Melicocceae (Sapindaceae): Melicoccus and Talisia - CNCFlora

Organization for Flora Neotropica 

Melicocceae (Sapindaceae): Melicoccus and Talisia 

Author(s): P. Acevedo-Rodríguez 

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Source: Flora Neotropica, Vol. 87, Melicocceae (Sapindaceae): Melicoccus and Talisia (May 22, 

2003), pp. 1-178 

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FLORA NEOTROPICA MONOGRAPH 87 

MELICOCCEAE (SAPINDACEAE): 

MELICOCCUS AND TALISIA 

P. ACEVEDO-RODRiGUEZ 

F LORAt 

NEOTROPICAof, 

12001( Of CA021COIN4 

Publishedfor 

Organization for Flora Neotropica 

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The New York Botanical Garden 

Bronx, New York 

Issued 22 May 2003

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Library of Congress Cataloging-in-Publication Data 

Flora neotropica. - Monograph no. 1 - New York: Published 

for Organization for Flora Neotropica by The New York 

Botanical Garden, 1968v.: 

ill.; 26 cm. 

Irregular. 

Each issue has distinctive title. 

Separately catalogued andi classified in LC before monograph no. 40. 

ISSN 0071-5794 = Flora neotropica. 

1. Botany - Latin America - Classification - Collected works. 

2. Botany - Tropics Classification - Collected works. 3. Botany 

Classification - Collected works. I. Organization for Flora Neotropica. 

II. New York Botanical Garden. 

QK205.F58 581.98'012-dcl9 85-647083 

Library of Congress U8508i 

ISBN 0-89327-450-X 

03 0405 06070&-0910/98 765432 1

MELICOCCEAE (SAPINDACEAE): 

MELICOCCUS AND TALISIA 

P. ACEVEDO-RODRIGUEZ 

CONTENTS 

A bstract .. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2 

Resumen ......................................2 

Introduction ......................................3 

Taxonomic History .....................................4 

Morphology and Anatomy ...5..................................5 

Habit ......................................5 

Indumentum .......................................8 

Leaves ................................8 

Cataphylls ................................8 

Inflorescences .....................................8 

Flowers .......................................12 

Fruits and seeds ..................................... 

Embryo .......................................16 

Anatomy ......................................18 

14 

Pollen ..................................... 18 

Generic Relationships ..................................... 19 

Cladistic Analysis ..................................... 20 

Ecology and distribution ..................................... 23 

Conservation Status ..................................... 28 

Uses ...................................... 28 

Key to the American genera of Melicocceae ..................................... 28 

Taxonomic Treatment ..................................... 

Melicoccus ......................................28 

28 

Excluded Taxa ...................................... 50 

Talisia ...................................... 50 

Key to the subgenera ...................................... 50 

Talisia subgenus Pitombaria ..................................... 51 

Key to the species of Talisia subgenus Pitombaria ..................................... 51 

Key to the species of Talisia subgenus Talisia ..................................... 80 

Talisia subgenus Talisia ..................................... 88 

Excluded Taxa ..................................... 166 

Acknowledgments ..................................... 166 

Literature Cited ..................................... 167 

Numerical List of Taxa ..................................... 169 

List of Exsiccatae ..................................... 

169 

Index of Local Names ..................................... 

176 

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

177 

1

2 FLORA NEOTROPICA 

ABSTRACT 

Acevedo-Rodriguez, P. (Smithsonian Institution, Dept. of Systematic Biology (Botany), 

National Museum of Natural History, MRC-166, Washington, DC 20560-0166, USA). 

Melicocceae (Sapindaceae): Talisia and Melicoccus. Flora Neotropica Monogr. 87: iv + 1-179. 

2003. The tribe Melicocceae is characterized by the presence of a single apotropous, basal 

ovule per carpel; compound leaves with a distal rudimentary leaflet or process; actinomorphic 

flowers; non-arillate seeds, sometimes with a sarcotesta; and indehiscent unlobed fruits, not 

showing individual carpels. The American genera of Melicocceae (Talisia and Melicoccus) are 

further characterized by the presence of a sarcotesta. Melicoccus is characterized by an arboreal 

habit; paripinnately compound leaves with one, two or three pairs of leaflets; a distal 

rudimentary leaflet or process; racemes, panicles or thyrses; actinomorphic 4-(5-) merous 

flowers; bi- or tricarpellate gynoecium, with complete or less often incomplete septae; baccate 

fruits with coriaceous mesocarp; seeds with sarcotesta; and a lomatorrhizal embryo with 

erect cotyledons and punctiform radicle. The natural distribution of Melicoccus ranges from 

southern Mexico (Yucatan Peninsula) to South America on the periphery of the Amazon 

area, south to northern Argentina, Bolivia, and Paraguay and a disjunct species in Dominican 

Republic. Melicoccus bijugatus and M. oliviformis are distributed throughout the tropics 

through cultivation, while the other two species are largely restricted to their natural ranges. 

Melicoccusjimenezii of the Dominican Republic is an endangered species. 

Talisia is characterized by an arboreal or shrubby habit; pinnately compound leaves with 

a distal process or rudimentary leaflet; panicle-shaped or racemiform thyrses with flowers 

arranged in lateral dichasia; actinomorphic 5-merous flowers; petals with a single adaxial 

petal-like appendage; tricarpellate gynoecium with complete septa; baccate fruits; seeds with 

sarcotesta; and notorrhizal or lomatorrhizal embryos. The genus is distributed throughout 

continental tropical America, from southern Mexico (Yucatan Peninsula) to Paraguay, except 

Argentina, Chile, Uruguay, and the high Andes. The vast majority of species of Talisia are 

found in moist, non-flooded forest, either in the understory or part of the canopy layer. Talisia 

contains a few, widely dispersed species complexes, exhibiting considerable foliar variation. 

The remaining species are relatively vegetatively uniform with restricted distributions. 

The following new taxa are described and illustrated in this monograph: Melicoccus 

antioquensis, Melicoccus aymardii, Melicoccus espiritosantensis, Melicoccus novogranatensis, 

Melicoccus petiolulatus Talisia clathrata subsp. canescens, Talisia croatii, Talisia 

douradensis, Talisia equatoriensis, Talisia ghilleana, Talisia granulosa, Talisia hexaphylla 

subsp. elegans, Talisia hexaphylla subsp. multinervis, Talisia lanata, Talisia morii, Talisia 

parviflora, and Talisia velutina. 

Key Words: Sapindaceae, Melicocceae, Talisia, Melicoccus, tree architecture, Neotropics, 

cladistics, endangered species. 

RESUMEN 

Acevedo-Rodriguez, P. (Smithsonian Institution, Dept. of Sistematic Biology (Botany), 

National Museum of Natural History, NHB-166, Washington, DC 20560-0166, USA). 

Melicocceae (Sapindaceae): Talisia and Melicoccus. Flora Neotropica Mongr. 87: iv + 1-179. 

2003. La tribu Melicocceae esta caracterizada por la presencia de ovulos solitarios, basales 

apotropicos; hojas pinnaticompuestas con una hojuela terminal rudimentaria; flores 

actinomorfas; semillas no ariladas, a veces con presencia de sarcotesta; y frutos indehiscentes, 

no lobados. Los generos americanos de la Melicocceae (Melicoccus y Talisia), se distinguen 

adema's por la presencia de sarcotestas. Melicoccus se caracteriza por tener habito arboreo; 

hojas paripinnadas con uno, dos o tres pares de hojuelas y una hojuela distal rudimentaria; 

racimos, paniculas o tirsos; flores actinomorfas, 4(5)-meras, gineceo bi o tricarpelar con septas 

completas o con menos frecuencia incompletas; frutos carnosos con mesocarpo coriaceo; 

semillas con sarcotesta; embrion lomatorrizo, con cotiledones erectos y radicula punctiforme. 

Melicoccus se encuentra distribuido naturalmente desde el sur de Mexico (peninsula de

INTRODUCTION 3 

Yucatan) hacia Sur America a lo largo de la periferia de la region amazonica hasta Argentina, 

Bolivia y Paraguay y por una especies disjuncta en la RepRepublica Dominicana. La 

mayoria de las especies de Melicoccus se encuentran en bosques secos formando parte del 

dosel del bosque. Melicoccus bijugatus y M. oliviformis estan distribuidas a traves de los 

tr6picos por medio de cultivo mientras que las otras especies tienen en su mayoria 

distribuciones naturales, restringidas. Melicoccus]jimenezii de la Repu'blica Dominicana es 

una especie endemica amenazada. 

El genero Talisia se caracteriza por el habito arboreo o arbustivo; hojas pinnaticompuestas 

con una hojuela distal rudimentaria; tirsos paniculiformes o racemiformes con flores arregladas 

en dicasios laterales o con menos frecuencia paniculas; flores actinomorfas, 5-meras; gineceos 

tricarpelares, con septas completas; frutos carnosos conteniendo una semilla con sarcotesta, 

embrion notorrizo o lomatorrizo. Talisia esta distribuida a traves de America continental 

tropical desde el sur de Mexico hasta Paraguay, con excepcion de Argentina, Chile Uruguay, 

y la zona alta de los Andes. La gran mayoria de las especies de Talisia se encuentran en 

bosques h'umedos no inundables, ya sea en el sotobosque o formando parte del dosel de los 

bosques. Talisia contiene algunos complejos de especies ampliamente distribuidos y con 

gran variacion en los caracteres foliares. Las especies restantes son relativamente uniformes 

y tienen distribuciones bastantes restringidas. 

Los siguientes taxones nuevos son descritos en esta obra: Melicoccus antioquensis, 

Melicoccus. aymardii, Melicoccus espiritosantensis, Melicoccus novogranatensis, 

Melicoccus petiolulatus, Talisia clathrata subsp. canescens, Talisia croatii, Talisia 

douradensis, Talisia equatoriensis, Talisia ghilleana, Talisia granulosa, Talisia hexaphylla 

subsp. elegans, Talisia hexaphylla subsp. multinervis, Talisia lanata, Talisia morii, Talisia 

parviflora, y Talisia velutina. 

Palabras claves: Sapindaceae, Melicocceae, Talisia, Melicoccus, arquitectura de arboles, 

neotr6pico, cladistica, especie amenazada. 

INTRODUCTION 

The tribe Melicocceae (Sapindaceae), as currently 

recognized, contains New World and Old World elements. 

In the New World it is represented by the genera 

Melicoccus P. Browne and Talisia Aublet, which 

clearly form a cohesive unit sharing numerous morphological 

characters, but especially the presence of indehiscent 

fruits and seeds with a sarcotesta. The Old 

World elements differ greatly from the New World 

genera and perhaps do not belong in the Melicocceae at 

all. Since resolution of this problem would require an 

in-depth analysis of the whole Sapindaceae, the present 

paper deals only with the New World Melicocceae. 

Talisia is the largest of the two genera ofAmerican 

Melicocceae, with 52 species and 5 subspecies. Vegetatively, 

Talisia may be confused with numerous arboreal 

Sapindaceae or with other compound, altemateleafed 

trees in various families. However, it is possible 

to vegetatively distinguish Talisia from other Sapindaceae 

genera as follows: Talisia has leaflets with entire 

margins, whereas most Cupania Linnaeus, some 

Averrhoidium Baillon, Diatenopteryx Radlkofer, and 

Ungnadia Endlicher, have leaflets with serrate or dentate 

margins; from Vouarana and many species of 

Matayba, Talisia may be distinguished by the absence 

of domatia on secondary vein angles; and from Athyana 

(Grisebach) Radlkofer, Talisia lacks a terminal leaflet. 

Some previous taxonomic experience is required in 

order to differentiate other genera of Sapindaceae with 

similar suites of characters from Talisia, especially 

needed would be flowers or fruit. Occasionally, species 

of Euplassa (Proteaceae), Ophiocaryon (Sabiaceae), 

Simaba (Simaroubaceae), Guarea (Meliaceae), and 

Fabaceae are mistaken for Talisia. The former three are 

only reliably distinguished from Talisia when fertile, 

while Guarea is easily distinguished by its leaves with 

a pair of terminal minute, immature leaflets (vs. a single 

rudimentary one in Talisia), and the Fabaceae by the 

presence of stipules. 

Species of Talisia are usually distinguishable by their 

vegetative characters, with the exception of a small 

number of species that are highly variable with respect 

to foliar characters. In the absence of flowers, T cerasina 

is easily mistaken for T retusa or sometimes for T 

macrophylla, and T macrophylla is easily confused with 

T nervosa or T eximia, and T nervosa with T sylvatica. 

The inadequacy of collections often makes the interpretation 

of species difficult. This phenomenon is usually 

a result of the tendency of collectors to press only 

leaves that easily fit into their plant press, or to collect 

only fragments of a leaf without properly recording its 

characteristics. As a result, new taxa have been described 

based on unusually large leaves or leaflets.


Melicoccus, on the other hand, being a smaller, Tristira and Tristiropsis. The position of Tristira and 

uniform genus, is easily recognized by the jugate, Tristiropsis within the Melicocceae, however, is at its 

bijugate or trijugate leaves. Its species are well differ- best dubious. 

entiated, perhaps as a result of geographic isolation. Melicoccus was described by P. Browne in 1756 in 

Since the last monograph of Melicoccus and Talisia his account of the natural history of the island of Ja- 

(Radlkofer, 1931-34), much more material has become maica. The description of the new genus was based on 

available, making necessary a reinterpretation of spe- the commonly cultivated genip tree (Melicoccus 

cific concepts. Morphological gaps have been filled bijugatus), which was introduced to Jamaica from 

calling for the lumping of species, and much more varia- Surinam. In 1760, Jacquin described M. bijugatus, the 

tion has become apparent making necessary the descrip- first specific epithet for the genus. Two years later, 

tion of new taxa. On these grounds, a revision of the Linnaeus published the orthographic variation 

American Melicocceae is fully justified. 

Melicocca biuga citing P. Browne (1756) and the manuscript 

of Jacquin's Select. Stirp. Amer. Hist., tab. 72. 

TAXONOMIC HISTORY (as 71) as the source of his name. Following Linnaeus' 

publication, numerous specific names have been pub- 

In 1888, Radlkofer published a synopsis of Sapin- lished in Melicocca, the orthographic variation introduced 

daceae, in which he recognized 15 tribes, one of which by him. The original orthography of the genus has been 

was the Melicocceae, containing the genera Melicoccus, used only twice, once by Hasskarl in 1842 [Melicoccus 

Talisia, Glenniea Hook. f., Castanospora F. von 

amoenus Hasskarl = Lepisanthes amoena (Hasskarl) 

Muller, Lecaniodiscus Bentham, Eriandrostachys Leenhouts] and more recently by Kitanov in 1979 

Baillon, Macphersonia Blume, Tristiropsis Radlkofer, (Melicoccus bijuga f. alata =Melicoccus bijugatus). 

and Tristira Radlkofer. In a later revision of his familial However, Hasskarl a year later, changed the spelling 

classification (Radlkofer, 1931-34), he added of his name to conform with Linnaeus' spelling. In 

Hedyachras Radlkofer to the Melicocceae and trans- addition, 14 specific names have been published under 

ferred Lecaniodiscus, Eriandrostachys, and Macpher- Melicocca, all of which (except M. oliviformis and M. 

sonia into the closely related tribe Schleichereae. The lepidopetalus) belong to other genera or are synonyms. 

genus Strophiodiscus Choux was added posthumously To add confusion to the correct spelling of 

to the Melicocceae by the editors of his 1931-34 revi- Melicoccus, Index Kewensis had listed the original 

sion for a total of eight genera. 

spelling as the orthographic variant Melicocca. In ad- 

Capuron (1969) reduced Strophiodiscus to a syn- dition, there has been a proposal (Repert. Spec. Nov. 

onym of Plagioscyphus, which he placed in the tribe Regni Veg. 53: 178. 1944) to conserve the spelling 

Schleichereae. Glenniea was placed by Leenhouts Melicocca vs. Melicoccus, which was rejected by the 

(1975) in the Lepisantheae by implication when he re- Special Committee for Pteridophyta and Phanerogamae 

duced the genus Crossonephelis to a synonym of by the IAPT (Taxon 3: 241. 1954). As a result, the spell- 

Glenniea. Muller and Leenhouts (1976), who proposed ing Melicocca is to be corrected to Melicoccus. 

a modified system of classification for the Sapindaceae, In 1997, I transferred Talisia jimenezii into 

retained only five genera in Melicocceae. They found Melicoccus increasing the number of species to three. 

the New World genera Talisia and Melicoccus to have In this revision five new species are described and a 

pollen types (type A & Al) different from those of the transfer is made, increasing the total number of species 

Australasian genera Castanospora, Tristira, and in Melicoccus to ten. 

Tristiropsis (type B). This distinct separation of the Talisia was described in 1775 from French Guiana 

Melicocceae into two groups, along with their complete by Aublet, who derived the name from the vemacular 

absence from Africa, made them cast some doubts on name Toulichi. The first species to be described in the 

the monophyly of the tribe. 

genus was Talisia guianensis. In 1798 Vahl described 

In 1999, Klaassen analyzed the wood anatomy of Talisia hexaphylla from South America, and proposed 

Castanospora and found it to deviate from the remain- T rosea as a more appropriate name for Talisia 

ing members of Melicocceae. Although she did not ex- guianensis because of the pinkish inflorescence axis 

clude Castanospora from the Melicocceae, she sug- and immature calyx. De Candolle (1824) described 

gested it might belong in the tribe Lepisantheae. In Talisia glabra, a synonym of T guianensis. He noted 

addition, she cast some doubt on the placement of that the recently described genus Acladodea of Ruiz 

Tristira and Tristiropsis within the Melicocceae, how- and Pavon (1798) was congeneric with Talisia, transever, 

she did not exclude them from the Melicocceae. ferring Acladodea pinnata into Talisia as T acladodea. 

Currently, the Melicocceae contains the American gen- However, such a transfer is contrary to the code and was 

era Talisia and Melicoccus and the Old World genera corrected by Radlkofer in 1878 to T. pinnata. In 1829,

MORPHOLOGY AND ANATOMY 5 

Cambessedes described T. mollis from French Guiana, axillary or terminal on short, lateral branches and do 

noting that this was the same species treated by de 

Candolle in 1824 as T guianensis. In 1850 Bentham 

not seem to develop further into sympodial axes. 

described T laxiflora, a taxon currently recognized as a 

Talisia 

subspecies of Matayba guianensis Aublet. A few years Species of Talisia are trees (usually with buttressed 

later, Triana and Planchon (1862) transferred Comato- bases), treelets, or shrubs (Fig. la& b), described by 

glossum strictum to Talisia as T stricta, noting that three architectural models (Halle et al., 1978). About 

the newly described genus Comatoglossum of Karsten 40% of the species are treelets (Fig. la) i.e., unbranched 

& Triana (Triana, 1854) was essentially Talisia. shrubs described by the Chamberlain model. The habit 

In 1878, Radlkofer published a forerunner to his construction of these species is modular through linear 

monumental monograph of Sapindaceae. In this paper sympodial growth, where successive meristems contreating 

Talisia, he published 18 new species, trans- tribute to the construction of a seemingly unbranched 

ferred eight others into Talisia, and synonymized stem. The system operates through the succession of 

Racaria, a generic name published simultaneously with branches with determinate growth bearing a terminal 

Talisia by Aublet. As a result, he increased the total inflorescence. Once the inflorescence reaches maturity, 

number of known species of Talisia from four to 30, an axillary meristem basal to the inflorescence develand 

adopted an infrageneric classification that he fol- ops into an orthotropic shoot that displaces the inflolowed 

throughout his career. This paper was equally rescence to the side. In this way, the new shoot contribimportant 

for highlighting the distinctions between utes to the construction of the leader stem producing a 

Talisia and other neotropical arboreal genera of Sapin- seemingly unbranched shrub (Fig. 2b and 2d). 

daceae. This giant leap was followed by a moderate The remaining species do not seem to conform to 

series of contributions. Between 1900 and 1914, any of the architectural models proposed by Halle et al. 

Radlkofer published eight new species and made one (1978). The information available for the arboreal spenew 

transfer in Talisia. When his Sapindaceae mono- cies i.e., bearing a trunk and lateral branches, is rather 

graph was posthumously published from 1931 to 1934, limited as it comes from field observation (and confirforty-one 

species of Talisia were recognized, includ- mation with herbarium specimens) from T hemidasya, 

ing T. princeps published by Oliver in 1888 and T. T hexaphylla, and T praealta. Accordingly, they present 

panamensis (a synonym of T hexaphylla subsp. a complex system that changes through the different 

hexaphylla) published by Pittier in 1918. 

developmental phases of the plant. The early develop- 

Since Radlkofer's death in 1927, an additional 32 ment of the plant conforms to the Chamberlain model 

new taxa have been published in Talisia by 13 authors. showing a pattem of orthotropic sympodial growth by 

Of these, Steyermark described the highest number with substitution due to meristems with determinate growth. 

a total of eight, followed by GuarimNeto (1979, 1983) This phase is followed by the proleptic production of 

with five, Standley (1930, 1931a, 1931b, 1933) with orthotropic vegetative axes with terminal and axillary 

four, and the remaining authors with one or two spe- inflorescences. This mixed model seems to describe 

cies each. Of these 32 taxa, only 12 are recognized in about 50% of the species of Talisia. 

this treatment. The total number of taxa so farpublished Talisia angustifolia, a species with shrubby habit 

in Talisia is 78, however, only 45 of these are recog- and horizontal underground stems, seems to have an 

nized herein. In addition, 12 novelties are described for architectural model different from that previously dea 

total of 52 species and five subspecies. 

scribed. My knowledge of T angustifolia development 

is very limited, as I have no information on its 

MORPHOLOGY AND ANATOMY 

early stages of development. The only information 

available to me comes from herbarium specimens. Accordingly, 

the plagiotropic underground stems are of 

HABIT 

higher magnitude than the orthotropic ones. The pla- 

Melicoccus 

Species of Melicoccus are medium-sized trees with 

continuous and diffuse branching (Fig. Ic & d). The 

giotropic axis seems to be the result of sympodial growth 

by substitution with orthotropic branches arising 

proleptically. The orthotropic branches have determinate 

growth ending in an inflorescence but also bear 

little information I have on its architecture (observa- axillary inflorescences. This model is mixed and doesn't 

tions on seedlings and juveniles of M bijugatus) indi- seem to conform to any ofthe models proposed by Halle 

cates that the leading stem is the result of a single apical et al. (1978). 

meristem. Axillary meristems develop proleptically into In general, architectural models in Talisia are imlateral, 

plagiotropic branches. Inflorescences are either perfectly known because the necessary underlying


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Fig. 2. Vegetative features of Talisia. A. rudimentary distal leaflet or process in T hemidasya. B & D. Chamber- 

lain architecture model in T hemidasya. C. cataphylls in T megaphylla. (A, B, D from Acevedo R. et a! 4966; C 

from Acevedo R. et al. 4798).


developmental data are not available. At present, the best 


estimate of this character is inferred from specimen data 

or field observations. 

Leaves in Talisia are spirally arranged, paripinnately 

or imparipinnately compound, and devoid of stipules 

INDUMENTUM 

(Fig. 2d). New leaves have been documented to carry 

green leaflets in most species, with the exception of T 


hemidasya, which produces reddish juvenile leaflets 

(Fig. 2d). Petioles are usually pulvinate and the rachis 

Species of Melicoccus are essentially glabrous, with 

always ends in a rudimentary leaflet or process (Fig. 

scattered papilliform trichomes on the abaxial surface 

2a). Leaflets are alternate, opposite, or subopposite with 

of leaflets and along inflorescence axes. Bracts, sepals, 

entire margins. Their number varies from strictly two 

and petals are woolly-pubescent along their margins. 

as in T squarrosa to variably up to thirty as in T macro- 


phylla. Leaflet size is quite variable ranging from 3 cm 

long in T angustifolia to 79 cm long as in T bullata. 

Species of Talisia are either glabrous or exhibit sev- Venation is brochidodromus (sensu Hickey, 1979; Fig. 

eral different kinds of indumentum, which are pre- 

5a-b), adaxially impressed and abaxially prominent or 

dominantly simple, non-glandular trichomes (Figs. 3 less often plane on both surfaces and therefore inconand4). 

Less frequently, multicellular-glandular orpapilli- spicuous. Primary veins are moderate (Fig. Sb) to stout 

form trichomes are present. Trichomes occur as a pure (Fig. 5a) and unbranched. Secondary veins are alterstand 

or as a mixture of different types (Fig. 4c) de- nate, sometimes some of them opposite (Fig. 5a-b), 

pending on the species or the organ of the plant upon uniformly curved (Fig. Sb) or straight (Fig. 5a), and 

which they are found. Papilliform trichomes are usu- sometimes collected into a conspicuous vein that runs 

ally restricted to the adaxial surface of petals and abaxial parallel to the margin (Figs. 5a, 6a). Tertiary veins form 

surface of petal appendages, but they are sometimes a reticulum [= reticulate] (Fig. 5b, 6a) or run perpenfound 

on bracts and inflorescence axes. Sericeous in- dicular or nearly so to the midvein [= clathrate or subdumentum, 

although found throughout a wide range clathrate respectively] (Fig. 5a, 6b). Leaflet pubescence 

of structures, is predominantly found on the adaxial is variable but usually restricted to the abaxial surface 

surface of petal appendages. The remaining types of or the midvein, and commonly represented by simple 

indumentum are widely distributed and may be clas- trichomes. Glandular trichomes are present in only a 

sified as puberulent (Fig. 4a-b), appressed-pubescent few species. Several species exhibit a granular texture 

(Fig. 3a), tomentose (Fig. 4c), tomentulose, sericeous, on the abaxial surface when examined with a dissectvelutinous, 

setigerous (Fig. 4e), pubescent [= soft, ing scope at 20-50 x magnifications. Observation of 

short trichomes] (Fig. 3e-f), hirsute, woolly (Fig. 4d), this feature with SEM revels it to be stomata (Fig. 7). 

pilose, pilosulous, or furfuraceous (Fig. 3c-d). Type Petiolules are pulvinate, moderately to greatly enlarged, 

of indumentum is a useful diagnostic character at the bulbiform or slender and cylindrical, and usually adspecies 

level since indumentum variation within spe- axially furrowed. 

cies is rather uniform. For example, the presence of Foliar characters are of moderate taxonomic value, 

glandular hairs is characteristic of T hemidasya and because only a few species can be distinguished based 

T morii, while the presence of setigerous hairs on solely on these characters. This would include the numleaves 

is unique to T setigera. 

ber and size of leaflets and the shape of the leaf rachis. 

LEAVES 

However, many species of Talisia are vegetatively similar 

and in some cases identical, requiring the presence 

of fertile material for their identification. Notable are 


the difficulties in differentiating sterile material of T 

Leaves in Melicoccus are spirally arranged, jugate, macrophylla from T eximia and T cerasina, and T 

bijugate, or trijugate and devoid of stipules. Petioles sylvatica from T nervosa. 

are obscurely pulvinate, flattened, and occasionally 

winged or nearly terete. Rachises are flattened, winged 

or nearly terete, and end in a rudimentary leaflet or process, 

although a fully developed distal leaflet is some- 

CATAPHYLLS 


times produced. Leaflets are opposite or sometimes Cataphylls in Melicoccus (referred by Radlkofer as 

alternate, with entire margins. Venation is brochido- perules) are scale-like, ovate to nearly deltate with disdromus, 

with a stout or less often moderate, unbranched sected margins, and are clustered on axillary buds and at 

midvein. Secondary veins are alternate and uniformly the base of inflorescences. There is no significant variacurved; 

tertiary veins form a reticulum. 

tion of this structure in Melicoccus and therefore they


Fig. 3. Indumentum of Talisia. A-B. T angustifolia. A. trichomes on petiole (appressed pubescent). B. detail of 

trichomes. C-F. T furfuracea. C. trichomes on stem (furfuraceous). D. detail of trichomes. E. trichomes on abaxial 

surface of leaflet (pubescent). F. detail of trichomes. [A-B from Hatschbachz 159JJ (US); C-D from Acevedo R. 

& Grimes 4863 (US); E-F from Sabatier & Provost 2202 (US)].


Fig. 4. Indumentum of Talisia. A. trichomes on petiolule of T acutifolia (puberulent). B. trichomes on stem of 

T~ clathrata subsp. canescens (puberulent). C. trichomes on stem of T hemidasya (tomentose). D. trichomes on 

abaxial leaflet surface of T lanata (woolly). E. trichomes on abaxial leaflet surface of T setigera (setigerous). (A 

from Acevedo R. et al. 8456 (US); B from Acevedo R. et al. 4956 (US); C from Boschwezen 2574 (MO); D from 

Garz6n et al. 145 (COAH); E from Cornejo & Bonifaz 3878 (US).


1 cm 

a b 

Fig. 5. X-rays of Talisia leaflets. A. T furfuracea. B. 

T firma. (A from Sabatier & Prevost 2202 (US); B from 

Liesner 8883). 

Fig. 6. X-rays of Talisia leaflets. A. T nervosa. B. T 

clathrata subsp. clathrata. (A from Faber-Langendoen 

& Hurtado 1934; B from Klug 2509). 

i. .- S a n a e f f i o o ( d G il 

Fig. 7. Stomata on abaxial leaflet surface of Talisia longifolia (from de Granville 7033).


Fig. 8. Dichasia in Talisia and Melicoccus. A. simple 

dichasium in T nervosa. B. dichasium reduced to a single 

flower in M. aymardii. (A from Cuatrecasas 17010; B 

from Aymard & Ortega 2521). By B. Angell). 

are of no taxonomic value within the genus. However, 

they are useful in distinguishing Melicoccus from Talisia. 


Cataphyll was the term utilized by Radlkofer 

(1931-34) in reference to the perules or bud scales in 

Talisia. They are clustered at the ends of branches, on 

axillary buds, and at the base of inflorescences. Their 

size and form vary considerably, in some species they Fig. 9. Inflorescences of Talisia. A. T firma. B. T 

are large (up to 25 cm long) resembling undeveloped acutifolia. (A from Acevedo R. et al. 7963; B from 

leaves (Fig. 2c), while in others they are pinnatifid 

and a few cm in length (Fig. 2b), or even reduced to 

scale-like or spine-like structures less than 1 cm long. 

Acevedo R. et al. 8362). 

The latter are the most common type of cataphylls 

present in Talisia. Cataphylls, although not universally 

present in Talisia, are useful for differentiating Talisia 

vegetatively from other arboreal Sapindaceae and from 

trees in general with spirally arranged, compound 

leaves. They also can be of diagnostic value at the species 

level. 

often cauliflorous or ramiflorous. Inflorescence type 

and shape are of limited taxonomic value since there is 

widespread variation throughout the genus. 

Bracts are minute, scale-like structures subtending 

a dichasium. Bracteoles on the other hand are minute, 

persistent or deciduous structures subtending a secondary 

axis of the dichasia. Bracts and bracteoles are es- 

INFLORESCENCES 

sentially similar in shape and structure; however, bracts 

are usually bigger than the bracteoles. 


Inflorescences in Melicoccus are panicles or racemes 

Pedicels have an abscission zone or articulation 

anywhere from base to near the apex. 

with solitary flowers distributed along the inflorescence 

axis (Figs. 12d, 14a) or less often paniculate thyrses 

FLOWERS 

where some of the dichasia (mostly the distal ones) are 


reduced to a single flower. Flowers are subtended by a 

small deciduous to persistent bract. Pedicels are articulate 

or not. 

Flowers in Melicoccus are actinomorphic or nearly 

so, staminate (Fig. 12c & d) or pistillate, with rudimentary 

stamens or pistil. The calyx is 4- or 5-merous, with 

imbricate, nearly free sepals, sometimes produced in 


two unequal pairs. Petals are 4 or 5, distinct, with im- 

Inflorescences in Talisia are thyrses with flowers bricate aestivation, inserted on the base of an extragrouped 

in compound or simple dichasia (Fig. 8a). They staminal nectary disc. They are erect or reflexed, often 

are panicle-shaped (Fig. 9a-b), fasciculate (Fig. lOa-b), with woolly-pubescent margins, lacking appendages, 

or racemiform, terminal, axillary, supra-axillary, or less sometimess with a pair of marginal projections or

MORPHOLOGY AND ANATOMY 

__ 

- - 1E 

__ - 

_i _ 

= i i L I . 

1_ ,_ [, 

''I | 2i. 

. .I 

I 

E_ z ! 

_ 

_8_ 

_ 

I 

| ' 

_ _X_ .l 

_^ | 

l l 

_ I I i 

Fig. 10. Inflorescences of Talisia. A. T. clathrata subsp. canescens. B. T. megaphylla. (A from Acevedo R. 5012; 

B from Mori et al. 23121, photo by Carol Gracie). 

appendages above or at the base. The disc is extra- 

staminal, swollen, glabrous, annular, with a 4-5-lobed 

outline. Stamens are 8, the filaments are unequal, gla- 

brous or less often wooly pubescent, anthers are 

basifixed or dorsifixed, elliptic or ovate. The ovary is 

bi- or tricarpellate, with as many locules as carpels or 

unilocular (the septa incomplete), with a single basal, 

campylotropous or apotropous ovule per carpel; the style 

is solitary, short to very short, with a discoid-bilobed 

or trilobed, papillate stigmatic surface. 

13


Melicoccus is often regarded as dioecious but there 

is no literature dealing with its sexuality. Individual trees 

are often collected bearing either staminate or pistillate 

flowers with practically no overlap between sexes. 

However, I have been able to find one individual tree 

bearing flowers of both sexes at the same time (Acevedo 

R. & Clubbe 10956). Whether this is an isolated case 

of monoecy or a fairly common, undocumented phenomenon 

cannot be determined at present. Further studies 

are needed to elucidate whether Melicoccus is a sequentially 

monoecious or not. 


Flowers in Talisia are actinomorphic, functionally 

staminate or pistillate. Staminate flowers are distinguished 

by the presence of dehiscing anthers and a pistillode, 

while pistillate flowers by the presence of nondehiscing 

anthers and a fully developed pistil. With 

f~~~~~~ 

respect to the perianth, disc, and pubescence, staminate 

and pistillate flowers are essentially similar. Perianth is 

5-merous. Sepals are imbricate, free or connate to various 

degrees at base, and usually of similar size. Petals 

are distinct, with imbricate aestivation, inserted on the 

base of an extrastaminal nectary disc. They are erect or 

reflexed, entire, with a single, adnate, adaxial petaloid 

appendage [subgenera Talisia (Figs. 1 lb and 12b) and 

Pitombaria (Figs. 1 la and 12a)].The petaloid appendages 

are comparable to the petals in size and shape. They 

are simple or rarely bifid, and very often sericeous or 

tomentose. The disc is extrastaminal, annular or cupshaped, 

with a 5-8-lobed outline, glabrous or with various 

indumenta, and perhaps responsible for the production 

of nectar as a reward to pollinators. The number Fig. 11. Flowers in Talisia. A. T: subgenus Pitombaria 

of stamens is usually 5 or 8 but there is considerable (T clathrata subsp. canescens). B. T: subgenus Talisia 

variation with species containing from 5 to 8 or from 7 (T: bullata). (A from Mori et al. 21365; B from Eggers 

15004). By B. Angell. 

to 10 stamens. Filaments are glabrous or variously pubescent, 

of equal lengths or nearly so, or unequal and 

erect. Anthers are basifixed, with either oblong to linear 

or elliptic to ovate outline. Talisia subgenus Talisia and pistillate flowers. This breeding system has been 

contains 5-8 stamens with equal or nearly equal length, documented for other members of Sapindaceae as 

with oblong to linear anthers while T. subgenus well (Bawa, 1976; Subba Reddi et al., 1983; Acevedo- 

Pitombaria contain 8 or rarely 10 stamens with unequal Rodriguez. 1993; Genise V. Somner, unpub. data) 

lengths and elliptic to ovate anthers. The ovary is invariably 

3-carpellate, 3-locular, with a single basal, 

FRUITS AND SEEDS 

campylotropous or apotropous ovule per locule, and 

the style is solitary, short to elongated, with 


elongatetrigonous 

or short-capitate stigmatic surfaces. 

Fruits of Melicoccus are very much like those of 

There is little overlap between the temporal occur- Talisia, being indehiscent, unilocular and bearing one 

rence of pistillate and staminate flowers; herbarium seed (resulting from the abortion of I carpel) or two 

specimens only exceptionally contain flowers of both seeds, with a leathery pericarp (Fig. 13d). Fruits are 

sexes. The plants are either male or female at any one time. ellipsoid or ovoid (Fig. 13b &14b), or less often glo- 

My own observations in the field indicate that species bose-bilobed when containing two seeds. Seeds have 

of Talisia are sequentially monoecious, where individual a fleshy, edible testa and closely conformn to the shape 

plants go through the altemate production of staminate of the fruit.

MORPHOLOGY AND ANATOMY 1 5 

Fig. 12. Talisia and Melicoccus. A. portion of inflorescence in T firma. B. flowers in T megaphylla. C. inflo- 

rescence branch in M. oliviformis subsp. intermedius. D. inflorescence branch in Melicoccus bijugatus. (A from 

Acevedo R. et al. 7963; B from Mori et al. 23121, photo by Carol Gracie; C from Acevedo R. et al. 3473; D from 

Acevedo R. 5441).


Fig. 13. Fruits of Talisia and Melicoccus. A. infructescence in T mollis. B. infructescence in M. oliviformis 

subsp. intermedius. C. fruit with mesocarp cut away showing seed with fleshy testa in T mollis. D. fruit with me- 

socarp cut away showing seed with edible fleshy testa in M. oliviformis subsp. intermedius. (A & C from Acevedo 

R. et al. 4914; B & D from Acevedo R. et al. 3515). 


Fruits in Talisia are baccate, i.e., indehiscent, unilocular 

or less often 2-locular (resulting from the abortion of 

2 or 1 carpels) with a leathery to woody (Fig. 13c) pericarp. 

Fruits are usually ellipsoid or ovoid (Fig. 13a) but 

in a few species they are depressed-globose. There is a 

single seed per locule, which possesses a more or less 

fleshy, edible testa, commonly referred to as sarcotesta 

(Fig. 13c). Seeds closely conform to the shape ofthe fruit, 

except in the case where there are two seeds. In this case, 

the surface where both seeds meet is flat, with the out- 

line of both seeds conforming to the shape of the fruit. 

EMBRYO 


Embryos in Melicoccus are predominantly 

lomatorrhizal with erect straight or slightly curved coty-


Fig. 14. Melicoccus bijugatus. A. inflorescence. B. infructescence. (Both from Acevedo R. 5441). 

ledons (Fig. 1 5g). Melicoccusjimenezii has notorrhizal ferentiated and show a suture only on their distal portion 

embryos with the upper cotyledon being much bigger (Fig. 15d). Lomatorrhizal embryos (Fig. 15f) have straight, 

than the lower one. 

erect cotyledons of similar size parallel to the radicle. The 

radicle in both types of embryos is short, flattened and 


well defined, placed in a small depression or radicle pocket. 

Embryos in Talisia are fleshy and have been defined Embryo characters are of little taxonomic value 

as notorrhizal or lomatorrhizal. Notorrhizal embryos have within Talisia. Most species have notorrhizal embryos 

a suture between the cotyledons positioned perpendicular and the few species that have lomatorrhizal embryos 

to the radicle. Cotyledons of notorrhizal embryos are hori- also have notorrhizal embryos as well. However, 

zontally superimposed (Fig. l5a-b) or obliquely so (Fig. Talisia with its notorrhizal embryo can easily be told 

1 Sc) or rarely laterally oblique (Fig. 1 Se) and of equal or apart from Melicoccus which mostly has lomatorrhizal 

unequal size. Sometimes the cotyledons are poorly dif- embryos with erect cotyledons.


1cm 

Fig. 15. Embryos of Talisia and Melicoccus. A-B. T japurensis. A. dorsal view. B. lateral view. C. T bullata, 

lateral view. D. T acutifolia, lateral view. E-F. T stricta exhibing two types of embryo configurations, notorrhizal 

and lomatorrhizal respectively. G. M. bijugatus, dorsal view of lomatorrhizal embryo. (A-B from AssunQao & 

Pereira 261; C from Cogollo et al. 4572; D from Adis 7; E-F from Fernandez A. et al. 5484). 

ANATOMY 

Melicoccus in an effort to evaluate the tribal classifica- 

Generalities on the vegetative anatomy of Melicoccus 

and Talisia were studied by Metcalfe and Chalk (1957) 

in their monumental Anatomy oftheDicotyledons. Wood 

anatomy of Talisia was studied by Mennega (1972) in 

tion of Sapindaceae. She found that the Melicocceae 

sensu Muller and Leenhouts (1976) was rather homogeneous 

and that the genera of Melicocceae share numerous 

features with other tribes of Sapindaceae. 

an effort to provide additional taxonomic characters for 

the classification of the genus. She found that Talisia 

POLLEN 

was well defined by its wood anatomy and therefore, 

easily distinguished from other genera of New World 

Sapindaceae. Talisia can be recognized by numerous 

anatomical features, in particular by the aliform, aliformconfluent 

and confluent banded parenchyma. However, 

she did not find the variation significant enough to distinguish 

species from one another. She also noticed that 


Pollen in Melicoccus is of type A (Fig. 1 8d-f), i.e, 

tricolporate, the colpi elongated, extended to polar region; 

the polar view is obtusely triangular, with convex 

or straight sides; the equatorial view is rounded in 

the polar areas. Melicoccus pollen is similar to those of 

Talisia wood is similar to that of Melicoccus, except Talisia subgenus Pitombaria. 

that Melicoccus has broader and more regular parenchyma 

bands and rays which are 2-3 cells wide (vs. 


homocellular in Talisia). More recently, Klaassen Pollen grains in Talisia can be classified into two types 

(1999) examined the wood anatomy of Talisia and as defined by Muller and Leenhouts (1976). Pollen type

GENERIC RELATIONSHIPS 19 

Fig. 16. Scanning Electron Micrographs of Talisia 

pollen (A-D = type Al; E-F = type A). A-B. T guianensis. 

A. polar view. B. equatorial view. C-D. T marleneana. 

C. polar view. D. equatorial view. E-F. T. velutina. 

E. polar view. F. equatorial view. (A-B from Cowan 

38809; C-D from Davidson 10073; E-F from Schunke 

V 2637). 

Fig. 17. Scanning Electron Micrographs of Talisia 

pollen (A-B = type Al; C-F = type A). A. T dasyclada, 

polar view. B. T ghilleana, polar view. C-D. T croatii. 

C. polar view. D. equatorial view. E-F. T hexaphylla 

subsp. hexaphylla. E. polar view. F. equatorial view. (A 

from Vasquez & Criollo 5811; B from Silva et al. 2153; 

C-D from Barbour 5720; E-F from Haught 1608). 

Al (Figs. 16a-d and 17a-b) is tricolporate, the colpi are ters, i.e., a single apotropous, basal ovule per carpel; 

short, not extending into polar areas; the polar view is compound leaves with a distal rudimentary leaflet or 

obtusely triangular, with convex or concave sides; the process; actinomorphic flowers; non-arillate seeds; and 

equatorial view is slightly flattened in polar areas. This indehiscent, unlobed fruits, the combination of which 

pollen type is exclusively found in T subgenus Talisia. is unique within the Sapindaceae. The Melicocceae have 

PollentypeA(Figs. 16e-f, 17c-fand 18a-c) istricolporate, been distinguished from "closely related" tribes by 

the colpi elongated, extended to polar region; the polar means of carpological characters. However, it is unview 

is obtusely triangular, with convex or straight sides; likely that these were uniquely derived characters, castthe 

equatorial view is rounded in the polar areas. This type 

of pollen is predominant in T subgenus Pitombaria, but 

may be found in T subgenus Talisia as well. 

ing doubts on the monophyly of the Melicocceae. Of 

the seven Old World genera originally included in the 

Melicocceae, only three (Castanospora, Tristira, and 

Tristiropsis) are currently retained in the tribe. How- 

GENERIC RELATIONSHIPS 

ever, in my opinion, they do not belong in the 

Melicocceae as they do not share the basic fruit mor- 

MELICOCCEAE 

phology of the type genus Melicoccus. Retention of 

these genera within the Melicocceae is not strongly 

The tribe Melicocceae as originally circumscribed supported by morphological data, nor refuted by wood 

by Radlkofer (1888), was based on a suite of charac- anatomy (Klaassen, 1999). Resolution of this problem


Fig. 18. Scanning Electron Micrographs of Talisia and 

Melicoccus pollen (A-F = type A). A-C. T morii. A. polar 

view. B-C. equatorial views. D. M. espiritosantensis, 

polar view. E-F. M. bijugatus. E. polar view. F. equato- 

rial view. (A-C from Herrera et al. 981; D from Folli 

30; E-F from Smith 1705). 

awaits a comprehensive phylogenetic analysis of the 

genera of Sapindaceae. 

Evaluation of the Melicocceae based on fruit characters 

suggest different alliances. Melicoccus and 

Talisia are undoubtedly closely related as their fruits 

are morphologically similar. However, the remaining 

three genera are not clearly related to them as they have 

completely different kinds of indehiscent fruits. Tristira 

has trigonous-winged, trilocular, woody fruits that 

seem to be derived from capsular ancestors, perhaps 

within the Cupanieae. Tristiropsis has true drupes with 

a trilocular stony layer, clearly different from the core 

of Melicocceae and from any other genus of Sapindaceae. 

Wood anatomy of Castanospora was found by 

Klaassen (1999) to deviate from the remaining members 

of Melicocceae, suggesting a closer alliance with 

the Lepisantheae. On the other hand, Klaassen (1999) 

did not find any wood anatomical distinctions between 

the Asiatic genera Tristira and Tristiropsis, and the 

American genera Talisia and Melicoccus, with the re- 

sult that a close relationship based on these features 

cannot be excluded. However, Klaassen (1999) did 

find similarities between the Melicocceae and the 

Sapindeae (Sapindus Linnaeus) and the Lepisantheae 

(Lepisanthes Blume and Zollingeria Kurz) suggest- 

ing the possibilities of different alliances. Resolution 

of these problems awaits a global generic revision of 

Sapindaceae. 

CLADISTIC ANALYSIS 

MATERIALS AND METHODS 

The phylogenetic relationships among the species 

of Melicoccus and Talisia were analyzed cladistically 

using the computer program PAUP (Phylogenetic 

Analysis Using Parsimony, ver. 4.Ob3a; Swofford, 

1999). Twenty-eight characters were selected for this 

analysis (Table 1). Character state polarities were automatically 

established by PAUP through the method of 

outgroup comparison. 

The selection of a sister group and outgroups proved 

to be difficult. As already pointed out, the remaining 

genera currently included in the Melicocceae were determined 

to be unsuitable as they posses characters that 

greatly differ from those of Melicoccus and Talisia. The 

genus Sapindus was determined as the best possible 

outgroup based on its overall similarity to the in-group 

taxa. Alectryon was chosen as the sister-group, based 

on the unpublished data of Johnson and Chase 

(Klaassen, 1999). Their cladistic analysis, based on 

molecular data, regards Alectryon as closely related to 

Talisia. Examination of morphological characters 

proves to be consistent with their phylogenetic interpretation 

supporting a close relationship of Alectryon 

with Talisia. Melicoccuspetiolulatus and Talisiagranulosa 

were excluded from this analysis because many 

of their characters are not known. 

A discussion of the characters and their polarization 

follows. Numerous characters are considered 

plesiomorphic for Talisia because they are found 

throughout the sister-group (Alectryon) and the 

outgroup (Sapindus). All characters were treated as 

unordered, without application of weight. Characters 

present in the outgroups are considered plesiomorphic. 

CHARACTERS 

Quantification of characters was based on extensive 

sampling of herbarium specimens, unless only 

scanty material was available for the analysis. The qualification 

of characters was based on little sampling due 

to the scanty data available.

CLADISTIC ANALYSIS 21 

TABLE 1 

Data matrix for cladistics analysis of Talisia and Melicoccus. 

Sapindus OOOaO 00000 aOaOO OalOO OOOaO 110 

Alectryon 00010 OOaal b2000 OaOOa aOOOO 001 

T. acutifolia 10010 10000 10210 10011 00011 100 

T. bullata OlalO 00000 10210 10011 00010 102 

T. carinata lOalO OaOOO 10201 10011 00011 102 

T. caudata 11010 OaOOO 10200 10011 000?1 102 

T. cerasina 11010 00000 10200 10011 00001 102 

T. croatii llalO 10000 20200 10011 00010 102 

T. cupularis 10010 00000 20200 10011 00011 102 

T. dasyclada 11010 OaOOO 10200 10011 00001 102 

T. douradensis 00010 00000 10200 10011 000?1 102 

T. equatoriensis 00010 ?0000 10200 11001 000?0 102 

T. eximia 11010 00000 10200 10011 00011 102 

T. ghilleana lOalO 00000 10210 10011 00011 102 

T. guianensis 10010 aOOOO 10201 10011 00011 102 

T. hemidasya 01010 00000 10200 10010 00001 102 

T. hexaphylla OlalO OaO20 10200 11011 00000 102 

T. japurensis OOalO 00000 00200 11000 00000 102 

T. laevigata 00010 00000 10200 11011 0000? 102 

T. Iongifolia 10010 00000 10201 10011 00011 102 

T. macrophylla 10010 00000 10201 10011 00001 102 

T. marleneana 10110 00000 10200 10011 00011 102 

T. megaphylla lOalO aOOOO 20200 11011 00001 102 

T. mollis 11010 00000 10200 10011 00011 102 

T. morii 01011 aOOOO 10200 10011 000?0 102 

T. nervosa 10010 00000 10201 10011 0001? 102 

T. obovata 10010 00000 10201 10011 00001 102 

T. oedipoda ?0010 OOaOO 00200 11011 000?1 102 

T.pachycarpa 10010 00000 20200 10011 0000? 102 

M. petiolulatus 10010 ?0000 ?0??? 1?0?? 000?? 102 

T. pilosula 01010 aOaOO 10201 10011 00011 102 

T. pinnata llalO 10000 10200 10011 00010 102 

T. princeps 11010 10000 10200 10011 00010 102 

T. retusa OOalO 00000 10200 10011 00001 102 

T. setigera 10010 aOaOO 10210 10011 00000 102 

T. stricta 10010 10000 20210 10011 00010 102 

T. sylvatica lOalO aaOOO 10211 10011 00010 102 

T. velutina ?Oala 01020 10210 11001 000?0 102 

T. angustifolia 20alO 00000 00210 11001 00000 102 

T. chartacea 00010 00000 00200 11001 000?0 102 

T. clathrata OOall OaO20 00200 11001 00000 102 

T. coriacea OOalO OOaOO 00200 11001 00010 102 

T. esculenta OOalO 00010 00200 11001 00010 102 

T. firma OOalO OOaOO 00200 11000 00010 102 

T. floresii 00010 00000 10200 11000 00000 102


T. furfuracea 01010 00000 10200 11001 00000 102 

T. granulosa 00010 ?0??0 ?0??? 1?0?? 0000? 102 

T. lanata 00110 00020 00200 10000 0001? 102 

T. microphylla OOalO 00000 00200 10000 00010 102 

T. parviflora 00010 00000 00200 11001 000?0 102 

T. praealta 00010 OOalO 00200 11001 00000 102 

T. simaboides OOalO 00000 00200 11001 00010 102 

T. squarrosa 00020 00000 00210 11001 00000 102 

T.subalbens 10010 00000 10200 11001 00010 102 

T. veraluciana OGalO 00000 00200 11001 00010 102 

M. antioquensis 00010 Oa120 OOaOO 00011 000?? 102 

M. ayma rdii 00010 001?0 00100 01000 0002? 102 

M. espiritosan tensis OQalO 00000 00100 01001 00020 102 

M. novogranatensis 00010 Oa120 OOaOO 00001 000?? 102 

M. oliviformis 00010 OOa20 00110 0100? 00020 102 

M. pedicellaris 01010 OOalO 00100 01001 00020 102 

M. bijugatus OOOaO 00101 01000 01100 11130 102 

M. Iepidopetalus 00020 00111 lalO 01000 1103? 102 

M. jimenezii 00010 00111 01000 01000 1113? 102 

Polymorphic characters represented by a (=O& 1) and b (=1&2). 

1. Habit. Trees (0); treelets (1); shrubs (2). Although 14. Disc shape. Annular (0); cup-shaped (1). 

treelets and shrubs are widespread throughout the 15. Stamen number. Eight (0); five or predominantly 

Sapindaceae, they seem to be derived within the less than eight (1); more than eight (2). 

Melicocceae. In general, architectural models in the 16. Stamen orientation. Spreading (0); erect (1). 

Melicocceae are imperfectly known because the un- 17. Filaments. Equal lengths (0); unequal lengths (1). 

derlying developmental data are not available. At 18. Anther attachment. Basifixed (0); dorsifixed (1). 

present, the best estimate of this character is inferred 19. Anther outline. Elliptic or ovate (0); oblong, linear 

from specimen data or field observations. 

2. Glandular pubescence. Absent (0); present (1). 

or lanceolate (1). 

20. Anther apex. Obtuse or retuse (0); apiculate (1). 

3. Leaves. Paripinnate (0); imparipinnate (1). 21. Number of carpels. Three (0); two (1); more than 

4. Leaf rachis. Winged (0); unwinged (1); absent (2). three (2). 

5. Tertiary venation. Reticulate (0); clathrate (1). Subclath- 22. Gynoecium locules. With as many locules as carpels 

rate tertiary venation was sometimes found in a few (0); unilocular (1). 

species, and because it is intermediate between the 23. Style. Elongated (0); short to absent (1). 

two stages (reticulate and clathrate) it was codified 24. Stigma. Elongated (0); capitate (1); trilobed (2); 

as having both stages. 

discoid-bilobed (3). 

6. Cataphylls. Simple (0); pinnatifid (1). 

25. Pollen grains (sensu Muller & Leenhouts, 1976). 

7. Inflorescence position. Axillary or terminal (0); Type A (0); type Al (1). The characterization of 

Cauliflorous or ramiflorous (1). 

pollen grains occurring in each species is largely 

8. Flower clusters. Dichasia (0); solitary (1). 

based on one sample. However, the consistency of 

9. Inflorescence shape. Paniculate (0); racemose (1); 

fasciculate (2). 

10. Pedicel. Articulate (0); non-articulate (1). 

11. Sepals. Free (0); connate at base (1); connate to half 

pollen types within a species was tested by expanding 

the sample size to two or three specimens in the 

following species: M. oliviformis, T angustifolia, T 

firma, T ghilleana, T hexaphylla subsp. hexaphylla, 

way (2). 

12. Sepal number. Five (0); four (1); more than five 

T. japurensis, T. longifolia, T. macrophylla, T. 

simaboides, T. squarrosa, and T. subalbens. 

(2). 

26. Fruit. Dehiscent (0); indehiscent (1). 

13. Petals. Without appendages (0); with basal or mar- 27. Number of carpels per fruit. Polycarpic (0); monoginal 

appendages (1); with well developed adaxial 

appendage (2). 

carpic (1). 

28. Sarcotesta. Absent (0); partial (1); and complete (2).

ECOLOGY AND DISTRIBUTION 23 

RESULTS 

and T hexaphylla. Although, there are a few recogniz- 

A heuristic, most parsimonious analysis was ini- able subgroups within subgenus Talisia, it seems more 

tially run, using Sapindus and Alectryon as outgroups. appropriate to treat the species alphabetically because 

The resulting cladograms had Alectryon nested within there is no resolution for most of them. 

Melicoccus, rendering the in-group polyphyletic. A 

second analysis was done, using Alectryon as the only ECOLOGY AND DISTRIBUTION 

outgroup, with a maximum number of trees retained Talisia is essentially an Amazonian genus (sensu 

limited to 16,500. The shortest cladograms produced Prance, 1977) which has a distribution that extends to 

had 133 steps. A majority rule consensus tree was gen- the north through Central America to southern Mexico 

erated producing the topology shown in Fig. l9and one 

(south of the Isthmus of Tehuantepec) and to the south 

of the cladograms is reproduced in Fig. 20. There are 

to southeastern Brazil and Paraguay (Fig. 21). Talisia 

several branches that are strongly supported as they are 

esculenta (Saint- Hilaire) Radlkofer, is the exception, 

present in at least 90 % of the trees generated in this 

and may be found outside this range through cultivaanalysis. 

This branching pattem is not in contradiction 

tion. Species of Talisia occur from sea level to 1750 m 

with my previous notions and characterization of two 

elevation, however, most of the species are found within 

subgenera within Talisia. Accordingly, the subgenus 

the 200-800 m range. They are predominantly found 

Talisia is monophyletic while subgenus Pitombaria is 

in moist, non-flooded, primary forests on various types 

paraphyletic. Although there are numerous competing 

of soils, and they are either treelets that inhabit the uncladograms, 

I feel that the consensus tree presents some 

derstory, or medium to large-sized trees that reach the 

level of useful structuring at the subgeneric level. From 

canopy. A small number of species are found in the 

this tree, it becomes clear that relationships at the speseasonally 

flooded varzea or igapo forests, in dry forcies 

level, with a few exceptions, are unresolved. Alests 

or in shrubby savannas (cerrado vegetation) and 

though species have enough autoapomorphic characin 

forest islands surrounded by savanna. A few speters 

to define them, there are not enough synapomorphic 

cies are able to thrive in undisturbed 

characters to resolve their 

forests as well as 

relationships at the species 

level. The 

in 

relationships of species within the subgensecondary 

forests. Individuals of Talisia are not 

era is left unresolved until more characters (preferably 

uncommon, and typically can be found in almost any 

molecular) becomes available. 

tropical lowland undisturbed forested region of the 

Subgenus Talisia, being the most derived of the South American continent. 

groups, is recognized by numerous apomorphies. Sub- In an attempt to recognize the center(s) of distribugenus 

Pitombaria is basal to subgenus Talisia and tion for the genus, the distribution of its taxa were plotcontains 

numerous plesiomorphic characters, although ted against the phytogeographic regions recognized by 

it is morphologically recognizable. 

Prance (1977) for the Amazon area (Fig. 22). From the 

The analysis also showed Melicoccus to group with results, it is apparent that distribution of taxa (except 

Talisia oliviformis and T. pedicellaris and with 5 other for the endemics) is not limited to particular regions, 

species that are described in this work. This clade has but instead occupy many of them. This information 

numerous apomorphic characters that clearly distinguish could ultimately serve to re-evaluate Prance's phytoit 

from the remaining taxa. However, the recognition geographic regions and to postulate similarities among 

of this monophyletic clade expands the traditional them. However, performing an analysis of this type 

circunscription of Melicoccus to include trilocular (vs. would be beyond the scope of the present work as it 

unilocular) ovaries. 

would require a multi-taxa approach. From this exer- 

Treatment of species within the three groups herein cise, various centers of distribution become apparent. 

recognized, follows the order of species as suggested The main center is located in the Guianas-Amapa' rein 

the analysis only when their phylogenetic relation- gion [the Atlantic coastal region (1) of Prance, 1977] 

ships are fully resolved. The relationships of Melicoccus with 26 taxa ( 23 species and 2 subspecies), nine of 

are fully resolved and, therefore, the species are ordered which are endemics. A second center is located in southfrom 

more plesiomorphic to the most apomorphic. The western Amazonas (region 7) with 22 taxa (19 species 

relationships in Talisia subgenus Pitombaria, as and 3 subspecies) including four endemics, and a third 

shown in the consensus tree, are weakly supported and in central-south Amazonas region (Prance's Roraimatherefore 

unresolved. Although the species in this sub- Manaus, 4b) with 19 taxa (18 species and 1 subspegenus 

have numerous autoapomorphies, there are not cies), three of which are endemics. The remaining five 

enough synapomorphic characters to resolve their phy- regions contain 8 to 12 taxa each. 

logenetic relationships. Talisia subgenus Talisia can be The two subgenera of Talisia are widely dispersed, 

expanded to include two peripheral species, T laevigata more widely so in subgenus Pitombaria with species


lr* ? 

Fig. 19. Majority rule consensus tree. indicates strongly supported clades, Alectryon= 

outgroupu Melcoccus; 

Talisia.


2~~~~~~~~: 

W.ih 

Wmm 

IFI 

Fig. 20. Selected cladogram, showing Melicoccus and Talisia as sister groups; Talisa is further divided into 

subgenera, Pitombaria and Talisia.


l 

f.,~~~~~~~~~~~~~~~1 

a_, 


I IO9 W ! OD S W ! ; Ke,> 

7 0W ! 

SA A 

6 

OW ! 

s o W 1 4 

0 

W ! 

TA~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~ 

ISI tt 

TALISIA 

* endemic species 

A widely distributed species 

/ig. 2ibnTaeeA 

SAL~~~~~~~~~~~ 

A A A~ AL 

AL 

VLLL(A 

4. 

K ~~AU 

? 

s/JAL 

Fig. 23. Distribution of Talisia endemics vs. widely distributed species. 

occuring in the Amazon and beyond the Amazon Fruits of T megaphylla are reported by Roosmalen 

(Figs. 41, 45, 50, 54, & 62). Subgenus Talisia is pre- (1985) as being eaten and dispersed by capuchin mondominately 

Amazonian but extends into northern South keys, T microphylla by howler and tamarin monkeys, 

America and as far north as Nicaragua in Central and T sylvatica by spider and howler monkeys. Fruits 

America, and to the south into southeastern Brazil (Figs. of T. esculenta, according to Lorenzi (1992), are eaten 

67, 72, 76, 80, 84, 89, 93, 98, 103, 107). Its distribution by birds. 

is narrower than subgenus Pitombaria and does not ex- Melicoccus, on the other hand, contains ten species 

tend into the southern portions of North America, i.e., of trees with narrower ranges, naturally distributed in 

southern Mexico. Talisia subgenus Talisia evidently dry vegetation such as coastal forests, fields with sandy 

originated in South America and later expanded into soils or less often in periodically flooded forests. Most 

Central America. This is a likely explanation deduced species of Melicoccus occurs naturally in SouthAmerica 

from its modern distribution, where most of the spe- around the periphery of the Amazon area (Fig. 29, 36), 

cies are located in South America and only a few, mostly with the exception of M. oliviformis and the endemic 

widespread taxa, extend to Central America. M jimenezii and M. lepidopetalus which occur outside 

Endemism in Talisia is widely dispersed (Fig. 23). of this range. Melicoccus oliviformis is found in the 

Although most endemic species are found in the Ama- Yucatan Peninsula in seemingly natural conditions inzon 

region, there are a few endemics outside this region. dicating a possible disjunction with northern South 

These include one species in theYucatan peninsula, and America. However, it is not clear whether this distriseveral 

in Panama-Choc6-Ecuador, and a few in the butional pattern is natural or is the result of ancient in- 

Planalto and southeastern regions of Brazil. 

troductions by the people that inhabited the area. 

1 ON


Melicoccusjimenezii being endemic to the coastal dry are often sold under the name of pitomba or pitomba 

forest of southeastern Hispaniola and M. lepidopetalus da mata. A few species of Talisia (T esculenta, T 

endemic to the cerrado-chaco vegetation of south-cen- furfuracea, T hexaphylla, T squarrosa, and T stricta) 

tral South America are clearly disjuntfrom the remain- are reported as being used as piscicides in SouthAmerica 

ing species ofMelicoccus. Melicoccusjimenezii is the (Acevedo-Rodriguez, 1990). Talisia squarrosa is reonly 

naturally occurring member of the tribe in the ported as one of the major timber tree species of Guyana 

GreaterAntilles. Its presence in the Dominican Repub- (Polak, 1992) and T esculenta as a minor timber tree 

lic is clearly a relict population as the genus has been 

recorded from macrofossils for the middle Oligocene 

for interior construction work. 

in Puerto Rico (Graham, 1996). The current distribu- TAXONOMIC TREATMENT 

tion ofM bijugatus as a widespread coastal or dry forest 

Key to the genera of American Melicocceae 

species in Central America, the West Indies (Fig. 36), 

and areas of the Paleotropics is only the result of culti- 1. Petals with 2 marginal projections or appendages, 

on or above the base, or lacking appendages; 

vation for the last 250 years. 

stamens spreading, broadly exposed ....... Melicoccus 

CONSERVATION STATUS 

1. Petals with a well-developed, single, adaxial, 

petaloid appendage; stamens mostly erect, 

A number of Melicoccus and Talisia species are 

known only from the type or from less than three colpartially 

exposed (hidden by the petal's 

appendages) ............................ Talisia 

lections, indicating that they may possibly be uncommon 

or rare species. The following species are in this MELICOCCUS P. BROWNE 

category: Melicoccus antioquensis, Melicoccus MELICOCCUS P. Browne, Civ. Nat. Hist. Jaaymardii, 

Melicoccus espiritosantensis, Melicoccus maica. 210. 1756. Type. Melicoccus bijugatus Jacquin. 

petiolulatus, Talisia douradensis, Talisia granulosa, The name is derived from the Latin mel (= honey), and 

Talisia lanata, Talisia laevigata, Talisia parviflora, coccus (= berry), in reference to the flavor of the fruit. 

Talisia oedipoda, and Talisia pilosula. However, this 

may be the result of collection artifacts, since large areas 

in the Neotropics have not been thoroughly collected. 

The Dominican Republic endemic, Melicoccus 

jimenezii, is an extremely rare species restricted to a 

small population on the southeastern side of Hispaniola 

(Garcia & Mejia, 1996; Acevedo-Rodriguez, 1997). 

Melicocca Linnaeus, Sp. PI. ed. 2, 1: 495. 1762, 

nom. illeg. (illegit. typographical variant). 

Casimira Scopoli, Intr. Hist. Nat. 234. 1777, nom. 

illeg. (nomenclatural synonym). 

Talisia sect. Cotopais Radlkofer, Sitzungsber. Math. 

Phys. Cl. Konigl. Akad. Wiss. Miunchen 8: 342. 

1878. Type: Melicoccus oliviformis Kunth. 

This species may become endangered as the area where 

it is found is subjected to pressures imposed by un- Small trees. Leaves alternate, pari-pinnately comregulated 

tourism. 

pound, distal leaflet rudimentary or wanting; leaflets 2, 

4, or 6, opposite, or less often alternate; rachis winged 

USES 

or nearly cylindric; stipules wanting. Inflorescence of 

terminal or axillary panicle-shaped thyrses, panicles or 

Melicoccus bijugatus, M. lepidopetalus, and M. racemes. Flowers 4- or 5-merous, actinomorphic, unioliviformis 

are cultivated for their edible fruits sexual (the plants dioecious or monoecious); pedicels 

(sarcotesta). Melicoccus bijugatus and M. oliviformis articulate or not articulate; sepals imbricate, connate at 

have been introduced into Central America, the West base, shorter or as long as the petals; corolla of distinct 

Indies, and the Old World tropics, while M. petals with imbricate aestivation, inserted on the base 

lepidopetalus is only locally cultivated. Fruits of M. of an extrastaminal nectary disc, the petals erect or rebijugatus 

are available during the fruiting season in flexed, elliptic to obovate, without appendages, the 

markets in Colombia, French Guiana, Guyana, New margins sometimes with an elongated projection on each 

York City, Puerto Rico, and Venezuela. An alcoholic side above the base; nectary disc prominent, annular or 

drink called "bili" is made in Vieques Island (Puerto lobed; stamens 8, spreading, filaments of equal or un- 

Rico) by aging rum with the fruits. 

equal length, longer than the petals, glabrous or pubes- 

Most species of Talisia have seeds with an edible cent, the anthers dorsifixed, elliptic or ovate, retuse at 

sarcotesta, which has a slightly sour taste, that is con- apex; ovary 2 or 3 carpellate, with as many locules as 

sumed locally. Many of them are gathered from the for- carpels or unilocular, each carpel with a single campyest 

or cultivated in backyards or communal lands. Talisia lotropous or apotropous ovule, the stigma 2 or 3-lobed 

esculenta has fruits whose quality and quantity allow to subcapitate, sessile. Fruits baccate, indehiscent, elpeople 

to sell them in local markets. In Brazil, its fruits lipsoid, ovoid, or globose, with leathery mesocarp. Seed

TAXONOMIC TREATMENT 29 

1(-2), with a fleshy, edible testa. Embryo fleshy, 

lomatorrhizal with straight erect cotyledons of similar 

size, or notorrhizal, with cotyledons diagonally superimposed, 

the radicle punctiform or elongated. Pollen 

Key to the species of Melicoccus 

colporate, nearly spherical, oblate, isopolar, trangular 

or trigonous with straight to convex sides. 

A genus often species native to South America, 

and Hispaniola. 

1. Inflorescence a thyrse (most of the dichasia simple or compound). 

2. Petals glabrous; disc annular or pentagonal. 

3. Inflorescence panicle-shaped; stems glabrous; fruit smooth. 

4. Leaflets 4.5-10 cm long; petiolules 3-5 mm long ............................................... 1. M. espiritosantensis 

4. Leaflets 14-19 cm long; petiolules 1-2.5 cm long ....................................................... 3. M. petiolulatus 

3. Inflorescence racemiform; stems puberulent; fruit rugulose . .............................. 2. M. pedicellaris 

2. Petals puberulent to sparsely-woolly; disc cup-shaped, 10-lobed ............................................. 4. M. oliviformis 

1. Inflorescence a panicle or a raceme (all dichasia reduced to a single flower). 

5. Pedicels articulate; gynoecium tricarpellate, 3-locular; stigma trilobed. 

6. Petals subulate or lanceolate, entire, without appendage-like margins; filament equal; leaflets 

elliptic to oblanceolate, 7-23.5 cm long. 

7. Stems ferruginous-furfuraceous; leaflets acuminate to abruptly acuminate at apex; petals 

lanceolate, woolly to sparsely woolly on both surfaces; anthers lanceolate ......... 5. M. antioquensis 

7. Stems glabrous; leaflets obtuse at apex; petals subulate, puberulent on both surfaces; 

anthers ovate ...................................................... 6. M. novogranatensis 

6. Petals rhombate, with extended appendage-like margins; filaments unequal; leaflets narrowly 

elliptic, 4-9 cm long ............................................................... 7. M. aymardii 

5. Pedicels not articulate; gynoecium bicarpellate, unilocular, stigma bilobed. 

8. Leaves with 2 leaflets; petals with a pair of minute marginal appendages .................. 8. M lepidopetalus 

8. Leaves with 4 leaflets; petals lacking appendages. 

9. Leaflets acute or acuminate at apex; leaf rachis usually winged or margined; fruit green at 

maturity ...................................................... 9. M bijugatus 

9. Leaflets obtuse or rounded at apex; leaf rachis unwinged, slender; fruits yellow at 

maturity .10. M jimenezii 

1. MELICOCCUS ESPIRITOSANTENSIS 

canaliculate, wider at base. Thyrses panicle-shaped, con- 

Acevedo-Rodriguez, sp. nov. 

gested on distal portion of stem, to 13 cm long, branches 

Type. Brazil. Espirito Santo: Mun. Linhares, Reserva to 5 cm long; cataphylls deltate, minute, at base of inflo- 

Florestal de CVRD, Flamengo Station, tall forest, 17 

rescence; axes angled, sulcate, puberulent; bracts minute, 

Aug 1978 (fl), D.A. Folli 30 (holotype, US; isotype, 

early deciduous, minutely tomentose; dichasia compound 

CVRD, n.v., NY). Figs. 18d, 24a-g 

or simple; peduncle 5-7 mm long; pedicels ca. 1.5 mm 

long, articulatenearthe base. Calyx green, 2-2.5 mm long, 

A M. aymardii Acevedo-Rodriguez inflorescentiis tomentulose, the sepals free to the base, concave, ovate, 

thyrsoideis et filamentis glabris differt. 

adaxially tomentulose, obtuse at apex, margins ciliate; 

petals white, asymmetrically rhombate, ca. 2.5 mm long, 

Tree 18-26 m tall; tunk to 65 cm in diam., with scaly glabrous or with few scattered hairs on abaxial surface, 

bark. Stems terete to obtusely angled, glabrous, striate, the adaxial surface with a few scattered hairs, the apex 

lenticellate. Leaves paripinnate or less often imparipinnate; obtuse, the base cuneate, the margins elongated into a subdistal 

process aciculate, 3-5 mm long; leaflets 2-4(5) filiform projection, densely covered with elongated feropposite 

or altemate, elliptic, oblong-elliptic to nearly lan- ruginous hairs; appendages wanting; disc annular, 5-lobed, 

ceolate, 4.5-10 x 2-3.1 cm, chartaceous, the adaxial sur- glabrous, ca. 1 mmtall; stamens 8, spreading, the filaments 

face glabrous, slightly lustrous with sunken midvein, the of unequal lengths, glabrous, the anthers ca. 1 mm long, 

abaxial surface dull, glabrous except for the puberulously elliptic-ovate, glabrous, apiculate or retuse at apex; ovary 

prominent midvein, the venation brochidodromus, adaxi- nearly conical, with a few scattered hairs, the stigma trially 

inconspicuous, abaxially prominent, tertiary veins lobed, tomentulose, papillate. Fruits ellipsoid, to 2.2 cm 

reticulate, the margins entire or slightly undulate, slightly long, glabrous, smooth, the pericarp chartaceous, ca 1 mm 

revolute, the apex acuminate, glandular-mucronulate, the thick, the endocarp glabrous. Seed one per fruit, ellipsoid. 

base asymmetrical, acute to obtuse; petiolules slender, 3- Embryo with cotyledons superimposed, of equal size. 

5 mm long, puberulent; rachis 2-3 cm long, trullate, pu- The specific epithet refers to the Brazilian state where 

berulent; petioles 1.2-5.4 cm long, slightly flattened, the type was collected.


mm3 

2mm[ .~~~~3mm 

Fig. 24. Melicoccus espiritosantensis. A. flowering branch. B. simple dichasium. C. abaxial and adaxial views of 

petal with marginal projections. D. staminate flower with perianth removed showing disc, stamens, and detail of stamen 

(left) and I.s. same (right). E. leaf. F. pistillate flower. G. pistillate flower with perianth removed showing disc, sterile 

stamens, and pistil (right), and l.s. same with detail of sterile stamen (right). (A-D from Folli 30; E-G from Silva 255). 

Phenology. Collected in flower during August and 

in fruit during October. 

Distribution and Ecology. (Fig. 29) Know only 

from eastern Brazil. 

Representative specimens examined. BRAZIL. 

Bahia: Mun. Prado, forest reserve of Brazil de Holanda 

Industrias, entrance at km 18 E of Itamaraju on road to 

Prado, 22 Oct 1993 (fr), Thomas et al. 10087 (US). 

Espirito Santo: Mun. Linhares. Reserva Florestal da 

CVRD, Station Oiticica, before km 151, tall forest, 18 

Aug 1985 (fl), Silva 255 (US). 

Melicoccus espiritosantensis is closely related to 

M aymardii as they share numerous morphological features. 

Melicoccus espiritosantensis can be distinguished 

from M. aymardii by its widely rhombate petals, with 

two elongated (nearly as long as the petal) marginal appendages 

(vs. narrowly rhombate petals with very short 

marginal appendages); and by its glabrous filaments (vs. 

filament pilose or woolly on lower half). 

2. MELICOCCUS PEDICELLARIS (Sagot ex 

Radlkofer) Acevedo- Rodriguez, comb. nov. Talisia


cm j ~~~~~~~~~~~ 

fl 0 eG ] ~~~~~~~Imm{ 

Fig. 25. Melicoccus pedicellaris. A. flowering branch. B. staminate flower. C. adaxial, abaxial, and lateral views 

of petal with marginal projections. D. staminate flower with perianth removed showing disc and stamens (right), 

l.s. same (left). E. pistillode. F. infructescence. G. embryos, lateral and dorsal views. (A-E from Mori et al. 19144; 

F-G from Kuhlman & Jimbo 349). 

pedicellaris Sagot ex Radlk., Sitzungsber. Math.- 

Phys. Cl. Konigl. Bayer Akad. Wiss. Miinchen 8: 

342. 1878. Type. French Guiana. Acarouany, 1858 

(fl), Sagot 1188 (lectotype, P, here designated; iso- 

lectotypes, K-2, P-7; frag. at M). Syntype. French 

Guiana. Maroni river, without date (fr), Melinon 

s.n. (M, P). Fig. 25a-g 

. 

Talisia pulverulenta Radlkofer, Sitzungsber. Math.- 

Phys. Cl. KonigI. Bayer Akad. Wiss. Miinchen 8: 

342. 1878. Type. French Guiana. Without spe- 

cific locality, 1864 (fr), Melinon s.n. (holotype, 

B-destroyed, photo (F neg. 5681) at MO, NY, 

US); lectotype, M, here designated; isotypes, NY, 

P-3, US-2).


Tree 12-14 m tall, with stout lateral branches on upper est, including transition rain forest-savanna, forest on 

one-third of trunk; trunk reaching 25 cm in diam.; but- clay, primary forest, disturbed forest, trailside, riverine 

tressed at base; bark brown or reddish brown, rough, forest on clayish soil, and virgin terra firme forest. 

thin, peeling off in irregular plates; inner bark reddish 

brown. Branches terete, puberulent, becoming 

Common name. Surinam: Zwarte pintolokus. 

lenticellate. Leaves paripinnate; distal process acicular, Representative specimens examined. SURINAM. 

ca. 3 mm long, tardily deciduous leaving a truncate base Commewijne: Brokopondo, 8 Oct 1966 (fl), van 

ca. 0.6 mm long; leaflets 4 or 6, opposite or altemate, Donselaar 3762 (MO, P, VEN); Jodensavanne-Mapan, 

ovate, widely oblong, elliptic, or widely elliptic, charta- 5 May 1953 (fl), Lindeman 3828 (US), 7 Oct 1953 (fr), 

ceous, 4-14.5 x 1.8-6.6 cm, adaxially glabrous or some- Lindeman 4886 (K, US). Saramacca: Nassau, 8 Mar 

times the midvein ferruginous, appressed-pubescent, 1949 (st), Lanjouw & Lindeman 2523 (NY). 

abaxially glabrous to densely puberulent, very often with 

FRENCH GUIANA. Without specific locality, 1863 

(fl), Melinon s.n. (P). Bassin de L'Approuague: Saut 

midvein and secondaries ferruginous-hirsutulous, the 

Parare, 10 Sep 1983 (st), Barrier 4263 (CAY); Station 

venation brochidodromus, tertiary veins reticulate, the 

des Nouragues, 20 Jul 1989 (st), Sabatier & Prevost 2816 

apex long-acuminate to caudate, the base obtuse, acute, (CAY), 9 Aug 1991 (fl), Sabatier 3510 (CAY, NY, P, U, 

to nearly rounded, sometimes slightly inequilateral; peti- US). Bassin du Maroni: Maroni river, without specific 

olules cylindrical, 2-5 mm long, glabrous, puberulent locality or date (fl), Wachenheim 66 (K, P-4). Piste de 

or pubescent, less often with scattered ferruginous erect St. Elie: Station Biologique ORSTOM, 29 Apr 1992 (st), 

hairs; rachis 1.2-5.4 cm long, terete or nearly so, with Acevedo R. 4868 (CAY, US), 12 May 1992 (st), Acevedo 

same indumentum as the petiolules; petioles 2-11 cm R. 4939 (CAY); between Sinnamary & Counama rivlong, 

adaxially slightly flattened, with same indumen- ers, 5 Jun 1991 (st), Sabatier & Prevost 3601 (CAY). 

tum as the rachis, slightly enlarged at base. Thyrses Region de Saul: Mont Galbao trail, 250 m, 24 Aug 1988 

(fl), 

racemiform, solitary and supra-axillary or terminal and 

Mori et al. 19144 (CAY, MG, NY, P, US). 

BRAZIL. Amaph: Serra da Arumanduba, 150 m, 25 

congested, 4-8 cm long; cataphylls clustered, acicular, 

Jul 1961 (yfr), Egler & Irwin 45995 (IAN, MG, US); 

1.5-2 mm long; axes sulcate, puberulent, sometimes with 

Araguari river, 135 m, 28 Aug 1961 (yfr), M. Pires et 

glandular papillae; bracts and bracteoles deltate or subu- al. 50549 (IAN, MG, NY, US, VEN). Amazonas: Mun. 

late, < 0.6 mm long; dichasia simple or the flowers soli- Itapiranga. Uatumi river, 28 Aug 1979 (fr), C. Ferreira 

tary (due to aborted lateral flowers), especially toward et al. 822 (INPA, MG, NY). Park: Altamira, 19 Aug 

distal portion of inflorescence; peduncle slightly flattened, 1978, (fr), Bahia 91 (MG, NY), 16 Aug 1978 (fr), Ba- 

1-2 mm long, with same indumentum as the axes; hia 61 (MG, NY); Belem-Brasilia km 93, 16 Oct 1959 

pedicels 0.5-1 mm long, articulate at base. Calyx pale (fr), Kuhlmann & Jimbo 349 (INPA, K, MG, US); Belemgreen, 

2-2.5 mm long, puberulent, margins ciliate, the Brasilia: hwy., 7 Aug 1963 (fr), Maguire et al. 56058 

sepals unequal, free to the base or nearly so, oblong to (K, MO, NY, P, US); Santarem, 10 Sep 1969 (fr), Silva 

ovate; petals white, rhombate, 2-2.5 mm long, glabrous, & Souza 2535 (MG, MO, NY, P, US), 10 Sep 1969 (fr), 

Silva & Souza 2540 

the margins ciliate, extended, inrolled (appendage-like) 

(K, MG, MO, NY, US). 

at base, the apex obtuse, the base cuneate; appendages 

wanting; disc annular, unlobed or slightly lobed, gla- 

3. MELICOCCUS PETIOLULATUS Acevedobrous, 

ca. 0.5 mm tall; stamens 8, spreading, the filaments 

of unequal length, 2-3 mm 

Rodriguez, sp. nov. 

long, sparsely woollypubescent 

to nearly glabrous, the anthers wide-elliptic, Type. Peru, Loreto: Puerto Arturo, lower Huallaga 

ca. 0.5 mm long, shortly apiculate ornearly retuse; ovary river, below Yurimaguas, ca. 135 m, dense forest, 24conical, 

glabrous, the stigma trilobed, sometimes 25 Aug 1929 (fr), Killip, E.P & A.C. Smith 27762 (hosparsely 

pubescent. Fruit ellipsoid, apiculate, 1.8-2 cm lotype, US). Fig. 26a-c 

long, glabrous, the pericarp coriaceous, granulose, ca. 

0.7 mm thick; endocarp glabrous. Seed 1(2) per fruit, A M. pedicellari (Radlkofer) Acevedo- Rodriguez 

ellipsoid, with a thin fleshy testa. Embryo with cotylefoliolis 

nervatibus adaxialiter planis, sepalis caducis, et 

dons of similar size, lying transversely over each other. 

thyrsis paniculiformibus differt. 

Melicoccus pedicellaris is easily distinguished by Shrub 5-6 m tall. Stems terete, smooth, glabrous, 

its cylindrical, elongated petiolules. The specific epi- grayish, lustrous. Leaves paripinnate; distal process early 

thet seems to refer to the conspicuous pedicels. deciduous; leaflets 4, opposite, elliptic, 14-19 x 6.5-8.3 

cm, chartaceous, glabrous on both surfaces, the venation 

Phenology. Flowers from May to October and fruits 

brochidodromus, midvein prominent especially on abaxial 

in August to January. 

surface, secondary veins adaxially plane, abaxially promi- 

Distribution and Ecology. (Fig. 29) Surinam, nent, tertiary venation reticulate, the margins slightly 

French Guiana, and Brazil in non-flooded moist for- undulate, the apex long-acuminate, the acumen sometimes


X~~~~~~~~~~~~~~~~~~~~~ Icm.. 

Fig. 26. Melicoccus petiolatus. A. fruiting branch. B. detail of infructescence showing fruit. C. embryo, dorsal 

view. (A from Killip & Smith 27762; B-C from Williams 4272).


obtuse, the base obtuse, slightly to strongly unequal; 

petiolules slender, 1-2.5 cm, glabrous; rachis ca. 6 cm 

long, slender, terete, glabrous; petioles 8-11.5 cm long, 

terete, glabrous, pulvinate at base. Thyrses panicleshaped, 

terminal; cataphylls minute, acicular; axes to 23 

cm long, slightly angled, puberulent; dichasia seemingly 

simple. Flowers unknown. Fruit ellipsoid, glabrous, ca. 

2 cm long (immature), smooth. 

Melicocuspetiolulatus seems to be most closely related 

to M. pedicellaris Radlkofer in that both possess 

similar vegetative morphology. However, M. petiolulatusa 

differs from M. pedicellaris by having leaflets 

with adaxially plane venation and early deciduous sepals 

(vs. impressed venation and sepals persistent at the 

base of fruits). The specific epithet refers to the relatively 

elongated petiolules that characterize this species. 

4. MELICOCCUS OLIVIFORMIS Kunth in Humboldt, 

Bonpland & Kunth [as Melicocca 

olivaeformis], Nov. Gen. Sp. 5: 100. 1821 Talisia 

oliviformis (Kunth) Radlk., Sitzungsber. Math.-Phys. 

Cl. Konigl. Bayer Akad. Wiss. Miinchen 8: 342. 

1878. Type. Colombia. Bolivar: vic. of Turbaco, 360 

m, without date (fr), HBK 1490 (holotype, P-HBK, 

photo (F neg. 36020) at MO, US; isotypes, P-2). 

Diclinous tree, 5-35 m tall, forming a dense crown; 

trunk reaching 20 to 50 (100) cm in diam. Stems sulcate, 

glabrous, puberulent or pubescent, lenticellate, ashcolored. 

Leaves paripinnate; leaflets 2 or 4, opposite, 

subopposite or less often altemate, 2.7-13.5(19.5) x 

1.3-5.3(9.3) cm; distal process acicular, ca. 5 mm long, 

early deciduous, leaving a 2-3 mm long base with truncate 

apex; leaflets glabrous, the adaxial surface dull, 

lighter than the abaxial surface, with secondary veins 

plane or slightly sunken, the abaxial surface with prominent 

primary and secondary veins, the venation brochidodromus, 

with secondary veins alternate or subopposite, 

arching toward the margin, tertiary veins 

reticulate, the margins slightly undulate, the apex obtuse, 

rounded or less often obtusely apiculate acute or 

acuminate, the base obtuse, cuneate or rounded, slightly 

asymmetrical; petiolules 2-12 mm long, nearly cylindrical, 

wrinkled when dry; rachis (0.5)1.5-4 cm long, 

striate, nearly terete, glabrous, lenticellate; petioles (0.8) 

1-5 (10.5) cm long, striate, glabrous, lenticellate, enlarged 

at the very base. Thyrses fasciculate, axillary or 

distal, 2-20 cm long; cataphylls deltate-acicular, 3-5 

mm long, pubescent or puberulent, on each leaf axil; 

axes angled or slightly flattened; bracts and bracteoles 

deciduous, 1-1.5 mm long; dichasia simple or compound, 

sometimes with aborted lateral flowers; peduncle 

0-4 mm long; pedicels 1-1.5 mm long, pubescent, 

articulate at base or up to the upper third. Calyx 

2-2.7 mm long, ferruginous-tomentose ortomentulose, 

the sepals free to the base or nearly so, concave; petals 

rhombate, 2.2-4.5 mm long, obtuse at apex, cuneate at 

base, erect at anthesis, abaxially scattered woolly-pubescent, 

especially along margins, adaxially puberulent 

to scattered woolly-pubescent; appendages a short prolongation 

of the margins and densely woolly-pubescent, 

or absent; disc cup-shaped; stamens (7) 8, spreading, 

the filaments pilose to woolly on the lower '/2 to 2/3, in 

two sets of unequal length, the shorter ones 2.5-3.5 

mm long, the longer ones 3-4.3 mm long, the anthers 

Phenology. Collected in fruit in late August. ovate or ellipsoid, ca. 1 mm long, glabrous, obtuse at 

apex; ovary densely and minutely whitish or ferrugi- 

Distribution and Ecology. (Fig. 29) Known only 

nous-tomentose, the stigma trilobed, papillate. Fruits 

from the Department of Loreto, Peru. 

ellipsoid to ovoid, 2-3 cm long, obtuse at apex, with short 

Representative specimens examined. PERU. apiculum, the pericarp ca. 0.5 mm thick. Seed ellipsoid, 

Loreto: Fortaleza to Yurimaguas, 29 Oct 1929 (st), Wil- 1.8-2 cm long. Embryo with cotyledons superimposed, 

liams 4272 (F). 

the upper cotyledon slightly larger than the lower one. 

The specific epithet refers to the olive-shaped 

fruits of this species. 

Key to the subsp. of M. oliviformis 

1. Mature fruits brownish yellow, ferruginoustomentose; 

seed with orange or yellow 

sarcotesta ............ M oliviformis subsp. oliviformis 

1. Mature fruits green, grayish or ferruginoussericeous; 

seed with white sarcotesta 

............................ M oliviformis subsp. intermedius 

4a. MELICOCCUS OLIVIFORMIS subsp. 

OLIVIFORMIS Fig. 27a-g 

Toulicia brachyphylla Radlkofer, Sitzungsber. Math.- 

Phys. Cl. K6nigl. Bayer Akad. Wiss. Miinchen 

20: 232. 1890. Type. Venezuela. Carabobo: Puerto 

Cabello, May 1874 (fl), Kuntze 1737 (holotype, 

NY; isotype US). 

Cupania congestiflora Cuatrecasas, Rev. Acad. 

Colomb. Cienc. 8: 475. 1952. Type. Colombia. 

Magdalena: Santa Marta, Jul 1903 (fl), Smith 

1695 (holotype, F; isotypes, K, MO, NY, US-2). 

Bark smooth to rough, grayish. Leaflets 2-4, 2.7-9 

(19.5) x 1.3-5.3(9.3) cm elliptic, ovate, lanceolate, 

oblanceolate, obovate, or oblong, chartaceous to sub- 

coriaceous; petiolules 2-10 mm long, glabrous to pu- 

bescent. Thyrses racemiform or panicle-shaped; axes 

puberulent; bracts oblong-subulate, puberulent; pe- 

duncle pubescent; pedicels 1-1.5 mm long, pubescent, 

articulate at base or up to the upper third; sepals ovate 

to lanceolate, acute to obtuse at apex; disc 8-1 0-lobed,


F 1 

L~~~~~~~~~~~L 

:4~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~z 

2E. 

2 2mm. 

Fig. 27. Melicoccus oliviformis subsp. oliviformis. A. flowering branch. B. staminate flower. C. i.s. staminate 

flower showing disc, pistillode, and stamens. D. flower with perianth removed showing disc and stamens. E. adaxial 

and lateral views of petal with marginal projections. F. stamen. G. pistillode. (All from Dugand & Jaramillo 4120).


glabrous, orpuberulent, 0.7-0.8 mm tall. Fruits brown- of Playa del Carmen, decidous forest, 28 May 1994 (fr), 

ish-yellow, nearly smooth, densely ferruginous-seri- Duran et al. 2133 (CICY); Ciudad Vallarta, 17 km E 

ceous. Seed with sweet, orange or yellow sarcotesta. Puerto Morelos, 8 May 1983 (fl), Cabrera & de Cabrera 

4652 (MO); 4 km W of Playa del Carmen, 21 Jun 1984 

Phenology. Flowering from January to July, and (fr), Cabrera & de Cabrera 6446 (MO). Veracruz: 

fruiting from March to September. 

Cascada de Texolo, 1000 m, 7 Mar 1977 (fr), Calzada 

3206 (F). Yucatin: Becanchen, on road from Tekax to 

Distribution and Ecology. (Fig. 29) Melicoccus Peto, 16 Jan 1956 (st), Enriquez 356 (US); without speoliviformis 

ssp. oliviformis is found in lowlands of cific locality, 1917-1921 (fl), Gaumer 24137 (MO, NY), 

southern Mexico, Belize, Guatemala, Honduras, El 1917-1921(fl), Gaumer 24189 (MO, US). 

Salvador, Colombia, Venezuela, and Peru. However, it GUATEMALA. Petkn: La Libertad & vicinity, 11 Mar 

seems to be native only to Colombia and Venezuela, 1934 (fl), Aguilar 373 (K, NY); Remate, 31 Mar 1960 

where it has been collected in seemingly natural habi- (fl), Contreras 767 (NY, US); Macanche, 27 May 1966 

(fr), Contreras 5876 

tat. In the southern portions of the Yucatan Peninsula, 

(F, US); Melchor de Mencos road, 

250 m, 19 Mar 1965 (fl), Tun Ortiz 90 (NY, US); Santa 

it has been collected in seemingly natural habitats or in 

Elena, 14 Mar 1970 (fl), Tun Ortiz 761 (F, NY, US). 

late secondary forest, indicating that the species is pos- 

BELIZE. Corozal District: 1931-1932 (fl), Gentle 

sibly native there too. However, the Maya Indians who 402 (F, US). Orange Walk District: Indian Church, 

have occupied the peninsula for many centuries may Mayan ruin, 17 May 1976 (fr), Arnason & Lambert 

have introduced the species there due to its edible fruits. 17065 (MO); Honey Camp, Oct 1929 (fl), Lundell 558 

The species is known as huayum or wuayum in Mayan (NY, US); 22 Oct 1929 (st), Lundell 638 (MO, NY, US); 

language and is very likely the source of origin for the Jun 1931 (fl), Meyer 185 (K). 

Spanish word guaya or huaya. The species has been HONDURAS. Comayagua: Humuya river, 23 Apr 

collected in the state ofYucatan, Mexico but only from 

cultivated individuals and early secondary habitats. It 

is known to occur in dry scrubland and forest on sandy 

soils, forest on limestone substrate, riparian forest on 

clayish soil, dry tropical forest, disturbed moist forest, 

savanna, and primary forest on clayish-sandy soils. This 

1956 (fr), Molina 6774 (F, US). Francisco Morazin: 

Yeguare river, 17 Mar 1951 (fl), Molina 3936 (MO, US); 

vic. of El Zamorano, 17 Mar 1951 (fl), Morton 7090 (US). 

EL SALVADOR. AhuachapAn: Vic. Ahuachapan, 

700-1100 m, 6 Jan 1947 (st), Standley & Padilla 2467 

(MO); 800-1000 m, 2-27 Jan 1922 (st), Standley 19712 

(MO, NY, US). 

subspecies is widely cultivated in urban areas and around PUERTO RICO: San Juan. Jardin Botanico UPR, 

homesteads throughout the Neotropics. 

cultivated, 1 Feb 1985 (st), Liogier 35365 (NY), 13 May 

Common names and uses. Mexico: Guaya, huaya, 

1985 (yfr), Liogier 35626 (NY). 

WINDWARD ISLANDS. St. Vincent: Botanical 

guaya pais, huayuim, wuayum. Belize: guaya, kinap. Garden, cultivated, 21 Apr 1892 (fl), Powell 72 (K). 

Guatemala: guaya. El Salvador: tapaljocote. Trinidad: COLOMBIA. Atlintico: Vic. Juan de Acosta, 3 Feb 

yellow chenip. Colombia: mamon cutupli, juria, 1963 (fl), Dugand 6190 (COL, NY, US); Along road to 

cotopris, mamon de Maria, mamon de mico, mamoncillo 

de monte, catapu, cutupli, cutuplis, mamon 

cotopri. Venezuela: cotoperi, cota prisa; cotopriz. 

Puerto Colombia, 4-6 Dec 1963 (st), Dugand 6570 (US), 

(st), Dugand 6572 (US); Lena Candelaria, 30-50 m, 1 Jan 1941 (fl), Dugand & Jaramillo 2782 (US); Los 

Pendales, Hac. Riodulce, 20-50 m, 21-26 Jan 1946 (fl), 

Representative specimens examined. MEXICO. Dugand & Jaramillo 4120 (COL-2, US); Without spe- 

Campeche: Sucopo, 25 Mar 1956 (fl), Enriquez 514 cific locality, 30-40 m, 10 Mar 1933 (fl), Dugand & 

(US); Izamal, without date (fl), Gaumer 406 (MO, NY- Standley 375 (US); Barranquilla, Jan 1937 (fl), Elias 1508 

2, US); Chichankanab, without date (fl), Gaumer 1749 (US); Puerto Guiraldo, 5-10 m, 29 Apr 1960 (st), Mora 

(MO, US); Kancabconot, Feb 1917 (fl), Gaumer et al. 1450 (COL). Antioquia: Mun. Turbo, Lomas Aisladas, 

23577 (MO, NY, US); Chichankanab Lake, Apr 1917 50 m, 8 Feb 1985 (fr), Renteria et al. 3600 (JAUM). 

(fl), Gaumer et al. 23635 (MO, NY, US); Conhuas, 23 Bolivar: Mun. San Juan Nepomuceno, National Sanctu- 

Feb 1932 (fl), Lundell 1376 (US); Terrenos Brason, vic. ary Los Colorados, 230-250 m, 13 Jan 1988 (st), Gentry 

of Escarcega, 60 m, 14 Mar 1968 (fl), Pennington et al. et al. 60659 (MO), Gentry et al. 60665 (MO); Vic. Carmen 

9553 (K, NY). Chiapas: Tuxtla Gutierrez, 9 Mar 1904 de Bolivar, 19 Jan 1963 (st), Romero C. 10009 (COL). 

(fl), Goldman 747 (US); San Fernando, near Tuxtla Cesar: Manaure-La Paz road, 4 km from La Paz, 200 m, 

Gutierrez, 11 May 1966 (fr), Laughlin 911 (F); Tuxtla, 6 May 1984 (fl), Forero et al. 9976 (COL). La Guajira: 

cultivated, 27 Jan 1952 (fl), Miranda 7341 (US). Plains SE of Riohacha by road to Maicao, 10 m, 30 Jan 

Oaxaca: Chapingo, Campo Experimental Forestal "El 1964 (fl), Dugand 6651 (COL, US); Vic. Villanueva, 30 

Tormento", 80 m, 2 Dec 1967 (st), Pennington & Jan 1965 (fl), Ferreyra 1 (COL); Manure, 5 km W of 

Sarukhdn 9386 (K, NY). Quintana Roo: Isla Cozumel, Manure, 440-460 m, 13 Jan 1988 (fl), Gentry et al. 60709 

1 km NW of Faro de la Punta Celarain, 4 Jun 1986 (fr), (MO); Maicao, Correg. Albania, Campamento Tabaco- 

Cabrera & de Cabrera 11437 (MO); Calica, 7.5 km S Los Remedios, Fca. Los Laureles, 200 m, 4 May 1988


(fl), Roldan et al. 1003 (HUA, US); Road between Alba- ern Watershed Reserve, 21 Mar 1924 (fl), Broadway 

nia and Rancheria river, 120 m, 30 Apr 1988 (fl), Rolddn 5237 (F), 15 May 1931 (fl), Russell 12552 (K). 

et al. 945 (HUA); Aremasain, 25 K from Haucha river, ECUADOR. Sucumbios: Lago Agrio. Reserva 

near Rancheria river, 7 Feb 1963 (st), Saravia 2210 (US); Faunistica Cuyabeno, Laguna Imuya, 230 m, 3 Oct 1991 

Between Barrancas & Annerutau, 7 Mar 1962 (fl), Saravia (fr), Palacios et al. 8059 (NY). 

2447 (COL); Serrania La Macuira, Bosque de Nahi, 500- PERU. Loreto: Prov. Maynas. Alpahuayo. Estaci6n 

1700 m, 11 Apr 1964 (fl), Saravia & Saravia 3547 (US). Experimental del IIAP, 150-180 m, Nov 1990 (st), 

Magdalena: Along road to Fundacion, 10 Mar 1957 (fl), Vdsquez & Jaramillo 14935 (MO), Nov 1990 (st), 

Fernandez P. 5283 (COL); Cienaga, 17 Mar 1957 (fr), 

Fernandez P. 5319 (COL); Neguanje. Parque Natural 

Nacional Tayrona, 100 m, 12 Jun 1993 (st), Gentry & 

Vdsquez & Jaramillo 15000 (MO). 

Ortiz 79889a (US); Papayal, 150 m, 23 Feb 1944 (fl), 4b. MELICOCCUS OLIVIFORMIS Kunth subsp. 

Haught 4012 (K, NY, US); Vic. of Barrancas, 200 m, 12 INTERMEDIUS (Radlkofer) Acevedo-Rodriguez, 

Mar 1944 (fl), Haught 4035 (US); Neguanje. Parque 

Natural Nacional Tayrona, 14 Sep 1976 (fr), Lozano & 

Schnetter 2825 (COL, NY); North side of Sierra Nevada, 

1 Feb 1948 (fl), Romero C. 670 (COL, US); North side of 

Sierra de Santa Marta, nevado de Santo Martes, 1 Feb 

1949 (fl), Romero C. 711 (US); Santa Marta, 76 m, Dec 

1898-1901 (fl), Smith 784 (MO, NY, US), Dec 1898- 

1901 (fl), Smith 785 (MO, NY, US). Sucre: Coloso, Primate 

station, 300 m, 25 Oct 1989 (st), Gentry & Cuadros 

68178 (US). 

comb. nov. Talisia intermedia Radlkofer in Martius, 

Fl. Bras. 13(3): 536. 1900. Type. Brazil. Minas 

Gerais: Serra de Caraca, Sep 1884 (fl), Glaziou 

15447 (lectotype, C, here designated; isolectotypes, 

K, P, B-destroyed, photo of B (F neg. 5675) at NY; 

frag. at M). Syntypes. Brazil. Minas Gerais or 

Espirito Santo: 1829, Wied-Neuwied s.n. (M); Venezuela. 

without specific locality or date (st), Boos 20 

(G, n.v., W). Figs. Ic, 12c, 13b&d, 28a-g 

VENEZUELA. Amazonas: Atabapo. Ocamo river, 

vic. of raudal Arata, 270 m, Jan 1990 (fl), Fernandez 6586 

(NY). Anzoategui: Km 227 on Caracas-Barcelona hwy., 

16 km E of Boca de Uchire, 4 Jul 1975 (st), Gentry & 

Leaflets 4, 5-13.5 x 2-4.7 cm, elliptic, ovate or 

obovate, coriaceous, secondary veins straight or slightly 

arching toward the margin, tertiary veins inconspicu- 

Berry 14829 (MO); Boca de El Tigre, 21 Mar 1940 (fl), ous; petiolules 4-12 mm long, puberulent, glabrescent, 

Pittier 14298 (US); Cantaura, 15 Mar 1949 (fl), Smith 63 adaxially furrowed; rachis bicanaliculate along adaxial 

(US). Aragua: Limon, El Sombrero, 3 Feb 1935 (fl), surface, puberulent, glabrescent; petioles slightly flat- 

Archer 3013 (US); Colonia Tovar, 1856-1857 (fr), 

Fendler 2477 (K); La Trinidad de Maracay, 440 m, Jan- 

Feb 1913 (fl), Pittier 5806 (US); Lim6n, 550 m, Feb 1948 

(fl), Pittier 15727 (US); Lim6n, 450 m, 25 Mar 1938 

(fl), Williams 9977 (US); San Francisco, Villa de Cura, 

500 m, 27 Jan 1940 (fl), Williams 12343 (K, US-2). 

Carabobo: Valencia, cultivated, Feb 1920 (fl), Pittier 

8775 (NY, US). Cojedes: Guari, Tamanaco river, 6 Mar 

tened along adaxial surface. Thyrses panicle-shaped; 

axes sometimes sulcate, ferruginous- tomentose or 

tomentulose; bracts subulate or lanceolate, with same 

indumentum as the axis; pedicels ca. 0.1 mm long. 

Sepals ovate, obtuse at apex; disc cup-shaped, 5-lobed, 

tomentulose, ca. 0.5 mm tall. Fruits ellipsoid to ovoid, 

green at maturity, ferruginous or grayish sericeous. Seed 

1947 (fl), Curran 704 (NY). Distrito Federal: Los with sour, white sarcotesta. 

Cavacas, toward aqueduct, Feb 1959 (fl), Aristeguieta 

3825 (NY, US); Cabo Blanco, Curucuti, 7 Apr 1922 (fl), 

Pittier 10276 (US); Vargas, E of Osma, 50 m, 11 Mar 

Phenology. Flower in January, March, September, 

and October, and fruits in April, March and September. 

1973 (fl), Steyermark et al. 106882 (F, US). Falc6n: 

Talquarto, 6 Apr 1917 (fl), Curran & Haman 723 (K, 

NY, US). Guirico: San Juan de los Morros, 550 m, without 

date (fr), Cardona 2126 (US). Lara: 20 km W of 

Carora toward Maracaibo, 20 Jun 1975 (st), Gentry & 

Distribution and Ecology. (Fig. 29) Lowlands of 

Venezuela, Guyana, and Brazil, in xerophytic areas, 

dense forest, gallery forest, shrubby savanna, forest 

margins, savanna, and forested slopes. 

Puig R. 14246 (MO); San Pablo, Barquisimeto-Carora Common names. Guyana: Tacho, wild genip. Braroad, 

1 Sep 1964 (yfr), Trujillo 6840 (F). Miranda: 12 zil: pitomba amarela,pitomba, pitombeira. Venezuela: 

km E Cupira, 100 m, 4 Mar 1980 (fl), Liesner & Gonzalez 

mamon cutuplis, cotopalo. 

9151 (MO, NY). Nueva Esparta: Isla Margarita, 4 Jul 

1903 (fl), Johnston 299 (US). Zulia: Isla de San Carlos, Representative specimens examined. VENE- 

Sabaneta de Montiel, 8 May 1917 (fl, fr), Curran & ZUELA. Distrito Federal: Topo-Tacagua, Jan 1953 (fl), 

Haman 781 (NY, US-2); Santa Rosa, San Carlos del Zulia, Tamayo 33961 (VEN). Falc6n: Silva, Cerro Chichiriviche, 

2 May 1968 (fl), L6pez P. 1913 (MO, NY); Mara, along 20-200 m, 6 Sep 1974 (st), Steyermark & Manara 

road, 5-8 km S of Corpozulia, 50-90 m, 31 May 1980 110938 (MO, NY). 

(fl), Steyermark et al. 122991 (MO, NY). 

GUYANA. Upper Takutu/ Upper Essequibo: 

TRINIDAD. Botanic Gardens, cultivated, 5 Aug Karasabai, 100-110 m, 9 Mar 1990 (fl), Acevedo R. et al. 

1928 (yfr), Broadway s.n. (F, MO); cultivated, South- 3512 (CAY, NY, US), 9 Mar 1990 (fr), Acevedo R. et al.


'


I 9OWI WWL . v 6I 50W1 40wI 4owl 

4 

-v+ v~ 1sd 4 / w; ,~S~ '4 

* Melicoccus espiritosantensis - t) j '1 ' 

,. itK> X,'S r\X i 

0 

* Melicoccus pedicellaris \ ., . ,; Y 

A Melicoccus petiolulata S ,20S ~ $ 

+ Melicoccus olivaeformis ssp. olivaeformis A . .. 

v Melicoccus olivaeformis ssp. intermedia J $ E.. 

3515 (CAY, NY, US); Rupununi Savanna, Mt. Shiriri, 200- 

400 m, 2 Mar 1985 (fl), Jansen-Jacobs et al. 536 (CAY, 

F, MG, NY, US), 2 Mar 1985 (fr), Jansen-Jacobs et al. 

537 (CAY, F, MG, NY, US); Shea Village, 24 Sep 1997 

(fl), Jansen-Jacobs et al. 5596 (CAY, NY, US); Kanuku 

Mts. drainage of Moku-Moku creek, 150-400 m, 31 Mar- 

16 Apr 1938 (fr), Smith 3504 (F, IAN, MO, NY, P, US). 

BRAZIL. Amazonas: Rio Branco, Porto Alegre, 23 

Mar 1948 (fr), Froes 23108 (IAN, US). Espirito Santo: 

Mun. Linhares. Reserva Florestal da CVRD, 29 Sep 1972 

(fl), Lino 105 (RB); Linhares, 50 m, 31 Oct 1983 (fr), 

Martinelli & Soderstrom 9751 (RB, US); Estasco Farinha 

Seca, 21 Oct 1982 (fl), Silva 355 (NY, US). Minas 

Gerais: Faz. Ibituruna Figueira, Rio Doce, 3 Sep 1930, 

(fl, fr), Kuhlmann 294 (INPA, RB, US). Rio de Janeiro: 

Rio de Janeiro. Horto da Florestal, [cultivated?] 1929 

(fr), Anonymous 572 (RB). 

5. MELICOCCUS ANTIOQUENSIS Acevedo- 


Fig. 29. Distribution of species of Melicoccus (in part). 

Type. Colombia. Antioquia: Mun. San Luis, Canion 

del Rio Claro, northern sector, 325 m, 5?53'N, 

74?39'W, 2 Apr 1984 (fr), Cogollo, A. 1568 (holotype, 

JAUM; isotype, HUA). Fig. 30a-f 

A M. novogranatensi Acevedo-Rodriguez caulibus 

et foliolis fuifraceis, foliolis abrupte acuminatis differt. 

Tree 12-26 m tall. Young stems ferruginous-furfuraceous, 

slightly angled, becoming terete and lenticellate 

with age. Leaves paripinnate; distal process filiform, ca. 

7 mm long, early deciduous leaving a truncate base 2-3 

mm long; leaflets 4, opposite or the lower pair alternate, 

9.5-23.5 x 3.2-7.5 cm, oblong, elliptic, to nearly oblanceolate, 

subcoriaceous, the adaxial surface glabrous, 

slightly shiny or dull, with sunken or prominulous 

midvein, the abaxial surface ferruginous-furfuraceous, 

sparsely papillate, with prominent primary and secondary 

veins, the venation brochidodromus, secondary veins 

alternate or subopposite, arching toward the margin, tertiary 

veins finely reticulate, inconspicuous, the margins


N ~ ~ ~ ~ ~ ~ ~ ~ ~ " 

E ~~~~(1 ]imm ~~~~2mm 

Fig. 30. Melicoccus antioquensis. A. flowering branch. B. detail of inflorescence. C. flower with subtending 

bracteoles and bract. D. immature staminate flower with perianth removed, showing disc and stamens (left) and 

same showing pistillode. E. adaxial and abaxial views of petal, with reduced marginal projections. F. immature 

stamens lateral and ventral views. (All from Cogollo 974).


entire, the apex abruptly acuminate, the base slightly mary humid forest, 260 m, 05 1'S, 76?15'W, 25-30 

asymmetrical, one side attenuate the other obtuse; peti- Mar 1994 (fr), Dik 1179 (holotype, US; isotype, MO). 

olules nearly cylindrical, enlarged, wrinkled when dry, 

Fig. 3 1a-c 

6-10 mm long, ferruginous firfuraceous; rachis 3-7 cm 

A M. antioquensi Acevedo-Rodriguez caulibus et 

long, terete, striolate, ferruginous-furfuraceous to foliolis glabris, foliolis apice obtusis differt. 

tomentulose; petioles 5-12 cm long, with same indumentum 

as rachis, striate or less often lenticellate, adaxially 

Tree 12-30 m tall; trunk reaching 70 cm in diam.; 

flattened, enlarged at base. Inflorescence a fasciculate 

bark dark brown with numerous lenticels. Stems glapanicle, 

axillary or ramiflorous; 

brous, 

cataphylls acicular, furfuslightly 

angled, lenticellate. Leaves paripinnate; 

raceous, 3-7 mm long, 

distal process 

clustered at base of inflorescence; 

filiform, 5-10 mm long, early deciduous; 

leaflets 2 or4, opposite, 

axes 

7-23.5 (35) x 

1.5-7 cm 

3-10.5 (12) 

long, angled, ferruginous-tomentulose; 

cm, elliptic to oblanceolate, subcoriaceous, glabrous on 

bracts lanceolate, persistent, 1.5-1.8 mm long, with same 

both surfaces, the venation brochidodromus, slightly 

indumentum as the axis, bracteoles oblong-elliptic, ca. 1 

sunken or plane on adaxial surface, prominent on abaxial 

mm long, with same indumentum as the axis; dichasia 

surface, secondary veins alternate or subopposite, archreduced 

to a single flower subtended by a bract and two 

ing toward the margin, sometimes bifurcating, tertiary 

bracteoles; pedicels ca. 0.5 mm long, articulate at base. 

veins finely reticulate, the margins entire, undulate, the 

Calyx ca. 1.5 mm long, ferruginous-tomentulose, adaxiapex 

obtuse, the base slightly asymmetrical, one side 

ally puberulous, the sepals ovate-lanceolate, free to the 

attenuate the other obtuse; petiolules nearly cylindrical, 

base, obtuse at apex; petals lanceolate, ca. 1.5 mm long, 

enlarged, wrinkled when dry, 6-10 mm long, glabrous 

sparsely woolly on both surface, densely so on adaxial or puberulent; rachis 3-7 cm long, terete, striolate, glalowerportion, 

the apex obtuse, the base obtuse-cuneate; brous or puberulent; petioles 2.7-7 cm long, glabrous 

appendages wanting; disc annular, 5-lobed, glabrous, ca. or puberulent, striate, lenticellate, adaxially flattened, 

0.2 mm tall; stamens 8, the filaments woolly, apparently and enlarged at base. Inflorescence a fasciculate panicle, 

equal in length, the anthers lanceolate, 0.8 mm long, gla- axillary, ramiflorous or terminal; cataphylls lanceolate, 

brous, apiculate at apex; pistillode woolly. Fruits glo- furfuraceous, ca. 4 mm long, clustered at base of inflobose-ellipsoid, 

yellow, smooth, ferruginous-sericeous, rescence; axes 2.5-4 cm long, angled, ferruginous-fur- 

2-2.2 cm, apiculate at apex, the pericarp crustose, ca. furaceous to tomentulose; bracts ovate, persistent, 2-3 

0.6 mm thick, the endocarp granulate, glabrous. Seed mm long, with same indumentum as the axis, bracteglobose-ellipsoid, 

ca. 1.5 cm long, with tasty, yellow oles linear or elliptic, ca. 1 mm long, with same indusarcotesta. 

Embryo with erect cotyledons of similar size. mentum as the axis; dichasia reduced to a single flower 

Melicoccus antioquensis seems to be closely related subtended by a bract and two bracteoles; peduncle ca. 

to M. novogranatensis because they share numerous 0.5 mm long; pedicels ca. 0.5 mm long, articulate at 

morphological features. Melicoccus antioquensis dif- base. Calyx 1.5-1.7 mm long, ferruginous- tomentulose, 

fers from the latter by the ferruginous furfuraceous adaxially puberulous, the sepals free to the base, ovate, 

stems and abaxial surface of leaflets (vs. glabrous), and obtuse at apex; petals subulate, 1-1.2 mm long, puberuthe 

leaflets with abruptly acuminate apex (vs. obtuse). lent except for the ciliate margins and the adaxial thick- 

The specific epithet refers to the Colombian Depart- ened lower portion, the apex acute, the base truncatement 

where the type was collected. 

cuneate; appendages wanting; disc annular, unlobed, 

glabrous, ca. 0.2 mm tall; stamens 8, the filaments gla- 

Phenology. Collected in flower during November 

brous to sparsely woolly, apparently of same length, 

and in fruit during April. 

the anthers ovate, ca. 0.6 mm long, glabrous, apiculate; 

Distribution and Ecology. (Fig. 36) Known only pistillode woolly. Fruits ovoid, yellow (bluish when 

from the type locality. 

immature fide Romoleroux), smooth, ferruginous-sericeous, 

2.2-3 cm, apiculate, the pericarp crustose, ca. 

Representative specimens examined. COLOMBIA. 

0.6 mm thick, the endocarp smooth, glabrous. Seed 

Antioquia: Mun. San Luis, Canion del Rio Claro, NW 

sector, 340-425 m, 30 Nov 1983 (fl), Cogollo 974 

ovoid, ca. 2 cm long, with cream sarcotesta. Embryo 

(JAUM, HUA). 

with erect cotyledons of equal size. 

Melicoccus novogranatensis is morphologically 

similar to M. antioquensis and M. oliviformis. It can be 

6. MELICOCCUS NOVOGRANATENSIS distinguished from M. antioquensis by the glabrous 

Acevedo-Rodriguez, sp. nov. 

stems and leaflets (vs. ferruginous-furfiuraceous), and 

from M oliviformis by the single-flowered dichasia (vs. 

Type. Ecuador. Napo: Cant6n Aguarico. Huaorani dichasia simple or compound), calyx 1.5-1.7 mm long 

Ethnic Reserve, Maxus Oil Pipe road, km 86-89, pri- (vs. 2-2.7 mm), and by its subulate petals with entire


Fig. 31. Melicoccus novogranatensis. A. portion of branch with leaf. B. fruiting branch. C. embryo. (A from 

(Romoleroux 3068; B-C from Romero C. 4153). 

margins (vs. petals rhombate with appendage-like mar- 

gins). The specific epithet refers to the colonial name 

where the species occurs. 

Phenology. Collected in flower in July and Novem- 

ber and in fruit during January, April, October, and 

November. 

Distribution and Ecology. (Fig. 36) Known from 

Colombia and Ecuador in terra firme forest on loamish 

soil, humid primary forest, and primary moist forest. 


CaquetA: Along Caqueta river, upstream from mouth 

of Caguan river, 29 Apr 1953 (fr), Romero C. 4153 

(COL). Meta: Sierra La Macarena, in front of Isla Grande, 

460 m, 15 Nov 1975 (fr), Idrobo 8497 (COL). Tolima: 

Mariquita, 250 m, Renteria et al. 5410 (HUA). 

ECUADOR. Napo: Orellana, Yasuni, Estaci6n 

Cientifica Yasuni, Tiputini river, NE of confluence with 

Tivacuno river, km 20, 250 m, 28-30 Jul 1993 (st), 

Aulestia & Grefa 167 (MO, US), 28-30 Jul 1993 (fl), 

Aulestia & Grefa 192 (QCNE, MO, US), km 53-60, 230 

m, 6 Oct 1993 (fr), Dik 721 (QCNE, MO, US), km 44 

along Maxus road, 200-300 m, 22 Jan 1998 (fr), 

Romoleroux et al. 3068 (QCA, US).


2 mm.G ~ J \ ~ i 

Fig. 32. Melicoccus aymardii. A. flowering branch with detail of abaxial leaf surface. B. staminate flower with 

filaments not yet expanded. C. mature staminate flower. D. staminate flower with perianth removed showing disc 

and stamens. E. l.s. staminate flower showing pistillode and stamens. F. abaxial and adaxial views of petal with 

marginal projections. G. stamens. ( All from Aymard & Ortega 2521). 

7. MELICOCCUS AYMARDII Acevedo- 


Type. Venezuela. Portuguesa: Dtto. Araure, 5 km 

NE of Agua Blanca, 25 km NW of Acarigua, 

afloramientos calc'areos, 250 m, 9041'N, 69004'W, 5 

May 1984 (fl), G. Aymard & F Ortega 2521 (holo- 

type, US; isotype, PORT, n.v.). Fig. 32a-g 

A Melicoccus espiritosantensis Acevedo-Rodriguez 

inflorescentia racemosis, filamentis pilosis differt. 

Tree 5 m tall. Stems terete, lenticellate, puberulent, 

dark grey. Leaves paripinnate; distal process minute, 

inconspicuous; leaflets 2-4, opposite, 4-9 x 1.4-2.3 

cm, elliptic or lanceolate, chartaceous, glabrous, dull, 

venation brochidodromus, tertiary veins finely reticulate, 

inconspicuous except for the adaxially sunken,


abaxially prominent midvein, the margins entire or obtuse, rounded, less often acute or mucronulate, the base 

slightly undulate, the apex acute or less often obtuse, inequilateral, one side obtuse or truncate the other acute 

the base obtuse sometimes asymmetrical; petiolules 2- or cuneate; petiolules less than 1 mm long, drying dark 

6 mm long, glabrous to pubescent, nearly cylindrical, brown; petioles 1-3 cm long, glabrous, flattened, cuneate, 

adaxially furrowed, wrinkled when dry; rachis 1-1.5 cm or less often distally winged, narrowing toward the base. 

long, tnrllate, puberulent, dull, grayish; petioles 1.2-3 cm Inflorescence a terminal simple or basally branched 

long, nearly terete, striolate, the base slightly enlarged and raceme, 2-4.8 cm long, the axes glabrous, with a few 

flattened along adaxial surface. Inflorescence of scattered papillae; cataphylls broadly ovate, 1-1.5 mm 

racemes, fasciculate on distal portion ofbranch, 2-7.5 cm long; bract triangular or subulate, 0.5-1 mm long, ciliate 

long; cataphylls wanting; axes angled, sparsely ferrugi- at margins; pedicels 3-4 mm long, not articulate. Flownous-tomentulose; 

bracts ovat'e-deltate, persistent, 0.6 ers fragrant; calyx 4(-5)-merous, the sepals in 2 unequal 

mm long, with same indumentum as the axis; pedicels pairs, 1.5-2 mm long, greenish, connate at base, ovate, 

2.5-3 mm long, tomentulose, not articulate, with a pair concave, glabrous, ciliate at margins; petals 4(-5), 

of opposite or alternate bracteoles, ca. 0.5 mm long. Calyx whitish to yellowish, obovate, 2.5-4 mm long, concave, 

2-2.5 mm long, tomentulose, the sepals free to the base, rounded at apex, woolly-pubescent along margins, with 

lanceolate-deltate; petals narrowly rhombate, ca. 2.2 mm a pair ofminute (1-1.5 mm long), linear, woolly-pubeslong, 

whitish woolly-pubescent along margins; append- cent appendages; disc 4-lobed, swollen, glabrous, brownages 

of two minute, suprabasal-marginal projections, ish; stamens 8, spreading, the filaments of unequal 

densely whitish woolly-pubescent; disc annular, gla- lengths, glabrous, 3.5-6 mm long in male flowers, and 

brous, ca. 0.5 mm tall; stamens 7-9, spreading, the fila- ca. 1 mm long in female flowers, the anthers elliptic, ca. 

ments subulate, pilose to woolly on the lower ?2, of un- 0.5 mm long, retuse at apex; ovary glabrous, ovoid, the 

equal lengths, 1.5-2 mm long, the anthers ovate-deltate, style 1.5-2 mm long, the stigmas bilobed, papillate. Fruit 

ca. 0.5 mm long, glabrous, apiculate at apex. Pistillate ellipsoid, light yellow, 2.5-3.3 cm long, the pericarp 

flowers and fruits unknown. 

coriaceous, ca. 1 mm thick, the endocarp woolly-pubes- 

Melicocus aymardii seems to be closely related to cent. Seeds solitary, 1-1.8 cm long, with a yellowish, 

M. espiritosantensis. However, M. aymardii differs fleshy, edible fleshy testa. Embryo with erect, or siightly 

from M. espiritosantensis by: racemose inflorescences curved, similar-sized cotyledons. 

(vs. paniculate); dichasia reduced to a single flower (vs. The specific epithet refers to the petals bearing 

dichasia simple); petals with shallow appendage-like "scales" or appendages. 

margins (vs. petals with margins deeply cleft); and pilose 

filaments (vs. glabrous). The specific epithet hon- 

Phenology. Flowers in September and October, and 

ors Gerardo Aymard (PORT), collector of the type and 

fruits from October to January. 

eminent Venezuelan botanist. 

Distribution and Ecology. (Fig. 36) Bolivia (Santa 

Phenology. Collected in flower in May. 

Cruz), Brazil (Mato Grosso do Sul), Paraguay and 

northem Argentina. Along gallery and flooded forests, 

Distribution and Ecology. (Fig. 36) Known only and shrubby savanna on sandy and clayish soils. Culfrom 

the type collection. 

tivated for its fruit and as a timber species. 

8. MELICOCCUS LEPIDOPETALUS Radlkofer, [as 

Common names and uses: Bolivia: Motoyoe; 

Paraguay: Yvapovo, Papamundo. 

Melicocca lepidopetala] Sitzungsber. Math.-Phys. Representative specimens examined. BRAZIL. 

Cl. K6nigl. BayerAkad. Wiss. Miinchen 8: 344. 1878. Mato Grosso: Faz. Miranda, vic. of Aquidauana river, 

Type. Bolivia. Santa Cruz: Chiquitos, d'Orbigny 818 

(holotype, G, photo at US). Fig. 33a-f 

Dioecious tree 6-20 m tall; bark dark brown, in small 

18 Sep 1980 (fr), M. Pires & Furtado 17165 (MG). 

BOLIVIA. Santa Cruz: Prov. Andres Ibaniez, 12 km 

E of Santa Cruz on rd. to Cotoca, 375 m, 12 Aug 1987 

(fl), Nee 35630 (US), 7 km SE of Naranjillos, 480 m, 

plates; trunk reaching to 1 m in diam. Stems glabrous, 30 Sep 1990 (fl), Nee 38948 (US). 

angled. Leaves paripinnate; distal process early decidu- PARAGUAY. Without specific locality. Without date 

ous, rarely developing into a terminal leaflet; leaflets 2, (fl), Hassler 7244 (US). Central: Jardin Botanico & 

opposite, strongly asymmetrical, one half ofblade ellip- 

Zool6gico, Trinidad, Asunci6n, natural reserve, Oct- 

Nov 1991 (fr), Blas P. 1268 

tic, the other oblong, or obovate, 4-11.5 x 

(US); Ypacaray, Aug 1913 

2-4.5 cm, 

(fl), Hassler 12221 (US); San Lorenzo, 15 Oct 1893 (fl), 

chartaceous, glabrous on both surfaces, the abaxial sur- Lindman 2205 (US); Ypacaray river, 17 Sep 1893 (fl), 

face sometimes minutely puberulent, the venation brochi- Malme 962 (US); Without specific locality, 1888 (fr), 

dodromus, prominent on both surfaces, tertiary vena- Morong 817 (US-2). Concepci6n: 24 Oct 1991 (fr), 

tion reticulate, the margins entire, undulate, the apex, Basualdo 3683 (MO).


4 X \I N 2mm 

Fig. 33. Melicoccus lepidopetalus. A. flowering branch, with detail of lealfet's venation. B. staminate flower with 

detail of stamen, and 1.s. of same. C. flower bud. D. pistillate flower, and l.s. of same showing ovules. E. adaxial and 

abaxial views of petal. F. fruiting branch. (A, D-E from Malme 962; B-C from Hassler 12221; F from Perez 1268). 

ARGENTINA. Corrientes: Corrientes, 6 Oct 1978 

(fl), Ferrucci 69 (US); Empedrado, Estancia La Yela, 20 

Sep 1952 (fl), Pedersen 1833 (US). 

9. MELICOCCUS BIJUGATUS Jacquin, Enum. 

Syst. P1. 19. 1760. Lectotype, Jamaica, Jacq., Select. 

Stirp. Amer. Hist., tab. 72. 1763, designated by 

Howard, 1989. Figs. 12d, 14a-b, 18e-f, 34a-d 

Melicocca bijuga Linnaeus, Sp. P1. ed. 2, 1: 495. 

1762. nom. illeg., (orth. variant). 

Melicocca carpopodea Juss., MWm. Mus. Hist. Nat. 

3: 187, fig. 4. 1817. nom. illeg., (superfluous 

renaming). 

Melicocca bijuga L.[as Melicoccus bijuga] f. alata 

B. P. Kitanov, Phytology (Bulgaria) 11: 48. 1979. 

Type. Cuba. Oriente: Siboney, vic. of Santiago de 

Cuba, 50-100 m, 9 Apr 1970, Kitanov 82105 

(SO, n.v.).


N BI 

Fig. 34. Melicoccus bijugatus. A. flowering branch, with detail of winged leaf rachis. B. staminate flower and l.s. 

of same. C. pistillate flower and l.s of same showing ovule. D. portion of infructescence and l.s. of fruit. [Reprinted 

from Acevedo-Rodriguez (1996)]. 

Dioecious ormonoecious tree 10-25 m tall; bark light greenish, 1.5-2.2 mm long, oblong, concave, glabrous 

grey, smooth, with horizontal markings. Stems glabrous, except for the woolly margins; petals 5, cream to yelnearly 

terete. Leaves paripinnate; distal process subu- lowish, obovate, 2-2.5 mm long, narrowed at base, 

late, early deciduous; leaflets (2) 4, opposite or rounded at apex, woolly to sparsely woolly at margins, 

subopposite, asymmetrical, elliptic, oblong, ovate or lacking appendages; nectary disc annular, swollen, lobed, 

obovate, 4-14(20) x 2.2-5(7) cm, chartaceous, glabrous 

on both surfaces, the venation brochidodromus, adaxially 

prominulent, abaxially prominent, tertiary venation 

reticulate, the margins entire, slightly wavy, the apex acute 

or obtuse, the base attenuate on to the petiolule; petiolules 

drying brownish, not pulvinate, 2 mm long; rachis 1.5- 

3(4.5) cm long, glabrous, winged to broadly winged, or 

less often marginate; petioles 2.5-7 cm long, flattened, 

subterete or less often winged to broadly winged, glabrous. 

Panicles terminal, 5-12 cm long; cataphylls foglabrous; 

stamens 8, spreading, the filaments ofunequal 

length, glabrous, ca. 4 mm long in male flowers, and 1 

mm long in female flowers, the anthers dorsifixed, elliptic, 

ca. 0.6 mm long; ovary glabrous, cylindric-ellipsoid, 

the style ca. 0.5 mm long, the stigmas bilobed orbilobeddisciform, 

papillate. Fruit subglobose or ellipsoid, green, 

2-3.5 cm long, the pericarp coriaceous, ca. 1 mm thick. 

Seeds 1(2), 1.5-2.5 cm long, with a tan, fleshy testa. 

Embryo with erect, similar-sized cotyledons. 

The specific epithet refers to the bijugate leaves. 

liaceous, broadly ovate, 4-5 mm long; axes, glabrous, Phenology. In the Greater Antilles and Central 

angled, striate, the staminate inflorescences with 4-8 America flowering occurs between March and August 

lateral branches, slightly shorter than the main axis, the (rarely as late as November), followed by fruiting from 

pistillate ones with 3-4 mostly basal, short branches; April to August. In northern South America and the 

bracts triangular, minute, ciliate at margins; pedicels (2)5- Leeward Lesser Antilles, flowering has been recorded 

7 mm long, not articulate. Flowers fragrant; sepals 4, from October to May and fruiting from January to June.


Distribution and Ecology. (Fig. 36) Native to north- HAITI. Massif de la Selle, 1 May 1927 (fl), Ekman 

em SouthAmerica, perhaps from dry forest. Commonly H-8033 (US); Vic. of Fond Parisien, 5-13 May (fl), 

planted and naturalized along roads and in secondary dry 

forest throughout northern South America, Central 

America, the West Indies, Cameroon, Gabon (Fouilloy 

& Halle, 1973a, 1973b), and the Pacific Islands. In secondary 

forest on Thomas and St. John (US Virgin Islands) 

is sometimes the dominant species. 

Leonard 4175 (US). 

DOMINICAN REPUBLIC. Samani: Vic. of 

Sanchez, 29 May 1922 (fl), Abbott 2427 (US). Santo 

Domingo: Coastal plain, cultivated, 13 May 1929 (fl), 

Ekman 19447 (US). 

PUERTO RICO. Coamo, 19 Apr 1933 (fl), Britton 

& Britton 10138 (US); San German, 12 Aug 1950 (fr), 

Common names and uses: Colombia: Mamoncillo, 

Little 13604 (US); Sabana Grande, 16 Mar 1935 (fl), 

Sargent 219 

mamon; Cuba: mamoncillo; Dominica: kenip; Domini- 

(US); Fajardo, 20 Jun 1895, (fl), Sintenis 

1597 (M); Peiiuelas, 12 Jul 1886 (fl), Sintenis 4788 (M); 

can Republic: limoncillo; El Salvador: mel6n; Haiti: 

Guanica, 30 Aug 1915 (fl), Stevenson 3014 (US). 

canape; Honduras: mamon; Jamaica: genipe; Panamal: UNITED STATES VIRGIN ISLANDS. St. Croix: 

mam6n; Puerto Rico: quenepa; United States (Florida): Without locality, 5 Sep 1901 (st), Britton & Cowell 10 

Spanish lime (fide Popenoe, 1920); Venezuela: mam6n; (US). Altona, E of Christianted, 29 May 1977 (fl), 

Virgin Islands: genip, kenep, keneppy tree. Commonly Fosberg 56774 (US); Estate Thomas, Experimental 

cultivated for is delightful fruits (fleshy testa), which Forest, 21 Jun 1966 (st), Little 21517 (US); Buck Isare 

available during the fruiting season along roadsides land. 8 Apr 1967 (st), Little 22039 (US). St. John: 

and markets in Colombia, French Guiana, Guyana, New 

York City, Puerto Rico, and Venezuela. An alcoholic 

drink called "bili" is made in Vieques Island (Puerto 

Rico) by aging rum with the fruits. 

Lameshur to Reef Bay ruins, 29 May 1993 (fl), Acevedo 

R. & Siaca 5441 (US). St. Thomas. Without locality, 

26. May 1921 (fl), Morrow 121 (US). 

BRITISH VIRGINS ISLANDS. Jost van Dyke: 12 

Apr 1967 (fl), Little 21968 (US). Virgin Gorga: Span- 

Representative specimens examined. UNITED ish Town, 13 Nov 1999(fl), Acevedo R. & Clubbe 10956 

STATES. Florida: Key Largo, Hwy 1, cultivated, 19 (US); Without locality, 20 Jun 1969 (st), Little 23767 (US). 

Mar 1958 (st), Stern & Chambers 220 (US). 

LEEWARD ISLANDS. Antigua: Near Jennings, 14 

HONDURAS. Morazin: Escuela Agricola Panamer- May 1938(fl), Box 1442 (US); Without locality, 4-16 

icana Campus, cultivated, 800 m, 23 Mar 1970 (fl), Feb 1913 (fl), Rose et al. 3454 (US). Dominica: 

Molina R. & Molina 25601 (US); 13 Apr 1964 (yfr), Canefield Estate, 6 Jun 1977 (fr), Nicolson 4219 (US); 

Molina R. 13703 (US). 

St. Paul, Massacre, 16 Apr 1992 (yfr), Whitefoord 7053 

EL SALVADOR. San Salvador: Without specific (US). Guadeloupe: Without locality, 1895 (fl), Duss 

locality, Mar 1923 (fl), Calder6n 1526 (US). 

3718 (US); Basse Terre, Botanic Garden, 17-22 Apr 1979 

NICARAGUA. Managua: Managua, no data, 10 Mar (yfr), Howard & Howard 19449 (US). 

1926 (fl), Chaves 171 (US); Villa Stimson, Apr 1932 COLOMBIA. Antioquia: Mun. Amalfi. Arenas 

(yfr), Garcia 1063 (US). 

Blancas trail, ca. 1 100 m, 12-13 Apr 1994 (fl), Fonnegra 

PANAMA. Canal Zone: Balboa, Nov 1923-Jan 1924 et al. 4815 (US); Venecia, km 1.5 along road Bolombolo- 

(st), Standley 29248 (US). Panama: Taboga Island, Venecia, cultivated, 700 m, 13 Mar 1987 (fl), Zarucchi 

planted, Dec 1923 (st), Standley 27931 (US). & Echeverry 4673 (US). Bolivar: San Martin de Loba, 

BAHAMAS. South Bimini: Without specific local- Apr-May 1916 (fl), Curran 84 (US); Vic. Turbaco, Hac. 

ity, May 1948 (st), Howard 10053 (US). 

Coloncito, 200-300 m, 9 Nov 1926 (fl), Killip & Smith 

CUBA. Camaguey: Vic. of Nuevitas, 21 Mar 1909 14349 (US). La Guajira: La Macuira, Jassai, 4 Mar 1963 

(fl), Shafer 1007 (US). Habana: Vic. Santiago de las (fr), Saravia T 2359 (US); Nazareth, Cerro Itojoro, 300- 

Vegas, 17 May 1905 (yfr), Cook 41 (US); 18 Jul 1904 400 m, 15 Apr 1964 (st), Saravia T & Saravia 3667 

(st), Wilson 251 (US); Isla de Pinos: Nueva Gerona, 15 (US); Uribia, 19 Oct 1963 (fl), Saravia T 2876 (US); 

Apr-1 Jun 1904 (fl), Curtiss 444 (US). Las Villas: Santa Marta, ca. 75 m, Mar 1898-1901(fl), Smith 1705 

Cienfuegos, Harvard Tropical Garden, cultivated, 21 Mar 

1926 (st), Jack 4366 (US), 16 Mar 1926 (fr), Jack 4321 

(US-2). Magdalena: Neguange, Tayrona National Park, 

100 m, 12 Jun 1993 (fr), Gentry & Ortiz 79870 (US). 

(US); 11 Apr 1927 (fl), Jack 5151 (US). Oriente: Santiago Tolima: El Guama, road to Saldafia, 400 m, 15 Aug 1950 

de Cuba, between Caban and Punta de Sal, escaped, 12 (fl), Garcia-Barriga, 13488 (US); Vic. of Prado, ca. 

Apr 1919 (fl), Ekman 9480 (US); Valley of Matamoros 400 m, 9 Feb 1944 (fr), Little 7160 (US). 

river, S of Holguin, 11 Apr 1909 (yfr), Shafer 1324 (US), VENEZUELA. Bolivar: Ciudad Bolivar, 9 Jan 1921 

15 Apr 1909 (fl), Shafer 1383 (US), 13 Apr 1909 (fl), (yfr), Bailey & Bailey 814 (US); Carabobo: Valencia, 

Shafer 1341 (US). 

400-800 m, 19 Feb 1920 (fl), Pittier 8776 (US). Distrito 

JAMAICA. Manchester: Mandeville, Alligator Federal: Vic. of Caracas, Oct 1919 (fr), Rose & Rose 

Pond, 600 m, 20 Mar 1931 (st), Miller 1401 (US). St. 21625 (US), 19 Oct 1916 (st), Rose & Rose 21678 (US). 

Andrew: Near road between Gordon Town and Guava Guirico: Road to Calabozo, vic. of El Sombrero, 19 Feb 

Ridge, ca. 600 m, 3 Jul 1966 (fr), Anderson & Sternberg 1924 (fl), Pittier 11451 (US). Miranda: Ocumare del Tuy, 

3006 (US); Dallas Mountain, Lindos Gap, ca. 500 m, 300-1200 m, 1 May 1918 (fl), Pittier 7814 (US). 

18 Jun 1963 (fl), Crosby et al. 204 (US). St. Elizabeth: TRINIDAD. St. Amis. Cultivated, 20 Apr 1920 (fl), 

Pepper, 19 Mar 1931 (st), Miller 1369 (US). 

Broadway s.n. (US).


F J cmC 

Fig. 35. Melicoccus jimenezii. A. flowering branch. B. staminate flower, and 1.s. of same. C. adaxial view of 

petal. D. pistillate flower. E. fruiting branch. F. embryo, lateral view. (A-C from Garcia & Rodriguez 5773a; D 

from Jimenez 3312; E-F from Jimenez 2703). 

FRENCH GUIANA. Bassin du Maroni: Village 

Assici, 15 Dec 1988 (st), Fleury 758 (CAY). Ile de Cayenne: 

Cayenne, cultivated, 19 Apr 1994 (fl), Loubry 

1956 (CAY). 

BRAZIL. Park: Belem, Horto Museu Goeldi [cultivated], 

26 Jul 1957 (fr), Cavalcante, P. 271 (IAN). 

FRENCH POLYNESIA. Makatea, Vaitepaua Village, 

[cultivated] 20 Feb 1982 (fl), Florence 4047 (US). 

HAWAII. Oahu, Wimanalo Research Station, cultivated, 

16 Nov 1976 (fr), Shearard & Spence 2 (US). 

SOCIETY ISLANDS: Moorea Island. Oponohu Bay, 

cultivated, 12 Jul 1981 (st), Fosberg 60936 (US). 

10. MELICOCCUS JIMENEZII (Alain) Acevedo- 

Rodriguez, Moscosoa, 9: 60. 1997. Talisiajimenezii 

Alain, Phytologia47(3): 181. 1980. Type. Dominican 

Republic. La Altagracia: Bayahibe, coastal scrub, 

sea level, 2 Oct 1976 (fr), Alain Liogier, P Liogier 

& J.J. Jimenez 25442 (holotype, UPR; isotype, 

MO, NY). Fig. 35a-f 

Dioecious tree 7-8 m tall; bark light grey, fissured, 

trunk reaching 25-30 cm in diam. Stems terete, glabrous, 

striate, brownish. Leaves paripinnate; distal process 

minute, subulate, persistent, occasionally a terminal 

leaflet is produced; leaflets 4, opposite, 4-9 x 2-4.5 

cm, elliptic to broadly elliptic, chartaceous, glabrous, 

dull, venation brochidodromus, abaxially prominent, 

tertiary veins finely reticulate, the margins entire, 

strongly revolute, the apex obtuse or rounded, the base 

asymmetrical, one side obtuse the other rounded; petiolules 

ca. 1 mm long, drying brownish, glabrous; ra-


W 9ow -+8atY X; ; 70WI 64wl o50W 40wi 

> 

u ~ 

l 

~~ ,_r ~~ w +' '> >-x s 

N~~~~~~~~~~~~~~~~~~~~~~~~~O 

- ' -- 

~t t'^ '-'4*, '- -+' 

, 

aS,~~~ ;, 

- # . . c^ ` 

4 

(r t2. 

. iON 1 

o~ 

> 4 t~~~~~~~~~~~~~~~~~~~, 

x Melicoccus antioquensis -\4X s 1> 

*: Melicoccus aymardii Q , * 

_ + ~~~~Melicoccus bijugatus 

v Melicoccus jimenezii 8\ >/ L os X'' 

*Melicoccus lepidopetalus ;f->-- ?- , - 

| * Melicoccus novagranatensis t , , K 

Fig. 36. Distribution of species of Melicoccus (in part). 

chis 1-3.5 cm long, subterete, or rarely narrowly The specific epithet honors Jose J. Jimenez, emiwinged, 

striate, glabrous; petioles 2.4 cm long, gla- nent Dominican botanist, who accompanied the author 

brous, subterete, pulvinate at base. Inflorescence a (Alain) to the type locality. 

raceme, 5-6 cm long, on distal portion of branch; 

cataphylls broadly ovate, 1-2 mm long; axes angled, 

glabrous; bracts early deciduous; pedicels 6-7 mm 

Phenology. Collected in flower in January and May 

and in fruit from May to October. 

long, not articulate. Calyx 4-merous, the sepals green, Distribution and Ecology. (Fig. 36) An endanovate, 

concave, 1-1.2 mm long, ciliate along margins; gered, endemic species, known only from the Dominipetals 

4, oblong to wide-elliptic, whitish, ca. 2.2 mm can Republic from coastal scrubs in Bayahibe, on the 

long, ciliate along margins, lacking appendages, or with eastern side of the island, and in the central mountaina 

pair of minute, basal appendages; disc annular, 4- ous region of Jacagua in the vicinity of Santiago de los 

lobed, glabrous; stamens 8, spreading, the filaments Caballeros. 

glabrous, slightly unequal, ca. 2.5 mm long, the anthers 

oblong-elliptic, ca. 0.4 mm long, retuse at apex; pistillode 

minute, glabrous; ovary glabrous, ovoid, the stigma 

Common names and uses. Cotoperi, cotoperiz. 

Fruits are consumed locally. 

nearly sessile, 2-lobed. Fruits yellowish, ellipsoid or Representative specimens examined. DOMINIovoid, 

ca. 2.5 cm long, the pericarp leathery, ca. 0.7 CAN REPUBLIC. La Altagracia: 300 m E of 

mm thick. Seed solitary, ca. 1.5 cm long, with fleshy, Bayahibe, 10 m, 19 May 1995 (fl), Garcia etal. 5773a 

yellowish, edible, sour fleshy testa. Embryo with coty- (JBSD, US); 2 Oct 1976 (fr), Liogier et al. 8247 (NY, 

ledons diagonally superimposed, the upper one much US); 21 Feb 1976 (st), A. & P Liogier 24886 (NY); 

larger than the lower one. 

ca. 400 m NE of Navy Headquarters, 0 m, 21 May 1992 

20X'o _P',S s.............ivz\ ' 

,'2


(fr), Garcia & Jimenez 3861 (JBSD, US); 21 May underground stem. Leaves spirally arranged, pari- or 

1992 (fr), Garcia & Ramirez 3930 (JBSD, US); 19 impari-pinnately compound; distal leaflet rudimentary, 

Sep 1996 (fr), Acevedo R. et al. 8477 (JBSD, US). represented by a minute process; leaflets 2-30, oppo- 

Santiago: Jacagua, vic. Santiago de los Caballeros; site or alternate, usually with an enlarged, pulvinate 

cultivated, 1954 (fr), Jimenez 2703 (US); 28 Jan 1956 petiolule; rachis short to elongated, cylindrical, keeled, 

(fl), Jimenez 3312 (US-2). 

furrowed, flattened or angled; petiole with similar shape 

EXCLUDED TAXA 

as the rachis, and usually with a markedly enlarged base; 

stipules wanting. Inflorescence of terminal, axillary or 

Melicoccus amoenus Hasskarl = Lepisanthes amoena cauliflorous, panicle-shaped or racemiform thyrses, 

(Hasskarl) Leenhouts (Sapindaceae) 

with flowers in lateral, simple or compound dichasia. 

Melicocca apetala Poiret = Doratoxylon mauritianum Flowers 5-merous, actinomorphic, functionally stami- 

Thouars ex Bojer (Sapindaceae) 

nate or pistillate (plants sequentially monoecious); 

Melicocca australis Steudel = Diploglottis australis pedicels articulate; sepals imbricate, connate at base, 

(Cunningham) Radlkofer (Sapindaceae) 

shorter to longer than the petals; corolla of distinct pet- 

Melicocca dentata Jussieu = Hippobromus pauciflorus als with imbricate aestivation, inserted on the base of 

(Linnaeus f.) Radlkofer (Sapindaceae) 

an extrastaminal nectary disc, the petals erect or reflexed, 

Melicocca diphylla Bojer = Doratoxylon mauritianum 

oblong, lanceolate, elliptic, or spatulate, and containing 

Thouars ex Bojer (Sapindaceae) 

an adaxial petaloid appendage, petaloid appendages 

Melicocca diversifolia Jussieu = Doratoxylon 

simple or bifid, erect, sericeous or tomentose, as long 

mauritianum Thouars ex Bojer (Sapindaceae) 

as or shorter than the petals; disc extrastaminal, annu- 

Melicocca geniculata Sprengel = Protium aracouchili 

March 

lar or cup-shaped, with a 5-lobed outline, glabrous or 

(Burseraceae) 

with 

Melicocca javanica Hasskarl = 

various indumenta; stamens 5-8(-10), spreading 

Elattostachys verrucosa 

(Blume) Radlkofer (Sapindaceae) 

or erect, in one series, the filaments of equal or unequal 

Melicocca paniculata Jussieu = Exothea paniculata 

length, glabrous or with various indumenta, the anthers 

(Jussieu) Radlkofer (Sapindaceae) 

basifixed, dorsally with ovate, oblong, lanceolate, or 

Melicocca pubescens de Candolle = Schleichera oleosa linear outline; ovary 3- carpellate, each carpel with a 

(Loureiro) Oken (Sapindaceae) 

single campylotropous or apotropous ovule, the style 

Melicocca trijuga Jussieu = Schleichera oleosa solitary, short to elongated, the stigmatic surface elon- 

(Loureiro) Oken (Sapindaceae) 

gate and trigonous or short and capitate. Fruits inde- 

Melicocca triptera Blanco = Tristira triptera (Blanco) hiscent, ellipsoid, ovoid, or globose, with leathery to 

Radlkofer (Sapindaceae) 

woody mesocarp. Seed 1(-3), with a more or less fleshy 

and usually edible testa. Embryo fleshy, notorrhizal with 

TALISIA AUBLET 

cotyledons superimposed, horizontally, or obliquely so, 

TALISIAAublet, Hist. PI. Guiane. 1: 349. 1775. Type. 

Talisia guianensis Aublet. The name is derived from 

Toulichi, the common name of this species in the 

Carib language. 

or rarely laterally oblique; or lomatorrhizal with straight, 

erect cotyledons of similar size, the radicle elongated. 

Pollen colporate, nearly spherical, isopolar, triangular 

or trigonous, with straight to convex sides, or brevicolporate, 

oblate, isopolar, triangular, with straight or 

concave sides 

A continental Neotropical genus of 52 species. 

Racaria Aublet, Hist. P1. Guiane. II, Suppl. 24. 1775, 

Tab. 382. Type: Talisia sylvatica (Aublet) 

Radlkofer. 

Acladodea Ruiz & Pav6n, Syst. Fl. Peruv. 1: 262. 

1798. Type: Talisia pinnata (Ruiz & Pav6n) 

Radlkofer. 

Meleagrinex Arruda da Camara ex Koster, Travels 

Brazil 496. 1816. Type: unknown. 

Comatoglossum Karsten & Triana in Triana, Nuev. 

Jen. Esp. 10. 1854. Type: Talisia stricta (Triana) 

Triana & Planchon. 

Tapirocarpus Sagot, Ann. Sci. Nat. Ser. VI, 13: 292. 

1882, pro parte vegetativa; fruit cf. Protium 

(Burseraceae). Type: Tapirocarpus talisia Sagot. 

Diclinous, sequentially monoecious, understory 

palm-like treelets, medium-sized to large emergent trees, 

or less frequently shrubs with a prostrate, elongated 

Key to the subgenera 

1. Anthers elliptic or ovate in outline (length/ 

width ratio < 2); filaments of unequal lengths; 

sepals free to the base of calyx (or nearly so); 

pollen colporate, nearly spherical, isopolar, 

triangular or trigonous, with straight to convex 

sides ....... . T subgenus Pitombaria page 51 

1. Anthers oblong, linear or lanceolate in outline 

(length/width ratio > 2); filaments of equal or 

nearly equal length (except T. laevigata and T. 

hexaphylla); sepals connate at base for at least 

?/4 of the calyx length; pollen brevi-colporate, 

oblate, isopolar, triangular, with straight or 

concave sides ......... T. subgenus Talisia page 88

TAXONOMIC TREATMENT 5 1 

I. TALISIA subgenus PITOMBARIA (Radlkofer) 

Inflorescence of terminal, axillary or cauliflorous, 

Acevedo-Rodriguez, stat. nov. Talisia subsection 

panicle-shaped, racemiform, or congested thyrses. Se- 

Pitombaria Radlkofer, Sitzungsber. Math.-Phys. 

pals usually free to the base, or less often (T hexaphylla) 

Cl. K6nigl. Bayer Akad. Wiss. Miinchen 8: 344. 

connate at base. Petals with an adaxial petaloid append- 

1878. Type. Talisia esculenta (Cambessedes) 

age. Stamens (7) 8 (-10), erect, the filaments of un- 

Radlkofer. Figs. I la, 41, 45, 50, 54, & 62. 

equal lengths; anthers elliptic or ovate in outline (length/ 

Medium-sized to large emergent trees or treelets, width ratio < 2). Pollen colporate, nearly spherical, 

less often shrubs, or exceptionally shrubs with hori- isopolar, triangular or trigonous, with straight to conzontal, 

elongated underground stems. Leaflets 2-16. vex sides. 

Key to the species of Talisia subgenus Pitombaria 

1. Leaflets with abaxial surface white-granulose, and scattered, appressed-pubescent . ........................ T granulosa 

1. Leaflets with abaxial surface otherwise. 

2. Anthers apiculate at apex. 

3. Shrubs with underground horizontal stems and numerous erect, aerial stems with deterrninate 

growth reaching 30 to 40 cm in height. 

4. Leaflets abaxially glabrous or with appressed yellowish hairs; dichasia sessile or short 

pedunculate; petals oblanceolate; disc tomentulose . ........................................................ T angustifolia 

4. Leaflets abaxially albo-sericeous; dichasia long-pedunculate; petals elliptic; disc hirsute ..T. subalbens 

3. Trees or treelets without underground horizontal stems, with one main trunk. 

5. Inflorescence fasciculate, axillary, ramiflorous or cauliflorous. 

6. Stems velutinous; leaflets with clathrate-reticulate tertiary venation ............................... T velutina 

6. Stems pubescent or puberulent; leaflets with clathrate tertiary venation ...................... T clathrata 

5. Inflorescence paniculate or racemiform, axillary or terminal, never cauliflorous nor fasciculate. 

7. Inflorescence racemiform. 

8. Stems puberulent; petals adaxially papillate; disc hispidulous; anthers barbate at 

apex ............................................................... T praealta 

8. Stems glabrous; petals adaxially villous; disc glabrous; anthers glabrous at apex ...... T esculenta 

7. Inflorescence paniculate. 

9. Leaflets abaxially glabrous. 

10. Stems puberulent; filaments woolly; anthers elliptic ....................................... T parviflora 

10. Stems glabrous; filaments glabrous; anthers ovate. 

11. Petals elliptic, adaxially glabrous, abaxially woolly-pubescent on lower half; 

appendage elliptic; fruit tomentose .................................................. T coriacea 

11. Petals oblong-ovate, adaxially papillate, abaxially glabrous or puberulent; 

appendage lanceolate; fruit glabrous ........................................................... T chartacea 

9. Leaflets abaxially puberulent, appressed-pubescent, or albo-sericeous. 

12. Leaflets abaxially puberulent. 

13. Leaflets often with adaxially raised conical insect galls or necrotic round spots; 

the apex obtuse, retuse or less often nearly acuminate . .......................... T simaboides 

13. Leaflets without insect galls or necrotic spots, the apex long acuminate or 

caudate ........................................................7'. japurensis 

12. Leaflets abaxially appressed-pubescent or albo-sericeous. 

14. Leaflets abaxially albo-sericeous, often cordate at base . .......................... T subalbens 

14. Leaflets appressed-pubescent, never cordate at base. 

15. Stems glabrous; leaves with 6-12 leaflets; leaflets abaxially appressed 

ferruginous-pubescent; leaf rachis sharply angled, sparsely tomentose. 

..................................................................................................................... T veraluciana 

15. Stems ferruginous-sericeous; leaves with 4-8 leaflets; leaflets abaxially 

appressed-pubescent; leaf rachis terete or obtusely angled ............... T furfuracea 

2. Anthers obtuse or retuse at apex. 

16. Stems tomentose or sericeous-tomentose. 

17. Leaflets abaxially glabrous or puberulent, prominent intramarginal vein lacking ............ T floresii 

17. Leaflets abaxially ferruginous-woolly, prominent intramarginal vein present ................. ... Tl anata 

16. Stems glabrous or puberulent. 

18. Leaves with only 2 leaflets ......................................................................' T squarrosa 

18. Leaves with 2-8 leaflets (some leaves with more than two leaflets always present). 

19. Disc tomentose or villose; petioles 1.7-8 cm long ............................. .... T firma 

19. Disc glabrous or puberulent; petioles 1.2-3(4) cm long ......................................... T microphylla

5 2 FLORA NEOTROPICA 

Fig. 37. Talisia angustifolia. A. flowering branch. B. staminate flower. C. abaxial and lateral views of petal with 

adnate appendage. D. staminate flower with perianth removed showing disc, stamens and detail of anther (right), 

I.s. same (left). E. infructescence. (A-D from Toledo & Gehrt 43167; E from Kuhlmann 59056). 

1. TALISIAANGUSTIFOLIA Radlkofer, Sitzungsber. 

Math.-Phys. Cl. Konigl. BayerAkad. Wiss. Miinchen 

8: 345. 1878. Type. Brazil. Goials: Without locality 

or date (fl), Burchell 6195 (holotype, B-destroyed; 

isotype, K; frag. at M). Figs. 3a-b, 37a-e 

Talisia pygmaea Radlkofer, Bull. Herb. Boiss. ser. 2, 

362. 1907. Type. Paraguay. Caaguazu, vic. of Yhu 

river, Oct 1905 (fl, fr), Hassler 9503 (holotype, 

B-destroyed; isotypes, K, MO, NY, P; frag. at M). 

Talisia humilis Mattos, Loefgrenia 70: 3. 1976. Type. 

Brazil. Sao Paulo: Itirapina, 13 Sep 1962 (fl), 

Felippe 43 (holotype, SP; isotypes, K, US-2). 

2mm 

Shrub with underground horizontal stems and numerous 

erect, aerial stems with determinate growth reaching 

40 cm in height, and forming a dense clump reaching 

3 m across. Underground stems are thicker than the 

aerial ones, producing scattered adventitious roots; aerial 

stems striate, appressed ferruginous-puberulent, mature 

stems lenticellate. Leaves paripinnate or less often 

imparipinnate; distal process filiform, 2-5 mm long; leaflets 

6-16, altemate or opposite, lanceolate or oblong-elliptic, 

(3) 4.5-15 x 1.2-2 (-4.2) cm, coriaceous, the 

adaxial surface glabrous, dull or slightly shiny, the abaxial 

surface densely to sparsely covered with appressed,


yellowish hairs, or glabrous, the venation brochido- Colaris 1086 (U); Sao Carlos, 10 Nov 1954 (fl), 

dromus, with primary and secondary veins prominent Kuhlmann 59056 (US); Campo Alegre, 24 Sep 1940 (fl), 

on abaxial surface, tertiary veins reticulate, the margins Toledo & Gehrt 43167 (US). 

entire, the apex acute or obtusely acuminate, the base PARAGUAY. Amambay: Estancia Primera, Feb 

1932 (st), Jorgensen 4594 (F, NY, US); 300 m, 10 Feb 

obtuse, slightly unequal; petiolules pulvinate, 1-1.5 mm 

1982 (fl, fr), Solomon et al. 6856 (MO, NY); Parque 

long, puberulent; rachis 3.5-22 cm long, terete, bisulcate 

Nacional Cerro Cori, in front of Cerro Muralla, 11 Dec 

along adaxial surface, puberulent; petioles 1.5-8 cm long, 1989 (fl), Vanni et al. 1308 (MO). 

flattened-canaliculate along adaxial surface, puberulent, 

slightly enlarged at the base. Thyrses panicle-shaped or Talisia pygmaea was treated as distinct from T anwith 

few lateral branches, axillary or terminal, to 23 cm gustifolia by Radlkoferbecause of its cinnamomeus (yellong; 

cataphylls wanting; axes to 15 cm long, angled, lowish brown) branches (vs. subferruginous light redstriate, 

appressed ferruginous-pubescent and usually fur- dish brown), its ovate-lanceolate leaflets with sessile or 

furaceous; bracts subulate, persistent, 2-2.5 mm long; decurrent bases and glabrous adaxial surface (vs. landichasia 

simple or compound, sessile or shortly pedun- ceolate, petiolulate and sparsely pilose) and its fewculate; 

pedicels 1.5-2.5 mm long, articulate atupperthird. branched thyrses (vs. many-branched). However, none 

Calyx 2.5-3 mm long, tomentulose to tomentose, the of these characters have proven to be consistent nor do 

sepals free to the base, deltate or deltate-elliptic, concave, they correlate with each other; they represent extremes 

obtuse at apex; petals oblanceolate, 4-4.5 mm long, erect in variation of these characters. Furthermore, differences 

at anthesis, sparsely papillate and puberulent on adaxial in pubescence on many of the specimens examined is 

surface, puberulent on lower third of abaxial surface, the age related. Talisia subalbens Radlkofer seems to be 

apex apiculate, the base clawed-attenuate; appendages closely related to T angustifolia, due to its similar habit 

as long as the petals, adnate to petal along half of their and general appearance. However, T subalbens can be 

length, lanceolate, erect, villose on adaxial surface, pu- distinguished from T angustifolia by the subcordate or 

berulent-papillate on abaxial surface; disc cup-shaped, cordate base of the leaflets. Talisia veraluciana Guarim 

5-lobed, tomentulose, ca. 0.5 mm tall; stamens 8, the fila- Neto is morphologically very similar to T angustifolia, 

ments of unequal length, 0.6-2.3 mm long, pubescent, however, T. veraluciana can be distinguished from it by 

the anthers 0.9 mm long, ovate, glabrous, apiculate at its arboreal habit and by its larger flowers. 

apex; ovary ovoid, ferruginous, tomentulose, the stigma 

trigonous, tomentulose, papillate. Fruits ovoid, densely 

ferruginous-tomentose, 1.8-2 x 1-1.7 cm, depressed at 2. TALISIA CHARTACEA Acevedo-Rodriguez, 

apex, with a short apiculum, the pericarp 2.8-3.2 mm BioLlania Edicion Esp. 6: 144. 1997. Type. Venethick, 

the endocarp glabrous. Seed one per fruit, known zuela. Bolivar: Municipio Raul Leoni, humid forest 

only in immature stage. 

on consolidated sandy grounds, 3 km from foothills 

The specific epithet refers to the distinctive narrow of Cerro Camaron (SW of Guaiquinima Tepui comleaflets 

of this species. 

plex), 42.5 km SE of Entre Rios, 250 m elevation, 

ca. 5?43'N, 64?07'W, 30 Oct-2 Nov 1988 (fr), G. 

Phenology. Flowers from August to February and 

Aymard & A. Fernandez 7176 (holotype, PORT; 

fruits in October, January and February. 

isotypes, MO, NY, US). Fig. 38a-e 

Distribution and Ecology. (Fig. 41) Brazil and 

Paraguay, in sandy cerrado, campo-cerrado, campo- Tree to 30 m tall. Stems terete, smooth, puberulent to 

limpo, and open grassy fields and savannas. glabrous at maturity, dark brown to grayish. Leaves 

paripinnate; distal process early deciduous, leaving a trun- 


cate base ca. 2 mm long; leaflets 4, altemate to opposite, 

Goihs: Anapolis, 20 Nov 1975 (fl), Hatschbach 37742 

(M); Serra do Caiap6, 60 km S of Caiaponia on road to lanceolate to elliptic-lanceolate, 6-13 x 2.5-5.7 cm, chart- 

Jatai, 800-1000 m, 27 Oct 1964 (fl), Irwin & Soderstrom aceous, concolorous, glabrous on both surfaces, slightly 

7451 (K, NY, US), 30 Oct 1964 (fr), Irwin & Soderstrom shiny on adaxial surface, the venation brochidodromus, 

7581 (IAN, MO, NY, P, US). Mato Grosso do Sul: Nova prominulous on both surfaces especially the midvein, 

Andradinha, 11 Nov 1975 (fl), Hatschbach 37393 (MO, tertiary venation reticulate, the margins slightly undu- 

NY); Tucuru; 16 Dec 1983 (fl), Hatschbach & Callejas late, the apex long-acuminate or less often obtuse, the 

47313 (US); Maracaju, 20 km W, 25 Oct 1988 (fl), base obtuse or acute-obtuse and slightly unequal; peti- 

Hatschbach & Cervi 52622 (US). Par*: Xingu river; 

olules slender, adaxially canaliculate, 0.7-1 cm long, 

Vict6ria, along road to Ponta Nova, 7 Aug 1918 (fl), 

Ducke 17176 (MG). Parani: Campo Mourao, 14 Oct 

glabrous or puberulent; rachis 2-3(4) cm long, slender, 

1965 (fl), Hatschbach 13002 (MO, NY, P, US, WIS), terete to slightly angled, faintly striate, puberulent to gla- 

25 Jan 1967 (fl, fr), Hatschbach 15911 (NY, P, US). Sio brous; petioles 3-5(6.5) cm long, slightly flattened ad- 

Paulo: Uba, along road to Sao Carlos, 12 May 1969 (st), axially, slightly thickened at base. Thyrses paniculate,


3 cm. 

Fig. 38. Talisia chartacea. A. flowering branch. B. staminate flower. C. lateral, adaxial, and abaxial views of petal 

with adnate appendage. D. staminate flower with perianth removed showing disc and stamens. E. fruiting branch. 

(A-D from Croizat 44; E from Aymard & Fernandez 7176).


axillary or terminal on lateral branches, to 13 cm long,; cate; leaflets (6) 8-10 (-12), alternate, subopposite or 

cataphylls minute, early deciduous; axes slightly angled, opposite, 9-24 (44) x (2.5) 5-9 (-12) cm, elliptic, obpuberulent 

or glabrous; bracts triangular, early deciduous, long or oblanceolate, chartaceous to coriaceous, the 

0.5-0.7 mm long; dichasia simple or compound, nearly adaxial surface glabrous, with prominent midvein and 

sessile; pedicels ca. 2 mm long, articulate at the middle. slightly impressed secondary veins, the abaxial surface 

Calyx 3-3.5 mm long, tomentulose, the sepals free to puberulent along midvein or glabrous, the venation 

the base, oblong-ovate, concave, rounded at apex; petals brochidodromus, midvein and secondary veins promioblong-ovate, 

3.5-4 mm long, erect, papillate on adaxial nent, secondary veins alternate or subopposite, arched, 

surface, glabrous to puberulent on abaxial surface, the forming a 20? 40? angle with the midvein and a strong 

apex obtuse, the base attenuate; appendages as long as submarginal loop, tertiary veins clathrate, the margins 

the petals, lanceolate, erect, obtuse at apex, adnate at base entire or wavy, revolute, the apex acuminate to caudate, 

along 1/4 of the petal length, woolly on both surfaces; usually mucronate, the base attenuate or obtuse, asymdisc 

cup-shaped, 5-lobed, tomentose, 1 mm tall; stamens metrical; petiolules nearly cylindrical, 6-15 mm long, 

8, the filaments of unequal length, 1.2-2 mm long, gla- glabrous or puberulent, wrinkled when dry; rachis 19brous, 

the anthers ca. 1 mm long, ovate, glabrous, api- 36 cm long, puberulent, striate, adaxially bicarinate on 

culate at apex; ovary not seen. Fruits ellipsoid, glabrous, distal portion, slightly flattened on proximal portion; 

purplish green, 3.5-3.7 cm long, smooth, mammiliform petioles striate or less often lenticellate, adaxially flatat 

apex, the pericarp 1-2 mm thick, the endocarp gla- tened. Thyrses fasciculate, cauliflorous or ramiflorous; 

brous. Seed one per fruit, known only in immature stage. cataphylls acicular, 2-4 mm long; axes 2-7 cm long, 

Talisia chartacea seems to be most closely related angled, ferruginous-pubescent; bracts subulate, persisto 

Tfirma Radlkofer in that both possess similar floral tent, ca. 2 mm long, with same indumentum as the axis; 

morphology and 4-foliolate leaves. However, T. dichasia simple or compound, nearly sessile; pedicels 

chartacea differs from Tfirma by having chartaceous, ca. 1 mm long, articulate at upper third. Calyx 2.5-3.2 

concolorous, abaxially shiny leaflets (vs. coriaceous, mm long, the sepals free nearly to the base, elliptic, 

discolorous, abaxially dull leaflets), a slender, greenish obtuse at apex; petals elliptic or less often ovate, obto 

light brown leafrachis (vs. robust andblackishbrown), tuse at apex, (3-) 4-5.8 mm long, reflexed at anthesis, 

and glabrous fruits with coriaceous (ca. 1 mm thick) abaxially glabrous; appendages as long as the petals, 

pericarp (vs. flavido-tomentose with semi-woody, 2-2.2 elliptic, erect, abaxially puberulent and papillate, adaximm 

thick pericarp). The specific epithet refers to the ally villous; disc cup-shaped, 5-lobed, ca. 1 mm tall; 

chartaceous texture of the leaflets. 

stamens 8, the filaments in two or three sets of unequal 

lengths, the shorter 2-2.5 mm long, the longer 4-5 mm 

Phenology. Collected in flower in December and July, 

long, glabrous, the anthers ca. 0.6 mm long, ovate, glaand 

in fruit inAugust and late Octoberto earlyNovember. 

brous, apiculate at apex; ovary ovoid, densely sericeous, 

Distribution and Ecology. (Fig. 41) Known from the stigma elongated, trigonous, pubescent. Fruits el- 

Venezuela and Brazil in terra firme forest, humid for- lipsoid to brownish-yellow, nearly smooth, densely seriest, 

and along seasonally flooded river banks. ceous but glabrous at maturity, 2-3 x 1.1-2 cm long, 

Representative specimens examined. VENEZUELA. obtuse at apex, with short apiculum, the pericarp 1-3 

Amazonas: Between Atabapo river and La Esmeralda, mm thick, the endocarp granulate, glabrous. Seed el- 

3 Jul 1951 (fl), Croizat 44 (F). Bolivar: SW of Icabaru, lipsoid, 1-1.3 cm long, with sweet, white sarcotesta. 

along Los Brazileros creek, 480 m, 16 Dec 1978 (fl), Embryo with cotyledons superimposed, of equal size. 

Steyermark et al. 117702 (VEN). 

The specific epithet refers to the tertiary venation 

BRAZIL. Amazonas: Watoriketheri village, Perimetral which is perpendicular to the secondary veins. 

Norte Hwy, km 21 1, Aug 1994 (fr), Milliken 2031 (K). 

Key to the subspecies of Talisia 

clathrata. 

3. TALISIA CLATHRATA Radlkofer, Sitzungsber. 


8: 349. 1878. Type. Brazil. Without specific locality, 

without date (fr), Martius s.n. (holotype, M, photo 

1. Calyx ferruginous-tomentulose or less often 

whitish puberulent; petals abaxially appressedpubescent 

on lower half, disc ferruginoustomentose; 

leaflets coriaceous, oblanceolate, 

(F neg. 6019) at NY, US; isotype, M). 

with secondary veins collected near the 

Tree, (4-)10-30 m tall; trunk to 20 cm in diam. 

Young stems striate, pubescent or puberulent, becoming 

terete and lenticellate with age. Leaves paripinnate 

margin, the basal secondary veins sometimes 

forming an acute angle with the 

midvein Talisia clathrata subsp. clathrata 

1. Calyx sericeous-canescent; petals abaxially 

or less often imparipinnate; distal process minute, trun- glabrous; disc canescent-tomentulose on upper


portion; leaflets chartaceous, elliptic, with second- 

ary veins forming a marginal loop 5-10 mm from 

the margin ..... Talisia clathrata subsp. canescens 

3a. TALISIA CLATHRATA subsp. CLATHRATA 

Figs. 6b, 39a-g 

Leaflets coriaceous, oblanceolate or less often elliptic, 

7.5-30 (44) x 3-9 (12) cm, with secondary veins 

collected near the margin, the basal secondaries veins 

sometimes forming an acute angle with the midvein, 

the apex abruptly acuminate, the base obtuse or attenuate. 

Calyx ferruginous-tomentulose or less often canescent-puberulent; 

petals elliptic or less often ovate, 

abaxially appressed-pubescent at lower half; disc ferruginous-tomentose. 

Phenology. Collected in flower from December to 

August and in fruit from May to November. 


Colombia, Peru and Brazil in understory of primary, 

non-flooded, moist, mature forest on clayish and sandy 

soils, primary terra firme forest, seasonally flooded 

tahuampa forest, caatinga forest on clayish-sandy soils, 

igapo forest on clayish soils, dense forest and adjacent 

campina on white-sand soils. 


Amazonas: Leticia, along trail to Tarapaca, 7 Apr 1975 

(bd), Cabrera 3329 (COL). 

PERU. Loreto: University Arboretum near Puerto 

Almendras, 27 Jul 1972 (fl), Croat 18550 (MO); vic. of 

Llanchama lake, close to Nanay river, 2 Aug 1972 (fl), 

Croat 18759 (F, MG, NY, USM); Zungaro Cocha near 

Iquitos, 150 m, 5 Nov 1964 (st), Dodson & Torres 2924 

(MO); 4 km S of Mishana, 13 Aug 1980 (fl), Foster 4429 

(F, U); Yanamono Explorama Tourist Camp, between 

Indiana and mouth of Napo river, 120 m, 28 Aug 1983 

(st), Gentry et al. 43759 (MO); Alpahuayo, km 20 

Iquitos-Nauta, 130 m, 17 Feb 1987 (bd), Gentry et al. 

56030 (F, MO); Iquitos-Nauta road, km 30, 140 m, 11 

Feb 1989 (st), Gentry et al. 65604 (MO); Iquitos, along 

Penia Negra road, 100 m, 3 Aug 1929 (fl), Killip & Smith 

27024 (F, NY, US); Mishuyacu near Iquitos, 100 m, Oct- 

Nov 1929 (fr), Klug 18 (F, US), Jan 1930 (fl), Klug 811 

(F, NY, US); Santa Maria de Nanay, along Nanay river, 

80-110 m, 30 Sep 1990 (fr), Pipoly et al. 12630 (US); 

Iquitos, along Penia Negra road, 120-140 m, 13 Jun 

1984 (fl), Rimachi 7499 (US, USM); Mun. Iquitos. 

Caserio Nina Rumy, Nanay river, 123 m, 6 Feb 1986 

(st), Ruiz & Balseca 816 (MO); Iquitos-Nauta road, km 

50, 22 Dec 1988 (fl), Ruiz & Melendez 1269 (K-2); 

Mazan river, Gamitanacocha, 100-125 m, 1 Feb 1935 

(fl), Schunke 303 (F-2, NY, US-2); Nauta, 160 m, 2 Jun 

1984 (fr), Vasquez & Jaramillo 5070 (MO); Rio Blanco 

Biological Station, 150 m, 19 Sep 1985 (fr), Vasquez et 

al. 6759 (MO); Canchahuayo (Ucayali river), 500 m, 

25 Nov 1985 (fr), Vasquez et al. 6918 (MO, NY); Puerto 

Almendras, 2 Oct 1986 (fr), Vasquez et al. 8068 (MO, 

NY),122 m, 16 Jun 1989 (fl), Vdsquez & Soto 12346 

(US, USM); Iquitos, along road Iquitos-Nauta, 130 m, 

23 May 1990 (fr), Vasquez et al. 13745 (US). 

BRAZIL. Acre: Basin of JuruA river, Humaita creek, 

22 Mar 1992 (fl), Daly et al. 7571 (US); Cruzeiro do Sul, 

vic. new airport, 17 Feb 1976 (fr), Monteiro & Damido 

461 (INPA, MG); Mun. Cruzeiro do Sul. without spe- 

cific locality, 20 Feb 1976 (fr), Monteiro & Damido 525 

(INPA). Amazonas: Vila Bittencourt, Japura river, 17 

Nov 1982 (fr), Amaral et al. 543 (INPA, NY, US). 

3b. TALISIA CLATHRATA subsp. CANESCENS 

Acevedo-Rodriguez, subsp. nov. Type. French 

Guiana. Region de Saul: Eaux Claires; Sentier 

Botanique, 2-5 kln from entrance, non-flooded moist 

forest on gentle slopes, 250 m, 3?37'N, 53?12'W, 

24 May 1992 (fl), P. Acevedo-Rodriguez et al. 5012 

(holotype, US; isotypes, A, CAY, CTES, F, HUA, 

K, NY, P, U). Figs. 4b, lOa, 40a-g 

A Talisia clathrata subsp. clathrata calice 

canescentibus, 

petalis glaberis differt. 

Leaflets chartaceous, elliptic or seldom oblanceolate, 

8.5-20.5(35) x 2.4-5-8 (9.5) cm, with secondary veins 

forming a marginal loop 5-10 mm from the margin, 

the apex long-acuminate, the base obtuse. Calyx sericeous-canescent; 

petals abaxially glabrous; disc canescent-tomentulose 

on upper portion. 

The newly described subspecies differs from the 

typical subspecies by the characters shown in the key 

to the subspecies. The subspecific epithet refers to the 

canescent sepals of this subspecies. 

Phenology. Flowers from May to July, and fruit- 

ing from June to November. 

Distribution and Ecology. (Fig. 41) Non- 

flooded, moist lowland forest in Guyana, French 

Guiana and Brazil. 

Common name. Guyana: Sand mora, wa-i. 

Representative specimens examined. GUYANA. 

Barima/Waini: Barama River, Kariako, 145 m, 25 Sep 

1996 (st), van Andel 26sls5 (US). 

FRENCH GUIANA. Bassin de L'Approuague: 

Regina. Eastern plateau of Montagne Tortue, 11 km WNW 

of Approuague river, 16 Jun 1988 (fl), Feuillet et al. 

10087 (NY, US); Station des Nouragues, 16 Jul 1989 (fr), 

Sabatier & PrJvost 2792 (CAY, NY, P, US), 24 Jun 1994 

(st), Sabatier & Prevost 4237 (CAY, US). Bassin du 

Maroni: Antecume Pata, Nov 1975 (fr), Moretti 364 

(CAY-2); Grand Inini river, 26 Jul 1990 (st), Sabatier & 

Prevost 3433 (CAY), 26 Jul 1990 (st), Sabatier & Prevost 

3436 (CAY). Region de Saul: Route de Belizon, ca. 2 km 

S from Eaux Claires, ca. 200 m, 19 May 1992 (st), Acevedo 

R. et al. 4956 (US), 26 May 1992 (fl), Acevedo R. & Angell 

5021 (CAY, US-2); Layon Eaux Claires, Sentier Botanique, 

ca. 700 m from Route de Belizon, 230 m, 22 May 1992 

(bd), Acevedo R. et al. 4996 (CAY, US); Montagne La


3 mm. 

t t X ~~~ ~~~~~2 mm. ~ 

Fig. 39. Talisia clathrata subsp. clathrata. A. sterile branch. B. staminate flower. C. lateral and adaxial views of 

petal. D. staminate flower with perianth removed showing disc and stamens. E. infructescence. F. embryo, lateral 

view. (A from Killip & Smith 27024; B-D from Schunke 303; E-F from Monteiro & Damido 461).


2mm~~~~~~~~~~~~~~~~ 

X tE. 

Fig. 40. Talisia clathrata subsp. canescens. A. flowering branch. B. staminate flower. C. adaxial and lateral 

views of petaf with adnate appendage. D. staminate flower with perianth removed showing disc and stamens (left) 

and l.s. same (right). E. top views of disc. F. infructescence. G. embryo, lateral view. (A-E from Mori et al. 21365; 

F-G from Mori et al. 14961).


9.MwI O.- 7WNI 60W 50W! 40W 

N~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~O 

X,~~> a/, _1'" 

"A 

Fi. 1 Dsriuio f peis nTaiiasbgnu itmara(i ar) 

f*. 

A Talisia chartacea \ L ; 

* Talisia clathrata spp. clathrata ............. . 

tj 

X t - * ' 

Talisia clathrata ssp.canescens , ,< , 

Fig. 41' DistributionofspeciesinTalisiasubgenusPito i/a 0n p 

~~~~~~~~~~~~~~ 

Fumee, 200-400 m, 21 Sep 1982 (fr), Mori & Boom 14961 glabrous on both surfaces, the venation brochidodromus, 

(CAY, MO, NY), 16 Aug 1982, (fr), Mori & Boom 14711 

(MO, NY); Layon Boeuf Mort, 250-300 m, 15 Jun 1988 

(yfr), Mori & Gracie 18951 (CAY, NY, P); Saul, 18 May 

1973 (fl), Oldeman 3304 (CAY, MO, NY, US). 

BRAZIL. Amazonas: Manaus. Disto. Agropecuario, 

BDFF, km 41, 50-125 m, 12 Jul 1990 (fl), Mori et al. 

21365 (US). 

the midvein plane or sunken on adaxial surface, prominent 

on abaxial surface, secondary veins inconspicuous, 

or less often prominulous on abaxial surface, tertiary 

veins reticulate, the margins entire or slightly undulate, 

slightly revolute, the apex acute to long-acuminate, glandular- 

mucronulate, or less often obtuse, obtusely apiculate, 

retuse, or nearly rounded, the base rounded, obtuse 

or acute, sometimes asymmetrical or slightly so; 

4. TALISIA CORIACEA Radlkofer, Sitzungsber. petiolules canaliculate, 2-5 mm long, puberulent or gla- 

Math.-Phys. Cl. K6nigl. Bayer Akad. Wiss. brous; rachis 0.7-2.5 cm long, slightly flattened, striate, 

Munchen 8: 346. 1878. Type. Brazil. Bahia: Ilheus, lenticellate, puberulent; petioles 2-5 cm long, slightly 

1840 (fl), Luschnath s.n. (holotype, B-destroyed, flattened, lenticellate, canaliculate, wider at base. Thyrses 

photo (F neg. 5673) at MO, NY, US; isotype, BR- 

2 sheets, photos at US). Figs. lb, 42a-d 

paniculate, terminal, to 15 cm long, the branches to 10 

cm long; cataphylls lacking; axes angled, sulcate, minutely 

tomentose; bracts minute, early deciduous, mi- 

Tree 7-30(?) m tall; trunk to 140 cm in diam. Stems nutely tomentose; dichasia simple, sessile, sometimes 

terete to obtusely angled, puberulent, becoming glabrous reduced to a single flower; pedicels ca. 1.5 mm long, arand 

lenticellate with age. Leaves paripinnate or less of- ticulate below the middle or at the base. Calyx 2.3-3.2 

ten imparipinnate; distal process truncate, minute; leaflets mm long, tomentose, the sepals free to the base, concave, 

(2) 4-6, opposite or alternate, elliptic to lanceolate, or ovate, obtuse at apex; petals elliptic, 3.2-5.5 mm long, 

less often oblanceolate, 2.5-10 x 1.2-4.5 cm, coriaceous, reflexed, woolly-pubescent on lower abaxial surface,


t * 

~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~2 -mlM 

cm.D 2;mB X 

Fig. 42. Talisia coriacea. A. flowering branch. B. staminate flower and flower with part of perianth cut away 

showing disc and stamens. C. adaxial and abaxial views of petal with adnate appendage. D. flower with perianth 

removed showing disc, stamens, and pistillode. (All from Luschnath 1840). 

adaxially glabrous, ciliate at margins, the apex obtuse, bryo with cotyledons superimposed, of equal size. 

the base tapering; appendages slightly shorter than the The specific epithet refers to the coriaceous texture 

petals, elliptic, erect, densely woolly-pubescent adaxial- of the leaflets. 

ly, minutely woolly-pubescent on abaxial surface; disc 

cup-shaped, 5-lobed, minutely tomentose, ca. 1 mm tall; 

stamens 8, the filaments of unequal length, 0.8-2 mm 

long, glabrous, the anthers ca. 1 mm long, elliptic, glabrous, 

apiculate at apex; ovary nearly conical, tomen- 

Phenology. Flower from November to January and 

in May, collected in fruit in July. 

Distribution and Ecology. (Fig. 45) In tropical moist, 

terra firme forest in lowlands of Brazil and Venezuela. 

tose, the stigma obconical, tomentulose, papillate. Fruits Representative specimens examined. VENEellipsoid, 

tomentose, 2.2 cm long (immature), green, ZUELA. Amazonas: Metacuni river, 20 Jan 1990 (fl), 

shortly apiculate at apex, the pericarp 1 mm thick, woody. Stergios & Velazco 13978 (PORT, NY, US). Barinas: 77 

Seeds ellipsoid, ca. 1.7 cm long, with fleshy testa. Em- km from Barinas along road to San Cristobal, 350 m, 7 

2 mm.


L ~ ~ ~ L 

?.m2mL 7 

Fig. 43. Talisia esculenta. A. flowering branch. B. detail of leaflet apex showing venation. C. staminate flower. 

D. adaxial and abaxial views of petal with adnate appendage. E. staminate flower with perianth removed showing 

disc, stamens and pistillode. F. branch with infructescence. (A-E from Daly et al. 635; F from Nee & Coimbra 36952). 

May 1964 (fl), Breteler 3936 (F, NY, P, U, US, VEN-2). 

BRAZIL. Amazonas: Mun. Manaus. Manaus- 

Itacoatiara road, km 148, 18 May 1972 (fl), Loureiro 

et al. s.n. (INPA-35821, US), km 175, trail 23, 4 Jul 

1968 (fr), Rodrigues etal. 21329 (INPA). Bahia: Without 

specific locality, 1841 (fl), Luschnath s.n. (NY); 

1857 (fl), Martius 1851 (M). Mun. Una. Maruim, km 

33 road Oliven9a-Buerarema,I00 m, 13 May 198 1(st), 

Mori et al. 13975 (NY). Rio de Janeiro: Mun. Rio de 

Janeiro. Rio de Janeiro. Horto Florestal, [cultivated?] 15 

Dec 1931 (fl), Lourenqo 571 (INPA, U); road to Vista 

Chinesa, km 1, 24 Nov 1965 (fl), L. Sobre 1155 (US). 

5. TALISIA ESCULENTA (Cambessedes) Radlkofer, 

Sitzungsber. Math.-Phys. Cl. Konigl. Bayer Akad. 

Wiss. Miunchen 8: 345. 1878.-Sapindus esculentus 

Cambessedes in Saint-Hilaire, PI. Usuel. Bras. t. 68. 

Aug. 1828. Type. Brazil. Minas Gerais: Sertao du 

Rio Sao Francisco, 1824 (fl), Saint-Hilaire s.n. (holotype, 

MPU; isotype, P). Fig. 43a-f


Treelet or tree, 5-10 (20) m tall; trunk to 50 cm in Distribution and Ecology. (Fig 45) Brazil, Paradiam.; 

bark brownish, flaky. Twigs nearly terete, gla- guay, and Bolivia. In gallery, terra firme, varzea, 

brous, lenticellate with age. Leaves paripinnate or less caatinga, and dry forests, island forest surrounded by 

often imparipinnate; distal process minute, filiform, 4- savanna, and disturbed secondary forest, also widely 

5 mm long, early deciduous leaving a truncate base; cultivated for its edible fruits (with fleshy testa). 

leaflets (2) 6 to 8, opposite to alternate, elliptic or less 

Common names and uses. Brazil. Without specific 

often oblong-elliptic, lanceolate-elliptic or oblanceolate, 

locality: Olhio de boi, pitomba de macaco (fide Pio 

4.5-15 x 1.7-7.5 cm, chartaceous, the adaxial surface 

Correa 1984, and Lorenzi, 1992); Amazonas: pitomba, 

slightly shiny, glabrous, the abaxial surface glabrous 

pitombeira; Bahia: feijao cru', pitomba; Maranhao: 

or puberulous, the venation brochidodromus, pitomba; Rio de Janeiro:pitombeira,pitomba da mata. 

prominulous on both surfaces, tertiary veins reticulate, This species is commonly cultivated for its edible fruit 

the margins entire, undulate, the apex obtuse, acute, ob- (with fleshy testa), which is sold in local markets. The 

tusely acuminate or sometimes emarginate, the base sap is employed as a fish poison and the wood is used 

obtuse, rounded or rarely truncate, slightly asymmetri- in the construction of boxes (Pio Correa 1984). 

cal; petiolules slender, 1-3 mm long, puberulent or 

pubescent, usually drying dark brown; rachis (1) 3.5- 

Note: There is a strong indication that the protologue 

13 cm long, obtusely angled, puberulent or pubescent; 

of this species is the work of Cambessedes. Firstly, in 

petioles (3) 4.5-6.5 (9) cm long, slightly flattened along 

the description of the pistillate flowers and the fruits 

Saint-Hilaire is credited as the source of this 

adaxial surface, thickened and darker at the 

informavery 

base, 

tion. It is unlikely that Saint-Hilaire would cite himself 

with same indumentum as the rachis. Thyrses simple, 

as the source of his description. Secondly, the locality 

racemiform, distal or axillary on lateral branches, 15information 

is written in the third person and the usage 

25 cm long; cataphylls acicular, 3-4 mm long, congiven 

for the plants refers again to Saint-Hilaire. Accordgested, 

axillary in young growth; axes nearly terete to 

ing to Taxonomic Literature II (Stafleu & Cowan, 1983) 

angled and sulcate, puberulent or pubescent; bracts 

parts 9-14 of this book were co-authored by 

oblong, 3-4 mm long with same indumentum as the 

Cambessedes and de Jussieu. Parts 1-8 were written 

axis; dichasia compound or less often simple; peduncles 

exclusively by Saint-Hilaire, and in contrast with the later 

flattened, 2-6 mm long, pubescent to tomentulose; 

parts, it is written in first person, indicating the differpedicels 

2.5-4 mm long, articulate at the lower third, 

ence of style between Saint-Hilaire and his co-authors. 

with same indumentum as the peduncle. Calyx 3.5-4 

Since Cambessedes wrote the treatnent of Sapindaceae 

mm long, appressed-ferruginous-tomentulose, the sefor 

Flora Brasiliae Meridionalis (1824-1833) of Saintpals 

free to the base or nearly so, ovate, concave, slightly 

Hilaire, it would be logical to assume that he was the 

unequal; petals elliptic or elliptic-lanceolate, 4-5 mm 

person responsible for the description of S. esculentus. 

long, reflexed at anthesis, white, villose adaxially, appressed-pubescent 

on lower half abaxially, the apex Representative specimens examined. BRAZIL. 

acute or obtuse, the base narrowed; appendages with 

Without specific locality, 1825-1830 (fr), Burchell 7971 

(K, P); without data (fl), Sello s.n. (K). Amazonas: 

same shape and length as the petals, puberulent abaxially, 

Cachoerinha-Igarape do 40, 4 Nov 1955 (fl), Mello 2275 

villous adaxially, adnate along margins of lower half of 

(INPA, MG). Bahia: Barreiras, along road to Santo 

the petal to form a pocket, the apex obtuse or bifid; disc Desiderio, 23 Dec 1954 (fr), Black 54-17714 (IAN, U); 

annular, 5-lobed, glabrous, ca. 0.6 mm tall; stamens 8, Formosa do Rio Preto, 13 Oct 1908 (fl), Dias et al. 117 

spreading, the filaments of unequal length, white, 3-4 (US); Mun. Itacare, Faz. Monte Alegre, along Contas 

mm long, pilose to glabrous, the anthers ca. 1.2 mm river, cultivated, 18 Aug 1996 (fr), Jardim 952 (NY, 

long, elliptic, glabrous, apiculate at apex; ovary coni- US); Alvorada, 270 km from Brasilia, Correntes river, 2 

cal, ferruginous-tomentose, the stigma cylindric-capitate, 

tomentulose, papillate. Fruits ovoid to nearly 

rounded, minutely puberulent and yellow when ripe, 

2-3 cm long, shortly apiculate at apex, the pericarp ca. 

1 mm thick Seeds 1-2 per fruit, ellipsoid, 1.7-2 cm 

long, with a fleshy, clear white cover. Embryo with 

Jul 1964 (fl), M. Pires 58132 (K, MO, NY, US). Cearal: 

Without specific locality, 13 Apr 1918 (fr), Curran 30 

(US); Serra de Maranguabe, 14 Sep 1908 (bd), Ducke 

1634 (MG); Without specific locality, Aug-Nov 1838 

(fl), Gardner s.n. (NY); Between Reden9ao & Palmacea, 

10 Oct 1980 (fl), Nunes & Martins 8996 (F); Serra de 

Baturite, Sep 1910 (fl), Ule 9064 (K, US). Goias: Withcotyledons 

superimposed, of equal size. 

out specific locality, 1896 (fl), Glaziou 20862 (K, P). 

The specific epithet refers to the fruits (edible, fleshy Maranhao: Between Viana & Bandeirantes, 17 Oct 1980 

seed coat) of this species. 

(fl), Daly et al. 635 (K, MG, NY, US); Mun. Tuntum, 

Palmeirinhas, between P. Duarte-Barro do Corda, 26 Feb 

Phenology. Flower from July to February and fruit 

from June to April. 

1983 (fr), Taylor et al. E-1036 (F, MG, MO, NY, US). 

Mato Grosso: San Luis de Caceres, Sep 1911 (fl), Hoehne


4507 (M); Cuiaba, 19 Sep 1902 (fl), Malme 2300 (US); 

without date (fl), Manso 273 (M); Serra do Roncador, 

50 km N of Chavantina, Vau river, 9 Oct 1964(fl), Prance 

& Silva 59317 (M, MO, NY, P, US). Minas Gerais: With- 

out specific locality, s.d. (fl), Claussen s.n. (K); Apr- 

Aug 1840 (fl), Claussen s.n. (P, K-2); 1838 (fl), Claussen 

500 (P); Mun. Varzea de Palma, along road from Varzea 

de Palma to Serra do Cabral, 16 Jan 1996 (fr), 

Hatschbach & Silva 64141 (NY, US); Reserva de Grande 

Sertao, Veredas, Rio Preto, 11 Jun 1989 (fr), Ratter et 

al. 6366 (NY, US); Sao Francisco river, Sep 1834 (fl), 

Riedel & Lund 2645 (NY, US). Para: Alcantara, 26 Sep 

1903 (bd), Ducke 424 (MG); Santa Izabel, 20 Oct 1908 

(fl), Anonymous, MG 9743 (MG-2); Concei9ao do 

Araguaia, 16 Aug 1955 (fl), Macedo 4048 (US). 

Pernambuco: Ipojuca, Praia do Cupe, 10 Dec 1970 (fl), 

Andrade Lima 70-6231 (F); Island of Itamarica, Dec 

1837 (fl, fr), Gardner 951 (F, K-2, NY, P-2); Tapera. 

without specific locality, Dec 1930 (fl), Pickel 146 (F); 

thicket, 9 Jan 1932 (fl), Pickel 306 (IAN, US); Brejo 

Madre de Deus, 4 Feb 1995 (fl), Rodal & Sales 460 

(US); Vicencia, Eng. Canavieiras, 10 Dec 1964 (fl), 

Teixeira 2590 (US). Piaui: Mun. Valencia, Faz. Experi- 

mental da Universidade de Piagaba, 30 Oct 1984 (yfr), 

Freire 3709 (US); Sertao, Oct 1838 (fl), Gardner 1501 

(K-2, NY, P-2); Parnagua, 16 Aug 1980 (fl), Santino 

279 (MG); Mun. Teresina, Teresina, 11 Oct 1985 (fl), 

Sousa et al. 4230 (US). Rio de Janeiro: Santa Cruz, 

road to Sepatiba, 22 Nov 1980 (fl), V Freire 100 (K); 

Without specific locality, Feb 1874 (fl), Glaziou 6491 

(K, P), Sep 1876 (fl), Glaziou 8309 (K, P), Nov 1879 

(fl), Glaziou 10424 (K, P). 

BOLIVIA. Beni: Ballivian. 63 km from Santa Rosa 

toward Puerto Teresa, 170 m, 17 Sep 1993 (fl), de Michel 

& Beck 1237 (LPB, US-2); Don Lorenzo, 200-400 m, 

27 Jan 1995(fr), Mostacedo et al. 2680 (LPB, USZ); Prov. 

Chiquitos, Cordillera Santiago, Sep (fl), Orbigny 894 (P), 

Orbigny 896 (P); Beni River, Jul 1886 (fr), Rusby 1390 

(NY, US). Santa Cruz: Prov. Velasco. Las Gamas, 850 

m, 15 Nov 1993 (fr), Arroyo et al. 282 (US); Camp El 

Refugio, 27 Jun 1994 (fr), Guillen 1954 (US, USZ), 30 

Jun 1994 (st), Guilltn & Coria 2005 (MO, USZ); Reserva 

Forestal Bajo Paragua, Cerro Pelao, 450 m, 19 Oct 1994 

(fr), Killeen et al. 7033 (US, USZ); Prov. Nuflo de Chavez. 

10 km E of Concepci6n, 500 m, 13 Dec 1994 (fr), Killeen 

& Jardim 7424a (LPB, US); Prov. Andres Ibafiez, 450 

m, 29 Nov 1988 (fr), Nee & Coimbra 36952 (US); El 

Venao, 29 Aug 1987 (fl), M. Saldias P. 43 (USZ); Prov. 

Velasco. San Juancito, 20 km N of San Ignacio, 1 Jan 

1990 (fr), Seidel 3148 (LPB, US); Prov. Sara, 350 m, 8 

Oct 1926 (fl), Steinbach 7604 (K, MO, NY), Buenavista, 

8 Oct 1924 (fl), Steinbach 6595 (K); Prov. Velasco. Parque 

Nacional N. Kempff Mercado, trail to WCS antenna, 250 

m, 2 Oct 1995 (fr), Vargas et al. 4008 (MO). 

PARAGUAY. Without specific locality, Sep 1901- 

1902 (fl), Hassler 7413 (K, MO, NY, P-2, US). Amambay: 

Estancia Tres Hermanos, 9 Dec 1991 (fr), Basualdo 3830 

(MO); Cerro Lorito II, 400 m, 11 Dec 1978 (fr), Bernardi 

19128 (F, NY); Apa river, 29 Jul 1910 (fl), Fiebrig 4178 

(K, M); Concepci6n. Sep 1901-1902 (fl), Hassler 7413a 

(K, MO, P, US); Bella Vista, 15 Dec 1983 (fr), Vanni et 

al. 282 (F, MO). Central: Asuncion, Botanical Garden 

& Zoo, cultivated, Oct-Nov 1991 (fl), Perez 1288 (US), 

Dec 1991 (fr), Perez 1436 (US). 

6. TALISIA FIRMA Radlkofer, Sitzungsber. Math.- 


346. 1878. Type. Venezuela. Amazonas: Casiquiare, 

Vasiva & Pasimoni rivers, 1853-54 (fl), Spruce 

3311 (holotype, B-destroyed; isotypes, K-3, NY, 

P-2; frag. at M). Figs. Sb, 9a, 12a, 44a-f 

Talisia multinervis Radikofer, Sitzungsber. Math.- 


346. 1878. Type. Brazil. Uapes river, vic. of 

Cachoeira Ipanore [as Panure], Oct 1852-Jan 

1853 (fl), Spruce 2421 (holotype, B-destroyed; 

isotypes, C, K-2, NY, P; frag. at M). 

Tree 10-20 (30) m tall, branching on upper portion; 

trunk to 27 cm in diam.; bark light grey, smooth. Twigs 

nearly terete to obtusely angled, sulcate, glabrous, 

lenticellate with age. Leaves paripinnate or less often 

imparipinnate; distal process filiform, ca. 4 mm long, 

early deciduous; leaflets (2) 4 (5), opposite, lanceolate, 

oblong or elliptic, 5.5-17 x 3-7.8 cm, subcoriaceous 

to coriaceous, glabrous, the adaxial surface slightly 

lustrous, with plane midvein, the abaxial surface dull, 

drying darker, with prominent midvein, the venation 

brochidodromus, prominulous on both surfaces, tertiary 

veins reticulate, the margins entire, undulate, the 

apex obtuse, acute to long-acuminate, the base asymmetrical, 

one side obtuse the other rounded; petiolules 

pulvinate, slender, 5-10 mm long, slightly flattened on 

adaxial surface, glabrous; rachis 1-4 cm long, nearly 

terete, bisulcate along adaxial surface, glabrous; petioles 

1.7-8 cm long, slightly flattened along adaxial 

surface, slightly enlarged at base, glabrous. Thyrse 

panicle-shaped, axillary or distal, 5-25 cm long; 

cataphylls absent; axes angled, sulcate, puberulent; 

bracts early deciduous, oblong-lanceolate, ca. 3 mm 

long, with same indumentum as the axis; bracteoles 

similar but smaller; dichasia simple to compound, sometimes 

only one flower developing; peduncles slightly 

flattened, 0-2 mm long, pubescent or tomentulose; 

pedicels 3-4 mm long, articulate at upper third, with 

same indumentum as the peduncle. Calyx green, 3-4.5 

mm long, puberulent to pubescent, the sepals free to 

the base, oblong-lanceolate to ovate, slightly concave; 

petals elliptic-deltate, lanceolate-deltate or deltate, 3.6- 

5.5 mm long, reflexed at anthesis, white, adaxially villose 

on upper half, abaxially appressed-pubescent on 

lower half, the apex acute or obtuse, the base cuneate; 

appendages similar to the petals in shape, as long as or


Fig. 44. Talisia firma. A. flowering branch. B. staminate flower. C. I.s. staminate flower showing disc, stamens, 

and pistillode. D. adaxial and abaxial views of petal with adnate appendage. E. stamens. F. portion of infructescence. 

(A-E from Liesner 8866; F from Gentry 47377).


SOW? SGAI TO~~7W? SOW SAN? 40W 


Delgado 8458 (PORT, NY); Isla Rat6n, 1971 (fl), Bossio base; leaflets 24, opposite or less often alternate, ovate, 

s.n. (US); Vic. of Manare, 28 km S of San Carlos de Rio lanceolate or elliptic, 5.5-11.5 x 2.6-6.6 cm, coriaceous, 

Negro, along Rio Negro, 119 m, 22 Apr 1981 (fl), Clark 

& Maquirino 7931 (MO, NY); Casiquiare river, El 

Porvenir, 13 Feb 1991 (fl), Colella & Guayamare 2138 

(US); Brazo Casiquiare, between the mouth of Pasimoni 

& Culimacare rivers, 80 m, 25 Jul 1984 (fl), Davidse 

27875 (MO), 26 Jul 1984 (fl), Davidse 27960 (MO); 

Mawarinuma river, 140 m, 27 Apr 1984 (st), Gentry & 

Stein 47010 (MO); Baria river, 130 m, 11 May 1984 

glabrous, the adaxial surface lustrous, the abaxial surface 

dull, drying lighter, the venation brochidodromus, 

with tertiary veins highly reticulate, these plane or 

slightly sunken on adaxial surface, prominent on abaxial 

surface, tertiary veins reticulate, the margins entire, undulate, 

revolute, the apex obtuse, rounded, acute or 

acuminate, the base rounded or obtuse, sometimes asym- 

(fr), Gentry & Stein 47377 (MO); Cafno de Cholo, 16 metrical; petiolules cylindrical, rugose, grayish, 5-10 

km NE of San Carlos de Rio Negro, 120 m, 29 Jan 1980 mm long, furrowed along adaxial surface, appressed- 

(fl), Liesner 8866 (MG, MO, VEN), 29 Jan 1980 (fl), puberulous or pubescent; rachis 1.5-7 cm long, nearly 

Liesner 8883 (NY, MO, VEN); Paciba river, between 

Lago Paciba & Casiquiare river, 130 m, 5 Apr 1953 (fl), 

Maguire & Wurdack 34869 (MO-2, NY, VEN); Vic. of 

Piedra Arauicaua, along Pacimoni-Yatua river, 130 m, 

16 Jul 1959 (fl), Wurdack & Adderley 43476 (MG, MO- 

2, NY, US, VEN); Near Solano along Casiquiare river, 9 

Jul 1959 (fl), Wurdack & Adderley 43363 (K, MG, MO, 

NY, P, US, VEN); Yatua river, 110 m, Apr 1991 (fr), 

terete, bisulcate along adaxial surface, appressed-puberulent, 

lenticellate; petioles 3-6.5 cm long, slightly 

flattened along adaxial surface, slightly enlarged at base, 

appressed-puberulent, lenticellate. Thyrses panicleshaped, 

axillary or distal, 10-20 cm long; cataphylls 

wanting; axes slightly flattened, sulcate, ferruginoustomentose; 

bracts early deciduous, oblong-lanceolate, 

Velazco 1745 (MO-2, PORT). 

ca. 4 mm long, with same indumentum as the axis; brac- 

PERU. Loreto: Prov. Maynas: Iquitos, Quistococha, teoles similar but smaller; dichasia compound; pe- 

100 m, 29 Oct 1979 (st), Ayala 2004 (MO); Mishana, duncles terete, 3-4 mm long, tomentose; pedicels < 1 

Nanay river, half way between Iquitos & Sta. Maria de 

Nanay, 140 m, 23 Mar 1979 (st), Gentry et al. 26114 

(MO); Yanamono Explorama Tourist Camp, 130 m, 1 Jul 

1983 (st), Gentry et al. 42476 (MO); Moronillos, vic. of 

Iquitos, 116 m, 15 Sep 1986 (fr), Visquez & Jaramillo 

7913 (MO, NY). 

BRAZIL. Amazonas: Mun. Barcelos. 10 km N of 

Barcelos, 100 m, 6 Aug 1996 (fl), Acevedo R. et al. 7963 

(INPA, US), (fl), Acevedo R. et al. 7988 (INPA, US); 

mm long, articulate at the apex, with same indumentum 

as the peduncle. Calyx yellowish, ca. 5 mm long, tomentose, 

the sepals 3-4 mm long, ovate-deltate, slightly 

concave; petals narrowly elliptic, 3.5-4.5 mm long, 

reflexed at anthesis, yellowish, adaxially glabrous, 

abaxially villose, the apex acuminate, the base clawed; 

appendages as long as the petals, adaxially villous, 

abaxially glabrous, adnate along margins of lower half 

Lateral channels of Rio Negro toward entrance to Araga of the petal to form a pocket, the apex acuminate; disc 

river, 100 m, 7 Aug 1996 (fl), Acevedo R. et al. 8053 annular, 5-lobed, glabrous, 1-1.5 mm tall; stamens 8, 

(INPA, US); Upper Rio Negro, 100 m, 16 Aug 1996 (fr), the filaments of unequal length, 3-4 mm long, glabrous, 

Acevedo R. et al. 8390 (INPA, US); Rio Preto, 17 Aug the anthers 1-1.2 mm long, ovate, glabrous, obtuse at 

1996 (yfr), Acevedo R. et al. 8448 (INPA, US); Vila 

Bittencourt, along Japura river, 19 Nov 1982 (fr), Amaral 

et al. 579 (INPA, MG, MO, NY, US); Cauaburi river, 18 

Jun 1976 (fr), Coelho 495 (INPA, MG); Mun. Barcelos. 

Along Araga river, 29 Jul 1985 (fl, yfr), Cordeiro 318 

(F, INPA, NY, US); Carauari, Pogo Jurua, Jul 1980 (fr), 

da Silva et al. 905 (MG); Mun. Manaus. Reserva Ducke, 

Manaus-Itacoatiara, km 26, 7 Aug 1995 (fr), Sothers et 

apex; ovary conical, ferruginous-tomentose, the stigma 

cylindric-trigonous, tomentulose. Fruits asymmetrically 

depressed-ovoid, woody, densely ferruginous-sericeous, 

brown when ripe, 3.5-4.2 x 4.5-5 cm, the pericarp 

2-5 mm thick Seeds one per fruit, ovoid, ca. 3 cm 

long, with a fleshy white cover. Embryo with cotyledons 


al. 539 (US). 

The specific epithet honors Dr. Roman S. Flores, 

collector of the type specimen. 

7. TALISIA FLORESII Standley, Tropical Woods 

26: 14. 1931. Type. Mexico. Yucatain: Progreso, 

Mar 1931 (Fr), R. S. Flores s.n. (holotype, F 

633290). Fig. 46a-h 

Tree 6-30 m tall, branching on upper portion; trunk 

to 1 m in diam. Twigs nearly obtusely angled, sulcate, 

ferruginous-tomentose, becoming terete, glabrous, and 

noticeable lenticellate with age. Leaves paripinnate; distal 

process filiform, early deciduous, leaving a truncate 

Phenology. Known to flower from April to August 

and to fruit in February, July and November. 

Distribution and Ecology. (Fig. 50) Endemic to 

theYucatan Peninsula (Guatemala and Mexico), in tall, 

moist and rain forests. 

Common name and uses. Guatemala. Peten: Poloc. 

Fruits are reported as edible (Standley, 1931); dry fruits 

are used as rattles and the woody mesocarp is used in 

the construction of toys, similar to a whirligig.


3mm.~~~~~~~~~~~~~~~~~H 

LI 7D WG 

(7 

(1)~~~~~~~~~~~~/1 

Fig. 46. Talisia floresii. A. flowering branch. B. dichasium. C. staminate flower. D. adaxial and abaxial views 

of petal with adnate appendage. E. staminate flower with part of perianth cut away showing disc and stamens. 

F. flower with perianth removed showing disc and stamens (left) and ls. same (right). G. pistillode. H. portion of 

infructescence. (A-G from Bartlett 12680; H from Enriquez 392).


Representative specimens examined. MEXICO. branches to 12 cm long; cataphylls absent; axes slightly 

Campeche: Calakmul, Zon-Laguna, flooded forest, 15 obtusely angled, sulcate, ferruginous-papillate inter- 

May 1997 (fl), Alvaro 721 (CICY); Calakmul, tall, moist mixed with soft, long hairs; bracts persistent, oblongforest, 

28 Jun 202 (fr), Acevedo-Rdgz. & Tapia 12234 lanceolate, ca. 4 mm long, with same indumentum as 

(CICY, US); La China, 1.5 km SE of Campeche, remnant 

the axis; bracteoles similar but smaller; dichasia simple, 

tree along roadside, 27 Jun 2002 (fr), Acevedo et al. 12200 

sessile; pedicels 1.5-2 mm long, articulate on lower half, 

(CICY, US); Tuxpefia, 8 Nov 1931 (fr), Lundell 917 (K, 

MO, US). Quintana Roo: 2 km N of Laguna Guerrero, with same indumentum as the axis. Calyx pale green, 

1 Jul 1984 (fr), Cabrera & Cabrera 6569 (F); Othon 4-4.5 mm long, papillate-sericeous to tomentose, the 

Blanco, 4 km NW of Estero Franco, flooded forest on sepals ca. 3 mm long, ovate to elliptic, slightly concave; 

organig soils, 17 May 1985 (fl), Cabrera 8360 (CICY). petals elliptic-ovate, 4.5-6 mm long, white, adaxially 

Tabasco: Reforma, Balancafn, 22 May-6 Jun 1939 (fl), papillate, abaxially densely appressed-pubescent on 

Matuda 3193 (F, K). Yucatin: Becanchen, vic. of Aguada lower half, the apex obtuse, the base clawed; append- 

Xcruz-Akal, 3 Feb 1956 (fr), Enriquez 392 (US). ages deltate, as long as the petals, adaxially densely 

GUATEMALA. Peten: Tikal National Park, on ru- sericeous on upper half, abaxially papillate, adnate along 

ins, 11 Nov 1959 (fr), Contreras 375 (MO); Between margins of lower half of the petal to form a pocket, the 

San Jose & El Remate, along Laguna Peten, 5 Aug 1967 

apex obtuse; disc annular, 5-lobed, fleshy, tomentulose, 

(fl), Contreras 6991 (MO); San Andres, 4 May 1933 

1-1.2 mm tall; stamens 8, the filaments of unequal 

(fl), Lundell 3121 (US); Kantetul Bajo, 2 May 1933 (fl), 

length, 2.7-4.5 mm long, glabrous, the anthers ca. 1 

Lundell 3185 (K). 

mm long, ellipsoid, glabrous, obtuse at apex; ovary 

conical, ferruginous-tomentose, the stigma cylindric- 

8. TALISIA FURFURACEA Sandwith, Kew Bull. trigonous, papillate. Fruits ovoid or ellipsoid, woody, 

1935. 121. 1935. Type. Guyana. Mazaruni: Mazaruni densely ferruginous-sericeous, yellowish brown, 2.5- 

Station, 1 Mar 1934 (fl), Davis 359 (Forest Dept. 2.8 cm long, the pericarp ca. 3 mm thick Seeds one per 

ofBritish Guiana 2352) (holotype, K, n.v.; isotypes, fruit, ellipsoid, ca. 2 cm long, with a thin fleshy cover. 

NY, K-3). Figs. 3c-f, 5a, 47a-g Embryo with cotyledons superimposed, the upper cotyledon 

slightly larger than the lower one. 

The specific epithet seems to refer to the indumentum 

found on this species, however, it is better described 

as sericeous or appressed-pubescent, rather than 

furfuraceous. 

Tree to 30 m tall, branching on upper portion to 

produce a dense crown; trunk to 45 cm in diam., bark 

grayish brown, rough. Young growth conspicuously 

ferruginous-tomentose intermixed with short papilliform 

hairs. Twigs nearly terete, sulcate, rough, glabrescent 

with age. Leaves paripinnate; distal process cylindrical, 

truncate, ca. 4 mm long, early deciduous; leaflets 

4-8, opposite, elliptic to oblanceolate, 6.5-21 x 2.6-8 

cm, rigid-coriaceous, the adaxial surface slightly lustrous, 

glabrous, depressed along both sides of the 

prominulous midvein, sometime with scattered, raised, 

conical insect galls, the abaxial surface dull, sparingly 

appressed-pubescent, especially along the strongly 

prominent mid and secondary veins, the venation 

brochidodromus, forming a marginal loop, tertiary venation 

reticulate, the margins entire, revolute to strongly 

revolute, the apex obtuse, or shortly acuminate, the base 

obtuse, cuneate or rounded sometimes slightly asymmetrical; 

petiolules cylindrical, slightly enlarged at base, 

rugose, grayish, 4-10 mm long, furrowed along adaxial 

surface, puberulous; rachis 1.5-14 cm long, nearly terete, 

or obtusely angled on young leaves, bisulcate along 

adaxial surface, ferruginous-papillate intermixed with 

soft, long hairs, becoming glabrous with age; petioles 

3.5-9 cm long, slightly flattened along adaxial surface, 

slightly enlarged at base, with same indumentum as the 

rachis. Thyrses panicle-shaped, axillary or congested 

on distal portion of branches, to 21 cm long, the lateral 

Phenology. Known to flower from September to 

March and to fruit from October to May. 


the Guianas, in mixed forests on sandy soil or loam. 


Cuyuni/Mazaruni: Mazaruni Station, 30 m, 14 May 

1936 (fr), Davis 501 (K-2). Upper Demerara: Moraballi 

Creek, Essequibo river, 8 Sep 1939 (fl), Fanshawe 251 

(K-3, NY); Mabura Hill, 10 Dec 1994 (fr), Jansen- 

Jacobs 1134 (US). West Demerara: Makauria Creek, 

Essequibo river, 15 Oct 1940 (yfr), Fanshawe 576 (K, 

NY-2). 

SURINAM. Brokopondo: Brownsweg, nature park, 

Feb 1977 (fl), Roberts & v. Troon 14806 (MO, NY). 

FRENCH GUIANA. Bassin du Sinnamary: 

Sinnamary river, 8 Sep 1988 (st), Sabatier & Prevost 

2202 (CAY, US). Piste de Saint Elie: Station Biologique 

ORSTOM, 29 Apr 1992 (st), Acevedo R. & Grimes 4863 

(CAY, NY, US), 1 May 1992 (st), Acevedo R. & Grimes 

4874 (CAY). 

9. TALISIA GRANULOSA Acevedo-Rodriguez, sp. 

nov.


G fro Janen Jcobs1134 

G from Jansen-Jacobs 1134).~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~ 

Fig. 47. Talisia furfuracea. A. flowering branch. B. staminate flower. C. abaxial, adaxial and lateral views of 

petal with adnate appendage. D. staminate flower with perianth removed showing disc, stamens, and detail of 

anther. E. portion of infructescence. F. l.s. fruit. G. embryo, dorsal view. (A-D from Roberts & Troon 14806; B- 

Type. Brazil. Amazonas: Mun. Manaus. Reserva 

Ducke, Manaus-Itacoatiara Hwy, km 26, plateau forest, 

2053'S, 59058'W, 15 Jul 1997 (fr), PA. Assunfao et al. 

538 (holotype, US; isotype, INPA, n.v.). Fig. 48a-b 

A Talisia floresii Standley foliolis abaxiale albo- 

granulosis differt. 

Tree; trunk furrowed; bark reddish brown, with 

numerous lenticels, peeling off in irregular plates; inner 

bark light orange. Young stems striate, minutely 

appressed-pubescent, becoming terete and lenticellate. 

Leaves paripinnate; distal process acicular, deciduous, 

leaving a 1 mm long base with truncate apex; leaflets 

4-6, opposite or subopposite, 5-9 x 1.7-4, oblanceolate 

or obovate, coriaceous, the adaxial surface glabrous, 

slightly lustrous, with sunken primary and secondary 

veins, the abaxial surface albo-granulose, scattered, 

appressed-pubescent, with prominent primary and 

secondary veins, the venation brochidodromus, 

orangish, with secondary veins alternate or opposite, 

straight or slightly arching toward the margin, tertiary 

veins reticulate, the margins strongly revolute, the apex 

obtusely apiculate, the base obtuse; petiolules 1-5 mm 

long, minutely appressed-pubescent, adaxially furrowed; 

rachis 1.4-3.5 cm long, striate, minutely appressed-pubescent; 

petioles 2.5-4.2 cm long, minutely


Fig. 48. Talisia granulosa. A. sterile branch and detail of abaxial leaflet surface. B. fruit and lateral view of 

embryo. (A-B from Assun9ao 538). 

appressed-pubescent, lenticellate, slightly flattened and 

bicanaliculate along adaxial surface, slightly enlarged 

at base. Inflorescence and flowers unknown. Fruits 

depressed-globose, brownish, smooth, minutely ferruginous-sericeous, 

2 x 3.1 cm long, the pericarp 

woody, ca. 2 mm thick. Seed ca. ellipsoid, ca. 2.7 cm 

long, with fleshy testa. Embryo peanut-shaped; cotyledons 


Talisia granulosa is similar to T floresii, however, 

T granulosa can be distinguished from the latter by 

the leaflets with albo-granulose and scattered, appressed-pubescent 

abaxial surface (vs. glabrous). The 

specific epithet refers to the granulate appearance of the 

abaxial surface of the leaflets. 

Phenology. Collected in fruit during July. 


from the type collection. 

3 cm. 

10. TALISIA JAPURENSIS (C. de Candolle) 

Radlkofer in Martius, Fl. Bras. 13(3): 560. 1900. 

Guareajapurensis C. de Candolle in Martius, Fl. 

Bras. 11(1): 199. 1878. Type. Brazil. Amazonas: 

Japura river, Martius s.n. (holotype, G?,n.v.; 

isotype, M). Fig. 49a-f 

Tree 3-14 m tall. Stems terete, light brown, striate, 

ferruginous-tomentose, glabrescent and lenticellate with 

age. Leaves imparipinnate or paripinnate; distal process 

early deciduous, leaving a truncate base 2-3 mm long; 

leaflets (2)3-8, alternate, subopposite or opposite, elliptic, 

(4)6-23.5 x (1.8) 3-9.5 cm, chartaceous, slightly 

discolorous, the adaxial surface glabrous with 

prominulent midvein and slightly impressed secondary 

veins, the abaxial surface puberulent especially along 

veins, glabrescent, venation brochidodromus, prominent 

on abaxial surface, especially the midvein and


Fig. 49. Talisia japurensis. A. denuded fruiting branch. B. inflorescence. C. staminate flower. D. lateral, adaxial, 

and abaxial views of petal with adnate appendage. E. staminate flower with perianth removed showing disc and 

stamens. F. fruiting branch. (A from Martius s.n.; B-E from Ducke 18906; F from Martius s.n.).


9owl 7rWI s H owI 50WI 5RV 40W 

lp . o's. , S ,


of cachoeira Alta, 20 Sep 1954 (fl), Chagas s.n., herb. ellipsoid, glabrous, retuse at apex; ovary ferruginous- 

127 (IAN, INPA); Mindu creek, 27 Aug 1956 (fl), sericeous, the stigma capitate. Fruits not known. 

Coelho s.n., herb. 4130 (INPA-2); Mun. Presidente Talisia lanata can be distinguished from all other 

Figueredo along road to hydroelectric project, close to 

species of Talisia by the woolly pubescence of inflothe 

airport, 16 Jul 1986 (fr), C. Ferreira 7597 (US); Mun. 

rescences and abaxial surface of leaflets. No other spe- 

Borba. vic. of Urucurituba, 4-6 Sept 1934 (fl), Krukoff 

5960 (F, MO, NY, US); Manaus, 12 Sep 1955 (fl), Mello 

cies of Talisia is known to have this kind of pubess.n., 

herb. 1883 (INPA); Manaus, road to Aleixo, at INPA cence on its vegetative parts. The specific epithet refers 

grounds, 9 Oct 1976 (yfr), Monteiro 1285 (INPA); Rio to the indumentum. 

Negro, vic. of Cuieiras river, 12 Oct 1972 (fl), Pires & 

Piata 266 (INPA, US); Manaus, 3 Oct 1961 (fr), 

Phenology. Collected in flower during January. 

Rodrigues & Chagas 2557 (INPA); Manaus-Porto Velho Distribution and Ecology. (Fig. 54) Known only 

road, between Castanho and Tupana rivers, 17 Jul 1972 

(bd), Silva et al. 827 (INPA). Pari: Tapaj6s river, 14 

Aug 1923 (fl), Ducke s.n., herb. 18906 (RB, U). 


12. TALISIA MICROPHYLLA Uittien, Recueil 

Trav. Bot. Neerl. 34: 483. 1937. Type. Surinam. 

11. TALISIA LANATAAcevedo-Rodriguez, sp. nov. Brownsberg, 22 Mar 1924 (fl), Boschwezen 6445 

Type. Colombia. Amazonas: Araracuara, Inchi, disturbed 

vegetation, 17 Jan 1985 (fl), NC. Garzon, P 

(holotype, U, photo at US; isotypes, K, IAN, MO, 

NY, U). Fig. 52a-g 

Palacios & J. Mirana 145 (holotype, COAH). Tree 20-25 m tall, with small crown; trunk reaching 

Figs. 4d, 51a-d 35 cm in diam.; buttressed at base; bark light grey or 

light brown, rough lenticellate. Stems terete, glabrous or 

A T. furfuracea Sandwith foliolis et inflorescentiis 

lanatis differt. 

puberulent, striate, dark brown, sometimes minutely 

lenticellate. Leaves paripinnate or less often imparipinnate; 

Tree 8 m tall. Stems nearly terete, sulcate, ferrugi- distal process acicular, 4-5 mm long, carinate along 

nous-tomentose. Leaves imparipinnate; distal process adaxial surface, early deciduous leaving a truncate base; 

early deciduous; leaflets 7, alternate, with lowermost pair leaflets (2) 4 (5) opposite or alternate, obovate, elliptic, 

opposite, elliptic or nearly so, 10.7-15.5 x 4.2-5.1 cm, oblong or lanceolate, 4.5-12 x 1.5-4.5 cm, chartaceousrigidly 

coriaceous, bullate, adaxially glabrous and lus- coriaceous, usually with scattered necrotic round spots, 

trous, with sunken primary and secondary veins, glabrous on both surfaces, discolorous (abaxial surface 

abaxially densely ferruginous-woolly over the promi- darker), the venation brochidodromus, inconspicuous, 

nent primary and secondary veins, scattered so on the plane on both surfaces, midvein prominent on both surreticulate 

tertiary veins, the venation brochidodromus, faces, tertiary veins reticulate, the margins entire or 

forming a strong continuous intramarginal vein, the slightly undulate, slightly revolute, the apex obtusely 

margins entire, strongly revolute, the apex obtuse, with acuminate to caudate, glandular-mucronulate, the base 

a short apiculum, the base obtuse, slightly unequal; peti- usually asymmetrical, obtuse, rounded or obtuse-cuneate; 

olules cylindrical, 7-10 mm long, densely ferruginous- petiolules slightly thickened, 2-5 mm long, glabrous or 

tomentose; rachis ca. 10.5 cm long, terete, ferruginous- puberulent, adaxially canaliculate; rachis 2-4(-6) cm 

tomentose, striate; petioles ca. 11.5 cm long, terete, long, obtusely angled, adaxially carimnate, puberulent; petferruginous-tomentose, 

striate, slightly enlarged at the ioles 1.2-3 (4) cm long, slightly flattened, striate, swolvery 

base. Thyrses fasciculate, axillary, ca. 9 cm long; len at the very base. Thyrses panicle-shaped, terminal 

cataphylls apparently lacking; axes nearly terete, ferrugi- on lateral branches, 8-12 cm long, branches 2-4 cm long; 

nous-woolly; bracts oblong, puberulent, ciliate at mar- cataphylls deltoid, 1.5-2 mm long, puberulent; axes 

gins, 2.5-3 mm long; dichasia compound, sessile; angled, sulcate, minutely ferruginous pubescent; bracts 

pedicels ca. 1 mm long, articulate at the middle, woolly. and bracteoles subulate-lanceolate, 1-1.5 mm long, per- 

Calyx ca. 4 mm long, tomentose, the sepals free to the sistent, minutely ferruginous pubescent; dichasia combase, 

ovate, obtuse at apex; petals (immature) elliptic, ca. 

3 mm long, adaxially papillate, abaxially glabrous, except 

pound or the distal ones simple; peduncle 2-3 mm long, 

minutely ferruginous pubescent or tomentulose; pedicels 

for the pilose base, the apex obtuse, the base cuneate- ca. 1.2 mm long, articulate at the base. Calyx 2-2.6 mm 

attenuate; appendages nearly as long as the petals, broadly 

deltate, adaxially sericeous, abaxially glabrous at base, 

sericeous toward the apex; disc annular, tomentose, ca. 

0.5 mm tall; stamens (immature) 8 or 10, the filaments 

of equal length, glabrous, the anthers ca. 0.8 mm long, 

long, ferruginous-tomentulose or pubescent, the sepals 

free to the base, concave, ovate, obtuse or acute at apex, 

adaxially tomentulose; petals rhombic-ovate, 3-4 mm 

long, glabrous or sparsely pubescent on both surfaces, 

the apex acute, the base cuneate; appendages as long as


~~~~~~~~y ~ ~ ~ ~ .. 

2mm fi- f 

Fig. 51. Talisia lanata. A. flowering branch and detail of abaxial and adaxial surfaces of leaflets. B. flower bud. 

C. abaxial and adaxial views of petal with adnate appendage. D. pistillate flower with perianth removed showing 

disc, stamens and pistil. (A-D from Garzon et al. 145).


2 mm. 

L Im. 

Eimm} ~~5mm 

iW~~~~~~~~~~~ n 

Fig. 52. Talisia microphylla. A. flowering branch. B. staminate flower. C. adaxial and abaxial views of petal with 

adnate appendage. D. staminate flower with perianth removed showing disc, stamens and pistillode. E, stamen. 

F. pistillode (left) and i.s. pistillode (right). G. leaf. (A-F from Forest Dept. British Guiana 5355; G from Wood 

Herbarium Surinam 321). 

the petals or only 2/3 the length of petals, rhombic-ovate, capitate, 3-lobed, tomentulose. Fruits ovoid, ferruginousabaxially 

papillate, adaxially woolly-pubescent, fused to tomentose, minutely rugose, 2-2.4 cm long, shortly apithe 

petal along its basal margins to form a pocket, trun- culate at apex, the pericarp 1 mm thick, coriaceous, the 

cate at apex; disc annular, 10-lobed, glabrous or spar- endocarp sparingly pubescent or glabrous. Seeds ellipingly 

puberulent, ca.0.8 mm tall; stamens 8, the filaments soid, with pale pink or whitish edible, fleshy testa. Emsubulate, 

of equal or unequal lengths, 0.8-2 mm long, bryo not seen. 

glabrous, the anthers 0.5-0.6 mm long, ovoid, glabrous, The specific epithet refers to the small size of the 

retuse at apex; ovary nearly conical, tomentose, the stigma leaflets in this species. 

.....


Phenology. Collected in flower from March to May to 17 cm long; cataphylls lacking; axes angled, sulcate, 

and in September, and in fruit in June. 

puberulent; bracts early deciduous; dichasia compound, 

Distribution and Ecology. (Fig. 54) The Guianas, 

Ecuador, and Brazil, in primary, lowland, moist, nonflooded 

forest. 

sessile; pedicels ca. 1 mm long, articulate at base. Calyx 

1.5-2 mm long, tomentose, the sepals free to the base, 

concave, lanceolate, obtuse at apex; petals ca. 3 mm long, 

lanceolate, clawed at lower half, pilose abaxially at claw, 

Common name. Surinam: zwarte pintolobus. the blade abaxially glabrous, adaxially papillate, the apex 

obtuse; appendages as long as the petals, elliptic, erect, 

Representative specimens examined. GUYANA. densely sericeous, adaxially, sparsely sericeous- papil- 

Upper Demerara/Berbice: Moraballi creek, Essequibo 

late on abaxial surface; disc 5-lobed, minutely woollyriver, 

45 Apr 1946 (fl), Forest Dept. of British Guiana 

pubescent; stamens 8, the filaments of unequal lengths, 

5355 (K-2, NY, US); Bartica, on the Essequibo river, 

0.7-1.8 mm long, woolly-pubescent, the anthers ca. 0.6 

17 Jun 1952 (fr), Forest Dept. of British Guiana 6966 

(K, NY). Essequibo Islands/West Demerara: Groete mm long, elliptic, glabrous, apiculate at apex. Pistillate 

creek, lower Essequibo river, 20 Jun 1944 (fr), Fanshawe flower unknown. Fruits ellipsoid, ferruginous-sericeous- 

1982 (NY). 

tomentose, 2.2 cm long (immature), green, shortly api- 

SURINAM. Sakali(?), May 1945 (st), Wood her- culate at apex, the pericarp 1 mm thick, leathery, the enbarium 

Surinam 321 (NY). Nickerie: Fallawatra, 8 Nov docarp tomentulose. Seeds ellipsoid, ca. 1.3 cm long, 

1971 (st), Jimenes-Saa 1571 (CAY). 

with fleshy testa. Embryo with cotyledons superimposed, 

FRENCH GUIANA. Piste de Saint Elie: Station of equal size. 

Biologique ORSTOM, 4 May 1992 (st), Acevedo R. 4890 

Talisia parviflora seems to be closely related to T 

(CAY), 1 Nov 1991 (st), Prevost & Sabatier 2976 

coriacea because of their morphological similarity. 

(CAY). 

ECUADOR. Sucumbios: Reserva Cuyabeno, 1 km 

However, T parviflora differs by: calyx 1.5-2 mm long 

N of Laguna Grande, 265 m, 1 Apr 1988 (st), Valencia with lanceolate sepals (vs. calyx 2.3-3.2 mm long, with 

et al. 67728 (QCA). 

ovate sepals); petals ca. 3 mm long, (vs. 3.2-5.5 mm 

BRAZIL. Amazonas: Manaus. Distr. Agropecuario long); filaments woolly-pubescent (vs. glabrous). The 

da Suframa, Faz. Esteio, Nov 1979 (st), Rankin et al. 84 specific epithet refers to the relatively small flowers of 

(INPA). Para: Tocantins river, 10 Apr 1981(st), Maciel 

et al. 576 (MG); Tucurui, 22 Sep 1983 (fl), Ramos 910 

(INPA, NY), 25 Nov 1983 (st), Ramos 1169 (INPA, NY). 

this species. 

Phenology. Collected in flower during May and 

August, and in fruit during October. 

13. TALISIA PARVIFLORA Acevedo-Rodriguez, 

sp. nov. 

Type. Brazil. Amazonas: Manaus-Caracarai road, 

km 121, sandy soils, 8 Aug 1974 (fl), A. Loureiro et 

al. s.n. (holotype, MO). Fig. 53a-e 

A T. coriacea Radlkofer floribus minoribus et 

filamentis pubescentibus differt. 

Tree 10-20 m tall; trunk to 30 cm in diam. Stems 

terete, puberulent, striate, becoming glabrous and 

lenticellate with age. Leaves paripinnate; distal process 

truncate, minute; leaflets 4, opposite, lanceolate, oblong 

or ovate, 4-10 x 1.9-4.5 cm, coriaceous, glabrous on 

both surfaces, the venation brochidodromus, the midvein 

plane or sunken on adaxial surface, prominulous on 

abaxial surface, secondary and tertiary veins inconspicuous, 

the margins slightly undulate, the apex obtuse to 

long-acuminate, the base rounded or obtuse, slightly 

asymmetrical; petiolules canaliculate, 3-5 mm long, puberulent 

or glabrous; rachis 0.8-3 cm long, slightly flattened 

adaxially, striate, puberulent; petioles 2.5-5 cm 

long, slightly flattened adaxially, striate, pulvinate at base. 

Thyrses panicle-shaped, terminal, to 18 cm long, branches 

Distribution and Ecology. (Fig. 54) In moist, terra 

firme forest on sandy or clayish-sandy soils in low- 

lands of Brazil. 


Amazonas: Manaus-Itacoatiara road, km 104, trail to 

INPA, 23 May 1967 (fl), Byron & Elias 67-51 (INPA); 

Carabinani river, 22 Oct 1976 (fr), Ramos s.n. (INPA). 

14. TALISIA PRAEALTA Sagot ex Radlkofer, 

Sitzungsber. Math.-Phys. Cl. K6nigl. Bayer Akad. 

Wiss. Muinchen 8: 345. 1878. Cupania praealta 

(Radlkofer) Sagot, Ann. Sci. Nat. Bot. Ser. 6, 12: 

199. 1881. Type. French Guiana. Acarouani, Mar 

1857 (fl), Sagot 1047(holotype, B-destroyed, photo 

(F neg. 5679) at US; lectotype, P, here designated, 

photo at US; isotypes, K-2, NY, P-3, U, n.v.; frag. 

ex B at M). Fig. 55a-f 

Tree to 30 m tall; trunk reaching 25 cm in diam., 

buttressed at base; bark light brown to grayish, rough, 

with numerous horizontal rows of lenticels, peeling off 

in large irregular plates; inner bark reddish brown. 

Branches terete, striate, puberulent. Leaves paripinnate;


2mm 

C ~~~~mm. 

Fig. 53. Talisia parviflora. A. flowering branch. B. staminate flower. C. abaxial and adaxial views of petal with 

adnate appendage. D. staminate flower with perianth removed showing disc and stamens. E. stamens. (All from 

Loureiro et al. s.n.). 

distal process acicular, 2-4 mm long; leaflets 2-4, opposite 

or subopposite, elliptic or obovate, chartaceous to 

subcoriaceous, 5-12.5 (-20.5) x 1.9-5.5 (8.2) cm, glabrous, 

often with scattered, necrotic round spots, midvein 

impressed adaxially, prominent abaxially, secondary 

veins prominulous abaxially, tertiary veins reticulate, the 

apex obtuse or obtusely acuminate and retuse, the base 

acute or obtuse, decurrent on to the petiolule, sometimes 

slightly asymmetrical; petiolules slender, 2-6 (15) mm 

long; rachis 0.7-3.5 (5.5) cm long, slightly flattened along


9owl I' . 

70WI 60w 50w1 40W 

* Talisia lanata 

V Talisia microphylla[ 

* Talisia parviflora 

K iON 

*4/,J Y.'. . ;--' . 7 t, 5'1O 

A 

Fig. 54. Distribution of species in Talisia subgenus Pitombaria (in part). 

~~3mm. 

~~~f 

A. F.4t ~~~~~~~E. 

Fig. 55. Talisia praealta. A. fruiting branch. B. staminate flower. C. adaxial and abaxial views of petal with adnate 

appendage. D. staminate flower with perianth removed showing disc and stamens (left) and I.s. same showing pistillode 

(right). E. stamens. F. pistillode (left) and I.s. pistillode (right). (A from Mori et al. 20822; B-F from Wachenheim 308). 

2Q


adaxial surface, striate, puberulent; petioles 0.7-2.5 (7.5) (st), Villiers & Feuillet 2020 (CAY). Bassin du Maroni: Haut 

cm long, similar to rachis, slightly enlarged at base. Th- Maroni, Aug 1979 (fr), Moretti 971 (CAY-3); Saut du 

yrses racemiform, 2-8 cm long, axillary, congested or Litani, vic. Antecume Pata, 4 Aug 1989 (fr), Moretti 1189 

less often solitary; cataphylls wanting; axes puberulent; (CAY); Grand Inini river, 13 Jul 1990 (st), Sabatier & 

bracts and bracteoles subulate, 0.5-1 mm long, early Prevost 3254 (CAY); Godebert, 14 Jun 1921 (fl), 

deciduous; dichasia simple, sessile, or of a solitary flower Wachenheim 308 (P). Bassin de L'Oyapock: Armontabo 

due to the abortion of lateral flowers; peduncle 1-2.2 creek, between Saut Bace & Saut Canori, 4 Jul 1969 (yfr), 

mm long, tomentose; pedicels 1-2 mm long, tomentose, Oldeman T-285 (CAY, P, US); opposite Colonia Agricola 

articulate at the middle. Calyx 2.5-3 mm long, ferrugi- do Oiapoque, about 4 km N of mouth of Cricu river, 12 

nous-tomentose, the sepals free to the base or nearly so, Aug 1960 (fr), M. Pires 47465 (F, NY). Bassin du 

oblong or elliptic; petals elliptic, ca. 2.2 mm long, abaxially Sinnamary: Tigre creek, 7 m, 14 Jun 1994 (fr), Bordenave 

tomentose on lower half, adaxially papillate; appendages 1013 (CAY). Piste de Saint Elie: Station Biologique 

deltate, as long as the petals, sericeous-tomentose on both ORSTOM, 1 May 1992 (st), Acevedo R. et al. 4873 (CAY, 

surfaces; disc annular, 5-lobed, minutely hispidulous; US), 3 May 1992 (st), Acevedo R. 4882 (CAY, US); Interstamens 

(7)8, the filaments of unequal length, 2-3 mm section of Sinnamary-Counamari rivers, 12 Jan 1991 (st), 

long, sparsely to densely pilose, the anthers oblong-el- Pr&vost & Sabatier 2978 (CAY), 4 Oct 1991 (st), Sabatier 

lipsoid, apiculate and barbate at apex, ca. 0.7 mm long; & Prevost 3894 (CAY). Region Littorale: Kourou, 

ovary ferruginous, sericeous-tomentose, the stigma elon- Passoura creek, 6 May 1992 (st), Acevedo R. et al. 4905 

gated, papillate. Fruits ellipsoid, light green, to 2.2 cm (CAY, US). Region de Saul: Montagne La Fumee, 300 m, 

long, minutely ferruginous, sericeous, the pericarp nearly 5 Sep 1989 (fr), Mori et al. 20822 (CAY, US-2). 

woody, ca. 1 mm thick, smooth to slightly granulate, the BRAZIL. Amazonas: Mun. Itapiranga, Pitinga river, 

endocarp glabrous. Seed one per fruit, nearly ellipsoid, 

ca. 1.7 cm long, with slightly fleshy testa. Embryo with 

24 Aug 1979 (fr), C. Ferreira et al. 689 (INPA, NY, US). 

cotyledons lying dorsiventrally over each other, the upper 

one slightly larger than the basal one. 

15. TALISIA SIMABOIDES Kramer, Act. Bot. 

Talisia praealta vs. T squarrosa: T praealta has Neerl. 21: 676. 1972. Type. Surinam. North Westcongested 

short inflorescences and (2) 4-6-foliolate ern Surinam, Snake creek a tributary of Marataka 

leaves, while T squarrosa has long, paniculate thyrses river, 17 May 1965 (fl), RJ.M. Maas 10818 (holoand 

bifoliolate leaves. Talisiapraealta vs. T coriacea, type, U, n.v.; isotypes, K, NY). Fig. 56a-h 

although very similar at times seems to overlap, the 

shape of leaflets seems to be different; filaments are 

glabrous in T coriacea, pilose in T praealta. 

The specific epithet meaning very high, refers to the 

relatively large size of this species. 

Tree to 30 m tall; trunk reaching 35 cm in diam., 

buttressed at base to 1 m high; bark grayish brown, 

smooth, with lenticels in rows; inner bark orange or 

reddish brown. Branches sulcate, tomentulose, becoming 

terete and lenticellate with age. Leaves paripinnate 

Phenology. Collected in flower in February, June or imparipinnate; terminal process truncate, ca. 2 mm 

and July; fruiting from July to September. 

long; leaflets 5-8, alternate or subopposite, oblong, 


Venezuela, Surinam, French Guiana and the State of 

Amapa, Brazil, in moist, non-flooded forest and gallery 

forest often in primary or disturbed habitats. 

elliptic, lanceolate, or nearly obovate, chartaceous to 

subcoriaceous, 4-9(14) x 1.5-3.4(4.5) cm, discolorous, 

the venation brochidodromus, tertiary veins finely reticulate, 

the adaxial surface glabrous, often with scattered, 

raised, conical insect galls, or necrotic, round 

Common names: Venezuela: Mamon de cerro; spots, with impressed midvein and prominulous sec- 

French Guiana: singabassou, tapuyma; Brazil: pitombarana. 

Representative specimens examined. VENE- 

ZUELA. Barinas: Pedraza, 18 Feb 1954 (st), Bernardi 

1147 (K, VEN). 

SURINAM. Brokopondo: Brownsweg, Feb 1978 (fl), 

Roberts 16313 (CAY, K, NY, US). Saramacca: Nassau, 

21 Feb 1949 (st), Lanjouw & Lindeman 2218 (NY). 

FRENCH GUIANA. Bassin de L'Approuague: Saut 

Sarari, 7 Sep 1984 (fr), Riera 783 (CAY-2); Station des 

Nouragues, 1 Nov 1987 (st), Sabatier & Prevost 1932 

(CAY, US); Arataye river, 70 km SW of Regina, 6 Oct 1983 

ondaries, the abaxial surface usually drying brownish, 

minutely puberulent, especially along the prominent 

midvein, secondary veins inconspicuous or less often 

prominulous, the apex obtuse, retuse, sometimes nearly 

acuminate, the base obtuse or acute, decurrent onto the 

elongated petiolule; petiolules nearly cylindrical, slender, 

glabrous, 3-7 mm long, striate, adaxially 

canaliculate; rachis 3.5-7(11) cm long, puberulent, terete, 

striate; petioles 2.5-5.5 cm long, puberulent, slightly 

flattened adaxially, striate, minutely lenticellate, slightly 

enlarged at base. Thyrses panicle-shaped, 9-25 cm long, 

terminal on branches; cataphylls wanting, the axes ob-


Fig. 56. Talisia simaboides. A. sterile branch. B. flowering branch. C. pistillate flower. D. pistillate flower with 

perianth removed showing pistil and sterile stamens (left) and l.s. same (right). E. adaxial, abaxial, and lateral 

views of petal with adnate appendage. F. lateral and ventral views of stamen. G. portion of infructescence. H. 

embryo, dorsal view. (A-F from Sabatier & Prevost 3958; G-H from Sabatier & Prevost 3820). 

tusely angled, striate, minutely, velvety pubescent; above the base, abaxially glabrous; disc annular, crenate, 

bracts and bracteoles triangular, tomentose, ca. 0.5 mm tomentose, 0.9 mm tall; stamens 8, the filaments of 

long, early deciduous; dichasia simple, nearly sessile; unequal lengths, glabrous or puberulent, the anthers 

pedicels 1.5-2 mm long, ferruginous-tomentulose, ar- ovoid, 0.5-0.75 mm long, apiculate; ovary ovoid, toticulate 

at the middle or below. Calyx pale green, 2.5- mentose, the stigma trigonous-capitate, papillate. Fruit 

3 mm long, minutely tomentose or pubescent, the se- ellipsoid to ovoid, ca. 2 cm long, densely appressed 

pals free to the base, ovate or deltate, concave; petals ferruginous pubescent, the pericarp woody, ca. 1 mm 

white, rhombo-ovate or ovate, ca. 3 mm long, abaxially thick, the endocarp sparsely woolly-pubescent. Seed 

appressed-pubescent at base, adaxially glabrous, cu- solitary, with thin, dry testa. Embryo with cotyledons 

neate at base, obtuse at apex; appendages as long as the of similar size, lying transverse over each other. 

petals, ovate-deltate, fused at base to the basal margins The specific epithet seems to refer to the Simabaof 

the petal to form a pocket, adaxially white-sericeous like (Simaroubaceae) appearance of the leaves.


Phenology. Flowering from May to August, and 

fruiting in September. 


French Guiana, in lowland, moist, primary, non- 

flooded forest. 

Representative specimens examined. FRENCH 

GUIANA. Bassin de L'Approuague: Station des 

Nouragues, 26 Oct 1987 (st), Sabatier & Prevost 1852 

(CAY), 26 Jul 1989 (st), Sabatier & Prevost 2885 (CAY). 

Bassin du Maroni: Godebert, 14 Jun 1921 (fl), 

Wachenheim 371 (P-2). Bassin du Sinnamary: Sinnamary 

river, along Courcibo river, vic of Saut Caouene, 500 m, 

10 Aug 1967 (fl), Oldeman B-1194 (CAY, P, US). Piste 

de Saint Elie: Station Biologique ORSTOM, 28 Apr 1992 

(st), Acevedo R. et al. 4857 (CAY, US); Intersection of 

Sinnamary-Counamamari rivers, 27 Jul 1991 (fl), 

Sabatier & Prevost 3658 (CAY, NY, U, US), 27 Sep 1991 

(fr), Sabatier & Prevost 3820 (CAY, U, US); Station 

Biologique ORSTOM, 6 Aug 1991 (st), Sabatier & Prevost 

3717 (CAY). Region du Saul: Montagne La Fumee, 200 

m, 15 Sep 1982 (st), Boom & Mori 1688 (CAY, NY); 25 

Oct 1982 (st), Boom & Mori 2379 (NY). 

16. TALISIA SQUARROSA Radlkofer, Sitzungsber. 

Math.-Phys. Cl. K6nigl. Bayer Akad. Wiss. 

Miinchen 8: 346. 1878. Type. Guyana. Roraima, 

1842-43 (fl), Rob. Schomburgk 738 (holotype, BM 

?; isotypes, P, NY; frag. at M). Fig. 57a-e 

0.6-0.7 mm tall; stamens 8, the filaments glabrous, of 

unequal length, 1-2 mm long, the anthers ovate, obtuse at 

apex, 0.7-0.8 mm long; ovary conical, sericeous-tomentose, 

the stigma elongated, trigonous. Fruits (immature) 

ellipsoid or ovoid, sericeous-tomentose, the pericarp 

woody, smooth. Seed ellipsoid, with white juicy sarcotesta 

(fide Polak 1992). 

According to Steam (1983), squarrosa means rough 

with scales, a term that does not seem to describe any 

morphological features in this species. The stems of 

this species are rough however, due to the presence of 

numerous lenticels. 

Phenology. Flowers from March to October and 

produces fruit from May to October. 


Guyana and French Guiana, in non-flooded, moist forest 

on sandy or clayish soils. According to Polak (1992), 

this species is frequent in the Wallaba and the Clump- 

Wallaba forest on white sands. 

Common names and uses. French Guiana: Grigri, 

singabassou; Guyana: candlewood, karimora, (fide 

Brunner et al. 1994), muro-balli, moroballi, sand mora. 

The leaves, fruits, wood, and heartwood are used for 

fish poisoning in Guyana. 


Without specific locality, Demerara river, 1 Mar 1943 

(st), Fanshawe 1215 (K);Without specific locality or date 

Tree to 20-30 m tall; trunk reaching 30 cm in diam., 

(fl), Schomburgk 736 (K), (fl), Schomburgk 1351 (M, 

buttressed at base; bark silvery grey, inner bark reddish 

US). Barima/Waini: Amakura river, 23 Mar 1923 (fl), 

brown. Branches terete, rough, lenticellate, glabrous. de la Cruz 3475 (MO, NY). Potaro: Potaro river, be- 

Leaves paripinnate; distal process early deciduous; leaf- low Tukeit, 16 May 1944 (fl), Maguire & Fanshawe 

lets 2, opposite, elliptic, coriaceous, 5.3-9(19) x 2.4- 23510 (K, MG, MO, NY, P, US). Upper Demerara/ 

4.5(7.5) cm, glabrous, dull, the venationbrochidodromus, Berbice: Yawakuri forest, Berbice-Rupununi trail, Jul 

the adaxial surface with prominulous venation, the abaxial 1919 (fl), Abraham 219 (K, NY); Mora-mora-bisi creek, 

surface with prominent primary and secondary veins, ter- Courantyne river, 7 May 1917 (fr), Hohenkerk 715 (K, 

NY); Moraballi Creek, Essequibo river, 24 Sep 1929 

tiary veins reticulate, prominulous, the apex obtuse or 

(fr), Sandwith 333 (K-2, NY, P), 28 Sep 1929 (fl), 

acute, the base acute or obtuse, asymmetrical; petiolules 

Sandwith 357 (K, NY, P); Vic. of Mabura hill, 28 Oct 

pulvinate, glabrous, 3-7 mm long, drying dark brown; 1982 (fr), Stofers et al. 87 (NY, US). 

petioles 2-5.5 cm long, glabrous, flattened along adaxial FRENCH GUIANA. Bassin de L'Iracoubo: 

surface, lenticellate. Thyrses panicle-shaped, to 27 cm long, Rocoucoua, close to Iracoubo, 8 Sep 1986 (bd), Prevost 

terminal or axillary on distal portion ofbranches; cataphylls 2170 (NY). Bassin de la Mana: Organabo, ca. 2 km W 

wanting; axes obtusely angled, sulcate, puberulent; bracts of Organabo river, north of road, 9 Jul 2000 (st), Acevedo 

and bracteoles subulate, < 1 mm long, early deciduous; R. 11117 (CAY, US); Camp Charvein-Acarouany, km 4, 

dichasia compound; peduncles 2-4 mm long, puberulent 

29 Sep 1954 (yfr), Anonymous 226 m (CAY), (fl), Anonymous 

227m (CAY); Camp Charvein, 14 Oct 1913 (fl), 

orminutelytomentose; pedicels 2-3 mm long, tomentulose, 

Benoist 54 (P); Organabo Station, 12 Jun 1995 (st), Loubry 

articulate at the middle. Calyx 3-4 mm long, tomentose, 

2322 (CAY), 16 May 1993 (st), Loubry 2373 (CAY, US); 

the sepals free to base, elliptic to ovate, concave; petals Cayenne-St. Laurent road, km 171.5, 3 Sep 1986 (fl), 

white, rhombo-elliptic or wide elliptic, 2.8-3 mm long, Sastre et al. 8222 (CAY, P, U, US). 

adaxially minutely papillate, abaxially appressed-pubescent 

on lower half, the base cuneate to nearly clawed, the 

apex obtuse; appendages as long as the petals, deltate, as 17. TALISIA SUBALBENS (Martius) Radlkofer, 

wide as the petals ornanrower, abaxiallypapillate, adaxially Sitzungsber. Math.-Phys. Cl. Konigl. Bayer Akad. 

tomentose; disc cup-shaped, 5-lobed, minutely tomentose, Wiss. Miinchen 8: 345. 1878. Cupania subalbens


IA 0X 

A.~~~~~~~~~~~~~m 

I X 

t 

3 mm. 

Fig. 57. Talisia squarrosa. A. flowering branch. B. staminate flower. C. adaxial and abaxial views of petal with 

adnate appendage. D. staminate flower with perianth removed showing disc and stamens. E. stamen. (All from 

Sandwith 490).


9oWI 80 ?, 70WI 60W 5CWI 40W 

.~~~~~~~~~~~~~. ..' . ... 2 

21~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~ 

_ %ff;s tse vo-t 1 -> ;-s! 2Q~~~~~~~~ 

NI 

}xv XI_;Q gg- '.2 

7 Nj,~~~4 

S~~~~~~~~~~~~~~~~~~~~~~ 

tTalisia squarrosa ,', , .--' .. ' ' as u X' ' 

Fig. 58. Distribution of species in Talisia subgenus Pitombaria (in part). 

Martius, Flora 21(2): 72. 1838. Type. Brazil. Mato 

Grosso: Serra a Cima, close to Cuiaba', Oct (without 

year) (fl), Martius Herbarium 264 (holotype, 

M; isotype, K). Fig. 59a-g 

ly papillate and puberulent, abaxially puberulent at base 

and lower margins, the base attenuate or nearly clawed, 

the apex obtuse sometimes pubescent, slightly retuse; 

appendages as long as the petals, adnate to petal half of 

Shrub or treelet, 0.3-2 m; stems terete, cano- or fer- their length, elongate-deltate, erect, adaxially villose 

ruginous-tomentulose, striate. Leaves paripinnate; dis- along margins and upper half, abaxially puberulenttal 

process filiform, 3-4 mm long, truncate at apex; leaf- papillate; disc annular, 5-lobed, tomentose, 0.6-0.7 mm 

lets 8-12, opposite to alternate, asymmetrically tall; stamens 8, the filaments of unequal length, 2-3 mm 

oblong-elliptic, sub-lanceolate, 4-1O x 1.5-3.2 cm, co- long, tomentose, the anthers 1.2-1.6 mm long, ellipriaceous, 

the adaxial surface puberulent, dull, with soid or oblong-ovoid, glabrous, obtuse or apiculate at 

midvein slightly sunken, the abaxial surface albo-seri- apex; ovary conical, ferruginous-tomentose, the stigma 

ceous, with prominent midvein and prominulous sec- trigonous to capitate, tomentulose, papillate. Fruits 

ondary veins, the venation brochidodromus, tertiary ovoid, minutely ferruginous-tomentose, 2-2.5 cm long, 

veins reticulate, the margins entire, revolute, the apex shortly apiculate at apex, the pericarp woody, 3-4 mm 

obtuse or short- acuminate, with obtuse acumen, the thick. Seed solitary, the testa thin, apparently not fleshy. 

base unequal, obtuse, rounded or cordate; petiolules Embryo with cotyledons laying slightly obliquely dorslightly 

pulvinate, 1-2 mm long, minutely pubescent, siventrally over each other, the upper one slightly larger 

adaxially canaliculate; rachis 12-17 cm long, terete, than the lower one. 

bicanaliculate along adaxial surface, minutely pubes- The specific epithet refers to the whitish appearance 

cent; petioles 3-6 (1 1) cm long, slightly flattened along of the abaxial surface of leaflets. 

adaxial surface, striate, slightly enlarge at the very base. Phenology. Flowers from August to November and 

Thyrses simple or panicle-shaped, axillary or terminal, fruits in October and November. 

10-25 cm long; cataphylls acicular, minutely tomen- 

Distribution and 

tose; axes 

Ecology. (Fig. 62) Endemic to the 

angled, striate, minutely ferruginous-tomentose; 

bracts subulate, deciduous, ca. 2 mm long; dicha- 

Chapada dos Guimaraes and adjacent areas in the state of 

Mato 

sia compound; peduncle 2-10 mm; pedicels 2-4 mm 

Grosso, Brazil, in sandy cerrado and gallery forest. 

long, articulate at the middle. Calyx 3.5-4 mm long, Common name. Brazil: Cancudo. 

abaxially tomentose, adaxially appressed-pubescent, the 

sepals 2.5-3 mm long, elliptic, concave, obtuse at apex; 

Representative specimens examined. BRAZIL. Without 

specific locality or date (fl), Martius 146 (MO, P). 

petals elliptic to ovate, 3.5-4.5 mm long, erect, adaxial- Mato Grosso: Caxipozinho river, vic. of Cachoeira Veu


'7 

N~~~~~~~~~~~~~ 

~~~~~~~~V72~~~~~~~~~~~~~~~~~~~3m 

Fig. 59. Talisia subalbens. A. flowering branch. B. staminate flower. C. adaxial and abaxial views of petal with 

adnate appendage. D. staminate flower with perianth removed showing disc and stamens. E. lateral and ventral 

views of stamen. F. pistillode (left) and I.s. same (right). G. fruit (right) and c.s. fruit (left). (A-F from Hatschbach 

37543; G from Santos & Rosario 400). 

de Noiva, 21 Oct 1985 (fr), C. Ferreira et al. 6556 (MG, 

US); Riberao da Salgadeira, 14 Nov 1975 (fl), Hatschbach 

37543 (US); Rio Claro, 23 Oct 1993 (fl), Macedo et al. 

3331 (INPA); Chapada dos Guimaraes, Colegio Buriti 

field, 720 m, 11 Oct 1973 (fl), Prance et al. 18815 (K, 

US); Chapada dos Guimaraes, along road to Cachoeira 

Veu de Noiva, 21 Oct 1985 (fl), Pirani 1354 (US); Cuiaba, 

without specific locality, Aug 1927 (fl), Riedel 1106 (US); 

Santa Ana da Chapada, 20 Sep 1902 (fl), Robert 554b 

(K); Chapada dos Guimaraes, 21 Nov 1982 (fr), Santos 

& Rosario 400 (IAN, MG); Santo Ant6nio de Levenger, 

70 km W of Cuiaba; 5 km SW of Sao Vicente, 23 Oct 

1985 (fl), Thomas 

et al. 4553 (MO, US). 

18. TALISIAVELUTINAAcevedo-Rodriguez, sp. nov. 

Type. Peru. Loreto: Prov. Coronel Portillo. Dtto 

Iparia. Bosque Nacional de Iparia, tropical dry forest, 

along Ucayali river, 250-300 m, 20 Aug 1968 (fl), 

Schunke Vigo 2637 (holotype, F; isotypes, NY, US). 

Figs. 16e-f, 60a-e


2 mm. 

Fig. 60. Talisia velutina. A. leaf. B. inflorescence. C. staminate flower. D. lateral, abaxial, and adaxial views of 

petal with adnate appendage. E. staminate flower with perianth removed showing disc and stamens, and detail of 

stamen (left). (A from Schunke V 2637; B-E from Renteria 4841).


A T. clathrata Radlkofer foliis pilosis vel velutinis 30 m, 4 Apr 1985 (fr), Renteria 3741 (JAUM); Quebrada 

et caulibus velutinis differt. 

la Puerca and Malagon, 10 m, 24 Mar 1986 (fl), Renteria 

et al. 4758 (JAUM); Uraba-Chigorod6-Malagon, 

Shrub 3-7 m tall. Stems slightly sulcate, nearly ter- Cocuelo to Mono Macho, 10 m, 27 Mar 1986 (fl), 

ete, ferruginous-velutinous. Leaves imparipinnate or 

paripinnate; distal process acicular, to 1.5 cm long, early 

deciduous, leaving a truncate base; leaflets 8-11, alter- 

Renteria et al. 4841 (JAUM). 

nate, opposite or subopposite, 19-75 x 7-19 cm, elliptic 19. TALISIA VERALUCIANA G. Guarim Neto, 

to oblanceolate, chartaceous, the adaxial surface com- Acta Amazonica 9: 239. 1979. Type. Brazil. 

pletely glabrous or velutinous along the sunken primary Amazonas: Mun. HumaitA, terra firme forest, 1934 

and secondary veins, the abaxial surface pilose to (fr), B.A. Krukoff6886 (holotype, BR, n.v.; isotypes, 

sparsely velutinous, especially along veins, the vena- A, n.v., BM, n.v., F, K, MO, U, US-2). 

tion brochidodromus, with secondary veins alternate 

Fig. 61a-f 

or subopposite, tertiary veins clathrate-reticulate, prominent 

on abaxial surface, the margins wavy, the apex 

abruptly acuminate, mucronate, the base obtuse to 

Tree to 27 m tall; trunk buttressed at base. Stems angled 

to terete, smooth, striate, lenticellate, glabrous. Leaves 

rounded, slightly asymmetrical; petiolules 8-15 mm imparipinnate or paripinnate; distal process truncate, 2-3 

long, velutinous, nearly cylindrical; rachis 16-32 cm mm long; leaflets 6-12, alternate or opposite, oblong, 

long, velutinous, striate, nearly terete; petioles 27-56 ovate, elliptic or less often oblanceolate, 5.5-14.5 x 2-5 

cm long, velutinous, striate, terete, slightly enlarged at cm, coriaceous, adaxially glabrous, lustrous or opaque, 

base. Thyrses fasciculate, cauliflorous; cataphylls ac- with sunken midvein, abaxially densely appressed, fericular, 

to 1.2 cm long, velutinous, congested in supra- ruginous-sericeous, dull, with prominent midvein and 

axillary buds; axes 2-15 cm long, angled, velutinous- secondary veins, tertiary veins reticulate, inconspicuous, 

tomentose; bracts subulate, persistent, 2-4 mm long, the venation brochidodromus, with secondaries formsparsely 

tomentose to glabrous; bracteoles similar to ing a 60?-70? angle with midvein, tertiary veins reticubracts 

but smaller; dichasia compound, sessile; pedicels late, the margins entire, revolute, the apex abruptly acumica. 

1 mm long, articulate above or below the middle. nate, mucronate, the base obtuse, or rounded, sometimes 

Flowers congested along inflorescence axes. Calyx unequal; petiolules cylindrical, lenticellate, 3-7 mm long, 

(2.5) 4-5.2 mm long, sericeous-tomentose, the sepals minutely ferruginous-sericeous; rachis 3-11 cm long, 

ovate-deltate, ca. 1.5-2.3 mm long, obtuse at apex; sharply angled, minutely sericeous, striate, sometimes 

petals elliptic to nearly spatulate, obtuse at apex, long- lenticellate, and adaxially bisulcate on distal portion; petattenuate 

at base, 5-6.7 mm long, abaxially glabrous, ioles 3-9 cm long, flattened along adaxial surface, striadaxially 

papillate; appendages as long or slightly longer ate, sparsely tomentulose, slightly enlarged at the very 

than the petals, oblong or deltate, erect, adnate to at- base, sometimes lenticellate. Thyrses panicle-shaped, 

tenuate base of petal, adaxially villous, abaxially axillary or terminal, to 30 cm long,; cataphylls acicular, 

sparsely villous and papillate; disc poculiform, 5-lobed, ca. 4 mm long, minutely sericeous, clustered, axillary, 

hirsute, ca. 1 mm tall; stamens 8, the filaments unequal, early deciduous; axes angled, striate, minutely ferrugi- 

2.5-3.5 mm long, glabrous, the anthers 0.7-1.2 mm 

long, ellipsoid, glabrous, apiculate at apex. Fruits 

(young) ellipsoid, ferruginous, sericeous-velutinous, 

the pericarp thick-woody. 

This species differs from T. clathrata by its 

velutinous pubescence (vs. abaxially glabrous leaflets; 

puberulent stems and leaf rachis). The specific epithet 

refers to the indumentum of stems, leaf rachis and inflorescence 

axes. 

nous-sericeous; bracts subulate, tomentulose, 1-1.5 mm 

long, early deciduous; dichasia simple or compound; peduncle 

0.5-3 mm long, tomentulose; pedicels ca. 1.5 mm 

long, articulate at the middle or at base. Calyx 3-3.5 mm 

long, tomentose, the sepals free to the base, oblong or 

elliptic, obtuse at apex; petals elliptic, 5.5-5.7 mm long, 

erect, adaxially glabrous, abaxially glabrous, except for 

the appressed-pubescent lower third, the apex obtuse, 

the base cuneate to attenuate; appendages as long as the 

Phenology. Collected in flower in March and Sep- petals, adnate to the petal half oftheir length, deltate, erect, 

tember, and fruit in April. 

adaxially sericeous, abaxially puberulent papillate; disc 


from the lowlands of Colombia. [upland mixed Guadua 

forest] 

5-lobed, tomentulose, ca. 0.6 mm tall; stamens 8 (10), 

the filaments of unequal length, 3-4.5 mm long, pilose, 

the anthers ca. 1 mm long, ellipsoid, glabrous, apiculate 

at apex; ovary ovoid, ferruginous, shortly tomentose, the 

Representative specimens examined. COLOMBIA. stigma trigonous-capitate, papillate. Fruits ellipsoid, 

Antioquia: Mun. Chigorod6. Bohios trail, Fca. La Cabafia, densely ferruginous-sericeous, 1.8-3 cm long, apiculate


Fig. 61. Talisia veraluciana. A. flowering branch. B. detail of inflorescence. C. staminate flower. D. abaxial, 

adaxial, and lateral views of petal with adnate appendage. E. staminate flower with perianth removed showing 

disc and stamens. F. fruiting branch. (A-B from Oliveira et al. 242; C-E from Cavalcante 1840; F from Ramos 

& Mota 1675).


.ow 

m- , 7. . 0.W SN 5oI 4.W 

"


panicle-shaped thyrses, or exceptionally cauliflorous. mens 5-8, erect, the filaments of essentially equal length, 

Calyx with sepals connate to various degrees, 1/4 to 23 glabrous or with various indumenta, the anthers obthe 

length of calyx; petals with well developed petaloid long, lanceolate, or linear. Pollen brevi-colporate, obappendages 

with densely sericeous adaxial surface; sta- late, isopolar, triangular, with straight or concave sides. 

Key to the species of Talisia subgenus Talisia 

1. Stamens 5 (sometimes up to 8 in same plant). 

2. Leaflets with a conspicuous intramarginal vein 

3. Plant glabrous or nearly so. 

4. Leaflets 4-6, obovate or oblanceolate; disc glabrous .............................................................. T obovata 

4. Leaflets 8 or more, elliptic, narrowly elliptic, oblong-elliptic, oblong, or less often oblanceolate 

or falcate; disc variously pubescent. 

5. Leaflets abaxially minutely pubescent along veins, the apex acuminate, abruptly acuminate 

or apiculate; petiolules minutely pubescent; leaf rachis minutely pubescent, terete or 

sometimes obtusely angled, canaliculate or slightly compressed on distal portion; 

fruits 1-2 cm long .......................................................... T sylvatica 

5. Leaflets abaxially glabrous, the apex long-acuminate, acuminate or obtuse; petiolules 

glabrous; leaf rachis glabrous, slightly flattened dorsiventrally, sharply angled toward 

the margins on distal portion; fruits 1.8-2.5 cm long ..................................................... T nervosa 

3. Plant pilose, tomentose or minutely pilose. 

6. Leaf rachis terete, striate, pilose to tomentose; petals obovate, abaxially appressed-pubescent, 

clawed at base; filaments pilose ............................................................ T pinnata 

6. Leaflet rachis slightly flattened, minutely hirsute; petals elliptic, abaxially glabrous, cuneate or 

attenuate at base; filaments glabrous ........................................................................ T . pilosula 

2. Leaflets lacking a conspicuous intramarginal vein. 

7. Petals abaxially pubescent or puberulent, clawed or nearly so at base; disc tomentose. 

8. Leaf rachis trigonous, striate; leaflets elliptic, oblong or oblanceolate ............. ................. Tl ongifolia 

8. Leaf rachis pentagonous, bisulcate, not striate; leaflets oblong-elliptic or elliptic ............ T carinata 

7. Petals abaxially glabrous, cuneate at base; disc glabrous or pubescent .............................. T macrophylla 

1. Stamens 8 (seldom 7 or 9 in same plant). 

9. Petals abaxially pubescent or tomentose. 

10. Petals abaxially sericeous-tomentose or appressed-pubescent throughout. 

11. Leaf rachis sharply triangular; anthers lanceolate ............................................................ T marleneana 

11. Leaf rachis terete; anthers oblong. 

12. Stems puberulent; filaments of equal length, pilose; leaflets 6-8; petiolules not drying 

dark brown; calyx 2.5-3 mm long, the sepals deltate ............................................. T douradensis 

12. Stems glabrous; filaments of unequal length, glabrous; leaflets 11-15; petiolules drying 

dark brown; calyx 4-7 mm long, the sepals ovate or rounded ............................... T. megaphylla 

10. Petals abaxially pubescent or tomentose only at base. 

13. Filaments glabrous. 

14. Leaflets 2-7; leaflets abaxially glabrous; calyx puberulous or pubescent; petals adaxially 

papillate ......................................................... T ghilleana 

14. Leaflets 11-24; leaflets abaxially pubescent along veins; calyx tomentose; petals adaxially 

glabrous ............................................................... 

T croatii 

13. Filaments pilose, pubescent or tomentose. 

15. Stems glabrous. 

16. Leaflets 6-8, oblanceolate, or less often elliptic, abruptly acuminate at apex, abaxially 

glabrous; rachis terete; calyx puberulent; petals elliptic, adaxially glabrous; disc 

puberulent .......................................................7'. equatoriensis 

16. Leaflets 11-14, oblong or oblong-elliptic, obtuse at apex, abaxially puberulent along 

veins; rachis obtusely quadrato-rhombic; calyx tomentulose; petals obovate, adaxially 

puberulent; disc tomentose .............................................................T. bullata 

15. Stems pubescent. 

17. Inflorescence axes, bracts and calyx with scattered glandular hairs. 

18. Leaflets with clathrate tertiary venation ............................................................ . T morii 

18. Leaflets with reticulate tertiary venation. 

19. Stems cano-pubescent; disc tomentose at apex; anthers lanceolate; fruit glabrous, 

the pericarp to 5 mm thick .............T.................................... T princeps 

19. Stems minutely hirsute; disc hirsute; anthers oblong; fruit appressed-puberulent, 

the pericarp ca. 1 mm thick ..................................................T. pinnata


17. Inflorescence axes, bracts or calyx lacking glandular hairs. 

20. Leaflets lanceolate or elliptic-lanceolate; petals adaxially papillate, abaxially puberulent 

at base; petal appendage shorter than the petal .......................................... T acutifolia 

20. Leaflets oblanceolate or oblong; petals adaxially glabrous, abaxially tomentose at 

base; petal appendage as long as the petal. 

21. Stems, abaxial side of leaflets, and leaf rachis canescent; filaments sparingly 

wooly pubescent; fruits appressed ferruginous-pubescent . ............................ T stricta 

21. Stems, abaxial side of leaflets, and leaf rachis setigerous; filaments tomentose; 

fruits puberulent .................................................................T. setigera 

9. Petals abaxially glabrous. 

22. Stems and abaxial side of leaflets glabrous; leaf rachis glabrous or puberulent. 

23. Filaments of unequal length. 

24. Inflorescence fasciculate, axillary, ramiflorous or cauliflorous, less often simple or 

panicle-shaped; sepals connate for l/2 to 3/4 the length of calyx ................................. T hexaphylla 

24. Inflorescence solitary, axillary or terminal, never cauliflorous nor fasciculate; sepals 

connate for less than l/2 the length of calyx. 

25. Tree to 16 m tall; stems terete, lacking lenticels; leaf rachis terete or slightly angled, 

puberulent ...................................................T. laevigata 

25. Shrub; stems slightly sulcate, lenticellate; leaf rachis cross-section crescent-shaped, 

adaxially bicanaliculate on distal portion, glabrous ................................................ T oedipoda 

23. Filaments of equal length. 

26. Calyx poculiform .......... ................................................ T cupularis 

26. Calyx cup-shaped. 

27. Filaments, ovary and disc glabrous (disc sometimes hispidulose). 

28. Treelet; leaflets 16-22, with acuminate apices; petals 6-7.5 mm long, abaxially 

glabrous ................................................T. guianensis 

28. Tree; leaflets 2-12, with rounded, retuse and mucronate apices; petals 4.8-6.2 

mm long, abaxially appressed pubescent at base ............................................... T. retusa 

27. Filaments pilose, ovary sericeous or tomentose, and disc pubescent or tomentose. 

29. Petals oblong; disc tomentose, tomentulose, pubescent or less often glabrous; 

anthers oblong; leaflets strongly asymmetrical at base ....................................... T cerasina 

29. Petals ovate-lanceolate; disc tomentose; anthers lanceolate; leaflets symmetrical or 

slightly asymmetrical ............................................................. T eximia 

22. Stems, abaxial side of leaflets, and leaf rachis tomentose or hirsute. 

30. Leaflets with a strong intramarginal vein; thyrses terminal or cauliflorous ................... T dasyclada 

30. Leaflets without a strong intramarginal vein; thyrses terminal or axillary. 

31. Leaflets abaxially minutely hirsute over the entire surface or less often only on veins. 

32. Inflorescence axes, bracts, bracteoles and calyx hirsute; petals oblong, cuneate-attenuate 

at base; disc minutely hirsute at apex; fruits ovoid to globose, minutely hirsute ..... T mollis 

32. Inflorescence axes, bracts, bracteoles and calyx velutinous; petals elliptic, longattenuate 

at base; disc velutinous at apex; fruits trigonous (in x.s.), sericeoustomentose 

.............................................. T pachycarpa 

31. Leaflets abaxially glabrous or pubescent only along veins. 

33. Stems tomentose; leaflets 10-14, acuminate at apex; rachis minutely hirsute; petiolules 

pulvinate; dichasia compound, sessile ........................................ T hemidasya 

33. Stems tomentulose; leaflets 16-24, caudate at apex; rachis tomentulose; petiolules 

cylindrical; dichasia simple, short pedunculate ........................................ T caudata 

20. TALISIAACUTIFOLIA Radlkofer, Sitzungsber. pelago Anavilhanas, 3 Mar 1976 (fl), Medri 33 

Math.-Phys. Cl. K6nigl. BayerAkad. Wiss. Miinchen (holotype, INPA). 

8: 349. 1878. Type. Brazil. Amazonas: Uanauca- 

Gapo, Dec 1851 (fl), Spruce 1992 (lectotype, K, 

here designated, photo at US; isolectotype, K-2, 

P). Syntypes. Brazil. Amazonas: Manaus, along 

Taruma-Aqu creek, Feb 1855 (fr), Spruce 1992 (K- 

3); Rio Negro, s.d. (fl), Martius s.n. (M). 

Figs. 4a, 9b, 63a-h 

Treelet 1.5 to 4 m tall, sometimes with few upright 

branches; trunk ca. 2.5 cm in diam. Stems terete, 

smooth or minutely lenticellate, puberulent or pubescent. 

Leaves paripinnate; distal process acicular, ca. 3 mm long, 

early deciduous, leaving a truncate base; leaflets (4) 6-8 

(10), alternate to opposite, lanceolate to elliptic-lanceolate, 

6-3 1 x 2.4-10 cm, coriaceous, glabrous except 

Talisia ,nedri G. Guarim Neto, Acta Amaz. 9: 235. for the abaxially, minutely pilose midvein, the venation 

1979. Type. Brazil. Amazonas: Rio Negro, Arqui- brochidodromus with a slightly conspicuous marginal


3mm. 

B ff 

mm. 3 

]3cm. 

Fig. 63. Talisia acutifolia. A. flowering branch and leaf. B. staminate flower. C. adaxial and abaxial views of 

petal with adnate appendage. D. staminate flower with perianth removed showing disc and stamens. E. stamen. 

F. pistillate flower with perianth removed showing pistil and sterile stamens (left) and l.s. same (right). G. steril 

stamen. H. fruit. (A-G from Liesner 4162; H from Liesner 8687).


loop, plane on adaxial surface, prominent on abaxial Common names and uses. Known in Brazil as 

surface, tertiary veins reticulate, the margins sometimes pitomba, pitomba amarela, pitombeira, and olhio de 

revolute, the apex long-acuminate, acuminate or less venado. The seeds have a fleshy, edible coat, consumed 

often acute, the base acute-obtuse, sometimes unequal; by the local people. 

petiolules pulvinate, 0.5-1 cm long, minutely pubes- Representative specimens examined. VENEZUELA. 

cent; rachis 3-22 cm long, terete, minutely pubescent, Amazonas: San Carlos de Rio Negro, 120 m, 3 Dec 1977 

glabrescent; petioles 7.5-15 cm long, gradually thick- (fl), Liesner 4162 (MO, NY, VEN), 120 m, 25 Jan 1980 

ened toward base. Thyrses panicle-shaped, sometimes (fr), Liesner 8687 (MO, NY, VEN); Confluence of Negro 

fasciculate, terminal or supra-axillary; cataphylls flat- & Casiquiare rivers, 120 m, 15 May 1979 (fr), Liesner 

tened-acicular or sub-pinnate, 1.7-2 cm long, tomen- 7439 (MO, VEN); Casiquiare river above Chapez6n, 

between Solano and Boca, 120 m, 31 Jan 1980 (fr), 

tose; axes to 20 cm long, terete, ferruginous-tomentose; 

Liesner & Clark 8906 (MO, VEN); Rio Negro, vic. San 

bracts triangular, persistent, 1.5-2 mm long; dichasia 

Carlos de Rio Negro, 23-29 Jul 1982 (fr), Stergios & 

simple or compound, sessile or shortly pedunculate; Aymard 4213 (PORT). 

pedicels to 1.5 mm long, articulate at the apex. Calyx BRAZIL. Amazonas: Rio Negro, ca. 100 m, 16 Aug 

2.5-3 mm long, tomentulose, the sepals 1-1.5 mm long, 1996 (fl), Acevedo R. et al. 8362 (INPA, NY, US); 16 

oblong, concave to slightly keeled, rounded at apex; Aug 1996 (fr), Acevedo R. et al. 8414 (INPA, US); Rio 

petals oblanceolate to elliptic, 4-4.5 mm long, reflexed Preto, ca. 100 m, 17 Aug 1996 (fl), Acevedo R. et al. 

8442 (INPA, US), (fl), Acevedo R. et al. 8456 (INPA, 

at anthesis, papillate on adaxial surface, glabrous on 

US); Rio Taruma-Mirim, 29 Apr 1976 (fr), Adis 7 (MG); 

abaxial surface except for a few hairs near the base, the 

Parana' de Sao Jose de Ariranha, 11 Feb 1944 (fl), 

apex sometimes slightly retuse, the base attenuate; ap- Baldwin 3290 (F, IAN, NY, US); Reserva Ducke, 

pendages slightly shorter than petals, oblong, erect, Manaus-Itacoatiara Hwy, km 26, 80 m, 19 Jan 1990 (st), 

entire or bifid at apex, sericeous-tomentose on both sur- Gentry & Nelson 69247 (MO); Reserva Ducke, Nov 1972 

faces; disc cup-shaped, 5-lobed, hirsute, ca. 0.8 mm (st), Rodrigues 9227 (INPA); Maues, Parauari river, 

tall; stamens 8, the filaments of equal length or slightly Mucaja, 7 Dec 1961 (fl), Rodrigues & Coelho 3925 

unequal, 4-5 mm long, minutely pilose, the anthers (INPA-2, NY); Upper Rio Negro, Ilha da Bateria, 29 Nov 

1947 (fl), Schultes & L6pez 9246 (IAN); Rio Negro, ca. 

1.2-1.6 mm long, oblong-lanceolate, glabrous, apicu- 

Ilha da Silva, 16 Jan 1978 (fl), Steward et al. 378 (K, 

late at apex; ovary nearly conical, tomentose, the stigma 

NY); Manaus, Cachoeira Grande, 27 Dec 1874 (fl), Traill 

obconical to capitate. Fruits ellipsoid to ovoid, glabrous, 125 (K); Barcelos, along Rio Negro, 1 Jan 1902 (fl), 

(trigono-obovoid and appressed-pubescent when Ule 6037 (INPA, K, M). 

young), turning from green to yellow, 1.7-2.5 (3) cm long, 

minutely granulate to smooth, apiculate, the pericarp 

0.7-0.8 mm thick, the endocarp granulate, glabrous. 21. TALISIA BULLATA Radlkofer in Martius, Fl. 

Seeds ellipsoid, with fleshy testa; cotyledons superim- Bras. 13(3): 556. 1900. Type. Ecuador. Manabi: 

posed horizontally or obliquely, the lower one slightly Rosario, Jul 1893 (fl), Eggers 15004 (lectotype, 

larger to twice as large as the upper one, sometimes M, here designated; isolectotypes, B-destroyed, C, 

poorly differentiated in the area adjacent to the radicle. n.v., K, US, photo of B (F neg. 5671) at US, frag. 

Two collections with different dates and localities at MO. Syntype. Ecuador. without locality, Aug 

were distributed under Spruce 1992 (both annotated by 1893 (fl), Zapullo s.n. (?). Fig. 64a-g 

Radlkofer). The first is from Taruma-Agu creek, col- 

Treelet 8-12 m tall; bark grayish, lenticellate. Stems 

lected in February of 1855. The second collection is 

terete, glabrous, lenticellate. Leaves paripinnate or less 

from Uanauca-Gapo and was collected in December 

often imparipinnate; distal process acicular, deciduous, 

of 1851. I have designated the 1855 collections as lec- leaving a truncate base ca. 4 mm long; leaflets 11-14, 

totypes and isolectotypes because there are more du- alternate to opposite, oblong to oblong-elliptic, 23-79 

plicates available for this collection than for the 1851 x (7.5) 9-21.5 cm, the distal and basal leaflets smaller 

collection. The specific epithet refers to the character- than the median ones, rigidly coriaceous, bullate, the 

istic leaflets with acute or acuminate apices. adaxial surface with deeply sunken veins, glabrous, 


February and in August; known to fruit in January, May 

and August. 

the abaxial surface with prominent, puberulent veins, 

the venation brochidodromus, with secondary veins alternate, 

arching, nearly parallel, forming a strong submarginal 

vein, the margins strongly revolute, the apex 

Distribution and Ecology. (Fig. 67) From central obtuse, the base obtuse-rounded, unequal or slightly 

to westem Amazon toward Rio Negro area, in varzea so; petiolules pulvinate, slightly flattened along adaxial 

and igapo forests. 

surface, 1.2-1.7 x 0.5-1.0 cm, puberulent to glabrous;


Fig. 64. Talisia bullata. A. habit. B. leaf. C. inflorescence with detail of denuded dichasium. D. flower bud. E. 

mature pistillate flower. F. lateral, abaxial, and adaxial views of petal with adnate appendage. G. pistillate flower 

with perianth removed showing, disc, sterile stamens, and pistil (left) and ls. same showing ovary, ovules, and 

sterile stamens with detail of sterile stamen (right). (A-B from Acevedo R. et al. 6909; C-G from Eggers 15004).


rachis 37-77 cm long, obtusely quadrato-rhombic, to (fl), Poiteau s.n. (K-2, P); without specific locality 

transverse-rhombic, firrowed, puberulent or glabrous; 

or date (fl), Richard s.n. (P-3). Fig. 65a-h 

petioles 30-45 cm long, terete, striate, greatly enlarged Talisia carinata forna sericea Radlkofer in Martius, 

at base, slightly depressed along adaxial surface. Th- Fl. Bras. 13 (3): 549. 1900. Type. French Guiana. 

yrses panicle-shaped, apparently terminal; cataphylls Montagne de Kaw, Roura, 1858 (fl), Sagot s.n. 

(lectotype, P, here designated; isolectotype, M). 

acicular, 4-5.5 cm long; axes more than 50 cm long, 

Syntypes. French Guiana. Cayenne, without date 

secondary branches to 20 cm long, angled, sulcate, 

(fl), Martin s.n. (K, n.v., P); without specific lodensely 

covered with minute, cinnamomeus hairs, and cality or date (fl), Poiteau s.n. (K-2, P); without 

scattered glandular hairs; bracts ovate to deltate, per- specific locality or date (fl), Richard s.n. (P-3). 

sistent, 1-1.6 mm long; dichasia compound, sessile, 

congested along axes; pedicels to 1.5 mm long, tomen- Treelet to 3 (7) m tall, unbranched or with a few uptose, 

articulate at the apex. Calyx 3-3.5 mm long, right branches on distal portion, these with determinate 

tomentulose, the sepals ca. 2 mm long, ovate, concave, growth due to a terminal inflorescence; trunk to 5 cm in 

rounded at apex, ciliate at margins; petals obovate, 6- diam., bark grayish, smooth. Stems terete, minutely 

6.5 mm long, reflexed at anthesis, puberulent on adaxial lenticellate, puberulent, glabrescent. Leaves paripinnate 

surface, tomentose on lower half of abaxial surface, the or less often imparipinnate; distal process acicular, early 

apex rounded, the base attenuate-clawed; appendages deciduous leaving a truncate base 2-3 mm long; leaflets 

as long as the petals, lanceolate, erect, sericeous on both 8-16 (26), alternate or subopposite, oblong-elliptic or 

surfaces; disc cup-shaped, 5-lobed, tomentose, ca. 1 mm elliptic, (5) 7.2-21 (27.5) x (2.5-) 3-5.5 (-7.9) cm, 

tall; stamens 8, the filaments of equal length, ca. 1.5 mm chartaceous to subcoriaceous, glabrous, the adaxial surlong, 

pubescent, the anthers ca. 1.3 mm long, oblong, face with a prominulous midvein, and slightly sunken 

glabrous, apiculate at apex; ovary ovoid, densely fer- secondary veins, the abaxial surface with prominent, 

ruginous-tomentose, the stigma nearly capitate. Fruits puberulent veins, the venation brochidodromus, with 

ellipsoid, granulate, puberulent or glabrous, 2.6-3 cm secondary veins altemate or subopposite, arching, terlong, 

apiculate at apex, the pericarp coriaceous, ca. 1 tiary veins reticulate, the margins undulate, the apex acumm 

thick, the endocarp puberulent. Seed one per fruit, minate to long-acuminate, the base oblique, one side 

ellipsoid. Embryo with cotyledons laying obliquely obtuse the other attenuate, tapering on to the petiolule, 

dorsiventral over each other, the upper one slightly distal leaflets usually larger than the proximal ones; petilarger 

than the lower one. 

olules pulvinate at base, canaliculate along adaxial sur- 

The specific epithet refers to the blistered appear- face, 3.5-10 mm long, glabrous to puberulent, the pulviance 

of the leaflets. 

nus obconical to cylindrical, 3.5-5 mm long; rachis (12) 

18-52 cm long, pentagonous, bisulcate, obtusely to 

Phenology. Collected in flower during July and narrowly angled, nearly terete toward base, glabrous or 

August, and in fruit in May and September. puberulent, smooth; petioles 10-17 (27) cm long, ter- 

Distribution and Ecology. (Fig. 67) In coastal, non- ete, pulvinate at base. Thyrses panicle-shaped, terminal 

flooded rainforest of westem Colombia and Ecuador. or axillary on distal portion of main stem and branches, 

or less often cauliflorous; cataphylls acicular, clustered, 

Representative specimens examined. COLOMBIA. supra-axillary, 5-12 mm long, persistent or deciduous; 

Antioquia: San Luis, along Medellin-Bogota hwy, ca. km axes (13.5-) 19-25 (-68) cm long, the secondarybranches 

25, 500-630 m, 23 May 1990 (fr), Cogollo et al. 4572 

6-10 (-21) cm long, angular, usually sulcate, reddish 

(MEDEL). Choc6: Mun. Nuqui; Corregimiento Termales, 

tinged, sparsely to densely covered with minute, whitcoastal 

zone between Piedra Piedra & Terco river, 9 Sep 

ish hairs; bracts ovate, persistent, ca. 0.5 mm long; di- 

1994 (st), Acevedo R. et al. 6909 (F, HUA, K, NY, US). 

ECUADOR. Esmeraldas: Eloy Alfaro. San 

chasia 

Miguel, 

compound, sessile or shortly pedunculate; pedicels 

Cayapas river, 100 m, 3-5 Sep 1993 (fr), Palacios & 0.5-1 mm long, articulate at the middle. Calyx 1-1.8 mm 

Tirado 11097 (US). 

long, puberulent to pubescent, the sepals 0.5-0.6 mm 

long, ovate, concave, rounded at apex; petals white, oblong, 

truncate at base, 2.5-3.6 (-4.2) mm long, reflexed 

22. TALISIA CARINATA Radlkofer, Sitzungsber. at anthesis, adaxially glabrous or papillate, abaxially ap- 

Math.-Phys. Cl. K6nigl. Bayer Akad. Wiss. pressed-pubescent, especially on lower half, the apex 

Miinchen 8: 349. 1878. Type. French. Guiana. rounded, the base cuneate-obtuse; appendages as long 

Montagne de Kaw, Roura, 1858 (fl), Sagots.n. (lec- as or a little shorter than the petals, lanceolate, erect, 

totype P, here designated; isolectotype, M). Syntypes. entire at apex, adaxially sericeous-tomentose, abaxially 

French Guiana. Cayenne, without date (fl), Martin sparsely sericeous-tomentose or glabrous; disc annus.n. 

(K, n.v., P-2); without specific locality or date lar, 5-lobed, pubescent to tomentose, 0.4-0.5 mm tall;


Fig. 65. Talisia carinata. A. inflorescence. B. staminate flower. C. i.s. staminate flower showing disc, stamens, 

and pistillode. D. adaxial and lateral views of petal with adnate appendage. E. staminate flower with perianth 

removed showing disc and stamens. F. leaf. G. fruiting branch. H. seed. (A-E from Mori et al. 19109; F-H from 

Mori et al. 14955). 

2 mm.


stamens 5 (6), the filaments of equal length, 1.5-2.2 mm 

long, pilose, the anthers 1.1-1.3 mm long, oblong, glabrous, 

apiculate at apex; ovary ovoid, densely ferruginous-tomentose, 

the stigma capitate. Fruits 1-2-seeded, 

ellipsoid to obovoid, turning from green to orange-yellow, 

sparselypubescent, glabrescent, granulate, 1.5-2.1(- 

2.5) cm long, rounded at apex, shortly apiculate, the 

pericarp 0.4-1.2 mm thick, the endocarp granulate, glabrous. 

Seeds ellipsoid, to 1.5 cm long, with fleshy, edible, 

cream testa; cotyledons superimposed, of equal size. 

Talisia carinata seems to be closely related to T 

sylvatica. However, T carinata differs from T sylvatica 

by: secondary veins 8-11 (vs. 11-20); abaxial leaf 

surface without granules (vs. punctate or granulate); 

venation and leaf rachis glabrous (vs. puberulent); and 

petals pubescent sericeous adaxially (vs. glabrous). In 

addition, Talisia sylvatica on the average, has larger 

leaflets and fewer per leafthan T carinata. A few specimens 

of T carinata with large leaflets resemble T 

acutifolia, however, T carinata differs by its angled 

leaf rachis (vs. terete), chartaceous and more numerous 

leaflets (vs. coriaceous and fewer leaflets), and by 

its flowers with abaxially appressed-pubescent petals 

(vs. glabrous), and 5 stamens (vs. 8). 

The specific epithet refers to the abaxially keeled 

midvein of the leaflets. 

Phenology. Known to flower from May to Octo- 

ber and to fruit all year round. 

Distribution and Ecology. (Fig. 67) Guianas and 

Brazil. An understory treelet of terra firme and seasonally 

flooded, lowland, rain to dry forests. 

Common names and uses. French Guiana. Creole: 

Dayacoussa, gaulette indienne, bois flambeau, 

payacoussa, picoussa, petite gaulette, tit-gaulette; 

Oyampi: touliatan; Wayapi: tuliata. Surinam. 

Bosknippa. The fruits, although of small size have an 

edible juicy testa covering the seed. 

Representative specimens examined. SURINAM. 

Nickerie: Kabalebo Dam project, 30-130 m, 20 Sep 

1980 (fr), Lindeman et al. 501 (MO). 

FRENCH GUIANA. Without locality, 3 May 1901 (fl), 

Geay 993 (P-2), 18 Aug 1969 (fl), Irwin et al. 47630 (IAN, 

NY, P, US-2); s.d., Poiteau s.n. (K). Bassin de 

L'Approuague: Station des Nouragues, 26 Jan 1989 (fr), 

Loubry 163 (CAY, US); Arataye river, top of Saut Parar6, 

40 m, 3 Feb 1969 (fr), Oldeman 2922 (CAY, P, US); 

Approuague river toward Parepou creek, 25 Sep 1968 (fl), 

Oldeman B-1871 (CAY, P, US). Bassin de la Camopi: 

D6grad Claude, Tamouri, 20 Mar 1974 (fr), Lescure 169 

(CAY-2, P); Roche Jose, 7 Dec 1967 (fr), Oldeman 2597 

(CAY-2, P, US); Grand Tamouri, 200 m from Camopi river, 

13 Feb 1968 (fr), Oldeman & Sastre 219 (CAY-2, IAN, 

NY, U-2). Bassin de L'Inini: Without specific locality, 7 

Apr 1986 (fr), Feuillet 3692 (CAY); Montagne Bellevue 

de L'Inini, 550 m, 15 Aug 1985 (fr), de Granville et al. 

7500 (CAY-2, U), 550 m, 30 Aug 1985 (fl), de Granville 

et al. 7919 (CAY, P, US); Monts Atachi Bakka, 440 m, 8 

Jan 1989 (fr), de Granville et al. 10480 (CAY, US); Grand 

Inini river, ca. mouth of Sai creek, 26 Aug 1970 (fl), 

Oldeman B-3578 (CAY). Bassin de L'Iracoubo: Iracoubo 

river, old trail to Saut Franconie, 12 Aug 1966 (fl), Oldeman 

B-595 (CAY); Maniga village, 18 Aug 1966 (fl), Oldeman 

2223 (CAY-2, P). Bassin de la Mana: Mana river, vic. of 

Saut Fracas, 22 Jul 1981 (fl), Cremers 7275 (CAY, P); 

Baboune creek, 15 km from La Mana, 31 Jul 1981 (fl), de 

Granville 4710 (CAY, P); Montagnes de la Trinite, 430 m, 

6 Feb 1984 (fr), de Granville et al. 6536 (CAY-2, NY, P, U, 

US-2). Bassin de L'Oyapock: Gabaret creek, 5 Sep 1996 

(bd), Blanc 120 (CAY, US); Pic du Grand Croissant, 40 

km N of Camopi, 8 Dec 1983 (fr), Feuillet 1221 (CAY, P); 

Camopi, 1900 (fr), Geay 852 (F, P); Upper Oyapock river, 

W of Trois Sauts, Mont Saint Marcel, 21 Jul 1975 (bd), de 

Granville T-1169 (CAY-2), 28 Jul 1975 (st), de Granville 

T-1205 (CAY-2, P); Zidock village, 3 Nov 1973 (fr), 

Grenand 58 (CAY); Trois Sauts, Oyapock river, 2 Jan 1975 

(fr), Grenand 618 (CAY); Maripa, 18 May 1965 (fr), 

Oldeman 1278 (CAY-2, P, US); Montagne Trois Pitons- 

Crique Ouanary, 6 Aug 1969 (fl), Oldeman B-2582 (CAY, 

P); Eureupoucigne creek, affluent of Oyapock river, 22 

May 1970 (fr), Oldeman B-3321 (CAY, P, US); Upper 

Oyapock, Zidock village, 12 Aug 1985 (fl), Prevost & 

Grenand 1934 (CAY, P, US). Bassin du Maroni: Village 

Touinke, 12 Apr 1977 (fr), Cremers 4689 (CAY-2); Gobaya 

Soula, 1.5 km from L'Itany, 100 m, 6 Jan 1989 (fr), de 

Granville et al. 10439 (CAY); 6 km E of Gobaya Soula, 

420 m, 10 Jan 1989 (fr), de Granville et al. 10537 (CAY, 

NY, US); 9 km SE of Gobaya Soula, 15 Jan 1989 (fr), de 

Granville et al. 10657 (CAY, US), 9 Mar 1971 (fr), de 

Granville C-148 (CAY-2, P); Comte river, Roche Fende, 

18 Jan 1977 (fr), Mori & Veyret 8929 (CAY, MO, NY). 

Bassin du Sinnamary: Camp Eugene, 70 m, 4 Feb 1995 

(fr), de Granville & Cremers 12752 (CAY, NY, P, US); 80 

m, 29 Nov 1994 (fr), de Granville et al. 12634 (CAY, US); 

Sinnamary river, 1 May 1969 (fl), Oldeman B-2322 (CAY, 

P). Bassin de la Yaloupi: Saut Tainoua, 17 Apr 1970 (fr), 

de Granville 376 (CAY-4, IAN, NY); Yaloupi river, near 

Saut Coueki, 30 Apr 1970 (fr), de Granville 512 (CAY-2); 

Saut Tainoua, 17 Mar 1970 (fr), Oldeman T-514 (CAY, P, 

US). Ile de Cayenne: Grand Matoury, NE side, Reserve de 

La Mirande, 21 Apr 1992 (fr), Acevedo R. & Grimes 4797 

(CAY, K, NY, US-2); Mont Rorota, 24 Apr 1992 (fr), 

Acevedo R. & Grimes 4837 (CAY, US); Mont Cabassou, 

100 m, 2 Apr 1996 (fr), Cremers et al. 14360 (CAY, NY, P, 

US), (fr), Cremers et al. 14361 (CAY-2); Cayenne, Martin 

s.n. (K); Montagne Tigre, 140 m, 3 Feb 1965 (fr), Oldeman 

1052 (CAY-2, US-2); Montagne Mahoury, 12 km E of 

Cayenne, 250 m, 6 May 1965 (fl, fr), Oldeman 1265 (CAY- 

3, NY, P, US); Mont Rorota, 23 Mar 1984 (fr), Prevost 

1496 (CAY-2, US). Montagne de Kaw: Route de Belizon, 

19 Feb 1988 (fr), Billiet & Jadin 4439 (CAY-2); Roura 

along road to Gabrielle creek, 2 Feb 1977 (fr), Cremers 

4243 (CAY-2, MO, NY); Montagne de Kaw, 225-270 m, 

13 Dec 1954 (fr), Cowan 38795 (NY-2, US-2). Piste de 

Saint Elie: Station Biologique ORSTOM, 27 Apr 1992 (fr),


Acevedo R. et al. 4842 (CAY, US); between Sinnamary & cylindrical, 5-10 mm long, puberulent; rachis > 35 cm 

Counama rivers, 23 Apr 1982 (fr), Anonymous 374 (CAY- long, terete, densely covered with minute ferruginous 

2); Station Biologique ORSTOM, Mar 1980 (fr), Lescure hairs; petioles ca. 12 cm long, terete, slightly pulvinate 

893 (CAY-2); between Sinnamary & Counama rivers, 19 

at base. Thyrses panicle-shaped, terminal on distal por- 

Aug 1994 (fl), Prevost 3055 (CAY, US). Region des 

Emerillons: Massif des Emerillons, 200 m, 5 Sep 1980 (fl), 

tion of main stem; cataphylls acicular, 4-5 mm long, 

de Granville 3738 (CAY-2). Region Littorale: Kourou. early deciduous; axes to 15 cm long, the secondary 

Passoura, near Passoura creek, 29 Apr 1992 (fr), Acevedo branches to 7 cm long, angled, sulcate, densely cov- 

R. et al. 4872 (CAY, NY, US), 6 May 1992 (fr), Acevedo ered with glandular, multiseptate, yellowish hairs and 

R. et al. 4915 (CAY, K, NY, US). Region de Paul- Isnard: with minute, erect, simple hairs; bracts lanceolate, per- 

Paul Isnard, vic. of Citron, 11 Feb 1983 (fr), Cremers s.n. sistent, 6-7 mm long, with same indumentum as the 

(US); Paul Isnard, base of Montagne Lucifer, 430 m, 11 axis; dichasia simple, shortly pedunculate, sometimes 

Feb 1983, (fr), Cremers, s.n. (US); Citron, 11 Feb 1983 

reduced to a single flower due to the abortion of lateral 

(fr), Cremers 7961 (CAY-2, P, US); along recent road to 

Citron, 11 

flowers; pedicels 0.5-1 mm 

Feb 1983 (fr), Feuillet 698 

long, articulate at the apex. 

(CAY); Paul Isnard, 

base of Montagne Lucifer, 430 m, 9 Nov 1982 (fr), de Calyx urceolate, pinkish, ca. 3 mm long, with same in- 

Granville 5235 (CAY-2, P). Region de Saul: Route de dumentum as the inflorescence axes, the sepals 2-2.5 

Belizon, Eaux Claires, 26 Mar 1981 (fr), de Granville 4475 mm long, ovate, concave-carinate, obtuse at apex; pet- 

(CAY, P); Mont Belv6dere, 160 m, 22 Nov 1984 (fr), de als ovate-lanceolate, white, 4.5-5.5 mm long, erect, 

Granville 6952 (CAY, P, U), 7 Dec 1984 (fr), de Granville adaxially puberulent with a few glandular hairs, 

7161 (CAY); Mont Galbao, 570 m, 22 Jan 1986 (fr), de abaxially glabrous, the apex obtuse, the base truncate; 

Granville et al. 8890 (CAY, P), 27 Jul 1979 (fl), de Granville appendages 2-3 mm long, triangular-lanceolate, erect, 

B-5507 (CAY); Montagne Grand Fosse, 20 Aug 1971 (fl), 

entire at apex, villous on both surfaces; disc annular, 5- 

Oldeman B-4051 (CAY-2, P); Chamberlain, km 3 from 

lobed, tomentulose, ca. 0.5 mm tall; stamens 7 or 8, the 

Galbao, 8 Feb 1978 (fr), Prevost 184 (CAY). 

BRAZIL. Amapl: Oiapoque, 92 km SE of Oiapoque, filaments of equal length, ca. 0.3 mm long, glabrous, 

5 Dec 1984 (fr), Daly et al. 3822 (K, MG); Oiapoque, 13 the anthers 1.4-1.5 mm long, oblong, glabrous, apicu- 

Oct 1950 (fr), Froes 26617 (IAN); Mt. Tipac, 0-200 m, late at apex. Pistillate flowers unknown. Fruits ellip- 

13 Oct 1960 (fr), Irwin 48688 (MG, MO, NY, US); soid (trigonous when young), yellowish brown, 2.8-3 

Oiapoque river, vic. of Tres Saltos Fall, 10 Sep 1960 (fr), cm long, puberulent, granulate, rounded at apex, with 

Irwin et al. 48154 (IAN, MG, NY, US), Agua Azul, 26 Jul short apiculum, the pericarp 1 mm thick, the endocarp 

1962 (fl), M. Pires & Cavalcante 52291 (F, IAN, MG, 

granulate, sparsely furfuraceous. Seeds ellipsoid, to 2.4 

NY, US), 30 Jul 1962 (fr), M. Pires & Cavalcante 52298 

cm 

(F, IAN, MG, NY); 84 km SE of Oiapoque, 5 Dec 1984 

long; cotyledons superimposed, of equal size. 

(fr), Rabelo & Cardoso 2888 (MG, NY). 

The specific epithet refers to the caudate apices of 

the leaflets. 

23. TALISIA CAUDATA Steyermark, Ann. Missouri 

Bot. Gard. 75: 1072. 1988. Type. Venezuela. 

Phenology. Collected in flower during November 

and in fruit during August and October. 

Amazonas: Cerro Sipapo, mixed montane forest, Distribution and Ecology. (Fig. 67) A treelet en- 

125 m, 25 Jan 1949 (fr), Maguire & Politi 28615 demic to Venezuela, known only from river banks in 

(holotype, MO-2 sheets; isotypes, MG, NY, US). laja and rain forests. 

Fig. 66a-i 

Common name. Kaerinuae ukwaewaesa. 

Treelet 2-3.5 m tall. Stems terete, puberulent. Leaves 

paripinnate; distal process filiform, 5-13 mm long; leaflets 

16-24, alternate to opposite, oblong to oblong-lanceolate, 

9-13.5 x 2-3.5 cm, chartaceous, glabrous, the 


ZUELA. Amazonas: Atures, Alto Carinagua, 4 Sep 1995 

(fr), Contreras 94 (VEN); Atures, Cerro Uchonhua, 120- 

150 m, 9 Nov 1980 (fl), Gudnchez 383 (TFAV); Left 

adaxial surface with prominulous midvein, the abaxial bank, tributary of Cuao river, 350-400 m, 15 Aug 1985 

surface with prominent venation, the midvein puberu- (st), Zent 885-20 (US), 400-450 m, 5 Oct 1985 (fl), 

lent, sometimes with multicellular glandular trichomes, 

the venation brochidodromus, with secondary veins 

alternate or subopposite, arching, forming a submar- 

Zent 1085-03 (US). 

ginal loop, smaller loops formed between the margin 24. TALISIA CERASINA (Bentham) Radlkofer, 

and the submarginal loops, tertiary veins reticulate, the Sitzungsber. Math.-Phys. Cl. Konigl. Bayer Akad. 

margins entire, the apex caudate (this 1.8-2.2 cm long), Wiss, Miinchen 8: 349. 1878. Sapindus cerasinus 

the base oblique, one side obtuse the other attenuate, Bentham, J. Bot. (Hooker) 3: 197. 1851. Type. Brazil. 

tapering into an elongated petiolule; petiolules nearly Para: vicinity of Santarem, Apr-Aug 1850 (fl),


Fig. 66. Talisia caudata. A. leaf. B. inflorescence. C. detail of inflorescence branch and trichome. D. staminate 

flower. E. adaxial and abaxial views of petal with adnate appendage. F. I.s. staminate flower with perianth removed 

showing disc, stamens, and pistillode and same (entire). G. pistillode. H. portion of infructescence. I. dorsal view 

of seed (left) and embryo (right). (A-G from Zent 1085-03; H-I from Zent 0885-24).


n 

9oW! 

< 

\; 

YS 

- . 70WI 60W 

'.~~~~~~~~~~~~~~~~\ 

~ ~ ~ ~ P 

~~~~~~~r 

50W 40W 

I ON 

o _W......t >*. 

* Talisia acutfWolia < 0 

A Talisia buflata 

vTalisia 

; - 

carinata ~ I ~ r ~I' 

* Talisia caudata \ ( ) ?" 

Fig. 67. Distribution of species in Talisia subgenus Talisia (in part). 

Spruce 1035 (lectotype, K, here designated; 

isotypes, K-2, M, P, photo of M (F neg. 6018) at 

NY and US). Fig. 68a-h 

hispidulose to tomentose, the secondary branches to 

17 cm long,angled, sulcate; bracts subulate to deltate, 

persistent, 2.5-5 mm long, with same indumentum as 

Sapindus oblongus Bentham, J. Bot. (Hooker) 3: 198. the axis, sometimes with glandular hairs; dichasia simple 

1851. Type. Brazil. Pari: vicinity of Santarem, Apr 

1850 (st), Spruce s.n. (holotype: K, photo at US). 

or compound; peduncles 0.5-5 mm long; pedicels 2-3 

mm long, articulate at or slightly above the middle. Calyx 

Treelet or small tree, 2-4(12) m tall. Stems terete, 

smooth or minutely lenticellate, glabrous. Leaves 

paripinnate; distal process truncate, 2-3 mm long; leaflets 

(6)10-16(18), alternate or subopposite, 5-32(40) 

x 1.2-7.2 (18) cm, falcate, elliptic, oblong, lanceolateelliptic, 

ovate-elliptic or oblanceolate, chartaceous to 

coriaceous, glabrous, the adaxial surface glossy, with 

prominent midvein and sunken or plane secondary veins, 

the abaxial surface with numerous glandular dots and 

prominent midvein and secondary veins, the venation 

brochidodromus, lighter than the blade, with secondary 

veins alternate or subopposite, arching toward the 

margin, tertiary veins reticulate, the margins entire, revolute 

or slightly so, the apex short- to long-acuminate, 

the base asymmetrical to strongly asymmetrical, one 

side obtuse the other attenuate; petiolules pulvinate at 

base or less often nearly cylindrical, 0.5-1.5 cm long, 

1.5-2.5 (-3.5) mm long, appressed-pubescent to nearly 

tomentose, the sepals 1.5-3 mm long, ovate, oblong, 

or less often deltate, concave, obtuse or acute at apex; 

petals oblong, (4) 5.5-7 (-8) mm long, reflexed at anthesis, 

adaxially papillate, abaxially glabrous or sometimes 

appressed-pubescent at base, usually ciliate at margins, 

the apex obtuse, the base cuneate or obtuse; 

appendages as long as the petals, triangular-lanceolate, 

erect, simple or seldom bifid at apex, adaxially sericeous, 

abaxially glabrous or papillate; disc annular, 5- 

lobed, tomentulose, tomentose, puberulent or rarely glabrous, 

0.6-0.7 mm tall; stamens 8, the filaments of equal 

length, 1.7-2.7 mm long, pilose or less often glabrous, 

the anthers 1.4-2.3 mm long, oblong, glabrous, apiculate 

at apex; ovary ovoid, densely sericeous, the stigma 

elongate, papillate. Fruits 1(2)-seeded, ellipsoid to nearly 

globose, glabrescent, yellow, orange or red, 2-3(3.5) 

glabrous, usually drying dark brown, the pulvinus coni- cm long, nearly smooth, rounded at apex, with short 

cal, 4-6 mm long; rachis 10-55 cm long, terete, stri- apiculum, the pericarp 0.8-1.2 (3) mm thick, the enate, 

slightly angled on distal portion, glabrous; peti- docarp smooth, glabrous. Seed ellipsoid, 1.8-2 cm 

oles 9-28 cm long, terete, striate, slightly pulvinate at long; cotyledons superimposed, the upper cotyledons 

base. Thyrses panicle-shaped, terminal or axillary; slightly larger than the lower one. 

cataphylls acicular, 4-5 (10) mm long, early decidu- Talisia cerasina is a highly variable species that can 

ous; axes 20-60 cm long, puberulent, minute be recognized by the presence of falcate leaflets with


Fig. 68. Talisia cerasina. A. flowering branch. B. detail of inflorescence. C. leaflet. D. staminate flower. E. adaxial, 

abaxial, and lateral views of petal with adnate appendage. F. staminate flower with perianth removed showing 

disc and stamens and detail of stamens (left) and l.s. of same showing disc, stamens and pistillode (right), and 

pistillode (far right). G. pistillate flower with perianth removed showing disc, sterile stamens, and pistil with detail 

of sterile stamen (left) and l.s. same (right). H. fruit. (A-B, D-F from Vieira et a!. 579; C, G from Viera et al. 

416; H from Silva 1266).


acute to acuminate apices; leaf rachis glabrous, striate 

and slightly angled distally; long-pedunculate inflorescences 

with relatively large flowers, bearing 8 stamens 

with pilose to puberulent filaments; and disc pilose or 

puberulent. Talisia cerasina and T retusa are vegetatively 

very similar, however, T cerasina can be distinguished 

by its pubescent disc (vs. glabrous in T retusa). 

The specific epithet refers to the cherry-like fruits, as 

originally described by Spruce in the type collection. 

Phenology. Collected in flower from June to March, 

with most collections from September to December; 

fruiting throughout the year. 

Distribution and Ecology. (Fig. 72) From Costa 

Rica to Bolivia, extending into the Amazon and east 

into northeastern and southeastern Brazil. In terra 

firme, igapo, varzea, gallery, and secondary forests, 

and campina. 

Common names and uses. Peru. Loreto: tambor 

caspi, carachupa caspi, virote huayo. Brazil. Acre: breu, 

pitomba; Amazonas: pitomba, pitomba brava, pitombeira; 

Minas Gerais: camboata' da folha grande; Para: 

pitomba; Pemambuco: pitombinha; Rondonia:pitomba 

do mato. Bolivia. Beni: Pit6n. Cultivated for its edible 

fruits (fleshy seed testa). 

Representative specimens examined. COSTA 

RICA. Puntarenas: Golfito, 100-300 m, 27-28 Jan 

1967 (fl), Burger & Matta 4757 (MO, NY, US); Reserva 

Biol6gica Carara, El Salto creek, 130 m, 10 Jan 1990 

(bd), Zuniga 70 (F-2). San Jos6: Santa Rosa de Puriscal, 

17 Jan 1973 (bd), Poveda 446 (MO). 

PANAMA. San Blis: Cordillera San Blas, vic. of Isla 

Tubuala, 200-400 m, 11 Mar 1993 (fr), Herrera 1311 (US). 

COLOMBIA. Antioquia: Cerro de Pava, valley of the 

Cristalina, 1 Sep 1994 (st), Acevedo-Rodriguez & Callejas 

6772 (US); Mun. San Luis. Corregimiento El Prodigio; 

Cano el Tigre, 350 m, 26 Jun 1990 (fl), Cirdenas et al. 

2914 (COL), trail Las Confusas, 300-770 m, 7 Mar 1990 

(fr), Cardenas & Ramirez 2566 (COL, JAUM); Mun. Turbo. 

Corregimiento Alto Mulatos; Finca El Bosque, 350, 19 Dec 

1990 (fr), Callejas et al. 9771 (NY-2); Mun. San Luis. 

Quebrada La Cristalina, 550-770 m, 27 Jan 987 (fl), 

Ramirez & Cardenas 519 (COL, JAUM), 27 Oct 1987 (bd), 

Ramirez & Cdrdenas 1873 (JAUM). Choc6: Riosucio, 

along Truand6 river, 24 Jul 1970 (fr), Arciria 169 (COL). 

Guaviare: San Jose de Guaviare, Guayabero river, 300 m, 

27 Jan 1990 (fl), Marulanda & Mdrquez 1799 (HUA). 

Meta: Sierra de la Macarena, 650 m, 19 Dec 1949 (fl), 

Philipson & Idrobo 1802 (COL, US); Cafio Ciervo, 600 

m, 19 Jan 1950 (fl), Philipson et al. 2142 (COL). 

Putumayo: Puerto Ospina, 23 Mar 1953 (fr), Gutierrez 

2582 (MEDEL). Vaupks: Raudal Alto, margins of Inirida 

river, 180 m, 3 Feb 1953 (fr), Fernarndez 2107 (COL, US); 

Bocas del Ariari, along Guaviare river, 18 Feb 1968 (fl), 

Pinto & Sastre 919 (COL). 

VENEZUELA. Amazonas: San Carlos de Rio Negro, 

20 km from the confluence with Casiquiare, 119 m, 21 

Mar-17 Apr 1981 (fr), Delascio et al. 9572 (VEN), 21 

Mar 1981 (fr), Delascio et al. 9695 (VEN); Atures. Sipapo 

river, 110 km from confluence with Guayapo river, 120 

m, May 1989 (fr), Foldats & Velazco 9249 (PORT, NY). 

ECUADOR. Napo: Orellana. Yasuni Forest Reserve. 

10 km E of Pontificia Universidad del Ecuador Scientific 

Station, along road, 240-310 m, 27 Jun 1995 (st), 

Acevedo R. & Cedeno 7559 (US), 200 m, 30 Jun 1995 

(st), Acevedo R. & Cedeno 7601 (US), 250 m, 27 Jul 1993 

(fr), Aulestia 110 (MO); Aguarico. Reserva Huaorani, 250 

m, 21-25 Oct 1993 (fl), Aulestia & Andi 908 (US); Sector 

Payamino, Via Loreto, 4 km W of Payamino river, 

250 m, 3 Aug 1986 (st), Palacios & Neil 1132 (MO). 

PERU. Amazonas: Prov. Bagua. Marafion river, 

Yamayakat, 320 m, 26 Jan 1996 (fr), Jaramillo & Jaramillo 

981 (US); Cenepa river, Chigkan island, ca. 250 m, 18 

May 1973 (fr), Kayap 761 (MO); Marafion river, 

Yamayakat, 16 Jul 1994 (bd), Vasquez et al. 18734 (US). 

Cuzco: La Convenci6n, Distr. Echarati, 400 m, 15 Feb 

1997 (st), Nunez et al. 19631 (USM). Loreto: Samiria 

river, 20 Oct 1982 (fl), Ayala et al. 3908 (MO, NY); 

Requena. Espejo Cocha, 27 Nov 1987 (fr), Ayala et al. 

5822 (US); Amazonas river 2 km W of Indiana, 130 m, 

12 Feb 1987 (st), Gentry et al. 55713 (MO); Negro Urco, 

Napo river, 160 m, 21 Jan 1983 (fr), Gentry & Emmons 

39627 (F, NY); Amazonas river, Hamburgo-Cafio Pasta 

Cocha, 122 m, 30 Apr 1985 (fr), Grdndez & Vasquez 240 

(K, MO); Samiria river, 11 May 1985 (fr), Grdndez & 

Vasquez 403 (K); Prov. Alto Amazonas. Fortaleza, near 

Yurimaguas, 140 m, Dec 1932 (fl), Klug 2798 (F, MO, 

NY, US); Prov. Mariscal Castilla. Yavari river, 16 miles 

above Curaga river, 25 Oct 1976 (fl), Prance et al. 24113 

(K, MG, MO, NY, US); Prov. Loreto. Amazonas river, 3 

Mar 1977 (fr), Rimachi 2877 (F, MO-2); Prov. Maynas. 

Amazonas river, Yanayacu creek (black waters), 15 Aug 

1977 (fr), Rimachi 3176 (F); Prov. Requena. along Ucayali 

river, 23 Nov 1981 (fr), Spichiger & Encarnaci6n 1201 

(MO); San Jose de Parinari, 5 Nov 1982 (fr), Vasquez & 

Jaramillo 3353 (MO, NY); Esperanza, Tahuayo river, 

140 m, 26 Jan 1981 (fr), Vasquez & Jaramillo 1275 

(F-2, MO-2, NY); Yanamono-Explorama Lodge, 106 

m, 3 May 1987 (fr), Vasquez & Jaramillo 9126 (F, NY); 

Explorama Reserve, 106 m, 9 Nov 1989 (fl), Vasquez 

& Jaramillo 13131 (US), 17 Apr 1985 (fr), V6squez & 

Jaramillo 6358 (MO); Prov. Requena, Sapuena, Jenaro 

Herrera, 170 m, 17 Nov 1987 (fl), Vasquez & Jaramillo 

10107 (K-3); Samiria river, 130 m, 11 May 1985 (fr), 

Vasquez et al. 6492 (MO); Prov. Ucayali. Canchahuayo, 

200 m, 29 Nov 1985 (fl), Vdsquez et al. 7028 (MO, NY); 

Yurimaguas, Huallaga river, 155-210 m, Oct-Nov. 1929 

(fl), Williams 3901 (F); Fortaleza, near Yurimaguas 29 

Oct 1929 (fl), Williams 4303 (F); Masana, Mazan river, 

125 m, 5 Apr 1930 (fr), Williams 8141 (F). 

BRAZIL. Acre: Mun. Bujari. Basin of Purus river, 

Antimari river, Floresta Estadual de Antimari, 12 Mar 

1997 (fr), Daly et al. 9497 (US); Mun. Sena Madureira, 

vic. of Sena Madureira, 27 Sep 1980 (fl), C. Ferreira & 

Nelson 2575 (INPA, MG); Mun. Mancio Lima, road to 

Isac, 24 Mar 1992 (fr), C. Ferreira et al. 10932 (US); Rio


Branco, 26 Sep 1990 (fl), Lowrie et al. 199 (MG, NY); 

Basin of Purus river, along Icaco river, 28 Oct 1993 (fl), 

Silveira et al. 677 (NY); Seringal Sao Francisco, Acre river, 

Oct 1911 (fl), Ule 9517 (K-2, MG, US). Amazonas: Mun. 

Limoeiro; Japura river, 23 Nov 1977 (fl), Damido 2742 

(INPA); Manaus, Ilha do Careiro, cultivated, 15 Jan 1987 

(st), Grenand 2797 (INPA); Mun. Borba; Near 

Urucurituba, 4 Sep 1934 (fr), Krukoff 5963 (K, MO, NY); 

Mun. Humaita; Madeira river near Tres Casas, 11 Oct 1934 

(fl), Krukoff 6504 (K); Mun. Sao Paulo de Olivenqa; near 

Palmares, 11 Sep-26 Oct 1936 (fl), Krukoff 8603 (K, MO, 

NY); Mun. Librea. Purus river, 5 km S of Labrea, 31 Oct 

1968 (fl), Prance et al. 8117 (F, INPA, MG, NY, P, US); 

Janauari, Rio Negro opposite Manaus, 1 Apr 1971 (fr), 

Prance et al. 11245 (F, K, MG, NY, US); Serra de Araca, 

60 m, 20 Mar 1984 (fr), Rodrigues et al. 10557 (INPA, 

NY); Solimoes river, Santo Ant6nio do Ica, 19 Oct 1968 

(fl), Silva 2111 (MG). Bahia: Mun. Una. Reserva 

Biol6gica Mico-leao, 8-12 Mar 1993 (fr), Amorim et al. 

1103 (NY, US); Mun. Uru9uca; km 7.3 on road Serra 

Grande-Itacare, Faz. Lagoa, 7 Sep 1991 (fl), de Carvalho 

et al. 3624 (NY); Reserva Biologica de Una-REBIO, 20 

Jan 1999 (fl), Jardim et al. 1962 (NY); Mun. Una. 

Reserva Biol6gica Mico-leao, 50-75 m, 26 Feb 1978 (fl), 

Mori et al. 9303 (K, NY); Mun. Santa Cruz de Cabralia, 

2-3 km W of Sta. Cruz de Cabralia, 6 Apr 1979 (fr), Mori 

et al. 11694 (NY). Espirito Santo: Reserva Florestal Porto 

Seguro, 1 Jan 1990 (fl), Folli 1043 (NY, US). Minas 

Gerais: Mun. Tombos, Faz. Cachoeira, 29 Jul 1935 (st), 

Mello Barreto 1791 (F-2). Pari: Braganca, 27 Dec 1955 

(fr), Bordallo s.n., herb 22655 (INPA, MG); Santarem, 

mouth of Tapajos river, 11 Dec 1966 (fl), Cavalcante & 

Silva 1674 (MG), 11 Dec 1966 (fr), Cavalcante & Silva 

1675 (MG); Madeira falls, 11 Aug 1902 (fl), Ducke 2875 

(MG); Almeirim, 9 Dec 1902 (fl), Ducke 3037 (INPA, 

MG); 13 Dec 1902 (fl), Ducke 3049 (MG); without spe- 

cific locality, 11 Sep 1907 (fl), Ducke 8703 (MG); Cacaoal 

Imperial, 6 Mar 1909 (fl), Ducke 10205 (MG); 

Cuminamirim-Ariramba, 18 Dec 1910 (fl), Ducke 11462 

(MG); without specific locality, 19 Sep 1927 (fl), Ducke 

20902 (US); Madeira falls, Oct 1886 (fl), Rusby 2527 (F, 

NY-2, US); Santarem, Porto Paricatuba, 6 Dec 1948 (fl), 

Silva 117 (IAN-2, U); Oriximina, along Trombetas river, 

28 Jan 1968 (fr), Silva 1266 (MG); Sao Jose do Gurupi, 

4 Mar 1980 (fr), Teixeira 64 (MG). Pernambuco: Usina 

Mussurepe, along road to Aldeia, 20 Nov 1951 (yfr), 

Ducke & Andrade-Lima 25 (INPA). Rondonia: Madeira 

river, Calama, Apr 1980 (fr), Goulding 174 (MG); 

Guapore river, Mamore-Corte do Indio, 27 Aug 1988 

(fl), Lisboa & Maciel 4452 (MG); Ji-Parand river, 23 Oct 

1979 (fl), Vieira et al. 579 (INPA, MG, MO). 

BOLIVIA. Beni: Prov. Ballivian, 200 m, 18 Oct 1980 

(fl), Beck 5171 (LPB, MO, US); Prov. Vaca Diez, 23 km 

S of Riberalta, vic. of Tumi Chucua, along Tumi Chucua 

lake, 29 Sep 1991 (fl), Beck 20054 (LPB, US, USZ); 

Prov. Yacuma, ancient arm of Yacuma river, 189 m, 6 

Nov 1989 (fl), Moraes & Sarmiento 1111 (LPB, US); 

Espiritu, Sicuri forest, 189 m, 17 Nov 1989 (fr), Moraes 

& Sarmiento 1164 (LPB, US); Prov. Itenez, 60 km NE 

of Trinidad, 30 Oct 1993 (yfr), Moraes et al. 1378 (LPB); 

Prov. Marban, San Joaquin, 2 Nov 1993 (fl, fr), Moraes 

et al. 1475 (US); Prov. Mamore, 25 km N of San 

Ram6n, 7 Nov 1993 (yfr), Moraes et al. 1570 (LPB); 

Pampas, near Lake Rogagua, 11 Mar 1921 (fl), Rusby 

1409 (K, NY); Prov. Ballivian. ca. 2 km S of Curiraba 

river, 250 m, 8 Nov 1985 (fl, yfr), Solomon 14623 (MO- 

2); Trinidad-Misiones, Guarayos, 250 m, Sep 1926 (fl), 

Werdermann 2474 (LPB). Cochabamba: Prov. Carrasco, 

Experimental Station El Chapare, 288 m, 14-19 Oct 

1991 (fl), Killeen et al. 3527 (US). Santa Cruz: Prov. 

Concepci6n, 475 m, 28 Dec 1986 (fr), Nee 33356 (US); 

Prov. 1uflo de Chavez; San Ramon, 505 m, 21 Feb 1991 

(fr), Quevedo & Centuri6n 301 (MO); Prov. Santistevan, 

Estacion Experimental Agricola de Saavedra, 15 km 

NNE of Montero, 265 m, 9 Oct 1990 (fl), Saldias P. & 

Menacho 1231 (MO, NY). 

25. TALISIA CROATII Acevedo-Rodriguez, sp. 

nov. 

Type. Panama. Canal Zone: Barro Colorado Island. 

Wheelertrail, 2 Oct 1970 (fl), Croat 12494 (holotype, MO- 

2 sheets; isotypes, F, MO). Figs. 17c-d, 69a-g 

A T. pinnata (Ruiz & Pav6n) Radlkofer foliolis et 

inflorescentiis pubescentibus vel tomentulosis et petalis 

longioribus differt. 

Treelet 3.5-4.5(-9) m tall. Stems terete, sulcate, 

puberulent, becoming glabrous and lenticellate with age. 

Leaves paripinnate or less often imparipinnate, spirally 

arranged on distal portion of stem; distal process filiform, 

0.3-1.5 cm long, deciduous; leaflets 11-26, alternate 

or subopposite, oblanceolate or less often oblong to 

nearly elliptic, coriaceous, 9.5-22 (41) x 3-6 (12) cm, 

the adaxial surface glabrous, the abaxial surface finely 

pubescent or tomentulose along the prominent veins, the 

venation brochidodromus, tertiary veins reticulate, the 

apex long-acuminate to caudate, the base inequilateral, 

one side obtuse the other acute, the margins entire or 

undulate, slightly revolute; petiolules sub-terete, slightly 

compressed along adaxial surface, 3-5 (13) mm long, 

pubescent to tomentulose; rachis 23-55 (72) cm long, 

terete, striate, puberulent, pubescent to tomentulose, 

sometimes with a few glandular hairs; petioles 15-35 

cm long, terete, striate, puberulent, slightly flattened toward 

the pulvinate base. Thyrses panicle-shaped, 38- 

53 cm long, terminal; cataphylls pinnate, (often with reduced 

acicular leaflets), puberulent to pilose, numerous, 

congested at distal portion of stem or within the regular 

leaves, developed from supra-axillary bud, 8.5-25 cm 

long when terminal, 1-2 cm long when axillary; axes 

sharply angled, ferruginous-puberulent to -tomentulose; 

bracts subulate, puberulent, early deciduous; bracteoles 

subulate, 0.5-1 mm long, persistent, tomentulose; dichasia 

compound; peduncle ca. 2 mm long, flattened, ferruginous-tomentose; 

pedicels 0.5-2 mm long, articulate


Fig. 69. Talisia croatii. A. portion of leaf. B. cataphyll. C. pistillate flower. D. adaxial and abaxial view of petal 

with adnate appendage. E. inflorescence & pistillate flower with perianth removed showing disc, sterile stamens 

and pistil. F. detail of sterile stamen. G. infructescence. H. fruit and diagrams of seed and embryo. (A, C-F from 

Zetek 3570; B from Croat 8820; G-H from Allen 5310).


at the base. Calyx 2.5-4 mm long, abaxially ferrugi- Darien: Cerro Yaviza river, vic. of Yaviza, 1966 (fl), 

nous-tomentose intermixed with glandular papillae or Duke & Bristan 431 (US); Without specific locality, s.d. 

pubescent, adaxially tomentulose, the sepals 1-2 mm (fr), Duke & Bristan 8183 (MO); El Real, 7 Jan 1975 

long, oblong to rounded, concave, dorsally keeled, im- (yfr), Gentry 13454 (MO, US). Panami: Isla Alto de Maje, 

newly formed lake at the Bayano river, 100 m, 19 Apr 

bricate, ciliate; petals white, ca. 5-6.5 mm long, obovate, 

1976 (st), Croat 34375 (MO); Los Santos. Upper Pedregal 

reflexed at anthesis, abaxially sericeous-pubescent along 

river, 350-400 m, 29 Apr 1976 (fr), Croat 34511 (MO, 

dorsal portion and base, adaxially glabrous, rounded at NY); Vic. of Torti Arriba, 200-300 m, 6 Dec 1977 (yfr), 

apex, clawed to attenuate at base; appendage as long as Folsom et al. 6857 (MO); Camp. Gorgas, 11 May 1976 

the petal or slightly longer, oblong to deltate, densely (fr), Garibaldi 106 (MO). 

sericeous on both surface; disc cup-shaped, 5-lobed, to- COLOMBIA. Antioquia: Rio Le6n, 27 Jan 1962 

mentose, 0.6-1 mm tall; stamens 8, the filaments of (st), Cain 59 (COL); Santa FP, 10 km NW of de Antioquia, 

slightly unequal length, pilose or sparsely so, ca. 2.8- 1570 m, 4 Mar 1992 (fr), Gentry et al. 76186 (US). 

3.4 mm long, filiform, the anthers lanceolate, ca. 1.8 mm, Bolivar: San Martin de Loba, Apr-May 1916 (fr), 

apiculate at apex; ovary conical, sericeous, the stigma Curran 203 (US). Choc6: Truand6 river, 18 May 1967 

capitate. Fruit yellow to yellowish orange at maturity, (fr), Duke 11074 (NY, US). Valle: Anserma, Finca La 

ellipsoid to ellipsoid-ovoid, 1.8-2.2 cm long, finely Aurora, 28 Jan 1983 (st), Franco et al. 2036 (COL). 

ECUADOR. Napo: San Jose de Payamico, 40 km 

granulate, sparsely appressed-puberulent, apiculate at 

W of Coca, 300-600 m, 27 Apr 1984 (st), Irvine 1012 

apex, the pericarp coriaceous to woody, 0.5-0.7 mm 

(F). Pichincha: Reserva Maquipucuna, along Umachaca 

thick, the endocarp glabrous. Seed 1, ellipsoid, to 2 cm 

river, 1300-1400 m, 31 Aug 1989 (fl), Webster et al. 

long, with thin, fleshy testa; cotyledons about the same 27287 (QCA). 

size, lying transversely over each other. 

PERU. Huanuco: Pachitea, Puerto Inca, 250-300 

Talisia croatii is morphologically similar to T m, 12 Sep 1982 (fl), Foster 8714 (USM). Madre de Dios: 

pinnata, however, its leaflets, branches, inflorescence Confluence of Tambopata & La Torre rivers, 260 m, 15 

axes and calyx are pubescent to tomentulose, (vs. hir- Jun 1980 (fl), Barbour 5720 (F, USM), 39 km SW of 

sute) and the filaments are glabrous (vs. woolly). Puerto Maldonado, 10 Oct 1983 (fr), Crandlemire-Sacco 

Talisia croatii is also vegetatively similar to T princeps, 82 (MO, US); Tambopata Reserve, 22 May 1986 (fl), 

however, it differs from the latter by the long-pedicelled Funk et al. 8109 (US); Tambopata, Lodge camp site 

flowers (vs. nearly sessile), the tomentulose inflores- 

# 1,200 m, 14 Jun 1989 (fl), Nunez et al. 10903 (MO). 

BRAZIL. Acre: Macauhan river, 4 Sep 1933 (fr), Krukoff 

cence (vs. tomentose), the oblong sepals (vs. ovate), 

5787 (F, K, M, MO, NY, US). Amazonas: Near mouth 

and the ellipsoid fruits (vs. ovoid). The specific epithet 

of Embira river, 15 Jun 1933 (fl), Krukoff 4840 (F, M, 

honors Dr. Thomas B. Croat from the Missouri Bo- MO, NY, US); Mun. Sao Paulo de Oliven9a. Near Palnares, 

tanical Garden, collector of the type and many repre- 11 Sep-26 Oct 1936 (fr), Krukoff 8587 (F, K, MO, NY); 

sentative specimens of this species. 

Mun. Sena Madureira. Seringal Fonte Boa, along Icaco 

Phenology. Flowers from May to November, and river, 28 Oct 1993 (fr), Silveira et al. 675 (NY, US). 

fruiting from January to October. 


Costa Rica, Panama, Colombia, Ecuador, Peru, and Brazil, 

in terra firme, v'arzea, and gallery forests. 

Common names and uses. Panama: Mamon de 

monte, sapatero; Brazil: pitombarana. The fruit has 

been recorded for Peru (Crandlemire-Sacco 82) as a 

food source for the primate Saguinusfuscicolis. 

Representative specimens examined. COSTA 

RICA. Puntarenas: Cant6n Osa, vic. Jalaca farm, Golfo 

Dulce area, 8 Jun 1949 (fr), Allen 5310 (F, US); Palmar 

Norte, 100-200 m, 20 May 1976 (st), Croat 35123 (MO). 

PANAMA. Canal Zone:. Barro Colorado Island, 

Lutz trail, 21 Mar 1969 (st), Croat 8820 (MO); 20 m S 

of Snyder-Molino, 24 Mar 1970 (st), Croat 9034 (MO); 

Shannon trail, 15 Jul 1970 (st), Croat 11270 (MO); Along 

road between locks & Ft. Sherman, 10 Jul 1971 (st), 

Croat 15366 (MO); Barro Colorado Island, Aug 1927 

(st), Kenoyer 646 (US); 13 Nov 1931 (fl), Shattuck 377 

(F, MO), 11 Oct 1935 (fl), Zetek 3570 (F, MO, US). 

26. TALISIA CUPULARIS Radlkofer, Sitzungsber. 


Miinchen 8: 350. 1878. Type. Brazil. Amazonas: 

Prov. Rio Negro, close to Barra [Manaus], Apr 

1851 (yfr), Spruce 1785 (holotype, P; isotypes, K- 

2, P; frag. at M). Fig. 70a-h 

Treelet 3-8 m tall; trunk reaching 8 cm in diam.; bark 

light brown, fissures, with dark brown lenticels, inner 

bark cream. Stems obtusely angled to terete, glabrous, 

lenticellate. Leaves paripinnate; distal process truncate, 

2-3 mm long; leaflets 6-16, usually drying reddish brown 

on abaxial surface, alternate or less often opposite, oblong, 

elliptic or less often lanceolate, (7.5) 10-26 (37) x 

2.8-8.5 (14) cm, coriaceous, glabrous on both surfaces, 

the venation brochidodromus, prominent on abaxial surface, 

midvein prominent on adaxial surface, secondary 

veins plane on adaxial surface, tertiary veins reticulate, 

the margins entire, slightly revolute, the apex acute or


if)~~~~~m 

Fig. 70. Talisia cupularis. A. leaf B. leaflet. C. inflorescence with detail of dichasium. D. staminate flower. 

E. abaxial and adaxial views of petal with adnate appendage. F. staminate flower with perianth removed showing 

disc and stamens with detail of stamen (right) and pistillode (left). G. portion of infructescence. H. lateral view of 

embryo. (A from Mori & Assun,ao 21369; B-F from Prance et al. 3785; G-H from Prance et al. 2624).


acuminate, the base strongly asymmetrical, one side airport, 2 Sep 1979 (fl), C. Ferreira et al. 964 (INPA, K- 

obtuse the other attenuate; petiolules pulvinate, 0.5-1.5 2, NY); Along Uatuma river, 1 1 Mar 1986 (fr), C. Ferreira 

(2.5) cm long, glabrous, the pulvinus 2-7 mm long, et al. 6732 (INPA, K-2, NY); Balbinas, 19 Sep 1986 (fl), 

bulbous; rachis (9)32-47 cm long, terete to obtusely C. Ferreira et al. 8209 (NY); Dtto. Agropecuario, BDFF, 

reserve 1501, 50-125 m, 15 Aug 1990 (fl), Freitas & 

angled distally, glabrous; petioles 7-30 cm long, terete, 

Cardoso 14 (NY); Road to Taruma, 10 May 1953 (fl), 

glabrous, striate, enlarged at base. Thyrses panicle- 

Froes 29610a (IAN, U); 12 Aug 1949(fr), Froes 25002 

shaped, terminal or axillary, to 45 cm long, branches to (IAN); Manaus, 24 Jun 1882 (fl), Glaziou 3757 (M); 

22 cm long; cataphylls acicular, ca. 5 mm long; axes Manaus, 12 Aug 1936 (fr), Krukoff 7962 (F); Sao Paulo 

terete to angled, sulcate, puberulent to tomentose; bracts de Oliven9a; near Palmares, 11 Sep-26 Oct 1936 (fl), 

and bracteoles subulate, tardily deciduous, minutely to- Krukoff 8463 (K, MO, NY, P); Uatuma river, 20 Nov 1983 

mentose; dichasia compound; peduncles 2-4.5 (6) mm (fl), Lima 654 (INPA); Dtto. Agropecuario, BDFF, reserve 

long; pedicels < 0.5 mm long, articulate just below the 1501, 50-125 m, 14 Jul 1990 (fl), Mori et al. 21369 (US); 

flower. Calyxpoculiform, 4-7 mm long, minutely tomen- Manaus-Itacoatiara highway, km 138, 15 Jun 1972 (fr), 

tose, sometimes intermixed with glandulartrichomes, the Pires & Lima 156 (INPA); Mun. Manaus; Reserva Ducke, 

6 Sep 1966 (fr), Prance et al. 2202 (F, INPA, K-2, MG, 

sepals 2-4 mm long, concave, oblong, rounded at apex; 

MO, NY, P, US), 10 Oct 1966 (fr), Prance et al. 2624 (K, 

petals spatulate, cream, 5-8 mm long, reflexed at anthe- 

MG, NY, US); Manaus-Itacoatiara highway, km 204, 21 

sis, glabrous except for the ciliate lower margins and Dec 1966 (st), Prance et al. 3785 (K, MG, NY, US); 

papillate adaxial surface, the apex rounded, the base Aleixo, along road, 30 Aug 1973 (fr), Prance et al. 18769 

clawed; appendages as long as the petals or slightly (K, NY, US); Reserva Ducke, vicinity of dormitories, 3 

shorter, elliptic, erect, densely sericeous on both surfaces; May 1994(fl), Ribeiro, et al. 1299 (US); Manausdisc 

cup-shaped, 5-lobed, minutely sericeous, ca. 1 mm Itacoatiara highway, km 170, 20 Oct 1965 (fl), Rodrigues 

tall; stamens 8, the filaments of equal length, ca. 3 mm 7257 (INPA); Mun. Sao Gabriel da Cachoeira; Iana river, 

long, glabrous, the anthers ca. 1.5 mm long, lanceolate, 5 Nov 1987 (fr), Rodrigues 10863 (INPA, US); Binda 

glabrous, apiculate at apex; ovary nearly conical, sen- creek, 25 May 1961 (fl), Rodrigues & Chagas 2634 

(INPA, US); Reserva Ducke, 1 Aug 1961 (fl), Rodrigues 

ceous-tomentose, the stigma obconical, papillate. Fruits 

& Coelho 2270 (INPA), along Barro Branco creek, 16 

ovoid, glabrous, 2.5-3.2 cm long, yellow, shortly apicu- 

May 1963 (fl), Rodrigues & Coelho 5228 (INPA-unlate 

at apex, the pericarp 2 mm thick, woody. Seeds ob- mounted); Manaus-Caracarai highway (BR-174), km 142, 

long-elliptic, ca. 1.5 cm long, with fleshy testa. Embryo 21 Feb 1974 (fr), Steward et al. P-20411 (K, M, MG, 

with cotyledons superimposed, the upper one slightly MO, P, US). Bahia: Mun. Una. 20 km N of Una along 


road to Ilheus, evergreen, 0-100 m, 23 Jan 1977 (fl), 

The specific epithet refers to the vase-shaped calyx, Harley 18206 (K, NY); Reserva Biologica do Mico-leao, 

which is formed from connate sepals. 

6 Jun 1997 (fr), Jardim et al. 1071 (US). Espirito Santo: 

Santa Teresa, Santa Lucia Biological Station, 650-800 

Phenology. Flowering and fuiting troughout the year. m, 9 Nov 1993 (st), Thomaz 722 (US); 2 Mar 1993 (st), 

Distribution and Ecology. (Fig. 72) Brazil, French 

Guiana, and Peru. In moist, lowland, terra firme, and 

seasonally flooded forests on clayish or sandy soils, 

tall caatinga forest, and campina on white sand. 

Thomaz 1379 (US). Park: Mapuera river, above Carana 

Beira, 6 Dec 1907 (fl, fr), Ducke 9049 (MG); Mun 

Oriximina. Cachorro river, Cabeqa de Onqa creek, 25 Aug 

1986 (fl), C. Ferreira et al 8023 (NY); Trombetas river, 

Monte Branco, 6 Oct 1982 (fr), Revilla et al. 6958 (INPA, 

Common names. Brazil. Amazonas: Pitomba, 

pitomba do mato; Bahia: Combat. 

US). Rio de Janeiro: Quinta de Sao Christovao, 15 May 

1882 (fl), Glaziou 13616 (K, P). 

Representative specimens examined. FRENCH 

GUIANA. Bassin de L'Approuague: Station des 

Nouragues, Jun 1993 (fr), Zhang 18 (US). 

PERU. Loreto: Prov. Maynas, San Miguel, 2 km 

below Indiana, 130 m, 12 Feb 1989 (yfr), Gentry et al. 

65695 (US). 

BRAZIL. Amazonas: Manaus-Itacoatiara road km 26, 

21 Jun 1996 (fl), AssunraCo et al. 315a (US), 20 Feb 1998 

(fl), Assun9do & Pereira 800 (US); Aleixo, INPA grounds, 

28 Apr 1972 (fl), Coelho 156 (INPA-2); Manaus; along 

Bolivia creek, 27 Jul 1956 (fl), Coelho & Coelho 4000 

(INPA); Mun. Itapiranga. Utama river, by St. Luzia creek, 

16 Aug 1979 (fl), C. Ferreira et al. 423 (INPA, K, NY); 

Manaus-Caracarai highway (BR-174), km 97, close to 

27. TALISIA DASYCLADA Radlkofer, Sitzungsber. 


Miinchen 8: 348. 1878. Type. Brazil. Amazonas: 

close to Borba, Aug 1828 (fl), Riedel 1367 (holo- 

type, LE, n.v., photo at US; isotype, LE, n.v., photo 

at US; frag. at M). Figs. 17a, 7 la-i 

Talisia amazonica G. Guarim Neto, Acta Amaz. 9: 

233. 1979. Type. Brazil. Amazonas: Manaus- 

Itacoatiara road, 4 Nov 1970 (fl), Rodrigues 

8989 (holotype, INPA). 

Treelet 2-7 m tall, sometimes with a few upright 

branches on distal portion, theses with determinate

TAXONOMIC TREATMENT 

C 

\AJr 

~~~3cm ~~~~~~~~~ F.~~3m 

3mm 2mm~~~~~mm 

Fig. 71. Talisia dasyclada. A. portion of leaf. B. inflorescence. C. flower bud. D. pistillate flower. E. adaxial view 

of petal. F. pistillate flower with perianth removed showing disc, sterile stamens, and pistil (left) and l.s. same (right). 

G. sterile stamen. H. portion of infructescence. I. seed. (A-G from Rimachi 8965; H-I from Vasquez et al. 11982). 

107


,,Wi ,_, A , , K 

:~ ~~~~ * *s ass 

~~ 

* Talisia cerasina/ 

* Talisia croatil 

* Talisia cupuEn lar}! is\ -> 

* Talisia dasycdada.. vnA 

70W] sow 

47-~~~~~~~~~~7 


growth due to a terminal inflorescence. Stems terete, 5-lobed, pilose or glabrous, ca. 0.8 mm tall; stamens 8, 

smooth, velutinous on young parts. Leaves paripinnate; the filaments of equal length, 1.8-3 mm long, glabrous, 

distal process filiform, to 1 cm long; leaflets 6-10, the the anthers 1.5 mm long, oblong, glabrous, apiculate at 

distal ones opposite, the others sub-opposite to alternate, apex; ovary nearly conical, tomentose, the stigma capielliptic 

to oblanceolate, 15-47 x 6.4-14 cm, coriaceous, tate, tomentulose, papillate. Fruits narrow ellipsoid, glathe 

adaxial surface, glabrous, with impressed midvein brescent, 2-2.2 cm long, yellow, granulose, shortly apiand 

secondary veins, the venation brochidodromus, culate at apex, the pericarp ca. 1 mm thick, the endocarp 

prominent on abaxial surface, forming a marginal loop, granulate, glabrous. Seeds ellipsoid, 1.7 cm long, with 

tertiary veins reticulate, the margins entire, the apex short fleshy cover. Embryo with cotyledons superimposed, the 

to long-acuminate, the base obtuse to rounded; petiolules upper one twice as large as the lower one. 

thickened, 1-1.5 cm long, woolly-pubescent; rachis to The specific epithet refers to the shaggy stems of 

55 cm long, terete, canaliculate, velutinous; petioles 24- this species. 

30 cm long, terete, canaliculate, thickened at base, with 

same indumentumum as the rachis. Thyrses panicleshaped, 

terminal or cauliflorous, pyramidal, to 30 cm 

Phenology. Flowering in August, October, and 

January, and fruiting in February and April. 

long; cataphylls acicular, ca. 1 cm long; axesangled, sul- Distribution and Ecology. (Fig. 72) Peruvian and 

cate, minutely velutinous to tomentulose; bracts ovate, Brazilian Amazon, in moist, lowland, non-flooded 

tomentose with a few glandular hairs, persistent, 1.5-2 forests. 

mm long; dichasia compound, basal ones pedunculate, 

to 4 mm long; pedicels 1.5-4 mm long, articulate at upper 

third to near the apex. Calyx 2.5-3 mm long, tomentose, 

with a few glandular hairs, the sepals 1.8 mm long, 

ovate, rounded at apex; petals obovate, 5.5-7 mm long, 

reflexed, sparsely papillate on adaxial surface, glabrous 

on abaxial surface, ciliate at margins, the apex rounded, 

the base narrowed; appendages as long as the petals, ob- 

Common name. Brazil. Amazonas: Pitomba. 

Representative specimens examined. PERU. 

Loreto: Amazonas river, along Tahuayo creek, 120 m, 

26 Oct 1988 (fl), Rimachi 8965 (US); Alpahuayo, 20 Oct 

1984 (fl), Vasquez & Criollo 5811 (MO, NY); Iquitos- 

Nauta road, km 32, 150 m, 20 Aug 1986 (fl), Vasquez 

& Jaramillo 7876 (MO), km 44, 6 Apr 1989 (fr), Vdsquez 

et al. 11982 (USM); Saboya, Pintuyacu river, 150 m, 

long, erect, villous on both surfaces; disc cup-shaped, 19 Apr 1986 (fr), Vdsquez et al. 7430 (MO, NY). 

5ow1 

40W


BRAZIL. Amazonas: Mun. Coari; Lago do Coari, 

23 Feb 1972 (fr), Coelho 500 (INPA); 

Manaus-Itacoatiara highway, km 64, 28 Mar 1967 (fr), 

Prance et al. 4712 (INPA). Para: Alto Tapaj6s, Vila Nova, 

vic. of Chacaras falls, 19 Jan 1952 (fr), M Pires 3968 (US). 

28. TALISIA DOURADENSIS Acevedo-Rodriguez, 

sp. nov. 

Type. Brazil. Para': Rio Jari, Monte Dourado, terra 

firme forest, 22 Nov 1968 (fl), Silva, TN. 1437 (holotype 

NY; isotypes, F, K). Fig. 73a-d 

A T. guianensi Aublet foliolis paucis, petalis 

minoribus, sericeis et filamentis pilosis differt. 

ily distinguished by the elongated petiolules and abaxially 

sericeous petals. The specific epithet refers to Monte 

Dourado, the locality where the species was collected. 

Phenology. Collected in flower in October. 


from terra firme forest in Monte Dourado, Brazil. 


Park: Monte Dourado, 23 Oct 1968 (bd), Silva 1294 

(F, IAN, K, NY). 

29. TALISIA EQUATORIENSIS Acevedo- 


Tree 10 m tall, stems 8-10 cm in diam. Distal por- Type. Ecuador. Pichincha: Between Atenas and 

tion of stems terete, blackish brown, puberulent, smooth. 

Leaves paripinnate; distal process minute, conical; leaflets 

6-8, opposite or alternate, elliptic, coriaceous, 

Sapullo, Km 17, 1750 m, 2 Nov 1991 (fl), J. 

Jaramillo & E. Grijalva 14473 (holotype, NY; isotype, 

QCA, n.v.). Fig. 74a-f 

13.5-20 x 4.5-6.5 cm, glabrous, glossy on adaxial 

surface, midvein yellowish, prominent on abaxial sur- 

A T. eximia Kramer foliolis abrupte acuminatis, petalis 

tomentulosis et fructibus ovoideis vel globosis differt. 

face, tertiary veins reticulate, the apex acute or acuminate, 

the base inequilateral, one side obtuse the other 

acute, narrowing into the petiolule; petiolules 10-20 

mm long, with a conical, reddish brown, pulvinate base; 

rachis 20 cm long or longer, terete, puberulent, dark 

reddish brown, dull; petioles 9-17 cm long, pulvinate 

at base. Thyrses panicle-shaped, 17-33 cm long, axillary; 

cataphylls clustered, axillary, acicular, ca. 2 mm 

long, puberulent; axes terete, striate, minutely pubescent; 

bracts persistent, deltate, 0.6-1 mm long, minutely pubescent; 

dichasia compound; peduncles ca. 2-2.5 mm 

long, flattened, minutely pubescent; pedicels 1.7-2.2 

Tree 5-25 m tall. Stems terete, glabrous, lenticellate. 

Leaves paripinnate, 1.5-3 m long; distal process acicular, 

ca. 1 cm long; leaflets 6-12, opposite or altemate, 

oblanceolate or less often elliptic, 40-55 x 6.5-20.5 

cm, sub-coriaceous, glabrous, flattened to slightly bullate, 

dull on both surfaces, the venation brochidodromus, 

not forming a sub-marginal loop, the adaxial surface 

with a prominent midvein and slightly raised secondaries, 

the abaxial surface with strongly prominent midvein, 

secondary and tertiary veins prominulous, tertiary veins 

reticulate, the margins undulate, the apex abruptly acuminate, 

the base slightly asymmetrical, obtuse; petimm 

long, articulate at the middle, minutely pubescent. olules woody, enlarged, 0.6-1.6 x 0.5-0.9 cm, glabrous, 

Calyx green, 2.5-3 mm long, minutely pubescent, the adaxially canaliculate; rachis terete, striate, glabrous; 

sepals 1.2-1.6 mm long, deltate, concave; petals reflexed petioles at least 21 cm long, nearly terete, striate, glaat 

anthesis, oblong-elliptic, 3.5-4 mm long, white, sen- brous, bulbous at base. Thyrses panicle-like, terminal; 

ceous on abaxial surface, papillate on adaxial surface, cataphylls minute, scale-like; dichasia simple or comthe 

base cuneate; appendages deltate, as long as the pound; pedicels ca. 2.5 mm long, articulate at the middle. 

petals, adaxially sericeous-tomentose, abaxially seri- Calyx 2.5-3 mm long, puberulent, the sepals 1-1.7 mm 

ceous on upper half; disc 5-lobed, sericeous; stamens long, ovate, concave, obtuse at apex; petals elliptic, 5.5-6 

6-8, the filaments pilose, slightly unequal, 2-2.3 mm mm long, white, adaxially glabrous, abaxially tomentulose 

long, the anthers elliptic, 1.3-1.7 mm long, apiculate at at base, ciliate at margins, the apex rounded-obtuse, the 

apex. Pistillate flowers, fruits and seeds unknown. base cuneate; appendages oblong-lanceolate, as long 

Talisia douradensis resembles T guianensis in gen- as the petal, villose sericeous-tomentose on both sureral 

aspect. However, T douradensis differs by its leaves faces, adaxially glabrous at base; disc cup-shaped, 5- 

with fewer leaflets (6-8 vs. 16-22) and puberulent lobed, puberulent, ca. 1 mm tall; stamens 8 (9), the filarachis 

(vs. glabrous); its deltate, minutely pubescent (vs. ments of unequal length, 3-3.5 mm long, slightly 

subulate, appressed-pubescent) bracts and bracteoles; flattened, woolly-pubescent, the anthers 1-1.5 mm long, 

minutely pubescent calyx with deltate sepals (vs. puberu- elliptic, glabrous, apiculate at apex. Pistillate flower 

lent with ovate sepals); abaxially sericeous, 3.5-4 mm unknown. Fruits brown, ovoid to globose, 3-3.2 cm 

long petals (vs. abaxially glabrous, 6-7.5 mm long); long, glabrous, rugulate, the pericarp woody, ca. 4 mm 

sericeous disc (vs. glabrous or hispidulose); and stamens thick, the endocarp glabrous, smooth. Seeds solitary, 

with pilose (vs. glabrous) filaments. This species is eas- nearly globose, the testa apparently not fleshy. Embryo


t l'Y 

3. 

.J?~~~ 3mm. 

2mm / ~~~~~~~~~~ 

Fig. 73. Talisia douradensis. A. inflorescence with subtending leaf. B. staminate flower. C. lateral, adaxial and 

abaxial views of petal with adnate appendage. D. staminate flower with perianth removed showing disc, stamens, 

and pistillode (left), I.s. same (right). (All from Silva 1437). 

4


3. j~~~~~~~~~~~~~~~~~~~~~~~~~~m 

Fig. 74. Talisia equatoriensis. A. portion of leaf. B. staminate flower. C. lateral, adaxial, and abaxial views of 

flower. D. detail of stamen, staminate flower with perianth removed showing disc and stamens, and I.s. same, 

showing pistillode. E. branch with infructescence and x.s. of fruit. F. lateral view of embryo. (A, E-F from Mori 

& Kallunki 5199; B-D from Dodson et al. 8420).


sub-globose, the cotyledons laying obliquely transverse 0.3-1.2 cm, glabrous, adaxially sub-canaliculate to 

over each other, of similar size. 

strongly canaliculate; rachis at least 23 cm long, terete to 

Talisia equatoriensis is morphologically similar to transversely depressed-obovate in x.s., striate, minutely 

T eximia, however, the former differs from the latter by: lenticellate, glabrous; petioles to at least 17 cm long, 

abruptly acuminate leaflets (vs. obtuse to acuminate); nearly terete, striate, glabrous, bulbous at base. Thyrses 

abaxially tomentulose petals (vs. glabrous); and the axillary or distal, panicle-like, ca. 50 cm long; cataphylls 

ovoid or globose fruits (vs. ellipsoid). The specific epi- acicular, clustered supra-axillary, 4-6 mm long; 

thet refers to the country where the type was collected. axesangled, puberulent, with minute papillate glands; 


Panama, Colombia and Ecuador, in wet or humid, lowland 

forests. 

Representative specimens examined. PANAMA. 

Col6n: Guanche river, ca. 3 km upriver from bridge on 

road to Portobelo, 25 m, 24 Mar 1975 (fr), Mori & 

Kallunki 5199 (MO). 

COLOMBIA. Without specific locality, Vereda 

bracts triangular to nearly lanceolate, 0.5-0.7 mm long, 

puberulent; dichasia simple or compound; peduncles ca. 

0.5 mm long, puberulent; pedicels 2.5-4 mm long, articulate 

at upper third, puberulent to pubescent. Calyx 

2-2.5 mm long, puberulent, the sepals ovate, ca. 1.5 mm 

long, keeled along dorsal ridge, ciliate at margins; petals 

ovate-lanceolate, 5.5-6 mm long, reflexed at anthesis, 

white, papillate adaxially, glabrous abaxially, ciliate at 

Bohios, toward La Patricia. 30 m, 5 Aug 1985 (fr), margins, the apex obtuse, the base cuneate; appendages 

Renteria & Cdrdenas 4339 (MEDEL). 

lanceolate, slightly shorter than the petal, villose adaxially 

ECUADOR. Carchi: Cant6n Maldonado. Parroquia on upper half and along margins, glabrous or sparsely 

Tobar Donoso, Reserva Awa, 900 m, 22 Nov 1992 (fl), appressed-pubescent abaxially, the apex obtuse, the base 

Aulestia et al. 641 (US). Guayas: Chong6n-Coloche, 

narrowed; disc annular, 5-lobed, orange, tomentulose, 

600 m, 26 Aug 1997 (st), Cornejo & Bonifaz 5720 (US). 

ca. 1 mm tall; stamens 8, the filaments of equal length, 

Manabi: Pedernales. Cerro Pata de Piajaro, 300-700 m, 

19-21 Jun 1996 (fr), Clarke et al. 2650 (MO, NY, US). 

pilose, the anthers ca. 1 mm long, lanceolate, glabrous, 

Pichincha: El Centinela, crest of Montafias de Ila on road apiculate at apex; ovary conical, ferruginous-tomentulose, 

to Patricia Pilar to 24 de Mayo at km 12, 600 m, 15 July the stigma 3-lobed-capitate, papillate. Fruits yellow, el- 

1979 (fl), Dodson 8420 (MO), 23 May 1983 (fr), Dodson lipsoid, ca. 2.8 cm long, glabrous, minutely rugulate, the 

& Benzing 13911 (MO), 30 Jul 1984 (st), Dodson et al. pericarp woody, ca. 2 mm thick, the endocarp glabrous, 

14634 (MO). 

rugulate. Seeds unknown. 

While the specific epithet means magnificent or exceptional, 

it is not clear how it would apply to this species. 

30. TALISIA EXIMIA Kramer, Act. Bot. Neerl. 21: 

677. 1972. Type. Surinam. Brokopondo District: E 

of Brokopondo, high forest, 24 Jan 1966 (fl), J. 

van Donselaar 3047 (holotype,U-2 sheets) 

Fig. 75a-e 

Talisia morilloi Steyermark, Acta Bot. Venez. 10: 

237. 1975. Type. Venezuela. Distrito Federal, 

Tacamahaco, humid forest, 900-1000 m, 16 Jul 

1973 (fl), G. Morillo et al. 3339 (holotype, VEN, 

n.v.; isotypes, MO, US). 

Treelet to 8 m tall; trunk to 6 cm in diam.; bark light 

brown, lenticellate. Stems terete, glabrous, vemicose. 

Leaves paripinnate; distal process acicular, ca. 4 mm long; 

leaflets 4-8 (or more), opposite or altemate, oblanceolate, 

spatulate or elliptic, 18.5-66 x 6.5-20 cm, sub-coriaceous, 

glabrous, flattened to bullate, slightly lustrous on 

both surfaces, the venation brochidodromus, forming a 

sub-marginal loop, tertiary veins reticulate, the adaxial 

surface with a prominent carinate midvein and slightly 

raised secondaries, the abaxial surface with strongly 

prominent midvein, secondaries and tertiary prominulous, 

the margins undulate, slightly revolute, the 

apex obtuse to acuminate, the base slightly asymmetrical, 

cuneate to obtuse; petiolules woody, bulbous, 0.5-1.5 x 

Phenology. Collected in flower in Febnruy and July. 


Venezuela and Surinam, in lowland, tall forest. 

Common names. Surinam: Gaan tatoe, tatoe, 

wanhede. 

Representative specimens examined. VENEZUELA. 

Distrito Federal: Tamahaco, Quebrada Rio Grande, 900- 

100 m, 16 Jul 1973 (fl), Morillo et al. 3339 (MO, US). 

SURINAM. Brokopondo: Brownsberg, along trail 

to Irene Falls, ca. 6 km from road, 18 Feb 1977 (fl), 

Lindeman et al. 72 (K, MO, NY). Saramacca: Voltzberg 

Nature Reserve, Coppename river, 24 Feb 1977 (fl), 

Lindeman et al. 137 (MG, MO, VEN). 

31. TALISIA GHILLEANA Acevedo-Rodriguez, sp. 

nov. 

Type. Brazil. Amazonas: Rio Cuieras, just below 

mouth of Rio Brancinho, savanna forest on sand, 25 

Sep 1971 (fl), G. T Prance, D.F Coelho & O.P Monteiro 

14816 (holotype, MO; isotypes, INPA, n.v., K, MG, 

NY, U, US, S, n.v.). Figs. 17b, 77a-g


- -...--- 1\\YMFW ~~~~~~~2mm. 

/~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~ 

3 cm.~ 

/ .~ ~ ~ ~ ~ ~ ~ ~~~3m 

Fig. 75. Talisia eximia. A. portion of leaf showing leaflets and petiole. B. inflorescence. C. pistillate flower. D. 

abaxial and adaxial views of petal. E. pistillate flower with perianth removed showing disc, sterile stamens and 

pistil and l.s. same. (A from Lindenman et al. 72); B-E from Lindenman 137).


I 

|OW, ,. .\;A: ... 

70W 6saw 5OWi 40W 

*Talii douradenis 

7 ~~~~~~~~~~~ 


eximia'i ,, ,,,, - ;os 

* Talsiadoradeis.76 Fig. 76.-Disributin Disr.ibuin of secies i Talisa subgnus Talsia (i part) 

fspce in al/ i sugeu Taii/inpr) 

A T. cerasina (Bentham) Radlkofer foliolis paucis, 

petalis pubescentibus differt. 

I ON 

margins ciliate, the base cuneate, the apex rounded; 

appendages deltate, as long as the petals, adaxially sericeous-tomentose 

above the cuneate base, abaxially glabrous; 

disc cup-shaped, ca. 1 mm tall, puberulent to 

nearly tomentose; stamens 8, the filaments glabrous, 

Treelet, 2-7 m tall. Stems terete, puberulent, minutely 

lenticellate, becoming glabrous with age. Leaves 

paripinnate or less often imparipinnate; distal process 

deciduous, leaving a truncate base 2-3 mm long; leaf- slightly unequal, 1.5-3 mm long, the anthers lanceolate, 

lets (2) 4-6 (7), opposite or altemate, lanceolate, ovate, ca. 1.1 mm long, apiculate at apex; ovary conical, serioblong-elliptic, 

or elliptic, coriaceous, 6.5-24 x 2.2- ceous, the style elongated, the stigmatic surface elon- 

7(13.2) cm, glabrous, the adaxial surface slightly glossy, gate capitate, papillate. Fruit orange-yellow to red at 

with prominent to plane primary and secondary veins, maturity, ellipsoid, apiculate, ca. 2 cm long, glabrous at 

the abaxial surface dull, with very prominent, yellow- maturity, the pericarp coriaceous, granulose, ca. 0.6 mm 

ish midvein and prominent secondary and tertiary veins thick, the endocarp glabrous. Seeds 1 per fruit, nearly 

forming a reticulum, the margins revolute, the apex long- ovoid, with fleshy testa. Embryo with cotyledons layacuminate 

or less often caudate, the base obtuse, some- ing obliquely transverse to each other, the upper one 

times inequilateral, decurrent on to the petiolule; peti- slightly smaller than the lower one. 

olules flattened along adaxial surface, 0.5-2 cm long, Talisia ghilleana seems to be closely related to T 

pulvinate at base, glabrous; rachis 2-14 cm long, nearly cerasina in general habit. However, T ghilleana difterete 

or obtusely angled, puberulent, dark reddish 

fers from T cerasina by: leaves with 4-6 leaflets (vs. 

brown or dark grey. Thyrses panicle-shaped, terminal 

10-16); peduncle 0.1-1.5 mm long (vs. 0.5-5 mm 

or supra-axillary; 12-35 cm long; cataphylls clustered 

long); and petals elliptic to obovate, abaxially appressedat 

base of inflorescence, acicular 2-3 mm long, puberupubescent 

at base, and 3.5 mm long, (vs. oblong, 

lent; axes obtusely angled, sulcate, puberulent; bracts 

abaxially glabrous, and 5-5.5 mm long). Talisia 

and bracteoles persistent, subulate, concave, 1-1.2 mm 

ghilleana honors Dr. Ghillean T. Prance, collector of 

long, puberulent; dichasia compound; peduncles 0.1the 

type specimen and eminent Amazonian botanist and 

1.5 mm long, puberulent; pedicels 1.5-2 mm long, arexplorer 

who has advanced the botanical knowledge 

ticulate at upper portion, puberulent. Calyx green, 2.5of 

this vast region. The name T prancei is already oc- 

3.2 mm long, puberulent to pubescent (white hairs), 

cupied by a synonym of Matayba guianensis Aublet 

the sepals ca. 2 mm long, broadly ovate, concave, cili- 

(T prancei G. Guarim Neto). 

ate at margins; petals elliptic to obovate, ca. 3.5 mm 

long, greenish white, appressed-pubescent on lower half Phenology. Flowers from April to October and 

on abaxial surface, papillate on adaxial surface, the fruits from October to May.


c. G 

3cm.[~~~~~~~~~3m 

1?~~~~~ E 

Fig. 77. Talisia ghilleana. A. flowering branch. B. staminate flower. C. abaxial, adaxial, and lateral views of petal 

with adnate appendage. D. staminate flower with perianth removed showing disc and stamens (right) and l.s. same with 

detail of anther (left). E. flower with immature fruit. F. flower with perianth removed showing disc, sterile stamen 

and immature fruit. G. portion of infructescence. (A-F from C. Ferreira et al. 953; G from Plowman et al. 12664).


Distribution and Ecology. (Fig. 80) Known only alternate, oblong, oblong-elliptic, or elliptic, nearly cofrom 

the State ofAmazonas, Brazil in terra firme forest riaceous, 8-18(44) x 2.1-6(13.5) cm, glabrous and 

on white sandy or clayish soils, campina forest on white glossy on both surfaces, midvein yellowish, prominent 

sand, and non-flooded caatinga. 

on abaxial surface, tertiary veins reticulate, the apex acu- 

Common names. Brazil: Pitomba,pitomba brava, 

pitomba da mata. 

minate, the base inequilateral, one side obtuse the other 

acute, abruptly narrowing into the petiolule; petiolule 

3-9 mm long, with a conical, reddish brown, pulvinate 

Representative specimens examined. BRAZIL. base; rachis 10-58 cm long, terete, or slightly striate on 

Amazonas: Manaus-Caracarai hwy., km 61, Biological distal area, glabrous, dark reddish brown, glossy; pet- 

Reserve Campinas, 2 Oct 1974 (fl), deAlbuquerque 1098 ioles 9-38 cm long, pulvinate at base. Thyrses panicle- 

(INPA), 9 Oct 1974 (fr), deAlbuquerque 1110 (INPA); shaped, 10-25 cm long, terminal or axillary on distal 

Mun. Manaus. CachoeiraAlta, 2 Sep 1955 (fl), Chagas portion of branches; cataphylls clustered, supra-axils.n., 

herb. 1798 (INPA); along Passarinho creek, 17 Jun lary, acicular or dendroid, 4-17 mm long, appressed- 

1955 (fl), CoeAlho 1205 (INPA-2); along road to Mindfi, pubescent; axes reddish tinged, obtusely angled, sulcate, 

19 Aug 1955 (fl), Coelho 1667 (INPA-2); Manaus- glabrous to puberulent; bracts persistent, subulate, 3-5 

Caracarai hwy., km 97, 1 Sep 1979 (fl), C. Ferreira et mm long, appressed-pubescent; dichasia compound; 

al. 953 (INPA, MG, NY); Manaus-Itacoatiara road, km peduncles 3-5 mm long, puberulent; pedicels 1-1.5 mm 

19, secondary, 15 Sep 1955 (fl), Mello, s.n., herb 1918 long, articulate at the upper third, puberulent, reddish. 

(INPA-unmounted); Road to Paredao, 24 Apr 1956 (fl), Calyx reddish tinged, 2-3 mm long, puberulent, the 

Mello & Coelho s.n., herb 3778 (INPA); Manaus- sepals 1.5-2 mm long, ovate, concave, ciliate at mar- 

Caracarai hwy. km 60, INPA reserve, 24 Nov 1982 (fr), gins; petals reflexed at anthesis, oblong-elliptic, 6-7.5 

Mesquita et al. 825 (MG); Manaus-Caracarai hwy., km mm long, white, glabrous on abaxial surface, papillate 

45, Biological Reserve Campinas, 26 Dec 1982 (fr), on adaxial surface, the base truncate-cuneate, the mar- 

Plowman et al. 12664 (F, MG, NY); Manaus-Caracarai gins ciliate; appendages deltate-lanceolate, as long as 

hwy., km 2, 14 Nov 1966 (fr), Prance et al. 3124 (F, the petals, adaxially sericeous-tomentose, abaxially 

MG, NY, US); Manaus-Caracarai hwy., km 46, 16 Sep glabrous and papillate, adnate at the base of petal; disc 

1977 (fl), Ribamar & Ramos 274 (INPA); Experimental annular, 5-lobed, glabrous or hispidulose; stamens 

Reserve, 22 Sep 1977 (fl), Ribamar & Ramos 363 (5)6-8, the filaments glabrous, equal or slightly unequal, 

(INPA); Reserva Ducke, 26 Aug 1957 (st), Rodrigues 2.5-4.5 mm long, the anthers oblong, 1.3-1.5 mm long, 

546 (INPA); along creek of Parque 10, 3 Aug 1959 apiculate at apex; ovary conical-trilobed, glabrous, the 

(fl), C. Rodrigues 1243 (INPA), Buiao creek, 5 May style elongated, appressed-pubescent, the stigma capi- 

1962 (fr), Rodrigues & Chagas 4682 (INPA-2); tate, papillate. Fruit orange-yellow at maturity, ellipsoid- 

Manaus-Caracarai hwy., km 8, 8 Sep 1960 (st), ovoid, apiculate, 2-2.5 cm long, glabrous, the pericarp 

Rodrigues & Coelho 1738 (INPA); Leao creek, 17 Oct coriaceous to woody, granulose, ca. 1 mm thick. Seeds 

1961 (fr), Rodrigues & Lima 2665 (INPA). 1-3 per fruit, nearly ellipsoid, with yellowish, fleshy 

testa. Embryo with cotyledons diagonally superimposed, 

the upper one larger than the lower one. 

32. TALISIA GUIANENSIS Aublet, Hist. P1. Guiane 

1: 349. 1775. Type. French Guiana. Without specific 

locality, at riverbank, without date (fl), Aublet 

s.n. (holotype, BM). Figs. 16a-b, 78a-f 

This species is easily recognized by the attenuate 

leaflet base decurrent onto the elongated petiolule. The 

specific epithet refers to the locality where the type was 

collected. 

Talisia rosea Vahl, Eclog. 2: 30. 1798, nom. illegit., Phenology. Flowers from September to December, 

a superfluous renaming of TJ guianensis Aublet and fruits from November to May. 

Talisia glabra DC., Prodr. 1: 609. 1824. Type. French 


Guiana. Cayenne, without date (fl), Anonymous, 

s.n. [IDC microfiche 2842] (holotype, G-DC, n.v.; Guyana, French Guiana, and Amapa, Brazil, in moist, 

microfiche; frag. at M). 

terra firme forest. Expected from Surinam. 

Treelet, 2.5-10 m tall, unbranched or with few ascending, 

sympodial branches on distal portion; bark dark 

brown, lenticellate. Distal portion of stem nearly terete 

or sulcate, glabrous or puberulent, smooth. Leaves 

paripinnate, spirally arranged on distal portion of 

stem(s); distal process 5-7 mm long, acicular, deciduous 

leaving a truncate apex; leaflets 16-22, opposite or 


East Berbice/Corentyne: Mapenna creek, 20 May 1950 

(fr), Fanshawe 2955 (NY, US); Courantyne river, 

Mapenna creek, 20 May 1950 (st), Fanshawe 6285 (K, 

US); Canje river, 250 m, 9 Apr 1987 (fr), Pipoly et al. 

11348 (US). Upper Demerara/ Berbice: Mabura hill, 

180 km SSE of Georgetown, 0-50 m, 14 Dec 1988 (st), 

Steege & de Jager 562 (U).


3 ml (n , / X~~~~~~~~~~3mm 

S A 

Fig. 78. Talisia guianensis. A. portion of leaf. B. pistillate flower. C. adaxial and lateral views of petal with 

adnate appendage. D. pistillate flower with perianth removed showing disc, sterile stamens, and pistile with detail 

of anther (right) and I.s. same, showing ovules (left). E. fruiting branch. F. seed. (A from Mori et al. 14926; B-D 

from Mori & Boom 15164; E-F from Acevedo R. et al. 4871).


FRENCH GUIANA. Without specific locality, 1838 acute; petiolules pulvinate, 3-5 mm long, minutely hir- 

(fl), Le Prieur s.n. (P-2). Bassin de la Comte: Comte river, sute with scattered glandular hairs; rachis (5.5) 13-40 

20-24 km S of bridge, along road to Brazil, 15 Jan 1977 cm long, terete, or slightly angled distally, slightly stri- 

(fl), Mori & de Granville 8914 (CAY, MO, US-2). Bassin 

ate, minutely hirsute, sometimes with scattered setade 

la Mana: Montagnes de la Trinite; Inselberg NW, 500 

m, 17 Jan 1984 (fr), de Granville et al. 6093 (CAY, U). 

ceous or glandular hairs; petioles (5.5) 8-19 cm long, 

Bassin du Maroni: Camp Maipouri, road to Apatou, km enlarged at the base. Thyrses panicle-shaped, 6-31 cm 

40, 25 Apr 1984 (fr), Sabatier 859 (CAY). Bassin du long, terminal or axillary on distal portion of branches; 

Sinnamary: Research station between Silvolab and cataphylls 5-25 mm long, acicular, sericeous-tomentose; 

Paracou, 16 Oct 1995 (fl), Molino 1469 (CAY); Paracou axes pubescent to sericeous-tomentose, with scattered 

Forest; experimental site C.T.F.T., 5 Feb 1988 (fr), Riera to numerous glandular hairs; bracts early deciduous, 

1459 (CAY-2, US). Montagne de Kaw: Montagne de subulate, ca. 5 mm long, with same indumentum as the 

Kaw, 14 Dec 1954 (fl), Cowan 38809 (NY-2). Piste de axes; bracteoles persistent, similar to the bracts but 

Saint Elie: Arbocel, near St. Elie, May 1980 (st), de smaller; dichasia compound, sessile; pedicels 1-1.2 mm 

Foresta 146 (CAY-2); Station Biologique ORSTOM, 13 

long, appressed-pubescent, articulate at the base. Ca- 

Apr 1983 (st), de Foresta 341 (CAY-2), 19 Nov 1983 

(fl), de Foresta 419 (CAY-3); Piste de St. Elie, 9 Sep 

lyx nearly conical 2.5-3.5 mm long, tomentose, with 

1981 (st), Halle 2756 (CAY); Valley between Sinnamary- scattered glandular hairs, the sepals 1-2.2 mm long, 

Counamama rivers, 14 May 1991 (fr), Sabatier 3572 oblong or ovate-deltate, obtuse at apex; petals whitish, 

(CAY, US). Region Littorale: Kourou. Passoura creek, reflexed at anthesis, 5.5-6 mm long, nearly elliptic to 

6 May 1992 (fr), Acevedo R. et al. 4916 (CAY, US), 6 obovate, the abaxial surface glabrous, the adaxial sur- 

May 1992 (fr), Acevedo R. et al. 4922 (CAY, US); Roche face with scattered glandular, papilliform hairs, ciliate 

Degonde, 5 Feb 1995 (st), Cadamuro & Solacroup 173 

(CAY). Region de Saul: Monts La Fumee, 200-400 m, 

5 Nov 1982 (fl), Mori & Boom 15164 (CAY-2, MG, NY), 

220 m, 14 Sep 1982 (bd), Mori et al. 14926 (NY). 

BRAZIL. Amapi: Oiapoque river, 2 Feb 1950 (fr), 

Froes 25807 (IAN). 

at margins, the apex rounded, the base attenuate; appendages 

as long as the petals, deltate, the abaxial surface 

glabrous with a few glandular hairs, the adaxial 

surface hirsute on the upper half; disc annular, 5-lobed, 

ca. 1 mm tall, minutely hirsute; stamens 8, the filaments 

glabrous, of same length, ca. 4 mm long, the anthers 

oblong, ca. 1.2 mm long, apiculate at apex; ovary coni- 

33. TALISIA HEMIDASYA Radlkofer, Sitzungsber. 


8: 349. 1878. Type. Surinam. Without specific locality 

or date (fl), Hostmann 1274 (holotype, P; 

isotypes, C, n.v., K-2, MO, NY, P; frag. at M). 

Figs. 2a-b&d, 4c, 79a-h 

Talisia glandulifera Steyermark, Ann. Missouri Bot. 

Gard. 75: 1072. .1988. Type. French Guiana. 

Saul, Mont La Fumee, 200-400 m, 3?37'N, 

cal, sericeous-tomentose, the stigma cylindric-trigonous, 

papillate, ca. 1 mm long. Fruit yellow to orange-yellow 

at maturity, globose-ovoid or obovoid, apiculate, ca. 2 

cm long, densely hirsutulous, the pericarp woody, ca. 3 

mm thick. Seed one per fruit, nearly globose, ca. 1.5 cm 

long, with a thin, fleshy, orange-yellow testa. Embryo 

with cotyledons superimposed, of equal size. 

The specific epithet refers to the "partly shaggy" 

pubescence of stems and inflorescence axes. 

53012'W, 1 Oct 1982 (fl), S.A. Mori et al. 15027 

(holotype, MO; isotypes, CAY, MG-2, NY). 

Phenology. Flowers from August to October and 

in May, and fruits throughout the year. 

Tree 12-18 m tall, branching on upper portion to 

produce a dense crown; trunk to 23 cm in diam.; bark 

grayish, thin, smooth, peeling off in irregular plates; 

Distribution and Ecology. (Fig. 80) Venezuela, the 

Guianas, and Brazil, in non-flooded moist, tall forest. 

inner bark reddish. Young growth conspicuously fulvotomentose, 

intermixed with numerous glandular hairs. 

Twigs nearly terete, glabrescent with age. Leaves 

paripinnate, spirally arranged on distal portions of 

stem(s); distal process cylindric, truncate, 2-4 mm long; 

leaflets (6) 10-14, opposite to alternate, oblong-elliptic, 

chartaceous, 6-23.5 x 2-5.7(8.5) cm, both surfaces 

glabrous, except for the minutely pubescent (sometimes 

with glandular hairs) midvein, the abaxial surface rarely 

sparsely minutely hispidulous, the venation brochidodromus, 

flattened or slightly impressed, adaxially, prominent 

abaxially, tertiary veins reticulate, the apex acumi- 

Common names. Surinam: Zwarte pintolokus; 

Venezuela: Cotoperis montanero, cotuperia, cotoperis; 

Jatoama. 

Representative specimens examined. VENEZUELA. 

Amazonas: Atabapo, Orinoco river, 5 km N of 

confluence with Orinoquito river, hills, 400 m, s.d. (fl), 

Marin 1608 (NY, PORT, US), Oct 1991 (fl), Marin 1654 

(NY, PORT, US). Bolivar: Botanamo-Los Yagrumales, 

60 m, 14 Jul 1959 (fr), Bernardi 7998 (MER); Vic. of 

El Paraiso, 1 Aug 1965 (fr), Blanco 303 (NY-2, VEN); 

El Dorado, 9 Apr 1957 (fr), Couret 301 (US); Along El 

Tigre-La Soledad road, 29 May 1965 (fl), Marcano B. 

nate, the base inequilateral, one side obtuse the other 677 (F, IAN, NY, U, US, VEN); El Palmar, 75 m, 24 Nov


3


0Wl ._ OW=Lt, 7 

_WI eaN 50W! 40W 

*Talisia ghilleana L2 > N 

V Talisia guianensis/ 

* Talisia hemidasya \ ;, 

! 

( (Y I~~~~~~~~~ 


1967 (fr), Wessels Boer 2090 (MER, U, VEN). Delta 

Amacuro: East of Rio Grande, near Bolivar border, Nov 

1965 (fr), Blanco 368 (MER-2, MO, US); Tucupita, ESE 

of Los Castillos de Guayana, 50-200 m, 28 Mar-2 Apr 

1979 (fl), Davidse & Gonzdlez 16385 (NY, US, VEN); 

Vic. of Los Castillos, 30 m, 12 Jul 1960 (fr), Little 17676 

(MER, VEN); East of Rio Grande, near Bolivar border, 

29 Nov-18 Dec 1964 (fr), Marcano B. 457 (MER, MO, 

NY, US, VEN). 

GUYANA. Demerara/Mahaica: Demerara river, 

May 1889 (fl), Jenman 4925 (K-2). Cuyuni/Mazaruni: 

Takutu creek to Puruni river, 24 Oct 1944 (fl), Fanshawe 

2046 (K-3, NY, P, US-2). Potaro/Siparuni: Northern 

Pakaraima Mts., Ciong valley, 9 km N of Kato Village, 

900 m, 31 May 1995 (fr), Mutchnick & Tiwari 1440 

(CAY, US). Upper Takutu/Upper Essequibo: Maparri 

river, 6 June 1996 (fr), Clarke 2015 (US). 

SURINAM. Brokopondo: Brownsberg, 18 Oct 1924 

(fl), Zaandam 6705 (NY). Nickerie: Corantijne river, 

vic. of Wonotobo, 14 Oct 1916 (fl), Boschwezen 2574 

(K, MO, NY); Wilhelmina Gebergte, 3.5 km SE of 

Juliana Top, 11.5 km N of Lucie river, 450 m, 8 Aug 

1963 (fl), Irwin et al. 54600 (NY); 3 km S of Juliana 

Top, 12 km N of Lucie river, 500 m, 24 Aug 1963 (fl), 

Irwin et al. 55015 (F, NY, US); 14 km N of Lucie river, 

900 m, 13 Aug 1963 (st), Irwin et al. 54729 (NY); 

Kabalebo Dam project, 30-130 m, 13 Sep 1980 (fl), 

Lindeman et al. 406 (CAY, MG, NY, US). Saramacca: 

Coppename river near Raleigh falls, 13 Sep 1933 (fl), 

Lanjouw 812 (K, IAN, MO); Tafelberg, 11 Aug 1944 

(fl), Maguire 24341 (NY), 1400 m, 8 Sep 1944 (fl), 

Maguire 24690 (K-3, NY-2, US-2, VEN). 

FRENCH GUIANA. Bassin de la Mana: Montagnes 

de la Trinite, 5 Feb 1984 (fr), de Granville et al. 6516 

(CAY, NY, P, U-3,. US). Bassin du Maroni: Grand Inini 

river, 5 Sep 1970 (fl), de Granville B-3695 (CAY, P). 

Ile de Cayenne: Cayenne, without specific locality, with- 

out date (fl), Martin s.n. (K). Region de Saul: Route de 

Belizon, 20 May 1992 (st), Acevedo R. et al. 4966 (NY, 

US); Monts La Fumee, 26 Oct 1982 (st), Boom & Mori 

2416 (CAY), 4 Oct 1973 (fl), de Granville B-5074 (CAY- 

2, P), 200-400 m, 24 Sep 1982 (fl), Mori et al. 14988 

(CAY, F, MO, NY, US); Route de Belizon, 13 km S of 

Eaux Claires, 200-300 m, 8 Oct 1991 (fl), Mori et al. 

22006 (CAY, NY, P, US); Sentier Botanique, 22 Sep 1995 

(fl), Mori et al. 24182 (CAY, NY, US); Monts La Fumee, 

500 m, 3 Nov 1971 (fl), Oldeman 3241 (CAY-2, NY, 

P); Montagne Boeuf Mort, 30 Oct 1971 (fl), Oldeman 

B-4180 (CAY-2, P, US-2). 

BRAZIL. Maranhao: Moncao, Kaapor Indian re- 

serve, 6 May 1986 (st), Balee 2185 (NY). Park: Itacaiunas 

river, 2 Dec 1981 (fr), Daly et al. 1672 (INPA, MG, NY). 

2QU 

34. TALISIA HEXAPHYLLA Vahl, Eclog. Amer. 

2: 29. 1798. Type. South America. Without specific 

locality or date (fl), von Rohr 99 (lectotype, C-505, 

here designated). Syntype. South America. without 

specific locality or date (fl), von Rohr 98 (C-503, 

C-504, MO). 

Tree (4-)10-30 m tall, usually branched on the upper 

portion. Young stems striate, puberulent or pubescent, 

becoming terete and lenticellate. Leaves paripinnate or 

less often imparipinnate; distal process filiform, ca. 5 mm 

long, early deciduous leaving a truncate base 2-3 mm 

long; leaflets 6-12, altemate, subopposite or opposite,


6.5-23.5 (-68) x 2.4-7.1 (23) cm, chartaceous to co- termediate between those defining subgenus Pitombaria 

riaceous, the adaxial surface glabrous, with prominent and subgenus Talisia and it reseambles T clathrata in 

or sunken midvein and slightly impressed secondaries, numerous morphological aspects, suggesting that they 

the abaxial surface glabrous or puberulent, with may be closely related and perhaps belong together in 

prominent midvein and secondary veins, the venation subgenus Pitombaria. 

brochidodromus, secondary veins alternate or 

subopposite, arching toward the margin, the margins Key to the subspecies of Talisia hexaphylla 

entire or wavy, slightly revolute; petiolules nearly cy- 1. Leaflets with 18-28 pairs of secondary veins; 

lindrical or enlarged and slightly flattened, wrinkled tertiary veins reticulate-clathrate 

when dry, 6-17 mm long, glabrous or puberulent; ra- . .............................. T hexaphylla subsp. multinervis 

chis 5-36 cm long, adaxially bicarinate on distal portion, 1. Leaflets with 8-17 pairs of secondary veins; 

slightly flattened on proximal portion, abaxially striate, tertiary veins reticulate. 

puberulent; petioles 4-18 (25) cm long, striate or less 2. Leaflets oblanceolate or elliptic, the apex 

often lenticellate, adaxially flattened, enlarged at base. 

Thyrses simple or panicle-shaped, fasciculate, axillary, 

ramiflorous or cauliflorous; cataphylls acicular, pubescent, 

5-7 mm long, clustered at base of inflorescence 

or wanting; axes 2-27 cm long, angled, ferruginous or 

canescent-pubescent, or puberulent; bracts subulate or 

abruptly acuminate, obtuse or seldom 

retuse T. hexaphylla subsp. hexaphylla 

2. Leaflets oblong to lanceolate, the apex 

long-acuminate to caudate 

..................................... T hexaphylla subsp. elegans 

filiform, persistent, 0.5-2 mm long, with same indumentum 

as the axis; dichasia simple or compound, 34a. TALISIA HEXAPHYLLA subsp. HEXAsessile 

or nearly so; pedicels 1-3 mm long, articulate PHYLLA Figs. 17e-f, 8la-g 

from the middle to the apex. Calyx 2.5-4.6 (6) mm long, 

grayish pubescent to ferruginous-tomentose on both 

surfaces, sometimes with a few glandular hairs, the sepals 

ovate, 1/3 to 1/2 the length of the calyx, acute or 

Talisia panamensis Pittier, Contr. U.S. Natl. Herb. 

20: 129. 1918. Type. Panama. Darien: Pinogana, 

in forest, 16 Apr 1914 (fl), Pittier 6534 (holotype, 

US; isotypes, MO, US-2) 

obtuse at the apex; petals elliptic to oblanceolate, white, Talisia cararensis Cuatrecasas, Rev. Acad. Colomb. 

3-5.8 (6.5) mm long, reflexed at anthesis, abaxially glabrous 

or pubescent at base and sometimes ciliate along 

lower margins, adaxially puberulent-papillate, the apex 

obtuse or rounded, the base tapering; appendages as long 

Cienc. 8: 306. 1951. Type. Colombia. Santander: 

Puerto Berrio, between Carare & Magdalena rivers, 

100-700 m, 28 Mar 1935 (fl), Haught 1608 

(holotype, F, photo at US; isotypes, NY, US). 

as the petals, or little shorter, elliptic to long-deltate, Leaflets elliptic, oblanceolate, or less often obovate, 

erect, entire or notched at apex, adnate along its lower sometimes with scattered wart-like protuberances, the 

half to the petal, abaxially puberulent and papillate, adaxial surface sometimes drying grayish, the abaxial 

adaxially sericeous; disc cup-shaped, 5-lobed, pubes- surface sometimes papillate, usually drying brownish, 

cent or tomentose at the apex, or less often glabrous, the venation lighter than the blade, tertiary venation 

1-1.3 mm tall; stamens (7)8, the filaments glabrous, in reticulate, the apex abruptly acuminate, obtuse or seltwo 

or three sets of unequal lengths, the shorter ones dom retuse, the base asymmetrical, attenuate or obtuse. 

2-3 mm long, the longer ones 4-5 mm long, or some- Fruits sometimes reaching 4 cm long. 

times nearly equal, the anthers ellipsoid to oblong, 0.6- Talisia cararensis is considered here to be a synonym 

1.2 mm long, glabrous, apiculate or seldom obtuse at of T hexaphylla subsp. hexaphylla as the only characapex; 

ovary ovoid, densely sericeous, the stigma elon- ter differentiating it from the latter is the size of the leaves 

gated, papillate. Fruits ellipsoid to ovoid, brownish-yel- and leaflets. Leaf size is a variable character in T 

low or yellow, nearly smooth, densely ferruginous-seri- hexaphylla and in general is not well represented in 

ceous, (1.5) 2.5-3 x 1.3-2.2 cm, obtuse at apex, with herbarium collections. Although leaf and leaflet size in 

short apiculum, the pericarp woody, 2-3 mm thick, the the type specimen of T cararensis falls outside the range 

endocarp granulate, sparsely to densely woolly. Seed of variation present in T hexaphylla, I do not feel that 

ellipsoid, 1-2 cm long, with sweet, white fleshy testa. this variation alone is enough to recognize T cararensis 

Embryo with cotyledons superimposed, of equal size. as a separate entity. The leaf gigantism in T cararensis 

The specific epithet refers to the leaves with 6 leaf- apparently represents a collection artifact. 

lets as shown in the type specimens. 

Phenology. Collected in flower during March. 

Note: I have placed T hexaphylla in Talisia subgenus 

Talisia with some reservations. Its anthers (ellip- Distribution and Ecology. (Fig. 84) From Panama 

soid or oblong with apiculate or obtuse apex) are in- south to Bolivia, in terra firme, vaLrzea, gallery, and sec-


X 

/mm~ 

n~Imm it4m 3mm~ ~ 

Fig. 81. Talisia hexaphylla subsp. hexaphylla. A. fruiting branch. B. inflorescence. C. staminate flower and 

i.c. same. D. staminate flower with perianth removed showing pubescent disc, stamens, and detail of obtuse anther. 

E. staminate flower with perianth removed showing glabrous disc, stamens, and detail of apiculate anther. P adaxial 

and abaxial views of petal with adnate appendage. E. l.s. staminate flower showing disc, stamens, pistillode, and 

ovules. F. pistillate flower. G. lateral, adaxial, and abaxial viewa of petal with adnate appendage. staminate flower 

with perianth removed showing disc, stamens, and detail of anther. H. pistillate flower with perienth removed 

showing disc, sterile stamens, and pistil and detail of anther, I.s. same. I. fruit. (A-E from Pittier 8814; F-H from 

Haught 1608; I from Steyermark 91248). 

3cm.


ondary forests, dry semideciduous-evergreen forests, 

and wooded areas bordering savanna. 

Common names and uses. Bolivia: Japunaki, 

piton. Guyana: mauraballi, muro-balli, sand mora, 

morokwe. Peru: shatonilla, shatona. Trinidad: cacao 

macaque. Venezuela: Cotopalo, cotoperiz, cotoplis, 

mamon cutuplis. Used in Guyana as a fish poison plant; 

the seeds are reported to have a sweet, edible testa. 


Without data, Mutis 1154 (US). Antioquia: San Luis, 

Canion Rio Claro, northern area, 4 Jun 1984 (bd), 

Cogollo 1823 (HUA, JAUM); Sons6n, Concesi6n de 

Cementos Rio Claro, 450 m, 17 May 1990 (st), Cogollo 

& Cardenas 4476 (JAUM). Caldas: La Dorada, 200 m, 

23 Apr 1963 (fl), Espinal 1201 (COL, MEDEL). 

VENEZUELA. Aragua: Quebrada de Quiterio, Feb 

1966 (fl), Aristeguieta 5994 (NY, VEN); La Trilla, ca. 

km 36 of road from Maracay to Ocumare, 325 m, 12 Mar 

1990 (st), Edwards 335 (US); La Cerilla, Valle de Ocumare, 

100 m, 6 Jun 1939 (fr), Pittier 14253 (VEN-2). Bolivar: 

Takutu/Upper Essequibo. Arawau river, Bonasika Land- 

ing, 16 Jul 1934 (fl), Archer 2309 (NY, US). 

FRENCH GUIANA. Bassin de L'Approuague: Sta- 

tion des Nouragues, 29 Jul 1989 (st), Sabatier & Prevost 

2949 (CAY, US). Bassin du Maroni: Grand Inini river, 

14 Jul 1990 (st), Sabatier & Prevost 3269 (CAY, US). 

Bassin du Sinnamary: Above Petit Saut, between Plomb 

and Tigre creeks, 500 m, 4 Sep 1993 (st), Mori et al. 23588 

(NY). Piste de Saint Elie: Station Biologique ORSTOM, 

2 May 1992 (St), Acevedo R. 4879 (CAY), 12 May 1992 

(st), Acevedo R. 4940 (CAY, US); Saut Vata-Saut Berard, 

22 Sep 1965 (fr), Oldeman 1531 (CAY). Region 

Littorale: Passoura creek, 9 May 1992 (st), Sabatier & 

Prevost 4010 (CAY). 

PERU. Loreto: Prov. Maynas. vic. Sucusari, 

Explornapo Camp, 100-140 m, 3 Mar 1991 (st), Pipoly 

et al. 14227 (MO); Ucayali, 250 m, 20 Oct 1982 (st), 

Reynel 689 (K). San Martin: Huallaga, Cuzco, 25 Aug 

1986 (st), Alban 2396 (F); Bellavista, Huallaga Valley, 

200-300 m, 5 Sep 1948 (fl), Ferreyra 4749 (US); 

Tarapoto, 7 Oct 1984 (fr), Maas et al. 5973 (K, NY, U, 

USM); Pucacaca, Huallaga river, 30 Apr 1976 (fr), Plow- 

man 6016 (F); Chazuta, 264 m, 4 Oct 1963 (fl), Schunke 

V 6335 (F, US); Semi-cultivated land, 345 m, 19 Dec 

1974 (fl), Zegarra 11 (US). 

BRAZIL. Acre:. Tarauaca. Tarauaci river, Seringal 

Mucuripe, 19 Sep 1994 (fr), Daly et al. 8256 (US); 

Macauhan river, 25 Aug 1933 (fl), Krukoff 5630 (F, MO, 

NY, US), 1 Sep 1933 (fr), Krukoff 5725 (F, M, MO, NY, 

US); Cruzeiro do Sul, 28 Feb 1976 (fr), Ramos & Mota 

152 (INPA). Amazonas: Boca do Acre, Purus & Acre 

rivers, Lago da Cobra, 19 Sep 1966 (fl), Prance et al. 

2425 (F, K, MG, MO, NY, P, U, US). 

BOLIVIA. Beni: Prov. Ballivian. Serrania Pil6n Lajas, 

300 m, 8-9 May 1991 (st), Killeen et al. 3244 (US); Prov. 

Itenez. Road to Puerto El Carmen, 126 km NE of Trinidad, 

29 Oct 1993 (fr), Moraes et al. 1360 (LPB, US); Yacuma, 

Cuberene river, above San Juan, 200 m, 28 Nov 1991 

Isla del Lago Guri, 40 km S of camp, 270 m, 26 Feb 1992 

(st), Aymard et al. 10231 (PORT), Aymard et al. 10271 

(PORT); Calzeta de la Botella, El Dorado, 100 m, 14 Apr 

1957 (bd), Bernardi 6536 (NY); La Paragua forest reserve, 

Asa river, Jun 1970 (fr), Blanco 791 (F), 42-50 km from 

La Paragua, Jan 1984 (st), Guevara & Rojas 357 (MER); 

Represa Guri, 220-250 m, 7 Apr 1981 (fl), Liesner & 

Gonzalez 11327 (MO, NY, TFAB, VEN); Botanamo river, 

40 SE of Tumeremo, 100 m, 5 Jul 1960 (fr), Little 17598 

(MER-3, VEN); Altiplanicie de Nuria, 570 m, 20 Jul 1960 

(st), Steyermark 86555 (NY); Pica, La Lira near El Dorado, 

220 m, 25 Jul 1960 (fr), Steyermark 86624 (NY, US, VEN); 

Cefro Cotorra (El Vigia), 5 Aug 1960 (st), Steyermark 86857 

(NY, VEN). Carabobo: Selva de Guaremales, 2 May 

1920 (fl), Pittier 8814 (P, M, US, VEN). Delta Amacuro: 

Cuyubini river, 100-200 m, 18 Nov 1960 (fr), 

Steyermark 87623 (NY, VEN); Guanamo river, 300 m, 

22 Nov 1955 (st), Wurdack & Monachino 39719 (NY). 

Distrito Federal: Without specific locality, 900 m, 31 

Jul 1982 (fr), Manara s.n. (VEN-2). Miranda: Cerros del 

Bachiller, 18-20 m, 18-19 Mar 1978 (fl), Steyermark 

116373 (VEN). Sucre: Peninsula de Paria, Cerro Patao, 

N of Puerto Hierro, NE of Giliria, rocky creek, 350 m, 

23 Jul 1962 (fr), Steyermark & Agostini 91248 (K, US, 

VEN). Yaracuy: Quebrada El Charal, Aug 1970 (fr), 

(st), Seidel et al. 6040 (LPB); Trinidad-Misiones, 

Guarayos, 250 m, Sep 1926 (fl), Werdermann 2560 (LPB, 

MO, U). La Paz: Prov. Sud Yungas, Alto Beni, concesi6n 

of Sapecho Cooperative, 600 m, 16 Nov 1991 (fr), Seidel 

et al. 5618 (US), 530 m, 1 Dec 1993 (fr), Seidel & Vaquiata 

7496 (LPB, US). Santa Cruz: Buena Vista, 450 m, 15 

Oct 1924 (fl), Steinbach 6562 (F). 

Blanco 930 (MER, VEN). 

TRINIDAD. Without data, 20 Mar 1989 (fl), Anony- 34b. TALISIA HEXAPHYLLA subsp. ELEGANS 

mous 3633 (US-2); St. Amis Botanic Gardens, cultivated, Acevedo-Rodriguez, subsp. nov. 

15 Apr 1925 (fl), Broadway 5598 (K-3, MO-2); Belmont, 

without locality, 26 May 1922 (fl), Broadway 10138 Type. Colombia. Antioquia: Mun. Turbo. Tapon del 

(K, NY); St. Clair Royal Botanic Gardens, Apr 1938 (st), Darien road, Rio Leon-Lomas Aisladas area, km 37, 

Dean 13326 (K-2); without data, 10 Apr 1861 (fl), disturbed primary forest on soggy soils, 20 m, 28 Feb 

Grueger 211 (K-2), 1786-91 (st), von Rohr 98 (MO). 

GUYANA. Barima/Waini: Koriabo river, 20 Jul 

1934 (fr), Archer 2378 (US). Essequibo Islands/West 

1984 (fl), Brand & M. Gonzalez 961 (holotype, COL; 

isotypes, HUA, JAUM-2 sheets) Fig. 82a-g 

Demerara: Essequibo river, Kamwatta Ck., 31 Jul 1918 A T. hexaphylla subsp. hexaphylla foliolis oblongis 

(yfr), Hohenkerk 3a (K). East Berbice/Corentyne: New vel lanceolatis, apicibus longe acuminatis vel caudatis 

river, steep slope, 4 Oct 1952 (st), Guppy 361 (NY). Upper differt.


~~~~~~~~ .~~~~~~~~~~~~m 

2mm ~ ~ ~ ~ ~ ~ ~ ~ ~ ~~~~~~~~~~ m 

Fig. 82. Talisia hexaphylla subsp. elegans. A. sterile branch. B. inflorescence. C. staminate flower. D. lateral, abaxial, 

and adaxial views of petal. E. staminate flower with perianth removed showing disc and stamens. F. infructescence. 

G. embryo, lateral view. (A, F-G from Brand & Cogollo 79; B-E from Brand & Lozano 923).


Leaflets oblong, lanceolate or seldom nearly elliptic, 

the adaxial surface drying grayish, the abaxial surface 

drying brownish, tertiary venation reticulate, the apex 

long-acuminate to caudate, the base asymmetrical, one 

side obtuse, the other rounded. Calyx ca. 5 mm long, 

ferruginous-tomentose, the sepals 1.5-2 mm long. Fruit 

ellipsoid, minutely sericeous, glabrescent. 

The new subspecies differs from the typical subspecies 

by the characters shown in the key to subspecies; 

the subspecific epithet refers to the elegant foliage. 


February, and in fruit in May, July and October. 


from Colombia and Panama in primary or disturbed, 

seasonally flooded forest. 

river and mouth of Napo river, 120 m, 16 Jul 1983 (st), 

Gentry et al. 43173 (MO); Llachapa, Napo river, 130 

m, 20 Jan 1983 (st), Vdsquez & Jaramillo 3770 (MO, 

NY); Madre de Dios: Prov. Manu. Parque Nacional del 

Manu, Manu river, 350 m, 4 Aug 1984 (fl), Foster 9747 

(MO, USM); Prov. Tambopata. Tambopata Reserve, 30 

km S of Puerto Maldonado, 260 m, 8 Nov 1984 (fr), 

Young & Stratton 135 (MO). 

35. TALISIA LAEVIGATA Acevedo-Rodriguez, 

BioLlania Edicion Esp. 6: 145. 1997. Type. Ven- 

ezuela. Bolivar: La Paragua Forest Reserve, vic. 

of Forestry camp "Potreritos", Feb 1970 (fr), 

Blanco, C. 652 (holotype, US; isotypes, K, MER, 

P, VEN). Fig. 85a-b 

Representative specimens examined. PANAMA. Tree to 16 (?)m tall. Stems terete, smooth, glabrous 

Without data, 1966 (fr), Bristan 184 (COL). at maturity, light brown. Leaves paripinnate; distal pro- 

COLOMBIA. Antioquia: Turbo. Road to Tap6n del 

Darien, Rio Leon-Lomas Aisladas sector, km 11, 10-20 

m, 20 May 1984 (fr), Brand 938 (JAUM); 28 May 1984 

(fr), Brand 1204 (JAUM), 24 Dec 1983 (fl), Brand & 

Escobar 713 (JAUM), 24 Oct 1983 (fr), Brand & 

Gonzdlez 518 (JAUM), km 37, 20 m, 26 Feb 1884 (fl), 

Brand & Lozano 923 (JAUM); Barranquillita, 2 km after, 

80 m, 9 Jul 1981 (fr), Brand & Cogollo 79 (COL, HUA, 

cess truncate, 2-3 mm long; leaflets 6-1 0, altemate to 

subopposite, strongly involute, elliptic, 4.5-9 x 2-3.5 

cm, coriaceous, slightly discolorous, involute, glabrous 

on both sides, shiny on upper surface, the venation 

brochidodromus, prominulous on both surfaces, especially 

the midvein, tertiary venation reticulate, inconspicuous, 

the apex obtuse, mucronulate, the base ob- 

JAUM); Acandi, El Paramo trail, headwaters of La Qui- tuse sometimes asymmetrical and decurrent onto the 

eta creek, 150 m, 22 May 1989 (fr), Fonnegra et al. petiolule; petiolules pulvinate, 1.5-2 mm long, glabrous; 

2795 (HUA, NY). 

rachis 4-6.5 cm long, slender, terete to slightly angled, 

puberulent; petioles pulvinate at base, 2.5-4 cm long. 

34c. TALISIA IEXAPHYLLA subsp. MULTINER- 

VIS Acevedo-Rodriguez, subsp. nov. 

Thyrses panicle-shaped, to 30 cm long, terminal on 

lateral branches; cataphylls minute, acicular, minute, 

early deciduous; axes angled, puberulent to tomentulose; 

Type. Peru. Loreto: Itaya river, above Iquitos, 14 bracts triangular, early deciduous, ca. 1 mm long; dicha- 

Aug 1972 (fr), TB. Croat 19160 (holotype, US; sia seemingly simple and sessile; pedicels 1.5-3 mm long, 

isotypes, F, MG, MO, NY, USM). Fig. 83a-e articulate at middle. Calyx ca. 2.5 mm long, tomentulose, 

A T. hexaphylla subsp. hexaphylla foliolis numerosis, 

nervis tertiaris clathratis differt. 

Leaflets elliptic or seldom oblanceolate; secondary 

veins 18-28 pairs; tertiary veins clatirate, the apex longacuminate 

to caudate, the base asymmetrical, one side 

obtuse the other rounded. 

The new subspecies differs from the typical subspecies 

by the characters shown in the key to subspecies. 

The subspecific epithet refers to the numerous conspicuous 

secondary venation present on leaflets. 

the sepals 1.7-2 mm long, ovate, concave, rounded at 

apex, ciliate; petals elliptic, ca. 4 mm long, reflexed, papillate 

on adaxial surface, glabrous on abaxial surface, the 

apex rounded, the base attenuate; appendages erect, 

slightly longer than the petal, lanceolate, sericeous-pubescent 

on both surfaces, obtuse at apex, adnate at base 

along 1/4 of the petal length; disc 5-lobed, glabrous, 0.5 

mm tall; stamens 8, the filaments of unequal length, 2-2.2 

mm long, glabrous, the anthers ca. 1.7 mm long, oblong-lanceolate, 

long-apiculate at apex; ovary not seen, 

the stigma elongated, ferruginous-papillate. Fruits (im- 

Phenology. Collected in flower in August, and in 

fruit in November. 


from the Departments of Loreto and Madre de Dios, 

Peru, in lowland, seasonally flooded forest. 

mature) obovoid, glabrous, smooth. 

Talisia laevigata is perhaps closer to T cupularis 

Radlkofer than to any other species of Talisia since they 

share many morphological features. Both species have 

coriaceous, slightly discolorous, glabrous leaflets with 

pulvinate petiolules. Talisia laevigata, however, differs 

Representative specimens examined. PERU. from T cupularis by its smaller (4.5-9 cm long) involute 

Loreto: Yanamono, Amazonas river, between Indiana (vs. 7-20 cm long and flattened, and revolute) leaflets,


IT" ~ ~ ~ ~ ~ 1 

Fig. 83. Talisia hexaphylla subsp. multinervis. A. portion of leaf and branch. B. staminate flower. C. lateral and 

adaxial views of petals. D. staminate flower with perianth removed showing disc and stamens. E. portion of 

infructescence. (A from Gentry et al. 43173; B-D from Foster 9747; E from Young & Stratton 135).


sowl 70WI 60W 5dowl 40W 

I\)r~~~~~~~~~~~~~I 

Fig.m84.Distributif 

Talises ssi m 

iin 

14 

A Talsia hxaphylaFis. elegan uto fsece nTlii ugnu aii (npr) 

and glabrous (vs. hirsute) floral disc. Immature fruits 

in T laevigata are completely glabrous, suggesting that 

the ovary may have been glabrous too. Under this as- 

sumption, Ti cupularis would also differ from T. 

laevigata by its hirsute ovaries. Flowers were described 

from flower remnants at the base of fruits. The specific 

epithet refers to the shiny upper surface of leaflets. 

Phenology. Collected in fruit in February. 

Distribution and Ecology. (Fig 89) Known only, 


Common names. Palo azul, mamoncillo. 

36. TALISIA LONGIFOLIA (Bentham) Radlkofer, 


Wiss Miinchen 8: 348. 1878. Cupania longifolia 

Bentham, Hooker's J. Bot. Kew. Gard. Misc. 2: 211. 

1850. Type. Brazil. Para: without specific locality, 4 

Jul-Aug 1849 (fl), Spruce s.n (lectotype, K, here 

designated; isotype, K; frag. at M). Figs. 7, 86a-h 

Slender tree or treelet, (3)9-20 m tall (ca. 50 m fide 

Bentham), unbranched or with few ascending, sympodial 

branches on distal portion. Trunk to 20 cm in diam; 

bark brown, with numerous black, protruding lenticels; 

inner bark orangish. Distal portion of stem sulcate, glabrous, 

lenticellate. Leaves paripinnate, spirally arranged 

on distal portion of stem(s); distal process minute, truncate 

at apex; leaflets 8-16 (20), opposite or alternate, 

elliptic, oblong or less often oblanceolate, coriaceous, 

(6.5)11.5-30 (54) x (2.3) 3-8.5 (14) cm, glabrous, the 

adaxial surface with prominent midvein, the abaxial 

surface with scurfy-like punctations, with prominent 

midvein and secondary veins, tertiary veins reticulate, 

the apex acute to acuminate or less often rounded, the 

base inequilateral, one side obtuse the other acute or 

attenuate; petiolules 0.5-1 x 0.2-0.7 cm long, conical, 

pulvinate; rachis (19) 25-45(60) cm long, glabrous, 

triangular at base, trullate and sharply angled towards 

apex; petioles 30-45 cm long, sharply trigonous, striate, 

gradually swollen toward base. Thyrses panicle-shaped, 

tenninal or axillary on distal part of branches, 19-50 cm 

long, lateral branches to 17 cm long; cataphylls acicular 

to subulate, persistent, 1-2 cm long; axes puberulent, 

obtusely angled, sulcate; bracts deltate, ca. 1 mm long; 

dichasia compound, sessile; pedicels 1-2 mm long, tomentose, 

articulate on distal portion. Calyx yellowish 

green, 1.2-2.5 mm long, pubescent to tomentulose, the 

sepals 0.5-1 mm long, oblong or ovate, concave; petals 

white, 2.5-4.5 mm long, oblong to oblong-lanceolate, 

appressed-puberulent or pubescent on the lower half 

2Q


Fig. 85. Talisia laevigata. A. fruiting branch. B. fruit. (All from Blanco 652).


cm. 3mm. 

Fig. 86. Talisia longifolia. A. portion of leaf. B. portion of inflorescence. C. pistillate flower. D. adaxial and 

abaxial views of petal with adnate appendage. E. pistillate flower with perianth removed showing disc, sterile 

stamens, and pistil. F. infructescence. G. fruit, seed, and embryo. H. fruit. (A from Spruce, July 1849; B-E from 

Irwin et al. 55885; F-G from Schubert 2234; H from Oldeman 185).


of the abaxial surface, sometimes completely pubes- Station des Nouragues, 25 Jul 1986 (st), Sabatier & 

cent, glabrous adaxially, ciliate at margins, rounded at Prevost 2872 (CAY), 26 Jun 1994 (st), Sabatier & Prevost 

apex, clawed at base; appendage as long as the petals, 

long-triangular, hirsute on the distal half of the adaxial 

surface, glabrous abaxially, the base cuneate to clawed; 

disc annular, tomentose; stamens 5, the anthers lanceolate, 

apiculate, glabrous, 1.2 mm long, the filaments 

pilose or glabrous, white, of equal length; ovary 

4291 (CAY, US). Bassin de la Camopi: Mont Belvedere, 

160 m, 27 Nov 1984 (fr), de Granville 7033 (CTES, CAY, 

NY, P, U, US); Camopi river, 12 Feb 1968 (fr), Oldeman 

& Sastre 185 (CAY, IAN, NY, P). Basin de L'Inini: 

Montagne Bellevue de L'Inini, 600 m, 29 Aug 1985 (st), 

de Granville et al. 7899 (CAY, MG, MO, P). Bassin du 

Maroni: Grand Inini river, Sai creek, 26 Aug 1970 (bd), 

densely appressed- and grayish tomentose, the style Oldeman B-3577 (CAY, P); near Village Fourmi, 20 Aug 

elongate with an elongated, trilobate, papillate, pinkish 1970 (fl), Oldeman B-3521 (CAY, P, US). Bassin de la 

stigma. Fruit usually 1-3-locular, orange-yellow at Waki: Ouaqui river, 4 Jul 1973 (bd), de Granville 4880 

maturity, trigonous-ellipsoid, 2-2.5 cm long, puberulent, 

the pericarp coriaceous, ca. 2 mm thick, granulose, 

the endocarp usually scattered pilose. Seeds ellipsoid, 

1 or 2 per fruit; cotyledons superimposed, the upper 

cotyledon larger than the lower one. 

This species is characterized by its sharply triangu- 

(CAY, P). Region de Saul: Monts La Fumee, 200 m, 10 

Sep 1982 (st), Boom & Mori 1633 (CAY, NY); Saul, 

220 m, 21 Jun 1988 (st), Gentry et al. 63037 (MO, NY). 

BRAZIL. Acre: Cruzeiro do Sul, secondary, 23 Feb 

1976 (yfr), Monteiro & Damido 606 (MG, INPA). 

Amapi: Serra do Navio, along trail from Porto 

Terezinha to Reservatorio, 70-300 m, 13 Nov 1954 (fl), 

lar petioles and lower parts of the rachis; the distal por- Cowan 38296 (IAN, K, NY, P); Vic. of Porto Terezinha, 

tion of the rachis is trullate; glabrous foliage; petals that 70 m, 22 Nov 1954 (fl), Cowan 38491 (F-2, US); Mun. 

are pubescent or puberulent on lower abaxial one-half; 

5 stamens with lanceolate anthers; and tomentulose disc. 

It is closely related to T marleneana, which has different 

flowers (8 stamens with ellipsoid anthers; petals 

that are appressed-pubescent abaxially; and leaflets that 

are elliptic with prominent primary and secondary veins. 

The specific epithet refers to the long leaves. 

Macapa; Riozinho, 122 km NW of Porto Grande, 1 Jan 

1985 (fr), Mori & Souza 17605 (K, NY); Mun. Mazagao. 

Jari river, Morro do Felipe V, 23 Oct 1985 (fl), Pires et 

al. 696 (INPA, MG-2); Araguari river, 8 Sep 1961 (fl), 

M. Pires et al. 50774 (MG-2, NY); Amapari river, 1 Oct 

1961 (fl), M. Pires et al. 51412 (IAN); Falsino river, 

upstream of confluence with Araguari river, 2 Oct 1983 

(fl), Rabelo et al. 2415 (MG, NY, US). Pari. Carapara, 

Phenology. Flowers from June to November and 

fruits from July to March. 

7 Sep 1908 (fl), Anonymous (MG-9625); Acara. Faz. 

Borba Gato vic. of Acara river, 7 Nov 1980 (fr), Daly et 

al. 872 (MG, NY); Cagancho creek, ca. 1 km E of dam 

Distribution and Ecology. (Fig. 89) Known from of Tucurui, 29 Oct 1981 (fr), Daly et al. 1040 (INPA, 

Colombia, the Guianas and Brazil in moist, non-flooded 

forest and riverine forest. 

K-2, MG, US); Vic. of Tucurui, 7 Nov 1981 (fr), Daly et 

al. 1245 (IAN, INPA, K, MG); Belem, Sao Braz, 4 Aug 

1895 (fl), Huber 9 (MG-2); Belem-Brasilia road, km 98, 

Common names. Brazil: Pitomba brava; Colom- 6 Aug 1963 (fl), Maguire et al. 56031 (MO-2, NY, US); 

bia:jicacara; French Guiana: gaulette, gaulette indien, Tome A9u, vic of Ipiranga creek, 3 Jan 1978 (fr), 

paicoussa. 

Nascimento 418 (K, MG); Cachoeira, km 96, 29 Oct 

1965 (fr), Prance & Pennington 1787 (F, IAN, K, M, 

Representative specimens examined. COLOMBIA. NY, US); Vic. of Paragominas, Bel6m-Brasilia hwy., km 

Amazonas: Caqueta river, W of Cafio Paujil, 30 Nov 161, 28 Aug 1964 (fr), Prance & Silva 58919 (F, NY, 

1988 (st), Sanchez et al. 1814 (COAH). 

US); Belem, Dec 1965 (fr), Schubert 2234 (K, IAN, US); 

SURINAM. Brokopondo: Brownsweg, 9 Sep 1917 Santarem, km 70 along road to Palhao, vic of Guarana 

(fi), Wessels Boer 3227 (K-2, MO, NY). Marowijne: 

Tumuc Humac Mts., 300 m, 16 Aug 1993 (fl), Acevedo 

R. et al. 6053 (CAY, F, K, NY, P, US), 600 m, 18 Aug 

1993 (fl), Acevedo R. et al. 6080 (CAY, K, MG, MO, 

creek, 9 Sep 1969 (fl), Silva & Souza 2524 (K, MG, MO, 

NY, US), km 35 on road to Palhao, vic of Curupira creek, 

4 Sep 1969 (fr), Silva & Souza 2500 (MG, NY, US). 

NY, P, US), w of Talouaken Peak, 600 m, 23 Aug 1993 

(fl), Acevedo R. et al. 6127 (CAY, K, MG, NY, US). 37. TALISIA MACROPHYLLA (Martius) 

Nickerie: Zuid river, near confluence with Lucie river, Radlkofer, Sitzungsber. Math.-Phys. Cl. K6nigl. 

220-250 m, 20 Sep 1963 (fl), Irwin et al. 55885 (F-2, BayerAkad. Wiss Miinchen 8: 347. 1878. Cupania 

NY-2,.US-2). Saramacca: Tafelberg, Table Mts., 17 Sep 

1944 (fl), Maguire 24797 (K-2, NY, US-2). 


Confluence of Alice & Tampok creeks, 24 Mar 1977 

(fr), Moretti 658 (CAY-2); Parepou creek, 25 Sep 1968 

(fl), Oldeman B-1883 (CAY, P, US); Approuague river, 

28 Jan 1967 (fr), Oldeman 2390 (CAY, NY, P); Arataye 

river, Feb 1969 (fr), Oldeman 2994 (CAY, IAN, NY, P); 

macrophylla Martius, Flora 21, Suppl.: 74. 1838. 

Type. Brazil. Bahia: Ilheus, 1819 (fl), Luschnath 

s.n. (Martius 483) (holotype, M-2 sheets, photo at 

US; isotype M). Fig. 87a-e 

Talisia elephantipes Sandwith, J. Bot. 78: 256. 1940. 

Type. Guyana. Cuyuni river, 20 Jul 1933 (fl), 

Tutin 405 (holotype, BM; isotypes, K, US).


Fig. 87. Talisia macrophylla. A. leaf and detail of leaflets. B. detail of dichasium. C. adaxial view of petal with 

adnate appendage. D. staminate flower with perianth removed showing disc and stamens (left) and pistillode (right). 

E. infructescence. (A from Acevedo R. et al. 4967; B-D from Mori & Pipoly 15533; E from Acevedo R. et al. 4967).


Talisia allenii Croat, Ann. Missouri Bot. Garden 63: 

527. 1976. Type. Panama. Darien: Tiura river, 

between Punusa & Mangle rivers, 1 Oct 1967 

(fl), Duke, J.A. 14578 (holotype, MO; isotype, 

PMA, n.v.). 

Talisia pentantha Steyermark, Ann. Missouri Bot. 

Gard. 75: 1075. 1988. Type. Venezuela. Bolivar: 

entire; disc annular, 5-lobed, glabrous or puberulent, 

0.4-0.5 mm tall; stamens 5, the filaments of equal length 

or slightly unequal, 1-2.2 mm long, glabrous or less 

often puberulous, the anthers 1.1-1.4 (2) mm long, 

oblong, oblong-lanceolate to linear, glabrous, apiculate 

at apex; ovary glabrous, the stigma elongate, nearly 

Gran Sabana, Canaima, W of Avensa Camp, gal- terete, papillate. Fruits 1(2)-seeded, ellipsoid to globose, 

lery forest on terra firme, 500 m, 4 Oct 1974 green, glabrescent, smooth, 1.7-3 x 1.1-3 cm, rounded 

(bd), Ehrendorfer 74104-23 (holotype, VEN). at apex, shortly apiculate, the pericarp 0.9 mm thick. 

Talisia sancarlosiana Steyermark, Ann. Missouri Bot. Seeds ellipsoid, to 1.5 cm long. Embryo with cotyledons 

Gard. 75: 1075. 1988. Type. Venezuela. Amazonas: 


Between San Carlos & Solano, 11-17 Mar 1970 

Talisia macrophylla seems to be closely related to 

(fl), Marcano Berti & P. Alcedo 119-979 (holotype, 

MER). 

T carinata. However, T macrophylla differs from T 

carinata by the following characters: leaflets larger; se- 

Treelet, 4 to 12 (25) m tall; trunkto 15 cm in diam. pals ovate-deltate (vs. ovate); petals smaller, oblong, 

Branches terete, smooth, minutely lenticellate, glabrous. abaxially glabrous and truncate at base (vs. elliptic, seri- 

Leaves paripinnate; distal process 2-3 mm long, trun- ceous and attenuate at base); disc glabrous (vs. tomencate 

at apex; leaflets (6)8-16(30), alternate, opposite or tose); filaments glabrous and shorter than the anthers; 

subopposite, oblong to elliptic, 10-45 x 3.7-13, chart- (vs. tomentose and longer than the anthers); anthers 

aceous to sub-coriaceous, glabrous, the adaxial surface oblong (vs. lanceolate); and fruits smooth (vs. granulose). 

with prominulous, plane or slightly sunken primary and Talisia macrophylla looks vegetatively similar to T 

secondary veins, the abaxial surface with prominent, cerasina (Bentham) Radlkofer. However, T macrophylla 

primary and secondary veins, the venation brochido- can be distinguished by its nearly sessile dichasia, 

dromus, with secondary veins alternate or subopposite, smaller flowers (corolla ca. 3 mm long, vs. 5 mm long) 

arching, tertiary veins reticulate, the margins undulate, with 5 stamens (vs. 8 stamens), and glabrous ovaries 

the apex obtuse, acuminate or seldom long-acuminate (vs. sericeous). In addition, leaflets of T macrophylla 

and mucronate, the base asymmetrical to strongly asym- are less strongly asymmetrical basally than those of T 

metrical, obtuse-cuneate, to nearly rounded (then one cerasina. Leaflets of Talisia macrophylla differ from 

side higher than the other, tapering into the petiolule; those of large-leaved T nervosa by having lower secpetiolules 

pulvinate, 4-12(35) mm long, glabrous, ob- ondary veins that do not form a closed loop at margins 

long-conical, canaliculate; rachis reddish or dark brown (all secondaries forming a closed loop). The specific 

20-58 (115) cm long, obtusely to narrowly rhombic epithet refers to the large leaflets of this species. 

distally, nearly terete and striate at base, glabrous, shiny, 

smooth; petioles terete, 27-37 cm long, gradually swol- 

Phenology. Flowers and fruits throughout the year. 

len toward base. Thyrses axillary, panicle-shaped, Distribution and Ecology. (Fig. 89) From Costa 

terminal on distal portion of main stem and branches, Rica to Colombia, Venezuela, Peru, French Guiana, and 

28-35 (100) cm long, the secondary branches 18(35) Brazil. In lowland, moist, primary or disturbed, terra 

cm long; cataphylls acicular, glabrous, reddish brown firme, varzea or igapo forests, caatinga on sandy soil 

5-8 mm long, in groups of 5 or 6; axes angled, sulcate, and palm swamp forest. 

sparsely puberulent; bracts nearly triangular-lanceolate, 

Common names and uses. Brazil: Pitomba do mato, 

persistent, 1-2 mm long; dichasia simple or compound, 

pitomba, pitombarana. Costa Rica: huesillo. French 

nearly sessile to shortly pedunculate; pedicels 0.3-0.7 mm 

Guiana: bois-flambeau. Panama. Choco: yarrepatado; 

long, articulate near the middle to just below the flower, 

fruits are said to be eaten by local people. Peru. Pasco: 

usually pinkish when immature, green at anthesis. Calyx 

copal rosado de hoja grande. Venezuela: cafecillo, 

pinkish in immature flowers, green at anthesis, 1.2-2.2 

sangre de lapa. 

(3) mm long, puberulent, the sepals 1.5-2.5 mm long, 

concave to nearly keeled, ovate-deltate to lanceolate, Representative specimens examined. COSTA 

obtuse or rounded at apex, the margins sometimes cili- RICA. Puntarenas: Palmar Norte, trail to Buenos Aires, 

ate with simple or glandular hairs; petals oblong or 830 m, 17 Feb 1951 (fl), Allen 5917 (F, US); Along road 

between Rinc6n & Osa, 150-260 m, 3 Mar 1985 (fl), 

oblong-lanceolate, 3-3.7(5) mm long, reflexed at an- 

Croat & Grayum 59760 (NY); Aguabuena, 3.5 km W 

thesis, papillate on adaxial surface, glabrous on abaxial 

of Rinc6n, 350 m, 18 Nov 1992 (st), Thomsen 675 (C). 

surface, rounded at apex, cuneate at base; appendages PANAMA. Without locality, 1967 (fr), Bristan 1511 

a little shorter than the petals, lanceolate, erect, villose (MO-2). 

on adaxial surface, papillate or glabrous on abaxial sur- COLOMBIA. Amazonas: Caqueta river, 3 km upface, 

sometimes with appressed hairs at base, the apex stream from Sumaeta island, 200-300 m, 5 Apr 1990 (fr),


Alvarez et al. 286 (NY); Igara-ParanA river, 15 km down- 3253 (CAY). Bassin de L'Oyapock: Roche Touatou, 120 

stream of La Chorrera, 7 May 1975 (fr), Idrobo 8091 m, 23 May 1995 (fr), Cremers & de Granville 14113 

(COL); Apaporis river, between Pacoa & Kananari rivers, (CAY, US). Bassin de la Yaloupi: Yaroupi river, vic. Saut 

Soratama, 250 m, 21 Jun 1951 (fr), Schultes & Cabrera Tainoua, 16 Apr 1970 (fl), de Granville 371 (CAY, P). 

12767 (US); Caqueta' river, north margin in front of Ile de Cayenne: without specific locality, Jul 1824 (fr), 

Mariname, Jul 1989 (st), Urrego et al. 887 (COHA). Poiteau s.n. (K). Piste de Saint Elie: Station Biologique 

Antioquia: Mun. Turbo. Turbo. Along Tap6n del Darien ORSTOM, 3 May 1992 (st), Acevedo R. 4885 (CAY). 

road, between Rio Le6n & Lomas Aisladas, 10-20 m, 25 Region de Saul: Route de Belizon, 2-4 km S of Eaux 

Oct 1983 (fr), Brand & Gonzalez 529 (COL-2, HUA); Claires, 160 m, 20 May 1992 (fr), Acevedo R. et al. 4967 

Along Tap6n del Darien road, between Rio Le6n & Lomas (CAY, NY, US); Eaux Claires, Sentier Botanique, ca. 200 

Aisladas, 10-20 m, 27 Nov 1983 (fr), Brand & Narvaez m from entrance, 25 May 1992 (fr), Acevedo R. et al. 

622 (JAUM), 15 Jan 1985 (fr), Brand et al. 1313 (JAUM); 5030 (NY, US); Saul. 220 m, 20 Jun 1988 (st), Gentry 

Cerro del Cuchillo, 14 Sep 1987 (fl), Cardenas 485 et al. 62957 (MO, NY); Eaux Claires, Sentier Botanique, 

(MEDEL); Uraba. Villa Arteaga, 50 m, 16-20 Feb 1953 250 m, 1 Feb 1990 (fl), Mori & Gracie 21079 (CAY, 

(fr), Schultes & Cabrera 18660 (US). Caqueth: Araracuara, NY); Monts La Fumee West, 200-400 m, 6 Apr 1983 

28 Jan 1989 (fl), Gentry et al. 65295 (MO); Cartagena, 20 (fl), Mori & Pipoly 15533 (CAY, MO-2). 

Apr 1953 (fr), Romero C. 4029 (COL). Choc6: Quibd6- ECUADOR. Carchi: Tulcan. Reserva Etnica Awa, 

Istmia road, between Yuto & Certegui, 11 Sep 1976 (fl), Canumbi river, 1150 m, 19-28 Feb 1993 (fl), Grijalva 

Forero & Jaramillo 2758 (COL); Quibdo-Tutunendo road, et al. 556 (US). Esmeraldas: Eloy Alfaro. Reserva 

ca. 3 km W of Tutunendo, 80 m, 6 Jan 1981 (st), Gentry et Cotacachi-Cayapas, Santiago river, 250 m, 23-27 Oct 

al. 30216 (MO); Sucio river, Parque Natural Los Katyos, 1993 (fr), Tirado et al. 552 (US), 28-31 Oct 1993 (fr), 

170-250 m, 18 Nov 1976 (fl, fr), Le6n 425 (COL, MO), Tirado et al. 683 (US). 

80-100 m, 9 Dec 1976 (fr), Le6n 721 (COL, MO). Guania: PERU. Loreto: Puerto Almendras, 130 m, 22 Feb 

Outskirts of San Felipe, 120 m, 8 Apr 1984 (st), Gentry & 1979 (st), Gentry et al. 24875 (MO); Explorama Inn, 

Stein 46473 (MO). Valle: Bajo Calima Concession, ca. 20 ca. 2 km W of Indiana on Amazonas river, 130 m, 12 

km n of Buenaventura, 50 m, 10 Jul 1987(st), Faber- Feb 1987 (st), Gentry et al. 55700 (MO); Mom6n river, 

Langendoen & Renteria 1258 (MO), 300 m, 27 Jun 1988 ca. 1 km above junction with Nanay river, 12 Jan 1976 

(st), Faber-Langendoen & Hurtado 1496 (MO). (fl), McDaniel & Rimachi 20424 (K); Vic. of Iquitos, 

VENEZUELA. Amazonas: Dpto. Casiquiare. Guainia. 120 m, 1977 (fl, yfr), Revilla 3596 (K, MO); Estacion 

Along road from Maroa to Yavita, ca. 700 m from Yavita, Biol6gica Rio Blanco, 150 m, 21 Sep 1985 (fr), Vasquez 

19 Feb 1998 (yfr), Acevedo R. et al. 10242 (PORT, US); et al. 6773 (MO); Puerto Almendras, 122 m, 25 Sep 1985 

Dpto. Atabapo. Upper Orinoco river, 35 km SE of La (st), Visquez & Jaramillo 6815 (MO); Vic. of Sucusari 

Esmeralda, 180 m, 16 Feb 1990 (fr), Aymard & Delgado along Napo river, 130 m 21 Feb 1989 (fl), Vdsquez & 

7915 (PORT, US); Vic. of Yavita, 125-140 m, 6-9 Jul Jaramillo 11807 (MO); Iquitos. Iquitos-Nauta road, 150 

1969 (fr), Bunting et al. 3827 (U); Mawarinuma river, m, 7 Apr 1989 (fr), Vasquez et al. 12008 (US); Yanamono- 

140 m, 22 Apr 1984 (st), Gentry & Stein 46857 (MO); Explorama Lodge, 108 m, 14 Feb 1990 (fr), V4squez et 

San Carlos de Rio Negro, ca. 20 km S of confluence of al. 13475 (US); Santa Maria de Nanay, Mishana, 5 Sep 

Rio Negro and Casiquiare rivers, 120 m, 5 May 1979 1990 (st), Vasquez et al. 14297 (MO). Pasco: Shirigamazu, 

(fr), Liesner 7222 (MO); Dpto. Atures. 5-8 km NW of ca. 20 km S of Iscozacin, Palcazu river Valley, 300 m, 6 

Yutaje, 700-1000 m, 10 Mar 1987 (bd), Liesner & Jul 1988 (st), Gentry et al. 63320 (MO). San Martin: 

Holst 21840 (MO). Bolivar: Sifontes; Concesion Juan Jui. Upper Huallaga river, 400-800 m, Apr 1936 

Minera Cristina Cuatro, 180 m, 11 Aug 1985 (fr), (fl), Klug 4303 (F, MO, US); San Martin. Chazuta; ridge 

Aymard et al. 4154 (MO, NY); Rauil Leoni, 230 m, 3 to W of Quebrada Chazuta, 200-300 m, 21 Sep 1986 

Nov 1988 (fr), Aymard & Fernandez 7327 (MO, PORT). (fr), Knapp 8351 (F, MO); Campanilla, San Eliseo, 395 

Distrito Federal: Tributary of Camuri Grande river, 1140 m, 19 Aug 1970 (fr), Schunke V 4275 (F, MO, NY, US); 

m, 20-22 May 1992 (bd), Meier et al. 2306 (VEN). Prov. Mariscal Caceres, 3 Sep 1970 (fl), Schunke V 4318 

GUYANA. Without Specific locality. Along Bartica- (F-2, MO, NY, P, US-2); Tocache Nuevo, Puerto Pizana, 

Potaro road, 26 Oct 1947 (fr), Fanshawe 2718 (K-2, NY- 2 Jan 1971 (fr), Schunke V 4613 (IAN, K, MO, NY, US); 

2, US-2). Upper TakutulUpper Essequibo: Chodikar creek Huinguillo, 500 m, 3 Mar 1962 (fl), Woytkowski 7172 

bank, ca. 300 m, 24 Nov 195? (st), Guppy 674 (NY). (MO-2, US), 3 Mar 1962 (fl), Woytkowski 7173 (MO-2). 

SURINAM. Marowijne: Tumuc Humac Mts. BRAZIL. Amaph: Maraca, 7 May 1982 (yfr), Rosa & 

Talouaken, 330 m, 8 Aug 1993 (fr), Acevedo R. et al. Martins 4324. (MG). Amazonas: Barcelos, Dermini river, 

5935 (CAY, K, US), 26 Aug 1993 (fr), Acevedo R. et al. 100 m, 10 Aug 1996 (st), Acevedo R. et al. 8218 (US); 

6145 (CAY, K, US). 

Manaus-Itacoatiara road, km 26, 3 Jun 1998 (fr), Assunado 

FRENCH GUIANA. Bassin de L'Approuague: 854 (US); Manaus-Itacoatiara road, km. 104, 15 Apr 1968 

Approuague river, Saut Parare, 13 Feb 1981 (fr), Barrier (fl), Bayron 68-111 (INPA); Mun. Barcelos, along Araci 

& Feuillet 2589 (CAY). Bassin de la Comtk: Montagne river, ca. Foz do Rio Jauri, riverside, 4 Jul 1985 (fr), 

Tortue, km 25 along Route de Belizon, 23 Apr 1992 Cordeiro 151 (NY); Serra de Araca, 15 Jul 1985 (fr), 

(fr), Acevedo R. et al. 4828 (CAY, US). Bassin du Cordeiro 215 (NY, US); Mun. Tonantins, Vila Velha, 

Maroni: Itani river, 16 Apr 1977 (fl), Moretti 715 (CAY); Tonantins river, 15 minutes upstream by motorboat from 

Grand Inini river, 13 Jul 1990 (st), Sabatier & Prevost confluence with Solimoes river, 18 Nov 1986 (fr), Daly et


al. 4345 (NY); Upper Rio Negro, 19 Nov 1929 (fl), Ducke 

s.n., herb 25218 (RB, U); Manaus, 27 Apr 1932 (fl), Ducke 

s.n. (RB 25219); Mun. Pres. Figueredo, Balbina hydroelectric, 

9 Mar 1986 (fl), C. Ferreira et al. 6676 (INPA, K, 

US); Uatuma river, 22 Mar 1986 (fr), C. Ferreira et al. 

6952 (INPA, NY); Mun. Sao Paulo de Olivenca, along road 

to Bom Fim, 25 Nov 1986 (fl), C. Ferreira et al. 8565 

(INPA, US); Mun. Atalaia do Norte, 18 Feb 1989 (bd), C. 

Ferreira et al. 9865 (NY); Curuca river, 5 km from 

confluence with Javari river, 1-18 Jan 1989 (fr), C. Ferreira 

et al. 9970 (NY); Mun. Borba. Transamazonica hwy., 1- 

5 km upstream from Sucunduri, 9 May 1985 (st), 

Henderson et al. 415 (INPA, MG, MO, NY, US); Manaus, 

Arara river, 26 Apr 1973 (fr), Loureiro et al. s.n. (INPA- 

37737); Rio Negro, close to Arara river, 29 Apr 1973 (fr), 

Loureiro et al. s.n. (INPA-37878, US); Road to Aleixo km 

7, 14 Mar 1956 (fl), Mello & Coelho s.n. (INPA-3597), 

Mello & Coelho s.n. (INPA-3599), Manaus, 19 Mar 1956 

(fl), Mello & Coelho s.n. (INPA-3634); Mun. Manacapuru. 

Manacapuru lake, 2 Apr 1976 (fl), Mello & Mota 47 (INPA, 

MG); Manaus-Porto Velho road, km 220, 28 Apr 1976 

(fl), Monteiro & Ramos 985 (INPA); Reserva Ducke 11 

Jan 1977 (st), Nascimento 392 (INPA); Serra de Araca, 4 

Mar 1984 (fr), Pipoly & Cress 6797 (INPA, NY, US), 

middle slopes of western massif, 20 Jul 1985 (fr), Prance 

& Cordeiro 29706 (INPA, MG, MO, NY, US); Manaus- 

Itacoatiara road, Km 8, Colonia Santo Ant6nio, 8 Sep 1966 

(fr), Prance et al. 2216 (K, MG, NY); Road to Aleixo, km 

19, 30 Mar 1967 (st), Prance et al. 4716 (US-2); Manaus- 

Itacoatiara road, km 211, 30 May 1968 (fr), Prance et al. 

4906 (F, K-2, INPA, MG, NY, US); Manaus-Igarape Leao 

road, 5 km from Manaus-Caracarai road, 26 Jan 1971 (fl), 

Prance et al. 11458 (K-2, NY, US); Ituxi river, vicinity of 

Boca do Curuquete, 10 Jul 1971 (fl), Prance et al. 14085 

(NY); Reserva Ducke, 17 Apr 1962 (fr), Rodrigues & 

Chagas 4399 (INPA); Cachoeira do Taruma, 26 Jul 1961 

(fr), Rodrigues & Lima 2242 (INPA); Rio Preto, 30 Jan 

1962 (fl), Rodrigues & Lima 4159 (INPA); road to Ponta 

Negra, Hotel Tropical, along Rio Negro, 26 Mar 1993 (fl), 

Rodrigues et al. 11089 (US); Serra de Aracia south of the 

central portion, 60 m, 20 Mar 1984 (fl), Rodrigues et al. 

10562 (INPA, NY); Road to Aleixo, 26 May 1972 (fl), 

Silva et al. 103 (INPA-2); Jauari river after Pretinho creek, 

4 Jul 1985 (fr), Silva 268 (NY-2); Between Castanho and 

Araca rivers, 13 Jul 1972 (fr), Silva et al. 626 (INPA); Rio 

Icana, Nazare, 10 May 1973 (fr), Silva et al. 1470 (INPA- 

2); Manaus-Itacoatiara road, km 26, 19 Jun 1997 (fr), de 

Souza 380 (US). Pari: Tucurui, without specific locality, 

11 Nov 1980 (fr), Lisboa et al. 1560 (MG, NY); Tapaj6s, 

km 183 along road Torquato-Tapajos, 5 Apr 1975 (fr), 

Loureiro et al. s.n. herb. 48443 (INPA); Mun. Oriximina, 

along road BR-163, vic. Cumina-Mirim river, 60 m, 3 Jun 

1980 (fr), Martinelli et al. 6761 (INPA); Basin of Xingu 

river, just downstream from mouth of Bacaja river, 28 Nov 

1980 (fl), Prance et al. 26521 (MG, US); Jari. Road to 

Munguba, 21 May 1969 (st), Silva 2010 (IAN, K, NY); 

Porto Trombetas, Ilha do Carana, 3 Mar 1986 (fl), Soares 

105 (INPA). Rondonia: Serra Pacaas Novas, westernmost 

hill, 175 m, 9 Apr 1987 (bd), Nee 34702 (NY); Basin of 

Madeira river, track from Mutuparana to Madeira river, 

30 Nov 1968 (fl), Prance et al. 9009 (F, K, M, MG, US). 

Roraima: Vic. Uaica, airstrip, Uraricoeira river, 1 Mar 1971 

(st), Prance et al. 10802 (K, MO, NY, P, U, US), Vic. of 

Uaicd, 1 Mar 1971 (fl), Prance et al. 10810 (INPA, K, M, 

MG, MO, NY, P, US). 

38. TALISIA MARLENEANA (G. Guarim Neto) 

Acevedo-Rodriguez, stat. nov. Talisia mollis var. 

marleneana G. Guarim Neto, Acta Amazonica 13: 

497. 1983. Type. Brazil. Para: Pocoes, Oriximina', 

12 Jun 1957 (fl), WA. Egler 585 (holotype, MG). 

Figs. 16c-d, 88a-h 

Slender tree or treelet, 1.5-7 m tall, unbranched or 

with few ascending, sympodial branches on distal por- 

tion. Trunk 2.5-8 cm in diam. Distal portions of stem 

nearly terete, smooth, puberulent. Leaves imparipinnate, 

spirally arranged on distal portions of stem(s); distal 

process early deciduous leaving a truncate patch at apex; 

leaflets 3-15 alternate or opposite, elliptic or oblong, 

chartaceous, 12-35.5 x 6-13 cm, glabrous, the adaxial 

surface with prominulous midvein and sunken secondary 

veins to produce a weakly bullate lamina, the abaxial 

surface with prominent midvein, secondary and tertiary 

veins, tertiary veins reticulate, the apex acuminate, the 

base inequilateral, one side rounded the other obtuse; 

petiolules to 8 mm long, pulvinate (conical), drying dark 

brown; rachis 20-75 cm long, glabrous, triangular at 

base, trullate and sharply angled towards apex; petioles 

17-35 cm long, sharply triangular, striate, gradually 

swollen toward base. Thyrses panicle-shaped, terminal 

or axillary on distal part of branches, or less often 

cauliflorous, 11-30 cm long; cataphylls acicular, minute; 

axes puberulent to nearly glabrous, obtusely to sharply 

angled, sulcate; bracts and bracteoles deltate, 0.5-1 mm 

long, tomentulose; dichasia compound, sessile; pedicels 

1.5-2 mm long, tomentulose, articulate above the middle. 

Calyx cream-colored, 1.5-2.7 mm long, pubescent, the 

sepals 1-1.5 mm long, ovate, concave; petals white, 

3.5-5 mm long, lanceolate, abaxially appressed-pubescent, 

adaxially papillate, obtuse at apex, cuneate at base; 

appendage slightly shorter than the petals, triangular, 

adaxially sericeous, abaxially glabrous or papillate; disc 

annular, tomentose; stamens 8, the anthers lanceolate, 

apiculate at apex, glabrous, 1-1.3 mm long, the filaments 

glabrous, of equal or slightly unequal lengths, 1.2-1.4 

mm long; ovary ferruginous-sericeous, the stigma elongated, 

capitate, papillate. Fruit 1-locular, orange-yellow 

at maturity, ellipsoid, glabrescent, granulate, 2-2.5 cm 

long, the pericarp coriaceous, ca. 1 mm thick, the endocarp 

usually glabrous. Seeds ellipsoid, 1 or 2 per fruit. 

Embryo with cotyledons superimposed, the upper cotyledon 


This species is characterized by its sharply triangular 

petioles and lower parts of rachises; distal portion 

of the rachis trullate; essentially glabrous foliage; pet-


Fig. 88. Talisia marleneana. A. leaf. B. flowering branch. C. flower. D. adaxial and abaxial views of petal with 

adnate appendage. E. staminate flower with perianth removed showing disc, stamens, and detail of stamen. F. pistil- 

late flower with perianth removed showing disc, sterile stamens, and pistil (left) and l.s. same showing detail of sterile 

stamen (right). G. portion of infructescence. H. dorsal-lateral view of embryo. (A-F from Acevedo R. et al. 81S0; 

G-H from Prance et al. 24727).


NW.[ 5-w^ SOW -, ?~ 70w 4-W 

Om 

l --.,, f 

Al 

* Tai l,aeXv;gata 

* Talisia marleneana 


als appressed-pubescent on abaxial surface; stamens 8 

with lanceolate anthers; and disc tomentose. The species 

is closely related to T longifolia, however, it differs 

from it by its chartaceous, dull, elliptic or oblong, 

sub-bullate (vs. coriaceous, lustrous, narrow elliptic to 

oblanceolate, flattened) leaflets with abaxially smooth 

surface and prominent tertiary veins (vs. surface with 

scurfy-like punctations and tertiary veins not prominent 

or only slightly so). The specific epithet honors Dra. 

Marlene Freitas da Silva from the Instituto Nacional de 

Pesquisa da Amaz6nia, Brazil. 

Phenology. Flowers from April to November and 

fruits from July to February. 


from the Brazilian states ofAmazonas and Para, in non- 

flooded, moist, terra firme and igapo forests. 

Common names. Brazil: pitomba,pitomba da mata. 


AmapA: Falsino river, ca. 10 km upstream from con- 

fluence with Araguari, 29 Sep 1987 (fl), Pruski et al. 3311 

(NY). Amazonas: Upper Rio Negro, W of Romao, along 

Araca river, 120 m, 9 Aug 1996 (fl), Acevedo R. et al. 

8150 (INPA, US); Pitinga river, 28 Sep 1979 (fr), C. 

Ferreira et al. 888 (INPA, NY); Mun. Presidente 

Figueredo. Balbina, 16 Aug 1986 (fr), Freitas et al. 276 

(INPA); Urubui river, between Iracema falls & Manaus- 

Itacoatiara road, 8 Jun 1968 (fl), Prance et al. 5092 (K, 

NY). Park: Itaituba. Santarem-Cuiaba' road, km 1208, 

Serra Mazi, 18 May 1983 (fl), Amaral et al. 1296 (INPA, 

MG); Altamira, 22 Jul 1971 (fr), Cavalcante & Silva 

2775 (MG); Boa Vista, Tapaj6s river, May-Jun 1929 (fl), 

Dahlgren & Sella 120 (F); Oriximini-Obidos road, 55 

km from Oriximina, 50 m, 5 Jun 1980 (fl), Davidson & 

Martinelli 10073 (INPA, K, MG, NY, US); Faro, 26 Aug 

1907 (fr), Ducke 8507 (MG); Ilha Salgado, castanhal, 24 

Nov 1907 (st), Ducke 8886 (MG); Oriximina. Trombetas 

river, Erepecu lake, 18 Jul 1980 (fr), C. Ferreira et al. 

1637 (INPA, NY), 18 Jul 1980 (fr), C. Ferreira et al. 1655 

(INPA); Porto Trombetas, 29 Aug 1980 (fr), C. Ferreira 

et al. 1892 (INPA, NY); Paru do Oeste river, 10 Sep 1980 

(fr), C. Ferreira et al. 2359 (INPA); Oriximina--6bidos, 

vic. of Cumina-Mirim river, 14 Sep 1980 (fr), C. Ferreira 

et al. 2499 (MG, MO, NY, US); Oriximini-6bidos road, 

55 km from Oriximini, 50 m, 5 Jun 1980 (fl), Martinelli 

6791 (INPA, MG, NY, US); Ilha de Breu, 19 Sep 1965 

(fr), Prance et al. 1355 (IAN, K, NY, US), 27 Sep 1965 

(fr), Prance et al. 1482 (F, IAN, NY, US); Altamira- 

Itaituba road, 31 Oct 1977 (fr), Prance et al. 24727 (MG, 

MO-2, NY, US); Tucurui, 19 Jun 1978 (fl), Rosa 2418 

(MG); Tucurui, Tocantins river, 10 Apr 1978 (fl), Silva 

& Bahia 3478 (MG, NY); Victoria, along Xingui river, 

Jun 1909 (st), Snethiage 10416 (MG); Porto Trombetas, 

20 Jun 1986 (fl), Soares 132 (INPA). 

39. TALISIA MEGAPHYLLA Sagot ex Radlkofer, 

Sitzungsber. Math.-Phys. Cl. Konigl. Bayer Akad. 

Wiss. Miinchen 8: 350. 1878. Type. French Guiana. 

Acarouany, 1857 (yfr), Sagot 1194 (lectotype, P, 

here designated; isolectotypes K-2, P-3; frag. at M). 

Syntypes. French Guiana. without date (st ), Poiteau 

s.n. (K); Surinam. without date (fl), Hostmann 1149 

(K-2, P). Figs. 2c, lOb, 12b, 90a-e 

Slender tree or treelet, 4-10 m tall; trunk to 8 cm in 

diam. Stems sulcate, glabrous, lenticellate, usually hollow, 

inhabited by ants. Leaves paripinnate or imparipinnate,


5mm. 

Fig. 90. Talisia megaphylla. A. leaf and detail of leaflets. B. staminate flower. C. I.s. staminate flower showing 

petals, disc, stamens, and sterile stamens. D. adaxial and lateral views of petal with adnate appendage. E. infructescence. 

(A from Mori & Boom 15138; B-D from Mori et al. 23121; E from (Mori & Boom 15138).


spirlly arranged on distal portions of stem; distal process Common names. Brazil: pitomba. French Guiana: 

acicular, 5-7 mm long; leaflets (7)11-15, opposite or al- bte. Surinm: bosknipa, kraskrastiki; Pemu:pishico huayo. 

ternate, oblanceolate, oblong-elliptic, sometimes obovate Representative specimens examined. VENEZUELA. 

or falcate, chartaceous, (14-) 17-43 x 4.5-10.5 cm, the Amazonas: Rio Negro, s.d. (fl), Schomburgk s.n. (K). 

adaxial surface glabrous, with prominulous midvein and Aragua: Cumboto, Valle de Ucumare, 2 Oct 1947 (fr), 

slightly sunken secondary veins (sometimes bullate), the Pittier 15615 (NY). 

abaxial surface glabrous or sparingly pubescent, primary GUYANA. Without specific locality or date (bd), 

and secondary veins prominent, tertiary veins reticulate, Schomburgk 859 (M). 

prominulous, the apex abruptly acuminate to nearly cau- SURINAM. Brokopondo: Kaboerie, 2 Aug 1922 (fl, 

fr), Boschwezen 5956 (IAN, K-2). Commewijne: Mapana 

date, the base inequilateral, one side obtuse the other acute 

creek, Nov 1957 (fl), Schulz 8073 (NY-2). Marowijne: 

or cuneate, the margins undulate; petiolules browmsh (dry- Moengo, Grote Zwiebelzwamp, 22 Sep 1948 (fr), Lanjouw 

ing darkbrown), pulvinate, adaxially furrowed, 7-14 mm & Lindeman 388 (IAN, K, NY). Nickerie: Zanderij, 12 

long; rachis 20-54 cm long, terete, striate to bicarinate on Aug 1916 (st), Boschwezen 231 (K-3, NY); Sipaliwini. 

distal portion, glabrous, yellowish brown, glossy; peti- Kuruni river, N side of Kuruni island, 180 m, 9 Nov 1994 

oles 20-29 cm long, terete, striate, glabrous, enlarged at (fr), Evans et al. 1924 (P, US); Kabalebo Dam project, 17 

base. Thyrses panicle-shaped, 20-42 cm long, terminal Nov 1976 (fr), Heyde & Lindeman 131 (MO); Lucie river, 

or axillary on distal part of stem, lateral branches to 15 cm lower slopes of Juliana Top, 500-600 m, 14 Aug 1963 

long; cataphylls acicular or pinnate, 1.5-5 cm long, clus- 

(fl), Irwin et al. 54770 (NY, US); Wilhelmina Gebergte, 3 

km S of Juliana Top, 300 m, 24 Aug 1963 (fl), Irwin et al. 

tered at leaves axils; axes angled, sulcate, sericeous-to- 

55046 (NY-2); Kabalebo Dam project, 30-130 m, 20 Sep 

mentose; bracts deltate, sericeous-tomentose, ca. 1 mm 1980 (fr), Lindeman et al. 523 (K, MO, VEN); Corantijne 

long; bracteoles similar to the bracts but smaller; dichasia river, Kaboerie, Sep 1920 (fl), Pulle 526 (MO). 

simple or compound; peduncle 5-8 mm long on proximal Saramacca: Saramacca, Jun 1945 (st), Boschwezen 330 

dichasia, ca. 1 mm on distal dichasia; pedicels 1-1.5 mm (K, NY); 15 Dec 1917 (fr), Boschwezen 3538 (MO). 

long, articulate near the middle, sericeous-tomentose. FRENCH GUIANA. Without specific locality, Apr 

Calyx 4-7 mm long, abaxially ferruginous-sericeous- 1961 (fl), Aubreville 258 (P); 1863 (fl), Melinon s.n (P-3); 

tomentose, the sepals 2-3.5 mm long, ovate to rounded, 4 May 1922 (st), Wachenheim 377 (P). Bassin de 

concave, imbricate, abaxially glabrous around the woolly- 

L'Approuague: Arataye river at Saut Parare, 9 Sep 1983 

(st), Barrier 4228a (CAY); 12 Oct 1983 (st), Feuillet 1046 

pubescent margins, adaxially glabrous, the outer sepals 

(CAY); 70 km SW of Regina, forest plot, 5 Oct 1983 (st), 

shorter than the inner ones; petals white to orangish, 8-9 Villiers 2010 (CAY); 5 Oct 1983 (st), Villiers & Feuillet 

mm long, elliptic, reflexed at anthesis, abaxially ferrugi- 2009 (CAY), 5 Oct 1983 (st), Villiers & Feuillet 2000 (NY). 

nous-sericeous-tomentose except along upper margins, Bassin de L'Inini: Palofini creek, 22 Aug 1970 (fl), de 

adaxially papillate, the margins woolly to ciliate, clawed at Granville C-54 (CAY). Bassin du Maroni: Grand Inini 

base, the claw ca. 3.5 mm long, slightly fleshy, white, gla- river, Degrad Fourmi, 150 m, 8 Aug 1985 (fl), de Granville 

brous; appendage as long as or shorter than the petal, white, et al. 7384 (CAY, P, U); Maroni, 1864 (st), Melinon s.n. 

erect, connivent, strap-shaped or elongated deltate, seri- (US); Acarouany, 30 m, 1855 (st), Sagot s.n. (P); Marouini 

ceous-tomentose on both surfaces; disc 5-lobed, com- river, ca. Maroni river, 23 Apr 1975 (fr), Sastre et al. 3929 

pletely or only apically tomentose, ca. 1 mm tall; stamens 

(CAY, P, US). Bassin du Sinnamary: CTFT plots, 13 Jun 

1985 (fl), Feuillet 2318 (CAY, US). lie de Cayenne: Mont 

8, the filaments white, glabrous, of unequal length, 4-5 

Grand Matoury, 21 Apr 1992 (fr), Acevedo R. & Grimes 

mm long, the anthers oblong-lanceolate, 1.5-1.7 mm long, 4798 (CAY, US), 50 m, 5 Apr 1995 (fr), Cremers et al. 

apiculate at apex; ovary feruginous-tomentose, the stigma 13854 (CAY). Piste de Saint Elie: Station Biologique 

ellipsoid, papillate. Fruit maturing yellow, orange-yellow ORSTOM, 4 May 1992 (st), Acevedo R. et al. 4888 (CAY, 

or red, oblong-ellipsoid to nearly globose, apiculate, 2- US); Piste de St. Elie, 4 Mar 1982 (st), Sabatier 213 (CAY). 

3.2 cm long, velvety pubescent, becoming glabrous at ma- Region Littorale: Charvein, 12 Dec 1913 (fr), Benoist 337 

turity, the pericarp ca. 1.5 mm thick, coriaceous, smooth (P). Region de Saul: Route de Belizon, between Sail & 

or less often granulate. Seed solitary, ellipsoid with lilac, Eaux Claires,19 May 1992 (st), Acevedo R. et al. 4957 

sweet fleshy testa. Embryo with cotyledons diagonally (US); Sail, 220 m, 21 Jun 1988 (st), Gentry et al. 62981 

overlapping, the lower one larger han the upper one. 

(MO-2, NY), 23 Jul 1979 (fl), de Granville 3150 (CAY); 

Monts La Fumee, 200-400 m, 27 Oct 1982 (fr), Mori & 

The specific epithet refers to the large leaflets. 

Boom 15138 (CAY, MO, NY); Mont Galbao trail, 7 Aug 

Phenology. Flowers from June to November and 1987 (fl), Mori & Gracie 18673 (CAY, NY, P); Route de 

in February and April, and fruits from September to Belizon, between Sail & Eaux Claires, 200 m, 1 1 Nov 1990 

December and from February to April. 

(fr), Mori et al. 21608 (CAY, NY); 5 Aug 1993 (fl), Mori 

et al. 23121 (CAY, NY, US); Montagne Boeuf Mort, 10 

Distribution and Ecology. (Fig. 93) Venezuela, the Aug 1971 (fl), Oldeman B-4008 (CAY, P, U, US). 

Guianas, Ecuador, Peru, and Brazil in non-flooded, ECUADOR. Without locality, 4 May 1897 (fr), 

moist, lowland terra firme forest. 

Eggers 15764 (F). Napo: Yasuni Forest Reserve, hill E


of PUCE Scientific Station, 225 m, 19 Jun 1995 (st), 

Acevedo R. & Cedenio 7380 (US); Eosin Forest Reserve, 

28-30 Jul 1993 (fl), Aulestia & Grefa 123 (US). 

PERU. Amazonas: Huambisa, valley of Santiago 

river, ca. 65 km N of Pinglo, 200 m, 19 Dec 1979 (fr), 

Huashikat 1595 (MO); Prov. Bagua. Yamayakat, 320 m, 

25 Feb 1996 (fr), Jaramillo et al. 1306 (US). Cuzco: 

Camisea, vic of community Malvinas, 500 m, 17 Sep 

1997 (st), Acevedo R. et al. 9729 (US, USM). Loreto: 

Tigre river, Nuevo Canaan, along Lamas-Tipishca lake, 

15 Dec 1979 (fr), Ayala et al. 2539 (MO, NY); Alto 

Amazonas, Pastaza river, 210 m, 21 Nov 1980 (fr), 

Vasquez & Jaramillo 825 (MO). 

BRAZIL. Acre: Serra do Canta, Oct 1951 (fl), Black 

51-13951 (U). Amapi: Oiapoque river, 3 km E of Ponta 

dos Indios, 2 Aug 1960 (fl), Irwin et al. 47325 (IAP, MG, 

MO); Iaue river, 3 km E of confluence with Oiapoque 

river, 26 Aug 1960 (fl), Irwin et al. 47853 (IAP, NY, US); 

18 Oct 1960 (fl), Irwin 48803 (IAN); Cachoeira Santa 

Maria, Murure river, 21 Aug 1961 (fr), M. Pires et al. 

50406 (IAN, MG, NY, US). Roraima: Evangelic Mis- 

sion at Paumiu, 2 Mar 1979 (fr), M. Pires et al. 16850 

(INPA, MG); Apiaui river, 28 Jan 1967 (fr), Prance et al. 

4135 (F, MG, NY, US-2); Uraricoeira river, between 

Cutalba creek & Uaica, 24 Feb 1971 (fl, fr), Prance et al. 

10674 (NY, US); Mucajai, 13 Mar 1971 (fr), Prance et 

al. 10920 (K, NY, US); Mucajai river, 17 Mar 1971 (fr), 

Prance et al. 11044 (K, M, MG, NY, US); Ilha de Maraca, 

SEMA ecological reserve, 6 Oct 1987 (fl), Pruski et al. 

3390 (NY), Fuma9a trail, 26 Oct 1988 (fl), Ratter et al. 

5894 (K); Boa Vista, Oct 1908 (fl), Ule 7679 (K, M, MG). 

40. TALISIA MOLLIS Kunth ex Cambessedes, 

Mem. Mus. d'Hist. Nat. Paris 18: 48. 1829. Type. 

French Guiana. Cayenne, 1820 (fl), Anonymous 347 

(holotype, B-destroyed, photo (F neg. 5677) at US). 

Epitype. French Guiana. Saul: Along Limonade trail, 

by km. 1, 30 Oct 1971 (fl), Oldeman B-4183 

(epitype, CAY-2, here designated; iso-epitypes, P, 

US). Figs. la, 13a&c, 91a-k 

Tapirocarpus talisia Sagot, Ann. Sci. Nat. Ser. VI, 

13: 292. 1882, pro parte vegetativa. Type. French 

Guiana. Acarouany, 1857 (st), Sagot s.n. (lecto- 

type, P, here designated; isolectotype, P). 

Talisia guianensis sensu de Candolle, Prodr. 1: 609. 

1824, non Aublet, 1775. 

Slender tree or treelet, 4-10(13) m tall, unbranched 

or with few ascending, sympodial branches on distal 

portion; trunk to 8 cm in diam.; bark grayish to dark 

brown, smooth or lenticellate. Stems terete, minutely 

hirsute, sometimes hollow in the center and inhabited by 

ants. Leaves paripinnate, spirally arranged on distal portion 

of stem(s); distal process acicular, early deciduous, usually 

curved; leaflets chartaceous, 10-16 (32), opposite 

to altemate, oblong, elliptic or lanceolate, (5.5) 9-27(53) 

x (2)3.5-8.5(12) cm, the venationbrochidodromus, tertiary 

veins reticulate, the adaxial surface with impressed 

venation, glabrous or the midvein minutely tomentose, 

the abaxial surface minutely hirsute, with prominent venation, 

the apex acuminate to caudate, the base 

inequilateral, one side obtuse or rounded the other attenuate 

or cuneate, the margins entire slightly revolute; 

petiolules pulvinate, 4-6 mm long, minutely hirsute to 

glabrescent; rachis 22-60 cm long, terete, or slightly 

angled on distal portion, minutely hirsute; petioles 12- 

27 (40) cm long, terete, minutely hirsute, greatly enlarged 

at base. Thyrses panicle-shaped, 30-60 cm long, terminal 

or axillary on distal part of branches; cataphylls acicular, 

5-10 mm long, hirsute, axes terete or slightly 

angled, hirsute; bracts and bracteoles lanceolate or subulate, 

hirsute, 1.3-3.5 mm long; dichasia simple, sessile; 

pedicels < 1 mm long, articulate at the apex. Calyx light 

green, urceolate, 2.3-4 mm long, minutely hirsute, sometimes 

intermingled with glandular hairs, the sepals 1.6- 

3 mm long, ovate-deltate; petals white, 5.5 mm long, 

oblong, abaxially glabrous, adaxially papillate, rounded 

at apex, cuneate or cuneate-attenuate at base; appendage 

as long as the petal, long-deltate, abaxially papillate, 

adaxially hirsute on upper half; disc 5-lobed, minutely 

hirsute at apex, ca. 0.8 mm tall; stamens 8, the filaments 

of equal or nearly equal length, glabrous, 2-4 mm long, 

nearly filiform, the anthers oblong or linear-lanceolate, 

1.5-1.8 mm long, apiculate at apex; ovary conical, sericeous-hirsute, 

the stigma elongate-trigonous, ferruginous-papillate. 

Fruit yellow to orange-yellow at maturity, 

ovoid to globose, 2-2.5 cm long, densely to sparsely 

hirsutulous, the pericarp woody, smooth, to 2.5 mm thick. 

Seed ellipsoid, the testa woody, with a thin, fleshy, yellowish 

to orange-yellow upper layer, rugulate when 

dried. Embryo with cotyledons lying dorsiventrally to 

obliquely dorsiventrally over each other, the lower one 

almost twice as large as the upper one. 

Specimens of Talisia mollis are sometimes difficult 

to tell apart from those of T hemidasya because many 

of the characters may overlap. In general, T mollis has 

stems that are more densely pubescent than those of T 

hemidasya, and which lack glandular hairs. The abaxial 

side of leaflets in T. mollis are always minutely 

hispidulous, while those of T hemidasya are glabrous 

or puberulent, only rarely minutely hispidulous. These 

distinctions may sound weak in the absence of the habit 

character which is clearly different in both species. 

Talisia mollis is an unbranched, palm-like understory 

shrub while T hemidasya is a large tree with profuse 

branching which forms a dense crown. Talisia mollis 

is vegetatively similar to T pachycarpa; however, they 

are easily distinguished by their flower morphology. 

The calyx in T mollis has sepals that are free nearly to 

the base, while those of T pachycarpa are connate to 

form a cupular calyx. The specific epithet refers to the 

soft pubescence of this species. 

The holotype of T mollis at B was destroyed dur-


8 cm1 

3 cmI.' 

cmg4 __ 

3mm[ 

Fig. 91. Talisia mollis. A. leaf. B. detail of leaflets. C. portion of inflorescence. D. flower bud and detail of tri- 

chomes. E. pistillate flower (left) and l.s. of same (right). F. lateral and adaxial views of petal with adnate appendage. 

G. pistillate flower with perianth removed showing disc, sterile stamens, and pistil. H. sterile stamen. I. fruiting branch. 

J. two-seeded fruit, I.s. K. seeds. (From Mori et al., 2003. Guide to the Vascular Plants of Ce ntral french Guiana.)


ing World War II. The only surviving material of this de Granville B-4222 (CAY-2, P, US); New road to vilcollection 

appears to be a fragment in the Cambessedes lage water supply, 260 m, 25 May 1986 (fr), Mori & 

herbarium at MPU. Represented only by an inflorescence 

fragment, this collection is not an adequate specimen 

to typify the species. For this reason, I am choosing 

a recent collection as epitype for T mollis. 

Pennington 18140 (CAY, K, US); Route de Belizon, between 

Eaux Claires & entrance to Grand Boeuf Mort, 

250 m, 4 Jan 1996 (fl), Mori & Pepper 24260 (CAY, 

US); Route de Belizon, between Saul & Eaux Claires, 

200-300 m, 11 Nov 1990 (fl), Mori et al. 21607 (CAY, 

Phenology. Flowers from September to January and 

fruits from November to July. 

NY, US); Cochon creek valley, along Belvedere trail, 2 

Sep 1971 (fl), Oldeman B-4094 (CAY-4, NY, P). 

BRAZIL. AmapA: Mazagao, 21 Dec 1984 (fr), Daly 

Distribution and Ecology. (Fig. 93) The Guianas et al. 3951 (MG, NY); Mun. Oiapoque. Oiapoque river, 

and Brazil in understory of non-flooded forest. 12 Oct 1960 (fl), Irwin 48680 (IAN, M, MG, NY, US-2); 

BRi56, between Cal9oene & Oiapoque, by Ca9ipore 

Common names. Brazil: pitomba. French river, 1 Jan 1985 (fr), Mori & Souza 17301 (MG, US); 

Guiana: payacoussa, paiacoussa, tuliata, flambeau Ariramba, 26 Jun 1982 (fr), Rosa 4416 (INPA, MG, NY); 

dur, tatu-tatu. 

Jarn river, between Monte Dourado & Munguba, 13 Dec 


Without specific locality. Basin of Essequibo, 212 m, 1 

Oct 1952 (st), Forest Dept. of British Guiana 7305 (NY). 

SURINAM. Brokopondo: Brownsweg, 15 Aug 1924 

(fl), Boschwezen 6575 (US), 1 Jul 1924 (fr), Boschwezen 

6595 (IAN, K, MO, NY); Jodensavanne Mapan, Nov 

1960 (yfr), Schulz 8387 (K, NY). 

FRENCH GUIANA. Without specific locality, 1792 

(fl), Leblond 448 (P-2), 1864 (fl), Melinon s.n. (P-2, US). 

Bassin de L'Approuague: Arataye river at Saut Parare, 8 

Sep 1983 (st), Barrier 4200 (CAY). Bassin de la Comte: 

Route de Belizon, Montagne Tortue, 23 Apr 1992 (fr), 

Acevedo R. et al. 4823 (CAY, US). Bassin de la Mana: 

Montagnes de la Trinite, 400 m, 12 Jan 1984 (fr), de 

Granville et al. 5903 (CAY-3, U, US). Bassin du Maroni: 

Maripasoula, 12 Mar 1986 (st), Fleury 100 (CAY-2, US); 

Maroni, 1864 (fl), Melinon s.n. (NY, P, US-2), 1862 (fr), 

Melinon s.n. (P-2); Camopi river, Saut Ouasseye, 13 Dec 

1967 (fl), Oldeman 2666 (CAY-4); E of Montagne 

1968 (fl), Silva 1542 (F, K, IAN, NY, US). Maranhao: 

Along road Br 222, between Santa Ines e A9ailandia, 300 

m, 17 Dec 1978 (fr), Jangoux & Bahia 562 (MG, NY). 

Pari: Serra dos Carajas, 15 Oct 1977 (fr), Berg et al. 

548 (MG, MO-2, NY, US); Obidos, Col6nia Curucamba, 

27 Dec 1904 (fr), Ducke 6945 (MG-2); Castanhal, 24 

Nov 1907 (fl), Ducke 8887 (MG);12 Dec 1906 (fl), Ducke 

7940 (MG); Trombetas river, 18 Jul 1980 (fr), C. Ferreira 

et al. 1660 (INPA, NY); Ilha de Breu, 5 Oct 1965 (fl), 

Prance et al. 1544 (F, IAN, M, NY, US); Tucurui, 8 Apr 

1981 (fr), Rosa et al. 4083 (MG); Almeririm, Monte 

Dourado, 28 Nov 1978 (fl), M. R. Santos 431 (NY); 

Bujaru, 25 Apr 1982 (fr), Silva 5936 (INPA, MG, NY); 

Tucurui, 2 Jun 1980 (fl), Silva & Rosario 5328 (MG, 

NY); Gari river, between Planalto and Braxo, 22 Jan 1969 

(fl), Silva 1665 (F, IAN, NY); Tucurui, trail side, 17 Dec 

1979 (fl), da Silva et al. 445 (INPA, MG); Altamira, 13 

Oct 1986 (fl), Vasconcelos et al. 255 (MG, NY). 

Yanioue, 8 Feb 1968 (st), Oldeman & Sastre 104 (CAY- 

2, IAN, NY, P, VEN), 8 Feb 1968 (fr), Oldeman & Sastre 

41. TALISIA MORII Acevedo-Rodriguez, sp. nov. 

111 (CAY-3, P). Bassin de L'Oyapock: Trois Sauts, Type. Panama. Darien: North slopes of Cerro Pirre; 

Kanikani creek, 25 May 1974 (fr), Grenand 364 (CAY); wet forest, 300-700 m, 4 Apr 1975 (fl), Mori, S.A. & J. 

Zidock village, 20 May 1974 (st), Grenand 243 (CAY), 

31 Oct 1974 (fl), Lescure 380 (CAY). Bassin du 

Sinnamary: Sinnamary river at intersection with 

Marouina creek, 14 Feb 1979 (fr), Cremers 5434 (CAY- 

Kallunki 5382 (holotype, MO). Figs. 18a-c, 92a-h 

A T. croatii Acevedo-Rodriguez inflorescentiis 

dense glanduliferis, et discis 5-glandulosis differt. 

2, P); Plomb creek, 11 Jul 92 (fr), Loubry 1894 (CAY, Tree 4-5 m tall. Stems nearly terete, ferruginous- 

NY, U, US); Gregoire creek, 16 Jul 1970 (fr), Oldeman 

tomentulose, glabrescent and lenticellate with age. 

B-3481 (CAY-2, P, US). Ile de Cayenne: Mont Grand 

Matoury, 13 Jul 2000 (st), Acevedo R. 11125 

Leaves paripinnate; distal process acicular, ca. 3 mm, 

(CAY, US). 

Piste de Saint Elie: Station Biologique ORSTOM, 27 deciduous; leaflets 10-14, opposite when distal, alter- 

Apr 1992 (fr), Acevedo R. et al. 4844 (CAY, US), 2 May nate or subopposite when proximal, oblong, oblong- 

1992 (fr), Acevedo R. & Grimes 4877 (CAY, US); Piste elliptic or oblanceolate, 1 1-40 x 3.5-12 cm, chartaceous, 

de St. Elie, 10 Sep 1981 (st), Halle 2773 (CAY), km 14, the adaxial surface glabrous except for the puberulent 

23 Oct 1980 (fl), Prevost 1079 (CAY-2, P), km 14.8, 21 midvein, the abaxial surface flavo-pilosulous along 

May 1981 (fr), Prevost 1109 (CAY), 29 Jul 1991 (st), 

Sabatier & Prevost 3676 (CAY). Region Littorale: 

Kourou, 6 May 1992 (fr), Acevedo R. et al. 4914 (CAY, 

US). Region de Saul: Route de Belizon, between Saul 

& Eaux Claires, 160 m, 19 May 1992 (fr), Acevedo R. 

et al. 4953 (CAY, NY, US); Saul. Without specific locality, 

220 m, 22 Jun 1988 (st), Gentry et al. 63091 (MO); 

Limonade creek, 300 m from village, 13 Jan 1972 (fr), 

veins, intermixed with scattered glandular hairs, the 

venation brochidodromus, plane on adaxial surface, 

prominent on abaxial surface, tertiary venation clathrate, 

the margins entire, slightly undulate, the apex longacuminate, 

usually abruptly so, the base asymmetrical, 

one side acute the other obtuse; petiolules nearly cylindrical 

to pulvinate, slightly compressed along adaxial


I 'I''~~~~~~~~~~~~~~~~~~~~~7 

1.. ~ ~ ~ I 

2mmm.~ f 

Fig. 92. Talisia morii. A. flowering branch. B. section of inflorescence branch showing flower. C. staminate 

flower. D. abaxial and lateral views of petal with adnate appendage. E. staminate flower with perianth removed 

showing disc and stamens. F. detail of disc lobe and subtending stamen. G. detail of leaflets. H. fruit with mesocarp 

broken away showing portion of seed. (A-F from Herrera et al. 981; G-H from Romero C. 5049).


_w rx 70WI oiwlV 4W 

~~~~~t4Z I~~~~~~~~~~~ 

* Talisia megaphylla / A 

* Talisia mori 

A Talisia sollis > a 


surface, 0.5-1 cm long, flavo-pilose; rachis terete, pubescence. Seed one per fruit, ovoid-ellipsoid, ca. 2 cm 

flavo-pilosulous with scattered glandular hairs, 24-34 long. Embryo with cotyledons superimposed, the upcm 

long; petioles 17-20 cm long, terete, striate, flavo- per one slightly larger than the lower one. 

pilosulous, thickened at base. Thyrses panicle-shaped, Talisia morii is distinguished from all other Talisia 

compound, terminal, pyramidal, to 25 cm long; by its densely glandular-pubescent inflorescence axes 

cataphylls acicular or dendroid, tomentose, numerous, and its nectary disc with 5 prominent, tooth-shaped lobes. 

congested at base of inflorescence or supra-axillary, Talisia morii seems to be closely related to T croatii; 

reaching 4 cm long; axes sulcate, scattered pilosulous however, it can be distinguished from the latter by its 

to puberulent, densely glandular-pubescent; bracts subu- glandular-pubescent inflorescences axes (vs. puberulent 

late, puberulent, glandular-pubescent, early deciduous; to tomentose without glandular hairs) and by the deeply 

bracteoles deltate-subulate, 0.5-1 mm long, persistent, 5-lobed or 5-glandular disc (vs. disc shallowly 5-lobed). 

pilosulous, glandular-pubescent; dichasia compound, The specific epithet honors Dr. Scott A. Mori, Senior 

simple or the flowers solitary (due to aborted lateral Scientist at the New York Botanical Garden, collector 

flowers) along secondary branches; peduncle 1-7 mm of the type, and eminent plant systematist. 

long; pedicels 1-3 mm long, pilosulous, with scattered 

glandular hairs, articulate at the middle. Calyx 3-4.5 

mm long, sparsely tomentulose with scattered glandular 

hairs, the sepals 1.5-3 mm long, ovate or oblong, concave, 

rounded at apex, ciliolate; petals ovate, ca. 7 mm 

Phenology. Collected in flower during April and 

May and in fruit during October. 


from Panama and Colombia, in moist, lowland forest. 

long, reflexed at anthesis, adaxially glabrous, abaxially Representative specimens examined. PANAMA. 

tomentulose on lower half, the apex rounded, the base Darien: Parque Nacional Darien, along trail from Casa 

clawed, the margins ciliate on lower half; appendages as Vieja to Cerro Sapo, 180-500 m, 23 May 1991 (fl), 

long as the petals, erect, oblong, adaxially ferruginous- Herrera et al. 981 (US). 

sericeous-tomentose, abaxially sparsely sericeous-tomen- COLOMBIA. Antioquia: Mun. San Luis, Fca. Las 

tose; disc 5-lobed, tomentulose, the lobes tooth-shaped, 

Confusas, 350-500 m, 11 Apr 1990 (fl), Cardenas et al. 

2700 (JAUM). Santander: Ca. 80.2 km SE of Barranca 

ca. 1.6 mm tall; stamens 8, the filaments densely pilose, 

Bermeja, 3 km from the left side of Op6n river, ca. 

of unequal lengths, 3-4 mm long, the anthers oblong- 

200 m, 14 Oct 1954 (fr), Romero C. 5049 (US-2). 

elliptic, ca. 2 mm long, glabrous, apiculate at apex. Pistillate 

flowers unknown. Fruits ellipsoid, ca. 3 cm long, 

glabrous, granulate, pericarp slightly woody, ca. 2 mm 42. TALISIA NERVOSA Radlkofer, Smithsonian 

thick, endocarp nearly glabrous, with scattered woolly Misc. Collect. 61(24): 4. 1914. Type. Panama. Prov. 

2Qh 

/1Q


Colon: Loma de la Gloria, back of Fato, 23 Aug late at the middle, with the same indumentum as the axis. 

191 1 (fr), H. Pittier 4249 (holotype, US; frag. at M). Flowers buds angular-ovoid. Calyx 2-3 mm long, light 

Figs. 6a, 94a-e green, puberulent to tomentose, the sepals 1-1.2 mm 

Talisia grandifolia Cuatrecasas, Rev. Acad. Colomb. long, oblong-rounded or ovate, slightly concave; pet- 

Cienc. 8: 306. 1951. Type. Colombia. Valle: Costa als white, 4.5-7 mm long, lanceolate or less often obdel 

Pacifico, Cajambre river, Barco, 5-80 m, long, glabrous or adaxially papillate, obtuse or rounded 

21-30 Apr 1944 (fl), Cuatrecasas 17010 (holoat 

apex, cuneate at base, the margins usually ciliate on 

type, F; isotype, US). 

lower portion; appendage as long as the petal or slightly 

Talisia tirirensis Steyermark & Maguire, Mem. New 

York Bot. Gard. 17: 448. 1967. Type. Venezuela. shorter, long deltate, abaxially papillate or less often 

Bolivar: Tirica river, above Techine-meru6, 470 m, glabrous, adaxially hirsute above the base; disc 5-lobed, 

16 Jan 1955 (fl), Steyermark & Wurdack 116 puberulent, tomentose or sparsely hirsute, 0.7-1 mm 

(holotype, NY; isotype, MO). 

tall; stamens 5, the filaments of equal length, pilose to 

Talisia dwyeri Croat, Ann. Missouri Bot. Gard. 63: 528. glabrous, 2.2-2.5 mm long, slightly flattened, the an- 

1976. Type. Panama. Dari6n: NE of Cerro Azul, thers oblong to lanceolate, 1.3-2.2 mm long, apiculate 

premontane rain forest, 10 Dec 1974 (fl), Mori at apex; ovary tomentose, conical, the stigma elongate- 

& Kallunki 3652 (holotype, MO; isotype, PMA). capitate, ferruginous-papillate. Fruits yellow to orange- 

Talisia amaruyana Steyermark, Ann. Missouri Bot. 

yellow at maturity, ellipsoid or ovoid-ellipsoid, apicu- 


late, 1.8-3 cm long, glabrescent, minutely rugulate, the 

Amaruay-tepui, 550-800 m, 5?55'N, 62?15'W, 

20 May 1986 (fl), Liesner & Holst 20394 (holo- pericarp woody, 1-3 mm thick, the endocarp smooth, 

type, MO-4 sheets); isotype, VEN). 

sparsely tuberculate and sparsely appressed- or woollypubescent. 

Seed 1(2) per fruit, ellipsoid, the testa woody, 

Treelet or tree, with sympodial branching, 2-8 (30) m with a thin, fleshy, upper layer. Embryo with cotyletall; 

truk to 8 (35) cm in diam.; bark grayish to dark brown, dons lying dorsiventrally to obliquely dorsiventrally 

smooth or lenticellate. Stems nearly terete, striate, gla- over each other, both cotyledons of similar size or the 

brous, becoming lenticellate, sometimes hollow in the lower one larger than the upper one. 

center and inhabited by ants. Leaves paripinnate, spi- Talisia nervosa can be distinguished by its flowers 

rally arranged on distal portion of stem(s); distal pro- (5-stamens, pilose filaments and puberulent to 

cess woody, truncate, early deciduous, 2-4 mm long; hispidulose disc), fruits with pubescent endocarp and 

leaflets (2) 8-16 (32), opposite or altemate, oblong, leaflets with strong intramarginal veins. The specific 

narrow-oblong, elliptic, narrow-elliptic, obovate, oblan- epithet refers to the conspicuous venation of leaflets. 

ceolate or less often falcate, chartaceous to coriaceous, 

Phenology. Flowers from October to May, and 

14-36 (-61) x 3.5-10.4 (-21) cm, flattened or less often 

fruits throughout the year. 

bullate, the adaxial surface glabrous, with prominent 

midvein but impressed secondaries and tertiaries, the Distribution and Ecology. (Fig. 98) From Nicaraabaxial 

surface glabrous or less often puberulent, with gua south to Colombia, Venezuela, Ecuador, Peru, and 

prominent brochidodromus venation, the secondary French Guiana, in understory of wet, non-flooded forest. 

veins altemate, forming a prominent intramarginal vein, Common name. Peru: Juapina. 

tertiary venation reticulate, the apex long-acuminate or 

Representative specimens examined. NICARAGUA. 

less often obtuse, the base asymmetrical to very strongly 

Rio San Juan: Bocas de Sabalo, 70-100 m, 14 Mar 

asymmetrical, one side obtuse the other cuneate or at- 

1987 (st), Moreno 26743 (MO-2). Zelaya: El Achote, 

tenuate, the margins entire or undulate, revolute or slightly 200 m, 25 Aug 1982 (fr), Araquistain 3137 (MO); Beso; 

petiolules enlarged, cylindrical or bulbous (longer tween Nueva Guinea and Verdun, 240 m, 17 Aug 1983 

than wider), 7-12 mm long, glabrous or puberulous, (fr), Miller & Sandino 1117 (MO); Buena Vista, 100adaxially 

canaliculate; rachis glabrous or puberulous, 150 m, 23 May 1984 (st), Robleto 621 (MO); El Zapote, 

(7.5-) 21-41 cm long, slightly flattened dorsiventrally, 40 km NE of Nueva Guinea, 25-31 Mar 1984 (fl), 

sharply angled toward the margins on distal portion, Sandino 4898 (MO). 

nearly terete on proximal portion; petioles 14-41 (56) COSTA RICA. Heredia: La Selva, OTS Field Stacm 

long, terete, striate, glabrous, less often lenticellate, tion, forest, 100 m, 5 Apr 1982 (fl), 

enlarged at base. Thyrses panicle-shaped, 30-60 cm 

Hammel & Schatz 11578 (F). Puntarenas: Finca La 

Lola, Apr 1949 (fl), Holdridge 2537 (US-2); Parque 

long, terminal or axillary on distal part of branches; 

Llorona Forest, 0-150 m, 16 Jan 1989 (fr), Kernan & 

cataphylls acicular, 5-15 mm long, appressed-pubes- 

Phillips 905 (MO); Peninsula de OSA, Aguabuena, 3 

cent; axes slightly angled, striate or sulcate, puberulent km W of Rinc6n, 100 m, 4 May 1993 (fl), Thomsen 360 

to tomentose; bracts and bracteoles deltate, puberulent (C, US), 2 km W of Rinc6n, 200 m, 8 Mar 1995 (fl), 

to tomentose, ca. 0.5 mm long; dichasia compound; Thomsen 1279 (C); Uvita, 100-200 m, 14 Apr 1988 (fl), 

peduncle 1-2 mm long; pedicels < 1 mm long, articu- Zamora et al. 1488 (MO).


D. ~~ ]3mm. 

Fig. 94. Talisia nervosa. A. portion of leaf. B. portion of inflorescence. C. dichasium. D. adaxial view of petal 

and adnate appendage. E. staminate flower with perianth removed showing disc and stamens (right) and l.s. same 

with detail of anther (left). (All from Cuatrecasas 17010).


PANAMA. Canal Zone: Barro Colorado Island, 

Armour trail, 305 m, 14 Jun 1970 (fr), Croat 10873 (MO); 

Balboa trail, 550 m, 20 Mar 1969 (fl), Croat 8800 (MO- 

2); Lutz trail, 200 m, 27 Sep 1968 (fr), Croat 6498 (MO, 

US); Shannon trail, 500 m, 27 Feb 1969 (bd), Croat 8236 

(F, MO), 5 Jul 1971 (fr), Croat 15251 (MO-2); Snyder 

Molino trail, 250 m, 22 Feb 1969 (fl), Croat 8095 (MO); 

Wheeler trail, 20 Mar 1969 (fl), Croat 8782 (MO-2); Zetek 

trail, 22 Mar 1971 (st), Croat 14027 (MO-2, US-2); 29 

Mar 1970 (fl, yfr), D'Arcy 3916 (F, MO); Fort Sherman, 

Dec 1965 (st), Dwyer 6940 (MO); Apr 1965 (fl), Dwyer 

& Robyns 140 (MO, NY); 1 Jul 1960 (fr), Ebinger 240 

(US); Balboa trail, 1960 (fr), Ebinger s.n. (MO); 4 mi. N 

of Gamboa; Pipeline Road, 18 Dec 1971 (st), Gentry 3165 

(MO); Barro Colorado Island, s.d. (fl), Hladick 361 (MO- 

2); Gatun Lake, 6 Aug 1955 (fr), Johnston 1557 (MO); 

Lim6n Bay, Gatun Locks & Gatun Lake, 3 Apr 1956 (fr), 

Johnston 1778 (MO); Lutz trail, 21 Jul 1927 (st), Kenoyer 

426 (US); Margarita swamp, 15 Jun 1923 (fr), Maxon & 

Valentine 7044 (US-2); Frijoles, 19 Dec 1923 (fl), 

Standley 27575 (US); 17 Jan 1924 (st), Standley 31421 

(US), Jul 1931 (st), Starry 260 (MO); Shannon trail, 20 

Feb 1932 (fl), Woodworth & Vestal 631 (MO); 17 Feb 

1939 (fl), Zetek 4335 (F, MO-2, US-2). Col6n: East Santa 

Rita Ridge, 16 Jan 1968 (fl), Correa & Dressler 629 (MO- 

2); East Ridge, 23 Feb 1968 (fr), Duke 15250 (MO, US); 

Santa Rita Ridge, E of Pan-American hwy, 300-500 m, 

16 Dec 1972 (fl), Gentry 6603 (MO-2); 22-25 km from 

Pan-American Hwy, 400-500 m, 21 Oct 1981 (fl), Knapp 

et al. 1703 (MO); Santa Rita Ridge, 500 m, 11 Jan 1987 

(fl), McPherson 10261 (NY-2); Santa Rita Ridge road, 

17.1 km from B-R highway, 21 May 1975 (fr), Mori & 

Crosby 6318 (MO-2). Darien: Between Sasardi & Morti, 

ca. 400 m, 14 Feb 1967 (fr), Duke 10024 (MO, US). 

Panama: Cerro Jefe, ca. 850 m, 1 Jan 1972 (fl), Gentry 

& Dwyer 3492 (MO); Between Cerro Azul-Cerro Jefe, 

km 22 Pan-American hwy., 11 Jun 1975 (fr), Mori et al. 

6537 (MO); Llano-Carti road, km 8-11 Pan-American 

hwy., 300-400 m, 13 Aug 1975 (fr), Mori 7738 (MO-2); 

Cerro Azul to Cerro Jefe, 9 Jul 1966 (fr), Tyson et al. 4336 

(MO). San Blas: Veraguas. Ensenada Santa Cruz. Coiba 

Island, woods at airport, 16 Aug 1961 (fr), Dwyer 1548 

(MO-2), 7 Aug 1962 (fr), Dwyer 2332 (MO, US), 27 Aug 

1970 (fr), Foster 1619 (F, MO); Nusigandi-El Llano Carti 

road, 300-350 m, 27 Mar 1987 (fl), McPherson 10761 

(NY-2); Camp Nusagandi, km 19.1 on Pan-American 

hwy., 350 m, 19 Mar 1993 (fl), Paredes 953 (US). 

COLOMBIA. Bolivar: Rio Viejo; Norosi, Norosi- 

Tiquisionuevo road, 200 m, 9-14 Apr 1985 (fr), 

Cuadros 2176 (COL); La Raya, Quebrada La Culebra, 

between junction of Cauca & Magdalena rivers, 80-110 

m, 5 May 1987 (st), Gentry & Cuadros 57395 (MO); 

Between Monte Libano & San Pedro, 28 May 1949 (fr), 

Romero C. 1792 (COL). Choc6: Jequed6, 42 km W of 

Las Animas, E of Pato river on Pan American hwy., 250 

m, 11 Jan 1979 (fr), Gentry & Renteria 23991 (MO-2, 

NY); Quibd6-Tutunendo road, 14 km NE of Quibd6, 

trail to Tubad6, 90 m, 17 Jan 1979 (st), Gentry & 

Renteria 24291 (MO). Valle: Bajo Calima Concession, 

ca. 20 km NW of Buenaventura, 50 m, 2 Aug 1988 (st), 

Faber-Langendoen & Hurtado 1934 (MO). 

VENEZUELA. Amazonas: Guainia, along road 

from Maroa to Yavita, ca. 700 m from Yavita, 20 Feb 

1998 (st), Acevedo R. et al. 10286 (US); Atabapo, Cerro 

Huachamacari, E slope, 600-700 m, 3 Nov 1988 (fr), 

Liesner 25739 (US); Cerro de la Neblina, Yatua river, 

700 m, 2 Jan 1958 (fl), Maguire et al. 42590 (K-2, MG, 

MO-2, NY-2, US-3); Casiquiare river, 5 May 1942 (fr), 

Williams 15174 (US-2); Cuao river, 350-400 m, 27 Jul 

1986 (fr), Zent 586-47b (MO). Bolivar: Piar, 550-800 

m, 20 May 1986 (fr), Liesner et al. 20935 (MO-2, NY, 

US); Chimanta-tepui, 1000-1400 m, 15 May 1953 (fr), 

Steyermark 75374 (F, NY-2). 

FRENCH GUIANA. Bassin du Maroni: St. Jean, 

20 Nov 1914 (fl), Benoist 789 (P). 

ECUADOR. Carchi: Tulcan. San Marcos, 750 m, 

20-30 Apr 1993 (fl), Mendez et al. 248 (MO). 

Esmeraldas: Cayapa river, Zapallo Grande, 150 m, 3 

Aug 1982 (fr), Kvist & Asanza 40820 (QCA). 

PERU. Huanuco: Tingo Maria, Rio Subte Grande, 

680 m, 2 Mar 1945 (st), Burgos 10 (F). Loreto: Corrientes 

river at the Ecuador border, between T. L6pez and Puesto 

Avanzado, 280-350 m, 4 Apr 1977 (fr), Gentry et al. 

19029 (F, MO); Prov. Maynas. across Nanay river from 

Puerto Almendras, 130 m, 3 Mar 1979 (st), Gentry et al. 

25360 (MO); Mishana, along Nanay river, l/2 way between 

Iquitos and Santa Maria de Nanay, 130 m, 1979 

(fr), Ramirez 31 (F, MO); Estaci6n Biol6gica Rio Blanco, 

150 m, 16 Sep 1985 (fr), Vasquez et al. 6741 (MO); 

BRAZIL. Amazonas: Rio Negro, Sao Felipe, 20 Apr 

1947 (fr), Froes 22187 (IAN); Serra do Ferro, 6 Mar 

1975 (fr), Silva 3908 (IAN). 

43. TALISIA OBOVATA A.C. Smith, Brittonia 2: 

154. 1936. Type. Brazil. Amazonas: Near mouth of 

Embira river, basin of Jurua river, 17 Jun 1933 (fl), 

B.A. Krukoff 4981 (holotype, NY-2 sheets). 

Fig. 95a-g 

Treelet 3-4 m tall. Stems terete, glabrous or puberulent, 

lenticellate, striate. Leaves paripinnate; distal process 

acicular, ca. 2 mm long; leaflets 4 or 6, opposite, 

obovate, oblanceolate or elliptic, chartaceous, 5.5-22 x 

2.5-8.5 (15.2) cm, glabrous, the adaxial with prominent 

midvein and slightly impressed secondaries, the abaxial 

surface with prominent brochidodromus venation, the 

secondary veins straight, forming a strong submarginal 

loop, tertiary veins reticulate, the apex obtuse to abruptly 

acuminate, the base asymmetrical, attenuate or obtuse, 

the margins entire, slightly revolute; petiolules 5-12 mm 

long, pulvinate at base, glabrous, adaxially canaliculate; 

rachis 6-8 cm long, terete or nearly so, glabrous; petioles 

5-12.5 cm long, terete, glabrous, less often 

lenticellate, bulbous at base. Thyrses panicle-shaped, 

8-20 cm long, terminal or axillary on distal portion of 

branches; cataphylls acicular, 2-4 mm long, appressedpubescent, 

supra-axillary; axes slightly angled, striate 

or sulcate, puberulent; bracts and bracteoles subulate, 

pubescent, ca. 1 mm long; dichasia compound or simple


I~~~~~~~~~~~~~~~~~~~~~~~~~ m 

Fig. 95. Talisia obovata. A. flowering branch. B. staminate flower. C. abaxial and adaxial views of petal with 

adnate appendage. D. staminate flower with perianth removed showing disc, stamens, and pistillode (left) and i.s. 

same (right). E. stamens. F. pistillode. G. portion of leaf with narrow leaflets. (A-F from Krukof 4981; G from 

Prance 14330).


toward distal portion of axis; peduncle less than 1 mm INPA, MG, NY), Carreira et al. 526 (INPA, MG, NY); 

long; pedicels 1-1.5 mm long, articulate above the Vic. of Cachoeira Santo Antonio, Curuquete river, 14 Jul 

middle. Flower buds angular-ovoid. Calyx 1.5-2.5 mm 1971 (fl), Prance 14330 (INPA, MG, MO, NY). 

long, light green, pubescent to pilose, the sepals 1-1.5 

mm long, deltate or ovate-deltate, concave, adaxially 

glabrous; petals white, 4.5-5 mm long, oblong-lan- 

44. TALISIA OEDIPODA Radlkofer, Sitzungsber. 

ceolate or lanceolate, glabrous except for the ciliate lower Math.-Phys. Cl. Konigl. Bayer Akad. Wiss. 

margin, obtuse at apex, cuneate at base; appendage as Miinchen 8: 347. 1878. Type. Brazil. Mato Grosso 

long as the petal, long deltate, abaxially glabrous or do Sul: Rio Pardo at open, dry, sandy field, 1824sparsely 

papillate, adaxially hirsute on upper half; disc 1829 (fl), Riedel 522 (holotype, LE?; isotypes, NY- 

5-lobed, glabrous, 0.54.7 mm tall; stamens 5, the fila- 2, US; frag. at M). Fig. 96a-d. 

ments of equal length, glabrous, 1.2-2.2 mm long, Shrub. Stems slightly sulcate, glabrous, vemicose, 

slightly flattened, the anthers oblong, 1.2-2 mm long, mature stems lenticellate. Leaves paripinnate; distal proapiculate 

at apex; ovary glabrous or puberulent, conical, cess acicular, ca. 5 mm long, early deciduous leaving a 

the stigma elongate-trigonous, ferruginous-papillate. truncate base 2-3 mm long; leaflets (2-) 6 (8), opposite, 

Fruit ovoid or ellipsoid, ca. 2 cm long, yellow, the perielliptic, 

5.5-12 x 1.2-4 cm, coriaceous, the blade invocarp 

coriaceous, granulose, ca. 0.5 mm thick, the en- lute, glabrous, the adaxial surface shiny, the abaxial surdocarp 

glabrous. Embryo with cotyledons lying trans- face dull, the venation brochidodromous, prominent on 

versely over each other, both of similar size. both surfaces, tertiary veins reticulate, the margins en- 

The specific epithet refers to the shape of the leaflets tire, revolute, the apex obtuse to acuminate, mucronate, 

as found in the type collection. 

the base attenuate, decurrent into the petiolule, some- 

Phenology. Flowers April to October, and fruit from times slightly unequal; petiolules 2-10 mm, glabrous, 

January to March. 

bulbous at base; rachis 6-12 cm long, crescent-shaped 

in cross-section, distal portion bicanaliculate along 

Distribution and Ecology. (Fig. 98) Venezuela, 

adaxial surface, glabrous; petioles (1.5) 4.5-7 cm long, 

Peru, and Brazil, in non-flooded, primary, lowland forest 

flattened along adaxial surface, glabrous, bulbous at 

and vatrzea forest. 

base. Thyrses panicle-shaped, terminal, to 10 cm long, 

Common name. Brazil: pitomba. 

with few lateral branches; cataphylls acicular, ca. 2.5 

Representative specimens examined. VENEZUELA. mm long, apiculate, distally tomentose; axes sulcate, 

Amazonas: Atabapo, Upper Orinoco river, 180 m, 3 minutely puberulent; bracts subulate, persistent, ca. 0.6 

Mar 1990 (fr), Aymard & Delgado 8409 (PORT, US); mm long; dichasia simple or distal flowers seemingly 

Rio Negro, 2km S of San Carlos de Rio Negro, 140 m, solitary; peduncle 0.5-3.5, flattened; pedicels ca. 1.5 

29 Jan 1992 (yfr), Aymard et al. 9808 (PORT, US); San mm long, articulate on lower third. Calyx ca. 2.5 mm 

Carlos de Rio Negro, 4.5 km NE of, 120 m, 5 Apr 1979 long, pilose-pubescent, the sepals 1.6-2.2 mm long, 

(fl), Liesner 6179 (MO, NY). 

ovate or oblong, concave, obtuse at apex, the margins 

PERU. Loreto: Prov. Ramon Castilla along ciliate, sub-hyaline; petals seemingly spatulate, ca. 4 mm 

Pucaurquillo-Pebas trail, 140 m, 31 Jan 1987 (fr), Stein 

long, glabrous, except for the ciliate margins, the apex 

et al. 3994 (F); Prov. Maynas. Iquitos. Alpahuayo, Estaci6n 

Experimental del IIAP, 13 Sep 1990 (fl), Vasquez & 

rounded, the base clawed; appendages as long as the 

Jaramillo 14401 (MO). Pasco: Oxapam, Caboose de petals, adnate to the base, oblong, erect, abaxially gla- 

Mono, Iscozacin river, 320 m, 8 Jun 1983 (st), Gentry brous, adaxially tomentose; disc cup-shaped, 5-lobed, 

et al. 41623 (MO), 9 Jun 1983 (st), Gentry et al. 41729 glabrous, ca. 0.6 mm tall; stamens 8, the filaments of 

(MO); Shiringamazu, ca. 20 km S of Iscozacin, Palcazu unequal length, 2-2.5 mm long, glabrous, the anthers 

river valley, 300 m, 9 Jul 1988 (st), Gentry et al. 63470 0.7-0.9 mm long, oblong-lanceolate, glabrous, apicu- 

(MO); Palcazu valley, Selva Central, 9 May 1985 (st), late at apex. Pistillate flowers and fruits not known. 

Hartshorn et al. 2732 (F, MO). 

The specific epithet which means swollen (oedo) 

BRAZIL. Acre: Hill dividing Brazil from Peru, 4 Mar and foot (podo) seems to refer to the swollen petioles 

1976 (fr), Marinho 350 (NY). Amazonas: Rio Negro, 


between Sao Gabriel & Manaus, without data (yfr), Coelho 

& Monteiro 29 (INPA); Near mouth of Embira river, basin Phenology. Unknown. 

of Jurua river, 10 Jun 1933 (fl), Krukoff 4750 (MO, US- 

2); Manaus-Porto Velho road, between Castanho & Tupana 


rivers, 20 Jul 1972 (fl), Silva 966 (INPA); Manaus- from the type collection. 

Itacoatiara road, km 26, 2 Jun 1997 (fl), Sothers et al. 

998 (US). Mato Grosso: Humboldt Center, 10 Oct 1973 

(fl), Berg et al. P-18388 (MG, MO, NY, US). Rondonia: 45. TALISIA PACHYCARPA RadLkofer, Sitzungsber. 

Bom Sossego, 3 Feb 1983 (fr), Carreira et al. 540 (IAN, Math.-Phys. Cl. Konigl. BayerAkad. Wiss. Miinchen


cm. 

Fig. 96. Talisia oedipoda. A. flowering branch. B. staminate flower. C. staminate flower with perianth removed 

showing disc and stamens (right), and 1.s. same showing pistillode. D. pistillode. (A-D from Riedel 522). 

8: 350. 1878. Type. French Guiana. Cayenne, with- 

out date (fr), Poiteau s.n. (holotype, G, n.v.; isotype, 

K?, n.v.; frag. at M). Fig. 97a-e 

Slender tree or treelet, 5-12 m tall, monocaulous; 

trunk to 15 cm in diam. Stems terete or furrowed, puberulent 

to minutely tomentose. Leaves paripinnate; 

distal process acicular, early deciduous; leaflets chartaceous, 

6-10, opposite to alternate, elliptic, 6.5-29(45) 

x 2-9.5(12) cm, the adaxial surface with impressed 

venation, glabrous, the abaxial surface minutely hirsute, 

with prominent venation, tertiary veins reticulate, the 

apex acuminate to long-acuminate, the base inequilateral, 

one side obtuse the other attenuate, the margins entire, 

strongly revolute; petiolules pulvinate, 4-6 mm long, 

minutely hirsute to glabrescent; rachis 20-40 cm long, 

terete, or slightly angled on distal portion, minutely 

hirsute; petioles 12-33 cm long, terete, minutely hir- 

sute, greatly enlarged at base. Thyrses panicle-shaped,


3mm 2mm 

Fig. 97. Talisia pachycarpa. A. portion of leaf. B. distal portion of branch with inflorescence. C. staminate 

flower. D. lateral and adaxial views of petal. E. staminate flower with perianth removed showing disc and stamens, 

l.s. of same (right). (All from L. da Cunha & Assunqjo 214a).


, SOWI gm. \ 70W1 60W 50WI 

'' 

\ , 

_ / za. v^ . u ,> X 2 

1.-'. 

* Talisia nervosa 

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

v Talisia obovata 

-P Talisia oedipoda 

*Talisia pachycarpa 

- ' u, 

, ,. 

,' 

j 

;-,.' 

', . 

, 

. 

, 2 


at least 35 cm long, terminal or axillary on distal part of 

branches; cataphylls acicular, 5-10 mm long, velutinous; 

axes angled, velutinous; bracts and bracteoles ovatedeltate, 

velutinous, 1-2 mm long; dichasia simple, 

sessile; pedicels short, flowers essentially sessile, to 

0.5 mm in fruit. Flowers sessile, congested along inflorescence 

axis. Calyx 3.5-5 mm long, light green, 

sericeous-velutinous, the sepals 1.5-2 mm long, oblong; 

petals white, 6-7 mm long, elliptic, abaxially glabrous, 

adaxially papillate, rounded at apex, long-attenuate 

at base; appendage as long as the petal, strap-shaped, 

abaxially glabrous, adaxially sericeous except for the 

glabrous base; disc 5-lobed, velutinous at apex, ca. 1 mm 

tall; stamens 8, the filaments of equal length, glabrous, 

3-4 mm long, filiform, the anthers oblong or linearlanceolate, 

1.5-1.8 mm long, apiculate at apex ovary 

sericeous, the stigma elongate-trigonous, ferruginouspapillate. 

Fruit trigonous, 2-2.5 cm long, sericeous, glabrescent, 

the pericarp woody, smooth, to 2.5 mm thick, 

the endocarp pilose, glabrescent. Seed not known. 

The specific epithet refers to the thickened pericarp 


Phenology. Collected in flowers in May, July, Au- 

gust, and December, and fruit during June and August. 

4W 

1 ON 


French Guiana, Guyana, and Brazil in non-flooded 

forest. 

Common name. Brazil: Pitomba. 


Upper Takutu/Upper Essequibo: Basin of Essequibo, 

near mouth of Onoro creek, 15 Dec 1937 (fl), Smith 

2677 (K, NY, US). 

BRAZIL. Amazonas: Manaus. Dtto. Agropecuario. 

BDFF reserve, km 41, 50-125 m, 9 Jul 1989 (fl), L. da 

Cunha & Assunfao 214a (NY); Novo Aripuana, BR 230, 

Rod. Transamaz6nica, 400 km from Humaita, 4 May 1985 

(fl), C. Ferreira 6036 (K, MG, NY); Mun. Presidente 

Figueredo, along road to hydroelectric plant, ca. airport, 

16 Jul 1986 (fl), C. Ferreira 7591 (US); Mun. Oriximina. 

Mapuera river, vic. of cachoeira Sao Marcal, 13 Aug 

1986 (fr), C. Ferreira et al. 7720 (INPA, NY, US); Reserva 

Ducke, Manaus-Itacoatiara, Km 26, Tinga creek, 12 Aug 

1993 (fr), Ribeiro et al. 1126 (MG, NY, US); Manaus. 

Porto Velho, 12 Jul 1972 (fl), Silva et al. 519 (INPA);12 

Jul 1972 (fl), Silva et al. 530 (INPA-2). Rondonia: Hy- 

droelectric plant reserve, close to Japiim creek, 18 Jun 

1986 (yfr), C. Ferreira 7495 (US); Puerto Velho to 

Cuiaba hwy., vic. of Santa Barbara, 12 Aug 1968 (fl), 

Prance & Ramos 6909 (K-2, MG, NY, US).


46. TALISIA PILOSULA Sagot ex Radlkofer, Representative specimens examined. FRENCH 

Sitzungsber. Math.-Phys. Cl. K6nigl. Bayer Akad. GUIANA. Bassin du Sinnamary: Plomb creek, 20 m, 

Wiss. Miinchen 8: 349. 1878. Type. French Guiana. 16 Mar 1994 (fr), Bordenave 832 (CAY, US); Karouany, 

Acarouany, 1858 (fl), Sagot s.n. (holotype, B-de- 1859 (fl), Sagot s.n. (K). 

stroyed, photo (F neg. 5680) at US; isotypes, K, P- 

3; frag. at M). Fig. 99a-g 

47. TALISIA PINNATA (Ruiz & Pavon) Radlkofer, 

Tree to 23 m tall. Stems nearly terete, pubescent, 


becoming glabrous, and slightly glossy. Leaves 

Wiss. Miinchen 8: 351. 1878. Acladodea pinnata 

paripinnate; distal process acicular, tardily deciduous, 

Ruiz & Pav6n, Syst. Fl. Peruv. 1: 262. 1798. Talisia 

0.5-2 cm long; leaflets 16-18, subopposite oraltemate, 

(?)acladodea de Candolle, Prodr. 1: 609. 1824. Type. 

elliptic or oblong-elliptic, coriaceous, 5.5-12 x 1.4-3.2 

Peru. Huinuco: Pillao, forest at Chacahuas, Oct-Nov 

cm, the venation brochidodromus, the adaxial surface 

(fl), Pavon s.n. (holotype, MA, n.v., photo at US; 

with impressed venation, glabrous, the abaxial surface 

isotypes, BM; frag. ex MA at M). Fig. 100a-h 

pilose to minutely hirsute, with prominent venation, 

tertiary veins reticulate, the apex caudate, the base Treelet 1-6.5 m tall. Stems terete, densely hirsutuinequilateral, 

one side obtuse the other rounded, the lous to tomentose when young. Leaves paripinnate or less 

margins entire, revolute; petiolules cylindrical, ca. 2 mm often imparipinnate, spirally arranged on distal portion of 

long, abaxially pilose or glabrescent; rachis 16-28 cm stem; distal process filiform, ca. 1 cm long, persistent; 

long, nearly terete on proximal portion, slightly flattened leaflets 14-20, alternate or subopposite, oblong, oblongon 

distal portion, minutely hirsute orpilose; petioles 5.5- elliptic or oblanceolate, chartaceous, 6.5-25.5 x 2.2-5.7 

1 1.5 cm long, terete, pilose or minutely hirsute, slightly cm, the adaxial surface glabrous, except for the minutely 

enlarged at base. Thyrsespanicle-shaped, 1-12 cm long, tomentose impressed midvein and part of the secondary 

supra-axillary on distal portion of branch; cataphylls veins, the abaxial surface ferruginous-pilose, densely so 

acicular or pinnate, 4-10 mm long, ferruginous hirsute along the prominent veins, the venation brochidodromus, 

in a supra-axillary cluster; axes slightly angled, hirsute tertiary veins reticulate, the apex acuminate to caudate, 

with intermixed multicellular longer trichomes; bracts and the base inequilateral, one side obtuse the other acute, 

bracteoles subulate, 2-4 mm long, hirsute, with numer- the margins undulate, slightly revolute; petiolules slenous 

multicellular, elongated trichomes; dichasia simple, der, sub-terete, 3-7 mm long, densely ferruginous-pior 

the distal flowers seemingly solitary (lateral flowers lose; rachis 23-35 cm long, terete, striate, densely feraborted), 

sessile; pedicels 0.5-1 mm long, articulate at ruginous-pilose to nearly tomentose; petioles ca. 15 cm 

the apex. Calyx 1.5-3 mm long, minutely hirsute, interlong, 

terete, striate, densely ferruginous-pilose, enlarged 

mingled with multicellular, glandular trichomes, the seat 

base. Thyrses panicle-shaped, 14-33 cm long, termipals 

1-2 mm long, ovate or lanceolate, concave; petals 

nal or axillary on distal portion of stem; cataphylls hir- 

4-4.5 mm long, elliptic, abaxially glabrous, adaxially with 

sutulous to tomentose, transitional from leaves (pinnate) 

elongated glandular papillae, obtuse at apex, cuneate to 

to bracts (acicular), numerous, congested at distal porattenuate 

at base; appendage as long as the petal, elliptic, 

tion of stem, to 11 cm long; axes slightly angled, ferabaxially 

glabrous, adaxially hirsute onupperhalf, fused 

ruginous-pilose to -tomentose, intermixed with 

to the petals base margins to form a pocket; disc 5-lobed, 

multiseptate, glandular hairs; secondary axes to 10.5 cm 

hirsute, ca. 0.5 mm tall; stamens 5, the filaments of equal 

long at the base of inflorescence, gradually decreasing 

length or slightly unequal, glabrous, 3-3.3 mm long, 

toward distal 

filiform, the anthers oblong, 1-1.3 mm long, apiculate at 

portion; bracts and bracteoles subulate, 

apex; ovary hirsute, conical, the 

pilose, sometimes with glandular hairs, 1-2 mm long; 

stigma elongatetrigonous, 

ferruginous-papillate. Fruit 

dichasia 

ellipsoid, apicucompound; 

peduncle 5-8 mm long, flattened, 

late, ca. 1.8 cm long, puberulent, sub-tuberculate, the 

densely pilose; pedicels ca. 1 mm long, articulate at the 

pericarp coriaceous, ca. 1 mm thick, the endocarp glamiddle. 

Calyx 4-4.7 mm long, abaxially ferruginousbrous. 

Seed solitary, the testa fleshy. Embryo with cotyhirsutulous, 

intermixed with few glandular, septate hairs, 

ledons lying transversely over each other, the lower one 

adaxially sparsely pilose, the sepals 1.5-2 mm long, 

slightly larger than the upper one. 

ovate, rounded; petals white, 4.5-5.5 mm long, oval to 

The specific epithet refers to the pilose indumenobovate, 

abaxially appressed-pubescent dorsally and at 

tum of this species. 

base, adaxially glabrous, ciliate on lower margins, 

rounded at apex, clawed at base; appendage as long as 

Phenology. Flowering period unknown, fruits col- the petal, oblong, sericeous on both surface; disc cuplected 

in March. 

shaped, 5-lobed, hirsute on distal portion, ca. 1 mm tall; 

Distribution and Ecology. (Fig. 103) Known only stamens 5-8, the filaments filifonn, of equal or nearly 

in French Guiana from riverine forest. 

equal length, 2-3 mm long, pilose, especially on upper


Fig. 99. Talisia pilosula. A. portion of leaf and detail of abaxial surface of leaflet. B. inflorescence. C. detail of 

inflorescence branch. D. staminate flower. E. adaxial and abaxial views of petal with adnate appendage. F. staminate 

flower with perianth removed showing disc, stamens, and detail of anthers. G. pistillate flower with perianth removed 

showing disc, sterile stamens, and pistil. (All from Sagot 1898).


3 mm.I 

Fig. 100. Talisia pinnata. A. portion of leaf. B. inflorescence C. pistillate flower. D. adaxial, abaxial, and lateral 

views of petal with adnate appendage. E. staminate flower with perianth removed showing disc and stamens. F. pistillate 

flower with perianth removed showing disc, sterile stamens and pistil, and l.s. same, showing ovule. G. distal 

portion of branch with cataphylls and infructescence. H. lateral view of embryo. (A, B, E from Acevedo R. et al. 

1585; C, D, F from Vasquez 12293; G-H from Vdsquez et al. 4869). 

I Im 

X


half, the anthers oblong, ca. 1.5 mm long, apiculate at entire length, 0.5-1.2 cm long, minutely pilose; rachis 

apex; ovary conical, appressed-pubescent, the stigma terete, striate, puberulent to pubescent, sometimes with 

capitate. Fruit yellow at maturity, ellipsoid 3.2-3.5 cm few to many glandular hairs, 40 to 60 cm long; petioles 

long, granulate, sparsely appressed-puberulent, apicu- 14-31 cm long, terete, striate, puberulent, thickened at 

late at apex, the pericarp coriaceous, ca. 1 mm thick, the base. Thyrses panicle-shaped, compound, terminal, pyendocarp 

glabrous. Seed 1 or 2 per fruit, ellipsoid, with ramidal, to 80 cm long, lateral basal branches to 35 cm 

fleshy testa. Embryo with cotyledons lying dorsiventrally long; cataphylls transitional from leaves (leaf-like) to 

or obliquely dorsiventrally over each other, the lower bracts (acicular), numerous, congested at distal portion 

one slightly smaller than the upper one. 

of stem or supra-axillary, reaching 20 cm long when ter- 

The specific epithet refers to the pinnately com- minal, 3-5 cm long when axillary; axes nearly terete, 

pound leaves. 

striate to sulcate, appressed-pubescent to nearly tomen- 

Phenology. Known to flower from April to July, 

and in October and November, and to fruit in January 

and July. 

tose; bracts subulate, to 4 mm long, tomentose, persistent; 

bracteoles lanceolate, puberulent to pubescent, persistent, 

1.5-2 mm long; dichasia compound, sessile; 

pedicels < 0.5 mm long, tomentose. Calyx 2.8-3.2 mm 

Distribution and Ecology. (Fig. 103) Known only long, tomentose, the sepals 1.5-2 mm long, rounded or 

from Peru in understory of seasonally flooded forest. oblong-rounded, sometimes with less pubescence around 

Representative specimens examined. PERU. Loreto: 

Sapuena. Riparian, 200 m, 15 Apr 1987 (fl), Acevedo R. 

et al. 1585 (NY, US-2); Requena. Reserva Nacional 

the ciliate margins; petals obovate, 5-5.5 mm long, reflexed 

at anthesis, glabrous on adaxial surface, appressed-pubescent 

on abaxial surface at base, the apex 

Pacaya Samiria Santa Cruz, 3 May 1993 (fl), del Carpio rounded, the base clawed; appendages oblong, as long 

2016 (USM); Nanay river, slightly below Bellavista, 12 

Jul 1976 (fl, yfr), Rimachi 2408 (NY, USM); Ucayali. 

Jenaro Herrera. Quebrada Sapuay, 120 m, 7 Jun 1989 

(fl), Vasquez 12293 (US, USM); Yarina, Trapiche river, 

180 m, 11 Jan 1984 (fr), Vdsquez et al. 4869 (MO). 

as the petals, erect, villous on both surfaces; disc cupshaped, 

5-lobed, tomentose at apex, ca. 0.6 mm tall; stamens 

8, the filaments sparingly pilose, of equal length, 

the anthers lanceolate, glabrous, apiculate at apex; ovary 

conical, tomentose, the stigma conical-trilobed, papillate. 

Fruits ellipsoid to globose, glabrous, 3.5-4 cm long, 

48. TALISIA PRINCEPS Oliver, Hooker's Icon. 

Plant. Ser. 3, 18: 1769. 1888. Type. Venezuela?, 

Specimen from plant cultivated at Hort. Crawford, 

Oct 1878 (fl), J. Cook, s.n. (lectotype, K-4 sheets, 

here designated). Syntype. Unknown origin, plant 

cultivated at Kew Gardens, May 1886 (st), Anonymous, 

s.n. (K-3 sheets). Fig. 10la-i 

Theophrasta pinnata Jacq., Fragm. Bot. 49. 1800. 

Type. Venezuela. Tab. 64, fig. 2, 65, 66. 

Talisia erecta Radlkofer in Martius, Fl. Bras. 13(3): 

550. 1900. Type. Venezuela. A specimen from plant 

cultivated at Kew Gardens, Anonymous, s.n. (hogranulose, 

apiculate, yellow, the pericarp woody, ca. 5 

mm thick. Seed ellipsoid to nearly globose, 2-2.3 cm 

long. Embryo with cotyledons superimposed, the upper 

one slightly larger than the lower one. 

Talisia princeps is distinguished from many other 

Talisia species by its oblanceolate leaflets with long, 

abruptly acuminate apices, its long (to 7 cm) distal processes, 

and its petiolules enlarged throughout their 

length. In addition, T princeps has numerous cataphylls 

or rudimentary leaves on distal portions of stems and 

leaf axils; the cincinni are sessile, and the fruits woody, 

ovoid and large. The specific epithet means distinlotype, 

M, photo at US; isotypes, BM, n.v., M). guished or noble perhaps in reference to the graceful 

Treeletortree, 2-20mtall. Stems nearly terete, densehabit 


ly cano-pubescent when young, with numerous leaf Phenology. Collected in flower during December 

scars. Leaves paripinnate; distal process filiform, 1-7 and in fruit during December and January. 

cm long, persistent; leaflets (6)16-24, altemate to opposite, 

oblong-oblanceolate, oblanceolate or less often 

oblong, 9-42 x 3-8 cm, chartaceous to subcoriaceous, 

the adaxial surface glabrous, the abaxial surface minutely 

pilose (sometimes with glandular hairs) along veins, the 


Panama, Venezuela, Ecuador, and Peru in lowland, rain 

forest. 

Common names. Venezuela: Copito, limpia botella 

venation brochidodromus, plane on adaxial surface, (fide Steyermark, 1994). 

prominent on abaxial surface, with secondaries forming 

a 450 angle with midvein, tertiary veins reticulate, the 

margins entire, the apex long-acuminate, sometimes 

abruptly so, the base acute to obtuse, strongly to slightly 

unequal; petiolules nearly cylindrical, thickened for their 

Representative specimens examined. PANAMA. 

Bocas del Toro: Without specific locality, hill above railroad 

station at mile 7.5, 27 Jul 1971 (st), Croat 16376 (MO). 

VENEZUELA. Aragua: Turiamo road km 50-60, 

300 m, 10 Nov 1961(fl), Trujillo 5071 (NY). Distrito


Fig. 101. Talisia princeps. A. portion of leaf. B. distal portion of branch showing cataphylls. C. flower bud. 

D. pistillate flower. E. pistillate flower with perianth removed showing disc, sterile stamens and pistil. F. stamen. 

G. abaxial and adaxial views of petal with adante appendage. H. l.s. of fruit showing mesocarp and seed. I. fruit and 

ventral view of embryo. (A, C-G, from Cook, Oct 1878; B from Steyermark 119784; H-I from Cook, Sep 1879).


Federal: Km 18, along road from Caracas to La Guaira, base clawed, the apex rounded, the margins ciliate; ap- 

500 m, 28 May 1945 (st), Steyermark 62928 (F-3, US), pendage as long as the petal, deltate, sometimes bifur- 

Dec 1940 (fl), Tamayo 1500 (F, US, VEN). Falc6n: S of cate, abaxiallypapillate, adaxiallysericeous; disc 5-lobed, 

Punta Faustino, 15-25 m, 29 Aug 1974 (st), Steyermark glabrous, 0.8-1 mm tall; stamens (7) 8, the filaments 

& Manara 110418 (MO); Silva. Cerro Chichiriviche, 

glabrous, slightly unequal, 1.5-3.3 mm long, the anthers 

above La Soledad, 200 m, 5 Sep 1974 (st), Steyermark & 

Manara 110778 (MO). Miranda: Bri6n. 4.8 km E-NE 

oblong or oblong-lanceolate, 1.2 mm long, apiculate at 

of Pueblo Seco, along creek Agua Bendita, 75 m, 24-25 apex; ovary ovoid to globose, glabrous, the stigma cy- 

Mar 1973 (st), Steyermark & Carreno 106915 (F). lindric, papillate. Fruits ellipsoid, to nearly globose, 3.3- 

TAchira: Cerro Cuite, along creek La Colorada, 450- 3.7 cm long, orange-yellow or orange when ripe, gla- 

630 m, 9 Nov 1979 (st), Steyermark et al. 119784 (MO). brous, smooth, the pericarp coriaceous, ca. 2 mm thick, 

ECUADOR. Napo: San Jose de Payamico, 40 km W endocarp irregularly woolly-pubescent. Seeds solitary, 

of Coca, 300-600 m, 11 Jan 1984 (fr), Irvine 576 (F); the testa fleshy, brownish. Embryo of 2 nearly equal, 

Via Hollin-Loreto; km 60, ca. Guataraco river, 650 m, 

10 Dec 1987 (fr), Palacios 2182 (K-3, MO, NY, US-4). 

PERU. Loreto: Tasha Curaray river, 19 Sep 1972 

(bd), Croat 20433 (MO, NY). 

erect cotyledons, perpendicular to the radicle, or laying 

obliquely dorsiventrally over each other. 

Talisia retusa can be vegetatively similar to Talisia 

cerasina, however, the former can be distinguished 

from the latter by its leaflets which are always sym- 

49. TALISIA RETUSA Cowan, Brittonia 7: 403. 

1952. Type. Guyana. Komo Falls, Takutu river, 

Nov 1948 (fl), Wilson-Browne 508 (holotype, NY; 

isotype, K, photo at MO, NY). Fig. 102a-h 

metrical, with obtuse, rounded and usually retuse apex 

(vs. leaflets asymmetrical especially at base, short to 

long-acuminate at apex). The specific epithet refers to 

the retuse apices of the leaflets in this species. 

Talisia heterodoxa Steyermark, Ann. Missouri Bot. Phenology. Flower from April to December and fruit 


Represa Guri, islands 6-18 km S of dam, forest, 

7038'N, 62058'W, 220-240 m, 2 Apr 1981 (fl), 

Liesner & Gonzalez 11151 (holotype, MO; 

isotypes, NY, US, VEN). 

from September to April. 

Distribution and Ecology. (Fig. 103) Venezuela, 

Guyana, Surinam, Peru, Brazil, and Bolivia. In terra 

firme, vatrzea, dry, and semideciduous forests. 

Tree 20-25 m tall; trunk reaching 30-40 cm in diam., 

not buttressed. Branches terete, glabrous. Leaves 

paripinnate or imparipinnate; distal process acicular, truncate, 

2-4 mm long; leaflets (2-) 8-12, opposite to alternate, 

oblong-elliptic, oblong or oblanceolate, coriaceous, 

5.5-11.5 (-16) x 2-4 (-6.5) cm, glabrous, the adaxial 

surface with prominulous midvein, secondary veins 

plane or slightly impressed, the abaxial surface glandular-punctate, 

with prominent midvein and secondary 

veins, tertiary veins reticulate, the apex rounded, usually 

retuse and sometimes mucronate, the base attenuate or 

obtuse, decurrent on to the petiolule; petiolules pulvinate 

at base, 4-7 mm long; rachis 3-20 cm long, obtusely 

angled, adaxially bisulcate, abaxially striate, glabrous; 

petioles 1.7-11 cm long, slightly flattened along 

adaxial surface, abaxially striate, glabrous, enlarged at 

base. Thyrses panicle-shaped, 10-40 cm long, terminal 

or axillary; cataphylls supra-axillary, clustered, acicular, 

glabrous or puberulent, 2-5 mm long; axes nearly terete, 

sulcate, puberulent; secondary axes to 20 cm long; 

bracts and bracteoles subulate, early deciduous; dichasia 

simple or compound; peduncle 0.5-1 mm long, tomentose; 

pedicels 0.5-0.7 mm long, tomentulose, articulate 

at apex. Calyx ca. 2.2 mm long, puberulent on both 

surfaces, the sepals 1.5-2 mm long, ovate or roundedovate, 

ciliate at margins; petals white, oblong, or elliptic, 

4.8-6.2 mm long, abaxially glabrous except for the appressed-pubescent 

base, adaxially sparsely papillate, the 

Common names and uses. Brazil: Pitomba, 

pitomba de anta, pitombinha, macucu. Venezuela: 

7iestigo. The seeds have a fleshy edible testa. 


ZUELA. Bolivar: Mun. Piar, Isla Redonda, Lago Guri, 

15 km E of the damp, Feb 1992 (fr), Aymard et al. 10148 

(NY, PORT), Represa Guri, 220-240 m, 2 Apr 1981 (fr), 

Liesner & Gonzalez 11116 (MO, NY, PORT, US); Bajo 

Caura, Isla de Guayapo, 250-300 m, 16 Apr 1939 (fl), 

Williams 11833 (US-2, VEN). 

GUYANA. Upper Takutu/ Upper Essequibo: Nappi 

creek watershed, 350-600 m, 9 Feb 1993 (st), Hoffman 

& Foster 3599 (US); Wabuwak, Kanuku Mts., 615 m, 

Nov 1948 (fl), Wilson-Browne 437 (NY-2, US), 600 m, 

Nov 1948 (fl), Wilson-Browne 471 (NY). 

SURINAM. Nickerie: Sipaliwini, 325 m, 13 Nov 

1968 (fl), Oldenburger et al. 538 (K, NY). 

PERU. Huanuco: Pachitea, 300-400 m, 17 Feb 

1967 (fr), Schunke 1637 (F, NY, US). Madre de Dios: 

Tambopata Tourist Camp at junction of Tambopata & 

La Torre rivers, 280 m, 22 Jul 1985 (st), Gentry et al. 

51136 (MO). Puno: Candamo river, 810 m, 25 May 

1992 (st), Gentry et al. 77167 (MO). 

BRAZIL. Acre: Brasileia, 3 Nov 1980 (fr), C. 

Ferreira et al. 3114 (INPA, MG, MO, NY, US); Brasileia- 

Assis road, 2 Nov 1980 (fr), Lowrie et al. 684 (MG, US); 

Tarauaca. basin of Jurua river, Tarauaca' river. Seringal 

Macei6, 23 Sep 1994 (fl), Silveira et al. 876 (US). 

Maranhao: Island of Sao Luis, Feb-Mar 1939 (fr), 

Froes & Krukoff 11510 (MO, U); Alto Turi, 26 Oct 1962


4, 

/ 

Fig. 102. Talisia retusa. A. flowering branch. B. detail of abaxial surface of leaflet and apex showing mucron. 

C. staminate flower. D. I.s. staminate flower showing disc, stamens, and pistillode. E. adaxial and abaxial views of 

petal with adnate appendage. F. staminate flower with perianth removed showing disc and stamens. G. stamen. 

H. partly denuded infructescence with fruit. (A-G from Wilson-Browne 471; H from Vieira 801).


9 ;w WMj ' 7OWI 6(V 5 WVI 40W 


3mm3 

Fig. 104. Talisia setigera. A. portion of inflorescence and leaf. B. detail of abaxial side of leaflet showing 

petiolule, midvein, and pubescence. C. detail of inflorescence with denuded dichasium, and detail of trichome. 

D. staminate flower. E. adaxial and abaxial views of petal with adnate appendage. F. staminate flower with perianth 

removed showing disc and stamens (left) and pistillode (right). G. stamen. H. portion of infructescence. (A-G 

from Asplund 5412; from Dodson 5893).


slightly striate abaxially, with same indumentum as the Common name. Huevos de chivo. 

rachis, enlarged at base. Thyrses panicle-shaped, ca. 

Representative specimens examined. ECUADOR. 

35 cm long, terminal or axillary; cataphylls pinnate, or Esmeraldas: Pambil river near Estero Capuli, 150 m, 3 

less often acicular, 1-3 cm long, setigerous; axis angled, Jul 1966 (fr), Jativa & Epling 1057 (MO, NY, U, US). 

sulcate, sericeous-setigerous, the hairs ferruginous and Esmeraldas: 50 km SW of Atacames along the 

septate, intermixed with shorter, straight hairs; bracts Esmeraldas-Muisne road, 150 m, 16 Sep 1982 (fr), 

and bracteoles subulate, ca. 1 mm long, pubescent, early Balslev & Steere 3101 (QCA, NY). Los Rios: Hacienda 

deciduous; dichasia compound, simple or of a solitary Clementina on Pita river, 20 Mar 1939 (fl), Asplund 5412 

flower due to the abortion of lateral flowers; peduncle (K, US); Santo Domingo, Rio Palenque Biological Station, 

150-200 m, 11 Feb 1973 (fr), Dodson 5236 (US), 

flattened, 2-6 mm long; pedicels 1-2.5 mm long, ar- 

25 Jul 1975 (fr), Dodson 5893 (MO, QCA, US), 1 Mar 

ticulate at upper third, both with same indumentum as 

1976 (fl), Dodson 5998 (MO, QCA); Canton Vinces, km 

the axis. Calyx light green, 3-4.5 mm long, puberulent 70 along Quevedo-Palenque road, 100 m, 23 Mar 1980 

and sparsely setigerous, the sepals 2-3.5 mm long, (fl), Dodson & Gentry 9798 (F); Jauneche Forest, 100 m, 

oblong, elliptic or deltate, concave, obtuse to acute at 14 Jul 1979 (fr), Dodson et al. 7989 (MO); Montalvo, 

apex, ciliate at margins; petals white, 5-6 mm long, Bosque del Oro, Hac. Las Balsas, 30-400 m, 28 Jun 1964 

oblong to nearly spatulate, glabrous except for the (st), Jativa & Epling 636 (MO). Manabi: San Sebastian. 

abaxially tomentose base and ciliate lower margins, Parque Nacional Machalilla, 600 m, 20 May 1995 (fr), 

rounded at apex, cuneate at base; appendage as long as Cornejo & Bonifaz 3878 (US); Portoviejo-Pichincha 

road, 3 km E of San Placido, 150-200 m, 6 May 1985 

the petal, deltate, abaxially sparsely appressed-pubes- 

(fr), Harling & Andersson 24900 (QCA). 

cent sometimes nearly glabrous, adaxially sericeous 

above the base; disc cup-shaped, 5-lobed, hispidulous 

at apex, ca. 1 mm tall; stamens 8, the filaments of equal 51. TALISIA STRICTA (Karsten & Triana) Triana 

or nearly equal length, sparsely pilose, ca. 3.5 mm long, & Planchon, Ann. Sci. Nat. Ser. 4, 18: 369. 1862. 

the anthers oblong or oblong-lanceolate, ca. 1.5 mm Comatoglossum strictum Karsten & Triana in 

long, apiculate at apex; ovary ovoid, sericeous, the stigma Triana, Nuev. Jen. Esp. 11. 1854. Type. Colomoblong, 

trigonous, short, papillate. Fruit yellow, obov- bia. Bogoti: between Apulo & Piedras, 600 m, 

oid, apiculate, ca. 3 cm long, puberulent, granulate, the Mar 1853 (fl), Triana s.n. (lectotype, COL, here 

pericarp woody, ca. 2 mm thick. Seeds one or two per designated; isolectotype, COL). Syntypes. Colomfruit, 

the testa fleshy. Embryo with cotyledons super- bia. Bogota: between Apulo & Magalena, 800 m, 

imposed, the upper one larger than the lower one. s.d. (fl) Triana s.n. (K, P-2); Bogota: between 

A photograph of a likely holotype (Eggers 14704, Tocaima & Magalena, 300-800 m, s.d. (fl) Triana 

annotated by Radikofer) for T setigera., was distributed s.n. (K). Fig. 105a-h 

by the Field Museum (F neg. 6025) as being a Munich 

specimen. However, I have not been able to locate this Treelet 1-3 m tall. Stems terete, cano-velutinous, 

specimen at Munich, suggesting that the specimen be- smooth. Leaves paripinnate; distal process acicular, 4- 

longed to another herbarium. It is quite possible that 6 mm long; leaflets 12-22, opposite to alternate, oblanthis 

specimen may have been from Berlin, in which case 

ceolate to nearly oblong, 9-18 x 3.2-6 cm, chartaceous 

it would have been destroyed. Not being able to locate 

to coriaceous, the adaxial surface glabrous except for 

the sometimes canescent midvein, the abaxial surface 

this specimen, I have decided to lectotypify T setigera 

canescent, the venation brochidodromus, plane on 

with one of the remaining isotypes. 

adaxial surface, prominent on abaxial surface, tertiary 

Talisia setigera seems to be closely related to T. 

veins reticulate, the margins entire, revolute, ciliate, the 

pinnata as they share numerous morphological feaapex 

obtusely and abruptly acuminate or less often acutures. 

However, T setigera can be distinguished from 

minate, the base cuneate-obtuse, conspicuously asymthe 

latter by its setigerous branches and abaxial side of 

metrical; petiolules nearly cylindrical, velutinous, thickleaflets 

(vs. hirsutulose to tomentose branches and piened 

for their entire length, slightly flattened adaxially, 

lose abaxial surface of leaflets). The specific epithet 

3-4 mm long, velutinous; rachis terete, bicanaliculate 

refers to the setigerous indumentum in this species. 

toward distal portion, cano-pubescent, 16.5-20 cm 

Phenology. Collected in flower during March and 

in fruit in February, May, and July. 

long; petioles 9-13 cm long, terete, cano-pubescent, 

thickened at base. Thyrses panicle-shaped, pyramidal, 

terminal, to 50 cm long, lateral basal branches to 20 cm 

Distribution and Ecology. (Fig. 107) Known only long; cataphylls pinnate, 6.5-10 cm long, cano-velutinous 

from Ecuador, in understory of moist, seasonally to -pilose, congested at distal portion of stem; axes 

flooded forest. 

nearly terete, striate to sulcate, tomentulose to tomen-


5mm 

\~~~~~~~~~~~~~~~~~~~~~~~qo 

IIP, 

2mm 2m[m. ./A 

Fig. 105. Talisia stricta. A. portion of flowering branch and portion of leaf. B. staminate flower. C. adaxial and 

abaxial views of petal with adnate appendage. D. l.s. staminate flower showing disc, stamens and pistillode. E. stamen. 

F. l.s. pistillate flower showing ovules. G. pistillate flower with perianth removed showing disc, sterile stamens and 

pistil, and c.s. pistil. H. sterile stamen. (All from Killip 38319). 

3 cm.[


tose; bracts oblong, tomentulose to tomentose, persistent, 

3-4 mm long; dichasia compound; peduncles, 6- 

15 mm long, flattened; pedicels < 1 mm long, tomentose, 

articulate at base. Calyx 4.5-6mm long, tomentose, 

the sepals ovate, the 3 outer ones ca. 3 mm long, the 2 

Talisia carinata forma acutisepala Radlkofer in 

Martius, Fl. Bras. 13 (3): 549. 1900. Type. French 

Guiana. Without specific locality or date, Melinon 

357 (lectotype, P, here designated). Syntype. 

French Guiana. Acarouany, May 1858 (fl), Sagot 

s.n. (K, P-2). 

inner ones ca. 2 mm long; petals white, nearly spatu- Talisia micrantha Radlkofer, Fedd. Repert. Nov. Spec. 

late, 6-9 mm long, reflexed at anthesis, abaxially to- 9: 375. 1911. Type. Surinam. Tapanahony river, 

mentose on lower half, adaxially glabrous, the apex Aug 1904 (fl), Versteeg 807 (holotype, U). 

rounded, the base clawed; appendages oblong, as long 

Talisia reticulata Radlkofer, Repert. Spec. Nov. Regni 

Veg. 9: 376. 1911. Type. Surinam. Tapanahony 

or little shorter than the petals, bifid, erect, tightly conriver, 

vic. of Onie Haberteg, Jul 1904 (fl), 

vergent, villous on both surfaces; disc cup-shaped, 5- Versteeg 674 (holotype, U, photo at US; isotypes, 

lobed, hirsute, ca. 1 mm tall; stamens 8, the filaments M, P-2) 

sparingly woolly-pubescent, of equal length, ca. 2.5 mm 

Slender tree or treelet, 2-8 m tall, unbranched or 

long, the anthers oblong, ca. 2.2 mm long, glabrous, 

with few ascending, sympodial branches on distal porapiculate 

at apex; ovary conical, tomentose, the stigma 

tion; trunk to 8 cm in diam.; bark light brown, 

cylindrical, papillate. Fruits nearly ovoid, appressed 

lenticellate. Stems terete, puberulent or tomentulose, 

ferruginous-pubescent, ca. 3 cm long, minutely 

dull, becoming lenticellate and glabrous with age. 

granulose, apiculate, the pericarp woody, ca. 1.5 mm 

Leaves paripinnate or imparipinnate, spirally arranged 

thick, the endocarp puberulent. Seed solitary or less 

on distal portion of stem; distal process 2-3 mm long, 

often two per fruit, ovoid, the sarcotesta very thin, not 

acicular or truncate; leaflets 5-14, altemate or opponoticeable 

when dry. Embryo of 2 equal, erect cotylesite, 

elliptic or oblong-elliptic, chartaceous, 7.4-26 (40) 

dons, or these obliquely superimposed. 

x (2.2) 3.5-9(-16) cm, the adaxial surface glabrous or 

Talisia stricta is distinguished by its canescent 

sometimes with a few hairs along the prominulous 

young stems, leaf rachises and abaxial surface of the 

midvein, primary and secondary veins impressed, the 

leaflets. The specific epithet refers to the tightly conabaxial 

surface with puberulent, prominent primary 

vergent petal appendages. 

and secondary veins, secondary veins collected into a 

Phenology. Flowering from January to June and 

fruiting in March and August. 

strong intramarginal vein, tertiary veins prominulous, 

reticulate, the apex acuminate, abruptly acuminate or 

apiculate, the base usually inequilateral, one side ob- 

Distribution and Ecology. (Fig. 107) Known only tuse the other acute or attenuate, the margins entire to 

from Colombia in forest on hillsides and along slightly undulate; petiolules pulvinate, minutely puriverbanks. 

bescent, conical, 5-10 mm long, sometimes extend- 

Common name. Mata puerco. 

ing into the blade; rachis (14.5)19-32(48) cm long, 

minutely pubescent, terete or sometimes obtusely 

Representative specimens examined. COLOMBIA. angled or canaliculate on distal portion, lenticellate and 

Without specific locality or date (fr), Mutis 4195 (US). striate along proximal portion; petioles 12-21(28) cm 

Casanare: San Ignacio, 5 Jan 1879 (fl), Andre 582 (K- long, terete, minutely pubescent, lenticellate, greatly 

2), Andre 583 (K, NY). Cundinamarca: Narifio, 350- enlarged at base. Thyrses panicle-shaped, 15-30 cm 

450 m, 1 Mar 1986 (fr), Ferndndez A. et al. 5484 (COL); long, axillary on distal portion of branches, or sel- 

Quebrada Carmargo, 460-480 m, 5 May 1944 (fl, yfr), dom cauliflorous; cataphylls acicular to dendroid, 4- 

Killip et al. 38246 (US); Tocaima, Haz. El Cucharo, 7 

12 mm long, minutely pubescent, clustered, supra- 

May 1944 (fl), Killip et al. 38319 (US); Viota to Girardot, 

axillary; axes sulcate, pinkish tinged, minutely pilose; 

320-560 m, Aug 1964 (fr), Saravia 4660, (COL); 

Tocaima, Haz. El Cucharo, 350 m, 14 Jun 1952 (fl), 

bracts and bracteoles subulate, 1-1.5 mm long; dicha- 

Uribe U. & Garcia 2317 (JAUM, US). Tolima: Chicoral, sia compound; peduncle 1-2 mm long, minutely, 

450 m, 11 Mar 1949 (fl), Haught 6353 (US-2). densely pilose; pedicels 1-1.7 mm long, articulate on 

upper half. Flower buds 5-angled-ovoid. Calyx cupshaped, 

pinkish tinged, 1-1.5 (2.5) mm long, minutely 

52. TALISIA SYLVATICA (Aublet) Radlkofer, pilose, the sepals 0.5-0.6 (1) mm long, ovate-deltate, 

Sitzungsber. Math.-Phys. Cl. K6nigl. Bayer Akad. strongly concave to carinate; petals cream or white, 3.5- 

Wiss. Miinchen 8: 341. 1 878.-Racaria sylvatica 4.5 mm long, oblong-elliptic, reflexed at anthesis, 

Aublet, Hist. P1. Guiane Suppl. 24. Tab. 382. 1775. abaxially glabrous, adaxially papillate or sparsely papil- 

Type. French Guiana. Without specific locality or late, obtuse at both ends, ciliate along proximal pordate 

(fr), Aublet s.n. (holotype, BM). Fig. 106a-i tion of margins; appendage elongate-deltate, as long


Fig. 106. Talisia sylvatica. A. leaf. B. detail of leaflets. C. inflorescence. D. dichasium. E. staminate flower. F. 

adaxial and abaxial views of petal with adnate appendage. G. staminate flower with perianth removed showing 

disc and stamens (left) and l.s. same showing pistillode. H. portion of infructescence. I. dorsal view of embryo. (A- 

G from Mori & Gracie 18928; H-I from Oldeman 4159).


. 

.7< ;- g WtW \ ; WrI -O 5 WI 40W I 

| Talisia setigera 

v Talisia stricta ........... 

' 

_z-a;l' 

* Talisia sylvatica 4 z~ ,.*Fii'."0 * 

( 

t 

stION 

,B ? , o '\/ 


as the petal, adaxially hirsute on upper half, abaxially and 4) fruits 1-2 cm long (vs. 1.8-2.5 cm). The speglabrous 

for most of its length; disc hispidulous to to- cific epithet refers to the forested habitat where the spementose 

at apex, 5-lobed, 0.5 mm tall; stamens 5(6), cies is found. 

the filaments pilose or densely so, of similar length, 2- 

3 mm long, the anthers oblong to ellipsoid, 1-1.2 mm 

long, apiculate at apex; ovary ovoid, sericeous, the stigma 

Phenology. Flowers from May to November and 

fruits from September to May. 

capitate, papillate. Fruit turning from green to yelloworange, 

ellipsoid, apiculate, ca. 2 cm long, puberulent, 

the pericarp coriaceous, granulose, ca. 1 mm thick, the 

endocarp glabrous. Seed solitary, ellipsoid, with creamyellow 

fleshy testa. Embryo with cotyledons lying 

slightly obliquely dorsiventrally to each other, the upper 

slightly larger than the lower one. 

Distribution and Ecology. (Fig. 107) Surinam, 

French Guiana, Brazil, and Peru, in lowland, tenra finne, 

gallery, and periodically flooded forests. 

Common names. French Guiana: boisflambeau, 

gaulette indien, paicoussa, tepu, touliatan, tepuime. 

Surinam: bosknippa. 

Talisia sylvatica is vegetatively similar to T Representative specimens examined. SURINAM. 

nervosa, however, T sylvatica differs by the following Without specific locality, s.d. (fl), Hostmann 1123 (K), 

characters: 1) leaflets elliptic or oblong-elliptic, charta- Hostmann 1124 (K); Forest Reserve, 29 Jul 1933 (fl), 

ceous, abaxially minutely pubescent, the apex acuminate, Lanjouw 307 (MO). Commewijne: Vic. Jande creek, 23 

abruptly acuminate or apiculate (vs. oblong, narrowly Jul 1953 (bd), Lindeman 4465 (NY). Marowijne: 

oblong, elliptic, narrowly elliptic, oblanceolate or less 

often falcate, chartaceous to coriaceous, abaxially glabrous, 

the apex long-acuminate or less often obtuse); 

2) petiolules minutely pubescent (vs. glabrous); 3) leaf 

rachis minutely pubescent, terete or sometimes obtusely 

angled, canaliculate or slightly compressed on distal 

portion (vs. glabrous, slightly flattened dorsiventrally, 

Moengo, Cottica, 23 Jul 1953 (fr), Cowan 38983 (NY, 

US); Lely Mts., 550-710 m, 28 Sep 1975 (fr), 

Lindeman et al. 490 (K, MO, NY, VEN); Lely Mts., 175 

kms SSE of Paramaribo, 500-700 m, 18 Oct 1976 (fr), 

Mori & Bolten 8534 (NY); Para, 1 Aug 1920 (fl), Pulle 

110 (K-2). 


Arataye river at Saut Parare, Oct 1984 (fr), Forget 376 

sharply angled toward the margins on distal portion); (CAY); Approuague river near Saut Grand Canori, 13 

2


Jul 1968 (fl), Oldeman T-32 (CAY-3); Upper Approuague Boeuf Mort, 300 m, 14 Jun 1988 (fl), Mori & Gracie 

river, vic of Parapou creek, 10 Oct 1968 (fr), Oldeman 18928 (NY); Monts La Fumee Oeste, 200-400 m, 31 

B-1911 (CAY); Saut Fini, 13 Sep 1968 (fr), Oldeman Mar 1983 (fr), Mori & Pipoly 15464 (NY-2); Between 

2811 (CAY-2); Arataye river at Saut Parare, 15 Aug 1977 Eau Noire creek & Monts Galbao, 2 May 1973 (bd), 

(fr), Sastre 5669 (CAY). Bassin de la Camopi: Saut Oldeman 3292 (CAY-2, P, US); Montagne Grand-Fosse, 

Ouasseye, along Camopi river, alluvial plateau, 13 Dec 28 Oct 1971 (fr), Oldeman 4159 (CAY, NY). 

1967 (fr), Oldeman 2671 (CAY-2, P). Bassin de la Comtk: PERU. Loreto: Puerto Almendras, 130 m, 16 Jun 

Montagnes Soufflet, vic. of Comte river, 10 Jun 1975 1990 (fr), Gentry et al. 71196 (US); Pefia Negra, on the 

(fl), de Granville B-5277 (CAY-2). Bassin de L'Inini: road from Iquitos past Quistococha, 7 Dec 1979 (fr), 

Palofini creek, 22 Aug 1970 (fl), de Granville C-40 (CAY, Jones 9683 (WIS); Mishuyacu, near Iquitos, 100 m, Oct- 

P); Montagne Bellevue de L'Inini, 600-700 m, 3 Sep Nov 1929 (fl), Klug 104 (F, NY, US), (fl), Klug 389 (F, 

1985 (fr), de Granville et al. 8028 (CAY-2, P, US). Bassin US); Santa Maria de Nanay, 10 km W of Mishana, along 

de la Mana: Route de L'Acarouany, 9 Feb 1990 (st), Nanay river, 130 m, 15 Mar 1991 (fr), Pipoly et al. 

Cremers & Hoff 11313 (CAY). Bassin du Maroni: St. 15028 (US); Quistococha, vic. Iquitos, 25 Feb 1977 (fr), 

Jean, 8 May 1914 (fl), Benoist 1193 (P-2); Upper Revilla 2408 (MO, NY); Road to Peiia Negra, trail to 

Maroni, Antecume Pata, 19 Nov 1977 (fr), Cremers Itaya river, 120-140 m, 20 Jan 1984 (fr), Rimachi 7309 

5084 (CAY-2, P, US), 5 Nov 1977 (fl), Cremers 4963 (US); Alpahuayo, IIAP Station, 19 Oct 1984 (bd), 

(CAY-2, P, US); Anapaike, 23 Jul 1996 (yfr), Fleury 958 Vasquez & Criollo 5780 (MO); Puerto Almendras, 

(US); Grand Inini river, 5 Sep 1970 (fr), de Granville Nanay river, 30 Oct 1984 (yfr), Vasquez & Jaramillo 

B-3677 (CAY, P); Maroni at Saut Badjeri, 8 Sep 1970 5875 (MO); Quebrada Santa Cruz, 18 Mar 1982 (fr), 

(fr), de Granville B-3736 (CAY, P), 7 Aug 1985 (fr), de 

Vasquez & Ruiz 2917 (MO, NY); Iquitos, 130 m, 23 

Granville et al. 7307 (CAY); Upper Maroni, Antecume 

May 1990 (fr), Vdsquez et al. 13741 (US). 

Pata, Oct 1976 (fl), Moretti 353 (CAY), 20 Apr 1975 (fr), 

BRAZIL. Amapi: Oiapoque, 5 Oct 1949(fr), Black 

Sastre et al. 3832 (CAY-2, P, US); Maroni, s.d. (fl), 

49-8388 (IAN). Amazonas: Moro da Seis Lagoas, 400 

Wachenheim 26 (P). Bassin du Sinnamary: Sinnamary, 

m, 20 Sep 1990 (st), Nelson 2428 (MO). Mun. Humaita, 

10 m, 24 Nov 1989 (fr), Schafer 9221 (CAY). Bassin 

estrada Huimata'-Porto Velho, km 60, 30 Apr 1982 (fr), 

Teixeira et al. 89 (INPA, K, MG, NY). 

de la Yaloupi: Yaroupi river, near Saut Coueki, 8 Apr 

1970 (st), de Granville 493 (CAY). Bassin de la Waki: 

Rivier Ouaqui-Grigel, 4 Jul 1973 (yfr), de Granville 

EXCLUDED TAXA 

1685 (CAY-2, P, US-2). Ile de Cayenne: Mont Grand Talisia diphylla Standley = Exothea diphylla 

Matoury, 13 Jul 2000 (st), Acevedo R. 11123 (CAY, US). (Standley) Lundell 

Piste de Saint Elie: Station Biologique ORSTOM, 1 May Talisia jimenezii Alain = Melicoccus jimenezii 

1992 (st), Acevedo R. 4875 (CAY, US), 8 May 1992 (Alain) Acevedo-Rodriguez 

(fl), Acevedo R. 4925 (US-2), 9 Sep 1992 (fr), Prevost Talisia laxiflora Bentham = Matayba guianensis 

3028 (CAY, US); St. Elie road, km 17, 4 May 1990 (fl), 

Aublet f. laxiflora (Bentham) Radlkofer 

Sanoja & Loup 3523 (CAY). Region Littorale: Kourou, 

Talisia oliviformis (H.B.K.) Radlkofer =Melicocca 

Passoura creek, 6 May 1992 (fl), Acevedo R. et al. 4911 

(CAY, US). Region Paul Isnard: Paul Isnard road, km 

oliviformis H.B.K. 

70, 7 Nov 1982 (fr), Feuillet 255 (CAY, P, US). Region 

Talisiapedicellaris Sagot ex Radlkofer =Melicoccus 

de Saul: Route de Belizon, Eaux Claires to Saul, ca. 2 pedicellaris (Sagot ex Radlkofer) Acevedo-Rodriguez 

km S, 26 May 1992 (fl), Acevedo R. & Angell 5028 (US); Talisia peruviana Standley = Simaroubaceae, 

Eaux Claires, sentier Botanique, ca. 700 m from entrance, undescribed genus. 

230 m, 22 May 1992 (bd), Acevedo R. et al. 4999 (CAY, Talisia porteriana Johnst. ex Croat (invalid name, 

US-2), 22 May 1992 (fr), Acevedo R. et al. 5001 (US), published in synonymy)= Matayba glaberrima 

3-5 km from entrance, 350 m, 24 May 1992 (bd), Radlkofer 

Acevedo R. et al. 5014 (US); Grand Boeuf Mort, 300 m, 

8 Dec 1977 (st), Cremers 3906 (CAY); Saul, 220 m, 23 

Jun 1988 (fl), Gentry et al. 63137 (MO); Monts Galbao, 

10 km W/SW of Sail, 420 m, 17 May 1973 (fl), de 

Granville 1664 (CAY-3, P); La Fumee-La Douane, 3 Apr 

Talisia prancei G. Guarim Neto = Matayba 

guianensis Aublet 

Talisia svensonii Standley = Cupania seemannii 

Triana & Planchon 

1982 (fr), de Granville 5049 (CAY-2); Plateau La Douane, 

30 Sep 1974 (fr), de Granville B-5176 (CAY-2); Saul, ACKNOWLEDGMENTS 

along tributary of Mana river, 380 m, 13 Nov 1997 (fr), 

Gustafsson 319 (CAY, NY); Montagne Grand-Fosse, 7 

Nov 1974 (fr), Jacquemin 1438 (CAY-2); Carbet Mais 

trail, between Mt. La Fumee & Crique Nouvelle France, 

300 m, 9 Sep 1989 (fr), Mori et al. 20868 (CAY, NY, 

US); Eaux Claires, 6 Nov 1997 (fr), Mori et al. 24669 

(CAY, NY), (fr), Mori et al. 24671 (CAY, NY); Grand 

This publication was made possible through the 

collaboration of numerous people and by the financial 

support of the Smithsonian Institution. 

I am indebted to the following people who either 

gave their time and advice or performed numerous tasks 

during the completion of this study: Mark T. Strong

LITERATURE CITED 167 

(US) for performing numerous technical tasks, includ- Botany for funding the botanical illustrations; The 

ing management of exsiccatae, data gathering and en- Neotropical Lowlands Research Program for fundtry, 

acetolysis of pollen, preparation of SEM stubs and ing most of the field work, field supplies and some of 

light slides, SEM microscopy, leaflet x-rays, prepara- the field equipment; The Biological Dynamics of Fortion 

of image plates, proof reading, writing figure cap- est Fragments Program for providing financial suptions, 

and editing; Bobbi Angeli (NY) for rendering port for my field work in Amazonas, Brazil; and the 

precise and artistically pleasing illustrations, and for Biological Diversity of the Guianas Program for supaccompanying 

me in the field; Gerardo Aymard porting my field work in the Tumuc Humac region of 

(PORT) & Paul E. Berry (WIS) for their collabora- French Guiana-Surinam-Brazil. 

tion during field work in southem Venezuela; German I am also indebted to the curators of the following 

Carnevali, Jose Luis Tapia, and Filogonio May Pata herbaria for loaning their valuable collections, for send- 

(CICY) for their valuable collabortation during field ing numerous specimens as gift for determination and 

work in the Yucatan Peninsula; Jose A. Cedefio for or photos of type specimens and for valuable discushis 

assistance in the field; Ricardo Callejas (HUA) sions on the status of their collections: BM, BOL, BR, 

for facilitating field work and collecting permits; Dan 

Cole for helping generate the distribution maps; Ellen 

Far (US) for converting the coordinate data into a usable 

format for generating the distribution maps; 

C, CAY, CICY, COAH, COL, CVRD, F, HUA, IAN, 

INPA, JAUM, JBSD, K, M, MA, MEDEL, MER, MG, 

MO, NY, P, PORT, QCA, R, RB, U, UPR, USM, VEN. 

Christiane Feufllet (US) for his valuable help in translating 

French text; Vicki Funk (US) for her valuable 

help and discussions during the performance of the 

cladistic analyses; Ricardo Garcia and Melciades 

ABBREVIATIONS 

bd: flower buds 

yfr: young fruit 

Mejias (JBSD) for their assistance in the field and 

granting of collecting permits; Carol Gracie (NY) for LITERATURE CITED 

providing numerous Talisia photos, two of which are Acevedo-Rodriguez, P. 1990. The occurrence of 

piscicides and stupefactants in the plant kingdom. 

published in this monograph; Jean Jacques de Adv. Econ. Bot. 8: 1-23. 

Granville (CAY) for planning and collaborating on an . 1993. Systematics of Serjania (Sapindaceae). 

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Londofio for facilitating fieldwork in Colombia; Scott New York Bot. Gard. 67: 1-93. 

A. Mori (NY) for his support during field work in Saul, 

. . 1996. Flora of St. John, U.S.V.I. Mem. New 

York Bot. Gard. 78: 1-581. 

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discussions and advice on typification, nomenclature, nueva combinacion basada en Talisia jimenezii, 

derivation of Latin names, and for reviewing the Latin 

diagnoses; Joan Nowicke (US) for her valuable discussion 

on pollen morphology; Marie F. Pr6vost and 

Daniel Sabatier (CAY) for their assistance and hosespecie 

endemica de la Repu'blica Dominicana. 

Moscosoa 9: 58-61. 

. 1998. Novelties in Neotropical Sapindaceae II. 

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the Caribbean. Pp. 13-39. In: S. K. Donovan & T.A. of the Eocene Green River formation. Bull. Torrey 

Jackson (eds.), Caribbean Geology: An Introduction. Bot. Club 60: 479-524.

LIST OF EXSICCATAE 169 

1. MELICOCCUS 

1. Melicoccus espiritosantensis Acevedo-Rodriguez 

2. Melicoccus pedicellaris (Sagot ex Radlkofer) 

Acevedo-Rodriguez 

3. Melicoccus petiolulatus Acevedo-Rodriguez 

4a. Melicoccus oliviformis (H.B.K.) Radlkofer subsp. 

oliviformis 

4b. Melicoccus oliviformis subsp. intermediaus 

(Radlkofer) Acevedo-Rodriguez 

5. Melicoccus antioquensis Acevedo-Rodriguez 

6. Melicoccus novogranatensis Acevedo-Rodriguez 

7. Melicoccus aymardii Acevedo-Rodriguez 

8. Melicoccus lepidopetalus Radlkofer 

9. Melicoccus bijugatus Jacquin 

10. Melicoccus jimenezii (Alain) Acevedo-Rodriguez 

2. TALISIA 

1. Talisia angustifolia Radlkofer 

2. Talisia chartacea Acevedo-Rodriguez 

3a. Talisia clathrata Radlkofer ssp clathrata 

3b. Talisia clathrata ssp canescens Acevedo-Rodriguez 

4. Talisia coriacea Radlkofer 

5. Talisia esculenta (Cambessedes) Radlkofer 

6. Talisia firma Radlkofer 

7. Talisia floresii Standley 

8. Talisia furfuracea Sandwith 

9. Talisia granulosa Acevedo-Rodriguez 

10. Talisia japurensis (C. de Candolle) Radlkofer 

11. Talisia lanata Acevedo-Rodriguez 

12. Talisia microphylla Uittien 

13. Talisia parviflora Acevedo-Rodriguez 

14. Talisia praealta Sagot ex Radlkofer 

15. Talisia simaboides Kramer 

16. Talisia squarrosa Radlkofer 

17. Talisia subalbens (Martius) Radlkofer 

18. Talisia velutina Acevedo-Rodriguez 

19. Talisia veraluciana G. Guarim Neto 

NUMERICAL LIST OF TAXA 

LIST OF EXSICCATAE 

Abbott, W. L., 2427(1-9). 

Abraham, 219(2-16). 

Acevedo-Rodriguez, P., 4868(1-2); 4875(2-52); 

4879(2-34a); 4882(2-14); 4885(2-36); 4890(2-12); 

4925(2-52); 4939(1-2); 4940(2-34a); 11117(2-16); 

11125(2-40). 

Acevedo-Rodriguez, P. & B. Angell, 5021; 5025(2- 

3(2-3b); 5028(2-52). 

Acevedo-Rodriguez, P. & R. Callejas, 6772(2-24). 

Acevedo-Rodriguez, P. & J. A. Cedenio, 7380(2-39); 

7559; 7601(2-24). 

Acevedo-Rodriguez, P. & C. Clubbe, 10956(1-9). 

Acevedo-Rodriguez, P. & J. Grimes, 4797(2-22); 

4798(2-39); 4801; 4803; 4837(2-22); 4863(2-8); 

4867(2-15); 4877(2-40); 4863(2-8). 

Acevedo-Rodriguez, P. & J. L. Tapia, 12234(2-7). 

Acevedo-Rodriguez, P. et al., 1585(2-47); 3473; 3512; 

3515(1-4b); 4823(2-40); 4828(2-37); 4842(2-22); 

4844(2-40); 4857(2-15); 4871(2-32); 4872(2-22); 

4873(2-14); 4874(2-8); 4888(2-39); 4905(2-14); 

20. Talisia acutifolia Radlkofer 

21. Talisia bullata Radlkofer 

22. Talisia carinata Radlkofer 

23. Talisia caudata Steyermark 

24. Talisia cerasina (Bentham) Radlkofer 

25. Talisia croatii Acevedo-Rodriguez 

26. Talisia cupularis Radlkofer 

27. Talisia dasyclada Radlkofer 

28. Talisia douradensis Acevedo-Rodriguez 

29. Talisia equatoriensis Acevedo-Rodriguez 

30. Talisia eximia Kramer 

31. Talisia ghilleana Acevedo-Rodriguez 

32. Talisia guianensis Aublet 

33. Talisia hemidasya Radlkofer 

34a. Talisia hexaphylla Vahl subsp. hexaphylla 

34b. Talisia hexaphylla subsp. elegans Acevedo-Rodriguez 

34c. Talisia hexaphylla subsp. multinervis Acevedo- 

Rodriguez 

35. Talisia laevigata Acevedo-Rodriguez 

36. Talisia longifolia (Bentham) Radlkofer 

37. Talisia macrophylla (Martius) Radlkofer 

38. Talisia marleneana (G. Guarim Neto) Acevedo- 

Rodriguez 

39. Talisia megaphylla Sagot ex Radlkofer 

40. Talisia mollis Kunth ex Cambessedes 

41. Talisia morii Acevedo-Rodriguez 

42. Talisia nervosa Radlkofer 

43. Talisia obovata A.C. Smith 

44. Talisia oedipoda Radlkofer 

45. Talisia pachycarpa Radlkofer 

46. Talisia pilosula Sagot ex Radikofer 

47. Talisia pinnata (Ruiz & Pav6n) Radlkofer 

48. Talisia princeps Oliver 

49. Talisia retusa Cowan 

50. Talisia setigera Radlkofer 

51. Talisia stricta (Karsten & Triana) Triana & Planchon 

52. Talisia sylvatica (Aublet) Radikofer 

4911(2-52); 4914(2-40); 4915(2-22); 4916(2-32); 

4920(2-22); 4922(2-32); 4941; 4942(2-15); 

4953(2-40); 4954(2-33); 4956(2-3(2-3b); 

4957(2-39); 4960; 4966(2-33); 4967; 4989(2-37); 

4996(2-3(2-3b); 4999; 5001(2-52); 5012(2-3(2-3b); 

5014(2-52); 5019(2-3(2-3b); 5030(2-37); 5441(1-9); 

5935(2-37); 6127(2-36); 6145(2-37); 6909(2-22); 

7963; 7988; 8053(2-6)(2-6); 8150(2-38); 8218(2-37); 

8362(2-20); 8390(2-6)(2-6); 8414; 8442(2-20); 

8448(2-6)(2-6); 8456(2-20); 8477(1-10); 9729(2-39); 

10242(2-37); 10286(2-42); 12200(2-7). 

Adis, J., 7(2-20). 

Aguilar, M., 373(1-4a). 

Albin, C., J., 2396(2-34a). 

de Albuquerque, B., 1098; 1110(2-31). 

Allen, P., 5310(2-25); 5917(2-37). 

Alvarez, E., et al., 286(2-37). 

Alvaro, P., 721(2-7). 

Amaral, I. L. et al., 543(2-3(2-3a); 579(2-6)(2-6); 

1296(2-38).


Amorim, A. M. et al., 1103(2-24). 

Brand, J. & M. Escobar, 713(2-34b). 

van Andel, T., 26SIS5(2-3(2-3b). 

Brand, J. & M. Gonzalez, 518(2-34b); 529(2-37); 

Anderson, W. R. & D. C. Stemnberg, 3006(1-9). 961(2-34b). 

Andrade-Lima, 52-1190(2-49); 70-6231(2-5). Brand, J. & J. Lozano, 923(2-34b). 

Andre, E., 582; 583(2-51). 

Anonymous, 226 M; 227 M; 230 M; 245 M; 265 M(2- 

Brand, J. & M. Narvaez, 622(2-37). 

Brand, J. et al., 1313(2-37). 

16); 330(2-39); 374(2-22); 572(1-4b); 889(2-16); Breteler, F. J., 3936(2-4). 

3633(2-34a); s.n.(2-22); s.n.(2-48); MG-9625(2- 

36); MG-9743(2-5). 

Araquistain, M., 3137(2-42). 

Archer, W. A., 2309; 2378; 3013(1-4a). 

Arciria, A., 169(2-24). 

Aristeguieta, L., 3825(1-4a); 5994(2-34a). 

Arnason, T. & J. Lambert, 17065(1-4a). 

Arroyo, L. et al., 141(2-5); 282(2-24). 

Asplund, E., 5412(2-50). 

AssungAo, P. A. C. L., 76(2-19); 344a(2-26); 854(2-37). 

Assungao, P. A. C. L. & E. C. Pereira, 261(2-10); 

800(2-26). 

Assun9ao, P. A. C. L. et al., 315a(2-26); 538(2-9). 

Aubreville, 258(2-39). 

Bristan, N., 184(2-34b); 1511(2-37). 

Britton, N. L. & E. G. Britton, 10138(1-9). 

Britton, N. L. & J. F. Cowell, 10(1-9). 

Broadway, W. E., 5237(1-4a); 5598; 10138(2-34a); 

s.n.(1-4a); s.n.(1-9). 

Bunting, G. S. et al., 3827(2-37). 

Burchell, 6195(2-1); 7971(2-5). 

Burger, W. & G. Matta U., 4757(2-24). 

Burgos L., J. A., 10(2-42). 

Byron, J. L. & Elias, 67-51(2-13). 

Cabrera, E. & H. de Cabrera, 4652; 6446(1-4a); 

6569(2-7); 11437(1-4a). 

Cabrera, I., 3329(2-3a). 

Cadamuro, L. & Solacroup, 173(2-32). 

Aublet, s.n.(2-32); s.n.(2-52). 

Cain, S. A., 59(2-25). 

Aulestia, C. et al., 641(2-29). 

Calder6n, S., 1526(1-9). 

Aulestia, M., 110(2-24). 

Callejas, R. et al., 9771(2-24). 

Aulestia, M. & J. Andi , 908(2-24). 

Calzada, J. I., 3206(1-4a). 

Aulestia, M. & G. Grefa, 123(2-39); 167; 192(1-6). Cirdenas, D., 485(2-37). 

Ayala, F., 2004(2-6)(2-6). 

Cardenas, D. & J. G. Ramirez, 2566(2-24). 

Ayala, F. et al., 2539(2-39); 3908; 5822(2-24). Cardenas, D. et al., 2700(2-41); 2914(2-24). 

Aymard, G. & L. Delgado, L., 7915(2-37); 8409(2-43); Cardona, F., 2126(1-4a). 

845 8(2-6)(2-6). 

del Carpio, 2016(2-47). 

Aymard, G. & A. Fernindez, 7176(2-2); 7327(2-37). Carriera, L. et al., 526; 540(2-43) 

Aymard, G. & F. Ortega, 2521(1-7). 

de Carvalho, A. M. et al., 3624(2-24). 

Aymard, G. et al., 4154(2-37); 9808(2-43); 10148(2- Cavalcante, P., 1840(2-19). 

49); 1023 1; 10271(2-34a). 

Cavalcante, P. & M. Silva, 1674(2-24); 1675(2-24); 

Bahia, R. P., 61; 91(1-2). 

2775(2-38). 

Bailey, L. H. & E. Z., Bailey 814(1-9). 

Chagas, J., s.n. herb. 127(2-10); s.n. herb. 1798(2-31); 

Baldwin, J. T. Jr., 3290(2-20). 

s.n. herb. 3171(2-31). 

Balee, W., 2185(I-40) 

Chaves, D., 171(1-9). 

Balslev, H. & W. C. Steere, 3101(2-50). 

Clark, H. L. & P. Maquirino, 7931(2-6)(2-6). 

Barbour, P. J., 5720(2-25). 

Clarke, D., 2015(2-33). 

Barrier, S., 4200(2-40); 4228a(2-39); 4263(1-2). Clarke, J. et al., 2650(2-29). 

Barrier, S. & C. Feuillet, 2589(2-37). 

Claussen , P., 500(2-5); s.n.(2-5). 

Basualdo, I., 3683(1-8); 3830(2-5). 

Coelho, L. F., 156(2-26); 495(2-6); 500(2-27); 1205; 

Bayron, 68-111(2-37). 

1667(2-31); s.n. herb 4130(2-10). 

Beck, S. G., 1337(2-5); 5171; 20054(2-24); Coelho, L. F. & D. Coelho, 4000(2-26). 

Benoist, R., 54(2-16); 337(2-39); 789(2-42); 1193(2-52). Coelho, L. F. et al., 3922; s.n. herb. 21254(2-19). 

Berg, C. C. et al., 548(2-40); P-18388(2-43). Coelho, L. F. & 0. P. Monteiro, 29(2-43). 

Bernardi, L., 1147(2-14); 6536(2-34a); 7998(2-33); Cogollo, A., 974, 1568(1-5); 1823(2-34a). 

19128(2-5). 

Cogollo, A. & D. Cardenas, 4476(2-34a). 

Billiet, F. & B. Jadin, 4439(2-22). 

Cogollo, A. et al., 4572(2-22). 

Black, G. A., 49-8388(2-52); 51-13951(2-39); 54- Colaris, W. J. J., 1086(2-1). 

177 14(2-5). 

Colella, M. & E. Guayamare, 2138(2-6). 

Blanc, M., 120(2-22). 

Contreras, E., 375(2-7); 767; 5876(1-4a); 6991(2-7). 

Blanco C., 48; 303; 368(2-33); 652(2-35); 791; Contreras, J., 94(2-23). 

930(2-34a). 

Cook, J., s.n.(2-48). 

Boom, B. & S. A. Mori, 1610(2-15); 1633(2-36); 1688; Cook, M. T., 41(1-9). 

2379(2-15); 2255(2-3(2-3b); 2416(2-33). Cordeiro, I., 151; 215(2-37); 318(2-6). 

Boos, 20(1-4b). 

Cornejo, X. & C. Bonifaz, 3878(2-50); 5720(36). 

Bordallo, A., s.n. MG-22655(2-24). 

Correa A., M. & R. Dressler, 629(2-42). 

Bordenave, B., 832(2-46); 1013(2-14). 

Couret, 301(2-33). 

Boschwezen, 231; 330(2-39); 2574(2-33); 3538; Cowan, R. S., 38296; 38491(2-36); 38795(2-22); 

5956(2-39); 6445(2-12); 6575; 6595(2-40). 38809(2-32); 38983(2-52). 

Bossio, H., s.n.(2-6)(2-6). 

Crandlemire-Sacco, J. & R. Sacco, 82(2-25). 

Box, H. E., 1442(1-9). 

Cremers, G., 3906(2-52); 4243; 4689(2-22); 4963; 5084 

Brand, J., 938(2-34b); 1204(2-34b). 

(2-52); 5434(2-40); 7275; 7961(2-22); s.n.(2-22). 

Brand, J. & A. Cogollo, 79(2-34b). 

Cremers, G. & J. J. de Granville, 14113(2-37).


Cremers, G. & M. Hoff, 11313(2-52). 

Cremers, G. et al., 13854(2-39); 14360; 14361(2-22). 

Croat, T. B., 6498; 8095; 8236; 8782; 8800(2-42); 

8820; 9034(2-25); 10873(2-42); 11270; 12494(2- 

25); 14027; 15251(2-42); 15366(2-25); 16376(2- 

48); 18550; 18759(2-3a); 19160(2-34c); 20433(2- 

48); 34375; 34511; 35123(2-25). 

Croat, T. B. & M. H. Grayum, 59760(2-37). 

Croizat, L., 44(2-2). 

Crosby, M. R. et al., 204(1-9). 

de la Cruz, J. S., 3475(2-16). 

Cuadros, H., 2176(2-42). 

Cuatrecasas, J., 17010(2-42). 

L. da Cunha N. M. & C.L. Assunsao, 214a(2-45). 

Curran, H. M., 30(2-5); 84(1-9); 203(2-25); 704(1-4a). 

Curran, H. M. & M. Haman, 723; 781(1-4a). 

D'Arcy, W. G., 3916(2-42). 

Dahlgren, B. E. & E. Sella, 120(2-38). 

Daly, D. C. et al., 635(2-5); 872; 1040; 1245(2-36); 

1672(2-33); 1768(2-49); 3822(2-22); 3951(2-40); 

4345(2-37); 7571(2-3a); 8256(2-34a); 9497(2-24). 

Damiao, C., 2742(2-24). 

Davidse, G., 27875; 27960; 27999(2-6). 

Davidse, G. & A. Gonzalez, 16385(2-33). 

Davidsom, C., 3616(2-3a). 

Davidson, C. & Martinelli, 10073(2-38). 

Davis, T.A.W., 359; 501(2-12). 

Dean, R. E., 13326(2-34a). 

Defler, S., 540; 541(2-6). 

Delascio, F. et al., 9572; 9695(2-24). 

Delgado, L., 587; 689(2-6). 

Dias, B. J. et al., 117(2-5). 

Diaz, C. et al., 583(2-10). 

Dik, A., 721; 1179(1-6). 

Dodson, C. H., 5236; 5893; 5998(2-50); 8420(2-29). 

Dodson, C. H. & D. Benzing, 13911(2-29). 

Dodson, C. H. & A. Gentry, 9798(2-50). 

Dodson, C. H. & J. Torres, 2924(2-3a). 

Dodson, C. H. et al., 7989(2-50); 14634(2-29). 

van Donselaar, J., 3047(2-30); 3762(1-2). 

Ducke, A., 424; 1634(2-5); 2875; 3037; 3049(2-24); 

6945; 7940(2-40); 8507(2-38); 8703(2-24); 

8886(2-38); 8887(2-40); 9049(2-26); 10205(2-24); 

10488(2-19); 11462(2-24); 17176(2-4); 20902(2-24); 

s.n., RB-18901(2-19); s.n., RB-18906(2-10); s.n., 

RB-25218; s.n., RB-25219(2-37). 

Ducke, A. & Andrade-Lima, 25(2-24). 

Dugand, A., 6190; 6570; 6572; 6651(1-4a). 

Dugand, A. & R. Jaramillo, 2782; 4120(1-4a). 

Dugand, A. & P. Standley, 375(1-4a). 

Duke, J. A., 10024(2-42); 11074(2-25); 14578(2-37); 

15250(2-42). 

Duke, J. A. & N. Bristan, 431; 8183(2-25). 

Duran, R., 2133(1-4a). 

Duss, P., 3718(1-9). 

Dwyer, J. D., 1548; 2332; 6940(2-42). 

Dwyer, J. D. & A. Robyns, 140(2-42). 

Ebinger, J. E., 240; s.n.(2-42). 

Edwards, K. S. et al., 335(2-34a). 

Eggers, 14704(2-50); 15004(2-22); 15764(2-39). 

Egler, W. A., 585(2-38). 

Egler, W. A. & H. S. Irwin, 45955; 45995(1-2). 

Ehrendorfer, 74104-23(2-37). 

Ekman, E. L., H-8033; 9480; 19447(1-9). 

Elias, Br., 1508(1-4a). 

Enriquez, 0. G., 356(1-4a); 392(2-7); 514(1-4a). 

Espinal T., S., 1201(2-34a). 

Evans, R. et al., 1924(2-39). 

Faber-Langendoen, D. & J. A. Hurtado, 1496(2-37); 

1934(2-42). 

Faber-Langendoen, D. & E. Renteria, 1258(2-37). 

Fanshawe, D. B., 251; 576(2-8); 1215(2-16); 1982 

(2-12); 2046(2-33); 2718(2-37); 2955(2-32); 

3402(19); 6285(2-32). 

Felippe, G. M., 43(2-1). 

Fendler, A., 2477(1-4a). 

Fernandez, A., 2107(2-24); 6586(1-4a). 

Fernandez Alonso, J. L. et al., 5484(2-51). 

Femrandez Perez, A., 5283; 5319(1-4a). 

Cid Ferreira, C. A., 7495; 7591(2-45). 

Cid Ferreira, C. A. & B. W. Nelson, 2575(2-24). 

Cid Ferreira, C. A. et al., 423(2-26); 689(2-14); 822(1- 

2); 888(2-38); 953(2-31); 964(2-26); 1637; 1655(2- 

38); 1660(2-40); 1666; 1892; 2359; 2499(2-38); 

3114; 4805; 4905(2-49); 6036(2-45); 6524(2-49); 

6556(2-17); 6676(2-37); 6732(2-26); 6952(2-37); 

7597(2-10); 7720(2-45); 8023; 8209(2-26); 8565; 

9865; 9970(2-37); 10932(2-24). 

Ferreira, L. V., 105; 152(2-20). 

Ferreyra, M., 1(1-4a). 

Ferreyra, R., 4749(2-34a). 

Ferrucci, M.S., 69(1-8). 

Feuillet, C., 255(2-52); 698(2-22); 1046(2-39); 1221(2- 

22); 2318(2-39); 3692(2-22); 10087(2-3b). 

Fiebrig, K., 4178(2-5). 

Fleury, M., 100(2-40); 758(1-9); 958(2-52). 

Florence, J., 4047(1-9). 

Foldats, E. & J. Velazco, 9249(2-24). 

Folli, D. A., 30(1-1); 416(1-4b); 1043(2-24). 

Folsom, J. P. et al., 6857(2-25). 

Fonnegra, R. et al., 2795(2-34b); 4815(1-9). 

Forero, E. & Jaramillo, 2758(2-37). 

Forero, E. et al., 9976(1-4a). 

de Foresta, 146; 341; 419(2-32). 

Forest Department British Guiana, 3849(2-40); 5355; 

6966(2-12); 7305(2-40). 

Forget, 376(2-52). 

Fosberg, F. R., 56774; 60936(1-9). 

Foster, R., 1619(2-42); 4429(2-3a); 8714(2-25); 

9747(2-34c). 

Franco, P. et al., 2036(2-25). 

Freire, F. M. T., 3709(2-5). 

Vieira Freire, 100(2-5). 

Freitas, C. A. A. et al., 276(2-38). 

Freitas, M. A. & R. M. Cardoso, 14(2-26). 

Froes, R.L., 22187(2-42); 23108(1-4b); 25002(2-26); 

25807(2-32); 26617(2-22); 29610a(2-26); 

32186(2-19). 

Froes, R. L. & B. A. Krukoff,11510(2-49). 

Funk, V. A. et al., 8109(2-25). 

Galeano, G. & A. Mirafia, 1720(2-10). 

Garcia, A., 1063(1-9). 

Garcia, R. & F. Jimenez, 3861(1-10). 

Garcia R. & N. Ramirez, 3930(1-10). 

Garcia R. et al., 5773a(1-10). 

Garcia Barriga, H., 13488(1-9). 

Gardner, 951; 1501; s.n.(2-5). 

Garibaldi, C., 106(2-25). 

Garz6n N. C. et al., 145(2-11). 

Gaumer, G. F., 406; 1749; 24137; 24189; 24446(1-4a). 

Gaumer, G. F. et al., 23577; 23635(1-4a). 

Geay, F., 852; 993(2-22).


Gentry, A., 3165; 6603(2-42); 13454(2-25); 49083 

(2-19). 

Gentry, A. & P. Berry, 14829(1-4a). 

Gentry, A. & H. Cuadros, 57395(2-42); 68178(1-4a). 

Gentry, A. & J. Dwyer, 3492(2-42). 

Gentry, A. & L. Emmons, 39627(2-24). 

Gentry, A. & B. Nelson, 69247(2-20). 

Gentry, A. & R. Ortiz, 79870(1-9); 79889a(l-4a). 

Gentry, A. & L. Puig-Ross, 14246(1-4a). 

Gentry, A. & E. Renteria A., 23991; 24291(2-42). 

Gentry, A. & B. Stein, 46420(2-6); 46473; 46857(2- 

37); 47010; 47377(2-6). 

Gentry, A. et al., 19029(2-42); 24875(2-37); 25360(2- 

42); 26063(2-10); 26114(2-6); 30216(2-37); 41623; 

41729(2-43); 42476(2-6); 43173(2-34c); 43759(2- 

3a); 51136(2-49); 55700(2-37); 55713(2-24); 

56030(2-3a); 60659; 60665; 60709(1-4a); 62957(2- 

37); 62981(2-39); 63037(2-36); 63091(2-40); 

63137(2-52); 63320(2-37); 63470(2-43); 65295(2- 

37); 65604(2-3a); 65695(2-26); 71196(2-52); 

76186(2-25); 77167(2-49). 

Glaziou, A., 3757(2-26); 6491; 8309; 10424(2-5); 

13616(2-26); 15447(2-39); 20862(2-5). 

Goldman, E. A., 747(1-4a). 

Goulding, M., 174(2-24). 

Grindez, C. & R. Vasquez, 240; 403(2-24). 

de Granville, J. J., 371(2-37); 376(2-22); 493(2-52); 

512(2-22); 1664; 1685(2-52); 3150(2-39); 3738; 

4475; 4710(2-22); 4880(2-36); 5049(2-52); 5235; 

6952(2-22); 7033(2-36); 7161; B-3677(2-52); B- 

3695(2-33); B-3736(2-52); B-4222(2-40); B-5074 

(2-33); B-5176; B-5277(2-52); B-5507(2-22); C- 

40(2-52); C-54(2-39); C-148; T-1169; T-1205(2-22). 

Herrera, H. et al., 981(2-41). 

Heyde, N. M. & J. C. Lindeman, 131(2-39). 

Hladick, 361(2-42). 

Hoehne, 4507(2-5). 

Hoffman, B. & R. Foster, 3599(2-49). 

Hohenkerk L. S., 3a(2-34a); 715(2-16). 

Holdridge, L. R., 2537(2-42). 

Hostmann, 1123; 1124(2-52); 1149(2-39); 1274(2-33). 

Howard, R. A. & E. S. Howard, 19449(1-9). 

Huashikat, V., 1595(2-39). 

Huber, J., 9(2-36). 

Idrobo, J. M., 8091(2-37); 8497(1-6). 

Irvine, D., 576(2-48); 1012(2-25). 

Irwin, H. S., 48680(2-40); 48688(2-22); 48803(2-39). 

Irwin, H. S. & T. Soderstrom, 7451; 7581(2-1). 

Irwin, H. S. et al., 47325(2-39); 47630(2-22); 

47853(2-39); 48154; 48242; (2-22); 54600; 

54729(2-33); 54770(2-39); 55015(2-33); 

55046(2-39); 55885(2-36). 

de Granville, J. J. & Cremers, G., 12752(2-22). 

Jack, J. G., 4321; 4366; 5151(1-9). 

Jacquemin, H., 1438(2-52). 

Jangoux, J. & R.P. Bahia, 562 (2-40). 

Jansen-Jacobs, M. J., 1134(2-8). 

Jansen-Jacobs, M. J. et al., 536; 537; 5596(1-4b). 

Jaramillo, N. & M. Jaramillo, 981(2-24). 

Jaramillo, N. et al., 1306(2-39). 

Jardim, J. G., 952(2-5). 

Jardim, J. G. et al., 1071(2-26); 1962(2-24). 

Jativa, C. & C. Epling, 636, 1057(2-50). 

Jenman, 4925(2-33). 

Jimenes-Saa, H., 1571(2-12). 

Jimenez, A., 3888(2-42). 

Jimenez, J.J., 2703; 3312(1-10). 

Johnston, I. M., 1557; 1778(2-42). 

de Granville, J. J. et al., 5903(2-40); 6093(2-32); 6516(2- Johnston, J. R., 299(1-4a). 

33); 6536(2-22); 7307(2-52); 7384(2-39); 7500(2- Jones, J., 9683(2-52). 

22); 7899(2-36); 7919(2-22); 8028(2-52); 8132; J6rgensen, P., 4594(2-1). 

8890; 10439; 10480; 10537; 10657; 12634(2-22). Josse, C., 698(2-50). 

Grenand, P., 58(2-22); 243; 364(2-40); 618(2-22); 

2797(2-24). 

Grijalva et al., 556(2-37). 

Grueger, 211(2-34a). 

Guanchez, F., 383(2-23). 

Guevara & Rojas, 357(2-34a). 

Guillen, R., 1954(2-5). 

Guillen, R. & S. Coria, 2005(2-5). 

Guppy, N. G. L., 361(2-34a); 538(2-19); 674(2-37). 

Gustafsson, M. H., 319(2-52). 

Gutierrez, G. V., 2582(2-24). 

Halle, F., 2756(2-32); 2773(2-40). 

Hammel, B. & G. Schatz, 11578(2-42). 

Harley, R. M., 18206(2-26). 

Kayap, R., 761(2-24). 

Kenoyer, L. A., 426(2-42); 646(2-25). 

Kernan, C. & P. Phillips, 905(2-42). 

Killeen, T. & A. Jardim, 7424a(2-5). 

Killeen, T. et al., 3244(2-34a); 3527(2-24); 7033(2-5). 

Killip, E. P. & A. C. Smith, 14349(1-9); 26946; 27024 

(2-3a); 27762(1-3). 

Killip, E. P. et al., 38246; 38319(2-51). 

Klug, G., 18(2-3a); 104; 389(2-52); 811; 1044; 1160; 

1558; 2509(2-3a); 2798(2-24); 4303(2-37). 

Knapp, S., 8351(2-37). 

Knapp, S. et al., 1703(2-42). 

Krukoff, B. A., 4750(2-43); 4840(2-25); 4981(2-43); 

5630; 5725(2-34a); 5787(2-25); 5960(2-10); 5963; 

Harley R. M. et al., 10982; 10984(2-49). 

6504(2-24); 6886(2-19); 7962; 7982; 8463(2-26); 

Harling, G. & L. Andersson, 24900(2-50). 

Hartshorn, G. et al., 2732(2-43). 

8587(2-40); 8603(2-24). 

Kuhlmann, J. G., 294(1-4a). 

Hassler, E., 7244(1-8); 7413; 7413a(2-5); 9503(2-1); 

12221; 12221a(1-8). 

Hatschbach, G., 13002; 15911; 37393(2-1); 37543(2- 

17); 37742(2-1). 

Hatschbach, G. & R. Callejas, 47313(2-1). 

Hatschbach, G. & Cervi, 52622(2-1). 

Hatschbach, G. & Silva, 64141(2-5). 

Haught, O., 1608(2-34a); 4012; 4035(1-4a); 6353(2-51). 

HBK, 1490(1-4a). 

Henderson, A. et al., 415(2-37). 

Herrera, H., 1311(2-24). 

Kuhlmann, M., 59056(2-1). 

Kuhlmann, M. & S. Jimbo, 349(1-2). 

Kuntze, O., 1737(1-4a) 

Kvist, L. & E. Asanza, 40820(2-42). 

Lanjouw, J., 307(2-52); 812(2-33). 

Lanjouw, J. & J. Lindeman, 388(2-39); 2218(2-14); 

2523(1 -2). 

Lanna Sobre, J. P., 1155(2-4). 

Larpin, D., 986(1-2). 

Laughlin, R., 911(1-4a). 

Leblond, 448(2-40).


Le6n, H., 425; 721(2-37). 

Leonard, E. C., 4175(1-9). 

Le Prieur, s.n.(2-32). 

Lescure, 169(2-22); 380(2-40); 893(2-22). 

Liesner, R., 4162(2-20); 6179(2-43); 7222(2-37); 7439; 

8687(2-20); 8866; 8883(2-6); 25739(2-42). 

Liesner, R. & H. L. Clark, 8906(2-20). 

Liesner, R. & A. Gonzailez, 9151(1-4a); 11116; 

11151(2-49); 11327(2-34a). 

Liesner, R. & B. Holst, 20394(2-42); 21840(2-37). 

Liesner, R. et al., 20935(2-42). 

Lima, J., 654(2-26). 

Lindeman, J. C., 3828(1-2); 4465(2-52); 4886(1-2). 

Lindeman, J. C. & J. de Haas, 4504(2-1). 

Lindeman, J. C. et al., 72(2-30); 137(2-40); 406(2-33); 

490(2-52); 501(2-22); 523(2-39). 

Lindman, C. A. M., 2205(1-8). 

Lino, A. M., 105(1-4b). 

Liogier, A., 35365; 35626 (1-4a). 

Liogier, A. & P. Liogier, 25442(1-10). 

Liogier, A. et al., 8247(1-10). 

Lisboa, P. & U. Maciel, 4452(2-24). 

Lisboa, P. et al., 1560(2-37). 

Little, E. L. Jr., 7160; 13604(1-9); 17598(2-34a); 

17676(2-33); 21517; 21968; 22039; 23767(1-9). 

Lohmann, L. G., 1-22(2-26). 

L6pez Palacios, S., 1913(1-4a). 

Loubry, D., 135(1-2); 163(2-22); 1894(2-40); 

1956(1-9); 2322; 2373; 2377; 2531(2-16). 

Loureiro, A et al., 37737; 48443(2-37); s.n. herb. 

35821(2-4); s.n., herb. 37878(2-37); s.n.(2-13). 

Lourenco, 571(2-4). 

Lowrie, S. R. et al., 199(2-24); 684(2-49). 

Lozano C., G. & R. Schnetter, 2825(1-4a). 

Lundell C. L., 558; 638(1-4a); 917(2-7); 1376(1-4a); 

3121; 3185(2-7). 

Luschnath, s.n.(2-4); s.n.(2-37). 

Maas, P. J. M., 10818(1-22). 

Maas, P. J. M. et al., 2196(2-22); 5973(2-34a). 

Macedo, A., 4048(2-5). 

Macedo, A. et al., 3331(2-17). 

Macedo, M., 1412(2-19). 

Maciel, U. N. et al., 576 (2-12). 

Maguire, B., 24340; 24341; 24690(2-33); 24797(2-36). 

Maguire, B. & D. Fanshawe, 23510(2-16). 

Maguire, B. & L. Politi, 28615(2-23). 

Maguire, B. & J. J. Wurdack, 34869(2-6). 

Maguire, B. et al., 42590(2-37); 56031(2-36); 

56058(1-2). 

Malme, G. 0. A., 962(1-8); 2300(2-5). 

Manara, B., s.n.(2-34a). 

Manso, 273(2-5); 275(2-5). 

Marcano-Berti, L., 457; 677(2-33). 

Marcano-Berti, L. & P. Alcedo, 119-979(2-37). 

Marin, E., 1608; 1625; 1654(2-33). 

Marinho, L. R., 350(2-43). 

Martius, 146; 264(2-17); 483(2-37); 1851(2-4); s.n.(2- 

3(2-3a); s.n.(2-10); s.n.(2-20). 

Martin, s.n.(2-22); s.n.(2-33); s.n.(2-40). 

Martinelli, G., 6791(2-38). 

Martinelli, G. & T. Soderstrom, 9751(1-4b). 

Martinelli, G., et al. 6761(2-37). 

Marulanda, 0. & S. Marquez, 1799(2-24). 

Matuda, E., 3193(2-7). 

Maxon, W. R. & A. Valentine, 7044(2-42). 

McDaniel, S. & M. Rimachi Y., 20424(2-37). 

McPherson, G., 10261; 10761(2-42). 

Medri, 33(2-20). 

Meier, W. et al., 2306(2-37). 

Melinon, M., 357(2-52); s.n.(1-2); s.n.(2-39); s.n.(2-40). 

Mello Barreto, 1791(2-24). 

Mello, F., s.n., herb. 1791(2-24); s.n., herb. 1883(2- 

10); s.n., herb. 1918(2-31); s.n., herb. 2275(2-5). 

Mello, F. & G. Mota, 47(2-37). 

Mello, F. & L. Coelho, s.n., herb. 3597; s.n., herb. 

3599; 3634(2-37); s.n., herb. 3778(2-31). 

Mendez, P. et al., 248(2-42). 

Mesquita, A. L. et al., 825(2-31). 

Meyer, 185(1-4a). 

de Michel, R. & St. G. Beck, 1237(2-5). 

Miller, G., 1396; 1401(1-9). 

Miller, J. & J. C. Sandino, 1117(2-42); 1369; 1401(1-9). 

Miller, O.O. & J.R. Jonhston, 128(1-9). 

Milliken, 2031(2-2). 

Miranda, F., 7341(1-4a). 

Molina R., A., 3936; 6774(1-4a); 13703(1-9). 

Molina R., A. & Molina, 25601(1-9). 

Molino, J.F., 1469(2-32). 

Monsalve B., M., 784(1-5). 

Monteiro, 0. P., 1285(2-10). 

Monteiro, 0. P. & C. Damiao, 461; 525(2-3(2-3a); 

606(2-36). 

Monteiro, 0. P. & J. Ramos, 985(2-37). 

Mora, L. E., 1450(1-4a). 

Moraes, M. & J. Sarmiento, 1111; 1164(2-24). 

Moraes, M. et al., 1360(2-34a); 1378; 1475; 1570(2-24). 

Moreno, P., 26743(2-42). 

Moretti, C., 353(2-52); 364(2-3b); 658(2-36); 715(2- 

37); 971; 1189(2-14). 

Mori, S. A., 7738(2-42). 

Mori, S. A. & B. Boom, 14711; 14961(2-3b); 15138(2- 

39); 15164(2-32); 15234(2-52); 15246(2-32). 

Mori, S. A. & M. Crosby, 6318(2-42). 

Mori, S. A. & C. Gracie, 18673(2-39); 18928(2-52); 

18951(2-3b); 21079(2-37); 23853(2-33). 

Mori, S. A. & J. de Granville, 8914(2-32). 

Mori, S. A. & J. Kallunki, 3652(2-42); 5199(2-29); 

5382(2-41). 

Mori, S. A. & T. Pennington, 18140(2-40). 

Mori, S. A. & A. Pepper, 24260(2-40). 

Mori, S. A. & J. Pipoly, 15464(2-52); 15533(2-37). 

Mori, S. A. & R. Souza, 17301(2-40); 17605(2-36). 

Mori, S. A. & Y. Veyret, 8929(2-22). 

Mori, S. A. et al., 6537(2-42); 9303; 11694(2-24); 

13975(2-4); 14926(2-32); 14988; 15027(2-33); 

15653(2-52); 19144(1-2); 20822(2-14); 20868(2- 

52); 21365(2-3b); 21369(2-26); 21605(2-33); 

21607(2-40); 21608(2-39); 22006(2-33); 22025(2- 

52); 23121(2-39); 23588(2-34a); 23954; 24182(2- 

33); 24669; 24671(2-52). 

Morillo, G. et al., 3339(2-30). 

Morong, T., 817(1-8). 

Morrow, C. F., 121(1-9) 

Morton, C. V., 7090(1-4a). 

Mostacedo, B. et al., 2607; 2680 (2-5). 

Mutchnick, P. & Tiwari, 1440(2-33). 

Mutis, J. C., 1154(2-34a); 4195(2-51). 

Nascimento, C. G., 418(2-36). 

Nascimento, J. R., 392(2-37). 

Nascimento, 0. C., 825(2-6). 

Nee, M., 33356(2-24); 34702(2-37); 35630; 38948(1-8). 

Nee, M. & G. Coimbra S., 36952(2-5).


Nelson, B. 2428(2-52). 

Nicolson, D. H., 4219(1-9) 

Nunes, E. & P. Martins, 8996(2-5). 

Nuniez, P. et al., 10903(2-25); 19631(2-24); 23976 

(2-34c). 

Oldeman, 1052; 1265; 1278(2-22); 1531(2-34a); 

2223(2-22); 2390(2-36); 2597(2-22); 2666(2-40); 

2671; 2811(2-52); 2922(2-22); 2994(2-36); 

3241(2-33); 3292(2-52); 3304(2-3b); 4159(2-52); 

4183(2-40); B-595(2-22); B-1194(1-22); B-1871 

(2-22); B-1883(2-36); B-1911(2-52); B-2322; B-2582; 

B-3321(2-22); B-3481(2-40); B-3521; B-3577 

(2-36); B-3578; B-4051(2-22); B-4094(2-40); B- 

4180(2-33); B-4008(2-39); T-32(2-52); T-285(2-14); 

T-5 14(2-22). 

Oldeman & Sastre, 104; 11 1(2-40); 185(2-36); 219 

(2-22). 

Oldenburger F. H. F. et al., 538(2-49). 

Oliveira, A. et al., 242(2-19). 

Oliveira, E., 4218(2-19); 4237(2-40); 6611; 6727(2-49). 

d'Orbigny, 818(1-8); 894; 896(2-5). 

Pab6n, M., 985(2-10). 

Padilla, S. A., 438(1-4a). 

Palacios, W., 2182(2-48). 

Palacios, W. & Neil, 1132(2-24). 

Palacios, W. & M. Tirado, 11097(2-21). 

Palacios, W. et al., 8059(1-4a). 

Paredes, R., 953(2-42). 

Pav6n, H., s.n.(2-47). 

Pedersen, T., 1833(1-8). 

Pennington, T. D. & J. Ruiz, 12450(2-52); 12456(2-37). 

Pennington, T. D. & Sarukhan, 9386(1-4a). 

Pennington, T. D. et al., 9553(1-4a). 

Perez, B., 1288; 1436(2-5). 

Philipson, W. R. & J. Idrobo, 1802(2-24). 

Philipson, W. R. et al., 2142(2-24). 

Pickel, D. B., 146; 306(2-5). 

Pinto E., P. & C. Sastre, 919(2-24). 

Pipoly, J. J. & A. Cress, 6797(2-37). 

Pipoly J. J. et al., 11348(2-32); 12630(2-3a); 14227 

(2-34a); 15028(2-52). 

Pirani, J., 1354(2-17). 

Pires et al., 696(2-36). 

Pires, 0. & A. B. C. Piata, 266(2-10). 

Pires, 0. & J. Lima, 156(2-26). 

Murqa Pires, J., 3968(2-27); 47465(2-14); 58132(2-5). 

Murqa Pires, J. & P.P. Furtado, 17165(1-8). 

Murqa Pires J. & N. T. Silva, 4434; 4441; 11346(2-19). 

Murqa Pires, J. & P. B. Cavalcante, 52291; 52298(2-22). 

Murca Pires, J. et al., 16850; 50406(2-39); 50549(1-2); 

50774; 51412(2-36). 

Pittier, H., 4249(2-42); 5806; 6534(2-34a); 7814; 

8776(1-9); 8775(1-4a); 8814(2-34a); 10276(1-4a); 

11451(1-9); 14253; 14298(2-34a); 15615(2-39); 

15727(1-4a). 

Plowman, T., 6016(2-34a). 

Plowman, T. et al., 12664(2-31). 

Poiteau, M., s.n.(2-22); s.n.(2-37); s.n.(2-39); s.n.(2-45). 

Poveda, L. J., 446(2-24). 

Powell, 72(1-4a). 

Prance, G. T., 14330(2-43). 

Prance, G. T. & I. Cordeiro, 29706(2-37). 

Prance, G. T. & T. Pennington, 1787(2-36). 

Prance, G. T. & J. F. Ramos, 6909(2-45). 

Prance, G. T. & N. T. Silva, 58919(2-36); 58947(2-49); 

59317(2-5). 

Prance, G. T. et al., 1355; 1482(2-38); 1544(2-40); 

2202(2-26); 2216(2-37); 2425(2-34a); 2624(2-26); 

3124(2-31); 3785(2-26); 4135(2-39); 4712(2-27); 

4716; 4906(2-37); 5092(2-38); 8117(2-24); 9009 

(2-37); 10674(2-39); 10802; 10810(2-37); 10920; 

11044(2-39); 11245(2-24); 11458; 14085 (2-37); 

14816(2-31); 18769(2-26); 18815 (2-17); 24113 

(2-24); 24727(2-38); 26521(2-37). 

Prevost, M. F., 184(2-22); 583(2-22); 1079(2-40); 

1109(2-40); 1496(2-22); 2170(2-16); 3028(2-52); 

3055(2-22). 

Prevost, M. F. & P. Grenand, 1934(2-22). 

Prevost, M. F. & D. Sabatier, 2638(1-2); 2976(2-12); 

2978(2-14); 2979(2-8). 

Pruski, J. et al., 3311(2-38); 3390(2-39). 

Pulle, A., 110(2-52); 526(2-39). 

Quevedo, R. & T. Centuri6n, 301(2-24). 

Rabelo, B. V. & J. Cardoso, 2888(2-22). 

Rabelo, B. V. et al., 2415(2-36). 

Ramirez, J. G. & Cirdenas, D., 519; 876; 1455; 1873 

(2-24). 

Ramirez, R., 31(2-42). 

Ramos, J., 910; 1169(2-12); s.n.(2-13). 

Ramos, J. & C. D. A. Mota, 152(2-34a); 1675(2-19). 

Ramos, J. & E. F. Lima, 1567(2-19). 

Rankin, J. et al., 84(2-12). 

Ratter, J. A. et al., 5894(2-39); 6366; 6402(2-5). 

Renteria, E., 3741(2-18). 

Renteria, E. & D. Cardenas, 4339(2-29). 

Renteria, E. et al., 3600(1-4a); 4758; 4841(2-18); 

5410(1-6). 

Revilla, J., 2408(2-52); 3596(2-37). 

Revilla, J. et al., 6958(2-24). 

Reynel, C., 689(2-34a). 

Ribamar, J. & J. Ramos, 274; 363(2-31). 

Ribeiro, J.E.L.S. et al., 1126(2-45); 1299(2-26). 

Richard, L. C., s.n.(2-22). 

Riedel, L., 522(2-44); 1106(2-17); 1170(2-5); 1367 

(2-27). 

Riedel, L. & Lund, 2645(2-5). 

Riera B., 783(2-14); 1459(2-32). 

Rimachi Y., M., 2408(2-47); 2877; 3176(2-24); 7309 

(2-52); 7499(2-3a); 8965(2-27). 

Robert, A., 554b(2-17). 

Roberts, L., 16313(2-14). 

Roberts, L: & F. v. Troon, 14806(2-8). 

Robleto, W., 621(2-42). 

Rodal, M. J. N. & M. F. Sales, 460(2-5). 

Chagas Rodrigues, J., 1243(2-31). 

Rodrigues, R. S., 8812(2-49). 

Rodrigues, W. A., 546(2-31); 7257(2-26); 9227(2-20); 

8989(2-27); 10863(2-26). 

Rodrigues, W. A. & J. Chagas, 2557(2-10); 2634(2-26); 

4399(2-37); 4682(2-31). 

Rodrigues, W. A. & D. Coelho, 1738(2-31); 2270(2-26). 

Rodrigues, W. A. & L. Coelho, 3925(2-20); 5228(2-26). 

Rodrigues, W. A. & J. Lima, 2242(2-37); 2665(2-31); 

4159(2-37). 

Rodrigues, W. A. et al., 10562(2-37); 10557(2-24); 

11089(2-37); 21329(2-4). 

van Rohr, J. P. B., 98; 99(2-34a). 

Roldan, F. J. et al., 945; 1003(1-4a). 

Romero Castanieda, R., 670; 71.1(1-4a); 1792(2-42); 

4029(2-37); 4153(1-6); 5049(2-41); 10009(1-4a). 

Romoleroux, K., 3068(1-6). 

Rosa, N. A., 2418(2-38); 4416(2-40).


Rosa, N. A. & J. Martins, 4324(2-37). 

Rosa, N. A. et al., 4083(2-40). 

Rose, J. N. & Rose, 21625; 21678(1-9). 

Rose, J. N. et al., 3454(1-9). 

Ruiz, J. & W. Balseca, 816(2-3a). 

Ruiz, J. & W. Melendez, 1269(2-3a). 

Rusby, H. H., 1390(2-5); 1409; 2527(2-24). 

Russell, 12552(1-4a). 

Sabatier, D., 213(2-39); 859(2-32); 1114(2-14); 

3510(1-2); 3572(2-32). 

Sabatier, D. & M. F. Prevost, 1373(1-2); 1852(1-22); 

1932(2-14); 2202(2-8); 2792(2-3b); 2816(1-2); 

2872(2-36); 2885(1-22); 2949(2-34a); 3253(2-37); 

3254(2-14); 3269(2-34a); 3433; 3436(2-3a); 

3601(1-2); 3631; 3658(1-22); 3663(2-8); 3676(2- 

40); 3717; 3820(1-22); 3855(2-12); 3894(2-14); 

4010(2-34a); 4196(1-2); 4237(2-3b); 4291(2-36). 

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Sanoja, E. & C. Loup, 3523(2-52) 

Santino, 279(2-5). 

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Santos, J. U. & C. S. Rosario, 400(2-17). 

Saravia T., C., 2210(1-4a); 2359(1-9); 2447(1-4a); 

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Saravia T., C. & M. Saravia, 3547(1-4a); 3667(1-9). 

Sargent, F. H., 219(1-9). 

Sastre, C., 5669(2-52). 

Sastre, C. et al., 3832(2-52); 3929(2-39); 8222(2-16). 

Schafer, 9221(2-52). 

Schomburgk, Richard, 1351(2-16). 

Schomburgk, Robert, 736; 738(1-23); 859; s.n.(I-46). 

Schubert, B. G., 2234(2-36). 

Schultes, R. E. & I. Cabrera, 12612; 12663(2-6); 

12767(2-37); 13539(2-6); 18660(2-37). 

Schultes, R. E. & F. L6pez, 9246(2-20). 

Schulz, J. P., 8073(2-39); 8387(2-40). 

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6335(2-34a). 

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Seidel, R., 3148(2-5). 

Seidel, R. & D. Vaquiata, 7496(2-34a). 

Seidel, R. et al., 5618; 6040(2-34a). 

Sello, s.n.(2-5). 

Shattuck, O., 377(2-25). 

Shafer, J. A., 1007; 1324; 1341; 1383(1-9). 

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da Silva, A. S. & C. Rosario, 4659(2-49). 

da Silva, A. S. et al., 905(2-6). 

Silva, I. A., 255(1-1); 355(1-4b). 

Silva, J. A., 268(2-37). 

Silva, M., 1266; 2111(2-24). 

Silva, M. & R. Souza, 2500; 2524(2-36); 2535; 

2540(1-2). 

Silva, M. et al., 2153(2-31); 966(2-43). 

Silva, M. F., 626(2-37); 827(2-10). 

Silva, M. F. et al., 103(2-37); 519; 530(2-45); 1470 

(2-37). 

Silva, M. G., 5036; 5936(2-40). 

Silva, M. G. & C. Rosario, 5328(2-40). 

Silva, M. G. & R. Bahia, 3478(2-38). 

Silva, N. T., 117(2-24); 1294; 1437(2-28); 1468(2-19); 

1542; 1665(2-40); 3908(2-42). 

Silva, N. T. & M. R. Santos, 4616(2-38). 

Silveira, M. et al., 675(2-25); 677(2-24); 876(2-49). 

Sintenis, P., 1597; 4788(1-9). 

Smith, A. C., 2677(2-45); 3504(1-4b). 

Smith, F. D. Jr., 63(1-4a). 

Smith, H. H., 784; 785; 1695(1-4a); 1705(1-9). 

Snethiage, E., 10416(2-38). 

Soares, E., 105(2-37); 132(2-38). 

Solomon, J. C., 14623(2-24). 

Solomon, J. C. et al., 6856(2-1). 

Sothers, C. A. et al., 539(2-6); 998(2-43). 

Sousa, A. B. et al., 4230(2-5). 

de Souza, M. A. D. et al., 380(2-37). 

Sperling, C. R. et al., 5976(2-49). 

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s.n.(2-24); s.n.(2-36). 

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29248(1-9); 31421(2-42). 

Standley, P. C & E. Padilla V., 2467(1-4a). 

Starry, D. E., 260(2-42). 

Steege, H. & P. de Jager, 562(2-32). 

Stein, B. A. et al., 3994(2-43). 

Steinbach, J., 2905; 3511; 6562(2-34a); 6595; 7604 

(2-5). 

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Stergios, B. & J. Velazco, 13978(2-4). 

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Stevenson, J. A., 3014(1-9). 

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Steyermark, J. A. & B. J. Manara, 110418; 110778(2-48); 

1 10938(1-4b). 

Steyermark, J. A. & J. J. Wurdack, 116(2-42). 

Steyermark, J. A. et al., 106882(1-4a); 110938(1-4b); 

117702(2-2); 119784(2-48); 122991(1-4a). 

Stofers, A. L. et al., 87(2-16). 

Swabey, C., 12394(1-4a). 

Tamayo, F., 1500(2-48); 33961(1-4b). 

Tavares, S., 1121(2-49). 

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Teixeira, G., 2590(2-5). 

Teixeira, L. 0. A., 64(2-24). 

Teixeira, L. 0. A. et al., 89(2-52). 

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Tirado, M. et al., 552; 683(2-37). 

Toledo, J. F. & A. Gehrt, 43167(2-1). 

Traill, J. W. H., 125(2-20). 

Triana, J., s.n.(2-51). 

Trujillo, B., 5071(2-48); 6840(1-4a). 

Tuin Ortiz, R., 90; 761(1-4a). 

Tutin, T. G., 405(2-37). 

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377(2-39). 

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Vasquez, R., 12293(2-47). 

Vasquez, R. & G. Criollo, 5780(2-52); 5811(2-27); 

7066(2-52). 

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(2-24); 3770(2-34c); 5070(2-3a); 5875(2-52); 6358 

(2-24); 6815(2-37); 7876(2-27); 7913(2-6); 8923 

(2-52); 9126; 10107(2-24); 11807(2-37); 13131 

(2-24); 14401(2-43); 15023(1-4a). 

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7082(2-52); 7430(2-27); 8068(2-3a); 8082(2-52); 

11982(2-27); 12008; 13475(2-37); 13741(2-52); 

13745(2-3a); 14245(2-10); 14297(2-37); 14324 

(2-10); 18734(2-24). 

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Wied-Neuwied, M.A.P., s.n.(1-4b). 

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24); 9977(1-4a); 11833(2-49); 12343(1-4a); 

15174(2-42). 

Wilson, P., 251(1-9). 

Wilson-Browne J., 437; 471; 508(2-49). 

Wood herbarium Surinam, 321(2-12). 

Woodworth, R. &. P. Vestal, 631(2-42). 

Woytkowski, F., 7172; 7173(2-37). 

Wurdack, J. J. & L. Adderley, 43363; 43476(2-6). 

Wurdack, J. J. & Monachino, 39719(2-34a). 

Young, H. & D. Stratton, 135(2-34c). 

Zaandam, C., 6705(2-33). 

Zamora, N. et al., 1488(2-42). 

Zapullo, s.n.(2-21). 

Zarucchi, J. & B. Echeverry, 4673(1-9). 

Zegarra, 11(2-34a). 

Zent, S., 586-47b(2-42); 885-20; 885-24; 1085-03 

(2-23). 

Zetek, J., 3570(2-25); 4335(2-42). 

Zhang, S. Y., 18(2-26). 

Zufiiga, R., 70(2-42). 

Be, 138 

Bois-flambeau, 96, 132, 165 

Bosknipa, 96, 138, 165 

Breu, 101 

Cacao macaque, 123 

Cafecillo, 132 

Camboata', 101 

Camboata da folha grande, 101 

Canape, 47 

Cancudo, 83 

Candlewood, 81 

Carachupa caspi, 101 

Cascudo, 88 

Catapu', 36 

Cedrillo, 65 

Combat, 106 

Copal rosado de hoja grande, 132 

Copito, 155 

Cota prisa, 36 

Cotopalo, 37, 123 

Cotoperi, 49, 36 

Cotoperis, 118 

Cotoperis montaniero, 118 

Cotoperiz, 49, 123 

Cotopris, 36 

Cotopriz, 36 

Cotuperia, 118 

Cutupli, 123 

Cutuplis, 37 

Dayacoussa, 96 

Feijao cru, 62 

Flambeau dur, 141 

Gaan tatoe, 112 

Gaulette indienne, 96, 130, 165 

Genipe, 47 

INDEX OF LOCAL NAMES 

Grigri, 81 

Guaya, 36 

Guaya pais, 36 

Huaya, 36 

Huayo, 36 

HuayuTn, 36 

Huesillo, 132 

Huevos de chivo, 161 

Juapina, 144 

Japunaki, 123 

Jatoama, 118 

Jicacara, 130 

Juria, 36 

Kaeriniuae ukwaewaesa, 97 

Karimora, 81 

Kenep, 47 

Keneppy tree, 47 

Kenip, 47 

Kinap, 36 

Kraskrastiki, 138 

Limoncillo, 47 

Limpia botella, 155 

Macucu, 157 

Mamon, 47 

Mam6n cotopri, 36 

Mam6n cutupui, 36 

Mam6n cutuplis, 36, 37, 123 

Mam6n de cerro, 79 

Mam6n de Maria, 36 

Mam6n de mico, 36 

Mam6n de monte, 104 

Mamoncillo, 47, 127 

Mamoncillo de monte, 36 

Matapuerco, 163 

Mauraballi, 123 

Mel6n, 47 

Miiverezu, 72 

Moroballi, 81 

Morokwe, 1.21 

Muro-balli, 81 

Motoyoe, 44 

Olhio de boi, 62 

Olhio de venado, 92 

Palo azul, 127 

Paicoussa, 96, 130, 139, 165 

Papamundo, 44 

Pau de espeto vermelho, 88 

Payacoussa, 96, 141 

Petit gaulette, 96 

Pishico huayo, 138 

Pitomba, 37, 62, 72, 92, 101, 106, 

108, 116, 132, 138, 141, 148, 

151 

Pitomba amarela, 37, 92 

Pitomba brava, 101, 116, 130 

Pitomba da anta, 157 

Pitomba da mata, 62, 116, 136 

Pitomba de macaco, 62 

Pitomba do mato, 101, 106, 132 

Pitombarana, 79, 88, 104, 132 

Pitombeira, 37, 62, 92, 99 

Pitombinha, 101, 157 

Piton, 101, 121 

Poloc, 66 

Quenepa, 47 

Sand mora, 56, 81, 123 

Sangre de lapa, 132 

Sapatero, 104 

Shantona, 123

INDEX OF SCIENTIFIC NAMES 177 

Shantonilla, 123 

Singabassou, 79, 81 

Spanish lime, 47 

Tacho, 37 

Tambor caspi, 101 

Tapaljocote, 36 

Tapuyma, 79 

Tatoe, 1 12 

Acladodea, 4, 50 

pinnata, 4, 153 

Athyana, 3 

Averrhoidium, 3 

Alectryon, 20 

Burseraceae, 50 

Casimira, 28 

Castaspora, 4, 19, 20 

Comatoglossum, 5, 50 

strictum, 5, 161 

Crossonephelis, 4 

Cupania, 3 

congestiflora, 34 

longifolia, 127 

macrophylla, 130 

praealta, 76 

seemannii, 166 

subalbens, 81 

Diatenopteryx, 3 

Diploglottis 

australis, 50 

Doratoxylon 

mauritianum, 50 

Elattostachys 

verrucosa, 50 

Eriandrostachys, 4 

Euplassa, 3 

Exothea 

diphylla, 166 

paniculata, 50 

Fabaceae, 3 

Glenniea, 4 

Guarea, 3 

japurensis, 70 

Hedyachras, 4 

Hippobromus 

pauciflorus, 50 

Lecaniodiscus, 4 

Lepisantheae, 4 

Lepisanthes, 20 

amoena, 4 

Macphersonia, 4 

Matayba, 3 

glaberrima, 166 

guianensis, 5, 115, 166 

Meleagrinex, 50 

Meliaceae, 3 

Melicocca, 28 

apetala, 50 

australis, 50 

bijuga, 4, 45 

carpopodea, 45 

dentata, 50 

diphylla, 50 

Tatu-tatu, 141 

Tepu, 165 

Tepuime, 165 

Tit-gaulette, 96 

Toliatan, 96, 165 

Tuliata, 96, 139 

Virote huayo, 101 

Wai, 56 

INDEX OF SCIENTIFIC NAMES 

diversifolia, 50 

geniculata, 50 

javanica, 50 

lepidopetala, 44 

olivaeformis, 34 

paniculata, 50 

pubescens, 50 

trijuga, 50 

triptera, 50 

Melicocceae, 4, 19, 20 

Melicoccus, 3, 4, 28 

amoenus, 4, 50 

antioquensis, 29, 39, 41 

aymardii, 30, 43 

bijuga f. alata, 4, 45 

bijugatus, 4, 44 

espiritosantensis, 29, 44 

jimenezii, 48, 166 

lepidopetalus, 4, 44 

novogranatensis, 41 

oliviformis, 4, 34, 41 

oliviformis subsp. oliviformis, 34 

oliviformis subsp. intermedius, 

37 

pedicellaris, 30 

petiolulatus, 32 

Ophiocaryon, 3 

Plagioscyphus, 4 

Proteaceae, 3 

Protium, 50 

aracouchili, 50 

Racaria, 50 

sylvatica, 163 

Sabiaceae, 3 

Sapindus, 20, 23 

cerasinus, 97 

esculentus, 61 

oblongus, 99 

Simaba, 3 

Simaroubaceae, 3, 166 

Schleichera 

oleosa, 50 

Strophiodiscus, 4 

Talisia, 3, 4, 50 

acladodea, 4, 152 

acutifolia, 90, 94 

allenii, 132 

amaruyana, 144 

amazonica, 106 

angustifolia, 52 

bullata, 92 

carinata, 94 

carinata f. acutisepala, 163 

carinata f. sericea, 94 

Wanhede, 112 

Wild genip, 37 

Wuayu'n, 36 

Yarre patado, 132 

Yellow chenip, 36 

Yvapovo, 44 

Zwarte pintolobus, 76 

Zwarte pintolokus, 32, 118 

cararensis, 121 

caudata, 97 

cerasina, 97, 115 

chartacea, 53 

clathrata, 55, 86 

clathrata subsp. clathrata, 56 

clathrata sp. canescens, 56 

coriacea, 59, 79 

croatii, 102, 143 

cupularis, 104, 128 

dasyclada, 106 

diphylla, 166 

douradensis, 109 

dwyeri, 144 

equatoriensis, 109 

elephantipes, 130 

erecta, 155 

esculenta, 51, 61 

eximia, 112 

firma, 55, 63 

floresii, 66, 70 

furfuracea, 68 

ghilleana, 112 

glabra, 115 

glandulifera, 118 

guianensis, 4, 50, 88, 109, 116, 

139 

grandifolia, 143 

granulosa, 68 

hemidasya, 118 

heterodoxa, 157 

hexaphylla, 120 

hexaphylla subsp. elegans, 123 

hexaphylla subsp. hexaphylla, 

121 

hexaphylla subsp. multinervis, 

125 

humilis, 52 

intermedia, 37 

japurensis, 70 

jimenezii, 4, 48, 166 

laevigata, 126 

lanata, 73 

laxiflora, 5, 166 

longifolia, 127 

macrophylla, 130 

marleneana, 130, 134 

medri, 90 

megaphylla, 136 

micrantha, 163 

microphylla, 73 

mollis, 139 

mollis var. marleneana, 134 

morii, 141


morilloi, 112 

multinervis, 63 

nervosa, 143, 165 

obovata, 146 

oedipoda, 148 

oliviformis, 34, 166 

pachycarpa, 148 

panamensis, 121 

parviflora, 76 

pedicellaris, 166 

pentantha, 132 

peruviana, 166 

pilosula, 152 

pinnata, 50, 88, 152, 161 

porteriana, 166 

praealta, 76, 79 

prancei, 114, 166 

princeps, 104, 155 

pulverulenta, 31 

pygmaea, 52, 53 

reticulata, 163 

retusa, 101, 157 

rosea, 116 

sancarlosiana, 132 

setigera, 159 

simaboides, 79 

squarrosa, 79, 81 

stricta, 50, 161 

subalbens, 53, 81 

svensonii, 166 

sylvatica, 50, 88, 96, 163 

tirirensis, 144 

velutina, 84 

veraluciana, 53, 86 

sect. Cotopais, 28 

sect. Eutalisia, 88 

subsect. Acladodea, 88 

subsect. Pitombaria, 51 

sect. Racaria, 88 

subgenus Pitombaria, 14, 23, 50, 

51 

subgenus Talisia, 14, 23, 50, 88 

Tapirocarpus, 50 

talisia, 50, 139 

Theophrasta 

pinnata, 155 

Toulicia 

brachyphylla, 34 

Tristira, 4, 19, 20 

triptera, 50 

Tristiropsis, 4, 19, 20 

Ungnadia, 3 

Vouarana, 3 

Zollingeria, 20

Melicocceae (Sapindaceae): Melicoccus and Talisia - CNCFlora

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