Vines and Climbing Plants of Puerto Rico and the Virgin Islands

SMITHSONIAN INSTITUTION 

Contributions from the United States National Herbarium 

Volume 51: 1-483 

Vines and Climbing Plants 

of 

Puerto Rico and the Virgin Islands 

by 

Pedro Acevedo-Rodríguez 

Department of Botany 

National Museum of Natural History 

Washington, DC 

2005

ABSTRACT 

Acevedo-Rodríguez, Pedro. Vines and Climbing plants of Puerto Rico and the Virgin Islands. 

Contributions from the United States National Herbarium, Volume 51: 483 pages (including 

184 figures). The present book constitutes an illustrated field guide to the native, naturalized, or commonly 

cultivated vines and lianas of Puerto Rico and the Virgin Islands. It includes nomenclatural and taxonomic revisions, 

discussions on the distribution and conservation status, as well as full descriptions and illustrations for the 

species. A total of 386 species distributed in 64 families and 187 genera are treated. Of these 274 are native, 34 

are endemic, 64 have been naturalized and approximately 49 are commonly cultivated in gardens. A new 

combination of Aristolochia oblongata Jacq. ssp calciformis (Urb.) R.Rankin & Acev.-Rodr. is made. 

KEY WORDS: vines, lianas, climbing plants, scandent plants, Puerto Rico, Virgin Islands, flora, Caribbean, 

Greater Puerto Rico, Greater Antilles, West Indies. 

RESUMEN 

Acevedo-Rodríguez, Pedro. Vines and Climbing plants of Puerto Rico and the Virgin Islands. 

Contributions from the United States National Herbarium, Volume 51: 483 pages (including 

184 figures). La presente obra constituye una guía para los bejucos y plantas trepadoras, nativas, naturalizadas 

o comúnmente cultivadas que se encuentran en Puerto Rico y las Islas Vírgenes. En ésta se han actualizado la 

nomenclatura y taxonomía y se incluyen datos de la distribución y estado de conservación de las especies. Cada 

especie es descrita e ilustrada. En total se tratan 386 especies, distribuidas en 64 familias y 187 géneros. De 

éstas, 274 son nativas, 34 son endémicas, 64 son naturalizadas y alrededor de 49 son comúnmente cultivadas en 

nuestros jardines. La nueva combinación Aristolochia oblongata Jacq. ssp calciformis (Urb.) R.Rankin & Acev.- 

Rodr. es efectuada. 

PALABRAS CLAVES: bejucos, lianas, plantas trepadoras, plantas sarmentosas, Puerto Rico, Islas Vírgenes, 

flora, Caribe, Macro Puerto Rico, Antillas Mayores, Indias Occidentales. 

DATE OF PUBLICATION: February 2005 

Cover Design by Alice Tangerini based on an illustration of Margravia rectiflora and M. sintenisii by Bobby 

Angell. 

Illustrations prepared and copyrighted © by Bobbi Angell, 2003. 

Translated from Spanish into English by Duane Kolterman, 2004. 

Contributions from the United States National Herbarium (ISSN 0097-1618) Department of Botany, MRC 

166, National Museum of Natural History, Smithsonian Institution, Washington, DC, 20013-7012, USA. 

POSTMASTER: Send address changes to Contributions U.S. National Herbarium, Department of Botany 

MRC 166, National Museum of Natural History, Smithsonian Institution, Washington, DC, 20013-7012, 

USA. 

The paper used in this publication meets the minimum requirements of the American National Standard for 

Permanence of Paper for Printed Library Materials Z39.48—1984. 

Contributions from the U.S. National Herbarium was first published in 1890 by The United States Department 

of Agriculture. From July 1, 1902 forward it was published as a Bulletin of the United States National Museum. 

The series was discontinued after volume 38, 1974, and has been revived with volume 39, as a venue for 

publishing longer taxonomic papers, checklists, floras, and monographs, produced by staff and associates at 

the U.S. National Herbarium. It is externally peer reviewed, and published at irregular intervals. Subscriptions 

and other correspondence should be addressed to CUSNH, Department of Botany, MRC 166, National Museum 

of Natural History, Smithsonian Institution, Washington, DC, 20013-7012, USA. e-mail: CUSNH@si.edu. 

The present issue is available for free while supplies last.

PREFACE 

To the future generation 

of Caribbean botanists 

More than 15 years have passed since the publication of Los Bejucos de Puerto Rico Vol. 1., the 

precursor to the present edition. The original intention had been to prepare a comprehensive guide to 

the native and naturalized climbing plants in Puerto Rico. This task was interrupted, however, following 

my departure to pursue graduate studies in the area of systematic botany. Persuaded by Dr. Ariel E. 

Lugo, I decided to publish what at that time constituted the manuscript, and was therefore called the 

first volume in order to indicate its partial or incomplete character. The idea of completing the task 

survived various projects that required my full dedication. Having finished them, I have found the time 

necessary to complete this long-awaited work. Nevertheless, this has not required the preparation of a 

second volume, but rather a total revision, in keeping with current taxonomy, that includes all the 

species of climbing plants of Puerto Rico. At the same time, it has been expanded to include the species 

found in the Virgin Islands, a natural appendage to the Puerto Rican biota. As a result, the title of the 

present work reflects the botanical and geographical diversity represented here. It also contains new 

illustrations more in keeping with its professional character. Although the title “Climbing plants of 

Puerto Rico and the Virgin Islands” might seem more appropriate for this book, its present title maintains 

a clearer connection with its precursor. 



December, 2002 

The present work is a translation with corrections of the book Bejucos y plantas trepadoras de 

Puerto Rico e Islas Vírgenes. Most corrections deal with formatting, some with spelling, and very few 

with content. The major change involving content is the removal of Sabicea sp. a (Rubiaceae) as its 

record in Puerto Rico was based on sterile collections of Tetrapterys inaequalis. The removal of this 

species reduces the total number of vine species in Puerto Rico and the Virgin Islands from 386 to 385. 

Bibliographical citations for the accepted names of taxa have been added in order to facilitate literature 

searches, and a few references have been added. 



September, 2004

CONTENTS 

INTRODUCTION .................................................................................................................................... 7 

General Characteristics ...................................................................................................................... 7 

Stem Morphology ............................................................................................................................... 7 

Climbing and Attachment Mechanisms .......................................................................................... 11 

Diversity and Distribution ............................................................................................................... 13 

Methods ............................................................................................................................................. 15 

Acknowledgments ............................................................................................................................ 15 

GENERAL REFERENCES ................................................................................................................... 17 

SYSTEMATIC TREATMENT .............................................................................................................. 21 

Key to the Divisions ......................................................................................................................... 21 

Pteridophytes ........................................................................................................................... 21 

Dicotyledons ............................................................................................................................ 41 

Monocotyledons ..................................................................................................................... 403 

COLLECTIONS EXAMINED ............................................................................................................447 

ILLUSTRATIONS VOUCHERS ......................................................................................................... 456 

GLOSSARY .........................................................................................................................................465 

INDEX TO SCIENTIFIC NAMES .....................................................................................................470 

INDEX TO VERNACULAR NAMES ................................................................................................481

Vines and Climbing Plants of Puerto Rico and the Virgin Islands 7 

Vines and Climbing Plants of Puerto Rico and the Virgin Islands 

Pedro Acevedo-Rodriguez 1 

GENERAL CHARACTERISTICS 

Vines can be defined as climbing plants that 

are rooted in the soil and whose stems are 

incapable of maintaining themselves erect, so that 

they need external support. Although the term 

vine is used to describe any herbaceous or woody 

climber, it usually denotes an herbaceous climber 

with limited secondary growth. When, on the 

other hand, climbing plants are markedly woody, 

they are known as lianas. 

Shrubs are distinguished from vines by 

having rigid stems capable of maintaining 

themselves erect. Nevertheless, this distinction is 

not always easy to make, because there are 

intermediate forms between lianas and shrubs that 

have a tendency to climb or support themselves 

on nearby objects. These are known as clambering, 

scrambling or scandent plants. These climbing 

plants are usually characterized as vines in the 

broad sense of the word, and for this reason they 

are included in the present book. 

Climbing plants are found in numerous 

ecosystems, but are more abundant in lowelevation 

tropical forests than in any other habitat. 

According to Gentry (1991), climbing plants in 

temperate forests represent on average 7% of the 

local flora, while in tropical forests this number 

reaches 20%. Lianas are characteristic of tropical 

forests, where at least 50% of the trees contain 

lianas. These can constitute a significant portion 

of the biomass of the forest, since their crowns 

can be as large as that of the tree that supports 

them. In some moist forests or rain forests in 

continental tropical areas, lianas can represent up 

INTRODUCTION 

to 40% of the plant species present in the 

ecosystem (Jacobs, 1988), so that some of these 

forests are known locally as liana forests. 

Despite the ecological importance and the 

great number of species of climbing plants, they 

have received relatively little attention on the part 

of ecologists and forest engineers. The objective 

of this book is to encourage the understanding of 

this interesting group of plants. 

STEM MORPHOLOGY 

This section does not purport to be a treatment 

of the morphology of climbing plants, but rather 

to present general characteristics that are useful 

in their identification. Some of the most 

conspicuous morphological traits found in many 

species of climbers that are useful in 

distinguishing different taxonomic groups at the 

level of family, genus, or species are discussed 

below. In general, climbing plants present 

numerous morphological and anatomical 

characteristics that distinguish them from other 

forms of plant life. Among these characteristics 

are the anatomical structure of the stems and the 

climbing and attachment mechanisms. 

Vines have long and flexible stems that depend 

on external support to maintain themselves erect 

or to reach illuminated areas in their habitat. Their 

stems are characterized by the scarcity of 

supporting cells (fibers) and an increase in the 

diameter of the xylem vessels, which may be 

visible to the naked eye. The increase in the 

diameter of the xylem vessels triplicates the 

conduction of water, making lianas able to 

maintain a great quantity of leaves in relation to 

1 United States National Herbarium, National Museum of Natural History, MRC-166, Smithsonian Institution, Washington, DC 

20013-7012, U.S.A. e-mail: acevedop@si.edu

8 


Fig. 1. Stem cross sections. A. Pisonia aculeata. B. Ipomoea violacea. C. Pinzona coriacea. D. Chamissoa altissima. E. 

Securidaca virgata. F. Hyperbaena domingensis.


the total diameter of their stems. These stems that 

are specialized for the conduction of water are 

known only in plants that possess xylem vessels 

(elements with perforated walls), and are absent 

in those that have only tracheids or imperforate 

elements. Imperforate elements obviously 

represent an obstacle to the free flow of water, 

slowing it down and making water transport over 

great distances difficult. 

The stems of climbing plants face structural 

challenges that differ from those experienced by 

trees and shrubs. They are subjected to tensile and 

compacting forces, due to the movement of the 

structures (usually trees) that support them. For 

this reason their stem construction, with an 

alternation of vascular and parenchymatous 

tissues, gives them considerable flexibility to 

withstand these types of pressure. In addition, the 

stems of climbing plants are subject to friction 

against the host trees that can lacerate their bark 

and thus wound the phloem tissue. Many lianas 

have encountered a solution to this problem by 

having phloem tissue inside the xylem (Dobbins 

and Fisher, 1986). The arrangement of phloem 

tissue in relation to the xylem can produce patterns 

sometimes considered anomalous, which serve to 

characterize families or genera of lianas (Schenk, 

1893; Carlquist, 1991; Gentry, 1991; Acevedo- 

Rodríguez, 1993). Due to their taxonomic utility, 

these patterns form part of the species descriptions 

presented in this book. 

For the purposes of this chapter, a brief 

summary is presented below to characterize in 

general terms the most common patterns that are 

known for the stems of the lianas of Puerto Rico 

and the Virgin Islands. These are described based 

on cross sections of mature stems, without taking 

developmental processes into consideration. 

1. Discrete vascular bundles. This pattern is the 

result of the activity of successive bands of 

cambium, which produce discrete bundles of 

xylem and phloem surrounded by parenchyma 

cells. The resulting pattern is that of collateral 

bundles dispersed in connective tissue 

(parenchyma). Examples of this type of pattern 

are found in the genera Bougainvillea and Pisonia 

(Fig. 1A) of the family Nyctaginaceae. 

2. Alternation of bands of vascular tissue with 

connective tissue. 

Concentric bands. This pattern is the result of 

the activity of successive bands of cambium that 

produce a band of vascular tissue (xylem and 

phloem) accompanied by a band of connective 

tissue (parenchyma). The cambial activity is 

repeated to produce successive concentric bands 

of vascular tissue and connective tissue. The 

connective tissue can be as wide as the vascular 

tissue, thus producing a conspicuous pattern of 

alternating bands. This pattern can be observed 

in members of the genus Ipomoea (Fig. 1B) of 

the family Convolvulaceae and in the genera 

Pinzona (Fig. 1C) and Doliocarpus of the family 

Dilleniaceae. On the other hand, the band of 

connective tissue can be very narrow and hardly 

discernible, giving the appearance of xylem made 

up of successive concentric layers. Examples of 

this pattern can be observed in the genera 

Chamissoa (Fig. 1D) and Pfaffia of the family 

Amaranthaceae. 

Non-concentric bands. This pattern, like the 

previous one, is the result of the activity of 

successive bands of cambium. In this case, 

however, the activity of the cambial tissue gives 

rise to asymmetric bands, which develop primarily 

toward only one sector of the stem, thus producing 

a stem whose pith is not in a central position. 

Examples of this pattern are seen in the genus 

Securidaca (Fig. 1E) of the family Polygalaceae, 

in the genus Machaerium of the family Fabaceae, 

and in the genus Hyperbaena (Fig. 1F) of the 

family Menispermaceae. 

3. Polystelic stems. 

A. These can be produced by stems with 

supernumerary cambial tissue whose secondary 

growth forms a stem composed of numerous 

vascular cylinders. In this pattern the peripheral 

vascular cylinders are of a smaller diameter than 

the central cylinder and can be seen both in young 

stems and in mature ones. Examples of this 

pattern are found in the genera Paullinia (Fig. 

2A) and Serjania (Fig. 2B) of the family 

Sapindaceae. 

B. These are also produced by the successive 

production of cambial tissue within the cortical 

tissue, which develops into peripheral vascular

10 


Fig. 2. Stem cross sections. A. Paullinia pinnata. B. Serjania polyphylla. C. Chiococca alba. D. Turbina corymbosa. E. 

Rhynchosia phaseoloides. F. Marcgravia rectiflora.


cylinders of different diameters. This pattern is 

visible only in mature stems. Examples are found 

in the genus Chiococca (Fig. 2C) of the family 

Rubiaceae and in the genus Turbina (Fig. 2D) of 

the family Convolvulaceae. 

4. Compressed stems. This pattern, with the 

vascular cylinder in a central position, is obtained 

through asymmetrical secondary growth, in which 

the stem grows laterally in two opposing 

directions. Examples of this pattern are found in 

the genera Rhynchosia (Fig. 2E) and Dioclea of 

the family Fabaceae. 

5. Deeply lobed stems. This pattern is achieved 

by secondary growth of the stem in particular 

regions, giving rise to the lobes. Examples of this 

pattern are found in the genus Marcgravia (Fig. 

2F) of the family Marcgraviaceae, in the genus 

Distictis (Fig. 3A) of the family Bignoniaceae, 

and in the genus Passiflora (Fig. 3F) of the family 

Passifloraceae. 

6. Stems with the xylem forming a cross of 4- 

12 arms. This pattern is produced through 

differential secondary growth of the xylem and 

phloem, with the result that there is more xylem 

tissue in the area of the arms. This pattern is very 

common in climbers of the family Bignoniaceae, 

e.g., Arrabidaea, Cydista (Fig. 3B) and 

Amphilophium (Fig. 3C). 

7. Cylindrical stems with lobed xylem. These 

are formed by the pronounced growth of xylem 

tissue in certain regions of the stem. This pattern 

can be seen in the genus Passiflora of the family 

Passifloraceae. In the genus Macfadyena (Fig. 3D) 

of the family Bignoniaceae, this pattern is 

obtained through differential growth of the xylem 

and phloem, producing more xylem in the area 

of the lobes. 

8. Cylindrical stems with conspicuous rays. A 

cross section of the stem shows the presence of 

wide parenchymatous rays inserted in the xylem 

tissue. Examples of this pattern can be seen in 

the genera Pristimera (Fig. 3E) and Hippocratea 

of the family Celastraceae and in the genus 

Psiguria of the family Cucurbitaceae. 

CLIMBING AND ATTACHMENT 

MECHANISMS 

Climbing plants achieve their objective of 

climbing on and attaching themselves to host 

plants by means of different active or passive 

mechanisms. Some species have active 

mechanisms for both tasks, while others are 

passive in one or both of them. Twining plants, 

as well as those that have tendrils or sensitive 

stems, possess active mechanisms that permit 

them to achieve both objectives. Climbers with 

recurved spines or adventitious roots do not have 

active climbing mechanisms, but these structures 

represent an active mechanism for the task of 

attaching them to the host plant. Clambering 

plants represent an extreme case in which both 

mechanisms are passive. The different climbing 

and attachment mechanisms are listed and 

discussed below. 

1. Tendrils. Tendrils are sensitive, usually 

filamentous appendages with which some plants 

climb on and attach themselves to host plants. 

These are developed from various structures of 

the plant body, and are discussed below under the 

following categories. 

A. Axillary tendrils. These are homologous to 

short axillary branches; examples are found in 

the Cucurbitaceae (Fig. 4A) and Passifloraceae. 

In the genus Gouania (Rhamnaceae) they may 

develop at the end of a short axillary branch, 

which sometimes produces a single leaf (Fig. 4B). 

B. Tendrils opposite the leaves. This type of 

tendril is probably homologous to the distal end 

of the main stem of the plant; consequently, the 

branches form a sympodial system. Examples of 

this type are found in the Vitaceae (Fig. 4C). 

C. Tendrils in the inflorescence. In the climbing 

Sapindaceae, the tendrils are homologous to the 

basal lateral branches of the inflorescences. They 

are present in pairs in the basal flowering portion 

of the inflorescence. Examples of this type are 

found in the genera Paullinia and Serjania (Fig. 

4D). 

D. Foliar tendrils. In many climbing genera of 

Bignoniaceae, the tendrils are found to replace

12 


Fig. 3. Stem cross sections. A. Distictis lactiflora. B. Cydista aequinoctiale. C. Amphilophium paniculatum. D. Macfadyena 

unguis-cati. E. Pristimera caribaea. F. Passiflora multiflora.


the terminal leaflet of the leaves (Fig. 4E). They 

are simple, trifurcate, or sometimes are found to 

be modified into a harpidium or small claw or 

into small adventitious discs. 

E. Tendrils derived from the leaf sheath. In the 

Smilacaceae (Fig. 4F), the tendrils represent a 

prolongation of the leaf sheath. 

2. Twining plants (Fig. 4G). Twining plants, 

commonly called vines, have active mechanisms 

to climb on and attach themselves to the host 

plant. They present a circumnutational movement 

in which their stems, somewhat arching in the 

distal portion, rotate on their own axis, rather like 

the hands of a clock. This movement is essential 

so that the vine can locate a structure on which it 

can climb and thus use as a source of support. 

The principal shoot of a vine rotates freely until 

it runs into some structure that presents enough 

resistance. Once this has happened, the stem of 

the vine continues its rotational motion, but in a 

smaller spiral, thus twining around the host plant. 

The greater the diameter of the host structure, the 

more difficult is the ascent, because the vine’s stem 

is subject to the force of gravity. Trees of excessive 

diameter represent an obstacle for the vine’s 

ascent. 

3. Sensitive branches or leaves. This category is 

analogous to tendrils, but does not present any 

type of modification; in this case the branches or 

leaves are sensitive (prehensile or twining) 

without having any kind of structural 

modification. This mechanism can be considered 

to be active both in the task of climbing and in 

that of attachment to the host plant. Examples of 

species with sensitive stems are found in the 

family Celastraceae, and species with sensitive 

leaves are found in the genus Clematis (Fig. 4H) 

of the family Ranunculaceae. 

4. Adventitious roots. Adventitious roots do not 

really represent an active climbing mechanism 

except in young plants, but in adult plants they 

are very effective in the task of holding onto the 

host plant. Examples are found in the genus 

Marcgravia of the family Marcgraviaceae and the 

genus Schlegelia of the family Schlegeliaceae. 

5. Cauline or foliar spines. The presence of 

spines on the stems or leaves of some climbing 

species may be considered to be an active 

mechanism for holding onto the host plant but 

not necessarily for the task of climbing. Examples 

of these structures are found in the cacti, some 

species of the genera Mimosa and Acacia of the 

family Fabaceae, the genus Celtis of the family 

Ulmaceae, and the genus Berylsimpsonia of the 

family Asteraceae. 

6. Clambering plants. Clambering plants are 

climbing plants that manage to climb on and gain 

support from other plants by means of passive 

mechanisms, that is, leaning against and growing 

upon the host plant. Examples of this type of plant 

are found in the genus Lepidaploa of the 

Asteraceae and the genus Senna of the Fabaceae. 

DIVERSITY AND DISTRIBUTION 

The total number of species of climbing plants 

that are found in Puerto Rico and the Virgin 

Islands amounts to 386, distributed in 64 families 

and 187 genera. Of these, 274 are native, 34 are 

endemic, 64 are naturalized, and about 49 are 

commonly cultivated in our gardens. Among the 

endemic vines is Neorudolphia, the only plant 

genus endemic to Puerto Rico. 

The naturalized species are usually tropical 

weeds with a very wide distributional range, not 

only in Puerto Rico and the Virgin Islands but 

throughout the tropics. These species are generally 

of rapid growth and are typically found in areas 

of disturbed vegetation, such as abandoned farms, 

roadsides, pastures, or secondary forests and 

thickets. Examples of these species are: Asystasia 

gangetica, Thunbergia alata, Thunbergia 

fragrans, Cryptostegia grandiflora, Momordica 

charantia, Lablab purpureus, Pueraria 

phaseoloides, Vigna hosei, and Jasminum 

fluminense. 

The native species, as a general rule, 

have a less widespread distributional range than 

the naturalized species. The range varies from 

species with a widespread neotropical distribution 

to species that are restricted to a group of islands. 

Similarly, within Puerto Rico and the Virgin 

Islands these species can have quite a widespread 

range or may be restricted to particular 

ecosystems. Species with a widespread neotropical

14 

Vines and Climbing Plants of Puerto Rico and the Virgin Islands


distribution are usually found in areas of disturbed 

or secondary vegetation, and thus their 

distribution in Puerto Rico is quite widespread. 

Examples of species that show this distributional 

range are: Mikania cordifolia, Mikania congesta, 

Macfadyena unguis-cati, Cayaponia americana, 

Melothria pendula, Ipomoea setifera, Ipomoea 

tiliacea, Merremia dissecta, Merremia 

quinquefolia, Merremia umbellata, Acacia retusa, 

Centrosema pubescens, Vigna luteola, 

Cissampelos pareira, Passiflora foetida, 

Passiflora rubra, Passiflora suberosa, Paullinia 

pinnata, and Cissus verticillata. 

The endemic species have, by definition, the 

least widespread distributional range, but their 

distribution varies considerably from species to 

species. Some are found throughout the island(s), 

while others are restricted to one or several 

localities. Examples of the former are: Forsteronia 

portoricensis, Chromolaena borinquensis, 

Lepidaploa borinquensis, Mikania fragilis, 

Mikania odoratissima, Mikania pachyphylla, 

Piptocarpha tetrantha, Clusia gundlachii, 

Neorudolphia volubilis, Heteropterys wydleriana, 

Marcgravia sintenisii, and Stigmaphyllon 

floribundum (shared with the Virgin Islands). 

Species restricted to several localities are the 

following: Metastelma monense, Matelea 

sintenisii, Bidens urbanii, Mikania porosa, 

Mikania stevensiana, Passiflora tulae, and Rubus 

florulentus. Some of the endemic vines have such 

a limited distributional range that they may be 

characterized as threatened species; these are the 

following: Marsdenia elliptica, Marsdenia 

woodburyana, Piptocoma acevedoi, Cordia 

bellonis, Gonocalyx concolor, and Galactia 

eggersii (endemic to the Virgin Islands). 

METHODS 

The present work is based on the study of 

herbarium specimens, supplemented with fresh 

material. This applies both to the descriptions and 

to the data on phenology and distribution. The 

descriptions and data presented come from plants 

collected in Puerto Rico and the Virgin Islands. 

On rare occasions, the descriptions or 

phenological data have been supplemented with 

material from other Caribbean islands or 

monographic studies of the species in question. 

The descriptions that are presented are quite 

representative, although it is possible that the 

study of fresh material, flowers in particular, 

might show larger dimensions than those 

mentioned here. The habit dimensions are mostly 

based on the study of fresh material and are 

estimations, since it is impractical to give absolute 

measurements of stem length. In cases where the 

species was not seen in the field, information on 

the habit comes exclusively from herbarium labels 

or literature reports. The distributions have been 

supplemented with various floristic publications 

on forests or keys and islands belonging to Puerto 

Rico (see general references). In particular, the 

publications on Cayo Santiago, Desecheo, and 

Vieques have been of great help, since I have not 

sampled any of these islands. 

ACKNOWLEDGMENTS 

This book would not be possible without the 

collaboration and encouragement of many friends 

and colleagues, whose contribution to this work I 

acknowledge. This has been a lengthy process that 

has taken several decades to complete. Roy O. 

Woodbury gave me the original inspiration and 

guided me through the processes of botanical 

learning. Ariel E. Lugo has been instrumental 

since the beginning of this work. His visionary 

interest and support for the publication of both 

the first volume and the present one have been 

essential for this book to see the light of day. 

Numerous friends, colleagues, or family members 

have collaborated in one way or another in 

explorations of Puerto Rico and the Virgin Islands. 

Among these are my wife Amneris Siaca, my 

daughter Liana, and my nephew Javier, as well 

as my friends and colleagues Migdalia Álvarez, 

Franklin S. Axelrod, Gary Breckon, Miguel 

Canals, J. Arnaldo Cedeño, Danilo Chinea, Livia 

Colón, José M. (Papo) López, George R. Proctor, 

Lillian Ramírez, Anne Reilly, Jeanine Vélez, Ivan 

Vicens, and Papo Vives. 

Numerous colleagues have contributed to the 

discussion of various aspects of this work. Dan 

Nicolson (US) has been an invaluable resource in 

discussions related to nomenclatural problems, as 

well as several aspects of the classification of the 

Araceae. Gisela Sancho (LP) edited the whole

16 


book and contributed greatly to it through her 

corrections and suggestions on the first draft. 

Similarly, I thank various colleagues, experts in 

specific taxonomic groups, for the help they gave 

me; among them are Christiane Anderson 

(MICH), Daniel F. Austin (FAU), Barry T. 

Hammel (MO), Dave Lellinger (US), James 

Luteyn (NY), Mike Nee (NY), Lauren Raz (NY), 

Harold Robinson (US), and Charlotte M. Taylor 

(MO). I also express my appreciation to those 

colleagues who have assisted diligently in locating 

numerous collections deposited in their respective 

herbaria and in making them available for study 

through loans or electronic images: Jackie 

Kallunki (NY), Thomas A. Zanoni (NY), Eugenio 

Santiago (UPR), J. Carlos Trejo (UPR), Frank 

Axelrod (UPRRP), José L. Chabert (SJ), and Marc 

Pignal (P). I thank Christiane Feuillet (US) for 

her help in locating some of the original 

descriptions deposited in the library of the 

Department of Agriculture in Beltsville, 

Maryland, Scott A. Mori (NY) and James Luteyn 

(NY) for the loan of various illustrations for use 

in this book, and Alice Tangerini for her support 

in the processing of electronic images, designing 

the front cover, assisting on questions regarding 

images, and handling and modifying some of the 

original illustrations prepared by Bobbi Angell. 

My special thanks to Mark T. Strong (US), 

who has diligently accomplished the numerous 

tasks related to the care and handling of the 

herbarium specimens as well as the database, the 

conversion of the illustrations to electronic format, 

the incorporation of the figure legends, and 

proofing of the English language edition. Finally 

I thank Bobbi Angell (NY) for having agreed to 

illustrate the present work both accurately and 

artistically, as well as her splendid company and 

friendship during field studies. 

This book was made possible by the financial 

support of the Smithsonian Institution, 

Washington, DC, through the Biological Surveys 

and Inventories Program, the Walcott Fund of the 

Department of Botany, which financed a large 

number of the illustrations, the International 

Institute of Tropical Forestry, Río Piedras, Puerto 

Rico, and the Atherton Seidell Grant Program at 

the Smithsonian Institution for providing funding 

for the translation of this work from Spanish into 

English.


Acevedo-Rodríguez, P. 1984. Nuevo record para 

el Nogal de las Antillas (Juglans jamaicensis 

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70. 

———.1985. Los bejucos de Puerto Rico Vol. 1. 

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——— et al., 1996. Flora of St. John, U.S. Virgin 

Islands. Mem. New York Bot. Gard. Vol. 78: 

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——— and F. S. Axelrod. 1999. Annotated checklist 

for the tracheophytes of Río Abajo Forest 

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Ackerman, J. D. 1992. The orchids of Puerto Rico 

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848 pp. 

Breckon, G. 2000. A revision of the flora of 

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of dicotyledon wood. Springer-Verlag, New 

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García, R. G. 1991. Relaciones taxonómicas y 

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serpentina en Susúa, Puerto Rico y Río Piedras, 

Gaspar Hernández, República Dominicana. 

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of climbing plants. Pp. 3-51. In: Putz, F. 

E. and H. A. Mooney (eds.) The biology of 

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Antilles. Arnold Arboretum of Harvard University, 

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tropics. Federal Experimental Station in Puerto 

Rico, Mayagüez, P.R. Circular No. 31. U.S. 

Department of Agric. Washington, D.C. 72 pp. 

Jacobs, M. 1988. The tropical rain forest. Springer- 

Verlag. Berlin. 

Krebs, H. 1847. Et Bidrag til St. Thomas’ Flora. 

Naturhist. Tidsskr. 1847: 291-302. 

Liogier, H. A. 1965. Nomenclatural changes and 

additions to Britton and Wilson’s ‘’Flora of 

Porto Rico and the Virgin Islands”. Rhodora 

67: 315-361. 

———. 1967. Further changes and additions to 

the flora of Porto Rico and Virgin Islands. 

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———. 1985-1997. Descriptive flora of Puerto 

Rico and adjacent islands, Spermatophyta. Vols. 

1-5. Editorial de la Universidad de Puerto Rico, 

Río Piedras, P.R. 

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Rico and adjacent islands: a systematic synopsis. 

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Virgin Islands). U.S. Forest Service Research 

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trees of Puerto Rico and the Virgin Islands. 

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Service, Washington, D.C. 556 pp. 

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the world. Hearthside Press Inc., Great Neck, 

New York. 410 pp. 

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68: 441-546. 

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1945. Catálogo de los nombres vulgares y 

científicos de algunas plantas puertorriqueñas. 

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names in Pteridophyta. Royal Botanic 

Gardens, Kew. 

Proctor, G. R. 1989. Ferns of Puerto Rico and the 

Virgin Islands. Mem. New York Bot. Gard. 53: 

1-389. 

———. 1994. Manual de manglares de Vieques. 

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Ambientales. San Juan, Puerto Rico. 72 pp. 

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Barro Colorado Island, Panama. Ecology 65: 

1713-1724. 

Quevedo, V., S. Silander, and R.O. Woodbury. 

1990. Plantas críticas y en peligro de extinción 

en el bosque de Guánica. Acta Científica 4: 137- 

150. 

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of Taxonomic Literature; http:// 

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KRHomeExt.html


Schenk, H. 1893. Biologie und Anatomie der 

Lianen. II Theil. Beiträge zur Anatomie der 

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Rico. Tomo I-III. (Second Edition). Federal 

Emergency Relief Admin., San Juan, P.R. 881 

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I. (ed.) Symbolae antillanae. Vol 4: 1-771. 

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Virgin Islands English Creole. Published by the 

author. St. John, U.S. Virgin Islands. 

———. 1990. What a Pistarkle. A glossary of 

Virgin Islands English Creole. New supplement. 

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Virgin Islands. 

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P.R. 497 pp. 

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Croix med en kort Udsigt over St. Thomas, St. 

Jean, Tortola, Spanishtown, og Crabeneiland. 

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of Puerto Rico: a committee report. Department 

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cooperation with the Puerto Rican Department 

of Natural Resources, San Juan, P.R. 165 pp. 

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Reef National Monument (U.S. Virgin 

Islands). U.S. Forest Service Res. Paper ITF- 

19: 1-27. 

———, L. F. Martorell, and J.G. García-Tudurí. 

1977. The flora of Mona and Monito Islands, 

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Rico, Agricultural Experimental Station Bulletin 

252. 60 pp. 

——— and B. M. Marriott. 1994. Catalog of the 

vascular plants of Cayo Santiago, Puerto Rico. 

Caribb. J. Science 30: 130-142.

20 



SYSTEMATIC TREATMENT 

KEY TO THE DIVISIONS 

1a. Sexual reproduction by sporangia, clustered in sori, which are arranged on the abaxial surface or 

the margin of the blade of the fronds ............................................. PTERIDOPHYTES (Page 21) 

1b. Sexual reproduction by flowers. ........................................................................................................ 2 

2a. Flowers 5-6-merous; leaf blade with reticulate venation, the last free veins of the network forming 

an areole; the petioles not forming a sheath around the stem ............................................................ 

........................................................................................................... DICOTYLEDONS (Page 41) 

2b. Flowers usually trimerous; leaf blade usually with parallel venation, the last-order nerves forming 

a network, never free within the areole; petioles sheathing, covering the stem 

..................................................... ........................................ MONOCOTYLEDONS (Page 403) 

PTERIDOPHYTES 

Key to the families 

1a. Fronds microphyllous (minute), with only one vein................................................Selaginellaceae 

1b. Fronds megaphyllous; venation well developed, with primary, secondary, or tertiary veins.........2 

2a. Sori marginal or submarginal on the fronds ........................................................ Dennstaedtiaceae 

2b. Sori on the abaxial surface of the frond ............................................................................................ 3 

3a. Fronds monomorphic, the fertile and sterile ones similar ............................................................... 4 

3b. Fronds dimorphic, the fertile ones significantly different from the sterile ones ............................. 7 

4a. Fern scandent or climbing by the twining leaf rachis, with indeterminate growth ........................ 5 

4b. Fern climbing by adventitious roots .................................................................................................. 6 

5a. Sori naked (without indusia), dispersed on the abaxial surface of the frond, usually near the midvein 

................................................................................................... Gleicheniaceae 

5b. Sori covered with an indusium in the form of a pocket, produced on digitiform marginal projections 

of the pinnae .................................................................................................................. Schizaeaceae 

6a. Blades of the fronds entire ............................................................................................ Davalliaceae 

6b. Blades of the fronds pinnatisect ................................................................................ Polypodiaceae 

7a. Sori dispersed on the lower surface, not grouped; indusium absent. ................. Lomariopsidaceae 

7b. Sori in elongate-linear groups, lateral to the midvein; indusium entire, elongate ..... Blechnaceae

22 


1. Family BLECHNACEAE 

1. BLECHNUM 

Terrestrial ferns, hemiepiphytic or less frequently epiphytic; rhizomes elongate, erect or scandent, 

sometimes forming a caudex or stolon, densely covered with scales. Fronds sterile or fertile, monomorphic 

or dimorphic; blades glabrous, pinnatisect, simple-pinnate or less frequently entire; veins free, bifurcated, 

except for the minor veins on the fertile fronds; pinnae entire or serrate. Sori linear, parallel to the 

midvein; indusia linear, opening toward the midvein. A cosmopolitan genus of about 180 species, 

mostly in the Southern Hemisphere. 

1. Blechnum fragile (Liebm.) C. V. Morton & 

Lellinger, Amer. Fern J. 57: 68. 1967. 

Fig. 5. A-C 

BASIONYM: Lomaria fragilis Liebm. 

Slightly woody fern, hemiepiphytic, scandent 

by means of adventitious roots, attaining 2 m in 

length. Rhizomes thick, 0.8-1.5 cm wide, densely 

covered with subulate, slihtly undulate, ciliate, 

ferruginous scales, 8-17 mm long. Fronds more 

or less clustered, arching, ascendent or pendulous, 

dimorphic; blades pinnate, membranaceous. 

Sterile fronds 40-60 × 4-12 cm, with an elliptical 

outline, the apex acuminate or caudate, the base 

acute (the pinnae gradually reduced in size toward 

the base); pinnae 25-75 pairs, oblong-deltate, the 

apex acuminate, the margins revolute, sometimes 

with minute callus-like projections; the upper 

2. Family DAVALLIACEAE 

1. OLEANDRA 

surface dull, with a prominent midvein; the lower 

surface dull, paler than the upper surface, with a 

prominent midvein; stipe 10-18 cm long, blackish 

or brown, with some vestigial pinnae. Fertile 

fronds slightly smaller than the sterile ones; 

pinnae 28-41 pairs, linear; indusia half as wide 

as the pinna. 

Phenology: Found in fertile condition 

throughout the year. 

Status: Native, common. 

Distribution: In moist forests at middle and 

upper elevations along the Cordillera Central and 

the Sierra de Luquillo. Also in Jamaica, Cuba, 

Hispaniola, southern Mexico, and Central 

America. 

Public Forests: Carite, El Yunque, Guilarte, 

and Toro Negro. 

Epiphytic or terrestrial ferns with very elongate rhizomes, erect, creeping, or climbing, sometimes 

branched, densely covered with peltate scales. Fronds sterile or fertile, monomorphic; blades simple, 

entire; veins free and parallel; stipe articulate. Sori circular, usually adjacent to the midvein; indusia 

circular or reniform, peltate. A tropical genus of about 40 species. 

1. Oleandra articulata (Sw.) C. Presl, Tent. 

Pterid. 78. 1836. 

Fig. 5. D-E 

BASIONYM: Aspidium articulatum Sw. 

SYNONYM: Oleandra nodosa (Willd.) C. Presl 

Herbaceous fern, epiphytic or climbing by 

adventitious roots, attaining 5 m in length. 

Rhizomes slender, densely covered with 

lanceolate, ferruginous scales. Fronds pendulous; 

blades (15) 23-39 (42) × (2.5) 3-6 cm, 

membranaceous, oblong, elliptical, or 

sublanceolate, the apex acute, acuminate, or 

caudate, the base acute or acuminate, sometimes 

unequal, the margin crenulate or crenate, slightly 

revolute and ferruginous; the upper surface shiny,

Vines and Climbing Plants of Puerto Rico and the Virgin Islands 23

24 


with a prominent, sulcate midvein; the lower 

surface dull, with a prominent, blackish or shiny 

gray midvein, covered with ferruginous scales; 

stipe (4) 8-16 cm long, blackish or gray, glabrous, 

articulated near the base. Sori circular, light 

brown, distributed along both sides of the midvein. 





upper elevations along the Cordillera Central and 

the Sierra de Luquillo. Also in Jamaica, Cuba, 

Hispaniola, the Lesser Antilles, and throughout 

the Neotropics. 


Maricao, Río Abajo, and Toro Negro. 

Reference: Maxon, W. R. 1914. The American species of Oleandra. Contrib. U.S. Natl. Herb. 17: 

392-398. 

3. Family DENNSTAEDTIACEAE 

Key to the genera 

1a. Rhizomes without scales; sori marginal or submarginal, at the tips of the veins, covered by the 

reflexed margins of the blade ......................................................................................... 1. Hypolepis 

1b. Rhizomes covered with scales; sori in marginal pockets ......................................... 2. Odontosoria 

1. HYPOLEPIS 

Terrestrial ferns; rhizomes creeping, woody, elongate, quite branched, covered with multicellular 

hairs and without scales. Fronds monomorphic, separated, elongate, usually clambering and spiny; 

blades 2-5-pinnate; pinnules lobed or pinnatifid, glabrous or pubescent, sometimes with viscous glands; 

veins free; stipe and rachis elongate, robust, not articulate, sometimes spiny. Sori circular, marginal or 

submarginal, solitary at the tips of the veins; indusium false, formed by the reflexed margins of the 

blade. A pantropical genus of about 45 species. 

Key to the species of Hypolepis 

1a. Stipe and rachis densely spiny........................................................................................................... 2 

1b. Stipe and rachis unarmed or with some scattered spines ................................................................. 3 

2a. Leaf rachis brown, with some of the spines recurved and dark brown; blades 4-pinnate-pinnatifid; 

pinnules distally emarginate .................................................................................... 1. H. nigrescens 

2b. Leaf rachis creamy yellow, with straight, pale spines; blades 3-pinnate-pinnatifid; pinnules distally 

obtuse, entire ................................................................................................................... 2. H. repens 

3a. Leaf blade membranaceous with light brown venation; the upper surface with a flat midvein; the 

lower surface glabrous or puberulent ....................................................................... 3. H. tenerrima 

3b. Leaf blade chartaceous with yellowish venation; the upper surface with a sulcate midvein; the 

lower surface pubescent ................................................................................................ 4. H. urbanii


1. Hypolepis nigrescens Hook., Sp. Fil. 2: 66, t. 

90c. 1852. 

Fig. 6. A 

Clambering fern. Rhizomes branched, dark 

brown, pubescent. Fronds 4-pinnate-pinnatifid, 

with indeterminate growth, 5-10 m long; blades 

chartaceous, narrowly oblong-lanceolate in 

outline; upper surface glabrous or less frequently 

puberulent; lower surface with scattered hairs 

along the main veins; pinnae opposite or 

subopposite, broadly lanceolate, up to 35 cm long; 

primary pinnules alternate; blades 18-24 cm long; 

stipe and rachis reddish to dark brown, with 

numerous dark brown straight or recurved spines. 

Sori circular, submarginal; indusia very narrow, 

glabrous. 

Phenology: Collected in fertile condition in 

February, July, and September. 

Status: Native, rare. 

Distribution: Known from three localities in 

Puerto Rico, in the interior or at the edges of moist 

forests. Also in the Greater Antilles and from 

Mexico to Bolivia. 

Public forest: Toro Negro. 

2. Hypolepis repens (L.) C. Presl, Tent. Pterid. 

162. 1836. 

Fig. 6. B 

BASIONYM: Lonchitis repens L. 

SYNONYM: Hypolepis aculeata (Spreng.) J. Sm. 

Clambering fern. Rhizome covered with 

brown hairs. Fronds ascending, clambering or 

arching, 3-pinnate-pinnatifid, 2-4 m long; blades 

coriaceous, triangular in outline, up to 1.5 m wide; 

upper surface with scattered hairs along the 

midvein; lower surface shiny, pubescent; pinnae 

lanceolate-triangular, 20-50 cm long; primary 

pinnules subopposite, 7-22 cm longstipe and 

rachis yellowish or light brown, with straight 

spines, the stipe darker toward the base. Sori 

circular, submarginal; indusia membranaceous, 

whitish, subentire. 


August and September. 


Distribution: In moist secondary and disturbed 

forests along the Cordillera Central and the Sierra 

de Luquillo. Also in the Antilles (except Jamaica) 

and continental tropical America. 

Public forests: Carite, El Yunque, Guilarte, 

Maricao, and Toro Negro. 

3. Hypolepis tenerrima Maxon, J. Wash. Acad. 

Sci. 14: 196. 1924. 

Fig. 6. C 

Clambering fern. Fronds weakly ascending or 

clambering, 2-3-pinnate-pinnatifid, up to 2.5 m 

in length; blades membranaceous, elongatetriangular 

in outline, 1-1.5 m wide at the base; 

glabrous or puberulent, with articulate or 

glandular hairs on both surfaces, the venation 

light brown; pinnae subopposite or alternate, up 

to 80 cm long, obliquely oblong-deltoid to oblongovate, 

acuminate; rachis puberulent, unarmed or 

with some scattered straight spines; primary 

pinnules subopposite or alternate, 16-20 cm long, 

narrowly oblong-lanceolate in outline; secondary 

pinnules alternate, 2.5-4 cm long, light to dark 

brown, oblong, obtuse; stipe and rachis light to 

dark brown, glabrescent, with non-pointed 

outgrowths. Sori submarginal; indusia 

membranaceous, greenish yellow, crenate. 


March. 

Status: Native, uncommon. 

Distribution: Known in Puerto Rico from two 

collections made by Paul Sintenis at the end of 

the nineteenth century from primary forests in 

Adjuntas (Sintenis 4105) and Utuado (Sintenis 

6454). Also in Hispaniola. 


4. Hypolepis urbanii Brause in Urb., Symb. 

Antill. 9: 344. 1925. 

Fig. 6. D 

Clambering fern. Rhizomes covered with 

brown articulate hairs. Fronds clambering, 3pinnate-pinnatifid, 

3 m or more in length; blades 

coriaceous, pubescent on the lower surface, 

triangular in outline, up to 1 m wide at the base, 

the venation yellowish; pinnae subopposite, up 

to 50 cm long, oblong-deltoid, acuminate; rachis 

glabrescent or slightly muricate; primary pinnules 

subopposite or alternate, 3-15 cm long, narrowly 

oblong-deltoid in outline; stipe and rachis light 

brown, dark toward the base, glabrescent, 

unarmed. Sori submarginal; indusia 

membranaceous, yellowish, glabrous, crenate.

26 




February. 


Distribution: Known in Puerto Rico from only 

2. ODONTOSORIA 

one collection from Monte Jayuya (Proctor 

40130). Also in Hispaniola. 


Clambering ferns, terrestrial with short, slender, creeping rhizomes, covered with scales. Fronds 

monomorphic, elongate, clambering, usually spiny; blades narrowly triangular to linear, 2-5-pinnate; 

pinnae and pinnules forming a right angle with the axis or rachis; pinnules linear to flabellate. Sori in 

marginal pockets, containing three sporangia; indusia absent. A neotropical genus of 12 species. 

Key to the species of Odontosoria 

1a. Rachis of the fronds spiny .......................................................................................... 1. O. aculeata 

1b. Rachis of the fronds unarmed ................................................................................... 2. O. scandens 

1. Odontosoria aculeata (L.) J. Sm., Cult. Ferns 

67. 1857. 

Fig. 7. A-C 

BASIONYM: Adiantum aculeatum L. 

SYNONYMS: Davallia aculeata (L.) Sm. 

Davallia dumosa Sw. 

Clambering fern. Rhizomes short, creeping, 

densely covered with reddish brown scales at the 

apex. Fronds 2-3-pinnate, glabrous, clambering, 

up to 3 m in length; pinnae numerous, opposite, 

20-40 cm long, ovate-deltoid; pinnules alternate; 

secondary pinnules (2) 3 (5) times laciniate, 

cuneiform, the apex usually revolute; rachis spiny. 

Phenology: Probably found in fertile condition 



Distribution: In secondary forests and moist 

disturbed areas from sea level to 950 m. Also in 

St. John, St. Thomas, and Tortola; Cuba and 

Hispaniola. 



2. Odontosoria scandens (Desv.) C. Chr., Ind. 

Fil. 354, 465. 1906. 

Fig. 7. D-F 

BASIONYM: Humata scandens Desv. 

SYNONYM: Odontosoria uncinella (Kunze) Fée 

Clambering fern. Rhizomes creeping, scarcely 

elongate, densely covered with reddish brown 

scales at the apex. Fronds 3-pinnate, glabrous, 

clambering or twining, up to 2 m in length; pinnae 

numerous, opposite or subopposite, 10-18 cm 

long, lanceolate or oblong; pinnules subopposite 

or alternate; secondary pinnules bilobed or 

laciniate, cuneiform, coriaceous, the apex not 

revolute; rachis yellowish, unarmed (rarely with 

scattered small spines), smooth. 




Distribution: Sporadic along the Cordillera 

Central, in moist secondary and disturbed forests. 

Also in Cuba and Hispaniola. 

Public forests: El Yunque, Guilarte, Maricao, 


Reference: Barcelona, J. F. 2000. Systematics of the genus Odontosoria sensu lato (Lindsaeaceae). 

Doctoral thesis, Miami University, Oxford, Ohio

28 


4. Family GLEICHENIACEAE 


1a. Leaf blade with secondary venation (between the midvein and the margin) 2-5 times bifurcate; 

rhizomes and fronds without scales ....................................................................... 1. Dicranopteris 

1b. Leaf blade with secondary venation (between the midvein and the margin) only once bifurcate; 

rhizomes and fronds with scales (fronds with scales at least at the apical meristems)....... 2. Gleichenia


1. DICRANOPTERIS 

Terrestrial ferns; rhizomes elongate, creeping with setulose trichomes, without scales. Fronds 

monomorphic, with periodic indeterminate growth, 4-5 times bifurcate; pinnae, pinnules, and secondary 

pinnules opposite, the secondary venation 2-4 times bifurcate, the lower surface glaucous; petioles 

brittle and glabrous. Sori rounded, without an indusium, on the secondary veins, with 6-15 sessile 

sporangia. A tropical genus of about 10 species. 

Key to the species of Dicranopteris 

1a. Area of frond bifurcation with a pair of accessory pinnae; tertiary rachis of the fronds slightly 

angular; lower surface pale green, glabrous ............................................................... 1. D. flexuosa 

1b. Area of frond bifurcation without accessory pinnae or sometimes with accessory pinnae only in the 

primary bifurcations; tertiary rachis of the fronds with narrow longitudinal keels; lower surface 

glaucous, glabrous, or sometimes with scattered stellate hairs ................................ 2. D. pectinata 

1. Dicranopteris flexuosa (Schrad.) Underw., 

Bull. Torrey Bot. Club 34: 254. 1907. 

Fig. 8. A-E 

BASIONYM: Mertensia flexuosa Schrad. 

SYNONYM: Gleichenia flexuosa (Schrad.) Mett. 

Terrestrial fern. Rhizomes creeping, with 

deciduous setose trichomes. Fronds clambering 

or decumbent, 1-2 m long, forming a dense thicket 

difficult to penetrate; pinnae 4 times bifurcate, 

with a pair of accessory pinnae of reduced size 

and a tuft of rigid, reddish trichomes in the area 

of bifurcation; rachis cylindrical and brittle, the 

primary rachis with proleptic indeterminate 

growth, the secondary or tertiary rachis of 

determinate growth, the latter angular; secondary 

pinnules oblong-deltoid to linear, glabrous, the 

secondary venation 2-4 times bifurcate, the apex 

obtuse, the margins revolute; lower surface pale 

green, glabrous. Sori reddish brown, 0.2-0.4 mm 

in diameter. 



Status: Native, very common. 

Distribution: Widely distributed in moist 

disturbed areas along varying elevational 

gradients. Also in the Antilles, the United States 

(Florida), Central America, and tropical South 

America. 

Public forests: Carite, El Yunque, Maricao, 

Río Abajo, Toro Negro, and Tortuguero. 

2. Dicranopteris pectinata (Willd.) Underw., 

Bull. Torrey Bot. Club 34: 260. 1907. 

Fig. 8. F-G 

BASIONYM: Mertensia pectinata Willd. 

Terrestrial fern. Rhizomes creeping, scabrous, 

light brown, with deciduous articulate hairs. 

Fronds clambering or decumbent, 1-2 m long, 

forming a dense thicket difficult to penetrate; 

pinnae unequally bifurcate, with a tuft of rigid, 

reddish trichomes in the area of bifurcation, but 

without accessory pinnae or these only in the 

primary bifurcations; axes cylindrical and brittle, 

primary rachis proleptic, with sympodial growth, 

tertiary rachis with two narrow longitudinal keels; 

leaf segments oblong-deltoid, the secondary 

venation 3-5 times bifurcate, the apex retuse, the 

margins revolute; lower surface glaucous, 

glabrous, or sometimes stellate-pubescent. Sori 

yellowish, 0.5-0.7 mm in diameter. 




Distribution: Widely distributed in moist 

disturbed areas at various elevations. Of wide 

distribution in the Neotropics. 


and Río Abajo.

30 



2. GLEICHENIA 

Terrestrial ferns; rhizomes creeping, elongate, slender, covered with scales and setose or pilose 

trichomes. Fronds monomorphic, elongate, with indeterminate growth, bifurcate, bipinnate, usually 

clambering; blades with secondary veins free, only once bifurcate; area of bifurcation covered with 

scales; rachis elongate, cylindrical, and rigid. Sori rounded, without an indusium, containing 3-6 

sporangia. A pantropical and subtropical genus of about 110 species. 

Key to the species of Gleichenia 

1a. Lower surface of the pinnae glaucous, glabrous except for the main veins, which are covered with 

reddish scales or cilia ............................................................................................... 3. G. rubiginosa 

1b. Lower surface of the pinnae tomentose or woolly ............................................................................ 2 

2a. Lower surface of the pinnae densely ferruginous-tomentose to woolly; principal rachis of the young 

pinnae densely covered with pale brown, ciliate scales on the upper surface, of woolly 

appearance.........................................................................................................................1. G. bifida 

2b. Lower surface of the pinnae white-woolly; principal rachis of the young pinnae covered with a few 

brown-purple, short-ciliate, and deciduous scales on the lower surface ................. 2. G. brevipubis 

1. Gleichenia bifida (Willd.) Spreng., Syst. Veg. 

ed. 16, 4: 27. 1827. 

Fig. 9. A-C 

BASIONYM: Mertensia bifida Willd. 

Terrestrial fern. Rhizomes creeping, reddish 

brown, with attenuate, ciliate scales at the apex. 

Fronds clambering or decumbent, 1-2 m in length, 

forming a dense thicket difficult to penetrate; 

pinnae 3 times bifurcate, with small accessory 

pinnae and densely covered with scales in the 

bifurcation; rachis cylindrical, those of the young 

pinnae covered with pale brown ciliate scales; 

primary rachis with proleptic indeterminate 

growth, the secondary ones short; terminal pinnae 

20-35 × 3-5 cm, the midvein densely covered with 

ferruginous scales on the lower surface; secondary 

pinnules oblong-deltoid, the apex obtuse; upper 

surface glabrous; lower surface densely covered 

with ferruginous scales. Sori ferruginous. 




Distribution: Widely distributed in Puerto 

Rico, in moist disturbed areas at middle to upper 

elevations. Also throughout the Neotropics. 

Public forests: Carite, Guilarte, El Yunque, 

Maricao, and Río Abajo. 

2. Gleichenia brevipubis C. Chr., Bull. Herb. 

Boissier Ser. 2, 6: 280. 1906. 

Fig. 9. D 

SYNONYM: Sticherus brevipubis (C. Chr.) A.R. Sm. 


brown, glabrous or with some scattered shortciliate 

scales. Fronds clambering or decumbent, 

1-2 m in length, forming a dense thicket, difficult 

to penetrate; pinnae 2-3 times bifurcate, sparsely 

covered with scales in the bifurcation; rachis 

cylindrical, glabrescent, the primary rachis with 

proleptic indeterminate growth, the secondary 

ones short; terminal pinnae 20-35 cm long, with 

a few brown-purple, short-ciliate, and deciduous 

scales along the midvein on the lower surface; 

secondary pinnules oblong-deltoid, the apex 

obtuse; the upper surface glabrous; the lower 

surface white-woolly. Sori ferruginous. 


August. 


Distribution: Known in Puerto Rico from two 

recent collections (Proctor 39420 and 41419). 

Also from Mexico to Costa Rica. 

Public forest: Toro Negro.

32 



3. Gleichenia rubiginosa Mett., Ann. Sci. Nat. 

Bot. Sér. 5, 2: 267. 1864. 

Fig. 9. E 

SYNONYM: Dicranopteris rubiginosa (Mett.) Maxon 


brown, with attenuate, ciliate scales at the apex. 

Fronds clambering or creeping, 1-2 m in length; 

pinnae 2-3 times bifurcate, with a tuft of scales in 

the bifurcation; rachis cylindrical, with some 

scattered hairs, the primary rachis with proleptic 

indeterminate growth, the secondary ones not 

developed; pinnae 10-25 cm long, the lower 

5. Family LOMARIOPSIDACEAE 


surface glaucous, glabrous except for the main 

veins, which are covered with reddish scales or 

cilia; secondary pinnules oblong, the apex retuse. 

Sori yellowish. 


February and August. 


Distribution: Restricted to the highest areas 

of the Cordillera Central, in moist disturbed areas, 

on roadsides and in forests. Also in mountainous 

areas of Venezuela, Colombia, Ecuador, and Peru. 

Public forests: Guilarte and Toro Negro 

1a. Secondary venation reticulate.................................................................................. 1. Lomagramma 

1b. Secondary venation free............................................................................................. 2. Lomariopsis 

1. LOMAGRAMMA 

Terrestrial ferns; rhizomes elongate, climbing by adventitious roots, covered with scales. Fronds 

short-stipitate, dimorphic, the fertile ones with narrower pinnae than the sterile ones; blades pinnate; 

pinnae membranaceous, glabrous, with serrate margins and reticulate venation (free only near the 

margin); lateral pinnae deciduous by means of a basal articulation; terminal pinna not articulate. 

Fertile fronds with sporangia scattered on the abaxial surface, rarely on the margins; indusia absent. A 

genus of 20 species, Paleotropical with the exception of L. guianensis, which is widely distributed in 


1. Lomagramma guianensis (Aubl.) Ching, 

Amer. Fern J. 22: 17. 1932. 

Fig. 10A-B 

BASIONYM: Polypodium guianense Aubl. 

Terrestrial fern. Rhizomes climbing by aerial 

roots, reddish brown, 3-7 mm wide, with deltoidattenuate 

scales, 1-1.5 mm long. Fronds 

horizontal or pendulous. Sterile fronds 0.45-1 m 

long; stipes 6-20 cm long, dark brown, glabrous 

or with some scattered scales; rachis cylindrical 

toward the base, narrowly winged toward the 

apex; pinnae 20-50, alternate or subopposite, 

elliptical to lanceolate, chartaceous, 6-14 × 1-2.3 

cm, the apex long-acuminate, the base cuneate, 

unequal, the margins serrate, glabrous or with 

some scattered hairs on the lower surface; midvein 

prominent on both surfaces, secondary venation 

reticulate, slightly prominent, yellowish. Fertile 

fronds smaller than the sterile ones; pinnae linear, 

2-5 × 2-4 mm, the apex obtuse, the base obtuse; 

lower surface densely covered with ferruginous 

sporangia. 


May. 


Distribution: In moist forests at lower and 

middle elevations. Also in Cuba, Hispaniola, 

Venezuela, French Guiana, and Brazil. 

Public forest: Maricao.

34 



2. LOMARIOPSIS 

Terrestrial or hemiepiphytic ferns; rhizomes climbing by aerial roots, elongate, somewhat 

compressed, covered with scales. Fronds short-stipitate, dimorphic, the fertile ones with narrower 

pinnae than the sterile ones; blades pinnate, pinnae membranaceous, with serrate margins and free 

venation, simple or bifurcate, the lateral pinnae deciduous by means of a basal articulation, the terminal 

pinna not articulate. Fertile fronds densely covered with sporangia on the lower surface; indusia absent. 

A pantropical genus of 45 species, 15 of which are found in the Neotropics. 

Key to the species of Lomariopsis 

1a. Pinnae of the sterile fronds unequal at the base, with inconspicuous venation; pinnae of the fertile 

fronds ca. 2 mm wide ........................................................................................ 1. L. amydrophlebia 

1b. Pinnae of the sterile fronds equilateral at the base, with prominent venation on the lower surface; 

pinnae of the fertile fronds 3-8 mm wide .......................................................................................... 2 

2a. Margin of the sterile pinnae acutely incised-serrate; rhizomes sparsely covered with reddish brown, 

lanceolate-caudate, denticulate to ciliate scales, 2-5 mm long; pinnae 20-46.......... 2. L. kunzeana 

2b. Margin of the sterile pinnae finely serrate; rhizomes densely covered with light brown, narrowly 

lanceolate-caudate, ciliate scales, 8-12 mm long; pinnae 30-54 .............................. 3. L. sorbifolia 

1. Lomariopsis amydrophlebia (Sloss. ex 

Maxon) Holttum, Kew Bull. 1939: 617. 1940. 

Fig. 10. C 

BASIONYM: Stenochlaena amydrophlebia Sloss. ex Maxon 


roots, dark brown, 4-8 mm wide, densely covered 

with ferruginous, lanceolate-caudate scales, 6-9 

mm long. Sterile fronds 30-70 cm long; stipes 

12-18 cm long, dark brown, sulcate, with some 

scattered glandular hairs and scales; rachis 

narrowly winged; pinnae 22-34, alternate, oblonglanceolate, 

falcate, chartaceous, 3.5-9 × 1-1.7 cm, 

puberulent on both surfaces, the venation 

inconspicuous, parallel, free, the apex obtuse to 

acuminate, the base cuneate, unequal, the margins 

crenate to finely serrate, revolute; lower surface 

with the midvein lighter in color. Fertile fronds 

smaller than the sterile ones; pinnae linear, 4-9 

cm × ca. 2 mm, the apex obtuse, the base obtuse; 

lower surface densely covered with light brown 

sporangia. 


April. 


Distribution: In moist forests of the Cordillera 

Central and the Sierra de Luquillo. Also in the 

Dominican Republic. 

Public forests: Carite, El Yunque, and Toro 

Negro. 

2. Lomariopsis kunzeana (Underw.) Holttum, 

Kew Bull. 1939: 617. 1940. 

Fig. 10. D 

BASIONYM: Stenochlaena kunzeana Underw. 


roots, light brown, slightly compressed or obtusely 

triangular, with longitudinal keels, 5-10 mm wide, 

sparsely covered with ferruginous, lanceolatecaudate 

scales, 2-5 mm long. Sterile fronds 40- 

90 cm long; stipes 8-18 cm long, light brown, 

sulcate, with some scattered glandular hairs and 

scales; rachis narrowly winged; pinnae 20-46, 

alternate or subopposite, lanceolate, chartaceous, 

4-11 × 1-1.5 cm, the apex acuminate, the base 

cuneate, equilateral, the margins acutely incisedserrate, 

slightly revolute; the upper surface 

glabrous, the midvein compressed; lower surface 

puberulent, the midvein rather prominent, 

yellowish, the secondary venation conspicuous, 

parallel, free. Fertile fronds smaller than the sterile 

ones; pinnae oblong, 4-8 mm wide, the apex acute, 

the base obtuse; lower surface densely covered 

with light brown sporangia.

36 



July and August. 


Distribution: Known in Puerto Rico from few 

collections from the region of Utuado, Lares, and 

Bayamón, probably from forests on mogotes. Also 

in the United States (Florida), Cuba, and 

Hispaniola. 

3. Lomariopsis sorbifolia (L.) Fée, Mém. Foug. 

2: 69. 1845. 

Fig. 10. E 

BASIONYM: Acrostichum sorbifolium L. 

Terrestrial fern, 1-1.5 m long. Rhizomes 

scandent by aerial roots, light brown, slightly 

compressed or obtusely triangular, with 

longitudinal keels, sometimes sulcate, 5-10 mm 

wide, densely covered with light brown, narrowly 

lanceolate, ciliate, imbricate scales, 8-12 mm long. 

Sterile fronds 25-115 cm long; stipes 5-30 cm 

long, light brown, sulcate, with scattered scales; 

rachis narrowly winged; pinnae 30-52, alternate 

or subopposite, ovate to lanceolate, chartaceous, 

6-13 × 1.5-2.3 cm, with the midvein prominent 

on both surfaces, the apex long-acuminate to 

caudate, the base cuneate, equilateral, the margins 

finely serrate, slightly revolute; lower surface with 

the secondary venation conspicuous, parallel, free. 

Fertile fronds smaller than the sterile ones; pinnae 

linear, 4-8.5 × 3-5 mm, the apex obtuse, the base 

cuneate; lower surface densely covered with dark 

brown sporangia. 

Phenology: Collected in fertile condition from 

May to November. 


Distribution: Of wide distribution in all of 

Puerto Rico, especially in the northwest. Also in 

St. Thomas; Hispaniola and the Lesser Antilles. 

Public forests: El Yunque, Maricao, and Río 

Abajo. 

Reference: Moran, R.C. 2000. Monograph of the neotropical species of Lomariopsis 

(Lomariopsidaceae). Brittonia 52: 55-111. 

6. Family POLYPODIACEAE 


1a. Blades simple, entire............................................................................................... 1. Microgramma 

1b. Blades pinnatisect ....................................................................................................... 2. Polypodium 

1. MICROGRAMMA 

Epiphytic or rarely saxicolous ferns. Rhizomes elongate, creeping or climbing, covered with 

lanceolate or acicular scales, peltate and not clathrate. Fronds monomorphic or dimorphic, with the 

fertile ones narrower and longer than the sterile ones, stipitate, glabrous, pubescent or with scales; 

blades simple, with entire margins, the venation reticulate, the areoles with free venules. Sori rounded 

or oblong, in a single row between the rachis and the margin; indusia absent; paraphyses present, 

consisting of narrow scales or pluricellular trichomes. Spores bilateral, verrucose, rugose, or tuberculate. 

A genus of 25 species in the New World and 2-3 species in Africa. 

Key to the species of Microgramma 

1a. Blades with numerous minute, subulate scales; paraphyses conspicuous (longer than the 

sporangia).............................................................................................. 3. M. piloselloides 

1b. Blades glabrous; paraphyses inconspicuous ..................................................................................... 2


2a. Rhizomes 0.5-2 mm wide; blades chartaceous, with deeply crenate margins ... 1. M. heterophylla 

2b. Rhizomes 2-4 mm wide; blades coriaceous, with entire or slightly crenate margins.............................. 

.............................................................................................................................2. M. lycopodioides 

1. Microgramma heterophylla (L.) Wherry, 

South. Fern Guide 346. 1964. 

Fig. 11. A-B 

BASIONYM: Polypodium heterophyllum L. 

SYNONYM: Polypodium serpens Sw. 

Polypodium exiguum Heward 

Epiphytic or less frequently epilithic fern. 

Rhizomes scandent by minute aerial roots, 1-2 m 

long, filiform, cylindrical, 0.5-2 mm wide, densely 

covered with appressed, ferruginous, attenuate 

scales, 5-7 mm long, peltate above the base. 

Fronds dimorphic, 3-19 × 0.4-1.7 cm; stipes 

slender, 4-20 mm long, yellowish to light brown, 

glabrous; blades entire, chartaceous, elliptical, 

oblong or lanceolate, glabrous, the midvein 

prominent, the secondary venation conspicuous, 

widely reticulate, the apex obtuse or acute, the 

base long-attenuate, the margins crenate. Sori 

rounded, ferruginous, forming a row on each side 

of the midvein; paraphyses inconspicuous. 



Status: Native, quite common. 

Distribution: In moist or dry forests, in the 

coastal lowlands and in the zone of mogotes. Also 

in Vieques, St. Croix, St. John, and Virgin Gorda; 

the United States (Florida), the Bahamas, the 

Greater Antilles, the Cayman Islands, and the 

northern Lesser Antilles. 

Public forests: El Yunque and Río Abajo. 

2. Microgramma lycopodioides (L.) Copel., Gen. 

Fil. 185. 1947. 

Fig. 11. C-D 

BASIONYM: Polypodium lycopodioides L. 

Epiphytic or less frequently epilithic fern. 

Rhizomes scandent by aerial roots, 1-2 m in 

length, more or less cylindrical, 2-4 mm wide, 

densely covered with appressed, ferruginous, 

linear-attenuate scales, 6-10 mm long, peltate 

above the base, caudate at the apex. Fronds 5-13 

× 0.5-2 cm; stipes slender, up to 5 mm long, 

yellowish to light brown, glabrous; blades entire, 

coriaceous, lanceolate or elliptical, glabrous, the 

midvein prominent, the secondary venation 

conspicuous, loosely reticulate, the apex obtuse 

or acute, the base long-attenuate, the margins 

entire or slightly crenate. Sori rounded, 

ferruginous, forming a row on each side of the 

midvein; paraphyses inconspicuous. 




Distribution: In moist forests, along the 

cordilleras and in the zone of mogotes. Also in 

St. John and Tortola. Pantropical. 


and Río Abajo. 

3. Microgramma piloselloides (L.) Copel., Gen. 

Fil. 185. 1947. 

Fig. 11. E-F 

BASIONYM: Polypodium piloselloides L. 

Epiphytic or less frequently terrestrial fern. 

Rhizomes scandent by aerial roots, up to 4 m long, 

more or less cylindrical, 0.5-1 mm wide, more or 

less covered with divaricate, ferruginous, 

attenuate scales that are 3-4 mm long and caudate 

at the apex. Fronds dimorphic, the sterile ones 

elliptical or elliptical-lanceolate, 1.5-8 × 1-1.8 cm. 

Fertile fronds 3.5-10 × 0.3-1.1 cm; stipes slender, 

1.5-4 cm long, light brown, glabrous; blades 

entire, chartaceous, oblong or linear-lanceolate, 

sparsely covered with minute, ferruginous, 

subulate scales, the midvein prominent on the 

lower surface, the secondary venation 

conspicuous, loosely reticulate, the apex obtuse, 

the base long-attenuate, the margins entire or 

slightly crenate. Sori rounded, ferruginous, 

forming a row on each side of the midvein; 

paraphyses longer than the sporangia. 




Distribution: In moist forests, along the 

cordilleras, in the zone of mogotes, or less 

frequently in the coastal lowlands. Also in St. 

Thomas and Tortola; the Greater and Lesser 

Antilles and South America. 

Public forests: El Yunque, Maricao, Río Abajo, 

and Toro Negro.

38 



2. POLYPODIUM 

Epiphytic, epilithic, or rarely terrestrial ferns. Rhizomes short or elongate, creeping or less frequently 

climbing, covered with concolorous or discolorous scales, peltate, clathrate or not clathrate, or less 

frequently glabrous. Fronds articulate at the base, monomorphic, usually stipitate, glabrous, pubescent, 

or with scales; blades pinnatisect or simple-pinnate, the venation branched, free or reticulate, the 

areoles with a single free venule. Sori rounded or elliptical, usually terminal on the free venules; 

indusia absent; paraphyses present or absent. Spores bilateral, verrucose, rugose, or tuberculate, reniform, 

yellow. Polypodium “sensu stricto” is a cosmopolitan genus of about 125 species. 

1. Polypodium loriceum L., Sp. Pl. 1086. 1753. 

Fig. 11. G 

Terrestrial fern. Rhizomes scandent by aerial 

roots, 1-2 m in length, greenish or whitish, with 

light brown spots, cylindrical, 2-5 mm wide, 

sparsely covered with appressed, light brown, 

oblong-circular, peltate scales, 2-4 mm long. 

Fronds 30-80 × 5.5-15(25) cm; stipes 7-24(39) 

cm long, yellowish to light brown, slightly sulcate, 

glabrous; blades chartaceous, oblong-deltate or 

deltate-lanceolate, deeply pinnatisect; segments 

15-40 pairs, alternate or subopposite, oblongdeltate, 

the apex acuminate, the margins entire; 

7. Family SCHIZAEACEAE 

1. LYGODIUM 

upper surface glabrous, the midvein compressed; 

lower surface glabrous or puberulent, with the 

venation somewhat prominent, yellowish, the 

secondary venation conspicuous, reticulate. Sori 

rounded, yellowish, forming a row (rarely two) 

on each side of the midvein. 

Phenology: Apparently in fertile condition 



Distribution: In moist forests in the 

cordilleras. Also throughout the Antilles and 

continental tropical America. 



Terrestrial ferns; rhizomes creeping, short, branched, pubescent. Fronds monomorphic, elongate, 

with indeterminate growth, climbing with twining rachis; pinnae bifurcate, with a latent shoot at the 

apex of the petiolule; secondary pinnules alternate, with free veins or areolate. Sporangia pear-shaped, 

with an apical annulus, individual (not clustered in sori), borne at the apices of the veins, arranged in 

2 rows on marginal projections of the blade. A genus of about 40 species, the great majority tropical, 

75% of them of the Old World. 

1. Lygodium japonicum (Thunb. ex Murray) Sw., 

J. Bot. (Schrader) 1800: 106. 1802. 

Fig. 12. A-C 

BASIONYM: Ophioglossum japonicum Thunb. ex Murray 

Clambering fern. Rhizomes elongate, 

creeping, slender, densely pubescent. Fronds 

tripinnate, puberulent, clambering, 1-2 m in 

length; pinnae bifurcate, with a tuft of trichomes 

in the area of bifurcation; rachis subcylindrical, 

with two longitudinal keels, glabrous or 

puberulent, the primary rachis with indeterminate 

growth, the secondary ones very short, the tertiary 

ones narrowly winged, up to 5 cm long; secondary 

pinnae 5-7 cm long, the distal ones 3-5-dissected, 

the basal ones pinnately compound; secondary

40 



pinnules puberulent, oblong, elongate, with 

prominent venation, the apex obtuse to rounded. 

Sporangia on the lower surface of digitiform 

marginal projections of the basal secondary 

pinnules or on fertile pinnae, markedly dissectedlobed; 

indusium forming a small pocket, which 

opens toward the apex of the leaf blade. 

8. Family SELAGINELLACEAE 

1. SELAGINELLA 



Status: Exotic, naturalized, uncommon. 

Distribution: Native to Japan, cultivated and 

apparently naturalized in several localities in 

Puerto Rico. Widely cultivated in tropical and 

subtropical areas. 

Public forest: Río Abajo. 

Terrestrial herbs, usually creeping or decumbent, rarely erect or clambering. Leaves numerous, 

microphyllous, with a single vein; monomorphic and spirally arranged or dimorphic and distichous. 

Sporophylls in sessile spikes, compact or separated; sporangia axillary on the sporophylls; spores 

dimorphic. A genus of 600-700 species, mostly tropical. 

1. Selaginella willdenovii (Desv. ex Poir.) Baker, 

Gard. Chron. 1867: 783, 950. 1867. 

Fig. 12. D-G 

BASIONYM: Lycopodium willdenovii Desv. ex Poir. 

Clambering herb, 6-10 m in length. Stems 

brittle, with numerous secondary branches up to 

50 cm long, these again branched into short 

branches with determinate growth, which attain 

13 cm long and give the appearance of a 3pinnate, 

triangular leaf. Primary and secondary 

branches greenish or with a reddish shade. Leaves 

oblong-ovate, imbricate and in two layers, the 

upper layer of leaves ca. 1 mm long, the lower 

DICOTYLEDONS 

Key to the families of Dicotyledons 

layer of leaves 3-4 mm long, the apex obtuse, the 

base auriculate to subcordiform, sessile. Strobili 

1.5-2 cm long, green, at the tips of the tertiary 

branches. 


February. 


Distribution: Native to tropical Asia and 

Indonesia, introduced in Puerto Rico as an 

ornamental where it is found to be naturalized 

(Cayey, El Verde, Río Piedras, and Mayagüez). It 

has also become naturalized throughout the 

Neotropics. 

Public forest: El Yunque. 

1a. Plants without leaves or with inconspicuous, minute (< 2 mm long), ovate-triangular 

leaves........................................................................................................................... Key 1 

1b. Plants with well developed leaves (> 5 mm long), simple or compound ........................................ 2 

2a. Leaves compound ..................................................................................................................... Key 2 

2b. Leaves simple or unifoliolate .................................................................................................. Key 3

42 


Key 1: LEAVES ABSENT OR INCONSPICUOUS 

1a. Plants not parasitic, without leaves; stems green, with clustered spines ........................ Cactaceae 

1b. Plants parasitic, with haustoria that penetrate the tissues of the host plant; leaves very reduced, 

ovate-triangular; stems yellow or yellowish green, without spines.................................................. 2 

2a. Inflorescences spicate; anthers dehiscent by two small windows that open upwards; perianth of 3 

tepals; style solitary ........................................................................................................... Lauraceae 

2b. Inflorescences cymose; anthers dehiscent by longitudinal sutures; perianth of a calyx and a tubular 

corolla; styles 2 ............................................................................................................... Cuscutaceae 

Key 2: LEAVES COMPOUND 

1a. Leaves alternate ................................................................................................................................. 2 

1b. Leaves opposite .................................................................................................................................. 8 

2a. Plants climbing by tendrils ................................................................................................................ 3 

2b. Plants twining or scandent, without tendrils .................................................................................... 5 

3a. Tendrils opposite the leaves .................................................................................................. Vitaceae 

3b. Tendrils axillary or lateral to the leaves ........................................................................................... 4 

4a. Leaves palmately lobed or palmatifid; tendrils lateral to the leaves, filiform, simple or branched, 

spiral-shaped .............................................................................................................. Cucurbitaceae 

4b. Leaves pinnate, biternate, or triternate; tendrils axillary, in pairs, spirally twisted, usually at the 

base of the inflorescence ................................................................................................ Sapindaceae 

5a. Plants clambering, with numerous recurved spines ......................................................................... 6 

5b. Plants twining or with twining branches, without spines ................................................................ 7 

6a. Flowers with numerous pistils (apocarpous carpels) that develop into concrescent fleshy follicles to 

form an aggregate fruit ....................................................................................................... Rosaceae 

6b. Flowers with a single unicarpellate pistil that develops into a dry fruit that dehisces by the ventral 

suture (legume) .................................................................................................................... Fabaceae 

7a. Lateral branches sometimes twining and with determinate growth, resembling a tendril; flowers 

with 5 apocarpous pistils, of which usually only one develops (the rest abort) into a fleshy 

follicle........................................................................................................................ Connaraceae 

7b. Lateral branches twining, with indeterminate growth, not forming a structure similar to a tendril 

(except for Dalbergia); flowers with a single unicarpellate pistil that develops into a dry fruit that 

dehisces by the ventral suture (legume) ............................................................................. Fabaceae 

8a. Plants climbing by tendrils .......................................................................................... Bignoniaceae 

8b. Plants scandent, twining, or with the leaf rachis twining, without tendrils ................................... 9 

9a. Plants scandent; flowers minute, yellow, aggregated on a common receptacle (head or 

capitulum)..................................................................................................................... Asteraceae 

9b. Plants twining or with the leaf rachis twining ............................................................................... 10


10a. Leaf rachis not twining; corolla gamopetalous............................................................................. 11 

10b. Leaf rachis twining; corolla of free petals ............................................................. Ranunculaceae 

11a. Plant woody, 5-10 m in length, not fetid; corolla hypocrateriform, 1.5-2.5 cm long...... Oleaceae 

11b. Plant herbaceous, 1.5-2 m in length, fetid; corolla tubular, ca. 1.5 mm long ........ Valerianaceae 

Key 3: LEAVES SIMPLE OR UNIFOLIOLATE 

1a. Leaves opposite .................................................................................................................................. 2 

1b. Leaves alternate ............................................................................................................................... 23 

2a. Plants scandent or clambering. ......................................................................................................... 3 

2b. Plants twining or climbing by adventitious roots. ............................................................................ 8 

3a. Perianth differentiated into a calyx and corolla; corolla gamopetalous. ......................................... 4 

3b. Perianth not differentiated into a calyx and corolla, consisting of 5 tepals. ................................... 7 

4a. Stipules present (sometimes early deciduous, but leaving a scar); ovary inferior or superior ........ 5 

4b. Stipules absent; ovary superior .......................................................................................................... 6 

5a. Ovary inferior .................................................................................................................... Rubiaceae 

5b. Ovary superior ............................................................................................................. Buddlejaceae 

6a. Corolla infundibuliform, the stamens slightly or not at all exserted; fruit capsular, opening in two 

longitudinal halves to reveal 2-4 seeds ........................................................................ Acanthaceae 

6b. Corolla hypocrateriform, the stamens twice as long as the corolla, exserted; fruit drupaceous, with 

4 pyrenes ......................................................................................................................... Verbenaceae 

7a. Plant herbaceous or slightly woody, without spines; tepals in two series, free; fruit circumscissile, 

membranaceous, not glandular ................................................................................. Amaranthaceae 

7b. Plant woody, with axillary spines; tepals united, forming a cone; fruit an anthocarp, with stipitate 

glands on its outer surface ......................................................................................... Nyctaginaceae 

8a. Plants climbing by adventitious roots ............................................................................................... 9 

8b. Plants twining .................................................................................................................................. 10 

9a. Corolla white; plants with prominently elongate stipules............................................... Rubiaceae 

9b. Corolla pink; plants without stipules .......................................................................... Bignoniaceae 

10a. Plants with white or yellowish milky latex ................................................................................... 11 

10b. Plants without colored latex .......................................................................................................... 13 

11a. Latex of the plant yellowish; stamens numerous; fruit a multilocular capsule ................. Clusiaceae 

11b. Latex of the plant white; stamens 5; fruit a unilocular follicle .................................................... 12 

12a. Corona present; stamens fused around the ovary and united to the stigma to form a gynostegium 

........................................................................................................................... Asclepiadaceae 

12b. Corona absent; stamens free or concrescent around the style, without forming a 

gynostegium........................................................................................................... Apocynaceae

44 


13a. Flowers with tepals (not differentiated in a calyx and corolla); cross section of the stem with areas 

of phloem included in the xylem .............................................................................. Amaranthaceae 

13b. Flowers with a calyx and corolla; cross section of the stem with the phloem external to the xylem 

.................................................................................................................................................................................14 

14a. Corolla of free petals ...................................................................................................................... 15 

14.a Corolla gamopetalous ....................................................................................................................17 

15a. Calyx elongate, tubular; petals pink to red .............................................................. Combretaceae 

15b. Calyx short, not tubular; petals yellow, green, yellowish green, or pink .................................... 16 

16a. Plant with short lateral branches that intertwine like tendrils; flowers minute (< 5 mm wide); 

calyx crateriform, with minute lobes; petals green or yellowish green; fruit capsular, with numerous 

winged seeds .................................................................................................................. Celastraceae 

16b. Plant lacking branches that resemble tendrils; flowers medium-sized (≥ 8 mm wide); calyx reduced 

to a pair of glands per sepal; petals yellow or pink; fruit a samaroid schizocarp, with only one seed 

per mericarp ............................................................................................................... Malpighiaceae 

17a. Ovary inferior ................................................................................................................................. 18 

17b. Ovary superior ............................................................................................................................... 20 

18a. Flowers aggregated in heads or capitula (with a receptacle in common), the heads forming a 

compound paniculiform or corymbiform inflorescence .................................................. Asteraceae 

18b. Flowers in panicles or aggregated in the leaf axils, not clustered in heads ................................19 

19a. Stipules present; corolla < 8 mm long, actinomorphic, with 5 similar lobes ............... Rubiaceae 

19b. Stipules absent; corolla > 2 cm long, zygomorphic, bilabiate (4 lobes united and one separate) 

..................................................................................................................... Caprifoliaceae 

20a. Stipules intrapetiolar, glandular; flowers or inflorescences not showy; corolla cream-colored; 

stamens 5, concrescent around the stigma; fruits of two follicles, with numerous seeds crowned 

with long, silky hairs .................................................................................................... Apocynaceae 

20b. Stipules absent; flowers or inflorescences showy; corolla of various colors, but not cream-colored; 

stamens 2 or 4; fruits capsular, fleshy, or dry, not follicular; seeds without long, silky hairs ...... 21 

21a. Fruits capsular, opening in two longitudinal halves; placental tissue persistent, projecting like a 

spur .............................................................................................................Acanthaceae 

21b. Fruits fleshy, indehiscent; placental tissue inconspicuous ........................................................... 22 

22a. Stamens 4; fruits with 2 or 4 pyrenes, containing 2 seeds ......................................... Verbenaceae 

22b. Stamens 2; fruits without pyrenes, containing a single seed .......................................... Oleaceae 

23a. Plants climbing by adventitious roots ........................................................................................... 24 

23b. Plants twining, scandent, clambering, or climbing by tendrils ................................................... 25 

24a. Plant with abundant milky latex; inflorescence a pear-shaped syconium (resembling a fruit) 

................................................................................................................Moraceae 

24b. Plant without latex; inflorescence umbelliform, with large nectaries in the center of the inflorescence 

.......................................................................................................................... Marcgraviaceae


25a. Plants climbing by tendrils ............................................................................................................ 26 

25b. Plants twining or scandent or clambering .................................................................................... 30 

26a. Tendrils opposite the leaves ................................................................................................ Vitaceae 

26b. Tendrils axillary (or in axillary inflorescences) or lateral to the leaves ...................................... 27 

27a. Tendrils lateral to or subopposite the leaves, usually branched ............................. Cucurbitaceae 

27b. Tendrils axillary ............................................................................................................................. 28 

28a. Base of the petiole forming an ocrea around the stem; tendrils representing a modification of the 

distal portion of the inflorescence axis ....................................................................... Polygonaceae 

28b. Base of the petiole not covering the stem to form an ocrea; tendrils not forming part of the 

inflorescences ................................................................................................................................... 29 

29a. Tendrils spiral-shaped; flowers with a corona and an androgynophore ................ Passifloraceae 

29b. Tendrils spirally twisted; flowers with neither a corona nor an androgynophore, stamens and 

ovary sessile on the receptacle ....................................................................................... Rhamnaceae 

30a. Plants clambering or scandent ...................................................................................................... 31 

30b. Plants twining ................................................................................................................................ 37 

31a. Ovary inferior ................................................................................................................................. 32 

31a. Ovary superior................................................................................................................................ 33 

32a. Flowers aggregated in a head or capitulum, with a receptacle in common; fruits dry, crowned with 

a tuft of scales or plumose hairs ...................................................................................... Asteraceae 

32b. Flowers in corymbs; fruits fleshy, crowned by the calyx lobes ...................................... Ericaceae 

33a. Flowers apetalous or with tepals (not discernible as a calyx and corolla)................................... 34 

33b. Flowers with a calyx and corolla................................................................................................... 36 

34a. Branches and main stem with a pair of spines in the leaf axils; fruits orange when ripe .... Ulmaceae 

34b. Branches without spines; fruits violet or yellowish when ripe .................................................... 35 

35a. Perianth of 5 yellowish tepals, ovate-lanceolate, concave, with a keel along their dorsal portion; 

stigmas 2, elongate, recurved; fruits membranaceous, circumscissile, yellowish ... Amaranthaceae 

35b. Perianth of 4 white tepals, oblong-elliptical, without a dorsal keel; stigma sessile, capitate; fruits 

fleshy, indehiscent, dark purple ................................................................................ Phytolaccaceae 

36a. Corolla gamopetalous; stamens 5; ovary sessile on the floral receptacle; fruit an indehiscent 

berry...................................................................................................................... Boraginaceae 

36b. Corolla of free petals; stamens numerous; ovary borne on a gynophore; fruit a dehiscent follicle 

........................................................................................................................ Capparaceae 

37a. Corolla absent ................................................................................................................................ 38 

37b. Corolla present ............................................................................................................................... 39 

38a. Calyx well developed, elongate, tubular, the limb expanded or with a pendulous tail; flowers 

usually solitary ........................................................................................................ Aristolochiaceae 

38b. Calyx minute; flowers usually aggregated in an inflorescence with showy bracts ....... Euphorbiaceae

46 


39a. Corolla gamopetalous ....................................................................................................................40 

39b. Corolla of free petals...................................................................................................................... 43 

40b. Flowers clustered in heads or capitula .......................................................................... Asteraceae 

40b. Flowers in various kinds of inflorescences, but not in heads ....................................................... 41 

41a. Stamens dehiscent by terminal pores ............................................................................ Solanaceae 

41b. Stamens dehiscent by longitudinal sutures ................................................................................... 42 

42a. Fruits fleshy, indehiscent; stamens with short filaments or the anthers sessile, united to the corolla 

in its middle to upper portion ...................................................................................... Boraginaceae 

42b. Fruits dry, capsular; stamens with more or less long filaments, united to the base of the corolla 

......................................................................................................................... Convolvulaceae 

43a. Flowers actinomorphic; petals white, all similar ......................................................................... 44 

43b. Flowers zygomorphic; petals pink or red, unequal, modified into a standard, wings, and a keel. 

............................................................................................................................................... 46 

44a. Stamens numerous; fruit capsular ............................................................................... Dilleniaceae 

44b. Stamens 4-6; fruit indehiscent ...................................................................................................... 45 

45a. Flowers unisexual; stamens 4 or 6; fruits fleshy, without a persistent style ....... Menispermaceae 

45b. Flowers bisexual; stamens 5; fruits dry, crowned with 3 persistent styles ................. Basellaceae 

46a. Fruit a legume ................................................................................................................... Fabaceae 

46b. Fruit an indehiscent samara ...................................................................................... Polygalaceae 

1. Family ACANTHACEAE 


1a. Plants with axillary spines; leaves up to 2 cm long, coriaceous; corollas 1.2-1.5 cm long; stamen 2 

................................................................................................................................... 2. Oplonia 

1b. Plants without spines; leaves > 2.5 cm long, chartaceous to coriaceous; corollas > 2.3 cm long; 

stamens 4 ............................................................................................................................................ 2 

2a. Plants twining; stems cylindrical ................................................................................ 3. Thunbergia 

2b. Plants decumbent or clambering, not twining; stems quadrangular ............................ 1. Asystasia 

1. ASYSTASIA 

Perennial herbs, erect, decumbent, or clambering. Leaves opposite; blades simple, with numerous 

linear cystoliths on the upper surface, the secondary veins conspicuous; stipules absent. Flowers bisexual, 

zygomorphic, arranged in spikes or one-sided, terminal racemes, accompanied at the base by 2 small 

bracts and 2 bracteoles. Calyx of 5 linear or lanceolate sepals of equal size; corolla zygomorphic, pale 

violet, pink, white, or pale yellow, infundibuliform, with 5 rounded lobes; stamens 4, didynamous;


ovary superior, bicarpellate, bilocular, the stigmas bilobed or capitate. Capsules ellipsoid or claviform; 

seeds 2-4, lenticular. A genus of about 40 species, of paleotropical origin. 

1. Asystasia gangetica (L.) T. Anderson in 

Thwaites, Enum. Pl. Zeyl. 235. 1860. 

Fig. 13. A-E 

BASIONYM: Justicia gangetica L. 

Decumbent herb, creeping or clambering, 1- 

3 m in length. Stems quadrangular. Leaves 2.5- 

11 × 2.5-6.5(8) cm, ovate to lanceolate, 

chartaceous, pubescent on both surfaces, 

especially on the veins, the apex acuminate, the 

base obtuse, truncate, rounded, or cordiform, the 

margins crenulate; petioles 0.5-3(4.5) cm long. 

Flowers arranged in terminal spikes on axillary 

branches; bracts and bracteoles ovate, ciliate. 

Calyx green, 5-7 mm long, the sepals lanceolate; 

corolla light yellow or pale violet, 2.3-5 cm long, 

2. OPLONIA 

with glandular hairs on the outer surface; stamens 

and pistil inserted on the floral tube. Capsules 2- 

2.5 cm long, claviform, light brown when ripe, 

densely covered with glandular hairs; seeds light 

brown, with irregular margins. 

Phenology: Collected in flower and fruit 


Status: Exotic, ornamental, naturalized in 

Puerto Rico and the Virgin Islands, quite common. 

Distribution: On roadsides and in open places. 

Also on Vieques, St. Croix, St. John, St. Thomas, 

and Tortola. A species of Asiatic origin, cultivated 

throughout the tropics. 

Public forests: Ceiba, El, Yunque, Río Abajo, 

and Tortuguero. 

Erect or clambering shrubs, usually with opposite, axillary spines. Leaves opposite, small, with 

short petioles or sessile, the secondary veins inconspicuous; stipules absent. Flowers bisexual, 

zygomorphic, heterostylous, axillary, solitary or in short, condensed racemes. Calyx of 5 acicular sepals; 

corolla infundibuliform, zygomorphic, with 5 lobes, one of them wider than the others; stamens 2, 

inserted or exserted; ovary superior, bicarpellate, bilocular, the style filiform, the stigma bilobate. Fruit 

capsular, claviform; seeds 2 or 4, lenticular. A genus of 19 species, native to tropical America and 

Madagascar. 

1. Oplonia spinosa (Jacq.) Raf., Fl. Tellur. 4: 65. 

1838. 

Fig. 13. F-K 

BASIONYM: Justicia spinosa Jacq. 

SYNONYM: Anthacanthus spinosus (Jacq.) Nees 

Clambering shrub, 2-4 m in length. Stems 

slender, obtusely quadrangular, densely covered 

with minute, whitish hairs when young, with 

numerous lateral branches; spines axillary, 0.4- 

1.2 cm long, recurved or rarely straight, 

perpendicular to the stem. Leaves opposite, 

usually clustered at the nodes; blades 0.5-2.2 × 

0.4-2 cm, elliptical, obovate, or narrowly 

elliptical, rigid-coriaceous, glabrous, the apex 

obtuse or rounded, slightly sulcate, usually with 

a tuft of minute hairs, the base cuneate or 

attenuate, the margins revolute; upper surface 

shiny, usually with cystoliths; lower surface dull, 

pale green; petioles minute. Flowers solitary or 

clustered in condensed racemes; bracts minute, 

triangular. Calyx green, infundibuliform, ca. 3 

mm long, of 5 lanceolate sepals; corolla lilac or 

lavender, 1.2-1.5 cm long. Capsules 1.2 cm long, 

claviform, glabrous; seeds 4, brown, lenticular, 

3-4 mm long. 

Phenology: Flowering from December to July 

and fruiting from April to July. 


Distribution: In coastal thickets and in the 

zone of mogotes. Also on Vieques, Cayo Icacos, 

Buck Island, St. John, St. Croix, St. Thomas, 

Tortola, and Virgin Gorda; the Bahamas, Cuba, 

Hispaniola, and the Lesser Antilles. 

Public forests: Guajataca, Guánica, Maricao, 

Piñones, Río Abajo, and Susúa.

48 


3. THUNBERGIA 

Herbaceous or woody vines or less frequently erect shrubs. Leaves opposite; blades simple; stipules 

absent. Flowers bisexual, zygomorphic, axillary, usually solitary, with long pedicels and a pair of 

foliaceous bracts covering the lower portion of the corolla. Calyx short, cupuliform, truncate or dentate; 

corolla infundibuliform or hypocrateriform, large and showy, regular or irregular; stamens didynamous;


ovary superior, bicarpellate, bilocular. Fruits capsular, globose at the base, with the apex in the form 

of a beak; seeds subglobose. Approximately 200 species of the Paleotropics, 12 of which have been 

cultivated and have become naturalized in the Neotropics. 

Key to the species of Thunbergia 

1a. Petioles winged; corolla orange or pale yellow, with the center dark violet .................... 1. T. alata 

1b. Petioles not winged; corolla white or violet, with the center white or light yellow ................2 

2a. Herbaceous vine, 2-3 m in length; corolla white, with the limb 4-5 cm in diameter... 2. T. fragrans 

2b. Woody vine, 10-15 m in length; corolla lilac or whitish, with the limb 6-7 cm in diameter ................ 

.............................................................................................. 3. T. grandiflora 

1. Thunbergia alata Bojer ex Sims, Bot. Mag. 

52, t. 2591. 1825. 

Fig. 14. G-L 

Susana, Viuda, Black-eye susan, Clock vine 

Herbaceous vine, creeping or climbing, 

twining, 2-3 m in length. Stems cylindrical, 

slender (ca. 2 mm), puberulous. Leaves opposite; 

blades 4.5-10.5 × 3.2-6 cm, ovate, lobed, 

chartaceous, the apex acute, the base 

subcordiform; upper surface dark green, dull, 

pubescent; lower surface pale green, dull, with 

prominent venation; petioles 4-8 cm long, winged, 

pubescent. Flowers axillary, solitary; pedicels 

pubescent, 4-5 cm long; bracts green, ovate, 

pubescent, 1.5 cm long, covering the calyx and 

the corolla tube. Calyx yellowish green, with 12 

filiform lobes, ca. 4 mm long; corolla orange, pale 

yellow, or less frequently whitish, 

infundibuliform, with 5 lobes, the tube ca. 2.5 cm 

long, narrow at the base, dark violet inside, the 

lobes ca. 2.5 cm long with the apex truncate, the 

limb ca. 5 cm in diameter; stamens with glandular 

hairs on the basal portion. Capsules ca. 4 mm 

long, depressed-globose to 4-lobed at the base, 

the upper half in the form of a beak, dehiscent by 

two valves; seeds 2 or 4, 1.2-1.5 mm long, 

semicircular, reticulate. 

Phenology: Flowering sporadically throughout 

the year. 

Status: Exotic, very common, naturalized in 

Puerto Rico. 

Distribution: Throughout Puerto Rico, 

especially in moist disturbed areas, at lower to 

upper elevations. Also on St. Croix. Native to 

eastern Africa, but introduced throughout the 

tropics. 

Public forests: Cambalache, Carite, El 

Yunque, Guilarte, Maricao, Río Abajo, Toro 

Negro, and Tortuguero. 

2. Thunbergia fragrans Roxb., Pl. Coromandel. 

1: 47. 1796. 

Fig. 14. A-F 

SYNONYM: Thunbergia volubilis Pers. 

Susana blanca, Flor de nieve 

Herbaceous vine, twining, 2-3 m in length. 

Stems cylindrical, striate, slender, puberulous. 

Leaves opposite; blades 6.5-11 × 1.8-6 cm, ovate 

to lanceolate, chartaceous, the apex acute, the base 

truncate or subcordiform; margins undulate and 

ciliate; upper surface dark green, glabrous or 

somewhat scabrous; lower surface pale green, dull, 

puberulous, with prominent venation; petiole 2- 

3.5 cm long, slender, pubescent, sulcate, with the 

base somewhat dilated. Flowers axillary, solitary 

or in pairs; pedicels pubescent, 5-7 cm long, 

striate; bracts green, membranaceous, ovate, 

pubescent, 1.6-2 cm long, covering the calyx. 

Calyx green, of 15-20 sepals, lanceolate, 3-5 mm 

long; corolla white, infundibuliform, with 5 lobes, 

the tube 2.5-4 cm long, narrow at the base, yellow 

inside, the limb 4-5 cm in diameter. Capsules 1- 

2.5 cm long, depressed-globose at the base, the 

upper half in the form of a beak, dehiscent in two 

halves; seeds 4, globose, ca. 5 mm in diameter, 

pubescent, with a depression at the base. 

Phenology: Flowering and fruiting throughout 

the year.

50 



Status: Exotic, common, naturalized in Puerto 

Rico and the Virgin Islands. 

Distribution: In areas of disturbed vegetation 

throughout Puerto Rico. Also on Vieques, St. 

Croix, St. John, St. Thomas, and Tortola. Native 

to India and Sri Lanka, but naturalized in the 

tropics. 

Public forests: Maricao, Piñones, Río Abajo, 


3. Thunbergia grandiflora (Roxb. ex Rottler) 

Roxb., Hort. Geng. 45. 1814. 

Fig. 15. A-F 

BASIONYM: Flemingia grandiflora Roxb. ex Rottler 

Tunbergia 

Woody vine, twining, 10-15 m in length. 

Stems cylindrical, up to 2.5 cm in diameter, 

striate, puberulous; cross section of the stem with 

the pith hollow and the xylem tissue with wide 

rays. Leaves opposite; blades 15-26 × 13-30 cm, 

ovate or broadly ovate, chartaceous, the apex acute 

or acuminate, the base cordiform or subcordiform, 

the margins lobate-dentate, ciliate; upper surface 

dark green, shiny, puberulous, with slightly 

prominent venation; lower surface light green, 

dull, glabrous or puberulous, with prominent 

venation; petioles 6-12 cm long, sulcate. Flowers 

arranged in axillary cymes; pedicels robust, 

cylindrical, 4-6 cm long; bracts light green, 

membranaceous, ovate, ca. 4 cm long, covering 

the calyx and the corolla tube. Calyx green, in 

the form of a ring, 4-5 mm long; corolla lilac or 

white, infundibuliform, with 5 lobes, the tube 6- 

7 cm long, light yellow inside, narrow at the base, 

the limb 6-7 cm in diameter. Capsule ca. 3 cm 

long, subglobose at the base, the upper half in the 

form of a beak, dehiscent in two halves. 

Phenology: Flowering throughout the year and 

fruiting sporadically from May to July. 

Status: Exotic, very aggressive, widely 

cultivated and naturalized in Puerto Rico. 

Distribution: In moist disturbed areas at 

moderately low to middle elevations. Native to 

India, but widely cultivated in the tropics. 

Cultivated on St. Croix and St. Thomas. 

Public forests: Maricao and Río Abajo. 

References. Stearn, W. T. 1971. A survey of the tropical genera Oplonia and Psilanthele 

(Acanthaceae). Bull. Br. Mus. Nat. Hist. (Bot.) 4: 261-323. 

2. Family AMARANTHACEAE 


1a. Leaves alternate; tepals glabrous....................................................................................................... 2 

1b. Leaves opposite; tepals densely covered with tufts of long hairs on the outer surface ................... 3 

2a. Clambering herbs, 0.5-1 m in length; utricle with numerous naked seeds .................... 1. Celosia 

2b. Scandent lianas or shrubs, 5-10 m in length; utricle with only one seed, covered by a fleshy arillode 

............................................................................................................................. 2. Chamissoa 

3a. Clambering herbs, 1.5-2 m in length; stigma divided in 2 divergent filiform branches....3. Iresine 

3b. Scandent shrubs or twining lianas, > 5 m in length; stigmas bilobate.............................. 4. Pfaffia 

1. CELOSIA 

Herbs or shrubs, erect or sometimes clambering or scandent. Leaves alternate, petiolate; blades 

simple; stipules absent. Flowers bisexual, pedicellate or sessile, clustered in cymes, which are arranged 

along terminal or axillary spikes, panicles, or thyrses. Tepals 5, subequal, free; stamens 5, the filaments 

united at the base to form a short crateriform tube; interstaminal appendages dentate or absent; ovary

52 



superior, unilocular, subglobose, ovoid or cylindrical, the style elongate or the stigmas elongate or 

capitate; ovules 2 or more numerous. Fruit a membranaceous, circumscissile utricle; seeds 2 to many, 

ellipsoidal, naked. About 50 species, widely distributed in the tropics and subtropics. 

1. Celosia nitida Vahl, Symb. Bot. 2: 44. 1791. 

Fig. 16. A-E 

Herb or subshrub, erect or sometimes 

clambering, 0.5-1 m in length, with numerous 

branches that are borne on a woody base. Branches 

cylindrical, glabrous, striate. Leaves alternate; 

blades 2-6 × 1-3 (4) cm, deltate-lanceolate or 

lanceolate, chartaceous, glabrous, the apex acute, 

the base unequal, truncate, abruptly narrowed 

toward the petiole, the margins entire or slightly 

undulate; upper surface dull, with numerous 

whitish dots; petioles slender, 1-1.5 cm long. 

Flowers arranged in terminal, simple or branched 

spikes, 2-10 cm long; bracts and bracteoles 

triangular, with a hyaline margin. Tepals whitish 

with the apex somewhat reddish, oblonglanceolate, 

ca. 4 mm long, glabrous; stamens 5, 

2. CHAMISSOA 

the filaments compressed, united at the base to 

form a short staminal tube; ovary ovoid, the 

stigmas 3, ca. 2 mm long. Capsule subglobose, 

somewhat compressed, ca. 3 mm long; seeds 9- 

19, dark brown, shiny, ca. 1.2 mm long, lenticular, 

naked. 




Distribution: In understory and thickets in 

sandy coastal areas and in disturbed areas in the 

interior at middle elevations. Also on Caja de 

Muerto, Cayo Ratones, Culebra, Mona, Vieques, 

St. Croix, St. John, St. Thomas, Tortola, and 

Virgin Gorda; in the southern United States 

(Florida), Mexico, the Antilles, and northern 

South America. 

Public forests: Guánica, Mona, Piñones, and 

Río Abajo. 

Subshrubs, scandent shrubs, or lianas. Leaves alternate, petiolate; blades simple; stipules absent. 

Flowers bisexual or functionally unisexual; clustered in cymes, which are arranged in terminal or 

axillary panicles. Tepals 5, ovate or lanceolate, free; stamens 5, the filaments united at the base to form 

a short staminal tube; interstaminal appendages absent; ovary superior, unilocular, uniovulate, the 

style short, with 2 or 3 stigmas. Fruit a membranaceous, circumscissile utricle; seed solitary, sometimes 

covered by a fleshy arillode. A genus of 2 species, widely distributed in the Neotropics, from Mexico to 

northern Argentina, including the Antilles. 

1. Chamissoa altissima (Jacq.) Kunth in Humb. 

Bonpl. & Kunth, Nov. Gen. Sp. (quarto ed.) 

2: 197. 1817. 

Fig. 1. D, 16. F-K 

BASIONYM: Achyranthes altissima Jacq. 

Liana or woody shrub, clambering, 3-15 m in 

length, with numerous pendulous branches. Stems 

angular when young and cylindrical when mature, 

up to 3 cm in diameter, glabrous; cross section of 

the stem with bands of vascular tissue alternating 

with very narrow bands of parenchymatous tissue. 

Leaves alternate; blades 8-14 × 3-7 cm, elliptical, 

ovate, or elliptic-ovate, chartaceous, the apex 

acute or acuminate, the base rounded, the margins 

undulate; upper surface slightly shiny, puberulous; 

lower surface pale green, dull, with prominent 

venation; petioles slender, usually reddish, 2-4.5 

cm long. Flowers in terminal panicles, 4-16 cm 

long. Perianth white to greenish, with five ovate 

tepals, concave, acute, ca. 3 mm long, glabrous; 

stamens 5; stigma bifid. Capsule ovoid, truncate 

at the apex, membranaceous, turning from green 

to white, 3-4 mm long; seeds dark brown, shiny, 

completely covered by a white arillode. 



Status: Native, very common.

54 



Distribution: In moist secondary forests, at 

middle and lower elevations. Also on Vieques, St. 

Croix, St. Thomas, and Tortola; in the Antilles 

and continental tropical America. 

3. IRESINE 

Public forests: Cambalache, El Yunque, 

Guilarte, Maricao, Río Abajo, and Toro Negro. 

Erect or clambering herbs, shrubs, and less frequently small trees. Leaves opposite or subopposite, 

petiolate; blades simple; stipules absent. Flowers unisexual or bisexual, pedicellate, clustered in cymes 

along axillary or terminal panicles; bracts and bracteoles persistent. Tepals 5, with a tuft of hairs at the 

base on the outer surface; stamens 5, the filaments united at the base to form a short staminal tube; 

interstaminal appendages present or absent; ovary superior, uniovulate, rounded, the stigmas divided 

in 2 elongate, filiform branches, sessile or subsessile. Fruit a membranaceous, subglobose, circumscissile 

utricle. Seed solitary, globose or lenticular, shiny, naked. A genus of approximately 40 species of 

neotropical distribution. 

Key to the species of Iresine 

1a. Bracts and bracteoles with a dorsal keel, aristate at the apex; bracteoles > 1.5 mm long, of the same 

size as the tepals; leaves lanceolate ........................................................................ 1. I. angustifolia 

1b. Bracts and bracteoles not keeled or aristate; bracteoles < 1 mm long, smaller than the tepals; leaves 

lanceolate or ovate ............................................................................................................ 2. I. diffusa 

1. Iresine angustifolia Euphrasén, Beskr. Ste. 

Barthél. 165.1795. 

Fig. 17. A-F 

SYNONYMS: Iresine elatior Rich. ex Willd. 

Iresine celosioides Sw. 

Herb, clambering or less frequently creeping, 

or sometimes a subshrub attainig 2 m in length, 

much branched from a woody base. Stems 

cylindrical, glabrous, with the nodes somewhat 

swollen. Leaves opposite; blades 2-13 × 0.5-4.5 

cm, lanceolate, chartaceous, the apex short or 

long-acuminate, the base obtuse or acute, usually 

unequal, the margins entire, slightly revolute; 

upper surface glabrous or puberulous; lower 

surface dull, with prominent venation; petioles 

slender, 0.5-2 cm long. Flowers bisexual, in 

terminal panicles, up to 30 cm long; bracts and 

bracteoles lanceolate, with a dorsal keel and the 

apex aristate, the bracteoles of the same size as 

the tepals. Tepals 5, white to greenish, elliptical, 

ca. 1.5 mm long, with a tuft of undulate hairs at 

the base; stamens 5; stigma bifid. Capsule 

subglobose, membranaceous, ca. 1 mm long; seeds 

black, shiny, subglobose, 0.8-1 mm long. 




Distribution: In disturbed areas at lower 

elevations. Also on Cayo Santiago, Culebra, 

Desecheo, Vieques, St. Croix, St. John, St. 

Thomas, and Tortola; from Mexico to Ecuador, 

including the Antilles. 

Public forests: Guánica and Río Abajo. 

2. Iresine diffusa Humb. & Bonpl. ex Willd., Sp. 

Pl. 4: 765. 1805. 

Fig. 17. J-L 

SYNONYMS: Iresine paniculata (L.) Kuntze 

Iresine celosioides L. 

Iresine celosia L. 

Herb or subshrub, clambering or less 

frequently creeping, 1-3 m in length, much 

branched from the woody base. Stems glabrous 

or pubescent, with the nodes compressed, usually 

pilose. Leaves opposite; blades 3-9 (-11) × 1-5 

cm, lanceolate or ovate, chartaceous, the apex 

acute or obtuse, the base obtuse, acute, or truncate, 

the margins slightly undulate and revolute; upper 

surface dull, puberulous or glabrous; lower surface 

dull, pubescent or glabrous, with prominent 

venation; petioles slender, 0.5-3.0 cm long.

56 


Flowers unisexual, in terminal or axillary 

panicles, pendulous, 7-15 cm long, the axes 

pubescent or glabrous; bracts and bracteoles not 

keeled, acute or short-acuminate at the apex, the 

bracteoles smaller than the tepals. Tepals creamcolored 

or whitish; ovate or oblong, with a tuft of 

undulate hairs at the base, 0.5-0.8 mm long in 

the staminate flowers, oblong, 0.9-1.1 mm long 

in the pistillate flowers; stamens 5, the filaments 

4. PFAFFIA 

unequal. Capsule ellipsoid, membranaceous, ca. 

1 mm long; seeds reddish brown, shiny. 

Phenology: Flowering and fruiting from 

September to May. 


Distribution: In disturbed areas at lower or 

middle elevations. Also on Desecheo and Vieques; 

the Lesser Antilles, Colombia, and Venezuela. 

Public forests: Ceiba, El Yunque, Maricao, Río 

Abajo, Toro Negro, Tortuguero, and Vega. 

Erect or clambering herbs, scandent shrubs, or twining lianas. Leaves opposite, petiolate; blades 

simple, stipules absent. Flowers bisexual, arranged in glomerules, spikes, or axillary or terminal panicles 

on short axillary branches; bracts and bracteoles persistent. Tepals 5, free, concave, with a tuft of hairs 

at the base on the outer surface or the hairs borne on the pedicel; stamens 5, the filaments united at the 

base to form a short staminal tube; interstaminal appendages present or absent; ovary superior, unilocular, 

uniovulate, ovoid, the stigma bilobed, sessile or subsessile. Fruit an indehiscent utricle, covered by the 

perianth. A genus of approximately 35 species of neotropical distribution. 

1. Pfaffia aurata (Mart.) Borsch, Novon 5: 230. 

1995. 

Fig. 17. G-I 

BASIONYM: Trommsdorffia aurata Mart. 

SYNONYMS: Trommsdorffia argentata Mart. 

Iresine argentata (Mart.) D. Dietr. 

Achyranthes nodosa Bertero ex Mart. (pro syn.) 

Achyranthes argentata (Mart.) Moq. 

Iresine argentata var. latifolia Suess. 

Pfaffia grandiflora sensu Alain, non (Hook.) R. E. Fr. 

Twining liana, 15-20 m in length, with 

numerous pendulous branches. Stems cylindrical, 

with swollen nodes, densely silvery- pubescent 

when young, glabrous and lenticellate when 

mature, up to 10 cm in diameter; bark fissured, 

brown-cream; cross section of the stem with bands 

of vascular tissue alternating with very narrow 

bands of parenchymatous tissue. Leaves opposite; 

blades 3-10 × 2.7-4.2 cm, elliptical, ovate or 

lanceolate, chartaceous to coriaceous, the apex 

acuminate, acute, obtuse, or retuse, the base obtuse 

or rounded, sometimes unequal, the margins 

entire, slightly undulate and revolute; upper 

surface dull, glabrous or puberulous; lower surface 

dull, glabrous or puberulous, with slightly 

prominent venation; petioles slender, puberulous, 

3-10 mm long. Flowers in panicles at the tips of 

short axillary branches, ascending, up to 30 cm 

long, the axes strigose or tomentose; bracts and 

bracteoles widely ovate to deltoid, hyaline, villous 

when young, 0.5-0.7 mm long. Tepals 5, creamcolored, 

oblong, 1.7-2.2 mm long, with a tuft of 

white, erect hairs at the base, these of the same 

length as or slightly longer than the tepals; 

stamens 5, the filaments alternating with short 

obtuse or rounded appendages, ca. 0.1 mm long; 

stigma bilobed, turbinate, papillose. 

Phenology: Collected in flower in January. 

Status: Native, locally common. 

Distribution: In moist areas along the 

Cordillera Central, along rivers and at the base 

of mogotes in the zone where they occur. Also in 

the Lesser Antilles, Panama, Colombia, 

Venezuela, Ecuador, Peru, and Brazil. 


Commentary: This species was originally 

described for Puerto Rico as Trommsdorffia 

argentata by Martius in 1826 and then transferred 

to Iresine argentata. Studies by T. Borsch in 1995 

consider T. argentata as belonging within the 

variation observed in Pfaffia aurata sensu lato 

and therefore as a synonym of the latter species.


58 


References: Borsch, T. 1995. Three new combinations in Pfaffia (Amaranthaceae) from the New 

World tropics. Novon 5: 230-233. Eliasson, U. H. 1987. Amaranthaceae. In: G. Harling and L. Anderson 

(eds.) Flora of Ecuador 28: 1-138. Sohner, S. H. 1977. Chamissoa (Amaranthaceae) en la República 

Dominicana. Moscosoa 1: 1-8. 

3. Family APOCYNACEAE 


1a. Upper surface of the leaves with numerous cystolithic hairs that give it a scabrous texture; 

inflorescence opposite the leaf ....................................................................................... 2. Anechites 

1b. Upper surface of the leaves glabrous; inflorescences axillary, terminal or pseudo-terminal.......... 2 

2a. Plants with watery latex; corolla cream-colored or greenish ........................................... 3. Echites 

2b. Plants with abundant milky (white) latex; corolla yellow, white, red, or purple. ........................... 3 

3a. Corolla < 1 cm long, cardinal red, the lobes oblong, long (as long as or longer than the tube of the 

corolla) ......................................................................................................................... 4. Forsteronia 

3b. Corolla > 2.5 cm long, white, yellow, or purple, the rounded lobes short (much shorter than the 

tube of the corolla). ............................................................................................................................ 4 

4a. Corolla white, with a yellow center ......................................................................... 6. Rhabdadenia 

4b. Corolla yellow or purple. ................................................................................................................... 5 

5a. Leaves opposite; stems copper-colored; corolla tube 3-6 cm long ............................. 5. Pentalinon 

5b. Leaves whorled; stems grayish; corolla tube 7-9 cm long .......................................... 1. Allamanda 

1. ALLAMANDA 

Lianas, shrubs, or trees, producing abundant milky latex. Leaves opposite or in whorls of 3-4 (5); 

blades simple; petioles more or less elongate, with intrapetiolar glands. Flowers actinomorphic, bisexual, 

produced on short axillary or pseudo-terminal cymes; bracts minute, deciduous. Calyx of 5 foliaceous 

sepals; corolla infundibuliform, showy, large, the lobes 5, rounded, contorted in the floral bud; stamens 

inserted, the anthers free; ovary superior, with an annular nectary at the base, of two unilocular carpels, 

with numerous ovules, the stigma bicapitate. Fruits capsular, globose or subglobose, with numerous 

spiny projections; seeds numerous, compressed, winged. A genus of approximately 18 species distributed 

throughout tropical South America, some of them of wide distribution. 

1. Allamanda cathartica L., Mant. 214. 1771. 

Fig. 18. A-D 

Canaria, Cantiva, Allamanda 

Shrub or woody vine, clambering or 

sometimes twining, much branched, 2-8 m in 

length, with abundant milky latex. Stems grayish, 

cylindrical, glabrous or puberulous. Leaves in 

whorls of 3 or 4; blades 8-13 × 1.5-3.5 cm, oblong, 

elliptical or oblanceolate, coriaceous, the apex 

acuminate, the base acute, the margins undulate 

and revolute; upper surface glabrous, dark green, 

shiny, with a prominent midvein; lower surface


yellowish green, with the midvein thickened, 

prominent, and puberulous; petioles 5-10 mm 

long; stipules transformed into 4 small 

intrapetiolar glands. Flowers in axillary cymes, 

few-flowered. Calyx greenish, of 5 lanceolate 

sepals, 12-18 mm long; corolla infundibuliform, 

yellow, the tube 7-9 cm long, the limb ca. 8 cm in 

diameter, with five rounded, revolute lobes. 

Capsules ellipsoid, with numerous spines on the 

outside, infrequent; seeds numerous, oval, 

compressed, 1.2-1.5 cm long, with a discolorous, 

wing-like margin. 

2. ANECHITES 

A monotypic genus, characterized by the following species. 

1. Anechites nerium (Aubl.) Urb., Repert. Sp. 

Nov. Regni Veg. 16: 150. 1919. 

Fig. 18. E-H 

BASIONYM: Apocynum nerium Aubl. 

SYNONYMS: Echites lappulacea Lam. 

Anechites asperuginis Griseb. 

Twining vine, woody or herbaceous, attaining 

10 m in length. Stems cylindrical, sparsely 

strigulose, glabrous and corky lines when mature. 

Leaves opposite or, when solitary, opposite the 

inflorescence; blades 5.5-10 × 2.5-4.5 cm, 

elliptical, oblong or ovate, chartaceous, the apex 

acuminate, the base obtuse, truncate, or 

subcordiform, the margins entire or undulate; 

upper surface dull, scabrous, covered with rigid 

cystolithic hairs; lower surface with small 

scattered protuberances and with the venation 

puberulous; petioles 15-22 mm long; stipules 

transformed into small intrapetiolar glands. 

Flowers bisexual, actinomorphic, in simple or 

bifurcate racemes, opposite a leaf, 6-15 (25) cm 

long, the axis sparsely strigulose. Calyx of 5 

lanceolate sepals, 2.5-3 mm long; corolla 

hypocrateriform, the tube ellipsoid, ca. 6 mm long, 

3. ECHITES 


fruiting from April to May. 

Status: Exotic, naturalized in Puerto Rico. 

Distribution: Widely cultivated in Puerto Rico, 

Vieques, and the Virgin Islands. Native to South 

America, but found throughout the tropics due to 

its cultivation as an ornamental. 

Public forests: Carite, El Yunque, Piñones, Río 

Abajo, and Tortuguero. 

Cultivated species: Allamanda blanchetii DC., 

with a purple corolla, is often cultivated in Puerto 

Rico and the Virgin Islands, but apparently it has 

not become naturalized. 

yellowish, pilose inside, the lobes 5, white, 8-12 

mm, oblong, rounded at the apex; stamens 5, the 

filaments short, connate with the corolla, the 

anthers apiculate; ovary superior, bicarpellate, the 

style elongate, the stigmas 2, elongate. Fruit of 

(1) 2 fusiform follicles, densely strigose, especially 

in the distal portion; seeds (according to 

Grisebach, 1862) numerous, with a tuft of hairs 

at the apex. 

Phenology: Flowering from May to July and 

fruiting from June to July, according to the 

material examined (from Puerto Rico, Cuba, and 

Colombia). 

Status: Native, extremely rare. 

Distribution: Known from Puerto Rico from 

a single collection made by Dr. Agustín Stahl in 

1888 in the area of Vega Baja. A surviving 

duplicate of this collection (Stahl 870) is deposited 

in the United States National Herbarium of the 

Smithsonian Institution. Recently recollected by 

C. Trejo (2025) in Mata de Plátano reserve in Bo. 

Dominguito, Arecibo. In addition, this species has 

been collected in Cuba, Hispaniola, Jamaica, 

Panama, Colombia, and Ecuador. 

Twining vines or lianas, with abundant milky or watery latex. Leaves opposite; blades simple; 

petioles with or without glands; stipules glandular, intrapetiolar and interpetiolar. Flowers bisexual, in 

axillary cymes, elongate or compact. Calyx campanulate, of 5 elongate sepals, each with a glandular 

appendage at the base on the inner surface; corolla hypocrateriform, not ringed in the throat; stamens

60 



5, inserted, the anthers concrescent around the 

stigma, the filaments united to the corolla; ovary 

superior, with 5 glandular nectarines at the base; 

carpels 2, free at the base, united at the apex by a 

style in common. Fruits of 2 cylindrical follicles; 

seeds numerous, with a tuft of long hairs at the 

apex. A neotropical genus of approximately 35 

species. 

1. Echites agglutinata Jacq., Enum. Syst. Pl. 13. 

1760. 

Fig. 20. A-F 

SYNONYM: Prestonia agglutinata (Jacq.) Woodson 

Babeiro 

Herbaceous vine, twining, 1-5 m in length, 

with watery latex. Stems green, cylindrical, 

slender, lenticellate, becoming hollow on drying. 

Leaves opposite; blades 4-19 × 3-9 cm, oval, ovate, 

rounded, or sometimes lanceolate, glabrous, 

chartaceous, the apex obtuse, acute, shortacuminate, 

or subrounded and mucronate, the base 

obtuse, rounded, or truncate, lacking glands and 

4. FORSTERONIA 

usually asymmetrical, the margins entire or 

undulate, revolute; upper surface dull green; lower 

surface pale green, dull, with a prominent 

midvein; petioles 0.5-4 cm long, canaliculate, 

pubescent; stipules intrapetiolar and interpetiolar, 

minute, acicular. Flowers in small axillary or 

terminal cymes. Calyx green, 2-3 mm long, with 

5 ovate lobes, ca. 2 mm long; corolla 

infundibuliform, the tube greenish or creamcolored, 

4-6 mm long, with 4 ribs, pilose inside, 

the limb pale yellow or cream-colored, with 5 

oblong lobes, reflexed, 2-4 mm long. Follicles 

linear, 10-25 cm long, divaricate when mature; 

seeds numerous, 5-7 mm long, light brown, with 

cream-colored, silky hairs. 

Phenology: Flowering from May to December 

and fruiting from June to March. 


Distribution: Abundant in thickets and coastal 

forests. Also on Desecheo, Mona, Vieques, 

Anegada, St. Croix, St. John, St. Thomas, and 

Tortola; Hispaniola. 

Public forests: Guánica, Maricao, Mona, 

Piñones, Río Abajo, and Susúa. 

Twining vines or rarely subshrubs, with abundant milky latex. Leaves opposite or whorled in groups 

of 3 or 4; petiolate; blade simple, usually with domatia in the axils of the secondary veins, the midvein 

usually with glands at the base; stipules minute, intrapetiolar or interpetiolar. Flowers bisexual, 

actinomorphic, in axillary or terminal corymbiform or paniculiform cymes. Calyx of 5 sepals; corolla 

hypocrateriform or campanulate, with 5 lobes; stamens 5, inserted or exserted, the anthers concrescent 

around the stigma, with or without apical appendages; ovary superior, with 5 glandular nectarines at 

the base; carpels 2, free or rarely united, the stigma fusiform or subcapitate. Fruits of 2 slender follicles; 

seeds numerous, truncate, with a tuft of long hairs at the apex. A neotropical genus of about 50 species. 

1. Forsteronia portoricensis Woodson, Ann. 

Missouri Bot. Gard. 21: 618. 1934. 

Fig. 19. A-F 

SYNONYM: Forsteronia corymbosa sensu Britton & P. Wilson 

Bejuco de San Juan, Sanjuanera, 

Leche de perra 

Woody vine, twining, much branched from the 

base, 7-15 m in length, with abundant creamcolored 

latex. Stems dark brown, glabrous, 

cylindrical, lenticellate. Leaves opposite; blades 

4-8 × 1.8-3.8 cm, elliptical or broadly elliptical, 

coriaceous, glabrous, the apex acute, the base 

obtuse or rounded, the margins entire, slightly 

undulate, revolute; upper surface dark green, 

shiny, with a slightly prominent midvein, without 

glands; lower surface yellowish green, shiny, with 

a prominent midvein, domatia absent; petioles 5- 

7 mm long, slightly winged or sulcate; stipules 

interpetiolar, minute, brown, acicular. Flowers in 

terminal corymbiform cymes, 3-8 cm wide; 

pedicels pink, 7-12 mm long, with a pair of minute 

bracteoles at the base. Calyx cardinal red, 

glabrous, the sepals ovate, free to the base, 1.5- 

1.7 mm long, ciliate; corolla cardinal red, 

infundibuliform, 5-6 mm long, the lobes oblong,

62 


as long as or longer than the tube; stamens 

exserted. Follicles straight, ca. 8 cm long; seeds 

numerous, with long, silky hairs. 

Phenology: Flowering from April to January 

and fruiting from January to June. 

Status: Endemic to Puerto Rico, quite 

common. 

5. PENTALINON 

Twining vines or scandent shrubs, with abundant milky latex. Leaves opposite, petiolate; blades 

simple, without glands, stipules glandular, intrapetiolar. Flowers in axillary cymes. Calyx campanulate, 

of 5 elongate sepals, with digitiform glandular hairs (colleters) at the base; corolla infundibuliform; 

stamens 5, inserted, the anthers concrescent around the stigma, with apical appendages, the filaments 

very short; ovary superior, with 5 glandular nectaries at the base; carpels 2, free or united by a common 

style. Fruits of 2 cylindrical follicles, connate at the apex; seeds numerous, narrowly elliptical, with a 

tuft of long hairs at the apex. A neotropical genus of two species. 

1. Pentalinon luteum (L.) Hansen & Wunderlin, 

Taxon 35: 167. 1986. 

Fig. 19. G-J 

BASIONYM: Vinca lutea L. 

SYNONYM: Urechites lutea (L.) Britton 

Babeiro amarillo, Wild allamanda 


base, up to 5 m in length, with abundant milky 

latex. Stems smooth, cylindrical, pubescent when 

young, lenticellate; bark thin, copper-colored. 

Leaves opposite; blades 3-7 × 1.5-3.7 cm, elliptical 

or obovate, coriaceous, glabrous, the apex rounded 

and mucronate, the base rounded to obtuse, the 

margins entire, revolute; upper surface shiny with 

the midvein yellowish; lower surface dull, with a 

prominent midvein, puberulous; petioles ca. 5 mm 

long, cylindrical, pubescent. Flowers in axillary 

cymes, few-flowered; pedicels cylindrical, 

6. RHABDADENIA 


upper elevations, along the Cordillera Central and 

the Sierra de Luquillo and in the zone of mogotes. 

Public forests: Carite, El Yunque, Guajataca, 

Guilarte, Maricao, Río Abajo, Susúa, and Vega. 

pubescent, 1.5-2 cm long. Calyx light green, 

pubescent, of five lanceolate sepals, ca. 12 mm 

long; corolla yellow, infundibuliform, ca. 3 cm 

long, with cardinal red lines inside, the limb 3 

cm in diameter, with 5 rounded lobes. Follicles 

woody, linear, divaricate; seeds numerous, with 

long, silky hairs. 




Distribution: Coastal forests and thickets. Also 

on Cayo Icacos, Culebra, Mona, Vieques, 

Anegada, Guana Island, St. Croix, St. John, St. 

Thomas, Tortola, and Virgin Gorda; the Antilles 

and the United States (Florida). 


Río Abajo. 

Twining lianas or less frequently erect shrubs, with abundant milky latex. Leaves opposite, petiolate; 

blades simple, without glands; stipules interpetiolar, deciduous or persistent. Flowers bisexual, 

actinomorphic, in small axillary cymes. Calyx of 5 free sepals, elongate, without appendages; corolla 

infundibuliform; stamens 5, inserted, the anthers concrescent around the stigma, the filaments united 

to the corolla; ovary superior with 5 glandular nectaries at the base; carpels 2, free. Fruits of 2 cylindrical 

follicles; seeds numerous, linear, with a tuft of long hairs at the apex. A neotropical genus of 4 or 5 

species.


64 



1. Rhabdadenia biflora (Jacq.) Müll. Arg. in 

Mart., Fl. Bras. 6(1): 175. 1860. 

Fig. 20. G-K 

BASIONYM: Echites biflora Jacq. 

SYNONYM: Rhabdadenia paludosa (Vahl) Miers 

Twining liana, glabrous, 7-10 m in length, 

with abundant milky latex. Stems green, 

cylindrical, smooth. Leaves opposite; blades 5-8 

× 1.5-4.5 cm, elliptical, obovate, oblong, 

oblanceolate, and sometimes ovate, glabrous, the 

apex mucronate, obtuse, truncate, or sometimes 

retuse, the base obtuse or attenuate, the margins 

entire, slightly revolute; upper surface dull green; 

lower surface glaucous, with obscure venation; 

petioles 0.5-2 cm long; stipules not visible. 

Flowers in short axillary cymes. Calyx of oblong, 

mucronate sepals, 5-5.5 mm long, green; corolla 

infundibuliform, white, sometimes tinged with 

pink, with the center yellow inside, 4.5-5.7 cm 

long. Follicles linear, 6-10 cm long, parallel or 

slightly divaricate when mature; seeds numerous, 

linear, ca. 2.5 cm long, with a tuft of silky hairs 



fruiting mostly from March to September. 


Distribution: Abundant in mangroves and 

marshy coastal forests. Cited for St. Thomas by 

Britton, but probably extirpated since most of its 

mangroves have been destroyed. Also in the 

Antilles, the United States (Florida), Mexico, 

Central America, and South America. 

Public forest: Piñones. 

References: Morales, J. F. 1997. A reevaluation of Echites and Prestonia sect. Coalitae 

(Apocynaceae). Brittonia 49: 328-336. Sakane, M. and G. J. Shepherd. 1986. Uma revisão do gênero 

Allamanda (Apocynaceae). Rev. Brazil. Bot. 9: 125-149. 

4. Family ARISTOLOCHIACEAE 

1. ARISTOLOCHIA 

Herbaceous or woody vines, less frequently herbs or shrubs. Leaves alternate, petiolate; blades 

simple, entire or trilobed; pseudostipules usually present. Flowers bisexual, zygomorphic, solitary and 

axillary or clustered and cauliflorous. Calyx inflated in the basal portion (utricle), the apex of the 

utricle narrow, forming a tube that expands toward the margins into an entire, circular or bilobed limb, 

the lobes generally unequal, some forming a long tail; corolla absent; stamens 5-6, sessile; ovary 

inferior or half-inferior, of 5-6 united carpels; ovules numerous; styles 5-6, connate, the stigma capitate. 

Fruit a septicidal capsule, with numerous light, compressed seeds. A tropical genus of 450-550 species. 

Key to the species of Aristolochia 

1a. Leaves trilobed ............................................................................................................. 7. A. trilobata 

1b. Leaves entire, not lobed ..................................................................................................................... 2 

2a. Pseudostipules absent......................................................................................................................... 3 

2a. Pseudostipules present ....................................................................................................................... 4 

3a. Leaves lanceolate, hastate, or less frequently ovate; floral limb 4-6 cm in diameter, without a tail 

...................................................................................................................... 5. A. odoratissima 

3b. Leaves ovate or broadly ovate; floral limb 12-31 cm in diameter with a long tail, 7-150 cm long 

......................................................................................................................... 3. A. grandiflora

66 


4a. Leaves oblong, oblong-lanceolate, lanceolate-deltoid or less frequently ovate; floral limb 2.5 cm 

wide or smaller ................................................................................................................................... 5 

4b. Leaves broadly ovate, reniform or orbicular; floral limb > 5 cm wide ............................................ 6 

5a. Leaves oblong or oblong-lanceolate, the apex obtuse to rounded; lower surface tomentulose, with 

prominently reticulate venation; limb sub-bilabiate, the lower lip not well developed, the upper lip 

laterally expanded .................................................................... 4. A. oblongata subsp. calceiformis 

5b. Leaves lanceolate-deltoid or ovate, the apex acute to obtuse; lower surface puberulous, the venation 

not prominent; floral limb unilabiate, the erect lip forming a right angle with the tube..1. A. anguicida 

6a. Limb of the flower bilabiate (the lips elongate); pseudostipules 2.5-5 cm long .......... 6. A. ringens 

6b. Limb of the flower peltate, crateriform, the lower margin retuse; pseudostipules 1-1.5 cm long .. 7 

7a. Limb of the flower concave; capsules straight, 5-6 cm long ........................................ 2. A. elegans 

7b. Limb of the flower flat to convex; capsules curved, 7-10 cm long ...................... 5. A. odoratissima 

1. Aristolochia anguicida Jacq., Enum. Syst. Pl. 

30. 1760. 

Fig. 21. A-D 

Slightly woody vine, twining, attaining 5 m 

in length. Branches puberulous, sulcate. Leaves 

alternate; blades 4.5-15 × 3.5-9 cm, lanceolatedeltoid 

or ovate, chartaceous, the apex acute to 

obtuse, the base cordiform or subtruncate, the 

margins entire; upper surface glabrous or 

puberulous; lower surface puberulous, the 

venation not prominent; petioles 2-6.5 cm long, 

puberulous; pseudostipules reniform or circular, 

1.5-3.5 cm long. Flowers cream-colored with 

cardinal red spots, solitary; peduncle ca. 4.5 cm 

long; utricle obovoid, 6-13 mm long, the tube 1-2 

cm long, expanding toward the limb into an 

infundibuliform structure, with one lobe or 

elongate lip erect on the upper margin, the limb 

up to 2.5 cm wide. Capsule cylindrical, 2.2-3.8 

cm long; seeds membranaceous, broadly ovate, 

5.5-6.8 mm long, with a surrounding wing. 



Status: Native to South America, apparently 

cultivated on St. Croix. 

Distribution: Known from St. Croix from a 

single collection; Central America, Colombia, 

Trinidad, and Martinique. 

2. Aristolochia elegans Mast, Gard. Chron. II, 

24: 301. 1885. 

Fig. 22. A-F 

SYNONYM: Aristolochia littoralis sensu Pfeifer, non Parodi 

Woody vine, twining, attaining 7 m in length. 

Stems glabrous, cylindrical. Leaves alternate; 

blades 7-9 × 6-10 cm, reniform or broadly ovate, 

chartaceous, the venation palmate, the apex obtuse 

or rounded, the base cordiform, the margins 

entire; upper surface glabrous, shiny; lower 

surface with prominent venation, glaucous; 

petioles 1-3 cm long, glabrous; pseudostipules 

reniform, 1-1.5 cm long. Flowers solitary, 

pendulous; peduncle ca. 5 cm long; utricle 

ellipsoid-cylindrical, ca. 3 cm long, the tube 2.5- 

3.5 cm long, curved at the base, ascending, the 

limb asymmetrically peltate, crateriform, the 

lower margin retuse, 5-7 cm in diameter, light 

green or cream-colored with a cardinal red 

reticulum. Capsule cylindrical with 5 longitudinal 

ribs, 5-6 cm long; seeds membranaceous, 

triangular or ovate, 6-7 mm long. 

Phenology: Collected in flower in February 

and March and in fruit in February. 

Status: Exotic, cultivated and naturalized, 

uncommon. 

Distribution: On the edge of dry secondary 

forests. Native of South America but cultivated 

throughout the Antilles. Naturalized on St. John 

and Tortola, cultivated on St. Croix. 

3. Aristolochia grandiflora Sw., Prodr. 126. 

1788. 

Fig. 23. A-C 

Woody vine, twining, glabrescent, 5-7 m in 

length. Stems cylindrical. Leaves alternate; blades


68 



70 


6-20 × 6-16 cm, ovate or broadly ovate, 

chartaceous, the venation semi-palmate (with 

three main veins), the apex acute or acuminate, 

the base cordiform-sagittate, the margins 

undulate; upper surface glabrous, dull; lower 

surface with prominent venation, glaucous, 

puberulent; petioles 5-11 cm long, glabrous; 

pseudostipules absent. Flowers solitary, 

pendulous; peduncle 12-23 cm long; utricle 

asymmetrically obovoid, 6-12 × 2-3.5 cm, the tube 

8-13 cm long, curved, broadened toward the apex; 

limb peltate, obovate to elliptical, 12-31 cm in 

diameter, cream-colored with a cardinal red 

reticulum, the apex cordate, the base with an 

appendage or tail, 7-150 cm long, the throat 

cardinal red. Capsule more or less cylindrical, 

with 5 longitudinal ribs, 13-15.5 cm long; seeds 

membranaceous, broadly ovate, 1-1.3 cm long. 

Phenology: Collected in flower in December, 

February, April, May, and August and in fruit in 

April. 

Status: Exotic, cultivated and escaped from 

cultivation, uncommon. 

Distribution: In areas of secondary vegetation. 

Also in Jamaica, Cuba, Mexico, Central America, 

Colombia, and Ecuador; cultivated throughout the 

tropics. 

4. Aristolochia oblongata Jacq. subsp. 

calceiformis (Urb.) R. Rankin & Acev.-Rodr., 

comb.nov. 

Fig. 21. E-J 

BASIONYM: Aristolochia calceiformis Urb., Symb. Antill. 1: 

300. 1899. 

SYNONYMS: Aristolochia bilabiata sensu Pfeifer, non L. 

Slightly woody vine, twining, 10-15 m in 

length. Stems subcylindrical, attaining 4 cm in 

diameter, the bark corky with numerous 

longitudinal furrows; cross section of the stem 

showing a xylem dissected by multicellular rays. 

Branches puberulous, sulcate. Leaves alternate; 

blades 5-13 × 2.4-3.6 cm, oblong, oblonglanceolate, 

or less frequently elliptical, 

chartaceous, venation reticulate, the apex obtuse 

or rounded, the base subcordiform or subtruncate, 

the margins entire, slightly revolute; upper surface 

glabrous, with some scattered resinous dots; lower 

surface with prominent venation, tomentulose; 

petioles 1-1.5 cm long, puberulous; pseudostipules 

reniform, 0.8-1 cm long. Flowers solitary or in 

axillary racemes; peduncle ca. 3 cm long; utricle 

ellipsoid, brown, ca. 1 cm long, geniculate at the 

base, the tube ca. 1.5 cm long, expanding toward 

the limb into an infundibuliform structure, subbilabiate, 

with two short lobes, of which the upper 

lobe is larger and folded, covering the throat after 

anthesis, the throat bright yellow, pubescent. 

Capsule ellipsoid or subglobose, 2-2.5 cm long; 

seeds membranaceous, triangular, ca. 5 mm long. 

Phenology: Collected in flower in February, 

March, and September and in fruit from March 

to September and in December. 


Distribution: In moist forests at lower and 

middle elevations in the Cordillera Central and 

in the northern limestone zone. Also in Cuba and 

Hispaniola. 


5. Aristolochia odoratissima L., Sp. Pl. ed. 2, 

1362. 1763. 

Fig. 24. A-B 

SYNONYM: Aristolochia pandurata Jacq. 


Stems cylindrical, glabrous, smooth. Leaves 

alternate; blades 6.5-11 × 3.5-6 cm, lanceolate, 

hastate, or less frequently ovate, chartaceous or 

coriaceous, the venation palmate, the apex 

acuminate, the base deeply cordate, the margins 

entire; upper surface dull, glabrous; lower surface 

puberulous, the venation not prominent; petioles 

1-3.5 cm long; pseudostipules inconspicuous or 

ovate, up to 1.5 cm long. Flowers solitary, 

yellowish with a cardinal red reticulum; peduncle 

ca. 3 cm long; utricle obovoid, ca. 1 × 0.5 cm; 

tube slightly curved, 0.7-1.7 cm long, almost 

forming a right angle with the utricle; limb 4-6 

cm in diameter. Capsule curved, 7-10 cm long, 

subcylindrical, with 5 longitudinal ribs, the apex 

and base acute; seeds numerous, membranaceous, 

triangular, ca. 3 mm long. 



Status: Apparently exotic, naturalized on St. 

John; uncommon. 

Distribution: In areas of secondary vegetation, 

in coastal thickets on St. John. Collected on St. 

Thomas but possibly from a cultivated individual; 

Mexico, Central America, and South America.


6. Aristolochia ringens Vahl, Symb. Bot. 3: 99. 

1794. 

Fig. 24. C-F 

SYNONYMS: Aristolochia grandiflora sensu Vahl, non Sw. 

Aristolochia galeata sensu authors, non Mart. & Zucc. 

Gallito, Panitos, Pelicanos 


in length. Stems cylindrical, slender, glabrous, 

with the pith hollow. Leaves alternate; blades 5- 

16 × 6-18 cm, broadly ovate, reniform or orbicular, 

chartaceous, with prominently reticulate venation, 

the apex obtuse or rounded, the base deeply 

cordate, the margins entire; upper surface dark 

green, dull; lower surface glaucous, glabrous, with 

numerous scattered dots; petioles 3-11 cm long, 

sulcate or compressed, broadened at the base; 

pseudostipules foliaceous, ovate-rounded, 2.5-5 

cm long. Flowers solitary, pendulous; peduncle 

7.5-17 cm long; utricle obovoid, 5-7 × 2.5-4 cm, 

the tube straight, 3-4 cm long, almost forming a 

right angle with the utricle; limb bilabiate, 

yellowish with a cardinal red reticulum, the upper 

lip spathulate, 6-9 cm long, the lower lip 

lanceolate, 10-15 cm long. Capsule 6-11 cm long, 

oblong or oblanceolate, with 6 ribs, the apex 

mucronate, the base acute; seeds numerous, 

rhomboid, winged, 7-15 mm long. 



Status: Exotic, uncommon, cultivated and 

escaped. 

Distribution: On roadsides and in open areas. 

Native to Brazil; widely cultivated in the 

Neotropics. 

7. Aristolochia trilobata L., Sp. Pl. 960. 1753. 

Fig. 22. G-H 

Bejuco de Santiago, Cachimbo, Panitos, 

Tobacco pipe 

Slightly woody vine, twining, 3-5 m in length. 

Stems slender, shiny, cylindrical, glabrous, pink 

on the younger portions. Leaves alternate; blades 

trilobed, 8-14 × 10-16 cm, coriaceous, the lobes 

shallow to quite deep, the apex rounded or obtuse, 

the base subcordiform or truncate, the margins 

entire or sinuate; upper surface dark green, shiny, 

glabrous, with palmate venation, somewhat 

sunken and yellowish; lower surface pale green, 

puberulous, the venation reticulate, prominent and 

puberulous; petioles robust, cylindrical, 2-4.5 cm 

long, glabrous, dilated at the base; pseudostipules 

rounded or reniform, 1-1.5 cm long, covering the 

stem. Flowers solitary, pendulous; peduncle up 

to 7 cm long; utricle ellipsoid, 2.5-4 cm long; tube 

straight, 4.5-7.5 cm long, ascending, forming an 

angle of 50° to 60° with the utricle; limb unilobed, 

greenish yellow with dark violet spots, the upper 

lobe with a caudate appendage, 10-31 cm long, 

ascending-arcuate. Capsules oblong, 6-8 cm long, 

with 6 longitudinal ribs; seeds numerous, 

membranaceous, triangular, ca. 7 mm long, 

compressed. 

Phenology: Flowering and fruiting almost 



Distribution: In forests and coastal thickets. 

Also on Cayo Santiago, Vieques, and St. John; 

along the Antilles, from Belize to Panama, 

Colombia, and the Guiana’s. 

Public forest: Piñones. 

References: González, F. A. 1990. Aristolochiaceae. Flora de Colombia. 12: 1-185; Rankin Rodríguez, 

R., 1998. Aristolochiaceae. Flora de la República de Cuba. Fascículo 1-2, 29 pp; Rankin Rodríguez, R. 

& W. Greuter. 1999. Taxon 48: 677-688. Rankin Rodríguez, R. 2001. La familia Aristolochiaceae en 

Cuba. Ph. D. Thesis, Universidad de La Habana, Cuba.

72 



5. Family ASCLEPIADACEAE 


1a. Stamens with the filaments free; corolla violet, 5 cm long or longer, the floral tube 2 cm long or 

longer ......................................................................................................................... 1. Cryptostegia 

1b. Stamens with the filaments connate into a tube that is united to the style or to the stigma to form a 

gynostegium; corolla white, cream-colored, pale yellow, or pale green, < 2.5 cm long, the floral 

tube < 5 mm long ............................................................................................................................... 2 

2a. Pollinium erect ................................................................................................................................... 3 

2b. Pollinium horizontal or pendulous.................................................................................................... 4 

3a. Leaves fleshy, broadly elliptical; inflorescences of 25 or more flowers; lobes of the corona horizontal, 

adnate to the staminal tube .................................................................................................... 4. Hoya 

3b. Leaves coriaceous, elliptical; inflorescence of 8-20 flowers; lobes of the corona erect, each of them 

borne on the basal portion of the dorsal side of the anthers ........................................ 5. Marsdenia 

4a. Petals 1.5-2 cm long, lanceolate or oblong ....................................................................................... 5 

4b. Petals < 0.5 cm long, ovate, lanceolate, oblong, or deltoid ............................................................. 6 

5a. Petals ca. 1.5 mm wide at the base; stems with soft, whitish hairs; leaves cordiform at the base, with 

5 acicular glands at the union with the petiole .......................................................... 8. Oxypetalum 

5b. Petals ca. 4 mm wide at the base; stems with reflexed, somewhat rough hairs, glabrous when 

mature; leaves deeply cordiform at the base, with 2 triangular glands at the union with the petiole 

.............................................................................................................................. 3. Gonolobus 

6a. Inflorescences with peduncles 1.5 cm long or longer; pedicels 1 cm long or longer..2. Funastrum 

6b. Inflorescences with short or subsessile peduncles; pedicels < 1 cm long ........................................ 7 

7a. Corolla rotate; pollinium horizontal ................................................................................ 6. Matelea 

7b. Corolla campanulate; pollinium pendulous ............................................................... 7. Metastelma 

1. CRYPTOSTEGIA 

Twining lianas or scandent shrubs, producing abundant milky latex. Stems cylindrical, glabrous, 

lenticellate. Leaves opposite, petiolate; blades simple, without glands; stipules interpetiolar, minute. 

Flowers in terminal cymes. Calyx campanulate, of 5 more or less elongate sepals; corolla infundibuliform; 

corona of 5 filiform lobes; stamens 5, inserted at the base of the corolla, the filaments short and free, 

the anthers lanceolate, concrescent around the stigma; ovary superior, bicarpellate, the stigma capitate. 

Fruit of 2 divergent, woody, boat-shaped follicles, with numerous ovate seeds, crowned with a tuft of 

long hairs. A genus of 2 species, native to the Old World tropics. 

Key to the species of Cryptostegia 

1a. Corona with bifid lobes; follicles 10-15.4 cm long ............................................... 1. C. grandiflora 

1b. Corona with simple lobes; follicles 5.8-13 cm long ................................... 2. C. madagascariensis

74 


1. Cryptostegia grandiflora R. Br., Bot. Reg. tab. 

435. 1820. 

Fig. 25. E-H 

Canario morado falso, Purple alamanda, 

Rubber vine 

Woody vine, twining, attaining 6 m in length, 

with abundant milky latex. Stems cylindrical, 

glabrous or pilose in the area of the nodes, dark 

brown, with numerous minute lenticels. Leaves 

opposite; blades 5-10 × 2-6.3 cm, elliptical to 

rounded, membranaceous, glabrous, the apex 

acute or obtuse, the base obtuse to rounded, 

decurrent on the petiole, without glands at the 

union with the petiole, the margins entire; 

venation pinnate, with 11-13 pairs of secondary 

veins; upper surface dull; lower surface pale, with 

obscure venation; petioles glabrous, 17-10 mm 

long; stipules interpetiolar and intrapetiolar. 

Flowers 6-9, arranged in dichasial cymes, 

pedunculate; bracts foliaceous, ovate, 1-1.2 cm 

long. Calyx green, campanulate, the sepals ovate, 

pubescent, 1-1.5 cm long; corolla infundibuliform, 

5-7 cm long, violet, whitish inside, the tube ca. 2 

cm long; corona with bifid, lanceolate lobes, ca. 

1.5 cm long. Follicles 2, divergent, brown when 

mature, 10-15.4 cm long, boat-shaped (keeled), 

woody; seeds reddish brown, ovate-lanceolate, 3- 

4 mm long, with long, white, silky hairs. 




naturalized. 

Distribution: Along the coast. Also on Vieques. 

Native to Madagascar, but widely cultivated in 


2. Cryptostegia madagascariensis Bojer ex 

Decne. in DC., Prodr. 8: 492. 1844. 

Fig. 25. A-D 

2. FUNASTRUM 

Rubber vine 

Twining woody vine or scandent shrub, 6-8 

m in length, with abundant milky latex. Stems 

cylindrical, glabrous, reddish brown, with few 

lenticels. Leaves opposite; blades 4-10 × 2-4.7 cm, 

elliptical, oblong, or ovate, coriaceous, glabrous, 

the apex short-acuminate, obtuse, or rounded, the 

base obtuse to rounded, short- decurrent on the 

petiole, without glands at the union with the 

petiole, the margins entire; venation pinnate, with 

14-16 pairs of secondary veins; upper surface dull; 

lower surface pale, with obscure venation; petioles 

glabrous, 0.6-1.5 cm long; stipules minute, 

intrapetiolar. Flowers few or many in pedunculate 

cymes; bracts foliaceous, lanceolate, ca. 5 mm 

long. Calyx green, campanulate, the sepals ovate, 

pubescent, 0.5-1.5 cm long; corolla 

infundibuliform, 3-6 cm long, violet, the tube 

darker inside, the lobes abaxially whitish in the 

overlapping portion; corona violet, with subulate, 

simple lobes, ca. 1 cm long. Follicles 2, divergent, 

brown when mature, 5.8-13 cm long, winged or 

keeled, woody; seeds reddish brown, ovatelanceolate, 

ca. 3 mm long, with long, creamcolored, 

silky hairs. 

Phenology: Collected in flower in August and 

from December to February and in fruit in August. 

Status: Exotic, common, cultivated and 

naturalized in Puerto Rico and the Virgin Islands. 

Distribution: In thickets and coastal forests. 

Also on Vieques, St. Croix, St. John, St. Thomas, 

and Tortola; native to Madagascar, but of wide 

distribution in the Neotropics. 

Public forests: Guánica and Río Abajo. 

Commentary: This species was treated 

erroneously for the flora of St. John (Acevedo- 

Rodríguez, 1996) under the name of Cryptostegia 

grandiflora R.Br. 

Herbaceous or slightly woody vines, twining, with abundant milky latex. Stems slender, glabrous 

or puberulous. Leaves opposite, petiolate; blades simple, the upper surface with 2-8 glands at the union 

with the petiole; pseudo-stipules absent. Inflorescences of subaxillary umbelliform cymes. Calyx deeply 

lobed, the sepals with or without glands at the base on the inner surface; corolla rotate, slightly 

campanulate, the lobes short; corona of 5 inflated, ovoid or ellipsoid, free and erect vescicles; stamens 

5, erect, with distal appendages adpressed to the apex of the stigma; pollinium pendulous; ovary superior,


bicarpellate, the gynostegium sessile or stipitate, the stigma forming a conical or apiculate head. Fruit 

of (1) 2 follicles, asymmetrically fusiform; seeds usually numerous, obovate, flattened, with a tuft of 

hairs on the apical portion. A genus of about 15 species, of tropical and subtropical areas of the New 

World.

76 


1. Funastrum clausum (Jacq.) Schltr., Repert. 

Spec. Nov. Regni Veg. 13: 283. 1914. 

Fig. 32. G-J 

BASIONYM: Asclepias clausa Jacq. 

SYNONYM: Sarcostemma clausum (Jacq.) Schult. 



glabrous, slender. Leaves opposite; blades 3-6 × 

0.8-2 cm, elliptical or elliptic-lanceolate, the apex 

obtuse with a small, tapering point, the base obtuse 

or rounded, the margins entire; upper surface 

puberulous, with a prominent midvein, with a pair 

of triangular glands at the union with the petiole; 

lower surface pale green, dull, puberulous, with a 

prominent midvein; petioles 0.5-3 mm long, 

pubescent; stipules triangular, minute. Flowers 

3. GONOLOBUS 

Herbaceous or slightly woody vines, twining, with abundant milky latex. Leaves opposite, petiolate; 

blades simple, the upper surface with minute triangular glands where they join the petiole; intrapetiolar 

stipules minute. Flowers in axillary, umbelliform, or racemose, pedunculate or subsessile cymes. Calyx 

of 5 elongated sepals, with a gland at the base on the inner surface; corolla rotate or subcampanulate, 

sometimes forming a short tube at the base, the petals linear, oblong, lanceolate, or rounded; corona 

with a single series of lobes; stamens 5, inserted at the base of the corolla, the filaments connate at the 

base to form a short tube, the anthers with a fleshy dorsal appendage, pollinia horizontal or pendulous; 

ovary superior, bicarpellate, the stigma 5-angled, depressed. Fruit of 1 angular or winged follicle; 

seeds numerous, crowned with a tuft of long hairs. A genus of approximately 100 species, mostly 

native to Mexico and Central America, with several species in the Antilles. 

1. Gonolobus stephanotrichus Griseb., Cat. Pl. 

Cub. 177: 1866. 

Fig. 26. A-G 

SYNONYM: Vincetoxicum stephanotrichum (Griseb.) Britton 

Herbaceous or slightly woody vine, twining, 

attaining 10 m in length, with abundant milky 

latex. Stems cylindrical, pubescent with reflexed, 

somewhat rough hairs, glabrescent when mature. 

Leaves opposite, pendulous; blades 3.5-12 × 1.5- 

5.5 cm, ovate or oblong-ovate, membranaceous, 

glabrous, the apex acuminate, the base deeply 

cordiform, with a pair of minute triangular glands 

at the union with the petiole, the margins entire 

or crenate; upper surface dark green, dull; lower 

surface pale, dull, with a prominent midvein; 

petioles cylindrical, 4-6 cm long, pubescent. 

Flowers 2 or more in sessile or subsessile cymes; 

pedicels pilose, ca. 5 mm long. Calyx green, 

numerous (10-20), arranged in umbelliform 

cymes; peduncle 4-12 cm long; pedicels 1-2 cm 

long, pubescent. Calyx green, the sepals 

triangular, pubescent on the outer surface, ca. 3 

mm long; corolla rotate, white, pubescent outside, 

the lobes ovate, ca. 7 mm long; corona of oblongobovoid 

vescicles, white. Follicles cylindrical, 5- 

8 × 1 cm. 

Phenology: Flowering from January to 

October and fruiting from November to January. 

Status: Exotic, cultivated, although 

uncommon in the Virgin Islands. 

Distribution: Cited from St. Croix by West 

(1793) and by Eggers (1879) and from St. Thomas 

by Krebs (1847). From southern North America 

to Argentina, including the Antilles. 

tinged with pink, the sepals ovate-lanceolate, ca. 

5 mm long; corolla rotate, pale purple or green 

tinged with purple on both surfaces, the petals 

narrowly lanceolate, ca. 15 mm long, abaxially 

pilose; corona purple, double, with 5 outer lobes 

and 2 short inner lobes; pollinium pendulous; 

ovary sessile, the stigma purple. Follicles green, 

fusiform, 15-17 cm long, with 5 longitudinal ribs; 

seeds light brown, ovate, ca. 6.5 mm long, with 

long, silky hairs at the apex. 

Phenology: Collected in flower in October and 

January and in fruit in April, September, and 

November. 


Distribution: In moist or wet forests of the 

Cordillera Central, the Sierra de Luquillo, and 

the zone of mogotes. Also in Cuba and Hispaniola. 

Public forests: El Yunque, Guilarte, Río Abajo, 

and Toro Negro.


4. HOYA 

Twining vines or epiphytes, herbaceous or slightly woody, sometimes subshrubs, producing abundant 

milky latex. Leaves opposite; blades simple, fleshy, the upper surface with minute triangular glands 

where they join the petiole; intrapetiolar stipules minute or absent. Flowers usually numerous, in 

axillary, umbelliform, pedunculate or sessile racemes. Calyx of 5 free sepals that have a gland at the 

base on the inner surface; corolla rotate or campanulate, the tube short, cylindrical or urceolate, the 

petals fleshy or waxy; corona simple, with 5 horizontal segments adnate to the staminal tube; stamens 

5, inserted at the base of the corolla, the filaments fused to a gynostegium, which is not markedly 

capitate, the anthers with a distal appendage; pollinium erect and solitary; ovary superior, bicarpellate, 

the stigma conical. Fruit of 2 slender follicles; seeds numerous, crowned with a tuft of long hairs. A 

genus of approximately 300 species, native to southern Asia, Malaysia, and Australia, some of which 

are cultivated as ornamentals throughout the tropics. 

1. Hoya australis R. Br. ex Traill, Trans. Hort. 

Soc. London 7: 28. 1830. 

Fig. 26. H-K 

Mata de cera, Wax plant 

Slightly woody vine, twining, 6-7 m in length, 


puberulous, green. Leaves opposite; blades 5-12 

× 2.5-5.5 cm, oval, elliptical, or ovate, fleshy, 

glabrous, acute, acuminate, or subrounded at the 

apex, the base obtuse or rounded, with 3-5 minute 

triangular glands at the union with the petiole, 

the margins entire, the venation pinnate; upper 

surface dull, the midvein sunken; lower surface 

dull, with a prominent midvein and puberulous; 

petioles subcylindrical, 1-2 cm long, pubescent. 

Flowers numerous, arranged in umbelliform 

racemes, clustered at the tip of a peduncle; basal 

5. MARSDENIA 

flowers early deciduous; bracts overlapping and 

persistent on the peduncle; peduncle 1.5-3 cm 

long; pedicels 2-4 cm long, pale green. Calyx 

whitish green, of lanceolate sepals, ca. 3 mm long; 

corolla rotate, white, with a bright pink to purple 

center, the lobes broadly ovate, 4-5 mm long, 

densely papillose inside; corona white, the 

segments 3 mm long. Follicles linear, cylindrical, 

6-13 cm long; seeds oblong, ca. 4 mm long, with 

long, whitish, silky hairs. 

Phenology: Collected in flower in December. 

Status: Exotic, commonly cultivated in Puerto 

Rico and the Virgin Islands. 

Distribution: Native to Australia; widely 

cultivated in the Neotropics. 

Commentary: This species has been cited as 

Hoya carnosa (L.) R.Br. by Britton and P. Wilson 

(1925) and by Liogier (1995). 

Herbaceous or woody vines, twining, less frequently erect shrubs, with abundant milky latex. Stems 

cylindrical or angular, lenticellate or with abundant cork when mature. Leaves usually opposite, petiolate; 

blades simple, the upper surface with 2 or more minute glands where they join the petiole; intrapetiolar 

stipules minute. Flowers in subaxillary or terminal, umbelliform or racemose, dichasial or paniculiform 

cymes, with few to many flowers. Calyx of 5 deep or short sepals, with 1-2 (3-5) glands at the base on 

the inner surface, or without glands; corolla rotate, campanulate, urceolate, or hypocrateriform; corona 

absent or of 5 segments, each of them borne on the basal portion of the dorsal side of the anthers; 

stamens with the filaments fused to a sessile or stipitate gynostegium; stamens 5, the anthers erect, 

usually rectangular; pollinia erect; ovary superior, bicarpellate, the stigma conical or depressed. Fruit 

a fusiform or ovoid, ellipsoid, or angular follicle, winged at the margin; seeds numerous, crowned with 

a tuft of long hairs. A tropical and subtropical genus of approximately 300 species.

78 



Key to the species of Marsdenia 

1a. Leaves with 5 or more minute glands on the portion adjacent to the petiole; corolla hypocrateriform 

or campanulate, white ........................................................................................................................ 2 

1b. Leaves with 2 acicular glands on the portion adjacent to the petiole; corolla rotate, purple-pink 

..................................................................................................................... 3. M. woodburyana 

2a. Leaves usually obovate; corolla campanulate, the tube ca. 3 mm long, the lobes ca. 5 mm long 

............................................................................................................................ 1. M. elliptica 

2b. Leaves elliptical; corolla hypocrateriform, the tube 2-2.5 cm long, the lobes ca. 1 cm long 

..........................................................................................................................2. M. floribunda 

1. Marsdenia elliptica Decne. in DC., Prodr. 8: 

616. 1844. 

Fig. 27. A-D 



glabrous, dark brown, with some scattered 

lenticels. Leaves opposite; blades 5-13 × 2-6 cm, 

obovate or less frequently elliptical or oblongelliptical, 

coriaceous, glabrous, the venation not 

prominent, the apex short-acuminate, the base 

obtuse to rounded, the margins entire, revolute; 

upper surface dull, with 5-10 minute, dark brown 

glands on the portion adjacent to the petiole; lower 

surface pale; petioles glabrous, 1-2 cm long. 

Flowers 8-15, arranged in subaxillary cymes, 

glabrous, shorter than the petiole; peduncles ca. 

5 mm long; pedicels ca. 3 mm long. Calyx 

crateriform, the sepals ovate, obtuse, ca. 3 mm 

long, glabrous, ciliate on the margin; corolla 

campanulate, white, ca. 8 mm long, glabrous, 

short-tubular (ca. 3 mm) at the base, the lobes 

ovate, obtuse, ca. 5 mm long; corona of simple, 

subulate, erect, connivent segments; stigma 

turbinate. Follicles fusiform, green, ca. 15 x 1 cm. 

Phenology: Collected in flower in June and 

in fruit in November. 

Status: Endemic to Puerto Rico, very rare. 

Distribution: Originally known from two 

collections, the type collection, made by Wydler 

(no. 308) in 1827 but without a specific locality, 

and the second collection, made by Sintenis (no. 

1452) in June of 1885 from Monte Jiménez in the 

Sierra de Luquillo. Recently rediscovered in the 

Río Abajo Forest, Monte Torresilla and in Sabana 

Hoyos. 


Commentary: In sterile condition, this species 

can be confused with Matelea variifolia; 

nevertheless, Marsdenia elliptica is distinguished 

by the presence of coriaceous leaves with 5-10 

glands of various forms on the upper surface in 

the portion where they join the petiole (vs. 

membranaceous with 2 acicular glands in M. 

variifolia). 

2. Marsdenia floribunda (Brongn.) Schltr. in 

Urb., Symb. Antill. 1: 275. 1899. 

Fig. 27. E-H 

BASIONYM: Stephanotis floribunda Brongn. 

Estefanotis, Florador, Madagascar jasmine, 

Wax plant 

Woody vine, twining, 3-5 m in length, with 

abundant milky latex. Stems cylindrical, glabrous, 

striate, lenticellate. Leaves opposite; blades 5-12 

× 2.2-3.5 cm, elliptical, coriaceous, glabrous, the 

apex short-acuminate, the base rounded to 

subcordiform, the margins entire, markedly 

revolute; upper surface dull, with 5 minute, 

conical glands, on the portion adjacent to the 

petiole; lower surface pale green, dull, with a 

prominent midvein; petioles 1-2 cm long, 

glabrous; intrapetiolar stipules acicular, minute. 

Flowers numerous, fragrant, in axillary umbels, 

ascending; peduncle 2-2.5 cm long; pedicels 2.5- 

3 cm long. Calyx crateriform to rotate, green, the 

sepals oblong, obtuse, sometimes minutely ciliate, 

glabrous; corolla white or cream-colored, 

hypocrateriform, 2.5-5 cm long, glabrous, the 

lobes oblong, obtuse, ca. 1 cm long; corona absent;

80 



anthers sessile, lanceolate; stigma conical. 

Follicles 7.5-10 cm long. 

Phenology: Collected in flower in September. 

Status: Exotic, common, cultivated. 

Distribution: Native to Madagascar. 

Commonly cultivated in gardens for its attractive, 

fragrant flowers. Also in St. Croix. 

3. Marsdenia woodburyana Acev.-Rodr., 

Brittonia 51: 167. 1999. 

Fig. 28. A-F 

Woody vine, twining, 5-8 m in length, with 

abundant milky latex. Stems thick, cylindrical, 

glabrous, sparsely lenticellate, grayish, attaining 

ca. 3 cm in diameter at the base. Leaves opposite; 

blades 6-6.5 × 3.5-4 cm, ovate, coriaceous, the 

apex acute, obtuse, or short-acuminate, the base 

rounded, the margins entire, markedly revolute; 

upper surface glabrous, with 2 acicular glands on 

the portion adjacent to the petiole; lower surface 

pale green, glabrous or minutely puberulous; 

petioles 1.5-2 cm long, glabrous, sulcate on the 

upper surface; stipules absent. Flowers about 20, 

6. MATELEA 

arranged in umbelliform racemes; peduncles ca. 

1 cm long; pedicels 7-10 mm long. Calyx light 

green, rotate, of 5 imbricate sepals, ovate-rounded, 

obtuse, 3-3.5 mm long, puberulous on the abaxial 

surface, ciliate on the margins; corolla rotate, the 

lobes ovate, 5.5-7 mm long, glabrous, light green 

in the central portion, purple-pink toward the 

markedly revolute margins, the apex reflexed; 

corona with segments of staminal origin (the distal 

portion of the filaments), fleshy, asymmetricpyramidal, 

ca. 3 mm long, projecting into a short 

distal appendage, triangular, inflexed, pink; 

anthers oblong, ca. 2 mm long; stigma flat. 

Follicle lanceolate-fusiform, slightly compressed, 

woody, 11-13 × 2-3 cm; seeds numerous, 

membranaceous, ovate, ca. 1 cm long, with a long 

tuft of brown-cream hairs. 

Phenology: Collected in flower in May and in 

fruit in January and July. 

Status: Endemic to Puerto Rico, extremely rare 

or threatened. 

Distribution: Known only from Caña Gorda, 

Guánica (Acevedo-Rdgz. and L. Ramírez 10174; 

Proctor 39358; L. Ramírez 48). 

Public forest: Guánica. 

Erect subshrubs or twining vines, with abundant milky latex; with or without an indumentum of 

glandular hairs. Leaves opposite, petiolate; blades simple, with glands at the base; stipules absent. 

Flowers in axillary cymes, with short peduncles or subsessile. Calyx with 5 elongated sepals, usually 

glandular inside at the base; corolla subrotate, more or less lobed; corona annular, of staminal origin; 

stamens 5, inserted at the base of the corolla, the filaments connate, forming a short tube, the anthers 

concrescent around the style, pollinia horizontal or subpendulous; ovary superior, of two free carpels, 

the stigma capitate, projecting beyond the anthers. Fruit a fusiform to ovoid follicle, smooth or muricate; 

seeds numerous, with a tuft of hairs at the apex. A genus of 300 species, distributed from southern 

North America to South America, including the Antilles. 

Key to the species of Matelea 

1a. Plants robust, slightly woody; stems 4 mm or more in diameter; leaves densely pubescent on the 

lower surface; follicles ovoid-fusiform, verrucose .................................................... 1. M. maritima 

1b. Plants herbaceous; stems 2 mm or less in diameter; leaves glabrous or puberulous on the lower 

surface; follicles elongate-fusiform, smooth ..................................................................................... 2 

2a. Leaves rounded, truncate, or subcordiform at the base, never obtuse or acute; sepals ovate, ca. 1.5 

mm long; lobes of the corolla lanceolate-triangular, not overlapping ..................... 2. M. sintenisii 

2b. Leaves variable at the base, sometimes rounded, but always with some obtuse or acute; sepals 

oblong-ovate, 2-2.5 mm long; lobes of the corolla rounded, imbricate .................. 3. M. variifolia

82 


1. Matelea maritima (Jacq.) Woodson, Ann. 


Fig. 29. A-D 

BASIONYM: Asclepias maritima Jacq. 

SYNONYM: Ibatia maritima (Jacq.) Decne. 

Guanabanilla cimarróna, Pompón, 

Beach milk vine 


with abundant milky latex. Stems densely 

pubescent and sulcate when young, whitish and 

corky when mature, 4 mm or more in diameter. 

Leaves opposite; blades 4-10.5 × 3-7.5 cm, ovate, 

chartaceous, the apex acute or acuminate, the base 

cordiform, the margins entire; upper surface dull, 

pubescent, with 2 (3-7) acicular glands in the area 

adjacent to the petiole; lower surface pale, densely 

pubescent, the venation yellowish; petioles 

cylindrical, densely pubescent, 2-7 cm long. 

Flowers few, in small axillary cymes. Calyx green,


rotate, ca. 2 mm in diameter, the sepals ovate or 

lanceolate, pubescent; corolla rotate, greenish, 4- 

6 mm in diameter, the petals ovate, not imbricate; 

corona yellowish green, with short and rounded 

lobes. Follicles ovoid-fusiform, green, 5-8 × 3.5- 

4.5 cm, pubescent, verrocose; seeds numerous, 

brown, with long, white, silky hairs. 

Phenology: Flowering from October to 

November and fruiting from December to March. 



Also on Culebra, Desecheo, Mona, Vieques, St. 

Croix, St. John, St. Thomas, Tortola, and Virgin 

Gorda; Cuba, Hispaniola, the Lesser Antilles, and 

from Panama to northern South America. 


Susúa. 

2. Matelea sintenisii (Schltr.) Woodson, Ann. 


Fig. 29. E-H 

BASIONYM: Gonolobus sintenisii Schltr. 

SYNONYM: Vincetoxicum sintenisii (Schltr.) Britton 


with little or no milky latex. Stems slender, 

cylindrical, pubescent with retrorse hairs, 2 mm 

or less in diameter. Leaves opposite; blades 3-6.5 

× 1-3 cm, ovate or ovate-lanceolate, 

membranaceous, the apex acute or acuminate, the 

base rounded, truncate, or subcordiform, the 

margins entire or crenate; upper surface dark 

green, with two acicular glands at the union with 

the petiole; lower surface light green, glabrous or 

puberulous, with notable reticulate venation; 

petioles 1-2 cm long, with 2 lines of hairs along 

the upper surface. Flowers few, arranged in sessile 

or subsessile cymes; pedicels ca. 5 mm long, 

pubescent. Calyx green, crateriform, of 5 ovate 

sepals, ca. 1.5 mm long; corolla rotate, light green, 

the petals free to the base, lanceolate-triangular, 

not imbricate, 3.5-5 mm long; corona annular. 

Fruits fusiform, green, to 14 cm long, smooth; 

seeds numerous, spathulate, dark brown, ca. 4 mm 

long, with the margin verrocose and a long tuft 

of white hairs. 

Phenology: Collected in flower in March and 

April. 

Status: Endemic to Puerto Rico, uncommon. 

Distribution: In moist forests in the cordilleras 

and the zone of mogotes. 



3. Matelea variifolia (Schltr.) Woodson, Ann. 


Fig. 29. I-L 

BASIONYM: Gonolobus variifolius Schltr. 

SYNONYMS: Vincetoxicum variifolium (Schltr.) Britton 

Matelea borinquensis Alain 


with abundant milky latex. Stems slender, 

cylindrical, puberulous, glabrescent, 2 mm or less 

in diameter; pubescence of curved, reflexed hairs. 

Leaves opposite; blades 4.5-10 × 2-4.5 cm, 

lanceolate, elliptical, or oblong, membranaceous, 

the apex acute or acuminate, the base acute, 

rounded, or cordiform, the margins entire; upper 

surface dark green, glabrous, with 2 acicular 

glands at the union with the petiole; lower surface 

pale green, glaucous, glabrous; petioles 0.8-4 cm 

long, with 2 lines of hairs along the upper surface. 

Flowers few, arranged in short-pedunculate 

cymes; pedicels 6-8 mm long. Calyx green, rotate, 

of 5 oblong-ovate sepals, 2-2.5 mm long; corolla 

rotate, pale green with dark green venation, 10- 

14 mm in diameter, the lobes rounded, imbricate, 

ca. 5 mm long; corona annular, green. Fruits 

unknown. 

Phenology: Flowering from May to October. 


Distribution: In moist forests in the cordilleras 

and on mogotes. 


Río Abajo, and Toro Negro. 

Commentary: After a careful study of the type 

specimen of Matelea borinquensis Alain, I have 

come to the conclusion that it represents Matelea 

variifolia. The type specimen of M. borinquensis 

(Alain et al. 35111) is too poor to characterize its 

morphology adequately. The size and shape given 

for the petals cannot be considered as valid 

because they are based on floral buds. In addition, 

the characterization of the corona (“lobis 

triangularibus 1.5 mm longis”) can not be 

critically evaluated because it is not discernible 

on the specimen.

84 



7. METASTELMA 

Herbaceous vines, twining, with abundant milky latex. Leaves opposite, small, petiolate, persistent 

or deciduous; blades simple, the portion of the upper surface adjacent to the petiole glandular or not, 

the glands (if present) sometimes deciduous; stipules minute, interpetiolar. Flowers small, produced in 

axillary, sessile or pedunculate cymes. Calyx of 5 more or less deep sepals, usually with glands at the 

base on the inner surface; corolla campanulate, the petals usually pubescent on the inner surface; 

corona of 5 lobes; stamens 5, inserted at the base of the corolla, concrescent around a conical style; 

pollinia pendulous. Fruit of 2 linear or fusiform follicles, divaricate, with numerous seeds, with a tuft 

of hairs at the apex. A neotropical genus of about 150 species. 

Commentary: The distinction between Metastelma and Cynanchum is somewhat imprecise, because 

the majority of the characters used to distinguish them overlap. The characters of the corona are the 

only ones that permit a clear differentiation. Metastelma has the corona lobes fused to the corolla, 

alternate with the petals, and not fused to the gynostegium, while Cynanchum has the corona lobes 

fused only to the gynostegium. Metastelma is restricted to the New World; Cynanchum, on the other 

hand, is found in the Old World. This distinction is not recognized by all authors, and in some floristic 

works Metastelma is considered to be a synonym of Cynanchum. In this book, both genera are recognized 

as valid to maintain concordance with other floristic works for the Caribbean region. 

Key to the species of Metastelma 

1a. Vine deciduous or with very few leaves during the flowering period; leaves without glands at the 

union with the petiole ........................................................................................................................ 2 

1b. Vine with abundant leaves during the flowering period; leaves with or without glands at the union 

with the petiole ................................................................................................................................... 3 

2a. Lobes of the corolla glabrous ................................................................................ 3. M. leptocladon 

2b. Lobes of the corolla strigulose on the inner surface .................................................. 5. M. monense 

3a. Upper surface without glands at the union with the petiole ............................................................ 4 

3b. Upper surface with acicular glands at the union with the petiole ................................................... 6 

4a. Leaves linear ................................................................................................................. 4. M. lineare 

4b. Leaves oblong or oblanceolate........................................................................................................... 5 

5a. Leaves oblong to oblanceolate; lobes of the corolla ovate-lanceolate, acute to obtuse; gynostegium 

sessile; segments of the corona lanceolate, almost as long as the lobes of the corolla... 5. M. monense 

5b. Leaves oblanceolate; lobes of the corolla lanceolate, acuminate; gynostegium stipitate; segments of 

the corona spiniform, much shorter than the lobes of the corolla........................ 1. M. anegadense 

6a. Leaves ovate, oblong-ovate, or lanceolate ......................................................................................... 7 

6b. Leaves linear ................................................................................................................. 4. M. lineare 

7a. Gynostegium long-stipitate (1.5-2 mm long); corolla 3-3.5 mm long ............... 6. M. parviflorum 

7b. Gynostegium sessile or short-stipitate (< 0.3 mm long); corolla ca. 2 mm long ... 2. M. decipiens

86 


1. Metastelma anegadense Britton, Bot. Porto 

Rico. 6: 97. 1925. 

Fig. 30. A-D 

SYNONYM: Cynanchum anegadense (Britton) Alain 

Herbaceous vine, twining, glabrous, attaining 

1 m in length, with abundant milky latex. Stems 

with branches that intertwine like a braid; 

branches green, cylindrical, smooth, < 1 mm in 

diameter. Leaves perennial, opposite, usually 

clustered on short lateral branches; blades 1-3.5 

× 0.2-0.8 cm, oblong to oblanceolate, coriaceous, 

the apex rounded to apiculate, the base acute, the 

margins entire, revolute; upper surface dull, with 

the midvein sunken, without glands at the union 

with the petiole; lower surface pale, with a 

prominent midvein, the secondary veins not 

visible; petioles 2-3 mm long; stipules 

interpetiolar, triangular, 0.5 mm long. Flowers 

few, in sessile cymes. Calyx green, crateriform, 

the sepals ovate-lanceolate, ca. 1 mm long; corolla 

yellowish, the lobes lanceolate, acuminate, 2-4 

mm long, puberulous toward the apex, strigose 

in the center on the inner surface; corona with 

spiniform lobes, ca. 0.5 mm long; gynostegium 

short-stipitate, ca. 0.7 mm long. Follicles 

fusiform, 4-5 cm long, glabrous; seeds numerous, 

light brown, ca. 4 mm long, elliptical, with a tuft 

of long hairs at the apex. 

Phenology: Collected in flower in January, 

February, and July and in fruit in June. 

Status: Endemic to Anegada and Tortola, 

uncommon. 

Distribution: Known from few collections 

from the sandy coasts of Anegada and Tortola. 

Commentary: This species was cited by 

Woodbury et al. (1977) and by Liogier and 

Martorell (1982) for Mona Island based on the 

erroneous identification of Metastelma monense. 

2. Metastelma decipiens Schltr. in Urb., Symb. 

Antill. 1: 249. 1899. 

Fig. 30. E-H 

SYNONYMS: Cynanchum decipiens (Schltr.) Alain, non C.K. 

Schneid. 

Metastelma fallax Schltr. 

Metastelma grisebachianum Schltr. 

Cynanchum grisebachianum (Schltr.) Alain 

Metastelma albiflorum Schltr. 

Metastelma decaisneanum Schltr. 

Cynanchum decaisneanum (Schltr.) Alain 

Cynanchum cheesmanii Woodson 



cylindrical, puberulous, glabrous when mature, 

1-2 mm in diameter, much branched, intertwining 

like a braid. Leaves perennial, opposite; 1-3(5) × 

0.5-1.5(2.5) cm, oblong-ovate, ovate or lanceolate, 

chartaceous, the venation pinnate, the apex acute 

or obtuse and apiculate, the base rounded, 

subcordiform, or truncate, the margins entire, 

revolute, and usually ciliate; upper surface dull, 

the venation flat, with a pair of acicular glands at 

the union with the petiole; lower surface pale, 

usually punctate, with slightly prominent 

venation; petioles 0.2-1 cm long; stipules 

interpetiolar, deltoid, ca. 0.4 mm long. Flowers 

few, in sessile or short-pedunculate cymes; 

pedicels 1.5-2 mm long. Calyx crateriform, green, 

the sepals ovate, ca. 1 mm long, more or less 

ciliate; corolla campanulate, light green outside, 

white or cream-colored inside, ca. 2 mm long, 

the lobes oblong-lanceolate, ca. 1.5 mm long, 

puberulous in the interior and on the margins, 

usually strigose toward the center; corona white, 

with linear or linear-spathulate segments, ca. 1.5 

mm long; gynostegium sessile (< 0.3 mm long), 

green in the stigmatic portion. Follicle fusiform, 

3.5-5.5 cm long, glabrous; seeds numerous, with 

a long tuft of hairs at the apex. 




Distribution: Common in coastal thickets and 

in the zone of mogotes. Also on Culebra, 

Desecheo, Cayo Icacos, Cayo Lobos, Cayo Luis 

Peña, Vieques, St. Croix, St. John, St. Thomas, 

and Virgin Gorda; Cuba, Hispaniola, the Lesser 

Antilles, and Tobago. 

Public forests: Boquerón, Ceiba, Guajataca, 

Maricao, Piñones, Río Abajo, and Tortuguero. 

Commentary: Metastelma decipiens and M. 

fallax were described by Schlechter in 1899 in 

the same publication, as doubtfully different. 

These two entities cannot be maintained as 

different species, because the characters used to 

distinguish them overlap. For this reason, Britton 

and P. Wilson (1923) placed M. fallax in 

synonymy with M. decipiens. Since neither 

specific epithet was previously used in the genus 

Metastelma, the decision of Britton and P. Wilson, 

to apply the name M. decipiens to this taxon, 

should be maintained, obviously having priority


over the decision of Liogier (1995) to place M. 

decipiens in synonymy under M. fallax. 

I have decided to place Metastelma 

grisebachianum Schltr. in synonymy under M. 

decipiens, because the characters used to separate 

the two species also overlap. These include the 

pubescence on the petals and the form of the 

corona segments, which in my view are not 

sufficient to distinguish two biological entities. 

In addition, the disjunction in the distribution of 

M. decipiens does not make biological sense, 

because it has only been cited for Puerto Rico, St. 

Thomas, and Tobago, being absent in the Lesser 

Antilles. By recognizing M. grisebachianum as a 

synonym of M. decipiens, the distribution of M. 

decipiens is expanded to include the Lesser 

Antilles. 

3. Metastelma leptocladum (Decne.) Schltr. in 


Fig. 30. I-O 

BASIONYM: Vincetoxicum leptocladum Decne. 

SYNONYMS: Cynanchum leptocladum (Decne.) Jiménez 

Metastelma ephedroides (Griseb.) Schltr. 

Amphistelma ephedroides Griseb. 

Cynanchum ephedroides (Griseb.) Alain 



cylindrical, glabrous, ca. 1 mm in diameter, with 

numerous pendulous branches, which sometimes 

intertwine like braids. Leaves opposite, deciduous 

during the flowering period; blades 0.4-1.5 cm × 

ca. 0.7 mm, narrowly elliptical to linear, the 

secondary venation not visible, the apex acute, 

the base attenuate, the margins entire, revolute; 

upper surface glabrous, without glands at the 

union with the petiole; lower surface puberulous; 

petioles ca. 1.5 mm long; intrapetiolar stipules 

acicular, ca. 0.2 mm long. Flowers few, in axillary 

cymes; pedicels 3.5-4.5 mm long. Calyx 

crateriform, green, the sepals oblong, puberulous, 

0.5-0.7 mm long; corolla creamy yellow, ca. 2 mm 

long, the lobes oblong, glabrous, free to the base; 

corona crateriform, with triangular segments, ca. 

0.3 mm long, shorter than the gynostegium; 

gynostegium sessile. Follicles fusiform, glabrous, 

4-6 cm long; seeds numerous, with a tuft of hairs 


Phenology: Collected in flower in March and 

from July to November. 

Status: Native, relatively common. 

Distribution: In moist or dry forests in central 

and western Puerto Rico. Also in Cuba, Jamaica, 

and Hispaniola. 

Public forests: Guilarte, Maricao, Río Abajo, 


4. Metastelma lineare Bello, Anales Soc. Esp. 

Hist. Nat. 10: 292. 1881. 

Fig. 31. A-F 

SYNONYM: Cynanchum lineare (Bello) Alain 

Slender vine, twining, 2-5 m in length, with 

abundant milky latex. Stems green, cylindrical, 

more or less glabrous, 1 mm or less in diameter. 

Leaves perennial, opposite or subwhorled; blades 

1.5-4.5 cm × 1.5-2 mm, linear, the apex acute or 

obtuse, subapiculate, the base obtuse or acute, the 

margins entire, revolute; upper surface glabrous 

with the midvein sunken, with a short acicular 

gland (or sometimes without a gland) where it 

joins the petiole; lower surface pale, with a 

prominent midvein, the secondary venation 

inconspicuous; petioles ca. 2 mm long, slender; 

intrapetiolar stipules acicular, 0.2 mm long. 

Flowers few, in subsessile cymes; pedicels slender, 

1.5-2.5 mm long. Calyx green, crateriform, the 

sepals ovate, ca. 1 mm long; corolla white inside, 

yellowish green outside, 3.5 mm long, the lobes 

lanceolate, 2.5-3 mm long, reflexed at the apex, 

the inner surface puberulous from the middle 

portion toward the apex, strigose from the middle 

portion toward the base; corona white, the 

segments narrowly lanceolate, 1.2-2 mm long, 

longer than the gynostegium; gynostegium sessile. 

Follicles linear, 3-5 cm long, glabrous, splitting 

in half to expose numerous dark brown, ovate 

seeds, 5-5.5 mm long, with long, silky hairs at 

the apex. 


fruiting from September to March. 


Distribution: In secondary forests at middle 

and upper elevations, in central and western 

Puerto Rico. Also on Mona and St. Thomas; Cuba. 

Public forests: Guánica, Maricao, Mona, Río 

Abajo, Susúa, and Toro Negro.

88 



5. Metastelma monense Britton, Bot. Porto Rico 

6: 97. 1925. 

Fig. 31. G-K 

SYNONYM: Cynanchum monense (Britton) Alain 

Herbaceous vine, twining, evergreen or 

partially deciduous during the flowering period, 

much branched, attaining 5 m in length, with 

abundant milky latex. Stems green, slender, 

cylindrical, glabrous, 1 mm or less in diameter. 

Leaves opposite or clustered on short axillary 

branches; blades 7-16 × 1.5-3 mm, oblanceolate, 

oblong or obovate, coriaceous, glabrous, the apex 

obtuse, rounded, or truncate and apiculate, the 

base obtuse or attenuate, the margins entire, 

revolute, slightly ciliate; upper surface dull, with 

the midvein sunken and whitish, without glands 

at the union with the petiole; lower surface pale 

green, with a prominent midvein, the secondary 

venation slightly conspicuous; petioles light green 

or yellowish, compressed and sulcate, 1.5-2 mm 

long; stipules interpetiolar, triangular, ca. 0.4 mm 

long. Flowers few, fragrant, arranged in sessile 

cymes; pedicels 1-1.5 mm long. Calyx crateriform, 

green, the sepals ovate, ca. 1 mm long, papillose 

or glabrous outside; corolla white, ca. 2.5 mm 

long, the lobes ovate-lanceolate, free to the base, 

the inner surface densely short-pubescent from 

the middle to the distal portion, strigulose in the 

central basal portion; corona white, with 

lanceolate segments, fleshy, almost as long as the 

lobes of the corolla; gynostegium sessile. Follicles 

fusiform, ca. 2.5 cm long, glabrous. 

Phenology: Collected in flower from 

December to May. 

Status: Endemic to Puerto Rico. 

Distribution: Locally common on Mona Island 

and recently discovered by Franklin Axelrod 

(UPRRP) in the Guánica Forest. 

Public forests: Guánica and Mona. 

8. OXYPETALUM 

6. Metastelma parviflorum (Sw.) R. Br., Mem. 

Wern. Soc. 1: 52. 1809. 

Fig. 31. L-P 

BASIONYM: Cynanchum parviflorum Sw. 

SYNONYM: Cynanchum parviflorum (R. Br.) Alain, nom. 

illegit. 



cylindrical, glabrous or puberulous, 2-3 mm in 

diameter. Leaves perennial, opposite; blades 1.4- 

3.5 × 0.8-1.5 cm, lanceolate or ovate, chartaceous, 

the venation pinnate, the apex acute or obtuse, 

usually mucronate, the base rounded, the margins 

entire or slightly undulate, revolute; upper surface 

glabrous, green, dull, with the midvein sunken, 

with a pair of acicular glands at the union with 

the petiole, these sometimes deciduous; lower 

surface pale green, dull, glabrous, with prominent 

venation. Flowers few, in sessile or shortpedicellate 

cymes; pedicels 1- 2.5 mm long. Calyx 

crateriform, green, puberulous, the sepals ovate, 

ca. 1 mm long, ciliate; corolla campanulate, 3- 

3.5 mm long, greenish outside, the lobes oblong, 

acute, ca. 2 mm long, pubescent inside, white 

along the margin and green in the central portion; 

corona white, the segments linear, ca. 2 mm long; 

gynostegium stipitate (1.5-2 mm long). Follicles 

fusiform, 4-6 cm long, divaricate, glabrous; seeds 

numerous, lanceolate, brown, ca. 5 mm long, with 

a tuft of silky hairs at the apex. 

Phenology: Flowering from October to May 

and fruiting from October to April. 


Distribution: Throughout Puerto Rico in areas 

of secondary or disturbed vegetation, at lower 

elevations. Also on St. Croix, St. Thomas, Tortola, 

and Virgin Gorda; the Lesser Antilles, Trinidad, 

and Isla Margarita. 

Public forests: Guánica, Piñones, Río Abajo, 


Twining, herbaceous vines or subshrubs, with abundant milky latex. Leaves opposite, small, petiolate; 

blades simple, glandular at the union of the petiole with the upper surface; stipules interpetiolar, 

minute. Flowers small, produced in axillary or terminal, umbelliform, sessile or pedunculate cymes. 

Calyx of 5 sepals, with or without glands at the base on the inner surface; corolla campanulate, the 

petals usually elongate; corona of 5 segments adnate to the corolla or free from it; stamens 5, inserted 

at the base of the corolla, the filaments connate at the base, forming a short tube; pollinium solitary, 

pendulous; stigma usually bifid. Fruit of 1-2 follicles, commonly ovate or ventricose, ribbed or winged,

90 



with numerous ovate or elliptical seeds, marginally dentate, with a tuft of hairs at the apex. A genus of 

about 100 species, distributed in the subtropics and temperate zones of the New World. 

1. Oxypetalum cordifolium (Vent.) Schltr. in 


Fig. 32 A-F 

BASIONYM: Gothofreda cordifolia Vent. 

Herbaceous vine, twining, attaining 4 m in 

length, with abundant milky latex. Stems 

cylindrical, pubescent, ca. 3 mm in diameter. 

Leaves opposite; blades 4-10 × 4-6 cm, ovate, 

pubescent, the apex acuminate or cuspidate, the 

base deeply cordiform, the margins entire; upper 

surface dull green, with 5 acicular glands at the 

union with the petiole; lower surface pale green, 

dull, with slightly prominent venation; petioles 

pubescent, slender, 2-3 cm long. Flowers few, 

arranged in umbelliform cymes; peduncle ca. 2 

cm long; pedicels 1-1.5 cm long. Calyx 

crateriform, green, the sepals linear-lanceolate, 

subulate, pubescent on the outer surface, 3.5-4 

mm long; corolla campanulate, yellowish or light 

green, pubescent outside, the tube ca. 4 mm long, 

the lobes linear-attenuate, 1.5-2 cm long; corona 

of cuneate-spathulate segments, fleshy, ca. 2.5 mm 

long; pollinia with an acicular appendage in the 

area where they connect to the translator arms; 

stigmatic branches ca. 5 mm long. Follicles 

fusiform, ca. 8 cm long. 

Phenology: Collected in flower in May. 



a single collection of Read (according to Urban, 

1910). Also on St. Thomas (according to Urban, 

1910); Cuba, Hispaniola, Jamaica, and from 

Mexico to South America. 

References: Forster, P. I., 1990. Hoya R. Br. (Asclepiadaceae) in Australia - an alternative 

classification. Austrobaileya 3: 217-234; Forster, P. I. and D. J. Liddle, 1991. Variation in Hoya australis 

R. Br. ex Traill (Asclepiadaceae). Austrobaileya 3: 503-521; Marohasy J. and P. I. Forster, 1991. A 

taxonomic revision of Cryptostegia R. Br. (Asclepiadaceae: Periplocoideae). Stevens, W. D. 2001. 

Asclepiadaceae. pags. 234-270, in: Stevens, W. D. et al. (eds.), Flora de Nicaragua. Mon. Syst. Bot. 

Missouri Bot. Gard. 85(1). 

6. Family ASTERACEAE 


1a. Leaves opposite .................................................................................................................................. 2 

1b. Leaves alternate ................................................................................................................................. 7 

2a. Capitula heterogamous, radiate, the central flowers with actinomorphic, tubular corollas, the 

peripheral ones zygomorphic, ligulate (with one of the lobules elongate in the form of a ligule). 3 

2b. Capitula homogamous, discoid, with all the flowers actinomorphic, tubular. ................................ 4 

3a. Leaves compound ................................................................................................................ 2. Bidens 

3b. Leaves simple ........................................................................................................ 11. Sphagneticola 

4a. Capitula of 4 phyllaries and 4 flowers ............................................................................. 6. Mikania 

4b. Capitula of more than 4 phyllaries and more than 4 flowers........................................................... 5

92 



5a. Receptacle (of the capitulum) with paleas at the base of each flower; branches of the style with 2 

stigmatic lines along their entire length ......................................................................... 10. Salmea 

5b. Receptacle without paleas; branches of the style with one stigmatic line, only on the lower half, or 

stigmatic lines absent ......................................................................................................................... 6 

6a. Capitula with several series of deciduous phyllaries, overlapping to form a cylindrical involucre; 

receptacle conical ...................................................................................................... 3. Chromolaena 

6b. Capitula with persistent phyllaries, in 1-2 series, forming a crateriform involucre; receptacle flat or 

convex ...................................................................................................................... 4. Koanophyllon 

7a. Capitula heterogamous, radiate, the peripheral flowers zygomorphic, with orange ligules, the central 

flowers actinomorphic, with yellow corollas ....................................................... 9. Pseudogynoxys 

7b. Capitula homogamous, discoid, with all the flowers tubular ........................................................... 8 

8a. Corollas bilabiate; margin of the leaves usually spinulose; stem with an obtuse or spiny, usually 

bifurcate, hardened or thickened area (callosity) at the base of the leaf ............. 1. Berylsimpsonia 

8b. Corollas tubular, 4-5-lobed; margin of the leaves entire, crenate, or denticulate; stem not forming a 

callosity nor spiny .............................................................................................................................. 9 

9a. Leaves with scales or with stellate hairs ......................................................................................... 10 

9b. Leaves with simple hairs ............................................................................................. 5. Lepidaploa 

10a. Pappus composed of bristles and short, irregular scales; corollas pale violet .......... 8. Piptocoma 

10b. Pappus composed exclusively of bristles, without scales; corollas white .............. 7. Piptocarpha 

1. BERYLSIMPSONIA 

Woody shrubs, clambering, with spiny pseudostipules, bifurcate and recurved, at the nodes. Leaves 

alternate, petiolate; blades simple, with entire, serrate, or spinulose margins. Inflorescences of capitula 

arranged in axillary cymes. Capitula homogamous, discoid, of 3-6 flowers; involucre turbinate, formed 

by 3-4 series of phyllaries; receptacle pubescent, without paleas. Flowers bisexual; corollas yellow, 

bilabiate; stamens 5, the anthers connate, exserted, the apex acute; ovary inferior, bicarpellate, unilocular, 

the style filiform, with 2 reflexed stigmatic branches. Fruit a fusiform or oblanceolate achene, pubescent, 

pilose, or glandular; pappus composed exclusively of barbate bristles. A genus of 2 species, distributed 

in Cuba, Hispaniola, and Puerto Rico. 

1. Berylsimpsonia vanillosma (C. Wright) B. L. 

Turner, Phytologia 74: 352. 1993. 

Fig. 33. A-C 

BASIONYM: Proustia vanillosma C. Wright 

SYNONYMS: Proustia krugiana Urb. 

Proustia stenophylla Urb. & Ekman 

Clambering shrub, woody, 2-5 m in length. 

Stems cylindrical, broadened at the base of the 

leaf to form an obtuse or spiny, usually bifurcate, 

hardened or thickened area (callosity). Leaves 

alternate, distichous; blades 4-7.5 × 1.5-3.7 cm, 

elliptical, oblong or ovate, chartaceous or 

subcoriaceous, the apex acute, obtuse, or 

acuminate, the base acute to subcordiform, the 

margins denticulate or spinulose or less frequently 

entire; upper surface scabrous, (strigulose), the 

venation somewhat sunken; lower surface 

appressed-pubescent (sericeous), with prominent 

venation; petioles 1-3 mm long, puberulous. 

Capitula of few flowers, subsessile, clustered in 

cymes to form a paniculiform or racemiform 

inflorescence; involucre cylindrical, ca. 1 cm high, 

phyllaries in 3-4 series. Corollas bilabiate, yellow, 

8-10 mm long, the inner lip 2-dentate, the outer 

one 3-dentate. Achene 3-6 mm long, linear, 

papillose; pappus of numerous barbate setae, 

yellowish, 6-8 mm long.

94 


Phenology: Flowering from November to July 

and fruiting from February to September. 


1. Bidens reptans (L.) G. Don. in Sweet, Hort. 

Brit. ed. 3, 360. 1839. 

Fig. 33. D-G 

BASIONYM: Coreopsis reptans L. 

Herbaceous vine, climbing or clambering, 2- 

5 m in length. Stems cylindrical, glabrous or 

puberulous, striate. Leaves opposite, 3-5-foliolate 

or 3-5-lobed; leaflets lanceolate or elliptical, 

membranaceous, the apex acuminate, the base 

acute in the terminal leaflet and unequal in the 

lateral ones, the margins serrate; upper and lower 

surface puberulous; petioles 1-1.8 cm long, 

glabrous. Capitula (1-3) in corymbs, produced at 

the tips of short axillary branches; involucre ca. 

7 mm high, phyllaries in two series, revolute; 

corollas yellow, those of the margin with ligules 

ca. 1 cm long; anthers dark brown; stigmas yellow. 

Achenes linear, 6-10 mm long, compressed, 4angled, 

with ascending hairs on the margins; 

pappus of 2 setae with retroflexed hairs. 

Phenology: Flowering from October to July. 


Distribution: Abundant in forests in the 

Cordillera Central, also in the zone of mogotes, 

in moist areas at middle and upper elevations. 

2. BIDENS 

Distribution: In forests and pastures on the 

southern slope of the Cordillera Central and in 

southwestern Puerto Rico, at middle and lower 

elevations. Also in Cuba and Hispaniola. 

Annual or perennial herbs, less frequently shrubs or vines. Stems usually striate. Leaves opposite; 

blades simple or compound; petioles and rachis narrowly winged or absent. Inflorescences terminal, 

with one to numerous pedunculate capitula; capitula heterogamous, generally radiate or discoid, with 

numerous flowers; involucre crateriform, with 2 series of unequal phyllaries, the outer ones green and 

narrow, the inner ones brown, with hyaline margins. Marginal flowers 5-12, usually sterile, with the 

corolla ligulate, yellow, white, or orange. Disc flowers few or numerous, bisexual, usually yellow, 

tubular; stamens 5, the anthers generally black, connate, exserted; ovary inferior, unilocular, the style 

filiform, with 2 stigmatic branches. Achenes slightly compressed, glabrous or pubescent; pappus 

composed of 1-6 erect bristles, retrorsely barbate. A genus of about 75 species, mostly of Mexico, also 

from North America to South America and in the Antilles. 

Key to the species of Bidens 

1a. Leaves 3-5-lobed or 3-5-foliolate, slightly pubescent; involucre ca. 7 mm high ........ 1. B. reptans 

1b. Leaves 2-3 times pinnatisect, glabrous; involucre 8-10 mm high ............................... 2. B. urbanii 

Also in Cuba, Jamaica, the Lesser Antilles to St. 

Vincent, and from Mexico to Venezuela. 

Public forests: Guajataca, Maricao, Río Abajo, 

and Susúa. 

2. Bidens urbanii Greenm., Field Mus. Bot. 2: 

271. 1907. 

Fig. 33. H-J 

SYNONYM: Bidens reptans var. urbanii (Greenm.) O.E. 

Schulz 

Bidens portoricensis Bello, non Spreng. ex DC. 

Herbaceous vine, climbing or clambering, 4- 

5 m in length. Main stem cylindrical, glabrous or 

puberulous, with numerous lateral branches along 

its length. Leaves opposite; blades simple, 2-3 

times pinnatisect, the segments chartaceous, longacuminate 

at the apex, the margins serrate, 

revolute; upper surface green, shiny, glabrous; 

lower surface pale green, dull, glabrous, pitted, 

with the pinnate venation darker, forming a 

marginal vein; rachis slender, marginate; petioles 

2-3.5 cm long, slender, sulcate, puberulous. 

Capitula (1-3) in corymbs, produced at the tips of 

short axillary branches; involucre 8-10 mm high,


96 


the phyllaries in 2 series, revolute; corollas yellow, 

those of the margin with ligules ca. 1.7 cm long; 

anthers dark brown; stigmas yellow, reflexed. 

Achenes linear, ca. 10 mm long, compressed, 4angled, 

with ascending trichomes on the margins; 

pappus of 2 retrorsely barbate setae. 

Phenology: Flowering from November to 

February. 


Distribution: In thickets and disturbed areas 

on serpentine soil. Known from Puerto Rico, 

eastern Cuba, and Campeche in Mexico. Has been 

cited erroneously for Jamaica, based on B. dissecta 

(O.E. Schultz) Sherff. 

Public forests: Maricao and Susúa. 

3. CHROMOLAENA 

Commentary: I agree with Greenman in 

recognizing this biological entity at the species 

level, because it is sufficiently different from B. 

reptans (see key). The argument presented by O.E. 

Schultz to recognize this entity at the level of 

subspecies is based on the variation presented by 

the leaves of other species of Bidens, specifically 

B. alba (L.) DC.; nevertheless, since the latter 

species is not close to B. reptans, the knowledge 

of its leaf morphology should not be applied to 

taxonomic decisions in B. reptans “sensu lato” 

(H. Robinson, pers. comm.). The characters that 

are used here to recognize B. urbanii at the 

specific level have likewise been used to recognize 

B. dissecta (from Jamaica), another species closely 

related to B. reptans. 

Erect or clambering herbs or shrubs. Stems usually pubescent. Leaves opposite, petiolate; blades 

simple, usually trinerved from the base. Capitula homogamous, discoid, with 10-40 flowers, arranged 

in corymbiform cymes. Involucre cylindrical; phyllaries deciduous, overlapping in 4-7 series. Flowers 

bisexual; corollas actinomorphic, tubular, shortly 5-lobed, the lobes usually papillose or glandular; 

stamens 5, the anthers inserted, connate; ovary inferior, the style filiform, with two linear stigmatic 

branches, elongate, exserted, usually with a sterile papillose appendage. Fruit a turbinate achene, 

usually with 5 longitudinal ribs; pappus with ca. 40 filiform bristles without scales. A genus of 166 

species distributed from the southern United States to southern South America, including the Antilles. 

1. Chromolaena borinquensis (Britton) H. Rob., 

Phytologia 20: 199. 1970. 

Fig. 34. A-D 

BASIONYM: Osmia borinquensis Britton 

SYNONYM: Eupatorium borinquense (Britton) B.L. Rob. 

Clambering shrub, slightly woody, much 

branched, 1.5-2.5 m in length. Stems cylindrical, 

glabrous, and smooth. Leaves opposite; blades 3- 

5 × 0.5-2 cm, chartaceous, deltate-lanceolate or 

narrowly lanceolate, 3-nerved from the base, the 

apex acuminate, the base cuneate or rounded, the 

margins with a marginal vein, revolute, remotely 

dentate-serrate; upper surface glabrous, with the 

venation sunken; lower surface glabrous, 

glandular-punctate; petioles 2-5 mm long, 

glabrous. Capitula numerous, arranged in 

corymbiform cymes; peduncles 1-2 cm long; 

involucre cylindrical, 0.5-1 cm high; phyllaries 

green, in 5 series, the outer much shorter than 

the inner, ovate, glabrous. Corollas pale violet, 

tubular, ca. 5 mm long, glabrous; anthers almost 

white, stigmatic branches pale violet. Achenes 

cuneiform, 3.5-4 mm long, triangular in cross 

section, glabrous, black, with the margins white; 

pappus of numerous white barbate setae, 3-3.5 

mm long. 


November to April. 


Distribution: On slopes of forests or thickets 

in the zone of mogotes. 

Public forest: Río Abajo.


4. KOANOPHYLLON 

Erect or scandent shrubs, small trees, or less frequently vines. Stems little or much branched, 

cylindrical. Leaves opposite or rarely alternate, petiolate; blade simple, with entire or serrate margins, 

with pinnate or trinerved venation. Capitula homogamous, discoid, with 5-20 flowers, arranged in 

pyramidal panicles or corymbs; involucre campanulate, with 2-4 unequal or subequal series of persistent 

phyllaries; receptacle without paleas. Flowers bisexual; corollas pale yellow or greenish or less frequently 

violet, actinomorphic, infundibuliform or tubular, glandular-pubescent on both surfaces or pubescent 

on the outer surface; stamens 5, the anthers connate; ovary inferior, the style filiform, with thickened 

branches, usually without glands, with or without a stigmatic line on the lower half. Fruit a prismatic 

achene, with 5 longitudinal ribs; pappus composed exclusively of bristles. A neotropical genus of about 

114 species. 

1. Koanophyllon polyodon (Urb.) R. King & H. 

Rob., Phytologia 32: 262. 1975. 

Fig. 34. E-F 

BASIONYM: Eupatorium polyodon Urb. 

SYNONYM: Eupatorium cordifolium sensu Bello, non Sw. 

Erect or clambering shrub, scarcely branched, 

1-1.5 m in length. Stems cylindrical, ferruginouspubescent, 

striate. Leaves opposite; blades 3-7 × 

1.8-4.5 cm, chartaceous, ovate, 3-nerved from the 

base, the apex acute, the base obtuse, rounded or 

cordiform, the margins denticulate-serrate; upper 

surface scabrid, with the venation sunken; lower 

surface ferruginous-puberulous, with prominent 

venation; petioles 3-7 mm long, puberulous. 

Capitula numerous in terminal corymbs; 

5. LEPIDAPLOA 

peduncles 1-2.5 cm long, ferruginous-pubescent. 

Involucre campanulate, ca. 2.5 mm high; 

phyllaries green, in 1(2) series, of equal size, 

puberulous. Corollas whitish, tubular, ca. 3 mm 

long, the lobes papillose; stigmatic branches 

exserted. Achenes prismatic, ca. 2 mm long, 

pentagonal in cross section, papillose, light 

brown; pappus of numerous white setae, barbate, 

3-3.5 mm long. 

Phenology: Collected in flower and fruit from 

November to January and in June. 


Distribution: On slopes in forests or thickets 

in the zone of mogotes and serpentine. 

Public forests: Maricao, Río Abajo, and Susúa. 

Annual or perennial herbs or subshrubs, erect or less frequently clambering. Stems usually much 

branched. Leaves alternate, sessile or petiolate; blades simple. Capitula homogamous, discoid, with 8- 

35 flowers, each more or less sessile, solitary or in terminal or axillary cymes; involucre campanulate, 

the phyllaries persistent, in 3-6 series. Flowers bisexual; corollas actinomorphic, narrowly campanulate, 

violet or less frequently white, usually pubescent or glandular, especially on the lobes; stamens 5, the 

anthers connate; ovary inferior, the style usually hispidulous, with 2 stigmatic branches. Fruit an 

angular achene, usually with 8-10 longitudinal ribs; pappus in 2 series, the inner of bristles and the 

outer of short, irregular scales. A neotropical genus of about 120 species. 

1. Lepidaploa borinquensis (Urb.) H. Rob., Proc. 

Biol. Soc. Wash. 103: 483. 1990. 

Fig. 34. G-K 

BASIONYM: Vernonia borinquensis Urb. 

SYNONYMS: Vernonia borinquensis Urb. var. stahlii Urb. 

Vernonia borinquensis Urb. var. hirsuta Gleason 

Vernonia borinquensis Urb. var. resinosa Gleason 

Slightly woody shrub, usually clambering, 1- 

4 (5) m in length. Stems slender, cylindrical, 

densely pubescent and striate when young, 

glabrescent. Leaves alternate, distichous; blades 

3-6 × 1.8-2.8 cm, coriaceous or rigid-coriaceous, 

ovate to lanceolate, the apex acute or acuminate, 

the base rounded or subcordiform, the margins

98 



ciliate, slightly undulate, and revolute; upper 

surface shiny, pubescent, scabrid, the venation 

usually sunken, covered with yellowish simple 

hairs; lower surface pale green, dull, with 

glandular dots, pubescent or sericeous, the 

venation prominent and pubescent; petioles 2-4 

mm long, pubescent. Capitula of 13-22 flowers, 

sessile or short-pedunculate, solitary, axillary at 

the end of short and flexuous branches; involucre 

campanulate, 5-9 mm high; phyllaries subulate, 

the basal series smaller than the distal ones. 

Corollas white or pale violet, 5-6 mm long, with 

6. MIKANIA 

Twining herbaceous or woody vines or less frequently erect shrubs. Stems cylindrical or hexagonal. 

Leaves opposite, petiolate; blades simple; pseudostipules present or absent. Capitula homogamous, 

discoid, numerous, composed of 4 flowers, arranged in spicate, paniculiform, or corymbiform, terminal 

or axillary cymes; involucre cylindrical with subequal and overlapping phyllaries, subtended by a 

subinvolucral bract, which is usually narrower than the phyllaries. Flowers bisexual; corollas 

actinomorphic, tubular, or infundibuliform, cream-colored or white; stamens 5, the anthers connate, 

exserted; ovary inferior, the style elongate, ascending, cream-colored, the apical portion with a long 

sterile appendage. Fruit a usually prismatic achene with 5 ribs, brown or black; pappus numerous, 

composed of bristles, without scales. A genus of about 400 neotropical species, only several in North 

America and the Old World tropics. About 10% of the species are shrubby; the great majority of these 

are found in the shrubby savannas of central and southern Brazil. 

Key to the species of Mikania 

five lanceolate to linear petals; anthers exserted; 

style pubescent, bifid. Achene ca. 1.3 mm long, 

conical, pubescent; pappus of bristles 5-6 mm 

long and scales ca. 0.8 mm long. 

Phenology: Flowering from November to July 

and fruiting from February to September. 

Status: Endemic to Puerto Rico, common. 

Distribution: In forests and pastures of the 

Cordillera Central and the zone of mogotes. 

Public forests: Carite, Guajataca, Guilarte, 


1a. Pseudostipules present ....................................................................................................................... 2 

1b. Pseudostipules absent ........................................................................................................................ 7 

2a. Pseudostipules entire, rounded, reflexed, 8-10 mm wide; leaves thick, fragile, slightly 

aromatic................................................................................................................. 3. M. fragilis 

2b. Pseudostipules divided into lanceolate or filiform segments ........................................................... 3 

3a. Capitula arranged in glomerules; lower surface densely covered with resinous dots. 1. M. congesta 

3b. Capitula arranged in corymbs; lower surface without dots or sparsely punctate ............................ 4 

4a. Involucre 3-4 mm high ............................................................................................. 4. M. micrantha 

4b. Involucre 6-9 mm high ...................................................................................................................... 5 

5a. Leaves pubescent on both surfaces; stems hexagonal .............................................. 2. M. cordifolia 

5b. Leaves glabrous or puberulous; stems subcylindrical or obscurely angular .................................... 6 

6a. Leaves coriaceous, the upper surface scabrid, the lower surface puberulous ....... 8. M. stevensiana 

6b. Leaves membranaceous, glabrous ........................................................................ 5. M. odoratissima 

7a. Blade of the leaves 3-6 cm long, with the venation pinnate, the margins entire ......... 7. M. porosa 

7b. Blade of the leaves 5-9 cm long, with 3 main veins from the base, the margins denticulate 

..................................................................................................................... 6. M. pachyphylla

100 


1. Mikania congesta DC., Prodr. 5: 197. 1836. 

Fig. 35. A-D 

SYNONYMS: Mikania micrantha var. congesta (DC.) L. B. 

Rob. 

Mikania scandens sensu A. Stahl, non Willd. 

Guaco falso 


Stems cylindrical or angular, striate, puberulous, 

or pilose. Leaves opposite; blades 2.5-14 × 1-10 

cm, deltate-ovate or oblong-ovate, chartaceous, 

with 3-5 main veins, the apex acute or acuminate, 

the base cordiform, the margins entire or weakly 

dentate or undulate; upper surface dull green, 

glabrous or puberulous; lower surface puberulous, 

densely covered with resinous dots; petioles 1.5- 

14 cm long, puberulous; pseudostipules forming 

a ring of lanceolate segments, early deciduous. 

Capitula numerous, sessile or short-pedunculate 

(4-5 mm long), arranged in dense, globose or 

corymbiform glomerules, terminal or at the end 

of short lateral branches; subinvolucral bract 

acicular, 2.5-4 mm long, puberulous; phyllaries 

oblong, 3-4 mm long, pilose, light green. Corollas 

white, infundibuliform, 2.5-3.5 mm long, with 

resinous dots at the apex. Achenes 3-3.5 mm long, 

black, cuneiform, with resinous dots; pappus of 

33-37 bristles, white, scabrous, ca. 3 mm long. 


November to March. 


Distribution: Of wide distribution, in disturbed 

areas at lower and middle elevations. Also in 

Jamaica, Martinique, northern South America, 

Peru, Bolivia, and Brazil. 

Public forests: Guánica, Maricao, Piñones, 

Río Abajo, and Susúa. 

Commentary: Mikania congesta is very 

similar to M. micrantha; nevertheless, it can be 

distinguished from M. micrantha by the 

subinvolucral bracts usually of the same length 

or longer than the phyllaries (vs. shorter) and by 

the capitula arranged in subglobose glomerules 

(vs. capitula in corymbs). 

2. Mikania cordifolia (L.f.) Willd., Sp. Pl. 3: 

1746. 1804. 

Fig. 35. E-H 

BASIONYM: Cacalia cordifolia L.f. 

SYNONYMS: Mikania gonoclada DC. 

Mikania convolvulacea DC. 

Guaco 


Stems pubescent, octagonal, striate. Leaves 

opposite; blades 7-12 × 3.5-7.4 (9) cm, ovate, 

slightly aromatic, chartaceous, the apex acute, the 

base cordiform or subcordiform, the venation with 

three main veins, the margins serrate, upper 

surface dull green, puberulous; lower surface 

glaucous, densely pubescent, with prominent 

venation; petioles 5-4 cm long, sulcate; 

pseudostipules divided in lanceolate segments, 

whitish. Capitula numerous, arranged in axillary 

corymbiform cymes; subinvolucral bract linear to 

lanceolate, 4-5 mm long, pilose; phyllaries 

elliptical or narrowly ovate, 6-8 mm long, green, 

pubescent; corollas white, infundibuliform, 

aromatic; anthers brown, connate, exserted. 

Achenes prismatic, with 5 longitudinal ribs, ca. 

3 mm long, grayish, scabrid; pappus of numerous 

white setae, scabrous, ca. 4.5 mm long. 




Distribution: In pastures or along rivers or 

roads, at middle and lower elevations. Also on 

Vieques and the Virgin Islands; throughout the 

Neotropics. 


Maricao, Río Abajo, Toro Negro, Tortuguero, and 

Vega. 

3. Mikania fragilis Urb., Symb. Antill. 1: 464. 

1899. 

Fig. 36. A-D 

SYNONYMS: Mikania fragilis Urb. var. leptodon Urb. 

Mikania rotunda sensu A. Stahl, non Griseb. 

Guaco redondo* 


Stems angular, with numerous lateral branches, 

pubescent when young, grayish with numerous 

lenticels when mature, the inner bark reddish, 

aromatic. Leaves opposite; blade 5-11 × 4-10 cm, 

broadly ovate, fragile, fleshy, slightly aromatic, 

not glandular, the apex acute, obtuse, or rounded, 

the base cordiform, the margins serrate, revolute; 

upper surface dark green, shiny, with the reticulate 

venation sunken, scabrid; lower surface pale 

green, dull, sometimes with some resinous dots, 

venation prominent; petioles thick, 2-4 cm long, 

pubescent, sulcate; pseudostipules broadly ovate,


102 



reflexed, 8-10 mm wide. Capitula few, 

pedunculate, in axillary corymbiform cymes; 

bracts of the involucre lanceolate to narrowly 

ovate, ca. 12 mm long, green to whitish; corollas 

white or with a reddish or greenish tinge, 

infundibuliform; anthers violet, connate, exserted. 

Achenes cuneiform, ca. 3.5 mm long, with 5 

longitudinal ribs, glabrous; pappus of numerous 

bristles, 7-8 mm long. 

Phenology: Flowering from September to April. 

Status: Endemic to Puerto Rico, rather common. 

Distribution: In forests of the cordilleras, at 

middle and upper elevations. 



* Common name taken from the work of Dr. 

Agustín Stahl (1937). 

4. Mikania micrantha Kunth in Humb. Bonpl. 

& Kunth, Nov. Gen. Sp. (quarto ed.) 4: 134. 

1818. 

Fig. 36. E-H 

Guaco falso 

Herbaceous vine, twining, attaining 10 m or 

more in length. Stems pubescent, angular, striate. 

Leaves opposite; blades 6-13 × 3-8 cm, ovate, 

chartaceous, 3-5-nerved, the apex acuminate, the 

base cordiform, the margins dentate; upper surface 

slightly shiny, puberulous; lower surface 

puberulous, with the venation somewhat 

prominent; petioles sulcate, glabrous or 

puberulous, as long as or shorter than the blade; 

pseudostipules forming an interpetiolar ring of 

lanceolate segments, early deciduous. Capitula 

numerous, pedunculate, arranged in corymbiform 

or paniculiform cymes, terminal on short lateral 

branches; subinvolucral bracts subulate, 2-3 mm 

long, glabrous; phyllaries ovate-oblong or 

elliptical, green, 3-4 mm long; corolla white, 

infundibuliform, slightly fragrant, 2.5-3 mm long; 

anthers exserted. Achenes ca. 1.2 mm long, dark 

gray to black, pentagonal; pappus of numerous 

white bristles, 2-3 mm long. 


November to February. 


Distribution: In disturbed areas, such as 

pastures, along rivers, and marshy areas, at low 

elevation. Also throughout tropical America. 

Public forest: Maricao, Río Abajo, and Susúa. 

5. Mikania odoratissima Urb., Symb. Antill. 1: 

464. 1899. 

Fig. 37. A-C 


Stems slender, striate, puberulous, aromatic. 

Leaves opposite; blades 4-10 × 3-8.5 cm, 

chartaceous, aromatic, ovate, triangular, or 

trilobed, glabrous or puberulous, 3-nerved, the 

apex acute or acuminate, the base truncate to 

cordiform, the margins dentate; upper surface 

dark green, shiny, with the venation flat or slightly 

sunken; lower surface light green, dull, venation 

prominent; petioles 1.5-3.5 cm long, sulcate, 

puberulous; pseudostipules forming a ring of 

lanceolate segments, pubescent, deciduous. 

Capitula in groups of 3, pedunculate to subsessile, 

arranged in terminal corymbiform cymes; 

subinvolucral bract obovate to oblong, 3-4 mm 

long; phyllaries 7-9 mm long, oblong-elliptical, 

green, puberulous; corollas white or pale green, 

infundibuliform, ca. 6.5 mm long. Achenes 4-4.5 

mm long, prismatic, scabrid, cream-colored; 

pappus of numerous scabrous bristles, creamcolored 

or light brown, 6.5-7 mm long. 




Distribution: In moist forests, at middle and 

upper elevations, along the Cordillera Central. 


6. Mikania pachyphylla Urb., Symb. Antill. 1: 

463. 1899. 

Fig. 37. D-G 


Stems hexagonal, 8-10 mm in diameter, glabrous, 

with numerous lateral branches, the bark grayish, 

smooth, and aromatic. Leaves opposite; blades 

3.5-9 × 2-3 cm, ovate, fleshy, brittle, slightly 

aromatic, 3-nerved, the apex acuminate, the base 

obtuse or rounded, the margins serrate-mucronate; 

upper surface pale green, shiny, glabrous, foveate, 

with the venation sunken; lower surface pale 

green, dull, glabrous, with prominent venation 

and whitish; petioles 1-2 cm long, glabrous,

104 



yellowish; pseudostipules absent. Capitula 

numerous, sessile or subsessile, arranged in 

terminal or axillary paniculiform cymes; 

subinvolucral bracts ovate, involute, 0.6-0.7 mm 

long, with some scattered reddish hairs; phyllaries 

oblong, 3-4.5 mm long, puberulous. Corollas 

white, infundibuliform, 3-3.2 mm long, with 

scattered resinous dots. Achenes cuneiform, 2-2.3 

mm long, light brown; pappus of numerous 

cream-colored bristles, ca. 2.7 mm long. 

Phenology. Flowering and fruiting throughout 

the year, especially from April to July. 


Distribution: In forests at upper elevations 

along the Cordillera Central and the Sierra de 

Luquillo. 



7. Mikania porosa Urb., Symb. Antill. 1: 465. 

1899. 

Fig. 38. A-C 

SYNONYM: Mikania swartziana sensu A. Stahl, non Griseb. 

Guaco de cabra 

Herbaceous vine, twining, deciduous, 

attaining 8 m in length. Stems slender, 4-6angled, 

glabrous, aromatic, with numerous pores. 

Leaves opposite; blades 1.5-6 × 0.7-2.5 cm, ovate, 

oblong, lanceolate, or sometimes trilobed or 

trifoliolate, the apex acute or acuminate, the base 

rounded or slightly cuneate, the margins entire, 

revolute, the venation pinnate, with 2 basal veins 

united to the midvein; upper surface dull or 

slightly shiny, glabrous, with numerous pores or 

sparse resinous dots; lower surface pale, dull, with 

numerous pores or resinous dots, the venation 

slightly prominent; petioles 5-15 mm long; 

pseudostipules absent. Capitula sessile or 

subsessile in subglobose or spiciform units, 

arranged in terminal paniculiform cymes; 

subinvolucral bracts ovate, ca. 0.5 mm long, 

puberulous; phyllaries oblong or subelliptical, 2.5- 

2.7 mm long, puberulous. Corollas white, 

infundibuliform, ca. 2.8 mm long. Achenes 

7. PIPTOCARPHA 

cuneiform, prismatic, 1.7-2.2 mm long, light 

brown or cream-colored; pappus of numerous 

bristles, white or cream-colored, scabrous, 2-3 mm 

long. 


February and fruiting in January and February. 


Distribution: In moist forests, in the zone of 

mogotes and on serpentinitic soils. 


8. Mikania stevensiana Britton, Bull. Torrey Bot. 

Club 43: 458. 1916. 

Fig. 38. D-F 

Guaco 


Stems slender, striate or angular, glabrous or 

puberulous. Leaves opposite; blades 3-5 × 1.9-4 

cm, 3-lobed, triangular-ovate or less frequently 

5-lobed, coriaceous, brittle, aromatic, trinerved, 

the apex acuminate or acute, the base 

subcordiform, the margins dentate, revolute, 

ciliate; upper surface dark green, shiny, with short 

bulbous hairs that give it a rough texture, the 

venation sunken; lower surface slightly shiny, 

puberulous, with prominent venation; petioles ca. 

3 cm long, sulcate, puberulous; pseudostipules 

forming a ring of filiform segments, deciduous. 

Capitula 3, forming pedunculate cymes that form 

a terminal corymbiform cyme; subinvolucral 

bracts oblanceolate, puberulous, 3-3.5 mm long; 

phyllaries oblong, 6-7 mm long, green or with a 

reddish tinge. Corollas white, infundibuliform, ca. 

6.5 mm long; anthers inserted; stigmatic branches 

long-exserted. Achenes cuneiform, ca. 5 mm long; 

pappus whitish, ca. 7 mm long, scabrous. 

Phenology: Flowering in August and from 

December to February. 

Status: Endemic to Puerto Rico, known from 

very few localities. 

Distribution: Uncommon in moist forests on 

serpentine soils. 

Public forest: Maricao. 

Lianas or woody shrubs, erect or clambering, with stellate or scaly pubescence. Leaves alternate, 

petiolate; blades simple. Capitula homogamous, discoid, with 2-20 flowers, arranged in corymbiform,

106 



umbelliform, spicate, or paniculiform cymes; involucre campanulate or cylindrical-campanulate, with 

several series of phyllaries. Flowers bisexual; corollas actinomorphic, tubular, 5-lobed; stamens 5, 

connate, the anthers sagittate; ovary inferior, the style slender with 2 stigmatic branches. Achenes with 

10 ribs, glabrous, truncate; pappus composed exclusively of bristles, without scales. A genus of 40 

species, distributed in the Neotropics. 

1. Piptocarpha tetrantha Urb., Symb. Antill. 1: 

457. 1899. 

Fig. 39. A-C 

Liana or clambering shrub, 10 m or more in 

length. Stems pendulous, flexible, with numerous 

white scales when young, slightly compressed, 

striate and lenticellate when mature, attaining 2.5 

cm in diameter. Leaves alternate, distichous; 

blades 6-12 × 2-5 cm, oblong, oblanceolate, or 

elliptical, coriaceous, the apex acute or obtuse, 

the base unequal, obtuse-acute, the margins 

smooth or denticulate; upper surface light green, 

slightly shiny, with reticulate venation, covered 

with small scales; lower surface completely 

covered with white scales, venation prominent; 

petioles 1-1.6 cm long, sulcate, whitish. Capitula 

in cymes, forming a compound axillary or 

8. PIPTOCOMA 

terminal racemiform inflorescence; involucre 

tubular, 7-8 mm long; phyllaries in 6 series, green, 

pilose at the apex and ciliate at the margins, the 

series gradually increasing in size toward the 

interior. Corolla white, infundibuliform, 5-6 mm 

long. Achenes cuneiform, cream-colored, 4-4.5 

mm long; pappus of numerous cream-colored 

bristles, ca. 5.8 mm long. 

Phenology: Flowering and fruiting probably 

throughout the year, but with more intensity from 

January to March. 


Distribution: In forests and on forest margins 


Luquillo. 

Public forests: Carite, Guilarte, El Yunque, 


Woody shrubs, erect or clambering. Stems angular, tomentose, glabrescent. Leaves alternate, 

petiolate; blades simple, the lower surface with stellate trichomes. Capitula homogamous, discoid, 

with 4-12 flowers, clustered in short-pedunculate glomerules to form corymbiform cymes; involucre 

cylindrical; the phyllaries, in several series, graduated; paleas present or not. Flowers bisexual; corollas 

actinomorphic, infundibuliform, 5-lobed; stamens 5, the anthers connate, spurred; ovary inferior, the 

style filiform, with 2 stigmatic branches, abaxially hispidulous. Fruit a turbinate achene, 5-angled, 

glabrous; pappus composed of bristles and short, irregular scales. A genus of 18 species, distributed in 


1. Piptocoma acevedoi Pruski, Novon 6: 98. 

1996. 

Fig. 40. A-H 

Woody vine or clambering shrub, 3-7 m in 

length. Stems much branched, pendulous, 

tomentose. Leaves alternate; blades 2-9.5 × 1-4.3 

cm, elliptical to obovate, chartaceous, the apex 

acute, obtuse, or rounded, the base acute or 

attenuate, the margins entire or undulate; upper 

surface glandular-punctate; lower surface whitish, 

densely stellate-tomentose; petioles 5-6 mm long, 

slightly winged, whitish. Capitula 4-9, with 6-10 

flowers each, in glomerules arranged in terminal 

cymes; involucre cylindrical or campanulate, 5- 

6.5 mm high, the phyllaries imbricate, in 3-4 

graduated series, green, pubescent; paleas present 

or not. Corollas pale violet, tubular, 5-6-lobed, 5- 

6.3 mm long, with glandular papillae on the tube 

and the apex of the lobes; anthers exserted, creamcolored; 

style with two recurved stigmatic 

branches, abaxially hispidulous. Achene turbinate, 

with 10 slight ribs, glabrous, 2.5-4 m long, light 

brown; pappus biseriate, the outer series of minute 

irregular-dentiform scales, the inner series of 

barbate bristles, 2.5-3.5 mm long.

108 




October to January. 

Status: Endemic to Puerto Rico, extremely 

rare. 

9. PSEUDOGYNOXYS 

Distribution: Known from two localities in 

Puerto Rico, in forests on limestone or serpentine 

substrate. 

Public forests: Guajataca and Maricao. 

Twining, herbaceous vines. Stems striate, puberulous. Leaves alternate, petiolate; blades simple, 

entire or serrate. Capitula heterogamous, radiate, longpedunculate, with numerous flowers, solitary 

or in terminal cymes; involucre campanulate or hemispherical, canaliculate; phyllaries uniseriate, 

oblong. Disc flowers bisexual; corollas actinomorphic, yellow, long-tubular, 5-lobed; stamens 5, the 

anthers connate, sagittate; ovary inferior, the style filiform, with 2 hirsute stigmatic branches on the 

distal portion. Ray flowers pistillate; corollas zygomorphic, ligulate, the ligules orange. Fruit a cylindrical 

achene; pappus of numerous bristles, without scales. A genus of 14 species, distributed in the Neotropics. 

1. Pseudogynoxys chenopodioides (Kunth) 

Cabrera, Brittonia 7: 54. 1950. 

Fig. 39. D-G 

BASIONYM: Senecio chenopodioides Kunth 

SYNONYM: Senecio confusus Britten 


Stems striate, subcylindrical, glabrous or 

puberulous. Leaves alternate; blades 5-8 × 2.5-5 

cm, lanceolate, membranaceous, the apex 

acuminate or acute, the base truncate, obtuse, or 

slightly cordiform, frequently unequal, the 

margins dentate-mucronate; upper surface dull, 

glabrous; lower surface dull, glabrous, with 

prominent venation; petioles 1.5-2.2 cm long. 

Capitula 2-6, pedunculate, in terminal 

corymbiform cymes; peduncles 2-5 cm long, 

puberulous; involucre crateriform, ca. 6 mm long, 

10. SALMEA 

the phyllaries green, lanceolate, ca. 4 mm long. 

Disc flowers with yellow tubular corollas, 9-10 

mm long; stigmatic branches yellow. Ray flowers 

with orange corollas, ligulate, the ligule elliptical, 

retuse at the apex, 2-2.5 cm long. Achenes 

turbinate, hispidulous, ca. 4 mm long; pappus of 

numerous white bristles, 5-7 mm long, scabrous. 


January to April. 

Status: Naturalized exotic, uncommon. 

Distribution: Exotic species, cultivated for its 

showy flowers; naturalized in more or less moist 

areas in Puerto Rico. Native to Central America, 

cultivated and naturalized in the tropics and 

subtemperate climates. Also on St. Croix and St. 

Thomas. 

Public forests: El Yunque, Río Abajo, and 

Vega. 

Erect or clambering shrubs. Stems striate, cylindrical. Leaves opposite, petiolate; blades simple, 

entire or dentate. Capitula homogamous, discoid, with numerous flowers, in terminal, pedunculate 

cymes; phyllaries in 1-6 series; receptacle conical, paleate. Flowers bisexual, each subtended by a 

palea; corollas actinomorphic, tubular, 5-lobed; stamens 5, the anthers connate, sagittate; ovary inferior, 

the style filiform, with 2 papillose-hirtellous stigmatic branches. Fruit a compressed, cuneiform achene; 

pappus of 2 short bristles. A genus of 2-4 species, distributed in the Neotropics. 

1. Salmea scandens (L.) DC., Cat. Hort. Monspel. 

141. 1813. 

Fig. 41. A-C 

BASIONYM: Bidens scandens L. 

Bejuco de miel, Bejuco de muela 


striate, cylindrical, glabrous or puberulous. Leaves 

opposite; blades 4.5-9.5 × 1.7-4.2 cm, lanceolate, 


obtuse, attenuate, or rounded, the margins

110 



revolute, entire, denticulate to dentate; upper 

surface dull, glabrous; lower surface dull, 

glabrous, with prominent venation; petioles 6-17 

mm long, glabrous. Capitula numerous, of few 

flowers, in terminal, pedunculate, corymbiform 

cymes; peduncles 1-10 mm long, puberulous; 

phyllaries 2-4-seriate, green, the outer elliptical, 

ca. 2 mm long, the inner spathulate, ca. 5 mm 

long; corollas white, tubular, 2.5-3 mm long; 

anthers exserted. Achenes cuneiform, compressed, 

ca. 2 mm long, glabrous, strigose on the margins; 

pappus of 2 barbate bristles, 1.5-1.7 mm long. 

11. SPHAGNETICOLA 




Distribution: Uncommon in moist thickets and 

forests at lower and middle elevations; locally 

common in the area of mogotes. Also in the 

Greater Antilles, Trinidad, and continental 

tropical America. 

Public forests: Carite, El Yunque, and Río 

Abajo. 

Decumbent herbs, sometimes scandent. Stems with adventitious roots at the nodes. Leaves opposite, 

petiolate; blades simple and serrate. Capitula heterogamous, radiate, with numerous flowers, pedunculate, 

solitary or clustered in a secondary terminal inflorescence; phyllaries in 2-3 series, foliaceous; receptacle 

paleaceous. Disc flowers bisexual, subtended by a palea; corollas yellow or orange, actinomorphic, 

tubular, 5-lobed; stamens 5, the anthers black, connate; ovary inferior, the style filiform, with 2 papillosehirtellous 

stigmatic branches. Ray flowers pistillate; corollas zygomorphic, ligulate, yellow or orange. 

Fruit a tuberculate achene, black; pappus forming a fimbriate “corona,” minute. A genus of 4 species, 

3 of them native to the New World. 

1. Sphagneticola trilobata (L.) Pruski in Acev.- 

Rodr., Mem. New York Bot. Gard. 78: 114. 

1996. 

Fig. 41. D-I 

BASIONYM: Silphium trilobatum L. 

SYNONYM: Wedelia trilobata (L.) Hitchc. 

Wedelia, Wild marigold 

Creeping or decumbent herb, sometimes 

scandent, attaining 2 m in length. Stems obtusely 

quadrangular or cylindrical, puberulous. Leaves 

opposite; blades 3-10 × 2.5-6 cm, oblanceolate or 

rhombic, usually 3-lobed, chartaceous, the apex 

obtuse or acute, the base obtuse or attenuate, the 

margins serrate; upper surface pubescent or 

puberulous; lower surface pubescent or 

puberulous, with glandular dots, with prominent 

venation and pubescent; petioles 0-5 mm long. 

Capitula of numerous flowers, solitary, terminal; 

peduncles 3.5-14 cm long, puberulous; involucre 

turbinate; phyllaries in one series, 12-14, unequal, 

10-14 cm long, oblanceolate or oblong, green; disc 

corollas tubular, yellow, 4.5-5.5 mm long, 

papillose on the outer surface of the lobes; ray 

corollas ligulate, the ligules ca. 15 mm long, 

subelliptical, trilobed at the apex. Achenes 

pyriform, ca. 2 mm long; pappus forming a 

corona, ca. 1 mm high. 



Status: Ornamental, cultivated and 

naturalized, very common. 

Distribution: Cultivated in gardens and parks 

in Puerto Rico and the Virgin Islands. Native to 

the New World, but naturalized throughout the 

tropics and subtropics. 

Public forests: El Yunque, Maricao, and Río 

Abajo. 

References: Holmes, W. C. 1993. The genus Mikania (Compositae: Eupatorieae) in the Greater 

Antilles. Sida Bot. Misc. 9: 1-69. Keeley, S. C. 1978. A revision of the West Indian Vernonias 

(Compositae). J. Arnold Arb. 59: 360-413. Pruski, J. F. 1996. Compositae of the Guayana highland-X. 

Reduction of Pollalesta to Piptocoma (Vernoniae: Piptocarphinae) and consequent nomenclatural 

adjustments. Novon 6: 96-102. Turner, B. L. (1993) Berylsimpsonia (Asteraceae: Mutisieae), a new 

genus of the Greater Antilles. Phytologia 74: 349-355.

112 



7. Family BASELLACEAE 


1a. Corolla with petals connate only at the base, neither accrescent nor fleshy; stamens inserted at the 

base of the corolla ............................................................................................................. 1. Anredera 

1b. Corolla urceolate, tubular, 5-lobed, accrescent, fleshy, surrounding the fruit; stamens inserted on 

the upper portion of the corolla tube .................................................................................. 2. Basella 

1. ANREDERA 

Herbaceous vines, clambering or twining. Stems usually fleshy. Leaves alternate, slightly succulent, 

petiolate; blades simple; stipules absent. Flowers actinomorphic, bisexual or functionally unisexual, 

produced in pendulous, axillary or terminal racemes. Calyx of 2 free sepals, much shorter than the 

corolla, united to the petals at the base; corolla not accrescent, with 5 petals connate at the base; 

stamens 5, the filaments free or connate and united to the petals, the anthers oblong, dehiscent by 

longitudinal sutures; ovary superior, of 3 carpels, the styles 3, free or connate at the base, as long as the 

ovary. Fruit a utricle. A New World genus of about 12 species. 

Key to the species of Anredera 

1a. Pedicellar glands persistent; petals 2-3 mm long, turning blackish on drying; stigmatic branches as 

long as the style ........................................................................................................... 1. A. cordifolia 

1b. Pedicellar glands deciduous; petals 1.5-2.2 mm long, turning cream-colored on drying; stigmatic 

branches 2 times as long as the style ........................................................................... 2. A. vesicaria 

1. Anredera cordifolia (Ten.) Steenis, Fl. Males., 

Ser. 1, 5: 303. 1957. 

Fig. 42. H-J 

BASIONYM: Boussingaultia cordifolia Ten. 

Madeira vine 

Herbaceous vine, fleshy, twining, attaining 5 

m in length. Roots tuberous. Stems glabrous, 

cylindrical, up to 2.5 cm in diameter, producing 

solitary or clustered bulbils. Leaves alternate; 

blades 2.2-10 × 1.4-6.2 cm, ovate or broadly ovate, 

glabrous, fleshy, the apex obtuse or acute, the base 

cordiform, decurrent on the petiole, the margins 

entire, hyaline or reddish; upper surface dull, 

waxy, with the venation somewhat sunken; lower 

surface lighter than the upper surface, with the 

venation slightly prominent; petioles 6-12 mm 

long, slightly winged. Flowers aromatic, turning 

blackish on drying, apparently bisexual, in 

terminal or axillary racemes or pendulous 

panicles, up to 65 cm long; pedicels with 

persistent bracts. Calyx of 2 white sepals, ovate 

or broadly elliptical, 1.3-2.2 mm long; corolla of 

5 white petals, ovate or elliptical, 2-3 mm long, 

connate at the base; stamens 5, the filaments 

connate at the base, 2-3.5 mm long, the anthers 

lanceolate, 0.7-0.8 mm long; ovary globose or 

obovoid, glabrous, stigmatic branches 3, as long 

as the style. Utricles globose, slightly compressed 

or triangular, 0.9-1.1 mm long, crowned by the 

base of the fleshy style. 


September. Rarely produces fruits in Puerto Rico; 

propagated by means of bulbils or underground 

tubers. 

Status: Exotic, cultivated as an ornamental, 

naturalized, uncommon. 

Distribution: In disturbed areas, at middle and 

lower elevations. Native to the New World, but 

extensively cultivated and naturalized throughout 

the tropics and subtemperate zones. 

Public forests: Guánica and Susúa. 

Commentary: This species has been identified 

erroneously as Anredera baselloides (Kunth) 

Baill. by many botanists of the Caribbean flora. 

Anredera baselloides is a different species, which

114 


is neither found either in Puerto Rico or the Virgin 

Islands. 

2. Anredera vesicaria (Lam.) C.F. Gaertn, Suppl. 

Carp. 176, t. 213. 1807. 

Fig. 42. A-G 

BASIONYM: Basella vesicaria Lam. 

SYNONYMS: Boussingaultia leptostachys Moq. 

Anredera leptostachys (Moq.) Steenis 

Suelda con suelda, Consuelda, Madera vine 


length. Roots tuberous. Stems slender, glabrous, 

cylindrical, sometimes with a reddish tonality. 

Leaves alternate; blades 3-11 × 2-6.5 cm, ovate, 

glabrous, fleshy, the apex acute or acuminate, the 

base rounded, obtuse, or cordiform, the margins 

entire or crenate; upper surface dull, with 

subconspicuous venation; lower surface slightly 

shiny, with prominent venation; petioles 0.4-2 cm 

long, slightly winged. Flowers aromatic, 

functionally unisexual, in axillary pendulous 

racemes, 10-20 cm long, or in terminal pendulous 

panicles, 30-50 cm long; pedicels with deciduous 

1. Basella alba L., Sp. Pl. 272. 1753. 

Fig. 42. K-N 

SYNONYM: Basella rubra L. 

Espinaca de Nueva Zelandia, Acelga trepadora 

Twining vine, climbing or creeping, 

herbaceous, glabrous, attaining 10 m in length. 

Stems somewhat succulent, cylindrical, usually 

with a reddish to purple tonality, up to 2.5 cm in 

diameter. Leaves alternate; blades 3-15 × 1.2-14 

cm, green, reddish, or purple, ovate, orbicular, or 

oblong, fleshy, the apex obtuse, rounded, or acute, 

the base acute, subtruncate, rounded, or cordiform, 

2. BASELLA 

bracts. Calyx white, of ovate sepals, ca. 0.4 mm 

long; corolla white, the petals ovate, 1.5-2.2 mm 

long, turning cream-colored or remaining white 

on drying; stamens 5, the filaments connate at 

the base, 3-4 mm long, the anthers lanceolate, 

0.2-0.4 mm long; ovary glabrous, globose, 

stigmatic branches twice as long as the style. 

Utricles obovoid, slightly triangular, 1-1.3 mm 

long, crowned by the fleshy style. 

Phenology: Flowering from September to 

April. The populations of this species in 

Hispaniola, Puerto Rico, the Virgin Islands, and 

the Lesser Antilles contain staminate flowers only 

and never produce fruits. The reproduction of this 

species on the islands mentioned seems to be 

exclusively asexual by means of underground 

tubers (Sperling, 1987). 


rather common. 

Distribution: In disturbed areas, at middle and 

lower elevations. Also on Cayo Santiago, Vieques, 

St. Croix, St. John, St. Thomas, and Tortola; from 

the southern United States to northern South 

America, including the Antilles. 

Public forests: Susúa and Tortuguero. 

Herbaceous, succulent, twining vines or decumbent or creeping herbs. Stems fleshy, much branched. 

Leaves alternate, succulent, petiolate; blades simple; stipules absent. Flowers bisexual, produced in 

axillary, simple or branched spikes. Flowers actinomorphic, bisexual, without aroma. Calyx of 2 white 

or pale green sepals, free to the base, much shorter than the corolla, united to the petals at the base; 

corolla tubular, urceolate, 5-lobed, succulent; stamens 5, with short filaments, inserted on the upper 

portion of the corolla tube, the anthers dehiscent by longitudinal sutures; ovary superior, globose or 

pyriform, of 3 carpels, uniovulate, the styles 3, free to the base. Fruit a utricle, completely covered by 

the persistent or accrescent corolla. A genus of 5 species native to Africa or Madagascar. 

decurrent on the petiole, the margins entire, 

undulate, usually reddish or purple; upper surface 

dull, with slightly conspicuous venation; lower 

surface dull, with slightly prominent venation; 

petioles green or reddish, 0.5-5 cm long, slightly 

winged. Flowers arranged in simple or branched 

spikes, axillary, erect, 15-25 cm long. Calyx white 

or with a reddish tinge, the sepals 2, ovate or 

elliptical, 3.2-4 mm long; corolla fleshy, white, 

with a reddish or violet tinge toward the apex, 

3.3-4.1 mm long. Utricles purple, fleshy, 4-lobate, 

4-5 mm long, covered by the accrescent corolla, 

5-7 mm long.


116 



August to March. 

Status: Exotic, widely cultivated as a 

vegetable, because its leaves are edible, like 

spinach. 

Distribution: Probably native to Africa, but 

widely cultivated and naturalized throughout the 

tropics. 

Reference: Sperling, C. R. 1987. Systematics of the Basellaceae. Doctoral thesis, Harvard University. 

8. Family BIGNONIACEAE 


1a. Leaves imparipinnate, without tendrils ............................................................................................ 2 

1b. Leaves 2- or 3-foliolate, usually with a terminal tendril .................................................................. 4 

2a. Corolla violet-pink; stamens inserted; calyx > 10 mm long ........................................ 8. Podranea 

2b. Corolla orange or reddish orange; stamens exserted; calyx < 8 mm long ............... 11. Tecomaria 

4a. Tendrils bifid or trifid ........................................................................................................................ 5 

4b. Tendrils simple, spiral or spiral-shaped, sometimes absent ............................................................. 9 

5a. Branches acutely hexagonal, with obtuse ribs .................................................................................. 6 

5b. Branches cylindrical or quadrangular ............................................................................................... 7 

6a. Leaflets cordiform or truncate at the base; calyx double, the interior margin short, the exterior 

margin undulate, elongate; corolla white with a purple tinge ...............................1. Amphilophium 

6b. Leaflets rounded at the base; calyx simple; corolla orange.......................................... 9. Pyrostegia 

7a. Tendrils trifid, forming a small claw (harpidium); corolla yellow ........................... 5. Macfadyena 

7b. Tendrils bifid or trifid, not forming a claw; corolla white, lilac, or purple ..................................... 8 

8a. Branches quadrangular; interpetiolar zones not glandular; leaves without odor; tendrils trifid, with 

one division deciduous and two developing into an adventitious disc; corolla white, with the tube 

yellow inside ....................................................................................................................... 4. Distictis 

8b. Branches cylindrical; interpetiolar zones glandular; leaves usually with a strong garlic odor; tendrils 

trifid, each division simple, perennial, without an adventitious disc; corollas lilac or 

purple...................................................................................................... 6. Mansoa 

9a. Corolla bilabiate, white, ca. 4 mm long ....................................................................... 12. Tynanthus 

9b. Corolla with 5 lobes of similar size, lilac, violet, or purple, > 1.5 cm long .................................. 10 

10a. Stems lepidote, with pseudostipules; leaflets cuneate at the base, upper and lower surface lepidote, 

with domatia in the axils of the basal secondary veins; corolla 8-9 cm long ................10. Saritaea 

10b. Stems glabrous, without pseudostipules or these inconspicuous; leaflets truncate, rounded, or 

subcordiform at the base; upper surface glabrous, lower surface glabrous or lepidote, without domatia; 

corolla < 8 cm long .......................................................................................................................... 11


11a. Calyx 2.1-4.1 cm long, in the form of a spathe, open distally along 2/3 of its length; corolla lilac 

with the lobes purple and the center white or purple ............................................7. Phryganocydia 

11b. Calyx 3-10 mm long, crateriform, truncate or denticulate at the apex; corolla violet or lilac with 

the center yellow .............................................................................................................................. 12 

12a. Corolla infundibuliform, 1.5-3 cm long, violet; mature stems quadrangular, sulcate; cross section 

of the stem with 4 dark arms formed by the phloem tissue; foliage turning reddish on drying 

............................................................................................................................. 2. Arrabidaea 

12b. Corolla tubular-campanulate, 2.5-7.5 cm long, lilac with the center yellow; mature stems cylindrical; 

cross section of the stem with 8 dark arms formed by the phloem tissue; foliage remaining green or 

turning brown on drying ..................................................................................................... 3. Cydista 

1. AMPHILOPHIUM 

Lianas. Stems hexagonal with conspicuous obtuse ribs; cross section of the mature stem with a 

cross of 4 (-5) arms formed by the phloem tissue; interpetiolar zones not glandular; pseudostipules 

foliaceous, persistent or deciduous. Leaves usually 3-foliolate, with the terminal leaflet replaced by a 

trifid tendril. Flowers produced in terminal panicles, usually on short axillary branches. Calyx 

campanulate, double, with the inner margin thick, short and the outer undulate; corolla purple, tubular, 

bilabiate; stamens 4, didynamous, the anthers glabrous; ovary superior, ovate-cylindrical, with 2 locules. 

Fruit an oblong-elliptical capsule, the valves parallel to the septum; seeds with 2 hyaline wings not 

very differentiated from the body. A neotropical genus of 7 species distributed from Mexico to northern 

Argentina, including the Antilles. 

1. Amphilophium paniculatum (L.) Kunth in 

Humb. Bonpl. & Kunth, Nov. Gen. Sp. 

(quarto ed.) 3: 149. 1819. 

Fig. 3. C, 43. A-C 

BASIONYM: Bignonia paniculata L. 

Liana that climbs by tendrils and attains 5-10 

m in length. Stems up to 10 cm in diameter, the 

bark fibrous, vertically striate; cross section of the 

mature stem cylindrical, with a cross of 4(-5) dark 

arms, formed by the phloem tissue; branches 

acutely hexagonal, with ribbed margins, lepidote 

or with dendroid pubescence; pseudostipules 

foliaceous, 3-10 mm long, early deciduous. Leaves 

2-foliolate, with a terminal tendril, filiform, trifid, 

early deciduous; leaflets 2.5-16 × 2.1-10.7 cm, 

ovate to suborbicular, membranaceous, the apex 

acuminate or obtuse, the base truncate or 

asymmetrically cordiform, the margins entire; 

upper surface with minute, discoid, peltate scales, 

sometimes pubescent; lower surface with minute 

discoid, peltate scales, with the veins pubescent; 

petioles and petiolules hexagonal, with lepidote 

or dendroid pubescence; petiole 2.3-7.1 cm long; 

petiolules 0.6-4.3 cm long. Flowers fragrant, 

produced in terminal panicles, with deciduous 

bracts, the axes lepidote or with dendroid 

pubescence. Calyx double, the interior margin 2or 

3-labiate, short, the outer 5-labiate, undulate, 

7-12 mm long, densely lepidote; corolla white 

with a purple tinge, tubular, bilabiate, 2.4-3.5 cm 

long, the tube split for 1/3 - 2/3 of its length, the 

2 upper lobes united, ca. 1 mm long, the 3 lower 

lobes united, 1.3-1.8 cm long; stamens inserted; 

ovary ovate-cylindrical, 2-3 mm long, with an 

annular-pulvinate disc at the base. Capsules 

elliptical, compressed, 3.9-9.5 cm long, verrucoselenticellate, 

lepidote; seeds numerous, 1-1.9 cm 

wide, with a membranaceous wing on each side. 

Phenology: Collected in flower from August 

to October. 


Distribution: In secondary forests and thickets, 

at middle to upper elevations, especially in the 

zone of mogotes. From Mexico to northern 

Argentina, including the Antilles. 

Public forest: Río Abajo.

118 


2. ARRABIDAEA 

Lianas or less frequently small trees or shrubs. Stems quadrangular or cylindrical; cross section of 

the mature stem with a cross of 4 arms formed by phloem tissue; interpetiolar zones usually glandular; 

pseudostipules small and inconspicuous. Leaves opposite, usually 3-foliolate, with the terminal leaflet 

replaced by a long, filiform, spiral-shaped tendril. Flowers produced in long axillary or terminal panicles. 

Calyx crateriform, simple, truncate, bilabiate or minutely 5-denticulate at the apex; corolla reddish, 

pink, or violet, hypocrateriform, infundibuliform, or campanulate, pubescent outside; stamens 4, 

didynamous; ovary superior, with 2 locules and a crateriform disc at the base. Fruit a linear capsule, 

compressed, with the valves parallel to the septum; seeds with 2 hyaline wings. A neotropical genus of 

about 70 species. 

1. Arrabidaea chica (Humb. & Bonpl.) Verl., 

Rev. Hort. 40: 154. 1868. 

Fig. 43. D-G 

BASIONYM: Bignonia chica Humb. & Bonpl. 

SYNONYM: Adenocalymna portoricensis A. Stahl 

Liana that climbs by tendrils and attains 15 

m in length. Stems quadrangular, striate, glabrous, 

4-sulcate, up to 6 cm in diameter; cross section of 

the mature stem with a cross of 4 dark arms, 

formed by the phloem tissue. Pseudostipules 

inconspicuous. Leaves turning reddish on drying, 

2-3-foliolate, the terminal leaflet replaced by a 

simple, deciduous, filiform, spiral-shaped tendril; 

lateral leaflets 3.4-11 × 1.5-5 cm, ovate, 

chartaceous, glabrous or with some scattered 

hairs, the apex acuminate or acute, the base 

truncate, rounded, or cordiform, the margins 

entire; lower surface with a prominent midvein; 

petioles 1.5-7 cm long, petiolules 0.3-4.5 cm long, 

both cylindrical and turning blackish on drying. 

3. CYDISTA 

Flowers fragrant, produced in terminal panicles. 

Calyx simple, 3-5 mm long, puberulent, 

crateriform, truncate or minutely denticulate; 

corolla violet, zygomorphic, infundibuliform, 1.5- 

3 cm long, with the lobes rounded; stamens and 

staminodia inserted, borne near the base of the 

corolla; ovary cylindrical, lepidote, with a small 

annular disc at the base. Capsules linear or 

narrowly elliptical, 12-23 cm long, smooth, 

glabrous, semi-woody, acute or obtuse at both 

ends; seeds numerous, 7-9 mm long, the wings 

lateral. 

Phenology: Collected in fruit in March. 

Status: Native, rare, probably extirpated in 

Puerto Rico, since it has not been collected there 

in more than a century. 


two collections made in 1885 in Bayamón, Stahl 

79 and Sintenis 1096. Also on St. John; 

throughout the Neotropics. 

Lianas with tendrils. Stems cylindrical or quadrangular; cross section of the mature stem with a 

cross of 8 dark arms formed by the phloem tissue; interpetiolar zone not glandular; pseudostipules 

present or absent. Leaves opposite, simple or 2-foliolate, with a filiform, terminal tendril. Flowers in 

terminal or axillary racemes or panicles. Calyx crateriform, simple, truncate or slightly bilobate at the 

apex; corolla white, pink, lilac, or purple, tubular or infundibuliform; stamens 4, didynamous; ovary 

superior, cylindrical, with 2 locules, without a disc. Fruit a linear capsule, compressed, with the valves 

parallel to the septum; seeds compressed, with 2 brown wings. A neotropical genus of 6 species. 

1. Cydista aequinoctialis (L.) Miers, Proc. Roy. 

Hort. Soc. London 3: 191. 1863. 

Fig. 3. B; 44. A-D 

BASIONYM: Bignonia aequinoctialis L. 

Bejuco blanco, Liana de la sierra, 

Guard wiss 

Liana that climbs by tendrils and attains 15 

m in length. Stems cylindrical, up to 6 cm in 

diameter, the bark grayish, vertically striate; cross


120 


section of the mature stem with a cross of 8 dark, 

marginal arms, formed by the phloem tissue; 

branches quadrangular, glabrous, pseudostipules 

absent. Leaves opposite, 2-foliolate, with a 

terminal tendril, simple, filiform, early deciduous; 

lateral leaflets 5.5-16 × 2.5-9 cm, ovate, oblong, 

or elliptical, chartaceous, the apex obtuse, acute, 

or acuminate, the base obtuse to subcordiform, 

the margins entire; upper surface glabrous; lower 

surface with punctiform scales, without domatia; 

petioles and petiolules quadrangular, pubescent 

or glabrous, the petiole 0.9-4.5 cm long, the 

petiolules 0.8-4 cm long. Flowers fragrant, 

produced in axillary or terminal panicles. Calyx 

yellowish, crateriform, simple, 4-10 mm long, 

pubescent or with punctiform scales, the apex 

truncate or denticulate; corolla zygomorphic, 

1. Distictis lactiflora (Vahl) DC., Prodr. 9: 191. 

1845. 

Fig. 3. A; 44. E-H 

BASIONYM: Bignonia lactiflora Vahl 

SYNONYM: Macrodiscus lactiflorus (Vahl) Bureau 

Pega palo, Liana fragante, Viuda 

Woody vine that climbs by tendrils with 

adhesive discs and attains 5 m or more in length. 

Mature stems cylindrical with the bark light 

brown, shredded; cross section with peripheral 

phloem forming a cross of 4 arms. Branches 

quadrangular, striate, puberulous, with thickened 

nodes; pseudostipules foliaceous, 3-4 mm long, 

deciduous. Leaves opposite, 2-foliolate, with a 

terminal tendril with 3 branches, 2 of them 

modified into an adhesive disc; leaflets 2.7-8 × 

2-4.5 cm, oblong, elliptical to subrounded, 

coriaceous, with the reticulate venation 

conspicuous, the apex obtuse, acute, or acuminate, 

4. DISTICTIS 

tubular-campanulate, 2.5-7.5 cm long, the tube 

slightly compressed, narrow at the base, white, 

yellow inside, the lobes rounded, lilac, the throat 

yellow with violet lines; stamens inserted; ovary 

cylindrical, 2-3 mm long, without a disc at the 

base. Capsules linear, 21-43 × 1.5-2.4 cm, 

prominent on the margins; seeds numerous, 

semicircular, membranaceous, 1.5-3.8 cm long, 

with two membranaceous wings. 

Phenology: Flowering throughout the year. 


Distribution: In river-bank forests, on the 

edges of mangrove swamps, and in coastal forests. 

Also on St. Croix, St. John, and St. Thomas; Cuba, 

Hispaniola, the Lesser Antilles, and continental 



Lianas with tendrils. Stems hexagonal or quadrangular, cross section of the mature stem with 

peripheral phloem forming a cross of 4 arms, or cylindrical and not forming a cross; interpetiolar zone 

not glandular; pseudostipules present. Leaves opposite, 2-foliolate, frequently with a terminal trifid 

tendril. Flowers in terminal racemes or panicles. Calyx crateriform, simple, more or less truncate, 

usually glandular near the apex; corolla white, cream-colored, or red-violet, tubular-campanulate or 

broadly tubular, pubescent outside; stamens 4, didynamous; ovary superior, oblong, with 2 locules and 

an annular-patelliform disc at the base. Fruit a convex or biconvex capsule, elliptic-oblong, with the 

valves parallel to the septum; seeds compressed, with elongate irregular protuberances, in 2 rows. A 

genus of 6 species distributed from Mexico to Brazil, including the Antilles. 

the base rounded or acute, unequal; upper surface 

shiny, glabrous, puberulous, or minutely lepidote; 

lower surface pale green, dull, covered with 

minute transparent scales, venation prominent; 

petioles 1-2 cm long, the petiolules 5-10 mm long, 

both with 2 rows of crimped hairs. Flowers 

fragrant, in pendulous, terminal racemes or 

panicles, 10-30 cm long; peduncles and pedicels 

pubescent. Calyx simple, green, crateriform, 5-7 

mm long, truncate at the apex or minutely 

denticulate; corolla white, infundibuliform, 2.5- 

5.5 cm long, the tube yellow inside, the lobes 5, 

rounded, ca. 1.5 cm long. Capsule woody, 

elliptical, 6.5-13 × 1.7-3.5 cm, somewhat 

compressed or subcylindrical, with the apex and 

base acute; seeds numerous, compressed, ca. 2.5 

cm long, the central circular body ca. 7 mm in 

diameter, with a surrounding marginal hyaline 

wing.


122 



throughout the year, especially from November 

to July. 

Status: Native, very common in southwestern 


5. MACFADYENA 

Lianas with tendrils. Stems cylindrical or quadrangular; cross section of the mature stem with 

multilobed xylem, the lobes alternating with radially arranged phloem tissue; glandular interpetiolar 

zone present or absent; pseudostipules small, lanceolate to ovate. Leaves opposite, 2-foliolate, with a 

terminal trifid tendril in the form of a claw. Flowers in axillary cymes or panicles. Calyx campanulate, 

simple; corolla yellow, zygomorphic, tubular or campanulate; stamens 4, didynamous; ovary superior, 

linear-cylindrical, with 2 locules and an annular disc at the base. Fruit a narrow capsule, elongate and 

compressed, with the valves parallel to the septum; seeds numerous, with 2 wings. A genus of 4 species 

of neotropical distribution. 

1. Macfadyena unguis-cati (L.) A. H. Gentry, 

Brittonia 25: 236. 1973. 

Fig. 3. D; 45. A-C 

BASIONYM: Bignonia unguis-cati L. 

SYNONYMS: Batocydia unguis (L.) DC. 

Doxantha unguis-cati (L.) Miers 

Uña de gato, Liana uñada, Cat-claw 

Woody vine, which climbs by tendrils and 

attains 10-15 m in length. Stems cylindrical, 

lenticellate, up to 6 cm in diameter; cross section 

of the mature stem with multilobed xylem, the 

lobes alternating with radially arranged phloem 

tissue; nodes thickened; interpetiolar zone not 

glandular. Pseudostipules ovate, ca. 5 mm long. 

Leaves opposite, 2-foliolate, with a terminal 

tendril, trifid like a claw, generally of short 

duration; leaflets 6-16 × 1.2-7 cm, elliptical, 

oblong, or obovate, chartaceous or coriaceous, 

glabrous or with punctiform scales, the apex acute 

or acuminate, the base acute, rounded, or unequal, 

the margins undulate or rarely denticulate; upper 

surface dark, shiny, with sunken venation; lower 

surface light green, dull, with prominent venation; 

6. MANSOA 

Distribution: In forests and coastal thickets. 

Also on Cayo Luis Peña, Vieques, St. Croix, and 

St. Thomas; Cuba and Hispaniola. 

Public forests: Cambalache, Ceiba, Guajataca, 

Guánica, Río Abajo, Susúa, and Vega. 

petioles 1-4.5 cm long, petiolules 0.5-2.5 cm long, 

both glabrous. Flowers solitary or in pairs, 

axillary; pedicel ca. 2 cm long. Calyx green, 

campanulate, 12-16 mm long, with 5 unequal 

lobes; corolla brilliant yellow, infundibuliform, 

4-8 cm long, the limb 3-6 cm in diameter, with 5 

unequal lobes, rounded; stamens 4, didynamous, 

inserted; ovary covered with punctiform scales. 

Capsule linear, somewhat woody, brown, 25-95 

cm long; seeds numerous, 1-3.5 cm long, with 2 

membranaceous wings. 





lower elevations, from the coast to the Cordillera 

Central. Also on Culebra, Desecheo, Vieques, St. 


Gorda; throughout the Antilles and tropical and 

subtropical America. 

Public forests: Cambalache, Ceiba, Desecheo, 

El Yunque, Guánica, Maricao, Río Abajo, Susúa, 

Toro Negro, and Vega. 

Lianas with tendrils. Stems cylindrical or subquadrangular; cross section of the mature stem with a 

cross of 4 or 8 dark arms formed by the phloem tissue; interpetiolar zone glandular or not; pseudostipules 

short, conical-obtuse. Leaves opposite, 2- or 3-foliolate, with a terminal trifid or simple tendril; petioles 

sometimes with an apical glandular zone. Flowers in terminal panicles or racemes or in axillary corymbs. 

Calyx crateriform, tubular, or campanulate, simple, truncate or 5-dentate; corolla white, lilac, purple, 

or reddish, zygomorphic, tubular-campanulate or infundibuliform; stamens 4, didynamous; ovary


superior, cylindrical, glandular-papillose, puberulent, with 2 locules, the ovules 2(3-4) seriate. Fruit a 

linear-oblong capsule, compressed or convex, sometimes tuberculate, with the valves parallel to the 

septum; seeds numerous, with 2 membranaceous wings or without wings. A genus of 15 species 

distributed from Mexico to Brazil. 

Key to the species of Mansoa 

1a. Leaflets 9-30 cm long; lower surface with a group of minute punctiform glands in the axils of the 

secondary veins (frequently of the basal and sub-basal veins); corolla 7.5-9 cm long; fruit 30-35 cm 

long, woody, swollen, with a very prominent rib along each valve ............................ 1. M. alliacea 

1b. Leaflets 6-10 cm long; lower surface without glands in the axils of the secondary veins; corolla 4.5- 

6.5 cm long; fruits 15-25 cm long, coriaceous, compressed, with a slight rib along each valve 

.......................................................................................................................2. M. hymenaea 

1. Mansoa alliacea (Lam.) A. H. Gentry, Ann. 


Fig. 45. H 

BASIONYM: Bignonia alliacea Lam. 

SYNONYM: Adenocalymna obovatum Urb. 

Bejuco de ajo, Mata de ajo 

Woody vine, which climbs by tendrils and 

attains 10-15 m in length. Young stems 

quadrangular, with the angles ribbed; mature 

stems cylindrical or subcylindrical, lenticellate; 

cross section with a cross of 4 arms, formed by 

the phloem tissue; nodes slightly compressed with 

a glandular interpetiolar zone; pseudostipules 

ovate, ca. 5 mm long. Leaves opposite, 2-foliolate, 

sometimes with a terminal tendril, trifid, 

deciduous, 20-25 cm long; leaflets 9-30 × 4.5-18 

cm, elliptical or broadly elliptical, chartaceous, 

the apex obtuse or obtusely acuminate, the base 

acute, rounded, or obtuse and sometimes unequal, 

the margins undulate or crenate; upper surface 

dull, glabrous, with the venation slightly 

prominent; lower surface light green, dull, 

minutely lepidote, with the venation slightly 

prominent, the axils of the secondary veins with 

a group of minute punctiform glands; petioles and 

petiolules glabrous, the petioles 7-30 mm long, 

the petiolules with two keels formed by the 

decurrent base of the blade, 7-30 mm long. 

Flowers few, in axillary racemes; pedicels 7-15 

mm long. Calyx green, campanulate or broadly 

campanulate, simple, puberulous, 5-8 mm long, 

truncate, slightly crenate at the apex; corolla 

violet-pink or lavender, infundibuliform, 7.5-9 cm 

long, the limb 4-5 cm in diameter, with five 

rounded lobes; stamens inserted; ovary 

cylindrical, lepidote. Capsule oblong, woody, light 

brown, 30-35 × 2.5- 3 cm, swollen, with a 

longitudinal rib on the middle portion of each of 

the valves; seeds asymmetrically ovate, 

compressed, thick, woody, 1.5-2 cm long. 

Phenology: Collected in flower from October 

to May and in fruit in October. 

Status: Exotic, cultivated in Puerto Rico. 

Distribution: Cited by Britton and P. Wilson 

(1925) for the Agricultural Experiment Station 

in Trujillo Alto. Species native to central South 

America, from the Amazonian regions of Peru 

and Brazil, also in Guyana. 

2. Mansoa hymenaea (DC.) A. H. Gentry, Ann. 


Fig. 45. D-G 

BASIONYM: Bignonia hymenaea DC. 

Bejuco de ajo, Mata de ajo 

Woody vine, with a strong odor of garlic, 

which climbs by tendrils, 3-5 m in length. Young 

stems angular to subcylindrical, striate, 

puberulous to glabrous; mature stems cylindrical, 

lenticellate; cross section with a cross of four arms, 

formed by the phloem tissue; nodes slightly 

compressed, with a glandular interpetiolar zone; 

pseudostipules keeled, 2-3 mm long. Leaves 

opposite, 2-foliolate, with a terminal trifid tendril, 

early deciduous; leaflets 6-10 × 2.5-6 cm, elliptical 

or ovate, chartaceous, the apex acute, obtuse, or 

acuminate, the base acute, obtuse, or rounded, 

sometimes unequal, the margins undulate; upper 

surface dull, glabrous, with the venation slightly 

prominent; lower surface light green, dull, 

sparsely punctate-glandular, without glands in the 

axils of the secondary veins, the venation slightly

124 


prominent; petioles and petiolules puberulous; 

petioles 1.3-3.5 cm long, the petiolules slightly 

keeled by the decurrent base of the blade, 1-2.6 

cm long. Flowers few in axillary racemes, 5-20 

cm long; peduncles keeled. Calyx green, 

campanulate, puberulous, 8-9 mm long, truncate 

or denticulate, ciliate and with a purple tinge at 

the apex; corolla violet-pink or lavender, 

infundibuliform, 4.5-6.5 cm long, the tube lighter 

than the lobes, the limb 2.5-4 cm in diameter, 

with 5 rounded lobes; stamens inserted; ovary 

cylindrical, lepidote. Capsule linear, long- 

7. PHRYGANOCYDIA 

acuminate, compressed, coriaceous, light brown, 

15-25 × 1.7-2 cm, with a longitudinal vein in the 

middle portion of each of the valves; seeds 

numerous, 2-winged, oblong, membranaceous, ca. 

3.3 cm long. 

Phenology: Flowering from December to May 

and fruiting from January to May. 


uncommon. 

Distribution: Along roads. Species native to 

the Neotropics, distributed from Mexico to 

southeastern Brazil. 

Lianas with tendrils. Stems cylindrical, glabrous or lepidote; cross section of the mature stem with 

phloem forming a cross of 8 (4) arms; interpetiolar zone not glandular; pseudostipules inconspicuous 

or absent. Leaves opposite, simple or 2-foliolate, usually with a terminal simple tendril. Flowers few, 

in terminal or axillary panicles, scarcely branched. Calyx simple, in the form of a spathe, split along 

T‡of its length; corolla lilac to purple, infundibuliform, lepidote outside; stamens 4, didynamous, the 

anthers glabrous; ovary superior, conical or cylindrical, lepidote, with 2 locules, the ovules 2-seriate 

per locule; disc absent. Fruit a linear capsule, oblong or ovoid (slightly compressed), coriaceous, with 

valves parallel to the septum; seeds numerous, with or without 2 membranaceous wings. A genus of 3 

species of tropical America, distributed from Costa Rica to Bolivia. 

1. Phryganocydia corymbosa (Vent) Bureau ex 

K.Schum. in Engl. & Prantl, Nat. 

Pflanzenfam. 4, Abt. 3b: 224, fig. 89H. 1894. 

Fig. 46. A-C 

BASIONYM: Spathodea corymbosa Vent. 

Liana that climbs by tendrils, attaining 10 m 

in length. Stems subcylindrical, finely striate, 

glabrous, up to 5 cm in diameter; cross section of 

the mature stem with a cross of 8 arms formed by 

the phloem tissue. Leaves opposite, 2-foliolate and 

sometimes with a terminal, simple tendril, 7-16 

cm long; leaflets 4-22 × 1.9-11 cm, elliptical or 

ovate-elliptical, chartaceous or coriaceous, 

glabrous, the apex acute, obtuse, or acuminate, 

the base obtuse or rounded, usually trinerved, the 

margins undulate; upper surface dark, shiny, with 

the venation somewhat sunken; lower surface 

slightly shiny, punctate, with prominent venation; 

petioles 0.6-3.3 cm long, the petiolules 0.5-3 cm 

long, slightly winged, both finely lepidote; 

pseudostipules absent or inconspicuous and early 

deciduous. Flowers in terminal or axillary 

panicles, the axes finely lepidote and somewhat 

compressed. Calyx green, in the form of a spathe, 

split along 2/3 of its length, 2.1-4.1 cm long; 

corolla lilac with the center white or purple, 

infundibuliform, 4.6-9.5 cm long, the limb 4-8 

cm in diameter, the lobes purple, crenate; stamens 

4, didynamous, inserted; ovary cylindrical, 

covered with punctiform scales. Capsule linearoblong, 

subwoody, grayish, 13-53 cm long, 

densely lepidote; seeds numerous, 1.5-2 cm long, 

with a membranaceous wing, not differentiated 

from the body of the seed 


Status: Exotic, cultivated, uncommon. 

Distribution: Native to continental tropical 

America, from Costa Rica to Bolivia. Cultivated 

on St. Croix and St. Thomas.


126 


8. PODRANEA 

Lianas or climbing shrubs. Stems cylindrical; cross section of the mature stem with normal peripheral 

phloem; interpetiolar zone not glandular; pseudostipules absent. Leaves opposite, pinnately compound, 

imparipinnate, without tendrils. Flowers in terminal panicles. Calyx simple, campanulate, 5-dentate; 

corolla zygomorphic, violet-pink, tubular-campanulate; stamens 4, didynamous; ovary superior, linear, 

tetragonal, glabrous, with 2 locules and an annular disc at the base, the ovules 6-seriate per locule. 

Fruit a linear capsule, coriaceous, with valves perpendicular to the septum; seeds numerous, with 

membranaceous wings at the ends. A genus of 1 or 2 species of tropical eastern and southern Africa. 

1. Podranea ricasoliana (Tanfani) Sprague in 

Thiselton-Dyer, Fl. Cap. 4 (2): 450. 1904. 

Fig. 46. D-G 

BASIONYM: Tecoma ricasoliana Tanfani 

Liana or clambering shrub, 3-4 m in length. 

Stems cylindrical, smooth; cross section with 

peripheral phloem not forming a cross. Leaves 

opposite, imparipinnate, 7-9-foliolate, without 

tendrils; leaflets 2.5-3.8 × 1.5-2.0 cm, ovate, 

chartaceous, glabrous, the apex obtuse or shortacuminate, 

the base truncate, cuneate, or 

attenuate, the margins serrate; upper surface dull, 

with the venation flat, dark; lower surface light 

green, punctate, dull, with prominent venation; 

rachis glabrous, marginate, petioles and petiolules 

glabrous, the petioles 2.5-5 cm long, the petiolules 

4-8 mm long; pseudostipules absent. Flowers in 

terminal panicles; pedicels 1.5-2 cm long. Calyx 

9. PYROSTEGIA 

green with a pink tinge, campanulate, 1.5-2 cm 

long, with 5 apiculate lobes; corolla violet-pink, 

campanulate, 5-7 cm long, the tube white with 

reddish purple lines, the limb ca. 5 cm in diameter, 

with 5 rounded lobes, unequal, stamens inserted; 

ovary glabrous. Capsule linear, coriaceous, 

cylindrical, 25-30 cm long; seeds numerous, ca. 

1.5 cm long, brown, with 2 marginal wings, ca. 5 

mm long. 


to May and in fruit in February. 

Status: Exotic, cultivated and sometimes 

naturalized. 

Distribution: Native to South Africa, but 

widely cultivated throughout the tropics. 

Cultivated along the Cordillera Central. 

Cultivated and possibly naturalized on St. Croix, 

St. John, and St. Thomas. 

Lianas with tendrils. Stems obtusely hexagonal or subangular with 6-8 prominences; cross section 

of the mature stem with normal peripheral phloem; interpetiolar zone not glandular; pseudostipules 

inconspicuous. Leaves opposite, 2-3-foliolate, with a terminal trifid tendril. Flowers in terminal or 

axillary panicles. Calyx simple, crateriform, truncate or 5-denticulate, more or less glandular-lepidote; 

corolla red-orange, zygomorphic, narrowly tubular-infundibuliform; stamens 4, didynamous, exserted; 

ovary superior, bicarpellate, linear-tetragonal, lepidote, with an annular-pulvinate disc at the base. 

Fruit a linear capsule, compressed, smooth, with the valves parallel to the septum; seeds numerous, 

with 2 brown wings with hyaline margins. A genus of 3 or 4 species naturally distributed from southern 

Colombia to Brazil and Paraguay. 

1. Pyrostegia venusta (Ker Gawl.) Miers, Proc. 

Roy. Hort. Soc. 3: 188. 1863. 

Fig. 47. A-E 

BASIONYM: Bignonia venusta Ker Gawl. 

SYNONYM: Pyrostegia ignea (Vell.) K. Presl 

Liana that climbs by tendrils, 5-7 m in length. 

Stems angular to subcylindrical, striate, 

puberulent or lepidote, interpetiolar zone not 

glandular; cross section of the mature stem with 

peripheral phloem tissue not forming a cross. 

Leaves opposite, 2- or 3-foliolate, sometimes with


128 


a trifid, deciduous tendril; leaflets 4-6 × 2.5-4 cm, 

ovate, chartaceous, lepidote or puberulent, with 

numerous resinous dots on both surfaces, the apex 

acute, the base rounded, the margins undulate, 

revolute; upper surface dark, slightly shiny, with 

the venation slightly prominent; lower surface 

light green, dull, with a prominent midvein, 

without domatia; petioles and petiolules glabrous 

or puberulous, the petioles 1.5-2.5 cm long, 

angular, the petiolules 1-1.5 cm long, sulcate on 

the upper surface; pseudostipules inconspicuous. 

Flowers numerous in terminal panicles; pedicels 

0.6-15 cm long. Calyx green, broadly 

campanulate, 4-4.5 mm long, subtruncate or 5denticulate 

and ciliate at the apex; corolla redorange, 

tubular, 5.5-6.5 cm long, the tube glabrous 

10. SARITAEA 

externally, puberulent inside, the limb 2.9-4 cm 

in diameter, with 5 lobes, 1-1.5 cm long; stamens 

exserted, yellow; ovary oblong, ca. 4 mm long, 

glabrous or lepidote. Capsule linear, compressed, 

coriaceous, light brown, 25-30 cm long, with the 

midvein slightly prominent; seeds numerous, 1.2- 

1.4 cm long, the wings brown with the margins 

hyaline. 

Phenology: Collected in flower in January and 

February. 

Status: Exotic, cultivated, common. 

Distribution: Native to southern Brazil, 

Paraguay, and northern Argentina; cultivated 

throughout the tropics. Widely cultivated along 

the Cordillera Central. 

A monospecific genus native to Colombia and Ecuador, characterized by the following species. 

1. Saritaea magnifica (W. Bull) Dugand, 

Caldasia 3: 263. 1945. 

Fig. 47. F-H 

BASIONYM: Bignonia magnifica W. Bull 

Liana that climbs by tendrils, 3-7 m in length. 

Stems cylindrical, lepidote, compressed at the 

nodes, interpetiolar zone not glandular; cross 

section of the mature stem normal. Leaves 

opposite, 2-foliolate, sometimes with a simple 

tendril, of short duration; leaflets 4.2-11.5 × 3.1- 

6.4 cm, obovate, chartaceous, with the venation 

slightly prominent on both surfaces, the apex 

obtuse, the base cuneate or decurrent, the margins 

entire; upper surface dull, minutely lepidote; lower 

surface dull, sparsely lepidote, with domatia in 

the axils of the basal secondary veins; petioles 

and petiolules lepidote, the petioles 1.7-2.8 cm 

long, the petiolules 0.3-1.6 cm long; 

pseudostipules foliaceous, 0.6-4.2 cm long. 

11. TECOMARIA 

Flowers few, in corymbiform panicles, usually 

terminal; pedicels 4-6 mm long. Calyx yellowish 

green, simple, crateriform, 7-8 mm long, truncate, 

lepidote; corolla purple-pink, tubularcampanulate, 

8-9 cm long, glabrous or minutely 

glandular, the tube pubescent inside, the throat 

white inside with purple-pink lines, the lobes 

unequal, 2.2-3.1 cm long; stamens 4, didynamous, 

inserted; ovary linear, ca. 4 mm long, glandularlepidote, 

with two locules, the ovules in 2 series 

per locule; disc hypocrateriform-pulviniform, 1 

mm high. Capsule linear, compressed, coriaceous, 

brown, 20-25 × 1-1.2 cm; seeds numerous, oblong, 

2-winged, the hyaline wings membranaceous. 


September to December. 

Status: Exotic, cultivated, uncommon in 


Distribution: Native to Colombia and Ecuador 

but cultivated throughout the tropics. 

Lianas or climbing shrubs. Stems cylindrical, interpetiolar zone not glandular; pseudostipules absent; 

cross section of the mature stem with normal peripheral phloem. Leaves opposite, pinnately compound, 

imparipinnate, without tendrils. Flowers in axillary racemes. Calyx simple, campanulate, 5-dentate; 

corolla orange or bright red, zygomorphic, tubular, curved; stamens 4, of the same length, exserted;


130 


ovary superior, oblong, glabrous, with 2 locules. 

Fruit a linear capsule, coriaceous, with valves 

perpendicular to the septum; seeds numerous, flat, 

with 2 hyaline-membranaceous wings. A genus 

of 2 species of tropical eastern and southern 

Africa. 

1. Tecomaria capensis (Thunb.) Spach, Hist. Nat. 

Veg. Phan. 9: 137. 1840. 

Fig. 48. A-C 

BASIONYM: Bignonia capensis Thunb. 

SYNONYM: Tecoma capensis (Thunb.) Lindl. 

Clambering or semi-erect shrub, 3-4 m in 

length. Stems cylindrical, lenticellate, puberulous; 

cross section of the mature stem with peripheral 

phloem not forming a cross. Leaves opposite, 

imparipinnate, 7-11-foliolate, without tendrils; 

leaflets 1.5-4.2 × 1-3 cm, elliptical to subrounded, 

membranaceous, sessile, puberulent, the apex 

rounded, the base rounded or abruptly cuneate, 

the margins serrate; upper surface dull, pale, with 

12. TYNANTHUS 

slightly prominent venation; lower surface light 

green, dull, punctate, with slightly prominent 

venation, forming a conspicuous network, with 

tufts of hairs in the axils; petioles 1.5-2.5 cm long; 

pseudostipules absent. Flowers numerous, in 

axillary racemes; pedicel 6-10 mm long. Calyx 

green, crateriform, 5-7 mm long, 5-dentate, 

ciliate, puberulent; corolla orange or reddish 

orange, tubular, curved, 3.5-5 cm long, with 5 

oblong, unequal lobes, the 2 upper lobes smaller 

than the 3 lower; stamens 4, of equal length, 

exserted; ovary superior, oblong, glabrous, with 

a crateriform disc at the base. Capsule linear, 5- 

11 cm long and 7-8 mm wide; seeds in 2 rows, 

slender, 2-winged, the wings hyalinemembranaceous. 


and March. 


Distribution: Cultivated in gardens along the 

Cordillera Central, also on St. Croix and St. 

Thomas. Native to South Africa, but widely 

cultivated throughout the tropics. 

Small trees or lianas with tendrils. Stems cylindrical or occasionally quadrangular, interpetiolar 

zone not glandular; pseudostipules absent or foliaceous but early deciduous; cross section of the mature 

stem with a cross of 4 arms formed by the phloem tissue. Leaves opposite, 2- or 3-foliolate, the terminal 

leaflet usually replaced by a terminal, simple or trifid tendril. Flowers numerous, in terminal or axillary 

panicles. Calyx simple, crateriform, subtruncate, usually 5-denticulate; corolla white, zygomorphic, 

infundibuliform, bilabiate, divided up to half its length, pubescent outside; stamens 4, didynamous, 

subexserted; ovary superior, bicarpellate, conical, densely pubescent, the ovules in 2-4-series per locule. 

Fruit a linear capsule, compressed, smooth, with the valves parallel to the septum; seeds numerous, 

flat, with 2 hyaline-membranaceous wings. A genus of 14 species of tropical America distributed from 

southern Mexico to Brazil and Bolivia. 

1. Tynanthus polyanthus (Bur.) Sandw., Kew 

Bull. 1953: 465. 1954. 

Fig. 48. D-I 

BASIONYM: Schizopsis polyanthus Bur. 

SYNONYMS: Bignonia caryophyllea Bello 

Tynanthus caryophylleus (Bello) Alain 

Bejuco de clavo 

Liana that climbs by tendrils, 10-20 m in 

length. Stems more or less quadrangular, 

tomentose, interpetiolar zone not glandular, 

usually with an interpetiolar line, attaining 3 cm 

or more in diameter; cross section of the mature 

stem with a cross of 4 arms formed by the phloem 

tissue, with a strong odor of cloves; pseudostipules 

foliaceous, ca. 1 cm long, deciduous. Leaves 

opposite, 2-foliolate, sometimes with a simple 

tendril, in the form of a spiral, up to 15 cm long; 

leaflets 6-13 × 3.5-10 cm, broadly ovate to 

subcircular, membranaceous, the apex cuspidate 

or short-acuminate, the base truncate, rounded, 

or subcordiform, the margins entire, undulate or 

crenulate; upper surface with the venation flat, 

glabrous; lower surface puberulent, especially on


132 


the veins; petioles and petiolules puberulent or 

short-pubescent, the petioles 1.7-4 cm long, deeply 

canaliculate, the petiolules 1.2-3 (-5) cm long. 

Flowers numerous in terminal or axillary panicles, 

bracteate, the axes pubescent; pedicels ca. 1 mm 

long. Calyx green, crateriform, 1-1.5 mm long, 

subtruncate, minutely 5-denticulate, puberulent; 

corolla white, tubular, bilabiate, ca. 4 mm long, 

puberulent outside, the 2 upper lobes smaller than 

the lower, the 3 lower lobes 2-3 mm long; stamens 

didynamous, inserted; ovary conical, ca. 0.8 mm 

long, densely appressed- pubescent; annular disc 

pubescent. Capsule linear, compressed, somewhat 

woody, dark brown, 15-27 × 0.5-0.7 cm, with the 

midvein inconspicuous; seeds numerous, brown, 

4-5 mm long, the wings hyaline-membranaceous. 

Phenology: Collected in flower in July and in 

fruit in April. 


Distribution: Native to western Amazonia, 

sporadically cultivated in Puerto Rico, Jamaica, 

Cuba, and the Dominican Republic. 

References: Gentry, A. H. 1982. Bignoniaceae. Flora de Venezuela. Ediciones Fundación. Venezuela. 

Hawk, W. D. 1997. A review of the genus Cydista. Ann. Missouri Bot. Gard. 84: 815-840. 

9. Family BORAGINACEAE 


1a. Corolla campanulate; style bifid, each branch again bifid, for a total of 4 stigmas .......... 1. Cordia 

1b. Corolla infundibuliform; style simple, bilobed .......................................................... 2. Tournefortia 

1. CORDIA 

Shrubs or trees and sometimes scandent shrubs. Leaves alternate, petiolate; blades simple; stipules 

absent. Flowers bisexual, usually heterostylous, in axillary or terminal cymes, panicles, corymbs, or 

spikes. Calyx campanulate or tubular, 5-dentate or 5-10 lobed, usually striate and accrescent in the 

fruit; corolla actinomorphic, campanulate or hypocrateriform; stamens 5, inserted on the corolla tube, 

shorter or longer than the corolla tube; ovary superior, bicarpellate, each carpel with 2 uniovulate 

chambers, the style branched into 4 stigmatic branches. Fruit a globose, fleshy drupe, with 1-4 pyrenes. 

A genus of about 300 species of pantropical distribution. 

Key to the species of Cordia 

1a. Inflorescences of unbranched glomerules; lobes of the corolla almost as long as the tube .............. 

........................................................................................................................................ 1. C. bellonis 

1b. Inflorescences of paniculate or corymbose heads; lobes of the corolla much shorter than the tube 

........................................................................................................................ 2. C. polycephala 

1. Cordia bellonis Urb., Symb. Antill. 1: 393. 

1899. 

Fig. 49. A-F 

SYNONYM: Varronia bellonis (Urb.) Britton 

Clambering or recumbent shrub, 2-3 m in 

length. Stems appressed-pubescent when young, 

glabrous when mature. Leaves alternate; blades 

2-6 × 1-2 cm, elliptical, oblong to sublanceolate, 

chartaceous, the apex acute, the base acute or 

obtuse, the margins dentate or denticulate, 

especially on the distal portion; upper surface with 

short appressed hairs that give it a scabrid texture, 

the midvein slightly prominent; lower surface pale


green, strigulose-puberulous, with the venation 

slightly prominent; petioles appressed-pubescent, 

4-7 mm long. Flowers few, sessile, arranged in 

short axillary cymes; peduncle < 6 mm long. 

Calyx green, turbinate, strigulose, 2-2.5 mm long; 

corolla white, infundibuliform, glabrous, ca. 4 mm 

long, the lobes ovate to oblong, almost as long as 

the tube; stamens and stigmas inserted. Drupe red, 

ovoid, ca. 3 mm in diameter, with the persistent 

accrescent calyx covering the lower 1/3 of the 

fruit. 


to January and in fruit from November to January. 

Status: Endemic, listed in the Federal Register 

of the United States as an endangered species. 

Locally common at the base of mogotes. 

Distribution: Along the banks of rivers and 

streams or in the interior of forests on substrates 

of limestone (mogotes) or serpentine. 

Public forests: Maricao, Río Abajo, and Susúa. 

2. Cordia polycephala (Lam.) I.M. Johnst., J. 

Arnold Arbor. 16: 33. 1935. 

Fig. 49. G-L 

BASIONYM: Varronia polycephala Lam. 

SYNONYM: Varronia corymbosa (L.) Desv. 

Basora, Palo de perico, Saragüero, Black sage 

Woody shrub, clambering, up to 5 m in length. 

Stems pubescent when young, cylindrical, 

attaining 1.5 cm in diameter. Leaves alternate; 

blades 4-15 × 2-5.5 cm, ovate, lanceolate, or 

2. TOURNEFORTIA 

elliptical, coriaceous, the apex acute or acuminate, 

the base obtuse or rounded, sometimes unequal, 

the margins serrate; upper surface scabrous, the 

venation sunken; lower surface pale green, more 

or less pubescent, with prominent reticulate 

venation; petioles pubescent, 5-10 mm long. 

Flowers numerous, sessile, arranged in terminal 

or axillary paniculate or corymbose heads. Calyx 

yellowish green, crateriform, pubescent, 2.5-3 mm 

long; corolla white, campanulate, 4-5 mm long, 

the lobes much shorter than the tube, the tube 

pubescent inside; stamens and stigmas inserted. 

Drupe red, globose, 3-4 mm in diameter, 

completely covered by the accrescent calyx. 


the year, particularly in June and July. 


Distribution: In weedy areas and along paths, 

at lower and middle elevations. Also on Vieques, 

St. Croix, St. Thomas, Tortola, and Virgin Gorda; 

Hispaniola, the Lesser Antilles, and northern 


Public forests: Cambalache, Carite, Ceiba, El 

Yunque, Guajataca, Guánica, Maricao, Río Abajo, 

Susúa, Toro Negro, and Tortuguero. 

Commentary: Cordia wagnerorum Howard 

was originally described as a clambering shrub; 

nevertheless, the clambering habit seems to be 

the exception to the rule, since I have only 

observed this species as an erect shrub. For this 

reason, this species has been excluded from the 

present work. 

Vines, lianas, shrubs, or trees. Leaves alternate, petiolate; blades simple; stipules absent. Flowers 

actinomorphic, bisexual, in terminal scorpioid cymes. Calyx crateriform, with 5 deep lobes; corolla 

hypocrateriform, the tube slightly broadened at the base, the lobes reflexed; stamens 5, inserted on the 

corolla tube, shorter than the corolla; ovary superior, with 2 carpels, each carpel with 2 uniovulate 

chambers, the style simple, bilobed. Fruit a fleshy drupe, globose or 2- or 4- lobed, with 1-4 pyrenes. A 

genus of about 150 species of pantropical distribution. 

Key to the species of Tournefortia 

1a. Leaves rigid-coriaceous, scabrous on the upper surface ................................................. 5. T. scabra 

1b. Leaves chartaceous or coriaceous, not rigid, pubescent, hirsute or glabrous on the upper surface, 

not scabrous ........................................................................................................................................ 2 

2a. Twining vines; fruits white ................................................................................................................ 3 

2b. Clambering subshrubs; fruits orange .......................................................................... 3. T. maculata

134 


3a. Fruits completely white. .................................................................................................................... 4 

3b. Fruits with 2-4 circular black spots. ................................................................................................. 5 

4a. Stems, leaves, and inflorescences hirsute or tomentose ........................................ 2. T. hirsutissima 

4b. Stems strigose, becoming glabrous when mature; leaves glabrous or puberulous on the venation; 

inflorescences strigose .....................................................................................................1. T. bicolor 

5a. Leaves 1.5-3 (3.5) cm long, the apex acute or obtuse ............................................ 4. T. microphylla 

5b. Leaves (3.5) 4-10 cm long, the apex acute or acuminate ............................................ 6. T. volubilis


1. Tournefortia bicolor Sw., Prodr. 40. 1788. 

Fig. 50. A-E 

SYNONYM: Tournefortia laevigata Lam. 


Stems cylindrical, strigose, glabrous when mature, 

attaining 4-5 cm in diameter. Leaves alternate; 

blades 4-15 × 2.5-8 cm, lanceolate or elliptical, 

chartaceous, the apex acute or short-acuminate, 

the base cuneate or obtuse, the margins entire or 

crenulate; upper surface glabrous or appressedpubescent 

or scabrid, puberulous on the venation, 

lower surface glabrous, with prominent 

puberulous venation; petioles sulcate, strigulose, 

5-25 mm long. Flowers numerous, sessile, in 

scorpioid cymes, branched, the axes strigose. 

Calyx green, crateriform, strigose, the sepals 

subulate, 1.5-2 mm long; corolla white, 

hypocrateriform, 4-6 mm long, strigose outside, 

the lobes ovate, spreading, ca. 1 mm long; stamens 

inserted; ovary ovoid, with 4 chambers, the stigma 

sessile. Drupe globose, white, ca. 8 mm in 

diameter, separating into 4 pyrenes on drying. 


February to August. 


Distribution: In moist forests and along stream 

banks. Also on St. John and St. Thomas; 

throughout tropical America. 

Public forests: El Yunque, Río Abajo, and Toro 

Negro. 

2. Tournefortia hirsutissima L., Sp. Pl. 140. 

1753. 

Fig. 50. F-J 

Nigua, Chiggernit, Chiggernit grape, Giniper 

Twining liana or climbing shrub, 6-10 m in 

length. Stems woody, up to 2 cm in diameter, the 

bark somewhat corky, cream-colored, usually with 

persistent petioles; branches fragile, cylindrical, 

hirsute or tomentose, usually hollow in the center 

on drying. Leaves alternate; blades 12-22 × 2-11 

cm, elliptical, oblong, or sometimes obovate, 


acute, the margins entire, ciliate; upper surface 

dark, dull, hirsute, with the venation sunken; 

lower surface pale green, dull, hirsute, with 

prominent venation; petioles cylindrical, hispid, 

10-12 mm long. Flowers numerous, fragrant, 

subsessile, in branched scorpioid cymes, 

corymbiform, terminal. Calyx green, crateriform, 

pubescent, of 5 lanceolate sepals, ca. 3 mm long; 

corolla white, hypocrateriform, 4-5 mm long, 

strigose outside. Drupe white, subglobose, 5-6 mm 

in diameter. 


November to September. 


Distribution: In pastures or areas of disturbed 

vegetation, in moist zones at middle and lower 

elevations. Also on Mona, Vieques, St. Croix, St. 

John, St. Thomas, and Tortola; the Greater and 

Lesser Antilles, Central America, South America, 

and the United States (Florida). 


Mona, Piñones, Río Abajo, Toro Negro, 

Tortuguero, and Vega. 

3. Tournefortia maculata Jacq., Enum. Syst. Pl. 

14. 1760. 

Fig. 51. A-F 

SYNONYM: Tournefortia laurifolia Vent. 

Tournefortia peruviana Poir. 

Bejuco de masa 

Vine or clambering or arcuate subshrub, 2-5 

m in length. Stems cylindrical, glabrous, not 

lenticellate. Leaves alternate; blades 5-12 (16) × 

2.8-8 cm, elliptical, oblong, ovate, broadly ovate, 

or lanceolate, chartaceous or membranaceous, the 

apex acute, the base rounded, acute, or sometimes 

unequal, the margins entire; upper surface 

glabrous, shiny, with a prominent midvein; lower 

surface glabrous, shiny with the venation slightly 

prominent; petioles 1-1.8 cm long. Flowers 

numerous, pedicellate, in cymes with scorpioid 

branches, terminal, with puberulous axes. Calyx 

green, crateriform, 2-3 mm long, the sepals ovate 

or lanceolate, puberulous; corolla yellow, 

infundibuliform, 5-6 mm long, puberulous or 

glabrescent outside, minutely pubescent-papillose 

inside, the lobes subulate, long-acuminate, slightly 

reflexed; stamens inserted, the filaments very 

short, the anthers apiculate at the apex. Drupes 

depressed-globose, 2-4-lobed, ca. 8 mm in 

diameter, orange, sometimes with 2 or 4 circular 

black spots. 

Phenology: Flowering from February to 

August and fruiting from March to August. 

Status: Native, locally common.

136 



Distribution: In moist forests on substrates of 

limestone or serpentine. Also in Jamaica and 

Hispaniola. 

Public forests: Guajataca, Guilarte, Maricao, 


Commentary: The populations of this species 

in Puerto Rico and the Dominican Republic have 

been considered as a distinct species (T. laurifolia) 

based on the relative length of the corolla lobes. 

Nevertheless, this character is very variable across 

the range of distribution of T. maculata, and 

therefore the recognition of these populations at 

the species level is not justified.

138 


4. Tournefortia microphylla Bertero ex Spreng., 

Syst. Veg. 1: 644. 1824. 

Fig. 52. A-D 

SYNONYM: Tournefortia volubilis L. var. microphylla 

(Bertero ex Spreng.) A. DC. 

Twining vine, 2-5 m in length. Stems 

cylindrical, slender, < 8 mm in diameter, 

pubescent or glabrescent, lenticellate when 

mature. Leaves alternate; blades 1.5-3.5 × 0.5- 

1.5 cm, elliptical, ovate, or lanceolate, 

chartaceous, the apex acute or obtuse, usually 

mucronate, the base obtuse or rounded, usually 

unequal, the margins entire; upper surface 

puberulous, with the venation flat; lower surface 

puberulous, with the venation slightly prominent; 

petioles cylindrical, glabrous, ca. 4 mm long. 

Flowers numerous, pedicellate, in scorpioid 

cymes, branched, terminal. Calyx green, 

crateriform or urceolate, pubescent, 1-1.5 mm 

long, the sepals subulate, 0.8-1 mm long; corolla 

white, hypocrateriform, 2-3 mm long, the tube 

narrow in the middle, strigulose outside, the lobes 

linear to subulate, 1-1.5 mm long; stamens 

inserted, the anthers sessile. Drupes ovoid, globose 

or 4-lobed, ca. 4 mm in diameter, white, with 2-4 

circular black spots. 

Phenology: Flowering and fruiting apparently 



Distribution: In thickets and dry forests at 

lower elevations. Also on Culebra, Culebrita, 

Desecheo, Mona, Vieques, Anegada, St. Croix, 

St. John, St. Thomas, Tortola, and Virgin Gorda; 

Hispaniola and the Lesser Antilles. 

Public forests: Boquerón, Guánica, and Mona. 

5. Tournefortia scabra Lam., Ill. 1: 417. 1791. 

Fig. 52. E-G 

Erect to scandent shrub, 2-5 m in length. 

Stems cylindrical, slender, < 5 mm in diameter, 

strigose when young, striate and glabrescent when 

mature. Leaves alternate; blades 2-5 × 0.5-1.2(1.7) 

cm, linear, oblong, or narrowly lanceolate, rigidcoriaceous, 

the apex acute, obtuse, rounded, or 

retuse, the base obtuse, rounded, or subcordiform, 

the margins revolute, undulate; upper surface 

scabrous, with the venation sunken; lower surface 

scabrous-pubescent, with the reticulate venation 

prominent; petioles strigose, 1.5-5 mm long. 

Flowers few, sessile, in scorpioid cymes, scarcely 

branched, terminal, 5 cm or less long, with 

strigose axes. Calyx green, crateriform, strigulose, 

the sepals ovate, ca. 2 mm long; corolla white or 

yellowish, urceolate-tubular, ca. 2.5 mm long, the 

tube strigose on the distal portion, the lobes ovate, 

very short. Drupes subovoid, 5-6 mm in diameter, 

white. 



Status: Native; uncommon. 

Distribution: In thickets and dry forests at 

lower elevations in southern and northwestern 

Puerto Rico. Also in Cuba and Hispaniola. 

Public forests: Guajataca and Guánica. 

6. Tournefortia volubilis L., Sp. Pl. 140. 1753. 

Fig. 52. H-K 

SYNONYM: Tournefortia ferruginea Lam. 

Twining vine, 2-7 m in length. Stems 

cylindrical, slender, ferruginous-pubescent to 

glabrous, lineate-lenticellate when mature. Leaves 

alternate; blades 3.5-10 × 0.6-3.5 cm, lanceolate, 

ovate, or less frequently oblong, chartaceous, the 

apex acuminate or less frequently acute, the base 

rounded, subcordiform, or less frequently obtuse, 

sometimes unequal, the margins entire; upper 

surface pubescent or glabrous, sometimes 

pustulose, with the venation slightly sunken; 

lower surface pubescent or less frequently 

glabrous, pale, with prominent venation; petioles 

2-7 mm long. Flowers numerous, sessile or 

subsessile, in terminal branched scorpioid cymes. 

Calyx green, crateriform, the sepals subulate, 1- 

2 mm long, strigose; corolla greenish white or 

greenish yellow, hypocrateriform, ca. 2 mm long, 

the tube pubescent outside, the lobes subulate, 1- 

1.5 mm long; stamens inserted, the anthers sessile. 

Drupes ovoid, usually 2-4-lobed, ca. 3 mm in 

diameter, white, with 2-4 circular black spots. 




Distribution: In coastal thickets and dry 

forests, in southern Puerto Rico, at lower and 

middle elevations. Also on Vieques, St. Croix, and 

Virgin Gorda; throughout tropical America, 


Public forest: Guánica.


140 


Commentary: This species can be 

distinguished with certainty from T. microphylla 

by the presence of at least some leaves with 

acuminate apices (some individuals present leaves 

with acute apices, but leaves with acuminate 

apices will always be found). Tournefortia 

volubilis, as a general rule, has the lower surface 

of the leaves with rather conspicuous venation, 

while T. microphylla, on the contrary, has 

inconspicuous venation.


10. Family BUDDLEJACEAE 

1. BUDDLEJA 

Erect, sometimes scandent shrubs, or less frequently trees, pubescent with simple, stellate trichomes 

or peltate scales. Leaves opposite or sometimes alternate, petiolate or sessile; blades simple, serrate or 

less frequently entire or lobed; stipules present or absent. Flowers usually functionally unisexual, 4merous, 

arranged in cymes that in turn form terminal or axillary heads or panicles. Calyx campanulate 

or tubular, of short sepals; corolla gamopetalous, white, yellow, orange, pink, or purple, the lobes 

shorter than the tube; stamens 4, inserted on the upper portion of the corolla tube, the anthers linear, 

inserted; ovary superior with 2 connate carpels, the style short, the stigma claviform or globose. Fruit 

a septicidal capsule or less frequently a berry. A genus of about 100 species of amphitropical distribution. 

1. Buddleja madagascariensis Lam., Encycl. 1: 

513. 1785. 

Fig. 53. A-E 

Scandent shrub, sometimes twining, 2-5(10) 

m in length. Branches obtusely quadrangular, 

tomentose, glabrescent. Leaves opposite; blades 

8-16 × 2.7-4.5(6.2) cm, lanceolate, ovate, or 

elliptical, coriaceous, the apex acuminate or less 

frequently acute, the base rounded, the margins 

entire or denticulate; upper surface sparsely 

tomentulose; lower surface lanate-tomentose, 

whitish or ferruginous, with prominent venation; 

petioles 5-15 mm long; stipules early deciduous. 

Flowers short-pedicellate to subsessile, in dichasia 

grouped in terminal thyrses, 5-25 cm long. Calyx 

Reference: Norman, E., 2000. Buddlejaceae. Flora Neotropica Monograph 81. 225 pp. 

11. Family CACTACEAE 


campanulate or broadly campanulate, whitetomentose, 

2.5-4 mm long, the sepals deltate, 0.5- 

1 mm long; corolla yellow-orange, tubular, 8.5- 

11 mm long, white-tomentose outside. Fruit 

unknown in the collections from Puerto Rico. 

Phenology: Collected in flower in April and 

September. 

Status: Exotic, cultivated, escaped or 

persistent, uncommon. 

Distribution: Ornamental, cultivated in the 

gardens of the Cordillera Central, escaped or 

persistent in the Villalba area. Native to 

Madagascar, but widely cultivated in the 

subtropics. 


1a. Stems dark brown, cylindrical, without ribs; leaves present ........................................... 2. Pereskia 

1b. Stems green, angular, compressed, or cylindrical, but then with longitudinal ribs; leaves absent.2 

2a. Stems cylindrical, with 5-8 slightly prominent ribs; spines acicular; hypanthium densely lanatepubescent 

..................................................................................................................... 3. Selenicereus 

2b. Stems angular, with 3-4 prominent ribs; spines conical or subconical; hypanthium glabrous 

....................................................................................................................... 1. Hylocereus

142 


1. HYLOCEREUS 

Lianas, climbing by adventitious roots, with numerous lateral branches. Stems green, angular, with 

3 or 4 prominent longitudinal ribs. Leaves absent; areoles surrounded by conical spines along the 

margin of the ribs. Flowers bisexual, actinomorphic, solitary, sessile, usually at the ends of the branches; 

outer tepals scale-like; inner tepals petaliferous; stamens numerous, the filaments shorter than the 

perianth; ovary inferior, forming an elongate, infundibuliform hypanthium, the style elongate, not 

exserted, the stigmas numerous, filiform. Fruit a berry with persistent tepals; seeds numerous, minute. 

A Caribbean genus of about 20 species. 

Key to the species of Hylocereus 

1a. Stems with 3 or 4 longitudinal ribs, with the groups of spines on small mounds ..... 1. H. trigonus 

1b. Stems with 3 longitudinal ribs, forming a depression in the areas where the groups of spines are 

borne ...............................................................................................................................2. H. undatus 

1. Hylocereus trigonus (Haw.) Saff., Annual Rep. 

Board Regents Smithsonian Inst. 1908: 553. 

1909. 

Fig. 54. A-C 

BASIONYM: Cereus trigonus Haw. 

Pitahaya, Chickenet, Strawberry pear 

Much branched vine, which climbs by aerial 

roots and attains 10 m in length. Stems fleshy, 

1.4-2 cm wide, lenticellate, triangular, with 3 or 

4 prominent, undulate, longitudinal ribs, with a 

group of spines on each of the protuberances that 

are found along the margin; spines conical, 4-7 

mm long, grayish or brown, usually in groups of 

8. Flowers solitary, pendulous, 20-24 cm long, 

axillary to the group of spines or terminal on the 

branches; perianth infundibuliform; outer tepals 

fleshy, green, the basal ones ovate, ca. 2 cm long, 

the distal ones oblong, 3.5-7.5 cm long, the inner 

tepals white, turning yellowish when mature. 

Berries ovoid, pink-purple, 12-14 × 5-7 cm, with 

tepals of the hypanthium persistent; seeds 

numerous, minute, black, covered with a white 

gelatinous matrix. 

Phenology: Flowering and fruiting from July 

to December. 


Distribution: In dry forests or coastal thickets. 

Also on Culebra, Vieques, St. Croix, St. John, St. 

Thomas, Tortola, and Virgin Gorda; Hispaniola. 

Public forests: Ceiba, Guajataca, Guánica, and 

Río Abajo. 

2. Hylocereus undatus (Haw.) Britton & Rose in 

Britton, Fl. Berm. 256. 1918. 

Fig. 54. D-E 

BASIONYM: Cereus undatus Haw. 

Pitahaya 

Much branched vine, creeping or climbing by 

aerial roots, attaining 7 m in length. Stems fleshy, 

2-5 cm wide, trigonal, with 3 longitudinal ribs 

with undulate margins, with depressions in which 

are found groups of 1-5 subconical brown spines, 

3-4 mm long. Flowers solitary, up to 30 cm long, 

axillary to the group of spines or terminal on the 

branch; perianth glabrous, infundibuliform; outer 

tepals greenish yellow, sometimes with a reddish 

tinge, those of the hypanthium ovate-lanceolate, 

the distal ones linear-lanceolate, the inner tepals 

petaliferous, white, longer than the outer ones; 

stamens yellow; inserted; stigmas greenish yellow. 

Berry red, ellipsoid, fleshy, 6-12 × 5-7 cm, with 

persistent tepals; seeds numerous, minute, black, 

covered with a red pulp. 

Phenology: Flowering from July to September 

and fruiting from September to December. 


naturalized. 

Distribution: Cultivated from the coast to the 

Cordillera Central, naturalized in dry or coastal 

areas. Native to Mexico, but widely cultivated in 

the tropics. Also on St. John and St. Thomas. 

Public forest: Guánica.


144 


2. PERESKIA 

Shrubs, trees or scandent shrubs. Stems cylindrical, without ribs, with axial areoles surrounded by 

spines; glochids absent. Leaves alternate, persistent, succulent, petiolate; blades simple. Flowers 

actinomorphic, unisexual, solitary or in racemes, panicles, or cymes; receptacle with areoles and 

bracteoles in 2 series; perianth of sepaloid and petaloid tepals; stamens numerous, the filaments usually 

unequal, shorter than the perianth; ovary inferior or half-inferior, the style thick, the stigmas numerous, 

digitiform, papillose. Fruit a berry with persistent bracts and areoles; seeds few, small. A neotropical 

genus of approximately 17 species. 

1. Pereskia aculeata Miller, Gard. Dict. ed. 8. 

1768. 

Fig. 55. A-E 

Hortensia de bejuco, Barbados gooseberry, 

Lemon vine 

Clambering shrub or liana attaining 15 m in 

length, much branched from a woody base. Stems 

woody, dark brown, cylindrical, without ribs, 2-3 

cm in diameter, with numerous areoles; spines 

dimorphic, up to 1.5 cm long, usually deciduous; 

branches scandent or pendulous. Leaves alternate, 

persistent; blades 4-7 × 1.5-5 cm, lanceolate, 

ovate, or oblong, fleshy, glabrous, the apex shortacuminate, 

the base cuneate or rounded, the 

margins entire; upper and lower surface glabrous; 

3. SELENICEREUS 

petioles 3-7 mm long, with a pair of stipular 

spines, recurved at the base. Flowers few, in 

terminal racemes; hypanthium crateriform, with 

several bracteoles and areoles with acicular spines. 

Tepals white, 2.5-3 cm long; stamens numerous, 

in two series, yellow; ovary inferior. Berry yellow 

or orange, globose, ca. 2 cm in diameter, with 

several bracteoles and persistent spines; seeds few, 

4-5 mm wide, lenticular or rounded, dark brown. 


in fruit in January and February. 


uncommon. 

Distribution: In dry thickets. Also on Vieques, 

St. Croix, St. John, and St. Thomas; throughout 

the Neotropics, its place of origin unknown. 

Lianas or creeping shrubs, which climb by adventitious roots. Stems green, elongate, with 4-12 

angles or ribs. Leaves absent; areoles minute, usually forming a small promontory, densely covered 

with lanate hairs and acicular spines. Flowers actinomorphic, bisexual, nocturnal, solitary; perianth 

elongate, infundibuliform; outer tepals scale-like, with areoles densely covered with long, lanate hairs; 

inner tepals petaliferous; stamens numerous, inserted on the perianth; ovary inferior, the style thick, 

the stigmas numerous; ovary inferior. Fruit a large, reddish berry, covered by basal bracts and several 

spines; seeds numerous, minute. A Caribbean genus of about 16 species. 

1. Selenicereus grandiflorus (L.) Britton & Rose, 

Contr. U.S. Natl. Herb. 12: 430. 1909. 

Fig. 55. F-G 

BASIONYM: Cactus grandiflorus L. 

Liana with numerous lateral branches, which 

climbs by aerial roots, attaining 6 m in length. 

Stems fleshy, 2-2.5 cm in diameter, cylindrical, 

with 5-8 ribs; areoles prominent, along the ribs, 

covered with spines and whitish lanate hairs, the 

spines acicular, 5-8 mm long, yellowish, in groups 

of 5. Flowers solitary, pendulous, axillary to the 

group of spines or terminal, 15-18 cm long; 

perianth infundibuliform, the tube elongate, 

densely lanate outside; outer tepals linear, salmoncolored; 

inner tepals whitish, obovate; stamens 

numerous, in two series, inserted. Berry ovoid, 

red or yellowish, ca. 8 cm long; seeds numerous, 

minute, black, covered with a white pulp. 

Phenology: Collected in flower in June. 


locally common.


146 


Distribution: In coastal forests and thickets. 

Also on St. Croix, St. John, St. Thomas, and 

12. Family CAPPARACEAE 

1. CAPPARIS 

Tortola; species native to Jamaica and Cuba, but 

widely cultivated throughout the tropics. 

Small trees or shrubs, erect or exceptionally scandent. Leaves alternate, petiolate; blades simple, 

usually lepidote; axillary extrafloral nectaries usually present; stipules absent. Flowers actinomorphic, 

4-merous, bisexual, pedunculate, solitary or in short axillary racemes; sepals valvate or imbricate; 

petals free, white, pink, or yellowish; stamens numerous, much longer than the petals; ovary superior, 

with two carpels, borne on a gynophore; ovules few or numerous, with parietal placentation. Fruit an 

elongate follicle, woody or coriaceous, indehiscent or irregularly dehiscent; seeds numerous, in 2 rows, 

with a fleshy covering. A tropical and subtropical genus of 150-350 species.


1. Capparis flexuosa (L.) L., Sp. Pl. ed. 2, 722. 

1762. 

Fig. 56. A-E 

BASIONYM: Morisonia flexuosa L. 

SYNONYM: Capparis saligna Vahl 

Burro, Palo de burro, Palinguam, Palawang, 

Black witty, Bottle wiss, Goat wiss, 

Limber caper 

Woody shrub, erect or clambering, much 

branched from the base, 2-10 m in length. 

Branches pendulous, glabrous, light brown, 

smooth, attaining 20 cm in diameter at the base. 

Leaves alternate; blades 4-10 × 1.5-2.5 cm, oblong 

or linear, coriaceous, glabrous, the apex rounded 

or emarginate, the base rounded, the margins 

entire, revolute; upper surface yellowish green, 

shiny or dull; lower surface pale green, dull, with 

a prominent midvein; petioles 5-8 mm long, light 

brown. Flowers fragrant, nocturnal, clustered in 

13. Family CAPRIFOLIACEAE 

1. LONICERA 

terminal corymbs. Calyx yellowish green, of 4 

imbricate sepals, rounded, fleshy, 6-10 mm long; 

petals green or yellowish green, 4, elliptical, 1.5- 

3 cm long, concave; stamens numerous, white, 

4-6 cm long. Follicles fleshy, cylindrical-torulose, 

6-8 cm long, light brown, with longitudinal 

dehiscence, the inner walls red-orange; seeds 

pendulous from the parietal placenta, oblong, ca. 

1 cm long, with a white, fleshy covering. 

Phenology: Flowering from March to October 

and fruiting in August and September. 


Distribution: In coastal forests and thickets 

or in semi-humid forests on karst limestone. Also 

on Culebra, Desecheo, Vieques, Mona, and the 

Virgin Islands; the Antilles and from the United 

States (Florida) to South America. 

Public forests: Ceiba, Guánica, Mona, 

Piñones, and Susúa. 

Vines, shrubs, or trees. Leaves opposite or rarely in whorls of 3, sessile or petiolate; blades simple; 

stipules absent. Flowers bisexual, 5-merous, 2 or 3 in axillary cymes. Calyx tubular, 5-dentate or rarely 

truncate at the apex, adnate to the ovary; corolla zygomorphic, tubular, infundibuliform, or campanulate, 

with the limb bilabiate, with 2 long lobes and 3 short lobes; stamens 5, subequal, exserted; ovary 

inferior, with 2-3(-5) locules, with axile or rarely parietal placentation, the ovules pendulous, 3-8 per 

locule. Fruit a fleshy berry, with few ovate seeds. A genus of about 200 species, the majority of the 

Northern Hemisphere. 

1. Lonicera japonica Thunb. ex Murray, Syst. 

Veg. (ed. 14), 216. 1784. 

Fig. 57. A-E 

Madreselva, Honeysuckle 

Woody vine, twining or clambering, much 

branched, 5-8 m in length. Stems strong, flexible, 

cylindrical, pilose, brown, smooth. Leaves 

opposite; blades 4-7.5 × 1.2-3 cm, lanceolate, 

oblong-lanceolate, or elliptic-lanceolate, 

chartaceous, glabrous except for some hairs on 

the midvein, the apex acute or obtuse, abruptly 

acuminate or mucronate, the base rounded, 

subtruncate, or obtuse, the margins entire, 

revolute, ciliate; upper surface dark green, dull; 

lower surface pale green, dull, with a prominent 

midvein; petioles 5-12 mm long, brown, 

pubescent. Flowers fragrant, sessile, in pairs at 

the end of short axillary branches; bracts 2, at the 

base of each flower, ovate, rounded, ca. 0.7 mm 

long, ciliate. Calyx green, tubular, ca. 3 mm long, 

the sepals 5, subulate, ca. 1 mm long; corolla 

white, turning yellowish when mature, 2.2-2.5 cm 

long, infundibuliform, the tube puberulous 

outside; the limb with 5 lobes, one of which is 

longer (ca. 2.2 cm) and free; stamens 4, exserted; 

style exserted, with the stigmatic surface 

claviform. Berry black, 6-7 mm long, ellipsoid,

148 


although in the Americas it apparently does not 

frequently produce fruits. 




locally common. 

Distribution: Native of Asia, but naturalized 

in the Cordillera Central in Puerto Rico in the 

Villalba area (Cerro Maravilla sector) and 

throughout the New World. This species seems 

not to have been naturalized in Puerto Rico at the 

end of the nineteenth century, because it was only 

known from a single collection made by Sintenis 

in 1886 from a plant cultivated in the Adjuntas 

area. At present this species is naturalized along 

Highway 143 in the Toro Negro Forest Reserve. 

It seems to reproduce in Puerto Rico exclusively 

by vegetative means, since it has never been 

collected with fruits. 

Public forest: Toro Negro.


14. Family CELASTRACEAE 


1a. Leaves slightly shiny on the upper surface, chartaceous, the margins crenulate or serrate, the tertiary 

veins inconspicuous; filaments of the stamens broadened gradually toward the base, adnate to the 

floral disc; petals pubescent on the distal portion ...................................................... 1. Hippocratea 

1b. Leaves dull on the upper surface, coriaceous, the margins subentire, denticulate, or crenulate, the 

tertiary veins conspicuous; filaments of the stamens more or less of the same width along their 

length, inserted on the floral disc; petals glabrous ....................................................... 2. Pristimera 

1. HIPPOCRATEA 

Lianas with opposite branches, elongate, twining. Leaves opposite, petiolate; blades simple, crenate, 

or serrulate; petioles slender; stipules absent. Flowers minute, bisexual, actinomorphic, in axillary 

cymes, panicles, corymbs, or pseudoracemes; bracts small, chartaceous. Calyx rotate, of 5 rounded 

sepals; corolla of 5 free petals; disc fleshy, pulviniform; stamens 3, the filaments adnate to the upper 

portion of the disc; ovary superior, 3-carpellate, obtusely triangular, trilocular, with 4-6 ovules per 

locule, the style subulate, the stigma inconspicuous. Fruit of 3 divergent, flattened mericarps, each 

dehiscent by a medial suture; seeds compressed, with a basal wing. A genus of 3 species, 2 in Africa 

and 1 in the Neotropics. 

1. Hippocratea volubilis L., Sp. Pl. 1191. 1753. 

Fig. 58. A-H 

SYNONYM: Hippocratea ovata Lam. 

Bejuco prieto 

Woody vine, climbing, which supports itself 

on other plants by twining lateral branches and 

attains 10-15 m in length. Stems green, 

quadrangular, becoming grayish and cylindrical 

when mature; branches opposite, twining, 

developing before the leaves. Leaves opposite; 

blades 4-17 × 2-8 cm, elliptical or oblong, 

coriaceous, the apex acute or acuminate, the base 

obtuse or rounded, sometimes unequal, the 

margins crenulate or slightly serrate; upper 

surface shiny, glabrous, with the veins slightly 

sunken; lower surface pale green, dull, glabrous, 

with the venation slightly prominent; petioles 3- 

7 mm long; stipules absent. Flowers numerous, 

in axillary corymbs. Calyx yellowish green, of 5 

rounded sepals, ca. 1 mm long; corolla of 5 

yellowish or whitish petals, free, oblanceolate, 2.5- 

3 mm long, pubescent on the distal portion; 

stamens 3, the filaments connate into a conical 

disc, which almost totally surrounds the ovary; 

ovary superior, tricarpellate, the style one, with a 

punctiform stigma. Mericarps 3, divergent, 

flattened, conspicuously veined, 4-8 cm long; 

seeds few, winged, ca. 4 cm long, the seed-bearing 

portion elliptical, the wing oblong. 


fruiting from October to January. 


Distribution: In secondary forests and along 

roadsides at middle and lower elevations in moist 

areas. Also on Vieques; the Antilles, Central 

America, South America, and the United States 

(Florida). 

Public forests: Cambalache, Ceiba, El Yunque, 

Guajataca, Maricao, Río Abajo, Susúa, and Vega.

150 



2. PRISTIMERA 

Lianas or scandent shrubs; glabrous, branches usually opposite. Leaves opposite, petiolate; blades 

simple; petioles slender; stipules absent. Flowers minute, bisexual, in axillary panicles, corymbs, or 

pseudoracemes; bracts small, chartaceous. Calyx rotate, of 5 rounded sepals; corolla of 5 free petals; 

disc annular, inconspicuous; stamens 3; ovary superior, 3-carpellate, obtusely triangular, trilocular, 

with 2-8 ovules per locule, the style short, the stigma obtuse or capitate. Fruit of 3 divergent, flattened 

mericarps, each dehiscent by a medial suture; seeds compressed, with a basal wing. A genus of 24 

species of pantropical distribution. 

1. Pristimera caribaea (Urb.) A. C. Sm., 

Brittonia 3: 378. 1940. 

Fig. 59. A-F 

BASIONYM: Hippocratea caribaea Urb. 

SYNONYM: Hippocratea domingensis Urb. & Ekman 

Woody vine, climbing, which supports itself 

on other plants by twining lateral branches and 

attains 10 m or more in length. Stems grayish, 

dark brown and lenticellate when mature. Leaves 

opposite; blades 6-11 × 2-6 cm, elliptical or ovateelliptical, 

coriaceous, glabrous, the apex acute or 

acuminate, the base acute to obtuse, the margins 

entire, denticulate or crenulate, the veins 

prominent on both surfaces. Flowers numerous, 

in dichasial cymes, dichotomous, axillary, 1.5-4 

cm long; pedicels ca. 1 mm long. Calyx greenish 

yellow, of 5 triangular or broadly ovate sepals, 

0.5-0.8 mm long, erose or ciliate-glandular at the 

margin; petals 5, elliptical or oblong, greenish 

yellow, 1.5-2.6 mm long; disc annular, 0.8-1.2 

mm wide; stamens ca. 1 mm long; ovary glabrous, 

conical. Mericarps 3, compressed, elliptical, 3.5- 

5 cm long; seeds usually 4 per locule, ca. 1.5 cm 

long, winged. 


November to January. 


Distribution: Of wide distribution in Puerto 

Rico, in forests in the zone of mogotes, the 

Cordillera Central, Sierra Bermeja, and Cerro de 

La Pandura. Also in Hispaniola, the Lesser 

Antilles, and Guyana. 

Public forests: Guajataca, Maricao, and Río 

Abajo. 

Commentary: Hippocratea, Pristimera, and 

various genera have been considered as belonging 

to the Family Hippocrateaceae. Nevertheless, 

recent taxonomic evidence suggests that the 

Hippocrateaceae is an artificial group not 

distinguishable from the Celastraceae. For this 

reason, these have been added to the Celastraceae. 

References: Smith, A. C. 1940. The American species of Hippocrateaceae. Brittonia 3: 341-555. 

van Gorts, A. R. A. and A. M. W. Mennega. 1994. Hippocrateaceae. Flora of the Guianas. 16: 1-81. 

15. Family CLUSIACEAE 

1. CLUSIA 

Trees or erect shrubs, usually stranglers or less frequently scandent, dioecious, with abundant 

yellowish or cream-colored latex. Leaves opposite, usually petiolate; blades simple, thick, coriaceous. 

Flowers unisexual, actinomorphic, solitary or in terminal cymes. Calyx of 4-6 sepals; corolla of 4-9 

free petals; stamens numerous, fertile in the staminate flowers, sterile (staminodia) in the pistillate 

flowers, the filaments usually united at the base; ovary superior, with 4-12 carpels, the ovules numerous, 

with axile placentation, the stigma peltate. Fruit a fleshy capsule, valvicidally dehiscent; seeds numerous, 

with a fleshy, reddish covering. A neotropical genus of about 150 species.

152 


1. Clusia gundlachii A. Stahl, Estud. 2: 122. 

1884. 

Fig. 60. A-G 

Cupey de altura, Bejuco de cupey, 

Cupeillo de altura 

Clambering shrub attaining 7 m in length, 

with abundant cream-colored latex. Trunk 

cylindrical, up to 10 cm in diameter at the base; 

branches pendulous, opposite, subcylindrical or 

slightly compressed, glabrous, with long, 

pendulous aerial roots. Leaves opposite; blades 

5.5-10 × 2.1-6 cm, oblanceolate, elliptical, oval, 

or ovate, coriaceous, glabrous, the apex acute or 

obtuse, the base cuneiforme to obtuse, the margins 

entire, revolute; upper surface dark, slightly shiny,


with the midvein yellowish; lower surface 

yellowish green, with a prominent midvein; 

petioles thick, 1-2 cm long. Flowers numerous, 

unisexual, arranged in dichasial cymes to form a 

terminal thyrse, as long as wide. Calyx green, 

crateriform, of 4 sepals, concave, rounded, in 2 

series, the 2 outer sepals ca. 3 mm long, the inner 

ones ca. 4 mm long; corolla tubular, ellipsoid or 

ovoid, 2-3 mm long; staminate flowers with 10 

stamens included within the corolla; pistillate 

flowers with sterile stamens, ovary urceolate, 4- 

5-locular, the stigmas sessile. In addition to the 

fertile inflorescences, this species produces sterile 

inflorescences, pendulous, much branched, with 

numerous green bracts, opposite and decussate 

along the axes. Capsule fleshy, ovoid or ellipsoid, 

1.6-2 cm long, greenish, opening by 4-5 valves, 

each of which contains a row of minute, elliptical 

seeds, covered with a sticky orange matrix. 



Status: Endemic to Puerto Rico, rather 

common. 

Distribution: Principally in moist forests along 

the Cordillera Central and the Sierra de Luquillo, 

but extending toward the dry forests of the south 

coast. 

16. Family COMBRETACEAE 

1. COMBRETUM 

Public forests: Carite, El Yunque, Guánica, 

Guilarte, Maricao, Río Abajo, Susúa, and Toro 

Negro. 

Commentary: This species, described by Dr. 

Agustín Stahl, was dedicated to Dr. Juan 

Gundlach, a German naturalist resident in Cuba, 

who had visited Puerto Rico in 1873 and in 1875- 

76. This species is superficially similar to Clusia 

minor L., which can be distinguished from C. 

gundlachii by the following key. 

1a. Clambering shrub with long, pendulous 

branches; leaves oblanceolate, elliptical, oval, 

or ovate, the apex acute or less frequently 

obtuse; flowers functionally unisexual, the 

pistillate ones with staminodia; corolla tubular, 

ellipsoid or ovoid, 2-3 mm long, closed at the 

apex to form a calyptra; fruit ovoid or ellipsoid 

………..………………….....…C. gundlachii 

1b. Erect shrub; leaves oblanceolate or spathulate, 

the apex rounded or obtuse; flowers 

hermaphroditic or pistillate (the latter with a 

resinous ring in place of the staminodia); 

corolla of free petals, rounded, ca. 1.5 cm long; 

fruit ovoid or subglobose..........……C. minor 

Trees, erect or scandent shrubs, or lianas. Leaves opposite or sometimes alternate or whorled; 

blades simple; petioles articulated at the base. Flowers 4-5-merous, bisexual, sessile or short-pedicellate, 

arranged in heads, racemes, spikes, or compound paniculiform inflorescences; bracts foliaceous or 

reduced. Calyx forming an infundibuliform, tubular, or cupular hypanthium that projects beyond the 

ovary, with the sepals on the distal portion of the hypanthium; corolla of free petals or absent; stamens 

8-10, in one or two whorls, exserted or inserted; ovary inferior, with 2-6 ovules, the style simple, free 

or adnate to the hypanthium. Fruit dry, indehiscent or dehiscent, with 4 or 5 wings or ribs; seed one. 

About 270 species of tropical distribution, with numerous species in Africa. 

Key to the species of Combretum 

1a. Hypanthium infundibuliform, red-purple; stamens exserted, in a single whorl; inflorescences spicate, 

with the flowers densely clustered ........................................................................ 1. C. grandiflorum 

1b. Hypanthium long-tubular, green; stamens inserted in two whorls; inflorescences racemose, with 

the flowers dispersed ..................................................................................................... 2. C. indicum

154 


1. Combretum grandiflorum G. Don, Edinburgh 

Philos. J. 346. 1824. 

Fig. 61. A-D 

Twining liana, 3-10 m in length. Branches 

cylindrical, somewhat compressed in the area of 

the nodes, sparsely pilose or glabrous, usually with 

the pith hollow. Leaves opposite; blades 7.5-14.5 

(25) × 3.5-7(12) cm, elliptical, obovate, or ovate, 

chartaceous or coriaceous, the apex obtuse or 

rounded, the base obtuse, slightly asymmetrical, 

the margins entire; upper surface glabrous, with 

the midvein appressed-pubescent; lower surface 

glabrous except for some papillae along the 

primary and secondary veins, with prominent


venation; petioles 10-13 mm long, geniculate at 

the base, appressed-pubescent. Flowers 4-5merous, 

ascending, numerous, in axillary 

racemes, horizontal, 6-12 cm long; pedicels 3-6 

mm long. Hypanthium infundibuliform, 8-12 mm 

long, glabrous, red-purple, puberulous; sepals 

triangular, ca. 3 mm long; petals red-purple, 

spathulate, 1-1.2 cm long; stamens exserted, the 

filaments unequal; style simple, free, exserted. 

Fruit oval in outline, retuse at the apex, 5-winged, 

pale yellow or stramineous, 3-4 cm long, stipitate 

at the base. Seed one, pentagonal, 7-10 mm long. 

Phenology: Collected in fruit in February. 

Status: Exotic, cultivated, locally naturalized, 

uncommon. 

Distribution: Ornamental plant, native to 

central Africa, cultivated, naturalized in the Cerro 

Las Mesas area in Mayagüez. 

2. Combretum indicum (L.) Jongkind, Fl. Gabon 

35: 48. 1999. 

BASIONYM: Quisqualis indica L. 

Fig. 61. E-G 

Twining liana, attaining 6 m in length. Stem 

much branched from the base; branches 

cylindrical, somewhat compressed in the area of 

the nodes, glabrous or puberulous, usually with 

the pith hollow; adult stems deciduous, with 3 

persistent spines at the nodes. Leaves opposite or 

subopposite; blades 6-17.5 × 2.2-7 cm, elliptical, 

oblong, or lanceolate, chartaceous, the apex 

acuminate, the base rounded, the margins entire; 

upper surface glabrous, with a prominent midvein; 

lower surface ferruginous-tomentulose or 

puberulous, with prominent venation; petioles 5- 

12 mm long. Flowers sessile or subsessile, in 

terminal spikes or racemes on short lateral 

branches. Hypanthium green, tubular, 4-6.5 mm 

long, pubescent; sepals green, triangular, ca. 2 

mm long; petals 5, oblong or oblanceolate, 1-1.5 

cm long, pink, turning red when mature; stamens 

exserted, the filaments unequal; disc absent; style 

exserted. Fruit elliptical in outline, angular, with 

5 narrow wings, ca. 3 cm long. 

Phenology: Flowering from July to March. 


uncommon. 

Distribution: Ornamental plant, native to 

tropical Asia, naturalized along roadsides. Also 

on St. Croix, St. John, and St. Thomas. 


References: Jongkind, C. C. H. 1991. Novitates Gabonenses 6. Some critical observations on 

Combretum versus Quisqualis (Combretaceae) and description of two new species of Combretum. Bull. 

Mus. Natn. Hist. Nat., Paris, Ser. 4 12: 275-280; Jongkind, C. C. H. 1999. Combretaceae. In: Flora du 

Gabon. Vol. 35. 115 pp. París. 

17. Family CONNARACEAE 

1. ROUREA 

Vines, shrubs, or trees. Leaves alternate, compound, imparipinnate or unifoliolate, petiolate; stipules 

present. Flowers 5-merous, bisexual, actinomorphic, in terminal, subterminal, or axillary panicles. 

Calyx of imbricate sepals, usually glandular; corolla of 5 petals, free or connate at the base; stamens 

10, in two series, the filaments glabrous, connate at the base to form a short tube, the anthers dehiscent 

by longitudinal sutures; ovary superior, with 5 free carpels, each with two ovules, the stigma capitate. 

Fruit a follicle, dehiscent by a longitudinal suture, with the calyx accrescent and persistent at the base; 

seed one, black, arillate at the base. A pantropical genus of about 85 species, 42 of which are distributed 

throughout tropical America.

156 



1. Rourea surinamensis Miq., Linnaea 26: 221. 

1853. 

Fig. 62. A-H 

SYNONYMS: Rourea frutescens sensu Bello, non Aubl. 

Rourea glabra sensu Griseb., non Kunth 

Juan caliente, Bejuco de garrote 

Woody vine, twining, attaining more than 20 

m in length. Stems strong, flexible, grayish or 

brown, up to 2 cm in diameter; lateral branches 

short, twining like tendrils. Leaves alternate, 

pinnately compound or less frequently 

unifoliolate; leaflets 3 or 5, opposite or 

subopposite, 4-15 × 1.5-6.5 cm, elliptical, oblong, 

or ovate, coriaceous, glabrous, the apex 

acuminate, the base rounded (or in young plants 

cordiform), the margins entire, revolute; upper 

surface dark, shiny, with the midvein sunken, the 

secondary veins slightly prominent; lower surface 

pale green, dull, with the reticulate venation 

prominent; petioles 3-8 cm long, with the base 

swollen; petiolules swollen, ca. 5 mm long; 

stipules minute. Flowers numerous, in clustered, 

axillary panicles or racemes, 5-10 cm long. Calyx 

crateriform, 2.5-3 mm long, of 5 oblong sepals, 

ca. 2 mm long; petals white, spathulate, ca. 4 mm 

long, early deciduous; stamens 10, in two series, 

the filaments unequal, the anthers ellipsoid; 

gynoecium of 5 free carpels, pubescent, the style 

elongate, the stigma peltate or bilobed. Follicles 

ellipsoid, 1-1.5 cm long, reddish when mature; 

seeds black, with a white aril at the base. 

Phenology: Flowering from March to 

September and fruiting from December to April. 

Status: Native, rather common. 

Distribution: in mature or secondary moist 

forests, at middle elevations. Also in Hispaniola, 

the Lesser Antilles, Trinidad, the Guianas, and 

eastern Venezuela. 


Reference: Forero, E. 1976. A revision of the American species of Rourea subgenus Rourea 

(Connaraceae). Mem. New York Bot. Gard. 26: 1-119. 

18. Family CONVOLVULACEAE 


1a. Flowers arranged in panicles; corolla white, 4-6 mm long......................................... 8. Poranopsis 

1b. Flowers solitary or in simple or compound cymes; corolla of diverse colors, when white they exceed 

2.5 cm long, when violet-pink, lavender, or yellow they are ca. 5 mm long................................... 2 

2a. Stigmas globose, subglobose, biglobose, or bilobate. ....................................................................... 3 

2b. Stigmas elongate (filiform or oblong). ............................................................................................ 11 

3a. Sepals conspicuously unequal, the outer ones broader, concealing the inner ones .......... 1. Aniseia 

3b. Sepals more or less equal or slightly unequal, the outer ones usually smaller than the inner ones.........4 

4a. Fruits indehiscent, irregularly dehiscent or dardily dehiscent by the dissolution of the pericarp. . 5 

4b. Fruits capsular, 4-valvate, sometimes with the exocarp operculate. ................................................ 9 

5a. Lower surface of the leaves densely pubescent; axes of the inflorescences densely pubescent or 

tomentose. ........................................................................................................................................... 6 

5b. Lower surface of the leaves and axes of the inflorescences glabrous or puberulous; fruits indehiscent 

or irregularly or tardily dehiscent, fibrous or crustaceous. ............................................................... 7 

6a. Fruit indehiscent, fleshy, covered by the accrescent sepals; corolla violet-pink ............ 2. Argyreia 

6b. Fruit with the exocarp separating like an operculum, endocarp opening irregularly, sepals not 

covering the fruit; corolla white or white with a yellow center .................................. 7. Operculina

158 



7a. Fruit covered by the accrescent, overlapping sepals; corolla violet-pink, 6-8 cm long; fruit fibrous, 

tardily dehiscent by the dissolution of the pericarp .................................................... 9. Stictocardia 

7b. Fruit exposed, the sepals although accrescent, not covering the fruit; corolla white or yellow, 2.5-3 

cm long; fruit crustaceous, indehiscent or irregularly dehiscent ..................................................... 8 

8a. Fruit ellipsoid, indehiscent, 1-1.2 cm long, with only one seed inside; corolla white with the center 

violet ................................................................................................................................. 10. Turbina 

8b. Fruit ovoid, irregularly indehiscent, 1.5-2.5 cm long, seeds generally 4; corolla yellow ................. 

................................................................................................................... 6. Merremia (in part) 

9a. Leaves narrowly lanceolate or linear, 0.5-6 mm wide, with the base hastate or auriculate; corolla 

yellow, 1-2 cm long ..................................................................................................... 11. Xenostegia 

9b. Leaves of diverse forms, if lanceolate and hastate then with the corolla white, 3-4 cm long. ..... 10 

10a. Anthers twisted after opening; sepals accrescent at the base of the fruit; leaves usually palmately 

compound (except in the case of M. umbellata); corolla yellow or white with a purple center 

.................................................................................................................. 6. Merremia (in part) 

10b. Anthers straight (not twisted) after opening; sepals generally not accrescent; leaves simple, or if 

palmately compound then with the corolla violet-pink or red ........................................ 4. Ipomoea 

11a. Stigmas filiform; corolla white ................................................................................. 3. Convolvulus 

11b. Stigmas oblong; corolla blue, white, or red ...........................................................5. Jacquemontia 

1. ANISEIA 

Herbaceous vines, twining, producing watery latex. Leaves alternate; blades simple; stipules absent. 

Flowers 5-merous, bisexual, actinomorphic, solitary or in axillary dichasia; bracts 2, on the distal 

portion of the peduncle. Calyx of unequal sepals, the 2 outer ones much broader, concealing the inner 

ones; corolla white, infundibuliform, the limb 5-dentate or entire, with 5 lines of pubescence outside; 

stamens 5, shorter than the corolla, the filaments adnate to the corolla; ovary superior, glabrous, bilocular, 

each locule with two ovules, the style solitary, the stigma bilobed. Fruit a globose capsule that opens 

along 4 valves, covered by the accrescent sepals; seeds 4 per fruit. A genus of neotropical origin, of 5 

species, of which one has been introduced into the Old World tropics. 

1. Aniseia martinicensis (Jacq.) Choisy, Mém. 

Soc. Hist. Nat. Genève 8: 66. 1837. 

Fig. 63. A-D 

BASIONYM: Convolvulus martinicensis Jacq. 

Herbaceous vine, twining, climbing, or 

creeping, 2-4 m in length. Stems glabrous or 

puberulous when young, with watery latex. Leaves 

alternate; blades simple, 4-9 × 2.4-3.7 cm, 

narrowly lanceolate, elliptical, or oblanceolate, 

membranaceous, the apex obtuse and mucronate, 

the base obtuse, acute, or attenuate, the margins 

entire or crenate; upper surface dull, glabrous, 

granular, with the midvein slightly prominent; 

lower surface pale green, glabrous, dull, with the 

venation slightly prominent and puberulous; 

petioles puberulous, slender, 1-2 cm long. Flowers 

solitary, axillary; peduncles 3-5 cm long, with a 

pair of minute bracteoles on the distal portion; 

pedicel ca. 1 cm long. Calyx green, of 5 unequal 

sepals, ovate, acuminate, the two outer ones 12- 

17 mm long, broader than the 3 inner ones; corolla 

white, campanulate, 2.5-3 cm long; stamens not 

exserted. Capsules 4-valvate, ovoid, 1.3-1.5 cm 

long, partially covered by the accrescent sepals; 

seeds dark brown to almost black, ca. 6.5 cm long, 

with two sides flat and one convex, sparsely 

covered with minute, squamose hairs, especially 

on the angles.

160 



and April and in fruit from December to February 

and in July. 


2. ARGYREIA 

3. CONVOLVULUS 

Distribution: In marshy areas near Laguna 

Tortuguero and at the mouth of the Humacao 

River. Throughout the Neotropics. 

Public forest: Tortuguero. 

Twining lianas. Leaves alternate, petiolate; blades simple; stipules absent. Inflorescences of simple 

dichasial cymes. Flowers 5-merous, bisexual, with actinomorphic symmetry. Calyx of 5 sepals, variable 

in shape and size, usually pubescent and accrescent in fruit; corolla purple, pink, or white, 

infundibuliform, campanulate, or tubular, the limb almost entire to deeply 5-lobed; stamens 5, inserted 

or exserted; ovary superior, glabrous or pubescent, 2-4-locular, the style solitary, inserted or exserted, 

the stigma biglobose. Fruits indehiscent, ellipsoid to globose, fleshy, red, orange, or yellowish; seeds 4 

or fewer per fruit. A genus of 90 species native to continental tropical Asia, Malaysia, and northern 

Australia; the following species is cultivated in the New World. 

1. Argyreia nervosa (Burm. f.) Bojer, Hort. 

Maurit. 224. 1837. 

Fig. 63. E-F 

BASIONYM: Convolvulus nervosus Burm. f. 

SYNONYMS: Rivea nervosa (Burm. f.) H. Hallier 

Argyreia speciosa (L.f.) Sweet 

Twining liana, without latex, attaining more 

than 5 m in length. Stems cylindrical, densely 

canescent when young, becoming glabrous when 

mature. Leaves alternate; blades simple, 12-22 

(27) × 8.5-20 cm, cordiform, coriaceous, the apex 

obtuse to rounded, sometimes mucronate, the base 

cordiform, the margins entire; upper surface dark, 

dull, glabrous, with the venation slightly sunken; 

lower surface densely canescent, with the pinnate 

venation prominent; petioles 5-15 cm long, 

densely canescent. Flowers few, in axillary simple 

or double dichasial cymes; bracts foliaceous, 

ovate, acuminate, 2-5 cm long, canescent on the 

lower surface, forming an involucre at the base 

of the dichasia; peduncles densely canescent, up 

to 15 cm long. Calyx crateriform, canescent, 

accrescent, of 5 ovate sepals, obtuse, mucronate, 

1.5-2 cm long; corolla lavender, dark violet in 

the throat, infundibuliform, 6-6.5 cm long, 

canescent outside, the limb with 5 slightly 

prominent, rounded lobes; stamens and style not 

exserted. Indehiscent fruits ovoid, 1-1.5 cm long, 

puberulous, the pericarp thick, subtended by the 

subwoody, persistent sepals, slightly shorter than 

the fruit; seeds light brown, ca. 5 mm long, 

densely appressed short-pubescent, with two sides 

flat and one convex, the hilum forming a navel. 

Phenology: Flowering from April to August, 

collected in fruit in November. 

Status: Exotic, cultivated in Puerto Rico and 

the Virgin Islands, uncommon. 

Distribution: Native to India, cultivated and 

naturalized in the Antilles and throughout the 

tropics. 

Erect or prostrate herbs, shrubs, or twining vines. Leaves alternate, petiolate; blades simple, entire 

or lobed; stipules absent. Flowers bisexual, actinomorphic, solitary or clustered in involucrate heads. 

Calyx of 5 equal or unequal sepals; corolla campanulate or infundibuliform, the limb slightly 5-lobed; 

stamens 5, exserted; ovary superior, with a solitary style and 2 filiform stigmas. Fruit a 4-valvate 

capsule, with each of the valves opening in two; seeds 4 per fruit, with two sides flat and one convex. 

A genus of about 250 species, mostly native to the Old World.


1. Convolvulus nodiflorus Desr. in Lam., Encycl. 

3: 557. 1792. 

Fig. 63. G-K 

SYNONYMS: Jacquemontia nodiflora (Desr.) G. Don 

Convolvulus albiflorus Vahl 


in length. Stems cylindrical, tomentose, with 

white or golden hairs, trifurcate, producing watery 

latex. Leaves alternate; blades simple, 2-4.5 × 1- 

3 cm, lanceolate, ovate, elliptical, or rounded, 

chartaceous, the apex obtuse, acute, acuminate, 

or retuse, usually mucronate, the base rounded, 

truncate, or cordiform, the margins entire; upper 

surface dull, with the midvein slightly prominent, 

sparsely covered with 2- or 3-fid trichomes; lower 

surface densely pubescent with white or golden, 

bi- or trifid hairs, with the pinnate venation 

prominent; petioles slender, 0.5-1.5 cm long. 

Flowers few, in axillary cymes; peduncle densely 

4. IPOMOEA 

tomentose, up to ca. 1 cm long. Calyx light green, 

crateriform, sepals ovate, equal, not accrescent, 

3-3.5 mm long, rounded at the apex; corolla white, 

infundibuliform, 1.2-1.5 cm long, the limb with 

5 obtuse lobes; stamens white, the filaments 8-10 

mm long; ovary white, the stigmas exserted. 

Capsules ovoid, ca. 5 mm long, the pericarp thin, 

light brown; seeds brown, ca. 3 mm long, with 2 

sides flat and one convex, puberulous. 


October to April. 


Distribution: In thickets along the coastal 

zone. Also on Caja de Muerto, Culebra, Desecheo, 

Isla Ramos, Vieques, Guana Island, St. Croix, St. 

John, St. Thomas, Tortola, and Virgin Gorda; 

throughout the Antilles, Central America, and 


Public forests: Boquerón, Ceiba, Guajataca, 

Guánica, and Susúa. 

Twining, creeping vines or rarely shrubs, usually with abundant milky latex. Leaves alternate, 

petiolate; blades simple, entire, lobed, palmately or pinnately compound; stipules absent. Flowers 

bisexual, actinomorphic, solitary or in axillary simple or dichasial cymes; peduncles slender or thick, 

shorter or longer than the petioles. Calyx of 5 equal or unequal sepals, free to the base; corolla 

campanulate, infundibuliform, or hypocrateriform, the limb entire to deeply 5-lobed; stamens 5, inserted 

or rarely exserted, the filaments usually unequal, the anthers lanceolate; ovary superior, 2-4-locular, 

the style solitary with a subglobose or biglobose stigma. Fruits capsular, 4-valvate, ovoid or globose; 

seeds 4 per fruit, with two sides flat and one convex, glabrous or pubescent. A pantropical genus of 


Key to the species of Ipomoea 

1a. Corollas hypocrateriform ................................................................................................................... 2 

1b. Corollas infundibuliform or campanulate ......................................................................................... 9 

2a. Corolla white ...................................................................................................................................... 3 

2b. Corolla cardinal red, red-orange or bright pink ............................................................................... 5 

3a. Leaves pinnatisect ......................................................................................................15. I. quamoclit 

3b. Leaves entire, simple. ........................................................................................................................ 4 

4a. Sepals lanceolate, aristate and revolute at the apex; corolla 10-12 cm long, the tube light green 

inside, the limb with 5 light green lines forming a star; stamens exserted ........................ 1. I. alba 

4b. Sepals ovate, concave, obtuse or rounded at the apex; corolla 5-7 cm long, the tube white, with 

cardinal red fringes inside at the base; stamens not exserted .................................... 23. I. violacea 

5a. Corolla cardinal red ........................................................................................................................... 6 

5b. Corolla bright pink or red-orange ..................................................................................................... 7

162 



6a. Leaves entire, reniform, with the apex retuse, 0.3-1.8 cm long ................................ 18. I. steudelii 

6b. Leaves pinnatisect, 2-10 cm long ..............................................................................15. I. quamoclit 

7a. Corolla bright pink; sepals pink; leaf with the base truncate, obtuse, or sometimes subcordiform.........8 

7b. Corolla red-orange; sepals green; leaf cordiform at the base................................... 5. I. hederifolia 

8a. Corolla with the limb entire or with 5 shallow, ovate, and expanded lobes ...... 10. I. microdactyla 

8b. Corolla with the limb deeply 5-lobed, the lobes oblong and reflexed ........................16. I. repanda 

9a. Leaves palmately compound............................................................................................................ 10 

9b. Leaves simple, entire, or trilobed .................................................................................................... 11 

10a. Corolla bright pink or crimson, thick, ca. 6 cm long; calyx smooth; peduncle thick, straight ...... 

............................................................................................................................ 6. I. horsfalliae 

10b. Corolla pale violet or purple, membranaceous, 1.5-2.5 cm long; calyx verrucose; peduncle filiform, 

spirally twisted .............................................................................................................. 24. I. wrightii 

11a. Plants creeping or sometimes the younger portions of the stem twining, producing roots in the 

area of the nodes that are in contact with the soil .......................................................................... 12 

11b. Plants ascending, twining, rarely producing roots at the nodes (I. setifera) ............................... 14 

12a. Plants strictly creeping; common on sandy coasts of the littoral zone ........................................ 13 

12b. Plants with the basal portion creeping, the younger portions ascending, twining; plants cultivated 

or escaped from cultivation .............................................................................................. 2. I. batatas 

13a. Corolla pink or lavender; leaves oblong to rounded, the apex deeply emarginate and 

mucronate........................................................................................................ . 13. I. pes-caprae 

13b. Corolla white with a yellow center; leaves oblong to lanceolate, the apex emarginate and mucronate 

.............................................................................................................................. 7. I. imperati 

14a. Leaves reniform or lyrate, < 1.5 cm long, clustered on short axillary branches; petioles usually 

longer than the leaf blade ............................................................................................... 4. I. eggersii 

14b. Leaves not reniform or lyrate, alternate, not clustered on short axillary branches ..................... 15 

15a. Corolla yellow with the center purple ...................................................................... 12. I. ochracea 

15b. Corolla of other colors ................................................................................................................... 16 

16a. Plant with at least some or all of the leaves trilobed .................................................................... 17 

16b. Leaves entire, never lobed ............................................................................................................. 21 

17a. Corolla 1.4-1.6 cm long ................................................................................................ 22. I. triloba 

17b. Corolla 1.8 cm long or longer ....................................................................................................... 18 

18a. Calyx glabrous or puberulous; corolla 5-7 cm long, violet to violet-pink .................... 8. I. indica 

18b. Calyx hirsute; corolla 1.5-4.5 cm long, lilac, pale blue, whitish, pink, or violet-blue ............... 19 

19a. Corolla 1.5-3 cm long; inflorescences of double dichasia resembling a head; bracts 1.5-3 cm long, 

forming an involucre at the base of the inflorescences .................................................... 9. I. meyeri 

19b. Corolla 4-4.5 cm long; inflorescences of simple dichasia or sometimes the flower solitary; bracts 

4-4.5 mm long, not forming an involucre ....................................................................................... 20

164 


20a. Sepals with apices long-acuminate, elongate, much longer than the body ....................... 11. I. nil 

20b. Sepals with apices acute or slightly acuminate .......................................................14. I. purpurea 

21a. Leaves elliptical, oblong, or lanceolate, hastate or auriculate at the base ........... 19. I. tenuissima 

21b. Leaves ovate, broadly ovate, or oblong, cordiform or truncate at the base ................................. 22 

22a. Corolla pink or violet-pink; sepals with the margins not hyaline ............................................... 23 

22b. Corolla blue, turning violet when mature; sepals with hyaline margins .................. 22. I. tricolor 

23a. Sepals with three parallel veins, prominent, like a keel ............................................17. I. setifera 

23b. Sepals without prominent veins .................................................................................................... 24 

24a. Young stems 4-winged; sepals obtuse or retuse at the apex ....................................... 3. I. calantha 

24b. Young stems cylindrical; sepals apiculate at the apex .............................................. 20. I. tiliacea 

1. Ipomoea alba L., Sp. Pl. 161. 1753. 

Fig. 64. A-E 

SYNONYMS: Calonyction aculeatum (L.) House 

Calonyction bona-nox (L.) Bojer 

Bejuco de puerco, Bejuco de vaca, Claro de 

luna, Flor de luna 

Twining vine, slightly woody, climbing or 

creeping, 5-20 m in length, with scarce milky or 

watery latex. Stems cylindrical, slender, fragile, 

glabrous, green, with the nodes mulberry-colored 

and with spiniform projections. Leaves alternate; 

blades simple or sometimes 3-5-lobed, 7-18 × 8- 

16 cm, broadly ovate, chartaceous or 

subcoriaceous, glabrous, the apex acute or 

acuminate, the base cordiform, the margins 

undulate and slightly revolute, entire or 3-7lobate; 

upper surface dark green, dull; lower 

surface pale green, with prominent venation; 

petioles 6-24 cm long, fragile, usually mulberrycolored, 

with the base somewhat broadened. 

Flowers fragrant, nocturnal, solitary or few in 

axillary racemes; peduncle cylindrical, 4-15 cm 

long. Calyx green, the sepals 2.5-3 cm long, 

lanceolate, long-aristate and revolute at the apex; 

corolla white, hypocrateriform, 10-12 cm long, 

the tube narrow, light green inside, the limb 5lobed, 

9-12 cm in diameter, with 5 light green 

lines outside forming a star; stamens and stigmas 

white, exserted. Capsules ovoid-conical, 2-3 cm 

long, with a persistent style; seeds 4, dark brown 

to black, glabrous, ca. 1 cm long, with two sides 

flat and one convex. 

Phenology: Flowering and fruiting 

sporadically throughout the year. 

Status: Possibly native, rather common. 

Distribution: In moist areas, along roads and 

in pastures at middle and lower elevations. Also 

on Vieques. Cultivated throughout the tropics but 

native to continental tropical America. 

Public forests: El Yunque, Guánica, Maricao, 


2. Ipomoea batatas (L.) Lam., Encycl. Méth. 6: 

14. 1804. 

Fig. 64. F-G 

BASIONYM: Convolvulus batatas L. 

Batata, Camote, Sweet potato 

Creeping vine, with the younger portions 

twining and ascending, 3-4 m in length, with 

abundant milky latex. Roots tuberous, edible. 

Stems cylindrical, slender, flexible, glabrous or 

pubescent, usually producing aerial roots in the 

area of the nodes. Leaves alternate; blades simple 

or 3-7-lobed, 6-13 × 6-10 cm, when simple 

broadly ovate, deltoid, chartaceous, glabrous, the 

apex acute, obtuse, or acuminate, mucronulate, 

the base cordiform, lyrate, or truncate, the margins 

entire; upper surface dark green, dull; lower 

surface pale green, with prominent venation; 

petioles 5-12 cm long, glabrous. Flowers few, in 

axillary dichasial cymes; peduncle cylindrical, 7- 

12 cm long. Calyx green, not accrescent, the 

sepals oblong, unequal, 8-12 mm long, aristate 

at the apex; corolla violet-pink, infundibuliformcampanulate, 

3-4.5 cm long, the limb ca. 4 cm in 

diameter, with 5 rounded lobes; stamens and 

stigmas not exserted. Capsules depressed-globose,


166 


ca. 5 mm long, pilose; seeds black, dull, 

puberulous, ca. 3.5 mm long, with two sides flat 

and one convex. 


November to January. 



Distribution: Widely cultivated in Puerto Rico 

and the Virgin Islands. Native to the Neotropics 

but cultivated throughout the tropics and 

subtropics. 

3. Ipomoea calantha Griseb., Cat. Pl. Cub. 202. 

1866. 

Fig. 65. A-C 

Twining vine, slightly woody, 3-4 m in length. 

Young stems 4-winged, puberulous, cylindrical 

and striate when mature. Leaves alternate; blades 

simple, 5.5-14 × 4.7-10.5 cm, broadly ovate, 

chartaceous, the apex acute, obtuse, or acuminate, 

mucronulate, the base cordiform or truncate, the 

margins entire; upper surface dark green, dull, 

glabrous, with a prominent midvein; lower surface 

pale green, with prominent venation and 

puberulous; petioles 3-13 cm long, puberulous. 

Flowers solitary or few, in simple dichasial and 

axillary cymes; peduncle 4-13.5 cm long. Calyx 

green, not accrescent, the sepals oblong, 

puberulous, unequal, 8-12 mm long, the 2 outer 

ones shorter than the 3 inner ones, obtuse at the 

apex, the inner ones retuse at the apex; corolla 

violet-pink, infundibuliform, 6.5-7 cm long, the 

limb ca. 5 cm in diameter, with 5 rounded lobes; 

stamens and stigmas not exserted. Capsules ovoidelongate, 

1-1.3 cm long, light brown, glabrous; 

seeds reddish brown, ellipsoid, ca. 8 mm long, 

with a row of long silky hairs at the margins. 

Phenology: Apparently flowering and fruiting 

sporadically throughout the year. 


Distribution: In thickets and disturbed areas 

in southern Puerto Rico. Also in Cuba, Hispaniola, 

Colombia, and Venezuela. 

Public forest: Guánica (according to Quevedo 

et al. 1990) and Maricao (according to Cedeño, 

1991). 

4. Ipomoea eggersii (House) D. F. Austin, Ann. 


Fig. 65. D-E 

BASIONYM: Exogonium eggersii House 

SYNONYM: Ipomoea arenaria sensu Urb. 

Twining vine, slightly woody, much branched 

from the base, attaining 5 m in length, with 

abundant milky latex. Tuberous roots subglobose 

(resembling a beet). Stems cylindrical, slender, 

reddish brown, glabrous. Leaves alternate, 

clustered on short axillary branches; blades 6-15 

× 4-13 mm, reniform, lyrate or lobed, chartaceous, 

glabrous, the apex retuse, mucronate, the base 

cordiform, truncate, or rounded, the margins 

sinuate; upper surface dark green, dull, glabrous, 

with the veins flat; lower surface pale green, with 

the midvein thick and prominent; petioles 5-30 

mm long, glabrous, slender. Flowers solitary at 

the end of short branches; bracts oblong, minute. 

Calyx pale green, not accrescent, the sepals ovate 

to rounded, 5-6 mm long, glabrous, unequal; 

corolla infundibuliform, 4-5 cm long, the tube pale 

green outside, white inside, the limb pink or pale 

violet, 3-4 cm in diameter, with 5 obtuse lobes; 

stamens and stigmas white, not exserted. Capsules 

ellipsoid, ca. 1.5 cm long, light brown; seeds 4 

per fruit, dark brown, ca. 6 mm long, with a tuft 

of long silky hairs at the margins. 

Phenology: Collected in flower from January 

to April and in fruit from January to March. 


Distribution: In thickets and coastal forests 

on Vieques, St. Croix, St. John, St. Thomas, 

Tortola, Virgin Gorda, and Water Island; the 

Lesser Antilles. 

Commentary: Ipomoea eggersii seems to be 

closely related to I. steudelii, because both species 

share numerous morphological characters. The 

leaves and seeds are almost indistinguishable. 

However, the corollas of both species are 

drastically different and their distribution does not 

overlap: I. eggersii is distributed from the Virgin 

Islands to the Lesser Antilles, while I. steudelii is 

found on Culebra, Vieques, Puerto Rico, and 

probably in Hispaniola. 

5. Ipomoea hederifolia L., Syst. Nat. ed. 10, 2: 

925. 1759. 

Fig. 65. F-H 

SYNONYM: Quamoclit coccinea sensu Urb.


Twining vine, slightly woody, attaining 5 m 

in length, with watery latex. Stems cylindrical, 

slender, glabrous. Leaves alternate; blades 4-13 

× 4.5-11 cm, ovate or deeply 3-5-lobed, 

chartaceous, glabrous, the apex of the lobes 

acuminate, the base cordiform or sagittate; upper 

surface dark green, dull, glabrous, punctate, with 

the veins slightly prominent; lower surface pale 

green, with the primary and secondary veins 

prominent; petioles 4.5-9 cm long, glabrous, 

slender. Flowers in dichasial cymes with one-sided 

racemose branches. Calyx pale green, not 

accrescent, the sepals 5-8 mm long, lanceolate, 

aristate at the apex; corolla hypocrateriform, 

bright red or red-orange, 4-4.5 cm long, the limb 

ca. 2.5 cm in diameter, with 5 obtuse lobes, 

shallow; stamens and stigmas white or pink, 

exserted. Capsules globose, ca. 8 mm in diameter, 

the pericarp thin; seeds 4 per fruit, light brown, 

ca. 5 mm long, pubescent. 


October to January. 

Status: Apparently native, moderately 

common. 

Distribution: In thickets, forest margins, and 

disturbed areas of the coast or the interior of Puerto 

Rico. Also on Vieques, St. Croix, St. John, and 

St. Thomas; the Antilles, Central America, and 

South America, introduced in the Old World 

tropics. 


6. Ipomoea horsfalliae W. Hook., Bot. Mag. t. 

3315. 1854. 

Fig. 66. A-B 

Twining vine, slightly woody, attaining more 

than 10 m in length, with watery latex. Stems 

cylindrical, slender, glabrous. Leaves alternate, 

palmately compound; leaflets 5, elliptical, 5.5-10 

× 1.6-2.7 cm (the central leaflet larger than the 

lateral ones), chartaceous or membranaceous, 

glabrous, the apex acuminate or long-acuminate, 

the base acute or acuminate; upper surface 

glabrous, with the midvein slightly prominent; 

lower surface glabrous, punctate, with a 

prominent midvein; petioles 4.5-6 cm long, 

glabrous, slender. Flowers in double dichasial 

cymes. Calyx with a reddish tinge, not accrescent, 

the sepals unequal, 7-10 mm long, elliptical to 

ovate, obtuse at the apex, the outer ones shorter 

than the inner ones; corolla infundibuliform, 

violet-red or bright pink, 4-6.3 cm long, the limb 

3.5-4.5 cm in diameter, with 5 rounded lobes; 

stamens and stigmas not exserted. Capsules 

ellipsoid, ca. 8 mm in diameter, the pericarp thin; 

seeds 4 per fruit, light brown, ca. 5 mm long, 

glabrous. 


to January. 

Status: Possibly exotic and naturalized, 

uncommon. 


Central and in the zone of mogotes. This species 

was originally described based on material 

cultivated in the gardens at Kew, but of uncertain 

origin. The species is considered by some as native 

to Jamaica and introduced in other places due to 

its showy flowers. 

Public forests: Maricao and Toro Negro. 

Commentary: This species forms hybrids with 

I. repanda and produces plants with 5-digitate 

leaves and flowers very similar to those of I. 

repanda. These are cultivated throughout the 

tropics. 

7. Ipomoea imperati (Vahl) Griseb., Cat. Pl. Cub. 

203. 1866. 

Fig. 66. C-E 

BASIONYM: Convolvulus imperati Vahl 

SYNONYMS: Ipomoea stolonifera J. F. Gmel. 

Convolvulus littoralis L. 

Ipomoea littoralis (L.) Boiss, non Blume 

Erect herb or creeping or sometimes ascending 

vine, slightly woody, attaining 2 m in length, with 

milky latex. Stems cylindrical, slender, glabrous, 

producing adventitious roots in the area of the 

nodes. Leaves alternate, usually clustered on short 

axillary branches; blades simple, 2-10 × 1-2.5 cm, 

lanceolate, oblong, or deltoid, sometimes 

subsagittate, lyrate, or deeply trilobed, fleshy when 

fresh, chartaceous on drying, glabrous, the apex 

obtuse, rounded, or sometimes emarginate, 

mucronate, the base cuneiform, truncate, 

subcordiform to subsagittate; upper surface dull, 

glabrous, with the veins flat; lower surface 

glabrous, with the primary and secondary veins 

prominent; petioles 1-6.5 cm long, sulcate, pilose 

at the apex. Flowers solitary or in ascending 

dichasial cymes; peduncles 1.5-4.5 cm long. Calyx

168 



green, not accrescent, the sepals 10-15 mm long, 

elliptical, aristate at the apex, the margin hyaline, 

the two outer ones slightly longer than the inner 

ones, glabrous; corolla white, with a yellow center, 

infundibuliform, 2.5-5 cm long, the limb 3-6 cm 

in diameter, with 5 shallow, obtuse lobes; stamens 

and stigmas not exserted. Capsules globose or 

conical, 1.2-1.5 cm long; seeds 1-4 per fruit, ca. 

5 mm long, lanate-pubescent. 


to March. 


Distribution: Along the sandy coasts of the 

northern littoral zone. Also on Culebra and 

Vieques. A pantropical species, distributed along 

the sandy coasts of the littoral zone. 

Public forest: Guánica and Piñones 

8. Ipomoea indica (Burm.) Merr., Int. Rumph. 

Herb. Amb. 445. 1917. 

Key to the varieties 

1a. Plants glabrous; leaves usually trilobed 

……………. I. indica var. acuminata 

1b. Plants appressed-pubescent; leaves cordiform, 

not lobed............…. I. indica var. indica 

8a. Ipomoea indica (Burm.) Merr. var. 

acuminata (Vahl) Fosberg, Bot. Not. 129: 38. 

1976. 

Fig. 66. F-I 

BASIONYM: Convolvulus acuminatus Vahl 

SYNONYMS: Ipomoea cathartica Poir. 

Ipomoea portoricensis (Spreng.) G. Don 

Bejuco de gloria 


with abundant milky latex. Stems slender, 

glabrous, cylindrical, smooth, greenish. Leaves 

alternate; blades 6-11 × 4-8 cm, glabrous, ovate 

or trilobed, chartaceous, the apex acute or shortacuminate, 

the base cordiform, the margins entire; 

upper surface with the venation slightly 

prominent; lower surface with prominent 

venation, glabrous or puberulous; petioles 6-12 

cm long, cylindrical, glabrous. Flowers solitary 

or in simple dichasial cymes, axillary. Calyx 

green, accrescent, glabrous, the sepals subequal, 

15-18 mm long, lanceolate, chartaceous, long- 

acuminate at the apex; corolla violet-pink with 

the center darker, infundibuliform, 5-7 cm long, 

the limb ca. 7 cm in diameter, with 5 rounded 

lobes; stamens not exserted; stigmas white, 

slightly exserted. Capsule globose, 10-15 cm in 

diameter, subtended by the accrescent sepals, 

subwoody; seeds black, elliptic-triangular, ca. 5 

mm long, smooth, with a line of hairs at the 

margins. 

Phenology: Flowering from July to April and 

fruiting from December to July. 



littoral zone and in moist areas at middle 

elevations. Also on Cayo Santiago, Culebra, 

Desecheo, Vieques, St. Croix, St. John, St. 

Thomas, and Tortola; throughout the Antilles and 


Public forests: Cambalache, Ceiba, El Yunque, 

Guilarte, Piñones, Río Abajo, Tortuguero, and 

Vega. 

8b. Ipomoea indica var. indica 

Herbaceous vine, twining. Stems appressedpubescent, 

cylindrical. Leaves alternate; blades 

simple, 6-8 × 5-6 cm, ovate, chartaceous, the apex 

acuminate, the base cordiform, the margins entire; 

upper surface glabrous; lower surface with the 

venation slightly prominent, appressed-pubescent; 

petioles 5-6 cm long, cylindrical. Flowers solitary 

or in simple dichasial cymes, axillary. Calyx 

green, accrescent, appressed-pubescent, sepals 

subequal, ca. 2 cm long, lanceolate, chartaceous, 

long-acuminate at the apex; corolla violet-pink, 

infundibuliform, ca. 5 cm long, the limb ca. 3 cm 

in diameter; stamens not exserted; stigmas white, 

slightly exserted. Capsule globose, 10-15 mm in 

diameter, subtended by the accrescent sepals, 

subwoody; seeds black, elliptic-triangular, ca. 5 

mm long, smooth, with a line of hairs along the 

margins. 


Status: Native, very rare. 

Distribution: Known from a single collection 

in Piedras Chiquitas in Coamo, Puerto Rico. This 

variety predominates on the coasts of the Indo- 

Pacific region, and apparently is also found from 

Mexico to Brazil and Argentina.

170 



9. Ipomoea meyeri (Spreng.) G. Don, Gen. Syst. 

4: 275. 1838. 

Fig. 67. A-E 

BASIONYM: Convolvulus meyeri Spreng. 


Stems slender, glabrous or pubescent, cylindrical, 

with scarce milky latex. Leaves alternate; blades 

simple, 4-9.5 × 3-7 cm, ovate, broadly ovate, 

deltate, lyrate, or trilobed, membranaceous, the 

apex and the lobes acute or acuminate, the base 

cordiform or sagittate, the margins entire; upper 

surface with the venation slightly prominent, 

strigose; lower surface with prominent venation, 

glabrous; petioles 2.5-6.5 cm long, slender, 

hirsute, especially at the apex. Flowers solitary, 

or 2 or 3 in axillary cymes, involucrate; peduncle 

2-17 mm long, strigose to glabrescent; bracts of 

the involucre oblong, 1.5-2 cm long, sparsely 

hirsute. Calyx green, accrescent, the sepals 

unequal, 1.2-1.7 cm long, lanceolate, hirsute, 

especially near the base, long-acuminate at the 

apex; corolla pale blue, pale violet, or whitish, 

with the center white, infundibuliform, 1.5-3 cm 

long, the limb 2.5-3 cm in diameter, with 5 

rounded lobes; stamens and stigmas not exserted. 

Capsules globose, ca. 8 mm in diameter, with the 

sepals persistent, erect, and accrescent; seeds 4 

per capsule, reddish brown, elliptic-cuneiform, ca. 

5 mm long, finely pubescent. 


to January and in fruit from December to January. 


Distribution: Common in southern and 

western Puerto Rico. Also in the Antilles, Central 

America, and northern South America. 

Public forests: Guánica. 

10. Ipomoea microdactyla Griseb., Cat. Pl. Cub. 

204. 1866. 

Fig. 67. F-J 

SYNONYMS: Exogonium microdactylum (Griseb.) House 

Ipomoea repanda var. microdactyla (Griseb.) Powell 

Herbaceous vine, twining, 1.5-2 m in length, 

without latex. Stems slender, cylindrical, glabrous, 

with spinescent projections on the more mature 

portions. Leaves alternate; blades simple, 3-10 cm 

long, glabrous, ovate, lanceolate, or deeply 3-7lobed, 

the apex and the lobes obtuse to acuminate, 

mucronate, the base cordiform, cuneate or 

truncate, the margins undulate; upper surface with 

the venation flat; lower surface punctate or 

foveolate, with prominent venation; petioles 1.5- 

4.5 cm long, slender, glabrous. Flowers 2-3, in 

axillary cymes; peduncles 0.8-4.5 cm long. Calyx 

pink, the sepals unequal, 4.5-7 mm long, rounded, 

glabrous, with the margins hyaline; corolla bright 

pink, hypocrateriform, 3-4.5 cm long, the limb 

with 5 ovate lobes, obtuse, expanded; stamens and 

stigmas exserted. Capsules conical, stramineous, 

1-1.5 cm long, apiculate; seeds 4 per capsule, 

reddish brown, 5-6 mm long, with two sides flat 

and one convex, with a row of silky hairs longer 

than the seed at the margins and the apex. 




Distribution: On the central plateau of Mona 

Island. Also in the Bahamas, Cuba, and the United 

States (Florida). 

Public forest: Mona. 

11. Ipomoea nil (L.) Roth, Catal. Bot. 1: 36. 1797. 

Fig. 67. K-O 

BASIONYM: Convolvulus nil L. 

Twining vine, attaining 5 m in length, with 

scarce watery latex. Stems cylindrical, slender, 

strigose. Leaves alternate; blades simple, 4.5-13.3 

× 5.7-15 cm, chartaceous, strigose, deeply 

trilobed, the lobes ovate or lanceolate, acuminate 

at the apex, the base cordiform, sagittate, or deeply 

cordiform; upper and lower surface with the veins 

slightly prominent; petioles 2-8 cm long, strigose. 

Flowers solitary or in simple dichasia, axillary; 

peduncles as long as the petioles or slightly 

shorter; bracts subulate, 4-5 mm long, not forming 

an involucre. Calyx green, not accrescent, of 5 

equal sepals, 2-3 cm long, lanceolate, chartaceous, 

long-acuminate, externally strigose, hirsute 

outside on the basal portion; corolla pale blue, 

lavender, or purple with the center white, 

infundibuliform, 4-4.5 cm long, the limb with 

shallow, rounded lobes; stamens and stigmas 

white, not exserted. Capsule subglobose, ca. 5 mm 

long, the pericarp thin, color yellowish brown, 

glabrous, with the chartaceous sepals persistent 

at the base; seeds 4 per fruit, ca. 4 mm long, dark 

brown, finely pubescent. 


April and fruiting from January to April.

172 



Status: Exotic, naturalized. 

Distribution: Common in southwestern Puerto 

Rico. Also on Desecheo, Vieques, St. Croix, St. 

John, St. Thomas, and Tortola; native to Mexico 

but found distributed throughout the tropics. 

Public forest: Boquerón and Guánica. 

12. Ipomoea ochracea (Lindl.) G. Don, Gen. Hist. 

4: 270. 1838. 

Fig. 68. A-C 

BASIONYM: Convolvulus ochraceus Lindl. 


in length, with scarce watery latex. Stems 

cylindrical, slender, pubescent. Leaves alternate; 

blades simple, 3-10 × 2.5-7.5 cm, ovate, 

chartaceous, glabrous except for some hairs on 

the veins, the apex acuminate and usually 

mucronate, the base cordiform, the margins entire 

or sinuate; upper and lower surface with the veins 

slightly prominent, the lower surface usually 

glaucous; petioles 2-6 cm long, pubescent. 

Flowers solitary or in double dichasia, axillary; 

peduncles slightly longer than the petioles; bracts 

ovate, ca. 1.6 mm long. Calyx green, not 

accrescent, of 5 subequal sepals, 5-7 mm long, 

chartaceous, ovate or oblong-ovate, glabrous, 

punctate; corolla pale yellow, with the base of the 

tube purple inside, infundibuliform, 3-4 cm long, 

the limb with 5 shallow, rounded lobes; stamens 

and stigmas white, not exserted. Capsule ovoid, 

glabrous, with a thin pericarp, 1.3-1.6 cm long, 

stramineous, with the sepals persistent, not 

accrescent at the base; seeds 4 per fruit, 4-5 mm 

long, black, dull, glabrous. 



Status: Exotic, naturalized. 

Distribution: Common in southwestern Puerto 

Rico. Also on St. Croix and St. John; probably 

native to tropical Africa, widely cultivated 


Public forests: Guánica, Maricao, and Susúa. 

13. Ipomoea pes-caprae (L.) R. Br. in Tuckey, 

Narr. Exped. Zaire 477. 1818. 

Fig. 68. G-H 

BASIONYM: Convolvulus pes-caprae L. 

Bejuco de puerco playero, Beach moning glory, 

Goat foot 

Vine, creeping or sometimes ascending, up to 

10 m in length, with scarce milky latex. Stems 

cylindrical, glabrous, woody in the older portions, 

the young portions herbaceous, producing 

adventitious roots in the area of the nodes. Leaves 

alternate; blades simple, 6.5-12 × 4-7.3 cm, 

oblong, reniform to subrounded, coriaceous, 

glabrous, the apex emarginate and mucronate, the 

base rounded, cordiform, or truncate, the margins 

entire; upper surface with the veins flat, scarcely 

visible; lower surface with the veins slightly 

prominent; petioles 3-8 cm long, glabrous, thick, 

slightly compressed on the upper surface, swollen 

at the base. Flowers solitary or in simple dichasia, 

axillary; peduncles shorter than the petioles; 

bracts subulate, ca. 5 mm long. Calyx yellowish 

green, not accrescent, the sepals unequal, 1.2-1.4 

cm long, coriaceous, ovate, the apex rounded and 

mucronate; corolla pink or lavender, 

infundibuliform, 4.5-5 cm long, the limb up to 6 

cm in diameter, with 5 rounded lobes; stamens 

and stigmas not exserted. Capsule ovoid to 

globose, 1.5-1.8 cm long, light brown, with a thin 

pericarp and with the sepals persistent and 

reflexed at the base; seeds 4 per fruit, ca. 8 mm 

long, dark brown, pubescent, with two sides flat 

and one convex. 




Distribution: On the sandy coasts of the littoral 

zone, probably on all the islands and keys of 

Puerto Rico and the Virgin Islands. A pantropical 

species, very common on the sandy coasts of the 

littoral zone. 

Public forests: Guánica, Mona, and Piñones. 

14. Ipomoea purpurea (L.) Roth, Bot. Abh. 27. 

1787. 

Fig. 68. D-F 

BASIONYM: Convolvulus purpureus L. 


Stems cylindrical, slender, pilose or hirsute. 

Leaves alternate; blades simple, 2-10 × 2-10 cm, 

cordiform or deeply trilobed, the lobes ovate or 

lanceolate, chartaceous, strigulose on both 

surfaces, the apex acuminate, the base cordiform, 

the margins entire or slightly sinuate, ciliate; 

upper and lower surface with the veins slightly

174 



prominent; petioles 2.5-6 cm long, slender, 

strigulose, sulcate. Flowers solitary or in simple 

dichasia, axillary; peduncles longer than the 

petioles; bracts subulate, ca. 3 mm long, not 

forming an involucre. Calyx green, of 5 subequal 

sepals, 8-16 mm long, chartaceous, oblonglanceolate, 

the outer ones slightly broader than 

the inner ones, acute at the apex, hirsute outside 

on the basal portion; corolla blue, purple, pink, 

or with lines (forming a star) of these colors on a 

white background, infundibuliform, 4-4.5 cm 

long, the throat white, limb with shallow, rounded 

lobes; stamens and stigmas pink, not exserted. 

Capsule depressed-globose, 9-10 mm in diameter, 

glabrous, the pericarp thin, with the chartaceous 

sepals persistent at the base; seeds 4 per fruit, 

pyriform, 3-4 mm long, black, glabrous. 

Phenology: Flowering from June to January. 

Status: Exotic, uncommon. 

Distribution: Along roads and in disturbed 

places. Also on Vieques, St. Croix, St. Thomas, 

and Tortola. Probably native to Mexico, but found 

distributed throughout the tropics. 

15. Ipomoea quamoclit L., Sp. Pl. 159. 1753. 

Fig. 69. A-B 


with watery latex. Stems cylindrical, slender, 

glabrous. Leaves alternate or clustered on short 

axillary branches; blades deeply pinnatisect, ovate 

or elliptical in outline, with 9-19 pairs of linear 

segments, alternate or opposite, 1-9 cm long, 

chartaceous, puberulous or glabrous; petioles not 

evident. Flowers solitary or 2-6 in cymes; 

peduncles as long as or longer than the leaves; 

bracts ovate, ca. 1 mm long. Calyx green, not 

accrescent, of 5 unequal sepals, 4-7 mm long, 

chartaceous, ovate, the outer ones shorter than 

the inner ones, glabrous, obtuse at the apex, with 

a mucro that is borne below the apex; corolla red 

(sometimes white), hypocrateriform, 2-3 cm long, 

the limb with deep lobes, deltate, obtuse; stamens 

and stigmas pink, exserted. Capsule ovoid to 

conical, with a thin pericarp, lepidote outside, 8- 

10 mm long; seeds 4 per fruit, cuneiform, 7-8 mm 

long, dark brown to black, with tufts of irregularly 

scattered hairs. 

Phenology: Flowering from August to 

February, collected in fruit in February. 


uncommon. 

Distribution: In disturbed areas. Cultivated on 

St. Croix and St. Thomas. Possibly native to 

Mexico, but today found distributed throughout 

the tropics. 

Public forests: Cambalache and Tortuguero. 

16. Ipomoea repanda Jacq., Enum. Syst. Pl. 13. 

1760. 

Fig. 69. C-F 

SYNONYM: Exogonium repandum (Jacq.) Choisy 

Bejuco rosado, Bejuco colorado, Mari-de-Lugo 


length, usually producing abundant milky latex. 

Roots tuberous, elongate or rounded. Stems 

slender, glabrous, cylindrical, striate when young, 

lenticellate on the mature portions. Leaves 

alternate; blades simple, 5-18 × 1.5-13 cm, 

glabrous, chartaceous, ovate, lanceolate, or 

sometimes trilobed, the apex and lobes acute or 

acuminate, mucronate, the base cordiform, 

truncate, or rounded, the margins revolute, entire 

or slightly undulate; upper surface dark green, 

shiny; lower surface pale green, punctate, with a 

prominent midvein; petioles 1.5-6 cm long, 

sulcate, glabrous. Flowers in compound dichasial 

cymes, pendulous; bracts deciduous. Calyx pink, 

crateriform, not accrescent, the sepals unequal, 

8-10 mm long, ovate, obtuse at the apex; corolla 

bright pink, hypocrateriform, 4-5.5 cm long, the 

limb with 5 deep lobes, oblong, obtuse, reflexed; 

stamens and stigmas pink or white, exserted. 

Capsules conical to ellipsoid, the pericarp thin, 

pale brown, 1.2-1.5 cm long, apiculate; seeds 4 

per capsule, black, 5-6 mm long, with two sides 

flat and one convex, with a row of silky hairs 

longer than the seed at the margins and the apex. 


the year, especially from June to January. 



Central, in the Sierra de Luquillo, and in the zone 

of mogotes. Also on St. John, St. Thomas, and 

Tortola; the Lesser Antilles. 

Public forests: Carite, El Yunque, Guajataca, 

Guilarte, Maricao, Río Abajo, and Toro Negro.

176 



17. Ipomoea setifera Poir. in Lam., Encycl. 6: 

17. 1804. 

Fig. 70. A-B 

SYNONYM: Ipomoea rubra (Vahl) Millsp. 

Bejuco de puerco, Gloria de la mañana, 

Wild morning glory 

Herbaceous vine, twining, climbing, or 

creeping, attaining 12 m or more in length, with 

abundant milky latex. Stems cylindrical, slender, 

pilose, sometimes with a pink tinge, usually with 

numerous aerial roots where they enter in contact 

with the soil. Leaves alternate; blades simple, 4.5- 

16 (24) × 2.7-6.4, glabrous, chartaceous, oblong 

or ovate, the apex obtuse, retuse and mucronate, 

the base cordiform or hastate, the margins 

revolute, undulate, usually with a pink or purple 

tinge; upper surface dark green with the midvein 

slightly prominent; lower surface pale green, 

minutely punctate, with prominent venation, 

conspicuously reddish; petioles 10-21 cm long, 

slender, glabrous, sulcate, usually reddish. 

Flowers in compound dichasial cymes, axillary; 

peduncle thick, shorter than the petioles; bracts 

ovate, 1.5-2 cm long. Calyx light green, 

crateriform, the sepals ovate, unequal, 12-17 mm 

long, with 3 parallel veins and prominent like a 

keel, the apex acuminate, the outer sepals longer 

than the inner ones; corolla violet-pink with the 

center darker, infundibuliform, 4-7 cm long, the 

limb 4-5 cm in diameter, with 5 rounded lobes; 

stamens pink, not exserted; stigmas pink, 

exserted. Capsule ovoid, ca. 1 cm in diameter, 

the pericarp thin, covered by the persistent sepals 

at the base; seeds 4 per fruit, with two sides flat 

and one convex, ca. 6 mm long, dark brown to 

black, tomentulose. 


April and fruiting from January to April. 


Distribution: In moist disturbed areas. Also 

on St. John; throughout the Antilles, continental 

tropical America, naturalized in western Africa. 

Public forests: Carite, Cambalache, Ceiba, El 

Yunque, Guilarte, Maricao, Río Abajo, Toro 


18. Ipomoea steudelii Millsp., Field Mus. Bot. 

2: 86. 1900. 

Fig. 70. C-G 

SYNONYMS: Exogonium arenarium Choisy 

Ipomoea arenaria (Choisy) Steud., non Roem. & Schult. 

Twining vine, slightly woody, much branched 

from the base, 2-3 m in length, with scarce milky 

latex (almost imperceptible). Stems cylindrical, 

slender, light brown, glabrous, striate. Leaves 

alternate, clustered on short axillary branches; 

blades simple, 3-18 × 1.3-14 mm, lyrate, reniform, 

deltate, or deeply 3-5-lobed, membranaceous, 

glabrous, the apex retuse, with the midvein 

projecting like a mucro, the base subcordiform, 

truncate, or cuneate, the margins slightly sinuate; 

upper surface dark green, dull, glabrous, with the 

veins flat; lower surface pale green, with the 

midvein thick and prominent; petioles 2-15 mm 

long, glabrous, slender. Flowers solitary, at the 

end of minute axillary branches, which present 

the scars of the deciduous leaves. Calyx green, 

the sepals glabrous, unequal, 7-11 mm long, ovate, 

obtuse at the apex; corolla crimson, 

hypocrateriform, 3.5-4 cm long, the limb ca. 2.5 

cm in diameter, with 5 more or less deep lobes, 

narrowly deltate; stamens and stigmas exserted. 

Fruits ellipsoid to subconical, 1-1.2 cm long, light 

brown; seeds 4 per fruit, dark brown, ca. 5.5 mm 

long, with a row of whitish silky hairs longer than 

the seed, at the apex and on the back. 

Phenology: Flowering from November to May 

and fruiting from January to July. 


Distribution: In thickets and dry forests of the 

coast and in the dry limestone zone. Also on 

Culebra, Culebrita, and Vieques; cited for 

Hispaniola, perhaps in error. 

Public forests: Boquerón, Guánica, Maricao, 


19. Ipomoea tenuissima Choisy ex DC., Prodr. 

9: 376. 1845. 

Fig. 71. A-C 


length. Stems cylindrical, slender, glabrous or 

puberulous. Leaves alternate; blades simple, 2-6 

× 0.3-1(2) cm, membranaceous, glabrous, 

lanceolate, the apex obtuse, mucronate, the base 

hastate or auriculate, the margins entire; upper 

and lower surface with the veins slightly 

prominent; lower surface punctate; petioles 1-2.2 

cm long, slender, glabrous. Flowers solitary or 

rarely in pairs, axillary; peduncles as long as or

178 


slightly longer than the petioles; bracts subulate, 

ca. 2.5 mm long. Calyx green, not accrescent, of 

5 subequal sepals, 6-7 mm long, chartaceous, 

elliptical, obtuse and aristate at the apex, ciliate 

at the basal margins, the midvein slightly 

prominent; corolla violet-pink, infundibuliform, 

ca. 3 cm long, the limb with shallow, rounded 

lobes; stamens and stigmas not exserted. Capsule 

ovoid-conical, ca. 5 mm long, glabrous, with a 

thin pericarp and with the chartaceous sepals 

persistent at the base; seeds 4 per fruit, cuneiform, 

ca. 3.7 × 2.5 mm, dark brown, finely pubescent 

at the margins. 


Status: Probably exotic.


Distribution: Known from Puerto Rico by a 

single collection from Joyuda (Cabo Rojo) in 1913. 

This species is known from Cuba, Hispaniola 

(whence it was described), and the United States 

(Florida). 

20. Ipomoea tiliacea (Willd.) Choisy in A.DC., 

Prodr. 9: 375. 1845. 

Fig. 71. D-H 

BASIONYM: Convolvulus tiliaceus Willd. 

Bejuco blanco, Bejuco de puerco, Goat foot, 

Wild potato, Wild vine 

Slightly woody vine, twining, 5-10 m in 

length, with abundant milky latex. Stems glabrous 

or pubescent, striate, cylindrical. Leaves alternate; 

blades simple, 5-20 × 4-14 cm, ovate, chartaceous, 

sometimes 3-5-lobed, the apex acuminate, the base 

cordiform, the margins undulate, ciliate; upper 

surface dark green, dull, usually with the venation 

sunken, glabrous except for the pilose midvein; 

lower surface pale green, with prominent 

venation, slightly pubescent; petioles 4-25 cm 

long, sulcate, puberulous. Flowers in double 

dichasia, axillary; peduncles shorter than the 

petioles. Calyx green, not accrescent, sepals 

unequal, 8-12 mm long, oblanceolate, glabrous, 

apiculate at the apex; corolla violet-pink to pale 

pink, with the center darker, infundibuliform, 5- 

6 cm long, the limb with 5 rounded lobes; stamens 

and stigmas not exserted. Capsule depressedglobose, 

8-10 mm in diameter, the pericarp thin, 

light brown; seeds 4 per fruit, glabrous, dark 

brown, dull, angular, 3.5-4 mm long. 




Distribution: One of the most common species 

of Ipomoea in Puerto Rico. It is found on roadsides 

and river banks, on fences, in pastures, and in 

forests at lower and middle-upper elevations. Also 

on Culebra, Vieques and the Virgin Islands; the 

Antilles, the United States (Florida), the Bahamas, 

and from Mexico to Brazil. 

Public forests: Carite, Ceiba, El Yunque, 

Guánica, Guilarte, Maricao, Piñones, Río Abajo, 

Toro Negro, Tortuguero, and Vega. 

21. Ipomoea tricolor Cav., Icon. 3: 5, t. 208. 

Fig. 71. I-K 

Herbaceous vine, twining or creeping, 

attaining 5 m in length, without milky latex. 

Stems glabrous, cylindrical, slender. Leaves 

alternate; blades simple, 5-12 × 4-14 cm, ovate, 

chartaceous, the apex acuminate and mucronate, 

the base deeply cordiform, the margins entire; 

upper and lower surface glabrous, minutely 

punctate, with the venation slightly prominent; 

petioles 2.5-10 cm long, glabrous. Flowers in 

simple or double dichasia, axillary; peduncle as 

long as or shorter than the petioles. Calyx green, 

not accrescent, sepals equal, 6-8 mm long, 

lanceolate, glabrous, obtuse at the apex, with the 

margin hyaline; corolla blue, turning violet when 

mature, infundibuliform, 6-7 cm long, the tube 

white outside and yellow inside, the limb with 5 

rounded lobes; stamens and stigmas white, not 

exserted. Capsule conical, 1.3-1.6 cm long, light 

brown; seeds 4 per fruit, glabrous, black, dull, 9- 

10 mm long. 


February. 


Distribution: On roadsides and in disturbed 

areas. Also on St. Croix, St. John, and St. Thomas. 

Native to Central America, but distributed 

throughout the tropics because of its cultivation. 

22. Ipomoea triloba L., Sp. Pl. 161. 1753. 

Fig. 72. A-C 

Bejuquillo de puerco 

Herbaceous vine, twining or creeping, 2-3 m 

in length, with scarce milky latex. Stems green, 

cylindrical, smooth, slender, puberulous. Leaves 

alternate; blades simple, 3-6 (9) × 3-5 (6.5) cm, 

ovate to subcircular, usually 3-5-lobed, 

chartaceous, the apex mucronate, acute, or 

emarginate, the base cordiform or sagittate, the 

margins smooth, slightly revolute; upper surface 

with minute hairs on the veins; lower surface 

glabrous, with prominent venation; petioles 

glabrous, somewhat sulcate, 2-4 cm long, with 

two glandular depressions where they join the 

blade. Flowers in simple or compound dichasial 

cymes, axillary; peduncles longer than the 

petioles, quadrangular, pubescent, tuberculate. 

Calyx green, not accrescent, the sepals pilose, 

subequal, ca. 8 mm long, ovate to oblanceolate, 

the apex acuminate or acute; corolla violet-pink

180 



or lavender, usually reddish at the base in the 

center, infundibuliform or campanulate, 1.4-1.6 

cm long, the limb ca. 1 cm in diameter; stamens 

sagittate, white, two of them shorter, not exserted; 

stigmas globose, white, not exserted. Capsule 

depressed-globose, brown, ca. 8 mm in diameter, 

pilose, with the style persistent; seeds 4 per fruit, 

3-5 mm long, glabrous, dark brown to black. 

Phenology: Flowering and fruiting throughout the 

year. 


Distribution: In grasslands and pastures and on 

roadsides. Also on Icacos, Caja de Muerto, 

Desecheo, Mona, Vieques, St. Croix, St. John, 

St. Thomas, Tortola, and Virgin Gorda. 

Naturalized in tropical regions of the Old 

World. 

Public forests: Boquerón, Guánica, Mona, 

Piñones, and Río Abajo. 

Commentary: In the absence of the corolla, this 

species can be confused with I. tiliacea in 

individuals with relatively large and unlobed 

leaves. Nevertheless, I. triloba is distinguished 

by having pilose sepals, while those of I. 

tiliacea are glabrous. 

23. Ipomoea violacea L., Sp. Pl. 161. 1753. 

Fig. 1. B; 72. I-K 

SYNONYMS: Convolvulus tuba Schltdl. 

Calonyction tuba (Schltdl.) Colla 

Ipomoea tuba (Schltdl.) G. Don 

Ipomoea macrantha Roem. & Schult. 

Convolvulus grandiflorus L.f. 

Operculina grandiflora (L.f.) House, pro parte 

Bejuco de vaca, Flor de luna, Coast moon vine, 

Beach morning glory 

Slightly woody vine, twining, up to 15 m in 

length, with abundant milky latex. Stems 

cylindrical or subtriangular, grayish, with 

numerous lenticels usually arranged in lines; adult 

stems up to 3 cm in diameter, cross section with 

numerous alternating concentric rings of xylem 

and phloem. Leaves alternate; blades 9-15 × 7-11 

cm, glabrous, broadly ovate, sometimes trilobed, 

fleshy-coriaceous, usually involute, the apex acute 

or acuminate, mucronate, the base cordiform or 

deeply cordiform, the margins undulate; upper 

surface light green, with the venation yellowish; 

lower surface pale green, shiny, with prominent 

venation; petioles cylindrical, glabrous, yellowish 

green, longer than the blade, with a pair of 

triangular glands at the base of the blade, swollen 

at the base. Flowers nocturnal, solitary or in 

simple dichasial cymes, axillary; peduncles 

cylindrical, up to 7 cm long, with two bracteoles 

in the upper middle portion. Calyx yellowish 

green, accrescent, the sepals unequal, 2-2.5 cm 

long, fleshy, concave, glabrous, the apex obtuse 

to rounded; corolla white, hypocrateriform, 

coriaceous, 5-7 cm long, the tube externally 

yellowish, internally with numerous transparent 

hairs and cardinal red fringes at the base, the limb 

6-7 cm in diameter, with 5 obtuse lobes; stamens 

and stigmas not exserted. Capsules globose or 

depressed-globose, yellowish brown, 2.5-3 cm in 

diameter, with accrescent sepals, persistent at the 

base; seeds 4, obtusely angular, 1-1.2 cm long, 

brown, velvety, with a row of hairs along two 

angles. 




zone. Also on Cayo Ratones, Culebra, Desecheo, 

Mona, Vieques, St. Croix, St. John, St. Thomas, 

and Tortola; the Antilles, the Bahamas, the 

Cayman Islands, the United States (Florida), and 

from Mexico to the Guianas. Introduced in the 

tropics of the Old World. 

Public forests: Guánica, Mona, and Piñones. 

24. Ipomoea wrightii A. Gray, Syn. Fl. N. Amer. 

2(1): 213. 1878. 

Fig. 72. D-H 

SYNONYM: Ipomoea pulchella sensu Griseb. 

Herbaceous vine, annual, twining, climbing, 

or creeping, 3-4 m in length. Stems cylindrical, 

slender, glabrous, sometimes with minute 

spiniform projections. Leaves alternate, palmately 

compound; leaflets 5, lanceolate or oblonglanceolate, 

1-4 cm long, glabrous, chartaceous, 

the apex acute, the base acuminate, the margins 

entire; upper surface sparsely punctate, with the 

midvein slightly prominent; lower surface pale 

green, with prominent venation, punctate or 

foveolate; petioles 2-6 cm long, slender, glabrous, 

with minute spiniform projections. Flowers 

usually solitary, axillary; peduncles 2-4 cm long, 

filiform, sinuate or spirally twisted; pedicels thick, 

claviform, 0.6-1.5 cm long, usually verrucose on

182 


the upper portion; bracts deltate, ca. 1 mm long. 

Calyx light green, not accrescent, the sepals equal, 

5-7 mm long, ovate, glabrous, verrucose, the apex 

obtuse or rounded and mucronate; corolla pale 

violet or purple, infundibuliform, 1.5-2 cm long, 

the limb 1.5-1.7 cm in diameter, with 5 obtuse 

lobes; stamens and stigmas not exserted. Capsule 

conical, 1-1.5 cm long, glabrous, the pericarp thin; 

seeds 4 per fruit, elongate, obtusely triangular, 

tomentulose, ca. 1 cm long. 

Phenology: Collected in flower in November 

and in fruit in February. 

Status: Apparently exotic, rare. 

Distribution: Known only from three 

collections in Puerto Rico, one from the Guánica 

Lagoon (Sintenis 3619), another from Sabana 

Grande (Sintenis 7019), and the last from Dorado 

(Woodbury, s.n.). A pantropical species of 

unknown origin, widely distributed in the New 

World. 

Additional Species: Ipomoea eriocarpa R.Br. 

and Ipomoea pestigris L. have been collected in 

5. JACQUEMONTIA 

Twining or creeping vines, herbs, or decumbent shrubs, apparently without milky latex. Leaves 

alternate, petiolate; blades simple, entire or lobed, usually punctate; stipules absent. Flowers bisexual, 

actinomorphic, in compound or less frequently simple dichasial cymes, axillary; peduncles usually 

elongate. Calyx of 5 free sepals, not accrescent, equal or unequal; corolla campanulate, infundibuliform, 

rotate, or hypocrateriform, the limb entire to deeply 5-lobed; stamens 5, inserted or exserted, the filaments 

usually unequal, the anthers lanceolate; ovary superior, 2-locular, the style solitary with two elongate 

stigmas, oblong to ellipsoid, slightly compressed. Fruits capsular, 4-valvate, but each valve opening in 

two; seeds 4 per fruit, triangular, glabrous or pubescent. A predominantly neotropical genus of about 

100 species. 

Key to the species of Jacquemontia 

Puerto Rico from cultivated plants; nevertheless, 

neither of these is common in our gardens, nor 

are they found naturalized. 

Doubtful Species: Ipomoea krugii Urb. is 

known only from the type collection made by 

Leopoldo Krug (no. 776) in Mayagüez between 

1868 and 1876. The type collection of this species, 

like its illustration (Krug, Flora Portoricensis 

icones, Fig. 128) were deposited in the Botanical 

Museum of Berlin, but were destroyed during the 

Second World War. Today, the only surviving 

material of this species is a photograph of the 

illustration (distributed by the Field Museum in 

Chicago). The description of this species agrees 

in numerous aspects with Ipomoea triloba and, 

since the presumed species has not been collected 

for more than a century, it is possible that it 

represents a teratological collection of I. triloba 

with white flowers or some other ornamental 

species that has not become naturalized in Puerto 

Rico. 

1a. Corollas tubular, red or crimson................................................................................ 5. J. solanifolia 

1b. Corollas infundibuliform or rotate, white, blue, pink, or lavender. ................................................. 2 

2a. Cymes compact, forming a head; bracts foliaceous, > 1.5 cm long, forming an involucre at the base 

of the inflorescence; plant hirsute ..............................................................................6. J. tamnifolia 

2b. Cymes open, not forming a head; bracts minute, not forming an involucre at the base of the 

inflorescence; plant tomentose, pubescent, or glabrous. .................................................................. 3 

3a. Corollas rotate, blue, the limb pentagonal in outline. ......................................................................4 

3b. Corollas infundibuliform, white, lavender, or pink, the limb deeply lobed. ................................... 5 

4a. Plant ferruginous-tomentose; corolla violet-blue...................................................... 2. J. cumanenis 

4b. Plant sparsely pubescent; corolla brilliant blue ........................................................4. J. pentanthos


184 


5a. Inflorescences pedunculate; corolla white, sometimes with a pink or lavender tinge. ................... 6 

5b. Inflorescences sessile; corolla lavender or pink ...................................................... 7. J. verticillata 

6a. Leaves coriaceous or subfleshy, the apex rounded or less frequently acute, emarginate and mucronate, 

the base acute, obtuse, or cuneate; lateral branches numerous, short, densely leafy, persistent on the 

main stem even after the loss of the leaves; inflorescences of simple dichasial 

cymes......................................................................................................................1. J. cayensis 

6b. Leaves coriaceous, the apex obtuse and mucronate, the base truncate or rounded, unequal; lateral 

branches elongate or short, sparsely leafy, deciduous; inflorescences of double dichasial 

cymes................................................................................................................ 3. J. havanensis 

1. Jacquemontia cayensis Britton in Britton & 

Millsp., Fl. Bahamas 349. 1920. 

Fig. 73. A-C 

Slightly woody vine, twining, much branched 

from the base, attaining 2 m in length. Stems 

slender, cylindrical, copper-brown, densely 

pubescent, with whitish stellate hairs. Leaves 

alternate or clustered on short axillary branches; 

blades simple, (1) 2-3 × 0.5-1.5 (2) cm, elliptical, 

ovate, obovate, or rounded, coriaceous or sub 

fleshy, the apex rounded or less frequently acute, 

emarginate and mucronate, the base acute, obtuse, 

or cuneate, the margins revolute, reddish, with 

stellate hairs; upper surface yellowish green, 

glabrous or with some scattered stellate hairs, the 

midvein sunken; lower surface yellowish green, 

dull, with a prominent midvein, covered with 

stellate hairs; petioles 5-12 mm long, reddish, 

densely covered with stellate hairs. Flowers 

arranged in simple dichasial cymes; peduncles ca. 

4 mm long, stellate-pubescent; pedicels 6-8 mm 

long, stellate-pubescent. Calyx green, crateriform, 

the sepals ovate, 2.5-3 mm long, acute at the apex, 

the margins brown; corolla white, 

infundibuliform, ca. 1 cm long, deeply lobed, the 

lobes ovate, ca. 5 mm long; stamens white, 

exserted; stigmas white, bilobate. Capsule ovoid, 

ca. 4 mm in diameter, light brown, glabrous; seeds 

pyriform-triangular, 2-3 mm long, puberulous or 

glabrous. 

Phenology: Collected in flower and fruit in 

mid-September. 


Distribution: Along the south coast of Puerto 

Rico. Also on Mona and Anegada; the Bahamas, 

Cuba, and the Turks Islands. 


Commentary: Jacquemontia cayensis can be 

confused with J. havanensis; both species are 

quite variable, to the point that it is difficult to 

identify some specimens as one species or the 

other. As a general rule, J. cayensis has leaves 

rounded at the apex and flowers clustered in 

reduced cymes. Jacquemontia havanensis, on the 

other hand, has leaves with acute or obtuse apices 

and the cymes contain 4 or more flowers. 

2. Jacquemontia cumanensis (Kunth) Kuntze, 

Revis. Gen. Pl. 2: 441. 1891. 

Fig. 73. D-F 

BASIONYM: Convolvulus cumanenis Kunth 

Slightly woody vine, twining, much branched 

from the base, up to 2 m in length, without latex. 

Stems slender, cylindrical, ferruginous-tomentose. 

Leaves alternate; blades simple, 1.2-4 × 0.8-2.6 

cm, broadly ovate, chartaceous, the apex obtuse 

or acute, the base cordiform, the margins sinuate; 

upper and lower surface tomentose; petioles 0.8- 

2 cm long, tomentose. Flowers in compound 

dichasia; peduncles 1.5-3 cm long; bracts 

elliptical, ca. 5 mm long. Calyx green, 

crateriform, the sepals 8-10 mm long, tomentose, 

broadly ovate or deltate, acuminate at the apex; 

corolla violet-blue to almost violet, with white 

lines forming a star inside, rotate, the limb 2.2- 

2.5 cm in diameter, pentagonal in outline and with 

the margins reflexed; stamens and pistil white, 

exserted. Capsule globose, ca. 5 mm in diameter, 

brown, glabrous, with the sepals persistent at the 

base; seeds obtusely triangular, ca. 4 mm long, 

brown, glabrous. 


September to March.




littoral zone. Also on Vieques, St. Croix, St. John, 

St. Thomas, and Tortola; the Lesser Antilles and 

Venezuela. 


3. Jacquemontia havanensis (Jacq.) Urb., Symb. 

Antill. 3: 342. 1902. 

Fig. 73. G-I 

BASIONYM: Convolvulus havanensis Jacq. 

SYNONYM: Jacquemontia jamaicensis (Jacq.) H. Hallier ex 

Soler. 

Aguinaldo de costa 


in length, without milky latex. Stem branched 

from the base, slender, cylindrical, densely 

covered with white, stellate hairs. Leaves 

alternate; blades simple, 1.5-5 × 0.5-1.5 cm, 

lanceolate, oblong, ovate, or linear, coriaceous, 

sparsely stellate-pubescent, the apex obtuse and 

mucronate, the base truncate or rounded, usually 

unequal, the margins revolute; upper surface with 

the venation sunken; lower surface with 

prominent venation; petioles slender, 6-12 mm 

long, slightly compressed, stellate-pubescent. 

Flowers in compound dichasial cymes, axillary, 

shorter than the leaves. Calyx green, crateriform, 

the sepals unequal, 4-5 mm long, puberulous, 

ovate to oblanceolate, the apex apiculate; corolla 

white, sometimes with a pink tinge at the margins, 

infundibuliform, 1-1.3 cm long, the limb with 

deep, obtuse lobes; stamens and stigmas white, 

slightly exserted. Capsules ovoid or ellipsoid, 5- 

6 mm long, with the sepals persistent at the base; 

seeds triangular, 2-2.3 mm long, brown, glabrous, 

with a short marginal wing, membranaceous. 


throughout the year, particularly from November 

to March. 


Distribution: In dry thickets along the 

southern littoral zone. Also on Mona, Cayo 

Ratones, Cayo Icacos, Culebra, Vieques, St. Croix, 


the Antilles, the United States (Florida), the 

Bahamas, southern Mexico, and Belize. 


4. Jacquemontia pentanthos (Jacq.) G. Don, Gen. 

Hist. 4: 283. 1838. 

Fig. 74. A-D 

BASIONYM: Convolvulus pentanthos Jacq. 

Aguinaldo azul, Clashie-melanie 


in length, without latex. Stems cylindrical, 

slender, sparsely covered with white trifid hairs, 

glabrous when mature. Leaves alternate; blades 

simple, 1.8-6 × 1.2-3.7 cm, ovate or lanceolate, 

chartaceous, sparsely covered with white trifid 

hairs on both surfaces, the apex obtuse, acute, or 

mucronate and sometimes retuse, the base 

cordiform or truncate, the margins entire or 

sinuate; upper surface dark green; lower surface 

pale green, with prominent venation; petioles 

slender, pubescent or glabrous, 1-4 cm long. 

Flowers in congested dichasial cymes; peduncles 

longer than the leaves; bracts elliptical, ca. 5 mm 

long. Calyx green, the sepals broadly ovate or 

deltoid, 8-10 mm long, pubescent, acuminate at 

the apex; corolla blue or brilliant blue, with the 

center white, forming a star of 5 arms, rotate, the 

limb pentagonal in outline, 2.2-2.5 cm in 

diameter, the margins reflexed; stamens and 

stigmas white, exserted. Capsules globose, ca. 5 

mm in diameter, brown, with the sepals persistent 

at the base; seeds obtusely triangular, ca. 4 mm 

long, brown, glabrous. 




Distribution: In thickets and dry forests and 

on sandy coasts of the littoral zone. Also on Cayo 

Santiago, Culebra, Desecheo, Mona, Vieques, St. 


Gorda; throughout the Neotropics, introduced in 

Malaysia and Sri Lanka. 


Guánica, Maricao, Mona, Río Abajo, and Susúa. 

5. Jacquemontia solanifolia (L.) H. Hallier, Bot. 

Jahrb. Syst. 16: 542. 1893. 

Fig. 74. E-H 

BASIONYM: Ipomoea solanifolia L. 

SYNONYMS: Ipomoea filiformis Jacq. 

Exogonium solanifolium (L.) Britton

186 



Slightly woody vine, twining, branched from 

the base, attaining 2 m in length and not 

producing latex. Stems cylindrical, slender, 

densely covered with white trifid hairs when 

young. Leaves alternate; blades simple, 3-6 × 1.5- 

3.7 cm, lanceolate, chartaceous, sparsely covered 

with white trifid hairs on both surfaces, the apex 

obtuse, acute, or mucronate, the base cordiform, 

rounded, obtuse, or truncate, the margins revolute; 

upper surface dark green; lower surface pale 

green, with prominent venation; petioles slender, 

pubescent, 1.5-4 cm long, canaliculate. Flowers 

sparse, in elongate cymes; peduncles 2-4 cm long; 

bracts subulate, ca. 5 mm long. Calyx green, 

crateriform, the sepals unequal, 4-5 mm long, 

ovate or rounded, glabrous; corolla 

hypocrateriform, crimson, 2-2.5 cm long, with 5 

deep lobes, expanded, obtuse, mucronate at the 

apex; stamens and stigmas white, exserted. 

Capsules ovoid or conical, 7-8 mm in diameter, 

brown, with the sepals persistent at the base; seeds 

obtusely triangular, ca. 4 mm long, brown, 

glabrous, with a short marginal wing at the 

vertices. 




Distribution: In dry forests and thickets of the 

littoral zone. Also on Mona, Vieques, St. Croix, 

St. John, and St. Thomas; the Lesser Antilles. 

Public forests: Guajataca, Guánica, Mona, 

Piñones, and Río Abajo. 

6. Jacquemontia tamnifolia (L.) Griseb., Fl. Brit. 

W. I. 5: 474. 1862. 

Fig. 74. I-K 

BASIONYM: Ipomoea tamnifolia L. 

SYNONYM: Thyella tamnifolia (L.) Raf. 

Aguinaldo peludo 

Slightly woody vine, twining, climbing, or 

creeping, attaining 5 m in length, with watery 

latex. Stems cylindrical, slender, hirsute or 

glabrescent, with malpighiaceous or simple hairs. 

Leaves alternate; blades simple, 3-10 × 1.5-3.7 

cm, ovate, chartaceous, the apex acute or 

acuminate, the base truncate to cordiform, the 

margins entire or sinuate; upper surface green, 

punctate; lower surface pale green, with 

prominent venation; petioles slender, pubescent 

or glabrous, 1-4 cm long. Flowers in compact 

cymes, forming a head; peduncles longer than the 

petioles; bracts foliaceous, ovate, ca. 2.5 cm long, 

forming an involucre. Calyx green, the sepals 

lanceolate, 10-15 mm long, hirsute, acuminate at 

the apex; corolla white, turning lavender or pale 

blue, infundibuliform, as long as the sepals; 


globose, 4-5 mm long, light brown, glabrous, with 

the sepals persistent at the base; seeds obtusely 

triangular, ca. 2 mm long, light brown, glabrous. 





littoral zone. Also in the southeastern United 

States, Cuba, Hispaniola, the Bahamas, the Lesser 

Antilles, Central America, South America, Africa, 

and the Mascarenes. 

Public forests: Boquerón and Guajataca. 

7. Jacquemontia verticillata (L.) Urb., Symb. 

Antill. 3: 339. 1902. 

Fig. 74. L-N 

BASIONYM: Ipomoea verticillata L. 


with watery latex. Stems cylindrical, slender, 

appressed-pubescent, with white trifid hairs. 

Leaves alternate; blades simple, 2.2-4.2 × 0.7- 

1.5 cm, lanceolate, chartaceous, the apex obtuse, 

acute, or mucronate, the base cordiform, the 

margins entire or repand, appressed-pubescent on 

both surfaces; upper surface dark green; lower 

surface pale green, with the venation 

inconspicuous; petioles slender, pubescent or 

glabrous, 0.7-1 cm long. Flowers in compact 

dichasial cymes, sessile or subsessile; bracts 

minute. Calyx green, the sepals subequal, 2-3 mm 

long, ovate-lanceolate, pubescent or glabrous, 

acuminate at the apex; corolla pink, lavender, or 

rarely white, infundibuliform, ca. 6 mm long, the 

limb deeply 5-lobed; stamens not exserted; 

stigmas exserted. Capsules ellipsoid, ca. 4 mm 

long, brown, with the sepals persistent at the base; 

seeds obtusely triangular, ca. 2 mm long, brown, 

glabrous. 


November and December. 

Status: Native, rare.

188 



Distribution: Known from the area of mogotes. 

Also Cuba, Hispaniola, Jamaica, and the 

Bahamas. 

6. MERREMIA 

Twining vines, with milky or watery latex, sometimes with glandular hairs. Leaves alternate, 

petiolate; blades simple, palmately lobed or compound; stipules absent. Flowers bisexual, actinomorphic, 

5-merous, solitary or in compound dichasial cymes, axillary; peduncles usually elongate. Calyx usually 

accrescent, the sepals equal or unequal; corolla campanulate or infundibuliform, the limb entire or 

slightly 5-lobed; stamens inserted, the filaments subequal, the anthers lanceolate, twisted after opening; 

ovary superior, 2-locular, the style solitary, with two subglobose stigmas. Fruits capsular, ovoid to 

globose, 4-valvate or irregularly dehiscent; seeds 4 per fruit, triangular, glabrous or velvety. A pantropical 

genus of about 80 species. 

Key to the species of Merremia 

1a. Leaves with simple blades; flowers in umbelliform cymes ......................................6. M. umbellata 

1b. Leaves palmately compound or palmatilobed; flowers solitary or in few-flowered dichasial cymes......2 

2a. Leaves palmatilobed .......................................................................................................................... 3 

2b. Leaves palmately compound ............................................................................................................. 4 

3a. Plants glabrous; margin of the leaf entire or slightly undulate; corolla brilliant yellow 

............................................................................................................................ 5. M. tuberosa 

3b. Plants hispid; margin of the leaf sinuate or sinuate-dentate; corolla white with reddish or pink 

center ............................................................................................................................. 3. M. dissecta 

4a. Leaflets with the margin entire. ........................................................................................................ 5 

4b. Leaflets with the margin serrate ........................................................................... 4. M. quinquefolia 

5a. Sepals hispid, acute at the apex................................................................................... 1. M. aegyptia 

5b. Sepals glandular-pubescent, acuminate at the apex .................................................. 2. M. cissoides 

1. Merremia aegyptia (L.) Urb., Symb. Antill. 

4: 505. 1910. 

Fig. 75. A-E 

BASIONYM: Ipomoea aegyptia L. 

Slightly woody vine, twining or creeping, 

attaining 3 m in length, with scarce milky latex. 

Stems cylindrical, slender, hispid, with simple 

yellowish hairs. Leaves alternate; blades 5palmately 

compound, chartaceous; leaflets 4-14 

× 2-6 cm, oblanceolate or elliptical, the apex and 

base acuminate, the margins entire, ciliate; upper 

and lower surface hispidulous; petioles slender, 

sulcate, hispid, 6-8 cm long. Flowers in dichasial 

cymes; peduncles shorter than the petioles; bracts 

deciduous. Calyx green, the sepals unequal, 1.5- 


2 cm long, lanceolate or elliptical, hispid outside 

on the basal portion, acute at the apex; corolla 

white, infundibuliform, 2.5-3 cm long, the limb 

slightly pentagonal, 4-4.5 cm in diameter; 


subglobose, ca. 1 cm in diameter, light brown, 

glabrous, with the sepals persistent, accrescent, 

and expanded; seeds obtusely triangular, 5-6 mm 

long, pale brown, glabrous. 


November to May. 


Distribution: Occasional in disturbed areas at 

lower and middle elevations. Also on Mona, St. 


Gorda; throughout the tropics.

190 


Public forests: Boquerón, Cambalache, and Mona. 

2. Merremia cissoides (Lam.) H. Hallier, Bot. 

Jahrb. Syst. 16: 552. 1893. 

Fig. 75. F-G 

BASIONYM: Convolvulus cissoides Lam. 

SYNONYM: Ipomoea cissoides (Lam.) Griseb. 


with scarce watery latex. Stems cylindrical, 

slender, glandular-pubescent and frequently 

hispidulous, glabrescent. Leaves alternate; blades 

5-palmately compound, 2-7.2 × 1.2-3.3 cm, 

chartaceous; leaflets elliptical, ovate, or ovatelanceolate, 

the apex obtuse, the base acute, the 

margins entire or slightly dentate; upper surface 

glabrous; lower surface glandular-pubescent or 

glabrous; petiolules ca. 1 mm long, glandularpubescent; 

petioles slender, hispidulous, 

glandular-pubescent, 2-3 (6) cm long. Flowers in 

simple or double dichasial cymes; peduncles 

longer than the petioles; bracts persistent, 

subulate. Calyx green, the sepals unequal, 1-1.5 

cm long, ovate or ovate-lanceolate, glandularpubescent 

externally, acuminate at the apex; 

corolla white, with the center purple, 

infundibuliform, 2-3 cm long, the limb 5-lobed, 

(2.2)3-4 cm in diameter; stamens and stigmas 

white, not exserted. Capsules globose, 6-8 mm in 

diameter, light brown, glabrous, with the sepals 

persistent; seeds ellipsoid, ca. 6 mm long, grayish, 

lanate. 

Phenology: Collected in flower in February. 


Public forest: Guánica (according to Quevedo 

et al., 1990). 

Distribution: Occasional in disturbed areas at 

lower elevations in eastern Puerto Rico. Also in 

Cuba, continental tropical America, and tropical 

Asia. 

3. Merremia dissecta (Jacq.) H. Hallier, Bot. 

Jahrb. Syst. 16: 552. 1893. 

Fig. 75. H-K 

BASIONYM: Convolvulus dissectus Jacq. 

Slightly woody vine, twining, much branched, 

2-5 m in length, with scarce milky or watery latex. 

Stems cylindrical, slender, striate, hirsute when 

young, glabrous when mature. Leaves alternate; 

blades simple, 4-7 × 1.2-3 cm, 7-9-palmatilobed, 

divided almost to the base, chartaceous, glabrous, 

the lobes sinuate to sinuate-dentate, lanceolate or 

elliptical in outline, acuminate and mucronate 

toward the apex, the lower lobes sometimes again 

lobate; petioles slender, pilose or glabrous, 2.5-4 

cm long. Flowers solitary or occasionally in 

cymes; peduncles as long as the petioles; bracts 

deciduous. Calyx green, the sepals unequal, 2- 

2.2 cm long, lanceolate to rounded, glabrous, 

hyaline; corolla white, with the center reddish, 

infundibuliform, 3-4 cm long, the tube light 

yellow, the limb pentagonal, 3.5-4 cm in diameter; 

stamens and stigmas yellow, not exserted. 

Capsules depressed-globose, 1-1.5 cm wide, 

opening by 4 hyaline valves, brown, glabrous, with 

accrescent sepals, ca. 3 cm long, persistent; seeds 

obtusely and asymmetrically pyramidal, 5-7 mm 

long, dull black, glabrous. 


fruiting from August to December. 



elevations. Also on Vieques, St. Croix, St. John, 

St. Thomas, and Tortola; throughout the tropics. 

Public forests: Guajataca, Guánica, and Susúa. 

4. Merremia quinquefolia (L.) H. Hallier, Bot. 

Jahrb. Syst. 16: 552. 1893. 

Fig. 76. A-D 

SYNONYM: Ipomoea quinquefolia L. 

Batatilla blanca 


twining, 2-3 m in length, with scarce watery latex. 

Stems slender, cylindrical, sparsely hirsute or 

glabrous, sometimes with a reddish or mulberrycolored 

tonality. Leaves alternate, (3-)5-palmately 

compound, chartaceous; leaflets 2-7 × 0.5-1.2 cm, 

elliptical or lanceolate, glabrous, the apex 

acuminate, mucronate, the base acute and sessile, 

the margins serrate; upper surface green, dull, 

glabrous, with the venation sunken; lower surface 

pale green, dull, glabrous, with prominent 

venation; petioles slender, 1-3 cm long, hispid or 

glabrous. Flowers solitary or in simple dichasia; 

peduncles as long as the petioles, glandularpubescent; 

pedicels slender, 8-15 mm long; bracts 

minute. Calyx green, the sepals unequal, ca. 1 

cm long, glabrous, oblong, the apex obtuse; 

corolla pale yellow or white, infundibuliform, 2-


192 


2.2 cm long, the limb with obtuse lobes, slightly 

reflexed; stamens and stigmas pale yellow, not 

exserted. Capsules depressed-globose, opening by 

4 valves, 0.7-1 cm long, light brown, glabrous; 

seeds obtusely trigonal, 3-4.5 mm long, black, 

covered with whitish lanate hairs. 

Phenology: Flowering throughout the year, 

especially from November to May. 


Distribution: In disturbed areas, on roadsides 

and in pastures, at lower elevations. Also on 

Culebra, Mona, Vieques, St. Croix, St. John, St. 

Thomas, Tortola, and Virgin Gorda; throughout 



Guánica, Mona, Río Abajo, Susúa, and Toro 

Negro. 

5. Merremia tuberosa (L.) Rendle in Dyer, Fl. 

Trop. Afr. 4: 104. 1905. 

Fig. 76. E-G 

BASIONYM: Ipomoea tuberosa L. 

Flor de palo 

Woody vine, climbing, twining, 10-15 m in 

length, with abundant milky latex. Stems thick, 

cylindrical, glabrous. Leaves alternate; blades 

simple, 7-12 × 6-11 cm, 7-palmatilobed, the lobes 

elliptical, long-acuminate at the apex, the base 

cordiform, the margins revolute, slightly sinuate; 

upper surface dark green, slightly shiny, glabrous, 


green, dull, glabrous or puberulous, with the 

venation yellowish, prominent; petioles as long 

as the blade, cylindrical, glabrous or puberulous. 

Flowers functionally unisexual, solitary or in 

simple dichasia. Calyx yellowish green, the sepals 

unequal, 2-3 cm long, fleshy, accrescent and 

woody once the fruit is formed; corolla yellow, 

infundibuliform, 4-5 cm long, the limb 4-5 cm in 

diameter; stamens exserted, the anthers white; 

stigma bilobed, green, exserted. Capsules ovoid, 

opening irregularly, 1.5-2.5 cm long, light brown, 

with the sepals persistent and accrescent at the 

base; seeds 4 per fruit, black, obtusely trigonal, 

1-1.5 cm long, velvety. 


December and fruiting from November to March. 


uncommon. 

Distribution: Ornamental plant, sometimes 

naturalized in disturbed areas. Also on St. Croix 

and St. John. Native to tropical America but found 

widely distributed throughout the tropics. 

6. Merremia umbellata (L.) H. Hallier, Bot. 

Jahrb. Syst. 16: 552. 1893. 

Fig. 76. H-L 

BASIONYM: Convolvulus umbellatus L. 

SYNONYM: Ipomoea polyanthes Willd. ex Roem. & Schult. 

Aguinaldo amarillo, Yellow morning glory 

Slightly woody vine, climbing or creeping, 

twining, attaining 5 m in length, with scarce milky 

latex. Stems slender, cylindrical, glabrous or 

pubescent, green or copper-colored, with a pair 

of spiniform projections at the nodes. Leaves 

alternate; blades simple, 4-17 × 5-12 cm, ovate 

or lanceolate, chartaceous, glabrous, the apex 

obtuse, acute, or short-acuminate and mucronate, 

the base cordiform or sagittate, the margins 

undulate; upper surface yellowish green, dull, 

glabrous, with the venation sunken, covered with 

minute white hairs; lower surface glabrous, with 

the veins puberulous; petioles cylindrical, usually 

longer than the blade, pubescent. Flowers in 

umbelliform cymes, axillary; peduncles shorter 

than the petioles, angular or cylindrical, 

puberulous, sometimes with winged projections; 

pedicels ca. 2 cm long, puberulous. Calyx green, 

the sepals unequal, 1-1.5 cm long, glabrous, 

overlapping, ovate or rounded; corolla brilliant 

yellow, infundibuliform, 2.5-3 cm long, the limb 

ca. 3 cm in diameter, with obtuse lobes; stamens 

white, not exserted; stigmas greenish, slightly 

exserted. Capsule globose, 4-valvate, ca. 1 cm 

long, brown, covered by the persistent sepals; 

seeds obtusely trigonal, 5-8 mm long, brown, 

velvety, with a line of longer hairs on two of the 

margins. 


November to April and sometimes later in the year. 


Distribution: In disturbed areas, on roadsides 

and in pastures and vacant lots, at lower to middle 


St. Thomas, and Tortola; throughout tropical 

America. 

Public forests: Guánica and Susúa.


194 


7. OPERCULINA 

Twining vines, with milky latex. Stems, petioles, peduncles, and pedicels usually winged. Leaves 

alternate, petiolate; blades simple, entire or lobed; stipules absent. Flowers bisexual, actinomorphic, 

solitary or in simple dichasial cymes, axillary; peduncles longer than the petioles. Calyx campanulate, 

of 5 equal or unequal sepals, usually not green, accrescent and woody in fruits, irregularly dentate at 

the margins; corolla campanulate or broadly infundibuliform, the limb entire or pentagonal; stamens 

5, inserted, the anthers twisted after opening; ovary superior, 2-locular, the style solitary, with two 

subglobose stigmas. Fruits capsular, ovoid to globose, circumscissile, with the exocarp separating like 

an operculum, the remaining layers of the fruit wall opening irregularly, or 4-valvate; seeds 4 per fruit, 

glabrous or velvety. A pantropical genus of about 12 species, with the majority of the species native to 

Central America and northern South America. 

1. Operculina turpethum (L.) J. Silva Manso, 

Enum. Subst. Braz. 16. 1836. 


1a. Leaves (mature) 5-16 cm long; corollas 3-4.5 

cm long……... O. turpethum var. turpethum 

1b. Leaves (mature) 16-24 cm long; corollas 7-8 

cm long………. O. turpethum var. ventricosa 

1a. Operculina turpethum var. turpethum 

Fig. 77. A-C 

BASIONYM: Convolvulus turpethum L. 

SYNONYMS: Convolvulus triqueter Vahl 

Operculina triquetra (Vahl) H. Hallier 

Slightly woody vine, twining. Stems slender, 

angular, sulcate or 3-5-winged, pubescent when 

young, glabrous when mature. Leaves alternate; 

blades simple, 5-16 × 5-15 cm, broadly ovate, 

orbicular, ovate-lanceolate, or lanceolate, 

chartaceous, the apex acuminate, acute, obtuse, 

or rounded and mucronate, the base cordiform or 

hastate, the margins entire or slightly undulate; 

upper surface glabrous or appressed-pubescent, 

dull, with the venation slightly prominent; lower 

surface densely appressed-pubescent, with 

prominent venation; petioles cylindrical, 2.5-5 cm 

long, appressed-pubescent. Flowers solitary or in 

simple dichasial cymes; peduncles 2-18 cm long, 

pubescent, cylindrical; bracts oblong, 2-3 cm long, 

at the base of the pedicels. Calyx light green, the 

sepals unequal, 1.5-2.5 cm long, the outer ones 

appressed-pubescent, ovate or broadly ovate, 

obtuse and mucronate at the apex; corolla white 

or with the center yellowish, broadly 

infundibuliform, glabrous, 3-4.5 cm long, the limb 

ca. 3 cm in diameter, with obtuse lobes; stamens 

and stigmas not exserted. Capsule depressedglobose, 

ca. 1.5 cm long, operculate, with the 

inner layers opening irregularly, covered by the 

accrescent and persistent sepals; seeds black, 

glabrous, subglobose, keeled, with a prominent 

hilum, ca. 6 mm long. 


Status: Exotic, cultivated, uncommon, 

probably not naturalized. 

Distribution: St. Croix, St. John, and St. 

Thomas. Native to the Old World. 

1b. Operculina turpethum var. ventricosa 

(Bertero) Staples & D. F. Austin, Brittonia 33: 

595. 1981. 

BASIONYM: Convolvulus ventricosus Bertero 

SYNONYM: Operculina ventricosa (Bertero) Peter 

Woody vine, robust, twining, attaining 10 m 

or more in length, with milky latex. Stems 

cylindrical, glabrous, with the pith hollow in dried 

specimens. Leaves alternate; blades simple, 16- 

24 × 5-15 cm, broadly ovate, chartaceous, the apex 

acuminate or obtuse and mucronate, the base 

cordiform, the margins entire or slightly undulate; 

upper surface glabrous, dull, with the venation 

slightly prominent; lower surface glabrous or 

puberulous, the prominent venation appressedpubescent; 

petioles cylindrical, 8-20 cm long, 

appressed-pubescent. Flowers solitary or in simple 

dichasial cymes; peduncles 15-20 cm long, thick, 

appressed-pubescent, cylindrical; bracts ovate, ca. 

3.5 cm long, deciduous. Calyx yellowish green,


the sepals subequal, 3.5-4 cm long, externally 

appressed-pubescent or glabrous, oblong-ovate, 

obtuse and mucronate at the apex; corolla white, 

broadly infundibuliform, glabrous, 7-8 cm long, 

the limb 8-10 cm in diameter, with obtuse lobes; 

stamens and stigmas not exserted. Capsule 

globose, ca. 3 cm long, operculate, with the inner 

layers opening irregularly, covered by the 

accrescent and persistent sepals; seeds black, dull, 

8. PORANOPSIS 

A monospecific genus, characterized by the following species. 

1. Poranopsis paniculata (Roxb.) Roberty, 

Candollea 14. 26. 1953. 

Fig. 77. D-G 

BASIONYM: Porana paniculata Roxb. 

Velo de novia, Christmas vine 


with scarce watery latex. Stems slender, 

cylindrical, canescent. Leaves alternate; blades 

simple, 4-9 × 3-6.5 cm, chartaceous, ovate, 

palmatinerved, the apex acute, acuminate, or 

cuspidate, the base cordiform, the margins entire 

or slightly sinuate; upper surface pubescent, with 


densely pubescent, with prominent venation; 

petioles cylindrical, pubescent, 2-3.2 cm long. 

Flowers bisexual, actinomorphic, in axillary or 

terminal panicles, ascending, up to 17 cm long, 

with the axes tomentose; bracts oblong or 

lanceolate, minute, tomentose. Calyx green, 

canescent-tomentose, accrescent, the sepals free 

9. STICTOCARDIA 

glabrous, obtusely trigonal, with a prominent 

hilum. 


March and fruiting in January and March 


Distribution: Species native to the Old World, 

introduced to the Antilles as an ornamental, where 

it can be found naturalized. Also on St. Croix and 

St. Thomas. 

to the base, ca. 1.5 mm long, lanceolate; corolla 

white, infundibuliform, glabrous, 4.5-6 mm long, 

the limb 5-lobed; stamens 5, not exserted, the 

filaments very short, of equal size; ovary superior, 

with an annular disc at the base, unicarpellate, 

with two basal ovules, the style very short, the 

stigma globose or bilobate, green, not exserted. 

Fruit indehiscent, ovoid-globose, 5-6 mm long, 

with the three outer sepals accrescent; seed usually 

one, ovoid or subglobose. 



Status: Exotic, cultivated and naturalized in 

disturbed areas, uncommon. 

Distribution: Species native to India, 

cultivated throughout the tropics, where it can be 

found naturalized in disturbed areas. Also on 

Vieques, St. Croix, and St. Thomas. 

Twining vines, glabrous or pubescent, usually with scarce milky latex. Leaves alternate, petiolate; 

blades simple, entire, punctate on the lower surface; stipules absent. Flowers bisexual, actinomorphic, 

solitary or in simple dichasial cymes, axillary. Calyx of 5 subequal sepals; corolla infundibuliform; 

stamens 5, inserted, the anthers lanceolate; ovary superior, 2-locular, the style solitary, with two 

subglobose stigmas, not exserted. Fruits indehiscent, 4-locular, with fibrous walls, tardily dehiscent by 

the dissolution of the pericarp, covered by the accrescent sepals; seeds 4 per fruit, obtusely triangular, 

velvety. A Paleotropical genus of about 12 species.

196 



1. Stictocardia tiliifolia (Desr.) H. Hallier, Bot. 

Jahrb. Syst. 18: 159. 1893. 

Fig. 78. A-D 

BASIONYM: Convolvulus tiliaefolius Desr. 

SYNONYM: Rivea campanulata sensu House, non (L.) House 

Woody vine, creeping or climbing, twining, 

attaining 5 m in length, with scarce milky latex. 

Stems slender, cylindrical, glabrous or pubescent, 

with numerous short lateral branches. Leaves 

alternate; blades simple, 5-17 × 5-12 cm, ovate 

to subrounded, chartaceous, glabrous, the apex 

acute, obtuse, or short-acuminate and mucronate, 

the base cordiform or sagittate, the margins 

undulate; upper surface dull with the venation flat; 

lower surface dull, punctate, with prominent 

venation; petioles 10-15 cm long, subcylindrical, 

slightly sulcate, glabrous. Flowers solitary or in 

simple cymes, axillary. Calyx green, the sepals 

glabrous, unequal, 1.7-2 cm long, overlapping, 

ovate or rounded; corolla pink or lavender, 6-8 

cm long, the limb up to 6 cm in diameter, with 

rounded lobes; stamens and stigmas pink, not 

exserted. Fruit globose, 2.5-3 cm long, brown, 

tardily dehiscent by the dissolution of the pericarp, 

covered by the accrescent sepals; seeds obtusely 

triangular or rounded, ca. 1 cm long, brown, 

velvety. 



Status: Exotic, naturalized in disturbed areas, 

uncommon. 

Distribution: In areas of low elevation, on the 

north and west coasts. Also on Cayo Santiago, 

Vieques, St. Croix. St. John, St. Thomas, and 

Tortola. Native to tropical Asia, but dispersed 

throughout the tropics because of its cultivation.

198 


10. TURBINA 

Woody vines, twining, glabrous or pubescent, which produce scarce watery latex. Leaves alternate, 

petiolate; blades simple, cordiform; stipules absent. Inflorescences of axillary cymes, terminal panicles, 

or the flowers solitary; sepals unequal, ovate or lanceolate, accrescent in the fruit; corollas infundibuliform 

or hypocrateriform, white, greenish, pink, or crimson; stamens inserted, the filaments glandularpubescent 

at the base; ovary superior, bilocular, the style simple, with two globose stigmas. Fruits 

indehiscent, dry, woody to crustaceous, ellipsoid or globose, unilocular; seed usually 1, ovoid to ellipsoid, 

pubescent. A pantropical genus of about 12 species. 

1. Turbina corymbosa (L.) Raf., Fl. Tellur. 11: 

81. 1838. 

Fig. 2. D; 79. A-G 

BASIONYM: Convolvulus corymbosus L. 

SYNONYM: Rivea corymbosa (L.) H. Hallier 

Aguinaldo blanco, Corona de novia 


base, attaining 10 m or more in length, with scarce 

watery latex. Stems glabrous or pubescent, 

cylindrical when young, compressed, angular, or 

sulcate when mature, up to 3 cm in diameter, the 

bark grayish, with numerous dark lenticels; cross 

section of the stem polystelic, with peripheral 

vascular cylinders of different diameters. Leaves 

alternate; blades 3-10 × 2.7-5.5 cm, ovate, 

chartaceous, the apex acute, short-acuminate, or 

acuminate, the base cordiform, the margins entire, 

slightly undulate; upper surface dark green, 

slightly shiny, glabrous, the venation flat; lower 

surface pale green, dull, glabrous or puberulous, 

with a prominent midvein; petioles 1.5-6 cm long, 

11. XENOSTEGIA 

slender, cylindrical, glabrous. Flowers numerous 

in axillary corymbs. Calyx green, the sepals 

oblong, 10-12 mm long, coriaceous, acute at the 

apex; corolla 2.5-3 cm long, white, 

infundibuliform, the tube yellow internally, except 

for the dark violet base, the limb with 5 yellowish 

bands forming a stellate design; stamens white, 

not exserted, the anthers sagittate; stigmas white, 

globose, not exserted. Fruits indehiscent, ellipsoid 

to subovoid, crustaceous, 1-1.5 cm long, the sepals 

persistent, unequal, accrescent, woody. Seed one, 

ellipsoid to subglobose, 4-5 mm long, minutely 

pubescent, light brown. 

Phenology: Flowering from the end of October 

to January and in May, and fruiting in January 

and February. 


Distribution: On roadsides or in recent 

secondary forests. Also throughout the Antilles, 

from Mexico to Bolivia, and the United States 

(Florida), introduced in the Old World tropics. 

Public forests: Cambalache, Río Abajo, and 

Susúa. 

A genus of two species, native to tropical Asia. The species that is described below characterizes the 

genus. 

1. Xenostegia tridentata (L.) D. F. Austin & 

Staples, Brittonia 32: 533. 1980. 

Fig. 79. H-M 

BASIONYM: Convolvulus tridentatus L. 

SYNONYM: Ipomoea angustifolia Jacq. 

Merremia angustifolia (Jacq.) H. Hallier 

Ipomoea filicaulis Willd. 

Slightly woody vine, twining, much branched, 

climbing or creeping, 1-2 m in length, with scarce 

milky latex. Stems glabrous or pubescent, angular. 

Leaves alternate; blades 2-10 × 0.5-0.6 cm, 

chartaceous, linear, oblong, or lanceolate, the apex 

acute or acuminate and mucronate, the base 

hastate or auriculate, the margins 1-3-dentate in 

the basal portion; upper surface dark green, dull,


200 


glabrous, the venation flat; lower surface pale 

green, dull, glabrous or puberulous, with a 

prominent midvein; petioles 1-1.5 mm long, 

slender, cylindrical, glabrous. Flowers solitary or 

in simple cymes, axillary; peduncles slender, 2.5- 

4 cm long. Calyx green, the sepals unequal, 

oblong or ovate-oblong, 4-10 mm long, the two 

outer ones smaller, obtuse-truncate at the apex; 

corolla white or pale yellow, campanulate, 1-2 cm 

long, the limb with 5 deep lobes; stamens not 

exserted; stigmas not exserted. Capsule 

subglobose, 4-9 mm long, 4-valvate, the sepals 

persistent; seeds 1-4, obtusely ovoid, 2-3 mm long, 

glabrous, light brown, dull, squamulose. 




Distribution: Native to the Old World tropics 

(Africa-Pacific), naturalized in some localities of 

the northern littoral zone (Manatí-Santurce), on 

sandy substrates. Also in South America and the 

Old World tropics. 

Public forest: Tortuguero. 

References: Austin, D. F. 1982. Convolvulaceae. Flora de Venezuela. Vol. 8. Ediciones Fundación, 

Venezuela. Austin, D. F. and S. Demissew. 1997. Unique fruits and generic status of Stictocardia 

(Convolvulaceae). Kew Bull. 52: 161-169. Austin, D. F. Genera of Convolvulaceae. WWW. Fau.edu/ 

divdept/biology/protologues.htm. Staples, G. W. and D. F. Austin. 1981. Changes in the West Indian 

Operculina (Convolvulaceae). Brittonia 33: 591-596. 

19. Family CUCURBITACEAE 


1a. Fruits < 7 cm long.............................................................................................................................. 2 

1b. Fruits > 10 cm long ........................................................................................................................... 8 

2a. Fruits capsular, dehiscent, with numerous pendulous seeds, covered by a fleshy red aril 

............................................................................................................................ 11. Momordica 

2b. Fruits indehiscent, the seeds without an aril .................................................................................... 3 

3a. Fruits red; corolla white, the limb 3-5 cm in diameter ................................................... 3. Coccinia 

3b. Fruits green, yellowish, or orange; corolla of various colors, when white then the limb 1-1.5 cm in 

diameter .............................................................................................................................................. 4 

4a. Fruits spinulose, spiny, or smooth, 4-5 cm long .......................................................4. Cucumis 

4b. Fruits smooth, 1-4 cm long ............................................................................................................... 5 

5a. Infructescences short, with the fruits densely clustered; stems articulate; roots tuberous 

................................................................................................................................. .6. Doyerea 

5b. Infructescences elongate, racemose, or the fruits solitary; stems not articulate; roots not tuberous......6 

6a. Plants herbaceous, usually 1-2 (4) m long; stems slender, < 5 mm in diameter; corolla yellow 

............................................................................................................................. 10. Melothria 

6b. Plants robust, usually 5 m or more in length; stems > 5 mm in diameter; corolla orange, creamcolored, 

white, or greenish yellow ..................................................................................................... 7 

7a. Corolla orange; stems cylindrical................................................................................... 12. Psiguria 

7b. Corolla cream-colored, white, or greenish yellow; stems angular...............................1. Cayaponia


8a. Fruits dry when mature...................................................................................................................... 9 

8b. Fruits fleshy, juicy when ripe .......................................................................................................... 11 

9a. Fruits with woody walls, not fibrous inside .................................................................................... 10 

9b. Fruits with thin and fragile walls on drying, the interior formed by a conglomeration of fibers that 

form a network ........................................................................................................................ 9. Luffa 

10a. Plants hispid-pubescent; petioles with prominent glands; fruits indehiscent, elongate, 20-35 cm 

long; seeds ovate-oblong, compressed, ca. 1.5 cm long ................................................8. Lagenaria 

10b. Plants glabrous; petioles without glands; fruits dehiscent by an operculum that occupies 1/3 of the 

fruit, this with 3 sutures that are united at the apex, subglobose, 8-12 cm in diameter; seeds circular, 

slightly compressed, 3-6 cm in diameter........................................................................... 7. Fevillea 

11a. Fruits white or light green, pyriform, spinulose ...........................................................13. Sechium 

11b. Fruits dark green, orange, reddish brown, or with light green bands alternating with dark green, 

not pyriform, smooth........................................................................................................................ 12 

12a. Fruits cylindrical (3 to 4 times longer than wide), reddish brown, aromatic ................ 14. Sicana 

12b. Fruits almost as wide as long, green, orange, or with dark green bands .................................... 13 

13a. Corolla campanulate, the limb ca. 10 cm in diameter ................................................ 5. Cucurbita 

13b. Corolla rotate, the limb ca. 2 cm in diameter ................................................................ 2. Citrullus 

1. CAYAPONIA 

Herbaceous vines, monoecious or dioecious, glabrous or pubescent, with tendrils axillary or lateral 

to the leaves; latex watery, usually abundant. Leaves alternate, petiolate; blades simple, entire, lobed, 

palmatilobed or palmatifid; stipules absent. Flowers unisexual, actinomorphic, solitary or in racemes, 

panicles or fascicles, axillary. Calyx campanulate, with 5 minute lobes; corollas tubular, the lobes 

reflexed or expanded; staminate flowers with a pistillode and 3 exserted stamens, the filaments adnate 

to the corolla, the anthers sigmoid, concrescent; pistillate flowers with 3 staminodia; ovary inferior, 3carpellate, 

trilobed, the style simple, with 3 stigmatic branches. Fruit a small berry; seeds 2-3, compressed. 

A predominantly neotropical genus of about 60 species. 

Key to the species of Cayaponia 

1a. Calyx 5-9 mm long; lobes of the corolla 12-15 mm long; margin of the leaves entire or crenate........... 

.................................................................................................................... 1. C. americana 

1b. Calyx 3-4 mm long; lobes of the corolla 3-5 mm long; margin of the leaves spinulose .................. 

...................................................................................................................................... 2. C. racemosa 

1. Cayaponia americana (Lam.) Cogn. in A.DC. 

& C.DC., Monogr. Phan. 3: 785. 1881. 

Fig. 80. A-B 

BASIONYM: Bryonia americana Lam. 

Bejuco de torero 

Herbaceous vine, slender, climbing or 

creeping, with axillary tendrils, attaining 10 m 

in length. Stems angular, glabrous or puberulous, 

swollen at the nodes; tendrils simple or branched, 

usually longer than the leaves. Leaves alternate; 

blades 5-20 × 5-18 (25) cm, chartaceous, ovate, 

palmatilobed, with 3-5 more or less deep lobes, 

oblong to linear, the apices acute, acuminate, or 

rounded, the base cordiform or lyrate, the margins

202 


entire or crenate; upper surface dark green, shiny, 

hispidulous, with the venation sunken and 

pubescent; lower surface yellowish green, dull, 

with the venation reticulate, prominent, and 

pubescent; petioles sulcate, puberulous, 2-7 cm 

long. Flowers solitary or in short racemes. Calyx 

green, campanulate, 5-9 mm long; corolla white 

or greenish yellow, minutely pubescent, the tube 

ca. 1 cm long, the lobes 12-15 mm long, oblong, 

reflexed; stamens exserted, the filaments greenish, 

pubescent at the base, the anthers yellow. Berry 

ellipsoid, 1.5-2 cm long, smooth, olive-green, 

turning orange or red-orange when ripe; seeds 

ovate, 0.8-1.2 cm long, not arillate. 


February to December. 

Status: Native, very common throughout 


Distribution: On roadsides and in dry forests 

and coastal thickets. Also on Culebra, Vieques, 


Virgin Gorda; the United States (Florida), Cuba, 

Hispaniola, and the Lesser Antilles. 

Public forests: Carite, Ceiba, El Yunque, 


2. Cayaponia racemosa (Mill.) Cogn. in A.DC. 

& C.DC., Mon. Phan. 3: 768. 1881. 

Fig. 80. C-G 

BASIONYM: Bryonia racemosa Mill. 

2. CITRULLUS 

Climbing herbaceous vine, with axillary 

tendrils, attaining 10-15 m in length. Stems green, 

cylindrical, striate, glabrous, swollen at the nodes; 

tendrils trifid, 15 cm long or longer. Leaves 

alternate; blades entire or 3-7-lobed, 4-20 × 3-17 

(25) cm, ovate, chartaceous or subcoriaceous, the 

lobes oblong, the apex acuminate or acute, the 

base cordiform or reniform, the margins 

spinulose; upper surface scabrid; lower surface 

light green, dull, with the reticulate venation 

prominent, puberulous; petioles sulcate or slightly 

winged, glabrous, 4-7 cm long. Flowers unisexual, 

in axillary racemes. Calyx pale green, crateriform 

or campanulate, 3-4 mm long, the lobes 

triangular; corolla greenish white, tubular, the 

tube ca. 3 mm long, pubescent inside, the lobes 

3-5 mm long, reflexed. Berry ovoid, 10-12 mm 

long, green, turning orange when ripe; seeds 2- 

3, elliptical, ca. 8 mm long, not arillate. 





Distribution: In pastures and on roadsides at 

lower elevations. Cited for Tortola by Britton 

(1925); also in the Greater Antilles, Central 

America, Barbados, Trinidad, and northern South 

America. 

Public forests: Guilarte, Maricao, Río Abajo, 

Susúa, and Toro Negro. 

Herbaceous vines, creeping or climbing, monoecious, pubescent, with axillary tendrils with 2-5 

branches or simple; latex watery. Leaves alternate, petiolate; blades simple, entire, lobed-pinnatifid; 

stipules absent. Flowers unisexual, actinomorphic, solitary or rarely in axillary fascicles; calyx 

campanulate, with 5 narrow lobes; corollas rotate or broadly campanulate, deeply 5-lobed; staminate 

flowers with a pistillode and 3 exserted stamens, the filaments free, adnate to the base of the corolla, 

the anthers concrescent or free; pistillate flowers with 3 staminodia; the ovary inferior, ovoid, 

tricarpellate, the styles 3, the stigmas bilobate. Fruit a fleshy berry, indehiscent, large; seeds numerous, 

oblong, compressed. An Old World genus of about 4 species, some of which are cultivated for their 

edible fruits. 

1. Citrullus lanatus (Thunb.) Matsum. & Nakai, 

Cat. Sem. Spor. Hort. Bot. Univ. Imp. Tokyo 

30, no. 854. 1916. 

Fig. 81. E-J 

BASIONYM: Momordica lanata Thunb. 

SYNONYM: Cucurbita citrullus L. 

Melón, Sandía 

Creeping or climbing herbaceous vine, with 

axillary tendrils, attaining 1.5-3 m in length. 

Stems green, sulcate, pilose or lanate, glabrous 

when mature; tendrils with 2-5 branches, 15 cm 

long or longer. Leaves alternate; blades 6-16 × 3- 

11 cm, ovate-triangular in outline, deeply 3-5-


204 


pinnatifid-lobed, membranaceous, the lobes 

oblong or ovate, lobate, obtuse or rounded at the 

apex, the base cordiform or reniform, the margins 

undulate or irregularly dentate; upper surface 

lanate-pubescent; lower surface light green, dull, 

with the reticulate venation prominent, 

puberulous, scabrous in mature leaves; petioles 

lanate-pubescent, 1-11 cm long. Staminate and 

pistillate flowers of similar size, solitary; 

peduncles 2-4 cm long, pubescent. Hypanthium 

villous. Calyx green, campanulate, ca. 1 cm long, 

villous, lobes linear-lanceolate, 3-4 mm long; 

corolla yellow or greenish yellow, campanulate, 

3. COCCINIA 

1-1.5 cm long, villous outside, limb ca. 2.3 cm in 

diameter, the lobes deep. Berry globose or 

cylindrical, up to 40 cm long, green, mottled or 

with light green lines, the endocarp thick, pink, 

white, or yellowish; seeds numerous, elliptical, 

5-10 mm long. 


and in March. 

Status: Exotic, cultivated or spontaneous, 

uncommon. 

Distribution: Species cultivated for its edible 

fruits. Spontaneous in sandy areas and on 

roadsides. Native to tropical Africa, but widely 


Herbaceous vines, dioecious, with axillary tendrils, simple or bifid; latex watery. Stems striate, 

glabrous. Leaves alternate, petiolate; blades simple, entire or lobed; stipules absent. Flowers unisexual, 

actinomorphic, solitary or in short racemes; calyx campanulate, with 5 minute lobes; corolla campanulate, 

5-lobed, with the venation dark; staminate flowers with 3 stamens, the filaments free from one another 

or distally connate, the anthers connate, triplicate; pistillate flowers with 3 staminodia; ovary inferior, 

smooth, trilocular, with numerous horizontal ovules, the style with 3 stigmatic branches. Fruit a globose 

berry, fleshy; seeds numerous, ovate. An African genus of about 30 species. 

1. Coccinia grandis (L.) Voigt, Hort. Suburb. 

Calcutt. 59. 1845. 

Fig. 81. A-D 

BASIONYM: Bryonia grandis L. 

SYNONYM: Coccinia cordifolia sensu Britton, non (L.) Cogn. 

Subwoody vine, climbing by axillary tendrils, 

which attains 20 m in length. Stems much 

branched, angular, with white dots, pubescent in 

the area of the nodes; tendrils simple or bifid, 

shorter than the leaves. Leaves alternate; blades 

simple, 4-11 × 4.2-10 cm, entire or deeply 5-lobed, 

the apex obtuse or acute, the base cordiform, the 

central lobe lanceolate, the lateral ones 

asymmetrical, ovate-lanceolate, the margins 

minutely dentate; upper surface dull, scabrid; 

lower surface pale green, dull, slightly scabrid, 

4. CUCUMIS 

with prominent venation; petioles 1.5-2 cm long, 

sulcate, glabrous. Flowers usually solitary, 

axillary, the staminate and pistillate flowers 

similar. Hypanthium ellipsoid. Calyx turbinate, 

green, ca. 5 mm long, glabrous; corolla white, 

with the center pale yellow, campanulate, 5-lobed, 

the tube 2-2.5 cm long, the limb 3-5 cm in 

diameter. Berry ellipsoid, smooth, 3-6 cm long, 

red when ripe; seeds numerous, compressed, ca. 

6 mm long. 

Phenology: Collected in flower in August. 

Status: Exotic, naturalized, locally common. 

Distribution: In southwestern Puerto Rico and 

in Quebradillas. Also on St. Croix, and cited for 

St. Thomas (Britton and P. Wilson, 1925). Native 

to Africa, but naturalized throughout the tropics. 

Herbaceous vines, monoecious, with axillary tendrils; latex watery. Leaves alternate, petiolate; 

blades entire, dentate or palmatilobed; stipules absent. Flowers unisexual, actinomorphic, solitary or 

in axillary fascicles; calyx campanulate, with 5 minute lobes; corolla yellow, campanulate; staminate 

flowers with a pistillode and 3 stamens, the filaments adnate to the base of the corolla, the anthers


206 


concrescent or free, distally with the connective prolonged into an appendage; pistillate flowers with 3 

staminodia; ovary inferior, trilocular, with numerous horizontal ovules, the style with 3-5 stigmatic 

branches, bilobed. Fruit an ellipsoid or cylindrical berry, fleshy; seeds numerous, elliptical. An Old 

World genus of about 30 species, some of which are widely cultivated for their edible fruits. 

Key to the species of Cucumis 

1a. Leaves deeply 3-5-palmatilobed .................................................................................... 1. C. anguria 

1b. Leaves ovate, entire or slightly 3-lobed. ........................................................................................... 2 

2a. Fruits densely spiny (spines ca. 1 cm long) ...............................................................2. C. dipsaceus 

2b. Fruits smooth, not spiny ..................................................................................................... 3. C. melo 

1. Cucumis anguria L., Sp. Pl. 1011. 1753. 

Fig. 82. A-D 

Pepinillo silvestre, Cocombro, Pepineto, 

Wild cucumber 

Herbaceous vine, creeping or climbing by 

axillary tendrils, which attains 0.25-2 m in length. 

Stems branched from the base, slender, angular, 

hirsute; tendrils simple, shorter than the leaves. 

Leaves alternate; blades 3-10 × 3.5-10 cm, deeply 

3-5-palmatilobed, the lobes oblong or 

oblanceolate, the apices obtuse or rounded, the 

base lyrate, the margins ciliate, crenate or 

denticulate; upper surface dull, scabrid; lower 

surface pale green, dull, scabrid, with prominent 

venation and hispidulous; petioles sulcate, 

hispidulous, 6-12 cm long. Flowers solitary or in 

axillary fascicles, unisexual. Calyx campanulate, 

yellowish, 5-6 mm long, villous-spinulose; corolla 

pale yellow, ca. 1 cm long, the lobes acute. Berry 

ellipsoid or obovoid, spinulose, 4-5 cm long, 

greenish yellow, edible; seeds numerous, elliptical, 

cream-colored, 1-1.3 cm long. 


January to March and in July and November. 


Distribution: Along the coast or in coastal 

thickets, in southern and southwestern Puerto 

Rico. Also on Culebra, Mona, Vieques, Anegada, 

St. Croix, St. Thomas, and Tortola. Native to 

Africa, but naturalized in the Antilles, Central 

America, and South America. 

Public forest: Guánica and Mona. 

2. Cucumis dipsaceus Ehrenb. ex Spach, Hist. 

Nat. Veg. 6: 211. 1838. 

Fig. 82. E-F 


axillary tendrils, which attains 1.5 m in length. 

Stems much branched, slender, angular, sulcate, 

hispid on the angular margins; tendrils simple, 

longer than the leaves. Leaves alternate; blades 

2.5-7.5 × 2-7 cm, ovate or trilobed, the lobes 

obtuse, the base cordiform, the apex obtuse, the 

margins dentate or entire; upper surface dull, 

scabrid; lower surface pale green, dull, scabrid, 

with prominent venation and hispidulous; petioles 

sulcate, hispidulous or hispid, 1.5-5 (12) cm long. 

Flowers solitary, the staminate flowers sometimes 

in axillary fascicles; calyx campanulate, 3.6-5.2 

mm long, hispidulous, the lobes narrowly oblong 

or linear, 1.6-4 mm long, hispidulous; corolla pale 

yellow, campanulate, hispidulous outside, the tube 

1-1.5 mm long, the lobes obovate, acute, 5.5-8.5 

mm long. Berry ellipsoid or globose, densely spiny 

(spines ca. 1 cm long), 3-6.5 cm long, pale yellow; 

seeds numerous, elliptical, cream-colored, 4-5 mm 

long. 

Phenology: Collected in flower in April and 

in fruit from April to June. 


Distribution: In coastal pastures in southern 

Puerto Rico. Also on Tortola. Native to Africa, 

but naturalized throughout the tropics. 

3. Cucumis melo L., Sp. Pl. 1011. 1753. 

Fig. 82. G-I 

Creeping herb or herbaceous vine, creeping 

or climbing by axillary tendrils, attaining 2 m in 

length. Stems branched from the base and along 

the main stems, slender, angular, sulcate, hispid; 

tendrils simple, hispid, as long as the leaves. 

Leaves alternate; blades 2.2-13 × 2-13 cm, ovate,


broadly ovate, or trilobed, the lobes obtuse, the 

base cordiform or lyrate, the margins denticulate; 

upper surface dull, scabrid; lower surface pale 

green, dull, scabrid, with prominent venation and 

hispidulous; petioles 1-9 cm long, sulcate, hispid. 

Staminate flowers in axillary fascicles; pistillate 

flowers solitary; calyx campanulate, yellowish 

green, 2.8-4(5.6) mm long, hispidulous, the lobes 

linear to triangular, 1.2-3.6 mm long; corolla 

yellow, infundibuliform, 7-1.4 mm long, the lobes 

obtuse. Berry ellipsoid, smooth, 4.5-5.5 cm long, 

green with whitish spots, turning yellow-orange 

5. CUCURBITA 

when ripe, the pulp extremely bitter; seeds 

numerous, elliptical, cream-colored, 4-5 mm long. 





elevations, collected in Gurabo and Lajas. An 

African species that has given rise to several races 

with edible fruits through artificial selection. 

Among these are the “cantaloupe” and the 

“honeydew.” 

Herbaceous vines, annual, monoecious, creeping or climbing by axillary tendrils; stems sulcate, 

producing watery latex; tendrils axillary, with 2-5 branches. Leaves alternate, petiolate; blades subentire, 

rounded or deeply lobed; stipules absent. Flowers unisexual, actinomorphic, solitary, the staminate 

flowers also in axillary fascicles; calyx campanulate, rarely cylindrical, with (4-)5(-7) minute lobes; 

corolla campanulate, deeply lobed; stamens 3, the filaments fleshy, inserted at the base of the floral 

receptacle, the anthers linear, connate into a cylindrical column; ovary inferior, 3-5-locular, with 

numerous horizontal ovules, the style simple, short, with 3-5 stigmatic branches, bilobed or bifurcate. 

Fruit a fleshy, fibrous, or woody berry, large; seeds numerous, elliptical. A New World genus of about 

30 species, some of which are widely cultivated for their edible fruits. 

1. Cucurbita moschata Duchesne ex Poir., Dict. 

Sci. Nat. 11: 234. 1818. 

Fig. 83. A-B 


axillary tendrils, 5-10 m in length. Stems 

branched from the base and along the main stems, 

flexible, angular, pubescent with soft or slightly 

rigid hairs, elongate, not pungent; tendrils with 

4 branches, pilose, shorter than the petiole. Leaves 

alternate; blades 15-25 × 15-25 cm, broadly ovate, 

slightly lobed, the lobes obtuse, the apex obtuse, 

the base cordiform or hastate, the margins finely 

serrate; upper surface puberulous, usually with 

irregular whitish spots; lower surface shortpubescent, 

with prominent venation; petioles 11- 

39 cm long, striate, puberulous to densely 

pubescent, with unicelular hairs intermingled with 

multicellular trichomes. Flowers solitary; calyx 

campanulate, yellowish green, 3.5-4 cm long, 

hirsute, the lobes linear to oblong, 2.5-3 cm long; 

corolla brilliant yellow, campanulate, 7-9 cm long, 

the lobes obtuse, revolute; peduncle thick, angular, 

sulcate. Berry variable, soft or hard, depressedglobose 

to globose, green, turning yellowish when 

ripe, 25-30 cm long; mesocarp orange, fleshy, 

thick; seeds numerous, elliptical, 1.5-2 cm long, 

cream-colored to light brown. 




common. 

Distribution: On roadsides, along trails, and 

in pastures. 

Public forests: Maricao, Piñones, and Río 

Abajo. 

Commentary: In Puerto Rico two other species 

of Cucurbita (C. pepo L. and C. maxima Duchesne 

ex Lam.) have been reported; nevertheless, these 

are rarely cultivated and are not found naturalized 

in Puerto Rico (pers. comm. Dr. Linda Beaver, 

University of Puerto Rico, Mayagüez Campus).

208 



6. DOYEREA 


1. Doyerea emetocathartica Grosourdy, Med. 

Bot. Criollo 2: 338. 1864. 

Fig. 83. C-F 

SYNONYMS: Corallocarpus emetocatharticus (Grosourdy) 

Cogn. 

Anguria glomerata Eggers 

Herbaceous vine, dioecious, climbing by 

tendrils, attaining 10 m or more in length. Stems 

green, cylindrical, fleshy, fragile, glabrous, 

producing abundant watery latex, much branched 

from the base, which bears tuberous roots; tendrils 

axillary, simple, shorter than the leaves. Leaves 

alternate; blades 5-10 × 5.5-10 cm, ovate or 

broadly ovate, usually 3-lobed, chartaceous, the 

apex acute, obtuse, or acuminate, usually 

mucronate, the base cordiform or lyrate, the 

margins minutely dentate; upper surface scabrid; 

lower surface pubescent, with the reticulate 

venation prominent; petioles sulcate, pubescent, 

ca. 4 cm long. Flowers unisexual, in axillary 

7. FEVILLEA 

cymes. Calyx green or pale orange, campanulate, 

3.5-4 mm long, puberulous, the lobes oblong, 

reflexed; corolla white, yellowish green, or pale 

orange, campanulate, the tube ca. 1 mm long, the 

lobes 1-1.2 mm long, pilose, oblong, erect or 

reflexed; stamens 2, the filaments short, 

concrescent; ovary inferior, elongate, bicarpellate, 

the style simple, bifurcate near the apex, the 

stigmas bifid, subglobose and exserted. Berries 

asymmetrically ellipsoid, 1-1.2 cm long, smooth, 

pale green, turning orange when ripe; seeds few, 

ovoid, ca. 4 mm long, not arillate. 


August, and September and in fruit in August. 



Also on Culebra, Vieques, St. Croix, St. John, and 

St. Thomas; the Antilles and from Mexico to 

northern South America. 

Public forests: Boquerón and Guánica. 

Herbaceous or slightly woody vines, dioecious, with simple axillary tendrils; latex watery. Leaves 

alternate, petiolate; blades entire, cordiform; stipules absent. Flowers unisexual, actinomorphic, in 

axillary panicles; calyx campanulate, with 5 minute lobes; corolla rotate, the lobes with an appendage 

on the adaxial surface; staminate flowers with 5 stamens, the filaments recurved; pistillate flowers 

with 5 minute staminodia, the ovary inferior, globose, tricarpellate, the styles 3, connate at the base, 

the stigmas forming a trilobed capitulum. Fruits globose, dehiscent by an operculum, which occupies 

1/3 of the fruit, pericarp subwoody; seeds numerous, large, circular, compressed, not arillate. A 

neotropical genus of about 10 species. 

1. Fevillea cordifolia L., Sp. Pl. 1013. 1753. 

Fig. 84. A-F 

Pepita amarga, Uyama, Secua 

Slightly woody vine that climbs by axillary 

tendrils and attains 10 m or more in length. Stems 

green, subcylindrical, glabrous, striate when 

young, with numerous pendulous lateral branches; 

tendrils simple or bifurcate, up to 16 cm long. 

Leaves alternate; blades entire, 8-16 × 4-12 cm, 

ovate or pentagonal, coriaceous, glabrous, the 

apex acuminate, the base cordiform, the margins 

revolute, entire or denticulate; upper surface dark 

green, shiny; lower surface light green, dull, the 

venation palmate, prominent; petioles usually 

curved, sulcate, glabrous, 4-6 cm long. Staminate 

and pistillate flowers similar, in axillary panicles, 

20-60 cm long; pedicels ca. 8 mm long. Calyx 

crateriform, puberulous, the sepals ovate, ca. 2 

mm long; corolla yellow-pink, rotate, the limb 

ca. 1 cm in diameter, with rounded lobes; stamens 

5, ca. 1.5 mm long; ovary globose, puberulous.

210 



Fruit subglobose, green, 8-12 cm in diameter, 

operculate, with three sutures united in the distal 

portion, the pericarp subwoody; seeds few, 

compressed, subcircular, 3-6 cm in diameter, 

cream-colored. 


September and fruiting from June to September. 

8. LAGENARIA 


1. Lagenaria siceraria (Molina) Standl., Field 

Mus. Bot. 3: 435. 1930. 

Fig. 84. G-K 

BASIONYM: Cucurbita siceraria Molina 

SYNONYM: Lagenaria vulgaris Ser. 

Güiro, Güicharo, Marimbo 

Herbaceous vine, monoecious, which climbs 

by tendrils, 5-10 m in length. Stems herbaceous, 

puberulous or densely pubescent, striate; tendrils 

axillary, bifid. Leaves alternate; blades simple, 6- 

15 × 7-22 cm, ovate or pentagonal, angular or 

lobed, chartaceous, the apex apiculate, the base 

cordiform, the margins sinuate-dentate; both 

surfaces puberulous; petioles elongate, with a pair 

of lateral glands, corniculate where they are joined 

to the blade; stipules absent. Flowers white, 

nocturnal, unisexual, actinomorphic, usually 

solitary, axillary; staminate flowers larger than 

the pistillate flowers; peduncles longer than the 

accompanying petiole in the staminate flowers or 

as long as the petiole in the pistillate flowers. 

9. LUFFA 

The seeds are dispersed by water currents and are 

frequently found on river banks and beaches. 


Distribution: In disturbed areas along rivers, 

roads, and moist forest margins in central Puerto 

Rico. Also in Jamaica, Cuba, Hispaniola, 

Trinidad, and continental tropical America. 


Calyx infundibuliform to campanulate, with 5 

triangular to linear lobes, 3-4 mm long; corolla 

yellow, of free petals, expanded, 2-4 cm long, 

obovate to oblong-ovate, with the apex apiculate 

and emarginate; stamens 3, the filaments free, 

inserted on the base of the floral receptacle, the 

anthers concrescent; hypanthium < 1 cm long; 

ovary inferior, tricarpellate, ovoid to cylindrical, 

tomentose, with numerous horizontal ovules, the 

style short, the stigmas trilobate. Fruit a berry, 

very variable in form, 20-35 cm long, the pericarp 

woody, smooth, glabrous; seeds numerous, ovateoblong, 

ca. 1.5 cm long, not arillate. 


September. 


uncommon. 

Distribution: In disturbed areas along roads 

and moist forest margins in central Puerto Rico. 

Native to the Old and New World tropics, where 

it is rather frequently cultivated. 


blades simple, 5-7-lobed; stipules absent. Flowers unisexual, actinomorphic; calyx campanulate, with 

5 elongate lobes; corolla rotate, the lobes deep; staminate flowers in axillary racemes; stamens 3-5, not 

exserted, the filaments free; pistillate flowers solitary, with 3 staminodia; ovary inferior, elongate, 

tricarpellate, the styles 3, connate at the base, the stigmas forming a trilobed capitulum. Fruits elongate, 

dehiscent by apical pores, the mesocarp with a fibrous reticulum; seeds numerous, compressed, winged 

or not, not arillate. An Old World genus of about 10 species, some of which are cultivated throughout 

the tropics for their useful fruits. 

Key to the species of Luffa 

1a. Stamens 3; fruits with 10 longitudinal ribs; seeds rugose....................................... 1. L. acutangula 

1b. Stamens 5; fruits trigonal, slightly sulcate longitudinally; seeds smooth .............. 2. L. aegyptiaca

212 



1. Luffa acutangula (L.) Roxb., Hort. Beng. 70. 

1814. 

Fig. 85. D-E 

BASIONYM: Cucumis acutangula L. 

Esponja, Estropajo 

Herbaceous vine, monoecious, creeping or 

climbing by axillary tendrils, attaining 5-10 m in 

length. Stems green, angular, scabrous; tendrils 

trifid. Leaves alternate; blades 15-20 cm long, 5- 

7-palmatilobed, chartaceous, the lobes more or 

less deep, the apex acute or acuminate, the base 

cordiform or hastate, the margins sinuate-dentate 

or denticulate; upper surface scabrous; lower 

surface pale green, scabrous; petioles 8-10 cm 

long. Flowers unisexual, actinomorphic. Calyx 

urceolate, with keeled lobes, 10-12 mm long, 

triangular; corolla pale yellow, the lobes deep, 

obtuse. Staminate flowers in racemes; stamens 3, 

the filaments free, 3-4 m long, villous. Pistillate 

flowers solitary, with a hypanthium < 1 cm long; 

staminodia 3, minute, glandular; ovary inferior, 

tricarpellate, claviform, 10-angled, with numerous 

horizontal ovules, the style short, the stigmas 

globose. Fruit claviform, with 10 longitudinal ribs, 

15-30 cm long, the pericarp crustose, dehiscent 

by apical pores; seeds numerous, ovate, 11-12 mm 

long, blackish, rugose. 

Phenology: Collected in fruit in July. 

Status: Exotic, cultivated and naturalized 

(according to Britton, 1925), uncommon. 

Distribution: Probably native to the 

paleotropics, in disturbed areas along roads and 

moist forest margins in central Puerto Rico. 

Cultivated throughout the tropics and subtropics. 

2. Luffa aegyptiaca Miller, Gard. Dict. ed. 8. 

1768. 

Fig. 85. A-C 

10. MELOTHRIA 

SYNONYM: Luffa cilindrica M. Roem. 

Esponja, Estropajo, Sponge cucumber, 

Strainer vine 

Herbaceous vine, monoecious, climbing by 

axillary tendrils, attaining 10 m in length. Stems 

green, slender, subcylindrical or angular, ribbed, 

glabrous or puberulous; tendrils trifid. Leaves 

alternate; blades 11-25 (35) × 7-25 (32) cm, 3-7palmatilobed, 

chartaceous, the lobes lanceolate or 

ovate, the apices acute or acuminate, the base 

cordiform, the margins entire or serrate; upper 

surface scabrid; lower surface pale green, scabrid; 

petioles as long as or longer than the blade. 

Flowers unisexual, actinomorphic. Calyx green, 

campanulate, the lobes 12-15 mm long, 

lanceolate; corolla pale yellow, 2.5-4.5 cm long, 

the lobes ovate, obtuse. Staminate flowers in 

racemes 12-15 cm long; stamens 5, not exserted, 

the filaments free, 3-4 m long, villous. Pistillate 

flowers solitary, with a hypanthium ca. 3 cm long; 

ovary inferior, tricarpellate, ellipsoid, with 

numerous horizontal ovules, the style short, the 

stigmas globose. Fruit trigonal, slightly sulcate, 

20-45 cm long, the pericarp crustose, dehiscent 

by apical pores, the mesocarp forming a network 

of fibers; seeds numerous, elliptical, 10-13 mm 

long, black, smooth. 


January and fruiting in January and from July to 

August. 


uncommon. 

Distribution: Native to the paleotropics, 

cultivated, escaped, or naturalized, in disturbed 

areas along roads and moist forest margins in 

central Puerto Rico. Also on St. Croix, St. John, 

and St. Thomas. Cultivated throughout the tropics 

and subtropics. 


blades simple, entire or lobed; stipules absent. Flowers unisexual, actinomorphic; calyx crateriform, 

with 5 minute lobes; corolla campanulate, short-tubular, the lobes more or less deep, expanded; staminate 

flowers in axillary racemes; stamens 3, the filaments free, the anthers free or concrescent; pistillate 

flowers solitary, with 3 staminodia; ovary inferior, ovoid or fusiform, tricarpellate, the style short, 

simple, the stigmas 3, linear. Fruit a smooth berry, small, ovoid or ellipsoid; seeds numerous, minute, 

without an aril. A neotropical genus of about 10 species.

214 



1. Melothria pendula L., Sp. Pl. 35. 1753. 

Fig. 86. A-F 

SYNONYMS: Bryonia guadalupensis Spreng. 

Melothria guadalupensis (Spreng.) Cogn. 

Pepinillo cimarrón 

Herbaceous vine, much branched, which 

climbs by tendrils, attaining 1-2 (4) m in length. 

Stems slender, 2-3 mm in diameter, green, slightly 

striate, puberulous and with some minute hairs; 

tendrils axillary, simple, filiform, 6-9 cm long. 

Leaves alternate; blades ovate or 3-5-lobed, 

membranaceous, the apex acute or acuminate, 

mucronate, the base lyrate or cordiform, the 

margins crenate, repand, or denticulate; upper 

surface green, shiny, scabrid, with the venation 

flat; lower surface light green, dull, scabrid, with 

the venation slightly prominent; petioles shorter 

than the blade, sulcate. Staminate and pistillate 

flowers in the same raceme; peduncles 2-3 cm 

11. MOMORDICA 

long. Calyx yellowish green, campanulate, ca. 5 

mm long; corolla pale yellow, campanulate, ca. 5 

mm long, with 5 deep lobes; stamens subsessile, 

adnate to the corolla tube, the anthers free, the 

pistil with an annular disc at the base. Berry ovoid, 

smooth, 1.5-1.8 cm long, olive green when ripe; 

seeds numerous, elliptical or ovate, ca. 5 mm long, 

not arillate. 




Distribution: In disturbed areas such as 

pastures, at upper to lower elevations. Also on 

Vieques, St. Croix, St. John, St. Thomas, Tortola, 

and Virgin Gorda; throughout the Antilles, the 

Bahamas, the southern United States, and 

continental tropical America. 


Toro Negro, and Vega. 

Herbaceous vines, monoecious, with axillary tendrils, without latex. Leaves alternate, petiolate; 

blades simple, entire or palmatilobed; stipules absent. Flowers unisexual, actinomorphic; calyx 

campanulate or infundibuliform, with 5 minute lobes; corolla campanulate or rotate. Staminate flowers 

in axillary racemes or solitary; stamens 3, the filaments free, the anthers free or concrescent; pistillodes 

absent or glandular. Pistillate flowers solitary, on long peduncles with foliaceous bracts; staminodia 

absent; ovary inferior, ellipsoid, tricarpellate, with numerous horizontal ovules, the style terminal, 

simple, the stigmas 3, linear, bilobate. Fruit a fleshy capsule, which opens by 3 valves; seeds numerous, 

compressed, arillate. A predominantly African genus of about 40 species. 

1. Momordica charantia L., Sp. Pl. 1009. 1753. 

Fig. 86. G-K 

Cundeamor, Jumbee pumpkin, 

Maiden apple, Old maid 


axillary tendrils, attaining 8 m in length. Stems 

green, slender, subcylindrical, striate, villous; 

tendrils simple, 9-11 cm long. Leaves alternate; 

blades 3-11 × 3-10 cm, 5-7-lobed, membranaceous 

to chartaceous, the apex obtuse or acute, the base 

cordiform, the margins deeply undulate or dentate; 

upper surface dark green, dull, puberulous; lower 

surface yellowish green, dull, with prominent 

venation, puberulous; petioles slender, sulcate or 

slightly winged, villous, 3-6 cm long. Flowers 

solitary, axillary; peduncle elongate, with a foliose 

bract below the middle. Calyx yellowish green, 

campanulate, 10-12 mm long, the lobes 

lanceolate; corolla pale yellow, rotate, ca. 3.5 cm 

in diameter. Capsule ellipsoid-angular to fusiform, 

muricate, 3-5 cm long, changing from green to 

yellow-orange when mature, dehiscent at the apex 

by 3 valves; seeds pendulous, compressed, covered 

by a red and fleshy aril. 



Status: Exotic, naturalized, very common. 

Distribution: On fences and roadsides and in 

coffee plantations and pastures at middle and 

lower elevations. Also on Culebra, Mona, Vieques, 


Virgin Gorda; native to China and Asia but found 

naturalized throughout the tropics and subtropics 

of the New World.

216 



Public forests: Cambalache, El Yunque, 

Maricao, Mona, Río Abajo, Susúa, and Vega. 

Commentary: Momordica charantia has at least 

three forms that are cultivated for their edible 

fruits. These are known as “bitter melon” and 

12. PSIGURIA 

“bitter gourd” and may attain up to 60 cm long. 

In Puerto Rico the wild form, not a cultivated 

one, is found, although there exists one 

collection (Sintenis 919), made in 1885, with 

fruits ca. 15 cm long, apparently from a 

cultivated plant. 

Herbaceous vines, monoecious or dioecious, climbing by simple axillary or lateral tendrils, without 

latex. Leaves alternate, petiolate; blades simple, entire, palmatilobed or 3-5-foliolate; stipules absent. 

Flowers unisexual, actinomorphic; calyx campanulate or infundibuliform, with 5 minute lobes; corolla 

campanulate or rotate. Staminate flowers in axillary cymes with elongate peduncles; stamens 2, sessile, 

inserted on the tube of the receptacle, the anthers free, linear, the connective prolonged into an appendage; 

pistillodes absent. Pistillate flowers solitary or in groups of 2-4; staminodia 2, minute; ovary inferior, 

oblong, bicarpellate, with numerous horizontal ovules, the style terminal, simple, the stigmas 2, bifurcate. 

Fruit a cylindrical or ovoid berry, quadrangular or sulcate; seeds numerous, compressed, oblong. A 

neotropical genus of about 12 species, distributed from Mexico to Paraguay, including the Antilles. 

Key to the species of Psiguria 

1a. Leaves 3-lobed. .................................................................................................................................. 2 

1b. Leaves 3-5-foliolate. .......................................................................................................................... 3 

2a. Calyx of the staminate flowers urceolate, with deltate lobes; corolla papillose outside; anthers 

broadly lanceolate............................................................................................................. 2. P. pedata 

2b. Calyx of the staminate flowers cylindrical, with lanceolate-acuminate lobes; corolla puberulous 

outside; anthers subulate ............................................................................................. 1. P. ottoniana 

3a. Pistillate flowers solitary or in pairs; calyx cylindrical-urceolate, ca. 12 mm long; petals 6-7 mm 

long ....................................................................................................................... 2. P. pedata 

3b. Pistillate flowers in groups of 2-4, axillary; calyx fusiform, 3-3.2 cm long; petals ca. 2.8 cm long 

............................................................................................................................. 3. P. trifoliata 

1. Psiguria ottoniana (Schltdl.) C. Jeffrey, Kew 

Bull. 33: 352. 1978. 

Fig. 87. A-B 

BASIONYM: Anguria ottoniana Schltdl. 

Herbaceous vine, attaining 3 m in length and 

climbing by axillary tendrils. Stems slender, 

glabrous; tendrils simple, filiform, ca. 15 cm long. 

Leaves alternate; blades 9-17 × 8-17 cm, deeply 

3-5-lobed, glabrous, the lobes acute or acuminate, 

mucronate, the base deeply cordiform, the margins 

entire, bidentate toward the apex of the lobe; 

petioles ca. 3.5 cm long, slender, glabrous. 

Staminate flowers 3-8, in axillary racemes; 

peduncle longer than the accompanying leaf; 

calyx green, cylindrical, slightly dilated at the 

base, the lobes lanceolate-acuminate; corolla 

orange, the lobes oblong, acute at the apex, 

puberulous outside; anthers subulate. Pistillate 

flowers and fruits unknown. 

Phenology: Unknown. 

Status: Native, known from Puerto Rico from 

a single collection (Plée 42) at the beginning of 

the 19th century. This collection was identified 

by Cogniaux, in his time a specialist in the 

Cucurbitaceae, and is found deposited in the 

Museum of Natural History in Paris. 

Distribution: Its distribution in Puerto Rico is 

unknown, since the collection of Plée did not 

include the specific locality. It is also found in 

Cuba and the Bahamas (according to Jeffrey and 

Trujillo, 1992).

218 


2. Psiguria pedata (L.) R. A. Howard, J. Arn. 

Arb. 54: 441. 1973. 

Fig. 87. C-G 

BASIONYM: Cucumis pedatus L. 

SYNONYMS: Psiguria trilobata (L.) R. A. Howard 

Cucumis trilobatus L. 

Anguria trilobata Jacq. 

Anguria plumeriana Schltdl. 

Monoecious vine, herbaceous, 3-4 m in length 

and climbing by tendrils. Stems green, slender, 

striate, glabrous, somewhat fleshy; mature stems 

fleshy, with the bark very thick, corky, and 

fissured, with winged longitudinal projections; 

tendrils lateral, simple, filiform, 10-15 cm long. 

Leaves alternate; blades 7-13.5 × 5-13.5 cm, 

broadly ovate, deltoid, 3-lobed, 3-5-foliolate, 

membranaceous or chartaceous, the venation 5palmate, 

the apex acute or acuminate, the lobes 

obtuse, the base cordiform or hastate, the margins 

undulate, denticulate, ciliate; upper surface dark 

green, dull, puberulous, pustulose; lower surface 

puberulous, sometimes pustulose, light green, 

dull, with prominent venation; petioles slender, 

glabrous, 2.5-5 cm long. Staminate flowers in 

axillary racemes; peduncle as long as or longer 

than the accompanying leaf; calyx yellowish 

green, urceolate, 5-7 mm long, the lobes deltate; 

corolla orange, tubular, ca. 11 mm long, papillose 

outside, the lobes oblong, ca. 2 cm long; anthers 

sessile, compressed, lanceolate, 5-8 mm long; 

pistillode absent. Pistillate flowers solitary or in 

pairs, axillary, peduncles shorter than the 

accompanying leaf; calyx and hypanthium 

fusiform or tubular, 3-3.2 cm long, the lobes 

subrounded, apiculate, ca. 2 mm long; corolla 

orange, ca. 2.8 cm long, the lobes free almost to 

the base, obovate. Berry ellipsoid, smooth, ca. 4 

cm long; seeds numerous, elliptical, ca. 7 mm 

long. 

13. SECHIUM 


May, July, and October and in fruit in July and 

November. 


Distribution: In moist forests along the 

Cordillera Central and in the zone of mogotes. 

Cited for St. Croix by Cogniaux (1916); also in 

Hispaniola, Cuba, and the Bahamas. 

Public forest: Río Abajo and Vega. 

3. Psiguria trifoliata (L.) Alain, Phytologia 47: 

192. 1980. 

Fig. 89. H 

BASIONYM: Anguria trifoliata L. 

SYNONYM: Anguria plumeriana var. trifoliata Cogn. 

Anguria cookiana Britton 

Herbaceous vine, which climbs by tendrils. 

Stems thick, striate, glabrous, somewhat fleshy; 

tendrils lateral, simple, filiform, as long as the 

accompanying leaf. Leaves alternate, 3-foliolate, 

glabrous; leaflets chartaceous, acute, mucronate, 

the margins entire or crenate-serrate, the central 

leaflet 7-10 × 3-5 cm, elliptical, the lateral leaflets 

5-9 × 2-4 cm, asymmetrically ovate-lanceolate; 

petioles slender, glabrous, 1-4 cm long. Staminate 

flowers 10-20 in axillary racemes; peduncle as 

long as or longer than the accompanying leaf; 

calyx obovoid to subcylindrical, glabrous, 8-12 

mm long, the lobes triangular, ca. 1 mm long; 

petals oblong, 1-2 cm long. Pistillate flowers in 

groups of 2-4, axillary, peduncles short. Berry 

ovoid-elongate, ca. 4 cm long, smooth, variegated; 

seeds numerous, elliptical, ca. 7 mm long, without 

an aril. 

Phenology: Collected in fruit in November. 


Distribution: Known from few old collections 

from Coamo, Vega Baja, and Cayey. Also in 

Hispaniola. 

Herbaceous vines, monoecious, climbing by axillary tendrils, 3-5-fid; without latex; stem tuberous 

at the base. Leaves alternate, petiolate; blades simple, palmatilobed; stipules absent. Flowers unisexual, 

actinomorphic; calyx campanulate, with 5 minute lobes; corolla campanulate or rotate, the lobes deep. 

Staminate flowers in axillary racemes or panicles with elongate peduncles; stamens 5, the filaments 

united to one another to form a central column, the anthers sigmoid; pistillodes absent. Pistillate 

flowers solitary or in pairs, smaller than the staminate flowers, axillary to the staminate inflorescence; 

ovary inferior, obovoid, unicarpellate, with a pendulous ovule, solitary, the style terminal, simple,


220 


short, the stigma subglobose. Fruit indehiscent, 

fleshy, pyriform, sulcate, spinulose; seed one, 

large, compressed, germinating inside the fruit. 

A genus native to Central America, of about 5 

species. 

1. Sechium edule (Jacq.) Sw., Fl. Ind. Occ. 1150. 

1800. 

Fig. 88. A-E 

BASIONYM: Sicyos edulis Jacq. 

Chayote, Cho-cho 

Herbaceous vine climbing by axillary tendrils, 

10 or more m in length. Stems much branched, 

greenish, striate, pubescent when young, glabrous 

when mature, sometimes pubescent at the nodes; 

tendrils axillary or lateral to the leaves, thick, with 

4 or 5 branches that intertwine in a spiral. Leaves 

alternate; blades 6-22 × 6-22 cm, ovate or 

rounded, 3-5-lobed-angled, chartaceous, the 

venation 5-9-palmate, the lobes acute at the apex, 

the base cordiform, the margins entire or 

denticulate; upper surface dark green, scabrous, 

14. SICANA 

with the venation slightly prominent; lower 

surface yellowish green, with prominent venation, 

pubescent; petioles thick, 4-15 cm long. Staminate 

flowers in axillary racemes; calyx green, 

campanulate, with 5 minute lanceolate lobes; 

corolla yellowish green, the lobes oblong, 6-8 mm 

long; stamens yellow. Pistillate flowers solitary 

on a long pedicel axillary to the staminate 

inflorescence; calyx green, campanulate, with 

oblong lobes, ca. 4 mm long; corolla yellowish 

green, with five deep lobes, oblong, 10-12 mm 

long; ovary inferior, obovoid-fusiform, pubescent. 

Fruit fleshy, pyriform, sulcate, 8-16 cm long, 

green or white, usually spinulose. Seed one, 

lenticular, germinating inside the fruit. 



Status: Exotic, cultivated and naturalized, very 

common. 

Distribution: In disturbed areas. Native to 

Central America, widely cultivated throughout the 

tropics for its edible fruits. 


Herbaceous vines, monoecious, climbing by axillary tendrils, 3-5-fid. Leaves alternate, petiolate; 

blades simple, 5-9-lobed; stipules absent. Flowers unisexual, actinomorphic; calyx campanulate, 

with 5 reflexed lobes; corolla campanulate, with 5 deep oblong-ovate lobes. Staminate flowers solitary; 

stamens 3, the filaments short, inserted on the receptacle, the anthers sigmoid, concrescent in a 

capitulum; pistillodes absent. Pistillate flowers solitary, similar to the staminate flowers; staminodia 

3, minute; ovary inferior, subovoid, tricarpellate, with numerous horizontal ovules, the style terminal, 

simple, short, the stigmas 3, slightly bilobate, papillose. Fruit a cylindrical berry, smooth; seeds 

numerous, oblong, compressed, not arillate. A genus of 4 species, one of which is cultivated throughout 

Latin America; of the remaining species, one is endemic to Jamaica, another to Trinidad and Tobago, 

and another to the Dominican Republic. 

1. Sicana odorifera (Vell.) Naudin, Ann. Sci. Nat. 

Bot. Sér. 4: 18. 181. 1862. 

Fig. 88. F-G 

BASIONYM: Cucurbita odorifera Vell. 

Pepino angolo, Casabana, Curuba, Pepino 

Herbaceous vine, climbing by tendrils, 

attaining 8 m or more in length. Stems green, 

thick, sulcate, puberulent or glabrescent; tendrils 

axillary, 5-fid, 9-11 cm long. Leaves alternate; 

blades 10-20 × 10-20 cm, 3-5 (7)-lobed, 

chartaceous, the lobes deep, acute, the base 

cordiform, the margins undulate or denticulate; 

upper surface dark green, slightly shiny, glabrous; 

lower surface pale green, with prominent 

venation, glabrous; petioles 7-13 cm long, slender, 

canaliculate or slightly winged, villous. Flowers 

unisexual, ascending, solitary, axillary; peduncle 

shorter than the accompanying leaf. Calyx green, 

campanulate, 6-8 mm long, tomentose, the lobes


222 


reflexed, 10-12 mm long, lanceolate; corolla 

yellow, campanulate, 2.5-4 cm long, tomentose; 

stamens 3; staminodia 3; ovary ovoid, 5-6 mm 

long, puberulent. Berry pendulous, cylindrical or 

ellipsoid, 30-50 cm long and ca. 10 cm in 

diameter, yellowish brown or mulberry-colored; 

seeds, numerous, asymmetrically ovate, 

compressed, with the margin darker, 13.5-15 mm 

long. 


during August. 

Status: Exotic, cultivated, common. 

Distribution: Species native to South America, 

cultivated in Puerto Rico and the Greater Antilles 

for its edible fruits, which are found for sale in 

the public markets. 

References: Cogniaux, A. 1916. Cucurbitaceae-Fevilleae et Melothrieae. Pp. 1-246. In: A. Engler 

(ed.). Das Pflanzenreich. IV. 275. Jeffrey, C. and B. Trujillo. 1992. Cucurbitaceae. Flora de Venezuela. 

Vol. 5(1): 11-201. Kirkbride, J. H. 1993. Biosystematic monograph of the genus Cucumis (Cucurbitaceae). 

Parkway Publishers. Boone, North Carolina. Purseglove, J.W. 1982. Tropical crops. Dicotyledons. 

Longman Group Ltd.; Whitaker, T.W. and G.W. Bohn. 1950. The taxonomy, genetics, production and 

uses of the cultivated species of Cucurbita. Econ. Bot. 4: 52-81. 

20. Family CUSCUTACEAE 

1. CUSCUTA 

Herbs with more or less long twining stems, without chlorophyll, parasites by means of haustoria. 

Leaves alternate, rudimentary, reduced to minute scales; stipules absent. Flowers minute, bisexual, 

actinomorphic, in axillary heads or spikes; calyx crateriform, of 4-5 free or connate sepals; corolla 

white or pink, tubular, with 4-5 lobes; stamens 4-5, the filaments adnate to the corolla tube, with a 

basal appendage on the ventral surface, the anthers longitudinally dehiscent; ovary superior, bicarpellate, 

each carpel with 2 ovules, the styles 2, free. Fruit a capsule, circumscissile or with irregular dehiscence, 

or an indehiscent berry; seeds (1)2-4 per fruit, minute. 

1. Cuscuta americana L., Sp. Pl. 124. 1753. 

Fig. 89. A-H 

Fideillo, Tente en el aire 

Herbaceous parasitic vine, attaining 5 m in 

length, with numerous lateral twining branches 

that adhere by means of haustoria. Stems yelloworange, 

cylindrical, smooth, glabrous, 1.5-2.5 mm 

in diameter. Leaves vestigial, 1-2 mm long, 

yellowish, in the form of scales. Flowers bisexual, 

sessile, in short axillary cymes. Calyx crateriform, 

whitish, membranaceous, 2.3-2.5 mm long, with 

5 minute lobes; corolla whitish, crateriform to 

tubular, ca. 2.3 mm long, with 5 minute and 

rounded lobes; stamens 5, adnate to the upper 

portion of the corolla tube, the filaments with a 

scalloped scale at the base; ovary depressedglobose. 

Capsule thin-walled, 1.5-2 mm long, 

opening longitudinally in the area between the 

two styles. Seed one or rarely two per fruit, 

subglobose or lenticular, ca. 1.5 mm long, smooth, 

light brown. 



Status: Native, very common 

Distribution: In coastal thickets. Also on 

Desecheo, Mona, Vieques, St. Croix, St. John, St. 



Public forests: Ceiba, Guánica, and Mona.


21. Family DILLENIACEAE 


1a. Leaves acuminate at the apex; ovary unicarpellate; inflorescence fasciculate; fruits subglobose, 

pubescent …..……………...…………………...………………......................1. Doliocarpus 

1b. Leaves obtuse or apiculate at the apex; ovary bicarpellate; inflorescence paniculate; fruits bilobate, 

glabrous ..……………………………………………………….……....................….2. Pinzona 

1. DOLIOCARPUS 

Twining vines; stems thick, sometimes attaining 30 cm in diameter; cross section with concentric 

rings of xylem alternating with concentric rings of parenchyma. Leaves alternate, simple, entire or 

serrate, petiolate; stipules absent or of short duration. Inflorescences of fascicles or glomerules. Flowers 

bisexual, actinomorphic. Calyx of 3-6 free sepals, subequal, generally the inner ones larger, imbricate, 

persistent; petals 2-6, free, early deciduous, white; stamens numerous, the filaments unequal, the anthers

224 


dehiscent by longitudinal sutures; ovary superior, unicarpellate, with two basal ovules, the style terminal, 

filiform, the stigma punctiform to peltate. Fruit a berry or capsule with irregular dehiscence; seeds 

usually 2, reniform, black, covered by a white arillode, membranaceous or fleshy. A neotropical genus 

of about 40 species.


1. Doliocarpus brevipedicellatus Garcke, 

Linnaea 22: 47. 1849. 

Fig. 90. A-J 

Twining liana attainig 5 m or more in length. 

Branches angular, appressed-pubescent. Leaves 

5-16 × 2.5-8 cm, lanceolate or less frequently 

elliptic-lanceolate, coriaceous or subcoriaceous, 

the upper surface pilose on the primary vein, the 

lower surface strigose on the secondary veins, the 

apex acuminate, the base cuneate or rounded, the 

margins deeply dentate-mucronate; petioles 0.5- 

1.2 cm long, strigose. Inflorescences fasciculate, 

racemose, with 3-5 flowers; peduncles and 

pedicels appressed-pubescent. Calyx of 4-5 

unequal sepals, pilose outside, the outer ones 1.5- 

2 mm long, lanceolate, the inner ones 3-3.5 mm 

long, ovate-elliptical; petals 2-3 mm long, white; 

stamens 25-40; pistil glabrous. Capsules 4-7 mm 

in diameter, irregularly dehiscent. Seed one. 

Phenology: Collected in flower in March. 

Status: Native, uncommon in Puerto Rico. 

2. PINZONA 

A monotypic genus characterized by the following species. 

1. Pinzona coriacea Mart. & Zucc. in Zucc., Abh. 

Math.-Phys. Cl. Königl. Bayer. Akad. Wiss. 

1: 371. 1832. 

Fig. 1C; 91. A-K 

SYNONYM: Doliocarpus calinoides (Eichl.) Gilg. 

Bejuco de agua 

Twining liana attainig 30 m in length; stems 

cylindrical, up to 10 cm in diameter, producing 

abundant potable water when cut; bark reddish 

brown, peeling in a scaly manner; cross section 

with concentric rings of xylem alternating with 

rings of parenchyma. Branches puberulent, 

angular, scabrous, glabrescent, and cylindrical 

when mature. Leaves alternate, 8.5-22 × 3-12.5 

cm, broadly elliptical, ovate to obovate, 

coriaceous, the apex rounded, sometimes shortapiculate, 

the base rounded to subcordiform, the 

margins revolute, sinuate, or dentate-mucronate; 

upper surface scabrid, sometimes with the veins 

Distribution: Known from the zone of 

mogotes. Its distribution in the Neotropics is 

rather widespread, being found in Hispaniola, 

Nicaragua, Panama, Colombia, Venezuela, the 

Guianas, Brazil, and Bolivia. 

Commentary: This species was treated by 

Liogier (1994) in his “Descriptive Flora of Puerto 

Rico and adjacent islands, Vol. 3” as a synonym 

of Doliocarpus major Gmel. This error was 

initiated by Hunter in 1965, but corrected by 

Kubitzki in 1971. Doliocarpus major is a South 

American species that is distinguished from D. 

brevipedicellatus, among other things, by having 

longer inflorescences and papillae on the lower 

surface of the leaves. 

Excluded species: Doliocarpus olivaceus 

Sprague & R.O. Williams ex Standl. was reported 

for Puerto Rico by Liogier (1994) based on the 

erroneous identification of a specimen of Pinzona 

coriacea. This species is known from Venezuela, 

the Guianas, Brazil, and Bolivia, but not from 

the Antilles. 

appressed-pubescent; lower surface with 

prominent venation, papillose; petioles thick, 1- 

3.5 cm long, winged, with the base decurrent to 

half the diameter of the branch. Inflorescences of 

axillary panicles, 3-7 cm long, pilose; bracts 

oblong to ovate, 1-2 mm long. Calyx of 3-4 

subequal sepals, ca. 2 mm long; petals 3, obovate, 

longer than the sepals; stamens 25-30, the 

filaments sinuate; ovary superior, bicarpellate, 

biglobose, glabrous. Fruit capsular, bilobate, 

bilocular, crustose, tardily dehiscent. Seeds 2 per 

fruit, with an orange arillode. 

Phenology: Collected in flower in May and 

June and in fruit in November. 


Distribution: In moist, mature forests of 

montane areas, in the Cordillera Central and the 

Sierra de Luquillo. From Belize to Bolivia, 


Public Forests: Carite and El Yunque.

226 



References: Aymard, G.A., 1998. Dilleniaceae. In: Steyermark et al. (eds.), Flora of the Venezuelan 

Guayana. Vol. 4; Aymard, G.A. 1997. Dilleniaceae Nova Neotropicae, V. El género Doliocarpus en 

Colombia. Anal. Jardín Bot. Madrid 55: 17-30; Hunter, G.E. 1965. Flora of Panama Part VI. Family 

118. Dilleniaceae. Ann. Missouri Bot. Gard. 52: 580-598; Kubitzki, K. 1971. Doliocarpus, Davilla 

und verwandte Gattungen (Dilleniaceae). Mitt. Bot. München 9: 1-105. 

22. Family ERICACEAE 


1a. Corolla tubular or tubular-campanulate, 1.4-1.8 cm long, white, pink, or red; leaves rounded or 

rounded-ovate, 1-2.7 cm long, the lateral veins inconspicuous; flowers solitary (in our 

species)…………………………..…………………………………………........…1. Gonocalyx 

1b. Corolla campanulate, 6-8 mm long, yellowish green; leaves 4-11 cm long, usually ovate, with the 

secondary and tertiary veins visible; flowers in terminal racemes ………...…. 2. Vaccinium 

1. GONOCALYX 

Epiphytic or terrestrial shrubs, erect or clambering. Leaves alternate, entire, coriaceous, shortpetiolate. 

Inflorescences of terminal racemes or the flowers solitary or in pairs. Flowers bisexual, 

actinomorphic. Calyx tubular or tubular-campanulate, ribbed or smooth, with 4-5 lobes at the apex; 

corolla campanulate to tubular, red, pink, or white, with 4-5 lobes; stamens 8-10, the filaments shorter 

than the anthers, adnate to the base of the corolla, the anthers with two thecae, whose distal portion is 

elongate and tubular, with a terminal pore or with longitudinal sutures; disc 10-lobate; ovary inferior, 

4-5-locular, the style terminal, single, the stigma truncate. Fruit a berry, usually with 4-5 longitudinal 

ribs; seeds numerous, minute. A genus of 8 species, distributed in Hispaniola, Puerto Rico, the Lesser 

Antilles, Costa Rica, and Colombia. 

Key to the species of Gonocalyx 

1a. Calyx green; corolla tubular-cylindrical, white or pink, with the lobes erect; stigma white, slightly 

exserted; leaves rounded or ovate-elliptical, the margins markedly revolute (the blade convex), the 

apex rounded .……………………………………………...............................…2. G. portoricensis 

1b. Calyx red; corolla tubular-campanulate or campanulate, red, with the lobes expanded; stigma red, 

exserted; leaves ovate, ovate-elliptical, or rarely rounded, the margins slightly revolute (the blade 

almost flat), the apex acute or obtuse .….……………………………..............1. G. concolor 

1. Gonocalyx concolor Nevling, J. Arn. Arb. 

51: 222. 1970. 

Fig. 92. A-D 

Epiphytic or terrestrial shrub, usually 

clambering, attainig 5 m in length. Stems much 

branched, slender, cylindrical, dark brown, the 

twigs minutely dense-pubescent. Leaves alternate, 

ovate, ovate-elliptical, or less frequently rounded, 

flat, rigid-coriaceous, 1.5-2.7 × 1-2 cm, the apex 

obtuse or acute, sometimes rounded, mucronate, 

the base obtuse or rounded, the margins slightly 

revolute or flat, entire or crenulate; upper surface 

glabrous, with the midvein sunken; lower surface

228 


pale green, shiny, with scattered papilliform hairs, 

the midvein prominent, the lateral venation 

inconspicuous; petioles 2-2.5 mm long. Flowers 

solitary, axillary, pendulous; peduncle 7-11 mm 

long. Calyx campanulate, red, with 5 longitudinal 

ribs, 3-3.5 mm long; corolla tubular-campanulate, 

red, ca.18 mm long, with 5 broadly deltate, 

expanded lobes; stamens 8, the filaments ca. 2 

mm long, pubescent, the anthers with the distal, 

tubular portion of the theca 4-5 times longer than 

the theca, projecting slightly outside the corolla; 

ovary inferior, the style thick, the stigma red, 

exserted. Fruits subglobose-5-angled, wine-red, 

truncate, 6.5-9 mm long (Kolterman et al. 

unpublished data). 

Phenology: Collected in flower during April 

and July. 

Status: Endemic to Puerto Rico, quite rare. 

Distribution: Known only from the Carite 

forest. 

Public Forest: Carite. 

2. Gonocalyx portoricensis (Urb.) A.C. Smith, 

Contr. U. S. Natl. Herb. 38: 354. 1932. 

Fig. 92. E-I 

BASIONYM: Thibaudia portoricensis Urb. 

SYNONYM: Ceratostema portoricensis (Urb.) Hoerold 

Clambering shrub attainig 3 m in length. 

Stems much branched, slender, cylindrical, dark 

2. VACCINIUM 

brown, the twigs reddish, densely pubescent. 

Leaves alternate, reddish when young, circular 

or broadly ovate, convex, rigid, 0.8-2 × 0.6-1.2 

cm, the apex obtuse or rounded, mucronate, the 

base obtuse or rounded, the margins markedly 

revolute, entire or denticulate; upper surface 

green, shiny, glabrous, with the midvein sunken; 

lower surface pale green, shiny, glabrous or with 

scattered papilliform hairs, with the midvein 

prominent and the lateral venation inconspicuous; 

petioles 1.5-3 mm long. Flowers solitary, axillary, 

pendulous; peduncle 6-12 mm long. Calyx 

campanulate, with 5 longitudinal ribs, 3-3.5 mm 

long; corolla tubular, pale pink or white, ca. 16 

mm long, with 5 erect, deltate lobes; stamens 10, 

the filaments ca. 1 mm long, pubescent on the 

upper margin, the anthers with the distal, tubular 

portion of the theca 4-5 times longer than the 

theca, not exserted; ovary inferior, the style thick, 

the stigma white, slightly exserted. Fruits 

depressed-globose, white or red, 4-7 mm in 

diameter. 

Phenology: Flowering and fruiting from the 

end of January to August. 

Status: Endemic to Puerto Rico, rather 

common. 

Distribution: On mountaintops, usually in 

dwarf forest, along the Cordillera Central and the 

Sierra de Luquillo. 



Shrubs, erect or less frequently clambering, or rarely trees. Leaves alternate, pinnately veined, 

entire or serrate, coriaceous, petiolate. Inflorescences of axillary racemes, less frequently the flowers 

solitary or in pairs; pedicels usually bibracteate. Flowers bisexual, actinomorphic. Calyx tubular to 

globose, with 4-5 lobes at the apex; corolla cylindrical, urceolate, or campanulate, white, green, red, or 

yellowish, with 4-5 more or less deep lobes; stamens 8-10, as long as the corolla; ovary inferior or 

partially inferior, 4-5-locular, the style terminal, single, the stigma simple or capitate. Fruit a berry, 

with the sepals persistent on the apical portion; seeds 5-numerous, minute. A genus of about 300 

species, mostly of the Northern Hemisphere and in the montane forests of the tropics. 

1. Vaccinium racemosum (Vahl) Wilbur & 

Luteyn, Brittonia 29: 275. 1977. 

Fig. 92. J-M 

BASIONYM: Hornemannia racemosa Vahl 

SYNONYMS: Symphysia racemosa (Vahl) Stearn 

Thibaudia krugii Urb. & Hoerold 

Woody shrub, scandent or clambering, attainig 

3-5 m in length. Stems much branched, coppercolored, 

more or less cylindrical, striate, attaining 

up to 3 cm in diameter, the nodes swollen. Leaves 

alternate, reddish when young, glabrous, 

coriaceous, slightly brittle, ovate or elliptical, 4-


230 


14 × 2-8 cm, in general convex, the venation 

pinnate, usually from near the base, the apex 

acuminate, the base rounded or obtuse, the 

margins entire or undulate, revolute; upper surface 

dark green, shiny, with the venation generally 

sunken, translucent and yellowish; lower surface 

yellowish green, with the palmate venation 

prominent; petioles 5-7 mm long, thick, rugose. 

Inflorescences of terminal corymbiform racemes; 

pedicels thick, 10-12 mm long. Calyx 

campanulate, ca. 5 mm long, green, smooth, with 

five minute lobes; corolla campanulate, early 

deciduous, yellowish green, ca. 8 mm long, the 

lobes ovate, ca. 5 mm long, reflexed; stamens 10, 

the filaments white, flattened, adnate to the base 

of the corolla, the anthers golden yellow, the distal 

tube as long as the theca; ovary inferior, the style 

simple, the stigma subcapitate. Fruit fleshy, 

globose, 5-8 mm in diameter, cardinal red, with 

the calyx persistent on the apical portion. Seeds 

numerous, ca. 1 mm long, foveate, obtusetrigonal. 

Phenology: Flowering and fruiting from June 

to March. 


Distribution: From high and moist regions 


Luquillo. Also throughout the Antilles. 



References: Luteyn, J.L. 1976. Notes on neotropical Vaccinieae (Ericaceae). 1. Gonocalyx-A genus 

new to Central America. Brittonia 28: 37-41. Luteyn, J.L. and R.L. Wilbur. 1977. New genera and 

species of Ericaceae (Vaccinieae) from Costa Rica and Panama. Brittonia 29: 255-276. Luteyn, J.L. 

1996. Ericaceae. In: Harling, G. and L. Andersson (eds.) Flora of Ecuador. Vol. 54. 

23. Family EUPHORBIACEAE 


1a. Inflorescences of cymes with two trilobate foliaceous bracts at the base, forming a pseudanthium; 

staminate flowers with numerous stamens; leaves deeply trilobate (in our species) 

…………….………..………………………….………………..................…1. Dalechampia 

1b. Inflorescences of racemes, lacking foliaceous bracts; staminate flowers with 2-3 stamens; leaves 

simple (in our species) ..………………………..……..………….....................………. 2. Tragia 

1. DALECHAMPIA 

Twining vines or less frequently shrubs, monoecious, usually covered with stinging hairs, producing 

scarce watery latex when wounded. Leaves simple, alternate, 3-5-lobate, petiolate, with a pair of stipels 

at the base of the blade; stipules present. Flowers unisexual, actinomorphic, apetalous, clustered in 

bisexual cymes, with long peduncles and two foliaceous bracts, forming a pseudanthium. Staminate 

flowers in distal cymules; bracteoles with numerous resinous glands; calyx 4-6-valvate; stamens 

numerous, grouped on a short stipe to form a head, the anthers short, opening along longitudinal 

sutures; pistillode absent. Pistillate flowers in basal cymules; calyx with 8-12 lobes with glandular 

margins; ovary superior, trilobate, tricarpellate, each carpel with a solitary ovule, the style simple, with 

a capitate or peltate stigma. Fruit a trilobate capsule, with one seed per locule. A genus of about 100 

species, mostly of the Neotropics.


1. Dalechampia scandens L., Sp. Pl. 1054. 

1753. 

Fig. 93. A-E 

Herbaceous vine, twining, attainig 5 m in 

length. Stems cylindrical, slender, pilose. Leaves 

alternate, membranaceous, deeply trilobate, 5-7 

× 6-12 cm, the apex acute or acuminate, the base 

cordiform, the margins denticulate; upper surface 

puberulent, with slightly prominent venation; 

lower surface pilose, with prominent venation; 

petioles 6-8 cm long, pilose; stipules ovate or 

lanceolate, 0.5-1 cm long. Pseudanthium on 

peduncles 4 cm long; bracts 3(-5)-lobate, up to 

2.5 cm long, foliaceous, with the margins serrate, 

with stinging glandular hairs. Staminate flowers 

2. TRAGIA 

in distal cymules. Calyx of 4 sepals, ca. 1.5 mm 

long; stamens numerous, forming a head. 

Pistillate flowers in basal cymules. Calyx of 10- 

12 sepals; ovary trilobate. Capsule trilobate, 

puberulent, ca. 5 mm long. Seeds globose, ca. 3 

mm in diameter, with black spots. 




Distribution: In coastal or dry forests and 

thickets. Also on Vieques, St. Croix, St. John, St. 


the Antilles (except for Jamaica) and from Mexico 

to South America. 

Public Forests: Boquerón, Guánica, and 

Susúa. 

Twining vines or less frequently erect herbs, monoecious, covered with stinging hairs, producing 

scarce watery latex when wounded. Leaves alternate, entire or 3-lobate, petiolate, the margins serrate 

or entire; stipules minute, deciduous. Flowers unisexual, actinomorphic, apetalous, in bisexual racemes, 

axillary or terminal. Staminate flowers short-pedicellate, numerous. Calyx 3-4-valvate; stamens (2-)3- 

5, the filaments connate at the base; pistillode minute. Pistillate flowers long-pedicellate. Calyx of 3 or 

6 sepals; ovary tricarpellate, each carpel with a solitary ovule, the style simple, with 3 stigmatic branches. 

Fruit a trilobate capsule with explosive dehiscence, with one seed per locule. A genus of about 150 

species, of tropical, subtropical, and subtemperate distribution. 

1. Tragia volubilis L., Sp. Pl. 980. 1753. 

Fig. 93. F-J 

Pringamosa, Bran nettle 

Climbing or creeping vine, twining, attainig 

1-3 m in length. Stems slender, with appressed 

and glandular stinging hairs. Leaves alternate, 

usually clustered on short lateral branches, 

membranaceous, oblong, ovate, or lanceolate, 2- 

8 × 0.9-4 cm, the apex acuminate, the base 

subtruncate or cordiform, the margins serrate; 

upper surface pale green, with stinging glandular 

hairs, the midvein slightly prominent; lower 

surface pale green, with prominent venation, 

puberulent; petioles slender, 0.3-6 cm long; 

stipules subulate, ca. 4 mm long. Flowers 

greenish, in axillary racemes. Staminate flowers 

numerous, along the length of the inflorescence. 

Calyx 0.8-1 mm long; stamens 2-3. Pistillate 

flowers on long pedicels at the base of the raceme. 

Calyx ca. 1 mm long, pilose; ovary hispid. Capsule 

with three rounded lobes, ca. 7 mm in diameter, 

covered with stinging hairs. Seeds globose, ca. 2 

mm in diameter, brown, smooth. 




Distribution: In pastures and secondary 

forests, at lower elevations, in the limestone zone 

and in dry areas. Also on Culebra, Vieques, St. 


Gorda; throughout tropical America. 

Public Forests: Cambalache, Ceiba, 

Guajataca, Guánica, Maricao, Río Abajo, and 

Susúa.

232 



24. Family FABACEAE 

Key to the subfamilies 

1a. Flowers zygomorphic; corolla of free petals; inflorescences usually racemose.................……...……2 

1b. Flowers actinomorphic; corolla of connate petals; inflorescences usually of densely flowered heads 

or spikes...............................…………………………………………………..…Mimosoideae 

2a. Corolla of 5 equal or almost equal petals ……..……………………….... Caesalpinioideae 

2b. Corolla with the central petal (standard) elongate, 2 lateral ones connate into a keel and 2 lateral 

ones smaller .......................…………….………………………….…………………..... Faboideae 

24a. Subfamily CAESALPINIOIDEAE 


1a.Leaves bipinnate; rachis lacking stipitate glands; fruits slightly flattened....1.Caesalpinia 

1b. Leaves pinnate; rachis with a stipitate gland; fruits cylindrical …...............……..…………2. Senna 

1. CAESALPINIA 

Trees or erect or clambering shrubs. Stems usually spiny. Leaves bipinnate; pinnae opposite, the 

leaflets opposite or alternate; petioles and rachis lacking stipitate glands; stipules minute to foliaceous. 

Flowers unisexual or bisexual, in axillary or terminal racemes; pedicels articulate in the distal portion. 

Calyx campanulate, 5-lobate; corolla of various colors, the petals 5, free; stamens 10, the filaments 

flattened, free, of equal length, the anthers dehiscent along longitudinal sutures; ovary unilocular, 

superior, sessile or short-stipitate, with numerous ovules. Fruit a legume of various forms, dehiscent or 

indehiscent; seeds solitary or numerous, of various forms. A tropical genus of about 100 species. 

Key to the species of Caesalpinia 

1a. Stipules foliaceous, up to 1.5 cm long, persistent; seeds gray ………………. 1. C. bonduc 

1b. Stipules minute, < 5 mm long, deciduous; seeds yellow-orange, brown, or black……...…2 

2a. Legumes not spiny……………………………………………………………………...….3 

2b. Legumes spiny……………………………………………………………………………..5 

3a. Stems spiny; leaflets oblong; legume 2-3 cm broad; seeds oblong ………. 4. C. decapetala 

3b. Stems not spiny; leaflets elliptical or broadly elliptical; legume ca. 4 cm broad; seeds almost globose 

...........................................................................................................................................4

234 


4a. Rachis of the leaf with minute spines; leaves with 3-5 pairs of pinnae; seeds dark brown or almost 

black......................…………………………...……………………………..…6. C. portoricensis 

4b. Rachis of the leaf not spiny; leaves with 7 pairs of pinnae; seeds yellow-orange 

.......................................................................................................................... 3. C. culebrae 

5a. Leaflets 4-8 cm long, acute or short-acuminate at the apex; petals ca. 15 mm long 

..............……………………………………………………………………………….. 5. C. major 

5b. Leaflets 1-3 cm long, obtuse, rounded, or emarginate at the apex; petals 7-8 mm long 

..............……………………………………………………………………………….… 2. C. ciliata 

1. Caesalpinia bonduc (L.) Roxb., Fl. Ind. ed. 

2, 2: 362. 1832. 

Fig. 94. A-C 

BASIONYM: Guilandina bonduc L. 

SYNONYM: Caesalpinia crista sensu Urb. and sensu 

Britton & Wilson 

Mato de playa 

Scandent shrub, much branched along the 

main stem, attainig 2-6 m in length. Stems 

cylindrical, up to 2.5 cm in diameter, densely 

spiny. Leaves bipinnate, paripinnate, 30-75 × 10- 

34 cm; pinnae 4-8 pairs, opposite; leaflets 3-7 

pairs per pinna, opposite, 2.5-5(6.8) × 1.2-3 cm, 

ovate, lanceolate, oblong, or elliptical, 

chartaceous, glabrous or puberulent, the apex 

obtuse, mucronate, the base truncate, rounded to 

almost cordiform, the margins entire, revolute, 

ciliate; upper surface dull; lower surface with the 

midvein prominent; petiolules short, cylindrical; 

petioles and rachis spiny; stipules foliaceous, up 

to 1.5 cm long, persistent, divided into 3-5 

segments. Flowers functionally unisexual, in 

axillary or terminal racemes; rachis tomentose and 

densely spiny; bracts lanceolate, 8-14 mm long, 

persistent. Calyx campanulate, 4-6 mm long, 

tomentose, the sepals reflexed, oblong. Petals 

yellow, 5.5-8 mm long, spathulate; stamens 10, 

of which 4 are fertile in the staminate flowers, all 

sterile in the pistillate flowers; ovary sessile, 

rudimentary in the staminate flower, the stigma 

terminal. Legumes oblong, 4-7.5 × 2-4 cm, semiinflated, 

densely spiny, tardily dehiscent along 

both sutures. Seeds usually 2, ovoid or almost 

globose, 1.5-2 cm long, gray. 


October to March. 



littoral zone. Also on Culebra, Mona, Vieques, 

Anegada, St. Croix, St. John, St. Thomas, Tortola, 

and Virgin Gorda; along the tropical littoral 

coasts. 

Public Forests: Guánica, Mona, and Piñones. 

2. Caesalpinia ciliata (Bergius ex Wikstr.) 


Fig. 94. D-F 

BASIONYM: Guilandina ciliata Bergius ex Wikstr. 

SYNONYM: Caesalpinia bonduc sensu Urb. 

Guilandina melanosperma Eggers 

Caesalpinia divergens Urb. 

Mato de playa 

Scandent shrub, with many lateral branches 

along the main stem, attainig 6 m in length. Stems 

cylindrical, up to 2.5 cm in diameter, densely 

spiny. Leaves bipinnate, paripinnate, 20-35 × 10- 

28 cm; pinnae 3-10 pairs, opposite; leaflets 4-8 

pairs per pinna, opposite, 1.5-3 × 1-1.7 cm, 

elliptical, ovate, or rounded, chartaceous, glabrous 

or puberulent, the apex rounded, mucronate, the 

base obtuse or cuneate, slightly unequal, the 

margins entire, revolute, ciliate; upper surface 

dull; lower surface with the midvein prominent; 

petiolules short, cylindrical, tomentose; petioles 

and rachis spiny; stipules lanceolate, acuminate, 

0.5-1 mm long. Flowers functionally unisexual, 

in axillary or terminal racemes; rachis tomentose 

and densely spiny; bracts lanceolate, 3.5-6 mm 

long, deciduous. Calyx campanulate, 4-5 mm 

long, tomentose, the sepals reflexed, oblong. 

Petals yellow, 4.5-6.5 mm long, spathulate; 

stamens 10, four of which are fertile in the 

staminate flowers, all sterile in the pistillate 

flowers; ovary sessile, rudimentary in the 

staminate flower, the stigma terminal. Legumes 

oblong to almost rounded, 5-7.5 × 3-5.5 cm, semiinflated, 

sparsely spiny, tardily dehiscent along 

both sutures. Seeds 1-2, ovoid or rounded, 1.5-2 

cm long, yellow to orange-brown or less frequently 

black.


Phenology: Flowering from March to August 

and fruiting in June. 



littoral zone. Also on Caja de Muerto, Cayo 

Ratones, Culebra, Icacos, Mona, Vieques, 


and Virgin Gorda; the Bahamas, Hispaniola, and 

throughout the Lesser Antilles. 


3. Caesalpinia culebrae (Britton & Wilson) 

Alain, Bull. Torrey Bot. Club 90: 187. 1963. 

Fig. 94. G 

BASIONYM: Guilandina culebrae Britton & Wilson 

Scandent shrub, with many lateral branches 

along the main stem, attainig 2 m in length. Stems 

cylindrical, unarmed, finely pubescent. Leaves 

bipinnate, paripinnate, ca. 22 × 14 cm; pinnae 7- 

8 pairs, opposite; leaflets 5-7 pairs per pinna, 

opposite, 1-1.5 × 0.9-1.4 cm, elliptical or ellipticobovate, 

chartaceous, the apex rounded, 

emarginate, the base unequal, one side rounded, 

the other obtuse, the margins entire, revolute, 

ciliate; upper surface glabrous, slightly punctate; 

lower surface puberulent, with the midvein 

prominent; petiolules short, tomentulose; petioles 

and rachis unarmed, tomentulose; stipules minute, 

deciduous. Flowers in axillary racemes, 6-10 cm 

long; rachis tomentulose; bracts lanceolate, with 

the apex caudate, ca. 4 mm long, deciduous. Calyx 

campanulate, ca. 8 mm long, tomentose, the sepals 

reflexed, oblong. Petals yellow, 7-8 mm long, 

spathulate. Legumes elliptic-circular, ca. 7 × 4 

cm, flattened, unarmed, apiculate at the apex. 

Seeds globose, ca. 1.7 cm in diameter, yelloworange. 


March. 

Status: Endemic, known from Culebra from 

very few collections. 

Distribution: Sandy dunes on Culebra and 

Cayo Diablo. 

4. Caesalpinia decapetala (Roth) Alst. in 

Trimen, Handb. Fl. Ceylon 6, Suppl. 89. 

1931. 

Fig. 95. A-C 

BASIONYM: Reichardia decapetala Roth 

SYNONYMS: Caesalpinia sepiaria Roxb. 

Biancaea sepiaria (Roxb.) Todaro 

Zarza de cercas, Zarza 

Vine or woody shrub, scandent, attainig 2-3 

m in length. Stems puberulent, striate, cylindrical, 

with numerous lenticels and recurved spines, 

attaining up to 2 cm in diameter. Leaves bipinnate, 

paripinnate, 30-45 cm long; pinnae 4-10 pairs, 

opposite; leaflets 7-12 pairs, opposite, 

chartaceous, oblong, 8-25 × 8-10 mm, the apex 

rounded or retuse, the base obtuse to rounded, the 

margins entire; upper surface dark green, dull; 

lower surface pale green, dull, punctate, with the 

midvein prominent, scattered-pubescent, 

especially along the midvein; petiolules minute; 

rachis usually with a reddish tinge, spiny, 

pubescent; petioles spiny, ca. 7 cm long, with the 

base broadened; stipules triangular, ca. 1 mm 

long, caducous. Flowers in subaxillary or terminal 

racemes, up to 35 cm long; rachis unarmed or 

with some minute spines at the base, pubescent, 

with stipitate glands; pedicels 2-2.5 cm long; 

bracts minute. Calyx green, the sepals oblong, ca. 

1 cm long, except for one that is almost twice the 

size of the other four. Petals yellow, almost 

rounded, 1-1.5 cm long; stamens exserted, 

filaments unequal, pubescent on the lower half; 

ovary pubescent. Legume woody, oblong, ca. 7 × 

2 cm, dehiscent along the upper suture. Seeds 

oblong, ca. 1 cm long, dark brown. 


September to June. 

Status: Exotic, naturalized, common. 

Distribution: On roadsides, along rivers, or 

in pastures, at lower and middle elevations. Native 

to India, but naturalized throughout the 

Neotropics and western Africa. 

Public Forests: Maricao, Río Abajo, and Toro 

Negro. 

5. Caesalpinia major (Medic.) Dandy & Exell, 

J. Bot. Brit. & For. 76: 180. 1938. 

Fig. 95. D-F 

BASIONYM: Bonduc majus Medic. 

SYNONYMS: Guilandina bonduc sensu Britton & Wilson, 

& Urb. 

Caesalpinia globerulum Bakh. f. & van Royen 

Liana or scandent shrub, which attains 15 m 

in length. Stems cylindrical, up to 2.5 cm in

236 



diameter, densely spiny. Leaves bipinnate, 

paripinnate, up to 60 cm long; pinnae 3-10 pairs, 

opposite; leaflets 4-8 pairs per pinna, opposite, 

3.5-7.7 × 2.5-3.7 cm, elliptical, ovate, or oblong, 

chartaceous, glabrous or puberulent, the apex 

obtuse or acute, mucronate, the base unequal, 

rounded, or one side obtuse, the margins entire, 

revolute, ciliate; upper surface glabrous, slightly 

shiny; lower surface with the midvein prominent, 

glabrous or puberulent; petiolules short, 

puberulent; petioles and rachis spiny, puberulent; 

stipules minute, deciduous. Flowers in axillary 

racemes, up to 30 cm long; rachis tomentulose, 

unarmed; bracts lanceolate, long-acuminate, ca. 

1 cm long, deciduous. Calyx campanulate, 6-7 

mm long, tomentose, the sepals reflexed, oblong. 

Petals yellow, 6-7 mm long, spathulate; stamens 

10, the filaments unequal, shorter than the petals; 

ovary pubescent, the style and stigma glabrous. 

Legumes broadly oblong, 6-12 × 3-6 cm, sparsely 

spiny. Seeds ovoid or rounded, 1.5-2.5 cm long, 

yellow-orange. 



Distribution: In the zone of mogotes. Also in 

Cuba, Jamaica, the Bahamas, and Hispaniola. 

Public Forest: Río Abajo. 

2. SENNA 

6. Caesalpinia portoricensis (Britton & 

Wilson) Alain, Bull. Torrey Bot. Club 90: 

187. 1973. 

Fig. 95. G-I 

BASIONYM: Guilandina portoricensis Britton & Wilson 

Scandent shrub attainig 6 m in length. Stems 

cylindrical, striate, puberulent, unarmed. Leaves 

bipinnate, paripinnate, ca. 15 cm long; pinnae 3- 

5 pairs, opposite; leaflets 3-5 pairs per pinna, 

opposite, 1-2 cm long, ovate or broadly elliptical, 

chartaceous, glabrous, the apex rounded or 

emarginate, the base unequal, rounded on one 

side, obtuse on the other, the margins entire; 

petioles and rachis puberulent, with some spines; 

stipules deciduous. Legumes broadly oblong, 5-6 

× cm, semi-woody, unarmed, dehiscent along one 

suture. Seeds ovoid or rounded, 1.5-2 cm long, 

dark brown to almost black, shiny, not striate. 

Phenology: Collected in fruit during March. 

Status: Endemic, very rare. 


from Salinas de Guánica. 

Public Forest: Guánica. 

Trees, erect, scandent, or clambering shrubs, or less frequently herbs. Stems unarmed. Leaves 

pinnate; leaflets opposite; petioles and rachis sulcate, usually with a stipitate gland; stipules persistent. 

Flowers bisexual, in axillary or terminal racemes or panicles or solitary; bracts minute to foliaceous; 

pedicels elongate or short. Calyx of 5 more or less elongate sepals; corolla yellow, of 5 free petals, 

unequal, unguiculate; stamens 10, three of which are usually smaller and sterile, the filaments flattened, 

free, the anthers basifixed, dehiscent by terminal pores; ovary unilocular, superior, short-stipitate, 

with numerous ovules. Fruit a legume of various forms, dehiscent or indehiscent; seeds numerous, of 

various forms. A genus of about 260 species, predominantly neotropical. 

Key to the species of Senna 

1a. Leaves with 2-5(6) pairs of sub-fleshy leaflets ……..…...……...….....….1. S. bicapsularis 

1b. Leaves with 2 pairs of chartaceous leaflets ……………………..……..………..2. S. nitida 

1. Senna bicapsularis (L.) Roxb., Fl. Ind. ed. 

2, 2: 342. 1832. 

Fig. 96. A-F 

BASIONYM: Cassia bicapsularis L. 

SYNONYM: Adipera bicapsularis (L.) Britton & Rose 

Hoja de sen, Sen del país, Stiver bush 

Clambering shrub, with many lateral 

branches, attainig 3 m in length. Stems slender, 

cylindrical, smooth, grayish or brown, with some

238 



lenticels. Leaves alternate, bipinnate, 3-8 cm long; 

leaflets 2-5 pairs, 1.5-4 × 1-2.5 cm, obovate to 

oblanceolate, sub-fleshy, glabrous, the apex 

truncate, emarginate and usually mucronate, the 

base asymmetrical, cuneate or subcordiform, the 

margins entire or slightly undulate; upper surface 

dark green, dull, sometimes glaucous; lower 

surface pale green, dull, with the midvein slightly 

prominent, glaucous; petiolules ca. 2 mm long; 

rachis sulcate, slender, with a stipitate, ellipsoid 

gland between the basal pair of leaflets; petioles 

short, glabrous or puberulent, with the base 

swollen; stipules minute, lanceolate, early 

deciduous. Inflorescences of axillary racemes, 

few-flowered; pedicels 1-3.5 cm long, articulate; 

bracts lanceolate, deciduous. Calyx forming a 

turbinate hypanthium, 1.5-3.4 mm long, the sepals 

5, oblong-elliptical, yellow, 5-8 mm long, 

concave, glabrous. Petals yellow, concave, 1-1.5 

cm long, cuneate at the base, the central, basal 

petal obovate, the lateral ones oblong, shorter than 

the central, basal petal; stamens 10, dimorphic, 3 

of which are longer and fertile, the remaining ones 

sterile; ovary glabrous, the style curved. Fruit an 

oblong, cylindrical legume, 8-19 × 1-2 cm, 

septate, indehiscent. Seeds numerous, 4-6 mm 

long, ellipsoid, brown, shiny. 


November to May. 


Distribution: In disturbed areas, usually on 

roadsides and in pastures, at lower elevations. Also 

on Culebra, Vieques, Anegada, St. Croix, St. 

Thomas, Tortola, and Virgin Gorda; the Antilles, 

tropical continental America, and Bermuda. Has 

been introduced into the Old World tropics. 

Public Forest: El Yunque. 

2. Senna nitida (L.C. Rich.) Irwin & Barneby, 

Mem. New York Bot. Gard. 35: 159. 1982. 

Fig. 96. G-J 

BASIONYM: Cassia nitida L.C. Rich. 

SYNONYMS: Chamaefistula antillana Britton & Rose 

Cassia antillana (Britton & Rose) Alain 

Hediondilla 

Clambering or scandent shrub, which attains 

2-5(10) m in length. Stems up to 2.5 cm in 

diameter, grayish, almost cylindrical; branches 

green, striate, glabrous or puberulent. Leaves 

alternate, pinnate; leaflets 4, opposite, ovate or 

elliptical, 6-12 × 2-4.5 cm, the distal pair larger, 

the apex acuminate, the base unequal or rounded, 

the margins slightly undulate, revolute; upper 

surface shiny, glabrous or puberulent, with the 

midvein sunken; lower surface glabrous, yellowish 

green, dull, punctate; petiolules swollen; rachis 

with a linear stipitate gland between the leaflets; 

petioles longer than the rachis, sulcate, swollen 

at the base. Inflorescences of axillary racemes, 

produced toward the end of the branches. Sepals 

oblong or ovate, ca. 5 mm long. Petals yellow, 

rounded, 1-2 cm long. Legume 10-18 × 1-1.5 cm, 

almost cylindrical, compressed along the suture, 

fleshy, producing a yellow exudate, oily and 

rancid. Seeds ca. 10 mm long, flattened, elliptical. 


December. 


Distribution: On roadsides and in forests and 

pastures, in moist areas, at middle and lower 

elevations. Also on St. Thomas and Tortola; St. 

Kitts and probably Haiti. 

Public Forests: Carite, El Yunque, Guajataca, 

and Maricao. 

Reference: Irwin, H. and R. Barneby. 1982. The American Cassiinae. A synoptical revision of 

Leguminosae tribe Cassieae subtribe Cassiinae in the New World. Mem. New York Bot. Gard. Vol. 35: 

1-918. 

24b. Subfamily FABOIDEAE 


1a. Leaves unifoliolate or pinnately compound. …………………………………………..……….2 

1b. Leaves trifoliolate………………………………………………………………………….……8

240 



2a. Leaves unifoliolate…………………………………………………………………..………....… 3 

2a. Leaves paripinnate or imparipinnate…………………………………………………..……….. 4 

3a. Leaflets cordiform, with a pair of stipels at the base; flower 3-5 cm long; calyx and corolla bright 

red; legume oblong, 6-8 cm long ……………………………...………. 15. Neorudolphia 

3b. Leaflets ovate to oblong, without stipels at the base; flower < 1 cm long; calyx green; corolla pink; 

legume almost circular, 2-2.3 cm long ……………………....7. Dalbergia ecastaphyllum 

4a. Leaves paripinnate; seeds subglobose, brilliant red with a black spot at the base …...… 1. Abrus 

4b. Leaves imparipinnate; seeds brown, black, or cream-colored…………………………….…… 5 

5a. Leaflets opposite, with stipels at the base; legume oblong……………………………………. 6 

5b. Leaflets alternate, without stipels at the base; corolla pink; legume circular or almost circular in 

outline …………………………………………………………………………………...........…… 7 

6a. Leaves with 5-7 leaflets; calyx with the dorsal lobes connate at the base; corolla blue-violet, the 

standard broadly ovate or circular, the wings as long as or longer than the keel; anthers yellow 

…………………………………………………..………………..…................ 6. Clitoria (in part) 

6b. Leaves with 11-21 leaflets; calyx with the dorsal lobes free to the base; corolla red-orange, the 

standard oblong, elongate, the wings shorter than the keel; anthers white ………. 2. Barbieria 

7a. Stipules spiny, persistent; fruit semicircular, woody ……………………….… 12. Machaerium 

7b. Stipules minute, deciduous, not spiny; fruit flattened, circular, chartaceous 

…………………………………………………………………………........ 7. Dalbergia monetaria 

8a. Legume obtusely 4-angular; seeds sticky …..………………………….….... 6. Clitoria falcata 

8b. Legume flattened or almost cylindrical, not quadrangular; seeds not sticky………………...… 9 

9a. Leaves lacking stipels at the base of the leaflets ……………………………..…… 4. Canavalia 

9b. Leaves with stipels at the base of the leaflets………………………………………………..... 10 

10a. Standard oblong, elongate; legumes with rigid, elongate, stinging hairs ………..... 14. Mucuna 

10b. Standard circular to reniform in outline; legumes lacking stinging hairs………………….... 11 

11a. Standard 3-4 cm wide, more than twice as long as the keel and the wings .…… 5. Centrosema 

11b. Standard < 2.5 cm wide, less than twice as long as the keel and the wings…………….…… 12 

12a. Legume cylindrical or almost cylindrical, sometimes compressed in the portion between the 

seeds………………………………………………………………………………………….......... 13 

12b. Legume not cylindrical, flattened or compressed laterally………………………………..… 15 

13a. Corolla maroon or purple ………………………………………………….... 13. Macroptilium 

13b. Corolla pink, lavender, or yellow…………………………………………………………… 14 

14a. Stipules auriculate; nodes of the rachis of the inflorescence swollen, with extrafloral nectaries; 

corolla yellow ……………………………………..……………......................… 21. Vigna luteola 

14b. Stipules truncate at the base; nodes of the rachis of the inflorescence not swollen, lacking extrafloral 

nectaries; corolla lavender or pink ..……..…………………………....................... 17. Phaseolus 

15a. Legumes conspicuously septate between the seeds……………………………………….…. 16 

15b. Legumes not septate between the seeds………………………………………………….….. 19

242 


16a. Legume with the apex straight ……..….………………………………………… 18. Pueraria 

16b. Legume with the apex recurved………………………………………………………..……. 17 

17a. Legumes ca. 3 mm wide; terminal leaflet elliptical or lanceolate ……………… 20. Teramnus 

17b. Legumes > 4mm wide; terminal leaflet rhombic or ovate……………………………..……. 18 

18a. Margin of the leaflets entire or sinuate ………………….…………….…… 3. Calopogonium 

18b. Margin of the leaflets serrate-mucronate ……………..……………………… 16. Pachyrhizus 

19a. Legumes ca. 5 cm wide, with 2 or 3 circular seeds, 3-4 cm in diameter ..…………. 9. Dioclea 

19b. Legumes


8-15 pairs, 0.8-1.5 × 0.3-0.7 cm, oblong or 

oblanceolate, membranaceous, the apex rounded 

and mucronate, the base rounded, the margins 

entire; upper surface glabrous, dull, with 

inconspicuous venation; lower surface puberulent, 

dull, with the midvein prominent; petiolules 

minute, slender; rachis without glands, 

puberulent, with a minute stipel at the base of 

each leaflet; petioles minute, slender, with the base 

slightly swollen; stipules filiform, 2-3 mm long, 

persistent. Inflorescences of small axillary or 

terminal pseudoracemes, with 5-7 flowers 

clustered on the swellings of the rachis; bracts 

minute, deciduous. Calyx campanulate, green, 3- 

5 mm long, puberulent; corolla pink, the standard 

ovate, with the center dark pink, up to 1 cm long, 

concave, the apex acute, the wings and keels as 

2. BARBIERIA 


1. Barbieria pinnata (Pers.) Baill., Hist. Pl. 2: 

263. 1870. 

Fig. 97. E-I 

BASIONYM: Galactia pinnata Pers. 

SYNONYMS: Clitoria pinnata (Pers.) R.H. Sm. & G.P. 

Lewis 

Clitoria polyphylla Poir. 

Barbieria polyphylla (Poir.) DC. 

Enredadera 

Woody vine or clambering shrub, twining, 

attainig 4-10 m in length. Stems slender, much 

branched, striate, hirsute, glabrescent, coppercolored, 

with some whitish lenticels. Leaves 

alternate, imparipinnate; leaflets 11-21, 

chartaceous, oblong, 2-5.5 × 1.2-2.2 cm, the apex 

rounded and mucronate, the base rounded or 

obtuse, the margins entire, ciliate; upper surface 

pale green, dull, with inconspicuous venation; 

lower surface green, silky or whitish, pubescent, 

with the midvein prominent, yellowish, covered 

with minute hairs; petiolules 1-2 mm long, 

ferruginous-pubescent; petioles 1-4 cm long, 

pilose; rachis pilose, with a pair of stipels at the 

base of each pair of leaflets; stipels linear, 3-5 

mm long; stipules lanceolate, 5-9 mm long, 

pubescent, persistent. Inflorescences of axillary 

racemes of few distal flowers, 4-16 cm long; bracts 

long as the standard, unguiculate. Legumes 

oblong, 2-4.5 × 1-1.5 cm, slightly inflated, with 

the apex elongate and recurved and the margins 

slightly undulate, opening along the two sutures. 

Seeds ellipsoid, brilliant red, with a black spot at 

the base, 6-8 mm long. 

Phenology: Flowering from October to June 

and fruiting from November to September. 

Status: Exotic, naturalized, rather common. 


secondary forests or along trails. Also on Culebra, 

Vieques, and the Virgin Islands. Native to the 

paleotropics, but found naturalized throughout the 

Neotropics. 

Public Forests: Cambalache, Ceiba, Guánica, 

Piñones, and Susúa. 

lanceolate, pubescent, forming an involucre at the 

base of the calyx. Calyx tubular, reddish, striate, 

2.5-3.5 cm long, with five subulate sepals, 

acuminate, 0.8-1 cm long, the ventral sepals 

slightly longer than the lateral ones; corolla red 

or red-orange, the standard oblong-oblanceolate, 

unguiculate, 5-6 cm long, the wings oblong, 9- 

13 mm long, the keel elliptic-oblong, 17-23 mm 

long; stamens 10, staminal column white, the 

anthers white; ovary superior, linear, 8-11 mm 

long, white-pubescent. Legume subsessile, 

oblong, flattened, laterally compressed between 

the seeds, 5-7 × 0.5-0.6 cm, pubescent, with the 

margins sinuate and the calyx persistent at the 

base. Seeds 4-9 per fruit, dark brown or black, 

oblong, 4-6 mm long. 


November to January and in fruit from January 

to May. 

Status: Native, not very common. 

Distribution: In forests in the zone of mogotes, 

in secondary forests, and on roadsides in moist 

places at middle elevations. Also in Cuba, 

Hispaniola, Central America, and South America. 

Public Forests: El Yunque, Maricao, and Río 

Abajo.

244 



3. CALOPOGONIUM 

Slightly woody vines, twining. Leaves alternate, trifoliolate; stipels and stipules minute. 

Inflorescences of axillary pseudoracemes or fascicles; the flowers clustered on the nodal swellings of 

the rachis; bracts and bracteoles minute; pedicels short. Calyx campanulate-tubular, with the two upper 

lobes united to form a lip, the 3 lower lobes lanceolate; corolla blue or violet, the standard obovate, 

auriculate at the base, the wings narrow, the keel shorter than the wings; stamens 9-10, one of them 

free, the rest with the filaments united into a long tube; ovary superior, sessile, hirsute, with numerous 

ovules, the style curved, glabrous, the stigma capitate. Fruit a flattened legume, linear, dehiscent, 

septate between the seeds; seeds circular. A genus of 6-8 species, native to tropical continental America, 

naturalized in the paleotropics. 

Key to the species of Calopogonium 

1a. Terminal leaflet rhombic or rounded-rhombic; stems villous, glabrescent when mature; bracteoles 

ca. 2 mm long, deciduous; legumes 7-10 mm wide ................................…………..1. C. coeruleum 

1b. Terminal leaflet ovate or rhombic-ovate; stems hirsute; bracteoles 4-6 mm long, persistent; legumes 

3-5 mm wide ...........................................…………………………..………….... 2. C. mucunoides 

1. Calopogonium coeruleum (Benth.) Sauv., 

Anal. Acad. Ci. Habana 5: 337. 1869. 

Fig. 98. A-F 

BASIONYM: Stenolobium coeruleum Benth. 

Slightly woody vine, twining, with many 

lateral branches, attainig 10 m in length. Stems 

slender, cylindrical, villous-pubescent on the 

younger portions. Leaves alternate, trifoliolate; 

leaflets chartaceous, 5-10.5 × 3-8 cm, the apex 

obtuse or less frequently rounded, the base of the 

distal leaflet cuneate, that of the lateral ones 

asymmetrical and obtuse-truncate, the margins 

undulate, revolute; upper surface dark green, dull, 

pubescent, especially on the sunken venation; 

lower surface pale green, pubescent, with the 

venation yellowish, prominent. Inflorescences of 

axillary pseudoracemes, 12-45 cm long, with 4-5 

flowers grouped on small mounds along the 

rachis; rachis thick, cylindrical, pubescent; 

pedicels ca. 5 mm long, pubescent. Calyx 

campanulate, 5-7 mm long, appressed-pubescent, 

the sepals lanceolate, 2.5-4 mm long; corolla blue, 

the standard ca. 1 cm long, oblanceolate. Legume 

tomentose, 3-8 × 0.7-1.0 cm, flattened, with the 

margins compressed between the seeds and the 

calyx persistent at the base. Seeds almost square, 

reddish brown, shiny, 4-5 mm wide. 


February and fruiting from February to April. 

Status: Exotic, naturalized, common. 

Distribution: In moist pastures and disturbed 

areas, at lower and middle elevations. Also in 

Cuba, Hispaniola, St. Vincent, Trinidad and 

Tobago, and tropical continental America. 

2. Calopogonium mucunoides Desv., Ann. Sci. 

Nat. (Paris) Sér. 1, 9: 423. 1826. 

Fig. 98. G-I 

SYNONYM: Calopogonium orthocarpum Urb. 

Jícama 

Slightly woody vine, twining or creeping, 

much branched, attainig 3-5 m in length. Stems 

slender, cylindrical, hirsute. Leaves alternate, 

trifoliolate; leaflets chartaceous, 2-10 × 1.5-6.5 

cm, the apex obtuse to almost rounded, the base 

of the terminal leaflet cuneate to rounded, and 

that of the lateral leaflets markedly asymmetrical 

(obtuse-subtruncate), the margins entire; both 

surfaces dull, appressed-pubescent; lower surface 

with prominent venation. Inflorescences of 

axillary pseudoracemes, up to 10 cm long; rachis 

hirsute; bracts subulate, persistent, ca. 7 mm long. 

Calyx campanulate, 7-8 mm long, hirsute, the 

sepals subulate, caudate at the apex; corolla blue 

or purple, the standard 6-7 mm long, emarginate. 

Legume oblong-linear, flattened, hirsute, 2-4 × 

0.3-0.5 cm. Seeds almost quadrangular, ca. 3 mm 

wide, reddish brown, shiny.

246 




December to March. 

Status: Exotic, naturalized, relatively 

common. 


pastures and roadsides, in moist zones at lower 

4. CANAVALIA 

and middle elevations. Also in Cuba, Hispaniola, 

Jamaica, tropical America, and naturalized in 

Africa and Asia. 


Woody or herbaceous vines, twining or creeping. Leaves alternate, trifoliolate; stipels absent; 

stipules minute, deciduous. Inflorescences of axillary or terminal pseudoracemes; the flowers clustered 

on the nodal swellings of the rachis; bracts minute, in pairs; pedicels short. Calyx campanulate, the 

lobes 5, unequal, 2 larger; corolla pink, violet, or purple, the standard obovate, unguiculate, auriculate 

and thickened at the base, the wings and the keel of similar size; stamens 10, the filaments united into 

a long tube; ovary superior, sessile or stipitate, pubescent, with numerous ovules, the style filiform, the 

stigma capitate. Fruit an elongate legume, flattened or slightly inflated, dehiscent or indehiscent, not 

septate between the seeds, with a longitudinal keel along both sutures or along the ventral suture; seeds 

3 or more, usually oblong. A genus of 50 species, the great majority native to the New World. 

Key to the species of Canavalia 

1a. Seeds brown or white; legumes with a keel on each side of both sutures (each valve with three 

longitudinal ribs); flowers in groups of 2-3 on the nodal swellings along the rachis 

............................................................................................................................................. 2 

1b. Seeds red; legumes with a rib on each side of the ventral suture; flowers in groups of 3-5 on the 

nodal swellings along the rachis ..........................…………………………………..….…2. C. nitida 

2a. Plants climbing, cultivated or in areas of secondary vegetation; legumes up to 30 cm long; seeds 

white ……………………………….........................................……...1. C. ensiformis 

2b. Plants creeping or climbing, common on the sandy coasts of the littoral zone; legumes 7-15 cm 

long; seeds brown .........................………………………………………………….… 3. C. rosea 

1. Canavalia ensiformis (L.) DC., Prodr. 2: 

404. 1825. 

Fig. 99. A-E 

BASIONYM: Dolichos ensiformis L. 

Slightly woody vine, twining, attainig 1-2 m 

in length. Stems cylindrical, puberulent, 

glabrescent, with the pith hollow. Leaves 

alternate, trifoliolate; leaflets chartaceous, broadly 

ovate or broadly elliptical, 6-20 × 5-12 cm, the 

apex obtuse or rounded, short-apiculate, the base 

obtuse, asymmetrical on the lateral leaflets, the 

margins entire; both surfaces strigulose, punctate; 

petiole and rachis cylindrical, glabrescent; 

petiolules swollen, ca. 8 mm long. Inflorescences 

of axillary pseudoracemes, 25-36 cm long, with 

2-3 flowers grouped on the nodal swellings along 

the rachis; pedicels 1-2 mm long, glabrescent. 

Calyx green, campanulate, 12-14 mm long, 

bilabiate; corolla pale violet, ca. 2 cm, the standard 

and the wings white at the base, violet on the distal 

portion. Legume linear, up to 30 × 3.5 cm, woody, 

slightly curved toward the apex, each valve with 

three longitudinal ribs. Seeds 15-20, ellipsoid, up 

to 3 cm long, white or cream-colored. 


Status: Exotic, known only in cultivation, not 

very common. 

Distribution: Native to the Neotropics, but 

known only in cultivation. This species was 

described based on material collected by Sloane 

in Jamaica.

248 


2. Canavalia nitida (Cav.) Piper, Contr. U. S. 

Natl. Herb. 20: 562. 1925. 

Fig. 99. F-G 

BASIONYM: Clementea nitida Cav. 

SYNONYM: Canavalia rusiosperma Urb. 

Mato colorado, Haba de burro de jardín 

Woody vine, twining, with numerous 

pendulous branches, attainig 15-18 m in length. 

Stems cylindrical, glabrous or puberulent, thick, 

the bark light brown, almost smooth. Leaves 

alternate, trifoliolate; leaflets coriaceous, 

elliptical, oval, or oblong, involute, 6.5-12 × 3- 

5.5 cm, the apex obtuse, acute, or rounded, the 

base rounded, asymmetrical on the lateral leaflets, 

the margins slightly revolute, thickened, both 

surfaces glabrous, with the venation slightly 

prominent; petiolules cylindrical, pubescent, 4-5 

mm long; petioles 4-5 cm long, with the base 

swollen; stipels absent; stipules minute, 

deciduous. Inflorescences of axillary 

pseudoracemes, 13-23 cm long, the flowers in 

groups of three or four on each nodal swelling 

along the rachis; pedicels 5-6 mm long, 

puberulent. Calyx greenish, shiny, 15-18 mm 

long, pubescent; corolla violet-pink, the standard 

orbicular, ca. 2.5 cm long, with the center pale 

yellow, the keel and the wings 

ca. 1.5 cm long. Legume woody, brown, up to 

20 × 5 cm, the valves with two slight ribs on the 

sides of the suture. Seeds ca. 2 cm long, almost 

circular or ellipsoid, crimson with the hilum 

black. 


August to February. 


Distribution: In forests in moist areas, at 

middle and lower elevations in central, northern, 

and western Puerto Rico. Also on St. John and St. 

Thomas; Hispaniola. 

3. Canavalia rosea (Sw.) DC., Prodr. 2: 404. 

1825. 

5. CENTROSEMA 

Fig. 99. H-L 

BASIONYM: Dolichos roseus Sw. 

SYNONYMS: Dolichos maritimus Aubl. 

Canavalia maritima (Aubl.) Urb. 

Canavalia maritima Thouars 

Dolichos obtusifolius Lam. 

Canavalia obtusifolia (Lam.) DC. 

Haba de playa, Habichuela playera, Mato de 

playa, Bay bean, Canavalia 

Woody vine, creeping or twining, much 

branched from the base, attainig 2-6 m in length. 

Stems smooth, glabrescent, green, mulberrycolored 

at the nodes, with adventitious roots. 

Leaves alternate, trifoliolate; leaflets 4-10.4 × 3- 

10 cm, chartaceous, oblong, ovate, or orbicular, 

involute, the apex obtuse, rounded, retuse and 

mucronate, the base cuneate or rounded, the 

margins entire; upper surface glabrous except for 

the puberulent veins; lower surface puberulent, 

yellowish green, dull, with prominent venation; 

rachis sulcate, villous, 3-4.5 cm long; petiolules 

swollen, villous, ca. 1 cm long; petioles sulcate, 

villous, swollen at the base, 4-6 cm long; stipels 

absent; stipules triangular, persistent. 

Inflorescences of axillary pseudoracemes, 25-30 

cm long, the flowers in pairs on each nodal 

swelling along the rachis; bracts minute, 

deciduous. Calyx 8-11 mm long, yellowish green, 

with five lobes, three small and two large, 

puberulent; corolla violet-pink, the standard 

broadly elliptical, 2-2.5 cm long, reflexed, white 

in the center, the wings and the keel shorter than 

the standard. Legume 10-17 × 2.3-2.5 cm, oblong, 

not flattened, subwoody, with three keels on each 

valve. Seeds numerous, 1.5-1.8 cm long, ellipsoid, 

brown, with the hilum white. 





littoral zone. On all the islands and keys of Puerto 

Rico and the Virgin Islands; throughout the 

tropical and subtropical coasts of the planet. 


Herbaceous vines, twining. Leaves alternate, trifoliolate; stipels present; stipules minute, persistent. 

Flowers axillary, solitary or grouped on a bracteate peduncle; bracts appressed to the calyx; pedicels 

more or less elongate. Calyx campanulate, the lobes 5, unequal or almost equal; corolla pink, lavender,


250 


or white, the standard ovate or rounded, unguiculate, the wings and the keel of similar size, shorter 

than the standard; stamens 10, diadelphous or monadelphous; ovary superior, almost sessile, with 

numerous ovules, the style curved, pubescent, the stigma capitate or truncate. Fruit a linear legume, 

flattened, with the margins ribbed, dehiscent by valves that twist on drying, not septate between the 

seeds; seeds numerous, small, oblong. A genus of about 45 species, of pantropical distribution. 

Key to the species of Centrosema 

1a. Corolla pink-burgundy; standard 4.5-6 cm wide; legume 7-10 mm wide ….1. C. plumieri 

1b. Corolla pale violet; standard 2-4.5 cm wide; legume 4-6 mm wide……………….…….. 2 

2a. Lobes of the calyx very unequal, the lateral ones deltate, as long as the tube of the calyx, the central 

ones subulate, much longer than the tube; legume ca. 6 mm wide................................. 2. C. pubescens 

2b. Lobes of the calyx subequal, subulate, longer than the tube of the calyx; legume ca. 4 mm wide 

...................………………………………………………..………………….. 3. C. virginianum 

1. Centrosema plumieri (Turp. ex Pers.) Benth., 

Comm. Leg. Gen. 54. 1837. 

Fig. 100. A-F 

BASIONYM: Clitoria plumieri Turp. ex Pers. 

SYNONYM: Bradburva plumieri (Turp. ex Pers.) Kuntze 

Flor de conchitas 

Non-woody vine, twining, attainig 5 m in 

length. Stems cylindrical, green, smooth, glabrous 

or puberulent. Leaves alternate, trifoliolate; 

leaflets 4-11 × 3-8 cm, the apex acute, obtuse, or 

short-acuminate; margins undulate; upper surface 

slightly shiny, with the venation sunken, 

puberulent; lower surface dull, puberulent, with 

prominent venation; terminal leaflet deltate, 

ovate, or elliptical, with the base cuneate or 

cuneate-rounded; lateral leaflets oblong or 

elliptical, with the base asymmetrical, roundedtruncate; 

petiolules swollen, 3-4 mm long; petioles 

6-10 cm long, slender, glabrous, with the base 

swollen; stipels linear, up to 10 mm long; stipules 

oblong-lanceolate, 10-12 mm long, with the 

venation conspicuously parallel. Flowers solitary 

or in pairs at the end of long axillary peduncles; 

bracteoles ovate, convex, 10-14 mm long. Calyx 

campanulate, 5-8 mm long, glabrous, with three 

lobes, ca. 2.5 mm long; corolla pink to cardinal 

red, abaxially appressed-pubescent, the standard 

4.5-6 cm wide, rounded, cardinal red in the center 

and pink or whitish toward the edges, the wings 

ca. 3 × 1.5 cm, with the apex cardinal red, the 

keel white, ca. 3.5 cm long. Legume flattened, 

linear, 10-15 × 0.6-1 cm, the apex longacuminate, 

the margins thickened. 


January and fruiting from January to February. 

Status: Apparently native, relatively common. 


roadsides and pastures, at lower elevations in 

moist areas. Throughout the Neotropics, 

introduced in Africa and tropical Asia. 

Public Forests: Guajataca and Río Abajo. 

2. Centrosema pubescens Benth., Ann. Wiener 

Mus. Naturgesh. 2: 119. 1837. 

Fig. 100. G-I 

SYNONYM: Bradburya pubescens (Benth.) Kuntze 

Flor de pito, Flor de conchitas 

Slightly woody vine, twining, attainig 8 m in 

length. Stems slender, appressed-pubescent. 

Leaves alternate, trifoliolate; leaflets ovate, 

oblong, or elliptical, 3-8 × 1.6-4.5 cm, the apex 

acute or obtuse, the base rounded or obtuse, the 

margins entire; both surfaces glabrous to 

pubescent; upper surface with the venation 

sunken; lower surface with prominent venation; 

petiolules thickened, pubescent; petioles sulcate, 

almost quadrangular, 2-5 cm long, puberulent; 

stipels subulate, 2-2.5 mm long; stipules ovate, 

2-4 mm long. 

Flores solitary or in pairs at the end of 

bracteate, axillary peduncles, 5-10 cm long; bracts 

oblong, slightly longer than the calyx. Calyx 

asymmetrically campanulate, the tubular portion 

5-7 mm long, with five very unequal lobes, the 

lateral ones deltate, ca. 1 mm long, the central


ones subulate or linear, ca. 6 mm long; corolla 

pale violet, lavender, or less frequently yellowish 

or whitish with purple lines, the standard 2-4.5 

cm wide, abaxially velvety, the wings and the keel 

shorter than the standard. Legume linear, 10-20 

× 0.6 cm, the apex acuminate, with the margins 

ribbed, dehiscent by valves that twist on drying. 

Seeds 8-15 per fruit, oblong-quadrangular, dark 

brown, dull, 5-6 mm long. 

Phenology: Flowering predominantly during 

the months from November to December, but may 

be found in flower from September to March. 



abandoned fields, pastures, or roadsides, at middle 

and lower elevations. Native to tropical America, 

introduced in Asia and Africa. 

Public Forests: Maricao, Río Abajo, and 

Susúa. 

3. Centrosema virginianum (L.) Benth., Comm. 

Legum. Gen. 56. 1837. 

Fig. 100. J-M 

BASIONYM: Clitoria virginiana L. 

SYNONYM: Bradburya virginiana (L.) Kuntze 

Conchita de Virginia, Flor de conchitas, Blue 

wiss, Butterfly pea 


in length. Stems slender, strong, cylindrical, finely 

pubescent. Leaves alternate, trifoliolate; leaflets 

2.5-7 × 0.5-2 cm (the terminal one longer than 

the lateral ones), lanceolate, oblong, ovate, or 

linear, chartaceous, the apex acute or obtuse, 

mucronate, the base cuneate to rounded, the 

6. CLITORIA 

margins entire or crenate, ciliate, both surfaces 

glabrous or puberulent; lower surface with 

prominent venation; rachis 0.7-1.5 cm long; 

petioles 2.3-2.8 cm long, slender, sulcate, 

thickened; stipels narrowly subulate, 4-5 mm 

long; stipules subulate, 3-5 mm long, persistent, 

markedly veined. Flowers 1-4, at the end of a 

bracteate axillary peduncle; bracts ovate or ovatelanceolate, 

persistent, notably veined. Calyx 

green, asymmetrically campanulate, with five 

subulate, subequal lobes, the tubular portion ca. 

2 mm long, the lateral lobes ca. 5 mm long, the 

central lobes 7-8 mm long; corolla lilac or almost 

white, the standard broadly ovate, 3-4 cm wide, 

white in the center with purple lines, the wings 

and keel unguiculate, much shorter than the 

standard. Legume linear, 10-13 × 0.3-0.4 cm, the 

apex acuminate, the margins ribbed, dehiscent by 

valves that twist on drying. Seeds numerous, 3-4 

mm long, oblong, brown or light brown with dark 

brown spots. 





roadsides, fences, and pastures, in thickets and 

dry forests, usually along the coasts, in the zone 

of mogotes, and in areas of the Cordillera Central. 

Also on Culebra, Culebrita, Icacos, Mona, 

Vieques, and the Virgin Islands; throughout the 

Antilles and from southern North America to 

Argentina. Introduced in Africa. 

Public Forests: Cambalache, Ceiba, El 

Yunque, Guánica, Maricao, Mona, Piñones, Río 

Abajo, Susúa, Toro Negro, and Tortuguero. 

Twining herbaceous or woody vines, trees, or shrubs. Leaves alternate, trifoliolate or imparipinnate; 

leaflets 3-9, opposite; stipels present; stipules minute, persistent. Flowers solitary or grouped in axillary 

racemes; bracteoles appressed to the calyx, persistent. Calyx campanulate, with 5 equal or almost 

equal lobes; corolla blue-violet, white, yellow, or red, the standard rounded, rugose, longer than the 

wings and the keel, the wings longer than the keel; stamens 10, diadelphous or monadelphous; ovary 

superior, stipitate, the style curved, pubescent, the stigma truncate. Fruit a linear or oblong legume, 

flattened, dehiscent along both sutures, not septate between the seeds; seeds few, rounded to oblong. A 

genus of about 60 species, the great majority of tropical to subtropical distribution.

252 



Key to the species of Clitoria 

1a. Leaves 3-foliolate; corolla light yellow, white, or white with lilac lines; legumes obtusely 

quadrangular; seeds sticky ……………...…………………………………….. 1. C. falcata 

1b. Leaves 5-7-foliolate; corolla blue-violet; legumes flattened; seeds not sticky 

………………………………………………………………………………...2. C. ternatea 

1. Clitoria falcata Lam., Encycl. Méth. Bot. 2: 

51. 1786. 

Fig. 101. A-C 

SYNONYMS: Clitoria rubiginosa Juss. 

Martiusia rubiginosa (Juss.) Britton 

Ahoga gallina, Conchitas, Flor de pito 


length. Stems slender, flexible, tomentose, 

glabrescent when mature. Leaves alternate, 

trifoliolate; leaflets coriaceous, 2.2-9 × 1.2-4.5 cm, 

elliptical, ovate, or lanceolate, the apex acute or 

obtuse, mucronate, sometimes retuse, the base 

obtuse or rounded, the margins smooth, revolute; 

upper surface dark green, dull, with the venation 

pale green, glabrous; lower surface sericeous, with 

prominent venation; stipels lanceolate, ca. 7 mm 

long, pubescent; petiolules minute, ferruginoustomentose, 

thickened; rachis 0.5-1.5 cm long, 

ferruginous-tomentose; petioles ferruginoustomentose, 

4-6 cm long, with the base thickened; 

stipules elliptical or ovate, ca. 7 mm long, with 

notable parallel veins. Inflorescences of axillary 

cymes with 2-4 flowers at the end of a peduncle, 

tomentose, 2-11 cm long; bracts hirsutulous, ca. 

8 mm long, oblong, green, with notable parallel 

veins; pedicels ca. 4 mm long, tomentose. Calyx 

campanulate, yellowish green, the tube 12-16 mm 

long, the lobes deltate, 7-12 mm long; corolla pale 

yellow or white with pale violet lines, the standard 

rounded, ca. 5 cm long, plicate, retuse, with violet 

or cardinal red lines on the central portion inside, 

the wings and keels ca. 2 cm long, longer than 

wide. Legume 4-6 cm long, pubescent, oblong, 

obtusely quadrangular, with a rib along each of 

the valves, the apex acuminate. The seeds 4-6 per 

fruit, almost spherical, ca. 3 mm in diameter, 

covered with a sticky, gelatinous matrix. 


September to January. 



pastures, fences, and along trails, at middle and 

lower elevations. Native to tropical America; 

introduced in western Africa. 

Public Forests: El Yunque and Río Abajo. 

Commentary: Fantz (1990) recognizes two 

varieties belonging to this species, of which only 

the typical variety is found in Puerto Rico. Within 

this variety he recognizes two forms based on 

characters of the fruit, which are found in Puerto 

Rico. Clitoria falcata var. falcata f. falcata 

possesses legumes with ribs that extend along the 

length of the valves, while in C. falcata var. 

falcata f. heteromorpha (Griseb.) Fantz the ribs 

are absent or extend along at most 2/3 of the length 

of the valve. 

2. Clitoria ternatea L., Sp. Pl. 753. 1753. 

Fig. 101. D-H 

Bejuco de conchitas, Papitos, Deleite, 

Conchitas, Blue vine, Butterfly pea 

Woody vine, twining, attainig 1-3 m in length. 

Stems slender, cylindrical, with lines of minute 

trichomes. Leaves alternate, 5-7-foliolate; leaflets 

opposite, 1.5-4.5 × 1-3.5 cm, elliptical or less 

frequently ovate or oblong, chartaceous, the apex 

rounded, obtuse, sometimes retuse or mucronate, 

the base obtuse, the margins entire; upper surface 

dark green, dull, puberulent, with the midvein 

sunken; lower surface pale green, dull, puberulent, 

with prominent venation; rachis 2-7 cm long; 

petiolules minute, pubescent; petioles 2-4 cm long, 

pubescent, with the base thickened; stipels 

filiform, ca. 1.5 mm long; stipules lanceolate, 

pubescent, ca. 4 mm long. Flowers solitary, on 

short peduncles; pedicel ca. 1 cm long, pubescent, 

with a pair of bracteoles in the middle. Calyx 

campanulate, 1.5-2.2 cm long, green, puberulent, 

the lobes lanceolate-ovate, 8-10 mm long, with 

the midvein conspicuous; corolla blue-violet, the 

standard broadly ovate, 3.5-5 cm long, retuse, with 

the base pale yellow and the center pale yellow 

inside. Legume 9-11 × ca. 1 cm, oblong, ribbed

254 



along both margins, the apex acuminate. Seeds 

numerous, 5-6 mm long, oblong, flattened, dark 

brown. 




relatively common. 


pastures or roadsides. Native to Africa but found 

widely distributed throughout the tropics and 

7. DALBERGIA 

subtropics of the New World. Also on Vieques, 

St. Croix, St. John, St. Thomas, and Tortola. 

Public Forests: Guánica and Río Abajo. 

Commentary: According to Fantz (1990), the 

following two varieties are found in Puerto Rico: 

Clitoria ternatea var. ternatea with simple flowers 

(only one standard per flower) and C. ternatea 

var. pleniflora Fantz with double flowers (4-5 

standards per flower). 

Trees, clambering shrubs, or lianas. Leaves alternate, imparipinnate or unifoliolate; leaflets alternate; 

stipels absent; stipules minute, deciduous. Inflorescences of axillary or terminal racemes or panicles; 

bracts and bracteoles minute, deciduous or persistent. Calyx campanulate, with 5 elongate lobes, equal 

or unequal; corolla white, yellow, or pink, the standard rounded or ovate, retuse, narrow at the base, 

the wings usually longer than the keel; stamens 9-10, diadelphous or monadelphous; ovary superior, 

stipitate, pubescent, the style usually curved, the stigma minute. Fruit a linear to rounded legume, 

indehiscent, usually with the margins membranaceous; seeds small, lenticular. A tropical genus of 


Key to the species of Dalbergia 

1a. Leaves unifoliolate; calyx ferruginous-tomentose; plants usually of the littoral zone 

...............………………………………………………………...……………… 1. D. ecastaphyllum 

1b. Leaves 3-5-foliolate; calyx puberulent; plants usually along rivers in the interior of the island 

.................………………….………………………………………………...…2. D. monetaria 

1. Dalbergia ecastaphyllum (L.) Taub in Engl. 

& Prantl, Nat. Pflanzenfam. 3(3): 335. 

1894. 

Fig. 102. A-E 

BASIONYM: Hedysarum ecastaphyllum L. 

SYNONYMS: Ecastaphyllum ecastaphyllum (L.) Britton 

Ecastaphyllum brownei Pers. 

Clambering or scandent shrub, much branched 

from the base, attainig 1-5 m in length. Stems 

cylindrical, white-pubescent. Leaves alternate, 

unifoliolate (exceptionally with 3-foliolate leaves 

on the same plant); leaflets 6-12 × 4.5-8 cm, ovate 

or oblong, coriaceous, the apex obtuse to 

acuminate, the base rounded to almost cordiform, 

the margins entire; upper surface dark green, dull, 

glabrous, with the venation sunken; lower surface 

light green, puberulent, with prominent venation; 

petioles 5-10 mm long, pubescent; stipels absent; 

stipules ovate to lanceolate, 4-10 mm long, white- 

sericeous, deciduous. Inflorescences of axillary 

panicles 1-4 cm long, densely ferruginoustomentose; 

bracts minute, deciduous. Calyx 

campanulate, 2.5-3.5 mm long, green, 

ferruginous-tomentose; corolla pink or white, the 

standard rounded, ca. 7 mm long, retuse, 

unguiculate, the wings longer than the standard. 

Legume 2-2.3 × 1.5-2 cm, oblong to almost 

circular, flattened, indehiscent, brown when ripe. 

Seed solitary, up to 1.7 cm long. 





littoral zone. Also on Mona, Vieques, St. Croix, 


throughout tropical America and Africa. 

Public Forests: Guánica, Mona, Piñones, and 

Tortuguero.

256 


2. Dalbergia monetaria L.f., Suppl. Pl. Syst. 

Veg. 317. 1781. 

Fig. 102. F-I 

SYNONYM: Securidaca volubilis L. pro parte 

Membrillo, Palo de brasilete 

Liana with pendulous branches, which 

supports itself on other plants by means of short 

branches that entangle like tendrils, attaining 10 

m or more in length. Stems flexible, cylindrical, 

up to 10 cm in diameter; bark light brown; 

branches cylindrical, pubescent on the younger 

portions. Leaves alternate, 3- to 5-foliolate; 

leaflets alternate, 5-12 × 3-7 cm, ovate or 

elliptical, involute, the apex acuminate, the base 

obtuse or rounded, the margins entire or slightly 

undulate, revolute; upper surface dark green, 

shiny, glabrous; lower surface pale green, dull, 

puberulent, with prominent venation; petiolules 

1. Desmodium axillare (Sw.) DC., Prodr. 2: 

333. 1825. 

Fig. 103. A-C 

BASIONYM: Hedysarum axillare Sw. 

SYNONYMS: Meibomia axillaris (Sw.) Kuntze 

Meibomia umbrosa Britton 

Meibomia sintenisii (Urb.) Britton 

8. DESMODIUM 

thickened, puberulent; rachis puberulent, up to 

11 cm long; petioles 1.2-5 cm long, puberulent; 

stipules minute, ovate, caducous. Inflorescences 

of short, clustered, axillary racemes. Calyx 

campanulate, green, 2-3 mm long, puberulent; 

corolla white, the standard rounded, 5-10 mm 

long, retuse, unguiculate at the base; the wings 

and the keel as long as the standard. Legume 

circular, flattened, 3-4 cm long. Seed one. 


October and fruiting from June to March. 


Distribution: Along the banks of rivers and 

streams, in the interior of the island, at middle 

and lower elevations. Also throughout the Antilles 

and in tropical South America. 

Public Forests: El Yunque, Guajataca, 

Maricao, Río Abajo, and Susúa. 

Erect, prostrate, or clambering herbs, perennial. Leaves alternate, usually trifoliolate; stipels minute; 

stipules minute, deciduous or persistent. Inflorescences of axillary or terminal pseudoracemes or panicles; 

bracts and bracteoles minute, deciduous or persistent. Calyx campanulate, with 5 short, almost equal 

lobes; corolla yellow, pink, or bluish, the standard oblong to rounded, retuse, narrow at the base, the 

wings and the keel of the same length; stamens 10, diadelphous or monadelphous; ovary superior, 

stipitate or sessile, pubescent, with few ovules, the style inflexed, the stigma minute. Fruit a linear 

legume, flattened or spiral, with the ventral margin or both margins deeply sinuate between the seeds, 

indehiscent, but separating in segments containing a single seed which adheres to the clothing or the 

fur of animals; seeds small, oblong. A genus of about 300 species of almost cosmopolitan distribution. 

Key to the species of Desmodium 

1a. Stems cylindrical; legume crenate only along the ventral margin ……..………………… 2 

1b. Stems trigonal; legume crenate along both margins ..…………………….... 3. D. intortum 

2a. Legume with 2-3 segments; leaflets with the apex acute or acuminate or less frequently obtuse, 

distal leaflet cuneate at the base ..........………………..………………….....1. D. axillare 

2b. Legume with 5-8 segments; leaflets with the apex obtuse or acute, distal leaflet obtuse or rounded 

at the base ...............……………………..………..……..……………………2. D. incanum 

Decumbent or scandent herb, scarcely 

branched from the woody base, attainig 2 m in 

length. Stems more or less cylindrical, producing 

aerial roots in the area of the nodes, pubescence 

mixed, of uncinate trichomes and minute, erect 

trichomes, glabrescent with age. Leaves alternate, 

trifoliolate; leaflets 3-9 × 2-6.5 cm, chartaceous, 

the apex obtuse, acute, or acuminate, usually


mucronate, the margins entire, strigose; upper 

surface dark green, glabrous, puberulent on the 

venation; lower surface grayish green, strigose, 

with prominent venation; distal leaflet rhombic, 

with the base cuneate; lateral leaflets 

asymmetrical, one side elliptical, the other ovate,

258 


the base obtuse or rounded; petiolules 2-5 mm 

long, pilose; petioles 4.5-10.5 cm long, uncinatepubescent, 

striate; rachis 1-6 mm long, pilose, 

canaliculate; stipules ovate to lanceolate, 5-10 mm 

long, deciduous; stipels filiform, ca. 2 mm long. 

Inflorescences of axillary racemes, erect, 30-45 

cm long, the axis pubescent, dark brown, with 

flowers paired along its length; pedicels 12-17 

mm long. Calyx purple, campanulate, ca. 2 mm 

long, pilose; corolla pink or pale violet, ca. 4 mm 

long. Legume uncinate-pubescent, 0.7-1(2) cm 

long, crenate on the ventral suture, with (1)2(3) 

fertile segments. Seeds ca. 3.5 mm long, oblongelliptical, 

light brown. 


January to August. 


Distribution: On the ground in dry to moist 

forests and in areas of disturbed vegetation along 

the Cordillera Central. Also on St. Croix; 

distributed throughout the Antilles, Central and 


Public Forests: El Yunque, Guilarte, Maricao, 


Key to the varieties in Puerto Rico 

1a. Leaflets acute or acuminate at the apex....... 2 

1b. Leaflets obtuse… D. axillare var. axillare 

2a. Plants densely pubescent ........………………. 

.…………… D. axillare var. acutifolium 

2b. Plants puberulent..........…………………… 

........…………. D. axillare var. stoloniferum 

2. Desmodium incanum DC., Prodr. 2: 332. 

1825. 

Fig. 103. D-G 

SYNONYM: Hedysarum canum J.F. Gmel. 

Desmodium canum (J.F. Gmel.) Schinz & Thell. 

Hedysarum supinum Sw. 

Meibomia supina (Sw.) Britton 

Decumbent or scandent herb, much branched 

from the woody base, attainig 1-1.5 m in length. 

Stems more or less cylindrical, appressedpubescent 

or pilose, glabrescent with age. Leaves 

alternate, trifoliolate; leaflets 3-9 × 0.8-3.8 cm, 

elliptical, oblong, ovate, lanceolate, or 

exceptionally rounded, chartaceous, involute, the 

apex obtuse or acute, mucronate, the base obtuse 

or rounded, the margins entire; upper surface dark 

green, puberulent; lower surface grayish green, 

strigulose, with prominent venation; petiolules 

1.5-3 mm long, pilose; petioles 0.7-3 cm long, 

pilose, canaliculate; rachis 3-10 mm long, pilose, 

canaliculate; stipules ovate to lanceolate, 6-10 mm 

long, persistent; stipels minute, subulate. 

Inflorescences of terminal pseudoracemes, 10-18 

cm long, pubescent; bracts minute, persistent. 

Calyx green, campanulate, 2.5-3 mm long, 

strigulose; corolla pink or pink-violet, the 

standard ovate, 5-6 mm long, retuse, unguiculate 

at the base, the wings and the keel as long as the 

standard. Legume uncinate-pubescent, 2-4 cm 

long, crenate on the ventral suture, with 5-8 fertile 

segments. Seeds 2.5-3.5 mm long, oblongelliptical, 

light brown. 



Status: Apparently native, very common. 

Distribution: In the understory of dry to moist 

forests and in areas of disturbed vegetation. Also 

on Mona, Vieques, St. Croix, St. John, St. Thomas, 

Tortola, and Virgin Gorda; distributed throughout 

the tropics as a weed. 


Guánica, Maricao, Mona, Río Abajo, and Susúa. 

3. Desmodium intortum (Mill.) Urb., Symb. 

Antill. 8: 292. 1920. 

Fig. 103. H-M 

BASIONYM: Hedysarum intortum Mill. 

Decumbent or ascendent herb, branched from 

the base, attainig 3 m in length. Stems trigonal, 

densely covered with uncinate hairs. Leaves 

alternate, trifoliolate; leaflets 4-7 × 2-5 cm, ovate, 

broadly ovate, or elliptical, chartaceous, the apex 

obtuse or acute, the base rounded, the margins 

entire; upper surface dark green, strigulose; lower 

surface pale green, more or less strigose, with 

prominent venation; petiolules 2.5-3 mm long, 

densely pilose; petioles 2-3.5 cm long, pilose, 

striate; rachis ca. 5 mm long, pilose; stipules 

ovate, acuminate, ca. 7 mm long, deciduous; 

stipels minute, subulate. Inflorescences of 

terminal or axillary pseudoracemes, uncinatepubescent. 

Calyx green, campanulate, puberulent, 

the tubular portion ca. 3 mm long, the inner sepal 

ca. 3.5 mm long, lanceolate, the rest ca. 1.5 mm 

long, deltate; corolla pink-violet, the standard ca.


9 mm long, retuse at the apex, unguiculate at the 

base, the wings and the keel as long as the 

standard. Legume uncinate-pubescent, ca. 2.5 cm 

long, crenate on both margins, with 6-11 fertile 

segments. 


during January and February. 

9. DIOCLEA 


Distribution: On the edges of forests along the 

Cordillera Central. Also in Hispaniola and 

Jamaica; southwestern United States, Mexico, and 

Central America to Peru. 

Public Forest: Toro Negro. 

Woody vines, twining, occasionally erect subshrubs. Leaves alternate, trifoliolate; stipels minute; 

stipules conspicuous, persistent, or absent. Inflorescences of axillary pseudoracemes, fasciculate, the 

flowers grouped on the swellings along the rachis; bracts and bracteoles minute, deciduous. Calyx 

campanulate, with 4 lobes, the upper lobe entire or emarginate; corolla purple or rarely white, the 

standard reflexed, emarginate, auriculate and usually with 2 callosities at the base, the wings free, 

sometimes spurred, the keel distally fused, rostrate; stamens 10, pseudomonadelphous; ovary superior, 

villous, stipitate or sessile, with 1 or more ovules, the style flattened or cylindrical, the stigma capitate, 

terminal or subterminal. Fruit a linear, oblong, obovate, or almost circular legume, compressed or 

turgid, coriaceous or woody, dehiscent or indehiscent; seeds large, compressed to globose, with a linear, 

elongate hilum. A genus of about 55 species, distributed throughout the tropics, with the greater number 

of species in South America. 

1. Dioclea reflexa Hook. f., Niger Fl. 306. 1849. 

Fig. 104. A-H 

Bejuco de mato, Maya prieta 

Woody vine, twining, attainig 10 m in length. 

Stems pilose, cylindrical, glabrescent when 

mature, attaining 1 cm in diameter. Leaves 

alternate, trifoliolate; leaflets 7-16 × 5-10.5 cm 

(the terminal leaflet larger than the lateral ones), 

elliptical or less frequently oblong, chartaceous, 

the apex rounded, usually terminating in a short 

apiculus, the base rounded, unequal on the lateral 

leaflets, the margins entire or undulate, slightly 

revolute; upper surface green, dull, punctate, 

strigulose, especially on the midvein, the lateral 

veins sunken; lower surface pale green, dull, 

strigulose, with prominent pilose venation; 

petiolules thickened, 5-6 mm long, pilose; stipels 

8-10 mm long, linear; petioles 7-9 cm long, 

sulcate, pilose, with the base thickened; stipules 

subulate, ca. 1 cm long, sericeous. Inflorescences 

of axillary or terminal pseudoracemes; rachis ca. 

40 cm long, thick, woody, sericeous; flowers in 

groups of three on each of the nodal swellings 

along the rachis; bracts subulate, ca. 1.5 cm long, 

reflexed, sericeous. Calyx ca. 1.5 cm long, 

ferruginous-strigulose, with five lobes, one of 

which is lanceolate and longer, the rest ovate; 

corolla cardinal red or violet, the standard ca. 1.2 

cm long, with the center whitish inside, the wings 

and the keel unguiculate, as long as the standard. 

Legume 10-15 × 5-6 cm, oblong, laterally 

compressed between the seeds, densely pubescent, 

glabrescent on drying, the margins thickened. 

Seeds 1-3, circular, 2-3.5 cm long, brown with a 

black hilum along 2/3 - 3/4 of the circumference. 


to December and in fruit from January to March. 


Distribution: In forests and pastures at lower 

or middle elevations. Also in Cuba, Hispaniola, 

Jamaica, Central America, South America, and 

tropical western Africa. 


Commentary: In this work I follow Maxwell 

(1999), who considers D. reflexa as a species 

distinct from the Asian species, D. hexandra 

(Ralph) Mabberley. 

The seeds of D. reflexa superficially resemble 

those of Mucuna urens, since they are of similar 

size and both possess a hilum that covers a large 

portion of its circumference. Nevertheless, the 

seeds of D. reflexa are distinguished from those 

of the latter species by their light brown color with 

brown spots (vs. uniformly reddish brown) and 

by the presence of a dark brown hilum (vs. a black 

hilum).

260 



262 


10. GALACTIA 

Herbaceous or slightly woody vines, twining. Leaves alternate, trifoliolate or unifoliolate; stipels 

minute or absent; stipules minute, deciduous. Inflorescences of axillary or terminal pseudoracemes, 

the flowers grouped on the swellings along the rachis; bracts and bracteoles minute, deciduous or 

persistent. Calyx campanulate, with 4 lobes, elongate; corolla pink, lavender, white, or less frequently 

red, the standard elliptical or rounded, reflexed, narrow at the base, the wings appressed to the keel; 

stamens 10, monadelphous or diadelphous, unequal; ovary superior, sessile, pubescent, with numerous 

ovules, the style curved, glabrous, the stigma capitate. Fruit a flattened, linear legume, slightly curved, 

with a beak at the apex, dehiscent by twisting valves; seeds small, few, ovoid, brown. A genus of about 

50 species, the majority of the New World tropics. 

Key to the species of Galactia 

1a. Corolla brilliant red; standard 14-18 mm long; species endemic to the Virgin Islands 

…………………………………………………………………………...............…… 2. G. eggersii 

1b. Corollas pink or lavender; standard 5-15 mm long; species of wide distribution ……….. 2 

2a. Leaflets linear or linear-oblong; inflorescences usually of a single flower; calyx strigose 

..……………………………………………………….………………..….. 3. G. longifolia 

2b. Leaflets ovate, elliptical, oblong, or lanceolate; inflorescences of 4 or more flowers; calyx pilose or 

pubescent (sometimes strigulose) …………………………………………….…........... 3 

3a. Legumes 5-5.5 mm wide; standard 12-15 mm long ………………………..…... 1. G. dubia 

3b. Legumes 6-9 mm wide; standard 8-10 mm long ………………………...….… 4. G. striata 

1. Galactia dubia DC., Prodr. 2: 238. 1825. 

Fig. 105. A-E 

SYNONYMS: Galactia dubia DC. var. ehrenbergii Urb. 

Galactia dubia DC. var. guanisensis Urb. 

Iron weed 


length. Stems slender, green, appressedpubescent. 

Leaves alternate, trifoliolate; leaflets 

1-4 × 0.8-2 cm, subcoriaceous, elliptical, oblong, 

or obovate, involute, the apex emarginate, 

rounded, mucronate, the base rounded, the 

margins entire; upper surface dark green, shiny 

or dull, puberulent or pubescent; lower surface 

pale green, dull, strigose or sericeous-pubescent, 

with the midvein prominent; petioles slender, 

pubescent, 1-2 cm long; stipules lanceolate, 2-3 

mm long; stipels absent. Inflorescences of axillary 

pseudoracemes, few-flowered; rachis 1-4 cm long; 

flowers 2-3, clustered on the swellings of the 

rachis; bracts minute, persistent. Calyx almost 

campanulate, green, pilose, 5-10 mm long, the 

sepals 4, ovate, acuminate, 4-7 mm long, two of 

which are longer; corolla pale pink or lavender, 

the standard oblong-elliptical, 12-15 mm long, 

reflexed, greenish at the base on the inner surface, 

the wings and the keel 10-12 mm long. Legume 

3-6 × 0.5-0.6 cm, oblong-linear, flattened, slightly 

curved, pubescent. Seeds ca. 4 mm long, oblong, 

dark brown. 

Phenology: Flowering from January to 

October and fruiting from January to May. 


Distribution: In areas of disturbed vegetation, 

at lower elevations, mostly near the coast. Also 

on Culebra, Culebrita, Desecheo, Icacos, Mona, 

Vieques, St. Croix, Little St. James, St. John, St. 

Thomas, and Tortola; the Lesser Antilles. 

Public Forests: Cambalache, Guánica, Mona, 


2. Galactia eggersii Urb., Symb. Antill. 2: 311. 

1900. 

Fig. 105. F-H



length. Stems slender, green, appressedpubescent, 

glabrous when mature. Leaves 


subcoriaceous, elliptical to rounded, involute, the 

apex emarginate, rounded, mucronate, the base 

rounded, the margins entire; upper surface dark 

green, slightly shiny, puberulent; lower surface 

pale green, dull, pubescent, with the midvein 

prominent; petioles 0.5-2 cm long; stipules 

lanceolate, 2-3 mm long; stipels subulate. 

Inflorescences of axillary pseudoracemes, of one 

or a few flowers; rachis 1-2.5 cm long, pubescent; 

flowers 1-3, clustered on the swellings of the 

rachis; bracts minute, persistent. Calyx almost 

campanulate, green, pilose, 8-12 mm long, the 

sepals unequal; corolla brilliant red, the standard 

elliptical, 14-18 mm long, reflexed, the wings and 

the keel as long as the standard; style white. 

Legume 5-6 × 0.5-0.6 cm, oblong-linear, flattened, 

slightly curved, pubescent, with the margins 

slightly sinuate. Seeds ca. 5 mm long, reniform, 

dark brown, shiny. 



Status: Endemic, uncommon. 

Distribution: In coastal thickets or in areas of 

disturbed vegetation, along the littoral zone. 

Endemic to Guana Island, St. John, St. Thomas, 

and Tortola. 

3. Galactia longifolia (Jacq.) Benth., Ann. 

Wien. Mus. 2: 127. 1838. 

Fig. 105. I-L 

BASIONYM: Galega longifolia Jacq. 

Decumbent herb, branched from the base, the 

branches twining in the distal portion, attaining 

2 m in length. Stems slender, cylindrical, 

sericeous, glabrescent with age. Leaves alternate, 

trifoliolate; leaflets 2-6.5 × 0.4-1(1.3) cm, 

subcoriaceous, linear, oblong or narrowly 

elliptical, the apex obtuse or acute, mucronate, 

the base rounded, the margins entire; upper 

surface dark green, dull, strigulose; lower surface 

pale green, dull, strigose to almost sericeous, with 

the midvein prominent, the lateral leaflets smaller 

than the terminal one; petioles slender, pubescent, 

1-10 mm long; rachis pubescent, longer than the 

petiole; stipules subulate, 1.5-3 mm long; stipels 

subulate, ca. 1 mm long, deciduous. Inflorescences 

of reduced, axillary pseudoracemes, usually with 

a single flower; rachis 1.7-5 cm long, strigulose 

or glabrescent. Calyx almost campanulate, green, 

strigose or strigulose, 3.5-5.5 mm long, the sepals 

4, elongate, lanceolate, acuminate; corolla pink 

to pale violet, the standard obovate, 5-7 mm long, 

reflexed, the wings and the keel as long as the 

standard. Legume 2.5-3.7 × ca. 0.4 cm, oblonglinear, 

flattened, slightly curved, appressedpubescent. 

Seeds ca. 2.8 mm long, elliptical, light 

brown. 


March to September. 

Status: Apparently native, but known from 

very few collections. 

Distribution: Known only from the Guayama 

area. Also in Hispaniola, the Lesser Antilles, the 

United States (Texas), and the Gran Chaco of 

Argentina and Paraguay. 

4. Galactia striata (Jacq.) Urb., Symb. Antill. 

2: 320. 1900. 

Fig. 105. M-O 

BASIONYM: Glycine striata Jacq. 

SYNONYMS: Galactia striata (Jacq.) Urb. var. tomentosa 

(Bertol.) Urb. 

Galactia berteriana DC. 

Galactia striata (Jacq.) Urb. var. berteriana (DC.) Urb. 


in length. Stems slender, pubescent. Leaves 

alternate, trifoliolate; leaflets 3-6 × 2-3 cm, 

elliptical or ovate, chartaceous, the apex rounded 

or less frequently obtuse, mucronate, the base 

rounded, the margins entire; upper surface dark 

green, dull, pubescent; lower surface pale green, 

dull, pilose or velutinous, with prominent 

venation; petiolules thickened, ca. 4.5 cm long, 

pubescent; stipels minute; rachis ca. 1 cm long; 

stipules filiform, ca. 3 mm long, pubescent. 

Inflorescences of axillary pseudoracemes; rachis 

4-14 cm long; flowers 2-3, grouped on the 

swellings along the rachis; bracts minute, 

persistent. Calyx green, ca. 1 cm long, pubescent, 

the sepals 4, one of them larger; corolla pink or 

lavender, the standard 8-10 mm long, elliptical, 

purple with white lines, yellow at the base on the 

inner surface, the wings and the keel as long as 

the standard. Legume 4-8 × 0.6-0.9 cm, oblong, 

flattened, pubescent, the apex with a recurved

264 



point. Seeds 4-5 mm long, flattened, reniform to 

oblong, dark brown, dull. 


August to March. 



at lower and middle elevations. Also on Culebra, 

11. LABLAB 


1. Lablab purpureus (L.) Sweet, Hort. Brit. ed. 

1, 481. 1826. 

Fig. 106. A-F 

BASIONYM: Dolichos purpureus L. 

SYNONYM: Dolichos lablab L. 

Chícharos, Frijol caballero, Habas de caballo, 

Bona wiss, Sweet pea 


in length. Stems slightly angular, pilose. Leaves 

alternate, trifoliolate; leaflets 5-16 × 4.5-12 cm, 

broadly ovate or rhombic, chartaceous, the apex 

acute or acuminate, the base cuneate or truncate 

on the central leaflet, unequal on the lateral ones, 

the margins entire, ciliate; upper surface dull, 

pubescent, with prominent venation; lower surface 

with the venation reticulate, prominent, 

pubescent; petiolules thickened, pubescent; 

petioles 6-20 cm long, canaliculate, laterally 

flattened, thickened at the base; stipules 

lanceolate, ca. 5 mm long, persistent; stipels 

subulate. Inflorescences of axillary 

pseudoracemes, erect, up to 25 cm long, the 

12. MACHAERIUM 




Public Forests: Cambalache, Guánica, 

Guajataca, Mona, Río Abajo, and Tortuguero. 

flowers 2-3, grouped on the swellings along the 

rachis; pedicels 4-5 mm long. Calyx campanulate, 

green, pubescent, 6-7 mm long, the sepals 4 or 5, 

unequal, lanceolate; corolla white or pale violet, 

the standard rounded, 1.4-2 cm long, the wings 

oblanceolate, the keel as long as the wings; 

stamens 10, diadelphous; ovary flattened, the style 

curved, the stigma terminal. Legume 5-10 × 2-3 

cm, almost oblong, broader in the portion near 

the apex, the upper margin with numerous 

callosities, the apex acuminate, tardily dehiscent. 

Seeds 3-5, up to 1 cm long, ovate or elliptical, 

flattened, light brown, with a white funiculus. 




Distribution: In disturbed areas, such as along 

roadsides, on fences, and in pastures. Probably 

native to Africa, found distributed throughout the 

tropics. Also on Vieques, St. Croix, St. John, and 

St. Thomas. 

Public Forests: El Yunque Maricao, Río Abajo, 


Trees, shrubs, or scandent lianas, usually with short, prehensile branches, spiny and with red 

latex. Leaves alternate, imparipinnate; leaflets alternate; stipels absent; stipules spinescent, persistent. 

Inflorescences of axillary or terminal racemes or panicles; bracts minute; bracteoles broadly ovate, 

paired at the base of the calyx. Calyx asymmetrically campanulate, 5-lobate or subtruncate; corolla 

violet-pink, white, or yellow, the standard rounded or reniform, narrowed at the base, the wings and 

the keel subequal; stamens 10, diadelphous or monadelphous; ovary short-stipitate, with 1-2 ovules, 

the style curved, the stigma punctiform or capitate. Fruit an indehiscent legume, usually samaroid, 

with a terminal wing, or flattened, straight, curved, or in the form of a half-moon, circular in outline, 

without a wing, or the wing reduced; seeds solitary, reniform, ovate, or orbicular. A genus of about 130 

species, the majority of the New World tropics.

266 


1. Machaerium lunatum (L.f.) Ducke, Arch. 

Jard. Bot. Rio de Janeiro 4: 310. 1925. 

Fig. 106. G-L 

BASIONYM: Pterocarpus lunatus L.f. 

SYNONYM: Drepanocarpus lunatus (L. f.) G.F.W. Meyer 

Escambrón, Palo de hoz 

Scandent shrub attainig 15 m in length. Stems 

cylindrical, attaining up to 12 cm in diameter, 

with numerous persistent, recurved stipular 

spines. Leaves imparipinnate; leaflets 5-7, 

alternate or rarely opposite, 2-7 × 0.5-3 cm, oblong 

or oblanceolate, chartaceous, glabrous, the apex 

retuse, the base obtuse or rounded, the margins 

entire; petiolules 2-3 mm long; stipules recurved, 

spiniform, up to 1 cm long. Inflorescences of 

axillary or terminal racemes, with few to many 

13. MACROPTILIUM 

flowers, 5-15 cm long; bracts minute, persistent. 

Calyx 3.5-5 mm long, glabrous; corolla pink to 

violet-pink, the standard up to 8 mm long, retuse, 

the wings falcate, the keel incurved. Legume 2- 

3.7 cm wide, in the form of a half-moon, circular 

in outline, indehiscent. Seed 1, reniform. 

Phenology: Collected in flower from February 

to August and in fruit from March to December. 


Distribution: In the coastal swamps that border 

on the mangroves. Also on Vieques, St. Croix, St. 

John, and St. Thomas; Hispaniola, the Lesser 

Antilles, tropical continental America and tropical 

Africa. 

Public Forest: In all probability, found in 

Piñones. 

Erect, creeping, or clambering herbs or herbaceous vines. Leaves trifoliolate; stipules and stipels 

minute. Inflorescences of axillary pseudoracemes, the flowers grouped in pairs on the swellings along 

the rachis; bracts minute. Calyx almost campanulate or tubular, of 5 equal or unequal sepals; corolla 

usually cardinal red, the standard rounded, reflexed, narrowed at the base, the wings narrowed at the 

base, much longer than the other petals, the keel narrowed at the base, twisted in the distal portion, 

fused to the staminal tube; stamens 10, diadelphous; ovary superior, subsessile, flattened, pubescent, 

with many ovules, the style thickened at the base, twisted, the stigma punctiform. Legume linear, 

cylindrical, dehiscent by valves that twist on drying; seeds numerous, small, oblong-cylindrical. A 

genus of 20 species, native to the New World. 

Key to the species of Macroptilium 

1a. Twining vine; terminal leaflet usually trilobate; lower surface densely white-pubescent; corolla 

purple ……..……………………………………………..…..................... 1. M. atropurpureum 

1b. Erect or clambering herb; terminal leaflet not lobed; lower surface puberulent; corolla maroon, 

pink, or white …………………………………….………...……........... 2. M. lathyroides 

1. Macroptilium atropurpureum (DC.) Urb., 

Symb. Antill. 9: 457. 1928. 

Fig. 107. A-D 

BASIONYM: Phaseolus atropurpureus DC. 

Herbaceous vine, twining, attainig 3-5 m in 

length. Stems cylindrical, strigose. Leaves 


coriaceous, upper surface dull, pubescent; lower 

surface densely white-pubescent, with prominent 

venation, the apex acute or obtuse; terminal leaflet 

ovate-rhombic, sometimes bilobate, the base 

cuneate; the lateral leaflets asymmetrical, 

unilobate, the base cuneate-obtuse; petiolules 

slender, white-pubescent; petioles 1-7 cm long, 

white-pubescent; stipules subulate to triangular, 

white-pubescent, 2.5-3 mm long; stipels subulate, 

white-pubescent, ca. 2 mm long. Inflorescences 

of axillary pseudoracemes, erect, 13-26 cm long, 

the flowers grouped in pairs on the swellings on 

the distal portion of the rachis; pedicels ca. 2 mm 

long. Calyx tubular-campanulate, green, with a 

reddish tonality, strigose-sericeous, 5-7 mm long, 

the sepals subequal or unequal, or all of them


268 


shorter than the tubular portion of the calyx; 

corolla purple, the standard rounded, ca. 2 cm 

long, the wings, shorter than the standard, 

unguiculate, the keel recurved in a spiral; stamens 

10, diadelphous; ovary flattened, sericeous, the 

style slightly curved, the stigma terminal, 

punctiform. Legume 5-8 × 0.2-0.4 cm, linear, 

flattened, sericeous. Seeds numerous, ca. 3.6 mm 

long, dark brown, dull. 


January to July. 

Status: Exotic, uncommon, a relatively recent 

introduction, since it was not reported for Puerto 

Rico by Britton and Wilson in 1924. 

Distribution: In disturbed, ruderal areas. Also 

on St. Thomas. Native to tropical continental 

America, but distributed throughout the tropics. 

Public Forest: Piñones. 

2. Macroptilium lathyroides (L.) Urb., Symb. 

Antill. 9: 457. 1928. 

Fig. 107. E-J 

BASIONYM: Phaseolus lathyroides L. 

SYNONYM: Phaseolus semierectus L. 

Habichuela parada, Wild bush bean 

Erect or clambering herb, scarcely twining, 

with numerous basal or lateral branches, attainig 

1.5-2(3) m in length. Stems almost cylindrical, 

up to 5 mm in diameter, pubescent, glabrescent. 

Leaves alternate, trifoliolate; leaflets elliptical, 

ovate, or lanceolate, 3-5 × 2-3 cm, chartaceous, 

14. MUCUNA 

the apex mucronate, acute or obtuse, the base 

obtuse or cuneate, the margins entire; upper 

surface glabrous, dark green, dull; lower surface 

pale green, dull, puberulent, with prominent 

venation; petiolules thickened, pubescent, 2-3 mm 

long; petioles sulcate, pubescent, sometimes 

reddish, with the base thickened; stipules 

lanceolate, 5-10 mm long, with notable parallel 

venation; stipels subulate, minute. Inflorescences 

of axillary pseudoracemes, 15-30 cm long, the 

flowers paired on the swellings on the distal 

portion of the rachis. Calyx campanulate, 5-7 mm 

long, strigose or puberulent, the sepals lanceolate, 

two of which are slightly smaller than the rest; 

corolla maroon, pink, and sometimes white, the 

standard 1-1.5 cm long, the wings longer than 

the standard, concave, the keel shorter than the 

standard, recurved in the form of a spiral. Legume 

linear, somewhat flattened and recurved, 6-12 cm 

× ca. 3 mm, dehiscent by valves that open in a 

spiral. Seeds numerous, 2-3 mm long, oblongreniform, 

dark brown. 



Status: Apparently native, very common. 

Distribution: In areas of disturbed vegetation 

throughout the island. Also on Vieques, St. Croix, 

St. John, St. Thomas, and Tortola; throughout the 

Antilles, United States (Florida), Central and 

South America, and the tropics of the Old World. 

Public Forests: Guánica, El Yunque, Maricao, 


Lianas or twining vines. Leaves alternate, trifoliolate; stipels absent or present; stipules deciduous. 

Inflorescences of pendulous axillary pseudoracemes, usually with a long peduncle; bracts foliaceous, 

deciduous. Calyx campanulate, bilabiate, with 4 lobes, one of which is smaller; corolla violet, violetpink, 

bluish, or yellow, the standard oblong, elongate, narrowed at the base, auriculate, the wings and 

the keel subequal, longer than the standard; stamens 10, diadelphous; ovary superior, sessile, villous, 

with few ovules, the style filiform, the stigma punctiform. Fruit an oblong legume, coriaceous, usually 

covered with stinging hairs, dehiscent; seeds oblong, circular, rounded, with an oblong, elongate hilum. 

A genus of about 120 species distributed throughout the tropics. 

Key to the species of Mucuna 

1a. Seeds oblong (in the form of a bean), with a short hilum; legume 4-9 cm long, ca. 1.5 cm wide, 

cylindrical, densely hispid …………………………………………....… 1. M. pruriens 

1b. Seeds circular, flattened, with a linear, black hilum, extending over 2/3 to 3/4 of the circumference; 

legume 8-20 cm long, 2-5 cm wide, more or less flattened, falcate, sparsely hispid............ 2


270 


2a. Lower surface of leaflets sericeous-pubescent; corolla yellow…………….….2. M. sloanei 

2b. Lower surface of leaflets glabrous; corolla bluish or violet-pink .…………..…3. M. urens 

1. Mucuna pruriens (L.) DC., Prodr. 2: 405. 

1825. 

Fig. 108. A-E 

BASIONYM: Dolichos pruriens L. 

SYNONYM: Stizolobium pruriens (L.) Medic. 

Pica-pica, Cow-itch 


in length. Stems pubescent, cylindrical, striate, 

glabrescent. Leaves alternate, trifoliolate; leaflets 

chartaceous, the apex acute or obtuse, mucronate, 

the upper surface dark green, dull, puberulent, 

the lower surface pale green, strigulose, with 

prominent venation, the margins slightly sinuate; 

terminal leaflet rhombic-ovate, (5.5)7.5-15 × 4-6 

cm, the base obtuse, rounded to almost cuneate; 

lateral leaflets markedly asymmetrical, one side 

oblong, the other ovate, 5)7.5-13(18) x (2.8)4.5- 

7.2(10.3) cm, the base obtuse on one side, truncate 

on the other; petiolules pubescent, 4-6 mm long; 

rachis 1.2-2 cm long, strigose; petioles 4.5-16.5 

cm long, strigose, with the base swollen; stipules 

minute, caducous; stipels linear, 4-5 mm long. 

Inflorescences of axillary pseudoracemes, 10-30 

cm long, with 1-3 flowers per node. Calyx grayish 

green, campanulate-asymmetrical, 1-1.5 cm long, 

sericeous-pubescent, mixed with some stiff hairs, 

the sepals unequal, triangular, deep; standard 

ovate, 1.5-2 cm long, violet, the wings purple, 

oblong, ca. 4 cm long, the keel pale violet; 

staminal column almost white, exserted, the 

anthers violet. Legume more or less cylindrical, 

with the curved extremities forming an ‘s’, 4-10 

× 1-1.4 cm, densely covered with stiff, extremely 

stinging hairs, dehiscent by valves that open in a 

spiral. Seeds ellipsoid, brown with black spots, 

ca. 1 cm long, with a cream-colored hilum, ca. 5 

mm long. 


to April and in fruit from October to January. 

Status: Probably native, rather common. 


pastures, forest edges, and roadsides, at lower and 

middle elevations. Also on St. Croix; of 

widespread distribution throughout the tropics. 

Public Forests: Guajataca, Río Abajo, and 

Susúa. 

2. Mucuna sloanei Fawc. & Rendl., J. Bot. 55: 

36. 1917. 

Fig. 108. F-H 

SYNONYMS: Dolichos urens Jacq., non L. 

Mucuna urens sensu Urb., non (L.) DC. 

Matos, Matos del monte, Ojo de buey 

Twining liana attainig 10-15 m in length. 

Stems cylindrical, strigulose. Leaves alternate, 

trifoliolate; leaflets 8-12.5 × 5.7-7.5 cm, 

chartaceous, the obtuse apex usually terminating 

in a short acumen, the margins entire; upper 

surface strigulose, dark green, dull; lower surface 

pale green, dull, sericeous-pubescent (silvery), 

with prominent venation; terminal leaflet oblongelliptical 

or ovate, the base obtuse, rounded to 

almost truncate; lateral leaflets very asymmetrical, 

one side lanceolate, the other elliptical, the base 

obtuse, truncate, or almost cordiform; petiolules 

thickened, strigose, ca. 5 mm long; petioles 

sulcate, glabrescent, up to 11 cm long; stipules 

deciduous; stipels subulate, ca. 4 mm long. 

Inflorescences of axillary pseudoracemes, short, 

with few flowers at the end. Calyx yellowish 

green, broadly campanulate, 1.5-2 cm long, 

sericeous; corolla yellow, the standard 2.5-4 cm 

long, the wings and the keel, elongate, falcate, 

longer than the standard; style appressedpubescent. 

Legume oblong, flattened, 8-18 × 4-5 

cm, densely covered with stiff stinging hairs. 

Seeds 2-4, circular, 2-3 cm long, brown with the 

black hilum along ¾ of the circumference. 



Distribution: Known from few collections 

from the western or central-western area (Aguada, 

Mayagüez, Sabana Grande, Lares). Also in 

Jamaica, Cuba, Hispaniola, some of the Lesser 

Antilles, Central and South America. 

Public Forests: Maricao and Susúa. 

3. Mucuna urens (L.) DC., Prodr. 2: 405. 1825. 

Fig. 108. I-M 

BASIONYM: Dolichos urens L. 

SYNONYM: Mucuna altissima (Jacq.) DC. 

Tortera, Mato, Pica-pica, Ox-eye-bean


272 



Stems cylindrical, glabrous, smooth, that turn 

blackish on drying, attaining up to 10 cm in 

diameter at the base and producing a watery sap 

with a pink color. Leaves alternate, trifoliolate, 

usually pendulous: leaflets coriaceous, glabrous, 

7-14 × 4.5-7 cm, the margins entire, upper surface 

dark green, shiny, glabrous; lower surface light 

green (purple on juvenile leaflets), shiny, glabrous, 

with prominent puberulent venation; terminal 

leaflet elliptical or oblong, the apex acuminate, 

the base rounded; the lateral leaflets asymmetrical, 

elliptic-lanceolate, the apex acuminate, the base 

unequal, truncate-obtuse; petiolules slightly 

thickened, 5-6 mm long, glabrous or puberulent; 

stipels absent; petioles 4-9 cm long, striate, 

thickened at the base; stipules deciduous. 

Inflorescences of pendulous pseudoracemes; 

peduncles 1-1.5 m in length. Calyx 1.3-1.7 cm 

long, ferruginous-pubescent, with one of the 

sepals longer than the rest; pedicel ca. 2 cm long, 

ferruginous-pubescent; corolla blue-violet or 

15. NEORUDOLPHIA 

cardinal red, the standard 3-4 cm long, the wings 

violet-pink, the keel yellowish, shorter than the 

wings. Legume woody, more or less oblong, 

flattened, 12-23 × 5-6 cm, the apex with a long 

point, the valves with prominent venation, like 

transverse ribs, which join at the ventral margin 

to form a sinuate projection, covered with rigid, 

rust-colored hairs, which separate with great 

facility and produce a sharp pain on contact. Seeds 

ca. 3.5 cm wide, circular, hard, brown with a black 

hilum along more than ¾ of the circumference. 


to April and in fruit in February. 


Distribution: On river banks, in moist forests 

at middle and lower elevations, in the zone of 

mogotes. Reported for St. Thomas by Krebs 

(1847), probably in error. Also in Jamaica, Cuba, 

Hispaniola, the Lesser Antilles, Trinidad, Central 

America, and northern South America. 

Public Forests: El Yunque, Guilarte, Maricao, and 

Río Abajo. 

A monospecific genus, endemic to Puerto Rico, characterized by the following species. Very similar 

to the genus Rhodopis of the Dominican Republic; nevertheless, both differ markedly in the morphology 

of their flowers. 

1. Neorudolphia volubilis (Willd.) Britton in 

Britton & Wilson, Bot. Porto Rico 5: 426. 

1924. 

Fig. 109. A-H 

BASIONYM: Rudolphia volubilis Willd. 

Bejuco prieto, Bejuco colorado, Bejuco de 

alambac, Bejuco de alambre 

Woody vine, twining, attainig 5-10 m in 

length. Stems strong, flexible, cylindrical, slightly 

flattened and with numerous lenticels when 

mature. Leaves alternate, unifoliolate; leaflets 

coriaceous, (5)6-13 × 4-9 cm, ovate or broadly 

ovate, the apex long- or short-acuminate, the base 

cordiform or less frequently truncate, the margins 

entire or sinuate, revolute; upper surface dark 

green, slightly shiny, with slightly prominent 

venation and puberulent; lower surface yellowish 

green, with the reticulate venation prominent; 

petiolule thickened, pubescent, 4-5 mm long, with 

a pair of stipels at the base; petioles 2.5-5 cm long, 

sulcate, puberulent, with the base broadened. 

Inflorescences of cauliflorous or axillary 

pseudoracemes, pendulous, 12-24 cm long, the 

flowers in groups of 2-3 per node of the 

inflorescence; pedicels 10-12 mm long, pink; 

bracts minute, lanceolate. Calyx 2.5-3 cm long, 

pink, fleshy, almost campanulate, the sepals 5, 

two of which are broad and rounded, the 

remaining three lanceolate; corolla brilliant red, 

the standard ca. 5 cm long, oblong, folded in half 

longitudinally, enclosing the stamens, the 

remaining 4 petals (the wings and two petals 

homologous to the keel, which are not fused) ca. 

1.5 cm long, linear and recurved; stamens 10, 

diadelphous, ca. 4 cm long, slightly exposed; 

ovary short-stipitate, with numerous ovules, the 

style slender, elongate, sericeous, the stigma 

punctiform. Fruit an oblong legume, flattened, 6- 

12 × 1-1.5 cm, pubescent, dehiscent by valves that


274 


open in a spiral. Seeds oblong or ovoid, 6-8 mm 

long, light brown. 

Phenology: Flowering from June to December 

and fruiting from November to April. 

Status: Endemic, very common. 

16. PACHYRHIZUS 

17. PHASEOLUS 

Distribution: In the moist or wet forests of the 

Cordillera Central, the Sierra de Luquillo, and in 

the area of mogotes, at middle and upper 

elevations. 

Public Forests: Carite, Guajataca, Guilarte, 

El Yunque, Maricao, Río Abajo, Susúa, and Toro 

Negro. 

Herbaceous or slightly woody vines, twining, with tuberous roots. Leaves alternate, trifoliolate; 

stipels filiform; stipules lanceolate, persistent. Inflorescences of axillary or terminal pseudoracemes, 

long-pedunculate; bracts minute. Calyx campanulate, bilabiate, with 5 lobes; corolla blue or violet, the 

standard broadly obovate, auriculate, oblong, the wings oblong-falcate, with a curved appendage at the 

base of the inner margin, adhering to the keel along the basal portion of the inner margin, the keel as 

long as the wings, recurved; stamens 10, diadelphous; ovary superior, subsessile, with many ovules, 

the style curved, the stigma globose. Fruit an oblong legume, coriaceous, flattened, dehiscent, the 

valves septate internally between the seeds; seeds flattened, with a hilum of reduced size. A genus of 6 

species, distributed throughout the Neotropics. 

1. Pachyrhizus erosus (L.) Urb., Symb. Antill. 

4: 311. 1905. 

Fig. 110. A-E 

BASIONYM: Dolichos erosus L. 

SYNONYM: Cacara erosa (L.) Kuntze 

Jícama 


length. Tuberous roots thick. Stems cylindrical, 

slightly ribbed, pilose. Leaves alternate, 

trifoliolate; leaflets chartaceous, the apex shortacuminate, 

the margins deeply dentate, the teeth 

mucronate; upper surface dark green, dull, 

sparsely appressed-pubescent, with slightly 

prominent venation; lower surface pale green, 

sparsely appressed-pubescent, with the reticulate 

venation prominent; terminal leaflet 8-16 × 9-19 

cm, rhombic, the base cuneate or sub-reniform; 

lateral leaflets asymmetrical, broadly ovateelliptical, 

the base cuneate-subtruncate; petiolules 

thickened, pubescent, with a pair of stipels at the 

base; rachis 3-4 cm long, pilose; petioles 8-13 cm 

long, sulcate, pilose, with the base broadened. 

Inflorescences of axillary pseudoracemes, erect, 

30-45 cm long, the flowers in groups of 1-5 per 

node of the inflorescence; pedicels 6-8 mm long; 

bracteoles minute, oblong. Calyx 9-11 mm long, 

green, campanulate, the sepals 4, one larger that 

the others; corolla blue or blue-violet, the standard 

semicircular, ca. 1.5 cm long, emarginate, the 

wings and the keel as long as the standard; 

stamens 10, diadelphous. Fruit an oblong legume, 

flattened, 7-15 × 1.1-2 cm, pubescent. Seeds 

quadrangular, flattened, ca. 7 mm long, dark 

brown. 

Phenology: Collected in flower during 

September and in fruit in January, February, and 

August. 



Distribution: In areas of degraded vegetation, 

along trails and in secondary forests. Native of 

southern Mexico, widely cultivated throughout the 

tropics for its edible tuberous roots. 

Public Forest: Guajataca. 

Erect herbs or twining vines, with a pubescence of uncinate hairs. Leaves alternate, trifoliolate, 

with the rachis more or less elongate; stipules striate, truncate at the base, persistent; stipels minute. 

Inflorescences of axillary racemes, with the nodes not swollen and lacking extrafloral nectaries; bracts


minute, persistent. Calyx campanulate, bilabiate, with 5 minute lobes at the apex; corolla white, pink, 

red, purple, or yellow, the standard symmetrical, rounded, unguiculate, reflexed, the keel spirally twisted, 

narrow; stamens 10, diadelphous; ovary almost sessile, linear, with one or more ovules, the style 

spirally twisted, (1.5-2 turns), barbate, the stigma terminal. Fruit a linear or oblong legume, straight, 

dehiscent; seeds oblong. A genus of 200 species, of cosmopolitan distribution. 

Key to the species of Phaseolus 

1a. Legume falcate, curved, up to 2 cm wide; leaflets truncate at the base ……….............. 1. P. lunatus 

1b. Legume oblong, straight or slightly curved, up to 1 cm wide; leaflets rounded or cuneate at the 

base………..……....……………………………………...…………........................…. 2. P. vulgaris 

1. Phaseolus lunatus L., Sp. Pl. 724. 1753. 

Fig. 110. F-I 


length. Stems obtuse-pentagonal or cylindrical, 

puberulent. Leaves alternate, trifoliolate; leaflets 

chartaceous, the apex acute or short-acuminate, 

the margins sinuate; upper surface dark green, 

dull, glabrous, with slightly prominent venation; 

lower surface pale green or glaucous, glabrous, 

with the primary and secondary venation 

prominent; terminal leaflet 5.5-11 × 3.5-7.5 cm, 

rhombic or deltate, the base cuneate or truncate; 

lateral leaflets asymmetrically deltate, the base 

truncate; petiolules thickened, 3-5 mm long, 

pubescent; rachis 1.5-2.5 cm long; petioles 6.5-9 

cm long, sulcate, puberulent, the base slightly 

broadened. Inflorescences of axillary 

pseudoracemes, erect, 3-30 cm long, the flowers 

in groups of 2 per node of the inflorescence; 

pedicels 6-9 mm long, pilose; bracteoles minute, 

oblong. Calyx 2-2.5 mm long, green, 

campanulate, pilose, the sepals deltate, subequal; 

corolla white or lilac, pink or bluish, the standard 

semicircular, 7-10 mm long, abaxially pilose, the 

wings obovate, unguiculate, as long as the 

standard, the keel spirally twisted, ca. 1 cm long; 

stamens 10, diadelphous, the vexillar stamen 

broadened at the base; ovary with hispidulous 

pubescence, intermingled with uncinate hairs. 

Fruit an oblong-falcate legume or in the form of 

a half-moon, flattened, 5-7 × 1-2 cm, puberulent 

with uncinate hairs or glabrescent, dehiscent by 

valves that open in a spiral. Seeds reniform, 

flattened, ca. 7 mm long, reddish brown, with dark 

spots. 


December to June. 




along trails, on fences, and in thickets. Also in 

the Antilles. Probably native to tropical 

continental America, where it is widely cultivated. 

Introduced in the tropics of the Old World. 

2. Phaseolus vulgaris L., Sp. Pl. 723. 1753. 

Fig. 110. J-M 

Habichuelas, Frijoles 

Herbaceous vine, climbing or creeping, 

attainig 3 m in length, or sometimes an erect herb. 

Stems obtuse-pentagonal or cylindrical, 

puberulent. Leaves alternate, trifoliolate; leaflets 

chartaceous, the apex acuminate, the margins 

entire; upper surface dark green, dull, appressedpuberulent, 

with slightly prominent venation; 

lower surface pale green, appressed-puberulent, 

with prominent venation; terminal leaflet 6-10 × 

5-7.5 cm, deltate or rhombic, the base cuneate or 

truncate; lateral leaflets asymmetrically deltate, 

the base truncate-obtuse; petiolules thickened, 4- 

6 mm long, pilose; rachis 1.8-2.8 cm long, 

marginate or slightly winged; petioles 8-15 cm 

long, adaxially flattened, marginate, puberulent; 

stipules triangular, ca. 4 mm long; stipels oblong, 

ca. 3 mm long. Inflorescences of axillary 

pseudoracemes, the rachis up to 8 cm long, 

uncinate-pubescent, the flowers usually 4, in 

groups of 2 per node, this with an ovate bract, 

persistent at the base; pedicels 7-8 mm long, 

pilose; bracteoles elliptic-ovate, rounded at the 

apex, up to 7.5 mm long. Calyx 3.5-4 mm long, 

green, obliquely campanulate, puberulent, the 

sepals unequal; corolla white to lilac, pink, or

276 



purple, the standard semicircular, ca. 10 mm long, 

glabrous, the wings obovate, unguiculate, longer 

than the standard, the keel spirally twisted, ca. 1 

cm long; stamens 10, diadelphous, the vexillar 

stamen broadened at the base; ovary strigulose. 

Fruit an oblong legume, slightly flattened, straight 

or slightly curved, 9-12 × 1-1.2 cm, glabrous, 

long-apiculate at the apex, dehiscent by valves 

that separate longitudinally. Seeds oblongreniform, 

1-1.2 mm long, of various colors, 

reddish brown, grayish with dark spots, pink. 


May to July and in November. 

18. PUERARIA 


uncommon. 


along trails, on fences, and in thickets. Also 

throughout the Antilles. Native to Mesoamerica, 

with numerous forms cultivated for millennia in 

tropical continental America. Widely cultivated 

throughout the world. 

Excluded Species: 

Phaseolus polystachyos (L.) Britton was 

reported for Puerto Rico by Liogier (1982, as P. 

polystachios), based on an erroneous 

identification of Phaseolus lunatus (Liogier, et al. 

31642). 

Herbaceous or woody vines, twining. Leaves alternate, trifoliolate; stipules ovate to linear, persistent; 

stipels minute. Inflorescences of axillary or terminal pseudoracemes; bracts minute. Calyx campanulate, 

with 5 unequal lobes; corolla blue or violet, the standard obovate, retuse at the apex, unguiculate and 

auriculate at the base, the wings unguiculate, with a curved appendage at the base of the inner margin, 

the keel slightly longer than the wings; stamens 10, monadelphous or diadelphous; ovary superior, 

sessile, with several ovules, the style glabrous, curved, the stigma capitate. Fruit a linear legume, 

flattened, dehiscent by valves that twist on opening; seeds oblong, numerous. A genus of 6 species 

native to southern Asia, with one species introduced in the New World as a forage plant. 

1. Pueraria phaseoloides (Roxb.) Benth., J. 

Linn. Soc., Bot. 9: 125. 1865. 

Fig. 111. A-D 

BASIONYM: Dolichos phaseoloides Roxb. 

Kudzu tropical, Corsú 

Herbaceous vine, twining, much branched, 

attainig 15 m in length. Stems cylindrical, pilose. 

Leaves alternate, trifoliolate; leaflets 3-12(14) × 

2.9-8.7(13) cm, chartaceous, ovate or rhombic, 

the lateral ones asymmetrical, the apex acute, the 

base cuneate on the central leaflet, rounded-obtuse 

on the lateral ones, the margins entire; upper 

surface dark green, dull, pubescent, especially on 

the veins; lower surface pale green, strigose, with 

prominent venation; petiolules swollen, 4-5 mm 

long, pubescent; petioles sulcate, pubescent, up 

to 12 cm long, with the base swollen; stipules 

narrowly lanceolate, 3-5 mm long; stipels 

subulate, minute, persistent. Pseudoracemes 

axillary, up to 25 cm long, with 2-3 flowers per 

node; bracts minute, persistent; peduncles 

pubescent. Calyx campanulate, ca. 5 mm long. 

Legume linear, flattened, 6-9 cm long, slightly 

curved, dehiscent by valves that open in a spiral, 

the valves septate between the seeds. Seeds 

numerous, ca. 3 mm long, oblong, dark brown to 

almost black. 



Status: Exotic, naturalized, very aggressive 

and very common. 

Distribution: In moist disturbed areas, at 

middle and lower elevations. Also on St. John. 

Native to tropical Asia, introduced in Africa and 

the Americas as a forage plant. 

Public Forests: Carite, El Yunque, Maricao, 

Río Abajo, and Tortuguero. 

Commentary: Apparently originally 

introduced on lands of the Agricultural 

Experiment Station in Mayagüez in Puerto Rico 

in 1940 from material from Malaya; today this 

species has a wide distribution throughout Puerto 

Rico.

278 


19. RHYNCHOSIA 

Herbaceous or woody vines, twining. Leaves alternate, trifoliolate, the lower surface with numerous 

yellow resinous dots; stipules deciduous; stipels minute. Inflorescences of axillary racemes, with the 

flowers sparse or clustered; bracts minute, persistent or deciduous. Calyx campanulate, with 4-5 elongate 

lobes; corolla yellow, the standard obovate or rounded, slightly retuse at the apex, unguiculate and 

auriculate at the base, the wings unguiculate, with a curved appendage at the base of the inner margin, 

the keel scarcely longer than the wings; stamens 10, monadelphous or diadelphous; ovary superior, 

short-stipitate, with few or numerous ovules, the style glabrous, curved, the stigma capitate. Fruit an 

oblong legume, flattened, apiculate at the apex, usually dehiscent by valves that twist on opening; 

seeds few or numerous, flattened, rounded or elliptical. A genus of about 200 species, of pantropical 

distribution. 

Key to the species of Rhynchosia 

1a. Robust lianas that attain 8-10 m in length; mature stems flattened, forming a ribbon up to 2.5 cm 

wide; seeds red and black ….………………………………….... 2. R. phaseoloides 

1b. Slightly woody vines, usually less than 5 m in length; mature stems cylindrical, less than 5 mm in 

diameter; seeds brown ……………………………………………………………. 2 

2a. Lower surface of the leaflets with the tertiary venation not prominent; calyx 2.5-3 mm long, the 

sepals short-subulate, as long as or twice as long as the tubular portion of the calyx 

..............…………………..…..……………………………...……………………1. R. minima 

2b. Lower surface of the leaflets with the reticulate tertiary venation prominent; calyx 6-10 mm long, 

the sepals linear-lanceolate, three times as long as the tubular portion of the calyx 

...................……………..…….…………………………………….…………….3. R. reticulata 

1. Rhynchosia minima (L.) DC., Prodr. 2: 385. 

1825. 

Fig. 111. E-H 

BASIONYM: Dolichos minimus L. 

SYNONYMS: Rhynchosia minima var. lutea Eggers 

Rhynchosia minima var. pauciflora Kuntze 

Dolicholus minimus (L.) Medic. 

Frijolillo, Habilla 

Non-woody vine, twining, attainig 1-4 m in 

length. Stems slender, densely pubescent, striate, 

cylindrical, with numerous resinous dots in the 

furrows. Leaves alternate, trifoliolate, 

chartaceous; margins entire, slightly revolute; 

upper surface dull, glabrous or white-pubescent, 

especially on the veins, with numerous orange 

resinous dots; lower surface pale green, glabrous, 

puberulent or pubescent, with minute orange 

resinous dots and the primary and secondary 

venation prominent; terminal leaflet rhombic, 0.9- 

2.8(7.5) × 0.6-2.8(6.3) cm, the apex acute or 

obtuse, the base cuneate; lateral leaflets 

asymmetrical, elliptical-ovate, slightly smaller 

than the terminal leaflet, the base obtuse; 

petiolules densely pubescent, 1-1.5 mm long; 

petiole striate, densely pubescent, 1.5-2 cm long; 

stipules subulate, 2-2.5 mm long, densely 

pubescent; stipels subulate, ca. 3 mm long. 

Inflorescence a raceme, as long as or longer than 

the leaves. Calyx green, the lobes lanceolate, 2-3 

mm long; corolla yellow with purple or brown 

venation, the standard 5 mm long, orbicularovate; 

wings oblong, unguiculate, 4-5.5 mm long, 

with resinous dots. Legume falcate to oblongovate, 

flattened, minutely villous, 9-20 × 3-5 mm, 

with a beak at the apex. Seeds 2-3, ovate-reniform, 

dark brown to almost black, 3-4 mm long. 



Status: Probably exotic, naturalized, very 

common. 

Distribution: Abundant on fences and in 

weedy places at middle and lower elevations. Also 

on Mona, Vieques, St. Croix, St. John, St. Thomas, 

Tortola, and Virgin Gorda; generalized in the 

tropics and subtropics of the world. Although this


species is very abundant in the New World, it is 

believed to be native to the Old World, because in 

the latter region it exhibits a great diversity of 

differentiation at the varietal level. 

Public Forests: Cambalache, Guánica, 

Guajataca, Mona, and Río Abajo. 

2. Rhynchosia phaseoloides (Sw.) DC., Prodr. 

2: 385. 1825. 

Fig. 112. A-H 

BASIONYM: Glycine phaseoloides Sw. 

SYNONYM: Dolicholus pyramidalis sensu Britton & 

Wilson, non Lam. 

Bejuco de paloma 


length. Stems cylindrical, sericeous, mature stems 

flattened like a ribbon, attaining 2.5 cm wide, 

glabrous. Leaves alternate, trifoliolate, 

chartaceous; margins entire or slightly sinuate; 

upper surface dark green, dull, strigulose or 

glabrous; lower surface pale green, lanatepubescent, 

with minute orange resinous dots, the 

venation prominent; terminal leaflet rhombic or 

deltate, 3.5-11.5 × 2.4-9.2 cm, the apex 

acuminate, the base cuneate to almost rounded; 

lateral leaflets asymmetrical, lanceolate-ovate, 

slightly smaller than the terminal leaflet, the base 

obtuse-rounded to cordiform; petiolules densely 

pubescent, 2-4 mm long; petiole densely 

pubescent, 4-7 cm long; stipules deciduous. 

Inflorescence a raceme, as long as or shorter than 

the leaves. Calyx green, puberulent, 3.5-4 mm 

long, asymmetrical, one of the lobes lanceolate, 

ca. 3 mm long, the rest ovate, ca. 1.5 mm long; 

corolla yellow, the standard 7-8 mm long, obovate, 

emarginate, punctate, yellow with a reddish tinge; 

wings oblong, unguiculate, 7-7.5 mm long, with 

resinous dots; ovary minutely sericeous. Legume 

oblong, flattened, with the margin sinuate between 

the seeds, with glandular dots, minutely sericeous, 

1.2-2.4 × 0.8-1.2 cm, the apex short-apiculate. 

Seeds usually two per fruit, ellipsoid, ca. 5 mm 

long, shiny, the longitudinal half nearer to the 

hilum red, the other half black. 



Distribution: In secondary forests in the zone 

of mogotes. Also on St. Thomas; throughout the 

Antilles, southern Panama, and South America. 

Public Forests: Maricao and Río Abajo. 

Commentary: The seeds of Abrus precatorius 

are apt to be confused with those of Rhynchosia 

phaseoloides, since both have a similar coloration. 

Nevertheless, in R. phaseoloides the hilum is on 

the red portion of the seed, while in A. precatorius 

the hilum is on the black portion. 

3. Rhynchosia reticulata (Sw.) DC., Prodr. 2: 

385. 1825. 

Fig. 111. I-K 

BASIONYM: Glycine reticulata Sw. 

SYNONYM: Dolicholus reticulatus (Sw.) Millsp. 

Frijolillo, Habilla 


length. Stems slender, angular, striate, sometimes 

almost triangular, ferruginous-tomentose when 

young. Leaves alternate, trifoliolate, chartaceous; 

margins entire or slightly sinuate, revolute; upper 

surface dark green, dull, puberulent, with the 

venation sunken; lower surface pale green, lanatepubescent, 

with minute orange resinous dots, the 

venation prominent; terminal leaflet rhombic, 2- 

12 × 1-7 cm, the apex obtuse or rounded, the base 

cuneate or obtuse; lateral leaflets asymmetrical, 

elliptical-ovate, smaller than the terminal leaflet, 

the base obtuse-cuneate; petiolules tomentose, 1- 

3 mm long; rachis 4-8 mm long, tomentose; 

petiole tomentose, 2-5 cm long; stipules 

lanceolate, up to 10 mm long, sericeous; stipels 

minute, only on the distal leaflet. Inflorescence a 

raceme, up to 10(25) cm long, with few flowers. 

Calyx green, pilose, 6-10 mm long, the lobes 

lanceolate; corolla yellow, the standard 6-10 mm 

long, obovate to orbicular, emarginate, glabrous 

or puberulent; wings obovate, unguiculate, longer 

than the standard; ovary minutely sericeous. 

Legume oblong-ovate, falcate, flattened, with one 

margin straight and the other curved, minutely 

sericeous, 1.5-3 × 0.8-1.1 cm, the apex shortapiculate. 

Seeds usually two per fruit, circular, 

flattened, ca. 5 mm in diameter, light brown with 

dark brown spots. 





pastures, weedy places, roadsides, and fences, at 

lower and middle elevations. Also on Culebra, 


Thomas, Tortola, and Virgin Gorda; also

280 



282 


throughout the Antilles, tropical continental 

America. 

Public Forests: Cambalache, Carite, Ceiba, 

Guajataca, Guánica, Maricao, Mona, Río Abajo, 



Rhynchosia caribaea (Jacq.) DC. has been 

cited for Puerto Rico by Liogier (1982, 1988), but 

is excluded from the present work. This species 

is native to Africa and very similar to R. minima, 

with which it has been confused in the New World. 

20. TERAMNUS 

Herbaceous vines, twining, climbing, or creeping. Leaves alternate, trifoliolate; stipules subulate, 

persistent; stipels minute. Inflorescences of axillary pseudoracemes; bracts small. Calyx campanulate, 

with 4-5 lobes of similar size; corolla white, yellow, or lavender, the standard obovate, the wings 

unguiculate, the keel shorter than the wings; stamens 10, monadelphous, the anthers unequal; ovary 

superior, sessile, with numerous ovules, the style short, pubescent, the stigma capitate. Fruit a linear 

legume, flattened, curved at the apex, dehiscent by valves that twist on opening; seeds numerous, 

flattened, oblong. A genus of 8 species, of pantropical distribution. 

Key to the species of Teramnus 

1a. Stems pilose; leaflets elliptical, ovate to almost rounded, strigose on the lower surface; legume 

pilose or glabrous, 2-5 cm long ....................………..…………………………..….1. T. labialis 

1b. Stems tomentose or sericeous; leaflets oblong or lanceolate, sericeous on the lower surface; legume 

ferruginous-tomentose, 4-7 cm long ........................…..………………..…2. T. uncinatus 

1. Teramnus labialis (L.f.) Spreng., Syst. Veg. 

3: 235. 1826. 

Fig. 113. A-E 

BASIONYM: Glycine labialis L.f. 

Frijolillo 

Twining vines, 3-5 m in length. Stems slender, 

pilose, almost hirsute when young. Leaves 

trifoliolate; leaflets 1.5-6.2 × 0.6-3.5 cm, 

chartaceous, the apex mucronate, obtuse or 

rounded, the base obtuse or rounded, the margins 

ciliate; upper surface appressed-pubescent; lower 

surface strigose; terminal leaflets elliptical; lateral 

leaflets asymmetrical, elliptic-oblong or ellipticovate; 

petiolules 2-3 mm long, pilose; stipules 

lanceolate, up to 3 mm long; stipels ca. 1.5 mm 

long, linear. Inflorescences of axillary 

According to Grear (1978), R. caribaea has been 

collected in Puerto Rico, and can be distinguished 

from R. minima by its viscid-hirsute fruits and by 

its larger flowers. The record of R. caribaea for 

Puerto Rico is based on two collections made in 

the nineteenth century, one of which was from a 

cultivated plant. This species has not been 

collected again in Puerto Rico, which suggests 

that it was unable to establish itself or to become 

naturalized. Rhynchosia caribaea is also 

unknown in our gardens; for this reason, it is 

excluded from our flora. 

pseudoracemes that attain 11 cm long, the flowers 

in pairs on the nodes of the rachis. Calyx 

campanulate, 3-5 mm long, strigose; corolla 

white, pink, or pale violet, the standard obovate, 

ca. 5 mm long, the wings and the keel shorter 

than the standard. Legume 2-5 × 0.3-0.4 cm, 

linear, slightly flattened, strigulose, glabrescent, 

with a curved beak at the apex, dehiscent by valves 

that twist back in a spiral. Seeds 8-10, oblong, 

2.5-3 mm long, reddish brown. 





pastures, forest edges, and roadsides, at lower and 

middle elevations. Also on Mona, Vieques, St. 

Croix, St. John, St. Thomas, Tortola, and Virgin


Gorda; throughout the Antilles and Central 

America. 

Public Forest: Guánica, Mona, and Río Abajo. 

2. Teramnus uncinatus (L.) Sw., Nov. Gen. Sp. 

Pl. 105. 1788. 

Fig. 113. F-I 

BASIONYM: Dolichos uncinatus L. 

Cresta de gallo blanca, Frijolillo peludo 


in length. Stems cylindrical, ferruginoustomentose 

or sericeous. Leaves alternate, 

trifoliolate; leaflets chartaceous, oblong, 

lanceolate, or less frequently elliptical, 5-12 × 

(0.6)0.9-3.2(6) cm, the apex acute, obtuse, or

284 


rounded, mucronate, the base obtuse or rounded, 

frequently unequal on the lateral leaflets, the 

margins entire; upper surface strigulose; lower 

surface sericeous-pubescent, with prominent 

venation, ciliate; petiolules tomentose, 2-4 mm 

long; rachis ferruginous-tomentose, sulcate, 0.2- 

2 cm long; petioles ferruginous-tomentose, 

sulcate, 3-11.5 cm long; stipules lanceolate, 

pubescent, 3-4 mm long; stipels linear, 1-2 mm 

long. Inflorescences of axillary pseudoracemes, 

10-15 cm long, with 2 or 3 flowers grouped at the 

nodes; peduncle tomentose or sericeous. Calyx 

campanulate, ca. 6 mm long, tomentose, the lobes 

linear; corolla pink or lavender, the standard ca. 

4 mm long, oblong or rounded, the wings longer 

21. VIGNA 

than the keel, the keel whitish. Legume flattened, 

linear, 4-7 × 0.3-0.4 cm, ferruginous-tomentose, 

with a curved beak at the apex, dehiscent by valves 

that twist back in a spiral. Seeds oblong, 4-5 mm 

long, reddish brown. 


to January and in fruit from October to March. 



roadsides, pastures, secondary forests, and 

thickets, at middle and lower elevations. Also in 

the Greater Antilles, tropical continental America 

and introduced in tropical Africa. 

Public Forests: Carite, Guajataca, Maricao, 


Herbaceous vines, twining, climbing, or creeping. Leaves alternate, trifoliolate; stipules of various 

forms, persistent; stipels subulate. Inflorescences of axillary or terminal pseudoracemes; bracts small. 

Calyx campanulate, bilabiate, with 5 lobes, two of which are almost completely united; corolla yellow, 

pink, or pale blue, the standard rounded, the wings unguiculate, with a narrow projection or spur at the 

base, the keel recurved, as long as the wings; stamens 10, diadelphous, the anthers of the same size; 

ovary sessile, with numerous ovules, the style curved, pubescent on the distal portion, the stigma 

lateral. Fruit an oblong or linear legume, flattened to cylindrical, dehiscent by valves that twist on 

opening; seeds flattened, quadrangular or almost reniform. A genus of about 100 species, the majority 

of tropical Africa and Asia. 

Key to the species of Vigna 

1a. Corolla uniformly brilliant yellow………………………………………………………..2 

1b. Corolla pink or violet-pink, changing to white or pale yellow with purplish lines .……..5 

2a. Standard ca. 1.5 cm long; legumes 4-7 cm long…………………………………………..3 

2b. Standard 6-10 mm long; legumes 1-4 cm long……………………………………………4 

3a. Leaflets chartaceous, ovate, acute at the apex; stipules sagittate at the base; legumes 4-7 cm long 

..............………………….....…………………………………………..………5. V. luteola 

3b. Leaflets fleshy, obovate, obtuse, or rounded at the apex; stipules truncate at the base; legumes 5-6 

cm long .........................……………………………………………………….....6. V. marina 

4a. Standard ca. 6 mm long; legume 8-15 mm long, with 1-3 seeds …………………. V. hosei 

4b. Standard ca. 1 cm long; legume 3-4 cm long, with 5-7 seeds……………....4. V. longifolia 

5a. Standard < 1 cm long; legumes ascendant …………..…………………..7. V. peduncularis 

5b. Standard 1.5-3 cm long; legumes sparse or pendulous…………………………………….6 

6a. Style spirally twisted……………………………………………………………………... 7 

6b. Style erect or sigmoid…………………………………………………………………….. 8


7a. Stipules truncate at the base; peduncle of the inflorescence as long as or shorter than the accompanying 

leaf; legume oblong, 8-9 mm wide, curved in the middle...................…………..1. V. adenantha 

7b. Stipules asymmetrical, the base auriculate on only one side; legume linear, curved in the distal 

portion, 3-4 mm wide…….............…….…………………………………..….2. V. antillana 

8a. Plant more or less glabrous; legume 10-40 cm long …………………..….8. V. unguiculata 

8b. Plant ferruginous-pilose; legume 7-11 cm long ………………………….......9. V. vexillata 

1. Vigna adenantha (G. Meyer) Marechal, 

Mascherpe & Stanier, Taxon 27: 202. 1978. 

Fig.114. E-H 

BASIONYM: Phaseolus adenanthus G. Meyer 

Habichuela cimarrona 

Slightly woody vine, twining, climbing, 

attainig 3-5 m in length. Stems slender, almost 

cylindrical, glabrous or pilose. Leaves alternate, 

trifoliolate; leaflets chartaceous, with three 

principal veins from the base, the apex acute, 

obtuse, or acuminate, the upper surface glabrous, 

sometimes sparsely appressed-pubescent, the 

lower surface pale green, appressed-pubescent, 

especially on the prominent veins, the margins 

deeply sinuate; terminal leaflet lanceolate or ovate, 

3-9.5(15.5) × 1.5-5.3(9.6) cm, the base cuneate 

or obtuse; lateral leaflets markedly asymmetrical, 

with one side elliptical and the other ovate or 

lanceolate, the base cuneate-obtuse; petiolules 

pubescent, 2-5 mm long; rachis 1-2 cm long, 

striate, pilose; petioles canaliculate, striate, pilose 

or glabrous, 3-9.5 cm long; stipules triangular, 

3-4 mm long, glabrous, conspicuously veined, 

truncate at the base; stipels oblong, ca. 2 mm long. 

Inflorescence of axillary pseudoracemes, 4-12 cm 

long, with 2 flowers per node; peduncle appressedpubescent 

or glabrous; pedicels 1.6-2 mm long, 

glabrous; bracts in pairs, lanceolate, deciduous. 

Calyx green, glabrous, campanulate, 5-7.5 mm 

long, with 3 lanceolate lobes, longer than the two 

central ones, which are rounded and fused; corolla 

pale violet; standard reniform, retuse at the apex, 

2.2-3 cm long and up to 2.5 cm wide, pale violet, 

whitish toward the margins, with the vexillar 

callus yellow; wings 2.2-2.5 cm long, obovate, 

twisted, enclosing the keel, pale violet, dark violet 

toward the apex, or sometimes pale yellow; keel 

up to 3 cm wide, pale violet to whitish, spirally 

twisted at the apex (½-1½ turns). Legume oblong, 

curved, flattened, the area of the seeds slightly 

prominent, 7-11 cm long, 8-9 mm wide, 

puberulent, scabrous along the margins, the apex 

acute, recurved. Seeds 11-16, almost circular, 

flattened, ca. 6 mm wide, dark brown. 


December to April. 

Status: Probably native, rather common. 

Distribution: Moist disturbed areas at lower 

and middle elevations. Native to tropical America, 

cultivated in Africa and Asia. 

Public Forests: El Yunque, Guajataca, and Río 

Abajo. 

2. Vigna antillana (Urb.) Fawcett & Rendle, Fl. 

Jam. 4(2): 69. 1920. 

Fig. 114. A-D 

BASIONYM: Phaseolus antillanus Urb. 

Herbaceous vine, twining, climbing, attainig 

3-5 m in length. Stems cylindrical, glabrous, 

reddish on the young portions. Leaves alternate, 

trifoliolate; leaflets chartaceous, 3-9 × 2-7 cm, 

the apex acuminate, the margins entire, slightly 

revolute; upper surface glabrous or puberulent, 

with slightly prominent venation; lower surface 

glabrous or puberulent, with prominent venation; 

terminal leaflet broadly ovate or deltate, the base 

cuneate or truncate, with three main veins; lateral 

leaflets markedly asymmetrical, with one side 

oblong and the other broadly ovate, the base 

truncate; petiolules puberulent, 4-5 mm long; 

rachis 2-2.5 cm long, marginate, puberulent or 

glabrous; petioles marginate, puberulent or 

glabrous, 3-10 cm long; stipules lanceolate, 6-8 

mm long, asymmetrical at the base, one side 

auriculate; stipels subulate, minute, persistent. 

Inflorescence of axillary pseudoracemes, up to 30 

cm long, with 2 flowers per node; peduncle 

glabrous or puberulent; pedicels 2-3 mm long, 

glabrous; bracts minute, persistent. Calyx green, 

puberulent, campanulate, 4-5 mm long, the lobes 

more or less of the same length, 3 of them 

triangular, the 2 central ones rounded and fused;

286 


corolla pale violet to violet-pink; standard 

obovate, rounded at the apex, 1.5-1.8 cm long, 

pale violet; wings longer than the standard, 

oblong, unguiculate, pale violet; keel as long as 

the standard, twisted in the form of an ‘s’, pale 

violet. Legume linear, curved toward the apex, 

flattened, the area of the seeds slightly prominent, 

6-14 cm long, 3-4 mm wide, puberulent, the apex 

terminating in a beak, dehiscent by valves that 

twist back on opening. Seeds 15-20, oblong, 

slightly flattened, 2-3 mm long, brown or 

blackish. 




Distribution: In more or less dry and disturbed 

areas, at lower elevations. Also on Caja de Muerto, 

Vieques, St. Croix, St. John, and Tortola; 

throughout the Antilles. 


Commentary: This species was treated 

erroneously as Phaseolus peduncularis in Flora 

of St. John, U.S. Virgin Islands (Acevedo-Rdgz. 

et al., 1996). 

3. Vigna hosei (Craib) Backer ex Heyne in Nutt., 

Pl. Nederl. Ind. ed. 2(2): 840. 1927. 

Fig. 114. I-L 

BASIONYM: Dolichos hosei Craib 


attainig 2-3 m in length, often forming a dense 

cover on the ground. Stems cylindrical, ca. 1 mm 

in diameter, greenish, ferruginous-pilose, much 

branched and producing roots in the area of the 

nodes. Leaves alternate, trifoliolate; leaflets 

chartaceous, with three main veins from the base, 

2-4(5) × 1.5-2 cm, the apex obtuse or acute, both 

surfaces strigulose, the margins slightly undulate; 

terminal leaflet elliptical, obovate, or lanceolate, 

the base obtuse or rounded; lateral leaflets 

asymmetrical, with one side wider than the other, 

the base obtuse-rounded; petiolules pilose, ca. 1 

mm long; rachis 4-10 mm long, striate, pilose; 

petioles 2-5 cm long, striate, pilose; stipules 

lanceolate, ca. 3 mm long, hastate at the base; 

stipels oblong or subulate, minute, persistent. 

Inflorescence of axillary pseudoracemes, 2-3(7) 

cm long, with 1-4 flowers in the distal portion 

(1-2 flowers per node); pedicels ca. 2 mm long. 

Calyx campanulate, ca. 1.5 mm long, the lobes 

ca. 0.7 mm long, triangular-subulate, ciliate; 

corolla pale yellow; standard broadly orbicular, 

7-9 × 8-11 mm, the wings obovate, ca. 8 mm long, 

the keel ca. 7 mm long; ovary with two ovules, 

the stigma lobate. Legume oblong, almost 

cylindrical, 8-15 mm long, ca. 4 mm wide, 

pubescent, slightly compressed between the seeds. 

Seeds 1-2, oblong, ca. 5 mm long, dark reddish 

brown. 


Status: Exotic, naturalized, locally common. 

Distribution: In moist disturbed areas. Native 

to Borneo and Java, introduced in the Antilles. 

Public Forests: Carite, El Yunque, and Río 

Abajo. 

4. Vigna longifolia (Benth.) Verdc., Kew Bull. 

24: 541. 1970. 

Fig. 115. A-C 

BASIONYM: Phaseolus longifolius Benth. 

SYNONYMS: Phaseolus lanceolatus Bello 

Phaseolus ovatus Benth. 

Habichuela cimarrona 


attainig 2-3 m in length. Stems very slender, 0.5- 

1.5 mm in diameter, greenish, ferruginous-pilose 

or glabrous when mature, sometimes with roots 

in the area of the nodes. Leaves alternate, 

trifoliolate; leaflets chartaceous, with three main 

veins from the base, 2-6.2 × 0.4-2.2(3) cm, 

lanceolate or linear, the apex acute, the base 

rounded, the margins slightly undulate; both 

surfaces more or less strigulose; lateral leaflets 

slightly asymmetrical at the base; petiolules 

broadened, pilose, 1-2 mm long; rachis 2-6 mm 

long, canaliculate, pilose; petioles 2-5 cm long, 

canaliculate, pilose; stipules peltate in the center, 

glabrous, persistent, both ends oblong, up to 4 

mm long, of the same length or one longer than 

the other; stipels oblong, ca. 0.5 mm long, 

persistent. Inflorescence of axillary 

pseudoracemes, 4-12(22) cm long, the flowers 2 

per node, which are found in the distal portion of 

the inflorescence; pedicels ca. 2 mm long; 

bracteoles in pairs, at the base of the calyx, oblong, 

sparsely pilose, 3-5 mm long, deciduous. Calyx 

asymmetrically campanulate, 1.5-3 mm long, the 

lobes short, obtuse, similar; corolla pale yellow; 

standard obovate, ca. 8 mm long, the wings and 

the keel more or less of the same length as the


288 


standard. Legume oblong, compressed, 2.5-4 cm 

long, 6-7 mm wide, appressed-pubescent. Seeds 

5-7, asymmetrically ellipsoid, ca. 5 mm long, 

reddish brown, shiny, the hilum whitish, ca. 2.5 

mm long. 


March and in fruit in May. 

Status: Apparently exotic, naturalized, locally 

common. Although the existence of this species 

in Puerto Rico has been since the end of the 

nineteenth century, it seems to have been 

introduced, because this constitutes the only 

record for the species in the Antilles. 

Distribution: Known only on the margins of 

the Tortuguero Lagoon, on a substrate of white 

sands. Native to tropical continental America, 

from Panama to Argentina and Uruguay. 

Public Forest: Tortuguero. 

5. Vigna luteola (Jacq.) Benth. in Mart., Fl. Bras. 

15(1): 194. 1859. 

Fig. 115. D-H 

BASIONYM: Dolichos luteolus Jacq. 

SYNONYMS: Dolichos repens L. 

Vigna repens (L.) Kuntze, non Baker 

Frijol silvestre, Goat wiss 


twining, attainig 10 m in length. Stems branched 

from the base, slender, cylindrical, ferruginouspilose. 

Leaves alternate, trifoliolate; leaflets 

chartaceous, ovate or lanceolate, 2.5-7.5 × 1-5 cm; 

upper surface glabrous, with slightly prominent 

venation; lower surface sparsely appressedpubescent, 

especially on the prominent veins, the 

apex acute, obtuse, or less frequently acuminate, 

the base cuneate to rounded, slightly asymmetrical 

on the lateral leaflets, the margins entire, ciliate; 

petiolules 2-3 mm long, pubescent; rachis 9-13 

mm long, marginate or narrowly winged; petioles 

2-9 cm long, marginate or narrowly winged; 

stipels oblong, 2-2.5 mm long; stipules lanceolate, 

ca. 3 mm long, auriculate at the base. 

Pseudoracemes axillary, up to 30 cm long, the 

flowers 1-2 per node, which are found in the distal 

portion of the inflorescence; pedicels 4-5 mm 

long, erect, with the calyx in a horizontal position 

or reflexed. Calyx green, campanulate, 4-5 mm 

long, the sepals unequal, the longer (3 mm) ones 

subulate, the shorter (1.5-2 mm) ones deltate, 

ciliate. Corolla brilliant yellow; standard rounded, 

1.5-2 cm long; the wings and the keel as long as 

the standard. Legume pointing downward, linear, 

almost cylindrical, slightly curved, with the 

margins slightly sinuate between the seeds, 4-7 

cm long, ca. 5 mm wide, glabrous or pubescent, 

blackish on drying. Seeds oblong or rectangular, 

ca. 5 mm long, reddish brown. 

Phenology: Flowering almost throughout the 

year. 



zone, also in disturbed areas such as roadsides 

and pastures. Also on Culebra, Mona, Vieques, 

St. Croix, St. John, St. Thomas, and Tortola; 

throughout tropical and subtropical America, 

especially along the coasts of the littoral zone, 

also in Africa and Asia. 


Guánica, El Yunque, Mona, Piñones, Río Abajo, 

Toro Negro, and Tortuguero. 

6. Vigna marina (Burm.) Merr., Interpret. 

Rhumph. Amboin. 285. 1917. 

Fig. 115. I-J 

SYNONYM: Vigna retusa (E. Meyer) Walp. 

Herbaceous vine, creeping, twining, attainig 

2 m or more in length. Stems slender, cylindrical, 

striate, pubescent, glabrous when mature. Leaves 

alternate, trifoliolate; leaflets slightly fleshy, 

obovate, elliptical, or almost rounded, 2.5-7 × 1.5- 

4 cm; upper surface appressed-pubescent or 

glabrous; lower surface appressed-pubescent or 

glabrous, the venation prominent, the apex obtuse 

or rounded, the base cuneate or rounded, slightly 

asymmetrical on the lateral leaflets, the margins 

entire; petiolules 1-2 mm long, glabrous or 

sericeous; rachis ca. 11 mm long, sulcate, 

glabrescent; petioles ca. 3 cm long, sulcate, 

glabrescent; stipels oblong, ca. 1 mm long; 

stipules lanceolate, 2-3 mm long, truncate at the 

base. Pseudoracemes axillary, 6-8(20) cm long, 

the flowers few per node, which are found in the 

distal 1/2 or 1/3 of the inflorescence; pedicels ca. 

4 mm long. Calyx green, campanulate, ca. 4 mm 

long, the sepals unequal, ciliate. Corolla yellow; 

standard obovate, ca.1.5 cm long, emarginate at 

the apex; the wings and the keel as long as the 

standard. Legume almost cylindrical, with the 

margins slightly sinuate between the seeds, 5-6 

cm long, ca. 5 mm wide, glabrous, blackish on


290 


drying. Seeds rectangular, ca. 4.5 mm long, black, 

shiny. 

Phenology: Collected in fruit in March. 



from Boca de Cangrejos, apparently along the 

sandy coast. Predominant along the tropical coasts 

of the Old World. In the New World, it is known 

from a few collections from Panama and Brazil. 

Commentary: The application of the name V. 

marina is rather confused, and needs an 

explanatory note. This epithet was originally 

published by Burman based on the work of 

Rumphius (Index Alt. Herb. Amboin. 5: 391, tab. 

141 fig. 2, 1750). As a result, the name Vigna 

marina is typified by the illustration presented in 

that work. Nevertheless, the examination of this 

illustration reveals that in reality it represents the 

species that today bears the name V. luteola. Due 

to the code of nomenclature’s principle of priority, 

the name V. luteola should be replaced with V. 

marina. Nevertheless, the name Vigna luteola 

could be maintained if the name V. marina was 

conserved with a different type, so as to preserve 

the traditional use of both names. In this treatment 

I use the traditional nomenclature in anticipation 

of a proposal to conserve the name V. marina with 

a different type. As a result, V. retusa becomes a 

synonym of V. marina. 

7. Vigna peduncularis (Kunth) Fawcett & 

Rendle, Fl. Jam. 4(2): 68. 1920. 

Fig. 116. A-D 

BASIONYM: Phaseolus peduncularis Kunth 

Herbaceous vine, creeping or climbing 

twining, attainig 2 m in length. Stems branched 

from the base, slender, almost cylindrical, 

puberulent, usually producing roots in the area of 

the nodes. Leaves alternate, trifoliolate; leaflets 

chartaceous, ovate-deltate or lanceolate, 3-8 × 1.5- 

5 cm, both surfaces glabrous, the venation slightly 

prominent on the lower surface, the apex acute, 

acuminate, or less frequently obtuse, the base 

cuneate, the margins entire, slightly ciliate; 

terminal leaflet larger than the lateral ones; lateral 

leaflets asymmetrical at the base, one side obtuse, 

the other cuneate or truncate; petiolules ca. 3 mm 

long, puberulent; rachis 5-10 mm long, striate; 

petioles 2-7 cm long, striate, slender, glabrous; 

stipels obovate, ca. 1 mm long; stipules oblong or 

lanceolate, 2-2.5 mm long. Pseudoracemes 

axillary, up to 26 cm long, the flowers 2 per node, 

in the distal portion of the inflorescence; 

peduncles thick, glabrous or puberulent; pedicels 

ca. 1 mm long, erect, with the calyx in an 

ascending position. Calyx green, campanulate, 

puberulent, 2.2-2.5 mm long, the sepals more or 

less equal, broadly deltate, except for the abaxial 

one, which is narrower and slightly longer; corolla 

pale pink, standard obovate, unguiculate, ca. 8 

mm long, the wings and the keel as long as the 

standard, the wings oblong, sinuate, unguiculate, 

the keel spirally twisted. Legumes ascendent, 

linear, flattened, straight, with the margins 

slightly undulate, 4-5.5 cm long, ca. 3 mm wide, 

puberulent, punctate. Seeds oblong, ca. 3.5 mm 

long, light brown with blackish spots, the hilum 

white, ca. 2 mm long. 


November and January. 


Distribution: In disturbed places like roadsides 

and pastures. Also in the Antilles, Central 

America, and South America. 


8. Vigna unguiculata (L.) Walp., Repert. Bot. 

Syst. 1: 779. 1842. 

Fig. 116. E-F 

BASIONYM: Dolichos unguiculatus L. 

Frijol, Cowpea 

Herbaceous vine, climbing and twining, 

attainig 3-5 m in length. Stems up to 5 mm in 

diameter, cylindrical, glabrous, wine-colored 

when young. Leaves alternate, trifoliolate; leaflets 

chartaceous, 5-14.6 × 3-10.5 cm, both surfaces 

glabrous, with slightly prominent venation, the 

apex obtuse or acute, mucronate, the margins 

sinuate to lobate; terminal leaflet rhombic-ovate 

or rhombic-lanceolate, the base cuneate, with the 

point rounded or sometimes subcordiform; lateral 

leaflets markedly asymmetrical, one side oblong, 

the other ovate or lanceolate, the base cuneate or 

truncate; petiolules 4-5 mm long, puberulent, 

winged; rachis 0.5-3.5 cm long, canaliculate, 

marginate or winged; petioles 1-10 cm long, 

canaliculate, marginate or winged, glabrous, 

thick, broadened at the base; stipels oblong or 

obovate, ca. 2 mm long; stipules peltate,


constricted in the zone of insertion, the distal 

portion lanceolate, 6-10 mm long, the basal 

portion 2-3 mm long. Pseudoracemes axillary, up 

to 40 cm long, the flowers 2 per node, in the distal 

portion of the inflorescence; peduncles thick, 

glabrous; pedicels ca. 2 mm long, erect, with the 

calyx in an ascending position. Calyx green, 

campanulate, glabrous, ca. 8 mm long, the sepals 

slightly unequal, subulate, as long as the tubular 

portion. Corolla pale violet, white, greenish, or 

yellowish; standard rounded, 1.5-3 cm long, 

unguiculate, with the base auriculate; the wings 

and the keel shorter than the standard, the keel 

not spirally twisted. Legumes ascendent or 

horizontal, linear, slightly compressed, straight, 

15-20 × ca. 1 cm, glabrous, the valves with 

partitions between the seeds. Seeds 10-15 per fruit, 

oblong, 9-10 mm long, reddish brown or white 

with a black spot, the hilum white, ca. 2 mm long. 




uncommon. 


and pastures. Cultivated throughout the tropics 

for food; apparently native to the tropics of the 

Old World. 


9. Vigna vexillata (L.) A. Rich. in Sagra, Hist. 

Fis. Polit. Nat. Cuba 11: 191. 1845. 

Fig. 116. G-K 

BASIONYM: Phaseolus vexillatus L. 

Frijol cimarrón 


twining, attainig 4-5 m in length. Stems slender, 

green, ferruginous-pilose. Leaves alternate, 

trifoliolate; leaflets chartaceous, 3.5-11.5 × 1.5- 

5.2 cm, lanceolate, the upper surface strigulose, 


strigulose or strigose, with prominent venation, 

the apex acute or acuminate, the base obtuse or 

rounded on the terminal leaflet, unequal (obtusetruncate) 

on the lateral ones, the margins revolute, 

ciliate; petiolules 2-4 mm long, pilose, marginate; 

rachis canaliculate, pilose, 3-20 mm long; petioles 

canaliculate, pilose, swollen at the base, 1.6-10 

cm long; stipels subulate, 2-3 mm long; stipules 

lanceolate, 6-9 mm long, sagittate at the base. 

Pseudoracemes axillary, 10-31 cm long, the 

flowers 2 per node, in the distal portion of the 

inflorescence; peduncle thick, sparsely pilose, 

glabrescent; pedicels 2.5-3 mm long, erect; 

bracteoles subulate, ca. 5 mm long, pubescent. 

Calyx green, campanulate, puberulent, 12-15 mm 

long, the sepals slightly unequal, subulate, as long 

as the tubular portion of the calyx. Corolla pale 

violet or pink; standard ca. 2 cm long, the wings 

1 cm wide, with the base mulberry-colored. Fruit 

a linear legume, cylindrical, 7-11 cm long, with 

the apex acute, slightly recurved, ferruginouspilose-pubescent. 

Seeds oblong, black or gray. 



Status: Apparently native, relatively common. 


and pastures. Also in the Antilles, Central 

America, South America, Africa, and tropical 

Asia. 



Vigna juruana (Harms) Verdc. was cited for 

Puerto Rico by Liogier (1998) based on the 

erroneous identification of a specimen of Vigna 

luteola (Alain 27987). 

References: Burkart, A. 1971. El género Galactia (Legum.-Phaseoleae) en Sudamérica, con especial 

referencia a la Argentina y países vecinos. Darwiniana 16: 662-802. Fantz, P.R. 1990. Clitoria 

(Leguminosae) Antillarum. Moscosoa 6: 152-166. Fantz, P.R. 1996. Resegregation of Barbieria from 

Clitoria (Leguminosae: Phaseoleae: Clitoriinae). Sida 17: 55-68. Grear, J.W. 1978. A revision of 

Rhynchosia (Leguminosae-Faboideae). Mem. New York Bot. Gard. 31: 1-168. Maxwell, R. H. 1999. 

Dioclea. pp. 310-315. In: J. A. Steyermark et al. (eds.) Flora of the Venezuelan Guayana. Vol. 5.

292 



24c. Subfamily MIMOSOIDEAE 


1a. Inflorescences racemes of spikes, 4-5 cm long…………………..…...………….. 2. Entada 

1b. Inflorescences racemes of heads or solitary heads …………………………………….…. 2 

2a.Corolla white, cream-colored, or yellowish ..………….………………………..............… 1. Acacia 

2b. Corolla pink ……………………………………...…………………………...........….... 3. Mimosa 

1. ACACIA 

Trees, shrubs, or lianas; stems unarmed or spiny. Leaves alternate, bipinnate; pinnae opposite; 

leaflets small, numerous, opposite; petioles and rachis usually with nectariferous glands; stipules usually 

spiny, persistent; stipels absent. Flowers bisexual, produced in heads or spikes grouped in axillary or 

terminal racemes or panicles; bracts small. Calyx campanulate, of 5 united or free sepals; corolla 

yellow or white, tubular, with 4-5 lobes; stamens numerous, exserted, the filaments free or united at the 

base; ovary superior, sessile or stipitate, with several ovules, the style filiform, curved, pubescent on 

the distal portion. Fruit an oblong or linear legume, straight or curved, flattened to almost cylindrical, 

dehiscent or indehiscent; seeds of diverse forms, usually flattened. A genus of about 500 species, the 

majority of tropical America, tropical Africa, and Australia. 

Key to the species of Acacia 

1a. Plant spiny; stems obtusely quadrangular; legume 1.5-2 cm wide …………................. 1. A. retusa 

1b. Plant not spiny; stems more or less cylindrical; legume 2.5-3 cm wide ….............. 2. A. vogeliana 

1. Acacia retusa (Jacq.) R. A. Howard, J. Arnold 

Arbor. 54: 459. 1973. 

Fig. 117. A-F 

BASIONYM: Mimosa retusa Jacq. 

SYNONYMS: Mimosa paniculata West ex Vahl, non Willd. 

Acacia westiana DC. 

Senegalia westiana (DC.) Britton & Rose 

Acacia riparia sensu Britton & Wilson, non Kunth 

Zarza brava, Katch and keep 

Liana that supports itself by means of recurved 

spines and attains 15 m in length. Stems 

cylindrical, quadrangular and sulcate when 

mature, attaining 10 cm in diameter, sometimes 

splitting into 4 longitudinal sections; branches 

angular, with numerous recurved spines. Leaves 

bipinnate, 6-13 cm long; pinnae 4-8 pairs; leaflets 

16-32 pairs per pinna, 3-5 mm long, oblong, 

chartaceous, the apex obtuse, the base cordateobtuse, 

unequal, the margins entire; primary 

rachis flattened, usually aculeate, pubescent, with 

two stipitate annular glands, one in a distal 

position and another in a basal position; secondary 

rachis cylindrical, unarmed; petioles ca. 2 cm 

long, with the base slightly swollen; stipules 

minute, early deciduous. Inflorescences of 

terminal panicles of globose heads, 1-1.5 cm in 

diameter. Calyx campanulate, 0.6-1 mm long, 

glabrous; corolla white, campanulate, 2-2.5 mm 

long; stamens white, exserted. Legume flattened, 

6-15 × 1.5-2 cm, glabrous or tomentose, 

chartaceous, the margin thickened, unarmed, 

dehiscent along both sutures. 

Phenology: Flowering from July to December 

and fruiting from October to March. 


Distribution: In dry disturbed areas, at middle 

and lower elevations. Also on Culebra, Vieques, 

St. Thomas, St. John, Tortola, and Virgin Gorda; 

also in Hispaniola, the Lesser Antilles, and 

northern South America.

294 


Public Forests: Cambalache, Ceiba, Guánica, Río 

Abajo, and Susúa. 

2. Acacia vogeliana Steud., Nomencl. Bot. ed. 2, 

1: 9. 1840. 

Fig. 117. G-J 

SYNONYMS: Lysiloma vogeliana (Steud.) Stehlé 

Acacia ambigua Vogel, non Hoffmans. 

Lysiloma ambigua (Vogel) Urb. 

Liana, not spiny, that climbs by means of 

twining lateral branches and attains 10 m in 

length. Stems cylindrical, striate, puberulent. 

Leaves bipinnate, 6-13 cm long, puberulent; 

pinnae in 5 pairs, pulvinate, with a pair of linear 

glands above the pulvinus; rachis sulcate, 

puberulent, with an annular gland between the 

two pairs of basal and distal pinnae; leaflets 15- 

16 pairs per pinna, 5-10 mm long, oblonglanceolate, 

chartaceous, the apex obtuse, 

mucronulate, the base obtuse-rounded, unequal, 

the margins entire; both surfaces glabrous or 

minutely puberulent; stipules minute, early 

2. ENTADA 

deciduous. Inflorescences of terminal panicles of 

heads, ca. 1 cm in diameter. Calyx campanulate, 

0.6-1 mm long, strigose; corolla white or creamcolored, 

campanulate, strigose, 2-2.5 mm long; 

stamens white, exserted. Legumes 6-16 × 2.5-3 

cm, oblong, flattened, chartaceous, glabrous, 

stipitate, with the seeds prominent, the margins 

thickened and slightly undulate. Seeds 4-6 mm 

long, elliptical, light brown. 


in fruit in January. 



in Puerto Rico (at the entrance to La Parguera) 

and another on St. John (entrance toward 

Bordeaux); also in Hispaniola and probably on 

Martinique. 

Commentary: The collections (Acevedo 1, 3, 

3242 and 5047) by which this species is known 

from Puerto Rico and the Virgin Islands differ 

from the type collection (from Hispaniola) in 

having a climbing habit. The collections known 

from Hispaniola are all of trees or shrubs. 

Lianas; stems unarmed or spiny. Leaves alternate, bipinnate, sometimes with a terminal tendril; 

pinnae opposite; leaflets numerous, opposite; petioles with or without nectariferous glands; stipules 

small, setaceous; stipels minute. Flowers bisexual, produced on spikes arranged in paniculate 

inflorescences; bracts minute. Calyx campanulate or crateriform, of 5 small sepals; corolla of free 

petals; stamens 10, exserted, the filaments free at the base, the anthers with a deciduous apical gland; 

ovary superior, subsessile, with numerous ovules, the style filiform. Fruit an oblong legume, straight or 

recurved, flattened, articulate between the seeds, the margins thickened, separating from the valves 

when ripe; seeds flattened, circular or reniform. A genus of about 15 species, of pantropical distribution. 

1. Entada polystachya (L.) DC. var. polyphylla 

(Benth.) Barneby, Brittonia 48: 175. 1996. 

Fig. 118. A-D 

SYNONYM: Entada polyphylla Benth. 

Liana, not spiny, that climbs by means of foliar 

tendrils (homologous to the two distal pinnae) and 

attains 10 m in length. Stems more or less 

cylindrical, striate. Leaves bipinnate, 15-25 cm 

long; pinnae in 4-6 pairs; rachis slightly flattened 

along the upper surface, puberulent; secondary 

rachis carinate, puberulent, with a pair of stipels 

below the basal pinnae; leaflets 10-20 pairs per 

pinna, 10-16 mm long, oblong, coriaceous, the 

apex obtuse, rounded, or emarginate, the base 

asymmetrical, one side cuneate, the other truncate, 

the margins entire, revolute; upper surface dark 

green, glabrous or puberulent; lower surface pale 

green, strigulose, with the midvein prominent; 

petiole 3-3.5 cm long, with a long pulvinulus at 

the base; stipules minute, early deciduous. 

Inflorescences terminal, forming a raceme of 

cylindrical spikes, 4-5 cm long. Calyx crateriform, 

ca. 0.5 mm long, puberulent; petals oblong, green 

or reddish brown, 1.2-1.5 mm long; stamens 

white, slightly longer than the petals. Legumes


296 


25-35 × 6.5-7.5 cm, oblong, flattened, coriaceous, 

glabrous, stipitate, with the margins thickened and 

conspicuously undulate, separating into segments 

that contain a single seed. Seeds 4-6 mm long, 

elliptical, light brown. 


3. MIMOSA 

Status: Apparently native, extremely rare or 

extirpated. 


(Sintenis 1240), from the base of Monte de Mula 

in Fajardo. Also on Dominica and in northern 


Erect or scandent herbs or shrubs; stems spiny. Leaves alternate, bipinnate; pinnae opposite; leaflets 

small, numerous, opposite; petioles and rachis without nectariferous glands; stipules minute, deciduous 

or persistent; stipels minute or absent. Flowers bisexual or staminate, produced in heads, solitary or 

grouped in axillary or terminal racemes; bracts small, usually shorter than the corolla. Calyx minute, 

hypocrateriform, crowned by 5 minute sepals; corolla yellow or pink, infundibuliform, with 3-6 lobes 

(petals); stamens as numerous as or double the number of petals, exserted, the filaments free; ovary 

superior, stipitate, with several ovules, the style filiform, the stigma punctiform. Fruit an oblong legume, 

chartaceous, flattened, indehiscent or dehiscent by the walls that separate from the thickened margin, 

usually spiny; seeds flattened, lenticular or ovate. A genus of about 450 species, of pantropical 

distribution, the majority of the Neotropics. 

Key to the species of Mimosa 

1a. Fruits obtusely quadrangular, with numerous longitudinal ribs …..…......... 4. M. quadrivalvis 

1b. Fruits flattened, without longitudinal ribs …………………………….…………………........ 2 

2a. Leaves with a single pair of pinnae; leaflets 3-4 pairs, elliptic-oblong-lanceolate, acute at the 

apex...........…………………………………………………………………..…….......… 1. M. casta 

2b. Leaves with 2-7 pairs of pinnae; leaflets 3-20 pairs, obliquely obovate or oblong, rounded or obtuse 

at the apex..............................…………………………………………………………………… 3 

3a. Leaflets 15-20 pairs, 3-4 mm long, oblong; heads ca. 6 mm in diameter................3. M. diplotricha 

3b. Leaflets 3-8 pairs, 1-1.5 cm long, obliquely obovate; heads 1.3-1.7 cm in diameter......2. M. ceratonia 

1. Mimosa casta L., Sp. Pl. 518. 1753. 

Fig. 118. E-H 

Zarza 

Woody vine, climbing, scarcely branched, that 

supports itself on other plants by means of spines 

which are borne along the length of its stems and 

petioles, and attainig 1-2 m in length. Stems 

angular, glabrous, with numerous recurved spines. 

Leaves alternate, bipinnate, with a single pair of 

pinnae; leaflets 3-4 pairs, elliptical to oblonglanceolate, 

1-3 × 0.5-1.5 cm, the apex acute, the 

base asymmetrical, one side attenuate, the other 

rounded, the margins ciliate; upper surface 

glabrous; lower surface sparsely sericeous, with 

prominent venation; petioles up to 10 cm long, 

with numerous recurved spines, pulvinate at the 

base; stipules subulate, ca. 5 mm long. 

Inflorescences of globose heads, axillary; peduncle 

5-15 mm long. Calyx ca. 0.2 mm long, glabrous; 

corolla white, infundibuliform 1-2 mm long, 

glabrous, with 4 lobes; stamens 4, the filaments 

free, ca. 1 cm long. Legume flattened, oblong, 3- 

4 × ca. 1.2 cm, glabrous, with 4-5 articulations, 

the margin with recurved spines. 


January and February. 

Status: Exotic, adventive, apparently a recent 

introduction, locally common. 

Distribution: On roadsides and in pastures at 

lower and middle elevations. Also in the Lesser 

Antilles and from Panama to Brazil.


298 


2. Mimosa ceratonia L, Sp. Pl. 523. 1753. 

Fig. 119. A-D 

SYNONYM: Lomoplis ceratonia (L.) Raf. 

Zarza, Lamedora, Black amaret, Amarat 

steckel, Amarat 

Woody vine, climbing, much branched, that 

supports itself on other plants by means of spines 

that are borne along the length of its stem and 

leaf axes, attaining 2-6 m in length. Stems green 

or with a reddish tinge, obtusely quadrangular, 

glabrous, striate, with numerous recurved spines, 

becoming almost cylindrical and grayish when 

mature. Leaves alternate, 7-15 cm long, bipinnate; 

pinnae 4 or 5, pairs opposite; rachis sulcate, with 

numerous recurved spines; leaflets opposite, 3-8 

pairs per pinna, 1-1.5 cm long, obliquely obovate 

to almost rounded, chartaceous, the apex rounded, 

the base rounded-obtuse, asymmetrical, the 

margins slightly revolute, sometimes ciliate; upper 

surface dark green, dull, glabrous; lower surface 

pale green, dull, with prominent venation; stipules 

8-10 mm long, lanceolate. Heads 1.3-1.7 cm in 

diameter, in terminal racemes; peduncles 1-2 cm 

long, with numerous recurved spines. Calyx 0.7- 

1 mm long, glabrous; corolla pink, 

infundibuliform, ca. 2 mm long, with three petals 

or lobes; filaments white, 4-6 mm long. Legumes 

flattened, straight or slightly curved, 4-6 × 1.5- 

1.7 cm, coriaceous, dehiscent by the walls that 

separate from the thickened margin, covered with 

recurved spines. Seeds 7-8 mm long, oblongelliptical, 

flattened, dark brown. 

Phenology: Flowering from June to January 

and fruiting from December to March. 



lower and middle elevations. Also on Vieques, St. 


Gorda; Hispaniola and the Lesser Antilles as far 

as St. Vincent, reported for Venezuela. 

Public Forests: Cambalache, Carite, El 

Yunque, Guajataca, Maricao, Río Abajo, Susúa, 


3. Mimosa diplotricha C. Wright ex Sauv., Anal. 

Acad. Ci. Habana 5: 405. 1868. 

Fig. 119. E-H 

Scandent shrub, much branched from the base, 

that supports itself on other plants by means of 

spines that are borne along the length of its stem 

and leaf axes, attainig 1.5-2 m in length. Stems 

pentagonal, pilose, with numerous recurved spines 

along the angles. Leaves alternate, 5-13 cm long, 

bipinnate; pinnae 6 or 7, pairs opposite; leaflets 

opposite, 18-15 pairs per 

pinna, 2.5-4.2 mm long, oblong, chartaceous, 

the apex obtuse, the base asymmetrically obtusetruncate; 

both surfaces strigulose; lower surface 

pale green, dull, with the midvein sunken; rachis 

very slender, pilose, with some recurved spines; 

stipules subulate, ca. 4.5 mm long; stipels linear, 

ca. 1.5 mm long. Heads 7-10 mm in diameter, 

axillary; peduncles with numerous recurved 

spines, 6-15 mm long. Calyx ca. 0.2 mm long, 

glabrous; corolla pink, infundibuliform, ca. 1.2 

mm long, with 4(5) lobes; filaments pink, 3-5 mm 

long. Legumes usually 10 or more per head, 

oblong or linear, flattened, straight or slightly 

curved, 10-24 × 3-4.3 mm, chartaceous, covered 

with spines, especially along the margin. Seeds 

3-8, 2.3-3 mm long, flattened, ovate or rhombic, 

dark brown. 


Status: Exotic, adventive, of recent 

introduction, uncommon. 


lower and middle elevations. Also in Cuba, 

Hispaniola; Mexico, Central and South America. 


4. Mimosa quadrivalvis L. var. urbaniana 

Barneby, Mem. New York Bot. Gard. 65: 297. 

1991. 

Fig. 119. I 

SYNONYMS: Schrankia portoricensis Urb. 

Morongia portoricensis (Urb.) Britton 

Leptoglottis portoricensis (Urb.) Britton & Rose 

Schrankia leptocarpa sensu Bello, non DC. 

Schrankia distachya sensu A. Stahl, non DC. 

Zarzaparilla 

Erect or clambering shrub, scarcely branched, 

that supports itself on other plants by means of 

spines that are borne along the length of its stem 

and petioles, attaining 1-2 m in length. Stems 

slender, angular or ribbed, pilose, glabrescent, 

with numerous recurved spines along the angles. 

Leaves alternate, bipinnate, with 4-5 pairs of 

pinnae; leaflets 8-10 pairs, oblong, 2.5-4 mm 

long, the apex obtuse, the base very asymmetrical, 

truncate-obtuse, the margins ciliate; both surfaces


300 


glabrous or puberulent with numerous minute 

circular scales; rachis very slender, spiny; stipules 

subulate, 3-4 mm long; stipels spiniform, ca. 1 

mm long. Heads globose, axillary, solitary; 

peduncle 4-13(20) mm long. Calyx 0.3-0.5 mm 

long; corolla campanulate, pink, 2-3 mm long, 

with 5-6 lobes; stamens exserted, pink, ca. 6 mm 

long. Legumes 1-7 per head, erect, linear or 

oblong, quadrangular, 2-14 cm long, 2.5-6.5 mm 

25. Family LAURACEAE 

1. CASSYTHA 

26. Family MALPIGHIACEAE 


wide, pilose, with 4-5 longitudinal ribs, which 

have recurved spines. 

Phenology: Collected in fruit during 

December. 

Status: Variety endemic to Puerto Rico, 

uncommon. 

Distribution: Known from Piedras Chiquitas, 

Coamo. 

Twining vines, herbaceous, parasitic. Leaves reduced to minute scales. Flowers bisexual, sessile, 

in axillary spikes . Tepals 6, in two unequal series; fertile stamens 9, in three whorls; ovary superior, 

globose, minute. Fruit a fleshy berry. A cosmopolitan genus of 20 species. 

1. Cassytha filiformis L., Sp. Pl. 35. 1753. 

Fig. 120. A-F 

SYNONYM: Cassytha americana Nees 

Fideillo, Bejuco dorado, Cabellos de ángel, 

Fideos, Tente en el aire, Yellow dodder, Yellow 

love 

Herbaceous vine, twining, parasitic, that 

adheres to the host plant by means of small 

haustoria, attainig 1-5 m in length. Stems 

yellowish green or yellow, flexible, ca. 2 mm in 

diameter, cylindrical, profusely branched. Leaves 

1-2 mm long, lanceolate, sessile, almost 

imperceptible or absent. Inflorescences of short 

spikes with few flowers. Tepals white, ovate, ca. 

2 mm wide. Fruit globose, light green or white, 

5-7 mm in diameter. 




Distribution: In disturbed areas at low 

elevation, predominantly near the littoral zone. 

Also on Cayo Diablo, Isla Piñeiros, Mona, 


and Virgin Gorda; a cosmopolitan species. 

Public Forests: Guánica, Mona, Piñones, and 

Tortuguero. 

1a. Mericarp with a single dorsal wing well developed (lateral wings not developed)………..2 

1b. Mericarp with 4 wings (two pairs of lateral wings, the dorsal wing not developed) 

…………………………………………………………...………….….......…3.Tetrapterys 

2a. Stamens 10, subequal, fertile; styles obtuse at the apex …..……………..…1. Heteropterys 

2b. Stamens 10, unequal, 4-6 fertile, with the anthers thicker; styles with the apex broadened 

..………………………………………………….…..…………………….2. Stigmaphyllon


1. HETEROPTERYS 

Twining vines, shrubs, or small trees, usually with malpighiaceous hairs. Leaves simple, entire, 

opposite, usually with glandular depressions on the blade; petioles usually with a pair of stipitate 

glands; stipules minute or absent. Flowers bisexual, in axillary or terminal umbels, corymbs, or 

pseudoracemes. Calyx of 5 basally connate sepals, 4 of which have a pair of glands on the outer 

surface, or less frequently eglandular. Petals yellow, pink, or white; stamens 10, unequal, fertile, the 

filaments connate at the base, the anthers ellipsoid to oblong; ovary superior, of 3 connate carpels, the 

styles 3, free, erect, thick. Fruit a schizocarp of 3 samaras with a dorsal wing. A genus of approximately 

125 species, the great majority distributed throughout the Neotropics. 

Key to the species of Heteropterys 

1a. Petals pink; leaves 1.5-2 cm long ………………………..…..…………...........… 2. H. purpurea 

1b. Petals yellow; leaves 7-15 cm long…………………………………………………..............…… 2 

2a. Stems not lenticellate; leaves with the lower surface densely appressed-ferruginous-pubescent; 

petioles with two glands in the middle ..……………………..................… 3. H. wydleriana 

2b. Stems densely lenticellate; leaves with the lower surface glabrous; petioles without glands 

…………………………………………………………………………...........….. 1. H. laurifolia

302 


1. Heteropterys laurifolia (L.) A. Juss., Ann. Sci. 

Nat. Bot. Sér. 2. 13: 176. 1840. 

Fig. 121. A-F 

BASIONYM: Banisteria laurifolia L. 

Bejuco de buey, Paralejo velludo, 

Dragon wiss 

Vine or sometimes scandent shrub, woody, 

twining, attainig 5-10 m in length. Stems flexible, 

cylindrical, lenticellate, attaining 3.5 cm in 

diameter. Leaves opposite, 5-15(23.5) × 2-9(13.5) 

cm, coriaceous, glabrous, oblong or elliptical, the 

apex acuminate or rounded, the base acute, the 

margins undulate or entire; upper surface shiny, 

with the midvein prominent, yellowish green; 

lower surface yellowish green, with the midvein 

prominent; petioles 1-1.2 cm long, without 

glands. Inflorescences of panicles, terminal or 

axillary, the axes densely ferruginous-pubescent; 

pedicels ca. 7 mm, ferruginous-pubescent, 

articulated above the base; bracts ca. 2 mm long, 

at the base of the pedicel; bracteoles in pairs, 

minute, inserted on the articulation of the pedicel. 

Calyx green, ferruginous-pubescent, of 5 oblong 

sepals, ca. 4 mm long, 4 of them with a pair of 

glands at the base of the outer surface, which are 

1.2-2 mm long. Petals 5, yellow, unguiculate, 4-7 

mm long. Mericarps 3, ferruginous-tomentose in 

the seminiferous portion, glabrous toward the 

wing, 2-4 cm long, with a dorsal wing, curved, 

ascendent or divaricate. 





Distribution: In secondary forests or pastures 

or along roads or rivers. Also throughout the 

Antilles, from Central America to Bolivia. 

Reported for St. Croix by Niedenzu (according to 

Britton and Wilson, 1924). 

Public Forests: Ceiba, El Yunque, Guajataca, 

Guánica, Maricao, Río Abajo, Susúa, and Toro 

Negro. 

2. Heteropterys purpurea (L.) Kunth in Humb., 


5: 164. 1821 [1822]. 

Fig. 121. G-K 

BASIONYM: Banisteria purpurea L. 

Bejuco de paralejo, Bejuco de toro, 

Red wiss 


Stems slender, cylindrical, copper-colored, 

puberulent, lenticellate, with numerous lateral 

branches. Leaves opposite, chartaceous, 1.5-2 × 

1-1.5 cm, oblong, elliptical, or ovate, the apex 

obtuse or rounded, sometimes mucronate, the base 

obtuse or rounded, the margins revolute; upper 

surface shiny, glabrous or puberulent; lower 

surface yellowish green, dull, puberulent; petioles 

3-10 mm long, puberulent, with a pair of glands 

in the middle; stipules absent. Inflorescence of 

axillary racemes; bracts minute, without glands; 

pedicels 5-6 mm long, slender, ferruginouspubescent; 

articulated near the base. Calyx of 5 

sepals, elliptical or ovate, ca. 3 mm long, 

appressed-pubescent, 4 of which have a pair of 

glands at the base of the outer surface. Petals 5, 

pink, broadly ovate, unguiculate, 4.5-5 mm long, 

the apex rounded. Mericarps 1.5-3 cm long, with 

an ascendent dorsal wing. 


fruiting from December to March. 


Distribution: In dry forests and disturbed areas 

at lower elevations, usually along the coast. Also 

on Culebra, Vieques, St. Croix, St. John, St. 

Thomas, Tortola, and Virgin Gorda; the United 

States (Florida), the Antilles, Trinidad, and 

Venezuela. 

Public Forests: Cambalache, Ceiba, Guánica, 

Guajataca, and Susúa. 

3. Heteropterys wydleriana A. Juss., Ann. Sci. 

Nat. Bot. Sér. 2. 13: 275. 1840. 

Fig. 121. L-S 

SYNONYMS: Banisteria wydleriana (A. Juss.) C.B. Robinson 

Heteropterys bellonis Urb. 

Ahorca caballo 


Stems cylindrical, ferruginous-pubescent when 

young, attaining 5 cm in diameter, the bark 

grayish. Leaves opposite, coriaceous, 6.5-15 × 3.6- 

7 cm, oblong or elliptical, the apex acute or obtuse 

and sometimes mucronate, the base obtuse or 

rounded; upper surface dark green, slightly shiny, 

puberulent, the midvein ferruginous-pubescent; 

lower surface densely ferruginous-sericeoustomentose, 

the venation prominent; petioles ca. 1 

cm long, with a pair of glands at the middle, 

ferruginous-pubescent. Inflorescences of axillary


304 


dichasial cymes; bracts minute, triangular; 

pedicels 9-11 mm long, ferruginous-strigose, 

articulated in the middle. Calyx ferruginousstrigose, 

the sepals 5, triangular, 3-4 mm long, 

four of them with a pair of glands at the base of 

the outer surface. Petals 5, yellow, oblong, 

unguiculate, ca. 4 mm long. Mericarps (1-)3, 

ferruginous-tomentose, 4-5 cm long, with a distal 

dorsal wing, ascendent. 

2. STIGMAPHYLLON 


January and fruiting in January and February. 

Status: Endemic to Puerto Rico, locally 

common. 

Distribution: In secondary and mature forests 

in the zone of mogotes and serpentine. 

Public Forests: Maricao, Río Abajo, and Susúa. 

Twining vines, usually with simple or malpighiaceous trichomes. Leaves simple, entire, opposite, 

usually with a pair of glands at the base or on the petiole; stipules minute, eglandular. Flowers bisexual, 

in axillary or terminal umbels, corymbs, or pseudoracemes; bracts eglandular; bracteoles glandular or 

eglandular. Calyx of 5 basally connate sepals, 4 of which have a pair of glands on the outer surface. 

Petals yellow, sometimes with a reddish tinge, unguiculate, the posterior petal usually with the 

unguiculate base larger and the limb smaller than the other petals; stamens 10, unequal, the filaments 

connate at the base, the anthers glabrous or pubescent; ovary superior, of 3 connate carpels, the styles 

3, free, with the stigmatic portion on the inner (adaxial) portion. Fruit a schizocarp of 3 samaras with 

a dorsal wing, distal, on a pyramidal receptacle. A genus of approximately 100 species, distributed 

from Mexico to Argentina, including the Antilles. 

Key to the species of Stigmaphyllon 

1a. Mericarp with a dorsal wing, divaricate (projecting horizontally); leaves coriaceous….... .2 

1b. Mericarp with a distal wing, ascendent or rudimentary; leaves chartaceous……………… 3 

2a. Young stems strigulose; leaves glabrescent on both surfaces, the tertiary veins inconspicuous; samaras 

glabrescent ........................................……...…………………..…….……2. S. emarginatum 

2b. Young stems tomentose; leaves densely appressed-pubescent on the lower surface, the tertiary veins 

conspicuously reticulate; samaras tomentose ..…..………............. 3. S. floribundum 

3a. Leaves acute, obtuse, or sometimes apiculate at the apex; wing of the mericarp rudimentary, reduced 

to an apical crest of the seminiferous locule, 4-9 mm long..…………........… 1 S. bannisterioides 

3b. Leaves acuminate at the apex; wing of the mericarp distal, ascendent, well developed, 2.6-3.7 cm 

long ..................…………………………………………………………....……….. 4. S. puberum 

1. Stigmaphyllon bannisterioides (L.) C. 

Anderson, Taxon 41: 328. 1992. 

Fig. 122. A-C 

BASIONYM: Malpighia bannisterioides L. 

SYNONYMS: Banisteria ovata Cav. 

Stigmaphyllon ovatum (Cav.) Niedenzu 

Brachypterys ovata (Cav.) Small 


Stems cylindrical, sericeous when young. Leaves 

opposite, ovate, elliptical, or lanceolate, 4-12 × 

1.5-5.5 cm, chartaceous, the apex acute, obtuse, 

or sometimes apiculate, the base attenuate, obtuse, 

or truncate, with a pair of annular glands in the 

area adjacent to the petiole, the margins entire; 

upper surface glabrous or glabrescent; lower 

surface sparsely sericeous; petiole slightly sulcate, 

sericeous, 0.4-1.8 cm long; stipules minute, 

triangular. Inflorescences of umbels with (3)4(6) 

flowers, axillary, solitary or in dichasial cymes, 

with a pair of broadly ovate leaves of reduced size 

at the base; peduncles 0.2-2.5 mm long; pedicels 

15-30 mm long, cylindrical; bracteoles 0.8-1.6 

mm long, ovate, elliptical, or triangular. Calyx of 

5 ovate sepals, sericeous, ca. 3 mm long, four of


them with a pair of elongate glands outside at the 

base. Petals yellow, rounded or obovate, unequal, 

9-12 mm long, unguiculate; stamens unequal; 

styles equal or almost equal. Mericarps with the 

wing reduced to an apical crest, 4-9 mm long, 

the seminiferous portion with 4-6 ribs. 

Phenology: Flowering from December to July 

and fruiting from February to July. 

Status: Native, quite rare. 

Distribution: Known only from the Naguabo 

area on the edge of a mangrove between the mouth 

of the Santiago River and the Antón Ruiz River. 

Also on Vieques; the Greater Antilles, Martinique, 

St. Lucia, and Barbados, from the Caribbean coast 

of Mexico to the north coast of Brazil, east of the 

mouth of the Amazon, and from the Pacific coast 

of Colombia to northern Peru. 

2. Stigmaphyllon emarginatum (Cav.) A. Juss., 

Ann. Sci. Nat. Bot., II, 13: 290. 1840. 

Fig. 122. D-G 

BASIONYM: Banisteria emarginata Cav. 

SYNONYMS: Stigmaphyllon periplocifolium (Desf.) A. 

Juss. 

Stigmaphyllon lingulatum (Poir.) Small 

Bejuco de San Juan 


length. Stem up to 8 cm in diameter; much 

branched from the base; bark very rough, reddish 

brown. Branches copper-colored, glabrous, more 

or less cylindrical, lenticellate, the young portions 

strigulose. Leaves opposite, coriaceous, 

lanceolate, ovate, oblong, linear, or less frequently 

rounded, 2.5-10(13) × 1-5.5(10.5) cm, sparsely 

appressed-pubescent when young, glabrescent, the 

apex acute or obtuse, emarginate-mucronate, the 

base truncate or cordiform, the margins entire or 

undulate; upper surface shiny, with the reticulate 

venation prominent; lower surface pale green, 

dull, with the midvein prominent; petioles 

yellowish, flattened on the upper surface, 0.2-2 

cm long, with two discoid glands in the area 

adjacent to the blade; stipules triangular, minute. 

Inflorescences of axillary umbels, corymbs, or 

pseudoracemes, with 15-35 flowers; peduncles 

pubescent, 0.2-2.5 cm long; pedicels 3-23 mm 

long, articulated above the base; bracts minute, 

eglandular. Calyx of 5 ovate-triangular sepals, 

4(5) of which have a pair of glands outside at the 

base. Petals 5, yellow, rounded, unguiculate, 11- 

15 mm long; stamens unequal, glabrous or 

puberulent; anterior style shorter; stigmas green. 

Mericarps 1.6-2.2 cm long, with an asymmetrical 

dorsal wing, curved on the upper margin and 

crenate on the lower margin. 


fruiting from January to August. 


Distribution: In disturbed areas and dry forests 

near the littoral zone. Also on Caja de Muerto, 

Culebra, Desecheo, Icacos, Mona, Vieques, 


Virgin Gorda; Jamaica, Hispaniola, and the Lesser 

Antilles southward to Martinique. 

Public Forests: Ceiba, Guánica, Maricao, 

Mona, Piñones, and Susúa. 

3. Stigmaphyllon floribundum (DC.) C. 

Anderson, Syst. Bot. 11: 128. 1986. 

Fig. 122. H-M 

BASIONYM: Banisteria floribunda DC. 

SYNONYMS: Stigmaphyllon tomentosum (Desf.) Niedenzu 

Tetrapterys paniculata sensu A. Stahl 

Heteropterys nitida var. obtusifolia Kuntze 

Bejuco de toro, Bejuco de menta, 

Bull wiss, Red wiss 


Stems much branched from the base, cylindrical, 

attaining 2-3 cm in diameter; bark rough, reddish 

brown. Branches lenticellate, cylindrical, 

tomentose. Leaves opposite, yellowish or reddish 

when young, elliptical, oblong, linear, lanceolate, 

or rounded, 4-18 × 2.5-15.5 cm, coriaceous, the 

apex obtuse or truncate, emarginate-mucronate, 

the base obtuse, truncate to cordiform, the margins 

entire, revolute, ciliate; upper surface dark green, 

glabrous or sparsely pubescent; lower surface pale 

green, dull, densely appressed-pubescent or 

sericeous, the venation prominent; petioles 

cylindrical, ferruginous-pubescent, 1-2.5 cm long, 

with a pair of prominent discoid glands in the 

area adjacent to the blade; stipules triangular, 

minute. Inflorescences of terminal panicles, up 

to 22 cm long, composed of umbels or 

pseudoracemes, with 20-30(50) flowers; 

peduncles strigose; pedicels strigose, 1-3 cm long. 

Calyx of 5 rounded sepals, 2-2.7 mm long, 

strigose, four of them with a pair of glands outside 

at the base. Petals yellow, rounded, 11-15 mm 

long, unguiculate; stamens unequal; anterior style

306 


shorter, the stigmas green. Mericarps 1.8-3.2 cm 

long, strigose, especially the seminiferous portion, 

with a rudimentary apical wing and a well 

developed dorsal wing, divaricate, more or less 

oblong, with the upper margin curved. 


fruiting sporadically. 

Status: Endemic to Puerto Rico and the Virgin 

Islands, rather common. 

Distribution: In coastal forests and disturbed 

areas of the littoral zone. Also on Mona, St John, 

and Virgin Gorda. 

Public Forests: Cambalache, El Yunque, 

Guánica, Guajataca, Guilarte, Maricao, Mona, 

Río Abajo, Susúa, and Tortuguero. 

4. Stigmaphyllon puberum (Rich.) A. Juss., Ann. 

Sci. Nat. Bot., II, 13: 289. 1840. 

Fig. 123. A-G 

BASIONYM: Banisteria pubera Rich. 


length. Stems cylindrical, sericeous when young. 

Leaves opposite, lanceolate, elliptical, ovate, or 

less frequently rounded, (6.8)8.2-13 × 3-6 cm, the 

apex acuminate, the base obtuse, slightly decurrent 

on the petiole, with a pair of prominent discoid 

glands, 1-1.8 mm in diameter, in the area near 

the petiole, the margins revolute, with scattered 

minute sessile glands; upper surface glabrous or 

sericeous on the midvein; lower surface sericeous 

or sparsely sericeous, the venation prominent; 

petiole canaliculate, sericeous, 1.5-4 cm long; 

stipules triangular, minute, deciduous. 

Inflorescences of axillary umbels, with 8-15 

flowers, grouped in dichasial cymes or thyrses; 

peduncles 1.5-4.8 mm long; pedicels cylindrical, 

3. TETRAPTERYS 

2.5-7.5 mm long; bracts triangular, 1.5-2.2 mm 

long, with a pair of glands at the base. Calyx of 5 

ovate sepals, sericeous, four of them with a pair 

of prominent oblong glands outside at the base. 

Petals yellow, sometimes with a reddish tonality, 

rounded, 7.5-13 mm long, unguiculate, the 

posterior petal 5-7 mm long; stamens unequal; 

anterior style longer, the stigmas with leafy 

appendages. Mericarps 2.6-3.7 cm long, glabrous, 

with an ascendent wing, in a distal position. 

Phenology: Known from Puerto Rico only in 

sterile condition. 


Distribution: Known from few localities in 

eastern Puerto Rico, from Fajardo to Naguabo. 

Also in Jamaica, the Dominican Republic, the 

Lesser Antilles, Central America, and northern 



Stigmaphyllon cordifolium Niedenzu (= S. 

diversifolium (Kunth) A. Juss.) was reported for 

St. Thomas by Niedenzu (1928), in error. His 

report is based on two collections, i.e., Read 88 

and Eggers 390, which have been identified as S. 

emargiatum in the recent monograph of C. 

Anderson (1997). 

Cultivated Species: 

Stigmaphyllon retusum Griseb. has been 

cultivated at the Agricultural Experiment Station 

in Mayagüez, from seeds from Vera Cruz, Mexico. 

The available collection of this species dates from 

1952 and includes flowers and fruits. Since this 

species can reproduce itself sexually in Puerto 

Rico, it is possible that it may manage to establish 

itself on the island some day. 

Twining vines, or less frequently shrubs. Leaves simple, entire, opposite; petioles sometimes with 

glands; stipules minute, simple, interpetiolar. Flowers bisexual, in umbels or corymbs, grouped in 

axillary or terminal cymes or panicles; bracteoles usually on the distal portion of the peduncle. Calyx 

of 5 basally connate sepals, ovate or oblong, usually 4 of which are biglandular or less frequently all 

biglandular or eglandular. Petals yellow, unguiculate, glabrous, rounded, ovate, or elliptical, with the 

margins entire, denticulate, or crenate; stamens 10, slightly unequal, the filaments connate at the base, 

the anthers glabrous or pubescent; ovary superior, of 3 connate carpels, the styles 3, free, erect, usually 

one of them more slender, the stigmatic surface on the inner (adaxial) portion. Fruit a schizocarp with 

2-3 samaroid mericarps with a narrow dorsal wing and 2 pairs of lateral wings forming an ‘x’. A genus 

of 90 species, distributed from Mexico to Argentina, including the Antilles.


308 


1. Tetrapterys inaequalis Cav., Diss. 433. 1790. 

Fig. 123. H-N 

SYNONYMS: Tetrapterys paniculata Bello 

Tetrapterys citrifolia sensu Alain, non (Sw.) Pers. 

Bejuco de paralejo, Bejuco de sopla 


length. Stems cream-colored, cylindrical, 

sericeous-tomentose when young, glabrous when 

mature. Leaves opposite, chartaceous, elliptical 

oblong or ovate, 4-15 × 3-6(8.9) cm, the apex 

obtuse, acute, or acuminate, the base obtuse, 

rounded, or subcordiform, sometimes unequal, the 

margins undulate or entire; upper surface dark 

green, slightly shiny, glabrescent; lower surface 

dull, glabrous or puberulent, with prominent 

venation and usually sericeous-tomentose; petioles 

0.6-1.5 cm long, sericeous-tomentose or glabrous, 

slightly sulcate; stipules oblong to lanceolate, 5- 

6 mm long, sericeous-tomentose, early deciduous, 

leaving an annular scar. Inflorescences formed by 

short branches with numerous umbels of 4 

flowers; peduncles sericeous-tomentose; pedicels 

sericeous-tomentose, 6-9 mm long, articulated in 

the middle; bracts minute. Calyx of 5 green sepals, 

ovate, 2-3 mm long, 4 of which have a pair of 

glands outside at the base. Petals yellow, 5-7 mm 

long, the base unguiculate, orange, the limb ovate. 

Mericarps 2.5-3 cm long, with two pairs of wings, 

the distal ones ca. 20 × 5 mm, the basal ones ca. 

10 × 2.5 mm. 


September to November and in fruit from 



Distribution: In disturbed areas like along 

roads and rivers and in secondary forests, at lower 

and middle elevations, found along the northern 

limestone zone. Also on Vieques; reported for St. 

Thomas and St. Croix (Britton and Wilson, 1924) 

but not confirmed; the Lesser Antilles. 

Public Forest: Cambalache and Maricao. 

Note: Sterile specimens of this species have 

been identified by Woodbury and treated by 

Liogier (1982, 1997) as Sabicea cinerea Aubl. 

and in the Spanish edition of this book as Sabicea 

sp. a. This error was apparently started by Werham 

(1914) who cited S. cinerea as occurring in Puerto 

Rico in his Monograph of the genus Sabicea. This 

mistake is corrected in the present edition. 

Excluded Genus: 

The report of Banisteriopsis lucida (A. Rich.) 

Small for Puerto Rico is based on a bibliographic 

error. Originally this species was reported as 

doubtful for Puerto Rico by Otto Kuntze (1891), 

as Stigmaphyllon nigrescens (A. Juss.) Kuntze, 

based on the transfer of Banisteria nigrescens A. 

Juss. Some years later, Niedenzu (1899) placed 

Stigmaphyllon nigrescens as a synonym of 

Banisteria lucida A. Rich., and cited Kuntze as 

evidence of the presence of this species in Puerto 

Rico. In that publication, Niedenzu disassociated 

the name S. nigrescens from its basionym, 

presenting it as a new species of Kuntze. Britton 

and Wilson (1924), adopted the opinion of 

Niedenzu and cited the presence of Banisteriopsis 

lucida in Puerto Rico based on a supposed 

specimen of Kuntze. This error acquired a new 

dimension when Bates, in 1982, cited S. 

nigrescens as a synonym of Banisteriopsis lucida 

and adjudicated a specimen of Otto Kuntze (s.n.) 

collected in Puerto Rico from a cultivated plant 

and deposited in the herbarium of the New York 

Botanical Garden. The examination of the 

publication of Kuntze clearly reveals that the 

author did not collect this species and that his 

report for Puerto Rico was characterized as 

doubtful. I therefore conclude that the report of 

this species for Puerto Rico is based on the 

perpetuation of an error and, for this reason, it is 

excluded from the flora of Puerto Rico in the 

present work. 

References: Anderson, C. 1997. Monograph of Stigmaphyllon (Malpighiaceae). Syst. Bot. Mon. 

51: 1-313. Bates, B. 1982. Banisteriopsis, Diplopterys (Malpighiaceae). Fl. Neotropica 30: 1-237. 

Kuntze, O. 1891. Revisio Generum Plantarum. Paris. Niedenzu, F. 1899. De genere Stigmatophyllo. 

Brunsberg. Niedenzu, F. 1928. Malpighiaceae. In: A. Engler (ed.). Das Pflanzenreich. IV. 141. Leipzig.


310 


27. Family MARCGRAVIACEAE 

1. MARCGRAVIA 

Clambering shrubs, usually epiphytic, with dimorphic branches and leaves; juvenile branches 

climbing by means of aerial roots, the internodes short, with leaves of reduced size; adult branches 

usually pendulous, with leaves 3-4 times as long as the juvenile ones. Leaves simple, entire, alternate, 

usually with dark glandular dots on the blade or the margin; petioles usually short; stipules absent. 

Flowers bisexual, in terminal umbelliform racemes; the peripheral flowers fertile, the central ones 

sterile, usually replaced by elongate nectaries with a cavity or pocket in which abundant nectar 

accumulates; bracteoles appressed to the calyx, similar in form to the sepals. Calyx of 4 decussate 

sepals; corolla in the form of a calyptra, deciduous; stamens 10 or more numerous, the filaments free; 

ovary superior, with 4-12 carpels, with numerous ovules. Fruit a berry, globose or ovoid, with numerous 

seeds surrounded by a reddish aril. A genus of 45 species, distributed throughout the Neotropics. 

Key to the species of Marcgravia 

1a. Adult leaves with glandular dots only along the margin; inflorescences with 6-25 flowers; pedicels 

4-7 cm long, usually densely lenticellate; flower straight; calyx 8-9 mm wide; nectaries light brown, 

4-5 mm wide ...........................…..………………………………...….1. M. rectiflora 

1b. Adult leaves with glandular dots distributed along the lower surface; inflorescences with ca. 50 

flowers; pedicels ca. 3 cm long, not lenticellate or with sparse lenticels; flower geniculate; calyx 3- 

6 mm wide; nectaries reddish, 1-1.3 cm wide ..........................................………..2. M. sintenisii 

1. Marcgravia rectiflora Triana & Planchon, 

Ann. Sci. Nat. Sér. 4, 17: 364. 1862. 

Fig. 2F; 124. A-G 

SYNONYMS: Marcgravia umbellata Griseb. 

Marcgravia brittoniana Alain 

Bejuco de rana, Pegapalma, Bejuco de paloma, 

Bejuco de lira, Bejuco de sapo 

Liana with two different morphological 

phases. Juvenile plants and branches climbing, 

ca. 1 m in length, sparsely branched, climbing by 

means of adventitious roots; stems flattened; 

leaves 1.5-2.5 cm long, ovate to oblong, retuse at 

the apex, very closely spaced. Adult plant 

attaining 10 m or more in length and with 

scandent or pendulous branches. Stems 

cylindrical, lenticellate, woody, flattened when 

adult, bilobate in cross section, attaining 2.5 cm 

wide. Leaves oblong or elliptical, 7-12.5 × 2.8-4 

cm, coriaceous, glabrous, the apex acuminate, the 

base acute, obtuse, or less frequently rounded; the 

margins entire, with minute black dots; upper 

surface dark green, slightly shiny; lower surface 

yellowish green, with the midvein prominent, the 

secondary veins inconspicuous; petioles 2-3 mm 

long. Inflorescence an umbelliform raceme with 

6-25 flowers and 1-3 nectaries in a central 

position, 3-3.5 × 0.4-0.5 cm; pedicels straight, 

3.5-6 cm long, lenticellate; bracts 2, rounded, 

smaller than the sepals. Calyx green, 5-7 mm 

long, of 4 rounded sepals; corolla green, forming 

a calyptra, ca. 1 cm wide, early deciduous; stamens 

numerous, the filaments white; ovary green. Fruit 

globose, fleshy, light brown, lenticellate-rugose, 

1.5-2 cm in diameter. Seeds numerous, minute, 

surrounded by a red pulp. 

Phenology: Flowering from March to 

December and fruiting from February to May. 


Distribution: In forests at lower and 

moderately high elevations. Also on Tortola; Cuba 





2. Marcgravia sintenisii Urb., Jahrb. Bot. Gart. 

Berlin 4: 245. 1886. 

Fig. 124. H-K 

Pegapalma 

Plant with the juvenile phase morphologically 

different from the adult phase. Juvenile plant ca. 

1 m in length, sparsely branched, climbing by 

means of adventitious roots; stems flattened; 

leaves 1.5-2 cm long, cordiform to almost 

rounded, with the margin crenate, very closely 

spaced. Adult plant developing into a woody vine, 

with scandent and pendulous branches, attainig 

5-10 m in length. Stems cylindrical, 2-5 cm in 

diameter. Leaves alternate, changing from reddish 

to green when mature, 4.5-9 × 1.3-4 cm, 

coriaceous, elliptical, lanceolate, or oblong, the 

base obtuse to rounded, the apex acute or obtuse, 

the margins revolute; upper surface dark green, 

dull; lower surface light green, dull, with 

numerous scattered black dots, the midvein 

prominent; petioles 4-5 mm long. Inflorescence 

an umbelliform raceme with about 50 flowers and 

7-9 reddish nectaries in a central position, 1.8-2 

× 1-1.2 cm; pedicels geniculate, 1.5-3.5 cm long, 

with few lenticels; bracts minute, opposite, not 

near the flower. Calyx reddish, of 4 rounded 

sepals, ca. 2 mm long; corolla calyptrate, 3-5 mm 

long, reddish or green with an orange tinge, 

deciduous; stamens 10-11; ovary light orange, the 

stigma green. Fruit globose, reddish, ca. 1 cm in 

diameter, fleshy. Seeds numerous, ca. 1 mm long, 

pink or green. 

Phenology: Flowering and fruiting from April 

to December. 


Distribution: In moist and wet forests at upper 

elevations. 



Reference: Dressler, S. 1997. Lectotypification of Marcgravia rectiflora (Marcgraviaceae). Taxon 

46: 109-110. 

28. Family MENISPERMACEAE 


1a. Slightly woody vine; leaves broadly ovate to rounded, with the lower surface pubescent and the 

base peltate; anthers sessile; fruits globose, red-orange, pilose ..…..1. Cissampelos 

1b. Liana; leaves ovate to broadly ovate, with the lower surface glabrous and the base not peltate; 

anthers on long filaments; fruits obovoid, purple ………………….2. Hyperbaena 

1. CISSAMPELOS 

Herbaceous vines, twining, dioecious, usually with simple trichomes. Leaves alternate, rounded to 

ovate, peltate or cordiform at the base, palmately veined; petioles elongate; stipules absent. Flowers 

minute, unisexual, pedicellate, in axillary inflorescences. Staminate flowers actinomorphic, in corymbs; 

sepals 4, free; corolla hypocrateriform; stamens 4, connate into a short tube, with sessile anthers. 

Pistillate flowers zygomorphic, in elongate cymes with foliaceous bracts; calyx and corolla of a single 

sepal and a single petal toward the same side of the flower; ovary superior, sessile, unilocular, with a 

single basal ovule, the stigma lobate. Fruit a globose drupe, fleshy, with the endocarp woody and 

verrucose; seed horseshoe-shaped. A genus of 19 species, of pantropical distribution.

312 



1. Cissampelos pareira L., Sp. Pl. 1031. 1753. 

Fig. 125. A-J 

Bejuco de mona, Pareira, 

Oreja de ratón, Velvet leaf 

Dioecious vine, slightly woody, twining, 

attainig 3-6 m in length. Stems slender, 

cylindrical, glabrous or pilose. Leaves alternate, 

chartaceous, broadly ovate to almost circular, 4- 

12 × 4-10 cm, the apex rounded, obtuse, 

emarginate, usually mucronate, the base 

cordiform, subtruncate, usually peltate; upper 

surface glabrous, yellowish green or grayish, dull; 

lower surface pale green, almost whitish, sparsely 

or densely pubescent, with the venation palmate, 

prominent, the margin revolute, crenate; petioles 

2. HYPERBAENA 

cylindrical, pubescent, 2-7 cm long, with the base 

swollen. Inflorescences of axillary cymes or 

cauliflorous; bracts of the pistillate inflorescences 

foliaceous, ovate, pilose. Flowers 1-1.5 mm long, 

greenish. Calyx pilose. Drupe globose, redorange, 

4-5 mm in diameter, fleshy, partially 

covered by a bract. Seed horseshoe-shaped. 



Distribution: In moist and disturbed areas 

throughout Puerto Rico. Also on Culebra, Vieques, 

St. Croix, St. John, St. Thomas, and Tortola; of 

pantropical distribution. 

Public Forests: Cambalache, Carite, Ceiba, El 

Yunque, Guajataca, Guilarte, Maricao, Río Abajo, 

Susúa, Toro Negro, and Tortuguero. 

Twining vines, scandent shrubs, or trees, glabrous or with a pubescence of simple hairs. Leaves 

alternate, coriaceous, rounded to ovate, peltate or cordiform at the base, pinnately veined; petioles 

elongate; stipules absent. Flowers minute, green or white, unisexual, pedicellate, in axillary spikes or 

panicles; sepals 6, unequal. Petals free, 6 or less frequently fewer or these completely absent. Staminate 

flowers with 6 stamens, longer than the petals; pistillate flowers with 6 staminodia; ovary superior, 

with 2-5 free carpels, ovoid to globose, the stigma sessile. Fruit a drupaceous monocarp with a single 

seed inside; endocarp woody; seed horseshoe-shaped. A genus of 21 species, distributed throughout the 

Antilles and from Mexico to Argentina. 

Key to the species of Hyperbaena 

1a. Leaves ovate or broadly ovate; monocarps 1.0-1.5 cm long …………….................. 1. H. domingensis 

1b. Leaves elliptical; monocarps 2.3-2.7 cm long ....................................... 2 H. laurifolia 

1. Hyperbaena domingensis (DC.) Benth., J. 

Proc. Linn. Soc., Bot. 5, Suppl. 2: 50. 1861. 

Fig. 1F; 126. A-H 

BASIONYM: Cocculus domingensis DC. 

Twining liana attainig 15 m in length, or less 

frequently a scandent shrub with numerous lateral 

branches. Stems cylindrical, glabrous or 

puberulous; cross section with concentric rings 

of xylem. Leaves alternate, coriaceous, ovate or 

broadly ovate, 4-20 × 2.5-12 cm, with the venation 

pinnate, the apex acute or acuminate, the base 

cordiform or truncate, the margins entire; upper 

surface glabrous or puberulent on the primary and 

secondary veins; lower surface glabrous, the 

venation prominent, the tertiary venation 

reticulate; petioles 1-6 cm long, swollen at the 

apex. Inflorescences racemose, of minute white 

or greenish flowers; staminate inflorescences 

clustered; pistillate inflorescences solitary. 

Staminate flowers: calyx 0.6-0.8 mm long; petals 

ca. 1 mm long; stamens 6, ca. 1 mm long. 

Pistillate flowers slightly larger; ovary of 3 

ellipsoid carpels. Monocarps obovoid, purple, 

glabrous, 1.0-1.5 cm long. 

Phenology: Collected in flower during June, 

July, and November and in fruit during October 

and December. 

Status: Native, uncommon.

314 


Distribution: In moist forests. Also on St. John 

and St. Thomas; in the Greater and Lesser Antilles 

and from northern South America to Bolivia. 

Public Forests: El Yunque, Guajataca, Guilarte, 


2. Hyperbaena laurifolia (Poir.) Urb., Symb. 

Antill. 1: 304. 1899. 

Fig. 126. I-O 

BASIONYM: Cissampelos laurifolia Poir. 

SYNONYMS: Hyperbaena glauciramis Urb. & Ekman 

Hyperbaena pellucida Urb. & Ekman


316 

Hyperbaena apiculata Urb. & Ekman 


Erect, decumbent, or scandent shrub attainig 

5-6 m in length. Stems cylindrical, glabrous, 

striate. Leaves alternate, coriaceous, elliptical or 

less frequently elliptic-lanceolate, glabrous, 8-16 

× 2.3-5 cm, with the venation pinnate, the apex 

acute, the base obtuse, the margins entire; upper 

surface with the venation pale; lower surface 

glabrous, the primary vein prominent, the 

secondary and tertiary venation slightly 

prominent, reticulate; petioles 0.7-1.7 cm long, 

pulvinate at the apex and the base. Inflorescences 

racemose, of minute greenish flowers; the axes 

ferruginous-strigulose; staminate inflorescences 

clustered, much longer than the petioles; pistillate 

inflorescences solitary or clustered, as long as or 

shorter than the petioles. Staminate flowers: calyx 

ca. 0.5 mm long; petals ca. 1 mm long; stamens 

6, almost sessile. Monocarps circular, slightly 

compressed, glabrous, 2.3-2.7 cm long. 


to August and in fruit in March. 


Distribution: In moist forests in the zone of 

mogotes. Also on St. Thomas and Hispaniola. 


Reference: Mathias, M. E. and W. L. Theobald. 1981. A revision of the genus Hyperbaena 

(Menispermaceae). Brittonia 33: 81-104. 

29. Family MORACEAE 

1. FICUS 

Trees or shrubs, erect, scandent, or strangling, that produce abundant milky latex when wounded. 

Leaves alternate, simple or lobate, coriaceous; petioles short or elongate; stipules deciduous, elongate, 

forming a conical hood that protects the apical meristem. Flowers unisexual, minute, produced in the 

interior of an axillary inflorescence, globose or ellipsoid, formed by an enlarged receptacle (syconium); 

calyx reduced, membranaceous; corolla absent. Staminate flowers with 2 stamens; pistillate flower 

with a unilocular ovary with one apical ovule. Fruit a syconium formed by an enlarged receptacle, 

globose or ellipsoid, containing numerous achenes in the interior. A genus of about 800 species, of 

pantropical distribution. 

Key to the species of Ficus 

1a. Plants with long pendulous or scandent branches, without adventitious roots; plants with a single 

phase; syconium globose, depressed-globose, or obovoid, 6-12 mm long....................... 1. F. citrifolia 

1b. Plants ascending by adventitious roots; plants with a dimorphic juvenile phase; syconium pyriform, 

4-6 cm long ..............................………………………………………………………… 2. F. pumila 

1. Ficus citrifolia Miller, Gard. Dict. ed. 8. 1768. 

Fig. 127. A-C 

Jagüey blanco, White fig 

Terrestrial or strangling (epiphytic) tree 

attainig 10 m in height, or less frequently a liana 

attainig 10 m in length, producing abundant milky 

latex. When it grows as a liana its branches are 

pendulous or scandent, the stems cylindrical, 

attaining 10 cm in diameter. Leaves alternate, 4- 

20 × 2-12 cm, elliptical, ovate, elliptic-lanceolate, 

or less frequently oblanceolate, chartaceous or 

subcoriaceous, glabrous, the apex obtuse or 

acuminate, the base obtuse, cordiform, or truncate,


the margins entire; upper surface with slightly 

prominent venation; lower surface with the 

reticulate venation prominent; petioles 1-7 cm 

long, glabrous, canaliculate; stipules conical, 

glabrous, with a reddish tinge. Syconium globose, 

depressed-globose, or obovoid, 6-12 mm long, 

greenish yellow, with reddish spots, turning red 

when ripe; peduncles 4-8 mm long. 

Phenology: Collected in fertile condition 

during August. 


Distribution: Although this is a tree of 

widespread distribution, the lianescent form has 

only been documented for the zone of mogotes in 

Puerto Rico. The arboreal form is also found on 

Caja de Muerto, Culebra, Desecheo, Mona, 

Vieques, Anegada, Jost van Dyke, St. Croix, St. 

John, St. Thomas, Tortola, and Virgin Gorda; the 

United States (Florida), the Greater and Lesser 

Antilles. 

Public Forests: Mona and Río Abajo. 

Commentary: The lianescent form of this 

species grows on the tops and sides of mogotes 

and produces long stems that hang down and 

reach the base of the mogotes. Once the stems 

have reached the base of the mogotes, they become 

creeping or scandent. It is possible that this growth 

form results from the etiolation of the stems due 

to the low light intensity, and that it does not 

constitute a fixed characteristic for this species. 

Nevertheless, this phenomenon has been observed 

rather frequently, which justifies the inclusion of 

this species in the present work. 

30. Family NYCTAGINACEAE 


2. Ficus pumila L., Sp. Pl. 1060. 1753. 

Fig. 127. D-L 

Hiedra 

Plant with the juvenile phase morphologically 

different from the adult phase. Juvenile plant 

attaining several meters in length, much 

branched, climbing by means of adventitious 

roots; stems flattened; leaves 1.5-2.5 cm long, 

ovate to oblong, retuse at the apex, very closely 

spaced. Adult plant developing into a much 

branched liana, with adventitious roots, attainig 

10 m in length and produces abundant white latex 

when wounded. Stems flattened, striate, 

tomentose, glabrescent when mature, with short 

pendulous branches. Leaves alternate, simple, 4- 

7 × 2.2-4 cm, oblong, oblanceolate, ovate, or 

elliptical, the apex obtuse, the base subcordiform, 

the margins entire; upper surface dark green, 

slightly shiny, with the venation notably lighter; 

lower surface light green, dull, with prominent 

reticulate venation; petioles 1.3-1.6 cm long, 

flattened on the upper surface, pubescent, light 

brown; stipules oblong-lanceolate to subulate, 

persistent, 1-1.2 cm long, brown, sericeous. 

Syconium green, pyriform, up to 6 cm long, soft. 

Apparently not forming seeds in Puerto Rico. 

Phenology: Collected in ‘flower’ during 

February. 

Status: Exotic, ornamental, common. 

Distribution: A plant widely cultivated in 


Public Forests: El Yunque and Maricao. 

1a. Herbaceous plants that attain 2-3 m in length; anthocarps (fruits) with a ring of stipitate glands on 

the distal portion .................……………………………..…………………............ 1. Boerhavia 

1b. Woody plants that attain 10 m or more in length; anthocarps lacking glands or with 5 or 10 

longitudinal lines of stipitate glands…………………………………………………...............…. 2 

2a. Leaves alternate; branches cylindrical; flowers produced in threes, each of which is subtended by a 

foliaceous bract of various colors (white, pink, red, or orange); flowers bisexual, 2-2.5 cm long, 

hypocrateriform, the tube angular, compressed in the central portion, the lobes revolute; stamens 

not exposed ..............................................………..........................2. Bougainvillea

318 



2b. Leaves opposite; branches quadrangular; flowers produced in axillary cymes, the bracts minute, 

green; flowers unisexual, 2.5-3 mm long, campanulate; stamens much longer than the perianth 

.........................……………...……………………...…………………………….…3. Pisonia 

1. BOERHAVIA 

Erect or prostrate herbs, quite branched from a taproot. Leaves opposite or subopposite, with 

prominent linear raphides. Flowers minute, in axillary or terminal panicles or in pedunculate heads; 

bracts and bracteoles hyaline; calyx constricted above the ovary, the upper portion with 5 lobes; stamens 

1-6, the filaments free; ovary superior, the stigma capitate or peltate. Anthocarps dry, cuneiform or 

fusiform, usually with 5 ribs. 

1. Boerhavia scandens L., Sp. Pl. 3. 1753. 

Fig. 128. A-F 

SYNONYM: Commicarpus scandens (L.) Standl. 

Clambering herb with pseudodichotomous 

branching, attainig 1-2 m in length. Branches 

cylindrical, brittle, glabrous. Leaves opposite, in 

pairs of the same size, 2-5 × 0.8-4 cm, deltoid, 

ovate, or broadly ovate, chartaceous, the apex 

obtuse or acute, the base cordiform or truncate, 

the margins undulate and ciliate; upper surface 

glabrous; lower surface puberulent; petioles 

canaliculate, 0.5-2.7 cm long. Inflorescences 

umbelliform, axillary; peduncles glabrous, 5-12 

cm long; bracts and bracteoles minute, lanceolate, 

deciduous; pedicels unequal, 2-8 mm long. Calyx: 

2. BOUGAINVILLEA 

basal part claviform, 1-1.5 mm long, glabrous, 

green; upper part or limb infundibuliform, white 

or yellowish green, green or with a pink tinge 

inside, 2-2.5 mm long; stamens 2, white; pistil 

white. Anthocarps long-pedicellate, green, 

claviform, dry, 12-13 mm long, with a ring of 

sticky stipitate glands on the distal portion. 

Phenology: Collected in flower in January and 

September and in fruit in January. 


Distribution: In pastures on roadsides, near 

the coast. Also on Anegada, St. Croix, St. John, 

St. Thomas, Tortola, and Virgin Gorda; from the 

southern United States to Guatemala, the Antilles, 

Venezuela, and from Colombia to Peru. 


Scandent or less frequently erect shrubs or small trees, usually with axillary spines. Leaves alternate 

or subopposite, simple; petioles slender; stipules absent. Flowers bisexual, in axillary dichasial cymes, 

with 3 flowers, each of which is subtended by 1 brilliantly colored foliaceous bract. Perianth tubular, 

consisting only of the calyx, with small imbricate lobes; stamens 5-8(10), included; ovary superior, 

stipitate, fusiform, the stigma unilateral. Fruit a fusiform anthocarp with 5 longitudinal ribs, not 

glandular. A genus of about 18 species, native to South America, cultivated throughout the tropics and 

subtropics. 

Key to the species of Bouganvillea 

1a. Stems and leaves pubescent with curved and short trichomes or glabrous; floral tube with ascendent 

and curved trichomes ......................…………………………………………….….1. B. glabra 

1b. Stems and lower surface of the leaves with numerous straight and elongate trichomes (pilose); 

floral tube with numerous straight and erect trichomes ............................………….2. B. spectabilis

320 



1. Bougainvillea glabra Choisy in DC., Prodr. 

13(2): 437. 1849. 

Fig. 129. A-F 

Trinitaria 

Woody vine or shrub, clambering, attainig 15 

m in length. Stem cylindrical, attaining 5-6 cm 

in diameter; cross section of the stem with discrete 

vascular bundles within the connective tissue. 

Branches pubescent, puberulent, or glabrous, with 

straight, axillary spines. Leaves alternate or 

subopposite, 3-6 × 1.5-3 cm, chartaceous, 

elliptical, puberulent, the apex acuminate, the base 

attenuate, slightly asymmetrical, the margins 

slightly sinuate; lower surface with the midvein 

prominent; petioles slender, 4-12 mm long. 

Dichasial cymes compound, on axillary branches 

or at the end of axillary spines, the axes glabrous 

or puberulent; bracts ovate, glabrous, cordiform 

at the base, 2.5-4 cm long, of showy colors, red, 

deep pink, orange, white, or purple. Perianth 

tubular-hypocrateriform, 2.5-3 mm long, 

pubescent, with 8-10 obtuse lobes, cream-colored. 

Anthocarp fusiform, glabrous, ca. 1.5 cm long. 


Status: Exotic, very common in our gardens. 

Distribution: Much cultivated throughout 

Puerto Rico. This is the most common species of 

its genus in Puerto Rico and the Virgin Islands. It 

grows well in dry areas and requires little 

watering, for which reasons it is cultivated along 

3. PISONIA 

avenues and highways. Also throughout the 

Antilles. Native to South America, but widely 

cultivated around the globe. 

2. Bougainvillea spectabilis Willd., Sp. Pl. ed. 

2, 348. 1799. 

Fig. 129. G-H 

Trinitaria 

Woody shrub, erect or clambering, attainig 5 

m in length. Branches pilose, with straight, 

axillary, pilose spines. Leaves alternate, 2.6-2.5 

× 2.7-5 cm, chartaceous, ovate, the apex acute, 

obtuse, or acuminate, the base obtuse, rounded, 

or attenuate, slightly asymmetrical, the margins 

sinuate; lower surface tomentulose, with 

prominent pilose venation; petioles slender, 1-2.3 

cm long, pilose. Dichasial cymes compound, on 

axillary branches or at the end of axillary spines, 

the axes pilose; bracts ovate, pubescent, pilose 

along the midvein, cordiform at the base, 3.6.5.5 

cm long, red or deep pink. Perianth tubularhypocrateriform, 

2.2-3 mm long, pilose. 



Distribution: Little cultivated in Puerto Rico. 

Also throughout the Antilles. Native to South 

America, but widely cultivated on a worldwide 

level. 

Trees, shrubs, or lianas, dioecious or monoecious. Leaves opposite or subopposite, sometimes with 

axillary spines. Flowers unisexual, small, white or pale yellow, in axillary or terminal panicles, corymbs, 

or cymes; bracts and bracteoles minute, persistent. Staminate flowers campanulate to obconical; stamens 

6-8, exserted, the filaments unequal, connate at the base. Pistillate flowers tubular, 5-lobate; ovary 

superior, sessile, the style exserted, the stigma branched. Fruit a dry anthocarp, oblanceolate or ellipsoid, 

with 5 ribs that bear sticky, stipitate glands. A genus of 10-15 species, of tropical distribution. 

1. Pisonia aculeata L., Sp. Pl. 1026. 1753. 

Fig. 1A; 128. G-M 

SYNONYM: Pisonia helleri Standl. 

Escambrón, Prickly-mampoo 

Liana or woody, clambering shrub, attainig 

20 m in length. Stems more or less cylindrical, 

glabrous or minutely pubescent, grayish or cream- 

colored, lenticellate, attaining 7 cm in diameter; 

cross section of the stem with discrete vascular 

bundles within the connective tissue. Branches 

short or elongate; short branches opposite, 

divaricate, persistent, with recurved spines, 

produced on the trunk; elongate branches at the 

end of the stem, glabrescent, quadrangular, with 

recurved axillary spines. Leaves opposite on the 

elongate branches or whorled on the short lateral

322 



branches, 6-14 × 3-6 cm, chartaceous, elliptical, 

ovate, oblong, or suborbicular, the apex acute, 

obtuse, rounded, or short-acuminate, the base 

obtuse, unequal or rounded, the margins crenulate; 

upper surface glabrous, dark green, shiny, with 

the midvein slightly prominent; lower surface pale 

green, shiny, glabrous or sometimes pubescent, 

with the midvein prominent; petioles slender, 1- 

3 cm long. Inflorescences of small axillary cymes, 

with minute sessile flowers. Staminate flowers 

campanulate, 2.5-3 mm long, yellowish, 

pubescent outside; stamen exserted, the filaments 

twice as long as the perianth. Pistillate flowers 

ca. 2.5 mm long, almost tubular, tomentose; 

stigma exserted, branched. Anthocarp ellipsoid 

or claviform, obtusely 5-angular, 10-12 mm long, 

tomentose, with one or two lines of stipitate glands 

in each of the angles. 

Phenology: Flowering from December to June 

and fruiting from February to August. 


Distribution: In disturbed areas, secondary or 

mature forests, moist to dry, at lower to middle 


and St. Thomas; throughout the Neotropics. 


Reference: Gillis, W. T. 1976. Bougainvilleas of cultivation (Nyctaginaceae). Baileya 20: 34-41. 

31. Family OLEACEAE 

1. JASMINUM 

Erect or scandent shrubs or twining vines. Leaves opposite or less frequently alternate, simple or 

pinnately compound; petioles present; stipules absent. Flowers bisexual, usually showy and fragrant, 

solitary, clustered in axillary or terminal cymes or panicles; calyx campanulate or infundibuliform, 

with 4-9 lobes; corolla hypocrateriform, with 4-9 imbricate lobes, the tube cylindrical; stamens 2, 

included in the tube; ovary superior, the style slender and elongate, the stigma bilobate or less frequently 

simple. Fruit a small berry. A genus of 450 species, native to Asia, Malasia, Africa, and Australia. 

Key to the species of Jasminum 

1a. Leaves compound………………………………………………………………………..…2 

1b. Leaves simple…………………………………………………………………………...….3 

2a. Leaves trifoliolate; sepals acuminate.……………………………………......... 1. J. fluminense 

2b. Leaves pinnately 5-7-foliolate; sepals filiform …………….………….....… 2. J. grandiflorum 

3a. Leaves subtruncate or cordiform at the base; lower surface pubescent….......... 3. J. multiflorum 

3b. Leaves rounded or cuneate at the base; lower surface glabrous, barbate in the axils of the secondary 

veins ...............................……………………………………………………………..… 4. J. sambac 

1. Jasminum fluminense Vell., Fl. Flumin. 10. 

1825 [1829]. 

Fig. 130. A-D 

SYNONYM: Jasminum azoricum sensu Urb., Britton, non 

L. 

Jazmín oloroso, Jazmín de canario, 

Jazmín de trapo 


Stems cylindrical, pubescent, attaining 1 cm in 

diameter, glabrescent when mature. Lateral 

branches numerous. Leaves opposite, trifoliolate, 

5-10 cm long; leaflets 2-5 × 2-3.5 cm (terminal 

leaflet larger than the lateral ones), broadly ovate,

324 


involute, the apex acute or acuminate, the base 

subtruncate, the margins entire; upper surface 

puberulent; lower surface with the midvein 

prominent, barbate in the axils of the secondary 

veins; petioles and petiolules pubescent, the 

petioles 0.5-2 cm long. Inflorescences of axillary 

cymes with numerous fragrant flowers; peduncles 

3-4 cm long; pedicels 3-4 mm long, densely 

pubescent. Calyx green, ca. 3 mm long, 

campanulate, with 4-9 small acuminate lobes; 

corolla white, hypocrateriform, 1.5-2.5 cm long, 

with 4-9 lobes; stamens 2; ovary 4-lobate, the 

stigma bilobate. Fruit a purple or almost black 

berry, shiny, globose, 5-8 mm in diameter. 

Phenology: Flowering from December to 

September and fruiting from January to August. 


Distribution: Along roads, in pastures, or in 

disturbed areas. Species native to Africa but 

described from material collected in Brazil. 

Naturalized on Vieques, St. Croix, St. John, and 

St. Thomas. Cultivated as an ornamental and 

naturalized throughout the tropics. 

Public Forests: Cambalache, Ceiba, and 

Guánica. 

2. Jasminum grandiflorum L., Sp. Pl. ed. 2, 9. 

1762. 

Fig. 130. E-J 

Jazmín, Jazmín de España 


in length. Stems octagonal to almost cylindrical, 

slender, glabrous, puberulent in the area of the 

nodes. Lateral branches numerous. Leaves 

opposite, imparipinnate, 5-7 cm long; leaflets 7 

or 9, opposite, 1-3 × 0.7-1.2 cm, the apex acute 

or obtuse, mucronulate, the margins entire; 

terminal leaflet larger than the lateral ones, 

elliptical, with the base obtuse; the lateral leaflets 

ovate, the base asymmetrical, obtuse, subrounded 

on the basal ones, the distal ones decurrent on 

the rachis; upper surface glabrous; lower surface 

with the midvein prominent, glabrous to 

puberulent; rachis narrowly winged; petioles 

glabrous, 12-14 cm long; petiolules puberulent. 

Inflorescences of axillary dichasial cymes with 3 

fragrant flowers; peduncles 2-5 cm long; pedicels 

8-20 mm long, glabrous, with a pair of minute 

bracteoles on the middle. Calyx green, ca. 1.5 mm 

long, campanulate, with 5 linear lobes, 2-4 mm 

long; corolla white, hypocrateriform, 2.2-2.5 cm 

long, the tube white or pink outside, the lobes 5, 

elliptical, 1.5-2 cm long; stamens 2, included; 

stigma bilobate, slightly exposed. Fruits not seen. 


December and January. 


Distribution: Although not very common, it 

is cultivated in our gardens. Species native to 

Arabia, but widely cultivated throughout the 

tropics. 

3. Jasminum multiflorum (Burm.f.) Andrews, 

Bot. Repos. 8: part 102, t. 496. 1807. 

Fig. 131. A-F 

BASIONYM: Nyctanthes multiflora Burm. f. 

SYNONYM: Jasminum pubescens (Retz.) Willd. 

Jazmín, Jazmín de papel 


slender, tomentose. Leaves opposite, simple, 3-7 

× 2-3.5 cm, ovate, the apex acute, the base 

subtruncate or cordiform, the margins entire; 

upper surface glabrous; lower surface tomentose, 

especially on the veins; petioles tomentose, 6-12 

mm long. Inflorescences of axillary cymes, sessile, 

with numerous fragrant sessile flowers; bracts and 

bracteoles densely pubescent. Calyx green, ca. 1 

cm long, infundibuliform, tomentose, with 

numerous linear lobes; corolla white, 

hypocrateriform, ca. 2 cm long, with 4-9 oblong 

lobes; stamens 2; ovary 4-lobate, the stigma 

bilobate. Fruits not observed on any of the 

collections from Puerto Rico or the Virgin Islands. 


to April. 



Distribution: Native to Asia, introduced as an 

ornamental, which is found naturalized in Puerto 

Rico, Vieques, St. John, and throughout the 

tropics. 

Public Forests: Maricao, Río Abajo, and Toro 

Negro.


326 



4. Jasminum sambac (L.) Soland. in Ait., Hort. 

Kew. ed. 1, 1: 8. 1789. 

Fig. 131. G-I 

BASIONYM: Nyctanthes sambac L. 

Diamela 

Erect or clambering shrub, 1-2 m in length. 

Stems slender, puberulent, glabrous when mature. 

Leaves opposite, simple, 3.5-10.5 × 3-5.8 cm, 

elliptical, the apex obtuse, mucronate, the base 

obtuse to rounded, the margins entire; upper 

surface glabrous, except for the midvein, which 

is puberulent; lower surface glabrous, with the 

midvein prominent, barbate on the secondary 

veins; petioles pubescent, 5-7 mm long. 

32. Family PASSIFLORACEAE 

1. PASSIFLORA 

Inflorescences of axillary cymes, longpedunculate, 

with few fragrant flowers; peduncle 

and pedicels pubescent. Calyx green, ca. 1 cm 

long, infundibuliform, tomentose, with 7-10 linear 

lobes; corolla white, hypocrateriform, the tube 1- 

1.4 cm long, the lobes 5, elliptical, 1-1.3 cm long. 

Fruits not observed. 


September. 


(according to Liogier, 1995), uncommon. 

Distribution: Species native to India, 

introduced as an ornamental. Also on Vieques; 


Herbaceous or woody vines that climb by means of axillary tendrils. Leaves alternate, simple, 

entire or palmately lobed; petioles usually with extrafloral nectaries; stipules usually small and deciduous. 

Flowers usually large and showy, bisexual or less frequently unisexual, actinomorphic, produced in 

axillary cymes or solitary, sometimes covered at the base by an involucre of foliaceous bracts; hypanthium 

short or tubular; calyx of (3-)5(-8) sepals, free or connate at the base. Petals as numerous as the sepals, 

alternating with them, rarely absent; corona of 1 or several whorls of free or connate appendages; 

stamens (4)5, the filaments free or on a stipe (androgynophore), the anthers dehiscent by longitudinal 

valves; ovary superior, usually on a gynophore, or rarely sessile, unilocular, with (2)3(-5) carpels, the 

placentation parietal, with numerous ovules, the styles 3, free or connate at the base, the stigmas 

capitate or oblanceolate. Fruit a berry or less frequently a capsule. A genus of 400 species, 350 of these 

native to tropical America, the rest to the paleotropics. 

Key to the species of Passiflora 

1a. Leaves trifoliolate ...............………………………………………………………… 1. P. berteroana 

1b. Leaves simple...............…………………………………………………………………...…… 2 

2a. Leaves entire, not lobed...............………………………………………………………………… 3 

2.b Leaves variably lobate..............………………………………………………………………….. 7 

3a. Stipules foliaceous, ovate or elliptical; petioles with 3 pairs of glands.........9. P. quadrangularis 

3b. Stipules filiform; petioles with one pair of glands...............…………………………………...… 4 

4a. Flowers subtended by a minute subulate bract or without bracts……………………..…… 5 

4b. Flowers subtended by an involucre of foliaceous bracts………………..……………… 6

328 


5a. Plants tomentose; flowers in groups of 2-6, axillary; petals linear, ca. 3.5 mm long; fruits depressedglobose 

....................………………………………………………………..…7. P. multiflora 

5b. Plants puberulent or glabrous; flowers solitary or in pairs, axillary; petals absent; fruits ovoid or 

ellipsoid......................………………………….……………………………...… 13. P. suberosa 

6a. Bracts of the involucre connate in the lower ½; leaves with three main veins that are borne almost 

from the base ...........................……….…………………………………………… 6. P. maliformis 

6b. Bracts of the involucre free to the base; leaves pinnately veined ..………… 5. P. laurifolia 

7a. Leaves with the lateral lobes divergent, as long as or longer than the central lobe (apex of the leaf), 

the leaf then bilobate or with the apex truncate or retuse. …………………...........…… 8 

7b. Leaves with the central lobe longer than the lateral lobes…………………………..…… 12 

8a. Leaves with the lateral lobes projecting well beyond the apex. ………………….……….9 

8b. Leaves with the lateral lobes as long as the central lobe (the apex), or slightly longer. ....10 

9a. Lateral lobes ca. 3 times longer than the central lobe (apex of the leaf), forming an angle of 

divergence of 45-93°; corolla green ……….…...………………………........… 2. P. bilobata 

9b. Lateral lobes 1.5-2 times longer than the central lobe, forming an angle of divergence of 108-160°; 

corolla brilliant red ……………………………………………............. 8. P. murucuja 

10a. Leaf cordiform at the base; flowers < 2.5 cm in diameter, the sepals green, the petals 

white……………………………………………………………………………………........… 11 

10b. Leaf rounded or cuneate at the base; flowers ca. 5 cm in diameter, the sepals and petals pink-violet 

........................….………………………..………..………………………………........ 14. P. tulae 

11a. Flowers solitary; sepals 2-2.5 cm long; fruit dehiscent, pyriform, reddish when ripe, ca. 2 cm in 

diameter ……………..……………………………………………….......... 10. P. rubra 

11b. Flowers grouped in axillary cymes; sepals ca. 1.5 cm long; fruit indehiscent, depressed-globose, 

purple when ripe, 5-8 mm in diameter ...……………………..….......… 12. P. sexflora 

12a. Leaves with margins serrate…………………………………………………..………....… 13 

12b. Leaves with margins entire or undulate ……………………………….......….. 13. P. suberosa 

13a. Leaves lobate-digitate, with 7 deep lobes; petioles with two pairs of stipitate, conical glands 

…………………….…………………………………….................….… 11. P. serrato-digitata 

13b. Leaves trilobate; petioles with a pair of prominent sessile glands or lacking glands .....….. 14 

14a. Plant pilose, foul-smelling; stipules laciniate, with numerous marginal glands; petioles lacking 

glands; bracts of the involucre deeply laciniate, with marginal glands; fruits ovoid, 2-2.5 cm long, 

orange when ripe ...............................…………………………………….….....…… 4. P. foetida 

14b. Plant glabrous, not foul-smelling; stipules filiform, without glands; bracts of the involucre ovate, 

not laciniate; fruits ovoid, 5-7 cm long, yellow when ripe........………………......... 3. P. edulis 

1. Passiflora berteroana DC., Prodr. 3: 325. 

1828. 

Fig. 132. A-D 

Herbaceous vine, attainig 1.5-4 m in length 

and climbs by means of axillary tendrils. Stems 

slender, angular, striate, glabrous. Leaves 

alternate or grouped on short axillary branches, 

trifoliolate, the leaflets usually deeply trilobate, 

1.2-3.0 × 2.0-4.0 cm, the lobes spathulate, the 

apices truncate and mucronate, the base cuneate, 

the margin undulate, revolute; upper surface 

glabrous; lower surface glabrous, with prominent


venation; petioles 2-3(6) mm long, canaliculate, 

without glands; stipules subulate, 2.5-4 mm long; 

tendrils simple, filamentous. Flowers axillary, 

solitary or in pairs; the bracts subulate, alternate, 

not forming an involucre; peduncle 8-11 mm long, 

articulated near the apex. Sepals subulate, green 

or whitish, 6-7 mm long; petals pale green or 

whitish, subulate, ca. 3 mm long; corona of two 

series of filiform filaments, the outer ones longer, 

yellowish, the inner ones much shorter than the 

outer ones, green; gynophore ca. 3 mm long, 

tubular; stamens 5; ovary ellipsoid, the styles 

recurved, the stigmas capitate. Fruit a fleshy berry, 

globose (fide Liogier, 1994), ca. 7 mm in diameter. 

Phenology: Collected in flower in September 

and December. 


Distribution: Known from two collections 

from Guánica, in dry forests and thickets of the 

south coast. Also in Hispaniola. 


2. Passiflora bilobata Juss., Ann. Mus. Hist. Nat. 

Paris 6: 107. 1805. 

Fig. 132. E-I 

Herbaceous vine, attainig 0.5-2 m in length 

and climbs by means of axillary tendrils. Stems 

slender, angular, striate, glabrous or puberulent. 

Leaves alternate, in the form of a horseshoe or a 

‘v’, with three main veins, coriaceous, with two 

divergent lobes, forming an angle of 45-93° 

between the, the lobes oblanceolate, oblong, or 

linear, 2.2-6.5 × 0.4-1.4 cm, with rounded or 

acuminate apices, the base cuneate or rounded, 

the margin undulate, revolute; upper surface 

puberulent; lower surface glabrous, with 

prominent venation; petioles 3-5 mm long, 

canaliculate, without glands; stipules filiform, ca. 

4 mm long; tendrils simple, filamentous. Flowers 

axillary, in pairs; the bracts subulate, not forming 

an involucre; peduncle ca. 10 mm long, articulated 

near the apex. Sepals oblong, green, 4-6 mm long; 

petals pale green, oblong, as long as the sepals; 

corona of two series of filiform filaments, pale 

green, as long as the sepals; gynophore ca. 2 mm 

long, tubular; stamens 5; ovary claviform, the 

styles recurved, the stigmas capitate. Fruit a fleshy 

berry, ovoid or globose, 1-1.4 cm long, almost 

black, dull. Seeds numerous, elliptical, ca. 2.3 mm 

long, with transverse striae. 


January and in fruit in January. 


Distribution: In dry forests and thickets of the 

south coast. Also in Hispaniola. 

Public Forest: Guánica, Maricao, and Susúa. 

3. Passiflora edulis Sims, Bot. Mag. 45: t. 1989. 

1818. 

Fig. 133. A-G 

Parcha, Water lemon, Passion fruit. 

Woody vine, glabrous, attainig 5-10 m in 

length and climbs by means of axillary tendrils. 

Stems cylindrical, attaining 2 cm in diameter. 

Leaves alternate, glabrous, coriaceous, 7-12 × 9- 

15 cm, deeply trilobate, the lobes oblong or 

elliptical, the apex acute, the base cordiform, the 

margins serrate; upper surface dark green, slightly 

shiny, with the venation yellowish; lower surface 

pale green, with prominent venation; petioles 3- 

6 cm long, with a pair of prominent sessile glands 

in the area where they join the blade; stipules 

filiform, ca. 5 mm long; tendrils simple. Flowers 

solitary, pendulous, axillary, subtended by an 

involucre of 3 ovate bracts, ca. 2 cm long; 

peduncle 4-6 cm long. Sepals oblong, green, 

whitish on the inner surface, 3-3.5 cm long; petals 

oblong, white on the inner surface, 2.5-3 cm long; 

corona with two series of appendages, with violet 

bands; gynophore green, ovoid, lobate, 5-7 mm 

long; stamens 5; ovary ellipsoid, green, the styles 

slightly reflexed, the stigmas capitate. Fruit ovoid, 

5-7 cm long, yellow when ripe, the pericarp 

coriaceous, thick. Seeds numerous, black, 

elliptical, foveate, ca. 5 mm long, covered with a 

juicy orange matrix. 

Phenology: Flowering from April to October 

and fruiting from June to December. 

Status: Exotic, cultivated and naturalized, very 

common. 

Distribution: In disturbed areas of secondary 

vegetation. Native to South America, but 

cultivated throughout the tropics for its edible 

fruits. 

Public Forests: El Yunque, Maricao, Río 

Abajo, Susúa, and Toro Negro.

330 



4. Passiflora foetida L., Sp. Pl. 959. 1753. 

Fig. 133. J-K 

Tagua-tagua, Love–in-the-mist, Papbush 

Slightly woody vine, glandular-pubescent, 

attainig 5 m in length and climbs by means of 

axillary tendrils. Viscid glandular hairs with a 

strong disagreeable odor. Stems cylindrical, 

slightly striate, glandular-pubescent. Leaves 

alternate, chartaceous, usually trilobate, with the 

central lobe longer than the lateral ones, or less 

frequently simple, 5-12 × 5-10 cm, both surfaces 

pubescent, the lobes elliptical to ovate, with the 

apex acute or acuminate, the base cordiform or 

subcordiform, the margins crenate-serrate; 

petioles slender, pubescent, 2-5.5 cm long, pilose, 

eglandular; stipules ovate in outline, pinnatifid; 

tendrils pubescent, simple, 15 cm or more in 

length. Flowers axillary, solitary or clustered in 

pairs or threes, subtended by an involucre of three 

pinnatifid bracts, 2-3 cm long, glandular-viscid; 

pedicels 4-4.5 cm long, pubescent. Sepals 5, 

oblong, green, whitish on the inner surface, 2- 

2.5 cm long, mucronate at the apex; petals white, 

oblong, 2-2.5 cm long; corona multiseriate, the 

segments filiform, 15-17 mm long, the innermost 

series with a violet band at the base; gynophore 

cylindrical, green, with reddish spots, ca. 7 mm 

long; stamens 5; ovary ellipsoid, green, sparsely 

pilose, the styles pilose, the stigmas capitate. Fruit 

a coriaceous or papery berry, ovoid, 1.5-2.5 cm 

long, orange when ripe, covered by the persistent 

bracts. Seeds numerous, oblong, truncate at both 

ends, foveate, ca. 5 mm long, covered by a yellow 

aril. 




Distribution: In dry or moist disturbed areas 

at lower elevations. Also on Culebra, Mona, 

Vieques, St. Croix, St. John, St. Thomas, and 

Tortola; common throughout the Neotropics. 

Public Forests: El Yunque, Maricao, Mona, 

and Susúa 

A very variable species, with 38 varieties 

recognized (Killip, 1938). In Puerto Rico 3 

varieties are found, and in the Virgin Islands 2 

are found. The following key describes the 

differences between them. 


1a. Plants essentially glabrous; fruits red (Puerto 

Rico; the United States (Florida), the 

Bahamas, Cuba, and Hispaniola)...............….. 

.........P. foetida var. riparia (Griseb.) Killip 

1b. Plants hirsute; fruits orange…………..2 

2a. Ovary pubescent; fruit pubescent or 

puberulent, less than 2.5 cm in diameter; 

bracts of the involucre slightly laciniate 

(Puerto Rico, St. Croix; Jamaica, the Lesser 

Antilles, and South America)................. 

……………….…. P. foetida var. foetida 

2b. Ovary glabrous; fruit glabrous, 2-3 cm in 

diameter; bracts of the involucre quite 

laciniate (Puerto Rico, Culebra, Mona, 

Vieques, St. Croix, St. John, St. Thomas, 

Tortola; the Antilles, South America, and the 

tropics of the Old World) ......................…… 

…..P. foetida var. hispida (DC.) Gleason 

5. Passiflora laurifolia L., Sp. Pl. 956. 1753. 

Fig. 133. H-I 

Parcha, Bell apple 

Woody vine, glabrous, attainig 2-5(-10) m in 

length and climbs by means of axillary tendrils. 

Young parts with a reddish tinge. Stems 

cylindrical, flexible. Leaves alternate, glabrous, 

coriaceous, 6.6-12 × 3.7-6.6 cm, oblong or 

elliptic-lanceolate, not lobate, the apex rounded 

or obtuse, sometimes mucronate or shortapiculate, 

the base almost cordiform, truncate to 

almost rounded, the margins entire; lower surface 

pale green, with the midvein prominent; petioles 

0.7-2.5 cm long, reddish, with a pair of globose 

glands in the area where they join the blade; 

stipules filiform, 8-10 mm long, glandular at the 

apex; tendrils simple. Flowers solitary, pendulous, 

axillary, subtended by a persistent involucre of 3 

ovate bracts, 2.5-4 cm long, with the margins 

serrate-glandular; peduncle 2-3.2 cm long. Sepals 

oblong-elliptical, 3-4 cm long, violet-pink on the 

inner surface, green externally, with a dorsal keel 

that projects beyond the apex; petals oblong, 

white, with numerous violet dots on the inner 

surface, 3.5-4 cm long; corona multiseriate, the 

segments filiform, with violet bands, shorter than 

the petals; gynophore cylindrical, yellow, ca. 1

332 



cm long; stamens 5; ovary ellipsoid, yellow, the 

styles curved, the stigmas capitate-bilobate. Fruit 

a coriaceous berry, ellipsoid, 4-6 cm long, green, 

with numerous yellow dots, with the involucre 

persistent at the base. Seeds numerous, elliptictriangular, 

foveate, cream-colored, covered by a 

juicy yellow matrix. 

Phenology: Collected in flower from June to 

August and in fruit in July and August. 

Status: Apparently exotic, uncommon. 

Distribution: Known in Puerto Rico only from 

the collection Sintenis 1890, which came from a 

plant cultivated in Fajardo. In the Virgin Islands 

(St. Croix, St. John, St. Thomas and Tortola) it is 

found in disturbed areas, thickets, and secondary 

forests at low elevation. Also throughout the 

Antilles and South America, extensively 

cultivated for its flowers and fruits. 

6. Passiflora maliformis L., Sp. Pl. 956. 1753. 

Fig. 134. A-B 

Parcha cimarrona 

Woody vine, glabrous or finely pilose, attainig 

2-5(15) m in length and climbs by means of 

axillary tendrils. Stems subangular, cylindrical 

when mature, up to 2.5 cm in diameter. Leaves 

alternate, glabrous, chartaceous, 6-12 × 4-10 cm, 

ovate or ovate-lanceolate, not lobate, the apex 

acute, acuminate, or less frequently rounded, the 

base truncate to almost cordiform, the margins 

entire or undulate; venation pinnate, but the two 

basal secondary veins extending beyond the 

middle of the blade, prominent on both surfaces, 

especially on the lower surface; petioles 1.5-5 cm 

long, with a pair of subsessile glands above the 

middle; stipules linear, 9-15 mm long, not 

glandular; tendrils simple. Flowers solitary, 


ovate bracts, green, 4-6 cm long, connate at the 

base; peduncles 3-7 cm long. Calyx campanulate, 

the tube ca. 1 cm long, the sepals oblong or 

oblong-lanceolate, 3-4 cm long, green, with a 

dorsal keel that projects beyond the apex; petals 

linear-lanceolate, green, with numerous violet 

dots on the inner surface, ca. 3 cm long; corona 

multiseriate, the 2 outer series of filiform 

segments, with violet bands, the inner series 

tuberculate; stamens 5; ovary oblong or 

subglobose, glabrous, the stigmas capitate. Fruit 

a crustaceous berry, globose, 3.5-4 cm in diameter, 

green, with the involucre not persistent. Seeds 

numerous, oblong-obcordate, 5-6 mm long, 

reticulate. 


and March and in fruit in June. 


Distribution: Known in Puerto Rico from 

several collections of Sintenis, one of which comes 

from a primary forest in Utuado. Also on St. 

Thomas; the Antilles, Venezuela, Colombia, and 

Ecuador. 

7. Passiflora multiflora L., Sp. Pl. 956. 1753. 

Fig. 3F; 134. C-F 

Slightly woody vine, tomentose, attainig 2-15 

m in length and climbs by means of axillary 

tendrils. Stems cylindrical, with numerous lateral 

branches; mature stems sulcate; cross section with 

4-lobate xylem. Leaves alternate, tomentose, 

chartaceous, 5-12 × 2-4 cm, oblong or lanceolate, 

not lobate, the apex rounded or obtuse, mucronate, 

the base obtuse or rounded, the margins entire, 

revolute; upper surface dull, pubescent; lower 

surface pale green, dull, pilose, with prominent 

venation; petioles 4-10 mm long, tomentose, with 

a pair of glands in the area where they join the 

blade; stipules filiform, 3-5 mm long; tendrils 

simple. Flowers in groups of 2-6, axillary, 

subtended by a minute subulate bract; peduncle 

5-15 mm long, tomentose. Sepals elliptical, green, 

ca. 3.5 × 1.5 mm; petals linear, white, 3.5 × 0.8 

mm; corona of numerous filiform segments, 

yellowish, ca. 2.5 mm long; gynophore 

cylindrical, broadened at the base, 0.7-0.8 mm 

long; stamens 5, yellowish; ovary ellipsoid, green, 

the stigmas capitate-bilobate, yellow. Fruit a fleshy 

berry, depressed-globose, 6-8 mm in diameter, 

glabrous, turning from green to purple when ripe. 

Seeds numerous, lenticular, rugose, ca. 2 mm 

long, light brown. 


December to March and in fruit from January to 

April. 

Status: Native, uncommon 

Distribution: In dry forests or in the zone of 

mogotes. Also on St. John, St. Thomas, Tortola; 

the United States (Florida), the Bahamas, Cuba, 


Public Forest: Río Abajo.

334 


8. Passiflora murucuja L., Sp. Pl. 957. 1753. 

Fig. 134. G-J 

Slightly woody vine, glabrous, attainig 2-5 m 

in length and climbs by means of axillary tendrils. 

Stems angular, striate. Leaves alternate, bilobate 

(in the form of a boomerang), with the lateral lobes 

1.5-2 times longer than the apex, these up to 4 

cm long, rounded and retuse at the apex, forming 

an angle of divergence of 108-160° between them 

(sometimes the central apex projecting like a small 

lobe), chartaceous, with 3 main veins from the 

base, the base cuneate, obtuse. or rounded, the 

margins entire; upper surface dull, glabrous; lower 

surface pale green, dull, glabrous, with prominent 

venation and a line of circular glands along the 

acroscopic side of the lobes; petioles 1-1.5 cm 

long, glabrous, without glands; stipules linearfiliform, 

2-4 mm long; tendrils simple. Flowers 

solitary or in pairs, axillary; peduncle 1-2.5 cm 

long, glabrous, articulated below the middle; 

bracts minute, below the articulation of the 

peduncle. Calyx tubular, brilliant red, 3-4 cm 

long, the sepals oblong, connate at the base; petals 

long-deltate, brilliant red, 1-2 cm long; corona 

tubular, brilliant red, 1-1.5 cm long, crenate or 

entire at the apex; stamens 5; ovary ovoid, 

glabrous. Fruit a fleshy berry, globose, 1-1.5 cm 

in diameter, black-purple when ripe. Seeds 

numerous, ovate, transversely sulcate, ca. 2.5 mm 

long. 



Status: Native, extremely rare in Puerto Rico. 

Distribution: Known from the mogotes in the 

zone of Quebradillas. Also in Hispaniola. 


9. Passiflora quadrangularis L., Syst. Nat. ed. 

10, 1248. 1759. 

Fig. 135. A 

Granadilla 

Slightly woody or herbaceous vine, glabrous, 

attainig 3-5 m in length and climbs by means of 

axillary tendrils. Stems trigonal or quadrangular, 

the angles winged, becoming cylindrical when 

mature. Leaves alternate, glabrous, chartaceous 

or membranaceous, 10-20 × 8-15 cm, broadly 

elliptical or broadly ovate, not lobate, rounded 

toward the abruptly acuminate apex, the base 

rounded, subtruncate, or cordiform, the margins 

entire; lower surface with the pinnate venation 

prominent; petioles 2-5 cm long, thick, 

canaliculate, with three pairs of sessile glands; 

stipules ovate or ovate-lanceolate, 2-3.5 cm long; 

tendrils simple. Flowers solitary, pendulous, 


ovate bracts, 3-5 cm long; peduncle 1.5-3 cm long, 

trigonal. Sepals ovate or oblong-ovate, 3-4 cm 

long, white, violet, or pink on the inner surface, 

green externally; petals oblong-ovate to oblonglanceolate, 

white with a pink tinge, 3-4.5 cm long; 

corona multiseriate, the segments filiform, as long 

as the sepals, with red-violet and white bands at 

the base, blue in the middle, and with numerous 

pink-blue spots above the middle; gynophore 

thick; stamens 5; ovary ovoid. Fruit a fleshy berry, 

oblong-ovoid, 20-30 × 12-15 mm, green, with 

three longitudinal furrows. Seeds numerous, 

cordiform or almost circular, 7-10 mm long, 

reticulate. 

Phenology: Collected in flower from March 

to September. 


uncommon. 

Distribution: In disturbed areas. Also on St. 

Croix; of uncertain origin, cultivated throughout 

the Neotropics for its edible fruits. 

10. Passiflora rubra L., Sp. Pl. 956. 1753. 

Fig. 135. B-D. 

Parcha colorada, Flor de pasión, 

Bat wings 

Herbaceous vine, that climbs by means of 

axillary tendrils and attains 2-5 m in length. Stems 

angular, almost triangular, pilose, striate, 

attaining 5 mm wide, with many lateral branches. 

Leaves alternate, simple, bilobate or less 

frequently with a third lobe in a central position, 

4-12 × 4-12 cm, chartaceous, pubescent, with 

three main veins that are borne from the base, 

the lobes acute, divergent, mucronate, the base 

cordiform, the margins entire; lower surface with 

the veins prominent, with a reddish tonality; 

petioles 2-4 cm long, pubescent, with a reddish 

tonality, without glands; stipules subulate, 

pubescent, ca. 5 mm long; tendrils simple, 

pubescent, up to 20 cm long. Flowers axillary,


336 



solitary or in pairs; pedicels pubescent, 3-5 cm 

long. Sepals oblong, 1.3-2.5 cm long, pilose and 

green on the outer surface, white on the inner 

surface; petals linear, whitish, 0.9-1.5 cm long; 

corona biseriate, the outer segments filiform, 1- 

1.3 cm long, pink to purple, the inner ones minute; 

gynophore cylindrical, 0.6-1 cm long; stamens 5, 

greenish; ovary ellipsoid-globose, green, hirsute, 

the styles reflexed, the stigmas capitate. Fruit a 

fleshy capsule, tardily dehiscent, up to 2 cm in 

diameter, ellipsoid or ovoid, red or red-pink, 

pilose, with 8 longitudinal ribs. Seeds numerous, 

elliptical, 3-4 mm long, transversely sulcate, 

covered by a white aril. 




Distribution: In disturbed areas, at lower to 

middle elevations. Also on Vieques, St. Croix, St. 

John, St. Thomas, Tortola, and throughout tropical 

America. 



11. Passiflora serrato-digitata L., Sp. Pl. 960. 

1753. 

Fig. 136. A-B 

Tagua-tagua 

Slightly woody vine that climbs by means of 

tendrils and attains 10-15 m in length. Stems 

cylindrical, brittle, attaining 1 cm in diameter; 

cross section with the pith hollow. Leaves 

alternate, 5-7-palmately lobed, 9-15 × 14-20 cm, 

the lobes oblong, acuminate at the apex, the base 

subcordiform, the margins finely serrate; upper 

surface shiny, glabrous; lower surface glabrous, 

with prominent venation; petioles 6.5-15 cm long, 

glabrous, with a pair of conical stipitate glands at 

the base of the blade and another in the middle; 

tendrils simple, axillary, longer than the leaves. 

Flowers solitary, axillary, subtended by an 

involucre of 3 foliaceous bracts, ca. 4 cm long, 

ovate, fleshy, with the veins notably pink. Sepals 

lanceolate, ca. 3 × 2.5 cm, greenish yellow with 

pink dots. Petals 5, white, oblong, ca. 3 × 1 cm; 

corona biseriate, the segments filiform, violet; 

anthers ca. 13 mm long; stigmas subglobose. Fruit 

a globose berry, crustaceous, 5-6.5 in diameter, 

yellow when ripe. Seeds numerous, more or less 

elliptical, ca. 6 mm long, foveate. 

Phenology: Flowering from June to September 

and fruiting from August to February. 


Distribution: Known in Puerto Rico from the 

Sierra de Luquillo, in moist gallery forests. Also 

in Hispaniola, the Lesser Antilles, and South 

America. 


12. Passiflora sexflora Juss., Ann. Mus. Hist. Nat. 

Paris 6: 110, t. 37, f. 1. 1805. 

Fig. 136. C-G. 

Parchita 

Herbaceous vine that climbs by means of 

tendrils and attains 6 m in length. Stems slender, 

pubescent, almost cylindrical or angular, attaining 

4 mm in diameter. Leaves alternate, simple, 

bilobate or less frequently with a third (short) lobe 

in a central position, 6-8(10) × 3.5-14 cm, 

chartaceous, with three main veins that are borne 

from the base, the lobes acute, divergent, the apex 

of the blade forming a central lobe, more or less 

deep, rounded, mucronate, the base subcordiform, 

the margins entire; upper surface pubescent, 

especially on the veins; lower surface with the 

veins prominent, pubescent; petioles 2-3 cm long, 

pubescent, with a violet tonality, without glands; 

stipules subulate, 3-5 mm long; tendrils axillary, 

simple, pubescent, ca. 6 cm long. Flowers minute, 

in axillary cymes; peduncles pubescent, 5-7 mm 

long; pedicels ca. 7 mm long, pubescent. Sepals 

yellowish green, 8-15 mm long, lanceolate, 

abaxially pubescent; petals 5, whitish, 8-9 × 2 

mm, linear; corona biseriate, the filaments white 

with mulberry-colored bands, 5-8 mm long; 

stamens 5; stigmas subglobose. Fruit a depressedglobose 

berry, dark violet, pubescent, 5-8 mm in 

diameter. Seeds numerous, ca. 2 mm long, 

depressed-ovoid, rugulose. 




Distribution: In forests and weedy places at 

middle and upper elevations in moist zones. Also 

on Tortola; Cuba, Hispaniola, Jamaica, the Lesser 

Antilles, the United States (Florida), and Mexico. 



338 



13. Passiflora suberosa L., Sp. Pl. 958. 1753. 

Fig. 137. A-E 

SYNONYM: Passiflora pallida L. 

Parcha yedra, Indigo berry, Ink berry 

Non-woody vine that climbs by means of 

tendrils and attains 1-3 m in length. Stems slender, 

cylindrical, pubescent or glabrous, usually with a 

reddish tinge; mature stems with 4 longitudinal 

wings, of cork or corky material. Leaves alternate, 

chartaceous, glabrous, 4-15 × 4-10 cm, extremely 

variable in form, trilobate, lanceolate, ovate, 

linear, or elliptical, with 3 or 5 main veins that 

are borne from the base, the lobes more or less 

deep, oblong, ovate to almost linear, the central 

lobe longer than the lateral ones, the apex obtuse 

or acute, the base cordiform, rounded, or obtuse, 

sometimes peltate, the margins slightly revolute, 

ciliate; upper surface shiny, glabrous or 

puberulent; lower surface glabrous or pubescent, 

with slightly prominent venation; petioles 15-20 

mm long, glabrous or pubescent, sulcate, reddish, 

with a pair of minute glands above the middle; 

tendrils axillary, simple, glabrous; stipules 

filiform, ca. 5 mm. Flowers erect, solitary or in 

pairs, axillary; pedicels articulate, 15-20 mm long; 

bracts absent. Sepals 5, lanceolate, 8-13 mm long, 

green outside, whitish or light green on the inner 

surface; petals absent; corona biseriate, the 

segments mulberry-colored, shorter than the 

sepals; gynophore cylindrical, ca. 5 mm long; 

stamens 5, green, ascendent; ovary ellipsoid, 

green, the styles reflexed, the stigmas capitate. 

Fruit an ovoid or ellipsoid berry, 10-12 mm long, 

dark violet when ripe. Seeds numerous, ellipsoid, 

ca. 3 mm long, covered by a green pulp. 




Distribution: In disturbed areas, on roadsides, 

in thickets, and in coastal forests. Also on Caja 

de Muerto, Cayo Santiago, Culebra, Desecheo, 

Icacos, Mona, Vieques, Anegada, St. Croix, St. 


throughout the Antilles, tropical America, and 

introduced in the tropics of the Old World. 

Public Forests: Cambalache, Guajataca, 

Guánica, Maricao, Mona, Piñones, Río Abajo, and 

Susúa. 

14. Passiflora tulae Urb., Symb. Antill. 1: 374. 

1899. 

Fig. 137. F-H 

Tagua-tagua serrana, Flor de pasión 


tendrils and attains 4-5 m in length. Stems slender, 

angular, green or mulberry-colored, striate, 

glabrous or puberulent on the younger portions. 

Leaves alternate, coriaceous, semicircular or 

semielliptical, sometimes in the form of a 

boomerang, 3-8 × 2.5-9 cm, with three main veins 

that are borne from the base, the apex more or 

less truncate, with two or three rounded lobes, 

the base rounded, the margins sinuate; upper 

surface dark green, shiny, glabrous, with the veins 

sunken; lower surface pale green, dull, puberulent, 

with prominent venation, with a row of circular 

glands on each side of the midvein; petioles 1.5- 

3 cm long, cylindrical, smooth, not glandular; 

tendrils axillary, simple, longer than the leaves. 

Flowers solitary or in pairs on an axillary 

peduncle, 3-5 cm long; sepals 5, oblong, 3-4 × 

0.7 cm, pink-violet; petals ca. 3 cm long, oblong, 

of the same color as the sepals; corona tubular, 

orange, ca. 1.5 cm long; gynophore ca. 3 cm long. 

Fruit a globose or ellipsoid berry, green, 1-2 cm 

long, with the sepals persistent at the base. 

Phenology: Flowering from December to April 

and fruiting from February to June. 

Status: Endemic to Puerto Rico, not very 

common. 

Distribution: Along the Cordillera Central and 

in some localities in the zone of mogotes. 

Public Forests: Carite, Guilarte, Maricao, and 

Río Abajo. 


Passiflora anadenia Urb. is excluded from the 

flora of Puerto Rico, since the report of this species 

is based on the erroneous identification of a 

collection of Passiflora bilobata from Guánica. 

Reference: Killip, E. P., 1938. The American species of Passifloraceae. Publ. Field Mus. Nat. Hist. 

Bot. Ser. 19: 1-613.

340 



33. Family PHYTOLACCACEAE 


1a. Fruits woody or subwoody…………………………………………………………….......…… 2 

1b. Fruits fleshy ……………………………………………………….……...............… 3. Trichostigma 

2a. Fruits indehiscent, with the sepals much longer than the fruit …………….…........ 1. Agdestis 

2b. Fruits capsular, with the sepals shorter than the fruit ……………………............ 2. Stegnosperma 

1. AGDESTIS 

A monotypic genus, characterized by the following species. 

1. Agdestis clematidea Moçiño & Sessé ex DC., 

Syst. Nat. 1: 543. 1818. 

Fig. 138. A-C 

Twining vine, herbaceous, attainig 15 m in 

length. Stems angular, reddish, striate, puberulent. 

Leaves with a strong, disagreeable odor, alternate, 

chartaceous, ovate or broadly ovate, 3-9 × 3-7.5 

cm, the apex obtuse or short-acuminate, 

mucronate, the base deeply cordiform or hastate, 

the margins crenulate; upper surface dark green, 

dull, with minute dots; lower surface light green, 

dull, puberulent, with prominent venation; 

petioles 1.5-9 cm long, with a reddish tinge, 

sulcate; stipules absent. Flowers bisexual, white, 

in axillary panicles, 6-17 cm long; peduncles 

2. STEGNOSPERMA 

glabrous or puberulent; pedicels 1-2 mm long, 

with a minute green bracteole, lanceolate, at the 

base. Sepals 4, white, 3-5 mm long, oblong or 

oblanceolate, with the parallel venation 

conspicuous; petals absent; stamens 13-20; ovary 

partially inferior, the style conical, with 4 recurved 

stigmatic branches; ovule solitary. Fruits 

coriaceous, indehiscent, turbinate, ca. 3 mm long, 

with the sepals persistent at the base. Seeds 

elliptical. 

Phenology: Flowering from May to December. 



Distribution: Occasional in our gardens. 

Native to Mexico, cultivated sporadically in the 

Antilles and South America. 

Erect or scandent shrubs; glabrous. Leaves alternate, simple; stipules minute, deciduous. Flowers 

bisexual, in terminal racemes; bracts and bracteoles minute, persistent. Calyx of 5 sepals; corolla of 5 

deciduous petals; stamens 10; ovary superior, with 3-5 uniovulate carpels, the styles free, as numerous 

as the carpels. Fruit a globose capsule, dehiscent from the apex toward the base; seeds 1-5, globose. A 

genus of 3 species, of the Antilles and Central America. 

1. Stegnosperma cubense A. Rich. in Sagra, Hist. 

Nat. Cub. 10: 309. 1845. 

Fig. 138. D-H 

Erect, clambering, or creeping shrub, with 

numerous lateral branches, attainig 1.5-5 m in 

length. Stems cylindrical, glabrous, dark gray, 

with minute lenticels. Leaves alternate, coriaceous

342 


or chartaceous, elliptical, 1.2-6 × 0.9-2.7 cm, the 

apex obtuse or less frequently acute or rounded, 

the base obtuse, the margins crenulate; upper 

surface dark green, dull, with inconspicuous 

venation; lower surface light green, dull, with the 

midvein prominent; petioles 4-10 mm long; 

stipules absent. Flowers fragrant, in racemes at 

the end of short axillary branches, up to 7 cm 

long; peduncle glabrous; pedicels 2-3 mm long. 

Sepals 5, white, ca. 2.5 mm long, elliptical, more 

or less concave; petals 5, white, elliptical, 2.5-3 

mm long; stamens 10, the filaments connate at 

the base; ovary superior, with 3-5 carpels, stigmas 

3-5, sessile, elongate. Fruit a reddish capsule, 

1. Trichostigma octandrum (L.) H. Walter, 

Pflanzenr. 4(83): 109. 1909. 

Fig. 138. I-M 

BASIONYM: Rivina octandra L. 

Bejuco de paloma, Bejuco de nasa, 

Bejuco de palma, Basket wiss, 

Black wiss, Hoopvine 

Clambering shrub or liana, with pendulous 

branches, attainig 5-10(-30) m in length. Stems 

cylindrical, 5-15 cm in diameter. Branches 

glabrous, with numerous lenticels. Leaves 

alternate, chartaceous, elliptical or oblong, 4- 

9(13) × 1.7-4(6) cm, the apex acute or acuminate, 

the base acute or obtuse, sometimes unequal, the 

margins crenulate; upper surface dark green, dull, 

glabrous, with the venation reticulate, 

inconspicuous; lower surface light green, dull, 

with the midvein prominent, sometimes the 

secondary veins, the tertiary veins inconspicuous; 

petioles 0.5-1.5(3) cm long, with a yellowish or 

pink tinge, sulcate, swollen at the base; stipules 

absent. Inflorescences of axillary racemes, up to 

15 cm long; peduncle glabrous or puberulent; 

pedicels 4-7 mm long, with a minute green 

3. TRICHOSTIGMA 

ovoid, 4-5.5 mm long, opening by 3-4 woody 

valves. Seeds ellipsoid, black, shiny, ca. 4 mm 

long, covered by a white to pink aril. 

Phenology: Collected in flower from June to 

September and from November to January and in 

fruit from January to March and from August to 

September. 

Status: Probably exotic, cultivated, probably 

naturalized. 


made by Britton and Horne (9628) in 1931 in 

Barrio Asomante near Aibonito. Also in Cuba, 

Hispaniola, Jamaica, and from Mexico to 

Nicaragua. 

Clambering shrubs or lianas with scandent branches; cross section of the stem with discrete vascular 

bundles, surrounded by parenchymatous tissue. Leaves alternate or subopposite; petioles long; stipules 

absent. Flowers bisexual, actinomorphic, in axillary racemes; bracts deciduous; bracteoles persistent; 

calyx of 4 free tepals; petals absent; stamens 8-16, in two whorls, the anthers dehiscent longitudinally; 

ovary superior, unilocular, with a single ovule, the stigma sessile, with numerous branches. Fruit a 

fleshy drupe with a single seed and the persistent sepals at the base. A genus of 3 species, distributed 

throughout the Neotropics 

bracteole, filiform, below or above the middle. 

Sepals 4, white, 3-5 mm long, oblong or elliptical, 

concave; petals absent; stamens 8-16; stigmas 

sessile, capitate, with numerous filiform branches. 

Fruit fleshy, globose or ellipsoid, purple, 5-7 mm 

long. 

Phenology: Flowering and fruiting from April 

to October. 


Distribution: In mature or secondary forests 

from the littoral zone to the Cordillera Central. 

Also on Culebra, Mona, Vieques, St. Croix, St. 


throughout the Antilles and tropical continental 

America. 


Yunque, Guánica, Mona, Guilarte, Río Abajo, 

Susúa, and Tortuguero. 

Commentary: In Guilarte and Río Abajo there 

are several populations of Trichostigma that differ 

from Trichostigma octandrum in having scabrous 

leaves with the tertiary and quaternary venation 

very conspicuous. The recognition of these 

populations at the specific or subspecific level is 

not possible because there are no other significant 

morphological differences.


344 


34. Family PIPERACEAE 

1. PEPEROMIA 

Terrestrial or epiphytic herbs, erect, prostrate, pendulous, or climbing; stems succulent, glabrous 

or pubescent. Leaves alternate, opposite or whorled, with palmate or pinnate venation; petioles elongate 

or short or less frequently absent. Flowers numerous, in terminal spikes or opposite the leaves; stamens 

2; ovary superior, sessile or stipitate, the stigma one, terminal or lateral. Fruit a minute drupe, ellipsoid 

or globose, sessile or stipitate. A genus of 1000 species, distributed throughout the tropics and subtropics. 

1. Peperomia rotundifolia (L.) Kunth in Humb. 


1: 65. 1815. 

Fig. 139. A-D 

BASIONYM: Piper rotundifolium L. 

Yerba de ratón, Yerba linda, Yerba de medio 

real, Bejuco de alcanfor 

Epiphytic herb, ascending by means of small 

aerial roots, attainig 1-3 m in length. Stems 

cylindrical, brittle, pubescent, ca. 1 mm in 

diameter, with numerous lateral branchlets. 

Leaves alternate, fleshy when fresh, aromatic, 

membranaceous on drying, 5-11 mm long, 

elliptical, obovate, or almost circular, the apex


rounded, the base obtuse or almost rounded, the 

margins entire, markedly revolute, ciliate; upper 

surface dark green, glabrous, puberulent, or 

minutely strigulose, with dark dots; lower surface 

concave, pale green, glabrous or puberulent, with 

dark dots; petioles thin, 1-5 mm long, with minute 

dark dots. Inflorescences of terminal spikes, 1- 

2.5 cm long, produced on the lateral branchlets. 

35. Family PLUMBAGINACEAE 

1. PLUMBAGO 



Distribution: On trees and rocks in moist 

forests of the Cordillera Central. Also in Cuba, 

Jamaica, the Lesser Antilles, Trinidad, and 

tropical continental America. 


Guilarte, Maricao, Río Abajo, and Toro Negro. 

Shrubs or herbs, erect, creeping, or climbing. Leaves simple, alternate, petiolate or sessile, with 

pinnate venation; stipules absent. Flowers bisexual, actinomorphic, short-pedicellate, bibracteate, in 

terminal racemes or panicles; calyx with 5 longitudinal ribs covered with stipitate glands, the lobes 

triangular; corolla hypocrateriform, the tube longer than the calyx, the lobes obovate, rounded, or 

truncate; stamens 5, the filaments free, included or slightly exserted, the anthers dehiscent by 

longitudinal valves; ovary superior, unilocular, with one basal ovule per carpel; style united, with 5 

stigmatic branches. Fruit a valvate capsule, membranaceous, with a single seed. A tropical genus of 

12-20 species. 

1. Plumbago scandens L., Sp. Pl. ed. 2, 215. 

1762. 

Fig. 140. A-F 

Slightly woody shrub, scandent, with 

numerous basal or lateral branches, attaining 2-3 

m in length. Stems cylindrical, striate, glabrous. 

Leaves alternate, chartaceous, 3-13 × 1-6 cm, 

ovate, oblong-lanceolate, spathulate, or 

oblanceolate, the apex acute, acuminate, or obtuse, 

the base obtuse, rounded, cuneate, or attenuate, 

sometimes unequal, the margins entire; upper 

surface glabrous; lower surface with lepidote dots; 

petioles 5-10 mm long. Inflorescences of terminal 

panicles, up to 30 cm long; peduncles glabrous, 

striate, sparsely covered with sessile glands; 

bracteoles elliptical, ca. 2 mm long; pedicels ca. 

1 cm long. Calyx green, 7-11 mm long, the ribs 

with sticky stipitate glands; corolla white, 

hypocrateriform, the tube 1.2-2 cm long, the lobes 

0.5-1 cm long, obovate, mucronate; stamens 

exserted, the anthers pale violet. Fruit an oblong 

capsule, covered by the persistent calyx and sticky. 


to August. 


common in our gardens. 

Distribution: Cultivated, rather common, also 

naturalized in disturbed areas, roadsides, and in 

pastures. Also on Desecheo, Mona, St. Croix, St. 

John, St. Thomas, and Tortola; throughout tropical 

America. 

Public Forest: Mona.

346 



35. Family POLYGALACEAE 

1. SECURIDACA 

Lianas or less frequently clambering shrubs or small trees. Leaves alternate, simple; petioles short; 

stipules absent; nodes usually glandular. Flowers bisexual, zygomorphic, in axillary or terminal racemes; 

bracts minute; calyx of 5 free sepals, deciduous, the two inner ones larger, petaloid; petals 3, deciduous, 

united at the base, the central petal in the form of a keel; stamens 8, the filaments united into a 

staminal tube, the anthers dehiscent by a subapical aperture; ovary superior, unilocular, with a single 

ovule, the stigma bilobate. Fruit a samara with a distal wing, the seminiferous locule basal, with a 

single seed. A genus of 50 species, distributed throughout the tropics and subtropics. 

Key to the species of Securidaca 

1a. Leaves narrowly ovate, ovate, or oblong, 3.5-12 cm long, the apex obtuse or shortacuminate..………..………………………………………………………....... 

1. S. diversifolia 

1b. Leaves ovate, elliptical, or obovate, 1-2 cm long, the apex rounded or emarginate......... 2. S. virgata 

1. Securidaca diversifolia (L.) Blake, Contr. U. 

S. Natl. Herb. 23: 594. 1923. 

Fig.141. A-D 

BASIONYM: Polygala diversifolia L. 

SYNONYMS: Elsota diversifolia (L.) Blake 

Securidaca erecta Jacq. 

Liana, with short twining lateral branches, 

attainig 5-10 m in length. Stems cylindrical, 

strigulose. Leaves alternate, coriaceous, 3.5-9(12) 

× 1.7-3.5 cm, narrowly ovate, ovate, or oblong, 

the apex obtuse or short-acuminate, the base 

obtuse, the margins entire, revolute; upper surface 

puberulent, with the midvein flat; lower surface 

with prominent venation, minutely strigulose; 

petioles 2-4 mm long. Inflorescences of racemes, 

terminal on short (4-12 cm long) lateral branches, 

these with rounded leaves, < 1 cm long, at the 

base; pedicels 4-5 mm long, strigulose; calyx of 5 

sepals, unequal, the 3 outer ones ca. 3 mm long, 

violet, rounded, strigulose, the inner ones petaloid, 

8-10 mm long, violet, rounded; keel violet, ca. 8 

mm long, the upper petals spathulate, violet, 

slightly shorter than the keel; ovary glabrous, the 

style curved, glabrous, the stigma truncate. 

Samara 3.5-4.2 cm long, hirsute, the wing with 

reticulate venation, 3-3.5 × 10-13 cm. 

Phenology: Collected in flower in March, 

April, and July. 

Status: Native or naturalized on St. Croix; 

cultivated in Puerto Rico and St. Thomas. 

Distribution: Uncommon in our gardens, 

cultivated at the Agricultural Experiment Station 

in Mayagüez. Its natural distribution is from 

Mexico to Peru and the Lesser Antilles. 

2. Securidaca virgata Sw., Prodr. 104. 1788. 

Fig. 1E; 141. E-M 

SYNONYM: Elsota virgata (Sw.) Kuntze 

Bejuco de sopla, Jaboncillo 

Woody vine, twining, with pendulous 

branches, attainig 5-10 m in length. Stems 

cylindrical, puberulent, slightly flattened when 

mature; cross section with bands of vascular tissue 

alternating with bands of parenchyma. Leaves 

alternate, 1-2 cm long, obovate or oblong, 

minutely strigulose, the apex rounded, retuse, the 

base cuneate or obtuse, the margins entire, 

revolute, the venation inconspicuous on both

348 



surfaces; petioles ca. 1 mm long, strigulose. 

Inflorescences of terminal racemes; pedicels ca. 

6 mm long, minutely sericeous; calyx of 5 sepals, 

unequal, the 3 outer ones 2-4 mm long, pink, 

rounded, strigulose, the inner ones petaloid, 8-12 

mm long, pink, rounded; keel white, ca. 8 mm 

long, the upper petals yellow, fleshy, slightly 

shorter than the keel. Samara 3-4 cm long, 

minutely sericeous, especially the seminiferous 

locule, the wing with reticulate venation, 2-3 × 1 

cm. 

36. Family POLYGONACEAE 

1. ANTIGONON 




Distribution: In disturbed areas, on roadsides, 

in pastures, and in secondary forests at lower and 

middle elevations. Also on Vieques, Cuba and 

Hispaniola. 


Yunque, Guajataca, Maricao, Río Abajo, and 

Susúa. 

Herbaceous or woody vines, that climb by means of the axes of the inflorescences, modified into 

tendrils. Leaves alternate, simple, entire; petioles elongate; stipules connate around the stem to form a 

tubular structure known as an ocrea. Flowers bisexual, actinomorphic, produced in ocreate fascicles 

along axillary or terminal racemes or panicles; perianth of 5 free petaloid tepals; stamens 8, the filaments 

connate at the base, the anthers dehiscent longitudinally; ovary superior, unilocular, with a single 

ovule, the styles 3, free, the stigmas peltate. Fruit an achene with a single seed, covered by the accrescent 

tepals. A neotropical genus of 8 species. 

Key to the species of Antigonon 

1a. Lower surface of leaves tomentose; outer tepals circular, 2.6-3.2 mm long, sagittate at the base, 

with the lobes overlapping; petioles < 1 cm long ...................……………..…1. A. guatemalensis 

1b. Lower surface of leaves puberulent or glabrous; outer tepals ovate, 5-7 mm long, subcordiform at 

the base, the lobes not overlapping; petioles > 1 cm long .............................….2. A. leptopus 

1. Antigonon guatemalense Meissn. in DC., 

Prodr. 14: 184. 1856. 

Fig. 142. A-F 

SYNONYMS: Antigonon macrocarpum Britton & Small 

Antigonon grandiflorum (Bertol) B.L. Rob. 

Bellísima 



puberulent, pentagonal. Leaves alternate, ovate, 

broadly ovate, or ovate-triangular, chartaceous, 

5-9.5 × 3.2-7.7 cm, the apex obtuse or acute, 

mucronate, the base cordiform, the margins 

crenulate; upper surface pubescent, dull, with the 

venation flat; lower surface tomentose, with 

prominent venation; petioles 8-10 mm long, 

tomentose, almost cylindrical. Flowers bisexual, 

in groups of 4-5 flowers, in axillary racemes or 

terminal panicles, the racemes 15-22 cm long, 

the axes densely pubescent with septate trichomes, 

the distal portion of the axis twisting like a tendril; 

pedicels 1.5-2 cm, articulated near the middle, 

the basal portion persistent, sometimes twisted 

like a tendril. Perianth pink, in two unequal series, 

the outer ones circular, 2.6-3.2 cm long, the base 

sagittate, with overlapping sinuses, the inner ones 

ca. 1.5 cm long, ovate; staminal column ca. 2 mm 

long. Achene ellipsoid, ca. 1.2 cm long. 


November.

350 


Status: Exotic, cultivated and spontaneous 

(according to Liogier, 1985). 

Distribution: Cultivated in Puerto Rico and 

the Virgin Islands (St. Croix and St. Thomas), 

although not very common. Native to Guatemala, 

but cultivated in Central and South America as 

well. 

2. Antigonon leptopus Hook. & Arn., Bot. 

Beechey Voy. 308, t. 69. 1838. 

Fig. 142. G-M 

Bellísima, Coralina, Coral, Coralillo, 

Coral vine, Mexican creeper, 

Coralita, Honeysuckle 



puberulent, pentagonal, with many lateral 

branches. Leaves alternate, ovate, triangularovate, 

or almost lanceolate, chartaceous, 5-14.5 

× 2-7 cm, the apex acute or acuminate, the base 

cordiform or truncate, the margins crenulate, 

sometimes ciliate; upper surface light green, 

37. Family RANUNCULACEAE 

1. CLEMATIS 

slightly shiny, puberulent, with the venation 

sunken; lower surface pale green, dull, puberulent 

or glabrous, with prominent venation; petioles 1- 

5 cm long, reddish, puberulent, cylindrical or 

subwinged. Flowers bisexual, in axillary racemes 

or terminal panicles, 10-20 cm long, puberulent, 

terminating in a pair of spiral tendrils; pedicels 

3-4(-10) mm long. Perianth 4-7 mm long, of 5 

ovate or elliptical tepals, intense pink or white; 

staminal column 2-3 mm long, of the same color 

as the tepals. Achene ovoid, 5-8 mm long. 


although more predominantly from June to 

December. 

Status: Exotic, naturalized, locally very 

common. 

Distribution: Originally cultivated, but 

escaped and naturalized, very prolific. In disturbed 

coastal areas, thickets, and dry forests. Also on 

Vieques, Anegada, St. Croix, St. John, St. Thomas, 

and Tortola. Native to Mexico but widely 

cultivated throughout the Antilles, South America, 

and the tropics in general. 


Woody vines that climb by means of twining leaves. Leaves opposite, trifoliolate, pinnate or biternate; 

petioles elongate; stipules absent. Flowers unisexual or bisexual, actinomorphic, produced in cymes or 

umbels, arranged in racemes; calyx of 4-5 free petaloid sepals; petals absent; stamens numerous, the 

filaments free, the anthers dehiscent longitudinally; ovary superior, of numerous free carpels, with a 

single ovule, the style terminal, slender, plumose. Fruit an achene with a persistent style, elongate and 

plumose, dispersed by the wind. A genus of about 250 species, the great majority of which are of the 

temperate zone. 

Key to the species of Clematis 

1a. Mature leaves trifoliolate…………………………………………………………………..2 

1b. Mature leaves 5- or 9-foliolate.………………………..………..….….3. C. flammulastrum 

2a. Plants essentially glabrous …………….……………………………………….1. C. dioica 

2b. Plants pubescent ………..……………………….………………………….2. C. polygama


352 


1. Clematis dioica L., Syst. Nat. ed. 10, 1084. 

1759. 

Fig. 143. J 

Cabellos de ángel 

Slightly woody vine, dioecious, that climbs by 

means of twining leaves and attains 3-10 m in 

length. Stems slender, sulcate, glabrous, almost 

cylindrical, with many lateral branches. Leaves 

opposite, trifoliolate; leaflets coriaceous, ovate or 

cordiform, 5-9 × 2-7 cm, the apex acute or shortacuminate, 

the base subcordiform or rounded, the 

margins entire, sometimes with 1-2 teeth; upper 

surface shiny, glabrous; lower surface pale green, 

dull, glabrous or puberulent toward the base, with 

slightly prominent venation; petiolules more or 

less glabrous, the terminal one 1.5-3 cm long, the 

lateral ones 0.7-2.5 cm long; petioles 3-7 cm long, 

glabrous. Inflorescences axillary or terminal on 

short lateral branches; flowers unisexual, in 

dichasial cymes; pedicels 1.2-4 cm long, with a 

pair of minute bracteoles below the middle. Sepals 

white or cream-colored, oblanceolate or obovate, 

4-10 mm long, more or less glabrous except for 

the lanate-pubescent margins; stamens 30-50, 

unequal; carpels 15-25, ca. 6 mm long. Fruit an 

ellipsoid achene, slightly compressed, 3-4 mm 

long, pilose, the margin not thickened, the style 

plumose, persistent, 2-6 cm long, dispersed by 

the wind. 


October to February. 



made by Sintenis (5843) in 1887 in the area of 

Barrio Callejones in Lares. Recently collected in 

a mogotes in Aguadilla. Known from southern 

Mexico, Belize, and the Greater Antilles. 

2. Clematis flammulastrum Griseb., Pl. Wright. 

153. 1860. 

Fig. 4.E, 143. E-I 



means of twining leaves and attains 3-4 m in 

length. Stems slender, almost cylindrical, brittle, 

puberulent, especially at the nodes; mature stems 

fissured, with corky bark, attaining ca. 1 cm in 

diameter. Leaves opposite, 5-9-foliolate; leaflets 

coriaceous, ovate, lanceolate, oblong, or 

sometimes deltate in outline, 2.5-6 × 0.8-3.6 cm, 

the apex acute or acuminate, the base rounded or 

subcordiform, the margins revolute, lobateserrate, 

especially on the basal leaflets; upper 

surface slightly shiny, glabrous or puberulent, with 

the venation sunken; lower surface pale green, 

dull, glabrous or puberulent, with prominent 

venation; petiolules puberulent, 5-15 mm long; 

petioles 2-7 cm long, puberulent. Inflorescences 

axillary or terminal on short lateral branches; 

flowers unisexual, in paniculiform dichasial 

cymes; pedicels 2.5-3.2 cm long, strigulose, with 

a pair of minute bracteoles almost at the base. 

Sepals oblanceolate or obovate, 8-10 mm long, 

sericeous on the surface; stamens 35-60, unequal; 

carpels 20-30(50), ca. 6 mm long. Fruit an 

elliptical achene, flattened, 3-4 mm long, pilose, 

with the margin thickened on one border, the style 

plumose, persistent, 1.5-3 cm long, dispersed by 



and in fruit in January. 


Distribution: Although not very common, this 

species is found throughout Puerto Rico, except 

for the wet areas. Also in the Greater Antilles. 

Public Forests: Guajataca, Guilarte, and 

Maricao. 

3. Clematis polygama Jacq., Enum. Syst. Pl. 24. 

1760. 

Fig. 143. A-D 



means of twining leaves and attains 10 m in 

length. Stems slender, cylindrical, sulcate, pilose 

or strigose, with many lateral branches. Leaves 

opposite, trifoliolate; leaflets chartaceous, ovate 

or lanceolate, 5-11 × 2-8 cm, the apex acute or 

acuminate, the base rounded or subcordiform, the 

margins entire, not lobate, sometimes with 1-3(5) 

teeth; upper surface dull, glabrous or sparsely 

strigulose; lower surface pale green, dull, glabrous 

or sparsely strigulose, with prominent venation; 

petiolules strigulose, the terminal one 0.7-3.5 cm 

long, the lateral ones 0.4-2 cm long; petioles 3- 

13 cm long, strigulose. Flowers unisexual, in 

axillary simple dichasial cymes, sometimes


354 


solitary; peduncles strigose; pedicels 1.2-3 cm 

long, strigose, with a pair of minute bracteoles 

near the base. Sepals white, cream-colored, or pale 

green, oblong or elliptical, 8-10 mm long, 

sericeous-pubescent on the outer surface; stamens 

16-66, unequal; carpels 20-60, ca. 6 mm long. 

Fruit an elliptical achene, compressed, ca. 4 mm 

long, pilose, the margin thickened, the style 

plumose, persistent, 4-6 cm long, dispersed by 



November and in fruit in December and January. 


Distribution: In disturbed areas, secondary 

vegetation, and thickets. Also from central Mexico 

to Panama and the Greater Antilles. 

Public Forests: Guajataca, Guilarte, Maricao, 


Commentary: The three species present in 

Puerto Rico had been treated as a single species 

under the name of Clematis dioica. The revision 

of Moreno (1993) reveals the presence of three 

species in Puerto Rico, with C. dioica limited to 

the Greater Antilles. The species that is found in 

the Lesser Antilles corresponds, according to her, 

to C. guadeloupae Pers. 

Reference: Moreno, N.P. 1993. Taxonomic revision of Clematis L. subgenus Clematis 

(Ranunculaceae) for Latin America and the Caribbean. Doctoral thesis, Rice University. 

38. Family RHAMNACEAE 

1. GOUANIA 

Woody vines that climb by means of axillary tendrils, basal to the inflorescences. Leaves alternate, 

chartaceous or coriaceous, usually serrate; stipules small, persistent. Flowers bisexual or less frequently 

unisexual, actinomorphic, in axillary or terminal spikes, racemes, or panicles; calyx conical to 

campanulate, with 5 lobes; petals 5, greenish yellow or whitish, unguiculate, concave; stamens 5, as 

long as the petals, usually hooded by the petal, the filaments adnate to the margin of the disc; disc 

crateriform; ovary subinferior, trilocular, the locules uniovulate, the stigmas 3, reflexed. Fruit a 

schizocarp, trilocular, three-winged, septicidal, that separates into 3 indehiscent mericarps; seeds obovate. 

A pantropical genus of about 50 species. 

Key to the species of Gouania 

1a. Leaves 4.5-7.5 cm long, glabrous or puberulent ………..…….….…………….. 1. G. lupuloides 

1b. Leaves 5-15 cm long, pubescent ……………………………………………..… 2. G. polygama 

1. Gouania lupuloides (L.) Urb., Symb. Antill. 

4: 378. 1910. 

Fig. 144. A-H 

BASIONYM: Banisteria lupuloides L. 

SYNONYM: Rhamnus domingensis Jacq. 

Gouania domingensis (Jacq.) L. 

Bejuco indio, Bejuco de sopla, 

Sopla, Chow stick, Soap stick, 

Soap vine, White root 

Woody vine that climbs by means of tendrils 

and attains 7-12 m in length. Stems with many 

lateral branches; bark grayish, fissured. Branches 

cylindrical, green, glabrous or puberulent; tendrils 

simple, semicircular in form, basal to the 

inflorescences or terminal on short lateral 

branches. Leaves alternate, 4.5-7.5(10.5) × 2- 

4(6.5) cm, chartaceous, ovate or elliptical, 

glabrous or puberulent, the apex acute or short-


acuminate, the base rounded or subcordiform, the 

margins serrate; upper surface dark green, shiny, 


green, slightly shiny, with prominent venation; 

petioles 0.5-1.5 cm; stipules 2-3 mm long. 

Inflorescences of racemes, terminal or on short 

axillary branches, 5-10 cm long, pubescent. Calyx 

pubescent outside, the sepals ca. 1 mm long, 

triangular, glabrous on the inner surface; petals 

yellowish, ca. 1 mm long, forming a hood that 

covers the stamen; stamens slightly shorter than 

the petals; disc ca. 1 mm high. Fruit a threewinged 

schizocarp, 5-7 mm long, dehiscent into 

3 trigonal mericarps. Seeds 3-4 mm long. 

Phenology: Flowering from August to March 

and fruiting from November to March. 


Distribution: In disturbed areas, secondary 

forests, and dry forests at lower and middle 


St. Thomas, and Tortola; throughout the Antilles, 

the United States (southern Florida) and from 

southern Mexico to northern South America. 

Public Forests: Cambalache, Ceiba, 

Guajataca, Guánica, Maricao, Río Abajo, and 

Susúa. 

2. Gouania polygama (Jacq.) Urb., Symb. Antill. 

4: 378. 1910. 

Fig. 144. I-N 

BASIONYM: Rhamnus polygamus Jacq. 

SYNONYM: Gouania tomentosa Jacq. 

Mascapelo 

39. Family ROSACEAE 

1. RUBUS 


and attains 8-15 m in length. Stems grayish, 

attaining 3 cm in diameter. Branches green, 

ferruginous-tomentose, slightly striate; tendrils 

simple, semicircular, basal to the inflorescences 

or distal on short lateral branches. Leaves 

alternate, 5-15 × 3.2-7.5 cm, broadly elliptical, 

oblong, or ovate, chartaceous, the apex obtuse, 

acute, or short-acuminate, the base subcordiform 

or rounded, the margins serrate; upper surface 

dark green, slightly shiny, pubescent; lower 

surface pale green, dull, pilose, with prominent 

venation; petioles 1-1.5 cm long, pubescent. 

Inflorescences of axillary or terminal racemes, 10- 

22 cm long. Calyx 1.5-2 mm long, cupuliform, 

whitish, tomentose, the sepals ovate, 0.7-1 mm 

long; petals whitish, ca. 1 mm long, spathulate, 

enclosing the stamens; stamens as long as the 

petals; disc almost pentagonal, fleshy; styles 3. 

Fruit a three-winged schizocarp, 8-12 mm long, 

dehiscent into 3 indehiscent mericarps, trigonal, 

with a wing on each side of the seminiferous 

center. Seeds ovate, light brown, shiny, ca. 3 mm 

long. 


to November and in fruit from December to 

January. 


Distribution: In disturbed areas, more or less 

moist, at middle elevations, in western and central 

Puerto Rico. Also in Cuba, Hispaniola, Tobago, 

Trinidad, and tropical continental America. 

Public Forests: Guánica, Guajataca, Guilarte, 


Erect, clambering, or decumbent shrubs, with numerous recurved spines. Leaves alternate, 

compound, trifoliolate or 5-9-pinnate; stipules small, conspicuous, usually adnate to the petiole, 

persistent. Flowers bisexual, actinomorphic, in axillary or terminal racemes or panicles; calyx with 5 

lobes, usually glandular; petals 5, free, white; stamens numerous, the filaments free; gynoecium of 

numerous coherent carpels, superior, uniovulate. Fruit a syncarp of numerous, minute, coherent drupes. 

A genus of about 250 species, the majority of the temperate zone of the Northern Hemisphere.

356 



1. Rubus florulentus Focke, Abh. Nat. Ver. 

Bremen 11: 411. 1800. 

Fig. 145. A-J 

Fresa de la montaña 

Clambering shrub or woody vine, climbing, 

that supports itself by means of recurved spines 

and attains 5-10 m in length. Stems cylindrical 

or angular, more or less glabrous, with a reddish 

tinge, with numerous recurved spines. Leaves 

alternate, trifoliolate; leaflets 4-11 × 3.3-8 cm, 

elliptical, ovate, broadly elliptical, or oblong, the 

terminal leaflet larger than the lateral ones, the 

apex obtuse, acute, or short-acuminate, the base 

rounded, truncate, or subcordiform, the margins 

serrate-mucronate; upper surface dark green, 

shiny, with the venation sunken, the midvein 

ferruginous-pubescent; lower surface pale green, 

dull, with prominent venation, the midvein 

yellowish, with recurved spines, puberulent, 

especially alongside the midvein and at the base 

40. Family RUBIACEAE 


of the secondary veins; petioles 2.5-6 cm long, 

swollen at the base, spiny; rachis spiny, 1-3 cm 

long; stipules linear, 3-4 mm long, adnate to the 

petiole. Inflorescences of terminal racemes or 

panicles; axes spiny, tomentose, up to 15 cm long; 

pedicels 4-7 mm long, tomentose, spiny. Sepals 

ovate, sericeous outside, free almost to the base, 

ca. 5 mm long, mucronate at the apex; petals 

white, obovate, 12-15 mm long; stamens 

numerous, 3-4.5 mm long; styles glabrous, stigma 

lateral on distal portion. Fruits aggregate, broadly 

ovoid, 1.5-2 cm long, formed of numerous fleshy 

achenes, dark violet in color. 


August. 

Status: Endemic to Puerto Rico, not very 

common. 

Distribution: Known only from the Cordillera 

Central, in Adjuntas, Jayuya, Maricao, and 

Orocovis. 

Public Forests: Guilarte, Maricao, and Toro 

Negro. 

1a. Twining vines or lianas ..................................................................................….. 2 

1b. Herbs or scandent shrubs, not twining……......................................................... 3 

2a. Fruits white, circular, laterally flattened; mature stems sulcate, with numerous vascular cylinders 

…………….............................................................................................................. 1. Chioccoca 

2b. Fruits red or wine-colored, ellipsoid or globose; mature stems terete with a single vascular cylinder 

…………………………………………….……...…........................................................ 6. Sabicea 

3a. Scandent shrubs or herbs, not bering aerial roots; corolla < 1 cm long ….................... 4 

3.b. Shrubs climbing by means of aerial roots; corolla > 2 cm long ...............................… 6 

4a. Leaves with a strong, fetid odor ….……………………………………….................... 4. Lasianthus 

4b. Leaves without fetid smell .................................................................................................. 5 

5a. Herbs; stems hispidulose, retrorsely pubescent along corners; fruits capsular, light 

brown…........................................................................................................................... 2. Diodia 

5b. Shrubs; stems glabrous, fruits of fleshy drupes, red …….............................................. 5. Psychotria 

6a. Flowers solitary; tube of the corolla 6-10 cm long; fruit capsular, green, 5-12 cm long 

.............................................................……………..........................….........................…. 3. Hillia 

6b. Flowers in heads; tube of the corolla 1.5-2.5 cm long; fruit a berry, white, ca. 1.5 cm long 

.…………………......................…..............................................….................… 7. Schradera

358 



1. CHIOCOCCA 

Twining vines or shrubs. Leaves opposite, coriaceous, entire; stipules interpetiolar, deciduous. 

Flowers bisexual or pistillate, 5-merous, actinomorphic, produced in axillary racemes. Calyx 5-lobate; 

corolla infundibuliform or campanulate, with reflexed or expanded lobes; stamens 5; ovary inferior, of 

two carpels, each carpel with a pendulous ovule, the style filiform. Fruit a flattened berry with two 

pyrenes. A neotropical genus of approximately 20 species. 

1. Chiococca alba (L.) Hitchc., Rep. Missouri 

Bot. Gard. 4: 94. 1893. 

Fig. 146. A-H 

BASIONYM: Lonicera alba L 

SYNONYMS: Chiococca racemosa L. 

Chiococca parvifolia Griseb. 

Chiococca alba (L.) Hitchc. var. parvifolia (Griseb.) Urb. 

Chiococca alba (L.) Hitchc. ssp. parvifolia (Griseb.) 

Steyerm. 

Bejuco de berac, Snow berry 

Woody vine or shrub, twining, attainig 3-10 

m in length. Stems sulcate, with numerous 

opposite lateral branches; cross section of the 

mature stem with several cortical vascular 

bundles. Leaves opposite, coriaceous, elliptical, 

lanceolate, ovate, or oblong, 3-8 × 1-3.5 cm, the 

apex obtuse, acute, or acuminate, the base obtuse 

or acute, the margins entire, revolute; upper 

surface dark green, dull; lower surface light green, 

dull, with the midvein prominent; petioles 4-7 mm 

long, glabrous or puberulent; stipules ca. 2 mm 

long, acicular. Flowers bisexual or pistillate, in 

axillary racemes, 2-5 cm long; pedicels 3.5-5 mm 

long. Hypanthium green, glabrous, laterally 

flattened, ca. 2 mm long, crowned by a crateriform 

calyx, 1-1.2 mm long, the sepals triangular, ca. 

0.5 mm long; corolla infundibuliform, pentagonal, 

intense yellow, usually with reddish lines on the 

angles (purple in the floral buds), the tube 4.5-6 

mm long, the lobes reflexed, 1.5-3 mm long; 

stamens slightly exserted, the filaments unequal, 

pilose, connate at the base into a short tube; style 

bilobate, exserted in the pistillate flowers. Fruit a 

circular drupe, laterally flattened, 5-7 mm long, 

fleshy, white when ripe. 

2. DIODIA 


fruiting from June to March. 


Distribution: In forests and on roadsides, in 

dry and moist areas at lower and middle 

elevations. Also on Mona, Vieques, St. Croix, St. 

John, St. Thomas, and Virgin Gorda; throughout 

the Antilles, the United States (Florida), and 




Susúa, and Tortuguero 

Commentary: There is too much variation in 

the characters that have been utilized to 

distinguish Chiococca alba from C. parvifolia, 

so that it is not possible to distinguish two species 

with certainty. For this reason, I consider C. alba 

in a broader sense, including C. parvifolia. On 

the other hand, in Puerto Rico there is another 

species of Chiococca that I consider to be in 

accordance with the original description of C. 

micrantha Johnst. This species is not included in 

this work, since it is of shrubby habit. It can be 

distinguished from C. alba by the following key. 

1a. Woody vine, twining; stems with cortical 

bundles; inflorescences 2-5 cm long, of 5-30 

flowers; corolla intense yellow, 4.5-6 mm long 

.................………...……..…… C. alba 

1b. Erect shrub with arcuate stems; stems without 

cortical bundles; inflorescences less than 1 cm 

long, with 2-3 flowers; corolla pale yellow, 

ca. 2 mm long ............………... C. micrantha 

Erect, decumbent, or scandent herbs. Stems quadrangular. Leaves opposite, entire, sessile or almost 

sessile; stipules interpetiolar, connate, and united to the petiole. Flowers bisexual, sessile, 4-(6)-merous, 

actinomorphic, solitary, in axillary spikes or heads. Calyx 2-4(-6)-lobate; corolla infundibuliform or 

hypocrateriform, with 4(-6) lobes; stamens 4(-6); ovary inferior, of two carpels, each carpel with a

360 


single ovule, the style filiform, the stigma bilobate or capitate. Fruit indehiscent or schizocarpous, dry. 

A genus of approximately 50 species, the majority of tropical America, some species of Africa. 

1. Diodia sarmentosa Sw., Prodr. 30. 1788. 

Fig. 146. I-L 

Scandent or decumbent herb, attainig 3-5 m 

in length. Stems quadrangular, hispidulous or 

puberulent, retrorsely pubescent on the angles. 

Leaves opposite, membranaceous, elliptical or 

oblong-lanceolate, 2.5-7.3 × 0.9-2.8 cm, the apex 

acute or acuminate, the base attenuate, the 

margins entire, revolute; upper surface dull, 

scabrous, with the venation sunken; lower surface 

scabrous, puberulent, with prominent venation; 

petioles < 1 mm long; stipules connate at the base, 

with numerous acicular projections up to 5 mm 

long. Flowers bisexual, in axillary fascicles, 

sessile. Hypanthium green, glabrous, obovoid, ca. 

2.5 mm long, crowned by 4 long-triangular lobes, 

ca. 1.5 mm long, pubescent; corolla tubular, ca. 

3. HILLIA 

Epiphytic shrubs, scandent, with long pendulous branches. Leaves opposite, thick-coriaceous, 

entire, petiolate; stipules intrapetiolar, membranaceous, deciduous. Flowers bisexual, actinomorphic, 

solitary, subsessile, terminal. Calyx with 2-9 foliaceous or truncate lobes; corolla hypocrateriform, the 

tube elongate, narrow, with 3-7 expanded lobes; stamens 4-7; ovary inferior, of two carpels, each 

carpel with numerous ovules, the style filiform. Fruit an elongate capsule, narrow, oblong or cylindrical, 

septicidal; seeds numerous, with an appendage of hairs on the apical portion. A neotropical genus of 24 

species. 

1. Hillia parasitica Jacq., Enum. Pl. Carib. 18. 

1760. 

Fig. 146. M-P 

SYNONYM: Hillia longiflora Sw. 

Hilia, Flor de sierra, Tibey trepador 

Clambering or epiphytic shrub, climbing by 

means of adventitious roots, attainig 2-3 m in 

length. Stems cylindrical, brown, with pendulous 

lateral branches. Leaves opposite, coriaceous, 

glabrous, 3.5-12 × 1.7-6.5 cm, elliptical or 

oblanceolate, the apex abruptly acuminate, 

revolute, the base acute, the margins entire, 

revolute; upper surface dark green, shiny, with 

the midvein yellowish; lower surface pale green, 

with the midvein slightly prominent; petioles 

cylindrical, 10-20 mm long; stipules oblong, 1- 

5.5 cm long, membranaceous, deciduous. Flowers 

2.5 mm long, white, the lobes 5, ca. 0.5 mm long; 

stamens included. Fruit schizocarpous, opening 

in two indehiscent longitudinal halves, each 

containing one seed inside. Seeds dark brown, 

oblong, slightly flattened on one side, ca. 3 mm 

long. 

Phenology: Flowering from July to April and 

fruiting from September to April. 


Distribution: In thickets and pastures and on 

forest edges and roadsides, in moist areas at lower 

and middle elevations. Reported for St. Thomas 

(Britton, 1925). Also in the Antilles, Mexico, 

Central America, northern South America, and 



Río Abajo, Susúa, and Tortuguero. 

solitary, terminal on lateral branches, subtended 

by a pair of oblong bracts, 15-18 mm long. Calyx 

tubular, 3-5 mm long, green; corolla white, the 

tube narrow, 6-10 cm long, the limb with 4-7 

slender lobes, 2-4 cm long, spreading, reflexed at 

the apex. Capsule cylindrical, 5-12 cm long, 

laterally compressed. Seeds numerous, elliptical, 

ca. 1 mm long, with a tuft of silky fibers, ca. 2 cm 

long, in the apical portion. 




Distribution: In moist upper regions along the 

Cordillera Central and in the Sierra de Luquillo. 

Also throughout the Antilles and in tropical South 

America. 


Maricao, and Toro Negro.


362 


1. Psychotria microdon (DC.) Urb., Symb. Antill. 

9: 539. 1928. 

Fig. 147. A-I 

BASIONYM: Rondeletia microdon DC. 

SYNONYM: Psychotria pinnularis Sessé & Moçiño 

4. LASIANTHUS 

Erect or clambering shrubs or small trees, usually with a fetid odor. Leaves opposite, decussate, 

simple, entire; stipules interpetiolar, triangular, persistent or deciduous. Flowers bisexual, 4-6-merous, 

actinomorphic, produced in axillary fascicles. Hypanthium urceolate, ovoid or subglobose, lobes 3-6; 

corolla tubular, hypocrateriform or infundibuliform, with 4-6 expanded or erect lobes; stamens 4-6, 

inserted on the throat, included; ovary inferior, of 4-12 carpels with one ovule, the style filiform, with 

4-10 stigmatic branches. Fruit a fleshy drupe, with 4-12 trigonal pyrenes, each containing a single 

seed. A genus of approximately 150 species, of southeastern Asia, Malasia, tropical Australia, and 

Africa, with two species in the Antilles and one in tropical continental America. 

1. Lasianthus lanceolatus (Griseb.) M. Gómez, 

Noc. Bot. Sist. Habana 86. 1893. 

Fig. 147. J-P 

BASIONYM: Hoffmannia lanceolata Griseb. 

SYNONYMS: Sabicea moralesii Griseb. 

Lasianthus moralesii (Griseb.) C. Wr. 

Mata de peo 

Slightly woody shrub or herbaceous, 

clambering or decumbent, attainig 2 m in length. 

Stems green, glabrous or sparsely lanate, 

cylindrical or slightly flattened or sulcate, 

attaining 5 mm in diameter. Leaves opposite, 

(4.5)6.5-13.3 × 2.3-5.7 cm, brittle and very foulsmelling 

when fresh, chartaceous on drying, 

oblanceolate, obovate, oblong, or elliptical, the 

apex acuminate or short-acuminate, the base acute 

or obtuse, the margins crenulate, revolute; upper 

surface glabrous, dull or shiny, foveate-reticulate, 


green, lanate-pubescent only on the primary and 

5. PSYCHOTRIA 

secondary veins, the venation reticulate, 

prominent; petioles sparsely lanate, 1.2-1.7(2.5) 

cm long; stipules oblong, ca. 3 mm long, early 

deciduous. Axillary buds lanate-pubescent. 

Flowers sessile, in axillary clusters. Calyx green, 

4-4.5 mm long, the lobes 3-5, ovate, ca. 2.5 mm 

long, strigulose; corolla infundibuliform, white, 

the tube ca. 3 mm long, the lobes 4-5, revolute, 

hirsute on the inner surface; ovary partially 

inferior, glabrous. Fruits urceolate, fleshy, with 5 

pyrenes, 5-7 mm long, white or yellowish, with 

the sepals fleshy and persistent at the apex. 


year and fruiting from December to July. 


Distribution: In the interior of moist and wet 

forests of the Cordillera Central and the Sierra 

de Luquillo. Also in the Greater Antilles. 



Erect or less frequently clambering shrubs, herbs, or small trees. Leaves opposite or less frequently 

3 or 4 per node, petiolate; stipules interpetiolar or intrapetiolar, persistent or deciduous. Flowers bisexual, 

4-6-merous, actinomorphic, sessile or pedicellate, in axillary or terminal panicles, corymbs, cymes, 

racemes or heads, with greenish axes. Calyx crateriform, dentate or truncate at the apex; corolla 

gamopetalous, hypocrateriform, infundibuliform, or campanulate, of various colors, but usually white; 

stamens 4-6, exserted or included; ovary inferior, with 2(-5) carpels, each carpel with one ovule. Fruit 

a fleshy drupe with 2(5) pyrenes, each containing a single seed. A pantropical genus of 1,000-1,500 

species. 

Woody shrub, clambering, attainig 2-2.5 m in 

length. Stems grayish, glabrous, slightly flattened 

or quadrangular; lateral branches decussate, short 

or elongate. Leaves opposite, 5-10 × 2-4.3 cm, 

chartaceous, obovate or oblanceolate, the apex


364 


acute or obtuse, the base acute, obtuse, or attenuate 

toward the petiole, the margins entire, revolute; 

upper surface glabrous, with the venation flat; 

lower surface pale green, glabrous or puberulent 

on the veins, the venation prominent; petioles 

glabrous, 0.5-2 mm long; stipules ovate, ca. 1.5 

mm long, early deciduous. Flowers almost sessile, 

in terminal corymbs at the ends of short lateral 

branches. Calyx green, 1.5-2 mm long, 

crateriform, glabrous, the lobes thin; corolla 

campanulate, white, the tube ca. 5 mm long, the 

lobes 3-3.5 mm long, spreading; stamens 5, 

6. SABICEA 

included. Fruits compressed-obovoid, 5-6 mm 

long, red when ripe. Pyrenes 2, sulcate. 

Phenology: Flowering from April to December 

and fruiting from July to March. 


Distribution: In disturbed areas, dry forests 

and thickets. Also on Isla Piñeiro, Vieques, Guana, 


Virgin Gorda; Cuba, Hispaniola, the Lesser 

Antilles, northern South America and along the 

Pacific coast of South America to Peru. 

Public Forests: Ceiba, Guánica, and 

Guajataca. 

Vines or twining lianas or less frequently small trees. Leaves opposite or whorled, entire; stipules 

interpetiolar, simple, persistent. Flowers bisexual, 4-6-merous, actinomorphic, produced in axillary 

cymes or fascicles. Calyx 3-6-lobate; corolla infundibuliform or hypocrateriform, the tube short or 

elongate, with 4-5 lobes; stamens 4-6, inserted on the throat of the corolla, exserted or included; ovary 

inferior, with (2)3-5 carpels, each carpel with numerous axile ovules. Fruit a berry with 2-5 locules; 

seeds numerous, minute, ovoid or angular, foveate. A genus of approximately 135 species, distributed 

in tropical America, Africa, and Madagascar. 

1. Sabicea villosa Roem. & Schult., Veg. 5: 265. 

1819. 

Fig. 148. A-D 

SYNONYM: Sabicea hirsuta Kunth 

Herbaceous or slightly woody vine, twining, 

attainig 5 m in length. Stem green, hirsute, 

cylindrical, glabrescent, attaining 5 mm in 

diameter. Leaves opposite, membranaceous, 6-12 

× 3.2-5.5 cm, ovate, oblong or elliptical, the apex 

acuminate, the base obtuse or less frequently 

attenuate, unequal, the margins entire, ciliate; 

upper surface sparsely hirsute, with the venation 

sunken; lower surface sparsely hirsute, 

particularly on the prominent veins; petioles 

hirsute, 7-14 mm long; stipules broadly ovate, 6- 

7 mm long, strigulose on the outer surface, 

persistent. Flowers sessile, in axillary clusters; 

bracts ovate, 3-5 mm long. Calyx green, strigose, 

2-2.5 mm long, the lobes 5, expanded, glabrous; 

7. SCHRADERA 

corolla tubular, white, strigose, 6-7 mm long. 

Berry globose or ellipsoid, fleshy, strigose, 7-10 

mm long, purple, with the sepals persistent at the 

apex. Seeds numerous, ca. 0.5 mm long, 

cuneiform. 

Phenology: Flowering and fruiting during 

most of the year. 


Distribution: In pastures and forests in moist 

and wet areas at middle elevations. Also tropical 

continental America. 



Excluded Species 

Sabicea cinerea and Sabicea sp. a. (of the 

Spanish edition of this books) are excluded as their 

records were based on misidentification of 

Tetrapteryx inaequalis of the Malpighiaceae. 

Trees or shrubs, epiphytic or climbing by means of adventitious roots. Leaves opposite, coriaceous, 

entire, petiolate; stipules interpetiolar, large, connate at the base, deciduous. Flowers bisexual, 5-10merous, 

actinomorphic, produced in involucrate and pedunculate heads, these solitary or in umbels.


366 


Calyx tubular or crateriform, truncate or with minute lobes; corolla white, infundibuliform, 

hypocrateriform, or tubular-campanulate, fleshy, villous on the throat, with 5-10 valvate lobes; stamens 

5-10, inserted on the throat of the corolla, included or exserted; ovary inferior, with 2-4 carpels, each 

carpel with numerous ovules, the style with 2-4 stigmatic branches. Fruit a berry with 2-4 locules; 

seeds numerous, minute, circular. A neotropical genus of 16-40 species. 

1. Schradera exotica (Gmelin) Standl., Field 

Mus. Pub. Bot. 4: 286. 1929. 

Fig. 148. E-H 

BASIONYM: Urceolaria exotica Gmelin 

SYNONYM: Schradera capitata Vahl 

Schradera vahlii Steyerm. 

Plant with a juvenile phase morphologically 

different from the adult phase. Juvenile plant ca. 

1 m in length, sparsely branched, which grows 

attached to the host plant by means of adventitious 

roots; leaves membranaceous, crenate. Adult plant 

developing into a woody vine, with scandent and 

pendulous branches, attainig 12 m in length. 

Stems cylindrical, up to 2.5 cm in diameter. 

Leaves opposite, coriaceous, glabrous, elliptical 

or oblong, 5-9.5 × 2.8-5.5 cm, the apex and the 

base obtuse, the margins entire, revolute; upper 

surface dark green, shiny, with the midvein 

yellowish green; lower surface pale green, with 

the midvein prominent; petioles yellowish green, 

thick, ca. 1.5 cm long; stipules interpetiolar, 

caducous, obovate, 1.5-2 cm long, reflexed at the 

apex. Inflorescences in terminal heads; involucre 

crateriform, thick, green, ca. 2.5 cm in diameter; 

flowers sessile; peduncles thick, 3-4 cm long. 

Calyx 10-15 mm long, crateriform, green, fleshy, 

truncate at the apex; corolla white, fleshy, tubular, 

with 9 or 10 oblong lobes, the tube pilose inside; 

stamens 6; style white, stigmas 2, green, exserted. 

Fruit a white berry, oblong, fleshy, ca. 1.5 cm long. 

Seeds numerous, minute. 

Phenology: Collected in flower from March 

to July and in November. 


Distribution: In moist and wet forests of the 

Cordillera Central and the Sierra de Luquillo. 

Also in Cuba, Hispaniola, and the Lesser Antilles. 



References: Steyermark, J. A. 1974. Rubiaceae. Flora de Venezuela. Vol IX (1-3): 1-2070. Edición 

Especial del Instituto Botánico. Caracas, Venezuela. Taylor, C. M. 1994. Revision of Hillia (Rubiaceae). 

Ann. Missouri Bot. Gard. 81: 571-609. 

41. Family SAPINDACEAE 


1a. Herbaceous vines, 2-3 m in length, without latex; stems simple (cross section with a single central 

vascular cylinder); fruits capsular, membranaceous, inflated; seeds black, with a white reniform or 

cordiform hilum ...........…..……………………………………........ 1. Cardiospermum 

1b. Woody vines, 5-10 m in length, with milky latex; stems simple or compound (cross section with a 

central vascular cylinder and three or more peripheral vascular cylinders); fruits capsular or 

schizocarpous, neither membranaceous nor inflated; seeds black or brown, with an aril at the base 

or with a light brown circular hilum..............................…………………………………………….2 

2a. Fruit red, capsular, dehiscent, exposing 1-3 black seeds with a white, fleshy aril at the 

base.................................................................................................................... 2. Paullinia 

2b. Fruit light brown, schizocarpous (indehiscent), which separates into three samaroid units with a 

basal wing; seeds not exposed, brown, with a circular scar, lacking an aril................ 3. Serjania


1. CARDIOSPERMUM 

Herbaceous vines or erect herbs, lacking latex. Stems simple (cross section with a single vascular 

cylinder), angular. Leaves alternate, trifoliolate or biternate; leaflets chartaceous; rachis and petioles 

not winged; stipules minute and caducous. Tendrils spirally twisted, in pairs on the axis of the 

inflorescence (this sometimes not developed), vestigial in the non-climbing species. Inflorescence of 

axillary thyrses, with flowers grouped in lateral scorpioid cymes; pedicels articulated at the base; calyx 

of 4 or 5 sepals; petals 4, with a petaliferous appendage on the inner surface; floral disc unilateral, with 

2 or 4 glands; stamens 8, the filaments unequal; ovary superior, 3-carpellate, the style simple, with 3 

stigmatic branches. Fruit an inflated capsule, membranaceous, septicidal; seeds one per locule, globose 

or ellipsoid, with a white reniform or cordiform scar at the base. A genus of neotropical origin, of 

approximately 15 species, 3 of which have a pantropical distribution. 

Key to the species of Cardiospermum 

1a. Petals 5 mm or more in length; glands of the floral disc 2, oblong, 1.2-2 mm long; capsules 

ellipsoid, up to 6 cm long………….…………….….………………………..2. C. grandiflorum 

1b. Petals less than 5 mm long; glands of the floral disc 4(2), ovoid or rounded, ca. 0.4 mm long; 

capsules subglobose or turbinate-trigonal, up to 3 cm long……………………………………2 

2a. Plants slightly robust; leaflets sinuate-dentate; inflorescences of 4 or more cincinni; capsules ellipsoid 

or globose-trigonal; seed with a reniform hilum .......…….....................1. C. corindum 

2b. Plants herbaceous; leaflets deeply lobate or laciniate; inflorescences of 3 cincinni; capsules trigonal, 

depressed at the apex; seed with a cordiform hilum …..……………3. C. halicacabum 

1. Cardiospermum corindum L., Sp. Pl. ed. 2, 

526. 1762. 

Fig. 149. M-N 

Slightly woody vine, much branched from the 

base, that climbs by means of tendrils and attains 

1.5-3 m in length. Stems with 5 longitudinal ribs, 

pubescent; cross section with a single vascular 

cylinder. Leaves alternate, biternate; leaflets 12- 

16 × 9-12 cm, chartaceous, the margins sinuatedentate; 

both surfaces more or less pubescent, 

especially on the prominent venation; terminal 

leaflet rhombic, acute or acuminate at the apex 

and attenuate at the base; lateral leaflets oblongovate, 

acute at the apex, rounded at the base; 

rachis and petioles not winged; petioles 2-3 cm 

long; stipules minute, early deciduous; tendrils 

in pairs, spirally twisted, at the end of short 

axillary axes (aborted inflorescences), from which 

an inflorescence usually develops. Flowers 

functionally unisexual, zygomorphic, in axillary 

racemiform thyrses; cincinni more than 4, usually 

in more than one whorl. Calyx light green, of 4 

sepals, the two outer ones ca. 1 mm long, the inner 

ones ca. 2.5-4 mm long; petals white, obovate, 

3.5-4.5 mm long; petaliferous appendages slightly 

shorter than the petals, fleshy and yellow at the 

apex, forming a hood that encloses the apex of 

the glands of the disc; disc unilateral, with 4 

rounded or ovoid glands, ca. 0.4 mm high; 

stamens 8, the filaments unequal, pubescent; 

ovary trilocular, with one style and 3 stigmas. 

Capsule ellipsoid to globose-trigonal, 2-3 cm long, 

inflated, stramineous when ripe. Seed one per 

locule, spherical, black, ca. 4 mm in diameter, 

with a white, reniform hilum. 


February and May. 

Status: Apparently native, rather common. 

Distribution: In disturbed areas and dry forests 

at low elevation. Also on Desecheo, Mona, 

Vieques, Guana Island, St. Croix, St. John, and 

St. Thomas; throughout the tropics and subtropics. 

Cultivated for its curious capsules. 

Public Forests: Guajataca, Guánica, and 

Mona.

368 


2. Cardiospermum grandiflorum Sw., Prod. 64. 

1788. 

Fig. 149. H-K 

Slightly woody vine, with numerous lateral 

branches, that climbs by means of tendrils and 

attains 5-8 m in length. Stems almost cylindrical, 

striate, glabrous; cross section with a single 

vascular cylinder. Leaves alternate, biternate; 

leaflets chartaceous, glabrous except for some 

hairs on the veins, the margins deeply serrate; 

terminal leaflet rhombic, 4.5-8 × 2-4 cm, the apex 

acute or acuminate, the base cuneate or attenuate; 

lateral leaflets oblong-lanceolate, 2.7-7 × 1.3-3 

cm, the apex acute or acuminate, the base obtuse 

or attenuate; rachis and petiole not winged, 

canaliculate; petioles 1-5 cm long; stipules 

minute, early deciduous; tendrils in pairs, spirally 

twisted, at the end of short axillary axes (aborted 

inflorescences), from which an inflorescence 

usually develops. Flowers functionally unisexual, 

zygomorphic, in axillary racemiform thyrses; 

cincinni more than 4, usually in more than one 

whorl. Calyx light green, of 4 sepals, the two outer 

ones ca. 1.7-3 mm long, the inner ones ca. 5-8 

mm long; petals white, obovate, 6-9 mm long; 

petaliferous appendages slightly shorter than the 

petals, fleshy and yellow at the apex, forming a 

hood that encloses the apex of the glands of the 

disc; disc unilateral, with 2 elongate glands, 

corniform, whitish, 1.2-2 mm long; stamens 8, 

the filaments unequal, glabrous or pubescent; 

ovary ovoid or ellipsoid, villous, with one style 

and 3 stigmas. Capsule membranaceous, inflated, 

ellipsoid or ovoid, 3-5.5 cm long, stramineous 

when ripe. Seed one per locule, spherical, black, 

4-5.5 mm in diameter, with a white, obtusetriangular 

hilum. 

Phenology: Collected in flower in December 

and in fruit in February. 

Status: Exotic, cultivated and locally 

naturalized in Puerto Rico, not very common. 

Distribution: Naturalized at the Experimental 

Station in Mayagüez. Reported for St. Croix and 

St. Thomas. Native to the Neotropics, apparently 


Cultivated for its curious capsules. 

3. Cardiospermum halicacabum L., Sp. Pl. 366. 

1753. 

Fig. 149. A-G 

Herbaceous vine, much branched from the 

base, that climbs by means of tendrils and attains 

1.5-2 m in length. Stems with 5 longitudinal ribs, 

glabrous or puberulent; cross section with a single 

vascular cylinder. Leaves alternate, biternate; 

leaflets chartaceous, puberulent or sparsely 

pubescent, the apex obtuse, acute, or acuminate, 

the base attenuate, the margins lobate or laciniate; 

terminal leaflet lanceolate or triangularlanceolate, 

rhombic or narrowly lanceolate in 

outline, 2-3.5(5) cm long; lateral leaflets ovate, 

lanceolate, or oblong in outline, 1-2.5 cm long; 

rachis and petiole not winged; petioles 2-3 cm 

long; stipules lanceolate, ca. 5 mm long; tendrils 


axillary axes (aborted inflorescences), from which 

an inflorescence usually develops. Flowers 

functionally unisexual, zygomorphic, in axillary 

racemiform thyrses, shorter than the 

accompanying leaf; cincinni usually in whorls of 

3. Calyx light green, of 4 unequal sepals, the outer 

ones ca. 1.2 mm long, the inner ones 3-3.5 mm 

long. Petals white, obovate, 2.5-3.5 mm long; 

petaliferous appendages slightly shorter than the 

petals, fleshy and yellow at the apex, forming a 

hood that encloses the apex of the glands of the 

disc; disc unilateral, with 4 rounded or ovoid 

glands, ca. 0.4 mm long; stamens 8, the filaments 

unequal, pubescent; ovary trilocular, with one 

style and 3 stigmas. Capsule membranaceous, 

subglobose or turbinate-trigonal, inflated, 

stramineous when ripe. Seed one per locule, 

spherical, black, 3-5 mm in diameter, with a white 

cordiform hilum. 


1. Capsules subglobose, as long as wide, 2.5-3.5 

cm long, with the apex retuse................... 

…...........C. halicacabum var. halicacabum 

1. Capsules turbinate-trigonal, wider than long, 

0.8-1.5 cm long, with the apex truncate............ 

............. C. halicacabum var. microcarpum 

3a. Cardiospermum halicacabum var. 

halicacabum L. 

Distinguished by its larger, subglobose fruits 

(see the key). 


December and March.


370 


Status: Apparently native, not very common. 

Distribution: In disturbed or open areas, like 

roadsides and thickets. In all probability native 

to the Neotropics, but distributed in the tropics of 

the Old World. Cultivated as an ornamental for 

its curious capsules. 


2b. Cardiospermum halicacabum var. 

microcarpum (Kunth) Blume, Rumphia 3: 

183. 1847. 

BASIONYM: Cardiospermum microcarpum Kunth 

2. PAULLINIA 

Distinguished from the typical variety by the 

presence of turbinate-trigonal fruits, truncate at 

the apex and of smaller size (see the key). 


November to July and in fruit from January to 

June. 

Status: Apparently native, rather common. 

Distribution: In disturbed, well illuminated 

areas, like roadsides and pastures. Also on St. 

Croix, St. John, and St. Thomas; throughout the 

tropics and subtropics. 

Public Forests: Mona and Tortuguero. 

Vines, woody or less frequently herbaceous, usually with abundant milky latex. Stems simple or 

compound (cross section with a single vascular cylinder or with a central cylinder and 3-5 peripheral 

cylinders). Leaves alternate, trifoliolate, 5-foliolate, biternate, or with numerous leaflets; leaflets 

chartaceous or coriaceous; rachis and petioles winged or not winged; stipules minute and caducous or 

large and persistent. Tendrils spirally twisted, in pairs at the base of the axis of the inflorescence (this 

sometimes not developed). Inflorescence of axillary thyrses, with flowers grouped in lateral scorpioid 

cymes; pedicels articulated at the base; calyx of 4 or 5 sepals. Petals 4, with a petaliferous appendage 

on its inner surface; floral disc unilateral, with 4 glands; stamens 8, with unequal filaments; ovary 

superior, tricarpellate, with one ovule per carpel, the style simple, with 3 stigmatic branches. Fruit a 

woody capsule, winged or without wings, septicidal; seeds one per locule, globose or ellipsoid, black, 

with a fleshy, white arillode on the lower portion. A neotropical genus of approximately 200 species, 

one of which is also found in Africa. 

Key to the species of Paullinia 

1a. Stems cylindrical, tomentose or tomentulose; cross section of the mature stems simple; leaves 

biternate (three trifoliolate pinnae, for a total of 9 leaflets); petioles not winged; fruits three-winged, 

1-1.5 cm long..........................................................................................2 

1b. Stems pentagonal, glabrous; cross section of the mature stems composed of 2-3 peripheral vascular 

cylinders; leaves 5-foliolate, pinnate; petioles winged; fruits pyriform, not winged, 1.5-2.5 cm long 

...............................................…..........................…….....................2. P. pinnata 

2a. Flowers sessile or almost sessile; calyx puberulent; lower surface with numerous glandular papillae, 

not lineate; fruits oblong in outline.......................................……........…1. P. fuscescens 

2b. Flowers pedicellate; calyx tomentose; lower surface not papillose, with dark lines; fruit elliptical or 

obovate in outline .............................…………………………..……………3. P. plumierii 

1. Paullinia fuscescens Kunth in Humb. Bonpl. 

& Kunth, Nov. Gen. Sp. (quarto ed.) 5: 120. 

1821. 

Fig. 150. A-D 


and attains 3-7 m in length. Stems cylindrical, 

tomentose or tomentulose, up to 5 mm in diameter, 

with slight watery latex when cut; cross section 

simple (with a single vascular cylinder). Leaves 

alternate, biternate; leaflets chartaceous, the 

margins serrate, revolute; upper surface sparsely 

pubescent on the midvein, green, shiny; lower 

surface green, dull, the venation prominent and


puberulent, with a tuft of hairs in the axils of the 

secondary veins, with glandular papillae; terminal 

leaflets 4-6 × 1.5-2.5 cm, lanceolate, the apex 

acuminate, the acumen obtuse, the base longattenuate; 

lateral leaflets 1-2 × 0.7-1 cm, almost 

elliptical, the apex obtuse, the base acute or obtuse; 

rachis narrowly winged, 1-1.6 cm long; petioles 

not winged, 7-12 mm long; stipules triangular, 

ca. 1 mm long; tendrils in pairs, spirally twisted, 

at the end of short axillary axes, from which an 

inflorescence usually develops. Inflorescences of 

axillary racemiform thyrses, with few sessile 

flowers, zygomorphic, in lateral cincinni; axes 

tomentose. Calyx whitish green, of 5 sepals, 2- 

2.5 mm long; petals white, cuneate at the base, 

ca. 2 mm long; petaliferous appendages slightly 

shorter than the petals, fleshy and yellow at the 

apex, forming a hood that encloses the glands of 

the disc; stamens 8, the filaments unequal; ovary 

trilocular. Capsule three-winged, red, 1-1.5 cm 

long. Seeds one per locule, black, obovoid, slightly 

laterally compressed, ca. 6 mm long, with a white 

and emarginate aril on the lower half. 


during February and March. 

Status: Exotic, naturalized, locally common 

on St. Croix. 


elevations; cultivated and locally naturalized at 

the Experimental Station in Mayagüez. Native to 

Cuba, Mexico, Central America, Trinidad, 

Tobago, and northern South America. 

2. Paullinia pinnata L., Sp. Pl. 366. 1753. 

Fig. 150. I-O 

Bejuco de costilla, Bejuco de paloma, Basket 

wiss 


and attains 6-10 m in length. Stems obtuse, 3-5angular, 

glabrous or pubescent on the young 

portions, glabrescent when mature, up to 6 cm in 

diameter and producing scarce milky latex when 

cut; bark smooth, light brown; cross section with 

a central vascular cylinder and 2 or 3 peripheral 

vascular cylinders, smaller than the central one. 

Leaves alternate, pinnately 5-foliolate; leaflets 

coriaceous, 5-16 × 2.2-6 cm (the distal leaflet 

larger), elliptical, ovate, or lanceolate, the apex 

acute, the base attenuate or acute on the terminal 

leaflet and obtuse or rounded and sometimes 

asymmetrical on the lateral ones, the margins 

deeply serrate; upper surface glabrous or 

puberulent along the midvein, green, shiny, with 

slightly prominent venation; lower surface 

glabrous or puberulent along the midvein, green, 

dull, the venation prominent, with a tuft of hairs 

in the axils of the secondary veins; rachis and 

petioles broadly winged; stipules subulate, ca. 5 

mm long; tendrils in pairs, spirally twisted, at the 

end of short axillary axes, from which an 

inflorescence usually develops. Flowers 

zygomorphic, in axillary racemiform thyrses; axis 

of the inflorescence tomentose. Calyx light green, 

puberulent, of 5 sepals, the two outer ones ca. 2 

mm long, the inner ones 4-5 mm long; petals 

white, obovate, 4-5 mm long; petaliferous 

appendages slightly shorter than the petals, fleshy 

and yellow at the apex, forming a hood that 

encloses the glands of the disc; disc unilateral, 

with 4 rounded to ovoid glands. Capsule pyriform, 

not winged, red, 1.5-3 cm long. Seeds 12-15 mm 

long, ellipsoid, with a white aril, covering the base 

and more than half of the two sides. 

Phenology: Flowering from April to 

November and fruiting from October to July. 


Distribution: In forests and pastures at middle 

and lower elevations, in moist or dry places. Also 

on Vieques and Tortola; throughout the Neotropics 

and tropical Africa. 


Maricao, Río Abajo, Susúa, and Toro Negro. 

3. Paullinia plumieri Triana & Planchon, Ann. 

Sci. Nat. Bot. Sér. 4: 18. 360. 1862. 

Fig. 150. E-H 


and attains 3-7 m in length. Stems cylindrical, 

tomentose or tomentulose, up to 0.5 cm in 

diameter; cross section simple (with a single 

vascular cylinder). Leaves alternate, biternate; 

leaflets chartaceous, the margins serrate, revolute; 

upper surface glabrous, puberulent on the 

midvein; lower surface with prominent venation 

and puberulent, with a tuft of hairs in the axils of 

the secondary veins, the tertiary venation dark,

372 



in the form of lines, sometimes with glandular 

papillae; terminal leaflets 4-9 × 2-3 cm, lanceolate 

or ovate, the apex obtuse or acuminate, the 

acumen obtuse, the base long-attenuate; lateral 

leaflets 2.5-5 × 1.2-2.2 cm, oblong, the apex 

obtuse or acute, the base obtuse or rounded; rachis 

narrowly winged, 1-3.2 cm long; petioles not 

winged, 1-5.5 cm long; stipules triangular, 1-2 

mm long; tendrils in pairs, spirally twisted, at the 

end of short axillary axes, from which an 

inflorescence usually develops. Inflorescences of 

axillary thyrses, with numerous pedicellate 

flowers, zygomorphic, in lateral cincinni; axes 

tomentose. Calyx densely tomentose, of 4 sepals, 

2.5-3.5 mm long; petals white, obovate, ca. 3.5 

3. SERJANIA 

mm long; petaliferous appendages slightly shorter 

than the petals, fleshy and yellow at the apex; 

ovary tomentose. Capsule three-winged, elliptical 

or obovate in outline, red, pubescent, 1.7-2 cm 

long. Seeds one per locule, black, elliptic-trigonal, 

ca. 6 mm long, with a white and emarginate aril 

at the base. 


February to April. 

Status: Apparently native, although 

uncommon. 

Distribution: Known from one side of Sage 

Mountain on Tortola. Also on some of the Lesser 

Antilles. 

Lianas or herbaceous vines, climbing by means of tendrils, usually with milky latex. Stems simple 

or compound (cross section with a single vascular cylinder or with several vascular cylinders). Leaves 

alternate, trifoliolate, pinnately 5-foliolate, biternate, or triternate; leaflets chartaceous or coriaceous; 

rachis and petioles winged or not winged; stipules minute and caducous. Tendrils spirally twisted, in 

pairs at the base of the axis of the inflorescence (this sometimes not developed). Inflorescence of 

axillary thyrses, with flowers grouped in lateral scorpioid cymes; pedicels articulated at the base; calyx 

of 4 or 5 sepals. Petals 4, with a petaliferous appendage on its inner surface; disc floral unilateral, with 

2 or 4 glands; stamens 8, with unequal filaments; ovary superior, of 3 carpels, the style simple, branched 

in 3 elongate stigmas. Fruit a schizocarp that separates into three samaroid mericarps, the seminiferous 

portion distal and the wing proximal. Seed globose or ellipsoid, with a triangular or rounded scar at the 

base. A neotropical genus of 230 species, the majority of savannas, thickets, and open areas. 

Key to the species of Serjania 

1a. Leaves 5-foliolate pinnate or less often biternate; stems with a single vascular cylinder 

..............…………………………………………………………………………….. 1. S. diversifolia 

1b. Leaves biternate triternate or bipinnate; stems with 5 or more vascular cylinders...............…... 2 

2a. Leaves biternate (9 leaflets); glands of the floral disc pubescent at the base; ovary tomentose; 

seminiferous locules of the fruit tomentose ..............………………..…….… 2. S. lucida 

2b. Leaves triternate or bipinnate (more than 9 leaflets); glands of the floral disc glabrous or puberulent; 

ovary glabrous or puberulent; seminiferous locules of the fruit glabrous or puberulent 

..............…………………………………………………...............………………..... S. polyphylla 

1. Serjania diversifolia (Jacq.) Radlk., Monogr. 

Serjania 136. 1875. 

BASIONYM: Paullinia diversifolia Jacq. 


and attains 6-10 m in length. Young stems 

obtusely pentagonal, glabrous or puberulent, 

reddish tinged, not producing milky latex; mature 

stems cylindrical, rough, fragile, light brown; 

cross section with a single vascular cylinder. 

Leaves alternate, 5- foliolate pinnate or less often

374 


biternate; leaflets coriaceous, the margins 

remotely serrate; upper surface glabrous, shiny, 

with light green venation; lower surface light 

green, dull, lineate, with the reticulate venation 

slightly prominent; distal leaflets rhombate, 4-9 

× 2-4.5 cm, the apex obtuse, retuse and sometimes 

slightly mucronate, the base long-attenuate; 

lateral leaflets oblong or elliptic, smaller than the 

distal ones, the apex obtuse, apiculate, the base 

acute or abruptly attenuate; rachis marginate; 

petioles 2-5-4 cm long, naked, glabrous; stipules 

minute, caducous; tendrils in pairs, spirally 

twisted, at the end of short axillary axes, from 

which an inflorescence usually develops. 

Inflorescence of axillary thyrses; flowers 

zygomorphic; calyx cream to whitish green; 

corolla cream. Fruit reddish tinged, stramineous 

when dry, splitting into 3 samaroid mericarps, 1.5- 

2 cm long. 

Phenology: Collected in fruit during March. 

Status: Quite rare. 

Distribution: Known in Puerto Rico from a 

steril collection of Sintenis (6999) from the 

vivinity of Utuado. Recently rediscoved in 

Guajataca (Acevedo-Rdgz. 13462) and Bayamón 

(Chinea s.n. and Acevedo-Rdgz. & Chinea 13751) 

in late secondary forests on limestone substrate. 

Also in Hispaniola, Cuba and the Bahamas. 

Public Forests: Guajataca. 

2. Serjania lucida Schum., Skr. Naturhist.-Selsk. 

3(2): 128. 1794. 

Fig. 151. A-F 

SYNONYM: Paullinia curassavica sensu West, non L. 

Bejuco de corrales, Bejuco de costilla, Basketwood, 

Black whithe, Cabrite rotting, White 

wist, Basket wiss, Black wiss 


and attains 6-10 m in length. Stems obtusely 

pentagonal or hexagonal, sulcate, glabrous or 

puberulent, producing abundant milky latex when 

wounded; mature stems cylindrical, smooth, 

deeply sulcate, light brown; cross section with a 

central vascular cylinder and 5-6(8) peripheral 

vascular cylinders. Leaves alternate, biternate (9 

leaflets); leaflets coriaceous, the margins serrate 

on the distal portion; upper surface glabrous, 

shiny, with slightly prominent venation; lower 

surface light green, dull, with the reticulate 

venation slightly prominent, barbate in the axil 

of the secondary veins, lineate; distal leaflets ovate 

or lanceolate, 3.5-9 × 1.7-3.8 cm, the apex acute 

or acuminate, the base rounded, abruptly 

attenuate; lateral leaflets lanceolate, 1.7-5 × 0.8- 

2.5 cm, the apex acute or obtuse, the base rounded 

or abruptly attenuate; rachis marginate; petioles 

1-4.5(8.5) long, not winged, glabrous; stipules 

minute, caducous; tendrils in pairs, spirally 

twisted, at the end of short axillary axes, from 

which an inflorescence usually develops. 

Inflorescences of axillary thyrses or forming a 

panicle when they are terminal on the branches; 

flowers zygomorphic, pedicellate, grouped in 

lateral cincinni. Calyx light green or whitish 

green, white-tomentose, of 5 sepals, 3-5 mm long. 

Petals white, obovate, 4-5 mm long; petaliferous 

appendages slightly shorter than the petals, fleshy, 

yellow, and bifurcate at the apex; disc unilateral, 

with 2-4 rounded or ovoid glands, pubescent at 

the base; stamens 8, the filaments pilose; ovary 

trilocular, tomentose. Mericarps samaroid, 1.5- 

2.5 cm long, with a wing on the lower portion, 

the seminiferous portion almost globose, 

reticulate, tomentose, with a single brown, 

ellipsoid seed inside. 



Status: Endemic to eastern Puerto Rico and 

the Virgin Islands, rather common. 

Distribution: On forest edges and roadsides, 

in dry forests and coastal thickets. In eastern 

(Ceiba) and northeastern (Vacía Talega to Fajardo) 

Puerto Rico, to the Virgin Islands (Culebra, 

Palominos, Vieques, Anegada, St. Croix, St. John, 

St. Thomas, Tortola, Virgin Gorda). 

Public Forests: Ceiba and Piñones. 

3. Serjania polyphylla (L.) Radlk., Monogr. 

Serjania 179. 1875. 

Fig. 151. G-J 

BASIONYM: Paullinia polyphylla L. 

SYNONYMS: Paullinia triternata Jacq. 

Serjania triternata (Jacq.) Willd. 

Bejuco de corrales, Bejuco de costilla 


and attains 5-10(20) m in length. Stems 

cylindrical, sulcate, puberulent, glabrescent, 

producing abundant milky latex when wounded; 

mature stems cylindrical, smooth, deeply sulcate,


376 


light brown, attaining 5 cm in diameter; cross 

section with a central vascular cylinder and (5)8- 

10 peripheral vascular cylinders. Leaves alternate, 

triternate (27 leaflets) or exceptionally bipinnate 

(more than 27 leaflets); leaflets coriaceous, the 

margins deeply serrate or lobate on the distal 

portion; upper surface glabrous, shiny, with 

slightly prominent venation; lower surface light 

green, dull, with the reticulate venation slightly 

prominent, barbate in the axil of the veins nearest 

to the base, lineate; distal leaflets ovate or 

rhombic, 1.7-5 × 0.7-3 cm, the apex acute, obtuse, 

truncate, or acuminate, the base abruptly 

attenuate; lateral leaflets elliptical, ovate, or 

oblong, 1-3 × 0.8-2.7 cm, the apex obtuse or 

truncate, the base rounded, abruptly attenuate; 

primary and secondary rachis winged or narrowly 

winged; petioles 1-3 long, not winged, glabrous 

or puberulent; stipules minute, caducous; tendrils 


axillary axes, from which an inflorescence usually 

develops. Inflorescences of axillary thyrses or 

forming a panicle when they are terminal on the 

branches; flowers zygomorphic, pedicellate, 

grouped in lateral cincinni. Calyx pale green, 

white-tomentose, of 5 sepals, 3-5 mm long. Petals 

white, obovate, 4-5 mm long; petaliferous 

appendages slightly shorter than the petals, fleshy, 

yellow, and bifurcate at the apex; disc unilateral, 

with 4 rounded or ovoid glands, glabrous or 

puberulent at the base; stamens 8, the filaments 

pilose; ovary trilocular, glabrous or puberulent. 

Mericarps samaroid, 1.5-2.5 cm long, with a wing 

on the lower portion, the seminiferous portion 

almost globose, reticulate, glabrous or puberulent, 

with a brown, ellipsoid seed inside. 


April and fruiting from October to April. 


Distribution: On forest edges and roadsides, 

in dry forests, coastal thickets, and forests in the 

zone of mogotes. Found from Cayey to Hispaniola, 

with collections in Utuado and the south and west 

coasts of Puerto Rico. Also on Caja de Muerto. 

Public Forests: Cambalache, Guajataca, 

Guánica, Maricao, Río Abajo, and Susúa. 

References: Acevedo-Rodríguez, P. 1993. Systematics of Serjania (Sapindaceae). Part I: A revision 

of Serjania section Platycoccus. Mem. New York Botanical Garden 67: 1-93. Radlkofer, L.T. 1875. 

Monographie der Sapindaceen-Gattung Serjania. Verlag der K.B. Academie. München. 

43. Family SCHLEGELIACEAE 

1. SCHLEGELIA 

Lianas or shrubs, climbing by adventitious roots. Stems cylindrical, interpetiolar zone not glandular; 

pseudostipules inconspicuous, conical, obtuse; cross section of the mature stem with normal peripheral 

phloem. Leaves opposite, simple, coriaceous, without tendrils. Flowers in axillary racemes or terminal 

bracteate panicles. Calyx simple, crateriform, truncate or irregularly lobed; corolla white, pink, purple, 

or reddish, zygomorphic, tubular-campanulate or tubular, more or less glabrous outside, the lobes 

lepidote inside; stamens 4, didynamous, inserted or exserted, 2 of them modified into staminodia; 

ovary superior, globose, bilocular, with a simple central placenta in each locule, the ovules multiseriate 

per locule. Fruit usually a globose berry with a thin pericarp and with a persistent calyx on the basal 

portion; seeds numerous, small, without wings, surrounded by a pulp. A genus of 14 species naturally 

distributed from Guatemala to Brazil, including the Antilles.


1. Schlegelia brachyantha Griseb., Cat. Pl. Cub. 

191. 1866. 

Fig. 152. A-E 

SYNONYMS: Schlegelia brachyantha var. portoricensis Urb. 

Schlegelia portoricensis (Urb.) Britton 

Schlegelia axillaris sensu A. Stahl, non Griseb. 

Higüerito de sierra, Tulipa 

Woody vine, 10-15 m in length, climbing by 

adventitious roots that are produced in the area 

of the nodes. Stems cylindrical, lenticellate, up to 

5 cm in diameter. Leaves opposite; blades simple, 

6-12 × 3-7 cm, elliptical, oblong, or obovate, 

coriaceous, fragile, glabrous, the apex obtuse or 

rounded, sometimes retuse, the base acute, obtuse, 

or rounded, the margins smooth, revolute; upper 

surface dark, shiny, with the midvein sunken; 

lower surface yellowish green, with a prominent 

midvein; petioles recurved, thick, grayish, 8-12 

mm long. Flowers in small axillary racemes; 

pedicels pubescent, 4-8 mm long. Calyx green, 

campanulate, truncate, ca. 5 mm long; corolla 

white or pink with mulberry-colored lines inside, 

tubular, 15-18 mm long, the limb with 5 rounded 

lobes, unequal; stamens 4, exserted, 2 fertile, 2 

staminodia. Berry globose, purple when ripe, 6- 

10 mm in diameter; seeds angular, surrounded 

by the pulp of the fruit. 


particularly from March to August, and fruiting 

in June. 



upper elevations. Also in Cuba and Hispaniola; 

cited for Panama and Venezuela. 


Maricao, Río Abajo, and Toro Negro.

378 


44. Family SCROPHULARIACEAE 

1. LOPHOSPERMUM 

Herbaceous vines, climbing by means of twining petioles, without latex. Basal leaves opposite, the 

distal ones alternate, simple, deltate or cordiform, the margins dentate or crenate; stipules absent. 

Flowers bisexual, solitary, axillary; peduncle elongate, erect or pendulous; calyx urceolate or inflated 

and campanulate, of 4 or 5 sepals; corolla gamopetalous, bilabiate or subtubular, the lobes 5, subequal 

or dimorphic; floral disc absent; stamens 4, didynamous, included, the filaments with glandular


trichomes; ovary superior, of 2 carpels, the style simple, the stigma conical, bilobate or with two short 

branches. Fruit a bivalvate capsule, ovate, globose, symmetrical or asymmetrical, with irregular 

dehiscence. Seeds numerous, circular. A genus of 10 species native from Mexico to Guatemala, the 

majority of montane cloud forests. 

1. Lophospermum erubescens D. Don in Sweet, 

Brit. Flow. Gard. Ser. 2 1: , t. 68, sub t. 75. 

1831 [1830]. 

Fig. 153. A-G 

Mijo del sol 

Herbaceous vine, branched from the base, that 

climbs by means of twining petioles and attains 

several meters in length. Stems cylindrical, with 

a pubescence of glandular trichomes. Leaves 

alternate, simple, deltate, chartaceous, 3.5-7.3(15) 

× 2.5-6.7, the apex acute, the base cordiform, the 

margins dentate-mucronate; upper surface and 

lower surface densely glandular-pubescent, the 

venation slightly prominent or not prominent; 

petioles 3-6.5 cm long, twining, densely 

glandular-pubescent; stipules absent. Flowers 

solitary, axillary; peduncles as long as or longer 

than the petioles, densely glandular-pubescent. 

Calyx green, usually purple tingeD, glandularpubescent, 

the sepals lanceolate, free, 2.5-3 cm 

long. Corolla infundibuliform, intense pink, 

abaxially glandular-pubescent, 6-7.5 cm long, 

with 5 rounded lobes, 1-1.4 cm long; stamens 4, 

didynamous; ovary bilocular, glandularpubescent. 

Capsule membranaceous, ovoid, 1.5- 

1.9 cm long. Seeds discoid, ca. 2 mm wide, 

sculptured in the center, surrounded by a 

translucent wing. 

Phenology: Flowering from January to July 

and fruiting from June to August. 



Distribution: Native to Mexico, but found 

cultivated and naturalized throughout the tropics. 

Reference: Elisens, W.J. 1985. Monograph of the Maurandyinae (Scrophulariaceae-Antirrhineae). 

Syst. Bot. Monographs 5: 1-97. 

45. Family SOLANACEAE 


1a. Corolla rotate, 1-2 cm in diameter, violet; anthers dehiscent by a terminal pore...........……….…2 

1b. Corolla cyathiform, greenish white or yellow, 15-23 cm long; anthers dehiscent by longitudinal 

sutures ..................................………...……………………………………………….…. 2. Solandra 

2a. Leaves deeply lobate; calyx of 5 sepals, without appendages ..................………………... 3. Solanum 

2b. Leaves entire; calyx with 10 linear appendages ..................……………………………. 1. Lycianthes 

1. LYCIANTHES 

Herbs, shrubs, or herbaceous or woody vines, without spines, glabrous or pubescent with simple or 

stellate hairs. Leaves alternate, simple, entire; stipules absent. Flowers actinomorphic, 5-merous, 

bisexual, solitary or fasciculate, axillary. Calyx campanulate or crateriform, truncate, usually with 5- 

10 linear appendages on or near the margin; corolla rotate, with the limb pentagonal or lobate; stamens

380 


5, the filaments shorter than the anthers, the anthers concrescent, dehiscent by terminal pores; ovary 

superior, of two connate carpels, with a single style, the placentation axile, with numerous ovules. 

Fruit a berry, usually globose, with the calyx accrescent at the base; seeds numerous, discoid. A pantropical 


1. Lycianthes virgata (Lam.) Bitter, Abh. Nat. 

Ver. Bermen 24: 307. 1919. 

Fig. 154. A-F 

SYNONYM: Solanum virgatum Lam. 

Berenjena de paloma 

Woody vine, twining, branched from the base, 

attainig 10 m in length. Stems cylindrical, stellatepubescent 

when young, glabrous when mature, 

attaining 2.5 cm in diameter. Leaves alternate, 

6.5-14 × 3.5-5.5 cm, oblong, ovate, or elliptical, 

chartaceous, involute, the apex acute, acuminate, 

or less frequently obtuse, the base obtuse or 

rounded, sometimes asymmetrical, the margins 

undulate; upper surface shiny, with some scattered 

stellate hairs, with the venation lighter; lower 

surface light green, dull, densely stellatepubescent, 

with prominent venation; petioles 

stellate-pubescent, 1-2 cm long. Flowers 4-5 in 

axillary fascicles; pedicels stellate-pubescent, 1- 

1.6 cm long. Calyx green, turbinate, stellate- 

2. SOLANDRA 

pubescent, 3-5 mm long, with 10 ribs that project 

beyond the margin like linear appendages; corolla 

pale violet or lilac, rotate, pentagonal, ca. 2 cm 

in diameter, with five thick greenish ribs on the 

outer surface, which alternate with the pale violet 

membranaceous tissue; anthers oblong, connivent, 

ca. 4 mm long, yellow, that open by a terminal 

pore. Berry globose, fleshy, orange, ca. 8 mm in 

diameter, with the calyx persistent at the base. 

Seeds numerous, circular, flattened, light brown, 

ca. 2 mm in diameter. 

Phenology: Collected in flower from July to 

August and in fruit in November. 


Distribution: In forests and weedy places at 

middle elevations, in the Cordillera Central and 

in the area of mogotes. Also in Cuba and 

Hispaniola. 

Public Forests: Guilarte, Río Abajo, and Toro 

Negro. 

Scandent shrubs or lianas, without spines, glabrous or pubescent with simple or branched hairs. 

Leaves alternate, simple, entire; stipules absent. Flowers actinomorphic, 5-merous, bisexual, solitary, 

terminal. Calyx tubular or campanulate, angular and sometimes inflated, 2-5-lobate; corolla 

infundibuliform or campanulate, with the limb 5-lobate; stamens 5, the filaments inserted on the tube, 

the anthers dehiscent by longitudinal sutures; ovary superior, of 4 connate carpels, partially inferior, 

the placentation axile, with numerous ovules, the style filiform, the stigma capitate. Fruit a coriaceous 

berry, enclosed by the accrescent calyx; seeds numerous, reniform. A neotropical genus of 8 species. 

1. Solandra grandiflora Sw., Königl. Vet. Acad. 

Handl. 8: 300, t. 11. 1787. 

Fig. 157. G-H 

Woody vine, attainig 8-10 m in length. Stems 

cylindrical, glabrous, with numerous short lateral 

branches; bark gray, rugose in dried specimens, 

slightly lenticellate. Leaves alternate or congested 

at the end of short lateral branches, 6.5-14 × 3.2- 

7.3 cm, obovate, spathulate, or broadly elliptical, 

glabrous, coriaceous, the apex almost rounded, 

projecting in a short acumen, the base attenuate, 

the margins revolute, entire; upper surface shiny, 

with the venation flat; lower surface light green, 

with prominent venation; petioles glabrous, 

slender, 2-5.5 cm long. Flowers solitary, at the 

end of short lateral branches, pedicels glabrous, 

thick, not distinguishable from the calyx. Calyx 

green, tubular-campanulate, glabrous, 4-10 cm 

long, inflated-angular; corolla cyathiform, 14-23 

cm long, greenish white or yellow, with several 

marron lines inside, the tubular portion shorter


382 


than or as long as the dilated portion, the limb 

with 5 sinuate lobes, more or less revolute; 

stamens as long as the corolla. Berry coriaceouswoody, 

depressed-ovoid, ca. 3 cm long, with the 

calyx split and persistent at the base. 


to December. 

Status: Apparently exotic, cultivated and 

naturalized, uncommon. 

Distribution: Cultivated or sometimes at the 

margin of forests along the Cordillera Central. 

Cultivated on St. Thomas. Apparently native to 

1. Solanum lanceifolium Jacq., Collectanea 2: 

286. 1788 [1789]. 

Fig. 155. A-D 

Woody vine, scandent, that supports itself on 

the host plants by means of recurved spines and 

attains 2-6 m in length. Stems cylindrical, stellatepubescent, 

with few or numerous recurved spines, 

yellowish. Leaves alternate, 10-18 × 3.7-7 cm, 

lanceolate, ovate, or elliptical, chartaceous, the 

apex acute, obtuse, or acuminate, the base obtuse 

or rounded, asymmetrical, the margins entire; 

upper surface dull, with some scattered stellate 

hairs, with the venation flat or slightly prominent; 

lower surface light green, dull, sparsely stellatepubescent, 

with prominent venation, the midvein 

with yellowish recurved spines; petioles stellate- 

3. SOLANUM 

Jamaica, today found throughout the Antilles, 

Panama, Venezuela, and Brazil. 

Public Forests: Guilarte and Maricao. 


Solandra longiflora Sw. has been collected on 

St. Croix from cultivated plants, and apparently 

does not grow there spontaneously. This species 

is distinguished from S. grandiflora by the longer 

corolla (23-32 cm long), with the tubular portion 

longer than the dilated portion. 

Herbs, shrubs, trees, or herbaceous or woody vines, usually with spines or prickles, glabrous or 

pubescent with simple or stellate hairs. Leaves alternate, simple or compound, entire or lobate; stipules 

absent. Flowers actinomorphic, 5-merous, bisexual or rarely unisexual, produced in racemes or solitary. 

Calyx deeply lobate; corolla usually rotate, with the limb pentagonal; stamens 5, the filaments shorter 

than the anthers, the anthers concrescent, dehiscent by terminal pores; ovary superior, of two connate 

carpels, the placentation axile, with numerous ovules, the style filiform, the stigmas bifid. Fruit a berry 

with numerous seeds inside. A cosmopolitan genus, of approximately 1,400 species. 

Key to the species of Solanum 

1a. Plants markedly spiny; corolla white …….…………………………..…....1. S. lancifolium 

1b. Plants not spiny or inconspicuously spiny; corolla lavender or pale violet……………..…2 

2a. Corolla deeply lobate, the limb 2-2.5 cm wide; anthers 3-5 mm long.........2. S. seaforthianum 

2b. Corolla rotate (the lobes short), the limb 3.5-6 cm wide; anthers 9-10.5 mm long.................. 

....……………………….…………..………………………………………....3. S. wendlandii 

pubescent, 1-5 cm long, usually with recurved 

spines. Flowers in lateral racemes, 2-4 cm long, 

stellate-pubescent; pedicels stellate-pubescent, 

1.5-1.7 cm long. Calyx green, campanulate, 

stellate-pubescent, 4.5-5 mm long, of 5 subulate 

sepals; corolla white, deeply lobate, stellatepubescent 

on the outer surface, the tube ca. 1.5 

mm long, the lobes 8-11 mm long, oblong, oblonglanceolate, 

or ovate, expanded or slightly reflexed; 

anthers subulate, connivent, 6.2-6.5 mm long, 

yellow; style projecting beyond the anthers, the 

stigma capitate. Berry globose, fleshy, brilliant 

red or orange when ripe, 0.9-1.3 cm in diameter, 

glabrous. Seeds numerous, circular, flattened, 

yellowish, 3.2-3.5 mm in diameter. 


January to August. 

Status: Native, uncommon.


Distribution: In forests and thickets. Vieques 

(according to Liogier, 1995), St. John, St. Thomas 

(according to Krebs, 1847), and Tortola; also in 

Hispaniola, the Lesser Antilles, Mexico, Central 

America, Colombia, Venezuela, and Ecuador. 

2. Solanum seaforthianum Andr., Bot. Repos. 

8, t. 504. 1807. 

Fig. 155. E-I 

Jazmín de Italia, Falsa belladona 

Slightly woody vine, scandent, attainig 3-6 m 

in length. Stems more or less cylindrical, 

puberulent, glabrescent, unarmed. Leaves 

alternate, trifoliolate, 5-7(9)-pinnately compound, 

5-7(9)-pinnatilobed, or less frequently simple, 4.5- 

9(14) × 2.6-10 cm, membranaceous; leaflets or 

segments elliptical or oblong-elliptical, the apex 

acute or acuminate, the base obtuse to almost 

cordiform, asymmetrical, the margins entire; 

upper surface dull, appressed-puberulent; lower 

surface light green, dull, appressed-puberulent, 

especially on the prominent venation; petioles 

appressed-puberulent, 2-6.5 cm long, slender. 

Flowers in terminal panicles but appearing lateral 

(sympodial by apposition), 6-23 cm long, 

puberulent; pedicels glabrous or puberulent, 8- 

12 mm long. Calyx green, 1-1.5 mm long, shortconical 

but attenuate toward the pedicel, glabrous 

or puberulent, almost truncate or with 5 very short 

lobes, obtuse; corolla lavender (rarely white), 

deeply lobate, the lobes 5-12 mm long, oblong or 

almost elliptical, expanded; anthers oblong, 3-4 

mm long, not connivent, the filaments slightly 

unequal; style projecting beyond the anthers, the 

stigma capitate. Berry globose, fleshy, crimson, 

9-16 mm in diameter. Seeds numerous, circular, 

flattened, ca. 3 mm in diameter. 


year, collected in fruit in January, February, and 

June. 

Status: Exotic, cultivated as an ornamental and 

naturalized, although not very common. 

Distribution: In disturbed areas, like roadsides 

and pastures. Also on Vieques, St. Croix, and St. 

Thomas. Native to South America, but cultivated 


Public Forests: Carite and Río Abajo. 

3. Solanum wendlandii Hook. f., Bot. Mag. t. 

6914. 1887. 

Fig. 155. J-L 

Campana de pascua, Flor de la mañana, Papa 

aérea, Giant potato climber 

Slightly woody vine, scandent, attainig several 

meters in length. Stems cylindrical, glabrous, 

unarmed or less frequently with some minute 

recurved spines. Leaves alternate, deeply (3)5-7pinnatilobate 

or sometimes simple, 4.5-7.5-17 × 

4.6-12 cm, membranaceous, glabrous, with 

numerous minute dots, elliptical when simple, 

4.5-14 × 4.6-7.5, the apex short-acuminate, the 

base obtuse to almost rounded; segments or lobes 

elliptical or oblong-elliptical, usually 

asymmetrical, the apex acute or acuminate, the 

margins entire; lower surface dull, the primary 

vein prominent, sometimes with recurved spines; 

petioles glabrous, 3-6.5 cm long, usually narrowly 

winged, sometimes with recurved spines. Flowers 

unisexual, in terminal pseudodichotomous 

panicles, 8-20 cm long, glabrous; pedicels 

glabrous, 1-1.7 cm long. Staminate flowers: calyx 

green, cyathiform, glabrous, 5-7 mm long, with 

5 ovate-deltate or ovate lobes, 1-3 mm long, 

apiculate at the apex; corolla lavender, rotate, 

pentagonal, 3.5-6 cm wide, the lobes apiculate, 

ciliate; anthers oblong, 9-10.5 mm long, not 

connivent, dehiscent by terminal pores and by 

longitudinal sutures, the filaments thick, unequal. 

Pistillate flowers and fruits not observed. 

Phenology: Collected in flower from April to 

October. 


escaped from cultivation (according to Liogier, 

1995), although not very common. The plants of 

this species that are found in cultivation seem to 

be all male, and apparently reproduce asexually. 

Distribution: In moist areas of the cordilleras. 

Native to Costa Rica, but widely cultivated in the 

tropics for the beauty of its flowers. 


Solanum pyrifolium Lam. was reported for 

Puerto Rico by Liogier (1995) based on one of his 

collections (Liogier 34137), from the area of the 

Cabezas de San Juan in Fajardo. This species is 

here excluded from the flora of Puerto Rico 

because his report is based on an erroneous

384 



identification of Solanum seaforthianum. Solanum 

pyrifolium is a species endemic to Hispaniola that 

is distinguished from S. seaforthianum by its 4-5 

mm long calyx, with rounded lobes ca. 1 mm long 

(vs. calyx 1-1.5 mm long, truncate, with obtuse 

lobes 0.1-0.2 mm long) and by simple and entire 

leaves (vs. leaves trifoliolate, 5-7(9)-pinnately 

compound, 5-7(9)-pinnatilobed, or less frequently 

simple and entire). The specimen Liogier 34137 

has almost all the leaves simple and entire, but a 

scrupulous examination reveals that it also has 

trifoliolate leaves. 

Reference: Bernadello, L.M. and A. T. Hunziker. 1987. A synoptical revision of Solandra 

(Solanaceae). Nord. J. Bot. 7: 639-652. 

46. Family TROPAEOLACEAE 

1. TROPAEOLUM 

Scandent or creeping herbs, annual or perennial, usually with tuberous roots. Leaves alternate, 

simple or palmately compound, entire or lobate, long-petiolate, peltate at the base; stipules minute. 

Flowers bisexual, 5-merous, solitary, axillary or less frequently in axillary umbels. Calyx deeply lobate, 

with a long basal spur; petals free; stamens 8, in two whorls, the filaments unequal, the anthers dehiscent 

by longitudinal sutures; ovary superior, of three connate carpels, the placentation axile-apical, with a 

single ovule per carpel, the style filiform, with three short stigmatic branches. Fruit schizocarpous, 

separating into three fleshy mericarps with a single seed inside. A genus of the New World, with 

approximately 86 species. 

1. Tropaeolum majus L., Sp. Pl. 345. 1753. 

Fig. 156. A-I 

Herbaceous vine, scandent or creeping, 


glabrous, fleshy. Leaves alternate, ascendent, 

simple, more or less circular, peltate, 5-8 cm in 

diameter, membranaceous, glabrous; upper 

surface dark green, dull, with the venation light; 

lower surface pale green; petioles glabrous, 5.5- 

15(25) cm long. Flowers solitary, axillary, 

ascendent; pedicels glabrous, as long as or longer 

than the petioles. Calyx yellow or pale orange, 

the sepals unequal, ovate, oblong, 2-2.3 cm long, 

the spur straight or slightly curved, up to 3 cm 

long; petals orange or red, obovate, longunguiculate, 

3-4 cm long, three of them longciliate 

on the margin adjacent to the claw; anthers 

lanceolate, the filaments slender, unequal; ovary 

trilobate, glabrous. Fruit tricoccal, ca. 10 mm 

long. 



naturalized, although not very common. 

Distribution: In moist areas of the Cordilleras 

Central. Native to the Andean cordillera, but 

widely cultivated for the beauty of its flowers. 

Public Forest: Guilarte.

386 



47. Family ULMACEAE 

1. CELTIS 

Shrubs, trees, or lianas, with spines or unarmed. Leaves alternate, simple, usually serrate and less 

frequently crenate or entire, petiolate; stipules minute and deciduous. Flowers 5-merous, unisexual or 

sometimes bisexual; the staminate flowers in axillary cymes; the pistillate flowers solitary or few in 

axillary cymes; calyx of free sepals. Petals absent; stamens 5; ovary superior, with a single locule; 

ovule solitary and pendulous; styles 2. Fruit a fleshy drupe. A genus of approximately 80 species, the 

majority of the Northern Hemisphere and South Africa. 

1. Celtis iguanaea (Jacq.) Sarg., Silva 7: 64. 1895. 

Fig. 157. A-G 

BASIONYM: Rhamnus iguanaeus Jacq. 

SYNONYMS: Momisia iguanaea (Jacq.) Rose & Standl. 

Celtis aculeata Sw. 

Azufaifo, Cock spur 

Scandent shrub or liana that climbs by means 

of short lateral branches with recurved spines, 

attainig 3-10(20) m in length. Stems glabrous or 

puberulent at the extremities, cylindrical, 

attaining 8 cm in diameter; bark grayish or light 

brown, with numerous lenticels. Leaves alternate,

388 


4-15 × 2.5-8.7 cm, chartaceous, ovate, oblong, or 

sometimes obovate, the apex short-acuminate or 

sometimes obtuse, the base rounded, cordiform, 

or truncate, the margins entire, crenate, or sparsely 

serrate; upper surface yellowish green, dull, 

glabrous or puberulent, punctate, with three main 

veins, sunken; lower surface yellowish green, dull, 

glabrous, the venation prominent, with small tufts 

of hairs in the axils of the secondary veins; petioles 

2-15 mm long. Flowers unisexual or bisexual, the 

staminate flowers in branched axillary cymes, the 

pistillate or bisexual flowers in simple axillary 

cymes. Calyx of discrete sepals, oblong, concave, 

48. Family VALERIANACEAE 

1. VALERIANA 

ciliate, 1-2 mm long; stamens 5, ascendent; ovary 

ovoid, with two pubescent styles. Drupe fleshy, 

ovoid or almost globose, 6-10(14) mm long, 

orange, with persistent styles. 

Phenology: Flowering from February to May 

and fruiting from May to September. 


Distribution: In dry areas along the coast, and 

in the area of mogotes. Also on Culebra, Desecheo, 

Icacos, Vieques, St. Croix, St. John, and St. 

Thomas; throughout the Neotropics. 

Public Forest: In all probability, found in 

Guánica. 

Herbs or less frequently herbaceous vines, twining or scandent, glabrous or pubescent, usually 

with a fetid odor. Leaves opposite, simple, entire, dentate, trifoliolate, pinnate, or pinnatisect; stipules 

absent. Flowers actinomorphic or slightly zygomorphic, bisexual or rarely unisexual, produced in 

terminal or axillary dichasial cymes; bracts and bracteoles present. Calyx tubular, with 5-20 lobes, 

dentate, setose or plumose; corolla infundibuliform, campanulate, or hypocrateriform, the tube short, 

the lobes 5, expanded; stamens 3(4), adnate to the throat, included or exserted, the anthers sessile; 

ovary inferior, 3-carpellate, the carpels with a single ovule, the stigma 2-3-lobate. Fruit a dry indehiscent 

achene with a single fertile carpel, crowned by the plumose sepals. An almost cosmopolitan genus, of 

approximately 250 species. 

1. Valeriana scandens L., Sp. Pl. ed. 2, 47. 1762. 

Fig. 158 A-E 

Valeriana 


length. Stems cylindrical, glabrous or puberulent 

at the nodes. Leaves opposite, trifoliolate; leaflets 

1.3-6 × 0.8-2.2 cm (terminal leaflet larger than 

the lateral ones), membranaceous, glabrous, 

lanceolate or ovate-lanceolate, the apex 

acuminate, the base obtuse, rounded, or truncate, 

the margins entire or crenate, the venation slightly 

prominent on both surfaces; petioles slender, 1.2- 

3 cm long. Flowers in dichasial or 

pseudodichotomous cymes with lateral 

monochasia. Hypanthium ovoid, ca. 1 mm long, 

crowned by 10-12 filiform sepals; corolla narrowly 

campanulate, 1.3-1.8 mm long, the lobes 0.2-0.4 

mm long; stamens included; style ca. 1 mm long, 

with three short stigmatic branches. Achene 

flattened, ribbed, ovate, ca. 2.5 mm long, crowned 

by a tuft of plumose sepals, ca. 4 mm long 


November to May and in fruit in March. 


Distribution: In moist areas of the Cordillera 

Central and in the area of mogotes. Also in tropical 

continental America, Cuba, Hispaniola, and the 

United States (Florida). 

Public Forests: Guilarte and Río Abajo.


390 


1. Clerodendrum aculeatum (L.) Schltdtl., 

Linnaea 6: 750. 1831. 

Fig. 159. A-E 

BASIONYM: Volkameria aculeata L. 

Erect or scandent shrub attainig 3 m in length. 

Stems cylindrical, usually striate, puberulent, with 

2 or 3 spines, opposite or whorled, in the area of 

the nodes; stipules absent. Leaves opposite or three 

per node, 1.5-7 × 0.5-2.6 cm, oblong or ellipticovate, 

chartaceous, glabrous or puberulent, the 

apex acute or obtuse, the base cuneate or rounded, 

the margins entire; upper surface dull, dark green; 

lower surface pale green, dull, with numerous 

49. Family VERBENACEAE 


1a. Flowers sessile, subtended by an involucre of 3(4) elongate bracts, expanded, pink or lilac at the 

base, much longer than the calyx.………………………………..….……..2. Congea 

1b. Flowers with minute bracts, not covering the calyx……………………...………………..2 

2.a Calyx gamosepalous, acetabuliform, orange, the sepals not apparent ….….3. Holmskioldia 

2b. Calyx of free sepals or sepals connate only at the base, of various colors, but not orange…………....3 

3a. Inflorescences corymbiform; calyx green, white, or pink, the sepals imbricate……1. Clerodendrum 

3b. Inflorescences racemose; calyx violet, the sepals expanded .................……………….…....4. Petrea 

1. CLERODENDRUM 

Trees, erect shrubs, or less frequently lianas or scandent shrubs, sometimes with axillary spines. 

Leaves simple, opposite or whorled, petiolate, with entire or dentate margins. Flowers showy, bisexual, 

zygomorphic, in axillary cymes or terminal panicles. Calyx campanulate or tubular, truncate, 5-lobate; 

corolla 5-merous, hypocrateriform; stamens 4, the filaments didynamous, adnate to the base of the 

corolla, exserted; ovary superior, 4-locular, each locule with a solitary ovule; style slender, crowned by 

a globose stigma. Fruit drupaceous, 4-lobate, with 4 pyrenes. A genus of approximately 400 species, 

the majority native to the tropics of the Old World. 

Key to the species of Clerodendrum 

1a. Branches with conical spines, opposite or whorled, in the area of the nodes................1. C. aculeatum 

1b. Branches unarmed....…………………………………………………………………………2 

2a. Calyx reddish, 1-1.5 cm long …............……….………..……...……………...….. C. × speciosum 

2b. Calyx white, 1.5-2.5 cm long ........….........………………..………………..…………. C. thomsonae 

dots; petioles 2.5-10 mm long, puberulent. 

Inflorescences of axillary cymes. Calyx 

campanulate, puberulent, 2.5-4 mm long, the 

sepals expanded; corolla white, hypocrateriform, 

1.4-2.7 cm long, the tube sometimes with a purple 

tinge; filaments pink, twice as long as the corolla; 

style purple, as long as the filaments. Drupe ovoid 

or depressed-ovoid, 5-7 mm long, opening in two 

halves when ripe. 


to March and from August to October and in fruit 

from January to August. 

Status: Native, locally common.


Distribution: In dry forests and thickets, along 

the littoral zone. Also on Mona, Anegada, Jost 

van Dyke, St. Croix, St. John, St. Thomas, Tortola, 

and Virgin Gorda; throughout the Antilles, 

Venezuela, and the Guianas. 

Public Forests: Guánica and Mona. 

2. Clerodendrum × speciosum Dombrain, Flor. 

Mag. 8, pl. 432. 1869. 

Fig. 160. E-G 

SYNONYMS: Clerodendrum thomsonae f. speciosum 

(Dombrain) Voss 

Clerodendrum umbellatum var. speciosum (Dombrain) 

Moldenke 


length. Stems obtusely quadrangular, striate,

392 


minutely puberulent; stipules absent. Leaves 

opposite, 7.5-15.5 × 5.7-8 cm, elliptical or ovate, 

chartaceous, the apex acuminate, the base rounded 

or cordiform, the margins entire or remotely 

sinuate; upper surface glabrous; lower surface 

glabrous, pale green, with numerous dots, the 

veins prominent and minutely puberulent; petioles 

1-2 cm long, sulcate, puberulent. Inflorescences 

of axillary dichasial cymes; bracts minute, 

subulate. Calyx more or less urceolate, 1.0-1.5 

cm long, intense pink to purple, puberulent, the 

sepals lanceolate, connate at the base, acuminate 

at the apex; corolla red or red-orange, 

hypocrateriform, 2.5-3 cm long, the tube quite 

narrow, the lobes rounded; filaments pink, twice 

as long as the corolla; style pink, as long as the 

filaments. Fruit unknown. 


September to February. 

Status: Exotic, cultivated, persistent and 

naturalized in some localities in Puerto Rico. 

Distribution: Common in gardens in Puerto 

Rico and the Virgin Islands. Cultivated 


Public Forests: Río Abajo and Tortuguero. 

Commentary: Discrepancies exist regarding 

the nature of this taxon. Originally (1869), it was 

described as a hybrid under the name of C. x 

speciosum. Later, Voss (1894) considered this 

taxon as a form of C. thomsonae and Moldenke 

(1937) as a variety of C. umbellatum Poir. 

Moldenke, in 1983, changed his opinion, 

recognizing it as a form of C. thomsonae, but 

pointed out that the studies necessary to discard 

its hybrid nature do not yet exist. Due to the fact 

2. CONGEA 

that the nature of this taxon has not been 

established with certainty, it seems to me 

appropriate to utilize the name of C. x speciosum, 

this being the most widely used name. 

3. Clerodendrum thomsonae Balf. , Edinburgh 

New Philos. J. 15(2): 233, t. 2. 1862. 

Fig. 160. A-D 


in length. Stems obtusely quadrangular, 

puberulent; stipules absent. Leaves opposite, 5.2- 

14 × 2.7-7 cm, elliptical or lanceolate, 

chartaceous, the apex acuminate, the base obtuse 

or rounded, the margins entire or remotely 

sinuate; upper surface puberulent, dark green, 

dull; lower surface puberulent, pale green, with 

numerous dots, the veins prominent; petioles 1- 

2.8 cm long, sulcate, puberulent. Inflorescences 

of axillary dichasial cymes; bracts minute, 

subulate. Calyx more or less urceolate, 1.5-2.5 

cm long, white, puberulent, the sepals lanceolate 

or elliptic-lanceolate, connate at the base, 

acuminate at the apex; corolla red or crimson, 

hypocrateriform, ca. 2 cm long, the tube quite 

narrow, the lobes rounded; filaments pink, twice 

as long as the corolla; style pink, as long as the 

filaments. Fruit unknown. 


Status: Exotic, cultivated. 


Rico and the Virgin Islands. Cultivated 


Lianas or clambering shrubs, without spines, pubescence of simple or stellate hairs. Leaves simple, 

opposite, petiolate, with entire margins; stipules absent. Flowers bisexual, zygomorphic, in head-like 

cymes, arranged in terminal panicles. Calyx ovoid-campanulate, 5-dentate; corolla 5-merous, tubular, 

bilabiate, the upper lip bilobate, the basal lip trilobate, shorter than the upper lip; stamens 4, exserted, 

the filaments didynamous, adnate to the corolla below the throat; ovary superior, imperfectly bilobate, 

each locule biovulate; style slender, as long as the filaments, stigma bifid. Fruit dry, obovoid, with a 

single seed. A genus of 10 species, native to southeastern Asia. 

1. Congea tomentosa Roxb., Pl. Coromandel. 3: 

90. 1820. 

Fig. 161. A-C 

Lluvia de orquídeas, Terciopelo 

Twining liana, with numerous lateral 

branches, short, divaricate, usually pendulous, 


sericeous-tomentose; stipules absent. Leaves


394 


opposite, 5-13(18.5) × 3-7.5(9.5) cm, ellipticovate 

or ovate, coriaceous, the apex shortacuminate, 

acute, or obtuse, the base rounded or 

subcordiform, sometimes asymmetrical, the 

margins entire or remotely sinuate; upper surface 

puberulent, dark green, dull; lower surface whitetomentose, 

with the venation quite prominent; 

petioles 5-13 mm long, lanate-tomentose. 

Inflorescences at the end of the short lateral 

branches, forming an inflorescence with a marked 

transition from leaves to bracts; bracts at the base 

of the branches of the inflorescence ca. 1 cm long, 

elliptical; axes lanate-tomentose, glabrescent; 

bracts of the involucre 3(4), expanded or reflexed, 

connate at the base, 2.3-4 × 0.8-1.4 cm, elliptical 

or oblong, pink or lilac toward the apex, green at 

the base, white-tomentose, with prominent 

venation. Flowers sessile, in groups of (5)7(9). 

3. HOLMSKIOLDIA 

Calyx infundibuliform, 5-7 mm long, light green, 

white-lanate-tomentose; corolla tubular, bilabiate, 

slightly longer than the calyx, white, glabrous, 

the upper lobes oblong, ca. 3 mm long, the basal 

lobes shorter and rounded; filaments longexserted, 

reddish brown or purple; ovary obovoid, 

glabrous, the style as long as the filaments. Fruit 

unknown. 


to March. 


although not very common. 


Rico and the Virgin Islands, and naturalized in 

disturbed vegetation in the Cordillera Central. 

Native to Indomalasia. Cultivated throughout the 

tropics. 

Erect or clambering shrubs, glabrous or pubescent, without spines or with axillary spines. Leaves 

simple, opposite, petiolate, with entire or dentate margins; stipules absent. Flowers bisexual, 

zygomorphic, in axillary cymes or racemes or congested and terminal. Calyx gamosepalous, rotate or 

acetabuliform, truncate or sinuate-dentate, usually of brilliant colors; corolla tubular, more or less 

bilabiate, the tube curved, the limb with 5 lobes; stamens 4, exserted, the filaments didynamous, connate 

to the corolla in the middle portion of the tube; ovary superior, 4-locular, each locule with a solitary 

ovule, the style elongate, exserted, the stigma bifid. Fruit drupaceous, 4-lobate, subtended by the 

accrescent calyx, with 4 pyrenes inside. A genus of 19 species, native to southern Asia, southeastern 

Africa, and Madagascar. 

1. Holmskioldia sanguinea Retz., Obs. 6: 31. 

1791. 

Fig. 161. D-E 

Paraguita chino, Platillo, Parasol 

Scandent or erect shrub, with numerous basal 

branches, attainig 2 m in length. Stems obtusely 

quadrangular or cylindrical, puberulent, grayish; 

stipules absent. Leaves opposite, 3-12 × 1.5-8.5 

cm, ovate or ovate-lanceolate, chartaceous, the 

apex acuminate or acute, the base cuneate or 

truncate, the margins serrate; upper surface 

puberulent, dark green, dull; lower surface 

puberulent, pale green, with numerous dots, the 

venation prominent; petioles 8-30 mm long, 

puberulent. Inflorescences of racemes or less 

frequently panicles, axillary or terminal, up to 5 

cm long; bracts minute; pedicels 2.5-8 mm long, 

slender, glandular-pubescent. Calyx 

acetabuliform, orange, 1.6-2.2 cm in diameter, 

with minute dots; corolla hypocrateriform, 

crimson, 1.5-2.5 cm long; stamens slightly 

exserted, the filaments pink; ovary glabrous, the 

style as long as the filaments. Fruit globose, 

brown, verrucose. 



Status: Exotic, cultivated, rather common, 

apparently naturalized in areas of the Cordillera 

Central. 


Rico. Native of southern Asia. Cultivated 

throughout the tropics.


396 


4. PETREA 

Tress, shrubs, or twining vines, without spines, glabrous or pubescent. Leaves simple, opposite, 

subopposite, or whorled, usually scabrous, petiolate, with entire or slightly crenate margins; stipules 

absent. Flowers bisexual, in axillary or terminal racemes. Calyx with a short cylindrical or campanulate 

tube and 5 deep lobes, elongate; corolla, hypocrateriform, blue, violet, or less frequently white, the tube 

urceolate, cylindrical, or infundibuliform, the lobes 5, expanded; stamens 4, the filaments didynamous, 

connate on the middle portion of the corolla; ovary superior, with a single fertile carpel, bilocular, each 

locule with a solitary ovule; style terminal, the stigma capitate. Fruit drupaceous, with 2 pyrenes, 

completely covered by the calyx. A genus of 11 species, native to the Neotropics. 

1. Petrea volubilis L., Sp. Pl. 626. 1753. 

Fig. 161. F-H 

Twining liana or shrub, scandent or less 

frequently erect, with numerous basal branches, 

attainig 2-4 m in length. Stems obtusely 

quadrangular, pubescent or glabrous; stipules 

absent. Leaves opposite, sometimes whorled, 3- 

25 × 1.5-11 cm, elliptical, chartaceous, the apex 

obtuse, rounded-acute, or short-acuminate, the 

base obtuse or rounded, sometimes slightly 

asymmetrical, the margins entire, crenate, or 

sometimes serrate; upper surface glabrous or 

puberulent, scaberulous, with prominent venation; 

lower surface puberulent or glabrous, pale green, 

scaberulous, with numerous resinous dots, the 

References: Abid, M. A., 1966. A revision of Congea (Verbenaceae). The Garden’s Bulletin Singapore. 

21: 259-314. Moldenke, H. N. and A. L. Moldenke. 1983. Verbenaceae. In: Dassanayake and Fosberg, 

(eds.). Flora of Ceylon. Vol. 4: 196-487. Rueda, R. M. 1994. Systematics and evolution of the genus 

Petrea (Verbenaceae). Ann. Missouri Bot. Gard. 81: 610-652. 

50. Family VITACEAE 


venation prominent; petioles 4-16 mm long, 

puberulent. Inflorescences of axillary racemes, 10- 

60 cm long; bracts minute; pedicels 7-18 mm 

long, slender, puberulent. Calyx infundibuliform, 

blue-violet or white, the sepals 1.6-2 cm long, 

connate at the base to form a tube, 3-7 mm long, 

glabrous or pubescent; corolla infundibuliform, 

blue-violet or white, 0.6-1.5 cm long, the limb 5lobate, 

one of the lobes larger; stamens included; 

ovary glabrous, the style short. Fruit unknown. 





Rico and the Virgin Islands. Native to tropical 

continental America. Cultivated throughout the 

tropics. 

1a. Stems generally herbaceous and fleshy; inflorescences ascendent or expanded toward the sides, as 

broad as long or broader than long; flowers 4-merous; petals persistent after anthesis 

..............................................…………………...............................................................1. Cissus 

1b. Stems woody when mature; inflorescences pendulous, much longer than broad; flowers 5-merous; 

petals concrescent at the apex, forming a deciduous aggregate during anthesis 

…………………..............................................………………...........................................2. Vitis


1. CISSUS 

Vines, usually fleshy, that climb by means of tendrils opposite the leaves. The stems fleshy or 

slightly woody, without anatomical anomalies, producing scarce watery latex when cut. Leaves simple 

or trifoliolate, alternate, with serrate or entire margins; petioles more or less long; stipules small and 

deciduous. Flowers 4-merous, actinomorphic, usually bisexual, produced in compound cymes, opposite 

the leaves; calyx campanulate, with 4 minute lobes; petals 4, reflexed during anthesis; stamens alternating 

with the petals; disc cupuliform, entire or lobate; ovary superior, bilocular, with a robust and elongate 

style. Fruit a fleshy berry, globose, ovoid, or depressed-globose. A pantropical genus of 350 species. 

Key to the species of Cissus 

1a. Leaves simple…………………………………………………………………………..……….2 

1b. Leaves trifoliolate……………………………………………………………………………….3 

2a. Leaves ovate, coriaceous ……………….……………………………….….….. 5. C. verticillata 

2b. Leaves elliptical, fleshy ………………………………………………………. 3. C. rotundifolia 

3a. Stems angular, 4-6-winged; terminal leaflet 4.3-15 cm long, usually elliptical, lanceolate, or ovate 

…..…………………………..……………..……...………………………….........… 1. C. erosa 

3b. Stems cylindrical, sometimes sulcate; terminal leaflet 1.6-4 cm long, rhombic or obovate….... 4 

4a. Terminal leaflets obovate, obtuse, or rounded at the apex, the margins crenate or scattered-dentate; 

flowers red…......…..……………………………………………................………. 2. C. obovata 

4b. Terminal leaflets rhombic, acute at the apex, the margins sinuate-dentate on the upper half; flowers 

yellowish green ………………...………………………………………................ 4. C. trifoliata 

1. Cissus erosa L.C. Rich., Act. Soc. Hist. Nat. 

1: 106. 1792. 

Fig. 162. A-J 

Caro de tres hojas 

Non-woody vine, that climbs by means of 


angular, quadrangular, or 4-6-winged, branched, 

glabrous or puberulent, the nodes swollen. Leaves 

alternate, trifoliolate; leaflets elliptical, lanceolate, 

or ovate, 4.3-10.5 × 3-5.5 cm (the terminal leaflet 

larger), glabrous or puberulent on the prominent 

veins, the apex acute or obtuse, the base acute, 

obtuse, or attenuate on the central leaflet and 

unequal on the lateral leaflets, the margins serrate; 

petiolules very short or absent; petioles 

canaliculate or slightly winged, 2-3(5) cm long; 

stipules deltoid, auriculate at the base, 2-4 mm 

long. Tendrils opposite the leaves, simple, 

filiform, 10-12 cm long, sometimes terminating 

in adventitious discs. Inflorescences of 

umbelliform cymes, opposite the leaves; peduncle 

longer than the accompanying leaf; pedicels 2- 

2.5 mm long, reddish. Calyx ca. 1 mm high, red 

or pink, truncate; petals red, glabrous, papillose; 

stamens green; disc reddish. Fruits globose, 5-7 

mm in diameter, black when ripe. Seed one, 5-7 

mm long. 



Distribution: In disturbed areas on roadsides, 

on fences, and in pastures at lower and middle 

elevations. Also on Anegada; Hispaniola and 



Río Abajo, and Tortuguero. 

2. Cissus obovata Vahl, Symb. Bot. 3: 19. 1794. 

Fig. 163. A-D 

SYNONYMS: Cissus caustica Tussac 

Cissus tuberculata Jacq. 


tendrils and attains 3-15 m in length. Stems much

398 



branched, subcylindrical, angular, fleshy, 3-8 mm 

in diameter, glabrous or puberulent, reddish and 

then grayish, flexuous, with the nodes swollen. 

Leaves alternate, trifoliolate, chartaceous, 

glabrous or pubescent; leaflets obovate, almost 

rounded, or elliptical, 0.7-6 × 0.5-3.5 cm (the 

terminal leaflet larger), the apex rounded, obtuse, 

mucronate and sometimes retuse, the base cuneate 

to rounded, unequal on the lateral leaflets, the 

margins crenate or scattered-dentate on the distal 

portion; petioles 0.8-5 cm long; stipules deltoid, 

early deciduous; tendrils opposite the leaves, 

minute, bifurcate, with an adventitious disc at the 

end. Inflorescences of cymes, opposite the leaves, 

with the peduncles glabrous or puberulent, 

reddish; pedicels reddish. Calyx reddish, ca. 1 mm 

long. Petals 4, reddish, 1-2 mm long, triangular, 

deciduous. Fruits globose to globose-obovoid, 4- 

5 mm in diameter, black or purple when ripe. Seed 

one, ca. 5 mm long. 

Phenology: Collected in flower in June and 

November and in fruit in July. 


Distribution: At lower and middle elevations 

in coastal areas and in the zone of mogotes. Also 

on Mona, St. Croix, St. John, and St. Thomas; 

Hispaniola and the Lesser Antilles. 


Guánica, Mona, and Río Abajo. 

3. Cissus rotundifolia (Forssk.)Vahl, Symb. Bot. 

3: 19. 1794. 

Fig. 163. E-F 

BASIONYM: Saelanthus rotundifolius Forssk. 


tendrils and attains more than 5 m in length. 

Stems fleshy, with abundant watery latex, more 

or less cylindrical, striate, with malpighiaceous 

hairs on the new growth, glabrescent. Leaves 

alternate, fleshy, oval, glabrous, 4-6.5 × 4-6 cm, 

involute, the apex truncate to almost rounded, the 

base cordiform, the margins serrate; petioles 0.8- 

1.2 cm long, sulcate; stipules ca. 2 mm long, 

oblong-lanceolate, early deciduous; tendrils 

opposite the leaves, thick, simple, slightly longer 

than the leaves, twisting in the form of a spiral. 

Inflorescences cymose, opposite the leaves, with 

the flowers grouped in umbels; pedicels 4-10 mm 

long, green. Calyx green, rotate or discoid, 1.5- 

1.8 mm wide; petals 4, pale green or whitish 

green, oblong-triangular, 2-2.5 mm long, 

deciduous; disc short-cupular, ca. 0.3 mm high; 

ovary sinuate-lobate at the base. Fruits globose, 

fleshy, ca. 0.8-1 cm in diameter, greenish. 

Phenology: Collected in flower in September 

and in fruit in April and September. 



Distribution: Cultivated in the Virgin Islands; 

escaped on St. Croix. Native to Yemen and Africa, 

but cultivated as an ornamental throughout the 

tropics. 

4. Cissus trifoliata (L.) L., Syst. Nat. ed. 10, 897. 

1759. 

Fig. 164. E-F 

BASIONYM: Sicyos trifoliata L. 

SYNONYM: Cissus acida L. 

Bejuco de caro, Sorrel vine 

Non-woody vine that climbs by means of 

tendrils and attains 2-15 m in length. Stems much 

branched, cylindrical, fleshy, 3-8 mm in diameter, 

glabrous or puberulent, reddish brown or grayish 

when mature, the nodes swollen. Leaves alternate, 

trifoliolate, fleshy, glabrous; leaflets ovate, 

obovate, or oblong, 2-3 × 1-2.6 cm, the apex 

rounded, acute, or truncate, the base cuneate or 

attenuate, the margins deeply dentate-mucronate 

on the upper half; upper surface and lower surface 

dull; petioles fleshy, sulcate, 1-3 cm long; stipules 

3.5-5 mm long, lanceolate; tendrils opposite the 

leaves, simple, filiform, 6-8 cm long, twisted in 

the form of a spiral. Inflorescences in cymes, 

opposite the leaves. Calyx yellowish green, 1.5-2 

mm long. Petals 4, yellowish or whitish, reflexed, 

2-2.5 mm long, lanceolate-triangular, deciduous. 

Fruits depressed-globose, 7-8 mm in diameter, 

shiny, metallic blue or black when ripe. 

Phenology: Flowering from March to July and 

fruiting from July to August. 


Distribution: At lower and middle elevations 

in coastal areas. Also on Caja de Muerto, Cayo 

Ratones, Mona, St. Croix, St. John, St. Thomas, 

and Tortola; the Bahamas, Cuba, Jamaica, 

Hispaniola, Colombia, Venezuela, and Ecuador. 

Public Forests: Guánica, Mona, and 

Tortuguero.

400 



5. Cissus verticillata (L.) Nicolson & Jarvis, 

Taxon 33: 727. 1984. 

Fig. 164. A-D 

BASIONYM: Viscum verticillatum L. 

SYNONYM: Cissus sicyoides L. 

Bejuco de caro, Lambrali, Pinna koop, Puddin 

vine 


tendrils and attains more than 10 m in length. 

Stems fleshy, with abundant watery latex, 

cylindrical when young, flattened when mature, 

attaining 5 cm in diameter, the nodes swollen. 

Leaves alternate, coriaceous, ovate, 5-12 × 3.8- 

6.5 cm, the apex acute or rounded, the base 

cordiform, the margins revolute, denticulate; 

upper surface green, shiny; lower surface green, 

dull, with prominent venation; petioles 2-5 cm 

long, sulcate; stipules 2.5-3.5 mm long, 

auriculate; tendrils opposite the leaves, simple or 

bifurcate, up to 25 cm long, twisting in the form 

of a spiral. Inflorescences of compound cymes that 

are borne opposite the leaves; pedicels ca. 3 mm 

long, yellowish green or reddish. Calyx yellowish 

green or reddish, 0.7-1 mm long; petals 4, 

yellowish or pink, oblong-lanceolate, 2-2.5 mm 

long, deciduous; disc annular, yellow, 0.5-0.8 mm 

high. Fruits globose, fleshy, 0.7-1 cm in diameter, 

shiny, dark violet or black, with one or two seeds 

inside. 





pastures and roadsides, at middle and lower 

elevations. Also on Culebra, Desecheo, Mona, 

2. VITIS 


and Virgin Gorda; from southern North America 

to northern South America, including the Antilles. 

Public Forests: Cambalache, Carite, Ceiba, El 

Yunque, Guánica, Maricao, Mona, Río Abajo, 

Susúa, and Toro Negro. 

Doubtful Species: 

Cissus intermedia A. Rich. was reported for 

Puerto Rico by Urban (1903-1911), based on the 

collections Stahl 602 and Sintenis 4797. The 

collection of Stahl has been re-identified by 

Lombardi (2000) as C. obovata. On the other 

hand, the collection of Sintenis (deposited in the 

herbarium of Berlin) seems to have been destroyed 

during the Second World War. Since the existence 

of any other specimen that might be referred to 

C. intemedia is unknown, I consider that this 

species is not found in Puerto Rico. Its distribution 

is limited to the Bahamas, Cuba, and Hispaniola. 

Commentary: Several species of Cissus (C. 

erosa, C. trifoliata, and C. verticillata) are 

affected by the parasitic fungus Mycosyrinx cissi 

(Poiret) G. Beck, which causes the inflorescences 

to grow in a monstrous form, producing numerous 

sterile branches. These deformed inflorescences 

look like a parasitic plant of the family 

Loranthaceae or Viscaceae. For this reason, the 

name C. verticillata has replaced the well known 

name C. sicyoides. It turns out that this species 

was first described by Linnaeus based on one of 

these deformed inflorescences, under the name 

of Viscum verticillatum. This basionym has 

priority over C. sicyoides, since it is older, and 

therefore it has replaced C. sicyoides. 

Lianas or woody vines, that climb by means of tendrils that are opposite the leaves. The stems 

woody, without anatomical anomalies, producing abundant water when cut. Leaves simple, usually 

palmately lobed, alternate, with serrate margins; petioles more or less elongate; stipules deciduous. 

Flowers 5-merous, actinomorphic, bisexual or unisexual, produced in panicles, opposite the leaves; 

calyx cupuliform, with 5 minute lobes; corolla of 5 petals, concrescent at the apex, that separate as a 

unit in anthesis; stamens alternate with the petals; disc cupuliform, 5-lobate; ovary superior, bilocular, 

the style short, conical. Fruit a fleshy berry, ellipsoid or globose. A genus of approximately 60 species, 

the majority of the Northern Hemisphere. The grape belongs to this genus.

402 



1. Vitis tiliifolia Roem. & Schult., Syst. Veg. 5: 

320. 1819. 

Fig. 164. G-J 

SYNONYM: Vitis caribaea DC. 


Liana that climbs by means of tendrils and 

attains 10-35 m in length. Stems cylindrical, 

woody, producing abundant potable water when 

cut and attaining 8 cm in diameter; bark dark 

brown, rough. Leaves simple, broadly ovate or 

trilobate, 6.5-14 × 6.5-12.5 cm, chartaceous, the 

apex acute to short-acuminate, the base cordiform 

to almost truncate, the margins serrate; upper 

surface puberulent; lower surface tomentose; 

petioles 2.5-8 cm long; stipules caducous. 

Inflorescences of pendulous axillary panicles, 9- 

17 cm long. Calyx green, discoid, the sepals ca. 2 

mm long. Petals 1.5-2.2 mm long, greenish 

yellow, oblong-obovate, persistent; disc ca. 0.2 

mm high. Berries globose, 5-10 mm in diameter, 

juicy, violet when ripe. Seeds 1-2, circular, ca. 4 

mm long. 

Phenology: Collected in flower during March 

and April and in fruit in September. 


Distribution: In moist forested areas, at middle 

elevations. Also on St. Croix, St. John, and St. 

Thomas; throughout the Antilles and from Mexico 

to northern South America. 


Abajo. 

Doubtful Genus: 

Ampelopsis arborea (L.) Koehne was treated 

by Liogier (1994) as present in Puerto Rico, based 

on Howard (1989), who included Puerto Rico in 

the distribution of the species. This report has not 

been confirmed, and it seems that, if it is found 

in our gardens, it is not very common; therefore, 

it is not included in the present work. 

Reference: Lombardi, J.A., 2000. Vitaceae. Gêneros Ampelocissus, Ampelopsis e Cissus. Flora 

Neotropica Mon. 80: 1-250. 

MONOCOTYLEDONS 

Key to the families 

1a. Plants climbing by means of adventitious aerial roots………………………………………….2 

1b. Plants twining, clambering, or climbing by means of tendrils……………………………...…..3 

2a. Flowers minute, < 3 mm long, produced in spadices, subtended by a spathe (foliaceous bract) 

……………………………………………..………….…..…………………………..... Araceae 

2b. Flowers showy, > 5 cm wide, solitary or in racemes……………...............….…... Orchidaceae 

3a. Plants like a fern, with verticillate phylloclades (needle-like) ..……...……….… Asparagaceae 

3b. Plants not like a fern, with true leaves whose blade is not needle-like……………..…………..4 

4a. Leaves petiolate (not forming a leaf sheath or this not enclosing the stem); blades ovate, rounded to 

cordiform at the base, with 5-7 parallel-arcuate main veins that are borne from the base (or nearly 

so).............................................................................................................................5 

4b. Leaves not petiolate, with a leaf sheath that encloses the stem; blades oblong, lanceolate, or rarely 

ovate, not cordiform at the base, with a single main vein…………………….………….6 

5a. Plants twining, lacking tendrils; fruits capsular or samaroid ..……………...….. Dioscoreaceae 

5b. Plants climbing by means of a pair of tendrils that are borne from the base of the petiole; fruits 

fleshy, almost globose………………………………….…………………..……… Smilacaceae

404 


6a. Stems triangular; leaf blades usually with cutting margins; leaf sheath closed; flowers subtended by 

a single scale ................………………………………………………………... Cyperaceae 

6b. Stems cylindrical; leaf blades with margins not cutting; leaf sheath open; flowers subtended by a 

pair of scales ………………………………….……………………………………... Poaceae 

1. Family ARACEAE 


1a. Flowers bisexual……………………………………………………………….………………..2 

1b. Flowers unisexual………………………………………………………………………………4 

2a. Spathes chartaceous and reflexed, the spadix completely exposed; stems up to 5 mm in diameter; 

leaves entire, green, not perforated ………….………………….............1. Anthurium 

2b. Spathes fleshy and erect, enclosing the spadix; stems up to 2 cm in diameter; leaves pinnately 

compound, or entire and variegated, or entire and perforated, not variegated………………….3 

3a. Leaves entire and variegated (green and yellow) or green and pinnately compound, not perforated; 

ovary unilocular; seeds reniform …..………………...……………...2. Epipremnum 

3b. Leaves with entire margins, the blade with large perforations; ovary bilocular; seeds ovoid or 

cylindrical ..………………………………….……...................................................3. Monstera 

4a. Leaves entire, cordiform or elliptical, or pinnately compound ....…………..….4. Philodendron 

4b. Leaves digitate ............................................................…...........……...........……...5. Syngonium 

1. ANTHURIUM 

Terrestrial or epiphytic herbs, erect or rarely climbing by means of adventitious roots; stems fleshy, 

elongate or very reduced. Leaves alternate, simple or digitate; petioles elongate, covered by a cataphyll 

in the juvenile stages. Spathe usually chartaceous, reflexed, usually perennial, green, white, pink, or 

red; spadix sessile or short-stipitate, cylindrical or conical, with numerous flowers from the apex to the 

base. Flowers bisexual, sessile; tepals 4; stamens 4; ovary bilocular, with 1 or 2 pendulous ovules per 

locule, the style short or absent, the stigma discoid or lobate. Fruit a bilocular berry, fleshy, red, white, 

or violet. Seeds oblong. A neotropical genus of about 700 species. 

1. Anthurium scandens (Aubl.) Engler in Mart., 

Fl. Bras. 3(2): 78. 1878. 

Fig. 165. A-F 

BASIONYM: Dracontium scandens Aubl. 

Herbaceous epiphytic vine, that climbs by 

means of adventitious roots and attains 1-2 m in 

length. Stems cylindrical, the internodes covered 

by a fibrous leaf sheath. Leaves simple, alternate, 

lanceolate-elliptical or elliptical, 7-13 × 2.5-5 cm, 

chartaceous, the apex acuminate, the base obtuse 

or attenuate, the margins entire; upper surface 

glabrous, foveolate; lower surface glabrous, with 

numerous dark dots; petioles 0.7-4.5 cm long; leaf 

sheath stramineous, fibrous, persistent, ca. 3 cm 

long. Inflorescences axillary, erect; spadix in


flower 1.5-2 cm long; spathe reflexed, 

membranaceous, green, with the venation 

mulberry-colored, 1-1.2 cm long. Tepals 4, green 

or brown, ca. 1 mm long, cuneiform, involute; 

anthers stramineous; ovary green. Berries 

depressed-globose, ca. 5 mm in diameter, juicy, 

white when ripe. Seeds ellipsoid, ca. 1 mm long. 

Phenology: Flowering and fruiting probably 



Distribution: Epiphytic herb in moist or rain 

forests. Of widespread distribution in the 

Neotropics. 


Maricao, and Río Abajo.

406 


2. EPIPREMNUM 

Robust vines, climbing by means of adventitious roots. Leaves alternate; blades simple, entire or 

pinnately compound; petioles pulvinate at the apex, the sheath forming a winged margin that usually 

decomposes into fibers. Spathe thick-coriaceous, straight, enclosing the spadix, deciduous; spadix 

cylindrical, thick, erect. Flowers bisexual, lacking a perianth; stamens 4; pistil prismatic, truncate at 

the apex, the ovary unilocular, with 2-8 basal ovules, the stigma linear. Berries fleshy, yellowish. Seeds 

reniform. A genus of 8 species, native to the area between southeastern Asia and the western Pacific, 


Key to the taxa of Epipremnum 

1a. Leaves green, not variegated, pinnately compound or the juvenile leaves entire; stems and petioles 

green; usually producing flowers and fruits ……………...……...……...1. E. pinnatum 

1b. Leaves variegated (green-yellow), entire or less frequently pinnately compound; stems and petioles 

yellow-orange; rarely in fertile condition ….…………………2. E. pinnatum ‘Aureum’ 

1. Epipremnum pinnatum (L.) Engler, Pflanzenr. 

IV (23B) 37: 60. 1908. 

Fig. 166. A-C 

BASIONYM: Pothos pinnata L. 

SYNONYM: Philodendron nechodomi Britt. 

Non-woody vine, glabrous, that climbs by 

means of aerial roots and attains 6-8 m in length. 

Stems flexible, cylindrical, green, with a 

papyraceous epidermis, up to 3 cm in diameter, 

producing scarce watery latex when cut. Leaves 

alternate, chartaceous, dimorphic, the juvenile 

leaves entire, the adult leaves pendulous, pinnately 

compound, up to 50 x 30 cm, the apex terminating 

in a leaflet or acuminate segment, the base 

asymmetrical, subcordiform; upper surface dull, 

glabrous, the venation flat; lower surface dull, 

with prominent venation, with a line of domatia 

on both sides of the midvein, this sometimes with 

small spiniform outgrowths; petioles up to 35 cm 

long, marginate along their length. Inflorescence 

axillary, solitary, erect; spathe almost cylindrical, 

up to 18 cm long, opening along one side, fleshy, 

yellowish, acuminate at the apex, deciduous; 

spadix sessile, more or less cylindrical, 12-15 cm 

long. 


January and February. 

Status: Exotic, cultivated although 

uncommon. 

Distribution: Cultivated in Puerto Rico, 

naturalized on St. Thomas. Native to southeastern 

Asia and Oceania. 

2. Epipremnum pinnatum ‘Aureum’ Nicolson, 

Allertonia 1: 347. 1978. 

Fig. 166. D-E 

SYNONYMS: Epipremnum aureum (Lindl. & André) 

Bunting 

Pothos aurea André 


This cultivated form is usually much more 

robust than the wild species, attaining more than 

10 m in length. It is also distinguished from the 

wild species by having yellow-orange stems and 

petioles and by the variegated (white or yellow) 

and predominantly entire leaves. The size of the 

leaves in the cultivated plants tends to be much 

smaller (7-10 cm long), while plants that have 

escaped from cultivation or have become 

naturalized have leaves with blades that attain 35 

cm long. This cultivated race seems to be sterile, 

since its fruits are not known. 

Phenology: Usually sterile, collected only once 

in flower in Puerto Rico. 


rather common.


408 


Distribution: Along roads and on abandoned 

farms. Known only in cultivation or naturalized, 


3. MONSTERA 

Public Forests: El Yunque, Río Abajo, and 

Toro Negro. 

Robust or slender vines, climbing by means of adventitious roots; stems smooth or verrucose, 

elongate. Leaves alternate; blades simple, entire or laciniate, usually perforated; petioles usually elongate 

and winged. Spathe very thick, convolute, enclosing the spadix, deciduous; spadix cylindrical, thick, 

erect. Flowers bisexual, lacking a perianth; stamens 4; pistil prismatic, truncate at the apex, the ovary 

bilocular, with two basal ovules in each locule, the stigma punctate or linear. Berries fleshy, green, 

white, yellow, or orange. Seed usually one, ovoid or cylindrical. A genus of 22 species, native to tropical 

continental America and the Lesser Antilles. 

1. Monstera adansonii Schott, Wiener Zeitschr. 

Kunst 1830: 1028. 1830. 

Fig. 167. A-C 



Stems flexible, cylindrical, green, 0.5-2.5 cm in 

diameter, producing scarce watery latex when cut. 

Leaves alternate, dimorphic, chartaceous, entire, 

asymmetrically elliptic-lanceolate, the basal 

leaves not perforated, the distal leaves with 

numerous perforations, 26-47 x 13-22.5 cm, the 

apex short- or long-acuminate, the base 

asymmetrical, cuneate-obtuse; upper surface dull, 

4. PHILODENDRON 

with a flat, broad midvein; lower surface pale 

green, with the midvein prominent and the 

secondary venation darker; petioles winged almost 

to the apex, shorter than the blade. Inflorescence 

axillary, solitary, erect; spathe almost cylindrical, 

up to 20 cm long, opening along one side, green 

outside, cream-colored on the inner surface. 



uncommon. 

Distribution: Cultivated in gardens and 

naturalized in areas of disturbed vegetation. 

Native to tropical continental America. 

Public Forests: El Yunque. 

Robust or slender vines, climbing by means of adventitious roots, or less frequently erect herbs; 

stems elongate, producing abundant watery and caustic sap when wounded. Leaves alternate; blades 

simple, lobate, divided or pinnately compound; petioles usually elongate. Spathe convolute, enclosing 

the spadix, usually thickened and not adhering to the spadix; spadix cylindrical, erect, usually almost 

sessile, with pistillate flowers on the basal portion. Flowers unisexual, lacking perianth; stamens 2-6; 

ovary usually bilocular, the locules with one or more ovules, the stigma sessile, entire or lobate. Fruits 

fleshy, with one or more seeds inside. Seeds ovoid or ellipsoid. About 275 species, native to the Neotropics, 

of which the majority are climbing. 

Key to the species of Philodendron 

1a. Petioles winged or marginate almost to the apical portion ….………...…….….4. P. lingulatum 

1.b Petioles not winged or with the wing only on the lower half…………………………...………2 

2a. Blades of the leaves 25-60 cm long; petioles 40-100 cm long; cataphylls persistent……...…...3 

2b. Blades of the leaves 13.5-35 cm long; petioles 10-35 cm long; cataphylls deciduous. ………..4


410 


3a. Stems 8-10 cm in diameter; spathe short-stipitate, ca. 45 cm long, thick, green on the outer surface, 

white on the inner surface …………………………………………...…2. P. giganteum 

3b. Stems 2-2.5 cm in diameter; spathe ca. 18 cm long, long-stipitate, moderately thick, whitish on the 

outer surface, maroon on the inner surface ...…………………………….……5. P. ornatum 

4a. Leaves oblong or lanceolate ...…………………………………………..…1. P. consanguineum 

4b. Leaves cordiform ……….........…………........................….....…………….…3. P. hederaceum 

1. Philodendron consanguineum Schott, Syn. 

Aroid. 88. 1856. 

Fig. 168. A-B 

SYNONYMS: Philodendron krebsii Schott 

Philodendron angustatum Schott 

Philodendron marginatum Urb. 

Bejuco de calabazón, Rasca garganta 


means of aerial roots and attains 10-30 m in 

length. Stems flexible, cylindrical, with an 

annular scar in the area of the node, attaining up 

to 2 cm in diameter and producing scarce watery 

and caustic latex when wounded; bark smooth, 

dark brown or grayish, membranaceous, 

exfoliating easily. Leaves alternate, oblong or less 

frequently lanceolate, coriaceous, 13.5-35 × 7.5- 

17.5 cm, the apex acute or acuminate, the base 

cordiform; upper surface shiny, with a broad 

midvein; lower surface yellowish green, shiny, 

with the midvein prominent and the secondary 

venation darker; petioles cylindrical, as long as 

the blade, with a winged margin on the lower half. 

Inflorescence axillary, solitary; spathe almost 

cylindrical, 9-13 cm long, erect, greenish, the 

spadix sessile. 


and March and in fruit in June. 


Distribution: In moist forests at lower to 

middle elevations. Also on St. Thomas; Cuba and 

Hispaniola. 



2. Philodendron giganteum Schott, Syn. Aroid. 

89. 1856. 

Fig. 169. A-D 

Terrestrial or epiphytic herb, erect or 

sometimes creeping, attainig 8 m in length. Stems 

cylindrical, 8-10 cm in diameter, producing 

watery latex when wounded. Cataphylls up to 60 

cm long, decomposing into persistent fibers. 

Leaves 25-60 × 17-50 cm, lanceolate or triangularlanceolate, 

coriaceous, the apex obtuse to acute, 

the base cordiform, the lobes not overlapping; 

lower surface slightly pale; petioles erect, almost 

cylindrical, attaining up to 1 m in length. 

Inflorescence axillary, solitary; peduncles robust, 

6-9 cm long; spathe 14-21 cm long, convolute, 

constricted in the central part, greenish outside, 

with the base pink-burgundy, white on the inner 

surface; spadix sessile, robust, as long as the 

spathe, the staminal portion whitish and the 

pistillate portion greenish yellow. Berries yellow 

or orange. 


to April. 


Distribution: Occasional in moist forests at 

the base of mogotes in the northern limestone 

zone. Also on St. John, St. Thomas, and Tortola; 

Hispaniola, the Lesser Antilles, Trinidad, and 

Venezuela. 


3. Philodendron hederaceum (Jacq.) Schott, 

Wiener Zeitschr. Kunst 1829: 780. 1829. 

Fig. 168. C-D 

BASIONYM: Arum hederaceum Jacq. 

SYNONYMS: Philodendron scandens Koch & Sello 

Philodendron micans Klotzsch ex Koch 

Philodendron oxycardium sensu Britton non Schott 

Philodendron isertianum Schott 

Paisaje 


means of aerial roots and attains 10 m in length. 

Stems cylindrical and smooth, producing watery 

and caustic latex when wounded; juvenile stems 

flexuous, with short internodes (ca. 10 cm); adult


412 



stems up to 2 cm in diameter, the internodes 

elongate and with nodes that present an annular 

scar; the lateral branches pendulous. Cataphylls 

up to 12 cm long, caducous. Leaves broadly ovate, 

coriaceous, 14-30 × 10-20 cm, the apex acuminate 

to cuspidate, the base cordiform, the margins 

entire or slightly undulate, pale; upper surface 

dull, sometimes glaucous, with the midvein 

prominent and broadened; lower surface yellowish 

green, dull, with the midvein and some of the 

lateral veins prominent; petioles arcuate or 

ascendant, 10-15 cm long, almost cylindrical. 

Inflorescence axillary, solitary, ascendant; 

peduncles robust, 5-9 cm long; spathe persistent, 

ca. 15 cm long, thick, convolute, almost 

cylindrical, yellowish outside and reddish inside 

when mature; spadix almost sessile, cylindrical, 

robust, whitish, as long as the spathe. 

Phenology: Collected in flower from May to 

August. 


Distribution: Abundant in moist forests at 

lower to middle elevations. Also on St. John, St. 

Thomas, and Tortola. Of widespread distribution 

in the Neotropics. 



4. Philodendron lingulatum (L.) C. Koch, Index 

Sem. Hort. Berol. App. 4[13]. 1853. 

Fig. 170. A 

BASIONYM: Arum lingulatum L. 

Bejuco de calabaza, Calabazón cimarrón 


means of aerial roots and attains 15 m or more in 

length. Stems flexible, cylindrical, dark green, 

sulcate along one side, 0.5-2.5 cm in diameter, 

producing scarce watery latex when wounded; 

epidermis exfoliating easily. Leaves simple, 

elliptical or ovate, subcoriaceous, 13-42 × 7-28 

cm, the apex acute or abruptly acuminate, the base 

cordiform, subcordiform, cuneate, or truncate; 

upper surface semi-shiny, with a broad midvein; 

lower surface semi-shiny, with the midvein 

prominent and the secondary venation lighter; 

petiole winged (except for the distal portion), 12- 

48 cm long, shorter or longer than the blade. 

Inflorescence axillary, solitary or sometimes 

paired; spathe cylindric-urceolate at the base, 10- 

23.5 × 2.9-5.3 cm, erect, light green or whitish 

on the inner surface, dark green outside; spadix 

stipitate, 8.7-21.2 cm long, the fertile staminate 

portion 0.9-1.7 in length, white to cream-colored, 

the sterile portion 0.6-1.3 cm long, the pistillate 

portion 2.1-4.7 cm long. Fruit and seeds unknown. 

Phenology: Collected in flower from May to 

July. 


Distribution: In wet and rain forests, in the 

Cordillera Central, the Sierra de Luquillo, and 

the zone of mogotes. Also on Tortola; Hispaniola 

and the Lesser Antilles. 

Public Forests: El Yunque, Guilarte, and Rio 

Abajo. 

5. Philodendron ornatum Schott, Oesterr. Bot. 

Wochenbl. 3: 378. 1853. 

Fig. 170. B-C 

Slightly woody vine, glabrous, that climbs by 


Stems cylindrical and smooth, with watery and 

caustic sap; stems 2-2.5 cm in diameter, the 

internodes elongate, the nodes with an annular 

scar. Cataphylls ca. 24 cm long, persistent. Leaves 

cordiform, chartaceous, glabrous, 32-60 × 24-46 

cm, the apex acuminate, the base deeply cordate, 

the margins undulate, slightly revolute; upper 

surface dull, dark green, the midvein and the 

secondary veins sunken; lower surface pale, with 

the midvein and the lateral veins prominent and 

reddish; petioles ascendant, 14-42 cm long, 

flattened along the upper surface, pulvinate in the 

apical portion, the pulvinule with small tubercles 

or protuberances. Inflorescence axillary, in groups 

of three, ascendant; peduncles robust, 11-13 cm 

long; spathe persistent, 16-18 cm long, thick, 

convolute, almost infundibuliform in outline 

(slightly constricted below the middle), aristate 

at the apex, whitish with a pink tinge outside, 

and cardinal red inside when mature; spadix 

cylindric-ellipsoid, robust, cream-colored, almost 

as long as the spathe. 

Phenology: Collected in flower in August. 

Status: Apparently introduced, naturalized in 

the area of Guajataca. 

Distribution: Rare, at the base of mogotes with 

abundant organic soil. Native to South America, 

Venezuela, Trinidad, Tobago, Surinam, and 

Brazil. 

Public Forest: Guajataca.

414 



5. SYNGONIUM 

Epiphytic or hemiepiphytic herbs, usually climbing by means of aerial roots. Stems producing 

abundant milky latex. Leaves simple or divided into 3-11 leaflets; petioles forming a leaf sheath at the 

base. Inflorescences axillary, solitary or in groups of up to 11; peduncles of the inflorescence erect, 

pendulous when fruiting; spathe fleshy, convolute, notably narrow in the middle, the tube ellipsoid, the 

blade whitish to greenish, opening when mature; spadix shorter than the spathe, erect, with the pistillate 

flowers in a basal position. Flowers unisexual, the perianth absent; stamens 3-4, united into a synandrium; 

ovary with (1)2(3) locules, which contain 1 or less frequently 2 ovules; stigma discoid or bilabiate. 

Fruit a berry, containing a single seed, united to form an ovoid syncarp. A neotropical genus of 33 

species, the majority climbing. 

1. Syngonium podophyllum Schott, Bot. Zeitung 

(Berlin) 9: 85. 1851. 

Fig. 171. A-C 

Vine attainig 10 m in length, climbing by 

means of adventitious roots that it produces in 

the area of the nodes. Stems cylindrical, glaucous, 

1-2 cm in diameter, producing milky latex when 

wounded. Juvenile plants with hastate leaves; 

adult plants with dimorphic leaves, the basal 

leaves hastate, the distal leaves digitate, with 3- 

11 leaflets, coriaceous, united or free at the base, 

the basal leaflets smaller and auriculate at the 

base, the middle leaflets 16-38 × 6-17 cm, obovate, 

elliptical, or lanceolate, with the apex acuminate; 

petioles 15-60 cm long, almost cylindrical. 

Inflorescences in groups of 4-11, ascendant; 

peduncles 8-9 cm long, slender; spathe ca. 10 cm 

long, convolute at the base to form a tube, the 

limb cream-colored on the inner surface, green 

outside, concave, ephemeral; spadix whitish, 

sessile, cylindrical, with a constriction between 

the area of pistillate flowers and the staminate 

flowers. Syncarp ovoid, red, reddish orange, or 

yellow, 3-5.5 cm long. 

Phenology: Collected in flower in February, 

July and August. 


Distribution: Species escaped from cultivation 

and naturalized, of rapid proliferation, found 

throughout Puerto Rico in moist disturbed areas. 

Also on St. John and St. Thomas. Native to 

Mexico, but widely cultivated throughout the 

Antilles and tropical continental America. 



References: Acevedo-Rodríguez, P. & D.H. Nicolson (in prep.) Araceae. In: Acevedo-Rodríguez, P. 

& M.T. Strong (eds.) Monocots of Puerto Rico and the Virgin Islands. Contrib. U.S. National Herbarium. 

Arias Granda, I. 1998. El género Philodendron Schott (Araceae) en Cuba. Feddes Repert. 109: 33-39. 

Croat, T. B. 1997. A revision of Philodendron subgenus Philodendron (Araceae) from Mexico and 

Central America. Ann. Missouri Bot. Gard. 84: 311-704. Madison, M. 1977. A revision of Monstera 

(Araceae). Contrib. Gray Herbarium 207: 3-100. Nicolson, D. H. 1987. Araceae. In: Dassanayake, M. 

D. and F. R. Fosberg (eds.) Flora of Ceylon. Vol. VI. Amerind Publishing Company, New Delhi.

416 



2. Family ASPARAGACEAE 

1. PROTASPARAGUS 

Shrubs or subshrubs, erect or scandent and twining, with tuberous roots. True leaves absent or 

reduced to small scales. The photosynthetic function is carried out by modified branchlets (phylloclades), 

needle-like or leaf-like, these whorled or in fascicles. Flowers bisexual, minute, in racemes or scattered 

among the phylloclades; perianth with 6 free segments; stamens 6; ovary superior, 3-locular, with 4-10 

ovules per locule, the stigmas three. Fruit a berry. Seeds globose, smooth, black. A genus of about 100 

species, distributed in southern and northern Africa and in Asia.

418 


1. Protasparagus setaceus (Kunth) Oberm., S. 

Afr. J. Bot. 2: 244. 1983. 

Fig. 172. A-E 

BASIONYM: Asparagopsis setacea Kunth 

SYNONYMS: Asparagus setaceus (Kunth) Jessop 

Asparagus plumosus Baker 

Semiwoody vine, twining, attainig 2-3 m in 

length. Stems cylindrical, green, spiny at the base. 

Phylloclades in dense whorls, needle-like, 3.5-6.5 

mm long. Flowers solitary, pendulous; pedicels 

green, 2-2.4 mm long, articulated near the base; 

perianth pale green, the segments oblong, obtuse, 

ca. 2.5 mm long. Berries globose, ca. 5 mm in 

diameter, olive-green or black. 


February. 


Distribution: Ornamental plant, very common 

in our gardens. Also on St. Thomas. Native to 

South Africa, but widely cultivated throughout the 

tropics. 

References: Obermeyer, A. A. 1983. Protasparagus. nom. nov., new combinations. S. Afri. J. Bot. 

2: 243-244. Obermeyer, A. A. 1985. The genus Protasparagus in South Africa. Bothalia 15: 548-549. 

3. Family CYPERACEAE 

1. SCLERIA 

Herbs, erect or infrequently ascendant. Rhizomes absent or less frequently elongate, short, or 

tuberous. Culms triangular. Leaves well developed along the culm (stem), the basal ones not developed; 

leaf blades linear, flat or somewhat involute, herbaceous, usually with the margins and the principal 

vein scabrous; leaf sheath trigonous, closed. Inflorescence a panicle or spike, terminal or terminal and 

axillary; involucral bracts similar to the leaves or absent; spikelets sessile or pedicellate, the staminate 

ones lanceolate or oblong, with numerous flowers, the scales spirally arranged; pistillate spikelets 

ovate or elliptical, the scales distichous; flowers unisexual; stamens 1-3; gynoecium with 3 stigmas, 

ovary usually with a hypogynium at the base. Fruit a subglobose achene, white or variegated with 

purple. A genus of tropical and subtropical zones, of about 200-225 species. 

Key to the species of Scleria 

1a. Contraligule entire; hypogynium conspicuously trilobate.………………………………...…..2 

1b. Contraligule scarious at the apex; hypogynium obscurely trilobate, discoid ………3. S. secans 

2a. Panicles not congested; margin of the leaves with thick retroflexed hairs (0.2-0.5 mm long), very 

cutting; scales of the spikelets dark brown or purple-brown; achenes ovoid-globose or globose, 2.6- 

3 mm long ...........………………………………………………………….1. S. canescens 

2b. Panicles congested; margin of the leaves with minute retroflexed hairs (almost not cutting); scales 

of the spikelets greenish brown; achenes globose, 2.3-2.6 mm long. ……...2. S. scindens 

1. Scleria canescens Böck., Cyp. Nov. 1: 37. 

1888. 

Fig. 173. A-B 

Cortadera, Cortadora 

Perennial herb, robust, climbing, forming 

large impenetrable colonies, and attaining 3.5 m 

in length. Culms ascendant, acutely triquetrous, 

3-10 mm wide, retrorsely and antrorsely scabrous 

on the corners, strigose on the distal portion. 

Leaves numerous, alternate, sparse, broadly linear,


25-60 × 0.4-1.0 cm, pleated or almost flat, with 

three main parallel veins, subrigid, retrorsely 

scabrous on the veins and the margins, the apex 

acuminate or obtuse; leaf sheath as long as the 

internode, scabrous on the angles. Panicles 

solitary, distal or axillary, pyramidal, the distal 

ones 6-14 cm long, with a short leafy bract at the 

base; axes minutely pubescent or glabrous; 

bracteoles filiform. Staminate and pistillate 

spikelets 4-6 mm long; sterile scales 4-5, ovatecircular, 

brown or reddish brown, finely ciliate 

on the margins, the apex apiculate. Achenes 

ovoid-globose or ovoid, 2.6-3.3 mm long, white, 

shiny or with a purple tinge; hypogynium 

trilobate, 1-1.8 mm long, whitish, the lobes 

semicircular, with the margins reflexed and 

appressed to the achene. 



Status: Apparently endemic to Puerto Rico. 

Locally very common. 

Distribution: In moist or rain forests of the 

cordilleras. Reported for Cuba, but no specimens 

seen or found. 



2. Scleria scindens Nees ex Kunth, Enum. Pl. 2: 

343. 1837. 

Fig. 173. C-E 

Razor grass 

Perennial herb, forming large colonies, erect 

or climbing, attainig 3.5 m in length. Culms 

acutely triquetrous, 3-8 mm wide retrorsely and 

antrorsely scabrid on the corners, strigulose on 

the distal portion. Leaves alternate, linear, 30-90 

× 0.5-1.5 cm, almost flat or pleated, with five 

parallel main veins, subrigid, retrorsely scabrid 

on the veins and the margins; leaf sheath scabrid 

on the angles. Panicles solitary, terminal, 

pyramidal, dense, 3-8 cm wide, sometimes with 

one or two small panicles at the base; bracts 

usually 3 on the terminal panicles. Staminate and 

pistillate spikelets intermingled, of the same size; 

sterile scales 2-3, 3-4 mm long, light brown, ciliate 

on the margins, apiculate at the apex. Achenes 

globose, 2.3-2.6 mm long, apiculate, white, shiny, 

sometimes with a purple tinge; hypogynium 

trilobate, 1-1.7 mm long, the lobes semicircular, 

slightly revolute, divergent (not appressed to the 

achene). 


February to August. 


Distribution: In moist or semi-moist disturbed 

areas in the Virgin Islands (St. John, Tortola, and 

Virgin Gorda). Also in Cuba, Hispaniola, and the 

Lesser Antilles. 

3. Scleria secans (L.) Urb., Symb. Antill. 2(2): 

169. 1900. 

Fig. 173. F-G 

BASIONYM: Schoenus secans L. 

Perennial herb, rhizomatous, climbing, 

attainig 5-10 m in length. Culms branched in the 

distal portion, acutely triquetrous, retrorsely 

spinulose on the corners, distally hirsute. Leaves 

alternate, linear, 10-40 × 0.2-0.7 cm, nearly flat 

or pleated, attenuate at the apex, the upper surface 

hirsute at the base, the lower surface sparsely 

hirsute to glabrous, retrorsely scabrous along the 

midvein and the margins; leaf sheath scaberulous 

on the angles. Inflorescences of few spikelets, 

composed of a terminal panicle and 1-3 axillary 

panicles that are borne from the ultimate leaf; 

terminal panicle 2.5-6 cm long; the lateral ones 

shorter. Staminate spikelets shorter than the 

pistillate ones; sterile scales 4, ovate or ovatelanceolate, 

2.5-6 mm long, light or dark brown, 

reddish or blackish on the margins. Achenes ovoid 

or oblong-ovoid, rounded at the apex, appressedpubescent, 

2.8-4 mm long, white, sometimes with 

greenish spots; hypogynium discoid, obscurely 

trilobate, 0.3-1 mm long, the lobes entire, reflexed. 


October to May. 


Distribution: In thickets, along trails, and at 

the margin of moist forests along the Cordillera 

Central and the Sierra de Luquillo. Also in the 

Antilles, Mexico, Central America, and South 

America. 

Public Forests: Carite and El Yunque. 

Reference: Strong, M. T. and P. Acevedo-Rodríguez. (in prep.) Cyperaceae. In: Acevedo-Rodríguez, 

P. & M.T. Strong (eds.) Monocots of Puerto Rico and the Virgin Islands. Contrib. U.S. National 

Herbarium.

420 



4. Family DIOSCOREACEAE 


1a. Fruit a trivalvate capsule, with three flattened locules .…………..……...….……1. Dioscorea 

1b. Fruit a unilocular samara, flattened, with a distal wing……………..………………2. Rajania 

1. DIOSCOREA 

Dioecious vines, herbaceous or slightly woody. Stems cylindrical, angular or winged, sometimes 

provided with spines. Leaves alternate or opposite, simple or palmately lobed, 3-7(-14)-veined (veins 

arcuate, parallel), with long petioles and axillary bulbils. Flowers unisexual, actinomorphic, produced 

in axillary spikes, racemes, or panicles. Perianth minute; staminate flowers with 6 stamens or sometimes 

with 3 stamens and 3 staminodia; pistillate flowers with 6 staminodia, the ovary inferior. Fruit capsular, 

chartaceous or coriaceous, 3-winged; seeds numerous, flattened and winged. A tropical and subtropical 


Key to the species of Dioscorea 

1a. Stems predominantly quadrangular or 4-winged (cylindrical at the base in D. alata)………….2 

1b. Stems cylindrical or triangular, not winged…………………………………………………….3 

2a. Leaves opposite, entire ...................…….......................................…….............….......1 D. alata 

2b. Leaves alternate, 3-6-lobate ………………………………...…….……………...…7. D. trifida 

3a. Leaves pilose on the lower surface along the venation ………………….....…5. D. pilosiuscula 

3b. Leaves glabrous……….......……………………………………………………...………………….4 

4a. Stems cylindrical………………………………………………………………………………..5 

4b. Stems acutely triangular, spiny ……..……………………………………...….…2. D. altissima 

5a. Stems unarmed; leaves alternate……………………………………………………………..…6 

5b. Stems usually aculeate; leaves usually opposite ……………………….………4. D. cayenensis 

6a. Staminate flowers in lateral cincinni along the axis of the inflorescences; fertile stamens 3; bulbils 

angular, smooth; base of the petiole with the margin slightly winged, this not extended on the stem 

like a pseudostipule ........………………………………………..…6. D. polygonoides 

6b. Staminate flowers solitary along the axis of the racemes; fertile stamens 6; bulbils rounded, verrucose; 

base of the petiole winged, projecting toward the stem on each side like a semicircular pseudostipules 

................................……...……………………………………………....3. D. bulbifera 

1. Dioscorea alata L., Sp. Pl. 1033. 1753. 

Fig. 174. A-E 

SYNONYM: Rajania flexuosa Bello 

Ñame, Ñame blanco, 

Water yam, White yam 

Non-woody vine, twining (toward the right), 

glabrous, attainig 10-15 m in length. Stems 

quadrangular, with 4 longitudinal projections, 

winged, undulate, green or reddish; mature stems 

(at the base) cylindrical and spiny. Leaves mostly 

opposite, sometimes alternate on branches of rapid

422 


growth, coriaceous, broadly ovate, 5-7-veined, 10- 

30 × 5-18 cm, the apex acute or acuminate, 

sometimes reflexed, the base cordiform; upper 

surface dark green, shiny, with the venation 

sunken; lower surface pale green, dull, with 

prominent venation; petioles 4-12 cm long, 4winged, 

forming an auriculate sheath at the base, 

with a pair of pseudostipules that enclose the stem; 

bulbils elongate, pendulous, attaining 15 cm long, 

produced when the leaves begin to wither. 

Inflorescences axillary, unisexual, pendulous; the 

staminate ones paniculate, 5-15 cm long, with 

numerous lateral and flexuous spikes that contain 

numerous male flowers; the pistillate ones 

racemose, with few flowers. Staminate flowers 

with the perianth 1-1.5 mm long; pistillate flowers 

with the perianth 2-2.8 mm long. Capsule 3locular, 

2-3 cm wide, each locule flattened like a 

wing, with two seeds inside. 


November and in fruit in January. 


Distribution: Very common in disturbed areas 

and moist secondary forests, at lower and middle 

elevations. Also on St. John. Native to 

southeastern Asia, extensively cultivated 

throughout the tropics for its edible bulbils. 


Maricao, Río Abajo, Susúa, and Toro Negro. 

2. Dioscorea altissima Lam., Encycl. Méth. Bot. 

3: 233. 1789. 

Fig. 174. F-G 

SYNONYM: Dioscorea chondrocarpa Lam. 

Ñame dunguey, Dunguey 

Slightly woody vine, twining, glabrous, 

attainig 10 m or more in length. Stems smooth, 

angular or triangular, attaining up to 1.5 cm in 

diameter, with recurved spines, ca. 1 cm long. 

Leaves alternate or less frequently subopposite, 

ovate or almost orbicular, 7-9-veined, 8-15 × 7- 

16 cm, the apex short-acuminate to caudate, the 

base cordiform, the margins slightly undulate; 

upper surface slightly shiny, with the venation flat 

or slightly prominent; lower surface dull, with 

translucent lines and dots, the primary venation 

prominent; petioles slender, as long as the blade, 

sometimes minutely aculeate, with a swelling at 

both extremities, forming a pair of spiny 

pseudostipules at the base. Inflorescences in 

simple spikes, the pistillate ones longer than the 

staminate ones. Perianth of oblong segments. 

Capsules up to 3 cm long and 1.5 cm wide. 

Phenology: Not observed. 

Status: Apparently native, uncommon. 

Distribution: Occasional in moist forests at 

lower to middle elevations. Also in Hispaniola, 

the Lesser Antilles, and Brazil. 

Public Forests: Carite and El Yunque. 

3. Dioscorea bulbifera L., Sp. Pl. 1033. 1753. 

Fig. 175. A-E 

Gunda 

Non-woody vine, twining, attainig 8 m or 

more in length. Stems glabrous, more or less 

cylindrical. Leaves alternate, ovate, 9-11-veined, 

membranaceous, 9-12.5(17) × 6.5-11(15.5) cm, 

the apex acuminate or caudate, the base cordiform; 

upper surface shiny, the venation slightly 

prominent; lower surface dull, the venation 

prominent; petiole 12-15 cm long, the base 

winged, enclosing the stem on each side like a 

semicircular pseudostipule; bulbils rounded, 5-6 

cm wide, verrucose. Inflorescences axillary, 

simple, fasciculate; the staminate ones usually in 

pairs, 9-15 cm long, with the flowers grouped in 

lateral stipitate cymes; the pistillate ones ca. 12 

cm long, with the flowers solitary. Flowers minute, 

sessile, white to pink; staminate flowers with the 

perianth ca. 1.2 mm long; fertile stamens 6, adnate 

on the corolla; pistillate flowers with the perianth 

ca. 1.4 mm long and the hypanthium ca. 2 mm 

long. Capsules 3-winged, ca. 2.5 cm long. 


August to October. 

Status: Exotic, locally common. 

Distribution: In moist secondary forests, at 

middle elevations, in the zone of mogotes and in 

the Sierra de Luquillo. Native to the tropics of 

the Old World. Introduced in the tropics and 

subtropics of the New World. 


4. Dioscorea cayenensis Lam., Encycl. Méth. 

Bot. 3: 233. 1789. 

Fig. 175. F-K 

Ñame de Guinea


424 


Slightly woody vine, twining (toward the 

right), glabrous, attainig 10 m in length. Stems 

flexible, cylindrical, usually with recurved spines. 

Leaves opposite, simple, more or less ascendant, 

coriaceous, ovate, 7-veined, 6-14 × 5-10 cm, the 

apex abruptly acuminate, the base cordiform to 

almost truncate, the margins entire, revolute; 

upper surface shiny, with the venation slightly 

sunken; lower surface dull, with translucent lines, 

the primary venation prominent; petioles 4.5-12 

cm long, pulvinate at both ends. Inflorescences 

axillary, fragrant, unisexual, in pendulous spikes, 

the staminate ones fasciculate, the pistillate ones 

solitary. Staminate flowers with the perianth 2- 

2.5 mm long, the stamens 6, all fertile; pistillate 

flowers with the perianth 1-1.5 mm long. Capsule 

elliptic-oblong in outline, 1.7 cm long. Seeds with 

a basal wing. 

Phenology: Collected in flower in July and in 

fruit in December. 



Distribution: In areas of disturbed or 

secondary vegetation. Although this species was 

described based on material from French Guiana, 

it is native to western Africa, where it has been 

cultivated for centuries (Burkill, 1985). Today it 

is cultivated throughout the tropics, where it has 

become naturalized. 

Public Forests: Río Abajo. 

5. Dioscorea pilosiuscula Bertero ex Spreng., 

Syst. Veg. ed. 16, 2: 152. 1825. 

Fig. 175. L-P 

Ñame de gunda, Gunda, Dunguey, Air yam, 

Wild yam 


length. Stems slender, strong, almost cylindrical 

or slightly angular, glabrous or puberulent. Leaves 

alternate, 5-7-veined, oblong-ovate or ovate, 5.5- 

12 × 2.5-6.5 cm, chartaceous, the apex longacuminate, 

the base cordiform, the margins 

undulate; upper surface slightly shiny, glabrous, 

the venation sunken; lower surface pale green, 

dull, with prominent venation, pilose; petioles 

slender, glabrous or puberulent, 1.5-4 cm long, 

sulcate and with both extremities swollen, 

provided with a pair of small ovoid tubercles in 

the axil. Inflorescences axillary, pendulous. 

Staminate flowers unknown in the populations of 

the Antilles; pistillate flowers with the perianth 

1.5-2 mm long. Capsule oblong, 1-2.5 cm long. 

Seeds ca. 8 mm long, with a basal wing. 


December to July. 


Distribution: Uncommon in disturbed areas 

of secondary vegetation, in moist zones at lower 

and middle elevations. Also on St. John, St. 




Abajo. 

6. Dioscorea polygonoides Humboldt & 

Bonpland ex Willd., Sp. Pl. 4: 795. 1806. 

Fig. 176. A-I 

Gunda 

Non-woody vine, twining, attainig 8 m or 

more in length. Stems glabrous, more or less 

cylindrical. Leaves alternate, ovate, 7-11-veined, 

chartaceous or membranaceous, (4)8-21 × (2.7)5- 

17 cm, the apex acuminate, the base cordiform; 

upper surface shiny, the venation slightly 

prominent; lower surface dull, the venation 

prominent; petiole 5-8 cm long, the base 

marginate, not extended like a pseudostipule; 

bulbils angular, up to 5 cm wide, smooth. 

Inflorescences axillary, pendulous; the staminate 

ones solitary or in pairs, simple or sometimes 

branched, up to 50 cm long, with the flowers 

grouped in sessile lateral cymes; the pistillate ones 

simple, up to 30 cm long, with the flowers solitary. 

Flowers minute, sessile, white or pale green; 

staminate flowers with the perianth ca. 1 mm 

long; fertile stamens 3; pistillate flowers with the 

perianth ca. 1 mm long and the hypanthium ca. 2 

mm long. Capsules 3-winged, 2.5-3.2 cm long, 

in a position almost perpendicular to the rachis 

of the spike. Seeds 2 per locule, almost circular, 

ca. 5 mm in diameter. 




Distribution: In forests of secondary 

succession, at middle elevations. Also throughout 

the Antilles and tropical America. 

Public Forests: El Yunque, Maricao, and Río 

Abajo.


426 



7. Dioscorea trifida L. f., Suppl. Pl. 427.1781. 

Fig. 176. J-L 

Non-woody vine, twining, attainig several 

meters in length. Stems glabrous or puberulent, 

obtusely quadrangular, with winged projections 

along the borders. Leaves alternate, palmate (5lobate), 

9-11-veined, 15-21 × 15-25 cm, 

membranaceous, the lobes ovate-lanceolate, 

acuminate at the apex, the base cordiform; upper 

surface glabrous, the veins slightly prominent; 

lower surface puberulent along the prominent 

veins; petioles 5-19 cm long, with 4 longitudinal 

wings, swollen at the base. Inflorescences axillary; 

the staminate ones paniculate; the pistillate ones 

in racemes. Staminate flowers in lateral cymes, 

sessile, perianth cream-colored, 1.5-6 mm long, 

the stamens 6; pistillate flowers solitary, 

hypanthium pubescent, the tepals ca. 2.5 mm long. 

2. RAJANIA 

Capsules oblong-elliptical in outline, 2.5-3.4 cm 

long by 1.5 cm wide. 


Status: Exotic, apparently naturalized, 

uncommon. 

Distribution: According to Britton and Wilson 

(1924) this species grows spontaneously in Puerto 

Rico in areas where it was formerly cultivated; I 

know collections only from the Agricultural 

Experiment Station. Also throughout the Antilles 

and South America. 


Dioscorea esculenta (Lam.) Burkill, D. 

floribunda Mart. & Gal., D. friedrichsthalii R. 

Knuth, and D. rotundata Poir. are cultivated in 

Puerto Rico, but no evidence exists that they have 

become naturalized or have escaped from 

cultivation. 

Dioecious vines, herbaceous, twining; bulbils elongate. Stems cylindrical. Leaves alternate, entire, 

petiolate. Inflorescences axillary, of racemes or racemiform thyrses, pendulous or ascendant. Flowers 

6-merous; the staminate ones commonly produced in lateral cymes along a racemiform thyrse; fertile 

stamens 6; the pistillate ones produced in racemes; styles bifid. Fruit an indehiscent samara, with a 

distal wing. An Antillean genus, of about 25 species. 

1. Rajania cordata L., Sp. Pl. 1032. 1753. 

Fig. 177. A-L 

SYNONYM: Rajania sintenisii Uline 

Guáyaro, Bejuco de guaragüao, Ñame 

gulembo, White yam 

Non-woody vine, climbing, twining, glabrous, 

attainig 3-10 m in length. Stems slender, 

cylindrical, strong, glabrous. Leaves alternate, 

glabrous, 4-11.5(20) × 2-6.5(12) cm, 5-9-veined, 

coriaceous or less frequently chartaceous, ovate, 

lanceolate, or less frequently hastate, the apex 

acute or acuminate, the base cordiform to almost 

truncate; upper surface slightly shiny, with the 

venation flat or slightly prominent; lower surface 

with prominent venation; petioles 2-7 cm long, 

cylindrical, pulvinate at the base. Inflorescences 

axillary, racemose, pendulous or ascendant. 

Staminate inflorescences solitary or fasciculate, 

4-25 cm long, the flowers grouped in lateral 

cymes, the cymes stipitate, 1-3 per node; flowers 

with the perianth white or cream-colored, 0.9-1.1 

mm long, the stamens 6. Pistillate inflorescences 

4-30 cm long, with solitary flowers distributed 

along the peduncle; flowers with the perianth 1- 

1.3 mm long, the hypanthium ca. 1.5 mm long. 

Fruits samaroid, (1.1)1.6-3.2 cm long, flattened, 

usually reddish or burgundy, with a wing in a 

lateral distal position. 

Phenology: Flowering from July to March and 

fruiting from September to June. 


Distribution: In moist forests at middle to 


in the northern limestone zone. Also on Vieques, 

St. Croix, and St. Thomas; throughout the 

Antilles. 


Guilarte, Maricao, Río Abajo, Susúa, and Toro 

Negro.

428 



References: Burkill, H.M., 1985. The useful plants of west tropical Africa. Vol. 1. Royal Botanic 

Gardens, Kew. Acevedo-Rodríguez, P. (in prep.) Dioscoreaceae. In: Acevedo-Rodríguez & M.T. Strong 

(eds.) Monocots of Puerto Rico and the Virgin Islands. Contrib. U.S. National Herbarium. 

5. Family ORCHIDACEAE 

1. VANILLA 

Herbaceous vines, climbing by means of aerial roots that are produced in the area of the nodes. 

Stems succulent, thick, glabrous, with abundant watery latex, usually caustic. Leaves alternate, articulate, 

succulent, coriaceous, lacking a leaf sheath. Inflorescences of short, axillary racemes, with numerous 

showy flowers. Flowers bisexual; calyx of free and expanded sepals; corolla of free and keeled petals; 

lip adnate to the semicylindrical and elongate column; ovary pedicellate, articulate; stigma with confluent 

lobes. Fruit elongate, indehiscent or partially dehiscent, with numerous minute seeds. A pantropical 

genus, of about 100 species. 

Key to the species of Vanilla (from Ackerman, 1995) 

1a. Leaves shorter than the internodes, early deciduous or persistent…………………………...…2 

1b. Leaves longer than the internodes, persistent…………………………………………………..5 

2a. Leaves persistent, as long as or longer than half the length of the internode, the blade flat; lip 

green, with cardinal red margins and veins, the margins irregular, the pubescence of yellowish 

hairs …………………………………………………………………………….…...6. V. poitaei 

2b. Leaves early deciduous (although some persistent), shorter than half the length of the internode; lip 

of a different color, the margins more or less entire or trilobat..........…………………3 

3a. Lip mostly red-purple; some leaves persistent, these with the apex curved in the form of a 

hook………………………….………………………………………………..…3. V. dilloniana 

3b. Lip completely white, white with reddish lateral spots, or reddish with a yellowish midrib; all the 

leaves deciduous.………………………………………………………………………..…..4 

4a. Lip trilobate (the lateral lobes as long as the central lobe), red with white margins, the midrib 

yellowish; leaves chartaceous, with the margins not revolute ………………….1. V. barbellata 

4b. Lip more or less simple, completely white or with lateral submarginal and sub-basal spots; leaves 

thick-coriaceous, with the margins revolute …………………………... 2. V. claviculata 

5a. Sepals and petals with the margins undulate and the apex reflexed; lip glabrous, with three keels, 

white with a yellow throat; stems less than 5 mm in diameter; leaves broadly elliptical or ovate, 

fleshy-coriaceous ……………………….……………………………..….4. V. mexicana 

5b. Sepals and petals with the margins and the apex straight; lip with a tuft of barbate hairs, yellowgreen 

or yellow-orange; stems 5-15 mm in diameter; leaves oblong or less frequently elliptical or 

ovate; fleshy-rigid……............………………………………………………………….…6

430 


6a. Floral bracts 12-25 mm long; lip entire ………………………………………….7. V. pompona 

6b. Floral bracts 5-10 mm long; lip trilobate ………..……….…………………...... 5. V. planifolia 

1. Vanilla barbellata Reichenb. f. , Flora 48: 274. 

1865. 

Fig. 178. A-E 

Non-woody vine, glabrous, scarcely branched, 

that climbs by means of aerial roots and attains 

5-7 m in length. Stems cylindrical, 3-9 mm in 

diameter, producing abundant watery and caustic 

latex when wounded; internodes 8-13 cm long; 

nodes swollen, with an annular scar, producing 1 

or 2 grayish roots; the lateral branches pendulous. 

Leaves early deciduous, sessile, linear-lanceolate, 

chartaceous, 4-5 × 0.8 cm, the apex acute, the 

margins entire, not revolute; upper surface dull; 

lower surface dull. Inflorescence a pendulous 

axillary raceme with few or many flowers; bracts 

fleshy, broadly ovate, 4-12 mm long. Sepals green, 

open, free, oblong to oblanceolate, 3-4 × 0.9-1.2 

cm. Petals similar to the sepals but slightly falcate, 

dorsally keeled and somewhat broader (1-1.3 cm); 

lip trilobate, greenish on the outer surface, red on 

the inner surface in the center, but turning white 

toward the margins, the center with a tuft of 

barbate and rigid hairs, the basal portion adnate 

to the column; column straight, almost cylindrical, 

2.3-3.3 cm long. Fruits pendulous, fusiform or 

cylindrical, slightly recurved, changing from 

green to yellow-orange when ripe, 7-9 cm long, 

9-13 mm in diameter. 

Phenology: Flowering from April to June 

(Ackerman, 1995), collected in fruit in January. 


Distribution: Uncommon in the southern and 

southwestern zone, in dry forests and coastal 

thickets at lower to middle elevations. Also on 

St. Thomas, St. John, and Virgin Gorda; Florida, 

Cuba, and the Bahamas. 

Public Forests: Guánica, Maricao, and Susúa. 

2. Vanilla claviculata (Wr.) Sw., Nova Acta 

Regiae Soc. Sci. Upsal. 6: 66. 1799. 

Fig. 178. F-H 

BASIONYM: Epidendrum claviculatum Wr. 


that climbs by means of aerial roots and attains 5 

m or more in length. Stems cylindrical, 3-8 mm 

in diameter, producing abundant watery and 

caustic latex when wounded; internodes 7-13 cm 

long; nodes swollen, with an annular scar, 

producing 1 or 2 grayish roots; the lateral 

branches pendulous. Leaves early deciduous, 

triangular to lanceolate, fleshy, 2.5-3.5 × 0.5-0.8 

cm, the apex reflexed, the margins revolute; upper 

surface dull; lower surface dull. Inflorescence an 

erect axillary raceme with few or many flowers; 

bracts fleshy, broadly ovate, 5-11 mm long. Sepals 

olive-green, expanded, free, 3.5-4.5 × 1-1.3 cm, 

the dorsal sepal concave, the lateral ones similar, 

but falcate. Petals olive-green, elliptic-spatulate, 

keeled, shorter and broader than the sepals; lip 

more or less entire, completely white or with some 

reddish spots at the base, the midvein with 

numerous bifurcate hairs, the basal portion adnate 

to the lower third of the column; column straight, 

almost cylindrical, pubescent on the portion near 

the stigma, 2.5-3 cm long. Fruits black when ripe, 

pendulous, fusiform to cylindrical, slightly 

recurved or straight, 7-11 × 0.9-1.2 cm. 

Phenology: Flowering from April to June and 

producing fruits infrequently (Ackerman, 1995). 


Distribution: In the dry limestone zone and 

on serpentine soils. Also in the rest of the Greater 

Antilles. 

Public Forests: Guajataca, Guánica, and 

Maricao. 

3. Vanilla dilloniana Correll, Amer. Orchid Soc. 

Bull. 15: 331. 1946. 

Fig. 179. A-C 




diameter, producing abundant watery and caustic 

latex when wounded; internodes up to 20 cm long; 

nodes slightly swollen, producing 1 or 2 grayish 

roots; the lateral branches pendulous. Leaves 

persistent, lanceolate-triangular, fleshy, sessile, 3- 

3.5 × 1-1.4 cm, the apex reflexed, the margins 

involute; upper surface dull; lower surface dull.


432 


Inflorescence a pendulous axillary raceme, fewflowered; 

bracts fleshy, broadly ovate, 5-7 mm 

long. Sepals light green, free, expanded, 4.8-5.5 

× 0.8-1.2 cm, the dorsal sepal concave, narrowly 

oblong-elliptical, the lateral ones similar but 

falcate, 4.5-5 × 0.9-13 cm. Petals light green, 

oblong-oblanceolate, keeled, 4.1-5 × 1.1-1.4 cm; 

lip crenulate, obscurely trilobate, purple-brown, 

pale toward the margins and the throat, the basal 

portion adnate to the lower half of the column; 

column arcuate, slender, 3.2 cm long. Fruits 

pendulous, fusiform-clavate, ca. 16 cm long and 

12 mm in diameter, changing from green to 

yellow-orange when ripe, partially dehiscent at 

the apex; seeds minute, numerous, black, in a 

white gelatinous matrix. 

Phenology: Flowering from March to May and 

fruiting from September to January. 


Distribution: In moist forests on serpentine 

soils, in the region of Sabana Grande, San 

Germán, and Yauco. Also in Florida, Cuba, and 

Hispaniola. 

Public Forests: Maricao and Susúa. 

4. Vanilla mexicana Miller, Gard. Dict. ed. 8, 

no. 1. 1768. 

Fig. 179. A-C 

SYNONYMS: Vanilla vanilla (L.) Britton, nom. inadmis. 

Vanilla inodora Schiede 



m or more in length. Stems cylindrical, 2-3(5) 

mm in diameter, producing watery and caustic 

latex when wounded; internodes 7.5-20 cm long; 

nodes swollen, with an annular scar, producing a 

single adventitious root per node; the lateral 

branches pendulous. Leaves persistent, elliptical 

or broadly ovate, fleshy-coriaceous, 13-25 × 6.5- 

12.5 cm, the apex acuminate, the margins entire, 

flat; both surfaces dull, glabrous, with prominent 

venation; petioles thick, 0.5-1 cm long. 

Inflorescence an axillary raceme, few-flowered; 

bracts fleshy, broadly ovate, 2.5 cm long. Sepals 

green, thick, rigid, free, expanded, up to 6.5 × 2 

cm, oblong-lanceolate to lanceolate-elliptical. 

Petals similar, but dorsally keeled and smaller; 

lip trilobate, white with a yellow crest, the basal 

portion adnate along 5 mm of the column; column 

white, slender, 2-2.5 cm long. Fruits pendulous, 

cylindrical, up to 27 cm long. 

Phenology: Flowering from March to May, 

producing fruits in Puerto Rico only by artificial 

pollination. 

Status: Exotic, cultivated and escaped, rare. 

Distribution: Escaped from cultivation by 

vegetative reproduction. In moist forests at middle 

elevations. Also on St. Croix; Mexico, Guatemala, 

Nicaragua, the United States (Florida), the 

Antilles, and northern South America. 


5. Vanilla planifolia Jackson in Andrews, Bot. 

Repos. 8: t. 538. 1808. 

Fig. 179. F-G 




diameter, producing watery and caustic latex when 

wounded; internodes up to 12 cm long; nodes 

slightly swollen, producing a single adventitious 

root per node; the lateral branches pendulous. 

Leaves persistent, oblong, elliptical, or ovate, 

fleshy, rigid, 14-25 × 4.5-8 cm, the apex acute or 

acuminate, the margins entire, slightly revolute; 

upper surface dull; lower surface dull; petioles 1- 

1.5 cm long, thick. Inflorescence a pendulous 

axillary raceme, few-flowered; peduncle flexuous; 

bracts fleshy, broadly ovate, 5-10 mm long. Sepals 

yellowish green, thick, free, expanded, 3.5-5.5 × 

1.3 cm, elliptic-oblanceolate. Petals similar, but 

dorsally keeled and smaller; lip greenish yellow, 

reflexed at the apex, with a retuse apical lobe, the 

basal portion unguiculate, adnate to the column; 

column arcuate, 3-3.5 cm long. Fruits black when 

ripe, pendulous, cylindrical, fragrant, up to 25 cm 

long. 

Phenology: Flowering from February to April, 

but rarely producing fruits, because its natural 

pollinators are not found in Puerto Rico. 

Status: Exotic, naturalized by asexual 

reproduction. 

Distribution: Formerly cultivated in Puerto 

Rico for the commercial production of vanilla. 

Today, some of these plantations persist, with 

some populations naturalized in moist forested 

areas at middle elevations. Species native to


434 


Mexico, but widely cultivated in the tropics. Also 

on St. Croix, St. John, and St. Thomas. 


6. Vanilla poitaei Reichenb. f., Linnaea 41: 66. 

1877. 

Fig. 180. A-F 

SYNONYMS: Vanilla eggersii Rolfe 

Vanilla correllii Sauleda & Adams 





wounded; internodes 8-20 cm long; nodes slightly 

swollen, producing 1-3 adventitious roots per 

node; the lateral branches pendulous. Leaves 

persistent, lanceolate to oblong-lanceolate, fleshy, 

rigid, flat, sessile, (3)7-12 × 1.4-3 cm, the apex 

markedly reflexed like a hook; upper surface dull; 

lower surface dull; petioles 3-4 mm long. 

Inflorescence a pendulous axillary raceme, with 

many flowers; bracts broadly ovate, 4-7 mm long. 

Sepals green, free, more or less expanded, the 

dorsal sepal oblanceolate, 5-5.5 × 1.1-1.3 cm, the 

lateral sepals similar but oblique. Petals similar, 

but dorsally keeled, obliquely oblanceolate, and 

smaller; lip greenish outside, white with cardinal 

red margins, and the throat yellow, the basal 

portion short-unguiculate, adnate along ¾ of the 

column; column arcuate, ca. 3 cm long. Fruits 

pendulous, fusiform-cylindrical, ca. 15 cm long. 

Phenology: Flowering from April to July, but 

rarely producing fruits. 


Distribution: In moist or rain forests, at lower 

to middle elevations. Also on Vieques, St. Thomas, 

the Bahamas, Cuba, and Hispaniola. 

Public Forests: El Yunque, Guajataca, Río 

Abajo, and Maricao. 

7. Vanilla pompona Schiede, Linnaea 4: 573. 

1829. 

Fig. 180. G-H 



m in length. Stems cylindrical, 12-15 mm in 


wounded; internodes up to 20 cm long; nodes 

swollen, with an annular scar, producing 1 

adventitious root per node; the lateral branches 

pendulous. Leaves persistent, oblong-ovate or 

oblong-elliptical, fleshy, rigid, flat, almost sessile, 

20-30 × 6-10 cm, the apex acute or obtuse, the 

base obtuse; both surfaces dull, with the parallel 

venation prominent; petioles ca. 1.5 cm long. 

Inflorescence an axillary raceme, with many 

flowers; bracts fleshy, broadly ovate, 1.2-2.5 cm 

long. Sepals greenish yellow or yellowish creamcolored, 

free, expanded, oblanceolate, obtuse, or 

rounded, ca. 9 × 1-2 cm. Petals similar, but 

dorsally keeled; lip yellow-orange, entire, 

unguiculate, adnate along the column; column 

slender, semicylindrical, ca. 7 cm long, barbate 

in the area near the stigma. Fruits pendulous, 

fusiform, ca. 18 × 2.5 cm. 

Phenology: Flowering from March to May, but 

not producing fruits in Puerto Rico because its 

natural pollinators are not found on the island. 

Status: Exotic, persistent, uncommon. 

Distribution: Formerly cultivated for the 

production of vanilla, and still found on 

abandoned farms. Species native to Mexico and 

Guatemala, but widely cultivated in the 

neotropics. 

Public Forests: Maricao. 

Reference: Ackerman, J. D. 1992. The orchids of Puerto Rico and the Virgin Islands. Editorial de 

la Universidad de Puerto Rico. Ackerman, J. D. 1995. An orchid flora of Puerto Rico and the Virgin 

Islands. Mem. New York Bot. Gard. Vol. 73: 1-203.


436 


6. Family POACEAE 


1a. Leaves of the culm different from the leaves of the branches; branches short, whorled in the area of 

the nodes.........………………………………………………………………….……………..2 

1b. Leaves of the culm similar to those of the branches; apex of the leaf sheath lacking setaceous 

appendages; branches short or elongate, alternate………………………………………………3 

2a. Leaves 7-13 per branch; apex of the leaf sheath with setose appendages ……1. Arthrostylidium 

2b. Leaves 15-40 per branch; apex of the leaf sheath lacking setose appendages ……..2. Chusquea 

3a. Leaf blades 0.2-3.4(4.6) cm wide; glumes not aristate.………………………………3. Lasiacis 

3b. Leaf blades 3-11 cm wide; glumes aristate, the arista 5-22 mm long ……………….…4. Olyra 

1. ARTHROSTYLIDIUM 

Herbs, caespitose, perennial, with pachymorphic and sympodial rhizomes. Culms cylindrical, 

lignified, hollow, elongate, scandent, with one main branch per node, the branches with three to many 

branches that are borne from a mound. Leaves of the culm different from those of the branches; foliaceous 

leaves with internal and external ligules; leaf sheath with a prominent or inconspicuous setaceous 

appendage. Inflorescences of spicate racemes, lacking bracts, the rachis straight or flexuous. Spikelets 

subsessile, with 2 or more flowers, bisexual; fertile flowers 1 or more numerous, deciduous along with 

the rachilla when mature; glumes 1 or 2, as long as half the length of the spikelet. About 30 species, 

distributed from Mexico to Bolivia and in the Antilles. 

Key to the species of Arthrostylidium 

1a. Leaves of the branches 0.7-2 mm wide ……………..………..……....……………1. A. farctum 

1b. Leaves of the branches 3-30 mm wide………………………………………………………….2 

2a. Foliaceous leaves 6-11 × 1-3 cm; culms markedly scabrous; rachis of the inflorescence straight 

……………………………………………………………………………….2. A. multispicatum 

2b. Foliaceous leaves 2.5-6 × 0.3-0.7 cm; culms smooth; rachis of the mature inflorescence flexuous 

………………………………………………………...............…….3. A. sarmentosum 

1. Arthrostylidium farctum (Aubl.) Soderstr. & 

Lourteig, Phytologia 64: 163. 1987. 

Fig. 181. A-C 

BASIONYM: Arundo farcta Aubl. 

SYNONYM: Arthrostylidium capillifolium Griseb. 

Climbing bamboo 

Climbing, small bamboo, attainig 3-5(15) m 

in length. Culms smooth, shiny, 1-5 mm in 

diameter, cylindrical, flexible, with internodes 5- 

20 cm long, the nodes with numerous branchlets, 

7-15 cm long, grouped in dense whorls. Leaves 

of the branches linear, 6-15 × 0.7-2 mm,


chartaceous, with the venation parallel; leaf sheath 

with the margins ciliate; setaceous appendages 

in two series, 1-5 mm long. Inflorescences 2-10 

cm long, spicate, with 3-7 spikelets. Spikelets 8- 

11 mm long, linear, appressed to the rachis. 

Phenology: Rarely seen in flower. The plant 

allegedly dies after flowering. 


Distribution: Uncommon on forested slopes, 

forest margins, and in dry thickets on limestone 

and serpentine. Also on Vieques, St. John, and 

St. Thomas; in the Bahamas, Cuba, and 

Hispaniola. This species was erroneously 

accredited to French Guiana in the original 

description. 

Public Forests: Maricao, Río Abajo, and 

Susúa. 

2. Arthrostylidium multispicatum Pilg., in Urb., 

Symb. Antill. 2: 341. 1901. 

Fig. 181. D-G 

BASIONYM: Arundinaria multispicata (Pilger) Hack. 

Climbing, small bamboo, attainig 5-10 m in 

length. Young culms scabrous, smooth when 

mature, cylindrical, flexible, attaining 3-5 mm in 

diameter, the internodes 2.5-10 cm long, the nodes 

with numerous branchlets, 20-40 cm long, 

grouped in dense whorls. Leaves of the branches 

lanceolate, 6-12 × 1-1.7 cm, chartaceous, with 

the venation parallel; leaf sheath puberulent; 

setaceous appendages 1-2 mm long, deciduous; 

internal and external ligules ca. 2 mm long. 

Inflorescences 4-12 cm long, spicate, with 4-10 

subsessile spikelets. Spikelets 2-3 cm long, linear, 

appressed to the rachis, with 3-5 fertile flowers; 

rachis straight. Fruit unknown. 

2. CHUSQUEA 



Distribution: Uncommon on forested slopes 

along the Cordillera Central. Also in Cuba, 

Hispaniola, and Trinidad. 

Public Forests: Carite, Guilarte, Maricao, and 

Toro Negro. 

3. Arthrostylidium sarmentosum Pilg., in Urb., 

Symb. Antill. 4: 108. 1903. 

Fig. 181. H-J 

Bambú trepador, Bambuilla 

Climbing, small bamboo, with numerous 

pendulous lateral branches, attainig 2-8 m in 

length. Stems glabrous, smooth, shiny, cylindrical, 

0.7-2.5(4) mm in diameter, the internodes 10-18 

cm long, the nodes with 15-30 branches, 10-15 

cm long, grouped in dense whorls. Leaves of the 

branches lanceolate, 1-5 × 0.2-0.5 cm, 

chartaceous, glabrous, with the venation parallel. 

Inflorescences terminal on the short lateral 

branches, racemose, 2-3.5 cm long, flexuous, with 

3-5 spikelets. Spikelets pubescent, 15-18 mm 

long, linear, appressed to the rachis, with 4-5 

flowers. 

Phenology: Rarely seen in flower, collected 

in flower in December. 



upper elevations; along the Cordillera Central and 

the Sierra de Luquillo. Also throughout the 

Antilles and in Venezuela. 


Maricao, and Toro Negro 

Small or medium-sized bamboos. Culms elongate, erect, arcuate or scandent, cylindrical, lignified, 

solid or occasionally hollow, with numerous branches per node. Leaves of the culm different from 

those of the branches; foliaceous leaves with internal and external ligules; leaf sheath lacking setose 

appendages. Inflorescences terminal, of panicles or less frequently racemes. Spikelets lanceolate or 

ovate, short-aristate or not aristate, with numerous flowers; glumes 2, short; 2 basal flowers sterile; 

distal flowers bisexual; palea bicarinate; stamens 3; stigmas 2. About 150 species, distributed from 

Mexico to Chile and Argentina, including the Antilles.

438 



1. Chusquea abietifolia Griseb., Fl. Brit. W. I. 

529. 1864. 

Fig. 182. A-D5 

Small bamboo, creeping or climbing, attainig 

5-10 m in length, forming dense thickets. Culms 

cylindrical, 2.5-3.5 cm in diameter, strong, 

scabrous when young, becoming smooth when 

mature, the internodes 6-14 cm long, the nodes 

with numerous minute whorled branches, 4-10 

cm long, these with 15-40 leaves. Leaves 

alternate, overlapping, narrowly lanceolate, 15- 

30 × 2-3 mm, the apex acute, the margins ciliate, 

scabrous; the outer ligule ca. 0.2 mm long, the 

inner ligule ca. 0.5 mm long; pseudostipules ca. 

3. LASIACIS 

1 mm long. Inflorescences of few-flowered 

panicles, ca. 5 cm long. Spikelets 5.5-7 mm long, 

short-pedicellate; glumes oblong, short-aristate, 

the lower ones 1-1.5 mm long, the upper ones 3- 

5 mm long; sterile flowers oblong-lanceolate, 6- 

7 mm long; fertile flowers lanceolate-ovate, 5.5- 

6.5 mm long, pubescent and ciliate at the apex; 

palea ca. 5 mm long. 

Phenology: Not observed. 


Distribution: Uncommon in moist forests at 

600-1205 m in elevation; along the Cordillera 

Central. Also in the rest of the Greater Antilles. 

Public Forests: Maricao and Toro Negro. 

Perennial herbs. Culms cylindrical, lignified, elongate, scandent, decumbent, or erect, branched. 

Leaves monomorphic, those of the culm similar to those of the branches; leaf sheath smooth, lacking 

setaceous appendages; ligules membranaceous; blades linear to ovate. Inflorescences in the form of a 

panicle, the primary branches divaricate, the rachis with a spikelet on the distal portion. Spikelets 

adaxial, cylindrical or globose, covered at the base by a glume, with 2 or more bisexual flowers. A 

neotropical genus of 16 species. 

Key to the species of Lasiacis 

1a. Ligules of the upper leaves 0.2-0.6 mm long …….….……………………….… 1. L. divaricata 

1b. Ligules of the upper leaves > 1.6 mm long……………………………………….…………….2 

2a. Primary branches reflexed or spreading; leaf sheath glabrous or pubescent …….....… 2. L. ligulata 

2b. Primary branches appressed or spreading; leaf sheath papillose, hispid, or pubescent.................... 

…………………………………………………………………………………...... 3. L. sorghoidea 

1. Lasiacis divaricata (L.) Hitchc., Contr. U.S. 

Natl. Herb. 15: 16. 1910. 

Fig. 182. E-G 

BASIONYM: Panicum divaricatum L. 

SYNONYMS: Panicum bambusoides Desv. ex Ham. 

Lasiacis harrisii Nash 

Perennial herb; culms 0.5-5 m in length, erect 

or scandent; brittle, cylindrical, up to 6 mm in 

diameter. Leaves distichous; leaf sheath glabrous; 

ligule 0.2-0.6 mm long; blades lanceolate to linear, 

flattened, (3-)5-12(-16) cm long by (3-)6-14(20) 

mm wide, glabrous except for some hairs on the 

midvein. Inflorescences ovate; primary axis 2- 

12(20) cm long; secondary axes 2-8(12) cm long. 

Spikelets solitary, obovoid, (3.5)3.7-4.3 × 2.1-2.6 

mm; lower glumes 1.4-2 mm long, 7-11-veined; 

upper glumes 3.5-4.5 mm long, 9-11-veined. 


September to February. 



along roads and in open and disturbed areas. Also 

on Caja de Muerto, Desecheo, Mona, Vieques, St. 


Gorda; throughout the Antilles, the United States 

(Florida), and from Mexico to South America.

440 





2. Lasiacis ligulata Hitchc. & Chase, Contr. U.S. 

Natl. Herb. 18(7): 337. 1917. 

Fig. 182 H-K 

SYNONYM: Panicum divaricatum L. var. puberulum 

Griseb. 

Perennial herb; culms 1-5 m in length, erect 

or scandent. Leaves distichous; leaf sheath 

glabrous or pubescent; ligule (1.6)2-3(3.7) mm 

long; blades broad to narrowly lanceolate, 

flattened, (5-)7-14(-17) cm × (6-)10-22(34) mm, 

scabrous, glabrous or pubescent on the upper 

surface. Inflorescences ovate; primary axis 2- 

17(21) cm long; secondary axes 1-8 cm long, 

reflexed or spreading. Spikelets solitary, obovoid, 

(3-)3.2-3.8 × 1.7-2.4 mm; lower glumes 0.7-2 mm 

long, 7-11-veined; upper glumes 2.7-3.5 mm long, 

9-11-veined. 


November to March and in June. 


Distribution: Uncommon in areas of secondary 

vegetation. Also on St. Croix, St. John, St. 


from Central America to Bolivia. 



4. OLYRA 

3. Lasiacis sorghoidea (Desv.) Hitchc., Contr. 

U.S. Natl. Herb. 18(7): 338. 1917. 

Fig. 183. A-D 

BASIONYM: Panicum sorghoideum Desv. 

SYNONYMS: Panicum divaricatum L. var. lanatum 

Schltdl. & Cham. 

Panicum fuscum Sieber ex Griseb. 

Lasiacis swartziana (Hitchc.) Hitchc. 

Perennial herb; culms 1-10 m in length, erect, 

decumbent, or scandent. Leaves distichous; leaf 

sheath pubescent, papillose or hispid; ligule 

(0.3)0.5-1.5(-2.6) mm long; blades lanceolate, 

ovate or linear, flattened, (6-)9-19(-23) cm × 

(6)12-34(46) mm, smooth, pubescent on the upper 

surface. Inflorescences ovate; primary axis (5-)9- 

25(-35) cm long; secondary axes 4-9 cm long, 

appressed or spreading. Spikelets solitary, obovoid 

or ellipsoid, (3-)3.4-4.1 × 2.5-3.2 mm; lower 

glumes (1.2-)1.5-2.1 mm long, 7-11-veined; upper 

glumes 3-4.3 mm long, 9-13-veined. 




Distribution: Uncommon in areas of secondary 

or disturbed vegetation. Also on Vieques, St. 

Croix, St. John, St. Thomas, and Virgin Gorda; 

throughout the Antilles and from Mexico to 

Argentina. 



Monoecious herbs, perennial, caespitose; culms erect or creeping or climbing. Leaves with well 

developed auricles; ligules membranaceous; blade broad. Inflorescences paniculate, solitary or numerous 

at the upper nodes of the stem. Spikelets with a single flower; pistillate spikelets usually with a claviform 

pedicel, separating as a unit or the flowers separating from the glumes, the glumes acuminate or 

aristate, membranaceous, pluriveined, longer than the flower; flowers lanceolate or ovate, whitish, 

persistent; spikelets staminate, smaller than the pistillate ones, linear or lanceolate, hyaline, deciduous, 

the glumes usually absent; stamens 3. 

1. Olyra latifolia L., Syst. Nat. ed. 10, 1261. 

1759. 

Fig. 183. E-H 

Perennial herb; culms erect or climbing, 1-6 

m in length, 3-12 mm in diameter, brittle, hollow 

inside, with numerous branches from the middle 

and upper nodes; nodes somewhat collapsed, 

pilose or glabrous. Leaf sheath pilose or glabrous; 

auricles up to 5 mm long; ligules 0.7-5 mm long; 

blades lanceolate to ovate, 10-32 × 3-11 cm, the 

base obtuse or subcordate, the apex acuminate. 

Panicles pyramidal, 7-20 × 4-14 cm, on peduncles 

ca. 15 cm long.


442 




October and January. 


7. Family SMILACACEAE 

1. SMILAX 

Distribution: In forests, pastures, and along 

rivers, in moist areas, at middle and lower 

elevations. 

Public Forests: Guajataca, Maricao, Río 

Abajo, and Susúa. 

Herbaceous or slightly woody vines, dioecious, with farinaceous rhizomes. Stems cylindrical, strong 

and flexible, sometimes with spines. Leaves alternate, simple, with the main veins arcuate, parallel; 

petioles with a pair of filamentous tendrils at their union with the leaf sheath. Flowers unisexual, 3merous, 

actinomorphic, produced in axillary umbels. Staminate flowers with 6 stamens, the anthers 

basifixed. Pistillate flowers usually with staminodia; ovary 3-carpellate, with 1 or 2 ovules per carpel. 

Fruit fleshy, with 1-6 seeds. A genus of tropical or temperate vines, of about 350 species. 

Key to the species of Smilax 

1a. Leaves coriaceous to rigid-coriaceous, with the margins usually spiny and the apex obtuse or acute; 

stems spiny ......................………..….........………….........…....................1. S. coriacea 

1b. Leaves chartaceous, with the margins entire and the apex acuminate; stems not spiny 

.........……………………………………………………………………….....2. S. domingensis 

1. Smilax coriacea Spreng., Syst. Veg. 2: 103. 

1825. 

Fig. 184. A-F 

SYNONYMS: Smilax ilicifolia sensu Boldingh non Desv. 

Smilax havanensis var. portoricensis A. DC. 

Smilax guianensis var. subarmata O.E. Schulz 

Smilax coriacea var. ilicifolia O.E. Schulz 

Smilax subarmata O.E. Schulz 

Dunguey, Dunguey blanco, Green briar 


tendrils and attains 5 m or more in length. Stems 

slender, strong, flexible, puberulent or glabrous, 

covered with small recurved spines. Leaves 

coriaceous or rigid-coriaceous, 5.5-12(18) × 2.6- 

9(12.5) cm, ovate, elliptical, oblong, lanceolate, 

or linear, 3-7-veined, the apex obtuse, acute, or 

less frequently rounded or slightly retuse, 

mucronate, the base rounded, obtuse, or 

cordiform, the margins revolute and usually spiny; 

upper surface dark green, shiny, glabrous, with 

the midvein prominent, the margins revolute and 

usually spiny; petioles 0.5-2 cm long, articulated 

at the apex, forming a sheath at the base, from 

which two simple tendrils are borne. 

Inflorescences axillary, umbelliform, these solitary 

or on short flexuous branches, lacking leaves. 

Flowers greenish or yellowish. Staminate flowers 

with pedicels ca. 5 mm long, tepals oblong, 

reflexed, 2-3 mm long. Pistillate flowers with 

pedicels ca. 3 mm long; tepals ovate, erect, 1.5- 

1.8 mm long. Fruits globose or depressed-globose, 

5-7 mm in diameter, black when ripe. 

Phenology: Flowering from January to March 

and from June to August and fruiting from 

September to January. 


Distribution: Of widespread distribution, in 

coastal thickets, dry forests, and moist forests of 

the Cordillera Central. Also on Vieques, Guana 

Is., St. Croix, St. John, St. Thomas, Tortola, Virgin 

Gorda; Hispaniola and the Lesser Antilles.

444 


Public Forests: Carite, El Yunque, Guánica, 

Guajataca, Maricao, Río Abajo, Susúa, Toro 


Commentary: In the first edition of this work, 

Smilax coriacea was considered a synonym of S. 

havanensis Jacq. This error is rectified in this 

edition, since the two species are different and 

can be distinguished by their pattern of the 

venation. The secondary veins on the leaves of S. 

coriacea form an angle with the principal vein 

that varies from 45º to 90º, while in S. havanensis 

the secondary veins form an angle with the 

principal vein that varies from 25º to 35º. Smilax 

havanensis is found in Cuba and in Hispaniola, 

but it is not found in Puerto Rico, nor in the Virgin 

Islands. 

2. Smilax domingensis Willd, Sp. Pl. 4: 783. 

1806. 

Fig. 184. G-K 

Bejuco de membrillo, Raíz de zarzaparilla, 

Zarzaparilla 



slender, strong, glabrous, smooth, scarcely 

branched. Leaves alternate, simple, chartaceous, 

5-12(15) × 2-6.5 cm, ovate, oblong, elliptical, or 

broadly elliptical, usually involute, 5-veined, the 

apex acuminate, the base obtuse, rounded, or less 

frequently subcordate; the margins entire; upper 

surface shiny, glabrous; lower surface dull, 

glabrous, papillose, with prominent venation; 

petioles 1-1.5 cm long, articulated at the apex, 

forming a sheath at the base, from which two 

simple tendrils are borne, 8-12 cm long. 

Inflorescence axillary, umbelliform, solitary. 

Flowers greenish, fragrant. Staminate flowers 

with pedicels 5-9 mm long; tepals elliptical, 3- 

3.5 mm long. Fruit globose, fleshy, ca. 1 cm in 

diameter, dark violet or almost black, with a single 

seed inside. 

Phenology: Flowering from May to August 

and fruiting from December to February. 




in the zone of mogotes. Also in the Greater 

Antilles, Mexico, Guatemala, and Belize. 

Public Forests: Cambalache, El Yunque, 

Guajataca, Maricao, Río Abajo, Susúa, and 

Tortuguero. 

Reference: Acevedo-Rodríguez, P. (in prep.) Smilacaceae. In: Acevedo-Rodríguez, P. & M. T. Strong 

(eds.) Monocots of Puerto Rico and the Virgin Islands. Contrib. U.S. Natl. Herb.


446 



BLECHNACEAE 

Blechnum fragile: Acevedo-Rdgz., P. 3064; 7815; 7828; 

10864; 11446. 

DAVALLIACEAE 

Oleandra articulata: Acevedo-Rdgz., P. 6967; 7739; 9324; 

Britton, N.L. 2060; 2204; 6863; 7570; Cowles, H.T. 406; 

Dale 51; Eggers, B. 1209; Heller, A.A. 1072; 4613; 

Hioram, B. 351; Johnston, J.R. 765; Otero, J. 52; Sargent, 

F.H. 604; Shafer, J.A. 3459; Sintenis, P. 1002; 1780; 

Webster, G.L. 8698. 

DENNSTAEDTIACEAE 

Hypolepis nigrescens: Acevedo-Rdgz., P. 9433; 11457; 

Proctor, G.R. 40131; 40629; Sargent, F.H. 3154. Hypolepis 

repens: Acevedo-Rdgz., P. 7740; Proctor, G.R. 39399; 

39644; 40628; 41285. 

Hypolepis tenerrima: Proctor, G.R. 40130; Sargent, F.H. 605; 

Sintenis, P. 4105; 6454. Hypolepis urbanii: Proctor, G.R. 

40130. 

Odontosoria aculeata: Acevedo-Rdgz., P. 2095; 7146; 9321; 

9434. Britton, N.L. 505; 1074; 1612; 6079; Cowles, H.T. 

255; Dale 5; Fisher, M.J. 71; Goll, G.P. 402; 403; 433; 

Heller, A.A. 126; Hioram, B. 108; Proctor, G.R. 39398; 

Shafer, J.A. 3146; Sintenis, P. 169; 2172; Underwood, L.M. 

25; Vélez, I. 1050. 

Odontosoria scandens: Acevedo-Rdgz., P. 7825; 9431. Britton, 

N.L. 1457; 2454; 2613; Proctor, G.R. 39422; 40812; 

Sargent, F.H. 403; 3139; Schubert 408; Sintenis, P. 4600; 

Webster, G.L. 8767. 

GLEICHENIACEAE 

Dicranopteris flexuosa: Acevedo-Rdgz., P. 9427; 9448; 

Britton, N.L. 8126; Heller, A.A. 4353; Hioram, B. 329; 

Proctor, G.R. 39416; 41104; 41158; Stevenson, J.A. 227; 

3259; Underwood, L.M. 1286. 

Dicranopteris pectinata: Acevedo-Rdgz., P. 7120; 9333; 9372; 

Britton, N.L. 584; Cowgill 630; Cowles, H.T. 398; Goll, 

G.P. 438; 938; Proctor, G.R. 39503; 41133; Sintenis, P. 

1768; Underwood, L.M. 266. 

Gleichenia bifida: Acevedo-Rdgz., P. 9322; 9373; 9447; 

10218; 10524; Britton, N.L. 503; 5389; 6561; 7573; 

Cowles, H.T. 399; Eggers, B. 1202; Gaibee; Goll, G.P. 413; 

435; 929; Hioram, B. 283; Proctor, G.R. 39418; Shafer, 

J.A. 3422; Sintenis, P. 429; 1769; 3988; Stimson 1282; 

Underwood, L.M. 288; 330; Webster, G.L. 8627. 

Gleichenia brevipubis: Proctor, G.R. 39420; 41419. 

Gleichenia rubiginosa: Acevedo-Rdgz., P. 9423; Gleason 33; 

Proctor, G.R. 39419; 40136; 42052. 

LOMARIOPSIDACEAE 

Lomagramma guianensis: Proctor, G.R. 41108; Sintenis, P. 

426; 526. 

Lomariopsis amydrophlebia: Proctor, G.R. 40712; Sintenis, 

P. 1795. 

COLLECTIONS EXAMINED 

Lomariopsis kunzeana: Britton, E.G. 5215; Underwood, L.M. 

& Griggs 843. 

Lomariopsis sorbifolia: Acevedo-Rdgz., P. 10816; Sintenis, 

P. 1762. 

POLYODIACEAE 

Microgramma heterophylla: Acevedo-Rdgz., P. 1907; 2037; 

10891. 

Microgramma lycopodioides: Acevedo-Rdgz., P. 263; 9412; 

11653; Britton, N.L. 4129; 7289; Fosberg, F.R. 51334; 

Goll, G.P. 404; 617; Heller, A.A. 1369; 4476; 6235; Proctor, 

G.R. 39396; 40837; Sargent, F.H. 102; Shafer, J.A. 3152; 

3615; Sintenis, P. 1784; 2834; Stimson 1237; Underwood, 

L.M. 70; 292. 

Microgramma piloselloides: Acevedo-Rdgz., P. 200; 3039; 

9319; 10810. 

Polypodium loriceum: Acevedo-Rdgz., P. 7826; 10523; 

10871. 

SCHIZAEACEAE 

Lygodium japonicum: Acevedo-Rdgz., P. 10849. 

SELAGINELLACEAE 

Selaginella willdenovii: Acevedo-Rdgz., P. 9328. 

ACANTHACEAE 

Asystasia gangetica: Acevedo-Rdgz., P. 3083; 7001; 7082. 

Oplonia spinosa: Acevedo-Rdgz., P. 760; 3975; 4116; 4147; 

5418; 7026; 9302; 10595; 11330; Britton, N.L. 2545; 

Heller, A.A. 4688; Johnston, J.R. 803; Prance, G.T. 29373; 

Sargent, F.H. 399; 693; Sintenis, P. 5202; Stevenson, J.A. 

2405; Wasshausen, D.C. 1303; Webster, G.L. 8974; 

Woodbury, R.O. 56. 

Thunbergia alata: Acevedo-Rdgz., P. 3050; 7002; 7135; 

Boom, B. 9895; Eggers, B. 760; Fisher, M.J. 21; 22; 23; 

Goll, G.P. 71; 152; Heller, A.A. 624; 6312; 6392; Liogier, 

A.H. 30010; Prey, N. 82; Sargent, F.H. 189; Sintenis, P. 

158; Stevenson, J.A. 98; 306; 3328; Underwood, L.M. 757. 

Thunbergia fragrans: Acevedo-Rdgz., P. 171; 529; 1928; 

4665; 5329; 7003; 9353; Boom, B. 9891; 9945; Goll, G.P. 

774; Sintenis, P. 2673; Stevenson, J.A. 2154. 

Thunbergia grandiflora: Acevedo-Rdgz., P. 7049; 9350; 9418. 

AMARANTHACEAE 

Celosia nitida: Acevedo-Rdgz., P. 1807; 4358; 6988; 11080; 

11293; Britton, N.L. 132; 1956; 5033; Heller, A.A. 6424; 

Johnston, J.R. 1246; Sargent, F.H. 3211; Shafer, J.A. 2756; 

Sintenis, P. 615; 3024; 3444; Velez, I. 3086a 

Chamissoa altissima: Acevedo-Rdgz., P. 3738; 7050; 7730; 

7849; 9410; 11872; Axelrod, F. 10982; Heller, A.A. 545; 

4521; 6197; Johnston, J.R. 1130; Sargent, F.H. 82; 461; 

704; Sintenis, P. 264; 2139; 5892; 6357; Stevenson, J.A. 

1130. 

Iresine angustifolia: Acevedo-Rdgz., P. 779; 2469;2568; 2726; 

3122; 3890; 6979; 10658; Raunkiaer, C. s.n. 

Iresine diffusa: Acevedo-Rdgz., P. 118; 7852; 9487; 9494; 

Britton, N.L. 1444; Goll, G.P. 600; 622; Heller, A.A. 349;

448 


Morton, C.V. 5740; Sargent, F.H. 221; Sintenis, P. 303; 

2422; 5844; Stevenson, J.A. 2410; Underwood, L.M. 342; 

457. 

Pfaffia aurata: Acevedo-Rdgz., P. 24; 7744. 

APOCYNACEAE 

Allamanda cathartica: Acevedo-Rdgz., P. 9338; Britton, N.L. 

1555; Fisher, M.J. 4; Goll, G.P. 389; 965; Sargent, F.H. 

748; 3010; Shafer, J.A. 3013; Sintenis, P. 832; 2499; 

Stevenson, J.A. 1271. 

Anechites nerium: Stahl, A. 870. 

Echites agglutinata: Acevedo-Rdgz., P. 902; 2045; 2665; 

3142; 3819; 3974; 4422; 5068; 5249; 7797; 10900; 

10978; 11322; 11403; 11558; 11726; Britton, N.L. 615; 

1296; 2277; Goll, G.P. 806; Proctor, G.R. 43988; Sintenis, 

P. 1943; 2457; Stevenson, J.A. 2944; 3503; Underwood, 

L.M. 666 Webster, G.L. 8601; 8896. 

Forsteronia portoricensis: Acevedo-Rdgz., P. 313; 319; 2957; 

5223; 7131; 10827. 

Pentalinon luteum: Acevedo-Rdgz., P. 368; 685; 686; 1835; 

2833; 4292; 11298; Johnston, J.R. 821; Sargent, F.H. 723; 

Shafer, J.A. 2737; Sintenis, P. 1891; Stevenson, J.A. 6583. 

Rhabdadenia biflora: Eggers, B. 1155; Sargent, F.H. b23; 

Sintenis, P. 935; Stevenson, J.A. 433. 

ARISTOLOCHIACEAE 

Aristolochia elegans: Acevedo-Rdgz., P.4136. 

Aristolochia oblongata subsp. calceiformis: Acevedo-Rdgz., 

P. 157; 2207; 11436; 11642; Britton, N.L. 6781; Johnston, 

J.R. 923; Sintenis, P. 3056. 

Aristolochia odoratissima: Acevedo-Rdgz., P. 4098. 

Aristolochia trilobata: Acevedo-Rdgz., P. 2215; 2217; 2431; 

4214; 5080; 6993; Britton, N.L. 5517; Shafer, J.A. 3808; 

Sintenis, P. 812. 

ASCLEPIADACEAE 

Cryptosgetia madagascariensis: Acevedo-Rdgz., P. Acevedo- 

Rdgz., P. 734; 1992; 4681; 6969; 6987. 6989; 10800; 

10829; 11238; 11463. 

Gonolobus stephanotrichus: Acevedo-Rdgz., P. 7785; 10596; 

10814; 11445; Sintenis, P. 4666. 

Hoya australis: Acevedo-Rdgz., P. 7044. 

Marsdenia elliptica: Acevedo-Rdgz., P. 7658; 10169; 11658; 

11739; Axelrod, F. 9399; Wydler, 308. 

Marsdenia woodburyana: Acevedo-Rdgz., P. 10174. 

Matelea maritima: Acevedo-Rdgz., P. 652; 903; 1034; 2737; 

4076; 4674; Britton, N.L. 1641; 1932; Goll, G.P. 750; 

Proctor, G.R. 46582; Shafer, J.A. 2657; Sintenis, P. 2174; 

3208. 

Matelea sintenisii: Acevedo-Rdgz., P. 7792; 7819; Britton, N.L. 

1076; 2040; 2600; Liogier, A.H. 28598. 

Matelea variifolia: Acevedo-Rdgz., P. 351; 11453; 11644. 

Metastelma anegadense: Acevedo-Rdgz., P. 10964; 11054. 

DArcy, W.G. 4809. 

Metastelma decipiens: Acevedo-Rdgz., P. 602; 900; 1815; 

1836; 1941; 2324; 2620; 3106; 3908; 3961; 4144; 4201; 

4258; 4659; 6973; 7083; 9297; 9486; 10944; 11024; 

11076; 11223; 11296; 11855; Axelrod, F. 10519; Britton, 

N.L. 129; 647; 883; 6817; Heller, A.A. 4603; Liogier, A.H. 

10219; 30185; Ostenfeld, C.H. 23; Sintenis, P. 1692; Rose, 

J.N. 3191. 

Metastelma leptocladon: Acevedo-Rdgz., P. 3754; 7714; 

10216; Axelrod, F. 9956; Sargent, F.H. 622. 

Metastelma lineare: Acevedo-Rdgz., P. 382; 3746; 7204; 

9361; 11414; Axelrod, F. 5386; Britton, E.G. 744; Britton, 

N.L. 2548; 2700; Sargent, F.H. 392; Stevenson, J.A. 2040; 

Underwood, L.M. 50; Webster, G.L. 8854. 

Metastelma monense: Acevedo-Rdgz., P. 4273. 

Metastelma parviflorum: Acevedo-Rdgz., P. 399; 1074; 3074; 

5386; 6174; 7919; 10177; Britton, N.L. 55; 311; 1430; 

1581; 1927; 2701; 5885; Fosberg, F.R. 58870a; 

Ricksecker, A.E. 170; Ricksecker, Mrs. J.J. 146; Sargent, 

F.H. 124; 3189; Sintenis, P. 861; 3813; Stevenson, J.A. 

518. 

ASTERACEAE 

Berylsimpsonia vanillosma: Heller, A.A. 6263; Sargent, F.H. 

375; Sintenis, P. 2920; Acevedo-Rdgz., P. 7068. 

Bidens reptans: Acevedo-Rdgz., P. 160; 3734; 7807; Britton, 

N.L. 1559; 7452; Goll, G.P. 572; Heller, A.A. 874; 6081; 

Sargent, F.H. 481; Sintenis, P. 2919; Stevenson, J.A. 2404. 

Bidens urbanii: Acevedo-Rdgz., P. 3735; 3745; 11730; Britton 

& Coewll, 1559; Kennedy, H. 4787. 

Chromolaena borinquense: Acevedo-Rdgz., P. 206; 7779; 

9367; Britton, N.L. 8486; Proctor, G.R. 43021; 44475; 

Sargent, F.H. 3292. 

Koanophyllon polyodon: Acevedo-Rdgz., P. 3744; 7063; 

10526; 10735. 

Lepidaploa borinquensis: Acevedo-Rdgz., P. 3053; 7065; 

7154; 7924; 9421; 10204; Heller, A.A. 4391. 

Mikania congesta: Acevedo-Rdgz., P. 2256; 7786; 11386; 

Axelrod, F. 3118; 7294; Britton, N.L. 2371; Liogier, A.H. 

10709; 32923; Shafer, J.A. 1972; Sintenis, P. 154; 2093; 

3873; Stevenson, J.A. 1704. 

Mikania cordifolia: Abbott 1532; Acevedo-Rdgz., P. 3739; 

4109; 4215; 7124; 7809; 9343; Boom, B. 7071; Britton, 

N.L. 368; 1669; 2349; 8578; Heller, A.A. 211; 4664; 6334; 

6379; Liogier, A.H. 10646; Shafer, J.A. 2644; Sintenis, P. 

204; 3052; 6194; Stevenson, J.A. 1154. 

Mikania fragilis: Acevedo-Rdgz., P. 3057; 3759; 6965; 7102; 

Boom, B. 6928; 7109; 7941; Sintenis, P. 4127; Wagner 

1749; Woodbury, R.O. s.n. 

Mikania micrantha: Acevedo-Rdgz., P. 13; 7052; 7136; 9406; 

10529; 10626; 10751; 11885; V‚lez, I. 3196. 

Mikania odoratissima: Acevedo-Rdgz., P. 7056; 7201; 7790; 

9370; 9371; 10671; 11450; 11754; Britton, N.L. 3899; 

9786; Proctor, G.R. 44339; Sargent, F.H. 8115. 

Mikania pachyphylla: Acevedo-Rdgz., P. 7111; 9435; Axelrod, 

F. 4288; Breckon, G. 4241; Liogier, A.H. 10814; Proctor, 

G.R. 43542; Sargent, F.H. 318. 

Mikania porosa: King, R.M. 10613; Acevedo-Rdgz., P. 7066; 

7212; 9356; Axelrod, F. 3980; Britton, N.L. 1678; 3892; 

Proctor, G.R. 47575; Stahl, A. 381; Stevenson, J.A. 1656. 

Mikania stevensiana: Acevedo-Rdgz., P. 7059; 7169. 

Piptocarpha tetrantha: Acevedo-Rdgz., P. 2974; 7095; 9449; 

Boom, B. 6899; 7104; 10077; Breckon, G. 4428; Britton, 

N.L. 2030; 5285; Liogier, A.H. 10024; Shafer, J.A. 3437; 

Sintenis, P. 1637; 5339. 

Piptocoma acevedoi: Acevedo-Rdgz., P. 5217; 7148; Cedeño, 

J.A. 196; 379. 

Pseudogynoxys chemopodioides: Liogier, A.H. 36008; 

Acevedo-Rdgz., P. 7238; 9420; 11352. 

Salmea scandens: Acevedo-Rdgz., P. 7798; 10862; 10879; 

Britton, N.L. 321; 1610; 1970; Eggers, B. 1159; Heller, 

A.A. 405; 4690; Liogier, A.H. 10711; Sintenis, P. 2346; 

6206; 6628; Stevenson, J.A. 495; Woodbury, s.n. 

Sphagneticola trilobata: Acevedo-Rdgz., P. 923; 2894.


BASELLACEAE 

Anredera vesicaria: Acevedo-Rdgz., P. 1870; 2512 4228; 

Axelrod, F. 9947. 

BIGNONIACEAE 

Amphilophium paniculatum: Acevedo-Rdgz., P. 10457; 


Arrabidaea chica: Acevedo-Rdgz., P. 2105; Sintenis, P. 1096; 

Stahl, A. 79. 

Cydista aequinictialis: Heller, A.A. 4447; Acevedo-Rdgz., P. 

2810; Axelrod, F. 9761; 9814; 10726; Britton, N.L. 8614; 

Heller, A.A. 1027; Sintenis, P. 945; 5764; Stevenson, J.A. 

1752; Underwood, L.M. 261; Woodbury, R.O. 7696. 

Distictis lactiflora: Acevedo-Rdgz., P. 982; 2203; 4660; 5403; 

7010; 10557; 11822; Britton, N.L. 1268; 1771; 5849; Goll, 

G.P. 500; 694; Heller, A.A. 341; Shafer, J.A. 2789; Sintenis, 

P. 577; 2116; 2173; 3091; 3187; 3304; 6819; Underwood, 

L.M. 459. 

Macfadyena unguis-cati: Acevedo-Rdgz., P. 2110; 2803; 

3958; 9352; 11272; Axelrod, F. 4740; Britton, N.L. 66; 

Heller, A.A. 1233; Howard, R.A. 15754; Sargent, F.H. 74; 

Shafer, J.A. 2001; 2349; Sintenis, P. 550; 862; 1242; 1285; 

1415; 3270; 5000; Underwood, L.M. 371; 782; Stevenson, 

J.A. 1757. 

Mansoa hymenaea: Acevedo-Rdgz., P. 7215; 10180; 10183. 

Phryganocydia corymbosa: Acevedo-Rdgz., P. 11500; 

Fosberg, F.R. 58885. 

Podranea ricasoliana: Acevedo-Rdgz., P. 7920. 

Saritaea magnifica: Acevedo-Rdgz., P. 6992. 

BORAGINACEAE 

Cordia bellonis: Acevedo-Rdgz., P. 7164; 10709. 

Cordia polycephala: Acevedo-Rdgz., P. 96; 822; 2059; 3054; 

3834; 5174; 10465; 10481; 11219; 11477. 

Tournefortia bicolor: Acevedo-Rdgz., P. 4121; Britton, E.G. 

2866; Britton, N.L. 339; 6269; Sargent, F.H. B20; Sintenis, 

P. 1420; 6211. 

Tournefortia hirsutisima: Acevedo-Rdgz., P. 2458; 2865; 

4162; 5076; 5154; 5340; 7240; 10224; 11202; 11205; 

Axelrod, F. 5053; Cowell, J.F. 586; Goll, G.P. 808; Heller, 

A.A. 515; Johnston, J.R. 875; Sargent, F.H. 32; Shafer, J.A. 

2693; Sintenis, P. 667; 1733; 6826; Stevenson, J.A. 1568; 


Tournefortia maculata: Acevedo-Rdgz., P. 218; 5222; 5237; 


4541; 5276; 5590; Eggers, B. 1298; Heller, A.A. 609; 

Liogier, A.H. 33133; Sargent, F.H. 184; Shafer, J.A. 2228; 

Sintenis, P. 1575; Webster, G.L. 8927. 

Tournefortia microphylla: Acevedo-Rdgz., P. 777; 1881; 

2326; 3793; 4404; 5361; 6977; 6981; 7771; 7866; 7893; 

10988; 11222; 11257; 11424; 11487; Axelrod, F. 4706; 

5055; 6328; Britton, N.L. 1578; 1580; 1658; 1803; 4626; 

4796; 5376; Liogier, A.H. 9163; Mill., G. 1668; Sargent, 

F.H. 166; 633; Shafer, J.A. 2695; 2868; Sintenis, P. 3404; 

4832; Stevenson, J.A. 5486; V‚lez, I. 3839. 

Tournefortia scabra: Acevedo-Rdgz., P. 7219; Britton, N.L. 

1285; Heller, A.A. 6274; Liogier, A.H. 31136; Sargent, F.H. 

B71; Shafer, J.A. 1982; Sintenis, P. 3378; Underwood, L.M. 

702; Woodbury, R.O. s.n. 

Tournefortia volubilis: Acevedo-Rdgz., P. 986; 1083; 10901; 

Britton, N.L. 271; Liogier, A.H. 31147; Sintenis, P. 640; 

1316; 4957; 5221. 

BUDDLEJACEAE 

Buddleja madagascariensis: Acevedo-Rdgz., P. 11454. 

CACTACEAE 

Hylocereus trigonus: Acevedo-Rdgz., P. 2874; 5091; 7042; 

11250; Britton, N.L. 1583; 1860; 1931; 2305; Hioram, B. 

12; Shafer, J.A. 2834. 

Hylocereus undatus: Acevedo-Rdgz., P. 11494. 

Pereskia aculeata: Acevedo-Rdgz., P. 4267; Gregory 209; 

Shafer, J.A. 2563. 

Selenicereus grandiflorus: Acevedo-Rdgz., P. 1874; 2835. 

CAPPARIDACEAE 

Capparis flexuosa: Acevedo-Rdgz., P. 1026; 1826; 1947; 

2312; 2351; 2521; 4664; 4687; 4688; 5202; 7079; 

Axelrod, F. 6338; Gregory 188; Heller, A.A. 6163; 

Johnston, J.R. 798; Little, E.L. 21636; Mill. 1667; Sargent, 

F.H. 46; Shafer, J.A. 2658; 2767; Sintenis, P. 869; 3334; 

3379; 3613; 3841; 3954; 4999; 5487; Stevenson, J.A. 

2502; 5490; Underwood, L.M. 586; 592; 645. 

CAPRIFOLIACEAE 

Lonicera japonica: Acevedo-Rdgz., P. 2986; 9439; Sintenis, 

P. 4467; Stevenson, J.A. 5106. 

CELASTRACEAE 

Hippocratea volubilis: Acevedo-Rdgz., P. 248; 569; 2999; 


1599; 1917; 4345; 5812; Eggers, B. 1160; Heller, A.A. 

266; 4424; Liogier, A.H. 10265; 30929; Pfeifer 2934; 

Shafer, J.A. 2876; 2899; 2910; 3247; Sintenis, P. 795; 

1854; 5164; 6392; Stevenson, J.A. 396; 2692; Underwood, 

L.M. 643. 

Pristimera caribaea: Acevedo-Rdgz., P. 7229; 7236; 7722; 

7803; 10617; Sargent, F.H. 16. 

CLUSIACEAE 

Clusia gundlachii: Acevedo-Rdgz., P. 2188; 3007; 7929; 9318; 

10634; 10865; 11443; Britton, N.L. 4214; 6374; Eggers, 

B. 1231; Howard, R.A. 16830; Liogier, A.H. 10011; 

10096; Little, E.L. 16328; Maguire, B. 60022; Sargent, 

F.H. 657; Shafer, J.A. 2259; 3174; 3284; 3550; Sintenis, 

P. 350; 351; 1529; Underwood, L.M. 404. 

COMBRETACEAE 

Combretum grandiflorum: Acevedo-Rdgz., P. 10841. 

Combretum indicum: Acevedo-Rdgz., P. 7140; 11217; 11493; 

Heller, A.A. 974; Sintenis, P. 996; 5134; Stevenson, J.A. 

54. 

CONNARACEAE 

Rourea surinamensis: Acevedo-Rdgz., P. 2135; 9340; 11684; 

Britton, N.L. 1692; 2705; Cede¤o, J.A. 121; Eggers, B. 

1246; Heller, A.A. 956; 997; Liogier, A.H. 9696; Pfeifer 

3047; Sargent, F.H. 229; Shafer, J.A. 3172; 3411; Sintenis, 

P. 668; 1040; 1496; 5728; 5984; Stevenson, J.A. 564; 

1755. 

CONVOLVULACEAE 

Aniseia martinicensis: Axelrod, F. 9936; Liogier, A.H. 31343; 

37389; 37405. 

Argyreia nervosa: Sargent, F.H. 754. 

Convolvulus nodiflorus: Acevedo-Rdgz., P. 2274; 2378; 2761; 

3165; 3960; 4000; 5362; 6188; 7008; 7870; 10171; 

Britton, N.L. 28; 1577; 1802; Goll, G.P. 514; 570; Heller, 

A.A. 877; 6060; Sargent, F.H. 148; 207; Shafer, J.A. 2384; 

2984; Sintenis, P. 2115; 2899; 3098.

450 


Ipomoea alba: Acevedo-Rdgz., P. 7918; 10531; 10791; Cowell, 

J.F. 788; Goll, G.P. 154; Heller, A.A. 375; Sargent, F.H. 


Ipomoea batatas: Goll, G.P. 590; 1062; Prey, N. 42; Stevenson, 

J.A. 2230; Acevedo-Rdgz., P. 3243; Stevenson, J.A. 2468. 

Ipomoea calantha: Sintenis, P. 3128. 

Ipomoea eggersii: Acevedo-Rdgz., P. 762; 1078; 2275; 2711; 

3095; 4043; Britton, N.L. 59; 508; Eggers, B. 31; Fosberg, 

F.R. 60856; Ogdon, N. s.n.; Ricksecker, A.E. 367: Rose, 

J.N. 3192; Smith, A.C. 10589. Ipomoea eriocarpa: 

Stevenson, J.A. 2278 

. Ipomoea hederifolia: Acevedo-Rdgz., P. 2347; 3031; 3082; 

3101; Eggers, B. 172; Fosberg, F.R. 59368; Ricksecker, 

A.E. 144; Stevenson, J.A. 5112; Rose, J.N. 3214. 

Ipomoea horsfalliae: Acevedo-Rdgz., P. 7837; 11451; 11675; 

11752. 

Ipomoea imperati: Acevedo-Rdgz., P. 7039; 9294; Britton, 

N.L. 199; 1544; 1716; Goll, G.P. 959; Heller, A.A. 42; 

Sargent, F.H. 726; Shafer, J.A. 2399; Sintenis, P. 976; 


Ipomoea indica var. acuminata: Acevedo-Rdgz., P. 526; 2264; 

4051; 4127; 4666; 6971; 7217; 7804; 9309; 9354; 10789; 

11455; Sintenis, P.442. 

Ipomoea indica var. indica: Axelrod, F. 3467. 

Ipomoea meyeri: Goll, G.P. 745; Heller, A.A. 6225; Sintenis, 

P. 5533. 

Ipomoea nil: Acevedo-Rdgz., P. 2315; 3100; 3161; 5372; 7243; 

Heller, A.A. 6310; Shafer, J.A.d 1973; Sintenis, P. 2912; 

3216. 

Ipomoea ochracea: Acevedo-Rdgz., P. 2607; 3096; 4099; 

5336; 6231; 7718; 10527. 

Ipomoea pes-caprae: Acevedo-Rdgz., P. 2052; 4380; 11057; 

Fosberg, F.R. 52279; Goll, G.P. 3; 957; Sargent, F.H. 107; 

Shafer, J.A. 2400; Sintenis, P. 86; Stevenson, J.A. 1688; 

Underwood, L.M. 924. 

Ipomoea purpurea: Heller, A.A. 1410. 

Ipomoea quamoclit: Britton, N.L. 7888; Goll, G.P. 41; 


Ipomoea repanda: Acevedo-Rdgz., P. 163; 620; 2400; 2559; 

2563; 2839; 3123; 3128; 7117; Britton, N.L. 1516; 7122; 

Eggers, B. 1321; Hartley 13328; Howard, R.A. 16812; 


Sintenis, P. 289; 5330; Stevenson, J.A. 1156; Vélez, I. 1480. 

Ipomoea reptans: Sargent, F.H. 

Ipomoea rubra: Sintenis, P. 962; Wetmore 163. 

Ipomoea setifera: Acevedo-Rdgz., P. 3094; 3764; 5433; 

Britton, N.L. 1392; 1506; 9428; Fisher, M.J. 7; Goll, G.P. 

251; 843; Heller, A.A. 376; 6364; Sargent, F.H. 108; 

Sintenis, P. 963; Stevenson, J.A. 291; 698. 

Ipomoea steudelii: Acevedo-Rdgz., P. 4761; 7766; 7867; 

10799; Britton, N.L. 30; 1828; 4813; Heller, A.A. 6170; 


Sintenis, P. 3226; 3637; 5540; Underwood, L.M. 544; 

Acevedo-Rdgz., P. 7143. 

Ipomoea tiliacea: Acevedo-Rdgz., P. 524; 2468; 2650; 3763; 

6207; 9306; 9332; 11230; Britton, N.L. 127; 1505; 1608; 

5529; Heller, A.A. 971; 6341; Johnston, J.R. 873; Prey, N. 

91; Shafer, J.A. 2449; Stevenson, J.A. 67; 292; Underwood, 

L.M. 127. 

Ipomoea tricolor: Acevedo-Rdgz., P. 3119. 

Ipomoea triloba: Acevedo-Rdgz., P. 2056; 2314; 3079; 3099; 


4975; 6060; Goll, G.P. 797; 800; Heller, A.A. 494; 6222; 

Prey, N. 51; H. Shafer, J.A. 2851; 3042; Sintenis, P. 827; 

3215; 3454; Stevenson, J.A. 2482. 

Ipomoea violacea: Acevedo-Rdgz., P. 2451; 4007; 4304; 6972; 

9458; Britton, N.L. 5043; Fosberg, F.R. 52281; Liogier, 

A.H. 10799; Sargent, F.H. 3226; Sintenis, P. 1873. 

Ipomoea wrightii: Sintenis, P. 3619. Jacquemontia cayensis: 

Acevedo-Rdgz., P. 7874; 10975; 11560; 11564; Liogier, 

A.H. 10794. 

Jacquemontia cumanensis: Acevedo-Rdgz., P. 3964; 3971; 

3973; 4237; 7882; 11423; Axelrod, F. 9562; Britton, N.L. 

1894; 1934; Liogier, A.H. 10605; 35705; Sargent, F.H. 

141; Shafer, J.A. 2918; Woodbury, R.O. 236. 

Jacquemontia havanensis: Acevedo-Rdgz., P. 2348; 4077; 

4177; 4392; 5294; 7007; 7142; 7220; 7763; 7770; 7896; 

10797; 10965; 11011; 11026; 11077; Axelrod, F. 9448; 

Britton, N.L. 217; 4696; 4988; Heller, A.A. 6425; Johnston, 

J.R. 737; Liogier, A.H. 10638; 1565; Sargent, F.H. 167; 

Shafer, J.A. 2815; Sintenis, P. 3453; Stevenson, J.A. 1244. 

Jacquemontia pentanthos: Acevedo-Rdgz., P. 687; 862; 1037; 

2014; 2306; 4024; 4037; 4100; 4102; 4410; 5365; 6974; 

7053; 10476; 11069; 11211; 11397; 11509. Britton, E.G. 

692; Britton, N.L. 13; 1570; 1965; 2820; 4072; 4823; 

5024; 8073; Goll, G.P. 594; 1018; Heller, A.A. 315; 4456; 

Johnston, J.R. 816; Sargent, F.H. 456; Shafer, J.A. 2612; 

2725; Sintenis, P. 31; 3160; Stevenson, J.A. 2455. 

Jacquemontia solanifolia: Acevedo-Rdgz., P. 2663; 3989; 

5414; 6999; 9303; 9366; 10804; Axelrod, F. 9437; 9456; 

Britton, N.L. 1943; 9524; Shafer, J.A. 2687; Sintenis, P. 

5681. 

Jacquemontia tamnifolia: Acevedo-Rdgz., P. 7024; Axelrod, 

F. 10844; Britton, N.L. 4011; Goll, G.P. 176; Heller, A.A. 

4400; 6437; Howard, R.A. 17306; Sintenis, P. 692; 

Stevenson, J.A. 271; 529. 

Merremia aegyptia: Acevedo-Rdgz., P. 527; 3080; 5360; 

7216; Axelrod, F. 10485. 

Merremia cissoides: Acevedo-Rdgz., P. 9348; Liogier, A.H. 

31319. 

Merremia dissecta: Acevedo-Rdgz., P. 691; 1987; 5413; 7006; 

7012; 11373; 11461; Cowles, H.T. 205; Webster, G.L. 

8984. 

Merremia quinquefolia: Acevedo-Rdgz., P. 690; 2296; 3081; 

3895; 5342; 7019; Garcia, J. Goll, G.P. 520; Liogier, A.H. 

10808. 

Merremia tuberosa: Acevedo-Rdgz., P. 5352; Liogier, A.H. 

31166. 

Merremia umbellata: Acevedo-Rdgz., P. 2467; 2472; 4019; 

4103; 5402; 6991; Britton, N.L. 10050; Goll, G.P. 499; 



Poranopsis paniculata: Acevedo-Rdgz., P. 9293; Goll, G.P. 

525; Liogier, A.H. 30239; Stevenson, J.A. 6924. 

Stictocardia tiliifolia: Acevedo-Rdgz., P. 2898; 3120; 4008; 

Axelrod, F. 10071; Shafer, J.A. 2583; Sintenis, P. 86; 

Stevenson, J.A. 3501; 5886. 

Turbina corymbosa: Acevedo-Rdgz., P. 5239; 7241; 9391; 

10774; Britton, N.L. 6013; Sargent, F.H. 109; Sintenis, P. 

5800. 

Xenostegia tridentata: Acevedo-Rdgz., P. 9478; Britton, N.L. 

1498; 5782; Heller, A.A. 1276; 6440; Sintenis, P. 6738; 

Stevenson, J.A. 1315; 1701; Strong, M.T. 417; Underwood, 

L.M. 982.


CUCURBITACEAE 

Cayaponia americana: Acevedo-Rdgz., P. 702; 1850; 2065; 

2625; 2735; 3856; 7853; 10792; 10860; 10896; 11320; 

Axelrod, F. 5064; Britton, N.L. 128; 923; 1240; 4146; 

7800; Heller, A.A. 4663; Liogier, A.H. 10468; 28920; 


2398; 5015; 5236; Stevenson, J.A. 5209; Wilson, P. 265. 

Cayaponia racemosa: Acevedo-Rdgz., P. 9355; 10805; 11444; 

Axelrod, F. 4441; 6087; Britton, N.L. 1784; Stevenson, 

J.A. 772. 

Citrullus lanatus: Acevedo-Rdgz., P. 9347. 

Coccinia grandiflora: Acevedo-Rdgz., P. 11460. 

Cucumis anguria: Acevedo-Rdgz., P. 4023; 5234; Axelrod, F. 

5939; 9695; Britton, N.L. 1265; Eggers, B. 627; Goll, G.P. 

646; Liogier, A.H. 29440; Shafer, J.A. 2663; Sintenis, P. 

601; Stevenson, J.A. 3258; Underwood, L.M. 575. 

Cucumis dipsaceus: Acevedo-Rdgz., P. 8519; 11073; Axelrod, 

F. 10474. Cucumis melo: Acevedo-Rdgz., P. 10460; 

Axelrod, F. 8452; Stevenson, J.A. 5310; Taylor, C.M. 8560. 

Doyerea emetocathartica: Acevedo-Rdgz., P. 1944; 4044; 

4226; 11276; Axelrod, F. 8750; Britton, N.L. 226. 

Fevillea cordifolia: Acevedo-Rdgz., P. 2218; 4724; 9419; 

Axelrod, F. 4276; Britton, N.L. 329; Sintenis, P. 986; 6604. 

Lagenaria siceraria: Acevedo-Rdgz., P. 11421; Britton, N.L. 

10101; Goll, G.P. 454; 974; Sintenis, P. 1922. 

Luffa aegyptiaca: Acevedo-Rdgz., P. 4066; 7231; Goll, G.P. 

829; Heller, A.A. 370; Sintenis, P. 716; 3174; Stevenson, 

J.A. 342. 

Melothria pendula: Acevedo-Rdgz., P. 2413; 5430; Britton, 

N.L. 1668; 2062; Goll, G.P. 180; 327; 328; 351; 608; 873; 

874; Kuntze, C.E.O. 376; Sintenis, P. 897; Stevenson, J.A. 

1797; Underwood, L.M. 35; 207; 349. 

Momordica charantia: Acevedo-Rdgz., P. 2678; 2734; 4671; 

Britton, N.L. 16; Goll, G.P. 60; 310; Heller, A.A. 153; 4458; 

Sargent, F.H. 123; Shafer, J.A. 2520; Sintenis, P. 726; 919; 


Psiguria pedata: Acevedo-Rdgz., P. 203; 7781; 10593; 10702; 

11659; Goll, G.P. 627; 762; Sintenis, P. 2131; Sintenis, P. 

2371. Sechium edule: Acevedo-Rdgz., P. 10528; Axelrod, 

F. 4738. Sicana odorifera: Acevedo-Rdgz., P. 12137; 


CUSCUTACEAE 

Cuscuta americana: Acevedo-Rdgz., P. 775; 1840; 2453; 

2466; 2569; 2751; 3774; 4048; 4190; 7071; 11045; 

11068. 

DILLENIACEAE 

Pinzona coriacea: Acevedo-Rdgz., P. 10821; 10863; Sintenis, 

P. 1442; 2629; 6173. 

ERICACEAE 

Gonocalyx concolor: Axelrod, F. 6643. 

Gonocalyx portoricensis: Axelrod, F. 6735; Luteyn, J. 5116; 

Sargent, F.H. 283; Shafer, J.A. 3641; Webster, G.L. 8697. 

Vaccinium racemosum: Acevedo-Rdgz., P. 3758; 7096; 7928; 

10875; McKee, H.S. 10640; Sargent, F.H. 533; Shafer, J.A. 

3343; 3649. 

EUPHORBIACEAE 

Dalechampia scandens: Acevedo-Rdgz., P. 600; 660; 1048; 

1882; 2325; Axelrod, F. 6231; Sintenis, P. 626; 3579; 


Tragia volubilis: Acevedo-Rdgz., P. 703; Britton, N.L. 36; 337; 

Goll, G.P. 539; 631; 1024; Heller, A.A. 340; Millspaugh 

696; Sargent, F.H. A75; Sintenis, P. 722; Stevenson, J.A. 

991; 1974; Underwood, L.M. 393; 728. 

FABACEAE 

Abrus praecatorius: Acevedo-Rdgz., P. 10486; 710; 2370; 

Britton, N.L. 150; Goll, G.P. 241; 521; 951; Heller, A.A. 

603; Sargent, F.H. 634; Shafer, J.A. 2340; Sintenis, P. 8; 

Stevenson, J.A. 508; Underwood, L.M. 405; 967; Wetmore 

234. 

Acacia retusa: Acevedo-Rdgz., P. 2624; 3791; 4055; 5050; 

5185; 11216; Britton, N.L. 15; 9776; Heller, A.A. 476; 

6191; Shafer, J.A. 2344; Sintenis, P. 45; 1951; 2994; 5216; 

5622; Stevenson, J.A. 2237; Underwood, L.M. 568; 665. 

Acacia vogeliana: Acevedo-Rdgz., P. 1; 3; 3242; 3794; 5047. 

Barbieria pinnata: Acevedo-Rdgz., P. 7074; Goll, G.P. 937; 

Sintenis, P. 151; 5710; 5942; Stevenson, J.A. 248. 

Caesalpinia bonduc: Acevedo-Rdgz., P. 807; 2570; 3932; 

4021; 11475; Britton, N.L. 209; 287; 1900; 4699; 8326; 

9007;Fosberg, F.R. 52287; Underwood, L.M. 701; Prey, 

N. s.n.; Ricksecker, A.E. 331; Sargent, F.H. 569; Shafer, 

J.A. 1587; 2730; Stevenson, J.A. 355; 2946. 

Caesalpinia ciliata: Acevedo-Rdgz., P. 4134; 4227; 4290; 

11522; Britton, N.L. et al. 116; 1678; Britton, N.L. 5047; 

Fosberg, F.R. 54063; Rose, J.N. et al. 3224; Shafer, J.A. 

2819 

Caesalpinia decapetala: Acevedo-Rdgz., P. 7069; Axelrod, F. 

6012; 6013; Liogier, A.H. 31396; Sargent, F.H. 560; 3284; 


Calopogonium caeruleum: Acevedo-Rdgz., P. 9461; Axelrod, 

F. 4255; Britton, N.L. 2822; Liogier, A.H. 10540; 


Calopogonium mucunoides: Acevedo-Rdgz., P. 7805; Britton, 

N.L. 400; 1007; Heller, A.A. 440; 4575; Stevenson, J.A. 

308; Telford 6. 

Canavalia ensiformis: Tracy 308. 

Canavalia nitida: Britton, N.L. 4151; Prey, N. s.n.; Shafer, J.A. 

2823; Sintenis, P. 452. 

Canavalia rosea: Acevedo-Rdgz., P. 776; 2018; 2911; 4320; 

5301; 6975. 

Centrosema plumieri: Acevedo-Rdgz., P. 10168; 10786. 

Centrosema pubescens: Acevedo-Rdgz., P. 256; 1054; 6985; 

9452; Axelrod, F. 10817; Britton, N.L. 2823; 6735; 7317; 

Heller, A.A. 210; 6084; Sargent, F.H. 122; Sintenis, P. 3080; 

5588; Wetmore 175. 

Centrosema virginianum: Acevedo-Rdgz., P. 715; 823; 1929; 

2318; 3896; 4409; 5426; 6959; 7851; 7881; 9414; 11227; 

11383; Britton, N.L. 7; 262; 8125; Fisher, M.J. 8; Goll, 

G.P. 8; 144; 377; 526; Heller, A.A. 926; 6415; McKee 

10603; Sargent, F.H. 355; Shafer, J.A. 2345; Sintenis, P. 

2306; 3573; Stevenson, J.A. 17; 30; 729; 2198. 

Clitoria falcata: Acevedo-Rdgz., P. 10564; Axelrod, F. 5631; 

Britton, N.L. 1468; 1621; Eggers, B. 1351; Goll, G.P. 299; 

373; Liogier, A.H. 10248; 10689; Sintenis, P. 74; 5737; 


Clitoria ternata: Acevedo-Rdgz., P. 2623; 5236; 5392; 

11489;Axelrod, F. 6378; Britton, N.L. 4875; Goll, G.P. 47; 

502; Heller, A.A. 6224; Hioram, B. s.n.; Sargent, F.H. 272; 

Shafer, J.A. 2358; Sintenis, P. 825; 1639; Stevenson, J.A. 

1835; 2244. 

Dalbergia ecastaphyllum: Acevedo-Rdgz., P. 805; 2039; 2041; 

9310; 6996; Axelrod, F. 5012; 9511; 10902; Britton, N.L. 

1712; 6824; Cowles, H.T. 219; Heller, A.A. 331; 4506;

452 



4862; Stevenson, J.A. 1705; 2132; Underwood, L.M. 657; 

699; 700; Woodbury, R.O. s.n. 

Dalbergia monetaria: Acevedo-Rdgz., P. 4715; 7174; 7700; 

9329; 11384; Britton, N.L. 1528; 2130; 2422; 8094; 

Gregory 175; Heller, A.A. 208; 979; Little, E.L. 16423; 

Sargent, F.H. 635; 667; Shafer, J.A. 3377; 3381; 3468; 

Sintenis, P. 328; 1421; 3898; 4498; 5265; 6293; Stevenson, 

J.A. 1282. 

Desmodium axillare: Acevedo-Rdgz., P. 194; 233; 7737; 7793; 

10586; 11439. 

Desmodium incanum: Acevedo-Rdgz., P. 864; 2291; 2518; 

2785; 4159; 9417; 10492; 10587; 10925. 

Desmodium intortum: Acevedo-Rdgz., P. 3761; 9430; 11447. 

Dioclea reflexa: Axelrod, F. 10825; Britton, N.L. 1677; 7815; 

Sintenis, P. 5040; 5311. 

Entada polyphylla: Sintenis, P. 1240. 

Galactia dubia: Acevedo-Rdgz., P. 808; 824; 1035; 1047; 

2331; 3114; 4429; 5351; 7862; 7864; 10798; 10990; 

11220; Axelrod, F. 6964; Britton, N.L. 146a; 207; Liogier, 

A.H. 10626. 

Galactia eggersii: Acevedo-Rdgz., P. 2483; 4138. 

Galactia longifolia: Axelrod, F. 9753; Liogier, A, 37048. 

Galactia striata: Acevedo-Rdgz., P. 716; 1032; 1851; 3816; 

5188; 7225; 7871; 9415; 10474; 10802; 10894; 11471; 

Britton, N.L. 29; 235; 1794; Goll, G.P. 497; 706; 725; 900; 

953; Heller, A.A. 6114; Liogier, A.H. 29965; McKee 

10622; Shafer, J.A. 2573; 2859; Sintenis, P. 1994; 2050; 

2927; Stevenson, J.A. 2168. 

Lablab purpureus: Acevedo-Rdgz., P. 2123; 2899; 2908; 

3030; 4714; 5350; 7085; 10625; 11258; 11261; 11464; 

Axelrod, F. 9506; 10855; Goll, G.P. 796; 995; Heller, A.A. 

36; Shafer, J.A. 2480; Sintenis, P. 5810; Stevenson, J.A. 

6125. 

Machaerium lunatum: Acevedo-Rdgz., P. 2829; Axelrod, F. 

9509. 

Macroptilium atropurpureum: Acevedo-Rdgz., P. 9312; 

11226; Axelrod, F. 10476. 

Macroptilium lathyroides: Acevedo-Rdgz., P. 367; 860; 1854; 

2117; 2757; 3879; 5187; 5393; 10715; 11362; 11498; 

Britton, N.L. 8786; Goll, G.P. 10; 149; Heller, A.A. 30; 


2672; Sintenis, P. 376; Stevenson, J.A. 59; Underwood, 

L.M. 134; 164. 

Mimosa casta: Acevedo-Rdgz., P. 10; 11; 6203; 10830. 

Mimosa ceratonia: Acevedo-Rdgz., P. 5152; 10512; Britton, 

N.L. 897; 1645; Goll, G.P. 406; Heller, A.A. 220; 6388; 

Shafer, J.A. 2717; Sintenis, P. 5050; Stevenson, J.A. 251; 


Mimosa diplotricha: Acevedo-Rdgz., P. 7045. 

Mimosa quadrivalvis: Sintenis 3162; 3199; Stahl. A., s.n. 

Mucuna pruriens: Acevedo-Rdgz., P. 7123; 10627; 10795; 

Axelrod, F. 11009; Goll, G.P. 698; 719; 721; Heller, A.A. 

4403; Liogier, A.H. 31882; Sargent, F.H. 609; 643; 

Sintenis, P. 148; 3107; Stevenson, J.A. 2115; 2170; 2174. 

Mucuna sloanei: Acevedo-Rdgz., P. 11385; Caminero, G. 306; 

Eggers, B. 1317; García, R. 3248; Prey, N. s.n.; Sintenis, 

P. 5748; 5941. 

Mucuna urens: Acevedo-Rdgz., P. 269; 7799; 9405; 10200; 

Axelrod, F. 9325; Sintenis, P. 4785; 5041. 

Neorudolphia volubilis: Acevedo-Rdgz., P. 2984; Britton, N.L. 

1059; 2010; Eggers, B. 1291; Shafer, J.A. 3227; Sintenis, 

P. 369; 1657; Stevenson, J.A. 240; Underwood, L.M. 91. 

Pachyrhizus erosus: Acevedo-Rdgz., P. 5225; 7228; Goll, G.P. 


Phaseolus lunatus: Acevedo-Rdgz., P. 3741; Britton, N.L. 

2068; Goll, G.P. 587; 673; 705; Liogier, A.H. 10810; 

Sintenis, P. 3110; 5778; Stevenson, J.A. 1979. 

Pueraria phaseoloides: Acevedo-Rdgz., P. 2411; 3740; 5432; 

6966; Axelrod, F. 5819; 9472; Hermann, F.A. s.n.; Liogier, 

A.H. 10539; 10574. 

Rhynchosia minima: Acevedo-Rdgz., P. 1957; 2337; 3921; 

5374; 5398; 7222; 7776; 11274; Britton, N.L. 1857; Goll, 

G.P. 574; 733; Heller, A.A. 32; 4571; 6288; Johnston, J.R. 

1157; Sargent, F.H. 587; Shafer, J.A. 2484; Sintenis, P. 117; 

3140; 3888; 5493; Stevenson, J.A. 1157; 1351; 

Underwood, L.M. 212; 712. 

Rhynchosia phaseoloides: Acevedo-Rdgz., P. 173; 220; 9365; 

10202; Britton, N.L. 2138; Sintenis, P. 2837; 4491; 5858; 

5923; 6623. 

Rhynchosia reticulata: Acevedo-Rdgz., P. 282; 880; 883; 

2118; 2304; 2879; 3227; 3889; 4426; 5199; 7202; 10193; 

10480; 11323; Britton, N.L. 229; 1605; Cowell, J.F. 754; 

Fosberg, F.R. 57539; Goll, G.P. 234; 246; 569; 585; 637; 

691; 722; Heller, A.A. 28; 4418; Johnston, J.R. 1316; 

McKee 10646; Millspaugh 252; Sargent, F.H. 494; Shafer, 

J.A. 2363; Sintenis, P. 375; 3724; Stevenson, J.A. 1316; 

Telford 4; Underwood, L.M. 454; 619. 

Senna bicapsularis: Acevedo-Rdgz., P. 705; 841; 2369; 2591; 

3917; 4032; Axelrod, F. 9507. 

Senna nitida: Acevedo-Rdgz., P. 2224; 7189; 10530; 11207. 

Teramnus labialis: Acevedo-Rdgz., P. 621; 648; 1956; 2367; 

5397; 6986; 11203; Britton, N.L. 1601; 1760; 6504; 

Liogier, A.H. 30403; McKee 10629; Sargent, F.H. 503; 

Shafer, J.A. 2446. 

Teramnus uncinatus: Acevedo-Rdgz., P. 3749; 7055; 7806; 

Cowell, J.F. 5490; Goll, G.P. 636; Heller, A.A. 6255; 

Liogier, A.H. 10421; Sargent, F.H. 3218; Sintenis, P. 110; 

2046; 2961; 5914; Stevenson, J.A. 5041; Telford 5. 

Vigna adenantha: Acevedo-Rdgz., P.9346; 9388; 10167; 


2367; Heller, A.A. 1221; Sintenis, P. 5627; Stevenson, J. 

A. 288. 

Vigna antillana: Acevedo-Rdgz., P. 3134; 3796; 3868. 

Vigna hosei: Acevedo-Rdgz., P. 7932; 9456; 10521; 10525; 

10677. 

Vigna longifolia: Britton, N.L. 6751. 

Vigna luteola: Acevedo-Rdgz., P. 2994; 4005; 5179; 7089; 

7915; 9389; 10773; 11240; Britton, N.L. 208; 1594; 5511; 

Goll, G.P. 16; 954; Heller, A.A. 240; 4682; Shafer, J.A. 



Vigna peduncularis: Acevedo-Rdgz., P. 310; 10567; 10723; 

10778; 11640. Vigna unguiculata: Acevedo-Rdgz., P. 9411; 

10859; Britton, N.L. 1464; Goll, G.P. 1061; Sintenis, P. 

2910; Stevenson, J.A. 2176. 

Vigna vexillata: Acevedo-Rdgz., P. 10166; 10794; 11375; 

Axelrod, F. 10478; Goll, G.P. 338; Sargent, F.H. 563; 


LAURACEAE 

Cassytha filiformis: Acevedo-Rdgz., P. 4372; 5299; 6169; 

9301; Britton, N.L. 1896; Fosberg, F.R. 52288; Heller, A.A. 

330; Little, E.L. 21671; Shafer, J.A. 2892; 2920; Sintenis, 

P. 625; 3325; 5675; Stevenson, J.A. 1723; Underwood, 

L.M. 931.


MALPIGHACEAE 

Heteropterys laurifolia: Acevedo-Rdgz., P. 3072; 5231; 5232; 

7080; 7157; 7230; 10197; 10840; Axelrod, F. 10757; 

10904; Britton, N.L. 426; 1510; Hartley 13368; Heller, 

A.A. 1320; 4406; Proctor, G.R. 44917; Sargent, F.H. 421; 

Sintenis, P. 101; 1559; 1693; 3286; 3809; 5068; Stevenson, 

J.A. 2187; Underwood, L.M. 481; 618. 

Heteropterys purpurea: Acevedo-Rdgz., P. 664; 872; 1033; 

2658; 5391; 7009; 7897; 10194; 11472; Britton, N.L. 54; 

1272; 1479; 1509; Goll, G.P. 542; Heller, A.A. 478; 

Sargent, F.H. 5; Shafer, J.A. 2551; Stevenson, J.A. 988; 


Heteropterys wydleriana: Acevedo-Rdgz., P. 2211; 4723; 

4793; 7193; 7708; 7794; Breckon, G. 4219; Britton, N.L. 

4201; Sintenis, P. 192. 

Stigmaphyllon bannisterioides: Acevedo-Rdgz., P. 9345. 

Stigmaphyllon emarginatum: Acevedo-Rdgz., P. 591; 790; 

992; 2017; 2897; 3965; 4042; 4355; 5206; 5366; 5439; 

7081; 9307; 10536; 11218; 11228; 11430; 11473. 

Stigmaphyllon floribumdum: Acevedo-Rdgz., P. 10796; 308; 

1062; 2854; 2855; 3742; 6994; 7013; 7195; 7239; 7765; 

10196; 10205. 

Stigmaphyllon puberum: Acevedo-Rdgz., P. 7072. 

Tetrapterys inaequalis Acevedo-Rdgz., P. 10182; 11676. 

MARCGRAVIACEAE 

Marcgravia rectiflora: Acevedo-Rdgz., P. 91; 581; 2198; 

3736; 7145; 7151; 9404; 9457; Britton, N.L. 470; 2216; 

Heller, A.A. 359; Sintenis, P. 269; 1592; 4461; 5057; 

Stevenson, J.A. 5249; Underwood, L.M. 57; 823; 835. 

Marcgravia sintenisii: Acevedo-Rdgz., P. 2958; 7114; Britton, 

N.L. 2169; Howard, R.A. 16813; Liogier, A.H. 10028; 

Nieves 1573; Shafer, J.A. 3310; 3645; Sintenis, P. 2222; 

4320; Webster, G.L. 8710. 

MENISPERMACEAE 

Cissampelos pareira: Acevedo-Rdgz., P. 826; 1868; 2219; 

2439; 3126; 5343; 7090; 11312; Axelrod, F. 6003; Britton, 

N.L. 126; 667; Heller, A.A. 332; 1283; Otero, J. 564; 

Sargent, F.H. 223; Shafer, J.A. 2482; 3255; Sintenis, P. 326; 

5186; 5354; Stevenson, J.A. 259; 435; 1971; Underwood, 

L.M. 305; 321; 350; 357; Wilson, P. 282. 

Hyperbaena domingensis: Acevedo-Rdgz., P. 3173; 5084; 

7030; 11698; Liogier, A.H. 10117; 10324; Shafer, J.A. 

3238; Sintenis, P. 4220. 

Hyperbaena laurifolia: Acevedo-Rdgz., P. 6998. 

MORACEAE 

Ficus citrifolia: Acevedo-Rdgz., P. 10635. 

Ficus pumila: Acevedo-Rdgz., P. 9336; 9459. 

NYCTAGINACEAE 

Boerhavia scandens: Acevedo-Rdgz., P. 2520; 3105; 4764; 

7892; 10801; 11291; 11499; 11532. 

Bougainvillea glabra: Acevedo-Rdgz., P. 11237. 

Bougainvillea spectabilis: Sintenis, P. 698; Stevenson, J.A. 322. 

Pisonia aculeata: Acevedo-Rdgz., P. 548; 4209; 4737; 7221; 

10651; 11480; Liogier, A.H. 35979; Shafer, J.A. 2572; 

Sintenis, P. 672; Underwood, L.M. 561. 

OLEACEAE 

Jasminum azoricum: Britton, N.L. 5818. 

Jasminum fluminense: Acevedo-Rdgz., P. 2892; 3839; 6978; 

7084; 7872; 11245; 11479. 

Jasminum grandiflorum: Acevedo-Rdgz., P. 10532; Prey, N. 

47. 

Jasminum multiflorum: Acevedo-Rdgz., P. 1892; 3854; 4672; 

10710; Britton, N.L. 1503; Croat, T.B. 60924; Goll, G.P. 

189; 360; 759; Heller, A.A. 383; Sargent, F.H. 327; Shafer, 

J.A. 3012; Sintenis, P. 171; 5422. 

Jasminum sambac: Stevenson, J.A. 255. 

PASSIFLORACAE 

Passiflora bilobata: Acevedo-Rdgz., P. 7681; 7755; 7762; 

11422. 

Passiflora edulis: Acevedo-Rdgz., P. 834; 5164; Liogier, A.H. 

10258; Stevenson, J.A. 6420. 

Passiflora foetida: Acevedo-Rdgz., P. 772; 1948; 5186; 5333; 

11229; Axelrod, F. 5050; Boom, B. 8008; Britton, N.L. 

2244; Goll, G.P. 393; Heller, A.A. 978; 1029; 6119; 

Johnston, J.R. 225; Liogier, A.H. 9805; Sargent, F.H. 193; 

573; Sintenis, P. 810; Underwood, L.M. 113; 346. 

Passiflora laurifolia: Acevedo-Rdgz., P. 927; 929; 1924; 2856; 

5104; 5169; 5355; 11478; Sintenis, P. 1890. 

Passiflora maliformis: Sintenis, P. 1166; 6560. 

Passiflora multiflora: Acevedo-Rdgz., P. 2484; 3857; 4002; 

11650; 11704; Britton, N.L. 1426; 6787; 7118; Sintenis, 

P. 687; 6654; Stevenson, J.A. 1248. 

Passiflora murucuja: Acevedo-Rdgz., P. 7032; 10817; Stevens, 

F.L. 1818. 

Passiflora quadrangularis: Acevedo-Rdgz., P. 10558; Sargent, 

F.H. 595; Shafer, J.A. 3483; Sintenis, P. 1022; Stevenson, 

J.A. 6715; Underwood, L.M. 778. 

Passiflora rubra: Acevedo-Rdgz., P. 103; 285; 905; 2438; 

4052; 9375; Axelrod, F. 5246; 5812; Britton, N.L. 824; 

Goll, G.P. 250; 303; 374; Heller, A.A. Shafer, J.A. 2987; 

Sintenis, P. 341; 2804; 6017; Stevenson, J.A. 153; 


Passiflora serrato-digitata: Axelrod, F. 5749; Eggers, B. 1372; 


Passiflora sexflora: Acevedo-Rdgz., P. 2679; 3752; 7847; 

9320; 9446; 10761; 10869; Axelrod, F. 5338; 6137; 

Britton, N.L. 450; 1398; 2019; 2607; 2756; Cowles, H.T. 

141; Goll, G.P. 440; Heller, A.A. 4475; Howard, R.A. 

16942; Sargent, F.H. 322; 599; Wetmore 209. 

Passiflora suberosa: Acevedo-Rdgz., P. 614; 630; 639; 898; 

2063; 2330; 2434; 3006; 3133; 4075; 4302; 5196; 6958; 

10498; 11374; 11474; 11510; Britton, N.L. 38; 5030; 

Fosberg, F.R. 57550; Goll, G.P. 185; 311; 331; 409; 634; 

Heller, A.A. 6068; Prey, N. 80; Sargent, F.H. 3208; Shafer, 

J.A. 2506; 2788; Sintenis, P. 644; 811; 1687; 3488; 5114; 

Stevenson, J.A. 1856; Underwood, L.M. 472; 601. 

Passiflora tulae: Acevedo-Rdgz., P. 3751; 10537; 11456; 

Britton, N.L. 937; 2588; 5539; Liogier, A.H. 28444; 

Sintenis, P. 4176; 5278. 

PHYTOLACCACEAE 

.Stegnosperma cubense: Britton, N.L. 9628. 

Trichostigma octandrum: Acevedo-Rdgz., P. 628; 4347; 4731; 

5354; 5447; 11350; Britton, N.L. 113; 1705; 2241; Cowell, 

J.F. 758; Heller, A.A. 1165; Sargent, F.H. 205; Shafer, J.A. 


2500 

PIPERACEAE 

Peperomia rotundifloia: Eggers, B. 1153; Sargent, F.H. 411; 

3294; Sintenis, P. 469; 1669.

454 


PLUMBAGINACEAE 

Plumbago scandens: Acevedo-Rdgz., P. 627; 1893; 2462; 

5341. 

POLYGALACEAE 

Securidaca diversifolia: Ricksecker, A.E. 338. 

Securidaca virgata: Acevedo-Rdgz., P. 3747; 6954; 7127; 

9453; Britton, N.L. 983; 4478; 5809; Cowell, J.F. 582; 

Eggers, B. Goll, G.P. 410; 586; 1055; Heller, A.A. 4385; 

Holdridge, L. 215; Sargent, F.H. 250; Shafer, J.A. 2380; 

Sintenis, P. 46; 1065; 5291; 6658; Stevenson, J.A. 102; 


POLYGONACEAE 

Antigonon leptopus: Acevedo-Rdgz., P. 1811; 5267; 11265; 

11468; Goll, G.P. 75; 648; Sargent, F.H. s.n.; Shafer, J.A. 


RANUNCULACEAE 

Clematis dioica: Sintenis, P. 5843. 

Clematis flammulastrum: Acevedo-Rdgz., P. 7224; 10221; 

Britton, N.L. 1762; Heller, A.A. 6156; Sintenis, P. 43; 

Stevenson, J.A. 1074; 1158. 

Clematis polygama: Acevedo-Rdgz., P. 7234; Axelrod, F. 8473; 

Sintenis, P. 2255; 2949; 6207. 

RHAMNACEAE 

Gouania lupuloides: Acevedo-Rdgz., P. 2295; 3812; 7011; 

7087; Goll, G.P. 230; 386; 582; Heller, A.A. 6104; Liogier, 

A.H. 10346; Shafer, J.A. 2356; 2860; Sintenis, P. 200; 

1261; 1960; 2924; 5508; 5550; Stevenson, J.A. 385; 697. 

Gouania polygama: Acevedo-Rdgz., P. 7139; 7811; Sargent, 

F.H. s.n.; 351; Sintenis, P. 36; 3566; 5577. 

ROSACEAE 

Rubus florulentus: Britton, N.L. 4550; Sargent, F.H. 639; 

3039; Sintenis, P. 4100; 4669. 

RUBIACEAE 

Chiococca alba: Acevedo-Rdgz., P. 162; 708; 1914; 2199; 

2432; 2592; 3129; 3821; 5077; 5173; 7020; 7033; 7075; 

7194; 7209; 9491; 10500; 11338; Axelrod, F. 4754; 

Britton, N.L. 1063; Fosberg, F.R. 44165; 51352; 

Holdridge, L. 232; Liogier, A.H. 9895; Sargent, F.H. 702; 

Shafer, J.A. 2945; 3792; Sintenis, P. 1; 278; 774; 1948; 

2393; 2483; 3421; 6199; 6550; Stevenson, J.A. 269; 1798; 

Underwood, L.M. 157; 907; Webster, G.L. 8869. 

Diodia sarmentosa: Acevedo-Rdgz., P. 7126; 7926; 11393; 

Britton, N.L. 1393; 1517; 2350; Heller, A.A. 272; 4387; 

Howard, R.A. 17023; Liogier, A.H. 29765; Proctor, G.R. 

44551; Sargent, F.H. 260; Sintenis, P. 5052; Stevenson, 


Hillia parasitica: Acevedo-Rdgz., P. 4657; 7112; 7931; Britton, 

N.L. 924; 2020; 2203; 2591; 5525; 6541; Fosberg, F.R. 

51339; Pfeifer 2485; Shafer, J.A. 3443; Sintenis, P. 525; 

Wagner 1752. 

Lasianthus lanceolatus: Acevedo-Rdgz., P. 6964; 7827; 7834. 

Psychotria microdon: Acevedo-Rdgz., P. 586; 2395; 2690; 

2876; 6171; 10472; 10932; Britton, N.L. 1507; 6152; 

Heller, A.A. 6102; Liogier, A.H. 9679; 10218; 10302; 

31514; Little, E.L. 21672; Shafer, J.A. 2335; 2529; 


Sabicea villosa: Sintenis, P. 263; Wilson, P. 232. 

Schradera exotica: Acevedo-Rdgz., P. 223; 9436; Boom, B. 

9794; Britton, N.L. 2276; Liogier, A.H. 10050; Shafer, J.A. 

3242; Sintenis, P. 1238; 1429. 

SAPINDACEAE 

Cardiospermum corindum: Acevedo-Rdgz., P. 1087; 1844; 

2345; 2373; 4154; 4345; 11085; Britton, N.L. 1586; 

Liogier, A.H. 35326; Shafer, J.A. 3017. 

Cardiospermum grandiflorum: Acevedo-Rdgz., P. 10843. 

Cardiospermum halicacabum: Acevedo-Rdgz., P. 523; 961; 

966; 967; 968; 969; 1088; 2429; 2474; 2501; 5235; 9463; 

11490; Britton, N.L. 1882; Goll, G.P. 1004; Liogier, A.H. 

10637; Prey, N. 24; Sargent, F.H. 530; Sintenis, P. 1725; 


Paullinia fuscescens: Acevedo-Rdgz., P. 5356; 10845; 11459. 

Paullinia pinnata: Acevedo-Rdgz., P. 304; 1090; 3073; 6230; 

6957; 6961; 7125; 9451; 11442; Boom, B. 7997; Britton, 

N.L. 1525; Eggers, B. 716; Goll, G.P. 369; Heller, A.A. 

1223; Sargent, F.H. 431; Shafer, J.A. 2905; 3404; Sintenis, 

P. 188; 1532; Underwood, L.M. 354; 469; 993. 

Paullinia plumierii: Shafer 1158. 

Serjania lucida: Acevedo-Rdgz., P. 629; 709; 1828; 2281; 

2313; 3146; 3765; 4658; 5210; 5334; 5358; 7086; 9305; 

11081; 11582; Heller, A.A. 56; 4661. 

Serjania polyphylla: Acevedo-Rdgz., P. 719; 2200; 7015; 

9392; 10195; 11204; 11212; Boom, B. 10022; Britton, 

N.L. 772; 1307; 4311; 4651; 4964; Goll, G.P. 692; Heller, 

A.A. 6063; Sintenis, P. 274; 3406; 5646; 5772. 

SCHLEGELIACEAE 

Schlegelia brachyantha: Acevedo-Rdgz., P. 2964; Britton, N.L. 

5534; 6383; Eggers, B. 1318; Liogier, A.H. 10762; 28533; 



Tecomaria capensis: Acevedo-Rdgz., P. 11353; Morrow, C.F. 

61; Shafer, J.A. 1474. 

SOLANACEAE 

Lycianthes virgata: Acevedo-Rdgz., P. 11437; 11643; Britton, 

N.L. 9440; Sargent, F.H. s.n.; Sintenis, P. 2091; 2620; 4235. 

Solandra grandiflora: Liogier, A.H. 35677; Sintenis, P. 364. 

Solanum lanceifolium: Acevedo-Rdgz., P. 906; 1857; 2417; 

2848; 4112; 5108. 

Solanum seaforthianum: Acevedo-Rdgz., P. 11476; Barker, 

E.E. 7859; Britton, N.L. 9867; Liogier, A.H. 29782; 34137; 

36499; Santiago-Blay, J.A. s.n.; Stevenson, J.A. 1829; 

Woodbury, R.O. s.n. 

TROPAEOLACEAE 

Tropaeolum major: Acevedo-Rdgz., P. 10201. 

ULMACEAE 

Celtis iguanaea: Acevedo-Rdgz., P. 635; 2021; 2361; 2691; 

3237; 3815; 5150; 6980; 7237. 

VALERIANACEAE 

Valeriana scandens: Acevedo-Rdgz., P. 9399; 10203; Britton, 

N.L. 495; 5401; Sintenis, P. 302; 4932. 

VERBENACEAE 

Clerodendrum aculeatum: Acevedo-Rdgz., P. 681; 1934; 

1994; 2818; 3766; 4026; 5140; 5409; 6976; 11059; 

11273; 11524. 

Clerodendrum x speciosum: Acevedo-Rdgz., P. 9413.


Congea tomentosa: Acevedo-Rdgz., P. 7121; 9460. 

Holmskioldia sanguinea: Acevedo-Rdgz., P. 10533; 10842; 

Fisher, M.J. 19. 

Petrea volubilis: Acevedo-Rdgz., P. 4239; 11266. 

VITACEAE 

Cissus erosa: Acevedo-Rdgz., P. 4720; 7171; 9471; 10809; 

11503; Britton, N.L. 4083; 6732; 9909; Sargent, F.H. 612; 

Sintenis, P. 190; 4716; 5346; Stevenson, J.A. 6351; 6723; 

Underwood, L.M. 85; 513. 

Cissus obovata: Acevedo-Rdgz., P. 10861; 11708; Britton, N.L. 

1687; Stevenson, J.A. 2621; Underwood, L.M. 550. 

Cissus rotundifolia: Acevedo-Rdgz., P. 11003; 11495. 

Cissus trifoliata: Acevedo-Rdgz., P. 1017; 1799; 2693; 4375; 

5127; 5363; 5446; 11251; 11497; Axelrod, F. 6346; 

Breckon, G. 4067; Sintenis, P. 4901; Acevedo-Rdgz., P. 

2015. 

Cissus verticillata: Acevedo-Rdgz., P. 169; 819; 833; 2088; 

2420; 2820; 3993; 4683; 5114; 5172; 7088; 10591; 

10929; 11681. Vitis tiliifolia: Acevedo-Rdgz., P. 198; 2128; 

Axelrod, F. 5996. 

ARACEAE 

Anthurium scandens: Acevedo-Rdgz., P. 7113; 7132; 9339; 

9395; 10661; 11880; Britton, N.L. 1031; 4138; Eggers, 

B. 1210; Heller, A.A. 6357; Proctor, G.R. 46929; Shafer, 

J.A. 3562; Sintenis, P. 1505; Stevenson, J.A. 5044; 


Epipremnum pinnatum: Acevedo-Rdgz., P. 7141; 10838; 

11260; Howard, R.A. 16921. 

Epipremnum pinnatum “aureum”: Acevedo-Rdgz., P. 9334; 

Birdsey, s.n. 

Monstera adansonii: Acevedo-Rdgz., P. 9313. 

Philodendron consanguineum: Breckon, G. 4264; Britton, 

N.L. 1656; 2586; 4313; Goll, G.P. 1032; Sargent, F.H. 342; 

Sintenis, P. 4347; Stevenson, J.A. 3494; Underwood, L.M. 

753. 

Philodendron giganteum: Acevedo-Rdgz., P. 4165; 10806; 

Britton, N.L. 1685; 4312; Sintenis, P. 757. 

Philodendron hederaceum: Acevedo-Rdgz., P. 2610; 2880; 

11360; Britton, N.L. 1691; 4489; 4544. 

Philodendron ligulatum: Acevedo-Rdgz., P. 11875; Britton, 

N.L. 2139; Sargent, F.H. 559; Sintenis, P. 1746. 

Philodendron ornatum: Proctor, G.R. 47073; 50206; 

Woodbury, R.O. s.n 

Syngonium podophyllum: Acevedo-Rdgz., P. 4061; 6968; 

7122; 9326; 9331; 9351. 

ASPARAGACEAE 

Protasparagus setaceus: Acevedo-Rdgz., P. 7067; 9349. 

CYPERACEAE 

Scleria canescens: Acevedo-Rdgz., P. 11449; González Más 

1410. Scleria scindens: Acevedo-Rdgz., P. 2606; 2884; 

5102; 7150. Scleria secans Britton, N. L. & Cowell 1402. 

DIOSCORACEAE 

Dioscorea alata: Acevedo-Rdgz., P. 7047; 7076; Goll, G.P. 137; 

1066; Sargent, F.H. B48; Sintenis, P. 448; 4509; Stevenson, 


Dioscorea bulbifera: Acevedo-Rdgz., P. 9369; 11701. 

Dioscorea esculenta: Axelrod, F. 10051. 

Dioscorea pilosiuscula: Acevedo-Rdgz., P. 399; 2419; 2645; 

Sintenis 1384b. 

Dioscorea polygonoides: Acevedo-Rdgz., P. 140; 2977; 7078; 

9317; 10812. 

Dioscorea trifida: Sargent, F.H. 500. 

Rajania cordata: Acevedo-Rdgz., P. 127; 5171; 5226; 6962; 

7035; 7104; 7147; 7664; 7922; 7925; 7927; 9403; 9437; 

10220; 10570; 10820; 10877; 10878; 11208; 11381; 

11406; Axelrod, F. 11069; Britton, N.L. 947; 1732; Goll, 

G.P. 238; 1033; Liogier, A.H. 9701; 9866; Sargent, F.H. 

385; 3159; Shafer, J.A. 2631; 3253; 3606; 3655; Sintenis, 

P. 1384a; Stevenson, J.A. 158; 3113; Underwood, L.M. 

739; Webster, G.L. 8738. 

ORCHIDACEAE 

Vanilla barbellata: Acevedo-Rdgz., P. 4142; 10953; 11389. 

Vanilla claviculata: 7205. 

Vanilla dilloniana: Acevedo-Rdgz., P. 7058; 7182. 

Vanilla mexicana: Axelrod, F. 5639; Eggers, B. 1322. 

Vanilla planifolia: Acevedo-Rdgz., P. 4058; 7077; 9323. 

Vanilla poitaei: Acevedo-Rdgz., P. 9387. 

POACEAE 

Arthrostylidium farctum: Acevedo-Rdgz., P. 2490; 2652; 

10742; 11639; Chase, A. 6248; Cowles, H.T. 1166; Shafer, 

J.A. 2625; Sintenis, P. 3891. 

Arthrostylidium multispicatum: Acevedo-Rdgz., P. 9422; 

11441; Britton, N.L. 953; 4222; 6542; 7300; Chase, A. 

6201; 6470; 6751; Sintenis, P. 209; 4106. 

Arthrostylidium sarmentosum: Acevedo-Rdgz., P. 6963; 

9327; Britton, N.L. 493; 2042; 2180; 4209; 5236; 5600; 

6096; 7283; Chase, A. 6190; 6223; 6468; 6730; 6731; 

6738; 6749; Clark, L.G. 229; Jones, G.N. 11014; Liogier, 

A.H. 9724; Nees, W.E. 116; Proctor, G.R. 42301; Shafer, 

J.A. 3490; Sintenis, P. 354; 4046; Soderstrom, T.R. 1804; 

1816; 2001; 2053. 

Chusquea abietifolia: Edelman, D.K. 58; 59; 60; Hess, W.E. 

116; Sargent, F.H. 3062; Stevens, F.L. 4755. 

Lasiacis divaricata: Acevedo-Rdgz., P. 213; 825; 1886; 2288; 

2415; 2638; 2681; 3169; 3795; 5218; 5247; 6984; 7155; 

10488; 10574; 10931; 11429; Barrett 9; Britton, N.L. 

2623; Chase, A. 6224; 6225; 6335; 6365; 6379; 6420; 

6521; 6532; 6543; 6560; 6578; 6587; 6610; 6683; 6726; 

6742; 6782; 6814; Edelman, D.K. 2; Hess, W.E. 454; 

Otero, J. 255; Sintenis, P. 2470; Underwood, L.M. 144. 

Lasciacis ligulata: Acevedo-Rdgz., P. 2688; 10822; Boom, B. 

6911; Britton, N.L. 678; 4490; Chase, A. 6454; 6734; 

6747; Liogier, A.H. 10418; Sintenis, P. 215; 5918; Wilson, 

P. 350. 

Lasiacis sorghoidea: Acevedo-Rdgz., P. 3844; 3853; Britton, 

N.L. 6450; Chase, A. 6218; 6419; 6457; 6728; 6760; 

6809; Heller, A.A. 4375; Hess, W.E. 75; Holm 74; Shafer, 

J.A. 2570; Sintenis, P. 2861; 3062. 

Olyra latifolia: Acevedo-Rdgz., P. 5123; 5153; 5246; 7156; 

Axelrod, F. 5080; Chase, A. 6175; 6200; 6334; 6416; 6570; 

6645; 6732; Heller, A.A. 4583; McKee 10601; Sintenis, 

P. 138; 2396; 4764; Soderstrom, T.R. 1802; Stimson 1229; 

Wetmore 172. 

SMILACACEAE 

Smilax coriacea: Acevedo-Rdgz., P. 3818; 7211; 9339; 11395; 

11418; 11848; Axelrod, F. 11103; 11160. 

Smilax domingensis: Acevedo-Rdgz., P. 183; 315; 330; 2184; 

7129; 9330; 9488; 11835; Axelrod, F. 9492; Sargent, F.H. 

362; Sintenis, P. 1417; Stevenson, J.A. 3458; Proctor, G.R. 

45653; 46291; 48067; 50442.

456 


Fig. 1. 

Pisonia aculeata: (A) Acevedo-Rdgz. s.n. 

Ipomoea violacea: (B) Acevedo-Rdgz. 4007. 

Pinzona coriacea: (C) Acevedo-Rdgz. 10863. 

Chamissoa altissima: (D) Acevedo-Rdgz. 4599. 

Securidaca virgata: (E) Acevedo-Rdgz. 9454. 

Hyperbaena domingensis: (F) Acevedo-Rdgz. 3173. 

Fig. 2. 

Paullinia pinnata (A) Acevedo-Rdgz. s.n. 

Serjania polyphylla (B) Acevedo-Rdgz. s.n. 

Chiococca alba: (C) Acevedo-Rdgz. 3129. 

Turbina corymbosa: (D) Acevedo-Rdgz. 9391. 

Rhynchosia phaseoloides: (E) Acevedo-Rdgz. 9365. 

Marcgravia rectiflora: (F) Acevedo-Rdgz. s.n. 

Fig. 3. 

Distictis lactiflora: (A) Acevedo-Rdgz. s.n. 

Cydista aequinoctiale: (B) Acevedo-Rdgz. 2810. 

Amphilophium paniculatum: (C) Acevedo-Rdgz. 10620. 

Macfadyena unguis-cati: (D) Acevedo-Rdgz. 3958. 

Pristimera caribaea: (E) Acevedo-Rdgz. 7236. 

Passiflora multiflora: (F) Acevedo-Rdgz. 11704. 

Fig. 4. 

Citrullus lanatus: (A) Whiteford 4047. 

Gouania lupuloides: (B) Woodbury 17003. 

Cissus trifoliata: (C) Acevedo-Rdgz. 5127. 

Serjania lucida: (D) Acevedo-Rdgz. 709. 

Arrabidaea chica: (E) Proctor 2731. 

Smilax coriacea: (F) Britton 1376 . 

Metastelma monense: (G) Acevedo-Rdgz. et al. 4273. 

Clematis flammulastrum: (H) Britton 1762. 

Fig. 5. 

Blechnum fragile: (A-C) Wilson 142. 

Oleandra articulata: (D-E) Britton & Bruner 7570. 

Fig. 6. 

Hypolepis nigrescens: (A) Proctor 40629. 

Hypolepis repens: (B) Proctor 39399. 

Hypolepis tenerrima: (C) Proctor 40130. 

Hypolepis urbanii: (D) Ekman H. 12842. 

Fig. 7. 

Odontosoria aculeata: (A, B) Acevedo-Rdgz. et al. 2095. (C) 

Dewolf 1889. 

Odontosoria scandens: (D, E) Webster et al. 8767. (F) Sánchez 

& Liogier 204. 

Fig. 8. 

Dicranopteris flexuosa: (A) Field sketch. (B-D) Britton & 

Britton 8126. (E) Proctor 39416. 

Dicranopteris pectinata: (F, G) Cowles 348. 

ILLUSTRATION VOUCHERS 

Fig. 9. 

Gleichenia bifida: (A, B) Stimson 1282. (C) Acevedo-Rdgz. 

9373. 

Gleichenia brevipubis: (D) Proctor 39420. 

Gleichenia rubiginosa: (E) Proctor 40136. 

Fig. 10. 

Lomagramma guianensis (A) Proctor 41108. (B) Ekman 

15083. 

Lomariopsis ampdrophlebia: (C) Woodbury s.n. 

Lomariopsis kunzeana: (D) Sargent 3107. 

Lomariopsis sorbifolia: (E) Bro. Hioram 331. 

Fig. 11. 

Microgramma heterophylla: (A) Grimes 3222. (B) Acevedo- 

Rdgz. et al. 2037. 

Microgramma lycopodioides: (C, D) Liogier & Martorell 

31012. 

Microgramma piloselloides: (E) Underwood & Griggs 352. 

(F) Wilson 245. 

Polypodium loriceum: (G) Acevedo-Rdgz. 10871. 

Fig. 12. 

Lygodium japonicum: (A-C) Tang Siu Ging 6968. 

Selaginella willdenovii: (D-G) Proctor 40184. 

Fig. 13. 

Asystasia gangetica: (A-E) Acevedo-Rdgz. 3083. 

Oplonia spinosa: (F, G, I-K) Heller 4688. (H) Photograph: 

Acevedo-Rdgz. 4147. 

Fig. 14. 

Thunbergia fragrans: (A-F) field sketch, Acevedo-Rdgz. 1928. 

Thunbergia alata: (G-L) Sketch form a cultivated plant. 

Fig. 15. 

Thunbergia grandiflora: (A-F) Sketch form a cultivated plant 


Fig. 16. 

Celosia nitida: (A-E) Woodbury 172. 

Chamissoa altissima: (F) Field sketch, Acevedo-Rdgz. 9410. 

(G-J) Sintenis 213a. (K) Acevedo-Rdgz. & Siaca 3738. 

Fig. 17. 

Iresine angustifolia: (A-F) Acevedo-Rdgz. 779. 

Iresine argentata: (G-I) Britton et al. 6371. 

Iresine diffusa: (J-L) Thompson 1124. 

Fig. 18. 

Allamanda cathartica: (A-D) Sketch form a cultivated plant 

at NYBG conservatory. Anechites nerium: (E-H) Nash 601. 

Fig. 19. 

Forsteronia portoricensis: (A-F) Acevedo-Rdgz. et al. 7131.


Pentalinon luteum: (G-I) Acevedo-Rdgz. 685. (J) Acevedo- 

Rdgz. 1835. 

Fig. 20. 

Echites agglutinata: (A-F) Zanoni 35972. 

Rhabdadenia biflora (G, I-K) Acevedo-Rdgz. et al 6347. (H) 

Photograph from Costa Rica. 

Fig. 21. 

Aristolochia anguicida: (A, D) Ricksecker 85. (B) Duss 4725. 

(C) Photograph de F. González (COL). 

Aristolochia oblongata subsp. calceiformis: (E, H) Fuertes 

1879. (F, J) Axelrod & Luckow 3417. (I) Ekman H. 7899. 

Fig. 22. 

Aristolochia elegans: (A-F) Acevedo-Rdgz. 4136 

Aristolochia trilobata: (G) Photograph from Guyana. (H) 

Marcano 8217. 

Fig. 23. 

Aristolochia grandiflora: (A) Morley & Whiteford 561. (B, 

C) Photograph of Gentry 32509 growing at NYBG 

conservatory). 

Fig. 24. 

Aristolochia odoratissima: (A) Nee 41844; Photograph of F. 

González 3574. (B) Britton & Shafer 338 

Aristolochia ringens: (C) Photograph of F. González; Heller 

861. (D) Allard 18080. (E, F) González s.n. 

Fig. 25. 

Cryptostegia madagascarensis: (A-D) Field sketch. 

Cryptostegia grandiflora: (E, G) Ekman 8500. (H) Acevedo- 

Rdgz. 8511. 

Fig. 26. 

Gonolobus stephanotrichus: (A-D, F) Field sketch, Acevedo- 

Rdgz. 10580. (B-C) Baker 3052. (G) Sintenis 4666. 

Hoya australis: (H-K) Acevedo-Rdgz. 7044. 

Fig. 27. 

Marsdenia elliptica: (A) Axelrod 9399. (B-D) Howard 6392. 

Marsdenia floribunda: (F-H) Cultivated, not vouchered. (E) 

Acuña & Roig 19968. 

Fig. 28. 

Marsdenia woodburyana: (A-D) L. Ramírez & A. Rosado 

27. (E-F) Proctor 39358. 

Fig. 29. 

Matelea maritima: (A-D) Acevedo-Rdgz. 2737. 

Matelea sintenisii: (E-H) Proctor 40302. 

Matelea variifolia: (I-L) Woodbury s.n. 

Fig. 30. 

Metastelma anegadense: (A-D) Britton & Fishlock 962. 

Metastelma decipiens: (E-H) Acevedo-Rdgz. 1815. 

Metastelma leptocladon: (I) Field sketch, not vouchered. (J- 

N) Acevedo-Rdgz. et al. 7714. (O) Acevedo-Rdgz. 3754. 

Fig. 31. 

Metastelma lineare: (A-E) Field sketch, Acevedo-Rdgz. 9361. 

(F) Acevedo-Rdgz. & Siaca 3746. 

Metastelma monense: (G-K) Acevedo-Rdgz. et al. 4273. 

Metastelma parviflorum: (L-P) Axelrod & Ackerman 268. 

Fig. 32. 

Oxypetalum cordifolium: (A-E) Cuatrecasas 14430. (F) Jack 

6943. 

Funastrum clausum: (G-J) Proctor 31087. 

Fig. 33. 

Berylsimpsonia vanillosma: (A-C) King & Proctor 10601. 

Bidens reptans: (D-G) Acevedo-Rdgz. 7807. 

Bidens urbanii: (H-I) Acevedo-Rdgz. & Siaca 3745. (J) 

Acevedo-Rdgz. & Siaca 3735. 

Fig. 34. 

Chromolaena borinquensis: (A) Acevedo-Rdgz. 9367. (B-D) 

Acevedo-Rdgz. & Axelrod 7779. 

Koanophyllon polyodon: (E-F) Axelrod et al. 3594. 

Lepidaploa borinquensis: (G-K) Acevedo-Rdgz. 7924. 

Fig. 35. 

Mikania congesta: (A-D) Axelrod 3118. 

Mikania cordifolia: (E-H) Duss 2814. 

Fig. 36. 

Mikania fragilis: (A, B) Field sketch, not vouchered. (C, D) 


Mikania micrantha: (E-H) Field sketch, Acevedo-Rdgz. 9406. 

Fig. 37. 

Mikania odoratissima: (A-C) Sargent 8115. 

Mikania pachyphylla: (D, F, G) Proctor 43542. (E) Axelrod 

4288. 

Fig. 38. 

Mikania porosa: (A-C) Field sketch, Acevedo-Rdgz. 9356. 

Mikania stevensiana: (D-F) Acevedo-Rdgz. 7169. 

Fig. 39. 

Piptocarpha tetrantha: (A-C) Acevedo-Rdgz. 2974. 

Pseudogynoxys chenopodioides: (D-G) Field sketch, not 

vouchered. 

Fig. 40. 

Piptocoma acevedoi: (A-H) Acevedo-Rdgz & Chinea 5217. 

Fig. 41. 

Salmea scandens: (A-C) Liogier 10711. 

Sphagneticola trilobata: (D-H) Acevedo-Rdgz. 2794. (I) 

Zanoni 10417 

Fig. 42. 

Anredera vesicaria: (A-G) Zanoni 37125. 

Anredera cordifolia: (H-J) Brown & Britton 78. 

Basella alba: (K-N) Schultes & Black 8138. 

Fig. 43. 

Amphilophium paniculatum: (A-C) Ernst et al. 2104. 

Arrabidaea chica: (D-F) Proctor 27391. (G) Saunders 604. 

Fig. 44. 

Cydista aequinoctialis: (A-C) Acevedo-Rdgz. 2810. (D) 

Questel 575. 

Distictis lactiflora: (E, F) Acevedo-Rdgz. 4660. (G) Sintenis 

3091. (H) Sintenis 3304.

458 


Fig. 45. 

Macfadyena unguis-cati: (A, B) Holdridge 1074. (C) 


Mansoa hymenaea: (D-G) Field sketch, not vouchered. 

Mansoa alliacea: (H) Ducke 7713. 

Fig. 46. 

Phryganocydia corymbosa: (A-C) Fosberg 58885. 

Podranea ricasoliana: (D-F) Acevedo-Rdgz. 7920. (G) Britton 

& Britton 9900. 

Fig. 47. 

Pyrostegia venusta: (A-D) Mori et al. 12294. (E) Regnell s.n. 

Saritaea magnifica: (F-H) Acevedo-Rdgz. et al. 6992. 

Fig. 48. 

Tecomaria capensis: (A-C) Morrow 61. 

Tynanthus polyanthus: (D, E) Abbott 330. (F-I) Kirkbride & 

Vera 52787. 

Fig. 49. 

Cordia bellonis: (A-F) Axelrod & Rivera 8469 

Cordia polycephala: (G-L) Axelrod & Axelrod 3064 

Fig. 50. 

Tournefortia bicolor: (A) Sintenis 6211. (B-E) Sargent B-70 

Tournefortia hirsutissima: (F-I) Acevedo-Rdgz. 2865. (J) 

Photograph, Acevedo-Rdgz. s.n. 

Fig. 51. 

Tournefortia maculata: (A) Sargent 184. Field sketch, not 

vouchered. (B-E) Sargent 184. (F) Britton et al. 2485. 

Fig. 52. 

Tournefortia microphylla: (A-C) Acevedo-Rdgz. 777. (D) 

Photograph, Acevedo-Rdgz. 2326. 

Tournefortia scabra: (E-G) Acevedo-Rdgz. 7219. 

Tournefortia volubilis: (H-K) Ernst 1636. 

Fig. 53. 

Buddleja madagascariensis: (A-E) Liogier 33103. 

Fig. 54. 

Hylocereus trigonus: (A-C) Acevedo-Rdgz. 2874. 

Hylocereus undatus: (D) Photograph, Acevedo-Rdgz. s.n. (E) 

Photograph of Juan T. Roig, Cuba 1912. 

Fig. 55. 

Pereskia aculeata: (A-C, E) Acevedo-Rdgz. 4267. 

(Photograph). (D) Redrawn form Memoirs of the NYBG. 

Selenicereus grandiflorus: (F) Rose 06.210, Photograph. (G) 


Fig. 56. 

Capparis flexuosa: (A, B) Acevedo-Rdgz. 4688. (C) Prance 

et al. 29290. (D) Woodbury 341 (left), Luteyn 5145. (E) 

Photograph, Acevedo-Rdgz. 2521. 

Fig. 57. 

Lonicera japonica: (A-E) Sketch of plant cultivated at NYBG 

conservatory 

Fig. 58. 

Hippocratea volubilis: (A-H) From S. Mori et al. 2002. Guide 

to the vascular plants of central French Guiana. Mem. New York 

Bot. Gard. 76. 

Fig. 59. 

Pristimera caribaea: (A, B, D-F) Acevedo-Rdgz. 10601. (C) 

Sargent 16 

Fig. 60. 

Clusia gundlachii: (A) Field sketch, not vouchered. (B-D) 

Howard 16830. (E-G) Photographs, Acevedo-Rdgz. 9318. 

Fig. 61. 

Combretum grandiflorum: (A-C) deWolf 2036. (D) Liogier 

31966. Combretum indicum: (E, F) Field sketch, not 

vouchered. (G) Ricksecker 337. 

Fig. 62. 

Rourea surinamensis: (A-E) Proctor 41586. (F-H) Field 

sketch, not vouchered. 

Fig. 63. 

Aniseia martinisensis: (A) Axelrod 5913. (B, C) Proctor 

37514. (D) Liogier 33579. 

Argyreia nervosa: (E) Arnoldo-Broeder 3117. (F) Leonard 

7447. 

Convolvulus nodiflorus: (G-K) Acevedo-Rdgz. 3960. 

Fig. 64. 

Ipomoea alba: (A-D) Field sketch, Acevedo-Rdgz. 10775. (E) 

Field sketch, not vouchered. 

Ipomoea batatas: (F-G) Ricksecker 228. 

Fig. 65. 

Ipomoea calantha: (A, B) Ekman 1471. (C) Ekman 7093. 

Ipomoea eggersii: (D-E) Acevedo-Rdgz. 4043. 

Ipomoea hederifolia: (F, G) Acevedo-Rdgz. 3031. (H) 


Fig. 66. 

Ipomoea horsfalliae: (A) Sintenis 450. (B) Acevedo-Rdgz. & 

Breckon 7837. 

Ipomoea imperati: (C, D) Acevedo-Rdgz. & Cedeño 9294. 

(E) Sintenis 976 

Ipomoea indica var. acuminata: (F) Acevedo-Rdgz. 4051. (G- 

I) Acevedo-Rdgz. 4127. 

Fig. 67. 

Ipomoea meyeri: (A, D) Heller 6225. (B) Ekman 2436. (C) 

Sintenis 5543. (E) Ekman 10916. 

Ipomoea microdactyla: (F-G) Curtiss 211. (H-J) Correll 

49280. 

Ipomoea nil: (K) Acevedo-Rdgz. 2315. (L-N) Acevedo-Rdgz. 

3080 (pickled). (O) Eggers s.n. 

Fig. 68. 

Ipomoea ochracea: (A, B) Acevedo-Rdgz. 10513 and field 

sketch. (C) Acevedo-Rdgz. 7718. 

Ipomoea purpurea: (D-E) Heller 1410 and from cultivated 

plant. (F) Cultivated plant. 

Ipomoea pes-caprae: (G) Field sketch, not vouchered. (H) Goll 

957.


Fig. 69. 

Ipomoea quamoclit: (A, B) Ricksecker 26. 

Ipomoea repanda: (C-F) Field sketch, not vouchered. 

Fig. 70. 

Ipomoea setifera: (A, B) Field sketch, not collected. 

Ipomoea steudelii: (C-E) Acevedo-Rdgz. 10783. (F, G) 


Fig. 71. 

Ipomoea tenuissima: (A-C) Curtiss 495. 

Ipomoea tiliacea: (D-H) Field sketch, not vouchered. 

Ipomoea tricolor: (I) Cultivated plant. (J, K) Goll 797. 

Fig. 72. 

Ipomoea triloba: (A, B) Acevedo-Rdgz. 3099. (C) Acevedo- 

Rdgz. 3079. 

Ipomoea wrightii: (D, F-H) Jack 6711. (E) Sintenis 3619. 

Ipomoea violacea: (I) Acevedo-Rdgz. 4007. (J, K) Fosberg 

55339. 

Fig. 73. 

Jacquemontia cayensis: (A, B) Correll & Popenoe 51309. 

(C) Millspaugh 9257. 

Jacquemontia cumanensis: (D) Axelrod & Escobar 2906. (E, 

F) Atha & Zanoni 723. 

Jacquemontia havanensis: (G, H) Acevedo-Rdgz. 4077. (I) 

Proctor 42607. 

Fig. 74. 

Jacquemontia pentanthos: (A, B) Acevedo-Rdgz. 4037. (C, 

D) Acevedo-Rdgz. 687. 

Jacquemontia solanifolia: (E, F) Field sketch, Acevedo-Rdgz. 

10788. (G, H) Sintenis 5681. 

Jacquemontia tamnifolia: (I-K) Heller 6437. 

Jacquemontia verticillata: (L) Leonard s.n. (M, N) Britton et 

al. 12927. 

Fig. 75. 

Merremia aegyptia: (A-E) Acevedo-Rdgz. 3080. 

Merremia cissoides: (F-G) Acevedo-Rdgz. 9348. 

Merremia dissecta: (H-K) Acevedo-Rdgz. & Reilly 1987. 

Fig. 76. 

Merremia quinquefolia: (A, D) Acevedo-Rdgz. 3081. (B, C) 


Merremia tuberosa: (E) Liogier 31166. (F) Zanoni 33435. 

(G) Feucht 629. 

Merremia umbellata: (H-J) Acevedo-Rdgz. 4019. (K, L) 


Fig. 77. 

Operculina turpethum: (A, B) Eggers 152. (C) Bristol 2331 

Poranopsis paniculata: (D-G) Field sketch, not vouchered. 

Fig. 78. 

Stictocardia tiliifolia (A-C) Acevedo-Rdgz 4008 and 3120. (D) 

Proctor 43440. 

Fig. 79. 

Turbina corymbosa: (A-D) Field sketch, Acevedo-Rdgz. 

10756. (E, G) Acevedo-Rdgz. 9391. 

Xenostegia tridentata: (H-J, M) Axelrod 5237. (J, K) Taylor 

7658. 

Fig. 80. 

Cayaponia americana: (A, B) Photograph, Acevedo-Rdgz. 

3856. 

Cayaponia racemosa: (C) Allard 14478. (D, E) Axelrod & 

Thomas 6087. (F) Molina 24700. (G) Stevenson 772. 

Fig. 81. 

Coccinia grandis: (A, B) Boom et al. 8040. (C) Fosberg 58912. 

(D) Hahn 4810. 

Citrullus lanatus: (E-I) Acevedo-Rdgz. 9347. (J) Field sketch, 

not vouchered. 

Fig. 82. 

Cucumis anguria: (A-D) Field sketch, Acevedo-Rdgz. 4023. 

Cucumis dipsaceus: (E) Ndegwa 297. (F) Burger 1345. 

Cucumis melo: (G-I) Cultivated plant, not vouchered. 

Fig. 83. 

Cucurbita moschata: (A, B) Field sketch, not vouchered. 

Doyerea emetocathartica: (C-E) Acevedo-Rdgz. 4044. (F) 


Fig. 84. 

Fevillea cordifolia: (A-D) Field sketch, Acevedo-Rdgz. 9419. 

(E) Cid & Nelson 2608. (F) Cuatrecasas 10915. 

Lagenaria siceraria: (G) Cultivated plant, not vouchered. (H- 

J) Photographs and Pedersen 8768. (K) Photos, Acevedo-Rdgz. 

11603. 

Fig. 85. 

Luffa aegyptiaca: (A-C) Acevedo-Rdgz. 4066. (B) B. León 

502. 

Luffa acutangula: (D) Shafer 480. (E) Unvouchered 

photograph. 

Fig. 86. 

Melothria pendula (A-E) and Momordica charantia: (F-K) 

From S. Mori, et al. 2002. Guide to the vascular plants of central 

French Guiana. Mem. New York Bot. Gard. 76(2). 

Fig. 87 

Psiguria ottoniana: (A) Rogel 390. (B) Pleé 42. 

Psiguria pedata: (C-F) Field sketch and unvouchered alcohol 

collection. (G) Axelrod 8405. 

Psiguria trifoliata (H) Goll 627. 

Fig. 88. 

Sechium edule: (A-E) Field sketch, not vouchered. 

Sicana odorifera: (F) Calderon 2062 and Webster 12752. (G) 

Photograph, Acevedo-Rdgz. s.n. 

Fig. 89. 

Cuscuta americana: (A-H) Acevedo-Rdgz. 2569. 

Fig. 90. 

Doliocarpus brevipedicellatus: (A-J) From S. Mori et al. 2002. 

Guide to the vascular plants of central French Guiana. Mem. 

New York Bot. Gard. 76(2). 

Fig. 91. 

Pinzona coriacea: (A-K) From S. Mori et al. 2002. Guide to 

the vascular plants of central French Guiana. Mem. New York 

Bot. Gard. 76(2).

460 


Fig. 92. 

Gonocalyx concolor: (A-D) Axelrod 6643. 

Gonocalyx portoricensis: (E, F) Webster & Mill. 8697. (G, H) 

Luteyn s.n. alcohol collection. (I) Field sketch, not vouchered. 

Vaccinium racemosum: (J-M) Field sketch, not vouchered. 

Fig. 93. 

Dalechampia scandens: (A-D) Acevedo-Rdgz. 1882. (B) 

Thompson 1089. (C) Bro. León 630. (E) Acevedo-Rdgz. 660. 

Tragia volubilis: (F-J) Acevedo-Rdgz. 703. 

Fig. 94. 

Caesalpinia bonduc (A) Acevedo-Rdgz. 2570 and field sketch. 

(B, C) Acevedo-Rdgz. 4021. 

Caesalpinia ciliata (D-F) Acevedo-Rdgz. 8227. 

Caesalpinia culebrae (G) Britton 79. 

Fig. 95. 

Caesalpinia decapetala: (A) Sargent 560. (B) Axelrod & Sastre 

6013. (C) Sargent 3284. 

Caesalpinia major (D-E) Sauleda 3698. (F) Brace 6779. 

Caesalpinia portoricensis (G-I) Britton et al. 4916. 

Fig. 96. 

Senna bicapsularis: (A-C) Field sketch. (D-F) Acevedo-Rdgz. 

3917. 

Senna nitida: (G-I) Nee 44141. (J) Liogier et al. 28351. 

Fig. 97. 

Abrus precatorius: (A) Acevedo-Rdgz. 710. (B, C) Yuncker 

17303. (D) Acevedo-Rdgz., s.n. , alcohol collection. 

Barbiera pinnata: (E-H) Almeda 7472. (I) Sintenis s.n. 

Fig. 98. 

Calopogonium caeruleum: (A-E) Field sketch, not vouchered. 

(F) Axelrod 4177. 

Calopogonium mucunoides: (G-I) Heller 440. 

Fig. 99. 

Canavalia ensiformis: (A) Yuncker 18150. (B) Thompson 598 

(C) F.C.I. 01931. (D, E) Duss 1077. 

Canavalia nitida: (F, G) Correll & Correll 50614. 

Canavalia rosea: (H-K) Acevedo-Rdgz. 776. (L) Zanoni 

17093. 

Fig. 100. 

Centrosema plumieri: (A-E) Field sketch, Acevedo-Rdgz. 

10770. (F) Zanoni et al. 18954. 

Centrosema pubescens: (G-I) Field sketch, not vouchered. 

Centrosema virginianum: (J) Mori 17093. (K-N) Acevedo- 

Rdgz. 1424. 

Fig. 101. 

Clitoria falcata: (A) Axelrod 5244. (B) Photograph, Acevedo- 

Rdgz. s.n. (C) Britton et al. 6662. 

Clitoria ternatea: (D) Unvouchered cultivated plant and Taylor 

9334. (H) Britton 920 (left) Acevedo-Rdgz. 2623 (right). 

Fig. 102. 

Dalbergia ecastaphyllum: (A, E) Acevedo-Rdgz. 2034. (B- 

D) Acevedo-Rdgz. 2041. 

Dalbergia monetaria: (F-I) Filed sketch. 

Fig. 103. 

Desmodium axillare var. acutifolium: (A, B) Woodbury et al. 

s.n. (C) Taylor 10160. 

Desmodium incanum: (D-G) Acevedo-Rdgz. 4159. 

Desmodium intortum: (H, J-M) Acevedo-Rdgz. 3761. (I) 

Axelrod 3770. 

Fig. 104. 

Dioclea reflexa: (A) Proctor 51109. (B) Sintenis 5311. (C-G) 

Axelrod & Royowitz 10825. (H) Britton & Shafer 1677 and 

Proctor 51109 (fruit). 

Fig. 105. 

Galactia dubia: (A-D) Acevedo-Rdgz. 1429. (E) Liogier 

10806. 

Galactia eggersii: (F) Eggers s.n. (G-H) Acevedo-Rdgz. 4138. 

Galactia longifolia: (I-K) Box 907. (L) Liogier 37048. 

Galactia striata: (M-O) Acevedo-Rdgz 10782, Field sketch and 

alcohol collection. 

Fig. 106. 

Lablab purpureus: (A-F) Acevedo-Rdgz. 2123. 

Machaerium lunatum: (G, K, L) Heller 823. (H-J) Acevedo- 

Rdgz. 2829. 

Fig. 107. 

Macroptilium atropurpureum: (A, D) Whiteford 7162. (B, C). 

Haught 6291. 

Macroptilium lathyroides: (E-I) Cultivated, unvouchered. (J) 


Fig. 108. 

Mucuna pruriens: (A-E) Field sketch and alcohol collection, 


Mucuna sloanei: (F, G) Eggers 1317. (H) Sintenis 5941. 

Mucuna urens: (I, K-M) Acevedo-Rdgz. 9405. (J) Zanoni 

44140. 

Fig. 109. 

Neorudolphia volubilis: (A) Field sketch, not vouchered. (B) 

Acevedo-Rdgz. 2984. (C) Boom 7069. (D-F) Axelrod 4865. 

(G-H) Grimes 3255. 

Fig. 110. 

Pachyrhizus erosus: (A-D) Stehlé 4507. (E) Acevedo-Rdgz. 

5225. 

Phaseolus lunatus: (F-H) Acevedo-Rdgz. 3741. (I) Sintenis 

5778. 

Phaseolus vulgaris: (J-L) Ekman 1793. (M) Goll 747. 

Fig. 111. 

Pueraria phaseoloides: (A) Liogier 28042. (B-D) Lavestre 

1826. 

Rhynchosia minima: (E-G) Shafer 2484. (H) Sintenis 1176. 

Rhynchosia reticulata: (I-K) Mori 17094. 

Fig. 112. 

Rhynchosia phaseoloides: (A-H) From S. Mori et al. 2002. 

Guide to the vascular plants of central French Guiana. Mem. 

New York Bot. Gard. 76(2). 

Fig. 113. 

Teramnus labialis: (A-D) Acevedo-Rdgz. 1956. (E) Acevedo- 

Rdgz. 716.


Teramnus uncinatus (F) Acevedo-Rdgz. 7055. (G-I) Sintenis 

2961. 

Fig. 114. 

Vigna antillana: (A) Acevedo-Rdgz 3796. (B-C) Acevedo-Rdgz 

3868. (D) Hess 5115. 

Vigna adenantha: (E-G) Acevedo-Rdgz 10772. (H) Acevedo- 

Rdgz 10167. 

Vigna hosei: (I-K) Acevedo-Rdgz. 10677. (L) Acevedo-Rdgz. 

9456. 

Fig. 115. 

Vigna longifolia: (A) Britton et al. 6751. (B, C) Stevenson 

2097. 

Vigna luteola: (D, F) Stimson 3158. (G, H) Fosberg 48305. 

Vigna marina: (I-J) Stevenson 1686. 

Fig. 116. 

Vigna peduncularis: (A-C) Acevedo-Rdgz 10762 (D) Acevedo- 

Rdgz 10778. 

Vigna unguiculata: (E, F) Acevedo-Rdgz. 9411; 10859. 

Vigna vexillata: (G-J) Axelrod 10478, field sketch and alcohol 

collection. (K) Sintenis 1095. 

Fig. 117. 

Acacia retusa: (A-C) Acevedo-Rdgz. 5050. (D-F) Acevedo- 

Rdgz. 4055. 

Acacia vogeliana: (G, J) Acevedo-Rdgz. 5047. (H, I) Acevedo- 

Rdgz. 3794. 

Fig. 118. 

Entada polystachya var. polyphylla: (A-C) Guedes 299 and . 

s.n., photo. (D) Henkel 3580. 

Mimosa casta: (E-G) Woodbury s.n. (H) Acevedo-Rdgz. 

10830. 

Fig. 119. 

Mimosa ceratonia: (A-C) Mori 17024. (D) Zanoni 29326. 

Mimosa diplotricha: (E-G) Acevedo-Rdgz. 7045. (H) 

Anderson 8803 (Brazil). 

Mimosa quadrivalvis var. urbaniana: (I) Liogier 36196. 

Fig. 120. 

Cassytha filiformis: (A) Acevedo-Rdgz. 4372. (B-F) Acevedo- 

Rdgz 3952. 

Fig. 121. 

Heteropterys laurifolia: (A-E) Acevedo-Rdgz. 5231. (F) 


Heteropterys purpurea: (G-K) Acevedo-Rdgz. 664 and 


Heteropterys wydleriana: (L-R) Acevedo-Rdgz. & Chinea 

2211. (S) Acevedo-Rdgz. et al. 4793 

Fig. 122. 

Stigmaphyllon bannisterioides: (A-C) Acevedo-Rdgz. 9345 

and photo. (B) Hahn 3826 (Guyana). 

Stigmaphyllon emarginatum: (D-G) Acevedo-Rdgz. 4042 and 

field sketch. 

Stigmaphyllon floribunda: (H-J, L, M) Acevedo-Rdgz. 10780 

and field sketch. (K) Acevedo-Rdgz. 10205. 

Fig. 123. 

Stigmaphyllon puberum: (A-G) Ernst 129a. 

Tetrapterys inaequalis: (H, I) Urban 2494. (J-L) Duss 1469. 

(M) Urban 3086. (N) Heller 3086. 

Fig. 124. 

Marcgravia rectiflora: (A, C-E) Acevedo-Rdgz. 9404 and field 

sketch. (B, F, G) Field sketch, not vouchered. 

Marcgravia sintenisii: (H-K) Field sketch, not vouchered. 

Fig. 125. 

Cissampelos pareira: (A) Mori 17021. (B-D) Acevedo-Rdgz. 

826. (E-J) Acevedo-Rdgz. 2439. 

Fig. 126. 

Hyperbaena domingensis (A, B, F) Daly 3301. (C, E) Diaz 

333. (G) Howard 19774. (H) Duss 3682. 

Hyperbaena laurifolia (I-O) Holdridge 24. (J, K) Séller 1355. 

(L, M) Stevenson 2186. (N) Llano s.n. (O) Britton 1918. 

Fig. 127. 

Ficus citrifolia: (A-C) Acevedo-Rdgz. 1967 

Ficus pumila: (D-L) Acevedo-Rdgz.9459. 

Fig. 128 

Boerhavia scandens: (A-D, F) Acevedo-Rdgz. 10801, Field 

sketch and pickled collection. (E) Heller 6090. 

Pisonia aculeata: (G-J) Acevedo-Rdgz. 4209. (K-M) Rose 

3598. 

Fig. 129. 

Bougainvillea glabra: (A-E) Field sketch and pickled 

collection. (F) Urban 698. 

Bougainvillea spectabilis: (G-H) Hermann 2788. 

Fig. 130. 

Jasminum fluminense: (A, D) Acevedo-Rdgz. 2892 and photo. 

(B, C) Howard 20334. 

Jasminum grandiflorum: (E) Acevedo-Rdgz. 10532. (F) 

Howard 19172. (G-I) Thompson 993. 

Fig. 131. 

Jasminum multiflorum: (A-F) Field sketch and pickled 

collection. 

Jasminum sambac (G, H) Liogier 36277. (I) Leonard 4947. 

Fig. 132. 

Passiflora berteroana: (A, C, D) Liogier 13915. (B) Liogier 

33732. 

Passiflora bilobata: (E, F) Liogier 11131. (G) Liogier 10495. 

(H, I) Acevedo-Rdgz. 7762. 

Fig. 133 

Passiflora edulis; (A-G) Acevedo-Rdgz. 834. 

Passiflora laurifolia: (H, I) Mori 17054. 

Passiflora foetida: (J, K) Acevedo-Rdgz. 1948. 

Fig 134. 

Passiflora maliformis: (A) Ekman 9796. (B) Leonard 4979. 

Passiflora multiflora: (C) Field sketch. (D, E) Acevedo-Rdgz. 

4002 and photo. (F) Acevedo-Rdgz. 3857 and photo. 

Passiflora murucuja: (G-J) Cultivated, unvouchered. 

Fig. 135. 

Passiflora quadrangularis: (A) Stevenson 6715.

462 


Passiflora rubra: (B) Field sketch. (C) , photo). (D) Axelrod 

5246. 

Fig. 136. 

Passiflora serrato-digitata: (A) Eggers 1372. (B) Axelrod 

5749. 

Passiflora sexflora: (C-E) Acevedo-Rdgz. 10745; Acevedo- 

Rdgz. 10761 and photo. (F, G) Field sketch. 

Fig. 137. 

Passiflora suberosa: (A, B, E) Acevedo-Rdgz. 10498. (C) 

Urban 871. (D) Acevedo-Rdgz. 4302. 

Passiflora tulae: (F-H) Acevedo-Rdgz. 3751. 

Fig. 138. 

Agdestis clematidea: (A-C) Palmer 50. 

Stegnosperma cubensis: (D-F) Lavestre 2240. (G, H) Liogier 

17818. 

Trichostigma octandrum: (I, M) Zanoni 25547. (J-L) Britton 

& Wheeler 113. 

Fig. 139 

Peperomia rotundifolia: (A) Mejia 15609. (B-D) Lavestre 

1553. 

Fig. 140. 

Plumbago scandens. (A-D) Hudson 754. (D-E) Rodríguez 

89. (F) Zanoni 27488. 

Fig. 141. 

Securidaca diversifolia: (A) Eggers 1219. (B) Rose 3638. (C- 

D) Liogier 31916. 

Securidaca virgata: (E-M) Field sketch and Acevedo-Rdgz. 

9453. 

Fig. 142. 

Antigonon guatemalense: (A-C) Ricksecker 16. (D-F) Archer 

433. 

Antigonon leptopus: (G-H, L, M) Acevedo-Rdgz. 1811. (I-K) 

Zanoni 24475. 

Fig. 143. 

Clematis polygama: (A) Axelrod 8473. (B-D) Liogier 31073. 

Clematis flammulastrum: (E, F) Axelrod 1547. (H, I) Axelrod 

3167. 

Clematis dioica: (J) Britton 1762. 

Fig. 144. 

Gouania lupuloides: (A-D) Acevedo-Rdgz. 2295. (E-H) 

Woodbury s.n. 

Gouania polygama: (I, K-L) Atha 752. (J) Sintenis 36. (M-N) 


Fig. 145. 

Rubus florulentus: (A, I, J) Gentry & Zardini 50448. (B-H) 

Sintenis 4100. 

Fig 146. 

Chiococca alba: (A, C-F) Acevedo-Rdgz. 5077. (B) Acevedo- 

Rdgz. 1914. (G-H) Acevedo-Rdgz. 3821. 

Diodia sarmentosa: (I, J, L) Liogier 29765. (K) Woodbury 

s.n. 

Hillia parasitica: (M) Acevedo-Rdgz. 2990. (N-P) Field sketch. 

Fig. 147. 

Psychotria microdon: (A-F) Liogier 15663. (G-I) Little 26127. 

Lasianthus lanceolatus: (J-L) Acevedo-Rdgz. 6964. (K) Field 

sketch. (M-P) Acevedo-Rdgz. 7834. 

Fig. 148. 

Sabicea villosa: (A-D) Woodbury s.n. 

Schradera exotica: (E, H) Field sketch. (F) Stevens 2356. (G) 

Jiménez 5096. 

Fig. 149. 

Cardiospermum halicacabum: (A-G) Field sketch, not 

collected. 

Cardiospermum grandiflorum: (H, I) Acevedo-Rdgz. 10841 

(photo) and Acevedo-Rdgz. 10843. (J-L) Liogier 35326. 

Cardiospermum corindum: (L) Acevedo-Rdgz. 4154. (M) 


Fig. 150. 

Paullinia fuscescens: (A, B, D) Fosberg 55298. (C) Britton 

103. 

Paullinia plumieri: (E, F, H) Stehlé 1591. (G) Stehlé 5719. 

Paullinia alata: (I, O) Filed sketch and Acevedo-Rdgz. 9451. 

(J) Woodbury s.n. (K-N) Underwood & Griggs 469. 

Fig. 151. 

Serjania lucida: (A) Acevedo-Rdgz. 8305. (B, F) Acevedo- 

Rdgz. 709. (C-E) Acevedo-Rdgz. 629. 

Serjania polyphylla: (G, J) Boom 10036. (H, I) Boom 10022. 

Fig. 152. 

Schlegelia brachyantha: (A-D) Field sketch, not collected. (E) 

Acevedo-Rdgz. 2964 

Fig. 153. 

Lophospermum erubescens: (A-G). Cultivated, unvouchered. 

Fig. 154. 

Lycianthes virgata: (A-E) Acevedo-Rdgz. 11437. (F) Urban 

2620. 

Solandra grandiflora: (G) Axelrod 5250 and photo. (H) 

Degener 18751. 

Fig. 155. 

Solanum lancifolium: (A-D) Acevedo-Rdgz. 5108 and field 

sketch. 

Solanum seaforthianum: (E-H) Acevedo-Rdgz. 11476. (I) 

Britton & Britton 9867. 

Solanum wendlandii: (J, L) Liogier 28469. (K) Zanoni 30535. 

Fig. 156. 

Tropaeolum majus: (A-I) Cultivated, unvouchered. 

Fig. 157. 

Celtis iguanaea: (A, B) Acevedo-Rdgz. 2691 and photo. (C-F) 

Acevedo-Rdgz. 2611, pickled. (G) Acevedo-Rdgz. 2021, 

pickled. 

Fig. 158. 

Valeriana scandens: (A, D) Acevedo-Rdgz. 9399. (B, C, E) 

Zanoni & Maas 34088


Fig. 159. 

Clerodendrum aculeatum: (A, E) Acevedo-Rdgz. 5140 and 

field sketch. (B, C, D) Acevedo-Rdgz. 2818, pickled. 

Fig. 160. 

Clerodendrum thomsonae:. (A-D) Acevedo-Rdgz. 9413. 

Clerodendrum x speciossimum: (E-G) Cultivated, not 

collected, field sketch. 

Fig. 161 

Congea tomentosa: (A-C) Liogier 32719 and Acevedo-Rdgz. 

s.n., photo. 

Holmskioldia sanguinea: (D, E) Acevedo-Rdgz. 10533 and 

Acevedo-Rdgz. s.n. , photo. 

Petrea volubilis: (F-H) Field sketch and pickled, not vouchered. 

Fig. 162. 

Cissus erosa: (A-J) From S. Mori et al. 2002. Guide to the 

vascular plants of central French Guiana. Mem. New York Bot. 

Gard. 76(2). 

Fig. 163. 

Cissus obovata: (A) Britton & Britton 9903. (B) Eggers 556. 

(C, D) Ekman 6050. 

Cissus rotundifolia: (E, F) Spellenberg 7370. 

Fig. 164. FSJ 

Cissus verticillata: (A) Acevedo-Rdgz. 5114, field sketch. (B- 

D) Acevedo-Rdgz. 4683, photograph and pickled. 

Cissus trifoliata: (E) Acevedo-Rdgz. 5127, field sketch. (F) 

Acevedo-Rdgz. 2693, pickled. 

Vitis tiliifolia: (G) Photograph from Escambray, Cuba and 

Augusto 1574. (H, I) Taylor 8086. (J) Augusto 1574. 

Fig. 165. 

Anthurium scandens: (A-F) Acevedo-Rdgz. 9395. 

Fig. 166. 

Epipremnum pinnatum: (A, B) Field sketch, not collected. (C) 


Epipremnum pinnatum cv. aureum: (D) Field sketch, not 

collected. (E) Acevedo-Rdgz. 7141. 

Fig. 167. 

Monstera adansonii. (A-C) From S. Mori et al. 1997. Guide 

to the vascular plants of central French Guiana. Mem. New York 

Bot. Gard. 76(1). 

Fig. 168. 

Philodendron consanguineum: (A) Engler 4347. (B) Liogier 

11161. 

Philodendron hederaceum: (C, D) Acevedo-Rdgz. 2880. 

Fig. 169. 

Philodendron giganteum: (A-D) Acevedo-Rdgz. 10806. 

Fig. 170. 

Philodendron lingulatum: (A) Axelrod 9192. 

Philodendron ornatum: (B) Gentry et al. 77672. (C) Proctor 

47073. 

Fig. 171. 

Syngonium podophyllum: (A,B) Mori 15131. (C) Acevedo- 

Rdgz. 4061. 

Fig. 172. 

Protoasparagus setaceus: (A-E) Acevedo-Rdgz. 9349. 

Fig. 173. 

Scleria canescens: (A, B) Heller 1090. 

Scleria scindens: (C) Acevedo-Rdgz. 5102. (D, E) Ernst 1577. 

Scleria secans: (F, G) Mejia 11127. 

Fig. 174. 

Dioscorea alta: (A, B) Acevedo-Rdgz. 7047. (C) Hansen et 

al. 9308. (D) Nee 44151. (E) Acevedo’s photos. 

Dioscorea altissima: (F) Gentry & Zardini 50388. (G) Allard 

13992. 

Fig. 175. 

Dioscorea bulbifera: (A, B) Acevedo-Rdgz. 9369 and field 

sketch. (C-E) Ekman 4732. 

Dioscorea cayenensis: (F, G) Acevedo-Rdgz. 11896, photo. 

(H) Howard 18920. (I-K) Webster & Gooding 3634. 

Dioscorea pilosiuscula: (L-P) Acevedo-Rdgz. 3991. 

Fig. 176. 

Dioscorea polygonoides: (A) Acevedo-Rdgz. 10812. (B) Boom 

6812. (C-E) Liogier 35001. (F-H) Boom 9595. (I) Shafer 3521. 

Dioscorea trifida: (J) Sargent 500. (K, L) Allard 14144. 

Fig. 177. 

Rajania cordata: (A) Stevens 2782. (B) Acevedo-Rdgz. 7147. 

(C) Grimes 3245. (D, F) Stevenson 1187 and pickled. s.n. (E) 

Howard 16165. (G-I) Axelrod 218. (J) Howard 16905. (K, L) 

Wagner 1911. 

Fig. 178. 

Vanilla barbellata: (A) Liogier 15640. (B) Liogier 15047. (C- 

E) Ackerman 2129. 

Vanilla claviculata: (F) Nee 44080. (G, H) Ackerman 2013. 

Fig. 179. 

Vanilla dilloniana: (A) field sketch. (B) Luer’s photo. (C) 

Stimson 1276. 

Vanilla mexicana: (D) Harris 8555. (E) Luer’s photo. 

Vanilla planifolia: (F, G) field sketch. 

Fig. 180. 

Vanilla poitaei: (A) Acevedo-Rdgz. 9387. (B) Pennington 

1836. (C-E) Ackerman 2016. (F) Acevedo-Rdgz. 7187. 

Vanilla pompona: (G) Duss 3487. (H) Photo. 

Fig. 181. 

Arthrostylidium farctum: (A-C) Ekman 9286. 

Arthrostylidium multispicatum: (D) Chase 6201. (E-G) Clark 

& Reiners 1508. 

Arthrostylidium sarmentosum: (H) Acevedo-Rdgz. 6963. (I, 

J) Soderstrom 2053. 

Fig. 182. 

Chusquea abietifolia: (A) Liogier 30906. (B-D) Ekman 8009. 

Lasiacis divaricata: (E-G) Goll 227. 

Lasiacis ligulata: (H-K) Acevedo-Rdgz. 2681. 

Fig. 183. 

Lasiacis sorghoidea: (A, B) Holm 74. (C, D) Cedeño 637. 

Olyra latifolia: (E-H) Mori 18848.

464 


Fig. 184. 

Smilax coriacea: (A) Martorell s.n. (B, E) Fuertes 444. (C) 

Britton 1376. (D) Liogier 35155. (F) Acevedo-Rdgz. 3818. 

Smilax domingensis: (G) Field sketch. (H, I) Proctor 48067. 

(J, K) Liogier 34627.


abaxial (abaxial)- in laminar organs such as leaves, 

sepals, and petals, refers to the lower surface 

accrescent (acrescente)- having additional growth after 

its formation 

achene (aquenio)- a one-seeded, dry, indehiscent fruit 

with the pericarp fused to the seed coat 

actinomorphic (actinomorfa)- having at least two 

planes of symmetry 

acuminate (acuminado)- terminating in an elongated 

point 

acute (agudo)- forming an angle less than 90° 

adaxial (adaxial)- in laminar organs such as leaves, 

sepals, and petals, refers to the upper surface 

adnate (adnato)- refers to the fusion (or close 

adherence) of different stuctures, e.g., a stamen 

and a petal 

adventitious (adventicio)- refers to any organ that 

develops from adult tissue 

aggregate fruit (fruto agregado)- a fruit that is made 

up of small fruitlets, the products of the apocarpous 

gynoecium of a single flower 

androgynophore (androginóforo)- a column that bears 

the stamens and the gynoecium 

anthocarp (antocarpo)- a structure consisting of a fruit 

together with the floral envelope or perianth 

apocarpous (apocárpico)- having the individual 

carpels separate 

appressed (adpreso)- pressed against something 

areole (areola)- a small cavity surrounded by the 

anastamosed (fused) veins 

arillode (arilodio)- a fleshy structure that resembles 

an aril, but whose origin is undetermined 

aristate (aristado)- terminating in a long, slender, rigid 

point 

articulate (articulado)- presenting a joint or 

articulation 

attenuate (atenuado)- narrowed 

auriculate (auriculado)- ear-shaped 

barbate (barbado)- having a beard-like tuft of hairs 

berry (baya)- a fleshy, indehiscent, simple or 

syncarpous fruit 

bicarinate (bicarinado)- having two keels or ridges 

bifurcate (bifurcado)- divided into two parts of the 

same size 

biglobose (bigloboso)- having a structure composed 

of two united but distinct spheres 

bilabiate (bilabiado)- refers to a calyx or corolla that 

is divided into two lips 

bipinnate (bipinnado)- refers to a twice-pinnate blade 

GLOSSARY 

biternate (biternado)- refers to a twice-ternate leaf 

blade, i.e., a ternate leaf in which each primary 

leaflet is divided into three secondary leaflets, for 

a total of nine secondary leaflets 

blade (lámina)- the laminar portion of the leaf, joined 

to the petiole 

bract (bráctea)- any foliaceous organ near the flowers, 

differing from the leaves by its size, coloration, 

form, or texture 

bracteole (bractéola)- diminutive of bract, referring 

to those bracts that are found on the secondary axes 

of inflorescences 

bulbil (bulbillo)- a small bulb borne in leaf axils, 

whose function is the accumulation of food reserves 

calyptra (caliptra)- a cap-like structure 

calyptrate (caliptrado)- cap-like 

cambium (cámbium)- a tissue composed of 

meristematic cells located between the wood and 

the liber (phloem) 

campanulate (campanulado)- bell-shaped 

canescent (canescente)- covered with short, white hairs 

capitate (capitado)- in the form of a head or arranged 

in capitula (heads) 

capitulum (capítulo)- an inflorescence, usually globose 

or convex, composed of sessile flowers 

caudate (caudado)- terminating in a long, tail-like 

point 

caudex (caudex)- the trunk of a palm or tree-fern 

chartaceous (cartáceo)- having a papery texture 

ciliate (ciliado)- bearing cilia 

cilium (cilio)- a fine hair along the margin of any 

laminar organ 

cincinnus (cincino)- a scorpioid cyme whose flowers 

are arranged in different planes 

circumcissile (circuncísil)- having circular dehiscence 

claviform (claviforme)- club-shaped, i.e., widening 

gradually toward the apex 

collateral (colateral)- borne on the side 

concolorous (concoloro)- of the same color 

concrescent (concrescente)- more or less united 

connate (connato)- refers to similar organs that are 

united, e.g., petals united to form a tubular corolla 

connective (conectivo)- the sterile portion of an anther, 

located between the two thecae (anther sacs) 

cordiform (cordiforme)- heart-shaped; in laminar 

organs, with the base auriculate and wider than 

the rest of the lamina 

coriaceous (coriáceo)- having a leathery texture 

corona (corona)- a group of petaloid appendages that 

form a whorl within the corolla

466 


cortical (cortical)- relative to the cortex or bark 

corymb (corimbo)- an inflorescence whose flowers are 

borne on different portions of the axis but are at 

the same height because their pedicels are of 

different lengths 

corymbiform (corimbiforme)- in the form of a corymb 

crateriform (crateriforme)- bowl-shaped 

crenate (crenado, festoneado)- having short, rounded 

protruberances along the margin, scalloped 

crenulate (crenulado)- diminutive of crenate 

crustaceous (crustáceo)- having a rigid and brittle 

texture, like an egg-shell 

culm (culmo)- the stem of the grasses and sedges 

cuneate, cuneiform (cuneado, cuneiforme)- wedgeshaped 

cupuliform (cupuliforme)- in the form of a little cup 

cyme (cima)- an inflorescence whose axis terminates 

in a flower and bears secondary axes 

cymose (cimoso)- in the form of a cyme 

deciduous (caducifolio)- refers to a plant that loses 

its leaves 

decumbent (decumbente)- lying or trailing on the 

ground, but with the tips ascending 

decurrent (decurrente)- extending below the point of 

insertion 

dehiscent (dehiscente)- opening 

deltoid (deltoide)- triangular 

dendroid (dendroide)- branching like a small tree 

dentate (dentado)- with teeth along the margin 

denticulate (denticulado)- diminutive of dentate, i.e., 

with small teeth along the margin 

dichasial (dicasial)- bearing dichasia 

dichasium (dicasio)- a cymose inflorescence whose 

axis terminates in a flower that is accompanied by 

two lateral flowers from the base 

didynamous (didínamo)- with four stamens, of which 

two are longer and two are shorter 

digitiform (digitiforme)- in the form of digits or 

fingers, i.e., deeply divided into diverging lobes 

dimorphic (dimorfo)- presenting two forms 

discoid (discoide)- disc-shaped 

discolorous (discoloro)- of two or more colors 

distichous (dístico)- arranged in two rows 

divaricate (divaricado)- applies to branches or 

secondary axes that form a very wide angle in 

relation to the principal axis 

domatium (domacio)- a small depression, usually in 

the axils of the secondary veins 

drupaceous (drupáceo)- referring to or like a drupe 

drupe (drupa)- a fleshy fruit with a stone, i.e., having 

a woody endocarp with the seed(s) inside 

ellipsoid (elipsoide)- in the form of an ellipse, but in 

three dimensions 

elliptical (elíptico)- in the form of an ellipse, i.e., 

widest in the central portion and narrowing toward 

both rounded ends 

endemic (endémico)- having a distribution limited to 

a particular place or region 

endocarp (endocarpo)- the inner tissue of the fruit wall 

epiphyte (epífito)- a plant that lives on another plant 

without parasitizing it 

exocarp (exocarpo)- the outer tissue of the fruit wall 

exserted (exserto)- exposed, projecting outward 

falcate (falcado)- sickle-shaped 

fascicle (fascículo)- a tight bundle; a tightly contracted 

cyme 

ferruginous (ferrugíneo)- rust-colored 

flabellate (flabelado)- fan-shaped 

flexuous (flexuoso)- twisted, more or less in the shape 

of a ‘z’ 

foveate (foveado)- with the surface bearing small pits 

frond (fronda)- a fern leaf 

fusiform (fusiform)- spindle-shaped 

gamopetalous (gamopétalo)- referring to a corolla 

composed of fused petals 

glabrescent (glabrescente)- losing hairs or pubescence 

at maturity 

glabrous (glabro)- lacking hairs or pubescence 

glaucous (glauco)- light green in color with a slightly 

bluish shade 

glume (gluma)- in grasses, a sterile bract at the base 

of the spikelet 

gynoecium (gineceo)- a collective term for the female 

parts of a flower 

gynophore (ginóforo)- a column that bears the 

gynoecium 

gynostegium (ginostegio)- a protective organ around 

the gynoecium that is distinct from the perianth 

harpidium (harpidio)- a hooked, claw-like tendril 

hastate (hastado)- referring to a laminar organ, pointed 

and with divergent lobes 

haustorium (haustorio)- an organ that functions as a 

sucker, through which vital juices are obtained 

hemiepiphyte (hemiepífito)- a plant that begins its 

life cycle as an epiphyte but later establishes 

contact with the soil, from which it obtains 

nutrients like a normal plant 

heterostylous (heterostilado)- having two or three 

kinds of individuals whose styles differ in length 

hilum (hilum)- a scar on the surface of the seed 

hirsute (hirsuto)- covered with rigid hairs and rough 

to the touch 

hispid (híspido)- covered with very rigid hairs and very 

rough to the touch 

hispidulous (hispiduloso)- with short, rigid hairs 

homologous (homólogo)- concordant, referring to 

organs that have the same origin 

hyaline (hialino)- transparent 

hypanthium (hipanto)- the sunken axis of a flower 

with an inferior ovary 

hypocrateriform (hipocrateriforme)- trumpet-shaped 

imbricate (imbricado)- referring to foliaceous organs 

that are closely spaced and overlap at their margins


incised (inciso)- divided more or less deeply into 

segments 

incurved (incurvado)- curved inward, with the 

concavity on the inner or upper side 

indehiscent (indehiscente)- not opening 

indeterminate (indeterminado)- growing indefinitely 

indusium (indusio)- an organ that covers and protects 

fern sporangia 

inflexed (inflexo)- curved inward 

infundibuliform (infundibuliforme)- funnel-shaped 

interstaminal (interestaminal)- between or within the 

stamens 

involucre (involucro)- a group of bracts that surrounds 

and more or less encloses the flowers 

laciniate (laciniado)- divided into deep, narrow, sharppointed 

segments 

lanceolate (lanceolado)- lance-shaped, referring to a 

laminar structure whose widest portion is below 

the middle, narrowing toward both ends 

lenticel (lenticelo)- a pore in the bark of a woody stem 

lenticellate (lenticelado)- having more or less 

conspicuous lenticels 

lenticular (lenticular)- lens- or lentil-shaped 

lepidote (lepidoto)- scaly or with trichomes in the form 

of scales 

lignescent (lignescente)- becoming woody at maturity 

limb (limbo)- in a gamopetalous corolla, the free 

portion of the corolla that forms a border at the 

end of the tube 

megaphyllous (megafilo)- with large leaves 

membranaceous (membranáceo)- like a membrane 

mericarp (mericarpo)- each of the units, composed of 

an entire carpel, into which a schizocarpic fruit 

separates 

mesocarp (mesocarpo)- the middle tissue of the fruit 

wall 

mogote (mogote) a limestone formation that results 

from the erosion and dissolution of the rock; in the 

strict sense, refers to small limestone bluffs 

surrounded by alluvion washed in from more or 

less remote areas; in the broad sense, includes other 

limestone bluff formations without alluvion at the 

base; in English, mogotes are sometimes called 

“haystack hills” 

monadelphous (monadelfo)- referring to stamens 

whose filaments are all fused 

monocarp (monocarpo)- a fruit formed from a single 

carpel 

monomorphic (monomorfo)- with all individuals 

presenting a single form 

mucronate (mucronato)- with an apex that terminates 

in a short, abrupt point 

multiseriate (multiseriado)- with several series of 

whorls 

oblanceolate (oblanceolado)- inversely lanceolate, 

with the widest portion above the middle 

oblong (oblongo)- much longer than wide and with 

parallel sides 

obtuse (obtuso)- forming an angle greater than 90° 

ocrea (ócrea)- a structure composed of a pair of connate 

stipules that forms a sheath surrounding the stem 

operculum (opérculo)- the upper portion that separates 

by circumscissile or transverse dehiscence 

orbicular (orbicular)- globose 

ovate (ovado)- egg-shaped, referring to a laminar organ 

overlapping (solapado)- referring to the characters 

used to distinguish species, these are tenuous when 

they overlap and usually show intermediate states, 

complicating the distinction of the limits between 

the forms 

ovoid (ovoide)- egg-shaped, in three dimensions 

palea (palea)- in grasses, the inner and upper of the 

two bracts that subtend a flower 

palmately compound (palmaticompuesto)- referring 

to a compound leaf whose three or more leaflets 

arise from the apex of the petiole 

palmately lobed (palmatilobado)- referring to a leaf 

whose lobes are palmately arranged 

panicle (panícula)- a compound inflorescence of 

indeterminate growth and pyramidal shape 

paniculiform (paniculiforme)- in the form of a panicle 

papilla (papila)- a very short trichome or protruberance 

of the epidermal cell membrane 

papillose (papiloso)- having papillae 

pappus (pappus)- in composites, the limb of the calyx 

transformed into simple or plumose hairs or rigid 

bristles 

paraphysis (paráfisis)- a trichome that accompanies 

the sorus 

parietal (parietal)- produced from or borne on the wall 

(of the carpel) 

patelliform (pateliforme)- shaped like a knee-cap or 

disc 

peduncle (pedúnculo)- the axis that subtends a solitary 

flower or an inflorescence 

peltate (peltado)- referring to a leaf with the petiole 

inserted in the center of a rounded blade 

pendulous (colgante)- hanging down 

perianth (perianto)- the floral envelope, i.e., sepals, 

petals, and/or tepals 

pericarp (pericarpio)- the fruit wall, composed of 

exocarp, mesocarp, and endocarp 

petaliferous appendage (apéndice petalífero)- a 

laminar structure adherent to the inner surface of 

the petal, making it appear double 

petiolule (peciólulo)- the petiole that subtends a leaflet 

of a compound leaf 

phenology (fenología)- the study of flowering and 

fruiting periods 

phloem (floema)- the portion of the conductive tissue 

responsible for food transport, usually found 

external to the xylem 

pinna (pinna)- synonym of a leaflet in compound leaves

468 


pinnate (pinnado)- referring to a leaf whose leaflets 

are arranged along an elongate rachis 

pinnately lobed (pinnatilobado)- referring to a leaf 

whose lobes are pinnately arranged 

pinnatisect (pinnatisecto)- referring to a leaf that is 

pinnately divided 

pinnule (pínnulas)- a leaflet of a bipinnate or tripinnate 

leaf 

pistillate (pistilado)- referring to female flowers 

pistillode (pistilodio)- the rudimentary pistil of a male 

flower 

plumose (plumoso)- like a bird’s feather 

pollinium (polinio)- a mass of pollen grains, 

characteristic of the families Asclepiadaceae and 

Orchidaceae 

prismatic (prismático)- prism-shaped 

proleptic (proléptico)- referring to a phenomenon 

observed in the shoot system in which lateral bud 

growth is supressed by the dominance of the apical 

meristem, so that the lateral buds develop only after 

the main shoot has attained a certain degree of 

development or maturity 

prophyll (profilo)- a bud-scale on a lateral branch 

pseudanthium (pseudanto)- an inflorescence that 

resembles a single flower due to the manner in 

which its flowers are arranged, typical of the family 

Euphorbiaceae 

pseudoracime (pseudoracimo)- an simple, racemose 

inflorescence whose flowers are borne on 

protuberances along the rachis, typical of the 

subfamily Faboideae of the family Fabaceae or 

Leguminosae 

pseudostipule (pseudoestípula)- a structure that 

occupies the position of a stipule but of different 

origin, representing a prophyll or a basal leaflet 

that surrounds the stem 

puberulent (puberulento)- having short, scattered hairs 

pubescent (pubescente, velloso)- hairy 

pulviniform (pulviniforme)- shaped like a pulvinus 

pulvinus (pulvino)- a swollen structure in the form of 

a small pillow 

pyrene (pirenio)- the bony portion of a drupe that 

contains one or several seeds 

raceme (racimo)- a simple, elongated inflorescence 

with stalked flowers 

racemose (racemoso)- in the form of a raceme 

rachis (raquis)- in a compound leaf, an axis bearing 

the petiolules or leaflets; in an inflorescence, the 

primary or secondary axis that bears the flowers 

ray (radio)- a row of radially arranged parenchymatous 

cells found in xylem tissue 

reflexed (reflexo)- bent downward 

reniform (reniforme)- kidney-shaped 

reticulate (reticulado)- forming a network or reticulum 

retroflexed (retroflexo)- curved downward 

retuse (retuso)- with a small cut or crack 

revolute (revoluto)- with the margins curved toward 

the lower surface 

rhizome (rizoma)- an underground stem 

rhombus (rombo)- a quadrilateral whose sides are 

parallel and of the same length, and whose opposite 

angles are equal 

rotate (rotáceo)- wheel-shaped, referring to a 

gamopetalous corolla with a very short tube and 

an evident limb 

rugose (rugoso)- wrinkled 

rugulose (ruguloso)- diminutive of rugose 

samara (sámara)- a dry, winged, usually 

membranaceous fruit that is wind-dispersed 

sarmentose (sarmentoso)- with long, slender, flexible 

woody branches that can support themselves on 

nearby objects 

scabrid (escábrido)- diminutive of scabrous 

scabrous (escabroso)- having a rough texture 

scandent (escandente)- climbing 

schizocarp (esquizocarpo)- an indehiscent fruit whose 

carpels separate at maturity 

secondary growth (crecimiento secundario)- growth 

in diameter of stems or other axial structures 

segment (segmento)- a portion of a pinnatisect or 

pinnately lobed leaf that reaches the midvein 

septicidal (septicida)- opening along the septum 

septum (septo)- the wall that divides the carpels in a 

syncarpous gynoecium 

sericeous (seríceo)- covered with short, appressed hairs 

serrate (aserrado)- saw-like, with small, sharp, closely 

spaced teeth 

seta (seta)- a long, stiff hair 

setose (setoso)- having a pubescence of long, stiff hairs 

setulose (setuloso)- diminutive of setose 

sheath (vaina)- a broadened leaf base that partially or 

completely surrounds the stem upon which it is 

inserted 

sigmoid (sigmoide)- in the form of an ‘s’ 

sorus (soro)- a group of sporangia located on a fern 

frond 

spadix (espádice)- a simple inflorescence with a fleshy 

central axis, surrounded by a spathe 

spathe (espata)- an elongate bract that surrounds the 

inflorescence 

spathulate (espatulado)- spatula- or spoon-shaped 

spicate (espigado)- in the form of a spike 

spike (espiga)- a simple inflorescence with sessile 

flowers 

spikelet (espícula)- the basic inflorescence of the 

grasses, composed of a very short axis that bears 

two sterile bracts (called glumes) at the base, 

followed by the flowers 

sporangium (esporangio)- a structure that contains the 

spores of a plant 

spurred (espolonado)- bearing a spur or a more or less 

deep sac 

squamose (escuamoso)- covered with scales


squamulose (escuamuloso)- diminutive of squamose, 

covered with small scales 

staminate (estaminado)- relating to the stamens or the 

male flower 

staminode (estaminodio)- a sterile stamen, more or 

less modified 

standard (estandarte)- the upper and usually the 

largest petal in the corolla of members of the 

subfamily Faboideae of the family Fabaceae or 

Leguminosae 

stellate (estrellado)- bearing radially branched, starlike 

hairs 

stipe (estípite)- the petiole of a fern frond 

stipel (estipela)- a small appendage at the base of some 

leaflets 

stipule (estípula)- a small appendage at the base of 

some leaves 

striate (estriado)- having lines, ridges, or grooves on 

the surface 

strigose (estrigoso)- bearing straight, rigid, appressed 

hairs 

strigulose (estriguloso)- diminutive of strigose 

strobilus (estróbilo)- a structure with a central axis 

bearing a group of bracts, which in turn bear or 

subtend seeds or spores 

subinvolucral (subinvolucral)- in a position below the 

involucre, referring to bracts that do not form part 

of the involucre 

subulate (subulado)- narrowing toward the apex and 

terminating in a fine point 

sulcate (sulcado)- furrowed 

supernumerary (supernumerario)- numerous 

suture (sutura)- the concrescent margin of the carpels 

syconium, syconus (sicono)- a fruit composed of a 

hollow, rounded or pear-shaped receptacle with an 

apical aperture; within the receptacle are found the 

flowers and later the fruitlets 

sympodial (simpódico)- referring to a growth pattern 

in which the principal axis shows determinate 

growth and, as a result, is displaced by an axillary 

or lateral axis 

syncarpous (sincarpo, sincárpico)- referring to ovaries 

whose carpels are fused 

tendril (zarcillo)- a touch-sensitive organ by means of 

which a vine holds on to its source of support 

tepal (tépalo)- members of a floral whorl that cannot 

be distinguished as either sepals or petals due to 

their similarity or position 

ternate (ternado)- refers to a leaf blade that is divided 

into three leaflets 

thyrse (tirso)- an inflorescence whose principal axis 

shows indeterminate growth but whose lateral units 

are cymose, i.e., with determinate growth 

thyrsoid (tirsoide)- in the form of a thyrse 

tomentose (tomentoso)- densely covered with short, 

simple or branched, interwoven hairs 

tracheid (traquea)- an imperforate xylem element 

whose principal function is the conduction of water, 

but which also functions in structural support 

translator (translator)- in some members of the family 

Asclepiadaceae, an organ that serves to transfer 

the pollen from one flower to another by means of 

insects; it is composed of the gland or clip 

(corpusculum) and the translator arms (retinacula 

or connectives) that hold the pollinia 

trichome (tricoma)- a hair 

trifid (trífido)- divided into three more or less equal 

parts 

tripinnate (tripinnada)- thrice pinnate 

triternate (triternado)- thrice ternate, resulting in 27 

leaflets, cf. ternate and biternate 

truncate (truncado)- with the end transversally flat, 

as though it had been cut off 

tuberculate (tuberculado)- with tubercles, i.e., more 

or less rounded protruberances 

turbinate (turbinado)- in the form of an inverted cone 

unicarpellate (unicarpelado)- with only one carpel 

uncinate (uncinado)- claw-shaped 

undulate (ondulado)- wavy, referring to the margin of 

a laminar organ 

unguiculate (unguiculado)- referring to petals with 

the basal portion very narrow, like a claw 

urceolate (urceolado)- urn-shaped 

utricle (utrículo)- a dry, indehiscent, syncarpous, oneseeded 

fruit, enclosed by a bladder composed of 

the concrescent prophylls 

vascular cylinder (cilindro vascular)- the portion of 

the stem composed of xylem, cambium, and phloem 

ventral suture (sutura ventral)- a suture opposite the 

midvein 

xylem vessel (vaso xilemático)- a perforated xylem 

element specialized for water conduction 

vine (bejuco)- a plant that climbs by means of active 

mechanisms or specialized structures 

voluble (voluble)- twining, by means of stems that 

are capable of spiral movement or circumnutation, 

which can be toward the right or the left, depending 

on the species 

wing (ala)- an elongate, membranaceous structure 

zygomorphic (zigomorfa)- having bilateral symmetry, 

i.e., with only one plane of symmetry

470 


Abrus......................................................................242 

precatorius L. ............................................................242 

Acacia .................................................................... 13, 293 

ambigua Vogel ........................................................ 294 

retusa (Aubl.) R.A. Howard ................................... 15, 293 

riparia sensu Britton & P. Wilson ................................... 293 

vogeliana Steud. ........................................................ 294 

westiana DC. .......................................................... 293 

ACANTHACEAE ................................................................. 46 

Achyranthes 

altissima Jacq. .............................................................. 53 

argentata (Mart.) Moq. ................................................... 56 

nodosa Bert. ex Mart. ............................................... 56 

Acrostichum 

sorbifolium L. ........................................................... 36 

Adenocalymna 

obovatum Urb. .......................................................... 123 

portoricense A. Stahl ..................................................... 118 

Adiantum 

aculeatum L. ................................................................ 27 

Adipera 

bicapsularis (L.) Britton & Rose ................................. 237 

Agdestis ...................................................................... 341 

clematidea Moçiño. & Sessé ..................................... 341 

Allamanda ...................................................................... 58 

blanchetii DC. ............................................................... 59 

cathartica L. ................................................................. 58 

AMARANTHACEAE ........................................................ 9, 51 

Ampelopsis 

arborea (L.) Koehne .................................................... 403 

Amphilophium ................................................................ 11, 117 

paniculatum (L.) Kunth ................................................. 117 

Amphistelma 

ephedroides Griseb. ....................................................... 87 

Anechites ....................................................................... 59 

asperuginis Griseb. .......................................................... 59 

nerium (Aubl.) Urb. ........................................................ 59 

Anguria 

cookiana Britton ............................................................ 218 

glomerata Eggers ............................................................ 209 

ottoniana Schltdl. ............................................................ 217 

plumeriana Schltdl. ..................................................... 218 

plumeriana var. trifoliata Cogn. ................................... 218 

trifoliata L. ...................................................................... 218 

trilobata Jacq. ................................................................. 218 

Aniseia ......................................................................... 159 

martinisensis (Jacq.) Choisy ......................................... 159 

Anredera .................................................................. 113 

baselloides (Kunth) Baill. ......................................113 

cordifolia (Tenore) Steenis ........................................ 113 

leptostachys (Moq.) Steenis ........................................... 114 

vesicaria (Lam.) C.F. Gaertn. ....................................... 114 

Anthacanthus 

spinosus (Jacq.) Nees .................................................... 47 

Anthurium .............................................................................. 404 

scandens (Aubl.) Engl. ................................................ 404 

INDEX TO SCIENTIFIC NAMES 

Antigonon ............................................................... 349 

grandiflorum (Bertol) B.L. Rob. ....................... 349 

guatemalense Meisn. ........................................... 349 

leptopus Hook. & Arn. ....................................... 350 

macrocarpum Britton & Small ................................... 349 

APOCYNACEAE ............................................................... 58 

Apocynum 

nerium Aubl. ................................................................ 59 

ARACEAE ........................................................................... 404 

Argyreia ................................................................................ 160 

nervosa (Burm. f.) Bojer ............................................. 160 

speciosa (L. f.) Sweet .................................................... 160 

Aristolochia ..................................................................... 65 

anguicida ..................................................................... 66 

bilabiata sensu Pfeifer .................................................... 70 

calceiformis Urb. ........................................................ 70 

elegans Mast. .............................................................. 66 

galeata sensu authors ...................................................... 71 

gandiflora Sw. ............................................................ 66 

grandiflora sensu Vahl ................................................... 71 

littoralis sensu Pfeifer ................................................ 66 

oblongata subsp. calceiformis (Urb.) R. Rankin & Acev.- 

Rdgz. .................................................................... 70 

odoratissima L. ...................................................... 70 

pandurata Jacq. ........................................................ 70 

ringens Vahl ............................................................ 71 

trilobata L. .............................................................. 71 

ARISTOLOCHIACEAE .................................................. 65 

Arrabidaea ....................................................................... 11, 118 

chica (H. & B.) Velot ....................................... 118 

Arthrostylidium .......................................................... 436 

capillifolium Griseb. ........................................... 436 

farctum (Aubl.) Sodestr. & Lourteig ............................ 436 

multispicatum Pilger .............................................. 437 

sarmentosum Pilger .................................................... 437 

Arum 

hederaceum Jacq. ........................................................... 410 

lingulatum L. .................................................................. 413 

Arundinaria 

multispicata (Pilger) Hack. ......................................... 437 

Arundo 

farcta Aubl. .................................................................. 436 

ASCLEPIADACEAE ............................................................ 73 

Asclepias 

clausa Jacq. ..................................................................... 76 

maritima Jacq. ................................................................ 82 

ASPARAGACEAE .............................................................. 417 

Asparagopsis 

setacea Kunth ................................................................ 418 

Asparagus 

plumosus Baker .............................................................. 418 

setaceus (Kunth) Jessop ................................................. 418 

Aspidium 

articulatum Sw. ................................................................. 22 

ASTERACEAE ................................................................ 13, 91 

Asystasia ................................................................................... 46


gangetica (L.) T. Anders. ....................................... 13, 47 

Banisteria 

emarginata Cav. .................................................... 305 

floribunda DC. ...................................................... 305 

laurifolia L. .................................................... 302 

lucida A. Rich. ................................................. 308 

lupuloides L. ......................................................... 354 

nigrescens A. Juss. ............................................ 308 

ovata Cav. ................................................................. 304 

pubera Rich. ............................................................... 306 

purpurea L. .................................................................... 302 

wydleriana (A. Juss.) C.B. Robinson ............................. 302 

Banisteriopsis 

lucida (A. Rich.) Small .................................................. 308 

Barbieria ............................................................................. 243 

pinnata (Pers.) Baill. ................................................... 243 

polyphylla (Poir.) DC. .................................................... 243 

Basella ............................................................................ 114 

alba L. ............................................................................ 114 

rubra L. ........................................................... 114 

vesicaria Lam. .................................................... 114 

BASELLACEAE .......................................................... 113 

Batocydia 

unguis (L.) DC. .............................................................. 122 

Berylsimpsonia ............................................................... 13, 93 

vanillosma (Wr. & Sauv.) B. Turner ............... 93 

Biancaea 

sepiaria (Roxb.) Todaro .................................................. 235 

Bidens ......................................................................... 94 

alba (L.) DC. ........................................................ 96 

dissecta (O.E. Schultz) Sherff. ....................................... 96 

portoricensis Bello ................................................ 94 

reptans (L.) G. Don ........................................... 94 

reptans var. urbanii (Greenman) O.E. Schulz ................. 94 

scandens L. .............................................................. 109 

urbanii O.E. Shulz .................................................. 15, 94 

Bignonia 

aequinoctialis L. ............................................................. 118 

alliacea Lam. ......................................................... 123 

capensis Thunb. ............................................................. 130 

caryophyllea Bello ...................................................... 130 

chica Humb. & Bonpl. .............................................. 118 

hymenaea DC. ......................................................... 123 

lactiflora Vahl .......................................................... 120 

magnifica W. Bull ........................................................... 128 

paniculata L. ................................................................. 117 

unguis-cati L ................................................................... 122 

venusta Ker-Gawl. .......................................................... 126 

BIGNONIACEAE ........................................................... 11, 116 

BLECHNACEAE .............................................................. 22 

Blechnum ................................................................................ 22 

fragile (Liebm.) Morton & Lellinger ......................... 22 

Boerhavia ............................................................................ 319 

scandens L. .................................................................... 319 

Bonduc 

majus Medic. ............................................................. 235 

BORAGINACEAE ............................................................ 132 

Bougainvillea ..................................................................... 9, 319 

glabra Choisy ................................................................ 321 

spectabilis Willd. .......................................................... 321 

Boussingaultia 

cordifolia Tenore ............................................................. 113 

leptostachys Moq. .................................................... 114 

Brachypterys 

ovata (Cav.) Small .................................................... 304 

Bradburya 

plumieri (Turp. ex Pers.) Kuntze .................................... 250 

pubescens (Benth.) Kuntze .................................. 250 

virginiana (L.) Kuntze ........................................... 251 

Bryonia 

americana Lam. ...................................................... 201 

grandis L. ............................................................ 204 

guadalupensis Spreng. ......................................... 215 

racemosa Mill. ......................................................... 202 

Buddleja ....................................................................... 141 

madagascariensis Lam. .......................................... 141 

BUDDLEJACEAE ...................................................... 141 

Cacalia 

cordifolia L. f. .................................................................. 100 

Cacara 

erosa (L.) Kuntze ........................................................... 274 

CACTACEAE ................................................................... 141 

Cactus 

grandiflorus L. ................................................................ 144 

Caesalpinia .................................................................. 233 

bonduc (L.) Roxb. ....................................................... 234 

bonduc sensu Urb. .......................................................... 234 

ciliata (Wikstr.) Urb. ............................................... 234 

crista sensu Urb. ........................................................... 234 

culebrae (Britton. & P. Wilson) Alain .......................... 235 

decapetala (Roxb.) Alst. ............................................. 235 

divergens Urb. ........................................................... 234 

globerulum Bakh. f. & van Royen ................................. 235 

major (Medik.) Dandy & Exell ...................................235 

portoricensis (Britton & P. Wilson) Alain .................... 237 

sepiaria Roxb. .............................................................. 235 

CAESALPINOIDEAE ..................................................... 233 

Calonyction 

aculeatum (L.) House .................................................. 164 

bona-nox (L.) Boj. ..................................................... 164 

tuba (Schltdl.) Colla ........................................................ 181 

Calopogonium ....................................................................... 245 

caeruleum (Benth.) Sauv. ............................................. 245 

mucunoides Desv. ......................................................... 245 

orthocarpum Urb. ........................................................... 245 

Canavalia ........................................................................ 247 

ensiformis (L.) DC. ................................................... 247 

maritima (Aubl.) Urb. .............................................. 248 

maritima Thouars .................................................... 248 

nitida (Cav.) Piper ................................................ 248 

obtusifolia (Lam.) DC. .................................................. 248 

rosea (Sw.) DC. ............................................................... 248 

rusiosperma Urb. ............................................................ 248 

CAPPARACEAE ................................................................. 146 

Capparis .............................................................................. 146 

flexuosa L. .................................................................. 147 

saligna Vahl ..................................................................... 147 

CAPRIFOLIACEAE ............................................................. 147 

Cardiospermum ..................................................................... 367 

corindum L. .................................................................. 367 

grandiflorum Sw. .................................................... 368 

halicacabum L. ............................................................. 368 

var. halicacabum L. ................................................... 368 

var. microcarpum (Kunth) Bl. .................................. 370 

microcarpum Kunth ....................................................... 370 

Cassia 

antillana (Britton & Rose) Alain ................................... 239 

bicapsularis L. ................................................................. 237

472 


nitida Rich. .................................................................... 239 

Casssytha ................................................................. 300 

americana Nees .................................................... 300 

filiformis L .................................................................... 300 

Cayaponia ................................................................... 201 

americana (Lam.)Cogn. ....................................... 15, 201 

racemosa (Mill.) Cong. ............................................. 202 

CELASTRACEAE .................................................... 11, 13, 149 

Celosia ......................................................................... 51 

nitida ................................................................ 53 

Celtis ............................................................................ 13, 387 

aculeata Sw. ................................................................. 387 

iguanea (Jacq.) Sarg. ..................................................... 387 

Centrosema ......................................................................... 248 

plumieri (Turp.) Benth. ................................................ 250 

pubescens Benth. ..................................................... 15, 250 

virginianum (L.) Benth. ............................................... 251 

Ceratostema 

portoricensis (Urb.) Hoerold ......................................... 228 

Cereus 

trigonus Haw. ............................................................ 142 

undatus Haw. ......................................................... 142 

Chamaefistula 

antillana Britton & Rose .................................................. 239 

Chamissoa ........................................................................ 9, 53 

altissima (Jacq.) Kunth .................................................. 53 

Chiococca ................................................................ 11, 359 

micrantha Johnst. .................................................... 359 

alba (L.) A.S. Hichc. ............................................. 359 

alba (L.) Hitchc. ssp. parvifolia (Griseb.) Steyerm. ...... 359 

alba (L.) Hitchc. var. parvifolia (Griseb.) Urb. ............. 359 

parvifolia Griseb. ........................................................... 359 

racemosa L. ..................................................................... 359 

Chromolaena ..................................................................... 96 

borinquensis (Britton) H. Robins. ......................... 15, 96 

Chusquea ................................................................... 437 

abietifolia Griseb. ........................................................ 439 

Cissampelos ................................................................... 311 

laurifolia Poir. .............................................................. 314 

pareira L. ................................................................. 15, 313 

Cissus ............................................................................. 397 

acida L. .......................................................................... 399 

caustica Tussac ................................................................ 397 

erosa Rich. ............................................................ 397, 401 

intermedia A. Rich. ...................................................... 401 

obovata Vahl. ......................................................... 397, 401 

rotundifolia Vahl ........................................................ 399 

sicyoides L. ................................................................. 401 

trifoliata (L.) L. ..................................................... 399, 401 

tuberculata Jacq. ......................................................... 397 

verticillata (L.) Nicols. & Jarvis ............................ 15, 401 

Citrullus ............................................................................... 202 

lanatus (Thumb.) Mats. & Nakai ................................... 202 

Clematis ......................................................................... 13, 350 

guadeloupae Pers. ............................................................ 354 

dioca L. ................................................................. 352 

flammulastrum Griseb. ............................................... 352 

polygama Jacq. ............................................................ 352 

Clementea 

nitida Cav. ............................................................... 248 

Clerodendrum ............................................................... 390 

aculeatum (L.) Schlecht. ........................................... 390 

thompsonae Balj. ................................................... 392 

thompsonae Balf. f. speciosum (Dombrain)Voss ........ 391 

umbellatum Poir. var. speciosum (Dombrain) ............. 391 

x speciosum van Gert. ........................................... 391 

Clitoria ......................................................................... 251 

falcata Lam .......................................................... 253 

falcata var. falcata f. falcata ................................. 253 

falcata var. falcata f. heteromorpha (Griseb.) Fantz ....... 253 

pinnata (Pers.) R.H. Sm. & G.P. Lewis .......................... 243 

plumieri Turp. ex Pers. .................................................. 250 

polyphylla Poir. ........................................................... 243 

rubiginosa Juss. .............................................................. 253 

ternata L. ...................................................................... 253 

ternatea var. pleniflora Fantz ......................................... 255 

ternatea var. ternatea ..................................................... 255 

virginiana L. ............................................................... 251 

Clusia ............................................................................. 151 

gundlachii A. Stahl ................................................ 15, 152 

CLUSIACEAE ....................................................................... 151 

Coccinia .............................................................................. 204 

cordifolia sensu Britton .................................................. 204 

grandis (L.) Voigt ........................................................ 204 

Cocculus 

domingensis DC. ........................................................ 313 

COMBRETACEAE ........................................................... 153 

Combretum ......................................................................... 153 

grandiflorum G. Don ..................................................... 154 

indicum (L.) Jongkind............................................. 155 

Commicarpus 

scandens (L.) Standl. ..................................................... 319 

Congea .................................................................................. 392 

tomentosa Roxb ................................................... 392 

CONNARACEAE .............................................................. 155 

CONVOLVULACEAE ............................................. 9, 11, 157 

Convolvulus ............................................................... 160 

acuminatus Vahl ........................................................... 169 

albiflorus Vahl .............................................................. 161 

batatas L. ................................................................ 164 

cissoides Lam. ............................................................... 190 

corymbosus L. ........................................................ 198 

cumanenis Kunth ........................................................... 184 

dissectus Jacq. .................................................... 190 

grandiflorus Jacq. .................................................... 181 

havanensis Jacq. .......................................................... 185 

imperati Vahl ............................................................ 167 

littoralis L. ........................................................... 167 

martinicensis Jacq. ................................................. 159 

meyeri Spreng. ..................................................... 171 

nervosus Burm. f. ..................................................... 160 

nil L. .................................................................... 171 

nodiflorus Desr ............................................................ 161 

ochraceus Lindl. ........................................................ 173 

pentanthos Jacq. ......................................................... 185 

pes-caprae L. ........................................................... 173 

purpureus L. ........................................................... 173 

tiliaceus Willd. .......................................................... 179 

tiliaefolius Desr. .............................................................. 197 

tridentatus L. .................................................................. 198 

triqueter Vahl ......................................................... 194 

tuba Schltdl. .................................................................. 181 

turpethum L. .................................................................. 194 

umbellatus L. .......................................................... 192 

ventricosus Bertero ...................................................... 194 

Corallocarpus 

emetocatharticus (Grosourdy) Cogn. ........................ 209 

Cordia ..................................................................................... 132


bellonis Urb. ......................................................... 15, 132 

polycephalla (Lam.) I.M. Johnst. ................................. 133 

wagnerorum Howard ..................................................... 133 

Coreopsis 

reptans L. .................................................................. 94 

Cryptostegia ................................................................... 73 

grandiflora R. Br .................................................. 13, 74 

madagascarensis Bojer ex Dcne. .................................. 74 

Cucumis .............................................................................. 204 

acutangula L. ............................................................. 213 

anguria L. ......................................................... 206 

dispsaceus Ehrenb. ex Spach ......................................... 206 

melo L. ................................................................... 206 

pedatus L. ....................................................................... 218 

trilobatus L. .................................................................... 218 

Cucurbita .............................................................................. 207 

citrullus L. .................................................................... 202 

maxima Duchesne ex Lam. ............................................ 207 

moschata (Duch.) Poir .................................................. 207 

odorifera Vell. ............................................................. 220 

pepo L. .................................................................... 207 

siceraria Molina ..................................................... 211 

CUCURBITACEAE .................................................. 11, 200 

Cuscuta ............................................................................. 222 

americana L. ................................................................ 222 

CUSCUTACEAE ......................................................... 222 

Cydista ..................................................................... 11, 118 

aequinoctialis (L.)Miers .............................................. 118 

Cynanchum 

anegadense (Britton) Alain .............................................. 86 

cheesmanii Woods. ...................................................... 86 

decaisneanum (Schltr.) Alain ............................................ 86 

decipiens (Schltr.) Alain ............................................... 86 

ephedroides (Griseb.) Alain ........................................... 87 

grisebachienum (Schltr.) Alain ................................ 86 

leptocladon (Decne.) Jiménez .................................... 87 

lineare (Bello) Alain ................................................ 87 

monense (Britton) Alain .......................................... 89 

parviflorum (R.Br.) Alain ............................................... 89 

parviflorum Sw. ...................................................... 89 

CYPERACEAE ............................................................. 418 

Dalbergia .............................................................................. 255 

ecastaphyllum (L.) Taub .............................................. 255 

monetaria L. .................................................................. 256 

Dalechampia ........................................................................ 230 


Davallia 

aculeata (L.) J.E. Smith ............................................. 27 

dumosa Sw. ..................................................................... 27 

DAVILLIACEAE ................................................................... 22 

DENNSTEADTIACEAE .................................................... 24 

Desmodium .................................................................... 256 

axillare (Sw.) DC. ......................................................... 256 

canum (J.F. Gmel.) Schinz & Thell. ............................ 258 

incanum DC. .............................................................. 258 

intortum (Mill.) Urb. .................................................... 258 

DICOTYLEDONS ............................................................ 41 

Dicranopteris ..................................................................... 29 

flexuosa (Schrad.) Underw. .......................................... 29 

pectinata (Willd.) Underw. ........................................... 29 

rubiginosa (Mett.) Maxon ........................................ 33 

DILLENIACEAE ........................................................... 9, 223 

Dioclea ...................................................................... 11, 259 

hexandra (Ralph) Mabberley ........................................259 

reflexa Hook. f. .................................................... 259 

Diodia ............................................................................... 359 

sarmentosa Sw. .......................................................... 360 

Dioscorea ......................................................................... 421 

alata L. ....................................................................... 421 

altissima Lam. .............................................................. 422 

bulbifera L. ................................................................ 422 

cayensis Lam. ........................................................ 422 

chondrocarpa Griseb. ............................................ 422 

esculenta (Lam.) Burkill ........................................... 427 

floribunda Mart. & Gal. ................................................ 427 

friedrichsthalii R. Knuth ......................................... 427 

pilosiuscula Bertero ex Spreng. polygonoides 

H. & B. .................................................................... 424 

polygonoides Humboldt & Bonpland ex Willd. .......... 424 

rotundata Poir. ........................................................ 427 

trifida L. .................................................................. 427 

DIOSCOREACEAE ....................................................... 421 

Distictis ................................................................... 11, 120 

lactiflora (Vahl) DC. ..................................................... 120 

Dolicholus 

minimus (L.) Medik. ................................................... 278 

pyramidalis sensu Britton & P. Wilson ................... 279 

reticulatus (Sw.) Millsp. ..................................... 279 

Dolichos 

ensiformis L. ......................................................... 247 

erosus L. ............................................................... 274 

hosei Craib ..................................................................... 286 

lablab L. ................................................................ 265 

luteolus Jacq. ............................................................... 288 

maritimus Aubl. .................................................. 248 

minimus L. ............................................................... 278 

obtusifolius Lam. .......................................................... 248 

phaseoloides Roxb. ....................................................... 277 

pruriens L. ........................................................... 270 

purpureus L. .................................................................. 265 

repens L. ................................................................ 288 

roseus Sw. ............................................................. 248 

uncinatus L. .................................................................... 283 

unguiculatus L. ............................................................. 290 

urens Jacq. .................................................................... 270 

urens L. ........................................................................ 270 

Doliocarpus ..................................................................... 9, 223 

brevipedicellatus Garcke ............................................ 225 

calineoides (Eichl.) Gilg. ................................................ 225 

major Gmel. ........................................................... 225 

olivaceus Sprague & R.O. William ex Standl. ............ 225 

Doxantha 

unguis-cati (L.) Miers. ................................................... 122 

Doyerea ................................................................................. 209 

emetocathartica Grosourd. ........................................ 209 

Dracontium 

scandens Aubl. ............................................................. 404 

Drepanocarpus 

lunatus (l. f.) G.F.W. Meyer .......................................... 266 

Ecastaphyllum 

brownei Pers. ........................................................ 255 

ecastaphyllum (L.) Britton ........................................... 255 

Echites .................................................................................... 59 

agglutinata Jacq. ........................................................ 61 

biflora Jacq. ............................................................ 65 

lappulacea Lam. ........................................................ 59 

Elsota

474 


diversifolia (L.) Blake ............................................... 347 

virgata (Sw.) Kuntze ............................................... 347 

Entada .................................................................................. 294 

polystachya (L.) DC var. polyphylla 

(Benth.) Barneby ........................................................ 294 

polyphyla Benth. .......................................................... 294 

Epidendrum 

claviculatum Wr. ........................................................... 430 

Epipremnum .......................................................................... 406 

aureum (Lindl. & André) Bunting ................................. 406 

pinnatum (L.) Engler .................................................... 406 

pinnatum cultivar Aureum ............................................ 406 

ERICACEAE ........................................................................ 227 

Eupatorium 

borinquense (Britton) B.L. Rob. ..................................... 96 

cordifolium sensu Bello ................................................ 97 

polyodon Urb. ............................................................... 97 

EUPHORBIACEAE ........................................................... 230 

Exogonium 

arenarium Choisy ....................................................... 177 

eggersii House ............................................................ 166 

microdactylum (Griseb.) House ..................................... 171 

repandum (Jacq.) Choisy ............................................... 175 

solanifolium (L.) Britton ............................................. 185 

FABACEAE ........................................................ 9, 11, 13, 233 

FABOIDEAE ....................................................................... 239 

Fevillea ............................................................................... 209 

cordifolia L. .................................................................. 209 

Ficus ................................................................................... 316 

citrifolia Miller ............................................................. 316 

pumila L. ....................................................................317 

Flemingia 

grandiflora Roxb. ex Rottler .............................................51 

Forsteronia ............................................................................. 61 

corymbosa sensu Britton & P. Wilson ........................... 61 

portoricensis Woods. ............................................ 15, 61 

Funastrum .............................................................................. 74 

clausum (Jacq.) Schltr. .................................................. 76 

Galactia ............................................................................... 262 

berteriana DC. ............................................................... 263 

dubia DC. ................................................................. 262 

dubia DC. var. ehrenbii Urb. ...................................... 262 

dubia DC. var. guanicensis Urb. .................................. 262 

eggersii Urb. ........................................................ 15, 262 

longifolia (Jacq.) Benth. .............................................. 263 

pinnata Pers. ............................................................... 243 

striata (Jacq.) Urb. ..................................................... 263 

striata (Jacq.) Urb. var. berteriana (DC.) Urb. ............ 263 

striata (Jacq.) Urb. var. tomentosa (Bertol.) Urb. ....... 263 

Galega 

longifolia Jacq. ............................................................ 263 

Gleichenia .............................................................................. 31 

bifida (Willd.) Spreng. ................................................... 31 

brevipubis Christ. ......................................................... 31 

flexuosa (Schrad.) Mett. .................................................. 29 

rubiginosa Mett. ........................................................... 33 

GLEICHENIACEAE ............................................................ 28 

Glycine 

labialis L. f. ................................................................. 282 

phaseoloides Sw. ......................................................... 279 

reticulata Sw. .................................................................. 279 

striata Jacq. ................................................................... 263 

Gonocalyx ............................................................................. 227 

concolor Nevling .................................................... 15, 227 

portoricensis (Urb.) A. C. Smith. .............................. 228 

Gonolobus ............................................................................ 76 

sintenisii Schltr. ............................................................. 83 

stephanotricus Griseb. ................................................ 76 

variifolius Schltr. ........................................................... 83 

Gothofreda 

cordifolia Vent. ............................................................... 91 

Gouania ..........................................................................11, 354 

domingensis (Jacq.) L. .....................................................354 

lupuloides (L.) Urb. ....................................................... 354 

polygama (Jacq.) Urb. ................................................. 355 

tomentosa Jacq. .......................................................... 355 

Guilandina 

bonduc L. ..................................................................... 234 

bonduc sensu Britton & P. Wilson ............................... 235 

ciliata Bergius ex Wikstr. ............................................. 234 

culebrae Britton & P. Wilson ....................................... 235 

melanosperma Eggers ................................................... 234 

portoricensis Britton & P. Wilson .................................. 237 

Hedysarum 

axillare Sw. ................................................................. 256 

canum J.F. Gmel ............................................................. 258 

ecastaphyllum L. ......................................................... 255 

intortum Mill. ............................................................. 258 

supinum Sw. .................................................................. 258 

Heteropterys .......................................................................... 301 

bellonis Urb. ................................................................... 302 

laurifolia (L.) A. Juss. ................................................. 302 

nitida var. obtusifolia Kuntze ........................................ 305 

purpurea (L.) Kunth. ................................................ 302 

wydleriana A. Juss. ............................................. 15, 302 

Hillia .................................................................................... 360 

longiflora Sw. .............................................................. 360 

parasitica Jacq. ............................................................ 360 

Hippocratea ...................................................................... 11, 149 

caribaea Urb. ..................................................................151 

domingensis Urb. & Ekm. .............................................. 151 

ovata Lam. ................................................................... 149 

volubilis L. .................................................................. 149 

Hoffmannia 

lanceolata Griseb. ........................................................ 362 

Holmskioldia ....................................................................... 394 

sanguinea Retz. ............................................................. 394 

Hornemannia 

racemosa Vahl ............................................................. 228 

Hoya ........................................................................................ 77 

australis R. Br. ex Traill ............................................... 77 

carnosa (L.) R.Br ......................................................... 77 

Humata 

scandens Desv. ................................................................ 27 

Hylocereus ............................................................................ 142 

trigonus (Haw.) Safford ............................................. 142 

undatus (Haw.)Britton ................................................ 142 

Hyperbaena ..................................................................... 9, 313 

apiculata Urb. & Elman ............................................... 316 

dominguensis (DC.) Benth. ....................................... 313 

glauciramis Urb. & Ekman ......................................... 314 

laurifolia (Poir.) Urb. ................................................. 314 

pellucida Urb. & Ekman ............................................. 314 

Hypolepis ................................................................................ 24 

aculeata (Spreng.) J. Smith ........................................... 25 

nigrescens Hook. .......................................................... 25 

repens (L.) C. Presl ....................................................... 25


tenerrima Maxon ......................................................... 25 

urbanii Brause ............................................................. 25 

Ibatia 

maritima (Jacq.) Decne. ................................................ 82 

Ipomoea ............................................................................ 9, 161 

aegyptia L. ................................................................... 189 

alba L. .......................................................................... 164 

angustifolia Jacq. ....................................................... 198 

arenaria (Choisy) Steud. ............................................... 177 

arenaria sensu Urb. ..................................................... 164 

batatas (L.) Lam. ......................................................... 164 

calantha Griseb .......................................................... 166 

cathartica Poir. ............................................................. 169 

cissoides (Lam.) Griseb. ................................................ 190 

eggersii Austin ........................................................... 166 

eriocarpa R.Br. ............................................................ 182 

filicaulis Willd. ........................................................... 198 

filiformis Jacq. ............................................................. 185 

hederifolia L. ................................................................. 166 

horsfalliae W. Hooker ................................................ 167 

imperati (Vahl) Griseb. ................................................ 167 

indica (Burm.) Merril ................................................... 169 

var. acuminata (Vahl) Fosberg ................................. 169 

var. indica .......................................................... 169 

krugii Urb. .................................................................... 182 

littoralis (L.) Boiss ....................................................... 167 

macrantha Roem & Schult. ........................................... 181 

meyeri (Spreng.) G. Don .............................................. 171 

microdactyla Griseb. ................................................ 171 

nil (L.) Roth. ................................................................. 171 

ochracea (Lindl.) G. Don ........................................... 173 

pes-caprae R.Br. .......................................................... 173 

pestigris L. .................................................................. 182 

polyanthes Roem. & Schult ............................................ 192 

portoricensis G. Don .................................................... 169 

pulchella sensu Griseb. ............................................... 181 

purpurea (L.) Roth. ................................................... 173 

quamoclit L. ................................................................. 175 

quinquefolia L. ............................................................. 190 

repanda Jacq. ............................................................... 175 

repanda var. microdactyla (Griseb.) Powell .................. 171 

rubra (Vahl) Millsp. .................................................... 177 

setifera Poir. ........................................................... 15, 177 

solanifolia L. ................................................................ 185 

steudelli Millsp. ............................................................. 177 

stolonifera J. Gmelin ................................................... 167 

tamnifolia L. ................................................................... 187 

tenuissima Choisy ..................................................... 177 

tiliacea (Willd.) Choisy ........................................... 15, 179 

tricolor Cav. .................................................................. 179 

triloba L. ................................................................ 179, 182 

tuba (Schltdl.) Don ........................................................ 181 

tuberosa L. ..................................................................... 192 

verticillata L. ................................................................. 187 

violacea L. ..................................................................... 181 

wrightii A. Gray ............................................................. 181 

Iresine ....................................................................................... 55 

angustifolia Euphras. .................................................. 55 

argentata (Mart.) D. Dietr .............................................. 56 

argentata var. latifolia Suess. .......................................... 56 

celosia L. ........................................................................ 55 

celosioides L. ................................................................ 55 

celosioides Sw. ................................................................. 55 

diffusa H. & B. ex Willd. .............................................. 55 

elatior Rich. ex Willd. .................................................. 55 

paniculata (L.) Kuntze .................................................. 55 

Jacquemontia ...................................................................... 182 

cayensis Britton .......................................................... 184 

cumanensis (Kunth) Kuntze ........................................ 184 

havanensis (Jacq.) Urb. ............................................ 185 

jamaicensis (Jacq.) H. Hallier ex Solereder .................. 185 

nodiflora (Desr.) G. Don .............................................. 161 

pentanthos (Jacq.) G. Don ........................................... 185 

solanifolia (L.) H. Hallier ........................................... 185 

tamnifolia (L.) Griseb. ................................................ 187 

verticillata (L.) Urb. ................................................. 187 

Jasminum ............................................................................. 323 

azoricum sensu Urb. .................................................... 323 

fluminense Vell. .................................................... 13, 323 

grandiflorum L. ....................................................... 324 

multiflorum (Brm.) Andr. .......................................... 324 

pubescens (Retz.) Willd. ................................................ 324 

sambac (L.) Soland. ..................................................... 327 

Justicia 

gangetica L. ................................................................... 47 

spinosa Jacq. ................................................................... 47 

Koanophyllon ......................................................................... 97 

polyodon (Urb.) R. King & H. Rob. ........................... 97 

Lablab ................................................................................. 265 

purpureus (L.) Sweet .......................................... 13, 265 

Lagenaria ................................................................................ 211 

siceraria (Molina)Standl. .............................................. 211 

vulgaris Ser. .................................................................. 211 

Lasianthus ........................................................................... 362 

lanceolatus (Griseb.) M. Gómez .................................. 362 

moralesii (Griseb.) C. Wr. ............................................. 362 

Lasiacis ................................................................................ 439 

divaricata (L.) Hitchc. ............................................... 439 

harrisii Nash ................................................................. 439 

ligulata Hitchc. & Chase ............................................. 440 

sorghoidea (Desv.) Hitchc. & Chase ......................... 440 

swartziana (Hitchc.) Hitchc. ......................................... 440 

LAURACEAE ...................................................................... 300 

Lepidaploa ....................................................................... 13, 97 

borinquensis Urb. ................................................... 15, 97 

Leptoglottis 

portoricensis (Urb.) Britton & Rose .............................. 298 

Lomagramma ........................................................................ 33 

guianensis (Aubl.) Ching .............................................. 33 

Lomaria 

fragilis Liebm ................................................................ 22 

LOMARIOPSIDACEAE ...................................................... 33 

Lomariopsis ............................................................................. 35 

amydrophlebia (Sloss. ex Maxon) Holttrum ................ 35 

kunzeana (Underw.) Holttrum ...................................... 35 

sorbifolia (L.) Fee ......................................................... 36 

Lomoplis 

ceratonia (L.) Raf. ........................................................ 298 

Lonchitis 

repens L. ......................................................................... 25 

Lonicera .................................................................................. 147 

japonica Thumb. ........................................................ 147 

alba L. ........................................................................... 359 

Lophospermum .................................................................... 378 

erubescens D. Don ...................................................... 379 

Luffa ..................................................................................... 211 

acutangula (L.) Roxb. ................................................. 213

476 


aegyptiaca Mill. ........................................................... 213 

cilindrica M.Roem. ......................................................... 213 

Lycianthes .............................................................................. 379 

virgata (Lam.) Bitter ................................................. 380 

Lycopodium 

willdenovii Desv. ex Poir. .............................................. 41 

Lygodium .............................................................................. 39 

japonicum (Thunb.) Sw. ............................................... 39 

Lysiloma 

ambigua (Vogel) Urb. ..................................................... 294 

vogeliana (Steud.) Stehlé ............................................ 294 

Macfadyena .................................................................... 11, 122 

unguis-cati (L.) A. Gentry ........................................ 15, 122 

Machaerium .................................................................... 9, 265 

lunatum (L. f.) Ducke ................................................... 266 

Macrodiscus 

lactiflorus (Vahl) Bureau ............................................... 120 

Macroptilium ...................................................................... 266 

atropurpureum (DC.) Urb. .......................................... 266 

lathyroides (L.) Urb. ................................................... 268 

Malpighia 

bannisterioides L. ........................................................... 304 

MALPIGHIACEAE .............................................................. 300 

Mansoa ................................................................................. 123 

alliacea (Beauv.) A. Gentry .......................................... 123 

hymenaea (DC.) A. Gentry ......................................... 122 

Marcgravia ............................................................. 11, 13, 310 

brittoniana Alain ......................................................... 310 

rectiflora Tria. & Planch. .......................................... 310 

sintenisii Urb. ........................................................ 15, 311 

umbellata Griseb. .......................................................... 310 

MARCGRAVIACEAE ............................................ 11, 13, 310 

Marsdenia ............................................................................. 77 

elliptica Decne. ....................................................... 15, 79 

floribunda (Brongn.) Schltr. ........................................ 79 

woodburyana Acev.-Rodr. ...................................... 15, 81 

Martiusia 

rubiginosa (Juss.) Britton ........................................... 253 

Matelea .................................................................................. 81 

borinquensis Alain ........................................................ 83 

maritima (Jacq.) Woods. ............................................ 82 

sintenisii (Schlecht.) Woods. .................................. 15, 83 

variifolia (Schlecht.) Woods. .................................. 79, 83 

Meibomia 

axillaris (Sw.) Kuntze ................................................. 256 

sintenisii (Urb.) Britton ............................................... 256 

supina (Sw.) Britton .................................................... 258 

umbrosa Britton ........................................................... 256 

Melothria .............................................................................. 213 

guadalupensis (Spreng.) Cogn. ...................................... 215 

pendula L. ................................................................ 15, 215 

MENISPERMACEAE ....................................................... 9, 311 

Merremia ............................................................................. 189 

aegyptia (L.) Urb. ....................................................... 189 

angustifolia (Jacq.) H. Hallier ........................................ 198 

cissoides (Lam.) H. Hallier ........................................ 190 

dissecta (Jacq.) H. Hallier .................................... 15, 190 

quinquefolia (L.) H. Hallier ................................... 15, 190 

tuberosa (L.) Rendle .................................................... 192 

umbellata (L.) H. Hallier ....................................... 15, 192 

Mertensia 

bifida Willd. .................................................................... 31 

flexuosa Schrad. ............................................................. 29 

pectinata Willd. .............................................................. 29 

Metastelma ............................................................................ 85 

albiflorum Schltr. ............................................................ 86 

anegadense Britton ....................................................... 86 

decaisneanum Schltr. ..................................................... 86 

decipiens Schltr. .......................................................... 86 

ephedroides (Griseb.) Schltr. ......................................... 87 

fallax Schltr. ................................................................... 86 

grisebachianum Schltr. ................................................... 86 

leptocladon (Dcne.) Schltr. ........................................... 87 

lineare Bello ................................................................... 87 

monense Britton ..................................................... 15, 89 

parviflorum R.Br. ......................................................... 89 

Microgramma ......................................................................... 36 

heterophylla (L.) Wherry ............................................. 37 

lycopodioides (L.) Copeland ......................................... 37 

piloselloides (L.) Copeland ........................................... 37 

Mikania ................................................................................. 99 

congesta DC. ...................................................... 15, 100 

convolvulacea DC. ..................................................... 100 

cordifolia (L.) Willd. .......................................... 15, 100 

fragilis Urb. ........................................................ 15, 100 

fragilis Urb. var. leptodon Urb. ................................... 100 

gonoclada DC. ............................................................ 100 

micrantha Kunth ...................................................... 103 

micrantha var. congesta (DC.) L.B. Rob. .................... 100 

odoratissima Urb. ................................................. 15, 103 

pachyphylla Urb. .................................................. 15, 103 

porosa Urb. ............................................................ 15, 105 

rotunda sensu A. Stahl .................................................. 100 

scandens sensu A. Stahl .............................................. 100 

stevensiana Britton ............................................. 15, 105 

swartziana sensu A. Stahl ............................................. 105 

Mimosa .......................................................................... 13, 296 

casta L. .......................................................................... 296 

ceratonia L. ................................................................. 298 

diplotricha C. Wr. ........................................................ 298 

paniculata West ex Vahl ............................................... 293 

quadrivalvis L. var. urbaniana Barneby ..................... 298 

retusa Jacq. .................................................................. 293 

MIMOSOIDEAE .................................................................. 293 

Momisia 

iguanaea (Jacq.) Rose & Standl. ................................. 387 

Momordica .......................................................................... 215 

charantia L. ......................................................... 13, 215 

lanata Thunb. ................................................................. 202 

MONOCOTYLEDONS ...................................................... 403 

Monstera ............................................................................... 408 

adansonii Schott ............................................................. 408 

MORACEAE ...................................................................... 316 

Morisonia 

flexuosa L. .................................................................... 147 

Morongia 

portoricensis (Urb.) Britton ............................................ 298 

Mucuna .............................................................................. 268 

altissima (Jacq.) DC. .................................................... 270 

pruriens (L.) DC. .......................................................... 270 

sloanei Fawc. & Rendl. ................................................ 270 

urens (L.) Medik. .......................................................... 270 

urens sensu Urb. ........................................................... 270 

Mycosyrinx 

cissi (Poiret) G. Beck ...................................................... 401 

Neorudolphia .................................................................. 13, 272 

volubilis (Willd.) Britton ....................................... 15, 272 

NYCTAGINACEAE ...................................................... 9, 317


Nyctanthes 

multiflora Burm. f. ..................................................... 324 

sambac L. .................................................................... 327 

Odontosoria .............................................................................. 27 

aculeata (L.) J. Smith ................................................... 27 

scandens (Desvaux) C. Christ. .................................... 27 

uncinella (Kuntze) Fée ................................................... 27 

OLEACEAE ........................................................................ 323 

Oleandra ................................................................................ 22 

articulata (Sw.) C. Presl. ............................................ 22 

nodosa (Willd.) C. Presl .................................................. 22 

Olyra ..................................................................................... 440 

latifolia L. ...................................................................... 440 

Operculina ............................................................................ 194 

grandiflora (Jacq.) House ................................................ 181 

triquetra (Vahl) H. Hallier ............................................ 194 

turpethum (L.) J. Silva. Manso .................................. 194 

turpethum var ventricosa (Bertero) Staples & 

D. F. Austin ................................................................ 194 

ventricosa (Bertero) Peter ........................................... 194 

Ophioglossum 

japonicum Thunb. .......................................................... 39 

Oplonia ................................................................................. 47 

spinosa (Jacq.) Raf. ...................................................... 47 

ORCHIDACEAE ................................................................... 429 

Osmia 

borinquenis Britton ........................................................ 96 

Oxypetalum ........................................................................... 89 

cordifolium (Vent.) Schlecnt. ....................................... 91 

Pachyrhizus ............................................................................. 274 

erosus (L.) Urb. ........................................................... 274 

Panicum 

bambusoides Desv. ex Ham. ........................................ 439 

divaricatum L. .............................................................. 439 

divaricatum L. var. lanatum Schltdl. & Cham. ............. 440 

divaricatum L. var. puberulum Griseb. ...................... 440 

fuscum Sieber ex Griseb. ........................................... 440 

sorghoideum Desv. ....................................................... 440 

Passiflora ...................................................................... 11, 327 

anadenia Urb. ................................................................ 339 

berteroana Balbis ex DC. ........................................... 328 

bilobata Juss. ................................................................. 329 

edulis Sims. ................................................................... 329 

foetida L. ................................................................... 15, 331 

var. foetida ....................................................... 331 

var. hispida (DC.) Gleason ................................. 331 

var. riparia (Griseb.) Killip ................................ 331 

laurifolia L. ................................................................... 331 

maliformis L. ................................................................ 333 

multiflora L. ................................................................ 333 

murucuja L. .................................................................. 334 

pallida L. ....................................................................... 339 

quadrangularis L. ........................................................ 334 

rubra L. ................................................................... 15, 334 

serrato-digitata L. ......................................................... 337 

sexflora Juss. ................................................................ 337 

suberosa L. ............................................................ 15, 339 

tulae Urb. .............................................................. 15, 339 

PASSIFLORACEAE .................................................... 11, 327 

Paullinia ................................................................... 9, 11, 370 

curassavica sensu West ................................................ 374 

diversifolia Jacq. ........................................................... 373 

fuscescens Kunth ......................................................... 370 

pinnata L. ................................................................ 15, 371 

plumierii Triana & Planchon .......................................... 371 

polyphylla L. ............................................................... 374 

triternata Jacq. ............................................................... 374 

Pentalinon ................................................................................ 62 

luteum (L.) Hansen & Wunderlin .................................. 62 

Peperomia ............................................................................. 344 

rotundifolia (L.) Kunth .............................................. 344 

Pereskia .................................................................................. 144 

aculeata Mill. ............................................................ 144 

Petrea ..................................................................................... 396 

volubilis L. ..................................................................... 396 

Pfaffia ................................................................................. 9, 56 

aurata (Mart.) Borsh ..................................................... 56 

grandiflora sensu Liogier ................................................ 56 

Phaseolus .............................................................................. 274 

adenanthus G. Meyer .................................................. 285 

antillanus Urb. ............................................................ 285 

atropurpureus DC. ......................................................... 266 

lanceolatus Bello ........................................................ 286 

lathyroides L. ............................................................... 268 

longifolius Benth. ......................................................... 286 

lunatus L. ..................................................................... 275 

ovatus Benth. .............................................................. 286 

peduncularis Kunth ...................................................... 290 

polystachyos (L.) Britton ............................................... 277 

semierectus L. ............................................................... 268 

vexillatus L. .................................................................... 291 

vulgaris L. ..................................................................... 275 

Philodendron ......................................................................... 408 

angustatum Schott .......................................................... 410 

consanguineum Schott ............................................... 410 

giganteum Schott .......................................................... 410 

hederaceum ............................................................... 410 

isertianum Schott .......................................................... 410 

krebsii Schott ............................................................... 410 

lingulatum (L.) C. Koch ............................................. 413 

marginatum Urb. ............................................................ 410 

micans Klotzsch ex Koch ............................................ 410 

nechodomi Britton ......................................................... 406 

ornatum Schott .............................................................. 413 

oxycardium sensu Britton ........................................... 410 

scandens Koch & Sello ................................................. 410 

Phryganocydia ........................................................................ 124 

corymbosa (Vent.) K. Schum. ...................................... 124 

PHYTOLACCACEAE .......................................................... 341 

Pinzona .......................................................................... 9, 225 

coriacea Mart. & Zucc. .................................................. 225 

Piper 

rotundifolium L. ............................................................ 344 

PIPERACEAE ....................................................................... 344 

Piptocarpha ............................................................................ 105 

tetrantha Urb. .......................................................... 15, 107 

Piptocoma .............................................................................. 107 

acevedoi Pruski ........................................................ 107 

Pisonia ................................................................................... 321 

aculeata L. .................................................................. 321 

helleri Standl. ................................................................. 321 

PLUMBAGINACEAE ........................................................ 345 

Plumbago ............................................................................. 345 


POACEAE ............................................................................ 436 

Podranea ................................................................................ 126 

ricasoliana (Tanf.) Sprague .......................................... 126 

Polygala

478 


diversifolia L. .................................................................. 347 

POLYGALACEAE ........................................................... 9, 347 

POLYGONACEAE ............................................................. 349 

POLYPODIACEAE ............................................................ 36 

Polypodium ............................................................................ 39 

exiguum Heward ........................................................... 37 

guianense Aubl. ............................................................. 33 

heterophyllum L. ............................................................... 37 

loriceum L. .................................................................... 39 

lycopodioides L. ............................................................. 37 

piloselloides L. ................................................................ 37 

serpens Sw. ....................................................................... 37 

Porana 

paniculata Roxb. ........................................................... 195 

Poranopsis .......................................................................... 195 

paniculata (Roxb.) Roberty ......................................... 195 

Pothos 

aurea André ................................................................. 406 

pinnata L. ........................................................................ 406 

Prestonia 

agglutinata (Jacq.) Woods. ............................................... 61 

Pristimera ......................................................................... 11, 151 

caribaea (Urb.) A.C. Smith ............................................ 151 

Protasparagus ....................................................................... 417 

setaceus (Kunth) Oberm. ................................................ 418 

Proustia 

krugiana Urb. ................................................................... 93 

stenophylla Urb. & Ekman .............................................. 93 

vanillosma C. Wright ..................................................... 93 

Pseudogynoxys ................................................................... 109 

chenopodioides (Kunth) Cabrera ............................... 109 

Psiguria ......................................................................... 11, 217 

ottoniana (Schlchet.) C. Jeffrey .................................. 217 

pedata (L.) R.A. Howard ............................................ 218 

trifoliata (L.) Alain ...................................................... 218 

trilobata (L.) Howard ................................................... 218 

Psychotria ............................................................................ 362 

pinularis Sessé & Moçiño ............................................. 362 

microdon (DC.) Urb. .................................................. 362 

PTERIDOPHYTES ................................................................ 21 

Pterocarpus 

lunatus L. f. .................................................................... 266 

Pueraria ................................................................................. 277 

phaseloides (Roxb.) Benth. ..................................... 13, 277 

Pyrostegia ............................................................................ 126 

ignea (Vell.) K. B. Presl ................................................. 126 

venusta (Ker-Gawl.) Miers ........................................... 126 

Quamoclit 

coccinea sensu Urb. ..................................................... 166 

Quisqualis 

indica L. ....................................................................... 155 

Rajania ................................................................................... 427 

cordata L. ..................................................................... 427 

flexuosa Bello .................................................................. 421 

sintenisii Uline ............................................................... 427 

RANUNCULACEAE .................................................... 13, 350 

Reichardia 

decapetala Roth ............................................................... 235 

Rhabdadenia ............................................................................. 62 

biflora (Jacq.) Muell. Arg. ............................................ 65 

paludosa (Vahl) Miers ................................................... 65 

RHAMNACEAE ............................................................ 11, 354 

Rhamnus 

domingensis Jacq. ......................................................... 354 

iguanaeus Jacq. ................................................................ 387 

polygama Jacq. .............................................................. 355 

Rhynchosia .................................................................... 11, 278 

caribaea (Jacq.) DC. ....................................................... 282 

minima (L.) DC. ............................................................. 278 

minima var. lutea Eggers ................................................ 278 

minima var. pauciflora Kuntze ..................................... 278 

phaseloides (Sw.) DC. ................................................. 279 

reticulata (Sw.) DC. .................................................... 279 

Rivea 

campanulata sensu House ............................................ 197 

corymbosa (L.) H. Hallier ............................................ 198 

nervosa (Burm. f.) H. Hallier ......................................... 160 

Rivina 

octandra L. ................................................................... 342 

Rondeletia 

microdon DC. ................................................................. 362 

ROSACEAE ......................................................................... 355 

Rourea .................................................................................. 155 

frutescens sensu Bello ................................................... 157 

glabra sensu Griseb. ........................................................ 157 

surinamensis Miq. ....................................................... 157 

RUBIACEAE ................................................................... 11, 357 

Rubus ................................................................................... 355 

florulentus Focke .................................................. 15, 357 

Rudolphia 

volubilis Willd ............................................................. 272 

Sabicea ................................................................................ 364 

cinerea Aubl. ........................................................... 308, 364 

hirsuta Kunth ................................................................... 364 

moralesii Griseb. ............................................................ 362 

villosa Roem. & Schult. ............................................. 364 

Saelanthus 

rotundifolius Forssk. .................................................... 399 

Salmea ................................................................................ 109 

scandens (L.) Urb. ....................................................... 109 

SAPINDACEAE ........................................................ 9, 11, 366 

Sarcostema 

clausum (Jacq.) R. & S. ................................................. 76 

Saritaea ................................................................................. 128 

magnifica (Sprague) Dugand ....................................... 128 

SCHIZAEACEAE ................................................................. 39 

Schizopsis 

polyanthus Bur. ............................................................. 130 

Schlegelia ....................................................................... 13, 376 

axillaris sensu A. Stahl ................................................... 377 

brachyantha Griseb. .................................................. 377 

brachyantha var. portoricensis Urb. .............................. 377 

portoricensis (Urb.) Britton .......................................... 377 

SCHLEGELIACEAE .................................................... 13, 376 

Schoenus 

secans L. ......................................................................... 419 

Schradera ................................................................................ 364 

capitata Vahl ................................................................. 366 

exotica (Gmelin) Standl. ................................................ 366 

vahlii Steyerm ................................................................. 366 

Schrankia 

distachya sensu A. Stahl ............................................... 298 

leptocarpa sensu Bello .................................................... 298 

portoricensis Urb. ......................................................... 298 

Scleria ..................................................................................... 418 

canescens Boeck. .......................................................... 418 

scindens Nees ex Kunth ................................................. 419 

secans (L.) Urb. ............................................................... 419


SCROPHULARIACEAE ...................................................... 378 

Sechium ................................................................................. 218 

edule (Jacq.) Sw. ............................................................. 220 

Securidaca ........................................................................ 9, 347 

diversifolia (L.) Blake .................................................... 347 

erecta Jacq. .................................................................... 347 

virgata Sw. ...................................................................... 347 

volubilis L. ....................................................................... 256 

Selaginella ............................................................................... 41 

willdenovii (Desv. ex Poir.) Baker .................................. 41 

SELAGINELLACEAE ........................................................... 41 

Selenicereus .......................................................................... 144 

grandiflorus (L.)Britton & Rose .................................. 144 

Senecio 

chenopodioides Kunth ................................................... 109 

confusus Britten .............................................................. 109 

Senegalia 

westiana (DC.) Britton & Rose ....................................... 293 

Senna .............................................................................. 13, 237 

bicapsularis (L.) Roxb. .................................................. 237 

nitida Rich. .................................................................... 239 

Serjania ...................................................................... 9, 11, 373 

diversifolia (Jacq.) Radlk. ............................................. 373 

lucida Schum. ............................................................... 374 

polyphylla (L.) Radlk. .................................................. 374 

triternata (Jacq.) Willd. ............................................... 374 

Sicana .................................................................................... 220 

odorifera (Vell.) Naud. .................................................. 220 

Sicyos 

edulis Jacq. .................................................................. 220 

trifoliata L. .................................................................. 399 

Silphium 

trilobatum L. .................................................................. 111 

SMILACACEAE ............................................................. 13, 443 

Smilax .................................................................................... 443 

coriacea Spreng. ............................................................ 443 

coriacea var. ilicifolia O.E. Schulz ................................. 443 

dominguensis Willd. ...................................................... 444 

guianensis var. subarmata O.E. Schulz .......................... 443 

havanensis Jacq. .......................................................... 444 

havanensis var. portoricensis A. DC. ............................ 443 

ilicifolia sensu Boldingh ............................................. 443 

subarmata O.E. Schulz ................................................... 443 

SOLANACEAE .................................................................... 379 

Solandra ............................................................................... 380 

grandiflora Sw. ............................................................ 380 

longiflora Sw. ............................................................... 382 

Solanum ................................................................................. 382 

lanceifolium Jacq. ......................................................... 382 

pyrifolium Lam. ................................................... 383, 385 

seaforthianum Andr. .......................................... 383, 395 

virgatum Lam. .............................................................. 380 

wendlandii Hook. f. ...................................................... 383 

Spathodea 

corymbosa Vent. ............................................................. 124 

Sphagneticola ........................................................................111 

trilobata (L.) Pruski .................................................... 111 

Stegnosperma ...................................................................... 341 

cubense A. Rich. ............................................................ 341 

Stenochlaena 

amydrophlebia Slosson ex Maxon ................................... 35 

kunzeana Underw. ........................................................... 35 

Stenolobium 

caeruleum Benth. .......................................................... 245 

Stephanotis 

floribunda Brongn. ........................................................... 79 

Sticherus 

brevipubis (Christ.) A.R. Smith ...................................... 31 

Stictocardia ............................................................................ 195 

tiliifolia (Desr.) H. Hallier .......................................... 197 

Stigmaphyllon ..................................................................... 304 

bannisterioides (L.) C. Anderson .................................. 304 

cordifolium Niedenzu ..................................................... 306 

diversifolium (Kunth) A. Juss. .................................... 306 

emarginatum (Cav.)A. Juss. .............................. 305, 306 

floribundum (DC.) C. Anderson ............................. 15, 305 

lingulatum (Poir.) Small ................................................ 305 

nigrescens (A Juss.) Kuntze ......................................... 308 

ovatum (Cav.) Niedenzu ............................................... 304 

periplocifolium (Desf.) A. Juss. ...................................... 305 

puberum (Rich.) A. Juss. ............................................ 306 

retusum Griseb. ............................................................ 306 

tomentosum (Desf.) Niedenzu ........................................ 305 

Stizolobium 

pruriens (L.) Medik. ....................................................... 270 

Symphysia 

racemosa (Vahl) Stearn ................................................... 228 

Syngonium ............................................................................ 415 

podophyllum Schott ..................................................... 415 

Tecoma 

capensis (Thunb.) Lindl. ............................................... 130 

ricasoliana Tanfani ........................................................ 126 

Tecomaria ............................................................................ 128 

capensis (Thunb.) Lindl. ................................................ 130 

Teramnus ............................................................................... 282 

labialis (L.) Spreng. ....................................................... 282 

uncinatus (L.) Sw. .......................................................... 283 

Tetrapterys .............................................................................. 306 

citrifolia sensu Alain ..................................................... 308 

inaequalis Cav. ....................................................... 308, 364 

paniculata Bello ............................................................. 308 

paniculata sensu A. Stahl .............................................. 305 

Thibaudia 

krugii Urb. & Hoerold ................................................... 228 

portoricensis Urb. .......................................................... 228 

Thunbergia .............................................................................. 48 

alata Bojer ex Sims ................................................. 13, 49 

fragrans Roxb. ......................................................... 13, 49 

grandiflora (Roxb. & Rottl.) Roxb. ............................. 51 

volubilis Pers. ................................................................ 49 

Thyella 

tamnifolia (L.) Raf. ....................................................... 187 

Tournefortia .......................................................................... 133 

bicolor Sw. .................................................................. 135 

ferruginea Lam. ............................................................... 138 

hirsutissima L. ............................................................... 135 

laevigata Lam. ............................................................... 135 

laurifolia Vent. ............................................................. 135 

maculata Jacq. .............................................................. 135 

microphylla Bert. ex Spreng. ...................................... 138 

peruviana Poir. ............................................................. 135 

scabra Lam. .................................................................. 138 

volubilis L. ..................................................................... 138 

volubilis L. var. microphylla (Bert. ex Spreng.) 

A.DC. ....................................................................... 138 

Tragia ................................................................................... 231 

volubilis L. ...................................................................... 231 

Trichostigma ........................................................................ 342

480 


octandrum (L.) Walt. ..................................................... 342 

Trommsdorffia 

argentata Mart. ................................................................. 56 

aurata Mart. ..................................................................... 56 

TROPAEOLACEAE ............................................................. 385 

Tropaeolum ............................................................................ 385 

majus L. .......................................................................... 385 

Turbina ........................................................................... 11, 198 

corymbosa (L.) Raf. ....................................................... 198 

Tynnanthus .......................................................................... 130 

polyanthus (Bur.) Sandw. ............................................ 130 

caryophylleus (Bello) Alain ........................................... 130 

ULMACEAE ................................................................... 13, 387 

Urceolaria 

exotica Gmelin .............................................................. 366 

Urechites 

lutea (L.) Britton .............................................................. 62 

Vaccinium ............................................................................. 228 

racemosum (Vahl) Wilber & Luteyn ............................ 228 

Valeriana .............................................................................. 388 

scandens L. .................................................................. 388 

VALERIANACEAE ........................................................... 388 

Vanilla .................................................................................... 429 

barbellata Rchb. ............................................................. 430 

claviculata (Wr.) Sw. .................................................. 430 

correllii Sauleda & Adams ........................................... 434 

dilloniana Corell ............................................................. 430 

eggersii Rolfe ................................................................. 434 

inodora Schiede ............................................................ 432 

mexicana Mill. .............................................................. 432 

planifolia G. Jackson ..................................................... 432 

poitaei Rchb. ................................................................... 434 

pompona Schiede ........................................................... 434 

vanilla (L.) Britton .......................................................... 432 

Varronia 

bellonis (Urb.) Britton ..................................................... 132 

corymbosa (L.) Desv. .................................................... 133 

polycephala Lam. ........................................................... 133 

VERBENACEAE .................................................................. 390 

Vernonia 

borinquensis Urb. ........................................................... 97 

borinquensis Urb. var. hirsuta Gleason .......................... 97 

borinquensis Urb. var. resinosa Gleason ......................... 97 

borinquensis Urb. var. stahlii Urb. .................................. 97 

Vigna .................................................................................. 284 

adenantha (Meyer) Mercht. et al. ............................... 285 

antillana (Urb.) Fawcett & Rendle ................................. 285 

hosei (Craib.) Back. ................................................ 13, 286 

juruana (Harms) Verdc. ................................................. 291 

longifolia (Benth.) Verdc. ............................................... 286 

luteola (Jacq.) Benth. ............................................. 15, 288 

marina (Burm.) Merr. .................................................... 288 

peduncularis Kunth ...................................................... 290 

repens (L.) Kuntze ......................................................... 288 

retusa (E. Meyer)Walp .................................................... 288 

ungiculata (L.) Walp. ..................................................... 290 

vexillata (L.) A. Rich. ................................................. 291 

Vinca 

lutea L. ............................................................................ 62 

Vincetoxicum 

leptocladon Decne. .......................................................... 87 

sintenisii (Schltr.) Britton ................................................ 83 

stephanotrichum (Griseb.) Britton ................................... 76 

variifolium (Schltr.) Britton .......................................... 83 

Viscum 

verticillatum L. .............................................................. 401 

VITACEAE .................................................................... 11, 396 

Vitis ........................................................................................ 401 

caribaea DC. .................................................................... 403 

tiliifolia H. & B. ex Roem. & Schult. ........................... 403 

Volkameria 

aculeata L. .................................................................... 390 

Wedelia 

trilobata (L.) Hitchc. ........................................................ 111 

Xenostegia .......................................................................... 198 

tridentata (L.) D.F. Austin & Staples ......................... 198

Acelga trepadora ............................... 114 

Aguinaldo amarillo ........................ 192 

Aguinaldo azul ............................... 185 

Aguinaldo blanco ............................. 198 

Aguinaldo de costa ......................... 185 

Aguinaldo peludo ............................ 187 

Ahoga gallina ................................. 253 

Ahorca caballo ................................ 302 

Air yam ............................................ 424 

Allamanda ......................................... 58 

Amarat ............................................. 298 

Amarat steckel ................................ 298 

Azufaifo .......................................... 387 

Babeiro ............................................... 61 

Babeiro amarillo ................................ 62 

Bambú trepador ................................ 437 

Bambuilla ......................................... 437 

Barbados gooseberry ........................ 144 

Basket wiss ....................... 342, 371, 374 

Basket-wood ................................... 374 

Basora ............................................. 133 

Bat wings ........................................ 334 

Batata ............................................. 164 

Batatilla blanca ............................... 190 

Bay bean ........................................ 248 

Beach milk vine .................................. 82 

Beach morning glory ........................ 181 

Bejuco blanco ......................... 118, 179 

Bejuco colorado ...................... 175, 272 

Bejuco de agua ................. 225, 403, 406 

Bejuco de ajo .................................. 123 

Bejuco de alambac .......................... 272 

Bejuco de alambre .......................... 272 

Bejuco de alcanfor ........................... 344 

Bejuco de berac .............................. 359 

Bejuco de buey ................................ 302 

Bejuco de calabaza ......................... 413 

Bejuco de calabazón ...................... 410 

Bejuco de caro ........................ 399, 401 

Bejuco de clavo .............................. 130 

Bejuco de conchitas .......................... 253 

Bejuco de corrales ............................ 374 

Bejuco de costilla .................... 371, 374 

Bejuco de cupey .............................. 152 

Bejuco de garrote ............................ 157 

Bejuco de gloria .............................. 169 

Bejuco de guaragüao ....................... 427 

Bejuco de lira .................................. 310 

Bejuco de masa ............................... 135 

Bejuco de mato ............................... 259 

Bejuco de membrillo ....................... 444 

Bejuco de menta ............................. 305 

Bejuco de miel ................................ 109 

Bejuco de mona ................................ 313 


INDEX TO VERNACULAR NAMES 

Bejuco de muela ............................... 109 

Bejuco de nasa ................................. 342 

Bejuco de palma ............................... 342 

Bejuco de paloma ..... 279, 310, 342, 371 

Bejuco de paralejo ................... 302, 308 

Bejuco de puerco .............. 164, 177, 179 

Bejuco de puerco playero ................. 173 

Bejuco de rana ................................ 310 

Bejuco de San Juan .................. 61, 305 

Bejuco de Santiago ............................ 71 

Bejuco de sapo ................................ 310 

Bejuco de sopla ................ 308, 347, 354 

Bejuco de torero ............................... 201 

Bejuco de toro ......................... 302, 305 

Bejuco de vaca ......................... 164, 181 

Bejuco dorado ................................. 300 

Bejuco indio .................................... 354 

Bejuco prieto ............................ 149, 272 

Bejuco rosado ................................... 175 

Bejuquillo de puerco ........................ 179 

Bell Apple ......................................... 331 

Bellísima .................................. 349, 350 

Berenjena de paloma ........................ 380 

Black amaret ................................... 298 

Black eye susan ................................. 49 

Black sage ........................................ 133 

Black whithe .................................... 374 

Black wiss ........................ 251, 342, 374 

Black witty ...................................... 147 

Blue vine ......................................... 253 

Blue wiss ......................................... 251 

Bona wiss ......................................... 265 

Bottle wiss ....................................... 147 

Bran nettle ......................................... 231 

Bull wiss ........................................... 305 

Burro ............................................... 147 

Butterfly pea ........................... 251, 253 

Cabellos de ángel .................... 300, 352 

Cabrite rotting .................................. 374 

Cachimbo ............................................ 71 

Calabazón cimarrón ........................ 413 

Camote .............................................. 164 

Campana de pascua ......................... 383 

Canaria ............................................... 58 

Canario morado falso ......................... 74 

Canavalia ........................................ 248 

Cantiva .............................................. 58 

Caro de tres hojas ............................ 397 

Casabana .......................................... 220 

Cat-claw ............................................ 122 

Chayote ........................................... 220 

Chícharos .......................................... 265 

Chickenet .......................................... 142 

Chiggernit ........................................ 135 

Chiggernit grape ............................... 135 

Cho-cho ........................................... 220 

Chow stick ....................................... 354 

Christmas vine ................................ 195 

Claro de luna .................................. 164 

Clashie-melanie ............................... 185 

Climbing bamboo ........................... 436 

Clock vine ......................................... 49 

Coast moon vine .............................. 181 

Cock spur ........................................ 387 

Cocombro ......................................... 206 

Conchita de Virginia ......................... 251 

Conchitas .......................................... 253 

Consuelda ........................................ 114 

Coral ................................................ 350 

Coral vine ......................................... 350 

Coralillo ........................................... 350 

Coralina .......................................... 350 

Coralita ........................................... 350 

Corona de novia .............................. 198 

Corsú ................................................ 277 

Cortadera ......................................... 418 

Cortadora ....................................... 418 

Cow-itch ........................................... 270 

Cowpea ........................................... 290 

Cresta de gallo blanca ..................... 283 

Cundeamor ....................................... 215 

Cupeillo de altura ............................. 152 

Cupey de altura ............................... 152 

Curuba ............................................ 220 

Deleite ............................................. 253 

Diamela ............................................ 327 

Dragon wiss ..................................... 302 

Dunguey ......................... 422, 424, 443 

Dunguey blanco ................................ 443 

Enredadera ........................................ 243 

Escambrón ............................... 266, 321 

Espinaca de Nueva Zelandia ............ 114 

Esponja ............................................ 213 

Estefanotis ......................................... 79 

Estropajo ........................................ 213 

Falsa belladona ................................ 383 

Fideillo .................................... 222, 300 

Fideos ............................................. 300 

Flor de conchitas ...................... 250, 251 

Flor de luna .............................. 164, 181 

Flor de nieve ...................................... 49 

Flor de palo ....................................... 192 

Flor de pasión ........................... 334, 339 

Flor de pito ............................... 250, 253 

Flor de sierra .................................... 360 

Flor de la mañana ............................ 383 

Florador .............................................. 79 

Fresa de la montaña ......................... 357

482 

Frijol ............................................... 290 

Frijol caballero ................................. 265 

Frijol cimarrón ................................. 291 

Frijol silvestre .................................. 288 

Frijoles .............................................. 275 

Frijolillo ........................... 278, 279, 282 

Frijolillo peludo ............................... 283 

Gallito ................................................. 71 

Giant potato climber ....................... 383 

Giniper ............................................. 135 

Gloria de la mañana .......................... 177 

Goat foot ................................. 173, 179 

Goat wiss ................................. 147, 288 

Granadilla ......................................... 334 

Green briar ....................................... 443 

Guaco ...................................... 100, 105 

Guaco de cabra ................................ 105 

Guaco falso ............................. 100, 103 

Guaco redondo ................................. 100 

Guanabanilla cimarróna .................... 82 

Guard wiss ........................................ 118 

Guáyaro .......................................... 427 

Güicharo ........................................... 211 

Güiro ............................................... 211 

Gunda ..................................... 422, 424 

Haba de burro de jardín .................. 248 

Haba de playa ................................. 248 

Habas de caballo ............................... 265 

Habichuela cimarrona .............. 285, 286 

Habichuela parada ...........................268 

Habichuela playera ......................... 248 

Habichuelas ..................................... 275 

Habilla ..................................... 278, 279 

Hediondilla ...................................... 239 

Hiedra .............................................. 317 

Higüerito de sierra ............................ 377 

Hilia ................................................. 360 

Hoja de sen ....................................... 237 

Honeysuckle ........................... 147, 350 

Hoopvine .......................................... 342 

Hortensia de bejuco .......................... 144 

Indigo berry ...................................... 339 

Ink berry ......................................... 339 

Iron weed ........................................ 262 

Jaboncillo ........................................ 347 

Jagüey blanco ................................. 316 

Jazmín ............................................... 324 

Jazmín de canario ........................... 323 

Jazmín de España ............................. 324 

Jazmín de Italia ............................... 383 

Jazmín de papel ............................... 324 

Jazmín de trapo ................................ 323 

Jazmín oloroso .................................. 323 

Jícama ...................................... 245, 274 

Juan caliente .................................... 157 

Jumbee pumpkin .............................. 215 

Jumbee seeds .................................... 242 

Katch and keep ................................. 293 

Kudzu tropical .................................. 277 

Lambrali ........................................... 401 

Lamedora ....................................... 298 

Leche de perra .................................... 61 

Lemon vine ...................................... 144 

Liana de la sierra ............................. 118 


Liana fragante ................................. 120 

Liana uñada ..................................... 122 

Limber caper ..................................... 147 

Lluvia de orquideas ......................... 392 

Love-in-the-mist ............................... 331 

Madagascar jasmin ........................... 79 

Madeira vine ........................... 113, 114 

Madreselva ...................................... 147 

Maiden apple ................................... 215 

Mari-de-Lugo .................................. 175 

Marimbo ........................................... 211 

Mascapelo ........................................ 355 

Mata de ajo ...................................... 123 

Mata de cera ....................................... 77 

Mata de peo ..................................... 362 

Mato ................................................ 270 

Mato colorado ................................ 248 

Mato de playa ........................ 234, 248 

Matos ............................................. 270 

Matos del monte .............................. 270 

Maya prieta ....................................... 259 

Melón ............................................... 202 

Membrillo ......................................... 256 

Mexican creeper ............................... 350 

Mijo del sol ..................................... 379 

Ñame ................................................. 421 

Ñame blanco .................................... 421 

Ñame de Guinea ............................... 422 

Ñame de gunda ................................. 424 

Ñame dunguey ................................ 422 

Ñame gulembo ................................. 427 

Nigua ............................................... 135 

Ojo de buey ...................................... 270 

Ojos de cangrejo .............................. 242 

Old maid .......................................... 215 

Oreja de ratón .................................. 313 

Ox-eye-bean .................................... 270 

Paisaje .............................................. 410 

Palawang .......................................... 147 

Palinguam ......................................... 147 

Palo de brasilete ............................... 256 

Palo de burro .................................... 147 

Palo de hoz ..................................... 266 

Palo de perico ................................... 133 

Panitos ............................................... 71 

Papa aerea ......................................... 383 

Papbush ............................................ 331 

Papitos ............................................. 253 

Paraguita chino ................................. 394 

Paralejo velludo ............................... 302 

Parasol ............................................. 394 

Parcha ....................................... 329, 331 

Parcha Cimarrona ........................... 333 

Parcha Colorada .............................. 334 

Parcha yedra .................................... 339 

Parchita ............................................ 337 

Pareira ............................................... 313 

Passion fruit ...................................... 329 

Pega palo ........................................... 120 

Pegapalma ................................ 310, 311 

Pelicanos ............................................. 71 

Pepineto ............................................ 206 

Pepinillo cimarrón ........................... 215 

Pepinillo silvestre ............................ 206 

Pepino .............................................. 220 

Pepino angolo .................................. 220 

Pepita amarga ................................. 209 

Peronías .......................................... 242 

Pica-pica .......................................... 270 

Pinna koop ....................................... 401 

Pitahaya ........................................... 142 

Platillo ............................................. 394 

Ponpón ............................................... 82 

Prickly-mampoo ............................... 321 

Pringamosa ...................................... 231 

Puddin vine ..................................... 401 

Purple alamanda ................................ 74 

Raíz de zarzaparilla ......................... 444 

Rasca garganta ................................ 410 

Razor grass ...................................... 419 

Red wiss .................................. 302, 305 

Rubber vine ....................................... 74 

Sandia .............................................. 202 

Sanjuanera ......................................... 61 

Saragüero .......................................... 133 

Scrubber ............................................ 242 

Secua ............................................... 209 

Sen del país .................................... 237 

Snow berry ...................................... 359 

Soap stick ........................................ 354 

Soap vine .......................................... 354 

Sopla ................................................. 354 

Sorrel vine ....................................... 399 

Sponge cucumber ............................. 213 

Stiver bush ....................................... 237 

Strainer vine ...................................... 213 

Strawberry pear ................................ 142 

Suelda con suelda ............................. 114 

Susana ................................................ 49 

Susana blanca ................................... 49 

Sweet pea .......................................... 265 

Sweet potato ...................................... 164 

Tagua-tagua ............................. 331, 337 

Tagua-tagua serrana ....................... 339 

Tente en el aire ......................... 222, 300 

Terciopelo ....................................... 392 

Tibey trepador ................................. 360 

Tobacco pipe ....................................... 71 

Tortera ............................................. 270 

Trinitaria .......................................... 321 

Tulipa .............................................. 377 

Tunbergia ............................................ 51 

Uña de gato ..................................... 122 

Uyama .............................................. 209 

Valeriana .......................................... 388 

Velo de novia .................................... 195 

Velvet leaf ......................................... 313 

Viuda ......................................... 49, 120 

Water lemon .................................... 329 

Water Yam ........................................ 421 

Wax plant ...................................... 77, 79 

Wedelia ............................................. 111 

White fig .......................................... 316 

White root ........................................ 354 

White wist ....................................... 374 

White yam ............................... 421, 427 

Wild allamanda ................................. 62 

Wild bush bean ................................ 268

Wild cucumber ................................. 206 

Wild Licorice ................................... 242 

Wild marygold .................................. 111 

Wild morning glory ......................... 177 

Wild potato ...................................... 179 

Wild vine ......................................... 179 


Wild yam .......................................... 424 

Yellow dodder ................................. 300 

Yellow love ..................................... 300 

Yellow morning glory ...................... 192 

Yerba de medio real ......................... 344 

Yerba de ratón ................................... 344 

Yerba linda ........................................ 344 

Zarza ............................... 235, 296, 298 

Zarza brava ....................................... 293 

Zarza de cercas ................................ 235 

Zarzaparilla ............................. 298, 444

Vines and Climbing Plants of Puerto Rico and the Virgin Islands

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