 fl (K, P, RB, U, US); RB 35554 fl (K, RB,
US); RB 35558fi (K, RB, U, US); Frdes21112 fl (IAN, K, NY, US); INPA 2163 fr (NY); 3228fr (NY);
6092 fr (INPA); GoeldiMG 3933 fl (RB 15130); Krukof8843 fr (BR, F, K, LE, NY, S); Mello INPA 2163
fr (INPA); Mello & CoelhoINPA 393I fl (IAN, INPA, NY); Pranceet al 4755 fl (INPA, NY); Rodrigues
1162 st (INPA 7477); 2695 fl (INPA 9202, NY); Rodrigues& Chagas2739fl (INPA 9282, NY); 4051 fr
(INPA Io616, NY); Rodrigues& Lima 2246 fl (INPA 9593); Schultes& Lopez 10183 fl (IAN, NY, US);
Ule 6I49 fl (K, MG 6022). Pard: DuckeMG 8536B fl (BM, MG, P, RB, US); MG 9125 fr (BM, MG,
RB); MG 17055 fl (BM, MG, P, RB, US) RB 1658 fr (US); MG 15844 fr (BM, MG, RB).

LOCALNAMES.Venezuela: Makara (Arekuna). Brazil: Jahaun (Maku), Tucriba.

This species is distinct from others with prominently reticulate leaves by the
paniculate inflorescence, the minute caducous bracts and stipules, and the fruit with
a glabrous verrucose epicarp.
C. racemosavar reticulataPilger is not C. canomensisas stated by Ducke (Arq. Serv.
Flor. Rio de Janeiro I(I): 29. I939). An examination of the type of var reticulata
(Ule 6149) shows that it falls well within the variation of this species in which there is
much variation in leaf shape and size. Consequently var reticulatais not maintained
here.

30. Couepia martinii Prance, sp nov Fig 38.

Arbor vel frutex?, ramulis juvenilibus glabris lenticellatis. Folia alternata,

petiolata; petiolo IO.0-I2.0 mm longo; lamina elliptica, I4.0-25.0 cm longa, 6.0-
10.5 cm lata, basi plus minusve cuneata, apive in acuminem brevem ad 8.0 mm
longum contracta, superne glabra, subtus sparse araneoso-griseo-tomentosa; costis
secundariis 10-12 jugis, subtus prominentibus. Stipulae parvae, caducae. Flores in
picem axillarem densiflorem ubique sparse ferrugineo-tomentosam dispositi.
 Couepia 2319
39

flower section, x 5; E, bracteole, X2.5.

240 Flora Neotropica

Bracteae bracteolaeque ad I7.0 mm longae, ovatae, acutae. Receptaculum cylindri-

cum, 20.0-25.0 mm longum, extus brunneo-tomentosum, intus prope faucem pilis
longis deflexis, versus basim glabrum. Calycis-lobi acuti. Petala 5, decidua, glabra,
praeter margines ciliolata. Stamina plus minusve 65, in orbem completum disposita.
Ovarium villosum. Stylus e basi ovarii ortus, dense pilosus. Fructus mihi ignoti.
TYPE. Martinsn, French Guiana, Cayenne, fl (holotype, NY; isotypes, BM, F,
K).
Known only from the type collection, which unfortunately lacks
DISTRIBUTION.
detailed field notes.
The material of this species has been erroneously filed in herbaria under the
name Couepiaparillo, but it does not resemble that species. Although C. martiniiis one
of the species of Couepiawith large persistent bracteoles, it is nevertheless distinct. The
inflorescence is completely different from the paniculate one of the C. bracteosa
species group which it resembles in flower structure. It may be distinguished from
other species with spiciform inflorescences by the long tubular receptacle bearing
large bracteoles at the base, as well as by the shape and size of the leaves.
The labels of some of the specimens examined are marked "collected by H. F.
Talbot and deposited in the Sims Herbarium." This is almost certainly the same
Martin collection from the pirated Martin specimens described by Steam, Bull.
Jard. Bot. Brux. 27: 243. I957.

3I. Couepia bondarii Prance, sp nov Fig 39 G-L.

Arbor?, ramulis juvenilibus glabris. Stipulae lanceolatae, membranaceae, ad

30mm longae, caducae. Folia alternata petiolata; petiolo 6.o-Io.omm longo,
tomentello, tereti; laminae oblongo-lanceolatae, I5.o-25. cm longae, 3.5-7.0 cm
latae, apice in acuminem plus minusve 5.0 mm longum contractae, basi subcuneatae,
superne glabrae, subtus dense rufo-brunneo-arachnoideo-pubescentes;costa media
supra glabra, leviter prominente; costis secundariis 17-25 jugis, supra planis, subtus
prominentibus. Flores 20.0-30.0 mm longi, in racemis fasciculatis terminalibus
axillaribusque dispositi, rachi ferrugineo-tomentelli.Bracteae bracteolaeque magnae,
ovatae, persistentes,ad 25.0 mm longae, extus dense brunneo-puberulae, intus sparse
puberulae. Receptaculum sessile, tubulosum, I2.o-I5.0 mm longum, extus rufo-
tomentellum, intus prope faucem pilis longis deflexis ornatum, versus basim glabrum.
Calycis-lobi acuti, rufo-tomentelli, Io.o-I6.o mm longi. Petala 5, extus sericea.
Stamina circa 65, in orbem completum disposita, basi ad 1.5 mm connata. Ovarium
ad faucemreceptaculiinsertum,pilosum. Styluse basi ovarii ortus, tomentosum. Fructus
mihi ignoti.
TYPE.Bondari863, Brazil, Bahia, without precise locality, fl (holotype, SP).
Known only from two collections, one from the State of Bahia,
DISTRIBUTION.
Brazil, and the other without any locality.
BRAZIL.Withoutlocality: Riedel68i fl (NY, US).

This species is most closely related to C. insignis and C. martinii, but is distinct
from both. It differs from C. insignisin the leaf veins which do not anastomose, the
much narrower leaves usually with subcuneate bases, the shorter petioles, the large
flowers, and the tubular receptacle. It differs from C. martiniiin the narrower oblong-
lanceolate leaves, the larger number of primary leaf veins, the much thicker rufous-
brown pubescence of the leaf underside, the denser pubescence of the inflorescenca
and flowers, and the shorter thicker receptacle tube.
Couepia 241

-.

i)

FIG 39. Species of Hirtella and Couepia.A-E, Hirtella bahiensis(Belem et al 2756, Duarte 8020);
A, habit, x 0.5; B, flower, x 4; C, flower section, x 4; D, young fruit and bract glands, X 0.5; E, petal,
x 5; F, ovary, x 12. G-L, Couepiabondarii(Bondar1863); G, flower, x i; H, flower section, X o.5; J,
ovary, x 8; K, petal, X 2; L, habit, x 0.5.
242 Flora Neotropica

This species is named after Gregorio Bondar whose collections from Bahia are
rich in novelties.

32. Couepia insignis Fritsch, Ann. Naturh. Mus. Wien 5: I I. 890.

Tree. Leaves oblong, I5.0-24.0 cm long, 8.0-9.5 cm broad, rounded at apex,

rounded to cordate at base, glabrous above, densely gray-arachnoidbeneath; primary
veins 21-25 pairs, impressed above, prominent beneath, arcuate and anastomosing
near the lamina margin; petioles 8.0- 1.0 mm lng, sparsely pubescent, terete.
Stipules linear, to I5.0 mm long, membraneous, caducous. Inflorescences few-
flowered racemes, the rachis with short rufous pubesence. Bracts and bracteoles
persistent, io.o-28. mm long, oblong, acuminate, membraneous. Receptacle
subcylindrical, 8.0-9.0 mm long, with short brown pubescence on exterior, glabrous
within except for deflexed hairs at throat; pedicels I.0-2.0 mm long. Calyx lobes
oblong-lanceolate, Io.o-I2.0 mm long, acute. Petals 5, ciliate on margins. Stamens
numerous (over 50). Ovary villous. Style pubescent for three fourths its length. Fruit
unknown.
TYPE. Blanchet32o0, Brazil, Bahia, fl (holotype, W; isotype, G).
DISTRIBUTION. Collected only in Bahia, but precise localities as yet unknown.
BRAZIL. Bahia: Blanchet215 st (P); Bondarsn st (F); Luschnathsn fl (BR, US). Without locality:
Riedelsn st (US).

LOCALNAME.Oiticoranga.
This species is nearest C. macrophylla, but differs in the shorter subcylindrical
receptacle, the acute calyx lobes, the ciliate margins, the smaller leaves, and in the
primary veins anastomosing at the lamina margin. Couepiainsignisdiffers from the
C. bracteosaspecies group in its larger bracteoles, in the racemose inflorescence, and
in the arcuately anastomosing leaf nerves.

33. Couepia recurva Spruce ex Prance, sp nov Fig 40 A-H.

Arbor?, ramulisjuvenilibus rufo-pubescentibuslenticellatis. Stipulae ad 3.0 mm

longae, caducae. Folia alternata, breviter petiolata, petiolo circa 2.0 mm longo
hirsuto; laminae oblongo-ovatae, 5.5-7.5 cm longae, 3.0-4.0 cm latae, margine
integrae, apice in acuminem ad 9.0 mm longum contractae, basi rotundatae, utrin-
que sparse hirsutae; costis secundariis 10-12 jugis, subtus prominentibus. Flores in
racemis axillaribus breviter rufo-brunneo-hirsutisdispositi. Bracteae bracteolaeque
caducae (haud visae). Receptaculum cylindricum, 20.0-25.0 mm longum, extus
rufo-brunneo-hirsutum; intus dense tomentosum; pedicelli 5.0-7.0 mm longi.
Calycis-lobi 5, rotundati. Petala 5, decidua, obtusa, margine glabra. Stamina circa
17, unilateralia, filamentis hirsutis. Ovarium ad faucem receptaculi insertum, dense
pubescens. Stylus e basi ovarii ortus, pilosus. Fructus mihi ignoti.
TYPE.Spruce5067, Ecuador, 'In Andibus Ecuadorensibus', fl (holotype, NY;
isotypes, BM, CGE, K, LE, OXF).
Known only from the type collection.
DISTRIBUTION.
This species is distinct from all other species of Couepiaby the hirsute filaments
and leaves. The receptacle is filled with hairs right to the base, but C. recurvais very
unlike the other species of Couepiawith this character. It does, in fact, resemble many
Couepia 243

v'',~~~~~~~~~;:?'.rpy

B
S:':

H~~~~~~~~~~
t~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~;
M . i

'
'4~~~~~~~~~~~

M~~~~~~~~~~~~~~~~~~~~~~~~~

A-H, C. recurva
FIG 40. Speciesof Couepia. (Spruce
5067); A, habit, x o-5; B, leaf surface(upper
surface above; lower surface below), x 2.5; C, flower, x 2,5; D, flower section, X 2.5; E, petal, X 2.5; F,
ovary and style, x 4; G, ovary and style section, x IO; H, diagram of filament base, x 2.5. J-0, C.
venosa(Hoehne17I97); J, habit, xo.-; K, flower, x2.5; L, flower section, X 2.5; M, petal, x2.5; N,
diagram of filament base, X 2.5; 0, ovary and style, x 4.
244 Flora Neotropica

species of Hirtellain most floral characters, but must be placed in Couepiabecause of

the number of stamens.

34. Couepia obovata Ducke, Arch. Inst. Biol. Veg. Rio de Janeiro 2: 35. 1935.

Small to medium-sized tree. Leaves obovate, 4.0-10.0 cm long, 2.0-5.0 cm

broad, subcuneate at base, bluntly acuminate at apex, with acumen 2.0-5.0 mm
long, glabrous above, with a sparse covering of appressed gray hairs beneath;
primary veins 8-IO pairs, prominent beneath; petioles 3.0-7.0 mm long. Stipules
1.0-2.0 mm long, lanceolate, caducous, Inflorescencesterminal and axillary racemes,
3.0-6.0 cm long, the rachis with sparse silver-gray pubescence. Bracts and bracteoles
1.5-3.5 mm long, ovate, persistent. Receptacle cylindrical, 7.0-9.0 mm long, curved,
with a few short appressed hairs on exterior, glabrous within except for deflexed
hairs at throat, subsessile. Calyx lobes rounded. Petals 5, white, ciliate on margins.
Stamens 16-21, inserted in a complete circle. Ovary lanate. Style giabrous except at
base. Fruit ovoid, 2.0-3.0 cm long, 1.5-2.5 cm broad; epicarp smooth, glabrous;
endocarp thick, hard, of a granular texture, glabrous within.
TYPE. DuckeRB 24996, Brazil, Amazonas, fl fr (holotype, RB; isotypes, IAN,
K, P, U, US); RB 24995 fl (paratypes, INPA, K, P, U).
DISTRIBUTION. Non-flooded forest in the Guianas and central and Colombian
Amazonia.
COLOMBIA. Vaupes: Schultes& Cabrera15662 fl (GH, NY, US). SURINAME. Irwin, Prance,
Soderstrom& Holmgren54802 fl (F, MO, NY, US). FRENCH GUIANA. Martin sn fl (K). BRAZIL.
Amazonas: Black 47-1632 fr (IAN, NY); ByronINPA 5566 fr (IAN, NY); CoelhoINPA 4137 fr (INPA,
NY); Ducke 406 fl (A, F, INPA, K, MO, NY, R, US); RB 35561 fl fr (K, RB); Rodrigues& Loureiro
5830 fl (INPA 14782, NY); 5832 fl (INPA 14784, NY); Wilson5566 fr (INPA). Para: Oliveira3735 fr
(IAN, NY).

LOCAL NAME.Colombia: Aseepee(Makuna).

35. Couepia williamsii Macbride, Candollea 5: 366. I934; Macbride, Fl.

Peru. 1082. 1938.

Small tree. Leaves oblong to oblong-lanceolate, 9.0-I7.0 cm long, 2.5-5.0 cm

broad, cuneate to subcuneate at base, abruptly acuminate at apex, with acumen
i2.0-22.0 mm long, glabrous above, densely appressed-lanate beneath; primary
veins I2-15 pairs, prominent beneath; midrib prominent above; petioles 5.0-7.0 mm
long, with a few stiff erect hairs, shallowly canaliculate above, with two inconspi-
cuous glands at lamina base. Stipules to 3.0 mm long, linear, caducous. Inflores-
cences terminal and axillary few-flowered racemes, the rachis sparsely puberulous.
Bracts and bracteoles to 3.0 mm long, early caducous. Receptacle slender and
tubular, I4.0-I9.0 mm long, curved, glabrous on exterior or with a few appressed
hairs, glabrous within except for deflexed hairs at throat; pedicels 3.0-5.0 mm long.
Calyx lobes rounded. Petals 5, white, ciliate on margins. Stamens ca 70, inserted in
a complete circle. Ovary villous. Style pubescent for two thirds its length. Fruit
globose, ca 6.5 mm in diameter; epicarp glabrous, verrucose; mesocarp thick, fleshy;
endocarp thin, fragile, granular, glabrous within.
TYPE. LI. Williams 2975, Peru, Loreto, fl (holotype, F).
DISTRIBUTION. Forest on non-flooded and periodically flooded ground in western
Amazonia. Flowering August to November.
Couepia 245

COLOMBIA. Amazonas: Black & Schultes46-59 fl (IAN, NY); Schultes & Cabrera17497 fl
(GH, NY, US). PERU. Loreto: Tessmann5382 fl (NY). BRAZIL. Amazonas: CapanemaRB 19204 fl
(RB); Cavalcante1376 fl (HB); Frdes21322 fl (IAN, US); 23833 fr (IAN, NY); Pires, Rodrigues& Silva
7495 fl (IAN); Pranceet al 2484 fl (INPA, NY). Acre: Pranceet al 2739 fr (INPA, NY).

36. Couepia chrysocalyx (Poeppig & Endlicher) Bentham ex Hooker f.,

Mart. Fl. Bras. 14(2): 42. 1867; Macbride, Fl. Peru io80. I938.
Poeppig & Endlicher, Nov. Gen. et Sp. P1. 3: 75. 1845.
Moquileachrysocalyx

Tree to Io.o m tall. Leaves oblong, Io.0-22.0 cm long, 4.0-8.5 cm broad,

rounded to cuneate at base, acuminate at apex, with acumen 4.0-12.0 mm long,
glabrous above (or with sparse appressed pubescence when very young), densely
silver-gray-arachnoid beneath; primary veins 15-20 pairs, prominent beneath;
petioles 4.0-7.0 mm long sparsely pubescent, terete, with two sessile glands near
base of lamina. Stipules to 7.0 mm long, linear, caducous. Inflorescences short
densely-flowered axillary corymbose racemes, the rachis with short reddish-brown
pubescence. Bracts and bracteoles 1.5-3.5 mm long; deciduous. Receptacle cylindi-
cal, slightly curved, 9.0-I4.0 mm long, with short reddish-brown pubescence on
exterior, glabrous within except for deflexed hairs at throat, with thick walls leaving
the cavity small; pedicels 2.0-3.0 mm long. Calyx lobes rounded. Petals 5, white,
sparsely ciliate or glabrous on margins. Ovary densely villous. Style densely villous
almost to apex. Stamens ca o00, inserted in a complete circle, the filaments far
exserted, forming a dense tangled mass. Fruit globose to oblong, to 9.0 cm long;
epicarp verrucose or smooth, glabrous; mesocarp thick, fleshy; endocarp thick,
fragile, granular-fibrous, glabrous within.
TYPE. Poeppig 2306, Brazil, Amazonas, fl (lectotype, W; isolectotypes, F, NY);
2177, Brazil, Amazonas, fl (paratype, P).
DISTRIBUTION. Non-flooded forest and secondary forest in Amazonia.
COLOMBIA. Putumayo: Klug 1854 fl (A, F, GH, K, MO, NY, US); Schultes3566 fl (GH).
Caqueta: Cuatrecasas8963 fr (COL); Killip 624 fl (US); Perez-Arbeldez624A fl fr (COL). Vaup6s:
Schultes,Baker & Cabrera18136 fl (GH, US). ECUADOR. Cazalet & Pennington7690 fl (FHO, K);
7760 fr (FHO, K, NY). PERU. Loreto: Killip & Smith 27865 fr (US); 27933 fl (F, NY, RB, US);
Spruce4614 fl (BM, BR, F, GH, GOET, K, LD, NY, OXF, P); LI. Williams5012 st (A, F, US). San
Martin: Klug 2656 fl (A, BM, F, K, MO, NY, US); 3971 fl (BM, F, GH, K, MO, NY, US); Woytkowski
5059 fl (NY, US). BRAZIL. Amazonas: DuckeRB 35557 fl (RB); Frdes21485 fr (IAN, US); Froes &
Addison28645 fl (IAN); Pires 834 fl (IAN); Rodrigues333 fr (INPA 443, NY); Schultes& Pires 8989 fl
(IAN, NY); Spruce3022 fl (K, P). Acre: Pranceet al 7934 fl (INPA, NY). Para: Baker57 fl (BM, GH, K,
LE, MO, NY, P, U, US); HuberMG 3664 fl (MG); MG 4843 fl (MG); MG 9359 fl (BM, F, MG, RB,
U); Oliveira2307 fl (IAN); Spruce844 fl (K, P).

LOCAL NAMES AND USES. Brazil and Peru: Parinari. Cultivated widely in the
Amazon basin for its edible fruit, and as an ornamental.

37. Couepia eriantha Spruce ex Hooker. f., Mart. Fl. Bras. 14(2): 45, t. I4.
1867.

Shrub or small tree. Leaves oblong to oblong-lanceolate, I0.0-I4.0 cn long,

3.0-5.5 cm broad, cordate to cuneate at base, apiculate or acuminate at apex, with
acumen to 5.0 mm long, glabrous on both surfaces; primary veins 15-20 pairs,
prominent beneath; petioles 3.5-6.o mm long, pubescent. Stipules to 3.0 mm long,
linear, caducous. Inflorescences terminal and axillary panicles, the rachis and
246 Flora Neotropica

branches densely brown-villous. Bracts and bracteoles ovate, to I5.0 mm long,

persistent. Receptacle cylindrical, I5.0-i8.o mm long, densely ferrugineous-villous
on exterior, glabrous within except for deflexed hairs at throat. Calyx lobes rounded,
sericeous. Petals 5, white, pubescent, ciliate on margins. Stamens ca 40, inserted in a
complete circle. Ovary densely villous. Style pubescent for three fourths its length.
Fruit (immature) long and narrow, with smooth glabrous epicarp.
TYPES. Spruce313, Brazil, Para, fl (lectotype, K; isolectotypes, CGE, GH,
OXF); 1138, Brazil, Amazonas, fl (paratypes, CGE, GOET, K, LD, LE, OXF).
DISTRIBUTION. Primary forest on periodically flooded ground, in central Amazon-
ian Brazil. Flowering November to March.
BRAZIL. Amazonas: Batten Poole 31 fl (K); CoelhoINPA 7259 fl (IAN, INPA, MG, NY);
Ducke908 fl (F, IAN, MG 17757, MO, NY, R, US); MG 7919 fl (BM, MG, RB); Pranceet al 8155 fl
(INPA, NY). Para: Black 47-1959 fl (IAN, NY); Ducke MG 8879 fl (BM, MG, RB); MG 11514 fl
(BM, MG); RB 18231 fl (RB); Froes31219 fl (IAN).

LOCAL NAME. Pajurd do igapo.

38. Couepia trapezioana Cuatrecasas, Brittonia 8: I97. I956.

Medium to large tree. Leaves ovate to oblong-elliptic, IO.O-I5.0cm long,

4.0-10.0 cm broad, rounded at base, acuminate at apex, with acumen 3.0-5.0 mm
long, glabrous above, dense gray or brown-appressed-arachnoidbeneath; primary
veins 14-18 pairs, prominent beneath; petioles 5.0-Io.o mm long, sparselypubescent,
shallowly canaliculate. Stipules to 2.0 mm long, linear, caducous. Inflorescences
terminal panicles, the rachis and branches with short light brown pubescence.
Bracts and bracteoles oblong-ovate, acute, 6.o-8.o mm long, caducous to subper-
sistent. Receptacle cylindrical, 7.0-8.0 mm long, densely light-brown-pubescent on
exterior, glabrous within except for deflexed hairs at throat; pedicels I.Omm long.
Calyx lobes narrow, rounded to acute. Petals 5, the margins ciliate. Stamens ca 24,
inserted in a nearly complete circle, the filaments connate at base. Ovary villous.
Style pubescent for half its length. Fruit (immature) ellipsoid; epicarp smooth,
glabrous; mesocarp thin, fleshy; endocarp thin, fragile, granular, glabrous within.
TYPE. Schultes8266, Colombia, Amazonas, fl (holotype, F; isotypes, IAN, U).
DISTRIBUTION. Forest in western Amazonia.
COLOMBIA. Amazonas: Pires 315 fr (NY). PERU. Ruiz & Pavon sn fl (BM). BRAZIL.
Amazonas: Schultes& Black 46-153 fl (NY).

This species is most closely related to C. schottii,the latter differing in the cordate
leaf bases, the subcampanulate receptacle, and the fewer stamens which are free to
their bases.

39. Couepia stipularis Ducke, Arch. Inst. Biol. Veg. Rio de Janeiro 4: 3. 1938.

Tree to 30.0 m tall. Leaves oblong to oblong-lanceolate, Io.o-i8.o cm long,

3.7-7.0 cm broad, rounded to subcordate at base, mucronate at apex, with acumen
3.0-4.0 mm long, glabrous above, with a few short gray hairs beneath; primary
veins 14-20 pairs, prominent beneath; petioles 9.0-I5.0 mm long, silvery-gray-
pubescent, terete, with two glands near lamina base. Stipules lanceolate, to 15.0 mm
long, persistent and adnate to base of petiole, silvery-gray-pubescent. Inflorescences
Couepia 247

terminal and axillary panicles 4.5-9.0 cm long, densely-flowered, with a thick, erect,
striated rachis, the rachis and branches with sparse appressed gray pubescence.
Bracts and bracteoles I.o-5.0 mm long, lanceolate, caducous. Receptacle ca 7.0 mm
long, subcylindrical and curved anteriorly, gray-pubescent on exterior, glabrous
within except for deflexed hairs at throat; pedicel 3.0 mm long. Calyx lobes rounded.
Petals 5, white, caducous, glabrous or with a few isolated hairs on margins. Stamens
27-35, inserted in a nearly complete circle. Ovary pilose. Style glabrous except for
base. Fruit ovoid, 3.5-5.0 cm long, 2.5-3.5 cm broad; pericarp thin, fragile, composed
of coarse fibers, glabrous within.
TYPE. Ducke RB 35583, Brazil, Amazonas, fl (holotype, RB; isotypes, INPA,
K, NY, P, U, US).
DISTRIBUTION. Known only from non-flooded forest in the vicinity of Manaus.
BRAZIL.Amazonas:Ducke1755fl fr (A, F, IAN, K, MG I7759, NY, R, US).

This species is distinct from all other members of the genus by the rather large
persistent stipules and the inflorescence consisting of a thick rachis with the flowers
clustered around it.

40. Couepia reflexa Ducke, Arch. Jard. Bot. Rio de Janeiro 5: 118. 1930.

Medium-sized tree. Leaves membraneous, oblong-elliptic, Io.o-I7.o cm long,

4.0-8.0 cm broad, subcuneate at base, acuminate at apex, with acumen 9.0-Io.o mm
long, glabrous on both surfaces; primary veins IO-I3 pairs, prominent on both
surfaces; petioles 5.o-8.o mm long, glabrous, terete. Stipules ca 2.0 mm long,
caducous. Inflorescences axillary reflexed racemose panicles to 8.o cm long, the
rachis and branches sparsely silver-graypubescent. Bracts and bracteoles 1.O-3.0 mm
long, lanceolate, persistent. Receptacle cylindrical, ca 7.0 mm long, sparsely silver-
gray-pubescent on exterior, glabrous within except for deflexed hairs at throat;
pedicels 3.0 mm long. Calyx lobes oblong, rounded. Petals 5, white, ciliate on margins.
Stamens ca 35, inserted in a complete circle. Ovary pilose. Style glabrous except at
base. Fruit unknown.
TYPE. Ducke RB I9757, Brazil, Para, fl (holotype, RB; isotypes, IAN, K, NY,
P, U).
Known only from the type gathering, made in primary forest on
DISTRIBUTION.
high ground at Juruti Velho.
This species is distinct by the sparsely pubescent deflexed inflorescence, and the
thin membraneous leaves.

41. Couepia longipendula Pilger, Notizbl. Bot. Gart. Berlin 6: 4I. I914.

Tree 5.0-30.0 m tall. Leaves oblong-elliptic, 6.0-I6.o cm long, 3.0-7.5 cm

broad, rounded to cuneate at base, caudately acuminate at apex, with acumen
I.0-I.5 cm long, glabrous on both surfaces, primary veins 5-8 pairs, almost plane;
petioles 4.0-8.0 mm long, shallowly weakly canaliculate, glabrous. Stipules lanceolate,
to 3.0 mm long, persistent. Inflorescences pendulous panicles on long (to 60.0 cm)
peduncles, mostly glabrous but sparsely pubescent towards apex. Bracts and brac-
teoles I.0-3.0 mm long, apiculate, caducous. Receptacle turbinate, Io.o-I5.0 mm
248 Flora Neotropica

long, with short light brown pubescence on exterior, completely glabrous within.
Calyx lobes rounded, with two sessile glands on exterior. Petals 5, rose, caducous,
ciliate on margins. Stamens ca 32, inserted in a nearly complete circle, with a few
toothed staminodes interspersedon distal side opposite ovary. Ovary densely villous.
Style glabrous, flattened at base. Fruit round to ellipsoid, 4.0-6.0 cm long, 4.0 cm
broad; epicarp with short sparse brown pubescence; pericarp hard, thick, fibrous,
not differentiated, densely lanate within.
TYPE. Ule 8854, Brazil, Amazonas, fl (holotype, B, lost; lectotype, K; isotypes, L,
LA, MG 13876).
DISTRIBUTION. Non-flooded forest in the lower Rio Negro region of Brazil.
to
Flowering August November.
BRAZIL. Amazonas: ArensINPA 410 fr (NY); Black & Ledoux50-10758 fr (IAN); ChagasIJPA
3309 fr (INPA, NY); INPA 4246 fl. (INPA, NY); INPA 5893 fl (INPA); CoelhoINPA 1733 fl (INPA,
NY); INPA 21129 fr (INPA, NY); Duarte 6880 fl (RB); Ducke 19 fl (A, F, IAN, K, MG, MO, NY, R,
US); 43 fl fr (A, F); RB 23592 fl (K, P, RB, U, US); RB 23593 fl (RB, US); Ferreira57-I fr (INPA
5050); Frdes26561 fl (IAN, UB); 28130 fl fr (IAN, P); INPA 5893 fl (INPA, NY); Mello INPA i816 fl
(IAN, INPA, MG, NY); 2026 fl (IAN, INPA, MG, NY); Prance2090 fr (NY); 4638 fr (INPA, NY);
RodriguesINPA 1602 fl (IAN, INPA, MG, NY); Rodrigueset al 2154 fr (INPA 8539, NY); 2332 fl
(INPA 9680, NY); 4649 fl (INPA 13158, NY); 4697 fl (INPA 1321 , NY); 6757 fl (INPA, NY); 71z6
fl (INPA 15778, NY); 7392 fr (INPA 16793, NY); 7605 fr (INPA 16908, NY); Spruce3825 st (K).

LOCALNAMESANDUSES.Castanhade galinha. The fruit contains much oil, and this

species is cultivated around Manaus for the edible cotyledons.
This species is remarkable for its long peduncles. This feature distinguishes it
from all other species of Couepia,as do the two sessile glands on each of the calyx lobes.
In the forest this species is a tall tree, but in cultivation, however, it is a low tree with
a much spreading crown which descends almost to ground level.

42. Couepia cognata (Steudel) Fritsch, Ann. Naturh. Mus. Wien 4: 60. 1889;
Sandwith, Kew Bull. 1931: 375. I93I; Kleinh. in Pulle, Fl. Suriname
2(I): 452. I939.
Hirtella cognataSteudel, Flora 26: 76I. 1843.
CouepiasteudelianaWalpers, Ann. 2: 463. 1852. Type. Hostmann795, Suriname, fl (CGE, GH, K,
LE, NY, P, U).
ParinarihostmanniiFritsch, Ann. Naturh. Mus. Wien 5: 13. I890. Type. Hostmann795.

Shrub or small tree I.o-Io.o m tall (rarely large trees). Leaves oblong to oblong-
lanceolate, 2.5-6.0(-I5.0) cm long, I.o-2.o(-4.5) cm broad, cuneate to rounded at
base, acuminate at apex, with acumen 2.0-I2.0 mm long, glabrous or sparsely
appressed-pubescent above, with gray-to brown-lanate pubescence beneath which
usually obscures the prominently reticulate parallel venation; midrib prominent
beneath, impressed above, pubescent on both surfaces; primary veins 7-IO(-I5)
pairs, prominent beneath; petioles 2.0 mm long, densely pubescent. Stipules to
9.0 mm long, linear-filamentous, deciduous. Inflorescences terminal panicles, the
rachis and branches brown-villous. Bracts and bracteoles lanceolate, to 6.0 mm long,
caducous. Receptacle cylindrical, curved anteriorly, 5.0-8.0 mm long, reddish-
brown and often villous on exterior, glabrous within except for deflexed hairs at
throat; pedicels I.0-3.0 mm long. Calyx lobes acute. Petals 5, white or pale pink,
glabrous. Stamens 12-25, unilateral, with a few teeth-like staminodes opposite them.
Ovary villous. Style pubescent. Fruit globose; epicarp pubescent when young,
verrucose and glabrous when mature; pericarp hard and thick, undifferentiated
densely lanate within.
Couepia 249

Key to the Varieties of Couepiacognata

i. Leaves thick and coriaceous; flowers shaggy-villous on exterior.
2. Leaves 2.5-6.0 cm long x 1.0-2.0 cm broad. a. var cognata.
2. Leaves 7.0-15.0 cm long x 3.0-4.5 cm broad. b. var major.
i. Leaves thin and membraneous; flowers with short ferrugineous pubescence on exterior.
c. var membranacea.

42a. Couepia cognata var cognata

Couepiacognata(Steudel) Fritsch, Ann. Naturh. Mus. Wien 4: 60. 1889.
ParinaricanescensGleason, Bull. Torrey Club 58: 370. I931. synon nov. Type. Tate 870, Venezuela,
Terr. Amazonas, fl (holotype, NY; isotype, K).
Licania exiguifolia Standley, Bull. Torrey Club 67: 285. 1940, synon nov. Type. Pinkus 245,
Guyana, fr (holotype, F; isotypes, MO, NY, S).

Leaves thick and coriaceous, 2.5-6.5 cm long, I.0-2.0 cm broad; flowers shaggy-
villous on exterior.
TYPE. Hostmann 795, Suriname, fl (holotype, P; isotypes, CGE, GH, K, LE, NY,
U).
DISTRIBUTION. Savanna, savanna forest, scrubland, and secondary forest in the
Guianas and adjacent Venezuela and Brazil.
VENEZUELA. Bolivar: Steyermark75526 fr (F, NY); Steyermark& Dunsterville93035 fr (NY).
GUYANA. Cowan39271 fr (GH, K, NY, US); De La Cruz 2107 fl fr (F, GH, K, MO, NY, US); 2215
fl (F, GH, MO, NY, US); Davis 387(F.D. 2383) fl fr (BRG, K, U); Fanshawe2489(F.D. 5225) fl (BRG,
K, NY); 2613(F. D. 5401) fl (BRG, K, NY); Harrison1239 fr (K); Jenman6031 fl (BRG, K). SURI-
NAME. B.W. 5529 fl (A, K, MO, NY); Focke455 fl (U); Heyligersr65 fl (U); Kegel 1155 fl (GOET);
Lanjouw& Lindeman1837 st (U); Lindeman4139 fr (NY, U); 4635 fl fr (NY, U); Schulz 8933 fl (U).
BRAZIL. Amazonas: Froes28080 fl (IAN, UB).

LOCAL NAMES.Venezuela: Acuri-yu-yek.Guyana: Inikiakia. Sluriname: Sabane

Kwepie.

42b. Couepia cognata var major Prance, var nov

A varietate cognatadiffert, foliis maioribus, 7.0-I5.0 cm longis, 3.0-4.5 cm latis.

Arbor magna ad 33.0 m alta.
TYPE. Fanshaweiooo(F.D. 3736), Guyana, Bartica-to-Potaro road mile IO7, fl
(holotype, NY; isotypes, BRG, K).
DISTRIBUTION. Primary forest in the Potaro River region of Guyana.
GUYANA. Kaieteur plateau: Cowan& Soderstrom
1789 fl (NY, US); 1992 fl (NY, US).

42c. Couepia cognata var membranacea Prance, var nov

A varietate cognatadiffert, foliis membranaceis, tenuibus; calycis-tubus lobisque

extus breviter rufo-tomentosis; calycis-lobis rotundatis.
TYPE. Fanshawe& Maguire32566, Guyana, Upper Mazaruni River, fl (holotype,
NY; isotypes, F, US).
DISTRIBUTION. Forest and savanna woodlands in the Upper Mazaruni region
of Guyana.
GUYANA. Fanshawe& Maguire32436 fl (F, NY, US).
250 Flora Neotropica

The difference between these varieties are constant but insufficient to merit
specific recognition. The varieties are ecologically but not geographically isolated
from each other.

43. Couepia multiflora Bentham, Jour. Bot. Hooker 2: 215. i840.

Moquileamultiflora(Bentham) Walpers, Rep. 2: 7. 1843.

Tree to 6.o m tall. Leaves oblong to oblong-elliptic, 9.o-15.0 cm long, 4.0-9.0

cm broad, rounded to cordate at base, rounded to short-acuminate at apex, with
acumen I.0-5.0 mm long, glabrous above, densely gray-arachnoid beneath; midrib
prominent beneath, pubescent above; primary veins I2-I4 pairs, prominent beneath,
slightly impressed above; petioles 4.0-7.0 mm long, pubescent, terete, eglandular.
Stipules linear, to 9.0 mm long, caducous, membraneous. Inflorescences terminal
panicles, the rachis and branches densely brown-tomentose. Bracts and bracteoles
2.0-5.0 mm long, oblong to lanceolate, caducous. Receptacle ca 7.0 mm long,
cylindrical, curved anteriorly in bud, with short thick brown tomentum on exterior,
glabrous within except for deflexed hairs at throat; pedicels I.o mm long. Calyx
lobes rounded. Petals 5, white, the margins glabrous or with a few isolated hairs.
Stamens ca 28, inserted in an arc of ca 240?, with a sterile portion of circle opposite
them. Ovary pilose, Style pubescent for half its length. Fruit unknown.
TYPE.Schomburgk 112, Guyana, fl (holotype, K; isotypes, BM, BR, CGE, F,
GH, L, LE, NY, OXF, P).
DISTRIBUTION. Savannas and savanna scrubland in Guyana and Terr. Roraima,
Brazil. Flowering October to April.
GUYANA. Appun1747 fl (K, P); Graham390 fl (K); 500 fl (K); Jen-nano188 fl (BRG, K); Schom-
burgk309 fl (BR, K); 321 fl (BM, CGE, F, K, OXF); A. C. Smith2502 fl (A, F, K, MO, NY, P, U, US).
BRAZIL. Terr. Roraima: Ducke 2950 fl (K); RB 15138 fl fr (K, P); Frdes 22997 fl (IAN, NY, U);
KuhlmannRB 2950 fl (F, INPA, K, P, RB, U, US); RB 15138 fl (K, RB, U, US); RB 17976 fl (K, RB,
U, US); Pranceet al 9 13 fl (INPA, NY); 9580 fl fr (INPA, NY); Ule 764z fl (K, MG 12756); 7885 fl
(K, MG 12977).

This species resembles C. ovalifoliaof southern Brazil, but differs in the larger
leaves and in the cylindrical, slightly curved receptacle. It is distinct from C. schottii
in the long, cylindrical receptacle. It should be noted that this species one of northerly
distribution has its closest affinities with these two species of the coastal savannas of
southern Brazil.

44. Couepia ulei Pilger, Verh. Bot. Ver. Brand. 47: 149. 1905; Macbride, Fl
Peru. 1082. 1938.

Tree to I5.o m tall, or shrub. Leaves elliptic, 5.0-Io.5 cm long, 2.5-4.5 cm

broad, rounded at base, acuminate at apex, with acumen 3.0-I I. mm long, glabrous
above, densely gray-to rufous-pubescent beneath; primary veins 10-13 pairs, promi-
nent beneath; petioles 3.0-7.0 mm long, pubescent, shallowly canaliculate, eglandu-
lar. Stipules to 3.0 mm long, linear, caducous. Inflorescencesterminal, little branched
panicles, the rachis and branches with short gray-brown pubescence. Bracts and
bracteoles I.0-2.0 mm long, ovate, caducous. Receptacle subcylindrical, 5.0-7.0 mm
long, shortly gray pubescent on the longitudinally striate exterior, glabrous within
except for deflexed hairs at throat; pedicels I.0-2.0 mm long, Calyx lobes rounded
Couepia 251

to acute. Petals 5, white, pubescent on exterior, ciliate on margins. Stamens ca 35,

inserted in a complete circle. Ovary villous. Style pubescent for half its length. Fruit
elongate ellipsoid, 5.0 cm long, 2.0 cm broad; epicarp glabrous, verrucose; mesocarp
fleshy, thin; endocarp thin, fragile, granular, densely pubescent within.
TYPE. Ule 5547, Brazil, Amazonas, fl (holotype, B, lost; lectotype, K; isotypes,
INPA, L, MG 5493, RB).
DISTRIBUTION.Riverine forest and non-flooded forest in western Amazonia.
Flowering continuously.
COLOMBIA. Amazonas: Duque-Jaramillo2316 fr (COL); Schultes6731 fr (COL, F). PERU.
Loreto: Asplund14320 fr (US); LI. Williams7 fl (F, US); 29 fr (F); 63 fl (F); 88 fl (F); 917 st (F, US);
1115 st (F, US); 2346 fl fr (F, US); 8157 fl fr (F, US). BRAZIL. Amazonas: Ducke 1541 fl fr (A, F,
IAN, INPA, MG, NY, R, RB, US); 1545 fl fr (A, F, IAN, MG, NY, R, RB 50642, US); Krukoff6037
fl (A, BM, BR, F, K, MO, NY, RB, U, US); Rodrigues5445 fl (INPA 14114); Rodrigueset al 6994 fl
(INPA I5656, NY); 71o6 st (INPA 15768, NY, RB); M. Silva 2048 fr (MG). Acre: Pranceet al 2968 fr
(INPA, NY).

LOCALNAME.Peru: Suichi-Parinari.

45. Couepia comosa Bentham, Jour. Bot. Hooker 2: 215. I840.

Moquileacomosa(Bentham) Walpers, Rep. 2: 6. I843.

Tree to 8.0 m tall or shrub. Leaves ovate-elliptic, 4.0-8.5 cm long, 2.0-3.8 cm

broad, short-acuminate at apex, with acumen I.0-5.0 mm long, rounded to cordate
at base, glabrous when young above, densely reddish-brown-pubescent beneath;
primary veins 9-12 pairs, prominent beneath; midrib pubescent on both surfaces;
petioles 2.0-4.0 mm long, short-pubescent, terete. Stipules linear, I.0-2.0 mm long,
caducous. Inflorescences compact little-branched terminal panicles, the rachis and
branches with short brown pubescence. Bracts and bracteoles to 3.0 mm long, ovate,
caducous. Receptacle subcylindrical, 8.o-Io.o mm long, with short brown pubes-
cence on exterior, glabrous within except for deflexed hairs at throat; pedicels 3.0
mm long. Calyx lobes rounded. Petals 5, white, sparsely pubescent on exterior,
ciliate on margins. Stamens ca 65, inserted in a complete circle, the filaments far
exserted. Ovary pilose. Style densely pubescent for two thirds its length. Fruit (in
single specimen examined) ellipsoid, 4.0 cm long, 1.8 cm broad; epicarp verrucose;
pericarp undifferentiated, hard, thick.
TYPE. Schomburgk28, Guyana, fl (holotype, K; isotypes, BM, BR, CGE, F, L,
LE, NY, P).
DISTRIBUTION.Rocky places beside rivers and on river islands in Guyana.
Flowering September to April.
GUYANA. Appun623 fl fr (K); Fanshawe1674 (F.D. 44o1) fl (BRG, K); 2180 (F.D. 4916) fl
(BRG, K); 2820(F.D . 5619) fl (K, NY); Hohenkerk 524 fl (BRG, K); Jenman761 fl fr (BRG, K); 969 fl
(BRG, K); o188 fl (BRG); 7627 fl (BRG); Linder15 fl (GH, NY); Sandwith687 fl (K, NY, P, RB, U,
US); Tutin 20 fl fr (K, US); Schomburgk244 fl (BM, OXF); 356 fl (K).

46. Couepia venosa Prance, sp nov Fig 40 J-O.

Arbor vel frutex, ramulisjuvenilibus glabris aetate lenticellatis. Stipulae parvae,

deciduae. Folia alternata, petiolata, petiolo 7.0-I3.5 mm longo, glabro; laminae
252 Flora Neotropica

oblongo-ellipticae, 8.5-15.0 cm longae, 3.5-6.0 cm latae, apice in acuminem ad

Io.o mm longum contractae, basi rotundatae vel subcuneatae, supra glabrae sat
nitidae, subtus dense tomentosae; costis secundariis I -I4 jugis, subtus prominenti-
bus. Flores in paniculis terminalibus brevissime pedunculatis dispositi, breviter
brunneo-tomentosi; pedicelli 2.0-3.0 mm longi. Bracteae bracteolaeque parvae,
oblongae, caducae. Receptaculum subcampanulatum, 6.0-7.0 mm longum, extus
dense brunneo-tomentosum, intus prope faucem pilis longis deflexis infra stamina
dispositis, versus basim glabrum. Calycis-lobi 5, rotundati. Petala 5, decidua, alba,
pubescentia. Stamina 37-50, in orbem completum disposita. Ovarium ad faucem
receptaculi insertum, dense pubescens. Stylus e basi ovarii ortus, pilosus. Drupa
ellipsoidea, extus glabra, laevis; mesocarpio tenue; endocarpio granulato, tenue,
fragilo, intus glabro.
TYPE. Hoehne 17197, Brazil, Sao Paulo, Alto da Serra, fl (holotype, NY; isotypes,
K, SP).
DISTRIBUTION. Mountain slopes in southern-central Brazil.
BRAZIL. Serra da cubatao, Burchell3694 fl (GH, K, P). Minas Gerais: Serra de Antonio Per-
reira, SchwackeI3682 fl (RB 47975); Ouro Preto, Schwacke10235 fl (NY, P, RB 47976). Sao Paulo: Alto
da Serra, Andrade69 fl (R); Emvgdio1526 fl (R); Kuhlmann3174 fr (SP); 3374 fl (SP); Schwebeli9 fl
(R, SP). Guanabara: Rio de Janeiro, Schwacke6653 fl. (RB 47827).

47. Couepia polyandra (Kunth) Rose, Contr. U.S. Natl. Herb. 5: I96. 1899;
Standley, Contr. U.S. Natl. Herb. 23(2): 344. I922; Standley & Steyer-
mark, Fieldiana Bot. 24(4): 443. I946.
HirtellapolyandraKunth, Nov. Gen. et Sp. 6: 246. 1823; DC., Prodr. 2: 529. 1825.
Hirtella dodecandra Mocifio & Sesse ex A. P. De Candolle, Prodr. 2: 529. 1825. Type. Mociiio &
Sess6 tabula 302 in A.L.P.P. De Candolle, Calq. des Dess. Fl. Mex., cf Herb. Mociiio &
Sesse 2282 fl (MA, F frag.) cf'Pavon' sn fl (BM). (see Prance, 1968, p 129).
MoquileakunthianaMartius & Zuccarini, Abh. Akad. Minchen i: 390. 1820.
Couepiakunthiana(Martius & Zuccarini) Bentham, Jour. Bot. Hooker 2: 216. I840.
Grymaniapolyandra(Kunth) Presl, Epim. Bot. I93. 1849.
Couepiafloccosa Fritsch, Ann. Naturh. Mus. Wien 5: 12. I890. Type. Friedrichsthal 256, Guatemala,
fl (holotype, W).
Couepiadodecandra(A. P. De Candolle) Hemsley in Hook. Ic. 27. 6: 2620, 2621. I9o0; Standl.,
Publ. Field Mus. Bot. 3: 274. 1930.

Tree to 15.o m tall. Leaves oblong to oblong-elliptic, 6.0-I3.o cm long (on

flowering branches, longer on sterile and fruiting branches), 2.5-5.5 cm broad,
rounded to cuneate at base, acuminate at apex, with acumen 2.0-10.0 mm long,
glabrous above when mature, with a caducous pubescence when young, densely
arachnoid beneath; midrib prominent above, pubescent when young; primary veins
8-I5 pairs, prominent on both surfaces; petioles 4.0-7.0 mm long, with two in-
conspicuous medial glands, terete. Stipules 2.0-4.0 mm long, linear, membraneous,
caducous. Inflorescences terminal and axillary panicles, the rachis and branches with
short light-brown pubescence. Bracts and bracteoles 1.0-3.5 mm long, ovate, cadu-
cous. Receptacle subcylindrical, ca 4.0 mm long, with short appressed pubescence on
exterior, glabrous within except for deflexed hairs at throat; calyx lobes rounded.
Petals 5, white, glabrous but with ciliate margins. Stamens 1-2I,
I inserted in an arc
of I80-240?, with a few staminodes opposite them. Ovary villous. Style pubescent for
half its length. Fruit ellipsoid to obovoid, 5.0-7.0 cm long, 3.0-4.5 cm broad when
mature; epicarp smooth, glabrous; mesocarp thick and fleshy; endocarp thin, fragile,
of a fibrous texture, glabrous within.
TYPE. Humboldt & Bonpland sn, Mexico, fl (P).
Couepia 253

DISTRIBUTION.Riverine forest and low woodland in Central America from

Sinaloa, Mexico, to Costa Rica.
Representative collections: MEXICO. Sinaloa: Montes & Salazar 712 fl (US); Ortega955 fl (K);
6198 fl (GH, US). Nayarit: McVaugh 19158 fr (US); Ortega58 fl (A, K, US); 6285 fl (K); Rose 1515 fr
(US); 3120 st (US); 3310 st (US); Rose et al I4430 fl (NY, US); Viereck1197 fl (US). Jalisco: Howell
10297 fr (US); McVaugh z186i fl (US); 11879 fl (MICH, NY); I5729 fr (MEXU); Nelson 4122 fl
(GH, US); Phillips sn st (MEXU);Rzedowski 16566 fl (MEXU, US). Colima: Jones32 fr (US); McVaugh
15729 fr (US). Michoacan: Hinton 13750 fl (GH, NY, US). Puebla: C. E. Smithet al 4104 fr (MEXU,
NY, US). Veracruz: Gomez-Pompa 947 fl (GH). Guerrero: Hinton 14008 fl (K, NY, US); 14126 fl (GH,
NY, US); Langman3341 fl (MEXU); Palmer 401 fl (A, F, GH, K, NY, US). Oaxaca: ElmoreD 14 fl
(F, GH, NY, US); Nelson 2372 fl (GH, US); Reho 3224 fr (US). Tabasco: Barlow 29/21 fl (GH);
Guzmdnsn fl (MEXU); Linden 1603 fl (K, LE); Matuda3102 fl (A, F, K, MEXU, NY); 3585 fr (A, F,
K, MEXU, MO, NY). Chiapas: Matuda 1ogo fl (A, MEXU, MO, NY); 3798 fr (A, F, MEXU, NY);
i6500 fl (F, MEXU, NY); Purpus51 fl (LE, US); 1oo51 fl (NY); 10531 fl (NY); 12074 fl (A, F, K, MO,
NY). Yucatan: Flores sn st (F); Gaumer23205 st (F); 24320 fl (F, GH); Liebmann1746 fl (US); Scholt
579 st (F). Without precise locality: Haenke1532 st (F); Rovirosa179 fl (US). GUATEMALA: Aguilar
H.36 fl (A, F); Lundell3267 fl (F, MEXU, UC); 3719 fl (A, US); 4881 fl (MO); Standley87330 st (A);
87457 st (F); 89034 st (F); 89549 st (US); Steyermark48074 st (A, NY, US); 44534 st (US); 52144 st
(US); 52145 st (A, US). BRITISH HONDURAS. Cook& Martin 22 fr (US); Gentle29 fl (F, K, NY);
Lundell4186 fl (BM, F); Robertson401 fl (BM); Schipp569 fr (A, BM, F, GH, K, NY); 717 fl (A, BM,
F, GH, K, MO, NY); Standley 54290 st (US). Honduras. Johansen 6 st (F); Standley 54290 st (A,
F, US); VanSeveren67 fl (US). EL SALVADOR. Allen 7279 (F, NY, US); Calderdn272 fl (GH, NY);
Standley 19666 st (GH, MO, NY, US); 21410 st (GH, NY, US); 22149 fl (GH, NY, US); 23190 st
(GH, NY, US); Velasco8961 fl (F, US). NICARAGUA. Garniersn fl (F); Levy 42 fl (K, P). COSTA
RICA. Cook & Doyle 68i fl (US); Standley& Valerio46556 st (US); Tonduz 13858 fl (BM, GH, K, P,
US); Valerioii6 fl (US).

LOCALNAMES. Mexico: Palo del Fraile, Pio, Sapotede nino, Usplo, 2apotillo, Guaya-
bitode tinta. British Honduras: Babooncap, Monkeycap. Honduras: Munzap, Zapotillo.
El Salvador: Ulazapote,Zapotillo,Zapotilloamarillo,Zapotebolo.Costa Rica: Olozapo,
Zapotillo.
The species cited in synonymy were all placed under C. polyandraby St-ndley
and Steyermark in their Flora of Guatemala (Fieldiana Bot. 24(4): 443. 1946), and
there is certainly only this single species of Couepiain Central America. The various
synonyms were delimited by number of stamens, but an unbroken range from I I to
21 was noted in the material examined. This species is most closely related to the
Colombian species of C. trapezioana.
Hirtelladodecandra,one of the synonyms of C.polyandra,is an example of the species
described by A. P. De Candolle from the Mocifio & Sesse drawings. Since the original
drawings, which should be the types, are missing, for the present the tracingspublished
by A.L.P.P. De Candolle in his Calquesare regarded as the types. The Mocifio and
Sesse herbarium material which was not seen by De Candolle, is not type material,
but is useful in the interpretation of the inadequate type tracings. In the case of
Hirtelladodecandra the drawing is in fact sufficient to place the species, because there
are so few species of Chrysobalanaceae growing in Mexico. Further details about the
Mocinioand Sesse drawings are given in Prance (i968).

48. Couepia platycalyx Cuatrecasas, Fieldiana Bot. 27: 66. I950.

Tree. Leaves obovate to obovate-orbicular, 8.5-12.0 cm long, 7.0-9.0 cm broad,

rounded at base and apex, glabrous above, with very sparse short appressed pubes-
cence beneath; primary veins 10-12 pairs, prominent beneath; petioles 7.o-8.o mm
long, glabrous, terete. Stipules cadcuous. Inflorescences terminal little-branched
 254 Flora Neotropica

panicles, the rachis and branches with a few minute appressed hairs only. Bracts and
bracteoles to 1.5 mm long, caducous. Receptacle broadly turbinate, flattened, almost
solid, oblique, 3.0 mm long, with appressed pubescence on exterior, with dense de-
flexed hairs at throat within filling the very small cavity; pedicels i.5 mm long.
Calyx lobes rounded. Petals glabrous, white. Stamens ca 54, inserted in a complete
circle. Ovary villous. Style pubescent at base only. Fruit unknown.
TYPE. Cuatrecasas & Perez-Arbeldez15336, Colombia, Cundinamarca, fl (holo-
type, F; isotypes, COL, US).
DISTRIBUTION. Known only from two gatherings on the eastern range of the
Andes in Colombia and Venezuela.
et al 103618 fl (NY, VEN).
VENEZUELA. Lara: Steyermark

LOCAL NAME. Venezuela: Mapurito montaniero.

As a species of Couepia,C. platycalyxis unique by its flattened, almost solid
receptacle. However, in all other features it is a Couepia.and I have followed Cuatre-
casas' placement of it in this genus.

49. Couepia rufa Ducke, Arq. Mus. Rio de Janeiro 22: 66. I9I9.
PleraginarufaArruda da Camara ex Koster, Trav. 499. I8i6, nom illegit, see Prance, Mem. N.Y.
Bot. Gard. I5(1): 133. 1966.
MoquilearufaBarbosa Rodrigues, Hort. Flum. 165. 1895 pro parte.
Couepiaglazioviana Pilger, Notizbl. Bot. Gart. Berlin 8: 537. I923, synon nov. Type. Glaziou
18217, Brazil, Guanabara, fl (holotype, B, lost; isotypes, BR, LE, NY, PR).

Large tree. Leaves elliptic, 10.0-27.0 cm long, 6.5- I.5 cm broad, subcordate
at base, rounded to short-acuminate at apex, glabrous above, densely rufous-arach-
noid beneath, the margins undulate and revolute; primary veins 15-25 pairs, promi-
nent beneath, impressed above; petioles 5.0-Io.o mm long, thick. Stipules linear, to
17.0 mm long, caducous. Inflorescences terminal many-flowered panicles to 15.0 cm
long, the rachis and branches striate, thick, shortly gray-brown-pubescent.Bracts and
bracteoles oblong to oblong-lanceolate, to 7.0 cm long, caducous. Receptacle tur-
binate, thick-walled, ca I2.0 mm long, shortly light-brown-pubescent on exterior,
glabrous within except for deflexed hairs at throat; pedicels 1.0-3.5 mm long. Calyx
lobes rounded, 5.0-7.0 mm long, Petals 5, white, with ciliate margins. Stamens ca 60,
inserted in a complete circle. Ovary pilose. Style pubescent for three fourths its
length. Fruit ovoid, to i I.o cm long and 6.0 cm broad; epicarp verrucose; mesocarp
thin, fleshy; endocarp thick and fragile.
TYPE.Lutz MG 3575, Brazil, Pernambuco, fl (BM, F, MG, P, R, RB, US).
DISTRIBUTION. Coastal primary forests of Pernambuco. Flowering September to
December.
BRAZIL. Pernambuco: BarrosCorreia4215 fl (IPA); Ducke2113 fl (IAN, MG 22659, NY, R, RB);
Lima 48-122 fl (IPA, RB); o100 fl (IAN); Pickel 831 fl (IPA); Prance 4807 fr (NY); Sobrinho614 fl
(IAN); sn fl (IPA). Rio deJaneiro (cult): Glaziou 1382 st (BR); 1383 st (BR).

LOCAL NAMES AND USES. Coro de Pernambuco,Oiti coro. Cultivated for the edible
fruit which is sold in the markets of Recife.
This distinct species is easily recognized by the undulate and revolute leaf
species group, but differs in the larger flower
margins. It is closest to the C. bracteosa
and fruit, the greater number of stamens, the verrucose epicarp of the fruit, etc. The
identity of Pleraginahas been fully discussed elsewhere (Prance I966a).
Couepia 255

50. Couepia robusta Huber, Bol. Mus. Emilio Goeldi 6: 76. I9IO.

Tree to 30.0 m tall. Leaves oblong to elliptic, 9.0-23.0 cm long, 4.0-I 1.0 cm
broad, rounded to cordate at base, rounded or shortly acuminate at apex, with
acumen I.0-5.0 mm long, glabrous above, densely rufous-brown-arachnoidbeneath;
primary veins 15-21 pairs, prominent beneath; petioles 5.0-8.0 mm long, with short
brown pubescence. Stipules to 5.0 mm long, lanceolate, caducous. Inflorescences
terminal little-branched panicles, the rachis and branches with reddish-brown
tomentum. Bracts and bracteoles 3.0-7.0 mm long, ovate, caducous. Receptacle
subcampanulate, 7.0-9.0 mm long, densely reddish-brown-pubescent on exterior,
glabrous within except for deflexed hairs at throat; pedicels 2.0 mm long. Calyx
lobes rounded, 6.0-7.0 mm long, densely reddish-brown-pubescent on exterior,
densely light-brown-pubescent within. Petals 5, white, ciliate on margins. Stamens
ca 43, inserted in an arc of ca 240?, with a toothed ridge of staminodes opposite them.
Ovary densely villous, often with two carpels developing. Style pubescent for half its
length. Fruit ovoid, 5.0-6.0 cm long, 3.5-4.0 cm broad; epicarp smooth, glabrous;
mesocarp thick, fleshy, fibrous; endocarp thin, fragile, granular, sparsely pubescent
within.
TYPE,R. S. RodriguesMG 965I, Brazil, Para, fl (holotype, MG; isotypes, BM,
NY, P, RB, U, US).
DISTRIBUTION. Non-flooded forest in Brazilian Amazonia. Flowering August to
December.
BRAZIL. Amazonas: Ducke 1267 fl (A, F, IAN, MG 17760, NY, R, RB, US); 2015 fl (A, COL,
IAN, INPA, MG, NY, R, RB 6oi 14, U, US); Frdes33627 fl (IAN); 34675 fl (IAN); INPA 6045 fr
(INPA); LoureiroINPA 16147 fl (INPA, NY); 16449 fl (INPA, NY); Pranceet al 3641 fl (INPA, NY);
Rodrigues7701 fr (INPA I7006); Rodrigues& Osmarino6034 fl (INPA 15085, NY); 6924 fl (INPA 15477,
NY); 7642 fr (INPA I6946, NY). Para: DuckeMG 15469 fl (BM, MG); MG I5484 fl (BM, MG, RB,
US); MG 15553 fr (MG, RB); Kuhlmann&Jimbo 289 fr. (IAN, INPA); Mus. Goeldi9721 fl (BM, MG).

LOCALNAMES.Pajurd, Mari bravo.

51. Couepia impressa Prance, sp nov Fig 41 F-J.

Arbor, ramulis juvenilibus glabris. Stipulae circa 2.0 mm longae, subulatae,

deciduae. Folia alternata, petiolata, petiolo 6.o-I2.0 mm longo, glabro, supra
canaliculato, eglanduloso; laminae oblongae vel oblongo-lanceolatae, I i.o-i8.5 cm
longae, 4.0-6.0 cm latae, apice in acuminem 3.0-7.0 mm longum contractae, basi
rotundatae, supra glabrae, subtus sparse pubescentes; costa media glabra; costis
secundariis 18-23, subtus prominentibus, supra impressis. Flores in paniculis ter-
minalibus dense griseo-pubescentibus dispositi. Bracteae bracteolaeque ovatae,
2.0-3.5 mm longae, caducae. Receptaculum subcampanulatum, 6.0-7.0 mm
longum, sursum curvatum, extus griseo-puberulum, intus sub-fauce pilis longis
deflexis dense tomentosum, versus basim glabrum. Calycis-lobi 5, acuti. Petala 5,
decidua, margine breviter ciliata. Stamina 15-17, in orbem 2/3 disposita, basi brevi-
ter connata. Ovarium ad faucem receptaculi insertum, villosum. Stylus e basi ovarii
ortus, medio pubescens. Fructus mihi ignoti.
TYPE. Lima 54-1897, Brazil, Pernambuco, Mata Dois Irmaos, fl (holotype,
IPA; isotype, IPA).
Primary forest of northeastern Brazil.
DISTRIBUTION.
BRAZIL. Pernambuco: Mata Dois Irmaos, Prance 4809 st (NY, from type tree). Alagoas:
Pontual 66-280 fl (IPA). Bahia: Coaraci, Belem & Pinheiro2948 fl (NY, UB).
256 Flora Neotropica

Couepiaimpressais nearest the Amazonian C. trapezioana,but the latter species

differs in the erect receptacle, the connate bases of the stamens, the yellow-brown
tomentum of the inflorescence, and the smaller number of primary leaf veins.
Couepiamultiflorais also similar, but differs in the yellow-brown tomentum of the
inflorescence, the greater number of stamens, the smaller number of primary leaf
veins which are plane with the upper surface of the lamina, the petioles which are
terete, and the midrib which is pubescent above.

52. Couepia meridionalis Prance, sp nov Fig 37 A-D.

Arbor?; ramulis juvenilibus lanato pubescentibus mox glabris. Stipulae parvae,

caducae. Folia alternata, petiolata, petiolo 5.0-Io.o mm longo, lanato pubescenti,
supra canaliculato; laminae ellipticae vel oblongo-ellipticae, 6.5-11.5 cm longae,
3.0-6.0 cm latae, apice in acuminem 3.0-5.0 mm longum contractae, basi rotundatae,
supra glabrae, subtus dense lanato-tomentosae; costis secundariis I0-I3 jugis, subtus
prominentibus. Flores in paniculis terminalibus pauciramosis dense sericeo-tomen-
tosis dispositi; pedicelli 1.5-2.0 mm longi. Bracteae bracteolaeque parvae, caducae.
Receptaculum cylindricum, ad 9.0 mm longum, extus dense sericeo-tomentosum,
intus prope faucem pilis longis deflexis infra stamina dense pilosum, versus basim
glabrum. Calycis-lobi 5, dense pubescentes. Stamina plus minusve 25, in orbem
completum disposita. Ovarium ad faucem receptaculi insertum, dense pilosum.
Stylus e basi ovarii ortus, pilosus. Fructus mihi ignoti.
TYPE.Mosen3377, Brazil, Sao Paulo, Santos, fl (holotype, S; isotypes, K, LE, P).
Known only from the type collection.
DISTRIBUTION.
Most closely related to the Amazonian species C. chrysocalyx, the present species
differs in the branched inflorescence, the smaller leaves with fewer primary veins,
the smaller flowers, the pubescent petals and the smaller number of stamens. It is
also near to C. ulei but differs in many small features, especially the pubescence of the
inflorescence and flowers, the longer petioles, and the larger flowers.
The specific name was suggested by the extremely southern distribution com-
pared with most of the genus.

53. Couepia pernambucensis Prance, sp nov Fig 37 K-O.

Arbor alta, ramulis juvenilibus puberulis mox glabris lenticellatis. Stipulae

parvae, caducae. Folia alternata, breviter petiolata, petiolo 4.o-5.0 mm longo,
lanato pubescenti, eglanduloso, supra canaliculato; laminae ovato-ellipticae vel
oblongae, 5.o-6.5 cm longae, 2.0-2.8 cm latae, apice in acuminem 2.0-6.0 mm
longum contractae, basi rotundatae, supra glabrae, subtus sparse araneoso-pubes-
centes; costa media basim versus pubescenti, costis secundariis 8-I0 jugis subtus
prominentibus. Stipulae ca I.0 mm longae, subulatae. Flores in paniculis terminali-
bus et axillaribuspauci-ramosisdispositi,rachi et ramulisdense tomentello-pubescenti-
bus. Bracteae bracteolaeque ad 2.5 mm longae, caducae. Receptaculum cylindricum,
5.o-6.o mm longum, sursum curvatum, extus dense tomentello-pubescens, intus
prope faucem infra stamina dense tomentosum, basim versus glabrum. Calycis-lobi
5, acuti. Pedicelli I.o mm longi. Petala 6, decidua, alba, marginibus ciliata. Stamina
14-I7, in orbem completum disposita. Ovarium ad faucem receptaculi insertum,
Couepia 257

G ~H

El ./'~~~'

UD E

FIG 41. Species of Couepia.A-E, C. froesii (Frdes32164); A, habit, xo.5; B, leaf undersurface,
x 0.5; C, petal, x 4; D, flower, x 2; E, flower section, x 2. F-J, C. impressa(Lima54-1897); F, habit,
x o.5; G, flower, x 2; H, flower section, x 2; J, petal, x 4.
258 Flora Neotropica

dense pilosum. Stylus e basi ovarii ortus, pilosus. Drupa ellipsoidia, extus glabra,
laevis; endocarpio granulato, tenue, fragilo, intus glabro.
TYPE. Lima53-1206, Brazil, Pernambuco, Recife, Mata Dois Irmaos, fl (holo-
type, IAN; isotype, IPA).
DISTRIBUTION. Known only from the type gathering, collected in primary forest
at Dois Irmaos.
This species is most closely related to the C. guianensissuperspecies but differs in
the dense pubescence of the inflorescence and flowers, and in the strongly curved
receptacle.

54. Couepia froesii Prance, sp nov Fig 41 A-E.

Arbor parva, ramulis juvenilibus glabris lenticellatis. Stipulae parvae, caducae.

Folia alternata, petiolata, petiolo 6.0-8.0 mm longo glabro eglanduloso, supra
canaliculato; laminae oblongae, I .o-23.o cm longae, 4.5-8.5 cm latae, apice in
acuminem 8.0-14.0 mm longum caudato-contractae, basi rotundatae, supra glabrae,
subtus lanis brevissimis griseis dense pubescentes; costa media glabra, costis secun-
dariis 20-24 jugis, subtusprominentibus,supra impressis.Flores in paniculis terminali-
bus axillaribusque pauci-ramosis dispositi, rachi ramulisque dense griseo-pubescenti-
bus. Bracteae magnae vel parvae, deciduae, bracteolae parvae, caducae. Receptacu-
lum subcampanulatum, 6.0 mm longum, extus griseo-pubescens,intus prope faucem
infra stamines dense tomentosum, basim versus glabrum. Calycis-lobi 5, acuti.
Pedicelli I.o mm longi. Petala 5, decidua, alba, marginibus breviter ciliata. Stamina
circa 30 in orbem completum disposita. Ovarium ad faucem receptaculi insertum,
dense pilosum. Stylus e basi ovarii ortus, medio pubescens. Drupa immatura ovalis,
extus laevis, pubescens mox glabra; endocarpio granulato, tenue, fragilo.
TYPE.Froes32164, Brazil, Para, Porto do Moz, fl (holotype, IAN; isotype, NY).
forest in central Brazilian Amazonia.
DISTRIBUTION. Non-flooded
BRAZIL.Amazonas:Rio Demini,Froes& Addison28964, fl (IAN). Par,: Portode Moz, Fr6es
32195 fl (IAN, NY).

This species is closest to C. excelsa,but differs in the smaller flowers, the glabrous
interior of the receptacle, the canaliculate petioles, and the glabrous upper surface of
the midrib.

55. Couepia parvifolia Prance, sp nov

Arbor parva, ramulis juvenilibus tomentellis mox glabris. Stipulae ad 2.0 mm

longae, caducae. Folia alternata petiolata, petiolo 5.0-7.0 mm longo, lanato-
pubescenti mox glabro, eglanduloso, supra canaliculato; laminae oblongae vel
oblongo-lanceolatae, 2.5-4.5 cm longae, o.8-I.3 cm latae, apice in acuminem
5.0-7.0 mm longum contractae, basi cuneatae, supra glabrae, subtus dense lanato-
tomentosae; costa media glabra, costis secundariis 7-9 jugis, subtus prominentibus.
Flores in paniculis terminalibus pauci-ramosis dispositi, rachi ramisque brunneo-
tomentosis. Bracteae bracteolaeque I.0-2.5 mm longae, ovatae, caducae. Receptacu-
lum subcampanulatum, 5.o-6.o mm longum, extus brunneo-tomentosum, intus
prope faucem infra stamina dense tomentosum, basim versus glabrum. Calycis-lobi
5, rotundati. Pedicelli brevi, 0.25 mm longi. Petala 5, dense pubescentia, marginibus
ciliata. Stamina circa 28, in orbem completum disposita. Ovarium ad faucem recep-
Couepia 259

taculi insertum, dense lanato-pubescens. Stylus e basi ovarii ortus. Fructus mihi
ignoti.
TYPE. Duarte1508, Brazil, Rio de Janeiro, Serra do Petr6polis, fl (holotype, NY;
isotype RB 65362).
DISTRIBUTION. Known only from the forests of Rio de Janeiro and Guanabara,
Brazil.
BRAZIL. Guanabara, near Horto Florestal, HortoFlorestal1012 fl (NY, RB I39762).

LOCAL NAME. Milho Cozido.

The small, oblong-lanceolate leaves clearly distinguish this species from all others.

Excluded Species
Couepialaurifolia Cuatrecasas, Brittonia 8: I98. 1956. =Persea laurifolia (Cuatr.)
Cuatr. (Lauraceae).
panamensisStandley, Trop. Woods 44: 22. 1935. =Maranthescorymbosa
Couepia Blume.

7. Hirtella Linnaeus, Sp. P1. 34. I753 (Hort. Cliff. 17. I738); Gen. P1. ed. 5.
20. I754; Lam., Illustr. 2: II3. 1793; Persoon, Syn. I: 249. I805; DC.,
Prodr. 2: 528. 1825; Dum., Anal. Fam. 40. 1829 (sub "Hirtellia");Mart.
& Zucc., Abh. Akad. Munchen 7: 372. I832; Zucc., Flora 15(Beibl. 2):
78. 1832; Hooker f. in Benth. & Hook., Gen. P1. I: 608. 1865; Hooker f,
Mart. Fl. Bras. 14(2): 27. i867; Focke in Engl. & Prantl, Nat. Pflanzenfam.
3(3): 59. I894; Pittier, Contrib. Fl. Venez. 23. 1923; Macbride, Fl. Peru
1074. 1938; Kleinhoonte in Pulle, Fl. Suriname 2(I): 444. I939; Prance,
in press.
TachibotaAublet, PI. Guiane I: 287. I775.
CauseaScopoli, Introd. n. 928. 2Io. I777.
SalmasiaSchreber, Linn. Gen. PI., ed. 8. I: 201. 1789 nom illegit.
CosmibuenaRuiz & Pavon, Prodr. Fl. Peruv. o0, t. 2. 1794; non Cosmibuena Ruiz & Pavon, Fl.
Peruv. Chil. 3: 2. I802, t. 198. 1789.
ThelyraThouars, Nov. Gen. Madag. 21. i8o6.
Brya Vellozo, Fl. Flum. 146. 1825, & 4 t. I. 1835; non Brya Browne (1756).
SphenistaRafinesque, Sylva Tellur. 90. 1838.
Zamzela Rafinesque, Sylva Tellur. go. 1838.

Trees or shrubs with hermaphrodite flowers 3.0-I I.o mm long. Leaves entire,
usually glabrous, or hirsute or with a few appressed hairs beneath, sometimes with
two large bulbous myrmecophilous inflations at the junction of the lamina with the
petiole. Petioles eglandular. Bracts and bracteoles commonly with few to many
stalked or sessile glands, or eglandular, not enclosing the young flowers in small
groups. Inflorescences panicles or elongate to fasciculate racemes, if paniculate then
frequently racemoid or thyrsoid, when branched the branches often bearing several
sterile bracts and terminating with a single flower or a few cymosely arranged flowers.
Receptacle usually campanulate, rarely elongate-cylindrical, hollow, usually glab-
rous inside at the base and with retrorsehairs at the throat, glabrous to tomentose or
hispid on exterior, lined by a well-defined disc. Calyx lobes acute to rounded, usually
reflexed. Stamens 3-9 (-io), unilateral in most species, in a complete circle in two
species, with staminodes present or absent; filaments usually far exceeding calyx
lobes, rarely only slightly exceeding calyx lobes. Ovary of one unilocular carpel,
inserted laterally at the mouth of the receptacle or less frequently midway up the
260 Flora Neotropica

receptacle; ovules 2. Style filiform, far exserted. Fruit a fleshy drupe, most often
ellipsoid; epicarp smooth and glabrous or rarely densely tomentose; endocarp thin,
hard, bony, exterior usually smooth to slightly channelled, hirsute within, usually
with 4-7 lines of weakness which allow the seedling to escape. Germination hypogeal,
the first leaves alternate.
TYPE SPECIES. Hirtella americanaLinnaeus. The name Hirtella is derived from the
Latin hirtus =hairy. This refers to the hirsute pubescence of the leaves, young
branches, and inflorescence of the type species.
Mexico and Central America. West Indies, Colombia, Ecuador,
DISTRIBUTION.
through the Guianas and Amazonia to Bolivia and Southern Brazil; also three species
in East Africa and Madagascar.
Taxonomic History of Hirtella
Among genera of the Chrysobalanaceae, Hirtellahas one of the longest histories,
having first appeared in the literature in Linnaeus' HortusCliffortianus (I738). Valid
publication was established in SpeciesPlantarumof Linnaeus (1753). Several species
were published by earlier workers on the American flora such as, Chamisso &
Schlechtendal, De Candolle, Kunth, Lamarck, and Swartz and, as is common in
such cases, there was much confusion about the circumscription of some of these
L. and H. racemosa
earlier species, e.g H. americana Lam. These problems are discussed
and clarified below under the species concerned.
In his monograph of the genus for the Prodromus, de Candolle (1825) included
several species belonging to the other genera which had been described by earlier
workers as Hirtella.De Candolle acknowledged that six of the twenty species he listed
were incompletely known to him. The last comprehensive monograph of the entire
genus is that published in two places by Martius and Zuccarini (1832) and Zuccarini
(1832). This monograph is very good for the 23 species known to that date. Martius
and Zuccarini redefined the generic boundaries of Hirtella and gave an accurate
description of the genus which remains valid to this day. They separated Hirtella
from Couepiaon the basis of the stamen number correlated with the type of fruit, a
distinction which I have followed here and elsewhere (eg Prance, in press). If authors
subsequent to Martius and Zuccarini had adhered to their definitions of Hirtella
much of the more recent confusion over the circumscription of the genus would have
been avoided. Martius and Zuccarini reassigned a number of species, originally
described in Hirtella to other genera of the family. Their monograph presented 23
species which were divided into two groups based on the type of inflorescence (ie
racemose or paniculate). Since the time of these authors, many additional species of
Hirtellahave been described, and the genus has been treated regionally in the various
floras. J. D. Hooker (1867) in his account for MartiusFlora Brasiliensis,included 26
Brazilian species. In the New World the genus has not been subdivided into any
supraspecific taxa. The first African species to be recognized as an Hirtellawas H.
zanzibaricaOliver (1876), although a Hirtellafrom Madagascar was described under
the generic name Thelyraby Du Petit-Thouars (I806). It was not until the present
century, after several species of Magnistipulawere described from Africa, that Hirtella
became confused with Magnistipula.Most species of the African genus Magnistipula
have at some time been transferred to Hirtella.R. Graham (1957) gave reasons for
keeping Magnistipulaapart from Hirtella.In elaborating upon these, I clearly defined
the differences (Prance, in press).
Hauman (I 95 ) is the only worker who divided the genus into infrageneric taxa.
He proposed two subgenera, Euhirtellaand Afrohirtella,but as the latter is in fact
referable to Magnistipula,all the species considered below belong to subgenusEuhirtella
of Hauman. Hauman divided this subgenus into two series:
Hirtella 261

Digitibracteatae.Bracteae bracteolaeque glandulis stipitatis ornatae. Examples:

H. americana,H. zanzibarica.
Bracteae bracteolaeque glandulis sessilibus ornatae. Example:
Integribracteatae.
H. hirsuta.
These series are furtherdiscussedin the next section. In my Synopsis(Prance, in press),
beyond circumscribing Hirtella, I indicated the differences between Hirtellaand re-
lated genera. Little change was made to the limits of the genus, except to confirm
that the African Magnistipulais a separate genus. Even when Magnistipulais segre-
gated from Hirtellathere are still two species of Hirtellafrom East Africa which have
obvious affinities with the American species. In the same work I also clarified the
separation of Hirtellafrom Couepiaand Grangeria,the two most closely related genera
apart from Magnistipula.
The Subdivision of Hirtella
As outlined above, the division made by Hauman (I951), the only one to date,
is completely artificial. The two types of glands were his only criterion, but they are
in fact quite variable. One type of gland merges with the other, and in some species
both stipitate and sessile glands occur. Indeed, the example of Integribracteatae, (H.
hirsuta =H. paniculata),cited by Hauman, frequently has stipitate glands. Further-
more, Hauman's subdivision does not account for over a third of the species of
Hirtellawhich have eglandular bracts and bracteoles. While the gland type is useful
as a key character, it certainly is not sufficiently reliable or meaningful to divide
Hirtellainto sections or series. Gland type is not correlated with any other character,
and some species may even be glandular or eglandular.
Most regional treatments and the early monographs divided Hirtella into two
groups based on the type of inflorescence (racemose or paniculate). The majority of
species fall clearly into one group, or the other, but in a few species the inflorescence
can be either paniculate or racemose (eg H. hebeclada,H. triandra).In my opinion,
the type of inflorescence cannot be used to divide the group into any meaningful in-
frageneric taxa. The only division which I have made is based on the myrmecophilous
swellings at the leaf base in six species. In the Chrysobalanaceae, this character is
confined, to these six obviously related species of Hirtella, although it occurs in a
number of unrelated families (eg Melastomataceae and Boraginaceae). Since the six
species with myrmecophilous swellings are otherwise closely related, I have created
a separate section for them. The majority of species, however, belongs to section
Hirtella,which cannot be reliably further subdivided.
Hirtella sect Myrmecophila Prance, sect nov
Folia basi cum duobus vesiculis formicariis munita.
Hirtella sect Hirtella
Leaf bases without swollen ant cavities.

Specific Characters in Hirtella

THE LEAF
The characters used are largely self explanatory. There is very little variation in
the leaf pubescence in Hirtella.The lower surface is either glabrous, hirsute, or with a
few sparse stiff appressed hairs.
THE GLANDS
One of the most useful characters in Hirtella is the glands of the bracts and
bracteoles, and of the pedicels. Some species have more than one type of gland, but
in the vast majority of species the gland type is constant. The gland types are:
I. Bracts and bracteoles with stipitate glands (Fig 42B). In this condition the
distinctly stalked glands are frequently abundant.
262 Flora Neotropica

A B

/ E

J2'okn
FIG 42. Bract glands in Hirtella. A, Sessile concave glands in H. racemosavar racemosa(Tillett et
al 45475) ; B, stipitate glands in H. glandulosa(Irwinet al 16745); C, solitary sessile glands in H. pilosissima
(Pires et al 889); D, sessile glands and translucent secretions in H. gracilipes (Pranceet al 59052); E,
solitary stipitate pedicel gland in H. burchellii(SpencerMoore I41).
Hirtella 263

2. Bracts and bracteoles with sessile glands (Fig 42A, C). These glands may be
extremely conspicuous, large and concave as in H. racemosa var racemosa,or they may
be plane and inconspicuous. However, if a plant is said to possess sessile glands in this
work, they can be clearly seen on at least 25 percent of the bracts and bracteoles.
There are usually fewer sessile glands. Most frequently they consist of a pair of
glands towards the base of the bracteoles.
3. Pedicels glandular (Fig 42E). A few species have one or more stipitate glands
arising from the pedicels. The bracts may be either eglandular or with sessile glands.
The pedicel glands vary from leaf-like bracts with an apical gland to solitary glands
on a long slender stalk. These glands are probably derived from a reduction of the
bracts borne on the pedicels of some species. In paniculate species the branches
naturally bear many bracts, and the pedicel is never glandular. Some species with
unbranched inflorescences which are called racemose here, bear bracts on the
pedicels which are perhaps the result of reduction from the paniculate type of
inflorescence. In a further stage of reduction the pedicels may bear only a single
stalked gland.
In addition to the glands on the bracts and pedicels, glands may occur on the
calyx lobes or on the stipules. I have not been able to determine the function of these
glands, but in some species they excrete a sticky substance which makes the in-
florescence and flowers distinctly viscous. In H. gracilipesand related species there are
very few glands, but the bracts and bracteoles are covered with translucent globules
of the viscous secretions exuded by secretory cells. (Fig 42D).
THE INFLORESCENCE
Two main types of inflorescence occur in Hirtella,racemose and paniculate. In
the majority of species the inflorescencesare clearly of one type or the other. The term
racemose, here generally restricted to a completely unbranched inflorescence of
pedicellate flowers, also includes the species whose inflorescencebranches are reduced
to bracts on the pedicels (see above under glands, section 3). In some species with
characteristically paniculate inflorescences, the simple racemose state may appear in
a few collections.
THE FLOWER
The floral characters used here for Hirtella are self-explanatory except for the
measurements. Flower size is measured from the tip of the longest calyx lobes to the
base of the receptacle, and pedicel length from the base of the receptacle to the
junction with the rachis or inflorescence branch.

Key to Species of Hirtella

I. Leaf base with myrmecophilous swellings. Section Myrmecophila.
2. Inflorescence fasciculate (bunched racemes) sometimes cauliflorous.
3. Stamens 4-5; ovary inserted at middle of receptacle; leaves 9.0-I9.0 cm long, coria-
ceous, the lower surface bullate. i. H. myrmecophila.
3. Stamens 6-7; ovary inserted at mouth of receptacle; leaves 17.0-30.0 cm long, mem-
braneous, the lower surface not bullate. 2. H. physophora.
2. Inflorescence an elongate raceme or panicle, never cauliflorous.
4. Inflorescence a panicle, sometimes only slightly branched at base.
5. Inflorescence only slightly branched at base; leaves oblong-lanceolate; bracts
eglandular. 3. H. vesiculosa.
5. Inflorescence with many small lateral branches; leaves ovate; bracts glandular.
4. H. dorvalii.
4. Inflorescence an elongate raceme, never branched.
6. Exterior of receptacle and calyx lobes hirsutulous; inflorescence 15.0-30.0 cm long.
5. H. guainiae,
264 Flora Neotropica

6. Exterior of receptacle and calyx lobes sparsely hispid only; inflorescence 2.0-12.0
cm long. 6. H. duckei.
i. Leaf base without myrmecophilous swellings.
7. Inflorescence paniculate (rarely only very slightly branched.) (For contrasting state-
ment see p 267).
8. Bracts and/or pedicels glandular (with stalked or sessile glands). (For contrasting
statement see p 266).
9. Leaves glabrous beneath when mature, or rarely with a few short stiff appressed
hairs.
o0. Stamens 6-7.
I . Bracts with few sessile glands towards apex only; inflorescence rachis and
branches and flower exterior glabrous or glabrescent. 7. H. macrosepala.
1 . Bracts with numerous stalked glands; inflorescence rachis and branches
and flower exterior pubescent.
12. Leaves 3.5-8.5 cm long, 1.4-4.5 cm broad; stipules linear to 6.0 mm
long; African. 86. H. zanzibarica.
85. H. megacarpa.
12. Leaves (6.4-)8.o-I3.0 cm long, 4.0-7.0 cm broad; stipules small,
deltoid to 2.0 mm long; American.
13. Calyx lobes glandular; bract glands small, ca 0.I mm in diameter;
rachis and branches of inflorescence sparsely puberulous; leaves
glabrous beneath. 8. H. ulei.
I3. Calyx lobes glandular; bract glands large, ca 0.5 mm in diameter;
rachis and branches of inflorescence tomentellous; leaves sparsely
pubescent beneath. 9. H. glabrata.
Io. Stamens 3-5.
14. Glands large and solitary on a long stalk, arising from pedicel or junction
of inflorescence; inflorescence distinctly thyrsoid. 10. H. carbonaria.
14. Glands usually numerous, long or short stalked, arising from the margins
of bracts and bracteoles; inflorescence not thyrsoid.
I5. Inflorescence a long rachis bearing small clusters of flowers on very
short branches; leaves oblong-lanceolate. I. H. araguariensis.
I5. Inflorescence spreading, with many distinct lateral branches; flowers
not in distinct clusters; leaves oblong to elliptic.
i6. Leaves 3.0-5.5 cm long, cordate at base; inflorescences scarcely
branched, hirsutulous. 12. H. cordifolia.
16. Leaves usually cuneate or subcuneate at base (if subcordate then
7.0-17.0 cm long and inflorescence glabrescent); inflorescences
usually much branched.
17. Bracteolar glands few, usually either sessile on margins or apex
terminating in a single gland; inflorescence usually pilose or if
glabrescent then corymbose; leaf apex acuminate.
I8. Leaves 12.0-17.0 cm long, rounded to subcordate at base; fer-
tile stamens 5. 13. H. insignis.
I8. Leaves 3.5-I5.0 cm long, cuneate at base; fertile stamens 3.
I9. Leaves I0.0-15.0 cm long; bracts membraneous, the
glands several, borne on margins; inflorescence 7.0-I8.0
cm long, terminal. 14. H. tocantina.
I9. Leaves 4.0-9.0 cm long; bracts coriaceous, with single
reflexed apical gland only; inflorescence 3.0-1.0 cm
long, axillary or terminal. 15. H. piresii.
17. Bracteolar glands numerous, borne on the margins, usually
shortly stalked; inflorescence spreading but not corymbose,
usually glabrescent, if tomentellous then leaf apex rounded to
acute.
20. Leaves I.8-4.5 cm long, rounded to acuminate at apex,
thickly coriaceous; primary veins 4-7 pairs. 37. H. bahiensis.
20. Leaves 6.0-I3.5 cm long, acuminate at apex, thinly coria-
ceous; primary veins 9-10 pairs.
21. Stamens 3; bracteoles coriaceous, ovate; petioles 3.0-4.0
mm long. I6. H. davisii.
2I. Stamens 4; bracteoles membraneous, lanceolate; petioles
1.0-2.0 mm long. 17. H. subglanduligera.
Hirtella 265

9. Leaves hirsute beneath at least on principal venation, usually more generally so.
22. Calyx lobes with small stipitate glands.
23. Leaves orbicular to ovate-elliptic, 3.0-6.5 cm long, retuse to mucronate
at apex (rarely acuminate); trunk with corky bark; receptacle pubescent
within to base. I8. H. ciliata.
23. Leaves oblong to ovate, 4.5-23.0 cm long, acute to acuminate at apex;
trunk with thin bark; receptacle glabrous within except at throat.
24. Venation of leaves prominulous above; inflorescence subcorymbose,
bearing numerous tightly clustered flowers; stamens 7. 19. H. hoehnei.
24. Venation of leaves impressed or prominulous above; inflorescence
lax, spreading, not subcorymbose, flowers loosely arranged; stamens
3-5.
25. Leaves subcordate at base, subconduplicate, the lower surface
merely sparsely appressed-pubescent. 29. H. adderleyi.
25. Leaves rounded to cuneate at base, rarely weakly subcordate but
never subconduplicate, the lower surface hirsute. 20. H. glandulosa.
22. Calyx lobes eglandular.
26. Venation of leaves impressed or bullate above.
27. Inflorescence and flowers rufous-tomentose; bracts bearing very few
sessile glands, these often obscured by pubescence. 42. H. obidensis.
27. Inflorescence and flowers tomentellous to puberulous, not rufous;
bracts bearing many obvious stalked or sessile glands.
28. Leaf surface usually distinctly bullate; base usually subcordate.
21. H. bullata.
28. Leaf upper surface plane but with impressed venation, base rounded
to subcuneate.
29. Bracteolar glands ca 0.5 mm in diameter, sessile or slightly-
stipitate; bracteoles distinct, ovate. 22. H. americana.
29. Bracteolar glands ca o. mm in diameter, on long slender stalks;
bracteoles reduced to a mass of glands with a slightly connate
base. 27. H. tentaculata.
26. Venation of leaves prominulous or plane above, never bullate.
30. Stamens 3.
31. Bracteolar glands large, numerous, apparent; inflorescence tomen-
tellous, many-flowered; epicarp glabrescent; endocarp thin and
bony. 22. H. americana.
3I. Bracteolar glands small, few; inflorescence hispid, few-flowered;
epicarp tomentose; endocarp thick and fibrous. 23. H. guatemalensis.
30. Stamens 4-7.
32. Epicarp tomentose; endocarp thick and fibrous; bracts with few
inconspicuous stipitate or sessile glands.
33. Inflorescence hispid, the flowers lax; bracts with small short-
stipitate glands; stamens 3-4. 23. H. guatemalensis.
33. Inflorescence tomentellous, bearing small clusters of flowers on
long thick rachis; bracts with a few sessile or a single terminal
gland; stamens 6. 24. H. eriandra.
32. Epicarp glabrescent; endocarp thin and bony; bracts with many
conspicuous stipitate glands.
34. Inflorecence hispid-hirsute; bracts with short-stalked glands
arising from their margins, the secretory tip small, ellipsoid.
35. Inflorescences predominantly terminal; leaf bases rounded to
weakly subcordate. 25. H. paniculata.
35. Inflorescences predominantly axillary; leaf bases deeply
cordate. 26. H. deflexa.
34. Inflorescence puberulous to tomentellous; bracts either reduced
to numerous, long-stalked clavate glands arising from the junction
of the pedicels with the stem, or ovate with glands borne on their
margins, the secretory tip large, flattened forming an expanded
head.
36. Glands arising from junction of pedicel and stem, with a long
slender stalk and a small secretory tip, the whole appearing
clavate; young stems puberulous or sparsely tomentellous.
28. H. macrophylla.
266 Flora Neotropica

36. Glands arising from the bracts, with a thick stalk and a large
secretory tip, which is flattened to form an expanded head;
young stems with short dense compact puberulence.
37. Leaves 6.5-13.0 cm long; inflorescence I2.0-I5.0 cm
long; stipules to 2.0 mm long, deltoid. 9. H. glabrata.
37. Leaves 5.0-7.0 cm long; inflorescence 3.5-8.o cm long;
stipules to 5.0 mm long, linear. 29. H. adderlyi.
8. Bracts and pedicels eglandular.
38. Leaf apex retuse, rounded, or acute, never acuminate.
39. Stamens 6; leaf apex predominantly rounded to retuse (or mucronulate).
30. H. punctillata.
39. Stamens 3; leaf apex acute.
40. Leaves ovate, thick-coriaceous; inflorescence compact, to 3.0 cm long.
31. H. corymbosa.
40. Leaves oblong, chartaceous; inflorescences lax and spreading, 4.0-8.0 cm
long. 36. H. triandra.
38. Leaf apex distinctly acuminate.
41. Stamens 3.
42. Leaf base subcordate; inflorescence 25.0-50.0 cm long, pendulous.
32. H. pendula.
42. Leaf base rounded to cuneate; inflorescence 1.0-15.0 cm long, erect.
43. Leaves lanceolate, at least three times longer than broad; inflorescence
and exterior of flowers glabrescent. 33. H. barrosoi.
43. Leaves oblong to elliptic, not exceeding 2.5 times longer than broad;
inflorescence and exterior of flowers usually pubescent, rarely glabrous.
44. Inflorescence and exterior of flowers glabrous or glabrescent, or
leaf apex distinctly cuspidate with a long fine acumen; inflorescence
I.0-4.0 cm long, predominantly axillary. 50. H. bicornis.
44. Inflorescence and exterior of flowers pubescent or tomentose; leaf
apex acuminate but never cuspidate; inflorescence 2.0-8.0 cm long,
predominantly terminal.
45. Inflorescence and young stem hispid; calyx tube ca 6.o mm, the
lobes ca 5.0 mm; corolla lobes ca 8.o mm long; stipules 6.o-
o0.0 mm long. 34. H. leonotis.
45. Inflorescence usually puberulous, rarely weakly hirsute; young
stem glabrous to tomentellous; calyx tube to 4.0 mm, the lobes
to 3.0 mm long; corolla lobes to 5.o mm long; stipules 2.0-6.0
mm long.
46. Inflorescence little-branched, almost racemose, but with a
few branches bearing two flowers, others one flower and a
single pair of bracts at junction with pedicel. 35. H. mutisii.
46. Inflorescence much-branched, the branches usually bearing
many bracts.
47. Leaves 4.0-14.5 cm long, 2.0-5.5 cm broad; pedicels
1.0-3.0 mm long. 36. H. triandra.
47. Leaves i5.0-I7.0cm long, 8.o-io.o cm broad; pedicels
0-0.5 mm long. 38. H. latifolia.
41. Stamens 4-9.
48. Inflorescence appearing racemose, but in fact either a long central rachis
bearing groups of flowers on short branches, or with a few branches with
only three flowers each; leaf base sometimes cordate.
49. Inflorescence almost a raceme, but with a few branches bearing two or
three flowers; flowers lax, not borne in distinct groups. 85. H. hebeclada.
49. Inflorescence a long central rachis with groups of flowers on short
branches.
50. Bracteoles large and persistent, 3.o-6.5 mm long, enclosing buds,
leaf base subcuneate; epicarp glabrous. 39. H. suffulta.
50. Bracteoles small not conspicuous, I.0-2.0 mm long; leaf bases sub-
cordate to subcuneate, epicarp tomentose or glabrous.
5I. Leaves subcordate at base, the lower surface glabrescent;
epicarp glabrous. 40. tI. elongata.
5 1. Leaves rounded to subcuneate at base, the lower surface sparsely
pilose-hirsute; epicarp tomentose. 24. H. eriandra.
Hirtella 267

48. Inflorescence a much-branched panicle without a long central rachis;

leaf base rounded to cuneate.
52. Inflorescence and exterior of flowers tomentose, hirsute, or hispid,
usually rufous to golden-brown.
53. Inflorescence and young branches sparsely hispid; leaves mem-
braneous. 41. H. rodriguesii.
53. Inflorescence and young branches tomentose or hirsute; leaves
coriaceous.
54. Venation of leaves slightly impressed above; inflorescence rufous-
tomentose. 42. H. obidensis.
54. Venation of leaves plane or prominulous above; inflorescence
brown-tomentose or-hirsute.
55. Leaves 2.5-6.0 cm long; inflorescence 0.8-4.0 cm long.
56. Inflorescence 0.8-i.5 cm long, clustered; leaves ovate,
rounded at base; stamens 7. 43. H. cowanii.
56. Inflorescence 2.0-4.0 cm long, lax; leaves orbicular to
oblong-orbicular, usually subcordate at base; stamens 5.
44. H. orbicularis.
55. Leaves 5.5-17.0 cm long; inflorescence 4.0-1 I.0 cm long.
57. Bracts 4.0-6.0 mm long, ovate, up to half the length of
receptacle; leaves ovate, thick-coriaceous, shortly mucro-
nate at apex; inflorescence tomentose. 45. H. guvanensis.
57. Bracts 1.0-3.0 mm long, oblong, to base of receptacle
only; leaves elliptic, subcoriaceous, acuminate (never
mucronate) at apex, inflorescence hirsutulous-tomentose.
46. H. lightioides.
52. Inflorescence and exterior of flowers gray-puberulous to glabrescent.
58. Inflorescence a subcorymbose panicle 4.0-6.5 cm long; petioles
7.0-8.0 mm long. 47. H. aramangensis.
58. Inflorescence a spreading panicle (not subcorymbose); petioles
0.5-5.0 mm long.
59. Laminas 9.0-I5.0 cm long, with distinct basal glands; inflores-
cence lax and spreading 7.0-I9.0 cm long. 48. H. rasa.
59. Laminas 2.5-8.0 (-9.5) cm long, the base eglandular or with
indistinct glands; inflorescence short and compact, 1.0-8.0 cm
long.
60. Inflorescence and exterior of flowers densely puberulous;
leaves thick and coriaceous, with prominent venation, the
apex acuminate but never cuspidate. 49. H. scabra.
60. Inflorescence and exterior of flowersglabrescent to puberulous;
leaves chartaceous, with plane to prominulous venation, the
apex acuminate to cuspidate.
50. H. bicornis.
7. Inflorescence a raceme.
6I. Glands present on bracts or pedicels (either stipitate or sessile). (For contrasting
statement see p 270).
62. Leaves narrowly linear-lanceolate, 5.5-I8.o cm long, 0.5-2.2 cm broad.
5I. H. angustissima.
62. Leaves ovate to oblong-lanceolate, 3.0-30.0 cm long, I.5-I0.0 cm broad.
63. Bracts bearing sessile glands only or with translucent glandular excretions,
glands often covering entire surface, most frequently paired towards base of
bracts and bracteoles; stipitate glands absent from bracts and pedicels, but
bract apex occasionally glandular.
64. Leaves oblong-lanceolate. 52. H. tenuifolia.
64. Leaves oblong to ovate.
65. Young stem and inflorescence thick-pilose or pilose-tomentose, the
stem with persistent pilosity 2.0-3.0 mm long; stemens 3-6.
66. Leaf margins hispid-ciliate; stamens 3-6; pedicels 1.5-3.0 mm
long; bracts axillary. 53. H. pilosissima.
66. Leaf margins eciliate; stamens 3; pedicels 3.0-6.o mm long;
bracts inserted on pedicels. 34. H. leonotis.
65. Young stem and inflorescence usually puberulous, sometimes tomentel-
lous; stamens 4-6.
268 Flora Neotropica

67. Leaves thin-membraneous, stamens 4; ovary inserted midway up

receptacle. 41. H. rodriguesii.
67. Leaves thick, coriaceous; stamens 5-6; ovary inserted at mouth of
receptacle.
68. Bracts and bracteoles caducous, with minute translucent secre-
tions, the sessile glands present or absent; pedicels 6.0-25.0 mm
long.
69. Bracts with sessile glands in addition to translucent secretions;
pedicels 6.0-i6.0 mm long, thick; inflorescence 4.5-I4.0 cm
long; leaves sparsely appressed-pubescent on venation or
glabrescent beneath. 54. H. gracilipes.
69. Bracts with glandular secretions only, other glands absent;
pedicels 12.0-25.0 mm long, very slender; inflorescence
1.5-5.0 cm long; leaves glabrous beneath. 55. H. brachystachva.
68. Bracts and bracteoles persistent, with sessile glands only; pedicels
1.5-I0.5 mm long.
70. Primary veins 6-Io pairs; leaves 3.5-I8.5 (-I9.5) cm long,
glabrous or rarely sparsely appressed-pubescent; mature
leaves plane, not slightlv bullate. 56 H. racemosa
70. Primary veins II-I5 pairs; leaves 19.5-25 o cm long or if
smaller (7.0-13.0 mm long) then hirsutulous beneath; mature
leaves slightly bullate.
71. Leaves 19.5-25.0 cm long, primary veins I3-15 pairs, lower
surface with few sparse appressed hairs. 57. H. juruensis.
71. Leaves 7.o-13.omm long; primary veins I1-13 pairs;
lower surface hirsutulous. 58. H. kuhlmannii.
63. Either bracts bearing stipitate glands, or with one or more solitary stipitate
glands arising from some pedicels, and bracts often glandular at apex only.
72. Bracts and bracteoles eglandular, or with sessile glands and a glandular
apex only; some pedicels with I-3 solitary stipitate glands.
73. Leaves ovate, cordate at base; bracts and bracteoles with a few sessile
glands. 59. H. standleyi.
73. Leaves oblong to oblong-lanceolate, usually rounded to subcuneate
at base, rarely subcordate; bracts and bracteoles eglandular or with
sessile glands.
74. Primary veins 17-19 pairs; leaves I9.o-26.ocm long; pedicels
8.0-12.0 mm long. 60. H. longifolia.
74. Primary veins 8-15 pairs; leaves 8.0-I8.o cm long (to 24.0 cm in
one species with short pedicels 0.5-2.0 mm long).
75. Pedicels slender 4.5-9.0 mm long (or if 3.5-5.0 mm then leaf
constricted above base); leaves glabrous beneath except for
pubescence on midrib and extreme base of margin; bracteoles
eglandular or glandular.
76. Fertile stamens 3; leaves oblong-elliptic 8.o-14.5 cm long;
bracts with apical glands. 6I. H. lemsii.
76. Fertile stamens 4-6; leaves elliptic to oblong-lanceolate;
bracts glandular or eglandular.
77. Leaves elliptic to oblong-elliptic, 3.5-8.0 cm long; bracts
usually with se;sile and apical glands. 56. H. racemosa.
77. Leaves oblong-lanceolate, 8.o-I8.o cm long; bracts
eglandular.
78. Leaves subauriculate because of slight constriction
above base; primary veins 11-13 pairs; ovary inserted
near mouth of receptacle; fertile stamens 5. 62. H. schultesii.
78. Leaves not constricted above base; primary veins 8-I5
pairs; ovary inserted at mouth or middle of receptacle.
79. Primary veins 12-15 pairs; tertiary venation slightly
impressed; ovary inserted midway up receptacle;
fertile stamens 4. 63. H. paraensis.
79. Primary veins 8-I o pairs; tertiary venation prominu-
lous; ovary inserted at mouth of receptacle; fertile
stamens 6. 64. H. sprucei.
75. Pedicels I.0-3.0 mm long, thick; leaves hirsute beneath or
Hirtella 269

glabrous with hirsute margin and midrib; bracteoles with

glandular apex and often sessile glands.
80. Leaves bullate above, with distinctly impressed venation,
secondary venation and reticulations extremely prominent
beneath. 65. H. lancifolia.
80. Leaves not bullate above, venation prominent or plane,
secondary venation and reticulations only prominulous be-
neath.
81. Leaves acuminate at apex, the lower surface glabrous to
sparsely hirsute. 66. H. burchellii.
81. Leaves usually mucronate at apex, the lower surface densely
hirsute or glabrous.
82. Inflorescence densely tomentose; leaves mucronate at
apex, hirsute beneath. 67. H. mucronata.
82. Inflorescence sparsely puberulous; leaves acuminate at
apex, glabrescent beneath. 56. H. racemosa.
72. Bracts and bracteoles with numerous stipitate glands; pedicels eglandular.
83. Receptacle cylindrical, elongate. 79. H. couepiflora.
83. Receptacle campanulate.
84. Leaves bullate above, with impressed venation. 21. H. bullata.
84. Leaves plane, not bullate above, with plane or prominulous vena-
tion.
85. Leaves orbicular, rounded to retuse at apex; pedicels 7.0-20.0
mm long. 68. H. longipedicellata.
85. Leaves oblong to elliptic, acuminate at apex; pedicels 4.0-I I.0
(-15.0) mm long.
86. Stipules bearing numerous stipitate glands. 69. H. glandistipula.
86. Stipules eglandular.
87. Inflorescence short, compact, densely-flowered, 3.0-7.0 cm
long; calyx bearing numerous glands; leaves drying gray.
70. H. martiana.
87. Inflorescence lax, 3.0-15.0 cm long; calyx eglandular or
the glands scarce; leaves drying green to brown.
88. Flowers 8.o-9.0 mm long; calyx lobes 5.o-6.o mm
long; stamens 7-8. 82. H. angustifolia.
88. Flowers 4.o-6.0 mm long; calyx lobes 1.5-4.0 mm
long; stamens 3-6.
89. Young branches and lower part of inflorescence
rachis hispid-setose.
90. Leaves 2.5-6.5 cm long; inflorescence 3.0-5.0 cm
long. 71. H. pimichina.
90. Leaves 6.0-15.0 cm long; inflorescence 6.o-
28.0 cm long.
9I. Leaves entirely glabrous beneath, the dried
material with silver metallic sheen.
72. H. subscandens.
9I. Leaves hirsute beneath, especially on principal
venation, the dried material without silver
metallic sheen.
92. Bracteoles oblong, broad towards base,
chartaceous, with short-stalked glands, and
apex terminating with a short-stalked gland.
25. H. paniculata.
92. Bracteoles linear-lanceolate, coriaceous,
with long-stalked glands, and apex always
terminating with a long-stalked gland.
73. H. hispidula.
89. Young branches tomentellous to puberulous; lower
part of inflorescences glabrous to puberulous or
tomentellous.
93. Inflorescences tomentellous, 4.5-27.0 cm long;
leaves 10.0-22.0 cm long, broadest above middle;
pedicels 4.0-9.0 cm long. 74. H. silicea.
270 Flora Neotropica

93. Inflorescences glabrescent or very sparsely setose,

3.0-I5.0 cm long; leaves 4.5-10.0 cm long,
broadest below middle; pedicels 5.0-I5.0 mm
long.
94. Stamens 3; calyx glandular. 75. H. excelsa.
94. Stamens 5-6; calyx eglandular.
95. Leaf apex gradually tapered to apex from
near base; pedicels 4.0-5.0 mm long.
76. H. adenophora.
95. Leaf apex cuspidate; pedicels 8.0-15.0 mm
long. 77. H. caduca.
6I. Bracts and pedicels eglandular.
96. Inflorescence clustered, fasciculate; leaves slightly bullate, 8.0-12.0 cm long;
ovary inserted near base of receptacle. 78. H. fasciculata.
96. Inflorescence of elongate racemes; leaves usually plane not bullate, if bullate
then small; ovary inserted at or near mouth of receptacle.
97. Receptacle elongate cylindrical; calyx lobes glandular.
98. Inflorescences densely tomentose; leaves I0.O-15.0 cm long; flowers ca
1.5 cm long. 79. H. couepiflora.
98. Inflorescences sparsely puberulous; leaves to 8.5 cm long; flowers to
I.o cm long. 80. H. tubiflora.
97. Receptacle campanulate; calyx lobes eglandular.
99. Leaves orbicular, predominantly retuse to rounded at apex; pedicels
7.0-20.0 mm long. 68. H. longipedicellata.
99. Leaves oblong-lanceolate to ovate, acuminate at apex; pedicels to 15.0
mm long, usually shorter.
1oo. Lower surface of leaf hirsute, pilose or hispid at least on primary and
secondary venation.
1o. Leaves oblong, oblong-lanceolate, or lanceolate, the venation
sometimes deeply impressed above; fertile stamens 7-9.
102. Leaves oblong, 3.5-7.5 cm long, venation deeply impressed
above; flowers 4.0 mm long. 81. H. floribunda.
102. Leaves lanceolate to oblong-lanceolate, 5.0-15.0 cm long;
venation level above; flowers ca 6.0 mm long. 82. H. angustifolia.
o10. Leaves ovate to elliptic, the venation usually prominent above
rarely slightly impressed but leaves broadly ovate; fertile stamens
3-9.
o03. Leaf venations slightly impressed above; leaves broadest near
base, 2.5-8.0 cm long, subcordate at base; inflorescence densely
crowded, with short rachis. 83. H. rugosa.
103. Leaf venation level to slightly prominent above; leaves broadest
at or above middle, subcuneate to rounded at base; inflores-
cence lax with longer rachis.
I04. Stamens 3. 35. H. mutisii.
104. Stamens 5-7.
I05. Leaf apex mucronate to abruptly acuminate; pedicels
2.0-3.0 mm long. 84. H. scaberula.
o05. Leaf apex acute to acuminate, but never mucronate or
abruptly acuminate; pedicels 4.0-10.0 mm long.
0o6. Inflorescence and exterior of flowers densely tomen-
tose; flowers 5.0-8.0 mm long 85. H. hebeclada.
0o6. Inflorescence and exterior of flowers puberulous,
flowers 4.0-5.0 mm long. 56. H. racemosa.
Ioo. Lower surface of leaf glabrous or with few sparse appressed hairs on
midrib and primary veins.
107. Leaves narrowly lanceolate, 5.5-I8.0 cm long; 0.5-2.2 cm broad.
5I. H. angustissima.
107. Leaves ovate to oblong.
o18. Fertile stamens 7-9; flowers 6.o-Io.o mm long.
I09. Leaf bases subcordate; calyx lobes 4.0-6.o mm long; young
branches sparsely hispid. 82. H. angustifolia.
o19. Leaf bases subcuneate to rounded; calyx lobes 1.5-2.5 mm
long; young branches puberulous. 86. H. enneandra.
Hirtella 271

io8. Fertile stamens 3-6; flowers 4.0-5.0 mm long.

I I. Fertile stamens 3; inflorescence either 2-3 flowered or
densely many-flowered and tomentose; filaments only
slightly exceeding calyx lobes.
I I. Leaves 7.0-13.0 cm long, 3.0-5.0 cm broad, chartaceous;
inflorescence 2-3 flowered. 87. H. pauciflora.
I I. Leaves 3.5-5.5 cm long, 1.5-2.7 cm broad, thick-coria-
ceous; inflorescence many-flowered. 88. H. glaziovii.
Io. Fertile stamens 4-6; inflorescence puberulous to tomentel-
lous, many-flowered; filaments far exceeding calyx lobes.
1I2. Leaves thick-membraneous; inflorescence 2.0-3.0 mm
long; fertile stamens 4. 4I. H. rodriguesii.
I 2. Leaves thick-coriaceous; inflorescence 4.0-15.0 cm long;
fertile stamens 5-7. 56. H. racemosa.

i. Hirtella sect Myrmecophila

This small section of closely related species is easily recognized by the ant-
inhabited cavities at the base of the leaves. The several species, although very closely
related, are nevertheless quite distinct.

I. Hirtella myrmecophila Pilger, Verh. Bot. Ver. Brand. 47: 148. 1905;
Suesseng., Fedde Repert. Nov. Sp. 39: 15. I935.
var. tetrandraHuber, Bol. Mus. Emilio Goeldi 5: 369. I909. Type. Ducke
Hirtella myrmecophila
MG 8557, Brazil, Para, fl. (holotype, MG; isotypes, BM, P, US).

Small tree to 7.0 m tall, the young branches sparsely hispid. Leaves coriaceous,
oblong to oblong-lanceolate, 9.0-19.0 cm long, 4.o-7.0(-8.5) cm broad, acuminate
at apex, with acumen 6.0-I 7.0 mm long, bearing swollen ant cavities at base, slightly
bullate beneath, with sparse hispid pubescence on principal venation; primary veins
I2-I4 pairs, prominent above; midrib prominent above. Stipules filamentous, to
12.0 mm long, subpersistent, hispid. Inflorescences fasciculate racemes, 1.0-3.5 cm
long, the rachis glabrescent with a few hispid hairs. Bracts and bracteoles linear, to
3.5 mm long, usually with paired sessile basal glands, hispid. Flowers ca 4.0 mm long.
Receptacle campanulate, sparsely hispid on exterior, glabrous within except for
deflexed hairs at throat; pedicels 4.0-5.0 mm long. Calyx lobes acute, sparsely hispid
on exterior, puberulous within. Petals 5, glabrous. Stamens 4-5, unilateral, the
filaments far exceeding the calyx lobes. Style glabrous except for hirsute base. Ovary
inserted midway up receptacle tube, pilose. Fruit not seen.
TYPE. Ule 6916, Brazil, Amazonas, fl (holotype, B, lost; lectotype, HBG).
DISTRIBUTION. Non-flooded forest in central Amazonian Brazil.
BRAZIL. Amazonas: CoelhoINPA i837 fl (INPA, NY); Coelho& ChagasINPA 4222 fl (INPA,
MG, NY); Ducke385 fl (A, F, NY, R, S, US); 2277 fl (MG, R, RB, SP); MG 7153 fl (MG); RB 25008
fl (K, P, S); Fr6es22544 fl (IAN); 28358 fl (IAN); 28951 fl (IAN); KuhlmannRB I7977 fl (K, RB, S);
Oliveira2701 fl (IAN); 2833 fl (IAN); Pranceet al 2201 fl (INPA, MG, NY); 4984 fl (INPA, NY);
Rodrigueset al 4552 fl (INPA I3052); 7233 fl (INPA I6136, NY); 8249 fl (INPA I7639, NY). Terr.
Roraima: KuhlmannRB 2972 fl (RB); Luetzelburg21380 fl (M, NY, R); 21381 fl (R). Para: DuckeMG
16459 fl (MG).

This species differs from H. physophoraprincipally in the smaller, narrower,

272 Flora Neotropica

thicker leaves with a slightly bullate undersurface, in the fewer stamens, and in the
tendency for a lower insertion of the ovary.
Huber's variety, var tetrandra Hub. distinguished by having 4 instead of 5 stamens,
cannot be maintained as the type has some flowers with 5 (not 4) stamens, and this
hardly offers a valid basis for recognizing a variety. There are no other differences to
distinguish var tetrandra.

2. Hirtella physophora Martius & Zuccarini, Abh. Akad. Minchen :374.

1832; Zucc., Flora i5(Beibl. 2): 79. 1832; Hook. f., Mart. Fl. Bras. 14(2):
29. 1867; Macbride, Fl. Peru 1076. 1938.
Hirtella caulifloraHuber, Bol. Mus. Emilio Goeldi 6: 73. I9I0, synon nov. Type. DuckeMG 4745,
Brazil, Terr. Amapa, fl (holotype, MG; isotypes, BM, K).
Hirtella cotticaensisKleinhoonte, Rec. Trav. Bot. N6erl. 22: 386. 1925; Kleinh. in Pulle, Fl.
Suriname 2: 445. 1937, synon nov. Types. GonggrijpBW 5333, Suriname, fl (holotype, U;
isotype, NY); Gonggrijp& StahelBW 5666, st (paratype, U).

Small tree to 6.0 m tall, the young stems sparsely hispid, lenticellate. Leaves
membraneous, oblong-elliptic to oblong, I7.o-30.0 cm long, 6.o-i I.o cm broad,
acuminate to mucronate at apex, with acumen 5.0-20.0 mm long, bearing swollen
ant cavities at base, hirsute on venation beneath; midrib prominent above; primary
veins 13-16 pairs, prominent beneath. Stipules filamentous, to I.5 cm long, sub-
persistent, hispid. Inflorescences fasciculate racemes, 2.0-4.0 cm long, hispid. Bracts
and bracteoles linear to filamentous, 2.0-8.0 mm long, hispid, persistent. Flowers
5.0-7.0 mm long. Receptacle campanulate, hispid on exterior, glabrous within
except for deflexed hairs at throat; pedicels 3.0-4.0 mm long. Calyx lobes oblong-
lanceolate, hispid on exterior, puberulous within near apex, glabrous beneath.
Petals 5, glabrous, twice as long as calyx lobes. Stamens 6, unilateral, the filaments
far exceeding calyx lobes. Style hirsute on lower portion up to half of its length. Ovary
inserted at mouth of receptacle, pilose. Fruit ellipsoid; epicarp smooth, glabrous;
mesocarp thin, fleshy; endocarp thin, hard, bony.
TYPE. Martiussn, Brazil, Amazonas, fl (holotype, M).
DISTRIBUTION. Non-flooded forest in the Guianas and Amazonia.
COLOMBIA. Vaupes: Ferndndez1950 fl (COL, US). VENEZUELA. Amazonas: Cowan &
Wurdack31439 fl (F, MG, NY, US); Croizat o108 fl (GH, NY, RB); Maguire & Politi 28623 fl (COL,
GH, LE, NY); Maguire et al 31822 fl (NY). PERU. Loreto: Klug 989 fl (F, NY, US). GUYANA.
Guppy250 (F.D. 7226) fl (NY). SURINAME. B.W. 6834 st (K); Irwin, Prance,et al 55520 fl (F, GH,
NY, US); Maguireet al 24103 fl (A, K, NY, US); 54257 fl (NY, US); WesselsBoer 1284 fl (U, US).
FRENCH GUIANA. Black et al 54-17631 St (IAN). BRAZIL. Amazonas: ChagasINPA 882 fl (INPA,
NY); Coelho& Mello INPA 3531 fl (INPA, NY); DuckeRB 23587 fr (K, P, RB, S, US); RB 25003 fl
(RB, US); 25004 fl (RB, US); Krukoff6874 fl (A, BM, BR, F, K, MICH, NY, RB, S, US); 8190 fl
(A, BR, F, LE, NY, P, S); Maguire,Steyermark& Maguire6o4ioA fl (NY); Mello INPA 2005 fl (INPA,
NY); Rodrigues5403 fl (INPA 14072, NY); 5427 fl (INPA 14096, NY); Sprucesn fl (LE, NY); 1438 fl
(K, P). Terr. Roraima: Pranceel al 10156 fl (INPA, NY). Para: Ducke RB 18242 fl (RB, S). Terr.
AmapA; Froes26766 fl (IAN, UB); Irwin & Westra47177 fl (GH, MICH, MG, NY, R, RB, US).

LOCAL NAME. Guyana: Fulishakuyefu.

Hirtella cotticaensis was separated from H. physophoramainly by the larger in-
florescence and smaller flowers. There is much variation in these characters, and they
certainly do not separate two species. Hirtellacauliflorais meant to differ mainly in
the cauliflorous habit. However, this is frequently seen in H. physophora,and I have
observed trees in the field in which the same individual bears both cauliflorous and
axillary flowers.
Hirtella 273

3. Hirtella vesiculosa Suessenguth, Fedde Repert. Nov. Sp. 42: 43. I937.

Tree to 8.0 m tall; the young branches hirsute. Leaves coriaceous, oblong-
lanceolate, I3.5-24.0 cm long, 2.5-6.0 cm broad, acuminate at apex, with acumen
5.0-10.0 cm long, bearing swollen ant cavities at base, hirsute on venation beneath;
midrib prominulous above; primary veins I5-20 pairs, prominent beneath. Stipules
filamentous, to 8.0 mm long, hispid, caducous. Inflorescences elongate racemose
panicles 8.0-I6.o cm long, branched slightly at base only, the rachis and branches
hirsute. Bracts and bracteoles linear, to 8.0 mm long, persistent, hirsute. Flowers
5.0-6.0 mm long. Receptacle campanulate, hispid on exterior, glabrous within
except for deflexed hairs at throat; pedicels 6.0-7.0 mm long. Calyx lobes ovate,
hispid on exterior, puberulous within near apex. Petals 5, exceeding calyx lobes,
glabrous. Stamens 6, the filaments far exceeding calyx lobes. Style hirsute at base only.
Ovary inserted at mouth of receptacle, pilose. Fruit not seen.
TYPES. Luetzelburg 22245, Colombia, Guainia, fl (holotype, M); 22226 st (para-
type, M).
DISTRIBUTION.
Known only from the type gatherings from San Fellipe on the
Rio Negro.
This species is closely related to H. guainiae,and thus far seems to differ only in
the inflorescence, which has two branches in the one specimen examined, and in the
narrower more lanceolate leaves. Further material is needed to determine if the
present species is distinct from H. guainiae.

4. Hirtella dorvalii Prance, sp nov Fig 43 D-G.

Frutex, ramulis juvenilibus hirsutis. Folia alternata, petiolata; petiolo 2.0-4.0

mm longo, eglanduloso, piloso, tereti; laminae ovatae vel oblongo-ovatae, coriaceae,
leviter bullatae, 8.5-I4.0 cm longae, 5.5-7.0 cm latae, apice in acuminem I.0-3.0
mm longum contractae, basi subcordatae, cum duobus vesiculis formicariis munitae;
costa media utrinque prominente, subtus hirsuta; costis secundariis 9-12 jugis,
subtus prominentibus, hirsutis. Stipulae filamentosae, circa 6.0 mm longae, hirsutae,
caducae, eglandulosae. Flores 4.5-6.0 mm longi, in paniculis elongatis 14.0-25.0 cm
longis dispositi, rachi ramisque hispido-pilosis. Bracteae bracteolaeque oblongae,
2.0-7.0 mm longae, persistentes, glandulis stipitatis munitae. Receptaculum cam-
panulatum, extus hispidum, intus basim versus glabrum; pedicelli 2.0-3.0 mm longi.
Calycis-lobi acuti, extus pilosi, intus puberuli. Petala 5, glabra, purpurea, calycis-
lobos excedentia. Stamina 6-7, unilateralia, filamenta glabra, longe exserta. Ovarium
ad faucem receptaculi insertum, pilosum. Stylus e basi ovarii ortus, basim versus
hirsutus. Fructus mihi ignoti.
TYPE. Ducke 1330, Brazil, Terr. Roraima, Caracarai, Rio Branco, fl (holotype,
NY; isotypes, A, IAN, MG, R, RB, US).
Known only from the type gathering from forest at a savanna
DISTRIBUTION.
margin.
This species is the most distinct within the section because of the branched
inflorescence and the bracts bearing stipitate glands. In H. dorvaliithe inflorescence
has many short branches attached to a central rachis, but in H. vesiculosawhich also
has a branched inflorescence, the branches are few and occur at the base only.
This species is named in honor of Dr. Dorval Magalhaes, Agronomist of Boa
Vista, Terr. Roraima, who has given me much help with field work in the territory,
and who also collected with Drs. Black and Ducke. His knowledge of the region and
of the flora has been a great help to several collectors who have worked in this area.
274 Flora Neotropica

'7
B C '1

Loker F

FIG 43. Speciesof Hirtella.A-C, H. araguariensis(Pireset al 5I537); A, habit, x o.5; B, flower,

x 5; C, flower section, x 5. D-G, H. dorvalii(DuckeI330); D, habit, x o.5; E, flower section, x 5;
F, flower, x 5; G, leaf basewith myrmecophilousswellings,x i .5.
Hirtella 275

5. Hirtella guainiae Spruce ex Hooker f., Mart. Fl. Bras. 14(2): 3I, t. IO.
1867; Suesseng., Fedde Repert. Nov. Sp. 42: 43. I937; Macbride, Fl.
Peru IO76. I938.

Shrub or small tree to 6.0 m tall, the young branches hispid-hirsute.Leaves thin-
coriaceous, oblong, I4.0-23.0 cm long, 5.5-9.5 cm broad, acuminate at apex, with
acumen 2.0-I8.0 cm long, bearing swollen ant cavities at base, hirsute on venation
beneath; midrib prominulous above; primary veins II-I6 pairs, prominent and
hirsute beneath. Stipules filamentous, to 2.0 cm long, hispid, subpersistent. Inflores-
cences elongate racemes, I5.0-30.0 cm long, the rachis hirsutulous pubescent.
Bracts and bracteoles lanceolate, 4.0-6.0 mm long, with a broad usually glandular
base. Flowers 5.0-6.0 mm long. Receptacle campanulate, hirsutulous-puberulous
on exterior, glabrous within except for deflexed hairs at throat; pedicels 3.0-8.0 mm
long. Calyx lobes oblong-ovate, hirsutulous on exterior, puberulous within, Petals 5,
purple, exceeding calyx lobes. Stamens 4-5, unilateral, the filaments far exceeding
calyx lobes. Style sparsely hirsute up to half its length. Ovary inserted near mouth of
receptacle. Fruit not seen.
TYPE.Spruce3523, Venezuela, Terr. Amazonas, fl (holotype, K; isotypes, BM,
BR, P).
DISTRIBUTION.
Non-flooded forest in western Amazonia.
COLOMBIA. Guainia: Schultes,Baker & Cabrera18059 fl (NY, US). Vaup6s: Schult,s & Cabrera
14522 fl (BR, COL, GH, US); Ferndndez2269 fl (COL, US). VENEZUELA. Terr. Amazonas: Maguire,
Wurdack& Maguire 41614 fl (NY). PERU. Loreto: Asplund14163 fl (BR, K, LD, NY, P, R, US);
DuckeRB 17975 fl (RB, S); Killip & Smith29947 st (F, US); Klug 27 fl (F, NY, US); 512 fl (F, NY, US).
BRAZIL. Amazonas: DuckeRB 25013 fl (RB). Acre: Pranceet al 3006 fl (INPA, NY). Terr. Rond6nia:
Pranceet al 6008 fl (INPA, NY).

Although Hooker in the original description of this species gave the number of
stamens as 3, it is in fact 4-5 in the type and all other material I have examined.

6. Hirtella duckei Huber, Bol. Mus. Emilio Goeldi 6: 74. I9IO.

HirtellaformicariaSuessenguth, Fedde Repert. Nov. Sp. 39: I4. 1935. Type. Krukoff4557, Brazil,
Amazonas, fl (isotypes, A, BM, F, K, LE, M, MICH, NY, S, US).

Shrub, the young branches hispid, lenticellate. Leaves oblong to elliptic, thin-
coriaceous, (I5.o-)20.0-30.0 cm long, 6.0-I I.0 cm broad, acuminate at apex, with
acumen 5.0-20.0 mm long, bearing swollen ant cavities at base, hirsute on venation
beneath; midrib prominulous above; primary veins 11-18 pairs, prominent and
hirsute beneath. Stipules filamentous, to i.o mm long, hispid, persistent. Inflores-
cences elongate racemes, 2.0-I2.0 cm long, the rachis hispid. Bracts and bracteoles
2.0-3.0 mm long, linear, hirsute, subpersistent,eglandular. Flowers 5.0-6.0 mm long.
Receptacle campanulate, hispid on exterior, glabrous within except for deflexed
hairs at throat; pedicels 4.0-6.0 mm long. Calyx lobes oblong, hispid on exterior,
puberulous within. Petals 5, pale purple, exceeding calyx lobes. Stamens 5-8,
unilateral, the filaments far exceeding calyx lobes. Style hirsute at base only. Ovary
pilose, inserted at or near mouth of receptacle. Fruit not seen.
TYPE.DuckeMG 6756, Brazil, Amazonas, fl (holotype, MG; isotypes, BM, P,
RB, US).
DISTRIBUTION.Non-flooded forest in Guyana and central and western Amazonia.
COLOMBIA. Vaupds: Schultes& Ldpez94ooB fl (US). GUYANA. A. C. Smith2999 fl (A, F, K,
NY, P, US). PERU. Loreto: Ducke MG 7598 st (MG). BRAZIL. Amazonas: Baldwin 3200 fl (IAN,
US); Cardona1370 fl (VEN); Ducke RB 25009 fl (K, P, RB, S, US); Frdes 28714 fl (IAN); Krukoff
276 Flora Neotropica

6834 fl (A, BM, BR, F, K, LE, MICH, NY, RB, S, US); Pranceet al 3320 fl (INPA, MG, NY). BOLI-
VIA. Pando: Pranceet al 8521 fl (INPA, NY).

This species is most closely related to H. guainiae,from which it differs in the

hispid flowers, the shorter inflorescence, and the smaller eglandular bracts. As the
types of H. duckeiand H. formicariamatch well, the latter cannot be regarded as a
distinct species.

2. Hirtella sect Hirtella

7. Hirtella macrosepala Sandwith, Kew Bull. 1939: 549. I940.

Tree to I8.o m tall, the young branches glabrous. Leaves oblong, coriaceous,
6.5-I7.0 cm long, 3.0-8.0 cm broad, cuspidate at apex, with acumen 2.0-6.o mm
long, rounded at base, glabrous beneath; primary veins 7-IO pairs, prominulous on
both surfaces; midrib prominulous above, prominent beneath; petioles 3.0-5.0 mm
long, glabrous, terete, eglandular. Stipules lanceolate, 2.0-5.0 mm long, glabrous
when mature, eglandular, persistent. Inflorescences axillary and terminal panicles
2.5-5.0 cm long, usually subcorymbose, the rachis and branches glabrous or glabres-
cent. Bracts and bracteoles ca 2.0 mm long, ovate, glabrescent, persistent, the apex
with I or 2 sessile glands. Flowers 6.o-8.o mm long. Receptacle campanulate,
glabrous or glabrescent on exterior, glabrous within except at throat; pedicels 2.0-
8.o mm long. Calyx lobes rounded, glabrous or glabrescent on exterior, tomentellous
within, the margins eglandular. Petals 5, white, glabrous. Stamens 6-7, unilateral,
the filaments glabrous, connate at base, far exceeding calyx lobes. Style hirsute up to
half its length. Ovary inserted near mouth of receptacle, villous. Fruit not seen.
TYPE. Davis 590 (F.D. 2631), Guyana, fl (holotype, K).
DISTRIBUTION. Primary forest in eastern Venezuela and the Guianas. Flowering
September to April.
VENEZUELA. Terr. Delta Amacuro: Berti375 fl (F, NY, US). Bolivar: Blanco59r fl (NY, VEN).
GUYANA. CAP 35 (F.D. 5344) fl (A, F, K, NY, P, US). FRENCH GUIANA. For. Serv. 7237 fl
(NY); 7259 fl (NY).

LOCALNAMES.Venezuela: Pica-pica, Pica-pica rosada. Guyana: Bokobokotokon.

8. Hirtella ulei Pilger, Verh. Bot. Ver. Brand. 47: I48. 1905.

Shrub to 4.0 m tall, the young branches glabrous becoming lenticellate with age.
Leaves coriaceous, ovate-elliptic, 6.5-13.0 cm long, 4.0-7.0 cm broad, acuminate at
apex, with acumen 2.0-6.0 mm long, rounded to subcordate at base, glabrous
beneath, with palisade glands frequent towards base; primary veins 8-12 pairs,
prominulous on both surfaces; midrib prominulous above, glabrous and prominent
beneath; petioles 3.0-7.0 mm long, puberulous, becoming glabrous with age,
eglandular, terete. Stipules I.0-2.0 mm long, deltoid, intrapetiolar, puberulous,
eglandular, persistent. Inflorescences terminal panicles IO0.-45.0 cm long, the rachis
and branches puberulous to glabrescent. Bracts and bracteoles 1.5-4.0 mm long,
ovate, persistent, puberulous, with numerous stipitate glands. Flowers 5.0-7.0 mm
Hirtella 277

long, Receptacle campanulate, puberulous on exterior, glabrous within except for

reflexed hairs at throat; pedicels 1.0-2.0 mm long. Calyx lobes acute, with small
stipitate glands on margins, glabrescent on exterior, pubescent within. Petals 5, pink,
glabrous. Stamens 6-7, unilateral, the filaments far exceeding calyx lobes. Style
hirsute on lower portion up to one third its length. Ovary inserted at mouth of
receptacle, pilose. Fruit oblong; epicarp glabrous; mesocarp very thin, fleshy;
endocarp thin, hard, bony.
TYPES. Ule 6150, Brazil, Amazonas, fl (holotype, B, lost; lectotype, HBG;
isotype, MG).
DISTRIBUTION. Sandy margins and islands of rivers, and savanna margins in the
upper Orinoco region of Venezuela and the Rio Negro region of Brazil. Flowering
throughout the year.
VENEZUELA. Terr. Amazonas: Licky I fl fr (VEN); Maguire29293 fr (NY); 29317 fr (NY);
Maguire, Wurdack& Keith 41687 fl (NY); Maguire & Wurdack34685 fl (NY); Steyermark 57825 fl (US);
Tate 287 fl fr (NY, US); Wurdack& Adderley43246 fl (NY). BRAZIL. Amazonas: Ducke 1745 fl (A, F,
IAN, K, MG, NY, R, US); MG 11194 fl (MG); MG 11530 fl (MG); RB 15079 fl (K, RB, S, US);
Frdes24909 fl (IAN); 28078 fl (IAN); 29207 fl (IAN); Pires 716 fl (IAN); Rodrigues& Coelho3291 fl fr
(INPA 9840, NY); Rodrigues& Lima 2053 fl (INPA 9397); Ule 8853 fl (K).

9. Hirtella glabrata Pilger, Notizbl. Bot. Gart. Berlin 6: I38. I914.

Shrub or small tree to 8.o m tall, the young branches tomentellous, soon becom-
ing glabrous and lenticellate. Leaves coriaceous, oblong-elliptic to elliptic, 6.5-
13.0 cm long, 4.0-6.2 cm broad, acute to acuminate at apex, with acumen 4.0-5.0
mm long, rounded to subcordate at base, the lower surface with few sparse appressed
hairs, with palisade glands frequent on lower portion; primary veins 9-12 pairs,
prominulous on both surfaces; midrib prominulous above, prominent and puberu-
lous beneath; petioles 4.0-6.0 mm long, tomentellous when young, terete, eglandular.
Stipules to 2.0 mm long, deltoid, persistent, intrapetiolar, eglandular. Inflorescences
terminal panicles I0.0-25.0 cm long, the rachis and branches velutinous-tomentel-
lous. Bracts and bracteoles 2.5-4.5 mm long, ovate, persistent, with numerous to few
large stipitate glands, or rarely the glands mostly sessile. Flowers 5.0-7.0 mm long.
Receptacle campanulate, sparsely tomentellous on exterior, glabrous within except
for deflexed hairs at throat; pedicels 0.5-2.0 mm long. Calyx lobes acute, eglandular,
sparsely tomentellous on exterior, gray-pubescent within. Petals 5, glabrous. Stamens
6-7, unilateral with short staminodes opposite them, the filaments far exceeding calyx
lobes. Style hirsute for half its length. Ovary inserted at mouth of receptacle, tomentel-
lous. Fruit ellipsoid; epicarp smooth, glabrous; mesocarp thin, fleshy; endocarp thin,
hard, bony, hirsute within.
TYPE. Ule 8984, Brazil, Amazonas, fl (holotype, B, lost; lectotype, K; isotypes,
L, US).
DISTRIBUTION. Open forest, savannas, and secondary forest in central Brazilian
Amazonia. Flowering April to September.
BRAZIL. Amazonas: CoelhoINPA I714 fl fr (IAN, INPA, MG, NY); Ducke465 fl (A, F, IAN, K,
MG, NY, R, RB, S, US); RB 23586 fl (INPA, K, P, RB, S, US); RB 25012 fl (US); RB 35580 fl
(K, US); Mello INPA 1632 fl (IAN, INPA, MG, NY); Mello & CoelhoINPA 3941 fl (IAN, INPA, MG,
NY). Pari: Ducke MG 8490 fl (INPA, MG); MG 11630 fl (MG); MG 11939 fl (MG); MG I5078 fl
(RB 15078, US).

This species is close to H. ulei which it replaces east of Manuas. It differs from
H. ulei in the eglandular calyx lobes, the larger bract glands, and in the more pubes-
cent inflorescence and leaf undersurface. Hirtella glabrata is also close to H. macrophylla,
278 Flora Neotropica

but differs in the short dense tomentellous pubescence of the inflorescence, in the
almost glabrous leaf underside, and in the shorter-stalked, larger bract glands. In
H. macrophylla the glands arise from the stem, rather than from the bracts, and the
glands have longer stalks with a small gland at the apex appearing clavate.

o. Hirtella carbonaria Little, Jour. Wash. Acad. Sci. 38: 88. I948.

Medium sized tree to Io.o m tall, the young branches sparsely hirsutulous, soon
becoming glabrous and lenticellate. Leaves coriaceous, elliptic to ovate-elliptic,
4.5-9.5 cm long, 2.5-4.5 cm broad, acuminate at apex, with acumen 4.0-Io.o mm
long, rounded at base, glabrous beneath except for a few appressed hairs; primary
veins 6-9 pairs, prominulous on both surfaces; midrib prominent on both surfaces,
sparsely hirsutulousbeneath. Stipules linear, puberulous, eglandular, subpersistent,
2.0-4.0 mm long. Inflorescences terminal and axillary panicles, 3.0-9.0 cm long, the
rachis and branches glabrescent, with a few stiff appressedhairs. Bracts and bracteoles
I.0-3.0 mm long, ovate, hirsutulous, persistent; solitary glands on a long stalk,
I.0-2.0 mm long, arising from the junction of the pedicel and branch or from pedicel.
Flowers 4.0-5.0 mm long. Receptacle campanulate, glabrous on exterior, glabrous
within. Calyx lobes rounded to acute, eglandular, glabrous on exterior, gray-
pubescent within. Petals 5, white, glabrous. Stamens 3, unilateral, with three short
filamentous staminodes opposite them; filaments far exceeding calyx lobes. Style
hirsute on lower portion. Ovary inserted near mouth of receptacle, tomentose. Fruit
ellipsoid; epicarp smooth, glabrous when mature; mesocarp very thin, fleshy;
endocarp thin, bony, densely hirsutulous within.
TYPE.Little6275,Ecuador, Esmeraldas,flfr (holotype, US; isotypes, A, F, K, MAD,
NY).
DISTRIBUTION. Coastal thickets and margins of mangrove swamps in areas
periodically flooded by sea water, on the Pacific coast of Ecuador and southern
Colombia. Flowering throughout the year.
COLOMBIA. Choc6: Fuchs21852 fl (NY, US); Haught5418 fl (COL, F, US); Killip & Cautre-
casas39130 fl (COL, F, NY, P, US); VonSneidern4905 fl (S); 4914 fl (S). Valle: Cuatrecasas14322 fl fr
(COL, F, US); 15926 fr (US); 15929 fr (COL, F); 17692 fl (F, P); 19908 fl (COL, F, P); Killip 11702
38965 fl (COL, F, GH, NY, S, US). Cauca: LehmannB. T. 646
fl fr (GH, NY, US); Killip & Cuatrecasas
fl (NY). Nariiio: Barclay855 (BM); RomeroC. 2211 fl (COL); 3149 fl (COL); Schulz & Rodriguez498 fl
(U). Without precise locality: Seemannsn fl (GH, GOET). ECUADOR. Esmeraldas: Jdtiva & Epling
712 fl (LA, NY).

LOCALNAME.Ecuador: Carbonero.

1. Hirtella araguariensis Prance, sp nov Fig 43 A-C.

Arbor 8.0 m alta, ramulis juvenilibus hispidis mox glabris. Folia alternata
petiolata; petiolo 2.0-3.0 mm longo, tereti, eglanduloso, juveniliter hispido; laminae
oblongo-lanceolatae, subcoriaceae, I0.0-7.0 cm longae, 3.5-5.3 cm latae, apice in
acuminem 5.0-I2.0 mm longum contractae, basi subcordatae; costa media utrinque
prominente, glabra; costis secundariis 10-13 jugis, utrinque leviter prominentibus.
Stipulae lineares ad 8.0 mm longae, pubescentes, eglandulosae, persistentes. Flores
5.0-7.0 mm longi, in paniculis IO.O-25.0 cm longis dispositi, ramis brevissimis I-3
floris gerentibus, rachi ramisque piloso-tomentellis. Bracteae bracteolaeque 3.0-
Hirtella 279

6.0 mm longae, oblongae, acutae, tomentellae, persistentes, marginibus eglandulosae

vel cum glandulis sessilibus munitae. Receptaculum campanulatum, extus tomen-
tellum, intus basim versus glabrum; pedicelli 0.5-I.0 mm longi. Calycis-lobi acuti,
extus tomentosi, intus puberuli, marginibus eglandulosis. Petala 5, alba, glabra.
Stamina 5, unilateralia; filamenta basim versus hirsuta, longe exserta. Ovarium ad
faucem receptaculi insertum, pilosum. Stylus e basi ovarii ortus, hirsutus. Fructus
mihi ignoti.
TYPE. Pires, Rodrigues & Irvine 51537, Brazil, Terr. Amapa, Rio Araguari, fl
(holotype, NY; isotypes, IAN, INPA, MG).
DISTRIBUTION. Known only from the type gathering, collected in upland forest,
and flowering in October.
In this distinct species, the inflorescence suggests H. eriandra, but that species is
very different in leaf shape.

12. Hirtella cordifolia Prance, sp nov Fig 44 D-F.

Frutex parva, ramulis juvenilibus puberulis mox glabris lenticellatis. Folia

alternata petiolata; petiolo circa i.o mm longo, tereti, eglanduloso, juvenili hirsuto;
laminae cordiformi-ovatae, coriaceae, 3.5-5.5 cm longae, 1.8-2.9 cm latae, apice in
acuminem 2.0-7.0 mm longum contractae, basi cordatae, utrinque praeter costam
glabrae; costa media supra leviter prominente, hirsuta; costis secundariis 7-9 jugis,
utrinque leviter prominentibus. Stipulae lineares, ad 3.0 mm longum, sparse pubes-
centes, eglandulosae, persistentes. Flores circa 6.o mm longi, in paniculis paucira-
mosis vel racemis simplicibus 2.5-5.5 mm longi dispositi, rachi ramisque hirsuto-
puberulis. Bracteae bracteolaeque 1.5-3.0 mm longae, acutae, persistentes, puberu-
lae, marginibus cum glandulis stipitatis munitae. Receptaculum campanulatum,
extus hirsutum, intus basim versus glabrum; pedicelli 3.0-6.0 mm longi. Calycis-
lobi acuti, extus sparse hirsuti, intus griseo-pubescenti. Petala 5, glabra, rosea.
Stamina 4-5, unilateralia, staminodiis dentiformis opposita; filamenta glabra, longe
exserta, basi connata. Ovarium ad medium receptaculi insertum, sparse pilosum.
Stylus e basi ovarii ortus, hirsutus. Drupa ellipsoidea, epicarpio laevi, glabro;
mesocarpio tenue, carnoso; endocarpio duro, tenue, osseo, intus hirsuto.
TYPE. Wurdack & Adderley43621, Venezuela, Terr. Amazonas, Rio Hechimoni,
Sabana Hechimoni, fl (holotype, NY).
DISTRIBUTION. Collected only from savannas of the Rio Hechimoni, Venezuela,
Flowering in July.
VENEZUELA. Terr. Amazonas: Rio Hechimoni, Sabana Hechimoni, Maguire, Wurdack&
Bunting37665 fl (NY).

This species is most closely related to H. deflexa, but differs in the smaller leaves
with fewer primary veins which are less prominent beneath, and in the glabrous
lower surface of the leaves. It differs from H. paniculata in the glabrous leaves with
cordate bases, and in the little-branched inflorescence.

13. Hirtella insignis Briquet ex Prance, sp nov Fig 45 A-D.

Arbor ad I5.0 m alta, ramulis juvenilibus sparse hispidis mox glabris lenticel-
latis. Folia alternata, petiolata; petiolo 3.o-5.0 mm longo, tereti, eglanduloso,
280 Flora Neotropica

C'

i X

A.. \

F//

FIG 44. Speciesof Hirtella.A-C, H. piresii(Blacket al 52-14653, Rodrigues et al 3168); A, habit

x 0.5; B, flower, x 5; C, flowersection, x 5. D-F, H. cordifolia et al 43621); D, habit, x o.5; E,
(Wurdack
flower, x 5; F, flowersection, x 5; G, bract, x 7.5.
Hirtella 281

maturitate glabro; laminae oblongae, coriaceae, 7.0-I7.0 cm longae, 3.6-6.0 cm

latae, apice in acuminem 2.0-7.0 mm longum contractae, basi rotundatae vel sub-
cuneatae, subtus fere glabrae et cum pilis longis appressis, basim versus glandulosae;
costa media supra leviter prominente, subtus prominente leviter hispida; costis
secundariis 10-I5 jugis, utrinque leviter prominentibus. Stipulae lineares, parvae.
Flores 5.0-6.0 mm longi, in paniculis terminalibus multiramosis 6.0-7.0 cm longis
dispositi, rachi ramisque glabris. Bracteae bracteolaeque oblongae, acutae, glabres-
centes, persistentes, apice glandulam longe stipitatum desinentes. Receptaculum
campanulatum, extus fere glabrum et cum pilis appressis paucis, intus basim versus
glabrum; pedicelli 7.0-I6.0 mm longi, eglandulosi. Calycis-lobi rotundati vel acuti,
extus glabri, intus brunneo-pubescentes, marginibus ciliatis. Petala 5, glabra.
Stamina 5, unilateralia, filamenta glabra, longe exserta. Ovarium ad faucem recepta-
culi insertum, pilosum. Stylus e basi ovarii ortus, versus basim hirsutus. Fructus mihi
ignoti.
TYPE. Blanchet 1508, Brazil, Bahia, without precise locality, fl (holotype, BM;
isotype, G).
DISTRIBUTION. Known only from two collections made in the coastal forests of
Bahia. Collected in flower in February.
BRAZIL. Bahia: Santa Cruz Cabralia, Belem & Pinheiro3310 fl (UB).

This species is quite distinct from, and thus not easily confused with, any other
species of Hirtella.I have adopted Briquet'smanuscript name to avoid later confusion.
The Geneva sheet of the Blanchet collection bears his writing and notes. Hirtella
insignisis most closely related to H. tocantina,but differs in the larger leaves with broad
rounded base, the much-branched inflorescence, the larger number of stamens, and
the long-stalked apical gland of the bracts.

14. Hirtella tocantina Ducke, Arch. Jard. Bot. Rio de Janeiro 3: 266. 1922.

Medium-sized tree, the young branches sparsely pubescent, soon glabrous and
lenticellate. Leaves coriaceous, oblong, Io.0-15.0 cm long, 3.0-5.5 cm broad,
acuminate at apex, with acumen 8.0-12.0 mm long, cuneate at base, the lower
surface with several palisade glands towards base, glabrous; primary veins 9-I2
pairs, prominulous on both surfaces; midrib prominulous and glabrous on both
surfaces; petioles 4.0-5.0 mm long, glabrescent, terete, eglandular. Stipules linear,
to 5.0 mm long, glabrescent, subpersistent, eglandular. Inflorescences terminal and
axillary panicles 7.0-I2.0 cm long, the rachis and branches with sparse appressed
pubescence. Bractsand bracteoles oblong-lanceolate, 2.5-3.5 mm long, membraneous,
persistent, sparsely puberulous, the margins with both sessile and short-stipitate
glands. Flowers 5.0-6.0 mm long. Receptacle campanulate, glabrescent on exterior,
glabrous within except for hairs at throat; pedicels I.0-3.0 mm long. Calyx lobes
acute, glabrescent on exterior, gray-pubescent within, the margins glandular. Petals
5, white, glabrous. Stamens 3, unilateral, with 3 short filamentous staminodes oppo-
site them, the filaments far exceeding calyx lobes. Style hirsute up to one third its
length. Ovary inserted at mouth of receptacle, pilose. Fruit not seen.
TYPE. Ducke MG I6274, Brazil, Para, fl (holotype, MG; isotypes, P, RB I5046,
US).
DISTRIBUTION. Non-flooded forest, collected from the banks of the Rios Tocan-
tins, Tapaj6z and Madeira.
BRAZIL. Para: DuckeRB 18236 fl (P, RB, S, US). BOLIVIA. Pando: Pranceet al 6468 fl (INPA,
NY).
282 Flora Neotropica

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FI 4:5- SeieofHrel.ADH.isgi(Baez58;Ahai,xo5B,fwr,x;C
flower sect~~~ion x 5;D ea,x5 -,H
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Hirtella 283

15. Hirtella piresii Prance, sp nov Fig 44 A-C.

Arbor parva vel media, ramulis juvenilibus glabrescentibus. Folia alternata

petiolata; petiolo I.O-2.0 mm longo, tereti, eglanduloso, juvenili tomentello; laminae
oblongae, coriaceae, 3.5-9.0 cm longae, 1.4-3.2 cm latae, subtus glabrae, pauci-
glandulosae, apice in acuminem 2.0-10.0 mm longum contractae, basi cuneatae;
costa media supra leviter prominente, subtus prominente, glabrescente; costis
secundariis 8-I I jugis, utrinque leviter prominentibus. Stipulae circa I.o mm longae,
lineares, eglandulosae, caducae. Flores 4.0-5.0 mm longi, in paniculis terminalibus
axillaribusque 3.0-I 1.0 cm longis dispositi, rachi ramisque glabrescentibus. Bracteae
bracteolaeque 2.0-5.5 mm longae, oblongae vel lanceolatae, puberulae vel glab-
rescentes, persistentes. Receptaculum campanulatum, extus glabrum, intus basim
versus glabrum; pedicelli 1.5-3.0 mm longi. Calycis-lobi acuti, extus glabri, intus
glabrescentes, marginibus ciliatis, eglandulosi. Petala 5, glabra. Stamina 3, unilatera-
lia, staminodiis dentiformibus opposita; filamenta glabra, longe exserta. Ovarium ad
faucem receptaculi insertum, pilosum. Stylus e basi ovarii ortus, hirsutus. Drupa
(immatura tantum visa) pyriformis; epicarpio laevi, glabro; mesocarpio tenue,
carnoso; endocarpio duro, tenue, osseo, intus hirsuto.
TYPE. Black, Cordeiro& Francisco52-I4653, Brazil, Terr. Rondonia, Porto
Velho, fl (holotype, NY; isotype, IAN).
DISTRIBUTION. Non-flooded forest in central Brazilian Amazonia, and southern
Guyana. Flowering May to September.
GUYANA. Guppy213(F. D. 7189) fl (NY). BRAZIL. Amazonas: Manaus-Itacoatiara road; Km
27, Reserva Florestal Ducke, Rodrigues& Osmarino6975 fl (INPA 15637, NY); Km 50, Rodrigues& Lima
2368 fl (INPA 9716); 3168 fl (INPA 9716, NY); Km 74, Rodrigues& Loureiro7102 fr (INPA 15764,
NY). Para: Breves; Pires, Froes & Silva 4914 fl (IAN, NY); 518o st (IAN); Rio Jar, Monte Dourado,
Oliveira3549 fr (IAN); 3853 fl (IAN); 4682 st (IAN, NY). Terr. Rond6nia: Porto Velho; Cordeiro&
Silva 231 fl fr (IAN, US).

This species is most closely related to H. tocantina,but differs in the smaller leaves
and inflorescence, the single curved apical gland of the bracts and bracteoles, and the
but differs in the single apical
smaller stipules. It is also close to H. subglanduligera,
gland of the bracts and bracteoles, the smaller coriaceous bracts, and the shorter
glabrescent inflorescence.

x6. Hirtella davisii Sandwith, Kew Bull. 1935: 125. 1935.

Tree to 30.0 m tall, the young branches sparselypilose, soon becoming glabrous
and lenticellate. Leaves oblong to oblong-elliptic, coriaceous, 6.o-I3.5 cm long,
2.3-5.0 cm broad, acuminate at apex, with acumen 5.0-Io.o mm long, subcuneate
at base, with several palisade glands towards base beneath; primary veins 9-10 pairs,
prominulous on both surfaces; midrib prominulous above, prominent and with
sparsely appressed pubescence beneath; petioles 3.0-4.0 mm long, puberulous to
glabrous, terete, eglandular. Stipules linear, 3.0-4.0 mm long, subpersistent,eglandu-
lar, glabrous. Inflorescences terminal and axillary panicles 5.0-I4.0 cm long, the
rachis and branches sparsely pilose, Bracts and bracteoles 1.0-3.5 mm long, ovate to
lanceolate, puberulous, persistent, coriaceous, with numerous stipitate glands.
Flowers 5.o-6.o mm long. Receptacle campanulate, puberulous on exterior, glabrous
within except at throat; pedicels 0.5-2.0 mm long. Calyx lobes rounded to acute,
puberulous on exterior, gray-pubescent within, the margins eglandular. Petals 5,
white, glabrous. Stamens 3, unilateral, the filaments far exceeding calyx lobes. Style
284 Flora Neotropica

hirsute up to one third its length. Ovary inserted at mouth of receptacle, pilose. Fruit
ellipsoid: epicarp glabrous, smooth; mesocarp thin, fleshy; endocarp thin, bony,
densely pilose-lanate within.
TYPE. Davis 352(F.D. 2345), Guyana, fl (holotype, K; isotype, FHO).
DISTRIBUTION. Primary forest on high ground in eastern Venezuela, Guyana,
and eastern Amazonian Brazil. Flowering March to May in the Guianas, September
to October in Brazil.
VENEZUELA. Bolivar: Bernardi7184 fl (G); LI. Williams 11626 fl (F, MICH, S, US, VEN).
Terr. Delta Amacuro: Berti 66 fl (BR, F, NY, RB, S); 94 fl (NY); 156 fl (NY, US); Blanco413 fl (NY,
VEN); 474 fr (NY, VEN); 481 fr (NY, VEN); Breteler3748 fl (NY, U, VEN); Wurdack& Monachino
39729 st (NY). GUYANA. Fanshawe1751 (F.D. 4493) fl (K, NY, P, US); I8o3(F. D. 4540) fl (K);
Maguire & Fanshawe22912 fr (A, F, K, NY, US). BRAZIL. Par,: Lima & Black 48-3118 fr (IAN);
48-3107 fr (IAN). Maranhao: Frdes1899 fl bud (A, BM, F, G, MICH, NY, S).

LOCAL NAMES.Venezuela: Ceniza negra, Guaray.

17. Hirtella subglanduligera Pilger, Notizbl. Bot. Gart. Berlin 6: 140. 1914;
Macbride, Fl. Peru 1078. 1938.

Tree to 25.0 m tall, the young branches sparsely pilose, becoming glabrous and
lenticellate with age. Leaves ovate-elliptic to elliptic, subcoriaceous, 6.0-9.0 cm long,
2.5-4.0 cm broad, acuminate at apex, with acumen 4.0-5.0 mm long, subcuneate at
base, glabrous beneath or bearing a few stiff appressed hairs, with a few palisade
glands towards base; primary veins 9-o1 pairs, prominulous on both surfaces;
midrib prominulous above, prominent and sparsely pubescent beneath; petioles
I.0-2.0 mm long, glabrous, terete, eglandular. Stipules linear, to 6.o mm long,
glabrous, subpersistent, eglandular. Inflorescences terminal and axillary panicles
8.0-22.0 mm long, the rachis and branches sparsely pilose. Bracts and bracteoles
oblong-lanceolate, membraneous, puberulous, persistent, the margins bearing
stipitate glands. Flowers ca 7.0 mm long. Receptacle campanulate, puberulous on
exterior, glabrous within except for throat; pedicels 1.5-3.0 mm long. Calyx lobes
rounded to acute, puberulous on both surfaces, the margins sparselyglandular. Petals
5, white, glabrous. Stamens 4, unilateral, with 4 short filamentousstaminodes opposite
them; filaments glabrous, far exceeding calyx lobes. Style hirsute up to one third its
length. Ovary inserted at mouth of receptacle, pilose. Fruit not seen.
TYPE. Ule 9414, Peru, Madre de Dios, fl (holotype, B, lost; lectotype, MG
I4263; isotypes, L, US).
DISTRIBUTION. Known only from the type gathering from the upper Rio Acre,
flowering in July.
This species is most closely related to H. davisii.Although both of these species
seem very closely related, they are geographically far removed from each other.
Hirtella subglanduligeradiffers from H. davisii in the 4 stamens, the narrower mem-
braneous bracts, and the shorter petioles. Their separate maintainence here is tenta-
tive, as the present species can hardly be adequately interpretedfrom extant material.

18. Hirtella ciliata Martius & Zuccarini, Abh. Akad. Miinchen I: 378. 1832;
Zucc., Flora 15(Beibl. 2): 86. 1832; Hook. f., Mart. Fl. Bras. 14(2): 36.
1867.
Hirtella rubraBentham, Jour. Bot. Hooker 2: 217. I840. Type. Schomburgk
113, Guyana, fl (holo-
type, K; isotypes, BM, CGE, GH, LE, NY, OXF, P).
Hirtella 285

Hirtella rotundataPilger, Notizbl. Bot. Gart. Berlin 6: 140. I914. Type. Ule 9043, Brazil, Cear,, fl
(holotype, B, lost; isotype, K).
GrangeriabrasiliensisHoffmansegg ex Martius & Zuccarini, Abh. Akad. Munchen I: 378. 1832,
nom nud in syn.

Tree to I2.0 m tall, with thick corky bark on trunk, the young branches tomen-
tellous, becoming glabrous with age. Leaves orbicular to ovate-elliptic, coriaceot s,
3.0-6.5 cm long, 1.5-3.6 cm broad, retuse to acuminate at apex, most frequently
mucronate, subcordate to rounded at base, hirsute beneath or with sparse appressed
pubescence; palisade glands present; primary veins 6-9 pairs, plane above, prominu-
lous beneath; midrib prominent on both surfaces; petioles 0.5-2.0 mm long, tomen-
tellous, terete, eglandular. Stipules ca I.o mm long, oblong, tomentellous, eglandular,
persistent. Inflorescences terminal and subterminal panicles 5.0-22.0 cm long, the
rachis and branches sparsely tomentellous. Bracts and bracteoles 1.0-3.0 mm long,
ovate to oblong, pubescent, persistent, with numerous stipitate glands. Flowers
3.5-5.0 mm long. Receptacle campanulate, puberulous on exterior, pubescent to base
within; pedicels 2.0-6.0 mm long. Calyx lobes acute, puberulous on exterior, gray-
pubescent within, the margins glandular. Petals 5, glabrous, greenish-white. Stamens
6-9, unilateral with a few short filamentous staminodes opposite them; filaments
slightly exceeding calyx lobes. Style hirsute at base only. Ovary inserted at mouth of
receptacle, pilose. Fruit round to ellipsoid; epicarp smooth and glabrouswhen mature;
mesocarp thin, fleshy; endocarp thin, bony, hirsute within.
TYPE.Martius 1722, Brazil, Bahia, fl (holotype, M; isotype, BR).
DISTRIBUTION.
Savannas of the Guianas and Amazonia, cerradosof the Planalto
of Central Brazil, and taboleiros
of northeasternBrazil.
GUYANA. Fanshawe2586(F. D. 5374) fl (F, GH, K, NY, S, US); 2952(F.D. 6282) fl (K, NY, S);
Im Thurnsn fl (K); Jenman5678 fl (K). FRENCH GUIANA. Black & Klein 54-17361 fl (IAN, NY).
BRAZIL. Para: Black 47-1833 fl (IAN); 55-18691 fl (IAN); 55-i8736 fl fr (IAN, NY); 57-20074 fl
(IAN, NY, UB); Ducke 2155 fl (R); MG 8088 fl (BM, MG, RB); MG 9917 fl(BM, MG, RB); MG
i5584 fl fr (MG); RB 15066 fl (RB); RB r5068 fl (RB); Fr6es20456 fl (IAN, NY); Riedel 1529 fl (US);
B. Rodriguessn fl (R7i6I); A. Silva 489 fl (IAN, NY, UB); N.T. Silva 647 fl (IAN); Siebersn fl (BR);
Sprucesn fl (C, F, GH, GOET, LD, LE, NY, S); 382 fl (K, P). Terr. Amapa: Bastos 40 fl (RB I36913);
Cowan38648 fl (NY); Pires, Rodrigues& Irvine51089 fl (IAN, MG, NY); Pires & Cavalcante52534 fr
(IAN, MG, NY). Maranhao: DuckeMG 764 fl (INPA, MG); Eiten 4605 fl (US); 5379 fr (US); 5467 fr
(US); LisbbaMG 2505 fl (BM, P, RB, US); Macedo4012 fl (K, IAN, RB, S, SP, US); Pires & Black
1633a fl (IAN,) 1721a fl (IAN). Goias: Gardner4125 fl (BR, K); Maguire,Pires, Maguire & Silva 56077
fl (NY); Pires & Black 2374 fl (NY); Prance & Silva 58456 fl (F, K, RB, NY). Piaui: Gardner2562
fl (BM, K, US); Luetzelburg194 fl (NY). Ceara: Allemdo & Cysneiros564 fl (R); Duarte 1435 fl (RB);
Ducke 2381 fr (IAN, MG, NY); Gardner1591a fl (BM, BR, F, GH, K, NY, P, S, US); Guedes632 fl
(MG); 638 fl (IAN, MG, NY); Lima 58-3272 fl (IPA); Lima & Magalhaes52-1x135 fl (R); Luetzelburg
25878 fl (M, NY). Rio Grande do Norte: Castellanos22986 fl (R); WurdackB 164 fl fr (NY). Paraiba:
Lima 50-743 fl fr (IPA); Pickel 3420 fl (IPA). Pernambuco: Lima 58-2882 fr (IPA); Tavares 1ooo fl
(HST, US). Bahia: Blanchet46 fl (P); 328 fl (P); i841 fl (BM, GH, LE, NY, P, R); Lima 50-433 fl
(IPA); Torrend311 fl (SP); Vidalsn fl (R 35180); sn fl (R 36623).

LOCAL NAMES.Guyana: Bokobokotokon.Brazil: Canoe, Chordo, Murtinha, Pau

Pombo.
In light of material now available, H. rubraand H. rotundatacannot be main-
tained. Hooker ( 867), recognized two varieties of H. ciliatabut he did not name them.
These varieties, based on small differences in leaf shape and pubescence, cannot be
maintained in light of the additional material studied.

I9. Hirtella hoehnei Pilger, Revista Mus. Paul. 13: 1251. 1922.

Shrub to 5.0 m tall, the young branches tomentellous. Leaves ovate to ovate-
286 Flora Neotropica

elliptic, coriaceous, 4.5-I5.0 cm long, 2.5-7.0 cm broad, acute to acuminate at apex,

with acumen 0-4.0 mm long, subcordate at base, hirsute along venation beneath,
with palisade glands towards base; primary veins 7- 1 pairs, plane to prominulous
above, prominent beneath; midrib prominulous above, prominent and hirsute be-
neath; petioles 2.0-4.0 mm long, villous-tomentellous, terete, eglandular. Stipules
lanceolate, tomentose, to 6.0 mm long, persistent, eglandular. Inflorescencesdensely
clustered terminal and subterminal panicles 8.0-20.0 cm long, the rachis and
branches hirsutulous. Bracts and bracteoles 2.0-5.0 mm long, acute, oblong, mem-
braneous, hirsutulous, persistent, the margins with numerous stipitate glands.
Flowers 7.0-8.0 mm long. Receptacle campanulate, hirsutulouson exterior, glabrous
within except for deflexed hairs at throat; pedicels 2.0-4.0 mm long, Calyx lobes
acute, hirsutulous on exterior, gray-pubescent within, the margins with stipitate
glands. Petals 5, white, glabrous. Stamens 7, unilateral, with short filamentous
staminodes opposite them, the filaments far exceeding calyx lobes. Style hirsute up to
one third its length. Ovary inserted at mouth of receptacle, pilose. Fruit ellipsoid;
epicarp smooth, glabrous; mesocarp thin, fleshy; endocarp thin, bony, densely
hirsute within.
TYPES.Hoehne2141, Brazil, Mato Grosso, fl (holotype,B, lost; photos, F, NY); 2154 fl
(paratype, SP 5596); 2155 fl (paratype, R).
DISTRIBUTION.Beside rivers and in wet open marshy places (brejos)on the Planal-
to of central Brazil. Flowering May to October.
BRAZIL. Mato Grosso: Irwin & Soderstrom 6459 fl fr (NY); Maguire,Pires, Maguire& Silva 56442
fl (NY); Malme 2028 fl (S); Pranceet al 59195 fl (NY, UB); Riedel598 fl (US). Goias: Macedo1827 fl
(RB, S, SP).

This species is closely related to H. glandulosa,but may be distinguished by the

densely crowded inflorescence, the leaf shape, and the prominent venation of the
upper surface of the leaf. I have not yet chosen a lectotype of this species because of
the poor condition of the two paratypes which I have examined.

20. Hirtella glandulosa Sprengel, Neue Entdeck. I: 303. I820; DC., Prodr.
2: 528. 1825; Zucc., Flora 15(Beibl. 2): 86. 1832; Hook. f., Mart. Fl. Bras.
14(2): 35. 1867.
Hirtella damazianaBeauverd, Bull. Herb. Boiss. II. 7: 706. I907. Type. Damazio 1594, Brazil,
Minas Gerais, fl (holotype, G).
HirtellahookeriPilger, Notizbl. Bot. Gart. Berlin 6: 139. 1914. Type. Gardner1947, Brazil, Piaui, fl
(holotype, B, lost; isotypes, BM, F, K, NY, SP, US).
Hirtella velutinaPilger, Notizbl. Bot. Gart. Berlin 6: I4I. 1914. Type. Schomburgk 1051, Guyana,
fl (holotype, B, lost; isotype, K).
Hirtella wachenheimii R. Benoist, Bull, Mus. Hist. Nat. Paris 29. 595. 1923. Type. Wachenheim 372
fl (holotype, P; isotype, K).

Tree to 25.0 m tall, usually much smaller, the young branches tomentose, be-
coming glabrous and lenticellate with age. Leaves oblong to ovate, thick-coriaceous,
4.5-23.0 cm long, 2.5- 1.5 cm broad, acuminate at apex, with acumen 2.0-5.0 mm
long, rounded to cuneate at base, rarely weakly subcordate, hirsute beneath, the
palisade glands rare, with slightly impressed venation above; primary veins 8-15
pairs, impressed above, prominent beneath; midrib prominulous above, prominent
and hirsute beneath, petioles I.5-5.0(-I2.0) mm long, tomentellous, terete, eglandu-
lar. Stipules linear, 4.0-8.0 mm long, tomentellous, eglandular, persistent. Inflores-
cences terminal and subterminal panicles 9.0-26.0 cm long, the rachis and branches
hirsute-tomentellous. Bracts and bracteoles ovate, coriaceous, persistent, pubescent,
Hirtella 287

with numerous stipitate glands. Flowers 6.0-8.0 mm long. Receptacle campanulate,

puberulous on exterior, glabrous within except for deflexed hairs at throat; pedicels
1.0-3.0 mm long. Calyx lobes acute, puberulous on exterior, gray-pubescent within,
the margins with stipitate glands. Petals 5, white, glabrous. Stamens 3-5, unilateral,
with a few filamentous staminodes opposite them, the filaments far exceeding calyx
lobes. Style hirsute on lower portion only. Ovary inserted at mouth of receptacle,
pilose. Fruit round to pyriform; epicarp smooth, glabrous when mature; mesocarp
thin, fleshy; endocarp thin, bony, densely hirsute within.
TYPE. Sellow 44, Brazil, probably Espirito Santo, fl (holotype, B, lost; lectotype,
G; isotype, fragment F), Sellow sn (probable isotype, LE).
DISTRIBUTION. Damp forest in open places, mainly gallery forest, river banks,
and savannna margins. Commonest in the Planalto of Central Brazil, but extending
into the savannas of Amazonia and the Guianas.
Representative collections: VENEZUELA. Bolivar: Steyermark86482 fr (VEN); LI. Williams
12015 fl (F, K, S, US, VEN). Terr. Delta Amacuro: Little 17679 fl (VEN). GUYANA. Fanshawe313
(F.D. 3049) fl (NY); 3356(F.D. 6885) fl (MICH, NY, S). FRENCH GUIANA. For. Serv.7726 fl (NY).
BRAZIL. Amazonas: CardonaI373 fl (F, NY, US, VEN); Ducke RB 3558I fl (RB, US); Pranceet al
3406 fl (INPA, MG, NY); Silva & Brazao 60693 fr (NY). Para: Black 48-3461 fl (IAN); Burchell9780
fl (GH, K, NY, P); CoelhoINPA 15883 fl (INPA); DuckeMG 16291 fl (MG); RB 18239 fl (RB); Killip &
Smith30452 fl (NY, RB, US); Pires 51920 fl (K, NY, US); Pires & Black 478 fl (GH, NY, P); Pires &
Silva 10727 fl (IAN, NY); Pranceet al 2069 fl (IAN, NY); Sampaio5549 fl (R); 555oA fl (R). Terr.
Amapa: Irwin & Westra47183 fr (F, GH, IAN, MG, NY, US); Irwin, Pires & Westra48463 fr (F, IAN,
MG, NY, US); Pires, Rodrigues& Irvine50418 fl (F, GH, IAN, MG, NY, S, US); 50499 fl (F, GH, IAN,
MG, NY, S, US); 51232 fl (F, GH, IAN, INPA, MG, NY, S, US). Maranhao: DuckeMG 381 fl (BM,
MG); 764 fl (BM, MG); Frdes 11877 fl (A, F, MICH, NY, US); Lisboa MG 2501 fl (BM, MG, RB);
Pires et al i600 fl (IAN); I7i9a fl (IAN). Piaui: Luetzelburg1381 fl (M, NY); 1741 fl (RB 5900). Ceara:
Dahlgren 951 fl (F); Duarte I447 fl (NY, RB, 6431o); Lima 58-3279 fl (IPA); sn fl (IPA). Paraiba:
Luetzelburg26780 fl (A, F, M, NY, US). Pernambuco: Gardner2837 fl (BM, GH, K, RB). Bahia:
Belem & Magalhaes 771 fr (IAN, NY, UB); Belem & Pinheiro3201 fl (NY, UB); Duarte 6645 fl (RB);
Ule 7306 fl (K); Zehntner632 fl (R). Goias: Belem 199i fl (NY, UB); 1994 fl (NY, UB); 2010 fl (IAN,
NY, UB); Burchell5969 fl (GH, K, NY, P, US); 7620 fl (GH, K, P); Glaziou o0706fl (C, K, P); 21110
fl (BR, K, P, S); Irwinet al 18009 fl (NY); LaneSP 69995 fl (SP); Macedo3607 fl (IAN, NY, RB, S, US);
3885 fl (K); Maguireet al 561oo fl (NY); Oliveirasn fl (R I16325); Pohl 1275 fl (F, NY, W); Prance &
Silva 58564 fl (NY); St. Hilaire 850 fl (P); Ule 179 fl (P, R). Dist. Federal: Cobra296fl (IAN, NY, UB);
CoelhoINPA 16712 fl (INPA, NY); Heringer9035 fl (NY, UB); Irwin et al 5368 fl (NY); 7824 fl (IAN,
NY); 7922 fl (IAN, NY); Pires & Mattos9862 fl (NY, UB); Prance& Silva 59055 fl (NY, RB, UB, US);
Ramos 6653 fl (K, NY); Sucre286 st (NY, RB, UB). Minas Gerais: Barreto6784 fl (R); 6785 fl (F);
9998 fl (F, R); 12127 fl (R); Claussensn fl (BM, BR, K); 255 fl (P); Heringer728 fl (IAN, NY, RB, UB);
Irwin et al 23454 fl (NY); Magalhaes13931 fl (NY); Maguire,Magalhaes & Maguire49286 fl (NY, US);
Markgraf3238 fr (F, RB); Nettosn fl (GH); Pohl snfl (BR); RegnellIII-r51 fl (BR, S); Riedelsn fl (US);
Santos & Castellanos24388 fl (NY); St. Hilaire 1973 fl (P); Warmingsn fl (C, NY, P. S). Mato Grosso:
CoelhoINPA 15883 fl (INPA, NY); Hunt 5597 fl (K, NY); 5923 fl (K, NY); Irwin et al 16137 fl (IAN,
NY); 16435 fl (IAN, NY); 16745 fl (IAN, NY); Maguireet al 56342 fl (NY); 56357 fl (GH, MICH, NY);
Malme 2028a fl (S); Prance & Silva 59116 fr (K, NY, RB, US); 59429 fl (K, NY, US); H. Smithsn fl
(R 7157).

LOCAL NAMES. Guyana: Bukotokon. Brazil: Vermelhtio.

This species extends over a wide geographic range and, as might be expected,
is rather variable in leaf shape. There is a tendency to larger leaves in the northern
and northeastern part of the range. However, smaller leaved specimens have been
taken there too, and there is a complete gradation in leaf size. Not only does this
smooth variation fail to offer any basis for recognition of infraspecific taxa, but I have
also observed in the field that leaf size is quite variable on the same tree. However, I
find no grounds for maintaining H. hookeri,H. velutina, and H. wachenheimeias separate
species, which represent the same species described from different parts of its range
without adequate comparison.
Hooker's (i867) two unnamed varieties, based on inflorescence length and
receptacle shape similarly cannot be recognized.
288 Flora Neotropica

Hirtellaglandulosahas sometimes been confused with H. bullata,but the latter

differs in the thicker more bullate leaves usually with cordate bases, the inflorescence
with a long tomentose rachis bearing only small lateral branches, and the larger
glands of the inflorescencewhich arise on the pedicels or are sessile on the bracts etc.

2I. Hirtella bullata Bentham, Jour. Bot. Hooker 2: 216. 1840; Hook. f., Mart.
Fl. Bras. 14(2): 35, t. II (i). I867.
Hirtella aureohirsutaPilger, Bot. Jahrb. 37: 538. 1906; Macbride, Fl. Peru. 1075. 1938, synon nov.
Type. Weberbauer 4487, Peru, Loreto, fl (holotype, B, lost; isotype, G; isotype, fragment F).
Hirtella amplexicaulisPilger, Notizbl. Bot. Gart. Berlin 6: I38. 1914, synon nov. Type. Ule 798r,
Brazil, Terr. Roraima, fl (holotype, B, lost; isotypes, L, K, MG).
Hirtella stipitadeniaRusby, Mem. N.Y. Bot. Gard. 7: 249. 1927, synon nov. Type. Rusby768,
Bolivia, fl (holotype, NY; isotype, US).

Shrub or small to medium-sized tree, with long horizontal branches often

sprawling over vegetation, the young branchlets tomentellous, becoming glabrous
and lenticellate with age. Leaves oblong to ovate, thick-coriaceous, 6.0-22.0 cm long,
3.5-10.0 cm broad, acute to short-acuminate at apex, with acumen 0-I2.0 mm long,
subcordate or rarely rounded at base, bullate above with impressed venation, hirsute
beneath; primary veins 11-17 pairs, impressed above, prominent beneath; midrib
prominulous above, prominent and hirsute-tomentose beneath; petioles 1.5-4.0 mm
long, tomentose, terete, eglandular. Stipules 3.0-6.0 mm long, linear-lanceolate,
tomentellous, eglandular, persistent. Inflorescences terminal and axillary panicles
9.0-35.0 cm long, the long central rachis bearing short lateral branches with I-3
flowers, or rarely reduced to racemes, 8.0-33.0 cm long, the rachis and branches
tomentose. Bracts and bracteoles I.0-4.0 mm long, oblong-lanceolate, bearing large
sessile glands or with numerous stipitate glands arising from the region of pedicel
articulation. Flowers 4.5-8.o mm long. Receptacle campanulate, tomentellous on
exterior, glabrous within except at throat; pedicels I.0-3.0 mm long. Calyx lobes
acute, tomentellous on exterior, sparsely gray-puberulous within, the margins
eglandular. Petals 5, white, glabrous. Stamens 4-6, unilateral with small toothed
staminodes opposite them, the filaments glabrous, purple, far exceeding calyx lobes.
Style pilose for up to half its length. Ovary inserted at mouth of receptacle, pilose.
Fruit ellipsoid; epicarp smooth and glabrous; mesocarp thin, fleshy; endocarp very
thin and bony, hirsute within.
TYPE. Schomburgk 132, Guyana, fl (holotype, K; isotype, BM).
DISTRIBUTION. Open habitats at river and savanna margins and savannas in
Guyana and western Amazonia to Bolivia.
COLOMBIA. Vaupds: Allen 3082 fl (US); Humbert27407 fl (F, P); Schultes,Baker & Cabrera
18140 fl (GH, NY, US). VENEZUELA. Terr. Amazonas: Cowan& Wurdack31454 fl (NY); Farinaset al
475 fl (NY, VEN); Maguire & Maguire35045 fr (NY); Phelps & Hitchcocksn fl (NY); Spruce3084 fl
(BM, BR, K, NY, P); Steyermark 57904 fl (F, NY); Tate 146 fl (NY); 372 fl (NY); LI. WilliamsI46I6 fl
(F, US, VEN); Wurdack& Adderley43231 fl (NY). Bolivar: Agostini250 fl (VEN); Bernardi2611 fl (NY);
6524 fl (NY); Cardona2449 fl (VEN); 2556 fl (VEN); 2827 fl (VEN); Lasser 1796 fl (VEN);
Maguire & Maguire 40480 fr (NY); Maguire & Wurdack33774 fl (NY); Steyermark74707 fl (F, NY);
90821 fl (NY); 93176 fr (VEN); Steyermark& Wurdack47 fl (NY). GUYANA. Appun 1004 fl (K);
R. Boyan53 (F.D. 7877) fl (K, NY); Grahamr24 fl (K); 439 fl (K); Maguire,Bagshaw & Maguire40660
fl (NY); 40676 fl (NY); Maguire & Fanshawe32181 fr (NY); 32196 fl (NY); 32216 fr (NY). PERU.
Junin: Killip & Smith24794 fl (BM, F, NY, US); 25276 fl (F, GH, NY, US); Macbride5532 fr (F, GH,
US); Schunke328 fl (F); 329 fl (F). Loreto: Woytkowski 5787 fl (GH, US). Amazonas: Wurdack1986 fl
(F, K, NY, US). BRAZIL. Amazonas: Black 48-2588 fr (IAN, NY); Cavalcante701 fl (MG, NY);
Frdes 22257 fl (IAN, NY); 28104 fl (IAN); Krukqff4916 fl (A, BM, K, LE, NY, S, US); Pires 491 fl
Hirtella 289

(IAN, NY). BOLIVIA. Fleischmann353 fl (S); 354 fl (S); Luetzelburg21531 fl (R); Rusby703 fl (GH,
LE, MICH, NY, US).

LOCALNAMESAND USES.Guyana: Bokobokotokon.Bolivia: Urarau. Edible fruit.

The three species reduced to synonymy under H. bullataare not at all distinct.
Hirtellaaureohirsutawas said to differ from H. bullatasolely in the indumentum
of the inflorescence. This more golden pubescence occurs in some collections, over
much of the range of H. bullataand thus offers no basis for the recognition of a
separate taxon. Hirtella amplexicaulisand H. stipitadeniawere not compared to H.
bullatain the original description and fall well within the range of the latter species.
Hirtella bullatais most closely related to H. glandulosa,and the differences are given
under that species.

22. Hirtella americana Linnaeus, Sp. P1. ed. I. 34. I753; [Hort. Cliff. 17.
1738]; Sandwith, Kew Bull. 1931: 376. I931; Standl., Publ. Field Mus.
Bot. I8: 480. I937; non. H. americanaAublet, nec Jacquin, nec Hooker f.,
nec Fritsch, nec Ducke etc.
Hirtella mollicomaKunth, Nov. Gen. et Sp. 7: 263, t. 658. 1825; DC., Prodr. 2: 528. 1825; Zucc.,
Flora I5 (Beibl. 2): 85. i832;Jennings, Ann. Carneg. Mus. II: I22. 1917. Type. Humboldt
& Bonpland1798, Colombia, Magdalena, fl fr (P; fragment F).
Hirtella mollissimaHemsley, Biol. Centr. Am. Bot. I: 366. 1880, sphalm.
Chrysobalanus americanus(Linnaeus) Morales, Ann. Acad. Ci. Med. Fisic. & Nat. Habana 47: 391.
1887.

Tree to 20.0 m tall, usually smaller, or shrub, the young branches tomentellous,
becoming glabrous with age. Leaves elliptic to oblong, coriaceous, 5.5-I9.0(-23.0)
cm long, 2.0-7.0 cm broad, acuminate at apex, with acumen 2.0-8.0 mm long,
rounded to subcuneate at base, the lower surface hirsute; primary veins 9-I3 pairs,
plane or slightly impressed above, prominent beneath; midrib prominulous above,
prominent and tomentellous beneath; petioles 2.0-3.0 mm long, tomentellous, terete,
eglandular. Stipules lanceolate, 2.o-8.o mm long, tomentellous, eglandular, persis-
tent. Inflorescences terminal and axillary panicles 5.0-20.0 mm long, the rachis and
branches tomentellous. Bracts and bracteoles I.o-5.o mm long, ovate, persistent,
tomentellous, bearing numerous large sessile or slightly stipitate glands. Flowers
4.0-6.o mm long, Receptacle campanulate, tomentellous on exterior, glabrous
within except for throat; pedicels 0.5-2.0 mm long. Calyx lobes rounded, tomentel-
lous on exterior, gray-pubescent within, the margins eglandular. Petals 5, glabrous,
white. Stamens 3, unilateral with small teeth opposite them; the filaments white to
purple, far exceeding calyx lobes. Style hirsute up to half its length. Ovary inserted
at mouth of receptacle, pilose. Fruit ellipsoid; epicarp sparselypubescent to glabrous;
mesocarp thin, fleshy; endocarp thin, hard, bony, hirsute within.
TYPE. Linnaeus,HortusCliffortianus, p I7, fl (holotype, BM).
DISTRIBUTION. Dry forest on elevated ground in Cuba, through Central America
from Tabasco, Mexico, to Panama, and in northern Colombia and Venezuela.
Representative collections : MEXICO. Tabasco: Matuda3270 fr (A, F, K, MEXU, MICH, MO,
NY); 3343 fl (A, F, MEXU, MICH, NY). Campeche: Pennington& Sarukhdn9632 fr (NY). GUATE-
MALA. Aguilar I45 fl (F); 329 fl (A, K, MICH, NY); Bartlett 12741 fl (A, F, GH, K, LE, MEXU,
MICH, NY, S, US); Egler 42-224 fl (F); 42-296 fl (F); Lundell2099 fl fr (MEXU, MICH, S); 2435 fl
(BM, F, MEXU, MICH); 2859 fr (F, MICH); 2956 fl (F, MICH, S); 4878 fl (F); Steyermark 39449 fl
(F); 39514 fl fr (F); 45743 fl (A, F, NY, S). BRITISH HONDURAS. Brownsn fl (F); Campbellsn fl
(K); Castillo 24 fl (F); Chanek213 fl (F, K, MEXU, MICH); Cons.of Forests95 fl (F); Gentle r150 fl
(A, F, K, MICH, MO, NY, P, S, US); 2456fl (A, F, K, MEXU, MICH, MO, NY); 4574 fr (MICH);
Hunt 473 fl (BM, F, K, US); Schipp82 fl (A, BM, F, GH, K, LE, MICH, MO, NY, S, US); 660 fl
290 Flora Neotropica

(BM, F, GH, K, MO, NY); WinzerlingII-io fl (US). HONDURAS. Molina R. 6905 fl fr (F, US);
11807 fl (F, NY, US); Standley18222 fl (F, GH, US); 54286 st (A, F, US); 56606 fl (A, F, US); 56685
fl fr (A, F, US); Wilson 481 fl (NY, US). NICARAGUA. Molina R. 2133 fl (US); 2383 fl (F, GH);
Proctor,Jones & Facey 27045 st (NY); 27332 fl fr (NY); Standley19358 fl (F, US). COSTA RICA.
Pittier 6510 fl (BM, G, P, US); Skutch3948 fl (A, K, NY, S, US); 4130 fl (A, K, MO, NY, S, US);
48o1 fl (F, US); Tonduz3824 fl (US); Valerio891 fl (F). PANAMA. Duke 8109 fl (MO); 8132 fl (MO);
Erlansono18 fr (GH, MICH, NY, US); 121 fr (GH); Hayes 646 fl (BM, BR, G, K, P); Johnston513 fl
(GH, MO, S, US); 546 fl (BM, Fl, GH, P); Pittier 2708 fl (BM, NY, US); Seeman544 fl (BM, GH, K);
Sternet al 1798 fl (MO); Tyson3713 fr (MO); R. S. Williams 247 fl (NY); Woodworth& Vestal728 fl
(A, F). CUBA & IS. OF PINES. Baber2980 fl (A, F, GH, NY); Blain 25 fr (F); Brittonet al 14238 fl
(F, NY, US); Curtiss260 fl (A, BM, F, GH, K, LE, NY, P, US); Earle 655 fl (F, NY, US); Ekmnan
10796 fl (NY, S); 12146 fl (S); 16362 fl (A, F, LD, S); Hitchcocksn fl (F, US); Jennings 208 fr (BM,
GH, NY, US); Killip 41313 fl (US); 42678 fl (US); Le6n & Victorin18874 fl (GH); Sagra sn fl (P);
Shafer 1 178 fl (F, NY, US); VanHerman56I fl (BM, F, NY, US); Victorin& Alain 51 fl (GH); Wright
2409 fl (BM, GOET, K, LE, NY, P, S, US). COLOMBIA. Bolivar: RomeroC. 1729 fl (COL). Magda-
lena: Allen 599 fl (F); Espina & GiacomettoAi5 st (F, US); Gonzdlez, Gutidrrez& Barkley 18C-o42 fl
(BM, F, NY); RomeroC. 255 fl (COL, F, US); 817 fl (COL, IAN, US); Schlim898 fl (BR, G, K, P,
US); H. H. Smith850 fl (BM, BR, COL, F, FI, GH, K, MICH, NY, P, S, US); 85oa fr (NY). Boyaca:
Espinal et al i668 fl (COL). Cundinamarca: Garcia-Barriga11710 fr (COL, US). Tolima: Mason 13778
fl (COL, GH, US); Pennell 3637 fl (GH, NY, US). Huila: Little 7919 st (COL, NY, US); 8066 fl
(COL, NY, US); 808i fr (COL, NY, US); Maguire & Maguire 44I93 fl (NY); Rusby& PennellI 30 fl
(GH, NY, US). Meta: Goudotsn fl (GOET); Idrobo & Schultes1292 fl (COL, US); Little & Little 8304
fl (COL, NY, P, US); S. G. Smith & Idrobo1447 fl (COL, F, MICH, US); Sprague157 fl (K, US);
Triana 4223 fl (BR, FI, K, NY, US). Without precise locality: Mutis 3551 fl (US); 3552 fl (US);
3553 fl (US); 3555 fl (US); 5047 fl (US); Linden1638 fl (G, K, NY, OXF, P). VENEZUELA. Zulia:
Lasser2611 fl (VEN); Plee 14 fl (P); Tejeraizo fl (GH, NY, US). Barinas: Bernardi1952 fl (NY, VEN);
Breteler3730 fl (NY, SP, VEN). Apure: Bernardi o80 fl (K, VEN).

LOCALNAMES. Mexico: Icatillo.Honduras: Pasa. British Honduras: PigeonPlum,

Wild CocoPlum. Nicaragua: Guaviluna.Cuba: Icacode Aura, Icacopeludo.Colombia:
Caritade negro,Garrapato. Venezuela: raya.
The identification of Linnaeus' species H. americanahas been much confused in
the past. Sandwith (Kew Bull.193I: 376. 1931) in an important contribution to our
knowledge of this species, cleared up the problem, by showing that H. mollicomais
synonymous with H. americana L. Much material previously identified as H. americana
belongs to H. racemosaLam. The confusion arose when Aublet (I775) misinterpreted
H. americana. Aublet's definition was followed by all later authors prior to Sandwith.
I have studied both the Linnaean type and the Aublet specimen, and concur with
Sandwith.
Hirtellaamericanahas also been much confused with H. glandulosa,but the latter
can easily be distinguished by the thicker leaves with deeply impressed venation on
the upper surface, the more copiously branched inflorescence, the smaller and much
more numerous long-stalked bracteal glands, the glandular margins of the calyx
lobes, the more numerous stamens, and the round (not flattened) fruit.
Differences between H. americanaand H. guatemalensis are discussed under the
latter species.

23. Hirtella guatemalensis Standley, Trop. Woods I :: 19. I927.

Small tree, the young branches hispid, becoming glabrous and lenticellate with
age. Leaves oblong, subcoriaceous, 1.0-22.0 cm long, 4.0-7.5 cm broad, acuminate
at apex, with acumen 8.o-20.0 mm long, rounded to weakly subcordate at base,
hirsute on primary venation beneath; primary veins I I-16 pairs, almost plane above,
prominent beneath; midrib prominulous above, prominent and hirsute beneath;
petioles 1.5-4.0 mm long, hispid when young, becoming glabrous with age, terete,
Hirtella 291

eglandular. Stipules ca 5.0 mm long, linear, pilose, persistent, eglandular. Inflores-

cences terminal and axillary little-branched panicles, with a long central rachis
9.0-I8.o cm long, bearing short few-flowered lateral branches, the rachis and
branches hispid. Bracts and bracteoles 3.0-5.0 mm long, oblong to oblong-lanceolate,
hispid, persistent, the margins of bracts bearing small stipitate glands. Flowers
6.o-8.o mm long. Receptacle campanulate, hispid on exterior, glabrous within except
at throat; pedicels 4.0-0I.0 mm long. Calyx lobes rounded, hispid on exterior,
puberulous within, the margins eglandular. Petals 5, glabrous, white. Stamens 3-4,
unilateral with short filamentous staminodes opposite them, the filaments glabrous,
far exceeding calyx lobes. Style hirsute up to one third its length. Ovary inserted at
mouth of receptacle, pilose. Fruit round to ellipsoid; epicarp tomentose; mesocarp
very thin, fibrous; endocarp 1.5-2.0 mm thick, hard, fibrous, hirsute within.
TYPES.Von Tuerckheim II-r114, Guatemala, fl (US); Von Tuerckheim 8680 (ed.
J. D. Smith), Guatemala, fl (US).
Forest on well drained soil in Guatemala and British Honduras.
DISTRIBUTION.
Flowering December to February.
GUATEMALA. Jones & Facey 3518 fr (NY); Steyermark41615 fl (F, US). BRITISH HON-
DURAS. For. Dept. I9 fr (F); Gentle4493 fr (MICH); 5229 fl (F, MICH); Peck937 fl (K); StevensonI62
fr (F); SchippIo98fl (A, BM, F, FI, GH, K, MICH, MO, NY, S).

LOCALNAMES.Guatemala: Icaco de montana. British Honduras: Achotilla, High

RidgeCocoaPlum,Icacomacho.
The material cited above was distributed under 4 different specific names,
indicative of the fact that this species has been much confused with others. Further-
more, some material of H. americanahas been distributed as H. guatemalensis. Hirtella
is
guatemalensis closely related to H. americana,but differs in the hispid inflorescence,
young branches, and petioles, the more chartaceous leaves with a longer acumen, the
more sparsely pubescent leaf undersurface, the larger flowers, and most significantly
in the large fruit with a densely tomentose epicarp and a much thicker fibrous
endocarp. The endocarp is much thicker and more robust than in most other species
of Hirtella.

24. Hirtella eriandra Bentham, Jour. Bot. Hooker 2: 2I7. I840; Hook. f.,
Mart. Fl. Bras. I4(2): 38. 1867.

Small tree or shrub, the young branches tomentellous, becoming glabrous with
age. Leaves oblong, chartaceous to subcoriaceous, 8.0-I9.0 cm long, 3.3-8.0 cm
broad, acuminate at apex, with acumen 3.0-15.0 mm long, rounded at base, hirsute
beneath at least on principal venation, usually on other veins as well, with palisade
glands frequent towards base; primary veins I0-13 pairs, plane or nearly so above,
prominent beneath; midrib prominent above, prominent and hirsute beneath;
petioles 2.0-4.0 mm long, tomentose, terete, eglandular. Stipules linear, to 5.0 mm
long, tomentellous, persistent, eglandular. Inflorescences terminal and subterminal
panicles, with a thick central rachis I2.0-50.0 cm long bearing short branches or
clusters of flowers, the rachis and branches tomentellous. Bracts and bracteoles 2.0-
6.0 mm long, oblong to lanceolate, tomentose, with a few large sessile glands or a
large terminal gland on some bracts only. Flowers 6.0-7.0 mm long. Receptacle
campanulate, tomentellous on exterior, glabrous within except at throat; pedicels
o.5-6.o mm long. Calyx lobes acute, sparsely tomentellous on exterior, sparsely
puberulous to glabrescent within, the margins eglandular. Petals 5, greenish-white,
292 Flora Neotropica

glabrous. Stamens 6, unilateral, the filaments pink, far exceeding calyx lobes, hir-
sutulous on lower portion. Style hirsute up to half its length. Ovary inserted at mouth
of receptacle, tomentose. Fruit rotund; epicarp tomentose, longitudinally striate
when mature; mesocarp thin, endocarp thin, fibrous, hard, hirsutulous within.
TYPE. Schomburgk 886, Brazil, fl (holotype, K; isotypes, BM, CGE, GH, NY, P).
DISTRIBUTION.
Riverine forest, savanna margins or cerradoand secondary forest
in Amazonian and central Brazil. Flowering August to December.
BRAZIL. Amazonas: Black 48-2435 fl (IAN, NY, R); Ducke MG 7181 fl (MG); Frdes26336 fl
(IAN, NY); 26463 st (IAN, NY); 2936 fl (IAN); Krukoff5979 fl (A, BR, F, K, LE, MICH, NY, S,
US); 7124 fl (A, K, NY, S, US); KuhlmannRB 17982 fl (RB, S, US); Pranceet al 3212 fr (INPA, MG,
NY); Rodrigueset al 2486 fl (INPA 9835); 3286 fl (INPA 9835, NY); 3920 fl (INPA 10485); 4121 fr.
(INPA io686, NY); 4795 fl (INPA 13322); 5781 fr (INPA 14493, NY); 7097 fl bud (INPA I5759,
NY). Para: Cavalcante204I fl (MG 35835); DuckeMG 4881 fr (MG); MG 7980 fl (MG, RB); RB 18235
fl (RB); Froes33939 fr (IAN); Kuhlmann1842 fr (RB 17968); Oliveira1202 fl (IAN); 3508 fl (IAN);
Pires & Silva 4579 fr (IAN); Pranceet al I474 fl (IAN, NY); I496 fl (IAN, NY); M. Silva I 113 fl (MG);
1455 fl (MG); 1708 fl (MG); 1721 fl (MG); N. T. Silva iioi fl (IAN, NY); 1138 fl (IAN, NY); 1442 fl
(IAN, NY); Spruce17 fl (K). Terr. Amapa: Bastos5 fl (RB 56891); I99 fl (F, IAN, INPA, RB 96384);
RB 96382 fl (RB); Cowan38527 fl (NY); 38624 fl (NY); DuckeMG 1966 fr (MG); Egler 626 fl (MG);
Frdes 27377 fl (IAN, LE); Frdes & Black 27265 fr (IAN); Oliveira3646 fl (IAN); Pereira3356 fl (RB
IOI669); Pires & Silva 4817 fr (US). Maranhao: Tavares1107 fl (HST, US); 1130 fl (HST, US). Terr.
Rond6nia: Duarte 7132 fl (RB); Maguire, Pires, Maguire & Silva 56603 fl (NY); Pranceet al 6869 fl
(INPA, NY); 8464 fr (INPA, NY); 8912 fr (INPA, NY); N. T. Silva 346 fr (IAN). Mato Grosso:
Maguire, Pires, Maguire & Silva 56954 fl (NY). BOLIVIA. Pando: Pranceet al 6495 fl (INPA, NY);
8361 fl (INPA, NY).

This species is closely related to H. elongata,but differs in the rounded leaf base,
in the more pubescent lower surface of the leaf, in the sessile glands of the bracts, in
the smaller and fewer bracts, and in the tomentose epicarp and thicker endocarp of
the fruit. The fruit of this species is similar to that of H. guatemalensis.

25. Hirtella paniculata Swartz, Prod. Veg. Ind. Occ. 51. 1788; Vahl, Symb.
Bot. 2:43, t. 31. 1791; Persoon, Syn. I: 250. I805; Roem. & Schult., Linn.,
Syst. Veg. ed. 9. 5: 272. I819; Zucc., Flora I5(Beibl. 2): 85. I832; Sandw..
Kew Bull. 1931: 377. 1931; Kleinh. in Pulle, Fl. Suriname 2(I): 450. 1939,
Hirtella hirsutaLamarck, Illustr. 2: I I4. 1793; Roem. & Schult., Linn., Syst. Veg. ed. 9. 5: 272.
1819; DC., Prodr. 2: 528. 1825, nom illegit.
Hirtella aggregataPoiret, Encyc., Suppl. 2: 53. 1813, nom illegit.
Hirtella indecoraSchott ex Sprengel, Syst. 4(App): 404. 1827, nom illegit.
Hirtella bracteosaSteudel, Flora 26: 761. I843. Type. Hostmann60o, Suriname, fl (holotype, P;
isotypes, BM, C, CGE, F, GH, GOET, L, LE, M, NY, S).

Shrub to 4.0 m tall or small tree, the young branches pilose, becoming glabrous
with age. Leaves oblong-lanceolate to ovate-orbicular, most frequently ovate,
coriaceous, 2.5-I3.5 cm long, 1.3-5.5 cm broad, acuminate at apex, with acumen
1.0-7.0 mm long, rounded to subcordate at base, hirsute beneath at least on venation,
with palisade glands present towards base; primary veins 7-9 pairs, plane or nearly
so above, prominent beneath; midrib prominulous above, prominent and hirsute
beneath; petioles .o0-2.0 mm long, terete, eglandular, sparsely pilose. Stipules
3.0-I2.0 mm long, linear, persistent, sparsely hirsute. Inflorescences terminal and
axillary little-branched panicles, 3.0-26.0 cm long, or rarely reduced to racemes, the
rachis and branches sparsely hirsute-hispid. Bracts and bracteoles 2.0-6.0 mm long,
oblong to lanceolate, membraneous, sparsely hirsute, persistent, the margins bearing
small stipitate glands or sessile rather inconspicuousglands. Flowers 5.0-7.0 mm long.
Receptacle campanulate, hirsute on exterior, glabrous within except for deflexed
Hirtella 293

hairs at throat; pedicels I.0-3.0 mm long. Calyx lobes acute, hirsute on exterior,
gray-puberulous within, the margins eglandular. Petals 5, white to purple, glabrous.
Stamens 5-6, unilateral with a toothed portion of the circle opposite them, the
filaments purple, far exceeding calyx lobes. Style hirsute up to half its length. Ovary
inserted at or near mouth of receptacle, pilose. Fruit ellipsoid; epicarp smooth,
glabrous; mesocarp thin, fleshy; endocarp thin, hard, bony, hirsute within.
TYPE. Rohrsn, French Guiana, fl (holotype, BM, Banks Herb.; isotypes, C, LE).
DISTRIBUTION.Open sandy places, savannas, savanna margins, sandy river
banks and islands, in St. Vincent, Trinidad, Venezuela, the Guianas, and northern
Brazil. Flowering throughout the year.
Representative Collections: WINDWARD ISLANDS. St. Vincent: Andersonsn fl (K); Guilding
sn fl (BR, K). COLOMBIA. Meta: Garcia-Barriga& Jaramillo 17111 fl (COL). Vaupes: Cuatrecasas
7516 fl (COL. US); RomeroC. 3474 fl (COL); 3489 fl (COL). VENEZUELA. Anzoategui: Aristeguieta
4597 fl (VEN); Croizat sn fl (VEN); Herndndez56 fl (VEN); Tamayo3586 fl fr (VEN). Monagas:
Aristeguietaet al 7250 fl (NY, VEN). Terr. Delta Amacuro: Bond, Gillin & Brown o18 fl (NY, US);
Gines5071 fl (US); 5141 fl (US). Bolivar: Aristeguieta5860 fl (VEN); Bailey & Bailey 1373 fl (A, GH,
NY, US); Cardona1736fl (US, VEN); 2600 fl (NY, US, VEN); Grosourdy 13 fl (P); Maguire, Wurdack&
Bunting35896 fl (NY); Phelps & Hitchcock306 fl fr (NY, VEN); Steyermark57647 fl (F, NY, VEN);
86268 fl (US, VEN); Ll. Williams11884 fl (F, K, S, US, VEN); Wurdack34378 fl (NY). Terr. Amazonas:
Cowan & Wurdack31460 fl (NY); Maguire & Politi 27361 fr (NY); 27443 fl (NY); Maguire & Maguire
35087 fl (NY); 35183 fl (NY). Without precise locality: Rusby & Squires88 fl (A, BM, F, GH, LE,
MICH, NY); Stevenssn fl (NY). TRINIDAD. Bot. Gardeno1756 fl (NY); Brittonet al 89 fl (GH, K, NY,
US); 725 fl (K, NY, US); 2095 fl (NY, US); Broadway2751 fl (BR, F, G); 5583 fl (A, K, S); Eggerssn
fl (GH); 937 fl (GOET); 1357 fl (A, C, S); Fendler o119 fl (K); Howard 10329 fl (A, MICH, NY);
Kuntze 1112 st (NY); Prance 2110 fl (NY); Purdie sn fl (GH); Simmonds15018 fl (K); Vocke6086 fl
(GOET); R. 0. Williams r2494 fl (K). GUYANA. Alston525 fl (K, NY, P, RB); Appun378 fl (K); De
La Cruz 1069 fl (NY, US); I585 fl (F, GH, NY, US); 1736 fl (F, GH, K, NY, US); 2257 fl (F, GH, K,
NY, US); 2538 fl (F, GH, NY, US); 2707 fl (F, GH, NY, US); 2804 fl (F, GH, NY, US); 3831 fl
(F, GH, K, NY, US); 4174 fl (F, GH, NY, US); Fanshawe64o(F.D. 3376) fl (K); Harrison798 fl (K);
o020 fl (K); Hitchcock17063 fl (GH, NY, US); Hohenkerk723 fl (K); Jenman Iiio fl (K); 4900 fl (K);
5326 fl (K); Linderi 6 fl (GH, NY, S); Maguire & Maguire46261 fl (NY); Maguire,Maguire & Wilson-
Browne 45978 fl (NY, US); 46o09A fl (NY); Mell & Mell 214 fl (NY, US); Persaud5 fl fr (F); 28 fl
(F, K, NY, S); Sandwith285 fl (K, NY); Schomburgk 7 fl (BM, BR, F, K, LE, NY, P, US); 135 (68) fl
(BM, BR, F, K, OXF, P); A. C. Smith3409 fl (A, F, IAN, K, LE, NY, P, S, US); Tate 190 fl (K, NY);
Wilson-Browne262(F.D. 5716) fl (K, NY). SURINAME. B.B.S. 245 fl (K); B.W. 1668 fl (A, C);
4356 fl (IAN, K, NY); 5478 fl (A); 6578 fl (NY); Focke 643 fl (GH, K, U); Hitchcock17063 fl (S);
Hostmann78 fl (BM, LE, NY, OXF); Irwin, Prance,Soderstrom & Holmgren55218 fl (F, GH, NY, S, US);
55370 fl (F, NY, US); 55460 fr (F, GH, MICH, NY, S, US); Lanjouw& Lindeman562 fl (K, NY, U);
81o fl (NY); 3048 fl (IAN, K, NY, U); 3327 fl (NY, U); Maguire 23820 fl (A, BR, F, K, NY, US);
Maguire,Schulz, Soderstrom& Holmgren53923 fl (F, GH, NY, US); 54008 fl (F, GH, NY, US); Stockdale
8819 fl (K); Samuels129 fl (A, GH, K, NY, P); 298 fl (GH, K); Van Emdensn fl (NY); Weigeltsn fl
(C, F, LE). FRENCH GUIANA. Broadway346 fl (GH, NY, US); 428 fl (GH, NY, US); 814 fl (G
NY, US); Capricesn fl (NY); Halle 450 fl (P); Leblondsn fl (K, P); Leprieursn fl (A, BM, BR, F, GH,
IAN, K, LA, NY, P, R, US); Martinsn fl (BM, F); Melinonsn (I864) fl (A, BR, F, GH, IAN, K, LA,
NY, P, R, US); 33 fl (F); 218 fl (BM, F, GH, K, P, US); 274 fl (BM); 419 fl (P, R); Poiteausn fl (K,
LE, P); Sagot 205 fl (BM, BR, C, K, NY, P, S); Wachenheim143 fl (BM, F, K, P, US); 213 fl (K, P).
BRAZIL. Amazonas: Silva & Brazao 60609 fl (NY). Terr. Roraima: AlvarengaRB 90573 fl (RB); Black
51-12711 fl (NY); Black & Magalhaes5s-12968fl (IAN, NY); Ducke1341 fl (A, IAN, MG, NY, R, US);
KuhlmannRB 3511 fl (K, RB, S); Pranceet al 4017 fl (INPA, MG, NY); 9119 fl (INPA, NY); 9536 fl
(INPA, NY); Ule 7585 fl (F, frag, K, MG, US). Para: Sampaio5656 fl (R). Terr. Amapa: DuckeMG
4760 fl (MG, RB 15052, US); Maguire,Pires & Maguire47128 fl (F, GH, IAN, LE, MG, NY, US).

LOCALNAMES.Venezuela: Icaco, Icacillo. Guyana: Buku-buku. French Guiana:

Bois Gaulette.
This species displays considerable variation in leaf shape but uniformity in the
flowers and inflorescence. As the foliar variation has no geographical correlation,
there is no basis for meaningful subspecific taxa.
In the original description, Swartz marked H. paniculatawith an asterisk, thus
indicating that the species was based on a type in the Banks Herbarium, rather than
294 Flora Neotropica

on a specimen of Swartz's own herbarium. Since Swartz worked with the Banks
Herbarium for some time, Rohrsn from the Banks Herbarium is the type for H.
paniculata,rather than the material of this species at Stockholm marked "Swartz
Herbarium". The Stockholm specimens may be isotypes, but they bear no collector's
name or date.
The name H. hirsuta,which has been much applied to this species, was created
by Lamarck, but is a later homonym and cannot be used.

26. Hirtella deflexa Maguire, Fieldiana Bot. 28: 255. 1952.

Small tree or shrub, the young branches pilose, becoming glabrous with age.
Leaves ovate, subcoriaceous, 4.o-8.0 cm long, 2.0-4.0 cm broad, acute to acuminate
at apex, with acumen 0-5.0 mm long, cordate at base, sparsely hirsute beneath;
primary veins 9-12 pairs, plane or almost so above, prominent beneath; midrib
prominulous above, prominent and hirsute beneath; petioles I.0-2.0 mm long, terete,
eglandular, hispid. Stipules to 6.0 mm long, linear, hispid, subpersistent,eglandular.
Inflorescences terminal and predominantly axillary little-branched panicles, 4.5-
I0.0 mm long, the rachis and branches sparsely hirsute. Bracts and bracteoles 2.0-
3.0 mm long, oblong, membraneous, persistent, the margins bearing slender stipitate
glands. Flowers 5.0-6.0 mm long. Receptacle campanulate, sparsely hirsutulous on
exterior, glabrous within except at throat; pedicels 2.0-3.5 mm long. Calyx lobes
rounded to acute, hirsutulous on exterior, gray-pubescent within, the margins
eglandular. Petals 5, glabrous, white. Stamens 4-6, unilateral, the filaments reddish
purple, far exceeding calyx lobes. Style hirsute on lower portion only. Ovary inserted
midway up receptacle, villous. Fruit not seen.
TYPES.Pinkus 46, Venezuela, Bolivar, fl (holotype, NY; isotypes, F, S, US);
Steyermark 60250, Venezuela, Bolivar, fl (paratypes, F, NY, US).
DISTRIBUTION. River banks in Bolivar, Venezuela. Known only from the type
collections. Flowering November to January.
This species is very close to H. paniculata,but differs in the deeply cordate leaf
bases, and in the predominantly axillary inflorescences.

27. Hirtella tentaculata Poeppig & Endlicher, Iov. Gen. & Sp. P1. 3: 75.
1827.

Tree to 20.0 m tall, the young branches tomentellous, becoming glabrous and
lenticellate with age. Leaves elliptic to oblong-elliptic, coriaceous, 7.0-24.0 cm long,
3.5-10.5 cm broad, acuminate at apex, with acumen 2.0-10.5 mm long, rounded at
base, with impressed venation above, hirsute beneath, with a few palisade glands to-
wards base; primary veins I0-13 pairs, impressed above, prominent beneath;
midrib plane above, prominent and hirsute beneath; petioles 3.0-8.0 mm long,
tomentellous, caducous, eglandular. Stipules 2.0-3.0 mm long, ovate, tomentose,
persistent. Inflorescencesdensely-floweredterminal and axillary panicles 6.0-25.0 cm
long, the rachis and branches tomentellous to puberulous. Bracts and bracteoles
i.o-6.o mm long, oblong to lanceolate; glands numerous, long-stalked, stipitate,
arising from pedicels and axils of flowers, on stalks up to 8.0 mm long. Flowers
4.0-6.0 mm long. Receptacle campanulate, puberulous on exterior, glabrous within
Hirtella 295

except at throat- Calyx lobes rounded to acute, puberulous on exterior, gray-pubes-

cent within, the margins eglandular. Petals 5, glabrous. Stamens 3-5, unilateral, with
short filamentous staminodes opposite them; filaments glabrous, far exceeding calyx
lobes. Style hirsute up to one third its length. Ovary inserted at mouth of receptacle,
pilose. Fruit ellipsoid; epicarp smooth, glabrous; mesocarp thin, fleshy; endocarp
thin, hard, bony, hirsute within.
TYPE.Poeppigsn, Brazil, Para, fl fr (holotype, W; isotype, LE; isotype, fragment
F).
DISTRIBUTION. Non-flooded or periodically flooded forest in eastern Venezuela,
Guyana, and Para, Brazil. Flowering May to November.
VENEZUELA. Bolivar: Bernardi769 fl (K, NY, VEN); Cardona1210 fl fr (F, NY, US, VEN);
1665 fl (NY, US, VEN); Steyermark60561 fl (F, NY, US); 60639 fr (F, NY). GUYANA. Maguire &
Fanshawe32217 fl (NY); Tillett & Tillett 44969 fl (NY); 45488 fl (NY); 45701 fl (NY). BRAZIL. Pari:
Baker MG 4I7 fr (BM, F, MG); Ducke 1251 fl (IAN, K, MG, NY, R, US); RB 18237 fl (RB); Fr6es
31771 fl (IAN); 31836 fl (IAN); HuberMG 3819 fl (MG); Museu Goeldi9792 fl (MG, US); Pires &
Silva 10259 fl (IAN); A. Silva 285 fl (IAN, NY, US); Snethlage71 fl fr (F, GH, US).

LOCALNAMES.Venezuela: Can-wadak-yek.Brazil: Ajuruirana.

This species was omitted by Hooker in Flora Brasiliensis.It is related to H.
but differs in the impressed venation of the upper surface of the leaf,
macrophylla,
and in the more pubescent lower surface.

28. Hirtella macrophylla Bentham ex Hooker f., Mart. Fl. Bras. I4(2): 36,
t. 12. 1867.

Small tree, the young branches tomentellous, soon becoming glabrous and
lenticellate. Leaves elliptic, coriaceous, io.o-i8.ocm long, 5.o-9. cm broad,
acuminate at apex, with acumen 3.0-10.0 mm long, rounded to subcuneate at base,
sparsely hirsute beneath on primary venation, with palisade glands frequent towards
base; primary veins 7-10 pairs, prominulous above, prominent beneath; midrib
plane above, prominent and hirsute beneath; petioles 3.0-10.5 mm long, tomen-
tellous, terete, eglandular. Stipules 2.0-3.0 mm long, subulate, persistent, tomentose.
Inflorescences terminal and axillary panicles IO.O-30.0cm long, the rachis and
branches tomentellous to sparsely puberulous. Bracts and bracteoles to 8.0 mm
long, oblong, puberulous, persistent, with numerous long-stalked glands arising from
pedicels and axils of flowers. Flowers 5.0-6.0 mm long. Receptacle campanulate,
puberulous on exterior, glabrous within except at throat. Calyx lobes rounded,
sparsely puberulous on exterior, gray-pubescent within, the margins eglandular.
Petals 5, glabrous. Stamens 4-6, unilateralwith a toothed portion of arc opposite them;
filaments glabrous, far exceeding calyx lobes. Style hirsute up to half its length. Ovary
inserted at mouth of receptacle, pilose. Fruit ellipsoid; epicarp smooth, glabrous;
mesocarp thin, fleshy; endocarp hard, thin, bony, hirsute within.
TYPE. Spruce3095, Venezuela, Terr. Amazonas, fl fr (holotype, K; isotypes,
BM, BR, CGE, LE, NY, OXF, P).
DISTRIBUTION. River banks and secondary forest in Amazonian Colombia,
Venezuela and Brazil.
COLOMBIA. Putumayo: Klug I948 fl (BR, COL, F, GH, K, MICH, NY, P, S, SP, US).
Amazonas: Schultes & Cabrera12669 fl (COL, US); 13622 fl (A, COL, GH, NY, US). BRAZIL.
Amazonas: CoelhoINPA 3486 fl (IAN, INPA, MG, NY); INPA 3885 fl (IAN, INPA, MG, NY); INPA
3975 fl (IAN, INPA, MG, NY); Ducke 705 fl (F, IAN, MG, NY, R, RB, US); Rodrigues& Chagas
2359 fl (INPA).
296 Flora Neotropica

is close to H. tentaculata
Hirtellamacrophylla and H. glabrata,but for differencessee
under those species.

29. Hirtella adderleyi Prance, sp nov Fig 48 D-G.

Frutex parva, ramulis juvenilibus tomentosis. Folia alternata petiolata, petiolo

i.0-2.0 mm longo, tomentoso, tereti, eglanduloso; laminae ellipticae, coriaceae,
5.5-7.5 cm longae. 2.5-3.7 cm latae, apice in acuminem obtusum 2.0-4.0 mm longum
contractae, basi rotundatae vel subcordatae, subconduplicatae, subtus cum pilis
sparse appressismunitae, supra reticulatae; costa media utrinque prominente, subtus
hirsuta; costis secundariis I0-I2 jugis, utrinque leviter prominentes. Stipulae
lineares ad 5.0 mm longum, pubescentes, eglandulosae, persistentes. Flores 6.0-7.0
mm longi in paniculis terminalibus 4.0-7.0 cm longis dispositi; rachi ramisque
tomentosi vel puberuli. Bracteae bracteolaeque 2.0-3.5 mm longae, ovatae, tomen-
tosae, persistentes, cum glandulis stipitatis numerosis instructae. Receptaculum
campanulatum extus tomentosum, intus glabrum; pedicelli circa 0.5 mm longi.
Calycis-lobi acuti, extus tomentosi, intus puberuli, eglandulosi vel glandulosi. Petala
5, rosea, glabra. Stamina 4-5, unilateralia, staminodiis dentatis opposita; filamenta
glabra, longe exserta. Ovarium ad faucem receptaculi insertum, pilosum. Stylus e
basi ovarii ortus, praeter basim glaber. Drupa ellipsoidea, epicarpio laevi, glabro;
mesocarpio tenue, carnoso; endocarpio tenue, duro, osseo, intus hirsuto.
TYPE. Wurdack & Adderley 43716, Venezuela, Terr. Amazonas: Rio Orinoco,
Sabana de Mogo, fl fr (holotype, NY).
Savannas of the upper Orinoco region of Venezuela, flowering
DISTRIBUTION.
June and July.
VENEZUELA. Terr. Amazonas: Rio Atabapo, Sabana Manacal, Wurdack& Adderley42973 fl
(NY).

This distinct species is not easily confused with the others having a glandular
paniculate inflorescence.

30. Hirtella punctillata Ducke, Arch. Jard. Bot. Rio de Janeiro 3: 268. 1922.

Tree to 20.0 m tall or shrub, the young branches puberulous, soon becoming
glabrous. Leaves orbicular to oblong, coriaceous, 2.2-8.0 cm long, 1.4-6.0 cm broad,
retuse to rounded or minutely mucronate at apex, rounded to subcordate at base, the
lower surface glabrous with frequent palisade glands; primary veins 5-7 pairs,
prominulcus on both surfaces; midrib prominulous above, prominent and sparsely
appressed-pubescentbeneath; petioles I.0-2.0 mm long, terete, eglandular, sparsely
puberulous when young. Stipules ca i.o mm long, lanceolate, glabrous, eglandular,
persistent. Inflorescences short terminal corymbose panicles I.0-4.0 cm long, the
rachis and branches tomentellous. Bracts and bracteoles I.0-2.5 mm long, deltoid,
acute, tomentellous, persistent, eglandular. Flowers 5.0-6.0 mm long. Receptacle
campanulate, sparsely tomentellous on exterior, glabrous within except for retrorse
hairs at throat; pedicels 0.5-I.5 mm long. Calyx lobes acute, sparsely tomentellous
on exterior, gray-puberulous within. Petals 5, glabrous, pink. Stamens 5-6,
unilateral with small filamentous staminodes opposite them. Style hirsute up to one
third its length. Ovary inserted at mouth of receptacle, tomentose. Fruit ellipsoid;
Hirtella 297

epicarp smooth, glabrous; mesocarp thin, fleshy; endocarp thin, hard, bony, densely
hirsute within.
TYPE.DuckeMG 16663, Brazil, Para, fl (holotype, MG; isotypes, K, NY, RB
15044, US).
DISTRIBUTION.Sandy savannas, open and low forest in sandstone areas, and open
river margins in Venezuela, the Guianas, and eastern Amazonian Brazil. Flowering
August to November in Guianas and March to August in Amazonia.
VENEZUELA. Bolivar: BernardiI647 fl (COL); Cardona840 fl (US, VEN); 854 fl (F, NY, US);
1756 fl (G); 1771 fl (NY, US, VEN); 1789 fl (VEN); sn fl (VEN); Maguire, Steyermark& Maguire
53690 fl fr (NY); Steyermark98191 fl (NY, VEN). Terr. Amazonas: Barnes 85 fl (VEN). GUYANA.
Maguire & Fanshawe32623 fl (NY). SURINAME. Maguire 24392 fl (A, F, K, NY, US). BRAZIL.
Amazonas: Froes 22840 fl (IAN, UB). Pari: DuckeRB 18233 fl (RB, S). Terr. AmapA: Froes & Black
27718 fl (IAN); Pires & Cavalcante52191 fl (IAN, MG, NY).

LOCALNAME.Venezuela: Wada (Maquiritare).

Hirtellapunctillatais close to H. scabra,but differs in the blunt leaf apex. It is
closest to H. corymbosa,but for differences see under that species.

3I. Hirtella corymbosa Chamisso & Schlechtendal, Linnaea 2: 545. 1827;

Zucc., Flora 15(Beibl. 2): 87. 1932; Hook. f., Mart. Fl. Bras. 14(2): 37.
1867.

Low shrub, the young branches tomentellous becoming glabrous with age.
Leaves elliptic, coriaceous, 3.0-5.5 cm long, 1.5-3.0 cm broad, acute at apex, rounded
to subcordate at base, glabrous beneath, with a few palisade glands towards base;
primary veins 7-8 pairs, prominulous on both surfaces; midrib prominent on both
surfaces, appressed-pubescentbeneath; petioles 0.5-I.O mm long, terete, eglandular,
sparsely tomentellous. Stipules ca i.omm long, ovate, tomentellous, eglandular,
caducous. Inflorescences short terminal corymbose panicles to 3.0 mm long, the
rachis and branches sparsely tomentellous. Bracts and bracteoles I.0-2.0 mm long,
oblong-triangular, acute, puberulous, persistent, eglandular. Flowers 4.0-6.0 mm
long. Receptacle campanulate, puberulous on exterior, glabrous within except for
retrorse hairs at throat. Calyx lobes rounded to acute, puberulous on both surfaces.
Petals 5, glabrous. Stamens 3-4, unilateral. Style glabrous except at base only. Ovary
inserted near mouth of receptacle, pilose. Fruit ellipsoid (young fruit only seen);
epicarp smooth, glabrous; mesocarp thin, fleshy; endocarp thin, bony, hirsute
within.
TYPE.Sellowsn, Brazil, fl (holotype, LE; isotype fragments CGE, K, P).
Known only from the State of Espirito Santo, eastern Brazil.
DISTRIBUTION.
Growing in open savanna.
BRAZIL. Espirito Santo: Belem3837 fl (IAN, NY); Duarte9747 fl (RB 13161); Kuhlmann6672 fl
(NY, RB 62934).

The Sellow material cited above is without collector's number. One Berlin sheet,
now lost, but seen as a photograph, bore the inscription Sellow 45. The British
Museum has a sheet of this species marked Sellow 856. It is uncertain whether the
Sellow material represents more than one gathering. In addition, there is a sheet of
H. corymbosain the Praha herbarium (PRC) marked "Herb Tausch", which is
probably part of the type material.
Hirtellacorymbosais closest to H. punctillata,but differs in the fewer stamens, the
acute leaf apex, and the glabrous style.
298 Flora Neotropica

32. Hirtella pendula Solander ex Lamarck, Encyc. 3: 134. I789; Roem. &
Schult., Linn. Syst. Veg. ed. 9. 5: 273. I819; DC., Prodr. 2: 528. I825.

Tree to I2.0 m tall, the young branches tomentellous becoming glabrous with
age. Leaves oblong, membraneous-chartaceous,6.5-I3.5 cm long, 2.5-4.5 cm broad,
acuminate at apex, with acumen 2.0-15.0 mm long, subcordate at base, the lower
surface glabrous or with a few appressed hairs when young, markedly papillose, with
a few palisade glands towards base; primary veins 9- 1 pairs, prominulous on both
surfaces; midrib prominulous above, prominent and hirsute beneath; petioles 2.0-
4.0 mm long, terete, eglandular, tomentellous. Stipules linear, 2.0-4.0 mm long,
puberulous, persistent, eglandular. Inflorescences long, pendulous, little-branched
terminal panicles 25.0-55.0 cm long, with a long central rachis bearing small lateral
branches, the rachis and branches tomentellous to sparsely puberulous. Bracts and
bracteoles I.0-3.5 mm long, lanceolate, tomentellous, subpersistent, eglandular.
Flowers 5.0-7.0 mm long. Receptacle obliquely campanulate, short-tomentellous on
exterior, glabrous within except at throat; pedicels 7.0-12.5 mm long. Calyx lobes
rounded, tomentellous on both surfaces. Petals 5, glabrous. Stamens 3, unilateral
with small teeth opposite them, the filaments glabrous, far exceeding calyx lobes. Style
hirsute up to one third its length. Ovary inserted near mouth of receptacle, pilose.
Fruit ellipsoid; epicarp smooth, glabrescent; mesocarp thin, fleshy; endocarp thin,
hard, fibrous, hirsute within.
TYPE. Thouin,par Aitonsn, Antilles, fl (holotype, P-JUSS).
DISTRIBUTION. Primary forest in the Lesser Antilles from Guadeloupe to St.
Lucia. Flowering April to July.
LEEWARD ISLANDS. Guadeloupe: Duss 3255 fl (C, F, NY, US); 3255a fr (NY); 4147 fl
(NY); 4209 fl (NY); Forsstromsn fl (S); Holdridge437 fr (NY); Stehle936 fl fr (US). WINDWARD
ISLANDS. Martinique: Duss 1904 fl (NY, US). St. Lucia: Anderson(Herb Forsyth)sn fl (G-DC, K);
J. Beard i66 fr (A, K, NY); P. Beard Io50 fl (GH, S, US); Box 1747 fl (BM); Cooley8735 fl (GH);
Howard 11640 fr (A, BM); Ramagesn fl (BM, K); Webster,Ellis & Miller 9296 fl (MICH).

LOCAL NAMES AND USES. Gaudeloupe: Icacquemontagne.St. Lucia: Pend oreille.Fresh

fruit is used as ear pendants for adornment in St. Lucia.
This species may be distinguished from all other species of Hirtella by the long
peduncles of the inflorescence in which the flowers are borne obliquely at the end.

33. Hirtella barrosoi Prance, sp nov Fig 46.

Arbor parva, ramulis juvenilibus glabris, lenticellatis. Folia alternata petiolata;

petiolo 2.0-3.0 mm longo, tereti, eglandulso, cum pilis appressis sparse pubescenti;
laminae coriaceae, lanceolatae, 4.5-8.5 cm longae, 1.3-2.0 cm latae, apicem versus
gradatim contractae in acuminem 8.0-20.0 mm longum, basi cuneatae, subtus cum
pilis appressispaucis munitae, pauciglandulosae; costa media supraleviter prominente,
glabrescente; costis secundariis IO-I4 jugis, utrinque leviter prominentibus. Stipulae
circa 1.5 mm longae, lineares, eglandulosae, caducae. Flores 4.0-5.0 mm longi, in
paniculis terminalibus axillaribusque 3.0-6.0 cm longis dispositi, rachi ramisque
sparse pubescentibus. Bracteae bracteolaeque 1.0-2.0 mm longae, acutae, ovatae,
eglandulosae, persistentes. Receptaculum campanulatum, extus sparse puberulum,
intus basim versus glabrum; pedicelli 1.0-2.0 mm longi. Calycis-lobi acuti, extus
sparse puberuli, intus puberuli. Stamina 3, unilateralia; filamenta glabra, longe
exserta. Ovarium ad faucem receptaculi insertum, pilosum. Stylus e basi ovarii
Hirtella 299

k~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~..

~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~..."' a

.......

FIG 46. Species of Hirtella. A-G, H. barrosoi (Horto Florestal RE 136922, Vitdrio RE 1369r9);~~~~~~
A, hait, 0.5 B, ruit X 05; C frut setion X 05; D floer, 4~;E, oary,X
F,sepa,.x.,," I ;

floersecio, 6. -M H.stula (irs 613 6g3) H K,flw?

flwe, g; , var, rg
secion x o;L, abi, 0.......ngfrit X

FIG 46. Species of Hirtella. A-G, H. barrosoi(Horto Florestal RB 136922, Vitorio RB 136919);
A, habit, x o.5; B, fruit, x o. 5; C, fruit section, x o.5; D, flower, x 4; E, ovary, x 15; F, sepal, x 2.5; G
flower section, x6. H-M, H. suffulta (Pires 6813, 6943); H, flower, x 5; J, ovary, x i5; K, flower
section, x o; L, habit, xo.5; M, young fruit, x i.
300 Flora Neotropica

ortus, glabrus basus exceptus. Drupa globosa, 1.5-2.0 cm diametro; epicarpio laevi,
glabro; mesocarpio tenue, carnoso; endocarpio tenue, duro, osseo, intus tomentello.
TYPE. HortoFlorestalRio de JaneiroRB 136922, Brazil, Guanabara, near Horto
Florestal, Rio de Janeiro, fl (holotype, NY; isotype, RB).
DISTRIBUTION. Known only from two collections from the coastal forest around
Rio de Janeiro, Brazil. Flowering May, fruiting August.
BRAZIL. Guanabara: Horto Florestal, Rio de Janeiro, VitorioRB 136919 fr (NY, RB).

This species is most closely related to H. bicornis,but differs in the narrow lanceo-
late leaves with a gradually tapered apex.
It is with pleasure that I name this species for Dra. Graziela M. Barroso,who is
contributing so much to the botany of central and southern Brazil.

34. Hirtella leonotis Pittier, Contr. Fl. Venez. 23. 1923.

Shrub or small tree, the young branches hispid glabrescent. Leaves oblong,
subcoriaceous, II.5-15.5 cm long, 3.5-6.0 cm broad, acuminate at apex, with acu-
men 7.0-I2.0 mm long, rounded at base, sparsely appressed-pubescenton venation
beneath, the palisade glands rare, papillose; primary veins IO-12 pairs, plane above,
prominulous beneath; midrib prominulous above, prominent and hirsute bzneath;
petioles 3.0-5.0 mm long, terete, eglandular, hispid. Stipules linear, tomentellous,
6.o-Io.o mm long, persistent, eglandular. Inflorescences terminal and subterminal,
scarcely branched panicles or simply racemes, I0.O-I5.0 cm long; the rachis and
branches hispid-hirsute. Bracts and bracteoles 3.0-5.0 mm long, oblong, acute,
hispid, persistent, the margins bearing few sessile glands or eglandular. Flowers
7.0-8.0 mm long. Receptacle campanulate, hirsute on exterior, glabrous within
except at throat; pedicels 3.0-6.0 mm long. Calyx lobes rounded to acute, hirsute on
exterior, gray-puberulous within. Petals 5, glabrous. Stamens 3, unilateral, with small
teeth opposite them, the filaments glabrous, far exceeding calyx lobes. Style hirsute
at base only. Ovary inserted at mouth of receptacle, densely pilose. Fruit unknown.
TYPE. Jahn 459, Venezuela, Dist. Federal, fl (holotype, VEN; isotypes, GH, NY,
US).
DISTRIBUTION. Known only from the type gathering collected in Valle de Puerto
La Cruz, Venezuela.

The inflorescence in this species is essentially a raceme with flowers borne on

long peduncles. A few peduncles, however, are branched and bear two flowers. This
species is close to H. triandra, but differs in the inflorescence, and in the much larger
flowers.

35. Hirtella mutisii Killip & Cuatrecasas, Fieldiana Bot. 27: II2. 1951.

Tree to 20.0 m tall, the young branches tomentellous. Leaves oblong to oblong-
ovate, subcoriaceous, 6.0-I4.0 cm long, 2.7-5.0 cm broad, acuminate at apex,with
acumen 5.0-Io.o mm long, rounded at base, the lower surface papillose, sparsely
hirsute, with a few palisade glands towards base; primary veins 9-I3 pairs, plane
above, prominulous beneath; midrib prominulous above, prominent beneath,
tomentellous on both surfaces; petioles 1.5-2.5 mm long, terete, tomentellous,
Hirtella 301

eglandular. Stipules linear, to 4.0 mm long, tomentellous, persistent, eglandular.

Inflorescences scarcely branched panicles 3.0-8.0 cm long, most branches bearing a
single flower and one pair of bracts at junction of branch and pedicel, the inflores-
cence thus appearing racemose, but a few branches bearing two flowers, the rachis
and brancheshirsutulous-tomentellous.Bractsand bracteoles .0-2.5 mm long, oblong,
acute, tomentellous, glabrous within except at throat; pedicels 2.0-5.0 mm long.
Calyx lobes rounded, tomentellous on exterior, gray-puberulous within. Petals 5,
white, glabrous. Stamens 3, unilateral, with short toothed staminodes opposite them;
filaments glabrous, far exceeding calyx lobes. Style hirsute up to one third its length.
Ovary inserted at mouth of receptacle. Fruit not seen.
TYPE. Mutis 3554, Colombia, fl (holotype, US); 23, Colombia, fl (paratype,
US).
DISTRIBUTION. Primary forest in central and southern Colombia and in Ecuador.
Flowering throughout the year.
COLOMBIA. Antioquia: Haught 4710 fl (COL, US). Santander: Haught 1468 fl (COL, GH,
K, NY, US, W). Without precise locality: Mutis 11og fl (US). ECUADOR. Cazalet & Pennington51o4
fl (K, US); Eggers 1428 fl (A, LE); Rimbach59 fl (A); 97 fl (A, F, K, NY, US).

This species is very close to H. triandra,but differs in the inflorescence which is

scarcely branched and thus has a racemose appearance. It is also close to H. leonotis,
but differs in the smaller flowers, stipules and leaves, and in the hirsute (not hispid)
inflorescence and young branches.

36. Hirtella triandra Swartz, Prod. Veg. Ind. Occ. 51. 1788; Fl. Ind. Occ.
I: 508. I797; Willd., Linn., Sp. P1. ed. 4. I: 1151. 1797; Persoon, Syn.
I: 250. i805; DC. Prodr. 2: 528. I825. Schlecht., Linnaea 2: 542. 1827;
Zucc., Flora I5(Beibl. 2): 84. 1832; Griseb., Fl. Brit. W. Ind. 230. i86o;
Hook. f., Mart. Fl. Bras. 14(2): 37. I867; Kleinh. in Pulle, Fl. Suriname
2(I): 449. I937; Macbride, Fl. Peru 1079. 1938.

Tree to I5.0 m tall, usually smaller, or shrub, the young branches pilose-tomen-
tellous to puberulous, becoming glabrous with age. Leaves oblong to elliptic, sub-
coriaceous to membraneous, 4.0-I4.5 cm long, 2.0-5.5 cm broad, acuminate
(very rarely acute) at apex, with acumen I.o-i8.o mm long, rounded to cuneate at
base, papillose beneath, with a few stiff appressed hairs, the palisade glands few;
primary veins 5-I3 pairs, plane or prominulous above; midrib prominulous above,
prominent and hirsute beneath; petioles 1.0-3.0 mm long, terete, eglandular,
tomentellous to puberulous. Stipules linear, 2.0-6.0 mm long, tomentellous, persis-
tent, eglandular. Inflorescences terminal and axillary panicles, 3.5-I7.0 cm long,
with most branches bearing more than one flower and more than one pair of bracts,
the rachis and branches usually short-tomentellous, less frequently short-pilose-
hispid. Bracts and bracteoles 1.0-3.5 mm long, lanceolate to ovate, acute, persistent,
eglandular, sometimes with numerous bracts along inflorescence branches. Recep-
tacle campanulate, tomentellous on exterior, glabrous within except at throat;
pedicels 1.0-3.0 mm long. Calyx lobes acute, puberulous on both surfaces. Petals 5,
glabrous. Stamens 3, unilateral with teeth or short filamentous staminodes opposite
them, the filaments glabrous, far exceeding calyx lobes. Style hirsute up to half its
length. Ovary inserted at mouth of receptacle, pilose-tomentose. Fruit ellipsoid;
epicarp glabrescent or puberulous, less frequently tomentose, slightly longitudinally
canaliculate when dry; mesocarp thin, fleshy; endocarp thin, hard, bony, hirsute
within.
302 Flora Neotropica

Key to the Subspecies of Hirtellatriandra

i. Leaves with acumen 5.0-20.0 mm long, the laminas 4.0-15.0 cm long, subcoriaceous;
primary veins 9-13 pairs. a. subsp triandra.
i. Leaves with acumen 0.4.0 mm long, the laminas 3.5-8.0 cm long, often thick-coriaceous, most
frequently with acute apices; primary veins 5-9 pairs.
2. Leaves thick-coriaceous, with abrupt acumen; primary veins 7-9 pairs; inflorescence
with a central rachis and small lateral branches. b. subsp punctulata.
2. Leaves subcoriaceous, with acute apex, primary veins 5-7 pairs; inflorescence small,
compact, and much branched. c. subsp media

36a. Hirtella triandra subsp triandra

Hirtella triandraSwartz, Prod. Veg. Ind. Occ. 51. 1788.
Chrysobalanus triandra(Swartz) Morales, Ann. Acad., Ci. Med. Fisic. & Nat. Habana 43: 390.
1887.
Hirtella americanaJacquin, Select. Am. 8, t. 8. 1763; non L. (1753). Type. not seen, Martinique.
Hirtella cosmibuenaLamarck, Illustr. 2: 114. I793; Roem. & Schult., Linn., Syst. Veg. ed. 9.
5: 273. 1819; DC., Prodr. 2: 528. 1825; Zucc., Flora I5(Beibl. 2): 85. 1832. Type. Pavon
sn, Ecuador or Peru, fl (BM, F, MA; photo US).
Hirtella racemosaRuiz & Pavon, Fl. Peru 3: 5, t. 227. 1802, nom illegit., non Lam (1789). Type.
Pavonsn.
HirtellaperuvianaPersoon, Syn. I: 250. I805. Type. Pavonsn.
Sphenistaperuviana(Ruiz & Pavon) Rafinesque, Sylva Tellur. 90. 1838. Type. Pavonsn.
Zamzela racemosaRafinesque, Sylva Tellur. 90. 1838. Type. Herb. Swartz sn.
Hirtella paniculataLamarck, Encyc. Suppl. 3: 33. I813, non Swartz (I788). Type. Dupuis sn,
Antilles, fl (holotype, P-LAM).
Hirtella castaneaA. P. De Candolle, Prodr. 2: 528. 1825; Zucc., Flora 15(Beibl. 2): 82. 1832,
synon nov. Type. Plate 304 in A.L.P.P. de Candolle, Calques de Dessins. Fl. Mex. 1874.
HirtellabracteataMartius & Zuccarini, Abh. Akad. Miinchen I: 384. 1832; Zucc., Flora 15(Beibl. 2):
84. 1832; Hook. f., Mart. Fl. Bras. 14(2): 38. 1867, synon nov. Type. Martius sn, Brazil,
Fara, fl (holotype, M; isotypes, BR, K).
HirtellajamaicensisUrban, Symb. Antill. 5: 355. I908; Fawcett & Rendle, Fl. Jamaica 3: 256.
I914, synon nov. Types. Hansensn, Jamaica, fr (C, LD); Harris5604, Jamaica, fr (A, BM, F,
NY, US).
Hirtella multifloraUrban, Symb. Antill. 5: 356. 1908; Fawcett & Rendle, Fl. Jamaica 3: 256.
1914, synon nov. Types. Harris5317, Jamaica, fl (BM, C, NY, P); 5417 (not seen); 5966 fi
(NY, S); Purdiesn fl (K, NY, S).

Leaves 4.0-I5.0 cm long, subcoriaceous, with acumen 5.0-20.0 mm long;

primary veins 9-13 pairs; inflorescence little to much branched.
TYPE.Herb.Swartzsn, West Indies, fl (holotype, S; isotype, M).
Occurring in a wide range of habitats, but most common in open
DISTRIBUTION.
forest, river margins, sea shores, and wooded slopes, from central Mexico through
Central America, the Greater and Lesser Antilles and northern and western South
America to Bolivia and Amazonian Brazil.
Representative collections: MEXICO. Veracruz: Miranda8324 fl (MEXU); 8324 bisfl (MEXU);
Santos 2870 fl (MICH). Oaxaca: Schulteset al 851 fl (ECON). Chiapas: Matuda 607I fl (F, MEXU);
Miranda 7736 fl (MEXU). Without precise locality: Herb Pavonsn fl (BM). GUATEMALA. Holdridge
2306 fl (F); Skutch2088 fl (A, NY, US); Steyermark 7659 fl (F, GH, K, NY,
46356 fr (F); VonTuerckheim
US); 7942 fl (GH, US); 8514 fl (F, GH, K, NY, US); Watsonsn fl (GH); 294 fl (US). HONDURAS.
Molina R. 606i fl fr (F, US); 6076 fl (F, US); L. O. Williams & Molina R. 17968 fl (F, MO, US);
Tuncker,Koepper& Wagner8760 fl (F, GH, MO, NY). BRITISH HONDURAS. Schipp 1257 fl (A,
BM, F, GH, K, MICH, NY); Stevenson86 fl (F). NICARAGUA. Bunting & Licht r91 fl (NY); Shank
& Molina R. 4776 fl (F, GH). COSTA RICA. Skutch5388 fl (MICH, US); Standley& Valerio44551 fl
(US); 45782 fl (A, US); 45916 fl (US); 46634a fl (US); Tonduz94o1 fl (BR, US). PANAMA. Allen
967 fl (GH, MO, NY, US); 1689 fl (F, GH, NY, US); 3453 fl (F, GH, K, MICH, MO, NY); Aviles 66
fl (F); 949 fl (F); Bailey & Bailey 407 fl (F); Bangham398 fl (A, F, MICH); Cooper286 fl (F, NY, US);
Hirtella 303

Cooper& Slater 279 st (F); Dodge & Allen 17316 fl (C, F, FI, K, MICH, MO, NY, US); Duke 4896 fr
(GH, MO); 5826 fr (MO); 5827 fl (GH, MO); Kluge 48 fl (F, US); Pittier 2680 fl (BM, C, F, GH, K,
NY, US); 2784 fl (BM, C, NY, US); Shattuck519 fl (F); Sternet al 133 st (A, F, MO, NY, US); Tyson
et al 4644 fl (MO); Wetmore& Abbe 198 fl (A, F, GH); R. S. Williams 667 fl fr (NY, US); Woodson&
ScheryIoo8 fl (F); Woodworth& Vestal370 fl (A, F); 647 fr (A, F). CUBA. Alain 3641 st (NY); 3643 st
(NY); Brittonet al 4731 fl (NY); Clemente2953 fl (GH); 5242 fl (GH, US); Eggers 4665 fl (A, GOET,
LE); Ekman 2614 fl (NY, US); 4003 fl (S); 4080 fl (S); Howard5622 fr (A, F, GH, MICH, NY);
Leon 10970 st (NY); 22663 fl (F, GH); Leon & Rocha7909 fl (NY, P); Luna212 fl (NY); Pollard,Palmer
& Palmer213 fl (NY, US); Senn351 fl (GH); Shafer1640 fl (F, NY, US); I76o fl fr (F, NY); 3676 fl fr
(F, NY, US); 4404 fl (NY); Wright 155 fl fr (BR, GH, GOET, K, LE, NY, P, US). JAMAICA.
Alexandersn fl fr (C, GH, GOET, K, NY, P); Britton3524 fl (NY); 4176 fl (NY); Campbell478 fl (F,
NY); Harris & Britton 10753 fl (F, K, NY, US); Macfaydensn fl (GH, K, NY); Marsh 644 fl (K);
Perkins1358 fr (GH, K); Proctor923I fl (A); 23363 fl (GH, MICH, NY); 23954 fl (A, MICH); Wilson
sn fl (GOET). HISPANIOLA. Haiti: Ekman 2583 st (S); 3250 St (S); 10221 fl fr (S, US); Holdridge
2158 fl (MICH); Leonard& Leonard12417 fr (A, GH, NY, US). Santo Domingo: Abbott269 fl (GH,
NY, US); 412 fl (GH, US); 1163 fl (US); 2155 fl (GH); 2531 fl (NY, US); 2808 fl (GH, US); Augusto
I457 fl (A, NY); Eggers 1549 fl (A, BR, C, F, GOET, LD, LE, MICH, NY, P, S, US); Ekman 12247
fl (A, C, K, S, US); Howard9789 fl (GH); Fuertes332 fl (A, LE); 1650 fl (A); PrendleloupI70 st (NY,
US); Rose, Fitch & Russell 4344 fl (NY, US); Taylor 285 fl (F, NY); Valeur395 fl (A, C, F, K, US);
org9 fr (F, NY, US). PUERTO RICO. Berterosn fl (G-DC); Brittonet al 1679 fl fr (F, NY, US); 2293
fl fr (C, F, GH, NY, US); 6277 fl fr (NY); 7463 fl (NY); 9944 fl fr (NY); Eggerssn fl (C); Garber56 fl
(GH, K, NY); Gregory12 fl (NY); Hess 5614 fl fr (NY); Hioramsn fl (NY); Howardet al 15595 fl (GH);
Plee sn fl (K, P); 210 fl (P); 805 fl (P); Sintenis965 fr (BM, GH, GOET, LD, LE, S, US); 965b fl (US);
i668 fr (F, K, LD, US); 2145 fl (K); Wagner549 fl (BM, LE, S); 580 fl (BM, LE). LEEWARD
ISLANDS. Saba: Boldingh2277 fl (NY); Stoffers4169 fl (GH, U). St. Kitts: Beard292 fl (A, K, NY);
Britton & Cowell I87 fl fr (K, NY, US); Masson sn fl (BM, S); Proctor I9645 fl fr (A). Nevis: Proctor
I9419 fl (A); A. C. Smith 10507 fl (A, NY, US). Montserrat: Howard Ir886 fr (A); Miller 9 fl (US);
Proctor18921 fl (A); Ryan sn fl fr (C); Shafer525 fl (F, NY, US). Guadeloupe: Duchassaingsn fl (A,
GOET, NY); Duss 363 fl (A, P); 2214 fl fr (C, F, GH, NY, US); Quentinsn fl (A); Questel2027 fl (P,
US); 2302 fl (P, US); 2558 fl (P, US); 4089 fl (P, US); 5114 fl (US); Rodriguez4721 fl fr (A); Stehle
i8i3fl (US); 2524 fl (US); 2763 fl (US). Marie Galante: Proctor20997 fl (A). WINDWARD ISLANDS.
Dominica: Beard 657 fl (A, K, NY); Eggers 639 fl (BR, GH, GOET, LE); Hodge579 fl fr (BM, GH,
NY, US); 2000 fr (GH, US); 2569 fr (GH); 3074 fl (GH, US); Howard 11769 fl (A); Imray301 fr
(GOET); 319 fl (GOET); sn fl fr (K); Lloyd338 fl (NY). Martinique: Duss I905 fl fr (F, NY, US);
Hahn 213 fl (BM, GH, US); 973 fl (BM, W); 1247 fl (BM, BR, F, K, W); Isertsn fl fr (C); Stehle5573
fl fr (US). St. Vincent: H. H. & G. W. Smith 748 fl (BM, GH, NY). COLOMBIA. Goajira: Haught
4264 fl (COL, F, K, NY, US). Magdalena: Angel 679 fl (US); Haught 3817 fl (COL, F, NY, US);
4024 fl (COL, F, K, NY, US); RomeroC. 622 fl (COL, F, GH, IAN, NY, US); 787 fl (COL, US);
H. H. Smith 442 fl (A, BM, BR, COL, F, GH, K, MICH, NY, P, US). Meta: Philipson & Idrobo1535
fl (BM, COL, NY); I797 fr (BM, COL); 2085 fl (BM, COL, NY). Without precise locality: Mutis
155 fl (US); 1375 fl (US). VENEZUELA. Zulia: Lasser2580 fl (VEN); Steyermark& Ferndndez99662
fl (NY, VEN); Tejera82 fl (GH, NY, US); 169 fl (GH, US). Lara: Aristeguieta4935 fr (VEN); Saersn
fl (W). Yaracuy: Aristeguieta& Pannier 1898 fl fr (NY, VEN); Brito 37 fl (VEN). Portuguesa: Brito
37(11) fl (VEN); Ll. Williams 12319 fl (F, IAN, K, S, US, VEN). Carabobo: Funck & Schlim556 fl
(BR, LD, US); Pittier 7623 fl fr (GH, VEN). Aragua: Agostini 215 fl (VEN); Alston 5820 fl (BM);
Aristeguieta& Saldina562 fl (VEN); Delgado 107 fl (A, F, US, VEN); 126 fl (US); Lasser2238 fl (VEN);
Matos 1o91 fl (VEN); Pittier 6072 fl (G, NY, US, VEN); 15205 fl fr (US, VEN); 15778 fl fr (VEN);
Pittier & Nakichenovitch 15388 fl fr (US, VEN); Vogel49 fl (IAN, M, NY, US, VEN); Ll. Williams9995
fl (F, VEN); 10753 fl (A, F, VEN); Ll. Williams & Alston I50 fl (US). Dist Federal: Fendler38 fl (BR,
GH, GOET, K, NY, US); Pittier 11087 fl (US, VEN). Miranda: Pittier 7827 fl (GH, US, VEN).
Anzoategui: Steyermark 6o058 fl (F, NY, US, VEN). Sucre: Steyermark 662833 fr (F, NY, VEN). Mona-
gas: Lasser& Vareschi4048 fl fr (VEN); Steyermark 62183 fl fr (F, NY, VEN). Barinas: Aristeguieta1697 fl
(NY, US, VEN); Bernardi1723 fl (FI, K, NY, P, VEN); 1762 fl (NY); Breteler3449 fl (NY, RB, SP,
US, VEN); 4592 fl (SP); Veillon i/V fl (K, VEN). Bolivar: Steyermark89116 fl (NY). TOBAGO.
Sandwith1737 fl (K, NY). TRINIDAD. Bot. Garden2062 fr (NY, US); Britton & Broadway2814 fl (GH,
NY); 10323 fl (K); Broadway6469 fl fr (F, K, S, US); Don 53 fl (LD); 8977 fl (BM, K); Crueger301 fl
(GOET); Fendler1021 fl (NY, P); Lockhart162 fl (K); 220 fl (K); Prance2104 fl fr (NY); Russell 12240
fl (K); Russell & Swabey12470 fr (K); R. O. Williams 12605 fl (K). GUYANA. Jenman7000 fl (K, NY);
A. C. Smith3431 fl (A, F, IAN, K, LE, NY, P, S, US); 3590 fl (A, F, IAN, K, LE, NY, P, US); Wilson-
Browne174(F. D. 5667) fl (K, NY); 363(F.D. 5777) fl (K, NY). SURINAME. Irwin, Prance,Soderstrom,
& Holmgren54904 fl (F, GH, NY, US); Maguire, Schulz, Soderstrom& Holmgren54381 fl (F, GH, NY,
US); Pulle 382 fl (NY, U). ECUADOR. Little 6668 fl (US). PERU. Loreto: Killip & Smith 29271 fl
fr (F, NY, US); Klug 2250 fl (A, BM, F, GH, K, NY, US); Tessmann3831 fl (NY); LI. Williams 4653
304 Flora Neotropica

fl (F). San Martin: Ferreyra4582 fl (US); Klug 3724 fl (A, BM, GH, K, NY, US). Madre de Dios:
Ule 9413 fl (K, MG). BRAZIL. Amazonas: Krukoff4655 fl (A, BM, F, K, LE, MICH, NY, S, US);
4760 fl (A, BM, K, LE, MICH, NY, S, US); 4852 fl (A, BM, F, K, LE, MICH, NY, S). Terr. Roraima:
Pranceet al 4562 fl (IAN, INPA, NY). Para: DuckeMG 15183 fl (MG); MG 17124 fl (MG); RB 18234
fl (RB, S); Frdes23513 fl (IAN, NY); 30200 fl (IAN); 30530 fl (IAN); Silva 178 fl (IAN, NY). Acre:
Ducke RB 25011 fl (INPA, RB, US); Krukof 5246 fl (A, BM, F, FI, K, MICH, NY, S, US); 5321 fl
(A, BM, F, K, LE, MICH, NY, S); 5428 fl (A, BM, F, K, LE, MICH, NY, US); Pranceet al 7295 fl
(INPA, NY). Terr. Rond6nia: Pranceet al 5671 fl (INPA, NY). Mato Grosso: Kuhlmann1988 fl (R,
RB). Bahia: Belem & Magalhaes633 fl fr (IAN, NY, UB). BOLIVIA. Krukoffo1638 fl fr (A, F, K, MICH,
NY, S, US); 10658 fr (A, F, K, MICH, NY, S); Pearcesn fl (K); Pranceet al 6255 fr (INPA, NY);
Rusby565 fr (NY); 590 fr (NY); 1448 fl (F, LE, MICH, NY, US); 600ofl (F, K, GH, NY, US); 2492
fl (NY); Steinbach2183 fl (GH, NY); 6449 fl (F, GH, K, S).

LOCALNAMES. Panama: Camaroncillo, Carapato,Chicharrdn,Conejo.Cuba: Icacillo,

Siguapa.St. Kitts: Pigeonberry.Guadeloupe: Icaquea poils, Icaquepoileux.Dominica:
Bois poil. Colombia: Carita negro. Venezuela: Caramacatenegro,Carbonera,Hierrito,
Icaquillo,Fruta de paloma, Palo blanco.Guyana: Bokotokon,Waroma.Peru: Isiguiro-ey
(Huitoto).
The Stockholm herbarium contains two sheets of Hirtella triandralabelled
"Swartz Herbarium", but both these sheets are without data. Hirtella triandrawas
obviously based on this material which must therefore be regarded as the type. In
addition, Swartz must have seen other material such as a sheet from the Banks
herbarium at the British Museum. This sheet contains three separate collections all
from the Caribbean: De Ponthieusn, Fr. Mapa sn; J. Ryansn. Perhaps the material in
the Swartz herbarium originated from one or more of these collections. Stockholm
has the De Ponthieu material, but it is not marked as belonging to the Swartz her-
barium. Swartz' description is not followed by the asterisk used to indicate material
from the Banks herbarium, as has been mentioned under H. paniculata.For these
reasons the material in the Swartz herbarium is considered the type. Both the Swartz
herbarium and the Banks herbarium contain material of the minor variant which
Urban described as H. jamaicensis.They also contain material of what has generally
been interpreted as typical H. triandra.Since H. jamaicensisis not distinct from H.
triandra,this does not affect the nomenclature.
I have not seen the holotype of Hirtellacosmibuena (which is also the holotype of
H. racemosaRuiz & Pavon and H. peruviana)at Madrid. However, there is a photo-
graph of it in the herbarium at Washington (US) and a duplicate isotype at Chicago.
These match well a sheet marked "Pavon" at the British Museum which is pre-
sumably another isotype.
Hirtellacastaneais one of the plants described by De Candolle from the Mocifio
& Sesse drawings, see p253. There is a specimen in the Mocifio & Sesse herbarium
which matches both the drawing and the description of H. castaneafairly well. This
specimen is certainly H. triandra,and as there are so few species of Hirtellain Mexico
it seems certain that H. castaneais equal to H. triandra,the only triandrous species of
Hirtellaknown in that country. The specimen of H. triandramarked "Pavon" at the
British Museum is almost certainly a duplicate of the Mocifio & Sesse collection.
Pavon never visited Mexico, but distributed Mocinio & Sesse Mexican plants under
his name. Further details about the difficulties of Mocifio & Sesse collections have
been discussed elsewhere (Prance, 1968).

FIG 47. Leaf shape variation and geographical distribution of Hirtella triandra.? H. triandra
subsp triandra; ? H. triandrasubsp media; A H. triandrasubsp punctulata.A, LI. Williams 9480; B,
Proctoret al 27119; C, Harris 10753; D, Duss 1905; E, Aristeguieta1898; F, Krukoff10638; G, Glaziou8666.
all leaves x 0.25.
 Hirtella 305

-i~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~2

?
o .],-.., .',
)/I ~ ~~~~~~~~~~~~~~~~~~~~~ I ' ,o '~
~

?'
-

_. I
?~"'"~II

I~~~~,?
,_,. /
. _ ~. ?

,0 70 0'
F
">"I~
i~~~~~~~~~~~~....
'

!,..i,,,/ G~`

~..........~
.~,:w~
:''?? ~~ i'
306 Flora Neotropica

Hirtellabracteatawas distinguished from H. triandraby the numerous bracts, and

the broad leaves. Both characters merge into those of H. triandraand occur sporadi-
cally throughout the geographic range. Thus, H. bracteatacannot be regarded as a
separate species. Similarly, it is not possible to keep H. jamaicensisand H. multifJora
apart from H. triandra.Specimens bearing these names were studied with the use of
scatter diagrams which showed no gaps in the variation. Although at the extremes
these species look distinct from typical H. triandra,their variation merges completely
with that of H. triandra,and there is no correlation between the different characters.
For example, the fruit of H. jamaicensisappears rather distinct by virtue of its tomen-
tose exterior and blunt apex. Identical fruit occurs in several collections from Panama
(eg Pittier5523, Williams667) which in all other respects is typical H. triandra.Hirtella
jamaicensisand H. multifJora both have distinctly smaller and thicker leaves, a character
which occurs in Panamanian and Colombian material where there is complete
gradation into that of typical H. triandra.H. jamaicensisis also rather different in the
largely axillary inflorescences. This type of inflorescence also occurs in the Panama-
nian material where there is complete gradation again into that of typical H. triandra.

36b. Hirtella triandra subsp punctulata (Miquel) Prance, stat nov

HirtellapunctulataMiquel, Linnaea 19: 439. I847.
Hirtella triandrasensu Hooker f., Mart. Fl. Bras. I4(2): 37. I867.
Hirtella hevandaMartius ex Colla, Herb. Pedem. 2: 289. I834. Type. Martiussn, Brazil, fl (holo-
type, TO; isotype, BM).
Hirtella ellipticaPresl, Symb. Bot. 2: 23. 1858. Type. Herb. Presl. sn, Brazil, Guanabara, fl (holo-
type, PRC).

Leaves 3.5-7.0 cm long, thick-coriaceous, with a short abrupt acumen I.0-4.0

mm long; primary veins 7-9 pairs; inflorescence with a central rachis bearing small
lateral branches.
TYPE. Claussen 205612, Brazil, Guanabara, fl (G).
DISTRIBUTION. Wooded slopes and coastal forest in eastern Brazil. Flowering
July to January.
BRAZIL. Bahia: Duarte 644 fl (RB I 13215). Minas Gerais: Langsdorfsn fl (K, OXF); withoutcol.
sn fl (C). Espirito Santo: Freire73 fl (R); Kuhlmann381 fl (RB 136920); WIied-Neuwied sn fl (BR). Rio
de Janeiro and Guanabara: Bowie & Cunningham 572 fl (BM); CapanemaRB 19199 fl (RB); RB 19207
fl (RB); Gardner864 fl (BM, GH, K); Gaudichaud1020 fl (F, P); o023 fl (GH, P); Glaziou 6199 fl (C,
K, P); 8666 fl (BR, C, K, LE, P); Gomessn fl (NY); Hans sn fl (RB 35576); Luid 371 fl (C); Luschnathsn
fl (BR, OXF); Lutz 1230 fl (NY, R); Miers sn fl (BM, NY, P); 3776 fl (K); Nadeaudsn fl (NY, P);
Netto, Gla;.iou& Schwackesn fl (R 7171); Ottosn fl (C); Passarellisn fl (R 35575); Pulle & Lutz 1130 fl
(BM, R); Riedel sn fl (NY, OXF, R, US); 566 fl (US); 567 fl (NY, US); Schenck2303 fl (C); Schott
4969 fl (NY); Sellow sn fl (LE); St. Hilaire i fl (P); 7 fl (P); 542 fl (P); Tweediesn fl (OXF); 1268 fl
(K); Ule 4484 fl (NY, R); Vauthiersn fl (GH, OXF); 508 fl (F, GH, P); lVidgrensn fl (S); 704 fl (S)
'ied-Neuwiedsn fl (LD). Without locality: Raddisn fl (FI). Since so many of the above collections lack
the precise locality, I have not attempted to separate those from the two states of Rio de Janeiro and
Guanabara.

Hirtellapunctulatacannot be regarded as a species separate from H. triandraas

was recognized by Hooker (i 867) since he reduced the former species into synonymy.
However, although H. punctulata merges into H. triandra, the majority of specimens
are quite distinct in leaf shape and texture, and it is also geographically apart from
the rest of H. triandra. (see Fig 47). For these reasons, I recognize this taxon as a
geographical subspecies of H. triandra.
12
Cited as 2056 in the protologue, but marked 2o5b in the Geneva herbarium.
Hirtella 307

36c. Hirtella triandra subsp media (Standley) Prance, stat nov

HirtellamediaStandley,Publ. Field Mus. Bot. 23: 9. 1943.

Leaves 3.5-8.0 cm long, subcoriaceous, acute at apex, primary veins 5-7 pairs;
inflorescence small, compact, and much branched.
TYPES.C. & W. von Hagen 1378, Honduras, fl (holotype, F; isotype, NY);
Liebmanni75i, Mexico, Oaxaca, fl (paratypes, C, F); LI. Williams9480, Mexico,
Oaxaca, fl (paratypes, A, BM, K, M, MAD, MICH, NY, P, S, US).
Primary forest from southern Mexico to Nicaragua.
DISTRIBUTION.
Jones& Facey27119,fl (NY).
NICARAGUA:Proctor,

LOCAL NAMES. Mexico: Jicaco; Nicaragua: Barazon.

Hirtella media merges with H. triandra,but at the extremes is easily recog-
nized. Its geographical range overlaps with that of H. triandra,which suggests sub-
specific status. Subspecies mediais a large forest tree, whereas subspecies triandrais
most frequently a small tree.

37. Hirtella bahiensis Prance, sp nov Fig 39.

Arbor media, ramulis juvenilibus puberulis, mox glabris lenticellatis. Folia

alternata petiolata; petiolo 1.0-2.0 mm longo, tereti, eglanduloso, primo tomentello
mox glabro; laminae oblongae vel oblongo-ellipticae, 1.8-4.5 cm longae, 1.2-1.9 cm
latae, apice rotundatae vel acutae, basi cuneatae, supra glabrae, subtus glabrae vel
cum pilis longis appressis paucis instructae, pauciglandulosae; costa media supra
prominente, subtus prominente, glabrescente; costis secundariis 4-7 jugis, supra
planis, subtus leviter prominentibus. Stipulae circa .o mm longae, puberulae,
persistentes, eglandulosae. Flores in paniculis terminalibus axillaribusque 5.0-
Io.o cm longis dispositi, ramis brevibus; rachi ramisque sparse tomentellis. Bracteae
bracteolaeque 2.5-5.0 mm longae, ovatae, acutae, persistentes, tomentellae, cum
glandulis stipitatis munitae. Receptaculum campanulatum, extus tomentellum,
intus basim versus glabrum; pedicelli 0.2-1.5 mm longi. Calycis-lobi acuti, utrinque
puberuli, eglandulosi. Petala 5, glabra. Stamina 3, unilateralia, 4-5 staminodiis
opposita; filamenta glabra, longe exserta. Ovarium ad faucem receptaculi insertum,
pilosum. Stylus e basi ovarii ortus, hirsutus. Drupa immatura ellipsoidea; epicarpio
puberulo mox glabro; mesocarpio tenue, duro, osseo, longitudinaliter canaliculato,
intus hirsuto.
TYPE.Belem & Pinheiro2756, Brazil, Bahia, Santa Cruz Cabralia, fl (holotype,
NY; isotype, UB).
DISTRIBUTION. Known only from coastal forest in the State of Bahia, Brazil.
Flowering October to February.
BRAZIL. Bahia: Maraf, Belem& Pinheiro
3094 fr (NY, UB); Porto Seguro,Duarte8029 fl fr
(NY, RB).

This species is quite distinct from any other species of Hirtella,but is probably
closest to H. triandrasubsp punctulata.It differs from that subspecies in the glandular
bracts and bracteoles, the smaller leaves with rounder apices and fewer primary veins,
and the smooth not papillose undersurface of the leaves.
308 Flora Neotropica

38. Hirtella latifolia Prance, sp nov Fig 45 E-H.

Arbor media, ramulis juvenilibus tomentellis mox glabris. Folia alternata,

petiolata; petiolo 2.0 mm longo, tereti, eglanduloso, tomentello; laminae ovato-
ellipticae, coriaceae, I5.o-I7.0 cm longae, 8.o-Io.o cm latae, apice in acuminem
3.0-6.0 cm longum contractae, basi rotundatae, subtus costis venisque hirsutae;
costa media supra leviter prominente, tomentella, subtus prominente, hirsuta; costis
secundariis I2-13 jugis, supra plana, subtus prominentibus. Stipulae lanceolatae,
eglandulosae, persistentes, puberulae. Flores circa 6.0 mm longi, in paniculis ter-
minalibus 6.0-I2.0 cm longis dispositi, rachi ramisque tomentelli, rachi crassi.
Bracteae bracteolaeque ovatae, numerosae, persistentes, tomentellae. Receptaculum
campanulatum, extus tomentellum, intus basim versus glabrum; pedicelli 0-0.5 mm
longi, Calycis-lobi acuti vel rotundati, utrinque tomentelli. Petala 5, glabra. Stamina
3, unilateralia, filamenta glabra, longe exserta. Ovarium ad faucem receptaculi
insertum, pilosum. Stylus e basi ovarii ortus, basim versus hirsutus. Fructus mihi
ignoti.
TYPE. Duke 8012, Panama, Prov. Panama, Cerro Jefe, fl (holotype, MO;
isotype, MICH).
DISTRIBUTION. Collected only in forest at the type locality.
PANAMA. Prov. Panama: Cerro Jefe, Dwyer, Gauger& Baker 7259 fl (MO).

This species is most closely related to H. triandra,but differs in the much broader
leaves, the thicker rachis of the inflorescence, and the subsessile flowers.

39. Hirtella suffulta Prance, sp nov Fig 46.

Arbor media vel magna, ramulisjuvenilibus sparse puberulis mox glabris. Folia
alternata petiolata; petiolo 6.o-Io.o mm longo, tereti, eglanduloso, sparse appresso-
pubescenti; laminae oblongae, subcoriaceae, 5.5-1 I.5 cm longae, 2.0-5.0 cm latae,
apice in acuminem 2.0-6.0 mm longum abrupte contractae, basi subcuneatae, subtus
fere glabrae et cum pilis longis paucis, eglandulosae; costa media supra leviter
prominente, subtus prominente, cum pilis appressis munita; costis secundariis 8-9
jugis, supra planis, subtus prominentibus. Stipulae 2.0-3.0 mm longae, deltatae,
persistentes, eglandulosae. Flores 5.0-6.0 mm longi, in paniculis terminalibus sub-
terminalibusque dispositi, rachi 5.0-I5.0 cm longi, ramis brevissimisdenseflorentibus,
rachi ramisque sparse puberulis. Bracteae bracteolaeque 3.0-6.0 mm longae, per-
sistentes, alabastris includentes, membranaceae, sparse pubescentes, eglandulosae
vel interdum cum apice glandulosae. Receptaculum campanulatum, extus puberulum
intus basim versus glabrum; pedicelli circa 0.5 mm longi. Calycis-lobi acuti, extus
sparse puberuli, intus tomentelli, marginibus eglandulosis. Petala 5, glabra. Stamina
5-6, unilateralia, filamenta glabra, longe exserta. Ovarium ad faucem receptaculi
insertum, pilosum. Stylus e basi ovarii ortus, hirsutus. Drupa immatura ellipsoidea;
epicarpio glabra, laeve; mesocarpio tenue, carnoso; endocarpio tenue, duro, osseo,
intus hirsuto.
TYPE. Pires 6943, Brazil, Para. Pirelli Forest near Belem, fl fr (holotype, IAN;
isotypes, NY, UB).
DISTRIBUTION. Collected in non-flooded forest in the States of Amazonas and
Para, Brazil, flowering July.
BRAZIL. Amazonas: Rodrigues7705 fl (INPA I7010). Para: Pirelli Forest, Belem, Pires 6813 fl
(IAN, NY, UB); 6885 fl (IAN, UB); 7004 fl (IAN, NY, UB); Belem-Brasilia road, Km Ioo, N.T.
Silva 613 fl (IAN).
Hirtella 309

This species is distinct from any other by the large persistent bracteoles. It is
closest to H. eriandra,but, in addition to the bracteoles, it differs in the more densely
crowded inflorescence, the less pubescent leaves and inflorescence, and the fruit
with a glabrous epicarp and bony endocarp.

40. Hirtella elongata Martius & Zuccarini, Abh. Akad. Miinchen I: 384.
1832; Zucc., Flora 15(Beibl. 2): 85. I832; Hook. f., Mart. Fl. Bras.
14(2): 24, t. I (2). 1867; Macbride, Fl. Peru 1076. I938.

Tree to 12.0 m tall or shrub, the young branches puberulous, becoming glabrous
and lenticellate with age. Leaves elliptic to oblong-elliptic, subcoriaceous, 8.5-23.0
cm long, 4.5-8.0 cm broad, abruptly acuminate at apex, with acumen 3.0-I5.0 mm
long, usually subcordate at base, less frequently rounded, hirsute beneath along
principal venation only, with a few palisade glands towards base; primary veins
10-13 pairs, plane above, prominent beneath; midrib plane or nearly so and pubes-
cent above, prominent and hirsute beneath; petioles 3.0-4.0 mm long, terete, eglan-
dular, tomentellous. Stipules linear-lanceolate, to 7.0 mm long, caducous, eglandular.
Inflorescences terminal panicles, 10.0-30.0 cm long, with a long central rachis and
many short few-flowered lateral branches, the rachis and branches short-puberulous
to glabrescent. Bracts and bracteoles 2.0-5.0 mm long, often numerous on lateral
branches, lanceolate to deltoid, persistent, eglandular, puberulous. Flowers 5.o-6.o
mm long. Receptacle campanulate, puberulous on exterior, glabrous within except at
throat. Calyx lobes acute, puberulous on both surfaces. Petals 5, white, glabrous.
Stamens 6-7, unilateral, the filaments far exceeding calyx lobes, glabrous. Style
glabrous except at base. Ovary inserted at mouth of receptacle, pilose-tomentose.
Fruit oblong; epicarp smooth, glabrous; mesocarp thin, fleshy; endocarp thin, hard,
bony, hirsutulous within.
TYPE.Martiussn Brazil, Amazonas, fl (holotype, M; isotypes, BR, K, M).
DISTRIBUTION. River banks and riverine forest in western Amazonia. Flowering
throughout the year.
COLOMBIA. Boyac,: Haught 2604 fl (A, COL, F, IAN, S, US). Meta: Cuatrecasas7817 fl
(COL, US); Fosberg20193 fl (NY, US); Jaramilloet al 1ooI fl (COL); 1174 fl (COL); 121I fl (COL);
Karstensn fl (LE); Killip 34298 fl (COL, US); Pinto et al 262 fl (P); Schwabesn (COL). Vaupes: Cuatre-
casas 6915 fl (COL, F, US); Ferndndez2066 fl (COL, IAN, US); 2095 fl (COL, US); Gutierrez&
Schultes510 fl (COL); Schultes& Cabrera12558 fl (COL, GH, NY, US); I3807 fl (BM, COL, GH, NY,
US); 16612 fl (F, GH, NY, US); 18405 fl (COL, GH); 19514 fl (GH, US); Uribe 2916 fl (COL).
Amazonas: Schultes3854 fl fr (COL, ECON, GH, K, US); Schultes& Cabrera16459 fl (COL, GH, NY,
US). VENEZUELA. Guarico: Croizat203 fl (F). Bolivar: Wurdack& Monachino40935 fl (F, GH, NY,
US). Terr. Amazonas: Maguire,Cowan & Wurdack30757 fl fr (F, GH, NY, US); Maguire & Wurdack
34845 fl fr (NY); Maguire, Wurdack& Maguire42641 fi (NY); Spruce3196 fl (BM, BR, GH, K, NY, P);
Wurdack& Adderley430oo fl (NY); LI. WilliamsI4464 fl (F, VEN). PERU. Loreto: Klug 726 fl (F, NY,
US); 916fl fr (F, NY, US); 2530 fl (F, NY, S, US); 2575 fr (F, US); Schunke23 fl fr (A, F, NY, US);
Tessmann5297 fl (NY). BRAZIL. Amazonas: Ducke 1878 fl (A, F, IAN, K, MG, NY, R, RB, US);
MG 6785 fl (BM, MG, RB); MG 7365 fl (BM, MG, RB); RB 19763 fl (RB); Elias 314 fl (MG); Frdes
20892 fl (IAN, NY, US); 34788 fl (IAN); Krukof 8ioo fl (A, BM, BR, F, FI, K, LE, NY, S); 8966 fl
(A, BR, F, K, LE, NY, P, S); Lima 48-o03 fl (IPA); Luetzelburg22502 fl (R); 22758 fl (R); Pires 1351
fl (IAN); Poeppig2502 (4502) fl fr (F, GOET, LD, NY, P, W); RomeroC. 3575 fl (COL); Spruce2520
fl (BM, BR, C, F, GH, GOET, K, LD, LE, NY, P); Ule 5654 fl (K, MG). Acre: Pranceet al 2999 fl
(INPA, NY).

LOCALNAME.Colombia: Eenawdla (Yukuna), Garrapato

Closely related to H. eriandra,but for differencessee under that species.
310 Flora Neotropica

41. Hirtella rodriguesii Prance, sp nov Fig 48 A-C.

Arbor, ramulisjuvenilibus hispidis. Folia alternata, petiolata; petiolo 1.0-3.0 mm

longo, tereti, eglanduloso, juvenili hirsuto; mox glabro; laminae oblongae, vel ovato-
ellipticae membranaceae, 3.5-7.5 cm longae, 1.5-3.8 cm latae, apice in acuminem
3.0-9.0 mm longum contractae, basi rotundatae vel subcuneatae, subtus glandulosae,
praeter costam glabrae; costa media supra leviter prominente, subtus prominente
sparse hirsuta; costis secundariis 6-9 jugis, supra planis, subtus leviter prominentibus.
Stipulae lineares, circa 2.0 mm longae, hispidae, persistentes. Flores 4.0-5.0 mm
longi, in paniculis vel racemis axillaribus terminalibusque 2.5-7.0 cm longis dis-
positi, rachi ramisque sparse hispidis. Bracteae bracteolaeque 1.5-2.5 mm longae,
lanceolatae, puberulae, eglandulosae, vel cum glandulis sessilibus paucis munitae,
persistentes. Receptaculum campanulatum, extus hispidum, intus basim versus
glabrum; pedicelli 1.5-3.0 mm longi. Calycis-lobi acuti, extus sparse hispidi, intus
sparse puberuli. Petala 5, alba, glabra. Stamina 4-5, unilateralia, filamenta glabra,
basim versus connata, longe exserta. Ovarium ad medium receptaculi insertum,
pilosum. Stylus e basi ovarii ortus, hirsutus. Fructus mihi ignoti.
TYPE. W. Rodrigues& Osmarino6735, Brazil, Amazonas, Manaus, Reserva
Florestal Ducke, fl (holotype, NY; isotype, INPA 5I85).
DISTRIBUTION. Non-flooded forest in central Amazonian Brazil. Flowering June
to September.
BRAZIL. Amazonas: Manaus,Colonia Joao Alfredo, Ducke 976 fl (IAN, K, MG, NY, R, RB,
US); Reserva Florestal Ducke, CoelhoINPA 5984 st (INPA); INPA 5985 st (INPA); Rio Embira, basin
of Rio Jurua, Krukof 4962 fl (A, F, LE, MICH, NY, RB, S, US).

This well isolated species is most closely related to Hirtella glandistipula,but

differs in the bracts which are either eglandular or have sessile glands only, the
eglandular stipules, and the tendency toward a branched inflorescence. It differs
from H. bicornisin the hispid inflorescence and young branches, and in the thinner
leaves with acuminate (not cuspidate) apices. H. rodriguesiidiffers from H. triandra
in the greater number of stamens, in the little branched inflorescence, in the hispid
branches, and in the membraneous leaves.

42. Hirtella obidensis Ducke, Arch. Jard. Bot. Rio de Janeiro 3: 267. 1922;
Kleinh. in Pulle, Fl. Suriname 2(I): 448. I939.
Hirtella lanceolataKleinhoonte, Rec. Trav. Bot. N6erl. 22: 388. 1925; in Pulle, Fl. Suriname
2(I): 448. 1939, synon nov. Type. B.W. 5338, Suriname, fl (holotype, U).
Hirtella coppenamensis Kleinhoonte, Rec. Trav. Bot. N6erl. 30: i8o. I933. Type. B.W. 6167,
Suriname, fl (holotype, U; isotypes, A, C, IAN, K, NY).

Tree to 25.0 m tall, the young branches tomentose, becoming glabrous with
age. Leaves oblong-lanceolate to elliptic, thick-coriaceous, 5.0-I3.0 cm long,
2.5-5.0 cm broad, acuminate at apex, with acumen 2.0-9.0 mm long, rounded to
subcuneate at base, hirsutulous on venation beneath; venation of upper surface
impressed; primary veins 9-I2 pairs, impressed above, prominent beneath; midrib
plane or nearly so above, prominent beneath; tomentose on both surfaces; petioles
I.0-3.0 mm long, tomentose, terete, eglandular. Stipules ca 2.0 mm long, subulate,
tomentose, eglandular, persistent. Inflorescences short, terminal and subterminal,
little-branched panicles 4.0-I I.o cm long, the rachis and branches rufous-tomentose.
Bracts and bracteoles 1.5-2.5 mm long, oblong to ovate, persistent, tomentose,
with few sessile glands which are usually obscured by the pubescence. Receptacle
campanulate, tomentose on exterior, glabrous within except at throat; pedicels
Hirtella 311

A-:x

IN

D E
J.-Lokmv

FIG 48. Species of Hirtella. A-C, H. rodriguesii(Rodrigues5512); A, habit, x 0.5; B, flower, x 3;

C, flower section, x 3. D-G. H. adderleyi(Wurdacket al 42973, 437i6); D, habit, x 0.5; E, bract, X 7.5;
F, flower section, x 3; G, flower, x 3.
312 Flora Neotropica

0.5-2.0 mm long. Calyx lobes acute, rufous-tomentose on exterior, gray-pubescent

within. Petals 5, glabrous. Stamens 5-7, unilateral with short filamentous staminodes
opposite them, the filaments glabrous, far exceeding calyx lobes. Style hirsute up to
half its length. Ovary inserted near mouth of receptacle, pilose. Fruit oblong-rotund;
epicarp tomentellous when young becoming smooth and glabrous with age; mesocarp
thin, fleshy; endocarp very thin, hard, bony, tomentellous, within.
TYPE. Ducke RB I7113, Brazil, Para, fl (holotype, RB).
DISTRIBUTION.Non-flooded forest in the Guianas and eastern Amazonian Brazil.
GUYANA.Guppy 322 st (NY). SURINAME.Lindeman 5014 St(NY, U); Schulz7147st (NY, U);
8013 st (NY, U); Van Donselaar1870 st (U); 2930 st (U). BRAZIL. ParA: DuckeRB 19764 fl (INPA,
K, P, RB, S, US); Oliveira809 fl (IAN, UB); 4702 fl (IAN); Prance& Silva 58964 fr (NY, UB); Silva
90o fl (IAN, NY); o036 fl (IAN, NY).

This species is closest to H. hebeclada,

but differs in the branched inflorescence.

43. Hirtella cowanii Prance & Maguire, sp nov Fig 49 D-F.

Frutex, ramulis juvenilibus tomentellis mox glabris lenticellatis. Folia alternata

petiolata; petiolo 0.5-1.5 mm longo, tomentoso, tereti, eglanduloso; laminae ovatae
vel oblongo-ovatae, coriaceae, 2.0-6.3 cm longae, 1.5-3.2 cm latae, apice in acumi-
nem I.o-5.o mm longum contractae, basi rotundatae, subtus dense appresso-pubes-
centes, supra praeter costam mediam glabrae, reticulatae; costa media supra leviter
prominente subtus prominente, utrinque tomentella; costis secundariis 6-9 jugis,
utrinque leviter prominentibus, ad marginem anastomosantibus. Stipulae circa 1.5
mm longae, subulatae, tomentosae, eglandulosae, subpersistentes.Flores 4.0-5.0 mm
longi in paniculis brevissimis fasciculatis 0.8-I.5 cm longis dispositi, rachi ramisque
ferrugineo-tomentosis.Bracteae bracteolaeque circa 2.0 mm longae, oblongae, acutae,
tomentosae, eglandulosae, persistentes. Receptaculum campanulatum, subsessile,
extus tomentosum, intus basim versus glabrum. Calycis-lobi acuti, utrinque tomen-
telli, eglandulosi. Petala 5, alba, glabra. Stamina 7, unilateralia, staminodia fila-
mentosa aliquando opposita; filamenta glabra calycis-lobos paullo excedentes.
Ovarium ad faucem receptaculi insertum, piloso-tomentosum. Stylus e basi ovarii
ortus, basim versus hirsutus. Drupa (immatura tantum visi) rotundato-oblonga;
epicarpio tomentello; mesocarpio tenue, carnoso; endocarpio tenue, duro, osseo,
intus hirsuto.
TYPE. Cowan & Wurdack31237, Venezuela, Terr. Amazonas, Serrania Paru,
Rio Ventuari, fl (holotype, NY).
DISTRIBUTION. One of the few species of the family found at higher altitudes
(I600-2000 m alt.), growing in open rocky places, in the Rio Ventuari region of
Venezuela. Flowering February to April.
VENEZUELA.Terr. Amazonas:SerraniaParu; Rio Ventuari,Cowan& Wurdack
31o84 fl bud
(NY); 31149 fl (NY); 31383 fl (NY); Phelps& Hitchcock
514 fl (NY).

The small compact inflorescence of H. cowaniidistinguishes it from most other

species of Hirtella. It is closest to H. orbicularis,but for distinguishing characteristics
see under that species.

44. Hirtella orbicularis Prance, sp nov Fig 49 A-C.

Frutex vel arbor parva, ramulis juvenilibus tomentellis mox glabris. Folia
Hirtella 313

alternata petiolata; petiolo 1.0-2.0 mm longo, tomentello, tereti, eglanduloso;

laminae orbiculares vel ovatae, coriaceae, 2.5-5.8 cm longae, 1.6-4.6 cm latae, apice
breviter acuminatae vel mucronatae vel retusae, basi subcordatae vel raro rotun-
datae, subtus appresso-pubescentes, glandulosae; costa media supra leviter promi-
nente, subtus prominente, utrinque hirsuta; costis secundariis 7-9 jugis, utrinque
leviter prominentibus. Stipulae parvae, lanceolatae, ad 2.0 mm longae, eglandulosae.
Flores 5.0-6.0 mm longi, in paniculis 2.0-4.0 cm longis dispositi, rachi ramisque
rufo-tomentosis. Bracteae bracteolaeque 1.0-3.0 mm longae, ovatae, acutae, per-
sistentes, tomentellae, eglandulosae. Receptaculum campanulatum, extus tomentel-
lum, intus basim versus tantum glabrum supra pubescens; pedicelli 0-0.5 mm longi.
Calycis-lobi acuti, extus tomentelli, intus puberuli. Petala 5, alba, glabra. Stamina 5,
unilateralia, staminodiis dentatis opposita; filamenta glabra, longe exserta. Ovarium
ad faucem receptaculi insertum, pilosum. Stylus e basi ovarii ortus, hirsutus. Drupa
oblonga; epicarpio juvenili sparse tomentello, mox glabro; mesocarpio tenue, car-
noso; endocarpio tenue, duro, osseo, intus hirsuto.
TYPE. Maguire, Cowan & Wurdack29622, Venezuela, Terr. Amazonas, Cerro
Duida, Rio Cunucunuma, fl (holotype, NY).
DISTRIBUTION.Slopes and open places, 1500-3000 m altitude on Cerro Duida,
Venezuela. Like the preceding species, this is another high altitude endemic.
Flowering in November.
VENEZUELA. Terr. Amazonas: Cerro Duida; Rio Cunucunuma, Maguire,Cowan & Wurdack
29682 fl (NY); 29703 fl fr (NY).

This species is closest to H. cowanii,but differs in the orbicular leaves with sub-
cordate bases, the longer more lax inflorescence, the fewer stamens, and the more
pubescent style and interior of the receptacle. It differs from H. punctillatain the
acuminate leaf apex, and in the tomentose inflorescence which has thicker branches.
It differs from H. scabrain the leaf shape, the pubescence of the inflorescence, and in
the subsessile flowers.

45. Hirtella guyanensis (Fritsch) Sandwith, Kew Bull. 1931: 377. 1931.
Parinari
guyanense (Aublet)Lem6e.
Fritsch,Ann. Naturh.Mus. Wien 5: I4. I890, non P. guyanense

Tree to 30.0 m tall, the young branches tomentellous, soon becoming glabrous
and lenticellate. Leaves oblong to ovate-orbicular, coriaceous, 5.5-I4.0 cm long,
3.3-8.0 cm broad, abruptly acuminate at apex, with acumen 2.0-Io.o cm long,
subcordate, rounded, or subcuneate at base, usually with an appressed pubescence
beneath, rarely glabrous; primary veins 7-9 pairs, prominulous above, prominent
beneath; midrib prominulous above, prominent and pubescent beneath; petioles
3.0-5.0 mm long, tomentellous when young, terete, eglandular. Stipules ca 4.0 mm
long, lanceolate, pubescent, eglandular, caducous. Inflorescences terminal and sub-
terminal panicles 4.0-7.0 cm long, the rachis and branches tomentose. Bracts and
bracteoles 2.5-5.5 mm long, numerous, ovate, tomentose, persistent, eglandular.
Flowers 7.0-I0.0 mm long. Receptacle cylindrical-campanulate, subsessile, tomen-
tose on exterior, glabrous within except at throat and beneath style. Calyx lobes acute,
tomentose on both surfaces. Petals 5, white, glabrous. Stamens 6-8, unilateral, with
short filamentous staminodes opposite them, the filaments glabrous, slightly exceed-
ing calyx lobes. Style hirsute up to one third its length. Ovary inserted at mouth of
receptacle, pilose. Fruit not seen.
TYPE.Schomburgk i68, Guyana, fl (holotype, W; isotypes, K, P).
314 Flora Neotropica

/',/ 1

SD '
E".
CJLokenr E

FIG 49. Species of Hirtella. A-C, H, orbicularis(Maguireet al 29622); A, habit, x 0.5; B, flower,
x 5; C, flower section, x 3. D-F, H. cowanii (Cowanet al 30o84, 31237); D, habit, xo.5; E, flower
section, x5; F, flower, x5.
Hirtella 315

DISTRIBUTION. Primary forest on high ground in eastern Venezuela and Guyana.

Flowering August to February.
VENEZUELA. Bolivar: Aristeguieta3739 fl (VEN); Bernardi3001 fl (FI, VEN); 7321 fl (K, NY,
US, VEN). GUYANA. Fanshawe3o6(F.D. 3042) fl (K, NY, US); 1148 (F.D. 4184) fl (K); Jenman
4734 fl (BM, K); Tillett & Tillett 45048 fl (F, NY, US).

LOCAL NAME. Venezuela: Yucare.

46. Hirtella lightioides Rusby, Bull. N. Y. Bot. Gard. 4: 350. I907.

Hirtella acuminataRusby, Mem. N. Y. Bot. Gard. 7: 248. I927, synon nov. Type. Rusby 652,
Bolivia, fl (holotype, NY; isotypes, GH, K, US).

Shrub or small tree, the young branches tomentellous, becoming glabrous with
age. Leaves elliptic, subcoriaceous, 9.0-I7.0 cm long, 2.5-7.3 cm broad, acuminate
at apex, with acumen 5.0-Io.o mm long, rounded to subcuneate at base, sparsely
hirsute beneath, with a few palisade glands; primary veins Io-I4 pairs, plane above,
prominent beneath; midrib prominulous above, prominent beneath, tomentellous
on both surfaces; petioles 3.0-5.0 mm long, terete, eglandular, tomentellous. Stipules
3.0-5.0 mm long, linear, persistent, pubescent, eglandular. Inflorescences terminal
and axillary panicles 7.0-1 1.0 cm long, the rachis and branches hirsutulous-tomen-
tellous. Bracts and bracteoles 2.0-4.0 mm long, linear to oblong, tomentellous,
persistent, eglandular. Flowers 6.0-8.0 mm long. Receptacle campanulate, tomen-
tellous on exterior, glabrous within except at throat; pedicels 2.0-5.0 mm long.
Calyx lobes acute, tomentellous on exterior, puberulous within. Petals 5, white,
glabrous. Stamens 4-6, unilateral with a few short filamentous staminodes opposite
them, the filaments glabrous, far exceeding calyx lobes. Style hirsute up to half its
length. Ovary inserted at mouth of receptacle, villous. Fruit not seen.
TYPE. Bang 2418, Bolivia, fl (holotype, NY; isotypes, C, F, LE, M, S, US).
DISTRIBUTION. Primary forest, in northern and central Bolivia.
BOLIVIA. Krukoff10733 fl (A, F, MICH, NY, S, US); R. S. Williams 1568 fl (K, NY, US).

47. Hirtella aramangensis Prance, sp nov Fig 50 D-F.

Arbor, ramulis juvenilibus tomentellis mox glabris. Folia alternata petiolata;

petiolo 7.0-8.0 mm longo, tereti, eglanduloso, tomentello; laminae ellipticae,
coriaceae, 7.0-12.5 cm longae, 3.5-5.0 cm latae, apice in acuminem 3.0-Io.o mm
longum contractae, basi rotundatae, subtus appresso-pubescentes,pauciglandulosae,
haud papillosae, costa media supra leviter prominente, subtus prominente, utrinque
tomentella; costis secundariis I0-13 jugis, utrinque leviter prominentibus. Stipulae
parvae, lineares, tomentellae, eglandulosae, caucae. Flores circa 6.0 mm longi, in
paniculis subcorymbosis 4.0-6.5 cm longis dispositi, rachi ramisque tomentellis.
Bracteae bracteolaeque ovatae, tomentellae, glandulosae, persistentes.Receptaculum
campanulatum, extus tomentellum, intus basimversusglabrum; pedicelli circa 0.5 mm
longi. Calycis-lobi acuti, extus tomentelli, intus griseo-puberuli. Petala 5, glabra,
alba. Stamina 6. unilateralia, staminodiis filamentosis brevis opposita; filamenta
glabra, longe exserta. Ovarium ad faucem receptaculi insertum, pilosum. Stylus e
basi ovarii ortus, versus basim hirsutus.Drupa oblonga; epicarpio breviter tomentoso;
mesocarpio tenue; endocarpio tenue, duro, fibroso, intus sparse hirsuto.
316 Flora Neotropica

TYPE.Woytkowski 5625 Peru, Amazonas, Aramango, fl fr (holotype, S; isotypes,

GH, R 116328, US).
DISTRIBUTION. Known only from the type gathering, collected in forest, flowering
in April.
This species is close to H. rasa, but differs in the rounded leaf bases without the
characteristic constriction of H. rasa, in the far fewer palisade glands, the more
pubescent leaf undersurface, the tomentellous branches and petioles, the subcorym-
bose inflorescense, and the persistent bracts and bracteoles.

48. Hirtella rasa Standley, Publ. Field Mus. Bot. 17: 252. 1937; Macbride,
Fl. Peru 1077. I938.

Tree to 25.0 m tall, the young branches puberulous, soon becoming glabrous
and lenticellate. Leaves elliptic, coriaceous, 9.0-I5.0 cm long, 3.5-6.5 cm broad,
acuminate at apex, with acumen 2.0-7.0 mm long, subcuneate at base, papillose
and bearing only stiff appressed hairs beneath, with many palisade glands towards
base; primary veins 8-Io pairs, prominulous on both surfaces; midrib plane to
prominulous above, prominent beneath, puberulous to glabrescent on both surfaces;
petioles 4.0-5.0 mm long, terete, eglandular, glabrescent. Stipules ca 3.0 mm long,
linear, puberulous, caducous, eglandular. Inflorescences terminal lax panicles
7.0-I9.0 cm long, the rachis and branches sparsely puberulous. Bracts and bracteoles
2.0-4.5 mm long, ovate, puberulous, eglandular, caducous. Flowers ca 6.o mm long.
Receptacle campanulate, with short gray-tomentum on exterior, glabrous within
except at throat; pedicels 0.5-I.0 mm long. Calyx lobes rounded to acute, shortly
tomentellous on exterior, gray-puberulouswithin. Petals 5, white, glabrous. Stamens
6, unilateral, with short filamentous staminodes opposite them; filaments glabrous,
far exceeding calyx lobes. Style hirsute up to half its length. Ovary inserted at mouth
of receptacle, villous-tomentose. Fruit (young fruit only seen) ellipsoid; epicarp
tomentellous; pericarp thin, hard, bony, sparsely hirsutulouswithin.
TYPES.Klug 4249, Peru, San Martin, fl (holotype, F; isotypes, A, BM, K, NY,
S, US); Krukoff5525,Brazil, Acre, fl (paratypes,A, BM, F, K, LE, M, MICH, NY, RB,
S, SP, US, WIS).
DISTRIBUTION. Known only from the type gatherings made in primary forest in
Amazonian Peru and western Amazonian Brazil.

49. Hirtella scabra Bentham, Jour. Bot. Hooker 2: 2 18. 1840; Hook. f., Mart.
Fl. Bras. 14(2): 39. 1867.

Shrub, the young branches tomentellous, becoming glabrous and lenticellate

with age. Leaves ovate, thick-coriaceous, 2.5-7.ocm long, 1.5-4.0 cm broad,
acuminate at apex, with acumen 2.0-I2.0 mm long, rounded at base, sparsely
appressed-pubescent to glabrescent beneath, the palisade glands frequent, promi-
nently reticulate-venose above; primary veins 7-9 pairs, prominulous on both
surfaces; midrib prominulous above, prominent and hirsute beneath; petioles
2.o-3.o mm long, terete, eglandular, tomentellous when young. Stipules I.0-3.0
mm long, linear, caducous, eglandular. Inflorescences terminal and axillary panicles
2.0-9.0 cm long, the rachis and branches short-tomentellous. Bracts and bracteoles
o0.-30. mm long, acute, persistent, tomentellous, eglandular. Flowers 4.0-6.0 mm
Hirtella 317

FIG 50. Species of Hirtella. A-C, H. angustifolia(Lofgrenet al I595); A, habit, x o.5; B, flower,
x 3; C, flower section, x 3. D-F. H. aramangensis
( Woytkowski5625); D, habit, x o.5; E, flower, x 4; F,
flower section, x 4.
318 Flora Neotropica

long. Receptacle campanulate, tomentellous on exterior, glabrous within except at

throat; pedicels 1.5-4.0 mm long. Calyx lobes acute, tomentellous on exterior, gray-
pubescent within. Petals 5, white, glabrous. Stamens 5-7, unilateral; filaments
glabrous, far exceeding calyx lobes. Style hirsute up to one third its length. Ovary
inserted at mouth of receptacle, pilose. Fruit ellipsoid; epicarp smooth, glabrous;
mesocarp thin, fleshy; endocarp thin, hard, bony, hirsute within.
TYPE.Schomburgk 1051, Guyana, fl (holotype, K; isotypes, BM, CGE, F, NY).
DISTRIBUTION. Savannas and scrub forest in Venezuela and Guyana. Flowering
October to April.
VENEZUELA. Bolivar: Bernardi6620 fl (G); Cardona1870 fl (NY, US); Lasser I704 fl fr (F, K,
NY, US, VEN); Maguire33236 fl fr (NY); 33602 fl (NY); Steyermark 59311 fl (F, NY); 59395 fl (F, NY,
VEN); Tamayo3237 fl (VEN); 4308 fl (VEN); Ule 86i6fl (MG). Terr. Amazonas: Maguire,Cowan&
Wurdack30577 fl fr (NY); 30805 fl (NY); 36575 fr (NY); Spruce3548 fl (BM, BR, NY, P); Wurdack&
Adderley43693 fl (NY). GUYANA. Schomburgk 570 fl (F, K, NY, OXF, P); 865 fl (F, K); Tillett &
Tillett 45656fl (NY). BRAZIL. Amazonas: Frdeset al 22730 fl (IAN, NY); 29112 fl (IAN, UB).

The type collection of this species; Schomburgk o105, bears the same number as
that of H. velutina,=H. glandulosa(no. 20). These two species are very different and
the duplication no doubt merely reflects a mistake in the collector's records.

50. Hirtella bicornis Martius & Zuccarini Abh. Akad. Miinchen I: 377.
1832; Zucc., Flora Is(Beibl. 2): 80. 1832; Hook. f., Mart. Fl. Bras.
14(2): 39. i867. Fig. 5I.

Tree to 25.0 m tall, or shrub, the young branches puberulous to glabrescent,

lenticellate. Leaves membraneous to subcoriaceous, oblong to ovate, 3.5-9.5 cm
long, 1.7-4.0 cm broad, acuminate and often cuspidate at apex, with acumen 2.0-
15.0 mm long, rounded to subcuneate at base, glabrous beneath or sparselyappressed-
pubescent, the palisade glands few; primary veins 8- 3 pairs, prominulous on both
surfaces; midrib plane or nearly so above, prominent and glabrescent or puberulous
beneath; petioles 0.5-3.0 mm long, terete, eglandular, puberulous when young.
Stipules I.0-2.0 mm long, lanceolate, tomentellous, eglandular, caducous. Inflores-
cences predominantly axillary panicles I.O-4.5 cm long, the rachis and branches
glabrescent to tomentellous. Bracts and bracteoles 1.0-2.0 mm long, acute, persistent,
eglandular. Flowers 3.0-5.0 mm long. Receptacle campanulate, puberulous to
glabrescent on exterior, glabrous within except at throat; pedicels 3.0-6.0 mm long.
Calyx lobes acute, puberulous or glabrescent on exterior, gray-puberulous within.
Stamens 3-5, unilateral, the filaments glabrous, far exceeding calyx lobes. Style
hirsute up to one third its length. Ovary inserted at mouth of receptacle, pilose. Fruit
ellipsoid, the immature compressed longitudinally along center between apex and
base, appearing two-chambered with two lobes at the extreme apex; epicarp smooth,
glabrous; mesocarp thin, fleshy; endocarp thin, hard, bony, hirsute within.

Key to the Varieties of Hirtellabicornis

mm long, thick and obtuse,
I. Leaves ovate to elliptic, acuminate at apex, with acumen 4.0-Io.o
usually rounded at base; inflorescences glabrescent; shrubs or small trees of open beaches.
a. var bicornis.
i. Leaves usually oblong, usually cuspidateat apex, with acumen 7.0-15.0 mm long, thin,
usuallysubcuneateat base; inflorescencesusuallytomentellous;large foresttrees.
b. var pubescen.s.
 Hirtella 319

20

A Inft. pubgsct't
Leaf base 5ubcunecatd

4 Infl. pubcscemt

15. aLeaf bas vounted

InLt. glabrous
& b ra var. pubtscens .- LeoJ base roudet

S ,a
A

- /" vovar.
bicormis

1-5 2 5 3-0 5-5 -4

Lea Breacdt. - cms.

FIG 51. Pictorialized scatter d;agram of Hirtella bicornis.

5oa. Hirtella bicornis var bicornis

Hirtella bicornisMartius & Zuccarini, Abh. Akad. Minchen I: 377. 1832.
HirtellapraealtaSagot, Ann. Sci. Nat. Bot. VI. 15: 304. I883. Type. Sagot 792, French Guiana,
fl (holotype, P; isotypes, BM, BR, GOET, K, NY, S).

Leaves ovate to elliptic, acuminate at apex, with acumen 4.0-I0.0 mm long,

thick and obtuse, usually rounded at base; inflorescences glabrous or glabrescent;
shrub or small tree of open beaches.
TYPE. Martius sn, Brazil, Para, fl (holotype, M; isotypes, BR, M).
DISTRIBUTION. Sandy river banks and beaches, and sandy savannas in French
Guiana, eastern Amazonian and northeastern Brazil. Flowering July to December.
BRAZIL. Amazonas: Frdes 25333 fl (IAN, LE); SchwackeIII-542 fl (GOET, R). Parf: Black
47-1744 fl (IAN, NY); 55-18730 fl (IAN, NY); 57-20068 fl (IAN, NY); Black & Ledoux50-10612 fl
(IAN, NY); Black et al 57-19453 fl (IAN); Cavalcante223 fl (MG); Ducke 2154 fl (IAN); MG 2907 fl
(MG); MG 3729 fl (MG); MG 6902 fl (MG); MG 8034 fl (MG); MG 1910o fl (MG); MG 11313 fl
(MG); Huber MG 10457 fl (MG); Pires & Silva 4356 fl (IAN, NY, P); Riedel 1578 fl (OXF, US); sn
fl (LE); M. Silva 1627 fl (MG); 1659 fl (MG); Sprucesn fl (BM, NY); 322 fl (K, P); LI. Williams &
Silva 18246 fl (IAN, NY). Terr. AmapA: Frdes & Black 27564 fl (IAN); Pires & Cavalcante52246 fl
(IAN, MG, NY). Pernambuco: Lima 65-4367 fl (IPA, NY).

5ob. Hirtella bicornis var pubescens Ducke, Arch. Jard. Bot. Rio deJaneiro
3:269. 1922.
Hirtella caudataKleinhoonte, Rec. Trav. Bot. Neerl. 22: 389. 1925; in Pulle, Fl. Suriname 2(I):
450. I939. Type. B. W. 4376, Suriname, fl (lectotype, U).

Leaves usually oblong, usually cuspidate at apex, with acumen 7.0-15.0 mm

long, thin, usually subcuneate at base; inflorescences usually tomentellous; large
forest tree.
320 Flora Neotropica

TYPE. DuckeMG 15299, Brazil, Para, fl (holotype, MG; isotype, BM, P).
DISTRIBUTION. Non-flooded forest in Colombia, Venezuela, the Guianas and
Amazonian Peru and Brazil. Flowering July to November in the Guianas, May to
October in Amazonia.
COLOMBIA. Santander: RomeroC. 4930 fl (COL, US). VENEZUELA. Zulia: Bernardi2184
fl (NY, VEN). GUYANA. A. C. Smith2607 fl (A, NY, US). SURINAME. B. W. 1203 fl (U)13; 3007 fl
(A, C, IAN, U)13; 3061 fl (A, C, IAN, U)13; 3900 fl (U)13; 3976 fl (NY, U)13; 6243 fl (K, NY, U);
Heyligers448 st (U); Stahel 86 fl (NY); 206a fl (A, IAN, NY); VanDonselaar3693 fl (U). FRENCH
GUIANA. Irwin 48594 fl (IAN, MG, NY); Melinonsn (1863) fl (BR, GH, K, P, US); Wachenheim 183
fl (NY, P). PERU. ? Huanuco: Sandeman3406 fl (F). BRAZIL. Amazonas: Corner ro fl (IAN);
Froes21272 fl (IAN, K, NY, US); 25484 fl (IAN, NY, RB); Krukoff5099fl (A, F, K, MICH, NY, S, U,
US); Rodrigueset al 1778 fl (IAN, INPA 8141, MG, NY); 7103 fl (INPA 15765, NY); Spruce1708 fl
(GH, K, LE). Parl: Pires 6792 fl (IAN); 6863 fl (IAN); 7069 fl (IAN); 7122 fl (IAN, UB); Pires &
Black 340 fl (IAN, NY); Pires, Frdes & Silva 5826 fl (IAN, NY); Prance & Silva o0916 fl (IAN, NY);
Prance & Pennington1517 st (NY); Silva o112 fl (IAN, NY); ii6I fl (IAN, NY); 1167 fl (IAN, NY);
1376 fl (IAN, NY). Terr. Amapa: Frdes26625 fl (IAN, LE); Pires, Rodrigues& Irvine50273 fl (F, MG,
NY, US); 505I8 fl (F, GH, IAN, MG, NY, US); 50650 fl (F, IAN, MG, NY, US); 51271 fl (F, GH,
IAN, INPA, MG, NY, US). Terr. Rond6nia: Pranceet al 5434 fl (INPA, NY); 58io fl (INPA, NY);
J. F. Silva 28 fl (IAN). Mato Grosso: Lima58-3184 fl (IPA, NY, RB).

LOCAL NAMES. Suriname: Bokobokotokon.Brazil: Goiabinha.

The material brought together here under Hirtellabicornisundoubtedly belongs

to the same species. It was analyzed carefully, leading me to concur with Ducke who
designated the forest population a distinct variety. This variety cannot be considered
a distinct species because of the obvious morphological overlap with var. bicornis.The
majority of collections fall easily into one of the varieties, but the overlap of each
character is complete. Weak separations were obtained on scatter diagrams. Figure
51 is a pictorialized scatter diagram of the characters used to separate the varieties of
H. bicornis.Hirtellacaudataand H. praealtaare not distinct and must be reduced to
synonymy. The wide and broken geographical distribution indicates that this is a
widespread but apparently uncommon species.
Hirtella bicornisis most closely related to H. scabra,but differs in the thinner
leaves with a more pointed cuspidate acumen, and in the inconspicuous venation of
the upper surface of the leaf.

51. Hirtella angustissima Sandwith, Kew Bull. 1939: 10. 1939.

Tree to 6.o m tall, the young branches sparsely tomentellous, soon becoming
glabrous. Leaves narrowly lanceolate, coriaceous, 5.5-I8.0 cm long, 0.5-2.2 cm
broad, gradually tapering to a narrow acuminate or acute tip at apex, rounded at
base, glabrous beneath, with frequent palisade glands; primary veins 10-I2 pairs,
slightly prominulous and inconspicuous on both surfaces; midrib prominulous above,
prominent beneath; petioles 1.5-3.0 mm long, terete, eglandular, tomentellous when
young, soon becoming glabrous. Stipules 2.0-3.0 mm long, linear, persistent, eglan-
dular, pubescent. Inflorescences terminal and axillary racemes Io.o-I5.0 cm long,
the rachis sparsely tomentellous. Bracts 0.5-1.5 mm long, eglandular or with sessile
glands, persistent, acute, puberulous. Flowers ca 5.0 mm long. Receptacle campanu-
late, sparsely puberulous to glabrescent on exterior, glabrous within; pedicels 7.0-
I i.o mm long, sometimes with a solitary stipitate gland arising near base. Calyx
lobes acute, glabrescent on exterior, sparsely puberulous within, Petals 5, blue,
13 Paratypes of H. caudata.
Hirtella 321

glabrous. Stamens 4, unilateral, the filaments far exceeding calyx lobes, glabrous.
Style hirsute on lower portion only. Ovary inserted at mouth of receptacle, pilose.
Fruit not seen.
TYPES. Sandwith I246, Guyana, fl (holotype, K; isotype, NY); Tutin 6o0,
Guyana, fl (paratypes, BM, K, NY); Jenman884, Guyana, fl (paratype, K); 970 fl
(paratype, K).
DISTRIBUTION. River banks and islands in open places, confined to the Potaro
River region of Guyana.
GUYANA.Appunsn fl (BM); Cowan& Soderstrom
2o01 fl (NY, US); Maguire& Fanshawe
23030
fl (NY); Whitton
8i fl (K, NY, S).

This species is closely related to H. racemosa,but is easily distinguished from all

other species of Hirtellaby its narrowly lanceolate leaves.

52. Hirtella tenuifolia Prance, sp nov Fig. 52 A-E.

Arbor parva, ramulisjuvenilibus sparse tomentellis mox glabris. Folia alternata

petiolata; petiolo 2.0-3.0 mm longo, tereti, eglanduloso, juvenili hispido; laminae
lanceolatae, coriaceae, 9.0o-4.0 cm longae, 2.1-3.5 cm latae, apice in acuminem
io.o-I8.o mm longum contractae, basi subcordatae, subtus glabrae, versus basim
glandulosae; costa media supra leviter prominente, subtus prominente, glabra;
costis secundariis I 1-13 jugis, utrinque leviter prominentibus. Stipulae ca 2.o mm
longae, lineares, filamentosae, hispidae, eglandulosae, persistentes. Flores 4.0-5.0
mm longi, in racemis terminalibus 9.0-I4.0 cm longis dispositi, rachi sparse puberuli.
Bracteae bracteolaeque I.0-2.0 mm longae, lanceolatae, persistentes, tomentellae,
cum glandulis sessilibus munitae, Receptaculum campanulatum, extus puberulum,
intus versus basim glabrum; pedicelli 5.0-Io.o mm longi, eglandulosi. Calycis-lobi
acuti, utrinque puberuli. Petala 5, rosea, glabra. Stamina 5, unilateralia; filamenta
glabra, longe exserta, Ovarium tomentellum, ad faucem receptaculi insertum. Stylus
e basi ovarii ortus, basim versus hirsutus. Fructus mihi ignoti.
TYPE. Maguire,Pires & Maguire47110, Brazil, Terr. Amapa, Rio Oiapoque,
southeast of Clevelandia, fl (holotype, NY; isotype, IAN).
DISTRIBUTION. Primary and secondary forest in French Guiana and Territorio
de Amapa, Brazil.
FRENCH GUIANA. Melinonsn fl (K). BRAZIL.Terr.AmapA:Rio Oiapoque;Rio Pontanari,
Irwin,Egler& Pires47295fl (IAN, NY).

Closely related to Hirtellaparaensis,but differs in the eglandular pedicels, in the

bracteoles with sessile glands, the shorter receptacle-tube with the ovary inserted
terminally, the shortly tomentellous ovary, the one extra stamen, and the tertiary
venation of the leaf which is prominulous and not impressed as in H. paraensis.

53. Hirtella pilosissima Martius & Zuccarini, Abh. Akad. Munchen I: 373.
I832; Zucc., Flora s5(Beibl. 2): 78. 1832; Hook. f., Mart. Fl. Bras.
14(2): 30. 1867; Macbride, Fl. Peru Io77. 1938.
Hirtellapurusana
Huber, Bol. Mus. Emilio Goeldi 6: 72. 19o0. Type. GoeldiMG 3913, Brazil,
Amazonas,fl (holotype,MG; isotypes,BM, RB; fragmentF).

Tree to 7.0 m tall or shrub, the young branches densely pilose, becoming
322 Flora Neotropica

E~~~~~i~

x5.

X5.
Hirtella 323

glabrous with age. Leaves oblong, chartaceous, 7.0-I 7.0 cm long, 2.5-6.5 cm broad,
acuminate at apex, with acumen 7.0-16.0 mm long, subcordate to rounded at base,
hirsute beneath on venation, pilose above when young, the margins pilose, with
palisade glands scarce; primary veins 10-14 pairs, plane, or nearly so above, promi-
nent beneath; midrib prominulous and pilose above, prominent beneath; petioles
I.0-2.0 mm long, pilose, eglandular, terete. Stipules to i.o cm long, linear, pilose,
eglandular, persistent. Inflorescences terminal racemes 7.0-28.0 cm long, the rachis
pilose-tomentellous. Bracts and bracteoles 1.5-3.5 mm long, linear, tomentellous,
with a few sessileglands, persistent. Flowersca 4.0 mm long. Receptacle campanulate,
sparsely pilose on exterior, glabrous within except for deflexed hairs at throat;
pedicels 1.5-3.0 mm long. Calyx lobes acute, sparsely pilose on exterior, puberulous
within. Petals 5, glabrous. Stamens 3-6, unilateral, with a toothed portion of the
circle opposite them, the filaments far exceeding calyx lobes, glabrous. Style hirsute
up to half its length. Ovary inserted at mouth of receptacle, pilose. Fruit oblong-
pyriform; epicarp smooth, glabrous; mesocarp thick, fleshy; endocarp thin, fibrous,
densely tomentellous-hirsutewithin.
TYPE.Martius3039, Brazil, Amazonas, fl (holotype, M; isotypes, BR, K).
DISTRIBUTION. Non-flooded forest in western Amazonia. Flowering June to
November.
COLOMBIA. Putumayo: VogelmannI305 fl (U). Vaupes: Schultes& Cabrera15311 fr (F, GH,
NY, US). PERU. Loreto: LI. Williams 804 fl (A, F, US); rI83 fl (F, US). BRAZIL. Amazonas:
Krukoff6983 fl (A, K, NY); Pires & Black 889 fl (GH, IAN, NY). Terr. Rond6nia: Pranceet al 5404 fl
(INPA, NY). BOLIVIA. Pando: Pranceet al 5793 fl (INPA, NY); 5861 fl (INPA, NY).

LOCAL NAME. Peru: raku-shimbillu.

This species is close to H. racemosa, but differs in the extreme pilosity of the leaves,
branches, and inflorescence. It is also close to H. strigulosa,but differs in the sessile
glands of the bracts, and again greater pilosity, especially of the leaf margins.
in

54. Hirtella gracilipes (Hooker f.) Prance, stat nov

Hirtella americanaAublet var. gracilipesHooker f., Mart. Fl. Bras. 14(2): 34. I867, pro parte,
exsiccatum peruvianum exclusum.

Tree to 6.o m tall or shrub, the young branches sparsely pilose to glabrescent.
Leaves oblong, coriaceous, 5.0-I .5 cm long, 2.0-4.2 cm broad, gradually tapering
from middle to the acuminate apex, with acumen 4.0-I6.o mm long, subcordate to
rounded at base, glabrous or with a few stiff appressed hairs beneath on primary
venation, the palisade glands scarce; primary veins 6-8 pairs, prominulous on both
surfaces; midrib prominulous above, prominent and sparsely appressed-pubescent;
petioles 1.0-3.0 mm long, terete, eglandular, sparselypuberulous. Stipules ca I.o mm
long, subulate, caducous. Inflorescences terminal and axillary racemes 4.5-14.0 cm
long, the rachis sparsely hirsutulous to puberulous or glabrescent. Bracts 0.5-2.0 mm
long, ovate to lanceolate, membraneous, caducous, the margins bearing sessile glands
with translucent glandular secretions, sparsely appressed-pubescent or glabrous on
exterior. Flowers 5.0-7.0 mm long. Receptacle campanulate, sparsely appressed-
pubescent to glabrous on exterior, glabrous within except at throat; pedicels 6.o-
I6.0 mm long, slender. Calyx lobes acute, conspicuously reflexed, glabrescent on
exterior, gray-puberulous within. Petals 5, white to pale purple, glabrous. Stamens
4-6, unilateral, the filaments far exceeding calyx lobes, dark purple, connate at base
and up to I.o mm. Style hirsute on lower one-third. Ovary inserted at or near mouth
of receptacle, pilose. Fruit ellipsoid; epicarp smooth, glabrous, longitudinally striate;
mesocarp thin, fleshy; endocarp smooth, hard, bony, hirsute within.
324 Flora Neotropica

TYPES. Gardner2565, Brazil, Piaui, fl (lectotype, K; isolectotypes, BM, CGE,

F, NY, P, US); 3708, Brazil, Goias, fl (paratypes, BM, CGE, GH, K, NY, P); Pohl
2282, Brazil, Goias, fl (paratypes, BR, NY, W).
DISTRIBUTION. Riverine forest and forest on slopes in the Planalto of central
Brazil and its fringes. Flowering throughout the year.
Representative collections: BRAZIL. Par,: AlvarengaRB 90574 fl (RB); Bockerman181 fl (IAN,
SP); Silva 711 fl (IAN). Piaui: Snethlage66i fl (F). Mato Grosso: Collenettei45 fl (NY); Dorrien-Smith
132 fl (K, NY); Hoehne2033 fl (R); 2094 fl (R); Hunt 5600 fl (K, NY); 5682 fl (K, NY); Irwin et al
16292 fl (NY); Kuhlmann300 fl (R); 301 fl (R); 469 fl (RB 17986, US); Kuntzesn fl (NY); Lima 58-
3078 fl (IPA, NY, RB); Malme 1170 fl (S); 1773 fl (LD, S); Maguire,Pires, Maguire & Silva 56326 fl
(NY); Prance& Silva 59127 fl (NY, UB); 59146 fl (NY, UB); 59434 fl (NY, UB); Moore78 fl (BM, K,
NY). Goias: Belem 1996 fl (IAN, NY, UB); Burchell5964 fl (K, NY); 7079 fl (GH, NY, P, US); 7398
fl (K); Dawson 14270 fl (NY); Duarte9369A fl (RB); 9393 fl (RB); Glaziou 2zzz1 fl (BR, C, K, P);
Irwin et al 13722 fl (NY); 14980 fl (NY); LaneSP 69989 fl (SP); Ule 178 fl (P, R). Dist. Federal: Coelho
IJNPA16714 fl (INPA, NY); Duarteet al 82 fl (NY); Irwin et al 5324 fl (NY, UB); 5815 fl (NY, UB);
6197 fl (NY); 7887 fl (NY); 8465 fl (NY, UB); I5656 fl (NY); 15895 fl (NY); Pires et al 9403 fl (NY,
UB); Prance& Silva 59052 fl fr (NY, UB); Ramos6718 fr (K, NY). Bahia: Luetzelburg1882 fl (RB 61 14);
Zehntner40I fl (R, RB). Minas Gerais: Assis o09 fl (GH, R); Barreto6787 fl (F, R); 9158 fl (F, R);
Claussen1097 fl (NY, P); Duarte5697 fl (F, RB); Glaziou 14680 fl (A, BR, C, LE, P); 1468I fl (BR, LE);
I5944 fl (BR, C, GH, LE, P); Heringer5140 fl (NY, UB); Hoehne6078 fl (R); Macedo416 fl (BM, SP,
US); 2524 fl (S); Magalhaes214 fl (IAN, US); Markgraf3415 fl (F, RB); RegnellIII-I50 fl (BR, C, K,
LD, LE, R, S, US); Riedel802 fl (US); Sena, Herb. Schwacke2457 fl (R); Warmingsn fl (C). Sao Paulo:
Hemmendorf194 fl (S); Kuhlmann1593 fl (SP). BOLIVIA. Cardenas3012 fl (F); Rusby1371 fl (BM, GH,
LE, MICH, NY, US).

This species is very close to Hirtellabrachystachya,which differs in the longer, more

slender pedicels, the shorter inflorescence, the less pubescent leaf undersurface, and
in the absence of sessile glands on the bracteoles. Hirtellagracilipesand H. brachystachya
stand alone in the genus with the glandular secretions of the bracteoles and fre-
quently the calyx lobes. The secretions make the fresh inflorescence viscous. These
are the only New World species of Chrysobalanaceaewith such viscosity, a condition
common among the African Hirtellas and some species of Acioa. Hirtellagracilipes
and H. brachystachya are also closely related to H. caduca,which differs in bearing
stalked glands on the bracteoles, in the more cuspidate leaf apex, and the absence of
viscous secretions on the bracts and calyx lobes.
I have elevated Hooker's variety H. americana(=H. racemosa)var gracilipesto
specific rank because it is significantly different from the very variable H. racemosa.
Since H. racemosa is so variable, its boundaries had not been well defined owing to the
previous lack of critical study of the large amount of material which exists. Hirtella
gracilipesdiffers from H. racemosain the presence of viscous secretions on the bracts
and calyx lobes, in the caducous bracts and bracteoles, the lax inflorescence with
longer pedicels, and the glabrous rachis of the inflorescence.
When Hooker described H. americanavar gracilipes,he cited in addition to the
three Brazilian type collections listed above, Peruvian material of Ruiz & Pavon.
This last material better fits H. brachystachya, and is considered under that species in
this work.

55. Hirtella brachystachya Spruce ex Hooker f., Mart. Fl. Bras. 14(2): 34,
t. 9(2). 1867.

Tree to 6.o m tall or shrub, the young branches glabrescent. Leaves oblong to
oblong-elliptic, coriaceous, 6.o-Io.5 cm long, 2.3-3.5 cm broad, acuminate at apex,
with acumen I.o0-15.0 mm long, rounded to subcordate at base, glabrous beneath,
Hirtella 325

with palisade glands frequent towards base; primary veins 7-IO pairs, prominulous
on both surfaces; midrib prominent and glabrous on both surfaces; petioles I.0-2.0
mm long, terete, eglandular, glabrous. Stipules ca I.o mm long, subulate, caducous.
Inflorescences terminal and axillary racemes 1.5-5.0 cm long, the rachis glabrous.
Bractsand bracteoles0.5-1.5 mm long, ovate, glabrescent, caducous, with translucent
glandular excretions on margins. Flowers 4.0-6.0 mm long. Receptacle campanulate,
glabrous on exterior, glabrous within except at throat; flowering pedicels I3.0-
25.0 mm long, slender, glabrous. Calyx lobes acute, conspicuously reflexed, glabrous
on exterior, gray-pubescent within. Petals 5, white, glabrous. Stamens 6, unilateral,
the filaments far exceeding calyx lobes, glabrous. Style pilose up to half its length.
Ovary inserted at mouth of receptacle, pilose. Fruit ellipsoid; epicarp smooth,
glabrous; mesocarp thin, fleshy; endocarp thin, hard, bony, hirsute within.
TYPE. Spruce2268, Brazil, Amazonas, fl (holotype, K; isotypes, BM, BR, C,
CGE, GH, GOET, LD, LE, NY, OXF, P, RB).
Riverine forest in western Amazonia. Flowering April to Septem-
DISTRIBUTION.
ber.
COLOMBIA. Vaupes: Cuatrecasas 6909 fl (COL); 7 12 fl (COL); Schulteset al 24172 fl (ECON).
VENEZUELA. Terr. Amazonas: Maguire, Cowan & Wurdack31018 fl (NY); Wurdack& Adderley
43I79 fl (NY); 43311 fl (NY). PERU. without locality: Ruiz & Pavonsn14 fl (BM, G, K). BRAZIL.
Amazonas: DuckeRB 35576 fl (K, RB); Fres 21 I28 fl (F, IAN, K, NY, S); 28193 fl (IAN); Luetzelburg
22125 fl (R).

This species is close to H. gracilipes;see note under that species.

56. Hirtella racemosa Lamarck Encyc. 3: I33. 1789. Roem. & Schult., Linn.
Syst. Veg. ed. 9. 5:274. I819; DC., Prodr. 2: 529. 1925; Sandw., Kew Bull.
1931: 376. I93I; Kleinh. in Pulle, Fl. Suriname 2(I): 446. I939; Macbride,
Fl. Peru Io77. 1938, non H. racemosa Ruiz & Pavon (I802). Fig 53.
Hirtella americanaauct. non Linnaeus sensu Aublet, P1. Guiane I: 247, t. 98. 775; Persoon, Syn.
I: 250. I805; Zucc., Flora 15(Beibl. 2): 82. I932; Benth., Jour. Bot. Hooker 2: 216. 1840;
Hook. f., Mart. Fl. Bras. I4(2): 33. I867; Standley, Contrib. U.S. Natl. Herb. 23: 344.
1922; tom. cit. 27: i88. 1928. Type. Aubletsn, French Guiana, fl (P-Rousseau).

Shrub or small tree, or occasionally medium-sized tree, the young branches

usually puberulous but varying from tomentellous to glabrous or sparsely hispid.
Leaves elliptic to oblong, coriaceous, 3.5-I6.5 (-I9.5) cm long, 1.5-7.0 cm broad,
acuminate at apex, with acumen I.0-I4.0 (-25.0) mm long, subcordate to cuneate at
base, the lower surface glabrous or sparsely appressed-pubescent; primary veins
6-Io pairs, prominulous above, prominent beneath; petioles I.0-3.0 mm long, terete,
eglandular, glabrous to puberulous. Stipules 1.5-5.o mm long, linear, eglandular,
persistent, glabrous to hirsutulous. Inflorescences terminal and axillary racemes
5.0-29.0 cm long, the rachis puberulous to glabrescent. Flowers 3.5-6.0 mm long.
Bracts and bracteoles 0.5-3.0 mm long, sparsely appressed-pubescent, persistent,
usually with sessile glands mostly towards base, or with a single large rotund concave
gland, glandular or eglandular at apex, if glandular then bract with a distinct
flattened lamina, the bracts very rarely eglandular. Receptacle campanulate, sparsely
puberulous to glabrescent on exterior, glabrous within except at throat; pedicels
1.5-I .0 mm long. Calyx lobes acute, sparsely puberulous on exterior, puberulous
within. Petals 5, glabrous, pink. Stamens 5-7, unilateral, the filaments far exceeding
calyx lobes, glabrous or sparsely hirsute towards base. Style hirsute up to one third
14
Paratype of H. americana(H. racemosa)var gracilipesHooker f.; see note under H. gracilipes.
326 Flora Neotropica

0.5
/,' Ax Lw9ge hollow glands
o1~~~~0
1>8~~~
A Al
//^~, ^ ,Leaf basc CIcak,t,
var. hexandrca A Snmall
plane 9land
4, A, A A A A, A ea base. cunseat
A A
, 4 Large hollow glands
Ar
,."-'-.., Leaf basa rouned
8 Al AlA
A.
A A A Lacrrgqhottowglands
A)A A| A A, , Leaf base subcordale
E 7 , >^^ , A A, A-' s*A""^^
), AA A) ' ?A A " A
t Smalltp glands
CE 7. QF^*- Leaf base.- ound
A A A S
A matt plane glands
,,flekrmediate
A AAAAAAAAA ^ Leaf base subcordaat
6A) *A) AAAA7 AAA
4A A , AA AAA A 49
^ ^ A A A
5 AA
A ??AAAA A AA
AAA,'-
A"AAAAA AAA A A AA
^ A A A
~C_ ,^, ,
44- - AAAAAA ' A.
A
AA- & A AA
A A A A A
AAAAAAAAAA
6AAAAAAA A AAA AAA
A&A A A94
.A A A

var. racemosa
A.,A
AAA A
'''AAA
5s ' ' ' A A I
4 5 6 7 8 1 10 11 12 13 14 15 16 17 18 1q
Leaf length - cms.

FIG 53. Pictorialized scatter diagram of Hirtella racemosa.

its length. Ovary inserted at mouth of receptacle, pilose-tomentose. Fruit ellipsoid;

epicarp smooth, glabrous; mesocarp thin, fleshy; endocarp thin, hard, bony, hirsutu-
lous within.

Key to the Varieties of Hirtella racemosa

i. Pedicels eglandular, thick or slender, 1.5-10.5 mm long; bracteoles with large sessile concave
glands or with small plane glands or apical glands; leaves 3.5-19.5 mm long.
2. Leaves 7.0-I9.5 cm long, usually subcordate at base, less frequently rounded to cuneate;
leaf venation inconspicuous above; pedicels usually thick, 1.5-7.0 (-9.0) mm long;
bracteoles mostly bearing large sessile concave glands, or reduced to a single large concave
gland, less frequently the glands smaller and plane. a. var racemosa.
2. Leaves 3.5-9.5 (- 1.5) cm long, usually rounded to cuneate (very rarely subcordate) at
base; leaf venation usually prominently reticulate above; pedicels slender, 4.0-10.5 mm
long; bracteoles with small sessile plane glands or an apical gland only. b. var hexandra.
i. Pedicels with a single large gland arising near base, thick, 2.0-3.0 mm long; bracteoles with
large sessile concave glands; leaves I 1.5-I4.0 cm long. c. var glandipedicellata.

56a. Hirtella racemosa var racemosa

Hirtella racemosaLamarck, Encyc. 3: I33. 1789.
Hirtella americana auct, non Linnaeus, sensu Aublet, P1. Guiane I: 247, t. 98. 1775; Benth., Jour.
Bot. Hooker 2: 216. 1840; Hook. f., Mart. Fl. Bras. 14(2): 33. 1867, pro parte quoad var
americanaet var oblongifoliapro parte.
Tachibota guianensis Aublet, P1. Guiane I: 287. I775, see Sandwith, Kew Bull. 17: 261. 1963.
Type. Aubletsn, French Guiana, st (P-Rousseau, BM).
Salmasia guianensis (Aublet) G. F. Gmelin, Linn., Syst. Nat. ed. 3. 2: 500. 1791.
Salmasia racemosa Willdenow, Linn., Sp. ed. 4. I: 1502. 1797, nom illegit.
Hirtella violaceaSteudel, Flora 26: 761. I843. Type. Hostmann860, Suriname, fl (holotype, P;
isotypes, BM, K, L, LE, M, NY, S, U).
Hirtellla 327

Hirtella strigulosaSteudel, Flora 26: 761. I843. Type. Hostmann85ib, Suriname, fl (holotype, P;
isotypes, LE, NY).
HirtellascandensHoffmansegg ex Roemer & Schultes, Linn., Syst. Veg. ed. 9. 5: 274. 1819; DC.,
Prodr. 2: 529. 1825. Type. Siebersn, Brazil, Pari, fl (holotype, B, Herb, Willdenow 4849;
photo, NY).
Hirtellanemorosa Hoffmannsegg ex Roemer & Schultes, Linn., Syst. Veg. ed. 9. 5: 274. I819; DC.,
Prodr. 2: 529. 1825. Type. Siebersn, Brazil, Para, fl (holotype, B, Herb. Willdenow 4848;
photo, NY).
Hirtella racemosavar oblongifolia(A. P. De Candolle) Standley, Publ. Field Mus. Bot. 17: 252.
1937 pro parte, Typum De Candollei exclusum.
Hirtella racemosavar metallicaCuatrecasas, Fieldiana Bot. 27: 60. 1950. Type. Cuatrecasas19925,
Colombia, Valle, fl (holotype, F; isotypes, COL, MAD).

Leaves 7.0-I9.5 cm long, usually subcordate at base, less frequently rounded to

cuneate; venation inconspicuous above; pedicels usually thick, I.5-7.o(-9.o) mm
long, eglandular; bracteoles mostly bearing large sessile concave glands, or reduced
to a single large concave gland, less frequently glands smaller, plane and not concave,
or the bracteoles with apical glands.
TYPE. Badier sn, French Guiana, fl (holotype, P-LAM).
DISTRIBUTION. Primary forest especially beside rivers and streams, but also in
understory of forest on non-flooded ground from Panama and Colombia through the
Guianas and Amazonia to northern Bolivia.
Representative collections: PANAMA. Johnston 1673 fr (A); I749 fl fr (A). COLOMBIA.
Antioquia: RomeroC. 1547 fl (COL). Santander: Haught 1528 fl (US); RomeroC. 4948 fl (COL, US).
Valle: Cuatrecasas16262 fl (F). Cauca: Gorgona Is. Barclay926 fl (BM). Nariiio: RomeroC. 2598 fl
(COL). Vichada: RomeroC. 1279 fl (COL). Amazonas: Schultes& Black 8415 fl (COL, US); Schultes&
Cabrera13995 fl (COL, NY, US); 14167 fl (COL). Vaup6s: Ferndndez2005 fl (COL, US). VENEZUELA
Monagas: Breteler5140 fl (U); Wurdack& Monachino39431 fl (NY); 39507 fl (LE, NY). Terr. Delta
Amacuro: Gines4937 fl (US); Rusby& Squires261 fl (A, BM, F, GH, K, MICH, NY, S, US); Steyermark
87209 fr (GH, NY, US); Wurdack& Monachino39574 st (NY). Bolivar: Blanco 453 fl (NY); Breteler
3872 fl bud (NY, U, US, VEN); 5z11 fl (U); Cardona1927 fl (NY, US, VEN); 2095a fl (US); Killip
37465 fl (A, IAN, NY, US, VEN); Lasser1626 fl (F, K, NY, US, VEN); Steyermark 74639 fl fr (F, GH,
NY, VEN); 86380 fl (VEN); 86480 fl (VEN); 87927 fl fr (NY); 88406 fl (F, NY, US); 88410 fl fr
(F, NY, US); LI. Williams 1556 fl (F, US). TRINIDAD. Brittonet al 423 fl (GH, NY, US); Broadway
5591 fl (A, F, K); 6619 fl (BM, F, K, S); 6844 fl (BM, F, K); 7549 fl (BM, S); 7704 fl (US); 9373 fl
(A, FI, K, P); Cruegersn fl (GH, GOET, LE, NY); 2 fl (K); 300 fl (GOET); Fendler1o2oB fl (BM, K,
NY); Kuntzesn fr (NY); Richardson931 fl (K); A. C. Smith10083 fl (A, NY, US); R. O. Williams11897
fl (K, NY, P). TOBAGO. Beard 112 fl (A, NY); Broadway4133 fl (C); 4457 fl (BM, F, G, GH, NY).
GUYANA. Abraham o03 fl (NY); Cowan39357 fr (F, NY, RB, US); De La Cruz 1847 fl (F, GH, K, NY,
US); 1933 fl (F, GH, K, NY, US); 2531 fl (F, GH, K, NY, US); 2566 fl (F, GH, K, NY, US); 3748
fl (F, GH, K, NY, US); 4005 fr (F, GH, NY, US); 4320 fr (F, GH, K, NY, US); 4397 fl (F, GH, K,
MICH, NY, US); Fanshawe505(F.D. 324I) fr (K, NY); For. Dept. 6767 fl (MICH, NY); Gleason208
fl (GH, K, NY, US); 233 fl (NY); Hitchcock17061 fl (GH, NY, S, US); Lang & Persaud41 fl (F, NY,
US); Linder 119 fl (GH, NY, S); Maguire & Fanshawe23364 fl fr (A, BR, F, K, NY, US); 23515 fl
(BR, GH, F, K, NY, US); 32451 fr (NY); Persaud1617 fl (K, NY); Sandwith253 fl (K, NY); Schomburgk
23 fl (BM, BR, GH, K, LE, NY, P); 840 fl (BM, F, K, NY, P, W); A. C. Smith 2660 fl (A, F, K, NY,
P, S, US); 3625 fl (A, F, IAN, K, LE, NY, P, US); Tillett & Tillett 45475 fl (F, GH, NY, US); Tutin
30 fl (BM, US); Wilson-Browne203(F.D. 5686) fl (K, NY). SURINAME. B.W. 2319 fl (K); 2644 fl
(IAN, NY); 3367 fl (A, C); 5421 fl (K); Florschiitz1520 fl (C, NY); Focke1228 fl (U); Guppy& Lindeman
sn fl (NY); Hostmanni85 fl (BM, G, K, NY); Irwin, Prance,Soderstrom& Holmgren55164 fl (F, GH, NY,
S, US); 55205 fl (F, GH, NY, US); 55288 fl (F, GH, NY, US); 55543 fl (F, GH, NY, US); Lanjouw
1233 fl (NY); Lanjouw& Lindeman1124 fl (NY); Mennega136 fl (C); 433 fr (A, C); Maguire24947 fl
(NY); Samuels518 fl (GH, K, NY); Versteeg728 fl (U); VanDonselaar1152 fl (U); Wullschlaegel807 fl
(GOET). FRENCH GUIANA. Cowan38847 fl (GH, MICH, NY, S); 38857 fl (NY, P); For. Serv.7058
fl (NY); Irwin et al 47474 fl (IAN, NY); 48327 fl (IAN, MG, NY, US); 48589 fr (NY); Leprieursn st
(F, G, GH, K, P); Martin sn fl (BM, F, K, P, R); Melinon sn (I845) fl (F, GH, P, R); sn (I862) fl
(F, GH, P, R); sn (I864) fl (F, K, LA, P); 75 fr (BM, BR, F); 79 fr (F); 327 fl (A, BM, F, K, LA, P,
US); Poiteausn fl (NY, P); Sagot 915 fl (K, NY, P); 917 fl (BM, BR, K, NY, P, S). PERU. Loreto:
Killip & Smith 29795 fl bud (A, NY, US); Klug 203 fl (F, NY, US); 1333 fl (F, NY, US); 1989 fl (F,
GH, K, MICH, NY, S, US); Tessmann3115 fl (NY, S, US); Ll. Williams2358 fr (F); 2765 fl (F, US);
2990 fl (F, US). San Martin: Klug 2704 fl (A, BM, F, GH, K, NY, S, US); 2714 fl (A, BM, F, GH, K,
328 Flora Neotropica

NY, S,US). Madrede Dios: Rauhetal 1595 fl (F). Without locality: Mathews 1599 fl (OXF). BRAZIL.
Amazonas: Baldwin3286 fl (US); ChagasINPA 1252 fl (IAN, INPA, NY); Emygdio 1845 fl (R); Frdes
22556 fl (IAN); Krukoff4977 fr (A, K, MICH, NY, S, US); 5887 fl (A, K, MICH, NY, S, US); 6688
fl (A, BR, F, K, MICH, NY, RB, S, US); Luetzelburg22162 fl (R); Pires 221 fl (IAN, NY); 774 fr
(IAN); Pranceet al 2314 fl (INPA, MG, NY); 2432 fl (INPA, MG, NY); 2485 fl fr (IAN, INPA, MG,
NY); SchwackeIII-151 fl (GOET); Ule 5653 fl (GOET, K, PRC). Terr. Roraima: Black et al 51-13371
fl (IAN); 51-13516 fl (IAN); 51-13803 fl (IAN); 51-13918 fl (IAN, NY); R. Boyan 231 fl (INPA);
Pranceet al 3970 fl fr (INPA, MG, NY); M. Silva o04 fl (MG). Para: Black 47-1687 fr (IAN, NY);
48-3384 fl (IAN); 49-8109 fl (IAN, NY); Burchell9143 fl (K); Frdes & Black 24632 fl (IAN, LE, NY);
Guedes1234 fl (BM, RB, S); Guppy566(F.D. 7581) fl (NY); 596(F.D. 7611) fl fr (NY); Killip & Smith
30577 fl fr (NY, US); Krukoff 0o59 fl (A, K, NY, P); 1064 fl (A, K, NY, P); Mus. Goeldi o0858fl (BM,
MG, S); Pires & Silva 4594 fl (IAN, NY, P); Sampaio5059 fl (R); 5156 fl (R); 5465 fl (R); Snethlage99
fl (F, GH, US); Sprucesn fl (GH, NY); 222 fl (K, P); Zernysn fl (W). Terr. AmapA: Cowan38586 fl
(NY); Egler & Irwin 46506 fl (NY); Frdes & Black 27365 fl (IAN, NY); Irwin 48765 fl (F, GH, IAN,
MG, NY, S, US); Irwin et al 47439 fl (IAN, NY); Luetzelburg20252 fl (R); Pires et al 50334 fl (F, GH,
IAN, MG, NY, US); 5091o fl (F, IAN, MG, NY, US); 511z8 fl fr (IAN, NY); 52290 fl (IAN, MG,
NY); 52395 fl (IAN, MG, NY). Maranhao: Frdes 1933 fl (A, BM, F, K, MICH, NY, P, S); Snethlage
305 fl (F). Piauf: Netto 32 fl (NY, R). Acre: Emygdio 1869 fl (R); Krukoff5354 fl (A, BM, F, FI, LE,
MICH, NY, S); Pranceet al 6423 fl (INPA, NY); 7323 fr (INPA, NY); 76o0 fl (INPA, NY); 7635 fl
(INPA, NY); Ule 9412 fl (FI, GH). Terr. Rond6nia: Cordeiro& Silva 400 fl (NY); Pranceet al 5244 fl
(INPA, NY); 6286 fl (INPA, NY); 6505 fl (INPA, NY). BOLIVIA Rusby704 fl (BM, F, GH, K, LE,
MICH, NY, P, US); Krukoff10876 st (A, F, K, MICH, NY, U); Pranceet al 5715 fl (INPA, NY);
6089 fl (INPA, NY); 6467 fr (INPA, NY).

LOCALNAMES. Venezuela: Carbonero,Cenisa negra,Jicaquillo, Piquapiquamorada.

Guyana: Bokobokotokon, Suriname: KleinbladigeAnaura,Proewaipoeroe.French Guiana:
Akianteici, Gris-gris.
For discussion see notes at end of species.

56b. Hirtella racemosa var hexandra (Willdenow ex Roemer & Schultes)

Prance, comb nov
Hirtella hexandraWilldenow ex Roemer & Schultes, Linn., Syst, Veg. ed. 9. 5: 274. I819; DC.,
Prodr. 2: 529. 1825.
Hirtella americanaAublet var. hexandra(Willdenow ex Roemer & Schultes) Hook. f., Mart. Fl.
14(2): 33. I867.
Hirtella nitida Humboldt & Bonpland ex Roemer & Schultes, Lin., Syst. Veg. ed. 9. 5: 274.
1819; DC., Prodr. 2: 529. 1825. Type. Humboldt& Bonplandsn,Cuba, fl (holotype, B, Herb
Willdenow 4846; isotype, P; photo, NY).
Hirtella acayacensisMocifio & Sesse ex A. P. de Candolle, Prodr. 2: 529. 1829; Zucc., Flora 15
(Beibl. 2): 87. 1832. Type. Mociiio & Sess6, Icones florae Mexicane (ined.), plate 301
(G; photo, NY).
Hirtella oblongifoliaA. P. de Candolle, Prodr. 2: 529. 1825; Zucc., Flora I5(Beibl. 2): 82. I832.
Type. Herb. Lagasca78, Mexico, fl (holotype, G-DC).
Hir'ella coriaceaMartius & Zuccarini, Abh. Akad. Munchen i: 383. 1832; Zucc., Flora 15(Beibl.
2): 83. I832. Type: Martius sn, Brazil, Minas Gerais, fl (holotype, M).
Hirtella roseaMociiio & Sess6 ex A. P. de Candolle, Prodr. 2: 529. 1825, nom nud in syn.
HirtellafiliformisPresl, Symb. Bot. 2: 23. 1832. Type. Herb. Presl sn, Mexico, fl (holotype, PRC).
Hirtella americanaAublet var oblongifolia(A. P. de Candolle) Hooker f., Mart. Fl. Bras. 14(2): 34.
1867 pro parte quoad typum tantum.
Hirtella racemosaLamarck var oblongifolia(A. P. de Candolle) Standley, Publ. Field Mus. Bot.
17: 252. 1937 pro parte quoad typum tantum.

Leaves 3.5-9.0 (-i 1.5) cm long, usually rounded to cuneate (very rarely subcor-
date) at base; venation usually prominently reticulate above; pedicels slender
4.0-10.5 mm long, eglandular; bracteoles with small sessile plane glands, or with a
single apical gland only.
TYPE. Humboldt& Bonplandsn, Venezuela, fl (holotype, B, Herb Willdenow
4850; isotype, P; photo, NY).
Hirtella 329

DISTRIBUTION. Growing in a wide range of open habitats, especially open river

banks and rocky islands, lower forest slopes, savanna margins, and in disturbed
secondary forest. From central Mexico through Central America to Panama, in
Colombia, Venezuela, the Guianas, Amazonia to Bolivia, and in northeasternBrazil.
Representative collections: MEXICO. Nayarit: Feddema994 fl (MICH); McVaugh 18778 fl
(BM, GH, MICH, US); 19242 fl (MICH, US); Rzedowski14383 fl (MEXU). Michoacan: Langlasse
647 fl (G, GH, K, US). Veracruz: Gilly & Herndndez57 fl (GH, MICH); King 979 fl (MICH); Mell sn
fl (NY, US); Miranda 8545 fl (MEXU); Orcutt3047 fl (MO); C. L. Smith o08I fl (GH); LI. Williams
8401 fl (F, MICH). Guerrero: Hinton 10773 fl (BM, F, GH, K, MEXU, MO, NY, S, US); 11040 fl
(GH, K, NY, US); Palmer485 fl (A, F, GH, MICH, MO, NY, US); Paray 4035 fl (MEXU). Oaxaca:
Alexander142 fl (MEXU, MICH, NY, US); Andrieux372 fl (GH); McVaugh22244 fl (MICH); Mexia
9236 fl (F, GH, K, NY, S, US); Nelson 2715 fl (US); Orcutt3047 fl (F); 3232 fl (BM, F, GH, K, US);
Schultes& Reko948 fl (ECON, GH); Reko3405 fl (US); LI. Williams9340 fl (F, MICH, P, US); 9246
fl (F); 9487 fl (A, F, MICH, NY, US); 9568 fl (F, MICH, P). Tabasco: Barlow 6/5 fl (GH, MEXU);
30/62 fl (MEXU, MICH); 30/151 fl (BM, F, MEXU, MO); E. P. Johnsonsn fl (GH, NY); Matuda
3097 fl (F, MEXU, MICH, NY); Rovirosa85 fl (US). Chiapas: Enriquez7310 fl (MEXU); Matuda
388 fl (A, MEXU, MICH, MO, NY, US); 68i fl (MEXU, MICH, US); 16454 fl (BR, F, MEXU,
MICH); 16887 fl (F, MEXU); Nelson 3853 fl (GH, US). Without precise locality: Jurgensen924
fl (GH); Mocino & Sesse 87 fl (BM); 876 fl (F). GUATEMALA. Aguilar 190 fl (A, MICH); 1716 fl
(F); Bernouilli & Cario 77 fl (K); 2960 fl (GOET, S); 2961 fl (GOET); H. Johnson 1284 fl (F, US);
Kellerman6672 fl (F, MEXU, MICH); Lundell2217 fl (BM, F, MICH, S, US); 3049 fl (A, MICH);
3079 fl (F, MICH); 4874 fl (F); Standley24107 fl (GH, NY, US); 72343 st (F); 73122 fr (A, F); 73139
fl (F); 78957 fl (A, F); 87349 fr (F); 87466 fl (F); 87801 fl (A, F); Steyermark 38057 fl (F); 38982 fl (F);
39560 fl (F); 44667 fl (F, MICH); 45147 fl (F, MICH, US); 45466 fl (F, MICH); SuttonHayes 30 fl
(GH, US); VonTiirckheim4087 fl (ECON, F, LD, S, US); 7620 fl (A, NY, US); C. L. Wilson365 fl (F).
BRITISH HONDURAS. Bartlett 11537 fr (GH, MICH, NY, S, US); 11840 fr (A, F, MICH);
11932 fl (F, K, MICH, US); 12974 fl fr (MICH, NY, US); Campbellsn fl (NY); 23 fl (K); Castillo47
fl (F); Gentle65 fl (F, MICH, NY, S); 943 fl (A, K, LD, MICH, NY, P, S); 1876 fl (MICH, NY);
2406 fl (A, F, K, MEXU, MICH); 7598 fl (C, F); Hunt 182 fl (BM, F, US); Lundell20 fl (F, MICH);
565 st (F, NY, US); 619 fl (A, F, GH, K, NY, US); Meyer 126 fl (F); Molina R. 74 fl (F, GH); Pelly I
fr (F); 84 fl (F, K); RecordBH33 fl (NY, US); Schipp33 fl (A, BM, F, GH, K, LE, MICH, NY, S, US);
Stocker9 fl (A, F). HONDURAS. Molina R. 11619 fl (F, NY); Standley54273 fl (A, F, US); Thieme5221
fl (A, F, FI, GH, K, LE, MICH, NY, US); P. Wilson211 fl (F, GH, NY, US); VonHagen 1353 fl (F,
NY). EL SALVADOR. Standley20116 fl (GH, NY, US). NICARAGUA. Long o06fl (F); Molina R
i85I fl (F, GH); 14918 fl (F, NY); 15204 fl (F); S. S. White5308 fl (MICH); Shank & Molina R. 4559
fl (GH); Standley20057 fl (F). COSTA RICA. Brenessn fl (NY); 1912 fl (NY); 3606 fl (A, F, NY);
4i83 fl (F, NY, P);5028fl (F, NY); 14927 fl (F, NY); 69gi8fl (F, NY); I7003 fl (F); 17009 fl (F, NY);
18889 fl (F, NY); 20284 fl (A, F, NY); 20323 fl (F, NY); 20463 fl (F, NY); Jimenez M. 327 fl (F);
Lankester1336 fl fr (F); Pittier 12088 fl (US); 12164 fl (US); QuirosC. 858 fl (F); Tonduz3982 fl (BR, F,
US); 6708 fl (BM, BR, US); L. 0. Williams et al 24215 fl (F, US); 24523 fl (F); WinzerlingII-2I fl
(US). PANAMA. Allen I603 fl (F); 4156 fl (F, MICH); Andersson sn fl (S); Billberg sn fl (S); Dwyer 1399
fl (MO, NY); Frost 70 fl (F); Harlow i fl (US); 94 fl (GH, US); Harvey5187 fl (F); Johnston226 fl
(GH); 1250 fl (FI, GH, US); Macbride2645 fl (F, US); Piper5575 fl (GH, S, US); 5596f (F, NY, US);
Pittier 2702 fl (NY, US); Seeman291 fl (GH); C. E. & H. M. Smith3263 fl (BM, F); Steyermark& Allen
17500 fl (BR, C, FI, K, MICH, MO, P, S); Wetmore& Abbe o05fl (A, F, GH); R. S. Williams91 fl fr
(NY, US); Woodworth& Vestal332 fl (F, MO). COLOMBIA. Boyaca: Haught 2630 fl (A, COL, F,
MICH, US). Meta: Cuatrecasas4097 fl (COL, F); Hermann11176 fl (COL). Cauca: Gorgona Is. St.
GeorgeExped.583 fl (S, US). Vaupes: Cuatrecasas7132 fl (COL, F, US); Gutierrez& Schultes985 fl (GH),
Schultes& Cabrera12678 fl (COL, F, GH, NY, US). Without precise locality: Linden1637 fl (BM, BR,
K, LE, OXF, P, S); Mutis 157 fl (US); 3969 fl (US). VENEZUELA. Zulia: Pittier 10602 fl (G, GH,
NY, P, US, VEN); Tejera22 fl (GH, US); 227 fl (US). Lara: Saer 711 fl (F, VEN). Carabobo: Saer
793 fl (NY, US, VEN). Guirico: Aristeguieta4493 fl (NY, VEN); Pittier 12358 fl (NY, US, VEN).
AnzoStegui: Croizat 167 fl (F); Herndndez102 fl (VEN); Steyermark60979 fl (F, NY, VEN). Barinas:
Aristeguieta1566 fl (F, US, VEN); Bernardi1107 fl (VEN). Apure: Vdlez2265 fl (VEN); 2259 fl (US);
2323 fl (VEN). Bolivar: Agostini320 fl (NY); Bailey A2 fl (NY); Cardona2387 fl (VEN); Croizat54 fl
(F); 55 fl (F); Curran33 fl (VEN); Killip 37283 fl (A, F, US, VEN); 37565 fl (US, VEN); Maguire
3272 fl (NY); Maguireet al 35864 fl (NY); 35999 fl (NY); LI. Williams 11203 fl (F, S, US, VEN);
11746 fr (US, VEN); 12653 fl (F, K, S, US, VEN); 12705 fr (F, K, S, US); 12718 fr (F, S, US);
13278 fl (F, K, S, US, VEN); Wurdack& Guppy33 fl (NY); Wurdack& Monachino39754 fl (NY);
39974 fl (NY); 41124 fl (NY). Terr. Amazonas: Breteler4689 fl (U); 4872 fl (U); Curran90 M fl (NY)
1784 fl (NY); i870 fl (NY); Cowan& Wurdack31487 fl (NY); Croizat 269 fl (NY); 669 fl (NY); Level
121 fl (NY); Maguire & Politi 27315 fl (NY); 28973 fl fr (NY); Maguire, Wurdack& Keith4155I fl
330 Flora Neotropica

(NY); LI. Williams 15004 fl (VEN); I5018 fl (F, US, VEN). GUYANA. Cook 151 fl (K, NY); Irwin
593 fl (US); Lanjouw& VanDonselaar928 fl (U); Schomburgk 80 fl (BM, BR, F, FI, K, P, US); A. C.
Smith 2189 fl (A, F, K, NY, P, S, US); Wilson-Browne307(F.D. 5736) fl (K, NY). PERU. Loreto:
Ferreyra4953 fl (US); 4978 fl (US); Killip & Smith 27549 fl (F, NY, US); 29651 fl (F, NY, US); Ule
6264 fl (K); 6453 fl (K, PRC); LI. Williams3829 fl (A, F, S); 3884 fl (A, F); 4041 fl (F, US); 7832 fl
(F, S, US); Woytkowski5782 fl (GH); 34432 fl (F). San Martin: Ferreyra4603 fl (US); 4636 fl (US);
Klug 4025 fl (BM, F, GH, K, NY, S, US); Scolnik1181 fl (P, US, W); LI. Williams5887 fr (F); 6154
fr (F, US); Woytkowski5405 fl (F, LE, P, S, US); 7256 fl (GH, US). BRAZIL. Amazonas: Black 48-
2415 fr (NY); Cardona 431 fl (F, US); Chagas i8 fl (INPA 6730, NY); 25 fl (IAN, INPA 6841, MG,
NY); INPA 428 fl (IAN, INPA, MG, NY); INPA 1827 fl (IAN, INPA, MG, NY); INPA 3622 fl
(IAN, INPA, MG, NY); INPA 7218 fl (IAN, INPA, MG, NY); Ducke2 fl (A, F, IAN, K, MG, NY,
R, S, US); Ferreira58-320 fl (INPA 6694, NY); Krukoff596 fl (A, BM, BR, F, LE, MICH, NY, RB,
S); Rodrigues& Coelho3402 fl (INPA 9952, NY); 6907 fl (INPA I5460, NY); Schwacke393 fl (GOET,
R); TackeuchiINPA 7813 fl (INPA, NY). Terr. Roraima: KuhlmannRB 2807 fl (K, RB, S, US);
Pranceet al 4123 fl (INPA, NY); 4543 fl (INPA, MG, NY); 673I fl (INPA, NY); 9418 fr (INPA, NY).
Para: Archer795I fl (F, IAN, NY, US); Baldwin 2748 fl (IAN, NY, US); Black 48-3448 fl (IAN);
50-9752 fl (IAN, NY); Burchll go91 fl (GH); 9278 fl (GH, K, NY, P, US); Drouet 2108 fl (F, GH,
MICH, S, US); Jobert & Schwacke27 fl (P, R); 273 fl (P, R); Kauffmann21 (F, US); Killip & Smith
30548 fl (NY, US); Mexia 6060 fl (A, BM, F, GH, K, US); Pranceet al 1276 fl (IAN, NY); Pires &
Black sn fl (GH); Herb. Richardsn fl (P); A. Silva 95 fl (IAN, NY); i66 fl (IAN, NY, US); 254 fl (IAN,
NY, US); Spruce1042 fl (K, NY, P). Maranhao: Black et al 54-16684 fl (NY); Don 32 fl (BR); 33 fl
(BR); Frdes 1809 fl (A, K, MICH, NY, P, S); i85i fl (A, BM, F, K, MICH, NY, P, S, US); 11533 fl
(A, F, K, MICH, NY, S, SP); 11643 fl (A, F, NY); 11707 fl (A, F, K, NY, S); 11867 fl (A, NY, US);
Jobert & Schwacke940 fl (P, R); Lisboa RB 4701 fl (RB, S); Registro2 fl (IAN, NY); Snethlage128 fl
(F, GH, US). Piaui: DuckeMG 807 fl (MG); Luetzelburg1214 fl (RB 5826). Ceara: Allemao& Cysneiros
567 fl (R); Dahlgren912 fl (F); 962 fl (A, F); DuckeMG 1429 fl (MG); Drouet2532 fl (F, GH, MICH,
NY, R, S, US); Gardner1590 (I591 some sets) fl (BM, GH, K, NY, S, US). Rio Grande do Norte:
Emygdio1738 fl (R); Lima 86 fl (NY, UB); Tavares127 fl (US); 267 fl (US); WurdackB 215 fl (NY).
Paraiba: Moraes 692 fl (INPA). Pernambuco: Gardner993 fl (BM, GH, K, P, S, US); Pickel 203 fl
(SP, US); 603 fl (R); Lima & Magalhaes52-1102 fl (R); Ridleyet al sn fl (BM); Schenck4123 fl (C);
Tavares479 fl (HST, US); Xaiz 283 fl (R). Terr. Rond6nia: Pranceet al 5360 fl (INPA, NY); 6656 fl
(INPA, NY); 6731 fl (INPA, NY); 6953 fl (INPA, NY). Mato Grosso: KuhlmannRB 17986 fl (S).
Bahia: Duarte9328 fl (RB I30647). Alagoas: Falcdoet al 1189 fl (RB). Minas Gerais: Glaziou 10705 fl
(C, K, P). BOLIVIA. Rusby702 fl (BM, F, GH, LE, K, MICH, NY, P, US).

LOCALNAMES.Mexico: Cerricillo,Escobilla, Jicaquillo. Guatemala: Aceituna.

Honduras: Pasta. British Honduras: Grenada,Wildpigeonplum, Wayamche(Mayan).
El Salvador: Icacomontes.Costa Rica: Murta, Serrecio.Venezuela: Jicaquillo.Brazil:
Ajurarana,Coracaode Jegro, Murtinha.Peru: Puccavarilla, Varillacaspi.
The following material is either intermediate between var racemosaand var
hexandra,or is inadequate to determine to variety.
NICARAGUA. Englesing 131 fl (F); Molina R. 1869 fl (F, GH); Proctoret al 27213 fl (NY).
COLOMBIA. Cauca, Gorgona Is.: St. GeorgeExped. 607 fl (US). Amazonas: Schultes& Cabrera13772
fl (GH, NY). VENEZUELA. Bolivar: Lasser 1492 fl (VEN); Steyermark& Gibson95764 fr (NY); LI.
Williams 12554 fr (F, US, VEN). Amazonas: Holt & Gehriger244 fl (NY, P, S, US, VEN); Maguire,
Wurdack& Bunting37471 fl (NY). GUYANA. Jenman4287 fl bud (K, US). FRENCH GUIANA. For.
Serv.5I I fl (P); Irwinet al 47338 fl (NY). BRAZIL. Para: Pranceet al 1344 fr (K, NY); 58731 fr (NY);
58669 fl (NY). Terr. AmapA: Pires & Cavalcante52619 fl (NY). Goias: Pranceet al 58642 fl (NY). Terr.
Rond6nia: Cordeiro& Silva 384 fl (NY). Alagoas: Nettosn fl (P).

56c. Hirtella racemosa var glandipedicellata Prance, var nov

Frutex. Folia I .5-I4.0 cm longa, basi subcuneata, supra venis inconspicuis.

Receptaculum campanulatum; pedicelli brevi, crassi, versus basim cum glande
unice stipitate ornati. Bracteolae cum glandibus sessilibus concavibus munitae.
TYPE. Von SneidernA I9go, Colombia, Caqueta, Morelia, fl (holotype, S; isotype,
COL).
Hirtella 331

DISTRIBUTION.
Known only from the type gathering.
This collection is near var racemosa,but differs in the glandular pedicels, the
longer inflorescences, and the subcuneate leaf bases. It may ultimately prove to be
specifically distinct, but I hesitate to describe yet another new species in this complex,
based on a single herbarium specimen, from such a poorly known area.
Hirtella racemosais one of the most widespread and commonest (hence much
collected) species of Chrysobalanaceae. Frequently misidentified and confused with
other species, it is a highly variable species. At the extremes of variation a casual
comparison of the material would suggest that at least two species are involved.
However, whichever characters are used, there are always a considerable number of
intermediates between the two types somewhere in the geographical range. The
characters all prove to be diflerential rather than diagnostic. It was found that two
distinct groups exist, but that they merge together linked by many intermediates.
The majority of collections fall readily into one of the groups, but since these groups
overlap, are defined entirely by differential characters, and are not geographically
isolated, they are treated here as varieties of the same species.
Figure 53, one of the scatter diagramsprepared for H. racemosa, shows the pattern
of variation quite clearly. When it is difficult to identify the variety of a collection, it is
helpful to measure the length of the longest leaf and the pedicel, and use fig 53
as a guide. The correct variety may be determined by which side of line A-B the
specimen in question falls. The two varieties, racemosaand hexandra,are not geogra-
phically isolated; they are in fact sympatric over much of their range, but occupy
different habitats, although not without some overlap. Variety racemosa tends to grow
in damp, heavily shaded primary forest, while hexandragrows in more open, often
drier places such as wooded slopes, savanna margins and secondary forest.
The relationship of H. racemosato H. gracilipesand other related species is dis-
cussed under each species.
Because of complex variation, Hirtellaracemosa has been much confused and mis-
interpreted by past workers. The name, H. racemosa, is used here since it is the oldest
name which may be legitimately applied to this species, although some of the synonyms
antedate it (eg Tachibotaguianensis).The epithet guianensisis preoccupied in H.
guianensis(Fritsch) Sandwith. The greatest confusion has resulted from Aublet's use
of the name americana for this species. HirtellaamericanaL. is a very different species;
see note under species 22. De Candolle (1825), in pointing out that Aublet's inter-
pretation was incorrect, cited H. americanaAublet as a synonym of H. racemosaand
treated H. americanaL. as a separate species. Later investigators differed and until
recently H. racemosa has been called H. americana, variously attributed to Linnaeus or
Aublet.
The name, H. acayacensis was proposed by A. P. de Candolle based on one of the
Mocinfoand Sesse tracings left at Geneva. This drawing is not one of those published
by A.L.P.P. de Candolle in the Calques,see p. 253. Consequently H. acayacensisis
typified by plate 301 of the Iconesflorae Mexicaneat Geneva, which is labelled H.
octandra.Judging from the drawing H. acayacensis falls well within the range of varia-
tion of H. racemosa var hexandrathe only species with a racemose inflorescence known
from Mexico.

57. Hirtella juruensis Pilger, Revista Mus. Paul. 13: 1250. 1922.

Tree, the young branches tomentellous, Leaves oblong, thin-corriaceous,

I9.5-25.0 cm long, 5.8-7.5 cm broad, broadest nearer to apex and tapering to base
332 Flora Neotropica

from above middle, acuminate at apex, with acumen 6.o-io.o mm long, subcuneate
at base, sparsely appressed-pubescent on venation beneath, with palisade glands at
base only; primary veins 13-I5 pairs, plane to slightly impressed above, prominent
beneath; midrib prominulous above, prominent beneath; petioles 2.0 mm long,
terete, tomentellous, eglandular. Stipules linear, persistent, puberulous, eglandular.
Inflorescences terminal racemes I4.0-I8.o cm long, the rachis sparsely puberulous.
Bracts and bracteoles I.o-2.5 mm long, oblong to oblong-lanceolate, puberulous,
persistent, with two large sessile glands towards base. Flowers 4.0-5.0 mm long.
Receptacle campanulate, puberulous on exterior, glabrous within except at throat;
pedicels I.0-I.5 mm long, eglandular. Calyx lobes acute, sparsely puberulous on
exterior, gray-puberulous within. Petals 5, glabrous. Stamens 5, unilateral, the
filaments far exceeding calyx lobes, glabrous. Style hirsute up to one third its length.
Ovary inserted at mouth of receptacle, pilose. Fruit not seen.
TYPE. Hoehnei809, Brazil, Mato Grosso, fl (holotype, B, lost; lectotype, R);
1816 fl (B lost); 4909 (B lost).
DISTRIBUTION.
Known only from the type gatherings collected at Juruena, Mato
Grosso.
Although most of the type material of H. juruensiswas lost in Berlin, the single
sheet examined suggests close affinity to H. racemosa.It differs from H. racemosa
in the
large leaves which usually taper from above the mid point, the greater number of
primary veins, and the slightly bullate mature leaves.

58. Hirtella kuhlmannii Pilger, Notizbl. Bot. Gart. Berlin 8: 538. I923.

Small tree, the young branches tomentose soon becoming glabrous with age.
Leaves oblong, coriaceous, 7.0-I3.0 cm long, 3.0-5.8 cm broad, acuminate at apex,
with acumen 3.0-6.0 mm long, rounded at base, slightly bullate above; hirsutulous
beneath, the palisade glands at base only; primary veins I I- 13 pairs, plane to slightly
impressed abDve, prominent and hirsutulous beneath; midrib prominulous and
tomentellous above, prominent and hirsutulous beneath; petioles 2.0-4.0 mm long,
terete, tomentellous, eglandular. Stipules linear, persistent, tomentellous, eglandular,
4.0-9.0 mm long. Inflorescences terminal racemes IO.O-25.o cm long, the rachis
sparsely puberulous. Bracts and bracteoles oblong to oblong-lanceolate, puberulous,
persistent, 2.0-5.0 mm long, with two large sessile glands towards base. Flowers
4.0-5.0 mm long. Receptacle campanulate, puberulous on exterior, glabrous within
except at throat; pedicels 2.0-4.0 mm long, eglandular. Calyx lobes acute, sparsely
puberulous on both surfaces, the margins often with sessile glands. Petals 5, glabrous.
Stamens 6, unilateral, the filaments far exceeding calyx lobes, glabrous. Style hirsute
at base only. Ovary inserted at mouth of receptacle, pilose. Fruit not seen.
TYPE. Kuhlmann g99o, Brazil, Mato Grosso, fl (holotype, B, lost; lectotype, R).
Known only from two collections from southern central Amazonia.
DISTRIBUTION.
Flowering June to July.
BRAZIL. Par,: Egler 832 fl (MG, NY).

This species is most closely related to H. juruensis,from which it differs in the

much smaller leaves which are hirsutulous beneath, and to H. racemosa,but differs
in the slightly bullate leaves, the hirsutulous lower surface of the leaves, and the
larger more persistent bracteoles.
Hirtella 333

59. Hirtella standleyi Baehni & Macbride in Macbride, Fl. Peru I078. 1938.

Tree to 4.0 m tall, the young branches tomentellous, becoming glabrous and
lenticellate with age. Leaves ovate, coriaceous, 4.5-9.5 cm long, 3. -6.o cm broad,
acuminate at apex, with acumen 2.0-4.0 mm long, cordate at base, sparsely hirsu-
tulous beneath on primary venation, with palisade glands rare; primary veins 8-Io
pairs, plane above, prominent beneath; midrib prominulous and puberulous above,
prominent and hirsutulous beneath; petioles I.0-I.5 mm l6ng, terete, eglandular,
tomentellous. Stipules ca 4.0 mm long, linear, pubescent, eglandular, persistent.
Inflorescencesaxillary and terminal racemes 12.0-I 7.0 cm long, the rachispuberulous.
Bracts and bracteoles 2.0-3.0 mm long, lanceolate, puberulous,persistent,eglandular.
Flowers ca 5.0 mm long, Receptacle campanulate, sparsely puberulous to tomen-
tellous on exterior, glabrous within except for retrorse hairs at throat; pedicels
1.0-2.5 mm long, some with a single large stipitate gland. Calyx lobes acute, reflexed,
sparsely hirsutulous on exterior, puberulous towards apex within. Petals 5, glabrous.
Stamens 6, unilateral, with a few short staminodes opposite them, the filaments far
exceeding calyx lobes, glabrous or sparsely hirsute towards base. Style hirsute up to
half its length. Ovary inserted at mouth of receptacle, pilose. Fruit ellipsoid; epicarp
smooth, sparsely puberulous; mesocarp thin, fleshy; endocarp hard, fibrous, ca 1.5
mm thick, hirsute within.
TYPES. Klug 3022, Peru, Loreto, fl (holotype, F; isotypes, A, K, NY, S, US);
2869, Peru, Loreto, fl (paratypes, A, GH, K, NY, S, US).
DISTRIBUTION. Known only from the type gatherings, collected in forest in the
vicinity of Balsa Puerto, Peru.

60. Hirtella longifolia Bentham ex Hooker f., Mart. Fl. Bras. 14(2): 3I. 1867.

Small tree, the young branches hispid. Leaves lanceolate, chartaceous, I9.0-
26.0 cm long, 4.0-5.8 cm broad, acuminate at apex, with acumen 8.0-20.0 mm
long, subcordate at base, hispid-hirsute beneath on principal venation, with palisade
glands present at base only, hispid on margins; primary veins I7-I9 pairs, prominu-
lous on both surfaces; midrib prominulous above, prominent and hispid beneath;
petioles 3.0-4.0 mm long, terete, eglandular, hispid. Stipules ca 8.0 mm long, fila-
mentous, persistent, hispid, eglandular. Inflorescences terminal and axillary racemes
I5.0-20.0 cm long, the rachis sparselypuberulous. Bracts to 4.5 mm long, lanceolate,
persistent, puberulous, eglandular or with a glandular apex. Flowers 5.0-6.0 mm
long. Receptacle campanulate, puberulous on exterior, glabrous within except at
throat; pedicels 8.0-i2.0 mm long, often with a single short-stalked gland arising
near base. Calyx lobes acute, puberulous on both surfaces, eglandular on margins.
Petals 5, glabrous, bluish. Stamens 5-6, unilateral, the filaments far exceeding calyx
lobes, glabrous. Style hirsute on lower portion. Ovary inserted at mouth of receptacle,
tomentose. Fruit not seen.
TYPE.Spruce2904, Brazil, Amazonas, fl (holotype, K; isotypes, BM, BR, CGE,
OXF, P).
DISTRIBUTION.Known only from the type gathering, collected in secondary forest
near Panure on the Rio Vaupes.
This poorly known species is distinct from all other species of Hirtellawith lanceo-
late leaves by the larger leaves with more numerous primary veins and by the densely
hispid leaf venation and young branches.
334 Flora Neotropica

6I. Hirtella lemsii L. O. Williams & Prance, sp nov Fig 52.

Arbor parva, ramulis juvenilibus hispido-tomentellis, mox glabris. Folia alter-

nata petiolata, petiolo 2.0-3.0 mm longo, tereti, eglanduloso, tomentoso; laminae
oblongae vel oblongo-ellipticae, membranaceae, 8.0-I4.5 cm longae, 3.0-6.0 cm latae,
basi subcordatae, subtusvenulis hirsutae; costa media supraleviterprominente, tomen-
tella, subtus prominente; costis secundariis IO-I2 jugis, supra planis, subtus promi-
nentibus. Stipulae 1.5-2.5 mm longae, lineares, tomentosae, persistentes, eglandu-
losae. Flores 5.0-6.o mm longi in racemis terminalibus subterminalibusque 24.o-
30.0 cm longis dispositi, rachi sparse tomentelli. Bracteae bracteolaeque I.0-2.0 mm
longae, lineari-filamentosae, persistentes, pubescentes, apicibus cum glandulo
unico munitis. Receptaculum campanulatum, extus puberulum, intus versus basim
glabrum; pedicelli 6.o-8.o mm longi, versus basim cum I-3 glandulis stipitatis
instructis. Calycis-lobi acuti, eglandulosi, utrinque puberuli. Petala 5, glabra. Stamina
3, unilateralia; filamenta glabra, longe exserta. Ovarium ad faucem receptaculi
insertum, pilosum. Stylus e basi ovarii ortus, hirsutus. Fructus mihi ignoti.
TYPE. Lems 5203, Costa Rica, Golfito, southwest of airstrip, fl (holotype, F;
isotype, NY).
DISTRIBUTION. Known only from the type gathering, collected in forest.
This species differsfrom related ones with similar pedicel glands in the 3 stamens,
the long inflorescence, and in the leaf shape. Hirtellalemsiiis also related to H. pendula,
but differs in the racemose inflorescence, the smaller flowers, and in the smaller leaf
acumen. The type specimen was kindly sent to me by Dr. L. O. Williams, who had
already noted its novelty, and who had proposed the manuscript name H. lemsii. I
am grateful for Dr. Williams' help with this species, and have named it for Kornelius
Lems, a promising ecologist who died in an auto accident in 1968.

62. Hirtella schultesii Prance, sp nov Fig 54 D-F.

Frutex alta, ramulis juvenilibus sparse tomentellis mox glabris. Folia alternata
petiolata; petiolo 2.5-4.0 mm longo, tereti, eglanduloso, juvenili tomentoso; laminae
oblongo-lanceolatae, coriaceae, 9.0-I6.0 cm longae, 2.2-4.3 cm latae, apice in
acuminem 9.0-I6.o mm longum contractae, basi leviter subcordatae, supra basim
leviter constrictae, subtus glabrae pauciglandulosae; costa supra leviter prominente,
subtus prominente cum pilis appressis paucis munita; costis secundariis I 1-13 jugis,
utrinque glabris leviter prominentibus. Stipulae 5.0-9.0 mm longae, lineares, per-
sistentes, pubescentes, eglandulosae. Flores 4.0-5.0 mm longi in racemis terminalibus
I0.0-I5.0 cm longis dispositi, rachi puberuli. Bracteae bracteolaeque circa 1.5 mm
longae, oblongae, persistentes, puberulae, apicibus saepius cum glandulis stipitatis
terminalibus. Receptaculum campanulatum, extus puberulum, intus versus basim
glabrum; pedicelli 3.0-5.0 mm longi, versus basim cum glandulo unico instructi.
Calycis-lobi acuti, utrinque puberuli. Petala 5, glabra. Stamina 5, unilateralia;
filamenta glabra, longe exserta. Ovarium tomentosum, ad faucem receptaculi inser-
tum. Stylus e basi ovarii ortus, basim verus hirsutus. Fructus mihi ignoti.
TYPE. Schultes& Ldpez 9344, Venezuela, Terr. Amazonas, Rio Negro, San
Carlos, fl (holotype, US).
DISTRIBUTION. Non-flooded forest in western Amazonia. Flowering December
to May.
COLOMBIA. Vaupes: Alto Vaupes, Nare, Gutierrez& Schultes84r fl (GH, NY). BRAZIL.
Amazonas: Sao Paulo de Olivensa, Creek Belem, Krukoff8727 st (A, BM, F, K, LE, NY, S, U); Rio
Hirtella 335

Negro near mouth of Rio Vaupes, Pires 413 fl (IAN); Rio Negro, Tapuruquara, Pires 600 fl (IAN);
Tabatinga, DuckeMG 6853 fl (MG); Rio Negro, San Gabriel, Spruce2367 fl (GH, GOET, LD, RB).

This species is very close to Hirtellaparaensisand H. spruceibut differs from both

in the distinctive leaf base. It further differs from H. spruceiin the larger number of
primary leaf veins, the absence of staminodes, and in the ovary inserted below the
mouth of the receptacle. Other differences from H. paraensisinclude the longer and
thicker pedicel glands, the extra stamen, and the puberulous branches and rachis.

63. Hirtella paraensis Prance, sp nov Fig 54 A-C.

Frutex vel arbor, ramulis juvenilibus hispidis mox glabris. Folia alternata
petiolata; petiolo 1.5-3.0 mm longo, tereti, eglanduloso, hispido; laminae oblongo-
lanceolatae, coriaceae, Io.o-I8.o cm longae, 2.1-4.7 cm latae, apice in acuminem
12.0-22.0 mm longum attenuatae, basi subcordatae, subtus glabrae pauciglandulosae
costa media supra leviter prominente, subtus prominente; costis secundariis 12-15
jugis, supra planis subtus prominentibus, glabris. Stipulae 5.o-8.o mm longae,
lineari-filamentosae, puberulae, eglandulosae, persistentes. Flores circa 5.0 mm longi
in racemis terminalibus axillaribusque 8.o-15.o cm longis dispositi, rachi sparse
hirsuti. Bracteae bracteolaeque 1.5-3.5 mm longae, lineares, pubescentes, eglandu-
losae, persistentes. Receptaculum campanulatum, extus sparse hirsutum, intus versus
basim glabrum; pedicelli 4.0-6.0 mm longi, versus basim glandulos I-2 stipitates
gerentes. Calycis-lobi acuti, extus sparse hirsuti, intus sparse puberuli. Petala 5,
glabra, purpurea. Stamina 4, unilateralia; filamenta glabra, longe exserta. Ovarium
ad medium receptaculi insertum, pilosum. Stylus e basi ovarii ortus, basim versus
hirsutus. Fructus mihi ignoti.
TYPE. Frdes 20377, Brazil, Para, Belem, Sao Miguel do Guama, fl (holotype, NY;
isotypes, IAN, K, US).
DISTRIBUTION.
Non-flooded forest in eastern Brazilian Amazonia. Flowering
August to February.
BRAZIL. Para: Belem; Sio Miguel do Guam,, Pires & Black 1Io fl (GH, IAN); 508 fl (IAN);
Pires 3132 fl (IAN, US); 3141 fl (IAN, NY); M. B. Silva 72 fl (F, IAN, K, NY, US); Rio Jari, Oliveira
3502 fl (IAN, UB); 3973 fl (IAN, UB); N. T. Silva 1533 fl (IAN, NY); Santarem, Jobert 863 fl (P, R);
Rio PacajA, Pranceet al 1403 fl (NY); Bel6m-Brasilia road, Km 147, Oliveira519 fl (IAN); Santa Rosa,
Fr6es31500 fl (IAN); Belem Utinga, Markgraf3816 fi (RB); Belem, grounds of IAN, Black & Ledoux
50-10223 fl (IAN); A. Silva 143 fl (IAN, US). Without precise locality: N. T. Silva i 6fl (IAN).

This species and the next three are very closely related. Hirtellaparaensisis
nearest to H. sprucei,but differs in the narrower more lanceolate leaves which grad-
ually taper to a long thin acumen, the greater number of primary veins (8-o1 pairs
in H. sprucei),the smaller number of fertile stamens, and the lower insertion of the
ovary. It differs from H. lancifoliain the much shorter pedicels, the smooth plane
venation of the leaf upper surface (impressed and slightly bullate in H. lancifolia),
the shorter petioles, and the lower insertion of the ovary. H. paraensisis cultivated in
Guanabara (Glaziou9786 fl K).

64. Hirtella sprucei Bentham ex Hooker f., Mart. Fl. Bras. 14(2): 3I, t. 9(I).
1867.
Small tree, the young branches hispid, soon becoming glabrous. Leaves oblong
336 Flora Neotropica

/ D

J Loken

FIG 54. Species of Hirtella.A-C, H. paraensis(Frdes20377, Pranceet al 1403); A, habit, x 0.45; B,

flower section, x 2.8; C, petal, x 2.8. D-F, H. schultesii(Gutilrrezet al 841, Krukof8727); D, habit, x 0.45;
E,petal, x 2.8; F, flower section, x 2.8.
Hirtella 337

to oblong-lanceolate, coriaceous, 8.0-1 7.0 cm long, 2.2-5.0 cm broad, acuminate at

apex, with acumen 5.0-I8.o cm long, subcordate at base, glabrous beneath, the
palisade glands rare; primary veins 8-Io pairs, prominulous and glabrous on both
surfaces; midrib prominulous above, prominent and glabrous beneath; petioles
2.0-3.5 mm long, terete, eglandular, glabrous or hispid when young. Stipules 4.0-
7.0 mm long, linear-filamentous, hispid to glabrous, eglandular, persistent. Inflores-
cences terminal and axillary racemes 12.0-27.0 cm long, the rachis hispid or glab-
rous. Bracts and bracteoles 1.5-3.0 mm long, oblong to lanceolate, hispid, eglandular,
persistent. Flowers 4.0-5.0 mm long. Receptacle campanulate, with a few stiff erect
hairs but otherwise glabrous on exterior, glabrous within except at throat; pedicels
4.0-5.5 mm long, most with one or more solitary stalked glands arising near base.
Calyx lobes acute, glabrescent on exterior, puberulous within, Petals 5, glabrous.
Stamens 6, unilateral, with short filamentous staminodes opposite them, the filaments
far exceeding calyx lobes, glabrous. Style hirsute up to one third its length. Ovary
inserted at mouth of receptacle, pilose. Fruit ellipsoid; epicarp smooth, glabrous;
mesocarp thin, fleshy; endocarp thin, hard, bony, costate, opening along longitudinal
lines of weakness, hirsute within.
TYPE.Blanchet3568, Brazil, Bahia, fl (holotype, K; isotypes, BM, BR, C, F, G,
GH, LE, P).
DISTRIBUTION.
Forests of northeastern and east central Brazil. Flowering July
to November.
BRAZIL. Pemambuco: Lima 58-3294 fl (IPA, NY). Mato Grosso; Maguire & Maguire 44525 fl
(NY); Prance& Silva 59426 fr (NY, UB). Bahia: Blanchet3975 fl (MG 3129). Sao Paulo: Riedel I666 fl
(C, NY, US). Rio deJaneiro and Guanabara: Hoehnesn fl (SP 24867); Lallemandsnfl (R 7142); Moreira
13z fl (R); Regnell54 fl (S); Vidalsn fl (R 11636). Without precise locality: Bowie & Cunningham
sn fl
(BM, NY); Freyreissn fl (S); Raddisn fl (FI); Riedelsn fr (US); Schreinersn fl (R 7148).

Hirtellaspruceisubsp meridionalis Hassler, does not belong with this species, but is
part of H. burchelliiBritton.
For relationship of the present species to Hirtellaparaensissee under the latter
name. Hirtellaspruceiis closely related to H. burchellii,but differs in the longer pedicels,
the terminal insertion of the ovary, the larger number of stamens, the eglandular
bracts (those of H. burchelliiusually bearing sessile and apical glands), in the glabrous
leaf undersurface, and in the smaller number of primary veins.

65. Hirtella lancifolia Ducke, Arch. Jard. Bot. Rio de Janeiro 3: 264. 1922.

Shrub or small tree, the young branches tomentellous. Leaves oblong, thick-
coriaceous, I0.0-24.0 cm long, 2.8-5.5 cm broad, acuminate at apex, with acumen
5.0-I5.0 mm long, rounded to subcuneate at base, hirsute beneath with palisade
glands frequent towards base, slightly bullate above; primary veins I4-I5 pairs,
slightly impressed above, prominent beneath, the secondary venation impressed
above with the interstices appearing bullate; midrib prominulous above, prominent
and hirsute beneath; petioles 3.5-7.0 mm long, terete, tomentellous, eglandular.
Stipules 5.0-7.0 mm long, linear, tomentellous, eglandular, persistent. Inflorescences
terminal and axillary racemes 8.0-15.0 cm long, the rachis short-hispid-tomentellous.
Flowers 4.0 mm long. Bracts and bracteoles 1.5-3.0 mm long, linear, pilose, eglandu-
lar, persistent. Receptacle campanulate, hirsute on exterior, glabrous within except
at throat; pedicels 0.5-2.0 mm long, many with a solitary stipitate gland arising near
base. Calyx lobes acute, hispid-hirsute on exterior, puberulous to glabrous within.
Petals 5, glabrous. Stamens 4, unilateral, with a toothed portion of arc opposite them,
338 Flora Neotropica

the filaments glabrous, far exceeding calyx lobes. Style hirsute up to half its length.
Ovary inserted at mouth of receptacle, pilose. Fruit not seen.
TYPES.Goeldi MG 15086, Brazil, Para, fl (lectotype, MG; isolectotype, US;
fragment F); DuckeMG 15646, Brazil, Para, fl (paratypes, MG, RB, US).
DISTRIBUTION.
Non-flooded primary forest and secondary forest, in the basins of
the rivers Tapaj6z and Tocantins, Para, Brazil. Flowering June to November.
BRAZIL. Para: DuckeRB 18240 st (RB, S); Frdes23512 fl (IAN, NY); Krukoff1244 fl (MICH,
NY, S).

LOCALNAME.Canella de velho.
This species differs from H. burchelliiin the impressed venation and slightly
bullate upper surfaces of the leaves, and in the terminal insertion of the ovary.

66. Hirtella burchellii Britton, Bull. Torrey Club 17: Io. Jan. 15, 1890.
HirtellapulchraFritsch, Ann. Naturh. Mus. Wien 5: 9. Feb. 1890. Type. Pohl 2181, Brazil, Goiis
fl (holotype, W; isotypes, BR, F, M, NY).
Hirtella collina S. Moore, Trans. Linn. Soc. II. 4: 351. I895. Type. SpencerMoore 141, Brazil,
Mato Grosso, fl (holotype, BM; isotype, NY).
Hirtella spruceiBentham ex Hooker f. subsp. meridionalisHassler, Fedde Repert. Nov. Sp. 7: 376.
I909. Type. Hassler o0700, Paraguay, fl (isotypes, A, BM, K, NY, P, W).
Hirtella plumbeaPilger, Notizbl. Bot. Gart. Berlin 6: I39. 1914; Macbride, Fl. Peru 1077. I938.
Type. Ule 79, Brazil, Acre, fl (holotype B, lost; isotype fragment F; photos, F, US).

Tree to 6.0 m tall or shrub, the young branches pilose-tomentellous. Leaves

oblong to oblong-elliptic, coriaceous, 0.0-20.0 cm long, 2.5-5.0 cm broad, acuminate
at apex, with acumen 5.0-I1.o mm long, subcordate to rounded at base, hirsute or
glabrous with hirsute primary venation beneath, the palisade glands frequent towards
base; primary veins 10-15 pairs, prominulous above, prominent beneath; petioles
2.0-3.0 mm long, pilose, terete, eglandular. Stipules 6.o-Io.o mm long, linear,
persistent, eglandular, sparsely hirsute. Inflorescences terminal and axillary racemes
9.0-30.0 cm long, the rachis pilose-tomentellous. Bracts and bracteoles 2.0-5.0 mm
long, linear-lanceolate, hirsute, persistent, with sessile glands and often with a
glandular apex. Flowers ca 5.0 mm long. Receptacle campanulate, puberulous to
pilose-hirsuteon exterior, glabrous within except at throat; pedicels 1.0-3.0 mm long,
most with a single stipitate gland arising near base. Calyx lobes 5, sparsely hirsute on
exterior, sparselypuberulous within. Petals 5, white to purple, glabrous. Stamens 4-5,
unilateral, with short filamentous or toothed staminodes opposite them, the filaments
glabrous, far exceeding calyx lobes. Style hirsute up to one third its length. Ovary
inserted midway up receptacle or towards its base, pilose. Fruit ellipsoid; epicarp
smooth, glabrous; mesocarp thin, fleshy; endocarp thin, hard, bony, hirsute within.
TYPES. Rusby 1222, Bolivia, fl (holotype, NY; isotypes, BM, F, GH, K, LE,
MICH, P, US); Burchell6331, Brazil, Goias, fl (paratypes, GH, K, P); 6416 fl (para-
types, GH, K); 657I fl (paratypes, K, P).
DISTRIBUTION.
Gallery forest and forest on high ground in Goias and Mato
Grosso, Brazil, west to Bolivia. Flowering throughout the year.
BRAZIL. Amazonas: Pranceet al 7308 fl (INPA, NY); 7449 fl (INPA, NY); 7537 fl (INPA, NY).
Acre: Pranceet al 7983 fl (INPA, NY); 8i86 fi (INPA, NY). Mato Grosso: Hoehne1342 fl (SP); I4zz fl
(R); Irwin et al 17I45 fr (NY); 17347 fl (NY); I742I fl (NY); Lindman2977 fl (S); Malme 2494 fl (LD,
S); Pereira 450 fl (NY, RB 85663); Riedel II35 fl (OXF, US); Robert490 fl (BM); 49ob fl (K); H.
Smithsn fl (R 35571). Goias: Glaziou 21109 fl (BR, C, K, P, S); Dawson I5r30 fl (NY).

There are no significant differences to distinguish specimens bearing the various

names placed in synonymy under H. burchellii.
Hirtella 339

67. Hirtella mucronata Prance, sp nov Fig 55 D-F.

Arbor, ramulis juvenilibus tomentosis. Folia alternata, petiolata; petiolo 2.5-

4.0 mm longo, tereti, eglanduloso, tomentoso; laminae ellipticae, coriaceae, I3.0-
17.0 cm longae, 4.5-6.5 cm latae, apice mucronatae, 6.O-I I. cm longae, basi
rotundatae vel subcuneatae, subtus dense hirsutae; costa media supra glabra, leviter
prominente, subtus prominente tomentello-hirsuta; costis secundariis 13-15 jugis,
supra prominulis subtus prominentibus. Stipulae 5.0-7.0 mm longae, lineares,
persistentes, tomentellae, eglandulosae. Flores circa 5.6 cm longi, in racemis axillari-
bus terminalibusque 8.0-15.0 cm longis dispositi, rachi tomentosi. Bracteae bracteo-
laeque 1.5-3.5 mm longae, lanceolatae, persistentes, tomentosae, apicibus saepius
glandibus stipitatis terminantae. Receptaculum campanulatum, extus sparse
tomentellum, intus versus basim glabrum; pedicelli 1.0-2.0 mm longi saepius versus
basim cum glandulo unico stipitate ornati. Calycis-lobi acuti, extus tomentelli, intus
puberuli. Petala 5, glabra, purpurea vel azurea. Stamina 5, unilateralia; filamenta
glabra, longe exserta. Ovarium ad medium receptaculi insertum, pilosum. Stylus e
basi ovarii ortus, hirsutus. Fructus mihi ignoti.
TYPE. A. C. Smith2718, Guyana, Essequibo River near mouth of Onoro Creek,
fl (holotype, NY; isotypes, A, F, K, MAD, P, S, US).
DISTRIBUTION. Non-flooded forest in Guyana and the lower Rio Negro region of
Brazil.
BRAZIL. Amazonas: Manaus; Estrada dos Francezes, Mello INPA 2852 fl (IAN, INPA, NY);
Rio Negro, Igarape do Arara. Rodrigues5523 fl (INPA 14196).

This species is very closely related to H. burchellii,but differs in the much broader
leaves with mucronate apices, and in the more pubescent lower surface of the leaf.

68. Hirtella longipedicellata Prance, sp nov Fig 55 A-C.

Arbor vel frutex parva, ramulisjuvenilibus puberulis mox glabris. Folia alternata
petiolata; petiolo 1.0-2.5 mm longo, juvenili tomentoso, tereti, eglanduloso; laminae
orbiculares vel lato-ovatae, coriaceae, 2.0-7.0 cm longae, 2.3-4.7 cm latae, apice
saepius rotundatae vel retusae nonnunquam acutae vel in acuminem brevem con-
tractae, basi subcordatae vel rotundatae, subtus cum pilis appressispaucis instructae,
versus basim glandulosae; costa media supra prominula, subtus prominente, pubes-
cente; costis secundariis 6-9jugis, utrinque prominulis. Stipulae circa 2.0 mm longae,
lineares, caducae. Flores 5.0-6.o mm longi in racemis terminalibus axillaribusque
corymbosis 2.0-3.0 cm longis dispositi, rachi sparse tomentello-puberuli vel glabres-
centi. Bracteae bracteolaeque I.0-2.0 mm longae, lanceolatae, persistentes, pubes-
centes, eglandulosae vel cum paucibus glandibus stipitatis munitae. Receptaculum
campanulatum, extus glabrescens,intus versus basim glabrum; pedicelli 7.0-20.0 mm
longi, pedicelli inferiores longiores quam pedicelli superiores. Calycis-lobi acuti
eglandulosi, extus glabrescentes, intus griseo-puberuli. Petala 5, glabra, alba vel
rosea. Stamina 6, unilateralia cum staminodiis dentatis opposita; filamenta glabra,
longe exserta. Ovarium ad faucem receptaculi insertum, sparse pilosum vel glabrum.
Stylus e basi ovarii ortus, versus basim hirsutus. Drupa ellipsoidea; epicarpio laevi,
glabro; mesocarpio tenue, carnoso; endocarpio tenue, duro, osseo, intus hirsuto.
TYPE. Maguire, Cowan & Wurdack 30978, Venezuela, Terr. Amazonas, Cerro
Huachamacari, Rio Cunucunuma, fl (holotype, NY).
Wooded slopes and savanna margins. Collected in flowerJanuary
DISTRIBUTION.
to March.
340 Flora Neotropica

VENEZUELA. Terr. Amazonas: Cerro Huachamacari, Rio Cunucunuma, Maguire, Cowan &
Wurdack30847 fl (NY); 30526 fl (NY); Caiio Asisa, Rio Ventuari, Cowan& Wurdack31453 fl (NY);
Caiio Atabapure, Rio Manapiare, Maguire & Maguire35043 fl (NY).

This species may be distinguished by the orbicular leaves with rounded to retuse
apices, by the corymbose inflorescence, and the long pedicels. It is related to H.
brachystachya and H. gracilipes,but differs in the leaf shape, the bract glands, and in
the distinctly corymbose inflorescence.

69. Hirtella glandistipula Ducke, Arch. Jard. Bot. Rio de Janeiro 3: 265.
1922.

Tree to i5.0 m tall, the young branches hispid, becoming glabrous with age.
Leaves oblong to elliptic, chartaceous, 4.0-6.5 cm long, 1.4-2.6 cm broad, bluntly
acuminate at apex, with acumen 3.0-Io.o mm long, rounded to subcordate at base,
with a few stiff appressed hairs beneath; primary veins 8-9 pairs, plane to slightly
impressedabove, prominent beneath; midrib prominulous above, hispid when young,
prominent and sparsely hispid beneath; petioles I.0-2.0 mm long, terete, eglandular,
hispid. Stipules 2.0-5.0 mm long, linear, persistent, hispid, with many stipitate
glands. Inflorescences terminal and axillary racemes 1.5-3.0 cm long, the rachis
short-hirsute. Bracts and bracteoles I.0-2.0 mm long, ovate, acute, puberulous,
persistent, with many stipitate and some sessile glands. Flowers ca 5.0 mm long.
Receptacle campanulate, sparsely hispid-hirsute on exterior, glabrous within except
at throat; pedicels 2.0-5.0 mm long, eglandular. Calyx lobes acute, hispid on exterior,
gray-puberulouswithin. Petals 5, white, glabrous. Stamens 3-4, with short staminodes
opposite them, the filaments far exceeding calyx lobes, glabrous to sparselypubescent
at base. Style hirsute up to two thirds its length. Ovary inserted midway up receptacle,
pilose. Fruit not seen.
TYPE. Ducke RB 15057, Brazil, Para, fl (holotype, RB; isotypes, K, S, US).
DISTRIBUTION. Primary and secondary forest on non-flooded ground in Para and
Terr. Amapa, Brazil. Flowering August to October.
BRAZIL. Para: Ducke RB 19761 fl (RB); Oliveira 86z fl (IAN, UB); Prance & Silva 58679 fl
(NY, UB); 58864 fl (NY, UB); 58944 fl (NY, UB). Terr. Amapa: Black 49-8312 fl (IAN, NY); 49-
8453 fl (IAN, NY).

This species is easily distintuished by the smaller chartaceous leaves and the
glandular stipules. There are only two other species (H. excelsa& H. pimichina)with
the latter character.

70. Hirtella martiana Hooker f., Mart. Fl. Bras. 14(2): 29. 1867.

Tree to Io.o m tall, the young branches lanate, soon becoming glabrous and
lenticellate. Leaves ovate-elliptic to oblong, coriaceous, 5.0-10.0 cm long, 1.8-3.8 cm
broad, acute to acuminate at apex, with acumen 2.0-6.0 mm long, rounded to
cuneate at base, glabrous beneath, with palisade glands at base only; primary veins
7-10 pairs, prominulous on both surfaces; midrib prominulous above, prominent
beneath; petioles 3.o-5.0 mm long, terete, eglandular, pubescent when young.
Stipules 2.0-4.0 mm long, erect, lanceolate, puberulous when young, persistent,
Hirtella 341

re

.aI l' .... :1

- -.

E F J.-Loken

FIG 55. Species of Hirtella.A-C, H. longipedicellata(Cowanet al 31453, Maguireet al 30978);

A, habit, x o.5; B, flowersection, x 2; C, petal, x 2. D-F, H. mucronata(A. C. Smith27I8); D, habit,
o.5; E, flower
xXo.,; section, x
flower section, petal, x
3; F, petal,
X3; X3.
3.
342 Flora Neotropica

eglandular. Inflorescencesaxillary and terminal densely crowded racemes 3.0-7.0 cm

long, the rachis sparsely tomentellous. Bracts and bracteoles 2.5-5.5 mm long, oblong
to ovate, persistent, puberulous, the margins with numerous stipitate glands. Flowers
6.o-Io.o mm long. Receptacle campanulate, sparsely tomentellous on exterior,
glabrous within except at throat; pedicels 5.o-1o.o mm long. Calyx lobes acute,
sparsely tomentellous on exterior, tomentellous within, the margins with small
stipitate glands. Petals 5, white, glabrous. Stamens 7-9, unilateral, the filaments
glabrous, far exceeding calyx lobes. Style hirsute up to three fourths its length. Ovary
inserted at mouth of receptacle, pilose. Fruit (immature) rotund; epicarp densely
tomentose; mesocarp thin; endocarp thin, hirsute within.
TYPES. Gardner4540, Brazil, Minas Gerais, fl (lectotype, K; isolectotypes, BM,
CGE, F, G, GH, M, NY, OXF, P, US); 3138, Brazil, Minas Gerais fl (paratype, K);
Martiussn, Brazil, Minas Gerais or Bahia fl (paratype, M).
DISTRIBUTION. Gallery and swamp forest of the Planalto of Central Brazil.
Flowering June to February.
BRAZIL. Goias: Burchell7494 fl (GH, K, NY, OXF, P, US); Prance& Silva 58246 fl (NY, S, US);
St. Hilaire 1654 fl (P). Dist. Federal: Irwinet al 6200 fr (NY); 18039 fl (IAN, NY); I8043 fl (IAN, NY);
Ramos6719 fl (K, NY); Ule I57 fl (P, R). Bahia: Luetzelburg153 fl (M, NY); Ule 7274 fl (HBG, K).
Minas Gerais: CapanemaRB 19203 fl (RB); Glaziou 12657 fl (BR, C, F, IAN, K, LE, P, R, US); 14676
fl (C, K, LE, NY); Heringer9493 fl (UB); St. Hilaire 1783 fl (P).

This distinctive species is probably most closely related to the two East African
species of this genus (H. megacarpa & H. zanzibarica),differing mainly in the racemose
inflorescence.

71. Hirtella pimichina Lasser & Maguire, Bol. Soc. Venez. Cienc. Nat.
I5: Io3. I954-

Small tree, the young branches hispid, soon becoming glabrous. Leaves oblong
to broadly lanceolate, coriaceous, 2.5-7.0 cm long, 1.1-3.2 cm broad, acuminate
at apex, with acumen 3.0-10o. mm long, subcordate at base, with a few stiff appressed
hairs beneath, sparsely hispid on margins and midrib, the palisade glands frequent
towards base; primary veins 6-9 pairs, prominulous on both surfaces; midrib promi-
nulous above, prominent and hispid-hirsute beneath; petioles I.O-I.5 mm long,
hispid, terete, eglandular. Stipules ca 2.0 mm long, linear, hirsute, persistent, with
stipitate glands or eglandular. Inflorescences terminal and axillary few-flowered
racemes 3.0-5.0 cm long, the rachis hispid. Bracts I .0-2.5 mm long, oblong, persistent,
hispid, with stipitate glands. Flowers 4.0-5.0 mm long. Receptacle campanulate
sparsely puberulous and hispid on exterior, glabrous within except at throat; pedicel
7.0-10.0 mm long, eglandular. Calyx lobes acute, sparsely hispid on exterior, puber-
ulous within. Petals 5, glabrous. Stamens 4-6, unilateral, the filaments far exceeding
calyx lobes, glabrous. Style hirsute up to half its length. Ovary inserted midway up
receptacle, pilose. Fruit not seen.
TYPE. LI. Williams 14437, Venezuela, Terr. Amazonas, fl (holotype, VEN;
isotypes, G, US).
DISTRIBUTION. Forest margins of the upper Rio Negro. Flowering September to
February.
VENEZUELA. Terr. Amazonas: Foldats 3636 fl (VEN). BRAZIL. Amazonas: Frdes 22737 fl
(IAN, NY); 29118 fl (IAN).
Hirtella 343

72. Hirtella subscandens Spruce ex Hooker f., Mart. Fl. Bras. 14(2): 32.
1867.

Shrub, often with trailing branches, the young stems sparsely hispid, soon be-
coming glabrous. Leaves oblong, thick-coriaceous, 7.0-12.0 cm long, 2.I-4.9 cm
broad, acuminate at apex, with acumen 4.0-Io.o mm long, cordate at base, glabrous
beneath, the palisade glands rare; primary veins IO-I4 pairs, prominulous above,
prominent beneath; midrib prominulous above, prominent and glabrous beneath;
petioles 1.0-2.0 mm long, terete, eglandular, hispid when young. Stipules ca 4.0 mm
long, linear, tomentellous, subpersistent, eglandular. Inflorescences terminal and
axillary racemes I7.0-25.0 cm long, the rachis puberulous above, hispid towards
base. Bracts and bracteoles I.0-3.0 mm long, oblong to lanceolate, persistent,
puberulous, with stipitate glands. Flowers 5.o-6.o mm long. Receptacle campanulate,
puberulous on exterior, glabrous within except at throat; pedicels 7.0-I 1.0 mm long.
Calyx lobes rounded to acute, glabrescent on exterior, gray-puberulous within.
Petals 5, pink, glabrous. Stamens 6-7, unilateral with toothed staminodes opposite
them, the filaments far exceeding calyx lobes, glabrous. Style hirsute at base only.
Ovary inserted at mouth of receptacle, tomentose. Fruit not seen.
TYPE. Spruce3533, Venezuela, Terr. Amazonas, fl (holotype, K; isotypes, BM,
BR, CGE, GH, LD, LE, NY, OXF, P).
Open forest in Amazonian Venezuela. Flowering April to August.
DISTRIBUTION.
VENEZUELA. Terr. Amazonas: Maguire & Wurdack35598 fl (NY); 35608 fl (NY); Ll. Williams
I4940 fl (A, F, US, VEN).

73. Hirtella hispidula Miquel, Stirp. Surin. Sel. 28, t. 7b. I850; Kleinh. in
Pulle Fl. Suriname 2(I): 445. 1937 (sub H. strigulosa Steudel).
Hirtella egensisFritsch, Ann. Naturh. Mus. Wien 5: Io. 1890, synon nov. Type. Poeppig2501,
Brazil, Amazonas, fl (holotype, W; isotypes, F, NY, OXF).
Hirtella subsetosaFanshawe & Maguire, Bull. Torrey Club 75: 38I. I948, synon nov. Type.
Fanshawe& Maguire23437, Guyana, fl (holotype, NY; isotypes, A, F, K, US).

Tree to 15.0 m tall, usually smaller, the young branches hirsute-hispid,becoming

glabrous with age. Leaves elliptic to oblong, coriaceous, 5.5-I5.0 cm long, 2.2-5.0
cm broad, acuminate at apex, with acumen 5.o- 5.0 mm long, subcordate to rounded
at base, hirsute on principal venation beneath, the palisade glands occasional; primary
veins 8- 2 pairs, prominulous on both surfaces; midrib prominulous above, promi-
nent and hirsute beneath; petioles 1.5-3.0 mm long, terete, eglandular, hispid.
Stipules 3.0-9.0 mm long, linear, persistent, hispid, eglandular. Inflorescences ter-
minal and axillary racemes 6.5-28.0 cm long, the rachis hispidulous on lower portion.
Bracts and bracteoles 1.5-3.5 mm long, linear-lanceolate, persistent, tomentellous-
hispid, terminating at apex in a long thin stipitate gland, usually with several
stipitate glands on margins. Flowers 4.0-5.0 mm long. Receptacle campanulate,
sparsely pilose on exterior, glabrous within except at throat; pedicels 2.0-8.0 mm
long, eglandular. Calyx lobes acute, sparsely pilose to glabrescent on exterior,
sparsely puberulous within. Petals 5, pinkish-purple, glabrous. Stamens 4-5, uni-
lateral with toothed staminodes opposite them, the filaments far exceeding calyx
lobes, glabrous. Style hirsute up to one third its length. Ovary inserted at mouth of
receptacle, pilose-tomentose. Fruit ellipsoid; epicarp smooth, glabrous; mesocarp
thin, fleshy; endocarp thin, hard, smooth, opening longitudinally, hirsute within.
TYPE. A. Fockesn, Suriname, fl (holotype, U).
344 Flora Neotropica

DISTRIBUTION. Primary forest on high ground in the Guianas and Amazonia

to northern Bolivia.
VENEZUELA. Bolivar: Bernardi 6486 fl (NY). Terr. Amazonas: Maguire & Politi 28621 fl
(NY). GUYANA. CAP 93(F.D. 6478) fl (K, NY, U); Maguire32092 fl (NY); 32109 fr (NY). SURI-
NAME. B.W. 6303 fl (K, NY, U); HostmannI75 fl (K, NY, OXF, P, S); Irwin, Prance,Soderstrom&
Holmgren55169 fl (NY, US); 55683 fl (F, GH, NY, S, US); 57592 fl (GH, NY, US); Lanjouw& Linde-
man 2949 fl (IAN, K, NY, U); 3377 fl (IAN, K, NY, U); Van Donselaar1947 st (U); 3035 st (U).
FRENCH GUIANA. LemoineF.D. 7757 fl (MG, NY, P); Melinon 240 fl (NY, P); Rohr sn fl (LE);
Sagot 1382 fl (K, S); Herb. Vahlsn fl (C). BRAZIL. Amazonas: ChagasINPA 1311 fl (IAN, INPA, NY);
Duarte7130 fl (INPA); DuckeRB 35579 fl (RB); Krukoff4684 fl (A, BM, F, LE, MICH, NY, S, US);
Maguire,Steyermark& Maguire 60076 fl (NY); Riedel I375 fl (US); Rodrigues538 fl (IAN, INPA 5732,
MG, NY). Pari: Capucho587 fl (F, IAN); DuckeMG 8374 fl(MG); Sampaio5I33 fl (R). Terr. Rond6nia:
Cordeiro& Silva 169 fl (IAN, NY); Pranceet al 5884 fl (INPA, NY); 6290 fl (INPA, NY); 6744 fl
(INPA, NY). BOLIVIA. Bang 1468 fl (A, F, FI, GH, LE, MICH, NY, S, US); Krukoffo1558 fl (A,
F, FI, K, MICH, NY, S, SP, US); 10884 fr (A, F, K, MICH, NY, S); Pranceet al 5769 fl (INPA, NY);
8427 fl (INPA, NY).

LOCAL NAMES.Suriname: Kleinbladige Anaura. French Guiana: Akianteifi-koko

(Taki-taki).
This species is most closely related to H. pilosissima,but differs in the long-
stipitate glands of the bracteoles, the almost glabrous leaf margins, and in the shorter
petioles. Hirtellahispiduladiffers from H. racemosa in the stipitate bract glands, and in
the pilose-hispid inflorescence and young branches.
Much of the Guiana material cited has been distributed as H. strigulosa,Hirtella
hispidulahaving been listed as a synonym of H. strigulosaby Kleinhoonte (in Pulle,
Fl. Suriname 2(I): 445. I939). However, the type of H. strigulosadiffers from H.
hispidula,as defined here, in the sessile bracteal glands, the larger petioles, and in the
sparsely hispid young branches. In all respects, material distinguished as H. strigulosa,
fits well into H. racemosa,and must be considered as part of that species, necessitating
in turn, resurrection of the name H. hispidulafor the present species. The confusion
between H. hispidulaand H. strigulosawas further complicated in that one collection
of H. hispidula,HostmannI75, was incorrectly labelled as the type of H. strigulosa.

74. Hirtella silicea Grisebach, Fl. Brit. W. Ind. 229. I864.

Hirtella melinoniiR. Benoist, Bull. Mus. Hist. Nat. Paris 29: 596. 1923. Type. MelinonI32, French
Guiana, fl (holotype, P; isotypes, NY, US).
Hirtella manigeraKleinhoonte, Rec. Trav. Bot. N6erl, 22: 387. 1925; in Pulle, Fl. Suriname
2(I): 446. I937, synon nov. Types. SammlerHerb. 117, Suriname, fl fr (U); Gonggrijp
& StahelB.W. 905, Suriname, fl bud (U).

Tree to Io.o m tall or shrub, the young branches tomentellous, becoming glab-
rous with age. Leaves oblong, chartaceous to thin-coriaceous, IO.0-22.0 cm long,
3.0-9.0 cm broad, acuminate at apex, with acumen 6.0-i5.0 cm long, subcordate to
rounded at base, sparsely appressed pubescent beneath, especially on venation, with
palisade glands at base only; primary veins 8-12 pairs, prominulous above, promi-
nent beneath; midrib prominulous and tomentellous above when young, prominent
beneath; petioles 1.5-3.5 mm long, terete, eglandular, tomentellous. Stipules 2.0-
3.0 mm long, linear, filamentous, persistent, tomentellous, eglandular. Inflorescences
terminal and axillary racemes 4.5-27.0 cm long, the rachis tomentellous. Bracts
2.0-6.0 mm long, oblong, tomentellous, persistent, with few to numerous stipitate
glands. Flowers ca 5.0 mm long, Receptacle campanulate, sparsely hirsute-tomen-
tellous on exterior, glabrous within except at throat; pedicels 4.0-9.0 mm long,
eglandular. Calyx lobes acute, sparsely tomentellous on exterior, gray-puberulous
Hirtella 345

within. Petals 5, glabrous, white to pink. Stamens 5-6, unilateral, with short filamen-
tous staminodes opposite them, the filaments glabrous, far exceeding calyx lobes.
Style hirsute up to one third its length. Ovary inserted at mouth of receptacle. Fruit
ellipsoid; epicarp smooth, glabrous; mesocarp thin, fleshy; endocarp thin, hard,
bony, hirsute within.
TYPE. Cruegersn, Trinidad, fl (K; fragment NY).
DISTRIBUTION. Forest on high ground in Venezuela, Trinidad and the Guianas.
Flowering throughout the year.
VENEZUELA. Sucre: Broadway565 fl (GH, NY, US); Steyermark& Rabe96100 fl (NY, VEN).
Terr. Delta Amacuro: Blanco 199 fl (NY, VEN); Steyermark87554 fr (NY). Bolivar: BernardiI4f6 fl
(NY); Steyermark 76039 fl (F, NY, VEN); 86384 fl (VEN). TOBAGO. Broadway3577 fl (BM, F, NY,
S, US); 4025 fl (BM, C, F, G, GH, K, NY, S, US); Cowan1477 fl (NY, US); Eggers5818 fl fr (A, K, P,
US); Sandwith 1721 fl (K); Websteret al 9810 fl (A, S, US); 9866 fl (S). TRINIDAD. Britton et al
1329 fl (GH, K, NY, US); Broadway6905 fl (F, K); Brooks12481 St (K); Fendler1020 fl (BM, K, NY,
P); Prance2108 fr (NY); A. C. Smith 10036 fl (A, K, NY, US). GUYANA. J. Boyan 69(F.D. 7753) fl
(K, NY); Fanshaweiii9(F.D. 3855) fl (K, NY); Jenman3980 fl (BM, F, K); Maguire & Cowan39298 fr
(K, NY, US); Maguire & Fanshawe22924 fl (NY); 23514 fl (A, F, K, NY, US); Sandwith92 fl (K, NY,
P, RB); Tillett & Tillett 45044 fl (NY); Ward8646 fl (K). SURINAME. Cowan & Lindeman39023 fl
(F, NY, P); Daniels & Jonker 784 fl (NY, U); Irwin, Prance,Soderstrom& Holmgren54594 fl (NY);
54889 fr (F, NY, US); Schulz 7500 st (NY, U); Van Donselaar 2260 St (U); 3121 st (U).

LOCAL NAMES. Venezuela: Candelita, Hija, NJaranjillo. Guyana: Bokobokotokon.

Suriname: Tijantesie.

As there are no differences among species labelled H. silicea, H. melinonii,and

H. manigera,the two latter species are herewith reduced to synonymy. The identity
of H. melinoniihas been somewhat confused in the past because Melinon collections
of two different species have been distributed under the same name. The holotype
and the original description of H. melinoniimatch H. silicea.However, some material
marked as the isotype of H. melinoniiat Kew and Paris belongs to another species
which is described here as H. tenuifolia.
Hirtellasilicea may be distinguished from related species by the large stipitate
glands, the large leaves, and the long inflorescence.

75. Hirtella excelsa Standley ex Prance, sp nov Fig 56 D-G.

Arbor magna vel media, ramulis juvenilibus glabrescentibus. Folia alternata

petiolata; petiolo 1.5-2.5 mm longo, tereti, eglanduloso, glabro; laminae oblongae
vel oblongo-ellipticae, 6.o-Io.o cm longae, 2.3-3.5 cm latae, coriaceae, apice in
acuminem 5.0-I5.0 mm longum contractae, basi cuneatae vel subcuneatae, subtus
venis cum pilis appressispaucibus instructae, versus basim glandulosae; costa media
supra prominula, glabrescente, costis secundariis 9-10 jugis, utrinque prominulis.
Stipulae circa 3.0 mm longae, lineares, eglandulosae, subpersistentes,glabrae. Flores
circa 6.o mm longi in racemis axillaribus terminalibusque 7.9-I5.0 mm longis dis-
positi, rachi juvenili sparse puberuli mox glabri. Bracteae bracteolaeque 1.5-3.5 mm
longae, oblongae vel ellipticae, membranaceae, persistentes, sparse tomentellae, cum
glandulis stipitatis numerosis munitae. Receptaculum campanulatum, extus sparse
puberulum, intus versus basim glabrum; pedicelli 7.0-10.0 mm longi,eglandulosi.
Calycis-lobi acuti, extus glabrescentes, intus griseo-puberuli, eglandulosi. Petala 5,
glabra. Stamina 3, unilateralia; filamenta glabra, longe exserta. Ovarium ad faucem
receptaculi insertum, pilosum. Stylus e basi ovarii ortus, hirsutus. Drupa ellipsoidea;
epicarpio laevi, juvenili sparse pubescenti mox glabri; mesocarpio tenue, carnoso;
endocarpio tenue, osseo, intus hirsuto.
346 Flora Neotropica

TYPE. Majin 58, Peru, Loreto, Calleria, Pcia Cnel., Portillo, fl (holotype, F;
isotype, NY, WIS).
DISTRIBUTION. Non-flooded forest in western Amazonia. PERU. Loreto:
Distrito Calleria, Pcia Cnel., Portillo, Majin72, fr (F, NY).
BRAZIL. Amazonas: Near mouth of Rio Embira; Krukoff4786fr (A, BM, F, FI, K, MICH, NY,
S, US). Acre: Near mouth of Rio Macauhan; Krukoff5491fr (A, F, K, LE, MICH, NY, SP, US).

LOCALNAME.Peru: Chuchuhuasha-masha.
Standley saw only the Krukoff material which is in the immature fruiting con-
dition. This material has been widely distributed under Standley's manuscript name
which is now validated here. Hirtellaexcelsais distinct and not easily confused.

76. Hirtella adenophora Cuatrecasas, Brittonia 8: 196. 1956.

Small tree, the young branches puberulous, soon becoming glabrous. Leaves
ovate-lanceolate, coriaceous, broadest below midpoint, 6.0-I2.0 cm long, 1.6-4.2 cm
broad, gradually tapering to an acute or acuminate apex, rounded to subcordate at
base, glabrous beneath except on midrib, with palisade glands towards base; primary
veins 6-9 pairs, prominulous and glabrous on both surfaces; midrib prominulous
above, prominent and puberulous beneath; petioles 2.o-4.o mm long, terete,
eglandular, glabrous or with a few stiff erect hairs. Stipules ca 2.0 mm long, linear,
subpersistent, glabrescent, with stipitate glands. Inflorescences terminal and axillary
racemes 6.0-I2.0 cm long, the rachis sparsely puberulous. Bracts and bracteoles
i.5-2.5 mm long, oblong, peristent, puberulous, with stipitate glands on apex and
margins. Flowers 5.o-6.o mm long. Receptacle campanulate, sparsely puberulous
on exterior, glabrous within except at throat; pedicels 3.0-5.0 mm long, eglandular.
Petals 5, pink, glabrous. Calyx lobes acute, sparsely puberulous on exterior, gray-
puberulous within, Stamens 5-6, unilateral with toothed staminodes opposite them,
the filaments glabrous, far exceeding calyx lobes. Style hirsute on lower portion.
Ovary inserted at mouth of receptacle, pilose. Fruit ellipsoid; epicarp smooth,
glabrous when mature, puberulous when young; mesocarp thin, fleshy; endocarp
hard, bony, hirsute within.
TYPE. Philipson& IdroboI887, Colombia, Meta, fl (holotype, US; isotypes, BM,
COL, NY).
DISTRIBUTION. Known only in scrubby montane forest of Meta, Colombia.
Flowering December to February.
COLOMBIA. Meta: Philipson & IdroboI969 fl (BM, COL, NY, US); Philipson2309 fr (BM).

may be readily distinguished from related species by the leaf

Hirtellaadenophora
shape.

77. Hirtella caduca Fanshawe & Maguire, Bull. Torrey Club 75: 379. 1948.
Tree to 8.o m tall, the young branches sparsely hispid, soon becoming glabrous.
Leaves oblong to oblong-elliptic, chartaceous, 4.5-8.5 cm long, 1.5-3.5 cm broad,
cuspidate to acuminate at apex, with acumen 7.0-18.0 mm long, subcordate to
rounded at base, glabrous beneath except on primary venation, with palisade glands
present towards base; primary veins 5-8 pairs, prominulous on both surfaces; midrib
 Hirtella 347

C'

?
\

J Loken E

FIG 56. Species of Hirtella. A-C, H. rugosa(Gleasonet al H-73; Garber82); A, habit, x 0.5; B
flower section, x 2.5; C, petal, 2.5; C', young fruit, x i. D-G, H. excelsa(Krukoff549, Majin 58); D
habit, x 0.5; E, flower section, x 3; F, petal, x 3; G, inflorescence, x 0.5.
348 Flora Neotropica

prominulous above, prominent and appressed-pubescent beneath; petioles I.O-3.0

mm long, terete, eglandular, sparsely hispid to puberulous, soon becoming glabrous.
Stipules 2.0-5.0 mm long, ovate-triangular, caducous, puberulous with glandular
margins. Inflorescences terminal and axillary racemes 3.0-6.0 cm long, the rachis
sparsely strigose to puberulous. Bracts and bracteoles 0.5-2.0 mm long, oblong,
persistent, puberulous, with stipitate glands on margins and apex. Flowers ca 5.0 mm
long. Receptacle campanulate, with a few stiff appressed hairs on exterior, glabrous
within except at throat; pedicels 8.0-15.0 mm long, eglandular. Calyx lobes acute,
reflexed, glabrescent on exterior, gray-puberulous within, the margins eglandular.
Petals 5, white, glabrous. Stamens 6, unilateral, the filaments glabrous, far exceeding
calyx lobes. Style hirsute up to half its length. Ovary inserted at mouth of receptacle,
pilose. Fruit ellipsoid; epicarp smooth, glabrous; mesocarp thin, fleshy; endocarp
thin, hard, bony, hirsute within.
TYPES.Maguire& Fanshawe23498, Guyana, fl (holotype, NY; isotypes, A, BR,
F, K, LE, M, MO); 23005a fl (paratypes, A, NY, US).
DISTRIBUTION. Forest in open places, such as savanna and river margins, in
Guyana and adjacent Venezuela. Flowering April to August.
VENEZUELA. Bolivar: Maguire, Steyermark& Maguire 46714 fl (NY). GUYANA. R. Boyan30
2234 fl (NY, US); Tillett, Tillett & Boyan45241 fl (NY).
(F.D. 7854) fl (NY); Cowan& Soderstrom

LOCALNAME.Guyana: Bokobokotokon.
but differs in the stipitate
This species is close to H. gracilipesand H. brachystachya,
glands of the bracts, in the absence of translucent glandular excretions, and in the
cuspidate leaves.

78. Hirtella fasciculata Prance, sp nov Fig 57 A-C.

Arbor media vel magna, ramulis juvenilibus tomentellis. Folia alternata,

petiolata; petiolo 2.0-3.0 mm longo, tereti, eglanduloso, tomentoso; laminae ellipti-
cae, coriaceae, 9.0-I2.5 cm longae, 5.o-6.5 cm latae, apice in acuminem abruptum
I.0-4.0 mm longum contractae, basi rotundatae, subtus hirsutae versus basim glandu-
losae, supra leviter bullatae; costa media supra prominula, subtus prominente,
hirsuta; costis secundariis 8-Io jugis, supra impressis, subtus prominentibus. Stipulae
circa 2.5 mm longae, lanceolatae, caducae, tomentosae, eglandulosae. Flores 6.0-
7.0 mm longi, in racemis fasciculatis axillaribus I.0-2.0 cm longis dispositi, rachi
tomentelli. Bracteae 2.0-3.5 mm longae, lineares, tomentellae, persistentes, eglan-
dulosae. Receptaculum campanulatum, extus hirsuto-tomentellum, intus glabrum;
pedicelli 1.0-2.0 mm longi, eglandulosi. Calycis-lobi acuti, extus tomentelli, intus
puberuli. Petala 5, glabra. Stamina 4, unilateralia; filamenta glabra, longe exserta.
Ovarium ad medium vel versus basim receptaculi insertum, pilosum, Stylus e basi
ovarii ortus, hirsutus. Fructus mihi ignoti.
TYPE.Rodrigues& Loureiro5947, Brazil, Amazonas, Manaus, Reserva Florestal
Ducke, fl (holotype, NY; isotype, INPA 1489I).
DISTRIBUTION. Non-flooded forest in Central Amazonian Brazil. Collected in
flower in July and September.
BRAZIL. Para: M. Silva & Souza 2530 fl (MG, NY).

This species may be distinguished by the short fasiculate inflorescence and the
slightly bullate leaves. It is not very closely related to any other species.
Hirtella 349

E D JiLokent

FIG 57. Species of Hirtella. A-C, H. fasciculata (Rodrigueset al 5947); A, habit, x 0.5; B, flower
section, x 2; C, petal, X 2. D-F, H. couepiflora(Irwinet al 47367); D, habit, x o.5; E, petal, x 2; F, flower
section x 2.
350 Flora Neotropica

79. Hirtella couepiflora Prance, sp nov Fig 57 D-F.

Arbor parva, ramulis juvenilibus tomentellis mox glabris lenticellatis. Folia

alternata, petiolata; petiolo 2.5-4.0 mm longo, tereti, eglanduloso, tomentello;
laminae oblongo-ellipticae, coriaceae, 10.0-14.5 cm longae, 4.o-6.5 cm latae, apice
in acuminem 5.0-10.0 mm longum contractae, basi subcuneatae vel cuneatae, subtus
venis cum pilis appressissparse munitae; costa media supra leviter prominula, subtus
prominente; costis secundariis 13-15 jugis supra planis, subtus prominentibus.
Stipulae circa 3.5 mm longae, lanceolatae, puberulae, subpersistentes,eglandulosae.
Flores I7.0-20.0 cm longi, in racemis terminalibus subterminalibusque 2.5-I I.0 cm
longis dispositi, rachi tomentelli. Bracteae bracteolaeque 2.0-6.5 mm longae, oblon-
gae vel lanceolatae, tomentellae, persistentes, eglandulosae vel cum paucis glandibus
stipitatis instructae. Receptaculum elongato-cylindricum, sessile, extus tomentellum,
intus glabrum. Calycis-lobi acuti, extus tomentelli, intus puberuli, eglandulosi vel
cum glandibus stipitatis muniti. Petala 5, alba, glabra. Stamina 8-9, unilateralia;
filamenta tomentosa, longe exserta. Ovarium ad faucem receptaculi insertum,
tomentosum. Stylus e basi ovarii ortus, versus basim hirsutus. Fructus mihi ignoti.
TYPE. Irwin, Egler & Westra 47367, Brazil, Terr. Amapa, Rio Oiapoque, near
Clevelandia, fl (holotype, NY; isotypes, IAN, MG).
DISTRIBUTION. Known only from primary forest in the Rio Oiapoque region.
Collected in flower July and August.
BRAZIL. Terr. AmapA: Rio Oiapoque, near Clevelandia, Maguire, Pires & Maguire 47073 fl
(IAN, MG, NY).

In this species, not easily confused with any others, the flowers have a long
tubular receptacle, unique except in H. tubiflora,which differs in many other charac-
ters. Superficially the general appearance of the present species suggests Couepia, but
the fewer stamens, the calyx glands, and the hirsute pubescence of the leaf venation
all place it in Hirtella.

80. Hirtella tubiflora Cuatrecasas, Fieldiana Bot. 27: 59. 1950.

Large tree, the young branches tomentellous, soon becoming glabrous. Leaves
elliptic-lanceolate, chartaceous, 4.0-8.6 cm long, 1.7-3.0 cm broad, acuminate at
apex, with acumen 6.o-15.o mm long, rounded to subcuneate at base, sparsely
appressed pubescent beneath, with palisade glands; primary veins 10-12 pairs,
prominulous on both surfaces, midrib prominulous above, prominent beneath;
petioles 1.5-2.0 mm long, terete, eglandular, appressed-puberulous when young.
Stipules ca 1.5 mm long, ovate, pubescent, eglandular, caducous. Inflorescences
terminal and axillary racemes 4.0-7.0 cm long, the rachis puberulous. Bracts and
bracteoles I.0-3.0 mm long, lanceolate, acute, puberulous, persistent, eglandular.
Flowers 7.0-9.0 mm long, Receptacle cylindrical, subsessile, elongate, puberulous on
exterior, glabrous within. Calyx lobes acute, puberulous on both surfaces, with
numerous small glands on margins. Petals 5, glabrous, white. Stamens 6, unilateral,
the filaments far exceeding calyx lobes, glabrous, Style hirsute up to three fourths its
length. Ovary inserted at mouth of receptacle, tomentose. Fruit ellipsoid; epicarp
smooth, glabrous; mesocarp thin, fleshy; endocarp thin, hard, bony, sparsely hirsute
within.
TYPE. Cuatrecasas21183, Colombia, Valle, fl (holotype, F; isotypes, COL, P).
DISTRIBUTION. Known only from the type gathering, collected in the Pacific
coastal forest of Colombia.
Hirtella 351

This species is easily distinguished by the elongate-cylindrical receptacle. It

differs from H. couepiflora,
also with this type of receptacle, in the smaller flowers, in
the sparsely puberulous exterior of the receptacle and calyx lobes, in the numerous
small glands on the calyx lobes, and in the smaller number of stamens.

8I. Hirtella floribunda Chamisso & Schlechtendal, Linnaea 2: 548. I827;

Zucc., Flora 15(Beibl. 2); 8i. 1832; Hook. f., Mart. Fl. Bras. 14(2): 32.
1867.

Shrub or small tree, the young branches tomentellous. Leaves oblong, 3.4-8.0
cm long, 1. -3.2 cm broad, acute to acuminate at apex, with acumen 0-5.0 mm long,
cuneate to subcuneate at base, hirsute beneath, the palisade glands absent; primary
veins 8-I o pairs, impressed above, prominent beneath; midrib impressed and pubes-
cent above, prominent and hirsute beneath; petioles 2.0-3.0 mm long, hispid-setose,
terete, eglandular. Stipules 2.0-3.0 mm long, linear, hirsute, persistent, eglandular.
Inflorescences terminal and axillary racemes 3.o-9.0 cm long, the rachis hirsute.
Bracts 0.7-I.5 mm long, oblong-lanceolate, persistent, tomentose, eglandular.
Flowers 4.0-5.0 mm long. Receptacle campanulate, sparsely hirsute and puberulous
on exterior, glabrous within except at throat; pedicels 8.o-I2.0 mm long, eglandular.
Calyx lobes acute, eglandular, sparsely hirsute on exterior, gray-puberulous within.
Petals 5, glabrous. Stamens 7-9, unilateral, inserted in a semicircle, the filaments
glabrous, far exceeding calyx lobes. Style hirsute up to one fourth its length. Ovary
inserted at mouth of receptacle, pilose. Fruit not seen.
TYPE. Sellowsn, Brazil, fl (holotype, LE; isotypes, BR, CGE, K, LE, M).
DISTRIBUTION. Minas Gerais in Brazil. None of the collections examined is
supported with details about the habitat.
BRAZIL. Minas Gerais: Ackermann sn fl (BR); Badini 3236 fl (RB); Claussensn fl (K, P); Io98 fl
(P); Gardner4541 fl (BM, F, G, GH, K, NY, P, US); Glaziou 19138 fl (C, P); Motta R 7145 fl (R);
8722 fl (R). Withoutprecise locality: Chamisso
Schwacke sn fl (K); Claussen
I47 fl (P); Gardner
4501 fl
(P).

82. Hirtella angustifolia Schott ex Sprengel, Linn. Syst. Veg. ed. i6. 4(Cur
Post): 341. 1827; Zucc., Flora 15(Beibl. 2): 83. I832; Hook. f., Mart. Fl.
Bras. 14(2): 30. 1867. Fig 50 A-C.

Shrub, the young branches hispid-setose. Leaves lanceolate to oblong-lanceolate,

coriaceous, 5.0-I I.5 cm long, 1.6-3.8 cm broad, acuminate at apex, with acumen 5.0-
15.0 mm long, subcordate at base, hirsute beneath, the palisade glands rare, at base
only; primaryveins 9- I pairs,plane above, prominulousbeneath; midribprominulous
and pubescent above when young, prominent and hirsute beneath; petioles 1.5-
3.0 mm long, terete, eglandular, hispid. Stipules 5.0-7.0 mm long, linear, hispid,
persistent, eglandular. Inflorescences terminal and axillary racemes 9.0-I5.0 cm
long, the rachis hispid-hirsute. Bracts 1.0-3.0 mm long, oblong to ovate, persistent,
pubescent, eglandular. Flowers 5.0-6.0 mm long. Receptacle campanulate, puberu-
lous and hirsute on exterior, glabrous within except for retrorse hairs at throat; pedi-
cels 8.0-I5.0 mm long, eglandular but bracteate. Calyx lobes acute, sparsely puberu-
lous on exterior, densely gray-puberulous within, margins eglandular. Petals 5, glab-
rous. Stamens 7-8, inserted in an arc of ca 270?, the filaments glabrous, far exceeding
352 Flora Neotropica

calyx lobes. Style hirsute up to half its length. Ovary inserted at mouth of receptacle,
pilose. Fruit not seen.
TYPE. Schott4970, Brazil, Rio de Janeiro or Guanabara, fl (holotype, W; iso-
types, BR, F, NY, OXF).
DISTRIBUTION. Coastal forests of eastern Brazil, known only from the states of
Rio de Janeiro and Guanabara. Flowering September to November.
BRAZIL. Minas Gerais (Cult) Barreto831 fl (RB 138213). Rio de Janeiro: Ducke & Kuhlmann
RB 19175 fl (RB, S, US); Glaziou I94I fl (BR, C, F, LE, P, R); 12658 fl (A, C, LE, P); Netto, Glaziou
& SchwackeR 7149 fl (NY, R); Pohl sn fl (BR, W). Without locality, Rio de Janeiro or Guanabara:
Gaudichaud1022 fl (F, G, P); Riedelsn fl (US).

83. Hirtella rugosa Thuillier ex Persoon, Syn. P1. I: 250. I805; Roem. &
Schult., Linn. Syst. Veg. ed. 9. 5: 274. I819; DC., Prodr. 2: 528. 1825;
Zucc., Flora I5(Beibl. 2): 82. I832; Urb., Symb. Antill. 4:26I. I905;
Britton & Wilson, Sci. Survey Porto Rico 5(3): 346. 1924. Fig 56 A-C.
Hirtella portoricensisHumboldt & Bonpland ex Roemer & Schultes, Linn., Syst. Veg. ed. 9.
5: 274. 1819. Type. Humboldt& Bonplandsn, Puerto Rico, fl (holotype, B, Herb. Willdenow
4845; photo, NY; isotype, P).
Zamzelarugosa(Thuillier ex Persoon) Rafinesque, Sylva Tellur. go. 1838.
Chrysobalanus rugosus(Thuillier ex Persoon) G6mez de la Maza, Dicc. Bot. Nom. Vulg. Cuba &
Puerto Riquefios Io. I889.

Tree to 9.0 m tall, or shrub, the young branches hispid-tomentellous, becoming

glabrous with age. Leaves ovate to oblong, thick-coriaceous, 2.5-IO.0 cm long, I.4-
5.0 cm broad, acuminate at apex, with acumen 5.0-17.0 mm long, subcordate to
rounded at base, sparsely hirsute beneath on principal venation, with palisade glands
absent, sometimes slightly bullate above; primary veins 7-10 pairs, impressed above,
prominent beneath; midrib slightly impressed above, prominent beneath; petioles
1.5-3.0 mm long, terete, eglandular, hispid when young. Stipules 6.0-9.0 mm long,
linear, persistent, pubescent, eglandular. Inflorescences short axillary and terminal
racemes 1.0-4.0 cm long, the rachis hispid when young. Bracts 1.5-3.5 mm long,
oblong to lanceolate, acute, tomentellous, persistent, eglandular. Flowers 6.0-7.0 mm
long. Receptacle campanulate, tomentellous on exterior, glabrous within except at
throat; pedicels 6.0-I I.Omm long, eglandular. Calyx lobes acute, tomentellous on
exterior, puberulous within. Petals 5, glabrous, pink. Stamens 3, unilateral, with
toothed staminodes opposite them, the filaments glabrous, far exceeding calyx lobes.
Style sparsely hirsute up to half its length. Ovary inserted at mouth of receptacle,
pilose. Fruit ellipsoid; epicarp smooth, puberulous when young, becoming glabrous
with age; mesocarp thin, fleshy; endocarp thin, hard, bony, hirsute within.
TYPE. Herb. Moquin-Tandon sn, Puerto Rico, fl (holotype, P).
DISTRIBUTION.Endemic to the montane forestsof Puerto Rico. Flowering through-
out the year, but mostly from February to July.
Representative collections: PUERTO RICO: Alain 9055 fl (NY, US); Britton et al 2284 fl fr
(F, GH, NY, S, US); 2435 fr (C, F, GH, NY, US); 7274 fr (NY); Eggers 849 fl (BR, F, GH, GOET,
HBG, LE, P, R); 1183 fl (C, US); Garber82 fl fr (GH, K, NY); Gleason& CookH 73 fl (NY); Heller
730 fl (F, K, NY, US); Home & Britton9724 fl (NY); Johnston850 fl (NY, US); Le Dru sn fl (G-DC,
P-LAM); Little 13093 fl fr (F); 13420 fl (F); Otero595 fl (A, F); 721 fl (A, NY); Riedlesn fl (P-JUSS);
SargentB 93 fl (US); Shafer3559 fl (F, NY, US); 3563 fl fr (F, GH, NY, US); Sintenis201b fl (GH,
GOET, K LD, LE, S, US); 232 fl (A, BM, F, GH, K, NY, P, US); o059 fl (BR, K, LE, US); 5256 fl
(GOET, K, LD, LE, US); 6067 fl (BM, GH, LD, NY, P, S); Vilez Io85 fl (NY); Io8 fl (NY); Wagner
231 fl (BM, LE); 325 st (BM, LE).

LOCALNAMES.Icacillo, Teta de burracimarrdn.

Hirtella 353

Although Persoon did not give any details of the type specimen in his description,
of the several early collections of H. rugosaat Paris in the various historical herbaria,
the specimen given here as the type is cited because it bears Thuillier's manuscript
writing. The Leiden herbarium also has a sheet without collectors name bearing the
inscription "Herb. Pers." This is probably an isotype of this species.
Hirtella rugosais quite distinct from other species of Hirtellawith a racemose,
eglandular inflorescence by the small leaves with impressed venation which are
broadest near to the base, the short densely crowded inflorescence, and the cordate
leaf bases.

84. Hirtella scaberula Spruce ex Hooker f., Mart. Fl. Bras. 14(2): 33. 1867.

Small tree, the young branches tomentellous, soon becoming glabrous. Leaves
ovate-elliptic, 6.0-9.0 cm long, 3.5-5.5 cm broad, abruptly acuminate at apex, with
acumen 3.o-5.0 mm long, rounded to subcuneate at base, sparsely appressed-
pubescent beneath, the palisade glands at base only; primary veins 7-9 pairs, plane
to prominulous above, prominent beneath; midrib prominulous above, prominent
and pubescent beneath; petioles 3.0 mm long, terete, pubescent, eglandular. Stipules
linear, persistent, eglandular, pubescent. Inflorescences terminal and axillary race-
mes, the rachis sparsely tomentellous. Bracts 1.0-2.0 mm long, oblong to lanceolate,
acute, puberulous, persistent, eglandular. Flowers 3.5-4.0 mm long. Receptacle
campanulate, sparsely tomentellous on exterior, glabrous within except at throat;
pedicels 2.0-3.0 mm long, eglandular. Calyx lobes acute, sparsely tomentellous on
exterior, glabrescent within except at margins. Petals 5, glabrous. Stamens 5-7, unila-
teral, the filaments glabrous, far exceeding calyx lobes. Style hirsute up to one third
its length. Ovary inserted at mouth of receptcle, pilose. Fruit not seen.
TYPE.Spruce2418 (2481 some sets), Brazil, Amazonas, fl (holotype, K; isotypes,
BM, BR, CGE, OXF, P).
DISTRIBUTION. Known only from the type gathering, collected in the Caatinga
forest near Panure on the Rio Vaupes.

85. Hirtella hebeclada Moricand ex A. P. de Candolle, Prodr. 2: 529. I825;

Zucc., Flora I5(Beibl. 2): 83. 1832; Hook. f., Mart. Fl. Bras. 14(2): 28.
1867.
Hirtella gardneriBentham, Jour. Bot. Hooker 2: 216. 1840. Type. Gardner370, Brazil, Rio de
Janeiro, fl (holotype,K; isotypes,BM, CGE, F, GH, NY, OXF, S, US).
Hirtella selloana Hooker f., Mart. Fl. Bras. 14(2): 29. 1867. Type. Sellow 2043, Brazil, Minas
Gerais,fl (holotypeB, lost; isotypefragments,K, P).
Schott ex Sprengel,Linn. Syst, Veg, ed. I6. 4(Cur. Post.): 34I. 1827. Type.
Hirtellatomentella
Schottsn Brazil,fl (holotypeB, lost; photos,F, GH, NY).

Tree to 15.0 m tall, the young branches tomentose, becoming glabrous with age.
Leaves ovate to elliptic, thick-coriaceous, 5.5-I6.0(-2I.o) cm long, 2.5-9.0 cm broad,
acute to acuminate at apex, with acumen 0-12.0 mm long, rounded to subcuneate at
base, hirsute beneath, rarely with two palisade glands at base; primary veins 8-I3
pairs, usually slightly impressed above, less often plane; prominent beneath; midrib
prominulous and tomentellous above, prominent beneath; petioles 3.0-6.0 mm long,
terete, eglandular, tomentose when young. Stipules 3.0-6.0 mm long, linear, sub-
354 Flora Neotropica

persistent, tomentellous, eglandular. Inflorescences terminal and axillary racemes,

or rarely slightly branched panicles, 3.o-15.0 cm long, the rachis tomentose to
tomentellous. Bracts 1.0-3.0 mm long, oblong to lanceolate, acute, persistent,
tomentellous, eglandular. Flowers 5.o-8.o mm long. Receptacle campanulate,
tomentose to tomentellous on exterior, glabrous within except at throat; pedicels
5.0-15.0 mm long, eglandular. Calyx lobes acute, tomentellous on both surfaces,
the margins eglandular. Petals 5, white, glabrous. Stamens 6, unilateral, often with
short filamentous staminodes opposite them, the filaments far exceeding calyx lobes,
glabrous. Style hirsute up to half its length. Ovary inserted at mouth of receptacle,
pilose. Fruit ellipsoid, tapering at base; epicarp smooth, glabrous when mature;
mesocarp thin, fleshy; endocarp thin, hard, bony, hirsute within.
TYPE.Sellowsn (in Herb. Moricand), Brazil, fl (holotype, G; isotypes, L, M).
DISTRIBUTION. Primary forest in eastern and southern Brazil from Minas Gerais
to Rio Grande do Sul. Flowering throughout the year, but mostly from November to
February.
BRAZIL. Minas Gerais: Barreto4759 fr (F, R); Gomes2482 fl (NY, RB 97678); KuhlmannRB
138215 fl (RB); Mexia 5473 fl (A, BM, F, GH, K, LA, MICH, NY, S, US); Pohl sn fl (BR). Espirito
Santo: DolbolmoRB 86219 fl (RB). Sao Paulo: Bowie & Cunningham sn fl (BM); Brade 12125 fl (R);
Faria 14 fl (SP); Gaudichaud868 fl (P); GehrtSP 12887 fl (SP); HoehneSP 7546 fl (SP); SP 28586 fl (A,
F, GH, NY, P, S, SP); KoscinskiSP 31640 fl (SP); KuhlmannSP 24062 fl (SP, US); Lofgren456 fl (C, SP);
Mosen2838 fl (C, K, LD, LE, R, S); Novaes690 fl (US); Palma R 63195 fl (R); Pickel5570 fl (IPA, US);
Riedeli665 fl (NY, US); Serv.Florestal31 fl (FI). Rio deJaneiro: Herb.Saldanhasn fl (R 63194); Goes&
Constantino 965 fl (RB 51668); Nunes299 fl (RB 47922); Pabst 7232 fl (HB, NY); Velloso200 fl (R 38616).
Guanabara: Duarte 4858 fl (RB I110053); Frajao RB i5511 fl (RB, S); Glaziou 2127 fl (BR, C, P, R);
2559 fl (BR, C, K, P); 11942 fl (BR, C, K, P); Hoehne4206 fl (GH, SP); HortoFlorestalRB 136916 fl
(RB); KuhlmannRB 15097 fl (K, RB, S, US); Porto306 fl (RB 15505); Schwacke68 fl (R 7154). Rio de
Janeiro or Guanabara: Ackermann sn fl (BR); Luschnathsn fl (BR, C, P); Miers sn fl (NY); PortoRB i5506
fl (RB, S); Riedel sn fl (NY, R, US); Rosa 129 fl (RB I38161); Schott 4972 fl (F, NY, W). Parana:
Dusen7441 fl (S); 8222a st (S, US); 8653 fl (BM, F, GH, K, NY, S); 10114 st (NY, S); 13536 fl (A, BR,
S); i6150 fl (GH, S, US); Hatschbach1121 fl (RB, SP, US). Santa Catarina: Gevieski7 fl (NY, S, US);
131 fr (S, US); Klein 148 (Eq. Ecol. 41) fl (NY, S, US); 754 fl (S, US); 871 fl (NY); 898 fl (NY); 921
fl (NY); 2169 fl (NY); 2304 fl (BR, F, K, US); Lourteig2340 fl (NY, P); Muller 215 fl (K); 322 fl (R);
Nadeaudsn fl (P); Reitz & Klein 1690 fl (NY, S, US); 3059 fr (NY); 3234 fl (S, US); 5778 fl (BR, GH,
K, NY); 5896 fi (S, US); 5973 fl (NY); 6091 fl (US); 8067 fl (BR, K); 8640 fl (US); 9278 fl (US).
Rio Grande do Sul: Dutra 300 fl (R); Henz 53330 fl (S, US); Malme 762 fl (S); Rambo37841 fr (NY);
38606 fl (NY); 39047 fl (BR, LE, US); 39194 fl (NY); 40100 fl (NY, P).

LOCALNAMES.Cinzeiro, Pau de lixa.

Hooker (Mart. Fl. Bras. 14(2): 28. 1867) divided this species into three varieties
which he left unnamed, basing them on small leaf differences. The further material
now available shows that the presumed differences do not exist. This distinct species
is not easily confused with others of Hirtella.

86. Hirtella enneandra Cuatrecasas, Fieldiana Bot. 27: 58. I950.

Tree 20.0 m tall, the young branches sparsely puberulous, soon becoming
glabrous and conspicuously lenticellate. Leaves oblong, coriaceous, 4.5-11.5 cm
long, 1.8-3.8 cm broad, acuminate at apex, with acumen 8.0-I8.0 mm long, rounded
to subcuneate at base, glabrous beneath, the palisade glands occasional; primary
veins 8- I pairs, prominulous on both surfaces; midrib prominent and glabrous on
both surfaces; petioles 2.0-3.0 mm long, terete, eglandular, glabrous. Stipules
2.0-3.0 mm long, lanceolate, caducous, pubescent, eglandular. Inflorescences
axillary and terminal racemes 3.0-6.0 cm long, the rachis sparselypuberulous. Bracts
0.5-2.0 mm long, acute, puberulous, persistent, eglandular. Flowers 6.0-7.0 mm long.
Hirtella 355

Receptacle campanulate, sparsely puberulous on exterior, glabrous within except

for retrorsehairs at throat; pedicels 5.0-6.0 mm long, eglandular. Calyx lobes acute,
sparsely puberulous on exterior, gray-puberulous within. Petals 5, white, glabrous.
Stamens 9, inserted in a complete circle. Style hirsute on lower portion. Ovary inserted
at mouth of receptacle, pilose. Fruit not seen.
TYPE.Cuatrecasas I8370, Colombia, Valle, fl (holotype, F; isotypes, COL, P).
DISTRIBUTION. Montane forest in western Colombia; known only from the type
collection.
This poorly known but distinct species, differs from most others of Hirtellain the
greater number of stamens which are arranged in a complete circle.

87. Hirtella pauciflora Little, Jour. Wash. Acad. Sci. 38: 88. I948.

Large tree, the young branches puberulous, becoming glabrous with age. Leaves
elliptic, chartaceous, 7.0-13.0 cm long, 3.0-5.0 cm broad, acuminate at apex, with
acumen 6.o-Io.o mm long, subcuneate at base, with a few stiff appressed hairs
beneath, the palisade glands absent; primary veins 7-I pairs, prominulous on both
surfaces; midrib prominulous above, prominent and puberulous beneath; petioles
2.5-3.5 mm long, terete, eglandular, tomentellous when young. Stipules ca 3.0 mm
long, linear, persistent, eglandular, pubescent. Inflorescences few-flowered racemes
3.0-6.0 cm long (3-flowered in specimen examined), the rachis puberulous to tomen-
tellous. Bracts persistent, eglandular. Flowers ca Io.o mm long. Receptacle cam-
panulate, tomentellous on exterior, glabrous within, borne on long thickened woody
pedicels ca I2.0 mm long, eglandular. Calyx lobes rounded, puberulous on both
surfaces. Petal 5, glabrous. Stamens 3, unilateral, the filamentsglabrous. Style hirsute
at base only. Ovary inserted at mouth of receptacle, pilose. Fruit not seen.
TYPE. Little 6475, Ecuador, Los Rios, fl (holotype, US; isotypes, K, NY, US).
DISTRIBUTION. Known only from the type gathering collected in primary forest
in Ecuador.
LOCAL NAME. Coquito.
This is an extremely poorly known species whose exact relationship within the
genus will only be determined when further material is collected. Judging by the
single inadequate collection examined, this species is distinct by the few-flowered
inflorescences,and the thick woody pedicel. I do not place much weight on the latter
character since it is possible that in the type the flowers were diseased.

88. Hirtella glaziovii Taubert, Bot. Jahrb. 15(Beibl. 34): 8. 1892.

Tree? Young branches sparsely tomentellous, becoming glabrous with age.

Leaves oblong-elliptic, coriaceous, 3.5-5.5 cm long, 1.5-2.7 cm broad, acute at apex,
cuneate at base, with few sparse appressed hairs beneath, the palisade glands fre-
quent especially towards base; primary veins 6-8 pairs, prominulous on both sur-
faces; midrib prominulous above, prominent and with an appressed pubescence
beneath; petioles 2.0-3.0 mm long, terete, eglandular, puberulous. Stipules 2.0-4.5
mm long, linear, tomentellous, eglandular, caducous. Inflorescences terminal and
subterminal racemes, the rachis rufous tomentose. Bracts 1.5-3.0 mm long, oblong,
acute, tomentellous, eglandular. Flowers 6.0-7.0 mm long, Receptacle campanulate-
cupuliform, tomentose on exterior, glabrouswithin except for retrorsehairs at throat;
356 Flora Neotropica

pedicels 1.5-3.0 mm long, eglandular, Calyx lobes acute, tomentellous on both

surfaces. Petals 5, glabrous. Stamens 3, unilateral, the filaments glabrous, shortly
exceeding calyx lobes. Style pilose up to half its length. Ovary inserted at mouth of
receptacle, pilose-tomentose. Fruit ovoid; epicarp sparsely tomentellous; mesocarp
thin, fleshy; endocarp thin, hard, hirsutulous within.
TYPE. Glaziou 4946, Brazil, Guanabara, fl (holotype B, lost; lectotype, C;
isotype fragment F; isotypes, K, NY, P, R, S).
DISTRIBUTION.
Eastern Brazil.
BRAZIL. Sao Paulo: HoehneSP I119 fl (NY, SP); Kuhlmann1577 fl (RB); i748 fr (NY, SP);
4328 fr (SP). Guanabara: Hoehne188 fl (SP); Markgraf3096 fl (NY, RB).

African Species

89. Hirtella megacarpa R. A. Graham, Kew Bull. 1957: 230. 1957.

TYPE. Drummond & Hemsley 2614, Tanzania, fl (holotype, K).

Confined to Tanzania and Mozambique.
DISTRIBUTION.

go. Hirtella zanzibarica Oliver, Hooker Ic. P1. 12: 8I, t. 1193. 1876.
Hirtella thouarsianaBaillon ex Lanessan, P1. Util. Colon. Fran9. 874. i886. Type. Du Petit Thouars
sn, Madagascar, fl (holotype, P).

TYPE. Kirk sn, Mafia Is., fl fr (holotype, K; isotypes, F, GH).

DISTRIBUTION. Tanzania, Kenya, Mozambique, and Madagascar.
In addition to the two African species listed here, Capuron (Publ. Centr. Tech.
For. Trop. Madag. 1966), described a third, H. tamenaka. As the detailed description
lacks a Latin diagnosis, this species is not yet validly published. I have not seen
material of H. tamenaka,and am unable to evaluate it at present.

Imperfectly Known Species (not included in key)

91. Hirtella cliffortiana Vellozo, Fl. Flum. 91. 1825, 2. t. I36. 1827; Arch.
Mus. Nac. Rio de Janeiro i88I: 87. I88I.

TYPE. (not seen).

I have not been able to typify this species, which has only a brief description.

92. Hirtella pohlii Hooker f., Mart. Fl. Bras. 14(2): 33. 1867.

TYPE.Pohl. sn, Brazil, Guanabara, (not seen).

A Pohl collection at Munich annotated H. pohlii does not match Hooker's
description at all. I have not yet been able to find further material of this species,
and there is none in the herbaria at Kew or Vienna.
The description of Hirtellapohlii is close to the species named H. burchelliihere,
and it is possible that the type of the name H. pulchra,a synonym of H. burchellii,is in
Hirtella 357

fact H. pohlii. However, the localities are different between that specimen and
Hooker's description of H. pohlii.

Name Without Description

Hirtellahostmanniana
Klotsch in Schomburgk, Faun. et Fl. Brit. Guian. I 98. 1848.
HirtellaroraimaeKlotzsch in Schomburgh, Faun. et Fl. Brit. Guian. II03. 1848.

Excluded Species
New World
i. Hirtella apetalaE. Mey., Nova Acta Acad.Leop.-Carol. 21: 803. I825. =Licania
apetala(E. Mey.) Fritsch.
2. H. canomensis (Mart.) Spreng., Linn., Syst. Veg. ed. I6. 4(Cur. Post.): 34I. I827.
=Couepiacanomensis (Mart.) Benth. ex Hook. f.
3. H. cognataSteud., Flora 26: 76I. 1843. =Couepiacognata(Steud.) Fritsch.
4. H. dodecandra Moc. & Sesse ex DC., Prodr. 2: 529. I825. =Couepiapolyandra
(Kunth) Rose.
5. H. glaberrimaSteud., Flora 26: 761. I843. =Crudiaglaberrima(Steud.) Macbr.
(Leguminosae).
6. H. octandra Hoffmgg. ex Roem. & Schult., Linn. Syst. Veg. 5: 274. I819. =Licania
octandra(Hoffmgg, ex Roem. & Schult.) Kuntze.
7. H. polyandraKunth, Nov. Gen. et Sp. 6: 246. 1823. =Couepiapolyandra(Kunth)
Rose.
Africa
i. Hirtella bangweolensis (R. E. Fries) Greenw. =Magnistipula bangweolensis(R. E.
Fries) R. A. Grah.
2. H. butayi(De Wild.) Brenan. = MagnistipulabutayiDe Wild.
3. H. conrauana(Engl.) A. Chev. =Magnistipulaconrauana Engl.
4. H. cupheiflora (Mildr.) Mildbr. =Magnistipulacupheiflora Mildbr.
5. H. eglandulosa Greenw. = Magnistipulaeglandulosa(Greenw.) R. A. Grah.
6. H. fleuryanaA. Chev. = Magnistipulafleuryana (A. Chev.) Hauman
7. H. fruticulosaHauman. = Magnistipulaeglandulosa(Greenw.) R. A. Grah.
8. H. glaberrima(Engl.) A. Chev. = Magnistipulaglaberrima Engl.
9. H. katangensis Hauman. = Magnistipulakatangensis(Hauman) Mendes.
o. H. montanaHauman. = Magnistipulamontana(Hauman) Prance.
II. H. olivaeformisPoir., Encyc. Suppl. 3: 53. = IcacinasenegalensisJuss.(Icacinaceae)
12. H. porosa(Boiv. ex Baill.) Drake. = GrangeriaporosaBoiv. ex Baill.
I 3. H. sapinii(De Wild.) A. Chev. =MagnistipulasapiniiDe Wild.
I4. H. zenkeri(Engl.) A. Chev. = MagnistipulazenkeriEngl.

8. Acioa Aublet, P1. Guiane 2: 698, t. 280. I775; Focke in Engl. & Prantl, Nat.
Pflanzenfam. 3(3): 60. 1894; Maguire, Brittonia 7: 27I. I951; Prance,
in press.
Dulacia Necker, Elem. Bot. 414. 1790. non Dulacia Vellozo, 1823.
Acia Schreber, Linn. Gen. P1. ed. 8. 2: 458. 1791 pro parte exl.syn Couepia
Moquileasensu Martius, Nov. Gen. et Sp. 79. 1826 pro parte quoad syn Acioa tantum; sensu
Blume, Mus. Bot. Lugd. Bat. 2: 92. I852 pro parte quoad syn Acioa tantum.
DactyladeniaWelwitsch, Apont. in Ann. Cons. Ultram. 55. 572. I859.
GriffoniaHooker f. in Bentham & Hooker f., Gen. P1. I: 608. 1865 non GriffoniaBaillon.
Couepiasensu Hooker f. in Bentham & Hooker f., Gen. PI. :: 608. I865 quoad syn Acioatantum.

Shrubs or small trees with hermaphrodite flowers I2.o-40.o mm long. Leaves

358 Flora Neotropica

usually with two glands towards the base of the lamina, glabrous or with stiff erect
rufous hairs beneath. (rarely white-lanate in some African species); petioles eglandu-
lar. Bracts and bracteoles eglandular (glands numerous and stipitate in some African
species only), not enclosing flowers in groups. Inflorescence of branched or dichoto-
mous racemes or less frequently of panicles. Receptacle usually elongate, obconic-
tubular, rarely campanulate, hollow, glabrous within, tomentose or glabrous on
exterior. Calyx lobes rounded or acute, puberulous within. Stamens IO-65, the
filaments ligulately connate for most of their length, inserted laterally at mouth of
receptacle; staminodes sometimes present and opposite stamens. Ovary inserted
laterally at mouth of receptacle; carpels unilocular. Fruit a hard or fleshy drupe, often
markedly tapered towards base or apex; epicarp often rufous-tomentose, less often
glabrous; mesocarp thin; endocarp usually hard, thin, with a roughish surface, hairy
within, with no special mechanism for seedling escape.
TYPE SPECIES.Acioa guianensis Aublet. The name Acioa is derived from the ver-
nacular name in French Guiana.
DISTRIBUTION. Largely in West Africa and the Congo Basin, with three little-
known South American species in the Guianas and Amazon basin.
This genus has been discussed elsewhere on a worldwide basis (Prance, in press)
as has been the South American element alone (Maguire, 195I). Since the appearance
of Maguire's account little extra information has been gathered, and Acioa in the
New World remains poorly known.

Taxonomic History of Acioa

Acioa was first described by Aublet (I775) from a single specimen which he
collected in French Guiana. Both Necker (1790) and Schreber (I791) created new
and consequently invalid names for Acioa.Necker renamed it Dulacia, and Schreber
renamed it Acia. During the nineteenth century the Aublet material was ignored by
most of the workers publishing on Acioa, with the result that Acioa was not recog-
nized as a distinct genus. Martius (I827) reduced Acioato synonymy under Moquilea.
Moquileasensu Martius largely contained species of Coupiatogether with the Aublet
type species of Acioaand Moquilea,a concept followed by several other workers, eg,
Zuccarini (I832), Meisner (1837), and Endlicher (I840). Hooker (1865), in Bentham
and Hooker's GeneraPlantarum,cited Acioaas a synonym of Couepia.In the same work
Hooker described a new genus from Africa which be named Griffonia.Baillon (1869)
reinstated Acioaas a genus and was the first to connect the American Acioaguianensis
with the African genus Grifoniaof Hooker. He cited Griffoniaas a synonym of Acioa.
Since the time of Baillon, Acioahas generally been regarded as a distinct genus. Many
new African species have been described, raising the total from that continent to 31.
Maguire (i951) described two more species from South America. The three species
from Malesia by Kostermans (I965b) differ from American and African Acioa in
important characters, and will be transferredto a new genus in Prance (in press).

Key to American Species of Acioa

i. Receptacle obconical-tubular; sepals pubescent within; staminal ligule I0.0-I2.0 mm long,
2.0-3.0 mm broad.
2. Leaf apex abruptly short-acuminate, the base cuneate or at least tapered at junction with
petiole; petioles 2.0-3.0 mm thick. i. A. guianensis.
2. Leaf apex rounded to retuse, the base broadly rounded; petioles 4.0-5.0 mm thick.
2. A. somnolens.
i. Receptacle campanulate and gibbous; sepals glabrous within; staminal ligule 8.0-Io.o mm
long, ca 5.0 mm broad. 3. A. schultesii.
Acioa 359

The American Species of Acioa

i. Acioa guianensis Aublet, P1. Guiane 2: 698, t. 280. I775; DC., Prodr. 2:
526. 1825. Fig 58 A-C.
Acia dulcisWilldenow, Linn., Sp. P1. ed. 4. 3: 7I7. 800onom. illegit.
Acioa dulcis (Willdenow) Steudel, Nom. ed. I. 9. 1821 nom illegit.
MoquileaaubletianaBlume, Mus. Bot. Lugd. Bat. 2: 92. 1852 nom illegit.
Licaniaaubletiana(Blume) Lem6e, Fl. Guyane Frans. 2: 23. 1952 nom illegit.

Medium-sized tree. Leaves ovate to oblong, 7.0-I 7.5 cm long, 3.0-8.3 cm broad,
cuneate or slightly tapered at base, abruptly short-acuminate at apex, with acumen
2.0-5.0 mm long, glabrous on both surfaces; primary veins 7-9 pairs, inconspicuous
on both surfaces; petioles 5.0-I2.0 mm long, 1.5-3.0 mm thick, glabrous, terete.
Stipules caducous (not seen). Inflorescencesterminal and axillary panicles, the rachis
and branches glabrous. Receptacle obconic-tubular, I0.0-30.0 mm long, glabrous on
exterior, glabrous within except for deflexed hairs at throat. Calyx lobes 5, pubescent
within. Petals 5, with minutely serrate margins. Stamens unilateral, I0-I2, the
filaments ligulately connate. Ovary inserted at mouth of receptacle, densely pilose.
Style pilose on lower portion. Fruit ellipsoid; epicarp glabrous and sparsely lenti-
cellate; mesocarp to I4.0 mm thick, fibrous; endocarp thick, hard, merging gradually
into mesocarp, with thick fibers, lacking mechanism for seedling escape, glabrous
within.
TYPE. Aublet sn, French Guiana, fl (BM).
DISTRIBUTION. Forest in French Guiana and Amazonian Brazil, apparently
extremely rare.
FRENCH GUIANA. Melinon32 fl (P); Perrotetsn fl (P); Poiteausn fl (K, P); Sagot sn fl (P).
BRAZIL. Amapd: Pires, Rodrigues& Irvine50490 fl (COL, F, K, LE, NY, P, RB, US). Guanabara:
O. A. da Silva RB 52237, cult fl fr (F, IAN, NY, R, RB).

LOCALNAMES.Brazil: Coco de Cotia, Castanha de Cotia.

This species may be distinguished from A. somnolensin a few leaf characters.
The differences in flower size given by Maguire have not proved valid in the further
material studied. The origin of the cultivated material in the Rio de Janeiro Botanical
Garden was the Rio Purus in Amazonian Brazil, indicating a wider distribution than
that of the collections cited here.
The material grown at Rio de Janeiro was introduced by Dr. Ducke. When
he was travelling on the Rio Purus he found the natives using fruits unknown to him
as a soap. The fruits were collected from the river where they were floating in large
quantities. When the contained seeds were germinated at Rio de Janeiro, the result-
ing plants proved to be Acioaguianensis.Although this species has never been collected
from the Rio Purus, I have, examined two collections of Acioafrom near that region.
These are not cited here because they are considerably different in leaf characters
from A. guianensisand from each other. These collections (L. Garciasn, from the Rio
Japura, and Rodrigues& CoelhoI408 from Tefe) are both fruiting. At this stage of our
knowledge of the genus it is impossible to say whether they are A. guianensisor whether
they represent one or even two undescribed species.

2. Acioa somnolens Maguire, Brittonia 7: 272. 195I.

Tree? Leaves broadly elliptic to oblong-elliptic, I0.0-15.o cm long, 6.0-9.0 cm
broad, rounded to slightly retuse at apex, rounded at base, glabrous on both
360 Flora Neotropica

---- -
i?-

JTLoken

FIG 58. Species of Acioa and their geographical listribution, A-C, A. guianensis (Pires et a
50490); A, habit, X o.5; B, leaf undersurface, x 5; C, flower section, x i. D-E, A. schultesii(Schultese
al 958); A, habit, x 0.5; B, flower section, x i. ? A. guianensis; ? A. somnolens;A A. schultesii.
Acioa 361

surfaces; primary veins 6-8 pairs, inconspicuous on both surfaces; petioles thick,
8.0-io.o mm long, 4.0-5.0 mm broad. Stipules caducous (not seen). Inflorescences
terminal and axillary panicles, glabrous. Bracts and bracteoles small, caducous.
Receptacle obconic-tubular, I8.o-20.0 mm long, glabrous on exterior, glabrous
within except for deflexed hairs at throat. Calyx lobes 5, pubescent within. Petals 5,
caducous. Stamens ca 15, unilateral, the filaments ligulately connate; ligule 2.0-3.0
mm broad, I0.0-12.0 mm long. Ovary pilose, inserted at mouth of receptacle. Style
sparsely hirsute for two-thirds its length. Fruit unknown.
TYPE.Melinon230, French Guiana, fl (holotype, P).
DISTRIBUTION.
Known only from French Guiana. The specimens examined lack
detailed field notes.
FRENCH GUIANA. Desvauxsn fl (P); withoutcollector'sname413 fl (R 63209).

3. Acioa schultesii Maguire, Brittonia 7: 272. I95I. Fig 58 D-E.

Small tree. Leaves broadly elliptic to oblong-elliptic, 8.o-I6.o cm long, 4.5-

8.5 cm broad, obtusely cordate at base, rounded to short-acuminate at apex, galbrous
on both surfaces; primary veins inconspicuous; petioles 3.o-6.0 mm long, glabrous,
thick. Stipules caducous (not seen). Inflorescences axillary and terminal panicles,
the rachis and branches glabrous. Bracts and bracteoles small, the margins ciliolate.
Receptacle broadly campanulate and gibbous, glabrous on exterior, glabrous within
except for deflexed hairs at throat. Calyx lobes 5, glabrous on both surfaces except
for ciliate margins. Petals 5, the margins ciliate. Stamens ca 15, unilateral, the fila-
ments ligulately connate, with ligules 5.0 mm thick. Ovary pilose, inserted at mouth
of receptacle. Style glabrous except for hirsute base. Fruit unknown.
TYPE.Schultes& L6pez9958, Brazil, Amazonas, Rio Dimite, fl (holotype, NY;
isotype, K).
Known only from the type gathering, collected from a steep
DISTRIBUTION.
mountain slope. This species is quite distinct from all other species of Acioa in the
campanulate and gibbous receptacle, and in the short broad staminal ligule. How-
ever, as it shows the very distinctive leaf morphology of all species of the genus, there
is no doubt that it is a species of Acioa.

Excluded Genera
LecostemonMocifio & Sesse ex A. P. de Candolle, Prodr. 2: 639. 1825. =Sloanea
(Elaeocarpaceae).
Lecostemonsensu Bentham, Jour. Bot. Kew Misc. 5: 295. I853. =Rhabdodendron
(Rhabdodendraceae).
362 Flora Neotropica

Table V

Phytogeographic Distribution of the Chrysobalanaceae

*3 " ^K c3
x . S-

U.S.A. Gulf states - I - -

Central America - 7 - - 3+i
Caribbean Islands - - - - 2
Pacific coastal Colombia & Ecuador 9 I - 3
Northern Colombia and Venezuela - 2
Central Colombia 2 - - I 2+i
Guayana highlands (over Iooo m. alt) - - 2
Guianas incl. Venezuela Bolivar &
D. Amacuro, Brazil Amapa & - 23+i 2 - - 7+iii 10 I
N. Roraima
widespread - I2+i - - - 6+i 6
Amazonia western 14 3 - 6 18 I
central - + ii I - 8 4
eastern 2 3 i 4
Bolivia - - - - - -
Planalto of central Brazil - 4 I 2 - 2 4
Northeastern Brazil 3 3
Eastern Brazil I +i I I - 7 9+i
Widespread in two or more of above
Central America, Colombia, Guianas i -ii
& Amazon
Caribbean &/or Trinidad and northern
coastal Venezuela 3
Colombia, Peru and Guianas,
- -
Venezuela Bolivar I - ---
Guianas and eastern Amazonia - 1 I I - - 3
Guianas and western or central Amazonia
or widespread Amazonia - 19+i 2 - 5 8+ii I
- I +iii - - - 2
Guianas, Amazon to S. Brazil
Amazon and E. central Brazil - I
Amazon and Bolivia - I
Eastern and northeast Brazil - i - - - i - -
S. coastal U.S.A., Caribbean,
Central Am. to S. Brazil -
Central Am., Cuba, Colombia &
Venezuela - - - - -
Amazon and northeastern Brazil - - i

arabic numerals = species

roman numerals = subspecific taxa
 NUMERICAL LIST OF TAXA

I. Chrysobalanus 47. L. subarachnophylla Cuatr.

I. C. icacoL. 48. L. salicifolia Cuatr.
2. C. cuspidatusGriseb. ex Duss 49. L. araneosaTaub.
50. L. silvaticaGlaziou ex Prance
2. Licania 5I. L. chocoensisCuatr.
I. L. michauxiiPrance 52. L. licaniaeflora(Sagot) Blake
2. L. boliviensisPrance 53. L. hirsutaPrance
3. L. maritimaPrance 54. L. costaricensisStandl. & Steyerm.
4. L. durifoliaCuatr. 55. L. krukoviiStandl.
5. L. macrocarpa Cuatr. 56. L. lasseriMaguire
6. L. salzmanii (Hook. f.) Fritsch 57. L. latifolia Benth. ex Hook. f.
7. L. klugii Prance 58. L. minusculaCuatr.
8. L. guianensis(Aubl.) Griseb. 59. L. operculipetala Standl. & L. 0. Wms.
9. L. retifoliaBlake 60. L. reticulataPrance
Io. L. longipedicellataDucke 61. L. arachnoidea Fanshawe & Maguire
I. L. tomentosa(Benth.) Fritsch 62. L. oblongifoliaStandl.
I2. L. pyrifoliaGriseb. 63. L. macrophylla Benth.
13. L. leucosepalaGriseb. 64. L. caudataPrance
I4. L. angustataPrance 65. L. latistipulaPrance
I5. L. platypus(Hemsl.) Fritsch 66. L. divaricataBenth.
I6. L. gonzalezii Miranda 67. L. glabrifloraPrance
I 7. L. egleriPrance 68. L. intrapetiolarisSpruce ex Hook. f.
8. L. minutiflora(Sagot) Fritsch 69. L. heteromorpha Benth.
19. L. maranhensis Prance a. var heteromorpha
20. L. fritschii Prance b. var glabra (Mart. ex Hook. f.) Prance
21. L. brittenianaFritsch c. var subcordataFritsch
22. L. unguiculataPrance d. var perplexansSandw.
23. L. longipetalaPrance 70. L. glaziovianaWarm.
24. L. wurdackiiPrance 71. L. littoralisWarm.
25. L. turbinataBenth. a. var littoralis
26. L. lata Macbr. b. var cuneataKuhlm.
27. L. apetala (E. Mey.) Fritsch 72. L. fanshawei Prance
a. var apetala 73. L. irwinii Prance
b. var aperta (Benth.) Prance 74. L. cyathodesR. Ben.
28. L. parvifoliaHuber 75. L. polita Spruce ex Hook. f.
29. L. maguireiPrance 76. L. silvae Prance
30. L. gardneri(Hook. f.) Fritsch 77. L. densifloraKleinh.
31. L. cuspidata(Rusby) Prance 78. L. cupreaSandw.
32. L. sparsipilisBlake 79. L. impressaPrance
33. L. emarginataSpruce ex Hook. f. 8o. L. dealbataHook. f.
34. L. calvescensCuatr. 81. L. pallida Spruce ex Sagot
35. L. persaudiiFanshawe & Maguire 82. L. gracilipesTaub.
36. L. sprucei(Hook. f.) Fritsch 83. L. parvifructaFanshawe & Maguire
37. L. sclerophylla(Mart. ex Hook. f.) Fritsch 84. L. cymosaFritsch
38. L. albifloraFanshawe & Maguire 85. L. ternatensisHook. f. ex Duss
39. L. longistyla(Hook. f.) Fritsch 86. L. membranacea Sagot ex Lanes.
40. L. fuchsii Prance 87. L. piresii Prance
41. L. humilisCham. & Schlecht. 88. L. hypoleucaBenth.
42. L. foveolataPrance a. var hypoleuca
43. L. octandra(Hoffmgg. ex Roem. & Schult.) b. varfoveolata Prance
Kuntze 89. L. boyaniiTutin
a. subsp octandra go. L. buxifoliaSandw.
b. subsp pallida (Hook. f.) Prance 9I. L. orbicularisSpruce ex Hook. f.
44. L. rigida Benth. 92. L. niloi Prance
45. L. arboreaSeem. 93. L. coriaceaBenth.
46. L. velataCuatr. 94. L. urceolarisHook. f.

363
364 Flora Neotropica

95. L. affinisFritsch 4. P. excelsaSabine

96. L. glauca Cuatr. 5. P. occidentalisPrance
97. L. davillaefoliaR. Ben. 6. P. spruceiHook. f.
98. L. ellipticaStandl. 7. P. pachyphyllaRusby
99. L. canescensR. Ben. 8. P. brasiliensis(Schott) Hook. f,
o00. L. couepifoliaPrance 9. P. klugii Prance
oI . L. trigonioidesMacbr. 10. P. maguireiPrance
102. L. cordataPrance 1. P. littoralisPrance
103. L.foldatsii Prance 12. P. parvifoliaSandw.
I04. L. hebanthaMart. ex Hook. f. 13. P. cardiophyllaDucke
I05. L. steyermarkii Maguire 14. P. parilis Macbr.
Io6. L. subrotundata Maguire I5. P. chocoensisPrance
107. L. crassiveniaSpruce ex Hook. f. i6. P. obtusifoliaHook. f.
Io8. L. majusculaSagot
109. L. alba (Bern.) Cuatr.
Io. L. hitchcockiiMaguire 4. Exellodendron
I I. L. sandwithiiPrance I. E. coriaceum(Benth.) Prance
112. L. laxifloraFritsch 2. E. cordatum(Hook. f.) Prance
I13. L. rufescensKlotzsch ex Fritsch 3. E. barbatum(Ducke) Prance
I 14. L. kunthianaHook. f. 4. E. gardneri(Hook. f.) Prance
I 5. L. bellingtoniiPrance 5. E. gracile (Kuhlmann) Prance
I 16. L. compactaFritsch
I I 7. L. ovalifoliaKleinh.
5. Maranthes
18. L. caldasianaCuatr. i. M. corymbosa
Blume
19. L. savannarum Prance
I20. L. microphylla Fanshawe & Maguire
2 I. L. triandraMart. ex Hook. f. 6. Couepia
I22. L. discolorPilg. I. C. guianensisAubl.
123. L. apiculataPrance 2. C. glandulosaMiq.
124. L. micranthaMiq. 3. C. paraensis(Mart. & Zucc.) Benth.
125. L. pruinosaR. Ben. a. subsp paraensis
126. L. nitida Hook. f. b subsp glaucescens(Spruce ex Hook. f.)
127. L. riedeliiPrance Prance
I28. L. bracteataPrance 4. C. leptostachya Benth. ex Hook. f.
129. L. parvifloraBenth. 5. C. maguireiPrance
I30. L. robustaSagot 6. C. sandwithiiPrance
I3I. L. lanceolataPrance 7. C. parillo DC.
132. L. spicataHook. f. 8. C. steyermarkii Maguire
133. L. strictaKleinh. 9. C. canomensis(Mart.) Benth. ex Hook. f.
134. L. leptostachya Benth. 10. C.foveolata Prance
35. L. incanaAubl. 11. C. magnoliifoliaBenth. ex Hook. f.
136. L. paraensisPrance 12. C. exflexaFanshawe & Maguire
I37. L. vaupesianaKillip & Cuatr. 3. C. habranthaStandl.
138. L. bahiensisPrance 14. C. spicataDucke
139. L. maximaPrance i5. C. bracteosaBenth.
I40. L. mollis Benth. i6. C. subcordataBenth. ex Hook. f.
14I. L. blackiiPrance I7. C. belemiiPrance
142. L. rodriguesiiPrance i8. C. caryophylloidesR. Ben.
143. L. indurataPilg. I9. C. excelsaDucke
144. L. hoehneiPilg. 20. C. uiti (Mart. & Zucc.) Benth. ex Hook. f.
145. L. cruegeriana Urb. 21. C. cataractaeDucke
146. L. belemiiPrance 22. C. macrophylla Spruce ex Hook. f.
I47. L. splendens(Korth) Prance 23. C. krukoviiStandl.
148. L. veneralensis Cuatr. 24. C. latifolia Standl.
I49. L. amapaensisPrance 25. C. ovalifolia (Schott) Benth.
150. L. tepuiensisPrance 26. C. schottiiFritsch
151. L. obtusifoliaFritsch 27. C. grandiflora(Mart. & Zucc.) Benth. ex
152. L. roraimensisStandl. Hook. f.
28. C. elata Ducke
3. Parinari 29. C. racemosa Benth. ex Hook. f.
I. P. campestrisAubl. 30. C. martiniiPrance
2. P. montanaAubl. 31. C. bondariiPrance
3. P. rodolphiiHuber 32. C. insignisFritsch
Acioa 365

33. C. recurvaSpruce ex Prance 35. H. mutisiiCuatr.

34. C. obovataDucke 36. H. triandraSw.
35. C. williamsii Macbr. a. subsp triandra
36. C. chrysocalyx(Poepp. & Endl.) Benth. b. subsp punctulata(Miq.) Prance
ex Hook. f. c. subsp media(Standl.) Prance
37. C. erianthaSpruce ex Hook. f. 37. H. bahiensisPrance
38. C. trapezioanaCuatr. 38. H. latifolia Prance
39. C. stipularisDucke 39. H. suffultaPrance
40. C. reflexaDucke 40. H. elongataMart. & Zucc.
4'. C. longipendulaPilg. 41. H. rodriguesiiPrance
42. C. cognata(Steud.) Fritsch 42. H. obidensisDucke
a. var cognata 43. H. cowaniiPrance & Maguire
b. var majorPrance 44. H. orbicularisPrance
c. var membranacea Prance 45. H. guyanensis(Fritsch) Sandw.
43. C. multifloraBenth. 46. H. lightioidesRusby
44. C. ulei Pilg. 47. H. aramangensis Prance
45. C. comosaBenth. 48. H. rasa Standl.
46. C. venosaPrance 49. H. scabraBenth.
47. C. polyandra(Kunth) Rose 50. H. bicornisMart. & Zucc.
48. C. platycalyxCuatr. a. var bicornis
49. C. rufa Ducke b. var pubescensDucke
50. C. robustaHuber 51. H. angustissimaSandw.
51. C. impressaPrance 52. H. tenuifoliaPrance
52. C. meridionalisPrance 53. H. pilosissimaMart. & Zucc.
53. C. pernambucensis Prance 54. H. gracilipes(Hook. f.) Prance
54. C.froesii Prance 55. H. brachystachya Spruce ex Benth.
55. C. parvifolia Prance 56. H. racemosaLam.
a. var racemosa
7. Hirtella b. var hexandra (Willd. ex Roem. &
I. H. myrmecophila Pilg. Schult.) Prance
2. H. physophora Mart. & Zucc. c. var glandipedicellata
Prance
3. H. vesiculosaSuesseng. 57. H.juruensis Pilg.
4. H. dorvaliiPrance 58. H. kuhlmanniiPilg.
5. H. guainiaeSpruce ex Hook. f. 59. H. standleyiBaehni & Macbr.
6. H. duckeiHuber 60. H. longifoliaBenth. ex Hook. f.
7. H. macrosepalaSandw. 6I. H. lemsii L. 0. Wms. & Prance
8. H. ulei Pilg. 62. H. schultesiiPrance
9. H. glabrataPilg. 63. H. paraensisPrance
o. H. carbonariaLittle 64. H. spruceiBenth.
1. H. araguariensisPrance 65. H. lancifoliaDucke
12. H. cordifoliaPrance 66. H. burchelliiBritton
13. H. insignisBriq. ex Prance 67. H. mucronataPrance
I4. H. tocantinaDucke 68. H. longipedicellataPrance
15. H. piresii Prance 69. H. glandistipulaDucke
16. H. davisii Sandw. 70. H. martianaHook. f.
I7. H. subglanduligera Pilg. 71. H. pimichinaLasser & Maguire
i8. H. ciliata Mart. & Zucc. 72. H. subscandens Spruce ex Hook. f.
I9. H. hoehneiPilg. 73. H. hispidulaSteud.
20. H. glandulosaSpreng. 74. H. silicea Griseb.
2 i. H. bullataBenth. 75. H. excelsaStandl. ex Prance
22. H. americanaL. 76. H. adenophora Cuatr.
23. H. guatemalensisStandl. 77. H. caducaFanshawe & Maguire
24. H. eriandraBenth. 78. H. fasciculata Prance
25. H. paniculataSw. 79. H. couepifloraPrance
26. H. deflexaMaguire 80. H. tubifloraCuatr.
27. H. tentaculataPoepp. & Endl. 8I. H. floribundaCham. & Schelcht.
28. H. macrophylla Benth. ex Hook. f. 82. H. angustifoliaSchott.
29. H. adderleyiPrance 83. H. rugosaThuill. ex Pers.
30. H. punctillataDucke 84. H. scaberulaSpruce ex Hook. f.
3I. H. corymbosa Cham. & Schlecht. 85. H. hebecladaMoric. ex DC.
32. H. barrosoiPrance 86. H. enneandraCuatr.
33. H. pendulaSoland. ex Lam. 87. H. paucifloraLittle
34. H. leonotisPittier 88. H. glaziovii Taub.
366 Flora Neotropica

89. H. megacarpaR. A. Grah. 8. Acioa

90. H. zanzibaricaOliv. I. A. guianensisAubl.
91. H. cliffortianaVeil. 2. A. somnolensMaguire
92. H. pohlii Hook. f. 3. .4. schultesiiMaguire
 LIST OF EXSICCATAE
The figures in parentheses refer to the genus number (first), and to the species
number (second), i.e. (genus number-species number). The following generic
numbers are used in the parentheses:
Acioa 8
Chrysobalanus I
Couepia 6
Exellodendron 4
Hirtella 7
Licania 2
Maranthes 5
Parinari 3
These numbers are also given in the numerical list of taxa.
Abbot, W. L., 269(7-36a); 412(7-36); 529(1-I);
I 63(7-36a); I286(1-I); 1378a(7-36a); 2155
(7-36a); 2213(1-I); 2249(1-I); 2531(7-36a);
2808(7-36a); 2863(1-I); 2882(1-1)
Abraham, A. A., 103(7-56a); 258(2-83)
Ackermann, sn(7-8i); sn(7-86)
Acuia,J., 12428(1- )
Agostini, G., 2I5(7-36a); 250(7-21); 320(7-56b)
Aguiar, J. et al., 241 (2-I )
Aguilar H., M., 36(6-47); 145(7-22); 176(7-36b);
I90(7-56b); 329(7-22); 1661(2-15); 17I7
(2-45)
Aguilar, J. I., 1716(7-56b)
Agr. Chefe, Cear,, RB 78265(2-44)
Ahles, H. et al., 18217(2-1)
Aitkin, J. B., 56(2-109)
Alain, Bro., 3641 (7-36a); 3643(7-36a); 9055(7-83)
Alencar, 16(2-44)
Alexander, E.J., 142(7-56b)
Alexander, R. C., sn(7-36a)
Allard, H. A., 15860(I-I)
Allemao, F. F. et al, 560(2-44); 562(2-II);
564(7-18); 567(7-56b)
Allen, Cy., 599(7-22)
Allen, P. H., 930(2-15); 941(2-15); 967(7-36a);
1603(7-56b); I689(7-36a); 174I(2-45); 3082
(7-21); 3453(7-36a); 4156(7-56b); 6030
(2-59); 6032(2-59); 6885(2-15); 7279(6-47)
Almeida, D. de, sn(I-i)
Almeida, H. de, RB 136929(2-126); RB 136931
(2-126)
Alston, A. H. G., 66(6-25); 136(6-25); 525(7-25);
5820(7-36a)
Aluisio, J., 166(2-81); I77(2-69a); I85(2-43b)
Alvarenga, M., RB 90573(7-25); RB 90574(7-54)
Ames,J. S., sn(i-i)
Anderson, A., 39(2-69a); 57(2-77); 232(1-I);
456(2-78); 586(2-135); sn(2-35); sn(7-25);
sn(7-32); sn(I-I)
Anderson, C. W., 554(6-15)
367
Anderson, N.J., sn(7-56b)
Andrade, A. G. et al, 351(6-27); 424(6-27);
443(7-54); 460(6-27); 551(2- 114); 554(2-30)
Andrade, N., 69(6-46)
Andrieux, G., 372(7-56b)
Angel, C. M., 679(7-36a)
Anisits, J. D., 2 82(6-20)
Appun, C. F., 1(2-109); 298(1-I); 378(7-25);
625(6-45); 707(2-8); 1004(7-21); 1747(6-43);
1752(2-93); si(7-51)
Araujo, S. et al, 488(I-I)
Archer, W. A., 421 (1- ); 2390(7-25); 247 (2-77);
7821(2-63); 7824(2-63); 7903(2-63); 7951
(7-56b); 8076(2-1I4)
Arens, K., INPA 410(6-41)
Aristeguieta, L. et al, 562(7-36a); 1566(7-56b);
I653(2-43a); I697(7-36a); I771(2-86); I898
(7-36a); 3004(2-27b) ;3005(6-3b); 3739(7-45);
4493(7-56b); 4505(2-27b); 4589(6-3b); 4597
(7-25); 4607(2-12); 4935(7-36a); 4988(2-12);
5084(6-3b); 5I31(2-27b); 5299(6-3b); 5860
(7-20); 5889(2-27b); 5940(2-27b); 5954
(2-27b); 7250(7-25)
Armond, A. F. das N., 175(3-4)
Asplund, E., 14163(7-5); 14320(6-44)
Assis, V., 109(7-54)
Atkinson, D.J., 88(2-129); 91(2-57)
Aublet, J. B. C. F., sn(8-I); sn(2-7); sn(2-I35);
sn(6-i); sn(3-I); sn(3-2); sn(3-3)
Aubreville, A., 24(1-I); 278(2-73); 280(7-56a)
Augusto, Bro. B., 595(I-I); 634(1-I); 638(1-I);
I457(7-36a)
Avila, i(6-3b)
Aviles, S., 66(7-36a); 949(7-36a)
Baber, 2980(7-22)
Badier, sn(7-56a)
Badini, J., 3236(7-81)
Bailey, I. W. et al, A 2(7-56b); 89(7-25); 407
(7-36a); 1373(7-25)
368 Flora Neotropica

Bailey, L. H., et al, 1422(2-27b); 1438(2-27b); B6na, R., I299(2-I 4)

I658(2-27b) Benoist, R., I46(2-135); I80(2-108); 270(3-I);
Baker, C. F., et al, 9(1-I); 57(6-36); 64(2-1); 278(2-135); 317(3-I); 555(2-I35); 625
MG 417(7-27); 2147(1-1); I4914(2-69a); (2-88a); 726(2-134); 778(2-74); 779(I-1);
sn(2-I) 950(2-52); 994(3-I); 1085(2-97); I089
Baldwin, J. T., 2748(7-56b); 3200(7-6); 3286 (2-I08); I498(2-69a); I519(3-1); I658(I-I);
(7-56a); 3447(2-140); 3453(2-69b); 3463 1709(1-I)
(4-i) Bernardi, A. L., 769(7-27); 1080(7-22); io8i
Bang, M., 1468(7-73); 2418(7-46) (2-27b); II07(7-56b); 1119(2-43a); 1215
Bangham, W. N., 398(7-36a) (3-7); I426(7-74); 1463(2-1 I4); I509(2-69a);
Barclay, G. W., 855(7-10); 926(7-56a) I538(2-I24); I540(3-4); I647(7-30); I723
Barkley, F. A., et al, 170356(1-I) (7-36a); I762(7-36a); 1767(7-36a); 1952
Barlow, F. D., I/I5(1-1); 6/5(7-56b); 7/3c(I-I); (7-22); 2138(3-3); 2I40(2-109); 2184(7-5ob);
29/4D(6-47); 29/21(6-47); 30/56A(I-I); 30/ 2346(2-43a); 2611(7-2I); 2658(2-81); 2737
62(7-56b); 30/15I (7-56b) (2-105); 2755(2-8I); 2951(2-77); 3001(7-45);
Barnes, V., 85(7-30) 3030(2-109); 3032(2-77); 5719(2-86); 6486
Barreto, M., 831(7-82); 4759(7-85); 6783(6-27); (7-73); 6491(2-77); 6524(7-21); 6620(7-49);
6784(6-27 and 7-20); 6785(7-20); 6787 7125(2-I3); 716(2-12-); ; 784(7-I6); 7275
(7-54); 7289(6-27); 8732(6-26); 9158 (2-112); 7321(7-45); 7505(2-109); sn(2-69a)
(7-54); 9227(6-26); 9998(7-20); 12127(7-20) Bernoulli, C. G., et al, 77(7-56b); 854(1-1); 2960
Barros Correia, 4215(6-49) (7-56b); 2961 (7-56b); 2966(1-1)
Barros, J. de, RB 47979(2-41) Bertero, C. G. L., sn(7-36a)
Barroso, G. M., 620(7-54) Berti, L. M., 66(7-16); 77(3-4); 81(2-o09); 84
Bartlett, A. W., 8309(2-135) (3-3); 94(7-i6); 120(2-27b); 124(3-4); I30
Bartlett, H. H., I1206(7-56b); iI218(i-I); (2-27b); I37(2-77); I56(7-i6); 172(2-109);
I I29(2-88a); I539(7-56b); I793(2-88a); I74(3-3); 184(2-77); I85(2-109); 244(2-113);
11840(7-56b); II932(7-56b); 12471(7-22); 290(2-88a); 315(3-4); 346(2-65); 375(7-7);
12741(7-22); I2974(7-56b); I3034(2-88a); 426(2-113); 566(3-4); 576(3-4); 588(3-3)
I3095(2-32); sn(I-i) Bettenfield, M., sn(2-63); sn(2-86)
Bastos, M., I(3-I); 5(7-24); 40(7-I8); 54(2-63); Beyrich, H. K., 132(2-I)
60(7-20 and 6-4); i62(2-69a); 199(7-24); Billberg, J. E., sn (7-56b)
4I4(2-69a); RB I3817(2-141); RB 96382 Biltmore Herb., 1949 a-f(2-I)
(7-24) Binot, P. M., 40(2-132)
Bates, G., sn(i-i); sn(2-I) Birkley, P., 2030(1-1)
Batten-Poole, 31(6-37) Black, G. A., et al, 46-59(6-35); 47-996(2-43a);
B. B. S. (Dienst's Lands Bosbeheer, Suriname), 47-I 67A(6-3a); 47-1458(7-56a); 47-1632
79(2-69a); I48(2-63); i96(2-88a); 197 (6-34); 47-I689(7-56a); 47-1738(2-67); 47-
(2-124); I99(2-135); 200(2-I35); 202(2-130); I744(7-5oa); 47-I758(4-1); 47-1765(6-3a);
245(7-25); 1027(2-117); I03I(2-63); 1032 47-I833(7-I8); 47-I854(2-57); 47-I9I2
(2-27a); II03(6-I); I 04(2-99); 1106(2-117); (2-67); 47-1959(6-37); 47-I975(2-97); 47-
1107(2-I30); 11I0(2-66); iiii(6-i); 1124 2150(2-62); 48-24I5(7-56b); 48-2435(7-24);
(2-27a) 48-2468(6-2); 48-247I (2-43b); 48-2588
Beard, J. S., II2(7-56a); 166(7-32); 192(2-85); (7-21); 48-2663(3-6); 48-270 (2-69a); 48-
292(7-36a); 488(1-2); 492(I-2); 657(7-36a) 2788(2-69b); 48-2828(6-2); 48-2902(2-69a);
Beard, P., I050(7-32); 1097(1-I) 48-2934(I-I); 48-3222(1-I); 48-3310(2-I35);
Beaumont, sn(2-I) 48-3313(2-37); 48-3360(2-39); 48-3386(2-7);
Beccari, sn (2-93) 48-3414(2-63); 48-3448(7-56b); 48-3461
Becerra, M. E., 9(2-44) (7-20); 48-3475(2-114); 48-3482(2-I41); 48-
Beckwith, F., 644(2-I) 35I9(1-I); 48-3614(1-I); 49-473(3-3); 49-
Belanger, C. P., 622(2-I2); 747(2-12); sn(2-I2) 4347(2-I); 49-4371(1-1); 49-8029(6-21); 49-
Belem, R. P. et al., 633(7-36a); 771(7-20); 1833 8io9(7-56a); 49-83 1(3-1); 49-8312(7-69);
(3-I ); 1926(3-I5); 1991(7-20); 1994(7-20); 49-8453(7-69) ;49-8500(2-69a) ;49-8547(3-3);
1996(7-54); 2005(7-54); 2010(7-20); 2I60 50-o074(2-27a); 50-8675(2-I35); 50-9752
(I-I); 2165(I-I); 2512(1-I); 2756(7-37); (7-56b); 50-9809(2-95); 50-9895 (2-52);
2837(6-25); 2846(6-25); 2948(6-51); 2978 50-9963(2-I8); 50-IOI78(7-56b); 50-10223
(6-25); 3082(2-86); 3084(6-25); 3094(7-37); (7-63); 50-10233(2- 14); 50-I06I2(7-50a);
3130(2-23); 3I32(6-I7); 3136(2-23); 3139 50-io634(2-27a); 50-o0690(2-27a); 50-10723
(6-I7); 3144(2-23); 3146(6-17); 3165(2-23); (2-69a); 50-I0749(6-3a); 50-10758(6-41);
3I86(2-9oa); 319I(2-23); 3201 (7-20); 3203 50-10909(7-20); 5I-12711(7-25); 51-12968
(2-88a); 3207(7-20); 3209(7-88a); 3211 (7-27); 5I-13371(7-56a); 5I-135I6(7-56a):
(2-146); 3227(6-25); 328 (6-255) 3310(7-I3); 51-i38o3(7-56a); 51-13885(2-122); 51-13918
3315(6-25); 3341(2-25); 3837(7-31). (7-56a); 52-I4472(2-37); 52-I4636(2-69a);
Bello, 6I78(2-43a) 52-14653(7-15); 52-I4661(2-93); 54-16684
List of Exsiccatae369

(7-56b); 54-16850(2-135); 54-16957(6-2); (2-88a); 4862(2-52); 4872(7-56a); 5111

54-17013(2-114); 54-I7I60(3-I); 54-I7342 (7-56a); 5140(7-56a)
(7-25); 54-1736i(7-i8); 54-I7520(7-56a); Brito, 37(7-36a); 37-I I (7-36a)
54-17631(7-2); 55-18691(7-18); 55-I8730 Britton, E. F., 6587(1-I)
(7-49a); 55-18736(7-18); 57-I9453(7-5oa); Britton, N. L., et al., 53(1-I); 89(1-I, 7-26);
57-I9589(2-I35); 57-I9664(7-20); 57-I9845 123(1-1); 124(1-1); I87(7-36a); 211(1-I);
(7-56a); 57-199I9(6-4); I9980(2-43b); 57- 274(2-I); 286(1-I); 423(7-56a); 528(1-I);
20068(7-5oa); 57-20069(7-56b); 57-20074 614(3-I); 69I (2-26b); 725(7-25); 1329(7-74);
(7-i8); sn(i-i) I571(I-I); I632(I-I); 1679(7-36a); 118I
Blain, J., 25(7-22) (2-69a); 2005(1-I); 2095(7-25); 2135(1-I);
Blake, S. F., 7553(7-56b); 7840(1-1) 2284(7-83); 2293(7-36a); 2435(7-83); 2516
Blanchet, J. S., 46(7-I8); 215(6-32); 216(2-43a); (I-i); 2692(1-I); 2709(1-1); 2435(7-83);
328(7-I8); 1254(1-I); 1415(I-1); 1508(7-13); 2814(7-36a); 3524(7-36a); 4176(7-36a); 4731
I822(I-I); 1847(7-I8); 2299(I-I); 2775 (7-36a); 6277(7-36a); 6504(I-I); 7274(7-83);
(6-20); 3200(2-84); 3209(6-32); 3290(2-126); 7463(7-36a); 7818(1-1); 9944(7-36a); 10309
3387(6-25); 3568(7-64); 3975(7-64) (I-i); 10323(7-36a); 14238(7-22)
Blanco, C., 30(2-88a); 33(2-77); 4I(2-88a); Brizicky, G. K., et al., 435(1-I)
46(7-56a); 51(7-20); 63(7-20); 75(2-88a); Broadway, W. E., 57(2-85); 346(7-25); 428(7-25);
I95(2-77); 199(7-74); 215(6-32); 254(2-77); 565(7-75); 814(7-25); 1024(1-I); 2169(1-I);
265(6-2); 29(2-88a);294(2-
77); 357(2- 13); 275I(7-25); 2884(2-I); 3577(7-74); 4025
361(2-109); 413(7-16); 453(7-56a); 474 (7-74); 4070(2-69a); 4133(7-56a); 4409
(7-16); 481 (7-16); 484(3-3); 591 (7-7) (2-I45); 4457(7-56a); 4462(2-145); 4466
Blanton, F. S., 6433(2-1) (2-69a); 4483(2-12); 5583(7-25); 559I
Blodgett, J. L., sn(i-i) (7-56a); 6160(3-1); 6445(2-69a); 6469(7-36);
Blume, C. L. von, sn(5-I) 6619(7-56a); 6634(2-69a); 6680(i-I); 6761
Bockerman, W., I40(7-56a); 81 (7-54) (2-27b); 6844(7-56a); 6905(7-75); 7549
Boechat, 21(6-26) (7-56a); 7659(I-I); 7704(7-56a); 8014(1-1);
Bojant, H., sn(i-i) 8977(7-36a); 9373(7-56a); sn(i-i)
Boldingh, I., 88oB(I-I); 2277(7-36a); 2290B Brooks, R. L., 12257(3-I); 12473(3-I); 12481
(i-i); 3871(2-135); 3883(2-I35) (7-74); I2484(2-69a)
Bond, F. E., et al, 58(i-i); 108(7-27) Brown, C. S., sn(7-22)
Bondar, G., 133(6-26); 1863(6-31); 2355(6-26); Browne, P., sn(i-i)
2511 (6-26); 3012 (6-26) Buchtien, 0., 741(2-55); I721(2-2)
Boole, 114I(2-I5) Budowski, G., 40(2-134); 40-I(2-134); 99(3-I)
Bordallo, A., MG 22634(7-56a) Bunting, G. S., et al, 717(2-45); 1191(7-36a)
Borgesen, F. C. E., sn(i-i) Burch, D., et al, I321(7-56b)
Botanic Garden Trinidad, i439(2-69a); 2059 Burchell, W. J., 3235(2-126); 3694(6-46); 5075
(i-i); 2060(1-I); 2061(7-25); 2062(7-36a); (2-41); 5130(2-4I); 5964(7-54); 5969(7-20);
3677(I-I); 5700(2-27b); 6420(2-86); 10756 6331(7-66); 6416(7-66); 6571(7-66); 7079
(7-25) (7-54); 7I85(2-27b); 7369(2-I14); 7398
Bourgeau, 3124(2-69a) (7-54); 7494(7-70); 7620(7-20); 7638(2-37);
Bowie, J. et al, 572(7-36b); sn(7-64); sn(7-85) 9I 0(7-56b); 9I43(7-56a); 9278(7-56b);
Box, H. E., 658(1-1); 1747(7-32) 93i6(2-43a); 9455(2-141); 9625(2-57); 9629
BoyanJ., 9(2-28); 39(2-8); 48(2-I 12); 69(7-76); (2-42a); 9778(2-57); 9870(7-20)
70(6-7); 75(2-69d); 88(2-77) Burger, D., 21(2-117)
Boyan, R., 28(2-42);
30(7-77); 44(2-112); 53 Burger, W. et al., 4066(2-45); 4131 (7-56b)
(7-21); 92(2-III); 23I(7-56a) Burgess, A. B., 718(I-I); 725(1-1); 768(1-I)
Bozeman, J. R., 9434(2-1) Busey, E. A., I(I-I)
Brace, L.J. K., 4569(1-I); 6980(I-I) Bush, B. F., 367(2-I)
Brade, A. C., 860(7-54); 7462(3-16); 11007(6-25); B. W. (Boschwezen Suriname), 11(2-130);
12125(7-85); 12758(6-25); I8344(6-25) 32(2-63); 49a(6-I); 5I(2-27a); 56(2-124);
Brass, L.J., 14991(2-1); 20404(1-I); 20509(1-1); I32(2-135); 525(2-135); 547(2-135); 628
20539(1-1) (2-o08); 1001(2-135); Ii36(2-88a); 1193
Bravo, H., 385(2-15) (2-69a); I203(7-50b); 1226(6-i); 1300
Brenes, A. M., 19I2(7-56b); 3606(7-56b); 4183 (2-130); 1301(2-66); I333(6-I); I339(6-I);
(7-56b); 5028(7-56b); 12I700(2-I5); I2170a I446(2-27a); 1460(2-130); 1496(2-66); 1507
(2-15); 12250(1-1); I4927(7-56b); 16918 (2-I24); 1509(2-66); 1538(2-69a); 1598
(7-56b); I7003(7-56b); 17009(7-56b); I8889 (2-69a); 1616(2-130); 1668(7-22); 1699
(7-56b); 20284(7-56b); 20323(7-56b); 20463 (2-108); I729(2-88a); 1738(2-108); 1748
(7-56b); sn(7-56b) (2-108); I755(6-i); 1916(2-124); I970
Breteler, F. J., 3449(7-36a); 3730(7-22); 3748 (2-69a); 2039(2-77); 2194(2-134); 2257
(7-i6); 3850(2-109); 3872(7-56a); 3927 (2-69a); 2302(2-124); 2312(2-69a); 2319
(2-43a); 4592(7-36a); 4689(7-36b); 4720 (7-56a); 2382(2-135); 24I8(2-88a); 2523
370 Flora Neotropica

(2-69a); 2531(6-2); 2604(2-69a); 2644 (7-56b); 2449(7-2I); 2556(7-21); 2557

(7-56a); 2658(2-130); 2686(2-130); 2729 (2-129); 2569(2-129); 2573(2-69a); 282I
(2-130); 2806(2-130); 2813(2-88a); 2817 (2-129); 2822(6-3b); 2827(7-21); sn(7-3o)
(2-135); 2818(2-130); 2825(2-66); 2881 Carleton, M. A., 629(1-I)
(2-135); 2882(2-130); 2906(2-130); 2909 Carlson, M. C., 1053(1-I); 1122(1-1)
(2-66); 2989(2-69a); 3007(7-49b); 3027(6-I); Castellanos, A., 25(2-I2); 523(2-43a); 22867
3033(3-I); 3060(2-88a); 306i(7-49b); 3067 (2-44); 22975(1-1); 22986(7-18); 23005
(2-134); 3080(6-I); 3098(2-130); 3126 (2-44); 24388(7-10)
(2-135); 3I74(3-I); 3352(2-27a); 3367 Castillo, A., 24(7-22); 47(7-56b)
(7-56a); 337I(6-1); 3392(2-88a); 3396 Castillo, E., sn(6-47)
(2-69a) ;3476347(6-);3410(2-24); 3463(2-33); Cavalcante, P. B., 222(6-4); 223(7-5oa); 234
3527(3-I); 3557(3-I); 358I(6-I); 3624 (2-10); 470(2-39); 472(7-56b); 523(7-56a);
(2-134); 3632(2-88a); 3662(2-99); 3687 623(2-140); 739(2-109); 769(2-27a); 1376
(2-63); 3768(2-130); 3791(2-112); 3817 (6-35); I497(2-69a); I669(6-3a); 1726(6-3a);
(2-75); 3824(2-134); 3860(2-88a); 3890 1746(2-57); 2041(7-24)
(7-25); 3900(7-50b); 3920(2-135); 3937 Cazalet, P. C. D. et al, 5104(7-35); 7690(6-36);
(2-69a); 3976(7-5ob); 3999(2-o08); 4018 7760(6-36)
(3-I); 4058(2-I24); 4106(2-27); 4307(2-63); Chaffanjon, J., 322(6-3b)
4351(3-I); 4356(7-25); 4366(2-135); 4376 Chagas, J. et al, I8(7-56b); 25(7-56b); INPA
(7-5ob); 4396(2-69a); 4422(3-I); 4479 I99(6-4); INPA 20I(2-69a); INPA 246
(2-I24); 4687(2-135); 4806(2-135); 4811 2-124); INPA 355(2-26); INPA 428(7-56b);
(2-27a); 4929(3-I); 5039(2-124); 5I05(6-I); INPA 784(6-9); INPA 865(6-15); INPA
5149(2-124); 5338(7-42); 5346(2-77); 5384 882(7-2); INPA 952(2-60); INPA 983
(2-135); 5385(2-124); 5387(2-130); 540I (6-29); INPA I252(7-56a); INPA 1311
(3-1); 542I(7-56a); 5427(6-2); 5478(7-25); (7-73); INPA 1457 (7-56b);INPA 1496
5529(6-42); 5659(I-69a); 5863(2-130); 5882 (6-i6); INPA I827(7-56b); INPA 1956
(2-88a); 6008(2-124); 6oI8(2-27a); 6034 (6-15); INPA 2137(6-15); INPA 3309(6-41);
(2-124); 6Io6(2-27a); 6108(2-124); 6113 INPA 3438a(2-26); INPA 3438b(2-62);
(2-88a); 6156(2-130); 6157(6-I); 6167(7-42); INPA 3470(2-140); INPA 3576(6-I I); INPA
6I8o(2-69a); 6184(2-63); 6i85(6-i); 6i86 4206(6-9); INPA 4246(6-41); INPA 4299
(2-I21); 6243(7-5ob); 6263(2-108); 6303 (6-9); INPA 5549(2-82); INPA 5893(6-41);
(7-73); 6350(6-18); 6455(2-108); 6457 INPA 6730(7-56b); INPA 7218(7-56b)
(2- 12); 6462(3-4); 6578(7-25); 6599(3-4); Chamisso, L. A. von, sn(7-8i)
6635(6-I); 6693(2-69a); 6761(6-I); 6777 Chanek, M., 213(7-22)
(3-4); 6796(2-69a); 6816(2-130); 68i8 Chapman, A. W., 840(1-I); I949(2-I); sn(2-1)
(2- 17); 68:34(7-2); 6846(2-88a) Chardon, C. E., 247(6-3b)
Byron (W. de Albuquerque) et al, 67-34(6-15); Chase, A., 7808(6-20)
67-57(2-142); 67-59(7-56a and 2-39); INPA Chickering, A. M., 50(I-I)
5566(6-34) Claussen, P., I47(7-81); 255(7-20); 256(6-27);
1097(7-54); 1098(7-81); 2023(6-25); 2056
Calder6n, S., 203(1-1); 272(6-47); 623(I-I); (7-36b); sn(7-2o); sn(7-8I); sn(6-27)
902(2-15); 2030(2-45) Clemente, Bro. A., 2953(7-36a); 5242(7-36a);
Calkins, W. W., sn(I-I) 5917(7-36); 6663(7-36a)
Campbell, E. J. F., 23(7-56b); 478(7-36a); Cobra, L. Q. et al, 43(7-56); 174(6-27); 296(7-20)
sn(7-22); sn(7-56b); sn(2-15); sn(6-47) Coche, 3132(2-I)
Campos, J. F., 30(2-41) Codazzi, L. et al, 9600(6-3b); sn(6-3b)
Campos, S. M., 62(6-27) Coelho, D. & L. et al, 9(2-83); 14(4-I); INPA
CAP, 35(7-7); 93(7-73) 1259(6-9); INPA
1286(2-11); INPA 1578
Capanema, RB 19199(7-36b); RB I9200(6-25); (3-4); INPA I714(7-9); INPA I733(6-4I);
RB 19023(7-70); RB I9204(6-35); RB 19207 INPA 1837(7-I); INPA 2907(6-29); INPA
(7-36b) 29II(2-26); INPA 2966(2-II); INPA 2974
Caprice, F., sn(7-25) (2-99); INPA 2998(2-26); INPA 301 (6-9);
Capucho, P., 347(2-52); 384(4-3); 460(6-3a); INPA 3057(2-26); INPA 3057b(2-62);
587(7-73) INPA 3486 (7-28); INPA 3531(7-2); INPA
Carcerelli, C., 62(2-69a); 76(2-69a); I 3(7-36b) 3885(7-28); INPA 3946(2-64); INPA 3953
C,rdenas, M., 3012(7-54) (6-29); INPA 3955(6-4); INPA 3975(7-28);
Cardona, F., 389(2-69a); 840(7-30); 854(7-30); INPA 4062 (2-I24); INPA 4099(3-4); INPA
1174(3-4); 1176(3-4); 1177(2-129); 1210 4137(6-34); INPA 4222(7-I); INPA 4261
(7-27); 1254(2-27b); I370(7-6); I373(7-20); (2-99); INPA 5I 78(2-43b); INPA 5984(7-4 );
I431(7-56b); 1479(2-124); I665(7-27); 1736 INPA 5985(7-41); INPA 5986(2-62); INPA
(7-25); 1756(7-30); I771(7-30); I789(7-30); 6004(2-I28);INPA 65I9(2-88a); INPA 7259
1870(7-49); I927(7-56a); 2600(7-25); 2095a (6-37); INPA 7624(2-27a); INPA 15883
(7-56a); 2203(2-75); 2303(2-89); 2387 (7-20); INPA 16712(7-20); INPA 16714
List of Exsiccatae 371

(7-54); INPA 20880(6-29); INPA 21129 Cuming, H., 1 I09(2-45); I272(2-I5); 1289(1-I)
(6-41) Curran, H. M., 33(7-56b); 9oM(7-56b); I39(3-7);
Collenette, C. L., I45(7-54); 472(7-56b) 146(2-I5); 231(6-20); 256(1-I); 303(2-45);
Collins,J. F., sn(I-I) 847(1-1); 913(3-7); I784(7-56b); 1853
Combs, R., 436(1-I) (6-3b); I870(7-56b)
Conrad, H. S., sn(2-I) Curtiss, A. H., 260(7-22); 280(1-I); 727(2-I);
Conservator of Forests, British Honduras, 95 728(1-I); 4212(2-I); 4905(2-1); 5779(2-I);
(7-22) :?'45(I-I)
Constantino, D., RB 29814(1-1) CdtC-r,H. C., 8264(2-44); 8375(2-44)
Conzatti, C., 3262(1-I); 4402(1-1); 4455(2-45);
4473(I-I) Dal:lgren, B. E., 876(2-27a); 912(7-56b); 932
Cook, C. D. K., 151(7-56b) (2-44); 951(7-20); 972(7-56b); 986(6-20);
Cook, O. F. et al, 22(6-47); 68I (6-47) sn(6-2o); sn(6-3a); sn(2-II); sn(2-6); sn
Cooley, G. R. et al, 2632(2-I); 4860(2-I); (I-I)
8365(1-I); 8735(7-32) Damazio, L. B., 1594(7-20); RB 47980(2-I43);
Cooper, G. P. et al, 256(2-15); 279(7-36a and RB 47985(2-41)
5-I); 286(7-36a) Daniel, Bro. J., 477(2-48)
Cooper, J. G., sn (I-i); sn(i-i) Daniels, A. G. H. et al, 784(7-74)
Cordeiro, E. et al, 87(2-17); I45(2-43b); i6i Danin,J., sn(I-I)
(6-3b); 169(7-73); 225(2-17); 23I(7-15); Dannouse, L., 6653(2-13); sn(2-I3); sn(3-I)
309(2-43b); 384(7-56); 400(7-56a) Davis, T. A. W., 87(6-6); I43(3-I); I50(2-108);
Corner, E.J. H., I 10(7-50) 259(2-18); 264(2-93); 280(2-109); 288
Correll, D. S., 5494(2-I); 6040(1-1); 6326(2-1) (2-88a); 239(2-135); 315(2-108); 350(2-35);
Cowan, R. S. et al, 1477(7-74); I597(1-I); 1789 352(7-1is); 369(2-88a); 383(2-69d); 387
(6-42); 1917(2-I35); 1948(2-I35); 1922 (6-42); 426(2-78); 429(6-15); 433(2-108);
(6-42); 201 (7-5I); 2140(2-I35); 2173(2-95); 437(6-7); 452(2-93); 587(2-66); 590(7-7);
2180(3-10); 2234(7-77); 3I084(7-42); 3 149 sn, F.D. 2309(?-Jo)
(7-43); 31237(7-43); 31383(7-43); 31439 Dawe, W. T., 429(1-I)
(7-2); 31453(7-68); 31454(7-21); 31460 Dawson, E. Y., I4270(7-54); 14812(2-88); 15130
(7-25); 31487(7-56b); 38031(2-135); 38045 (7-66); 26500(1-1); 26617(1-I); 26761(1-I)
(i-i); 38212(2-69a); 38260(3-2); 38270 Deam, C. C., 57(I-I)
(2-14); 38527(7-24); 38586(7-56a); 38624 Degener, 0., 18770(1-1); 18839(1-1)
(7-24); 38648(7-18); 38781(2-109); 38847 De Jai Cruz, J. S., 990(I-I); 1069(7-25); 1195
(7-56a); 38854(6-7); 38857(7-56a); 38875 ( - ); 1283(2-8); 1635(1-I); I585(7-25);
(2-135); 38890(1-I); 39023(7-74); 3927I 1670(2-35); 1736(7-25); I775(2-I35); I776
(6-42); 39273(2-135); 39277(2-78); 39357 (2-78); 1777(2-I35); I825(6-7); I845(7-56a);
(7-56a); 39392(2-69a) i 33(7-56a); 2015(1-1); 2032(2-108); 2050
Cowell, J. F., 599(1-I) (2-135); 2107(6-42); 2144(6-15); 2215(6-42);
Cowles, H. C., 569(2-I); 5570(1-1) 2218(6-15); 2257(7-25); 2370(3-I); 2377
Craighead, F. C., 645(I-1) (2-109); 253I(7-56a); 2538(7-25); 2542
Crans, sn(2-I ) (2-69a); 2566(7-56a); 2648(2-78); 2661
Croizat, L. C. M., 54(7-56b); 55(7-56b); 167 (2-135); 2707(7-25); 27i5(2-88a); 2720(3-I);
(7-56b); 203(7-40); 269(7-56b); 282(6-4); 2739(2-124); 2780(3-I); 2804(7-25); 2857
669(7-56b); 1018(7-2); I04oa(2-69a); sn (3-I); 2858(2-88a); 2865(2-78); 3031(2-8);
(7-25) 3408(2-89); 346.(6-7); 3484(2-8); 3499
Cronquist, A., 5332(2-I) (2-43a); 3656(1-I); 3748(7-56a); 3828
Crueger, H., 2(7-56a); I05(2-69a); 205 (2065) (2-135); 3829(2-69a); 3831(7-25); 3932
(2-145); 298(7-25); 300(7-56a); 301(7-36a); (I-I); 3963(2-27a); 4005(7-56a); 4174(7-25);
sn(7-56a); sn(7-75); sn(2-I2); sn(2-86); 4226(1-I); 4320(7-56a); 4349(2-109); 4384
sn(3-I) (2-109); 4397(7-56a); 4411(2-69a); 4419
Cuatrecasas, J., 4097(7-56b); 5336(6-48); 6837 (2-69a); 4443(1-I); 4524(2-69a); 4539
(2-137); 6909(7-55); 6913(6-2); 6915(7-40); (2-27a); 4591 (2-8); 4595(3-I)
6917(3-6); 6983(6-36); 7054(2-121); 7067 Delgado, E., 88(2-145); Io7(7-36a); I20(2-106);
(6-2); 7112(7-55); 7I32(7-56b); 7245(2-88a); I26(7-36a); 192(3-7); 399(3-7); 434(2-o06);
7304(2-140); 7516(7-25); 7817(7-40); io865 445(2-106)
(2-45); 14322(7-IO); 1533I(2-4); I578i Demaree, D., 33429(2-I); 35889(2-I)
(7-148); I5835(2-34); I5924(7-10); I5929 Dept. Lanbouw, Suriname, 244(2-134)
(7-o1); 15941(2-5I); 16177(1-1); 16262 Der. -:, sn(6-3a)
(7-56b); I7431(2-34); 17690(7-10); I7740 Desvaux, A. N., sn(8-2); sn(3-I)
(2-5); 18370(7-86); 19908(7-1O), 19925 de Vriese, sn(2-I35)
(7-56a); 21179(2-46); 2 I79a((246); 21183 De Wolf, G. P., 1779(2-I)
(7-80); 21234(2-58); 21455(2-51); 21546 Diogo, J. C., 301(6-20)
(2-58); 22251(2-4); 24219(2-5); 24876(3-7) Dodge, C. W. et al, I6524(7-36a); I73I6(7-36a)
372 Flora Neotropica

Dolbolmo, J., RB 862 19 (7-85) MG 7344(6-3b); MG 7365(7-40); MG 7372

Don, G., 32(7-56b); 33(7-56b); 53(7-36b); sn (3-4); MG 7598(7-6); MG 76i9(7-56b);
(3-4) MG 7623(2-43b); MG 7890(7-24); MG 7919
Dorrien-Smith, 132(7-54) (6-37); MG 7958(2-75); MG 7980(7-24);
Drake, I(I-I); sn(2-93); sn(2-28) MG 8034(7-50); MG 8088(7-18); MG 837I
Drouet, F. E., 2108(7-56b); 2455(1-I); 2532 (2-43a); MG 8373(7-56a); MG 8374 (7-73);
(7-56b); 2676(1- ) MG 8409(6-3a); MG 8490(7-9); MG 8514
Drummond, et al, 2614(7-89) (2-69a); MG 8531(7-56b); MG 8536(6-3b);
Drushel, J. A., 4649(2-I) MG 8536b(6-29); MG 8557(7-1); MG 86o1
Duarte, A. P., 82(7-54); 99(6-27); 823(3-I6); (2-69a); MG 8630(6-7); MG 8651(2-69a);
1435(7-I8); 1447(7-20); 1451(7-56b); 1508 MG 8879(6-37); MG 8915(7-56a); MG 8928
(6-65); 2251(2-I44); 2967(2-43a); 3269 (6-3a); MG 8929(2-134); MG 8948(7-56a);
(6-27); 4839(2-70); 4958(7-85); 4981(6-25); MG 8961(2-52); MG 8979(2-28); MG 8997
5697(2-54); 583I(2-43a); 5980(6-26); 60I4 (2-93); MG 9008(7-56a); MG 9052(2-75);
(7-36b); 6065(1-1); 6200(1-1); 6393(7-20); MG 9070(7-56a); MG 9087(2-69a); MG
6645(7-20); 6880(6-41); 7129(2-121); 7131 9125(6-29); MG 9899(6-2); MG 9917(7-18);
(7-56a); 7132(7-24); 7133(2-27a); 8005 MG 10242(4-1); MG 10243(6-3a); MG
(6-26); 8029(7-37); 8057(2-25); 8244(2-41); 10248(2-27b); MG 10249(4-1); MG 10257
8319(3-16); 8965(7-20); 9255(6-27); 9259A (7-56b); MG 10461(6-29); MG 10492(2-27b)
(7-20); 9328(7-56b); 9369A(7-54); 9393 MG 10672(2-129); MG 10884(3-4); MG
(7-54); 9747(7-31) 10910(7-50); MG 10912(2-52); MG 10922
Duchassaing (de Fontbressin), E. P., sn(2-i3) (2-56a); MG 11051(3-4); MG III92(7-56a);
sn(i-I); sn(7-36a) MG 11194((7-8); MG 11288(2-135); MG
Ducke, A. et al, 2(7-56b); I6(2-43a); I9(6-41) 11313(7-5oa); MG I1397(2-37); MG I1514
33(2-43a); 43(6-41); I56(6-ii); I84(6-I5); (6-37); MG 11530(7-8); MG I1580(2-27a);
I95(6-i6); 248(2-82); 293-1(2-128); 293-II MG 11630(7-9); MG 11658(6-29); MG
(2-I28); 303(2-62); 307(2-36); 385(7-I); 406 II665(6-3b); MG II708(2-69a); MG 11717
(6-34); 407(6-29); 464(6-9); 465(7-9); 488 (7-56a); MG I 1758(2-52); MG 1 794(2-27a);
(2-60); 534(3-4); 579(6-3b); 580(2-69a); MG 11939(7-9); MG 11969(2-37); MG
66I(6-13); 662(6-15); 705(7-28); 778(6-2); 12026(2-37); MG I2350(7-56b); MG 12563
779(6-4); 78i(3-1); 866(6-20); 908(6-37); (2-135); MG 12587(6-2); MG 12614(6-2);
921(6-28); 924(6-29); 976(7-4I); I04I(2-9); MG 14858(2-99); MG 14996(2-134); MG
I249(6-3b); 1251(7-27); 1265(3-I); I267 14998(2-99); MG 15019(2-75); MG 15142
(6-50); 1297(6-9); I329(6-3b); 1330(7-4); (2-136); MG 15173(2-21); MG 15183
I34i(7-25); I347(3-4); I54I(6-44); 1544 (7-36a); MG 15244(3-4); MG 15288(2-27b);
(3-4); 1545(6-44); I619(3-3); 1729(3-4); MG 15299(7-50b); MG I5305(3-3); MG
I742(6-3a); 1745(7-8); 1755(6-39); I824 I5336(6-4); MG 15438(6-I5); MG 15469
(3-4); I878(7-40); I910(2-24); 2015(6-50); (6-50); MG 15475(2-57); MG 15484(6-50);
2113(6-49); 2127(6-I3); 2137(2-114); 2142 MG 15527(2-8); MG 15529(2-63); MG
(6-29); 245(3-4); 249(2-69a); 2154 (7-5oa); 15553(6-50); MG 15566(2-27a); MG 15584
2155(7-I8); 2165(2-128); 2208(6-29); 2209 (7-I8); MG 15632(6-2); MG 15644(3-3);
(2-6o); 2211 (2-99); 22 12(2-27b); 2213(6-15); MG 15646(7-65); MG 15684(6-29); MG
22I7(6-3b); 2232(6-13); 2233(2-88a); 2277 15752(2-I 4); MG I5799(2-57); MG I5806
(7-I); 2279(3-2); 2381(7-18); 2418(3-4); (3-3); MG 15844(6-29); MG 15967(2-39);
2948(6-3a); 2950(6-43); MG 336(7-56b); MG I6223(2-43b); MG 16225(2-97); MG
MG 434(1-1); MG 525(7-56b); MG 614 I6274(7-14); MG 16291(7-20); MG 16299
(7-56b); MG 684(7-56b); MG 719(7-56b); (2-52); MG 16319(6-4); MG 16341(2-57);
MG 727(4-2), MG 764(7-20); MG 807 MG 16367(6-I5); MG 16388(6-20); MG
(7-56b); MG 1102(2-44); MG 1364(4-I); 16415(2-52); MG 16418(2-39); MG 16459
MG I429(7-56b); MG 1518(2-II); MG 1603 (7-I); MG I6590(2-27b); MG 16592(2-27b);
(2-44); MG 1616(2-II); MG 1700(4-1); MG 16648(3-4); MG 16663(7-30); MG
MG I956(7-56b); MG 1966(7-24); MG 2508 17055(6-29); MG I7124(7-36a); MG 17178
(1-I); MG 2898(2-99); MG 2907(7-50a); (6-4); MG 17179(6-4); RB 11051(3-4); RB
MG 2983(2-I29); MG 3061 (7-18); MG 3460 11658(6-29); RB 15037(3-4); RB 15057
(7-56b); MG 3690 (2-43a); MG 3729(7-5oa); (7-69); RB 15066(7-18); RB 15068(7-18);
MG 4732(7-56a); MG 4745(7-2); MG 4760 RB I5o69(7-36a); RB 15078 (7-9); RB 15079
(7-25); MG 4774(7-56a); MG 4829(2-69a); (7-8); RB 15098(6-2); RB 15119(6-4); RB
MG 4881(7-24); MG 4924(7-3a); MG 4945 15120(6-I6); RB 15133(6-29); RB 15136
(2-114); MG 6756(7-6); MG 6775(7-56a); (6-21); RB 15138(6-43); RB 15157(2-82);
MG 6785(7-40); MG 6853(7-62); MG 6902 RB i5I66(2-69a); RB I5I67(2-69a); RB
(7-5oa); MG 6904(2-27a); MG 6905(2-72a); 15173(2-114); RB I5i82(2-69a); RB 15187
MG 7III(2-69a); MG 7138(7-56b); MG (2-57); RB 15192(2-43b); RB 15216(2-27b);
7153(7-I);MG 7181 (7-24); MG 7205(7-56b) RB 15244(3-4); RB 16385(4-3); RB 17113
List of Exsiccatae 373

(7-42); RB I7975(7-5); RB 18224(4-3); RB Duckett, F., sn(2-i); sn(i-I)

I8225(3-2); RB 18226(3-I); RB 18227(6-2); Dugand, A. et al, 369:816(2-45); 3481(1-i);
RBi8229(6-4); RB 18230(6-4); RB 18231 6311 (2-45); 6314(2-45)
(6-37); RB I8233(7-30); RB I8234(7-36a); Duke, J. A., 4673(7-56b); 4896(7-36a); 5826
RB 18235(7-24); RB 18236(7-14); RB 18237 (7-36a); 5827(7-36a); 7009(7-83); 8o 2
(7-27); RB 18239(7-20); RB 18240(7-65); (7-38); 8o53(2-88a); 8093(2-88a); 8109
RB 18242(7-2); RB 18243(7-2); RB 18246 (7-22); 83I1(2-88a); 8132(7-22); 8201
(7-56a); RB 18337(3-3); RB I8755(2-43b); (7-36a); 859 (7-36a); 9275(7-36a); 9284
RB 18787(2-8); RB 18788(2-8); RB 18789 (7-36a); 9515(7-56b); 9659(2-3); 97I6(I-I);
(2-8); RB 1879I(2-39); RB I8795(2-43a); I0206(7-36a); I509(2-3); I 679(2-3); 11883
RB i8796(2-43a); RB 18808(2-129); RB (7-56b); I4143(7-56b)
18809(2-52); RB 18810(2-69a); RB 18812 Dumortier, F., sn(6-2)
(2-142); RB 18813(2-86); RB 18814(2-142); Dupuis, sn(7-36a)
RB 18815(2-130); RB 18816(2-75); RB Duque-Jaramillo, J. M., 2316(6-44)
I8817(2-69a); RB 18818(2-136); RB 18820 Dusen, K. H., 7441(7-85); 8222a(7-85); 8653
(2-69a); RB 18821(2-114); RB 18822(2-69a); (7-85); O1114(7-85); 13536(7-85); I6150
RB 18823(2-69a); RB i8824(2-43a); RB (7-85)
18825(2-81); RB 18830(2-69a); RB 19175 Duss, A., 153(2-13); 154(1-2); 363(7-36a); 1902
(7-82); RB I9750(6-9); RB 19754(6-3b); (2-85); I904(7-32); 1905(7-36a); 1906(I-I);
RB 19755(6-2); RB 19756(6-15); RB 19757 2212(1-I); 22I4(7-36a); 2730(1-I); 2731
(6-40); RB 19758(6-19); RB 19760(4-3); (i-I); 2868(2-85); 3255(7-32); 3255a(7-32);
RB 1976I (7-69); RB 19763(7-40); RB 19764 3476(I-2) ;3633(I-2) ;4147(7-32) ;4209 (7-32)
(7-42); RB I9766(2-69a); RB I9770(2-69a); Dutra, J., 300(7-85)
RB I9771(2-69a); RB 19773(2-141); RB Dwyer, J. D. et al, I399(7-56b); 1494(5-I);
19774(2-97); RB 19755(2-69a); RB 19777 I872(2-88a); 1973(2-88a); 7085(2-88a); 7259
(2-121); RB 19779(2-97); RB 19780(2-129); (7-38); 8o6o(5-1)
RB 19784(2-20) ;RB 19786(2-69a); RB 19787
(2-135); RB 19788(2-142); RB I9789 Earle, F. S., 655(7-22); 702(1-I); sn(2-I)
(2-69a); RB I9792(2-93); RB I9793(2-97); Eaton, A. A., 333(1-I); II03(I-I)
RB 19796(2-60); RB 19797(2-27a); RB Edwards, E. P., 15(2-45)
19798(2-43a); RB I9799(2-43a); RB 19800 Eggers, H. von, II3(2-69a); 134(1-I); 618(i-2);
(2-43a); RBI98oi(2-27a); RB 21325(2-63); 639(7-36); 742(2-85); 753(2-I3); 849(7-83);
RB 23586(7-9); RB 23587(7-2); RB 23589 937(7-25); 986(1-2); I051(3-I); 1060(2-13);
(6-I5); RB 23590(6-9); RB 23591(6-3b); 1073(2-85); 1084(1-I); 1092(7-25); 1183
RB 23592(6-41); RB 23593(6-41); RB 23594 (7-83); 1357(7-25); 1362(3-1); 1439(2-69a);
(2-52); RB 23595(2-27b); RB 23596(2-26); I458(2-69a); I549(7-36a); I64o(I-I); 2456
RB 23597(2-36); RB 23598(2-36); RB 23599 (i-i); 4060(1-I); 4665(7-36a); 5818(7-74);
(2-22); RB 23600(2-23); RB 23601(2-62); 5842(2-69a); 6722(1-I); 7145(1-1); 14218
RB 23602(2-60); RB 23603(2-10); RB 24181 (7-35); sn(7-25); sn(7-36a)
(3-4); RB 24182(3-13); RB 24183(3-I3); Egler, F. E., 39-200(1-I); 42-17(1-I); 42-224
RB 24987(6- I1); RB 24988(6-I ); RB 24989 (7-22); 42-296(7-22)
(6-9); RB 24990(6-15); RB 24991(6-28); Egler, W. A., 154(1-1); I70(7-56b); 260(2-135);
RB 24992(6-4); RB 24993(6-2); RB 24994 626(7-24); 828(2-17); 832(7-56a); 842
(6-2); RB 24995(6-34); RB 24996(6-4); (2-43a); 887(2-17); 925(6-9); 1258(6-13);
RB 25001(6-8); RB 25002(6-14); RB 25003 1354(7-56a); 465o6(7-56a)
(7-2); RB 25004(7-2); RB 25006(7-25); Eiten, G., 4605(7-18); 5379(7-I8); 5467(7-I8)
RB 25008(7-I); RB 25009(7-6); RB 25011 Ekman, E. L., 2583(7-36a); 2614(7-36a); 3250
(7-36a); RB 25012(7-9); RB 25013(7-5); (7-36a); 353I(I-I); 4003(7-36a); 4080
RB 25025(2-22); RB 25026(2-24); RB 25027 (7-36a); 580o1(-1); 9903(1-I); 10221(7-36);
(2-27a); RB 25028(2-43b); RB 25029 10796(7-22); 12146(7-22); 12205(1-I); 12247
(2-27b); RB 25030(2-119); RB 25031(2-60); (7-36a); I2477(I-I); 13330(1-I); I5775
RB 25032(2-79); RB 25033(2-69b); RB (i-i); 16362(7-22)
35185(4-I); RB 35553(2-62); RB 35554 Elias (de Paula), J., 244(7-56b); 3I3(2-94); 314
(6-29); RB 35555(3-4); RB 35557(6-36); (7-40)
RB 35558(6-29); RB 35559(6- I); RB 35560 Ellenberg, 2414(2-21); 2504a(2-45); 2504b(2-52)
(6-9); RB 35561(6-34); RB 35562(6-3b); Elmore, F. H., DI4(6-47); DI5(I-I); 645(1-I)
RB 35563(6-I5); RB 35564(2-20); RB 35565 Emmerich, M., 2906(2-41)
(2-19); RB 35566(2-49); RB 35567(2-128); Emygdio, L., 197(1-1); 1526(6-46); 1738(7-56b)
RB 35568(2-62); RB 35570(2-62); RB 3557I I845(7-56a); I869(7-56a); 2322(7-54)
(2-88b); RB 35573(2-82); RB 35575(2-112); Englesing, F. C., 131(7-56)
RB 35576(7-55); RB 35577(7-56a); RB Enriquez, 7291(2-88a); 7310(2-56b)
35578(7-56a); RB 35579(7-73); RB 35580 Erlanson, C. 0., 58(7-56b); 108(7-22); ii6
(7-9); RB 35581(7-20); RB 35583(6-39) (7-56b); 121(7-22)
374 Flora Neotropica

Erlanson,J. B. M., 78(I-I) Flores, R. S., sn(6-47)

Ernst, A., sn(2-24) Florschiitz, P. A., 640(2-135); 1520(7-56a)
Espina, R. et al, AI5(7-22); B2-A(3-7) Focke, A., sn(7-83)
Espinal, T. S., 1668(7-22) Focke, H. C., 455(6-42); 643(7-25); 1021(2-63);
Eugenio, J., 711 (2- I) I228(7-56a); 128I(2-I35); I368(2-27a)
Evans, A. W., sn(2-I2) Foldats, E., 3626(2-88a); 3633(2-I03); 3636
Ewan,J., 117117(1-I) (7-7I); 3709(2-103); 3750(3-I); 3857(2-103)
Forero, E. et al, P6423(7-56a); P702o(7-56a);
P705I(7-56a); P7o65(7-56a); P7129(7-56a);
Fairchild, A. S., sn(i-I) P7143(7-56b)
Falcao, J. I. A., et al, I I89(7-56b) Forest Department, British Guiana (Guyana),
Fanshawe, D. B., I5(1-I); I27(2-IO8); I33(2-78); 334(6-2); 6268(2-109); 6269(2-109); 6383
204(2-124); 242(6-I); 265(2-72); 280(3-3); (3-4); 6384(3-4); 6767(7-56a)
306(7-45); 313(7-20); 324(2-I24); 377(6-7); Forest Department, British Honduras, 19(7-23)
405(6-15); 505(7-56a); 584(6-6); 588(2-83); Forest Service, French Guiana, 22M(2-69a);
598(2-18); 6o7(2-69a); 6 (3)
(3-4); 633(2-35); 34M(3-I); 40(2-132); 46M(2-97); 53M
640(7-25); 650(2-72); 66I(2-88a); 670(3-4); (2-97); 56M(2-99); 62M(2-IO8); 64M
67I(2-77); 708(6-6); 709(6-6); 722(6-12); (2-117); 77M(2-io8); 132(2-97); I49M
722a(6-I2); 723(6-I2); 841(2-114); 84Ia (2-63); I69M(2-52); I92M(3-I); 251M(3-4);
(2-114); 902(6-7); 904(2-108); 937(2-II4); IOI6(2-I24); 1019(2-114); I23(2-69a);
944(2-72); 1000(6-42); IOI9(2-I12); I020 II6I(2-27a); II73M(2-97); 1174(2-124);
(2-II2); 1032(2-II2); 1047(2-69b); I059 II75(2-27a); I2II(6-I); I3I7(2-27a); 1321
(2-II2); 1078(2-120); 1086(2-35); 119 (6-15); I328(2-52); 5111(7-56); 6087(2-63);
(7-74); 1148(7-45); 1224(6-7); 1315(2-66); 6242(2-52); 7058(7-56a); 7110(6-I8); 7138
I333(6-I5); I336(2-69a); I358(2-69a); 1359 (6-i8); 7204(2-95); 7237(7-7); 7354(2-95);
(2-35); I420(2-69a); I495(2-6I); 1506(6-7); 7392(2-95); 7440(2-108); 7480(2-63); 7485
1555(2-57); 1564(2-28); 1674(6-45); 1678 (2-Io8); 7526(2-69a); 7630(3-I); 7676 (2-130);
(2-38); I680(2-68); 1681(2-68); 1700(2-35); 7697(2-117); 7700(3-I); 7726(7-20); 7730
1751(7-16); 1785(2-52); I803(7-I6); I98I (2-52); 7757(7-73); 7987(2-69a)
(2-69a); 2004(2-18); 2180(6-45); 2469 Ford, E. S. et al, 3887(2-I); 4416(2-I)
(2-69a); 2489(6-42); 2586(7-18); 2587 Forsstrom,J. E., sn(7-32); sn(I-I)
(2-43a); 2613(6-42); 2701(1-I); 2820(6-45); Fosberg, F. R., 20193(7-40)
2836(2-99); 2837(2-57); 2866(2-69a); 2946 Foster, M. B. et al, 1355(2-1); I356(2-I)
(2-114); 2952(7-I8); 3356(7-20); 34 7(2-35); Fox, W. B., 518(2-1)
3515(2-108); 3516(2-72); 23437(7-73); Frazao, A., RB 15511(7-85)
32436(6-42); 32566(6-42) Fredholm, A., 5251(2-I); 5438(2-I)
FAO-FYDEP, 21(2-I5) Freire, C. V., 73(7-36b)
Faria, R., I4(7-85) Freyreis, G. W., sn(7-64)
Farinias, M. et al, 475(7-21); 519(2-27a); 658 Friedrichstahl, E. R. von 256(6-47); sn(i-i)
(2-69b); 66I(2-27a); Fr6es, R. L. et al, 8(I-I); 2I(2-II4); 1735(2-I34);
Faris, J. A., 543(1-1) I8o9(7-56b); I85I(7-56b); I909(3-4); 1915
Feddema, C., 994(7-56b); 1832(1-I) (2-69a); I930(6-2); I933(7-56b); 1935(2-i8);
Felippe, G. M., 52(6-27) 1945(6-2); I96I(2-I8); I976(2-69a); 1982
Fendler, A., 38(7-35a); 107(1-i); 1019(7-25); (3-4); I992(I-I); 2004(2-I 14); 1533(7-56b);
1020(7-74); I020B(2-56b); I02I(7-37a); sn II643(7-56b); 11687(2-44); 11702(2-37);
(I-I) I 707(7-56b); II867(7-56b); 11877(7-20);
Fennell,J. L., 300(1-I); 374(2-I); 392(2-I) 11879(2-37); 11963(2- I); 12435(2-33);
Fernandez P., A., 244(1-I); 1950(7-2); 2005 20283(6-2I); 20377(7-63); 204I5(2-69a);
(7-56a); 2066(7-40); 2095(7-40); 2153(6-9); 20427(2-8); 20438(2-39); 20456(7-I8); 20892
2221(3-6); 2269(7-5); 2305(7-6); 2338(6-2) (7-40); 20910(2-61); 20984(6-3b); 21093
Ferreira, A. G. et al, 19(6-27); 86(7-53) (2-121); 21102(6-2); 2III2(6-29); 21128
Ferreira, E., 147(2-128); 57-I(6-41); 57-12(3-4); (7-55); 21133(2-22); 21204(6-9); 21272
58-320(7-56b); 87-57(6-7); INPA 5880 (7-49b); 21322(6-35); 21432(6-5); 2I485
(2-22); INPA 5912(2-62); INPA 5920(6-4) (6-36); 21495(2-39); 21548(6-2); 21590
Ferreyra, R., 4582(7-36a); 4603(7-56b); 4636 (i-i); 22061(2-27a); 22129(6-2); 22i68
(7-56b); 4953(7-56b); 4978(7-56b) (3-6); 22198(2-140); 22240(2-69b); 22257
Fiebrig, K., 1379(6-20) (7-2I); 22431(6-2); 22449(2-27a); 22510
Figueiras, M. L., 130(1-1) (2-75); 22544(7-I); 22549(2-81); 22556
Finch, R. G., 26(1-I) (7-56a); 22630(4-1); 22671(2-81); 22675
Fishlock, W. G., 22(2-85); 371(I-I) (6-4); 22694(6-3b); 22709(2-69b); 22730
Fittkau, E.J., et al, INPA 12816(2-134) (7-49); 22737(7-71); 22799(2-69a); 22840
Fleischmann, E., 353(7-21); 354(7-2 ) (7-30); 22863(2-27a); 22908(6-2); 22991
Flint, M. B., sn(2-I) (4-I); 22997(6-43); 23001(2-140); 23189
 List of Exsiccatae 375

(3-I); 23215(4-I); 23252(2-43b); 23272 Gandoger, A. M., 95(2-IO8); I 6(1-1)

(2-17); 23274(2-69a); 23319(3-2); 23329 Garber,A.P.,56(7-36a);82(7-83); sn(I-I);sn(2-I)
(2-88a); 23419(6-21); 23512(7-65); 23513 Garcia, L., sn(8-I ?)
(7-37a); 23738(3-4); 23833(6-35); 23856 Garcia-Barriga, H. et al, I 1710(7-22); 13312 ( -I);
(2-99); 23885(2-98); 23891(2-17); 23929 13441(I-I); 13782(2-140); 13863(2-26);
(2-69a); 24124(2-52); 24250(2-114); 24434 I41oo(2-27a); I4212(2-69b); I4282(6-2);
(7-53); 24553(2-134); 24632(7-56a); 24909 I4463(2-I24); 16090(7-40); 17111 (7-25);
(7-8); 24910(6-9); 25033(2-69a); 25040 I7145(2-47)
(2-I7); 25059(6-9); 25086(6-3a); 25148 Gardner, G., 370(7-85); 864(7-36b); 902(I-I);
(2-69c); 21173(2-17); 25I98(2-27a); 25204 992(2-II); 993(7-56b); 1056(6-20); 1149
(2-88a); 25226(2-99); 25228(7-56a); 25229 (2-25); 1150(1-1); I266(4-3); I284(6-20);
(2-88a); 25236(6-3a); 25240(2-I I9); 25244 I59o(7-56b); I59Ia(7-i8); 1592(2-44); I947
(2-129); 25271(2-27a); 25313(4-I); 25333 (7-20); 2559(2-I ); 2560(4-2); 2561(6-20);
(7-50a); 25353(2-57); 25442(2-69a); 25455 2562(7-I8); 2563(6-27); 2564(2-30); 2565
(2-69c); 25460(2-69c); 25462(2-27a); 25465 (7-54); 2836(2-80); 2837(7-20); 3138(7-70);
(2-69a); 25467(2-27a); 25484(7-5ob); 25556 3I39(4-4); 3137(3-I6); 3708(7-54); 4125
(2-129); 25612(2-I 4); 25622(2-114); 25745 (7-I8); 450I(7-81); 4538(6-27); 4539(2-30);
(2-69a); 25870(2-8); 25882(2-124); 25926 4540(7-70); 4541(7-81); 5448(2-114)
(2-63); 25958(2-69a); 26118(2-129); 26137 Garganta F., M. de, 1133(1-1)
(2-53); 26242(6-15); 26247(2-82); 26278 Garnier, F. A., 416I(I-I); sn(6-47)
(2-69a); 26295(2-114); 26300(2-62); 26303 Gates, sn(2-i)
(6-I5); 26336(7-24); 26376(2-69a); 26390 Gaudichaud, C., II3(2-43a); 868(7-85); I020
(2-8I); 26396(3-4); 26436(6-3b); 26451 (7-36b); 1022(7-82); o023(7-36b); 1170
(3-4); 26459(2-17); 26463(7-24); 26532 (2-1 I4)
(3-4); 26561(6-41); 26609(2-124); 26625 Gaumer, G. F., 23205(6-47); 24320(6-47)
(7-5ob); 26632(2-63); 26660(2-124); 26706 Gehrt, A., SPI2887(7-85); SP45851(6-20)
(2-109); 26766(7-2); 27278(3-4); 27365 Gentle, P. H., 29(6-47); 65(7-56b); 943(7-56b);
(7-56a); 27377(7-24); 27501 (7-56a); 27564 1150(7-22); ii82(2-88a); 1248(I-I); 1249
(7-50a); 27596(7-56a); 27677(7-56); 27705 (2-88a); I287(2-88a); I585(2-88a); 1876
(2-134); 27718(7-30); 27729(7-56a); 27799 (7-56b); 2406(7-56b); 2456(7-22); 2642
(7-20); 27889(2-91); 27934(2-24); 28078 (2-88a); 2744(2-88a); 3342(2-88a); 3535
(7-8); 28080(6-42); 28104(7-21); 28130 (2-32); 4I65(7-56b); 4365(2-88a); 4432
(6-4 ); 28133(2-27a); 28193(7-55); 28220 (2-88a); 4493(7-23); 4574(7-22); 4585
(2-119); 28358(7-1); 28524(2-43a); 28645 (2-88a); 4806(2-88a); 5229(7-23); 5570
(6-36); 28714(7-6); 28777(2-88a); 28951 (2-88a); 6I93(2-88a); 2028(2-88a); 7081
(7-I); 28964(6-54); 29037(2-88a); 29048 (2-88a); 7598(7-56b); 7930(2-88a); 8177
(6-I5); 29070(3-13); 29076(2-I24); 2911I (2-88a); 8584(2-32); 8709(2-88a); 9188
(7-49); 29I77(2-88a); 29118(7-67); 29207 (2-88a); 9744(2-88a)
(7-8); 293I6(7-24); 29594(4-I); 30059(6-3c); Gerhart, G. A., 10(2-I)
30162(6-21); 30200(7-36a); 30274(7-18); Gevieski, A., 7(7-85); 131(7-85)
30379(2-43a); 30530(7-36a); 30541(6-2); Geyskes, D. G., I8(2-69a)
30588(6-2); 30617(6-2); 30654(6-2); 30946 Gillis, W. T., 6373(1-I)
(7-5ob); 31219(6-37); 31500(7-63); 31771 Gilly et al, 57(7-56b)
(7-27); 31819(3-4); 31836(7-27); 31920 Gines, Bro. H., 4849(1-I); 4937(7-56a); 5071
(2-66); 31944(3-I); 3197 (3-4); 31996(2-98); (7-25); 5141(7-25); 517I(I-I); 5178(I-I);
32164(6-54); 32312(2-63); 32340(7-56a); 5232(1-1)
32374(2-124); 32438(2-63); 32508(2-124); Giovanni, F. di, sn(3-6)
32883(7-56b); 32896(3-4); 33059(2-39); Glaziou, A. F. M., 78(6-25); 237(2-II); 738(2-
33075(2-114); 336I (7-56b); 33I83(7-56b); I4);752(3-4); 896(6-25); I382(6-49); 1383
33223(7-69a); 33563(4-I); 33572(2-27a); (6-49); I384(6-25); I389(2-43a); 1514(2-11);
33585(2-52); 33627(6-50); 33653(6-2); 2127(7-85); 2126(6-26); 2128(3-4); 2559
33736(2-27a); 33939(2-24); 33991 (2-27a); (7-85); 2560(3-4); 2561(2-70); 2960(3-4);
34014(2-18); 34133(2-134); 34175(2-28); 4946(7-88); 5676(6-26); 6167(6-25); 6i68
34504(7-56a); 34506(2-134); 3453 (7-56b); (2-71) ;6I99(7-36b); 7692(3-4); 7603(2- 114);
34543(7-56b); 34558(7-56a); 34618(2-69a); 7876(6-26); 7879(2-II); 8399(2-132); 8666
34675(6-50); 34788(7-40); 34888(6-2) (7-36b); 9391(3-4); 9716(I-I); 9786(7-63);
Fromm, E. et al, I337(1-I); 1401(2-27a) 9788(2-36); 9789(2-69a); 10698(I-I); I0700
Frost, S. W., 70(7-56b) (2-44); I0701(2-27a); 10702(2-50); 10703
Fuchs, H. P. et al, 21793(3-I5); 21852(7-10); (2-II); I0705(7-56b); 10706(7-20); 11941
22036(2-40) (7-82); II942(7-85); 11943(2-132); II943a
Fuertes, P. M. D., 19I(I-I); 332(7-36a); I650 (2-124); 12657(7-70); 12658(7-82); I3433
(7-36a) (6-25); 1367I(2-69a); I371I(2-69a); 13796
Funck, N. et al, 536(7-36a); 556(7-36a) (3-4); I3798(6-3a); I3799(6-I5); 13800
376 Flora Neotropica

(2-82); 14676(7-70); I4677(4-4); 14678 Hall6, 450(7-25); 570(2-8); 576(7-25); 662(6-2);

(2-43a); 14680(7-54); 1468I(7-54); I5943 726(2-97)
(4-4); 15944(7-54); 16761(6-25); 18217 Handro, 0., 692(2-41); 890(2-41); SP 28197
(6-49); 19137(2-132); 19138(7-81); 19139 (2-114); SP 31063(2-114); SP 39715(2-I44);
(6-20); 20117(6-27); 21109(7-66); 21110 (7- SP 43305(3-i6)
20); 2 111(7-54); 21112(3-I6); 21113(2-80); Hans, D., 132(6-25); I62(6-25); sn(7-36b)
21114(2-114); 2I115(2-43a); 21116(2-41) Hansen, C. 0. E., sn(7-36a)
Gleason, H. A. et al, 73(7-83); 208(7-56a); Harbison, T. G., 2(2-I)
233(7-56a); 447(2-88a); 539(6-7) Harlow, J., I(7-56b); 94(7-56b)
Godfrey, R. K. et al, 53336(2-I); 55531(2-I) Harper, J. S., 204(2-I)
57022(2-I) Harper, R. M., 698(2-I); 983(2-I); I433(2-I)
Goeldi, E. A., MG 3882(3-4); MG 3913(7-53); Harris,J. A., 17257(1-I); I9973(2-I); 2I265(2-1);
MG 3933(6-24); MG 3948(7-56a); MG 3993 21271(1-I); C 21322(1-I); C 23329(1-I);
(6-9); MG 7278(7-56a); MG 8233(2-86); 23363(1-1); C I9IO52(2-I); sn(I-I)
MG 8234(6-4); MG 15086(7-65) Harris, W., 53I7(7-36a); 54I7(7-36a); 5604
Goes, O. C., 790(2-69a); 943(3-4); 965(7-85); (7-36a); 5966(7-36a); 6721(1-I); 8515(I-1);
1046(3-4) io753(7-36a); Ii66(I-i)
Goldman, E. A., 615(1-I); 6i6(i-I) Harrison, S. G., 741(1-I); 798(7-25); 1020(7-25);
Goll, G. P. et al, 864(1-1) 1239(6-42)
Gomes, I., sn(6-25) Harvey, D. R., 5187(7-56b)
Gomes, M., 1107(3-16); 2409(2-I27); RB 47989 Hassler, E., 10700(7-66); sn(I-I)
(2-127) Hatch, W. R. et al, sn(I-I)
Gomes Moura, J., sn(2-28) Hatschbach, G., 1121 (7-85); 2819(6-27)
Gomes, V., 2482(7-85); sn(7-36b) Haught, 0., 1468(7-35); I528(7-56a); 2604
G6mez, 12(6-3b) (7-40); 2629(2-47); 2630(7-56b); 3817
G6mez-Pompa, A., 887(2-15); 922(2-I5); 947 (7-36a); 4024(7-36a); 4264(7-36a); 4540
(6-47); 1216(2-45) (i-I); 4710(7-35); 5418(7-I0)
Gonggrijp, J. W. et al, B.W. 905(7-74); B.W. Hayes, S., 3o(7-56b); 45(I-I); 407(7-56b); 646
5333(7-2); B.W. 5666(7-2) (7-22); 900(5-I)
Gonziles, A., 27(3-7) Heinsdijk, D., 3(2-135); 55(2-I35); 62(7-63)
Gonzalez, L. C. et al, I8C-042(7-22) Heller, A. A., 7(1-I); 730(7-83)
Gonzalez, L. Q., 583(7-56b) Hemmendorf, 194(7-54)
Gooding, E. G. B., 336(1-1) Henz, 35330(7-85)
Goudot, J., sn(7-22) Herb. Lagasca, 78(7-56b)
Gouvea, J., INPA io66(2-43b) Herb. Meyer, F. G. W., 51(I-I)
Graf, L. J., 4(- I) Herb. Moquin-Tandon, sn(7-83)
Graham, V., 124(7-21); 147(2-135); 354(2-I34); Herb. Presl, sn(7-36); sn(7-56b)
390(6-43); 439(7-21); 500(6-43) Herb. Richard, sn(7-56b)
Gray, sn(2-I) Herb. Swartz, sn(7-36a)
Greenman,J. M. et al, 5739(1-1) Herb. Vahl, sn(7-73)
Gregory, L. E., 12(7-36a) Heriberto, Bro., 5(I-I)
Grosourdy, R. de, 13(7-25) Heringer, E. P., 514C(7-54); 2160(7-85); 2584
Grubb, P. J. et al, 1045(6-22) (2-I 14); 4087(6-27); 5368(6-27); 5415(7-54);
Grunwald, 0., sn(2-43a) 5472(7-54); 7218(7-20); 8799(6-27); 9035
Guedes, M., MG I234(7-56a); MG 2141(6-4); (7-20); 9272(2-41); 9288(6-27); 9493(7-70);
MG 2583(2-52); MG 2589(7-56b) 10466(2-41); 10474(7-20); 10529(6-27);
Guedes, T. N., 20(2-62); 30(2-69a); 225(6-15); Io665(2-27a); 8671/865(3-16); 8709/903
238(2-39); 256(6-4); 628(2-43a); 632(7-18); (3-i6)
638(7-18); 2141(6-4) Herman, F. J., III76(7-56b)
Guilding, L., sn(7-25); sn(i-I) Hernandez, A., 56(7-25); I02(7-56b)
Guppy, N. G. L., 226(6-7); 250(7-2); 322(7-42); Hess, W. E., 5614(7-36a)
408(2-93); 566(7-56a); 595(2-134); 596 Heyde, E. T. et al, 531(1-I); 3959(1-I); 6421
(7-56a); 653(2-99); 669(6-7); 7189(7-15); (2-I5)
sn(7-56a) Heyder, H. M., I6(7-56b)
Gutidrrez, G. et al, 510(7-40); 567(2-27a); Heyligers, P. C., 117(2-135); I64(2-69a); i65
661(2-140); 841(7-62); 862(2-129); 875 (6-42); 343(2-66); 448(7-5ob); 466(6-I);
(2-129); 882(2-129); 883(2-27a); 907(2-129); 524(2-66); 746(2-135)
985(7-56b) Hinds, R. B., sn(7-56b); sn(i-i); sn(2-88a)
GuzmAn, R., sn(6-47) Hinton, G. B., 3021(2-45); 3832(2-45); 5357
(2-45); 5957(2-45); I005(2-45); 10773
Haenke, T., 1532(6-47) (7-56b); I1040(7-56b); 13750(6-47); 14008
Hahn, L., 191(I-I); 213(7-36a); 528(1-I); 951 (6-47); 14I26(6-47)
(I-I); 973(7-36a); 1124(2-12); I247(7-36a) Hioram, Bro., sn(7-36a)
TLit of Fvcxsicrrtaep 77

Hitchcock, A. S., 87(1-1); 579(I-I); 16883(1-1); Humbert, H., 27407(7-21)

16913(2-135); 1706I(7-56a); 17063(7-25); Humboldt, A. von et al, I059(2-I2); I798(7-22);
sn(7-22) sn(6-47); sn(7-56b); sn(7-56b); sn(7-83)
Hodge, W. H., 578(2-85); 579(7-36a); 582(1-I); Hume, H. H., 22(2-I)
583(I-I); 584(1-I); 585(1-1); 960(I-I); Hunnewell, F. W., 5862(I-I); 735I(I-I); 8679
1237(2-85); 1319(2-85); I735(2-85); 2000 (2-I); I6474(7-56)
(7-37a); 2067(2-85); 2395(2-85); 2569 Hunt, D. R., i82(7-56b); 323(7-56b); 354(I-I);
(7-36a); 2672(2-85); 2696(1-1); 2840(2-85); 473(7-22); 5597(7-20); 5600(7-54); 5682
3067(1-1); 3074(7-36a); 335I(I-I); 3590 (7-54); 5923(7-20)
(2-85); 3750(I-I)
Hoehne, F. C., SP 1119(7-88); 1342(7-66); I41I Idrobo, J. et al, I292(7-22)
(7-66); 1496(2-144); I809(7-57); 1816(7-57); Im Thurn, E. F., sn(7-I8); sn(2-28); sn(2-43a);
2033(7-54); 2094(7-54); 2141(7-19); 2154 sn(2-I34)
(7-19); 2155(7-19); 2186(2-41); SP 4206 Imray, J., 129(2-85); 166(2-85); 301(7-36a);
(7-85); 4216(7-85); 4262(2-30); 4589(2-30); sn(I-2); sn(2-85); sn(7-36a)
4602(2-18); 4681(7-54); 4909(7-57); 6078 I.N.P.A., (Instituto Nacional de Pesquisas da
(7-54); SP 7546(7-85); SP 11978(3-16); Amaz6nia), 2I63(6-29); 3228(6-29); 3247
SP 17197(6-46); SP 22997(2-144); SP 27385 (4-I); 3710(4-I); 4062(2-41); 4141(2-60);
(2-I 4); SP 27563(2-114); SP 27716(6-27); 4883(2-112); 5730(2-60); 5893(6-4I); 6028
SP28586(7-85); SP 29359(I-I); SP 29539 (2-63); 6031(2-26); 6045(6-50); 6062(2-26);
(2-144); SP 36764(6-27); sn(7-64) 6065(6-15); 6092(6-29); 6117(2-129); 8078
Hohenkerk, L. S., 39A(2-8); 39B(2-69d); I39A (6-I5); 9040(6-28); 9041(6-4); 9043(6-II)
(2-69d); 524(6-45); 676(2-93); 676A(2-I26); Irwin, H. S. et al, 6(2-135); 8(6-15); 72(I-I);
705(2-66); 723(7-25); 80I(2-78); 816(2-83) I3I(2-66);2-6);8438(66);238(2-43a); 256(2-89);
Holdridge, L. R., 437(7-32); 2158(7-36a); 2306 383(2-66); 593(7-56b); 5058(3-16); 5257
(7-36a) (2-43a); 5324(7-54); 5368(7-20); 5548
Holn, H. T., sn(2-I) (6-27); 58I5(7-54); 6070(2-80); 6I97(7-54);
Holt, E. G. et al, 69(2-88a); 71(1-1); 88a(2-I2); 6200(7-70); 6423(6-27); 6430(2-41); 6459
244(7-56b); 517(2-135) (7-19); 6495(6-27); 6615(6-27); 665I(2-I7);
Holton, I. F., II(I-I) 6711(6-27); 6744(2-30); 7340(3-I6); 7444
Home, F. W. et al, 9724(7-83) (3-16); 7821 (2-43a); 7824(7-20); 7847(3-16);
Horto Florestal, Rio de Janeiro, 1oI2(6-55); 7887(7-54); 7922(7-20); 7968(6-27); 8229
RB 136916(7-85); RB I369 7(7-36b); 136922 (3-i6); 8465(7-54); 9I97(3-i6); 10176
(7-33); RB 136924(7-64); RB 136926(2-I14); (2-27a); 10580(3-16); 10813(3-16); IIIOI
RB I36932(2-114); RB I36934(2-II4); RB (3-I6); 11587(3-16); 11649(3-16); 11833
136939(2-4Ia); RB 139762(6-55) (2-49); 13435(2-4I); 13608(3-16); 13722
Hortus Botanicus Paris, sn(2-151) (7-54); 13880(3-16); 14469(3-16); 14483
Hostmann, W. R., 78(7-25); I75(7-73); I85 (4-2); I4521(2-80); 14591(4-4); 14694(4-2);
(7-56a); 363(2-26a); 476(1-I); 6o0(7-25); 147I9(2-80); 14700(3-I6); 14720(2-80);
794(2-135); 85Ib(7-56b); 859(6-2); 860 14813(4-2); I4980(7-54); 15332(2-80);
(7-56a); Io38(2-69a); 1119(6-2); I250(2-0o8) 15656(7-54); 15895(7-54); 16137(7-20);
1257(2-124); 1712(2-135); sn(I-I) 16292(7-54); 16435(7-20); 16718(2-30);
Hotchkiss, N. et al, 3856(2-I) 16888(2-30); 16893(7-20); 17034(2-30);
Howard, R. A. et al, 5597(I-I); 5622(7-36a); 7127(2-41); 17145(7-66); I7347(7-66);
6017(1-I); 9789(7-36a); I0087(I-I); I0329 17352(2-41); 17421(7-66); 16713(2-41);
(7-25); I0492(I-I); 11347(I-I); I1638(1-2); 17707(2-30); 17864(2-41); I7923(2-30);
1640(7-32); II769(7-36a); I I886(7-36a) 18009(7-20); 18039(7-70); 18043(7-70);
15496(7-83); 15545(7-83); I5595(7-36a) I8229(2-48); 23408(7-54); 23454(7-20);
Howell, A. H., 905(1-1); 928(1-I); I060(2-I); 23881(2-80); 47147(2-69a); 47177(7-2);
1075(2-I); I I44(2-I) 47183(7-20); 47184(2-69a); 47195(2-69a);
Howell, J. T., 10297(6-4) 47210(3-3); 47295(7-52); 47338(7-56); 47367
Huber, J. E., MG 749(7-24); MG 768(7-56a); (7-80); 47434(2-134); 47439(7-56a); 47470
MG 803(7-56b); MG 854(6-2); MG 946 (2-134); 47474(7-56a); 47617(2-134); 47766
(2-52); MG o059(7-56a); MG 1147(2-39); (6-2); 4778I(2-97); 47836(2-I25); 47862
MG 1876(3-4); MG I877(2-69a); MG I894 (2-125); 47914(2-125); 48037(2-97); 48122
(2-95); MG 2030(6-4); MG 2055(7-56b); (2-125); 48327(7-56a); 48344(2-125); 4836oa
MG 2617(2-39); MG 2725(7-56b); MG 3015 (2-69a); 48390(6-2); 48443(6-2); 48463
(2-II); MG 3664(6-36); MG 3819(7-27); (7-20); 48589(7-56a); 48594(7-5ob); 48622
MG 4843(6-36); MG 6982(2-57); MG 7045 (6-i8); 48695(6-18); 48741(2-77); 48765
(3-2); MG 9360(2-II); MG 9359(6-36); (7-56a); 54564(2-108); 54594(7-74); 54785
MG o1457(7-5oa); MG I5224(2-II) (2-o18); 54802(6-34); 54889(7-74); 54903
Huertas, G. et al, 4072(I-I) (6-18); 54904(7-36a); 54926(2-122); 54937
Hulk, J. F., 320(2-134) (2-o18); 54988(2-1o8); 55002(2-73); 55164
378 Flora Neotropica

(7-56a); 55169(7-71); 55205(7-56a); 55218 Karling, J. S., I9(1-I)

(7-25); 55224(I-I); 55226(3-I); 55288 Karsten, sn(7-36a)
(7-56a); 55370(7-25); 55374(3-I); 55375 Kauffman, E., 2I(7-56b)
(2-134); 55387(2-i08); 55409(4-3); 55425 Kegel, H., 250(7-25); 271(2-135); 303(7-25);
(2-69a); 55434(2-77); 55454(3-I); 55460 456(7-25); 470(I-I); 1154(2-I35); 1155
(7-25); 55520(7-2); 55543(7-56a); 55667 (6-42); 1346(7-56a); 1500(3-I)
(2-108); 55681(2-122); 55683(7-73); 55756 Kellerman, W. A., 4758(I-I); 4889(1-I); 5670
(2-I34); 55766(4-3); 55769(2-Io8); 55915 (2-15); 6672(7-56b); 7063(2-45)
(I-I); 55976(4-3); 55995(3-I); 57592(7-73) Killip, E. P. et al, 624(6-36); 336I(7-56b); 3409
Isert, P. E., sn(7-36a) (7-22); 4015(3-I); 4666(3-I); 4976(3-I);
11702(7-10); 14093(1-1); 24794(7-2I); 25276
Jack, J. G., 4429(1-I); 8243(2-1); 8250(1-1); (7-21); 27I84(6-3b); 27549(7-56b); 27865
8776(I-I) (6-36); 27933(6-36); 2927I((7-36a); 29651
Jahn, A., 459(7-34) (7-56b); 29795(7-56a); 29947(7-5); 29963
Jan, L. K., sn(7-7i) (6-3b); 30031(6-9); 30382(1-1); 30452
Jaramillo, J. D., I250(1-I) (7-20); 30546(3-I); 30548(7-56b); 30577
Jaramillo, R. et al, IooI(7-40); 1027(6-2); 1174 (7-56a); 30954(1-1); 31660(2-I); 32791 (2-);
(7-40); 1211(7-40) 33211(2-96); 34298(7-40); 37283(7-56b);
J,tiva, C. et al, 712(7-10) 37465(7-56a); 37470(2-69a); 37565(7-56b);
Jenman, G. S., 42(2-43a); 298(1-I); 540(1-I); 37668(1-I); 38965(7-10); 39000(2-96); 39130
76 (6-45); 884(7-51); 969(6-45); 970(7-51); (7-10); 41313(7-22); 41475(I-I); 42004
1088(6-43); 1088(6-45); 1110(7-25); I314 (2-I); 42080(3-1); 42611 (I-); 42678(7-22);
(2-I24); 2011(3-12); 2258(1-I); 2393(2-I09); 42884(1-I); 42911(7-22); 44231(2-1); 44449
3604(2-77); 3615(2-8); 3742(2-135); 3839 (I-I)
(2-109); 3842(2-8); 3902(2-69d); 3917 King, R. M., 979(7-56b)
(2-69a); 3952(6-15); 3979(3-I); 3980(7-74); Kings, 355(1-I),
3992(3-12); 4012(2-109); 4041(2-93); 4110 Kinloch, I7o(2-88a); 234(2-88a)
(2-90); 4184(6-7); 4232(6-15); 4282(2-8); Kirk, J., sn(7-9o)
4287(7-56); 4301(2-39); 4650(2-39); 4734 Klein, R. M., I48(7-85); 754(7-85); 871(7-85);
(7-45); 4865(2-43a); 4900(7-25); 5073(3-1); 898(7-85); 921(7-85); 2169(7-85); 2304
5326(7-25); 5678(7-I8); 6031(6-42); 6274 (7-85); 2731(2- 1)
(2-27a); 6300(2-78); 6580(6-2); 6624(2-95); Klug, G., 27(7-5); 142(6-7); 203(7-56a); 449
6625(3-I); 6627(2-I09); 663o(2-43a); 6967 (2-101); 462(2-140); 512(7-5); 587(2-26);
(2-8); 6998(2-8); 7000(7-38a); 7125(2-8); 726(7-40); 729(2-27a); 916(7-40); 989(7-2);
73i6(2-27a); 7336(2-69d); 7627(6-45); 773I 1333(7-56a); i444(6-3b); 1525(2-88a); I526
(3-I) (2-52); I854(6-36); 1948(7-28); i989(7-56a);
Jennings, 0. E., 9(I-I); 208(7-22) 2089(6-22); 2221(2-7); 225o(7-36a); 2520
Jewitt, A., sn(2-I) (6-36); 2530(7-40); 2775(7-40); 2656(6-36);
Jimenez, M. A., 327(7-56b); 1210(7-36a); 2848 2704(7-56a); 2714(7-56a); 2869(7-59); 3022
(i-I); 3010(2-45); 3515(7-36a) (7-59); 3724(7-36a); 374o(2-88a); 3870
Jobert et al, 27(7-56b); 273(7-56b); 863(7-63); (3-9); 3971(6-36); 4025(7-56b); 4249(7-48)
94o(7-56b) Kluge, H. C., 13(I-1); 3I(2-I5); 48(7-36a)
Johansen, H., 6(6-47); 14(1-I) Koscinski, M., SP 3i640(7-85)
Johnson, E. P., sn(7-56b) Kral, R. et al, 3959(I-1); 5060(2-I); 6751 (2-I)
Johnson, H., 1070(2-15); 1167(1-I); I284(7-56b) Kramer, K. U. et al, 2910(2-135); 2988(1-I)
Johnston, I. M., 226(7-56b); 292(1-1); 513(7-22); Krukoff, B. A., Io59(7-56a); Io64(7-56a); 1137
546(7-22); 659(2-15); 802(1-I); 850(7-83); (4-3); I244(7-65); 1362(6-2); 1452(2-62);
938(1-I); 1250(7-56b); I673(7-56a); I749 I50I(3-4); 4557(7-6); 4576(6-3b); 4648
(7-56a); I782(2-88a) (7-73); 4655(7-36a); 4684(2-44b); 4744
Jones, D. G. et al, 277(7-56b); 35I8(7-23) (2-69b); 4760(7-36a); 4796(7-69a); 4846
Jones, G. C. et al, 3059(2-88a); 3272(2-88a); (2-69b); 4880(2-53); 4852(7-36a); 4913
3288(2-88a) (6-29); 49I5(2-43b); 49I6(7-2I); 4962(7-41;)
Jones, S. B., 13074(2-1) 4977(7-56a); 5014(2-98); 5070(2-55); 5099
Jones, sn(2-I3); 32(6-47) (7-5ob); 5I65(2-2I); 5246(7-36a); 5252
Jonker, F. P., I49(2-I35) (2-69a); 5321(7-36a); 5354(7-56a); 5379
Jordan, sn(2-I35) (2-69a); 5428(7-36a); 5491(7-75); 5525
Joseph, Bro. A., sn(I-I) (7-48); 5629(2-43b); 5635(2-21); 5650(3-5);
Jurgensen, 924(7-56b); sn(I-i) 5726(6-22); 5786(7-75); 5862a(3-3); 5887
Jussieu, sn(2-I34) (7-56a); 596 (7-56b); 5979(7-24); 6005
Juzepczuk, S. V., 116(2-45) (6-24); 6007(6-23); 6037(6-44); 6139(2-21);
6208(6-23); 6306(2-21); 6487(2-69a); 66i8
Kappler, A., 439(2-I35); I824(3-1); I887(1-1); (2-57); 6668(2-75); 6670(2-129); 6688
I968)2-95); 2 I 7(2-27a); I241(6-2) (7-56a); 6703(6-3b); 6812(2-62); 68I 7(2-94);
List of Exsiccatae 379

6834(7-6); 6860(2-33); 6874(7-2); 6896 Lang, H. et al, 41 (7-56a); 127(2-129); 242(2-129)

(6-9); 6897(2-81); 6898(2-69a); 69I3(2-69a); Langlass6, E., 647(7-56b); 992(2-9)
6926(2-94); 6983(7-6); 7002(2-I29); 7028 Langman, 3341(6-47)
(2-69a); 7036(2-33); 7058(2-33); 7084(2-57); Langsdorff, G. H. von, sn(7-36b); sn(2-43a)
7109(6-9); 7124(7-24); 7216(2-62); 7237 Lanjouw,, J. et al, 152(2-135); 265(2-I35); 326
(2-129); 7252(6-13); 7267(2-140); 7271 (2-63); 327(2-95); 332(2-95); 357(2-99);
(2-69b); 7983(6-9); 8100(7-40); 8138(2-75); 401(2-124); 420(2-63); 438(2-135); 454
8190(7-2); 8276(2-75); 8445(2-13); 8451 (2-73); 562(7-25); 563(2-135); 574(I-I);
(2-124); 8507(2-75); 8553(2-43b); 8560 618(2-63); 709(6-2); 727(2-134); 732(2-135);
(2-43b); 8640(2-22); 8646(2-69a); 8727 792(2-I34); 805(3-I); 810(7-25); 895(2-134);
(7-62); 8755(2-94); 8751(2-94); 8761(2-33); 911(2-108); 916(2-69a); 928(7-56b); 968
8764(2-99); 8843(6-29); 8850(2-18); 8851 (6-2); I24(7-56a); I202(I-I); I233(7-56a);
(2-i6); 8869(2-33); 8886(2:94); 8907(2-33); I238(2-69a); 1245(2-135); I307(3-I); 1833
8962(2-69a); 8966(2-40); I0558(7-73); 10638 (2-66); 1837(6-42); I838(2-135); 2663(2-52);
(7-36a); 10658(7-36a); 10733(7-46); 0o876 2805(2-112); 2949(7-73); 3048(7-25); 3287
(7-56a); 10884(7-74); 10996(2-55); IIOOO (2-135); 3327(7-25); 3377(7-73); 3458(2-63);
(3-5); 1104I(2-2); IIII3(3-5); 11233(2-2); 3461(2-8)
11246(2-55); I1278(3-5); 11293(6-I); 12316 Lankester, C. H., I336(7-56b)
(2-135) Lasser, T., 895(3-7); I492(7-56); I626(7-56a);
Kuhlmann, J. G., 35(2-144); 121(2-27a); 154 1704(7-49); 1730(2-56); 1796(7-21); 2238
(3-4); 162(2-6); 174(2-126); 208(2-71); (7-46a); 2368(2-43a); 2580(7-36a); 2611
227(7-54); 231(7-20); 240(6-27); 279(4-5); (7-22); 4048(7-36a)
296(2-37); 298(7-54); 300(7-54); 301(7-54); L. B. B., 8358(2-135); 9131(2-135); 9185(2-117)
303(2-75); 304(2-75); 306(6-3a); 312(2-28); Leal, C. G. et al, I54(2-43a); 205(7-56b)
315(2-27b); 364(2-144); 38I(7-36b); 502 Leavenworth et al, 452(2-45); I448(2-45); I6I2
(2-27a); 927(2-27a); 928(2-122); 963(6-9); (2-45)
1ooI(2-11); I004(2-124); 1561(2-43a); I577 Leblond,J. B., 464(2-1 I4); sn(7-83)
(7-88) ;I842(7-24);I868(2-27b) ;I922(2-88a); Le Conte, L., sn(2-I); sn(2-I)
I988(7-36a); 1990(7-58); 1992(2-88); 6672 Ledoux, P., 287(2-40)
(7-3I); RB 2025(6-25); RB 2807(7-56b); Le Dru, A. P., sn(7-83)
RB 2809(2-27a); RB 2903(7-56b); RB 2950 Lehmann, F. C., 98(1-I); B.T. 646(7-10); 7556
(6-43); RB 2972(7-I); RB 3060(3-4); (I-I)
RB 3061(2-22); RB 35II(7-25); RB 4539 Le Conte, L., sn(2-I); sn(2-I)
(2-122); RB 15097(7-85); RB 15138(6-43); Ledoux, P., 287(2-40)
RB 17891(2-27a); RB 17966(6-16); RB Le Dru, A, P., sn(7-83)
17967(6-15); RB 17969(2-97); RB I7970 Lehmann, F. C., 98(1-I); B.T. 646(7-10); 7556
(2-75); RB 17973(2-129); RB 17974(6-3b); (I-I)
RB 17976(6-43); RB I7977(7-I); RB 17978 Leme6, A. M. V., sn(2-86); sn(I-i)
(2-27a); RB I7979(2-27a); RB 17980 Lemoine, FD 7757(7-73); FD 7786(6-2); FD
(2-27a); RB 17982(7-24); RB 17985(2-21); 7793(6-2); FD 7824(3-I); FD 7850(6-2)
RB 17986(7-54); RB 17986(7-56b); RB Lems, K., 5124(2-135); 5302(7-61); 650224
17987(6-16); RB 21027(4-I and 3-I); (2-88a); 650227(2-135); sn(I-I)
RB 41453(2-44); RB 48I 10(2-6); RB 138215 Lent, R. W., 345(1-I)
(7-85); RB 138217(2-114); RB 139760(6-25) Le6n, Bro. H. et al, 998(2-45); 5908(1-I); 7909
Kuhlmann, M. et al, 9(2-I1); 28(6-4); 68(6-4); (7-36a); 9249(1-I); I0970(7-36a); 16242
92(6-4); 95(2-8I); 117(2-57); 161(6-4); (I-I); 18874(7-22); 22659(7-36a); 22663
225(7-85); 229(3-3); 268(7-20); 289(6-50); (7-36a)
328(3-3); 342(7-56a); 1593(7-54); 1748 Leonard, E. C. et al, I2417(7-36a); 13950(1-I)
(7-88); 2354(3-4); 2736(3-4); 2738(3-4); Leprieur, F. R., sn(7-25); sn(7-65a); sn(i-I);
2983(1-1); 3174(6-46); 3374(6-46); 3455 sn(2-69a); sn(2-27a); sn(2-I34)
(2-4I); 3933(2-37); 4328(7-88); SP 24062 Le Roy, P. U., sn(i-i)
(7-85); SP 66047(2-11 4) Le Sueur, H., 553(1-I)
Kuntze, C. E. O., 980(1-I); 981(1-I); 1014 Level, J. S., 30(2-131); 93(6-2); I02(2-69a);
(2-27b); I112(7-25); sn(2-4I); sn(7-54); I2I(7-56b); I34(3-I)
sn(7-56a) Levy, P., 46(6-47); 62(1-I); 222(2-I5)
Kuyper, J., 109(2-69a); 561(2-135) Lichy, E., 1(7-8)
Liebmann, M. F., 1746(6-47); I751(7-36c)
Labroy, sn(2-36); sn(6-3b) Lightthipe, L. H., 585(2-I)
Lakela, O., 24102(2-I); 24962(2-I); 25975(1-1); Lima, D. A., 135(1-1); 1009(6-49); 3118(7-16);
29832(2-I) 48-103(7-40); 48-122(6-49); 48-3107(7-16);
Lallemand, sn(7-64) 48-3108(2-63); 49-209(2-71); 49-290(2-71);
Lane, F., 8(2-41); SP 69973(1-27); SP 69989 50-433(7-I8); 50-743(7-I8); 52-I I02(7-56b);
(7-54); SP 69995(7-20) 52-1135(7-18); 52-1200(2-71); 53-I206(6-53)
380 Flora Neotropica

54-I897(6-6I); 55-2072(2-7I); 55-2256(6-25) (7-55); 22162(7-56a); 22226(7-3); 22245

58-2882(7-18); 58-3078(7-54); 58-3083(2-30) (7-3); 22502(7-40); 22758(7-40); 25878
58-3139(2-17); 58-3I57(2-69a); 58-3 75 (7-i8); 26616(2-44); 26619(2-44); 26642
(2-I7); 58-3I84(7-50b); 58-3272(7-18); 58- (2-44); 26780(7-20); 28105(2-44); 28106
3279(7-20); 58-3294(7-64); 60-3434(2-7I); (2-44); 28107(2-44); 28108(2-44); 28109
61-3919(2-43a); 65-4311(2-II4); 65-4340 (2-44); 28112(2-44); 28 13(2-44)
(2-43a); 65-4367(7-50a); sn(7-20) Lutz, A., 671(6-25); 931(6-25); MG 3575(6-49)
Lima, F., MG Io858(7-56a) Lutz, B., 1230(7-36b)
Lima, N., 86(7-56b)
Lindeman, J. C. et al, 3671(6-18); 3894(2-135); Macbride, J. F., 2645(7-56b); 5532(7-2I)
3896(2-66); 4048(6-18); 4125(6-I); 4137 Macedo, A., 416(7-54); 702(2-41); 715(6-27);
(2-66); 4139(6-42); 4178(6-18); 4205(2-66); 1128(2-30); 1262(6-20); 1827(7-19); 2524
4599(2-124); 4635(6-42); 4636(2-135); 497I (7-54); 2550(2-30); 2558(6-20); 3466(3-16);
(2-I24); 5014(7-42); 5209(2-1I7); 5794 3607(7-20); 3624(2-37); 3885(7-20); 3887
(6-i); 5874(2-99); 5923(6-I); 5947(2-99); (7-20); 3905(6-27); 4012(7-18); 4035(6-3c);
6o09(6-18); 623o(2-43a); 637I(2-43a); 6375 4065(6-20); 4261(3-16)
(2-66); 6431(6-i); 6743(6-18); 6781(2-124); Macfayden, J., sn(7-36a)
6795(4-3); 687o(2-69a); 70II(2-52); 7024 Machado, D., RB 75835(6-25); RB 75836(2-7ia)
(2-52) Mackenzie, K. K., 4038(2-I)
Linden, J. L., 556(7-36a); I603(6-47); 1637 Magalhaes, G. M., 214(7-54); 706(7-54); 2298
(7-56b); I638(7-22); 2977(7-66); sn(I-I) (2-I44); 60I5(6-26); 9670(3-I6); 9854(3-16);
Linder, D. H., I5(6-45); 116(7-25); I 9(7-56a) I393I(7-20); 17996(2-41); I8I13(7-20);
Linnaeus, Hort. Cliffortianus, pI 7(7-22) 19240(2-41); 19005(3-16)
Lisb6a, A., MG 2377(2-27a); MG 2378(6-20); Maguire, B. et al, 22912(7-16); 22924(7-54);
MG 2428(2-27a); MG 2438(2-44); MG 2458 22949(2-8); 23005(7-51); 23005a(7-77);
(2-37); MG 2462(2-19); MG 2482(2-30); 23030(7-51); 23151(2-135); 23266(2-135);
MG 2489(7-54); MG 2494(6-27); MG 2501 23363(6-13); 23364(7-56a); 23390(3-I);
(7-20); MG 2505(7-18); RB 4701(7-56b); 23498(7-77); 235 I(2-II2); 23514(7-74);
RB 4702(7-20) 235I5(7-56a); 23569(6-13); 23651(2-135);
Little, E. L. et al, 6275(7-10); 6475(7-87); 23820(7-25); 24024(2-77); 24059(6-17);
6668(7-36a); 79I9(7-22); 8066(7-22); 8o8i 24066(2-98); 24068(2-98); 24103(7-2);
(7-22); 8304(7-22); I3093(7-83); 13420 24232(2-135); 24354(2-135); 24392(7-30);
(7-83); I59I2(2-I2); 15991(3-I); I6ooo 24438(2-I35); 24782(6-7); 2479 (2-35);
(2-77); 17615(3-3); I7674(2-77); I7679 24795(2-113); 24862(2-134); 24947(7-56a);
(7-20) 24982(2-135); 27315(7-56b); 27359(6-3b);
Lloyd, F. E., 338(7-36a); 613(1-1); 761(2-85) 27361(7-25); 27443(7-25); 27854(6-3b);
Lockhart, D., I62(7-36a); 220(7-36a); sn(i-i) 27953(2-69a); 28143(6-3b); 28208(2-69a);
Lodge, F. A., sn(2-69a) 28437(6-3b); 28621(7-73); 28623(7-2);
Lofgren, A. et al, 456(7-85); 930(2-41); I451 28973(7-56b); 28998(6-3b); 29000(2-69a);
(3-I6); I595(7-82); 4180(2-126); 4598(2-41) 29005(2-69a); 29021(6-3b); 29260(2-91);
Long L. E., io6(7-56b) 29293(7-8); 29312(6-9); 29317(7-8); 29448
Loureiro, A. et al, INPA 15512(2-40); INPA (2-18); 29487(2-69a); 29622(7-44); 29682
15514(2-43b); INPA I5518(2-43b); INPA (7-44); 29703(7-44); 29904a(6-3b); 30397
I5532(6-28); INPA I6I39(2-62); INPA (2-99); 30497(2-102); 30524(7-68); 30575
16147(6-50); INPA I6449(6-50); INPA (6-3b); 30577(7-49); 30757(7-40); 30769
16566(2-62) (2-27a); 30785(2-102); 30805(7-49); 30832
Lourteig, A., I777(7-56b); 2340(7-85) (6-3b); 30847(7-68); 30978(7-68); 30995
Lugo, 44(2-4) (2-140); 31008(2-68); 31018(7-55); 3IO19
Luna, A., 212(7-36a) (4-i); 31822(7-2); 32052(2-69b); 32092
Lund, P. W., 37I(7-36a); sn(2-4I) (7-73); 32109(7-73); 32181(7-21); 32196
Lundell, C. L., 20(7-56b); 565(7-56b); 619 (7-21); 32216(7-21); 32217(7-27); 32243
(7-56b); 620(1-1); 2099(7-22); 22 7(7-56b); (2-34); 32451(7-56a); 32603(2-135); 32623
2435(7-22); 2859(7-22); 2956(7-22); 3049 (7-30); 32714(6-5); 3272I(7-56b); 33236
(7-56b); 3079(7-56b); 3267(6-47); 3719 (7-49); 33389(2-150); 33602(7-49); 33774
(6-47); 3817(1-I); 4186(6-47); 4349(7-56b); (7-21); 34500(2-119); 34505(2-102); 34594
4352(1-1); 4874(7-56b); 4878(7-22); 4881 (2-27a); 34685(7-8); 3478I (2-27a); 34801
(6-47); 66oI(2-88a); 6963(2-88a) (2-129); 34845(7-40); 34862(2-69b); 34891
Luschnath, B., sn(7-36b); sn(7-85); sn(2-43a); (2-140); 34892(2-140); 34984(2-140); 34986
sn(6-32) (2-69b); 35043(7-68); 35045(7-21); 35087
Luetzelburg, P. von, I53(7-70); I94(7-I8); 1214 (7-25); 35183(7-25); 35555(6-5); 35556(2-68);
(7-56b); 1381(7-20); I741(7-20); I822(7-54); 35568(2-131); 35598(7-72); 35608(7-72);
I253I(2-44); 20232(2-97); 20252(7-56a); 35624(2-140); 35673(2-27a); 35723(2-88a);
21380(7-1); 21381(7-1); 21531(7-21); 22125 35729(2-I40); 35864(7-56b); 35890(2-122);
List of Exsiccatae 381

35896(7-25); 35999(7-56b); 36215(2-24); Mattos, J. R. et al, I091(7-36a); 8362(6-27);

36237(6-3b); 36238(2-24); 36255(2-24); 9709(2-71); I0788(I-I)
36378(2-88a); 36519(2-124); 36545(3-I); Matuda, E., 388(7-56b); 68i (7-56b); 1090(6-47);
36561(6-3b); 36575(7-49); 36615(2-102); 3054(2-88a); 3097(7-56b); 3102(6-47); 3110
36641(2-91); 36758(2-26); 36767(2-26); (2-88a); 3132(I-I); 3i42(2-88a); 3270(7-22);
37415(2-I29); 37420(3-1); 3747I(7-56); 3343(7-22); 3565(2-88a); 3585(6-47); 3798
37478(2-91); 37526(2-27a); 37537(2-140); (6-47); 4I37(7-56b); 607I(7-36a); 16380
37578(2-27a); 37579(2-91); 37584(2-119); (2-15); I6450(2-15); 16454(7-56); I6500
37604(2-119); 37652(2-68); 37665(7-I2); (6-47); I6887(7-56b); 30680(2-44); 31087
39298(7-74); 40480(7-21); 40493(2-109); (2-44)
40533(2-I 2); 40660(7-21); 40662(2-135); Maurice, 8I6(7-56b)
40676(7-21); 40827(2-15); 41487(2-102); Maury, 4698(2-48); 5313(2-45)
4I55I(7-56b); 41614(7-5); 41663(2-88a); Maxon, W. R. et al, 3787(7-56b)
41687(7-8); 41833(2-88a); 4189 (2-88a); McFarlin, J. B., 6383(2-1)
4I936(3-I); 42630(3-I); 42635(2-27a); 4264I McGrath, G., 8521(1-1)
(7-39); 44156(2-104); 44193(7-22); 44525 McKay, sn(2-69a)
(7-64); 44577(6-25); 45911(2-135); 45978 McVaugh, 8634(2-I); 1861(6-47); I 1879(6-47);
(7-25); 46o09A(7-25); 46136A(2-I 13); 15729(6-47); I8778(7-56b); 19158(6-47);
46247A(2-II2); 4626I(7-25); 46714(7-77); 19242(7-56b); 22244(7-56b)
46754(2-77); 4698I(2-69a); 47052(2-69a); Melinon, M., 4(3-I); I2(2-95); 15(4-3); I6
47073(7-79); 47081(2-139); 47108(2-87); (4-3); i8(6-2); 32(8-I); 33(7-25); 46(3-I);
47I10(7-52); 47128(7-25); 49286(7-20); 54(6-2); 60(2-86); 75(7-56a); 76(2-88a);
53500(2-27b); 53584(2-135); 53690(7-30); 79(7-56a); 17(2-63); 122(2-134); 130(1-I);
53747(2-69a); 53923(7-25); 54008(7-25); I32(7-52); 144(2-97); 148(2-I35); 149(6-2);
54037(6-18); 54100(2-38); 54257(7-2); I8I(2-27a); 187(2-1); 197(1-I); I99(2-27a);
54267(4-3); 54368(6-18); 54381 (7-36a); 202(2-109); 209(2-63); 218(7-25); 219(2-8);
56066(2-79); 56077(7-I8); 56097(2-141); 230(8-2); 240(7-73); 263(2-8); 274(7-25);
56100(7-20); 56I39(2-114); 56160(2-30); 287(2-95); 327(7-56a); 401(2-8); 413(2-8);
56234(2-41); 56300(6-27); 56326(7-54); 419(7-25); 422(2-52); 423(2-52); 433(2-8);
56342(7-20); 56354(2-141); 56357(7-20); 466(4-3); 495(2-8); 1090(2-63); sn(6-i);
56400(2-41); 56442(7-19); 56470(2-29); sn(6-i); sn(6-2); sn(6-I5); sn(7-25); sn
56538(2-17); 56603(7-24); 56004(2-62); (7-49b); sn(I845(7-56a); sn(I862(7-56a);
5668i (2-43); 56696(2-129); 56954(7-24); sn(I864(7-56a); sn(2-I8); sn(2-43a); sn(2-
60076(7-73); 6o4IoA(7-2) 52); sn(2-63); sn(2-69a); sn(2-95); sn(2-99);
Majin, R. L., 58(7-75); 6i (2-39); 72(7-75) sn(2-io8); sn(2-I30); sn(2-I35); sn(3-I)
Malme, G. O. A., 76I(7-85); 1170(7-54); 1757 Mell, C. D. et al, 214(7-25); sn(7-56b)
(2-30); I757a(2-3o); 1773(7-54); I992a Mello, F. et al, INPA 1632(7-9); INPA I808(4-I);
(3-I6); 2028(7-19); 2028a(7-20); 2356 INPA 1816(6-41); INPA 2005(7-2); INPA
(6-27); 2378(2-I 4); 2703(2-28); 2494 2026 (6-41); INPA 2086(2-36); INPA 2116
(7-66); sn(3-I6) (6-9); INPA 2163(6-29); INPA 2852(7-67);
Markgraf, F. et al, 3038(6-25); 3696(7-35); 3238 INPA 3335(2-27a); INPA 3490(6-13); INPA
(7-20); 3415(7-54); 3816(7-63) 3931(6-29); INPA 394I(7-9); INPA 4317
Marsh, 299(1-1); 644(7-36a) (2-27a); INPA 4399(2-27a)
Marshall, 11912(2-86); I 930(2-69a); 12678 Mennega, A. M. W., I36(7-56a); 3i6(2-27a);
(2-86); 12688(2-69a) 331(2-63); 357(2-27a); 433(7-56a)
Martin, J., 157(2-27a); sn(i-i); sn(6-3o); sn Metcalfe, sn(I-i)
(6-34); sn(7-25); sn(7-56a); sn(2-52); sn Mexia, Y., 5409(2-114); 5460(2-114); 5473(7-85);
(2-69a); sn(2-76); sn(2-II4); sn(3-I); sn 5996(2-69a); 6047(2-129); 6060(7-56b);
(3-3); sn(3-4) 9236(7-56b)
Martin, R. et al, 444(7-20); 1676(6-9) Meyer, E., sn(2-27a)
Martins, 0., MG 902(2-44); MG 16524(6-2) Meyer, W. C., 126(7-56b)
Martius, K. F. P. von, 449(2-43a); 511(I-1); 911 Michaux, A., sn(2-I)
(2-30); 1467(2-37); 1722(7-38); 2921(6-36); Miers, J., 2283(7-85); 3432(7-85); 3776(7-36b);
3039(7-53); sn(6-3a); sn(6-9); sn(6-2o); sn 3814(2-114); 3889(1-I); 3890(2-11); 4094
sn(6-27);
6-20); sn(6-27);sn(7-2); sn(7-36a); (7-85); 4095(2-132); 5448(2-114); sn(7-36b);
sn(7-36b); sn(7-4o); sn(7-50oa); sn(7-56b); sn(7-85); sn(2-II4); sn(2-I32)
sn(7-7o); sn (2-69b); sn(2-696c; sn(2-8o); Mikan, J. C., sn(2-43a);
sn(2-Io4); sn(2-i2I); sn(2-I29); sn(2-I37) Mille, 1059(1-I)
Martyn, 273(2-35) Miller, G. S.,9(7-36a); I131(7-36a); I837(7-56b);
Mason, 13778(7-22) 2035(7-56b)
Masson, F., sn(7-36a) Milligan,J. M., sn(2-I)
Mathews, A., 1599(7-56a) Millspaugh, C. F., 284(1-1); 1265(I-I); 2175(I-1)
Mattos, A., 136(2-41) 2280(1-I); 9202(I-I)
382 Flora Neotropica

Miranda, F., I9I5(2-44); 4955(2-44); 6448(6-47); 243(6-27); R 7149(7-82); sn(7-2o); sn

7045(6-47); 7256(2-15); 7291(2-88); 7310 (7-36b); sn(7-56)
(7-56b); 7736(7-36a); 8126(1-I); 8324(7-36); Northrop,J. I., 115(1-I)
8324bis(7-36); 85o6(2-88a); 8545(7-56b); Novaes, J. de C., 690(7-85)
9062(2-I6) Nunes, G. M., 299(7-85)
Miranda, V. C., MG 3180(7-56b) Nuiiez, C., sn(2-44)
Mitchell, 3(1-I) Nuttall, sn(2-I)
Mociiio, J. M. et al, 87(7-56b); 876(7-56b); Nyst, P., sn(7-36a)
2282(6-47); sn(7-9I)
Mocquerys, 3(7-36a); 991(7-36a) Occhioni, P., 1157(6-25)
Mohn, S. H., 138(1-I) Ojasti, sn(6-3b)
Mohr, C., sn(2-I) Oliveira, E., 6(2-27a); 32(2-114); 56(7-56a);
Moldenke, H. N., 333(2-I); 751(1-I); 87I(I-I); ii8(2-69a); I43(7-56b); 230(2-130); 272
961(2-I); 5442(I-I); 5463(2-I) (2-69a); 356(2-57); 379(2-86); 493(3-4);
Molina et al, I8V42(6-3b) 519(7-63); 555(2-67); 623(2-99); 793(2-99);
Molina R., A. et al, 74(7-56b); 853(2-45); 1819 809(7-42); 8i I (7-56a); 823(2-99); 86i (7-69);
(I-I); 185I(7-56b); 1869(7-56); 2133(7-22); 895(2-57); 9II(7-56b); 9I6(7-56a); 1202
2383(7-22); 3637(2-15); 6o6I(7-36a); 6076 (7-24); I598(7-56b); 1735(7-54); I787(6-21);
(7-36a); 6905(7-22); II619(7-56b); 11807 1920(7-25); 2009(7-56b); 2092(3-4); 2130
(7-22); II899(2-45); 14790(1-I); 14918 (6-2); 2285(2-22); 2307(6-36); 2619(2-57);
(7-56b); I5204(7-56b); I8120(7-22) 2701(7-1); 2833(7-I); 2842(2-27a); 2873
Monteiro da Costa, R. C., I09(4-3); I32(6-3a); (7-56a); 2979(6-28); 3136(2-114); 3193
277(6-3a); 279(2-37); 317(2-27b) (7-56a); 3502(7-63); 3508(7-24); 3533(2-69a);
Montes et al, 712(6-47) 3549(7-15); 3580(3-4); 3595(7-10); 36oi
Moore, S., 78(7-54); '41(7-64); 564(2-28) (7-56); 3636(2-39); 3646(7-24); 3649(6-4);
Moraes, J. C. de, 692(7-56b); 892(2-44); 894 3735(6-32); 3768(3-4); 3792(2-43a); 3795
(2-43a) (2-69a); 3850(2-43a); 3853(7-I5); 3891
Moreira, 131(7-64) (2-43a); 3933(7-63); 4095(2-134); 4098
Moritz, J. W. K., 1227(1-I) (2-43a); 411 (6-7); 4179(3-4); 4500(2-43a);
Morrow, C. F., I48(1-I) 4563(3-2); 4575(3-I2); 4606(3-4); 4682
Morton, C. V., 5658(I-I); 9912(1-I); 9965(1-I); (7-I5); 4651(2-134); 467I(2-69a); 4702
ioo61(1-i); I0072(7-22) (7-42); 4706(3-2); sn(7-2o)
Mos6n, H., 2811(3-4); 2837(2-I26); 2838(7-85); Oliveira, J. E., RB 67975(6-27)
3377(6-52) O'Neill, H., 8624(7-56b); sn(I-I); sn(2-I)
Moses, T., 38(2-18) Onishi, E., 813(7-54)
Mosier, C. A., I(I-1); 2(1-I); 232(1-I) Orcutt, C. R., 3047(7-56b); 3232(7-56b)
Moss, A. M., sn(6-4) Ortega, J. G., 41(2-16); 58(6-47); 955(6-47);
Motta, M. da, R 7145(7-8I) 6198(6-47); 6285(6-47)
Muenscher, W. C. et al, 14092(1-1) Otero,J. T., M I8(7-83); 595(7-83); 72I(7-83)
Muller, 215(7-85); 322(7-85) Osmarino (P. Monteiro), INPA 20738(2-43b);
Murrill, W. A., 420(2-I); 588(2-I) INPA 20739(6-4)
Museu Goeldi, Belem, 5156(7-56a); 5465(7-56a); Otto, E., 969(2-27b); 970(6-3a); sn(7-36b)
9614(7-56a); 9704(2-86); 9721(6-50); 9747
(7-56b); 978I(2-141); 9792(7-27); 9824 P. in Forest Department, British Guiana, 50
(2-69a); 10I23(2-39); 10I8I(2-99) (2-113); 69-F.D. 2841(2-129)
Museu Nacional, Rio de Janeiro, 36006(6-25); Pabst, G., 4959(3-I6); 5219(3-4); 7232(7-85);
6323I (2-132) 7376(3-i6)
Mutis, J. C., 23(7-35); I109(7-35); 1114(2- 118); Palacios et al, 3543(2-144)
1130(2-118); II55(7-36a); 1157(7-56b); Palma, R 63195(7-85)
I375(7-36a); 355I(7-22); 3552(7-22); 3553 Palmer, E. J., I41(2-I); 142(I-I); I73(I-I);
(7-22); 3554(7-35); 3555(7-22); 3868(2-I i8); 342(2-45); 40I(6-47); 428(2-15); 469(I-I);
3969(7-56b); 5047(7-22) 485(1-I and 7-56b); 27280(2-I); 27327
Myers, J. G., 5894(2-134) (2-I); sn(6-2o)
Pannier, F. et al, Io98(2-69a)
Nadeaud, J., sn(7-36b); sn(7-85); sn(3-4) Paray, 3365(2-44); 4035(7-56b)
Name, T. Van, sn(i-i) Parker, sn(i-i)
Nascimento, 28(2-114) Passarelli, sn(7-36b)
Nash, G. V. et al, 62(1-1); 394(I-I); 734(2-I); Patris,J. B., sn(2-27a)
-I); I846(2-I) Paul, Bro., 22(1-I); 284(7-22); 412(7-22); 6ii
1302(
Neiva, A., SP 24061(1-I) (7-36a)
Nelson, E. W., 2363(2-45); 2372(6-47); 2604(1-I); Pavon, J., sn(6-47); sn(7-36a); sn(7-36a)
27I5(7-56b); 3853(7-56b); 4122(6-47) Pearce, sn(7-36a)
Netto, L. do S. M. et al, 32(7-56a); 242(2-I26); Peck, M. E., 21(1-I); 174(I-I); 348(7-56b);
List of Exsiccatae 383

367(1-I); 368(1-I); 47I(2-88a); 858(2-32); 6813(7-39); 6863(7-5ob); 6885(7-39); 6925

937(7-23) (3-3); 6943(7-39); 7004(7-39); 7037(2-8I);
Pelly, R. S., I I(7-56b); 85(7-56b) 7069(7-5ob); 7122(7-50b); 7424(3-3); 7495
Pena, B. S., 20(2-52); 36(7-56b); 44(2-27a) (6-35); 7972(2-27a); 8I2I (2- 14); 8I 74(1-I);
Pennell, F. W., 3637(7-22) 8237(7-20); 9058(3-16); 9059(3-I6); 9199(2-
Pennington, T. D. et al, 7326(2-I5); 9277(2-15); 80); 9403(7-54); 9546(2-43a); 9640(2-43a);
9326(2-15); 9337(2-45); 9477(2-45); 9478 9725(6-9); 986i(7-54); 9862(7-20); 9958
(2-45); 9632(7-22) (7-56a); I0071(7-36a); 10259(7-27); 10306
Pereira, E., 450(7-66); i833(7-54); 2034(7-20); (7-47b); 10366(7-56b); 10514(7-56b); 10542
3239(7-56b); 3356(7-24); 3530(6-25); 4033 (2-86); 10543(2-99); 10590(2-52); 10602
(6-25); 4281(6-25); 4652(3-16); 7409(7-54) (2-69a); O6I11(2-52); 10648(6-4); 10649
P6rez-Arbelaez, E., 592(1-i); 624A(6-36) (2-86); io650(2-69a); 10654(2-18); 10663
Perkins, I358(7-36a) (7-56a); 10678(2-124); Io689(7-53a); 10690
Perkins, A. E., sn(2-I) (3-3); 10703(4-3); 10727(7-20); 10869(2-81);
Perrottet, G. S., sn(8-I); sn(2-27a); sn(2-114) 10900(3-3); I0901(3-3); I0916(7-49b);
Persaud, A.
C., 5(7-25); 8(2-28); 13(2-109); 1094I (3-2); 10942(3-2); 11002(2-114);
28(7-25); 41(2-124); 58(3-I); 79(2-I24); io 64(2-27a); 11070(2-39); III4o(2-27a);
94(2-69a); 95(2-35); 97(2-28); 104(2-66); 11201(7-20); 11226(2-27a); 11237(2-86);
110(3-I); I4I(6-I5); I4I(6-7); 161(2-35); 11285(2-86); 11299(2-27a); INPA 13241
209(3-I); I6I7(7-56a) (2-27a); 50273(7-5ob); 50334(7-56a); 50349
Peters, H. M., sn(I-I) (3-4); 50418(7-20); 50435(2-87); 50448
Phelps, K. D. et al, 306(7-25); 407(2-I10); 514 (3-I); 50490(8-I); 50499(7-20); 50515(2-83);
(7-43); sn(7-2I) 50518(7-50b); 50557(2-109); 50650(7-50b);
Philipson, W. R. et al, I535(7-36a); I797(7-36); 50652(2-97); 5o69o(2-69a); 50762(2-87);
1887(7-76); 1969(7-76); 2085(7-36a); 2309 50765(6-19); 50843(6-7); 50903(2-97);
(7-76) 5090o(7-56a); 51044(2-69a); 51045(6-4);
Phillips, A. R., sn(6-47) 51055(2-134); 51089(7-i8); 51181(7-56a);
Pickel, B., i8i(7-56b); 201(1-I); 203(7-56b); 51198(2-149); 51232(7-20); 5I248(6-19);
603(7-56b); 831(6-49); 1175(2-II); 1878 51271(7-49b); 51324(6-19); 5140o(2-69a);
(2- I); 284I(-I I); 3420(7-20); 3947(2-44); 51407(2-69a); 51418(2-69a); 51470(2-63);
5570(7-85); 5705(2-144) 5I474(3-3); 51537(7-II); 51558(3-4); 518i6
Pinho, R. A., 6(6-27); 26(6-27) (6-4); 5I8I8(2-130); 5I82I(2-69a); 51830
Pinkus, A. S., 46(7-26); 6I (2-152); 52, F.D. 2834 (2-99); 51851(3-3); 51856(2-27a); 51867
(2-93); 62(2-113); 89(2-93); 180(2-42); (2-39); 51870(2-63); 5I884(2-II4); 51898
191(2-129); I98(2-I29); 245(6-42); 260 (i-i); 5I920(7-20); 52191(7-30); 52228
(2-129) (2-83); 52246(7-50a); 52276(2-37); 52290
Pinto, P. et al, 255(2-88a); 262(7-40) (2-56a); 52395(7-56a); 52459(2-69a); 52476
Piper, C. V., 5575(7-56b); 5596(7-56b); 5654 (3-4); 52534(7-i8); 52619(7-56); 52565
(7-22); 5713(2-45) (2-69a); 56747(7-56a); 56823(2-17); 56825
Pires, J. M. et al, 2i(2-57); 8I(2-27a); IOI(7-63); (6-27); 56948(6-27); 57065(7-54); sn(7-56b)
I35(6-9); I63(6-3a); I69(6-3b); 176(2-129); Piper, C. V., sn(i-I)
I80(2-I29); 22I(7-56a); 23I(7-56b); 3I5 Pittier, H., 1899(2-45); 2075(7-56b); 2192
(6-3b); 340(7-5ob); 361(2-99); 413(7-62); (7-56b); 2641(7-56b); 2680(7-36a); 2702
458(2-69b); 469(2-128); 478(7-20); 491 (7-56b); 2708(7-22); 2784(7-36a); 284 (2-
(7-2I); 508(7-63); 600(7-62); 652(2-69b); 45); 3347(2-I5); 3869(7-36a); 4I78(7-36a);
716(7-8); 738(2-8); 774(7-56a); 834(6-36); 5486(7-56b); 5491 (2-88a); 5523(7-36a);
889(7-53); 936(2-128); 102I(2-26); I034 6o72(7-36a); 6510(7-22); 6577(7-36a); 6971
(2-36); I351(7-40); 1428(2-8); I473(2-39); (2-15); 6998(2-I5); 7623(7-36a); 7827
1486(2-39); 1600(7-20); 1633(7-20); I658 (7-36a); 7999(3-7); 10103(3-7); 10602
(2-30); I7I8a(6-3c); I719(7-20); I721(7-18); (7-56b); 1o87(7-36a); 1140(2-12); 12073
1745(2-37); 1840(2-63); 2374(7-18); 2672 (2-27b); I2088(7-56b); 12012(2-15); 12137
(2-8); 3132(7-63); 314i(7-63); 3142(2-52); (7-56b); 12163(7-56b); 12164(7-56b); 12185
3I93(3-3); 3194(3-3); 3433(I-I); 3565(6-21); (i-i); 12230(2-12); I2358(7-56b); I2402(2-
3821(2-99); 3839(7-56b); 4132(2-43a); 4134 12); 12433(1-1); I3556(3-7); I5205(7-36a);
(2-99); 4146(6-4); 4333(2-39); 4350(6-3a); I5388(7-36a); 15640(2-145); 15742(2-145);
4355(2-27a); 4356(7-50a); 4401(4-I); 4402 15778(7-36a); 63556(3-7)
(2-129); 4579(7-24); 4594(7-56a); 4614 P16e, A., 14(7-22); 210(7-36c); 805(7-36); sn
(6-2); 4620(2-134); 4627(2-37); 4638(2-69a); (7-36a); sn(i-i); sn(i-i)
4640(3-3); 4695(2-97); 4812(2-43a); 4817 Poeppig, E., 2177(6-36); 2305(6-36); 2501(7-73);
(7-24); 4860(6-4); 4914(7-I5); 5I80(7-I5); 2502(7-40); 2503(6-3b); 2531(2-114); 2770
5624(2-52); 5802(2-142); 5826(7-50b); 6039 (2-43b and 2-129); 2785(2-75); 2877(2-27b);
(2-135); 6320(2-27b); 6325(2-37); 6399 2880(2-27b); 2985(2-I35); 2988(1-I); 3007
(2-129);673 (2-2o);6792(7-47b);68oo(3-I2); (3-I); 3067(2-129); sn(7-27)
384 Flora Neotropica

Pohl, J. E., 1275(7-20); 2181(7-66); 2226(2-37); (7-5ob); 586 (7-53); 5884(7-73); 6000(2-37);
2282(7-54); 2283(2-30); 4402(2-69c); sn 6008(7-5); 6o89(7-56a); 6130(2-43b); 6255
(7-20); sn(7-82); sn(7-85); sn(7-92); sn(3-8) (7-36a); 6286(7-56a); 6290(7-73); 6467
Poiteau, P. A., sn(8-I); sn(I-I); sn(I-I); sn(7-25); (7-56a); 6468(7-14); 6475(2-99); 6495(7-24);
sn(2-8); sn(2-II4); sn(2-134); sn(7-56a); 6505(7-56a); 65'2(2-43b); 6532(2-136);
sn(3-I) 6559(6-3); 6656(7-56b); 6692(2-17); 6722
Pollard, C. L. et al, 196(1-i); 247(1-I); 213 (6-20); 6731 (7-56b); 6734(6-34); 6744(7-73);
(7-36a); 252(2-I); I20(2-I) 6833(2-69b); 6846(2-43b); 6858(6-38); 6869
Porter, T. C., sn(I-I) (7-24); 6953(7-56a); 6965(2-69a); 7000
Porto, P. C., 306(7-85); 729(2-I44); 2450(6-20); (2-I 5); 7020(7-56a); 7260(2-69a); 7295
RB I55o6(7-85) (7-36a); 7308(7-66); 7323(7-56a); 7449
Potter, 5189(1-1) (7-66); 7537(7-66); 7610(7-56a); 7635
Prance, G. T. et al, 1265(6-2); 1271(1-I); 1276 (7-56a); 7785(2-43b); 79I7(7-56a); 7934
(7-56b); 1287(2-135); 1304(1-1); 1344(7-56); (6-24); 7983(7-66); 8146(6-16); 8155(6-37);
1403(7-63); 1474(7-24); I496(7-24); 1513 8186(7-66); 82I5(2-69a); 8237(2-141); 8252
(2-69a); I517(7-5ob); 1564(2-114); 1579 (2-69a); 8257(2-99); 836I (7-24); 8427(7-73);
(6-15); 1653(2-39); i69o(2-27a); 1701(6-2); 8464(7-24); 8521(7-6); 8693(6-20); 8758
I734(2-I34); 1752(2-63); I762(2-134); I882 (6-15); 8769(2-129); 8793(6-I5); 8796(6-I5);
(2-69a); 1931(4-3); 1932(2-114); 1938(2-86); 8803(2-69a); 88I2(6-3b); 8821(3-4); 8912
1951(2-69a); 1994(3-3); 2005(2-86); 2022 (7-24); 8936(7-56a); 8996(2-69a); 9038
(2-114); 2033(2-69a); 2050(2-97); 2069 (6-4); 9101(2-27b); 9113(6-43); 9II9(7-25);
(7-20); 2077(2-69a); 2090(6-41); 2102(1-I); 9157(2-27a); 9279(2-27); 9359(2-122); 9418
2103(1-I); 2Io4(7-36a); 2105(2-86); 21o8 (7-56b); 9503(3-I); 9512(2-27b); 9536
(7-25); 2109(2-69a); 2110(7-25); 2111(3-I); (7-25); 9537(2-43b); 9566(7-56b); 9580
2112(2-12); 2113(2-15); 2II3A(2-44); 2201 (6-43); 9582(2-27b); 9762(2-69a); 10121
(7-I I); 2212(6-15); 2314(7-56a); 2339(6-24); (2-68); 10156(7-2); 10268(4-3); 10365(3-4);
2432(7-56a); 2484(6-35); 2485(7-56a); 2489 10416(2-8I); 10444(6-3b); 10457(2-I5);
(2-57; 2504(2-43b); 2524(2-2 I); 2534(2-76); 58246(7-70); 58456(7-18); 58479(2-37);
2596(2-69a); 2617(2-69a); 2648(6-3b); 2649 58512(2-141); 58519(6-3C); 58524(2-43a);
(6-3a); 2739(6-35); 2962(2-27b); 2964(6-3b); 58564(7-20); 58569(2-30); 58642(7-56);
2968(6-44); 2978(2-52); 2999(7-39); 3006 58643(2-114); 58669(7-56); 58679(7-69);
(7-5); 3023(6-3a); 3067(2-88b); 3077(6-I5); 58727(4-3); 58731(7-56); 58745(6-4); 58750
308I(2-69a); 3099(2-69a); 3101(6-4); 3144 (2-114); 58753(3-3); 58762(I-I); 58813
(2-69a); 3154(2-39); 3157(3-4); 3212(7-24); (2-63); 58827(6-2); 58861(2-86); 58864
3220(7-6); 3235(2-69a); 3282(2-39); 3339 (7-69); 58865(3-3); 58877(2-86); 58944
(2-140); 3350(6-3b); 3356(2-129); 3361 (7-69); 58948(2-27a); 58959(2-69a); 58964
(2-27b); 3369(6-3b); 3406(7-20); 3470(2-98); (7-42); 58968(2-76); 58989(2-141); 5899I
3615(2-69a); 3633(6-9); 3641(6-50); 3651 (2-17); 58995(2-80); 58996(3-I6); 59043
(2-69a); 3672(2-140); 3675(2-27a); 3691 (6-27); 59052(7-54); 59055(7-20); 59II6
(2-140); 3692(2-27a); 3718(6-13); 3728 (7-20); 59 17(2-30); 59127(7-54); 59129
(2-69a); 3747(6-3a); 3850(6-9); 3967(2-43a); (2-27b); 59144(6-20); 59I46(7-54); 59194
3970(7-56a); 3974(2-27a); 4017(7-25); 4020 (2-28); 59I95(7-19); 59212(2-37); 59394
(6-3b); 4039(2-129); 4049(2-27b); 4123(7- (2-27b); 59404(2-37); 594 I(2-27b); 59426
56b); 4131(3-4); 4133(2-27a); 4196(2-43a); (7-64); 59429(7-20); 59434(7-54); 5944I
4204(4-3); 4210(2-122); 4224(2-122); 4235 (2-30); 59471(2-114); 59514(2-30); 59553
(2-43a); 4270(3-4); 4271(2-27b); 4352 (3-16); 59614(2-114); 596I6(2-27b); 59622
(2-122); 4355(4-3); 4366(3-I); 4507(2-81); (2-27a); 59627(2-27b); 59629(2-27a); 59637
4520(2-43a); 4524(2-8I); 4543(7-56b); 4562 (2-27b); 59705(3-i6)
(7-36a); 4576(2-43a); 4590(2-I8); 4601 Prendleloup, L. A., I70(7-36a)
(3-I); 4624(2-114); 4636(6-13); 4638(6-41); Pringle, C. G., 6629(1-1)
4660(2-27a); 4696(6-11); 4706(6-15); 4732 Prior, R. C. A., sn(i-i)
(2-140); 4734(6-3a); 4746(2-88a); 4755 Proctor, G. P. et al, 3322(1-1); 923I(7-36a);
(6-29); 4756(6-3b); 4768(2-27a); 4807(6-49); 15223(1-1); I7535(1-I); 18778(1-1); 18921
4808(2-114); 4809(6-51); 4810(6-24); 4811 (7-36a); 19419(7-36a); I9645(7-36a); 19855
(2-II); 4875(2-88a); 4886(2-140); 4934 (i-i); 20997(7-36a); 213I6(I-I); 21621
(6-13); 4937(2-88a); 4984(7-1); 4992(2- 24); (1-2); 23363(7-36a); 23954(7-36a); 27045
5012(2-60); 5013(2-39); 5050(2-119); 5081 (7-22); 27119(7-36c); 27213(7-56); 27332
(2-88a); 5108(2-124); 5242(2-43b); 5244 (7-22)
(7-56a); 5307(7-56a); 5360(7-56b); 5404 Pulle, A., 56(2-135); 143(2-I35); 152(2-135);
(7-53); 5434(7-5ob); 5464(6-20); 5486(7-56a) 383(7-36a); 439(2-134); 452(2-77); 1130
5546(2-99); 5579(2-43b); 5624(2-43b); 5671 (7-36b)
(7-36a); 5678(2-43b); 57I5(7-56a); 575I Purdie, W., I05(2-69a); sn(7-25); sn(7-36a);
(2-43b); 5769(7-73); 5793(7-53); 58io sn(2-12); sn(2-86)
List of Exsiccatae 385

Purpus, C. A., 51(6-47); 342(1-1); 9183(2-45); Robert, A., 388(3-I6); 490(7-66); 490b(7-66);
0oo51(6-47); 10512(1-I); 10531(6-47); 497(2-41); 497b(2-41); 530(3-16)
12074(6-47) Robertson, 401(6-47); 5343(1-1)
Purseglove, J. W., 6387(I-I) Rodham, 0., sn(2-I)
Pursell, R. A., 9067(I-1) Rodrigues, J. B., 200(7-5oa); 202(7-18); sn(7-I8)
Rodrigues, J. S. et al, 87(2-140); 157(6-15)
Rodrigues, R. S., MG 4029(7-56a); MG 4030
Quelch et al, 308(2-27b)
(7-24); MG 4043(2-134); MG 4044(2-134);
Quentin, L. et al, 5541(1-I); sn(7-36a) MG 8274(6-4); MG 8275(2-86); MG 8276
Questel, A., 124(I-I); 754(2-I3); 876(2-13); MG 9644(6-4); MG 9647(2-69a); MG 9648
2027(7-36); 2302(7-36a); 2558(7-36a); 4089
(3-3); MG 9650(2-81); MG 9651(6-49);
(7-36a); 4745(2-85); 51 14(7-36a)
Quintero, L. G., 583(7-56b) (2-99); MG 8761(2-57); MG 8772(2-63);
MG 9684(3-3); RB 19776(2-69a)
Quir6s, C. M., 858(7-56b)
Rodrigues, W. A. et al, 207(2-27a); 273(6-3a);
288(6-3a); 289(2-27a); 300(2-119); 333
Raddi, G., sn(7-36b); sn(7-64); sn(7-85) (6-36); 446(2-27a); 538(7-73); 7i8(6-3b);
Ramage, G. A., sn(I-I); sn(7-32); sn(2-I3); 722(2-27a); 902(2-140); 957(6-I5); II62
sn(I888(2-85); sn(I889(2-85) (6-29); 1308(2-114); 1360(6-9); I404(2-I29);
Rambo, B., 37841(7-85); 38606(7-85); 39047 1408(8-I?); I423(2-43b); INPA I602(6-41);
(7-85); 39194(7-85); 40100(7-85); SP 5I365 1715(6-15); I778(7-5ob); I8o8(2-43b); I884
(7-85) (2-43b); 1889(1-I); 2040(2-27a); 2049(4-I);
Ramia, V., 586(2-12) 2053(7-8); 2io7(6-3a); 2I 0(7-56b); 2148
Ramos, J. F., 6653(7-20); 6718(7-54); 6719 (6-9); 2154(6-41); 2165(6-II); 2182(6-3a);
(7-70) 2206(2-129); 2219(6-3a); 2227(2-69a); 2238
Randolf, F. R., 108(2-I) (7-I); 2246(6-29); 2332(6-41); 2359(7-28);
Rauh, W. et al, I595(7-56a) 2368(7-15); 2390(6-15); 2476(6-3b); 2486
Ray, J. D. et al, 10898(2-I) (7-24); 2491(7-8); 2536(2-69a); 2577(7-56a);
R. B. (Botanical Garden, Rio de Janeiro), 19793 2659(6-29); 2686(6-9); 2699(3-2); 2739
(2-97) (6-29); 2770(6-I ); 2771(6-9); 2772(2-26);
Record, S. J., BH 33(7-56b) 2772a(2-64); 2785(6-9); 2805(2-26); 2946
Registro, 2(7-56b); 44(6-4) (2-99); 2948(2-69a); 2952(2-43b); 2965
Regnell, A. F., 54(7-64); 111-150(7-54); III-I5I (6-iI); 2974(6-I); 3168(7-15); 3286(7-24);
(7-20); III-152(6-27 and 2-41); III-I52A 3291(7-18); 3304(2-26); 3402(7-56b); 3486
(2-41) (6-9); 3571(6-9); 3585(6-9); 3863(2-26);
Rehder, A., 766(1-i); 767(1-I); 826(2-I); 822 3920(7-24); 3921(6-3b); 3970(2-123); 3998
(I-I) (2-123); 4051(6-29); 4058(6-9); 4086(2-26);
Reitz, P. R. et al, 1690(7-85); 3059(7-85); 3234 4105(2-26); 4I20(2-27a); 4121(7-24); 4196
(7-85); 5778(7-85); 5896(7-85); 5973(7-85); (2-26); 4552(7-I); 4649(6-41); 4697(6-41);
6091(7-85); 8067(7-85); 8640(7-85); 9278 4776(2-69a); 4786(2-27a); 4795(7-24); 4816
(7-85) (6-3a); 4827(2-69a); 4856(2-27a); 4857
Reko, 3224(6-47); 3405(7-56b) (6-3a); 4881(2-123); 489I(6-4); 4918(I-1);
Restinga I. (Segadas-Vianna et al.), 722(6-25); 4919(2-26); 4998(6-3a); 5003(2-69a); 5004
757(1-I); 803(1-I); 878(1-I); 1040(6-25) (2-27a); 5018(2-26); 5360(2-69a); 5367(6-9);
Rhoades, A. A., 8427(I-1) 5403(6-4 and 7-2); 5426(7-2); 5427(7-2);
Richard, L. C., sn(I-I); sn(6-7); sn(2-95); sn 5436(2-124); 5445(6-44); 5497(6-3b); 5512
(2-109); sn(2-I25); sn(3-3) (7-41); 5520(2-62); 552oa(2-8i) ;5523(7-67);
Richardson, W. D., 644(3-1); 915(I-1); 931 5557(2-39); 5563(2-124); 5572(2-112); 5689
(7-56a); 12668(2-86) (2-27b); 5699(2-27b); 5744(6-22); 5780
Ricksecker, A. E., 353(1-1) (4-I); 578i(7-24); 58i9(2-43b); 5828(2-17);
Ridley et al, sn(7-56b) 5829(6-28); 5830(6-34); 5832(6-34); 5844
Riedel, L., 459(3-I6); 482(2-41); 504(6-27); (2-53); 5937(6-9); 594I (2-39); 5947(7-78);
518(2-127); 567(7-36b); 598(7-19); 68i 5952(2-43b); 5974(2-43b); 6007(3-4); 6012
(6-31); 702(6-27); 802(7-54); 1028(6-27); (6-4); 6o28(2-69a); 6034(6-50); 6o35(6-3a);
'135(7-66); I304(2-69a); I375(7-73); 1406 6720(6-3b); 6735(2-41); 675I(2-69a); 6757
(4-i); 143I(2-36); 1529(7-18); i578(7-5oa); (6-41); 6775(6-3a); 6781(2-27a); 6814
I665(7-85); 1666(7-64); I667(3-4); 2218 (2-69a); 6823(2-69a); 6851(2-69a); 688i
(2-41); 2319(3-4); sn(i-i); sn(i-i); sn (2-69a); 6886(2-43b); 6894(2-64); 6907
(6-25); sn(6-32); sn(4-I); sn(7-2o); sn (7-56b); 69 1(2-93); 6924(6-50); 6928
(7-36b); sn(7-5oa); sn(7-64); sn(7-82); (2-124); 6941(2-81); 6944(2-124); 6946
sn(7-85); sn(2-25); sn(2-43a); sn(2-4) (2-39); 6951 (2-43b); 6962(6-28); 6969
Riedle, A., sn(7-83) (2-43b); 6975(7-15); 6994(6-44); 6997 (2-93)
Rimbach, A., 59(7-35); 97(7-35) 7o40(2-69a); 7097(7-24); 7102(7-15); 7103
Rizzini, C. T., RB 12994(7-20) (2-49b); 7I06(6-44); 7113(2-41); 7116
386 Flora Neotropica

(6-4I); 7123(2-69a); 7129(3-2); 7135(6-9); Saldanha, J. et al, 655(3-4); 4993(3-4); 5430

7138(2-129); 7170(6-15); 7233(7-I); 7245 (2-II)
(2-124); 7266(2-62); 7295(2-27a); 7357 Salvoza, F. M., 865(7-36)
(2-69a); 7392(6-41); 7398(2-62); 7605(6-4I); Salzmann, P., sn(2-6)
7638(6-28); 7698(2-79); 7699(3-2); 7700 Sammler Herb., 117(7-74)
(2-69a); 7701(6-50); 7702(2-124); 7771 Sampaio, A. J. de, 102(7-54); 4978(6-3a); 5133
(2-23); 7806(2-41); 7883(6-4); 7893(6-4); (7-73); 5206(6-3a); 5251(6-3a); 5451
8249(7-I); 8319(2-81); 8353(2-27a); 856I (2-27b); 5547(2-I34); 5549(7-20); 555oA
(7-56a); 8587(2-69a) (7-20); 56ioA(i-i); 5636(2-134); 5656
Rodriguez, L., 2749(7-36); 3218(1-I); 3403(1-1); (7-25); 5712(2-134); sn(2-52)
3620(1-1); 472I (7-36a) Samuels, J. A., 129(7-25); 223(2-135); 298(7-25)
Rogers, sn(2-I2) 518(7-56a)
Rohr, J. P. B. von, sn(7-25); sn(7-74) Sandeman, C., 3406(7-50b)
Rombouts, J. E., 2736(7-54) Sandwith, N. Y., 92(7-74); 139(3-12); I4o(2-69d);
Romero C., R., 255(7-22); 622(7-36a); 787 I84(3-I); 253(7-56a); 26i (2-109); 285(7-25);
(7-36a); 789(1-I); 817(7-22); 868(3-7); 299(2-109); 315(6-7); 327(2-90); 336(2-77);
1179(2-15); 1279(7-56a); 1547(7-56a); I729 341(2-77); 433(2-69d); 586(2-112); 604
(7-22); 22II(7-1
(7-); 2598(7-56a); 349(7-); (2-69a); 687(6-45); 711(2-112); 1044(2-135);
3474(7-25); 3489(7-25); 3497(6-9); 3562 1236(2-108); 1246(7-5I); I72I(7-72); 1737
(7-56b); 3575(7-39); 3964(2-43b); 4930 (7-36a)
(7-5ob); 4948(7-56a); 5368(I-I); 555I(I-I) Santoro et al, I0032((2-44)
Romero, M. de, sn(I-I) Santos, E., Io89(7-56b); 1092(2-7Ia); I695(3-16);
Rondon Commission, 2450(3-4) 27085(3-I6)
Ronna, A., Io5(7-36b) Santos, J. V., 2870(7-36a)
Rosa, F. R., 57(2-80) Santos, R. S., 24230(2-41); 24388(7-20)
Rosa, P., 129(7-85) Sargent, 893(7-83)
Rose, J. N. et al, 1515(6-47); 3I20(6-47); 3310 Sargent, A. R., sn (2-1)
(6-47); 4244(7-36a); 14430(6-47) Sargent, C. S., sn(i-i)
Roth, L., 894(2-144) Sargent, F. H., A29(I-I)
Rothrock,J. T., 228(1-I) Sastre, C., 238(7-25)
Rovirosa, J. N., 85(7-56b); I79(6-47); 729(1-I) Sauer,J. D. et al, 2509(1-1); 3273(I-I)
Rowlee, W. W. et al, 120(1-1) Schack, B. von, sn(i-i)
Rudge, sn(2-39) Schallert, P. O., 1014(2-I); 7428(2-I); 7498(2-I)
Rugel, F., 217(2-I); 474(2-I); sn(I-i); sn(2-I) Schenck, H., 2303(7-36b); 4123(7-56b)
Ruiz, H. et al, sn(6-38); sn(7-55) Schipp, W. A., 33(7-56b); 65(I-I); 82(7-22);
Rusby, H. H. et al, 88(7-25); 26I(7-56a); 423 102(2-32); 0o3(2-88a); 292(2-88a); 308
(2-109); 423a(3-I); 426(2-27a); 565(7-36a); (2-15); 569(6-47); 598(2-32); 660(7-22);
590(7-36a); 652(7-46); 702(56b); 703(7-21); 717(6-47); 932(1-I); o098(7-23); 1257
704(7-56a); 768(7-21); 1130(7-22); I222 (7-36a)
(7-66); I37 (7-54); I448(7-36a); i600 Schlim, L. J., 898(7-22)
(7-36a); 2442(2-21); 2492(7-36a); 2678 Scholt, 579(6-47)
(6-3b); sn(3-I) Schomburgk, Rob. & Rich, 7(7-25); 23(7-56a);
Ruschi, A., SP 45069(2-25) 28(6-45); 39(1-I); 41(2-134); 50(2-93);
Russell, 12240(7-36a); 12470(7-36a) 54(2-112); 65(4-I); 8o(7-56b); 92(1-I); 93
Ryan, J., sn(7-36a) (4-i); 111(2-I34); 112(6-43); 113(7-18);
Rzedowski, 14383(7-56b); 16566(6-47) 132(7-21); I35(7-25); I36(2-27b); i68
(7-45); 220(1-I); 230(2-27b); 244(6-45);
Saer de H.,J., 71 I(7-56b); 793(7-56b); sn(7-36a) 252(2-27b); 309(6-43); 321(6-43); 342
Safford, W. E. et al, 49(1-I) (2-134); 349(2-27b); 356(6-45); 388(2-135);
Sagot, P., 202(I-I); 203(2-7); 204(3-I); 205 402(2-27b); 463(2-66); 485(6-I5); 519
(7-25); 262(6-2); 462(2-63); 792(7-50a); (2-116); 535(3-I); 564(3-I); 570(7-49); 593
843(2-69a); 915(7-56a); 917(7-56a); 968 (2-27b); 601(2-113); 728(2-114); 785(3-4);
(2-I 4); 969(2-43a); 974(2-69a); o08I(2-86); 822(2-95); 824(2-69d); 840(7-56a); 848
II09(2-63); 128I(2-134); I382(7-73); sn (3-I); 865(7-49); 868(2-134); 873(2-69a);
(8-i); sn(2-74) 886(7-24); 897(2-27a); 905(2-8); 906(2-27a)
Sagra, R. de la, sn(7-22); sn(i-I) 9o8(2-69a); 910(2-140); 935(2-II3); 976
Saint George Expedition, 583(7-56b); 607(7-56) (2-I21); 977(2-129); 980(3-I); 986(2-69a);
Saint Hilaire, A., I(7-36b); 7(7-36b); 12(3-4); oo006(2-8); 1051(7-20 and 7-49); I279
23(7-86); 293(6-25); 499(2-41); C 499 (2-134); I357(2-69a); 1361(2-95); 1382
(2-126); 542(7-36b); 850(7-20); 869(3-I6); (2-8); I443(2-69d); 1599(2-69a); 17I9(2-8)
982(6-27); 1654(7-70); 1714(6-27); 1783 Schott, H. W., 4222(3-8); 4696(7-36b); 4970
(7-70); B i967(2-126); 1973(7-20); 2062 (7-82); 4972(7-85); sn(7-85); sn(6-25); sn
(2-43a) (6-26)
List of Exsiccatae 387

Schreiner, C., sn(7-64) 7I5(1-I); 1078(1-I); 0o79a(I-I) 164o(7-36a);

Schubert et al, 486(7-83) I760(7-36a); 2934(1-I); 3165(1-I); 3559
Schultes, R. E. et al, 85I(7-36a); 948(7-56b); (7-83); 3563(7-83); 3676(7-36a); 3995(1-I);
992(3-I); 3566(6-36); 3854(7-40); 3977 4404(7-36a); 7720(1-I); 8302(1-I); 10772
(2-69b); 5453(2-104); 6731(6-44); 8266 (i-i); iIOi8(I-I); 11718(7-22); 11759(1-I);
(6-38); 8288(2-27a); 8415(7-56a); 8588 Shank, P. J. et al, 4344(1-I); 4559(7-56b); 4776
(2-21); 8749(7-56a); 8895(2-69b); 8941 (7-36a)
(6-2); 8989(6-36); 9246(2-69b); 9280(2-69a); Shannon, W. C., 5034(7-56b)
9344(7-62); 9400B(7-6); 9472(2-69b); 9481 Shattuck, 0., 519(7-36a); 575(2-15); 823(2-88a);
(2-140); 97I9(2-88b); 9748(2-84); 9948 949(7-36)
(2-69a); 9958(8-3); 10029(3-I); IOI83(6-29); Shimek, B. et al, 95(1-I)
I0333(6-3b); I2558(7-40); 12669(7-28); Short, C. W., sn(2-i)
12678(7-56b); I2832(2-88a); I2788(2-88a); Sick, D. H., B34(6-20); B505(7-54); B7I7(7-56a)
12977(2-121); I3085(2-27a); I3145(6-2); Sidney (G. da Fonseca), 191(7-54); 221(2-37);
I3392(6-9); 13536(2-88a); 13622(7-28); 268(2-141); 269(7-20); 279(2-30); 298
13772(7-56); 13807(7-40); 13818(2-94); (7-54); 983(2-41); 985(6-27); 1049(7-20);
I3846(2-105); 13995(7-56a); I4167(7-56a); 1053(7-54); I 98(2-27a); I24o(2-27a); 1261
I4522(7-5); 14577 (3-6); 14693(2-I40) ; 4962 (6-27); I28I(2-141)
(6-9); 15291(6-29); I5311(7-53); I5563 Sieber, F. W., sn(6-3a); sn(2-43a); sn(7-18);
(2-137); I5599(2-17); I5662(6-34); I6004 sn(7-56a); sn(7-56a); sn(i-I)
(2-121); I6459(7-40); 16466(6-2); I6612 Silva, A., 95(7-56b); 143(7-63); i66(7-56b);
(7-40); I66I5(2-69b); I6662(2-27a); 16977 254(7-56b); 285(7-27); 474(2-113); 489
(2-121); I7025(2-69b); I7325(6-9); I7479 (7-I8); sn(2-8)
(6-35); 17644(6-2); 18059(7-5); 18136(6-36); Silva, A. de A., RB 13623(7-64)
18140(7-21); 18405(7-40); I9514(7-37); Silva, J. F., 28(7-50b); I69(7-73); I73(2-88a);
19768(2-69b); 24172(7-55); 2433I(7-56b); 384(7-56b); 400(7-56b)
24514(6-3b); 245i8(7-56a); 24559(2-43b); Silva, M. B., 72(7-63); Io4(7-56a); I07(2-27b);
46-153(6-38); 46-342(6-7) 19I(2-27b); 247(7-25); 343(2-44); 367
Schulz, J. P. et al, 498(7-10); 7147(7-42); 7212 (7-56b); 596(7-56b); III3(7-24); 1350
(6-i8); 7403(7-56a); 7500(7-74); 7630 (7-56b); 1455(7-24); 1486(7-56b); 1488
(2-43a); 7799(6-i8); 7947(2-I8); 8013 (7-56b); I520(7-56b); I627(7-5oa); I659
(7-42); 8933(6-42); 00oo50(6-18); ioio6 (7-5oa); 1708(7-24); 1721(7-24); 1915
(4-3); o1067(6-I8); 10343a(2-1o8) (7-56b): 2042(6-44); 2048(6-44)
Schunke,J. M., 23(7-40); 53(2-27a); 15I(2-52); Silva, N. T. da et al, 62(6-4); 116(7-63); 117
288(2-23); 328(7-21); 329(7-21); 344(2-7); (7-24); I78(7-36a); 222(7-56a); 322(7-5oa);
346(2-69b); 350(2-26) 323(6-4); 337(2-8); 346(7-24); 382(7-18);
Schwabe, W., sn(7-4o); sn(2-69b) 392(2-92); 405(2-27b); 479(2-124); 488
Schwacke, C. A., III-I05(2-69a); III-I5I (7-56a); (6-15); 496(2-I24); 554(2-43a); 590(2-99);
III-i63(2-27b); III-209(6-i6); III-292 593(6-4); 597(2-76); 611(2-76); 612(7-50);
(2-82); III-364(6-i6); 111-523(6-9); III- 6i3(7-39); 637(2-27a); 647(7-I8); 651 (2-2I);
542(7-47); III-641(6-3a); 68(7-85); 256 674(7-20); 676(6-21); 709(2-30); 711(7-54);
(3-4); 385(2-129); 393(7-56b); 434(2-36); 76I(7-56a); 811(7-54); 907(2-27a); 910
7Io(2-69b); 76I(2-69a); 3745(2-69a); 4242 (7-42); 926(3-4); 939(2-27a); IoIo(2-27a);
(6-26); 4256(3-4); 4476(2-41); 6653(6-46); o012(7-5ob); 1036(7-42); IOI(7-24); 1o06
8722(7-81); 10235(6-46); 13682(6-46); sn (2-43a); 1130(6-7); 1133(6-4); 1138(7-24);
(3-4); sn(2-43a) I I45(2-43a); i6i (7-5ob); 1167(7-50b);
Schwaneike, C., sn(7-83) I334(3-4); 1336(2-69a); I376(7-50b); 1442
Schwebel, 19(6-46); 1191(6-27); 11-81(2-143) (7-24); 1463(2-I 14); 1533(7-63); 1594(2-39);
Schweinfurth, sn(i- i) I602(2-69a); I69o(2-27a); 59729(2-22);
Scolnik, R. et al, I9An 549(I-I); II8I(7-56b) 59733(6-15); 60609(7-25); 60693(7-20);
Seemann, B. C., 291(7-56b); 508(2-45); 544 6o694(2-27a); 60705(2-109); 60791(2-119);
s n(I-)sn(
(7-22); 1095(I-1); n(7-o);sn (2-45) 60774(2-64); 60966(2-119); 60974(2-140);
Segadas-Vianna, 527(6-24); 2731(6-25); 3796 60976(2-119)
(6-25); 3813(6-25) Silva, 0. A. da, RB 52231(8-1)
Seler, G. E., I660(2-45); 1782(1-1); I992(2-45) Silva, P. da, 27743(2-6)
Sellow, F., 44(7-20); 299(3-4); 2043(7-85); 5713 Simmonds, N. W., 15018(7-25)
(2-144); sn(7-3I); sn(7-36b); sn(7-8I); sn Simpson, J. H., 284(1-1); 545(2-I)
(7-85); sn(2-4I) Sintenis, P. E. E., 20oB(7-83); 232(7-83); 587
Sena, Herb. Schwacke, 2457(7-54) (I-i); 965(7-36a); 965B(7-36a); 1059(7-83);
Senn, H., 35I(7-36a) I668(7-36a); 2I45(7-36252(7-52(7-36a); 267
Servico Florestal Sao Paulo, 31(7-85) (7-36a); 4307(7-36a); 4968(1-I); 5175
Sess6, M. et al, 2129(1-I) (7-36a); 5256(7-83); 5396(7-83); 5735(1-I);
Shafer, J. A., 378(1-1); 525(7-36a); 710(I-1); 5793(1-1); 6067(7-83); 6645(1-1)
388 Flora Neotropica

Skehan, sn(2-I) (2-129); 3314a(2-I40); 344I(2-I40); 3458

Skutch, A. F., 2o88(7-36a); 3948(7-22); 4130 (2-81); 3490(2-69a); 3503(2-69b); 3523
(7-22); 4810(7-22); 5388(7-36a) (7-5); 3533(7-72); 3539(2-68); 3548(7-49);
Small, J. K. et al, 13(1-I); 129(1-1); 7I11(2-1); 3681(6-2); 3696(2-88a); 3760(2-91); 3825
1315(I-I); 1448(1-I); I449(I-I); 2106(1-I); (6-4I); 4049(6-22); 4614(6-36); 5067(6-33);
2I08(I-I); 2110(I-I); 2479(2-I); 3166(1-I); sn(7-2); sn(7-I8); sn(7-5oa); sn(7-56a);
3312(1-I); 3377(2-I); 3383(2-I); 4130(1-I); sn(2-27b)
6505(2-I); 6559(2-I); 6923(2-I); 7244(1-1); Stahel, G., 4I(2-69a); 43(1-1); 6I(2-27a); 6ia
8760(1-I); 9988(2-I); II350(2-I); 11508 (2-43a); 80(2-124); 8oa(2-I3o); 84(3-I);
(2-I); sn(2-I) 86(7-5ob and 2-I24); 141(2-135); 143(2-63);
Smith, Au., I779(2-54) I50(2-88a); 158(2-66); I58a(2-66); 206a
Smith, A. C., 2138(2-93); 2I89(7-56b); 2226 (7-5ob); 257(2-75); 281(2-98)
(2-93); 2270(2-122); 2502(6-43); 2570 Stahl, A., 434(7-83)
(2-134); 2607(7-50b); 2609(4-3); 2653 Standley,J. P., 176(2-1); 302(1-1); 495(2-I)
(2-134); 266o(7-56a); 2696(2-28); 2718 Standley, P. C. et al, 91(2-45); I276(2-45);
(7-67); 2839(2-100); 2923(2-122); 2992 I290(2.45); 3143(2-45); 3401(2-45); 3564
(2-57); 2999(7-7); 3024(2-I34); 3095(2-108); (2-45); 5847(2-45); 7338(2-45); 9170(2-45);
3o07(2-43a); 3222(7-56b); 3275(2-27b); I 284(I-1); I 517(2-15); I8222(7-22); I8815
3302(2-43a); 3320(3-3); 3409(7-25); 3420 (2-15); 19358(7-22); 19492(1-1); I9666
(2-77); 343I(7-36a); 3519(2-27b); 3559 (6-47); 20035(2-45); 20057(7-56b); 201I6
(4-3); 3563(2-77); 3590(7-36a); 3625(7-56a); (7-56b'; 20o52(2-88a); 20327(2-45); 20429
10036(7-74); Ioo83(7-56a); i0507(7-36a) (2-45); 20532(2-15); 21206(2-45); 21410
Smith, C. E. et al, 3263(7-56b); 4104(6-47) (6-47); 22023(1-I); 22149(6-47); 22228
Smith, C. L., 1(1-1); Io81(7-56b) (2-15); 23190(6-47); 2370I(7-56b); 24107
Smith, H. H. et al, 198(1-I); 443(7-36a); 478 (7-56b); 24558(2-45); 25ro6(2-88a); 26366
(2-13); 748(7-36a); 811(1-1); 850(7-22); (7-56b); 27036(7-56b); 2737I(7-56b); 28017
85oa(7-22); 1566(2-13); 1730(2-85); 1773 (7-56b); 28370(7-56b); 29736(7-56b); 30107
(2-31); I775(3-7); I838(2-I3); sn(7-2o); (7-56b); 31135(7-36); 3I323(2-88a); 31742
sn(7-66) (7-56b); 4Io67(2-88a); 4455I (7-36a); 44901
Smith, J. D., 1417(1-1); 1481(1-1); 1804(1-I); (2-45); 45667(7-56b); 45719(7-56b); 45782
I812(7-56b); 8680(7-23); sn(2-I) (7-36a); 459i6(7-36a); 46556(6-47); 46634a
Smith, S. G. et al, 1447(7-22) (7-36a); 53007(1-I); 53774(2-88a); 53845
Snethlage, E. H., 7I(7-27); 99(7-56a); I28 (2-15); 53997(1-I); 54273(7-56b); 54286
(7-56b); I56a(i-I); 305(7-56a); 623(2-30); (7-22); 54290(6-47); 55732(2-15); 56606
661(7-54); 669(2-37); MG 9498(6-2); MG (7-22); 56685(7-22); 57763(I-I); 60I38
0oo90(7-56a) (2-45); 72343(7-56b); 726i5(2-88a); 72644
Sobrinho, V., 80(2-44); 614(6-49); sn(6-49); (2-15); 73038(1-I); 73052(2-88a); 73122
sn(7-56b); sn(i-i); sn(2-44) (7-56b); 73I39(7-56b); 75388(2-45); 76037
Soti, 12(6-26) (2-45); 78957(7-56b); 79082(2-88a); 87330
Sousa, M., sn(7-56b) (6-47); 87349(7-56b); 87457(6-47); 87466
Spence, 52(3-1) (7-56b); 8780 (7-56b); 89034(6-47); 89271
Splitgerber, F. L., 210(1-1) (2-45); 89549(6-47)
Sprague, T. A., 157(7-22) Steam, W. L., 1488(1-1)
Spruce, R., 117(7-24); I33(2-43a); 139(2-63); Steere, W. C., I900(1-I); I904(1-I); 3398(1-I)
174(6-2); 222(7-56a); 313(6-37); 322(7-5oa); Stehl6, H., 936(7-32); i8i3(7-36a); 2524(7-36a);
382(7-18); 457(2-57); 534(2-27a); 7I9(6-3a); 2763(7-36a); 4380(1-i); 5573(7-36a); 6519
840(2-37); 844(6-36); 877(2-43a); 987(2-37); (i-2); 6805(1-2),
Io14(6-3a); 1042(7-56b); 138(6-37); 1179 Steinbach, J., 2183(7-36a); 6449(7-36a)
(2-27a); I230(2-27a); I414(6-I1); I423 Stelling, 8410(2-145)
(6-i6); 1437(2-27a); I438(7-2); 147I(2-69a); Stern, W. L. et al, I33(7-36a); I35(2-45); 337
1496(6-I5); 1536(6-4); 1569(2-57); I576 (2-I); 7I7(7-36a); 725(7-36a); I7I3(7-36b);
(2-8i); I597(3-4); I635(2-8I); I649(2-69a); I798(7-22)
I708(7-50b); I752(6-3b); 1776(6-29); i8oi Stevens, E. P., sn(7-25)
(2-36); 1811(4-1); I83o(2-27a); I831(2-27a); Stevenson, J. A., 99(I-I); 245(1-I); 850(7-83)
1953(2-129); 2003(6-15); 2197(2-69b); 2262 Stevenson, N, S.. 8(7-56b); 86(7-36a); 162(7-23)
(6-2); 2268(7-55); 2318(6-9); 2322(6-29); Steyermark, J. A., 4(2-129); 8(2-69a); 47(7-21);
2367(7-62); 2422(2-94); 2481(7-84); 2490 62(6-o0); 26o(2-69a); 293(2-56); 484(2-69a);
(2-121); 2520(7-40); 2539(3-6); 2676(2-75); 1315(2-69a); 17500(7-56b); 30236(2-45);
2678(2-107); 2690(2-124); 2699(2-33); 2707 31399(2-45); 34529(1-1); 37783(1-1); 38046
(2-27a); 2762(2-75); 2885(2-129); 2904 (2-88a); 38o57(7-56b); 38357(2-32); 38717
(7-60); 3022(6-36); 3023(2-69b); 3072(6-2); (i-i); 38842(2-88a); 28982(7-56b); 39432
3084(7-21); 3095(7-28); 3196(7-40); 3232 (I-I); 39449(7-22); 39509(2-88a); 39514
(2-39); 3278(2-69a); 3302(2-43b); 33I4 (7-22); 3956o(7-56b); 39606(1-I); 41615
List of Exsiccatae 389

(7-25); 44534(6-47); 44638(2-88a); 44656 Tejera, E., 79(2-27b); 82(7-36a); 1IO(7-22);

(2-15); 44667(7-56b); 45I47(7-56b); 45350 I69(7-36a); 22I(7-56b); 227(7-56b)
(2-15); 45466(7-56b); 45525(2-88a); 45743 Tessmann, G., 3115(7-56a); 3215(2-21); 3250
(7-22); 46132(2-15); 46356(7-37a); 46465 (2-21); 3446(2-43a); 3690(6-7); 383I (7-36a);
(2-15); 47761(2-45); 48073(2-15); 48074 4502(2-14); 5219(2-52); 5297(7-40); 5300
(6-47); 49303(2-15); 52144(6-47); 52145 (6-3b); 5370(2-60); 5382(6-35); 5468(2-21);
(6-47); 52829(2-4); 56917(2-106); 57647 sn(2-89b)
(7-25); 57825(7-8); 57904(7-21); 5931" Thieme, C., 522I(7-56b); 5464(2-88a)
(7-49); 59360(2-56); 59395(7-49); 6ooo8 Thorne, R. F., 14909(1-I)
(2-105); 60025(6-8); 60o58(7-36a); 60250 Thouin, A. par Aiton, W., sn(7-32)
(7-26); 60277(2-88a); 60465(2-105); 6056I Tidestrom, I., 6997(1-1)
(2-27); 60639(2-27); 60758(2-124); 60768 Tillett, S. S. et al, 43846(2-I35); 44912(2-56);
(2-105); 6o979(7-56b); 61093(2- I14); 61435 44969(7-27); 54044(7-74); 45048(7-45);
(2-27a); 6I759(2-43a); 62 73(2-I I4); 62183 45234(2-56); 45241(7-77); 45475(7-56a);
(7-36a); 62833(7-36a); 62841(2-13); 74639 45488(7-27); 45573(6-9); 45578(2-69a);
(7-56a); 74707(7-21); 74709(6-o0); 75428 45656(7-49); 45701 (7-27)
(2-69a); 75526(6-42); 75555(6-0o); 75562 Toledo, J. F. et al, SP 8850(2-44) ;SP 43174(3-I6);
(2-69a); 76039(7-74); 86268(7-25); 86272 SP 43190(2-41)
(i-I); 86273(1-1); 86370(2-77); 86374(3-4); Ton, A. S., 3759(2-45)
86380(7-56a); 86384(7-74); 86451(3-I); Tonduz, A., 3824(7-22); 3982(7-56b); 6708
86459(2-88a); 86468(3-4); 86480(7-56a); (7-56b); 7002(1-I); 7023(I-I); 94I0(7-36a);
86482(7-20); 86592(2-122); 86644(2-77); 10003(I-I); I0004(I-I); 10023(I-I); I0403
86684(2-109); 86758(2-77); 86828(4-3); (i-i); 13808(2-45); I3858(6-47); I3890
86843(4-I); 86864(4-3); 86865(2-43a); (2-45)
86972(3-3); 86976(2-77); 87127(2-109); Torrend, C, 311(7-18)
87209(7-56a); 87248(3-3); 87282(2-69a); Torrey, J., sn(2-I)
8730I (2-8); 87409(2-69a); 87429(2-8); 87451 Tracey, S. M., 6853(2-I); 7450(I-I); 745I(2-I);
(2-69a); 87493(3-1); 87506(2-39); 87554 9241 (-1)
(7-74); 87557(2-72); 87592(2-77); 87735 Traill, J. W. H., 2Io(2-69a)
(2-8); 87907(2-67); 87927(7-56a); 88030 Traverse, A., 487(I-I); 593(I-I)
(2-77); 88io8(2-69a); 88406(2-56a); 88410 Triana, J.J., 214(2-96); 4223(7-22); sn(I-I)
(2-56a); 88460(3-7); 88489(3-4); 88582 Tucker, 915(2-45)
(2-112); 89076(2-77); 89i 6(2-36a); 89716 Tuerckheim, H. von, II-1141(7-23); II-II52
(2-135); 89727(2-145); 90088(2-86); 90409 (7-56b; II-227();;
-127) I458(7-56b); 4087
(2-81); 908I6(6-3b); 90821(7-21); 90840 (7-56b); 7620(7-56b); 7659(7-36a); 7942
(2-93); 90865(2-106); 91027(2-86); 91361 (7-36a); 85I4(7-36a); 8680(7-23)
(2-86); 91377(2-69a); 92373(2-56); 92952 Tulleken, 483(2-69a) ;519(2-I34)
(2-56); 93019(2-124); 93035(6-42); 93070 Tutin, T. G., 20(6-45); 30(7-56a); 94(1-I); 230
(2-27a); 93176(7-21); 9449I(3-7); 95223 (2-89); 232(2-66); 289(6-I5); 29I(6-I5);
(2-86); 95334(2-69a); 95764(7-56); 95871 292(2-135); 293(2-89); 294(2-66); 6oi (7-5I);
(2-o16); 96017(7-22); 96100(7-72); 98191 626(2-I35)
(7-30); 99662(7-36a); 103618(6-48) Tweedie, J., I268(7-36b); sn(7-36b)
Stockdale, J. A., 8819(7-25) Tyson, E. L. et al, 1973(7-56b); 3713(7-22); 3859
Stocker, C. L., 9(7-56b) (2-88a); 4644(7-36a)
Stoffers, 2410(1-I); 4I69(7-36a)
Stork, H. E., 529(1-1); 3194(1-I)
Sucre, D., 268(7-54); 286(7-20); 852(3-16) Ule, E., 79(7-66); I57(7-70); I78(7-54); I79
Sunkill, 527(2-I); 568(2-1) (7-20); 459(2-49); 965(6-27); 2881(6-27);
Swabey et al, 31I39(2-69a) 2885(2-49); 3187(2-41); 4484(7-36b); 5547
(6-44); 5653(7-56a); 5654(7-40); 5996
(7-56a); 6II8(2-69b); 6149(6-29); 6150
Tackeuchi, D. M., INPA 7813(7-56b) (7-8); 6264(7-56b); 6453(7-56b); 6916(7-I);
Talbot, sn(2-I 4) 7096(6-26); 7274(7-70); 7306(7-20); 7585
Tamayo, F., I44(2-I06); 1421(2-I2); 3237(7-49); (7-25); 7641(6-43); 7885(6-43); 7886(2-26b);
3439(6-3b); 3586(7-25); 4308(7-49) 7981(7-2I); 8393(2-22); 8616t7-49); 8853
Tate, G. H. H., 146(7-21); 190(7-25); 287(7-8); (7-8); 8854(6-4I); 8913(2-69a); 8984(7-9);
372(7-21); 870(6-42) 9043(7-I8); 9412(7-56a); 94I3(7-36a); 9414
Tavares, S., I27(7-56b); 267(7-56b); 479(7-56b); (7-17); 9415(7-56b); 9446(2-21); 9598(2-45)
683(2-44); o00o(7-I8); II07(7-24); IIIO Underwood, L. M. et al, 251(1-1); 947(I-I);
(2-114); I130(7-24); 1208(2-114); 52-64 948(1-1)
(I-I); 53-335(I-I) University of Guyana, Bio io6, 3(2-135)
Taylor, N., 285(7-36a) Uribe, U. L., 2916(7-40); 359I(7-40)
Teixeira, 2573(2-114) Usteri, A., 3576(3-I6)
390 Flora Neotropica

Valerio, J., I16(6-47); 492(2-45); 891(7-22) Weberbauer, A., 4487(7-21); 4649(6-22)

Valeur, E. J., 395(7-36a); IoI9(7-36a) Webster, G. L. et al, 344I(2-I); 3547(2-I); 3812
Van Donselaar, J., I040(2-98); 1078(2-134); (I-I); 5341(1-I); 9264(2-85); 9296(7-32);
II31(6-18); 1134(2-52); 1151(2-97); 1152 9334(2-85); 9530(2-85); 9810(7-74); 9866
(7-56a); 1172(2-52); I20o(2-88a); 1242 (7-74)
(6-i8); 1247(6-18); 1467(2-97); 1493(2-52); Weddell, H. A., 2457(7-37); 2936(I-I)
1519(2-63); 1752(7-26); I870(7-42); 1897 Weigelt, sn(7-25)
(2-88a); I947(7-73); 2668(2-69a); 2786 Wessels, Boer, J. G., 1284(7-2); 1356(7-25)
(7-26); 2915(2-88a); 2930(7-42); 2946(2-43a); West et al, sn(2-I)
3035(7-73); 3036(2-88a); 3685(7-26); 3693 Westgate, 3157(1-I)
(7-5ob) Westra, L. Y. T., 48552(2-134); 48556(2-63)
Van Emden, W. C., 1333(6-2); sn(6-i); sn(7-25); Wetmore, A. et al, 105(7-56b); I98(7-36a); 213
sn(2-io8) (I-i)
Van Hermann, H. A., 561(7-22); sn(I-I) White, S. S., 5308(7-56b); 5378(I-I)
Van Sever6n, 67(6-47) Whitford, H. N. et al, 3(2-I ); 68(2-88a)
Vareschi 7998(2-1 19) Whitton, B. A., 8i (7-51); 98(6-7) 288(6-7);
Vauthier, 508(7-36b); sn(7-36b) Wickham, A., sn(6-22)
Veillon, J. P., i/v(7-36a) Widgren, J. F., 704(7-36b); sn(7-36b); sn(3-4);
Velasco, 8961(6-47) sn(2-i)
Velez, J., 1085(7-83); 108(7-83); 2259(7-56b); Wied-Neuwied, M. A. P. Prinz zu, sn(7-36b);
2265(7-56b); 2274(6-3b); 2275(2-69a); 2323 sn(7-36b)
(7-56b); 2543(2-69a); 2545(6-3b) Wiegand, K. M. et al, 146 (2- ); 1462 (2- )
Vellozo, H. P., 200(7-85); 736(2-138); io65 Wight, A. E., 6i(I-i); 174(1-I)
(2-I ) ;sn,R38167(2-I32); sn,R 38619(2-132) Wilbur, R. L. et al, 7989(1-I); 8174(1-I)
Veloso, H., I201(2-4I); 1330(6-27); 14I (6-20) Williams, LI. et al, 7(6-44); 29(6-44); 63(6-44);
Versteeg, G. M., 172(6-2); 389(2-27a); 422 88(6-44); 50o(7-36a); 804(7-53); 888(6-22);
(2-27a); 728(7-56a); 751(6-2) 917(6-44); 1113(3-I4); II 5(6-44); 1140
Viana, 0. de C., 2(2-25a) (3-14); I183(7-54); I33I(2-43b); I490(6-3b);
Vice-Consul, Ceara, sn(2-44) 2346(6-44); 2358(7-56a); 2765(7-56a); 2975
Victorin et al, 51(7-22) (6-35); 2990(7-56a); 3780(6-7); 3829(7-56b);
Vidal, J., sn,R 35180(7-18); sn,R 36623(7-18) 3884(7-56b); 4041(7-56b); 4653(7-36a);
Viereck, I 97(6-47) 5012(6-36); 5064(2-2I); 5887(7-56b); 6154
Virgilis, V., sn(2-44) (7-56b); 7832(7-56b); 8157(6-44); 8401
Vit6rio, RB 48468(2-127); RB 136919(7-33); (7-56b); 8650(2-15); 8722(2-15); 9340(7-56b
RB I36938(2-43a); sn(7-64) 9426(7-56b); 9480(7-36c); 9487(7-56b);
Vocke, 6086(7-25) 9568(7-56b); 9586(7-56b); 9972(3-7); 9995
Vogel, C., 49(7-36a); 1437(7-36a) (7-36a); 10333(2-145); 10753(7-36a); 11203
Vogelmann, H. W. et al, I305(7-53) (7-56b); 11306(2-75); II556(7-56a); 11626
Von Hagen, V. W., I353(7-56b); 1378(7-36c) (7-16); 11746(7-56b); 11884(7-25); 11909
Von Sneidern, A., 1190(7-56c); 4905(7-10); (i-i); 1198o(6-3b); 12015(7-20); 12037
49I4( 7-I) (6-3b); 12063(2-129); I23I9(7-36a); 12554
(7-56); I2603(2-27a); I2653(7-56b); 12678
Wachenheim, H., 11(2-130); I2(3-I); 23(2-108); (2-27b); I2679(2-27b); 12694(2-12); 12705
43(2-99); 6o(7-56a); 74(2-134); 111(3-I); (7-56b); I27I8(7-56b); 12917(2-12); 13024
II3(2-97); I24(2-63); I29(3-I); I37(2-97); (2-69b); 13278(7-56b);; i3305(6-3b); 13848
138(2-134); I4I(2-74); I43(7-25); 180(2- (2-103); I3849(2-27a); I3849(2-I03); 13987
io8); i83(7-5ob); 185(2-63); I98(2-8); 208 (2-68); 14048(3-I); 14077(2-103); I4I19
(2-69a); 213(7-25); 217(2-134); 235(2-69a); (2-103); 14I8I(6-9); 14292(2-9I); I4356
252(2-69a); 274(6-18); 317(2-135); 349 (6-5); I4437(7-71); 14464(7-40); I4616
(2-63); 372(7-20); 401(2-99); 2-274(6-18); (7-21); 14640(6-2); I4707(2-69a); I4755
3-I (6-18); sn(2-8); sn(2-69a); sn(2-Io8); (2-129); 14913(2-I03); 14940(7-72); 14961
sn(2-I35); sn(3-3) (2-24); I5004(7-56b); 15018(7-56b); 15055
Wadsworth et al, 523(1-2); 575(1-2); 588(1-2) (6-9); I5077(2-27a); I5227(2-69a); 15254
Wagner, R.J., 231(7-83); 325(7-83); 549(7-36a); (2-140); 15602(2-69b); 15645(6-2); 15680
58o(7-36a) (2-75); 15772(2-12); I5937(6-3b); 15939
Walsh, J. J., sn(i-i); sn(i-i) (2-69a); i597o(6-3b); I7968(7-36a); 18232
Ward, 8646(7-74) (2-97); I8246(7-5oa); 24215(7-56b); 24523
Warming, J. E. B., I88(I-I); sn(7-2o); sn(7-53); (7-56b)
sn(2-4I); sn(6-27) Williams, L. O. et al, 7597(6-27); I2210(2-45);
Watson, S., 27(1-I); 294(7-36a); 437a(7-56b); 12563(2-45); 12830(2-45); I4465(1-I); 15224
sn(7-36a) (2-15); I5951(2-24); 18005(2-45); i8i6o
Weber, 62 8(1-1) (2-45); 242I5(7-56b); 24523(7-56b); 28757
Weber, G. F., sn(2-I) (7-56b)
List of Exsiccatae391

Williams, R. O., 1I38(2-69a); 11897(7-56a); (2-128); 306(2-39); 339(2-77); 352(3-I);

12052(2-I2); 12432(3-I); 12450(2-12); I2493 353(I-I); 1986(7-21); 2524(2-27a); 34378
(i-i); 12494(7-25); I2495(I-I); 12605 (7-36) (7-25); 39414(3-I); 3943I(7-56a); 39507
Williams, R. S., 91(7-56b); 247(7-22); 338(2-45); (7-56a); 39535(I-I); 39569(2-109); 39574
400(1-I); 667(7-36a); Iooo(7-36a); 1568 (7-56a); 39657(2-69a); 39675(3-3); 39729
(7-46) (7-i6); 39754(7-56b); 39893(6-3b); 39896
Wilson, 5566(6-34) (2-69b); 39974(7-56b); 40864(6-3b); 40931
Wilson, C. L., I4(I-I); 67(7-36a); 365(7-56b); (6-2); 40933(2-69a); 40935(7-40); 40966
39I(I-I) (2-140); 40971(2-27a); 41028(6-2); 41039
Wilson, N., sn(7-36a) (2-27a); 41040(2-83); 41099(2-33); 41124
Wilson, P., 211(7-56b); 481(7-22) (7-56b); 41138(2-27a); 41143(3-7); 41175
Wilson-Browne, G., I74(7-36a); 203(7-56a); (2-83); 41254(2-27b); 41350(2-24); 4I363
224(2-Io8); 262(7-25); 306(2-o08); 307 (2-69a); 42731(2-69b); 42735(2-27a); 42769
(7-56b); 363(7-36a); 380(2- 13); 467(2-135); (2-69a); 42885(3-I); 42967(2-119); 42968
480(2-122); 482(2-52); 484(2-122); 525 (6-3b); 43231(7-21); 43242(2-88a); 43246
(3-3); 595(1-I); sn, F.D. 6491(2-134) (7-8); 43266(3-6); 433 1(7-55); 43573
Winsor, sn(2-I) (2-69a); 436I2(2-88a); 43621(7-12); 43693
Winzerling, II-I0(7-22); II-21(7-56b) (7-49); 437i6(7-29)
Woodson, R. E. et al, 80(2-I); Ioo8(7-36a)
Woodworth, R. H. et al, 332(7-56b); 370(7-36a);
647(7-36a); 728(7-22) Xaiz, 283(7-56b)
Woytkowski, F., 5059(6-36); 5405(7-56b); 5625
(7-47); 5782(7-56b); 5787(7-21); 7256
(7-56b); 34432(7-56b) Young, J. P., 638(1-1)
Wright, C. et al, I55(7-36a); I78(7-36a); 194 Yuncker, T. G. et al, 4673(1-1); 876o(7-36a)
(I-I); I607(1-I); 2409(7-22)
Wulff, C. W., sn(2-I2)
Wullschlaegel, H. R., I64(I-I); 778(7-25); 807 Zabala, A., 1(2-65); 26(2-77); 3I(2-65); 33
(7-56a); 8I7(7-56a); 836(1-I); 964(2-135); (2-1 3); 35(2-II3); 42(2-65); 63(3-3);
965(7-56a); 1372(3-I); I44I(2-I35); I443 65(2-77); 69(2-77);
(7-56a); I444(7-56a); 1650(2-63); I651 Zehntner, 23(6-20); 401(7-54); 632(7-20); 1917
(2-63); I776(7-56b); 2058(7-25); sn(2-95) (6-20); 3001(6-20)
Wurdack, J. J. et al, 33(7-56b); BI64(7-I8); Zerny, H., sn(7-56b)
BI89(I-I); B2I5(7-56b); 2I5(2-27b); 216 Zetek, J., 435I (2-I5)

Without col., sn(7-36b)

 INDEX OF LOCAL NAMES

Aceituna330 Cajuru'19
Achotilla 291 Caligni 163
Aciri-yu-yek249 Camaroncillo304
Aiamoradani86, 220 Canadulce 87
Ajurarana330 Candelita345
Ajuru-branco19 Canellade velho 338
Ajurui-rana295 Canillomula 87
Akianteifi-koko344 Canoe285
Akiantiici 328 Can-wakak-yek295
Akoerieerepare163 Caramacate negro 304
Alauna o00 Caramio 19
Alcornque87 Carapato304
Amaree67 CarboneraI45, 304
Ambure400 Carbonero278, 328
Anauera o00 Caripe 66, 68, 76, 79, 84, 27, I34, 57, I62
Anaura 92, Ioo, 104, io8, 229 Cariperana68, 69, io8, 144
Anaura,hoogland229 Caritade negro 290
Anauraballi229 Caritanegro 304
Apacharana62 Castanha99
Apesia 209 Castanhade cotia 359
AraudanniII8 Castanhade galinha 248
Aruadan227 Castanheiradepedra 182
Ascindiuva145 Cenizanegra 284, 328
Aseepee244 Cerricillo330
Babooncap 253 Chicharron304
Bafo de boi 145 Chorao285
Barazon 307 Chozo 127
Belero 124 Chuchuhuasha-masha 346
BoehoeradaI17, 182 Chupa 55
Boehoeroedoe 182 Cinzeiro 354
Bohowoda182 Clavellino84
Bois baude182 Cocdo84
Bois diable 123 Cocoaplum 19
Bois gaulette 132, I33, 148, I52, 293 CococowIo8
Boisfer 123 Cocode cotia 359
Bois gris 123 Coffeewood io8
Bois de masse 123 Conejo304
Bois poil 304 Copuda92
Bois resolu 123 Coquito355
Bokobokokandiamaroe141 Corafaode negro 330
Bokobokotokon 276, 285, 289, 320, 328, 345, 348 Corodepernambuco254
BokohotonI86 Cumate214
Boliquin 210, 229 Cuparana221
Bongro I58 Cuyuy 210
BoroborelliI04 Disbota 112
Buhurada 182 Eehawdla 309
Buhurada,hill 191 Encina 87
Bukotokon287, 304 Erejooroe19
Buku-buku293 Escobilla 330
Buruburuli104, 105, Io8 Farinhaseca I84
Cacahoanantzin87 Fat pork 9
Cacahuananche 87 Foengoe I34, 141, 148, 152, 158, I82
Cacahuate87 Foengoe,braka 152
Cacco 19 Foengoe,bruinbladige229

392
Index of local Names 393

Foengoe,man 152 Kauta balli balli I09, 112

Foengoe,sabana 162, 163 Kleinbladigeanaura 328, 344
Foengoe,witte 134 Kobo 210
Foungouti 182, I86 Kodibi-assiballi19
Frailecillo 87 Koelimiro 9
Frutade ema 195 Koenatepie19
Fruta depaloma 304 Koenoto-epoe19
Fulishakuyefu272 Koepesienoe182
Garrapato 290, 309 Koko-baaka141
Gaulette92 Koko-lebi 133
Gaulettegris-gris 145 Komkow o08
Goajuru 9 Koo-yo 2I0
Goiabinha320 Kouebi 210
Gopherapple 43 Krikimauroeroe108
Gorogoro 158 Kuepi 157
Grenada330 Kunoko I 8
Gris-gris 92, 133, 148, 328 Kwefe I35
Gris-griscumateI00o Kwepi 68, 78, 84
Grootblad158 Kwepi, sabane 249
Groundoak 43 Kwepierang143
Guaray I 7, 182, 284 Kwepilan 163
Guarayblanco 84 Landhout143
Guaraynegro 199 Lebi koko 132
Guari I86, I89 Macara-yek 139, i86
Guaviluna290 Macucu 108, 214, 221
Guayabito138 Macucubobo 66
Guayabitode tinta 253 Macucuchiador99
GuayaboIo8 Macuc farinha seca 152
Hatcheballi220 Macucuz
ffo Io8
Hierrillo 127, 144 Macucusangue 120
Hierrilloblanco 127 Mahaicaballi 182
Hierrillonegro 144 Makara 221, 238
Hierrito 117, 139, I42, 143, 157, I62, 304 Makara-yek152
Hierro 142 Mamoncillo68
High ridgecocoaplum 291 Manja croite 19
Hija 345 Mapuritomontaiero254
Huaje 57 Mari bravo 255
Huipanupare157 Marirana 227, 228
Icacillo 19, 293, 304, 352 Marishiballi 92, 117, I2I, 128, I30, I52, 158, 163
Icaco 19, 293 Marishiballihariratoe134
Icacode aura 290 Marishiballitataro 14I
Icacode montana291 Marishiballithick-skinned128
Icacomacho291 Marmelitodo campo80
Icacomontes330 Merecure52, 184
Icacopeludo 290 Merecurede montanaI89
Icaquemontagne21, 298 Merecurillo 117, I82, 184, i86
Icaqued poils 304 Merecurilloblanco 79
Icaquepoileux 304 Mamoncillorebalsero69
Icaquito 139 Mesonsapote55
Icatillo 290 Milho cozido 60o, 170, 259
Ingibarki o8, I99 Milho torrado 171
Iron mary 127 Miwari-ek 157
Isiguiro-ey304 Monkeyapple 55
Itoeloetanojapopalli 104 Monkeycap 253
Japopare o8 Mooneka 164
Jicacillo 328, 330 Moshpin 55
Jicaco 307 Munzap 253
Juan-poop I67 Muri kautaballi 118
Kairiballi o8, 109 Murta 330
Kairiballihohorodihoro 209 Murtinha 285, 330
Kanhe 210 Nahaun 238
Kauta 68, 75, 84, 141, 143 Naranjillo 345
Kautaballi 141, I42 Oajurzi19
394 Flora Neotropica

Oenikiakia163 Serrecio330
Oenikiakiadjamaro127, 209 Siguapa 304
Oiti 47, 51, II2, 214, 231 Soberbio124
Oiti boi 235 Sorore84
Oiti coranga242 Sponshoedoe00
Oiti coro 47, 254 Sponshout 00
Oiti da praia 51 Suichi-parinari251
Oiti silvestre90 Sunza 55
Oiticica 86 Sunzapote55
Oitizeiro 47 Swampkauta 48
Olozapo 253 Tacamahaco69
Onitjatja 162, I63 Tajamama 182
Oroy-uara-yeko8 Teta de burracimarron352
Pajurd213, 225, 227, 255 Tifeuilles 21
Pajurdde mata 227 Tijantesie345
Pajurddepedra 182 Totopostle87
Pajurddo igapo 246 Tucriba 238
Pajurarana92, 99, 133 Uchi de cotia 79
Pajurazinho170 Uchi de tambagui i86
Palo blanco 304 Uchirana 186, 213
Pasa 157, 290 Uilho torrado145
Palo defraile 87, 253 Uiti 231
Palo de hierro 84, 171 Ulazapote 253
Parinari I84, 245 Umarirana213, 228
Pasta 330 Unikiakia 118, 121, 127, I30, 145, 150, 163, 249
Pau de lixa 354 Upachaumari 193
Pau pombo 285 Urraco55
Pendoreille 298 Urura 289
Pereguetano189 Uspio 253
Perone229 Varillacaspi 330
Perz I15 Vermelhdo287
Pianchirana92 Vonkhout141, 182
Pica-pica 276 Wadd 297
Pica-picarosada 276 Waroma304
Pigeonberry304 Wayamche330
Pigeonplum 290 Wee-roo210
Pigeonplum, bastard73 Wetikoko 148, I58
Pigeonplum, wild 330 Wild cocoa I o8
Pilon 213 Wild cocoaplum 290
Pil6n nazareno 103 Wokerebesoerie186
Pintadinha Ioo, 124, 152 ra-fee-a 67
Pio 253 Ya-ya 290
Piquapiquamorada328 raku-shimbillu323
Pitombade leite 237 rucuare 315
Proewaipoeroe328 Zandhout148
Pruim 9 Zapote 55
Puccavarilla 330 Zapoteamarillo 55
Querebere213 Zapotebolo 253
Quirindolcacahuananche 87 Zapoteborracha55
Ran-hoo67 capote cabello 55
Roble 87 Zapotillo 253
Rosenkwarie186 Zapotilloamarillo 253
Sabaneanaura104 Zicaque 19
Sangre 55 Zicaquedes bois 21
Santihoedoe152 Zinzapote55
Sapotede nino 253 Zwartefoengoe92
Sergeants-kloot00
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 CHRYSOBALANACEAE-SUPPLEMENT

Since completing work on this monograph I have now received material of 2 new
species and have examined more exsiccatae of other species. For completion this extra
information is presented here rather than in a separate publication; and footnotes
have been added to the keys in the appropriate places.

Licania Aublet

Iooa. Licania naviculistipula Prance, sp nov Fig 59 G-K.

Arbor circa I m alta, ramulisjuvenilibus tomentosismox glabris. Folia alternata

petiolata; petiolo 8.o-I2.0 mm longo, tomentoso mox glabro, tereti, prope laminam
cum glandulis duobus munito; laminae coriaceae, oblongo-ellipticae, io.o-i8.o mm
longae, 5.5-9.0 cm latae, basi rotundatae vel subcordatae, apice in acuminem 2.0-
Io.o mm longum contractae, supra glabrae, subtus cum cavis stomatalis faucis dense
lanato-pubescentibus, venulis glabris conspicuis; costa media supra plana, glabra;
costis secundariis 12-16 jugis, supra planis, subtus prominentibus glabrescentibus.
Stipulae ad I3.0 mm longum, triangulares, naviculares persistentes, coriaceae,
tomentosae, cum basi petiolo adnatae. Flores 6.0-7.5 mm longi, sessiles, in paniculis
racemosis terminalibus subterminalibusque dispositi, rachi ramisque ferrugineo-
tomentosis. Bracteae ad Io.o mm longum, tomentosae, persistentes saepe cum folio
vestigiali adnatae, bracteolae 1.0-3.0 mm longae, oblongae, tomentosae, persistentes.
Receptaculum anguste campanulatum, pro genere magnum, extus brunneo-
tomentosum, intus tomentosum. Calycis-lobi acuti, utrinque tomentosi. Petala nulla.
Stamina 8, unilateralia, staminodiis dentatis opposita; filamenta calycis-lobos
breviora, glabra. Ovarium ad basim receptaculi insertum, pilosum. Stylus e basi
ovarii ortus, calycis-lobos aequans, lanato-pubescens. Fructus mihi ignoti.
TYPE. R.P. Belem3814, Brazil, Espirito Santo, Linhares, Vale do Rio Doce, fl
(holotype, NY; isotypes, FHO, UB).

This species is known only from the type gathering. It belongs to section Licania,
but is most distinct within the genus on account of its large flowers, large navicular
stipules, and the leaves with extremely well-developed stomatal cavities. The only
other species in section Licaniawith comparable large flowers is L. couepifolia,which
differs in many other characters. The large navicular stipules distinguish it from all
other species of Licania.The inflorescence is interesting in that the lower branches
have much larger bracteolessubtended by a residualleaf, showing that these bracteoles
are morphologically equal to the stipules which they resemble in shape.
keys out at lead I 2 on account
In the key to species of Licania,L. naviculistipula
of its large flowers. Lead I112 may be amplified as follows:
I I2. Flowers 6.o-7.5 mm long; stamens often connate for half their length.
I I2a. Stamens connate for half their length; leaves prominently reticulate but
without stomatal cavities; petioles eglandular. L. couepifolia.
II2a. Stamens free to base; leaves with conspicuous stomatal cavities; peti-
oles with 2 glands near to lamina base. L. naviculistipula.

398
Parinari 399

o D

~~ //

7ci.

J K t 0 0 Z

FIG 59. Speciesof Parinariand Licania.A-F, Parinariromeroi (Romero C. 5348,5570). A, habit

x i; B, leaf undersurfacex 2; C, flower x 5; D, flower section x 5; E, fruit x i; F, fruit section
(Bel6m38I4). G, habit x 1; H, leafundersurfacex 2; J, flower x 5;
x j.G-K, Licanianaviculistipula
K, flower
K, section x 5.
flower section
400 Flora Neotropica

Parinari Aublet.

13a. Parinari romeroi Prance, sp nov Fig 59 A-F.

Arbor ad 20.0 m alta, ramulisjuvenilibus hispidulis mox glabris et prominente

lenticellatis. Folia oblonga coriaceae, 7.0-I5.5 cm longa, 4.0-9.5 cm lata, basi
subcordata, apice in acuminem obtusum I.0-3.0 mm longe contractae, supra glabra,
subtus cum cavis stomatalisgriseo-lanatopubescentia plenis; costa media supra plana;
costis secundariis 19-22 jugis, subtus prominentibus, supra planis; petiolo 3.0-4.0 mm
longo, juvenile tomentello, tereti vel leviter canaliculato, eglanduloso. Stipulae parvae,
ovatae, subpersistentes. Flores in paniculis brevis pauci-ramosis ad 4.0 cm longis;
rachi ramulisque brunneo-tomentosis. Bracteae bracteolaeque 3.0-5.0 cm longae,
persistentes, ovatae, extus tomentosae. Receptaculum campanulato-turbinatum
extus brunneo-tomentellum, sessile. Petala 5, calycis-lobis aequantes. Stamina fertilia
6-7, fere 7 staminodiis brevibus opposita. Ovarium basisque styli pilosus. Drupa
oblongo-ovoidea, circa 5.0 cm longa; endocarpio lenticellato; mesocarpio tenue,
carnoso; endocarpio duro, 5.0-I2.0 mm crasso, osseo-fibroso, extus tuberculato-
fibroso, intus lanato.
TYPE.RomeroC. 5570, Colombia, Narinio,Tumaco, Brazo del Mira, Guandal, fl
(holotype, COL).
DISTRIBUTION. Known only from the coastal delta region of Narifio, Colombia.
Collected in flower in February-March and in fruit in October.
COLOMBIA. Nariiio: Tumaco, Vicinity of Pifial Dulce, RomeroC. 5348 fr (COL).

LOCAL NAME. Ambure.

Parinariromeroiis nearest to P. cardiophylla,but differs in the oblong leaves, the

blunter apices, the shorter petioles and the much reduced inflorescence. It is quite
distinct from the other Pacific coastal species P. chocoensis, differing in the reduced
brown-pubescentinflorescence, the thicker broader leaves with a much blunter apex,
the subcordate leaf bases, the shorter petioles etc.
To
In the species key to Parinari,P. romeroikeys out at lead 8 to P. cardiophylla.
this lead may be added:
8. Leaf base distinctly cordate or subcordate.
8a. Leaves ovate, the acumen 10.0-I2.0 mm long; petioles 6.0-I2.0 mm long, glandular;
inflorescences much-branched, 5.0-6.0 cm long. 13. P. cardiophylla.
8a. Leaves oblong, the acumen I.0-3.0 mm long; petioles 3.0-4.0 mm long, egland-
ular; inflorescences little-branched, 2.0-4.0 cm long. I3a. P. romeroi.
8. Leaf base rounded to cuneate.
 SUPPLEMENTARY LIST OF EXSICCATAE

Adams, C. D., 7158(1-1) Hitchcock, A. S., 1097(2-I)

Argent, G. C. G., et al, 6388(2-30); 6390(7-20); Hunt, D. R., 5506(3-I6); 5866(6-3b); 6657(2-49)
6397(7-54); 6423(7-54); 6497(7-54); 6484 Im Thurn, E. F., sn(7-51)
(6-27); 6608(7-20); 6624(7-54); 666i (2-30); Irwin, H. S., et al, 22882(7-85); 23424(4-4)
R 6662(2-114); 6665(2-141); 6666(7-19); Jativa, C., et al, 1152(7-10); 2046(7-10)
R 6698(2-141); 6737(2-30); R 6806(7-19); Kappler, A., 82(6-2); I 8(7-56a)
R 6847(6-27) Kelly, I., 339(6-47)
Bel6m, R. P., 38I4(2-Iooa) Kimber, C., 1399(1-I)
Bernardi, A. L., 757(6-10); 889(7-56a); 904 Kramer, W. P., 62(7-36a)
(6-io); I624(3-4); 2127(3-3); 214I (2-69a); Labroy, sn(7-6)
2777(2-18); 2828(2-81); 2839(2-81); 6629 Lakela, O., 28679(1-I); 29623(2-I)
(2-89); 6637 (2-69a); 6676(2-89); 6682(3-4) Laughlin, R. M., 1705(2-15)
Blanco, C., 421(2-109); 664(2-276) Lewis, W. H., 7117(1-I)
Boer,J. de, 13(1-1) Lightthipe, L. H., 845(2-')
Brenes, A. M., 4188(7-56b) Linden, J., 1735(7-36)
Broadway, W. E., 2823(7-56a); 2838(7-36) Luetzelburg, P. H. von, 218(7-56b); 494(7-54);
Burchell, W. J. 5935(7-54) I825(7-54); I5597(7-20); 20201(2-97);
Byrne, R., 466(1-1) 20314(7-56a); 20552(7-56a); 21374(7-21);
Castro, R. de, 0o660(2-114) 21585(7-25); 26232(7-56b); 26411(7-I8)
Contreras, E., 5661(7-22) Maguire, B., et al, 56022(6-4); 56024(7-56a);
Cowlos, T. H., 2467(7-83) 56026(2-27a); 56035(6-50); 56726(6-3a);
Croat, T. B., 4763(7-22); 6o48(7-36a); 7302 56747(7-56a); 56764(2-27a); 56798(2-82);
(7-566);775I(7-22);8632(7-22);8695(2-I5) 56823(2-17); 56825(6-27); 56948(6-27);
Crosby, et al, 45(7-36a) 57046(2-27a); 57065(7-54); 571 I6(2-27a)
Crudy, sn(i-i); sn(7-36) Martius, K. F. P. von, 1840(3-16); 2695(1-I);
Cuming, H., I I 4(7-56b) 2922(2-27a); 2952(6-16); 3072(2-27a); 3246
Duke, J. A., et al, I11569(I-I) ; 11822(7-22) (7-2); 3278(6-9); sn(i-I); sn(2-44) ;sn(3-4);
Eggers, H. von, II I3(2-69a); 10156(7-36) sn(7-2o); sn(7-54); sn(7-56a); sn(7-63);
Evans, L., 21(7-83) sn(7-7o); sn(7-73)
Ferreira, A. R., sn(7-4o) McKee, H. S., Io72I(7-56a)
Focke, 1 (6-42); 206(7-25); 343(1-1); 386(7-25) Meebold, A., 26965(1-I); 27381(2-I); 27503
Forest Dept., British Guiana (Guyana), 6272 (2-I); 276I8(2-I); 27962(I-I)
(2-90); 6381 (2-90); 6382(2-90); 6385(3-4) Melinon, M., I78(7-50b)
Frazer, sn(i-i) Molina, R. A., 8368(7-22)
Fr6es, R. L., et al, 28539(2-43a); 29o88(2-27a); Morton, C. V., 4778(1-I); 5293(2-85); 5294
29277(6-29); 29527(2-27b); 29564(2-129); (2-85); 5759(2-85)
30387(2-37); 30552(6-3a); 31327(2-27a); N. R. L., 78A(2-45)
3I345(2-69a); 31622(2-97); 32301(2-28); Oldeman, et al, 73(2-124); 202(2-124); 2134
32336(6-3a); 32353(6-3b); 32882(2-57); (2-86); 2463(2-67); 2464(2-69a); 2562
32942(2-69a); 33743(2-27a); 34108(2-28); (7-56a); 2570(2-69b); 2607(2-88); 2665
34580(2-69a); 34685(4-3); 34877(6-3b) (7-56a); 27i6(7-56a); B 584(7-56a); B 757
Gaudichaud, C., sn(7-8i) (2-69a); B 773(2-95); B 804(1-I); B 831
Glaziou, A. F. M., 10699(7-18) (7-25); B 854(2-69a); B 903(2-134);
Gonzalez Q.I., 3505(7-36a) B 937(2-95); B 938(2-8); B 969(2-8); B 987
Guillemin 6(7-85) (2-69); B I111(2-39); B 1014(7-25); B I85
Harley, R. M., et al, 10094(7-54); 0o200(2-27); (2-64); B 119I(7-25); B I200(2-97); B 1208
10283(7-20); 10317(6-27); 10517(6-27); (2-69a); B I226(2-134); B 1245(2-88);
10525(2-27); 10731(2-29); Io83I(6-3b); B I366(2-I34); B I467(2-28); B 1468(2-26);
II156(6-3b); 1i 88(2-28) B 1528(7-25); B 1530(2-8); B 1616(2-69);
Hatschbach, G., 7463(7-85) T I20(7-52); T 259(7-74)
Haufe, et al, I6(7-36c) Oliveira, E., 668(2-69a)
Heringer, E. P., Io650(2-27a) Omawale, et al, 77(1-I)
Heyder, H. M., 17(7-22) Perera, A., 2(2-15)
Hilerio H., L., 32(7-56b) Petitbon,J., 69(7-25); I49(3-I)

401
402 Flora Neotropica

Philcox, D., et al, 3052(2-141); 3229(2-17); Silva, M., et al, 2345(7-56a); 2349(7-50b); 2471
3255(2-II4); 3655(2-27b); 4230(7-54); (7-56a); 2514(7-56b); 2521(6-4); 2530
4785(2-80) (8-78); 2548(7-2); 2554(2-14I); 2566
Pinto E., P. et al, 828(7-40) (2-69a); 2580(6-2); 2621(2-79); 2627
Pires,J. M., 3186(2-63); 3187(2-63); 3188(2-63); (7-5oa); 2632(2--69a)
3195(3-3); 6859(2-69a); 6981(2-81); 7006 Silverstone, P., 124(1-1)
(6-50); 7038(2-81); 7Io6(2-69a); 7113 Small, J. K., et al, 6340(2-I)
(2-124); 7124(3-3) Souza, R., R 589(2-37)
Pl1e, A., 445(7-83); 446(7-36a) Splitgerber, F. L., 605(7-25); 873(2-135); 915
Pohl,J. E., 123(7-36b); sn(6-25); sn(7-54) (7-56a)
Ramos,J. F., et al, 26(2-29); 34(2-30); 90(7-20); Spruce, R., 298a(2-63); 299(6-2); 300(2-43a);
151(2-141); I66(2-30); 176(7-20) 302(7-56b); 303(7-56a); 304(7-18); 305
Ratter, J. A., et al, 28(7-20); 36(7-54); 51 (2-41); (7-18); 306a(2-27a); 307(2-57); 379(7-I8);
115(2-30); I21(2-41); 33(7-54); 34(7-20); 799(2-43a); 800(2-37); 801(2-37); 829
136(2-30); 191(2-41); 2o (7-54);275(7-20); (6-36); 1175(6-37); 176(6-I I); 1177(6-I6);
441(2-141); 445(7-20); 449(7-54); 455 I180(7-2); I472(2-69a)
(2-30); 482(6-20); 510(7-20); 618(7-66); Stahel, G., 54(6-I); 322(2-135)
663(6-27); 1209(7-73); 13I6(7-54); 1322 Stevenson, N. S., sn(7-23)
(2-30); 1386(2-114); I516(2-41); i6o6 Steyermark, J. A., et al, 86779(2-18); 86826
(7-56a); I947(2-41); 20I2(2-17); 2023 (2-27); 101398(2-114); 101518(7-36);
(7-54); 2029(2-30); 2065(7-19); 2068(7-54); 101578(2-57); IO1639(2-II4); I01721
2142(6-27) (2-57); II01772(2-1 4); I01788(6-44);
Rensch, 4(2-27a) 101826(3-7); 102011(7-22); 103618(6-48)
Richard, L. C., sn(7-83) Suriname For. Dept., 1400(2-63); 1403(3-I);
Richards, P. W., et al, 499(2-37); 6498(6-3b); 1404(3-I); 1409(2-63); 1410(2-63)
6521(7-54); 6528(2-141) Thiebaud, 41(7-56b)
Romero C., R., 5348(3-I3a); 557o(3-I3a) Thomas, J., et al, 1536(7-22)
Sandeman, C., 2148(7-54) Traill, J. W. H., 213(7-24)
Santos, R. R. de, et al, 1491(7-73); I555(7-20); Tulleken,J. E., 363(7-25); 445(7-73)
I666(7-5ob); 1796 (2-141) Vogel, C., 65(1-1); 417(1-I); 1250(1-I); I438
Sargent, F. H., 8261(2-1); sn(2-I) (7-36a)
Sastre, C. & F., et al, 290(2-135); 291(1-1); Wachenheim, H., 239(7-50a)
641(3-14) Wadsworth, R. K., 199(1-I)
Sauer, J. D., et al, 2177(1-1); 2345(I-I); 2479 Wagner, M., sn(7-56b)
(I-I); 3009(1-1); 3299(1-I); 3673(1-I); Wagner, R. J., 568(7-83)
4142(1-1); 4268(1-1) Ward, D. B. & S. S., 1571(2-I)
Schlim, L., 1144(7-22) Weigelt, sn(I-I)
Schmidt, E., 5(7-54); 26(3-5) Whitford, H. N., et al, 66(2-45)
Shank, P. J., 78(7-22) Williams, LI., I5972a(2-I2)
Sidney, 350(2-29) Woytkowski, F., I400(7-36a)
Sieber, F. W., 96(1-I); 305(7-36); 31 (1-') Wullschlagel, H. R., 1294(I-I)
Zehntner, 4048(7-20)
 INDEX OF SCIENTIFIC NAMES

Acia 202, 357 Couepia (cont.)

amara 208 calophlebia 237
dulcis 359 canomensis 220
Acioa 357-36I caryophylloides 229
amara 208 cataractae 231
dulcis 359 chrysocalyx 245
guianensis 359 cognata 248
schultesii 361 var cognata 249
somnolens 359 var major 249
Angelesia 22 var membranacea 249
splendens 172 comosa 251
Balantium 178 dahlgrenii 231
cordifolium 18I divaricata 213
Batheogyne (section) 23 var strictiuscula 213
Byra 259 dodecandra 252
Causea 259 duckei 212
Chrysobalaneae 14 elata 237
Chrysobalanus 14-2I eriantha 245
americanus 21, 289 excelsa 230
cuspidatus 20 exflexa 225
ellipticus 6 floccosa 252
icaco i8 formosana 236
var ellipticus 6 foveolata 221
var genuinus 6 froesii 258
var pellocarpus I6 glandulosa 209
incanus Rafin. 21, 42, glaucescens 212
incanus (Aubl.) G6mez de la Maza 2I, I62 glazioviana 254
humilis 21, 79 grandiflora 236
macrophyllus 21, 235 guianensis 208
montanus 21, I82 habrantha 225
oblongifolius 21, 42 hypoleuca 234
orbicularis 16 impressa 255
ovalifolius 21, 234 insignis 242
pallidus 2I, 42 krukovii 233
pellocarpus I5 kunthiana 252
prunifolius 21, 43 latifolia 233
purpureus 15 laurifolia 259
retusus 21, 42 leptostachya 213
rugosus 21, 352 longipendula 247
savannarum 16 macrophylla 232
splendens 172 magnoliifolia 224
sublanatus 21, 79 maguirei 214
subundulatus 21 martiana 231
triandrus 21, 302 martinii 238
Coccomelia 22 meridionalis 256
nitida 172 multiflora 250
Cosmibuena 259 myrtifolia 209
Couepia 202-259 obovata 244
amazonica 227 ovalifolia 234
belemii 228 panamensis 202, 259
bondarii 240 paraensis 210
bracteosa 227 subsp cerradoana 213
var grandifolia 227 subsp glaucescens 212
var minor 227 subsp paraensis 212

403
404 Flora Neotropica

Couepia (cont.) Griffonia 357

paraguariensis 231 Grymania 201, 202
parillo 218 polyandra 252
parvifolia 258 Hedycrea 2
pauciflora 218 incana 162
pernambucensis 256 Hirsuta (section) 28, 92-96
platycalyx 253 Hirtella 259-357
polyandra 252 acayacensis 328
racemosa 237 acuminata 315
var reticulata 238
adderleyi 296
recurva 242
reflexa 247 adenophora 346
rivalis 212 aggregata 292
americana Aubl. 325
robusta 255
var gracilipes 323
rufa 254
var hexandra 328
sandwithii 216
schottii 235 var oblongifolia 328
americana Jacq. 302
speciosa 232 americana L. 289
spicata 226
steudeliana 248 amplexicaulis 288
steyermarkii 220 angustifolia 351
stipularis 246 angustissima 320
subcordata 227 apetala 66, 357
suberosa 236 araguariensis 278
surinamensis 208 aramangensis 315
thyrsiflora 209 aureohirsuta 288
trapezioana 246 bahiensis 307
uiti 231 bangweolensis 357
ulei 250 barrosoi 298
venosa 251 bicornis 3 8
versicolor 208 var bicornis 319
villosa 218 var pubescens 319
williamsii 244 brachystachya 324
Cymosa (section) 28, ii6-I27 bracteata 302
Dactyladenia 357 bracteosa 292
Dahuronia 2I bullata 288
Dugortia 178 burchellii 338
Dulacia 357 butayi 357
Eulicania (section) 23 caduca 346
Exellodendron 195-201 canomensis 220, 357
barbatum I99 carbonaria 278
cordatum 197 castanea 302
coriaceum 197 caudata 319
gardneri 199 cauliflora 272
gracile 200 ciliata 284
Exitelia 201 cliffortiana 356
corymbosa 202 cognata 248, 357
Ferolia 178 collina 338
amazonica I85 conrauana 357
brasiliensis 189 coppenamensis 310
campestris 18 cordifolia 279
cordata 197 coriacea 328
coriacea 197 corymbosa 297
excelsa 185 cosmibuena 302
gardneri I99 cotticaensis 272
montana 182, I83 couepiflora 350
obtusifolia 194 cowanii 312
pohlii 189 cupheiflora 357
sprucei I87 damaziana 286
Geobalanus 22 davisii 283
oblongifolius 42 deflexa 294
pallidus 42 dodecandra 252, 357
Grangeria brasiliensis 285 dorvalii 273
Index of Scientific Names 405

Hirtella (cont.) Hirtella (cont.)

duckei 275 nemorosa 327
egensis 343 nitida 328
eglandulosa 357 obidensis 31o
elliptica 306 oblongifolia 328
elongata 309 octandra 8I, 357
enneandra 354 olivaeformis 357
eriandra 291 orbicularis 312
excelsa 345 paniculata Lam. 302
fasciculata 348 paniculata Sw. 292
filiformis 328 paraensis 335
fleuryana 357 pauciflora 355
floribunda 35I pendula 298
formicaria 275 peruviana 302
fruticulosa 357 physophora 272
gardneri 353 pilosissima 32
glaberrima Steud. 357 pimichina 342
glaberrima (Engl.) A. Chev. 357 piresii 283
glabrata 277 plumbea 338
glandistipula 340 pohlii 356
glandulosa 286 polyandra 252, 357
glaziovii 355 porosa 357
gracilipes 323 portoricensis 352
guainiae 275 praealta 3I9
guatemalensis 290 pulchra 338
guyanensis 313 punctillata 296
hebeclada 353 punctulata 306
hevanda 306 purusana 321
hexandra 328 racemosa Lam. 325
hirsuta 292 var glandipedicellata 330
hispidula 343 var hexandra 328
hoehnei 285 var metallica 327
hookeri 286 var oblongifolia 327, 328
hostmanniana 357 var racemosa 326
indecora 292 racemosa Ruiz & Pav. 302
insignis 280 rasa 316
jamaicensis 302 rodriguesii 310
juruensis 331 roraimae 357
katangensis 357 rosea 328
kuhlmannii 332 rotundata 285
lanceolata 310 rubra 284
lancifolia 337 rugosa 352
latifolia 308 sapinii 357
lemsii 334 scaberula 353
leonotis 300 scabra 316
lightioides 315 scandens 327
longifolia 333 schultesii 334
longipedicellata 339 selloana 353
macrophylla 295 silicea 344
macrosepala 276 sprucei 335
manigera 344 subsp meridionalis 338
martiana 340 standleyi 333
media 307 stipitadenia 288
megacarpa 356 strigulosa 327
melinonii 344 subglanduligera 284
mollicoma 289 subscandens 343
mollissima 289 subsetosa 343
montana 357 suffulta 308
mucronata 339 tentaculata 294
multiflora 302 tenuifolia 321
mutisii 300 thouarsiana 350
myrmecophila 271 tocantina 281
var. tetrandra 271 tomentella 353
406 Flora Neotropica

Hirtella (cont.) Licania (cont.)

triandra 301 columbarum 177
subsp media 307 compacta 146
subsp punctulata 306 conduplicata 177
subsp triandra 302 coriacea 28, 130
tubiflora 350 var nitida 154
ulei 276 cordata 136
velutina 286 costaricensis 94
vesiculosa 273 costata Io8
violacea 326 couepifolia 34
wachenheimii 286 crassifolia 162
zanzibarica 356 crassivenia 139
zenkeri 357 cruegeriana 171
Hirtelleae 178 cuprea 117
Hymenopus (section), 23, 28, 96-I 16 cuspidata 72
Icaco I4 cyathodes 13
Lecostemon 36I cymosa 122
Lepidocarpa 178 dahlgrenii 67
Leptobalanus (section) 23, 28, 62-85 davillaefolia 132
Licania 21-178 dealbata 18
affinis Fritsch 131 densiflora 116
affinis Kuntze 68 discolor I50
alba 14I divaricata 28, 103
albiflora 77 duckei 120
amapaensis 174 durifolia 44
angelesia 172 egensis 84
angustata 53 egleri 56
anisophylla 152 elata 59, 65
aperta 68 elliptica 133
apetala 28, 66 emarginata 73
var apetala 67 exiguifolia 177, 249
var aperta 68 fanshawei I 2
var pendula 67 flavicans 177
apiculata 151 floribunda 67
arachnites 69 foldatsii 136
arachnoidea 98 foveolata 80
araneosa 89 fritschii 59
arborea 86 fuchsii 79
aspera 76 galibica 123
aubletiana 177, 359 gardneri 72
axilliflora i6x glabra Io8
bahiensis I65 glabriflora I04
belemii 172 glauca 132
bellingtonii 145 glazioviana Io
benthamii o6 gonzalezii 56
biglandulosa 0o6 gracilipes 120
blackii 167 gracilis 143
boliviensis 43 grisea 126
bothynophylla 83 guianensis (Aubl.) Griseb., 28, 48
boyanii 127 guianensis Klotzsch 177
bracteata I55 hebantha 138
bracteosa 91 helvola 152
britteniana 59 heteromorpha I05
bullatifolia 86 var divaricata I03
buxifolia 128 var glabra o8
caldasiana 148 var grandifolia Io
calvescens 74 var heteromorpha Io6
canescens 134 var perplexans o09
capinensis 91 var subcordata 0o8
caracasana 68 hitchcockii 142
caudata Ioo hirsuta 92
celativenia 177 hoehnei 170
chocoensis 90 hookeri 84
Index of Scientific Names 407

Licania (cont.) Licania (cont.)

hookeri (cont.) nitida 154
var obtusa 83 oblongifolia 99
hostmannii 140 obovata 95
huberiana 91 obovatifolia 91
humilis 79 obtusifolia 176
hypagyrea 144 octandra 81
hylaea 67 subsp octandra 83
hypoleuca 28, 124 subsp pallida 84
var foveolata 127 odorata 177
var hypoleuca I26 oligantha 20, 177
impressa 118 operculipetala 97
incana 28, 29, 162 orbicularis 129
var axilliflora I6 organensis I60
indurata 170 ovalifolia I46
intrapetiolaris 105 pachystachya I57
irwinii 1 3 pallida Spr. ex Sagot I19
kanukuensis 116 pallida Britton 59
klugii 47 pallida (Hook. f.) Kuntze 84
krukovii 94 pallidula 130
kunthiana 144 paniculata 133
kuntzeana 68 paraensis I63
lanceolata 158 parinarioides 28, 91
lasseri 95 var latifolia 91
lata 65 parviflora Benth. 156
latifolia 28, 95 var conduplicata 126, 157
latistipula 103 var grandifolia 157, 164
laurifolia 115 var pallida 19, 157
laxa 115 var subfalcata 157
laxiflora 143 var submembranacea 144, 157
leptostachya I6i parviflora (Blume) Lemee 176
var axilliflora I6 parvifolia Huber 69
var crassifolia 162 parvifolia Pittier 126
leucosepala 52 parvifructa 121
licaniaeflora 91 pendula 67
littoralis I I persaudii 75
var cuneata 12 piresii I24
var littoralis I I platypus 55
longifolia 141 poeppigii I 5
longipedicellata 50 polita I 5
longipetala 62 prismatocarpa Io6
longistyla 78 pruinosa 154
lucida 167 pubiflora 68
macrocarpa 46 pulchravenia 150
macrophylla 99 pyrifolia 52
maguirei 70 reticulata 97
majuscula 140 retifolia 50
maranhensis 58 retusa 86
maritima 44 riedelii I 55
mattogrossensis 72 rigida 28, 85
maxima i65 riparia 57
membranacea 123 robusta 157
michauxii 42 rodriguesii 168
micrantha I52 rondonii 57
var triandra 150 roraimensis 176
microcarpa 126 rufescens I44
microphylla 149 salicifolia 89
minuscula 96 salzmannii 47
minutiflora (Sagot) Fritsch 57 sandwithii 142
minutiflora Cuatr. 96 savannarum I49
mollis 167 schomburgkiana I77
naviculistipula 398 sclerophylla 76
niloi 129 var myristicoides 77
408 Flora Neotropica

Licania (cont.) Moquilea (cont.)

sclerophylla (cont.) leucosepala 52
var scabra 77 licaniaeflora 91
seleriana 86 longistyla 78
sellowiana 83 macrocarpa 52
silvae I15 mattogrossensis 72
silvatica go minutiflora 57
sparsipilis 73 multiflora 178, 250
spicata 60 organensis i 60
splendens 172 orinocensis 67
sprucei 75 pallida 84
stahelii I46 paraensis 178, 210
stenocarpa 84 parilla 178, 218
steyermarkii 138 parviflora 176
stricta I60 pendula 67
subarachnophylla 88 platypus 55
subcordata o8 pyrifolia 52
subrotundata 139 riparia 57
surinamensis 177 rufa 178, 254
takutuensis 83 salzmannii 47
tepuiensis 174 sclerophylla 76
ternatensis I22 var myristicoides 77
tomentosa 5I var pohliana 77
var angustifolia 51 var scabra 77
var latifolia 5I sprucei 75
triandra I50 steudeliana 178
trigonioides 135 tomentosa 51
turbinata 65 turbinata 65
turiuva 8I turiuva 67, 8i
ulei 79 uiti 178, 231
unguiculata 60 utilis 83
urceolaris 130 Moquilea (section) 28, 42-62
utilis 83 Moquilea (subgenus) 26, 42-91
vaupesiana 164 Myrmecophila (section) 261, 27I-"76
velata 88 Parinari 178-195
veneralensis 174 barbaturm 195, I99
venosa 14I brachystz.chya 185
wilson-brownei 91 brasilF sis 189
wurdackii 63 campestns 181
Maranthes 20I-202 canescens I95, 249
corymbosa 202 cardiophylla 193
Microdesmia (section) 23, 28, 85-91 chocoensis 193
Moquilea 21 cordatum 195, I97
aubletiana 177, 359 coriaceum 195, I97
bothynophylla 83 corymbosa 202
bracteosa I77 excelsa 185
canomensis 177, 220 gardneri '95, 199
chrysocalyx 177, 245 glazioviana 185
comosa I77, 251 gracilis I95, 200
couepia 177, 208 guyanensis Fritsch 195, 313
cuspidata 72 guyanensis (Aubl.) Lem6e 195
elata 59 hostmannii I95, 248
eliti 177 klugii 190
floribunda 67 krukovii 195, 209
gardneri 72 laxiflora I85
glandulosa 77, 209 var lata I85
grandiflora I77, 236 littoralis I9I
guianensis 48 lucidissima 183
humilis 79 maguirei 90
hypoleuca 177 montana 182, I83
inaequalis 178, 227 nitidum I 2
kunthiana I78, 252 obtusifolia 194
kuntzeana 68 occidencalis I86
Index of ScientificNames 409

Parinari (cont.) Pleragina (cont.)

pachyphylla I89 umbrosissima 85
pajura I82 Prunus
parilis 193 icaco 15
parvifolia I91 Pulverulenta (section) 28, I27-134
philippinensis 172 Salmasia 259
pilosa 195, 220 guianensis 326
pohlii I89 racemosa 326
rodolphii 183 Sphenista 259
romeroi 400 peruviana 302
silvestris I85 Tachibota 259
sprucei 187 guianensis 326
Parinariopsis (subgenus) 28, 9I-92 Thelyra 259
Petrocarya 178 Theobroma
brasiliensis I89 alba 141
campestris I8I Trichocarya 22
excelsa I85 splendens 172
montana I82, I83 Zamzela 259
Pleragina 202 racemosa 302
odorata 51 rugosa 352
rufa 254